The authors' approach has been shaped by careful study of the scholarly work of Dr Henriette Harmse (European Bioinformatics Institute) and Dr Marie Keet (University of Cape Town).
Dr Christopher Johann Muller rer. nat. (data modeler, IT architect, programmer)
Dr Edit Hlaszny PhD (domain expert, editor)
The VVO (Vitis Vinifera Ontology) is a controlled vocabulary that formally describes the morphological structures, phenological growth and developmental stages, quality parameters, dispositions, roles, and information entities of Vitis vinifera L., the common grapevine.
OWL/XML encoding (+several serialisations)
Dr Hlaszny Biosystems Engineering: In Vivo Integration of Neural Parenchyma
This ontology is distributed under an Attribution 4.0 International (CC BY 4.0). The License is: https://creativecommons.org/licenses/by/4.0/. Copyright: Dr Edit Hlaszny (2026-)
UDC 634.8 Viticulture. Grape vines. Vineyards
3MH precursor concentration (Cys-3MH + Glut-3MH) ranges from 50-150 µg/L in low-precursor cultivars to 400-1200 µg/L in high-precursor Sauvignon Blanc from Marlborough. Cysteine-3MH typically comprises 60-80% of total 3MH precursors, with glutathione-3MH as the biosynthetic intermediate. Wine 3MH concentrations of 200-8000 ng/L result from 1-5% conversion of grape precursors during fermentation, vastly exceeding the sensory threshold of 60 ng/L. The released 3MH shows distinct grapefruit (zesty citrus) and passionfruit (tropical) character, with perception varying by enantiomeric ratio ((R)-3MH more citrus, (S)-3MH more passionfruit). 3MH can be acetylated to 3-mercaptohexyl acetate (3MHA) during fermentation, adding boxwood and tropical notes with a threshold of 4 ng/L. Precursor accumulation increases during ripening, with sun exposure promoting accumulation through glutathione conjugation mechanisms.
A thiol precursor accumulation disposition that is the inherited capacity of a Vitis vinifera plant to accumulate cysteine and glutathione conjugates of 3-mercaptohexan-1-ol (3MH) in grape berries, with genotype-specific concentrations determining the potential for grapefruit, passionfruit, and tropical fruit aroma release during fermentation.
3MH and its acetate 3MHA are the primary contributors to Sauvignon Blanc tropical character, with concentrations typically 10-100× above threshold in aromatic examples. The precursor concentration in grapes sets the ceiling for potential wine thiol content, but fermentation conditions (yeast strain, nutrition, temperature) strongly modulate conversion efficiency. The dual origin of 3MH (from both (E)-2-hexenal and hexanal via conjugation) provides metabolic flexibility in precursor synthesis. Water deficit and nitrogen status affect precursor accumulation through effects on glutathione metabolism.
3-mercaptohexanol potential
3MH conjugate capacity
3MH-Vorläufer
Grapefruit-Thiol
Passionsfrucht-Aroma
grapefruit thiol precursor
Peyrot des Gachons C et al. (2002) Influence of water and nitrogen deficit on fruity and herbaceous aroma potential of Vitis vinifera L. cv. Sauvignon blanc in three terroirs. Journal of the Science of Food and Agriculture 82:1167-1174 Capone DL et al. (2010) The cysteine conjugate of 3-mercaptohexan-1-ol in grape juice. Food Chemistry 119:1326-1330
3MH Precursor Disposition
3MH precursor quality in Sancerre Sauvignon Blanc illustrates Loire Valley terroir expression, with Kimmeridgian marl soils producing grapes with 800-1500 μg/L Cys-3MH (versus 400-700 μg/L from siliceous soils), translating to wines with 1500-3500 ng/L free 3MH that provide the exotic fruit dimension complementing the region's characteristic minerality this precursor concentration difference between soil types contributes to the stylistic variation among Sancerre vineyard sites, demonstrating how 3MH precursor quality links terroir to aromatic expression.
3MH_Precursor_Quality represents the chemical quality characterizing the concentration of precursor conjugates for 3-mercaptohexan-1-ol (3MH) in grape berries, capturing the bound forms of the volatile thiol responsible for the distinctive passion fruit, grapefruit, and citrus zest aromas that define aromatic Sauvignon Blanc character. The 3MH precursors include S-3-(hexan-1-ol)-L-cysteine (Cys-3MH) and S-3-(hexan-1-ol)-glutathione (Glut-3MH), both formed in grapes through conjugation of (E)-2-hexenal with cysteine or glutathione via Michael addition reactions. Cys-3MH is the direct substrate for yeast β-lyase cleavage releasing 3MH, while Glut-3MH requires prior cleavage to the cysteine form. In Sauvignon Blanc grapes, total 3MH precursor concentration typically ranges from 200-2000 μg/L depending on cultivar clone, growing conditions, and harvest timing, with precursor accumulation increasing during late ripening. The released 3MH (3-mercaptohexan-1-ol) has an extraordinarily low sensory threshold of approximately 60 ng/L in wine, with typical concentrations in aromatic Sauvignon Blanc of 500-5000 ng/L providing levels 10-80 times threshold. This substantial suprathreshold concentration explains the dominant tropical fruit character of 3MH in finished wines. However, the conversion efficiency from precursor to free thiol is low (typically 1-10%), meaning that precursor concentration provides an upper limit but not a direct prediction of wine 3MH content. The 3MH released during fermentation can undergo acetylation to form 3-mercaptohexyl acetate (3MHA), which has an even lower threshold (approximately 4 ng/L) and contributes passion fruit and guava notes, making 3MH precursor content relevant to both 3MH and 3MHA potential in finished wines.
3MH_Precursor_Quality specifically addresses the bound precursor forms for the dominant tropical thiol in Sauvignon Blanc. The class encompasses both cysteine and glutathione conjugates that can ultimately yield 3MH. The dramatic concentration ratio between precursor (hundreds to thousands of μg/L) and released thiol (hundreds to thousands of ng/L) reflects the low conversion efficiency that makes yeast strain selection critical for thiol release. The further conversion of 3MH to 3MHA through acetylation during fermentation adds complexity to the precursor-to-aroma relationship.
3-Mercaptohexanol-Vorläufergehalt
3-mercaptohexanol precursor content
3MH conjugate content
3MH-Vorläufergehalt
Cys-3MH concentration
Cys-3MH-Konzentration
Passionsfrucht-Vorläuferqualität
passion fruit precursor quality
Subileau M et al. (2008) New Insights on 3-Mercaptohexanol (3MH) Biogenesis in Sauvignon Blanc Wines: Cys-3MH and (E)-Hexen-2-al Are Not the Major Precursors. Journal of Agricultural and Food Chemistry 56(19): 9230-9235 Capone DL et al. (2011) Evolution and Occurrence of 1,6-Dimethyl-1,2,3,4-tetrahydronaphthalene (TDN) in Aged Riesling Wines. Journal of Agricultural and Food Chemistry 59(11): 6112-6119
3MH Precursor Quality
4MMP precursor concentration (Cys-4MMP + Glut-4MMP) ranges from 5-20 µg/L in most cultivars to 30-80 µg/L in high-precursor Sauvignon Blanc clones. 4MMP precursors accumulate at 10-20× lower concentration than 3MH precursors, but the 75× lower sensory threshold (0.8 ng/L versus 60 ng/L for 3MH) means 4MMP significantly impacts wine aroma even at trace concentrations. Wine 4MMP concentrations of 5-50 ng/L are typical in aromatic Sauvignon Blanc, with the boxwood and blackcurrant bud character becoming dominant above 20 ng/L. 4MMP shows no acetylation (lacking the hydroxyl group of 3MH), maintaining its ketone form throughout wine aging. The precursor arises from conjugation of mesityl oxide with glutathione/cysteine, with biosynthesis pathway details less characterized than for 3MH. Precursor concentration shows strong clone and terroir effects within Sauvignon Blanc.
A thiol precursor accumulation disposition that is the inherited capacity of a Vitis vinifera plant to accumulate cysteine and glutathione conjugates of 4-mercapto-4-methylpentan-2-one (4MMP) in grape berries, with genotype-specific concentrations determining the potential for boxwood, blackcurrant bud, and broom aroma release during fermentation.
4MMP contributes the distinctive boxwood/blackcurrant character that, together with the grapefruit notes of 3MH, defines Sauvignon Blanc typicity. The extremely low threshold means 4MMP significantly impacts aroma even in wines with modest precursor levels. The blackcurrant bud character can be perceived as either desirable (varietal typicity) or undesirable (cat urine association at high concentrations) depending on context and concentration. Some Sauvignon Blanc terroirs (Sancerre) show higher 4MMP/3MH ratios than others (Marlborough), contributing to regional style differences.
4-mercapto-4-methylpentan-2-one potential
4MMP conjugate capacity
4MMP-Vorläufer
Buchsbaum-Thiol
Schwarze-Johannisbeere-Aroma
boxwood thiol precursor
Tominaga T et al. (1998) Development of a method for analyzing the volatile thiols involved in the characteristic aroma of wines made from Vitis vinifera L. cv. Sauvignon blanc. Journal of Agricultural and Food Chemistry 46:1044-1048 Helwi P et al. (2016) Vine nitrogen status and volatile thiols and their precursors from plot to transcriptome level. BMC Plant Biology 16:173
4MMP Precursor Disposition
4MMP precursor quality in Pouilly-Fumé Sauvignon Blanc contributes the smoky, gunflint character (fumé) that historically named the appellation, with Cys-4MMP concentrations of 150-350 μg/L in ripe fruit translating to 15-40 ng/L free 4MMP in finished wines this 4MMP level, approximately 10-50 times sensory threshold, provides the characteristic box tree-blackcurrant complexity that distinguishes Pouilly-Fumé from the more purely tropical-fruited Sauvignon Blanc styles of New Zealand, demonstrating how 4MMP precursor quality contributes to regional typicity.
4MMP_Precursor_Quality represents the chemical quality characterizing the concentration of precursor conjugates for 4-mercapto-4-methylpentan-2-one (4MMP) in grape berries, capturing the bound form of the volatile thiol responsible for the distinctive box tree (boxwood), blackcurrant bud, and broom aromas in Sauvignon Blanc and related varieties. The 4MMP precursor, S-4-(4-methylpentan-2-one)-L-cysteine (Cys-4MMP), is formed in grapes through conjugation pathways analogous to 3MH precursor formation. In Sauvignon Blanc grapes, Cys-4MMP concentration typically ranges from 50-500 μg/L, substantially lower than 3MH precursor levels, reflecting the compound's status as a secondary contributor to the varietal thiol profile. The released 4MMP (4-mercapto-4-methylpentan-2-one) has an extremely low sensory threshold of approximately 0.8-3 ng/L in wine, among the lowest known thresholds for wine aroma compounds. This remarkable potency means that even the low conversion efficiency (typically 1-5%) from precursor to free thiol yields perceptible concentrations, with typical 4MMP levels in aromatic Sauvignon Blanc of 5-50 ng/L providing 5-50 times threshold concentration. The box tree/blackcurrant bud aroma of 4MMP provides a distinctive green-sulfurous note that complements the tropical fruit character of 3MH, with the balance between these thiols contributing to wine style variation. Cultivar and clonal variation affects 4MMP precursor accumulation, with Sauvignon Blanc showing highest levels among commercial varieties. The accumulation of 4MMP precursor during ripening and its response to viticultural factors parallels 3MH precursor behavior, though the magnitude of accumulation is typically lower.
4MMP_Precursor_Quality specifically addresses the bound precursor form for the box tree/blackcurrant thiol that complements 3MH in varietal thiol profiles. The extremely low sensory threshold of released 4MMP (below 1 ng/L in some studies) makes even very low conversion efficiency yield perceptible aroma impact. The distinct sensory character of 4MMP (vegetal, box tree, blackcurrant) versus 3MH (tropical fruit, passion fruit) means their ratio affects overall thiol aroma profile beyond their individual contributions. Analysis of Cys-4MMP employs the same LC-MS/MS methods as other thiol precursors.
4-Mercapto-4-methylpentan-2-on-Vorläufergehalt
4-mercapto-4-methylpentan-2-one precursor content
4MMP conjugate content
4MMP-Vorläufergehalt
Buchsbaum-Vorläuferqualität
Cys-4MMP concentration
Cys-4MMP-Konzentration
box tree precursor quality
Tominaga T et al. (1998) Identification of New Volatile Thiols in the Aroma of Vitis vinifera L. var. Sauvignon Blanc Wines. Flavour and Fragrance Journal 13(3): 159-162 Peyrot des Gachons C et al. (2002) Influence of Water and Nitrogen Deficit on Fruity Aroma Potential of Vitis vinifera L. cv. Sauvignon Blanc in Three Terroirs. Journal of the Science of Food and Agriculture 82(11): 1233-1243
4MMP Precursor Quality
In Sauvignon Blanc vines in Marlborough, New Zealand, ABA accumulation in axillary buds progresses from baseline concentrations of 40-50 ng per gram fresh weight in February to 80-100 ng per gram by late March as photoperiod decreases below critical threshold, reaching maximum concentrations of 180-220 ng per gram fresh weight by late May coincident with full dormancy establishment, with NCED gene expression increasing 4-fold during this period.
ABA accumulation in Vitis vinifera represents the progressive increase in abscisic acid concentration within bud and other tissues during dormancy establishment, reflecting both increased biosynthesis and decreased catabolism of this key regulatory hormone that plays central roles in dormancy induction, maintenance, and cold hardiness development. This process occurs during autumn, typically from September through November in Northern Hemisphere viticulture, with ABA levels in bud tissues increasing several-fold from growing season baseline values to reach maximum concentrations coincident with full dormancy establishment. The biosynthesis of ABA during dormancy induction involves increased expression and activity of 9-cis-epoxycarotenoid dioxygenase enzymes that catalyze the rate-limiting step in ABA production from carotenoid precursors, with NCED gene expression increasing in response to both short-day signals and low temperature exposure. The physiological functions of accumulated ABA during dormancy establishment include direct inhibition of cell division and elongation in meristematic tissues through effects on cell cycle regulators and expansion-related gene expression, promotion of storage compound accumulation by redirecting metabolism from growth toward reserve synthesis, induction of cold hardiness through activation of cold-responsive gene expression, and maintenance of the dormant state by preventing premature growth resumption in response to temporary favorable conditions. The regulation of ABA accumulation involves integration of multiple environmental signals, with short photoperiod and low temperatures both promoting ABA biosynthesis while also reducing ABA catabolism through decreased activity of ABA 8'-hydroxylase enzymes, resulting in net ABA accumulation that reflects both production and turnover rates. The concentration of ABA in dormant buds typically reaches values 3-5 times higher than in actively growing buds, with maximum concentrations of 150-250 nanograms per gram fresh weight in fully dormant buds of most Vitis vinifera cultivars. The subsequent decline in ABA concentration during chilling accumulation and dormancy release represents an important component of the transition from endodormancy to ecodormancy, with decreasing ABA levels permitting the eventual growth response to favorable spring temperatures. Cultivar differences in ABA accumulation patterns contribute to variation in dormancy depth and cold hardiness among genotypes.
ABA_Accumulation represents the hormonal dimension of Dormancy_Establishment, specifically addressing the increase in abscisic acid that regulates dormancy. Distinguished from Metabolic_Adjustment which addresses broader metabolic changes, while ABA_Accumulation specifically addresses this key regulatory hormone. Involves increased NCED expression and decreased ABA catabolism. Typical maximum concentrations 150-250 ng per gram fresh weight in dormant buds. Duration spans 8-10 weeks during autumn. Subsequent ABA decline during chilling is associated with dormancy release.
ABA buildup
ABA-Akkumulation
ABA-Aufbau
Abscisinsäure-Anreicherung
Dormanzhormon-Akkumulation
abscisic acid accumulation
abscisic acid increase
dormancy hormone accumulation
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Zheng C et al (2015) Abscisic acid catabolism enhances dormancy release of grapevine buds. Plant Cell and Environment 38: 2490-2501 Or E et al (2000) The transduction of the signal for grape bud dormancy breaking induced by hydrogen cyanamide may involve the SNF-like protein kinase. Plant Molecular Biology 43: 483-494
ABA Accumulation
In Tempranillo vines in Rioja, Spain, ABA maximum level is reached in early December at approximately 190-210 ng per gram fresh weight in primary buds, representing a 4-fold increase from September baseline values of 45-55 ng per gram, with this peak concentration maintained for approximately 3-4 weeks before chilling-induced catabolism progressively reduces ABA levels through January, coinciding with gradual dormancy depth reduction as measured by decreasing forcing time requirements.
ABA maximum level in Vitis vinifera represents the peak concentration of abscisic acid achieved in bud tissues during deep endodormancy, reflecting the culmination of autumn ABA accumulation and constituting a key hormonal component of maximum dormancy depth. This peak concentration typically occurs during December in Northern Hemisphere viticulture, coincident with other indicators of maximum dormancy intensity, before chilling-dependent ABA catabolism begins reducing hormone levels as part of the dormancy release process. The quantitative values of ABA maximum level vary among cultivars but typically fall in the range of 150-250 nanograms per gram fresh weight in dormant buds, representing concentrations 3-5 fold higher than those present in actively growing buds during summer. The physiological significance of ABA maximum level relates to the multiple growth-inhibiting functions of this hormone, including direct inhibition of cell cycle progression through effects on CDK activity and cyclin expression, suppression of growth-related gene expression through ABA-responsive transcription factors, and promotion of storage and protective processes that characterize the dormant state. The achievement of ABA maximum level requires both active ABA biosynthesis through the NCED pathway that continues during early dormancy, and reduced ABA catabolism through decreased activity of ABA 8'-hydroxylase enzymes, with the balance between production and degradation determining the peak concentration achieved. The subsequent decline from ABA maximum level during chilling accumulation is an essential component of dormancy release, with chilling exposure activating ABA catabolic pathways and reducing biosynthetic gene expression, progressively lowering hormone concentrations toward levels compatible with growth resumption. The relationship between ABA maximum level and chilling requirement is complex, with some evidence suggesting that cultivars achieving higher ABA maximum levels may require more chilling for complete hormone reduction and dormancy release, though genetic variation in ABA sensitivity also affects this relationship. Environmental factors during dormancy induction can influence ABA maximum level, with stress conditions potentially elevating peak concentrations while suboptimal acclimation conditions might result in lower maximum levels.
ABA_Maximum_Level represents the hormonal peak within Deep_Endodormancy specifically addressing abscisic acid concentration maximum. Distinguished from ABA_Accumulation which addresses the process of increasing ABA during dormancy establishment. Typical peak values 150-250 ng per gram fresh weight depending on cultivar. Duration of peak plateau typically 2-4 weeks before chilling-induced decline begins. Subsequent ABA reduction is a key component of dormancy release mechanism.
ABA peak
ABA-Höchstwert
ABA-Maximalgehalt
Abscisinsäure-Maximum
highest ABA level
maximaler Abscisinsäure-Spiegel
maximum abscisic acid
peak ABA concentration
Zheng C et al (2015) Abscisic acid catabolism enhances dormancy release of grapevine buds. Plant Cell and Environment 38: 2490-2501 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Or E et al (2000) The transduction of the signal for grape bud dormancy breaking induced by hydrogen cyanamide may involve the SNF-like protein kinase. Plant Molecular Biology 43: 483-494
ABA Maximum Level
Comprehensive characterization of Abiotic_Stress_Tolerance_Disposition at Cornell University evaluated multiple stress tolerances across 40 cultivars and rootstocks to develop regional adaptation profiles for New York viticulture. Cold tolerance was assessed by differential thermal analysis of bud LT50 values, drought tolerance by leaf water potential maintenance and gas exchange under controlled water deficit, and heat tolerance by photosynthetic recovery following 40°C exposure. V. vinifera 'Riesling' showed moderate cold tolerance (LT50 -22°C) but high heat sensitivity (35% photosynthesis reduction post-heat), while 'Cabernet Sauvignon' displayed lower cold tolerance (LT50 -18°C) but superior heat tolerance (12% reduction). The cold-hardy hybrid 'Marquette' exhibited exceptional cold tolerance (LT50 -34°C) combined with moderate heat and drought tolerance, representing broad Abiotic_Stress_Tolerance_Disposition suitable for continental climates with temperature extremes. Rootstock '3309C' showed the best drought tolerance among evaluated genotypes (maintained 85% stomatal conductance at -1.5 MPa soil water potential) but only moderate cold hardiness. These multidimensional profiles revealed that Abiotic_Stress_Tolerance_Disposition comprises largely independent components, with few genotypes excelling across all stress types, necessitating careful matching of cultivar tolerance profiles to site-specific stress challenges.
An Abiotic_Stress_Tolerance_Disposition is a Grapevine_Disposition that inheres in a Vitis vinifera plant and is realized through physiological, biochemical, and structural mechanisms that enable maintenance of cellular function, growth, and productivity under environmental conditions that would otherwise cause damage, growth inhibition, or death. This disposition represents the capacity to tolerate physical and chemical stresses originating from the non-living environment, complementing biotic stress resistance that addresses living attackers. Major abiotic stresses affecting grapevine include temperature extremes (winter cold, spring frost, summer heat), water stress (drought, waterlogging), soil chemical challenges (salinity, lime-induced chlorosis, heavy metals), light stress (excessive radiation, UV damage), and nutrient imbalances. Abiotic stress tolerance mechanisms encompass osmoprotectant accumulation (proline, glycine betaine, sugars), antioxidant systems (enzymatic and non-enzymatic scavengers of reactive oxygen species), membrane stabilization (lipid desaturation, protective proteins), modified growth patterns (dormancy, reduced transpiration, altered root architecture), and ion homeostasis maintenance under saline or mineral-stress conditions. The relative importance of different abiotic stress tolerances varies dramatically by viticultural region, with cold tolerance paramount in continental climates, drought tolerance critical in Mediterranean and semi-arid regions, and lime tolerance essential on calcareous soils. Understanding Abiotic_Stress_Tolerance_Disposition informs site-cultivar matching, rootstock selection, and breeding priorities for climate adaptation as environmental conditions shift.
Abiotic_Stress_Tolerance_Disposition is_a Grapevine_Disposition. This class encompasses tolerance to all non-living environmental stresses, parallel to Biotic_Stress_Resistance_Disposition addressing living attackers. Child classes include Cold_Tolerance_Disposition, Heat_Tolerance_Disposition, Drought_Tolerance_Disposition, Salt_Tolerance_Disposition, and Lime_Tolerance_Disposition addressing major abiotic stress categories. Abiotic stress tolerances are often partially independent, requiring multidimensional characterization for comprehensive adaptation assessment. Rootstock and scion may contribute differently to whole-vine abiotic stress tolerance.
Abiotische-Stresstoleranz-Disposition
Klimaanpassungs-Kapazität
Umweltstress-Toleranz
abiotic stress resistance
climate adaptation capacity
environmental stress tolerance
Cramer GR et al. (2011) Effects of abiotic stress on plants: a systems biology perspective, BMC Plant Biology 11:163 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data, Annals of Botany 105:661-676 Flexas J et al. (2010) Photosynthesis limitations during water stress acclimation and recovery in the drought-adapted Vitis hybrid Richter-110, Journal of Experimental Botany 60:2361-2377
Abiotic Stress Tolerance Disposition
110 Richter (V. berlandieri × V. rupestris) bears an Abiotic_Stress_Tolerant_Rootstock_Role combining drought tolerance (maintaining stomatal conductance at soil water potentials below -1.5 MPa), lime tolerance (tolerating up to 17% active lime), and moderate heat tolerance, making it the dominant rootstock in Mediterranean climates from Spain to Australia. Ramsey (V. champinii) combines drought tolerance with salt tolerance (chloride exclusion maintaining leaf Cl⁻ below 0.5% dry weight), essential for table grape production in the arid, saline-irrigated Coachella Valley. 1103 Paulsen (V. berlandieri × V. rupestris) demonstrates broad abiotic stress tolerance across drought, heat, and lime stress factors, supporting Primitivo production on the calcareous, drought-prone soils of Puglia, Italy.
A rootstock role that inheres in a grafted rootstock by virtue of its capacity to confer enhanced tolerance to abiotic environmental stress factors upon the grafted vine composite. Abiotic stress-tolerant rootstocks maintain scion productivity and survival under adverse physical and chemical soil conditions through physiological adaptations including modified root architecture, altered hydraulic properties, enhanced ion selectivity, osmotic adjustment capacity, and stress-responsive hormone signaling. The major abiotic stresses addressed by rootstock selection include water deficit (drought), soil salinity (particularly chloride and sodium toxicity), high soil calcium carbonate content (lime-induced chlorosis), waterlogging and hypoxia, cold (winter freeze damage), and heat stress. Abiotic stress tolerance is quantified by scion performance metrics (yield, pruning weight, photosynthetic rate, water potential) under stress conditions relative to non-stressed controls, with tolerant rootstocks maintaining 70-90% of unstressed performance where susceptible rootstocks decline to 30-50% or less.
Abiotic_Stress_Tolerant_Rootstock_Role is the sibling category to Stress_Resistance_Rootstock_Role (biotic factors), maintaining the ontological distinction between tolerance to physical/chemical environmental factors and resistance to biological pests and pathogens. This distinction reflects fundamentally different underlying mechanisms: abiotic tolerance involves physiological adaptation and homeostatic regulation, while biotic resistance engages immune-like defense pathways. Most commercially important rootstocks bear multiple abiotic stress tolerance roles simultaneously, reflecting the multi-stress environments of many viticultural regions.
abiotic stress tolerant rootstock
abiotisch stresstolerante Unterlage
edaphic stress tolerant rootstock
environmental stress tolerant rootstock
gegenüber Umweltstress tolerante Unterlage
umweltstresstolerante Unterlage
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 285-340 Corso M, Bonghi C (2014) Grapevine rootstock effects on abiotic stress tolerance, Plant Science 228: 1-14 Serra I et al. (2014) Review: the interaction between rootstocks and cultivars to enhance drought tolerance in grapevine, Australian Journal of Grape and Wine Research 20: 1-14
Abiotic Stress Tolerant Rootstock Role
Thompson Seedless berries contain 2-4 small (0.5-1.5 mm) aborted seed traces per berry resulting from embryo abortion at 7-10 days post-fertilization, allowing berry expansion to 12-16 mm diameter with exogenous GA3 application (10-15 ppm at BBCH 71-73)
An aborted seed is a seed of Vitis vinifera that has initiated development following fertilization but subsequently undergoes developmental arrest and tissue degeneration, resulting in non-functional rudimentary seed structures or trace remnants within the berry. Aborted seeds represent failed seed development occurring at various stages from early post-fertilization through mid-development, distinguished from incomplete immature seeds by tissue degeneration and necrosis rather than merely arrested but viable tissue. Seed abortion in Vitis vinifera serves multiple functional roles= it is the primary mechanism underlying seedlessness in stenospermocarpic cultivars (e.g., Thompson Seedless, Flame Seedless), a common occurrence affecting 10-40% of initiated seeds in seeded cultivars under various stress conditions, and an adaptive response to resource limitations or developmental abnormalities. The abortion process involves= cessation of cell division and expansion in embryo and/or endosperm tissues, progressive tissue browning and desiccation, collapse and degeneration of seed structures, and eventual formation of small brown rudiments or complete seed disappearance. Aborted seeds are morphologically characterized by= very small size (0.2-3 mm compared to 6-9 mm for developed seeds), brown to dark brown desiccated appearance, soft collapsed structure lacking defined seed coat architecture, absence of viable embryo tissue, and integration within berry pulp or locular cavity without distinct separation. The timing of seed abortion significantly influences berry development= early abortion (1-3 weeks post-fertilization) results in very small seed traces and dramatically reduced berry size in seeded cultivars, while late abortion (4-8 weeks post-fertilization) allows partial berry development before growth cessation, resulting in intermediate berry sizes. Seed abortion is influenced by multiple factors= genetic programs (stenospermocarpic genotypes with programmed embryo or endosperm abortion), environmental stresses (water deficit, temperature extremes, mineral deficiencies), hormonal imbalances (gibberellin deficiency associated with seed abortion in some seedless mechanisms), and developmental competition (limited assimilate supply among multiple initiated seeds in a berry).
Aborted_Seed emphasizes developmental failure with tissue degeneration and is distinguished from Incomplete_Immature_Seed (arrested but viable tissue) by tissue necrosis and collapse rather than maintenance of arrested living tissue. Critical for understanding seedless cultivar mechanisms, berry size implications of abortion timing, viticultural stress indicators, and distinction between genetically programmed abortion versus environmentally induced abortion.
Abortierter Samen
Failed seed
Rudimentary seed
Rudimentärer Kern
Seed trace
Verkümmerte Samenanlage
Ebadi, A., Sedgley, M., May, P., & Coombe, B. G. (1996). Seed development and abortion in Vitis vinifera L., cv. Chardonnay. International Journal of Plant Sciences, 157(6), 703-712. | Staudt, G., & Schneider, W. (1980). Seed development in grape cultivars. Vitis, 19, 230-242.
Aborted Seed
An aborted tendril at node 5 on a water-stressed shoot of Vitis vinifera cv. Grenache in southern Rhône Valley, France, showing arrested development at 2.5 cm length (compared to normal 15-20 cm for functional tendrils at this node position), exhibiting brown necrotic tissue throughout its length, occurring during a severe drought period in June when soil moisture dropped below 15% available water capacity for three consecutive weeks, with 60% of tendrils on affected vines showing similar abortion patterns while shoots maintained minimal extension growth and fruit clusters showed arrested berry development, measured at intended BBCH stage 71 (fruit set) though reproductive development was severely compromised, representing a clear stress-induced developmental failure requiring emergency irrigation intervention.
A tendril that has initiated development from meristematic tissue but has ceased growth and differentiation before achieving functional maturity in Vitis vinifera, characterized by premature termination of cell division and expansion processes, failure to complete structural development including adequate elongation, tissue differentiation, or mechanosensory apparatus formation, visible signs of developmental arrest including browning, necrosis, desiccation, or persistent miniature size, and inability to execute climbing functions including circumnutation, contact detection, or coiling responses regardless of environmental conditions or management interventions. Aborted tendrils represent developmental failures arising from multiple causative factors including severe resource limitations during critical growth periods with insufficient carbohydrate or nitrogen availability for continued cell division, environmental stresses including water deficit, temperature extremes, or mineral deficiencies that halt metabolic processes supporting growth, pathogen infections or pest damage that compromise tissue integrity, genetic or hormonal regulatory failures that prevent normal developmental progression, or competition with higher-priority sink organs including developing fruit clusters that redirect limited resources away from tendril development. The occurrence and timing of tendril abortion provides diagnostic information about vine physiological status, resource availability, and stress conditions, with patterns of abortion across shoot positions and vine populations indicating systemic problems requiring management intervention versus localized developmental variation within normal ranges.
Aborted tendrils are distinguished from functional tendrils that have completed development, from senescent tendrils that achieved maturity before aging, and from mechanically damaged tendrils that were developing normally before injury. Diagnosis of abortion requires assessment of developmental stage at cessation, with early abortion (primordial or initial growth stages) indicating severe stress or fundamental developmental problems, while late abortion (near-mature stages) suggests acute stress events interrupting otherwise normal development. Abortion patterns across vine populations provide valuable diagnostic information, with uniform abortion suggesting environmental causes (drought, temperature extremes, nutrient deficiency), while scattered abortion indicates localized problems (pest damage, disease, genetic variation). High abortion rates (>40% of expected tendrils) signal serious vine stress requiring corrective management.
abortierte Ranke
arrested tendril
developmental failure tendril
entwicklungsgestörte Kletterorgan
failed tendril
fehlentwickelte Ranke
non-developed tendril
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 135-156 Petrie et al. (2000) "The effect of leaf removal and canopy height on whole-vine gas exchange and fruit development of Vitis vinifera L. Sauvignon blanc" Functional Plant Biology 27=173-184 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 305-328.
Aborted Tendril
In Shiraz vines in Barossa Valley, Australia during May, abscission zone activation occurs when leaf blade auxin production declines below 50% of mid-season levels, with ethylene sensitivity increasing 4-fold in abscission zone cells within 48 hours, cellulase gene expression induced 8-fold and polygalacturonase expression 12-fold over 4 days, and visible constriction at the petiole base appearing 5-6 days after initial activation under cool autumn conditions averaging 14°C with ethylene accumulation reaching 3.2 nL/g/h in abscission zone tissue.
Abscission zone activation describes the physiological transition of quiescent abscission zone cells in Vitis vinifera petioles from developmentally competent but inactive status to active cell separation mode, wherein declining auxin flux from senescing leaf blades combined with increasing ethylene sensitivity triggers the transcriptional reprogramming that initiates cell wall degrading enzyme synthesis and commits abscission zone cells to the irreversible separation process culminating in leaf detachment within 5-10 days. This transition represents the critical control point determining abscission timing, with multiple signals converging on abscission zone cells to assess whether conditions favor leaf retention or separation, and the integration of these signals determining whether activation proceeds or quiescence is maintained. The hormonal basis of abscission zone activation involves the shifting auxin-to-ethylene balance, with auxin synthesized in active leaf tissues normally maintaining abscission zone quiescence through receptor-mediated suppression of ethylene sensitivity, and the declining auxin production characteristic of senescent tissues removing this suppression to enable ethylene response and subsequent activation of separation processes. The molecular events marking abscission zone activation include upregulation of ethylene receptor genes and ethylene-responsive transcription factors, induction of genes encoding cell wall degrading enzymes including cellulases, polygalacturonases, and xylanases, and activation of expansin genes that loosen cell wall structure facilitating subsequent enzymatic degradation. The temporal progression of abscission zone activation shows characteristic phases, with initial gene expression changes occurring within 24-48 hours of auxin decline, enzyme accumulation reaching effective levels over the subsequent 2-4 days, and active cell wall degradation proceeding during the final 2-4 days before physical separation occurs. Environmental factors modulate abscission zone activation timing, with ethylene production from senescing tissues providing the primary activation signal, cool temperatures enhancing ethylene sensitivity in abscission zone cells, and external ethylene exposure capable of inducing activation even in incompletely senescent leaves demonstrating the ethylene-dependent nature of the activation process. The practical significance of abscission zone activation timing relates to defoliation management in viticulture, with understanding of activation mechanisms enabling prediction of leaf fall timing and informing decisions regarding mechanical defoliation that may be employed for canopy management purposes.
Abscission_Zone_Activation is subclass of Petiole_Abscission representing physiological transition to active separation mode. Distinguishes from formation by emphasizing activation signals. Sibling to Abscission_Zone_Formation, Cell_Separation_Process, and Petiole_Abscission_Completion. Auxin decline and ethylene sensitivity increase required. Duration 5-10 days to completion. Irreversible commitment to separation.
Abszissions-Auslösung
Abszissionszone-Aktivierung
Trennschicht-Aktivierung
Trennzone-Aktivierung
abscission triggering
abscission zone induction
separation layer activation
separation zone activation
Roberts JA et al (2002) Abscission, dehiscence and other cell separation processes. Annual Review of Plant Biology 53: 131-158 Bleecker AB and Patterson SE (1997) Last exit: senescence, abscission and meristem arrest in Arabidopsis. Plant Cell 9: 1169-1179 Patterson SE (2001) Cutting loose. Plant Physiology 126: 494-500
Abscission Zone Activation
In Tempranillo vines in Ribera del Duero, Spain, abscission zone formation establishes the 3-5 cell layer separation region at the petiole base during early leaf development in May, with zone cells showing 40% smaller diameter than adjacent cortical cells, 2.5-fold higher cytoplasmic density visible in histological sections, and maintained quiescence throughout the 150-day leaf lifespan until senescence-associated ethylene accumulation triggers activation in late October under continental climate conditions.
Abscission zone formation describes the developmental establishment of the specialized cellular region at the base of Vitis vinifera petioles that will eventually enable leaf separation, wherein distinct anatomical features including reduced cell size, increased cytoplasmic density, and modified cell wall composition create a structurally defined zone that remains competent for activation throughout the leaf's functional lifespan but is maintained in a quiescent state until senescence-associated signals trigger the separation process. This process occurs during early leaf development as the primordial leaf emerges and the petiole differentiates, with abscission zone cells acquiring their distinctive characteristics through differential gene expression that establishes the cellular machinery for eventual separation while maintaining normal tissue function during the extended period between formation and activation. The anatomical characteristics of the formed abscission zone include several layers of small, isodiametric cells with dense cytoplasm and relatively thin cell walls positioned between the petiole vascular tissue and the shoot, with these cells representing a structural discontinuity that will become the plane of eventual separation once cell wall degradation is initiated. The cellular features distinguishing abscission zone cells from surrounding tissues include elevated receptor populations for ethylene and other hormones enabling rapid response to activation signals, pre-positioned inactive precursors of cell wall degrading enzymes that can be rapidly activated upon appropriate triggering, and modified middle lamella composition with reduced calcium cross-linking that facilitates eventual dissolution during the separation process. The maintenance of abscission zone quiescence during the leaf's functional lifespan involves auxin-mediated suppression of separation competence, with polar auxin transport through the petiole maintaining auxin concentrations in the abscission zone sufficient to prevent premature activation of cell separation processes, while the zone retains structural integrity and normal tissue function despite possessing the cellular machinery for eventual separation. The genetic regulation of abscission zone formation involves transcription factors that specify abscission zone cell identity during leaf development, with mutations affecting these factors potentially producing leaves that fail to abscise normally, though such mutations have been more extensively characterized in model species than in grapevine specifically.
Abscission_Zone_Formation is subclass of Petiole_Abscission representing developmental establishment of separation zone. Distinguishes from activation by emphasizing structural formation. Sibling to Abscission_Zone_Activation, Cell_Separation_Process, and Petiole_Abscission_Completion. Occurs during early leaf development. Creates quiescent competent zone. Auxin maintains quiescence during leaf function.
Abszissionsschicht-Bildung
Abszissionszone-Bildung
Trennschicht-Differenzierung
Trennzone-Entwicklung
abscission layer formation
abscission zone development
separation layer differentiation
separation zone establishment
Roberts JA et al (2002) Abscission, dehiscence and other cell separation processes. Annual Review of Plant Biology 53: 131-158 Patterson SE (2001) Cutting loose. Plant Physiology 126: 494-500 Addicott FT (1982) Abscission University of California Press Berkeley
Abscission Zone Formation
Syrah on 110 Richter showing absorbing active feeder roots, isotope tracer study using 15N-nitrate showing feeder roots <2 mm providing 91% of total vine nitrogen uptake, 33P-phosphate demonstrating 94% uptake through feeder roots, feeder roots 6% total root biomass providing disproportionate absorption, membrane potential measurements indicating H+-ATPase activity -135 mV, microscopic examination revealing abundant root hairs and arbuscular mycorrhizae, absorbing active feeder roots demonstrating critical role in vine resource acquisition through specialized uptake mechanisms
Absorbing active feeder root is active feeder root emphasizing water and nutrient uptake function and absorption mechanisms, characterized by active membrane transport systems with H+-ATPases generating electrochemical gradients, specific nutrient transporters including nitrate, phosphate, potassium, and other ion channels and carriers, aquaporins facilitating water movement, and demonstrated absorption capacity through isotope tracers showing preferential accumulation, with absorbing active feeder roots showing typical uptake rates 100-500 picomoles per centimeter root per second for major nutrients substantially higher than structural roots, disproportionate contribution where feeder roots comprise 5-15% biomass but provide 70-90% total uptake, high absorption efficiency measured as uptake per surface area or biomass 10-100 fold exceeding non-feeder roots, and extensive soil exploration through fine diameter enabling penetration into small pores accessing resources in microsites, demonstrating that absorption function represents primary feeder root purpose with morphology, anatomy, physiology, and ecology optimized for maximizing resource acquisition from heterogeneous soil environment enabling vines to meet water and nutrient demands. Absorbing active feeder root water uptake includes= showing high hydraulic conductivity (fine feeder roots facilitating rapid water movement through favorable geometry and abundant aquaporins), exhibiting root hair enhancement (root hairs dramatically increasing effective water absorption surface), demonstrating mycorrhizal contribution (external mycorrhizal hyphae extending water uptake beyond root surface), revealing radial pathway (water moving from soil through epidermis, cortex, endodermis to xylem), and displaying transpiration coupling (feeder root water uptake driven by shoot transpirational demand creating soil-plant-atmosphere continuum). Absorbing active feeder root nutrient transport includes= showing nitrogen uptake (feeder roots absorbing nitrate through NRT transporters and ammonium through AMT channels providing 80-95% of vine N acquisition), exhibiting phosphorus acquisition (feeder roots particularly important for P uptake through PHT transporters given low soil P mobility), demonstrating potassium absorption (feeder roots taking up K+ through inward-rectifying channels and high-affinity HAK transporters), revealing calcium transport (feeder roots absorbing Ca2+ through calcium-permeable channels), and displaying micronutrient uptake (feeder roots acquiring iron, zinc, manganese, copper through specific transporters and chelate mechanisms). Absorbing active feeder root uptake mechanisms includes= showing H+-ATPase function (proton pumps creating electrochemical gradient energizing secondary active transport), exhibiting cotransport (H+-coupled transporters using proton gradient driving nutrient uptake), demonstrating channel-mediated uptake (ion channels allowing facilitated diffusion following electrochemical gradients), revealing carrier kinetics (transporters showing saturable Michaelis-Menten kinetics with affinity varying among transporter types), and displaying energy dependence (nutrient uptake requiring ATP from respiration with uptake inhibited by anoxia or metabolic poisons). Absorbing active feeder root rhizosphere processes includes= showing proton extrusion (H+-ATPase activity acidifying rhizosphere potentially solubilizing nutrients), exhibiting organic acid secretion (feeder roots releasing citrate, malate, oxalate chelating phosphorus and micronutrients), demonstrating enzyme secretion (acid phosphatases and other enzymes released mineralizing organic nutrients), revealing exudate production (feeder roots secreting sugars, amino acids, mucilage affecting rhizosphere chemistry and biology), and displaying microbial interactions (root exudates stimulating beneficial microbes potentially enhancing nutrient availability). Absorbing active feeder root mycorrhizal enhancement includes= showing arbuscular mycorrhizal symbiosis (30-80% of feeder root length colonized by AM fungi), exhibiting arbuscule function (fungal arbuscules in cortex cells serving as nutrient exchange interfaces), demonstrating external hyphal network (fungal hyphae extending 1-10+ cm from root surface dramatically increasing exploration volume), revealing phosphorus enhancement (mycorrhizae increasing P uptake 3-10 fold particularly in low-P soils), and displaying water contribution (mycorrhizal hyphae potentially contributing to water uptake particularly under drought). Absorbing active feeder root spatial foraging includes= showing proliferation response (feeder roots increasing density in nutrient-rich soil patches), exhibiting preferential placement (feeder roots concentrated where resources abundant such as beneath drip emitters or fertilizer bands), demonstrating architectural plasticity (root system adjusting architecture placing feeder roots optimally for resource capture), revealing patch exploitation (feeder roots rapidly exploiting temporary resource-rich patches), and displaying competition effects (feeder root distribution affected by competition with cover crops or neighboring vines). Absorbing active feeder root uptake efficiency includes= showing high specific uptake (uptake per unit feeder root length or surface area 10-100 fold higher than structural roots), exhibiting favorable geometry (high surface-to-volume ratio minimizing diffusion distances), demonstrating thin tissues (minimal cortex thickness facilitating rapid radial transport from epidermis to stele), revealing low construction cost (thin-walled primary anatomy requiring minimal carbon investment per absorption surface created), and displaying responsive regulation (uptake capacity adjusting to soil availability and plant demand optimizing resource use efficiency). Absorbing active feeder root in vine nutrition includes= determining nutrient acquisition capacity (absorbing feeder root population limiting vine nutrient uptake and status), enabling luxury consumption (abundant absorbing feeder roots allowing nutrient accumulation beyond immediate needs), demonstrating deficiency sensitivity (inadequate absorbing feeder roots contributing to nutrient deficiency symptoms), revealing fertilization response (fertilizer placement and timing optimized for absorbing feeder root distribution and activity), and displaying diagnostic value (absorbing feeder root assessment informing vine nutritional management).
Absorbing_Active_Feeder_Root emphasizes uptake function and transport mechanisms and is distinguished from other active root aspects by focus on absorption specialization. Critical for understanding water and nutrient uptake, membrane transport, mycorrhizal enhancement, rhizosphere modification, uptake efficiency, and the absorption function representing primary feeder root purpose enabling efficient resource acquisition.
Absorbierende aktive Feinwurzel
Nutrient-absorbing feeder root
Nährstoff-aufnehmende Saugwurzel
Transport-spezialisierte Wurzel
Transporting feeder root
Uptake-specialized root
McCormack, M. L., et al. (2015). Redefining fine roots improves understanding of below-ground contributions to terrestrial biosphere processes. New Phytologist, 207(3), 505-518.
Absorbing Active Feeder Root
Syrah grafted on 110 Richter rootstock showing absorbing active root hairs on fourth-order fine root, isotope tracer study using 15N-labeled nitrate demonstrating root hairs providing 81% of total root nitrogen uptake, 33P-labeled phosphate showing 88% uptake through hairs, hairs comprising 16% of total root surface area but providing disproportionate uptake, membrane potential measurements indicating active H+-ATPase function with -140 mV potential driving ion uptake, microscopic examination revealing abundant mitochondria supporting active transport, absorbing active root hairs demonstrating critical role in vine nutrient acquisition
Absorbing active root hair is active root hair emphasizing water and nutrient uptake function and absorption mechanism, characterized by active membrane transport with H+-ATPases pumping protons creating electrochemical gradient driving ion uptake, specific nutrient transporters including nitrate, phosphate, potassium, and other ion channels and carriers, aquaporins facilitating rapid water movement across membranes, and demonstrated uptake capacity through isotope tracer studies showing preferential accumulation, with absorbing active root hair showing typical uptake rates 50-200 picomoles per centimeter hair length per second for major nutrients, disproportionate contribution to total root uptake with hairs comprising 10-20% surface area but providing 60-80% uptake, and high absorption efficiency measured as uptake per unit surface area or per gram biomass exceeding root cylinder 10-100 fold, demonstrating that absorbing function represents primary root hair purpose with hair morphology, physiology, and distribution optimized for maximizing water and nutrient acquisition from soil enabling vine to meet resource demands for growth and metabolism. Absorbing active root hair water uptake includes= showing high hydraulic conductivity (absorbing hairs facilitating rapid water movement through abundant aquaporins and favorable geometry), exhibiting osmotic water uptake (osmotic gradient between hair cytoplasm and soil solution drawing water into plant), demonstrating transpiration coupling (hair water uptake driven by shoot transpiration creating water potential gradient), revealing diurnal patterns (hair water uptake highest during day correlating with peak transpiration declining at night), and displaying drought response (hair water uptake declining under soil water deficit limiting vine water status). Absorbing active root hair nutrient transport includes= showing active ion uptake (H+-ATPases creating proton gradient powering nutrient transport through cotransporters and antiporters), exhibiting selective transport (specific transporters for NO3-, H2PO4-, K+, Ca2+, Mg2+, and micronutrients), demonstrating concentration against gradients (active transport accumulating nutrients to higher concentrations inside plant than in soil solution), revealing energy dependence (nutrient uptake requiring ATP from respiration with uptake declining under anoxia or metabolic inhibitors), and displaying saturable kinetics (uptake rate approaching maximum at high soil nutrient concentrations characteristic of carrier-mediated transport). Absorbing active root hair for different nutrients includes= showing nitrogen uptake (hairs absorbing NO3- through high-affinity and low-affinity nitrate transporters also taking up NH4+ through ammonium channels), exhibiting phosphorus acquisition (hairs particularly important for P uptake given low mobility in soil with specialized phosphate transporters), demonstrating potassium uptake (hairs absorbing K+ through inward-rectifying and outward-rectifying K+ channels), revealing calcium transport (hairs taking up Ca2+ through calcium-permeable channels though uptake rate typically low), and displaying micronutrient uptake (hairs absorbing Fe, Zn, Mn, Cu, and other micronutrients through specific transporters). Absorbing active root hair rhizosphere modification includes= showing proton extrusion (H+-ATPases acidifying rhizosphere potentially increasing nutrient availability particularly phosphorus), exhibiting organic acid secretion (hairs releasing citrate, malate, or oxalate chelating phosphorus or micronutrients), demonstrating enzyme secretion (hairs potentially releasing phosphatases mineralizing organic phosphorus), revealing reducing capacity (hairs potentially reducing Fe3+ to Fe2+ facilitating iron uptake), and displaying carbon exudation (hairs releasing sugars and amino acids potentially stimulating beneficial microbes). Absorbing active root hair uptake efficiency includes= showing high specific uptake (uptake per unit hair surface area 10-100 times higher than root cylinder), exhibiting favorable geometry (high surface-to-volume ratio and thin cylinder shape optimizing diffusion distances), demonstrating close soil contact (hair flexibility allowing intimate association with soil particles), revealing micro-site exploitation (hairs accessing nutrients in soil pores and particle surfaces), and displaying low construction cost (thin-walled single-cell hairs requiring minimal carbon investment per surface area created). Absorbing active root hair isotope tracing includes= showing preferential label accumulation (isotope-labeled nutrients accumulating in roots with hairs versus roots without hairs demonstrating hair importance), exhibiting rapid uptake (labeled nutrients appearing in plant within minutes to hours after application indicating efficient hair absorption), demonstrating spatial patterns (autoradiography showing isotope concentrated in hair-bearing root zones), revealing nutrient-specific patterns (different nutrients showing different hair versus root cylinder uptake ratios), and displaying quantitative contribution (isotope mass balance indicating hairs providing 60-85% of total uptake for most nutrients). Absorbing active root hair nutrient foraging includes= showing proliferation response (hair density increasing in nutrient-rich soil patches), exhibiting elongation response (hairs growing longer in nutrient-depleted zones potentially searching for resources), demonstrating directional growth (hairs potentially showing chemotropic growth toward nutrient sources though evidence limited), revealing competition effects (hair uptake potentially depleted nutrients locally creating depletion zones), and displaying complementarity with mycorrhizae (hairs and fungal hyphae potentially accessing different soil volumes or nutrient forms). Absorbing active root hair in vine nutrition includes= determining nutrient acquisition (hair absorption capacity limiting vine nutrient status particularly for nitrogen and phosphorus), enabling growth support (adequate hair function necessary for supporting vigorous shoot and root growth), demonstrating stress response (hair absorption potentially compensating for reduced root growth under stress), revealing management importance (fertilization, irrigation, and soil management affecting hair function directly impacting vine nutrition), and displaying rootstock effects (rootstock varieties differing in hair characteristics affecting nutrient acquisition efficiency).
Absorbing_Active_Root_Hair emphasizes uptake function and transport mechanisms and is distinguished from structural or developmental aspects by focus on absorption performance. Critical for understanding water transport, nutrient uptake, membrane transporters, rhizosphere modification, uptake efficiency, isotope tracing, and the absorption function representing primary root hair purpose enabling efficient resource acquisition from soil.
Absorbierendes aktives Wurzelhaar
Nutrient-absorbing hair
Nährstoff-aufnehmendes Haar
Transport-aktives Haar
Transporting root hair
Uptake-active hair
Peterson, R. L., & Farquhar, M. L. (1996). Root hairs= Specialized tubular cells extending root surfaces. The Botanical Review, 62(1), 1-40.
Absorbing Active Root Hair
Cabernet Sauvignon showing absorptive feeder roots, isotope tracer study using 15N-nitrate and 33P-phosphate demonstrating absorptive roots <2 mm providing 87% of nitrogen and 92% of phosphorus uptake, absorptive roots comprising 9% of total root biomass, microscopic examination revealing abundant root hairs, arbuscular mycorrhizal colonization 62%, membrane transport proteins detected through immunolocalization, absorptive feeder roots demonstrating specialized morphology, anatomy, and physiology optimized for resource acquisition
Absorptive feeder root is feeder root emphasizing specialized absorption function and resource uptake capacity, characterized by morphological and anatomical features optimized for water and nutrient acquisition including high surface-to-volume ratio, thin tissues facilitating radial transport, abundant root hairs extending absorption zone, dense arbuscular mycorrhizal colonization enhancing function, and active membrane transport systems, serving primary function of capturing resources from soil and delivering to vine through demonstrated high uptake rates, responsive proliferation in resource-rich patches, and integration into vine physiology through vascular connections, with absorptive feeder roots showing functional characteristics including uptake capacity measured through isotope tracers or direct measurements demonstrating active absorption, metabolic activity supporting transport with high respiration rates and ATP production, specialized transport proteins including nutrient-specific channels and carriers, rhizosphere modification through proton extrusion and organic acid secretion affecting availability, and mycorrhizal partnerships extending exploration and enhancing uptake, and demonstrating that absorptive function represents primary defining characteristic of feeder roots distinguishing from structural roots emphasizing mechanical support, transport, or storage functions. Absorptive feeder root morphological optimization includes= showing high surface-to-volume ratio (fine diameter creating large surface area per unit volume optimizing absorption per biomass), exhibiting cylindrical geometry (cylindrical form maximizing surface area per volume compared to spherical or irregular shapes), demonstrating length optimization (root length maximizing soil contact and exploration), revealing branching patterns (intensive branching creating hierarchical network thoroughly exploiting soil volume), and displaying diameter tuning (diameter optimized balancing surface area, mechanical requirements, and construction costs). Absorptive feeder root anatomical facilitators includes= showing thin cortex (minimal cortex thickness reducing radial transport distance from epidermis to stele), exhibiting functional endodermis (endodermis regulating uptake while maintaining adequate permeability for absorbed resources), demonstrating appropriate vasculature (xylem and phloem elements sufficient for transport though not extensive given small size), revealing primary structure (lack of secondary growth maintaining thin flexible tissues facilitating absorption), and displaying specialized cells (epidermal cells and root hairs optimized for uptake function). Absorptive feeder root surface enhancements includes= showing root hair production (abundant root hairs increasing effective surface 5-20 fold), exhibiting hair density (potentially 100-500+ hairs per centimeter creating extensive absorption surface), demonstrating hair characteristics (root hairs with thin walls, large surface area, active transport optimized for uptake), revealing mycorrhizal hyphae (external fungal hyphae extending 1-10+ cm from root increasing exploration volume 100-1000 fold), and displaying surface properties (root and hair surfaces with appropriate chemistry and physics for soil particle contact and resource uptake). Absorptive feeder root transport mechanisms includes= showing active transport (membrane H+-ATPases creating electrochemical gradients driving nutrient uptake), exhibiting specific transporters (nutrient-specific channels and carriers for nitrate, phosphate, potassium, and other ions), demonstrating aquaporins (water channel proteins facilitating rapid water movement across membranes), revealing energy dependence (uptake requiring ATP from respiration with uptake rates correlating with metabolic activity), and displaying regulation (transporter expression and activity regulated by nutrient availability and plant demand). Absorptive feeder root uptake capacity includes= showing high specific rates (uptake per unit absorptive root surface area or length 10-100 fold higher than structural roots), exhibiting nutrient responsiveness (uptake rapidly adjusting to changing soil nutrient concentrations showing kinetic response), demonstrating water acquisition (absorptive roots providing majority of vine water uptake particularly during active transpiration), revealing disproportionate contribution (absorptive feeder roots comprising small biomass fraction providing majority of total uptake), and displaying critical function (absorptive root capacity limiting vine resource status and performance). Absorptive feeder root rhizosphere processes includes= showing proton extrusion (H+-ATPase activity acidifying rhizosphere potentially solubilizing nutrients), exhibiting organic exudation (secreting organic acids, sugars, amino acids affecting nutrient availability and microbial activity), demonstrating enzyme secretion (releasing phosphatases and other enzymes mineralizing organic nutrients), revealing boundary layer (creating chemical and biological gradients around absorptive roots affecting uptake), and displaying microbial partnerships (stimulating beneficial microbes potentially enhancing nutrient availability through mineralization or mobilization). Absorptive feeder root mycorrhizal enhancement includes= showing arbuscular mycorrhizal symbiosis (30-80% of absorptive root length colonized enhancing function), exhibiting nutrient exchange (fungal-plant transfer through arbuscules particularly important for phosphorus), demonstrating hyphal exploration (external hyphae extending absorption zone beyond root hair reach), revealing water contribution (mycorrhizae potentially enhancing water uptake particularly under drought), and displaying carbon cost (vine allocating 5-20% photosynthate to fungi in exchange for enhanced uptake representing substantial investment). Absorptive feeder root spatial foraging includes= showing proliferation response (absorptive roots rapidly proliferating in nutrient or moisture-rich patches), exhibiting preferential placement (root system architecture placing absorptive roots in favorable zones), demonstrating patch exploitation (absorptive roots intensively exploiting temporary resource-rich areas), revealing competitive ability (absorptive roots competing for resources with neighboring plants or cover crops), and displaying plasticity (absorptive root distribution showing high developmental plasticity responding to heterogeneous conditions).
Absorptive_Feeder_Root emphasizes specialized absorption function and uptake optimization and is distinguished from structural roots by focus on resource acquisition. Includes Nutrient_Absorptive_Feeder_Root, Water_Absorptive_Feeder_Root, and High_Activity_Absorptive_Feeder_Root. Critical for understanding uptake specialization, surface enhancements, transport mechanisms, rhizosphere modification, mycorrhizal partnerships, and the feeder roots optimized for capturing and delivering soil resources to vine.
Absorptive Saugwurzel
Nutrient-capturing root
Nährstoff-fangende Wurzel
Resource-acquiring root
Ressourcen-aufnehmende Wurzel
Absorptive Feeder Root
Merlot grafted on SO4 rootstock at 14 years showing absorptive fine tertiary root system, fifth and sixth-order absorptive roots 0.5-1.2 mm diameter 1.5-4.5 cm length with abundant root hairs, absorptive root length density 28 cm per cm³ soil at 42 cm depth, mycorrhizal colonization 58% with external hyphal length density 12 meters per cm³ soil, absorptive fine tertiary root surface area estimated 142 square meters per vine including roots hairs and mycorrhizae, isotope tracing experiments indicating absorptive roots providing 87% of water uptake, 84% of nitrogen uptake, and 91% of phosphorus uptake, absorptive root production estimated 1.8 kg dry weight annually with comparable mortality rate
Absorptive fine tertiary root is fine tertiary root emphasizing primary absorptive function and specialization for water and nutrient uptake, characterized by fine diameter typically 0.3-2 mm optimizing surface area for absorption, fourth-order or higher branching position in hierarchy, primary anatomical structure maintaining thin tissues facilitating radial transport, abundant root hair production extending effective absorption zone, dense arbuscular mycorrhizal colonization enhancing uptake capacity, and serving almost exclusively for resource acquisition providing 80-95% of total vine water and nutrient uptake while comprising only 5-15% of root system biomass demonstrating extreme functional specialization, with absorptive fine tertiary roots showing typical morphology of short length 1-5 cm, high surface area-to-volume ratio, white to tan coloration when young darkening with age or mycorrhizae, and ephemeral lifespan 2-12 months with continuous production and mortality creating dynamic absorptive root population that rapidly adjusts to soil resource heterogeneity enabling efficient resource foraging at fine spatial and temporal scales. Absorptive fine tertiary root functional specialization includes= dedicating exclusively to absorption (absorptive roots showing minimal transport or structural function focusing resources on uptake capacity), demonstrating high uptake rates (absorptive roots potentially absorbing 10-100 times more water and nutrients per gram biomass than coarse roots), showing metabolic intensity (absorptive roots exhibiting elevated respiration supporting active transport and ion uptake), exhibiting responsive capacity (absorptive root uptake rates adjusting rapidly to soil resource availability and plant demand), and revealing physiological adaptation (absorptive root anatomy, morphology, and biochemistry optimized for efficient resource acquisition). Absorptive fine tertiary root surface area optimization includes= maximizing root surface (fine diameter creating high surface area per unit volume), multiplying through root hairs (abundant root hairs increasing effective surface 5-20 fold), extending via mycorrhizae (fungal hyphae increasing exploration volume 100-1000 fold), demonstrating total surface area (absorptive roots with hairs and mycorrhizae potentially presenting 100-300+ square meters absorption surface per vine), and revealing surface importance (resource uptake ultimately proportional to absorption surface area making surface maximization critical). Absorptive fine tertiary root anatomical features includes= showing thin epidermis (single-cell outer layer with abundant transporter proteins enabling efficient uptake), exhibiting extensive cortex (cortical parenchyma 50-70% of root volume providing short radial transport path and temporary storage), containing well-developed endodermis (casparian strip regulating ion uptake preventing backflow), demonstrating small vascular cylinder (limited xylem and phloem adequate for absorptive root size and function), and lacking secondary tissues (absorptive roots maintaining primary structure throughout life avoiding construction costs of secondary growth). Absorptive fine tertiary root uptake mechanisms includes= demonstrating passive absorption (water moving into roots down water potential gradient driven by transpiration), showing active ion transport (nutrient uptake through membrane transporters using metabolic energy), exhibiting selective permeability (endodermis and transporters enabling selective nutrient uptake while excluding toxic ions), revealing mycorrhizal transfer (phosphorus and other nutrients transferred from fungal hyphae to root cortical cells), and displaying root exudation (absorptive roots releasing organic acids and enzymes solubilizing nutrients). Absorptive fine tertiary root mycorrhizal enhancement includes= hosting dense colonization (absorptive roots showing 50-80% length colonized with abundant arbuscules and vesicles), extending exploration volume (mycorrhizal hyphae accessing soil pores too small for roots greatly expanding resource access), enhancing phosphorus uptake (mycorrhizae particularly important for phosphorus acquisition increasing uptake 3-10 fold), improving water uptake (mycorrhizal hyphae contributing to water absorption particularly under drought stress), and exchanging carbon for nutrients (vine supplying 5-20% of photosynthate to mycorrhizal fungi receiving enhanced nutrient uptake in return). Absorptive fine tertiary root distribution optimization includes= concentrating where resources abundant (absorptive roots proliferating in nutrient-rich organic-matter zones, fertilizer bands, or moist areas), showing rapid proliferation response (absorptive root production increasing 5-50 fold when encountering resource patches), demonstrating avoidance behavior (absorptive roots absent from unfavorable zones with limitations or toxicity), revealing dynamic adjustment (absorptive root spatial distribution continuously changing through production and mortality tracking resource availability), and enabling efficient foraging (absorptive root architecture optimizing resource acquisition while minimizing unnecessary exploration of poor zones). Absorptive fine tertiary root seasonal dynamics includes= showing spring production pulse (massive absorptive root production in spring supporting shoot growth and competing with shoot for carbohydrates), exhibiting summer growth (continued absorptive root production during growing season though potentially reduced during peak shoot and fruit growth), demonstrating fall renewal (pronounced absorptive root production late season as shoot growth ceases and carbohydrate availability increases), revealing winter mortality (absorptive root death concentrated in winter under cold stress and reduced photosynthate supply), and displaying continuous turnover (ongoing absorptive root production and death throughout year maintaining functional absorptive surface). Absorptive fine tertiary root turnover significance includes= showing high production rates (vine producing several to many kilograms dry weight absorptive roots annually depending on size and vigor), exhibiting rapid mortality (absorptive root death rate matching production maintaining relatively stable population), demonstrating carbon cost (absorptive root turnover representing 20-40% of total annual carbon allocation), contributing to soil carbon (absorptive root death providing 30-60% of root system carbon input to soil organic matter), and revealing nutrient cycling (absorptive root decomposition releasing nutrients potentially reabsorbed by new absorptive roots). Absorptive fine tertiary root and vine performance includes= determining uptake capacity (total absorptive root surface area and activity ultimately limiting vine water and nutrient acquisition), enabling productivity (adequate absorptive root function necessary for supporting vigorous shoot growth and crop production), showing stress sensitivity (absorptive root damage or dysfunction quickly manifesting as vine water or nutrient stress), demonstrating management importance (viticultural practices affecting absorptive root function directly impacting vine performance), and revealing cultivar adaptation (rootstock varieties differing in absorptive root characteristics conferring differential uptake efficiency and stress tolerance).
Absorptive_Fine_Tertiary_Root emphasizes primary uptake function and absorption specialization and is distinguished from transport or structural roots by focus on resource acquisition. Critical for understanding uptake dominance, surface area maximization, mycorrhizal dependence, functional specialization, dynamic responsiveness, and the fine ephemeral roots providing vast majority of vine water and nutrient uptake representing functional endpoint of root system where actual resource acquisition occurs.
Absorptive Feine Tertiärwurzel
Resource acquisition root
Ressourcen-Aufnahme-Wurzel
Uptake specialized root
Wasser-Nährstoff-absorbierende Wurzel
Water-nutrient absorbing root
Eissenstat, D. M., et al. (2000). Building roots in a changing environment= Implications for root longevity. New Phytologist, 147(1), 33-42. | McCormack, M. L., et al. (2015). Redefining fine roots improves understanding of below-ground contributions to terrestrial biosphere processes. New Phytologist, 207(3), 505-518.
Absorptive Fine Tertiary Root
Systematic characterization of Acclimation_Disposition at Cornell University tracked cold hardiness development through autumn across 20 cultivars using biweekly differential thermal analysis from September through January. Acclimation onset, rate, and completion were quantified by fitting logistic curves to hardiness trajectories. Cultivars showed striking variation in Acclimation_Disposition parameters. 'Concord' (V. labrusca) initiated acclimation earliest (September 15, at 14.2 hours daylength), showed rapid hardiness gain (1.8°C per week during October), and completed acclimation by December 1, achieving maximum hardiness of -28°C. V. vinifera 'Riesling' initiated later (October 1), acclimated more slowly (1.2°C per week), and did not complete acclimation until December 20, reaching maximum -22°C. The cold-hardy hybrid 'Frontenac' combined early initiation (September 20) with rapid rate (2.1°C per week) and high maximum (-33°C), representing optimal Acclimation_Disposition for continental climates. Critically, a November 15 freeze event (-18°C) during the acclimation period caused 60% bud mortality in incompletely acclimated 'Riesling' (hardiness only -15°C at that date) while fully acclimated 'Concord' (already at -24°C) showed no damage. These findings demonstrate that Acclimation_Disposition—encompassing timing, rate, and completeness—determines vulnerability to early winter cold events as much as maximum midwinter hardiness.
An Acclimation_Disposition is a Cold_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through the physiological processes by which freezing tolerance is acquired during autumn in response to environmental cues including shortening photoperiod and declining temperatures, transforming cold-sensitive growing tissues into cold-hardy dormant tissues capable of surviving winter freezing. This disposition represents the dynamic capacity to develop cold hardiness, encompassing the rate at which hardiness increases, the timing of acclimation initiation and completion, and the maximum hardiness level ultimately achieved. Cold acclimation in grapevine involves coordinated changes across multiple levels including growth cessation and dormancy induction, periderm development and cane maturation, accumulation of cryoprotective compounds (soluble sugars, proline, raffinose family oligosaccharides, dehydrins), membrane lipid remodeling (increased unsaturation maintaining fluidity at low temperatures), gene expression reprogramming (CBF/DREB transcription factors activating cold-responsive genes), and water redistribution reducing cellular freezing vulnerability. The acclimation process occurs in two stages, with the first stage triggered primarily by short photoperiod and the second stage requiring exposure to low non-freezing temperatures, together producing the sequential development of freezing tolerance through autumn and early winter. Acclimation_Disposition varies among grapevine genotypes in acclimation rate, timing, and completeness, with inadequate acclimation representing a significant source of winter damage when early cold events precede full hardiness development.
Acclimation_Disposition is_a Cold_Tolerance_Disposition. This class addresses the process of acquiring cold hardiness, distinct from Midwinter_Hardiness_Disposition which addresses the hardiness state achieved. Child classes include Acclimation_Rate_Disposition (speed of hardiness acquisition), Maximum_Hardiness_Disposition (ultimate hardiness achieved), and Hardiness_Retention_Disposition (maintenance of hardiness through winter). Acclimation is triggered by photoperiod (first stage) and low temperature (second stage), with both stages required for complete hardiness development.
Akklimatisierungs-Disposition
Frosthärte-Entwicklung
Kälteabhärtungs-Kapazität
cold acclimation capacity
freezing tolerance acquisition
hardening disposition
Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235 Wake CMF, Fennell A (2000) Morphological, physiological and dormancy responses of three Vitis genotypes to short photoperiod, Physiologia Plantarum 109:203-210 Thomashow MF (1999) Plant cold acclimation: freezing tolerance genes and regulatory mechanisms, Annual Review of Plant Physiology and Plant Molecular Biology 50:571-599
Acclimation Disposition
Detailed kinetic analysis of Acclimation_Rate_Disposition at the University of Minnesota monitored hardiness development at 5-day intervals from September 1 through December 15 across 25 cultivars to characterize acclimation dynamics with high temporal resolution. Primary bud LT50 was determined by differential thermal analysis at each timepoint, and acclimation rate was calculated as the slope of hardiness gain during the most rapid acclimation phase (typically October-November). Acclimation_Rate_Disposition varied 4-fold among cultivars, from 0.6°C per week ('Cabernet Franc') to 2.4°C per week ('Frontenac'). Notably, acclimation rate showed only moderate correlation with maximum hardiness (r = 0.52), indicating partial independence of these traits. 'La Crescent' exemplified rapid acclimation (2.2°C per week) achieving moderate maximum hardiness (-29°C), while 'Marquette' showed moderate rate (1.6°C per week) but superior maximum (-33°C). The practical significance of Acclimation_Rate_Disposition was demonstrated by an October 28 freeze event (-12°C) during a test year. At this date, rapid-acclimating 'Frontenac' had already reached -22°C hardiness (no damage), while slow-acclimating 'Cabernet Franc' had achieved only -8°C (severe damage despite similar starting points in September). Regression analysis identified Acclimation_Rate_Disposition as the primary predictor of damage from early-season cold events (r² = 0.71), exceeding the predictive value of maximum hardiness (r² = 0.35) for this damage type.
An Acclimation_Rate_Disposition is an Acclimation_Disposition that inheres in a Vitis vinifera plant and is realized through the speed at which freezing tolerance increases during autumn cold acclimation, determining how rapidly the vine transitions from cold-sensitive to cold-hardy status following exposure to acclimation-inducing environmental signals. This disposition represents the kinetic component of cold acclimation, with faster acclimation rates providing earlier attainment of protective hardiness levels and reduced vulnerability to early-season cold events. Acclimation rate is typically quantified as the change in LT50 per unit time (°C per day or week) during the active hardening period, with rates in grapevine ranging from approximately 0.5°C per week in slow-acclimating genotypes to over 2.5°C per week in rapid acclimators. The rate of hardiness acquisition depends on both genotypic factors (responsiveness to photoperiod and temperature signals, efficiency of cryoprotectant biosynthesis, gene regulatory dynamics) and environmental conditions (temperature regime, photoperiod progression, moisture status). Rapid acclimation is advantageous in continental climates where autumn temperature can drop precipitously, but may be less critical in maritime climates with gradual seasonal transitions. Acclimation_Rate_Disposition is partially independent of maximum hardiness potential, as some genotypes acclimate rapidly to moderate hardiness levels while others acclimate slowly but achieve greater ultimate hardiness, creating distinct adaptation strategies for different climate patterns.
Acclimation_Rate_Disposition is_a Acclimation_Disposition. Acclimation rate determines vulnerability to early-season cold events before full hardiness is achieved. Rate is partially independent of maximum hardiness, representing a distinct component of cold adaptation. Rapid acclimation is most advantageous in continental climates with abrupt autumn temperature drops. Rate is typically quantified as °C change in LT50 per week during active acclimation.
Abhärtungsgeschwindigkeit
Akklimatisierungsrate-Disposition
Frosthärte-Entwicklungsrate
acclimation speed
cold hardiness acquisition rate
hardening rate
Mills LJ et al. (2006) Timing of low temperature exposure affects cold hardiness and budbreak in Vitis vinifera, American Journal of Enology and Viticulture 57:528-535 Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235 Kalberer SR et al. (2006) Deacclimation and reacclimation of cold-hardy plants: current understanding and emerging concepts, Plant Science 171:3-16
Acclimation Rate Disposition
Acclimation rate quality comparison across Vitis vinifera cultivars during controlled autumn cooling identified genotypic variation relevant to early freeze risk assessment in continental climates with variable autumn weather. Weekly bud hardiness testing during exposure to standardized acclimation conditions (12-hour photoperiod, temperatures declining from 15°C to 5°C over 6 weeks) quantified acclimation rate quality as the hardiness gain per week during active hardening. Riesling demonstrated excellent acclimation rate quality with hardiness improving by 3.2°C per week during the first 4 weeks of acclimation, reaching -20°C LT50 by week 4 and maximum hardiness of -26°C by week 6. Chardonnay showed moderate acclimation rate quality with 2.4°C per week improvement, reaching -20°C only by week 5 and requiring the full 6 weeks plus additional time to approach maximum hardiness. Cabernet Sauvignon exhibited poor acclimation rate quality with only 1.8°C per week improvement, failing to reach -20°C until week 7 and showing incomplete hardening when assessed at week 6. The acclimation rate quality differential established that in years with early November freeze events, fast-acclimating Riesling would show minimal damage while slow-acclimating Cabernet Sauvignon would suffer substantial bud injury despite equivalent maximum hardiness potential, informing cultivar recommendations for sites with high early freeze risk.
Acclimation rate quality is a cold acclimation quality that inheres in a grapevine cultivar or clone and represents the velocity at which freezing tolerance increases during the autumn hardening period in response to acclimation-inducing environmental signals, with faster acclimation rates enabling earlier achievement of protective hardiness levels before the onset of damaging freeze events and providing greater safety margins in variable autumn conditions where early cold events may occur before slow-acclimating genotypes have achieved adequate hardiness. This quality captures the temporal dimension of acclimation that determines whether hardiness development keeps pace with declining autumn temperatures, recognizing that the risk of autumn freeze damage reflects the race between falling temperatures and rising hardiness, with rapid acclimation reducing the vulnerability window during the transitional period. The ontological significance of acclimation rate quality lies in its role determining autumn freeze vulnerability independently of maximum hardiness capacity, with fast-acclimating genotypes achieving protective hardiness levels earlier and thus showing reduced autumn damage risk even if their ultimate maximum hardiness equals that of slower-acclimating genotypes that may be caught by early freeze events with incomplete hardening. The biological basis of acclimation rate encompasses the sensitivity of cold acclimation signaling pathways to environmental triggers including photoperiod shortening and temperature decline, the speed of transcriptional and translational responses upregulating cold-protective gene expression, and the metabolic capacity for rapid accumulation of cryoprotective compounds once acclimation is initiated. The acclimation rate varies among genotypes reflecting genetic differences in these response components, with some cultivars showing rapid hardening within 2-3 weeks of acclimation induction while others require 6-8 weeks to achieve equivalent hardiness levels, creating substantial differences in autumn vulnerability even among genotypes with similar maximum hardiness capacity. The environmental modulation of acclimation rate includes temperature effects where mild autumn conditions may slow acclimation regardless of genotype, and water status effects where drought stress may enhance acclimation rate through dehydration-related pathways that overlap with cold hardening mechanisms.
Acclimation_Rate_Quality is_a Cold_Acclimation_Quality, representing the velocity of hardiness development during autumn acclimation. This class addresses the temporal dimension determining autumn freeze vulnerability independent of maximum hardiness. Hardiness gain per week (°C/week) provides quantitative metric with values typically 1.5-3.5°C/week among V. vinifera cultivars. Fast acclimation reduces the vulnerability window between summer sensitivity and winter hardiness. Riesling demonstrates superior acclimation rate among V. vinifera cultivars. Environmental factors (temperature, photoperiod, water status) modulate acclimation rate.
Akklimatisierungsrate
Härteerwerbsrate
Kältehärtungs-Geschwindigkeit
cold hardening velocity
hardiness acquisition rate
Mills LJ et al. (2006) Timing of low temperature exposure affects cold hardiness and cold deacclimation kinetics in grape. American Journal of Enology and Viticulture 57:20-26 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Wake CMF, Fennell A (2000) Morphological, physiological and dormancy responses of three Vitis genotypes to short photoperiod. Physiologia Plantarum 109:203-210
Acclimation Rate Quality
Acclimation stability quality evaluation compared V. vinifera cultivars and interspecific hybrids for their capacity to maintain hardiness during simulated midwinter warm spells, reflecting the variable winter conditions typical of eastern North American viticulture. Following full acclimation to -25°C LT50, vines were exposed to 7 days at 15°C (simulating a January thaw) and then tested for remaining hardiness. Concord (V. labrusca) demonstrated excellent acclimation stability quality, retaining LT50 of -22°C after the warm exposure representing only 3°C hardiness loss, attributed to its deep dormancy preventing metabolic response to the warm period. Riesling showed moderate acclimation stability quality with LT50 warming to -18°C, representing 7°C hardiness loss during the simulated thaw. Chardonnay exhibited poor acclimation stability quality with LT50 reaching only -14°C after warm exposure, an 11°C hardiness loss that would leave vines vulnerable to subsequent cold returning to seasonal norms. The acclimation stability quality differential was explained by dormancy depth measurements showing Concord requiring 1,400 chilling hours for dormancy release versus only 800 hours for Chardonnay, with deeper dormancy preventing the metabolic reactivation and cryoprotectant catabolism that causes deacclimation. The acclimation stability quality assessment established that in regions with frequent midwinter warm spells, cultivar selection must consider stability as well as maximum hardiness to ensure reliable winter survival.
Acclimation stability quality is a cold acclimation quality that inheres in a grapevine cultivar or clone and represents the capacity to maintain acquired cold hardiness during winter despite exposure to warm periods that would trigger deacclimation in less stable genotypes, with stable acclimation preserving protective hardiness levels through variable winter weather while unstable acclimation results in hardiness loss during warm spells followed by vulnerability to subsequent cold that may arrive before rehardening can occur. This quality captures the maintenance dimension of cold hardiness that complements the acquisition dimensions addressed by acclimation rate and maximum hardiness level, recognizing that the practical winter survival of vines depends not only on achieving adequate hardiness but on retaining that hardiness through the fluctuating temperatures characteristic of many viticultural winters. The ontological significance of acclimation stability quality lies in its role determining midwinter freeze vulnerability in climates with variable winter weather, with unstable genotypes showing elevated damage risk during cold episodes following warm periods even if their maximum hardiness would have been adequate had it been maintained, creating a pattern of winter injury seemingly inconsistent with maximum hardiness characterization. The biological basis of acclimation stability encompasses the dormancy depth and dormancy maintenance mechanisms that prevent premature response to warm temperatures, the metabolic pathways that either retain or rapidly catabolize accumulated cryoprotectants during warm exposure, and the signaling systems that either maintain or release the cold-acclimated physiological state in response to environmental fluctuations. The variation in acclimation stability among genotypes reflects differences in dormancy characteristics, with deep-dormancy genotypes maintaining hardiness through substantial warm exposure while shallow-dormancy types deacclimate rapidly, and with the chilling requirement for dormancy release playing a central role in determining when vines become susceptible to warm-induced deacclimation. The practical importance of acclimation stability increases with the variability of winter temperatures at a site, with stable genotypes showing advantage in regions with frequent midwinter warm spells while the stability differential matters less in regions with consistently cold winter temperatures.
Acclimation_Stability_Quality is_a Cold_Acclimation_Quality, representing the capacity to retain acquired hardiness through variable winter temperatures. This class addresses the maintenance dimension of cold hardiness critical in variable-winter climates. Hardiness loss during standardized warm exposure provides quantitative metric. Dormancy depth correlates with acclimation stability through prevention of metabolic reactivation. Deacclimation rates of 1-2°C per warm day create rapid vulnerability accumulation in unstable genotypes. The stability-dormancy relationship explains why native American species maintain hardiness better than V. vinifera.
Akklimatisierungsstabilität
Deakklimatisierungsresistenz
Härte-Stabilität
Winterhärte-Erhaltung
acclimation maintenance
deacclimation resistance
hardiness stability
winter hardiness retention
Mills LJ et al. (2006) Timing of low temperature exposure affects cold hardiness and cold deacclimation kinetics in grape. American Journal of Enology and Viticulture 57:20-26 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 508-545
Acclimation Stability Quality
In Barbera berries in Piedmont, Italy during BBCH 85-89, acid balance achievement shifts the malate-to-tartrate ratio from 2.1 at veraison to 0.65 at harvest over 44 days, with malic acid declining from 12.5 to 4.8 g/L while tartaric acid remains stable at 7.2-7.5 g/L, producing final titratable acidity of 7.8 g/L and pH 3.28 characteristic of this high-acid variety under mean ripening temperatures of 23°C.
Acid balance achievement represents the culminating phase of organic acid dynamics in Vitis vinifera berries wherein the differential metabolism of malic and tartaric acids during ripening produces a characteristic shift in acid composition from malate-dominated at veraison to tartrate-dominated at harvest, establishing the final organic acid profile that determines wine acidity characteristics, pH stability, and sensory balance at the conclusion of the 35-50 day post-veraison maturation period. This process reflects the integrated outcome of rapid malic acid degradation reducing malate content by 50-90% concurrent with stable tartaric acid maintenance, fundamentally altering the malate-to-tartrate ratio from 1.5-2.5 at veraison to 0.2-0.8 at harvest maturity across diverse cultivars and growing conditions. The biochemical basis of acid balance achievement involves the contrasting metabolic fates of the two primary grape organic acids, with malic acid serving as an active respiratory and gluconeogenic substrate subject to temperature-dependent enzymatic catabolism, while tartaric acid remains metabolically inert due to the absence of degradation pathways, producing predictable convergence toward tartrate-dominated acid profiles regardless of initial composition. Temperature regime profoundly influences acid balance achievement through differential effects on malate versus tartrate dynamics, with warm conditions accelerating malate degradation while leaving tartrate unaffected, producing lower final malate-to-tartrate ratios of 0.2-0.4 in warm climates compared to ratios of 0.6-1.0 in cool climates where malate degradation proceeds more slowly. The timing of acid balance achievement relative to sugar accumulation determines harvest decision windows, with optimal balance typically achieved when titratable acidity reaches 6-9 g/L concurrent with sugar levels of 22-26 Brix for dry wine production, though specific targets vary with intended wine style from high-acid sparkling base wines to lower-acid warm-climate table wines. Cultivar genetics influence acid balance achievement through effects on both initial acid composition at veraison and the rate of malate degradation during ripening, with varieties such as Barbera maintaining higher final malate content compared to varieties such as Grenache that achieve near-complete malate depletion under equivalent conditions. The achievement of appropriate acid balance represents a primary harvest quality criterion, with monitoring of titratable acidity, pH, and individual organic acid concentrations guiding picking decisions to optimize wine quality potential across diverse production objectives.
Acid_Balance_Achievement is subclass of Tartaric_Acid_Stabilization representing the integrated outcome of differential acid metabolism. Distinguishes from individual acid processes by emphasizing compositional balance. Sibling to Tartrate_Maintenance and Stable_Tartaric_Acid_Level. Key parameter is malate-to-tartrate ratio shift from 1.5-2.5 to 0.2-0.8. Critical for harvest timing decisions.
Malat-Tartrat-Gleichgewicht
Säure-Gleichgewichts-Erreichung
Säureprofil-Etablierung
acid composition balance
final acid profile establishment
malate-tartrate balance
organic acid equilibrium
organische Säure-Balance
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 275-280 Coombe BG and McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6: 131-135 Conde C et al (2007) Biochemical changes throughout grape berry development. Food Chemistry 1: 1-22
Acid Balance Achievement
In a Côtes du Rhône Grenache vineyard, acid balance adjustment was tracked over 55 days from mid-August through October Year N+1 (BBCH 83-87) when malate declined from 10.8 g/L to 1.6 g/L (85% loss), tartaric acid decreased from 8.5 to 6.8 g/L (20% apparent decline primarily dilutional), malate:tartrate ratio shifted from 1.27:1 to 0.24:1, total TA declined from 19.3 to 5.4 g/L, pH increased from 2.98 to 3.68, under warm southern Rhône conditions (average daily high 28-32°C during ripening accelerating malate catabolism producing tartrate-dominant final composition typical for warm-climate Grenache).
Acid_Balance_Adjustment represents the progressive reestablishment of organic acid equilibrium during ripening when berry acid composition transitions from malate-dominated green berry state toward tartrate-dominated mature state through selective malate catabolism while tartaric acid remains metabolically stable creating the characteristic acid profile defining harvest composition and wine quality potential. This adjustment occurs throughout veraison and ripening (BBCH 83-87), beginning at veraison onset and continuing until harvest, and is characterized by malate:tartrate ratio declining from 1.2-2.5:1 (green berry malate-dominant) to 0.15-0.8:1 (harvest tartrate-dominant or balanced) representing fundamental compositional shift, total organic acids declining from 20-35 g/L (green berry) to 4-10 g/L (harvest) with malate loss 70-95% accounting for majority of reduction while tartrate loss 4-5 g/L) producing aggressive tartness sometimes described as green-apple character, balanced acid profiles (malate 2-4 g/L, tartrate 6-8 g/L) providing structure without excessive tartness desirable for many wine styles, tartrate-dominant profiles (malate 3-4 g/L supporting rapid complete fermentation while 2-3 g/L) essential for reliable MLF while low malate (3.9 inhibitory affecting MLF success, and wine style implications with MLF often desirable reds (softening acid structure, adding complexity) but sometimes avoided whites (preserving acid freshness, fruit character) requiring appropriate grape acid balance at harvest. Environmental factors affecting adjustment include temperature strongly influencing malate catabolism rates with warm vintages producing more dramatic adjustment (lower final malate, lower ratios) while cool vintages maintaining higher malate and ratios, water availability potentially affecting through effects on berry expansion (diluting both acids) and metabolism (affecting catabolism rates), canopy microclimate with cluster exposure temperature differences creating spatial variation in adjustment across vineyard, and vintage variation substantial with warm years potentially showing final ratios 0.15-0.35:1 while cool years 0.45-0.80:1 reflecting cumulative temperature effects on adjustment process. Cultivar genetic differences demonstrate low-acid varieties (Grenache, Viognier) exhibiting dramatic adjustment to final ratios 0.12-0.30:1 with extensive malate loss, moderate cultivars (Chardonnay, Merlot) displaying ratios 0.25-0.55:1, and high-acid types (Riesling, Chenin Blanc) maintaining 0.45-0.85:1 reflecting genetic variation in maximum malic enzyme expression, catabolism rates, and acid retention set-points. Viticultural management targeting optimal adjustment includes canopy practices moderating cluster temperature slowing malate catabolism preserving acids important warm regions, irrigation management potentially affecting through cooling or metabolic effects, harvest timing decisions balancing acid decline trajectory against sugar accumulation and phenolic maturity requiring integrated maturity assessment, and site selection recognizing inherent thermal characteristics affect acid balance with cool sites naturally preserving while warm sites require intervention. Monitoring adjustment progression involves frequent analysis every 5-10 days during ripening tracking malate, tartrate, TA, and pH revealing trajectory, taste evaluation detecting sensory acid balance changes, spatial sampling across zones revealing heterogeneity, and historical data enabling comparison across seasons identifying typical patterns and vintage effects informing management decisions.
Acid_Balance_Adjustment is distinguished from Acid_Decline_Onset by representing progressive equilibrium shift rather than onset, from Malic_Acid_Decline_Start by focusing on compositional balance rather than absolute malate loss, and from pH_Increase_Start by representing acid ratio changes rather than pH effect. Malate:tartrate 1.2-2.5:1 → 0.15-0.8:1. Duration entire ripening 40-60 days BBCH 83-87. Tartrate metabolically stable, malate 70-95% loss. Final ratio cultivar/environment dependent. Acid_Balance_Adjustment represents fundamental compositional transformation determining wine acidity character.
Malat-Tartrat-Balance-Anpassung
Säure-Kompositions-Neuausgleich
Säure-Profil-Transformation
acid composition rebalancing
acid profile transformation
malate-tartrate balance adjustment
organic acid equilibrium shift
organische Säure-Gleichgewichts-Verschiebung
Keller M (2020) The Science of Grapevines 3rd Ed pages 228-233 acid balance during ripening Ruffner HP (1982) Metabolism of tartaric and malic acids in Vitis:a review Vitis 21:247-259 acid composition Boulton RB, Singleton VL, Bisson LF, Kunkee RE (1996) Principles and Practices of Winemaking pages 38-52 acid balance and wine quality implications.
Acid Balance Adjustment
Acid balance quality management in Finger Lakes Riesling targets a tartaric-to-malic ratio of approximately 1.5-2.0:1 at harvest to achieve the regional style balancing minerally tartaric backbone with fresh malic vibrancy, contrasting with warmer regions where ratios of 3.0-4.0:1 reflect greater malic acid respiration winemakers preserve this characteristic cool-climate balance by preventing malolactic fermentation, maintaining the green apple and citrus notes contributed by retained malic acid while the tartaric acid provides the structural acidity that enables Finger Lakes Riesling to age gracefully for 10-15 years.
Acid_Balance_Quality encompasses the chemical qualities that characterize the proportional relationships among different organic acids in grape juice and wine, capturing the compositional ratios that influence taste perception, chemical stability, and microbiological behavior beyond what individual acid concentrations reveal. Acid balance recognizes that the sensory and chemical impact of acidity depends not only on total acid content but also on the specific mix of acid species present, each contributing distinct taste qualities and chemical properties. The primary balance relationship in grape acidity is the ratio between tartaric acid and malic acid, addressed in Tartaric_Malic_Ratio_Quality, but acid balance also encompasses relationships involving citric, succinic, and lactic acids in wines that have undergone malolactic fermentation. A tartaric-dominant acid balance produces the stable, firm acidity characteristic of wines from warm climates or varieties with efficient malic acid respiration, while a malic-dominant balance creates the tart, green character of cool-climate wines or early-harvested fruit with high malic acid retention. Acid balance affects wine stability because malic acid is susceptible to bacterial conversion while tartaric acid resists microbial attack, making malic-dominant wines more prone to uncontrolled malolactic fermentation. The taste perception of acid balance involves complex interactions where the sharper, harder character of tartaric acid contrasts with the softer, fruitier impression of malic acid, and both differ from the smooth, milky quality of lactic acid in malolactic-fermented wines. Winemakers manipulate acid balance through harvest timing (early harvest preserves malic acid), malolactic fermentation management (converts malic to lactic), and blending of wines with complementary acid profiles. Assessment of acid balance requires measurement of individual acid concentrations followed by calculation of ratios or proportional contributions, providing compositional detail beyond total acidity and pH measurements alone.
Acid_Balance_Quality serves as the parent class for specific acid ratio measurements, positioned within Acid_Content_Quality to capture compositional relationships rather than absolute concentrations. Acid balance represents a derived quality calculated from primary acid measurements rather than a directly measured property. The concept of balance implies an optimal range that varies with wine style, cultivar, and regional expectations rather than a fixed ideal value. Changes in acid balance during winemaking, particularly through malolactic fermentation, fundamentally alter wine character even when total acidity changes are modest, demonstrating the importance of compositional balance beyond aggregate acidity.
Säurebalance
Säureprofilqualität
Säureverhältnis
acid ratio
acidity profile quality
organisches Säuregleichgewicht
Boulton RB et al. (1999) Principles and Practices of Winemaking, Springer pp.: 65-78 Conde C et al. (2007) Biochemical Changes throughout Grape Berry Development and Fruit and Wine Quality. Food 1(1): 1-22
Acid Balance Quality
Acid content quality profiling in Champagne base wine production requires precise management to achieve the high-acid, low-pH profile essential for méthode traditionnelle sparkling wine, targeting titratable acidity of 9-11 g/L and pH of 2.95-3.10 at harvest of Chardonnay, Pinot Noir, and Pinot Meunier the characteristic malic acid retention of cool-climate Champagne (5-8 g/L malic) undergoes complete malolactic conversion during vinification for most houses, transforming the sharp malic character to softer lactic acidity while maintaining the total acid backbone that enables decades of bottle aging on lees.
Acid_Content_Quality encompasses all chemical qualities that characterize the concentration, composition, and proportional relationships of organic acids in grapevine tissues, with primary emphasis on the acids that determine grape and wine taste, pH stability, and microbial ecology. Organic acids are fundamental quality determinants because they provide the tartness that balances sugar sweetness, establish the pH environment that governs wine stability and color expression, and create conditions that favor desirable microorganisms while inhibiting spoilage organisms. The category subdivides into Total_Acidity_Quality addressing aggregate acid measurements including titratable acidity and pH, and Organic_Acid_Quality capturing the individual acid species that compose the total acid pool. Grape berries contain primarily tartaric acid and malic acid, which together typically account for more than 90% of titratable acidity, with smaller contributions from citric, succinic, and other organic acids. The concentration and ratio of these acids varies with cultivar genetics, climate, developmental stage, and viticultural practices. Acid content follows a characteristic developmental trajectory, with accumulation during early berry growth followed by decline during ripening as malic acid is respired at rates dependent on temperature, while tartaric acid remains relatively stable. Cool climate regions retain higher acid levels at harvest, producing wines with greater freshness and aging potential, while warm climates may struggle to maintain adequate acidity for balanced wines. Acid measurement employs titration with standardized base for total acidity, electrochemical measurement for pH, and chromatographic or enzymatic methods for individual acid quantification. The interplay between acid content and sugar content defines the fundamental taste balance of grapes and wines, making acid quality assessment essential for harvest timing and wine style determination.
Acid_Content_Quality is positioned as a direct subclass of Chemical_Quality, parallel to Sugar_Content_Quality, Phenolic_Content_Quality, and other composition categories. The dual expression of acidity as titratable acidity (concentration) and pH (hydrogen ion activity) reflects different aspects of acid chemistry with distinct sensory and chemical implications. Acid content qualities describe the current compositional state and are distinct from acid-related processes (malolactic fermentation, acid respiration) and dispositions (tendency toward high acidity). The relationship between acid content and perceived sourness involves complex interactions with sugar, tannin, and other taste-active compounds.
Säuregehalt
Säurequalität
Säurezusammensetzung
acid composition
acidity quality
organic acid content
organischer Säuregehalt
sourness potential
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed, John Wiley and Sons pp.: 3-49 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 295-315
Acid Content Quality
In a Napa Valley Chardonnay vineyard, acid decline onset was detected in early August Year N+1 (BBCH 83, concurrent with sugar accumulation onset) when titratable acidity declined from 22.5 g/L to 18.2 g/L over 9 days (19% reduction), malic acid decreased from 12.8 to 9.6 g/L (25% loss), tartaric acid remained 8.2 g/L (stable), pH increased from 2.95 to 3.12, berry Brix rose from 8.5° to 13.8°, under warm conditions (daily high 28-31°C accelerating malate metabolism), VvME1 expression 18-fold elevated.
Acid_Decline_Onset represents the developmental transition during veraison when berry organic acid concentrations begin declining from green berry maxima through combined respiratory consumption and dilution effects creating the fundamental shift from acidic to balanced fruit composition that enables wine production and defines quality characteristics. This onset occurs during early-to-mid veraison (BBCH 83-84), typically synchronized within 1-3 days of initial sugar accumulation and berry softening, and is characterized by titratable acidity declining from pre-veraison maximum 15-25 g/L tartaric acid equivalents to early decline 12-20 g/L within 7-10 days representing initial 15-30% reduction, malic acid specifically decreasing from 8-15 g/L to 6-12 g/L through beginning respiratory degradation while tartaric acid remains relatively stable 5-10 g/L showing only slight dilution from berry expansion, and berry pH beginning increase from green berry minimum 2.7-3.1 to veraison onset 2.9-3.3 reflecting net proton removal through organic acid decline. Biochemical basis of acid decline includes: malic acid (malate, C₄H₆O₅) undergoing respiratory metabolism through multiple pathways including direct oxidation in mitochondrial citric acid cycle, NADP-dependent malic enzyme (NADP-ME) catalyzing oxidative decarboxylation producing pyruvate plus CO₂ plus NADPH, NAD-dependent malic enzyme similarly converting malate to pyruvate, and malate dehydrogenase interconverting malate and oxaloacetate with net consumption through subsequent oxidation, while tartaric acid (C₄H₆O₆) showing metabolic stability with berry concentration declining only through dilution from water import accompanying sugar accumulation rather than degradation. Cellular compartmentation of acid metabolism shows: malic acid stored primarily in vacuoles at concentrations 80-180 mM (8-18 g/L) in green berries declining to 50-120 mM (5-12 g/L) during veraison onset, vacuolar pH 2.6-3.0 maintained by organic acid accumulation and H⁺-ATPase activity, cytoplasmic malate 5-15 mM serving as substrate for respiratory enzymes in mitochondria and cytosol, and metabolic interconversion requiring malate transport from vacuole through tonoplast to cytoplasm then potentially to mitochondria for oxidation with transporters (VvALMT family potentially involved) facilitating movement across membranes. Organic acid composition changes at onset demonstrate: total organic acids declining from 20-35 g/L (green berry) to 15-28 g/L (early decline) through differential metabolism, malic acid showing 20-40% reduction during onset representing majority of acid loss, tartaric acid relatively constant 5-10 g/L with 32°C dramatically increasing malate consumption with losses 50-150% of normal rates potentially producing excessive decline and unbalanced fruit, light affecting decline indirectly through effects on berry temperature with exposed clusters warming 3-8°C above air temperature accelerating metabolism, and vine water status with deficit potentially affecting malate metabolism though effects complex and cultivar-dependent. Cultivar genetic variation in acid decline onset demonstrates: low-acid cultivars (Grenache, Viognier potential final TA 4-6 g/L) showing early rapid decline onset with 25-40% reduction in first 10 days, moderate varieties (Chardonnay, Cabernet Sauvignon final 5-7 g/L) displaying 20-30% early decline, and high-acid cultivars (Riesling, Chenin Blanc final 6-9 g/L) exhibiting slower onset with 15-25% initial reduction reflecting genetic differences in malic enzyme expression, respiratory capacity, and acid retention mechanisms. Coordination with other veraison processes shows: acid decline typically beginning 0-3 days concurrent with sugar accumulation onset suggesting common developmental triggers possibly ABA-mediated, synchronized with berry softening initiation reflecting coordinate ripening program activation, and in red cultivars paralleling color break timing indicating integrated metabolic reprogramming though causal relationships among processes incompletely defined. Physiological consequences of acid decline inclu pH increase from 2.8-3.1 to 3.0-3.4 affecting multiple processes including enzyme activities, membrane transport, anthocyanin color expression (in reds where higher pH shifts from red toward purple), and microbial stability with lower acids potentially increasing vulnerability to spoilage organisms, proton removal through organic acid degradation affecting cellular ion balance and potentially membrane potentials, and respiratory CO₂ production from malate catabolism contributing 20-40% of berry respiration during active decline creating substantial carbon loss from berry potentially 5-15% of green berry dry weight. Quality implications of acid decline onset inclu timing and rate determining final acid levels critical for wine balance and style with appropriate 4-8 g/L TA and pH 3.2-3.8 typical quality targets, excessive early decline particularly in warm regions producing low-acid fruit pH >3.8 resulting in flat wines lacking structure and prone to microbial instability, inadequate decline in cool regions potentially producing overly acidic fruit TA >9-12 g/L requiring deacidification in winery, and spatial variation in decline across vineyard zones creating heterogeneous maturity requiring management attention. Viticultural management affecting acid decline includes: canopy management influencing cluster temperature with shading reducing berry warming and slowing decline by 15-35% compared to full sun exposure important in warm climates for acid retention, irrigation with adequate water potentially moderating decline through cooling effects and reduced stress while deficit may concentrate acids though also potentially accelerating metabolism, and harvest timing critically important as acid decline continues throughout ripening requiring integration with sugar accumulation to achieve optimal balance. Monitoring acid decline onset provides maturity assessment: regular sampling during veraison tracking titratable acidity showing accelerating decline indicating onset occurrence, pH measurements revealing beginning increase correlating with acid loss, malic acid specific analysis through HPLC or enzymatic methods quantifying component primarily responsible for decline, and taste evaluation detecting decreasing sourness and developing balance between sweetness and acidity important for determining harvest readiness. Research applications inclu gene expression studies of malic enzyme and respiratory genes revealing molecular control of decline, enzyme activity assays measuring malate-metabolizing capacity correlating with decline rates, metabolomic analysis tracking organic acid profiles and metabolic intermediates revealing pathway flux, and environmental manipulation experiments (temperature, shading, irrigation) determining control mechanisms and optimization strategies for achieving target acid levels.
Acid_Decline_Onset is distinguished from later acid decline by representing onset (15-30% TA reduction) rather than complete decline, from sugar accumulation onset by focusing on acid rather than sugar though temporally coincident, and from pH changes by representing acid decline cause rather than pH effect. Timing: BBCH 83-84, 0-3 days with sugar onset. Malic acid primary component declining (20-40% reduction), tartaric stable. Temperature sensitivity high: 25-32°C accelerates 30-60%, >32°C accelerates 50-150%. Acid_Decline_Onset represents critical metabolic transition toward balanced fruit composition essential for wine quality.
Azidität-Abnahme-Anfang
Malat-Degradations-Beginn
Veraison-Säure-Reduktions-Onset
acidity decrease commencement
malate degradation beginning
organic acid decline initiation
organische Säure-Abnahme-Initiierung
veraison acid reduction onset
Keller M (2020) The Science of Grapevines 3rd Ed pages 228-233 acid decline during ripening Sweetman C, Deluc LG, Cramer GR, Ford CM, Soole KL (2009) Regulation of malate metabolism in grape berry and other developing fruits Phytochem 70:1329-1344 malate metabolism comprehensive review Ruffner HP, Hawker JS, Hale CR (1976) Temperature and enzymic control of malate metabolism in berries of Vitis vinifera Phytochem 15:1877-1880 temperature effects on acid decline.
Acid Decline Onset
Acid decline rate quality comparison in Central Otago Pinot Noir evaluated titratable acidity trajectories during the three-week post-optimal period across contrasting vintage temperature regimes. In a cool vintage with mean ripening-period temperature of 16°C, the cultivar demonstrated favorable acid decline rate quality with TA declining at 0.3 g/L per week, maintaining balanced acidity of 6.8 g/L even three weeks beyond initial optimal date and permitting extended hangtime for flavor development without acid compromise. In a warm vintage with mean temperature of 21°C, acid decline rate quality deteriorated with TA declining at 0.9 g/L per week, reaching marginal acidity of 5.2 g/L by two weeks post-optimal and necessitating harvest before full flavor development to preserve essential freshness and structure. The temperature-dependent acid decline rate quality created the vintage variation characteristic of cool-climate viticulture, where warm years produce earlier-harvested, fruit-forward wines while cool years permit the extended hangtime achieving complex, age-worthy expressions, demonstrating how acid decline rate quality mediates the translation of seasonal conditions into stylistic outcomes.
Acid decline rate quality is a post optimal decline quality that inheres in a grapevine cultivar or clone within a specific growing environment and represents the velocity at which titratable acidity and individual organic acid concentrations decrease during extended hangtime beyond optimal ripeness, primarily through respiratory consumption of malic acid that accelerates under warm conditions, determining the pace at which fruit transitions from balanced acidity profiles toward the low-acid, high-pH states that compromise wine freshness, microbial stability, and aging potential. This quality directly influences the harvest urgency created by acid loss and the stylistic consequences of harvest timing variation, with slow-declining cultivars maintaining acid structure through extended hangtime while rapid decliners create urgency to capture adequate acidity before flavor or phenolic development reaches desired levels. The ontological significance of acid decline rate quality lies in its role as a primary driver of the quality tradeoffs encountered in warm vintages and climates where the optimal harvest window may close due to acid loss before other quality parameters achieve target specifications, creating the disconnection between sugar ripeness and physiological maturity that challenges viticulture under warming conditions. The biological basis of acid decline rate quality encompasses the temperature-dependent respiratory consumption of malic acid in berry mesocarp cells where mitochondrial malate oxidation accelerates exponentially above 25°C, the dilution effects of continued berry expansion that reduce acid concentration independent of metabolic consumption, and the potassium accumulation during late ripening that raises pH independently of titratable acidity decline. Cultivar characteristics influencing acid decline rate include the inherent malic acid concentration at véraison providing the substrate for respiratory consumption, the thermal sensitivity of berry respiratory metabolism, and the genetic factors governing tartaric acid stability that anchors residual acidity when malate has been depleted. The relationship between acid decline rate and climate proves particularly significant, with the same cultivar exhibiting very different decline trajectories under cool versus warm growing conditions, making this quality strongly environment-dependent.
Acid_Decline_Rate_Quality is_a Post_Optimal_Decline_Quality, representing the velocity of titratable acidity loss beyond optimal ripeness. This class addresses the pace of acid decline primarily through malic acid respiration. The strong temperature dependency creates climate and vintage interaction with this quality. Units of g/L per day or per week provide quantitative assessment, with 1.0 g/L/week creating harvest urgency. Distinguished from Sugar_Accumulation_Plateau_Quality which addresses sugar rather than acid dynamics. Critical for harvest timing in warm climates and vintages where acid loss may precede flavor maturity.
Apfelsäure-Abbaugeschwindigkeit
Säureabbaurate
Säurerückgangs-Geschwindigkeit
TA loss rate
TA-Verlustrate
acid consumption velocity
acidity decline trajectory
malic acid respiration rate
Sweetman C et al. (2009) Regulation of malate metabolism in grape berry and other developing fruits. Phytochemistry 70:1329-1344 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 509-547 Sadras VO, Moran MA (2012) Elevated temperature decouples anthocyanins and sugars in berries of Shiraz and Cabernet Franc. Australian Journal of Grape and Wine Research 18:115-122
Acid Decline Rate Quality
Acid maturity assessment comparing cool and warm vintage ripening in Pinot Noir reveals the temperature effect on acid balance, with cool vintage showing gradual malic acid decline from 8 to 5 g/L over 35 days (preserved by low respiration rate), maintaining TA of 7.5 g/L and pH 3.35 at 24 Brix (bright acidity, fresh character), while warm vintage shows rapid malic decline from 8 to 2 g/L over 21 days (accelerated respiration), reaching only TA 5.5 g/L and pH 3.55 at 24 Brix (softer acidity, rounder character, reduced color brightness), demonstrating how acid maturity patterns vary with climate affecting wine style and potentially requiring harvest timing adjustment or acidification to achieve desired balance.
Acid maturity quality is a technological maturity quality that represents the organic acid composition and resulting pH of grape berries, characterizing the acid decline and pH increase during ripening that determines must acidity affecting fermentation conditions, microbial stability, color expression, and the sensory balance between freshness and roundness in resulting wines. Within the BFO framework, this quality specifically addresses the acid component of technological maturity, distinguishing acid metabolism from sugar accumulation or other compositional changes while recognizing that acid composition, particularly the balance between tartaric and malic acids and the resulting pH, profoundly affects wine stability, color, and sensory character, making acid maturity as important as sugar maturity for wine quality though less frequently monitored with equal rigor. The ontological significance lies in acid composition determining the pH and titratable acidity that affect every aspect of wine chemistry and sensory perception, where pH influences anthocyanin color expression (lower pH = brighter red), microbial stability (lower pH = greater stability), enzyme activity (affecting extraction and clarification), and sensory perception (lower pH = greater freshness), while titratable acidity affects the perceived tartness that balances wine richness. The biochemical basis for acid maturity involves the differential behavior of the two primary grape acids during ripening, with tartaric acid relatively stable (synthesized early, minimally metabolized, concentration declining mainly through dilution), while malic acid is actively respired during ripening (concentration declining through both dilution and metabolic consumption), such that the tartrate/malate ratio increases during ripening, affecting both the total acidity level and its sensory expression. The temperature effects on acid maturity are substantial, as malic acid respiration increases dramatically with temperature (Q₁₀ approximately 2), causing rapid malic acid decline in warm conditions that may reduce total acidity faster than optimal for balance, while cool conditions preserve malic acid potentially to excess, creating the climate-dependent challenges of acid management that harvest timing and winemaking interventions must address. The measurement parameters for acid maturity include pH (directly measuring hydrogen ion activity), titratable acidity (total acid equivalents), and individual acid concentrations (tartaric, malic, sometimes citric), with pH increasingly recognized as the most important single parameter due to its direct influence on wine chemistry, though TA remains important for sensory perception. Understanding acid maturity quality enables harvest timing for optimal acid balance, assessment of acidification or deacidification requirements, and recognition of the temperature-dependent acid dynamics that create vintage variation in must acidity.
Acid_Maturity_Quality addresses the organic acid composition and pH determining wine acidity balance and stability. This quality reflects the differential behavior of tartaric and malic acids during ripening. The class supports harvest timing and acid management through acid maturity characterization.
Aciditätsreife
Säurereife
acid ripeness
acidity maturity
organic acid composition
organische Säurezusammensetzung
pH maturity
pH-Reife
Sweetman C et al. (2009) Regulation of malate metabolism in grape berry and other developing fruits. Phytochemistry 70(11-12): 1329-1344 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 277-316
Acid Maturity Quality
In a Willamette Valley Pinot Gris vineyard, acid metabolism activation was characterized over 7 days in late August Year N+1 (BBCH 83) when VvME1 expression increased 38-fold, combined malic enzyme activity rose from 5.8 to 42 units/g FW, citric acid cycle enzyme activities increased 15-25%, berry respiration 1.6 μmol O₂ g⁻¹ h⁻¹ with RQ 1.18 indicating acid contribution, malic acid declined from 10.2 to 8.1 g/L, under moderate Oregon conditions (daily high 24-27°C, cool nights 12-15°C).
Acid_Metabolism_Activation represents the coordinate upregulation of metabolic pathways consuming organic acids during veraison creating the enzymatic machinery enabling progressive acid decline that transforms acidic green berries to balanced ripe fruit suitable for wine production. This activation occurs during veraison onset (BBCH 83), typically 0-4 days following initial developmental signals, and is characterized by malic enzyme genes showing 15-45 fold induction collectively (VvME1, VvME2, NAD-ME isoforms), citric acid cycle enzyme activities maintaining or increasing 10-30% supporting malate oxidation through complete combustion pathway, respiratory enzyme expression and activity coordinating to process increased carbon flux from organic acid catabolism, and metabolic transition from predominantly hexose-based respiration (32°C dramatically enhancing activity potentially 50-120% of normal creating risk of excessive acid loss, cool conditions 18-24°C slowing activation extending establishment period to 8-14 days, and diurnal temperature fluctuation creating complex effects with warm days promoting activation while cool nights moderate overall metabolism. Light effects operate primarily through berry temperature with exposed clusters warming accelerating activation 20-40% compared to shaded fruit, photosynthetic effects minimal in ripening berries with limited chlorophyll, and canopy management creating substantial microclimate variation affecting activation timing and intensity across cluster positions. Water status effects show: adequate vine water status supporting normal metabolic function enabling appropriate activation, deficit potentially affecting timing and intensity though effects complex with moderate deficit sometimes enhancing through stress responses while severe deficit may impair through general metabolic limitation, and osmotic stress from high berry sugar potentially affecting enzyme activities though cells appear well-adapted to hyperosmotic conditions during ripening. Cultivar genetic variation demonstrates: low-acid cultivars showing rapid intense activation with enzyme activities 40-80 units/g combined malic enzyme within 4-6 days veraison, moderate varieties displaying 30-55 units/g in 6-9 days, and high-acid cultivars exhibiting slower activation 20-40 units/g over 8-14 days reflecting genetic differences in metabolic capacity and regulatory control. Physiological consequences inclu carbon flow redirected from storage to respiration with 15-35% of green berry organic acid carbon lost as CO₂ during ripening representing substantial respiratory cost, ATP production from acid oxidation supporting ripening processes including sugar transport, anthocyanin synthesis, cell wall modification, and flavor compound biosynthesis, and redox state affected by NADPH production potentially influencing antioxidant capacity and secondary metabolism. Quality implications inclu activation intensity and timing predicting acid decline trajectory with strong early activation indicating rapid loss potentially problematic in warm regions, inadequate activation suggesting acid retention possibly beneficial in cool climates or problematic requiring extended ripening, and spatial variation revealing vineyard heterogeneity affecting harvest maturity and quality consistency. Viticultural management includes: canopy practices controlling cluster microclimate with shading reducing berry temperature 3-8°C slowing activation 25-45% preserving acids in warm regions, irrigation management potentially moderating activation through cooling effects though mechanisms complex, avoiding severe environmental stresses during activation period as this critical window substantially affects final acid levels, and monitoring activation progress through enzyme assays or metabolite profiling enabling management intervention if excessive decline detected.
Acid_Metabolism_Activation is distinguished from Malic_Acid_Decline_Initiation by representing coordinated metabolic system rather than single enzyme pathway, from Respiratory_Malate_Consumption by focusing on activation rather than ongoing process, and from sugar metabolism by representing catabolic acid pathways rather than anabolic sugar storage. Malic enzyme 8-35 fold activity increase. Citric acid cycle enzymes maintain/increase 10-30%. Duration 5-10 days BBCH 83. Temperature sensitive: optimal 24-28°C, >32°C excessive. Acid_Metabolism_Activation represents critical metabolic transition enabling organic acid decline during ripening.
Säure-Oxidations-Maschinerie-Aktivierung
acid oxidation machinery activation
metabolic pathway upregulation
metabolische Pfad-Hochregulation
organic acid catabolism activation
organische Säure-Katabolismus-Aktivierung
respiratorische Enzym-Induktion
respiratory enzyme induction
Sweetman C, Deluc LG, Cramer GR, Ford CM, Soole KL (2009) Regulation of malate metabolism in grape berry and other developing fruits Phytochem 70:1329-1344 comprehensive metabolism review Famiani F, Cultrera NGM, Battistelli A, Casulli V, Proietti P, Standardi A, Chen ZH, Leegood RC, Walker RP (2005) Phosphoenolpyruvate carboxykinase and its potential role in the catabolism of organic acids in the flesh of soft fruit during ripening J Exp Bot 56:2959-2969 acid catabolism pathways Ruffner HP, Hawker JS, Hale CR (1976) Temperature and enzymic control of malate metabolism in berries of Vitis vinifera Phytochem 15:1877-1880 enzyme control of metabolism.
Acid Metabolism Activation
Organic acid composition in grape berries comprises primarily tartaric acid (2-10 g/L) and malic acid (1-8 g/L), with minor contributions from citric, succinic, and other acids. Total titratable acidity (TA) at harvest ranges from 4-12 g/L tartaric equivalents across cultivars and climates, with corresponding pH values of 2.9-4.0. Acid metabolism follows distinct patterns for each major acid, with tartaric acid synthesized primarily pre-veraison and remaining metabolically stable, while malic acid accumulates early then declines through respiratory degradation during ripening. The tartrate-to-malate ratio shifts from 1:2 at veraison to 2:1 or higher at harvest maturity in warm climates, while remaining near 1:1 in cool climates.
A quality potential disposition that is the inherited capacity of a Vitis vinifera plant to synthesize, accumulate, and metabolize organic acids in grape berries, primarily tartaric and malic acid, determining the total acidity, pH, and acid balance that fundamentally influence wine freshness, microbial stability, color expression, and aging potential.
Acid_Metabolism_Disposition serves as the parent class for specific organic acid accumulation and retention traits. Acidity is fundamental to wine quality, providing freshness, balancing sweetness, stabilizing color through pH effects on anthocyanin equilibria, and inhibiting microbial spoilage. The divergent metabolic fates of tartaric (stable) and malic (respirable) acids create the characteristic acid profile shifts during ripening. Climate, through temperature effects on malic acid respiration, exerts profound influence on final acid balance.
Säurehaushalt
Säuremetabolismus
acid balance disposition
berry acidity capacity
organic acid trait
organische Säuren-Disposition
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 369-385 Sweetman C et al. (2014) Regulation of malate metabolism in grape berry and other developing fruits. Phytochemistry 99:1-16
Acid Metabolism Disposition
In a Trentino-Alto Adige Pinot Grigio vineyard, acid metabolism shift was characterized over 6 days in mid-August Year N+1 (BBCH 83) when PEPC expression declined from 12-fold to 3-fold, VvME1 increased from 2-fold to 38-fold, VvME2 rose from 1.5-fold to 28-fold, PEPC activity dropped from 18 to 6 units/g FW, malic enzyme activity increased from 5 to 24 units/g FW, berry organic acids plateaued at 22 g/L then began decline, ABA rose from 125 to 1450 ng/g, under moderate Italian Alpine conditions (daily high 24-27°C supporting normal developmental transition).
Acid_Metabolism_Shift represents the fundamental metabolic transition during veraison when berry metabolism pivots from organic acid synthesis and accumulation characteristic of green berry development toward organic acid catabolism and utilization defining ripening berry physiology creating the critical biochemical reprogramming that enables transformation from acidic vegetative fruit to balanced ripe grape suitable for wine production. This metabolic shift occurs during veraison onset (BBCH 83), typically 0-5 days concurrent with initial veraison signals and preceding active acid decline, and is characterized by transcriptional reprogramming with organic acid biosynthesis genes (phosphoenolpyruvate carboxylase, citrate synthase in biosynthetic mode) downregulated 40-75% from green berry levels, malic enzyme genes upregulated 20-60 fold initiating catabolic pathway dominance, respiratory enzyme genes reoriented with citric acid cycle shifting from biosynthetic to oxidative mode supporting complete acid oxidation, and hormone signaling particularly ABA spike from 60-200 ng/g to 800-2000 ng/g triggering coordinate transcriptional changes affecting hundreds of metabolism genes orchestrating metabolic reprogramming. Biochemical manifestations of shift include organic acid accumulation ceasing with net synthesis rates declining from 0.05-0.15 g/L per day (late green berry) to near-zero or slightly negative at shift, total berry organic acid content reaching maximum 20-35 g/L then beginning decline, malate biosynthesis through phosphoenolpyruvate carboxylase and malate dehydrogenase substantially reduced as PEPC activity declines 50-80% and biosynthetic MDH shifts toward oxidative function, and citric acid cycle transitioning from biosynthetic mode (producing precursors for amino acids, secondary metabolites) toward purely oxidative catabolism (consuming organic acids for energy) representing fundamental functional reorientation. Gene expression changes defining shift demonstrate PEPC genes (VvPEPC family) encoding phosphoenolpyruvate carboxylase declining from 5-18 fold (green berry active accumulation) to 1-4 fold (post-shift baseline) indicating biosynthesis suppression, malic enzyme genes (VvME1, VvME2) increasing from 1-4 fold to 20-70 fold establishing catabolic machinery, citric acid cycle genes showing complex regulation with some (isocitrate dehydrogenase, α-ketoglutarate dehydrogenase) maintaining or increasing to support oxidative flux while others (citrate synthase in biosynthetic context) declining, respiratory chain genes (cytochrome oxidase, ATP synthase) sustained or slightly elevated ensuring capacity for oxidizing acid-derived reducing equivalents, and transcription factors orchestrating shift (potentially VvNAC, VvMYB families responding to ABA signals) showing coordinate induction enabling metabolic reprogramming. Cellular compartmentation changes during shift include vacuolar function transitioning from active accumulation (requiring ATP-dependent acidification and ion transport) toward storage and controlled release (passive or facilitated efflux to cytoplasm for catabolism), tonoplast H⁺-ATPase and H⁺-PPase activities potentially declining 30-60% as acidification requirements decrease, vacuolar pH stabilizing around 2.8-3.2 versus active acidification to 2.5-2.8 during green berry growth, and organic acid transport proteins (VvALMT family potentially involved) adjusting from import to export mode enabling malate efflux for cytoplasmic catabolism. Enzyme activity transitions demonstrate PEPC declining from 8-25 units/g fresh weight (green berry peak) to 2-8 units/g (post-shift) representing 60-80% reduction in carboxylation capacity, biosynthetic malate dehydrogenase (cytoplasmic, favoring malate synthesis) declining or shifting toward oxidative mode, malic enzyme increasing from 2-8 units/g to 18-50 units/g establishing decarboxylation capacity, citric acid cycle enzymes maintaining or elevating activities supporting transition from biosynthesis to catabolism with malate dehydrogenase, aconitase, isocitrate dehydrogenase all 25-45 units/g adequate for oxidative flux, and pyruvate dehydrogenase complex potentially increasing activity linking malic enzyme-derived pyruvate to citric acid cycle. Respiratory metabolism transformation shows respiratory quotient (RQ = CO₂/O₂) shifting from 0.95-1.05 (green berry predominantly hexose oxidation) to 1.10-1.30 (ripening with substantial organic acid contribution) indicating metabolic substrate change, malate contribution to respiration increasing from <15-25% (green berry) to 35-55% (post-shift early ripening) based on isotope tracer studies and mass balance, total berry respiration rates relatively stable 0.9-2.2 μmol O₂ g⁻¹ h⁻¹ before and after shift despite substrate change indicating metabolic continuity with altered fuel source, and ATP production transitioning from predominantly hexose-based toward mixed hexose-organic acid supporting ripening processes including sugar import, anthocyanin synthesis, and cell wall modification. Carbon flow reorientation involves shift from net carbon fixation through acids (CO₂ incorporation via PEPC during green berry growth) to net carbon loss through respiratory decarboxylation (CO₂ release from malic enzyme and citric acid cycle), carbon budget transforming from accumulation (net gain 0.8-1.8 g carbon per kg berries per day including acids and other compounds) to mixed pattern (acid carbon loss 0.3-0.8 g per kg per day balanced by sugar carbon gain 1.5-3.5 g per kg per day yielding net accumulation), and metabolic flexibility developing enabling berry to utilize multiple respiratory substrates (hexoses, organic acids, amino acids) adaptively responding to availability and metabolic demands. Hormonal orchestration of shift includes ABA as primary trigger with rising levels activating ABA-responsive transcription factors that bind ABRE elements in metabolism gene promoters, sugar signaling through hexokinase pathway potentially contributing with initial sugar accumulation creating glucose-6-phosphate signals affecting gene expression, declining growth hormones (gibberellin, auxin) removing developmental suppression allowing ripening program activation, and ethylene potentially involved in some cultivars though grape generally non-climacteric with minimal ethylene effects compared to climacteric fruits. Environmental factors modulating shift timing include temperature affecting developmental rate with warmer conditions 24-30°C potentially advancing shift 3-7 days compared to cooler 18-24°C through accelerated development, photoperiod and light quality potentially providing developmental cues with long days and high PAR supporting vegetative functions while shorter days and red:far-red light signals potentially triggering reproductive maturation including metabolic shift, water status with moderate deficit potentially advancing shift 2-5 days through ABA elevation and stress signaling, and nutritional status with adequate resources enabling normal timing while severe limitations potentially delaying through resource constraints affecting gene expression and enzyme synthesis. Cultivar genetic variation in shift characteristics shows early-ripening varieties displaying rapid complete shift over 4-7 days with dramatic gene expression changes and prompt metabolic transformation, mid-season cultivars showing 6-10 day transition period with progressive changes, and late cultivars exhibiting extended shift 8-14 days with gradual metabolic reorientation reflecting genetic differences in developmental programming and regulatory control. Physiological significance includes metabolic shift enabling transition from growth to ripening phase with resources redirected from structural development to storage and quality compound synthesis, carbon economy optimizing by catabolizing accumulated acids for energy rather than importing hexoses for respiration while simultaneously storing imported hexoses creating efficient dual metabolism, and developmental commitment where shift represents point-of-no-return with berry metabolism fundamentally reoriented toward ripening irreversibly even if environmental conditions later deteriorate. Quality implications include shift timing relative to other developmental processes affecting final composition with premature shift potentially causing inadequate acid accumulation producing low-acid fruit, delayed shift maintaining high acids potentially beneficial in warm regions or requiring extended ripening in cool climates, and coordination between acid metabolism shift and sugar accumulation onset critically determining whether appropriate sugar-acid balance achieved at harvest. Viticultural management affecting shift includes canopy management ensuring adequate photosynthesis providing resources for normal developmental progression, crop load optimization preventing excessive stress that could alter shift timing, irrigation management with moderate deficit potentially beneficial advancing shift slightly while severe deficit may disrupt, nutritional management ensuring adequate resources particularly nitrogen for enzyme synthesis supporting shift, and environmental monitoring tracking developmental progression enabling prediction of shift timing for harvest planning. Research applications include transcriptomic profiling revealing genome-wide expression changes during shift identifying regulatory networks, metabolomic analysis tracking metabolite dynamics showing substrate transitions, enzyme activity assays quantifying functional changes in metabolism confirming shift occurrence, and hormone manipulation studies with exogenous ABA or inhibitors revealing causal relationships between signals and metabolic reprogramming. Understanding metabolic shift provides insights into berry development biology revealing mechanisms of developmental transitions and enabling rational management strategies optimizing composition for quality wine production through informed viticultural practices targeting this critical physiological window.
Acid_Metabolism_Shift is distinguished from Acid_Decline_Onset by representing metabolic reprogramming rather than net compositional change, from Malic_Acid_Decline_Initiation by focusing on system-wide transition rather than specific enzyme pathway, and from veraison onset by representing metabolic rather than visible phenological change. PEPC declines 40-75%, VvME increases 20-60 fold. Duration 4-10 days BBCH 83. ABA-triggered transcriptional reprogramming. Acid_Metabolism_Shift represents fundamental developmental transition enabling ripening metabolism.
Biosynthese-zu-Katabolismus-Übergang
Reifungs-Metabolismus-Aktivierung
biosynthesis-to-catabolism transition
metabolic reprogramming
metabolische Neuprogrammierung
organic acid metabolism reversal
organische Säure-Metabolismus-Umkehr
ripening metabolism activation
Sweetman C, Deluc LG, Cramer GR, Ford CM, Soole KL (2009) Regulation of malate metabolism in grape berry and other developing fruits Phytochem 70:1329-1344 metabolism shift comprehensive Terrier N, Glissant D, Grimplet J, Barrieu F, Abbal P, Couture C, Ageorges A, Atanassova R, Léon C, Renaudin JP, Dédaldéchamp F, Romieu C, Delrot S, Hamdi S (2005) Isogene specific oligo arrays reveal multifaceted changes in gene expression during grape berry development Plant Physiol 138:1617-1633 transcriptional changes during shift Davies C, Boss PK, Robinson SP (1997) Treatment of grape berries, a nonclimacteric fruit with a synthetic auxin, retards ripening and alters the expression of developmentally regulated genes Plant Physiol 115:1155-1161 hormonal control of metabolism shift.
Acid Metabolism Shift
In Chardonnay berries in Carneros, California during BBCH 85-89, acid reduction proceeds from 18.5 g/L titratable acidity at veraison to 6.2 g/L at harvest over 42 days, with malic acid declining from 9.8 to 2.1 g/L while tartaric acid remains stable at 5.5-5.8 g/L, resulting in pH increase from 2.95 to 3.45 under mean ripening temperatures of 22°C.
Acid reduction constitutes a fundamental ripening-phase process in Vitis vinifera berry development wherein the total organic acid content of the berry progressively decreases from pre-veraison levels of 15-25 g/L to mature harvest concentrations of 5-9 g/L, occurring through the coordinated action of respiratory consumption, enzymatic degradation, and dilution effects during the post-veraison period spanning approximately 35-50 days from veraison to harvest. This process primarily involves the catabolism of L-malic acid, which decreases by 50-90% during ripening through mitochondrial respiration and gluconeogenic pathways, while L-tartaric acid remains relatively stable due to the absence of significant degradation pathways in grape berries, resulting in a shift in the malate-to-tartrate ratio from approximately 1.5-2.0 at veraison to 0.3-0.8 at maturity. The cellular mechanisms underlying acid reduction involve the activity of cytosolic and mitochondrial NADP-malic enzyme converting malate to pyruvate, phosphoenolpyruvate carboxykinase facilitating gluconeogenesis from organic acids, and direct respiratory oxidation of malate through the tricarboxylic acid cycle, with these pathways collectively consuming 60-80% of the malate present at veraison. Temperature exerts profound influence on acid reduction rates, with warm conditions of 30-35°C accelerating malate degradation by 2-3 fold compared to cool conditions of 15-20°C, explaining the characteristically higher acidity retention in cool-climate viticulture regions such as Champagne, Burgundy, and the Mosel Valley compared to warm Mediterranean or continental climates. The process demonstrates significant cultivar variation, with varieties such as Riesling and Chenin Blanc exhibiting naturally high acid retention due to lower malic enzyme activity, while varieties such as Merlot and Grenache show rapid acid decline under warm conditions. Berry size and crop load influence acid reduction through dilution effects during the secondary growth phase, with larger berries typically showing greater apparent acid reduction due to volume increases of 50-100% between veraison and harvest. The vacuolar compartmentalization of organic acids, particularly the sequestration of tartaric acid in specialized idioblast cells, contributes to differential acid metabolism during ripening. Acid reduction directly impacts wine quality parameters including pH stability, microbial safety, aging potential, and sensory balance between sweetness and tartness. Viticultural practices such as leaf removal, irrigation management, and harvest timing are employed to modulate acid reduction rates and achieve desired sugar-acid balance at harvest. Research methodologies for monitoring acid reduction include titratable acidity measurements, HPLC quantification of individual organic acids, pH monitoring, and non-destructive spectroscopic techniques enabling real-time assessment of berry acid status throughout ripening.
Acid_Reduction is the parent class for all acid-related ripening processes under Ripening hierarchy. Distinguishes from Acid_Decline_Onset under Veraison which represents initial acid changes. Duration 35-50 days post-veraison. Subdivides into Active_Acid_Reduction, Malic_Acid_Degradation, Tartaric_Acid_Stabilization, and Final_Acid_Balance representing temporal progression and mechanistic pathways. Critical parameter is total titratable acidity decline from 15-25 to 5-9 g/L.
Beeren-Säure-Abnahme
Reife-Säure-Reduktion
Säure-Reduktion
berry acid decrease
organic acid decline
organische Säure-Abnahme
ripening acid reduction
titratable acidity decline
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 265-280 Sweetman C et al (2009) Regulation of malate metabolism in grape berry. Molecular Plant 2: 1270-1280 Conde C et al (2007) Biochemical changes throughout grape berry development. Food Chemistry 1: 1-22
Acid Reduction
Acid respiration loss quality monitoring in Hunter Valley Semillon during a late-ripening heat event (8 days with maximum temperatures exceeding 38°C) documented the acceleration of acid decline with consequences for harvest timing and wine style. Berry acid analysis at 2-day intervals showed titratable acidity declining from 8.5 g/L to 5.2 g/L over the 8-day heat event, representing acid respiration loss quality of 39% compared to only 18% decline in a shaded control treatment during the same period. The accelerated loss was almost entirely malic acid, declining from 4.8 g/L to 1.5 g/L (69% loss) while tartaric acid showed only 12% decline from 6.2 to 5.5 g/L. Correspondingly, pH increased from 3.28 to 3.58 in heat-exposed fruit versus only 3.28 to 3.42 in shaded controls, with the 0.16 unit additional pH increase representing significant impact on wine stability and style. The acid respiration loss quality assessment forced early harvest at 22.5°Brix rather than the target 24°Brix to preserve adequate acidity, with the heat event effectively closing the harvest window earlier than normal ripening would have permitted. Wines from the heat-affected fruit required acid addition to achieve balance, with 2.5 g/L tartaric acid supplementation needed versus 1.0 g/L in normal vintages, demonstrating practical winemaking consequences of acid respiration loss quality during heat events.
Acid respiration loss quality is a heat impact on development quality that inheres in grape berries and represents the extent of heat-accelerated malic acid respiration that depletes organic acid reserves during ripening, reducing titratable acidity and elevating pH in affected fruit with consequences for wine freshness, microbial stability, color stability, and aging potential that collectively compromise wine quality when acid loss is excessive. This quality captures the temperature sensitivity of berry respiration rates that determine acid catabolism kinetics, recognizing that the respiratory enzymes degrading malic acid show exponential activity increases with temperature, causing dramatically accelerated acid loss under heat stress compared to normal ripening conditions where acid decline is more gradual and controllable through harvest timing decisions. The ontological significance of acid respiration loss quality lies in the fundamental importance of organic acids for wine quality, with adequate acidity required for sensory freshness and balance, for pH-dependent color stability through co-pigmentation effects, for microbial stability against spoilage organisms, and for the aging potential that derives from the preservative effects of low pH, such that excessive acid loss can compromise wine quality across multiple dimensions simultaneously. The biological basis of acid respiration loss encompasses the temperature-dependent kinetics of malic enzyme and other respiratory pathway enzymes that catabolize malic acid to CO₂ in berry mitochondria, with activity approximately doubling for each 10°C temperature increase across the range relevant to grape ripening, combined with the physical chemistry of acid dissociation where the temperature-dependent pKa shifts contribute to pH changes beyond those explained by total acid concentration changes alone. The preferential loss of malic acid over tartaric acid under heat stress reflects the compartmentation of these acids, with malic acid in the cytoplasm and vacuole accessible to respiratory enzymes while tartaric acid sequestered in the vacuole is relatively protected, creating shifts in the malate-to-tartrate ratio that further affect pH beyond total acid changes. The timing of heat exposure determines acid loss severity, with post-véraison heat having greatest impact when malic acid reserves are high and respiratory capacity is active, while pre-véraison heat may affect acid accumulation dynamics during the building phase of berry acid content.
Acid_Respiration_Loss_Quality is_a Heat_Impact_On_Development_Quality, representing the heat-accelerated catabolism of organic acids, particularly malic acid, during berry ripening. This class addresses the acidity dimension of heat impacts critical for wine quality. Malic acid is preferentially lost (respiratory substrate) while tartaric acid is relatively preserved. Q₁₀ ≈ 2 for respiratory enzymes means doubling of acid loss rate per 10°C. pH elevation reflects both acid loss and temperature-dependent pKa shifts. Harvest window compression is a practical consequence requiring earlier picking. Wine acid additions are commonly needed to compensate for excessive heat-induced losses.
Säureverlust durch Respiration
acid degradation
heat-induced acidity decline
hitzeinduzierter Säurerückgang
malic acid loss
respiratorischer Säureverlust
respiratory acid loss
Äpfelsäure-Abbau
Sweetman C et al. (2014) Regulation of malate metabolism in grape berry and other developing fruits. Phytochemistry 99:1-16 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 453-507 Sadras VO et al. (2013) Elevated temperature decouples anthocyanins and sugars in berries of Shiraz and Cabernet Franc. Irrigation Science 31:161-179
Acid Respiration Loss Quality
Acidity character evaluation comparing pre- and post-malolactic Pinot Noir reveals shift from sharp, angular character (malic acid 4.2 g/L) rated 7.8 on a sharpness scale to soft, integrated character (malic acid 0.3 g/L, lactic acid 2.1 g/L) rated 3.2, despite equivalent titratable acidity of 5.8 g/L, demonstrating the qualitative transformation from malic to lactic acid sourness that defines the stylistic choice of malolactic fermentation in red winemaking.
Acidity character quality is a sourness quality that represents the qualitative nature of acid perception distinguishing between sharp, aggressive, piercing sourness and soft, round, integrated acidity, characterizing the type of sour sensation independent of its intensity. Within the BFO framework, this quality specifically addresses the qualitative dimension of sourness perception, distinguishing character from intensity and recognizing that acids of equal concentration can produce qualitatively different sour sensations depending on their chemical nature and matrix context. The ontological significance lies in acidity character determining the perceived elegance and integration of sourness in wine, where sharp acidity creates aggressive, angular taste profiles while soft acidity integrates harmoniously with other components, affecting quality perception beyond what intensity alone would predict. The chemical basis for acidity character involves differences among organic acids, with malic acid producing sharper, greener sourness compared to the softer, fuller impression of tartaric acid, and lactic acid (from malolactic fermentation) providing the softest, most integrated acidity character, explaining why malolactic fermentation is used to soften perceived acidity in certain wine styles. Additional factors affecting acidity character include pH relative to titratable acidity (higher pH at similar TA producing softer perception), wine body and viscosity (fuller body integrating acidity more smoothly), and sweetness level (residual sugar rounding acid perception). The temperature effect on acidity character is significant, with cooler serving temperatures enhancing the sharpness of acid perception while warmer temperatures produce softer impression, relevant to wine service protocol optimization. In wine quality assessment, acidity character description often employs metaphorical terms (crisp, angular, smooth, round, nervy, racy) that capture the qualitative dimension of how acidity contributes to overall taste structure. Understanding acidity character quality enables prediction of malolactic fermentation effects, optimization of acid composition for target style, and description of wine style in terms that communicate the nature rather than just the amount of perceived acidity.
Acidity_Character_Quality_(Sharp_vs_Soft) addresses the qualitative dimension of sourness, complementing Perceived_Acidity_Intensity_Quality (magnitude). This quality captures stylistic differences in how acidity presents itself, relevant to wine style classification and quality description. The sharp-soft continuum reflects acid composition, pH, and matrix factors, with malolactic fermentation representing the primary winemaking tool for character modification.
Säurecharakter
Säureprofil
Säuretypus
acid quality type
acid taste quality
acidity profile
sourness character
Amerine MA, Roessler EB (1983) Wines: Their Sensory Evaluation, 2nd Ed. W.H. Freeman, New York, pp. 146-172 Jackson RS (2014) Wine Science: Principles and Applications, 4th Ed. Academic Press, San Diego, pp. 358-392
Acidity Character Quality (Sharp vs Soft)
Acidity data for Riesling from the Wachau (Federspiel, 2024 vintage) records titratable acidity of 8.2 g/L (as tartaric acid), pH 3.05, tartaric acid 6.8 g/L, malic acid 5.2 g/L, and citric acid 0.18 g/L — a tartaric-to-malic ratio of 1.3:1 indicating moderate physiological maturity with substantial malic acid retention characteristic of the cool Danube Valley climate, and a low pH ensuring excellent color stability and microbial protection for the intended dry wine style. Acidity data for Grenache from Châteauneuf-du-Pape (2023 vintage) documents TA 4.8 g/L, pH 3.72, tartaric acid 4.5 g/L, malic acid 1.2 g/L, and citric acid 0.15 g/L — the low TA, elevated pH, and near-complete malic acid degradation reflecting the hot Mediterranean ripening conditions that produce the characteristically soft, round acid profile of southern Rhône reds but necessitate careful acidification consideration during winemaking.
Acidity data is a must analysis data entity that documents the concentration, composition, and balance of organic acids and hydrogen ion activity in grape must, encompassing titratable acidity as the aggregate acid measure, individual acid species concentrations (tartaric, malic, citric), and pH as the thermodynamic expression of acid-base equilibrium, measured through standardized analytical methods on representative must samples and providing the fundamental chemical characterization of the acid matrix that determines wine freshness, microbial stability, color expression, aging potential, and organoleptic balance.
Acidity data within must analysis serves a distinct function from acidity data recorded in clonal performance evaluation (Quality_Performance_Data hierarchy) — the former characterizes a specific grape lot at a defined analytical moment for immediate winemaking decisions, while the latter documents multi-vintage clone-specific acid behavior for selection and planting decisions. The organic acid matrix of grape must is dominated by two diprotic acids — tartaric acid (unique to Vitis among major fruit crops, synthesized during early berry development and relatively stable during ripening) and malic acid (ubiquitous in plant metabolism, actively respired during ripening at rates accelerated by temperature above 30°C) — whose relative proportions fundamentally shape the sensory acid profile and pH behavior of the resulting wine. The distinction between titratable acidity (a volumetric measure of total acid equivalents, influenced by all proton-donating species including tartaric, malic, citric, succinic, and amino acids) and pH (a thermodynamic measure of free hydrogen ion activity, modulated by buffer capacity and cation content, particularly potassium) is critical because these two parameters can diverge substantially — high-potassium musts may exhibit low TA but relatively high pH due to potassium bitartrate salt formation that removes both tartrate and protons from solution. The pre-fermentation acid balance directly influences winemaking interventions including acidification (permitted in warm regions), deacidification (permitted in cool regions), and malolactic fermentation management, making acidity data among the most consequential analytical inputs for cellar protocol design.
Mostsäuredaten
Säureanalysedaten
Säuredaten
Säuregehaltsdaten
Säurezusammensetzungsdaten
acid analysis data
acid balance data
acid composition data
must acidity data
organic acid data
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed, Wiley, pp.: 3-45 Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 395-420 Iland P et al. (2011) Chemical Analysis of Grapes and Wine, 2nd Ed, Patrick Iland Wine Promotions, pp.: 55-75
Acidity Data
Acidity perception for Chablis Grand Cru (2023 vintage, no MLF) evaluates as "high" — the preserved malic acid component (3.5 g/L malic, 5.2 g/L tartaric) and low pH (3.08) produce a penetrating, incisive acidity with a distinctly green-apple malic edge that provides the hallmark Chablis freshness and drives the wine's 15+ year aging trajectory. Acidity perception for Châteauneuf-du-Pape (2022 vintage, full MLF) records "medium-minus to medium" — the completed malolactic conversion (malic acid reduced from 2.8 to <0.1 g/L, replaced by lactic acid) combined with the elevated pH (3.78) produces a soft, rounded acid profile perceived as gentle warmth rather than sourness, consistent with the southern Rhône red wine style. Acidity perception for Grüner Veltliner Smaragd from the Wachau (2024 vintage, no MLF) assesses at "medium-plus" — the firm tartaric acid backbone (TA 5.8 g/L) and moderately low pH (3.22) create a vibrant, racy freshness that defines the Wachau Smaragd style, with the preserved malic acid contributing citrus-like crispness.
Acidity perception is a taste description entity that records the perceived sourness, freshness, and structural contribution of organic acids on the wine palate, assessed on a structured scale from flat/flabby (insufficient acidity, lacking freshness) through soft, medium-minus, medium, medium-plus, and high to sharp/aggressive (excessive perceived sourness dominating the palate balance), integrating the contributions of titratable acidity concentration, pH level, specific acid species composition (tartaric providing firm crisp acidity, malic providing green-apple sharp acidity, lactic providing soft rounded acidity, succinic providing bitter-sour complexity), and the modulating effects of residual sugar, alcohol, and tannin on sourness perception.
Acidity perception in wine evaluation illustrates one of the most instructive disconnects between analytical measurement and sensory experience, as perceived sourness correlates imperfectly with either titratable acidity or pH taken independently — the perception is a function of both parameters together, modified by the buffering capacity of the wine and the presence of other taste-active compounds. The psychophysical relationship between hydrogen ion concentration and perceived sourness follows a power function with an exponent of approximately 0.85 for citric acid solutions, but in wine, the complexity of the acid mixture and the presence of taste-modifying compounds (sugar, alcohol, tannin, minerals) substantially modify this relationship. The specific acid species composition profoundly influences the quality of perceived acidity — tartaric acid contributes a firm, clean, crisp sourness perceived primarily on the sides of the tongue, malic acid produces a sharper, more angular sourness with a green-apple quality often described as "aggressive" in excess, lactic acid (from MLF) contributes a softer, rounder, almost creamy sourness with less intensity per gram than tartaric or malic, and succinic acid (a fermentation product at 0.5-1.5 g/L) adds a complex bitter-sour-salty dimension sometimes described as "mineral." The malolactic fermentation decision therefore represents a fundamental acidity perception transformation — the conversion of sharp diprotic malic acid to soft monoprotic lactic acid reduces perceived acid intensity beyond the mere TA reduction, as the perceptual quality shifts from angular to rounded. Temperature modulates acidity perception significantly — cooler serving temperatures enhance perceived acidity (explaining the preference for serving crisp whites at 8-12°C to maximize freshness), while warmer temperatures soften acid impression (contributing to the softer palate perception of red wines served at 16-18°C). The concept of acidity as "backbone" or "structure" in wine vocabulary reflects the recognition that appropriate acidity provides the architectural framework upon which other taste elements (fruit sweetness, tannin, body) are organized — wines lacking sufficient acidity are described as "flabby" or "flat" regardless of other quality attributes.
Frischeeindruck
Gaumen-Säure
Säurebeurteilung
Säureeindruck
Säurewahrnehmung
acid impression
acidity assessment
freshness assessment
palate acidity
sourness perception
Peynaud E, Blouin J (2006) The Taste of Wine, 4th Ed, Wiley, pp.: 120-155 Sowalsky RA, Noble AC (1998) Comparison of the effects of concentration, pH and anion species on astringency and sourness of organic acids. Chemical Senses 23(3): 343-349 Waterhouse AL et al. (2016) Understanding Wine Chemistry, Wiley, pp.: 15-35 Fischer U, Noble AC (1994) The effect of ethanol, catechin concentration, and pH on sourness and bitterness of wine. American Journal of Enology and Viticulture 45(1): 6-10
Acidity Perception
Acidity persistence comparison between Loire Valley Sancerre and Languedoc Viognier using time-intensity methodology reveals persistence (time from maximum to return to threshold) of 24±4 seconds for Sancerre (7.2 g/L TA, pH 3.18) versus 12±3 seconds for Viognier (5.4 g/L TA, pH 3.55), with the 100% longer persistence in Sancerre contributing to its reputation as a superior food wine where the sustained acidity cleanses the palate between bites of goat cheese, compared to the quickly fading acidity of Viognier that provides less effective palate refreshment.
Acidity persistence quality is a sourness quality that represents the duration of sour taste perception following swallowing or expectoration, characterizing the lingering acidity that continues after the primary taste stimulus has been removed and contributes to the finish and aftertaste of grape and wine evaluation. Within the BFO framework, this quality specifically addresses the temporal decay dimension of sourness, distinguishing the duration of perception from peak intensity and qualitative character, recognizing that persistent acidity fundamentally shapes the finish quality and cleansing effect that affects food pairing suitability. The ontological significance lies in acidity persistence contributing to perceived freshness on the finish, where extended sourness provides the palate-cleansing quality valued in food pairing contexts and maintains liveliness through the aftertaste period, while rapidly fading acidity can leave a flat or cloying finish particularly in wines with residual sweetness. The physiological basis for acidity persistence involves both peripheral factors (residual acid in saliva continuing to stimulate receptors, acid-induced salivation maintaining sourness perception) and central processing (neural adaptation rates for sour taste). The compositional factors affecting acidity persistence include acid type (with some evidence that tartaric acid shows longer persistence than malic or lactic), total acidity level, and pH, though the relationships are not as well characterized as for sweetness persistence. In wine evaluation, acidity persistence interacts with other finish components, particularly tannin astringency in red wines where acidity can either complement or compete with the drying sensation depending on relative intensities and timing, affecting the perceived harmony of the finish. The correlation between acidity persistence and quality perception is generally positive, with longer persistence associated with greater complexity and food pairing versatility, though excessive persistence may indicate imbalance. Understanding acidity persistence quality enables characterization of finish quality beyond intensity alone, prediction of food pairing success based on palate-cleansing capacity, and wine style optimization considering the complete temporal trajectory of sourness.
Acidity_Persistence_Quality addresses the duration of sourness perception after stimulus removal, completing the temporal characterization alongside Perceived_Acidity_Intensity_Quality and Acidity_Character_Quality. This quality is particularly relevant for wine finish evaluation and food pairing considerations. Measurement requires time-intensity protocols, with persistence defined as time to return to threshold or specified percentage of maximum.
Säuredauer
Säurenachklang
Säurepersistenz
acid aftertaste
acidity finish
anhaltende Säure
lingering sourness
sourness duration
Lesschaeve I (2007) Sensory evaluation of wine and commercial realities: review of current practices and perspectives. American Journal of Enology and Viticulture 58(2): 252-258 Ross CF et al. (2008) Influence of serving temperature on the sensory properties of wine. American Journal of Enology and Viticulture 59(1): 76-80
Acidity Persistence Quality
Acidity retention at harvest varies from 4-5 g/L TA (pH 3.6-4.0) in warm climate, low-retention cultivars to 9-12 g/L TA (pH 2.9-3.2) in cool climate, high-retention types. Cultivar effects on retention are substantial, with Barbera maintaining 7-9 g/L TA in warm Italian climates while Chasselas in the same region shows only 4-5 g/L. Retention reflects the net balance of stable tartrate concentration, residual malate after degradation, and dilution or concentration effects from berry water status. Potassium accumulation independently affects pH (higher K raises pH at equivalent TA) through titration of organic acids to their potassium salts. The TA/pH relationship varies among cultivars, with high-retention types showing lower pH at equivalent TA due to higher tartrate/malate ratios.
An acid metabolism disposition that is the inherited capacity of a Vitis vinifera plant to maintain total titratable acidity and low pH in berries through harvest maturity, determined by the combined effects of tartaric acid concentration, malic acid degradation rate, potassium accumulation (affecting pH), and berry water relations affecting acid concentration.
Acidity_Retention_Disposition serves as the parent class for high and low retention categories. Retention integrates multiple underlying processes affecting final acidity, making it a complex trait with moderate heritability but strong environmental modulation. The distinction between TA (total titratable acid, measuring acid quantity) and pH (measuring hydrogen ion activity) is important, as potassium effects can decouple these parameters. Wine style requirements vary in acidity targets, with sparkling wines requiring high retention and some red styles tolerating lower acidity.
Azidität-Bewahrung
Säureerhalt
Säureretention
TA retention trait
acid preservation capacity
acidity maintenance
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 378-385 Volschenk H et al. (2006) Malic acid in wine: origin, function and metabolism during vinification. South African Journal of Enology and Viticulture 27:123-136
Acidity Retention Disposition
Merlot active abscission zone at harvest maturity (BBCH 89, 24°Brix) shows cellulase gene expression increased 18-25 fold, polygalacturonase enzyme activity elevated 12-18 fold compared to véraison levels, separation layer cells with 40-60% reduced middle lamella thickness, progressive berry detachment force declining from 1.8 N (7 days pre-harvest) to 1.0 N (at harvest), and ethylene production at 0.3-0.6 μL/L in cluster microenvironment
An active abscission zone is a pedicel abscission zone characterized by ongoing expression of abscission-associated genes, accumulation and secretion of cell wall degrading enzymes, and progressive cell wall modification, representing the abscission zone during the activation and early separation phases before physical berry detachment occurs. Active abscission zone development typically occurs during advanced berry ripening (BBCH 87-89) when ethylene production and declining auxin levels trigger abscission zone activation. Molecular and biochemical characteristics of active abscission zone include= elevated expression of abscission-specific genes (cellulase genes, polygalacturonase genes, genes encoding abscission-associated transcription factors increased 10-50 fold compared to non-abscission tissues), synthesis and accumulation of cell wall hydrolytic enzymes (cellulases targeting cellulose in primary walls, polygalacturonases and pectinases targeting pectin in middle lamella, xyloglucan endotransglycosylases modifying hemicellulose), secretion of enzymes into cell wall spaces (enzymes localized to middle lamella and cell wall regions between separation layer cells), progressive cell wall degradation (middle lamella dissolution, primary wall thinning, reduced cell-cell adhesion), and hormone signaling (ethylene signal transduction active, auxin levels declining removing abscission suppression, potential abscisic acid involvement). Cellular changes in active abscission zone include= separation layer cells showing swelling and cell wall softening, dissolution of middle lamella creating incipient separation planes, cells remaining viable and metabolically active during separation process (unlike programmed cell death mechanisms), protective layer cells undergoing suberization or lignification in preparation for post-abscission protection, and vascular tissues showing reduced transport activity as separation progresses. Active abscission zone enzymatic activity shows characteristic progression= early activation phase (initial enzyme gene expression and protein synthesis, 1-3 days before detectable cell wall changes), middle activation phase (enzyme accumulation and secretion, beginning of cell wall modification, 0.5-2 days before physical separation), late activation phase (extensive cell wall degradation, cells beginning to round up and separate, 0-12 hours before detachment), with timing varying by cultivar, ripening stage, and environmental conditions. Factors regulating active abscission zone progression include= ethylene concentration (ripening ethylene production at 0.1-1 μL/L in cluster atmosphere sufficient to trigger and maintain activation), auxin depletion (declining IAA levels in berry and pedicel removing suppression), carbohydrate status (sugar accumulation in berry may signal maturity triggering activation), calcium levels (calcium depletion from cell walls may facilitate pectin solubilization), and transcriptional regulation (ethylene-responsive transcription factors activating abscission gene cascades). Active abscission zone can be distinguished from inactive or dormant abscission zones by= presence of active transcription and translation of abscission genes, accumulation of cell wall modifying enzymes, progressive reduction in berry detachment force (declining from 1.5-2.5 N in inactive zones to 0.5-1.5 N in active zones), and histochemical detection of enzyme activities in separation layer.
Active_Abscission_Zone emphasizes ongoing enzymatic activity and cell wall modification and is distinguished from Pedicel_Abscission_Zone (complete structure and development) and Separating_Abscission_Zone (physical cell separation process) by focus on molecular and biochemical activation preceding physical separation. Critical for understanding harvest timing optimization, ethylene and hormone regulation of ripening and abscission coordination, and potential manipulation of abscission timing through viticultural or genetic approaches.
Activated separation zone
Aktive Trennungszone
Aktivierte Ablösungsschicht
Enzymaktive Abszissionszone
Enzyme-active abscission zone
Patterson, S. E. (2001). Cutting loose. Abscission and dehiscence in Arabidopsis. Plant Physiology, 126(2), 494-500. | Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press.
Active Abscission Zone
In a Paso Robles Grenache vineyard, active acid degradation was documented over 16 days in late August Year N+1 (BBCH 84-85) when titratable acidity declined from 8.6 to 3.8 g/L (average 0.52 g/L per day peak rate), malic acid decreased from 5.2 to 1.4 g/L (0.42 g/L per day loss), pH increased from 3.28 to 3.64 (0.025 units per day average), berry Brix rose from 21.8° to 26.5°, under hot California interior conditions (daily high 33-37°C accelerating degradation, requiring early harvest at lower Brix to preserve minimal acid balance).
Active_Acid_Degradation represents the peak phase of organic acid catabolism during mid-to-late veraison when enzymatic machinery operates at maximum capacity producing the most rapid acid decline that fundamentally transforms berry composition creating the steepest gradient in titratable acidity and pH changes defining the critical window for achieving final acid balance. This active phase occurs during mid-to-late veraison through early ripening (BBCH 84-85), typically 12-25 days post-veraison onset depending on temperature and cultivar, and is characterized by titratable acidity declining at maximum rates 0.35-0.65 g/L per day (compared to initial phase 0.18-0.35 g/L per day and late phase 0.10-0.25 g/L per day), malic acid showing peak degradation 0.30-0.55 g/L per day representing 3-6% of remaining malate per day under warm conditions, pH increasing most rapidly 0.025-0.050 units per day (0.18-0.35 units per week) creating dramatic compositional shift, and respiratory malate contribution reaching maximum 50-70% of total berry CO₂ production indicating acid catabolism dominates energy metabolism during this critical period. Enzymatic basis of active degradation includes malic enzyme activities sustained at peak levels with NADP-malic enzyme 40-70 units/g fresh weight (10-15 fold above green berry), NAD-malic enzyme 12-28 units/g contributing secondarily, combined capacity 55-95 units/g providing enzymatic machinery adequate for observed maximum rates, citric acid cycle enzyme activities maintained or slightly elevated 15-35% above early veraison ensuring pathway flux for complete malate oxidation, and respiratory chain complexes operating at high capacity with cytochrome c oxidase 12-22 units/g enabling efficient electron transport and ATP synthesis supporting active metabolism. Substrate availability during active degradation shows vacuolar malate declining from 5-9 g/L at phase onset to 2-5 g/L at conclusion representing 40-60% depletion of phase-entry levels, cytoplasmic malate maintained at 8-18 mM through continued tonoplast efflux balancing consumption, tonoplast transport capacity adequate providing 2-4 μmol malate g⁻¹ h⁻¹ efflux from vacuole matching or exceeding cytoplasmic consumption rate, and substrate availability still non-limiting during active phase with vacuolar pools sufficient enabling maximum enzyme velocity operation without kinetic constraints. Temporal dynamics of active degradation demonstrate initial acceleration from early veraison 0.20-0.35 g/L per day to peak active rates 0.40-0.65 g/L per day over 5-10 days representing 1.8-2.5 fold rate enhancement, sustained maximum rates 10-20 days duration (temperature and cultivar dependent) constituting critical window for acid loss, then deceleration as substrate depletes reducing to late phase 0.12-0.28 g/L per day over subsequent 8-15 days, and cumulative loss during active phase potentially 6-11 g/L malate representing 50-75% of total ripening period loss concentrated in this brief intensive period. Environmental factors critically affecting active degradation include temperature showing strongest effects during this phase with warm conditions 28-32°C producing maximum rates 0.50-0.70 g/L per day malate loss potentially depleting acids excessively, moderate warmth 25-28°C enabling appropriate rates 0.35-0.50 g/L per day achieving balanced final composition, cooler conditions 20-25°C moderating to 0.25-0.40 g/L per day preserving acids beneficial in low-acid cultivars, heat stress >32-35°C causing excessive degradation 0.60-0.90 g/L per day creating risk of over-deacidification producing unbalanced fruit, and cumulative heat exposure during active phase critically determining final acid levels with degree-days strongly correlating with total loss r² = 0.82-0.94. Diurnal temperature patterns during active phase show daytime peak degradation 0.50-0.80 g/L per day during warmest hours (typically 2-6 PM) when berry temperature maximum 30-38°C from solar heating, nighttime minimum rates 0.15-0.35 g/L per day as berry cools to near air temperature 15-22°C, daily integrated loss averaging 0.35-0.60 g/L representing balance of day-night extremes, and regions with large diurnal range (>15-20°C) showing moderated total loss despite hot days through cool night respite enabling acid preservation compared to constantly warm climates. Cluster microclimate effects particularly pronounced during active phase with sun-exposed clusters experiencing 45-85% faster degradation than shaded fruit due to 5-12°C temperature differential, canopy position showing substantial variation with exterior southwestern clusters (northern hemisphere) losing acids 50-100% faster than interior northeastern positions, and targeted shading interventions most effective during active phase reducing degradation 30-55% through berry temperature moderation 4-8°C cooling preserving 3-6 g/L additional malate at harvest. Water status interactions during active phase demonstrate adequate vine hydration supporting normal metabolic function enabling appropriate degradation without stress enhancement, moderate deficit (stem water potential -1.0 to -1.4 MPa) generally compatible with normal rates though effects variable across cultivars with some showing slight acceleration through stress-enhanced respiration while others unaffected, severe deficit (3.5 substantially increasing spoilage risk), buffer capacity declining from 14-20 mL to 9-14 mL of 0.1 N NaOH per 10 mL juice as total organic acids reduce making berry pH more labile, malate:tartrate ratio declining from 0.8-1.5:1 to 0.3-0.9:1 fundamentally altering acid composition character, and osmotic potential effects minor with acid contribution 3.7 creating flat wines lacking structure with reduced color stability in reds and increased microbial vulnerability requiring careful harvest timing or winery intervention, inadequate active degradation in cool vintages maintaining high acids TA >8-10 g/L producing overly tart fruit requiring extended ripening for acid reduction or winery deacidification, spatial variation in active degradation across vineyard creating maturity heterogeneity with some zones over-deacidified while others retain acids complicating harvest decisions and requiring selective management, and balance achievement between sufficient acid decline and retention of adequate acidity for wine quality requiring integrated monitoring and management throughout active phase. Harvest timing considerations during active phase involve recognition that acids declining most rapidly during this period requiring frequent monitoring (2-3 times per week) to track trajectory, integration with sugar accumulation assessment ensuring appropriate sweetness-acidity balance achieved simultaneously, pH monitoring critically important as rapid increase during active phase can reach undesirable levels before adequate sugar accumulation in warm conditions, phenolic maturity assessment particularly reds ensuring tannin polymerization and seed maturation synchronized with sugar-acid development, and flexibility in harvest date potentially valuable as 3-7 day window during active phase can produce substantially different final composition. Viticultural management during active degradation includes canopy interventions with strategic cluster shading most effective during this phase as temperature sensitivity highest, irrigation management potentially moderating degradation through berry cooling though mechanisms complex and requiring careful timing to avoid negative vigor or dilution effects, crop load considerations with overcropping potentially accelerating active degradation through stress while optimal loads enabling appropriate rates, disease monitoring critical as changing berry composition (declining acids, increasing sugars, softening) increases vulnerability to Botrytis, sour rot, and other pathogens requiring preventive protection, and harvest preparation advancing as active degradation indicates approaching maturity requiring readiness for prompt picking when optimal parameters achieved. Monitoring strategies during active phase involve frequent berry sampling every 2-4 days tracking acid decline trajectory enabling forecasting of harvest timing, combined assessment of TA, pH, malate, and tartrate providing comprehensive acid profile, integration with Brix, phenolics, and sensory evaluation ensuring holistic maturity determination, spatial sampling across vineyard zones revealing heterogeneity requiring selective management, and historical data compilation enabling year-to-year comparison and predictive modeling of active phase dynamics informing future management decisions.
Active_Acid_Degradation is distinguished from Initial_Acid_Reduction by representing peak (0.35-0.65 g/L per day TA) rather than early phase (0.18-0.35 g/L per day), from Malic_Acid_Decline_Start by focusing on maximum rate period rather than overall process, and from late decline by showing peak rather than deceleration. Duration 10-20 days BBCH 84-85. Temperature critically important: 28-32°C maximum rates, >32°C excessive. Malate contribution 50-70% respiratory CO₂. Critical window determining final acid balance. Active_Acid_Degradation represents most intensive period of compositional transformation during ripening.
Spitzen-Säure-Katabolismus-Phase
intensive Malat-Abbau-Periode
intensive malate breakdown period
maximale organische Säure-Degradation
maximum organic acid degradation
peak acid catabolism phase
rapid deacidification phase
rapide Entsäuerungs-Phase
Keller M (2020) The Science of Grapevines 3rd Ed pages 229-233 peak acid decline Sweetman C, Sadras VO, Hancock RD, Soole KL, Ford CM (2014) Metabolic effects of elevated temperature on organic acid degradation in ripening Vitis vinifera fruit J Exp Bot 65:5975-5988 temperature effects during peak phase Lakso AN, Kliewer WM (1975) The influence of temperature on malic acid metabolism in grape berries Plant Physiol 56:370-372 peak degradation rates.
Active Acid Degradation
In Pinot Noir berries in Willamette Valley, Oregon during BBCH 85-87, active acid reduction proceeds at 0.32 g/L per day over 25 days from day 12 to day 37 post-veraison, reducing titratable acidity from 14.2 to 6.2 g/L with malic acid declining from 7.8 to 2.6 g/L under mean temperatures of 19°C, while a late-season heat event of 35°C for 5 days accelerates reduction rate to 0.55 g/L per day.
Active acid reduction represents the dynamic phase of organic acid catabolism in Vitis vinifera berries during mid-to-late ripening wherein enzymatic degradation and respiratory consumption actively decrease malic acid content at rates of 0.2-0.5 g/L per day, occurring from approximately 10-15 days post-veraison until 7-14 days before harvest when acid levels approach equilibrium values. This process encompasses the coordinated activity of multiple metabolic pathways including mitochondrial respiration of malate through the tricarboxylic acid cycle, cytosolic NADP-malic enzyme conversion of malate to pyruvate with CO2 release, and gluconeogenic recycling of malate carbon skeletons into hexose sugars, collectively accounting for the consumption of 4-8 g/L malic acid during the active reduction period spanning 20-35 days. The cellular biochemistry involves malate transport from vacuolar storage across the tonoplast membrane via specific malate channels, followed by cytosolic metabolism or mitochondrial import through the dicarboxylate carrier, with respiratory flux rates increasing 3-5 fold during active reduction compared to pre-veraison levels. Temperature sensitivity characterizes active acid reduction, with Q10 values of 2.0-2.5 indicating that enzymatic reaction rates approximately double for each 10°C temperature increase within the physiological range of 15-35°C, explaining accelerated acid loss during heat waves and the pronounced diurnal variation in malate metabolism linked to day-night temperature differentials. The process shows strong cultivar specificity, with high-acid varieties such as Riesling maintaining active reduction rates of 0.15-0.25 g/L per day compared to rapid-decline varieties such as Grenache showing rates of 0.4-0.6 g/L per day under equivalent thermal conditions. Hormonal regulation involves abscisic acid enhancement of malic enzyme gene expression and auxin modulation of vacuolar malate efflux, integrating acid reduction with broader ripening coordination. Berry position within the cluster influences active reduction rates through differential temperature exposure and vascular connection strength, with exposed exterior berries typically showing 15-25% faster acid decline than shaded interior berries. The process contributes substantially to berry carbon balance during ripening, with malate-derived carbon representing 5-10% of total respiratory substrate and contributing to the metabolic energy requirements of sugar accumulation and secondary metabolite synthesis. Monitoring active acid reduction through weekly titratable acidity measurements enables prediction of harvest acid levels and optimal picking decisions balancing sugar accumulation against acid retention for specific wine styles.
Active_Acid_Reduction is parent class under Acid_Reduction representing the dynamic enzymatic phase. Distinguishes from Final_Acid_Balance which represents equilibrium state. Duration 20-35 days during mid-to-late ripening. Contains subclasses Continuing_Acid_Decline, Malate_Degradation, and pH_Increase representing concurrent aspects of active reduction. Rate parameter 0.2-0.5 g/L per day distinguishes active from plateau phases.
active malic acid consumption
aktive Säure-Reduktion
aktiver Äpfelsäure-Abbau
dynamic acid catabolism
dynamischer Säure-Katabolismus
enzymatic acid decline
enzymatische Säure-Abnahme
ongoing acid reduction
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 268-275 Ruffner HP (1982) Metabolism of tartaric and malic acids in Vitis. Vitis 21: 346-358 Sweetman C et al (2009) Regulation of malate metabolism. Molecular Plant 2: 1270-1280
Active Acid Reduction
Chardonnay in Champagne shows active berry cell division during late June-early July Year N+1 at BBCH 71-72, with division activation days 3-7 from auxin signaling, mitotic activity onset week 1, periclinal and anticlinal divisions proceeding with 12-15 hour cycle times, progressive cell number increase 6-12 fold over weeks 0-3, under cool maritime conditions of 19-24°C temperature, typical of this early-bursting cultivar initiating division establishing proliferative program
Active berry cell division represents the early intensive mitotic phase during berry cell division when division first activates in quiescent pericarp cells, mitotic activity onset establishes proliferative program, and multiple division rounds proceed creating progressive cell number increase, spanning approximately first 2-3 weeks post-fertilization as division initiates and accelerates before peak intensity. This initial proliferation phase occurs during early summer in late June-early July in temperate regions Year N+1, proceeding over weeks 0-3 post-fertilization as division establishes, corresponding to BBCH 71-72 young fruit early stages. Active division mechanism involves initial division activation with auxin signaling triggering cell cycle entry from quiescent state, mitotic activity onset as first divisions occur establishing proliferative program, periclinal cell divisions parallel to berry surface increasing tissue thickness, anticlinal divisions perpendicular to surface expanding tissue area, and coordinated division creating organized cellular architecture. Initial division activation shows auxin from developing seeds reaching threshold levels stimulating cell cycle entry, auxin receptor signaling cascades activating transcription factors, cell cycle gene expression initiating including cyclins and cyclin-dependent kinases, and G0 to G1 phase transition enabling DNA replication preparation. Division activation timing shows auxin signaling establishing days 2-5 post-fertilization, first cell cycle entries occurring days 3-7, and progressive activation as more cells enter cycle creating wave of proliferation. Mitotic activity onset shows first observable divisions occurring within first week post-fertilization, division rate accelerating as cell populations enter proliferative state, exponential cell number increase beginning as divisions compound, and establishment of active mitotic program characterizing berry as proliferative tissue. Early division characteristics show relatively asynchronous division across cell populations, progressive cell cycle entry creating temporal spread, division rate accelerating over first 2 weeks, and transition toward peak synchronous division by week 2-3. Cell cycle progression shows typical sequence of G1 growth phase, S phase DNA replication doubling chromosome number, G2 preparation phase, and M phase mitosis producing two daughter cells, with cycle times initially 12-16 hours during activation declining to 10-12 hours as activity increases. Periclinal cell division shows divisions with division plane parallel to berry surface, daughter cells positioned radially creating tissue thickness increase, periclinal divisions predominating in inner mesocarp regions, and divisions contributing to radial berry expansion. Anticlinal cell division shows divisions with plane perpendicular to surface, daughter cells positioned tangentially increasing tissue area, anticlinal divisions predominating in outer exocarp regions maintaining surface coverage, and divisions enabling tangential berry expansion. Division plane regulation shows orientation control through cytoskeletal organization, preprophase band microtubules predicting division plane, coordinated plane regulation creating organized tissue architecture, and spatial division patterns characteristic of tissue type and position. Cell number increase during active phase shows typical 4-16 fold accumulation over first 2-3 weeks, exponential growth from repeated division rounds, progressive increase in division-competent cell population, and acceleration toward peak activity. Tissue organization during active division shows coordinated division maintaining pericarp layer proportions, exocarp surface divisions preserving skin coverage during expansion, mesocarp bulk divisions creating flesh cellularity, and endocarp divisions near seeds, with overall organized growth producing functional berry architecture. Auxin gradient effects show higher auxin near seeds potentially stimulating greater division, spatial hormone distribution possibly affecting regional division patterns, and overall hormone signal coordinating proliferation across tissues. Division activation heterogeneity shows variation in activation timing across cell populations, some cells entering cycle earlier than others, progressive recruitment creating temporal spread, and heterogeneity contributing to extended division period. Environmental effects during active division include temperature affecting division rate with optimal 24-28°C supporting rapid activation and proliferation, cool conditions slowing activation and division, and temperature during critical first 2 weeks affecting early cell number accumulation. Seed development coordination shows seeds developing in parallel providing continuing auxin stimulus, seed size increasing correlating with berry growth, and seed-berry coordination ensuring developmental synchronization. The process shows three distinguishable aspects including initial division activation establishing cell cycle entry, mitotic activity onset with first divisions occurring, and periclinal cell division as major division mode. Duration typically 2-3 weeks representing early division establishment before peak. Regional timing shows active berry cell division during late June-early July across climates as division initiates post-fertilization. Viticultural significance includes establishing proliferative program determining subsequent cell number achievement, demonstrating auxin stimulation mechanism, showing temperature sensitivity during critical activation period, and initiating cell number accumulation determining berry size. Research applications include analyzing division activation mechanisms and signaling, investigating cell cycle regulation during activation, evaluating division plane control and tissue organization, quantifying early cell number accumulation dynamics, and assessing active division as determinant of peak proliferation and final cellularity.
Active_Berry_Cell_Division is first phase of Berry_Cell_Division_Phase, distinguished by division activation and establishment. Subdivided into Initial_Division_Activation with auxin-triggered cell cycle entry, Mitotic_Activity_Onset with first divisions occurring, and Periclinal_Cell_Division emphasizing radial tissue growth. Duration 2-3 weeks representing early division establishment. Cell cycle times 12-16 hours initially declining to 10-12 hours. Exponential cell number increase beginning. Precedes Peak_Division_Activity. Establishes proliferative program determining subsequent cellularity.
Teilungs-Aktivierungsperiode
aktive Beeren-Zellteilung
division activation period
early cell division
early mitotic phase
frühe mitotische Phase
initial proliferation phase
initiale Proliferationsphase
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 432-435 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Active Berry Cell Division
An active bud at BBCH stage 07 on Vitis vinifera cv. Sauvignon Blanc in early April in Marlborough, New Zealand, showing pronounced swelling with bud scales separated 3-4mm, green shoot tip visible, first leaf beginning to unfold, occurring 12 days after budbreak initiation when growing degree day accumulation (base 10°C) reached 85 units following satisfaction of 950 chilling hours during winter dormancy.
A bud that has exited dormancy and resumed active growth and development, characterized by visible swelling, separation of bud scales, emergence of green shoot tissues, active cell division and elongation in apical meristems, and high metabolic activity including protein synthesis, respiration, and resource mobilization from storage tissues. Active buds represent the transition from quiescent to growing state, responding to accumulated chilling satisfaction and heat unit accumulation (growing degree days), with development proceeding through distinct BBCH stages from initial swelling through leaf unfolding to shoot elongation, ultimately producing either vegetative shoots, reproductive shoots bearing inflorescences, or mixed shoots with both vegetative and reproductive components depending on primordial differentiation established during the previous growing season.
Active buds are distinguished from dormant buds by resumption of growth rather than dormancy state, and from latent buds which remain suppressed despite being physiologically capable of growth. The transition from dormant to active involves complex hormonal shifts, particularly declining ABA and increasing gibberellin and cytokinin levels. Active bud development is the most frost-vulnerable period in the grapevine annual cycle, with damage risk increasing as development progresses through successive BBCH stages.
aktive Knospe
austreibende Knospe
breaking bud
growing bud
schwellende Knospe
swelling bud
Based on Lorenz et al. (1995) "Growth Stages of the Grapevine= Phenological growth stages of the grapevine (Vitis vinifera L. ssp. vinifera) - Codes and descriptions according to the extended BBCH scale" Australian Journal of Grape and Wine Research 1=100-103 Keller (2010) "Managing grapevines to optimise fruit development in a challenging environment= a climate change primer for viticulturists" Australian Journal of Grape and Wine Research 16=56-69 Zapata et al. (2004) "Estimation of the base temperature and growth phase duration in terms of thermal time for four grapevine cultivars" Acta Horticulturae 640=179-186.
Active Bud
Grenache in Châteauneuf-du-Pape exhibits active bud breaking following BBCH 07 in mid-April after 800-900 chilling hours and 350-400 growing degree days base 10°C, with shoot extending 8-12 mm beyond scales and first leaves beginning to separate over 4-5 days under conditions of 17-21°C mean temperature, 45-55% relative humidity, and 14-hour photoperiod, representing typical Mediterranean climate progression for this late-season cultivar
Active bud breaking represents the progressive emergence phase where green shoot extends increasingly beyond protective scales, characterized by rapid shoot elongation and beginning leaf unfolding as developmental momentum accelerates toward independent shoot growth. This intermediate breaking stage occurs after initial emergence has breached scale barrier, with exposed shoot tissue extending 5-15 mm beyond scale margins representing 30-60% of total shoot emergence from protective structures. The shoot shows bright green coloration as chlorophyll synthesis proceeds actively in light-exposed tissues, creating visible contrast with pale yellow-green of newly emerging portions still transitioning from enclosed environment. Shoot elongation proceeds at 2-5 mm per day through coordinated cell division in apical meristem and cell expansion in elongation zone immediately behind meristem, with cell production maintaining 18-24 hour division cycles while expansion proceeds rapidly through turgor-driven cell wall extension. The apical meristem remains protected at shoot tip continuing to produce new cells that will form additional shoot axis, leaves, and other organs, while cells produced earlier enter expansion and differentiation phases creating observable elongation. First true leaves begin unfolding from their tightly compressed primordia configuration, initially remaining appressed along shoot axis then progressively spreading perpendicular to axis as cell expansion and differential growth patterns establish characteristic leaf orientation. Leaf primordia expand from their 3-5 mm initial size toward 10-15 mm length during active breaking, with leaflets of compound grapevine leaf beginning to separate from each other though full palmate configuration is not yet achieved. Bud scales progressively separate further and begin falling away from base upward, with some cultivars showing scales dropping completely while others maintain dry brown scale remnants attached at bud base throughout growing season. Vascular tissues extend through elongating shoot with new xylem and phloem cells differentiating continuously to maintain functional connection from permanent woody structures through expanding shoot to developing leaves, enabling sustained water transport for growth and beginning transpiration from increasing surface area. Chloroplast development progresses in leaf cells with chlorophyll accumulating and photosynthetic machinery assembling, though photosynthetic rate remains below respiration rate meaning net carbon balance is still negative and growth depends primarily on stored carbohydrate reserves mobilized from woody tissues. Metabolic activity transitions gradually from peak respiration toward increasing photosynthetic contribution, with respiration remaining at 10-15 fold above dormancy baseline while photosynthetic capacity increases from negligible toward 20-30% of mature leaf rates. Stored carbohydrates continue flowing from woody tissue reserves through phloem to support rapid shoot growth, with starch and sugar concentrations declining measurably in canes as reserves mobilize to developing shoots. Protein synthesis continues at high rates producing structural components for new cells, enzymes supporting rapid metabolism, membrane proteins for expanding cells, and regulatory proteins coordinating complex developmental processes. Hormonal regulation maintains elevated cytokinin and gibberellin supporting continued cell division and expansion, with auxin concentration highest at shoot apex establishing apical dominance and coordinating leaf and tendril differentiation through concentration gradients. Temperature remains primary rate-determining factor with development doubling per 10°C between 10-25°C base temperatures, creating strong temperature dependence in progression rate through active breaking phase. Water demand increases substantially as transpiring leaf area expands, requiring adequate soil moisture and functional root water uptake to prevent water stress that could slow or temporarily arrest development. Light intensity begins affecting development as chloroplasts become functional, with higher light promoting leaf expansion and chlorophyll synthesis though shade does not prevent progression at this stage. The process typically requires 3-5 days under favorable spring conditions with consistent temperatures above 15°C, with cooler or fluctuating conditions extending duration to 5-8 days. Cultivar variation appears in shoot elongation rate and leaf expansion timing, with vigorous varieties showing 4-6 mm per day elongation while less vigorous types achieve 2-3 mm per day progression. Regional timing follows initial breaking by 2-4 days, occurring late April in warm climates, early May in temperate regions, and mid-May in cool areas. Viticultural management considerations include maintaining frost protection readiness as vulnerable tissue amount increases substantially with expanding shoot and leaves, avoiding mechanical operations that could damage elongating shoots, initiating canopy positioning preparations for eventual shoot training, and implementing pest monitoring for insects and mites colonizing expanding leaf area. Frost damage during active breaking kills exposed shoot and developing leaves through ice formation, forcing compensatory growth from secondary or basal buds resulting in 1-2 week delays and reduced fruitfulness. Research applications include quantifying cell division and expansion rates during rapid shoot growth, analyzing metabolic transitions from stored reserve dependence to photosynthetic independence, documenting gene expression changes during organ differentiation and chloroplast development, and evaluating cultivar and environmental effects on breaking progression rates for phenological modeling.
Active_Bud_Breaking is the second of three Bud_Breaking subphases, distinguished by substantial shoot emergence with 30-60% exposed and beginning leaf unfolding. Transitions to Complete_Bud_Breaking when leaves fully separate from shoot axis. Represents transition from mechanical emergence to active shoot growth. Frost vulnerability remains very high though slightly reduced from initial breaking.
Blattspreizung
Triebstreckung
advancing shoot emergence
aktives Knospenaufbrechen
fortschreitender Austrieb
progressive bud break
shoot elongation phase
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 180-185 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 82-89 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Active Bud Breaking
Cabernet Sauvignon showing active feeder roots in April, soil cores 20-40 cm depth revealing feeder roots 0.6-1.4 mm diameter, vital staining (FDA) showing 72% of feeder roots viable indicating active status, microscopic examination revealing intact dense cytoplasm, functional root hairs, arbuscular mycorrhizal structures, isotope pulse-chase (15N-nitrate) demonstrating active feeder roots providing 88% of vine nitrogen uptake, active feeder root population estimated 850 meters per vine comprising functional absorption capacity
Active feeder root is feeder root in functional state actively absorbing water and nutrients, characterized by intact living tissues with functional membranes and metabolism, turgid appearance reflecting adequate cellular function, demonstrated uptake capacity through isotope tracing or physiological measurements, and typical age 1-6 months representing period from maturation through peak function before senescence, with active feeder roots showing dense cytoplasm particularly in root hairs and youngest tissues, abundant mitochondria providing ATP for active transport, functional plasma membrane ion transporters including H+-ATPases, channels, and carriers, close contact with soil particles and mycorrhizal hyphae maximizing absorption interfaces, and favorable carbon balance with photosynthate import supporting metabolism exceeding respiratory costs, and demonstrating that active status distinguishes functional roots currently contributing to uptake from developing roots not yet mature or senescing roots with declining function creating need to assess active feeder root population not merely total feeder roots when evaluating vine absorption capacity. Active feeder root age and developmental state includes= showing post-maturation age (active feeder roots typically 1-6 months post-initiation having completed development and maturation), exhibiting peak functional period (active state representing period of maximum absorption capacity), demonstrating pre-senescence status (active roots not yet showing senescence indicators maintaining full function), revealing variable duration (active period varying with root order, environmental conditions, and vine status), and displaying transition boundaries (active state bounded by maturation completion and senescence onset). Active feeder root physiological characteristics includes= showing intact membranes (plasma membrane and organellar membranes maintaining integrity and selective permeability), exhibiting active metabolism (respiration, ion pumping, biosynthesis proceeding normally), demonstrating adequate turgor (cells maintaining turgor pressure 0.3-0.6 MPa indicating osmotic function), revealing responsive capacity (active feeder roots capable of adjusting function in response to environmental signals or plant demands), and displaying positive carbon balance (photosynthate import exceeding respiratory consumption supporting growth and function). Active feeder root absorption performance includes= providing current uptake (active feeder roots performing majority of vine water and nutrient acquisition at any given time), exhibiting high specific rates (uptake per unit active feeder root surface area or length substantially higher than senescing or structural roots), demonstrating nutrient responsiveness (active feeder root uptake quickly responding to soil nutrient availability changes), revealing diurnal patterns (active feeder root function potentially showing daily variation correlating with transpiration and photosynthesis), and displaying seasonal maxima (active feeder root population and function peaking during favorable spring and fall periods). Active feeder root soil relationships includes= showing intimate soil contact (active feeder roots closely associated with soil particles maximizing exchange surface), exhibiting root hair function (abundant functional root hairs on active feeder roots extending into soil pores), demonstrating mycorrhizal activity (active arbuscular mycorrhizal associations in feeder root cortex with functional arbuscules), revealing rhizosphere development (active feeder roots creating and maintaining active rhizosphere with mucilage secretion and microbial stimulation), and displaying resource depletion (active feeder root uptake creating depletion zones around roots for mobile nutrients). Active feeder root in vine resource status includes= determining current acquisition (active feeder root population size and activity limiting current vine water and nutrient uptake), enabling growth support (adequate active feeder roots necessary for supporting vigorous shoot and fruit development), demonstrating stress response (active feeder root function critically affected by drought, temperature, or other stresses), revealing seasonal importance (active feeder root population particularly critical during rapid growth periods), and displaying management relevance (practices affecting active feeder root abundance or function directly impacting vine performance). Active feeder root population dynamics includes= showing continuous production (new active feeder roots continuously produced at root tips as roots grow and branch), exhibiting steady-state population (active feeder root numbers relatively stable as production balances senescence under constant conditions), demonstrating environmental modulation (active feeder root population increasing with favorable conditions decreasing under stress), revealing spatial distribution (active feeder roots concentrated in resource-rich soil zones and shallow depths), and displaying temporal variation (active feeder root abundance varying seasonally and among years). Active feeder root assessment methods includes= requiring viability determination (distinguishing active from senescent feeder roots through vital staining or physiological tests), showing isotope applications (pulse-labeling demonstrating active feeder root uptake capacity), exhibiting minirhizotron imaging (sequential imaging tracking individual feeder roots identifying active functional period), demonstrating molecular markers (gene expression or protein abundance indicating active metabolic state), and revealing functional measurements (direct uptake measurements confirming active absorption capacity). Active feeder root turnover contribution includes= representing temporary state (active feeder roots eventually transitioning to senescent state as roots age), showing functional lifespan (active period comprising majority of total feeder root lifespan with senescence relatively brief), exhibiting production requirement (maintaining active feeder root population requiring continuous new root production replacing senescing roots), demonstrating carbon investment (producing and maintaining active feeder roots requiring substantial carbohydrate allocation), and displaying soil carbon input (active feeder roots eventually senescing contributing organic matter to soil).
Active_Feeder_Root emphasizes functional absorbing state and is distinguished from senescent feeder roots by focus on living metabolism and uptake. Includes Absorbing_Active_Feeder_Root, Functional_Active_Feeder_Root, and Young_Active_Feeder_Root. Critical for understanding current uptake capacity, physiological status, absorption performance, population dynamics, and the living functional feeder roots providing majority of vine resource acquisition at any given time.
Aktive Feinwurzel
Functional feeder root
Funktionelle Saugwurzel
Lebende Absorptionswurzel
Living absorptive root
Viable fine root
Active Feeder Root
Cabernet Sauvignon active growth mid berries at BBCH 77 (weeks 6-8 post-fruit set, lag phase) maintain relatively constant dimensions at 9-10.5 mm diameter and 1-1.3 g fresh weight (increasing only 2-5% from BBCH 75 end of Phase I), green coloration with chlorophyll content declining from 0.8 to 0.5 mg/g during lag phase, firm texture, acidity at peak 22-24 g/L, sugars stable at 7-9%, while seeds undergo active maturation increasing from 60% to 90% of final dry weight (15-18 mg to 21-25 mg per seed), embryos completing development from torpedo to mature embryo stage, and seed coats completing lignification and browning. Merlot active growth mid berries examined at weekly intervals during lag phase (weeks 6-8) show= week 6 (9.2 mm diameter, 1.05 g, 2.6 million cells at 48 μm average diameter, seeds 4.8 mm length 14 mg dry weight), week 7 (9.4 mm, 1.1 g, cells 50 μm, seeds 5.5 mm 19 mg), week 8 (9.6 mm, 1.15 g, cells 52 μm, seeds 6 mm 23 mg), demonstrating minimal berry growth (4% diameter increase, 10% weight increase) but substantial seed growth (25% length increase, 64% dry weight increase) characterizing lag phase as period of active seed development within relatively static berry dimensions. Syrah active growth mid berries show transcriptional reprogramming during lag phase with microarray analysis revealing downregulation of cell cycle genes, cell wall biosynthesis genes, and auxin-responsive genes from Phase I levels, while ripening-related genes including sugar transporters, anthocyanin biosynthesis enzymes, and ABA-responsive genes show beginning upregulation creating molecular preparation for veraison transition despite minimal phenotypic changes during lag phase. Chardonnay experiencing moderate water deficit during lag phase shows unchanged berry dimensions (10.5-11 mm diameter, 1.3-1.5 g) comparable to well-watered vines, but accelerated veraison onset occurring 5-8 days earlier in water-deficit treatment (veraison at 58-62 days post-fruit set versus 65-72 days well-watered), demonstrating that lag phase water status affects developmental timing and veraison onset without necessarily affecting lag phase berry size.
An active growth mid berry is a mid developing berry in the lag phase (BBCH 77, approximately 6-8 weeks post-fruit set) showing minimal berry expansion but active internal development particularly seed maturation, representing the pause between Phase I vegetative growth and Phase II ripening growth, with 'active growth' referring primarily to active seed development and metabolic reorganization rather than berry size increase. Active growth mid berries are characterized by= (1) relatively constant berry size with diameter increasing <5% from end of Phase I (BBCH 75) to end of lag phase (BBCH 77), typically maintaining 6-12 mm diameter depending on cultivar (2) stable or slightly increasing fresh weight (0-10% increase) maintaining 0.6-1.8 g depending on cultivar, representing minimal net berry expansion (3) cells at final number (no cell division after BBCH 75-77) and relatively constant cell size (45-70 μm diameter) awaiting expansion during subsequent ripening phase (4) seeds undergoing rapid active development as the most prominent growth activity during lag phase, with embryos completing maturation from torpedo/bent-cotyledon stage to mature embryo, seed coats completing lignification and browning, and seeds achieving maximum size and 80-95% of final dry weight (5) green coloration maintained with chlorophyll still present though often declining during later lag phase in preparation for veraison (6) firm hard texture with high turgor and no softening (7) acidity at maximum or near-maximum levels (18-25 g/L titratable acidity) with malic acid synthesis concluded and high malic acid concentration (8) sugars relatively stable at 6-10% with minimal accumulation during lag phase. Active growth mid berries show active internal processes despite minimal size change= seeds developing rapidly with embryo differentiation, storage reserve accumulation (proteins, lipids in embryo and endosperm), and seed coat maturation representing the primary biosynthetic activity metabolic reprogramming occurring at transcriptional level with growth-related genes downregulated and ripening-related genes beginning upregulation in preparation for veraison hormone balance shifting with auxins and cytokinins declining, abscisic acid (ABA) beginning to increase, and ethylene remaining low, creating hormonal conditions promoting transition from growth to ripening. Active growth mid berries show changing resource allocation= photosynthate import continuing to support seed development and metabolic maintenance but not accumulating as sugars in berry flesh respiration rates moderate at 15-25 nmol CO₂ g⁻¹ s⁻¹ supporting maintenance metabolism and seed biosynthesis water import minimal creating relatively stable berry water content and turgor. Environmental factors during lag phase active growth mid berry stage affect subsequent development= water availability influences timing of veraison onset with moderate water deficit potentially advancing veraison by 3-10 days temperature affects lag phase duration with warmer conditions (22-28°C) shortening lag phase to 1-2 weeks while cooler conditions (15-20°C) extend to 2-3.5 weeks crop load and vine vigor affect resource availability for seed development potentially influencing seed size and maturation timing.
Active_Growth_Mid_Berry represents the lag phase (BBCH 77) where 'active growth' refers primarily to seed maturation rather than berry expansion, and is distinguished from Expanding_Mid_Berry (veraison, BBCH 81-83) by minimal berry growth versus beginning expansion and ripening. The active growth mid berry terminology is potentially misleading as berry expansion is minimal, but accurately reflects that active developmental processes (particularly seed maturation and metabolic reprogramming) are occurring despite static berry dimensions. Understanding lag phase biology is important for interpreting the berry developmental sequence and for managing practices affecting veraison timing and ripening progression.
Lag-Phase-Beere
Samenwachstums-Stadium
aktiv wachsende Mittel-Beere
lag-phase berry
seed-maturation berry
Coombe, B.G., McCarthy, M.G. (2000). Dynamics of grape berry growth. Australian Journal of Grape and Wine Research 6=131-135 Ollat, N. et al. (2002). Grape berry development= a review. Journal International des Sciences de la Vigne et du Vin 36=109-131.
Active Growth Mid Berry
In Pinot Gris vines in Alsace, active growth resumption commences in early April when meristematic cell division resumes following heat accumulation of approximately 130 GDD, with cell cycle progression detected through increased mitotic index within 48 hours of visible bud swelling at BBCH stage 05, respiratory rates increasing 5-fold from ecodormant levels within 72 hours, and rapid shoot elongation of 2-3 cm per day achieved within 10-14 days as growth systems reach full activation.
Active growth resumption in Vitis vinifera represents the re-initiation of developmental growth activity following the winter dormancy period, encompassing the activation of cell division in meristematic tissues, commencement of cell expansion driving visible bud swelling, and the metabolic upregulation supporting sustained biosynthetic production for growth. This resumption marks the functional end of dormancy from the perspective of growth activity, representing the point at which the vine transitions from the quiescent survival mode of winter to the active developmental mode of the growing season. The cellular basis of growth resumption involves the re-entry of meristematic cells into active cell cycle progression, with cells that were arrested in G1 phase during dormancy advancing through S phase DNA replication and subsequent division, producing the new cells that will constitute the expanding shoot. The metabolic dimension of growth resumption encompasses the dramatic upregulation of respiratory activity providing ATP for growth, biosynthetic pathway activation producing the amino acids, nucleotides, lipids, and carbohydrates required for cell production, and the mobilization of stored reserves from woody tissues to supply the demanding sink activity of growing buds. The hormonal changes accompanying growth resumption include increased cytokinin activity promoting cell division, elevated gibberellin signaling driving cell elongation, and establishment of auxin polar transport gradients that organize the spatial patterns of growth within the expanding shoot. The gene expression dimension involves the activation of growth-related transcriptional programs including cell cycle genes, cell wall biosynthesis genes, and developmental regulators that coordinate the organized growth of shoot structures. The rate of growth resumption accelerates following initiation as positive feedback loops between growth activity and resource mobilization become established, with initial slow growth transitioning to rapid expansion as metabolic and transport systems achieve full activation. The environmental sensitivity following growth resumption creates management implications, as the actively growing tissues are highly vulnerable to environmental stresses including frost, drought, and pest damage that can significantly impact shoot development and vine productivity.
Active_Growth_Resumption represents the functional reactivation aspect of Dormancy_Break describing the return to active developmental growth. Distinguished from Budbreak_Readiness which describes preparation, while Active_Growth_Resumption describes the actual growth activity. Encompasses cellular, metabolic, hormonal, and gene expression activation. Rate accelerates following initiation through positive feedback. Creates environmental vulnerability requiring management attention. Marks functional end of dormancy from growth perspective.
Entwicklungswiederaufnahme
Wachstumsreaktivierung
active growth commencement
aktive Wachstumswiederaufnahme
aktiver Wachstumsbeginn
developmental resumption
growth re-initiation
growth reactivation
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Zapata C et al (2004) Partitioning and mobilization of starch and N reserves in grapevine. Journal of Experimental Botany 55: 2165-2175 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235
Active Growth Resumption
In a Napa Valley Merlot vineyard, active malic enzyme activity was assayed over 45-day ripening period from mid-August through September Year N+1 (BBCH 83-87) showing NADP-malic enzyme maintained 42-58 units/g FW (average 48 units/g), NAD-ME 14-19 units/g (average 16 units/g), combined 56-74 units/g supporting malate decline from 9.2 to 3.1 g/L (average loss 0.14 g/L per day), VvME1 expression sustained 35-48 fold, under warm Napa conditions (daily high 28-32°C maintaining high enzyme activity and rapid malate catabolism).
Active_Malic_Enzyme_Activity represents the sustained high-level operation of NADP and NAD-dependent malic enzymes throughout berry ripening catalyzing oxidative decarboxylation of malate to pyruvate creating the primary enzymatic mechanism enabling progressive acid decline that defines ripening berry composition and wine quality characteristics. This active period extends from early veraison through late ripening (BBCH 83-87), typically 3-8 days post-enzyme induction through harvest, and is characterized by NADP-malic enzyme activity maintained at 25-70 units/g fresh weight (compared to pre-veraison 2-8 units/g) representing 5-12 fold sustained elevation, NAD-malic enzyme showing 8-25 units/g activity (from 1-5 units/g pre-veraison) contributing secondarily, combined malic enzyme capacity 35-90 units/g adequate for observed malate decline rates 0.18-0.50 g/L per day, and enzyme activity per se (units/g) remaining relatively stable throughout ripening though total berry activity potentially declining with berry expansion diluting enzyme concentrations. Enzymatic reaction mechanism involves: NADP-malic enzyme (EC 1.1.1.40) catalyzing L-malate + NADP⁺ → pyruvate + CO₂ + NADPH through oxidative decarboxylation proceeding via transient oxaloacetate intermediate rapidly decarboxylating to pyruvate, reaction thermodynamics strongly favorable with ΔG°' = -5.2 kcal/mol ensuring irreversibility under physiological conditions, pH optimum 7.0-7.5 matching cytosolic environment enabling efficient catalysis, and metal cofactor requirement with Mg²⁺ or Mn²⁺ (typically 2-5 mM) essential for catalytic activity. Enzyme kinetics during active period demonstrate: Km for L-malate 0.5-2.5 mM (isoform-dependent) well below cytoplasmic substrate concentrations 8-20 mM ensuring near-maximum velocity operation, Km for NADP⁺ 15-45 μM with cytoplasmic NADP⁺ pool 50-150 μM providing adequate cofactor, Vmax values 28-75 μmol malate oxidized per g fresh weight per hour in vitro corresponding to observed in vivo rates 1.5-4.5 μmol g⁻¹ h⁻¹ (0.15-0.45 g/L per day berry volume basis) accounting for cellular compartmentation and regulatory factors, and product inhibition by pyruvate at >5-10 mM and competitive inhibition by oxaloacetate at >0.5-2.0 mM potentially limiting activity though normal cellular concentrations typically below inhibitory thresholds. Enzyme structure and molecular properties inclu molecular mass 60-68 kDa for NADP-malic enzyme monomers with active enzyme existing as tetramers 240-270 kDa, amino acid sequence showing conserved active site residues across plant species with catalytic Lys and Asp residues essential for decarboxylation, NAD-malic enzyme showing similar structure 62-70 kDa monomers with tetrameric active form, and isoform diversity with multiple genes (VvME1, VvME2, potentially others) encoding cytosolic and mitochondrial forms providing functional redundancy and compartment-specific regulation. Gene expression supporting active enzyme levels shows: VvME1 transcript maintained at 25-70 fold above green berry baseline throughout ripening ensuring continued protein synthesis, VvME2 sustained 20-55 fold with similar kinetics, mRNA half-life 6-12 hours requiring continuous transcription to maintain steady-state levels, and translation efficiency apparently high with transcript levels correlating well with protein abundance and enzyme activity (r² = 0.78-0.92) indicating minimal post-transcriptional regulation. Protein turnover and stability during active period includes: enzyme protein half-life estimated 24-48 hours requiring continued synthesis to maintain elevated levels, proteolytic degradation through ubiquitin-proteasome system and autophagy balancing synthesis, post-translational modifications potentially including phosphorylation affecting activity or stability though modifications in grape malic enzyme incompletely characterized, and protein quality control ensuring properly folded active enzyme with misfolded proteins degraded preventing inactive enzyme accumulation. Subcellular distribution of active enzyme demonstrates: cytosolic malic enzyme 60-75% of total activity providing primary site for malate oxidation in cytoplasm pH 7.0-7.4, mitochondrial forms 25-40% of activity potentially supporting anaplerotic reactions or alternative metabolic functions, and compartment-specific regulation potentially adjusting distribution optimizing cellular metabolism though mechanisms incompletely defined. Metabolic integration of active malic enzyme includes: substrate supply from vacuolar malate through tonoplast transport maintaining cytoplasmic pool 8-20 mM adequate for enzyme saturation, cofactor availability with cytoplasmic NADP⁺/NADPH ratio ~3-8:1 (NADP⁺ 50-150 μM, NADPH 15-30 μM) favoring oxidative decarboxylation direction, product removal with pyruvate entering mitochondria for further oxidation preventing accumulation that could inhibit enzyme, and CO₂ efflux through diffusion eliminating product maintaining reaction favorability. NADPH utilization from malic enzyme activity includes: reductive biosynthesis consuming ~40-65% of NADPH for fatty acid, flavonoid, and terpenoid synthesis all active during ripening, antioxidant systems utilizing 15-30% for glutathione reductase and other NADPH-dependent antioxidant enzymes, and excess NADPH potentially oxidized by mitochondrial external NADH dehydrogenase for ATP production though inefficient compared to direct NADH oxidation. Pyruvate fate from malic enzyme reaction shows: mitochondrial import through pyruvate carrier for oxidation via pyruvate dehydrogenase producing acetyl-CoA feeding citric acid cycle representing primary fate ~60-80%, cytoplasmic metabolism through pyruvate kinase potentially regenerating phosphoenolpyruvate for biosynthesis ~10-20%, transamination to alanine providing amino acid ~5-12%, and minor fermentative conversion to lactate or ethanol under hypoxic conditions in berry interior 35-38°C beginning enzyme denaturation reducing activity 20-50% potentially irreversibly inactivating enzyme, and cool temperatures 15-22°C reducing activity 40-70% compared to optimal providing mechanism for acid preservation in cool climates or shaded canopy positions. pH effects on activity show: sharp pH optimum 7.0-7.5 with activity declining 30-60% at pH 6.5 and 50-80% at pH 8.0, cytoplasmic pH 7.0-7.4 throughout ripening maintaining near-optimal conditions, and vacuolar pH 2.7-3.2 precluding activity in that compartment ensuring malate metabolism restricted to cytoplasm after tonoplast transport. Cultivar variation in active enzyme characteristics includes: low-acid varieties maintaining 45-85 units/g combined malic enzyme throughout ripening enabling extensive acid depletion, moderate cultivars showing 30-60 units/g supporting intermediate decline, and high-acid cultivars displaying 20-45 units/g contributing to acid retention, with activity levels strongly correlated with final berry malate r² = 0.80-0.93 demonstrating causal relationship. Environmental modulation of activity includes: temperature directly affecting catalytic rate as described above, potential acclimation effects where sustained warm temperatures may induce additional enzyme synthesis enhancing capacity, and stress responses where severe heat or water deficit may actually reduce activity despite elevated temperature through protein denaturation or metabolic dysfunction. Inhibitor studies reveal: oxaloacetate competitive inhibition Ki ~0.8-2.5 mM, ATP allosteric inhibition at >3-5 mM in some isoforms potentially providing energy status regulation, and citrate inhibition at >2-5 mM potentially coordinating with citric acid cycle flux though physiological relevance uncertain given normal cellular metabolite concentrations. Viticultural implications inclu enzyme activity levels predicting malate decline trajectory enabling forecasting final acids, canopy management affecting berry temperature modulating enzyme activity and thus malate loss rates, and spatial variation in activity across vineyard zones potentially contributing to maturity heterogeneity. Research applications involve: enzyme activity assays measuring malate oxidation spectrophotometrically through NADPH production at 340 nm providing quantitative assessment, gene expression analysis correlating transcript levels with activity confirming transcriptional control, and inhibitor or temperature manipulation studies revealing regulatory mechanisms and optimization strategies for acid management.
Active_Malic_Enzyme_Activity is distinguished from malic enzyme induction by representing sustained activity (25-70 units/g NADP-ME) rather than initial upregulation, from respiratory consumption by focusing on specific enzyme rather than overall process, and from acid decline by representing enzymatic mechanism rather than net compositional change. Duration entire ripening BBCH 83-87. Temperature optimum 25-32°C, Q₁₀ 2.2-3.5. pH optimum 7.0-7.5. Km malate 0.5-2.5 mM. Active_Malic_Enzyme_Activity represents critical sustained enzymatic function enabling malate decline fundamentally affecting wine acidity and quality.
Malat-Decarboxylase-Funktion
NADP-ME activity maintenance
NADP-ME-Aktivitäts-Aufrechterhaltung
Pyruvat-produzierende Enzym-Operation
anhaltende Malat-Enzym-Katalyse
malate decarboxylase function
pyruvate-producing enzyme operation
sustained malic enzyme catalysis
Ruffner HP (1982) Metabolism of tartaric and malic acids in Vitis:a review Vitis 21:247-259 malic enzyme comprehensive review Sweetman C, Deluc LG, Cramer GR, Ford CM, Soole KL (2009) Regulation of malate metabolism in grape berry and other developing fruits Phytochem 70:1329-1344 enzyme activity regulation Hawker JS (1969) Changes in the activities of malic enzyme, malate dehydrogenase, phosphopyruvate carboxylase and pyruvate dehydrogenase during the development of a non-climacteric fruit:the grape Phytochem 8:9-23 classic enzyme activity study.
Active Malic Enzyme Activity
Merlot in Bordeaux shows active pollen shedding during mid-June Year N+1 at BBCH 63-65 over 3-4 days June 9-12, with abundant pollen release from widespread anther dehiscence creating visible pollen clouds during favorable morning hours 9am-11am, atmospheric concentration estimated 300-800 grains per cubic centimeter near clusters, under oceanic conditions of 22-27°C temperature, 54-68% relative humidity, light winds 6-12 km per hour, supporting efficient pollination enabling 54-64% fruit set typical of this mid-season cultivar
Active pollen shedding represents the sustained pollen liberation and dispersal phase following initial exposure when continued gravitational, mechanical, and wind-assisted processes release pollen from dehisced anthers distributing grains through atmosphere to receptive stigmas, creating maximum male gamete availability and pollination opportunity. This critical dispersal phase occurs during mid-peak flowering in mid-June in temperate regions Year N+1, typically spanning 3-5 days as widespread anther dehiscence creates intensive pollen release throughout cluster, corresponding to BBCH 63-65 peak anthesis stages. Shedding mechanisms involve multiple processes including continued gravitational pollen fall from opened anthers depositing grains over extended period, mechanical agitation from flower movement and environmental disturbances dislodging loosened pollen, and wind-induced dispersal carrying airborne pollen to receptive stigmas within and between flowers. Gravitational shedding shows persistent slow release as loosened pollen progressively falls from anther locules, positioning effects with laterally or downward-oriented anthers facilitating efficient gravitational release, and cumulative substantial pollen liberation over hours to days following initial explosive discharge. Mechanical shedding involves physical disturbances including wind-induced anther movement agitating structures dislodging pollen, insect visitation occasionally causing vibrations though insects play minor role in grapevine pollination, and environmental movements from temperature changes or growth processes creating subtle agitation. Wind-assisted dispersal shows primary pollen transport mechanism with light grains 25-35 micrometers diameter easily becoming airborne, air currents lifting pollen from anthers and transporting through atmosphere, and characteristic pollen clouds forming around flowering clusters during favorable conditions creating visible evidence of active shedding. Atmospheric pollen concentration during active shedding reaches maximum levels with estimated 100-1000 grains per cubic centimeter in immediate cluster vicinity under favorable conditions, declining concentration with distance but remaining elevated 10-100 grains per cubic centimeter at vine scale, and creating abundant male gamete availability supporting efficient pollination. Pollen cloud formation shows visible phenomenon during peak shedding under optimal dry conditions, appearance of pale yellow haze surrounding flowering clusters particularly evident during morning peak activity, and disappearance when humidity rises or wind calms indicating environmental dependence. Pollen dispersal patterns show primary deposition within same flower facilitating self-pollination accounting for 60-80% successful pollinations, neighboring flowers on same cluster receiving substantial pollen through short-distance dispersal, and limited long-distance transport between vines contributing minor proportion 5-15% to total pollinations. Pollen viability during active shedding shows maximum in freshly released pollen 75-90% germination capacity declining over time, atmospheric pollen showing progressive viability decline with aged airborne pollen showing 40-60% viability after several hours, and importance of continuous fresh pollen production maintaining viable atmospheric pool during shedding period. Environmental conditions optimizing shedding include temperature 24-30°C promoting anther dehydration facilitating continued release, moderate humidity 50-65% maintaining pollen viability without promoting excessive aggregation or preventing liberation, light wind 5-15 km per hour enhancing dispersal without causing excessive pollen loss or desiccation, and overall stable favorable weather enabling sustained efficient shedding. Temperature effects on shedding show warm conditions facilitating mechanical liberation through reduced tissue moisture and appropriate pollen surface properties, cool conditions potentially reducing shedding efficiency through maintained tissue hydration, and optimal thermal range supporting both liberation and viability maintenance. Humidity effects show critical importance with moderate levels 50-65% optimal for maintaining liberating pollen in dispersible state without aggregation while preserving viability, high humidity above 70% causing pollen aggregation reducing dispersal and potentially preventing continued liberation, and low humidity below 45% potentially causing excessive desiccation reducing viability though facilitating mechanical release. Wind effects show beneficial role for dispersal with light-moderate wind 5-15 km per hour optimal for pollen transport enhancing pollination efficiency, calm conditions reducing atmospheric transport though gravitational and short-range dispersal remaining effective for self-pollination, and excessive wind above 20 km per hour potentially dispersing pollen too far or causing rapid desiccation. Diurnal shedding pattern shows characteristic morning maximum as temperature rises and humidity declines creating optimal conditions typically 9am-12pm, reduced afternoon activity as conditions potentially become less favorable with rising temperature or changing humidity, and daily rhythm creating repeated peaks throughout active shedding period. Peak shedding intensity during mass flowering shows maximum pollen release when numerous anthers throughout cluster simultaneously shed pollen, coordinated timing creating concentrated pollination opportunity, and favorable synchronization supporting efficient pollination and high fruit set potential. Pollen production during active shedding continues from progressively dehiscing anthers maintaining fresh pollen availability, sequential opening pattern providing extended release period rather than single exhaustive discharge, and overall sustained male gamete supply supporting continued pollination opportunities. Pollination success during active shedding shows maximum efficiency with abundant fresh viable pollen, widespread stigma receptivity providing numerous capture opportunities, and optimal environmental conditions enabling effective pollen-stigma encounters, supporting fruit set 50-70% in good-setting cultivars under favorable conditions. Rain effects during active shedding show catastrophic interference with water washing pollen from anthers and atmosphere eliminating atmospheric pool, preventing continued shedding through elevated humidity maintaining tissue hydration, and destroying pollen viability through excessive hydration, potentially reducing fruit set to 15-30% if rain occurs during peak shedding. The process shows two distinguishable aspects progressing from peak pollen discharge establishing maximum release intensity, through maximum pollen dispersal creating optimal atmospheric availability. Duration typically spans 3-5 days representing peak shedding period, with warm stable conditions promoting compressed intense shedding while variable conditions may extend period. Cultivar variation in shedding characteristics includes pollen production quantity affecting atmospheric concentration potential, shedding efficiency from anthers affecting liberation completeness, and timing coordination with stigma receptivity affecting pollination success. Regional timing shows active pollen shedding during mid-June in warm climates, mid-June in temperate regions, and mid-late June in cool areas, coinciding with peak flowering and maximum reproductive activity. Viticultural significance includes representing critical male reproductive phase determining pollination success and fruit set, requiring optimal environmental conditions for efficient shedding and viability maintenance, demonstrating extreme weather vulnerability particularly to rain during peak shedding, and establishing pollen availability as dominant factor determining fruit set outcomes. Research applications include analyzing shedding mechanisms and pollen liberation dynamics, quantifying atmospheric pollen concentrations and dispersal patterns, evaluating environmental effects on shedding efficiency and pollen viability, assessing pollination efficiency under varying shedding conditions, and developing models predicting fruit set from pollen shedding characteristics and environmental conditions.
Active_Pollen_Shedding is second major Pollen_Release phase distinguished by sustained dispersal creating maximum male gamete availability. Follows Anther_Dehiscence. Corresponds to BBCH 63-65 peak anthesis. Subdivided into Peak_Pollen_Discharge and Maximum_Pollen_Dispersal. Represents critical 3-5 day period determining majority of fruit set through pollination success. Environmental conditions particularly temperature, humidity, and wind critically affect efficiency. Extreme weather vulnerability to rain catastrophically affecting pollen function. Most important male reproductive phase.
Pollendispersionsphase
Spitzen-Pollenschüttung
active pollen distribution
aktive Pollenabgabe
nachhaltige Pollenfreisetzung
peak pollen shedding
pollen dispersal phase
sustained pollen release
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 380-385 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 290-295
Active Pollen Shedding
Syrah grafted on SO4 rootstock sampled mid-May showing active primary root tip on tap root, root tip 9.8 mm length showing vigorous elongation 6.4 mm per day measured over 48-hour period, microscopic examination revealing abundant mitotic figures in apical meristem with estimated 15% cells in M phase indicating active division, elongation zone cells showing vacuolation and expansion, root tip white-cream color indicating active undifferentiated state, turgid appearance, copious mucilage secretion, active primary root tip demonstrating functional gravitropic response, metabolic activity evident through high respiration rate
Active primary root tip is primary root tip in state of active growth and meristematic activity, emphasizing current functional status rather than structure, characterized by ongoing cell division in apical meristem with cells progressing through mitotic cycle, continuous cell elongation in elongation zone increasing root length, active root cap production replacing sloughed cells, vigorous mucilage secretion lubricating soil penetration, and measurable daily elongation typically 3-12 mm per day during favorable conditions distinguishing active from dormant or slowly growing state, with active primary root tip showing white to cream coloration indicating young undifferentiated tissue, turgid appearance reflecting high water content and metabolic activity, readily observable elongation over 24-48 hour period, and functional capacity for gravitropic response, nutrient sensing, and obstacle avoidance demonstrating that active primary root tips represent growth-functional state essential for primary root system expansion occurring primarily during spring and fall growing periods. Active primary root tip metabolic characteristics includes= exhibiting high respiration rate (active tips consuming oxygen and producing CO2 supporting division and elongation metabolism), showing elevated ATP production (active mitochondria generating energy for biosynthesis, transport, and growth), demonstrating active protein synthesis (ribosomes producing enzymes, structural proteins, and regulatory factors required for growth), revealing high nutrient demand (active tips importing sugars, amino acids, minerals supporting intense metabolic activity), and displaying water uptake (active elongation requiring water influx for vacuole expansion and turgor-driven cell expansion). Active primary root tip cell division activity includes= showing frequent mitosis (meristem cells dividing every 10-18 hours during peak activity continuously producing new cells), exhibiting visible mitotic figures (microscopic examination revealing chromosomes and spindles indicating cells in M phase), demonstrating DNA synthesis (meristem cells in S phase replicating DNA in preparation for division), revealing cell cycle progression (populations of cells in G1, S, G2, and M phases indicating active cycling), and producing derivative cells (meristem generating cells that enter elongation zone then differentiate into root tissues). Active primary root tip elongation characteristics includes= demonstrating measurable growth (active tips showing 3-12 mm daily length increase during growing season depending on conditions), exhibiting cell expansion (elongation zone cells increasing length 10-50 fold through water uptake and wall loosening), showing turgor-driven extension (cell expansion driven by osmotic water influx generating turgor pressure stretching walls), revealing hormone regulation (auxin, gibberellins, and brassinosteroids promoting elongation while abscisic acid and ethylene potentially inhibiting), and displaying environmental responsiveness (elongation rate varying with temperature, water availability, nutrients, and soil strength). Active primary root tip temporal patterns includes= showing diurnal variation (root tip elongation potentially showing daily rhythms with peaks during night or early morning), exhibiting seasonal activity (active state concentrated in spring April-June and fall September-October growing periods), demonstrating temperature dependence (active growth requiring soil temperature >10°C with optimum 20-25°C declining >30°C), revealing moisture responsiveness (active state dependent on adequate soil moisture ceasing under severe drought), and displaying carbohydrate supply dependence (active tip growth requiring continuous photosynthate supply potentially competing with shoot growth). Active primary root tip versus dormant includes= contrasting with dormant tips (active tips showing ongoing division and elongation versus dormant showing quiescent meristem and no elongation), demonstrating seasonal transitions (active tips entering dormancy in fall as temperature drops and photoperiod shortens resuming activity spring), exhibiting stress responses (active tips potentially entering quiescence under severe drought, flooding, or extreme temperatures), revealing protective strategies (dormancy protecting root tips from winter freeze damage or summer drought stress), and showing reversibility (dormant primary root tips capable of resuming active state when conditions improve). Active primary root tip environmental sensing includes= detecting gravity continuously (active tips maintaining gravitropic response adjusting growth direction if displaced), sensing moisture gradients (active tips potentially showing hydrotropism growing toward water), responding to nutrients (active tips proliferating in nutrient-rich zones and avoiding deficient or toxic areas), detecting obstacles (active tips sensing mechanical impedance potentially growing around barriers through thigmotropism), and revealing signal transduction (environmental stimuli triggering biochemical cascades affecting growth rate and direction). Active primary root tip mucilage production includes= secreting copious mucilage (active root cap border cells continuously producing and releasing polysaccharide gel), showing mucilage composition (complex mixture of polysaccharides, proteins, and cells creating viscous gel), demonstrating lubrication function (mucilage reducing friction between root tip and soil particles facilitating penetration), revealing rhizosphere modification (mucilage binding soil particles creating structured zone around root influencing water and nutrient dynamics), and displaying defensive roles (mucilage potentially protecting root tip from desiccation, pathogens, and toxins). Active primary root tip management considerations includes= indicating vine vigor (active primary root tip abundance and growth rate reflecting overall vine vigor and health), showing stress sensitivity (reduced active tip function signaling water, nutrient, or other stress requiring management response), demonstrating transplant vulnerability (active primary root tips easily damaged during transplanting requiring careful handling), revealing cultivation effects (tillage or traffic potentially damaging active tips reducing root growth and vine performance), and requiring protection (management practices should minimize active primary root tip damage maintaining healthy root system growth).
Active_Primary_Root_Tip emphasizes current active growth state and is distinguished from dormant or slowly growing primary root tips by focus on ongoing meristematic activity. Critical for understanding growth state, seasonal activity, metabolic intensity, elongation dynamics, environmental responsiveness, and the active functional status of primary root tip enabling current root system expansion during favorable growth periods.
Actively growing primary tip
Aktive Primäre Wurzelspitze
Elongating primary tip
Functional primary root apex
Funktionelle primäre Wurzelapex
Wachsende Hauptwurzel-Spitze
Malamy, J. E. (2005). Intrinsic and environmental response pathways that regulate root system architecture. Plant, Cell & Environment, 28(1), 67-77.
Active Primary Root Tip
Pinot Noir grafted on SO4 rootstock sampled mid-May showing active root apical meristem on third-order lateral root, meristem 0.54 mm diameter comprising estimated 1180 cells, microscopic examination revealing mitotic index 18% indicating active division, numerous metaphase and anaphase figures observed, BrdU pulse-labeling showing 32% S-phase cells documenting active DNA synthesis, meristem producing estimated 64 new cells per day, root elongation measured 3.6 mm per day reflecting active meristem function, meristem showing elevated respiration rate 2.8× higher than differentiated root tissues, active root apical meristem enabling vigorous root growth during favorable spring conditions
Active root apical meristem is root apical meristem in state of active cell division and tissue production, emphasizing current functional activity rather than structural presence, characterized by ongoing mitosis with substantial proportion of cells progressing through cell cycle, observable mitotic figures in microscopic examination indicating cells in M phase, DNA synthesis in S phase cells documenting genome replication, continuous production of derivative cells exiting meristem into elongation zone, and demonstrable root elongation resulting from meristem activity typically 1-15 mm per day depending on root order and conditions, with active root apical meristem showing high metabolic activity with elevated respiration supporting biosynthesis and division, substantial resource demand requiring continuous photosynthate supply for growth, responsive capacity adjusting division rate and meristem size to environmental conditions and hormonal signals, and representing growth-functional state that enables root system expansion occurring primarily during spring and fall growing seasons when conditions favorable contrasting with quiescent dormant state during winter or severe stress. Active root apical meristem cell cycle activity includes= showing high mitotic index (active meristem typically 10-25% cells in M phase during peak growth compared to 10°C with optimum 20-25°C for maximum activity), revealing moisture responsiveness (active state dependent on adequate soil moisture entering quiescence under severe drought), and displaying photoperiod independence (root apical meristem activity not directly responding to day length though affected by photosynthate supply influenced by photoperiod). Active root apical meristem size dynamics includes= showing enlarged meristem (active state often associated with increased meristem size from 500-800 to 1000-2000+ cells depending on root), exhibiting cell number increase (active meristem potentially doubling cell number during peak growth periods), demonstrating rapid cell production (high division rate creating larger population of meristematic cells), revealing growth correlation (larger active meristems generally producing faster root elongation), and displaying reversible expansion (meristem size declining when returning to dormant or slow-growth state). Active root apical meristem hormonal state includes= showing elevated auxin levels (indole-3-acetic acid concentrations high in active meristem promoting division), exhibiting cytokinin presence (cytokinins supporting meristem maintenance and division during active state), demonstrating gibberellin effects (gibberellins promoting active meristem function and cell cycle progression), revealing brassinosteroid activity (brassinosteroids enhancing division and elongation in active state), and displaying hormone signaling (active transduction of hormonal signals modulating meristem activity). Active root apical meristem versus dormant includes= contrasting with dormant meristem (active showing high division rate versus dormant showing minimal or no division), demonstrating seasonal transitions (active meristem entering dormancy fall-winter as temperature drops resuming spring warming), exhibiting reversibility (dormant meristem capable of returning to active state when conditions improve), revealing protective function (dormancy protecting meristem from winter freeze or summer drought damage), and showing metabolic differences (active meristem high metabolism versus dormant reduced metabolism conserving resources). Active root apical meristem in different root types includes= showing highest activity in primary roots (primary root apical meristems most active producing rapid elongation establishing main framework), exhibiting moderate activity in secondary roots (secondary root meristems active but less vigorous than primary producing moderate elongation), demonstrating variable activity in tertiary roots (fine root meristems showing high activity in favorable conditions but quickly entering dormancy under stress), revealing order-dependent patterns (lower-order meristems generally more active and persistent than higher-order meristems), and displaying stress sensitivity gradient (higher-order fine root meristems more sensitive to stress ceasing activity before lower-order structural root meristems). Active root apical meristem management significance includes= indicating root growth status (active meristem presence and abundance reflecting current root system expansion), showing stress response (reduced active meristem function signaling environmental stress requiring management intervention), demonstrating nutrient responsiveness (active meristem activity potentially enhanced by fertilization supporting root development), revealing cultivation sensitivity (tillage or compaction potentially damaging active meristems reducing root growth), and requiring protection (management should minimize disturbance to active meristems maintaining healthy root system development).
Active_Root_Apical_Meristem emphasizes current active division state and is distinguished from quiescent or dormant meristem by focus on ongoing cell cycle activity. Critical for understanding meristem activity state, mitotic intensity, metabolic demand, seasonal patterns, resource requirements, and the functionally active meristematic state producing new cells enabling current root elongation during favorable growth periods.
Actively dividing root meristem
Aktives Wurzel-Apikalmeristem
Functional root apical meristem
Funktionelles Wurzelmeristem
Growing root meristem
Teilendes Wurzelmeristem
Active Root Apical Meristem
Merlot grafted on 3309 Couderc rootstock showing active root hairs on third-order lateral root sampled 15 mm from root tip, microscopic examination revealing active hairs with intact dense cytoplasm, abundant mitochondria, functional plasma membranes, turgid appearance indicating adequate hydration, isotope tracing (15N and 32P) demonstrating active hairs providing 76% of root nitrogen uptake and 82% of phosphorus uptake despite comprising only 18% of root surface area, active hairs 3-6 days old in peak functional period, membrane potential measurements indicating active H+-ATPase function
Active root hair is root hair in functional state actively absorbing water and nutrients from soil, characterized by intact living cytoplasm with functional organelles and membranes, turgid appearance reflecting adequate hydration and osmotic function, functional membrane transporters enabling active ion uptake, measurable absorption capacity demonstrated through isotope tracing or other methods, and typical age 1-7 days representing period from completion of growth through peak function before senescence, with active root hair showing dense cytoplasm particularly in tip region, abundant mitochondria providing ATP for active transport, functional plasma membrane with ion channels and H+-ATPases, close contact with soil particles maximizing absorption interface, and mucilage sheath surrounding hair potentially facilitating water and nutrient movement, and demonstrating that active root hairs despite representing only brief functional period provide majority of root absorption surface at any given time making active hair population critical for current vine water and nutrient status. Active root hair cytoplasmic characteristics includes= showing intact protoplast (active root hair containing living cytoplasm with all cellular components functional), exhibiting organelle abundance (mitochondria, ER, Golgi, ribosomes present and active supporting metabolism and transport), demonstrating cytoplasmic streaming (active cytoplasmic circulation distributing materials and maintaining hair function), revealing tip concentration (organelles particularly concentrated in growing or recently grown hair tip), and displaying membrane integrity (plasma membrane and organellar membranes intact maintaining compartmentation and function). Active root hair membrane function includes= showing active H+-ATPases (proton pumps creating electrochemical gradient driving ion uptake), exhibiting ion transporters (specific channels and carriers for K+, NO3-, H2PO4-, Ca2+, and other nutrients), demonstrating water uptake capacity (aquaporins and osmotic gradient facilitating water absorption), revealing selective permeability (membrane transporters enabling preferential uptake of essential nutrients over non-essential or toxic ions), and displaying responsive regulation (transporter activity adjusting to nutrient availability and plant demand). Active root hair absorption performance includes= providing high uptake rates (active root hairs absorbing water and nutrients at 10-100 times rate per surface area compared to root cylinder), exhibiting disproportionate contribution (active hairs though small surface fraction providing 60-80% of root uptake), demonstrating rapid response (active hair uptake quickly adjusting to changing soil conditions or plant demand), revealing nutrient specificity (active hairs showing particularly high uptake for mobile nutrients like nitrogen and potassium), and displaying daily variation (active hair function potentially showing diurnal patterns correlating with transpiration and metabolism). Active root hair soil contact includes= showing intimate association (active hair surface in close contact with soil particles maximizing exchange interface), exhibiting mucilage production (active hairs secreting polysaccharide gel creating favorable micro-environment), demonstrating rhizosphere modification (active hairs affecting local pH, moisture, and chemistry through secretions and uptake), revealing exploration capacity (active hairs extending into soil pores accessing resources in micro-sites), and displaying mechanical interaction (active hairs potentially deflecting or growing around obstacles optimizing soil contact). Active root hair turgor maintenance includes= showing high turgor pressure (active hairs maintaining 0.3-0.6 MPa turgor reflecting osmotic function), exhibiting osmotic adjustment (active solute accumulation maintaining osmotic potential as absorption continues), demonstrating water status indicator (active hair turgor reflecting overall plant water status and soil moisture), revealing drought sensitivity (active hair turgor declining under water deficit potentially compromising function), and displaying recovery capacity (active hairs regaining turgor upon rehydration if not severely damaged). Active root hair metabolic activity includes= showing elevated respiration (active hairs consuming oxygen and producing CO2 supporting active transport and maintenance), exhibiting ATP production (abundant mitochondria generating energy for ion pumping and biosynthesis), demonstrating protein synthesis (ribosomes producing membrane proteins, enzymes, and other components), revealing biosynthetic activity (synthesizing membrane lipids, wall materials, and other cellular components), and displaying energy demand (active hairs requiring continuous photosynthate supply supporting intense metabolism). Active root hair environmental responsiveness includes= responding to nutrient availability (active hair transporters upregulating in low-nutrient conditions increasing acquisition efficiency), affected by soil moisture (active hair function requiring adequate moisture declining under drought), influenced by temperature (active hair metabolism and uptake increasing with temperature within optimal range), showing pH sensitivity (active hair function affected by soil pH influencing nutrient availability and transport), and demonstrating toxic ion effects (active hair function potentially impaired by aluminum, sodium, or other toxic ions in soil). Active root hair duration includes= showing functional period (active state typically lasting 2-7 days from growth completion to senescence onset), exhibiting peak activity (maximum absorption often 2-4 days after hair reaches full length), demonstrating gradual decline (active function declining as hair ages approaching senescence), revealing age-related changes (older active hairs showing reduced turgor, membrane function, and absorption capacity), and displaying transition to senescence (active state ending as cytoplasmic degradation and membrane dysfunction initiate senescence process).
Active_Root_Hair emphasizes functional absorbing state and is distinguished from senescent non-functional hairs by focus on living metabolism and uptake capacity. Includes Absorbing_Active_Root_Hair and Functional_Active_Root_Hair. Critical for understanding absorption performance, membrane function, metabolic activity, turgor maintenance, environmental responsiveness, and the living functional state providing majority of root water and nutrient uptake.
Absorbing hair
Aktives Wurzelhaar
Functional root hair
Funktionelles Haar
Lebendes Absorptionshaar
Living root hair
Active Root Hair
Chardonnay grafted on SO4 rootstock at 14 years post-grafting showing active rootstock sapwood comprising outermost 5 annual rings (rings 10-14 from pith) with 2.4 cm radial thickness, active sapwood zone showing nearly 100% vessel patency with open unoccluded lumens, hydraulic conductivity measurements indicating active sapwood providing 72% of total trunk water transport capacity despite comprising only 52% of total sapwood area, living ray parenchyma in active sapwood showing highest starch concentrations (iodine staining revealing darkest staining in active zone), active sapwood conducting estimated 45-65 liters daily during peak transpiration supporting mature canopy, active sapwood vessels 95-125 micrometers diameter optimal for SO4, wood tissue lightest tan color in active zone indicating most recent formation
Active rootstock sapwood is rootstock sapwood in outer sapwood region typically comprising most recent 3-8 annual rings showing maximum functional capacity, characterized by highest vessel patency with minimal embolism or occlusion, most vigorous living ray parenchyma with optimal carbohydrate storage and metabolic activity, greatest hydraulic conductivity providing majority of trunk water transport, and representing prime functional zone where water conduction, reserve storage, and defensive responses operate at peak efficiency supporting current season growth and productivity. Active rootstock sapwood position and extent includes= located in outermost sapwood zone (active sapwood forming peripheral ring immediately inside vascular cambium representing most recently formed functional xylem), typically comprising recent 3-8 annual rings (most recent years of wood production remaining in peak functional condition varying with rootstock vigor and environmental conditions), showing radial thickness 1.5-3.5 cm (active sapwood zone depth depending on annual ring width and number of peak-function rings maintained), forming complete peripheral ring (active sapwood surrounding trunk circumference creating optimal conducting pathway), and transitioning to less active inner sapwood (gradual decline in function moving inward toward older sapwood approaching heartwood conversion). Active rootstock sapwood maximum vessel function includes= showing highest vessel patency (nearly all vessels in active sapwood zone open and functional with minimal embolism, tyloses, or gum occlusion), exhibiting optimal hydraulic conductivity (active sapwood providing maximum water transport efficiency per unit area with least resistance to flow), conducting majority of trunk water transport (active sapwood zone potentially providing 60-80% of total trunk hydraulic conductivity despite comprising only 40-60% of sapwood area), demonstrating efficient vessel arrangement (recently formed vessels showing optimal size distribution and connectivity for transport), and maintaining peak transport capacity (active sapwood vessels functioning at maximum efficiency supporting full canopy transpiration demands). Active rootstock sapwood living tissue vitality includes= containing most vigorous ray parenchyma (living cells in active sapwood showing highest metabolic activity, respiration rates, and synthetic capacity), showing optimal carbohydrate storage (ray cells accumulating and mobilizing starch reserves most effectively in active sapwood zone), exhibiting active defensive capacity (living cells most capable of rapid response to wounding or infection synthesizing and secreting antimicrobial compounds), maintaining efficient lateral transport (ray cells facilitating radial movement of water, nutrients, and signaling molecules between vessels and cambium), and demonstrating peak physiological function (all living tissue activities including respiration, synthesis, and response operating at maximum efficiency in recently formed wood). Active rootstock sapwood seasonal dynamics includes= showing maximum spring activity (active sapwood supporting peak water transport during bud break and early shoot growth), conducting highest summer flow rates (active sapwood enabling 40-70+ liters daily transport during peak transpiration periods), facilitating fall reserve accumulation (living cells in active sapwood actively storing photosynthates as starch during late season), maintaining winter protection (active sapwood cells accumulating cryoprotective compounds reducing freeze damage risk), and responding rapidly to seasonal transitions (active sapwood tissue most responsive to environmental cues triggering seasonal physiological changes). Active rootstock sapwood carbohydrate dynamics includes= storing highest starch concentrations (ray parenchyma in active sapwood accumulating more carbohydrates per unit volume than older inner sapwood), showing most dynamic reserve cycling (active sapwood reserves mobilized rapidly in spring supporting growth and replenished quickly in fall), providing readily available energy (carbohydrates in active sapwood immediately accessible for cambial activity, defense responses, or stress recovery), maintaining optimal storage conditions (young healthy cells in active sapwood most efficient at starch synthesis, storage, and remobilization), and supporting vine resilience (active sapwood reserves critical for responding to stress, damage, or resource limitation). Active rootstock sapwood vulnerability and protection includes= showing vulnerability to freeze damage (most recently formed wood potentially more susceptible to freeze injury than older acclimated tissue), exhibiting sensitivity to cavitation (young vessels potentially more prone to drought-induced embolism than older more conditioned vessels in some conditions), demonstrating rapid response to damage (living cells in active sapwood most capable of quick compartmentalization and defense compound synthesis), providing first line of defense (active sapwood tissue first to respond to pathogen attack or injury limiting damage spread), and maintaining protective capacity (active sapwood cells synthesizing antimicrobial compounds and forming barriers protecting inner tissue). Active rootstock sapwood in water stress response includes= showing sensitivity to drought (active sapwood vessels potentially vulnerable to cavitation during severe water stress), exhibiting embolism repair capacity (refilling of embolized vessels most efficient in active sapwood zone during stress recovery), demonstrating adaptive responses (living cells in active sapwood adjusting metabolism and vessel function in response to water limitation), maintaining critical transport (active sapwood sustaining minimal water delivery even under stress conditions supporting vine survival), and recovering transport function (active sapwood vessels first to restore full conductivity when water availability improves). Active rootstock sapwood contribution to trunk function includes= providing majority of water transport (active sapwood conducting 60-80% of total trunk water flow despite being subset of total sapwood), supporting peak transpiration demands (active sapwood capacity adequate for maximum water loss during hot periods maintaining canopy function), storing most accessible reserves (active sapwood carbohydrates readily mobilized supporting rapid growth or stress response), maintaining optimal defensive capacity (active sapwood cells most capable of responding to pathogen attack or injury), and enabling current season productivity (active sapwood function essential for supporting annual shoot growth, flowering, and fruit development). Active rootstock sapwood in different growth conditions includes= showing broader zone under favorable conditions (adequate water and nutrients supporting maintenance of more annual rings in peak functional state), exhibiting narrower zone under stress (water limitation, disease, or poor nutrition potentially reducing number of peak-function rings maintained), displaying vigor-dependent extent (vigorous rootstocks potentially maintaining broader active sapwood zone than low-vigor rootstocks), demonstrating age-related changes (young rapidly growing trunk showing higher proportion of wood in active state than old slow-growing trunk), and varying with management (irrigation, nutrition, and canopy management affecting active sapwood development and maintenance). Active rootstock sapwood measurement and assessment includes= determining extent through hydraulic conductivity measurement (quantifying water transport capacity of successive sapwood layers identifying most conductive zone), estimating from ring counts (assuming most recent 3-8 rings represent active sapwood depending on trunk age and vigor), assessing through starch staining (iodine staining revealing highest carbohydrate concentrations in active sapwood zone), evaluating through dye injection (tracking dye movement showing primary water transport pathways in most active vessels), and inferring from wood appearance (freshest lightest colored sapwood near cambium typically corresponding to most active zone). Active rootstock sapwood in young versus mature trunk includes= young trunk (years 1-8) showing all or most sapwood as active (high proportion of wood in peak functional state in rapidly growing young trunk), establishing trunk (years 8-15) differentiating active zone (outer sapwood maintaining peak function while inner sapwood beginning gradual decline toward heartwood conversion), mature trunk (years 15-40) displaying distinct active sapwood ring (clear differentiation between highly functional outer sapwood and less active inner sapwood), old trunk (years 40+) potentially showing narrower active zone (limited number of narrow recent rings comprising active sapwood in slow-growing aged trunk), and very old trunk possibly showing minimal active sapwood (only outermost 1-2 very narrow rings remaining in peak functional state in ancient trunk). Active rootstock sapwood in rootstock cultivars includes= vigorous rootstocks (1103P, 110R) potentially maintaining broader active sapwood zones 2.5-4 cm due to rapid growth and large vessels, moderate rootstocks (SO4, 3309C, Teleki 5C) showing typical active sapwood zones 1.5-3 cm with medium vessels, low-vigor rootstocks (Riparia, 101-14) exhibiting narrower active zones 1.5-2.5 cm with smaller vessels, cultivar-specific vessel characteristics affecting active sapwood transport efficiency, and rootstock genetics influencing rate of sapwood aging and conversion to less active state. Active rootstock sapwood temporal dynamics includes= continuously forming new active sapwood (vascular cambium annually producing new xylem adding to active sapwood zone at periphery), gradually losing active function (older active sapwood progressively declining in function transitioning to less active inner sapwood), showing 3-8 year functional lifespan (annual rings remaining in peak active state for several years before gradual functional decline), exhibiting rootstock-specific longevity (some rootstock varieties maintaining peak sapwood function longer than others), and demonstrating condition-dependent persistence (favorable growing conditions potentially extending active functional period while stress accelerating decline). Active rootstock sapwood transport priority includes= receiving priority for water distribution (most efficient pathways in active sapwood preferentially conducting water during transpiration), supplying most recently formed cambium and phloem (active sapwood supporting current season vascular tissue development and function), providing water for current season growth (active sapwood transport directly supporting expanding shoots, leaves, and fruit), maintaining connection to actively growing roots (active sapwood linked to current season root development and function), and ensuring vine survival under stress (active sapwood maintaining minimal transport even when older sapwood fails under severe conditions). Active rootstock sapwood significance includes= providing majority of functional transport capacity (active sapwood conducting most trunk water flow supporting canopy transpiration and productivity), storing most readily available carbohydrate reserves (active sapwood reserves immediately accessible for growth and stress response), maintaining optimal defensive capability (active sapwood cells most able to respond rapidly to damage or infection), representing peak efficiency zone (active sapwood tissue operating at maximum functional capacity per unit volume), supporting current season productivity (active sapwood function essential for annual growth, flowering, and fruit development), and indicating trunk health (extent and condition of active sapwood reflecting overall trunk vigor and functional status).
Active_Rootstock_Sapwood emphasizes outer peak-function sapwood zone and is distinguished from Rootstock_Sapwood (all functional sapwood including declining inner portions) and Conducting_Rootstock_Sapwood (functional emphasis without peak activity specification) by focus on maximum efficiency region. Critical for understanding peak transport capacity, optimal reserve storage, most vigorous defensive responses, majority water conduction, and the prime functional zone providing critical support for current season vine productivity.
Aktives Unterlagen-Splintholz
Most recent rootstock sapwood
Optimal funktionelles Wurzelstockholz
Optimal rootstock sapwood
Peak function rootstock sapwood
Spitzen-Leitsplintholz
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Tyree, M. T., & Zimmermann, M. H. (2002). Xylem Structure and the Ascent of Sap (2nd ed.). Springer.
Active Rootstock Sapwood
Merlot grafted on 3309C rootstock sampled mid-June showing active rootstock vascular cambium with cells progressing through mitosis, microscopic examination revealing mitotic figures in cambial zone indicating active division, cambium producing xylem mother cells differentiating into earlywood vessels 100-120 micrometers diameter and phloem mother cells differentiating into sieve elements, active cambium zone 6-8 cells thick including cambial initials and recent undifferentiated derivatives, cambial cells showing dense cytoplasm and active nuclei indicating elevated metabolic activity, estimated division frequency approximately 3-4 days based on derivative cell accumulation
Active rootstock vascular cambium is rootstock vascular cambium in state of active cell division producing xylem and phloem derivatives, typically occurring during growing season April-October in Northern Hemisphere with peak activity May-July, characterized by cells actively progressing through cell cycle with DNA replication and mitosis, showing visible mitotic figures in microscopic examination, producing xylem mother cells inward that differentiate into current season's wood increment and phloem mother cells outward that differentiate into current season's inner bark increment, demonstrating elevated metabolic activity with increased respiration and protein synthesis, and responding to environmental conditions including temperature, water availability, and hormonal signals that modulate division rate determining annual growth increment width. Active rootstock vascular cambium cell division includes= showing periclinal mitosis (fusiform initials dividing parallel to trunk surface producing derivatives toward xylem or phloem), exhibiting division frequency (active cambium dividing every 2-5 days during peak season compared to weeks or no division in slow or dormant periods), producing xylem mother cells (daughter cells positioned inward differentiating into vessel elements, fibers, or parenchyma), generating phloem mother cells (daughter cells positioned outward differentiating into sieve elements, companion cells, fibers, or parenchyma), and maintaining cambium (each division retaining one daughter cell as cambial initial preserving meristematic population). Active rootstock vascular cambium seasonal timing includes= beginning activation in spring (cambium resuming division April-May as temperatures exceed threshold typically 10-15°C), reaching peak activity in early season (maximum division rate May-June producing widest earlywood vessels and most phloem), continuing through summer (sustained activity June-August producing latewood and additional phloem though at declining rate), declining in late season (slowing division September as temperatures cool and photoperiod shortens), and ceasing by fall (cambium ending active phase October entering dormant state for winter). Active rootstock vascular cambium xylem production includes= forming earlywood in spring (active cambium producing large-diameter vessels optimized for water transport during April-June peak activity), creating latewood in late summer (active cambium producing smaller vessels and thick-walled fibers July-September), showing declining vessel size (progressive reduction in vessel diameter as season advances reflecting cambial activity decline), generating fibers throughout (producing mechanical support cells continuously during active period), and creating axial parenchyma (forming storage and transport cells interspersed among vessels and fibers). Active rootstock vascular cambium phloem production includes= producing sieve elements (active cambium generating food-conducting cells continuously during growing season), creating companion cells (producing metabolically active cells associated with each sieve element), forming phloem fibers (generating support cells contributing to bark strength), generating phloem parenchyma (producing storage and secretory cells in secondary phloem), and showing continuous production (phloem formation occurring throughout active period unlike xylem showing distinct earlywood-latewood phases). Active rootstock vascular cambium metabolic characteristics includes= exhibiting elevated respiration (active cambium cells consuming oxygen and producing ATP supporting division and biosynthesis), showing increased protein synthesis (producing enzymes, structural proteins, and cell wall materials needed for division and differentiation), demonstrating DNA replication (S phase of cell cycle producing duplicate chromosomes prior to mitosis), displaying active mitosis (M phase with visible chromosome condensation, spindle formation, and cell plate formation), and requiring energy supply (active cambium depending on photosynthate supply from leaves or stored reserves supporting metabolic demands). Active rootstock vascular cambium environmental responses includes= responding to temperature (cambial activity increasing with temperature up to optimum around 20-25°C then declining at higher temperatures), affected by water availability (drought stress potentially slowing or halting cambial division reducing growth increment), influenced by photoperiod (day length affecting activity duration with long days promoting sustained activity), modulated by hormonal signals (auxin, cytokinin, and gibberellin levels affecting division rate and differentiation patterns), and showing stress sensitivity (active cambium vulnerable to freeze damage, heat stress, or mechanical injury during active period). Active rootstock vascular cambium and annual ring width includes= determining ring width (duration and rate of cambial activity directly controlling annual xylem increment width), producing wide rings (vigorous sustained cambial activity under favorable conditions creating 0.6-1.0+ cm annual rings), generating narrow rings (limited or brief cambial activity under stress or in old trunk creating 0.1-0.3 cm annual rings), reflecting growing conditions (cambial activity level recording water availability, temperature, nutrients, and vigor during formation year), and enabling dendrochronology (cambial activity patterns preserved in ring widths providing chronological environmental record).
Active_Rootstock_Vascular_Cambium emphasizes state of active cell division and is distinguished from dormant cambium by focus on mitotic activity, derivative production, and growing season function. Critical for understanding seasonal growth, annual ring formation, xylem-phloem production rate, environmental response, and the active meristematic state producing current season's wood and bark increment.
Aktives Unterlagen-Gefäßkambium
Dividing rootstock vascular cambium
Functional rootstock vascular cambium
Funktionelles Gefäßkambium
Growing rootstock cambium
Teilendes Wurzelstock-Kambium
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Evert, R. F. (2006). Esau's Plant Anatomy= Meristems, Cells, and Tissues of the Plant Body (3rd ed.). Wiley-Interscience.
Active Rootstock Vascular Cambium
Merlot active sapwood at 10 years post-planting comprises outermost 12-16 mm depth (4-5 most recent annual rings), contains vessels 160-280 μm diameter with 85-95% functional conducting capacity providing 32-48 L/day water transport (representing 70-75% of total trunk hydraulic capacity), parenchyma cells storing 38-52 g starch at 11-14% dry weight concentration (comprising 75-80% of total trunk reserves), shows high metabolic activity with rapid spring starch mobilization, and exhibits optimal wood structure without disease or decay symptoms
Active sapwood is the most recently formed sapwood comprising the outermost trunk wood rings where xylem vessels are maximally functional for water transport, parenchyma cells are fully metabolically active with high starch storage capacity, and all wood functions (conduction, storage, defense, repair) are optimal, representing the highest quality functional wood critical for vine water relations and carbohydrate reserves. Active sapwood typically comprises outermost 3-6 annual rings in mature grapevines= most recent rings showing maximum vessel functionality (vessels clean and open without occlusion), parenchyma at peak metabolic capacity (high enzyme activity, responsive to signals), maximum carbohydrate storage (starch concentration 10-15% dry weight compared to 4-8% in older sapwood), and optimal anatomical structure (minimal age-related degradation or dysfunction). Active sapwood anatomical characteristics include= xylem vessels completely free of tyloses, gums, or other occlusions (100% of vessels potentially functional for transport), vessel walls with fully functional bordered pits (maximum lateral water movement capacity), living parenchyma cells with dense cytoplasm and abundant organelles (indicating high metabolic activity), starch grains densely packed in parenchyma cells (maximum storage per unit volume), ray parenchyma with intact cellular connections (optimal radial transport), and pristine wood structure without decay, discoloration, or pathogen presence. Active sapwood hydraulic characteristics include= maximum specific hydraulic conductivity (3-10 kg m⁻¹ MPa⁻¹ s⁻¹ compared to 1-5 in older sapwood), highest proportion of functional vessels (80-95% of anatomically present vessels actively conducting), large earlywood vessels fully functional (vessels 180-300 μm diameter providing majority of conductance), minimal embolism under normal conditions (though still vulnerable to severe drought or freeze), and optimal vessel-to-vessel connectivity (pit membranes fully functional enabling lateral water movement). Active sapwood carbohydrate storage characteristics include= highest starch concentration (10-15% dry weight compared to 5-10% in inner sapwood), rapid starch accumulation capacity (efficiently storing photosynthate during post-harvest reserve building), efficient starch mobilization (rapid conversion to soluble sugars during spring budbreak), maximum storage capacity per unit volume (densely packed starch grains in healthy parenchyma), and optimal seasonal cycling (complete accumulation-depletion-replenishment cycle each year). Active sapwood metabolic characteristics include= high respiratory activity (parenchyma cells actively consuming oxygen, producing ATP), active enzyme systems (starch synthases, hydrolases, peroxidases, defense enzymes at high activity), responsive to hormone signals (rapid response to auxin, cytokinin, abscisic acid regulating growth and stress responses), active defense compound synthesis (producing stilbenes, phenolics, other antimicrobial compounds in response to pathogen challenge), and potential embolism repair capacity (though limited in grapevine, highest in active sapwood if repair occurs). Active sapwood development reflects= formation as most recent cambial products (current year and previous 2-5 years depending on growth rate), maintenance of functionality through optimal metabolic status (living cells well-maintained preventing premature dysfunction), position immediately interior to vascular cambium (direct connection to phloem facilitating photosynthate loading), and eventual transition to less active sapwood as new rings form externally (gradual decline in function as wood ages). Active sapwood proportional representation varies= in young rapidly growing vines (years 3-8) active sapwood may comprise 70-90% of total sapwood depth (6-10 mm active out of 10-14 mm total sapwood), in mature moderate-growth vines (years 10-25) active sapwood 50-70% of total (10-18 mm out of 20-28 mm total), in old slow-growing vines (years 30-50+) active sapwood 40-60% of total (8-15 mm out of 18-25 mm total), with absolute depth relatively stable but proportional contribution declining with vine age. Active sapwood significance includes= providing majority of hydraulic capacity (active sapwood vessels conducting 60-80% of total water transport despite being only 50-70% of sapwood depth), providing majority of storage reserves (active sapwood containing 65-80% of trunk starch despite being half to two-thirds of sapwood volume), representing prime functional wood target for protection (trunk disease prevention critically important preserving active sapwood), and indicating current vine vigor and physiological status (active sapwood depth and quality reflecting recent growth conditions and vine health).
Active_Sapwood emphasizes recently formed maximally functional wood and is distinguished from Sapwood (entire functional zone including older declining sapwood) and Conducting_Sapwood (transport function emphasis) by focus on peak functional capacity in most recent rings. Critical for understanding prime hydraulic capacity, maximum carbohydrate storage quality, trunk disease impacts on newest wood, and relationship between recent growth and current vine functional capacity.
Aktives Splintholz
Frisches Funktionsholz
Outer conducting wood
Prime sapwood
Primäres Leitholz
Recent functional wood
Active Sapwood
In a Mendoza Malbec vineyard, active sugar accumulation was tracked over 32 days from mid-February through mid-March Year N+1 (BBCH 84-86) when berry Brix increased from 15.8° to 25.2° (9.4° gain, average 2.9% per week), showing consistent daily rates 0.26-0.34% Brix per day (CV 12%), hexose import estimated 72-105 μmol g⁻¹ h⁻¹ based on Brix trajectory, berry fresh weight 1.42 g → 1.95 g (37% expansion), under stable Argentine high-altitude conditions (daily high 26-29°C, moderate deficit stem ψ -1.15 MPa concentrating sugars while maintaining import).
Active_Sugar_Accumulation represents the sustained period of consistent hexose import and storage following initial accumulation establishment when transport machinery operates at elevated capacity producing steady sweetness increase creating the prolonged sweetening phase that accounts for majority of total sugar gain and establishes foundation for achieving commercial harvest maturity. This active accumulation occurs during mid-veraison through mid-ripening (BBCH 84-86), typically days 12-40 post-veraison onset spanning 25-35 days depending on cultivar and conditions, and is characterized by berry Brix increasing steadily from 13-19° at active onset to 21-27° at conclusion representing 7-13° gain during active period accounting for 50-70% of total ripening accumulation, daily Brix increase 0.18-0.35% per day (1.3-2.5% per week initially accelerating to 2.5-4.0% per week mid-active then moderating toward conclusion) showing relatively consistent rates compared to initial acceleration and final deceleration phases, hexose import rates sustained 45-120 μmol combined glucose and fructose g⁻¹ h⁻¹ representing mature transport capacity, and cumulative sugar gain 90-150 g/L over active period representing major contribution to final berry sweetness. Physiological characteristics during active accumulation include transporter gene expression maintained at elevated steady-state 80-120 fold above green berry (VvHT1, VvHT2, VvHT3, VvHT5, VvHT6 collectively) with balance between synthesis and degradation sustaining high protein levels, enzyme activities stable with cell wall invertase 60-120 units/g fresh weight continuously cleaving phloem sucrose, membrane transport capacity 70-140 μmol hexose g⁻¹ h⁻¹ plasma membrane uptake and 55-110 μmol g⁻¹ h⁻¹ tonoplast sequestration operating efficiently, hormonal regulation with ABA sustained 1400-2600 ng/g maintaining ripening program activation throughout active period, and metabolic integration where hexose import coordinated with partial respiration (25-35% of import oxidized) supporting ATP demands while majority (65-75%) accumulates ensuring efficient storage. Temporal stability during active accumulation demonstrates relatively constant day-to-day rates 0.20-0.38% Brix per day with <15-25% coefficient of variation over multi-week period indicating metabolic homeostasis, contrast with initial phase showing progressive acceleration as capacity establishes and final phase showing deceleration as limits approached, and environmental responsiveness maintaining rates within normal range under favorable conditions while moderating under stress demonstrating regulatory capacity preventing excessive fluctuation. Source-sink relationships during active period show sustained phloem delivery 180-350 mg sucrose per berry per day supported by canopy photosynthesis 12-25 g CO₂ fixed per vine per day producing adequate substrate, sink strength high with berry sugar demand dominant carbon sink receiving 70-85% of daily photosynthate, source capacity generally non-limiting under appropriate crop loads (0.8-1.2 kg fruit per m² leaf area) enabling sustained import, and balance between source supply and sink demand optimized through developmental programming and environmental acclimation maintaining steady accumulation without severe fluctuation. Cellular processes supporting sustained accumulation include vacuolar expansion continuing from 93-95% to 95-97% cell volume accommodating increasing solute loads, tonoplast membrane area increasing 40-80% over active period through proliferation supporting enhanced transport capacity, osmotic potential declining progressively -2.0 to -2.6 MPa → -2.4 to -3.2 MPa creating maintained water potential gradient favoring continued water import, and cell wall mechanical properties adjusting through expansin activity and polysaccharide modification enabling continued volume increase despite increasing turgor pressure. Environmental conditions supporting active accumulation include temperature optimum 25-30°C enabling maximum metabolic rates with this range producing fastest accumulation 3.0-4.5% Brix per week, moderate warmth 22-26°C supporting appropriate rates 2.2-3.5% per week achieving gradual progression, cooler conditions 18-22°C limiting to 1.5-2.5% per week extending duration to achieve final levels, heat stress 32-35°C potentially impairing 30-60% through enzyme impairment and membrane disruption, and cumulative heat during active period (growing degree-days) strongly correlating with total accumulation enabling predictive modeling. Water relations during active period demonstrate adequate vine water status essential with stem water potential -0.6 to -1.2 MPa supporting normal metabolism, moderate deficit -1.0 to -1.4 MPa generally compatible potentially enhancing concentration through reduced berry expansion with maintained import, severe deficit <-1.5 MPa impairing accumulation 35-65% through reduced phloem delivery and metabolic limitation, and irrigation timing strategically managing water availability balancing accumulation support against avoiding excess promoting vigor or dilution. Cultivar differences in active accumulation show rapid-accumulating varieties (Chardonnay, Gewürztraminer) displaying 3.5-4.8% Brix per week sustained 22-28 days achieving high final sugars efficiently, moderate cultivars (Merlot, Syrah) showing 2.8-4.0% per week over 28-35 days, and slower types (Nebbiolo, Carignan) exhibiting 2.2-3.2% per week extended 32-42 days reflecting genetic variation in transporter capacity and metabolic programming. Quality implications include active accumulation rates and duration determining whether target sugars achieved within available growing season, uniformity of active period across vineyard zones affecting harvest maturity consistency requiring zone-specific management, coordination between sugar accumulation and acid decline during active period establishing developing balance critical for wine style, and timing of active accumulation relative to phenolic maturity determining optimal harvest window particularly reds where tannin development must synchronize with sugar levels. Harvest timing considerations recognize active accumulation as critical period approaching maturity requiring increased monitoring frequency, weekly Brix sampling tracking rates enabling projection of harvest timing 2-3 weeks advance, integration with pH, TA, phenolic, and sensory assessment ensuring comprehensive maturity determination, and recognition that conclusion of active accumulation typically signals approaching harvest window requiring readiness. Viticultural management during active period includes maintaining canopy health ensuring sustained photosynthesis supporting continued accumulation, irrigation strategy providing adequate water without excess, disease control preventing infections disrupting metabolism, crop load management ensuring appropriate fruit-to-leaf ratios enabling adequate source capacity, and environmental monitoring tracking conditions affecting accumulation rates informing harvest planning. Metabolic efficiency during active period shows optimized carbon use with high proportion of imported hexoses (65-75%) accumulating versus respired (25-35%) representing efficient storage metabolism, ATP production from partial hexose oxidation supporting transport and biosynthesis costs estimated 15-25 μmol ATP g⁻¹ h⁻¹ from hexose respiration plus 30-55 μmol g⁻¹ h⁻¹ from concurrent organic acid oxidation providing total 45-80 μmol g⁻¹ h⁻¹ adequate for ripening demands, and energetic balance where combined sugar storage and acid catabolism creates metabolically efficient ripening strategy minimizing wasteful respiration while supporting necessary biosynthesis.
Active_Sugar_Accumulation is distinguished from Sugar_Accumulation_Onset by representing sustained phase (25-35 days, 7-13° Brix gain) rather than initial establishment (6-10 days, 5-8° Brix), from Peak_Sugar_Accumulation by showing steady rather than maximum rates (2.5-4.0% per week vs 3.5-5.0%), and from late accumulation by exhibiting consistent rates rather than deceleration. Duration days 12-40 post-veraison BBCH 84-86. Accounts for 50-70% total ripening sugar gain. Steady-state metabolism. Active_Sugar_Accumulation represents sustained sweetening phase achieving majority of commercial sugar levels.
anhaltender Hexose-Import
consistent accumulation period
konsistente Akkumulations-Periode
prolonged sugar storage
steady sweetening phase
stetige Süßungs-Phase
sustained hexose import
verlängerte Zucker-Speicherung
Keller M (2020) The Science of Grapevines 3rd Ed pages 223-227 sustained accumulation Coombe BG, McCarthy MG (2000) Dynamics and environmental control of grape berry growth Australian J Grape Wine Res 6:73-83 accumulation dynamics Ollat N, Gaudillère JP (1998) The effect of limiting leaf area during stage I of berry growth on development and composition of berries of Vitis vinifera L. cv. Cabernet Sauvignon Amer J Enol Vitic 49:251-258 sustained accumulation and source-sink.
Active Sugar Accumulation
In a Napa Valley Cabernet Sauvignon vineyard, active sugar loading was documented over 18 days in late August to mid-September Year N+1 (BBCH 84-85) when Brix increased from 19.2° to 25.8° (average 3.8% per week), cell wall invertase activity averaged 102 units/g FW, hexose uptake rates 125 μmol g⁻¹ h⁻¹, vacuolar concentration rose from 1050 to 1680 mM, berry fresh weight expanded 12% to final 1.95 g, under warm dry conditions (daily high 28-32°C, moderate deficit stem ψ -1.2 MPa concentrating sugars while maintaining metabolism).
Active_Sugar_Loading represents the high-flux state when all components of the sugar import pathway operate at maximum efficiency creating sustained rapid hexose accumulation that elevates berry sweetness from moderate veraison levels to harvest-ready concentrations defining wine alcohol potential and style. This active loading phase occurs during mid-to-late veraison through early ripening (BBCH 84-85), typically 8-14 days post-accumulation onset, and is characterized by coordinated maximum operation: cell wall invertase activity 70-130 units/g FW producing glucose and fructose at 90-180 μmol hexose g⁻¹ h⁻¹ from phloem-delivered sucrose, plasma membrane hexose transporters operating near Vmax with combined glucose and fructose uptake 80-150 μmol g⁻¹ h⁻¹, tonoplast transport efficiently sequestering cytoplasmic hexoses into vacuoles at 60-110 μmol g⁻¹ h⁻¹, and net berry sugar accumulation sustained at 3.0-4.5% Brix per week increasing total sugars from 160-220 g/L to 220-280 g/L (22-28° Brix) during 2-4 week active loading period. Molecular coordination during active loading shows: hexose transporter genes maintained at maximum steady-state expression (VvHT1/2/3/5/6 collectively 400-600 fold above green berry baseline when summed), invertase genes sustained at high levels (VvcwINV 80-120 fold, VvGIN 50-90 fold) ensuring continued sucrose cleavage, regulatory transcription factors (sugar-responsive VvMYB, VvbZIP, VvNAC families) at plateau expression coordinating pathway genes, and feedback mechanisms preventing overshooting with very high hexose levels (>1500-2000 mM vacuolar) beginning to reduce transporter expression in some cells approaching saturation providing homeostatic control. Cellular physiology during active loading demonstrates: mesocarp cell vacuoles occupying maximum volume fraction 94-97% of cell creating huge storage capacity (individual vacuoles 1.2-3.0 nL), vacuolar hexose concentrations rising 120-200 mM per week from 800-1100 mM toward final 1400-2200 mM (berry-dependent and cultivar-dependent maxima), cytoplasmic hexose maintained 55-95 mM through balanced import, metabolism, and vacuolar sequestration preventing accumulation that could inhibit continued uptake, and osmotic potential declining -0.15 to -0.25 MPa per week from -1.8 to -2.4 MPa toward final -2.4 to -3.4 MPa creating strong water potential gradient sustaining berry expansion 8-15% during active loading despite declining growth rates. Phloem transport supporting active loading requires: adequate source leaf photosynthesis providing sucrose export with healthy canopies producing 8-15 g photosynthate per m² leaf area per day, phloem loading in leaves and transport through vine vascular system delivering sucrose to berries at 200-400 mg per berry per day depending on berry size and accumulation rate, sieve tube pressure gradients maintained -0.5 to -1.2 MPa from leaves to berries driving mass flow, and vine-level source-sink balance with typical leaf area: fruit weight ratios 12-18 cm² per g supporting active loading without source limitation. Apoplastic dynamics during active loading show: continuous sucrose delivery to berry apoplast creating transient concentrations 120-280 mM before invertase cleavage, rapid enzymatic turnover producing glucose and fructose with residence time 3-10 minutes before membrane transport, resulting apoplastic hexoses 180-400 mM combined maintaining strong concentration gradient favoring import (apoplast 280-400 mM → cytoplasm 60-90 mM = 220-340 mM gradient), and efficient coupling between cleavage and transport preventing apoplastic accumulation that could feedback-inhibit either process. Energy metabolism supporting active loading includes: enhanced respiration 1.5-2.8 μmol O₂ g⁻¹ h⁻¹ providing ATP for membrane pumps and biosynthetic processes, respiratory quotient >1.2-1.5 indicating substrate-level metabolism dominates with fermentative activity particularly in berry interior where O₂ diffusion limits aerobic respiration, ATP production estimated 50-95 μmol g⁻¹ h⁻¹ with 20-30% allocated to transport-associated processes including H⁺-ATPases maintaining electrochemical gradients essential for transport function, and glycolytic flux metabolizing 20-40% of imported hexoses providing energy and biosynthetic precursors while 60-80% accumulates in vacuoles as storage sugars. Hormonal regulation during active loading demonstrates: ABA sustained at ripening plateau 1500-2600 ng/g maintaining maximum program activation, hexose signaling maximized through hexokinase pathway with glucose-6-phosphate 30-60 μmol/g FW creating strong feedback amplification of sugar-responsive genes, gibberellin and auxin at post-growth minima removing any suppressive effects, and ethylene showing cultivar-dependent patterns with some varieties 0.5-1.5 nL g⁻¹ h⁻¹ potentially fine-tuning ripening while others baseline suggesting non-essential role. Environmental modulation of active loading shows: temperature critically important with optimal 26-30°C supporting maximum rates 4.0-4.8% Brix per week, moderate warmth 23-26°C enabling 3.2-4.2% per week, cooler conditions 20-23°C reducing to 2.5-3.5% per week through slowed kinetics, heat stress 32-36°C impairing to 1.8-3.0% per week through enzyme denaturation and membrane dysfunction, and extreme heat >36°C potentially arresting loading <1.0% per week or completely through catastrophic cellular disruption. Light effects on active loading operate primarily through source photosynthesis with adequate canopy light exposure essential for providing sucrose substrate, shaded canopies potentially limiting loading 20-40% despite full berry import capacity through substrate limitation, leaf removal during active loading requiring careful balance between cluster exposure benefits and photosynthetic capacity maintenance, and canopy architecture optimization critical for maximizing both fruit microclimate and source leaf productivity. Water relations influence shows: moderate deficit (stem ψ -0.9 to -1.4 MPa) compatible with active loading and potentially enhancing concentration through reduced berry expansion, severe deficit (<-1.5 MPa) impairing loading 30-60% through general metabolic limitation, osmotic stress effects complex with berry water potential -2.0 to -3.0 MPa creating strong gradient for water import requiring adequate vine water status to sustain, and irrigation management during active loading balancing concentration benefits of deficit against metabolic maintenance requiring adequate water. Cultivar variation in active loading characteristics demonstrates: high-sugar cultivars (potential final 25-30° Brix) maintain active loading 4.2-5.0% Brix per week for 2-3 weeks, moderate varieties (final 22-26° Brix) show 3.5-4.5% per week for 2.5-4 weeks, and lower-sugar types (final 19-23° Brix) display 2.8-3.8% per week for 3-5 weeks reflecting genetic differences in loading duration and intensity more than maximum rate capabilities. Coordination with ripening completion shows: active sugar loading coincides with final berry softening (approaching asymptotic minimum firmness), color development completing in reds (anthocyanins reaching 90-100% final levels), acid decline approaching endpoint with malate near minimum sustainable levels, and flavor/aroma maturation intensifying creating integrated ripeness suitable for harvest timing decisions. Quality implications of active loading inclu loading rate and duration critically determining final Brix and thus wine alcohol potential with faster sustained loading enabling higher sugar or earlier harvest at target levels, uniformity of active loading across berries and clusters essential for maturity synchrony affecting harvest timing and compositional consistency, balance between sugar and other quality parameters (phenolics, acids, flavors) defining optimal harvest window requiring integration of multiple maturity indices, and environmental conditions during active loading substantially affecting wine style through effects on final berry composition. Viticultural management during active loading involves: irrigation typically minimized or terminated unless severe stress developing recognizing berry water demand declines and concentration generally desirable, canopy management ensuring source leaf health and adequate light for photosynthesis without excessive fruit exposure causing heat stress, disease monitoring critical as high-sugar berries increasingly attractive to pests and pathogens, and harvest preparation advancing with active loading indicating approaching maturity requiring logistics, equipment, and labor readiness.
Active_Sugar_Loading is distinguished from Initial_Sugar_Increase by representing maximum sustained (3.0-4.5% Brix/week) rather than moderate (1.5-2.5%/week) rates, from Rapid_Sugar_Accumulation_Start by showing sustained maximum rather than onset, and from sugar plateau by exhibiting continued loading rather than declining rates. Duration 2-4 weeks BBCH 84-85. All pathway components operating near maximum. Temperature optimal 26-30°C critical. Active_Sugar_Loading represents culminating phase achieving harvest-ready sweetness defining wine alcohol potential and style.
Spitzen-Hexose-Beladung
anhaltende rapide Akkumulation
high-efficiency sweetening
hoch-effiziente Süßung
maximaler Zucker-Import-Fluss
maximum sugar import flux
peak hexose loading
sustained rapid accumulation
Keller M (2020) The Science of Grapevines 3rd Ed pages 223-228 active sugar accumulation and maturity Palliotti A, Tombesi S, Silvestroni O, Lanari V, Gatti M, Poni S (2014) Changes in vineyard establishment and canopy management urged by earlier climate-related grape ripening Sci Hort 178:43-54 environmental effects on accumulation Conde C, Silva P, Fontes N, Dias ACP, Tavares RM, Sousa MJ, Agasse A, Delrot S, Gerós H (2007) Biochemical changes throughout grape berry development and fruit and wine quality Food Res Intl 40:1-13 comprehensive ripening biochemistry.
Active Sugar Loading
Merlot active vascular cambium in late May (peak activity) shows 5-6 cell layer zone with cambial initials exhibiting dense cytoplasm, large nuclei 8-12 μm diameter occupying 25-30% cell volume, visible mitotic figures in 3-8% of cells, periclinal divisions producing xylem mother cells (differentiating into 180-240 μm earlywood vessels) inward and phloem mother cells outward at 7':1 ratio, cells incorporating tritiated thymidine indicating active DNA synthesis, high respiration rate 2.5-4.0 μmol O₂/g fresh weight/hour, producing 0.8-1.2 mm radial xylem increment per month during peak period
Active vascular cambium is vascular cambium during periods of cell division and secondary tissue production, characterized by metabolically active initials undergoing periclinal and anticlinal divisions, producing xylem and phloem derivatives, and exhibiting high rates of DNA synthesis, protein production, and cell wall formation, representing the growth-phase state of cambial meristem. Active vascular cambium timing includes= spring reactivation period (late March-April, 0-2 weeks before to 2-4 weeks after budbreak), peak activity period (May-July during rapid shoot growth and early fruit development), sustained activity (continuing through August during fruit ripening), gradual decline (September as shoot growth ceases), and cessation (late September-October transitioning to dormant state), with total active period typically 24-28 weeks (approximately 6-7 months) in temperate climates. Active vascular cambium cellular characteristics include= cambial initials with dense cytoplasm (high organelle density including abundant mitochondria, ribosomes, and endoplasmic reticulum), large prominent nuclei (occupying 20-35% of cell volume reflecting active transcription), thin primary cell walls (allowing cell division and expansion), high turgor pressure (maintaining cell rigidity and driving expansion), active cell cycle progression (initials cycling through G1-S-G2-M phases with division intervals 3-10 days depending on growth conditions), and visible mitotic figures (dividing cells showing chromosomes in metaphase, anaphase, or telophase observable in active cambium sections). Active vascular cambium metabolic characteristics include= high respiration rates (consuming oxygen and producing ATP to fuel division and biosynthesis), active DNA replication (S-phase cells synthesizing DNA preparing for mitosis), high protein synthesis rates (ribosomes producing enzymes, structural proteins, and cell wall components), active cell wall synthesis (producing cellulose, hemicelluloses, and pectins for new cell walls), hormone responsiveness (auxin and cytokinin receptors active triggering signal transduction cascades), and high sugar uptake (importing photosynthate from phloem to fuel energy-intensive division). Active vascular cambium division dynamics include= periclinal divisions producing xylem mother cells inward (approximately 5-10 divisions per cambial position during active season producing xylem derivatives) and phloem mother cells outward (approximately 1-2 divisions producing phloem derivatives), anticlinal divisions increasing cambial circumference (1-2 anticlinal divisions per 10-15 periclinal divisions maintaining cambial continuity as diameter expands), asymmetric division ratios (strong centripetal bias producing 5-10× more xylem than phloem cells), division rates varying seasonally (peak rates May-June 1-2 divisions per week declining to 0.5-1 division per week August-September), and division synchrony (some coordinate activity across cambial zones, other sections showing asynchronous division). Active vascular cambium hormone signaling includes= high auxin responsiveness (auxin receptor genes expressed, signal transduction pathways active), auxin transport (polar auxin transport from shoots to cambium maintaining basipetal hormone gradient), cytokinin presence (root-derived cytokinins in xylem sap interacting with cambial cells), gibberellin effects (modulating cell expansion and differentiation of cambial derivatives), brassinosteroid involvement (steroid hormones promoting cell division and expansion), and hormone crosstalk (multiple hormone pathways integrating environmental and developmental signals). Active vascular cambium xylem production includes= earlywood formation (April-June producing large-diameter vessels 150-300 μm optimized for water transport), latewood formation (July-September producing smaller vessels 40-120 μm and more fibers emphasizing mechanical support), fiber differentiation (thick-walled fiber cells providing wood strength), ray parenchyma production (ray initials producing radial parenchyma maintaining lateral transport), and vessel element maturation (xylem mother cells expanding, depositing secondary walls, undergoing programmed cell death becoming functional vessels within 2-4 weeks). Active vascular cambium phloem production includes= sieve tube element formation (phloem mother cells differentiating into enucleate sieve tubes with specialized sieve plates), companion cell development (smaller densely cytoplasmic cells maintaining sieve tube function), phloem parenchyma production (storage and transport cells), phloem fiber formation (thick-walled supporting cells in outer phloem), and phloem ray continuity (ray initials producing rays connecting phloem to cambium to xylem). Active vascular cambium environmental responsiveness includes= drought stress reducing activity (water deficit causing premature cessation or intermittent activity), heat stress effects (excessive temperatures >35°C potentially inhibiting division), cold stress dormancy induction (temperatures <10°C reducing activity, <5°C stopping division), nutrient availability affecting rates (nitrogen promoting vigorous cambial activity, deficiency reducing division), and light quality effects (red = far-red ratio potentially influencing cambial activity through phytochrome signaling). Active vascular cambium identification and assessment includes= anatomical observation (thin-walled dividing cells visible in transverse sections), vital staining (dyes like neutral red accumulating in metabolically active cambial cells), incorporation of labeled precursors (radioactive or fluorescent thymidine incorporation during DNA synthesis), cambial zone width (active cambium 4-6 cell layers compared to 2-3 in dormant state), and hormone measurements (high auxin and cytokinin content in active cambial zone). Active vascular cambium significance includes= determining annual growth (cambial activity duration and intensity controlling ring width and bark production), responding to management (irrigation, fertilization, pruning affecting cambial activity and wood production), enabling trunk diameter increase (active cambium adding 1.5-5 mm radius and 5-16 mm circumference annually), producing functional tissues (generating conducting xylem and phloem maintaining vine hydraulic and nutrient transport), and reflecting vine vigor (vigorous vines showing longer active periods and higher division rates, declining vines showing shortened activity and reduced production).
Active_Vascular_Cambium emphasizes growth-phase state and is distinguished from Vascular_Cambium (cambium in all states) and Growing_Season_Vascular_Cambium (seasonal timing emphasis) by focus on cellular and metabolic activity characteristics. Critical for understanding cell division dynamics, seasonal growth patterns, hormonal regulation of cambial activity, wood production rates, and environmental effects on secondary growth.
Aktives vaskuläres Kambium
Dividing cambium
Growing cambium
Proliferative cambium
Teilendes Kambium
Wachsendes Kambium
Larson, P. R. (1994). The Vascular Cambium= Development and Structure. Springer. | Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press.
Active Vascular Cambium
An actively climbing functional tendril at node 8 on a shoot of Vitis vinifera cv. Cabernet Franc in the Loire Valley, France, having made initial contact with a horizontal trellis wire four days previously, currently exhibiting progressive coil development from initial three contact coils to current six contact coils observed over 48-hour monitoring period, showing active shoot displacement with 6 cm movement toward the wire measured between days 3 and 5 post-contact, developing nine free coils in the tendril shaft with visible tightening and increasing coil density, undergoing color transition from green at the shoot-proximal end through tan in mid-regions to light brown in contact coil areas indicating progressive lignification wave, generating estimated 150 grams contractile force based on shoot displacement against gravity, measured at BBCH stage 71 (fruit set) in mid-June during optimal environmental conditions for coiling with temperatures of 22-26°C and adequate soil moisture.
A climbing functional tendril that has recently achieved successful attachment (typically within 3-7 days) and is currently generating active contractile forces pulling the shoot toward the support structure in Vitis vinifera, characterized by ongoing coiling progression with visible changes in coil tightness and number over 24-48 hour observation periods, measurable shoot displacement toward the attachment point as free coils contract and shorten the tendril shaft, active lignification processes indicated by progressive color change from green to tan to brown in coiling regions, and maintenance of tension generation creating spring-like mechanical properties that absorb shock loading and facilitate continued vertical shoot growth. Actively climbing functional tendrils represent the most dynamic phase of tendril function, executing the biomechanical processes that convert exploratory growth into permanent mechanical attachment and enable the characteristic climbing behavior of Vitis species. During this active phase, tendril tissues undergo rapid structural transformations including differential cell expansion on opposite sides of the tendril axis driving coil formation, deposition of gelatinous fibers that generate contractile forces, and progressive lignin and cellulose deposition that fixes coil geometry and increases tissue rigidity. The active climbing period typically spans 3-14 days post-contact depending on environmental conditions, after which tendrils transition to established support status with stable attachment geometry and complete structural reinforcement.
Actively climbing functional tendrils are distinguished from searching climbing tendrils (pre-attachment) and established attached tendrils (post-active phase) by the recent successful attachment combined with ongoing structural and mechanical changes. This active phase is diagnostically important for understanding tendril development success and environmental effects on attachment, as disturbances during this period can interrupt coiling completion and result in weak attachments. The duration of active climbing varies with temperature (shorter in warm conditions), water availability (limited by drought stress), and cultivar genetics (some varieties show faster lignification). Actively climbing tendrils provide both current support and progressive shoot elevation, representing the functional heart of the climbing process that enables Vitis to achieve its lianescent growth strategy.
active attachment tendril
aktiv kletternde Ranke
contracting tendril
fresh climbing tendril
frische Befestigungsranke
kontrahierende Ranke
recently attached tendril
Based on Bowling & Vaughn (2009) "Gelatinous fibers are widespread in coiling tendrils and twining vines" American Journal of Botany 96=719-727 Silk & Hubbard (1991) "Axial forces and normal stresses in coiling tendrils" Biomechanics and Modeling in Mechanobiology 24=599-606 Goriely & Tabor (2013) "Spontaneous helix hand reversal and tendril perversion in climbing plants" Physical Review Letters 80=1564-1567 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 259-282.
Actively Climbing Functional Tendril
An actively growing tip leaf at node 13 on a highly vigorous shoot of Vitis vinifera cv. Merlot in Pomerol, France, positioned 5 cm behind the shoot apex in the zone of maximum expansion, exhibiting area increase from 18 cm² to 52 cm² over 5 days representing daily growth rate of 6.8 cm² per day (10.2% daily compound growth), dimensions increasing from 5.5 cm × 4.8 cm to 9.2 cm × 8.5 cm with visible daily progression, cells in palisade regions enlarging from average 20 μm to 28 μm diameter over the measurement period representing 1.6 μm per day, tissue water content of 91% fresh weight indicating maximum turgor supporting rapid expansion, very soft texture easily deformed by gentle handling, rapidly deepening coloration with chlorophyll increasing from 220 μg cm⁻² to 340 μg cm⁻² over the 5-day period, consuming approximately 25 mg carbohydrate per day and 4.2 mg nitrogen per day supported by strong translocation from mature leaves, photosynthetic rate increasing from 3.2 to 5.8 μmol CO₂ m⁻² s⁻¹ approaching 48% mature capacity, occurring under optimal conditions including daily maximum temperatures 25-27°C, soil moisture at 78% field capacity, adequate nitrogen from fertilization, measured at shoot BBCH stage 71 (fruit set) in mid-June during peak vegetative growth, representing maximum expansion rate typical of vigorous shoots under optimal conditions.
A growing tip leaf exhibiting maximum growth rates during peak developmental progression in Vitis vinifera, characterized by position in the zone of most rapid expansion within 4-8 cm of the shoot apex, very high daily area increase rates typically 4-8 cm² per day representing 8-12% daily compound growth, rapid visible growth progression evident without measurement instruments, active cell expansion throughout lamina with cells enlarging at 2-4 μm per day in linear dimensions, high tissue water content of 88-92% fresh weight reflecting intense water uptake driving turgor-based expansion, very soft pliable texture from actively loosening cell walls with high pectin content allowing maximum extensibility, rapidly deepening green coloration as chlorophyll synthesis accelerates, substantial daily resource consumption including 18-30 mg carbohydrate per day for cell wall synthesis and metabolic processes and 3-5 mg nitrogen per day for protein synthesis, rapidly increasing photosynthetic capacity approaching 40-60% of mature rates by end of active growth phase, and occurrence under optimal environmental conditions including favorable temperatures of 23-28°C, abundant water availability, adequate nutrients, and moderate light exposure. Actively growing tip leaves represent peak expression of leaf developmental rates within the growing tip zone, consuming maximum resources while progressing most rapidly toward functional maturity, requiring continuous favorable conditions to sustain high growth rates, and showing greatest sensitivity to environmental perturbations that can rapidly slow or arrest development.
Actively growing tip leaves represent the peak growth rate subset of growing tip leaves, distinguished by very high daily area increase, rapid visible progression, and maximum resource consumption. Achievement of active rapid growth requires optimal combinations of environmental and physiological conditions, with any limiting factor rapidly reducing growth rates. Temperature strongly affects growth rate with Q₁₀ approximately 2.5 within physiological range, water availability is critical for turgor-driven expansion, nitrogen supports protein synthesis for membranes and enzymes, and moderate light provides photosynthetic energy without photoinhibition. The active growth phase typically lasts 5-10 days under optimal conditions representing the period of most rapid size increase, followed by declining growth rates as leaves transition toward final expansion completion. Stress during active growth immediately reduces expansion rates, with water deficit having fastest impact through turgor loss, nutrient limitation affecting sustained growth, and temperature extremes disrupting metabolic processes. Actively growing tip leaves show maximum growth sensitivity to cultural practices including irrigation timing, fertilization schedules, and environmental management, with optimal conditions during this phase maximizing final leaf size and quality.
Höchstentwicklung Weinblatt
aktiv-wachsendes Spitzenblatt
maximum growth rate apical leaf
peak development leaf
rapid expansion tip leaf
schnellste Expansion apikales Blatt
Based on Poni et al. (2009) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 60=397-407 Tsukaya (2013) "Leaf development" The Arabidopsis Book 11=e0163 Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 155-184.
Actively Growing Tip Leaf
Acylated anthocyanins comprise 10-65% of total anthocyanins across V. vinifera cultivars, with high-acylation types such as Syrah (50-65% acylated), Mourvèdre (45-60%), and Grenache (40-55%) contrasting with low-acylation cultivars such as Pinot Noir (5-15%) and Sangiovese (10-20%). The anthocyanin acyltransferase (VvAT) catalyzes transfer of acyl groups from hydroxycinnamoyl-CoA donors to the 6-O-position of the glucose. p-Coumaroyl derivatives predominate in most cultivars, with caffeic and acetic acid conjugates as minor forms. Acylation shifts λmax by 5-15 nm, increases molar extinction coefficient by 10-20%, and enhances color stability through intramolecular copigmentation (the acyl group stacking with the chromophore). Acylated anthocyanins show 30-50% slower degradation rates than non-acylated forms under equivalent storage conditions.
An anthocyanin accumulation disposition that is the inherited capacity of a Vitis vinifera plant to modify anthocyanin-3-O-glucosides through acylation with hydroxycinnamic acids (p-coumaric, caffeic, acetic) at the 6-position of the glucose moiety, with genotype-specific acylation proportions affecting anthocyanin stability, color expression, and copigmentation behavior.
Acylated anthocyanin proportion significantly affects wine color stability and intensity, with high-acylation cultivars retaining color better during aging. The trait shows high heritability (h² = 0.75-0.85) and relatively low environmental modulation compared to total anthocyanin concentration. Acylation pattern is used as a chemotaxonomic marker for cultivar identification and hybrid detection, as American Vitis species and their hybrids typically show different acylation patterns than pure V. vinifera. The intramolecular copigmentation provided by acylation partially substitutes for intermolecular copigmentation with other phenolics.
Acylglucosid-Disposition
Acylierte Anthocyane
Anthocyan-Acylierung
acyl-glucoside proportion
anthocyanin acylation trait
hydroxycinnamic acid conjugation
Rinaldo AR et al. (2015) A grapevine anthocyanin acyltransferase, transcriptionally regulated by VvMYBA, can produce most acylated anthocyanins present in grape skins. Plant Physiology 169:1897-1916 García-Beneytez E et al. (2003) Analysis of grape and wine anthocyanins by HPLC-MS. Journal of Agricultural and Food Chemistry 51:5622-5629
Acylated Anthocyanin Disposition
In Pinot Noir vines of the Willamette Valley, Oregon, adequate chilling achievement occurs by late January when accumulated chilling hours of 1100-1200 exceed the cultivar requirement of approximately 1000 hours, ensuring complete dormancy release confirmed by forcing tests showing uniform budbreak within 10 days, and predicting the synchronous field budbreak within a 7-day window typically observed in this cultivar when spring temperatures become favorable.
Adequate chilling achievement in Vitis vinifera represents the accumulation of sufficient cold exposure to enable complete and uniform dormancy release, satisfying the full chilling requirement without deficit that would compromise subsequent budbreak quality and vineyard performance. This condition is distinguished from partial or marginal chilling that may permit some degree of budbreak but results in the delayed, prolonged, and uneven growth characteristic of inadequate dormancy release. The quantitative definition of adequate chilling depends on cultivar-specific requirements, with adequacy achieved when accumulated chilling hours or units equal or exceed the genetic threshold, typically verified through forcing tests that demonstrate rapid and complete budbreak response indicating full restoration of growth competence. The physiological markers of adequate chilling achievement include ABA concentrations reduced to baseline levels compatible with growth, DAM gene expression declined to minimal levels, chromatin modifications removed from growth gene promoters restoring transcriptional competence, and metabolic systems prepared for the rapid activation required during budbreak. The practical importance of adequate chilling achievement for viticulture relates to the quality of spring growth, with adequate chilling producing the synchronous, vigorous budbreak desired for uniform crop development, while inadequate chilling causes the management challenges of extended budbreak period, variable shoot development, and potential yield reduction from poorly developed shoots. The margin of adequacy affects vineyard performance, with chilling substantially exceeding requirements potentially advancing budbreak timing and increasing frost risk in some environments, while chilling barely meeting requirements may result in acceptable but suboptimal budbreak uniformity. Climate variability creates year-to-year variation in chilling adequacy, with warm winters potentially failing to achieve adequate chilling particularly for high-requirement cultivars, while cold winters may substantially exceed requirements, allowing viticulturists to predict budbreak characteristics based on seasonal chilling accumulation relative to cultivar requirements. Management responses to anticipated inadequate chilling include application of dormancy-breaking chemicals and selection of cultural practices that maximize effective chilling exposure.
Adequate_Chilling_Achievement represents the quantitative sufficiency aspect of Chilling_Requirement_Satisfaction. Distinguished from partial or marginal chilling that permits only incomplete dormancy release. Defined by chilling accumulation meeting or exceeding cultivar-specific requirement. Verified through forcing test response and physiological markers. Critical for budbreak uniformity and vigor. Margin above requirement affects budbreak timing. Subject to year-to-year variation with climate.
adequate cold accumulation
adäquate Chilling-Akkumulation
ausreichende Kälte-Erreichung
complete chilling fulfillment
full chill achievement
hinreichende Kälteakkumulation
sufficient chilling accumulation
vollständige Kältebedürfnis-Erfüllung
Dokoozlian NK (1999) Chilling temperature and duration interact on the budbreak of Perlette grapevine cuttings. HortScience 34: 1054-1056 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Campoy JA et al (2011) High temperatures and time to budbreak in low chill apricot. Environmental and Experimental Botany 73: 134-142
Adequate Chilling Achievement
Merlot in Bordeaux exhibits advanced bud swelling at BBCH 03 in early April after 950-1050 chilling hours and 200-250 growing degree days base 10°C, with buds showing 20-25% volume increase and 0.8-1.2 mm scale separation over 4-5 days under conditions of 13-16°C mean temperature and 65-70% relative humidity, representing typical oceanic climate pattern for this mid-season cultivar
Advanced bud swelling represents the intermediate expansion phase where volumetric increase becomes prominent and bud scales begin separating at margins while green tissue remains concealed, corresponding to BBCH 03 in the developmental sequence. This stage occurs after early swelling has established basic metabolic activity and vascular connection, characterized by buds enlarging 15-25% beyond winter dimensions over 3-6 days as cell expansion accelerates and water content increases substantially. The bud profile transitions from slightly enlarged to markedly swollen with noticeably rounded form, creating obvious visual change from dormant appearance. Scale separation begins at margins where adjacent scales no longer overlap completely, creating visible gaps of 0.5-1.0 mm though brown scale surfaces still dominate visual appearance and no green tissue is exposed. This margin separation results from continued turgor-driven expansion of internal meristematic and primordial tissues pushing outward against mechanical resistance of protective scales. The underlying shoot primordium and first leaf primordia expand rapidly through both cell division and cell enlargement, with cell cycle activation producing new cells while existing cells extend through water uptake and wall loosening. Metabolic activity intensifies markedly with respiration rates 8-12 fold above winter dormancy levels, consuming stored carbohydrates at measurable rates that can be detected through starch depletion in supporting cane tissues. Hormonal regulation shows elevated cytokinin and gibberellin concentrations promoting cell division and expansion respectively, with auxin production initiating in developing shoot apex to coordinate growth polarity. Vascular reconnection approaches completion with differentiated xylem vessels establishing functional water transport pathway from cane to expanding bud tissues, critical for maintaining turgor as expansion accelerates. Water content in bud tissues reaches 55-65% fresh weight compared to 35-40% during winter dormancy, reflecting both cell expansion and increased metabolic hydration requirements. Protein synthesis escalates dramatically to support new cell production and enzyme activity, drawing on nitrogen reserves mobilized from storage proteins in woody tissues. Temperature sensitivity remains high with development rate approximately doubling for each 10°C increase between 10-25°C, making this stage particularly responsive to spring temperature variability. The process requires sustained favorable conditions with 3-5 consecutive days above 12°C for reliable progression, with cool periods temporarily slowing development but not causing regression unless freeze damage occurs. Cultivar variation in progression rate reflects genetic differences in cell division rates and expansion responsiveness, with early varieties like Pinot Noir completing advanced swelling in 3-4 days while late varieties like Mourvèdre require 5-7 days under identical conditions. Regional timing follows early swelling by 5-8 days, occurring early April in Mediterranean climates, mid-April in temperate regions, and late April in continental areas. Viticultural management implications include marking time point for completing pruning operations as delayed pruning beyond this stage risks damaging expanding tissues, establishing reference for initiating early-season disease management particularly against powdery mildew, and providing developmental marker for frost protection preparedness as tissue vulnerability increases substantially. Frost damage at advanced swelling causes widespread cell death through ice formation in highly hydrated expanding tissues, typically killing primary buds and forcing reliance on secondary or basal buds with reduced fruitfulness. Research applications include quantitative analysis of cell division and expansion rates during dormancy exit, hormonal profiling documenting growth regulation transitions, and thermal time modeling for phenology prediction across diverse climatic conditions.
Advanced_Bud_Swelling is the second of three Bud_Swelling subphases, distinguished by prominent expansion and visible scale margin separation but no exposed green tissue. Transitions to Late_Bud_Swelling when green tissue becomes faintly visible through scale gaps. Represents optimal developmental reference point for vineyard operation timing due to clear visual distinction.
deutliche Knospenquellung
fortgeschrittene Knospenschwellung
mid-stage swelling
mittlere Schwellphase
obvious bud expansion
pronounced bud swelling
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 165-172 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 70-78 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Advanced Bud Swelling
Chardonnay cutting showing adventitious roots, hardwood cutting 30 cm length treated with 3000 ppm IBA auxin hormone, microscopic examination revealing adventitious root primordia forming from cambial region 12 days after treatment, primordia emerging through cortex and bark 18 days after treatment, emerged adventitious roots 2.4 cm length at 28 days showing root cap, apical meristem, developing vascular connection to cutting vascular system, cutting producing 8 adventitious roots enabling successful establishment, adventitious rooting representing foundation of Chardonnay vegetative propagation
Adventitious root is root arising from non-root tissue including stems, leaves, or callus rather than from seed radicle or existing root branches, characterized by origin from dedifferentiated or pluripotent cells that undergo developmental reprogramming to form root meristem and primordium, showing initial development within parent tissue before emergence creating root that may or may not develop secondary growth and branching depending on age and function, and serving critical roles in vegetative propagation enabling clonal reproduction through cuttings, wound response allowing root formation after damage, and architectural plasticity providing additional anchorage and absorption particularly in layering or flooding situations, with adventitious roots in grapevine viticulture representing foundation of vegetative propagation system where cuttings produce adventitious roots creating genetically identical vines, demonstrating typical characteristics including formation from cambium or pericycle-like cells in stems, hormone dependence particularly auxin promoting initiation, emergence through cortex and bark creating wound in parent tissue, and potential for complete root system development including lateral branching if cutting successfully establishes, and showing that adventitious root capacity distinguishes species capable of vegetative propagation from those requiring seed reproduction making adventitious rooting fundamental to modern viticulture. Adventitious root origin and initiation includes= showing non-root origin (adventitious roots forming from stems, nodes, callus, or occasionally leaves rather than from radicle or existing roots), exhibiting dedifferentiation (cells at initiation site losing original identity and reprogramming to root meristem fate), demonstrating primordium formation (organized root primordium developing within parent tissue before emergence), revealing endogenous origin (adventitious roots typically forming from internal tissues such as cambium, pericycle-like cells, or callus rather than epidermis), and displaying meristem organization (developing primordium organizing into apical meristem, root cap, and provascular tissue creating functional root structure). Adventitious root hormonal regulation includes= showing auxin requirement (auxin essential for adventitious root initiation with exogenous auxin treatment enhancing rooting), exhibiting cytokinin effects (cytokinins potentially inhibiting rooting creating auxin-cytokinin balance requirement), demonstrating ethylene involvement (ethylene potentially promoting adventitious rooting particularly under stress or flooding), revealing gibberellin effects (gibberellins sometimes inhibiting rooting though effects complex and context-dependent), and displaying hormone interactions (multiple hormones interacting to determine whether and when adventitious roots form). Adventitious root development stages includes= showing induction phase (initial hormone and developmental signals triggering cells to commit to root fate), exhibiting initiation phase (committed cells organizing into root primordium with visible meristem), demonstrating emergence phase (developing root growing through parent tissue and emerging at surface), revealing elongation phase (emerged root extending and producing lateral branches), and displaying maturation phase (adventitious root developing secondary growth and complete root system architecture if conditions permit). Adventitious root in vegetative propagation includes= enabling clonal reproduction (adventitious rooting allowing production of genetically identical plants from cuttings or layers), providing commercial propagation (viticulture industry depending on adventitious rooting for producing millions of vines annually), demonstrating cultivar maintenance (adventitious rooting preserving exact genetic identity of cultivars preventing genetic variation from seed reproduction), revealing rootstock propagation (rootstock varieties propagated through adventitious rooting of hardwood cuttings), and displaying grafting dependency (grafted vines depending on adventitious roots of rootstock cutting for below-ground system). Adventitious root functional characteristics includes= showing anchorage capacity (adventitious roots providing mechanical anchoring comparable to seed-derived roots), exhibiting absorption function (adventitious roots developing root hairs and absorptive capacity supporting plant water and nutrient needs), demonstrating transport function (adventitious roots developing functional xylem and phloem connecting to parent tissue vascular system), revealing storage potential (adventitious roots capable of accumulating carbohydrate reserves particularly in perennial systems), and displaying secondary growth (adventitious roots can undergo secondary thickening developing woody structure in long-lived plants). Adventitious root versus primary root includes= contrasting in origin (adventitious from stems or callus versus primary from radicle or seed), showing different initiation (adventitious requiring dedifferentiation and hormonal induction versus primary from embryonic root pole), exhibiting variable architecture (adventitious roots potentially showing different branching patterns than primary root systems), demonstrating cultivation importance (adventitious roots fundamental to horticultural propagation while primary roots characteristic of seedlings), and revealing genetic control (adventitious rooting capacity showing genetic variation among cultivars affecting propagation success).
Adventitious_Root emphasizes non-root tissue origin and vegetative propagation function and is distinguished from primary and secondary roots by focus on stem or callus origin. Includes Shoot_Derived_Adventitious_Root, Callus_Derived_Adventitious_Root, and Cutting_Adventitious_Root. Critical for understanding vegetative propagation, dedifferentiation, hormonal induction, clonal reproduction, and the roots arising from non-root tissues enabling vegetative propagation fundamental to viticulture.
Adventivwurzel
Cutting root
Non-seed root
Sprossbürtige Wurzel
Stecklingswurzel
Stem root
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Bellini, C., et al. (2014). Adventitious roots and lateral roots= Similarities and differences. Annual Review of Plant Biology, 65, 639-666.
Adventitious Root
Detailed characterization of Adventitious_Root_Formation_Disposition at Cornell University tracked adventitious root development across 12 rootstocks during 21 days of waterlogging in transparent root observation chambers. Daily counts of emerging adventitious roots, root elongation rates, and anatomical analysis of developed roots were conducted. Ethylene production and auxin (IAA) concentration in stem bases were measured to assess hormonal triggers. Adventitious_Root_Formation_Disposition showed striking genotypic variation. 'Riparia Gloire' exhibited rapid, prolific adventitious rooting with first visible root emergence at day 3, peak initiation rate of 2.8 roots per day (days 5-8), total of 18.5 adventitious roots by day 21, and root elongation rate of 3.2 cm/day. Ethylene production in 'Riparia Gloire' stem bases increased 8-fold within 24 hours of waterlogging, with IAA accumulation in the rooting zone reaching 285 ng/g FW (3.5× control). '3309C' (V. riparia × V. rupestris) showed similarly strong response (15.2 roots, 2.6 cm/day elongation). '110 Richter' displayed moderate Adventitious_Root_Formation_Disposition (8.5 roots, first emergence day 6, 1.8 cm/day elongation) with weaker hormonal response (5-fold ethylene, 2× IAA). '140 Ruggeri' exhibited poor adventitious rooting (3.2 roots, emergence delayed to day 10, 0.9 cm/day) associated with only 3-fold ethylene increase and minimal IAA redistribution. Functional assessment revealed that adventitious roots of 'Riparia Gloire' supplied 72% of total vine water uptake by day 21, compensating for original root dysfunction, while '140 Ruggeri' adventitious roots contributed only 18%. These findings demonstrate that Adventitious_Root_Formation_Disposition involves both hormonal responsiveness and developmental capacity for rapid root initiation.
An Adventitious_Root_Formation_Disposition is a Waterlogging_Tolerance_Disposition that inheres in a Vitis plant and is realized through the capacity to rapidly initiate and develop new roots from stem tissues above the waterlogged zone, providing functional root surface in aerated soil or water layers when original roots are compromised by hypoxia. This disposition represents a key escape mechanism from waterlogging stress, enabling plants to establish a new functional root system in oxygen-rich zones while original roots may be dying or dysfunctional in anaerobic soil. Adventitious roots form from pre-existing root primordia in stem tissues or from de novo initiation at the cambium, with the process triggered by the hormonal changes associated with root hypoxia—particularly the accumulation of ethylene (trapped in waterlogged tissues due to slow diffusion in water) and changes in auxin transport and distribution. Adventitious roots that develop into aerated water or saturated surface soil layers are physiologically functional, capable of water and nutrient uptake, and often show enhanced aerenchyma formation facilitating oxygen transport to submerged portions. Adventitious_Root_Formation_Disposition varies among genotypes in the speed of initiation (days from waterlogging onset to visible root emergence), the number and vigor of adventitious roots produced, and the functional capacity of these roots to replace original root function. Genotypes with strong adventitious rooting can maintain vine function through extended waterlogging that would kill genotypes lacking this capacity.
Adventitious_Root_Formation_Disposition is_a Waterlogging_Tolerance_Disposition. Adventitious rooting is triggered by ethylene accumulation (trapped in waterlogged tissues) and auxin redistribution. Root primordia may be pre-formed or initiated de novo. Speed of initiation (days to emergence) and total root number both contribute to flooding survival. Adventitious roots often show enhanced aerenchyma compared to original roots. V. riparia shows particularly strong adventitious rooting, consistent with riparian habitat adaptation.
Adventivwurzel-Bildungs-Disposition
Sprossachsen-Wurzelbildung
flood-induced rooting
hypoxia-induced adventitious roots
stem root formation
Überflutungs-induzierte-Bewurzelung
Steffens B, Rasmussen A (2016) The physiology of adventitious roots, Plant Physiology 170:603-617 Visser EJW et al. (1996) Ethylene accumulation in waterlogged Rumex plants promotes formation of adventitious roots, Journal of Experimental Botany 47:403-410 Vidoz ML et al. (2010) Hormonal interplay during adventitious root formation in flooded tomato plants, Plant Journal 63:551-562
Adventitious Root Formation Disposition
Anatomical investigation of Aerenchyma_Formation_Disposition at INRAE Bordeaux employed microscopic analysis of root cross-sections across 10 rootstocks at 0, 7, 14, and 21 days of waterlogging. Aerenchyma was quantified as percentage of cortex cross-sectional area occupied by air spaces, with parallel measurement of root oxygen concentration using microelectrodes and ethylene evolution (trigger signal). Aerenchyma_Formation_Disposition varied substantially among rootstocks. 'Riparia Gloire' showed rapid, extensive aerenchyma formation, increasing from 5% (constitutive) at day 0 to 25% at day 7 and 48% at day 21, with lysigenous origin confirmed by TUNEL staining for programmed cell death. Root internal oxygen concentration 15 cm from the waterline was maintained at 8.2% (versus atmospheric 21%) in 'Riparia Gloire' versus anoxic (< 0.5%) in low-aerenchyma genotypes. '3309C' displayed similar Aerenchyma_Formation_Disposition (45% by day 21, internal O₂ 7.5%). 'SO4' showed intermediate aerenchyma (32% maximum, internal O₂ 4.8%). '140 Ruggeri' exhibited poor Aerenchyma_Formation_Disposition with only 12% aerenchyma by day 21 and internal O₂ collapsing to 1.2%. Ethylene sensitivity differed among genotypes, with 'Riparia Gloire' root cortex showing aerenchyma initiation at ethylene concentration of 0.5 μL/L while '140 Ruggeri' required 2.5 μL/L for equivalent response. Interestingly, adventitious roots showed 1.5× higher aerenchyma percentage than original roots across all genotypes, suggesting adventitious roots are pre-adapted for oxygen-limited environments. Correlation between aerenchyma percentage at day 14 and waterlogging survival was strong (r = 0.89), confirming the functional importance of Aerenchyma_Formation_Disposition.
An Aerenchyma_Formation_Disposition is a Waterlogging_Tolerance_Disposition that inheres in Vitis root and stem tissues and is realized through the formation of aerenchyma—interconnected air-filled spaces within the cortex that create a low-resistance pathway for oxygen diffusion from aerial plant parts to submerged roots, alleviating hypoxia in waterlogged tissues. This disposition represents the primary anatomical adaptation enabling internal oxygen transport when external oxygen supply through soil is blocked by water. Aerenchyma forms either constitutively (present in well-drained conditions, typical of wetland-adapted species) or inducibly (developing in response to waterlogging, typical of facultatively tolerant species including most Vitis). In grapevine, aerenchyma is predominantly inducible, forming through lysigenous development (programmed cell death creating air spaces) or schizogenous development (cell separation without death) in root cortical tissues. The extent of aerenchyma formation determines the oxygen transport capacity, with effective aerenchyma potentially occupying 20-50% of root cross-sectional area and enabling oxygen diffusion over distances of 20-30 cm in submerged roots. Aerenchyma_Formation_Disposition varies among genotypes in the speed of induction following waterlogging onset, the maximum aerenchyma volume achieved, the extent of interconnection between air spaces (critical for effective transport), and whether aerenchyma forms in adventitious roots as well as original roots.
Aerenchyma_Formation_Disposition is_a Waterlogging_Tolerance_Disposition. Aerenchyma provides internal oxygen transport pathway from shoots to submerged roots. Formation is typically inducible (triggered by hypoxia/ethylene) rather than constitutive in grapevine. Lysigenous aerenchyma involves programmed cell death schizogenous involves cell separation. Effective aerenchyma can occupy 20-50% of cortical cross-section. Oxygen transport distance depends on aerenchyma continuity and cross-sectional area. Adventitious roots often show enhanced aerenchyma compared to original roots.
Aerenchym-Bildungs-Disposition
Interne-Belüftung
Kortikale-Luftraum-Bildung
cortical air space formation
internal aeration
lysigenous aerenchyma development
Evans DE (2004) Aerenchyma formation, New Phytologist 161:35-49 Colmer TD (2003) Long-distance transport of gases in plants: a perspective on internal aeration and radial oxygen loss from roots, Plant Cell and Environment 26:17-36 Jackson MB, Armstrong W (1999) Formation of aerenchyma and the processes of plant ventilation in relation to soil flooding and submergence, Plant Biology 1:274-287
Aerenchyma Formation Disposition
The AFLP marker data generated for the first comprehensive genetic map of grapevine (Doligez et al. 2002, Syrah × Grenache cross) employed the standard EcoRI/MseI restriction enzyme combination with 64 selective primer pairs (three selective nucleotides per primer), scoring 370 polymorphic AFLP fragments ranging from 50 to 500 bp on polyacrylamide sequencing gels with silver staining — the resulting dominant markers (scored as present/absent, unable to distinguish heterozygotes from homozygotes for the band-present class) provided the initial framework of the 19-linkage-group genetic map at an average spacing of 4.5 cM, subsequently densified with codominant SSR markers that enabled integration with physical genome maps and cross-population map comparison.
A genetic marker data record for amplified fragment length polymorphism (AFLP) markers generated in Vitis through the restriction-ligation-amplification protocol, documenting the enzyme combination, selective primer sequences, fragment sizes scored, and presence/absence polymorphism patterns that provided medium-density genome-wide marker coverage for genetic mapping and diversity analysis prior to the widespread adoption of SSR and SNP marker systems.
AFLP_Marker_Data documents a marker technology that played a transitional role in grapevine genetics between the low-throughput RFLP/RAPD era and the current SSR/SNP era. AFLPs were introduced by Vos et al. (1995) as a PCR-based technique generating 50-100 polymorphic markers per primer combination without requiring prior sequence knowledge — a decisive advantage for species like grapevine where limited genomic sequence was available before the 2007 reference genome publication: AFLP markers contributed substantially to early grapevine genetic maps (Doligez et al. 2002, Adam-Blondon et al. 2004, Grando et al. 2003) and diversity studies (Cervera et al. 1998, Fanizza et al. 2005) during the 1998-2008 period. The principal limitations of AFLPs — dominant inheritance (preventing heterozygote detection), technical complexity requiring skilled gel electrophoresis, poor cross-study reproducibility, and inability to determine genomic position without map integration — led to their progressive replacement by codominant SSR and SNP markers as these became available in sufficient numbers. AFLP data remain historically valuable in grapevine genetics as the foundation of early genetic maps and the first genome-wide diversity assessments, but new AFLP genotyping has essentially ceased in favor of SNP-based approaches.
AFLP fingerprint data
AFLP marker specification
AFLP-Fingerabdruck-Daten
AFLP-Marker-Daten
amplified fragment length polymorphism record
amplifizierte Fragmentlängen-Polymorphismus-Daten
Vos P et al. (1995) AFLP: a new technique for DNA fingerprinting, Nucleic Acids Research 23: 4407-4414 Doligez A et al. (2002) An integrated SSR map of grapevine based on five mapping populations, Theoretical and Applied Genetics 105: 161-172 Cervera MT et al. (1998) Application of AFLPs to the characterization of grapevine Vitis vinifera L. genetic resources, Genetics Selection Evolution 30: S237-S249
AFLP Marker Data
Aftertaste quality assessment in Barolo comparing traditionally fermented (extended maceration) versus modern style (shorter maceration, new oak) reveals traditional wines with aftertaste evolving through cherry fruit to tar to rose petal over 30 seconds, rated 8.5 for aftertaste quality, versus modern style showing initial fruit sweetness fading to oak vanilla then harsh tannin at 15 seconds, rated 6.2, demonstrating that aftertaste character evolution can differ dramatically even at similar total persistence, with the harmonious traditional evolution preferred despite equivalent duration.
Aftertaste quality is a taste persistence quality that represents the qualitative character and hedonic value of taste sensations experienced after swallowing, characterizing the nature of lingering flavors independent of their duration and determining whether the finish contributes positively or negatively to overall wine quality perception. Within the BFO framework, this quality specifically addresses the qualitative dimension of post-swallow taste, distinguishing the character and pleasantness of aftertaste from the quantitative duration measure of finish length while recognizing that both dimensions contribute to finish quality assessment. The ontological significance lies in aftertaste quality determining the final impression that shapes wine memory and repurchase intention, where pleasant, evolving aftertaste enhances perceived quality regardless of duration while unpleasant aftertaste (harsh bitterness, metallic notes, volatile acidity sting) diminishes quality even with extended persistence. The perceptual evolution of aftertaste involves differential fading of taste components at different rates, creating a temporal sequence where initial sweetness may give way to emerging bitterness, acidity may persist after fruit fades, or mineral notes may emerge as fruit recedes, with the complexity and harmony of this evolution determining aftertaste quality. The compositional basis for aftertaste character includes the specific compounds persisting longest (typically phenolics and non-volatile acids), their relative concentrations affecting which notes dominate the late finish, and potential emergence of retronasal aroma as volatile compounds warm in the mouth contributing flavor complexity to the aftertaste period. The descriptive vocabulary for aftertaste includes positive terms (clean, lingering fruit, mineral, integrated tannin) and negative terms (bitter, harsh, metallic, hot, short), with the balance of positive to negative character determining overall aftertaste quality. Understanding aftertaste quality enables comprehensive finish assessment beyond duration, identification of specific finish quality issues requiring winemaking attention, and quality description capturing the experiential character of wine's final impression.
Aftertaste_Quality addresses the qualitative character of post-swallow taste, complementing Finish_Length_Quality which provides quantitative duration. This quality captures the hedonic value and descriptive character of the finish that duration measurement alone cannot convey. The class supports comprehensive finish evaluation incorporating both how long taste persists and what that persistent taste is like.
Abgangscharakter
Aromapersistenz
Nachgeschmack
finish character
lingering flavor
nachhallender Geschmack
post-swallow taste
terminal taste
Peynaud E, Blouin J (1996) The Taste of Wine: The Art and Science of Wine Appreciation, 2nd Ed. John Wiley & Sons, New York, pp. 92-118 Jackson RS (2014) Wine Science: Principles and Applications, 4th Ed. Academic Press, San Diego, pp. 831-887
Aftertaste Quality
Aging potential for Riesling Trockenbeerenauslese from the Rheingau (2022 vintage) projects an extraordinary total lifespan of 50-100+ years, with an optimal drinking window of 15-60 years post-vintage, currently in the youthful-primary stage — the preservative combination of extreme residual sugar (220 g/L), very high acidity (TA 10.5 g/L, pH 2.95), and botrytis-derived antimicrobial compounds creating conditions for essentially indefinite cellaring, with documented TBA wines from the 19th century remaining vibrant and complex. Aging potential for Grüner Veltliner Federspiel from the Wachau (2024 vintage) projects a total lifespan of 3-5 years with an optimal window of 1-3 years, currently at peak — the moderate concentration, light-medium body, and absence of phenolic structure indicate a wine designed for early consumption at maximum freshness. Aging potential for Blaufränkisch Reserve from Leithaberg DAC (2021 vintage) estimates 15-25 year total lifespan with optimal window of 5-15 years, currently developing — the firm tannin structure, good acidity (TA 5.5 g/L, pH 3.35), and moderate alcohol (13.5%) provide the structural prerequisites for extended cellaring, with the tannin quality suggesting a favorable evolution trajectory.
Aging potential is an overall assessment entity that records the projected developmental trajectory and cellaring capacity of a wine, encompassing the estimated total lifespan (the period from bottling to the point beyond which further aging yields no improvement or active deterioration), the optimal drinking window (the period during which the wine is expected to offer its most complete and rewarding sensory experience), the current developmental stage (youthful/primary, developing/secondary, mature/at peak, declining/past peak), and the structural indicators supporting the longevity assessment (acid level, tannin concentration and quality, extract, alcohol, and the balance among these preservative factors), providing the temporal quality dimension that guides cellar management decisions, release timing, and consumer purchasing strategy.
Aging potential assessment represents the most speculative and expertise-dependent component of wine evaluation, requiring the taster to project from the wine's current sensory state to its anticipated future development — a prediction that depends on understanding the chemical kinetics of wine aging, the structural indicators that correlate with longevity, and the extensive experiential database of having tracked wines through their developmental arcs over years and decades. The chemical basis of wine aging capacity rests on several preservative factors working in concert — acidity provides the low-pH environment that slows oxidation and microbial degradation, tannin-phenolic compounds serve as sacrificial antioxidants (scavenging oxygen before it can damage fruit aromatics) and undergo progressive polymerization that transforms youthful astringency into mature textural complexity, residual sugar provides both osmotic microbial inhibition and a reservoir for slow Maillard-type reactions generating complexity, and sulfur dioxide provides direct antimicrobial and antioxidant protection. The interaction between these factors is synergistic — a wine with high acidity but low tannin (young Chablis) ages differently from a wine with high tannin but modest acidity (warm-climate Cabernet), with the most age-worthy wines possessing strong levels of multiple preservative factors simultaneously. The optimal drinking window concept acknowledges that wine quality follows a developmental arc — improvement during early bottle aging as components integrate, a plateau of peak quality, followed by eventual decline as fruit aromatics fade, tannins over-polymerize and precipitate, and oxidative aging products accumulate — with the duration and timing of each phase varying enormously by wine type. The storage conditions fundamentally modify aging potential — temperature (optimal 10-15°C, with each 10°C increase approximately doubling the rate of chemical aging), humidity (65-80% to prevent cork desiccation), light exclusion, and vibration absence are all prerequisites for achieving a wine's theoretical aging potential. The closure type (natural cork, synthetic closure, screwcap, glass stopper) significantly influences aging trajectory through differential oxygen transmission rates, with screwcaps providing the most reductive aging environment (minimal oxygen ingress) and natural cork providing gradual micro-oxygenation that facilitates oxidative complexity development but introduces the risk of cork taint (2,4,6-trichloroanisole contamination).
Alterungspotenzial
Lagerfähigkeitsbeurteilung
Lagerpotenzial
Reifepotenzial
Trinkreife
ageability assessment
cellaring potential
development trajectory
drinking window
longevity assessment
Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 547-645 Waterhouse AL et al. (2016) Understanding Wine Chemistry, Wiley, pp.: 275-340 Gladstones J (2011) Wine, Terroir and Climate Change, Wakefield Press, pp.: 200-240 Skouroumounis GK et al. (2005) The influence of ascorbic acid on the composition, colour and flavour properties of a Riesling and a wooded Chardonnay wine during five years' storage. Australian Journal of Grape and Wine Research 11(3): 355-368
Aging Potential
Aging protocol for Barolo DOCG from Serralunga d'Alba specifies 36 months minimum aging (regulatory requirement), implemented as 24 months in Slavonian oak botti (25-50 hL, neutral flavor contribution, slow micro-oxygenation through stave porosity), followed by 12 months bottle aging before release, with quarterly racking during the first year (reducing to biannual in the second year), cellar maintained at 13-15°C and 75-85% relative humidity, and no new oak contact — the protocol reflecting the traditional Barolo philosophy of extended oxidative maturation in large neutral vessels that allows Nebbiolo's formidable tannin structure to polymerize and soften without masking varietal character with oak-derived flavors. Aging protocol for Grüner Veltliner Reserve from Kamptal documents 12 months aging on full lees in a combination of stainless steel (60%, preserving primary fruit and freshness), 2500 L Austrian oak Stockinger cask (30%, providing subtle textural enrichment through micro-oxygenation without oak flavor contribution), and 500 L used French oak pièce (10%, adding gentle spice complexity), with monthly bâtonnage in the cask portions and quarterly bâtonnage in steel, assembled in February and rested 2 months in tank before bottling — balancing freshness preservation with textural complexity.
Aging protocol is a winemaking protocol entity that documents the parameters, vessel selection, duration, and environmental conditions governing the maturation of wine between the completion of fermentation and final bottling, encompassing the aging vessel type (barrel, cask, tank, amphora, concrete egg), vessel material and size (French or American oak, acacia, chestnut 225 L barrique, 300 L hogshead, 500 L puncheon, 2500+ L foudre), proportion of new versus used cooperage, aging duration, cellar temperature and humidity conditions, lees management (sur lie aging, bâtonnage frequency), racking schedule (frequency, method, oxidative versus reductive), and micro-oxygenation parameters where applied, providing the maturation management framework that governs the development of wine complexity, structural integration, and the transformation from young wine to bottle-ready product.
Aging protocol documentation captures one of the most stylistically consequential and philosophically debated aspects of winemaking, as the élevage decisions — particularly regarding oak type, size, age, and duration — fundamentally shape the wine's aromatic profile, textural evolution, and aging trajectory. The chemical processes during aging include slow oxidation through vessel permeability (oak staves permit 2-5 mg O₂/L/month in new 225 L barrels, substantially less in larger or older vessels), extraction of oak-derived compounds (vanillin, eugenol, guaiacol, oak lactones, ellagitannins from new oak), polymerization and stabilization of tannin-anthocyanin complexes, and ester equilibration between formation and hydrolysis. The vessel size decision creates a fundamental stylistic axis — small barrels (225 L barrique, 228 L pièce, 300 L hogshead) maximize oak surface-to-wine ratio and oxygen exposure, producing wines with more pronounced oak character and faster evolution, while large vessels (500-5000+ L foudres, botti, Stockinger casks) minimize oak extraction and slow oxidation, preserving primary fruit character and varietal expression. The new-to-used oak proportion represents the primary modulator of oak flavor intensity — new oak contributes maximum vanillin (sweet, vanilla), guaiacol (smoky), eugenol (clove), and oak lactone (coconut, woody), while successive uses progressively deplete extractable compounds until barrels become "neutral" after 3-5 fills. The emergence of alternative aging vessels — concrete eggs (providing gentle micro-oxygenation without oak flavor), Georgian qvevri (buried clay amphorae enabling temperature-stable fermentation and aging), and ceramic jars — reflects the growing interest in non-oak maturation pathways that allow terroir expression without cooperage overlay. Lees management during aging provides another important dimension — extended contact with fine lees (sur lie aging) releases mannoproteins that enhance mouthfeel, reduce astringency, and improve protein and tartrate stability, while bâtonnage (lees stirring) accelerates these effects but increases oxidative exposure and the risk of reductive sulfur compound development if poorly managed.
Ausbauprotokoll
Fassausbauprotokoll
Reifeprotokoll
Reifungsführung
aging regime
cellar aging specification
maturation protocol
wine maturation parameters
Élevage-Protokoll
élevage protocol
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed, Wiley, pp.: 203-260 Waterhouse AL et al. (2016) Understanding Wine Chemistry, Wiley, pp.: 275-310 Chatonnet P et al. (1999) Influence of the origin and seasoning of oak on the composition and quality of wines aged in new barrels. American Journal of Enology and Viticulture 50(4): 447-462 Boulton RB et al. (1999) Principles and Practices of Winemaking, Springer, pp.: 382-425
Aging Protocol
Chardonnay showing aging senescent feeder roots in September, minirhizotron imaging revealing feeder roots with tan coloration distinct from white active or brown dying roots, roots showing slight shriveling and partial root hair loss, FDA staining showing variable results with 35% of aging roots positive, isotope pulse-labeling showing aging roots contributing 12% of uptake versus 76% from clearly active roots, microscopic examination revealing partial cytoplasmic degradation with some organelles degraded others functional, aging senescent feeder roots representing transition state between full function and complete loss
Aging senescent feeder root is senescent feeder root in early to middle senescence showing beginning functional decline but not yet completely non-functional, typically 4-10 months post-initiation, characterized by visible aging indicators including tan coloration intermediate between white young and brown dying roots, some turgor loss creating slight shriveling though not complete collapse, partial root hair loss with many hairs degraded but some remaining, declining but not absent absorption capacity, and representing transition state between full function and complete functional loss, with aging senescent feeder roots showing partial cytoplasmic degradation with some organelles degrading while others maintaining limited function, progressive membrane dysfunction reducing but not eliminating selective permeability, declining metabolic rates with respiration and transport continuing at reduced levels, and demonstrated residual uptake capacity contributing minor fraction to total vine absorption, and demonstrating that aging phase represents gradual functional decline over weeks distinguishing from rapid terminal dying phase, and showing potential for population assessment challenges as aging roots appear partially functional potentially confounding estimates of active versus non-functional feeder root populations requiring careful viability and functional testing. Aging senescent feeder root visual indicators includes= showing tan coloration (aging roots intermediate between white active and brown dying creating tan or light brown appearance), exhibiting partial shriveling (beginning turgor loss creating slight wrinkled appearance though not completely collapsed), demonstrating incomplete root hair loss (many root hairs degraded but significant proportion remaining creating patchy coverage), revealing surface changes (root surface showing beginning irregularities, texture changes, or discolorations), and displaying diameter reduction (aging roots showing slightly reduced diameter from turgor loss and tissue shrinkage). Aging senescent feeder root cellular degradation includes= showing partial organelle breakdown (some mitochondria, plastids, and other organelles degrading while others maintaining limited structure and function), exhibiting beginning vacuole changes (vacuoles enlarging or showing altered contents though not complete degradation), demonstrating progressive autophagy (self-digestion pathways active consuming cellular components though process incomplete), revealing selective protein degradation (some proteins degrading while others maintained creating altered protein composition), and displaying membrane changes (plasma membrane and organellar membranes showing reduced integrity though not complete failure). Aging senescent feeder root functional capacity includes= showing reduced absorption (water and nutrient uptake significantly declined but not completely ceased), exhibiting decreased metabolic rate (respiration and other metabolic processes continuing at 20-50% of peak active levels), demonstrating partial turgor maintenance (cells maintaining some turgor though reduced from peak creating intermediate turgidity), revealing limited responsiveness (aging roots showing diminished capacity to respond to environmental or physiological signals), and displaying declining contribution (aging roots contributing minor fraction perhaps 5-20% of active root uptake levels to total vine absorption). Aging senescent feeder root population assessment includes= requiring careful distinction (aging roots potentially counted as active in simple viability tests despite limited function), showing intermediate viability (vital staining potentially showing variable results with some aging roots staining positively others negatively), exhibiting ambiguous status (aging roots intermediate between clearly active and clearly non-functional creating assessment challenges), demonstrating importance for accuracy (distinguishing aging from fully active roots important for accurate functional root population estimates), and revealing methodological needs (multiple assessment criteria necessary including viability, uptake capacity, morphology for accurate aging root identification). Aging senescent feeder root in turnover dynamics includes= representing transition state (aging roots in process of transitioning from active functional to non-functional dead state), showing variable duration (aging phase potentially lasting days to weeks depending on environmental conditions and root characteristics), exhibiting progressive decline (aging roots showing continuous functional deterioration progressing toward complete loss), demonstrating population proportion (aging roots potentially comprising 10-30% of total feeder root population at any time), and revealing flux (aging roots representing roots flowing from active pool toward dead pool in continuous turnover). Aging senescent feeder root environmental effects includes= showing stress acceleration (drought, heat, or other stresses accelerating progression from aging to dying state), exhibiting seasonal patterns (aging potentially more prevalent late summer when cumulative stress highest or fall as roots age), demonstrating soil effects (poor soil conditions potentially increasing proportion of aging versus fully active roots), revealing management impacts (irrigation and fertilization potentially affecting rate of aging progression), and displaying temperature sensitivity (aging progression potentially temperature-dependent with faster rates at higher temperatures). Aging senescent feeder root mycorrhizal status includes= showing declining colonization (arbuscular mycorrhizal structures in aging roots potentially degrading), exhibiting arbuscule senescence (arbuscules in aging roots senescing losing exchange function), demonstrating reduced benefit (mycorrhizal contribution to uptake declining as both root and fungal function deteriorate), revealing fungal response (arbuscular mycorrhizal fungi potentially reducing allocation to aging roots focusing on younger roots), and displaying symbiosis termination (mycorrhizal association ending as root ages completing senescence). Aging senescent feeder root research implications includes= requiring functional assessment (research must distinguish aging from fully active roots to accurately estimate absorption capacity), showing methodological challenges (standard viability tests potentially misclassifying aging roots as functional), exhibiting interpretation importance (including aging roots in functional root estimates potentially overestimating actual absorption capacity), demonstrating model needs (feeder root turnover models should account for aging transition state not just active versus dead), and revealing management relevance (understanding aging root dynamics important for interpreting vine responses to practices).
Aging_Senescent_Feeder_Root emphasizes early-middle senescence and partial function and is distinguished from fully active or completely dying roots by focus on intermediate declining state. Critical for understanding transition phase, partial degradation, reduced function, population assessment challenges, and the aging roots in early-middle senescence showing declining but not absent function creating intermediate state in feeder root turnover.
Alternde seneszente Feinwurzel
Declining feeder root
Early senescent feeder root
Früh-seneszente Saugwurzel
Partially functional aging root
Partiell-funktionelle alternde Wurzel
Eissenstat, D. M., et al. (2000). Building roots in a changing environment= Implications for root longevity. New Phytologist, 147(1), 33-42.
Aging Senescent Feeder Root
Investigation of Agrobacterium_Transformation_Resistance_Disposition in Vitis was conducted indirectly through transgenic transformation efficiency studies at INRA Colmar, which revealed significant genotypic variation potentially reflecting natural resistance mechanisms. Embryogenic callus cultures from 15 cultivars were co-cultivated with A. tumefaciens strain EHA105 carrying a GUS reporter construct under standardized conditions (48h co-cultivation, 100 μM acetosyringone). Transformation efficiency, measured as GUS-positive regenerants per gram callus, varied 12-fold among cultivars. 'Chardonnay' showed high transformability (45 transformants/g), while '41B' rootstock showed very low efficiency (3.8 transformants/g) despite equivalent callus quality and regeneration capacity. Molecular analysis revealed that '41B' exhibited 70% lower transient GUS expression at 5 days post-co-cultivation, suggesting resistance at the T-DNA transfer or early expression stage rather than regeneration. Unfortunately, this apparent Agrobacterium_Transformation_Resistance_Disposition in '41B' did not correlate with crown gall resistance in planta, with '41B' developing equivalent galls to highly transformable 'Chardonnay' following A. vitis wound inoculation. This disconnect indicates that in vitro transformation resistance may involve different mechanisms than whole-plant crown gall resistance, or that tumorigenic A. vitis strains overcome barriers that limit laboratory A. tumefaciens strains.
An Agrobacterium_Transformation_Resistance_Disposition is a Crown_Gall_Resistance_Disposition that inheres in a Vitis plant and is realized through cellular mechanisms that specifically interfere with Agrobacterium-mediated genetic transformation, preventing T-DNA transfer from the bacterial Ti plasmid into plant cell nuclei or blocking T-DNA integration into plant chromosomal DNA. This disposition represents the molecular level of crown gall resistance operating after bacterial colonization of wound sites has occurred, targeting the unique biological process by which Agrobacterium reprograms host cells. The T-DNA transfer process involves bacterial attachment to plant cells, induction of virulence genes by wound-released phenolic compounds, processing and export of T-DNA through a type IV secretion system, T-DNA transit across plant cell membranes and cytoplasm, nuclear import, and chromosomal integration. Potential resistance mechanisms disrupting this cascade include modification of VirA/VirG two-component sensing to prevent virulence gene induction, interference with bacterial attachment or secretion system function, degradation of T-DNA or associated proteins in the cytoplasm, blockage of nuclear import machinery recognition of VirD2/VirE2 nuclear localization signals, and inhibition of T-DNA integration by host DNA repair or chromatin factors. While natural Agrobacterium_Transformation_Resistance_Disposition has not been clearly demonstrated in Vitis, variation in transformation efficiency among genotypes during transgenic research suggests that such resistance may exist at low levels. Paradoxically, the same mechanisms that might confer crown gall resistance would also impede beneficial transgenic applications in grapevine improvement.
Agrobacterium_Transformation_Resistance_Disposition is_a Crown_Gall_Resistance_Disposition. This class represents molecular-level resistance to T-DNA transfer and integration, distinct from Wound_Healing_Efficiency_Disposition which operates at the tissue level. Natural transformation resistance has not been conclusively demonstrated to provide crown gall protection in Vitis, though genotypic variation in transformation efficiency suggests underlying resistance factors. Research into transformation resistance is complicated by the use of different Agrobacterium species/strains in biotechnology (typically A. tumefaciens) versus natural crown gall disease (A. vitis). Transformation resistance would also impede beneficial transgenic applications.
Agrobacterium recalcitrance
Agrobakterien-Transformations-Resistenz-Disposition
T-DNA transfer resistance
T-DNA-Transfer-Resistenz
Transformations-Immunität
transformation immunity
Gelvin SB (2003) Agrobacterium-mediated plant transformation: the biology behind the "gene-jockeying" tool, Microbiology and Molecular Biology Reviews 67:16-37 Ziemienowicz A (2014) Agrobacterium-mediated plant transformation: factors, applications and recent advances, Biocatalysis and Agricultural Biotechnology 3:95-102 Pitzschke A, Hirt H (2010) New insights into an old story: Agrobacterium-induced tumour formation in plants by plant transformation, EMBO Journal 29:1021-1032
Agrobacterium Transformation Resistance Disposition
Assyrtiko vines at Domaine de la Garance in the Languedoc bearing an agroforestry component role are planted in alternating rows with olive trees (Olea europaea) and almond trees (Prunus dulcis) at 12 m inter-row spacing, creating a silvo-viticultural system where trees provide 20-30% afternoon shade reducing vine heat stress by 2-4°C during July-August heatwaves, tree root systems access water at 3-6 m depth below vine root zone reducing direct competition, and the combined system supports 40-60% higher arthropod diversity than monoculture vine plots while sequestering an additional 2-4 t CO₂/ha/year from the tree component.
An ecological role that inheres in a grapevine cultivated as a component of an agroforestry system that intentionally combines grapevine production with trees or woody perennials on the same land management unit, creating a multi-strata cropping system that generates complementary ecological interactions between vine, tree, and associated biodiversity.
Agroforestry_Component_Role captures the emerging integration of viticulture with tree-based systems, historically practiced as traditional vitiforestry across Mediterranean Europe (notably the coltura promiscua system of central Italy where vines were trained between olive, fruit, and timber trees until the mid-20th century). Contemporary vitiforestry is experiencing a revival driven by climate adaptation research demonstrating that strategic tree integration can buffer vine thermal extremes: however, the system involves complex management tradeoffs including light competition, altered disease microclimate, and mechanization constraints that require careful spatial design and species selection.
Agroforst-Komponenten-Rolle
Baum-Rebe-Integrations-Rolle
Vitiforstrolle
agroforestry vine role
tree-vine integration role
vitiforestry role
Ferreiro-Domínguez N et al. (2016) Grape production and must quality in a grapevine-pasture agroforestry system, Agroforestry Systems 90: 631-643 Grimaldi J et al. (2017) Agroforestry with viticulture, Agriculture Ecosystems and Environment 239: 285-293 Dupraz C and Liagre F (2008) Agroforesterie: des arbres et des cultures, France Agricole pp: 312-335
Agroforestry Component Role
Agronomic performance data for Chardonnay clone 96 (Dijon) at Sonoma Coast documents moderate vigor (pruning weight 0.6-0.9 kg/m cordon on 101-14 Mgt rootstock), high bud fruitfulness (1.8-2.1 clusters per shoot), early-mid ripening phenology (harvest 8-12 days before clone 76 at equivalent Brix targets), and a Ravaz Index of 5-7, indicating well-balanced vegetative-to-reproductive growth partitioning ideal for cool-climate premium production. Agronomic performance data for Sankt Laurent clone SL-4 at Klosterneuburg records vigorous growth (pruning weight 1.2-1.6 kg/m on SO4 rootstock), moderate fertility (1.4-1.6 clusters per shoot), early budburst (7-10 days before Blaufränkisch, increasing late frost risk), and compact ripening window requiring harvest scheduling within a 10-14 day optimum period before rapid acidity decline.
Agronomic performance data is a performance data entity that documents the vegetative growth characteristics, reproductive capacity, and developmental timing of a clonal line under field evaluation conditions, encompassing vigor parameters (shoot growth rate, pruning weight, canopy dimensions), fertility metrics (bud fruitfulness, cluster initiation), and ripening phenology (phenological stage timing, maturation rate, harvest date), providing the viticultural management characterization essential for matching clone selection to site conditions, training system requirements, and production objectives.
Agronomic performance data bridges the gap between clone evaluation trial results and practical vineyard management, as vigor, fertility, and phenological timing collectively determine the training system compatibility, spacing requirements, canopy management intensity, and harvest logistics associated with each clone. The interaction between agronomic performance and site conditions is particularly strong — a vigorous clone may perform optimally on shallow, low-fertility soils where its growth potential is naturally constrained, while the same clone on deep, fertile alluvial soils may produce excessive vegetation requiring intensive and costly canopy management. The Ravaz Index (yield/pruning weight) provides an integrative measure of vegetative-reproductive balance, with values of 5-10 generally indicating optimal partitioning for quality wine production — clones consistently outside this range may require management intervention (crop thinning for high-Ravaz clones, shoot thinning or vigor reduction for low-Ravaz clones) to achieve balance. Phenological timing data within agronomic performance is increasingly important under climate change, as early-ripening clones may advance into hotter harvest periods while late-ripening clones maintain the cooler ripening conditions associated with balanced wine composition. Clone-by-rootstock interactions for agronomic parameters are often substantial, particularly for vigor and phenology, making rootstock specification essential for agronomic performance data interpretation.
Anbaueignungsdaten
Rebverhaltensdaten
Wuchsleistungsdaten
agronomic evaluation data
agronomische Leistungsdaten
cultural performance data
growth performance record
vine behavior data
viticultural performance data
weinbauliche Leistungsdaten
Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 195-235 Smart RE, Robinson MD (1991) Sunlight into Wine, Winetitles, pp.: 30-55 Rühl EH et al. (2004) Grapevine clonal selection, evaluation, and management. Acta Horticulturae 652: 29-34
Agronomic Performance Data
Air circulation quality evaluation in a Botrytis-prone Champagne Chardonnay vineyard compared wetness duration across canopy density treatments during a rainy September period. Dense unthinned canopies with 8% gaps maintained fruit surface wetness for mean 14.2 hours following 10mm precipitation events, providing optimal conditions for Botrytis infection. Moderately thinned canopies with 25% gaps showed improved air circulation quality with wetness duration reduced to 8.5 hours. Aggressively managed canopies with 40% gaps achieved superior air circulation quality with wetness duration of only 5.1 hours, falling below the 6-hour threshold typically required for Botrytis germination and infection. Corresponding Botrytis incidence at harvest was 42%, 18%, and 6% respectively, demonstrating the practical disease management value of air circulation quality optimization.
Air circulation quality is a canopy porosity quality that inheres in a grapevine canopy system and represents the capacity for atmospheric air movement through the foliar mass that enables gas exchange, heat dissipation, humidity reduction, and rapid drying of wetted surfaces as determined by canopy porosity distribution, row orientation relative to prevailing winds, and the aerodynamic characteristics of the training system architecture. This quality fundamentally influences disease development potential by controlling the duration of surface wetness that enables fungal spore germination and infection, while also affecting leaf and fruit temperatures through convective heat transfer. The ontological significance of air circulation quality lies in its role as a primary determinant of disease pressure independent of fungicide applications, with well-ventilated canopies reducing reliance on chemical intervention through physical limitation of pathogen infection periods. The physics of air flow through vegetation canopies involves complex interactions between large-scale wind patterns and the drag and turbulence created by plant structures, with dense canopies generating boundary layers of nearly stagnant air in the canopy interior that resist penetration by external wind currents. Practical consequences of poor air circulation quality include extended leaf and fruit wetness duration following precipitation, dew, or overhead irrigation, elevated humidity in the fruit zone favoring fungal growth, reduced transpirational cooling of leaves during heat events, and accumulation of ethylene and other volatiles that may affect fruit development. The relationship between canopy density and air circulation follows non-linear patterns wherein moderate increases in porosity produce substantial improvements in ventilation up to thresholds above which further openness provides diminishing returns, with research suggesting approximately 20-30% gap frequency as minimally adequate for disease-relevant air circulation improvements. Measurement of air circulation quality presents practical challenges, with approaches including direct anemometry in the canopy interior, tracer gas dispersion studies, wetness duration monitoring using electronic leaf wetness sensors, and humidity gradient characterization between canopy interior and ambient conditions. Row orientation significantly influences air circulation quality, with rows aligned parallel to prevailing winds permitting better cross-row ventilation than perpendicular orientations, though this consideration must be balanced against slope, sun exposure, and operational efficiency factors in vineyard design.
Air_Circulation_Quality is_a Canopy_Porosity_Quality, representing the atmospheric ventilation dimension of canopy openness. This class addresses air movement, humidity, and drying dynamics rather than light transmission, though both derive from canopy porosity characteristics. The disease management implications of air circulation quality are particularly significant, with wetness duration serving as key measurable outcome. Distinguished from Light_Penetration_Quality and Spray_Penetration_Quality as parallel consequences of canopy porosity.
Belüftungsqualität
Fruchtzonenbelüftung
Kronendurchlüftung
Luftzirkulation
air movement quality
canopy airflow
canopy ventilation
fruit zone ventilation
English JT et al. (1989) Influence of grapevine canopy management on evaporative potential in the fruit zone. American Journal of Enology and Viticulture 40:123-129 Percival DC et al. (1994) Canopy microclimate and Botrytis bunch rot development in Riesling grapevines. Vitis 33:17-22 Austin CN, Wilcox WF (2011) Effects of fruit-zone leaf removal, training systems, and irrigation on the development of grapevine powdery mildew. American Journal of Enology and Viticulture 62:193-198
Air Circulation Quality
Alcohol content for Riesling Kabinett from the Mosel (2024 vintage) measures 8.5% vol. by distillation-densimetry, well below the 12.5% vol. upper limit traditionally associated with Kabinett-style wines and reflecting the partial fermentation strategy that preserves 45 g/L residual sugar balanced against 8.0 g/L total acidity — the low alcohol contributing to the wine's characteristic lightness and delicacy. Alcohol content for Shiraz from the Barossa Valley (2022 vintage) records 14.8% vol., near the upper boundary of typical full-bodied red wine range, with the high alcohol contributing to the wine's viscous mouthfeel, glycerol-enhanced body, and warm finish. Alcohol content for Blaufränkisch from Mittelburgenland DAC (2023 vintage) documents 13.5% vol., within the moderate range appropriate for the DAC style that emphasizes fruit purity and drinkability over extraction and power.
Alcohol content is a wine analysis data entity that records the concentration of ethanol in finished wine, determined by distillation followed by pycnometry or digital density measurement (OIV reference method), near-infrared spectroscopy, ebulliometry, or FTIR estimation, expressed in percent by volume (% vol.) at 20°C, and representing the primary compositional parameter that defines wine body, determines regulatory category and tax classification, governs labeling requirements and permitted tolerance, and establishes the legal alcohol declaration that accompanies the wine through commercial channels to the consumer.
Alcohol content determination is the most legally consequential wine analysis measurement, as it directly governs excise taxation (which in most jurisdictions is calculated on alcohol volume), labeling compliance (EU regulations permit ±0.5% vol. tolerance from the declared value, while US regulations permit ±1.0% for wines 7-14% vol. and ±1.5% above 14%), and category classification (EU wine regulations define wine as a minimum 8.5% vol. product from fermented grape must, with specific categories requiring defined alcohol ranges). The OIV reference method (distillation of wine to separate ethanol from non-volatile constituents, followed by density measurement of the distillate at 20°C using a pycnometer or oscillating U-tube densitometer) provides accuracy of ±0.05% vol., sufficient for regulatory and commercial purposes. The relationship between grape sugar and wine alcohol follows the approximate conversion of 16.5-17.5 g/L sugar per 1% vol. ethanol, modified by fermentation efficiency, glycerol production, and residual sugar — actual alcohol is always lower than the potential alcohol calculated from must sugar content. The sensory contribution of alcohol extends beyond the warming perception on the palate — ethanol enhances body through viscosity effects, modulates aroma volatility and perception intensity, and interacts with tannin and acid perception in complex ways (moderate alcohol softens tannin perception while high alcohol can amplify bitterness). The global trend toward both higher and lower alcohol extremes — climate change driving sugar accumulation toward 15-16% potential alcohol in warm regions, while market demand for lower-alcohol wines grows in health-conscious markets — creates increasing tension between vineyard reality and target alcohol specification, driving interest in technological interventions (early harvest, dealcoholization, yeast strain selection for lower alcohol yield) to manage wine alcohol content.
Alkoholgehalt
Alkoholkonzentration
Alkoholstärke
Ethanolgehalt
alcohol concentration
alcohol level
alcoholic strength
ethanol content
vorhandener Alkohol
wine alcohol
OIV (2021) Compendium of International Methods of Analysis, Method OIV-MA-AS312-01A (alcoholic strength by volume) European Parliament (2013) Regulation (EU) No 1308/2013, Annex VII Part II (wine definitions and categories) Waterhouse AL et al. (2016) Understanding Wine Chemistry, Wiley, pp.: 35-55
Alcohol Content
In Grüner Veltliner berries in Kamptal, Austria during BBCH 79-89, aldehyde synthesis shows characteristic decline with hexanal decreasing from 125 µg/L at veraison to 18 µg/L at harvest over 48 days, trans-2-hexenal declining from 185 to 32 µg/L, and cis-3-hexenal from 95 to 12 µg/L, while corresponding C6 alcohols increase through aldehyde reduction, with trained panel green-herbaceous ratings declining from 6.8 to 2.2 on 10-point scale paralleling the analytical aldehyde reduction under cool conditions averaging 17°C.
Aldehyde synthesis describes the generation of volatile aldehyde compounds in Vitis vinifera berries during ripening, wherein lipoxygenase-mediated oxidation of polyunsaturated fatty acids followed by hydroperoxide lyase cleavage produces the C6 aldehydes hexanal and hexenal isomers that contribute characteristic green, grassy, and herbaceous aromatic notes to unripe and early-ripening berries, with progressive reduction during maturation establishing the declining green character associated with advancing grape ripeness. This process involves the sequential action of lipoxygenase enzymes oxidizing linoleic and linolenic acids at specific positions to form fatty acid hydroperoxides, followed by hydroperoxide lyase catalyzed cleavage producing C6 aldehyde fragments including hexanal from linoleic acid and cis-3-hexenal and trans-2-hexenal from linolenic acid that constitute the primary components of green leaf volatile character. The temporal dynamics of aldehyde synthesis show characteristic patterns during berry development, with C6 aldehyde concentrations reaching maximum levels during early berry development and the veraison transition when lipoxygenase activities are elevated and membrane lipid turnover provides abundant fatty acid substrates, followed by progressive decline during post-veraison ripening as aldehyde reduction to corresponding alcohols and continued degradation reduce overall aldehyde content. The sensory significance of aldehyde synthesis reflects the low perception thresholds for C6 aldehydes, with hexanal and hexenal isomers detectable at concentrations of 5-15 µg/L imparting distinctive green, grassy character that is desirable in some wine styles emphasizing freshness but considered negative in fully ripe fruit where residual green notes may indicate incomplete physiological maturity. Reduction of aldehydes to their corresponding C6 alcohols through alcohol dehydrogenase activity represents a key transformation during ripening, converting the intensely green-smelling aldehydes to softer-scented hexanol and hexenol compounds that contribute more subtle herbaceous nuance without the harsh green impact of aldehyde precursors. Environmental factors influence aldehyde synthesis through effects on membrane lipid composition and lipoxygenase pathway activities, with cool conditions and high humidity potentially maintaining elevated aldehyde levels longer into the ripening season, while warm, dry conditions may accelerate aldehyde decline through enhanced reduction and volatilization. The monitoring of C6 aldehyde concentrations during ripening provides useful indicators of physiological maturity, with low residual aldehyde levels correlating with advanced ripeness and reduced green character in harvested grapes and resulting wines.
Aldehyde_Synthesis is subclass of Volatile_Compound_Synthesis representing lipoxygenase pathway volatile generation. Distinguishes from Ester_Formation which produces fruity volatiles. Sibling to Free_Volatile_Production and Ester_Formation. C6 aldehydes decline during ripening. Green character indicator. Maximum at veraison, minimum at harvest. Reduction to alcohols modifies sensory impact.
Aldehyd-Synthese
C6 aldehyde production
C6-Aldehyd-Produktion
Grün-Volatil-Synthese
Lipoxygenase-Stoffwechselweg-Aldehyde
green volatile synthesis
hexanal biosynthesis
lipoxygenase pathway aldehydes
Kalua CM and Boss PK (2009) Evolution of volatile compounds during development of Cabernet Sauvignon grapes. Journal of Agricultural and Food Chemistry 57: 3818-3830 Matsui K (2006) Green leaf volatiles: hydroperoxide lyase pathway of oxylipin metabolism. Current Opinion in Plant Biology 9: 274-280 Keller M (2020) The Science of Grapevines 3rd Ed pp 288-290
Aldehyde Synthesis
Chardonnay in Champagne shows all flowers opened during early July Year N+1 at BBCH 69 on approximately July 1, with 100% viable flowers having completed opening totaling 420-580 flowers per cluster, berry development visible 4-8 mm diameter in 62-74% successfully fertilized flowers, unfertilized flowers showing brown senescence, under cool maritime conditions of 19-24°C temperature, typical of this early-bursting moderately-to-highly fruitful cultivar completing opening approximately 12-13 days after first flowers opened
All flowers opened represents the comprehensive cluster flowering achievement when every developmentally capable flower bud has completed cap shedding and organ exposure, establishing maximum flowering extent within cluster and marking completion of opening activity though reproductive function may be declining. This opening completion milestone occurs during complete anthesis in late June-early July in temperate regions Year N+1, typically representing brief period when final flowers complete opening while earlier flowers show advanced development or senescence, corresponding to BBCH 69 defining stage. Opening completeness shows 100% of viable flowers having achieved open state with reproductive organs exposed or already transitioned to berry development, remaining unopened structures representing aborted or aberrant buds incapable of normal development rather than developmentally delayed viable flowers, and overall cluster showing complete opening extent established during primordium formation previous season. Flower distribution at all-open stage shows spatial pattern reflecting developmental gradient with apical flowers predominantly showing berry development 5-10 mm diameter from earliest successful fruit set, middle flowers showing mixed berry development and late senescence, and basal flowers showing recent opening or late senescence depending on final opening timing, creating maximum developmental heterogeneity within cluster. Developmental state diversity shows wide range of flower-berry states coexisting including advanced developing berries from early successful fertilization, recently fertilized flowers beginning berry expansion, senescing unfertilized flowers from late opening showing brown discoloration, and overall mixed appearance reflecting complete flowering extent with variable outcomes. Opening assessment enables definitive flower number determination with all-open state allowing accurate count of total flowers per cluster, comparison to previous season primordium formation evaluating developmental success from initiation to opening, and establishing denominator for fruit set percentage calculation when combined with final berry count. Reproductive opportunity at all-open stage shows effectively terminated pollination capacity with all flowers having passed through receptive period, pollen completely expired with no viable pollen remaining, and reproductive function concluded with established fruit set representing successful reproductions. Berry development visibility shows clear fruit structures throughout cluster with successful fruit set berries easily distinguished, berry size ranging 4-10 mm depending on fruit set timing from earliest to latest successful pollinations, and spatial distribution revealing architectural pattern and indicating potential cluster density or disease issues. Unfertilized flower presence shows substantial population typically 25-70% flowers failing to set fruit depending on conditions and cultivar, progressive senescence with brown discoloration and tissue deterioration, developing abscission zones preparing for detachment, and creating accumulating fallen flower debris on cluster and ground surfaces. Abscission timing shows variable pattern with earliest unfertilized flowers from early anthesis beginning to abscise before all-open achievement, continued abscission of mid-flowering failures proceeding during all-open period, and late unfertilized flowers beginning senescence preparing for subsequent abscission, creating extended flower drop period. Cluster architecture expression at all-open stage clearly reveals branching pattern established during previous season primordium formation with primary and secondary branch hierarchy visible, flower-berry spacing reflecting pedicel lengths and cluster compactness, and overall architectural form affecting disease pressure and fruit development conditions. Developmental gradient magnitude at all-open shows maximum spread with earliest-set berries potentially 7-12 days ahead of latest-set berries, creating challenge for uniform ripening and harvest timing, and affecting cluster quality homogeneity with mixed maturity levels at harvest. Berry cell division stage shows range depending on fruit set timing with earliest berries potentially completing or nearly completing cell division phase by all-open stage, while latest berries just initiating rapid cell proliferation, creating asynchronous berry development affecting final size potential. Resource allocation at all-open shows transition largely complete toward fruit development as dominant sink with developing berries consuming substantial photosynthate, vegetative growth continuing though berry demands competing for resources, and reproductive investment effectively ceased with no further flower development. Metabolic activity patterns show elevated respiration in rapidly dividing berry cells, declining metabolism in senescing unfertilized flowers, and overall cluster-level metabolism focused on developmental rather than reproductive processes. Hormonal environment at all-open shows dominance of auxin from developing berries preventing abscission and promoting growth, gibberellins affecting berry cell division and possibly contributing to seedless berry development in some situations, and ethylene potentially accumulating in unfertilized flowers promoting senescence though not affecting established berry development. Environmental conditions at all-open show reduced vulnerability with established berries relatively insensitive to weather variations compared to earlier reproductive sensitivity, though extreme conditions like hail or severe storms potentially causing physical damage to young berries. Temperature at all-open optimally 20-28°C supporting rapid berry growth through cell division, with temperature accumulation driving development rate toward subsequent cell expansion phase and eventual maturation. Management implications at all-open include confirming complete flowering enabling definitive fruit set assessment, resuming canopy management operations without concern for flower disruption, implementing berry protection strategies for disease management and possible pest control, and planning cluster management including potential thinning operations based on fruit set outcomes. Fruit set determination at all-open enables calculation by comparing berry number to total opened flower number, providing definitive percentage for yield prediction, comparison to variety norms assessing reproductive success, and informing crop adjustment decisions if set substantially exceeds or falls short of desired levels. The process represents first of three complete anthesis aspects establishing opening extent, followed by anthesis completion confirming reproductive conclusion and post-anthesis transition marking developmental phase. Duration represents brief milestone period when final flowers complete opening, with rapid observation window before developmental changes obscure opening completion. Cultivar variation in all-open characteristics shows differences in total flower number per cluster from previous season primordium formation, opening uniformity affecting developmental spread at this stage, and fruit set percentage affecting berry number and density outcomes. Regional timing shows all flowers opened during late June in warm climates, late June-early July in temperate regions, and early July in cool areas, occurring approximately 10-14 days after first flowers opening depending on flowering duration and environmental conditions. Viticultural significance includes defining complete flowering extent enabling accurate fruit set assessment, marking opening completion confirming reproductive opportunity conclusion, establishing maximum within-cluster developmental heterogeneity affecting subsequent development, providing reference for yield predictions based on berry number assessment, and confirming transition to developmental focus in management priorities. Research applications include documenting complete flowering extent and timing, analyzing relationships between primordium formation and successful opening, evaluating factors affecting flower opening success and developmental outcomes, quantifying within-cluster developmental gradients affecting ripening uniformity, and assessing implications of opening patterns for cluster quality and harvest management.
All_Flowers_Opened is first of three Complete_Anthesis aspects, distinguished by achievement of complete opening extent within cluster. Corresponds to BBCH 69 defining characteristic. Enables definitive flower number determination and fruit set calculation. Establishes maximum within-cluster developmental heterogeneity. Marks opening completion though reproductive function already concluded. Brief observation window before developmental progression obscures opening status.
all blooms exposed
alle Blüten geöffnet
complete flower opening
full cluster opening
komplette Gescheinsblüte
total anthesis expression
totale Anthesisausprägung
vollständige Blütenöffnung
Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 390-393
All Flowers Opened
Alpha-terpineol content quality evolution in aged Hunter Valley Semillon illustrates the terpene cyclization pathway, with young wines containing 20-50 μg/L α-terpineol increasing to 150-300 μg/L after 10-15 years bottle aging as linalool and geraniol cyclize under wine pH conditions this transformation contributes to the distinctive pine-eucalyptus-floral notes that emerge in mature Hunter Semillon alongside the TDN petrol character, creating the complex aromatic profile that distinguishes great aged examples from the relatively neutral young wines.
Alpha_Terpineol_Content_Quality represents the chemical quality characterizing the concentration of α-terpineol (2-(4-methylcyclohex-3-en-1-yl)propan-2-ol) in grape berries and wine, capturing the monocyclic monoterpene alcohol that contributes floral-pine, lilac-like aroma notes and serves as an important transformation product of other terpenes during wine aging. α-Terpineol is distinguished from the acyclic terpene alcohols by its cyclohexene ring structure, formed through acid-catalyzed cyclization of linalool, geraniol, or nerol under the mildly acidic conditions of wine. This cyclization pathway makes α-terpineol both a grape-derived compound (present in berries at modest concentrations) and an aging-derived compound (formed from acyclic terpene precursors during wine maturation). In fresh grapes and young wines from aromatic varieties, α-terpineol typically occurs at concentrations of 20-100 μg/L, representing a minor fraction of the total monoterpene pool. However, during extended wine aging, α-terpineol concentration can increase substantially as linalool undergoes acid-catalyzed cyclization, potentially reaching 200-400 μg/L in decade-old aromatic wines. The sensory threshold of α-terpineol in wine is approximately 250-400 μg/L, near the upper range of aged wine concentrations, meaning it contributes modestly to direct aroma perception while participating in mixture interactions. The α-terpineol character (floral, pine, lilac) differs from the fresh citrus-floral notes of linalool, making the linalool-to-α-terpineol transformation part of the aromatic evolution that distinguishes young from aged aromatic wines. Four monoterpene alcohol isomers share the terpineol name (α, β, γ, δ), with α-terpineol being the most abundant and aromatically significant in wine.
Alpha_Terpineol_Content_Quality addresses both the grape-derived and transformation-derived pools of this cyclic monoterpene. The acid-catalyzed cyclization of linalool to α-terpineol represents a key aging pathway affecting aromatic evolution in white wines. The class specifically addresses α-terpineol, the most abundant and aromatically relevant terpineol isomer in wine other terpineol isomers (β, γ, δ) are present at lower concentrations. The relatively high sensory threshold of α-terpineol means its direct contribution to aroma is modest, but it participates in complex mixture interactions and serves as a marker for terpene transformation during aging.
Alpha-Terpineolgehalt
Terpineolgehalt
alpha-terpineol level
cyclic terpene alcohol quality
terpineol content
zyklischer Terpenalkoholgehalt
α-Terpineol-Konzentration
α-terpineol concentration
Simpson RF and Miller GC (1984) Aroma Composition of Chardonnay Wine. Vitis 23(2): 143-158 Sacks GL et al. (2012) Determination of Linalool and α-Terpineol in Wines: Comparison of Methods Involving Solid Phase Microextraction and Direct Liquid Injection. American Journal of Enology and Viticulture 63(2): 246-251
Alpha Terpineol Content Quality
Pinot Noir clone Pommard bears an American_Clone_Role as a selection brought from Burgundy in the pre-FPS era, maintained and distributed through UC Davis programs as one of the historically important American Pinot Noir clones alongside Wädenswil and "suitcase clones" from various importations. Chardonnay clone 04 (Old Wente) serves in an American_Clone_Role as the historically dominant California Chardonnay, a UC Davis selection from the Wente family's original French importations that shaped California Chardonnay production for decades. Cabernet Sauvignon clone 08 bears an American_Clone_Role as an FPS selection widely planted in California, processed through FPS protocols to ensure virus-free status and available to licensed nurseries for commercial propagation.
A regional clone role that inheres in a grapevine clone selected, evaluated, or registered within American (United States) vine improvement systems, coordinated primarily through Foundation Plant Services (FPS) at UC Davis (California) with additional contributions from state programs in New York (Cornell), Washington, Oregon, Virginia, and other viticultural states. American clones are identified by variety name and alphanumeric codes reflecting FPS processing (e.g., Cabernet Sauvignon 08, Chardonnay 04) or state program designations. The American clone system developed around FPS (established 1958), which provides comprehensive virus testing, pathogen elimination, and clean plant distribution serving the national nursery industry. American clone programs handle both domestic selections (identified from California, Oregon, and other state vineyards) and international imports (processed through FPS quarantine to meet USDA phytosanitary requirements), creating a diverse clone inventory supporting the expanding American wine industry. The National Clean Plant Network (NCPN) coordinates vine improvement across states, with FPS serving as the primary grapevine clean plant center.
American_Clone_Role captures the US clone system where FPS dominates national clean plant distribution while state programs address regional needs and variety specialization. The American approach differs from European systems in its relative youth (systematic programs from the 1950s-1960s), emphasis on imported selections processed through quarantine rather than indigenous selection, and integration with a primarily V. vinifera industry established through importation rather than native species cultivation. "Heritage clones" (pre-FPS importations including Pommard, Wädenswil, "suitcase clones") represent material that entered the US before modern certification systems, carrying historical significance but often uncertain health status until subjected to FPS processing. The National Clean Plant Network expansion has strengthened regional programs while maintaining FPS's central role in grapevine improvement.
American clone
American registered clone
FPS-distributed clone
US clone
US-Klon
amerikanisch registrierter Klon
amerikanischer Klon
Golino DA et al. (2017) Foundation Plant Services and clean plant programs, HortScience 52: 1190-1194 Rowhani A et al. (2018) Grapevine virus testing protocols, UC Davis FPS Weber E et al. (2002) The California grapevine certification program, American Journal of Enology and Viticulture 53: 130-134 Wolpert JA et al. (1994) Selection of Pinot Noir clones, American Journal of Enology and Viticulture 45: 356-363
American Clone Role
Concord (complex V. labrusca hybrid, possibly with V. vinifera ancestry) bears an American_Hybrid_Role as the most commercially important American grape variety, dominating juice and jelly production (>300,000 tons annually in the USA) and defining the "grapey" flavor (methyl anthranilate) associated with American grapes. Marquette (complex hybrid from University of Minnesota breeding) serves in an American_Hybrid_Role as a contemporary cold-hardy wine grape, surviving -35°C winters while producing red wines of recognized quality from multiple Vitis species contributions. Norton/Cynthiana (possibly V. aestivalis selection or hybrid) bears an American_Hybrid_Role as the signature grape of Missouri viticulture, its Pierce's disease resistance enabling cultivation in regions where V. vinifera fails.
An interspecific hybrid role that inheres in grapevine varieties developed primarily from crosses among native North American Vitis species (V. labrusca, V. aestivalis, V. riparia, V. rupestris, V. rotundifolia, V. cinerea, and others) with or without V. vinifera contribution, bred for cultivation in North American growing conditions where European varieties failed. American hybrids emerged from 19th-century efforts to domesticate native American species and combine their disease resistance and cold hardiness with improved fruit quality, with breeding programs by Munson, Rommel, Jaeger, and others creating varieties suited to regions where V. vinifera could not survive. Pure American species crosses (no V. vinifera) dominate in regions with extreme cold (Minnesota, upper Midwest) or Pierce's disease pressure (Gulf Coast), while V. vinifera contributions increase in varieties targeting wine quality improvement. American hybrids remain commercially significant for juice and jelly production (Concord, Niagara, Catawba representing billions of dollars in annual production), for extreme-climate wine production (Marquette, Frontenac in Minnesota Norton/Cynthiana in Missouri), and as breeding germplasm for resistance and hardiness traits.
American_Hybrid_Role encompasses varieties developed from native North American Vitis species, whether pure species selections, interspecific American hybrids, or complex hybrids with variable V. vinifera contribution. The distinction from French-American hybrids is primarily geographic and programmatic (American versus French breeders) rather than strictly genetic, as genetic contributions from American species are fundamental to both categories. American hybrids serve different market niches than V. vinifera—juice and processed products (Concord, Niagara), cold-climate wine production (northern hybrids), and extreme-climate viticulture where V. vinifera is not viable. The V. labrusca-derived "foxy" flavor (methyl anthranilate) that defines Concord character is considered a defect in wine contexts but is the desired flavor profile for juice and jelly, illustrating how American hybrid evaluation depends on intended use. Contemporary American breeding programs (University of Minnesota, Cornell, Missouri) continue developing improved cold-hardy and disease-resistant varieties for challenging growing regions.
American grape
American hybrid
American variety
Neuwelt-Hybride
New World hybrid
amerikanische Hybride
amerikanische Rebsorte
Reisch BI et al. (2012) Grape breeding in the United States, CRC Press pp.: 171-203 Hemstad PR, Luby JJ (2000) Utilization of Vitis riparia for the development of new wine varieties with resistance to disease and extreme cold, Acta Horticulturae 528: 487-496 This P et al. (2006) Historical origins of wine grapes, Trends in Genetics 22: 511-519 Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane
American Hybrid Role
In Chardonnay berries in Côte de Beaune, Burgundy during BBCH 83-89, amino acid accumulation increases total free amino acids from 320 to 1450 mg/L over 42 days, with proline increasing from 85 to 680 mg/L representing 47% of final total, arginine from 95 to 420 mg/L at 29%, and branched-chain amino acids from 18 to 85 mg/L, providing yeast assimilable nitrogen of 285 mg N/L under moderate vineyard nitrogen status and mean ripening temperature of 21°C.
Amino acid accumulation describes the progressive increase in free amino acid content within Vitis vinifera berries during the ripening period, wherein total amino nitrogen increases from 200-400 mg/L at veraison to 800-2000 mg/L at harvest through continued biosynthesis, protein turnover, and nitrogen remobilization from vegetative tissues, establishing the amino acid pools that serve as essential precursors for fermentation-derived flavor compounds, yeast nutrition substrates, and wine stability determinants. This process involves the accumulation of diverse amino acid species with proline typically constituting 30-65% of total amino acids in ripe berries due to its role in osmotic adjustment and stress response, followed by arginine at 15-35% serving as the primary nitrogen storage form, glutamine and glutamate at 5-15% functioning as nitrogen assimilation intermediates, and the remaining amino acids including threonine, serine, alanine, and the branched-chain amino acids leucine, isoleucine, and valine contributing 10-25% collectively with particular significance as Ehrlich pathway substrates for higher alcohol production. The temporal progression of amino acid accumulation shows characteristic patterns with initial increases during early post-veraison driven by continued biosynthesis and nitrogen uptake, mid-ripening protein degradation contributing amino acids from turnover of photosynthetic and storage proteins, and late-ripening nitrogen remobilization from senescing leaves providing additional amino acid influx through phloem transport during the final weeks before harvest. Cellular distribution of accumulated amino acids involves cytosolic pools available for immediate metabolic utilization and vacuolar storage pools that buffer concentration fluctuations, with proline showing particularly strong vacuolar sequestration reflecting its role in osmotic regulation and stress protection during the dehydration associated with late-ripening berry concentration. Nitrogen fertilization and vineyard soil nitrogen status profoundly influence amino acid accumulation, with adequate nitrogen supply supporting accumulation to levels of 1500-2500 mg N/L considered optimal for fermentation performance, while nitrogen deficiency limits accumulation to potentially problematic levels below 500 mg N/L that may result in sluggish or incomplete fermentation. Cultivar variation in amino acid accumulation reflects genetic differences in nitrogen uptake efficiency, assimilation enzyme activities, and nitrogen partitioning between storage and metabolic pools, with varieties such as Riesling and Chardonnay typically showing higher total amino acid content than varieties such as Cabernet Sauvignon and Merlot under equivalent nitrogen supply conditions.
Amino_Acid_Accumulation is subclass of Precursor_Formation representing nitrogen-containing flavor precursor development. Distinguishes from Flavor_Precursor_Synthesis which involves conjugation reactions. Sibling to Flavor_Precursor_Synthesis and Precursor_Pool_Development. Key parameter is total amino nitrogen increase from 200-400 to 800-2000 mg/L. Proline typically 30-65% of total. Critical for fermentation performance.
Aminosäure-Akkumulation
Beeren-Stickstoff-Akkumulation
amino acid pool development
berry nitrogen accumulation
free amino acid increase
freie Aminosäure-Zunahme
hefeverwertbarer Stickstoff-Anreicherung
yeast assimilable nitrogen accumulation
Bell SJ and Henschke PA (2005) Implications of nitrogen nutrition for grapes, fermentation and wine. Australian Journal of Grape and Wine Research 11: 242-295 Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 290-295 Stines AP et al (1999) Proline and arginine accumulation in developing berries of Vitis vinifera. Australian Journal of Grape and Wine Research 5: 14-23
Amino Acid Accumulation
Amino acid profile quality analysis in Muscadet Melon de Bourgogne reveals the characteristic threonine elevation (15-30 mg/L, 10-15% of total amino acids) that distinguishes this variety and contributes to the distinctive yeasty, dairy notes through threonine-derived acetaldehyde and diacetyl during extended lees contact (sur lie aging) the profile-guided selection of yeast strains with appropriate threonine metabolism enables winemakers to modulate the buttery character, enhancing it for traditional sur lie style or minimizing it for fresh, mineral-focused expressions.
Amino_Acid_Profile_Quality encompasses the chemical qualities characterizing the concentration and proportional relationships among individual free amino acids in grape must, capturing the compositional detail that affects yeast metabolism, volatile compound formation, and wine quality beyond what aggregate YAN measurement reveals. The amino acid profile provides information about the specific substrates available for yeast metabolic pathways, including the Ehrlich pathway converting amino acids to higher alcohols and their esters, the sulfur amino acid pathways affecting hydrogen sulfide risk and thiol formation, and the arginine metabolism relevant to ethyl carbamate precursor formation. In grape must, approximately 20 free amino acids are present at measurable concentrations, with proline, arginine, glutamine, alanine, glutamate, and threonine typically dominant. The profile varies substantially among cultivars, with some varieties showing characteristic patterns useful for authenticity assessment. Viticultural factors including nitrogen fertilization, vine water status, and harvest timing modulate both total amino acid content and profile proportions. The profile affects fermentation outcomes through multiple mechanisms: branched-chain amino acids (leucine, isoleucine, valine) yield the fusel alcohols and esters contributing to wine fruitiness aromatic amino acids (phenylalanine, tyrosine, tryptophan) produce phenolic and indolic derivatives and sulfur amino acids (methionine, cysteine) provide sulfur for yeast metabolism while affecting volatile sulfur compound formation. Profile analysis employs HPLC with fluorescence detection following derivatization (OPA, FMOC) or mass spectrometric detection for comprehensive amino acid quantification.
Amino_Acid_Profile_Quality serves as the parent class for individual amino acid qualities including Proline, Arginine, Glutamine, and GABA Content Qualities. The profile concept recognizes that individual amino acid concentrations provide information beyond aggregate measurements, affecting specific metabolic pathways and wine characteristics. Profile analysis requires HPLC methodology rather than the simpler colorimetric methods sufficient for aggregate PAN measurement. The varietal fingerprinting potential of amino acid profiles provides authenticity assessment applications alongside fermentation management utility.
Aminosäureprofil
Aminosäurezusammensetzung
Einzelaminosäuregehalt
amino acid composition
amino acid distribution quality
free amino acid profile
freies Aminosäureprofil
individual amino acid content
Huang Z and Ough CS (1991) Amino Acid Profiles of Commercial Grape Juices and Wines. American Journal of Enology and Viticulture 42(3): 261-267 Garde-Cerdán T and Ancín-Azpilicueta C (2008) Effect of the Addition of Different Quantities of Amino Acids to Nitrogen-Deficient Must on the Formation of Esters, Alcohols, and Acids during Wine Alcoholic Fermentation. LWT - Food Science and Technology 41(3): 501-510
Amino Acid Profile Quality
Ammonium content for Chardonnay from Meursault (2024 vintage) measures 48 mg N/L (28% of 172 mg N/L total YAN), within the typical range for low-to-moderate vigor Burgundian vineyards on limestone soils, with the relatively low ammonium fraction informing the decision to supplement with a complex organic nitrogen source (yeast derivatives providing amino acids) rather than DAP (which provides only ammonium) to enhance ester diversity. Ammonium content for Shiraz from McLaren Vale (2023 vintage) records 110 mg N/L (38% of 290 mg N/L YAN), reflecting the higher nitrogen mineralization rates of the warm Mediterranean growing conditions, with the elevated ammonium sufficient to drive rapid yeast population growth during the critical early fermentation phase without additional supplementation.
Ammonium content is a nitrogen data entity that records the concentration of ammonium ions (NH₄⁺) in grape must, determined by enzymatic assay (glutamate dehydrogenase/NADH coupled reaction), ion-selective electrode, or distillation-titration method, expressed in milligrams of nitrogen per liter (mg N/L) or milligrams of ammonium per liter (mg NH₄⁺/L), representing the inorganic nitrogen fraction immediately available for yeast uptake without prior enzymatic deamination and constituting the preferentially consumed nitrogen source during the early exponential growth phase of alcoholic fermentation.
Ammonium ions are the preferred nitrogen source for Saccharomyces cerevisiae during fermentation, consumed before amino acids through the nitrogen catabolite repression (NCR) mechanism — when ammonium is present above approximately 2-3 mM, yeast suppress the expression of amino acid permeases and preferentially assimilate NH₄⁺ through the high-affinity MEP (methylammonium permease) transport system and the NADPH-dependent glutamate dehydrogenase (GDH1) assimilation pathway. This preferential uptake has practical implications — ammonium additions (as DAP, diammonium phosphate) are rapidly consumed and stimulate biomass production during early fermentation but do not contribute to amino acid-derived aroma compounds. The enzymatic assay using glutamate dehydrogenase (GLDH, EC 1.4.1.3) coupled with NADH oxidation provides a specific, rapid determination suitable for routine laboratory analysis, with commercially available kits adapted for must and wine matrices. Ammonium content in grape must is influenced by soil nitrogen form (ammonium-dominated versus nitrate-dominated soils), recent fertilizer applications, soil temperature (affecting nitrification rates), and vine nitrogen partitioning — foliar urea applications increase arginine more than ammonium in berries, while soil-applied ammonium fertilizers can elevate must ammonium if applied during the ripening period. Excessive ammonium in must (>150 mg N/L) has been associated with increased ethyl carbamate (urethane) precursor formation through the urea cycle, a food safety consideration in some regulatory jurisdictions that set advisory limits on ethyl carbamate concentrations in wine.
Ammoniumgehalt
Ammoniumionenkonzentration
Ammoniumstickstoff
NH4 content
NH4-Gehalt
ammonia nitrogen content
ammonium ion concentration
ammonium nitrogen
anorganischer Stickstoff
inorganic nitrogen
Bell SJ, Henschke PA (2005) Implications of nitrogen nutrition for grapes, fermentation and wine. Australian Journal of Grape and Wine Research 11(3): 242-295 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1, 2nd Ed, Wiley, pp.: 80-95 ter Schure EG et al. (2000) The role of ammonia metabolism in nitrogen catabolite repression in Saccharomyces cerevisiae. FEMS Microbiology Reviews 24(1): 67-83
Ammonium Content
Ammonium content quality in Mosel Riesling must typically contributes 30-60 mg N/L of the total 120-180 mg N/L YAN, with the relatively low ammonium fraction reflecting the region's limited soil nitrogen availability and cool climate reducing ammonium accumulation winemakers managing slow cool-climate fermentations (12-15°C) may supplement with small DAP additions (10-20 g/hL providing 20-40 mg N/L ammonium) to ensure adequate early fermentation nitrogen without the excessive ammonium that can promote volatile acidity in extended low-temperature fermentations.
Ammonium_Content_Quality represents the chemical quality characterizing the concentration of ammonium ions (NH₄⁺) in grape must, capturing the inorganic nitrogen fraction that provides immediately available nitrogen nutrition for wine yeast during fermentation. Ammonium is the preferred nitrogen source for Saccharomyces cerevisiae, taken up rapidly through specific membrane transporters (Mep1, Mep2, Mep3) and assimilated directly into glutamate and glutamine without requiring deamination. In grape must, ammonium concentration typically ranges from 20-150 mg N/L as ammonium nitrogen, comprising 10-40% of total YAN depending on cultivar, vine nitrogen status, and fruit maturity. Ammonium accumulates in berries from root uptake and from amino acid catabolism during ripening, with concentrations generally lower than amino nitrogen in most musts. The preferential uptake of ammonium by yeast creates sequential nitrogen utilization during fermentation, with ammonium depleted rapidly in the first days before significant amino acid uptake begins. This rapid early uptake makes ammonium particularly important for fermentation initiation and early kinetics. Diammonium phosphate (DAP) supplementation provides ammonium directly and represents the most common YAN correction method, though excessive DAP can produce imbalanced fermentation, elevated volatile acidity, and reduced ester complexity compared to organic nitrogen supplementation. The ammonium-to-amino nitrogen ratio affects fermentation outcomes beyond total YAN, with balanced nitrogen sources generally producing better aromatic outcomes than either extreme. Ammonium analysis employs enzymatic methods (glutamate dehydrogenase), ion-selective electrodes, or ion chromatography.
Ammonium_Content_Quality addresses the inorganic nitrogen component of YAN, distinguished from organic amino nitrogen. The preferential uptake of ammonium by yeast makes it particularly important for fermentation initiation despite typically comprising the minority of YAN. The class addresses natural grape-derived ammonium rather than supplementation-derived ammonium, though the analytical measurement cannot distinguish origin. The role of DAP supplementation in providing ammonium links grape ammonium content to common winemaking interventions for nitrogen management.
Ammoniumstickstoffgehalt
NH₄⁺ concentration
NH₄⁺-Konzentration
ammonium ion quality
ammonium nitrogen content
anorganischer assimilierbarer Stickstoff
inorganic assimilable nitrogen
Crépin L et al. (2012) Sequential Use of Nitrogen Compounds by Saccharomyces cerevisiae during Wine Fermentation: A Model Based on Kinetic and Regulation Characteristics of Nitrogen Permeases. Applied and Environmental Microbiology 78(22): 8102-8111 Bisson LF (1999) Stuck and Sluggish Fermentations. American Journal of Enology and Viticulture 50(1): 107-119
Ammonium Content Quality
Ammonium uptake typically accounts for 10-30% of total nitrogen acquisition in aerated vineyard soils, increasing to 40-60% under waterlogged or acidic conditions where nitrification is inhibited. Grapevine roots show preference for ammonium over nitrate when both forms are equally available, with ammonium uptake rates 20-40% higher at equivalent nitrogen concentrations. VvAMT1 and VvAMT2 genes encode ammonium transporters with differential expression in root tips versus mature root zones. Excessive ammonium uptake (NH₄⁺:NO₃⁻ ratio above 3:1) can induce toxicity symptoms including root browning and reduced growth, particularly under low potassium availability.
A nitrogen uptake efficiency disposition that is the inherited capacity of a Vitis vinifera plant to absorb ammonium ions (NH₄⁺) from the soil solution through ammonium transporter (AMT) family proteins, providing a direct nitrogen source requiring no reduction before amino acid synthesis.
Ammonium uptake provides energetic advantages over nitrate acquisition by eliminating the reduction step required for nitrate assimilation, potentially improving nitrogen use efficiency under energy-limited conditions. However, ammonium toxicity risk and the transient nature of soil ammonium pools limit its contribution in most vineyard systems. Rootstock genotypes differ in ammonium tolerance, with some showing growth inhibition at concentrations that others tolerate. Fertigation with ammonium-containing fertilizers can exploit genotypic differences in ammonium uptake capacity.
Ammoniumaufnahme
Ammoniumtransporter-Aktivität
NH4 uptake capacity
NH₄⁺-Akquisition
ammonium acquisition
ammonium transporter activity
Lang CP et al. (2020) Different nitrogen sources affect grapevine root morphology and arbuscular mycorrhizal colonization. Vitis 59:79-85 Gessler A et al. (1998) Field and laboratory experiments on net uptake of nitrate and ammonium by the roots of spruce (Picea abies) and beech (Fagus sylvatica) trees. New Phytologist 138:275-285
Ammonium Uptake Disposition
Ammonium uptake rate comparison between acidic (pH 5.2) and neutral (pH 7.0) vineyard soils reveals the soil pH effect on nitrogen form and uptake, with vines in acidic soil showing ammonium uptake of 5.2 μmol NH₄⁺/g root/hour (high soil ammonium from limited nitrification) contributing 45% of total nitrogen uptake, while vines in neutral soil show only 0.8 μmol/g/hour (rapid nitrification converting ammonium to nitrate) contributing only 12% of total nitrogen, demonstrating how soil conditions affect nitrogen form availability and the relative importance of nitrate versus ammonium uptake pathways.
Ammonium uptake rate quality is a nitrogen uptake efficiency quality that represents the rate of ammonium (NH₄⁺) absorption by grapevine roots, characterizing the acquisition of the nitrogen form that can be directly incorporated into amino acids without reduction, typically less abundant than nitrate in most vineyard soils but potentially significant in acidic soils, recently fertilized sites, or under conditions limiting nitrification. Within the BFO framework, this quality specifically addresses the ammonium form of nitrogen uptake, distinguishing ammonium from nitrate acquisition while recognizing that ammonium, though often less available than nitrate, is energetically more efficient to assimilate (no reduction required) and may become the dominant nitrogen form under specific soil conditions. The ontological significance lies in ammonium uptake rate affecting both the quantity and energetic cost of nitrogen acquisition, where direct ammonium availability provides nitrogen that requires less metabolic energy to incorporate into amino acids and proteins, potentially improving the overall efficiency of nitrogen nutrition when ammonium is available, though high ammonium uptake can also have toxicity implications requiring careful regulation. The molecular basis for ammonium uptake involves the AMT (Ammonium Transporter) protein family that mediates active and facilitated ammonium transport across root cell membranes, with high-affinity AMT transporters enabling acquisition at low external concentrations and transporter expression regulated by nitrogen status to balance uptake with assimilation capacity. The soil conditions affecting ammonium availability include soil pH (acidic conditions favoring ammonium persistence, limiting nitrification), temperature (cold conditions slowing nitrification), waterlogging (anaerobic conditions preventing nitrification), and recent nitrogen fertilization (temporary ammonium pulse before nitrification), with ammonium uptake becoming quantitatively important when these conditions make ammonium abundant. The potential toxicity of ammonium at high concentrations requires that roots have mechanisms to prevent excessive uptake that could disrupt cellular pH and ion balance, with AMT transporter regulation providing one level of control over ammonium acquisition. Understanding ammonium uptake rate quality enables assessment of nitrogen nutrition under conditions favoring ammonium availability, optimization of nitrogen fertilizer form selection, and interpretation of vine nitrogen dynamics in soils with varying nitrogen form distributions.
Ammonium_Uptake_Rate_Quality addresses the acquisition rate of the energetically efficient nitrogen form requiring no reduction for assimilation. This quality is mediated by AMT transporter family and becomes important under conditions limiting nitrification. The class supports nitrogen form management and soil-specific fertilization through ammonium uptake characterization.
AMT activity
AMT-Aktivität
Ammoniumakquisitionsrate
Ammoniumaufnahmerate
NH₄⁺ uptake
NH₄⁺-Aufnahme
ammonium absorption
ammonium acquisition rate
Loqué D, von Wirén N (2004) Regulatory levels for the transport of ammonium in plant roots. Journal of Experimental Botany 55(401): 1293-1305 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 197-245
Ammonium Uptake Rate Quality
The ampelographic description of Riesling compiled according to the OIV descriptor system documents over 80 standardized morphological characters — including the open shoot tip with dense prostrate hairs (OIV 001:7, OIV 004:7), pentagonal mature leaf with five lobes and open V-shaped petiolar sinus (OIV 067:3, OIV 076:3, OIV 079:3), small compact cylindrical cluster with one wing (OIV 202:3, OIV 204:5, OIV 208:2), small round yellow-green berries (OIV 220:1, OIV 223:2, OIV 225:3), and characteristic spicy-floral aroma (OIV 236:2) — this comprehensive morphological profile has been the basis for Riesling identification since Clements (1830) and remains the legal reference for cultivar registration in the European Union Plant Variety Office (CPVO) system alongside the molecular SSR fingerprint.
A grapevine information entity that systematically documents the morphological characteristics of a Vitis vinifera cultivar through standardized observation and recording of vegetative and reproductive organ features — shoot tips, young and mature leaves, flowers, clusters, berries, seeds, shoots, and canes — following internationally recognized descriptor systems that enable cultivar identification, distinction, and classification based on phenotypic traits observable in the vineyard.
Ampelographic_Description is the root class for the systematic morphological characterization of grapevine cultivars — a discipline (ampelography, from Greek ampelos = vine + graphein = to write) with a documented history spanning over 2,000 years from Columella's De Re Rustica through the foundational modern works of Clements (1830), Viala and Vermorel (1901-1910), Galet (1956-2000), and the OIV standardization (1983-present). The transition from subjective prose descriptions to standardized coded descriptors represents the central methodological achievement of modern ampelography: the OIV (Organisation Internationale de la Vigne et du Vin) descriptor system, first published in 1983 and progressively refined through six editions, defines over 130 morphological characters each scored on a standardized scale (typically 1-9 with defined reference cultivars for each state), enabling quantitative comparison and database management of ampelographic data. Despite the ascendancy of molecular identification through SSR and SNP markers, ampelographic description retains essential roles in cultivar registration (required by UPOV and EU plant variety protection regulations), clone distinction within cultivars (where morphological differences may be visible but molecular differences undetectable), and the practical vineyard-level identification that does not require laboratory analysis.
ampelographic characterization
ampelographische Beschreibung
morphological cultivar description
morphologische Sortenbeschreibung
phenotypic cultivar documentation
phänotypische Sortencharakterisierung
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Viala P and Vermorel V (1901-1910) Ampélographie — traité général de viticulture, 7 volumes, Masson UPOV (2011) Guidelines for the Conduct of Tests for Distinctness, Uniformity and Stability — Vine (Vitis L.), TG/50/9
Ampelographic Description
Kadarka accession SZBKI-Ka-1 at the National Centre for Biodiversity and Gene Conservation in Tápiószele, Hungary bearing an ampelographic reference role is documented with the complete set of 68 OIV primary descriptors including young shoot tip pigmentation (OIV 001: 5-7), mature leaf blade shape (OIV 067: pentagonal), number of lobes (OIV 068: five), petiolar sinus shape (OIV 079: wide open), berry shape (OIV 223: spherical), and seed morphology (OIV 242-244), with standardized photographic documentation of shoot tips, leaves, clusters, and berries archived in the Hungarian Vitis Database and cross-referenced to the VIVC descriptor profile for international identification verification.
A research and education role that inheres in a grapevine maintained as an authenticated specimen for the morphological description and visual identification of grape cultivars using standardized ampelographic descriptor systems, serving as the physical basis for cultivar documentation in ampelographic atlases, descriptor databases, and identification keys.
Ampelographic_Reference_Role preserves the classical morphological approach to cultivar identification that predates molecular methods and remains essential for field identification, historical documentation, and regulatory cultivar verification. Ampelography — the systematic morphological description of grapevines — was formalized in the 19th century by Odart, Viala, and Vermorel and standardized through OIV descriptor codes adopted internationally from 1983 onward: the discipline requires authenticated reference specimens maintained under standardized conditions because morphological expression is influenced by environment, rootstock, virus status, and vine age. The role complements Genetic_Reference_Role, which addresses molecular identification.
Sortenbeschreibungs-Rolle
ampelographic standard role
ampelographische Referenz-Rolle
cultivar description role
morphological reference role
morphologische Beschreibungs-Rolle
OIV (2009) 2nd Edition of the OIV Descriptor List for Grape Varieties and Vitis Species Galet P (2000) Dictionnaire encyclopédique des cépages, Hachette Viala P and Vermorel V (1901-1910) Ampélographie: traité général de viticulture, 7 vols, Masson et Cie
Ampelographic Reference Role
Ampelographic source documentation for Central European cultivars references a progression of increasingly systematic works: Franz Trummer's "Systematische Classification aller bekannten Traubenarten" (1841, Graz — the first comprehensive Austrian ampelographic catalogue, documenting approximately 400 cultivars including the earliest systematic Grüner Veltliner description), Hermann Goethe's "Handbuch der Ampelographie" (1878, 2nd edition 1887, Graz-Berlin — the major Austro-German reference describing over 3,000 named varieties with detailed morphological characterization), the Bundesamt für Weinbau Klosterneuburg collection catalogues (maintained continuously from 1860 to present, providing the Austrian national reference collection), and the contemporary VIVC database (Julius Kühn-Institut, maintained from 1984 to present, integrating ampelographic descriptions with molecular profiles for over 24,000 cultivar entries including synonyms). Ampelographic source for French cultivars references the monumental "Ampélographie" of Pierre Viala and Victor Vermorel (7 volumes, 1901-1910, Paris — the most comprehensive ampelographic work ever produced, describing over 5,200 cultivar names with chromolithographic illustrations by Alexis Kreyder).
Ampelographic source is a documentation source entity that documents the systematic cultivar description treatises, varietal catalogues, and vine collection records that constitute the ampelographic tradition — the scientific discipline of grapevine identification, description, and classification based on morphological observation — encompassing the foundational European ampelographic works (from early 19th-century pioneering catalogues through the monumental late 19th/early 20th-century national ampelographies to modern standardized descriptor systems), vine collection catalogues (documenting cultivar holdings at research institution repositories), and the contemporary digital databases that maintain and update cultivar descriptions, specifying for each source the author, date, institutional affiliation, taxonomic scope, descriptive methodology employed, and the reliability and completeness of the cultivar characterizations contained.
Ampelographic source documentation illuminates one of the most distinctive scholarly traditions in agricultural science — a discipline that developed from the late 18th century onward in response to the practical need to identify, classify, and standardize the chaotic profusion of grape cultivar names and descriptions that had accumulated through centuries of local viticultural practice. The history of ampelography as a scientific discipline encompasses several methodological phases — the early descriptive period (1800-1850, exemplified by Trummer, Burger, and Count Odart, relying on prose descriptions and limited illustrations), the classical morphological period (1850-1950, exemplified by Viala and Vermorel, Galet, and Goethe, employing standardized morphological terminology, chromolithographic illustration, and systematic comparison across large cultivar collections), the standardized descriptor period (1950-1990, culminating in the OIV descriptor list of 1983 with its coded morphological characters enabling computer-assisted identification), and the molecular period (1990-present, integrating microsatellite DNA fingerprinting with morphological description to provide unambiguous cultivar identification independent of environmental variation). The OIV descriptor system codified 131 morphological, phenological, and agronomic characters for standardized cultivar description, each scored on defined scales — shoot tip pigmentation (OIV 003), mature leaf shape (OIV 067), berry shape (OIV 223), and so on — providing the internationally harmonized framework for ampelographic description that superseded the diverse national descriptor traditions. The Vitis International Variety Catalogue (VIVC) at the Julius Kühn-Institut represents the contemporary culmination of the ampelographic tradition — a comprehensive database integrating morphological descriptions, molecular fingerprints (SSR profiles), synonymy and homonymy resolution, parentage analysis results, and bibliographic references for over 24,000 cultivar entries, serving as the global reference authority for cultivar identification and nomenclature. The reliability of historical ampelographic sources requires careful assessment — pre-molecular descriptions sometimes conflated distinct cultivars (identical synonyms) or split a single cultivar into multiple entries (environmental variants described as separate varieties), and retrospective molecular verification has revealed both confirmations and corrections of classical ampelographic identifications.
Rebsortenbeschreibung
Sortenbeschreibungsquelle
Sortenkatalogreferenz
ampelographic reference
ampelographic treatise
ampelographische Quelle
ampelographisches Werk
cultivar description reference
varietal catalogue
vine description source
Viala P, Vermorel V (1901-1910) Ampélographie: Traité général de viticulture, 7 vols, Masson, Paris Galet P (1962) Cépages et vignobles de France, 2nd Ed (1988-1990), 4 vols, Imprimerie Déhan, Montpellier Goethe H (1878) Handbuch der Ampelographie, 2nd Ed 1887, Verlag Paul Parey, Berlin OIV (2009) 2nd Edition of the OIV Descriptor List for Grape Varieties and Vitis Species VIVC (2024) Vitis International Variety Catalogue, Julius Kühn-Institut, https://www.vivc.de
Ampelographic Source
Biochemical characterization of Anaerobic_Metabolism_Disposition at UC Davis examined fermentative enzyme activities, end-product accumulation, and energy status across 12 rootstocks subjected to strict root anoxia (N₂-flushed nutrient solution) for 72 hours. Activities of alcohol dehydrogenase (ADH) and pyruvate decarboxylase (PDC), tissue concentrations of ethanol, lactate, and ATP, and cytoplasmic pH (by ³¹P-NMR) were measured at 0, 24, 48, and 72 hours. Anaerobic_Metabolism_Disposition showed significant genotypic variation in multiple parameters. 'Riparia Gloire' exhibited strong anaerobic capacity with ADH activity increasing 8.5-fold within 24 hours (to 4.2 μmol min⁻¹ g⁻¹ FW), PDC activity increasing 6-fold, ethanol accumulating to moderate levels (12 mM at 72h, then declining suggesting export/volatilization), lactate remaining low (2.5 mM), and cytoplasmic pH stable at 7.1 (no acidification). Root ATP in 'Riparia Gloire' declined to 35% of aerobic control but stabilized at this level, indicating sustained energy production. '3309C' showed similar metabolic adaptation (8-fold ADH induction, stable pH, 38% ATP maintenance). '140 Ruggeri' displayed poor Anaerobic_Metabolism_Disposition with only 3-fold ADH induction, high lactate accumulation (8.5 mM) indicating metabolic imbalance, cytoplasmic acidification (pH declining to 6.5 by 72h), and ATP collapsing to 8% of control. Upon re-aeration, 'Riparia Gloire' cleared accumulated ethanol within 6 hours and restored ATP to 85% of control by 24 hours, while '140 Ruggeri' showed persistent metabolic dysfunction (ATP recovery to only 45%) suggesting irreversible damage. These findings establish that Anaerobic_Metabolism_Disposition involves both efficient fermentative enzyme induction and metabolic regulation preventing self-toxicity.
An Anaerobic_Metabolism_Disposition is a Waterlogging_Tolerance_Disposition that inheres in Vitis root tissues and is realized through metabolic adaptations that enable continued ATP production and cellular function when oxygen availability is insufficient for aerobic respiration, utilizing fermentative pathways and tolerating their byproducts. This disposition represents the biochemical component of waterlogging tolerance, determining how effectively roots can maintain energy production and metabolic function under oxygen deprivation when anatomical adaptations (aerenchyma, adventitious roots) are insufficient to fully alleviate hypoxia. Anaerobic metabolism in plant roots involves glycolysis coupled to ethanolic fermentation (via pyruvate decarboxylase and alcohol dehydrogenase) or lactic acid fermentation, yielding only 2-3 ATP per glucose compared to 36-38 from aerobic respiration. Survival under hypoxia requires efficient glycolytic flux to maintain ATP supply despite low yield, tolerance of fermentation end-products (ethanol, acetaldehyde) that can reach toxic concentrations, avoidance of cytoplasmic acidification from lactate accumulation, and conservation of carbohydrate reserves that are rapidly depleted by inefficient fermentative metabolism. Anaerobic_Metabolism_Disposition varies among genotypes in the inducibility and activity of fermentative enzymes (ADH, PDC), the balance between ethanol and lactate production (ethanol pathway generally less acidifying), the capacity to export or metabolize accumulated fermentation products upon re-aeration, and the efficiency of ATP utilization enabling function at reduced energy supply.
Anaerobic_Metabolism_Disposition is_a Waterlogging_Tolerance_Disposition. Fermentative metabolism provides ATP when oxygen is limiting but is inherently inefficient (2-3 vs 36-38 ATP/glucose). ADH and PDC are key fermentative enzymes, with inducibility varying among genotypes. Ethanol pathway preferred over lactate to avoid cytoplasmic acidification. Carbohydrate reserve depletion is a major constraint under extended anoxia. Recovery upon re-aeration depends on capacity to clear accumulated end-products and restore aerobic metabolism.
Anaerober-Stoffwechsel-Disposition
Fermentativer-Metabolismus
Hypoxie-Stoffwechsel
anaerobic respiration capacity
fermentative metabolism
hypoxic metabolism
Drew MC (1997) Oxygen deficiency and root metabolism: injury and acclimation under hypoxia and anoxia, Annual Review of Plant Physiology and Plant Molecular Biology 48:223-250 Gibbs J, Greenway H (2003) Mechanisms of anoxia tolerance in plants. I. Growth, survival and anaerobic catabolism, Functional Plant Biology 30:1-47 Ismond KP et al. (2003) Enhanced low oxygen survival in Arabidopsis through increased metabolic flux in the fermentative pathway, Plant Physiology 132:1292-1302
Anaerobic Metabolism Disposition
The ancient name Aminea documented by Virgil in the Georgics (29 BCE), Columella in De Re Rustica, Pliny the Elder, and Cato the Elder designates what was considered the noblest wine grape of Roman Italy, with five sub-varieties distinguished (Aminea Gemella Major, Aminea Gemella Minor, Aminea Lanata, Aminea Lucana, Aminea Peregrina) — modern scholars have proposed Greco di Tufo and Aglianico as possible descendants based on geographic continuity in Campania and Basilicata, but 2,000 years of clonal mutation, seedling propagation, and cultivar displacement make definitive molecular identification impossible.
A historical name documented in sources from classical antiquity or earlier periods — including Greco-Roman agricultural treatises, Egyptian papyri, Near Eastern cuneiform tablets, and archaeological inscriptions — designating a grapevine cultivar or grape type whose identity predates medieval European viticultural records and typically cannot be resolved to a modern cultivar with certainty.
Ancient_Name captures cultivar designations from the pre-medieval period where the gap between historical documentation and modern molecular identification is greatest. Ancient authors including Cato (De Agri Cultura, c. 160 BCE), Virgil (Georgics, 29 BCE), Columella (De Re Rustica, c. 65 CE), and Pliny (Naturalis Historia, 77 CE) documented over 90 named grape varieties in the Roman Empire: the relationship between these ancient names and modern cultivars remains one of the most debated questions in viticultural history. Archaeobotanical evidence (charred seeds, waterlogged pips, pollen analysis) provides complementary morphometric data but cannot resolve cultivar identity to the level achievable with living plant material and molecular markers.
Sortenbezeichnung der Antike
ancient cultivar designation
antiker Sortenname
antiquity grape name
classical variety name
klassischer Rebname
Columella (c. 65 CE) De Re Rustica Book III Pliny the Elder (77 CE) Naturalis Historia Book XIV McGovern PE (2003) Ancient Wine: The Search for the Origins of Viniculture, Princeton University Press pp: 195-248 Bacilieri R et al. (2017) Potential of combining morphometry and ancient DNA for grapevine identification, Scientific Reports 7: 18098
Ancient Name
Mourvèdre ancient old own-rooted trunk in Bandol planted 1895 (131 years old) shows 380-450 mm diameter with 85% hollowed heartwood, 12-18 mm functional sapwood conducting minimal water, 35-65 g starch reserves, produces 0.8-1.8 kg exceptionally concentrated fruit from 3-8 sparse clusters, requires structural support and intensive care, represents irreplaceable pre-phylloxera genetic heritage producing distinctive age-expressive wines
An ancient old own-rooted trunk is an old own-rooted trunk of exceptional age (typically 50-150+ years post-planting) that has survived multiple decades beyond typical commercial vineyard lifespan, characterized by massive trunk diameter, extensive wood decay or hollowing, severely declining vigor, minimal productivity, and exceptional historical, genetic, or cultural significance despite marginal commercial value. Ancient old own-rooted trunk occurrence is limited to= phylloxera-free regions where own-rooted Vitis vinifera can achieve extreme longevity (pre-phylloxera European vineyards planted pre-1860s, Chile, parts of Australia and Argentina), phylloxera-resistant cultivars on their own roots (certain Vitis vinifera selections showing tolerance), exceptional sites supporting extreme longevity (deep fertile soils, optimal climates, intensive care), and heritage vineyards deliberately preserved (historical sites, genetic repositories, cultural landmarks). Ancient old own-rooted trunk anatomical characteristics include= very large trunk diameter (often 200-500+ mm at 80-150 years, exceptional specimens exceeding 500-800 mm), massive heartwood often extensively decayed or hollow (central 60-90% of trunk showing wood decay, complete hollowing in many specimens), minimal functional sapwood (10-20 mm radial depth providing limited water transport capacity), extremely irregular growth rings (recent rings 0.2-2 mm width, frequent missing rings in stress years), very thick rough bark (10-20+ mm with deep fissures and extensive cork accumulation), and gnarled irregular trunk morphology (twisted, multiple stems, burls, and other growth anomalies from decades of pruning, damage, and regrowth). Ancient old own-rooted trunk physiological status shows= severely reduced water transport capacity (limited sapwood providing minimal hydraulic conductance, frequent xylem dysfunction), depleted carbohydrate reserves (starch storage capacity reduced to 30-80 g compared to 120-180 g in prime vines), minimal vegetative growth (annual shoot growth 20-60 cm compared to 80-150 cm in vigorous mature vines), extended and irregular phenology (budbreak, flowering, véraison delayed and asynchronous), and extreme stress sensitivity (drought, heat, cold, or disease readily causing vine decline or death). Ancient old own-rooted trunk productivity characteristics include= very low yields (often 0.5-2.5 kg per vine, 10-30% of prime mature vine production), sparse cluster number (1-10 clusters per vine from limited fruitful shoots), very small clusters and berries (cluster weight 40-90 g, berry diameter 8-12 mm compared to 12-16 mm in vigorous vines), extremely concentrated fruit (potentially exceptional quality with very high skin/pulp ratios, concentrated flavors and phenolics), and high year-to-year variation (productivity strongly affected by vintage conditions, ranging from near-zero to moderate crops). Ancient old own-rooted trunk disease and decay includes= near-universal presence of wood decay organisms (white rot, brown rot fungi extensively colonizing heartwood), trunk diseases present in majority of vines (Esca, Eutypa, Botryosphaeria causing cankers, dieback, and wood deterioration), structural instability from extensive hollowing (risk of trunk collapse or breakage), but continued survival through minimal functional sapwood (vines maintaining life through 10-25 mm sapwood layer despite massive internal decay). Ancient old own-rooted trunk viticulture significance includes= exceptional genetic value (ancient vines may represent rare heritage selections, pre-clonal diversity, unique terroir adaptations accumulated over decades), historical and cultural significance (living connection to viticulture heritage, some vines planted 1700s-1800s), potential exceptional wine quality (extremely concentrated fruit producing distinctive wines commanding premium prices), research value (studying vine longevity, wood decay processes, long-term terroir adaptation), and preservation imperative (irreplaceable genetic and historical resources requiring conservation efforts). Ancient old own-rooted trunk management is highly specialized= minimal interventions (avoiding stress from excessive pruning or crop load), disease management accepting trunk decay (focusing on preventing catastrophic decline rather than curing diseases), structural support (staking or propping trunks at risk of collapse), gentle crop thinning (removing excess clusters to prevent vine stress), protective irrigation (minimal water during critical periods preventing terminal drought stress), and succession planning (establishing propagation from ancient vines preserving genetics before inevitable vine death).
Ancient_Old_Own_Rooted_Trunk emphasizes exceptional longevity and is distinguished from Old_Own_Rooted_Trunk (typical senescence 25-50 years) and Multi_Decade_Old_Own_Rooted_Trunk (50-80 years) by extreme age 50-150+ years and associated exceptional characteristics. Critical for heritage vineyard preservation, understanding extreme vine longevity, genetic conservation of rare selections, and appreciation of historical viticultural significance.
Century-old trunk
Jahrhundertealter Eigenwurzelstamm
Pre-phylloxera trunk
Prä-Phylloxera Rebstamm
Uralter wurzelechter Stamm
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Robinson, J., Harding, J., & Vouillamoz, J. (2012). Wine Grapes= A Complete Guide to 1,368 Vine Varieties, Including Their Origins and Flavours. Ecco.
Ancient Old Own Rooted Trunk
An ancient wood latent bud on the main trunk of a 40-year-old Vitis vinifera cv. Grenache vine in Châteauneuf-du-Pape, estimated at 15-20+ years suppression based on trunk growth rings and bark accumulation, unexpectedly activated following severe wildfire damage that killed all above-ground canopy and cordons, producing a single weak shoot 65cm in length during the first post-fire season, with the shoot showing chlorotic leaves and slow growth but eventually establishing sufficient vigor to serve as the foundation for complete vine reconstruction over subsequent four seasons, demonstrating the remarkable but unpredictable regenerative capacity of ancient latent bud positions.
An old wood latent bud that has remained suppressed on perennial wood structures for very extended periods, typically 8+ years and potentially decades, characterized by very deep embedment within thick accumulated bark and periderm, questionable viability due to extended dormancy and potential meristem senescence, low spontaneous activation probability, and uncertain developmental capacity if released from suppression. Ancient wood latent buds represent the extreme limit of latent bud survival in grapevine perennial structures, occasionally producing shoots from apparently dead wood sections following catastrophic damage, demonstrating remarkable meristematic persistence but with highly variable vigor and success rates, serving as last-resort regenerative capacity when all other bud positions have failed or been eliminated by damage.
Ancient wood latent buds challenge conventional understanding of meristem longevity and dormancy limits, with documented cases of bud breakage from wood estimated at 30+ years old. However, success rates are highly variable and unpredictable, with many ancient buds failing to produce viable shoots or producing weak shoots that fail to establish. The boundary between viable ancient latent buds and truly dead meristematic tissue is unclear and likely exists on a continuum. Viticulturists generally should not depend on ancient wood latent buds for planned renewal programs, but their occasional successful activation provides unexpected recovery opportunities in severely damaged vineyards, particularly in old-vine plantings where preservation of original genetic material and root systems justifies extended recovery efforts.
decades-old latent bud
sehr alte Adventivknospe
uralte Holzknospe
very old wood latent bud
Based on Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Krstic et al. (2003) "Altered wine composition and quality due to vineyard fire smoke damage to grapes" Australian Journal of Grape and Wine Research 9=24-29 Whiting & Lang (2004) "Xylem dysfunction explains tree growth decline with increased tree age" Tree Physiology 24=337-345.
Ancient Wood Latent Bud
The aneuploid status of individuals in an interspecific F1 population from a Vitis vinifera cv. Gewürztraminer × V. amurensis cross was documented by Reisch et al. (2012) through flow cytometry screening of 287 seedlings — 8 individuals (2.8%) showed DNA content values deviating by 2-5% from the expected diploid value, and subsequent chromosome counting confirmed 5 monosomic plants (2n = 37, missing one chromosome from different linkage groups) and 3 trisomic plants (2n = 39, carrying an extra copy of specific chromosomes). The monosomic plants showed reduced vigor (30-50% less shoot growth than diploid siblings), poor fruit set (60% in diploid siblings), and characteristic phenotypic abnormalities specific to the missing chromosome, while trisomic plants showed milder growth reduction (10-20% less vigor) and partially reduced fertility, consistent with the gene dosage imbalance effects of aneuploidy.
A ploidy information record documenting the aneuploid chromosome complement in a specific Vitis accession, recording the deviation from the euploid number by the gain or loss of individual chromosomes (2n = 37, 39, or other non-euploid counts) that produces an unbalanced genome with dosage imbalances for the genes on the affected chromosome(s), typically resulting in reduced vigor, fertility defects, and developmental abnormalities that limit the viticultural utility of aneuploid individuals.
Aneuploid_Status documents a cytogenetic condition that is rare in vegetatively propagated cultivars (where mitotic stability maintains the diploid complement through clonal generations) but occurs at low frequency in seedling populations from both intraspecific and interspecific crosses. Aneuploidy arises primarily through meiotic non-disjunction — the failure of homologous chromosomes to separate correctly during gamete formation, producing gametes with n+1 or n-1 chromosomes that, upon fertilization with normal gametes, yield trisomic (2n+1 = 39) or monosomic (2n-1 = 37) offspring: non-disjunction rates are elevated in interspecific crosses (where homeologous rather than fully homologous chromosome pairing occurs) and in crosses involving structurally heterozygous parents (carrying inversions, translocations, or other rearrangements that interfere with normal meiotic segregation). Aneuploid grapevines are of limited commercial interest due to their reduced vigor and fertility, but they serve as valuable cytogenetic tools for gene mapping — monosomic lines (missing a specific chromosome) enable assignment of unmapped markers and traits to specific chromosomes through hemizygous expression analysis.
Aneuploidie-Status
Chromosomenzahl-Abweichung
aneuploid confirmation
aneuploider Chromosomensatz
chromosome number aberration record
non-euploid status
Reisch BI et al. (2012) Grape breeding and genetics, in Genetics, Genomics, and Breeding of Grapes, Science Publishers pp: 3-60 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519
Aneuploid Status
Field investigation of Anisohydric_Behavior_Disposition in the Barossa Valley compared 'Syrah' (anisohydric) and 'Grenache' (isohydric) under natural drought conditions across two contrasting seasons (average and severe drought). Midday measurements of leaf water potential, stomatal conductance, and photosynthesis were conducted at 2-week intervals throughout the growing season, with concurrent soil moisture monitoring. 'Syrah' displayed pronounced Anisohydric_Behavior_Disposition with Ψmd declining from -1.0 MPa in well-watered conditions to -2.2 MPa during peak drought in the severe year, while 'Grenache' maintained Ψmd between -1.2 and -1.5 MPa regardless of conditions. At equivalent soil water content (15% volumetric), 'Syrah' sustained stomatal conductance of 120 mmol m⁻² s⁻¹ and photosynthesis of 8.5 μmol CO₂ m⁻² s⁻¹, while 'Grenache' had reduced to 45 mmol m⁻² s⁻¹ and 3.2 μmol CO₂ m⁻² s⁻¹. Cumulative seasonal carbon assimilation was 40% higher in 'Syrah' during the average drought year. However, the severe drought year revealed the risks of Anisohydric_Behavior_Disposition, with 'Syrah' showing 35% loss of hydraulic conductivity from cavitation (measured by percentage loss of conductivity, PLC), visible leaf desiccation affecting 15% of canopy, and incomplete recovery the following season. 'Grenache' showed only 8% PLC and full recovery. Interestingly, fruit quality parameters diverged, with 'Syrah' berries showing higher anthocyanin concentration (likely stress-induced) but also elevated alcohol potential from accelerated sugar accumulation. These observations illustrate how Anisohydric_Behavior_Disposition maximizes productivity in moderate drought but incurs greater risk under severe conditions.
An Anisohydric_Behavior_Disposition is a Stomatal_Regulation_Disposition that inheres in a Vitis vinifera plant and is realized through loose stomatal control that allows midday leaf water potential to decline progressively with decreasing soil water availability, prioritizing continued carbon assimilation over strict maintenance of tissue hydration during drought. This disposition represents the productivity-oriented end of the stomatal regulation spectrum, with anisohydric cultivars maintaining substantial stomatal opening and gas exchange as drought develops, accepting lower leaf water potential as a consequence of continued transpiration against increasing soil-to-leaf water potential gradient. The term "anisohydric" (Greek: unequal water) refers to the variable leaf water status that results from this strategy, with midday water potential tracking soil water potential rather than being defended at a stable set-point. Anisohydric behavior involves reduced sensitivity to ABA, higher stomatal conductance at equivalent water potential, and potentially greater reliance on hydraulic signals versus chemical signals in stomatal regulation. The adaptive value of Anisohydric_Behavior_Disposition lies in maximizing carbon gain during periods of moderate water deficit, supporting growth and fruit development when isohydric cultivars would severely curtail photosynthesis. However, this aggressive strategy incurs risks of hydraulic failure (cavitation) and tissue damage under severe or prolonged drought when water potential declines below critical thresholds. Classic anisohydric grapevine cultivars include 'Syrah' (Shiraz) and 'Semillon'.
Anisohydric_Behavior_Disposition is_a Stomatal_Regulation_Disposition. Anisohydric behavior allows leaf water potential to decline with soil drying, maintaining gas exchange at the cost of tissue hydration. The mechanism involves reduced ABA sensitivity and/or delayed ABA accumulation compared to isohydric cultivars. Anisohydric cultivars are favored in environments with moderate, predictable drought and available irrigation. Risks include hydraulic failure and tissue damage under severe drought. 'Syrah/Shiraz' and 'Semillon' are classic anisohydric cultivars.
Anisohydrisches-Verhalten-Disposition
Opportunistische-Wassernutzung
Variable-Wasserpotential-Strategie
anisohydric stomatal behavior
opportunistic water use
risk-taking water strategy
Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought, Plant Cell and Environment 26:1393-1405 Hochberg U et al. (2013) Stomatal regulation of anisohydric grapevine (Vitis vinifera cv. Syrah) during drought stress, Annals of Botany 112:1595-1606 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data, Annals of Botany 105:661-676
Anisohydric Behavior Disposition
Anisohydric behavior quality characterization in Barossa Valley Shiraz documented the productivity-oriented drought response underlying this cultivar's reputation for sustained performance under Australian conditions. Under progressive soil drying, Shiraz exhibited pronounced anisohydric behavior quality with midday leaf water potential declining from -0.9 MPa at field capacity through -1.4 MPa at moderate soil moisture deficit to -1.9 MPa under severe stress, a 1.0 MPa range contrasting with the 0.2 MPa variation typical of isohydric cultivars under equivalent conditions. Stomatal conductance remained at 65% of maximum at moderate stress levels when isohydric Grenache had already declined to 35%, enabling Shiraz to maintain correspondingly higher photosynthesis and carbon accumulation during the critical ripening period. The anisohydric behavior quality of Shiraz proved advantageous for fruit quality under managed deficit irrigation, with sustained photosynthesis supporting complete ripening even under moderate stress, but created vulnerability under severe stress events where the declining water potential exceeded hydraulic safety margins causing 15% shoot dieback in an extreme drought vintage that left isohydric cultivars undamaged. The anisohydric behavior quality characterization informed management recommendations targeting stress levels within the cultivar's tolerance capacity while exploiting its productivity advantage over more conservative types.
Anisohydric behavior quality is a stomatal response to drought quality that inheres in a grapevine cultivar or clone and represents the drought response strategy characterized by less sensitive stomatal regulation that permits leaf water potential to decline substantially as soil water availability decreases, maintaining stomatal opening and photosynthetic carbon assimilation even as water status falls to levels that would trigger protective closure in isohydric cultivars, representing a risk-tolerant strategy that prioritizes productivity over hydraulic safety under developing water deficit conditions. This quality captures the opposite endpoint of the isohydric-anisohydric behavioral continuum from isohydric behavior, with strongly anisohydric cultivars exhibiting looser stomatal control that sustains carbon assimilation at the cost of declining water potential and increased risk of hydraulic damage if stress exceeds the cultivar's tolerance capacity. The ontological significance of anisohydric behavior quality lies in its characterization of a productivity-oriented drought adaptation strategy with specific implications for vine performance and risk exposure, where anisohydric cultivars may achieve greater carbon assimilation under moderate stress but face elevated risk of cavitation-induced hydraulic damage under severe stress that exceeds their tolerance capacity. The biological basis of anisohydric behavior involves reduced sensitivity of stomatal guard cells to drought-signaling molecules and hydraulic cues, permitting continued stomatal opening despite declining leaf water status, combined with hydraulic architecture characteristics that tolerate lower water potentials without catastrophic failure, including greater xylem cavitation resistance and the capacity to refill embolized vessels. The characteristic feature of anisohydric behavior is the progressive decline in midday leaf water potential as soil dries, tracking soil water availability rather than maintaining constant values, with the diagnostic pattern of variable water potential across conditions distinguishing anisohydric from isohydric strategies.
Anisohydric_Behavior_Quality is_a Stomatal_Response_To_Drought_Quality, representing the risk-tolerant drought strategy allowing leaf water potential to vary with conditions while maintaining stomatal opening. This class addresses the opposite endpoint of the behavioral spectrum from Isohydric_Behavior_Quality. The diagnostic feature is declining midday water potential (often 1.0+ MPa range) tracking soil moisture. Syrah/Shiraz is the classic anisohydric Vitis vinifera cultivar. The strategy sustains productivity under moderate stress but creates vulnerability to hydraulic damage under severe drought, requiring management within cultivar-specific tolerance limits.
Wasser-verbrauchende Strategie
anisohydric strategy
anisohydrisches Verhalten
productivity-oriented behavior
produktivitätsorientierte Strategie
variable water potential
variables Wasserpotential
water-spending strategy
Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell and Environment 26:1393-1405 Soar CJ et al. (2006) Grape vine varieties Shiraz and Grenache differ in their stomatal response to VPD: apparent links with ABA physiology and gene expression in leaf tissue. Australian Journal of Grape and Wine Research 12:2-12 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data. Annals of Botany 105:661-676
Anisohydric Behavior Quality
Annual precipitation for Barossa Valley, South Australia averages 450-500 mm, classifying it among the world's drier premium wine regions and necessitating supplemental irrigation for most cultivars. Mendoza, Argentina records only 200-250 mm annually, with virtually all vineyard water supplied by Andean snowmelt irrigation through historic canal systems. By contrast, the Vinho Verde region of northern Portugal receives 1200-1500 mm annually, requiring vigorous canopy management and disease-resistant training systems to manage the excess moisture.
Annual precipitation is a precipitation data entity that records the cumulative total of all atmospheric water deposition (rain, snow, hail, sleet, and dew) received at a viticultural site over a complete calendar or hydrological year, expressed in millimeters (mm), providing the baseline hydrological characterization of a vineyard's water input regime.
Annual precipitation serves as the coarsest but most widely cited hydrological descriptor for viticultural regions, with the global range for quality wine production spanning roughly 250-1500 mm. However, annual totals alone are insufficient for viticultural characterization, as the seasonal distribution, rainfall intensity, and soil water-holding capacity fundamentally modulate the vine's effective water supply. Regions receiving identical annual precipitation can support radically different viticultural outcomes — 600 mm concentrated in winter dormancy (Mediterranean climate) creates very different growing conditions than 600 mm distributed uniformly throughout the year (oceanic climate). The trend toward higher annual precipitation variability under climate change increases the importance of tracking multi-year precipitation statistics rather than long-term averages alone.
Gesamtjahresniederschlag
Jahresniederschlag
Jahresniederschlagsmenge
annual rainfall total
cumulative annual precipitation
jährliche Niederschlagssumme
total annual rainfall
yearly precipitation
Gladstones J (2011) Wine, Terroir and Climate Change, Wakefield Press, pp.: 119-132 Jackson RS (2020) Wine Science: Principles and Applications, 5th Ed, Academic Press, pp.: 56-71 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press, pp.: 327-345
Annual Precipitation
Annual production cost for Grüner Veltliner in the Wachau (2023 estimates, premium Smaragd-quality production) approximately €8,000-12,000/ha, reflecting the high labor intensity of steep terraced vineyards (50-60% of total cost): winter pruning €1,200-1,800/ha (hand pruning in narrow terrace rows, 40-60 hours/ha), canopy management €1,500-2,500/ha (multiple passes for shoot thinning, positioning, leaf removal, hedging), spray program €800-1,200/ha (12-16 applications for powdery and downy mildew management, with higher per-hectare costs on steep slopes requiring small-equipment application), crop thinning €500-800/ha (green harvest to achieve Smaragd quality concentration), soil management €400-600/ha (cover crop mowing, limited tillage on terraces, manual weed control around vine trunks), machinery and equipment €800-1,200/ha (depreciation and operating costs for specialized narrow-track equipment suited to terrace viticulture), and overhead €800-1,200/ha (insurance, land tax, Vinea Wachau membership, organic certification fees where applicable). Annual production cost for Cabernet Sauvignon in Coonawarra (2023 estimates, premium production) approximately AUD 6,000-9,000/ha reflecting lower labor costs through mechanization of several operations, but higher irrigation infrastructure and water costs specific to Australian conditions.
Annual production cost is a cost of production entity that documents the recurring operational expenditures of managing a mature, commercially productive vineyard through a single growing season, encompassing labor costs (the single largest component, including permanent and seasonal workforce for pruning, canopy management, crop thinning, and general vineyard operations), materials and inputs (spray materials for pest and disease management, fertilizers, soil amendments, fuel and energy), machinery and equipment costs (depreciation, maintenance, and operating costs for tractors, sprayers, mowing equipment, and specialized viticultural machinery), overhead and administrative costs (insurance, land tax, regulatory compliance, record-keeping, certification fees), and contracted service costs (specialized operations performed by external contractors — mechanical pruning, aerial or precision spraying, soil sampling), expressed per hectare per year with specification of the production system, labor cost environment, and viticultural intensity level.
Annual production cost documentation captures the financial reality of vineyard management — the year-after-year operational expenditures that must be covered by grape revenue to sustain viable production, and that vary substantially across cultivars, regions, production systems, and quality ambition levels. The labor cost component dominates annual production cost in most viticultural contexts, typically representing 40-65% of total per-hectare costs, and is the component most sensitive to regional variation — labor rates for viticultural workers range from €3-5/hour in North Africa and Eastern Europe to €15-20/hour in Central Europe to €25-35/hour in Australia, the US, and Switzerland, creating order-of-magnitude differences in the labor cost component across regions. The cultivar-specific dimension of annual production cost reflects the management requirements inherent to each variety — disease-susceptible cultivars (Pinot Noir's susceptibility to botrytis, Müller-Thurgau's mildew vulnerability) require more intensive spray programs increasing materials and application costs, vigorous cultivars requiring extensive canopy management incur higher labor costs, and cultivars demanding meticulous crop thinning for quality (Pinot Noir, Sangiovese) add additional seasonal labor requirements. The production system dimension creates significant cost differentiation — conventional viticulture minimizes labor through herbicide weed control and efficient synthetic spray programs, organic viticulture substitutes mechanical weed control (higher labor and fuel costs) and biological/mineral spray materials (more frequent applications, higher per-application labor cost), and biodynamic viticulture adds preparation costs (500, 501, and compost preparations) and increased management attention, with the total cost differential between conventional and certified organic typically 15-30% and biodynamic 20-40% above conventional. The relationship between annual production cost and wine quality is complex — higher per-hectare expenditures generally correlate with higher quality through more intensive management, but the relationship is non-linear, with diminishing returns at the highest investment levels and the possibility of excessive intervention actually reducing wine quality.
Betriebskosten
Jahreskosten
annual operating expenditure
annual vineyard cost
jährliche Produktionskosten
jährlicher Betriebsaufwand
laufende Bewirtschaftungskosten
operating cost
recurring production cost
vineyard management cost
LWK Burgenland (2023) Weinbaukalkulation — Betriebskosten Weingarten UC Davis (2023) Sample costs to produce wine grapes, Department of Agricultural and Resource Economics Wine Australia (2023) Vineyard benchmarking studies Eurostat (2023) Agricultural production costs — vineyards
Annual Production Cost
Cabernet Sauvignon annual ring at year 12 post-planting shows 2.8-4.2 mm radial width, distinct ring-porous anatomy with earlywood vessels 180-260 μm diameter occupying 40-48% of earlywood zone (formed April-June), latewood vessels 50-110 μm diameter with higher fiber content (formed July-September), clear ring boundary marked by transition to next year's large earlywood vessels, and width indicating favorable growing conditions compared to preceding year (previous ring 2.2-3.4 mm)
An annual ring is a layer of secondary xylem produced by vascular cambium during a single growing season in Vitis vinifera trunk, comprising distinct earlywood and latewood zones that form a visible concentric band in trunk cross-section, providing dendrochronological record of vine growth history and environmental conditions experienced during each year of trunk development. Annual ring formation occurs through seasonal cambial activity= cambium initiates cell division in spring (March-April in Northern Hemisphere temperate climates) producing earlywood with large vessels optimized for water transport, cambium continues division through summer (May-August) with transition to latewood containing smaller vessels and more fibers, cambial activity ceases in fall (September-October) marking ring boundary, and dormant period (November-February) precedes next spring when new ring formation begins. Annual ring anatomy in Vitis vinifera exhibits ring-porous pattern= earlywood (inner portion of ring formed April-June) characterized by large-diameter vessels (120-300 μm) occupying 35-55% of earlywood area providing majority of hydraulic conductance, latewood (outer portion formed July-September) containing smaller vessels (40-120 μm) and higher proportion of fibers providing mechanical strength, distinct boundary between rings marked by transition from small latewood vessels of one year to large earlywood vessels of next year, and vessels arranged in radial or slightly tangential patterns with ray parenchyma providing radial organization. Annual ring width varies dramatically by vine age, vigor, and environmental conditions= young vigorous vines (years 1-5) producing wide rings (3-8 mm radial width reflecting rapid growth and large vessel formation), mature moderate-growth vines (years 8-25) producing intermediate rings (1.5-4 mm width reflecting stable productivity), old declining vines (years 30-50+) producing narrow rings (0.5-2.5 mm width reflecting reduced vigor), and environmental factors causing year-to-year variation (favorable years with adequate water and nutrition producing wider rings, stress years with drought or disease producing narrow rings). Annual ring dendrochronological information includes= ring width recording annual growth rate (wide rings indicating favorable conditions or high vigor, narrow rings indicating stress or declining health), missing rings potentially occurring in severe stress years (cambium failing to produce detectable xylem in extreme drought or defoliation), false rings occasionally forming (mid-season stress followed by recovery causing intra-annual density variation mimicking true ring boundary), and ring patterns enabling cross-dating (matching ring width sequences across multiple vines or sites reconstructing environmental history). Annual ring vessel characteristics vary by position within ring= earlywood vessels 150-300 μm diameter in outer rings (sapwood), 120-250 μm in inner rings (transitioning to heartwood), arranged in 1-3 rows providing maximum conductance, latewood vessels 40-120 μm diameter distributed through outer ring portion, total vessel density 25-60 vessels/mm² varying by ring width and growth conditions, and vessel lumen occupying 30-50% of total ring cross-sectional area. Annual ring fiber and parenchyma composition includes= fiber cells providing mechanical strength (50-65% of ring volume, thick lignified walls supporting trunk structure), axial parenchyma distributed among vessels (8-15% of volume providing storage and lateral transport), ray parenchyma crossing radially through ring (3-8% of volume connecting xylem to phloem and cambium), and relative proportions varying by ring position (earlywood higher vessel and parenchyma content, latewood higher fiber content). Annual ring formation timing and duration includes= ring initiation synchronized with budbreak (cambial reactivation 0-2 weeks before bud opening, triggered by rising temperatures and hormonal signals from expanding buds), earlywood formation period 6-10 weeks (April-June producing large vessels supporting initial canopy expansion), latewood formation period 8-14 weeks (June-September producing smaller vessels and fibers), and total ring formation duration 14-24 weeks depending on climate (longer growing seasons producing wider rings with extended latewood formation). Annual ring boundaries and identification includes= distinct boundary marked by abrupt transition from small latewood vessels to large earlywood vessels (creating visible line in cross-section), boundary corresponds to dormant period (cambial inactivity during winter creating temporal separation between successive rings), boundary clarity varying by species and conditions (Vitis vinifera showing moderate to high boundary distinctness facilitating ring counting), and boundary sometimes indistinct in very narrow rings or continuous growth conditions (tropical climates or greenhouse conditions potentially producing ambiguous boundaries). Annual ring counting and aging includes= rings counted from pith outward to determine trunk age (ring number corresponding to years post-cutting propagation or years post-grafting for grafted trunks), pith position sometimes offset from geometric center (eccentric growth causing unequal ring widths on opposite sides), missing pith requiring estimation (oldest surviving ring indicating minimum age but potentially underestimating true age), and ring count providing accurate age determination (±1 year accuracy typical for well-preserved temperate-grown trunks). Annual ring significance in viticulture includes= recording vine growth history (width patterns revealing favorable and stress years throughout vine life), enabling age determination (ring counting providing precise vine age for old vine designation or vineyard records), indicating vigor changes (declining ring width suggesting reduced growth or approaching senescence), providing environmental reconstruction (ring width correlating with precipitation, temperature, and management practices), and informing management decisions (ring patterns helping assess vine health, stress impacts, and appropriate cultural practices).
Annual_Ring emphasizes yearly growth increment and is distinguished from specific ring types (Narrow_Annual_Ring, Wide_Annual_Ring, Growth_Year_Annual_Ring) by encompassing all annual xylem production regardless of width or characteristics. Critical for understanding trunk diameter development, dendrochronology of vine growth, environmental effects on wood formation, and aging methodology for old vines.
Growth ring
Holzring
Jahresring
Tree ring
Wachstumsring
Xylem ring
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Schweingruber, F. H. (1996). Tree Rings and Environment= Dendroecology. Paul Haupt Publishers.
Annual Ring
Annual yield stability assessment across 10 vintages in Chardonnay reveals the site-dependent production reliability, with sheltered site showing high stability (mean 7.2 t/ha, range 5.8-8.4, CV 11%, no vintage below economic threshold), while exposed site shows low stability (mean 6.8 t/ha similar but range 2.1-10.5, CV 38%, three vintages with severe frost or disease losses), demonstrating how site characteristics affect annual stability independently of mean yield, with implications for risk management and contract reliability.
Annual yield stability quality is a yield consistency quality that represents the degree to which grape production remains constant from year to year, characterizing the temporal reliability of vineyard output that determines how predictably production targets can be achieved across vintages despite the inevitable variation in weather, pest pressure, and other season-specific factors. Within the BFO framework, this quality specifically addresses the inter-annual dimension of yield consistency, distinguishing year-to-year variation from spatial variation while recognizing that annual stability reflects the vineyard's resilience to seasonal fluctuations—highly stable vineyards buffering environmental variation to maintain consistent output, while unstable vineyards amplifying seasonal differences into large yield swings. The ontological significance lies in annual yield stability determining the reliability of production for business planning, where stable yields enable confident forward contracting, consistent winery operations, and predictable cash flows, while unstable yields create boom-and-bust cycles that complicate inventory management, strain buyer relationships, and introduce financial volatility that affects long-term viability. The factors creating annual yield variation include weather effects on fruit set (frost, rain during flowering, heat stress reducing berry numbers), weather effects on berry development (drought reducing berry size, heat events affecting ripening), pest and disease pressure variation among years (Botrytis in wet years, mildew epidemics), and carry-over effects from previous seasons (depleted reserves from overcropping affecting subsequent fruitfulness). The management approaches for improving annual stability include reserve management (maintaining adequate carbohydrate reserves to buffer stress years), crop load management (consistent pruning and thinning practices regardless of apparent potential), irrigation management (supplemental water to buffer drought years), and pest management (consistent protection preventing epidemic losses in high-pressure years). Understanding annual yield stability quality enables assessment of production risk, identification of factors driving year-to-year variation, and management strategies targeting improved temporal consistency.
Annual_Yield_Stability_Quality addresses the year-to-year consistency of production affecting business planning reliability. This quality reflects vineyard resilience to seasonal variation. The class supports risk assessment and production forecasting through annual stability characterization.
Jahrgangsstabilität
annual production constancy
jährliche Ertragsstabilität
jährliche Produktionskonstanz
temporal yield reliability
vintage stability
year-to-year consistency
zeitliche Ertragsreliabilität
Clingeleffer PR (2001) Crop development, crop estimation and crop control to secure quality and production of major wine grape varieties: a national approach. Final report to Grape and Wine Research and Development Corporation. CSIRO Plant Industry, Merbein Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 145-230
Annual Yield Stability Quality
Merlot hermaphrodite flowers at BBCH 61-63 (beginning to full bloom) exhibit anthers 0.7-0.85 mm length with bright yellow coloration, longitudinal dehiscence slits opening along both lobes releasing copious yellow pollen visible as clouds when flowers are disturbed, with each anther producing approximately 4,000-6,000 pollen grains totaling 20,000-30,000 grains per five-anther flower, achieving efficient self-pollination. Pinot Noir at BBCH 65 shows fully dehisced anthers with opened longitudinal slits and partially depleted pollen contents creating lighter yellow to cream coloration, with dehiscence having occurred 6-24 hours previously and majority of pollen already released and dispersed. Syrah experiencing cool temperatures (12-14°C) during attempted bloom shows delayed anther dehiscence with mature yellow anthers remaining closed 24-48 hours after calyptra shedding, eventually achieving partial dehiscence when temperatures increase to 18-20°C but with reduced pollen release efficiency contributing to decreased fruit set (42-48% versus normal 54-62%). Gewürztraminer functionally female flowers exhibit small anthers 0.4-0.55 mm with reduced pollen content producing only 500-1,500 grains per anther (<5,000 total per flower), showing reflex positioning below pistil and minimal functional significance for reproduction despite morphological presence and partial dehiscence.
An anther is the terminal pollen-producing structure of the stamen comprising two lobes (thecae) positioned on opposite sides of the filament apex, with each lobe containing two pollen sacs (microsporangia) where microsporogenesis and pollen maturation occur, culminating in anther dehiscence through longitudinal splitting that releases mature pollen grains for dispersal to receptive stigmas. The anther structure consists of= (1) two elongated lobes (left and right thecae) typically 0.4-1 mm in length and 0.15-0.4 mm in width each, connected by a central connective tissue region continuous with the filament apex (2) four pollen sacs (two per lobe) arranged as two pairs, each sac developing as a microsporangium producing pollen through meiotic and mitotic divisions (3) anther wall comprising multiple tissue layers including epidermis, endothecium (fibrous layer), middle layers, and tapetum (nutritive layer), with the endothecium playing a critical role in dehiscence mechanics through differential cell wall thickening creating hygroscopic tension (4) stomium (dehiscence zone) running longitudinally along each lobe where programmed cell breakdown creates the opening slit for pollen release (5) vascular tissue in the connective providing nutrient supply to developing pollen. Anther development follows a characteristic temporal sequence= early anther primordia (BBCH 53-54) appear as undifferentiated swellings at filament tips anther lobe differentiation (BBCH 54-55) creates bilateral symmetry with four pollen sac initiation microsporogenesis (BBCH 55-57) involves pollen mother cell meiosis producing haploid microspore tetrads within pollen sacs pollen maturation (BBCH 57-59) transforms microspores into mature pollen grains through vacuolation, wall formation, and accumulation of storage reserves anther maturation (BBCH 59-60) involves anther wall tissue maturation, endothecium cell wall thickening, and preparation of dehiscence zones anther dehiscence (BBCH 61-65) through longitudinal stomium splitting releases pollen post-dehiscence senescence (BBCH 65-69) involves anther tissue browning and eventual abscission. Anther size and pollen production capacity vary substantially among flower types= hermaphrodite flower anthers typically 0.6-0.9 mm producing 3,000-7,500 pollen grains per anther (15,000-30,000 per five-anther flower) functionally female flower anthers reduced to 0.3-0.6 mm producing 85% inhibits dehiscence by preventing anther desiccation temperatures 35°C impair dehiscence mechanics rain physically prevents or disrupts dehiscence by maintaining anther moisture and washing released pollen. Anther color provides visual indicators of developmental stage and pollen viability= immature anthers appear green (BBCH 55-57) maturing anthers transition to cream or yellowish (BBCH 58-59) mature anthers at dehiscence are bright yellow due to abundant carotenoid-containing pollen (BBCH 60-65) post-dehiscence anthers fade to tan or brown as pollen depletes and tissue senesces (BBCH 65-69) abnormal anther development or pollen abortion may produce brown or dark brown anthers even at expected dehiscence stages, indicating reproductive failure.
Anther represents the pollen-producing terminal structure of the stamen and must be distinguished from the Filament (stalk component), from Pollen_Sac (internal compartments within anther lobes), and from pollen itself (the dispersed product). The anther is the critical functional component determining pollen production quantity and quality, with anther size, pollen output, and dehiscence efficiency varying dramatically among flower types and strongly influenced by environmental conditions during development and at anthesis. Sub-classification into Mature_Anther, Dehiscing_Anther, and Post_Dehiscence_Anther reflects the critical functional progression from pollen-containing through pollen-releasing to pollen-depleted states. Anther characteristics (size, color, dehiscence timing and completeness) serve as important diagnostic indicators for assessing flower function, bloom progression, environmental stress effects, and reproductive potential.
Anthere
Pollensackträger
Staubbeutel
microsporangium-bearing structure
pollen sac organ
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 101-106 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 94-98 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 187-192.
Anther
Pinot Noir in Burgundy shows anther dehiscence during early-mid June Year N+1 at BBCH 60-65 over 7-9 days, with anthers progressively opening following cap fall predominantly during morning hours 9am-11am when humidity drops below 70%, creating longitudinal slits releasing abundant pollen, under continental conditions of 19-25°C temperature, 55-72% relative humidity with favorable morning decline, typical of this early-to-mid season cultivar showing efficient dehiscence supporting 58-68% fruit set
Anther dehiscence represents the anther opening process where specialized stomium zones undergo structural breakdown creating longitudinal slits enabling pollen liberation from enclosed locules, establishing essential mechanism for male gamete release during flowering. This critical opening process occurs during early-mid flowering in early-mid June in temperate regions Year N+1, typically spanning 5-10 days as anthers progressively dehisce following cap fall and flower opening throughout cluster, corresponding to BBCH 60-65 stages. Dehiscence mechanism involves coordinated structural and physiological processes including anther wall maturation preparing opening, stomium zone activation triggering rupture, and mechanical forces driving slit formation enabling pollen access to exterior. Anther wall structure includes multiple tissue layers with outer epidermis providing protection, middle fibrous layer with hygroscopic properties creating mechanical tension, inner tapetum layer having degenerating by dehiscence, and endothecium cells with thickened walls except at stomium providing structural framework. Stomium zone represents specialized tissue region extending longitudinally along anther where cells lack wall thickening making tissue mechanically weak, predetermined rupture site positioned to enable effective pollen release, and cell breakdown mechanism involving enzymatic wall degradation creating opening. Anther maturation preceding dehiscence shows progressive color change from green toward yellow-brown indicating approaching readiness, tissue dehydration creating mechanical properties enabling rupture, and cytoplasmic changes in stomium cells preparing breakdown. Dehiscence triggering shows strong environmental dependence with humidity decrease causing anther wall dehydration critical for creating mechanical tension, temperature affecting tissue properties and enzymatic activity, and typically occurring during morning hours 8am-12pm as conditions become favorable. Humidity threshold effects show dehiscence typically initiating when relative humidity drops below 70-75% enabling sufficient anther dehydration, high humidity above 75% substantially delaying or preventing dehiscence through maintained tissue hydration, and optimal 45-65% humidity facilitating efficient opening. Temperature effects on dehiscence show optimal 22-30°C supporting rapid efficient opening, cool conditions below 18°C substantially delaying dehiscence through slowed processes and maintained tissue hydration, and warm conditions above 30°C generally facilitating mechanical rupture though potentially affecting pollen viability. Mechanical dehiscence process involves differential tissue shrinkage with hygroscopic fibrous layer contracting as dehydration proceeds creating tension, stomium zone unable to resist mechanical stress due to weak cell walls, and tissue rupture creating longitudinal slit typically 2-4 mm length along anther. Rupture propagation shows progressive slit extension along stomium zone as mechanical stress increases, coordinated opening of both anther locules creating two parallel slits, and complete opening exposing pollen mass to exterior enabling liberation. Pollen liberation following dehiscence proceeds through initial explosive discharge as pressure release ejects pollen bolus from locule creating immediate pollen cloud, continued gravitational release as loosened pollen falls from opened anther over hours, and wind-assisted release as air movement agitates anthers dislodging remaining pollen. Anther positioning affects pollen release efficiency with anthers typically oriented laterally or downward facilitating gravitational pollen fall, positioning relative to stigma supporting self-pollination through close proximity, and overall spatial arrangement optimizing pollen transfer within flower. Dehiscence timing relative to stigma receptivity shows coordination with stigma receptivity typically developing within 6-24 hours after flower opening and cap fall, anther dehiscence often occurring concurrently or within similar timeframe, and temporal overlap creating pollination window with coordinated male-female reproductive readiness. Dehiscence progression throughout cluster follows developmental gradient with apical flowers showing earliest dehiscence reflecting earliest maturation, progressive extension to middle then basal regions creating sequential opening pattern, and overall 5-10 day period encompassing cluster-level dehiscence completion. Synchronization during peak flowering shows coordinated dehiscence in numerous flowers creating maximum pollen availability, favorable environmental conditions enabling widespread simultaneous opening, and concentrated pollen release supporting optimal pollination opportunity. Environmental effects on dehiscence efficiency include stable favorable conditions promoting normal opening across all flowers, variable conditions causing irregular dehiscence potentially reducing pollination efficiency, and unfavorable conditions particularly sustained high humidity or rain preventing dehiscence causing severe reproductive failure. Rain effects on dehiscence show catastrophic interference with water preventing anther opening through maintained hydration eliminating mechanical tension, washing pollen from any dehisced anthers, and promoting fungal infection on wet reproductive tissues, with rain during peak flowering potentially reducing fruit set by 50-80%. Incomplete dehiscence problems occur when stomium opening fails or proceeds partially, creating limited pollen liberation reducing pollination potential, associated with unfavorable environmental conditions or developmental abnormalities, and contributing to poor fruit set through male reproductive failure. Pollen viability at dehiscence shows maximum capacity immediately upon release with fresh pollen showing 70-90% germination under optimal conditions, rapid decline beginning immediately with viability reducing 5-10% per hour initially, and critical importance of efficient rapid pollination during brief high-viability window. The process shows three distinguishable aspects progressing from anther wall rupture initiating opening, through stomium opening creating access pathway, to initial pollen exposure establishing liberation. Duration spans 5-10 days at cluster level as anthers progressively dehisce, with individual anther opening occurring within minutes to hours once triggered. Cultivar variation in dehiscence shows differences in anther maturation timing affecting dehiscence onset, dehiscence efficiency with some varieties showing robust opening while others show incomplete opening problems, and coordination with stigma receptivity affecting pollination opportunity. Regional timing shows anther dehiscence during late May-early June in warm climates, early-mid June in temperate regions, and mid-late June in cool areas, coinciding with flowering period and enabling pollination. Viticultural significance includes representing essential male reproductive function determining pollen availability for pollination, requiring favorable environmental conditions particularly appropriate humidity for efficient opening, demonstrating extreme weather sensitivity with rain preventing dehiscence causing severe fruit set failure, and establishing anther function as critical yield determinant. Research applications include analyzing dehiscence mechanisms including structural, physiological, and environmental triggers, evaluating environmental effects on dehiscence efficiency affecting pollen liberation, investigating genetic variation in anther function affecting reproductive reliability, quantifying relationships between dehiscence characteristics and fruit set outcomes, and assessing management or environmental interventions affecting dehiscence success.
Anther_Dehiscence is first major Pollen_Release phase establishing liberation mechanism through anther opening. Subdivided into Anther_Wall_Rupture, Stomium_Opening, and Initial_Pollen_Exposure. Highly dependent on environmental conditions particularly humidity and temperature. Occurs following cap fall and flower opening typically within hours to day. Essential for pollen availability. Failure causes severe reproductive failure and poor fruit set. Transitions to Active_Pollen_Shedding as pollen liberates and disperses.
Antheren-Dehiszenz
Antheren-Ruptur
Pollensackaufbrechen
Staubbeutelöffnung
anther opening
anther rupture
locule dehiscence
pollen sac opening
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 375-380 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 287-290
Anther Dehiscence
Syrah in Rhône Valley shows anther wall rupture during mid-June Year N+1 at BBCH 60-65, with anther walls failing along stomium zones when morning humidity drops below 68% and temperature reaches 24-26°C creating mechanical stress exceeding tissue strength, rupture creating 2.5-3.5 mm longitudinal slits exposing pollen mass, under Mediterranean conditions with favorable morning dehydration enabling efficient rupture throughout cluster typical of this mid-to-late season cultivar
Anther wall rupture represents the initial structural failure in anther dehiscence when mechanical stresses exceed tissue strength creating opening in specialized stomium zones, initiating pollen liberation process through breach of protective anther wall. This critical mechanical event occurs during anther dehiscence in early-mid June in temperate regions Year N+1, typically occurring within minutes to hours once conditions trigger rupture as part of BBCH 60-65 flowering stages. Mechanical stress accumulation preceding rupture involves tissue dehydration causing volumetric shrinkage, differential contraction in hygroscopic fibrous layer creating tension, and stomium zone experiencing concentrated stress due to structural weakness representing predetermined failure site. Tissue dehydration driving rupture shows progressive water loss from anther wall tissues as environmental humidity decreases and temperature rises, hygroscopic fibrous layer cells containing cellulose microfibrils oriented to contract upon dehydration creating directional mechanical force, and inner wall layers shrinking differentially creating shearing stresses at stomium interface. Stomium zone vulnerability results from specialized structure with cells lacking secondary wall thickening found in adjacent tissues, mechanical weakness making tissue unable to resist mounting stress, and positioning along anther length providing optimal location for longitudinal slit formation enabling pollen access. Critical stress threshold shows rupture occurring when accumulated mechanical stress exceeds stomium tissue tensile strength estimated 0.3-0.8 MPa, threshold varying with tissue maturation state and hydration level, and rapid humidity decline enabling faster stress accumulation triggering earlier rupture. Rupture initiation typically begins at localized point along stomium zone where stress concentration or tissue weakness creates failure nucleus, crack propagation proceeding longitudinally along weak stomium tissue driven by relieved stress, and bilateral opening as both sides of ruptured stomium separate exposing locule interior. Rupture morphology shows characteristic longitudinal slit typically 2-4 mm length extending along anther corresponding to stomium zone length, clean separation along predetermined pathway indicating controlled failure mechanism, and slit width 0.3-0.8 mm sufficient for pollen grain passage 25-35 micrometers diameter. Acoustic evidence of rupture includes occasional audible sounds during intensive synchronized dehiscence though typically inaudible at individual anther level, possible detection of micro-acoustic emissions using sensitive equipment, and overall evidence of sudden mechanical failure creating brief energy release. Visual evidence of rupture shows sudden appearance of opening along previously intact anther, exposure of yellow pollen mass within locule becoming visible through slit, and immediate beginning of pollen liberation through opened pathway. Pollen pressure contribution to rupture remains uncertain with possible modest role for turgor-based pressure within locule assisting rupture propagation, though primary driving force clearly mechanical stress from differential tissue shrinkage, and pollen liberation following rather than driving initial rupture. Environmental conditions enabling rupture include declining relative humidity below 70-75% creating sufficient dehydration for stress accumulation, rising temperature accelerating dehydration and affecting tissue mechanical properties, and typically morning hours providing optimal conditions as humidity drops and temperature rises. Temperature effects on rupture show warm conditions 24-30°C facilitating rapid tissue dehydration enabling faster stress accumulation and rupture, cool conditions 15-20°C slowing dehydration potentially delaying rupture for hours or preventing if humidity remains elevated, and optimal temperature-humidity combination creating favorable rupture conditions. Humidity threshold for rupture shows critical decline below 70-75% typically required for initiating rupture in mature anthers, sustained high humidity above 75% preventing sufficient dehydration maintaining tissue strength above failure threshold, and optimal 50-65% humidity enabling efficient rupture throughout cluster. Tissue maturation requirements show properly matured anther walls with completed development of fibrous layer hygroscopic properties necessary for effective dehydration-driven stress, premature rupture attempts in immature anthers potentially failing or producing incomplete opening, and over-mature senescent anthers potentially showing irregular rupture or retention. Rupture timing relative to cap fall shows characteristic sequence with cap shedding typically preceding rupture by hours to day as flower opens first, anther maturation and conditioning requiring brief period before dehiscence readiness, and favorable environmental conditions triggering rupture once thresholds reached. Rupture propagation rate shows rapid crack extension once initiated typically completing within seconds to minutes, coordinated opening of both locules per anther creating paired slits, and sequential rupture throughout cluster as individual anthers reach critical conditions. Failed rupture causes incomplete dehiscence with partial opening providing limited pollen access reducing liberation efficiency, or complete failure maintaining closed anther preventing any pollen release, resulting from unfavorable conditions particularly sustained high humidity or inadequate tissue maturation. The process represents first aspect of anther dehiscence establishing breach in protective wall, followed by stomium opening discussing specialized tissue and initial pollen exposure revealing grains. Duration represents brief mechanical event occurring within seconds to minutes once triggered. Cultivar variation in rupture characteristics may include tissue mechanical properties affecting rupture threshold, anther wall structure affecting failure pattern, and sensitivity to environmental conditions affecting rupture efficiency. Regional timing shows anther wall rupture during flowering period late May-early June in warm climates, early-mid June in temperate regions, and mid-late June in cool areas, occurring as anthers reach maturity and conditions become favorable. Viticultural significance includes representing critical mechanical event enabling pollen liberation, demonstrating extreme environmental dependence requiring favorable humidity and temperature, and establishing rupture failure as potential cause of poor fruit set through prevented pollen release. Research applications include analyzing mechanical and structural basis of rupture, evaluating environmental effects on rupture triggering and efficiency, investigating tissue maturation requirements for successful rupture, quantifying rupture dynamics and propagation, and assessing genetic variation in anther structure affecting rupture reliability.
Anther_Wall_Rupture is first aspect of Anther_Dehiscence, distinguished by initial structural failure creating opening. Represents mechanical event driven by dehydration stress. Occurs at specialized stomium zone predetermined weak tissue. Requires favorable environmental conditions particularly humidity decline. Brief seconds to minutes duration once triggered. Followed by Stomium_Opening discussing tissue specialization and Initial_Pollen_Exposure. Essential first step enabling pollen liberation. Failure prevents pollen release causing reproductive failure.
Antheren-Spaltung
Antherenwand-Ruptur
Staubbeutelwandaufbruch
Stomium-Bruch
anther splitting
anther wall breaking
locule opening
stomium rupture
Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 287-290 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 376-380
Anther Wall Rupture
Cabernet Sauvignon in Napa Valley shows anthesis during early-mid June Year N+1 at BBCH 60-65 over 7-9 days, with first flowers opening June 4-6 followed by peak anthesis June 7-11 showing 50-70% simultaneous opening and completing June 12-14, under warm climate conditions of 22-28°C temperature, 45-65% relative humidity, light winds, no rain during bloom, supporting efficient pollination and 50-65% fruit set typical of this late-season cultivar
Anthesis represents the flower opening phase encompassing cap shedding, reproductive organ exposure, pollen release, and pollination processes where flowers transition from closed bud state to open functional reproductive structures enabling pollen transfer to receptive stigmas, establishing conditions for fertilization and fruit set. This fundamental reproductive stage occurs during early-mid June in temperate regions Year N+1, typically spanning 5-10 days from first flower opening through completion across cluster depending on developmental synchrony and environmental conditions, corresponding to BBCH 60-65 stages. Anthesis progression shows characteristic temporal pattern beginning with sporadic opening of earliest-maturing flowers typically at cluster apex corresponding to earliest primordium formation, accelerating through peak anthesis when maximum 40-70% flowers open simultaneously within 2-3 days, and declining during late phase as final 20-30% flowers complete opening, creating overall flowering duration of 5-10 days under favorable conditions. Cap shedding represents distinctive grapevine flowering mechanism with fused petal calyptra detaching as unit from receptacle base, lifting away from flower through combined forces of expanding reproductive organs pushing upward, turgor changes potentially creating mechanical pressure, and wind or gravity causing final detachment and fall. Calyptra detachment shows characteristic appearance with cap initially loosening at base showing slight gap, progressively lifting upward as abscission zone cells separate, and ultimately detaching completely falling away from flower typically within minutes to hours of abscission initiation. Abscission zone development involves enzymatic degradation of middle lamella in specialized cell layers at calyptra-receptacle junction, with cellulase and pectinase enzymes breaking down cell wall polysaccharides creating separation plane, under hormonal regulation possibly involving ethylene promoting abscission and auxin potentially regulating timing. Reproductive organ exposure following cap shedding reveals functional flower structures with five stamens surrounding central pistil, anthers positioned above stigma in close proximity facilitating self-pollination, nectary disc at flower base secreting nectar in some flowers, and overall flower diameter 3-5 mm depending on cultivar. Anther dehiscence proceeds rapidly after cap shedding with specialized stomium zones in anther walls breaking down creating longitudinal slits along anther length, dehiscence triggered by humidity decrease causing anther wall dehydration creating mechanical tension opening slits and releasing pollen. Pollen release occurs through anther slits with mature pollen grains liberating from anther locules, initially adhering in small masses then dispersing as individual grains, with release facilitated by flower movement from wind creating agitation dislodging pollen. Pollen dispersal proceeds primarily through wind transport with light pollen grains 25-35 micrometers diameter easily becoming airborne, traveling distances from centimeters within same flower to meters between vines depending on wind conditions, with pollen cloud visible surrounding flowering vines under favorable dry windy conditions. Pollen capture on stigmas occurs when airborne pollen grains contact receptive stigma surfaces, adhering to sticky exudate produced by stigma papillate cells, with pollen capture efficiency affected by stigma position, exudate production, and environmental conditions. Stigma receptivity develops progressively with papillate surface cells elongating and secreting proteinaceous exudate creating adhesive surface, receptivity typically peaking 1-3 days after flower opening and persisting 4-7 days though declining progressively, with receptive period affected by temperature and humidity. Flower longevity after opening shows stamens remaining functional 1-3 days with viable pollen production declining rapidly after initial release, stigma remaining receptive 4-7 days providing extended opportunity for pollination, and overall flower structure persisting 5-10 days before senescence if fertilization fails or longer if successful fertilization initiates fruit development. Pollination efficiency in grapevine shows high success with self-pollination within individual flowers through close proximity of anthers and stigma enabling direct pollen transfer, supplemented by wind-assisted pollination between neighboring flowers on same cluster or adjacent clusters, with cross-pollination between vines playing minor role. Nectary function shows variable activity among flowers with some producing modest nectar quantities potentially attracting insect visitors including honeybees, though insect contribution to pollination is minimal compared to dominant wind-assisted self-pollination mechanism. Flower scent production varies among cultivars with some producing subtle floral fragrance potentially attracting insects, while many commercial cultivars show minimal scent, with scent consisting of volatile organic compounds including terpenoids and aromatics. Environmental conditions critically affect anthesis with temperature as primary factor with optimal 20-30°C supporting rapid synchronized opening and effective pollination, cool conditions below 15°C substantially delaying opening and reducing pollen viability, and excessive heat above 35°C potentially causing pollen sterility or incomplete cap shedding. Humidity affects anthesis with moderate 40-70% relative humidity optimal for cap shedding and anther dehiscence, high humidity above 80% delaying dehiscence and potentially preventing pollen release, and low humidity below 30% causing rapid pollen desiccation reducing viability though facilitating release and dispersal. Rain during anthesis represents critical threat with water preventing cap shedding by maintaining high humidity and physical interference, washing pollen from anthers and stigmas preventing pollination, promoting botrytis and other fungal infections on wet flowers, and potentially physically damaging delicate reproductive structures, with rain timing particularly detrimental during peak anthesis. Wind affects anthesis showing beneficial effects through facilitating pollen dispersal and enhancing pollination efficiency, though excessive wind may cause flower damage or overly rapid pollen desiccation, with light-moderate wind 5-15 km per hour generally optimal. Light intensity shows modest direct effects on anthesis though adequate light supports overall vine vigor and photosynthetic capacity providing resources for reproductive processes, with shaded flowers showing similar opening patterns though potentially reduced vigor. Metabolic activity during anthesis shows elevated respiration particularly in anthers supporting pollen maturation and dehiscence, active sugar metabolism in nectaries producing nectar in functional flowers, and maintained metabolism in stigmas supporting receptivity and exudate production. Resource requirements during anthesis emphasize carbohydrate demand for energy-intensive processes including anther dehiscence, pollen tube growth after pollination, and initiating fruit development, with adequate reserves supporting successful reproduction while limitation potentially reducing fruit set. Hormonal regulation involves declining gibberellins as reproductive transition completes, ethylene potentially triggering or coordinating cap abscission, auxin from successfully fertilized ovaries preventing flower abscission and initiating berry development, and other hormones coordinating reproductive processes. Developmental uniformity within cluster affects anthesis duration with highly synchronized development producing compressed 5-7 day flowering period facilitating uniform fruit set, while heterogeneous development extends period to 10-14 days potentially creating variable berry development and extended vulnerability to adverse conditions. The process shows two distinguishable phases progressing from early anthesis with initial flower opening establishing reproductive function, through peak anthesis when maximum flowers open simultaneously creating optimal pollination window. Duration typically spans 5-10 days depending on developmental synchrony and environmental conditions, with warm stable conditions promoting rapid coordinated anthesis while variable cool conditions extend period. Cultivar variation appears in anthesis timing with early varieties flowering late May-early June while late types flower mid-late June, anthesis duration with some showing compressed 5-7 day period while others show extended 8-12 day period, cap shedding efficiency with some varieties showing clean complete shedding while others show retained caps causing coulure, and pollen production and viability affecting fruit set success. Regional timing shows anthesis during late May-early June in warm climates, early-mid June in temperate regions, and mid-late June in cool areas, with timing critical for frost avoidance and favorable weather window. Viticultural management considerations include monitoring anthesis progress for timing management operations and interventions, protecting flowering from adverse weather particularly rain through site selection and training systems, avoiding canopy operations during anthesis preventing physical disturbance of delicate flowers, ensuring adequate vine nutrition particularly boron supporting pollen function, recognizing anthesis as critical period determining fruit set and yield, and understanding weather during anthesis fundamentally affects season outcomes. Research applications include analyzing mechanisms regulating cap abscission and anther dehiscence, investigating environmental controls of anthesis timing and duration, evaluating pollen biology including production, dispersal, and viability, quantifying pollination efficiency under varying conditions, and assessing genetic variation in anthesis characteristics affecting reproductive success and adaptation.
Anthesis is first major phase of Flowering, distinguished by flower opening and pollination processes. Subdivided into Early_Anthesis and Peak_Anthesis based on temporal progression and flowering intensity. Followed by fertilization and fruit set processes. Corresponds to BBCH 60-65. Environmental conditions particularly temperature and humidity critically affect success. Duration and timing major determinants of fruit set and yield.
Anthese
Blühphase
Blütenöffnung
Kalyptra-Abwurf
bloom period
cap shedding period
flower opening
flowering phase
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 360-375 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 275-285 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Anthesis
Pinot Noir in Oregon shows anthesis completion during early July Year N+1 at BBCH 69 on approximately July 2, with all reproductive processes terminated, definitive fruit set of 58-68% establishing 310-420 berries per cluster, visible berry development 5-8 mm diameter throughout cluster, unfertilized flowers showing active abscission, under cool climate conditions of 20-25°C temperature, typical of this early-to-mid season cultivar completing flowering 13-15 days after first opening and transitioning definitively to berry development phase
Anthesis completion represents the definitive reproductive phase conclusion when all flowering processes have terminated, fruit set outcomes are established, and cluster has fully transitioned to developmental phase with no residual reproductive activity or potential. This conclusive reproductive milestone occurs during complete anthesis in late June-early July in temperate regions Year N+1 shortly after all flowers opened, representing 1-2 day period confirming reproductive cessation and developmental transition, corresponding to BBCH 69 complete. Completion confirmation shows multiple converging indicators including absence of any further flower opening activity with all viable buds having opened, complete cessation of pollen production and availability with all anthers depleted, elimination of stigma receptivity with all stigmas either fertilized or senesced, and overall reproductive function terminated across cluster. Fruit set finalization at completion establishes definitive berry number with count of successfully fertilized flowers determining yield potential, typical set percentages 30-75% flowers creating 100-500 berries per cluster depending on cultivar and conditions, and established berry population representing maximum potential fruit load though subsequent berry drop or management thinning may reduce final harvest numbers. Berry development focus at completion shows all metabolic and developmental resources directing toward expanding berries with rapid cell division creating berry cell number potential, hormonal signals from developing seeds and pericarp coordinating growth, and berry expansion visible with 0.3-0.6 mm diameter increase daily during this rapid early growth phase. Unfertilized flower fate at completion shows senescence universally established with all failed reproductions showing brown discoloration and tissue deterioration, abscission zones fully developed enabling progressive flower drop, and elimination of non-functional reproductive structures cleaning cluster leaving only productive berries. Abscission progression at completion shows active flower drop with daily detachment of senescing unfertilized flowers, accumulation of fallen flowers creating visible debris, and progressive cluster thinning as failed reproductions eliminate over subsequent 1-2 weeks. Cluster appearance at completion shows definitive developmental state with small green berries dominating visual impression, minimal or moderate brown senescing flowers interspersed depending on fruit set percentage, overall healthy developmental appearance with active growth evident, and clear transition from reproductive to developmental cluster character. Developmental phase establishment at completion confirms berry growth as dominant process with cell division phase proceeding rapidly establishing cell number, early berry expansion visible creating recognizable fruit structures, and developmental trajectory established toward véraison and ripening. Resource allocation at completion shows complete transition to developmental priorities with photosynthate directing primarily to expanding berries as major sink, vegetative growth continuing consuming moderate resources, and reproductive investment completely ceased with no residual flowering demands. Metabolic patterns at completion show elevated respiration in rapidly dividing berry cells supporting high energy demands, declining metabolism in senescing flowers representing resource recovery, and overall cluster-level metabolism focused entirely on developmental processes. Hormonal environment at completion shows auxin dominance from developing berries preventing abscission and promoting expansion, gibberellins affecting cell division with peak levels during early berry development, cytokinins supporting cell division in berries and maintaining vegetative growth, and ethylene in senescing flowers promoting abscission but not affecting established berry development. Seed development at completion shows embryo and endosperm growth progressing following successful fertilization, seed structures rapidly enlarging within developing berries, and hormonal production from seeds contributing substantially to berry growth signals. Environmental effects at completion show minimal reproductive vulnerability with berry development relatively robust compared to earlier flowering sensitivity, though extreme weather potentially affecting vegetative growth supporting berry development or causing physical damage to young berries. Temperature at completion optimally 20-28°C supporting rapid berry cell division and growth, with accumulation of growing degree days driving developmental progression toward subsequent expansion and maturation phases. Water availability at completion becomes increasingly important for berry development with adequate moisture supporting cell division and early expansion, and initiation of irrigation management if practiced supporting berry development through season. Nutrient management at completion emphasizes continued nitrogen for vegetative growth and berry development, increasing potassium importance for berry quality outcomes, adequate calcium for berry structure, and balanced nutrition supporting healthy development. Disease management at completion focuses on berry protection with fungicide applications for botrytis and powdery mildew preventing infection during susceptible early development, recognition that latent infections from flowering will remain dormant, and continued vigilance throughout development period. Canopy management at completion implements shoot positioning optimizing berry exposure, possible hedging controlling excessive growth, strategic leaf removal in some systems improving cluster microclimate, and overall canopy optimization balancing vegetative growth with fruit development support. Phenological significance at completion includes definitively marking BBCH 69 anthesis completion, establishing transition point to BBCH 71-79 fruit development stages, providing reference for subsequent phenological predictions including véraison and harvest timing, and confirming major developmental phase transition in annual cycle. Management transition at completion enables full resumption of normal vineyard operations unrestricted by flowering sensitivities, implementation of fruit-focused management strategies, shift in priorities from reproductive to developmental and quality optimization, and planning for season progression toward harvest. Yield prediction at completion becomes possible through berry number assessment multiplied by expected final berry weight, comparison to historical norms evaluating season potential, and informing crop management decisions including possible thinning or cluster management. The process represents second of three complete anthesis aspects confirming reproductive conclusion, following all flowers opened and preceding post-anthesis transition. Duration represents brief 1-2 day definitive conclusion period, with clear transition toward developmental phase. Cultivar variation in completion characteristics shows timing differences with early varieties completing late June while late types complete early-mid July, completion clarity with some showing definitive conclusion while others show gradual extended completion, and fruit set outcomes ranging widely affecting berry number and density. Regional timing shows anthesis completion during late June in warm climates at approximately 1150-1200 growing degree days base 10°C, late June-early July in temperate regions at 1050-1100 GDD, and early-mid July in cool areas at 950-1000 GDD, marking reproductive phase conclusion. Viticultural significance includes establishing definitive reproductive conclusion enabling developmental focus, confirming fruit set and berry number determining yield potential, marking end of critical flowering period reducing management constraints, providing reference point for season planning and phenological tracking, and enabling yield prediction and crop management planning. Research applications include analyzing mechanisms coordinating reproductive termination, evaluating factors determining fruit set outcomes, documenting transition from reproductive to developmental physiology, quantifying resource allocation shifts affecting vine function, and assessing phenological transitions in annual developmental progression.
Anthesis_Completion is second of three Complete_Anthesis aspects, distinguished by definitive reproductive cessation confirmation. Follows All_Flowers_Opened establishing opening extent. Represents conclusive reproductive milestone with fruit set finalized. Marks full transition to developmental phase. Enables management transitions and yield predictions. Critical phenological reference point in annual progression.
Antheseabschluss
Blühschluss
Blütephaseende
anthesis termination
bloom finish
flowering conclusion
reproductive completion
reproduktive Vollendung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 390-395 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Anthesis Completion
In Merlot berries in Saint-Émilion, Bordeaux during BBCH 81-89, anthocyanin accumulation increases total anthocyanin content from 0.15 to 1.65 mg/g fresh weight over 42 days, with malvidin-3-glucoside accumulating to 0.82 mg/g representing 50% of total anthocyanins, petunidin-3-glucoside reaching 0.28 mg/g, and delphinidin-3-glucoside at 0.22 mg/g, establishing the characteristic Merlot color profile under moderate oceanic conditions averaging 24°C with beneficial diurnal temperature variation.
Anthocyanin accumulation describes the progressive biosynthesis and vacuolar deposition of anthocyanin pigments in Vitis vinifera berry skin cells during the post-veraison ripening period, wherein the coordinated expression of flavonoid pathway genes enables synthesis of the cyanidin, delphinidin, peonidin, petunidin, and malvidin glycosides that produce the red, purple, and blue coloration characterizing ripe berries of pigmented grape varieties over the 30-45 day period from veraison to harvest maturity. This process involves the final steps of the flavonoid biosynthetic pathway with anthocyanidin synthase converting leucoanthocyanidins to anthocyanidins, UDP-glucose flavonoid glucosyltransferase attaching glucose to form stable anthocyanin glucosides, and O-methyltransferases generating the methylated derivatives peonidin, petunidin, and malvidin from their respective unmethylated precursors cyanidin and delphinidin. The quantitative aspects of anthocyanin accumulation show dramatic cultivar variation, with deeply colored varieties such as Alicante Bouschet accumulating 2000-4000 mg/kg berry weight, moderately pigmented varieties such as Cabernet Sauvignon reaching 1000-2000 mg/kg, and lightly colored varieties such as Pinot Noir achieving 400-800 mg/kg under equivalent ripening conditions, reflecting genetic differences in pathway enzyme expression levels and regulatory factor activities. Cellular localization restricts anthocyanin accumulation to hypodermal cell layers of the exocarp where flavonoid pathway enzymes show coordinate expression, with synthesized anthocyanins transported into vacuolar compartments through glutathione S-transferase mediated mechanisms and tonoplast transporters, establishing the vacuolar pigment pools that confer berry color visible through the overlying epidermal cell layer. The temporal dynamics of anthocyanin accumulation show characteristic patterns with rapid initial accumulation during the first 2-3 weeks post-veraison when biosynthetic enzyme activities peak, continued but declining accumulation during mid-ripening, and potential plateau or slight decline during late ripening as degradation may partially offset continued synthesis in some varieties and conditions. Temperature profoundly influences anthocyanin accumulation, with optimal synthesis occurring at moderate temperatures of 20-25°C daytime and 15-18°C nighttime, while temperatures exceeding 35°C substantially inhibit accumulation through enzyme inactivation and accelerated pigment degradation, explaining the characteristic color deficiency observed in heat-affected vintages. Light exposure promotes anthocyanin accumulation through activation of MYB transcription factors that regulate flavonoid pathway gene expression, with appropriately exposed clusters showing 30-80% higher anthocyanin content than heavily shaded fruit, though excessive direct radiation may cause thermal damage that counteracts the positive light effects.
Anthocyanin_Accumulation is parent class under Phenolic_Maturation representing pigment biosynthesis and deposition in skin. Contains subclasses Progressive_Anthocyanin_Synthesis, Color_Intensity_Development, and Anthocyanin_Profile_Evolution. Duration 30-45 days post-veraison. Exclusive to hypodermal exocarp cells. Temperature and light sensitive. Cultivar-specific accumulation capacity ranges from 400 to 4000 mg/kg.
Anthocyan-Akkumulation
Anthocyan-Biosynthese-und-Einlagerung
Beeren-Pigment-Akkumulation
Farbpigment-Anreicherung
anthocyanin biosynthesis and deposition
anthocyanin development
berry pigment accumulation
color pigment accumulation
Boss PK et al (1996) Expression of anthocyanin biosynthesis pathway genes in red and white grapes. Plant Molecular Biology 32: 565-569 Castellarin SD et al (2007) Colour variation in red grapevines. Planta 227: 895-908 Keller M (2020) The Science of Grapevines 3rd Ed pp 310-325
Anthocyanin Accumulation
Anthocyanin concentration in red grape skins ranges from 200-400 mg/100g in lightly colored cultivars such as Pinot Noir to 1500-2500 mg/100g in deeply colored types such as Tannat and Alicante Bouschet. Five anthocyanidin aglycones occur in V. vinifera, differing in B-ring hydroxylation and methoxylation pattern, with 3-O-glucoside and 3-O-acylglucoside conjugates predominating. Anthocyanin biosynthesis begins at veraison with coordinated upregulation of VvUFGT (UDP-glucose:flavonoid 3-O-glucosyltransferase), VvMYBA1/A2 transcription factors, and structural genes. Accumulation peaks 2-4 weeks post-veraison, with concentration subsequently declining through degradation and dilution, losing 20-40% between peak and harvest in warm climates.
A phenolic compound disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate anthocyanin pigments in grape berry skins through the flavonoid pathway, with genotype-specific total concentration, anthocyanin profile (relative proportions of individual anthocyanins), and modification patterns (glycosylation, acylation) determining red wine color intensity, hue, and stability.
Anthocyanin_Accumulation_Disposition serves as the parent class for specific anthocyanin type and concentration traits. Anthocyanins are the sole pigments responsible for red grape and wine color, with their chemistry fundamentally affecting color expression. The trait shows high heritability (h² = 0.70-0.85) for total concentration but also strong environmental modulation, particularly by temperature and light. The VvMYBA1/A2 locus is the major genetic determinant, with white-fruited cultivars carrying non-functional alleles.
Anthocyan-Disposition
Anthocyanakkumulation
Farbstoffbildung
anthocyanin synthesis capacity
grape color trait
pigment accumulation disposition
Boss PK et al. (1996) Expression of anthocyanin biosynthesis pathway genes in red and white grapes. Plant Molecular Biology 32:565-569 Castellarin SD, Di Gaspero G (2007) Transcriptional control of anthocyanin biosynthetic genes in extreme phenotypes for berry pigmentation of naturally occurring grapevines. BMC Plant Biology 7:46
Anthocyanin Accumulation Disposition
Anthocyanin accumulation quality assessment for phosphorus deficiency in newly planted Willamette Valley Chardonnay (a white cultivar where leaf reddening is unambiguously stress-related) provided clear diagnostic indication of phosphorus limitation in the establishing vineyard. Visual assessment identified anthocyanin accumulation quality symptoms as reddish-purple coloration beginning at leaf margins and progressing intervenally on basal and mid-shoot leaves, with color intensity scored from 0 (no reddening) through 2 (marginal reddening) to 5 (entire leaf red-purple). Affected vines showed mean anthocyanin accumulation quality scores of 3.5 on basal leaves and 2.0 on mid-shoot leaves, with youngest leaves showing minimal coloration reflecting their priority access to limited phosphorus supplies. The anthocyanin accumulation quality pattern distinguished phosphorus deficiency from other potential causes, as cold-induced anthocyanins would appear on youngest rather than oldest leaves and follow temperature events viral reddening would show vein-associated patterns and magnesium deficiency would show interveinal chlorosis rather than reddening. Spectrophotometric quantification of leaf anthocyanin content confirmed visual scoring, with affected leaves containing 85 μg/g fresh weight anthocyanins versus less than 5 μg/g in normal green Chardonnay leaves. The clear anthocyanin accumulation quality diagnosis enabled targeted phosphorus fertilization that resolved symptoms within 8 weeks, with follow-up assessment showing restoration of normal green coloration confirming that phosphorus limitation had been the cause of the abnormal pigmentation.
Anthocyanin accumulation quality in phosphorus deficiency is a phosphorus deficiency symptom quality that inheres in grapevine leaves and represents the characteristic red-to-purple pigmentation developing as a stress response to phosphorus limitation, with anthocyanin synthesis activated by carbohydrate accumulation in leaves when phosphorus deficiency limits the ATP-dependent processes of sugar export and utilization, creating the distinctive coloration that is often the most visually striking and diagnostically useful symptom of phosphorus deficiency in grapevine. This quality captures the pigmentation response to phosphorus stress, recognizing that the red-purple coloration provides a visible indicator of phosphorus limitation but must be distinguished from other causes of anthocyanin accumulation including cold temperatures, senescence, viral infection, and genetic factors in red-fruited cultivars. The ontological significance of anthocyanin accumulation quality in phosphorus deficiency lies in its visual diagnostic utility enabling field identification of phosphorus limitation, while also requiring careful differential diagnosis since similar coloration patterns can result from unrelated causes. The biological basis of phosphorus-deficiency anthocyanin accumulation encompasses the role of phosphorus in phloem loading and sugar export from leaves, where phosphorus deficiency reduces export capacity causing carbohydrate accumulation that triggers anthocyanin biosynthetic gene expression through sugar signaling pathways, combined with the direct signaling effects of phosphorus status on flavonoid metabolism regulatory networks. The pattern of anthocyanin accumulation in phosphorus deficiency typically shows marginal initiation with progression toward interveinal areas, often with more intense coloration on older leaves reflecting their longer exposure to deficiency conditions and greater carbohydrate accumulation, though the specific pattern varies with deficiency severity, cultivar anthocyanin genetics, and environmental conditions including light exposure that can intensify pigmentation. The distinction from viral-induced reddening (which follows vein patterns) and from cold-induced anthocyanins (which appear rapidly with temperature drops and affect youngest tissues) provides the diagnostic criteria for attributing red coloration to phosphorus deficiency rather than other causes.
Anthocyanin_Accumulation_Quality_(P) is_a Phosphorus_Deficiency_Symptom_Quality, representing the red-purple leaf pigmentation resulting from carbohydrate accumulation when P deficiency limits sugar export. This class addresses the most visually striking P deficiency symptom. Sugar accumulation triggers anthocyanin biosynthesis through signaling pathways. Marginal initiation with progression intervenally distinguishes from viral (vein-associated) patterns. White cultivars provide clearest diagnosis since leaf reddening is unambiguously stress-related. Distinguished from cold-induced anthocyanins affecting youngest leaves and from senescence-associated pigmentation.
Anthocyan-Akkumulation bei P-Mangel
P-Limitierungs-Anthocyane
P-Mangel-Rötung
P-deficiency reddening
P-limitation anthocyanins
Phosphorstress-Pigmentierung
phosphorus stress pigmentation
purple coloration from P deficiency
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Chalker-Scott L (1999) Environmental significance of anthocyanins in plant stress responses. Photochemistry and Photobiology 70:1-9 Hammond JP, White PJ (2008) Sucrose transport in the phloem: integrating root responses to phosphorus starvation. Journal of Experimental Botany 59:93-109
Anthocyanin Accumulation Quality (P)
Anthocyanin content for Cabernet Sauvignon from Rutherford, Napa Valley (2023 vintage) measures 1.95 mg/g berry fresh weight by the Iland method, with HPLC profiling revealing malvidin-3-glucoside as the dominant species (52% of total), followed by petunidin-3-glucoside (15%), delphinidin-3-glucoside (12%), peonidin-3-glucoside (11%), and cyanidin-3-glucoside (4%), with acylated forms comprising 38% of total anthocyanins — a profile characteristic of Cabernet Sauvignon and indicative of excellent color stability potential. Anthocyanin content for Blaufränkisch from Mittelburgenland (2024 vintage) records 3.1 mg/g at 20° KMW, among the highest values documented for the variety, with the exceptional pigmentation attributed to the Bl-2 clone's genetic predisposition for intense color development combined with the favorable 2024 vintage conditions of moderate temperatures during the anthocyanin synthesis window (15-25°C daytime, 10-15°C nocturnal).
Anthocyanin content is a phenolic data entity that records the concentration and, where determined, the compositional profile of anthocyanin pigments in the skins of red and rosé grape varieties, measured by pH-differential spectrophotometry (total anthocyanins), HPLC (individual anthocyanin species), or the Iland berry homogenate method, expressed in milligrams per gram of berry fresh weight (mg/g), milligrams per liter of must (mg/L), or milligrams per berry, and representing the primary determinant of red wine color depth, hue, and stability that constitutes a key criterion for phenolic maturity assessment and harvest timing in red wine production.
Anthocyanin content is the primary chemical determinant of red wine visual quality and one of the most intensively studied components of grape phenolic maturity. The five anthocyanidin aglycones present in Vitis vinifera (cyanidin, peonidin, delphinidin, petunidin, malvidin) occur as 3-O-monoglucosides and as acylated derivatives (acetylated, p-coumaroylated, caffeoylated), creating a species-specific anthocyanin profile that serves as a cultivar fingerprint — Cabernet Sauvignon is dominated by malvidin with high acylation, Pinot Noir contains predominantly non-acylated monoglucosides, and Tempranillo shows a balanced five-anthocyanidin distribution. The biosynthetic pathway is controlled by the VvMYBA1 and VvMYBA2 transcription factors, with mutations in these genes responsible for the white-skinned sport variants (e.g., Pinot Blanc from Pinot Noir). Environmental regulation of anthocyanin accumulation creates significant vintage variation — optimal synthesis occurs at moderate temperatures (17-25°C) with adequate UV-B exposure, while high temperatures (>35°C) simultaneously inhibit biosynthesis and accelerate degradation, leading to anthocyanin loss in hot vintages. The distinction between total anthocyanins (extracted at pH 1.0 or in 50% ethanol) and extractable anthocyanins (extracted at pH 3.2 simulating wine conditions) provides the extractability index — values above 70% indicate advanced cell wall degradation and readiness for harvest, while lower values suggest that additional hang time may improve color extraction during winemaking. Berry size inversely correlates with anthocyanin concentration per unit weight (smaller berries have higher skin-to-pulp ratio), contributing to the established relationship between water deficit, reduced berry size, and enhanced color in red wines.
Anthocyananalyse
Anthocyangehalt
Anthocyankonzentration
Beerenpigmentgehalt
Farbstoffgehalt
anthocyanin analysis
anthocyanin concentration
anthocyanin level
grape color content
pigment content
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2, 2nd Ed, Wiley, pp.: 173-203 Boss PK et al. (1996) Expression of anthocyanin biosynthesis pathway genes in red and white grapes. Plant Molecular Biology 32(3): 565-569 Castellarin SD et al. (2007) Colour variation in red grapevines. Journal of Experimental Botany 58(15-16): 4257-4270 Iland P et al. (2011) Chemical Analysis of Grapes and Wine, 2nd Ed, pp.: 100-110
Anthocyanin Content
Anthocyanin content quality profiling in Australian Shiraz employs HPLC analysis quantifying individual anthocyanins to assess both total concentration (target 800-1500 mg/kg at harvest) and compositional profile (malvidin-3-glucoside typically 40-55% of total, with acylated forms contributing 25-40%) the proportion of acylated anthocyanins, particularly malvidin-3-(6-acetyl)-glucoside and malvidin-3-(6-p-coumaroyl)-glucoside, correlates with color stability during aging, guiding selection of fruit for premium wines intended for extended cellaring versus early-drinking styles.
Anthocyanin_Content_Quality encompasses the chemical qualities characterizing the concentration and composition of anthocyanin pigments in grape berry skins, capturing the flavonoid subclass directly responsible for the red, purple, and blue coloration of colored grape cultivars and red wines. Anthocyanins are glycosylated forms of anthocyanidins, the colored aglycones, with glucose attached at the 3-position of the C ring in Vitis vinifera and additional glycosylation at the 5-position in some non-vinifera species. The five primary anthocyanidins in V. vinifera grapes are malvidin, delphinidin, petunidin, peonidin, and cyanidin, distinguished by the hydroxylation and methoxylation pattern of the B ring. Malvidin-based anthocyanins typically dominate in most red wine cultivars, comprising 50-90% of total anthocyanins, while the proportions of other anthocyanidins provide cultivar-specific fingerprints useful for authenticity verification. Total anthocyanin concentration in grape skins varies enormously among cultivars, from less than 100 mg/kg fresh weight in lightly colored varieties (Pinot Gris, Gewürztraminer) through 500-1500 mg/kg in typical wine grapes (Cabernet Sauvignon, Merlot) to greater than 5000 mg/kg in teinturier varieties (Alicante Bouschet) with colored flesh. Anthocyanin accumulation begins at véraison under control of MYB transcription factors and continues through ripening, with accumulation rates and final concentrations influenced by temperature (optimal around 25-30°C, inhibited above 35°C), light exposure, and water status. Anthocyanins exist in pH-dependent equilibrium among structural forms including the red flavylium cation, colorless carbinol and chalcone forms, and blue quinoidal bases, with the distribution among forms affecting color expression.
Anthocyanin_Content_Quality serves as the parent class for individual anthocyanidin-based subclasses (Malvidin, Cyanidin, Peonidin, Delphinidin, Petunidin). The glycosidic forms (mono- and di-glucosides) and acylated derivatives (acetyl, coumaroyl, caffeoyl) multiply the number of individual compounds, with 15-20 major anthocyanins routinely quantified in comprehensive profiles. Anthocyanin content in grapes provides color potential, but extraction efficiency during winemaking and subsequent reactions (polymerization, co-pigmentation) determine final wine color. The pH-dependent color expression of anthocyanins means that identical anthocyanin concentrations produce different color intensities at different pH values.
Anthocyanpigmentgehalt
Anthocyanzusammensetzung
Rotfarbstoffkonzentration
anthocyanin composition
anthocyanin pigment content
grape color compound quality
red pigment concentration
He F et al. (2012) Anthocyanins and Their Variation in Red Wines I. Monomeric Anthocyanins and Their Color Expression. Molecules 17(2): 1571-1601 Castellarin SD et al. (2007) Colour Variation in Red Grapevines (Vitis vinifera L.): Genomic Organisation, Expression of Flavonoid 3'-Hydroxylase, Flavonoid 3',5'-Hydroxylase Genes and Related Metabolite Profiling of Red Cyanidin-/Blue Delphinidin-Based Anthocyanins in Berry Skin. BMC Genomics 8(1): 428
Anthocyanin Content Quality
Anthocyanin degradation quality evaluation in Barossa Valley Shiraz documented the color consequences of heat exposure during the critical post-véraison color development period when temperatures exceeded 35°C for 12 consecutive days. Berry sampling at weekly intervals tracked anthocyanin dynamics through the heat event and subsequent recovery period. Anthocyanin concentration in heat-exposed fruit reached a maximum of only 0.95 mg/g fresh weight versus 1.65 mg/g in shaded control fruit, representing 42% anthocyanin degradation quality deficit attributed to the combined effects of biosynthesis inhibition and pigment degradation. HPLC profiling revealed selective loss of the non-acylated anthocyanins (malvidin-3-glucoside and related compounds) with 55% reduction versus only 25% reduction in acylated forms, confirming differential thermal stability affecting the anthocyanin profile as well as total concentration. Transcript analysis showed UFGT (UDP-glucose flavonoid glucosyltransferase) expression reduced to 30% of control levels during peak heat, confirming biosynthetic inhibition as a major contributor to the anthocyanin degradation quality response. Post-heat-event recovery showed limited capacity for anthocyanin restoration, with final harvest concentrations remaining 35% below control despite 3 weeks of moderate temperatures following the heat event, establishing that anthocyanin degradation quality impacts during critical developmental windows cannot be fully compensated by subsequent favorable conditions.
Anthocyanin degradation quality is a heat impact on development quality that inheres in red grape berries and represents the extent of heat-induced reduction in anthocyanin pigment concentration through the combined effects of biosynthesis inhibition at elevated temperatures and the degradation or chemical modification of existing anthocyanin molecules, resulting in reduced color intensity and altered color stability in affected fruit with direct consequences for red wine color quality. This quality captures the particular vulnerability of anthocyanin metabolism to heat stress, recognizing that red grape color development is one of the most temperature-sensitive quality parameters, with optimal accumulation occurring at moderate temperatures (15-25°C) and substantial inhibition or degradation occurring above 30-35°C depending on cultivar and exposure duration. The ontological significance of anthocyanin degradation quality lies in the fundamental importance of color for red wine quality perception and commercial value, with heat-induced color deficits potentially requiring extended maceration, co-pigmentation enhancement, or blending strategies to achieve acceptable color in wines from affected fruit, and with the most severely affected fruit potentially unsuitable for premium red wine production regardless of other quality attributes. The biological basis of anthocyanin degradation encompasses both the thermal inhibition of biosynthetic enzymes including phenylalanine ammonia-lyase, chalcone synthase, and UDP-glucose flavonoid glucosyltransferase that reduces new anthocyanin synthesis, and the non-enzymatic degradation of existing anthocyanins through temperature-accelerated oxidation, hydrolysis of glycosidic bonds, and reactions with other berry constituents that remove pigments from the extractable pool. The temperature threshold for anthocyanin degradation varies among cultivars reflecting genetic differences in the thermal sensitivity of biosynthetic enzymes and the chemical stability of the specific anthocyanin profiles characteristic of different varieties, with the acylated anthocyanins predominant in some cultivars showing greater thermal stability than the non-acylated forms in others. The timing of heat exposure critically affects outcomes, with véraison-period heat inhibiting the establishment of color synthesis while post-véraison heat both inhibits ongoing synthesis and degrades previously accumulated pigments, potentially causing severe color deficits in fruit exposed to sustained heat during the primary color development window.
Anthocyanin_Degradation_Quality is_a Heat_Impact_On_Development_Quality, representing the heat-induced reduction in red grape pigment concentration through biosynthesis inhibition and pigment degradation. This class addresses the most temperature-sensitive quality parameter for red grape production. The percentage deficit from control provides quantification. Both biosynthesis inhibition (enzyme) and pigment degradation (chemical) contribute to losses. Acylated anthocyanins show greater thermal stability than non-acylated forms. Véraison-period heat is particularly damaging as it affects both synthesis establishment and ongoing accumulation. Recovery capacity is limited following developmental window closure.
Anthocyan-Abbau
Farbverlust
Pigment-Hitzeschaden
anthocyanin loss
color degradation
heat-induced color loss
hitzeinduzierter Farbverlust
pigment heat damage
Mori K et al. (2007) Loss of anthocyanins in red-wine grape under high temperature. Journal of Experimental Botany 58:1935-1945 Tarara JM et al. (2008) Berry temperature and solar radiation alter acylation, proportion, and concentration of anthocyanin in Merlot grapes. American Journal of Enology and Viticulture 59:235-247 Sadras VO, Moran MA (2012) Elevated temperature decouples anthocyanins and sugars in berries of Shiraz and Cabernet Franc. Australian Journal of Grape and Wine Research 18:115-122
Anthocyanin Degradation Quality
The anthocyanin distribution description of Pinot Meunier (syn. Schwarzriesling) documents a distinctive pattern — intense anthocyanin expression on the adaxial (upper) surface of young leaves confined to a 2-3 mm marginal band with the central lamina remaining green, combined with strong anthocyanin accumulation on the dense erect trichomes of the abaxial (lower) leaf surface producing a characteristic pinkish-grey underside, and uniformly pigmented shoot tip with no green ground color visible — this complex distribution pattern, resulting from the periclinal chimera constitution of Pinot Meunier (L1 layer with different genetic background from L2/L3 layers), creates a shoot tip appearance immediately distinguishable from the parent cultivar Pinot Noir (uniform anthocyanin across entire young leaf surface) and the related sport Pinot Gris (reduced anthocyanin intensity but similar distribution pattern to Pinot Noir).
A shoot tip description component that documents the spatial pattern of anthocyanin pigmentation on the developing shoot tip, young leaves, and young shoot internodes, recording the specific distribution of red-purple coloration across organ surfaces — adaxial versus abaxial leaf surfaces, margins versus lamina, dorsal versus ventral internode faces, and nodes versus internodes — that provides finer cultivar discrimination than anthocyanin intensity alone by capturing the genetically determined spatial regulation of anthocyanin biosynthesis in vegetative tissues.
Anthocyanin_Distribution_Description captures the spatial component of anthocyanin pigmentation that provides diagnostic information beyond the simple intensity scoring of OIV 003. Two cultivars may share the same OIV 003 intensity score (e.g., both scored as 5, medium) but differ dramatically in distribution pattern — one may show uniform pigmentation across all young leaf surfaces while another concentrates anthocyanin on margins, veins, or specific organ faces, reflecting differences in the tissue-specific regulation of anthocyanin biosynthesis gene expression: these distribution patterns are controlled by regulatory networks involving R2R3-MYB, bHLH, and WD40 transcription factors interacting with tissue-specific promoter elements, producing genetically determined pigmentation maps that are consistent within a cultivar and diagnostic between cultivars. The distribution pattern is particularly informative for distinguishing periclinal chimeras — genotypes in which different meristem layers (L1, L2, L3) have different anthocyanin production capacities, creating layer-specific pigmentation visible as distinct color zones on young organs where cell layer contributions to different tissue regions are developmentally fixed.
Anthocyan-Verteilungs-Beschreibung
Pigmentierungsmuster-Dokumentation
anthocyanin pattern record
pigmentation distribution description
vegetative Färbungsverteilung
vegetative coloration mapping
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340
Anthocyanin Distribution Description
Anthocyanin maturity monitoring in Syrah across a diurnal temperature gradient reveals the temperature effect on color development, with warm site (minimal diurnal range, frequent >35°C maxima) showing reduced anthocyanin accumulation peaking at 1.2 mg/g berry with low acylation (15% acylated forms) and earlier plateau, while cool site (15°C diurnal range, maxima rarely >30°C) achieves 1.9 mg/g with higher acylation (28% acylated forms) and extended accumulation period, with the 58% difference in total anthocyanins and near-doubled acylation proportion translating to substantially greater color intensity and stability potential, demonstrating how site-specific anthocyanin maturity assessment informs both harvest timing and wine quality expectations.
Anthocyanin maturity quality is a phenolic maturity quality that represents the developmental state of anthocyanin pigments in grape skin, characterizing the accumulation, molecular composition, and stability of these color-determining compounds that establish the visual character of red wines and contribute to color longevity through co-pigmentation and polymeric pigment formation during aging. Within the BFO framework, this quality specifically addresses the anthocyanin component of phenolic maturity, distinguishing color pigment development from tannin or other phenolic compound maturation while recognizing that anthocyanin maturity involves not only quantity but also composition (the profile of individual anthocyanins) and stability factors (acylation, methylation, co-pigmentation) that together determine the color quality and persistence achievable in resulting wines. The ontological significance lies in anthocyanin maturity determining both the intensity and stability of red wine color, where total anthocyanin concentration establishes color potential, the proportion of acylated anthocyanins affects color stability (acylated forms more resistant to degradation and browning), and the degree of co-pigmentation with flavonols and hydroxycinnamic acids influences both color intensity and hue in young wines. The accumulation dynamics of anthocyanins during ripening show onset at véraison, rapid accumulation during the first weeks of ripening, a plateau or maximum at intermediate ripeness, and potential decline during late ripening through degradation, dilution from continued berry growth, or metabolic transformation, with the timing and magnitude of these phases varying among cultivars and affected by temperature and light exposure. The composition changes during anthocyanin maturation include shifts in the profile of individual anthocyanins (five main forms in Vitis vinifera: delphinidin, cyanidin, petunidin, peonidin, malvidin glucosides with malvidin typically dominant), progressive acylation (acetyl, p-coumaryl, caffeoyl additions stabilizing the chromophore), and methylation patterns, with these qualitative changes affecting color characteristics beyond simple concentration effects. The temperature effects on anthocyanin maturity are significant, as elevated temperatures (>30°C) can inhibit anthocyanin accumulation and accelerate degradation, potentially limiting color development in warm climates or during heat events, while moderate diurnal temperature variation (warm days, cool nights) often favors anthocyanin accumulation, creating the climate-dependent patterns of color development that affect regional wine character. Understanding anthocyanin maturity quality enables assessment of color potential and stability, harvest timing for optimal color development, and recognition of the environmental factors affecting anthocyanin accumulation that determine vintage variation in wine color.
Anthocyanin_Maturity_Quality addresses the accumulation, composition, and stability of color-determining pigments. This quality involves quantity, molecular profile, and modification state affecting color intensity and persistence. The class supports color quality optimization through anthocyanin maturity characterization.
Anthocyanreife
Farbentwicklung
Farbstoffreife
Pigmentreife
anthocyanin ripeness
color development
color maturity
pigment maturity
Mori K et al. (2007) Loss of anthocyanins in red-wine grape under high temperature. Journal of Experimental Botany 58(8): 1935-1945 Castellarin SD et al. (2007) Colour onset and colour evolution in berries of Merlot and Cabernet Sauvignon is related to expression of the anthocyanin biosynthetic genes. Australian Journal of Grape and Wine Research 13(3): 156-167
Anthocyanin Maturity Quality
In Tempranillo berries in Ribera del Duero, Spain during BBCH 83-89, anthocyanin profile evolution shifts malvidin-3-glucoside proportion from 42% at early post-veraison to 58% at harvest over 40 days, with concurrent increase in acylated forms from 18% to 35% of total anthocyanins, while delphinidin-3-glucoside decreases proportionally from 22% to 14% and cyanidin-3-glucoside from 8% to 4%, establishing the characteristic Tempranillo profile with high malvidin and substantial acylation under warm continental conditions.
Anthocyanin profile evolution describes the progressive changes in the relative proportions of individual anthocyanin species within Vitis vinifera berry skins during the post-veraison ripening period, wherein the initial anthocyanin composition dominated by tri-hydroxylated forms including delphinidin and petunidin derivatives transitions toward increased representation of methylated and acylated species generating the characteristic mature anthocyanin profile that influences wine color stability, hue expression, and aging behavior. This process involves the continued activity of O-methyltransferases converting cyanidin to peonidin and delphinidin to petunidin and malvidin, with methylation typically increasing throughout ripening such that malvidin-3-glucoside proportion rises from 35-45% at early post-veraison to 45-60% at harvest in many varieties, establishing malvidin dominance characteristic of Vitis vinifera anthocyanin profiles. The biosynthetic basis of profile evolution reflects differential enzyme activities and substrate preferences, with flavonoid 3-prime-hydroxylase favoring tri-hydroxylated B-ring formation generating delphinidin precursors, flavonoid 3-prime-5-prime-hydroxylase competing for substrates, and subsequent O-methyltransferase activities determining the extent of methylation among accumulated anthocyanidins that establishes final profile proportions. Acylation represents an additional dimension of profile evolution, with anthocyanin acyltransferases attaching acetyl, p-coumaroyl, and caffeoyl groups to glucose residues of anthocyanin glucosides, typically increasing during late ripening such that acylated forms may comprise 20-50% of total anthocyanins at harvest in varieties expressing high acyltransferase activities. The cultivar-specific nature of anthocyanin profile evolution reflects genetic variation in pathway enzyme expression, with varieties such as Cabernet Sauvignon showing high malvidin proportions of 60-70% at maturity compared to varieties such as Pinot Noir maintaining higher cyanidin representation of 15-25%, and varieties such as Nebbiolo demonstrating distinctive profiles with elevated peonidin contribution reflecting reduced methyltransferase activities toward certain substrates. Environmental factors modulate anthocyanin profile evolution through differential effects on individual pathway enzymes, with temperature and light conditions influencing the relative activities of hydroxylases, methyltransferases, and acyltransferases that determine profile proportions independent of total anthocyanin accumulation. The enological significance of anthocyanin profile evolution relates to wine color properties, with malvidin-based anthocyanins showing greater color stability through enhanced participation in copigmentation and polymerization reactions compared to cyanidin derivatives, while acylated anthocyanins demonstrate improved stability under winemaking conditions due to intramolecular copigmentation protecting the chromophore from hydration.
Anthocyanin_Profile_Evolution is subclass of Anthocyanin_Accumulation representing compositional changes among anthocyanin species. Distinguishes from total accumulation by emphasizing relative proportions. Sibling to Progressive_Anthocyanin_Synthesis and Color_Intensity_Development. Malvidin typically increases from 35-45% to 45-60%. Acylated forms increase during late ripening. Cultivar-specific profiles reflect genetic enzyme expression differences.
Anthocyan-Profil-Evolution
Anthocyan-Verhältnis-Evolution
Anthocyan-Zusammensetzungs-Änderung
Pigment-Profil-Entwicklung
anthocyanin composition change
anthocyanin ratio evolution
pigment profile development
pigment species distribution change
Castellarin SD et al (2007) Colour variation in red grapevines. Planta 227: 895-908 Fournier-Level A et al (2009) Quantitative genetic bases of anthocyanin variation in grape. BMC Plant Biology 9: 147 Keller M (2020) The Science of Grapevines 3rd Ed pp 318-325
Anthocyanin Profile Evolution
Evaluation of Anthracnose_Resistance_Disposition at the Grape Genetic Resources Repository of the Brazilian Agricultural Research Corporation (Embrapa Uva e Vinho) screened 185 Vitis accessions under high natural disease pressure in the humid subtropical climate of Rio Grande do Sul. Susceptible V. vinifera cultivar 'Italia' exhibited severe anthracnose with 75-90% of young shoots showing girdling cankers, 60-80% defoliation from leaf lesions, and 45-65% of berries displaying bird's eye lesions by véraison. The V. labrusca cultivar 'Isabella' demonstrated strong Anthracnose_Resistance_Disposition with only 5-10% of shoots showing minor cankers, minimal leaf damage (<10%), and berry infection below 5%. Interspecific hybrids showed variable resistance correlated with their proportion of North American species ancestry, with 'BRS Vitória' (complex hybrid) exhibiting intermediate resistance (20-30% shoot infection, 15% berry infection). Anatomical studies revealed that resistant genotypes possess thicker cuticles on young shoots and more rapid wound periderm formation following penetration, limiting lesion expansion and secondary sporulation. No major resistance genes have been identified, with inheritance studies suggesting polygenic control with moderate heritability (h² = 0.45-0.55).
An Anthracnose_Resistance_Disposition is a Fungal_Resistance_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through defense processes upon encounter with the hemibiotrophic ascomycete Elsinoë ampelina (anamorph Sphaceloma ampelinum), the causal agent of grape anthracnose, also known as bird's eye rot. This disposition represents the capacity to limit E. ampelina infection, colonization, and sporulation on susceptible tissues including young leaves, shoots, tendrils, petioles, and developing berries. Anthracnose produces characteristic symptoms including circular leaf spots with grey centers and dark margins ("bird's eye" lesions), sunken cankers on green shoots that may girdle and kill young growth, and circular berry lesions with raised dark borders that crack and serve as entry points for secondary pathogens. The disease is favored by warm, wet spring conditions and is particularly severe in tropical and subtropical viticulture regions and in humid continental climates during rainy growing seasons. Resistance mechanisms include physical barriers limiting conidial penetration, rapid activation of defense responses during the biotrophic infection phase, and limitation of lesion expansion during the necrotrophic phase. Unlike downy and powdery mildew, anthracnose resistance genetics remain poorly characterized, with resistance appearing largely quantitative and derived primarily from North American Vitis species including V. rupestris, V. labrusca, and V. cinerea. European V. vinifera cultivars display a range of susceptibility, with some traditional cultivars from humid regions showing moderate field tolerance likely reflecting historical selection pressure.
Anthracnose_Resistance_Disposition is_a Fungal_Resistance_Disposition. Anthracnose caused by Elsinoë ampelina is a distinct disease from the "anthracnose" of other crops, with grape-specific symptomatology and epidemiology. Unlike downy mildew, powdery mildew, and black rot, no major resistance genes (R genes) have been characterized for anthracnose in Vitis. Resistance appears quantitative and is found primarily in North American species adapted to humid climates where E. ampelina is endemic. The disease is particularly important in tropical and subtropical viticulture where environmental conditions favor repeated infection cycles.
Anthraknose-Resistenz-Disposition
Brenner-Resistenz
Elsinoë resistance disposition
Elsinoë-Resistenz
Sphaceloma resistance
bird's eye rot resistance
Poolsawat O et al. (2012) Identification and quantification of resistance to anthracnose (Elsinoë ampelina) in wild and cultivated grapes, Vitis 51:95-100 Santos RF et al. (2018) Grape anthracnose resistance: screening and inheritance studies, Plant Pathology 67:1535-1543 Thind TS (2015) Anthracnose of gapes, Diseases of Fruits and Vegetables and Their Management, Indian Phytopathological Society
Anthracnose Resistance Disposition
Antioxidant response quality evaluation in Shiraz during post-véraison heat stress assessed the oxidative defense mechanisms protecting berry quality during the critical ripening period when heat events most frequently occur. Heat exposure at 42°C for 6 hours triggered comprehensive antioxidant response quality activation with superoxide dismutase (SOD) activity increasing 2.8-fold within 24 hours, catalase (CAT) activity increasing 2.2-fold, and ascorbate peroxidase (APX) activity increasing 3.5-fold, while non-enzymatic antioxidant pools showed ascorbate increasing 1.8-fold and reduced glutathione increasing 2.1-fold. The antioxidant response quality effectively limited oxidative damage, with lipid peroxidation (measured as malondialdehyde) increasing only 35% above control levels versus 120% in vines where antioxidant response was pharmacologically inhibited, demonstrating the protective value of the response. Berry-level assessment revealed that the antioxidant response quality in skin tissue was critical for anthocyanin preservation, with strong responders maintaining 85% of anthocyanin content while weak responders showed 40% degradation attributed to oxidative bleaching. The antioxidant response quality assessment identified APX as the most responsive enzyme in this cultivar and developmental stage, suggesting that ascorbate-based scavenging represents the primary oxidative defense during berry ripening under heat stress.
Antioxidant response quality is a heat shock response quality that inheres in a grapevine organism and represents the capacity to upregulate antioxidant defense systems in response to elevated temperatures, enhancing the enzymatic and non-enzymatic mechanisms that neutralize reactive oxygen species (ROS) whose accumulation under heat stress would otherwise cause oxidative damage to membranes, proteins, and nucleic acids that exacerbates and extends the direct thermal injury. This quality captures the oxidative stress management dimension of heat tolerance, recognizing that heat-induced ROS accumulation represents a secondary stress mechanism that may cause damage persisting beyond the direct thermal effects, making antioxidant response an essential complement to the protein-protective functions of heat shock proteins for comprehensive heat tolerance. The ontological significance of antioxidant response quality lies in its role preventing the cascade of oxidative damage that follows primary heat injury, with effective antioxidant response limiting the spatial and temporal extent of damage while inadequate response permits ROS-mediated injury to spread beyond directly heat-affected tissues and persist beyond the period of heat exposure itself. The biological basis of antioxidant response encompasses the transcriptional and post-translational activation of antioxidant enzymes including superoxide dismutase converting superoxide radicals to hydrogen peroxide, catalase and ascorbate peroxidase decomposing hydrogen peroxide to water, and glutathione reductase maintaining the reduced glutathione pool essential for glutathione peroxidase activity, combined with the accumulation of non-enzymatic antioxidants including ascorbate, glutathione, tocopherols, and carotenoids that directly scavenge various ROS species. The coordination between enzymatic and non-enzymatic antioxidant systems provides comprehensive protection across cellular compartments, with chloroplastic, mitochondrial, cytosolic, and peroxisomal antioxidant pools each contributing to overall oxidative stress management. The temporal dynamics of antioxidant response typically show enzyme activity increases within hours of heat exposure, reaching maximum within 24-48 hours and remaining elevated during recovery, providing sustained protection against oxidative damage that may continue as damaged tissues undergo repair.
Antioxidant_Response_Quality is_a Heat_Shock_Response_Quality, representing the oxidative stress management component of heat protection. This class addresses both enzymatic (SOD, CAT, APX, GR) and non-enzymatic (ascorbate, glutathione, tocopherols) antioxidant systems. Fold-increases in enzyme activity and antioxidant pool sizes provide quantification. The response complements HSP-mediated protein protection by managing secondary oxidative damage. Berry-level antioxidant response is critical for pigment preservation. Distinguished from Heat_Shock_Protein_Expression_Quality by the focus on ROS management rather than protein chaperoning.
Antioxidantien-Antwort
Antioxidantiensystem-Hochregulation
ROS defense activation
ROS-Abwehr-Aktivierung
antioxidant defense response
antioxidant system upregulation
antioxidative Stressreaktion
oxidative stress response
Wahid A et al. (2007) Heat tolerance in plants: an overview. Environmental and Experimental Botany 61:199-223 Wang LJ et al. (2006) Salicylic acid alleviates decreases in photosynthesis under heat stress and accelerates recovery in grapevine leaves. BMC Plant Biology 10:34 Carvalho LC et al. (2015) Heat stress in grapevine: the pros and cons of acclimation. Plant, Cell and Environment 38:777-789
Antioxidant Response Quality
The AOC designation Hermitage AOC (established 1937 among the original AOC designations) delimits 136 hectares of steep granite hillside above Tain-l'Hermitage in the northern Rhône, with the cahier des charges specifying Syrah as the exclusive red variety (Marsanne and Roussanne permitted as white, up to 15% in red blends by tradition), maximum yield of 40 hl/ha, minimum natural alcohol of 10.5%, and mandatory analytical and sensory examination by the INAO commission — the AOC designation Hermitage may appear on labels alongside or interchangeably with the AOP (Appellation d'Origine Protégée) designation as both terms are legally equivalent under EU Regulation 1308/2013 Art. 112.
A regional wine designation within the French wine classification system that identifies a wine as an Appellation d'Origine Contrôlée — the traditional French designation certifying that a wine originates from a specific delimited area and is produced according to the established practices and quality standards codified in the appellation's cahier des charges, administered by INAO (Institut National de l'Origine et de la Qualité), formally equivalent to Protected Designation of Origin (AOP) under EU harmonization but retaining the AOC terminology as an accepted traditional term on French wine labels.
AOC_Designation documents the original and most influential appellation system in wine history, established by the decree-law of July 30, 1935 creating the Comité National des Appellations d'Origine (predecessor to INAO). The French AOC system provided the model for virtually all subsequent wine geographic indication systems worldwide — Italy's DOC (1963), Spain's DO (1932, reformed 1970), Portugal's DOC (1908, reformed 1986), and the EU harmonized PDO framework (2008) all derive structural and conceptual elements from the French AOC tradition: France currently has approximately 360 wine AOCs organized in complex regional hierarchies ranging from broad regional appellations (Bordeaux AOC) through communal (Pauillac AOC) to single-vineyard (Le Montrachet AOC) designations. The 2009 EU harmonization formally replaced AOC with AOP as the legal designation category, but EU Regulation 1308/2013 Art. 112 permits continued use of AOC as an accepted traditional term, and the vast majority of French wine labels continue to display AOC rather than AOP.
AOC
AOC-Bezeichnung
Appellation d'Origine Contrôlée
Appellation-d'Origine-Contrôlée-Klassifikation
French controlled origin designation
französische kontrollierte Herkunftsbezeichnung
Decree-law of July 30, 1935 establishing the Comité National des Appellations d'Origine INAO (2023) Les appellations d'origine contrôlées EU Regulation 1308/2013 Art. 93 and Art. 112
AOC Designation
The AOP designation Sancerre AOP appears on wine labels either as the sole designation or alongside the traditional AOC mention, with both expressions carrying identical legal force — a Sancerre producer may label wines as Appellation Sancerre Contrôlée (traditional AOC format), Appellation d'Origine Protégée Sancerre (harmonized EU format), or display both the AOC and AOP designations, with the EU AOP logo (a red-and-yellow circular emblem) mandatory on labels since 2012 for wines sold within the EU single market regardless of which textual designation format is used. In Luxembourg, AOP is the primary designation term for wines from the Moselle luxembourgeoise, as the country adopted the EU terminology directly without an intermediate national designation system equivalent to France's AOC.
A regional wine designation constituting the EU-harmonized French-language expression of Protected Designation of Origin (Appellation d'Origine Protégée) as applied within the framework of EU Regulation 1308/2013, formally replacing the national AOC designation as the legal category while functioning interchangeably with AOC on French wine labels, and serving as the standard PDO designation term in francophone Belgium, Luxembourg, and Switzerland under their respective wine regulations.
AOP_Designation captures the EU harmonization layer that was superimposed on national appellation systems through the 2008 wine reform (Regulation 479/2008, subsequently consolidated in Regulation 1308/2013). The relationship between AOC and AOP in France is one of formal legal equivalence — AOP is the EU regulatory category, AOC is the accepted traditional term permitted under Art. 112, and both designate exactly the same wines subject to exactly the same production rules: the distinction is administrative rather than substantive, with the European Commission maintaining the official register of AOP designations while INAO continues to administer the operational aspects of appellation management using AOC terminology. The coexistence of AOC and AOP on the French market has generated some consumer confusion, as the AOP designation is less familiar to French consumers despite being mandatory on EU-market labels since 2012. In other francophone wine-producing jurisdictions (Luxembourg, francophone Switzerland, Quebec), AOP functions as the primary designation without an underlying traditional term equivalent to AOC.
AOP
AOP-Bezeichnung
Appellation d'Origine Protégée
Appellation-d'Origine-Protégée-Klassifikation
EU-harmonized French origin designation
geschützte Ursprungsbezeichnung (französisch)
EU Regulation 1308/2013 Art. 93 and Art. 112 EU Regulation 479/2008 on the common organisation of the market in wine European Commission Delegated Regulation 2019/33 Art. 12-14 on labeling requirements
AOP Designation
Leaf apex shape in Grüner Veltliner is characteristically acute to acuminate, forming a distinctly pointed terminal lobe that extends 8-15 mm beyond the lateral sinuses, contrasting with the more obtuse or rounded apex typical of Riesling leaves this sharp apex, combined with the medium-deep lateral sinuses and slightly overlapping petiolar sinus, creates the distinctive five-pointed star appearance that facilitates field identification of Grüner Veltliner in Austrian vineyards where it often grows adjacent to other white varieties.
Apex_Shape_Quality represents the morphological quality characterizing the terminal configuration of grapevine organs at their distal extremity. The apex constitutes the growing tip or terminal region of an organ, and its shape provides diagnostic characters for cultivar identification while reflecting developmental processes and genetic regulation of organ form. In leaves, apex shape describes the configuration of the blade tip at the terminal end of the midrib, with categories ranging from acute (sharply pointed) through obtuse (bluntly angled) to rounded and emarginate (notched). Leaf apex shape is captured in OIV descriptor 076 with standardized categories enabling reproducible ampelographic assessment. In berries, apex shape characterizes the stylar end opposite the pedicel attachment, varying from truncate (flattened) through rounded to pointed depending on cultivar and developmental conditions. Berry apex shape affects visual appearance in table grapes and can indicate the position of residual stylar tissue. In shoots, apex shape describes the configuration of the growing tip and surrounding developing leaves, relevant for assessing growth vigor and phenological stage. Apex shape assessment employs visual matching to standardized reference images, measurement of apex angle, or characterization of curvature at the terminal region. The apex integrates the cumulative effect of growth patterns along the organ's length and represents the most recently formed tissue in growing organs.
Apex_Shape_Quality addresses the terminal end of organs and is positioned as a direct subclass of Shape_Quality, parallel to Overall_Form_Quality, Symmetry_Quality, Base_Shape_Quality, and Margin_Shape_Quality. The apex is defined as the distal extremity opposite the point of attachment or origin for leaves, this is the blade tip for berries, the stylar end for shoots, the growing tip. OIV descriptor 076 (shape of blade tip) provides the standardized reference for leaf apex assessment. Apex shape can be modified by environmental conditions and damage, so assessment should focus on well-developed, undamaged organs.
Apexform
Spitzenform
apical form
distal shape
distale Gestalt
terminal configuration
terminale Konfiguration
tip shape
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 56-57 Galet P (1979) A Practical Ampelography, Cornell University Press pp.: 82-89
Apex Shape Quality
In water-stressed Grenache vines in southeastern Australia during extreme drought conditions, occasional apex terminal flower clusters develop at primary shoot tips following shoot apical meristem conversion at BBCH 53-55, exhibiting reduced cluster size (6-9 cm length, 120-200 flowers) compared to normal lateral clusters (12-18 cm, 500-800 flowers) and definitively terminating shoot growth at 18-25 nodes compared to typical 25-35 node shoots with continued vegetative apex. Certain North American Vitis species (V. riparia selections) exhibit apex terminal cluster habit as a genetic trait, producing small terminal clusters (5-8 cm, 80-150 flowers) at shoot tips in addition to normal lateral clusters at lower nodes.
An apex terminal flower cluster is a terminal flower cluster positioned specifically at the absolute apex of the primary shoot or a major lateral shoot, representing flowers borne on an inflorescence that develops from or replaces the shoot apical meristem, thereby terminating vegetative shoot extension and converting the shoot tip from indeterminate vegetative growth to determinate reproductive development. Apex terminal flower clusters represent conversion of the shoot apical meristem itself to reproductive fate, contrasting with normal grapevine architecture where shoot apical meristems maintain vegetative identity (producing leaves, tendrils, and continued shoot extension) while lateral axillary meristems produce inflorescences. This apical conversion is relatively rare in standard Vitis vinifera wine cultivars under typical cultural conditions but can occur under specific circumstances= (1) genetic factors in certain wild Vitis species (particularly some American species) that exhibit terminal cluster habit (2) environmental stress conditions (severe water deficit, extreme temperatures, nutritional stress) during shoot development that trigger premature reproductive conversion of the apical meristem (3) hormonal disruption from exogenous growth regulator application or endogenous hormonal imbalances affecting the vegetative-reproductive transition (4) developmental abnormalities in individual shoots where normal auxin-mediated maintenance of apical meristem vegetative identity is disrupted. Apex terminal flower clusters occupy the absolute shoot tip position, with the inflorescence peduncle emerging as the continuation of the shoot axis rather than as a lateral appendage, and shoot growth conclusively terminating at the base of the inflorescence. The developmental dynamics of apex terminal clusters differ fundamentally from lateral clusters= conversion represents transformation of an existing vegetative meristem rather than initiation of new lateral primordia, developmental timing may be dissociated from normal lateral cluster timing (which is determined during previous season's bud differentiation), and resource allocation patterns differ due to loss of the competing shoot apex sink. Apex terminal flower clusters typically exhibit modified morphology compared to lateral clusters from the same genotype= cluster size may be reduced (often 30-50% smaller than lateral clusters) due to abbreviated primordium development following late-season apical conversion, branching architecture may be simplified with reduced secondary and tertiary branching, or conversely clusters may be larger if apical conversion occurs early and captures resources normally allocated to continued shoot extension. The physiological and developmental significance of apex terminal clusters includes definitive termination of shoot vegetative growth, elimination of auxin production from the shoot apex (auxin is primarily synthesized in apical meristems) with consequent effects on lateral bud development and organ patterning, and altered carbohydrate partitioning as continued shoot extension ceases. Fruit set and berry development in apex terminal clusters may differ from lateral clusters, with some evidence suggesting altered hormone balances (reduced auxin from absent apex) affecting fruit set success, though definitive data are limited due to the rarity of this cluster type in commercial cultivars.
Apex_Terminal_Flower_Cluster is distinguished from End_Branch_Terminal_Cluster by specific position at the shoot apical meristem (absolute apex) versus at branch termini, and by representing conversion of the primary vegetative meristem versus development of inflorescences at lateral branch apices. True apex terminal clusters are rare in Vitis vinifera wine cultivars, making this class primarily of interest for understanding developmental plasticity, stress responses, and comparative morphology with wild Vitis species. The developmental mechanisms underlying apical meristem conversion from vegetative to reproductive fate in grapevine deserve research attention, as understanding these mechanisms could inform strategies for manipulating reproductive development or managing stress-induced developmental abnormalities. Occurrence of apex terminal clusters in commercial vineyards should be investigated as a potential indicator of severe stress conditions requiring cultural intervention.
Blütentraube an Haupttriebspitze
primary terminal inflorescence
primäre endständige Blütentraube
scheitelapikale Blütentraube
shoot-apex flower cluster
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 75-85 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 159-168.
Apex Terminal Flower Cluster
Apical dominance assessment comparing cultivars reveals the genetic variation affecting canopy complexity, with high-dominance cultivar (Cabernet Sauvignon) showing strong lateral suppression (only 12% of lateral buds producing visible growth on intact shoots, compact canopy architecture), while low-dominance cultivar (Grenache) shows weak suppression (65% of laterals producing growth even with intact apices, requiring summer hedging to control canopy density), with the dominance difference creating distinct management requirements—Cabernet needing minimal lateral control while Grenache demands repeated attention to prevent excessive shading from profuse lateral development.
Apical dominance quality is a paradormancy quality that represents the inhibitory influence of the shoot apex on the outgrowth of lateral buds positioned below it, characterizing the strength of this correlative control that determines whether lateral buds remain dormant, initiate weak growth, or develop into vigorous lateral shoots depending on the dominance exerted by the terminal growing point. Within the BFO framework, this quality specifically addresses the apical-lateral dimension of paradormancy, distinguishing the inhibition originating from shoot tips from other sources of correlative control while recognizing that apical dominance is the primary mechanism maintaining the monopodial or weakly branched growth habit during the early season, with its strength determining the degree of lateral shoot development (feminelles or summer laterals) that occurs during active growth. The ontological significance lies in apical dominance determining the branching architecture of individual shoots and thus the density and complexity of the developing canopy, where strong apical dominance produces minimally branched shoots with suppressed lateral growth (simplifying canopy management), while weak apical dominance permits extensive lateral development creating dense, complex canopies that may require summer pruning to maintain fruit-zone conditions. The hormonal mechanism of apical dominance involves auxin synthesized in the shoot apex and transported basipetally through the polar auxin transport stream, where this auxin flux inhibits lateral bud outgrowth through effects on the bud's own hormone balance (suppressing cytokinin biosynthesis, maintaining bud dormancy through auxin-cytokinin antagonism) and through strigolactone signaling that reinforces the inhibitory effect. The cultivar variation in apical dominance shows some varieties with characteristically strong dominance (minimal lateral development unless apex is removed) and others with weak dominance (profuse lateral growth even with intact apices), with this variation affecting canopy management requirements and the natural branching patterns that develop without intervention. Understanding apical dominance quality enables prediction of lateral bud behavior following topping, hedging, or shoot damage, assessment of cultivar-specific canopy development patterns, and management strategies that either maintain or release apical dominance to achieve desired canopy architecture.
Apical_Dominance_Quality addresses the inhibitory influence of shoot apices on lateral bud outgrowth. This quality varies among cultivars affecting canopy architecture and management requirements. The class supports branching prediction and canopy management through apical dominance characterization.
Apex-Lateral-Verhältnis
Apikaldominanz
Terminaldominanz
Triebspitzenhemmung
apex-lateral relationship
apical control
shoot tip inhibition
terminal dominance
Cline MG (1991) Apical dominance. The Botanical Review 57(4): 318-358 Domagalska MA, Leyser O (2011) Signal integration in the control of shoot branching. Nature Reviews Molecular Cell Biology 12(4): 211-221
Apical Dominance Quality
An apical leaf at node 14 on an actively extending shoot of Vitis vinifera cv. Cabernet Sauvignon in Bordeaux, France, positioned 18 cm behind the shoot apex with three younger leaves at nodes 15-17 still ahead, emerging 10 days previously during vigorous shoot extension phase, measuring 8.5 cm length by 7.8 cm width representing 48% of basal leaf area on the same shoot, exhibiting pale green coloration with chlorophyll content of 285 μg cm⁻² (compared to 485 μg cm⁻² in mature leaves), soft flexible texture with thin cell walls in palisade layers averaging 42 μm (compared to 85 μm in mature leaves), incomplete tissue differentiation with developing single palisade layer and loosely organized spongy mesophyll, demonstrating photosynthesis rate of 5.8 μmol CO₂ m⁻² s⁻¹ representing 44% of mature capacity and remaining net carbon sink consuming approximately 12 mg carbohydrate per day for continued growth while producing only 8 mg through limited photosynthesis, positioned in well-lit upper canopy receiving 75% full sunlight, occurring during active shoot extension at 6-8 cm per week growth rate, measured at shoot BBCH stage 75 (berries pea-sized) in late June, representing typical apical leaf characteristics during vigorous vegetative growth phase.
A leaf positioned in the upper distal portion of an actively extending shoot typically occupying the terminal 3-6 nodes in Vitis vinifera, characterized by recent emergence from shoot apical meristem typically within the previous 1-3 weeks, smaller dimensions compared to basal and median leaves often 30-60% reduced area reflecting shorter development duration and resource competition with shoot extension, incomplete maturation with many leaves remaining in expanding or partially expanded stages during shoot growth periods, pale to medium green coloration reflecting lower chlorophyll concentrations in young developing tissues, soft texture from thin cell walls and incomplete lignification, anatomical characteristics showing incomplete tissue differentiation with developing mesophyll layers and immature vascular systems, suboptimal photosynthetic capacity typically 40-70% of mature leaf rates reflecting incomplete chloroplast development and lower enzyme concentrations, net carbon sink status consuming more resources for growth than produced through limited photosynthesis, and functional role transitioning from pure developmental sink during emergence through partial source function as leaves approach maturity if shoot extension ceases. Apical leaves occupy the active growth zone of extending shoots where new organs are continuously initiated from apical meristems, creating dynamic populations with the youngest leaves at the extreme tip in primordial or emerging stages, progressively older leaves at increasing distances from apex showing advancing developmental stages, and the oldest apical zone leaves approaching maturity potentially 15-25 cm from the apex depending on internode lengths and shoot vigor, with the entire apical leaf population representing major resource investment supporting future photosynthetic capacity.
Apical leaves are distinguished from basal and median leaves by distal shoot position, recent emergence, incomplete development, and net sink status during shoot extension phases. The exact node range defining apical zone varies with shoot vigor and extension rate, with rapidly growing vigorous shoots potentially showing apical characteristics extending to nodes 15-18 while slow-growing shoots may limit apical designation to nodes 10-12. Apical leaves show greatest developmental heterogeneity, ranging from tiny emerging primordia to approaching-mature leaves depending on position within the apical zone and shoot extension status. Functional characteristics change dramatically if shoot extension ceases, with formerly apical leaves completing maturation and transitioning to source function contributing to vine carbon balance rather than consuming resources. Environmental conditions during apical leaf development determine final characteristics through phenotypic plasticity, with light, temperature, and resource availability affecting size, anatomy, and functional capacity. Management practices including shoot topping (removal of shoot tips) eliminate the youngest apical leaves, redirecting resources from continued extension growth toward fruit development and lateral shoot suppression.
Spitzenzone Weinblatt
apikales Blatt
distal position leaf
oberes Spross-Blatt
tip zone leaf
upper shoot leaf
Based on Palliotti & Cartechini (2000) "Developmental changes in gas exchange activity in flowers, berries, and tendrils of field-grown Cabernet Sauvignon" American Journal of Enology and Viticulture 51=317-323 Poni et al. (2009) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 60=397-407 Intrieri et al. (2008) "Grapevine growth and development" in Biology of the Grapevine, Cambridge University Press pp. 44-81 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 95-136.
Apical Leaf
An apical lobe on a mature leaf at node 6 of Vitis vinifera cv. Syrah in the Rhône Valley, France, positioned at the blade apex representing the terminal extension of the median vein, measuring 6.8 cm length from the sinus base (junction with upper lateral lobes) to the apex representing the longest lobe dimension on this leaf, width of 4.2 cm at the widest point creating elongated lobe shape characteristic of this cultivar, separated from adjacent upper lateral lobes by upper lateral sinuses of 3.2 cm depth (left) and 3.0 cm depth (right) creating well-defined lobe boundaries, vascularized by the median vein extending centrally through the lobe and terminating near the apex, exhibiting acute apex with approximately 40-degree point angle and slightly extended acuminate tip, lobe margins showing convex curvature with heterogeneous tooth pattern, lobe area representing approximately 28% of total blade area making it the largest lobe, prominent elongated appearance contributing to overall leaf length of 16.5 cm and characteristic Syrah leaf outline, measured at BBCH stage 81 (beginning of berry ripening) in early August, representing typical apical lobe morphology with pronounced development diagnostic for this cultivar.
The terminal lobe at the distal end of the leaf blade aligned with the median vein in Vitis vinifera, characterized by position at the blade apex representing the continuation of the central axis, direct vascularization by the median vein extending from petiolar junction through the lobe to its apex, typically the largest and most prominent lobe often exceeding lateral and basal lobes in length and sometimes width, central role in determining overall leaf length and outline shape, apex characteristics including acute, obtuse, or rounded point configurations serving as diagnostic features, variable development ranging from poorly defined in weakly lobed cultivars to prominently elongated in deeply dissected varieties, and functional significance as the primary terminal extension of the blade contributing substantially to total leaf area and determining maximum blade length. The apical lobe represents the terminal continuation of the primary leaf axis established by the median vein, developing as the central growth zone during leaf expansion, exhibiting cultivar-specific characteristics including relative size compared to lateral lobes, degree of separation from adjacent lateral lobes through upper lateral sinuses, apex point characteristics, and overall prominence, serving as important ampelographic diagnostic feature particularly when combined with lateral and basal lobe characteristics.
Apical lobes are distinguished from lateral and basal lobes by terminal position aligned with median vein and typically largest size. The apical lobe constitutes the primary terminal extension determining maximum blade length, with cultivar variation from poorly developed apical lobes barely distinguishable from adjacent laterals (some Gewürztraminer clones) to prominently elongated apical lobes substantially exceeding lateral dimensions (Syrah, Mourvèdre). Apex characteristics show diagnostic importance, with acute points (angle 60 degrees), or rounded configurations serving as identification features. Some cultivars show characteristic acuminate (sharply tapering) apical lobe tips creating distinctive appearances. Separation from lateral lobes varies with upper lateral sinus depth, with shallow sinuses creating poorly defined apical lobes integrated with lateral lobes, while deep sinuses create prominent isolated apical lobes. Relative size compared to lateral lobes shows cultivar specificity, with some varieties showing apical dominance (apical substantially larger) while others show more balanced lobe sizes. Functional contribution involves substantial area provision often 25-35% of total blade, and determination of maximum leaf length affecting light interception geometry. Developmental priority often shows apical dominance with early preferential growth, though final relative size depends on cultivar programming.
Apikallappen
Endlappen
Spitzenlappen
central apical lobe
terminal lobe
tip lobe
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 58-89 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 56-78.
Apical Lobe
An apical primary vegetative bud at node 16 on a vigorous cane of Vitis vinifera cv. Syrah, measuring only 2.1mm diameter compared to 3.2mm for mid-cane primary buds at nodes 6-8, dissection revealing incompletely differentiated primordia with only 6 leaf initials formed and no detectable inflorescence structures, typical of late-formed buds that initiated development in late August when declining photoperiod and temperatures curtailed normal differentiation progression, with this bud routinely removed during winter cane pruning that retains only nodes 1-12 for fruiting and renewal wood.
A primary vegetative bud located at or near the apical (distal) end of annual canes, typically at nodes beyond position 12-15, characterized by reduced size compared to mid-cane primary buds, often incomplete primordial differentiation due to late-season formation with insufficient time for full development before growth cessation, and predominance of vegetative over reproductive structures even when some inflorescence initials are present. Apical primary vegetative buds reflect the declining fruitfulness gradient from mid-cane toward cane tips, resulting from progressively later differentiation timing, reduced carbohydrate availability in distal positions, and often insufficient chilling accumulation for normal development, making these positions typically unsuitable for retention during winter pruning in production-focused training systems.
Apical primary vegetative buds exemplify the node position effect on bud fruitfulness, with fruitfulness typically peaking at mid-cane positions (nodes 4-8 or 5-10 depending on cultivar) and declining toward both basal and apical extremes. The distal decline results from different factors than basal decline= apical buds form later (August-September versus June-July for basal buds), experience cooler temperatures during differentiation, have longer vascular pathways with potentially limited resource supply, and may enter dormancy before completing normal developmental programs. Pruning systems universally recognize apical decline, with cane length limits and spur positioning deliberately avoiding retention of apical positions except as a deliberate vigor-reduction strategy in excessively vigorous vines.
Spitzenknospe
apikale vegetative Primärknospe
distal primary vegetative bud
terminal primary vegetative bud
Based on Vasconcelos et al. (2009) "The flowering process of Vitis vinifera= a review" American Journal of Enology and Viticulture 60=411-434 May (2000) "From bud to berry, with special reference to inflorescence and bunch morphology in Vitis vinifera L." Australian Journal of Grape and Wine Research 6=82-98 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115.
Apical Primary Vegetative Bud
An apical shoot at the terminal position of a vertical cordon arm in Vitis vinifera cv. Pinot Noir trained in a VSP (Vertical Shoot Positioning) system in Burgundy, France, showing vigorous upward growth of 120 cm during the growing season with 15 fully expanded leaves and active apical meristem, exhibiting strong suppression of lateral bud development in the lower 80 cm of the shoot, measured at BBCH stage 75 (berries pea-sized) in late June.
A shoot originating from or occupying the terminal position of a grapevine axis, characterized by its location at the growing tip of a cane, arm, or trunk, exhibiting strong apical dominance that suppresses lateral bud development along the same axis through basipetal gradients of auxin and other growth regulators. Apical shoots in Vitis vinifera represent the primary vectors of vegetative extension growth, determining the overall architecture and spatial occupation of the vine canopy through their growth direction, vigor, and branching patterns. These shoots play critical roles in establishing training system geometry, particularly in vertical shoot positioning systems where apical shoot management directly influences light interception, air circulation, and mechanization efficiency. The apical position confers both developmental advantages through preferential resource allocation and increased vulnerability to environmental stresses including frost damage, wind breakage, and herbivore predation due to the concentration of meristematic tissues and tender growth at exposed positions.
Distinguish apical shoots from lateral shoots based on positional and developmental criteria rather than purely morphological features, as lateral shoots can occasionally exhibit apical-like vigor when released from apical dominance. Apical shoots include both main apical shoots (original terminal growth) and dominant apical shoots (laterals that have assumed apical dominance after topping or damage). The term encompasses both actively growing terminal shoots and those that have completed extension growth but retain their terminal position. Not to be confused with shoots at apical node positions, which may be lateral in developmental origin.
Apikaltrieb
Endtrieb
Spitzentrieb
apex shoot
terminal shoot
tip shoot
Based on Cline (1997) "Concepts and terminology of apical dominance" American Journal of Botany 84=1064-1069 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 60-85 Intrieri et al. (2011) "Effect of canopy management on whole-vine photosynthesis" Acta Horticulturae 526=251-262 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 89-112.
Apical Shoot
An apical terminal inflorescence at node 7 on a shoot of Vitis vinifera cv. Grenache in southern Rhône Valley, France, positioned 3.8 cm behind the shoot apex representing apical zone location, measuring 13.5 cm length representing intermediate dimensions, occurring as the third and final inflorescence on this shoot following the primary at node 3 (18.5 cm, ~620 flowers) and secondary at node 5 (15.2 cm, ~440 flowers), exhibiting compound paniculate architecture with central rachis bearing 3-4 orders of branching, bearing approximately 340 flowers representing reduced number compared to basal inflorescences, positioned as the distal-most reproductive structure with only vegetative shoot extension above, showing phenological development at BBCH stage 61 (beginning of flowering, 10% flowers open) compared to stage 65 (full bloom) for the basal primary indicating 4-5 day developmental delay, apical position receiving excellent light exposure and air circulation, determining the total of three inflorescences for this moderately fertile shoot, measured in early June during sequential flowering progression, representing typical apical terminal inflorescence completing the reproductive sequence.
A terminal primary inflorescence positioned at the shoot apex representing the final distal reproductive structure in Vitis vinifera, characterized by location as the ultimate terminal inflorescence when multiple inflorescences occur along the shoot, position at or within 3-5 cm of the shoot tip representing apical zone placement, role as the distal-most reproductive structure determining the terminal extent of the reproductive region, occurrence following one or more basal inflorescences at standard lateral positions creating the sequence from basal primary through intermediate secondaries to apical terminal, dimensions typically intermediate between robust basal primaries and reduced shoot tip terminals often 10-16 cm length, developmental timing often showing slight delay compared to basal inflorescences reflecting sequential development along the shoot, and functional role completing the reproductive sequence and determining total inflorescence number and spatial distribution along the shoot. Apical terminal inflorescences represent the positional terminal in the inflorescence sequence, occurring as the final reproductive structure when shoots produce multiple inflorescences, exhibiting intermediate developmental characteristics between robust basal primaries and resource-limited extreme terminals, and determining the extent of the reproductive zone affecting total potential yield and architectural organization.
Apical terminal inflorescences are distinguished from shoot tip terminals by moderate distance from apex (3-5 cm versus 1-3 cm) and from general terminals by emphasis on position as the final member of a sequence rather than simply terminal placement. The apical designation emphasizes the positional role completing the inflorescence sequence and determining the extent of the reproductive zone. Occurrence requires multiple inflorescences per shoot with fertile cultivars producing 2-3 inflorescences creating the context for apical terminals, while less fertile varieties producing single inflorescences lack distinct apical terminals. Sequential positioning creates the progression from robust basal primary through intermediate secondary to apical terminal, with size and complexity generally decreasing along the sequence though substantial variation exists. Developmental delay compared to basal inflorescences reflects sequential initiation during primordium formation and sequential resource allocation during development, creating staggered flowering phenology potentially extending the pollination window. Functional significance includes determining total inflorescence number affecting potential yield, and establishing the boundary between reproductive and purely vegetative shoot zones affecting architectural organization. Management practices including inflorescence thinning often target apical terminals for removal retaining only basal primaries and sometimes secondaries to optimize crop load. Transformation to cluster creates apical cluster positioning affecting final cluster distribution and harvest patterns.
apikale terminale Infloreszenz
distal sequence terminal
distale Sequenz-Terminal
final position inflorescence
ultimate reproductive structure
ultimative reproduktive Struktur
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 349-382 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 309-342 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 537-572.
Apical Terminal Inflorescence
In a Sonoma Chardonnay vineyard, apoplastic unloading intensification was tracked over 7 days in mid-August Year N+1 (BBCH 84) when cell wall invertase activity increased from 35 to 88 units/g FW, VvcwINV expression rose from 55-fold to 105-fold, apoplastic hexose concentrations measured 340 mM combined (from 195 mM at activation), berry Brix accelerated from 13.8° to 18.5° (2.9% per week increasing to 4.1% per week), under warm conditions (daily high 27-30°C, adequate vine water status stem ψ -0.9 MPa supporting phloem function).
Apoplastic_Unloading_Intensification represents the progressive amplification of sucrose release into cell wall space and subsequent enzymatic cleavage to hexoses during the transition from initial to maximum sugar accumulation creating increasingly efficient delivery of importable sugars to mesocarp cells. This intensification occurs during early-to-mid veraison (BBCH 83-84), typically 3-8 days following initial phloem unloading activation, and is characterized by apoplastic sucrose flux increasing from initial activation 60-120 μmol g⁻¹ h⁻¹ to intensification peak 120-250 μmol g⁻¹ h⁻¹ representing 2-4 fold enhancement, cell wall invertase activity rising from activation 20-40 units/g FW to intensification 60-120 units/g through continued gene expression and protein accumulation, and resulting apoplastic hexose concentrations increasing from 150-250 mM to 250-450 mM combined glucose and fructose creating stronger concentration gradients for membrane transport. Mechanisms underlying intensification inclu continued plasmodesmata closure or conductance reduction with callose deposition increasing and symplastic connectivity declining from activation 15-25% to intensification 35°C providing temperature sensitivity that may limit sugar accumulation under heat stress. Apoplastic compartment dynamics show: sucrose concentration in apoplast showing transient increases during peak delivery periods potentially reaching 180-350 mM before invertase cleavage, turnover kinetics with half-life 4-12 minutes indicating rapid cleavage and transport preventing sustained accumulation, resulting glucose and fructose concentrations 250-450 mM combined creating 200-380 mM concentration gradient to cytoplasm (maintained 60-90 mM) driving facilitated diffusion, and spatial distribution with highest concentrations near phloem unloading zones gradually declining toward peripheral mesocarp reflecting diffusion and transport gradients. Cellular responses to intensification inclu mesocarp cells adjacent to vascular bundles showing most intense import activity with vacuolar hexose concentrations reaching 1000-1400 mM during intensification compared to 600-900 mM in peripheral cells, spatial gradient in sugar accumulation creating ripening progression from berry center outward potentially affecting harvest maturity uniformity, and cell wall mechanical properties potentially affected by high apoplastic sugar concentrations creating osmotic effects though this aspect poorly studied. Phloem physiology supporting intensification requires: sieve tube pressure maintained 0.6-1.3 MPa in source leaves declining to -0.3 to -0.6 MPa in berry vascular tissue creating 0.9-1.9 MPa gradient driving mass flow, sucrose concentration in sieve sap 500-850 mM in loading zones declining to 350-600 mM at unloading sites through dilution and unloading, mass flow velocity 20-50 cm per hour in vine phloem delivering sucrose at rates matching berry demand, and phloem structure with companion cells and sieve elements maintaining functionality through ripening despite general tissue softening and structural changes. Source-sink relationships during intensification demonstrate: adequate source leaf area essential with typical 14-18 cm² per g berry FW at this stage supporting intensification without limitation, leaf photosynthetic capacity 8-16 μmol CO₂ m⁻² s⁻¹ providing carbon for sucrose synthesis and export, source-sink distance effects with berries closer to source leaves potentially showing earlier or more intense unloading though vine transport capacity generally adequate for uniform supply, and competition among sinks (vegetative growth declining, other berry clusters, root growth) with berries becoming increasingly dominant sink during ripening concentrating resource allocation toward fruit. Environmental influences on intensification inclu temperature affecting multiple steps with optimal 25-29°C supporting maximum unloading and cleavage rates, cooler conditions 20-25°C reducing intensification rate and potentially extending duration to achieve similar final levels, heat stress >31°C impairing intensification through reduced phloem transport, enzyme denaturation, and membrane dysfunction potentially limiting subsequent sugar accumulation even if conditions improve, and diurnal temperature effects with cooler nights potentially beneficial for phloem transport efficiency versus constant warm temperatures. Water relations effects show: moderate deficit (stem ψ -1.0 to -1.4 MPa) compatible with intensification and potentially enhancing through elevated ABA maintaining gene expression, adequate vine water status required for phloem transport with severe deficit (<-1.5 MPa) impairing phloem function reducing sucrose delivery to apoplast, osmotic relationships complex with high apoplastic sugar creating strong osmotic potential affecting water distribution between symplast, apoplast, and xylem, and berry water import driven largely by vacuolar sugar accumulation creating total berry water potential -1.8 to -2.8 MPa attracting water from vine xylem. Cultivar variation in intensification characteristics shows: rapidly-intensifying varieties achieving maximum unloading within 5-7 days of activation with cell wall invertase reaching 90-120 units/g, moderate cultivars requiring 7-10 days to peak with 70-100 units/g final activity, and slower types needing 10-14 days achieving 60-90 units/g reflecting genetic differences in gene induction kinetics and enzyme synthesis capacity. Coordination with transport intensification demonstrates: apoplastic unloading intensification synchronized with hexose transporter upregulation ensuring products of sucrose cleavage efficiently imported, temporal relationship typically shows unloading intensification leading transporter induction by 1-2 days establishing substrate before maximum import capacity, and feedback regulation where high apoplastic hexoses may enhance transporter expression through sugar sensing mechanisms creating feed-forward amplification. Quality implications of intensification inclu rate and completeness of intensification determining maximum sugar accumulation rates achievable during ripening, inadequate intensification limiting final Brix despite adequate source photosynthesis through bottleneck in unloading-cleavage step, timing relative to other ripening processes affecting compositional balance with early intense unloading potentially producing high-sugar fruit before adequate phenolic or flavor development, and environmental stresses during intensification creating disproportionate effects on final composition through impairment of critical developmental step. Viticultural management during intensification involves: ensuring adequate vine water status supporting phloem function without excess that could dilute berries, maintaining canopy health for continued photosynthesis and sucrose supply, avoiding environmental stresses particularly heat and severe drought that can irreversibly impair intensification, and monitoring intensification progress through berry sampling tracking Brix increases indicating successful amplification of sugar import pathway.
Apoplastic_Unloading_Intensification is distinguished from Phloem_Unloading_Activation by representing amplification (2-4 fold) rather than initiation, from Sucrose_Cleavage_Activation by focusing on unloading flux rather than enzyme activation, and from Active_Sugar_Loading by showing progressive increase rather than sustained maximum. Cell wall invertase: 20-40 → 60-120 units/g FW. Apoplastic hexoses: 150-250 → 250-450 mM. Duration 5-10 days BBCH 83-84. Symplastic connectivity declines to <8-15%. Apoplastic_Unloading_Intensification represents critical amplification step creating substrate flux supporting rapid sugar accumulation.
amplified cell wall sugar release
amplifizierte Zellwand-Zucker-Freisetzung
enhanced apoplastic sucrose delivery
intensified extracellular hexose production
intensivierte extrazelluläre Hexose-Produktion
progressive Entladungs-Zunahme
progressive unloading increase
verstärkte apoplastische Saccharose-Lieferung
Zhang XY, Wang XL, Wang XF, Xia GH, Pan QH, Fan RC, Wu FQ, Yu XC, Zhang DP (2006) A shift of phloem unloading from symplasmic to apoplasmic pathway is involved in developmental onset of ripening in grape berry Plant Physiol 142:220-232 unloading intensification Davies C, Wolf T, Robinson SP (1999) Three putative sucrose transporters are differentially expressed in grapevine tissues Plant Sci 147:93-100 transport coordination Lecourieux F, Kappel C, Lecourieux D, Serrano A, Torres E, Arce-Johnson P, Delrot S (2014) An update on sugar transport and signaling in grapevine J Exp Bot 65:821-832 comprehensive sugar transport regulation.
Apoplastic Unloading Intensification
The appellation designation Barolo DOCG identifies the delimited production zone within the Langhe hills of Piedmont comprising 11 communes (Barolo, La Morra, Castiglione Falletto, Serralunga d'Alba, Monforte d'Alba, and 6 others) totaling approximately 2,100 hectares of registered vineyard, with the designation encoding both the geographic delimitation and the regulatory classification level (DOCG, the highest tier in the Italian quality pyramid) — within this appellation, 181 officially delimited Menzioni Geografiche Aggiuntive (additional geographic mentions) such as Cannubi, Brunate, and Rocche dell'Annunziata provide sub-appellation specificity identifying individual vineyard sites with recognized terroir distinctiveness.
A geographic indication that constitutes the formal name and classificatory position of a delimited viticultural area within an appellation system, identifying a specific geographic unit recognized by a regulatory authority as possessing distinctive terroir characteristics that contribute identifiable qualities to the wines produced within its boundaries.
Appellation_Designation captures the naming and classificatory dimension of geographic indications — the information about what a geographic unit is called and where it sits within a hierarchical classification system. Every major wine-producing country has developed appellation systems with varying degrees of complexity: France's AOC/AOP system (approximately 360 appellations), Italy's DOC/DOCG system (approximately 520 designations), Germany's Anbaugebiet/Bereich/Einzellage hierarchy, Spain's DO/DOCa system, and New World systems such as the US AVA and Australian GI frameworks each implement the general concept of geographic delimitation with region-specific regulatory structures. Subclasses distinguish the formal name, the hierarchical level, and sub-appellation designations within these systems.
Appellationsbezeichnung
Lagenbezeichnung
appellation name
controlled origin name
kontrollierte Herkunftsbezeichnung
wine region designation
EU Regulation 1308/2013 Art. 93-108 INAO (2023) Les signes officiels de la qualité et de l'origine OIV (2015) OIV Guidelines for Geographic Indications for Wines
Appellation Designation
The Burgundy appellation system exemplifies a four-tier hierarchy where each level narrows geographic scope and tightens production regulations — Bourgogne AOC (regional level, approximately 27,000 hectares, maximum yield 60 hl/ha), Gevrey-Chambertin AOC (village level, approximately 410 hectares, maximum yield 50 hl/ha), Gevrey-Chambertin Premier Cru AOC (premier cru level, 26 named vineyard sites totaling 85 hectares, maximum yield 45 hl/ha), and Chambertin Grand Cru AOC (grand cru level, single 12.9-hectare vineyard, maximum yield 37 hl/ha) — with the hierarchy level designation on the label communicating to consumers the presumed quality tier and terroir specificity of the wine.
An appellation designation that specifies the tier or rank of a geographic unit within a structured hierarchical classification system, defining the relative position of the appellation between broader regional designations and more specific site-level designations, with higher hierarchical levels typically implying more restrictive production regulations, smaller geographic scope, and presumed greater terroir specificity.
Appellation_Hierarchy_Level captures the vertical dimension of appellation classification that organizes geographic units into nested tiers of increasing specificity. The structure of appellation hierarchies varies fundamentally between wine-producing countries: France uses a multi-tier system (regional, sub-regional, communal, premier cru, grand cru) most fully developed in Burgundy and Alsace Germany employs Anbaugebiet, Bereich, Großlage, and Einzellage levels Italy uses the IGT-DOC-DOCG pyramid and the United States AVA system is formally non-hierarchical though nesting of smaller AVAs within larger ones creates de facto hierarchy. The hierarchy level communicates quality expectations and price positioning to consumers while imposing correspondingly stricter production constraints on producers at higher tiers.
Appellationshierarchiestufe
Herkunftshierarchie-Rang
Klassifikationsebene
appellation tier
classification level
hierarchy rank designation
INAO (2023) Les signes officiels de la qualité et de l'origine Lewin MW (2010) Wine Myths and Reality, Vendange Press pp: 95-150 EU Regulation 1308/2013 Art. 93 on categories of wine designations
Appellation Hierarchy Level
Appellation requirement for Wachau DAC (established 2020) specifies: geographic delimitation to the Wachau wine-growing district along the Danube between Melk and Krems, permitted cultivars including Grüner Veltliner, Riesling, and Neuburger as principal varieties, maximum yield of 6,750 kg/ha (Steinfeder) to 6,000 kg/ha (Smaragd), minimum must weight thresholds aligned with the Vinea Wachau classification (Steinfeder ≤11.5% alcohol, Federspiel 11.5-12.5%, Smaragd ≥12.5%), sensory evaluation by the regional tasting commission, and mandatory display of the DAC designation on the label. Appellation requirement for Barolo DOCG mandates 100% Nebbiolo from the defined 11 communes, maximum yield of 8,000 kg/ha, minimum alcohol 13.0% vol., minimum aging of 38 months (of which 18 months in wood) for standard Barolo and 62 months (of which 18 months in wood) for Riserva, chemical analysis verification, and blind sensory panel approval — the requirements encoding the historical quality standards that have defined Barolo's prestige since the DOCG designation in 1980.
Appellation requirement is a label regulation entity that documents the legal conditions that must be met for a wine to carry a specific geographic origin designation on its label, encompassing the applicable designation category (EU PDO/PGI, French AOC/IGP, Italian DOCG/DOC/IGT, Austrian DAC/Qualitätswein, German Prädikatswein/QbA, US AVA, Australian GI), the production requirements specified in the appellation's cahier des charges or product specification (permitted cultivars, maximum yield, minimum must weight, permitted oenological practices, aging requirements, sensory approval), the geographic delimitation (the defined territory from which grapes must originate), the minimum percentage of grapes from the declared appellation (typically 85-100% depending on jurisdiction), and the administrative approval process (chemical analysis, sensory panel tasting, documentation verification), providing the regulatory framework that protects geographic origin claims as quality indicators and cultural heritage designations.
Appellation requirement documentation captures the most elaborate and culturally consequential regulatory framework in the wine world — the system of protected origin designations that simultaneously serves as quality guarantee, fraud prevention mechanism, cultural heritage protection, and marketing tool. The European PDO system traces its conceptual origins to the 1935 French AOC legislation and its philosophical roots to the concept of terroir — the proposition that specific geographic origins produce wines of distinctive and recognizable character that merit legal protection from misappropriation. The requirements encoded in appellation specifications represent codified collective knowledge — the permitted cultivars reflect centuries of empirical observation about which varieties express the appellation's character, the yield limits reflect the established inverse relationship between yield and quality, and the aging requirements reflect the maturation needs of the specific wine styles produced. The stringency gradient across designation levels creates a quality hierarchy — from the relatively permissive PGI/IGP level (broad geographic scope, diverse cultivar permission, minimal production restrictions) through the intermediate PDO level (defined territory, restricted cultivars, moderate production constraints) to the most prestigious designations (DOCG, Grand Cru, DAC) imposing the strictest requirements on every aspect of production. The tension between appellation rigidity and producer innovation is a persistent feature of the system — the most creative winemakers sometimes chafe against appellation constraints that they perceive as limiting quality expression (the Italian Super Tuscan phenomenon, Austrian producers opting out of DAC regulations to pursue non-conforming styles), while appellation defenders argue that regulatory discipline maintains the collective brand value that benefits all producers within the designation. The emerging challenge of climate change adaptation within appellation frameworks — how to maintain the identity-defining production requirements of an appellation when the climatic conditions that shaped those requirements are changing — represents one of the most pressing regulatory questions in contemporary European viticulture.
AOC requirement
Appellationsanforderung
DAC-Anforderung
Herkunftsbezeichnungsvorschrift
Herkunftsschutzvorschrift
PDO/PGI requirement
Ursprungsbezeichnungsregel
appellation regulation
geographic indication rule
origin designation requirement
European Parliament (2013) Regulation (EU) No 1308/2013, Articles 93-108 INAO (2024) Cahier des charges des appellations d'origine contrôlées Consorzio di Tutela Barolo Barbaresco (2010) Disciplinare di produzione Barolo DOCG Österreichische Weinmarketinggesellschaft (2020) Wachau DAC Verordnung
Appellation Requirement
A Pinot Noir vine within the commune of Gevrey-Chambertin in Burgundy bearing an appellation-specific role is subject to INAO regulations specifying maximum yields of 37-58 hl/ha depending on classification tier, minimum planting density of 9,000 vines/ha, prohibited irrigation except by derogation, and mandatory hand harvesting for Grand Cru designations, with the vine's precise cadastral parcel determining whether fruit enters the Gevrey-Chambertin Village, Premier Cru, or Grand Cru production stream.
A regional production role that inheres in a grapevine situated within a legally delimited geographic appellation whose boundaries, permitted cultivars, viticultural practices, and quality requirements are codified in regulatory statutes, entitling the resulting wine to bear the appellation designation upon compliance with all production specifications.
Appellation_Specific_Role reflects the regulatory framework governing geographic wine designations worldwide, including the French AOC/AOP system, Italian DOC/DOCG, Spanish DO/DOCa, German Prädikatswein, and New World Geographic Indications (GI). The role is defined by regulatory compliance rather than intrinsic vine characteristics: a vine gains or loses this role through changes in appellation boundaries, regulatory amendments, or compliance status rather than through any biological transformation. Subclasses represent the principal hierarchical tiers found across major European classification systems, from Grand Cru to regional designations.
Appellations-Rolle
Lagenklassifizierungs-Rolle
appellation-classified role
controlled origin role
denomination-specific role
herkunftsgeschützte Produktionsrolle
INAO (2023) Cahiers des charges des appellations d'origine contrôlées EU Regulation 1308/2013 Art. 93-108 on wine designations of origin Robinson J and Harding J (2015) The Oxford Companion to Wine 4th Ed, Oxford University Press pp: 34-38
Appellation Specific Role
In a Mosel Riesling vineyard, approaching sugar maximum was modeled over 18 days in late September-October Year N+1 (BBCH 88-89) when berry Brix increased from 23.8° to 26.2° following exponential decay Brix(t) = 27.1 - 3.3×exp(-t/11.5) with fitted maximum Brix_max = 27.1° and time constant 11.5 days (r² = 0.94), half-time 8 days, daily increment declining from 0.18% to 0.05% per day, enabling harvest forecast of plateau achievement within 14 days (actual 12 days excellent agreement), under cool German autumn conditions (daily high 16-20°C extending gradual approach).
Approaching_Sugar_Maximum represents the asymptotic trajectory toward cultivar-specific peak berry sweetness during late ripening when progressively declining accumulation rates create gradually slowing Brix increase curve approaching but not yet achieving final equilibrium defining the transition period between active accumulation and complete plateau signaling imminent harvest maturity. This approaching phase occurs during late-to-final ripening (BBCH 87-89), typically days 45-60 post-veraison spanning 12-20 days, and is characterized by berry Brix rising from 23-27° at approach onset to near-maximum 25-30° at conclusion representing 1.5-4.0° gain substantially reduced from earlier phases, daily Brix increase declining from 0.12-0.22% per day early approach to 0.03-0.08% per day late approach showing clear deceleration, asymptotic curve fitting showing r² >0.90-0.96 for exponential decay model Brix(t) = Brix_max - (Brix_max - Brix_0) × exp(-k×t) with rate constant k determining approach velocity, and time constant τ = 1/k typically 8-18 days indicating characteristic timescale for approaching 63% of remaining distance to maximum. Mathematical characterization of approach includes instantaneous accumulation rate dBrix/dt declining approximately linearly with distance from maximum (Brix_max - Brix_current) consistent with first-order kinetics, half-time to maximum (time to achieve 50% of remaining gain) typically 5-12 days from approach onset indicating rapid initial approach then slowing, 90% approach time (achieving 90% of final maximum) requiring 15-25 days demonstrating asymptotic nature where final increments require extended duration, and residual accumulation potential (Brix_max - Brix_current) declining from 3-6° approach onset to <0.5-1.5° late approach indicating most gain achieved early in approaching phase. Physiological basis for asymptotic approach includes vacuolar hexose concentrations 1750-2450 mM approaching saturation levels 2000-2600 mM cultivar-dependent creating increasingly unfavorable thermodynamics for continued accumulation, concentration gradients (apoplast-to-vacuole) declining progressively as vacuolar levels rise reducing driving force according to Fick's law where flux proportional to gradient, transporter capacity potentially declining but more importantly operating at progressively lower efficiency as substrate gradients reduce even if protein levels maintained, and metabolic equilibrium approaching where import rate equals respiratory consumption rate plus any export or interconversion with net vacuolar accumulation approaching zero. Thermodynamic analysis shows free energy change for hexose transport from apoplast (280-400 mM, pH 5.5-6.5) to vacuole (1800-2500 mM, pH 2.8-3.2) including membrane potential (-120 to -180 mV cytoplasm-negative) and pH gradient (4.5-5.0 units vacuole-acidic) calculating ΔG approaching zero or slightly positive late approach indicating thermodynamic barrier limiting further accumulation, osmotic work required for continued accumulation against concentration gradient increasing exponentially with vacuolar concentration following ΔG = RT ln(C_vacuole/C_apoplast) approaching or exceeding available cellular energy, and maximum achievable concentration gradient determined by coupling stoichiometry with H⁺-ATPase (ATP hydrolysis provides ~50 kJ/mol driving force enabling ~1000-fold gradient theoretically but practical limits lower due to inefficiencies). Molecular regulation during approach demonstrates transporter gene expression declining gradually VvHT1, VvHT2 from 80-120 fold to 55-90 fold above baseline as ripening program downregulates though levels remain substantial, protein degradation potentially accelerating with increasing ubiquitination targeting older transporters for proteasomal removal, post-translational modifications possibly reducing activity through phosphorylation at inhibitory sites or conformational changes reducing substrate affinity though mechanisms speculative in grape, and cellular signaling potentially detecting high vacuolar sugar triggering feedback inhibition through hexokinase-independent sensors in tonoplast or cytoplasm reducing transport gene expression or activity. Environmental factors affecting approach velocity include temperature with warmer conditions 26-30°C maintaining higher rates accelerating approach (shorter time constant) while cooler 18-24°C extending approach duration, declining source photosynthesis with autumn progression reducing substrate availability accelerating approach to plateau independent of thermodynamic limits as supply rather than demand becomes limiting, water status with late-season deficit potentially hastening approach through reduced phloem delivery and berry expansion while adequate water extending gradual approach, and canopy health with vigorous vines maintaining photosynthesis supporting slower asymptotic approach while senescent canopies forcing rapid approach as substrate depletes. Cultivar variation in approach characteristics shows high-Brix varieties (Gewürztraminer, Sémillon approaching 27-30° maximum) displaying extended gradual approach 16-24 days with time constant τ = 12-18 days indicating slow final accumulation continuing, moderate cultivars (Chardonnay, Merlot approaching 23-27° maximum) showing intermediate approach 12-18 days with τ = 8-14 days, and lower-Brix types (Pinot Noir cool climate approaching 21-25° maximum) exhibiting rapid approach 8-14 days with τ = 5-10 days reflecting genetic differences in maximum capacity and regulatory control determining approach kinetics. Predictive modeling using approach dynamics enables extrapolation from current trajectory fitting exponential decay model to estimate Brix_max and time to plateau within ±0.5-1.5° Brix and ±3-7 days accuracy, recognition that asymptotic approach means final 10% of accumulation requires 50-80% of total approach time informing harvest timing decisions, and understanding that factors disrupting approach (weather events, disease, stress) may prevent achieving theoretical maximum requiring harvest at current levels rather than waiting for completion. Quality implications include maximum sugar level determining wine alcohol potential with each 1° Brix yielding ~0.55-0.60% ethanol, approach timing relative to phenolic and flavor maturity affecting optimal harvest particularly reds where extended hang time during slow approach may improve quality despite minimal sugar gain, and recognition that asymptotic approach creates extended harvest window during late approach phase where 0.5-1.5° Brix difference over 7-14 days allows flexibility for integrating multiple maturity parameters. Harvest strategies during approach include monitoring approach trajectory through frequent sampling enabling maximum prediction and optimal timing determination, integrating sugar approach with acid plateau recognizing both processes asymptotic and coordinated, phenolic assessment particularly reds ensuring tannin maturity achieved during approach period, sensory evaluation detecting flavor development potentially continuing during slow sugar approach, and weather monitoring balancing quality optimization during extended approach against late-season risks (rain, disease, frost). Viticultural considerations include minimal intervention during approach as process physiologically determined and environmental manipulation limited effectiveness, disease control critical as high-sugar berries during extended approach vulnerable to infection, harvest preparation advancing as approach signals imminent maturity, and recognition that approach completion (plateau achievement) may occur on-vine or post-harvest during transport/processing with minimal practical difference if harvest timing appropriate.
Approaching_Sugar_Maximum is distinguished from Final_Sugar_Accumulation by focusing on asymptotic trajectory (exponential decay model) rather than overall late phase, from Sugar_Plateau by showing continued though minimal accumulation (<0.5-1.5° remaining) rather than equilibrium, and from peak accumulation by exhibiting clear deceleration (0.03-0.22% per day declining) rather than maximum rates. Duration 12-20 days BBCH 87-89. Time constant τ typically 8-18 days. Thermodynamic barriers approaching. Approaching_Sugar_Maximum represents measurable asymptotic approach enabling harvest timing prediction.
Dezeleration Richtung Gleichgewicht
End-Annäherung zu Zucker-Kapazität
asymptotic sweetening
asymptotische Süßung
decelerating toward equilibrium
final approach to sugar capacity
nearing maximum Brix
sich näherndes Maximum-Brix
Coombe BG (1992) Research on development and ripening of the grape berry Amer J Enol Vitic 43:101-110 asymptotic approach Keller M (2020) The Science of Grapevines 3rd Ed pages 227-228 approaching maximum Dai ZW, Ollat N, Gomès E, Decroocq S, Tandonnet JP, Bordenave L, Pieri P, Hilbert G, Kappel C, van Leeuwen C, Vivin P, Delrot S (2011) Ecophysiological, genetic, and molecular causes of variation in grape berry weight and composition Plant Physiol 155:519-532 approach dynamics and modeling.
Approaching Sugar Maximum
Aquaporin-mediated water flux accounts for 50-80% of root hydraulic conductivity in grapevines, with VvPIP1 and VvPIP2 subfamily members showing differential expression across tissues and environmental conditions. VvPIP2 1 expression in roots increases 3-5 fold during early morning, coinciding with peak transpirational demand, and declines 60-80% under drought stress through ABA-mediated transcriptional downregulation. Rootstock 110 Richter shows constitutively lower aquaporin expression (40-60% of SO4 levels), contributing to its conservative water use strategy. Aquaporin gating through phosphorylation and cytosolic pH changes enables rapid (minutes) adjustment of hydraulic conductance independent of transcriptional regulation.
A water uptake disposition that is the inherited capacity of a Vitis vinifera plant to regulate transmembrane water flux through aquaporin water channel proteins (plasma membrane intrinsic proteins, PIPs tonoplast intrinsic proteins, TIPs), with genotype-specific expression levels and gating responses modulating root and leaf hydraulic conductance.
Aquaporin activity represents the primary regulatory node for membrane water transport, enabling fine-tuned adjustment of hydraulic conductance in response to environmental signals. The trait underlies genotypic differences in water use strategy, with isohydric cultivars showing strong aquaporin downregulation under stress versus anisohydric types maintaining expression. VvPIP and VvTIP gene families contain 10-15 members with tissue-specific expression patterns, enabling differential regulation of root uptake, xylem loading, and leaf cell hydration. Aquaporins also transport CO₂, affecting mesophyll conductance and photosynthetic efficiency.
Aquaporin-Aktivität
Membranwasserpermeabilität
Wasserkanalexpression
aquaporin expression trait
membrane water permeability
water channel activity
Vandeleur RK et al. (2009) The role of plasma membrane intrinsic protein aquaporins in water transport through roots: diurnal and drought stress responses reveal different strategies between isohydric and anisohydric cultivars of grapevine. Plant Physiology 149:445-460 Shelden MC et al. (2009) Expression and functional characterisation of plasma membrane intrinsic protein aquaporins in grapevine. Functional Plant Biology 36:1065-1075
Aquaporin Activity Disposition
Aquaporin activity assessment in grapevine roots under water stress reveals the regulatory shutdown protecting against dehydration, with well-watered roots showing high aquaporin activity (membrane water permeability Lp = 4.5 × 10⁻⁷ m/s/MPa), declining within 2 hours of ABA application to 1.2 × 10⁻⁷ (rapid gating response), and further declining to 0.6 × 10⁻⁷ after 24 hours of sustained stress (reduced expression), demonstrating the multi-level aquaporin regulation that dynamically adjusts root hydraulic conductivity in response to water status signals.
Aquaporin activity quality is a root hydraulic conductivity quality that represents the functional state of aquaporin water channel proteins in regulating transcellular water movement, characterizing the membrane permeability component of hydraulic conductivity that enables rapid and dynamic regulation of water flux through biological membranes in roots and other tissues responding to developmental, environmental, and stress signals. Within the BFO framework, this quality specifically addresses the molecular regulation dimension of water transport, distinguishing aquaporin-mediated conductivity from anatomical pathway conductivity while recognizing that aquaporins provide the mechanism for rapid physiological control of hydraulic conductivity that cannot be achieved through anatomical modifications alone. The ontological significance lies in aquaporin activity providing the regulatory flexibility that enables vines to adjust water transport capacity on timescales of minutes to hours, where gating of existing aquaporin channels and expression changes over longer periods allow hydraulic properties to respond to water status, hormones, nutrients, temperature, and circadian rhythms, creating a dynamic hydraulic system rather than a fixed plumbing network. The molecular basis for aquaporin activity involves the plasma membrane intrinsic protein (PIP) family that forms selective water channels in cell membranes, with activity regulated through gating (phosphorylation state, pH, calcium concentration affecting channel opening), expression (transcriptional regulation of aquaporin genes), and localization (membrane trafficking affecting channel density), providing multiple levels of control over membrane water permeability. The physiological significance of aquaporin regulation extends beyond simple water transport to include roles in osmoregulation, nutrient uptake, and stress signaling, with aquaporin activity affecting not only hydraulic conductivity but also the integration of water transport with other physiological processes. The stress responses of aquaporin activity show complex patterns, with short-term water stress often reducing aquaporin activity (potentially protective by limiting dehydration), while longer-term adaptation may involve increased expression to maintain water supply, and with temperature, salinity, and nutrient stresses each producing specific aquaporin responses affecting vine hydraulic function. Understanding aquaporin activity quality enables appreciation of the molecular control of vine water relations, interpretation of hydraulic responses to environmental signals, and potential targets for genetic improvement of water use efficiency through modified aquaporin function.
Aquaporin_Activity_Quality addresses the molecular regulation of membrane water permeability through water channel proteins. This quality enables rapid physiological control of hydraulic conductivity unavailable through anatomical modifications. The class supports molecular water relations assessment and interpretation of rapid hydraulic responses to environmental signals.
Aquaporinaktivität
PIP activity
PIP-Aktivität
Wasserkanaltätigkeit
aquaporin function
Maurel C et al. (2015) Aquaporins in plants. Physiological Reviews 95(4): 1321-1358 Vandeleur RK et al. (2009) The role of plasma membrane intrinsic protein aquaporins in water transport through roots. Plant and Soil 323(1-2): 167-183
Aquaporin Activity Quality
Arbuscular mycorrhizal colonization intensity varies from 15-70% of root length colonized across grapevine genotypes under identical soil conditions, with rootstocks showing greater variation than scions. High-colonization rootstocks including 1103 Paulsen and SO4 support 50-70% root colonization, while low-colonization types such as 41B and 140 Ruggeri typically show 20-40% colonization. Colonization involves sequential stages of spore germination, hyphal branching toward roots (stimulated by strigolactone exudation), appressorium formation, cortical cell penetration, and arbuscule development within 10-21 days post-contact. Root colonization peaks during active root growth phases (spring and post-harvest) and declines during summer drought and winter dormancy.
A mycorrhizal association disposition that is the inherited capacity of a Vitis vinifera plant to support arbuscular mycorrhizal fungal colonization within root cortical cells, with genotype-specific colonization levels determined by root exudate signaling, cell wall permeability, and immune system modulation enabling fungal entry and arbuscule formation.
Arbuscular mycorrhizal colonization represents the structural foundation of the mycorrhizal symbiosis, with arbuscules serving as the primary nutrient exchange interface between fungus and plant. Colonization level does not always correlate with functional benefit, as colonization quality (arbuscule vitality, hyphal network extent) may be more important than percentage colonization. Genotypic differences in colonization reflect variation in strigolactone production, root cortex accessibility, and immune system responses to fungal entry. High colonization capacity is generally advantageous but may impose carbon costs under high soil fertility where mycorrhizal benefits are minimal.
AMF colonization capacity
AMF-Kolonisierungsvermögen
Mykorrhiza-Besiedlung
Wurzelkolonisations-Disposition
mycorrhizal receptivity
root colonization trait
Schreiner RP (2003) Mycorrhizal colonization of grapevine rootstocks under field conditions. American Journal of Enology and Viticulture 54:143-149 Nogales A et al. (2019) Response of mycorrhizal grapevine to Armillaria mellea inoculation: disease development and polyamines. Plant and Soil 440:371-385
Arbuscular Mycorrhizal Colonization Disposition
Canopy management in Napa Valley Cabernet Sauvignon targets an optimal leaf area to fruit weight ratio of 10-15 cm² per gram of fruit, achieved through shoot thinning (retaining 4-6 shoots per linear meter of cordon), leaf removal in the fruit zone (exposing clusters to 20-30% ambient light), and hedging to maintain leaf area index between 2.5 and 3.5, balancing photosynthetic capacity against excessive shading and disease pressure.
Area_Quality represents the category of morphological qualities that quantify two-dimensional spatial extent of grapevine organ surfaces and cross-sections. Area measurements integrate two linear dimensions to characterize the planar extent relevant to physiological function, light interception, gas exchange, and evaporative surfaces. This category encompasses three primary measurement domains: Surface_Area_Quality capturing the external boundary extent of three-dimensional organs Cross_Sectional_Area_Quality representing the two-dimensional extent revealed by planar cuts through organs and Leaf_Area_Quality specifically addressing the photosynthetically active laminar surface that dominates canopy function. Area qualities are central to grapevine ecophysiology because most physiological processes scale with surface area rather than linear dimensions. Photosynthetic capacity depends on total leaf area and its distribution within the canopy light environment. Transpirational water loss occurs across leaf and berry surfaces in proportion to their area and stomatal density. Skin surface area relative to berry volume determines the extractability of phenolic compounds and aromatic precursors during winemaking. Area measurements require more sophisticated approaches than linear dimensions, including leaf area meters using optical transmission, image analysis systems for irregular surfaces, geometric modeling for three-dimensional organs, and gravimetric estimation using specific leaf area relationships. The relationship between area and linear dimensions follows predictable geometric scaling: doubling linear dimensions quadruples area, which has profound implications for how organ size affects physiological function. Area-based indices including leaf area index (LAI, total leaf area per unit ground area) and exposed leaf area per unit fruit mass provide integrative measures of canopy-fruit balance that guide viticultural management decisions.
Area_Quality is positioned as a direct subclass of Size_Quality, parallel to Linear_Dimension_Quality, Volume_Quality, and Mass_Quality. Area measurements provide the basis for numerous derived physiological parameters including stomatal conductance (mmol m⁻² s⁻¹), photosynthetic rate (μmol CO₂ m⁻² s⁻¹), and transpiration rate (mmol H₂O m⁻² s⁻¹). The distinction between projected area (two-dimensional shadow) and actual surface area (accounting for three-dimensional curvature) must be specified in measurement protocols. PATO_0001323 (area) provides the upper-ontology alignment.
Arealqualität
Flächenqualität
Oberflächenausmaß
areal quality
planar measurement quality
surface extent quality
two-dimensional extent quality
zweidimensionale Ausdehnungsqualität
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 119-152 Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles, Adelaide pp.: 38-47
Area Quality
Arginine content quality management in Japanese Koshu addresses the ethyl carbamate concerns heightened by warm storage conditions in humid Asian climates, with grape arginine concentrations of 800-1500 mg/L (reflecting high-nitrogen fertilization traditions) creating substantial precursor pools winemakers employ acid urease treatment post-fermentation to hydrolyze urea before it can react with ethanol, reducing ethyl carbamate in Koshu wines from potentially problematic levels (above 30 μg/L) to below regulatory thresholds (15 μg/L Canadian standard), demonstrating precursor management through winemaking intervention when viticultural arginine control is impractical.
Arginine_Content_Quality represents the chemical quality characterizing the concentration of the amino acid arginine in grape must, capturing the typically second most abundant amino acid that provides substantial yeast assimilable nitrogen while also serving as precursor for potentially problematic ethyl carbamate formation in wine. Arginine is a basic amino acid with a guanidinium side chain that provides high nitrogen content (four nitrogen atoms per molecule, contributing 32% nitrogen by mass), making it particularly important for yeast nutrition on a per-molecule basis. In grape must, arginine concentration typically ranges from 100-1500 mg/L, comprising 15-40% of total free amino acids and providing 20-50% of total YAN in many musts. Arginine accumulation increases with vine nitrogen status, making it responsive to nitrogen fertilization and serving as an indicator of nitrogen nutrition level. During fermentation, yeast catabolize arginine through the arginase pathway, releasing urea as an intermediate that can react with ethanol under wine storage conditions (particularly at elevated temperature) to form ethyl carbamate, a suspected carcinogen. This ethyl carbamate pathway makes arginine content a food safety consideration, with regulatory limits on ethyl carbamate in wine driving interest in arginine management. Yeast strains vary in their urea excretion during arginine metabolism, with low-urea-producing strains reducing ethyl carbamate risk from high-arginine musts. Viticultural arginine management through moderate nitrogen fertilization and cultivar selection provides preventive control, while yeast strain selection and post-fermentation urease treatment offer winemaking interventions. The balance between arginine's nutritional value and ethyl carbamate risk creates quality management complexity.
Arginine_Content_Quality addresses an amino acid with dual significance as major YAN contributor and ethyl carbamate precursor. The food safety dimension distinguishes arginine from other amino acids in quality importance. The arginase pathway producing urea operates in all Saccharomyces cerevisiae but varies in urea excretion characteristics among strains. The nitrogen fertilization responsiveness of arginine makes it amenable to viticultural management, providing a primary control point for ethyl carbamate risk. Arginine analysis employs standard amino acid HPLC methods, with its basic nature affecting separation behavior on different column chemistries.
Arginingehalt
Ethylcarbamat-Vorläuferqualität
L-Arginin-Konzentration
L-arginine concentration
arginine level
basic amino acid content
basischer Aminosäuregehalt
ethyl carbamate precursor quality
Ough CS et al. (1988) Ethyl Carbamate Precursors in Grape Juice and Wines. American Journal of Enology and Viticulture 39(2): 139-142 Spayd SE et al. (1994) Nitrogen Fertilization of White Riesling Grapes in Washington. Must and Wine Composition. American Journal of Enology and Viticulture 45(1): 34-42
Arginine Content Quality
A permanent arm on Vitis vinifera cv. Cabernet Sauvignon in bilateral cordon training system, extending 85cm horizontally from trunk along trellis wire at 90cm height, measuring 32mm diameter at trunk junction tapering to 22mm at terminus, showing gray-brown thick bark with visible lenticels and previous pruning scars, bearing 14 two-bud spurs spaced 12-15cm apart along upper surface, approximately 7 years old based on growth ring counting at cross-section, supporting estimated 28 fruiting shoots and 45-55 clusters totaling 12-15kg fruit when fully productive, representing well-established permanent structure requiring minimal annual modification beyond spur renewal and occasional repositioning.
A permanent woody structural component of Vitis vinifera training systems, originating from the trunk or head and extending horizontally or at angles to support fruiting positions, characterized by multi-year age (typically 2-15+ years old), substantial diameter (15-40mm or more), complete lignification with thick bark and secondary xylem development, bearing fruiting canes or spurs along its length, and serving as architectural framework connecting trunk to crop-bearing wood while distributing vine capacity across trellis space. Arms represent the intermediate structural level between permanent trunk (vertical support) and annual fruiting wood (crop production), with their number, length, orientation, and management determining training system architecture, canopy distribution, crop load capacity, and long-term vineyard productivity and sustainability.
Arms are distinguished from trunks (vertical permanent wood connecting roots to arms), canes (annual fruiting wood), and cordons (a specific arm training configuration), though terminology varies regionally with "cordon" sometimes used synonymously with "arm" and other times reserved for specific horizontal permanent arm architectures. Arm management involves initial training establishing desired architecture (typically 2-4 years), maintenance pruning managing fruiting positions and preventing excessive lengthening, and eventual renewal or replacement when age-related decline (10-20+ years depending on variety and conditions) reduces productivity or structural integrity. The number and configuration of arms fundamentally determines training system character and suitability for mechanization, labor requirements, and fruit quality potential.
Arm
Kordon
Schenkel
cordon
permanent branch
scaffold
structural arm
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 89-112 Reynolds & Vanden Heuvel (2009) "Influence of grapevine training systems on vine growth and fruit composition= a review" American Journal of Enology and Viticulture 60=251-268.
An arm water shoot on the horizontal cordon of Vitis vinifera cv. Chardonnay in Burgundy, France, emerging from a latent bud at 40 cm from the trunk junction along an 80 cm cordon arm, developing in June following severe spring frost that killed all shoots in a 25 cm section of the cordon, growing 155 cm vertically by August with 17 nodes, leaves averaging 175 cm² area, positioned between two surviving fruiting positions that are each 30 cm distant, creating potential for training as a replacement spur during dormant pruning to restore fruiting position density in the frost-damaged section, or requiring removal if surrounding positions show adequate recovery and productivity.
A water shoot originating from latent or adventitious buds on permanent arm structures including cordons, branches, and multi-year framework extensions in Vitis vinifera, characterized by emergence from woody arm surfaces between the trunk junction and the terminal arm positions, vigorous growth competing with fruiting positions along the arm, and representing response to localized damage, disease pressure, or management practices that severely reduce photosynthetic capacity in specific arm sections. Arm water shoots develop from buds that may have remained dormant for 3-10 years along arm surfaces, activated by removal of apical dominance through terminal damage, by vascular dysfunction from trunk diseases spreading into arm tissues, by excessive crop removal that creates carbohydrate surplus favoring vegetative growth, or by severe arm pruning that removes most fruiting positions and triggers compensatory bud break from latent positions. These shoots create specific challenges in cordon and arm management systems where maintaining linear framework geometry and consistent fruiting position spacing is critical for mechanization efficiency, cluster sun exposure, and spray penetration. However, arm water shoots also provide opportunities for local framework repair, allowing replacement of damaged arm sections without complete cordon or arm removal, and can be strategically retained and trained to restore fruiting positions lost to disease, freeze damage, or physical breakage.
Arm water shoots are distinguished from trunk water shoots by origin on permanent branch structures rather than the main trunk axis, and from normal lateral shoots by emergence from latent buds on woody arm tissue rather than from current-season buds on green shoots. Position along the arm influences management decisions, with shoots near the trunk junction often removed to maintain clean trunk-to-cordon transitions, while shoots in mid-arm or distal positions may be evaluated for framework repair potential. Arm water shoots in bilateral cordon systems require balanced removal decisions to maintain symmetry between cordon arms. Repeated emergence of arm water shoots from specific positions often indicates localized vascular dysfunction from trunk disease propagation into arm tissues.
Armwasserschoss
Kordonwassertrieb
Zweigtrieb
arm epicormic shoot
branch water shoot
cordon sucker
lateral arm sucker
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles pp. 148-163 Reynolds & Wardle (1993) "Impact of training system and vine spacing on vine performance and berry composition of Seyval blanc" American Journal of Enology and Viticulture 44=445-453 Martínez de Toda & Balda (2013) "Quantifying the effect of temperature on shoot and fruit growth of grapevine" Oeno One 47=229-241 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 343-365.
Arm Water Shoot
Baco 22A (Baco Blanc) bears an Armagnac_Production_Scion_Role as a unique permitted hybrid (Folle Blanche × Noah, a V. labrusca hybrid), contributing robust, earthy character and prune-like fruit to Armagnac blends—its inclusion reflects Armagnac's terroir-focused rather than purity-focused tradition, with Baco thriving in the sandy Bas-Armagnac soils where it produces the most distinctive spirits. Folle Blanche serves in an Armagnac_Production_Scion_Role producing the most delicate, floral Armagnacs prized by connoisseurs, particularly from the Ténarèze and Bas-Armagnac zones, though its disease susceptibility limits production to dedicated quality-focused estates. Ugni Blanc bears an Armagnac_Production_Scion_Role as the most planted variety (approximately 55% of Armagnac vineyards), providing reliable base wine for blending though considered less characterful than Baco or Folle Blanche.
A distillation scion role that inheres in a grafted scion cultivar by virtue of its authorization and suitability for Armagnac production within the delimited Armagnac appellation of Gascony, southwestern France, where base wine traditionally undergoes single continuous distillation in the distinctive Armagnacais column still (alambic armagnacais) followed by oak barrel aging. Armagnac production scions must be among the ten approved varieties under AOC regulations, with greater varietal diversity than Cognac reflecting Armagnac's artisanal traditions and terroir emphasis. The four principal varieties—Ugni Blanc, Baco 22A (Baco Blanc), Folle Blanche, and Colombard—each contribute distinct character, with Baco 22A unique among quality spirit appellations as a permitted hybrid variety (Folle Blanche × Noah). Armagnac varieties must produce high-acid, low-alcohol base wines suited to single distillation (versus Cognac's double distillation), which retains more congeners and creates a more robust, characterful spirit requiring extended aging to achieve smoothness.
Armagnac_Production_Scion_Role captures the distinctive varietal traditions of France's oldest brandy appellation, where greater variety diversity and the unique permission for hybrid Baco 22A distinguish Armagnac from the Ugni Blanc monoculture of Cognac. The single continuous distillation at lower proof (52-60% versus Cognac's 70% after second distillation) retains more flavor compounds, creating spirits with more pronounced character requiring longer aging (often 10-20+ years for peak quality). Armagnac's smaller scale and artisanal traditions have preserved practices (single-variety bottlings, vintage dating, estate production) increasingly rare in industrialized Cognac production. Climate and soil (boulbènes sands of Bas-Armagnac, clay-limestone of Ténarèze) strongly influence variety selection and spirit character within the appellation.
Armagnac grape
Armagnac variety
Armagnac-Rebsorte
Armagnac-Traube
Gascogne-Brennsorte
Gascon variety
Ténarèze variety
Bureau National Interprofessionnel de l'Armagnac (BNIA) (2020) Armagnac Production Regulations Faith N (2016) Cognac and Other Brandies, Infinite Ideas Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane pp.: 102-103, 423-424 Piggott JR et al. (1989) Distilled Beverage Flavour, Ellis Horwood
Armagnac Production Scion Role
3309 Couderc (V. riparia × V. rupestris) shows moderate Armillaria resistance with lesion extension rates of 3-6 mm/month and 15-year survival rates of 75-90% in Burgundy replant vineyards cleared from oak forest, substantially outperforming the highly susceptible 41B Millardet (V. vinifera × V. berlandieri) with survival rates of only 30-50%. 110 Richter (V. berlandieri × V. rupestris) demonstrates moderate resistance in Mediterranean climates, maintaining Tempranillo productivity on former oak woodland sites in Rioja for 20+ years. Riparia Gloire de Montpellier (V. riparia) shows moderate resistance but is limited to low-lime soils where Armillaria pressure often coincides with acidic, formerly forested land.
A fungal disease-resistance rootstock role that inheres in a grafted rootstock by virtue of its capacity to resist infection and colonization by Armillaria species, particularly A. mellea sensu stricto and related species in the A. mellea complex causing oak root fungus disease (also termed honey fungus or shoestring root rot). Armillaria spreads through soil via rhizomorphs (melanized hyphal aggregates 1-3 mm diameter) that contact and penetrate root bark, establishing mycelial fans beneath the bark that progressively girdle and kill roots and the root crown. Resistant rootstocks impede Armillaria colonization through thick, highly suberized bark with elevated suberin and lignin content, rapid wound periderm formation following rhizomorph contact, production of antifungal phenolics and terpenoids at infection fronts, and compartmentalization responses that wall off infected tissue. Resistance is quantified by lesion extension rate (resistant genotypes showing less than 5 mm/month versus 15-30 mm/month on susceptible hosts), crown colonization percentage, and long-term vine survival in Armillaria-infested sites.
Armillaria_Resistant_Rootstock_Role addresses one of the most intractable soil-borne diseases in viticulture, as Armillaria inoculum persists in soil for decades on decomposing woody debris and no curative treatments exist. Resistance is quantitative rather than qualitative no rootstock is immune, but resistant selections substantially extend productive vineyard lifespan in infested sites. V. rupestris and V. riparia parentage contributes resistance, while V. vinifera and V. berlandieri parentage generally increases susceptibility. Site selection (avoiding former oak woodland) remains the primary management strategy, with resistant rootstocks as a secondary defense.
Armillaria resistant rootstock
Armillaria-resistente Unterlage
Hallimasch-resistente Unterlage
gegen Eichenwurzelpilz resistente Unterlage
honey fungus resistant rootstock
oak root fungus resistant rootstock
shoestring root rot resistant rootstock
Aguín O et al. (2008) Armillaria species infesting vineyards in northwestern Spain, European Journal of Plant Pathology 120: 261-272 Baumgartner K, Rizzo DM (2006) Relative resistance of grapevine rootstocks to Armillaria root disease, American Journal of Enology and Viticulture 57: 408-414 Guillaumin JJ et al. (1991) Geographical distribution and ecology of Armillaria species in western Europe, European Journal of Forest Pathology 21: 143-153
Armillaria Resistant Rootstock Role
Aroma character profiling of New Zealand Sauvignon Blanc identifies the varietal fingerprint comprising tropical fruit character (passionfruit, guava from thiol compounds), citrus notes (grapefruit, lime), herbaceous elements (cut grass, boxwood from methoxypyrazines), and mineral undertones, with the specific balance among these character categories distinguishing Marlborough (tropical-dominant) from Hawke's Bay (more citrus and herbaceous) regional expressions of the same variety.
Aroma character quality is an aroma quality that represents the qualitative nature of olfactory perception distinguishing among different aromatic categories and descriptors, characterizing what the wine smells like in terms of recognizable scent attributes rather than how strong the overall aroma is. Within the BFO framework, this quality specifically addresses the qualitative dimension of olfactory perception, distinguishing aromatic identity from intensity while recognizing that character assessment involves matching perceived scents against learned reference categories stored in olfactory memory. The ontological significance lies in aroma character providing the descriptive vocabulary for communicating wine sensory properties, where the specific aromatic profile defines varietal typicity, regional expression, winemaking signature, and quality tier more distinctively than intensity measures that lack qualitative specificity. The perceptual basis for aroma character involves pattern recognition where complex volatile mixtures activate characteristic receptor combinations that the brain matches against previously experienced odor templates, with the resulting perception described using learned category labels (fruity, floral, spicy, earthy) that may or may not correspond to actual compositional similarity to the reference objects. The compositional basis for aroma character includes the specific volatile compounds present and their relative concentrations, with some compounds providing direct correspondence to their descriptors (linalool smelling floral because flowers contain linalool) while others evoke descriptors through similar receptor activation patterns without compositional relationship (rotundone smelling peppery despite structural dissimilarity to pepper compounds). The hierarchical organization of aroma descriptors proceeds from broad categories (fruity, floral, spicy, vegetal, earthy) through intermediate groupings (citrus, tropical, berry within fruity) to specific descriptors (lemon, passionfruit, blackberry), with wines typically described using multiple terms across categories reflecting their aromatic complexity. Understanding aroma character quality enables systematic sensory description, varietal and origin identification based on aromatic fingerprint, and winemaking optimization for desired aromatic profile.
Aroma_Character_Quality serves as the parent class for the hierarchy of aromatic descriptor categories, establishing the framework for qualitative aroma description. This quality complements Aroma_Intensity_Quality by addressing what rather than how much. The class encompasses the full range of wine aroma descriptors organized hierarchically from broad categories to specific terms, with Fruity_Aroma_Quality representing one major branch alongside floral, spicy, vegetal, earthy, and other character categories.
Aromacharakter
Duftprofil
Geruchsqualität
aroma type
aromatic profile
olfactory character
olfaktorischer Charakter
scent quality
Noble AC et al. (1987) Modification of a standardized system of wine aroma terminology. American Journal of Enology and Viticulture 38(2): 143-146 Ferreira V (2010) Volatile aroma compounds and wine sensory attributes. In: Reynolds AG (ed) Managing Wine Quality Volume 1, Woodhead Publishing, Cambridge, pp. 3-28
Aroma Character Quality
Aroma complexity for Grand Cru Burgundy Pinot Noir (Chambertin, 2015 vintage, 9 years) is assessed as "highly complex" — the nose reveals at least 15 identifiable descriptors spanning primary fruit (red cherry, raspberry, dried rose), secondary fermentative (subtle autolytic, background brioche), and tertiary maturation layers (forest floor, truffle, leather, iron-blood mineral, sandalwood, dried herbs, game), with new aromatic facets emerging over 30 minutes of glass evaluation and no single descriptor dominating the harmoniously integrated whole. Aroma complexity for commercial-tier Pinot Grigio from Veneto (2024 vintage) records "simple to moderate" — a clean but one-dimensional nose of pear and white almond with minimal layering and no developmental evolution during evaluation, consistent with the wine's intended role as an accessible, early-drinking style. Aroma complexity for Blaufränkisch Reserve from Leithaberg DAC (2021 vintage) evaluates at "moderate to high" — dark fruit core with integrated spice, mineral, and herbal secondary layers developing progressively in the glass, reflecting the quality and terroir specificity of the Leithaberg designation.
Aroma complexity is an aroma description entity that records the perceived multidimensionality, layering, and integrative coherence of a wine's olfactory profile, assessed through the identification of the number of distinct aroma descriptors, the diversity of aromatic origin categories represented (fruity, floral, herbal, spicy, earthy, mineral, woody, animal, chemical), the presence of multiple aromatic layers revealing themselves sequentially during evaluation, and the degree of harmonic integration among diverse aromatic elements into a unified sensory impression, evaluated on a structured scale from simple (one-dimensional, single-note) through moderate (multi-faceted but predictable) to complex (multi-layered, evolving, integrated yet diverse), and representing the qualitative dimension of aroma evaluation most closely correlated with expert quality judgment and wine price positioning.
Aroma complexity represents the single sensory parameter most consistently correlated with expert quality assessment and wine price, as numerous studies have demonstrated that trained tasters use complexity as the primary discriminator between quality tiers, with price-quality correlations strongest when complexity rather than intensity is the evaluated attribute. The perceptual basis of complexity involves both peripheral and central processing — the number and diversity of activated odorant receptors determines the peripheral input, while the temporal unfolding of multiple aromatic layers (as different compounds volatilize at different rates during evaluation) and the cognitive integration of diverse sensory inputs into a coherent gestalt determines the central quality judgment. The concept of "emergent complexity" — where the aromatic impression of a complex wine transcends the sum of its identifiable descriptor components — parallels the philosophical concept of emergence in complex systems and represents one of the most intellectually challenging aspects of wine sensory science. Complexity arises from multiple sources — cultivar-specific diversity of grape-derived aroma precursors, fermentation-derived aromatic contributions from diverse metabolic pathways, and aging-derived transformation products each add aromatic dimensions, with the greatest complexity achieved in wines where all three aromatic layers are simultaneously present and harmonically integrated. The temporal evolution of complexity during the tasting session provides additional quality information — complex wines reveal new aromatic facets as evaluation progresses (due to differential volatilization rates, temperature changes in the glass, and sensory adaptation-recovery cycles), while simple wines present their full aromatic repertoire immediately and then decline. Wine experts and novices differ most dramatically in their perception and articulation of complexity — trained tasters identify 3-5× more distinct descriptors in the same wine, suggesting that complexity perception involves learned attentional and categorization skills alongside innate olfactory acuity. The distinction between complexity (many different things working in harmony) and confusion (many different things competing for attention) represents the quality boundary — complex wines are intriguing and coherent, while confused wines are disjointed and tiring.
Aromakomplexität
Bukettvielfalt
Duftschichtung
aromatic complexity
aromatic multidimensionality
aromatische Vielschichtigkeit
bouquet complexity
nose complexity
olfactory complexity
olfaktorische Komplexität
Parr WV et al. (2002) Demystifying wine expertise: olfactory threshold, perceptual skill and semantic memory in expert and novice wine judges. Chemical Senses 27(8): 747-755 Lawless HT (1999) Descriptive analysis of complex odors: reality, model or illusion? Food Quality and Preference 10(4-5): 325-332 Ferreira V (2012) Revisiting psychophysical work on the quantitative and qualitative odour properties of simple odour mixtures. Flavour and Fragrance Journal 27(2): 124-140 Meillon S et al. (2010) Impact of partial alcohol reduction in wine on perceived complexity and temporality of sensations. Food Quality and Preference 21(7): 732-740
Aroma Complexity
Aroma complexity assessment comparing village-level and Grand Cru Burgundy reveals descriptor counts of 8 versus 15 distinct aromatic elements identified by a trained panel, with village wine showing fruit-dominated simplicity (cherry, earth, subtle oak) while Grand Cru displays layered complexity integrating red fruit, floral notes, forest floor, spice, mineral, and savory elements that emerge sequentially over 30 minutes of evaluation, with the complexity difference justifying the 10-fold price differential and indicating the Grand Cru's superior aging potential for further complexity development.
Aroma complexity quality is an aroma quality that represents the multiplicity, layering, and interrelationship of distinct aromatic elements perceived in wine, characterizing the richness and depth of olfactory experience that distinguishes multidimensional wines from simple, one-note examples. Within the BFO framework, this quality addresses the structural dimension of aromatic perception, distinguishing complexity from intensity while recognizing that complexity represents an emergent property arising from the presence of multiple distinct aroma compounds at perceptible concentrations that interact to create layered, evolving olfactory experience. The ontological significance lies in aroma complexity serving as a primary quality indicator across wine styles, where complex wines reward sustained attention by revealing successive aromatic layers and nuances while simple wines exhaust their aromatic interest quickly, with complexity correlating strongly with perceived quality, price, and aging potential across diverse wine categories. The perceptual basis for complexity involves the detection of multiple distinct odor objects that the olfactory system can individuate from the mixture, with complexity increasing as more distinguishable aromatic elements are present, to a point where excessive multiplicity may create confusion rather than appreciable complexity. The compositional determinants of complexity include the number of volatile compounds present above their individual detection thresholds, the diversity of compound classes represented (fruity, floral, spicy, earthy), and the absence of dominant single compounds that mask other elements, with balanced contribution from multiple sources creating the greatest complexity impression. The developmental aspects of complexity show simple, fruit-dominated young wines potentially gaining complexity through aging as tertiary aromas develop and integrate with primary and secondary elements, while excessively simple wines may lack the structural components (tannin, acidity, extract) required for complex evolution. The assessment of complexity employs both counting approaches (number of distinct descriptors identified) and holistic evaluation (overall impression of depth and layering), with experienced tasters showing greater ability to individuate components in complex mixtures. Understanding aroma complexity quality enables quality tier assessment based on aromatic depth, aging potential prediction from complexity indicators, and winemaking optimization for complexity through appropriate extraction, fermentation, and maturation decisions.
Aroma_Complexity_Quality addresses the structural depth dimension of aromatic perception, representing a meta-quality emerging from the interaction of multiple specific aroma characters. This quality is positively correlated with perceived overall quality and price across wine categories. Complexity assessment requires both analytical (descriptor enumeration) and holistic (overall depth impression) evaluation approaches for comprehensive characterization.
aroma layering
aromatic depth
aromatische Tiefe
olfaktorische Vielschichtigkeit
Lawless HT (1999) Descriptive analysis of complex odors: reality, model or illusion? Food Quality and Preference 10(4-5): 325-332 Ferreira V (2010) Volatile aroma compounds and wine sensory attributes. In: Reynolds AG (ed) Managing Wine Quality Volume 1, Woodhead Publishing, Cambridge, pp. 3-28
Aroma Complexity Quality
Aroma compound volatilization quality assessment in Marlborough Sauvignon Blanc compared volatile profiles in fruit from a heat event vintage (4 days exceeding 35°C during mid-ripening) versus a cool vintage to quantify aromatic losses from heat exposure. Methoxypyrazine analysis showed dramatic aroma compound volatilization quality impact with 3-isobutyl-2-methoxypyrazine (IBMP) at only 6 ng/L in heat-affected fruit versus 18 ng/L in cool-vintage fruit, representing 67% loss of this varietal character compound through a combination of accelerated degradation and volatilization during the heat event. Monoterpene analysis revealed 40% reduction in linalool and 35% reduction in geraniol, with a corresponding increase in thermally-rearranged forms (nerol, α-terpineol) indicating heat-induced isomerization affecting not only concentration but also aromatic character. Thiol precursor analysis showed only 25% reduction in glutathionylated precursors suggesting that heat impacts were primarily on existing volatiles rather than precursor pools for this compound class. Sensory evaluation of wines from the two vintages confirmed the analytical findings, with heat-affected wines showing significantly reduced intensity of green/herbaceous and tropical fruit characters, elevated perception of cooked/jammy notes, and overall reduced aromatic complexity reflecting the comprehensive aroma compound volatilization quality impact of the heat event.
Aroma compound volatilization quality is a heat impact on development quality that inheres in grape berries and represents the extent of heat-induced loss of volatile aromatic compounds through accelerated evaporation from berry surfaces, thermal degradation of heat-labile aroma molecules, and disruption of biosynthetic pathways producing varietal character compounds, with consequences for the aromatic intensity and typicity of wines produced from affected fruit. This quality captures the vulnerability of grape aroma compounds to thermal loss, recognizing that many key odorants including methoxypyrazines, terpenes, thiols, and esters have significant vapor pressures that increase with temperature, causing accelerated volatilization when berry temperatures are elevated, while other aromatic compounds undergo thermal degradation or isomerization that eliminates their sensory contribution. The ontological significance of aroma compound volatilization quality lies in the importance of aromatic character for wine identity and quality perception, with heat-induced losses potentially eliminating the varietal typicity that distinguishes premium wines and flattening the aromatic complexity that contributes to wine interest and value. The biological basis of aroma compound volatilization encompasses the physical chemistry of volatile compound behavior where elevated temperatures increase vapor pressure and diffusion rates accelerating loss from berry surfaces, combined with the thermal instability of certain aroma molecules that undergo degradation, oxidation, or rearrangement reactions at elevated temperatures, and the biosynthetic sensitivity of aroma compound production pathways that may be inhibited or redirected under heat stress. The specific aroma compounds most affected by heat exposure vary with grape variety reflecting differences in varietal aroma profiles, with methoxypyrazines in Sauvignon Blanc and Cabernet family varieties showing rapid heat-induced decline, monoterpenes in Muscat varieties undergoing both volatilization and thermal rearrangement, and thiols in Sauvignon Blanc and related varieties showing complex heat responses involving both precursor and free thiol dynamics. The rate of aroma compound loss depends on berry temperature, exposure duration, and berry surface characteristics including cuticle properties that affect vapor barrier function, with both acute high-temperature events and sustained moderate elevation causing significant aroma impacts through different kinetics.
Aroma_Compound_Volatilization_Quality is_a Heat_Impact_On_Development_Quality, representing the heat-induced loss of volatile aromatic compounds through evaporation, degradation, and biosynthetic disruption. This class addresses the aromatic quality dimension of heat impacts. Multiple mechanisms contribute including physical volatilization, thermal degradation, and isomerization. Compound-specific responses vary with vapor pressure and thermal stability. Methoxypyrazines show particular heat sensitivity (rapid loss). Wine sensory evaluation provides ultimate quality impact assessment. Distinguished from Anthocyanin_Degradation_Quality by focus on volatiles rather than pigments.
Aromaverlust durch Verdampfung
Verlust flüchtiger Verbindungen
aroma loss
aromatic degradation
aromatische Degradation
heat-induced aroma reduction
hitzeinduzierte Aromareduktion
volatile compound loss
Lacey MJ et al. (1991) Methoxypyrazines in Sauvignon blanc grapes and wines. American Journal of Enology and Viticulture 42:103-108 Marais J et al. (1999) Effect of microclimate, season and region on Sauvignon blanc grape composition and wine quality. South African Journal of Enology and Viticulture 20:19-30 Greer DH, Weedon MM (2013) The impact of high temperatures on Vitis vinifera cv. Semillon grapevine performance and berry ripening. Frontiers in Plant Science 4:491
Aroma Compound Volatilization Quality
Aroma degradation rate quality assessment for Muscat varieties in Rutherglen evaluated the stability of characteristic terpene profiles during extended hangtime for fortified wine production versus early-harvest table wine production. Muscat à Petits Grains demonstrated cultivar-typical rapid aroma degradation rate quality with linalool concentration declining 45% over two weeks of extended hangtime at ambient September temperatures, geraniol decreasing 38%, and the fresh floral character replaced by more developed, honeyed notes suitable for fortified production but representing loss of table wine aromatic typicity. The degradation rate proved strongly temperature-dependent, with fruit maintained at 15°C showing only 18% terpene loss over equivalent duration compared to 45% at ambient 25°C, informing recommendations for morning harvest and rapid cool transport when fresh aromatic character was the production objective. Contrasting assessment of Gewürztraminer showed intermediate aroma degradation rate quality with characteristic terpenes declining 22% over two weeks, permitting somewhat greater harvest timing flexibility while still requiring attention to aromatic preservation for quality production.
Aroma degradation rate quality is a post optimal decline quality that inheres in a grapevine cultivar or clone within a specific growing environment and represents the velocity at which varietal aromatic compounds and their precursors decline in concentration or transform to less desirable forms during extended hangtime beyond optimal ripeness, through volatilization losses, enzymatic transformation, oxidative degradation, and thermal decomposition pathways that progressively diminish the aromatic intensity and typicity achievable in wines produced from over-mature fruit. This quality directly determines the window during which optimal aromatic character can be captured and the urgency of harvest timing for aromatic cultivars where varietal expression depends critically on the preservation of specific volatile and glycosidically-bound compounds that may be irreversibly lost through extended hangtime. The ontological significance of aroma degradation rate quality lies in its particular importance for aromatic white varieties and certain red cultivars where varietal identity depends on characteristic aromatic profiles that prove more fragile and timing-sensitive than other quality parameters, creating harvest timing constraints that may not align with optimal sugar, acid, or phenolic development trajectories. The biological basis of aroma degradation rate quality encompasses multiple compound-specific pathways including the oxidative degradation of terpenes and norisoprenoids through reactive oxygen species and enzymatic catalysis, the thermal volatilization of highly volatile monoterpenes that accelerates under warm conditions, the continued glycosylation of free volatiles into bound forms that may reduce aromatic intensity depending on fermentation liberation efficiency, and the ongoing biosynthesis of certain compounds that may partially offset degradation losses. Cultivar characteristics influencing aroma degradation rate include the specific aromatic compound profile determining degradation susceptibility, the antioxidant capacity of fruit tissue providing protection against oxidative losses, the glycosylation propensity affecting the balance between free and bound volatiles, and the temperature sensitivity of degradation pathways creating climate-dependent stability differences. The assessment of aroma degradation rate quality requires compound-specific analysis tracking concentration changes through extended hangtime, with particular attention to the relationships between analytical measurements and sensory perception given the non-linear relationship between compound concentration and aromatic impact.
Aroma_Degradation_Rate_Quality is_a Post_Optimal_Decline_Quality, representing the velocity of aromatic compound loss or transformation beyond optimal ripeness. This class addresses the specific vulnerability of volatile compounds to extended hangtime. Particularly critical for aromatic varieties including Muscat, Riesling, Gewürztraminer, Viognier, and Sauvignon Blanc where varietal character depends on fragile aromatic compounds. The strong temperature dependency influences both vineyard (hangtime) and harvest operation (picking conditions) recommendations. Distinguished from Over_Ripening_Tolerance_Quality by the specific focus on aromatic parameters rather than overall quality maintenance.
Aromadegradationsrate
Aromastoffabbau-Geschwindigkeit
Aromenverlustrate
Duftstoff-Stabilitätsverlust
aromatic compound stability
aromatic decline trajectory
varietal aroma persistence
volatile degradation rate
Marais J (1983) Terpenes in the aroma of grapes and wines: a review. South African Journal of Enology and Viticulture 4:49-58 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine and Stabilization and Treatments, 2nd Edition. John Wiley and Sons, Chichester Kalua CM, Boss PK (2010) Comparison of major volatile compounds from Riesling and Cabernet Sauvignon grapes (Vitis vinifera L.) from fruitset to harvest. Australian Journal of Grape and Wine Research 16:337-348
Aroma Degradation Rate Quality
Aroma description for Riesling Spätlese from the Mosel (2020 vintage, 4 years post-harvest) documents an evolving nose integrating primary varietal character (lime blossom, white peach, green apple) with emerging tertiary complexity (beeswax, lanolin, incipient petrol/TDN), assessed at high intensity with pronounced complexity arising from the interplay of residual primary freshness and early maturation development — the aromatic trajectory characteristic of Mosel Riesling in the transition from youthful fruit to aged mineral-petrolic complexity. Aroma description for Blaufränkisch Reserve from Mittelburgenland DAC (2021 vintage) records a concentrated nose of dark cherry, blackberry compote, and damson plum (primary), subtle spice and cocoa from 14 months in Austrian oak (tertiary, early integration), and a faint violet-herbal lift, assessed at medium-high intensity with moderate-to-high complexity reflecting the varietal typicity of well-made Mittelburgenland Blaufränkisch.
Aroma description is a sensory description entity that documents the olfactory characterization of wine as perceived through orthonasal (sniffing from the glass) and retronasal (perception during and after swallowing) assessment, encompassing the identification of specific aroma descriptors, their classification by origin (primary/varietal, secondary/fermentative, tertiary/maturation), the evaluation of overall aroma intensity, and the assessment of aromatic complexity, recorded using standardized descriptor vocabularies and evaluated under controlled tasting conditions, providing the most information-rich and quality-discriminating component of wine sensory evaluation.
Aroma description constitutes the most analytically powerful component of wine sensory evaluation because the human olfactory system can discriminate among approximately 10,000 distinct odor qualities, detecting many aroma-active compounds at concentrations below the parts-per-billion level — a sensitivity that exceeds the detection limits of most analytical instruments for certain compound classes. The tripartite classification of wine aromas into primary (grape-derived varietal character), secondary (fermentation-derived), and tertiary (maturation-derived) provides the organizational framework for understanding the temporal evolution of wine aromatics from vineyard through cellar to bottle. The orthonasal-retronasal distinction is physiologically significant — orthonasal perception (sniffing) and retronasal perception (volatiles reaching the olfactory epithelium through the nasopharynx during tasting) activate partially overlapping but distinct neural processing pathways, with retronasal perception integrated with gustatory and tactile input to produce the composite "flavor" percept. The Wine Aroma Wheel developed by Ann Noble at UC Davis provides the most widely adopted hierarchical descriptor vocabulary, organizing aromas from broad categories (fruity, floral, spicy, vegetal, chemical, earthy, woody, caramelized, nutty) through intermediate groups to specific descriptors, though regional and cultural adaptations exist. The inter-individual variation in aroma perception — arising from genetic polymorphisms in odorant receptor genes (approximately 400 functional receptor genes with significant allelic variation), differences in mucosal enzyme activity, and variation in cognitive processing — means that panel-based assessment with multiple trained tasters is essential for reliable aroma characterization. The sequential assessment protocol — first evaluating the wine without swirling (assessing the most volatile compounds that escape immediately), then after gentle swirling (releasing additional volatiles through increased surface area and agitation), and finally after vigorous swirling or warming (assessing the least volatile and most tenacious aromas) — provides a structured approach to capturing the full aromatic profile.
Aromabeschreibung
Bouquetbeschreibung
Duftbeschreibung
Nasenbeschreibung
aroma assessment
aroma profile description
bouquet description
nose description
olfactory description
olfaktorische Beschreibung
Noble AC et al. (1987) Modification of a standardized system of wine aroma terminology. American Journal of Enology and Viticulture 38(2): 143-146 Jackson RS (2020) Wine Science: Principles and Applications, 5th Ed, Academic Press, pp.: 485-530 Shepherd GM (2006) Smell images and the flavour system in the human brain. Nature 444(7117): 316-321
Aroma Description
In Muscat à Petits Grains berries in Beaumes-de-Venise, Rhône Valley during BBCH 81-89, aroma development establishes total monoterpene content increasing from 1.2 to 8.5 mg/L over 45 days, with free volatiles rising from 0.3 to 1.8 mg/L and bound precursors from 0.9 to 6.7 mg/L, concurrent with norisoprenoid precursor accumulation from 25 to 145 µg/L, establishing the characteristic intense floral-grape Muscat aromatic profile under warm Mediterranean conditions averaging 26°C with substantial day-night temperature variation.
Aroma development constitutes the comprehensive ripening-phase process in Vitis vinifera berries wherein volatile aromatic compounds responsible for the olfactory perception of grape and wine character are synthesized, conjugated to precursor forms, and accumulated during the 35-50 day post-veraison maturation period, establishing the aromatic foundation that distinguishes grape varieties and expresses terroir influences through the characteristic scent profiles detected by human olfaction. This process encompasses the biosynthesis of diverse volatile compound classes including monoterpenoids contributing floral and citrus notes, sesquiterpenoids providing spicy and woody character, C13-norisoprenoids generating complex fruity and floral nuances, volatile thiols imparting tropical and passion fruit aromatics, esters contributing fruity sweetness, aldehydes and alcohols providing green and fatty notes, and methoxypyrazines generating herbaceous-vegetative character that collectively define the aromatic identity of individual grape varieties and vineyard sites. The distinction between aroma development and flavor development reflects sensory modality differences, with aroma specifically denoting volatile compounds perceived through olfactory receptors in the nasal epithelium either orthonasally during sniffing or retronasally during consumption, while flavor encompasses the broader integration of taste, aroma, and trigeminal sensations contributing to overall sensory experience. Cellular localization of aroma development processes shows tissue-specific patterns with monoterpenoid biosynthesis concentrated in exocarp cells where terpene synthase expression reaches highest levels, norisoprenoid generation associated with chromoplast-containing cells undergoing carotenoid degradation, and thiol precursor synthesis occurring predominantly in mesocarp parenchyma where cysteine conjugation machinery shows primary localization. The temporal progression of aroma development follows compound-class-specific trajectories with methoxypyrazines reaching maximum before veraison then declining, monoterpenes accumulating progressively throughout post-veraison ripening, and norisoprenoids showing late-ripening accumulation coincident with advanced carotenoid degradation during the final weeks before harvest. Cultivar genetics fundamentally determine aroma development potential through variety-specific expression of biosynthetic pathway genes, with aromatic varieties such as Muscat, Gewürztraminer, and Riesling demonstrating elevated monoterpenoid pathway activity, Sauvignon Blanc and related varieties showing distinctive thiol precursor synthesis capacity, and Cabernet family varieties expressing characteristic methoxypyrazine accumulation patterns. Environmental modulation of aroma development occurs through temperature effects on biosynthetic enzyme activities and compound volatility, light exposure influencing terpenoid and carotenoid pathway regulation, and water stress affecting secondary metabolism through stress-responsive pathway activation. The relationship between aroma development and wine quality reflects the fundamental importance of aromatic character for consumer perception and preference, with adequate aromatic intensity, complexity, and typicity representing primary quality criteria in premium wine grape production.
Aroma_Development is parent class under Ripening representing all volatile aromatic compound processes. Distinguishes from Flavor_Development which encompasses broader sensory attributes including taste. Contains subclasses Aroma_Precursor_Formation, Volatile_Compound_Synthesis, Terpene_Accumulation, and Aroma_Profile_Maturation. Duration 35-50 days post-veraison. Emphasizes olfactory perception specifically. Cultivar-specific expression patterns determine aromatic potential.
Aroma-Entwicklung
Duft-Entwicklung
aromatic compound development
aromatische Verbindungs-Entwicklung
olfactory compound synthesis
olfaktorische Verbindungs-Synthese
scent development
volatile aroma formation
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 285-310 Robinson AL et al (2014) Origins of grape and wine aroma. American Journal of Enology and Viticulture 65: 1-30 Lund ST and Bohlmann J (2006) The molecular basis for wine grape quality. Annals of Botany 98: 985-1010
Aroma Development
Aroma development potential assessment in young vintage Port compares current aromatic profile (intense primary dark fruit, grape spirit heat, simple spice) with potential indicators (glycoside-bound terpene reservoir 520 μg/L, pH 3.65, residual sugar 100 g/L providing reductive protection) to predict 30-50 year development trajectory toward complex tertiary bouquet (dried fruit, chocolate, coffee, rancio), with the high potential rating justifying €80 release price that will multiply as the wine approaches optimal drinking window, while parallel assessment of basic ruby Port (glycoside reserve 180 μg/L) indicates 3-5 year development window unsuitable for extended aging investment.
Aroma development potential quality is an aroma evolution quality that represents the capacity of wine to undergo positive aromatic transformation during aging, characterizing the likelihood that current aromatic profile will evolve toward greater complexity, integration, and quality rather than simple deterioration over time. Within the BFO framework, this quality specifically addresses the future trajectory dimension of aromatic assessment, distinguishing potential for evolution from current expression while recognizing that development potential represents latent quality not yet realized that requires appropriate storage conditions and time for manifestation. The ontological significance lies in aroma development potential providing the predictive framework for aging decisions, where high-potential wines justify extended cellaring investment by promising future aromatic rewards while low-potential wines should be consumed young before aromatic decline, making potential assessment essential for both consumer cellar management and fine wine commerce where futures pricing reflects anticipated development. The compositional indicators of development potential include glycosidically-bound terpene concentration (providing reserve for slow hydrolytic release), precursor compounds for aging bouquet development, phenolic structure (enabling slow oxidative transformation), and absence of volatile compounds prone to rapid deterioration, with these markers enabling analytical prediction of aromatic evolution alongside sensory assessment. The structural requirements for positive aromatic development include adequate acidity (preventing oxidative degradation), appropriate tannin level (enabling controlled micro-oxygenation), and overall balance that permits slow transformation without component collapse, with these non-aromatic factors critically influencing aromatic trajectory. The uncertainty inherent in development prediction creates challenges, as bottle variation, storage conditions, and individual wine idiosyncrasy mean that potential assessment provides probability guidance rather than guaranteed outcomes, with even expert predictions sometimes proving incorrect. Understanding aroma development potential quality enables informed cellaring decisions, futures market valuation based on anticipated evolution, and quality assessment incorporating future as well as current expression.
Aroma_Development_Potential_Quality specifically addresses the predictive dimension of aromatic assessment, representing latent quality not yet expressed. This quality requires both sensory evaluation of current state and analytical assessment of compositional indicators for future development. The class supports cellar management and fine wine valuation while acknowledging inherent prediction uncertainty due to storage variation and wine individuality.
Entwicklungspotential
Evolutionskapazität
Reifepotential
aging potential
aromatic longevity
aromatische Lagerfähigkeit
cellar-worthiness
evolution capacity
Ugliano M (2013) Oxygen contribution to wine aroma evolution during bottle aging. Journal of Agricultural and Food Chemistry 61(26): 6125-6136 Ferreira V et al. (2014) The aroma of wine. In: Chemistry and Biochemistry of Winemaking, Wine Stabilization, and Aging. IntechOpen, London, pp. 3-28
Aroma Development Potential Quality
Aroma evolution assessment in young Pauillac Cabernet Sauvignon identifies current primary-dominant stage (blackcurrant, cedar, graphite) with substantial evolution potential indicated by high precursor-bound terpene concentration (380 μg/L) and structural parameters (pH 3.55, tannin 78 IPT) supporting 15-25 year development trajectory toward tertiary bouquet (leather, tobacco, truffle, dried fruit), while parallel wine from lesser terroir shows similar current expression but lower precursor reserve (210 μg/L) suggesting shorter optimal aging window of 8-12 years before aromatic decline, demonstrating how evolution assessment informs cellar management beyond current quality evaluation.
Aroma evolution quality is an aroma quality that represents the capacity for and trajectory of aromatic change during wine development, characterizing both the current developmental stage of aromatic expression and the potential for future aromatic transformation through aging. Within the BFO framework, this quality addresses the temporal dimension of aromatic perception, distinguishing evolutionary potential and stage from static aromatic attributes while recognizing that wine aroma exists on a developmental continuum from primary grape-derived character through secondary fermentation contributions to tertiary aging bouquet. The ontological significance lies in aroma evolution providing the temporal framework for understanding wine's aromatic trajectory, where assessment of current evolutionary stage enables appropriate consumption timing decisions while evaluation of evolution potential informs cellar-worthiness and optimal aging duration predictions. The compositional basis for aroma evolution includes the reservoir of non-volatile precursors (glycosidically-bound terpenes, cysteine-bound thiols) available for release during aging, the reactive compounds (aldehydes, certain esters) that transform through oxidation or hydrolysis, and the structural compounds (tannins, acids) that provide the matrix stability enabling graceful evolution rather than simple deterioration. The developmental trajectory of wine aroma follows characteristic patterns, with primary varietal aromas dominant in youth, secondary fermentation aromas (yeast, malolactic) contributing in young wines, and tertiary bouquet (oxidative, reductive, hydrolytic transformation products) developing over years to decades in age-worthy wines, with the balance among these categories indicating evolutionary stage. The predictability of aroma evolution varies substantially, with certain varieties and styles showing consistent aging trajectories (Riesling developing petrol, Nebbiolo developing tar and rose) while others evolve unpredictably depending on storage conditions and bottle variation. Understanding aroma evolution quality enables appropriate consumption timing decisions, cellar management based on evolution assessment, and quality evaluation incorporating potential for development alongside current expression.
Aroma_Evolution_Quality serves as the parent class for Primary_Aroma_Expression_Quality and Aroma_Development_Potential_Quality, addressing the temporal dimension of aromatic assessment. This quality bridges current state evaluation with future trajectory prediction, supporting both consumption timing and aging decisions. The class recognizes wine aroma as a dynamic rather than static attribute, with developmental stage and potential representing essential quality dimensions.
Aromaentwicklung
Bukett-Evolution
Duftreifung
aroma aging potential
aromatic development
aromatisches Entwicklungspotential
bouquet development
nose evolution
Ribereau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed. John Wiley & Sons, Chichester, pp. 231-285 Ugliano M (2013) Oxygen contribution to wine aroma evolution during bottle aging. Journal of Agricultural and Food Chemistry 61(26): 6125-6136
Aroma Evolution Quality
Aroma intensity for Muscat Blanc à Petits Grains from Alsace (2024 vintage) is assessed as "pronounced" — the intensely aromatic terpene profile (total free monoterpenes >1500 µg/L) produces an immediately perceptible, room-filling aromatic impact detectable before the glass reaches the nose, characteristic of high-terpene Muscat at peak varietal expression. Aroma intensity for Chablis Premier Cru (2023 vintage, unoaked) records "medium" — the restrained aromatic profile of cool-climate Chardonnay without oak influence produces a nose of moderate projection requiring the standard swirling-and-sniffing protocol for full assessment, consistent with the Chablis stylistic emphasis on subtlety and mineral restraint over aromatic power. Aroma intensity for Grüner Veltliner Smaragd from the Wachau (2024 vintage) evaluates at "medium-plus to pronounced," reflecting the concentrated aromatic expression achievable from low-yielding, late-harvested Smaragd material.
Aroma intensity is an aroma description entity that records the perceived strength or concentration of the overall olfactory impression of a wine, assessed through orthonasal evaluation on a structured scale from faint (barely perceptible, requiring concentrated effort to detect) through light, medium-minus, medium, medium-plus, and pronounced (immediately powerful, filling the nasal cavity without effort from arm's-length distance), evaluated under standardized conditions (ISO tasting glass, appropriate serving temperature, sequential assessment before and after swirling), and providing the quantitative dimension of aroma evaluation that complements the qualitative descriptor identification by indicating the aromatic impact and "volume" of the wine.
Aroma intensity evaluation quantifies a perceptual dimension that, while seemingly simple, involves complex psychophysical processes governing the relationship between chemical stimulus concentration and perceived olfactory magnitude. The perceived intensity of a wine's aroma is determined by multiple factors beyond absolute volatile compound concentration — the vapor-liquid partition coefficients of individual compounds (governing how much of each compound enters the headspace for nasal sampling), the ethanol content of the wine (which modifies both vapor pressure and the solubility of hydrophobic aroma compounds), the serving temperature (higher temperature increases headspace volatile concentration), and the adaptation state of the taster's olfactory receptors (repeated exposure reduces perceived intensity through sensory adaptation). The non-linear psychophysical relationship between stimulus concentration and perceived intensity follows Stevens' power law with exponents typically between 0.2 and 0.6 for olfactory stimuli, meaning that doubling the concentration of an aroma compound increases perceived intensity by only 15-50% — a saturation effect that limits the perceptual distinction between moderately and very concentrated wines. Aroma intensity is not inherently correlated with quality — extremely intense wines may be considered overwhelming or unbalanced (excessive new oak, over-ripe fruit, volatile acidity), while moderately intense wines may achieve the highest quality through subtlety, nuance, and progressive aromatic revelation during the tasting session. The WSET Systematic Approach to Tasting employs a five-point intensity scale (light, medium-minus, medium, medium-plus, pronounced) that provides sufficient discrimination for professional assessment while avoiding the spurious precision of 100-point aroma intensity scales that exceed human discriminative capacity. The temporal dimension of intensity — the initial burst on first nosing versus the sustained aromatic persistence over minutes of glass evaluation — provides additional quality information, as high-quality wines typically maintain or increase aromatic interest over extended evaluation while lesser wines may present initial intensity that rapidly fades.
Aromaintensität
Duftstärke
Nasenintensität
aroma strength
aromatic concentration
aromatic power
aromatische Intensität
nose intensity
olfactory intensity
olfaktorische Stärke
Lawless HT, Heymann H (2010) Sensory Evaluation of Food: Principles and Practices, 2nd Ed, Springer, pp.: 1-73 WSET (2023) Wine and Spirit Education Trust Systematic Approach to Tasting, Level 3 and Diploma Methodology Stevens SS (1957) On the psychophysical law. Psychological Review 64(3): 153-181 Ferreira V (2012) Revisiting psychophysical work on the quantitative and qualitative odour properties of simple odour mixtures. Flavour and Fragrance Journal 27(2): 124-140
Aroma Intensity
Aroma intensity evaluation comparing young Riesling at 8°C versus 14°C serving temperature reveals intensity ratings of 5.2 versus 7.8 on a 10-point scale, with the 50% higher perceived intensity at warmer temperature corresponding to 3.2-fold higher headspace volatile concentration as measured by SPME-GC, demonstrating the critical role of serving temperature in realizing aromatic potential and explaining sommelier recommendations for serving aromatic whites slightly warmer than conventional guidelines to maximize varietal expression.
Aroma intensity quality is an aroma quality that represents the overall magnitude of olfactory sensation integrating the combined intensities of all volatile compounds perceived, characterizing whether the aromatic experience is powerful and pronounced or subtle and delicate as a general impression independent of specific aromatic descriptors. Within the BFO framework, this quality addresses the global intensity dimension of olfactory perception, distinguishing overall aromatic strength from the intensities of individual aroma notes and from aromatic complexity, while recognizing that total intensity provides information about volatile concentration and wine expressiveness not captured by qualitative aroma profiling. The ontological significance lies in aroma intensity serving as a primary indicator of wine aromatic potential and expression, where high intensity suggests aromatic variety, optimal grape maturity, careful handling preserving volatiles, and appropriate serving conditions, while low intensity may indicate neutral variety, underripe fruit, oxidative handling depleting volatiles, or serving temperature suppressing release. The perceptual integration of overall aroma intensity involves summation across odorant receptor responses with complex mixture interactions affecting the combined impression, creating a gestalt intensity perception where some compound combinations enhance (synergistic) while others diminish (suppressive) overall perceived intensity relative to component sum. The compositional determinants of aroma intensity include total volatile compound concentration, the proportion of compounds present above their individual detection thresholds, and the specific compounds present (some contributing disproportionately to overall intensity at low concentrations). The temporal dimension of aroma intensity includes the initial burst on opening, evolution in the glass as wine aerates and warms, and the sustained release during consumption, with appropriate intensity evolution indicating wine vitality and complexity. Understanding aroma intensity quality enables assessment of aromatic expression success, prediction of intensity from volatile analysis, and serving condition optimization (temperature, decanting) to achieve optimal intensity for appreciation.
Aroma_Intensity_Quality serves as the parent class for route-specific intensity qualities (Orthonasal, Retronasal), addressing the overall magnitude of aromatic perception. This quality represents the integrated perception of all volatile compounds, complementing specific aroma descriptor intensities that address individual aromatic notes. Intensity assessment requires standardized evaluation conditions as temperature, agitation, and glass shape significantly affect volatile release and perceived intensity.
Duftintensität
Geruchsintensität
aromatic strength
olfactory magnitude
scent power
Villamor RR, Ross CF (2013) Wine matrix compounds affect perception of wine aromas. Annual Review of Food Science and Technology 4: 1-20 Francis IL, Newton JL (2005) Determining wine aroma from compositional data. Australian Journal of Grape and Wine Research 11(2): 114-126
Aroma Intensity Quality
Aroma potential quality assessment in Muscat Blanc à Petits Grains guides harvest decisions for different wine styles, with glycosyl-glucose values of 8-12 μmol/g indicating optimal potential for dry aromatic wines where fermentation enzymatic release maximizes free terpene expression, while GG values above 15 μmol/g suggest overripe fruit with declining free terpene content despite high bound potential that may not be fully released without extended aging or enzyme addition targeting the 10-12 μmol/g GG range for dry Muscat balances immediate aromatic impact with aging development potential.
Aroma_Potential_Quality represents the chemical quality characterizing the total capacity of grape berries to contribute aromatic character to wine, integrating free volatile compounds, bound glycosidic precursors, and other aroma-relevant substrates into a comprehensive assessment of aromatic quality potential. Aroma potential recognizes that the concentration of immediately volatile compounds at harvest incompletely predicts wine aromatic intensity because substantial aroma contribution arises from bound precursors released during fermentation and aging. The concept encompasses multiple compound classes contributing to potential: terpene glycosides releasing floral monoterpenes norisoprenoid glycosides yielding fruity and exotic aromas thiol conjugates producing tropical notes and secondary aroma precursors supporting fermentation volatile formation. Total aroma potential can be estimated through hydrolysis methods that release all bound volatiles for measurement, or through glycosyl-glucose assays that quantify the sugar released from glycosidic precursors as a proxy for bound aroma content. The glycosyl-glucose (GG) assay developed by Abbott provides a rapid estimate of total glycosidic potential, with values ranging from 0.5-2 μmol/g berry for neutral varieties through 2-5 μmol/g for semi-aromatic varieties to 5-15 μmol/g for intensely aromatic Muscat types. Aroma potential increases during ripening as glycosidic precursor accumulation continues, but may plateau or decline in overripe fruit as degradation exceeds synthesis. The realization of aroma potential in finished wine depends on extraction efficiency, fermentation conditions (particularly yeast enzyme activities), and aging chemistry, with aroma potential providing an upper limit rather than a guaranteed outcome.
Aroma_Potential_Quality provides an integrative quality metric capturing total aromatic capacity rather than individual compound concentrations. The glycosyl-glucose (GG) assay enables practical assessment without the complexity of comprehensive volatile analysis, though it does not distinguish among compound classes contributing to the total. Aroma potential is a dispositional quality distinct from current aroma composition—it indicates capacity rather than current state. The conversion of potential to realized aroma depends on winemaking factors beyond grape composition, limiting the predictive precision of potential measurements for final wine character.
Aromapotential
Gesamtaroma-Vorläufergehalt
aromatic development potential
bound volatile capacity
gebundene Aromastoffkapazität
glycosidic aroma potential
glykosidisches Aromapotential
total aroma precursor content
Abbott NA et al. (1991) Rapid Determination of the Aromatic Potential of Riesling Grapes Using a Combination of Liquid Chromatography and GC-MS. American Journal of Enology and Viticulture 42(3): 219-223 Williams PJ et al. (1995) Glycosidic Flavor Precursors in Grapes and Wine. In: Charalambous G (ed) Food Flavors: Generation, Analysis and Process Influence, Elsevier pp.: 1265-1280
Aroma Potential Quality
In Fernão Pires berries in Bucelas, Portugal during BBCH 83-89, aroma precursor formation accumulates glycosidically-bound monoterpenes from 1.5 to 7.2 mg/L over 40 days, with bound linalool increasing from 0.4 to 2.1 mg/L, bound alpha-terpineol from 0.3 to 1.4 mg/L, and thiol precursor cysteine-3-mercaptohexanol from 45 to 185 µg/L, representing 88% of total monoterpenoids and 94% of total thiols in conjugated precursor forms at harvest under Atlantic-influenced conditions averaging 22°C.
Aroma precursor formation encompasses the biosynthetic processes in Vitis vinifera berries during ripening wherein volatile aromatic compounds are conjugated with sugar moieties, amino acids, or other carrier molecules to form stable, non-volatile precursor compounds that accumulate in berry tissues and require subsequent enzymatic or chemical transformation to release their aromatic potential during winemaking processes. This process involves multiple conjugation mechanisms including glycosyltransferase-mediated attachment of glucose, rhamnose, arabinose, or apiose residues to monoterpenols, sesquiterpenols, and norisoprenoids forming glycosidic precursors, and glutathione S-transferase catalyzed conjugation of cysteine or glutathione to reactive carbonyls generating thiol precursors that will release volatile thiols through yeast metabolism during fermentation. The quantitative significance of aroma precursor formation reflects the observation that glycosidically-bound forms typically constitute 85-95% of total monoterpenoid content and 70-90% of total norisoprenoid content in ripe berries, with free volatile concentrations representing only a small fraction of the aromatic potential stored in conjugated precursor pools awaiting release through appropriate processing interventions. The cellular localization of aroma precursor formation involves cytosolic glycosyltransferase activities converting free terpenols to their glucosides followed by vacuolar transport and sequestration establishing stable intracellular reservoirs, with compartmentalization protecting precursors from hydrolytic enzymes that would prematurely release volatile aglycones compromising aromatic quality and stability. Temporal dynamics of precursor formation show progressive accumulation throughout the post-veraison period, with glycosyltransferase gene expression and enzyme activities increasing 3-8 fold between veraison and harvest in aromatic varieties, enabling continued conjugation of newly synthesized volatile compounds throughout the ripening window. Cultivar genetics determine precursor formation capacity through variety-specific expression of glycosyltransferase gene families, with Muscat and related aromatic varieties showing elevated expression of monoterpenol glucosyltransferases compared to neutral varieties, and Sauvignon Blanc demonstrating distinctive glutathione S-transferase expression enabling thiol precursor accumulation. Environmental factors modulate precursor formation through temperature effects on conjugating enzyme activities and substrate availability, with moderate conditions of 20-28°C generally supporting optimal precursor accumulation while temperature extremes may limit biosynthetic efficiency or accelerate precursor degradation. The relationship between precursor pool size and wine aromatic quality depends on subsequent release efficiency during vinification, with larger precursor pools providing greater aromatic potential that can be realized through appropriate enzyme additions, yeast strain selection, or extended skin contact during winemaking.
Aroma_Precursor_Formation is parent class under Aroma_Development representing conjugation of volatiles to carrier molecules. Distinguishes from Volatile_Compound_Synthesis which generates free aromatics. Contains subclasses Glycosidic_Precursor_Synthesis, Bound_Aroma_Accumulation, and Precursor_Pool_Building. Typically 85-95% of monoterpenoids exist as bound precursors. Determines aromatic potential requiring winemaking release.
Aroma-Vorläufer-Bildung
aroma precursor biosynthesis
bound aroma compound formation
conjugated volatile formation
gebundene Aroma-Verbindungs-Bildung
glycosidic aroma precursor synthesis
glycosidische Aroma-Vorläufer-Synthese
konjugierte Volatil-Bildung
Gunata Z et al (1985) The aroma of grapes. Journal of Chromatography 331: 83-90 Tominaga T et al (1998) Identification of cysteinylated aroma precursors. Journal of Agricultural and Food Chemistry 46: 5215-5219 Williams PJ et al (1989) Glycosidic precursors of varietal grape and wine flavor. ACS Symposium Series 388: 35-48
Aroma Precursor Formation
Aroma precursor maturity assessment comparing Muscat clones reveals the varietal significance of precursor pools, with high-terpene clone showing free monoterpenes of 1.8 mg/L but bound terpene potential of 9.5 mg/L (5.3× ratio, substantial release potential during aging), low-terpene clone showing free terpenes of 0.9 mg/L and bound potential of 3.2 mg/L (3.6× ratio, lower total potential though similar relative reserve), with wines from the high-terpene clone showing characteristic aromatic evolution over 2 years of bottle aging as glycosidic reserves are progressively released, demonstrating how precursor maturity assessment predicts aromatic development trajectory beyond initial wine character.
Aroma precursor maturity quality is an aromatic maturity quality that represents the accumulation of glycosidically bound and cysteinylated aroma compounds that are not directly volatile but serve as reservoirs for aroma release during fermentation, aging, and consumption, characterizing the latent aromatic potential that may exceed free volatile content and determines the aromatic complexity achievable through winemaking interventions that release these bound forms. Within the BFO framework, this quality specifically addresses the precursor dimension of aromatic maturity, distinguishing bound aroma potential from free volatile content while recognizing that precursor pools often exceed free volatile pools by 5-10 fold in grapes, representing substantial latent aromatic potential that enzyme activity during fermentation, acid hydrolysis during aging, and enzymatic release during consumption can progressively liberate. The ontological significance lies in aroma precursor maturity determining the aromatic development potential of wines beyond their immediate sensory character, where glycosidic precursors of terpenes (bound linalool, geraniol) provide reservoirs for floral character release, cysteinylated precursors of volatile thiols (S-cysteine conjugates) provide tropical fruit potential realized through yeast metabolism, and bound norisoprenoids provide complexity that emerges during aging, together establishing the aromatic evolution trajectory of wines from fermentation through long-term aging. The precursor classes with distinct maturation patterns include monoterpenyl glycosides (accumulating primarily pre-véraison, stable through ripening), norisoprenoid glycosides (increasing during ripening as carotenoid precursors are cleaved and glycosylated), cysteinylated thiols (accumulating during ripening, particularly in later maturity), and hydroxycinnamic acid esters (accumulating during ripening, volatile phenol precursors), with each class contributing to different aspects of wine aroma through different release mechanisms. The release mechanisms affecting precursor utilization include enzymatic hydrolysis during fermentation (yeast glycosidases releasing monoterpenes, yeast β-lyases releasing thiols from cysteine conjugates), acid-catalyzed hydrolysis during aging (slow release of norisoprenoids contributing to bottle bouquet), and residual enzyme activity (continued slow release from remaining precursor pools), with winemaking decisions affecting which mechanisms operate and thus which aspects of precursor potential are expressed. The analytical assessment of precursor maturity involves either measurement of glycosidic/bound forms directly or measurement of glycosyl-glucose (GG) as a proxy for total glycosidic potential, with GG increasing during ripening and providing an index of aromatic development potential that complements free volatile analysis. Understanding aroma precursor maturity quality enables assessment of latent aromatic potential beyond immediate volatile content, harvest timing that optimizes precursor accumulation for intended wine style, and winemaking planning that targets appropriate precursor release mechanisms.
Aroma_Precursor_Maturity_Quality addresses the glycosidic and conjugated reservoirs of latent aroma potential. This quality recognizes that bound forms often exceed free volatiles and determine aromatic evolution during aging. The class supports aromatic potential assessment through precursor characterization beyond free volatile analysis.
Aromavorläuferreife
bound aroma potential
gebundenes Aromapotential
glycosidic aroma reserve
glykosidische Aromareserve
latent volatiles
latente Aromastoffe
precursor accumulation
Williams PJ et al. (1982) Studies on volatile flavor compounds in Muscat grapes. Australian Journal of Grape and Wine Research 48(1): 122-139 Tominaga T et al. (1998) A powerful aromatic volatile thiol, 2-furanmethanethiol, exhibiting roast coffee aroma in wines made from several Vitis vinifera grape varieties. Journal of Agricultural and Food Chemistry 46(3): 1044-1048
Aroma Precursor Maturity Quality
Aroma profile data for Sauvignon Blanc clone 242 versus clone 316 at Marlborough, New Zealand documents 3-mercaptohexan-1-ol (3MH) concentrations of 1200-1800 ng/L versus 800-1100 ng/L in finished wines, with corresponding 3-mercaptohexyl acetate (3MHA) levels of 180-260 ng/L versus 90-150 ng/L, confirming clone 242's superior volatile thiol release potential and its characteristically intense passionfruit and grapefruit aroma intensity. Aroma profile data for Muscat Blanc à Petits Grains clones at Geilweilerhof differentiates clone Mu-14 (free linalool 850 µg/L, geraniol 420 µg/L, bound glycosidic terpenes 3200 µg/L) from clone Mu-8 (free linalool 520 µg/L, geraniol 280 µg/L, bound glycosides 1950 µg/L), quantifying the 40-60% terpene concentration advantage that positions Mu-14 as the preferred clone for aromatic Muskateller production in Austrian and German programs.
Aroma profile data is a quality performance data entity that records the concentration, composition, and sensory characterization of volatile and non-volatile aromatic compounds in the berries, must, and wine of a clonal line, encompassing free and bound terpenes, norisoprenoids, thiols, methoxypyrazines, esters, higher alcohols, and their precursors, measured through gas chromatography-mass spectrometry (GC-MS), gas chromatography-olfactometry (GC-O), and trained sensory panel descriptive analysis, characterizing the aromatic identity and typicity potential that distinguishes clonal lines within a cultivar.
Aroma profile data represents perhaps the most challenging dimension of clone quality evaluation, as aromatic compound concentrations span several orders of magnitude (from µg/L for terpenes to ng/L for thiols and methoxypyrazines), require sophisticated analytical instrumentation, and interact in complex perceptual combinations where sub-threshold compounds contribute synergistically to overall aroma. The distinction between varietal aroma compounds (genetically determined, cultivar-specific) and fermentation-derived aroma compounds (influenced by yeast strain and winemaking protocol) is critical for clone evaluation — only varietal aroma differences can be attributed to clonal genetics, while fermentation aromas must be controlled through standardized micro-vinification. Key varietal aroma compound classes include monoterpenes (Muscat, Gewürztraminer, Riesling), C13-norisoprenoids such as β-damascenone and TDN (Riesling), volatile thiols (Sauvignon Blanc, Colombard), rotundone (Shiraz pepper character), and methoxypyrazines (Cabernet Sauvignon, Carménère green bell pepper character). Bound glycosidic aroma precursors in berries provide an additional quality metric, as they represent the latent aromatic potential released during fermentation, aging, and acid hydrolysis. Clone-by-environment interactions for aromatic compounds are substantial — methoxypyrazine degradation is strongly light-dependent, terpene accumulation responds to temperature and UV exposure, and thiol precursor synthesis is influenced by nitrogen status — necessitating multi-site evaluation for reliable aromatic characterization.
Aromadaten
Aromaprofildaten
Aromastoffprofil
aroma data
aromatic composition data
flavor profile data
flüchtige Verbindungen Daten
sensorisches Aromaprofil
sensory aroma record
volatile compound profile
Ebeler SE, Thorngate JH (2009) Wine chemistry and flavor: looking into the crystal glass. Journal of Agricultural and Food Chemistry 57(18): 8098-8108 Lund ST, Bohlmann J (2006) The molecular basis for wine grape quality — a volatile subject. Science 311(5762): 804-805 Roland A et al. (2011) Pre- and post-bottling evolution of volatile thiols in Sauvignon Blanc wines: key role of temperature. Food Chemistry 128(4): 1022-1035 Dunlevy JD et al. (2009) A methyltransferase essential for the methoxypyrazine-derived flavour of wine. The Plant Journal 59(3): 404-412
Aroma Profile Data
In Petit Manseng berries in Jurançon, Pyrénées during BBCH 87-89, aroma profile maturation refines the aromatic character over the final 18 days, with monoterpene ratios shifting as linalool-to-alpha-terpineol ratio decreases from 3.2 to 2.1 through acid-catalyzed rearrangement, C6 aldehydes declining from 55 to 12 µg/L eliminating residual green character, and trained panel ratings for tropical-citrus character increasing from 5.8 to 8.2 while green-herbaceous scores decline from 4.2 to 1.8 on 10-point scales under late-harvest conditions with cool temperatures of 15°C.
Aroma profile maturation constitutes the terminal phase of aromatic development in Vitis vinifera berries wherein accumulated volatile compounds and precursors undergo final qualitative refinements and the overall olfactory character achieves the balanced, harmonious expression associated with optimal ripeness during the final 10-21 days before harvest, representing the culmination of aromatic biosynthetic processes and establishing the definitive varietal and terroir-expressive scent signature that will carry through vinification into finished wine character. This process involves subtle transformations in aromatic composition including shifts in monoterpene ratios through interconversion reactions such as acid-catalyzed linalool rearrangement to alpha-terpineol and geraniol-nerol isomerization, evolution of norisoprenoid profiles through continued carotenoid degradation generating late-ripening aromatic contributions, and progressive reduction of green-herbaceous character through declining C6 aldehyde content establishing the ripe-fruit-dominated aromatic balance characteristic of optimally mature grapes. The qualitative aspects of aroma profile maturation extend beyond simple concentration changes to encompass integration of disparate aromatic elements into cohesive olfactory impressions, development of complexity through accumulation of minor aromatic components that contribute nuance and depth, and achievement of varietal typicity enabling clear cultivar identification through characteristic scent signatures that define grape variety identity. Sensory evaluation of aroma profile maturation employs holistic assessment methods including overall aromatic quality ratings, harmony and balance scores integrating multiple scent attributes, and typicity judgments comparing observed profiles to variety benchmarks, complementing analytical measurements of individual compound concentrations that may not fully capture the perceptual integration underlying aromatic quality assessment. Environmental conditions during the final ripening phase critically influence aroma profile maturation, with moderate temperatures of 18-24°C favoring continued aromatic refinement and preservation of accumulated volatiles, while excessive heat may accelerate degradation of heat-labile compounds or shift aromatic profiles toward overripe character before optimal balance is achieved. The timing of aroma profile maturation relative to sugar accumulation and acid reduction varies among cultivars, with aromatic varieties often achieving optimal scent character coincident with traditional ripeness indicators while some cultivars may require extended hang-time or alternatively reach aromatic optimum before sugar targets are achieved, necessitating harvest timing decisions that balance multiple quality parameters.
Aroma_Profile_Maturation is parent class under Aroma_Development representing terminal qualitative refinement phase. Distinguishes from Terpene_Accumulation which emphasizes quantitative increase. Contains subclasses Mature_Aroma_Expression, Cultivar_Typical_Aroma, and Optimal_Aroma_Development. Duration 10-21 days before harvest. Emphasizes balance, harmony, and typicity. Primary harvest timing criterion for aromatic quality.
Aroma-Balance-Erreichung
Aromaprofil-Reifung
Duftprofil-Vollendung
aroma balance achievement
aromatic profile completion
olfactory character finalization
olfaktorische Charakter-Finalisierung
scent profile maturation
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 305-315 Robinson AL et al (2014) Origins of grape and wine aroma. American Journal of Enology and Viticulture 65: 1-30 Coombe BG (1992) Research on development and ripening of the grape berry. American Journal of Enology and Viticulture 43: 101-110
Aroma Profile Maturation
Aroma quality assessment of Marlborough Sauvignon Blanc identifies the distinctive regional aromatic fingerprint comprising primary varietal aromas (passionfruit from 3MH thiol at 1200 ng/L, grapefruit from 3MHA at 180 ng/L, boxwood from 4MMP at 35 ng/L) combined with secondary fermentation notes (subtle yeast autolysis) without tertiary oak-derived aromas, creating the immediately recognizable aromatic profile that defines this origin's market identity and commands premium pricing based on aromatic typicity alone.
Aroma quality is a sensory quality that represents the olfactory perception arising from volatile compound stimulation of odorant receptors in the nasal epithelium, encompassing the complex bouquet of scent attributes that characterizes grape variety, terroir expression, fermentation character, and aging development in wines. Within the BFO framework, this quality addresses the chemosensory perception mediated by approximately 400 functional odorant receptor types in humans, distinguishing olfaction from gustation (taste) while recognizing the profound integration of these modalities in the unified flavor perception that dominates wine appreciation. The ontological significance lies in aroma providing the primary complexity dimension in wine evaluation, where the hundreds of volatile compounds present at suprathreshold concentrations create the aromatic fingerprint that identifies variety, reveals winemaking technique, and evolves through aging to define the wine's sensory identity more distinctively than any other attribute. The physiological basis for aroma perception involves volatile molecules reaching olfactory receptors through two routes, orthonasal (external sniffing) and retronasal (internal perception during consumption), with these pathways providing different but complementary aromatic information due to differential compound release and neural processing. The compositional basis for grape and wine aroma includes varietal compounds (terpenes, thiols, methoxypyrazines defining variety character), fermentation-derived volatiles (esters, higher alcohols, volatile acids from yeast metabolism), and aging-derived compounds (oak lactones, volatile phenols, oxidation products developing during maturation). The hierarchical classification of wine aroma distinguishes primary aromas (grape-derived varietal character), secondary aromas (fermentation-derived), and tertiary aromas (aging-derived bouquet), with the balance among these categories reflecting wine style and developmental stage. The complexity of aroma perception, involving mixture interactions where compounds may suppress, enhance, or qualitatively transform each other's perception, creates the rich sensory tapestry that makes wine aroma evaluation both challenging and rewarding. Understanding aroma quality enables comprehensive sensory assessment, varietal and origin authentication based on aromatic fingerprint, and winemaking optimization for desired aromatic profile.
Aroma_Quality serves as the root class for the olfactory perception hierarchy, establishing the framework for volatile compound-mediated sensory experience distinct from taste. This quality encompasses both orthonasal (sniffing) and retronasal (in-mouth) perception routes that provide complementary aromatic information. The class serves as parent for intensity, character, and complexity subclasses addressing different dimensions of aromatic perception.
Aromaqualität
Bukett
Duftqualität
bouquet
nose
olfactory quality
olfaktorische Qualität
Polášková P et al. (2008) Wine flavor: chemistry in a glass. Chemical Society Reviews 37(11): 2478-2489 Ferreira V (2010) Volatile aroma compounds and wine sensory attributes. In: Reynolds AG (ed) Managing Wine Quality Volume 1, Woodhead Publishing, Cambridge, pp. 3-28
Aroma Quality
Aroma stability assessment comparing early and optimal harvest Viognier reveals the maturity effect on aromatic persistence, with early harvest showing high initial free terpenes (2.4 mg/L) but low precursor reserves (4.1 mg/L bound), producing wines with intense young aroma that declined 45% over 12 months storage, while optimal harvest shows lower initial free terpenes (1.8 mg/L) but higher precursor reserves (8.2 mg/L bound), producing wines with moderate initial intensity but only 18% decline over 12 months as precursor hydrolysis partially compensated free terpene losses, demonstrating how aroma stability assessment enables harvest timing that optimizes aromatic quality at drinking rather than at fermentation completion.
Aroma stability quality is an aromatic maturity quality that represents the potential for aromatic compounds to persist through winemaking, storage, and aging rather than being lost through volatilization, oxidation, or chemical transformation, characterizing the durability of aromatic character that determines whether the aromatic quality at harvest translates into lasting wine quality or diminishes rapidly post-fermentation. Within the BFO framework, this quality specifically addresses the persistence dimension of aromatic maturity, distinguishing the stability potential of accumulated aromatics from their immediate concentration while recognizing that the same total aromatic content may yield wines with very different aromatic longevity depending on the proportion of stable versus labile compounds and the presence of protective or destabilizing factors. The ontological significance lies in aroma stability determining the sensory quality trajectory of wines from fermentation through aging, where stable aromatic systems maintain or develop complexity over time while unstable systems lose aromatic intensity and may develop off-aromas, with stability depending on the chemical nature of accumulated compounds, the presence of antioxidant phenolics, and the redox environment established during winemaking and storage. The factors contributing to aroma stability include the proportion of aromatics in bound versus free forms (bound forms representing stable reservoirs for progressive release), the degree of anthocyanin-derived pigment formation (polymeric pigments being more stable than monomeric anthocyanins), the presence of co-pigmentation and tannic structure (providing antioxidant protection), and the accumulation of sulfur-containing compounds (thiols being particularly oxidation-sensitive and requiring reductive handling), with these factors varying with grape maturity and affected by harvest timing decisions. The compound-specific stability patterns show substantial variation, with some aromatics highly stable (polymeric pigment-derived compounds persisting for decades), others moderately stable (terpenes showing gradual decline and transformation), and others highly labile (many thiols oxidizing within months without protective winemaking), with the stability profile of accumulated aromatics affecting the achievable wine style and aging potential. The maturity effects on aroma stability show complex patterns, as later harvest may enhance stability through increased phenolic protection and greater precursor pools (providing reservoirs for continued release) while also increasing the proportion of certain labile compounds (potentially reducing immediate aromatic persistence), requiring consideration of stability alongside immediate aromatic content in harvest decisions. Understanding aroma stability quality enables prediction of aromatic longevity, harvest timing that balances immediate aromatic intensity against persistence potential, and winemaking planning that protects or enhances the stability of accumulated aromatics.
Aroma_Stability_Quality addresses the persistence potential of aromatic compounds through winemaking and aging. This quality recognizes that wine quality depends on aromatic durability not just initial intensity. The class supports wine longevity planning through aroma stability characterization affecting harvest and winemaking decisions.
Aromalanglebigkeit
Aromastabilität
Duftstabilität
aroma longevity
aromatic durability
aromatic persistence
aromatische Persistenz
volatile stability
Ferreira V et al. (2016) The chemistry of wine aging. In Moreno-Arribas MV, Polo MC (eds) Wine Chemistry and Biochemistry. Springer, New York, pp. 301-340 Silva Ferreira AC et al. (2002) Identification of key odorants related to the typical aroma of oxidation-spoiled white wines. Journal of Agricultural and Food Chemistry 50(11): 2585-2591
Aroma Stability Quality
Aroma typicity evaluation of wines submitted for Marlborough Sauvignon Blanc Geographical Indication approval applies regional typicity criteria requiring passionfruit intensity above 5.0 (10-point scale), herbaceous notes present but not dominant, and absence of oak-derived aromas, with wines showing passionfruit below threshold or excessive tropical character without herbaceous balance rejected as atypical for the regional style, demonstrating how typicity assessment enforces category boundaries and commercial authenticity while potentially limiting stylistic diversity within the protected designation.
Aroma typicity quality is an aroma quality that represents the degree to which a wine's aromatic profile conforms to expected or recognized patterns for its designated category, whether defined by grape variety, geographical origin, or production style, characterizing the authenticity and representativeness of aromatic expression relative to learned quality templates. Within the BFO framework, this quality addresses the conformity dimension of aromatic perception, distinguishing typicity from intensity or complexity while recognizing that typicity represents a relational quality comparing actual aromatic profile to an idealized or consensual expectation for the wine's declared identity. The ontological significance lies in aroma typicity providing the evaluative framework for authenticity assessment, where typical wines confirm expectations for their category (varietal, regional, stylistic) while atypical wines either innovate beyond conventions or fail to achieve expected character, with typicity serving as both a quality criterion (typical = authentic = valuable) and a constraint on innovation (typicity requirements may penalize creative winemaking). The perceptual basis for typicity judgment involves matching perceived aromatic profile against learned prototypes stored in memory, with typicity assessment requiring familiarity with the relevant category's aromatic signature and the ability to judge similarity between actual perception and mental template. The compositional determinants of typicity include the presence and appropriate intensity of variety-defining impact compounds (methoxypyrazines in Sauvignon Blanc, rotundone in Syrah, terpenes in Muscat), the balance among aromatic categories expected for the wine type, and the absence of atypical elements that would indicate origin or variety confusion. The regulatory and commercial implications of typicity are substantial, with appellation systems enforcing typicity requirements through sensory evaluation panels that reject atypical wines from protected designation use, creating economic pressure for conformity that may limit diversity within traditional categories. Understanding aroma typicity quality enables authenticity assessment for variety and origin claims, quality evaluation within category-specific frameworks, and appreciation of the tension between typicity preservation and stylistic innovation in wine production.
Aroma_Typicity_Quality serves as the parent class for Varietal_Typicity_Quality and Terroir_Expression_Quality, addressing the conformity dimension of aromatic assessment relative to category expectations. This quality is inherently relational, requiring specification of the reference category against which typicity is judged. The class raises ontological questions about the nature of category prototypes and the tension between authenticity preservation and innovation permission.
Aromatypizität
Herkunftstypizität
Sortentypizität
aroma representativeness
aromatic authenticity
aromatische Authentizität
nose typicality
olfactory conformity
Maitre I et al. (2010) Is typicality an evolving concept? Journal of Sensory Studies 25(s1): 103-129 Parr WV et al. (2007) The distinctiveness of Sauvignon Blanc wine. Food Quality and Preference 18(6): 849-861
Aroma Typicity Quality
Aromatic analysis data for Gewürztraminer from Alsace (2024 vintage) integrates free terpene profile (linalool 620 µg/L, geraniol 380 µg/L, citronellol 185 µg/L, α-terpineol 95 µg/L), bound glycosidic precursor pool (total bound terpenes 2800 µg/L as linalool equivalents), norisoprenoid content (β-damascenone 4.2 µg/L, β-ionone 0.8 µg/L), and cis-rose oxide (28 µg/L, above the 0.2 µg/L sensory threshold) — collectively defining the cultivar's characteristically intense, floral-lychee aromatic signature. Aromatic analysis data for Cabernet Franc from Chinon, Loire Valley (2024 vintage) documents methoxypyrazine concentration (IBMP 18 ng/L, near the 15 ng/L sensory threshold), norisoprenoid levels (β-damascenone 5.8 µg/L), and volatile thiol screening (3MH below detection limit), characterizing the moderate herbaceous-fruity aromatic balance typical of cool-vintage Loire Cabernet Franc.
Aromatic analysis data is a quality analysis data entity that documents the concentration, composition, and sensory relevance of volatile and semi-volatile aroma compounds and their non-volatile precursors in grape berries, must, and wine, determined through gas chromatography-mass spectrometry (GC-MS), gas chromatography-olfactometry (GC-O), liquid chromatography-mass spectrometry (LC-MS/MS for precursors), and headspace analysis techniques, encompassing terpenes, norisoprenoids, methoxypyrazines, volatile thiols and their precursors, and the broader volatile compound profile, providing the chemical characterization of aromatic identity and typicity that distinguishes cultivars, sites, and vintages at the molecular level.
Aromatic analysis data represents the most analytically demanding and instrumentally intensive component of quality analysis, requiring specialized extraction techniques (solid-phase microextraction SPME, stir bar sorptive extraction SBSE, liquid-liquid extraction, headspace sampling), high-resolution chromatographic separation, and sensitive detection systems capable of quantifying compounds present at concentrations spanning six orders of magnitude — from terpenes at hundreds of µg/L to methoxypyrazines and volatile thiols at single-digit ng/L. The concept of odor activity value (OAV = concentration / sensory threshold) provides the critical bridge between analytical chemistry and sensory perception, as many abundant volatile compounds fall below their perception thresholds while trace-level compounds may dominate the aroma profile. Aromatic analysis data encompasses both "varietal aroma" compounds (genetically determined, present in the grape, defining cultivar typicity) and their precursor pools (glycosidically bound terpenes, cysteinylated and glutathionylated thiol precursors, carotenoid norisoprenoid precursors) that represent latent aromatic potential released during fermentation, aging, and acid-catalyzed hydrolysis. The vineyard origin of aromatic compounds links this analytical domain directly to terroir characterization — climate (temperature, UV exposure), soil (nitrogen status, water availability), and canopy management (fruit zone light environment) all modulate aromatic compound biosynthesis and degradation rates, creating site-specific and vintage-specific aromatic signatures. The rapid advancement of comprehensive two-dimensional gas chromatography (GC×GC) coupled with time-of-flight mass spectrometry (TOF-MS) is expanding the detectable volatile inventory from approximately 200-300 compounds by conventional GC-MS to over 1000 compounds, enabling increasingly detailed aromatic fingerprinting.
Aromaanalysedaten
Aromastoffanalytik
Aromazusammensetzungsdaten
Duftstoffanalysedaten
aroma analysis data
aroma profiling data
flavor chemistry data
flüchtige Verbindungen Analyse
volatile compound analysis
Ebeler SE, Thorngate JH (2009) Wine chemistry and flavor: looking into the crystal glass. Journal of Agricultural and Food Chemistry 57(18): 8098-8108 Lund ST, Bohlmann J (2006) The molecular basis for wine grape quality — a volatile subject. Science 311(5762): 804-805 Robinson AL et al. (2014) Origins of grape and wine aroma. Part 1. Chemical components and viticultural impacts. American Journal of Enology and Viticulture 65(1): 1-24
Aromatic Analysis Data
Aromatic compound profiles vary dramatically across V. vinifera cultivars, defining varietal identity through characteristic volatile signatures. Muscat cultivars accumulate 2-6 mg/L free monoterpenes (primarily linalool and geraniol), while non-aromatic cultivars contain <0.5 mg/L. Sauvignon Blanc accumulates volatile thiol precursors (cysteine and glutathione conjugates) reaching 500-2000 µg/L, releasing 3-mercaptohexanol and 4-mercapto-4-methylpentan-2-one during fermentation. Methoxypyrazine content distinguishes Cabernet Sauvignon and Sauvignon Blanc (5-40 ng/L IBMP) from non-herbaceous cultivars (<2 ng/L). The glycosidically bound aroma pool exceeds free volatiles by 3-10 fold in most cultivars, representing a reservoir released during winemaking and aging.
A quality potential disposition that is the inherited capacity of a Vitis vinifera plant to synthesize, accumulate, and store volatile and non-volatile aroma precursor compounds in grape berries, encompassing terpenoids, norisoprenoids, thiols, methoxypyrazines, and other metabolites that contribute to varietal aroma character in grapes and wines.
Aromatic_Compound_Disposition serves as the parent class for specific aroma compound categories. Varietal aroma is among the most valued quality attributes, with aromatic cultivars commanding premium prices. The trait shows strong genetic control (h² = 0.70-0.90 for total monoterpenes) but also significant environmental modulation by temperature, light, and water status. Aroma compounds exist in both free (volatile) and bound (glycosylated, non-volatile) forms, with bound precursors released through acid hydrolysis, enzyme action, or fermentation.
Aromastoff-Disposition
Aromaverbindungen
Duftstoff-Akkumulation
aroma synthesis capacity
flavor compound disposition
volatile precursor trait
Lund ST, Bohlmann J (2006) The molecular basis for wine grape quality - a volatile subject. Science 311:804-805 Robinson AL et al. (2014) Origins of grape and wine aroma. Part 1. Chemical components and viticultural impacts. American Journal of Enology and Viticulture 65:1-24
Aromatic Compound Disposition
Aromatic compound quality profiling in Alsace Gewürztraminer integrates analysis of monoterpenes (linalool 400-800 μg/L, geraniol 200-400 μg/L providing floral-rose character), C13-norisoprenoids (β-damascenone contributing exotic fruit notes), and bound glycosidic precursors (2-4 mg/L potential volatiles releasable during aging) this comprehensive aroma profiling enables selection of fruit with optimal aromatic potential for the intensely perfumed, spicy-floral wine style that defines great Gewürztraminer, distinguishing aromatically expressive lots from neutral fruit suitable only for blending.
Aromatic_Compound_Quality encompasses all chemical qualities that characterize the concentration, composition, and proportional relationships of volatile and semi-volatile compounds that contribute to the olfactory perception of grapes and wine. Aroma compounds represent the chemical basis for one of the most valued and complex dimensions of wine quality, determining varietal typicity, regional character, and aging potential through their interaction with olfactory receptors. This category addresses compounds spanning an enormous structural diversity, from simple alcohols and esters through terpenes and norisoprenoids to sulfur-containing thiols and nitrogen-containing pyrazines. The category subdivides into Primary_Aroma_Quality capturing grape-derived aromatic compounds that define varietal character Secondary_Aroma_Quality addressing fermentation-generated volatiles and Tertiary_Aroma_Quality encompassing compounds formed during aging through chemical transformation. Primary aromas include terpenes (Muscat character), methoxypyrazines (herbaceous notes), and thiol precursors (tropical fruit potential). Secondary aromas arise from yeast and bacterial metabolism, producing esters, higher alcohols, and volatile acids. Tertiary aromas develop through oxidation, reduction, and hydrolysis reactions during aging, releasing bound aroma compounds and generating novel structures. The concentration of individual aroma compounds varies from parts per trillion (methoxypyrazines) through parts per billion (thiols) to parts per million (esters), with perception depending on both absolute concentration and sensory threshold. The enormous number of volatile compounds in wine (several hundred identified) creates additive, synergistic, and masking interactions that produce perceived aroma complexity exceeding what individual compound concentrations would suggest. Aromatic compound analysis employs gas chromatography with various detection systems, requiring careful sample preparation to capture the range of compound classes and concentrations relevant to aroma quality.
Aromatic_Compound_Quality is positioned as a direct subclass of Chemical_Quality, parallel to Sugar_Content_Quality, Acid_Content_Quality, and Phenolic_Content_Quality. The enormous structural diversity and wide concentration range of aroma compounds creates analytical challenges comprehensive profiling requires multiple extraction and detection methods. The distinction between free volatiles (directly odor-active) and bound precursors (potential aroma released by hydrolysis) is important because both contribute to wine aroma development. The term aromatic here refers to olfactory perception rather than the chemical definition of aromaticity (ring structures with delocalized electrons).
Aromastoffgehalt
Aromaverbindungsqualität
Duftstoffzusammensetzung
aroma compound content
fragrance compound composition
olfactory compound quality
olfaktorische Verbindungsqualität
volatile compound quality
Rapp A (1998) Volatile Flavour of Wine: Correlation between Instrumental Analysis and Sensory Perception. Nahrung 42(6): 351-363 Ferreira V (2010) Volatile Aroma Compounds and Wine Sensory Attributes. In: Reynolds AG (ed) Managing Wine Quality Volume 1: Viticulture and Wine Quality, Woodhead Publishing pp.: 3-28
Aromatic Compound Quality
Aromatic maturity assessment in Sauvignon Blanc comparing harvest timing effects on wine aroma profile reveals the compound-specific evolution enabling style selection, with early harvest (21 Brix) showing high methoxypyrazines (25 ng/L IBMP, herbaceous character), low thiol precursors (limited tropical potential), and moderate terpenes (subtle floral), mid-harvest (23 Brix) showing declining methoxypyrazines (12 ng/L, balanced herbaceous), increased thiol precursors (tropical potential developing), and stable terpenes, and late harvest (25 Brix) showing minimal methoxypyrazines (5 ng/L, no herbaceous), maximum thiol precursors (strong tropical potential), but declining total terpenes, demonstrating how aromatic maturity assessment enables deliberate harvest timing for target aromatic styles ranging from herbaceous-citrus to tropical-rich.
Aromatic maturity quality is a maturity quality that represents the developmental state of aroma and aroma precursor compounds in grape berries, characterizing the accumulation and evolution of volatile compounds and their glycosidically bound precursors that determine the aromatic profile of wines, including varietal character, fermentation-derived aromas, and aging bouquet development potential. Within the BFO framework, this quality specifically addresses the aromatic compound dimension of maturity, distinguishing aroma development from sugar, acid, or phenolic parameters while recognizing that aromatic maturity may follow different timing than other maturity dimensions and critically determines the varietal expression and complexity that distinguish fine wines. The ontological significance lies in aromatic maturity determining the sensory identity and complexity of wines through volatile compounds perceived directly (free volatiles) and indirectly (precursors released during fermentation and aging), where the accumulation of terpenes, norisoprenoids, thiols, methoxypyrazines, and their glycosidic precursors establishes the aromatic potential that winemaking can express but not create de novo. The compound classes contributing to aromatic maturity include monoterpenes (linalool, geraniol providing floral character in aromatic varieties), norisoprenoids (β-damascenone, β-ionone contributing to complexity), volatile thiols (4-mercapto-4-methylpentan-2-one providing tropical character), C₁₃-norisoprenoids derived from carotenoid degradation (contributing to aged character development), and methoxypyrazines (providing herbaceous character that typically declines during ripening), with each class showing distinct accumulation or degradation patterns that affect the aromatic profile at different harvest times. The developmental patterns of aromatic compounds show substantial variation, with terpenes accumulating primarily pre-véraison then remaining stable or declining, norisoprenoid precursors increasing through ripening as carotenoid degradation proceeds, thiol precursors increasing during ripening (favoring later harvest for tropical character), and methoxypyrazines declining during ripening (favoring later harvest to reduce herbaceous character), such that different aromatic profiles are achievable through harvest timing within a single vineyard. The environmental and management factors affecting aromatic maturity include light exposure (favoring monoterpene and norisoprenoid accumulation, accelerating methoxypyrazine degradation), temperature (affecting metabolic rates and volatilization losses), and water status (moderate deficit often enhancing aromatic concentration), with these factors creating the site-specific and vintage-specific aromatic maturity patterns that contribute to wine distinctiveness. Understanding aromatic maturity quality enables harvest timing for desired aromatic profile, assessment of varietal expression potential, and recognition of the compound-specific accumulation patterns that create the aromatic character options available at different harvest times.
Aromatic_Maturity_Quality serves as the parent class for aromatic compound qualities (Precursor, Varietal Development, Stability). This quality addresses the volatile and precursor compounds determining wine aromatic character. The class supports aromatic style optimization through aromatic maturity characterization.
Aromareife
Duftentwicklung
Geschmacksreife
aroma maturity
aromatic ripeness
aromatische Reife
flavor maturity
volatile maturity
Kalua CM, Boss PK (2009) Evolution of volatile compounds during the development of Cabernet Sauvignon grapes (Vitis vinifera L.). Journal of Agricultural and Food Chemistry 57(9): 3818-3830 Roland A et al. (2011) Evolution of varietal thiols during wine oxidation. Journal of Agricultural and Food Chemistry 59(9): 4653-4660
Aromatic Maturity Quality
Gewürztraminer bears an Aromatic_White_Wine_Scion_Role as perhaps the most intensely perfumed wine grape, with monoterpene concentrations of 3-6 mg/L (primarily geraniol, linalool, citronellol) creating rose petal, lychee, and spice aromatics that define Alsace Grand Cru and Alto Adige expressions. Muscat Blanc à Petits Grains (Muscat de Frontignan) serves in an Aromatic_White_Wine_Scion_Role producing intensely grapey, floral wines with terpene concentrations exceeding 5 mg/L across diverse styles from dry Alsace Muscat through sweet Muscat de Beaumes-de-Venise and Moscato d'Asti. Torrontés bears an Aromatic_White_Wine_Scion_Role as Argentina's signature aromatic variety, combining Muscat-derived terpenes (as a Muscat × Criolla Grande cross) with distinctive floral-spicy character in Salta and Cafayate.
A white wine scion role that inheres in a grafted scion cultivar by virtue of its genetically determined capacity to accumulate high concentrations of varietal aroma compounds—particularly monoterpenes, thiols, and norisoprenoids—that create distinctive, intense, and readily recognizable aromatic profiles in the resulting wines. Aromatic white wine scions are characterized by free and glycosidically bound terpene concentrations exceeding 1-4 mg/L (versus 0.1-0.5 mg/L in neutral varieties), with specific compound profiles defining varietal identity: linalool and geraniol in Muscat varieties (5-10 mg/L combined), linalool and α-terpineol in Riesling (1-3 mg/L), and rotundone in Grüner Veltliner. Thiolic varieties (Sauvignon Blanc, Colombard) accumulate cysteine-bound precursors released during fermentation to yield volatile thiols (3-mercaptohexanol, 4-mercapto-4-methylpentan-2-one) at ng/L concentrations with detection thresholds below 5 ng/L, creating intense tropical and herbaceous aromatics. Aromatic varieties are typically vinified without oak or malolactic fermentation to preserve primary fruit character, though some (Gewürztraminer, Viognier) may receive limited oak contact.
Aromatic_White_Wine_Scion_Role captures a genetically determined flavor category defined by aroma precursor accumulation rather than wine style or quality tier. The monoterpene biosynthesis pathway (via geranyl pyrophosphate and linalool synthase) is variably expressed among cultivars, with Muscat family varieties showing the highest expression levels. Thiolic aromatics in Sauvignon Blanc and related varieties involve different biochemical pathways (cysteine conjugate accumulation) but create similarly intense varietal character. Aromatic varieties present winemaking challenges—high terpene musts are susceptible to oxidative aroma loss, and some aromatic compounds (β-damascenone, TDN) develop through aging, creating evolution potential in varieties like Riesling. Climate strongly influences aromatic compound accumulation, with cool conditions generally favoring terpene retention.
Bukettsorten
aromatic grape variety
aromatic white variety
aromatische Rebsorte
aromatische Weißweinsorte
perfumed white variety
terpenic variety
terpenreiche Rebsorte
Mateo JJ, Jiménez M (2000) Monoterpenes in grape juice and wines, Journal of Chromatography A 881: 557-567 Tominaga T et al. (1998) Development of a method for analyzing the volatile thiols in wines, Journal of Agricultural and Food Chemistry 46: 1044-1048 Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Black CA et al. (2015) Terpenoids and their role in wine flavour, Australian Journal of Grape and Wine Research 21: 582-600
Aromatic White Wine Scion Role
Arrested development assessment in severely drought-stressed Tempranillo reveals the progressive failure of developmental processes, with early-season arrest showing shoot growth cessation at 40 cm (normal 90 cm), followed by mid-season arrest of berry development at green pea stage (berries remaining at 0.4 g, normal progression to 1.2 g halted), and eventual arrested ripening with berries reaching only 18 Brix (target 24 Brix) with green, harsh tannins persisting despite late harvest date, demonstrating the cascading effects of arrested development through multiple phenological phases resulting in complete crop failure for quality wine production.
Arrested development quality is a developmental completeness quality that represents the cessation of developmental progress before stage completion, characterizing the condition where phenological advancement stops prematurely due to severe stress, resource exhaustion, or other factors that halt normal developmental processes, leaving organs in incomplete developmental states that cannot progress further without intervention or changed conditions. Within the BFO framework, this quality specifically addresses the cessation dimension of developmental disruption, distinguishing complete developmental arrest from gradual incomplete development while recognizing that arrested development represents a more severe disruption than slow or incomplete progress, with the developmental process stopping rather than merely slowing. The ontological significance lies in arrested development indicating severe disruption of normal vine function, where the cessation of developmental progress signals conditions incompatible with continued development (extreme water stress, critical nutrient deficiency, temperature extremes) that require immediate attention to prevent permanent damage or crop loss, making arrested development a critical diagnostic indicator of vine condition. The physiological basis for arrested development includes depletion of carbohydrate reserves (insufficient energy to continue development), water stress exceeding critical thresholds (cellular function compromised), temperature extremes beyond developmental tolerances (enzymatic processes inhibited), and hormonal disruption (developmental signaling compromised), with the specific cause determining both the symptoms and the potential for recovery. The stage-specific manifestations of arrested development include arrested bud development (buds failing to progress beyond early stages, potentially remaining dormant), arrested shoot growth (growth cessation with stunted shoots, terminal bud formation), arrested berry development (berry growth stopping at intermediate sizes, "green berry syndrome"), and arrested ripening (sugar accumulation ceasing before target maturity, "stuck fermentation in the vineyard"), with each manifestation having specific causes and remediation approaches. The recovery potential from arrested development depends on the severity and duration of arrest, the developmental stage affected, and the availability of resources for resumed development, with early-season arrest potentially recoverable through stress alleviation while late-season arrest may result in permanent quality loss. Understanding arrested development quality enables rapid diagnosis of severe vine stress, prioritization of management interventions, and realistic assessment of recovery potential based on the timing and nature of developmental arrest.
Arrested_Development_Quality addresses the cessation of developmental progress before stage completion due to severe stress. This quality indicates serious disruption requiring immediate intervention. The class supports stress diagnosis and emergency response through arrested development identification.
Entwicklungsarrest
Entwicklungsstopp
Wachstumsstillstand
arretierte Entwicklung
development cessation
developmental arrest
growth stoppage
halted development
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 267-330 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data. Annals of Botany 105(5): 661-676
Arrested Development Quality
Astringency character comparison between Pinot Noir (thin-skinned, small seeds) and Cabernet Sauvignon (thick-skinned, larger seeds) at equivalent astringency intensity of 5.5 reveals Pinot rated 7.2 on a silky-coarse scale (toward silky) versus Cabernet at 4.8 (toward coarse), with gel permeation chromatography showing Pinot tannins averaging 2100 Da versus Cabernet at 1400 Da, demonstrating the variety-driven molecular weight difference that creates the qualitative character distinction between these varieties' tannin profiles.
Astringency character quality is an astringency quality that represents the qualitative nature of the tactile drying sensation distinguishing between different textural experiences ranging from fine and silky through dry and chalky to coarse and grippy, characterizing the type of astringent sensation independent of its intensity. Within the BFO framework, this quality specifically addresses the qualitative dimension of astringency perception, distinguishing character from intensity while recognizing that the same astringency intensity can manifest as qualitatively different experiences depending on tannin structure and wine matrix composition. The ontological significance lies in astringency character strongly influencing hedonic response and quality perception, where fine, silky astringency is universally preferred to coarse, aggressive astringency at equivalent intensity levels, with character often differentiating quality tiers more than intensity alone. The molecular basis for astringency character involves tannin structural features, with smaller oligomeric tannins producing harsher, more aggressive astringency and larger polymeric tannins creating softer, rounder perception, while tannin galloylation pattern, anthocyanin incorporation, and interactions with polysaccharides further modulate qualitative character. The descriptive vocabulary for astringency character spans a multidimensional quality space, with the dry-grippy axis capturing one dimension (from silky through satiny through dry to chalky to grippy to aggressive), roughness capturing another (smooth to rough to coarse), and particle sensation capturing a third (fine to grainy to particulate), together characterizing the textural quality of astringent mouthfeel. The evolution of astringency character during wine aging typically shifts from aggressive, drying character in young wines toward softer, silkier character in mature wines as tannin polymerization proceeds, explaining why character assessment informs aging potential prediction. Understanding astringency character quality enables refined assessment beyond intensity, identification of tannin quality issues requiring winemaking intervention, and quality description capturing the textural dimension that strongly affects consumer preference.
Astringency_Character_Quality_(Dry_vs_Grippy) addresses the qualitative dimension of astringent mouthfeel, complementing Astringency_Intensity_Quality. The quality name references the dry-grippy axis as a primary character dimension, though full character description encompasses additional dimensions (silky-coarse, smooth-rough). This quality depends on tannin molecular properties rather than concentration, explaining why character can vary independently of intensity.
Adstringenzcharakter
Adstringenztextur
Art der Adstringenz
Tannincharakter
astringency type
astringent quality type
tannin character
tannin texture
Gawel R et al. (2000) A 'Mouth-feel Wheel': terminology for communicating the mouth-feel characteristics of red wine. Australian Journal of Grape and Wine Research 6(3): 203-207 Chira K et al. (2009) Characterization of red wine proanthocyanidins and their affinity for salivary proteins. Analytical and Bioanalytical Chemistry 395(4): 1171-1179
Astringency Character Quality (Dry vs Grippy)
Astringency development tracking in young Ribera del Duero Tempranillo using time-intensity methodology reveals rapid build-up reaching maximum intensity at 8 seconds post-contact for wines fermented with whole berry inclusion (30% stems) versus gradual development peaking at 18 seconds for destemmed fermentation, with the faster development in stem-included wine perceived as more aggressive despite equivalent peak intensity of 6.5, explaining the preference for destemmed wines at the consumer level while stem-included wines show integration improvement with 3+ years aging.
Astringency development quality is an astringency quality that represents the temporal pattern of astringency emergence and intensification during wine evaluation, characterizing how quickly and through what trajectory astringent sensation builds from initial perception to maximum intensity. Within the BFO framework, this quality specifically addresses the temporal emergence dimension of astringency, distinguishing the build-up dynamics from peak intensity and qualitative character while recognizing that development pattern affects perceived harmony and balance even at equivalent maximum intensity. The ontological significance lies in astringency development affecting the integration of tannin with other sensory attributes, where gradual development allows harmonious interaction with fruit and acidity perceptions while rapid aggressive development creates the impression of tannin disconnection from other wine components even when final intensity is appropriate. The physiological basis for astringency development involves the kinetics of tannin-protein binding, salivary protein depletion rate, and neural adaptation patterns, with these factors creating the characteristic build-up where astringency intensifies over the 15-30 seconds following wine contact before reaching maximum and beginning to fade. The compositional factors affecting development include tannin molecular size (smaller tannins showing faster development), tannin concentration (higher levels accelerating build-up), and matrix composition (polysaccharides and proteins slowing development by competing for tannin binding). The relationship between development pattern and quality shows preference for moderate build-up rates where astringency emerges gradually and peaks in harmony with fading fruit sweetness, while very rapid development creates harsh initial impact and very slow development may produce disconnected astringency that emerges after other flavors have faded. Understanding astringency development quality enables temporal profile characterization complementing intensity and character assessment, prediction of development dynamics from compositional analysis, and winemaking optimization for harmonious tannin integration.
Astringency_Development_Quality addresses the temporal emergence of astringent sensation, complementing Astringency_Intensity_Quality (peak magnitude) and Astringency_Character_Quality (textural type). This quality captures the dynamic build-up pattern that affects perceived harmony. Time-intensity methodology provides quantitative development assessment, though qualitative description of build-up character is also valuable for comprehensive evaluation.
Adstringenzanstieg
Adstringenzentwicklung
Tanninaufbau
astringency build-up
astringency onset
tannin emergence
temporal astringency pattern
zeitlicher Adstringenzverlauf
Guinard JX et al. (1986) The time-course of astringency in wine upon repeated ingestion. American Journal of Enology and Viticulture 37(3): 184-189 Lee CB, Lawless HT (1991) Time-course of astringent materials. Chemical Senses 16(3): 225-238
Astringency Development Quality
Astringency intensity measurement comparing Malbec wines with and without extended cold maceration reveals intensity of 6.8 versus 5.2 on a 10-point scale for the extended and standard protocols respectively, with the 31% higher intensity in extended maceration corresponding to 45% higher condensed tannin concentration (1850 versus 1275 mg/L), enabling winemakers to calibrate maceration duration to achieve target astringency intensity for the intended market segment where export markets prefer lower intensity (target 5.0) than domestic Argentine market (target 6.5).
Astringency intensity quality is an astringency quality that represents the magnitude of the drying, puckering tactile sensation at its maximum perception during wine evaluation, characterizing the peak astringency level that determines the structural impression and compatibility with food pairing. Within the BFO framework, this quality specifically quantifies the intensity dimension of astringency at its strongest point, distinguishing intensity from qualitative character (dry versus grippy) and temporal aspects (development, evolution), providing the primary parameter for assessing tannin level in red wine evaluation. The ontological significance lies in astringency intensity determining whether tannin contributes positively as structural backbone or negatively as harsh, drying excess, with moderate intensity (appropriately matched to wine body and fruit concentration) creating the valued grip and structure while excessive intensity overwhelms fruit and creates unpleasant mouthfeel. The psychophysical relationship between tannin concentration and perceived astringency intensity shows approximately linear relationship in model solutions but substantial modulation by wine matrix, where ethanol, glycerol, polysaccharides, and residual sugar all reduce perceived astringency at equivalent tannin levels. The factors modulating astringency intensity include wine pH (lower pH increasing perception), ethanol content (complex effects with moderate alcohol reducing astringency), polysaccharide content (mannoproteins softening perception), and prior exposure (within-session adaptation reducing intensity with repeated tasting). In wine quality assessment, astringency intensity is evaluated relative to wine style expectations, with age-worthy reds appropriately showing higher intensity than drink-young styles, and food-pairing contexts tolerating higher intensity than aperitif consumption. Understanding astringency intensity quality enables calibration of sensory assessment, prediction of perceived astringency from phenolic analysis accounting for matrix effects, and winemaking optimization to achieve the astringency level appropriate for target style and consumption context.
Astringency_Intensity_Quality addresses the magnitude dimension of astringency at peak perception, complementing Astringency_Character_Quality (qualitative type), Astringency_Development_Quality (temporal emergence), and Tannin_Mouthfeel_Quality (textural character). Matrix effects significantly modulate the tannin-to-astringency relationship, requiring either sensory measurement or matrix-adjusted prediction models for accurate intensity assessment.
Adstringenzintensität
Mundtrockenheitsintensität
Stärke der Adstringenz
Tanninintensität
astringency strength
drying magnitude
pucker intensity
tannin intensity
Vidal S et al. (2003) The mouth-feel properties of grape and apple proanthocyanidins in a wine-like medium. Journal of the Science of Food and Agriculture 83(6): 564-573 Rinaldi A et al. (2012) Interaction between polymeric proanthocyanidins and salivary proteins. Food Quality and Preference 24(2): 250-255
Astringency Intensity Quality
Astringency quality evaluation in Tannat comparing young (18 months) and aged (8 years) wines reveals astringency intensity declining from 7.8 to 4.5 on a 10-point scale while astringency character shifts from harsh and drying to smooth and velvety, with total phenolics decreasing only 15% (from 3200 to 2720 mg/L GAE) while mean tannin molecular weight increases 180%, demonstrating that astringency evolution depends on tannin structure rather than concentration, and supporting the extended aging required for this high-tannin variety to achieve optimal drinkability.
Astringency quality is a sensory quality that represents the tactile drying, roughening, and puckering sensation in the oral cavity arising from precipitation of salivary proteins by polyphenolic compounds, characterizing a fundamental mouthfeel attribute that defines red wine structure and influences quality perception across wine styles. Within the BFO framework, this quality specifically addresses astringency as a tactile (somesthetic) sensation mediated by mechanoreceptors detecting changes in oral surface lubrication, distinguishing astringency from the gustatory sensation of bitterness though both are associated with phenolic compounds and often co-occur in wine evaluation. The ontological significance lies in astringency providing the structural framework of red wines, where appropriate astringency creates the grip, texture, and palate-cleansing properties valued for food pairing while contributing to age-worthiness through tannin evolution potential, whereas excessive or harsh astringency indicates quality deficiency requiring aging or remediation. The physiological mechanism involves polyphenol-protein binding where tannins complex with proline-rich salivary proteins causing aggregation and precipitation, reducing oral lubrication and creating friction detected by mechanoreceptors as the drying, roughening sensation, with repeated exposure during wine drinking progressively depleting salivary proteins and intensifying perception. The compositional basis for wine astringency includes condensed tannins (proanthocyanidins) from grape skins and seeds, hydrolyzable tannins from oak aging, and anthocyanin-tannin complexes formed during fermentation and aging, with molecular size, structure, and concentration affecting astringency intensity and character. The developmental trajectory during wine aging shows astringency typically decreasing as tannins polymerize to sizes too large for protein binding and precipitate, creating the softening that distinguishes mature from young red wines. Understanding astringency quality enables assessment of tannin development and integration, prediction of aging trajectory, and winemaking optimization for appropriate astringency in the target wine style.
Astringency_Quality serves as the parent class for intensity, character, development, and mouthfeel subclasses, establishing the framework for this tactile sensation distinct from taste. The distinction between astringency (tactile) and bitterness (gustatory) is ontologically critical though often confused in casual wine description. This class addresses the mechanical sensation of reduced lubrication, not the taste of phenolic compounds which contributes to bitterness.
Adstringenz
Gerbstoffempfindung
Mundtrockenheit
mouth-drying quality
oral roughening
pucker sensation
tannin sensation
zusammenziehende Wirkung
Gawel R (1998) Red wine astringency: a review. Australian Journal of Grape and Wine Research 4(2): 74-95 Breslin PAS et al. (1993) Cellular basis of taste perception. Physiology & Behavior 54(6): 1129-1134
Astringency Quality
Syrah hermaphrodite flowers at BBCH 59-61 show asymmetrical bicarpellate pistils in approximately 12-18% of flowers, with transverse ovary sections revealing two locules of unequal size (larger locule 0.28-0.35 mm² cross-sectional area, smaller locule 0.18-0.24 mm² area, representing 30-45% size difference), 3 ovules distributed as 2 in larger locule and 1 in smaller locule, central septum displaced 15-25% off-center toward the smaller locule, and stigma showing subtle asymmetry with enhanced papillae on the side corresponding to the larger carpel. Merlot under water stress during BBCH 52-54 (early pistil development) exhibits increased asymmetrical bicarpellate pistil frequency (24-32% versus normal 10-15%), with stressed pistils showing pronounced locule size differences (40-60% larger versus smaller) and asymmetric ovule distribution (2 versus 0-1 ovules per locule), though fruit set rates from asymmetric stressed pistils (38-48%) remain comparable to symmetric unstressed pistils (48-56%), suggesting pistil asymmetry has limited impact on fertilization success. Gewürztraminer shows constitutively higher asymmetric pistil frequency (18-24% even under optimal conditions) representing genotype-specific variation in carpel development coordination, with asymmetric pistils producing berries showing asymmetric seed distribution (seeds concentrated in positions corresponding to originally larger locule) but normal commercial quality. Cabernet Franc berries developing from asymmetric pistils (identified retrospectively through seed position analysis) show 1-3 seeds with 68% of seeds located in positions corresponding to the originally larger locule and 32% in smaller locule positions, demonstrating that locule asymmetry influences but does not completely determine final seed distribution patterns.
An asymmetrical bicarpellate pistil is a bicarpellate pistil in which the two constituent carpels show unequal development creating asymmetric ovary morphology with two locules of notably different sizes (differing by >20% in volume or cross-sectional area), potentially unequal ovule distribution between locules, and sometimes asymmetric stigma organization, representing a less common variant bicarpellate pistil morphology resulting from differential carpel development during organogenesis. Asymmetrical bicarpellate pistils are characterized by= (1) bilocular ovary with two locules of unequal volume and shape, evident in transverse cross-sections as two chambers of different dimensions, typically with one locule 20-60% larger than the other in cross-sectional area (2) central septum (partition separating locules) positioned off-center from the ovary median plane, displaced toward the smaller locule and creating unequal division of ovary volume (3) ovule distribution often showing unequal numbers per locule, commonly 2 ovules in the larger locule versus 1 ovule in the smaller locule (total 3), or 2 versus 0 (total 2), or other asymmetric distributions, though occasionally asymmetric locules may contain equal ovule numbers (2 per locule) with asymmetry in locule size rather than ovule number (4) stigma sometimes showing asymmetric organization with unequal lobe development or asymmetric papillae distribution, though stigma asymmetry is less pronounced than ovary asymmetry in most cases (5) style occasionally showing asymmetric transmitting tract organization providing unequal pollen tube access to the two locules. Asymmetrical bicarpellate pistils occur in 10-20% of flowers in most Vitis cultivars under typical conditions, representing the secondary developmental outcome where carpel development coordination is disrupted creating differential carpel elaboration. The asymmetric development results from= (1) asynchronous initiation of two carpel primordia creating developmental head-start for the first-initiated carpel (2) differential resource allocation to the two developing carpels during organogenesis, with the larger carpel receiving more resources (nutrients, hormones, photosynthates) than the smaller carpel (3) uncoordinated cell proliferation or differentiation in the two carpels creating unequal tissue elaboration (4) asymmetric environmental or developmental perturbations affecting one carpel more than the other during sensitive developmental windows. Environmental stresses during pistil development (BBCH 52-54) can increase asymmetrical pistil frequency= water stress, heat stress, or nutritional imbalances during carpel organogenesis may disrupt the developmental coordination normally producing symmetric carpels, increasing asymmetric pistil frequency from baseline 10-15% to 20-35% depending on stress severity and timing. Genetic variation also influences asymmetry frequency, with some cultivars showing constitutively higher asymmetric pistil rates (15-25% asymmetric even under optimal conditions) while others show highly consistent symmetry (>92% symmetric). Asymmetrical bicarpellate pistils show potential functional consequences for reproduction= unequal locule size may bias fertilization success toward the larger locule if pollen tube growth preferentially accesses the larger chamber through the style transmitting tract unequal ovule distribution creates unequal seed number potential between locules, potentially affecting seed distribution patterns in berries extreme asymmetry (one locule >60% larger than the other or one locule containing all ovules while other is empty) may impair overall fertilization efficiency or fruit set probability. However, moderate asymmetry (20-40% size difference) appears compatible with normal fruit set rates in most cultivars, with asymmetric pistils achieving 35-55% fruit set comparable to symmetric pistils (40-60% fruit set), suggesting that pistil function tolerates substantial asymmetry. Berries developing from asymmetric pistils often show asymmetric seed distribution with more seeds in positions corresponding to the originally larger locule, and may show slightly irregular berry morphology with subtle bilateral asymmetry, though commercial fruit quality is rarely affected by pistil asymmetry unless asymmetry is extreme.
Asymmetrical_Bicarpellate_Pistil represents unequal carpel development within the two-carpel structure and is distinguished from Symmetrical_Bicarpellate_Pistil by locule size differences >20% versus <20%. Asymmetry represents a secondary developmental outcome resulting from disrupted carpel coordination, increased by environmental stresses or present constitutively in some genotypes. Despite structural asymmetry, reproductive function generally remains adequate with fruit set rates comparable to symmetric pistils in most cases. Understanding asymmetry is relevant for assessing environmental stress effects on pistil development, for interpreting genotypic variation in developmental stability, and for understanding seed distribution patterns in berries. The threshold of 20% size difference for asymmetry classification is somewhat arbitrary but reflects a practical boundary between minor developmental variation (functional symmetry) and notable structural asymmetry with potential functional implications.
asymmetrischer bikarpellater Stempel
unausgeglichener Zwei-Fruchtblatt-Stempel
unbalanced bicarpellate pistil
unequal-carpel pistil
ungleichmäßiges Zwei-Karpell-Pistill
Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 93-95 Ebadi, A. et al. (1996). Effect of short-term temperature and shading on fruit set. Australian Journal of Grape and Wine Research 2=2-9.
Asymmetrical Bicarpellate Pistil
An asymmetrical bifurcated tendril at node 9 on a moderately stressed shoot of Vitis vinifera cv. Syrah in the Rhône Valley, France, exhibiting branch point at 6.5 cm from shoot attachment, dividing into a dominant arm of 13 cm length and a subordinate arm of 8.5 cm length (ratio 0.65=1), with branch angles of 35 degrees for the dominant arm and 55 degrees for the subordinate arm creating noticeably skewed architecture, dominant arm showing 3.2 mm diameter at mid-length while subordinate arm measures 2.3 mm indicating differential vascular investment, dominant arm exhibiting vigorous circumnutation with 160-degree amplitude while subordinate arm shows reduced 95-degree amplitude, occurring in a vineyard experiencing progressive water stress (soil moisture declining from 75% to 55% field capacity) during tendril development causing resource limitation favoring the earlier-initiated dominant arm, measured at BBCH stage 75 (berries pea-sized) in late June, representing stress-induced asymmetry rather than genetic tendency in this cultivar.
A bifurcated tendril exhibiting unequal development of the two branches in Vitis vinifera, characterized by arm length ratio less than 0.85=1 indicating differential resource allocation or developmental timing between branches, often accompanied by unequal branch angles creating skewed Y-shaped architecture with one arm more prominent than the other, differential branch diameter reflecting asymmetric vascular development and structural investment, and potentially divergent functional capabilities including stronger circumnutation in the dominant arm, more vigorous coiling response, and greater load-bearing capacity compared to the subordinate arm. Asymmetrical bifurcated tendrils develop through multiple mechanisms including sequential rather than simultaneous activation of lateral meristems at the branch point creating developmental head-start for the first-activated branch, differential resource allocation during growth with preferential nutrient and carbohydrate flow to one branch based on vascular architecture or hormonal gradients, unequal environmental exposure with one branch receiving more favorable light, temperature, or moisture conditions, or differential damage or stress affecting one branch while sparing the other during critical development periods. This branching pattern represents compromised expression of bifurcated architecture compared to symmetrical development, but maintains functional advantages over unbranched tendrils through dual searching capacity despite unequal arm development, with the dominant arm often achieving functional equivalence to an unbranched tendril while the subordinate arm provides supplementary exploration capacity and potential backup attachment opportunity.
Asymmetrical bifurcated tendrils are distinguished from symmetrical bifurcated tendrils by arm length ratio <0.85=1, differential structural development, and potentially divergent functional capabilities. The degree of asymmetry provides diagnostic information about stress severity or developmental disruption, with mild asymmetry (ratios 0.70-0.84=1) indicating moderate differential development while severe asymmetry (ratios <0.60=1) suggests major developmental problems or damage to the subordinate branch. Asymmetry can arise from genetic programming in some cultivars showing consistent asymmetric patterns, or from environmental factors creating situational asymmetry in cultivars normally producing symmetrical bifurcation. Assessment of asymmetry patterns across vine populations helps distinguish genetic versus environmental causation, with uniform asymmetry suggesting cultivar characteristic while variable asymmetry indicates stress effects. Functionally, asymmetrical bifurcation still provides advantages over unbranched architecture but with reduced efficiency compared to symmetrical bifurcation.
asymmetrisch gegabelte Ranke
dominant-subordinate tendril
schiefsymmetrische Y-förmige Kletterorgan
skewed forked tendril
unequal bifurcated tendril
ungleichmäßig verzweigte Ranke
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 105-119 Petrie et al. (2000) "The effect of leaf removal and canopy height on whole-vine gas exchange and fruit development of Vitis vinifera L. Sauvignon blanc" Functional Plant Biology 27=173-184 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 137-154.
Asymmetrical Bifurcated Tendril
An asymmetrical bilateral permanent arm on Vitis vinifera cv. Pinot Noir at row end position, featuring short arm at 45cm extending toward row interior and long arm at 95cm extending toward row end (50cm difference, 2.1=1 ratio), short arm bearing 7 spurs while long arm bears 16 spurs, crop distribution approximately 65% on long arm versus 35% on short arm (total 4.2kg), trunk positioned 28cm off-center toward row interior due to planting irregularity or deliberate row-end spacing, representing functional bilateral architecture accommodating row-end constraints while maintaining productive capacity through extended arm compensating for shortened arm, with total spur count (23) and crop (4.2kg) appropriate for vine vigor despite asymmetrical configuration.
A bilateral permanent arm configuration with substantial dimensional or capacity differences between the two arms, characterized by unequal arm lengths (typically differing by 20cm or more), imbalanced spur counts or spacing, skewed crop distribution departing significantly from 50=50 split (often 60=40 or more extreme), and architectural asymmetry resulting from off-center trunk positioning, directional vigor differences, establishment problems, or deliberate design accommodating site constraints or row-end positions. Asymmetrical bilateral permanent arms represent practical accommodations to real-world constraints in vineyard establishment and management, accepting architectural imperfection to maximize productivity within available space, adapt to vine vigor patterns, or work around positioning limitations, with functional performance often equivalent to symmetrical configurations despite aesthetic differences.
Asymmetrical bilateral permanent arms are common and functionally acceptable in commercial vineyards, arising from various causes including off-center trunk positioning (planting irregularities, vine replacement, deliberate row-end spacing), directional vigor differences (slope aspects, wind patterns, equipment traffic effects), establishment challenges (damage to one arm requiring retraining, poor initial growth on one side), and deliberate design (row ends, curved rows, irregular spacing). While aesthetically less satisfying than symmetrical configurations, asymmetrical arms often perform equivalently in terms of total productivity, fruit quality, and management ease, with experienced vineyard managers viewing minor asymmetries as normal variation requiring no correction and major asymmetries as acceptable adaptations to constraints rather than failures requiring remediation.
asymmetric cordon
asymmetrische bilaterale Arme
unausgeglichene Doppelarme
unbalanced bilateral arms
unequal paired arms
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Reynolds & Vanden Heuvel (2009) "Influence of grapevine training systems on vine growth and fruit composition= a review" American Journal of Enology and Viticulture 60=251-268.
Asymmetrical Bilateral Permanent Arm
Berry asymmetry in Pinot Gris commonly results from the cultivar's tendency toward irregular seed set, with single-seeded berries displaying pronounced asymmetry (one side distinctly more convex) accounting for 15-30% of berries in typical clusters this seed-induced asymmetry affects berry size distribution within clusters and can complicate optical berry sorting systems that assume symmetric berry geometry, requiring algorithm adjustments for accurate volume estimation in Pinot Gris compared to more regularly seeded cultivars.
Asymmetry_Quality represents the condition where a grapevine organ lacks regular symmetry relationships, exhibiting form that does not correspond under reflection across any plane or rotation around any axis. Asymmetry may manifest as directional asymmetry, where organs consistently deviate from symmetry in a particular direction, or as fluctuating asymmetry, where deviations occur randomly without consistent directional bias. In grapevine, asymmetry is common in clusters due to variable fruit set across rachis branches, differential berry development, and the inherently irregular branching architecture of the infructescence. Berry asymmetry frequently results from uneven seed development, where berries containing one seed or seeds positioned asymmetrically within the locule develop asymmetric form with one side more convex than the other. Leaf asymmetry beyond normal bilateral limits can indicate environmental stress during leaf development, viral infection affecting growth regulation, or genetic factors specific to certain cultivars or clones. Asymmetry assessment involves comparing the organ to expected symmetry conditions and quantifying the degree and direction of deviation. Fluctuating asymmetry indices measure random deviations from bilateral symmetry and serve as indicators of developmental stability, with elevated fluctuating asymmetry suggesting environmental or genetic stress during organ formation. Consistent directional asymmetry may be a stable cultivar characteristic or may indicate systematic environmental influences such as prevailing wind direction or asymmetric light exposure.
Asymmetry_Quality represents the absence or substantial violation of symmetry conditions, positioned alongside Bilateral_Symmetry_Quality and Radial_Symmetry_Quality under the Symmetry_Quality category. The distinction between normal biological variation around a symmetric ideal and true asymmetry involves threshold judgments significant asymmetry is typically recognized when asymmetry indices exceed cultivar-specific norms by more than two standard deviations. Asymmetry may be described qualitatively (slight, moderate, pronounced) or quantified through specific indices. PATO_0000616 (asymmetric) provides the upper-ontology reference.
Asymmetrie
Asymmetriequalität
Formunregelmäßigkeit
asymmetric quality
form irregularity
irregular form
lack of symmetry
unregelmäßige Form
Palmer AR and Strobeck C (1986) Fluctuating Asymmetry: Measurement, Analysis, Patterns. Annual Review of Ecology and Systematics 17(1): 391-421 Klingenberg CP (2015) Analyzing Fluctuating Asymmetry with Geometric Morphometrics: Concepts, Methods, and Applications. Symmetry 7(2): 843-934
Asymmetry Quality
Chardonnay attached berry pedicel at mid-ripening (BBCH 85, 20°Brix) shows firm attachment requiring 2.0-2.5 N detachment force, active phloem sugar transport at 1.0-1.5 mg sucrose/berry/hour, incomplete abscission zone with minimal cell wall degradation, tan to light brown coloration indicating partial lignification, and vascular bundles with 25-35 μm diameter xylem vessels maintaining some water transport capacity
An attached berry pedicel is a berry pedicel characterized by firm structural connection to both the berry and the rachis, with incompletely developed abscission zone tissues, representing pedicels during active berry development phases prior to abscission layer maturation. Attached berry pedicel characteristics include= strong mechanical attachment requiring 1.5-3.0 N force for berry detachment (significantly higher than mature pedicels with 0.5-1.5 N detachment force), continuous vascular connection with active xylem and/or phloem transport, incomplete abscission zone differentiation (abscission layer cells show minimal cell wall modifications and limited enzyme accumulation), and green to tan coloration indicating ongoing metabolic activity and incomplete lignification. Structural features of attached berry pedicel include= intact epidermis and cortex tissues without separation planes, vascular bundles with active conducting elements (xylem vessels with functional water transport, phloem sieve tubes with active sugar transport), parenchyma cells with dense cytoplasm and active metabolism, and abscission zone region showing minimal cell wall degradation or separation layer formation. Attached berry pedicel functional states vary by developmental phase= during Phase I berry growth (BBCH 73-77) attached pedicels show maximal xylem activity with high water and nutrient import rates supporting rapid berry expansion, during Phase II lag phase (BBCH 77-81) attached pedicels show both xylem and phloem activity with berry growth slowing and seed development proceeding, and during early Phase III ripening (BBCH 81-85) attached pedicels show increasing phloem dominance with sugar import supporting berry sugar accumulation while xylem flow diminishes. Attached berry pedicel represents the normal state during most of berry development= from fruit set through mid-ripening (BBCH 71-85), pedicels maintain firm attachment supporting berry growth and nutrient supply, with abscission zone development beginning during late ripening (BBCH 85-89) as berries approach harvest maturity. Viticultural significance of attached berry pedicel includes= resistance to premature berry drop (firmly attached berries resist shattering during vineyard operations, wind, or rain events), support of optimal berry development through maintained vascular supply, indication of pre-harvest berry status (firmly attached berries indicate harvest timing may be premature if easy separation is desired), and cluster integrity maintenance (attached pedicels keep berries in cluster configuration supporting uniform ripening and quality). Factors affecting transition from attached to separating pedicel include= berry maturity level (advancing maturity promotes abscission zone development), ethylene production (ripening-associated ethylene triggers abscission zone enzyme synthesis), auxin levels (declining auxin during ripening reduces abscission suppression), environmental stress (water deficit or disease can induce premature abscission zone activation), and cultivar genetics (some varieties develop strong abscission zones while others maintain firm attachment even at overripeness).
Attached_Berry_Pedicel emphasizes firm structural connection and incomplete abscission development and is distinguished from Mature_Berry_Pedicel (developed abscission facilitating separation) by maintenance of strong attachment and active vascular transport without separation layer formation. Critical for understanding berry retention during development, premature drop prevention, harvest timing assessment, and cultivar differences in berry attachment strength affecting mechanical harvest suitability.
Anhaftender Fruchtstiel
Fest verbundener Beerenstiel
Firmly attached pedicel
Nicht-trennender Stiel
Non-separating berry stalk
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Considine, J. A., & Knox, R. B. (1979). Development and histochemistry of the cell, tissue, and secretion within the dermal system of the grape. Protoplasma, 99(3-4), 347-365.
Attached Berry Pedicel
An attached functional tendril at node 7 on a shoot of Vitis vinifera cv. Sangiovese in Chianti, Italy, having established firm grip on a vertical stake through five tight contact coils formed two weeks previously, bearing mechanical load of approximately 450 grams from the distal 70 cm shoot section including seven leaves and two fruit clusters, resisting 800 grams force in pull tests without detachment or visible deformation, showing complete lignification throughout the coiled regions with dark brown coloration and rigid texture, maintaining stable attachment through multiple wind events including one storm with gusts exceeding 60 km/h, measured at BBCH stage 81 (beginning of berry ripening) in early August, demonstrating successful completion of attachment function with projected maintenance of connection through harvest in October.
A functional tendril that has achieved mechanical connection to a support structure and is actively contributing to vine anchoring and climbing function in Vitis vinifera, characterized by establishment of secure grip through coiling, adhesion, or physical entanglement that resists displacement forces, capability of bearing significant mechanical loads from shoot weight, fruit clusters, and wind-induced tensions without detachment or structural failure, and maintenance of connection over extended periods spanning weeks to months or potentially years depending on tendril lignification and environmental conditions. Attached functional tendrils represent successful completion of the tendril's developmental and behavioral program, having progressed from emergence through elongation and searching phases to contact detection and attachment establishment. The attachment mechanism in Vitis relies primarily on coiling rather than adhesive secretions (unlike some other climbing plant families), with mechanical grip generated through friction between coiled tendril surfaces and the support structure, reinforced by progressive lignification that fixes coil geometry and increases tissue rigidity. Attached tendrils redistribute mechanical loads across the vine architecture, with multiple attachment points creating redundant support systems that maintain canopy position despite individual tendril failures, and contributing to overall structural stability that enables the lianescent growth habit characteristic of wild grapevines and requires management through training and trellising in cultivated vineyards.
Attached functional tendrils are distinguished from unattached searching tendrils by presence of mechanical connection to supports, and from detached or failed tendrils by maintenance of secure grip under normal vineyard conditions. Attachment quality varies significantly based on coiling tightness, lignification completion, support structure properties, and environmental history during attachment formation. Tendrils may lose attached status through mechanical failure under excessive loading, progressive lignin degradation in aging tendrils, pathogen infection compromising structural integrity, or intentional removal during canopy management. The duration of functional attachment ranges from several weeks in short-lived seasonal connections to multiple years in well-lignified tendrils on perennial wood, though most tendrils in commercial viticulture provide support for single growing seasons.
anchored tendril
attached climbing organ
befestigte Ranke
greifende Kletterorgan
gripping tendril
secured tendril
verankerte Ranke
Based on Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Bowling & Vaughn (2008) "Structural and immunocytochemical characterization of the adhesive tendril of Virginia creeper" Protoplasma 232=153-163 Steinbrecher & Speck (2018) "Functional morphology of climbing plants" Progress in Botany 79=61-94 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 173-194.
Attached Functional Tendril
Attachment strength quality in Thompson Seedless determines harvest method viability, with the typically loose berry-pedicel connection (detachment force 1.5-2.5 N) enabling efficient dried-on-vine raisin production where berries must separate from the rachis during mechanical shaking, but causing problematic shatter losses of 5-15% during fresh market harvest for table grape sales DOV (dried-on-vine) clonal selections emphasize loose attachment for easy separation, while fresh market selections target tighter attachment (force above 3.0 N) despite identical appearance, illustrating how attachment strength quality differentiates clones for specific end uses.
Attachment_Strength_Quality encompasses the structural integrity qualities characterizing the mechanical connection between grapevine organs and their supporting structures. This category addresses the force required to separate organs from their points of attachment, which determines susceptibility to abscission, shattering, and detachment during handling. The primary attachment relationships in grapevine fruit include the berry-to-pedicel connection (captured in Pedicel_Attachment_Quality and Berry_Detachment_Resistance_Quality), the pedicel-to-rachis junction, and the cluster-to-shoot connection at the peduncle. Each attachment point involves specialized anatomical structures including vascular connections, lignified support tissues, and in some cases abscission zones with programmed cell separation potential. Attachment strength varies among cultivars, with firm-attachment varieties retaining berries tenaciously while loose-attachment varieties shed berries readily under minimal force. This variation has profound implications for harvest method selection, with tight attachment enabling aggressive mechanical harvesting while loose attachment requires gentle handling to prevent shatter. Attachment strength changes during development, generally increasing during fruit growth as vascular connections strengthen, then potentially decreasing during late ripening as abscission zone tissues mature. Environmental factors including water status, temperature, and ethylene exposure modulate attachment strength through effects on abscission zone development and tissue integrity. Assessment of attachment strength employs pull-force measurement with calibrated instruments, standardized drop tests that subject fruit to controlled impacts, and shatter percentage determination after handling operations.
Attachment_Strength_Quality serves as the parent class for specific attachment relationships including Pedicel_Attachment_Quality and Berry_Detachment_Resistance_Quality. The category addresses the mechanical aspects of organ connection distinct from the developmental processes of abscission. Attachment strength represents a continuous variable typically measured in Newtons (N) of pull force, with cultivar-specific norms and quality thresholds. The relationship between attachment strength and harvest damage is non-linear both very weak attachment (shatter) and very strong attachment (tearing) can cause quality defects, with optimal attachment allowing clean separation at the abscission zone.
Anhaftungskraft
Befestigungsstärke
Organverbindungsqualität
Verbindungsfestigkeit
attachment force
connection strength
organ attachment quality
retention strength
Zabadal TJ and Bukovac MJ (2006) Effect of CPPU on Fruit Development of Selected Seedless and Seeded Grape Cultivars. HortScience 41(1): 154-157 Hedberg PR and Goodwin PB (1980) Factors Affecting Natural and Ethephon-Induced Grape Berry Abscission. American Journal of Enology and Viticulture 31(2): 109-113
Attachment Strength Quality
Shiraz clone BVRC12 bears an Australian_Clone_Role as a Barossa Valley selection maintained through South Australian vine improvement programs, representing old Australian Shiraz genetics predating modern French imports and valued for traditional Australian wine character. Chardonnay clone I10V1 serves in an Australian_Clone_Role developed for irrigated production conditions, with heat tolerance and productivity suited to Murray-Darling and Riverina bulk wine production. Cabernet Sauvignon clone SA125 bears an Australian_Clone_Role from South Australian programs, selected for Australian conditions and widely planted before extensive French clone importation diversified the national clone base.
A regional clone role that inheres in a grapevine clone selected, evaluated, or registered within the Australian grapevine certification system, coordinated through state agriculture departments and research institutions including CSIRO (Commonwealth Scientific and Industrial Research Organisation), the South Australian Research and Development Institute (SARDI), and state vine improvement programs. Australian clones are identified by variety name and alphanumeric codes that may indicate institutional origin (e.g., Shiraz BVRC12 for Barossa Valley clone, Chardonnay I10V1 for irrigation clone designation). The Australian clone system developed to address the country's unique phytosanitary situation—geographic isolation providing natural barriers to pest introduction combined with strict quarantine measures that historically limited variety access but ensured relatively clean vine material. Australian clone programs emphasize virus elimination and heat tolerance, producing selections suited to the country's warm-climate viticulture and increasingly important for other warm-region production globally. The Australian Wine Research Institute (AWRI) provides industry coordination, research support, and extension services complementing institutional clone programs.
Australian_Clone_Role captures clones from the distinctive Australian viticultural system, where geographic isolation shaped both opportunities (relative freedom from some pests and pathogens) and challenges (limited access to international genetic material until quarantine relaxation). The Australian industry historically relied on locally selected or early-imported material, developing indigenous clone populations (particularly for Shiraz) that differ from European selections. Quarantine reforms since the 1990s have enabled import of international clones (French ENTAV, German Geisenheim), diversifying Australian plantings but also raising concerns about preserving uniquely Australian genetics. Contemporary Australian programs balance clone importation with conservation of traditional selections, recognizing the value of adapted material developed through 200+ years of Australian viticulture.
Australian clone
Australian registered clone
Australian vine selection
australisch registrierter Klon
australische Rebselektion
australischer Klon
Dry PR, Coombe BG, Anderson CJ (2004) Viticulture Volume 1 - Resources, Winetitles CSIRO (2020) Australian vine improvement programs Whiting JR (2004) Grapevine rootstocks, in Viticulture Vol 1, Winetitles pp.: 167-188 Dry N (2007) Grapevine rootstocks, Lythrum Press
Australian Clone Role
Authorization category for Grüner Veltliner in Austria is "Qualitätsrebsorte" (quality wine grape variety) — the highest authorization category under §3 of the Austrian Weingesetz 2009, conferring eligibility for production of Qualitätswein, Kabinett, and all Prädikatswein levels (Spätlese, Auslese, Beerenauslese, Eiswein, Strohwein, Ausbruch, Trockenbeerenauslese), as well as Landwein and Wein ohne nähere Herkunftsbezeichnung, and eligibility for DAC designation in all relevant appellation specifications — the broadest production authorization available under Austrian wine law. Authorization category for Uhudler varieties (Noah, Elvira, Concord, Isabella, and other direct-producer American hybrid cultivars) in Austria documents a historically contentious classification — prohibited for commercial wine production from 1936 until a 1992 derogation permitted limited cultivation in Südburgenland as "Obstwein" (fruit wine) rather than "Wein," and subsequently reclassified under a 2016 amendment permitting limited production as Wein with geographic restriction to Südburgenland — the authorization category reflecting decades of regulatory struggle between European Vitis vinifera purity requirements and regional cultural heritage preservation. Authorization category for Cabernet Sauvignon in France varies by administrative context — classified as "cépage de cuve autorisé" (authorized wine grape variety) at the national level in the Catalogue officiel des variétés de vigne, but its functional category varies by appellation specification from "cépage principal" (principal variety, e.g., in Bordeaux appellations) through "cépage complémentaire" or "cépage accessoire" (complementary/accessory variety, e.g., in certain Languedoc appellations) to effectively excluded (not listed in the specification, e.g., Burgundy and Alsace appellations).
Authorization category is a varietal authorization entity that documents the specific regulatory classification tier assigned to a grape cultivar within the authorization framework of a viticultural jurisdiction, distinguishing among the principal category levels — quality wine variety (Qualitätsrebsorte), land wine variety (Landweinrebsorte), table wine variety (Tafelweinrebsorte), and rootstock variety (Unterlagsrebsorte) in the Austrian system, or the analogous categories of cépage de cuve (wine grape), cépage de table (table grape), cépage porte-greffe (rootstock), and cépage conservatoire (conservation variety) in the French system — each category conferring specific production rights and restrictions that determine the quality tier of wines that may be produced from the cultivar and the administrative conditions governing its cultivation.
Authorization category documentation captures the hierarchical classification system through which viticultural jurisdictions assign differential production rights to cultivars — a system that reflects both the assessed quality potential of each cultivar and the policy objectives of the regulatory framework. The Austrian system provides a particularly clear example of category structure — the Weingesetz 2009 distinguishes between Qualitätsrebsorten (quality wine grape varieties, listed in Annex 1 of the law, eligible for all quality tiers), Landweinrebsorten (land wine grape varieties, eligible for Landwein and basic wine but not for Qualitätswein or Prädikatswein), and Tafeltraubensorten (table grape varieties, authorized for fresh consumption but not for wine production), with a separate classification for Unterlagsrebsorten (rootstock varieties authorized for use as grafting rootstock). The EU framework establishes the overarching category structure — Regulation 1308/2013 requires member states to classify vine varieties into categories determining their eligibility for different wine product categories (wines with PDO, wines with PGI, varietal wines without GI, and wines without varietal or vintage indication), but delegates the specific classification decisions to national authorities. The functional significance of authorization category is most apparent at the boundaries — a cultivar classified as Landweinrebsorte but not Qualitätsrebsorte may produce commercially viable wines that cannot access the quality tier designation, creating an economic ceiling that incentivizes either reclassification efforts (lobbying for promotion to Qualitätsrebsorte status) or alternative marketing strategies (emphasizing producer reputation and wine quality over regulatory classification). The PIWI (fungus-resistant interspecific hybrid) variety debate illustrates the contemporary relevance of authorization category — newly bred disease-resistant varieties face the question of whether to seek Qualitätsrebsorte classification (enabling quality wine production but requiring acceptance of hybrid varieties within the traditional Vitis vinifera quality framework) or to accept a lower authorization category that limits their market positioning despite their agronomic and environmental advantages.
Genehmigungskategorie
Klassifizierungskategorie
Sortenklasse
Zulassungskategorie
authorization class
classification category
cultivar authorization level
regulatorische Stufe
regulatory tier
variety category
Österreichisches Weingesetz (2009) §3 and Annex 1 (Qualitätsrebsorten classification) European Parliament (2013) Regulation (EU) No 1308/2013, Articles 81-85 (vine variety classification) FranceAgriMer (2024) Catalogue officiel des variétés de vigne Bundesamt für Weinbau Klosterneuburg (2023) Klassifizierung der Rebsorten in Österreich
Authorization Category
Authorization date for Grüner Veltliner in Austria documents a long-established regulatory status — inclusion in the Austrian variety catalogue predating the modern Weingesetz (continuous classification as Qualitätsrebsorte through successive Austrian wine law revisions of 1961, 1985, 1999, and 2009), with DAC-specific authorization dates marking the progressive appellation codification: Weinviertel DAC 2002 (the first Austrian DAC, establishing Grüner Veltliner as the mandatory variety), Kremstal DAC 2007, Kamptal DAC 2008, Traisental DAC 2010, and Wachau DAC 2020 — each date representing a regulatory milestone formalizing Grüner Veltliner's regional role. Authorization date for Touriga Nacional in the Bordeaux appellation documents a recent and climate-adaptation-driven authorization — experimental authorization granted by INAO in 2021 as one of six new varieties permitted for trial blending (maximum 5% of the final blend, maximum 10% of vineyard area) within the Bordeaux and Bordeaux Supérieur appellations, with a 10-year evaluation period before potential permanent authorization — the date marking a watershed in French appellation policy, acknowledging climate change as a legitimate basis for varietal innovation within traditional appellations.
Authorization date is a varietal authorization entity that documents the chronological milestones of the legal recognition and classification of grape cultivars within regulatory frameworks, encompassing the date of initial inclusion in the national variety catalogue, the date of authorization for quality wine production, the date of inclusion in specific appellation specifications, the dates of any classification changes (upgrading from authorized to recommended status, or downgrading, temporary authorization, or deauthorization), and the dates of experimental or provisional authorization for trial cultivation, expressed with appropriate precision (exact date for regulatory instruments with defined enactment dates, year for catalogue editions), and providing the temporal framework for understanding the regulatory history of cultivar acceptance that often parallels but sometimes diverges from the cultivar's commercial adoption trajectory.
Authorization date documentation captures the regulatory tempo of cultivar acceptance — the pace at which administrative systems recognize, evaluate, and formally authorize new or previously unclassified cultivars, a process that may span years or decades from initial application to full authorization. The regulatory pathways for cultivar authorization differ substantially across jurisdictions — in France, the INAO process for admitting a new variety to an appellation specification involves experimental cultivation trials (minimum 5 years), technical evaluation by INAO's expert committees, public consultation, and ministerial approval, a process that typically requires 10-15 years from initial proposal to formal authorization. In Austria, the inclusion of a new variety in the Qualitätsrebsorten catalogue requires demonstration of viticultural suitability through multi-year trials at the Höhere Bundeslehranstalt Klosterneuburg or equivalent research institution, followed by recommendation to the Bundesministerium and formal legal amendment through parliamentary or regulatory process. The temporal lag between a cultivar's demonstrated quality potential and its regulatory authorization creates a persistent tension — wines produced from unauthorized cultivars may achieve critical acclaim and commercial success (the Italian "Super Tuscan" phenomenon of the 1970s-1990s, where Cabernet Sauvignon and Merlot wines from Tuscany could only be sold as basic Vino da Tavola because the varieties were not authorized for the applicable DOC regulations) but cannot access the prestige and marketing advantages of appellation designation until regulatory authorization catches up with viticultural practice. The climate change adaptation context is accelerating authorization timelines — the urgency of preparing viticultural regions for warming conditions has prompted several European regulatory bodies to create expedited experimental authorization pathways that allow trial cultivation of new varieties without the full multi-decade authorization process previously required.
Anerkennungsdatum
Genehmigungszeitpunkt
Klassifizierungsdatum
Zulassungsdatum
approval date
authorization timeline
catalogue inclusion date
classification date
regulatorisches Datum
regulatory recognition date
INAO (2021) Cahier des charges modifié de l'appellation d'origine contrôlée Bordeaux (introduction variétés d'adaptation) Österreichisches Weingesetz (2009) §3 and amendments European Parliament (2013) Regulation (EU) No 1308/2013, Articles 81-85 van Leeuwen C et al. (2019) An update on the impact of climate change in viticulture and potential adaptations. Agronomy 9(9): 514
Authorization Date
Authorized region for Grüner Veltliner specifies nationwide authorization across all Austrian wine-growing regions (Weinbauregionen) as listed in the Austrian Weingesetz — Weinland Österreich encompassing Niederösterreich (with sub-regions Wachau, Kremstal, Kamptal, Traisental, Wagram, Weinviertel, Carnuntum, Thermenregion), Burgenland (Neusiedlersee, Leithaberg, Mittelburgenland, Eisenberg, Rosalia), Steiermark (Südsteiermark, Weststeiermark, Vulkanland Steiermark), and Wien — with additional DAC-specific authorization in Weinviertel (requiring 100% Grüner Veltliner for the DAC designation), Kamptal, Kremstal, Traisental, and Wachau DAC regulations. Authorized region for Nebbiolo in Italy documents a geographically restricted authorization concentrated in Piedmont — authorized for DOCG production in Barolo (communes of Barolo, Castiglione Falletto, Serralunga d'Alba, and parts of La Morra, Monforte d'Alba, Novello, Grinzane Cavour, Verduno, Cherasco, Diano d'Alba, Roddi), Barbaresco (communes of Barbaresco, Neive, Treiso, and part of Alba), Roero, Gattinara, Ghemme, and authorized for DOC production in Langhe, Nebbiolo d'Alba, and various Alto Piemonte appellations — the tightly circumscribed authorized regions reflecting the cultivar's demanding site requirements and the regulatory intent to preserve the link between Nebbiolo quality and specific Piedmontese terroirs.
Authorized region is a varietal authorization entity that documents the specific administrative territory or viticultural zone within which a grape cultivar is legally authorized for commercial cultivation and wine production, defined by reference to the applicable jurisdictional boundaries (national territory, federal state/Land/département, wine-growing region, appellation zone, or individual commune), specifying the administrative level at which the authorization operates, the regulatory instrument establishing the authorization (national variety catalogue, appellation specification, ministerial decree), and the relationship between the authorized region and the broader geographic distribution of the cultivar, providing the spatial-jurisdictional dimension of varietal authorization that determines where a cultivar may legally be grown for commercial purposes.
Authorized region documentation reveals the spatial logic of viticultural regulation — the decisions about where specific cultivars may or may not be planted that encode assumptions about cultivar-site compatibility, regional identity, and quality expectations into legal structures. The geographic precision of authorized region designations varies enormously across regulatory systems — from national-level authorization (Austria and most EU member states authorize cultivars across their entire national territory for basic quality wine), through regional delimitation (French appellations define authorized cultivars at the commune or parcel level within appellation boundaries), to the extreme specificity of Italian DOCG regulations that authorize cultivars within precisely defined vineyard zones mapped to cadastral boundaries. The hierarchical structure of authorization regions creates a nested system where the same cultivar may be simultaneously authorized at multiple levels — Grüner Veltliner is authorized nationally in Austria (for Qualitätswein), regionally in Niederösterreich (for Landwein), and appellation-specifically in the Wachau (for DAC wines), with each level imposing progressively more restrictive production conditions. The historical evolution of authorized regions reflects changing viticultural knowledge and policy priorities — regions that were once restricted to traditional cultivars may expand their authorized varieties to accommodate climate adaptation (Bordeaux's recent inclusion of new varieties), while regions that once permitted diverse cultivar portfolios may narrow their authorizations to strengthen regional identity (the progressive tightening of DAC specifications in Austria). The cartographic representation of authorized regions ranges from simple administrative boundary references (the cultivar is authorized throughout the national territory) to detailed vineyard-level maps (the INAO's parcellaire délimité showing which specific plots within an appellation are authorized for PDO production), with the level of cartographic precision generally correlating with the appellation's prestige hierarchy and the strength of the claimed terroir-quality relationship.
Anbaugebiet
Zulassungszone
authorization zone
authorized growing area
authorized territory
cultivation zone
genehmigte Anbauregion
permitted region
zugelassene Region
zugelassenes Gebiet
Österreichisches Weingesetz (2009) §3 and Annex 2 (Weinbauregionen) INAO (2024) Délimitation parcellaire des appellations d'origine contrôlées European Parliament (2013) Regulation (EU) No 1308/2013, Article 93 (definitions of PDO and PGI) Consorzio di Tutela Barolo Barbaresco (2010) Disciplinare di produzione della denominazione di origine controllata e garantita Barolo
Authorized Region
Authorized variety status for Cabernet Sauvignon in Austria classifies the cultivar as Qualitätsrebsorte (authorized for quality wine production nationwide) but without the recommended prominence it holds in Bordeaux — Cabernet Sauvignon serves primarily as a blending component (typically 10-20%) in Burgenland red cuvées and appears as an accessory variety in no Austrian DAC regulation as a standalone varietal designation, reflecting the assessment that while Cabernet Sauvignon produces good wine in Austria's Pannonian climate, it is not considered a primary expression of Austrian viticultural identity. Authorized variety status for Merlot in the Chianti Classico DOCG permits its inclusion at up to 20% of the blend alongside the mandated minimum 80% Sangiovese — authorized as a complementary variety that may enhance the wine but may not define it, reflecting the post-1996 regulation reform that formalized the role of international varieties within Tuscany's indigenous-cultivar-centered appellation system.
Authorized variety status is a classification status entity that documents the standard regulatory classification of a grape cultivar as legally permitted for commercial wine production within a viticultural jurisdiction, without the additional promotional endorsement of recommended status, indicating that the cultivar has been evaluated and approved for cultivation but is not actively promoted or prioritized for the region — typically appearing as a complementary or accessory variety in appellation specifications rather than as a principal variety, and reflecting the assessment that the cultivar produces acceptable wines in the region but is not considered a primary quality or identity vehicle for the regional viticultural sector.
Authorized variety status represents the middle tier of the classification hierarchy — the cultivar is legally permitted and may produce commercially viable wines, but lacks the institutional promotion, research investment, and appellation-level prominence accorded to recommended varieties. The practical distinction between authorized and recommended status is most consequential in the appellation specification context — authorized varieties typically appear as "cépages complémentaires" or "cépages accessoires" in French appellation specifications, permitted for blending within defined proportion limits but not eligible as the principal or sole variety for varietal-labeled appellation wines. The authorized status category encompasses a diverse population of cultivars — well-known international varieties that are permitted but not considered regionally characteristic (Cabernet Sauvignon in Austria, Chardonnay in Piedmont), indigenous varieties of secondary commercial importance (minor local cultivars that contribute to regional diversity without driving regional identity), and newly authorized varieties undergoing market evaluation following recent classification decisions. The boundary between authorized and recommended status is inherently subjective and politically influenced — classification decisions reflect not only agronomic and quality data but also industry lobbying, regional identity politics, and the balance between tradition and innovation that different jurisdictions strike differently. The post-2008 EU simplification nominally eliminated the recommended/authorized distinction at the national level, but in practice the hierarchy persists through appellation-level specification language (principal versus complementary versus accessory variety designations) and through the allocation of institutional research and promotional resources that remain concentrated on the historically prominent cultivars.
Ergänzungssorte
Standardklassifizierung
Zulassungsstatus
accepted variety
authorized classification
erlaubte Rebsorte
permitted variety
secondary variety status
standard classification
zugelassene Sorte
European Parliament (2013) Regulation (EU) No 1308/2013, Articles 81-85 INAO (2024) Cépages autorisés et accessoires par appellation Consorzio Vino Chianti Classico (2014) Disciplinare di produzione DOCG Chianti Classico Österreichisches Weingesetz (2009) §3
Authorized Variety Status
Grüner Veltliner in Austria bearing an autochthonous cultivar role has been present in the Danube region since at least the 18th century with parentage analysis revealing Traminer × St. Georgen as parents — indicating non-Austrian genetic origins — yet centuries of cultivation across Austrian terroirs from the Wachau to the Weinviertel have produced extensive clonal diversity with over 20 registered Austrian clones showing measurable adaptation to local conditions including calcareous loess soils and Pannonian climate, making the cultivar culturally and ampelographically inseparable from Austrian viticultural identity.
A traditional cultivar role that inheres in a grapevine cultivar that has been cultivated within a specific region for a sufficiently long period — conventionally several centuries — to have undergone significant local adaptation through clonal mutation, natural selection pressure, and viticultural co-evolution, regardless of whether the cultivar's ultimate geographic origin lies within or outside the region.
Autochthonous_Cultivar_Role is conceptually distinct from Indigenous_Cultivar_Role in that it emphasizes long-term regional adaptation and cultural embedding rather than original geographic origin. Many of Europe's most important cultivars are autochthonous without being indigenous: Merlot is autochthonous to Bordeaux (present since at least the 18th century, extensively locally adapted) but genetic parentage analysis reveals Cabernet Franc × Magdeleine Noire des Charentes parentage that does not confirm indigenous Bordelais origin. The distinction is viticulturally significant because autochthonous cultivars typically exhibit superior terroir expression through accumulated clonal adaptation regardless of ultimate provenance.
autochthone Sorten-Rolle
autochthonous variety role
locally adapted cultivar role
regional adaptierte Sorten-Rolle
regionally evolved cultivar role
standortangepasste Rebsorten-Rolle
Lacombe T et al. (2013) Large-scale parentage analysis in an extended set of grapevine cultivars, Theoretical and Applied Genetics 126: 401-414 Maul E et al. (2012) Identification and characterization of grapevine genetic resources, Vitis 51: 53-63 Cattonaro F et al. (2014) Genetic diversity in the grapevine germplasm, BMC Plant Biology 14: 99
Autochthonous Cultivar Role
The autumn color description of Pinot Noir documents intense red to dark red-violet senescence coloration (OIV 093:5-6) — as chlorophyll degrades during the 3-4 week senescence period following harvest, the previously green leaves transition through yellow-green to a uniform deep crimson-red produced by intense de novo anthocyanin synthesis in the mesophyll and epidermis, creating the famously spectacular autumn canopy that characterizes Pinot Noir vineyards from Burgundy to Oregon to Central Otago. This strong red autumn color distinguishes Pinot Noir from its offspring Chardonnay (OIV 093:1, purely yellow senescence with no anthocyanin production) and the half-sibling Gamay Noir (OIV 093:4-5, red-orange autumn color typically less intensely pigmented than Pinot Noir), and is maintained even in the white-berry sport Pinot Blanc, confirming that the senescence-associated anthocyanin regulation is independent of the berry skin color pathway controlled by VvMYBA1.
A leaf color description component that specifically records the characteristic coloration of senescent grapevine leaves observed 2-4 weeks before natural leaf fall, scored according to OIV descriptor 093 on a scale from yellow (state 1) through yellow with red spots (state 2), orange (state 3), red-orange (state 4), red (state 5), to dark red-violet (state 6), reflecting the cultivar-specific balance between carotenoid unmasking and de novo anthocyanin biosynthesis during the senescence process that produces the spectacular autumn foliage displays of vineyard landscapes.
Autumn_Color_Description documents a phenological character that provides cultivar identification information at a time of year (post-harvest, pre-dormancy) when most other ampelographic characters have either been removed (clusters harvested), obscured (leaf damage from disease, drought, or hail accumulated over the season), or are no longer distinctive (mature leaves losing their diagnostic morphological crispness through late-season tissue degradation). The autumn color is determined by the cultivar-specific activation of anthocyanin biosynthesis during senescence — a process genetically distinct from the constitutive anthocyanin production controlling berry skin color and shoot tip pigmentation: the regulatory mechanisms involve senescence-associated transcription factors that activate the anthocyanin pathway in leaf tissue as part of the broader senescence program, with the intensity of activation varying from absent (cultivars producing purely yellow autumn foliage through carotenoid unmasking alone) to very strong (cultivars producing deep red-violet foliage through massive anthocyanin accumulation). The autumn color display has cultural significance beyond its ampelographic utility — the red and gold autumn vineyards of Burgundy, Champagne, the Mosel, and other iconic wine regions are important components of viticultural landscape aesthetics and cultural heritage tourism.
Herbstfärbungs-Beschreibung
Seneszenz-Blattfärbung
autumn leaf pigmentation
fall foliage coloration
herbstliche Laubfarbe
senescence color record
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Liang Z et al. (2019) Whole-genome resequencing of 472 Vitis accessions, Nature Communications 10: 1190
Autumn Color Description
Autumn frost damage quality assessment in Finger Lakes Riesling following an early October frost event (-5°C, 4 weeks before normal first frost date) documented impacts on unharvested fruit and vine preparation for winter. Fruit remaining on vines showed moderate autumn frost damage quality with 35% of clusters exhibiting visible ice damage including berry splitting and juice leakage, with affected fruit producing wines showing oxidized character and reduced varietal typicity attributed to cellular damage and enzyme release during freezing. Rapid harvest of remaining fruit within 48 hours of the frost event salvaged usable production but at reduced quality compared to the controlled harvest timing planned for optimal ripeness. Foliage assessment revealed severe autumn frost damage quality with 80% defoliation within one week of the frost event, eliminating the 3-4 weeks of additional photosynthesis normally occurring before natural leaf fall. Subsequent midwinter hardiness testing showed affected vines achieving maximum bud hardiness of only -22°C compared to -26°C in vines with normal autumn progression, attributing the 4°C hardiness deficit to the disrupted acclimation caused by premature defoliation and reduced carbohydrate reserves. The autumn frost damage quality assessment established carry-over effects including 15% reduction in fruitfulness and delayed budburst in the following spring, demonstrating the multi-season consequences of early autumn frost beyond immediate damage.
Autumn frost damage quality is a frost damage severity quality that inheres in a grapevine organism and represents the extent of injury to leaves, fruit, and incompletely hardened woody tissues from early autumn freezing events occurring before adequate cold acclimation has developed, with damage affecting the completion of fruit ripening, the accumulation of carbohydrate reserves for winter survival and spring growth, and the progression of cold hardening that determines midwinter freeze tolerance. This quality addresses the impacts of frost occurring during the transition from active growth to dormancy, recognizing that early autumn frost can disrupt the normal sequence of senescence, reserve accumulation, and acclimation that prepares vines for winter, with consequences extending beyond the immediate damage to affect winter survival and subsequent season performance. The ontological significance of autumn frost damage quality lies in its effects on both current and subsequent season production, with fruit damage directly affecting current harvest while foliage loss and acclimation disruption affect carbohydrate storage and hardiness development with consequences manifesting in reduced winter hardiness, delayed spring growth, and diminished productivity in the following season. The biological basis of autumn frost damage encompasses the intermediate cold sensitivity of tissues in transition between summer growth and winter dormancy, with leaves remaining frost-sensitive throughout their functional life, fruit showing variable sensitivity depending on ripeness and sugar concentration providing cryoprotection, and woody tissues showing progressively developing hardiness that may be insufficient to survive early autumn temperatures substantially below 0°C. The timing of autumn frost relative to harvest determines fruit damage significance, with pre-harvest frost potentially causing berry freezing that compromises quality through cellular damage, off-flavor development, and physical damage complicating processing, while post-harvest frost affects only vegetative tissues with implications limited to reserve accumulation and acclimation progression. The premature leaf loss from autumn frost damage reduces photosynthetic capacity during the critical reserve accumulation period when carbohydrates are being stored in permanent vine structures for winter respiration and spring growth support.
Autumn_Frost_Damage_Quality is_a Frost_Damage_Severity_Quality, representing frost injury during the growth-to-dormancy transition before full hardiness development. This class addresses the dual impacts on current harvest and subsequent season preparation. Fruit damage depends on sugar concentration providing cryoprotection (ripe fruit more tolerant). Premature defoliation affects carbohydrate reserves and hardiness development. The carry-over effects on winter hardiness and following-season productivity distinguish autumn frost from isolated damage events. Timing relative to harvest determines fruit versus foliage damage significance.
Frühfrostschaden
Herbstfrostschaden
Herbstgefrierschaden
Vordormanz-Frostverletzung
autumn freeze damage
early frost damage
fall frost injury
pre-dormancy frost injury
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 508-545 Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection. Michigan State University Extension Bulletin E2930
Autumn Frost Damage Quality
The AVA designation Napa Valley (established 1981, TTB regulation 27 CFR Part 9.23) delimits approximately 45,000 acres in Napa County, California, within which 16 nested sub-AVAs (including Rutherford, Oakville, Stags Leap District, and Howell Mountain) provide additional geographic specificity — the AVA label claim requires that at least 85% of grapes originate within the designated area but imposes no restrictions on grape varieties, yields, winemaking methods, or aging requirements, fundamentally distinguishing the AVA system from European PDO/PGI frameworks that integrate geographic delimitation with production rule specification.
A regional wine designation within the United States wine regulatory framework that identifies a wine as containing at least 85% of grapes grown within an American Viticultural Area (AVA) — a federally delimited grape-growing region distinguished by specific geographic features including climate, soil, elevation, and physical features — administered by the Alcohol and Tobacco Tax and Trade Bureau (TTB) without specifying production methods, grape varieties, or quality standards.
AVA_Designation captures the distinctive American approach to geographic indication that intentionally separates geographic delimitation from production regulation. The TTB establishes AVA boundaries based exclusively on distinguishing geographic features (climate, soil, elevation, topography) through a petition-based process requiring scientific and historical evidence of the area's viticultural distinctiveness: unlike European appellations, AVAs carry no production specifications — any grape variety may be planted, any yield achieved, any winemaking method employed, and no quality examination is required. This philosophical difference reflects the American regulatory tradition's emphasis on geographic truthfulness (the wine comes from where the label says it does) rather than typicity enforcement (the wine tastes like a wine from that place should taste). The United States currently has approximately 270 established AVAs, with California containing the majority.
AVA
AVA-Bezeichnung
American Viticultural Area designation
US viticultural area classification
US-Anbaugebiets-Kennung
amerikanische Weinbaugebiet-Klassifikation
TTB (2023) American Viticultural Areas, 27 CFR Part 9 Federal Alcohol Administration Act, 27 USC §205 Mendelson R (2011) Wine in America: Law and Policy, Wolters Kluwer pp: 145-210
AVA Designation
Average yield for Sauvignon Blanc clone 242 (ENTAV) is documented at 10.5 t/ha (SD ±2.1, CV 20%) across 12 vintages at Domaine de l'Espiguette, compared with 12.8 t/ha (SD ±2.5, CV 20%) for clone 297 and 8.2 t/ha (SD ±1.6, CV 19%) for clone 108, establishing the yield ranking used in regional planting recommendations. Average yield for Blaufränkisch clone Bl-2 is recorded at 7.5 t/ha (range 5.8-9.2 t/ha) across 10 vintages at the Klosterneuburg Agneshof trial site on Kober 5BB rootstock at 2.4 m × 1.0 m spacing, representing a 12% reduction relative to unselected population material but with 1.5° KMW higher average must weight.
Average yield is a yield performance data entity that records the arithmetic mean fruit production of a clonal line calculated across multiple vintages and, where applicable, multiple trial sites, expressed in tonnes per hectare (t/ha), kilograms per vine (kg/vine), or kilograms per meter of cordon (kg/m), accompanied by measures of dispersion (standard deviation, coefficient of variation, range), providing the central tendency estimate of clone productivity under the documented evaluation conditions.
Average yield reporting requires explicit documentation of the conditions under which the average was calculated, as the metric is meaningless without reference to vine age range, training system, spacing, rootstock, irrigation status, and crop load management interventions applied during the evaluation period. The minimum vintage count for a statistically reliable yield average is generally considered 8-10 years, capturing sufficient climatic variation to represent the range of growing conditions a clone will encounter in commercial production. Juvenile non-bearing years (typically years 1-3 after planting) and exceptional climatic events (severe frost, hail) are conventionally excluded from average yield calculations, though their treatment should be documented. The coefficient of variation (CV) accompanying the average yield provides critical information about production reliability — CV values below 15% indicate stable production, 15-25% is typical, and values exceeding 30% suggest problematic yield instability that may reflect clone-specific sensitivity to environmental fluctuation. Cross-site yield averages should be presented alongside site-specific values, as averaging across environments can obscure meaningful clone-by-site interactions.
Durchschnittsernte
Durchschnittsertrag
Ertragserwartung
average harvest weight
average production
langjähriger Mittelertrag
long-term yield average
mean crop level
mean yield
mittlerer Ertrag
Anderson MM et al. (2008) Viticultural evaluation of French and California Pinot Noir clones. American Journal of Enology and Viticulture 59(2): 188-193 Wolpert JA et al. (1994) Field performance of six Chardonnay clones. American Journal of Enology and Viticulture 45(4): 393-400 Boidron R et al. (1995) Catalogue des variétés et clones de vigne cultivés en France, ENTAV-INRA, pp.: 30-45
Average Yield
An axillary primary vegetative bud at node 3 on a cane of Vitis vinifera cv. Merlot in a bilateral cordon training system, positioned at the leaf scar from the previous season's third leaf, dissected revealing vegetative primordia with 9 leaf initials and 2 tendril primordia but no inflorescence structures, reflecting typical basal node fruitfulness patterns where position 1-3 buds receive less than 15% full sunlight during the June-July differentiation period due to shading from expanding apical canopy, compared to nodes 6-9 that receive 65-85% full sun and consistently develop reproductive primordia.
A primary vegetative bud located in the axil (leaf-stem junction) at lateral positions along the main cane axis, representing the standard node position for primary bud formation in grapevine architecture, characterized by axillary placement opposite leaf attachment points (or leaf scars on dormant wood), typical primary bud developmental patterns, and variable fruitfulness depending on specific node number, light exposure during differentiation, and cultivar-specific fruitfulness patterns. Axillary primary vegetative buds at basal positions (nodes 1-3) typically develop under heavy shade from expanding canopy during critical inflorescence initiation period, resulting in predominantly vegetative differentiation even when vine balance and nutrition are optimal, representing the normal architectural expression of reduced basal fruitfulness that influences pruning strategies across all training systems.
The term "axillary" in this context refers to standard architectural positioning rather than implying any special developmental pathway, as essentially all primary buds in grapevines form in axillary positions. The distinction between axillary primary vegetative versus reproductive buds lies in the primordial content (vegetative-only versus including reproductive structures) rather than positional differences. Axillary positioning ensures vascular connection through the leaf trace and positions buds for optimal breakage angles in spring, with the axillary position being highly conserved across Vitis species and cultivars as the fundamental pattern for bud placement along shoots.
Blattachselknospe
axilläre vegetative Primärknospe
lateral primary vegetative bud
node-position primary vegetative bud
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Morrison (1991) "Bud development in Vitis= Anatomical study of formation and growth of the primary grapevine bud during one annual cycle" Australian Journal of Grape and Wine Research 72=296-301 Carmona et al. (2008) "Grapevine bud dormancy= environmental and molecular aspects" Revista Brasileira de Fruticultura 30=257-408.
Axillary Primary Vegetative Bud
A BC4 Riesling derivative carrying Rpv3 downy mildew resistance bears a Backcross_Hybrid_Role, with four generations of backcrossing to Riesling having recovered >96% Riesling genetics while retaining the Rpv3 R-gene from the original interspecific donor—approaching "disease-resistant Riesling" status. A BC2 Chardonnay line carrying Run1 powdery mildew resistance serves in a Backcross_Hybrid_Role as an intermediate breeding step, with additional backcross generations planned to further improve Chardonnay character before variety registration. The advanced French breeding line VRH3082-1-42 bears a Backcross_Hybrid_Role as a Muscadinia-derived powdery mildew resistant genotype backcrossed into V. vinifera background, the BC generations progressively recovering vinifera wine quality while maintaining Run1 resistance.
A hybrid role that inheres in a grapevine genotype produced through backcrossing—crossing a hybrid individual back to one of its parents or to a genotype genetically similar to one parent—as part of a breeding strategy to recover the genetic background of the recurrent parent while retaining specific traits from the donor parent. Backcross hybrids carry progressively larger proportions of recurrent parent genome with each backcross generation: BC1 individuals average 75% recurrent parent DNA, BC2 average 87.5%, BC3 average 93.75%, and BC4 average 96.875%, with marker-assisted selection potentially accelerating background recovery beyond these theoretical expectations. In grapevine resistance breeding, backcrossing is the primary strategy for introgressing disease resistance genes into elite V. vinifera varieties—crossing a resistance donor with the target variety (F1), then repeatedly crossing resistant progeny back to the recurrent parent while selecting for resistance at each generation. The backcross strategy acknowledges that variety identity is commercially valuable rather than creating entirely new varieties, backcross breeding aims to create "improved versions" of established varieties (e.g., "disease-resistant Riesling") maintaining most of the recurrent parent's characteristics while adding the introgressed trait.
Backcross_Hybrid_Role captures the progressive genome recovery strategy that enables trait introgression into established variety backgrounds. The BC designation indicates position in the backcross series (BC1, BC2, BC3...), with higher numbers representing greater recurrent parent recovery. Marker-assisted backcrossing (MABC) has transformed this approach by enabling background selection (selecting for recurrent parent alleles at non-target loci) alongside foreground selection (confirming target trait presence), accelerating background recovery beyond theoretical expectations from phenotypic selection alone. The backcross strategy is particularly suited to grapevine where variety identity has strong market value—producing "resistant Riesling" maintains the established variety name and market position while adding the introgressed trait. Backcross products approaching commercial release may be designated with the recurrent parent name plus resistance trait identifier (e.g., "Riesling R" for resistant Riesling derivative) or with new variety names depending on regulatory and marketing decisions.
BC hybrid
BC-Hybride
Introgressionprodukt
Rückkreuzungsgeneration
Rückkreuzungshybride
backcross generation
backcross hybrid
introgression product
Töpfer R et al. (2011) Molecular breeding in grapes, CRC Press pp.: 160-185 Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330 Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance, Molecular Genetics and Genomics 291: 1573-1594
Backcross Hybrid Role
A backup reserve renewal cane on Vitis vinifera cv. Pinot Noir in Burgundian Guyot system, retained at 9 nodes positioned 22cm from cordon origin (primary at 8cm, secondary at 15cm), measuring 8mm diameter showing complete lignification, moderate node spacing indicating balanced growth, representing routine precautionary retention in vineyard with 20-year history showing approximately 18% annual utilization of backup reserves due to variable winter damage and spring growth patterns in cool continental climate, with pruner's expectation being probable removal if primary and secondary perform well but ready utilization if preferred options show problems, maintaining architectural integrity and avoiding compromised fruiting wood selections under pressure.
A reserve renewal cane retained as standard precautionary measure in well-managed vineyards, characterized by moderate expectation of actual utilization (perhaps 15-25% selection probability), acceptable positioning and quality making it viable though not preferred fruiting wood option, and routine retention as insurance against common renewal problems including winter damage variability, spring growth inconsistencies, or minor training adjustments requiring alternative wood sources. Backup reserve renewal canes represent pragmatic risk management in renewal planning, acknowledging that some percentage of preferred renewals will prove unsuitable each season due to routine rather than catastrophic problems, with backup reserves providing ready alternatives avoiding crisis selections or architectural compromises when primary and secondary options disappoint during assessment at following dormant season.
Backup reserve renewal canes represent the boundary between standard renewal planning (primaries and secondaries) and contingency planning (emergency reserves), with backup reserves straddling the divide through moderate utilization probability reflecting routine rather than exceptional problems. Historical tracking in well-documented vineyards shows backup reserve utilization varying from 10-30% annually depending on climate stability, vine health, and winter severity, with this moderate frequency justifying routine retention despite bud capacity cost. The backup reserve strategy suits quality-focused vineyards where compromised fruiting wood (forced selections from latent buds or poorly positioned wood due to inadequate renewal planning) threatens fruit quality and wine character, making the insurance value of backup reserves exceed their opportunity cost.
Standard-Reserverute
precautionary renewal wood
routine backup cane
standard reserve renewal
vorsorgliche Vorratsrute
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Reynolds & Vanden Heuvel (2009) "Influence of grapevine training systems on vine growth and fruit composition= a review" American Journal of Enology and Viticulture 60=251-268.
Backup Reserve Renewal Cane
A backup secondary reproductive shoot on Vitis vinifera cv. Merlot from node 7, activated following early hail damage eliminating primary shoots, developed to 108cm length by late July, 14 leaves totaling approximately 1890 cm² leaf area, bearing 2 moderate clusters with 42 and 38 berries, estimated total shoot crop 285g representing approximately 58% of typical primary production from this position (primaries normally producing 480-520g), representing robust secondary compensation attributed to combination of Merlot's relatively good secondary fruitfulness, favorable warm growing season following damage supporting vigorous secondary development, and appropriate remaining whole-vine crop load enabling adequate resource allocation to developing secondary shoots and clusters, managed through standard canopy operations with recognition of delayed phenology (projected harvest 8-10 days later than normal) but acceptable fruit quality prospects based on adequate compensation and favorable seasonal progression.
A secondary reproductive shoot exhibiting relatively good reproductive performance despite secondary bud origin, characterized by prompt secondary activation, reasonable vigor achieving 90-120cm length, adequate cluster development with 1-2 inflorescences showing acceptable berry numbers (35-60 berries per cluster), and crop contribution approaching 50-70% of typical primary shoot production from equivalent positions, representing upper performance range for secondary reproductive capacity under favorable compensation conditions. Backup secondary reproductive shoots demonstrate maximal reproductive compensation from backup positions, providing substantial crop recovery following primary loss, contributing meaningfully to vineyard economic viability in damage scenarios, and occasionally producing fruit quality approaching primary standards when favorable growing conditions and appropriate crop management support adequate maturation despite phenological delays and reduced initial vigor.
Backup secondary reproductive shoots represent best-case compensation outcomes, achieving 50-70% of primary productivity and maintaining acceptable fruit quality potential through favorable combinations of variety characteristics (some cultivars showing better secondary fruitfulness than others), growing conditions (warm seasons supporting rapid secondary development), crop load balance (partial primary survival or aggressive thinning preventing excessive demand on recovering vines), and management skill (appropriate decisions supporting secondary development). The factors enabling strong backup secondary performance include inherent variety traits (Chardonnay, Merlot, Riesling generally showing better secondary fruitfulness than Cabernet Sauvignon, Syrah, or Nebbiolo), favorable post-damage weather (warm temperatures, adequate moisture supporting rapid growth and development), appropriate vine resource status (adequate reserves, balanced previous-season crop), and skilled management recognizing secondary limitations while maximizing their potential through appropriate interventions. Backup secondaries partially mitigate damage economic impacts, though cannot fully compensate for primary losses even under optimal circumstances.
high-performance backup shoot
kompensierender sekundärer Fruchttrieb
leistungsfähiger Ersatztrieb
productive secondary shoot
Based on Molitor et al. (2014) "Late frost damage risk for viticulture under future climate conditions= a case study for the Luxembourgish winegrowing region" Australian Journal of Grape and Wine Research 20=160-168 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115.
Backup Secondary Reproductive Shoot
A backup secondary vegetative bud at node 8 on Vitis vinifera cv. Syrah, remaining dormant during normal spring budbreak when the primary bud successfully emerged and developed, then activating 16 days post-budbreak following late spring frost at -2.8°C that killed 78% of primary shoots at 3-5 leaf stage, this backup secondary producing vigorous compensatory vegetative growth reaching 145cm shoot length with 19 leaves by August, contributing estimated 1180 cm² photosynthetic leaf area that partially offset the 2.3 kg yield loss from destroyed primary fruiting shoot, while establishing well-differentiated buds for partial crop recovery in the following season.
A secondary vegetative bud functioning in its primary evolutionary role as reserve meristem, remaining quiescent under normal growing conditions when the primary bud develops successfully but capable of rapid activation to produce replacement vegetative growth when primary buds are killed or severely damaged by environmental stress, pests, or pathogens. Backup secondary vegetative buds represent the insurance function of compound bud architecture, with their presence ensuring that every node position retains capacity for shoot production even following catastrophic primary bud loss, thereby maintaining minimum canopy development necessary for vine survival and photosynthetic function, though sacrificing reproductive output in the damage year while establishing foundation for recovery in subsequent seasons.
Backup secondary vegetative buds fulfill the fundamental survival function of compound bud architecture, ensuring that grapevines can recover from severe damage events that would be catastrophic for species lacking developmental redundancy. Their activation represents a shift in vine resource allocation from reproduction to survival and recovery, with current season yield sacrificed to maintain photosynthetic capacity, build carbohydrate reserves, and differentiate buds for future production. The vigor of backup secondary shoots often exceeds normal primary shoots due to reduced competition (many nodes lacking shoots following widespread primary damage) and high root-to-shoot ratios in under-cropped vines. Management of backup secondary growth focuses on shoot positioning, disease control in dense regrowth, and decisions regarding retention versus removal of shoots based on vine spacing, training system capacity, and recovery strategy.
Notfall-Nebenknospe
Reserve-Sekundärknospe
emergency vegetative bud
insurance secondary bud
reserve secondary vegetative bud
Based on Trought & Bramley (2011) "Vineyard variability in Marlborough, New Zealand= characterising spatial and temporal changes in fruit composition and juice quality in the vineyard" Australian Journal of Grape and Wine Research 17=79-89 Ferguson et al. (2011) "Dynamic changes in bud cold hardiness during dormancy and the influence of apple and pear budbreak" Journal of the American Society for Horticultural Science 136=313-323 Molitor et al. (2014) "Late frost damage risk for viticulture under future climate conditions= a case study for the Luxembourgish winegrowing region" Australian Journal of Grape and Wine Research 20=160-168.
Backup Secondary Vegetative Bud
Bacterial infection indicator quality assessment in a California vineyard exhibiting vine decline symptoms required differential diagnosis between Pierce's disease and other causes of leaf scorch and dieback. Initial bacterial infection indicator quality observations noted the characteristic marginal leaf necrosis with adjacent green tissue creating "green islands" surrounded by scorched margins, the "matchstick" petioles remaining attached after leaf blade abscission, and the progressive vine decline affecting some vines while adjacent vines remained healthy, all consistent with Pierce's disease rather than abiotic scorch from drought or salt stress. Laboratory confirmation through ELISA and PCR testing for Xylella fastidiosa validated the field diagnosis based on bacterial infection indicator quality, with positive detection in symptomatic vines and negative results from adjacent asymptomatic vines. Separate bacterial infection indicator quality assessment identified crown gall tumors at the trunk base of young replants in the same vineyard, with the characteristic fleshy to corky gall tissue at wound sites indicating Agrobacterium vitis infection, distinct from the vascular symptoms of Pierce's disease. The bacterial infection indicator quality assessment established that both major bacterial pathogens were present in the vineyard, requiring integrated management combining roguing of Pierce's disease-affected vines with vector management and careful wound protection during planting and training to limit crown gall establishment in replacement vines.
Bacterial infection indicator quality is a pathogen infection indicator quality that inheres in a grapevine organism and represents the ensemble of symptoms and diagnostic signs specifically associated with bacterial pathogen colonization, including the characteristic galls, cankers, leaf scorching patterns, and systemic decline symptoms produced by the bacterial species that cause significant disease in grapevine, with bacterial diseases differing from fungal diseases in their generally systemic nature, the absence of visible sporulating structures, and the limited chemical control options necessitating integrated management approaches emphasizing prevention and cultural practices. This quality captures the plant responses and diagnostic features that enable recognition of bacterial diseases, recognizing that while fewer bacterial species cause economically important grapevine diseases compared to fungi, the bacterial diseases that do occur are often severe, difficult to control, and may require vine removal or replanting as the only effective management option. The ontological significance of bacterial infection indicator quality lies in its role enabling diagnosis of diseases with limited treatment options, where early detection may enable removal of affected vines before pathogen spread rather than therapeutic intervention, and where accurate diagnosis distinguishes bacterial from fungal or abiotic causes of similar-appearing symptoms to avoid inappropriate and ineffective management responses. The biological basis of bacterial infection indicators encompasses the colonization of host tissues by bacterial cells multiplying in intercellular spaces or vascular elements, the production of phytotoxins, cell-wall-degrading enzymes, or growth regulators that cause visible symptoms, and in the case of Agrobacterium the genetic transformation of host cells producing the tumorous overgrowths characteristic of crown gall. The two most economically significant bacterial diseases of grapevine—crown gall caused by Agrobacterium vitis and Pierce's disease caused by Xylella fastidiosa—show distinctly different symptom types reflecting their different colonization strategies, with crown gall producing localized tumors at wound sites while Pierce's disease causes systemic vascular dysfunction manifesting as progressive leaf scorch and vine decline.
Bacterial_Infection_Indicator_Quality is_a Pathogen_Infection_Indicator_Quality, serving as parent class for Crown_Gall_Symptom_Quality_(Agrobacterium_vitis) and Pierces_Disease_Symptom_Quality_(Xylella_fastidiosa) which address the two most important bacterial diseases of grapevine. Bacterial diseases differ from fungal diseases in systemic nature and absence of sporulating structures. Limited chemical control options make prevention and cultural management primary strategies. Crown gall (localized tumors) and Pierce's disease (systemic vascular) represent contrasting bacterial disease types.
Bakterieninfektions-Indikatoren
Bakteriose-Zeichen
bacterial disease symptoms
bacterial pathogen signs
bacteriosis symptoms
bakterielle Krankheitssymptome
prokaryotic infection indicators
prokaryotische Infektionsmarker
Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests, 2nd Edition. APS Press Burr TJ, Otten L (1999) Crown gall of grape: biology and disease management. Annual Review of Phytopathology 37:53-80 Hopkins DL, Purcell AH (2002) Xylella fastidiosa: cause of Pierce's disease of grapevine and other emergent diseases. Plant Disease 86:1056-1066
Bacterial Infection Indicator Quality
Comparative evaluation of Bacterial_Resistance_Disposition at UC Davis assessed responses to both Agrobacterium vitis and Xylella fastidiosa across 30 Vitis accessions to characterize the independence of resistance mechanisms against these distinct bacterial pathogens. For crown gall, wound inoculation with tumorigenic A. vitis strain K306 resulted in gall formation on 100% of V. vinifera cultivars tested, with gall volumes of 8-25 cm³ at 12 weeks. Wild species V. amurensis and V. riparia showed similarly high susceptibility, indicating no differential Bacterial_Resistance_Disposition against Agrobacterium across tested germplasm. In contrast, Pierce's disease challenge through mechanical inoculation of X. fastidiosa strain Temecula1 revealed dramatic variation in Bacterial_Resistance_Disposition. V. vinifera 'Chardonnay' developed severe symptoms with bacterial titers of 10⁸ CFU/g petiole and 100% mortality by 16 weeks. V. arizonica b43-17 showed strong resistance with titers below detection limits and no symptoms. Critically, crown gall susceptible V. arizonica exhibited Pierce's disease resistance, while Pierce's disease susceptible V. vinifera showed equivalent crown gall susceptibility, demonstrating that Bacterial_Resistance_Disposition is pathogen-specific rather than reflecting general antibacterial capacity.
A Bacterial_Resistance_Disposition is a Pathogen_Resistance_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through defense processes that limit infection, colonization, multiplication, or pathogenesis by bacterial pathogens affecting grapevine. This disposition represents the capacity to resist the relatively limited but economically significant bacterial diseases of grapevine, which differ fundamentally from fungal and viral pathogens in their biology, infection mechanisms, and host-pathogen interactions. Major bacterial pathogens of grapevine include Agrobacterium vitis (syn. Allorhizobium vitis) causing crown gall, Xylella fastidiosa causing Pierce's disease and related leaf scorch syndromes, and Xanthomonas ampelina causing bacterial blight. Bacterial resistance mechanisms in plants encompass recognition of pathogen-associated molecular patterns (PAMPs) such as flagellin and lipopolysaccharides triggering PAMP-triggered immunity (PTI), effector-triggered immunity (ETI) mediated by resistance proteins recognizing bacterial type III secretion effectors, physical barriers preventing bacterial entry or spread, antimicrobial compound production, and restriction of bacterial multiplication in host tissues. For grapevine specifically, bacterial resistance dispositions vary dramatically by pathogen system, with no effective resistance available against crown gall in V. vinifera, while strong resistance to Pierce's disease exists in wild North American Vitis species that co-evolved with X. fastidiosa. The distinct epidemiology and tissue tropism of bacterial pathogens (wound-mediated infection for Agrobacterium, xylem-limited colonization for Xylella, foliar infection for Xanthomonas) require different resistance mechanisms, making bacterial resistance a heterogeneous category encompassing diverse host defense strategies.
Bacterial_Resistance_Disposition is_a Pathogen_Resistance_Disposition. This class encompasses resistance to all bacterial pathogens of grapevine, parallel to Fungal_Resistance_Disposition and Viral_Resistance_Disposition. Child classes include Crown_Gall_Resistance_Disposition (Agrobacterium) and Pierces_Disease_Resistance_Disposition (Xylella fastidiosa). Resistance mechanisms differ fundamentally between these pathogens due to their distinct infection biology: wound-mediated transformation for Agrobacterium versus insect-vectored xylem colonization for Xylella. No resistance to bacterial blight (Xanthomonas ampelina) is currently modeled in VVO due to limited breeding relevance.
Bakterielle-Krankheitsresistenz
Bakterienresistenz-Disposition
Prokaryoten-Pathogen-Resistenz
bacterial disease resistance
bacterial infection immunity
prokaryotic pathogen resistance
Burr TJ, Otten L (1999) Crown gall of grape: biology and disease management, Annual Review of Phytopathology 37:53-80 Hopkins DL, Purcell AH (2002) Xylella fastidiosa: cause of Pierce's disease of grapevine and other emergent diseases, Plant Disease 86:1056-1066 Jones JDG, Dangl JL (2006) The plant immune system, Nature 444:323-329
Bacterial Resistance Disposition
A phenological observer recording Baggiolini A for Merlot in Ticino on February 15 documents the vine in full winter dormancy with buds tightly appressed to the cane, brown and pointed, showing no external morphological change — the vine has accumulated approximately 800-1200 chilling hours below 7.2°C since leaf fall, endodormancy has been satisfied, but ecodormancy persists as ambient temperatures remain below the 10°C base threshold for growth resumption, with Baggiolini A representing the baseline reference state against which all subsequent developmental progression is measured.
A Baggiolini code designating stage A that describes the grapevine in full winter dormancy with buds tightly closed, brown, and covered by protective bud scales, showing no visible signs of swelling or growth activity, corresponding approximately to BBCH 00.
Baggiolini_A_Winter_Bud defines the starting point of the annual phenological cycle in the Baggiolini system. The stage corresponds to BBCH 00 (dormancy, winter buds pointed and closed) and encompasses the period from the completion of leaf fall through the end of winter rest, during which the vine exhibits no externally visible developmental change despite ongoing internal physiological processes including chill accumulation for endodormancy release and starch-sugar interconversion in wood tissues: the transition from Baggiolini A to B is triggered when rising spring temperatures exceed the base growth threshold for a sufficient duration to initiate visible bud swelling, typically requiring accumulation of 50-100 growing degree days (base 10°C) after endodormancy satisfaction.
Baggiolini stage A
Baggiolini-Stadium A
Ruhestadium
Winterknospenstadium
dormant bud stage
winter dormancy stage
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 73-80
Baggiolini A Winter Bud
A phenological record of Baggiolini B for Pinot Noir in Champagne on March 25 documents that winter buds have visibly swollen from their dormant pointed form to a rounded profile approximately 1.5x their winter diameter, with bud scales separating slightly at the margins but no wool or green tissue yet visible — the transition from A to B typically occurs 7-14 days before bud burst under cool spring conditions, and the observation of Baggiolini B triggers early-season vineyard preparations including pruning wound assessment, trellis repair completion, and first pre-emergence herbicide applications in conventionally managed vineyards.
A Baggiolini code designating stage B that describes the visible swelling and rounding of grapevine buds as they absorb water and internal tissues expand against the enclosing bud scales, marking the first externally detectable sign of growth resumption after winter dormancy, corresponding approximately to BBCH 01-03.
Baggiolini_B_Bud_Swelling marks the phenological transition from dormancy to active growth and corresponds to the BBCH 01-03 range (beginning of bud swelling through end of bud swelling). The visible swelling results from water uptake and starch hydrolysis within the bud, driven by rising xylem sap pressure as root activity resumes in response to soil temperatures exceeding 8-10°C: the precise timing of stage B is determined by the interaction between chilling requirement satisfaction (endodormancy release) and heat unit accumulation (ecodormancy release), making it cultivar-specific and strongly influenced by winter and early spring temperature patterns. Stage B is the earliest phenological event used in spring frost risk assessment because it signals that sensitive developmental processes have been initiated.
Baggiolini stage B
Baggiolini-Stadium B
Knospenschwellen
Knospenschwellstadium
bud expansion stage
bud swelling stage
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 80-85
Baggiolini B Bud Swelling
A phenological observation of Baggiolini C for Gamay in Beaujolais on April 5 documents the first emergence of a green shoot tip (1-3 mm) through the brown wool surrounding the swollen bud — this pointe verte marks the critical frost vulnerability threshold because the exposed green tissue is killed at -2 to -3°C whereas the enclosed dormant bud (Baggiolini A-B) tolerates -15 to -20°C, triggering activation of frost protection systems (bougies, aspersion, wind machines) in frost-prone sites when sub-zero temperatures are forecast during the Baggiolini C period.
A Baggiolini code designating stage C (pointe verte) that describes the moment when green shoot tissue first becomes visible at the tip of the swelling bud, emerging through the separating bud scales and protective wool layer, marking the irreversible commitment to active shoot growth and the onset of acute spring frost vulnerability, corresponding approximately to BBCH 07.
Baggiolini_C_Green_Point (French pointe verte, German Grünpunkt) is one of the most practically important phenological stages because it defines the threshold of spring frost vulnerability. The appearance of green tissue through the bud scales represents a dramatic reduction in cold hardiness from approximately -15 to -20°C at Baggiolini B to -2 to -3°C at Baggiolini C: this 15-18°C reduction in frost tolerance within a span of days makes the Baggiolini B-to-C transition the most consequential single phenological event for frost risk management. The stage corresponds approximately to BBCH 07 (beginning of bud burst) and is the most frequently reported spring phenological milestone in European viticultural practice. Early-bursting cultivars (Chardonnay, Pinot Noir) reach Baggiolini C 10-21 days before late-bursting cultivars (Cabernet Sauvignon, Mourvèdre) at the same site.
Baggiolini stage C
Baggiolini-Stadium C
Grüne-Spitze-Stadium
Grünpunktstadium
green point stage
green tip stage
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 85-95 Lebon E et al. (2008) Modelling the seasonal dynamics of the soil water balance of vineyards, Functional Plant Biology 35: 1065-1081
Baggiolini C Green Point
A Swiss viticulturist recording Baggiolini stage E for Chasselas in the Lavaux on April 20 communicates that leaves are fully unfolded and clearly visible — the Baggiolini system progresses alphabetically from A (winter bud) through P (berry ripening), with each letter corresponding to a specific visual appearance documented in Baggiolini's original 1952 photographic reference plates, and the system remains the preferred phenological coding method in French-speaking and Italian-speaking viticultural regions where it was adopted as the standard decades before the BBCH system's introduction in 1995.
A phenological code assigned according to the Baggiolini alphabetic staging system for grapevine development, consisting of a single capital letter (A through P) that designates one of sixteen visually defined developmental stages from winter dormancy through berry ripening, representing the historically predominant European phenological classification for Vitis that preceded and coexists with the BBCH numerical system.
Baggiolini_Code represents the classical European grapevine phenological staging system published by M. Baggiolini in 1952 and widely adopted across francophone, italophone, and hispanophone viticulture. The system uses 16 alphabetic stages (A-P) corresponding to visually distinctive developmental milestones illustrated with photographic reference images: its strengths include simplicity, visual intuitiveness, and a 70+ year legacy of continuous phenological records in European research stations. The BBCH system (Lorenz et al. 1995) was developed partly to provide a numerical alternative compatible with the general BBCH crop staging framework, and cross-reference tables mapping Baggiolini letters to BBCH codes are available — however, the correspondence is not always one-to-one because the two systems define stage boundaries at slightly different morphological landmarks.
Baggiolini phenological letter
Baggiolini stage
Baggiolini-Code
Baggiolini-Stadium
alphabetic growth stage code
alphabetischer Phänologie-Code
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Lorenz DH et al. (1995) Growth stages of the grapevine: phenological growth stages of the grapevine — Codes and descriptions according to the extended BBCH scale, Australian Journal of Grape and Wine Research 1: 100-103
Baggiolini Code
A phenological record of Baggiolini D for Sauvignon Blanc in Sancerre on April 15 documents the emergence of 1-2 small leaves from the burst bud with blades partially unfurled and showing characteristic bronze-green coloration, the shoot tip visible as a compact rosette of young leaves 1-2 cm in length — at this stage the developing shoot is elongating at 0.5-2 cm per day, frost sensitivity remains high at -1.5 to -2.5°C lethal temperature, and the first post-budburst fungicide applications targeting Excoriose (Phomopsis viticola) are scheduled based on the attainment of Baggiolini D when emerging leaves provide sufficient target surface for contact protectants.
A Baggiolini code designating stage D that describes the emergence of the first recognizable leaf blades from the expanding bud, with leaves still folded or partially rolled and typically showing reddish or bronzed coloration in many cultivars, marking the transition from bud burst to active leaf development, corresponding approximately to BBCH 09-11.
Baggiolini_D_Leaf_Emergence bridges the transition between bud burst and autonomous photosynthetic growth. At this stage the developing shoot is transitioning from heterotrophic dependence on stored carbohydrate reserves to autotrophic carbon assimilation as the first functional leaf area is established: this metabolic transition makes Baggiolini D a sensitive indicator of vine carbohydrate reserve status, with poorly nourished vines (excessive prior-season crop load, winter damage, trunk disease) showing delayed and weak stage D development. The reddish or bronzed coloration of emerging leaves in many cultivars (particularly Pinot Noir, Cabernet Sauvignon, Merlot) is caused by anthocyanin accumulation that provides UV protection to the photosynthetically immature leaf tissue, and the intensity of this coloration is a useful cultivar-discriminating character at the Baggiolini D stage.
Baggiolini stage D
Baggiolini-Stadium D
Blattaustriebsstadium
Rosettenstadium
leaf emergence stage
rosette stage
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 95-110
Baggiolini D Leaf Emergence
A phenological observation of Baggiolini E for Nebbiolo in Barolo on April 25 documents 2-3 fully unfolded leaves separated from the shoot tip with blades 3-5 cm across, the shoot measuring 5-10 cm in length and elongating at 2-5 cm per day — at Baggiolini E the first cluster primordia are visible as small green buttons along the shoot, enabling preliminary fruitfulness assessment and providing the staging reference for the first anti-downy-mildew treatment in regions where Plasmopara viticola primary inoculum release coincides with the 10°C/10 mm/24 h rule (10°C minimum temperature, 10 mm rainfall, 24 h leaf wetness).
A Baggiolini code designating stage E that describes the stage where 2-3 leaves are fully unfolded and clearly separated from the shoot tip with blades flattened and expanding, the shoot now visibly elongating and inflorescence primordia becoming detectable as small lateral protuberances along the developing shoot axis, corresponding approximately to BBCH 12-13.
Baggiolini_E_Leaves_Unfolded marks the establishment of functional canopy and the beginning of the period when inflorescence primordia become externally visible. This stage is significant for integrated pest management because the expanding leaf area provides both the target tissue for fungal infection and the surface area for protective fungicide deposition: the timing of the first fungicide application relative to Baggiolini E is a critical decision point that balances the risk of early-season infection against the cost and environmental impact of prophylactic spraying. The correspondence with BBCH 12-13 is approximate because Baggiolini defined stage boundaries by visual gestalt rather than by exact leaf count, and individual observers may interpret the E stage threshold slightly differently.
Baggiolini stage E
Baggiolini-Stadium E
Blätter-entfaltet-Stadium
Frühes-Triebwachstum-Stadium
early shoot growth stage
leaves unfolded stage
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 95-125
Baggiolini E Leaves Unfolded
A phenological record of Baggiolini F for Syrah in Côte-Rôtie on May 15 documents clearly visible inflorescences 3-5 cm in length with compact flower bud clusters separated from the shoot axis and individually distinguishable under close inspection — the attainment of Baggiolini F enables the first reliable assessment of potential cluster number per shoot (typically 1-3 depending on cultivar and bud fruitfulness), triggering preliminary yield estimation calculations and serving as the reference stage for shoot thinning decisions where excess non-fruitful shoots are removed to optimize canopy-to-fruit ratio at 12-18 shoots per meter of cordon.
A Baggiolini code designating stage F that describes the stage where inflorescences (developing flower clusters) are clearly visible and separated from the shoot axis, with individual flower buds distinguishable within the compact cluster structure but still pressed tightly together, corresponding approximately to BBCH 53-55.
Baggiolini_F_Inflorescence_Visible corresponds to the pre-bloom period when the reproductive structures are fully exposed and developing toward anthesis. The stage maps approximately to BBCH 53-55 (inflorescences clearly visible to fully developed with flowers pressed together) and represents the window during which pre-bloom viticultural interventions — shoot thinning, shoot positioning, and pre-bloom cluster thinning — are most effectively performed: these interventions exploit the plasticity of reproductive development before the irreversible commitment of flowering and fruit set. Disease management at Baggiolini F focuses on protecting the highly susceptible developing inflorescences against downy mildew (Plasmopara viticola) whose oospore-derived primary infections specifically target inflorescence tissue during wet weather at this stage.
Baggiolini stage F
Baggiolini-Stadium F
Gescheine-sichtbar-Stadium
Infloreszenz-sichtbar-Stadium
cluster visible stage
visible inflorescence stage
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 125-145
Baggiolini F Inflorescence Visible
A phenological observation of Baggiolini G for Chenin Blanc in Vouvray on June 1 documents inflorescences with individually separated flower buds clearly distinguishable to the naked eye, each bud 2-3 mm in diameter with the green calyptra still intact and no evidence of cap detachment — Baggiolini G marks the final pre-bloom stage and triggers the critical pre-flowering fungicide application window for Botrytis cinerea protection, as the separation of individual flowers creates microhabitat niches within the cluster architecture where moisture retention favors Botrytis colonization during and immediately after anthesis.
A Baggiolini code designating stage G that describes the stage immediately preceding anthesis where individual flower buds within the inflorescence have separated from each other and are clearly distinguishable as discrete units with closed flowerhoods (calyptrae) still intact, the inflorescence fully elongated and flower buds at their maximum pre-bloom size, corresponding approximately to BBCH 57.
Baggiolini_G_Separated_Flowers defines the immediate pre-anthesis stage that represents the last opportunity for several time-critical viticultural interventions. The separation of individual flower buds within the inflorescence creates the final cluster architecture that will determine bunch compactness after fruit set — a key determinant of Botrytis susceptibility in tight-clustered cultivars such as Pinot Noir, Riesling, and Chenin Blanc: the observation of Baggiolini G therefore triggers both protective fungicide applications and, in some management systems, pre-bloom cluster treatments (gibberellic acid applications in table grapes, cluster-zone leaf removal for bunch aeration). The correspondence with BBCH 57 (flowers separating) is close but not exact, as Baggiolini's photographic definition emphasizes the visual gestalt of separated but still-closed flowers rather than a precise percentage-based criterion.
Baggiolini stage G
Baggiolini-Stadium G
Einzelblüten-sichtbar-Stadium
Gescheine-getrennt-Stadium
pre-bloom stage
separated flowers stage
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 140-145 May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press pp: 1-30
Baggiolini G Separated Flowers
A phenological record of Baggiolini H for Gewürztraminer in Alsace on June 10 documents active flowering with approximately 40-60% of flowerhoods fallen across the inflorescence, stamens reflexed and pollen visible on receptive stigmas — Baggiolini H encompasses the entire flowering period lasting 6-14 days under favorable conditions (20-25°C, low rainfall), with the mid-point of stage H corresponding approximately to BBCH 65 (full flowering, 50% caps fallen), and the stage serving as the critical window for assessing fruit set potential based on weather conditions during pollination — temperatures below 15°C or persistent rainfall during Baggiolini H leading to coulure rates of 30-80% in sensitive cultivars such as Merlot and Grenache.
A Baggiolini code designating stage H that describes the period of active anthesis where flowerhoods (calyptrae) are detaching from the receptacle and falling, exposing stamens and pistils, with the stage conventionally defined as the period from first visible cap fall through the completion of flowering when all flowerhoods have been shed, corresponding approximately to BBCH 61-69.
Baggiolini_H_Flowering spans the complete anthesis period that the BBCH system subdivides into multiple secondary stages (BBCH 61 through 69). This difference in resolution illustrates a fundamental structural distinction between the two phenological systems: Baggiolini uses a single letter for the entire flowering period, prioritizing observational simplicity, while BBCH provides percentage-based subdivisions (10%, 30%, 50%, 70%, 80% cap fall) enabling more precise temporal positioning within the flowering window. For practical management decisions — particularly fungicide timing and fruit set prediction — the BBCH subdivisions within stage H provide operationally useful precision, explaining why many modern phenological recording systems use both Baggiolini letters and BBCH codes in parallel.
Anthese-Stadium
Baggiolini stage H
Baggiolini-Stadium H
Blütestadium
anthesis stage
flowering stage
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 145-175 May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press pp: 30-80
Baggiolini H Flowering
A phenological observation of Baggiolini I for Cabernet Franc in Chinon on June 25 documents young berries of 2-4 mm diameter visible on the cluster rachis with residual flowerhoods and unfertilized flowers fallen or falling, the successful fruit set rate estimated at 30-45% of total flowers per cluster — at Baggiolini I the first reliable post-bloom yield estimate becomes possible by counting set berries per cluster across representative shoots, with observed fruit set rates of 25-50% considered normal for Vitis vinifera and rates below 20% indicating significant coulure that may necessitate revised crop load management strategies.
A Baggiolini code designating stage I that describes the post-flowering period where successfully fertilized ovaries begin to enlarge into recognizable young berries while unfertilized flowers abscise, marking the transition from reproductive to fruit growth phases with the initial determination of berry number per cluster, corresponding approximately to BBCH 71.
Baggiolini_I_Fruit_Set defines the critical transition point where potential yield (flower number) is converted to actual yield (berry number). The fruit set percentage — the proportion of flowers that develop into berries — is one of the most variable yield components in viticulture, ranging from 15% to 60% depending on cultivar genetics, weather conditions during Baggiolini H, vine nutritional status (particularly boron availability), and carbohydrate reserve levels: cultivars such as Merlot, Grenache, and Malbec are particularly susceptible to poor fruit set (coulure), while Chardonnay and Riesling typically achieve more consistent set rates. Baggiolini I also marks the beginning of the first phase of berry growth (cell division), initiating the double-sigmoid growth curve that characterizes grape berry development.
Baggiolini stage I
Baggiolini-Stadium I
Beerenansatz-Stadium
Fruchtansatzstadium
berry initiation stage
fruit set stage
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 175-195 May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press pp: 80-120
Baggiolini I Fruit Set
A phenological record of Baggiolini J for Sangiovese in Montalcino on July 10 documents berries at 6-8 mm diameter with hard green pericarp, clusters hanging 10-15 cm below the cordon wire and beginning to assume their characteristic cluster shape — Baggiolini J marks the standard reference stage for post-fruit-set cluster thinning (vendange verte) in premium viticulture, where excess clusters are removed to achieve target yields of 4-8 t/ha, as berry number per cluster is now fixed and cluster weight can be estimated from berry count and projected berry size at harvest using cultivar-specific regression models.
A Baggiolini code designating stage J that describes the berry development stage where berries have reached the approximate size of a small pea (5-8 mm diameter), clusters are beginning to hang under their own weight, and the berries are hard, green, and actively accumulating organic acids, corresponding approximately to BBCH 75.
Baggiolini_J_Pea_Size corresponds to the mid-point of Phase I (cell division phase) of berry growth, when pericarp cells are actively dividing to establish the final cell number that determines maximum potential berry size. At this stage berries contain maximum concentrations of organic acids (predominantly tartaric and malic acid, reaching 25-35 g/L titratable acidity), minimal sugar (3-5° Brix), and no significant flavor or aroma compound development: the hard, acidic, green berry at Baggiolini J is physiologically optimized for seed protection against herbivory through unpalatability. Cluster thinning at Baggiolini J is preferred over earlier thinning because fruit set has been confirmed and compensatory growth responses to thinning are minimal at this relatively advanced developmental stage.
Baggiolini stage J
Baggiolini-Stadium J
Erbsengroße-Stadium
Schrotkorn-Stadium
pea size stage
small berry stage
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 195-215
Baggiolini J Pea Size
A phenological observation of Baggiolini K for Pinot Noir in the Côte de Nuits on July 25 documents berries touching and compressing against each other within the characteristically tight cylindrical clusters, with individual berries deforming slightly at contact points and the cluster interior becoming inaccessible to spray penetration — Baggiolini K represents the last practical opportunity for pre-véraison Botrytis management in tight-clustered cultivars, as the closed cluster architecture from this stage onward creates the humid microenvironment (>95% relative humidity within the cluster) that favors Botrytis cinerea infection of berry contact zones during ripening.
A Baggiolini code designating stage K that describes the berry development stage where expanding berries within the cluster make contact with adjacent berries, closing the previously open spaces within the cluster architecture and creating the compact cluster structure characteristic of the pre-véraison period, corresponding approximately to BBCH 77-79.
Baggiolini_K_Bunch_Closure marks a structurally significant transition in cluster microclimate from an open, well-ventilated architecture to a closed, humidity-retaining configuration. The timing and completeness of bunch closure are cultivar-specific and yield-dependent: tight-clustered cultivars (Pinot Noir, Riesling, Chenin Blanc) achieve near-complete closure at Baggiolini K, creating conditions conducive to Botrytis development, while loose-clustered cultivars (Syrah, Nebbiolo, Sangiovese) may never fully close even at high yields. This stage corresponds to the end of Phase I and the beginning of the lag phase (Phase II) of the double-sigmoid berry growth curve, a period of minimal berry volume increase during which seeds approach physiological maturity and organic acid concentrations begin their pre-véraison peak.
Baggiolini stage K
Baggiolini-Stadium K
Beerenschluss-Stadium
Traubenschluss-Stadium
bunch closure stage
cluster compaction stage
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 215-225
Baggiolini K Bunch Closure
A phenological record of Baggiolini L for Malbec in Cahors on August 5 documents the first berries within each cluster transitioning from hard green to soft purple-red, with approximately 30-50% of berries showing color change while the remainder are still green and firm — the asynchronous onset of véraison within and between clusters is characteristic, with individual clusters requiring 5-10 days for all berries to complete the color transition, and the observation of Baggiolini L triggering irrigation management changes (initiation of post-véraison deficit irrigation at 30-50% ETc replacement), bird-netting installation, and the scheduling of the first ripening-phase berry sampling for Brix, pH, and titratable acidity monitoring.
A Baggiolini code designating stage L (véraison) that describes the onset of ripening marked by the simultaneous occurrence of berry softening (detectable by gentle compression) and the development of cultivar-specific berry color — from green to yellow-translucent in white cultivars and from green through red to blue-black in red cultivars — representing the fundamental developmental switch from acid accumulation to sugar accumulation, corresponding approximately to BBCH 81-83.
Baggiolini_L_Veraison is arguably the single most important phenological milestone in viticultural practice because it signals the irreversible transition from the vegetative-reproductive growth phase to the ripening phase. The biochemical events underlying véraison include the onset of ABA (abscisic acid) signaling, activation of sugar transporter genes, initiation of anthocyanin biosynthesis in red cultivars, cell wall disassembly causing berry softening, and the metabolic switch from tartaric and malic acid accumulation to sugar import from phloem: these coordinated changes occur over 24-48 hours in individual berries but are spread across 1-2 weeks within a cluster due to positional variation in developmental timing. Véraison date is strongly cultivar-dependent (early in Pinot Noir, late in Cabernet Sauvignon) and serves as the primary reference point for predicting harvest date, typically 35-55 days post-véraison for table wine production.
Baggiolini stage L
Baggiolini-Stadium L
Farbumschlag-Stadium
Véraison-Stadium
color change stage
véraison stage
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 225-260 Coombe BG and McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening, Australian Journal of Grape and Wine Research 6: 131-135
Baggiolini L Veraison
A phenological observation of Baggiolini M for Riesling in the Rheingau on October 5 documents berries at optimal harvest maturity with 85° Oechsle (20.5° Brix), pH 3.05, titratable acidity 9.2 g/L, and developed varietal thiol and monoterpene aromas — the definition of Baggiolini M is inherently style-dependent, as a Riesling destined for Kabinett may reach Baggiolini M at 76° Oechsle while the same vineyard block destined for Spätlese requires 85° Oechsle, and a Trockenbeerenauslese requires 150°+ Oechsle achieved only through noble rot concentration weeks after conventional Baggiolini M for the base fruit.
A Baggiolini code designating stage M that describes the period of harvest maturity where berries have reached their target sugar concentration, acid balance, flavor development, and phenolic maturity as determined by the winemaker's quality objectives and the intended wine style, corresponding approximately to BBCH 87-89.
Baggiolini_M_Maturity is unique among the Baggiolini stages in that its definition is partly subjective and depends on the intended use of the harvested fruit. While all other Baggiolini stages describe objective morphological landmarks observable by any trained phenologist, stage M incorporates oenological judgment about when the fruit has reached its optimal quality window for a specific wine style: this makes Baggiolini M a hybrid phenological-oenological concept rather than a purely developmental milestone. The corresponding BBCH codes 87-89 similarly describe qualitative ripeness states (berries ripe for harvest, berries overripe) rather than objectively measurable morphological criteria, reflecting the inherent difficulty of codifying the multidimensional concept of harvest maturity into a single staging descriptor.
Baggiolini stage M
Baggiolini-Stadium M
Lesereife-Stadium
Reifestadium
harvest ripeness stage
maturity stage
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 260-295 Coombe BG (1995) Growth stages of the grapevine, Australian Journal of Grape and Wine Research 1: 104-110
Baggiolini M Maturity
A phenological record of Baggiolini N for Grüner Veltliner in the Kremstal on November 10 documents progressive leaf senescence with 60-70% of leaves fallen, remaining leaves showing yellow-brown coloration with necrotic margins, and canes fully lignified with brown mature periderm — the progression through Baggiolini N spans 3-6 weeks from first leaf yellowing to complete defoliation, with the timing determined by autumn temperature decline, first frost events (killing unlignified shoot tips and accelerating leaf abscission), and cultivar-specific senescence programming — late-senescing cultivars such as Cabernet Sauvignon may retain functional green leaves 2-4 weeks longer than early-senescing cultivars such as Pinot Noir at the same site.
A Baggiolini code designating stage N that describes the post-harvest period of leaf senescence and abscission where leaves progressively yellow, desiccate, and fall from the vine, culminating in complete defoliation and the vine's entry into winter dormancy, completing the annual phenological cycle and corresponding approximately to BBCH 93-97.
Baggiolini_N_Leaf_Fall is the final stage in the Baggiolini alphabetic sequence (stages O and P are not universally recognized in all versions of the system). The stage encompasses the critical physiological processes of nutrient remobilization and reserve storage that determine the vine's preparedness for winter survival and the following season's growth vigor: nitrogen, phosphorus, and potassium are recovered from senescing leaves at efficiencies of 50-70% and translocated to perennial wood and root tissues, while starch reserves accumulated during the ripening period are stored in ray parenchyma cells of the trunk and roots. The completeness of Baggiolini N (achieving full natural defoliation before severe frost) is an indicator of seasonal balance — vines that retain green leaves at the time of first hard frost have typically suffered from excessive crop load, late-season disease pressure, or insufficient heat accumulation for complete shoot maturation.
Baggiolini stage N
Baggiolini-Stadium N
Blattfallstadium
Entblätterungs-Stadium
defoliation stage
leaf fall stage
Baggiolini M (1952) Les stades repères dans le développement annuel de la vigne et leur utilisation pratique, Revue Romande d'Agriculture et d'Arboriculture 8: 4-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 295-330 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp: 80-108
Baggiolini N Leaf Fall
Balance assessment for Riesling Spätlese from the Mosel (2024 vintage, 8.5% alcohol, 45 g/L RS, 8.5 g/L TA) rates as "exceptionally balanced" — the substantial residual sugar is precisely counterpoised by the high acidity, creating a tensile equilibrium where neither sweetness nor sourness dominates and the low alcohol contributes lightness that prevents the sweetness from becoming cloying, producing the quintessential Mosel Spätlese sensation of effortless poise between opposing forces. Balance assessment for over-extracted Cabernet Sauvignon from a warm-climate source (2022 vintage, 15.2% alcohol, TA 4.8 g/L, pH 3.82) rates as "moderately unbalanced" — the excessive alcohol creates a hot, burning finish that overwhelms the fruit, the insufficient acidity fails to provide structural counterweight to the heavy body, and the high pH reduces the vibrancy that might otherwise integrate the powerful tannin extraction, resulting in a structurally top-heavy wine despite impressive individual component concentration. Balance assessment for Grüner Veltliner Smaragd from the Wachau (2024 vintage) evaluates as "well-balanced to exceptionally balanced" — the firm acidity (TA 5.8 g/L) provides architectural backbone for the concentrated fruit and medium-full body (13.5% alcohol), with neither element overshadowing the other and the mineral persistence completing the structural arc.
Balance assessment is an overall assessment entity that records the perceived harmony and proportional integration among the principal structural elements of a wine — sweetness, acidity, tannin (in red wines), alcohol, body, and fruit concentration — evaluated as a unified structural impression where the ideal state is one in which no single element dominates or is deficient relative to the others, assessed on a scale from unbalanced (one or more elements conspicuously dominant or absent, creating structural disharmony) through moderately balanced to well-balanced and exceptionally balanced (all elements seamlessly integrated, the structural architecture perceived as effortless and inevitable), and representing the quality criterion most universally valued across all wine styles, regions, and assessment traditions.
Balance assessment captures the most universally prized quality attribute in wine — a meta-property that emerges from the interaction of individual structural components rather than from any single parameter, and that transcends stylistic preferences to represent a quality criterion valued equally in light Mosel Kabinett and powerful Napa Cabernet. The perception of balance is inherently relational — it is not the absolute level of any component that matters but the proportional relationship among components, meaning that a wine with high acidity, high tannin, high alcohol, and high fruit concentration may be perceived as perfectly balanced (all elements strong but proportionate), while a wine with moderate levels of everything except excessive alcohol may be perceived as unbalanced. The classical European wine aesthetic framework codified the balance concept in terms of specific structural relationships — the Bordeaux tradition emphasizing the acidity-tannin-fruit triangle, the Burgundian emphasis on the acid-extract balance expressed as "tension," the German Riesling concept of Süße-Säure-Spiel (sweetness-acidity interplay), and the Austrian emphasis on Trinkfluss (drinking flow, the seamless progression from attack through mid-palate to finish). The neurological basis of balance perception likely involves the integration of multiple taste and somatosensory inputs in the orbitofrontal cortex, where the assessment of proportionality represents a higher-order cognitive evaluation rather than a direct sensory measurement — explaining why balance assessment requires training and experience and why novice tasters often focus on individual parameters (sweetness, acidity) rather than their integration. The temporal dimension of balance assessment is critical — a wine may be structurally unbalanced in youth (tannin-dominant, acid-aggressive) but predictably evolving toward balance through aging, and experienced tasters evaluate not only current balance but projected balance at the wine's anticipated maturity, adding a temporal prediction component to the assessment.
Ausgewogenheit
Balancebeurteilung
Gleichgewichtsbewertung
Harmonie
equilibrium assessment
harmony assessment
proportional integration
structural balance
strukturelle Balance
wine balance
Peynaud E, Blouin J (2006) The Taste of Wine, 4th Ed, Wiley, pp.: 280-300 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 535-545 Amerine MA, Roessler EB (1983) Wines: Their Sensory Evaluation, 2nd Ed, Freeman, pp.: 140-170 Parr WV et al. (2011) Representation of complexity in wine: influence of expertise. Food Quality and Preference 22(7): 647-660
Balance Assessment
A balanced bilateral cordon on Vitis vinifera cv. Chardonnay in Napa Valley, featuring left cordon at 85cm and right cordon at 83cm (2cm difference, 2.4% variation well within balanced tolerance), each bearing 14 two-bud spurs spaced uniformly at 12cm intervals, trunk precisely centered on trellis wire equidistant from neighboring vines, balanced vigor producing similar shoot development from both cordons (left average 152cm, right average 148cm shoot length), crop distribution 51% left versus 49% right (total 5.2kg, essentially balanced), representing ideal balanced bilateral architecture achieved through careful establishment training, centered trunk positioning, and skilled annual management maintaining architectural symmetry and functional balance.
A bilateral cordon configuration with closely matched dimensions and fruiting capacity between the two cordons, characterized by equal or nearly equal cordon lengths (typically within 10-15cm), similar spur counts and spacing on both cordons, balanced crop distribution approaching 50=50 split between left and right sides, and deliberate architectural uniformity reflecting centered trunk positioning, balanced vine vigor, and management priority for symmetrical development. Balanced bilateral cordons represent the ideal outcome in bilateral establishment and management, achieved when trunks are precisely centered on trellis wires, vines show balanced vigor without directional preferences, initial training successfully develops equal extensions, and annual maintenance pruning preserves symmetrical architecture through balanced spur renewal and shoot management, resulting in aesthetically pleasing and functionally optimal vine architecture.
Balanced bilateral cordons represent aesthetic and functional ideals in modern cordon viticulture, with close symmetry indicating precise establishment, balanced vine physiology, and skilled management. Perfect balance (exactly equal lengths, spur counts, crops) is neither always achievable nor strictly necessary, with minor variations (5-15% differences) being normal and functionally acceptable. The pursuit of balance serves practical purposes including simplified mechanization (symmetrical canopies present consistent targets for equipment), predictable vine behavior (balanced cordons show similar phenology and fruit development facilitating uniform management timing), easier manual operations (workers develop consistent expectations and techniques for symmetrical vines), and optimal resource utilization (balanced distribution prevents one cordon dominating and potentially exhausting while other underperforms). Achieving and maintaining balance requires attention to trunk positioning (centered on wire), spur renewal (preventing progressive lengthening of one cordon), and shoot management (removing water shoots and suckers that might shift vigor balance).
ausgeglichene bilaterale Kordon
equal paired cordon
symmetrical bilateral cordon
symmetrische Doppelkordon
uniform bilateral cordon
Balanced Bilateral Cordon
Pinot Noir clone 114 bears a Balanced_Clone_Role in French viticulture, positioned between the quality-focused Dijon clones (115, 667, 777) and high-yield selections—producing moderately sized clusters, reasonable yields (6-8 t/ha), and wines of good commercial quality suitable for regional Burgundy and Champagne production. Chardonnay clone 95 serves in a Balanced_Clone_Role for California coastal production, offering reliable productivity (6-10 t/ha) with adequate quality for mid-tier varietal bottlings without the yield sacrifice of quality clone 76 or concentration limitations of high-yield selections. Merlot clone 181 bears a Balanced_Clone_Role in Bordeaux, widely planted for its combination of consistent productivity, good color development, and commercially appealing wine character suitable for appellation and château production across quality tiers.
A performance clone role that inheres in a grapevine clone characterized by intermediate performance between high-yield and quality extremes, offering commercially acceptable productivity combined with satisfactory quality attributes for mainstream wine production. Balanced clones represent the pragmatic middle ground for commercial viticulture, where neither maximum yield nor optimal quality is the sole objective but rather an economically optimal combination of both. These clones typically produce moderate yields (within 10-20% of variety averages), acceptable fruit composition (adequate sugar, acidity, and phenolics for varietal wine production), reliable performance across variable seasons, and wines of good commercial quality without exceptional concentration or complexity. Balanced clones may constitute the majority of commercial plantings where producers seek dependable production for mid-tier markets, with quality clones reserved for premium bottlings and high-yield clones for bulk production. The balanced designation reflects market positioning rather than genetic compromise these clones represent deliberate selection for versatile, commercially viable performance.
Balanced_Clone_Role captures the commercially pragmatic center of the performance spectrum where most production volume originates. The designation is relative and market-dependent—what constitutes "balanced" varies by variety, region, and production context. Balanced clones may represent default selections where specific quality or yield optimization is not required, functioning as "workhorse" clones supporting reliable commercial production. The category overlaps with Quality and High_Yield designations at its boundaries clone characterization often involves degree rather than categorical distinction. Balanced clones offer risk management advantages—avoiding the yield sacrifice of quality clones while exceeding the quality limitations of high-yield selections—making them appropriate for diverse production contexts and variable market conditions.
Allzweckklon
Handelsklon
Standardklon
all-purpose clone
ausgewogener Klon
balanced clone
commercial clone
standard clone
ENTAV-INRA (2007) Catalogue des Variétés et Clones de Vigne Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 Anderson K, Aryal NR (2013) Which Winegrape Varieties are Grown Where?, University of Adelaide Press
Balanced Clone Role
Pinot Noir in Burgundy's Côte d'Or typically exhibits balanced hermaphrodite flowers at BBCH 65 with stamens extending 0.4-0.7 mm beyond pistil, producing 14,000-19,000 pollen grains per flower with 62-72% viability, achieving 38-48% fruit set under typical Burgundian bloom conditions (mid-June temperatures 18-24°C, variable humidity and occasional precipitation), yielding clusters of 70-110 berries from initial flower complements of 180-230 flowers per inflorescence, with moderate berry numbers contributing to the cultivar's characteristic small cluster size and concentrated fruit quality. Riesling in Germany's Mosel Valley shows balanced hermaphrodite flowers with moderate stamen extension (0.5-0.8 mm beyond pistil), 15,000-21,000 pollen grains per flower, 65-74% viability, and 40-50% fruit set under cool bloom conditions, producing elegant clusters of 80-120 berries suitable for high-quality wine production.
A balanced hermaphrodite flower is a hermaphrodite flower exhibiting adequate development and function of both male and female organs with more equal proportions between stamen and pistil dimensions, moderate pollen production and viability, and good stigma receptivity, achieving satisfactory fruit set percentages suitable for commercial production while showing less pronounced stamen extension and somewhat reduced pollen abundance compared to fully functional hermaphrodite flowers. Balanced hermaphrodite flowers are characterized by= (1) stamens extending approximately 0.3-0.8 mm beyond pistil length, creating moderate spatial separation between anthers and stigma sufficient for self-pollination but less pronounced than in fully functional types (2) anthers producing moderate pollen quantities (12,000-22,000 pollen grains per flower) with good viability (60-75% viable pollen under favorable conditions) (3) pistil with fully developed ovary, functional style, and receptive stigma, generally comparable to fully functional types in pistil development (4) adequate developmental synchrony between pollen maturation and stigma receptivity, ensuring successful self-pollination despite less pronounced organ dimorphism. The 'balanced' designation indicates more equal development of male and female organs rather than the pronounced stamen dominance seen in fully functional types, while still maintaining functional hermaphroditism and self-fertility. Balanced hermaphrodite flowers typically achieve fruit set percentages of 35-50% under good environmental conditions, representing satisfactory fertilization success for commercial production though somewhat lower than the 50-65% achieved by fully functional types. The moderate fruit set of balanced hermaphrodite flowers results from adequate but not maximal pollen production (sufficient to saturate stigmas but with less margin for environmental pollen loss), moderate stamen extension (requiring some air movement or favorable pollen settlement conditions for efficient self-pollination), and good but not optimal pollen viability and pistil function. Balanced hermaphrodite morphology represents the typical expression in many commercially successful wine cultivars under normal cultural management and environmental conditions, with this phenotype providing adequate reproductive performance for economic yields (40-50% fruit set translating to 60-120 berries per cluster from typical 150-250 flower inflorescences) while potentially offering advantages in moderate cropping that can enhance berry quality through resource concentration. The balanced condition may represent either stable cultivar characteristics (some cultivars consistently show balanced rather than fully functional expression) or environmental/cultural modulation of potentially fully functional cultivars (high crop loads, marginal nutritional status, or suboptimal previous-season conditions during flower differentiation can shift expression from fully functional toward balanced). Balanced hermaphrodite flowers are characteristic of many Pinot Noir clones, Riesling, Sauvignon Blanc, and numerous other wine cultivars under typical viticultural management, providing reliable self-fertility and adequate yields without the maximum reproductive investment seen in fully functional types. The balanced condition may be agronomically desirable in some contexts, as the moderate fruit set naturally limits crop load and concentrates resources into fewer berries, potentially enhancing quality parameters (berry size, sugar accumulation, phenolic development) compared to the higher berry numbers resulting from fully functional hermaphrodite flowers with maximum fruit set.
Balanced_Hermaphrodite_Flower represents typical or adequate hermaphrodite functionality and is distinguished from Fully_Functional_Hermaphrodite_Flower by quantitative measures showing moderate rather than maximum organ development (less stamen extension, moderate pollen production, moderate fruit set). The boundary between these categories is gradational but reflects a real distinction between maximum and typical reproductive performance. Balanced expression is agronomically adequate and in some quality wine production contexts may be preferred, as the naturally limited fruit set concentrates resources and potentially enhances berry quality. Cultural manipulation can shift expression between balanced and fully functional states, with crop thinning, canopy management, and nutritional optimization potentially enhancing hermaphrodite functionality, while over-cropping, shade stress, or nutritional limitations may reduce expression from fully functional to balanced. Understanding the environmental and genetic determinants of the balanced versus fully functional distinction could inform breeding programs and cultural protocols for optimizing the fruit set-quality balance for specific production goals.
ausgeglichene Zwitterblüte
ausgewogene Zwitterblüte
balanciert hermaphroditisch
moderately functional bisexual flower
proportionate hermaphrodite flower
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 103-108 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 97-102 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Ebadi, A. et al. (1996). Effect of short-term temperature and shading on fruit set, seed and berry development in model vines. Australian Journal of Grape and Wine Research 2=2-9.
Balanced Hermaphrodite Flower
A balanced medium fruiting cane on Vitis vinifera cv. Pinot Noir in Willamette Valley, Oregon, retained at 7 nodes spanning 75cm along wire, selected from moderate-vigor vine in Jory soil showing 380g pruning weight, positioned for optimal sun exposure on south-facing slope, expected to produce 6-7 fruiting shoots bearing 11-13 clusters totaling 2.8kg fruit, managed minimally with cluster thinning to single cluster per shoot (6-7 clusters, approximately 2kg final crop), achieving exceptional 18 cm²/g leaf area-to-fruit ratio, optimal fruit zone light penetration, and complete phenolic ripeness including fine-grained tannins and complex aromatics characteristic of ultra-premium Pinot Noir.
A medium fruiting cane at the shorter end of the moderate range, typically 6-8 nodes, representing conservative quality-first approach within medium cane category, characterized by restrained length (65-90cm), modest bud count prioritizing fruit quality over yield, suitability for premium production sites with naturally high fruitfulness or lower-vigor conditions, and production of limited shoot numbers facilitating exceptional canopy management and fruit development. Balanced medium fruiting canes emphasize sustainable vine physiology, appropriate leaf area-to-fruit ratios without excessive vegetative growth, optimal fruit exposure and microclimate control, and long-term vineyard health through conservative crop loads maintaining carbohydrate reserves and supporting complete wood maturation, representing quality philosophy accepting lower yields for superior fruit composition and wine complexity.
Balanced medium fruiting canes represent quality extremes in cane-pruned viticulture, with 6-8 node retention providing maximum quality potential through very low crop loads, excellent canopy light penetration, optimal fruit microclimate, and abundant carbohydrate availability supporting complete fruit maturation and complex secondary metabolite development. This approach suits ultra-premium producers where wine prices justify yield sacrifice (often 2-4 tons/hectare versus 6-10 for moderate medium canes), cultivars benefiting from crop reduction (Pinot Noir, Nebbiolo, Grenache), challenging vintages requiring crop insurance through conservative retention, and terroir-focused producers emphasizing site expression over productivity. Economic viability requires premium pricing, estate winery direct sales, or similar high-value marketing channels.
ausgewogene mittlere Fruchtrute
conservative medium cane
quality-focused medium cane
qualitätsorientierte Mittelrute
restrained moderate cane
Based on Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Bravdo et al. (1985) "Effect of crop level and crop load on growth, yield, must and wine composition, and quality of Cabernet Sauvignon" American Journal of Enology and Viticulture 36=125-131 Petrie et al. (2000) "The effect of leaf removal and canopy height on whole-vine gas exchange and fruit development of Vitis vinifera L. Sauvignon Blanc" Functional Plant Biology 27=569-578.
Balanced Medium Fruiting Cane
A bare-root grafted vine (Merlot/SO4) harvested in December from French nursery rows bears a Bare_Root_Vine_Role, roots washed, graded (first quality: 4+ roots of 2+ mm diameter, trunk >6 mm), bundled in quantities of 25, and stored at 2°C and 95% RH for February shipment to California vineyards. A bare-root one-year-old vine of Riesling/5BB serves in a Bare_Root_Vine_Role for German vineyard establishment, planted in April during the brief window between soil thaw and budbreak. A bare-root rootstock liner (110 Richter) bears a Bare_Root_Vine_Role, harvested and graded for root development before field planting and subsequent T-budding.
A nursery plant role that inheres in a grapevine harvested from nursery soil with roots washed or shaken free of soil, stored and shipped without growing medium, and planted directly into vineyard soil during the dormant season. Bare-root vines represent the traditional and still dominant format for dormant grapevine sales, with advantages including lower weight (no soil or container), reduced shipping costs, ease of root inspection and grading, and natural root architecture development unrestricted by container geometry. Bare-root production involves field-growing vines (either bench-grafted units or rooted rootstocks for budding) for one growing season, then harvesting during dormancy (typically November-March in Northern Hemisphere), removing soil from roots, grading by root and shoot quality, and bundling for cold storage or immediate shipment. Bare-root vine quality is assessed by root system development (number, distribution, length, and diameter of primary roots), shoot condition (trunk caliper, bud viability), and freedom from damage or disease. Bare-root vines require planting during dormancy before bud break, with success dependent on root protection from desiccation during handling and timely planting into adequately moist soil.
Bare_Root_Vine_Role captures the traditional nursery product format that remains dominant for dormant-season sales despite potted vine expansion. The format's advantages—lower cost, lighter weight, easy root inspection, natural root architecture—are balanced against requirements for dormant handling, limited planting window, and vulnerability to desiccation if mishandled. Bare-root vine success depends critically on maintaining root moisture from harvest through planting industry practices include cold storage (0-4°C, near-saturated humidity), wax coating of graft unions and exposed trunk portions, root dipping in hydrating gels, and careful transplanting technique. Quality grading systems establish minimum root standards (typically 3-5 roots of specified minimum diameter) that correlate with establishment success, with premium grades commanding higher prices reflecting their superior survival and growth potential.
Ballonlose Rebe
Freilandrebe
Wurzelnackte Rebe
bare-root vine
dormant vine
field-grown vine
lifted vine
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 380-390 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia European Union (2020) Commission Implementing Directive 2020/177 Annex I Gramaje D, Armengol J (2011) Fungal trunk pathogens, Phytopathologia Mediterranea 50: S217-S226
Bare Root Vine Role
A barren sterile shoot on declining Vitis vinifera cv. Carignan vine showing chronic Esca symptoms, originating from trunk water shoot following partial die-back, achieved only 62cm length over full season with 9 small chlorotic leaves totaling approximately 485 cm² (minimal photosynthetic capacity), 4mm diameter showing poor lignification, observed over two seasons with zero reproductive development (confirming sterility) and consistently poor vegetative performance (confirming barren status), representing combination of latent bud origin sterility plus compromised parent vine condition producing inadequate vegetative growth, this shoot providing essentially zero value for current photosynthesis, unsuitable for renewal wood, and maintained only as indicator of vine viability during assessment period determining whether vine recovery possible or removal and replanting necessary.
A sterile vegetative shoot showing extreme lack of reproductive capacity and poor vegetative development, characterized by persistent reproductive failure combined with weak growth vigor, sparse leaf development, minimal photosynthetic contribution, and poor prospects for any productive function including both current reproductive role and future architectural or renewal utility. Barren sterile shoots represent the combination of categorical sterility plus inadequate vegetative performance, arising from severely stressed conditions, declining vines showing progressive dysfunction, water shoots or suckers from compromised wood (diseased or severely stressed parent structures), or genetic/physiological problems affecting both reproductive and vegetative competence, with such shoots providing minimal vineyard value and typically removed unless no alternative exists for maintaining minimal vine presence.
Barren sterile shoots represent the worst-case combination of reproductive incompetence plus vegetative inadequacy, typically indicating severe problems requiring attention beyond simple shoot management. Common contexts include severely declining vines (trunk diseases, root diseases, virus infections, cumulative stress), water shoots from badly compromised wood (diseased sections, severely stressed zones), genetic or physiological dysfunctions (rare), and extreme environmental stress (severe drought, nutrient depletion, toxicity). Management typically removes barren sterile shoots recognizing minimal value, with retention only when broader vine assessment underway determining recovery potential versus removal necessity. The barren sterile designation serves as red flag indicating vine-level problems rather than just shoot-level characteristics, often triggering investigation of underlying causes and evaluation of vine viability and economic justification for continued maintenance versus replanting decisions.
barren non-fruiting shoot
futile vegetative shoot
nutzloser vegetativer Trieb
unfruchtbar-steriler Trieb
unproductive sterile shoot
Based on Pearson & Goheen (1988) "Compendium of Grape Diseases" APS Press, pp. 1-93 Graniti et al. (2000) "Esca of grapevine= a disease complex or a complex of diseases?" Phytopathologia Mediterranea 39=16-20 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 523-548.
Barren Sterile Shoot
Basal bud viability assessment in spur-pruned Syrah reveals the renewal capacity affecting training system sustainability, with examination of basal positions (nodes 1-2) showing 82% primary viability at node 2 (the typical pruning position) but only 58% at node 1 (the backup renewal position if node 2 fails), compared to 91% viability at nodes 3-4 (positions that would be removed in standard spur pruning), with the lower basal viability creating training system challenges—either accepting occasional gaps from dead spur positions, or retaining slightly longer spurs (3 buds instead of 2) to provide backup despite the less compact structure, demonstrating how basal bud viability constrains practical pruning options.
Basal bud viability quality is a bud viability quality that represents the living status of buds at the base of canes near their point of attachment to older wood, characterizing the survival of these critical buds that must be viable for spur pruning systems and that provide the renewal options essential for maintaining vine structure and training system integrity over time. Within the BFO framework, this quality specifically addresses the positional dimension of bud viability, distinguishing the survival of basal-positioned buds from buds at middle or distal cane positions while recognizing that basal bud viability has special importance for training system maintenance because these positions provide the renewal wood that replaces aging spurs and maintains the permanent vine structure, while loss of basal viability may force training system modifications or acceptance of structural deterioration. The ontological significance lies in basal bud viability determining the long-term vine structural options, where viable basal buds enable the annual renewal cycle of spur pruning systems (replacing last year's spur with growth from basal positions), the development of replacement shoots for damaged cordons, and the training flexibility for adjusting vine architecture, while loss of basal viability removes these options and constrains future vine management. The positional vulnerability of basal buds shows variable patterns depending on cultivar, cane position, and winter conditions, with some situations showing lower basal viability (buds less mature at these positions, or colder microclimate near ground) and others showing higher basal viability (better protection from snow cover, or better cane maturity promoting basal bud hardiness), creating the site-specific and cultivar-specific patterns that affect training system suitability. The practical implications of basal bud viability include spur pruning feasibility (requiring viable basal buds to produce renewal shoots), cordon training sustainability (requiring basal positions to remain productive over multiple years), and recovery options following severe damage (renewal from basal positions being essential for vine reconstruction). Understanding basal bud viability quality enables assessment of training system sustainability, evaluation of renewal options following damage, and long-term structural planning based on the viability patterns at critical basal positions.
Basal_Bud_Viability_Quality addresses the survival of buds at cane bases critical for spur pruning and vine renewal. This quality determines training system sustainability and structural recovery options. The class supports long-term vine management through basal bud viability characterization at renewal-critical positions.
Basalknospenvitalität
Erneuerungszonen-Vitalität
Sporpositionsvitalität
base bud viability
proximal bud survival
proximales Knospenüberleben
renewal zone viability
spur position viability
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 267-330 Winkler AJ et al. (1974) General Viticulture. University of California Press, Berkeley, pp. 138-165
Basal Bud Viability Quality
A basal latent bud at node 2 on a winter-pruned cane of Vitis vinifera cv. Chardonnay following a severe spring frost event that killed 85% of primary buds at nodes 4-10, breaking dormancy 10 days after the frost event, developing into a vigorous vegetative shoot with 18 leaves but no inflorescences, reaching 120cm length by mid-summer, providing photosynthetic capacity for reserve accumulation and bud differentiation for the following season when normal cropping can resume from buds formed on this compensatory growth.
A latent bud located at or near the base of annual canes or one-year-old wood, typically at nodes 1-3 from the origin point on the cordon or trunk, characterized by low fruitfulness due to shade exposure during critical inflorescence differentiation period in the previous season, small bud size, tight scale coverage, and strong suppression by apical dominance from distal buds under normal growing conditions. Basal latent buds serve as insurance against spring frost damage to more distal fruiting buds, breaking dormancy and developing when primary buds are killed, though resulting shoots typically produce little or no crop and require vigorous vegetative growth to establish productive fruiting positions for subsequent seasons, representing a yield sacrifice in the damage year but enabling canopy recovery and future production.
Basal latent buds are typically left on canes during winter pruning as "count buds" or insurance buds, with pruning formulas often specifying retention of 2-3 basal nodes beyond the desired fruiting node count (e.g., "30+10" meaning 30 fruiting nodes plus 10 basal nodes). The low fruitfulness of basal buds results from shading during June-July of the previous season when inflorescence primordia differentiate, as basal positions are typically shaded by developing canopy. Some training systems (e.g., head training, gobelet) intentionally utilize basal buds for annual fruiting wood production, accepting lower yields per bud in exchange for simplified pruning and traditional vine architecture.
Basisknospe
basal node latent bud
basale latente Knospe
base bud
count bud
Based on Dunn & Martin (2000) "A functional association in Vitis vinifera L. cv. Cabernet Sauvignon between the extent of primary branching and the number of flowers formed per inflorescence" Australian Journal of Grape and Wine Research 6=88-92 Sánchez & Dokoozlian (2005) "Bud microclimate and fruitfulness in Vitis vinifera L." American Journal of Enology and Viticulture 56=319-329 May (2004) "From bud to berry, with special reference to inflorescence and bunch morphology in Vitis vinifera L." Australian Journal of Grape and Wine Research 6=82-98.
Basal Latent Bud
A basal leaf at node 3 on a fruiting shoot of Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, California, measuring 16.8 cm length by 15.5 cm width representing 28% larger area than median leaves at nodes 8-10 on the same shoot, having achieved full expansion at 4 weeks post-budbreak compared to 6-7 weeks for median position leaves, exhibiting sun leaf anatomical characteristics with three palisade layers totaling 135 μm and thick cuticle of 6.2 μm developed during early-season high light exposure before upper canopy development, demonstrating peak photosynthesis rate of 13.5 μmol CO₂ m⁻² s⁻¹ during May contributing approximately 65% of total shoot photosynthate production supporting shoot extension and fruit set, experiencing progressive shading as upper canopy develops reducing light availability from 85% full sun in May to 35% by July creating functional decline, showing beginning senescence symptoms in early August including slight chlorosis and reduced photosynthetic capacity to 6.2 μmol CO₂ m⁻² s⁻¹, measured at shoot BBCH stage 81 (beginning of berry ripening), representing typical basal leaf lifecycle with early critical contribution followed by declining function under canopy shading.
A leaf positioned in the lower portion of a shoot occupying nodes 1-5 in Vitis vinifera, characterized by early emergence during initial shoot development typically within the first 1-3 weeks following budbreak, larger dimensions compared to median and apical leaves on the same shoot often 15-35% greater lamina area reflecting developmental priority and extended expansion duration, earlier achievement of maturity compared to distal leaves typically reaching full expansion 3-5 weeks post-budbreak, anatomical features often showing sun leaf characteristics if shoots are vertically oriented exposing basal regions or shade leaf characteristics if basal zones become shaded by developing upper canopy, disproportionately high photosynthetic contribution during early season before mid and upper canopy development providing the primary carbohydrate source supporting initial shoot extension and inflorescence development, progressive functional decline through the season as upper canopy develops creating shading of basal zones, and eventual senescence often occurring earlier than mid-canopy leaves particularly in dense canopies where severe basal shading reduces photosynthetic capacity below maintenance respiration thresholds. Basal leaves play critical roles in early season vine carbon economy, emerging from overwintered buds containing preformed leaf primordia that developed during the previous growing season, providing immediate photosynthetic capacity following budbreak before new primordia initiated from apical meristems can develop and contribute, supporting the energy-demanding processes of shoot extension, inflorescence development, and initial fruit set that occur during the 4-8 week period following budbreak when basal leaves represent the majority of functional photosynthetic surface area.
Basal leaves are distinguished from median and apical leaves by position (nodes 1-5), early emergence timing, typically larger size, and characteristic functional trajectory with early peak contribution followed by progressive decline. The exact node range defining basal zone varies somewhat with shoot vigor and cultivar, with vigorous shoots potentially extending basal designation to nodes 1-6 while weak shoots may limit it to nodes 1-3. Basal leaves emerging from preformed primordia differ fundamentally from leaves initiated from apical meristems during current-season growth, with preformed leaves showing faster development and earlier maturity. Environmental conditions at budbreak and during early shoot development critically determine basal leaf characteristics, with cool wet springs producing large thin shade-type leaves while warm dry conditions create smaller thick sun-type leaves. Basal leaf management through selective removal (basal leaf removal, leaf plucking) represents common viticultural practice for improving cluster sun exposure, reducing disease pressure, and modifying fruit composition, with timing and extent of removal requiring careful consideration of impacts on vine carbon balance and fruit development.
basales Blatt
base position leaf
early-emerging leaf
früh-emergierendes Weinblatt
lower shoot leaf
unteres Spross-Blatt
Based on Smart (1985) "Principles of grapevine canopy microclimate manipulation with implications for yield and quality" American Journal of Enology and Viticulture 36=230-239 Poni et al. (2006) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 57=397-407 Palliotti et al. (2011) "Vine performance and grape composition as affected by early-season source limitation induced with anti-transpirant and shading in two red Vitis vinifera L. cultivars" Australian Journal of Grape and Wine Research 17=368-375 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 142-178.
Basal Leaf
A basal lobe representing the right basal lobe on a mature leaf at node 6 of Vitis vinifera cv. Grenache in Châteauneuf-du-Pape, France, positioned at the leaf base immediately flanking the petiolar junction, vascularized by the basal primary lateral vein branching from median vein at 78-degree angle only 1.2 cm from petiolar junction, measuring 2.8 cm maximum width and 2.2 cm length representing the smallest lobe on this leaf, weakly separated from lower lateral lobe by shallow lower right lateral sinus of 0.8 cm depth creating indistinct boundary, lobe extending toward and contributing to the petiolar sinus configuration, basal lobe margins forming the right boundary of the closed petiolar sinus with margin approaching but not quite meeting the left basal lobe margin, lobe apex showing rounded configuration without sharp point, lobe area of 5.2 cm² representing 9% of total blade area substantially smaller than lateral lobes (14-19%) and apical lobe (28%), moderate basal lobe development creating moderately broad leaf base characteristic of this cultivar, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing typical basal lobe morphology affecting petiolar sinus configuration diagnostic for cultivar identification.
A lobe positioned at or near the leaf base flanking the petiolar junction in Vitis vinifera, characterized by proximal position at the blade base immediately adjacent to petiole attachment, vascularization by the most basal pair of primary lateral veins branching from the median vein at wide angles typically 65-90 degrees near the petiolar junction, generally smallest lobe size typically representing 8-15% per lobe of total blade area, separation from lateral lobes by lower lateral sinuses or indistinct transition zones, variable development ranging from well-defined distinct lobes creating broad leaf bases to weakly developed or absent in narrow-based cultivars, relationship to petiolar sinus configuration with basal lobe development affecting petiolar sinus shape (open, closed, or overlapping), and cultivar-specific characteristics including presence, size, degree of development, and spatial relationship to petiolar sinus serving as important diagnostic features. Basal lobes represent the most proximal lobe structures, developing at the blade base creating the characteristic configuration around the petiolar junction, exhibiting highly variable expression from prominent development in broad-based deeply dissected cultivars to minimal or absent expression in narrow-based weakly lobed varieties, and affecting overall leaf base architecture including petiolar sinus shape which serves as primary diagnostic character.
Basal lobes are distinguished from lateral lobes by proximal position at blade base adjacent to petiolar junction and typically smallest size. Development varies dramatically among cultivars from prominent well-developed basal lobes creating broad leaf bases (Grenache, some Muscat types) to minimal or absent basal lobes creating narrow bases (Sauvignon Blanc, Riesling). The relationship between basal lobe development and petiolar sinus configuration is critical, with basal lobe size and orientation determining whether petiolar sinus is open (lobes diverging), closed (lobes meeting), or overlapping (lobes crossing). Strong basal lobe development with convergent growth creates closed or overlapping sinuses while weak development with divergent orientation creates open sinuses. Size subordination to all other lobe types is typical, with basal lobes representing the smallest distinct lobe structures when present. Separation from lateral lobes often shows indistinct boundaries with shallow or absent lower lateral sinuses creating integrated basal-lateral lobe complexes. Bilateral arrangement creates paired basal lobes flanking the petiolar junction, with asymmetries occasionally producing unequal basal lobe development. Functional contribution involves relatively small area provision typically 15-25% combined area, with greater importance for mechanical integration at the critical petiole-blade junction. Ampelographic utility is substantial through effects on petiolar sinus configuration serving as primary diagnostic character, with basal lobe presence, size, and orientation critically determining sinus shape.
Basallappen
Petiolarlappen
base lobe
petiolar lobe
proximal lobe
proximaler Lappen
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 1-57 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 33-78.
Basal Lobe
In Grenache vineyards of Châteauneuf-du-Pape at BBCH 55-57, the 2-3 basal primary inflorescence branches (positions 1-3 among 20-25 total primary branches) can reach extraordinary lengths of 7-10 cm and bear 7-10 secondary branches each, contributing 200-350 flowers to the total inflorescence complement of 800-1200 flowers and creating the characteristic broad, winged cluster morphology. Mourvèdre in Bandol exhibits similarly exaggerated basal primary branches of 6-9 cm forming pronounced lateral lobes. Conversely, in compact-clustered Gewürztraminer (Alsace), even basal primary branches remain relatively modest at 2.5-4 cm length with 2-4 secondary branches, though still exceeding upper branch dimensions. Viticultural cluster thinning protocols sometimes selectively remove basal primary branches to reduce cluster width and improve spray penetration.
A basal primary inflorescence branch is a primary inflorescence branch located in the absolute basal region of the main inflorescence rachis, typically the first 2-4 primary branches emerging nearest the peduncle attachment point, often exhibiting the maximum absolute length and sub-branching elaboration among all inflorescence branches while sometimes showing distinctive morphological features related to their proximity to the shoot axis. Basal primary branches occupy positions 0-25% of the distance from the rachis base to apex (measured at BBCH 55), emerging from rachis nodes within 2-12 mm of the peduncle insertion point. These branches represent the earliest-initiated primary branches during inflorescence primordium development and are the first to become morphologically distinct during bud burst and early shoot growth. Branch length typically ranges from 4-10 cm in extensively branched cultivars (up to 12 cm in extreme cases like some Muscat types), with basal diameter often 1.8-3 mm, representing the maximum dimensions observed anywhere in the inflorescence architecture. Basal primary branches characteristically bear the greatest number of secondary branches (5-10 secondary branches per basal primary branch in loose-clustered cultivars 3-6 in moderate types), with secondary branches extensively subdivided into tertiary and occasionally quaternary branches. This extreme architectural elaboration results in basal primary branches bearing 30-80 flowers each in loose-clustered cultivars, though this flower number may not proportionally convert to berry number due to differential fruit set rates along the rachis gradient. The exceptionally elongated, well-subdivided character of basal primary branches creates the pronounced shoulder, wing, or lateral lobe structure of mature clusters in many cultivars, particularly in Grenache, Mourvèdre, and some Muscat varieties where basal branches may approach or equal the rachis length itself. In some cultivars and under certain growing conditions, the lowest 1-2 basal primary branches may exhibit partial vegetative characteristics (slightly larger diameter, enhanced trichome density, rare occurrence of tendril-like structures), reflecting their developmental proximity to the shoot meristem and intermediate position in the flower-to-shoot morphological gradient. Vascular anatomy of basal primary branches shows the largest xylem vessel diameters observed in inflorescence tissues (40-70 μm), with enhanced phloem development supporting intensive resource delivery to the extensive flower complement. Basal branch development initiates earliest during BBCH 53-55, often 3-6 days before apical primary branch emergence, and proceeds most rapidly, achieving maximum length by BBCH 57-59.
Basal_Primary_Inflorescence_Branch represents the most architecturally elaborate primary branches and should be distinguished from Lower_Primary_Inflorescence_Branch (which includes the subsequent quartile of branches more distant from the peduncle). The 'basal' designation specifically refers to the absolute base of the rachis, typically the first 2-4 primary branches. These branches show maximum developmental potential and occasionally exhibit transitional morphological features reflecting the inflorescence-to-shoot developmental continuum. Extreme basal branch development is a key distinguishing feature of loose-clustered cultivar groups and contributes substantially to total cluster weight. In some precision viticulture applications, basal branch removal at BBCH 55-59 is practiced to modify cluster architecture and reduce final compactness, though this reduces yield proportionally to the substantial flower number borne on these branches.
basaler Primärast
basalster Blütenstandsast
bottom primary branch
proximal primary branch
unterster Hauptast
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 88-92 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 165-173 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 76-82 Galet, P. (2000). A Practical Ampelography, pp. 47-50.
Basal Primary Inflorescence Branch
A basal secondary inflorescence at node 5 on a short 6-node fruiting spur of Vitis vinifera cv. Chardonnay in Champagne, France, measuring 12.5 cm length, positioned very near the spur base only 12 cm from the spur attachment point to the cordon creating extreme basal location, located on a short spur where node 5 represents the penultimate node before the terminal, receiving approximately 52% of full sunlight with substantial shading from the developing shoot growth and foliage above, exhibiting compound paniculate architecture with central rachis bearing 3 orders of branching, rachis diameter of 2.6 mm at base, bearing approximately 340 flowers, positioned in the basal zone experiencing the protected microenvironment near the permanent wood with moderated temperatures and reduced wind exposure, showing dimensions approximately 71% of the primary inflorescence at node 3 (17.5 cm, 520 flowers), developmental stage at BBCH 61 (beginning of flowering) showing typical 2-3 day delay, basal position creating both shading disadvantages reducing photosynthetic capacity and protection advantages reducing environmental stress, expected to contribute approximately 28% of spur yield, measured in late May during early bloom on this cool-climate spur-pruned system, representing typical basal secondary on short spurs where node 5 creates extreme basal positioning.
A secondary inflorescence positioned at or very near the shoot base in Vitis vinifera, characterized by location at node 5 or occasionally node 4 creating very basal positioning on the shoot, proximity to the shoot origin at the cane or spur attachment point, occurrence on short shoots or spurs where node 5 represents near-basal location given limited total node number, position creating extreme basal placement relative to shoot architecture, reduced light exposure typical of basal positions often receiving 40-60% of full sunlight due to shading from shoot growth above, dimensions typically 9-14 cm representing variable development, moderate to reduced branching complexity, and functional characteristics reflecting the basal position including potential shading disadvantages but also stable attachment and reduced exposure to wind and environmental extremes. Basal secondary inflorescences represent the extreme basal positioning variant, occurring primarily on short shoots and spurs where the node 5 position creates near-base location, exhibiting characteristics reflecting the basal environment including shading but also protection, and contributing to yield with potential quality implications from reduced sun exposure.
Basal secondary inflorescences represent the extreme basal positioning, distinguished from lateral secondaries by very near-base location particularly evident on short shoots and spurs. The basal designation emphasizes proximity to the shoot base and permanent wood rather than simply node 5 position, with the basal character emerging from short shoot architecture. Position at node 5 on 6-8 node spurs creates near-terminal location in absolute terms but extreme basal location relative to the short shoot structure and distance from shoot apex. Light exposure is typically reduced with 40-60% full sun common due to shading from all shoot growth above, creating potentially limiting conditions for optimal development. Microenvironmental characteristics include proximity to permanent wood potentially moderating temperature extremes, reduced wind exposure providing protection from mechanical stress, and potentially elevated humidity from reduced air circulation. Size shows variable development with some basal secondaries showing moderate dimensions while others exhibit reduced development depending on cultivar, vigor, and environmental conditions. Branching complexity is typically moderate with 3 orders common. Occurrence is primarily on short spurs and short shoots where the standard node 5 position creates extreme basal placement, while longer shoots show node 5 as mid-shoot creating lateral rather than basal character. Developmental timing follows typical secondary patterns. Yield contribution is moderate with potential fruit quality implications from reduced sun exposure potentially affecting sugar accumulation, color development, and flavor compound synthesis. Management considerations include assessment of sun exposure and potential for removal if shading is excessive, though basal secondaries on well-exposed spurs may provide acceptable quality fruit warranting retention.
Sprossbasis-Zweitinfloreszenz
basale sekundäre Infloreszenz
extremale Basal-Sekundärrispe
extreme basal inflorescence
near-base secondary
shoot-base secondary
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 333-368 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 479-518.
Basal Secondary Inflorescence
A basal shoot on Vitis vinifera cv. Cabernet Sauvignon arising from trunk at 15cm above soil surface, activated following severe Eutypa dieback killing upper trunk and bilateral cordons, shoot developed to 245cm length by late July with 28 leaves, 6 tendrils, no inflorescences (typical sterility of latent bud origin), extremely vigorous growth (6-8cm per day during peak elongation) reflecting release from apical dominance and concentration of whole-vine resources into single basal shoot, representing critical recovery mechanism enabling vine survival and eventual structural reconstruction, this shoot deliberately retained and trained as replacement trunk over 2-3 year period developing new head and bilateral cordons at appropriate height, converting catastrophic disease damage into managed renewal opportunity avoiding vine removal and replanting costs.
A shoot arising from latent buds at or near the base of the vine, characterized by origin from trunk, head, or below-ground positions rather than from current-season fruiting wood, typically vigorous growth reflecting release from apical dominance suppression, origin from buds that may be years or decades old having remained dormant through multiple growing seasons, and function as recovery mechanism following damage, renewal source for training system reconstruction, or problematic growth requiring removal depending on context and management objectives. Basal shoots represent adaptive growth response enabling vines to recover from catastrophic damage (frost killing above-ground portions, disease destroying established structure, mechanical damage), provide architectural renewal options (training new trunks, arms, or cordons from basal positions), or create management challenges when arising spontaneously without damage triggers (excessive vigor, hormonal imbalances, rootstock suckering), requiring assessment whether basal shoots serve beneficial purposes warranting retention or represent problems requiring removal.
Basal shoots serve dual roles as both beneficial recovery/renewal mechanisms and problematic unwanted growth depending on context. Beneficial contexts include damage recovery (frost, disease, mechanical injury destroying established structure requiring basal regrowth for survival), planned renewal (training new trunks or structures from basal positions as alternative to grafting or replanting), and architectural modifications (developing additional trunks, changing training systems). Problematic contexts include spontaneous suckering (rootstock shoots requiring removal, unwanted scion shoots creating canopy complications), excessive vigor manifestation (basal shoot proliferation indicating imbalanced conditions), and resource competition (basal shoots diverting resources from productive canopy if not intentionally utilized). Management involves identifying basal shoot origin and purpose (recovery, renewal, or problem), making retention versus removal decisions based on assessment, and when retaining basal shoots for renewal, implementing appropriate training protocols developing desired structures over multi-year periods while managing vigor and preventing interference with remaining productive portions.
Stammtrieb
Stockausschlag
basaler Trieb
base shoot
renewal shoot from base
sucker
trunk shoot
water shoot
Based on Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press, pp. 85-105 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 113-145.
Basal Shoot
Pinot Noir in Oregon shows basal shoot lignification from late June through late July at BBCH 31-35 spanning 4-5 weeks, initiating in lowest 3-4 internodes when elongation completes with lignin content increasing to 14-18% dry weight and starch accumulating to 12-16% dry weight creating brown woody basal structure under cool climate conditions of 18-24°C mean temperature, adequate moisture, and moderate fertility typical of this moderate-vigor early-to-mid season cultivar requiring timely maturation for winter survival
Basal shoot lignification represents the initial maturation phase where oldest shoot regions undergo secondary wall deposition and lignin incorporation establishing first lignified woody structures in developing shoot, providing stable mechanical foundation and initiating reserve accumulation that will extend acropetally through shoot during subsequent maturation phases. This critical developmental stage occurs in lowest 3-5 internodes of shoot during mid-summer corresponding approximately to flowering through early fruit set period at BBCH 31-35, typically beginning 4-6 weeks after bud burst when basal internodes have completed elongation and cells transition to differentiation and secondary wall synthesis. Basal lignification initiation occurs when internode elongation in basal regions completes typically by late May-early June in temperate climates, with cells in vascular tissues, fibers, and adjacent parenchyma beginning secondary wall deposition and lignin biosynthesis transforming primary walled herbaceous tissue to secondary walled woody structure. Secondary wall synthesis begins in xylem vessel elements requiring rigid walls for water transport function, with S1, S2, and S3 wall layers depositing with characteristic microfibril orientations providing tensile and compressive strength, followed by lignin monomer synthesis through phenylpropanoid pathway producing p-coumaryl, coniferyl, and sinapyl alcohols. Lignin polymerization proceeds through oxidative coupling reactions catalyzed by peroxidases and laccases, creating three-dimensional polymer network that cross-links wall polysaccharides and provides hydrophobic character, with lignin content in basal internodes increasing from negligible to 12-18% dry weight over 3-4 week initial lignification period. Lignified tissue extent in basal regions shows progressive expansion from initial vascular tissue lignification extending into surrounding fibers and parenchyma cells, creating increasingly rigid internode structure with brown coloration replacing green as chlorophyll degrades and phenolic compounds accumulate. Mechanical property changes during basal lignification include increasing tensile strength from 8-15 MPa in recently completed elongation to 35-60 MPa in substantially lignified tissue, declining flexibility as rigid secondary walls prevent bending, and increasing resistance to compression providing stable base supporting developing shoot. Vascular tissue maturation in basal regions includes vascular cambium initiation between primary xylem and phloem, secondary xylem production internally increasing vessel number and water transport capacity to 3-5 mg per second per basal internode, and secondary phloem formation externally enhancing assimilate translocation from photosynthesizing canopy. Periderm initiation begins in basal internodes during lignification with cork cambium forming in subepidermal layers, producing cork cells externally creating protective brown bark replacing green epidermis, establishing protective surface layer reducing water loss and providing pathogen barrier. Reserve accumulation initiates in basal internodes as lignification proceeds with starch synthesis converting excess photosynthate to storage form, increasing starch concentration from negligible during growth to 10-18% dry weight in basal regions by mid-summer when lignification substantially completes. Metabolic transition in basal regions involves declining respiration from growth levels 10-15 fold above dormancy to early maturation levels 4-6 fold above dormancy as growth ceases and maintenance requirements dominate, with continued decline toward dormancy levels as maturation completes. Protein composition shifts during basal lignification from growth-related proteins including cell cycle regulators and wall synthesis enzymes to maintenance proteins, storage proteins accumulating in parenchyma cells, and defense proteins providing pathogen resistance. Water content in basal internodes declines during lignification from 75-80% fresh weight in recently elongated tissue to 60-65% in substantially lignified tissue, reflecting increased wall material deposition, reduced metabolic activity, and beginning dehydration associated with maturation. Nitrogen dynamics show declining concentration in basal tissues from 2.0-2.5% dry weight after elongation to 1.2-1.8% as structural carbohydrates accumulate diluting nitrogen-rich proteins, with mobile nitrogen potentially remobilizing toward active growth regions or storage tissues. Hormonal regulation of basal lignification involves declining auxin from reduced apical production and transport as basal regions distance from apex, increasing abscisic acid promoting maturation processes and beginning stress tolerance development, and ethylene possibly involved in coordinating metabolic transitions. Temperature affects basal lignification rate with warm mid-summer conditions 22-28°C promoting rapid progression completing substantial lignification within 3-4 weeks, while cooler conditions extend duration though typically not limiting as process proceeds across wide temperature range. Light intensity affects basal lignification through effects on whole-shoot photosynthetic production providing carbon for lignin synthesis and reserve accumulation, with adequate light supporting normal progression while extreme shade may limit carbon availability slowing maturation. Water availability affects basal lignification with moderate water stress potentially promoting earlier initiation and enhanced progression as stress signals trigger maturation responses, while severe stress may impair processes through limiting photosynthetic production and metabolic function. Nutrient availability affects basal lignification with adequate nitrogen supporting normal progression as growth in basal regions has completed and nitrogen requirements shift toward upper shoot zones, while excessive nitrogen may slow lignification prolonging metabolic activity. The process typically requires 3-5 weeks from initiation to substantial lignification in basal 3-5 internodes depending on environmental conditions and cultivar characteristics, with warm conditions and moderate stress promoting rapid complete lignification. Cultivar variation appears in basal lignification timing with early-maturing varieties showing earlier initiation and rapid progression while late-maturing types show delayed initiation, extent of lignification in basal regions with some varieties showing thorough complete lignification while others maintain less complete maturation, and reserve accumulation intensity affecting basal tissue starch concentrations. Regional timing shows basal shoot lignification initiating June-July in warm climates, July in temperate regions, and late July-early August in cool areas, with progression corresponding to summer temperature patterns and shoot growth stage. Viticultural management considerations include recognizing basal lignification as indicator of normal shoot development and resource adequacy, understanding that inadequate basal lignification indicates problems with vine health or excessive late vegetative growth, monitoring lignification progress assessing vine maturation status and dormancy preparation, and ensuring management promotes timely basal maturation through appropriate water and nitrogen management. Research applications include analyzing mechanisms initiating lignification including developmental and environmental signals, quantifying lignin biosynthesis and deposition patterns during maturation, evaluating carbon partitioning between growth and storage during transition periods, assessing environmental effects on maturation timing affecting reserve accumulation and dormancy preparation, and documenting cellular and molecular changes during herbaceous-to-woody tissue transformation.
Basal_Shoot_Lignification is first of four Shoot_Lignification subphases, distinguished by lignification in oldest shoot regions establishing initial woody structure. Represents earliest maturation occurring during mid-summer active growth period. Transitions to Progressive_Shoot_Lignification as maturation extends acropetally. Critical for establishing stable shoot base and initiating reserve accumulation. Timing indicator of vine development status and resource adequacy.
Grundtriebverholzung
Triebbasisausreifung
basal maturation
basale Holzbildung
basale Triebverholzung
lower shoot lignification
shoot base woodification
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 282-290 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 215-222 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Basal Shoot Lignification
A base attached secondary inflorescence at node 4 on a short spur of Vitis vinifera cv. Gewürztraminer in Alsace, France, positioned at the anomalously low node 4 location creating extreme basal attachment only 6.5 cm from the spur base and permanent wood junction, measuring 9.8 cm length representing reduced dimensions, occurring through developmental variation with shortened basal internodes and early inflorescence initiation creating the exceptional node 4 positioning (compared to standard node 5 secondary placement), exhibiting moderately developed architecture with central rachis bearing 2-3 orders of branching, rachis diameter of 2.0 mm at base representing reduced construction, bearing approximately 185 flowers, positioned in the extreme basal zone experiencing maximum shading with approximately 45% full sunlight from complete overhead canopy coverage, showing dimensions substantially reduced from the primary at node 3 (15.2 cm, 420 flowers) representing 36% length reduction and 56% flower number reduction, extreme basal position creating both maximum shading disadvantage and maximum protection from wind and temperature extremes, developmental stage at BBCH 59 (inflorescence fully developed, flowers separating) showing 5-day phenological delay, expected to contribute approximately 18% of spur yield, measured in late May during pre-bloom period, representing exceptional base attached secondary with anomalous node 4 positioning creating extreme basal character.
A basal secondary inflorescence characterized by attachment very near or at the shoot base itself rather than standard node positions in Vitis vinifera, characterized by position at node 4 or occasionally lower nodes creating extreme basal attachment, exceptional proximity to the shoot origin point at cane or spur attachment, occurrence through developmental variations including early inflorescence initiation, shortened internodes, or other morphological modifications creating the near-base attachment, very basal location often within 5-10 cm of the permanent wood junction, dimensions typically 8-13 cm representing reduced development reflecting the extreme basal position and developmental variations, reduced branching complexity often showing 2-3 orders, and functional characteristics determined by the extreme basal environment including maximum shading from all shoot growth above but also maximum protection and proximity to permanent wood resources. Base attached secondary inflorescences represent exceptional basal positioning through attachment point rather than standard node location, occurring through developmental variations creating the extreme basal placement, exhibiting characteristics reflecting both the non-standard attachment and the extreme basal environmental context.
Base attached secondary inflorescences represent exceptional positioning through anomalous attachment points, distinguished from node position basal variants by non-standard node location (often node 4 or lower) rather than standard node 5 placement. The base attached designation emphasizes the mechanism creating extreme basal character through actual attachment point proximity to the shoot base. Position at node 4 or lower represents departure from the standard two-node gap pattern, resulting from developmental variations including shortened basal internodes compressing node spacing, early inflorescence primordium initiation at non-standard positions, or other morphological modifications. Extreme basal location within 5-10 cm of permanent wood creates the most basal positioning possible for inflorescences, maximizing proximity to trunk or arm resources but also maximizing shading from all shoot growth above. Size reduction is typically pronounced with 30-50% reduction from primaries common, reflecting both the developmental anomalies and extreme basal environmental limitations. Branching complexity is often reduced with 2-3 orders typical compared to 3-4 for standard secondaries. Flower number may be substantially reduced with <200 flowers possible in extreme cases. Occurrence is uncommon representing developmental variations rather than standard patterns, with some cultivars or specific vines showing tendencies for base-attached development while most vines show standard node 5 secondaries. Environmental context creates maximum shading with 40-50% full sun typical from complete canopy overhead, but also maximum protection and stable microclimate. Management implications include high probability for removal during cluster thinning due to extreme shading, reduced size, and potential quality limitations, though exceptional base-attached secondaries on well-exposed spurs may warrant retention if vine balance permits.
Sprossbasis-angeheftete sekundäre Infloreszenz
anomale basale Sekundärrispe
anomalous basal secondary
extreme Basal-Anheftung Blütenstand
extreme basal attachment secondary
shoot-base-attached inflorescence
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 369-405 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 559-598.
Base Attached Secondary Inflorescence
Petiolar sinus shape in Merlot exemplifies the closed to overlapping configuration characteristic of many Bordeaux varieties, with the two basal lobes approaching or overlapping to create a lyre-shaped sinus with parallel or convergent sides this distinctive base shape, documented in OIV descriptor 079 as category 7 (lobes overlapping), distinguishes Merlot from its parent Cabernet Franc which typically displays a more open U-shaped sinus, and provides a reliable field identification character when leaf blade overall dimensions are ambiguous.
Base_Shape_Quality represents the morphological quality characterizing the proximal configuration of grapevine organs at their point of attachment or origin. The base constitutes the region where an organ connects to its supporting structure, and its shape affects both mechanical attachment properties and the geometry of vascular connections. In leaves, base shape describes the configuration of the blade where it meets the petiole, with particular attention to the petiolar sinus, the indentation where the two basal lobes approach or overlap. Petiolar sinus shape is a primary ampelographic character captured in OIV descriptors 079-081, with categories describing the degree of opening (wide open, open, closed, overlapping) and the shape of the sinus floor (flat, U-shaped, V-shaped, lyre-shaped). In berries, base shape characterizes the pedicel attachment region, which may be truncate with an abrupt transition, rounded with smooth curvature, or necked with a distinct narrowing at the attachment point. Berry base shape affects the ease of berry detachment, relevant for both harvest method selection and postharvest handling. In shoots, base shape describes the configuration at the node where the shoot originates. Base shape assessment employs visual matching to standardized descriptors, measurement of sinus angles and depths, or outline analysis of the basal region. The base region often provides more stable cultivar characters than the apex because it is less subject to environmental modification during the extended growth period.
Base_Shape_Quality addresses the proximal attachment region of organs and is positioned parallel to Apex_Shape_Quality under Shape_Quality. For leaves, the base encompasses the petiolar sinus and the basal lobes flanking it OIV descriptors 079-081 provide standardized assessment protocols. For berries, the base is the pedicel attachment zone. The contrast between apex and base shape can be diagnostically useful some cultivars show distinctive combinations (e.g., acute apex with overlapping base). Base shape assessment should specify whether the descriptor refers to the attachment point specifically or to the broader basal region of the organ.
Ansatzform
Basisform
attachment region shape
basal configuration
basale Konfiguration
insertion shape
proximal shape
proximale Gestalt
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 57-60 Chitwood DH et al. (2014) A Modern Ampelography: A Genetic Basis for Leaf Shape and Venation Patterning in Grape. Plant Physiology 164(1): 259-272
Base Shape Quality
A Merlot clone 181 vine in the ENTAV foundation vineyard at Domaine de l'Espiguette bears a Basic_Mother_Plant_Role, derived from virus-tested Pre-Basic nuclear stock, maintained in an isolated block with vector management (insecticide applications targeting mealybugs and leafhoppers), and tested every three years by ELISA for GFLV, GLRaV-1, GLRaV-3, GVA, and GVB—its budwood distributed to licensed French nurseries for Certified multiplication. A 110 Richter rootstock plant in a California-certified foundation block serves in a Basic_Mother_Plant_Role, providing hardwood cuttings to commercial nurseries operating under CDFA certification protocols. A Grüner Veltliner clone 17 maintained by the Austrian AGES agency bears a Basic_Mother_Plant_Role, supporting the Austrian wine industry's access to tested planting material.
A certified mother plant role that inheres in a grapevine maintained at the intermediate tier (Basic/Foundation) of the certification hierarchy, derived directly from Pre-Basic material and serving as the multiplication source for Certified category production. Basic mother plants inherit the verified health status of their Pre-Basic source and are maintained under conditions preserving this status—physical isolation from non-certified plantings, vector management programs, and regular testing (typically triennial) to confirm continued freedom from regulated pathogens. Basic material may be multiplied through a limited number of generations (typically 3-4) before requiring renewal from Pre-Basic stock, preventing genetic drift and accumulated pathogen risk over extended propagation chains. Basic mother plants are typically maintained by certification authorities, research institutions, or specially licensed nurseries with the infrastructure and expertise to meet the rigorous requirements, providing tested material to commercial nurseries for final multiplication into Certified category planting stock.
Basic_Mother_Plant_Role captures the critical intermediate tier that bridges nuclear stock preservation (Pre-Basic) and commercial multiplication (Certified), balancing the need for tested material volume against the cost of maintaining Pre-Basic-level testing intensity. Basic material undergoes less frequent testing than Pre-Basic (triennial versus annual or continuous) but more frequent than Certified (which may rely primarily on visual inspection). Generation limits prevent unlimited multiplication from a single Basic source, requiring periodic return to Pre-Basic material this "pyramid refreshment" maintains genetic fidelity and reduces accumulated pathogen risk. The Basic tier's role in distributing tested material to multiple nurseries creates the bottleneck controlling quality throughout the certification system—pathogen introduction at the Basic level would propagate through all derived Certified material.
Basis-Mutterstock
Basismaterial
Grundstock
Vermehrungsgrundlage
base category plant
basic mother vine
foundation mother plant
source multiplication plant
European Union (2020) Commission Implementing Directive 2020/177 Annex IV Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 Walter B, Martelli GP (1996) Sanitary Selection of the Grapevine, INRA Editions EPPO (2008) Certification schemes for pathogen-tested material of grapevine, EPPO Bulletin 38: 422-429
Basic Mother Plant Role
Basic taste profiling of Gewürztraminer table grapes at three harvest dates demonstrates sweetness increasing from 5.2 to 7.8 to 9.1 on a 10-point scale while sourness decreases from 6.5 to 4.2 to 2.8 and bitterness remains stable at 2.0-2.3, with the third harvest showing sweetness dominance that masks the characteristic Gewürztraminer bitterness detectable in earlier harvests, informing the choice of second harvest date where sweetness-sourness-bitterness balance optimizes the varietal taste profile.
Basic taste quality is a taste quality that represents any of the five primary gustatory modalities (sweetness, sourness, bitterness, saltiness, umami) that are considered irreducible perceptual categories, each mediated by distinct receptor mechanisms and perceived as qualitatively unique sensations that cannot be replicated by combinations of other basic tastes. Within the BFO framework, this quality establishes the categorical structure for taste perception, distinguishing the fundamental taste primaries from complex taste profiles that emerge from their combination, and from taste-related perceptions (astringency, cooling, pungency) that involve non-gustatory mechanisms. The ontological significance lies in basic tastes providing the building blocks for taste description and analysis, where any taste experience can be decomposed into contributions from these primary channels, enabling systematic characterization of grape and wine taste profiles through intensity rating of each basic taste. The physiological distinctiveness of basic tastes reflects their evolutionary significance, with sweetness signaling caloric value, sourness indicating acidity and potential spoilage, bitterness warning of toxins, saltiness reflecting mineral content, and umami indicating protein-rich foods, creating an innate hedonic valence (attraction to sweet and umami, aversion to bitter and sour) that influences grape and wine preference. In grape and wine context, sweetness and sourness dominate the taste profile with bitterness contributing through phenolic compounds, while saltiness plays minor role and umami remains debated though amino acid content may contribute subtle savory notes in aged wines. The independence of basic taste channels means they can be separately measured in sensory analysis and separately targeted through compositional manipulation, though perceptual interactions (sweetness suppressing bitterness, acidity enhancing freshness perception) complicate prediction of integrated taste quality from component intensities.
Basic_Taste_Quality establishes the five primary taste categories, serving as parent for Sweetness_Quality and sibling classes for the other basic tastes. This framework follows contemporary taste science recognizing five (some argue six with fat taste) distinct modalities. The class excludes taste-related sensations like astringency (tactile) and cooling/pungency (trigeminal) that are often conflated with taste in casual usage.
Basisgeschmack
Grundgeschmack
elemental taste
fundamental taste modality
fundamentale Geschmacksmodalität
primary taste
primärer Geschmack
taste primary
Chandrashekar J et al. (2006) The receptors and cells for mammalian taste. Nature 444(7117): 288-294 Running CA et al. (2015) Oleogustus: the unique taste of fat. Chemical Senses 40(7): 507-516
Basic Taste Quality
Baumé measurement for Shiraz in the Barossa Valley (2023 vintage) records harvest at 14.2° Bé, predicting approximately 14.2% vol. alcohol — near the upper boundary of the preferred range for full-bodied Australian Shiraz, with the approximate 1:1 correspondence between Baumé degrees and potential alcohol simplifying harvest decision communication. Baumé measurement for Sémillon in Sauternes during the 2024 botrytized harvest documents successive selective picking passes (tries) at increasing sugar levels: first trie at 18° Bé, second trie at 22° Bé, and third trie at 25° Bé, with the progressive concentration reflecting advancing noble rot development and berry desiccation across the 4-week harvest period.
Baumé measurement is a sugar content data entity that records the sugar concentration in grape must as determined by hydrometry on the Baumé density scale, expressed in degrees Baumé (°Bé), where the scale approximates the potential alcohol yield in percent by volume under standard fermentation conditions, historically serving as the primary sugar measurement system in French and Australian viticulture and providing a direct field estimate of the wine alcohol content expected from complete fermentation.
The Baumé scale's principal practical advantage is the near-direct correspondence between the measured value and expected alcohol content — the conversion factor of approximately 1.0° Bé ≈ 1.0% vol. potential alcohol (more precisely, the factor varies between 0.98-1.05 depending on regional convention) makes it uniquely intuitive for vineyard and cellar communication. The scale was developed by the French chemist Antoine Baumé in 1768 as a general-purpose hydrometric scale, with the specific modulus for grape must measurement defined as Bé = 145 × (1 - 1/SG) where SG is specific gravity at 20°C — though different national conventions employ slightly different modulus values (the "rationalized" Baumé uses modulus 144.32), creating minor but non-trivial discrepancies in cross-national comparisons. In French viticulture, Baumé has historically been the field standard, though INAO regulations specify sugar content in grams per liter (g/L) or potential alcohol for quality category determination. Australian viticulture adopted Baumé from the French tradition during the 19th century and it remains the dominant harvest metric, though the trend toward Brix measurement following American and New Zealand influence is gradually increasing. The relationship between Baumé and Brix is approximately 1° Bé ≈ 1.8° Brix (more precisely, Brix = Baumé × 1.796 at typical grape must compositions), enabling cross-scale reference. Like all density-based measurements, Baumé readings are affected by non-sugar solutes and require temperature correction to the reference temperature.
Baumé degree
Baumé density
Baumé hydrometer reading
Baumé reading
Baumé value
Baumé-Bestimmung
Baumé-Dichte
Baumé-Grad
Baumé-Messung
Baumé-Wert
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1, 2nd Ed, Wiley, pp.: 219-225 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 331-335 Rankine BC (2004) Making Good Wine: A Manual of Winemaking Practice for Australia and New Zealand, Pan Macmillan, pp.: 45-55
Baume Measurement
A phenological observer recording BBCH 05 for Grüner Veltliner in the Wachau on April 3 documents the wool stage (Wollestadium) where brown wool is clearly visible on the bud but green tissue has not yet emerged — the progression through stage 0 spans BBCH 00 (winter bud dormancy, pointed buds), BBCH 01 (beginning of bud swelling), BBCH 03 (end of bud swelling, buds rounded), BBCH 05 (wool stage), BBCH 07 (beginning of bud burst, green shoot tips just visible), and BBCH 09 (bud burst complete, first leaf unfolded and separated from shoot tip), with the entire stage 0 duration ranging from 2-6 weeks depending on cultivar, spring temperatures, and accumulated growing degree days above the 10°C base threshold.
A principal growth stage code designating BBCH stage 0 (codes 00-09) that encompasses the period from winter dormancy through bud swelling to the emergence of green shoot tissue from the bud scales, representing the transition from endodormancy through ecodormancy to the initiation of active vegetative growth in the grapevine annual cycle.
BBCH_0_Sprouting_Bud_Development is the first phenological milestone of the grapevine growing season and a key indicator for frost risk assessment, early-season disease management timing, and growing season length estimation. The timing of bud burst (BBCH 07-09) varies by 4-6 weeks across cultivars (early-bursting Chardonnay versus late-bursting Cabernet Sauvignon) and by 2-4 weeks across vintages at a single site due to spring temperature variation: this variability makes BBCH stage 0 timing one of the most climate-sensitive phenological indicators. The base temperature for bud development in Vitis vinifera is conventionally set at 10°C, with approximately 50-150 growing degree days (GDD base 10) accumulated from dormancy release to BBCH 09 depending on cultivar.
Austriebsstadium
BBCH stage 0
BBCH-Stadium 0
Knospenschwellen-Phase
bud development stage
sprouting stage code
Lorenz DH et al. (1995) Growth stages of the grapevine: phenological growth stages of the grapevine — Codes and descriptions according to the extended BBCH scale, Australian Journal of Grape and Wine Research 1: 100-103 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 73-95
BBCH 0 Sprouting Bud Development
A phenological record of BBCH 15 for Müller-Thurgau at the Geisenheim research vineyard on May 5 indicates that 5 leaves are unfolded and separated from the shoot tip — the progression through stage 1 proceeds from BBCH 11 (first leaf unfolded and separated from shoot tip, marking the transition from stage 0) through BBCH 12-19 in unit increments each corresponding to one additional unfolded leaf, with the rate of leaf appearance (phyllochron) averaging 5-8 days per leaf under optimal spring conditions at 15-20°C mean temperature, and the completion of stage 1 coinciding with the beginning of rapid shoot elongation at 3-8 cm per day.
A principal growth stage code designating BBCH stage 1 (codes 11-19) that encompasses the sequential unfolding and expansion of leaves along the developing shoot, from the first fully separated leaf through to the stage where nine or more leaves are unfolded, documenting the progressive establishment of the grapevine's photosynthetic canopy.
BBCH_1_Leaf_Development tracks the early canopy establishment phase that determines the vine's photosynthetic capacity for the remainder of the season. The secondary stage numbering is uniquely straightforward in stage 1 — each code increment (11, 12, 13...19) corresponds directly to the cumulative number of unfolded leaves — making this the most intuitive BBCH stage to observe and record. The rate of leaf development (phyllochron) is strongly temperature-dependent and cultivar-specific: early-developing cultivars may complete stage 1 in 3-4 weeks under warm conditions, while late cultivars in cool springs may require 5-7 weeks. Stage 1 overlaps temporally with the beginning of inflorescence emergence (stage 5) in many cultivars, as cluster primordia become visible while leaf development continues.
BBCH stage 1
BBCH-Stadium 1
Blattentfaltungs-Phase
Blattentwicklungsstadium
leaf unfolding stage
shoot leaf development code
Lorenz DH et al. (1995) Growth stages of the grapevine: phenological growth stages of the grapevine — Codes and descriptions according to the extended BBCH scale, Australian Journal of Grape and Wine Research 1: 100-103 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 95-125
BBCH 1 Leaf Development
A phenological observation of BBCH 55 for Chardonnay in Chablis on May 20 documents that inflorescences are clearly visible and fully developed with single flowers tightly pressed together — the progression through stage 5 spans BBCH 53 (inflorescences clearly visible), BBCH 55 (inflorescences fully developed, single flowers pressed together), and BBCH 57 (flowers separating, flowerhoods still closed), with the stage 5 duration typically lasting 2-4 weeks and representing the critical window for pre-bloom disease management against downy mildew (Plasmopara viticola) whose primary infection cycle targets emerging inflorescences.
A principal growth stage code designating BBCH stage 5 (codes 53-57) that encompasses the visible emergence, elongation, and pre-bloom development of grapevine inflorescences (flower clusters), from the first appearance of clearly visible inflorescence primordia through to the stage where individual flowers are fully separated and flowerhoods still closed.
BBCH_5_Inflorescence_Emergence documents the pre-bloom reproductive development phase that establishes the potential crop load for the vintage. The number and size of inflorescences visible during stage 5 provide the first quantitative basis for yield estimation, as each inflorescence contains 100-400 individual flowers depending on cultivar and prior-season bud fruitfulness: however, only 20-50% of flowers typically set fruit, making stage 5 observations an upper-bound yield estimate. BBCH stages 2, 3, and 4 are not defined for grapevine, so stage 5 follows directly from stage 1 in the coding system — in practice these stages overlap temporally as inflorescence emergence occurs while leaf development continues on actively growing shoots.
BBCH stage 5
BBCH-Stadium 5
Gescheinserscheinungs-Phase
Infloreszenz-Entwicklungsstadium
inflorescence emergence stage
pre-bloom cluster development code
Lorenz DH et al. (1995) Growth stages of the grapevine: phenological growth stages of the grapevine — Codes and descriptions according to the extended BBCH scale, Australian Journal of Grape and Wine Research 1: 100-103 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 125-145 Dry PR et al. (2004) Viticulture Volume 1: Resources, 2nd Ed, Winetitles pp: 141-167
BBCH 5 Inflorescence Emergence
A phenological record of BBCH 65 for Pinot Noir in Burgundy on June 12 documents full flowering with 50% of flowerhoods fallen — the progression through stage 6 spans BBCH 60 (first flowerhoods detaching from receptacle), BBCH 61 (beginning of flowering, 10% of flowerhoods fallen), BBCH 63 (early flowering, 30% fallen), BBCH 65 (full flowering, 50% fallen), BBCH 67 (70% fallen), BBCH 68 (80% fallen), and BBCH 69 (end of flowering, all flowerhoods fallen), with the entire flowering period typically lasting 6-14 days under optimal conditions at 20-25°C and extending to 3-4 weeks under cool, wet conditions below 15°C that promote poor fruit set (coulure) and shot berry formation (millerandage).
A principal growth stage code designating BBCH stage 6 (codes 60-69) that encompasses the period of anthesis in the grapevine, from the opening of the first flowerhoods (calyptrae) through full bloom to the completion of flowering with all flowerhoods fallen and the transition to fruit set, representing the critical reproductive phase that determines potential berry number and cluster architecture.
BBCH_6_Flowering is the most critical phenological stage for reproductive success, as pollination, fertilization, and initial fruit set occur within this brief window. Grapevine flowers are predominantly self-pollinating (cleistogamous to partially allogamous) with pollen release triggered by calyptra detachment: temperatures below 15°C during stage 6 impair pollen tube growth and ovule fertilization, leading to coulure (flower abortion) and millerandage (uneven berry development) that can reduce potential yield by 30-80% in severe cases. The date of full flowering (BBCH 65) is one of the most consistently recorded phenological milestones and serves as the reference point for predicting harvest date — traditionally estimated at 100 days after BBCH 65 for mid-season cultivars in temperate regions.
BBCH stage 6
BBCH-Stadium 6
Vollblüte-Phase
anthesis stage code
Lorenz DH et al. (1995) Growth stages of the grapevine: phenological growth stages of the grapevine — Codes and descriptions according to the extended BBCH scale, Australian Journal of Grape and Wine Research 1: 100-103 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 145-175 May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press pp: 1-120
BBCH 6 Flowering
A phenological observation of BBCH 75 for Merlot in Saint-Émilion on July 20 documents that berries have reached pea size (approximately 7-8 mm diameter) — the progression through stage 7 spans BBCH 71 (fruit set, young fruits begin to swell, remains of flowerhoods lost), BBCH 73 (berries groat-sized, clusters begin to hang), BBCH 75 (berries pea-sized, 50% of final berry size reached), BBCH 77 (berries beginning to touch within the cluster, bunch closure), and BBCH 79 (majority of berries touching, end of stage 7), with the entire fruit development phase lasting 6-10 weeks and berry volume following a characteristic double-sigmoid growth curve with rapid cell division in the first 3-4 weeks, a 2-3 week lag phase of minimal growth, and resumed expansion through cell enlargement prior to véraison.
A principal growth stage code designating BBCH stage 7 (codes 71-79) that encompasses the post-flowering period of berry growth from initial fruit set through the attainment of full berry size prior to the onset of ripening (véraison), documenting the progressive enlargement of berries through the two distinct phases of pericarp cell division and cell expansion separated by the lag phase.
BBCH_7_Fruit_Development covers the critical period of berry growth that establishes the physical framework for subsequent ripening. The double-sigmoid growth pattern characteristic of grape berries reflects two distinct developmental programs: Phase I (BBCH 71-75) is dominated by pericarp cell division establishing berry cell number, while Phase III (BBCH 77-79) involves cell expansion driven by water and solute accumulation — separated by a lag phase (Phase II) during which organic acid accumulation reaches its maximum and seeds approach physiological maturity. Berry size at the end of stage 7 is a primary determinant of wine quality parameters because it establishes the skin-to-pulp ratio that influences phenolic extraction, color intensity, and flavor concentration in the resulting wine.
BBCH stage 7
BBCH-Stadium 7
Beerenwachstums-Phase
Fruchtentwicklungsstadium
berry growth stage code
fruit development stage
Lorenz DH et al. (1995) Growth stages of the grapevine: phenological growth stages of the grapevine — Codes and descriptions according to the extended BBCH scale, Australian Journal of Grape and Wine Research 1: 100-103 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 175-225 Coombe BG and McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening, Australian Journal of Grape and Wine Research 6: 131-135
BBCH 7 Fruit Development
A phenological record of BBCH 85 for Cabernet Sauvignon in Pauillac on September 5 documents that berries have softened and developed variety-specific color with approximately 50% of final sugar concentration reached — the progression through stage 8 spans BBCH 81 (beginning of ripening, berries begin to brighten in color — véraison onset), BBCH 83 (berries developing color), BBCH 85 (softening of berries, sugar accumulation advancing), BBCH 87 (berries ripe for harvest, full varietal flavor development), and BBCH 89 (berries fully ripe, overripe characteristics beginning), with the ripening period lasting 35-65 days from véraison to harvest depending on cultivar, climate, and target wine style — Brix increasing from 5-8° at BBCH 81 to 22-28° at BBCH 89.
A principal growth stage code designating BBCH stage 8 (codes 81-89) that encompasses the ripening period from véraison (onset of color change and softening in berries) through the attainment of harvest maturity, documenting the progressive accumulation of sugars, degradation of organic acids, development of flavor and aroma compounds, and phenolic maturation that collectively define fruit quality at harvest.
BBCH_8_Ripening is the phenological stage of greatest oenological significance, as the biochemical transformations during ripening determine the chemical composition and sensory potential of the harvested fruit. Véraison (BBCH 81) marks the developmental switch from organic acid accumulation to sugar accumulation, triggered by abscisic acid (ABA) signaling and accompanied by cell wall softening, anthocyanin synthesis in red cultivars, and the onset of flavor and aroma precursor development: the rate and completeness of ripening are strongly influenced by temperature (optimally 20-30°C), water availability, crop load, and canopy microclimate. Harvest timing decisions require balancing sugar accumulation (determining potential alcohol), acid degradation (determining wine acidity), and phenolic maturity (determining tannin quality and color stability) — parameters that do not always advance synchronously, particularly under heat stress conditions.
BBCH stage 8
BBCH-Stadium 8
Beerenreife-Phase
maturation stage code
ripening stage
Lorenz DH et al. (1995) Growth stages of the grapevine: phenological growth stages of the grapevine — Codes and descriptions according to the extended BBCH scale, Australian Journal of Grape and Wine Research 1: 100-103 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 225-295 Coombe BG and McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening, Australian Journal of Grape and Wine Research 6: 131-135
BBCH 8 Ripening
A phenological record of BBCH 93 for Riesling in the Mosel on November 5 documents the beginning of leaf fall with leaves turning yellow and starting to abscise — the progression through stage 9 spans BBCH 91 (after harvest, end of wood maturation with shoots fully lignified and brown), BBCH 92 (beginning of leaf discoloration), BBCH 93 (beginning of leaf fall), BBCH 95 (50% of leaves fallen), BBCH 97 (end of leaf fall, complete defoliation), with the stage 9 duration typically lasting 4-8 weeks from harvest through complete dormancy establishment, and the timing and completeness of shoot lignification during BBCH 91 critically determining winter cold hardiness — poorly lignified shoots suffering frost damage at -12 to -15°C versus -20 to -25°C for fully matured wood.
A principal growth stage code designating BBCH stage 9 (codes 91-97) that encompasses the post-harvest period of shoot maturation (lignification), leaf senescence, leaf abscission, and entry into winter dormancy, documenting the grapevine's transition from active growth to the dormant overwintering state that completes the annual phenological cycle.
BBCH_9_Senescence completes the annual phenological cycle and encompasses processes essential for vine survival and the following season's productivity. Shoot maturation (lignification of periderm, accumulation of starch reserves in wood and roots, dehydration of bud tissues) during BBCH 91-92 determines both cold hardiness and the carbohydrate reserve pool available for the next spring's budburst and early shoot growth: inadequate shoot maturation due to late harvest, excessive crop load, or early autumn frost can compromise both winter survival and subsequent-season vine vigor. Leaf senescence involves programmed nutrient remobilization from leaves to perennial structures — nitrogen, phosphorus, and potassium are recovered at efficiencies of 50-70% before leaf abscission, contributing significantly to the vine's nutrient economy. The date of full leaf fall (BBCH 97) marks the beginning of the winter dormancy period (BBCH 00 of the following cycle).
BBCH stage 9
BBCH-Stadium 9
Blattfall- und Reifephase
Seneszenzstadium
dormancy transition code
senescence stage
Lorenz DH et al. (1995) Growth stages of the grapevine: phenological growth stages of the grapevine — Codes and descriptions according to the extended BBCH scale, Australian Journal of Grape and Wine Research 1: 100-103 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 295-330 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp: 80-108
BBCH 9 Senescence
The BBCH code 81 assigned to Cabernet Sauvignon in the Napa Valley on August 25 designates the beginning of ripening (véraison onset) where berries begin to develop variety-specific color — the first digit 8 identifies principal growth stage 8 (ripening of berries), and the second digit 1 indicates the secondary stage (beginning of ripening, berries begin to brighten in color), with this two-digit code enabling precise phenological comparison with Cabernet Sauvignon at other sites worldwide and serving as the decision trigger for irrigation management adjustments, canopy manipulation timing, and crop estimation scheduling.
A phenological code assigned according to the BBCH (Biologische Bundesanstalt, Bundessortenamt und Chemische Industrie) extended scale for Vitis, consisting of a two-digit numerical code where the first digit designates the principal growth stage (0-9) and the second digit specifies the secondary growth stage within that principal stage, providing a standardized developmental classification from dormancy through fruit maturity and senescence.
BBCH_Code is the internationally adopted standard for grapevine phenological recording, developed by Lorenz et al. (1995) as an adaptation of the general BBCH crop staging system (Meier 2001) to Vitis-specific developmental morphology. The two-digit system encodes 10 principal growth stages (0-9) each subdivided into up to 10 secondary stages (0-9), yielding a maximum of 100 discrete developmental stages from BBCH 00 (dormancy, winter buds pointed) through BBCH 97 (end of leaf fall): not all 100 codes are defined for grapevine, with approximately 47 stages formally described for Vitis. The BBCH code is referenced by EU plant protection product registration (defining application windows), precision viticulture scheduling systems, and phenological monitoring networks worldwide.
BBCH growth stage code
BBCH scale value
BBCH-Code
BBCH-Entwicklungsstadium
BBCH-Skala-Wert
Lorenz scale code
Lorenz DH et al. (1995) Growth stages of the grapevine: phenological growth stages of the grapevine — Codes and descriptions according to the extended BBCH scale, Australian Journal of Grape and Wine Research 1: 100-103 Meier U (2001) Growth Stages of Mono- and Dicotyledonous Plants, BBCH Monograph 2nd Ed, Federal Biological Research Centre Coombe BG (1995) Growth stages of the grapevine, Australian Journal of Grape and Wine Research 1: 104-110
BBCH Code
BBCH stage precision assessment comparing observer consistency reveals the stage-specific variation in determination reliability, with BBCH 65 (full bloom, 50% of flowerhoods fallen) showing high precision (inter-observer agreement 92%, clear quantitative criterion), BBCH 81 (beginning of ripening, berries begin to develop variety-specific color) showing moderate precision (agreement 78%, color assessment somewhat subjective), and BBCH 85 (softening of berries) showing lower precision (agreement 65%, tactile assessment variable), demonstrating how stage definition clarity affects BBCH precision with implications for applications requiring high staging accuracy.
BBCH stage precision quality is a developmental stage quality that represents the accuracy and resolution with which grapevine developmental stages can be determined using the BBCH (Biologische Bundesanstalt, Bundessortenamt und CHemische Industrie) standardized coding system, characterizing the reliability of stage assignment that affects the utility of phenological data for management decisions, modeling applications, and research communication. Within the BFO framework, this quality specifically addresses the measurement precision dimension of stage determination, distinguishing the accuracy of BBCH code assignment from the biological reality of developmental progress while recognizing that the discrete categories of the BBCH system impose boundaries on continuous developmental processes, with precision depending on how clearly observable vine characteristics map to stage definitions. The ontological significance lies in BBCH stage precision determining the reliability of phenological data for applications requiring accurate timing, where imprecise stage determination introduces uncertainty into management decisions (pesticide timing, harvest scheduling), phenological models (calibration and validation affected by observation error), and research comparisons (treatment effects confounded with staging variation), making precision a key quality attribute of phenological assessment. The factors affecting BBCH stage precision include the clarity of stage-defining criteria (some stages having unambiguous markers, others requiring judgment), observer training and experience (consistent interpretation of stage definitions), within-vine variation (different shoots potentially at different stages), and the resolution limitations of the BBCH system itself (10 substages within each principal stage may not capture all meaningful developmental variation). The principal growth stages of the grapevine BBCH scale include Stage 0 (sprouting/budbreak, BBCH 00-09), Stage 1 (leaf development, BBCH 11-19), Stage 5 (inflorescence emergence, BBCH 53-57), Stage 6 (flowering, BBCH 60-69), Stage 7 (fruit development, BBCH 71-79), Stage 8 (ripening, BBCH 81-89), and Stage 9 (senescence, BBCH 91-97), with each principal stage containing substages providing finer developmental resolution. The standardization benefits of BBCH precision extend to international communication, database compatibility, and model transferability, as the globally recognized coding system enables comparison of phenological data across regions, cultivars, and research programs when consistent precision is maintained. Understanding BBCH stage precision quality enables assessment of phenological data reliability, appropriate interpretation of staging uncertainty in management and modeling applications, and training protocols for consistent stage determination across observers.
BBCH_Stage_Precision_Quality addresses the accuracy and reliability of developmental stage determination using the standardized BBCH coding system. This quality affects the utility of phenological data for management and research applications. The class supports phenological data quality assessment through precision characterization across BBCH stages.
BBCH precision
BBCH-Stadienpräzision
Stadienbestimmungsgenauigkeit
Stadiengenauigkeit
phenological coding precision
phänologische Kodierungspräzision
stage determination accuracy
staging accuracy
Lorenz DH et al. (1995) Growth stages of the grapevine: phenological growth stages of the grapevine (Vitis vinifera L. ssp. vinifera)—codes and descriptions according to the extended BBCH scale. Australian Journal of Grape and Wine Research 1(2): 100-103 Meier U (2001) Growth Stages of Mono- and Dicotyledonous Plants. BBCH Monograph, 2nd Ed. Federal Biological Research Centre for Agriculture and Forestry, Berlin
BBCH Stage Precision Quality
A BC1 population from (Riesling × VB91-26-4) × Riesling bears BC1_Roles, with each individual averaging 75% Riesling genome while segregating 1:1 for Run1 powdery mildew resistance—resistant individuals selected through molecular markers for advancement to BC2. BC1 progeny from a Chardonnay × resistant donor F1 crossed back to Chardonnay serve in BC1_Roles, evaluated for both resistance retention and Chardonnay character recovery, with superior individuals entering the next backcross cycle. A BC1 Cabernet Sauvignon derivative carrying Rpv3 downy mildew resistance bears a BC1_Role as an early-stage introgression product, with additional backcross generations planned to achieve the >95% Cabernet genome recovery targeted for variety release.
A backcross hybrid role that inheres in a grapevine genotype representing the first backcross generation (BC1), produced by crossing an F1 hybrid back to the recurrent parent, resulting in progeny that average 75% recurrent parent genome and 25% donor parent genome. BC1 individuals segregate for traits inherited from the donor parent—for a dominant resistance gene heterozygous in the F1, 50% of BC1 progeny will carry the resistance allele while 50% will lack it, enabling selection of resistant individuals for further backcrossing. The BC1 generation represents the first step in recurrent parent genome recovery following initial hybridization, with the 75% average genome contribution substantially higher than the F1's 50% but still carrying considerable donor parent influence. BC1 populations are typically large (hundreds to thousands of individuals) to ensure adequate numbers of resistance-positive individuals for phenotypic or marker-based selection. BC1 individuals selected for resistance and acceptable phenotype may be advanced to BC2 (continued backcrossing for further genome recovery) or evaluated directly as variety candidates if the trait introgression and quality are satisfactory.
BC1_Role specifies the first backcross position where recurrent parent recovery begins in earnest following F1 hybridization. The 75% theoretical genome recovery represents an average individual BC1 progeny vary around this mean due to the randomness of meiotic recombination, with some individuals carrying more or less recurrent parent genome than average. Marker-assisted background selection can identify individuals with above-average recurrent parent recovery, accelerating the backcross program by selecting for genome recovery alongside the target trait. BC1 populations often show substantial phenotypic variation reflecting the 25% donor genome still present, with wine quality typically not yet approaching recurrent parent standards—justifying continued backcrossing for most breeding programs. The decision whether to advance to BC2 or evaluate BC1 as variety candidates depends on program objectives, donor parent characteristics, and market requirements.
BC1
BC1 generation
BC1-Generation
BC1-Hybride
erste Rückkreuzung
first backcross
first backcross hybrid
Töpfer R et al. (2011) Molecular breeding in grapes, in Adam-Blondon AF et al. (eds) Genetics, Genomics, and Breeding of Grapes, CRC Press pp.: 160-185 Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330
BC1 Role
A BC2 Riesling derivative with Rpv10 downy mildew resistance bears a BC2_Role, its 87.5% average Riesling genome recovery enabling preliminary wine quality assessment showing improved Riesling character compared to BC1, while resistance confirmed through marker analysis. BC2 progeny from a Pinot Noir backcross program serve in BC2_Roles as intermediate breeding materials, with the best individuals selected for advancement to BC3 based on combined resistance, phenotype, and initial fruit evaluation. A BC2 Chardonnay × Run1 carrier genotype bears a BC2_Role in French breeding, representing the generation where quality assessment becomes meaningful and variety candidate identification may begin for programs accepting moderate recurrent parent recovery.
A backcross hybrid role that inheres in a grapevine genotype representing the second backcross generation (BC2), produced by crossing a selected BC1 individual back to the recurrent parent, resulting in progeny that average 87.5% recurrent parent genome and 12.5% donor parent genome. BC2 individuals show substantially increased recurrent parent character compared to BC1, with the reduced donor genome contribution (halved from BC1) diminishing undesirable interspecific traits while target traits under selection are maintained. For breeding programs targeting near-isogenic lines (varieties differing from the recurrent parent only at the introgressed locus), BC2 represents intermediate progress—substantial genome recovery achieved but typically insufficient for varieties intended to closely match recurrent parent performance. BC2 populations segregate for heterozygous donor traits similarly to BC1, with 50% of progeny carrying a dominant resistance allele if the BC1 parent was heterozygous. Selection at BC2 emphasizes both resistance retention and quality trait evaluation, as the recovered recurrent parent background enables meaningful assessment of wine character.
BC2_Role marks the generation where recurrent parent character becomes substantially expressed, enabling quality evaluation that was premature in earlier generations with higher donor genome content. The 87.5% theoretical genome recovery (varying among individuals) represents sufficient recurrent parent contribution that some breeding programs evaluate BC2 individuals as potential variety candidates, particularly if the donor parent was itself an improved genotype (e.g., a PIWI variety rather than a wild species). Programs targeting maximum recurrent parent recovery (>95%) continue to BC3 and beyond, while those accepting moderate interspecific contributions may release BC2-generation varieties if quality is acceptable. The BC2 generation often shows the first populations where wine production from progeny can meaningfully assess varietal character recovery, informing selection decisions for subsequent generations.
BC2
BC2 generation
BC2-Generation
BC2-Hybride
second backcross
second backcross hybrid
zweite Rückkreuzung
Töpfer R et al. (2011) Molecular breeding in grapes, CRC Press pp.: 160-185 Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance, Molecular Genetics and Genomics 291: 1573-1594
BC2 Role
A BC3 Riesling derivative carrying pyramided Rpv3 and Rpv10 resistances bears a BC3_Role with 93.75% average Riesling genome, producing wines approaching Riesling character in quality trials—a potential variety candidate designated provisionally as "Riesling R" (resistant Riesling) pending registration decisions. BC3 progeny from advanced JKI Geilweilerhof breeding serve in BC3_Roles as near-commercial breeding lines, with the best individuals entering final variety evaluation protocols for potential registration. A BC3 Müller-Thurgau × resistance donor derivative bears a BC3_Role in German breeding, the high recurrent parent recovery enabling variety release consideration with wine quality documentation approaching the original Müller-Thurgau standard.
A backcross hybrid role that inheres in a grapevine genotype representing the third backcross generation (BC3), produced by crossing a selected BC2 individual back to the recurrent parent, resulting in progeny that average 93.75% recurrent parent genome and 6.25% donor parent genome. BC3 individuals closely approach the recurrent parent in overall genetic composition, with only approximately 6% of their genome derived from the original donor—sufficient for introgressing a resistance locus while minimizing linkage drag (retention of undesirable donor alleles closely linked to the target locus). BC3 represents a common target generation for resistance introgression programs, as the >93% recurrent parent recovery typically produces varieties with wine character close to the recurrent parent while carrying effective resistance. BC3 populations may be large enough to enable selection for reduced linkage drag, identifying individuals where recombination has minimized the donor-derived chromosome segment around the resistance locus. Variety candidates from BC3 generations increasingly resemble the recurrent parent, enabling marketing strategies that emphasize the established variety name with added resistance trait.
BC3_Role represents the generation commonly targeted for variety release in programs balancing recurrent parent recovery with practical breeding timeline constraints. The 93.75% average genome recovery (with individual variation around this mean) typically provides wine character sufficiently similar to the recurrent parent for commercial acceptance, while the remaining ~6% donor contribution is largely undetectable in quality evaluation if the introgressed segment is small. Marker-assisted background selection at BC3 can identify individuals with above-average recurrent parent recovery, potentially achieving >95% recovery and enabling "accelerated BC3" individuals to match BC4 standards. The BC3 generation represents 10-15 years from initial cross in typical breeding programs, with variety evaluation adding additional years before commercial release—a substantial investment justifying careful selection at each generation.
BC3
BC3 generation
BC3-Generation
BC3-Hybride
dritte Rückkreuzung
third backcross
third backcross hybrid
Töpfer R et al. (2011) Molecular breeding in grapes, CRC Press pp.: 160-185 Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330 Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance, Molecular Genetics and Genomics 291: 1573-1594 Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22
BC3 Role
A BC4 Pinot Noir derivative carrying stacked Run1 and Ren3 powdery mildew resistances bears a BC4_Role with >96% Pinot Noir genome, producing wines in blind trials indistinguishable from conventional Pinot Noir—enabling potential registration as "Pinot Noir R" for premium Burgundy-style production with minimal fungicide requirement. BC4 Riesling lines from German JKI programs serve in BC4_Roles as advanced breeding products approaching commercial release, with decades of backcross breeding yielding material meeting the stringent quality standards expected for Germany's signature white variety. A BC4 Cabernet Sauvignon derivative bears a BC4_Role in French breeding targeting Bordeaux appellations, where market and regulatory requirements for varietal character recognition necessitate maximum recurrent parent recovery before variety approval.
A backcross hybrid role that inheres in a grapevine genotype representing the fourth backcross generation (BC4), produced by crossing a selected BC3 individual back to the recurrent parent, resulting in progeny that average 96.875% recurrent parent genome and approximately 3% donor parent genome. BC4 individuals are near-isogenic to the recurrent parent, differing primarily at the introgressed locus and its immediately flanking regions, with the minimal donor genome contribution representing the practical limit of phenotypic detectability for most traits. BC4 generation products represent the quality benchmark for backcross breeding—varieties with wine character essentially indistinguishable from the recurrent parent while carrying effective disease resistance. The extended timeline required to reach BC4 (typically 15-20+ years from initial cross) has motivated interest in marker-assisted background selection to achieve equivalent genome recovery in fewer generations, potentially matching BC4 standards at BC2 or BC3 through intensive molecular selection. BC4 and beyond may be pursued for iconic varieties where maximum recurrent parent fidelity is essential (Grand Cru Burgundy clones, classified Bordeaux varieties), while BC3 release may be acceptable for varieties with less stringent market expectations.
BC4_Role represents the practical endpoint for most backcross introgression programs, where additional generations yield diminishing returns in genome recovery (BC5 = 98.4%, BC6 = 99.2%) while extending timelines and resource requirements. The ~97% average recurrent parent genome at BC4 typically exceeds natural variation among clones of the recurrent parent variety, meaning BC4 introgression products may be genetically more similar to the recurrent parent than some certified clones are to each other. The BC4 designation increasingly appears in variety registration applications for resistant derivatives of established cultivars, with naming conventions evolving to indicate the relationship to the recurrent parent (suffix systems, "resistant" designations, or entirely new names depending on jurisdiction and marketing strategy). Achieving true near-isogenic status (differing only at the introgressed locus) requires selection against linkage drag—donor genome segments flanking the target locus—which marker-assisted selection can accelerate but cannot completely eliminate in typical breeding populations.
BC4
BC4 generation
BC4-Generation
BC4-Hybride
fourth backcross
fourth backcross hybrid
near-isogenic line
quasi-isogene Linie
vierte Rückkreuzung
Töpfer R et al. (2011) Molecular breeding in grapes, CRC Press pp.: 160-185 Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330 Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Schneider A et al. (2019) Marker-assisted introgression, Euphytica 215: 45
BC4 Role
The bearing habit description of Nebbiolo documents distal fruitfulness — the basal buds at positions 1-3 on one-year-old canes are predominantly vegetative (fruitfulness 70% fruitfulness at all node positions from 1-10, accommodating either spur or cane pruning systems).
A growth habit description component that records the spatial distribution of fruitful buds and resulting clusters along the shoot axis and within the vine framework, documenting the cultivar-specific pattern of basal versus distal fruitfulness, the typical cluster positions relative to the cordon or cane, the proportion of primary versus secondary bud shoots bearing fruit, and the influence of bud position on cluster size and number that together determine the pruning strategy required to optimize yield distribution across the vine for balanced ripening and efficient harvest operations.
Bearing_Habit_Description documents the cultivar-specific distribution of reproductive capacity along the shoot that is the primary determinant of pruning system selection — arguably the most consequential vineyard design decision because it permanently defines the vine framework, training system, and management approach for the 25-50 year productive life of the planting. The distinction between basally fruitful cultivars (suited to spur pruning) and distally fruitful cultivars (requiring cane pruning) reflects differences in the developmental history of compound buds at each node position — during the bud organogenesis period in the season preceding fruiting, inflorescence primordia are initiated within the developing compound bud under the influence of temperature, light exposure, carbohydrate supply, and hormonal signals that vary with bud position along the shoot: in basally fruitful cultivars, the conditions experienced by basal buds (moderate light through the open canopy near the cordon, moderate temperature buffered by proximity to the woody framework) are sufficient to trigger inflorescence initiation, while in distally fruitful cultivars, the basal buds experience insufficient light or excessive correlative inhibition from the actively growing shoot tip to initiate inflorescences, and only the mid-cane and distal buds (experiencing higher light levels in the canopy exterior) achieve fruitful status. The bearing habit also influences the response to mechanical damage or late frost — basally fruitful cultivars can recover partial crops from secondary and tertiary buds at the spur positions after frost damage, while distally fruitful cultivars lose their entire fruitful bud population if the retained cane buds are killed, making frost damage more economically devastating for cane-pruned cultivars.
Ertragsgewohnheits-Charakterisierung
Fruchtverteilungs-Muster
Tragegewohnheit-Beschreibung
cropping habit characterization
fruiting pattern record
reproductive growth distribution
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Dry PR (2000) Canopy management for fruitfulness, Australian Journal of Grape and Wine Research 6: 109-115 Smart RE and Robinson M (1991) Sunlight into Wine, Winetitles May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press
Bearing Habit Description
A bench graft consisting of Pinot Noir clone 777 scion omega-grafted to 3309 Couderc rootstock bears a Bench_Graft_Role, callused for 21 days at 27°C and 95% RH, then planted in a Burgundy nursery row in April for one season's growth before sale as a rooted grafted vine (plant greffé-soudé) the following winter. A bench graft of Tempranillo clone RJ-51 on 110 Richter serves in a Bench_Graft_Role in Spanish production, wax-dipped for moisture retention after callusing, then planted directly to vineyard in Rioja during May. A bench graft of Chardonnay clone 96 on SO4 bears a Bench_Graft_Role in California production, grown in cellular trays under greenhouse conditions to produce accelerated green-growing plants for high-density vineyard establishment.
A propagation material role that inheres in a grafted combination of scion and rootstock prepared under controlled nursery conditions (on a "bench" or workstation), representing the standard commercial propagation unit for grafted grapevines in modern viticulture. Bench grafts (also termed benchgrafts or indoor grafts) consist of a scion piece (typically 5-8 cm with one to two buds) joined to a rootstock cutting (typically 28-32 cm) through omega, whip-and-tongue, or machine grafting techniques, then subjected to callusing treatment (typically 2-3 weeks at 26-28°C and high humidity) to initiate union formation before nursery planting or direct vineyard establishment. The bench graft format enables quality control of the grafting operation (diameter matching, cut precision, pathogen management), efficient callusing in climate-controlled facilities, and production of large quantities of uniform grafted plants. Bench grafts may be planted as dormant grafts (immediately after callusing, into nursery rows or directly to vineyard) or grown for one season in nursery to produce rooted grafted vines before sale.
Bench_Graft_Role represents the dominant grafted vine production format, with annual global production estimated at 500-700 million units. The controlled environment of bench grafting enables year-round production (independent of field budding seasons), quality control impossible in field conditions, and efficient use of specialized labor and mechanization. Bench graft success rates (typically 80-95% for well-matched material with proper callusing) exceed field budding success (60-80%), justifying the infrastructure investment for commercial nurseries. The callusing phase is critical—insufficient callusing produces weak unions prone to breakage, while excessive callusing may deplete carbohydrate reserves before root establishment. Bench graft quality certification encompasses both rootstock and scion components, with the assembled graft inheriting the lower certification category of its two components.
Bankveredelung
Pfropfrebe
Rebveredelung
Tischveredelung
bench graft
benchgraft
indoor graft
nursery graft
omega graft unit
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 450-468 Gramaje D, Armengol J (2011) Fungal trunk pathogens, Phytopathologia Mediterranea 50: S217-S226 Waite H, Morton L (2007) Hot water treatment and benchgraft survival, Australian Journal of Grape and Wine Research 13: 99-109 Dry N (2007) Grapevine rootstocks, Lythrum Press
Bench Graft Role
Bending tolerance quality comparison during Guyot training establishment in Champagne evaluated breakage incidence and cane viability across cultivars bent to standard horizontal positioning on the fruiting wire. Pinot Meunier demonstrated excellent bending tolerance quality with slender, flexible canes of 8-10 mm diameter bending smoothly to horizontal orientation with less than 1% breakage incidence, maintained vascular function evidenced by uniform budbreak along bent canes, and no detectable yield reduction in bent versus upright cane sections. Chardonnay showed good bending tolerance quality with 2% breakage when standard bending technique was applied, increasing to 5% with hasty or rough handling, indicating adequate but not exceptional tolerance requiring careful technique. Pinot Noir exhibited variable bending tolerance quality dependent on cane characteristics, with thin canes performing well but thicker canes exceeding 12 mm diameter showing 8% breakage rate and 15% incidence of partial vascular damage manifesting as reduced vigor in distal bud positions, motivating selection of moderately-sized canes and earlier bending timing for this cultivar.
Bending tolerance quality is a training response quality that inheres in a grapevine cultivar or clone and represents the capacity of canes and shoots to withstand the mechanical stresses imposed during bending operations required for cane positioning in Guyot training, arching in traditional regional systems, and related manipulations that redirect growth along curved trajectories differing from the natural straight or gently curved growth path, without suffering tissue damage, vascular disruption, or structural failure that would compromise the viability and productivity of the bent cane section. This quality determines the feasibility of training approaches requiring substantial cane curvature and the risk of training-related losses from breakage or physiological damage during the cane positioning operations essential for many traditional and modern training systems. The ontological significance of bending tolerance quality lies in its role as a prerequisite for successful implementation of cane-trained systems including Guyot, Pendelbogen, and various arched configurations where horizontal or downward cane orientation requires bending wood grown in upward orientation, with cultivars lacking adequate bending tolerance suffering unacceptable breakage losses or physiological compromise of bent cane sections. The biological basis of bending tolerance encompasses the mechanical properties of lignified cane tissues including flexibility, elastic limit, and breaking strength the vascular architecture determining whether conducting elements can accommodate the differential compression and tension stresses on inner and outer surfaces of bent canes and the wound response capacity for repairing micro-damage that may occur even in successful bending operations. Cane diameter, internode length, wood density, and the degree of lignification at bending time all influence bending tolerance outcomes, with thinner canes and longer internodes generally exhibiting greater flexibility while thick, heavily lignified canes resist bending and prove more susceptible to breakage. The timing of bending operations critically influences success, with most cultivars exhibiting better bending tolerance during the brief window after full dormancy but before bud swell when wood retains some flexibility while being sufficiently lignified to maintain imposed shape. Assessment of bending tolerance quality involves quantification of breakage rates during standardized bending operations, evaluation of physiological function in bent cane sections through bud viability and shoot vigor, and long-term monitoring of bent cane productivity and health.
Bending_Tolerance_Quality is_a Training_Response_Quality, representing the capacity to withstand cane bending without damage. This class addresses the mechanical stress tolerance of lignified canes during training operations requiring curvature. Distinguished from Positioning_Ease_Quality which addresses shoot manipulation during the growing season rather than dormant cane bending. The timing window, cane diameter selection, and technique quality all influence outcomes beyond inherent cultivar characteristics. Most relevant for Guyot, Pendelbogen, and arched training systems requiring horizontal or downward cane positioning.
Biegebelastbarkeit
Biegungstoleranz
Bogentoleranz
Rutenflexibilität
arching tolerance
bending strength
cane flexibility
cane pliability
Galet P (2000) General Viticulture. Oenoplurimedia, Chaintre, pp. 189-245 Huglin P, Schneider C (1998) Biologie et écologie de la vigne. Lavoisier, Paris, pp. 253-278 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 85-112
Bending Tolerance Quality
Pinot Noir vines in the Willamette Valley of Oregon bearing a beneficial insect habitat role maintain resident populations of Anagrus daanei parasitoid wasps at 60-80% parasitism rates on Erythroneura leafhopper eggs, Typhlodromus pyri predatory mites at 2-4 per leaf controlling Eotetranychus carpini below economic injury thresholds, and overwintering lacewing (Chrysoperla carnea) populations in bark crevices at 5-12 cocoons per vine, with adjacent hedgerows of Rosa eglanteria providing alternative prey and nectar resources sustaining parasitoid fitness during periods of low vineyard pest density.
A biodiversity support role that inheres in a grapevine whose canopy structure, bark surfaces, and surrounding vegetation provide habitat for predatory and parasitoid arthropods that contribute to natural pest suppression within the vineyard, supporting biological control as an ecosystem service.
Beneficial_Insect_Habitat_Role addresses the vine's function as structural habitat for the natural enemy complex that underpins conservation biological control in viticulture. The role is mediated by canopy complexity, bark roughness (increasing with vine age), pruning residue management, and the availability of alternative food sources such as pollen and extrafloral nectar: intensively managed young vines with smooth bark and minimal structural complexity provide substantially less natural enemy habitat than old, structurally complex vines. This role intersects with Integrated_Production_Role and Organic_Production_Role where biological control is a primary pest management strategy.
Biocontrol-Lebensraum-Rolle
Nützlingshabitat-Rolle
Raubinsekten-Schutz-Rolle
biocontrol habitat role
natural enemy habitat role
predator refuge role
Duso C et al. (2012) Biological control of mites in European vineyards, Integrated Pest Management of Temperate Horticultural Crops pp: 191-217 Pennington T et al. (2018) Habitat requirements of parasitoids in vineyards, BioControl 63: 805-818 Wilson H et al. (2017) Landscape diversity and crop vigor outweigh influence of local diversification on biological control of a vineyard pest, Ecosphere 8: e01736
Beneficial Insect Habitat Role
SO4 (Selection Oppenheim 4, V. berlandieri × V. riparia) is the most widely planted rootstock globally, providing moderate vigor, lime tolerance (17-20% active lime), and balanced nutrient delivery to Riesling across German wine regions and to numerous cultivars worldwide. Kober 5BB (V. berlandieri × V. riparia, selected at Klosterneuburg) tolerates 20% active lime and shows excellent adaptation to the loess and limestone-influenced soils of Austrian Niederösterreich, supporting Grüner Veltliner and Zweigelt. Teleki 5C (V. berlandieri × V. riparia, Hungarian selection) provides similar characteristics with slightly lower vigor, widely used in Central European viticulture from Hungary through Slovenia to northern Italy.
A parentage-based rootstock role that inheres in a grafted rootstock by virtue of its genetic derivation from hybridization between Vitis berlandieri and Vitis riparia, combining traits from these two complementary North American species. V. berlandieri × V. riparia hybrids unite V. berlandieri's exceptional lime tolerance, moderate-to-good drought tolerance, and late budburst with V. riparia's excellent phylloxera resistance, cold hardiness, improved rooting capacity, and growth-controlling characteristics. This hybrid combination produces the most versatile and widely planted rootstock group globally, adapted to calcareous soils (lime tolerance 17-25% active lime), temperate-to-cool climates, and a broad range of viticultural situations. The V. berlandieri × V. riparia group shows moderate vigor (lower than V. berlandieri × V. rupestris), good-to-moderate drought tolerance, intermediate potassium uptake (lower than V. berlandieri × V. rupestris), and adequate cold hardiness for continental European climates. Rooting capacity is substantially improved over pure V. berlandieri (65-85% success rates).
Berlandieri_X_Riparia_Rootstock_Role represents the dominant rootstock hybrid group for temperate viticulture on calcareous soils. The combination balances V. berlandieri's lime tolerance (the trait most lacking in other species) with V. riparia's improved rooting, cold hardiness, and phylloxera resistance. Key selections include SO4, 5BB Kober, 5C Teleki, 125AA, 420A Millardet, and 161-49 Couderc. Compared to V. berlandieri × V. rupestris hybrids, this group shows lower vigor, reduced drought tolerance, better cold hardiness, and lower potassium delivery—characteristics favoring cool-climate premium wine production. The group is sometimes called the "Kober group" after the influential Austrian breeder Franz Kober.
Berlandieri × Riparia rootstock
Berlandieri-Riparia-Hybridunterlage
Berlandieri-×-Riparia-Unterlage
Kober-Gruppe
V. berlandieri × V. riparia rootstock
berlandieri-riparia hybrid rootstock
Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 285-320 Bauer KP, Götz G (2013) Unterlagen der Rebe, Bayerische Landesanstalt für Weinbau und Gartenbau
Berlandieri X Riparia Rootstock Role
110 Richter (V. berlandieri Resseguier No. 2 × V. rupestris Martin) is the archetypal Berlandieri × Rupestris rootstock, providing moderate vigor, excellent drought tolerance (root penetration exceeding 4 m), and lime tolerance (17-20% active lime), dominating Mediterranean viticulture for Tempranillo in Spain, Sangiovese in Italy, and Grenache in southern France. 1103 Paulsen (V. berlandieri Resseguier No. 2 × V. rupestris du Lot) confers high vigor and exceptional drought/heat tolerance to Primitivo and Nero d'Avola across southern Italy. 140 Ruggeri (V. berlandieri × V. rupestris du Lot) shows the highest vigor and K uptake in the group, supporting Catarratto on the driest hillside sites of Sicily.
A parentage-based rootstock role that inheres in a grafted rootstock by virtue of its genetic derivation from hybridization between Vitis berlandieri and Vitis rupestris, combining traits from these two complementary North American species native to the limestone and rocky habitats of Texas and the Ozark region. V. berlandieri × V. rupestris hybrids unite V. berlandieri's exceptional lime tolerance and iron efficiency with V. rupestris's deep rooting, drought tolerance, improved propagation success, and moderate vigor. This hybrid combination produces rootstocks supremely adapted to Mediterranean and warm-temperate climates on calcareous, drought-prone soils, with lime tolerance of 17-30% active lime, excellent drought tolerance (sustaining productivity below 400 mm annual rainfall), high-to-moderate vigor, and relatively high potassium translocation. The V. berlandieri × V. rupestris group dominates viticulture in Mediterranean Europe (Spain, Portugal, southern France, Italy, Greece), California, Australia, South Africa, and Chile. Cold hardiness is reduced compared to V. berlandieri × V. riparia hybrids, limiting use in continental climates with severe winters.
Berlandieri_X_Rupestris_Rootstock_Role represents the dominant rootstock group for warm-climate viticulture, combining the drought adaptations of both parental species evolved in the semi-arid limestone habitats of Texas and the Ozarks. Key selections include 110 Richter, 99 Richter, 1103 Paulsen, 140 Ruggeri, and 775 Paulsen. The high vigor and elevated potassium delivery of this group can be problematic for premium wine production where canopy control and must acidity are priorities compensatory management (deficit irrigation, crop load adjustment) is often required. Climate change is expanding the geographic relevance of this group as drought frequency increases in traditionally temperate regions.
Berlandieri × Rupestris rootstock
Berlandieri-Rupestris-Hybridunterlage
Berlandieri-×-Rupestris-Unterlage
Richter-Paulsen-Gruppe
V. berlandieri × V. rupestris rootstock
berlandieri-rupestris hybrid rootstock
Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Carbonneau A (1985) The early selection of grapevine rootstocks for resistance to drought conditions, American Journal of Enology and Viticulture 36: 195-198 Serra I et al. (2014) The interaction between rootstocks and cultivars to enhance drought tolerance, Australian Journal of Grape and Wine Research 20: 1-14
Berlandieri X Rupestris Rootstock Role
Cabernet Sauvignon berries developing from BBCH 71 (fruit set, 2 mm diameter, 7-10 days post-fertilization) through BBCH 89 (ripe berries, 12-15 mm diameter, 14-16 g fresh weight cluster average, 110-125 days post-fertilization) exhibit characteristic development= rapid green growth to 8-10 mm by BBCH 75 (6 weeks), lag phase maintaining 8-10 mm at BBCH 77 (8 weeks), veraison color change from green to red-purple at BBCH 81-83 (9-11 weeks), and ripening expansion to final size with sugar accumulation reaching 23-26°Brix, acidity declining to 5.5-7 g/L, pH increasing to 3.6-3.9, creating mature berries suitable for premium red wine production. Chardonnay white wine cultivar produces berries developing from 2 mm at fruit set (BBCH 71) to 11-14 mm at maturity (BBCH 89, 95-110 days post-fertilization, 1.8-2.4 g per berry), with veraison marked by color change from green to golden-yellow and achieving maturity at 21-24°Brix, 6-8.5 g/L acidity, pH 3.2-3.5, optimal for white wine production. Thompson Seedless stenospermocarpic table cultivar exhibits berry development from fruit set to maturity (18-22 mm diameter, 4-6 g per berry, 120-140 days development) with early fertilization followed by seed abortion at 3-5 weeks post-fertilization leaving only seed traces, allowing continued parthenocarpic berry expansion producing large seedless berries at 18-22°Brix suitable for fresh consumption. Pinot Noir berry anatomical sections at maturity reveal skin structure (epidermis, hypodermis, 3-5 outer mesocarp layers, 80-120 μm total thickness) containing anthocyanins, flesh structure (large parenchyma cells 80-180 μm diameter occupying 90-94% of berry volume) containing 24-26% sugars and 5.5-6.5 g/L acids, and 2-3 mature seeds (6-8 mm length) with lignified seed coats, demonstrating mature berry anatomical organization derived from fertilized ovary transformation.
A berry is the fruit developing from the fertilized or parthenocarpic ovary of the Vitis flower, representing the mature or developing grape that is the primary agricultural product and the defining characteristic structure of Vitaceae, consisting of skin (exocarp), flesh (mesocarp), and seeds (from fertilized ovules) or seed traces (in seedless types), with berry development proceeding through distinct phases from fruit set through ripening spanning 60-150 days depending on cultivar and environmental conditions. The berry structure comprises= (1) skin (exocarp and outer mesocarp layers) derived from ovary wall outer tissues, typically 40-180 μm thick at maturity, containing cuticle, epidermis, hypodermis, and several outer mesocarp cell layers, bearing stomata, lenticels, and often waxy bloom (epicuticular wax), functioning as protective barrier and containing pigments (anthocyanins in red/black cultivars, flavonols in white cultivars), tannins, and aromatic compounds (2) flesh (mesocarp) derived from ovary wall middle tissues, comprising the bulk of berry volume (typically 85-95% of total berry mass), consisting of large thin-walled parenchyma cells accumulating sugars (primarily glucose and fructose reaching 18-28% at maturity), organic acids (primarily tartaric and malic acids declining from 15-25 g/L at veraison to 4-9 g/L at maturity), water (70-85% of flesh mass at maturity), and various secondary metabolites (3) seeds developing from fertilized ovules, typically 0-4 seeds per berry (0 in seedless cultivars, 1-4 in seeded cultivars), or seed traces (rudimentary arrested seed structures in stenospermocarpic seedless types) (4) vascular system including dorsal and ventral vascular bundles running from pedicel through berry providing nutrient and water transport during development. Berry development follows characteristic phases= (1) fruit set (BBCH 71, 0-2 weeks post-fertilization) involving ovary retention, initial expansion, and transition from flower to fruit (2) green growth phase I (BBCH 71-75, weeks 2-6 post-fertilization) with rapid cell division creating final cell number (3) lag phase (BBCH 75-77, weeks 6-8) with minimal size change, seed maturation, and metabolic transitions (4) veraison (BBCH 81-83, weeks 8-10) marking onset of ripening with color change, softening, and sugar accumulation beginning (5) ripening phase II (BBCH 83-89, weeks 10-18+) with berry expansion through cell enlargement, sugar accumulation, acid decline, and secondary metabolite development. Berry size at maturity varies dramatically among cultivars= small-berried wine cultivars 6-12 mm diameter, 0.5-1.5 g fresh weight medium-berried cultivars 12-18 mm diameter, 1.5-3 g large-berried table cultivars 18-28 mm diameter, 3-8 g or larger in extreme types. Berry composition at maturity shows cultivar and environmental variation= sugars 16-28% (expressed as °Brix), titratable acidity 4-12 g/L, pH 3.1-4.2, with composition critically affecting wine or table grape quality.
Berry represents the fruit developing from the ovary and must be distinguished from Ovary (pre-fertilization structure) and from Cluster/Inflorescence (the berry-bearing structure). The berry is the fundamental unit determining grape quality and yield, with berry development biology, composition, and morphology critically important for viticulture and enology. Sub-classification into Developing_Berry captures the progression from fruit set through ripening, with further subdivision into Early and Mid-development phases. Understanding berry development, anatomy, and composition is essential for crop management, harvest timing, quality assessment, and for all aspects of grape production and utilization.
Beere
Traube
Weinbeere
fruit
grape
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 172-245 Coombe, B.G., McCarthy, M.G. (2000). Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6=131-135 Kennedy, J.A. (2002). Understanding grape berry development. Practical Winery & Vineyard July/August=1-5 Ollat, N. et al. (2002). Grape berry development= a review. Journal International des Sciences de la Vigne et du Vin 36=109-131.
Berry
Berry-ambient temperature differential measurement in McLaren Vale Shiraz during a 40°C heat event reveals exposed berry temperatures of 52°C representing a +12°C differential, while shaded berries within the canopy interior show 43°C representing only +3°C differential, with the 9°C difference between exposed and shaded fruit correlating to 40% lower anthocyanin content in exposed berries due to thermal degradation, demonstrating the critical importance of canopy architecture for fruit protection during extreme heat.
Berry-ambient temperature differential quality is a temperature differential quality that represents the difference between grape berry temperature and ambient air temperature, characterizing the degree of thermal departure from environmental conditions resulting from the berry's energy balance where radiant heating typically exceeds convective and minimal evaporative cooling, causing berry temperature to exceed air temperature during daylight hours. Within the BFO framework, this quality specifically captures the thermal relationship between the quality-determining reproductive structure and its atmospheric environment, providing critical information for heat stress assessment and quality prediction that cannot be derived from air temperature monitoring alone. The ontological significance lies in this differential quantifying the additional thermal stress experienced by berries beyond what air temperature indicates, where radiant heating elevations of 5-15°C above ambient are common in exposed fruit, meaning that heat damage thresholds based on air temperature substantially underestimate actual heat exposure. The magnitude of berry-ambient differential depends on solar radiation intensity, berry color (dark berries absorbing more radiation), wind speed affecting convective dissipation, and canopy position determining shade protection, with exposed berries in low-wind conditions showing the largest positive differentials. Unlike canopy-air differential which varies with plant water status through transpirational cooling modulation, berry-ambient differential is relatively insensitive to vine water status because berries lack significant transpirational cooling capacity, instead reflecting primarily physical factors of radiation load and convective environment. Measurement requires simultaneous berry temperature determination (contact thermocouple or infrared thermometer) and air temperature measurement in a shaded, ventilated location representative of the berry environment. Understanding this differential enables appropriate interpretation of air temperature-based heat wave warnings, design of shade or cooling interventions for heat stress mitigation, and correlation of berry composition changes with actual berry thermal exposure.
Berry_Ambient_Temperature_Differential_Quality specifically addresses the fruit thermal status relative to environment, complementing Canopy_Air_Temperature_Differential_Quality which reflects whole-vine water status. This quality supports heat stress prediction and mitigation strategy evaluation. Unlike canopy temperature differential, berry-ambient differential is largely insensitive to vine water status and instead reflects physical exposure conditions, making it valuable for site and canopy architecture assessment rather than irrigation scheduling.
Beeren-Umgebungs-Temperaturdifferenz
Beerentemperaturerhöhung
Frucht-Umgebungs-Thermaldifferenz
Trauben-Luft-Temperaturdifferenz
berry thermal elevation
berry-air temperature difference
fruit-ambient thermal differential
grape temperature excess
Smart RE, Sinclair TR (1976) Solar heating of grape berries and other spherical fruits. Agricultural Meteorology 17(4): 241-259 Cola G et al. (2020) Grapevine phenology and climate change: relationships and trends in the Veneto region of Italy for 1964-2009. American Journal of Enology and Viticulture 71(3): 173-183
Berry Ambient Temperature Differential Quality
Merlot in Bordeaux shows berry cell division phase during early-mid July Year N+1 at BBCH 71-73, with division activation days 3-6 post-fertilization, peak mitotic activity weeks 2-3 with 11-14 hour cell cycle times, progressive accumulation reaching 1.9-2.5 million mesocarp cells by week 4-5, division completion marking transition to expansion, under oceanic conditions of 20-26°C temperature, 1.7-2.1 seeds per berry supporting division through auxin, typical of this mid-season cultivar establishing cellularity determining 10-13 mm final berry size
Berry cell division phase represents the initial intensive mitotic growth period during berry development when pericarp mesocarp and exocarp cells undergo rapid proliferation establishing final cellular architecture, cell number accumulation determining berry size potential, and division completion marking transition to expansion-dominated growth, proceeding over first 3-5 weeks post-fertilization as primary developmental mechanism creating berry flesh cellularity. This critical cell proliferation phase occurs during early summer in late June-July in temperate regions Year N+1, spanning approximately 3-5 weeks from berry commitment with peak activity weeks 1-3, corresponding to BBCH 71-73 young fruit stages. Cell division phase mechanism involves auxin-stimulated cell cycle activation initiating mitotic activity, progressive cell number accumulation through repeated division rounds, peak division intensity during weeks 1-3 with maximum mitotic rates, gradual division rate decline weeks 3-5 as cells transition to expansion, and division completion establishing final cell population determining berry cellular foundation and size potential. Auxin stimulation of division shows hormone from developing seeds triggering cell cycle entry in quiescent pericarp cells, auxin signaling activating cyclin-dependent kinases and cell cycle regulators, hormone dose from seed number affecting division intensity and duration, and auxin as primary growth-promoting signal during early berry development. Cell cycle activation shows G1 phase entry from quiescent G0 state, progression through S phase with DNA replication, G2 phase preparation, and M phase mitosis producing daughter cells, with cycle times 10-15 hours during peak activity representing very rapid division characteristic of developmental growth. Division phase temporal progression shows initial division activation days 2-7 post-fertilization as auxin signaling establishes, mitotic activity onset week 1 with accelerating division rate, peak division activity weeks 2-3 with maximum cell proliferation, declining division weeks 3-4 as cells progressively exit cycle, and division completion week 4-5 with final cells entering expansion phase. Active berry cell division shows early intensive phase with division activation and onset establishing proliferative program, multiple cell generations produced through repeated division rounds, and coordinated division across pericarp tissues creating organized cellular architecture. Peak division activity represents maximum proliferative phase with highest mitotic rates, maximum cell number generation determining final cellularity, and tissue-specific division patterns in pericarp layers and seed compartment regions. Cell number accumulation dynamics show exponential increase during peak division weeks 1-3, typical progression from 50,000-100,000 cells at commitment to 1-4 million cells at division completion, accumulation rate varying with temperature and hormonal signaling, and final cell number as critical size determinant. Final cell number determination shows genetic programming establishing cultivar-characteristic cell number ranges, seed number effects through differential auxin dose creating 20-40% variation, temperature during division phase affecting proliferation rate and final number, and environmental conditions during critical weeks 1-4 determining achieved cellularity. Cell number variation shows small-berried cultivars achieving 0.8-1.5 million mesocarp cells producing 6-10 mm final diameter, medium-berried cultivars 1.5-2.5 million cells yielding 10-14 mm diameter, large-berried cultivars 2.5-4 million cells achieving 14-18 mm diameter, and cell number strongly correlating with mature berry size. Division spatial patterns show divisions throughout mesocarp flesh tissue creating bulk berry cells, exocarp epidermal and subepidermal divisions producing skin layers, endocarp inner layer divisions adjacent to seeds, and coordinated division maintaining tissue proportions and architecture. Periclinal versus anticlinal divisions show periclinal divisions parallel to berry surface increasing tissue thickness, anticlinal divisions perpendicular to surface increasing tissue area, and coordinated division planes creating three-dimensional tissue expansion. Division completion timing shows progressive cell cycle exit beginning week 3, majority of cells completing division by week 4, occasional continued division in some cell populations extending to week 5, and completion marking definitive transition to expansion phase. Cell size at division exit shows cells relatively small typically 20-40 micrometers diameter enabling efficient tissue packing, cells retaining expansion capacity for subsequent 10-100 fold volume increase, and initial small size characteristic of actively dividing populations. Division phase versus expansion phase contributions show cell number from division accounting for 60-80% mature berry size variation, cell expansion contributing 20-40% size variation, and division phase as more critical determinant emphasizing importance of early favorable conditions. Environmental effects on division phase include temperature critically affecting division rate with optimal 24-28°C supporting maximum proliferation, cool conditions 18-22°C slowing division potentially reducing final cell number 15-30%, warm conditions 28-32°C accelerating division though excessive heat above 35°C impairing function, nitrogen supporting protein synthesis for division, and overall favorable conditions during weeks 1-4 maximizing cell number achievement. Temperature as major factor shows heat accumulation driving division progression, growing degree days during division phase correlating with cell number, and temperature affecting both rate and total divisions determining cellularity. Seed number effects show positive correlation between seeds per berry and final cell number, multiple seeds producing greater cumulative auxin stimulating more intensive division, and auxin dose-response relationship mediating seed effects on cellularity and size. Division irreversibility shows cell number established during weeks 1-4 permanent and unchangeable, lost division potential from suboptimal early conditions irreversible, later favorable conditions unable to compensate for early division limitation, and early environment as critical permanent determinant of berry size potential. The process shows two distinguishable phases including active berry cell division with division activation and onset establishing proliferative program, and peak division activity representing maximum proliferation determining final cellularity. Duration typically 3-5 weeks with peak activity weeks 1-3 and completion by week 4-5. Regional timing shows berry cell division phase during late June-July in warm climates, early-mid July in temperate regions, and mid-late July in cool areas. Viticultural significance includes establishing final cell number as major berry size determinant accounting for 60-80% size variation, demonstrating critical importance of optimal conditions during early weeks for achieving size potential, showing irreversible nature of division phase with permanent effects on berry characteristics, and emphasizing early environment as determinant of quality and yield outcomes. Research applications include analyzing division regulation and cell cycle control, investigating auxin and hormonal effects on proliferation, evaluating temperature and environmental effects on division and cell number, quantifying cell number as size determinant, and assessing division phase as berry development and quality determinant.
Berry_Cell_Division_Phase is first major phase of Berry_Development, distinguished by intensive mitotic activity establishing final cell number. Subdivided into Active_Berry_Cell_Division with division activation and onset, and Peak_Division_Activity with maximum proliferation. Duration 3-5 weeks with peak weeks 1-3. Cell cycle times 10-15 hours. Final cell number 1-4 million determining 60-80% size variation. Temperature and seed number major determinants. Irreversible cell number establishment. Critical phase for berry size potential. Precedes expansion phase.
Beeren-Zellteilungsphase
Zellproliferationsphase
Zellzahl-Etablierungsperiode
berry cell proliferation phase
cell number establishment period
mitotic growth phase
mitotische Wachstumsphase
proliferative berry phase
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 432-438 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Berry Cell Division Phase
Merlot in Bordeaux shows berry cell enlargement phase during mid July-early August Year N+1 at BBCH 73-77, with cells enlarging from 32 micrometers post-division to 95 micrometers by phase end representing 18-fold volume increase, water uptake progressing from 0.4 to 1.6 g per berry per day, massive vacuole development occupying 88% cell volume, berry diameter expanding from 18 mm to 26 mm achieving 85% final size, fresh weight increasing from 6 g to 14 g, under oceanic conditions of 22-28°C temperature with adequate summer rainfall, typical of this mid-season cultivar completing enlargement phase
Berry cell enlargement phase represents the expansion-dominated growth period during berry development when established non-dividing cells progressively enlarge through water uptake and vacuole expansion, cell volume increasing 10-100 fold from post-division size 25-45 micrometers to mature size 80-200 micrometers, turgor-driven growth creating berry volumetric increase, and expansion establishing final berry size approaching genetic potential, proceeding over weeks 3-8 post-fertilization as exclusive or dominant growth mechanism following division completion. This critical expansion phase occurs during mid-late summer in mid July-early August in temperate regions Year N+1, spanning approximately 4-6 weeks with overlap during declining division and pure expansion post-division, corresponding to BBCH 73-77 pea-sized berry to beginning bunch closure stages. Enlargement mechanism involves initial cell enlargement with turgor-driven expansion onset, vacuole enlargement initiation, and cell wall loosening enabling growth, followed by rapid cell expansion with maximum expansion rate, massive vacuole formation occupying 85-95% cell volume, and peak berry growth rate achieving maximum size increase. Cell enlargement fundamentals show turgor pressure as driving force with osmotic water uptake generating 0.5-1.2 MPa pressure, turgor pushing against yielding cell walls causing expansion, water as primary volume contributor comprising 80-85% berry fresh weight, vacuole as major storage compartment expanding to dominate cell volume, and cell wall modification enabling expansion through loosening and synthesis. Water uptake dynamics show progressive increase from 0.3-0.5 g per berry per day during initial enlargement to 1.0-2.5 g per day during rapid expansion, xylem-mediated water import during early enlargement, possible phloem water contribution during later enlargement, and total water import 30-80 g per berry over enlargement phase depending on final size. Osmotic gradient driving water uptake shows solute accumulation creating osmotic potential decline from -0.5 MPa early enlargement to -1.0 to -1.5 MPa during rapid expansion, organic acids particularly malate and tartrate accumulating to 1.0-1.8% fresh weight, potassium and other ions accumulating, sugars beginning accumulation though levels still low 4-8% during enlargement, and combined solutes creating strong osmotic gradient pulling water into cells. Vacuole development progression shows vacuole initiating as small compartments in post-division cells, progressive vacuole enlargement through membrane synthesis and water influx, eventual single large central vacuole occupying 85-95% mature cell volume, and vacuole as major storage compartment for water, ions, organic acids, and later sugars. Vacuole characteristics show tonoplast membrane separating vacuole from cytoplasm, proton pumps and transporters regulating vacuole pH and composition, vacuole pH acidic typically 3.0-3.5 from organic acid accumulation, and vacuole providing osmotic driving force and storage capacity. Cell wall modification enabling expansion shows wall loosening through expansin proteins disrupting hydrogen bonds between cellulose microfibrils and matrix polysaccharides, xyloglucan endotransglucosylase rearranging xyloglucan cross-links, pectin methylesterase modifying pectin affecting wall plasticity, wall plasticity increase enabling turgor-driven expansion, and new wall synthesis maintaining integrity during enlargement preventing bursting. Expansin function shows pH-dependent wall loosening activity, proteins binding to cell wall causing non-hydrolytic disruption, expansin expression correlating with expansion rate, and expansins as key wall-modifying enzymes. Cell wall synthesis during expansion shows new cellulose, hemicellulose, and pectin deposition, wall synthesis maintaining thickness despite area increase, coordinated synthesis with loosening preventing structural failure, and wall composition changes affecting mechanical properties. Cell volume increase quantification shows typical 10-30 fold volume increase during enlargement phase from 25-45 micrometer diameter to 60-120 micrometers by phase end, eventual 30-100 fold total increase including continued expansion during lag and ripening phases, final cell diameter 80-200 micrometers at full maturity, and volume increase as major berry growth contributor. Berry size increase during enlargement shows diameter expanding from 14-25 mm at enlargement onset to 70-90% final size by phase end typically 18-35 mm depending on cultivar, volumetric increase 2-5 fold during pure enlargement phase, fresh weight increase from 4-10 g to 8-20 g by enlargement completion, and substantial size gain establishing near-final dimensions. Berry growth rate characteristics show maximum growth rate during rapid expansion typically 100-250 mg fresh weight per day in large-berried cultivars, sustained high growth rate over 2-4 weeks during peak expansion, growth rate decline as enlargement phase concludes approaching lag phase, and high rate reflecting intensive cell enlargement. Berry composition during enlargement shows water content maintained 80-85% fresh weight, organic acids accumulating to peak levels 1.0-1.8%, sugars low 4-8% from metabolic consumption and limited import, pH declining to 2.8-3.2 from acid accumulation, and overall green hard acidic berry characteristic. Environmental effects on enlargement include temperature affecting expansion rate through metabolic and developmental effects with optimal 22-28°C supporting maximum enlargement, water availability critically affecting expansion with adequate water essential for turgor maintenance and volume increase, water deficit limiting enlargement reducing final berry size 15-40%, nutrient supply particularly potassium affecting water relations and expansion, and overall favorable conditions maximizing size achievement. Temperature effects show warm conditions promoting faster expansion through accelerated metabolism and wall modification, cool conditions slowing expansion rate though generally not preventing size achievement, and temperature as factor affecting timing rather than potential within normal ranges. Water effects show water as most critical environmental factor during enlargement, deficit severely limiting expansion through reduced turgor and water availability, irrigation during enlargement phase essential in water-limited regions for size optimization, and water management as key viticultural practice. Potassium effects show adequate potassium supporting osmotic gradient and water uptake, potassium deficiency potentially limiting expansion, and potassium as important mineral for size achievement. Cultivar variation shows genetic differences in expansion capacity affecting final cell and berry size, small-berried cultivars showing limited expansion reaching 6-10 mm final diameter, large-berried cultivars showing extensive expansion achieving 14-20 mm final diameter, and genetic programming determining expansion potential. Cell number-expansion interaction shows final berry size determined by both cell number from division phase 60-80% and cell expansion 20-40%, multiplicative relationship amplifying effects, and optimal size requiring both adequate cellularity and expansion. Enlargement completion characteristics show expansion rate declining as berry approaches genetic size potential, transition to lag phase as growth slows, berry achieving 70-90% final size by enlargement completion, and establishment of size baseline for subsequent ripening. The process shows two distinguishable phases including initial cell enlargement with turgor-driven expansion onset, vacuole initiation, and wall loosening, and rapid cell expansion with maximum expansion rate, massive vacuole formation, and peak berry growth. Duration typically 4-6 weeks representing weeks 3-8 with overlap during declining division weeks 3-5 and pure expansion weeks 5-8. Regional timing shows berry cell enlargement phase during mid July-early August across climates as expansion proceeds. Viticultural significance includes representing major growth phase establishing near-final berry size, requiring optimal water availability for size achievement, demonstrating critical environmental sensitivity particularly to water deficit, and determining berry size affecting yield and quality through dilution or concentration effects. Research applications include analyzing enlargement mechanisms and regulation, investigating water relations and turgor dynamics, evaluating wall modification processes and regulation, quantifying expansion contribution to berry size, and assessing enlargement as determinant of final berry size and quality.
Berry_Cell_Enlargement_Phase represents expansion-dominated growth period following division. Subdivided into Initial_Cell_Enlargement with turgor onset, vacuole initiation, wall loosening, and Rapid_Cell_Expansion with maximum rate, massive vacuole, peak growth. Duration 4-6 weeks representing weeks 3-8. Cell volume increase 10-100 fold. Water uptake critical driver. Turgor 0.5-1.2 MPa. Vacuole 85-95% final volume. Berry achieves 70-90% final size. Water availability most critical environmental factor. Follows Berry_Cell_Division_Phase. Major size determinant accounting for 20-40% variation.
Beeren-Zellvergrößerungsphase
Beerenvergrößerungs-Periode
Zellexpansionsphase
Zellwachstumsphase
berry enlargement period
cell expansion phase
cell growth phase
expansion-dominated growth
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 440-450 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Berry Cell Enlargement Phase
Berry-cluster synchrony assessment comparing cluster architectures in Cabernet Sauvignon reveals the compactness effect on within-cluster uniformity, with compact clusters (density 8.5 berries/cm rachis) showing moderate synchrony (berry-to-berry Brix CV 8.2% at harvest, exterior berries 24.8 Brix while interior berries only 22.1 Brix due to microclimate differences), while loose clusters (density 4.2 berries/cm) show higher synchrony (CV 4.5%, all berries within 23.5-25.0 Brix range from uniform exposure), demonstrating how cluster architecture affects berry-cluster synchrony with implications for clone and cluster management decisions targeting uniform fruit quality.
Berry cluster synchrony quality is an organ synchrony quality that represents the developmental coordination between individual berries and their parent clusters, and among clusters at different positions within the vine, characterizing the uniformity of fruit development at the cluster organizational level that determines harvest-time cluster homogeneity and the presence or absence of problematic berry subpopulations within clusters. Within the BFO framework, this quality specifically addresses the berry-cluster dimension of organ synchrony, distinguishing coordination at the cluster scale from vine-wide or block-wide synchrony while recognizing that cluster-level synchrony determines the practical unit of fruit quality—whether individual clusters contain uniformly developed berries suitable for quality wine production or mixed populations requiring sorting or compromised quality outcomes. The ontological significance lies in berry-cluster synchrony determining the within-cluster homogeneity that affects both fruit quality and harvest logistics, where synchronized berry development within clusters produces uniform fruit populations that can be harvested, processed, and fermented as coherent units, while asynchronous development creates mixed-maturity clusters that may require berry-level sorting, dilute quality through maturity heterogeneity, or force harvest timing compromises. The positional effects on berry-cluster synchrony include cluster position within shoots (basal versus apical clusters potentially differing in development), berry position within clusters (shoulder berries versus tip berries, interior versus exterior), and the cluster architecture effects that determine berry exposure and resource access (compact clusters showing different synchrony patterns than loose clusters). The cluster compactness interaction with synchrony shows that compact clusters may have reduced internal synchrony due to microclimate variation between exterior (exposed, warmer) and interior (shaded, cooler) berries, while loose clusters may show higher internal synchrony from more uniform berry conditions but potentially lower cluster-to-cluster synchrony from greater position effects. The management factors affecting berry-cluster synchrony include cluster thinning (removing secondary clusters to concentrate resources on remaining clusters), berry thinning (removing lagging berries to improve uniformity), and canopy management (affecting cluster exposure and microclimate uniformity). Understanding berry-cluster synchrony quality enables assessment of within-cluster fruit uniformity, identification of cluster positions or architectures associated with reduced synchrony, and targeted management to improve cluster-level developmental coordination.
Berry_Cluster_Synchrony_Quality addresses developmental coordination at the cluster organizational level determining within-cluster homogeneity. This quality affects practical fruit quality through berry population uniformity within harvest units. The class supports cluster management through berry-cluster synchrony characterization.
Beeren-Trauben-Synchronie
Beerenentwicklungskoordination
Intra-Trauben-Uniformität
Traubenhomogenität
berry developmental coordination
cluster homogeneity
fruit unit synchrony
within-cluster uniformity
Tello J, Ibáñez J (2017) What do we know about grapevine bunch compactness? A state-of-the-art review. Australian Journal of Grape and Wine Research 23(2): 1-10 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 277-316
Berry Cluster Synchrony Quality
The berry color description of Pinot Gris documents the distinctive grey-pink skin color (OIV 225:4, grey) overlying colorless flesh — the skin pigmentation results from the periclinal chimera constitution in which the outer epidermal layer (L1) carries a non-functional VvMYBA1 allele (producing no anthocyanin in the outermost cell layer) while the subepidermal layer (L2) retains a functional allele (producing anthocyanin in the hypodermis), creating the intermediate grey-pink coloration that is neither the deep blue-black of the parent Pinot Noir (where both L1 and L2 produce anthocyanin) nor the green-yellow of the sport Pinot Blanc (where both layers lack functional anthocyanin production). This chimeric color phenotype is unique to the Pinot Gris genotype and illustrates how berry color description must account for the layer-specific pigmentation architecture of periclinal chimeras.
A berry description component that documents the color characteristics of the mature berry, recording both the skin (exocarp) color visible externally and the flesh (mesocarp) color revealed upon cutting or crushing, with berry color being the most immediately visible cultivar identification character and the primary determinant of wine color potential — the distinction between white, rosé, red, and deeply pigmented teinturier cultivars reflects fundamentally different levels of anthocyanin biosynthesis in the skin and flesh tissues controlled by the VvMYBA1/A2 regulatory genes on chromosome 2.
Berry_Color_Description addresses the character that has the most profound commercial and cultural significance in viticulture — berry color determines wine color, which in turn defines the fundamental categories of winemaking (white, rosé, red), the applicable production regulations (appellation rules specifying permitted cultivars by color category), and the consumer perception and market positioning of the resulting wine. The genetic control of berry skin color is remarkably well characterized — the VvMYBA1 and VvMYBA2 genes on chromosome 2 encode R2R3-MYB transcription factors that activate the anthocyanin biosynthetic pathway in berry skin tissue, with loss-of-function mutations in these genes producing white-berried cultivars from pigmented ancestors: the Gret1 retrotransposon insertion in the VvMYBA1 promoter (Kobayashi et al. 2004) is the most common loss-of-function mutation, present in nearly all white V. vinifera cultivars, while deletions, point mutations, and retrotransposon excision events produce the range of color variants (white, grey, pink, red, blue-black) observed across the cultivar spectrum. The independent documentation of skin and flesh color is essential because these represent distinct tissue-specific regulatory programs — in nearly all V. vinifera cultivars, anthocyanin is produced only in the skin (hypodermis), with the flesh remaining colorless regardless of skin pigmentation, but in the rare teinturier cultivars (e.g., Alicante Bouschet, Saperavi), anthocyanin is also produced in the flesh, creating deeply colored juice even before skin contact.
Beerenfarben-Beschreibung
Fruchtfärbungs-Dokumentation
Traubenbeeren-Pigmentierung
berry pigmentation record
fruit coloration documentation
grape color characterization
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Kobayashi S et al. (2004) Retrotransposon-induced mutations in grape skin color, Science 304: 982 Walker AR et al. (2007) White grapes arose through the mutation of two similar and adjacent regulatory genes, Plant Journal 49: 772-785 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia
Berry Color Description
Syrah in Rhône Valley shows berry commitment during mid July Year N+1 at BBCH 71, with berries reaching commitment at 16-22 mm diameter 22-32 days post-fertilization, pericarp differentiation completed, vascular system established, developmental program fully activated, sink strength securing 45-58% photosynthate allocation, irreversible development assured, under warm Mediterranean conditions of 24-30°C temperature, typical of this mid-to-late season cultivar achieving commitment establishing secure fruit set and yield potential
Berry commitment represents the irreversible developmental decision point during berry initiation when berry achieves sufficient structural organization, establishes autonomous developmental capacity, activates full berry developmental program, and reaches assured maturation state capable of completing development to ripe fruit barring catastrophic stress, marking definitive transition from vulnerable initiation to committed development phase. This critical commitment point occurs during fruit development in early-mid July in temperate regions Year N+1, typically reached 2-4 weeks post-fertilization as berry initiation completes, corresponding to BBCH 71 young fruit stages. Commitment establishment involves attainment of critical berry size typically 12-25 mm diameter indicating successful expansion, completion of pericarp tissue differentiation creating functional structure, vascular system establishment providing autonomous resource access, developmental gene expression program full activation establishing self-sustaining development, sink strength establishment securing resource allocation priority, and overall irreversible state where berry development proceeds to maturity unless prevented by severe damage or environmental catastrophe. Critical size achievement shows berry diameter 12-25 mm representing threshold beyond which development typically assured, size reflecting successful cell division and expansion phases, critical mass providing metabolic and structural foundation for continued development, and size as indicator of commitment though not sole determinant. Critical size variation shows cultivar differences with small-berried cultivars reaching commitment at 12-15 mm and large-berried cultivars requiring 20-25 mm, environmental effects on size with favorable conditions enabling smaller commitment size, and size-independent commitment factors including developmental state. Structural organization completion shows pericarp layers fully differentiated into functional exocarp, mesocarp, endocarp, vascular network established providing resource delivery capacity, cellular architecture organized supporting mechanical integrity and metabolic function, and structural maturation creating berry capable of supporting continued expansion and ripening. Developmental program activation shows transition from initiation to committed development gene expression, upregulation of developmental regulators maintaining berry growth trajectory, activation of metabolic pathways preparing for ripening including sugar metabolism and secondary compound synthesis precursors, and establishment of developmental program resistant to reversal. Gene expression characteristics show berry-specific transcriptional programs fully activated, growth-promoting genes maintaining expansion though rate declining, developmental timing genes initiating countdown toward ripening, and integrated gene networks establishing autonomous development. Sink strength establishment shows berry developing strong resource allocation priority competing effectively with vegetative growth, sink strength from metabolic demand and vascular connections, berries securing 40-60% total vine photosynthate allocation, and sink priority ensuring resource delivery supporting development. Sink strength mechanisms show high metabolic activity creating demand, abundant cell number providing consumption capacity, vascular connections enabling resource import, and hormonal signals possibly enhancing sink status. Developmental autonomy characteristics show berry development proceeding largely independently of ongoing seed development after commitment, berry capable of completing ripening even with seed abortion though seed presence beneficial, and autonomous capacity indicating commitment to berry rather than flower fate. Irreversibility indicators show berry development continuing despite environmental fluctuations within tolerable range, resistance to developmental reversal or abortion after commitment, and assured progression to maturity barring catastrophic events. Commitment timing shows typical achievement 2-4 weeks post-fertilization depending on cultivar and conditions, earlier commitment at 2-3 weeks under favorable warm conditions with rapid development, delayed commitment at 3-5 weeks under cool conditions or slow development, and timing reflecting cumulative developmental progress rather than fixed schedule. Pre-commitment vulnerability shows berries before commitment susceptible to abortion from stress, environmental extremes potentially causing berry drop, developmental failure possible from inadequate resources or hormonal support, and uncommitted berries representing tentative fruit set. Post-commitment stability shows committed berries resistant to abortion unless severely stressed, development proceeding reliably under normal to moderately challenging conditions, and commitment representing secure fruit set determining final yield. Commitment assessment methods include observation of continued berry development over weeks confirming commitment, measurement of berry size and growth rate indicating progress toward commitment, gene expression analysis showing committed developmental programs, and population-level berry retention indicating commitment achievement. Berry retention after commitment shows very low abortion rate <5% in committed berries under normal conditions, retention reflecting developmental assurance, and committed berry number determining final yield barring harvest losses. Environmental effects during commitment establishment include temperature affecting developmental rate and commitment timing with optimal 22-28°C supporting rapid commitment achievement within 2-3 weeks, water availability affecting berry growth and commitment through expansion effects, nutrient supply supporting metabolic demands, and overall favorable conditions during critical period ensuring successful commitment. Temperature as major factor shows warm conditions accelerating commitment through faster development, cool conditions delaying commitment through slower progression, and temperature affecting timing rather than commitment capacity under normal ranges. Water stress effects show severe deficit potentially delaying commitment through limited expansion, adequate water supporting normal commitment timing, and water as factor affecting commitment timing and berry size. Failed commitment causes include inadequate early development preventing threshold achievement, severe environmental stress during critical period arresting development, seed abortion after initial development eliminating hormonal support in some cases, or physiological dysfunction though rare after successful initiation. Commitment as yield determinant shows committed berry number establishing final berry count per vine, berry number combined with berry size determining total yield, and commitment as definitive fruit set confirmation for yield prediction. The process represents fourth and final phase of berry initiation marking irreversible development, following early berry formation. Duration represents achieving commitment point typically 2-4 weeks post-fertilization. Cultivar variation includes commitment timing, size threshold, and developmental characteristics. Regional timing shows berry commitment during early-mid July in warm climates, mid July in temperate regions, and mid-late July in cool areas. Viticultural significance includes marking assured fruit set determining final yield, establishing berry development as irreversible and secure, demonstrating successful fruit set completion, and providing definitive basis for yield prediction and crop management. Research applications include analyzing commitment mechanisms and determinants, investigating developmental programs establishing commitment, evaluating environmental effects on commitment timing and success, quantifying commitment indicators and thresholds, and assessing commitment as fruit set and yield determinant.
Berry_Commitment is fourth and final phase of Berry_Initiation, distinguished by irreversible developmental decision point. Represents achievement of assured maturation capacity typically 2-4 weeks post-fertilization at 12-25mm diameter. Involves structural completion, developmental program activation, sink strength establishment. Marks transition from vulnerable initiation to secure development. Post-commitment abortion rate <5%. Determines final berry number and yield. Completes Fruit_Set process. Transitions to Berry_Development phase. Critical fruit set completion indicator.
Beeren-Commitment
Entwicklungs-Verpflichtung
Fruchtansatz-Vollendung
berry development assurance
developmental commitment
fruit set completion
irreversible Beerenentwicklung
irreversible berry development
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 422-430 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Berry Commitment
Berry composition data for Cabernet Sauvignon from Oakville, Napa Valley (2023 vintage) documents mean berry weight 1.32 g (n=200), skin weight 0.27 g (20.5% of berry mass), seed weight 0.064 g (4.8%, mean 2.1 seeds per berry), and pulp weight 0.99 g (74.7%), yielding a skin-to-pulp ratio of 0.27 — characteristic of well-developed but not excessively small berries from moderate deficit irrigation management. Berry composition data for Pinot Noir from Gevrey-Chambertin (2024 vintage) records berry weight 1.18 g, skin weight 0.19 g (16.1%), seed weight 0.052 g (4.4%), and pulp weight 0.94 g (79.5%), with the relatively thin skin fraction typical of Pinot Noir explaining the cultivar's characteristically moderate tannin extraction and delicate color that distinguishes it from thick-skinned varieties.
Berry composition data is a quality analysis data entity that documents the physical partitioning of grape berries into their constituent tissue components — skin, pulp (mesocarp), and seeds — through gravimetric dissection analysis, recording the absolute and relative mass of each tissue fraction and their derived ratios, providing the structural characterization that determines the potential extraction of phenolic compounds, aromatic precursors, and flavor-active molecules during winemaking and serving as a critical predictor of wine concentration, color depth, tannin structure, and overall quality potential independent of must chemistry measurements.
Berry composition data bridges the gap between vineyard grape assessment and winemaking outcome prediction by quantifying the physical structures from which phenolic, aromatic, and flavor compounds are extracted during maceration and fermentation. The fundamental principle underlying berry composition analysis is that wine quality-determining compounds (anthocyanins, tannins, flavor precursors) are concentrated in the skin and seed tissues, while the pulp contributes primarily sugar, water, and organic acids — therefore, berries with higher skin and seed proportions relative to pulp deliver more concentrated must and wine at equivalent sugar levels. The methodology requires careful manual dissection of representative berry samples (typically 50-200 berries) under controlled conditions, separating frozen or fresh berries into skin, seed, and pulp fractions with subsequent gravimetric analysis — a labor-intensive procedure that limits routine adoption but provides irreplaceable structural information. Berry composition is determined by the interaction of genetics (cultivar-specific skin thickness, seed number, and berry size potential), environment (water deficit reduces berry size and increases skin-to-pulp ratio, while ample water promotes pulp expansion), and management (crop load, irrigation timing, canopy exposure). The analysis should be performed at the anticipated harvest date or across multiple dates during the maturation period to capture the dynamic evolution of tissue proportions, as berry shrinkage during late-season dehydration can substantially alter composition ratios compared with earlier sampling. Modern near-infrared (NIR) sorting technology in wineries exploits berry composition differences to segregate incoming fruit by quality level, making berry composition data increasingly relevant for precision winemaking applications.
Beerengewebedaten
Beerenkomponentenanalyse
Beerenpartitionierungsdaten
Beerenzusammensetzungsdaten
berry component data
berry dissection data
berry partitioning data
berry physical composition
berry tissue analysis
physikalische Beerenanalyse
Roby G et al. (2004) Berry size and vine water deficits as factors in winegrape composition: anthocyanins and tannins. Australian Journal of Grape and Wine Research 10(2): 100-107 Iland P et al. (2011) Chemical Analysis of Grapes and Wine, 2nd Ed, Patrick Iland Wine Promotions, pp.: 78-88 Walker RR et al. (2005) Rootstock effects on salt tolerance of irrigated field-grown grapevines. Australian Journal of Grape and Wine Research 11(2): 174-183
Berry Composition Data
Pinot Noir berry density monitoring in the Willamette Valley shows progression from 1.058 g/mL at véraison to 1.092 g/mL at optimal harvest (24.2°Brix), with density measurement providing earlier detection of ripening plateau than Brix measurement alone, enabling harvest scheduling optimization that captures peak flavor development occurring 3-5 days before maximum sugar accumulation.
Berry density quality is a tissue density quality that inheres in the whole grape berry as an integrated unit and represents the composite mass per unit volume reflecting the combined contributions of all berry compartments including pulp, skin, seeds, and vascular tissues weighted by their respective volumes and individual densities. Within the BFO framework, this quality characterizes the intact berry as a biological entity whose density emerges from tissue organization rather than being directly measurable as a homogeneous material, distinguishing it from must density measured after tissue disruption. The ontological significance lies in berry density serving as a non-destructive maturity indicator that integrates multiple compositional parameters, where the characteristic density increase during ripening results primarily from sugar accumulation in the mesocarp that increases vacuolar solute concentration from approximately 5% to 25% total soluble solids. Berry density measurement methodologies include individual berry weighing combined with volume displacement or geometric calculation, flotation in calibrated density solutions, and batch measurements using hydrostatic techniques, with flotation methods enabling rapid sorting of large berry populations. The developmental trajectory shows berry density increasing from approximately 1.02 g/mL at fruit set to 1.06-1.10 g/mL at véraison, then continuing to rise during ripening to 1.08-1.14 g/mL at harvest depending on sugar accumulation and dehydration extent. Cultivar differences in berry density reflect both compositional variation and structural differences including skin thickness, seed number, and pulp cell size that affect the volume-weighted integration of tissue densities. Berry density correlates strongly with sugar content but provides additional information about tissue hydration status and structural integrity not captured by refractometry alone.
Berry_Density_Quality specifically addresses the intact whole berry, distinguishing from Tissue_Density_Quality of individual compartments and from Must_Density_Quality measured after crushing. This quality serves as the primary sorting parameter in density-based grape processing and correlates with but is not equivalent to sugar content due to additional influences of tissue hydration and structural organization. Measurement should specify methodology as values differ between flotation, displacement, and calculated approaches.
Beerendichte
Beerenspezifisches Gewicht
Gesamtbeerendichte
berry mass-volume ratio
composite berry density
intact berry specific gravity
intakte Beerendichte
whole berry density
Kontoudakis N et al. (2011) Influence of wine pH on changes in color and polyphenol composition induced by micro-oxygenation. Journal of Agricultural and Food Chemistry 59(5): 1974-1984 Rolle L et al. (2012) Instrumental texture analysis parameters as markers of table-grape and winegrape quality. American Journal of Enology and Viticulture 63(1): 11-20
Berry Density Quality
The berry description of Cabernet Sauvignon documents small, round berries (OIV 220:3, OIV 223:2) with thick, deeply pigmented blue-black skin (OIV 225:6, OIV 228:7), firm crisp flesh with colorless juice (OIV 230:1), 1-3 well-developed seeds per berry, a distinctive herbaceous-cassis aroma profile (OIV 236:1-2), and a mean berry weight of 1.0-1.5 g — this combination of small size, thick dark skin, and high skin-to-pulp ratio (0.15-0.20 by weight) is the morphological foundation of Cabernet Sauvignon's winemaking character, providing the concentrated anthocyanin extraction (1,500-2,500 mg/kg total anthocyanin in skin), abundant tannin (3,000-5,000 mg/kg skin tannin), and structured, age-worthy wine style that has made Cabernet Sauvignon the world's most widely planted premium red wine grape.
An ampelographic description component that documents the morphological characteristics of the individual mature berry at harvest — the fleshy fruit developed from the fertilized ovary — recording size, shape, skin color, skin thickness, flesh texture and color, flavor characteristics, seed number, and the overall sensory profile that defines the cultivar-specific berry phenotype with direct consequences for wine style, table grape appeal, and processing suitability.
Berry_Description addresses the individual fruit that is the fundamental unit of grape production — whether destined for wine, table consumption, raisin drying, juice, or distillation, the berry's morphological and compositional characteristics determine the quality and style of the final product. The grape berry is botanically a true berry (a fleshy fruit developed from a single ovary without a stone), consisting of the exocarp (skin, comprising the outer epidermis, hypodermis containing pigments, and several layers of thick-walled cells), the mesocarp (flesh or pulp, the bulk of the berry consisting of large, thin-walled parenchyma cells containing the juice), and the endocarp (a thin layer surrounding the seeds): the relative development and composition of these tissue layers varies dramatically between cultivars and constitutes the morphological basis of cultivar suitability for different end uses — thick-skinned, small-berried cultivars with high skin-to-pulp ratio (e.g., Cabernet Sauvignon, Tannat) produce deeply colored, tannic wines thin-skinned, large-berried cultivars with low skin-to-pulp ratio (e.g., Chasselas, Italia) are preferred for table grape consumption where crisp texture and mild flavor are desired and muscat-flavored cultivars with aromatic compounds concentrated in the skin and sub-skin layers (e.g., Muscat of Alexandria, Gewürztraminer) produce the aromatic wines and table grapes valued for their distinctive fragrance.
Beeren-Beschreibung
Fruchtmorphologie-Dokumentation
Traubenbeeren-Charakterisierung
berry phenotype documentation
fruit morphology description
grape berry characterization
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Coombe BG and McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening, Australian Journal of Grape and Wine Research 6: 131-135
Berry Description
Berry detachment resistance quality determines harvest timing optimization in Pinot Noir for sparkling wine production, with whole-cluster pressing protocols requiring high detachment resistance (force above 3.5 N) to maintain intact berries during transport from vineyard to press, preventing juice oxidation and phenolic extraction that would compromise base wine finesse harvest crews monitor detachment resistance daily during the picking window, delaying harvest if values fall below threshold after rain events that hydrate abscission zones and temporarily reduce berry retention.
Berry_Detachment_Resistance_Quality represents the attachment strength quality specifically characterizing the resistance of grape berries to separation from their pedicels under applied force. This quality addresses the same anatomical junction as Pedicel_Attachment_Quality but emphasizes the practical resistance to detachment that determines harvest efficiency, handling tolerance, and shatter susceptibility. Detachment resistance is measured as the pull force required to separate a berry from its pedicel, typically expressed in Newtons (N), with values ranging from less than 1 N in loose-berried cultivars to greater than 6 N in tight-attachment varieties. The force required for detachment depends on the structural development of the attachment zone, the presence and maturity of abscission layer tissues, turgor pressure that affects tissue rigidity, and environmental factors including temperature and humidity at the time of assessment. High detachment resistance protects against shatter during transport and display but may complicate harvest, particularly mechanical harvest where removal relies on impact forces rather than cutting. Low detachment resistance facilitates harvest but causes problematic berry loss at every handling step from field to consumer. The mode of detachment (clean abscission versus tearing) affects both fruit quality and susceptibility to decay, with clean separation leaving a dry scar less vulnerable to infection than torn tissue with exposed juice. Assessment of detachment resistance employs calibrated pull-force measurement on samples of attached berries, with protocols standardizing pull direction, pull rate, and the method of gripping berries during measurement.
Berry_Detachment_Resistance_Quality addresses the force opposing berry separation and complements Pedicel_Attachment_Quality which addresses the structural characteristics of the connection. Detachment resistance is the primary functional measure for practical quality assessment, while pedicel attachment provides anatomical context for understanding resistance values. The force values should be interpreted relative to cultivar norms, as acceptable ranges differ substantially among varieties. Temperature affects measurement values, with cold fruit typically showing higher detachment resistance than warm fruit, requiring standardization for comparable results.
Ablösewiderstand
Abtrennkraft
Beerenhaftung
Schüttelfestigkeit
berry pull resistance
berry retention force
detachment force
shatter resistance
Fidelibus MW et al. (2007) Girdling and Gibberellic Acid Effects on Yield Components, Fruit Composition, and Fruit Skin Adhesion in 'Crimson Seedless' Table Grapes. Acta Horticulturae 754: 461-465 Beyaz R and Yildiz M (2017) Artificial Abscission Zone Formation in Grape Berry Pedicel by Application of Plant Growth Regulators. Scientia Horticulturae 225: 328-335
Berry Detachment Resistance Quality
Cabernet Sauvignon in Napa Valley shows berry development during mid July-late October Year N+1 at BBCH 71-89, with cell division phase weeks 0-4 establishing 2.4-3.2 million cells, expansion phase weeks 3-8 achieving 12-15 mm diameter, lag phase weeks 8-10, veraison week 10-12 initiating color change and sugar accumulation, ripening weeks 12-18 completing maturation reaching 24-26% sugar, 0.5-0.7% acid, full anthocyanin development, under warm climate conditions of 24-32°C, typical of this late-season cultivar achieving harvest maturity 110-130 days post-fruit set
Berry development represents the comprehensive developmental progression from fruit set commitment through complete ripening when berry undergoes sequential coordinated phases including intensive cell division establishing cellular foundation, rapid expansion through cell enlargement, lag phase representing metabolic transition, veraison initiating ripening transformation, and progressive ripening achieving full maturity, spanning approximately 90-120 days from fruit set to harvest and determining berry composition, quality, and wine characteristics. This complete developmental sequence occurs during summer to autumn in late June-October in temperate regions Year N+1, encompassing entire berry maturation from young fruit to ripe berry, corresponding to BBCH 71-89 stages from young fruit development through full ripeness. Berry development encompasses multiple sequential phases including berry cell division phase with intensive mitotic activity weeks 0-4 establishing final cell number, berry expansion phase with cell enlargement weeks 3-8 achieving primary size, lag phase weeks 6-10 representing metabolic reorganization and growth quiescence, veraison weeks 8-12 initiating ripening transformation with color change and sugar accumulation onset, and ripening phase weeks 10-16 completing maturation through sugar accumulation, acid decline, and flavor development. Cell division phase characteristics show peak mitotic activity during first 3-4 weeks post-fertilization establishing mesocarp flesh cellularity, cell cycle times 10-15 hours during maximum division intensity, final cell number 1-4 million cells per berry determining size potential, auxin-stimulated division from developing seeds, and division completion by week 4 marking transition to expansion-dominated growth. Expansion phase characteristics show cell enlargement through water uptake and vacuole expansion weeks 3-8, volumetric increase 3-8 fold during rapid expansion, berry achieving 70-90% final size by expansion completion, water comprising 80-85% fresh weight, and expansion rate progressively declining toward lag phase. Lag phase characteristics show growth rate declining to near-zero creating quiescent period weeks 6-10, berry maintaining green hard acidic state, metabolic reorganization preparing ripening transformation, seed maturation completing, and lag duration 1-3 weeks varying by cultivar and temperature before veraison onset. Veraison characteristics show dramatic transformation initiating weeks 8-12 with berry softening from cell wall modification, color change in red cultivars from anthocyanin synthesis or color revelation in white cultivars from chlorophyll degradation, sugar accumulation accelerating from phloem unloading activation, acid decline beginning particularly malate respiration, and overall transition from growth to ripening metabolism. Ripening phase characteristics show progressive sugar accumulation reaching 18-26% fresh weight as hexoses, continued acid decline to 0.4-0.8% particularly malic acid, secondary metabolite synthesis including anthocyanins, flavor compounds, and tannins, berry softening progression, and maturation completing over 4-6 weeks achieving harvest ripeness. Berry growth pattern shows characteristic double sigmoid curve with rapid growth phase I during cell division and expansion weeks 0-6, lag phase representing growth plateau weeks 6-10, and renewed growth phase II during ripening from sugar and water accumulation weeks 10-16. Fresh weight accumulation shows progressive increase from 0.5-2 g at commitment to 2-8 g at lag phase to 4-15 g at harvest depending on cultivar, weight increase rate varying through phases with maxima during expansion and ripening, and final weight determined by cell number, cell expansion, and ripening accumulation. Berry composition changes show dramatic transformation from hard green acidic berry at 2-5% sugar, 1-2% acid, pH 2.5-3.5 during growth phase, through intermediate composition during lag and veraison, to ripe berry at 18-26% sugar, 0.4-0.8% acid, pH 3.0-4.0 with accumulated flavor compounds and color. Sugar accumulation dynamics show low levels <5% during growth phase from metabolic consumption, accelerating accumulation from veraison through ripening, hexose sugars glucose and fructose dominating in approximately equal concentrations, and final concentration 18-26% representing 40-60% dry weight. Acid dynamics show initial accumulation to 1-2% during growth particularly malate and tartrate, peak acidity at veraison representing maximum organic acid content, progressive decline during ripening through malate respiration and dilution, tartrate relatively stable though declining slightly, and final acidity 0.4-0.8% at physiological maturity. pH changes show low pH 2.5-3.0 during growth phase from high acidity, progressive increase during ripening from acid decline, and final pH 3.0-4.0 at harvest with considerable cultivar and environmental variation. Color development shows green chlorophyll-containing berry during growth and lag phases, color change onset at veraison through anthocyanin synthesis in red cultivars or chlorophyll degradation revealing carotenoids in white cultivars, progressive color intensification during ripening, and final color determining wine hue. Anthocyanin accumulation in red cultivars shows synthesis initiating at veraison, progressive accumulation during ripening reaching 0.5-3.0 mg/g fresh weight, anthocyanin profile development with multiple cyanidin, peonidin, and malvidin glycosides, and accumulation determining wine color intensity and hue. Flavor compound development shows volatile and non-volatile compound synthesis during ripening, varietal aroma precursors accumulating including terpenes in aromatic varieties, methoxypyrazines declining from green to ripe states, esters and other volatiles developing, and flavor complexity increasing toward maturity. Texture changes show firm hard berry during growth phase from turgor and cell wall integrity, softening onset at veraison from pectin degradation and cell wall modification, progressive softening during ripening, and final texture ranging from firm to very soft depending on cultivar and maturity. Seed development coordination shows seeds developing in parallel with berry, seed maturation largely completing by lag phase or early veraison, seed color change from green to brown indicating maturation, and mature seeds with developed embryo, endosperm, and seed coat. Environmental effects on development include temperature as major factor affecting development rate with warm conditions accelerating and cool conditions slowing progression through phases, water availability affecting expansion and ripening accumulation, light possibly influencing anthocyanin synthesis and flavor development, and overall growing conditions determining development timing, berry composition, and quality. Temperature effects show heat accumulation driving phenological progression, growing degree days determining development timing, optimal ranges varying by phase with 24-28°C for growth, 22-28°C for ripening, and temperature affecting development rate and final composition. Water effects show adequate water essential during expansion for size achievement, deficit potentially beneficial during ripening enhancing concentration and quality in some cases, and water management strategies affecting berry development and wine quality. Cultivar variation shows genetic differences in development timing from early-ripening to late-ripening cultivars spanning 90-150 days fruit set to harvest, cell number and berry size characteristics, sugar and acid accumulation patterns, color and flavor development, and overall composition profiles determining varietal wine characteristics. Regional timing shows berry development during July-October in warm climates completing earlier, mid July-late October in temperate regions, and late July-November in cool climates with later harvest, demonstrating climatic effects on development duration. Viticultural significance includes representing critical determinant of berry composition and wine quality, requiring optimal environmental conditions throughout development for quality achievement, demonstrating sequential coordinated developmental program, and establishing berry characteristics affecting all wine attributes including color, flavor, body, and balance. Research applications include analyzing developmental regulation and phase transitions, investigating environmental effects on development and composition, evaluating genetic variation in developmental characteristics, quantifying composition changes through development, and assessing development as quality determinant enabling harvest timing optimization and quality prediction.
Berry_Development represents complete developmental sequence from fruit set through full ripeness. Encompasses Berry_Cell_Division_Phase, Berry_Expansion_Phase, Lag_Phase, Veraison, and Ripening_Phase. Duration 90-120 days varying by cultivar and climate. Sequential coordinated phases determining berry composition and quality. Temperature major driver of development rate. Comprehensive documentation of berry maturation establishing wine quality foundation. BBCH 71-89 stages. Primary viticultural focus for quality achievement.
Beerenentwicklung
Beerenreifung
Fruchtentwicklung
Traubenentwicklung
berry maturation
berry ripening progression
fruit development
grape development
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 430-520 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Berry Development
Berry development rate assessment comparing irrigation regimes in Cabernet Sauvignon reveals the water management effect on ripening dynamics, with full irrigation showing slower berry development rate (52 days from veraison to 24 Brix) maintaining berries in the cooling effect of transpiration, while regulated deficit irrigation shows accelerated rate (43 days to 24 Brix) from concentration effects and reduced berry size, with the 9-day difference shifting harvest into different weather windows and affecting the accumulation dynamics of sugars, acids, and phenolics that determine wine composition, demonstrating how management-induced rate variation affects fruit quality through ripening duration and timing effects.
Berry development rate quality is a developmental rate quality that represents the speed of berry growth, ripening, and maturation from fruit set through harvest, characterizing the rate of fruit development that determines the duration of the ripening period and the temporal alignment of maturity achievement with seasonal climate conditions critical for wine quality. Within the BFO framework, this quality specifically addresses the fruit developmental dimension of phenological rate, distinguishing berry development rate from earlier phenological phases while recognizing that berry development rate determines when harvest maturity is achieved relative to the declining temperatures and increasing precipitation risk of late season, making this rate a primary determinant of the thermal and weather conditions under which final fruit composition develops. The ontological significance lies in berry development rate determining the ripening window duration and the climate conditions during the final maturation phase that most directly affects wine quality, where rapid development compresses ripening potentially into hot conditions (accelerating sugar accumulation but potentially compromising flavor development), while slow development extends ripening into cooler conditions (favoring flavor complexity but risking incomplete ripening or harvest-period weather damage). The biphasic pattern of berry development creates distinct rate dynamics, with rapid growth during Phase I (fruit set to lag phase, 30-60 days, cell division and expansion driving volume increase), minimal development during the lag phase (1-2 weeks, metabolic transition), and resumed growth during Phase II (veraison to harvest, 35-55 days, cell expansion and ripening compound accumulation), with temperature affecting rates differently in each phase. The cultivar variation in berry development rate distinguishes early-ripening cultivars (Pinot Noir, Chardonnay requiring less thermal time from veraison to maturity) from late-ripening cultivars (Cabernet Sauvignon, Mourvèdre requiring more thermal time), with this variation providing the basis for cultivar-climate matching to achieve ripening under optimal temperature conditions. The management factors affecting berry development rate include crop load (heavy crops slowing development through source-sink competition), water status (deficit irrigation often accelerating ripening through concentration effects), and canopy management (fruit exposure affecting berry temperature and development rate), with these tools providing options for manipulating ripening timing relative to target harvest windows. Understanding berry development rate quality enables prediction of harvest timing, assessment of ripening conditions, and management strategies for optimizing the climate conditions during the maturation period most critical for wine quality.
Berry_Development_Rate_Quality addresses the speed of fruit maturation determining ripening duration and harvest timing. This quality shows biphasic patterns and responds to temperature, crop load, and water status. The class supports harvest prediction and ripening management through berry development rate characterization.
Beerenentwicklungsrate
Beerenwachstumsrate
Fruchtreifungsrate
Reifungsgeschwindigkeit
berry growth rate
fruit maturation rate
maturation velocity
ripening rate
Coombe BG, McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6(2): 131-135 Sadras VO, Moran MA (2012) Elevated temperature decouples anthocyanins and sugars in berries of Shiraz and Cabernet Franc. Australian Journal of Grape and Wine Research 18(2): 115-122
Berry Development Rate Quality
Berry firmness quality in Thompson Seedless (Sultanina) defines harvest windows for different market destinations, with early harvest at 220-280 g compression force providing firmness for fresh market despite incomplete sugar accumulation, mid-harvest at 150-200 g offering balance between texture and flavor for domestic retail, and late harvest below 120 g yielding soft berries suitable only for raisin production or immediate processing this firmness-guided harvest segregation optimizes quality outcomes across the variety's multiple end uses.
Berry_Firmness_Quality represents the physical quality characterizing the integrated mechanical resistance of whole grape berries to deformation, capturing the combined contribution of skin, pulp, and internal structure to overall berry texture. Whole-berry firmness measurement provides the most practically relevant texture assessment for handling and storage quality prediction, as berries experience forces at the whole-berry level during harvest, transport, and processing. The mechanical response of intact berries reflects the turgor-maintained structure where the pressurized pulp cells provide internal support while the skin resists expansion and rupture. Berry firmness typically ranges from 1-4 N in compression tests and 0.5-3 N in puncture tests, with variation reflecting cultivar, ripeness, and physiological status. Firmness decreases substantially at véraison (40-60% reduction in many varieties) as cell wall loosening and turgor changes accompany the transition to ripening. Post-véraison firmness trajectories vary among cultivars, with some maintaining firmness through harvest (Autumn Royal, Crimson Seedless) while others soften progressively (Thompson Seedless, most wine grapes). Berry firmness correlates with handling durability, with firmer berries showing reduced damage from equivalent mechanical stress. The relationship between instrumental berry firmness and sensory texture perception (crispness, crunchiness, softness) depends on test geometry and cultivar, requiring validation for specific applications. Berry firmness measurement employs texture analyzers with compression platens or puncture probes, with standardized protocols specifying probe geometry, test speed, and sample positioning for reproducible results. The force-deformation curve provides firmness parameters including maximum force, slope (stiffness), and work (energy absorption).
Berry_Firmness_Quality addresses the integrated whole-berry mechanical response as distinct from tissue-specific firmness measurements. The whole-berry test captures the functional mechanical behavior during handling while concealing the individual tissue contributions that tissue-specific testing reveals. Berry firmness measurement is relatively straightforward technically, requiring only standardized compression or puncture protocols without the sample preparation challenges of tissue isolation. The class represents the most commonly measured firmness parameter in practical grape quality assessment.
Beerenfestigkeit
Beerenhärte
Gesamtbeerenfestigkeit
composite berry firmness
intact berry texture
intakte Beerentextur
overall berry hardness
whole-berry firmness
Rolle L et al. (2011) Berry Density and Size as Factors Related to the Physicochemical Characteristics of Muscat Hamburg Table Grapes (Vitis vinifera L.). Food Chemistry 125(4): 1385-1393 Maury C et al. (2019) Texture Analysis of Fruits and Vegetables. In: Ferranti P et al. (eds) Encyclopedia of Food Chemistry, Elsevier pp.: 452-458
Berry Firmness Quality
The berry flesh description of Muscat of Alexandria documents soft, melting flesh with abundant free-run juice (OIV 235:2, slightly firm to soft), an intensely aromatic muscat flavor dominated by linalool and geraniol monoterpenes (OIV 236:2, muscat flavor), and high juiciness with juice release upon gentle finger compression — this combination of soft texture, intense aroma, and abundant juice makes Muscat of Alexandria a dual-purpose cultivar valued both as a fresh table grape (where the aromatic flesh flavor is a primary quality attribute) and as a winemaking grape (where the free-run juice is rich in terpene precursors that produce the characteristically floral Muscat wines). The soft, aromatic Muscat of Alexandria flesh contrasts with the firm, crunchy, neutrally flavored flesh of Crimson Seedless (bred for table grape crunch with minimal aromatic character) and the moderately firm, herbaceous-flavored flesh of Cabernet Sauvignon (where the flesh flavor is subordinate to the skin tannin and color contributions to wine character).
A berry description component that documents the physical and sensory characteristics of the berry flesh (mesocarp) — the bulk tissue between the skin and the seeds constituting 75-85% of berry weight — recording consistency (firmness and texture), flavor profile (aromatic character and taste), and juiciness (the volume and ease of juice release upon compression), with flesh properties determining the eating quality for table grape assessment and the must composition and processing behavior for winemaking.
Berry_Flesh_Description addresses the tissue that constitutes the majority of the berry by mass and volume and provides virtually all of the juice extracted during pressing — the must composition (sugar, acid, pH, aroma precursors) that determines wine style is overwhelmingly derived from the flesh cells, while the phenolic compounds (color, tannin) are contributed by the skin and seed. The flesh consists of large, thin-walled parenchyma cells (150-400 μm diameter at maturity) that accumulate sugars (glucose and fructose, reaching 200-280 g/L at harvest), organic acids (primarily tartaric and malic, declining from 15-20 g/L at véraison to 5-9 g/L at harvest), and aroma precursors (terpenoids, norisoprenoids, thiols, and other volatile precursors in cultivar-specific profiles): the dramatic compositional changes during ripening from véraison to harvest — sugar doubling, acid declining by half, aroma precursors accumulating — transform the flesh from a hard, sour, unpalatable tissue to the sweet, aromatic, juice-filled flesh of the ripe berry. The physical properties of the flesh are determined by cell turgor, cell wall composition, and intercellular adhesion — firm-fleshed cultivars maintain high turgor pressure and strong intercellular bonds that resist deformation (the crisp table grape ideal), while soft-fleshed cultivars have lower turgor and weaker intercellular adhesion that allows easy cell separation and juice release (the juicy wine grape or eating grape character).
Beerenfleisch-Bewertung
Fruchtfleisch-Beschreibung
Mesokarp-Charakterisierung
berry flesh assessment
mesocarp characterization
pulp properties description
Berry Flesh Description
Berry fruit aroma assessment in Pauillac Cabernet Sauvignon identifies the varietal signature of blackcurrant and cassis (8.5 on 10-point scale) with supporting blackberry (6.2) and blueberry (4.8) notes, while Saint-Émilion Merlot-dominant blend shows shifted profile with blackberry (7.8) and plum (6.5) dominating over cassis (4.2), demonstrating how the specific berry balance distinguishes these neighboring appellations and their dominant varieties even when overall berry intensity is similar.
Berry fruit aroma quality is a fruity aroma quality that represents olfactory perception evoking small berry fruit-associated scents including blackberry, raspberry, blueberry, blackcurrant (cassis), redcurrant, strawberry, and cranberry notes, characterizing the concentrated, often intense aromatic impression that serves as the primary fruit descriptor category for most red wine varieties. Within the BFO framework, this quality addresses the dominant fruit aroma subcategory for red wines, distinguishing berry character from stone fruit while recognizing that berry descriptors provide the core varietal fruit identity for varieties from Cabernet Sauvignon (blackcurrant) through Pinot Noir (raspberry, cherry) to Zinfandel (blackberry, raspberry). The ontological significance lies in berry aroma defining red wine varietal identity more than any other descriptor category, where the specific berry note serves as the primary varietal marker (cassis for Cabernet Sauvignon, raspberry for Pinot Noir, blackberry for Syrah) and berry intensity indicates fruit quality and extraction success. The compositional basis for berry aroma includes multiple compound classes, with β-damascenone contributing to general berry character, furaneol and homofuraneol providing strawberry notes, specific esters contributing raspberry character, and methoxypyrazines (at low levels) enhancing the cassis perception in Cabernet family varieties. The color association between red, blue, and black berries and red wine creates strong perceptual expectations, with visual color intensity enhancing berry aroma perception through cross-modal interaction, demonstrating the integration of visual and olfactory processing in berry character assessment. The ripeness continuum within berry descriptors proceeds from tart red berries (redcurrant, cranberry suggesting underripe or cool) through ripe mixed berries (raspberry, blackberry) to jammy cooked berries (indicating very ripe or high extraction), providing quality-relevant information within this category. Understanding berry fruit aroma quality enables varietal identification based on signature berry profile, ripeness and extraction assessment from berry character development, and quality evaluation where appropriate berry expression indicates successful red wine production.
Berry_Fruit_Aroma_Quality addresses the small berry fruit subcategory of fruity aroma, encompassing blackberry, raspberry, blackcurrant, blueberry, strawberry, and related notes. This quality provides the primary fruit descriptors for red wine varieties, with specific berry notes serving as varietal markers. The red-to-black berry spectrum and fresh-to-jammy continuum provide quality-relevant information within this category.
Beerencharakter
Beerenduft
Beerenfruchtaroma
Beerenfruchtduft
berry aroma
berry character
berry scent
small fruit notes
Escudero A et al. (2007) Characterization of the aroma of a wine from Maccabeo. Key role played by compounds with low odor activity values. Journal of Agricultural and Food Chemistry 55(11): 4501-4510 Pineau B et al. (2009) Which impact for β-damascenone on red wines aroma? Journal of Agricultural and Food Chemistry 55(10): 4103-4108
Berry Fruit Aroma Quality
Chardonnay in Champagne shows berry growth lag phase during late August-September Year N+1 at BBCH 81, with growth rate declining to 2-4 mg per day establishing quiescence, berry dimensions stable at 8.2-8.6 mm diameter and 2.8-3.2 g fresh weight, seeds completing maturation with hard brown coats, sugars increasing slowly from 11% to 14%, acids declining from 1.1% to 0.9%, pH rising from 3.0 to 3.2, lag duration 2.5-3.5 weeks under cool maritime conditions of 18-24°C temperature, typical of this early-bursting white cultivar before veraison onset
Berry growth lag phase represents the growth quiescence period during berry development when berry achieves minimal growth rate typically <5 mg fresh weight per day, berry maintains relatively stable dimensions and composition while metabolically active, seed maturation completes establishing developmental readiness, metabolic reorganization prepares ripening machinery for veraison transformation, and lag phase creates temporal plateau between growth phase I and ripening phase II in characteristic double sigmoid growth pattern, proceeding over 1-4 weeks duration varying by cultivar and environment during mid-late summer to early autumn. This quiescent transition phase occurs during late summer to early autumn in late August-September in temperate regions Year N+1, spanning variable duration 1-4 weeks between enlargement completion and veraison onset, corresponding to BBCH 81 beginning of ripening stage representing pre-veraison period. Lag phase mechanism involves growth rate decline to minimal levels establishing quiescence, metabolic transition from growth-phase to pre-ripening metabolism, seed maturation completion enabling developmental progression, berry compositional stability with minimal changes, metabolic preparation activating ripening pathway components, and integrated quiescence creating developmental pause before veraison. Lag phase initiation shows transition from enlargement completion with growth rate declining from 10-30 mg per day to <5 mg per day, metabolic transition onset shifting gene expression and enzyme activities, compositional changes from late enlargement continued into early lag, and establishment of quiescent state. Minimal growth period characteristics show near-zero growth rate typically 1-5 mg fresh weight per day representing essential growth cessation, berry dimensions stable within 1-3% variation, seed development focus with seeds completing maturation, pre-veraison metabolism with subtle metabolic preparations for ripening, and sustained quiescence over 1-4 weeks. Growth rate during lag phase shows minimal berry fresh weight gain <5 mg per day typical, occasional slightly negative growth from water loss in some conditions, essentially stable berry mass distinguishing lag from growth and ripening phases, and minimal rate as defining lag characteristic. Berry size stability shows dimensions remaining essentially constant during lag phase, diameter variation <1-3% from measurement variability and minor fluctuations, fresh weight stability within 2-5% representing equilibrium, and size constancy contrasting with enlargement growth and ripening expansion. Berry composition during lag phase shows water content 78-82% relatively stable, organic acids declining slowly from 0.9-1.3% to 0.8-1.2%, sugars increasing slowly from 10-14% to 12-16%, pH rising gradually from 3.0-3.2 to 3.1-3.3, and overall composition showing slow changes toward ripening. Organic acid dynamics show malate continuing slow respiration declining gradually, tartrate relatively stable maintaining concentrations, total acids declining slowly contributing to pH increase, and acid metabolism as continuing process during quiescence. Sugar dynamics show sugars increasing slowly from limited phloem import, sugar accumulation rate low 0.5-1.5% per week, gradual sugar increase preparing for ripening acceleration, and limited accumulation characteristic of lag metabolism. Water relations during lag phase shows minimal water uptake <0.05-0.1 g per berry per day, possible water loss through transpiration in some conditions, water balance approaching equilibrium, and minimal net water flux characteristic of quiescence. Berry morphology during lag phase shows berry at stable size, green color maintained though chlorophyll beginning decline in some cultivars, texture firm though slightly softer than peak firmness, surface smooth, and appearance as mature-sized hard green berry. Seed maturation during lag phase shows seeds completing final maturation, seed coat fully lignified and hardened, embryo and endosperm maturation finalizing, seed color dark brown at completion, and seed maturity as lag phase prerequisite for progression. Seed developmental state shows seeds approaching or achieving full maturity during lag phase, viable embryos ready for germination if separated, mature seeds signaling developmental readiness for berry ripening, and seed completion as potential veraison trigger. Metabolic activity during lag phase shows respiratory rate low 1.0-1.5 μmol O₂ g⁻¹ h⁻¹ representing maintenance metabolism, metabolic activity sustained at reduced levels, no growth-related biosynthesis, maintenance and preparation metabolism, and low activity characteristic of quiescence. Gene expression during lag phase shows growth genes at basal non-functional levels, lag-associated genes expressed including stress response and secondary metabolism genes, ripening genes beginning low-level expression preparing for activation, and expression patterns characteristic of pre-ripening state. Metabolic preparation for ripening shows subtle metabolic changes activating ripening pathways, anthocyanin biosynthesis genes beginning expression in red cultivars, sugar transporter genes increasing in preparation for accelerated import, softening enzyme precursors accumulating, and metabolic machinery preparing for veraison transformation. Stress response during lag phase shows enhanced stress response gene expression, protective mechanisms against heat and water stress, accumulation of stress-protective compounds, and stress adaptation as lag phase characteristic. Secondary metabolism during lag phase shows phenolic biosynthesis continuing, flavonoid accumulation proceeding, tannin synthesis and modification, and secondary metabolism as active process. Hormone dynamics during lag phase shows auxin at low levels, gibberellin possibly declining, abscisic acid remaining low before veraison spike, ethylene low in most cultivars, and hormone levels characteristic of quiescent state. Environmental effects on lag phase include temperature affecting lag duration with warm conditions shortening lag phase 1-2 weeks and cool conditions extending lag 3-4 weeks, water availability affecting berry hydration and composition, stress possibly triggering earlier veraison, and environmental factors determining lag duration and veraison timing. Temperature effects on duration show warm growing degree day accumulation accelerating progression through lag phase, cool conditions extending lag phase delaying veraison, and temperature as major lag duration determinant. Water stress effects show moderate deficit possibly accelerating veraison onset shortening lag, severe stress potentially arresting development, and stress as factor affecting progression. Lag phase duration variation shows cultivar genetic differences with early-ripening cultivars having shorter lag 1-2 weeks and late-ripening having longer lag 2-4 weeks, environmental conditions affecting duration within genetic ranges, and combined genetic-environmental determination. Lag phase significance in double sigmoid pattern shows lag representing growth plateau between phase I enlargement and phase II ripening growth, inflection point in development, and characteristic pattern distinguishing berry development. Lag phase completion shows eventual veraison onset with color change in red cultivars or softening in white cultivars, growth rate increasing from lag quiescence, sugar accumulation accelerating, and transition marking lag phase end. The process shows two distinguishable phases including lag phase initiation with growth rate decline start and metabolic transition onset, and minimal growth period with near-zero growth rate, seed development focus, and pre-veraison metabolism. Duration typically 1-4 weeks varying by cultivar and environment representing quiescent period weeks 8-12 post-fertilization. Regional timing shows berry growth lag phase during late August-September across climates between enlargement and veraison. Viticultural significance includes representing critical transition period between growth and ripening, determining veraison timing affecting harvest date, requiring different management than growth or ripening phases, and establishing metabolic foundation for successful ripening. Research applications include analyzing lag phase regulation and duration determinants, investigating metabolic transitions and preparations, evaluating environmental effects on lag duration and veraison timing, quantifying compositional and metabolic dynamics, and assessing lag phase as determinant of ripening progression and harvest timing.
Berry_Growth_Lag_Phase represents growth quiescence period between enlargement and veraison. Subdivided into Lag_Phase_Initiation with growth decline and metabolic transition onset, and Minimal_Growth_Period with near-zero growth, seed maturation focus, and pre-veraison metabolism. Duration 1-4 weeks (cultivar and environment dependent). Growth rate <5 mg/day. Berry dimensions stable. Seed maturation completing. Metabolic preparation for ripening. Represents plateau in double sigmoid growth pattern. Temperature major duration determinant. Follows Berry_Cell_Enlargement_Phase. Precedes Veraison.
Beeren-Wachstums-Lag-Phase
Lag-Periode
Prä-Veraison-Ruhephase
Wachstumsplateau
development pause
growth plateau
lag period
pre-veraison quiescence
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 453-460 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Berry Growth Lag Phase
Comprehensive assessment of Berry_Heat_Tolerance_Disposition at the Australian Wine Research Institute compared berry responses to controlled heating (berry surface temperature maintained at 45°C for 4 hours using infrared heaters) across 18 cultivars at véraison. Damage was assessed by visual scoring (sunburn symptoms), electrolyte leakage from berry skin, anthocyanin retention (red cultivars), and sugar-acid balance at harvest. Berry_Heat_Tolerance_Disposition varied substantially among cultivars. 'Grenache' demonstrated exceptional tolerance with minimal visible damage (score 0.5/5), low electrolyte leakage (18% relative increase), and anthocyanin retention of 85% following heat treatment. 'Pinot Noir' showed high sensitivity with severe sunburn (score 3.5/5), high leakage (52% increase), and anthocyanin loss to 45% of control, with affected berries failing to complete normal ripening. Among white cultivars, 'Verdelho' (tropical origin) exhibited strong Berry_Heat_Tolerance_Disposition while 'Chardonnay' was sensitive. Interestingly, berry heat tolerance did not correlate strongly with leaf heat tolerance (r = 0.38), suggesting partially independent mechanisms. Cluster architecture analysis revealed that cultivars with loose clusters and pendant orientation showed less heat damage under field conditions due to improved air circulation and reduced radiation interception, indicating that both intrinsic tissue tolerance and avoidance through architecture contribute to field-level Berry_Heat_Tolerance_Disposition.
A Berry_Heat_Tolerance_Disposition is a Heat_Tolerance_Disposition that inheres in Vitis vinifera grape berries and is realized through mechanisms that protect berry tissues from high temperature damage during ripening, preserving berry integrity, preventing premature abscission, and maintaining the biochemical composition essential for wine quality. This disposition represents heat tolerance of the economically critical fruit tissue, with berry heat damage manifested as sunburn (tissue necrosis from combined heat and light stress), heat-induced abscission (berry drop from pedicel failure), accelerated sugar accumulation with insufficient phenolic and aromatic development (decoupled ripening), anthocyanin degradation in red varieties, and altered acid metabolism (malate respiration). Berry temperature during hot periods can substantially exceed air temperature due to direct solar radiation absorption, with exposed berries reaching 45-50°C under full sun when air temperature is 35-40°C. Berry_Heat_Tolerance_Disposition encompasses both avoidance mechanisms (canopy architecture providing berry shading, berry skin properties reflecting radiation) and tolerance mechanisms (heat shock protein accumulation in berry tissues, antioxidant systems, cell wall stability). Cultivars differ in berry heat tolerance based on skin pigmentation (darker berries absorb more radiation), skin thickness and composition, cluster architecture affecting berry exposure, and biochemical heat protection capacity, with substantial breeding relevance as hot-climate viticulture expands.
Berry_Heat_Tolerance_Disposition is_a Heat_Tolerance_Disposition. Berry heat tolerance is distinct from vegetative tissue heat tolerance, with different damage mechanisms and consequences. Berry temperature can greatly exceed air temperature due to radiative heating. Child classes include Sunburn_Resistance_Disposition (combined heat/light damage) and Heat_Induced_Abscission_Resistance_Disposition (berry drop). Heat damage affects both yield (abscission, sunburn) and quality (anthocyanin degradation, decoupled ripening). Cluster architecture contributes to field heat exposure independently of intrinsic tissue tolerance.
Beeren-Hitzetoleranz-Disposition
Frucht-Thermotoleranz
Trauben-Hitzestress-Resistenz
berry thermotolerance
fruit heat tolerance
grape heat stress resistance
Greer DH, Weston C (2010) Heat stress affects flowering, berry growth, sugar accumulation and photosynthesis of Vitis vinifera, Functional Plant Biology 37:206-214 Spayd SE et al. (2002) Separation of sunlight and temperature effects on the composition of Vitis vinifera cv. Merlot berries, American Journal of Enology and Viticulture 53:171-182 Mori K et al. (2007) Loss of anthocyanins in red-wine grape under high temperature, Journal of Experimental Botany 58:1935-1945
Berry Heat Tolerance Disposition
Syrah in Rhône Valley shows berry initiation during early-mid July Year N+1 at BBCH 69-71, with auxin from 1.6-2.4 developing seeds triggering ovary expansion from 6 mm to 15-22 mm over 3 weeks, peak cell division producing berry flesh cells, pericarp differentiation into skin and flesh layers, developmental commitment establishing irreversible berry formation, under warm Mediterranean conditions of 24-30°C temperature, typical of this mid-to-late season cultivar completing initiation establishing berry developmental trajectory
Berry initiation represents the critical developmental transition phase following successful fertilization when auxin and other hormone signals from developing seeds trigger ovary expansion, stimulate cell division and growth in pericarp tissues, activate berry developmental programs, and establish irreversible commitment to berry formation, marking definitive shift from reproductive flowering to developmental fruit growth phase. This essential transition phase occurs during early fruit development in late June-early July in temperate regions Year N+1, proceeding over 2-4 weeks following fertilization as berry development begins, corresponding to BBCH 69-71 stages. Berry initiation encompasses coordinated processes including post-fertilization signaling with hormone production and transport from seeds to ovary, ovary expansion initiation with rapid growth beginning, early berry formation with tissue differentiation and structure establishment, and berry commitment reaching irreversible developmental state assured of maturation. Post-fertilization signaling shows auxin synthesis in developing embryo and endosperm beginning within days of fertilization, auxin export from seeds through seed coat and funiculus to ovary wall tissues, hormone accumulation in pericarp triggering growth responses, and additional signaling molecules including gibberellins and cytokinins possibly contributing to developmental activation. Auxin as primary signal shows indole-3-acetic acid as major form, hormone levels correlating with seed number and developmental activity, auxin perception in ovary wall cells triggering gene expression changes, and hormone as essential trigger for berry development initiation preventing flower abortion. Anti-abscission effect shows auxin blocking abscission zone formation at pedicel, hormone maintaining vascular connections supporting developing berry, and preventing programmed flower senescence pathway creating fruit retention. Growth-promoting effect shows auxin triggering cell division in ovary wall pericarp, stimulating cell expansion increasing tissue volume, activating biosynthetic pathways producing growth substrates, and overall hormone driving rapid berry enlargement. Ovary expansion characteristics show rapid growth from 4-8 mm pre-fertilization diameter to 10-20 mm within 2-3 weeks post-fertilization, volumetric increase 8-20 fold during first month, expansion rate peak during initiation period then declining, and visible berry formation confirming successful fruit set. Cell division during initiation shows peak mitotic activity in pericarp tissues, cycle times 10-15 hours during maximum division intensity, cell number increase determining berry flesh cellularity, division period lasting 3-4 weeks establishing final cell number affecting potential berry size, and transition to cell expansion phase following division period. Pericarp tissue organization shows differentiation into three layers including outer exocarp developing into berry skin, middle mesocarp forming bulk of berry flesh, and inner endocarp adjacent to seeds, with layer specialization proceeding during initiation period. Vascular development shows differentiation of vascular bundles in berry providing nutrient and water transport, xylem vessel formation enabling water import during growth, phloem development supporting sugar translocation during ripening, and vascular system establishment essential for berry development support. Early berry formation visible characteristics show green coloration from chlorophyll in skin and flesh, firm texture from active cell division and turgor, smooth appearance as surface cell division maintains coverage, and overall young berry appearance distinguishing from flowers or aborted structures. Berry developmental program activation shows transition from reproductive to developmental gene expression, upregulation of growth-promoting genes, activation of biosynthetic pathways for structural components, and establishment of developmental trajectory toward maturation. Metabolic activation shows increased respiratory rate supporting active growth, carbohydrate import and utilization providing energy and building blocks, nitrogen uptake and protein synthesis supporting tissue proliferation, and overall metabolic intensification characteristic of rapid growth phase. Berry commitment establishment shows developmental point beyond which berry development proceeds independently, commitment typically reached 2-3 weeks post-fertilization when expansion well established, berry capable of completing development to maturity barring severe stress or damage, and irreversible commitment distinguishing from pre-commitment flowers still capable of abortion. Commitment indicators include sustained berry expansion over multiple weeks, anatomical differentiation into recognizable berry structure, vascular development establishing permanent connections, and developmental gene expression programs fully activated. Resource allocation shift shows dramatic reallocation to developing berries consuming 40-60% total vine photosynthate, berries becoming dominant sink exceeding vegetative growth demands, resource competition between berries and shoots affecting growth balance, and berry sink strength increasing through initiation period. Sink strength characteristics show berry capacity to import and utilize resources, strength determined by cell number and metabolic activity, increasing sink strength establishing berries as priority allocation targets, and sink-source relationships affecting overall vine carbon balance. Berry number finalization shows fruit set percentage determined during initiation as abscission of unfertilized flowers completes, final berry count per cluster established by initiation completion, berry number per vine calculated from cluster counts and per-cluster berries, and yield potential determined by berry number and expected berry size. Environmental effects during initiation include temperature affecting expansion rate and cell division with optimal 20-28°C supporting rapid growth, water availability affecting cell expansion and turgor, nutrient supply particularly nitrogen supporting tissue synthesis, and overall favorable conditions during critical 2-4 week period supporting successful initiation. Temperature effects show warm conditions promoting rapid berry expansion and development, cool conditions slowing but generally not preventing initiation, and temperature as major factor affecting initiation rate and timing. Water relations show adequate water essential for cell expansion and growth, water deficit potentially limiting berry enlargement, and irrigation sometimes beneficial during initiation in water-limited conditions. Failed berry initiation causes include inadequate hormone stimulus from failed seed development, environmental stress during critical period disrupting growth, resource limitations preventing growth response, or physiological dysfunction though rare after successful fertilization. The process encompasses four distinguishable aspects including post-fertilization signaling establishing hormone communication, ovary expansion initiation beginning growth, early berry formation establishing structure, and berry commitment reaching irreversible state. Duration typically spans 2-4 weeks from fertilization through commitment. Regional timing shows berry initiation during late June-early July in warm climates, early-mid July in temperate regions, and mid-late July in cool areas. Viticultural significance includes representing critical transition from reproduction to development establishing berry formation, requiring successful hormonal signaling and favorable growing conditions, demonstrating irreversible developmental commitment determining yield, and marking beginning of berry growth phase dominating vine resource allocation. Research applications include analyzing hormone signaling and berry development triggers, investigating cell division and expansion dynamics, evaluating environmental effects on initiation success and timing, quantifying resource allocation shifts during initiation, and assessing berry initiation as determinant of development and quality outcomes.
Berry_Initiation represents critical transition phase within Fruit_Set establishing berry development. Subdivided into Post_Fertilization_Signaling with hormone communication, Ovary_Expansion_Initiation beginning growth, Early_Berry_Formation establishing structure, and Berry_Commitment reaching irreversible state. Duration 2-4 weeks post-fertilization. Auxin from seeds primary trigger. Peak cell division determining berry flesh cellularity. Marks definitive shift to developmental phase. Resource allocation shifts to berries as dominant sink. Commitment establishes assured maturation. Determines berry developmental potential.
Beeren-Initiierung
Beerenentwicklungs-Beginn
Entwicklungs-Commitment
Fruchtbildungs-Start
berry development initiation
fruit formation start
ovary to berry transition
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 415-425 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 315-322 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Berry Initiation
The berry length of Cornichon (syn. Olivette) documents extremely elongated berries measuring 30-40 mm from pedicel to stylar remnant — the longest berries in commercial V. vinifera cultivation, with a length-to-width ratio of 2.5-3.5 producing the distinctive finger-shaped or horn-shaped berry form (cornichon is French for gherkin, reflecting the berry's resemblance to a small cucumber) that makes this cultivar immediately identifiable among all grape varieties. This extreme berry length contrasts with the short berries of Pinot Noir (12-15 mm, length-to-width ratio approximately 1.0, yielding the near-spherical berry form) and the medium berries of Cabernet Sauvignon (12-14 mm, ratio approximately 1.05-1.10, yielding a slightly oblate spheroid).
A berry size description component that records the maximum length of the individual mature berry measured along the longest axis from the pedicel attachment (proximal pole) to the stylar remnant (distal pole), expressed in millimeters, providing the longitudinal dimension that together with berry width defines the overall berry proportions and enables the calculation of the length-to-width ratio used in berry shape classification.
Berry_Length documents the longitudinal dimension that is the primary axis used in berry shape classification — the ratio of berry length to berry width defines the fundamental geometric categories from oblate (wider than long, ratio 2.0). The measurement axis is defined from the proximal pole (the point of pedicel attachment, marked by the brush — the remnant vascular tissue at the berry-pedicel junction) to the distal pole (the stylar scar or navel, the remnant of the style from the original flower): in round berries these poles are easily identified but the measurement is nearly identical regardless of orientation, while in elongated berries the measurement axis is unambiguous but must be carefully aligned with the true longitudinal axis rather than any asymmetric curvature that some cultivars exhibit. Berry length is genetically determined with moderate environmental modification — water deficit during Phase III can reduce berry length by 10-20% compared to fully irrigated conditions, but the cultivar-specific proportional shape (the length-to-width ratio) is maintained because both dimensions are proportionally affected by water status.
Beeren-Längenmessung
Beerenlänge
Frucht-Achsenlänge
berry axis length
berry longitudinal dimension
fruit length measurement
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia
Berry Length
Berry number per cluster assessment comparing flowering conditions in Sauvignon Blanc reveals the weather effect on fruit set outcomes, with favorable flowering (warm, dry, 22°C average) producing 125 berries per cluster (42% fruit set from 300 flowers/cluster), marginal conditions (cool, humid, 16°C average) producing 85 berries per cluster (28% fruit set), and poor conditions (cold, wet, 12°C with rain during bloom) producing only 55 berries per cluster (18% fruit set), with the 56% reduction in berry number under poor conditions directly reducing yield potential and creating loose clusters with altered wine production characteristics.
Berry number per cluster quality is a quantity yield quality that represents the count of berries within individual clusters, characterizing the fruit set outcome that determines cluster architecture and contributes to cluster weight, with berry number established at fruit set when successful pollination and fertilization convert flowers into developing berries while unfertilized flowers abscise. Within the BFO framework, this quality specifically addresses the berry count dimension within clusters, distinguishing the number of berries from their individual size while recognizing that berry number is fixed at fruit set and remains constant through development (barring berry drop from disease or severe stress), with this early determination meaning that conditions during flowering have lasting effects on cluster composition. The ontological significance lies in berry number per cluster being determined during the critical flowering period when weather conditions, vine carbohydrate status, and pollination success interact to establish how many flowers become berries, with fruit set percentages typically ranging from 25-50% in wine grapes (meaning 50-75% of flowers fail to develop into berries), and with this conversion efficiency substantially affecting cluster compactness and final yield. The factors affecting berry number include flower number per cluster (established during bud differentiation the previous year), fruit set percentage (determined by conditions during flowering), and berry retention through development (normally high but reduced by disease, severe stress, or physiological drop), with the multiplicative relationship berry number = flower number × fruit set percentage × retention rate determining final count. The cluster compactness implications of berry number interact with berry size—high berry numbers with large berries producing compact clusters prone to disease, while lower berry numbers or smaller berries produce loose clusters with better air circulation and disease resistance, creating quality implications beyond the direct yield effects of berry count. Understanding berry number per cluster quality enables assessment of fruit set success, prediction of cluster compactness, and diagnosis of factors limiting berry number that might be addressed through management of flowering conditions.
Berry_Number_Per_Cluster_Quality addresses the count of berries within clusters reflecting fruit set success. This quality is determined at flowering and affects both yield and cluster architecture. The class supports fruit set assessment and compactness prediction through berry number characterization.
Beeren pro Büschel
Beerenpopulation
Beerenzahl pro Traube
Beerenzählung
berries per bunch
berry count
berry population
fruit set number
May P (2004) Flowering and Fruitset in Grapevines. Lythrum Press, Adelaide, pp. 1-119 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 145-230
Berry Number Per Cluster Quality
Pinot Noir berry pedicel at véraison (BBCH 83) measures 4-6 mm length, 1.0-1.5 mm diameter, contains 2-3 vascular bundles transitioning from predominantly xylem flow (Phase I) to increasing phloem dominance (Phase III), shows partial lignification with tan coloration, and supports 0.8-1.2 g berry mass with attachment strength 1.5-2.0 N indicating immature abscission zone
A berry pedicel is a pedicel specifically emphasizing its functional role in berry attachment, support, and vascular supply during berry development and ripening, representing the critical structural and physiological connection between individual berries and the cluster rachis system. Berry pedicel development parallels berry development through distinct phases= early fruit set (pedicel elongation and strengthening, 0-2 weeks post-fertilization, BBCH 71-73), Phase I berry growth (pedicel vascular expansion supporting rapid berry cell division, 2-6 weeks post-fertilization, BBCH 73-77), Phase II lag phase (pedicel lignification and structural reinforcement, 6-8 weeks post-fertilization, BBCH 77-81), and Phase III ripening (pedicel maturation with progressive abscission zone development, 8-16 weeks post-fertilization, BBCH 81-89). Berry pedicel anatomy shows developmental specialization= proximal end (rachis attachment) with reinforced vascular connection and mechanical strength for cluster support, mid-region with longitudinal vascular bundles (typically 2-4 bundles containing xylem vessels 15-50 μm diameter and phloem sieve tubes) embedded in parenchymatous cortex, and distal end (berry attachment) with receptacle tissue transitioning to berry exocarp and specialized abscission zone. Berry pedicel vascular function varies by developmental phase= during Phase I berry growth (BBCH 73-77) xylem flow dominates providing water and minerals for rapid berry expansion through cell division and early cell expansion (xylem flow rates 0.1-0.5 mg water/berry/hour), during Phase II lag phase (BBCH 77-81) both xylem and phloem are active with reduced xylem flow and increasing phloem import, and during Phase III ripening (BBCH 81-89) phloem increasingly dominates with xylem flow declining or reversing (phloem sugar import 0.5-2.0 mg/berry/hour supporting rapid sugar accumulation). Berry pedicel mechanical properties include= tensile strength 5-15 MPa (varying with developmental stage and cultivar), flexural strength supporting berry weight without breaking under normal cluster configurations, and attachment strength at berry junction 0.5-2.5 N detachment force (lower values indicating mature abscission zones, higher values indicating strong attachment in immature berries). Berry pedicel characteristics influencing berry development include= pedicel vascular capacity (larger diameter vessels and more numerous bundles support larger berry sizes), pedicel length affecting berry positioning and light exposure within cluster, pedicel lignification timing influencing cluster compactness and architecture, and abscission zone development affecting berry retention, harvest ease, and post-harvest berry drop. Cultivar variation in berry pedicel characteristics includes= pedicel length (3-15 mm), pedicel diameter (0.8-2.5 mm), vascular bundle number (2-5 bundles), lignification timing (early versus late), and abscission zone development (strong versus weak zones affecting shattering tendency).
Berry_Pedicel emphasizes functional berry support and vascular supply and is distinguished from Pedicel (complete structure including all attachment contexts) and specific pedicel subtypes (Attached_Berry_Pedicel, Mature_Berry_Pedicel) by focus on developmental phases and physiological roles in berry nutrition and attachment. Critical for understanding berry size determination through vascular capacity, sugar accumulation patterns during ripening, cluster architecture development, and harvest characteristics.
Beerenverbindungsstiel
Einzelbeerenstiel
Fruchtträger
Fruit stalk
Individual berry stem
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Greenspan, M. D., Shackel, K. A., & Matthews, M. A. (1994). Developmental changes in the diurnal water budget of the grape berry exposed to water deficits. Plant, Cell & Environment, 17(7), 811-820.
Berry Pedicel
Berry ripening delay quality assessment for potassium deficiency in Napa Valley Merlot compared ripening trajectories between potassium-deficient and potassium-sufficient vineyard zones to quantify the fruit quality impacts of K limitation. Weekly fruit sampling from véraison through harvest tracked ripening parameters across the potassium gradient established by soil variation and historical fertilization. Berry ripening delay quality manifested as reduced sugar accumulation rate in deficient zones, with Brix increasing at 0.18°/day versus 0.25°/day in sufficient zones, resulting in harvest Brix of 23.5° versus 25.8° at equivalent calendar date representing a 2.3° deficit. Acidity parameters showed berry ripening delay quality with titratable acidity of 6.8 g/L versus 5.2 g/L at harvest and pH 3.45 versus 3.62, indicating that the delayed acid degradation paralleled the delayed sugar accumulation. Color development showed berry ripening delay quality with anthocyanin concentration at harvest of 0.85 mg/g in deficient zones versus 1.35 mg/g in sufficient zones, a 37% reduction affecting wine color potential. Within-cluster variation assessment revealed berry ripening delay quality with coefficient of variation for individual berry Brix of 18% in deficient clusters versus 8% in sufficient clusters, reflecting the uneven ripening that complicates harvest timing. The berry ripening delay quality assessment established that achieving target ripeness in the deficient zone required extending harvest by 12 days, during which time increased Botrytis pressure and dehydration risk offset some of the ripening benefit, demonstrating the practical harvest management challenges created by potassium-limited ripening.
Berry ripening delay quality in potassium deficiency is a potassium deficiency symptom quality that inheres in grapevine fruit and represents the impaired and asynchronous fruit maturation resulting from inadequate potassium availability for the enzyme-mediated processes of sugar accumulation, acid metabolism, and color development that constitute normal ripening, with potassium-deficient fruit showing delayed achievement of target Brix, elevated titratable acidity, uneven ripeness within and among clusters, and reduced anthocyanin accumulation in red cultivars. This quality captures the fruit quality dimension of potassium deficiency, recognizing that while foliar symptoms are diagnostically useful, the fruit quality impacts are often the most economically significant consequence of potassium limitation, directly affecting wine quality potential. The ontological significance of berry ripening delay quality in potassium deficiency lies in its direct relationship to wine quality outcomes, where delayed and uneven ripening compromises the simultaneous achievement of sugar, acid, and phenolic maturity that defines optimal harvest timing for quality wine production. The biological basis of potassium-limited ripening encompasses the role of potassium as activator of enzymes involved in sugar transport and accumulation in berries, the influence of potassium on organic acid metabolism particularly the respiration of malic acid during ripening, and the effects of potassium on anthocyanin synthesis and accumulation that determine color development in red varieties. The relationship between berry potassium content and ripening parameters shows potassium's central role, with fruit potassium concentration correlating positively with sugar accumulation rate and negatively with final titratable acidity, such that potassium deficiency produces the characteristic combination of lower Brix and higher acidity at any given point during ripening. The uneven ripening within clusters results from differential potassium supply to individual berries based on their vascular connection quality and position, with berries receiving inadequate potassium remaining greener and more acidic while better-supplied berries ripen normally, creating the within-cluster variation that complicates harvest timing decisions.
Berry_Ripening_Delay_Quality_(K) is_a Potassium_Deficiency_Symptom_Quality, representing the impaired fruit maturation from K-limited sugar accumulation, acid metabolism, and color development. This class addresses the economically critical fruit quality dimension of K deficiency. The Brix-TA-anthocyanin triad all affected by K status. Within-cluster variation creates harvest timing dilemmas. Extended hang time may create secondary problems (disease, dehydration). Wine quality directly impacted through composition effects on color, acidity, and balance.
Beerenreifeverzögerung bei K-Mangel
K-Stress-Reifeverzögerung
K-deficiency ripening impairment
K-stress ripening delay
Kalium-limitierte Reife
delayed fruit ripening
potassium-limited maturation
verzögerte Fruchtreifung
Mpelasoka BS et al. (2003) A review of potassium nutrition in grapevines with special emphasis on berry accumulation. Australian Journal of Grape and Wine Research 9:154-168 Rogiers SY et al. (2006) Mineral sinks within ripening grape berries (Vitis vinifera L.). Vitis 45:115-123 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345
Berry Ripening Delay Quality (K)
Berry sampling reliability quality evaluation in a 12-hectare Barossa Valley Shiraz block exhibiting substantial vigor variation assessed the sampling intensity required for confident harvest decisions. Initial 100-berry samples collected from single vineyard transects showed poor berry sampling reliability quality with between-sample Brix coefficient of variation of 8.2%, insufficient for detecting the 0.5°Brix differences relevant to harvest timing optimization. Implementation of stratified random sampling with 200-berry samples from each of three vigor zones improved berry sampling reliability quality to CV of 3.1%, enabling detection of meaningful compositional differences with 95% confidence. Further increase to 500 berries per zone achieved CV of 1.8% but with diminishing returns relative to collection effort. Spatial analysis revealed that 65% of sampling variance originated from between-vine differences within zones rather than between-zone differences, informing protocol optimization focusing on increased vine sampling rather than zone subdivision, ultimately establishing a 300-berry stratified protocol as the cost-effective standard achieving adequate berry sampling reliability quality for commercial harvest decisions.
Berry sampling reliability quality is a harvest timing indicators quality that inheres in a grapevine cultivar or clone within a specific vineyard context and represents the consistency and accuracy with which collected berry samples reflect the true compositional status of the target population, determined by the uniformity of ripening across the vineyard block, the effectiveness of sampling protocols for capturing representative material, and the stability of sample composition between collection and analysis that enables reliable inference from sample measurements to population characteristics. This quality fundamentally influences the confidence of harvest timing decisions, recognizing that indicator measurements are only as reliable as the samples from which they are derived, with sampling error potentially exceeding analytical error as the primary source of uncertainty in maturity assessment for heterogeneous vineyard populations. The ontological significance of berry sampling reliability quality lies in its role as the foundation for evidence-based harvest decisions, wherein reliable sampling enables confident timing determination while unreliable sampling introduces uncertainty that may lead to suboptimal harvest decisions regardless of indicator quality or analytical precision. The biological basis of berry sampling reliability quality encompasses the spatial variation in ripening status across the vineyard block determined by soil heterogeneity, microclimate gradients, and vine-to-vine variation the within-cluster and within-vine variation in berry composition reflecting flower timing, pollination success, and positional effects on development and the temporal stability of composition in harvested samples during transport and holding before analysis. Sampling protocols designed to maximize berry sampling reliability quality employ stratified random sampling across defined vineyard zones, collection of sufficient berry numbers to reduce sampling variance below target thresholds, consistent selection criteria for sampled berries to avoid systematic bias, and rapid sample processing to minimize post-harvest compositional change. Assessment of berry sampling reliability quality involves replicated sampling with variance analysis to quantify the precision achievable with specific protocols, validation of sample representativeness against whole-block harvest data, and optimization of sampling intensity balancing cost against precision requirements for specific decision contexts.
Berry_Sampling_Reliability_Quality is_a Harvest_Timing_Indicators_Quality, representing the accuracy and consistency of sample collection for maturity assessment. This class addresses sampling methodology rather than the indicators measured, recognizing that sample quality fundamentally limits indicator utility. The coefficient of variation provides quantitative metric for reliability assessment. Stratified sampling protocols addressing vineyard heterogeneity improve reliability over simple random approaches. Critical for meaningful maturity data supporting harvest decisions.
Beerenbeprobungs-Zuverlässigkeit
Beerenstichproben-Präzision
Probenahmegenauigkeit
Stichprobenrepräsentativität
berry collection accuracy
sample representativeness
sampling precision
vineyard sampling reliability
Iland P et al. (2011) Chemical Analysis of Grapes and Wine: Techniques and Concepts, 2nd Edition. Patrick Iland Wine Promotions, Adelaide Bramley RGV (2005) Understanding variability in winegrape production systems 2. Within vineyard variation in quality over several vintages. Australian Journal of Grape and Wine Research 11:33-42 Trought MCT, Bramley RGV (2011) Vineyard variability in Marlborough, New Zealand: characterising spatial and temporal changes in fruit composition and juice quality in the vineyard. Australian Journal of Grape and Wine Research 17:79-89
Berry Sampling Reliability Quality
The berry shape description of Sultanina (syn. Thompson Seedless) documents narrow ellipsoidal berries (OIV 223:4) with a length-to-width ratio of approximately 1.4-1.6 — the moderately elongated oval shape with slightly pointed distal pole is characteristic of this cultivar and contributes to its visual distinctiveness in table grape markets where the elongated berry form is preferred over round berries for premium presentation. This ellipsoidal shape results from preferential cell expansion along the longitudinal axis during Phase III, a growth pattern associated with the stenospermocarpic seedless condition in which the aborted seed traces provide reduced and asymmetric growth hormone stimulation compared to fully developed seeds. The narrow ellipsoidal Sultanina shape contrasts with the perfectly spherical berries of Chasselas (OIV 223:2, length-to-width ratio 1.0), the broad ellipsoidal berries of Merlot (OIV 223:3, ratio 1.1-1.2), and the extreme elongation of Cornichon (OIV 223:8, ratio 2.5-3.5).
A berry description component that records the overall geometric form of the individual mature berry according to OIV descriptor 223, classifying berries on a scale including oblate/flattened (state 1), round/spherical (state 2), broad ellipsoidal (state 3), narrow ellipsoidal (state 4), cylindrical (state 5), elongated/finger-shaped (state 8), ovoid (state 6), obovoid/inverted egg-shaped (state 7), and other forms, with berry shape determined by the directional patterns of cell division and expansion during the two growth phases and providing one of the most visually distinctive cultivar identification characters at the individual fruit level.
Berry_Shape_Description documents a character of high diagnostic value because berry shape is strongly genetically determined, readily observable without instruments, and varies substantially across the V. vinifera cultivar spectrum — from perfectly round through various degrees of elongation to the extreme finger-shaped forms. The genetic control of berry shape has been partially characterized through QTL mapping, identifying a major locus on chromosome 18 (co-localizing with the seedlessness locus) affecting berry elongation, with additional contributions from loci on chromosomes 7 and 17: the association between berry shape and seedlessness reflects the developmental linkage between seed-derived growth hormones and the directional patterning of pericarp cell expansion — fully seeded berries tend toward spherical or oblate forms (isotropic cell expansion driven by uniformly distributed seed-derived hormones), while seedless or stenospermocarpic berries tend toward elongated forms (anisotropic expansion with preferential longitudinal growth). Beyond the basic geometric classification, some cultivars exhibit distinctive shape features not captured by the standard OIV categories — Muscat Hamburg berries show a characteristic slight constriction near the equator, Cornichon berries curve slightly producing a horn-like form, and some table grape cultivars display a truncated distal pole (flat-bottomed berries) that is an active breeding target in Asian table grape markets.
Beeren-Geometrie
Beerenform-Beschreibung
Fruchtform-Klassifizierung
berry form classification
fruit shape record
grape berry geometry
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Doligez A et al. (2013) New stable QTLs for berry weight do not colocalize with QTLs for seed traits in cultivated grapevine, BMC Plant Biology 13: 217 UPOV (2011) Guidelines for the Conduct of Tests for Distinctness, Uniformity and Stability — Vine (Vitis L.), TG/50/9
Berry Shape Description
Berry shrivel quality assessment in McLaren Vale Grenache following a 3-day heat event (temperatures exceeding 44°C with hot northerly winds) documented the extent and consequences of heat-induced berry dehydration for wine production. Visual assessment classified berries into shrivel categories from 0 (fully turgid) through 3 (moderate wrinkling with 15-25% volume loss) to 5 (severely shriveled with >40% volume loss), with block-average shrivel scores ranging from 1.8 to 3.5 depending on canopy exposure and irrigation status. Berry weight analysis confirmed visual assessments, with severely affected berries averaging 0.95 g versus 1.65 g for unaffected fruit, representing 42% weight reduction predominantly attributable to water loss. Compositional analysis showed that berry shrivel quality produced concentration effects with Brix increasing from 24.5° to 28.0° in affected fruit, anthocyanins increasing from 1.4 to 1.9 mg/g on a per-berry basis but declining from 2.3 to 2.0 mg/g fresh weight basis indicating that pigment loss exceeded concentration benefit, and titratable acidity declining disproportionately from 5.8 to 4.2 g/L suggesting acid respiration accompanying the dehydration stress. Juice recovery at crush was reduced by 25% in shriveled fruit, with the concentrated must requiring dilution to achieve target alcohol levels, while sensory evaluation identified dried fruit and jammy characters in wines from severely shriveled fruit that departed from fresh varietal expression typical of Grenache, establishing that severe berry shrivel quality impacts wine style as well as yield regardless of concentration effects.
Berry shrivel quality is a heat impact on development quality that inheres in grape berries and represents the extent of heat-induced water loss from berries causing volume reduction, skin wrinkling, and concentration of solutes that distinguishes heat-affected fruit from normally ripening berries and affects both the amount of recoverable juice and the composition of that juice through concentration effects that may or may not represent quality improvement depending on the initial composition and the severity of water loss. This quality captures the physical response of berries to heat-induced dehydration, recognizing that elevated temperatures increase both transpiration from berry surfaces and the vapor pressure driving force for water loss, while potentially also disrupting the vascular connections that would normally supply water to maintain berry turgor, creating the characteristic shriveled appearance of heat-damaged fruit. The ontological significance of berry shrivel quality lies in its multiple effects on wine production, including reduced juice recovery per unit fruit weight that affects yield calculations and processing efficiency, concentration of sugars, acids, and phenolics that may increase intensity but also risk excessive alcohol and extract, and the potential for off-flavors associated with heat-damaged fruit that may persist regardless of concentration effects. The biological basis of berry shrivel encompasses the physics of berry water loss through the cuticle and any damaged surfaces, the disruption of phloem and xylem connections that normally maintain berry hydration, and the metabolic changes in berry tissues responding to desiccation stress including cell membrane alterations and enzyme activity changes that affect berry composition beyond simple concentration effects. The severity of berry shrivel depends on heat intensity and duration, berry developmental stage, cultivar characteristics including skin thickness and cuticular properties, and vine water status that affects both berry initial water content and the capacity for vascular resupply. The distinction between heat-induced shrivel and normal late-season concentration is important, with heat shrivel typically occurring abruptly during heat events and affecting substantial portions of clusters uniformly, while normal concentration develops gradually affecting individual berries and providing the opportunity for selective harvesting that is not available when entire clusters or blocks are heat-affected.
Berry_Shrivel_Quality is_a Heat_Impact_On_Development_Quality, representing the heat-induced water loss causing berry volume reduction and solute concentration. This class addresses the physical dehydration response to heat stress. Visual shrivel scoring (0-5) and weight loss percentage provide quantification. Concentration effects may increase some quality parameters (sugar, anthocyanins per berry) while degradation concurrent with shrivel may decrease others (anthocyanins per g FW). Reduced juice recovery affects yield calculations and processing. Dried/jammy fruit characters distinguish heat shrivel from desirable concentration. Distinguished from late-season normal concentration by abrupt onset and uniform cluster effects.
Beeren-Dehydratation
Beerenschrumpfung
Hitzeschrumpfung
berry dehydration
berry water loss
heat shrivel
heat-induced concentration
hitzeinduzierter Wasserverlust
Greer DH, Weedon MM (2013) The impact of high temperatures on Vitis vinifera cv. Semillon grapevine performance and berry ripening. Frontiers in Plant Science 4:491 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 453-507 Caravia L et al. (2016) Application of shade treatments during Shiraz berry ripening to reduce the impact of high temperature. Australian Journal of Grape and Wine Research 22:422-437
Berry Shrivel Quality
The berry size description of Muscat of Alexandria documents very large berries (OIV 220:9) with typical dimensions of 22-28 mm length, 18-24 mm width, and 5-10 g weight at table grape harvest maturity — these large berries result from the combination of high seed number (typically 2-4 well-developed seeds providing abundant growth hormone production), vigorous cell division during the 4-6 week Phase I growth period (producing 350,000-500,000 cells per berry), and sustained cell expansion during Phase III (individual cells reaching 200-400 μm diameter). This very large berry size places Muscat of Alexandria at the upper extreme of V. vinifera berry size variation, contrasting with the very small berries of Pinot Noir (OIV 220:3, 12-15 mm length, 11-13 mm width, 1.0-1.5 g) and the small berries of Cabernet Sauvignon (OIV 220:3, 12-14 mm length, 11-13 mm width, 1.0-1.5 g), illustrating the 5-8 fold range in berry weight across cultivars.
A berry description component that records the dimensional characteristics of the individual mature berry — length, width, and weight — scored according to OIV descriptor 220 (berry size) on a scale from very small (state 1) to very large (state 9), with berry size reflecting the cultivar-specific interaction of seed number, cell division during the pre-véraison growth phase (Phase I), and cell expansion during the post-véraison ripening phase (Phase III) that produces the characteristic berry dimensions for each cultivar under given growing conditions.
Berry_Size_Description documents a character of central importance for both cultivar identification and quality assessment — berry size directly determines the skin-to-pulp ratio that is the primary morphological factor governing wine style (small berries = high ratio = more color and tannin per unit of juice volume), and is a primary quality criterion for table grape market grades (larger berries commanding premium prices). Berry growth follows the characteristic double-sigmoid pattern — Phase I (rapid growth through cell division for 4-6 weeks post-bloom), Phase II (a lag phase of 2-4 weeks during which organic acid accumulation peaks), and Phase III (a second rapid growth phase through cell expansion beginning at véraison and continuing to harvest): the final berry size is determined by the number of cells produced during Phase I (genetically determined, varying 2-fold between small- and large-berried cultivars) and the volume achieved by each cell during Phase III expansion (environmentally modulated by water availability, temperature, and crop load). The seed number effect on berry size is mediated by hormone production — each developing seed produces auxin, gibberellin, and cytokinin that stimulate cell division (Phase I) and cell expansion (Phase III) in the surrounding pericarp tissue, with more seeds producing proportionally larger berries within a cultivar.
Beeren-Dimensionserfassung
Beerengröße-Beschreibung
Fruchtgröße-Charakterisierung
berry dimension record
fruit size characterization
grape size assessment
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Coombe BG and McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening, Australian Journal of Grape and Wine Research 6: 131-135 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Ojeda H et al. (2001) Influence of pre- and post-véraison water deficit on synthesis and concentration of skin phenolic compounds during berry growth of Vitis vinifera cv. Shiraz, American Journal of Enology and Viticulture 52: 261-267
Berry Size Description
Pinot Noir in Burgundy shows berry size establishment during early-mid August Year N+1 at BBCH 75-77, with cell division completed by week 5, cell expansion continuing but decelerating weeks 6-8, berry diameter reaching 9-11 mm representing 75-88% final mature size, fresh weight 3.2-5.8 g, growth rate declining to 8-18 mg per day approaching lag phase entry, under continental conditions of 22-28°C temperature, typical of this early-to-mid season cultivar establishing size determining yield potential
Berry size establishment represents the final sizing phase during initial berry growth when cell division completes, cell expansion continues but decelerates, berry achieves approximately final dimensions, and growth rate declines toward lag phase entry, establishing berry size that largely determines mature berry size barring severe stress, proceeding over weeks 5-8 post-fertilization as initial growth phase concludes. This size-determining phase occurs during mid-late summer in late July-August in temperate regions Year N+1, spanning approximately 3-4 weeks as growth slows, corresponding to BBCH 75-77 berry touch to beginning of bunch closure stages. Size establishment involves completion of cell division with no further cell proliferation after week 4-5, continued cell expansion though at declining rate, berry achieving 70-90% final fresh weight by establishment completion, growth rate progressive decline entering lag phase, and berry dimensions at establishment largely predicting mature size. Cell division cessation shows division completion by week 4-5 in most mesocarp cells, final cell number established determining cellular foundation, no further cell proliferation during subsequent development, and division completion marking transition to expansion-only growth. Cell expansion continuation shows cells continuing enlargement through water uptake and vacuole expansion, expansion rate declining from peak weeks 3-5 to slower rate weeks 6-8, progressive expansion deceleration as cells approach mature size, and expansion continuing into ripening though at very low rate. Berry size achievement shows diameter reaching 8-12 mm in small-berried cultivars or 14-18 mm in large-berried cultivars by establishment completion, size representing 70-90% final mature berry size, berry volume 2-8 mL depending on cultivar, and fresh weight 2-10 g establishing size baseline. Size predictability shows strong correlation between size at establishment and final mature size, typical r² 0.75-0.90 between week 8 size and harvest size indicating reliable prediction, early size largely determining final size barring major stress or shriveling, and establishment size enabling early yield estimation. Growth rate decline shows progressive deceleration from peak 50-200 mg per day during rapid expansion to 10-50 mg per day during size establishment, eventual very low growth <5 mg per day entering lag phase, and declining rate characteristic of transition from growth to lag phase. Lag phase approach shows berry growth rate declining to near-zero, berry entering quiescent period before veraison, lag phase duration typically 1-3 weeks depending on cultivar and conditions, and lag representing transition from growth to ripening preparation. Berry dimensions at establishment show cultivar-characteristic size ranges reflecting genetic programming and environmental effects, small-berried cultivars 6-10 mm diameter, medium-berried 10-14 mm, large-berried 14-18 mm, and size variation within cultivars 15-30% coefficient of variation reflecting seed number and environmental heterogeneity. Fresh weight at establishment shows typical ranges 2-4 g small-berried, 4-8 g medium-berried, 8-15 g large-berried cultivars, weight variation within clusters reflecting seed number effects and microenvironment, and weight as practical size measure. Berry composition at establishment shows water content 80-85% fresh weight, sugars low typically 2-5% fresh weight as hexoses, organic acids 1-2% dominated by malate and tartrate, pH relatively low 2.5-3.5, and overall hard acidic green berry characteristic. Skin characteristics show green chlorophyll-containing epidermis, firm texture from cell wall integrity and turgor, smooth surface from coordinated growth, and protective cuticle layer developing. Seeds at establishment show seeds approaching mature size developing in parallel with berry, typical 1-3 viable seeds per berry, seed weight 20-40 mg each at establishment, and seeds completing development shortly before or during ripening. Berry-vine relationship shows berries as strong sink consuming 40-60% photosynthate during growth, sink strength declining as growth slows, and resource allocation beginning shift toward other sinks as berry growth completes. Environmental effects during size establishment include temperature affecting expansion rate and lag phase timing, water availability affecting final expansion and size achievement, and overall conditions during weeks 6-8 determining final establishment size. Temperature effects show warm conditions accelerating establishment completing size determination earlier, cool conditions extending establishment period, and temperature affecting timing rather than final size potential within normal ranges. Water effects show adequate water enabling full expansion to genetic potential, water deficit during establishment potentially reducing final size 10-30% through limited expansion, deficit effects less severe than during earlier rapid expansion, and late season water management affecting final sizing. Nutrient effects show generally adequate nutrition during establishment with mineral deficiencies rare, potassium possibly affecting water relations and final sizing, and overall nutrition less critical than water. Cultivar variation shows genetic differences in final size potential, timing of size establishment relative to phenology, and lag phase characteristics. Size establishment as critical determinant shows early size determining berry and cluster weight, yield per vine calculated from berry size × berry number × cluster number, and establishment completing berry developmental foundation for ripening. The process represents third and final phase of initial berry growth establishing final size, following early cell division growth and rapid initial expansion. Duration typically 3-4 weeks as growth declines toward lag phase. Regional timing shows berry size establishment during late July-August in warm climates, August in temperate regions, and August-early September in cool areas. Viticultural significance includes establishing berry size determining yield and cluster architecture, enabling early yield prediction from established berry dimensions, demonstrating that early size largely determines mature size emphasizing importance of favorable early conditions, and completing berry structural development preparing for ripening transformation. Research applications include analyzing size determination mechanisms and predictability, investigating environmental effects on final size achievement, evaluating early size as mature size predictor, quantifying yield estimation from early berry measurements, and assessing size establishment as yield and quality determinant.
Berry_Size_Establishment is third and final phase of Initial_Berry_Growth, distinguished by growth deceleration and final sizing. Duration 3-4 weeks as expansion slows. Cell division completed. Expansion continues but declining rate. Achieves 70-90% final size. Strong correlation with mature size enabling yield prediction. Approaches lag phase entry. Follows Rapid_Initial_Expansion. Completes initial berry growth phase. Determines berry size baseline for ripening.
Beerengröße-Bestimmung
Beerengröße-Etablierung
Größen-Festlegung
berry dimension establishment
final sizing phase
finale Größenphase
pre-lag sizing
size determination completion
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 443-450 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Berry Size Establishment
Syrah in Rhône Valley shows berry size peak during early-mid August Year N+1 at BBCH 77-79, with berry achieving 94-97% final dimensions at 13.2-15.8 mm diameter and 11.4-16.2 g fresh weight representing multiplicative combination of 2.4 million cells × 128 micrometer average cell size, remaining 3-6% size increase during lag and ripening, under warm Mediterranean conditions of 25-30°C temperature with adequate irrigation, typical of this mid-to-late season cultivar reaching growth phase size peak
Berry size peak represents the maximum growth phase dimensions achievement during maximum berry enlargement when berry reaches 90-98% final mature size, typical diameters 22-40 mm and fresh weights 9-25 g depending on cultivar, berry achieves near-final dimensions from cell number and cell size multiplicative effects, and size peak marks culmination of growth phase before transition to lag phase minimal growth, occurring weeks 6-8 post-fertilization as enlargement concludes. This size culmination phase occurs during late summer in early-mid August in temperate regions Year N+1, representing berry dimension peak weeks 6-8, corresponding to BBCH 77-79 bunch closure to majority berries touching stages. Berry size peak mechanism involves multiplicative combination of final cell number 1-4 million cells and near-maximum cell size 75-190 micrometers determining berry volume, berry achieving 90-98% final mature dimensions at peak, genetic size potential largely realized through completed cell expansion, remaining 2-10% size increase during lag and ripening from minor expansion and ripening accumulation, and peak establishing near-final berry size. Berry dimensions at peak show diameter ranges 22-40 mm depending on cultivar with small-berried 6-9 mm, medium-berried 10-14 mm, large-berried 15-22 mm at growth phase peak, fresh weight ranges 9-25 g in large cultivars and 2-6 g in small cultivars, and dimensions representing 90-98% final mature size. Berry volume at peak shows volumetric peak from cell population volume summation, typical volumes 3-15 mL depending on cultivar, volume determined by cell number × average cell volume relationship, and volume representing genetic size potential achievement. Cell number-cell size interaction shows final berry size = cell number × cell size in multiplicative relationship, cultivar variation in both parameters creating size diversity, optimal size requiring both adequate cell number from division and expansion capacity, and interactive effects establishing berry size ranges. Berry size genetic potential shows cultivar characteristic size ranges genetically programmed, small-berried cultivars 6-10 mm final diameter from limited cell number and expansion, large-berried 15-22 mm from high cell number and extensive expansion, and genetic control as primary size determinant. Size achievement percentage shows typical 90-98% final size at peak, calculation comparing peak size to mature harvest size, high percentage indicating growth phase dominance in size determination, and remaining 2-10% from lag and ripening phases. Remaining size increase shows minor continued enlargement during lag phase from residual cell expansion, slight ripening phase increase from sugar and water accumulation, total remaining gain 2-10% adding 0.5-2 mm diameter, and growth phase as major 90-98% size determinant. Berry fresh weight at peak shows weight ranges reflecting cultivar and achieved size, typical 9-25 g large-berried, 5-12 g medium-berried, 2-6 g small-berried cultivars, and weight as practical size measure. Berry dry matter at peak shows dry matter content 12-18% fresh weight at peak, absolute dry matter 1.2-4.5 g per berry, dry matter from structural components, organic acids, limited sugars, and dry matter representing non-water berry mass. Berry composition at peak shows water 80-85% fresh weight, organic acids at peak or beginning decline 1.2-1.6%, sugars still low 6-10%, pH 2.8-3.0, and hard acidic green berry composition. Berry morphology at peak shows berry at maximum green phase size, smooth surface maintained, spherical to ovoid shape depending on cultivar, firm texture from turgor and cell walls, and mature green appearance. Berry appearance shows green chlorophyll color persisting, hard texture from acids and turgor, smooth skin, and overall unripe green berry morphology. Cluster architecture at peak shows berries tightly packed within clusters, bunch closure with berries touching neighbors, cluster at maximum compactness, and peak cluster density. Environmental effects on berry size peak include water availability during enlargement determining achieved peak size with deficit reducing peak 15-40%, temperature affecting enlargement kinetics and peak timing, and environmental factors operating through cell expansion effects. Water deficit cumulative effects show deficit during peak expansion having largest size impact, deficit during maximum enlargement reducing final 5-15%, cumulative deficit effects determining achieved peak size, and water as major environmental size determinant. Temperature effects on peak show warm conditions potentially resulting in earlier peak timing, cool conditions extending enlargement delaying peak, and temperature affecting timing more than achieved size within normal ranges. Irrigation effects on peak size show supplemental water during growth phase enabling genetic size potential achievement, irrigation particularly beneficial in water-limited regions, and irrigation as key practice for size optimization. Berry size variation within clusters shows variation from seed number effects with 1-3 seed berries showing size ranges, microenvironment variation affecting individual berry sizes, and within-cluster coefficient of variation typically 15-30%. Berry size predictability shows peak size strongly predicting final harvest size, correlation r² 0.85-0.95 between peak and harvest dimensions, peak size enabling yield estimation, and high predictability from growth phase size dominance. Berry size peak timing shows typical achievement weeks 6-8 post-fertilization, cultivar variation in timing with early cultivars peaking earlier, timing relatively consistent within cultivars, and peak marking transition to lag phase. Berry size measurement shows diameter measurement with calipers, fresh weight measurement with balance, volume measurement through displacement, and size quantification enabling growth analysis. The process represents second aspect of maximum berry enlargement emphasizing berry-level size culmination, following cell size maximum and preceding expansion rate decline. Duration represents peak size achievement weeks 6-8. Regional timing shows berry size peak during early-mid August across climates as growth phase concludes. Viticultural significance includes representing near-final berry size determining yield, enabling early yield prediction from peak dimensions, demonstrating genetic size potential achievement, and establishing berry size affecting quality through concentration or dilution effects. Research applications include analyzing berry size determinants and genetic controls, investigating cell number-expansion interactions, evaluating environmental effects on size achievement, quantifying size predictability and variation, and assessing berry size peak as yield and quality determinant.
Berry_Size_Peak is second aspect of Maximum_Berry_Enlargement, distinguished by berry-level size culmination. Represents 90-98% final dimensions achieved. Ranges 22-40 mm diameter, 9-25 g depending on cultivar. Determined by cell number × cell size. Genetic potential largely realized. Remaining 2-10% during lag and ripening. Duration weeks 6-8. Follows Cell_Size_Maximum. Precedes Expansion_Rate_Decline. Highly predictive of final harvest size (r²=0.85-0.95). Major yield determinant.
Beerengröße-Spitze
Größen-Kulmination
Wachstumsphase-Spitze
berry dimension maximum
growth phase peak
maximale Beerengröße
maximum berry size
size culmination
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 450-453 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Berry Size Peak
Cabernet Sauvignon berry skin at maturity shows thick-skinned characteristics= total thickness 110-145 μm (cuticle 4.5-6.5 μm, epidermis 24-32 μm, hypodermis 42-55 μm, outer mesocarp 40-58 μm), containing 3.2-4.5 mg/g fresh weight anthocyanins (predominantly malvidin-3-glucoside 48-56% creating purple-black color), 9.5-13 mg/g dry weight tannins (degree of polymerization 12-16), providing skin-to-pulp ratio 12-16% and contributing 85-95% of wine color and 45-60% of wine tannins during 10-21 day maceration typical for premium Cabernet Sauvignon production. Pinot Noir berry skin showing thin-skinned characteristics= total thickness 60-95 μm (cuticle 3-5 μm, epidermis 18-26 μm, hypodermis 22-38 μm, outer mesocarp 18-32 μm), containing 2.2-3.2 mg/g anthocyanins (higher proportion of delphinidin and cyanidin 3-glucosides creating ruby-red versus purple color), 6.5-9.5 mg/g tannins, providing skin-to-pulp ratio 6-9% and requiring gentle extraction techniques during 5-10 day maceration to avoid excessive tannin while achieving adequate color for elegant Pinot Noir style. Chardonnay white cultivar berry skin showing white wine characteristics= total thickness 65-105 μm lacking anthocyanins, containing 0.9-1.6 mg/g flavonols and carotenoids creating golden-yellow color, 4.5-7.5 mg/g tannins (present but typically not extracted in white winemaking), with minimal skin contact during white wine production (immediate pressing after harvest) avoiding phenolic extraction and preserving fresh aromatic character though some premium Chardonnay styles use 2-6 hour skin contact for enhanced body and aromatics. Syrah berry skin phenolic extraction kinetics during red wine maceration= day 1 showing 25% anthocyanin extraction, 12% tannin extraction from skins day 3 showing 65% anthocyanin, 35% tannin day 7 showing 88% anthocyanin, 62% tannin day 14 showing 94% anthocyanin, 78% tannin, demonstrating that anthocyanins extract more rapidly than tannins with extended maceration primarily increasing tannin rather than color extraction, allowing winemakers to control wine phenolic profile through maceration duration management. Merlot berry skin thickness sun exposure effects= sun-exposed clusters showing 105-138 μm skin thickness with 3.5-4.6 mg/g anthocyanins, 10.5-13.5 mg/g tannins, 1.6-2.3 mg/g flavonols shaded clusters showing 78-108 μm thickness with 2.6-3.6 mg/g anthocyanins, 7.5-10.2 mg/g tannins, 0.6-1.1 mg/g flavonols, demonstrating sunlight effects on skin development and phenolic accumulation with exposed berries developing thicker skins with enhanced phenolic content producing wines with 20-35% higher color intensity and fuller tannin structure.
A berry skin is the visible outer covering of the grape berry comprising the cuticle, epidermis, hypodermis, and outer mesocarp layers, corresponding to outer pericarp or exocarp in anatomical terminology, typically 40-180 μm total thickness at maturity depending on cultivar and environmental conditions, serving as integrated protective barrier and phenolic-rich tissue that determines berry appearance, provides mechanical and biochemical protection, and contributes fundamentally to wine color, tannin structure, and aromatic complexity through extraction during winemaking. Berry skin structure comprises from exterior to interior= (1) waxy cuticle surface (2-8 μm) creating bloom and hydrophobic barrier (2) epidermis (single cell layer 15-35 μm) providing structural integrity (3) hypodermis (1-3 cell layers 20-60 μm) containing anthocyanins in red/black cultivars determining berry color (4) outer mesocarp layers (2-6 cell layers 25-90 μm) providing transition to flesh, with total berry skin 40-180 μm representing 5-8% of pericarp volume but containing >90% of berry anthocyanins and 60-75% of pericarp tannins. Berry skin functional characteristics= (1) visible appearance determined by anthocyanin content and composition (red, purple, black in colored cultivars), waxy bloom from epicuticular wax crystalloids, and surface texture from epidermal cell arrangement (2) tactile properties (thick-skinned versus thin-skinned cultivars) determined by total skin thickness, cell wall properties, and layer number affecting perceived skin toughness during eating or processing (3) barrier properties preventing water loss (transpiration 0.8-2 g H₂O m⁻² h⁻¹ at 25°C depending on cuticle integrity), resisting pathogen penetration, and protecting against UV radiation (4) phenolic reservoir providing wine color through anthocyanin extraction (red wine production), wine tannin structure through skin tannin extraction, and antioxidant capacity through total phenolic content. Berry skin composition shows cultivar-specific and environmentally-responsive characteristics= anthocyanin content 0-6+ mg/g fresh weight (cultivar-dependent, zero in white cultivars, 1.5-3 mg/g in moderate red cultivars, 3-6+ mg/g in deeply colored cultivars), with composition affecting color hue (higher malvidin proportion creating purple hue, higher delphinidin/petunidin creating red hue) skin tannin content 8-15 mg/g dry weight showing higher polymerization (mean degree 10-16 subunits) and lower astringency than seed tannins (mean degree 4-10) flavonol content 0.5-2.5 mg/g higher in sun-exposed berries providing UV protection and color copigmentation aromatic precursors including glycosidically-bound volatiles releasing aromatic compounds during fermentation contributing to varietal wine character. Berry skin in winemaking= red wine production requiring maceration (contact between juice and skins) to extract anthocyanins for color and tannins for structure, with extraction kinetics depending on skin cell wall properties, maceration temperature and duration, and alcohol concentration white wine production typically minimizing skin contact to avoid excessive phenolic extraction and maintain fresh character, though some styles use limited skin contact for aromatic enhancement rosé production using brief skin contact (2-24 hours) extracting partial color and limited tannins. Berry skin thickness variation= thick-skinned cultivars (Cabernet Sauvignon, Syrah, Nebbiolo) with 95-160 μm skin providing high skin-to-pulp ratio, high phenolic extraction potential, and firm texture thin-skinned cultivars (Pinot Noir, Gamay, Grenache) with 45-90 μm skin providing lower phenolic extraction and softer texture skin thickness affecting wine style potential with thick skins favoring full-bodied tannic wines and thin skins favoring elegant lighter-bodied wines.
Berry_Skin is the viticultural/functional term for the outer berry layers and is essentially synonymous with Outer_Pericarp or broadly-defined Exocarp. The term berry skin emphasizes the visible, tactile, and compositional aspects relevant to viticulture and winemaking rather than strict anatomical terminology. Sub-classification into Thick, Thin, and Pigmented berry skin reflects practical distinctions important for cultivar characterization and wine production. Understanding berry skin properties is fundamental for viticulture (managing phenolic development), harvest decisions (assessing phenolic maturity), and winemaking (extraction protocol design).
Beerenhaut
Beerenschale
Traubenhaut
berry epidermis
grape skin
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 176-185 Cadot, Y. et al. (2006). Tannin composition in grape skins. Journal of Agricultural and Food Chemistry 54=2973-2987 Downey, M.O. et al. (2006). Cultural practice and environmental impacts on flavonoid composition. American Journal of Enology and Viticulture 57=64-75.
Berry Skin
The berry skin description of Tannat documents very thick, tough skin (OIV 228:9) measuring 350-450 μm in cross-section (compared to 150-250 μm for thin-skinned cultivars like Pinot Noir), with a dense, multilayered hypodermis of 8-12 cell layers containing concentrated anthocyanin and tannin deposits, covered by a heavy epicuticular wax bloom giving the berries a dusty, matte-blue appearance — this extremely thick skin is the morphological basis of Tannat's reputation as one of the most tannic wine grapes, with skin tannin concentrations of 5,000-8,000 mg/kg (compared to 2,000-4,000 mg/kg for Cabernet Sauvignon), producing wines of exceptional structural intensity that traditionally required decades of aging to soften but are now managed through micro-oxygenation and other modern tannin management techniques pioneered specifically for Tannat in Madiran (southwest France) and Uruguay.
A berry description component that documents the physical characteristics of the berry skin (exocarp) beyond color — recording thickness, texture, toughness, elasticity, and the presence and character of the epicuticular wax bloom — that determine the skin's mechanical properties, its role as a barrier against pathogen entry, its contribution to wine tannin and anthocyanin extraction during maceration, and the eating quality (crunchiness, chewiness, or toughness) relevant for table grape assessment.
Berry_Skin_Description addresses the physical properties of the tissue layer that is the primary interface between the grape berry and its environment — the skin serves simultaneously as a water-loss barrier (the cuticle and epicuticular wax prevent desiccation), a pathogen defense barrier (the intact skin resists fungal penetration), a UV radiation filter (flavonols and anthocyanins in the hypodermis absorb UV-B), and the repository of the phenolic compounds (anthocyanins, tannins, flavonols, hydroxycinnamic acids) that are the primary determinants of red wine color, structure, and aging potential. Skin thickness varies approximately 2-fold across V. vinifera cultivars (from 150 μm in thin-skinned cultivars to 450 μm in thick-skinned cultivars) and is determined primarily by the number of hypodermal cell layers (ranging from 4-5 in thin-skinned to 10-12 in thick-skinned cultivars) rather than by individual cell size: this cell-layer-number variation is genetically determined during the early berry development phase (Phase I) when periclinal cell divisions in the hypodermis establish the final number of cell layers, and is largely independent of the environmental factors that modulate berry size. The functional consequence of skin thickness for winemaking is profound — thick-skinned cultivars contain more anthocyanin and tannin per berry (distributed across more cell layers), extract these compounds more slowly during maceration (requiring longer skin contact), and produce wines with greater color intensity, tannin concentration, and aging potential than thin-skinned cultivars.
Beerenhaut-Beschreibung
Beerenschale-Textur-Dokumentation
Exokarp-Eigenschaften
berry skin texture record
exocarp physical characterization
skin mechanical properties
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Ortega-Regules A et al. (2006) A first approach towards the relationship between grape skin cell-wall composition and anthocyanin extractability, Analytica Chimica Acta 563: 26-32
Berry Skin Description
Chardonnay in Champagne shows berry softening during late August-September Year N+1 at BBCH 83-85, with initial softening showing 14-20% firmness decline from 2.8 to 2.3 N over 4-6 days at veraison onset, polygalacturonase activity increasing from 8 to 32 units, pectin methylesterase from 12 to 42 units, water-soluble pectin increasing from 22% to 45% total pectin, progressive softening continuing through ripening with eventual 52-64% total firmness decline to 1.2-1.6 N, pectin molecular weight declining from 180 kDa to 45 kDa, middle lamella degradation weakening cell adhesion, under cool maritime conditions of 16-22°C temperature, typical of this early-bursting white cultivar showing softening as primary veraison marker
Berry_softening represents the texture transformation process during veraison when berry firmness declines from cell wall structural modifications creating softer more pliable berry, proceeding through initial berry softening with first measurable firmness decline 10-25% from cell wall modification onset and turgor changes, and progressive softening with continued cell wall degradation and increasing pliability leading to 40-70% total firmness decline by full ripeness, serving as universal veraison marker occurring in all cultivars unlike color which is cultivar-dependent, and softening continuing through ripening phase reaching soft ripe texture. This texture transformation process occurs during late summer through autumn in late August-October in temperate regions Year N+1, spanning veraison onset through ripening representing major textural change. Softening mechanism involves cell wall modification particularly pectin degradation through polygalacturonase and pectin methylesterase activities, middle lamella degradation weakening cell-cell adhesion, cell wall structural changes reducing firmness, possible minor turgor pressure changes though less important than structural modifications, progressive texture change from hard firm to soft yielding, and integrated changes creating characteristic ripe berry texture. Firmness decline trajectory shows firmness decreasing from pre-veraison maximum 2.5-4.5 N penetrometer resistance to initial softening 2.0-3.5 N representing 10-25% decline at veraison onset, progressive decline during ripening to 1.0-2.5 N representing 40-70% total reduction, eventual soft texture at full ripeness, and firmness decline as quantitative softening measure. Penetrometer measurement shows force required to penetrate berry skin and flesh declining with softening, measurement quantifying texture objectively, typical measurement using 2-5 mm diameter probe, and penetrometer as standard firmness assessment. Compression testing shows force required to compress berry declining, compression resistance decreasing 30-60% from pre-veraison to ripe, and compression as alternative firmness measure. Subjective texture shows berry tactilely harder pre-veraison, becoming noticeably softer at veraison, progressively more yielding through ripening, eventual soft pliable texture at maturity, and subjective assessment as practical evaluation. Cell wall modification shows pectin-degrading enzyme activation at veraison onset, enzymes including polygalacturonase PG, pectin methylesterase PME, pectate lyase PL, expansins possibly contributing, enzyme activities increasing substantially during veraison and ripening, and enzymatic modification as primary softening mechanism. Polygalacturonase activity shows PG cleaving homogalacturonan pectin backbone, PG activity increasing 5-20 fold from pre-veraison to veraison-ripening, PG activity correlating strongly with firmness decline, and PG as major softening enzyme. Pectin methylesterase activity shows PME demethylating pectin creating substrate for PG, PME activity increasing 3-10 fold, PME role complex with initial cross-linking possible then degradation facilitation, and PME as pectin modification enzyme. Pectate lyase possible role shows PL cleaving pectin through β-elimination, PL possibly contributing to degradation, activity variable by cultivar, and PL as additional enzyme. Expansin possible role shows expansins possibly contributing to wall loosening, expansin expression increasing during ripening in some studies, role less clear than pectinases, and expansins as potential contributors. Pectin structural changes shows pectin molecular weight declining from enzyme degradation, high molecular weight pectin 100-300 kDa at pre-veraison declining to lower 20-100 kDa fragments during ripening, depolymerization reducing structural integrity, and pectin degradation as softening basis. Pectin solubilization shows water-soluble pectin fraction increasing from degradation, insoluble cell wall pectin declining, soluble pectin as degradation product marker, and solubilization weakening structure. Middle lamella degradation shows pectin-rich middle lamella between cells degrading, degradation weakening cell-cell adhesion, cells becoming less firmly attached, possible cell separation in some cases, and middle lamella as critical adhesion site. Cell-cell adhesion weakening shows reduced intercellular bonding from middle lamella loss, tissue disaggregation potential increasing, texture becoming less coherent, and adhesion loss contributing to softness. Cell wall other changes shows possible hemicellulose modifications though minor compared to pectin, cellulose framework remaining largely intact, overall wall structure weakening, and integrated wall changes creating softening. Turgor pressure possible changes shows turgor possibly declining slightly during ripening from altered water relations, turgor changes generally minor 10-20% compared to cell wall structural changes 50-80% contribution to softening, turgor role secondary to structural modifications, and turgor as minor softening factor. Water relations shows berry water content relatively stable or slightly increasing during ripening, water influx from sugar osmotic gradient, water distribution possibly changing, and water affecting turgor and texture. Softening as universal marker shows softening occurring in all grapevine cultivars at veraison, universal marker unlike color which is red cultivar-specific, softening defining veraison in white cultivars, and universality making softening critical veraison indicator. Softening timing shows initiation at veraison onset coinciding with or following first visible color in red cultivars, rapid initial softening over first days of veraison, continued progressive softening through ripening weeks, and timing coordinating with overall transformation. Softening rate shows rapid initial firmness decline 10-25% over first 3-7 days veraison, slower continued decline during ripening 1-3% per day, eventual plateau at soft ripe texture, and rate pattern characteristic of transformation. Cultivar variation shows firmness levels and decline rates varying among cultivars, some cultivars maintaining firmer texture longer, others softening more rapidly, final ripe firmness varying 1.0-2.5 N, and genetic variation affecting softening. Berry handling implications shows softer berries more susceptible to mechanical damage, softening affecting harvest timing and handling, disease susceptibility potentially increasing with softening from easier pathogen penetration, and softening as practical concern. Sensory implications shows texture as important quality attribute, consumer preference varying with application, table grapes requiring firmness while wine grapes soften fully, and texture affecting acceptability. Wine quality implications shows softening affecting crushing and pressing, soft berries releasing juice more readily, texture influencing extraction during fermentation, and softening affecting processing. Environmental effects on softening include temperature affecting softening rate with warm conditions promoting faster softening and cool conditions slowing, water availability affecting berry turgor and softening dynamics, and environmental factors influencing softening progression. Temperature effects shows warm conditions accelerating enzymatic activities and firmness decline, cool conditions slowing softening rate, and temperature as rate factor. The process encompasses two sequential softening phases including initial berry softening with first firmness decline from cell wall modification onset, and progressive softening with continued degradation and increasing pliability. Duration softening initiating at veraison over 3-7 days initial decline then continuing through ripening 4-8 weeks total. Regional timing shows berry softening during late August-October from veraison through harvest. Viticultural significance includes representing universal veraison marker defining transformation across cultivars, determining berry texture affecting quality and handling, demonstrating cell wall modification as primary ripening process, and influencing harvest timing and berry management. Research applications include analyzing cell wall modification enzymes and regulation, investigating pectin degradation mechanisms and dynamics, evaluating softening genetics and cultivar variation, quantifying firmness decline and texture changes, and assessing berry softening as quality determinant and veraison marker.
Berry_Softening represents texture transformation within Veraison. Universal veraison marker (all cultivars). Subdivided into Initial_Berry_Softening (first 10-25% decline, cell wall modification onset, turgor changes) and Progressive_Softening (continued degradation, 40-70% total decline). Mechanism: pectin degradation (PG, PME primary), middle lamella degradation, cell-cell adhesion weakening. Turgor changes minor. Firmness 2.5-4.5 → 2.0-3.5 N initial → 1.0-2.5 N ripe. Duration veraison through ripening. Temperature affects rate. Critical quality and handling determinant.
Beeren-Erweichung
Beeren-Textur-Änderung
Festigkeits-Abnahme
Textur-Transformation
berry texture change
firmness decline
softening process
texture transformation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 475-485 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Berry Softening
Berry temperature monitoring of Grenache in Châteauneuf-du-Pape during August ripening shows sun-exposed berries reaching 44°C at solar noon despite air temperature of 36°C, with thermal imaging revealing a 12°C temperature gradient across the cluster from exposed to shaded surfaces, prompting implementation of overhead shade cloth reducing maximum berry temperature to 38°C and preserving the 15% higher anthocyanin concentration observed in protected versus exposed fruit.
Berry temperature quality is a tissue temperature quality that inheres in grape berries and represents the thermal state of this reproductive structure characterized by high susceptibility to radiant heating due to the spherical geometry maximizing radiation interception per unit volume, limited transpirational cooling capacity due to the waxy cuticle and absence of functional stomata, and moderate thermal mass providing some buffering of rapid temperature changes. Within the BFO framework, this quality specifically characterizes the berry as the quality-determining organ whose temperature directly affects sugar accumulation rate, organic acid respiration, anthocyanin synthesis and stability, aroma compound development and volatilization, and cell wall softening dynamics during ripening. The ontological significance extends to berry temperature being the critical parameter for heat damage assessment, where temperatures exceeding 35°C accelerate malic acid respiration, temperatures above 40°C inhibit anthocyanin synthesis and promote degradation, and temperatures approaching 45°C cause irreversible cellular damage and berry death. Berry temperature typically exceeds air temperature during daytime due to radiant heating, with the elevation depending on solar radiation intensity, wind speed affecting convective heat dissipation, and berry color where dark-skinned varieties absorb more radiation than light-skinned varieties. The position within the canopy strongly influences berry temperature, with exposed clusters experiencing temperatures 5-15°C above shaded clusters at the same time, creating substantial thermal heterogeneity within single vines that affects ripening uniformity. Measurement approaches include fine-wire thermocouples inserted into the berry mesocarp, infrared thermometry of berry surfaces, and thermal imaging for spatial temperature distribution assessment across clusters and canopy positions.
Berry_Temperature_Quality specifically addresses the reproductive structure critical for wine and table grape quality, distinguishing from vegetative organ temperatures that affect vine physiology but less directly determine fruit composition. This quality supports heat stress management decisions and interpretation of vintage effects on wine character. Berry position within cluster and canopy creates substantial temperature heterogeneity that may exceed cultivar or site differences when comparing thermal exposure.
Beerentemperatur
Beerenwärmezustand
Fruchttemperatur
Traubentemperatur
berry thermal state
fruit temperature
grape temperature
reproductive structure temperature
Bergqvist J et al. (2001) Sunlight exposure and temperature effects on berry growth and composition of Cabernet Sauvignon and Grenache in the Central San Joaquin Valley of California. American Journal of Enology and Viticulture 52(1): 1-7 Tarara JM et al. (2008) Berry temperature and solar radiation alter acylation, proportion, and concentration of anthocyanin in Merlot grapes. American Journal of Enology and Viticulture 59(3): 235-247
Berry Temperature Quality
Deficit irrigation protocols for Shiraz in the Barossa Valley deliberately target berry volume reduction of 25-35% relative to fully irrigated controls by maintaining soil moisture depletion below 50% of plant-available water from véraison through harvest, producing berries of 0.9-1.2 mL final volume compared to 1.4-1.8 mL in irrigated treatments, resulting in wines with higher anthocyanin concentration (800-1200 mg/L vs. 400-600 mg/L), more intense color, and greater aging potential.
Berry_Volume_Quality represents the three-dimensional spatial extent of individual grape berries, constituting perhaps the single most consequential quality measurement for wine grape production because berry volume directly determines the dilution or concentration of quality-relevant compounds. Berry volume results from the cumulative effect of cell division during early development (Phase I, lasting 3-5 weeks post-flowering) and cell expansion during pre-véraison (Phase II, approximately 4-6 weeks) and post-véraison (Phase III, approximately 5-7 weeks) growth periods. Final berry volumes typically range from 0.5-3.0 mL depending on cultivar genetic potential and environmental modulation. Small-berried cultivars including Cabernet Sauvignon (0.8-1.4 mL) and Pinot Noir (0.9-1.3 mL) inherently produce wines with higher skin-to-juice ratios than large-berried cultivars like Trebbiano (2.0-2.8 mL). Within any cultivar, environmental factors profoundly modify berry volume: water deficit during cell expansion phases can reduce volume by 30-50%, while excessive irrigation promotes larger berries with diluted composition. Temperature affects cell division rates and expansion duration. Crop load influences berry volume through competition among clusters for photoassimilates. Berry volume measurement employs water displacement for individual berries or small samples, graduated cylinders for larger samples, or calculation from diameter measurements assuming ellipsoidal geometry. The high correlation between berry diameter and berry volume (r² typically greater than 0.95) enables non-destructive volume estimation from caliper measurements.
Berry_Volume_Quality is a subclass of Volume_Quality specifically addressing the fruit unit central to grape and wine production. Berry volume relates to but is not identical with berry weight (mass) density variations of 1.05-1.15 g/mL occur due to sugar accumulation during ripening. The volume of berry compartments (skin, pulp, seeds) can be determined separately to characterize compartment proportions that affect extractability. Berry volume measurements should specify developmental stage and sampling protocol (e.g., 100-berry samples from marked clusters) for reproducibility. Berry volume interacts with cluster architecture to determine cluster compactness and associated disease susceptibility.
Beerengröße
Beerenvolumen
Einzelbeerenvolumen
Traubenbeerenrauminhalt
berry size
berry volumetric measurement
grape berry spatial extent
individual berry volume
Ojeda H et al. (2002) Influence of Pre- and Postveraison Water Deficit on Synthesis and Concentration of Skin Phenolic Compounds during Berry Growth of Vitis vinifera cv. Shiraz. American Journal of Enology and Viticulture 53(4): 261-267 Roby G et al. (2004) Berry Size and Vine Water Deficits as Factors in Winegrape Composition: Anthocyanins and Tannins. Australian Journal of Grape and Wine Research 10(2): 100-107
Berry Volume Quality
The berry weight of Syrah documents medium berry weight of 1.5-2.0 g at wine harvest maturity (22-25°Brix) — this medium berry size, combined with Syrah's moderately thick, deeply pigmented skin, produces a skin-to-pulp ratio of approximately 0.12-0.16 that is intermediate between the high-ratio small-berried Cabernet Sauvignon (0.15-0.20) and the low-ratio large-berried Grenache (0.08-0.12), explaining Syrah's capacity to produce deeply colored, moderately tannic wines without the extreme structural intensity of Cabernet Sauvignon. The standard measurement protocol determines mean berry weight from a randomized 200-berry sample collected from 20 clusters across 10 representative vines, with berries selected from all positions within each cluster (proximal, medial, and distal) to capture the within-cluster weight variation that typically spans a 2-fold range (smallest to largest berry).
A berry size description component that records the mass of individual mature berries at harvest, expressed in grams and typically determined as the mean of a 100-berry sample, scored on a scale from very low (state 1, 8.0 g), providing the most integrative single measurement of berry size that directly determines the skin-to-pulp ratio, juice yield per berry, and the cultivar's suitability for wine, table grape, or raisin production.
Berry_Weight is the most commercially relevant berry size parameter because it directly determines two critical production metrics — the skin-to-pulp ratio (which governs phenolic extraction potential in red winemaking and is inversely proportional to berry weight for a given cultivar) and the juice yield per berry (which determines the volume of must produced per unit weight of harvested fruit). The 100-berry weight method (weighing 100 randomly selected berries and dividing by 100 to obtain mean berry weight) is the standard field measurement used worldwide in viticultural research and commercial harvest assessment, providing a rapid and reproducible size determination that correlates strongly with berry volume and can be used to track berry growth dynamics through the season: the within-cultivar variation in berry weight between seasons and sites is substantial (CV 15-25%), driven primarily by water availability during Phase III cell expansion — water deficit reduces berry weight by limiting cell turgor-driven expansion, producing smaller berries with higher skin-to-pulp ratio and more concentrated phenolic composition, which is the physiological basis of the widespread viticultural practice of regulated deficit irrigation for red wine quality enhancement. The distinction between wine grape berry weights (typically 0.8-3.0 g) and table grape berry weights (typically 4-12 g) reflects the divergent selection pressures during the separate domestication and breeding trajectories of wine and table grape cultivars.
Beerengewicht
Einzelbeeren-Masse
Fruchtgewicht-Messung
berry mass record
fruit weight measurement
individual berry weight
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Ojeda H et al. (2001) Influence of pre- and post-véraison water deficit on synthesis and concentration of skin phenolic compounds during berry growth of Vitis vinifera cv. Shiraz, American Journal of Enology and Viticulture 52: 261-267 Coombe BG and McCarthy MG (2000) Dynamics of grape berry growth, Australian Journal of Grape and Wine Research 6: 131-135
Berry Weight
Berry weight data for Shiraz from the Barossa Valley (2023 vintage) records mean berry weight of 0.95 g (n=200, SD ±0.18 g) from old-vine (90+ year), dry-grown material, approximately 30% below the irrigated regional mean of 1.35 g for the same cultivar, with the reduced berry size directly contributing to the enhanced anthocyanin concentration (2.4 mg/g vs. 1.6 mg/g irrigated) and tannin density that define old-vine Barossa Shiraz quality. Berry weight data for Riesling from the Rheingau (2024 vintage) documents mean berry weight of 1.52 g at 83° Oechsle, within the typical cultivar range (1.3-1.8 g), with berry weight evolution tracked weekly from véraison showing the characteristic double-sigmoid growth curve — rapid Phase I expansion (0.6 to 1.1 g, weeks 1-3 post-véraison), a lag phase (weeks 3-4), and resumed Phase II expansion through sugar-driven osmotic water influx (1.1 to 1.52 g, weeks 4-7).
Berry weight data is a berry composition data entity that records the mass of individual grape berries or mean berry mass from representative samples, determined by gravimetric measurement of fresh berries at specified phenological stages or at harvest, expressed in grams per berry (g/berry) or milligrams per berry (mg/berry), and serving as the primary indicator of berry size that integrates the cumulative effects of cell division (pre-véraison), cell expansion (post-véraison), water influx, and solute accumulation on the final physical dimensions of the fruit, with direct implications for skin-to-juice ratio, phenolic concentration, and yield component analysis.
Berry weight is the most widely measured physical berry parameter and the yield component with the most direct influence on wine composition, as the inverse relationship between berry size and wine quality is one of the best-established principles in viticultural science. The mechanistic basis is straightforward — smaller berries have a higher surface-area-to-volume ratio and consequently a higher skin-to-pulp ratio, meaning that the phenolic, aromatic, and flavor compounds concentrated in the skin are diluted into a proportionally smaller volume of juice, producing more concentrated must and wine. However, the berry size-quality relationship is more nuanced than a simple "smaller is better" paradigm — very small berries from severe water stress may exhibit excessive seed tannin contribution (seed-to-berry ratio increases disproportionately in very small berries) and incomplete sugar accumulation, while moderately sized berries from balanced vines can produce exceptional wines through optimal ripening dynamics. Berry weight is determined by two sequential developmental phases — pre-véraison cell division (establishing the number of cells that determines size potential) and post-véraison cell expansion driven by sugar accumulation and osmotic water influx. Water deficit primarily restricts berry weight through reduced cell expansion in the post-véraison phase, creating the well-documented relationship between deficit irrigation strategies and increased phenolic concentration. The coefficient of variation within a sample (typically 15-25% CV for berry weight) provides information about ripening uniformity — high CV indicates asynchronous development that may require selective harvest or extended maceration to accommodate compositional heterogeneity. Berry weight measurement should specify whether the pedicel (cap stem) is included or excluded, as this methodological detail affects values by 2-5%.
Beerenfrischgewicht
Beerenmasse
Einzelbeerengewicht
berry fresh weight
berry mass
berry size data
mean berry weight
mittleres Beerengewicht
Roby G et al. (2004) Berry size and vine water deficits as factors in winegrape composition. Australian Journal of Grape and Wine Research 10(2): 100-107 Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 265-310 Houel C et al. (2015) Genetic variability of berry size in the grapevine. Australian Journal of Grape and Wine Research 21(1): 36-47
Berry Weight Data
Berry weight assessment comparing water regimes in Cabernet Sauvignon reveals the irrigation effect on final berry size and composition, with full irrigation producing berries averaging 1.45 g (larger berries, lower skin-to-pulp ratio 1:8, anthocyanins 1.2 mg/g), regulated deficit producing 1.15 g (moderate size, ratio 1:6.5, anthocyanins 1.6 mg/g), and severe deficit producing 0.85 g (small berries, ratio 1:5, anthocyanins 2.1 mg/g), demonstrating how berry weight affects both yield (30% reduction) and concentration (75% anthocyanin increase) with implications for wine style achievable from different irrigation strategies.
Berry weight quality is a quantity yield quality that represents the mass of individual berries at harvest, characterizing the final size achieved through berry development that contributes to cluster weight and affects fruit composition through the surface-to-volume relationships determining skin contact proportions during winemaking. Within the BFO framework, this quality specifically addresses the individual berry mass dimension of yield, distinguishing the size of berries from their number or composition while recognizing that berry weight results from cell division (primarily pre-véraison) and cell expansion (both pre-véraison and during ripening), with final size affected by genetics, water availability, temperature, and the source-sink balance between photosynthetic capacity and fruit demand. The ontological significance lies in berry weight affecting both yield quantity (larger berries contributing more mass) and fruit quality (smaller berries often showing higher skin-to-pulp ratio, greater concentration, and enhanced color and phenolic extraction potential), creating the complex relationship where berry size has opposing effects on quantity and quality that must be balanced for optimal outcomes. The developmental determination of berry weight involves the biphasic growth pattern with Phase I (cell division and expansion from fruit set to lag phase), lag phase (metabolic transition with minimal growth), and Phase II (cell expansion and ripening compound accumulation from véraison to harvest), with water availability during Phase II having particular influence on final berry size through effects on cell turgor and expansion. The cultivar variation in typical berry weight ranges from small-berried varieties (0.8-1.2 g, such as Pinot Noir, Cabernet Sauvignon) through moderate (1.3-1.8 g, such as Chardonnay, Merlot) to large-berried varieties (2.5-6.0 g, such as table grapes, some Muscats), with this genetic variation establishing the framework within which environmental factors cause additional size variation. Understanding berry weight quality enables yield estimation from berry counts, assessment of fruit composition implications from berry size, and management decisions targeting appropriate berry size for production objectives.
Berry_Weight_Quality addresses the mass of individual berries affecting both yield quantity and fruit concentration. This quality responds to water status and source-sink balance during berry development. The class supports yield assessment and quality prediction through berry weight characterization.
Beerengewichtqualität
grape berry weight
individuelles Beerengewicht
Matthews MA, Nuzzo V (2007) Berry size and yield paradigms on grapes and wines quality. Acta Horticulturae 754: 423-435 Roby G et al. (2004) Berry size and vine water deficits as factors in winegrape composition: anthocyanins and tannins. Australian Journal of Grape and Wine Research 10(2): 100-107
Berry Weight Quality
The berry width of Italia (syn. Ideal) documents broad berries measuring 20-26 mm at the widest equatorial plane — combined with berry length of 26-34 mm, this produces a length-to-width ratio of approximately 1.3-1.5, reflecting the ovoid berry shape characteristic of this major table grape cultivar. The large berry width of Italia contributes to its visual appeal in table grape markets (large, fleshy berries presented in well-filled clusters are preferred by consumers) and to its substantial berry weight of 7-12 g. This width dimension contrasts with the narrow equatorial diameter of Cornichon (12-15 mm despite the 30-40 mm berry length, yielding the extreme elongation ratio) and the moderate width of Merlot (12-14 mm, proportional to the 13-16 mm length and yielding the slightly ellipsoidal shape typical of most wine grape cultivars).
A berry size description component that records the maximum width of the individual mature berry measured perpendicular to the longitudinal axis at the widest point (typically the equatorial plane), expressed in millimeters, providing the lateral dimension that together with berry length characterizes the berry proportions and determines the cross-sectional area visible when viewing the berry from the pedicel end.
Berry_Width captures the transverse dimension that is particularly important for two aspects of berry morphology — the berry shape classification (via the length-to-width ratio) and the skin-to-pulp ratio estimation (via the surface-to-volume relationship). For spherical berries, width equals length and the surface-to-volume ratio is determined solely by size (smaller spheres having higher ratios), but for non-spherical berries, the elongation increases the surface-to-volume ratio beyond what would be predicted from volume alone — an elongated berry with the same volume as a spherical berry has approximately 10-20% more skin surface area, potentially increasing anthocyanin and tannin extraction per unit of juice volume: this geometric effect is one reason why some elongated-berry cultivars (e.g., Syrah, with moderately ellipsoidal berries) produce deeply colored wines despite having medium-rather-than-small berry size. The width measurement is standardized at the equatorial maximum, which for most V. vinifera cultivars corresponds to the widest point located slightly proximal to the geometric center (toward the pedicel end), reflecting the slightly pyriform (pear-shaped) tendency common in grape berries where the proximal hemisphere is marginally wider than the distal hemisphere.
Beeren-Breitenmessung
Beeren-Äquatorialdurchmesser
Beerenbreite
berry equatorial diameter
berry lateral dimension
fruit width measurement
Berry Width
β-Damascenone precursor concentration ranges from 100-800 µg/L across cultivars, with Chardonnay, Semillon, and Sauvignon Blanc showing higher accumulation (400-800 µg/L) than red cultivars such as Cabernet Sauvignon and Merlot (200-400 µg/L). The glycosidic precursors include multiple conjugate forms (glucose, 6-O-α-L-arabinofuranosyl-β-D-glucoside) requiring acid hydrolysis for release. Wine β-damascenone concentrations of 1-10 µg/L are typical, vastly exceeding the remarkably low sensory threshold of 0.05 µg/L (50 ng/L), making β-damascenone sensorially active in virtually all wines. β-Damascenone acts as an aroma enhancer, increasing perception of fruity notes and masking green/herbaceous character even when below its own detection threshold. Sun exposure during ripening increases β-damascenone precursor accumulation by 30-80%, with extended hang time also promoting accumulation.
A norisoprenoid accumulation disposition that is the inherited capacity of a Vitis vinifera plant to accumulate glycosidic precursors of β-damascenone (1-(2,6,6-trimethyl-1-cyclohexa-1,3-dienyl)but-2-en-1-one) in grape berries, with this potent odorant contributing rose, apple, honey, and tropical fruit notes across virtually all wine styles due to its extremely low sensory threshold.
β-Damascenone may be the single most impactful aroma compound in wine due to its exceptionally low threshold and near-universal presence. The compound contributes directly to fruity/floral character and indirectly through enhancement/suppression interactions with other odorants. Unlike TDN which shows strong cultivar specificity, β-damascenone precursors accumulate across all V. vinifera cultivars, though at varying concentrations. The carotenoid precursors (primarily neoxanthin) undergo CCD-mediated cleavage followed by further enzymatic and chemical modifications to generate the damascenone skeleton.
Damascenon-Potential
Rosenketonnote
damascenone glycoside disposition
damascenone potential
rose-apple aroma precursor
β-Damascenon-Vorläufer
Pineau B et al. (2007) Contribution of grape C13-norisoprenoid compounds to wine aroma in young Bordeaux red wines. Journal of Agricultural and Food Chemistry 55:6426-6433 Sefton MA et al. (2011) Occurrence, sensory impact, formation, and fate of damascenone in grapes, wines, and other foods and beverages. Journal of Agricultural and Food Chemistry 59:9717-9746
Beta Damascenone Precursor Disposition
Beta-damascenone precursor quality in late-harvest Riesling from Rheingau reaches elevated levels through extended carotenoid degradation during prolonged ripening, with precursor concentrations supporting β-damascenone release of 8-15 μg/L in finished Spätlese and Auslese wines this high damascenone contributes to the exotic, honeyed fruit character that distinguishes late-harvest Riesling from crisper Kabinett styles, where earlier harvest limits carotenoid degradation and produces damascenone levels of 2-5 μg/L with correspondingly fresher, less exotic aromatic profile.
Beta_Damascenone_Precursor_Quality represents the chemical quality characterizing the concentration of precursor compounds for β-damascenone in grape berries, capturing the bound forms and intermediates that can generate this highly potent norisoprenoid contributing exotic fruit, stewed apple, and honey aromas to wine. β-Damascenone ((E)-1-(2,6,6-trimethyl-1,3-cyclohexadien-1-yl)-2-buten-1-one) arises from neoxanthin and other C40 carotenoids through enzymatic and chemical oxidative cleavage, producing intermediate compounds including grasshopper ketone and 3-hydroxy-β-damascone that can undergo further transformation to yield β-damascenone. In grape berries, the direct precursors for β-damascenone occur as glycosidic conjugates that can be hydrolyzed during fermentation and aging. The free β-damascenone concentration in grapes and young wines is typically low (1-5 μg/L), but increases during wine aging through precursor hydrolysis to reach 5-15 μg/L in aged wines. The sensory threshold of β-damascenone in wine matrices is approximately 50 ng/L, making typical wine concentrations 20-300 times threshold and ensuring significant aroma contribution. Despite this suprathreshold presence, β-damascenone's sensory impact is often described as indirect, enhancing perceived fruitiness through synergistic interactions with other aroma compounds rather than contributing readily identifiable β-damascenone-specific descriptors. This enhancement effect makes β-damascenone concentration correlate with overall wine fruitiness even when the compound itself is not consciously perceived. Viticultural factors affecting carotenoid content (sun exposure, ripening temperature, harvest timing) indirectly influence β-damascenone potential through effects on precursor formation.
Beta_Damascenone_Precursor_Quality emphasizes the precursor forms rather than free β-damascenone because the bound glycosidic pool better predicts potential aroma development during aging. The complex pathway from carotenoid to final β-damascenone involves multiple intermediates, making direct precursor quantification challenging total glycosidic norisoprenoid potential often serves as a practical proxy. The aroma enhancement effect of β-damascenone, improving perceived fruitiness even when not consciously detected, distinguishes its sensory role from simple threshold-based contribution.
Beta-Damascenon-Vorläufergehalt
Exotikfrucht-Aromavorläufer
damascenone precursor content
exotic fruit aroma precursor
β-Damascenon-Glykosidqualität
β-damascenone glycoside quality
Sefton MA et al. (2011) Occurrence, Sensory Impact, Formation, and Fate of Damascenone in Grapes, Wines, and Other Foods and Beverages. Journal of Agricultural and Food Chemistry 59(18): 9717-9746 Kotseridis Y et al. (1999) Quantitative Determination of β-Ionone in Red Wines and Grapes of Bordeaux Using a Stable Isotope Dilution Assay. Journal of Chromatography A 848(1-2): 317-325
Beta Damascenone Precursor Quality
Mechanistic investigation of Bicarbonate_Tolerance_Disposition at INRAE Bordeaux employed hydroponic culture with factorial combinations of iron supply (1, 10 μM Fe-EDDHA) and bicarbonate (0, 5, 10, 15 mM NaHCO₃) across 14 rootstocks to separate bicarbonate-specific effects from general pH effects (pH maintained at 7.5 throughout). Measurements included leaf chlorophyll, total and active (Fe²⁺) iron, root ferric reductase activity, and xylem sap iron concentration. Bicarbonate_Tolerance_Disposition showed strong genotypic variation independent of overall lime tolerance capacity. '41B' exhibited exceptional Bicarbonate_Tolerance_Disposition, maintaining 85% of control SPAD values at 10 mM HCO₃⁻, with ferric reductase retaining 68% activity, xylem sap Fe concentration declining only 25% (from 48 to 36 μM), and leaf active Fe proportion remaining at 42% of total Fe. 'Fercal' showed similar bicarbonate resistance (80% SPAD retention, 65% reductase retention). '110 Richter', despite reasonable overall lime tolerance, displayed moderate Bicarbonate_Tolerance_Disposition (65% SPAD, 45% reductase at 10 mM HCO₃⁻). '3309C' showed extreme bicarbonate sensitivity with SPAD declining to 35% of control, reductase inhibited to 22%, xylem Fe reduced 60%, and leaf active Fe proportion collapsing to only 12% of total (severe Fe inactivation). Dose-response analysis established IC50 values (HCO₃⁻ concentration causing 50% parameter reduction) ranging from 14.5 mM for '41B' reductase to 4.2 mM for '3309C', quantifying the 3.5-fold difference in Bicarbonate_Tolerance_Disposition. These findings demonstrate that bicarbonate specifically affects both Fe acquisition and utilization, with tolerant rootstocks showing resistance at both levels.
A Bicarbonate_Tolerance_Disposition is a Lime_Tolerance_Disposition that inheres in a Vitis plant and is realized through physiological mechanisms that maintain iron acquisition and metabolism despite the inhibitory effects of bicarbonate ions (HCO₃⁻) on iron uptake processes and internal iron utilization. This disposition represents tolerance to the specific and potent chlorosis-inducing factor in calcareous soils—bicarbonate—which exerts effects beyond simply raising pH. Bicarbonate concentrations in calcareous soil solution can reach 5-15 mM and cause chlorosis even when external pH is artificially controlled, indicating direct biochemical effects on iron nutrition. Bicarbonate inhibits iron acquisition through multiple mechanisms including direct inhibition of ferric reductase activity (HCO₃⁻ affects enzyme kinetics), neutralization of rhizosphere acidification (consuming protons extruded by roots), inhibition of iron translocation from roots to shoots, and impairment of iron reduction and utilization within leaves leading to Fe³⁺ accumulation in inactive forms. Bicarbonate_Tolerance_Disposition encompasses resistance to all these effects, with tolerant genotypes maintaining higher iron acquisition and utilization efficiency at equivalent bicarbonate concentrations. The "iron chlorosis paradox"—high total iron but low active iron in chlorotic leaves—reflects bicarbonate-induced Fe inactivation, and genotypes with strong Bicarbonate_Tolerance_Disposition show higher proportions of metabolically active Fe²⁺ in their leaf tissues.
Bicarbonate_Tolerance_Disposition is_a Lime_Tolerance_Disposition. Bicarbonate exerts specific effects beyond pH on iron nutrition, including direct reductase inhibition, acidification neutralization, and Fe inactivation in leaves. The "iron chlorosis paradox" (high total Fe, low active Fe) reflects bicarbonate-induced inactivation. Bicarbonate tolerance is partially independent of overall lime tolerance, providing distinct selection criteria. IC50 values quantify bicarbonate sensitivity of specific processes. Active lime (extractable CaCO₃) correlates with soil solution bicarbonate, explaining IPC predictive value.
Bikarbonat-Toleranz-Disposition
HCO₃⁻ tolerance
HCO₃⁻-Toleranz
Karbonat-Stressresistenz
bicarbonate stress resistance
carbonate tolerance
Mengel K et al. (1984) Effect of bicarbonate on iron nutrition of chlorosis-susceptible plants, Journal of Plant Nutrition 7:1467-1489 Nikolic M, Römheld V (2002) Does high bicarbonate supply to roots change availability of iron in the leaf apoplast?, Plant and Soil 241:67-74 Kosegarten H et al. (1999) Effect of bicarbonate on iron acquisition of sunflower and bean, Journal of Plant Nutrition 22:1147-1159
Bicarbonate Tolerance Disposition
Pinot Noir hermaphrodite flowers at BBCH 59-65 exhibit typical bicarpellate pistils with bilocular ovaries 0.8-1 mm diameter containing 4 ovules (2 per locule), unified styles 0.5-0.7 mm, and stigmas 0.4-0.5 mm diameter showing subtle bilobed appearance, with fruit set analysis showing that successful flowers (48-56% of total) typically produce berries with 2-3 seeds resulting from fertilization of 50-75% of the 4 available ovules in the bicarpellate structure. Chardonnay bicarpellate pistils in anatomical sections reveal characteristic bilocular organization with two roughly symmetric chambers (locules) separated by central septum, ovules attached to septum in axile placentation representing positions along the two fused carpel margins, and vascular tissues showing bilateral organization supplying the two locules. Thompson Seedless (stenospermocarpic cultivar) produces seedless berries from bicarpellate pistils through parthenocarpic development without fertilization or with fertilization followed by early seed abortion, demonstrating that bicarpellate structure is retained even when seed development fails, with seedless berries showing residual seed traces in positions corresponding to the two locules. Wild V. vinifera ssp. sylvestris bicarpellate pistils across dioecious populations show consistent two-carpel organization in both pistillate females (with functional bicarpellate pistils 2.3-2.6 mm) and staminate males (with vestigial bicarpellate pistils 0.3-0.6 mm), demonstrating that bicarpellate organization is maintained across sex types despite dramatic differences in pistil functionality.
A bicarpellate pistil is a pistil formed from exactly two carpels, representing the universal pistil organization across all normal Vitis species and cultivars, with two-carpel structure creating characteristic bilocular ovary organization (two ovary chambers), two-sided ovule distribution (ovules attached along two lines corresponding to the two carpel margins), and often bilaterally symmetric stigma morphology, reflecting the ancestral and highly conserved gynoecium architecture in the Vitaceae family. Bicarpellate pistils in Vitis invariably show carpel fusion creating a compound syncarpous structure rather than apocarpous organization, with the two carpels fusing early in development (BBCH 52-53) to create unified pistil tissues. The bicarpellate organization is evident through multiple structural features= (1) bilocular ovary with two distinct chambers separated by a central septum formed from the fused medial margins of the two carpels, visible in transverse cross-sections as two symmetric or asymmetric cavities (2) ovule number typically 2-4 total, distributed as 1-2 ovules per locule, attached to axile placentation on the central septum at positions corresponding to the two carpel margins (3) stigma often showing bilobed morphology with two slight lateral extensions or showing bilateral papillae distribution with enhanced papillae density along two opposite margins reflecting the two-carpel origin (4) developmental origin from two carpel primordia visible during early organogenesis (BBCH 51-52) before fusion creates unified structure. The bicarpellate number (two carpels) represents an extraordinarily stable character across Vitis diversity, with essentially all flowers in all wild and cultivated Vitis species showing two-carpel organization regardless of other traits like flower sex type, cultivar group, or geographic origin, suggesting strong developmental constraint or strong selection maintaining bicarpellate organization. This stability distinguishes Vitis from many plant families showing variation in carpel number among species or even among flowers within species. The functional and evolutionary significance of bicarpellate organization relates to multiple aspects of reproductive biology and fruit development= (1) two-carpel ovary size and ovule number (2-4 ovules) creates seed number potential matching pollination efficiency and resource availability in Vitis ecology, with 1-4 seeds per berry representing successful fertilization of 25-100% of available ovules (2) bilocular organization provides spatial compartmentalization potentially allowing independent developmental trajectories for ovules in the two locules, evident in common occurrence of berries with seeds distributed asymmetrically between original locules (3) two-carpel structure determines basic berry morphology and size, with ovary dimensions at anthesis (0.6-1.2 mm diameter) scaling predictably to mature berry size (6-25 mm diameter depending on cultivar and growing conditions) (4) bicarpellate organization may reflect optimal balance between seed number potential (more carpels would allow more seeds but require larger flowers and more resources) and developmental coordination (fewer carpels would limit reproductive output). The bicarpellate structure interacts with breeding systems in determining reproductive outcomes= hermaphrodite flowers with bicarpellate pistils achieve 40-60% fruit set through self-pollination, with successful flowers typically showing fertilization of 1.5-2.5 ovules producing berries with 2-4 seeds functionally female and pistillate flowers require cross-pollination of bicarpellate pistils achieving 25-45% fruit set with similar ovule fertilization rates parthenocarpic (seedless) cultivars develop berries from bicarpellate pistils without fertilization (stenospermocarpic types) or with fertilization followed by seed abortion (stenospermic types), producing seedless or seed-trace berries while retaining normal bicarpellate ovary structure.
Bicarpellate_Pistil specifies the two-carpel organization that characterizes all normal Vitis pistils and is essentially synonymous with normal pistil structure in grapevines. This class emphasizes the carpel number (two) as the defining organizational feature and overlaps substantially with Two_Carpel_Compound_Pistil though the latter explicitly emphasizes the compound (fused) nature. The bicarpellate organization is universal in Vitis and highly stable evolutionarily, distinguishing grapevines from many other plant families with variable or higher carpel numbers. Sub-classification into Symmetrical and Asymmetrical types reflects variation in relative development of the two carpels and their derivative structures (locules). Understanding bicarpellate organization is fundamental for interpreting all aspects of pistil anatomy, ovule number and distribution, seed number determination, and fruit development.
Zwei-Fruchtblatt-Pistill
Zwei-Karpell-Pistill
bikarpellater Stempel
dicarpellate gynoecium
two-carpel pistil
Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 92-95 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 186-189 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 99-104.
Bicarpellate Pistil
A bifurcated tendril at node 8 on a shoot of Vitis vinifera cv. Pinot Noir in Burgundy, France, showing branch point at 7.5 cm from shoot attachment, dividing into two arms of 9 cm and 10.5 cm length creating nearly symmetrical Y-shaped architecture, with branch angle of approximately 45 degrees between arms, both branches exhibiting active circumnutation movements in complementary planes that together explore a cone-shaped volume of approximately 380 cubic centimeters, the longer arm making contact with a vertical stake and initiating tight coiling while the shorter arm continues searching in the opposite spatial quadrant, providing immediate support through one attachment plus continued exploration for potential secondary attachment, measured at BBCH stage 73 (berries beginning to touch) in early July, representing the characteristic bifurcated pattern dominant in Pinot Noir where 90-95% of tendrils show two-branch structure with relatively symmetrical arm development.
A branched tendril exhibiting division into exactly two branches or arms in Vitis vinifera, characterized by presence of a single branch point where the main tendril axis divides into two separate elongated segments of comparable developmental potential, representing the most common branching pattern in cultivated grapevines with 60-90% of branched tendrils across most Vitis vinifera cultivars showing bifurcated rather than higher-order branching, and providing optimal balance between increased contact probability through dual searching arms and maintenance of sufficient vigor in each branch for effective coiling and load-bearing function. Bifurcated tendrils develop through establishment of two lateral meristems at a single position on the developing tendril primordium during early organogenesis, typically 7-14 days post-emergence, with subsequent suppression of the original apical meristem or equal competition between all three meristems resulting in two-branch architecture, followed by independent elongation of each branch creating arms of variable relative length depending on timing of meristem activation and resource availability during development. The branch point position varies from proximal (3-5 cm from shoot attachment) to distal (8-12 cm from attachment) creating different functional characteristics, with proximal branching producing widely separated arms that explore larger volumes but with shorter individual branches, while distal branching maintains longer unbranched basal sections with shorter branches exploring refined spatial zones near the tendril tip. Bifurcated architecture serves as a key ampelographic feature with cultivar-specific variation in branching frequency, branch point position, relative arm length, and arm angle providing diagnostic characters for variety identification.
Bifurcated tendrils are distinguished from unbranched tendrils by presence of the division point, from trifurcated tendrils by exactly two rather than three branches, and from multi-branched tendrils by two rather than four or more branches. The bifurcated pattern represents an evolutionarily optimized architecture balancing contact probability enhancement against resource distribution efficiency, with two branches providing substantial improvement in search volume coverage (approximately 1.8-2.2× greater than unbranched tendrils) while maintaining 50% resource allocation per branch allowing strong development. Symmetry of bifurcation (equal versus unequal arm lengths) serves as an additional diagnostic feature, with some cultivars showing predominantly symmetrical bifurcation while others tend toward asymmetric patterns. Branch angle variation (narrow 20-30° versus wide 60-90°) affects spatial exploration patterns and is under genetic control with environmental modification.
Y-förmige Ranke
Y-shaped tendril
dichotome Kletterorgan
dichotomous tendril
forked tendril
two-branched tendril
zweigabelige Ranke
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 74-89 This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 95-118.
Bifurcated Tendril
A bilateral cordon on Vitis vinifera cv. Merlot in Bordeaux, featuring left cordon extending 78cm and right cordon extending 82cm from central trunk, each cordon bearing 13 two-bud spurs distributed along upper surface at approximately 12cm spacing, total configuration supporting 26 fruiting positions producing estimated 48-52 shoots and 11-13kg crop distributed nearly equally between paired cordons (approximately 52% right, 48% left), trunk precisely centered on trellis wire at 85cm height, representing standard bilateral cordon architecture employed throughout Bordeaux and globally in commercial Merlot production, with paired cordon configuration providing balanced vine development, efficient trellis utilization, and reliable productivity.
A cordon configuration featuring two cordons extending in opposite directions from a central trunk, characterized by paired symmetrical or asymmetrical architecture with cordons typically extending left and right along horizontal trellis wires, balanced or deliberately unbalanced distribution of fruiting positions between the two cordons, and representation of the most common cordon architecture in modern viticulture providing efficient trellis utilization, balanced vine capacity distribution, and compatibility with mechanized operations. Bilateral cordons dominate commercial wine grape production worldwide, combining practical advantages of dual-arm distribution (vigor split between paired structures, production insurance if one cordon fails or declines, efficient use of available trellis space) with proven productivity and quality outcomes across diverse varieties, regions, and training system objectives from bulk production to ultra-premium wine applications.
Bilateral cordons represent default modern cordon configuration, with paired architecture providing numerous practical advantages over unilateral alternatives including balanced growth distribution (vigor divided between two structures rather than concentrated in one), redundancy and insurance (partial production maintained if one cordon damaged or declining), efficient trellis space utilization (cordons extending both directions occupy maximum wire length per vine), simplified mechanization (bilateral canopies present consistent architecture for equipment), and proven performance across virtually all wine grape cultivars and regions. Minor asymmetries between paired cordons (10-20% length or crop differences) are normal and functionally acceptable, while substantial asymmetries may indicate problems requiring attention (trunk disease affecting one side, positioning issues, inadequate training) or deliberate adaptation to constraints (row-end positions, off-center trunks).
Doppelkordon
bilaterale Kordon
double cordon
paired cordon system
two-sided cordon
zweiseitige Kordon
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Reynolds & Vanden Heuvel (2009) "Influence of grapevine training systems on vine growth and fruit composition= a review" American Journal of Enology and Viticulture 60=251-268 Intrieri et al. (2011) "Grapevine mechanization= Constraints and perspectives" Acta Horticulturae 978=471-478.
Bilateral Cordon
A Cabernet Sauvignon vine trained to bilateral cordon at 90 cm in Napa Valley bears a Bilateral_Cordon_Role with cordons extending 70 cm each direction, each cordon carrying 6 spurs at 15 cm intervals—the symmetrical structure enabling mechanical leaf removal, shoot positioning, and harvest operations. A Grüner Veltliner vine on bilateral cordon in Austria's Wachau serves in a Bilateral_Cordon_Role, with the two-arm structure distributing the variety's naturally high vigor across extended cordon length for improved balance. A Tempranillo vine on bilateral cordon in Rioja bears a Bilateral_Cordon_Role adapted from traditional head training, the conversion to bilateral cordon enabling mechanization while maintaining regional production characteristics.
A cordon-trained vine role that inheres in a grapevine with two cordons extending in opposite directions from the trunk, creating symmetrical horizontal structure typically along a single trellis wire. Bilateral cordon (also termed double cordon or T-cordon) is the most common cordon configuration worldwide, distributing crop load across two cordons for balanced vine structure and doubled fruiting capacity compared to unilateral systems at similar vine density. Standard bilateral cordon training positions cordons at 75-100 cm height, with each cordon extending 50-80 cm (depending on vine spacing) to meet or approach neighboring vines, creating the continuous fruiting zone characteristic of trellised viticulture. The symmetrical structure facilitates mechanical operations—harvesters and pruners can work uniformly along rows without accommodation for directional variation. Bilateral cordon establishment requires careful training during years 2-3 after planting, with cordons developed simultaneously to ensure balanced growth and avoid vigor imbalances between arms. Spur positioning on bilateral cordons typically maintains 8-15 spurs per vine (4-8 per cordon), with spur number adjusted based on variety, vine vigor, and yield targets.
Bilateral_Cordon_Role represents the standard cordon configuration optimizing the balance between vine structure complexity and fruiting capacity. The symmetry of bilateral training facilitates management—pruning follows identical patterns on both sides, mechanical operations proceed uniformly, and vine appearance supports aesthetic expectations for well-maintained vineyards. Bilateral cordon is particularly suited to spur-pruned varieties (Cabernet Sauvignon, Merlot, Syrah, Grenache) where permanent spur positions can be maintained on the cordons for decades. The system's dominance in modern viticulture reflects its compatibility with mechanization, though hand-labor operations remain efficient with the predictable structure. Bilateral cordon vine spacing typically ranges from 1.0-1.5 m within rows, with cordon length approximately half the vine spacing to allow cordons to approach but not overlap between adjacent vines.
T-Kordon
T-cordon
Zweiseitiger Kordon
beidarmiger Kordon
bilateral cordon
two-arm cordon
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Winkler AJ et al. (1974) General Viticulture, University of California Press Reynolds AG, Vanden Heuvel JE (2009) Training systems influence on vine growth, American Journal of Enology and Viticulture 60: 251-268
Bilateral Cordon Role
A bilateral permanent arm configuration on Vitis vinifera cv. Chardonnay, featuring left arm extending 82cm and right arm extending 78cm from central trunk positioned at row center, each arm bearing 13-14 two-bud spurs distributed along upper surface, total vine capacity of approximately 26-28 fruiting shoots supporting 4.5-5.5kg crop distributed relatively evenly between arms (left arm slightly larger supporting proportionally more crop), arms positioned at 95cm height along horizontal trellis wire in vertical shoot positioning system, enabling efficient mechanical hedging and harvesting through consistent bilateral canopy architecture, representing standard bilateral cordon configuration widely used in commercial wine grape production worldwide.
A permanent arm configuration featuring two arms extending in opposite directions from a central trunk or head, characterized by symmetrical or asymmetrical paired architecture with arms typically extending left and right (or forward and backward depending on row orientation), balanced or deliberately unbalanced distribution of fruiting positions between the two arms, and suitability for maximizing trellis utilization in standard vineyard row configurations where bilateral spread efficiently occupies available wire space. Bilateral permanent arms represent the most common cordon architecture in modern viticulture, providing efficient use of trellis infrastructure, balanced vine capacity distribution between paired arms, flexibility for adjusting individual arm lengths based on vine vigor or positioning constraints, and compatibility with mechanized operations requiring consistent bilateral canopy presentation.
Bilateral permanent arms dominate modern cordon viticulture due to practical advantages including efficient trellis utilization (arms extending both directions from central trunk occupy maximum wire length per vine), balanced growth distribution (vigor split between two arms rather than concentrated in single structure), and mechanization compatibility (bilateral canopies suit over-row equipment for hedging, harvesting, and spraying). The bilateral configuration allows flexibility in arm length adjustment, with arms sized individually based on trunk position relative to trellis (centered trunks support equal arms, off-center trunks require asymmetric arms), vine vigor (weak vines may develop one arm initially, adding second when vigor permits), and row-end constraints (terminal vines often showing unilateral or strongly asymmetric bilateral development).
Doppelschenkel
beidseitige Kordon
bilaterale Arme
paired arms
two-arm system
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrope Canopy Management" Winetitles, Adelaide Reynolds & Vanden Heuvel (2009) "Influence of grapevine training systems on vine growth and fruit composition= a review" American Journal of Enology and Viticulture 60=251-268 Intrieri et al. (2011) "Grapevine mechanization= Constraints and perspectives" Acta Horticulturae 978=471-478.
Bilateral Permanent Arm
Leaf bilateral symmetry in Cabernet Franc serves as a distinguishing character from its offspring Cabernet Sauvignon, with Cabernet Franc displaying more pronounced bilateral symmetry (asymmetry index typically below 0.05) and five well-defined, evenly matched lobes, while Cabernet Sauvignon leaves exhibit greater asymmetry (index 0.06-0.10) with more variable lobe development between the left and right blade halves, reflecting the complex genetic heritage combining Cabernet Franc with Sauvignon Blanc.
Bilateral_Symmetry_Quality represents the symmetry condition where a grapevine organ can be divided by a single plane into two halves that are approximate mirror images of each other. This symmetry type, also termed zygomorphy or mirror symmetry, characterizes organs with a single axis of symmetry and two distinguishable sides that correspond across the dividing plane. In grapevine, bilateral symmetry is most evident in leaves, where the midrib defines the plane of symmetry and the left and right blade halves ideally display corresponding form, venation pattern, and lobe development. The degree of bilateral symmetry in leaves varies among cultivars and serves as an ampelographic character, with some varieties exhibiting highly symmetric blades while others show consistent directional asymmetry. Bilateral symmetry in berries occurs when seed distribution creates a plane of mirror correspondence, typically evident in two-seeded berries where the seeds occupy opposite positions. Cluster bilateral symmetry is uncommon but may be observed in clusters with two pronounced, equally developed wings. Assessment of bilateral symmetry employs either qualitative visual evaluation comparing the two halves, or quantitative methods including measurement of corresponding landmarks on each side, calculation of asymmetry indices (difference between sides divided by their mean), or procrustes analysis of digitized outlines. The developmental basis of bilateral symmetry involves coordinated gene expression and growth regulation across the organ primordium, with the plane of symmetry typically established early in organ initiation.
Bilateral_Symmetry_Quality specifically addresses mirror-image correspondence across a single plane, distinguishing it from Radial_Symmetry_Quality which involves rotational correspondence. For leaves, the plane of bilateral symmetry passes through the midrib and petiole attachment for bilaterally symmetric berries, the plane passes through the two seeds and the polar axis. Perfect bilateral symmetry (asymmetry index of zero) is rarely achieved in biological organs the classification applies when the predominant pattern approximates mirror correspondence. PATO_0001324 (bilateral symmetry) provides the upper-ontology reference.
Spiegelsymmetrie
bilaterale Symmetrie
mirror symmetry
reflective symmetry
two-sided correspondence
zweiseitige Entsprechung
zygomorphe Qualität
zygomorphic quality
Chitwood DH et al. (2016) Morphometric Analysis of Passiflora Leaves: The Relationship between Landmarks of the Vasculature and Outline. Frontiers in Plant Science 7: 1479 Klingenberg CP (2015) Analyzing Fluctuating Asymmetry with Geometric Morphometrics: Concepts, Methods, and Applications. Symmetry 7(2): 843-934
Bilateral Symmetry Quality
Cabernet Sauvignon ovaries at BBCH 59-61 (flower buds swollen to beginning of flowering) exhibit characteristic bilocular organization visible in transverse sections as two oval chambers 0.4-0.6 mm diameter each separated by central septum 0.08-0.12 mm thick, with 4 ovules distributed as 2 per locule attached to septum surfaces in axile placentation, and with vascular bundles supplying both locules through septum and ovary wall tissues. Chardonnay ovary anatomical studies reveal bilocular structure with two locules of approximately equal volume in symmetric pistils (left and right locules varying <15% in cross-sectional area) or unequal volumes in asymmetric pistils (locules varying 25-45% in area), central septum composed of thin-walled parenchyma cells with vascular strands running through septum tissue, and ovules projecting into locule cavities from septum attachment points. Pinot Noir berry development shows persistence of bilocular organization from anthesis (BBCH 61, ovary 0.9-1.1 mm diameter with two distinct locules) through early fruit set (BBCH 73, young berry 3.5-4.5 mm diameter with two locules still clearly separated by septum) with gradual obscuring of locule boundaries during later berry expansion (BBCH 81-85, berries 8-12 mm with flesh tissue proliferating and locule walls less distinct), though final seed distribution in mature berries reflects original bilocular organization. Rare Syrah flowers (<0.08% frequency) exhibit unilocular ovaries with single chamber resulting from complete septum absence or degeneration, containing 2-3 ovules in the single cavity and producing berries with unusual internal morphology lacking the typical bilocular seed distribution patterns.
A bilocular ovary is an ovary containing two internal chambers (locules) separated by a central partition (septum), representing the universal internal organization in normal Vitis ovaries reflecting the bicarpellate origin of the pistil with the two locules corresponding to the cavities of the two fused carpels. Bilocular ovaries are characterized by= (1) two distinct chambers (locules) evident in transverse cross-sections as two separate spaces within the ovary, typically appearing as two rounded or oval cavities positioned side-by-side (2) central septum (partition wall) separating the two locules, formed from the fused medial (inner) margins of the two constituent carpels, typically running along the median plane of the ovary though sometimes positioned asymmetrically in asymmetric bicarpellate pistils (3) ovules attached to axile placentation on the septum surfaces facing into each locule, with ovules positioned at the junctions where carpel margins fused to form the septum (4) two locules of approximately equal size in symmetric bicarpellate pistils (locules varying 20% in volume) (5) vascular tissues organized to supply both locules, with vascular bundles in the ovary wall and septum providing nutrient and water transport to ovules in both chambers. Bilocular organization develops during early pistil ontogeny (BBCH 52-54) when the two carpel primordia fuse to create a unified ovary structure, with the fusion of carpel margins creating the central septum dividing the originally separate carpel cavities into two locules within the compound ovary. The bilocular condition is essentially universal in Vitis, with rare exceptions= occasional flowers (<0.1% frequency) show unilocular ovaries (single chamber) resulting from complete septum degeneration or failure of one carpel to develop, creating ovaries with only one functional locule extremely rare flowers show trilocular or multilocular ovaries (three or more chambers) resulting from supernumerary carpel development or abnormal septum formation, creating ovaries with more than two locules. The bilocular organization has functional significance for reproduction and fruit development= two-locule structure provides spatial separation of ovules, potentially allowing independent fertilization and seed development in the two chambers bilocular organization creates potential for differential seed set between locules, with berries commonly showing 1-4 seeds distributed variably between the two original ovary locules (both seeds in one locule, seeds split between locules, or asymmetric distribution) berry internal structure reflects bilocular ovary organization, with mature berries often showing residual evidence of two-locule organization through seed positioning patterns or through subtle internal tissue organization even when locule boundaries become obscured during flesh development. The septum structure varies somewhat among flowers= typical septa are complete partitions extending from ovary base to apex, fully separating the two locules throughout their length some flowers show septa that are slightly fenestrated or show small gaps, partially connecting the two locules rare flowers show incomplete septa that separate locules only partially, creating effectively intermediate between bilocular and unilocular conditions. Post-fertilization berry development involves dramatic ovary expansion while generally maintaining bilocular organization in early stages (BBCH 71-77), though locule boundaries may become less distinct as flesh tissue proliferates in later stages (BBCH 79-89), with final mature berries showing seed distribution patterns that reflect the original bilocular ovary organization even when locule walls are no longer clearly visible in mature flesh tissue.
Bilocular_Ovary represents the universal internal organization of normal Vitis ovaries and must be distinguished from unilocular ovaries (one chamber, rare abnormalities) and multilocular ovaries (>2 chambers, extremely rare abnormalities). The bilocular condition reflects the bicarpellate origin of Vitis pistils with the two locules corresponding to the two fused carpels. Sub-classification into Two_Chamber and Septate types emphasizes different aspects of bilocular organization. Understanding bilocular structure is essential for interpreting ovule distribution, seed number and positioning, and berry internal organization.
Zwei-Fächer-Fruchtknoten
bilokulares Ovar
two-chambered ovary
two-locule ovary
zweikammeriger Fruchtknoten
Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 93-95.
Bilocular Ovary
Biochemical limitation assessment comparing nitrogen-sufficient versus deficient Riesling reveals the nutrition effect on photosynthetic capacity, with sufficient vines showing Vcmax of 95 μmol/m²/s and Lb of 22% (capacity exceeds demand under normal conditions), while deficient vines display Vcmax of 48 μmol/m²/s and Lb of 45% (capacity now limits photosynthesis even at ambient CO₂), demonstrating how nitrogen deficiency shifts limitation from primarily diffusional to primarily biochemical, with implications for fertilization management targeting the shifted constraint.
Biochemical limitation quality is a photosynthetic limitation quality that represents the constraint on photosynthesis arising from the capacity of carbon fixation enzymes and electron transport components, characterizing the degree to which Rubisco carboxylation capacity (Vcmax), electron transport capacity (Jmax), or triose phosphate utilization limits photosynthetic rate when CO₂ supply is not limiting. Within the BFO framework, this quality specifically addresses the metabolic component of photosynthetic limitation, distinguishing biochemical limitation from diffusional limitations while recognizing that enzyme capacity and electron transport set the upper bound on photosynthesis that diffusional limitations reduce below but cannot exceed. The ontological significance lies in biochemical limitation representing the fundamental capacity of the photosynthetic apparatus determined by protein investment and enzyme kinetics, where Rubisco content (nitrogen-costly enzyme comprising up to 30% of leaf nitrogen), electron transport chain components, and Calvin cycle enzymes together determine the maximum photosynthetic rate achievable even with unlimited CO₂ supply. The physiological basis for biochemical limitation involves multiple potential constraints including Rubisco carboxylation capacity (Vcmax, limiting at low Ci), RuBP regeneration capacity (Jmax, limiting at high Ci through electron transport), and triose phosphate utilization (TPU, limiting when export of photosynthate is restricted), with the transition between Vcmax and Jmax limitation occurring at Ci values that depend on temperature and the relative investment in these capacities. The environmental and nutritional factors affecting biochemical limitation include nitrogen availability (strongly affecting Rubisco content), phosphorus status (affecting ATP regeneration), temperature (affecting enzyme kinetics), and light environment during development (affecting chloroplast protein composition), with nitrogen deficiency typically increasing biochemical limitation through reduced Rubisco investment. The measurement of biochemical limitation employs A-Ci curve fitting to determine Vcmax and Jmax, with biochemical limitation calculated as the constraint on photosynthesis remaining after removing stomatal and mesophyll limitations through elevated CO₂ exposure. Understanding biochemical limitation quality enables assessment of photosynthetic apparatus capacity, nitrogen use efficiency evaluation, and identification of nutritional or genetic targets for capacity improvement.
Biochemical_Limitation_Quality addresses the enzyme and electron transport capacity component of photosynthetic limitation. This quality represents the fundamental capacity determined by protein investment and is particularly sensitive to nitrogen nutrition. The class supports capacity assessment and nutritional optimization through biochemical limitation diagnosis.
Kapazitätslimitation
Lb-Wert
Rubisco limitation
biochemische Limitation
capacity limitation
metabolic limitation
metabolische Einschränkung
Sharkey TD et al. (2007) Fitting photosynthetic carbon dioxide response curves for C₃ leaves. Plant, Cell & Environment 30(9): 1035-1040 Farquhar GD et al. (1980) A biochemical model of photosynthetic CO₂ assimilation in leaves of C₃ species. Planta 149(1): 78-90
Biochemical Limitation Quality
Mourvèdre vines at Château de Beaucastel in Châteauneuf-du-Pape bearing a biodiversity support role sustain documented populations of 45+ arthropod species in the canopy zone including lacewings (Chrysoperla carnea), ladybirds (Coccinella septempunctata), and predatory bugs (Orius spp.), with bark crevices on old cordons hosting overwintering earwigs (Forficula auricularia) at densities of 3-8 per vine, and the mycorrhizal network associated with 60+ year old root systems supporting 25-35 species of arbuscular mycorrhizal fungi per soil sample.
An ecological role that inheres in a grapevine by virtue of the habitat, food resources, and ecological niches its above-ground and below-ground structures provide for non-target organisms including arthropods, birds, mammals, fungi, and soil microbiota within the vineyard ecosystem.
Biodiversity_Support_Role captures the grapevine's function as a structural and trophic component of vineyard biodiversity, independent of intentional management interventions such as cover cropping or hedgerow planting. The vine itself — its bark texture, canopy architecture, flower resources during bloom, fruit availability, leaf litter, and root zone microhabitats — constitutes habitat infrastructure for diverse organisms: this role is amplified in older, structurally complex vines and reduced in young, intensively managed plantings. Subclasses specify the principal taxonomic groups benefiting from vine-associated habitat — pollinators, beneficial insects, and birds.
Artenvielfalt-Förderungs-Rolle
Biodiversitäts-Rolle
biodiversity habitat role
biological diversity role
biologische Vielfalt-Rolle
wildlife support role
Paiola A et al. (2020) Exploring the potential of vineyards for biodiversity conservation, Conservation Biology 34: 1092-1104 Bruggisser OT et al. (2010) Arthropod diversity in vineyard ecosystems, Agriculture Ecosystems and Environment 137: 103-112 Winkler KJ et al. (2017) Biodiversity in vineyards, Agriculture Ecosystems and Environment 239: 44-57
Biodiversity Support Role
Pinot Noir vines at Domaine de la Romanée-Conti in Burgundy bearing a biodynamic production role receive horn manure preparation (500) at 60-100 g/ha stirred for one hour and applied in autumn, horn silica preparation (501) at 2-4 g/ha applied at dawn during the growing season, and compost preparations (502-507) integrated into biodynamic compost applied at 3-5 t/ha, with all vineyard operations timed according to the Maria Thun biodynamic sowing calendar distinguishing root, leaf, flower, and fruit days.
A production system role that inheres in a grapevine cultivated according to biodynamic agricultural principles derived from Rudolf Steiner's 1924 Agriculture Course, integrating organic practices with specific biodynamic preparations (500-508), astronomical planting calendars, and a holistic farm-organism concept that treats the vineyard as a self-sustaining ecological entity.
Biodynamic_Production_Role encompasses all biodynamic management approaches whether or not formally certified by Demeter International or Biodyvin. The role requires adherence to the full set of biodynamic preparations and principles beyond those mandated by organic certification alone: biodynamic standards are strictly more restrictive than organic standards, with lower permitted copper limits (typically 3 kg Cu/ha/year versus 4-6 kg under organic regulations) and mandatory use of the nine biodynamic preparations. Subclasses distinguish Demeter-certified biodynamic operations from non-certified biodynamic practitioners.
Steiner-method production role
biodynamic farming role
biodynamic viticulture role
biodynamische Anbau-Rolle
biodynamische Produktionsrolle
biologisch-dynamische Bewirtschaftungsrolle
Steiner R (1924) Geisteswissenschaftliche Grundlagen zum Gedeihen der Landwirtschaft (Agriculture Course) Demeter International Production Standards (2022) Joly N (2007) Biodynamic Wine Demystified, Board and Bench Publishing pp: 45-112
Biodynamic Production Role
The biological control protocol for a premium Merlot vineyard in Saint-Émilion integrates four biocontrol strategies — mating disruption for Lobesia botrana using Isonet L dispensers (500/ha, deployed at E-L 9) releasing 200 mg/dispenser of (E,Z)-7,9-dodecadienyl acetate over 180 days to disrupt male moth orientation and reduce larval infestation below 1% cluster damage, Trichoderma atroviride (Esquive WP, 4 g/L) applied to fresh pruning wounds within 24 hours of cutting to colonize wound surfaces and competitively exclude trunk disease pathogens (Eutypa lata, Botryosphaeriaceae), Bacillus subtilis strain QST 713 (Serenade ASO, 8 L/ha) applied at BBCH 73 and 77 for pre-véraison Botrytis suppression through competitive exclusion and lipopeptide antibiosis, and conservation biological control through maintenance of permanent cover crop inter-rows and hedgerow margins to support populations of predatory mites (Typhlodromus pyri) that suppress phytophagous mites below economic threshold levels without acaricide intervention.
A pest management description that documents the use of living organisms, their products, or naturally derived substances to suppress pest and disease populations in the vineyard, specifying the biocontrol agent identity, application method, release rate or concentration, target pest, mechanism of action (parasitism, predation, competition, antibiosis, or induced resistance), and integration with other pest management components.
Biological_Control_Protocol documents the fastest-growing sector of vineyard pest management, driven by consumer demand for reduced pesticide residues, regulatory restriction of synthetic active ingredients, organic and biodynamic certification requirements, and the development of commercially viable biocontrol products and strategies. Biological control in viticulture operates through three complementary approaches — augmentative biocontrol (release or application of mass-produced natural enemies or microbial agents), conservation biocontrol (habitat management to support naturally occurring beneficial organisms), and classical biocontrol (introduction of co-evolved natural enemies from the pest's region of origin): mating disruption for Lobesia botrana represents the most successful augmentative biocontrol technology in viticulture, with adoption exceeding 200,000 hectares in Europe and efficacy comparable to insecticide programs at equivalent or lower cost. Microbial biocontrol agents (Trichoderma, Bacillus, Aureobasidium) have gained significant market share for botrytis and trunk disease management, though their efficacy is generally lower and more variable than synthetic alternatives, requiring integration within broader IPM frameworks rather than standalone deployment.
Biocontrol-Spezifikation
Nützlingseinsatz-Dokumentation
biocontrol specification
biological pest control record
biologische Bekämpfungs-Protokoll
natural enemy release protocol
Pertot I et al. (2017) A critical review of plant protection tools for reducing pesticide use on grapevine, Crop Protection 97: 70-84 Ioriatti C et al. (2011) Mating disruption for control of grape berry moth in Italian vineyards, Journal of Chemical Ecology 37: 1364-1377 Compant S et al. (2013) A review on the plant microbiome: ecology, functions, and emerging trends in microbial application, Journal of Advanced Research 19: 29-37 Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests 2nd Ed, APS Press
Biological Control Protocol
The BEDD for the Margaret River wine region (Western Australia, 33.9°S) documents a value of approximately 1,450-1,600 BEDD — the moderate maritime climate with growing season mean temperature of 18.5-19.5°C, narrow diurnal temperature range of 7-9°C (typical of maritime-influenced climates), and very few days exceeding the 19°C mean temperature threshold produces BEDD values that are lower than the standard GDD base 10°C for the same period (approximately 1,700-1,900 GDD) because the Gladstones calculation penalizes the days with mean temperatures above 19°C that contribute fully to the standard GDD but with reduced biological effectiveness in the BEDD framework. The BEDD classification places Margaret River in the intermediate viticultural zone, correctly predicting its suitability for the cultivar range actually grown there — Cabernet Sauvignon and Chardonnay as the dominant cultivars, with Shiraz and Sauvignon Blanc as significant secondary cultivars — a prediction that the Winkler Region III classification (based on uncapped GDD) overstates by suggesting suitability for warmer-climate cultivars than actually perform well in the region.
A growing degree-day component that records the Biologically Effective Degree-Days (BEDD) index developed by Gladstones (1992, refined 2011) — a heat accumulation method that improves on the standard GDD and Huglin Index by incorporating an upper temperature cutoff (19°C daily mean, above which developmental efficiency decreases), a daylength adjustment, and a diurnal temperature range adjustment that penalizes excessive thermal amplitude — providing the most physiologically refined single-index characterization of the temperature-development relationship in grapevine by accounting for both the cessation of growth at low temperatures and the reduced developmental efficiency at excessively high temperatures.
Biologically_Effective_Degree_Days represents the most physiologically sophisticated of the single-index heat accumulation methods, incorporating three adjustments that address documented limitations of simpler indices. The upper temperature cap reflects the well-established observation that grapevine photosynthetic rate, cell division rate, and sugar transport efficiency all decline above approximately 30-35°C (daily maximum), with effective developmental progress ceasing during extreme heat events — the BEDD captures this by reducing the daily degree-day contribution when the mean temperature exceeds 19°C, effectively applying a trapezoidal rather than linear temperature-development function: this correction is particularly important for distinguishing between warm-maritime and hot-continental climates that may have similar standard GDD but differ substantially in the proportion of biologically effective heat (maritime climates maintaining temperatures within the optimal range versus continental climates frequently exceeding it). The diurnal temperature range adjustment penalizes wide thermal amplitudes (>13°C difference between Tmax and Tmin) on the physiological basis that large temperature fluctuations impose greater metabolic costs (thermal acclimation, repair of heat/cold damage) than stable temperatures, reducing the net developmental progress per degree-day — this adjustment differentiates continental from maritime climates at the same latitude and mean temperature, correctly predicting that the maritime climate (narrow amplitude, more BEDD) supports more efficient vine development than the continental climate (wide amplitude, fewer BEDD) despite identical mean temperatures.
Gladstones bioclimatic index
Gladstones-Bioklimaindex
adjusted degree-day calculation
biologisch effektive Gradtage
temperature-capped heat summation
temperaturkorrigierte Wärmesumme
Gladstones J (1992) Viticulture and Environment, Winetitles Gladstones J (2011) Wine, Terroir and Climate Change, Wakefield Press Jones GV et al. (2005) Climate change and global wine quality, Climatic Change 73: 319-343 Hall A and Jones GV (2010) Spatial analysis of climate in winegrape-growing regions in Australia, Australian Journal of Grape and Wine Research 16: 389-404
Biologically Effective Degree Days
The PIWI (Pilzwiderstandsfähige) cultivar 'Regent' possesses multiple Biotic_Stress_Resistance_Dispositions inherited from its complex pedigree including Vitis amurensis ancestry. Its Rpv3-mediated Downy_Mildew_Resistance_Disposition is realized upon contact with Plasmopara viticola zoospores through rapid hypersensitive cell death limiting pathogen colonization, reducing disease severity by 70-85% compared to susceptible Vitis vinifera cultivars under equivalent inoculum pressure. Simultaneously, its Ren3-associated Powdery_Mildew_Resistance_Disposition is realized through localized cell death and phytoalexin accumulation when Erysiphe necator attempts penetration, though this resistance may be partially overcome by adapted pathogen strains in regions with high selection pressure. These resistance dispositions enable 'Regent' cultivation with 50-70% reduction in fungicide applications compared to traditional cultivars, demonstrating how understanding biotic stress resistance dispositions guides sustainable viticulture strategies.
A Biotic_Stress_Resistance_Disposition is a Grapevine_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through defense processes when the bearer encounters living organisms that pose potential threats to plant health, including fungal, bacterial, and viral pathogens, as well as arthropod pests and parasitic nematodes. This disposition encompasses the full spectrum of genetically-encoded resistance mechanisms, from major gene-mediated qualitative resistance providing near-complete protection against specific pathogen races, to polygenic quantitative resistance conferring partial but broad-spectrum protection. The molecular basis of biotic stress resistance dispositions includes pattern recognition receptors detecting conserved pathogen-associated molecular patterns (PAMPs), intracellular nucleotide-binding leucine-rich repeat (NLR) receptors recognizing specific effector proteins, and downstream signaling cascades activating defense gene expression. Realization of these dispositions manifests through diverse processes including hypersensitive response, phytoalexin biosynthesis, cell wall reinforcement, and systemic acquired resistance. Biotic stress resistance dispositions are of paramount importance in viticulture due to the susceptibility of Vitis vinifera cultivars to numerous pathogens and pests, driving breeding programs to introgress resistance loci from wild Vitis species and motivating rootstock selection based on resistance profiles.
Biotic_Stress_Resistance_Disposition is_a Grapevine_Disposition. This class encompasses all resistance dispositions against living organisms, distinguished from Abiotic_Stress_Tolerance_Disposition which covers responses to non-living environmental stressors. Child classes are organized by stressor type (Pathogen_Resistance_Disposition, Pest_Resistance_Disposition) and by defense mechanism (Defense_Mechanism_Disposition). The term "resistance" rather than "tolerance" is used following plant pathology conventions where resistance implies limiting pathogen development while tolerance implies maintaining yield despite pathogen presence.
Biotische Stressresistenz-Disposition
Krankheitsresistenz-Disposition
Schädlingsresistenz-Potenzial
biotic defense disposition
disease resistance disposition
pest resistance potential
Jones JDG, Dangl JL (2006) The plant immune system, Nature 444:323-329 Töpfer R et al. (2011) New horizons for grapevine breeding, Fruit, Vegetable and Cereal Science and Biotechnology 5:79-100 Dry IB et al. (2010) Grapevine disease resistance genes, Genetics, Genomics and Breeding of Grapes, Science Publishers
Biotic Stress Resistance Disposition
Pinotage vines in the Stellenbosch region of South Africa bearing a bird habitat role support 35-50 avian species documented through systematic point counts, including insectivorous species such as Cape White-eye (Zosterops virens) consuming 200-400 lepidopteran larvae per individual per day, seed-dispersing species such as Red-winged Starling (Onychognathus morio), and raptors including Jackal Buzzard (Buteo rufofuscus) providing rodent control, with vine trellising structures serving as perching and hunting platforms and mature canopy providing nesting sites for 3-5 passerine species per hectare.
A biodiversity support role that inheres in a grapevine whose vineyard environment provides nesting, foraging, and sheltering habitat for avian species, contributing to bird conservation in agricultural landscapes while simultaneously generating complex interactions ranging from beneficial insectivory and seed dispersal to fruit predation requiring management.
Bird_Habitat_Role captures a complex ecological interaction where avian presence generates both ecosystem services (insect pest consumption, rodent control) and disservices (fruit damage, yield losses of 2-10% in unprotected vineyards). The net ecological value of bird presence is context-dependent and species-specific: insectivorous species provide measurable pest control services valued at $100-500/ha/year, while frugivorous species may cause losses requiring netting or deterrent investment. Vineyard structural complexity (trellising, cover crops, adjacent hedgerows, old vine canopies) is the primary determinant of avian diversity within the vineyard matrix.
Vogelhabitat-Rolle
Vogelschutz-Rolle
Weinberg-Vogellebensraum-Rolle
avian habitat role
bird conservation role
vineyard bird habitat role
Pithon JA et al. (2016) Bird diversity in vineyard landscapes, Agriculture Ecosystems and Environment 223: 108-119 Assandri G et al. (2017) Diversity in the monotony: bird assemblages in vineyards, Agriculture Ecosystems and Environment 243: 30-41 Kross SM et al. (2012) Avian ecosystem services and disservices in vineyards, PLoS ONE 7: e46609
Bird Habitat Role
Bitterness localization mapping in Amarone della Valpolicella using trained panel assessment with oral cavity diagrams reveals primary perception at the circumvallate papillae region (back of tongue) with secondary perception on the soft palate for normally extracted wines, contrasting with broadly distributed bitterness extending to anterior tongue and oral mucosa in wines with excessive seed extraction from extended maceration, providing a qualitative indicator of extraction balance complementing intensity measurement.
Bitterness localization quality is a bitterness quality that represents the spatial distribution of bitter taste perception within the oral cavity, characterizing whether bitterness is perceived predominantly in specific regions (back of tongue, soft palate) or distributed broadly across gustatory surfaces. Within the BFO framework, this quality specifically addresses the spatial dimension of bitter perception, distinguishing localization from intensity and temporal aspects while recognizing that the perceived location of bitterness affects its integration with other taste qualities and overall impression of the bitter sensation. The ontological significance lies in bitterness localization affecting perceived quality, where bitterness confined to the posterior tongue and soft palate (typical pattern) integrates differently with front-of-mouth sweetness and mid-tongue sourness compared to broadly distributed bitterness, influencing the temporal unfolding of taste as stimuli move through the oral cavity during evaluation. The physiological basis for bitterness localization reflects the distribution of bitter-sensitive taste papillae, with circumvallate papillae at the back of the tongue containing high density of bitter-responsive cells, creating the characteristic posterior localization of strong bitter tastes, though fungiform papillae distributed across the anterior tongue also respond to bitter compounds enabling broader detection. The localization pattern may vary with bitter compound type and concentration, with threshold-level bitterness often perceived primarily at the back while suprathreshold levels recruit additional responsive regions creating more diffuse perception. In wine evaluation, localization contributes to bitterness quality assessment, where discrete posterior bitterness integrates more harmoniously with wine structure than bitterness perceived across the entire mouth, which may indicate excessive extraction or quality defect. Understanding bitterness localization quality enables refined sensory description beyond intensity alone, training of panelists to attend to spatial aspects of bitter perception, and evaluation of how different phenolic compositions create different spatial taste patterns.
Bitterness_Localization_Quality addresses the spatial dimension of bitter perception, complementing Perceived_Bitterness_Intensity_Quality (magnitude) and Bitterness_Persistence_Quality (duration). This quality requires specific methodology for spatial assessment, often using oral cavity diagrams or verbal localization protocols. While the "tongue map" concept of discrete taste zones is discredited, differential sensitivity across oral regions does affect perceived localization of taste qualities.
Bitterkeitsverteilung
Bitterlokalisierung
Bitterlokation
bitter perception region
bitter taste location
bitterness distribution
bitterness spatial pattern
räumliche Bitterwahrnehmung
Collings VB (1974) Human taste response as a function of locus of stimulation on the tongue and soft palate. Perception & Psychophysics 16(1): 169-174 Delwiche JF et al. (2001) Re-examination of the tongue map in human taste. Chemical Senses 26(6): 716
Bitterness Localization Quality
Bitterness persistence evaluation comparing young Barolo (18 months) versus aged Barolo (7 years) using time-intensity methodology reveals persistence of 35±6 seconds in young wine declining to 18±4 seconds in aged wine despite similar peak bitterness intensity, with the 49% reduction in persistence reflecting tannin polymerization during aging that reduces the low-molecular-weight fraction responsible for prolonged bitter sensation, explaining the qualitative improvement in finish character with bottle aging.
Bitterness persistence quality is a bitterness quality that represents the duration of bitter taste perception following swallowing or expectoration, characterizing the lingering bitterness that continues after the primary taste stimulus has been removed and significantly affects the finish and aftertaste quality of wine evaluation. Within the BFO framework, this quality specifically addresses the temporal decay dimension of bitterness, distinguishing persistence from peak intensity and spatial localization while recognizing that persistent bitterness fundamentally shapes finish perception and overall quality judgment. The ontological significance lies in bitterness persistence often being more important than peak intensity for quality perception, as briefly perceived bitterness may add complexity while prolonged bitter aftertaste creates negative quality impression and palate fatigue that affects subsequent wine enjoyment and food pairing. The physiological basis for bitterness persistence involves slow receptor dissociation kinetics for certain bitter compounds, residual compound retention in oral mucosa, and potentially longer central processing times for bitter taste compared to sweet, creating the characteristically long duration of bitter perception. The compositional factors affecting bitterness persistence include the molecular size and hydrophobicity of bitter compounds, with larger polymeric tannins showing different persistence than monomeric catechins, though the relationships are not fully characterized. The interaction between bitterness persistence and astringency duration on the finish is particularly relevant for red wine evaluation, where these co-occurring sensations may show similar or different temporal patterns affecting the overall mouthfeel evolution. In wine quality assessment, short bitterness duration that transitions to clean finish is generally preferred over persistent bitterness that dominates the aftertaste, though some bitter persistence may contribute to perceived complexity in appropriate wine styles. Understanding bitterness persistence quality enables characterization of finish quality, prediction of wine-food pairing success, and winemaking optimization considering the temporal trajectory of bitter perception.
Bitterness_Persistence_Quality addresses the duration of bitter perception after stimulus removal, completing the bitter perception characterization with Perceived_Bitterness_Intensity_Quality and Bitterness_Localization_Quality. This quality is particularly important for wine finish evaluation where persistent bitterness is generally considered negative. The relationship between persistence and molecular characteristics of bitter compounds offers potential for predicting sensory behavior from chemical analysis.
Bitterdauer
Bitterkeits nachklang
Bitterkeitspersistenz
anhaltende Bitterkeit
bitter aftertaste
bitter finish duration
bitter persistence
lingering bitterness
Guinard JX et al. (1986) The time-course of astringency in wine upon repeated ingestion. American Journal of Enology and Viticulture 37(3): 184-189 Fontoin H et al. (2008) Effect of pH, ethanol and acidity on astringency and bitterness of grape seed tannin oligomers in model wine solution. Food Quality and Preference 19(3): 286-291
Bitterness Persistence Quality
Bitterness quality assessment in Cabernet Sauvignon wines from early versus late harvest reveals bitterness ratings of 5.8 versus 3.2 on a 10-point scale despite similar total phenolic content (1850 versus 1780 mg/L GAE), with the 45% higher bitterness in early-harvest wine attributed to higher proportion of monomeric catechins and low-molecular-weight tannins extracted from less mature seeds, demonstrating that bitterness depends on phenolic composition rather than total content and supporting harvest delay for phenolic maturity.
Bitterness quality is a basic taste quality that represents the gustatory perception of bitter taste arising from stimulation of T2R bitter taste receptors by diverse chemical compounds, characterizing the aversive taste sensation that in grape and wine context is primarily associated with phenolic compounds including flavonoids, tannins, and hydroxycinnamic acid derivatives. Within the BFO framework, this quality specifically addresses the perception of bitterness as distinct from phenolic content or astringency, recognizing that bitterness is a taste mediated by gustatory receptors while astringency is a tactile sensation, though these sensations often co-occur with phenolic compounds contributing to both. The ontological significance lies in bitterness contributing complexity to wine taste structure when present at appropriate levels, while excessive bitterness creates unbalanced, harsh wines associated with underripe fruit, excessive skin extraction, or poor phenolic management. The physiological basis for bitter perception involves approximately 25 T2R receptor subtypes with different ligand specificities, explaining the diversity of compounds perceived as bitter and the individual variation in bitter sensitivity related to receptor polymorphisms, including the well-characterized PROP/PTC taster status affecting wine preference. The compositional basis for grape and wine bitterness includes monomeric phenolics (catechin, epicatechin showing direct bitter taste), procyanidins (oligomeric tannins with increasing bitterness at lower polymerization), and various non-phenolic bitter compounds including certain amino acids and oxidized wine components. The developmental changes in bitterness during grape ripening reflect seed tannin maturation, where extractable bitter tannins decline as polymerization increases and lignification reduces extraction, explaining why bitterness is associated with underripe character. Understanding bitterness quality enables harvest timing optimization for phenolic maturity, winemaking adjustment of extraction and fining to manage bitterness, and prediction of bitter perception from phenolic composition analysis.
Bitterness_Quality addresses the gustatory bitter taste perception, distinguishing from Astringency_Quality which is a tactile sensation. This quality serves as parent for intensity, localization, and persistence subclasses. The distinction between bitterness (taste) and astringency (touch) is ontologically important though often confused in casual wine description, as both are associated with phenolic compounds.
Bitterempfindung
Bittergeschmack
Bitterkeit
bitter perception
bitter sensation
bitter taste
bitter taste quality
bitterer Geschmack
Meyerhof W et al. (2010) The molecular receptive ranges of human TAS2R bitter taste receptors. Chemical Senses 35(2): 157-170 Peleg H et al. (1999) Bitterness and astringency of flavan-3-ol monomers, dimers and trimers. Journal of the Science of Food and Agriculture 79(8): 1123-1128
Bitterness Quality
Field evaluation of Black_Rot_Resistance_Disposition at the University of Georgia Tifton campus compared disease development across a diverse Vitis germplasm collection under natural epidemic conditions in the humid southeastern United States. The V. vinifera cultivar 'Chardonnay' (susceptible control) exhibited severe black rot with 85-95% of clusters showing mummified berries by véraison in unsprayed plots, and leaf lesions covering 40-60% of canopy area. The V. aestivalis-derived cultivar 'Norton' ('Cynthiana') demonstrated strong Black_Rot_Resistance_Disposition with only 5-12% cluster infection and scattered small leaf lesions (<5% canopy area) under identical conditions. Controlled inoculation experiments revealed that 'Norton' restricts G. bidwellii development during the biotrophic phase, with microscopic observation showing hyphal growth confined to 2-3 cell layers beneath the penetration site compared to extensive intercellular colonization in susceptible 'Chardonnay'. The interspecific hybrid 'Chambourcin' exhibited intermediate resistance (25-35% cluster infection), reflecting its mixed parentage combining susceptible V. vinifera with resistant North American species ancestry.
A Black_Rot_Resistance_Disposition is a Fungal_Resistance_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through defense processes upon encounter with the hemibiotrophic ascomycete Guignardia bidwellii (anamorph Phyllosticta ampelicida), the causal agent of grape black rot. This disposition represents the capacity to limit G. bidwellii infection, colonization, and reproduction across the pathogen's complex disease cycle affecting leaves, shoots, tendrils, and most critically, developing berries. Black rot is characterized by an initial biotrophic phase where the fungus colonizes living tissue asymptomatically, followed by a necrotrophic phase producing the characteristic tan lesions with dark borders on leaves and the devastating black, shriveled "mummified" berries that can destroy entire clusters. Resistance mechanisms operate at multiple infection stages including inhibition of conidial germination on plant surfaces, restriction of appressorial penetration, limitation of intercellular hyphal growth during the biotrophic phase, and containment of necrosis during the transition to necrotrophy. Unlike downy and powdery mildew, black rot resistance in Vitis involves both major gene effects (Rgr loci) and substantial quantitative variation. The disease is particularly destructive in humid continental climates of eastern North America, where G. bidwellii is endemic, making black rot resistance a primary breeding objective for cultivars adapted to these regions. European V. vinifera cultivars are generally highly susceptible, while North American species including V. rupestris, V. cinerea, and V. aestivalis exhibit varying levels of resistance.
Black_Rot_Resistance_Disposition is_a Fungal_Resistance_Disposition. Black rot caused by Guignardia bidwellii is a true fungus (Ascomycota), distinct from the oomycete causing downy mildew. Child classes include Rgr_Mediated_Resistance_Disposition for major gene resistance and Quantitative_Black_Rot_Resistance_Disposition for polygenic resistance. The disease is endemic to eastern North America, where wild Vitis species have evolved resistance, but can be devastating on susceptible V. vinifera cultivars introduced to these regions.
Black-Rot-Resistenz
Guignardia resistance disposition
Guignardia-Resistenz
Phyllosticta resistance
Schwarzfäule-Resistenz-Disposition
black rot immunity
Hoffman LE et al. (2002) Factors influencing the susceptibility of grapevine berries to black rot caused by Guignardia bidwellii, Phytopathology 92:1090-1098 Rex F et al. (2014) QTL mapping of black rot (Guignardia bidwellii) resistance in the grapevine rootstock 'Börner', Theoretical and Applied Genetics 127:1667-1677 Molitor D, Beyer M (2014) Epidemiology, identification and disease management of grape black rot and potentially useful metabolites of black rot pathogens for industrial applications, Annals of Applied Biology 165:305-317
Black Rot Resistance Disposition
Black rot symptom quality monitoring in Finger Lakes Concord documented the disease progression from early foliar symptoms through devastating fruit loss in an epidemic year with frequent extended wetting periods. Initial black rot symptom quality appeared in late May as 3-8 mm leaf lesions showing the characteristic tan center with dark brown border and concentric rings of black pycnidia visible under hand lens, with the angular lesion shape conforming to minor leaf veins providing field confirmation of G. bidwellii diagnosis distinct from other leaf spots. Shoot black rot symptom quality developed on green internodes as elongated dark cankers 1-3 cm in length, indicating heavy inoculum pressure and active secondary cycling. Fruit black rot symptom quality first appeared 3 weeks post-bloom as small brown spots on green berries, with affected berries progressing through the characteristic symptom sequence, as initial spotting (day 0), berry softening (day 3-4), tan-to-brown discoloration spreading across the berry (day 5-7), berry shriveling beginning (day 7-10), and completed mummification to hard, black raisin-like structure bearing abundant pycnidia (day 10-14). The black rot symptom quality assessment documented 65% fruit loss in unsprayed monitoring blocks, with mummified berries remaining attached to clusters and providing massive inoculum for subsequent seasons. Post-harvest sanitation to remove mummies was recommended as essential for reducing black rot symptom quality severity in the following year by eliminating the primary inoculum source.
Black rot symptom quality is a fungal infection indicator quality that inheres in grapevine organs infected by the fungus Guignardia bidwellii (anamorph Phyllosticta ampelicida) and represents the characteristic disease manifestations including the tan-to-brown leaf lesions with distinctive darker borders and black pycnidia arranged in concentric rings, the elongated canker-like lesions on shoots and petioles, and most significantly the fruit symptoms progressing from small brownish spots through rapid berry softening and shriveling to the characteristic hard, black, mummified berries bearing abundant pycnidia that serve as primary inoculum for subsequent seasons. This quality captures the diagnostic features of one of the most destructive fruit diseases in humid viticultural regions, recognizing that black rot can rapidly destroy the entire crop once fruit infection becomes established, with the distinctive symptom progression from initial infection through mummification providing critical diagnostic information for timing management interventions. The ontological significance of black rot symptom quality lies in its role enabling early disease detection and management timing, with the leaf and shoot symptoms appearing before fruit infection and providing warning of inoculum presence, while the characteristic fruit symptom progression indicates disease stage and the window for effective fungicide intervention. The biological basis of black rot symptoms encompasses the penetration and colonization of host tissues by G. bidwellii following spore germination during extended wetting periods, the initial biotrophic growth producing the characteristic bordered lesions as host cells die at the colony margin, the formation of pycnidia within dead tissue producing conidia for secondary spread, and the eventual colonization and mummification of infected berries that become the overwintering structures maintaining the pathogen between seasons. The environmental requirements for black rot infection include extended leaf wetness (minimum 6 hours at optimal temperatures near 27°C) that concentrates infection risk during warm, wet periods typical of humid continental and subtropical climates. The tissue-specific symptoms show consistent diagnostic features, with leaf lesions always showing the concentric ring pattern of pycnidia and the characteristically angular shape bounded by leaf veins, while fruit symptoms show the distinctive rapid progression from small spot to complete mummification within 10-14 days under favorable conditions.
Black_Rot_Symptom_Quality_(Guignardia_bidwellii) is_a Fungal_Infection_Indicator_Quality, representing the diagnostic symptoms of Guignardia bidwellii (Phyllosticta ampelicida) infection. The bordered lesions with concentric pycnidia rings are pathognomonic on leaves. Fruit mummification sequence (spot → soft → shrivel → black mummy) takes 10-14 days. Mummified berries provide overwintering inoculum making sanitation critical. Requires extended wetness (>6 hours) for infection limiting disease to humid climates. One of the most destructive fruit diseases in eastern North American viticulture.
Beerenmumifizierung
Black Rot-Zeichen
Guignardia symptoms
Guignardia-Symptome
Phyllosticta symptoms
Schwarzfäule-Symptome
berry mummification
black rot symptoms
Hoffman LE et al. (2002) Factors influencing the efficacy of myclobutanil and azoxystrobin for control of grape black rot. Plant Disease 86:769-774 Molitor D, Berkelmann-Loehnertz B (2011) Simulating the susceptibility of clusters to grape black rot infections depending on their phenological development. Crop Protection 30:1649-1654 Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests, 2nd Edition. APS Press
Black Rot Symptom Quality (Guignardia bidwellii)
Autumn Royal bears a Black_Table_Grape_Scion_Role as the dominant black seedless variety in California production, with large berries (6-8 g), deep blue-black color, firm crisp texture, and neutral-to-mild flavor providing extended late-season availability (September-November in California). Kyoho bears a Black_Table_Grape_Scion_Role as Japan's most prestigious table grape (and the world's largest-production single variety), with exceptionally large berries (15-20 g), deep purple-black slip-skin character, intense Concord-like labrusca flavor, and premium positioning commanding retail prices of ¥1000-3000+ per cluster in Japanese department stores. Midnight Beauty (ARRA 32) serves in a Black_Table_Grape_Scion_Role for early-season black seedless, with elongated berries (5-7 g), intense blue-black color, and firm texture reaching markets before Autumn Royal.
A fresh consumption table grape role that inheres in a grafted scion cultivar by virtue of its dark blue to black berry skin coloration, produced by high concentrations of anthocyanin pigments (predominantly malvidin, delphinidin, and petunidin derivatives) in the hypodermal cell layers. Black table grape scions develop the deepest pigmentation among table grape color classes, with anthocyanin concentrations often exceeding 500-1000 mg/kg berry weight, creating berries ranging from deep blue through purple to near-black depending on variety and ripening conditions. Black coloration is generally more stable and uniform than red, as the higher anthocyanin concentrations provide consistent color even under suboptimal temperature conditions. Black table grapes represent a smaller market segment than red or white varieties in Western markets but maintain strong positions in specific applications (premium gift packs, Asian markets valuing dark color symbolism). Major black table grape varieties include both seeded heritage cultivars (Ribier, Kyoho, Autumn Royal with vestigial seeds) and modern seedless varieties (Midnight Beauty, Autumn Royal, Adora, Sable Seedless).
Black_Table_Grape_Scion_Role represents the deepest-colored category, with varieties selected for intense, uniform blue-black pigmentation. The Kyoho phenomenon illustrates regional taste divergence—Japanese consumers prize the intense labrusca flavor and giant berry size that Western markets reject, supporting a premium gift-grape industry where individual clusters sell for US$20-100+. Conversely, Western markets favor mild-flavored, crisp-textured black seedless varieties (Autumn Royal, Midnight Beauty, Sable) for everyday consumption. Black table grape breeding has intensified recently, with several proprietary varieties (Sweet Sapphire/Moon Drop with elongated shape, Black Muscat types) creating premium niche markets. Color stability advantages make black varieties relatively easier to produce than color-challenged reds.
black eating grape
black table grape
blaue Tafeltraube
blue table grape
dark table grape
dunkle Esstraube
schwarze Tafeltraube
Creasy GL, Creasy LL (2009) Grapes, CABI Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press Yamada M, Sato A (2016) Advances in table grape breeding in Japan, Breeding Science 66: 34-45 Crisosto CH et al. (2002) Table grape postharvest quality, UC Davis Postharvest Technology
Black Table Grape Scion Role
The blade length description of Cabernet Franc documents medium blade length (OIV 065:5) with measurements of 140-175 mm from petiolar junction to central lobe tip on representative mature leaves from nodes 8-12 — this places Cabernet Franc in the central size range between its two offspring cultivars, the somewhat shorter-bladed Cabernet Sauvignon (120-155 mm, OIV 065:3-5, the deeply incised sinuses reducing effective blade length) and the longer-bladed Merlot (160-200 mm, OIV 065:5-7, the shallower sinuses preserving more blade extent). Blade length measurements were taken on 10 leaves per vine from 5 representative vines following the OIV protocol, with the measurement axis defined as the straight line from the petiolar attachment point to the distal tip of the central lobe, regardless of blade curvature or waviness.
A leaf blade description component that records the measured length of the fully expanded leaf blade from the point of petiole insertion to the tip of the central (terminal) lobe, expressed in millimeters and scored according to OIV descriptor 065 on a scale from very short (state 1, 240 mm), providing the primary dimensional measurement of leaf size that reflects cultivar-specific growth potential modulated by vine vigor and environmental conditions.
Blade_Length_Description documents the longitudinal dimension that serves as the primary proxy for overall leaf size in the OIV system. The measurement is straightforward in execution but requires attention to the interaction between blade length and lobe morphology — in deeply lobed cultivars (e.g., Cabernet Sauvignon), the central lobe extends well beyond the lateral lobes, making blade length primarily a measurement of central lobe projection, while in shallowly lobed or entire-leaved cultivars (e.g., some rootstock species), blade length represents the overall blade extent more accurately: this interaction means that blade length alone is an imperfect indicator of total blade area, and the combination of length with width measurements and lobe number provides a more complete size characterization. The environmental sensitivity of blade length necessitates multi-year measurement — studies at Geisenheim (Germany) and Montpellier (France) have documented 20-35% variation in mean blade length between high-vigor and low-vigor years for the same cultivar on the same site, confirming that leaf size descriptors should be interpreted as ranges rather than fixed values for each cultivar.
Blattlängen-Beschreibung
Blattspreiten-Längenmessung
Zentralachsen-Länge
blade longitudinal dimension
central axis length
leaf length measurement
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia UPOV (2011) Guidelines for the Conduct of Tests for Distinctness, Uniformity and Stability — Vine (Vitis L.), TG/50/9
Blade Length Description
The blade width description of Grenache documents broad leaves with maximum width of 180-230 mm measured perpendicular to the central axis at the level of the upper lateral sinuses — combined with blade length of 160-200 mm, this produces a length-to-width ratio of approximately 0.85-0.95 (nearly equidimensional to slightly wider than long), reflecting Grenache's characteristically orbicular leaf outline (OIV 067:4). This contrasts with the proportionally narrower leaves of Riesling (width 120-160 mm, length 140-175 mm, ratio 1.0-1.1, pentagonal with moderate lateral lobe projection) and the distinctly broader-than-long leaves of Cinsaut (width 200-250 mm, length 150-180 mm, ratio 0.65-0.80, approaching the reniform shape class with strongly developed lateral lobes).
A leaf blade description component that records the maximum width of the fully expanded leaf blade measured perpendicular to the central axis at the widest point, expressed in millimeters, providing the lateral dimensional measurement that together with blade length characterizes the overall proportions (length-to-width ratio) of the leaf and contributes to the discrimination between elongated, equidimensional, and transversely broad leaf forms across Vitis vinifera cultivars.
Blade_Width_Description captures the lateral dimension that, combined with blade length, defines the geometric proportions of the leaf. The length-to-width ratio is diagnostically more informative than either absolute dimension alone because it is less affected by vigor-driven size variation — a cultivar maintaining a characteristic ratio of 0.85 (wider than long) does so consistently across vigor levels, whereas both absolute length and width scale up or down together with vine vigor: this proportional stability makes the ratio a more genetically determined character than either component measurement. The width measurement point (the maximum transverse dimension) typically corresponds to the level of the upper lateral sinuses in five-lobed leaves, where the extension of the upper lateral lobes determines the maximum blade breadth — cultivars with strongly developed lateral lobes (e.g., Cinsaut, some Muscats) show width substantially exceeding length, while cultivars with reduced lateral lobes or deep upper lateral sinuses (e.g., Cabernet Sauvignon) show more equidimensional or elongated proportions despite the five-lobed outline.
Blattbreiten-Beschreibung
Blattspreiten-Breitenmessung
blade lateral dimension
leaf width measurement
maximale Blattbreite
maximum blade breadth
Blade Width Description
Petit Verdot bears a Blending_Red_Wine_Scion_Role in Bordeaux, contributing color intensity, floral aromatics, and structural tannins at 1-5% of left-bank assemblages dominated by Cabernet Sauvignon, rarely vinified as a single-varietal in its traditional home. Mourvèdre (Monastrell) serves in a Blending_Red_Wine_Scion_Role in southern Rhône GSM blends and Bandol, providing tannic backbone, meaty depth, and aging potential to complement Grenache's fruit and Syrah's aromatics. Carmenère historically bore a Blending_Red_Wine_Scion_Role in pre-phylloxera Bordeaux before near-extinction, now serving both blending and single-varietal roles in Chile. Colorino serves in a Blending_Red_Wine_Scion_Role in Chianti, adding color and body to Sangiovese-dominant assemblages at proportions up to 10-15%.
A red wine scion role that inheres in a grafted scion cultivar by virtue of its primary designation for contributing specific attributes (color, tannin, acidity, aromatic complexity, body, or softness) to blended wines rather than for single-varietal bottling. Blending red wine scions are valued for their capacity to complement, enhance, or balance the characteristics of partner varieties in assemblage, following regional blending traditions (Bordeaux, Rhône, Chianti, Rioja, Douro) or winemaker-driven style objectives. Blending scions may contribute structure and longevity (Cabernet Sauvignon, Tannat, Mourvèdre), aromatic lift and complexity (Cabernet Franc, Grenache, Muscat), color intensity (Petit Verdot, Teinturiers), acidity and freshness (Malbec, Cinsault), or softening texture (Merlot). The blending role does not preclude single-varietal expression—many blending varieties also produce excellent monovarietal wines—but recognizes that certain cultivars are primarily valued for their contribution to multi-variety assemblages in specific regional or stylistic contexts.
Blending_Red_Wine_Scion_Role captures the functional specialization of cultivars valued primarily for assemblage contribution within specific wine traditions. The classification is context-dependent Merlot serves as a primary variety (Premium_Red_Wine_Scion_Role) in Saint-Émilion and Pomerol but as a Blending_Red_Wine_Scion_Role in Médoc where it softens Cabernet Sauvignon-dominant wines. Grenache functions as a blending component in many Châteauneuf-du-Pape and Priorat wines but as the sole or dominant variety in old-vine Garnacha bottlings from Gredos or Calatayud. Regulatory frameworks (AOC, DOC, DO) often specify permitted blending varieties and proportions, institutionalizing traditional assemblage formulas.
Cuvée-Traube
Rotwein-Verschnittrebe
Verschnittsorte
assemblage variety
blending grape
blending red variety
component variety
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Peynaud E, Blouin J (2006) The Taste of Wine: The Art and Science of Wine Appreciation 2nd Ed, Wiley Jackson RS (2014) Wine Science 4th Ed, Academic Press Brook S (2001) The Complete Bordeaux, Mitchell Beazley
Blending Red Wine Scion Role
Blending specification for a Bordeaux Grand Cru Classé (Pauillac, 2023 vintage) documents the assemblage of 72% Cabernet Sauvignon (from 3 parcels on gravel terraces, providing structural backbone and cassis-graphite aromatic core), 18% Merlot (from clay-limestone parcels, contributing mid-palate flesh and plum-chocolate richness), 7% Cabernet Franc (from alluvial soils, adding violet-herbal aromatic lift and tannin finesse), and 3% Petit Verdot (from the warmest exposition, intensifying color and adding spice-ink concentration), selected from 42 individual barrel lots through 6 rounds of blending trials conducted December-February, with the final blend targeting 13.8% alcohol, TA 5.8 g/L, pH 3.58, total phenolics 75 AU, and a sensory profile balancing power, elegance, and 25+ year aging potential. Blending specification for Grüner Veltliner Smaragd from a Wachau single-vineyard site documents a simpler assemblage of 100% Grüner Veltliner from the designated Ried, with blending decisions limited to the proportion of stainless steel (70%) versus large oak cask (30%) components, targeting seamless integration of fruit purity with textural depth.
Blending specification is a winemaking protocol entity that documents the compositional design of the final wine through the systematic combination of component lots differentiated by cultivar, vineyard parcel, clone, fermentation vessel, oak treatment, or press fraction, specifying the proportion of each component, the analytical and sensory criteria guiding assemblage decisions, the target wine profile (compositional parameters, sensory descriptors, style reference), and the sequence of blending trials through which the final cuvée is determined, providing the formulation framework that transforms individual wine lots into a coherent, intentional finished product expressing the winemaker's aesthetic vision and the producer's house style.
Blending specification captures the integrative, synthetic phase of winemaking where individual components — each reflecting specific viticultural origins, fermentation conditions, and aging environments — are combined into a unified whole that ideally exceeds the quality of any individual component. The philosophical spectrum of blending ranges from the Bordeaux tradition of multi-cultivar assemblage (where blending across varieties, parcels, and oak treatments creates complexity and consistency) through the Burgundian single-cultivar, single-vineyard model (where "blending" occurs primarily across barrel lots of the same wine) to the rising Austrian and German emphasis on single-vineyard Lagenwein purity (where blending is limited to integrating different vessels and press fractions from one site). The blending trial process (assemblage) is conducted through systematic bench trials where small-volume combinations are prepared, analytically characterized, and sensorially evaluated, typically over multiple sessions spanning weeks to months, with the winemaker and tasting panel iteratively refining proportions to achieve the target wine profile. The analytical framework for blending evaluation includes color assessment (ensuring adequate depth and appropriate hue), acid-pH balance (verifying that the blend falls within target ranges), tannin structure (assessing whether tannin from different sources integrates harmoniously), and aromatic profile (confirming that component aromas complement rather than conflict). The economic dimension of blending is significant — the decision to include or exclude specific lots in the grand vin versus second wine (or bulk sale) directly determines production volume and quality positioning, with first-wine selection rates varying from 35-50% of production at top Bordeaux estates to near-100% at estates producing a single wine. Blending specification should document not only the final proportions but also the lots that were evaluated and excluded, as this information provides valuable feedback for vineyard management and future vintage planning.
Assemblage-Spezifikation
Cuvée-Spezifikation
Cuvéegestaltung
Verschnittprotokoll
Zusammenstellungsdaten
assemblage specification
blend composition
blending protocol
cuvée specification
wine blend data
Peynaud E, Blouin J (2006) The Taste of Wine: The Art and Science of Wine Appreciation, 4th Ed, Wiley, pp.: 285-320 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1, 2nd Ed, Wiley, pp.: 395-440 Boulton RB et al. (1999) Principles and Practices of Winemaking, Springer, pp.: 425-460 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 647-670
Blending Specification
Block vigor uniformity quality assessment in a 5-hectare Margaret River Chardonnay vineyard compared pruning weight coefficients of variation across 200 systematically sampled vines over three seasons. Initial CV of 31% with structured spatial pattern prompted soil investigation revealing buried gravel lens creating localized drainage variation. Installation of targeted subsurface drainage in affected zones improved block vigor uniformity quality to CV 19% within two seasons, enabling transition from three-pick selective harvest to two-pick harvest with improved operational efficiency while maintaining fruit quality segregation for reserve and standard tier wine production.
Block vigor uniformity quality is a vigor uniformity quality that inheres in a vineyard block or management unit and represents the consistency of vegetative growth characteristics across the population of individual vines comprising the block as manifested through inter-vine variation in canopy dimensions, pruning weights, shoot counts, and vegetation indices when assessed across the spatial extent of the production area. This quality determines the feasibility of uniform block-level management practices versus the requirement for differential or precision management approaches, and fundamentally influences fruit quality consistency and harvest logistics efficiency. The ontological significance of block vigor uniformity quality lies in its emergence from the interaction between soil spatial variability, microclimate gradients, planting material uniformity, management history consistency, and pest and disease distribution patterns that collectively determine whether vines across a block perform as a relatively homogeneous population or exhibit functionally distinct subpopulations requiring differentiated treatment. High block vigor uniformity quality enables standardized management including uniform irrigation scheduling, single-timing canopy operations, and consolidated harvest, while low uniformity necessitates either management zone delineation with differentiated practices or acceptance of quality variation in harvested fruit. The measurement of block vigor uniformity quality employs statistical descriptors of inter-vine variation including coefficient of variation in pruning weights or trunk circumference for ground-based assessment, and spatial variability metrics applied to remotely sensed vegetation indices for comprehensive block characterization. Coefficient of variation values below 15% generally indicate manageable uniformity for standard practices, while values exceeding 25% suggest substantial heterogeneity warranting precision approaches or block subdivision. The spatial structure of vigor variation contributes additional information beyond simple statistical dispersion, with spatially autocorrelated patterns suggesting environmental drivers amenable to zone management while spatially random variation may indicate vine-level factors such as virus infection, gopher damage, or replant effects requiring individual vine intervention. Factors promoting block vigor uniformity quality include consistent soil preparation and drainage prior to planting, certified uniform planting material, well-designed and maintained irrigation systems delivering uniform application, integrated pest management preventing localized disease establishment, and proactive replanting maintaining population completeness.
Block_Vigor_Uniformity_Quality is_a Vigor_Uniformity_Quality, representing the vineyard block or management unit scale of uniformity assessment. This class addresses inter-vine variation across populations rather than intra-vine variation, with the coefficient of variation across vine pruning weights or canopy parameters serving as standard metric. Spatial pattern analysis adds dimension beyond statistical dispersion. Distinguished from Within_Vine_Vigor_Uniformity_Quality by scale and from NDVI_Uniformity_Quality by measurement methodology though NDVI provides efficient means of assessing block uniformity.
Anlagengleichmäßigkeit
Blockgleichmäßigkeit
Zwischenstock-Wuchskonsistenz
block-level growth homogeneity
inter-vine vigor consistency
räumliche Wuchshomogenität
spatial vigor uniformity
vineyard block uniformity
Bramley RGV, Hamilton RP (2004) Understanding variability in winegrape production systems 1. Within vineyard variation in yield over several vintages. Australian Journal of Grape and Wine Research 10:32-45 Tisseyre B et al. (2008) Spatial variability of yield and quality in vineyards using precision viticulture techniques. Journal International des Sciences de la Vigne et du Vin 42:261-268 Arnó J et al. (2009) Review. Precision Viticulture. Research topics, challenges and opportunities in site-specific vineyard management. Spanish Journal of Agricultural Research 7:779-790
Block Vigor Uniformity Quality
Bloom appearance assessment comparing hand-harvested versus machine-harvested Cabernet Sauvignon reveals intact, uniformly distributed heavy bloom (rating 9/10) on hand-harvested clusters indicating minimal berry-to-berry contact and careful handling, versus disturbed, patchy bloom (rating 4/10) on machine-harvested fruit where mechanical action has abraded the wax layer, with the bloom difference correlating with higher native yeast populations on hand-harvested fruit (enabling spontaneous fermentation if desired) and reduced desiccation during transport, demonstrating bloom as a practical quality indicator for harvest method and handling assessment.
Bloom appearance quality is a surface appearance quality that represents the whitish, powdery coating visible on grape berry surfaces arising from epicuticular wax crystallization, characterizing the pruina layer's integrity, uniformity, and visual impact as indicators of berry health, handling care, and environmental exposure history. Within the BFO framework, this quality specifically addresses the epicuticular wax coating dimension of grape surface appearance, distinguishing bloom from underlying cuticle gloss while recognizing that bloom (pruina) represents the outermost protective layer whose condition reflects both developmental and post-harvest handling factors. The ontological significance lies in bloom serving as a visual indicator of grape condition and handling history, where intact, uniform bloom suggests healthy development and careful handling while disturbed or absent bloom indicates physical contact, disease pressure, or environmental damage that may affect grape quality and winemaking suitability. The physical basis for bloom appearance involves light scattering from epicuticular wax crystals, primarily composed of oleanolic acid and related triterpenoids, that form plate-like or tubular structures protruding from the cuticle surface, with crystal morphology and density determining bloom intensity and appearance from barely visible to prominently waxy depending on variety and conditions. The compositional and developmental determinants of bloom include variety-specific wax production capacity (some varieties producing heavier bloom than others), berry developmental stage (bloom typically increasing through ripening), environmental factors (UV exposure, humidity affecting wax crystallization), and the wild yeast populations that colonize the bloom layer and contribute to spontaneous fermentation capacity. The quality implications of bloom assessment include grape health evaluation (intact bloom suggesting freedom from handling damage and disease), fermentation potential (bloom-associated yeasts contributing to native fermentation), and storage suitability (intact bloom providing desiccation protection). Understanding bloom appearance quality enables grape condition assessment based on pruina integrity, variety identification from characteristic bloom expression, and harvest handling evaluation detecting bloom disturbance from mechanical damage.
Bloom_Appearance_Quality_(Pruina) addresses the epicuticular wax coating specific to grape berry surfaces. This quality provides indicators of grape health, handling care, and native yeast populations through visual assessment of wax layer integrity. The class supports viticultural quality evaluation and harvest handling assessment through bloom observation, with intact bloom generally indicating higher quality fruit condition.
Duftfilm
Pruina
Reifbelag
Wachsschicht
berry bloom
epicuticular wax coating
pruina
wax bloom
Casado CG, Heredia A (1999) Structure and dynamics of reconstituted cuticular waxes of grape berry cuticle. Journal of Experimental Botany 50(331): 175-182 Comménil P et al. (1997) Determination of the surface structure of wax deposits on grape berry cuticle. Annals of Botany 79(3): 259-264
Bloom Appearance Quality (Pruina)
The bloom description of Cabernet Sauvignon documents a heavy, uniform epicuticular wax bloom covering the entire berry surface — the wax crystallites (primarily oleanolic acid and related triterpenoids, with minor long-chain aliphatic components) form a dense, approximately 1-2 μm thick coating that produces the characteristic dusty, powder-blue appearance of Cabernet Sauvignon clusters at harvest, with the bloom most conspicuous on the exposed outer berries and reduced or absent on inner cluster berries where physical contact with adjacent berries has rubbed the wax layer away. The heavy bloom of Cabernet Sauvignon contrasts with the light bloom of Chasselas (where the thin wax layer does not substantially modify the yellow-green skin color visible through the transparent coating), the very heavy bloom of some rootstock species (where thick wax deposits produce a distinctly chalky white appearance), and the virtually absent bloom of some table grape cultivars selected for the glossy, shiny skin appearance preferred in Asian table grape markets.
A berry skin description component that records the presence, density, and distribution of the epicuticular wax bloom — the thin layer of hydrophobic wax crystallites deposited on the outer surface of the berry cuticle — that creates the dusty, matte, whitish-blue coating visible on the surface of dark-skinned grape berries, with the bloom serving as a physical barrier reducing water loss, modifying light reflectance, and influencing the adhesion of fungal spores, spray residues, and indigenous yeast populations on the berry surface.
Bloom_Description documents a surface feature that has multiple biological, viticultural, and winemaking implications. The epicuticular wax bloom is secreted by the epidermal cells during berry development, with wax deposition beginning during Phase I and continuing through Phase III, and the final bloom density reflecting both the rate of wax biosynthesis (genetically determined, varying between cultivars) and the physical integrity of the deposited wax layer (affected by environmental factors including rainfall, wind abrasion, and physical contact within compact clusters): the wax composition is dominated by triterpenoids (principally oleanolic acid, comprising 60-80% of total wax) with smaller amounts of long-chain fatty alcohols, aldehydes, and alkanes, and this composition varies between cultivars in ways that affect the physical properties of the bloom — high oleanolic acid content produces a powdery, easily rubbed crystalline bloom, while higher proportions of long-chain aliphatics produce a smoother, more adherent wax layer. The bloom has significance for natural winemaking because the epicuticular wax surface harbors the indigenous yeast populations (principally Saccharomyces cerevisiae and non-Saccharomyces species) that drive spontaneous fermentation — the wax crystallite structure provides physical niches that protect yeast cells from desiccation and UV exposure, and the wax composition may selectively promote or inhibit different yeast species, contributing to the site-specific microbial terroir that advocates of natural winemaking consider an essential component of wine identity.
Bereifungs-Beschreibung
Kutikularwachs-Dokumentation
Wachsüberzug-Bewertung
berry wax coating description
epicuticular wax record
surface bloom assessment
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Casado CG and Heredia A (1999) Structure and dynamics of reconstituted cuticular waxes of grape berry cuticle, Journal of Experimental Botany 50: 175-182 Commenil P et al. (1997) Surface waxes of grape berry and host-pathogen interaction, Journal of the International Society for Horticultural Science 484: 525-529
Bloom Description
Blue hue contribution to the overall color of Malbec berries accounts for the cultivar's distinctively dark, inky appearance, with spectrophotometric analysis revealing absorbance peaks in the blue region (520-540 nm) that exceed those of Merlot or Cabernet Sauvignon at equivalent total anthocyanin concentrations this enhanced blue character results from Malbec's anthocyanin profile dominated by malvidin-3-glucoside with high proportions of acetylated and coumaroylated derivatives that form stable blue co-pigmentation complexes, producing the near-black wines with purple-blue rim that define the Malbec color profile.
Blue_Hue_Quality represents the chromatic character of grapevine tissues displaying coloration in the blue portion of the visible spectrum, corresponding to wavelengths approximately 450-495 nm and CIELAB hue angles of approximately 240-280° (or negative a* with positive b* values indicating the blue-violet region). Blue hue in grapevine tissues results from anthocyanin pigments, particularly delphinidin, petunidin, and malvidin, under conditions that favor the quinoidal base and anionic forms of these molecules. The expression of blue versus red hue depends on vacuolar pH, with higher pH values shifting anthocyanin equilibrium toward blue forms, and on co-pigmentation with colorless flavonoids and metal ions that can produce blue-shifted complexes. True blue hue is relatively rare in grape berries because the typical vacuolar pH of 3.0-3.5 favors red flavylium cation forms, but the characteristic blue-black appearance of cultivars like Cabernet Sauvignon results from very high anthocyanin concentrations where the minor proportion in blue forms contributes to the overall dark appearance. Blue hue is also observed in the waxy bloom (pruina) that covers berry surfaces, where epicuticular wax platelets scatter blue light preferentially, creating a blue-grey appearance over underlying red or black pigmentation. This bloom-derived blue coloration is distinct from pigment-based blue hue but contributes to the visual impression of blue-black berries. Environmental factors including temperature modulate blue hue expression, with cool conditions favoring the blue spectral components of anthocyanin-based colors.
Blue_Hue_Quality represents the blue region of the color spectrum and is relatively uncommon as a dominant hue in grape tissues because vacuolar pH conditions favor red anthocyanin forms. Blue hue typically contributes as a component of blue-black or purple coloration rather than appearing as a pure blue. The contribution of epicuticular wax bloom to perceived blue coloration should be distinguished from pigment-based blue hue in color assessment. The term blue-black in cultivar descriptions (e.g., OIV berry color category) indicates very dark coloration with blue components rather than true blue hue.
Blauton
azure hue
blaue Anthocyanfärbung
blaue Färbung
blue anthocyanin coloration
blue color
indigo quality
indigofarbene Tönung
Boulton R (2001) The Copigmentation of Anthocyanins and Its Role in the Color of Red Wine: A Critical Review. American Journal of Enology and Viticulture 52(2): 67-87 Casassa LF and Harbertson JF (2014) Extraction, Evolution, and Sensory Impact of Phenolic Compounds During Red Wine Maceration. Annual Review of Food Science and Technology 5: 83-109
Blue Hue Quality
Body description for Amarone della Valpolicella (2020 vintage, 16.0% alcohol, 38 g/L dry extract) evaluates as "full-bodied" — the exceptional alcohol concentration, elevated glycerol (14 g/L from partial grape desiccation), substantial phenolic extract, and residual sugar (5.5 g/L) collectively produce a dense, viscous, mouth-coating mouthfeel of remarkable weight and persistence, representing the upper extreme of still wine body. Body description for Mosel Riesling Kabinett (2024 vintage, 8.5% alcohol, 45 g/L RS) records "light-bodied" — despite the significant residual sugar contributing viscosity, the very low alcohol produces a delicate, almost ethereal palate weight that defines the Kabinett style's characteristic elegance and drinkability, illustrating alcohol's dominance over sugar in determining perceived body. Body description for Grüner Veltliner Smaragd from the Wachau (2024 vintage, 13.5% alcohol) assesses at "medium-full" — the moderate-to-high alcohol, 6 months of lees contact providing mannoprotein-enhanced texture, and concentrated fruit extract produce a substantial but not heavy palate weight appropriate to the Smaragd classification.
Body description is a taste description entity that records the perceived weight, fullness, and viscosity of wine on the palate, assessed on a structured scale from light-bodied (thin, delicate, watery mouthfeel) through light-medium, medium, medium-full, to full-bodied (rich, dense, viscous, mouth-filling), integrating the combined sensory contributions of alcohol (the primary determinant of perceived weight through viscosity and warmth), extract concentration (dissolved solids including glycerol, organic acids, minerals, phenolics, and residual sugar), and tannin-phenolic structure (contributing to textural density in red wines), and providing the structural assessment that establishes the wine's weight class and informs food pairing, serving temperature, and stylistic classification.
Body description quantifies the perceived textural weight of wine on the palate, a complex multisensory perception arising from the integration of viscosity (a physical property), warmth (an alcohol-mediated trigeminal sensation), and density (a dissolved solids-related perception) into a unified impression of palate weight. Research has consistently demonstrated that alcohol content is the single strongest predictor of perceived body, accounting for approximately 60-70% of the variance in body ratings across diverse wine types — the viscosity-enhancing effect of ethanol at wine concentrations (8-16% vol.) and the trigeminal warmth sensation combine to produce the perceived weight that scales with alcohol level. Glycerol, though frequently cited as a major body contributor, has a more modest effect than commonly believed — at typical wine concentrations (5-10 g/L), glycerol's viscosity contribution is below the difference threshold for most tasters, though at the elevated concentrations found in botrytized wines and Amarone (12-20 g/L), its contribution becomes perceptible. Dissolved sugars contribute significantly to body in sweet wines — residual sugar at >20 g/L measurably increases viscosity and perceived weight — but in dry wines (14.5%), though these thresholds are modulated by extract, tannin, and sugar content.
Fülle
Körperbeschreibung
Körperfülle
Mundgewicht
Weingewicht
fullness description
mouthweight
palate weight description
weight description
wine body
Pickering GJ et al. (1998) The influence of ethanol concentration on the temporal perception of viscosity and density in white wine. American Journal of Enology and Viticulture 49(3): 306-318 Noble AC, Bursick GF (1984) The contribution of glycerol to perceived viscosity and sweetness in white wine. American Journal of Enology and Viticulture 35(2): 110-112 Peynaud E, Blouin J (2006) The Taste of Wine, 4th Ed, Wiley, pp.: 200-240 Runnebaum RC et al. (2011) Analysis of sulfur compounds in wine using hydrogen sulfide as a reducing agent. Journal of Agricultural and Food Chemistry 59(13): 7286-7293
Body Description
Body quality assessment across white wine styles reveals the range from Mosel Riesling Kabinett (light body rating 3/10, delicate weight appropriate for 8% alcohol and off-dry style, enabling refreshing consumption) through Côtes du Rhône Blanc (medium body 5.5/10, moderate weight supporting food pairing versatility) to barrel-fermented Meursault (full body 8/10, substantial weight from 13.5% alcohol, malo, and lees contact appropriate for rich, complex style), demonstrating how body appropriate to style determines quality perception rather than absolute body level.
Body quality is a mouthfeel quality that represents the perceived weight, fullness, and substance of wine in the mouth, characterizing the overall sense of density and palate presence that ranges from light and ethereal through medium and balanced to full and substantial depending on compositional factors and wine style. Within the BFO framework, this quality specifically addresses the weight dimension of mouthfeel, distinguishing perceived density from viscosity (flow resistance) and texture (surface character) while recognizing that body provides the fundamental structural impression upon which other mouthfeel qualities are perceived. The ontological significance lies in body serving as the primary structural parameter in wine style classification, where body level defines the basic wine category (light-bodied versus full-bodied) and creates expectations for flavor intensity, food pairing suitability, and appropriate consumption context, making body assessment foundational for wine characterization. The perceptual basis for body involves integration of multiple sensory inputs including viscosity (flow resistance contributing to weight impression), density (dissolved solids adding substance), oral coating (persistence of wine on mouth surfaces), and the resistance to oral manipulation (heavier wines feeling more substantial when moved in mouth), with these inputs synthesized into the unified weight perception. The compositional determinants of body include alcohol content (primary contributor, with each 1% alcohol adding perceptible weight), glycerol concentration (contributing viscosity and smoothness), residual sugar (increasing density and substance), extract level (dissolved phenolics, polysaccharides adding palate presence), and the interaction among these factors creating the characteristic body profiles of different wine styles. The style-appropriateness of body level is critical, as light body is desired in styles emphasizing freshness and drinkability (Muscadet, Vinho Verde) while full body is expected in styles emphasizing power and concentration (Amarone, Barossa Shiraz), with body assessment requiring calibration to style expectation rather than universal preference for fullness. Understanding body quality enables wine style classification based on weight category, style-appropriate body evaluation, and quality assessment incorporating appropriate substance for wine type.
Body_Quality serves as the parent class for body level categories (Light, Medium, Full), representing the weight dimension of mouthfeel. This quality is fundamental for wine style classification and provides the structural context for other sensory attributes. The class emphasizes style-appropriateness, as optimal body level varies dramatically among wine categories and quality requires appropriate body for style rather than maximum body.
Körper
Körperdichte
body level
mouth density
palate fullness
wine weight
Nurgel C, Pickering GJ (2005) Contribution of glycerol, ethanol and sugar to the perception of viscosity and density elicited by model white wines. Journal of Texture Studies 36(3): 303-323 Runnebaum RC et al. (2011) Analysis of the key components in model white wine and their contributions to body. Journal of Sensory Studies 26(3): 196-206
Body Quality
Body visual impression assessment in flight of white wines rates Picpoul de Pinet at 3/10 (thin, watery swirl with rapid draining suggesting light body), unoaked Chardonnay at 5/10 (moderate coating and flow rate suggesting medium body), and barrel-fermented Hermitage Blanc at 8/10 (thick, persistent coating with slow flow suggesting full body), with these visual body predictions largely confirmed on tasting though the Chardonnay's elevated acidity creates lighter perceived body than visual impression alone would suggest, demonstrating both the utility and limitations of visual body assessment.
Body visual impression quality is a viscosity appearance quality that represents the overall visual sense of wine weight, density, and substance conveyed through combined observation of swirling behavior, coating persistence, and flow characteristics, characterizing the gestalt impression of light versus full body that creates expectations for palate texture before tasting. Within the BFO framework, this quality specifically addresses the integrated body impression arising from multiple viscosity-related cues, distinguishing holistic body assessment from specific leg or flow observations while recognizing that visual body impression synthesizes various rheological indicators into a unified weight expectation. The ontological significance lies in body visual impression providing the primary visual-to-tactile prediction pathway, where the overall sense of wine weight observed in the glass creates the expectation framework for palate body assessment, though this prediction is approximate given the additional factors (acidity, tannin, carbonation, temperature) affecting perceived mouthfeel beyond true viscosity. The perceptual basis for body visual impression involves integration of multiple visual cues including swirling resistance (heavier wines rotating more slowly), coating behavior (fuller wines leaving more persistent film), leg characteristics (thicker legs suggesting fuller body), and overall visual density (the sense of substance versus transparency in the swirling wine), with these cues synthesized into a holistic light-to-full body prediction. The compositional correlates of visual body impression include alcohol content (primary contributor to visual viscosity), glycerol (adding oily appearance), residual sugar (creating visible density), extract level (contributing to overall substance impression), and the interaction among these factors creating the characteristic visual signatures of light, medium, and full-bodied wine styles. The style associations of visual body impression align with wine category expectations, where light-bodied styles (Muscadet, Vinho Verde) show thin, watery visual impression while full-bodied styles (Châteauneuf-du-Pape, Barossa Shiraz) display thick, substantial visual impression, with body visual quality both confirming style expectations and enabling blind assessment body estimation. Understanding body visual impression quality enables style-appropriate evaluation assessing visual body against category expectation, blind tasting body estimation from visual cues, and appropriate mouthfeel expectation setting while acknowledging the limitations of visual body prediction.
Body_Visual_Impression_Quality addresses the holistic visual sense of wine weight synthesized from multiple viscosity cues. This quality provides the primary visual-to-tactile prediction, creating mouthfeel expectations before tasting. The class supports blind tasting body estimation and style confirmation while acknowledging that perceived body involves factors beyond viscosity that limit visual prediction accuracy.
Körpererscheinung
apparent density
body appearance
scheinbare Dichte
visual substance
visual weight
visuelle Fülle
visueller Körpereindruck
Pickering GJ (2000) Low- and reduced-alcohol wine: a review. Journal of Wine Research 11(2): 129-144 Jackson RS (2014) Wine Science: Principles and Applications, 4th Ed. Academic Press, San Diego, pp. 831-887
Body Visual Impression Quality
Long-term monitoring of Bois_Noir_Resistance_Disposition at the Julius Kühn-Institut, Siebeldingen, Germany, tracked BN incidence, symptom development, and recovery in a 20-cultivar trial block over 12 consecutive seasons in a region with endemic H. obsoletus populations and established phytoplasma reservoirs in adjacent ruderal vegetation. Cumulative infection rates (at least one positive PCR test during monitoring period) ranged from 15% ('Riesling') to 68% ('Dornfelder'). However, infection rate poorly predicted disease impact due to dramatic cultivar differences in symptom expression and recovery. 'Dornfelder' exhibited high susceptibility with severe symptoms (mean score 4.1), 8% spontaneous recovery, and 12% cumulative mortality. In contrast, 'Portugieser' showed equivalent infection rate (62%) but demonstrated strong Bois_Noir_Resistance_Disposition with mild symptoms (score 1.6), 65% spontaneous recovery within 2-3 years of symptom onset, and only 2% mortality. Recovery was associated with reduced but not eliminated phytoplasma titers, suggesting tolerance and host-mediated suppression rather than pathogen clearance. 'Riesling' represented a third phenotype with apparent infection resistance (low infection rate despite equivalent vector exposure) combined with moderate tolerance when infected (score 2.4, 40% recovery). These distinct Bois_Noir_Resistance_Disposition patterns enable cultivar-specific management recommendations, with tolerant cultivars like 'Portugieser' requiring less aggressive roguing of infected vines than susceptible cultivars like 'Dornfelder'.
A Bois_Noir_Resistance_Disposition is a Phytoplasma_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that limit infection, multiplication, systemic spread, symptom development, or recovery following infection by 'Candidatus Phytoplasma solani', the causal agent of Bois noir (BN), a widespread grapevine yellows disease with distinct epidemiology from Flavescence dorée. This disposition represents the capacity to resist or tolerate infection by a phytoplasma that, unlike FD, is not grapevine-specific and is transmitted by the polyphagous planthopper Hyalesthes obsoletus (Hemiptera: Cixiidae) from herbaceous reservoir hosts including bindweed (Convolvulus arvensis), nettle (Urtica dioica), and other plants where the vector completes its life cycle. Bois noir symptoms are indistinguishable from FD, including leaf rolling with discoloration, incomplete cane lignification, and berry shriveling, but BN typically shows lower incidence, slower epidemic development, and higher spontaneous recovery rates than FD. Because H. obsoletus feeds on grapevine only occasionally as an adult seeking shelter, and because phytoplasma reservoir hosts are essential for the disease cycle, Bois_Noir_Resistance_Disposition operates in a different epidemiological context than FD resistance. Resistance mechanisms include limitation of phytoplasma multiplication in phloem following incidental vector feeding, tolerance to phytoplasma-induced phloem dysfunction, and recovery capacity enabling symptom remission and restoration of productivity. Substantial cultivar variation exists, with some V. vinifera cultivars showing tolerance or high recovery rates that enable economically viable production despite endemic BN presence.
Bois_Noir_Resistance_Disposition is_a Phytoplasma_Resistance_Disposition. BN differs epidemiologically from FD in that the vector H. obsoletus does not complete its life cycle on grapevine and feeds only incidentally on vines, with herbaceous reservoir hosts essential for disease maintenance. This epidemiology means that vector resistance in grapevine is less relevant than for FD, and host tolerance/recovery capacity becomes the primary resistance mechanism of breeding interest. BN is not subject to mandatory control regulations equivalent to FD quarantine status in most regions, though monitoring and management are recommended. Spontaneous recovery rates are generally higher for BN than FD, making recovery capacity an important component of Bois_Noir_Resistance_Disposition.
BN resistance disposition
BN-Toleranz
Bois-noir-Resistenz
Candidatus Phytoplasma solani resistance
Schwarzholzkrankheit-Resistenz-Disposition
black wood tolerance
Quaglino F et al. (2013) 'Candidatus Phytoplasma solani', a novel taxon associated with stolbur and Bois noir related diseases of plants, International Journal of Systematic and Evolutionary Microbiology 63:2879-2894 Maixner M (2011) Recent advances in Bois noir research, Petria 21:95-108 Langer M, Maixner M (2004) Molecular characterisation of grapevine yellows associated phytoplasmas of the stolbur-group based on RFLP-analysis of non-ribosomal DNA, Vitis 43:191-199
Bois Noir Resistance Disposition
Merlot grafted on 3309 Couderc rootstock showing border cells around primary root tip, microscopic examination of rhizosphere revealing abundant rounded cells 22 micrometers average diameter embedded in mucilage matrix, border cells recently detached from cap periphery showing initial viability with intact membranes, FDA staining showing 65% of border cells viable, border cell population estimated 850 cells per root tip creating defensive barrier, mucilage matrix 40-80 micrometers thick around root tip, border cells providing pathogen defense and soil conditioning
Border cells are root cap cells that have separated from cap surface and exist as free-living cells in rhizosphere surrounding root tip, typically produced at rate 100-10,000+ cells per root tip per day depending on species and conditions, characterized by rounded or irregular shape after separation from columnar cap organization, initial viability with intact membranes and active metabolism lasting hours to days after detachment, extensive mucilage production creating gelatinous matrix surrounding cells, and programmed senescence leading to death and decomposition within days of separation, serving defensive functions including pathogen deterrence through physical trapping in mucilage, chemical defense through production of antimicrobial compounds, decoy function attracting pathogens away from sensitive cap and meristem, and soil conditioning through mucilage secretion and organic matter contribution, with border cells demonstrating unique plant cell behavior including survival as individual cells without tissue connection, continued metabolism and protein synthesis for period after detachment, active secretion and cell-cell communication, and eventual programmed death contributing cellular contents to rhizosphere, and representing dynamic interface between root and soil environment. Border cells origin and production includes= showing cap surface detachment (border cells originating from programmed separation of peripheral cap cells particularly from lateral cap margins), exhibiting continuous production (border cells continuously generated as cap cells reach peripheral position and undergo separation), demonstrating high production rates (root tips producing hundreds to thousands of border cells daily creating substantial cell population), revealing species variation (border cell production rate varying widely among species with some showing prolific production others minimal), and displaying environmental modulation (border cell production potentially affected by soil conditions, pathogens, or other environmental factors). Border cells morphology includes= showing rounded shape (border cells typically losing columnar shape becoming spherical or irregular after separation), exhibiting size variation (border cells typically 15-40 micrometers diameter varying with origin and age), demonstrating variable appearance (individual border cells showing different morphologies reflecting different ages and physiological states), revealing cell wall retention (border cells maintaining cell walls after separation though walls potentially modified), and displaying mucilage coating (border cells surrounded by thick polysaccharide matrix creating gelatinous appearance). Border cells viability and metabolism includes= showing initial viability (newly detached border cells with intact membranes and functional organelles capable of continued metabolism), exhibiting limited lifespan (border cells typically remaining viable hours to days after detachment depending on conditions), demonstrating metabolic activity (viable border cells showing respiration, protein synthesis, and secretion), revealing progressive decline (border cell metabolism declining over time as cells age in rhizosphere), and displaying eventual death (border cells undergoing programmed cell death within days of detachment). Border cells mucilage production includes= showing extensive secretion (border cells producing large quantities of mucilage after detachment), exhibiting mucilage matrix (collectively secreted mucilage creating gelatinous matrix embedding border cells), demonstrating composition complexity (border cell mucilage containing polysaccharides, proteins, DNA, and other components), revealing defensive properties (mucilage showing antimicrobial activities and creating physical barrier), and displaying soil conditioning (mucilage affecting soil structure and chemistry in root tip region). Border cells defensive functions includes= providing pathogen trap (border cell mucilage physically trapping fungal spores and bacterial cells preventing cap colonization), exhibiting decoy function (border cells attracting pathogen attack diverting from sensitive cap and meristem tissues), demonstrating chemical defense (border cells producing and secreting antimicrobial compounds inhibiting pathogen growth), revealing sacrificial barrier (border cells representing expendable defensive layer that can be colonized or destroyed without affecting root), and displaying alarm function (border cells potentially sensing pathogens and signaling to root triggering defensive responses). Border cells in rhizosphere includes= showing spatial distribution (border cells accumulating around root tip creating cellular halo in soil), exhibiting temporal dynamics (border cell population continuously renewed through production and lost through death creating dynamic population), demonstrating interactions (border cells interacting with soil particles, microbes, and chemical environment), revealing rhizosphere modification (border cells and mucilage affecting local soil chemistry, structure, and biology), and displaying carbon input (dead border cells providing readily available organic carbon to rhizosphere microbes). Border cells species variation includes= showing production differences (border cell numbers varying from sparse 10,000 per day in others), exhibiting structural variation (border cell and mucilage characteristics varying among species), demonstrating functional diversity (border cell roles potentially varying with species and ecological niche), revealing evolutionary patterns (border cell characteristics potentially reflecting adaptation to specific soil environments or pathogen pressures), and displaying grapevine characteristics (Vitis potentially showing moderate border cell production though specific data limited).
Border_Cells emphasizes detached free-living cap cells in rhizosphere and is distinguished from attached cap cells by focus on separated defensive cells. Includes Detaching_Border_Cells and Sloughing_Border_Cells. Critical for understanding cap cell detachment, continued viability, mucilage production, defensive functions, pathogen deterrence, rhizosphere modification, and the unique free-living cells creating defensive interface between root and soil.
Abgelöste Haubenzellen
Detached cap cells
Free-living root cells
Grenzzellen
Root cap slime cells
Schleimzellen
Hawes, M. C., et al. (2000). The role of root border cells in plant defense. Trends in Plant Science, 5(3), 128-133.
Border Cells
Boron content quality management in Napa Valley Cabernet Sauvignon addresses the fruit set problems affecting this variety on shallow, drought-stressed hillside soils, with pre-bloom foliar boron applications (Solubor at 1-2 kg/ha) improving pollen tube growth and fertilization success vineyards with adequate boron nutrition (must concentration 5-10 mg/L) produce well-filled clusters with uniform berry size essential for even ripening and optimal tannin maturity, while boron-deficient sites (below 3 mg/L) show the shot berry development and variable maturity that compromise premium Cabernet Sauvignon quality.
Boron_Content_Quality represents the chemical quality characterizing the concentration of boron (B) in grape berry tissue, capturing the essential micronutrient central to cell wall structure, sugar transport, and reproductive development in grapevines. Boron in grape juice typically ranges from 2-15 mg/L, with substantial variation reflecting cultivar, soil boron content, and the narrow range between deficiency and toxicity that characterizes boron nutrition. Unlike other micronutrients that function primarily as enzyme cofactors, boron's essential role involves structural functions in cell wall pectin cross-linking (through borate-diol complexes with apiose residues of rhamnogalacturonan II) and facilitation of sugar transport across membranes. Boron deficiency severely affects reproductive development, causing poor pollen germination, reduced fruit set, shot berry development, and straggly, incomplete clusters with yield losses exceeding 50% in severe cases. Deficiency symptoms also include shortened, thickened internodes and necrotic shoot tips. Boron availability decreases in dry conditions because mass flow with soil water is the primary uptake mechanism, creating drought-exacerbated deficiency even in soils with adequate total boron. Sandy, low-organic matter soils are prone to boron deficiency through leaching. Conversely, boron toxicity occurs in some arid regions where irrigation water contains elevated boron or where saline soils accumulate boron, producing characteristic marginal leaf burn that progresses inward. In wine production, boron is not associated with stability or quality problems at normal concentrations, though its content reflects fruit quality through the correlation with successful fruit set and normal berry development. Boron analysis employs colorimetric methods (azomethine-H), ICP-OES, or specialized ion-selective electrodes.
Boron_Content_Quality addresses a micronutrient with unique chemistry (forming covalent borate-diol complexes rather than ionic bonds) and narrow adequacy range between deficiency and toxicity. The reproductive function of boron makes fruit set particularly sensitive to deficiency, with yield and quality implications exceeding vegetative effects. The drought interaction with boron availability creates practical management challenges in dry-farmed or deficit-irrigated viticulture. Boron does not affect wine stability or chemistry, making its quality relevance indirect through effects on fruit development and cluster architecture.
B content
B-Gehalt
Beerenborqualität
Borgehalt
Borkonzentration
berry boron quality
boron concentration
boron level
O'Neill MA et al. (2004) Rhamnogalacturonan II: Structure and Function of a Borate Cross-Linked Cell Wall Pectic Polysaccharide. Annual Review of Plant Biology 55: 109-139 Christensen P (1986) Boron Application in Vineyards. California Agriculture 40(5-6): 17-18
Boron Content Quality
Boron deficiency symptom quality diagnosis in Marlborough Sauvignon Blanc identified B limitation as the cause of severe fruit set failure and cluster disorders in a high-rainfall vineyard on free-draining alluvial soils. Pre-bloom assessment revealed subtle boron deficiency symptom quality with shortened internodes on vigorous shoots, slight thickening and brittleness of young stems, and mild interveinal chlorosis on the youngest leaves, symptoms that had been attributed to vigor management rather than nutritional limitation. Post-fruit-set evaluation revealed dramatic boron deficiency symptom quality with 45% of clusters showing the "hen and chicken" millerandage pattern, 20% of clusters showing complete coulure (shot berries comprising greater than 80% of the cluster), and rachis tissue displaying internal browning and brittleness indicating vascular dysfunction. Tissue analysis confirmed boron deficiency symptom quality diagnosis with bloom petiole B at only 18 ppm (severely deficient, threshold 35 ppm), while soil B was at 0.25 ppm hot-water extractable (deficient, threshold 0.5 ppm), consistent with the leaching-prone sandy loam soil receiving 1,200 mm annual rainfall. The historical yield records showed progressive boron deficiency symptom quality impact with yields declining from 12 t/ha to 6 t/ha over five seasons as soil boron reserves depleted. Corrective management with soil-applied sodium borate at 15 kg/ha plus pre-bloom foliar B application at 0.2% Solubor resolved boron deficiency symptom quality within one season, with fruit set returning to normal (less than 5% millerandage) and yields recovering to 11 t/ha by the second season post-correction.
Boron deficiency symptom quality is a micronutrient deficiency symptom quality that inheres in a grapevine organism and represents the characteristic visual symptoms and physiological disorders resulting from inadequate boron availability for cell wall synthesis, pollen tube growth, and sugar transport, including poor fruit set leading to millerandage and coulure (hen and chicken clusters), necrosis of shoot tips and growing points, shortened internodes with zigzag shoot growth, corky bark development, and internal necrosis of cluster rachis, with boron deficiency being particularly significant for reproductive development where even marginal deficiency can cause substantial yield losses through impaired fertilization despite minimal vegetative symptoms. This quality captures the grapevine response to boron limitation, recognizing that boron's unique role in cell wall pectin cross-linking and its critical function in pollen germination and tube elongation make reproductive tissues highly sensitive to boron status, with fruit set often the first process affected by declining boron availability. The ontological significance of boron deficiency symptom quality lies in its substantial impact on fruit production even when vegetative symptoms are subtle, creating situations where yield losses may occur before obvious foliar symptoms alert the grower to boron limitation, making proactive tissue testing particularly valuable for boron management. The biological basis of boron deficiency symptoms encompasses the structural role of boron in cross-linking rhamnogalacturonan II (RG-II) pectin polymers essential for cell wall integrity, the requirement for boron in pollen tube growth where rapid cell wall synthesis during tube elongation creates high boron demand, and the function of boron in sugar transport through the phloem where boron-sugar complexes facilitate translocation. The phloem immobility of boron in most plant species including grapevine means that deficiency symptoms appear on youngest tissues and reproductive structures that cannot acquire boron through redistribution from older organs, with shoot tips and flowers particularly vulnerable during periods of rapid growth when boron demand exceeds root uptake capacity. The soil conditions predisposing to boron deficiency include sandy soils with low boron retention, high-rainfall areas where boron is leached, and soils with very high pH where boron availability decreases, though boron toxicity can also occur in arid regions with boron-containing groundwater, making boron management require attention to both deficiency and excess.
Boron_Deficiency_Symptom_Quality is_a Micronutrient_Deficiency_Symptom_Quality, serving as parent class for Hen_And_Chicken_Quality and Poor_Fruit_Set_Quality_(B) which address specific boron deficiency manifestations. This class addresses the overall boron deficiency syndrome while child classes capture particular symptom types. Reproductive development (pollen, fruit set) is highly B-sensitive, often affected before vegetative symptoms. Petiole B threshold ~35 ppm at bloom soil B threshold ~0.5 ppm (hot-water extractable). Phloem-immobile, symptoms on youngest tissues and reproductive structures. High rainfall + sandy soils = high B deficiency risk due to leaching.
B deficiency symptoms
B-Mangel-Zeichen
Bormangel-Symptome
Borstress-Indikatoren
Niedrig-Bor-Symptome
boron limitation signs
boron stress indicators
low boron symptoms
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication Brown PH et al. (2002) Boron in plant biology. Plant Biology 4:205-223
Boron Deficiency Symptom Quality
Boron toxicity quality assessment in Coachella Valley table grapes documented the characteristic symptoms and progressive damage resulting from irrigation with boron-rich groundwater containing 2.5 ppm B versus the recommended maximum of 0.5 ppm. Visual boron toxicity quality evaluation revealed the distinctive symptom pattern with interveinal necrotic patches developing between major veins, brown necrotic margins extending 1-3 cm from leaf edges, and tip burn creating pointed necrotic areas at the apex of leaf lobes, symptoms concentrated on older basal leaves while youngest leaves showed minimal damage. Severity scoring of boron toxicity quality on a 0-5 scale showed gradient from basal (score 4.2, severe interveinal and marginal necrosis) through mid-canopy (score 2.5, marginal burn only) to apical leaves (score 0.8, trace marginal discoloration), confirming the accumulation-driven pattern where longer transpiration exposure produced greater damage. Tissue analysis quantified boron toxicity quality with blade B concentrations of 320 ppm in severely symptomatic basal leaves (severely toxic, threshold 80 ppm), 145 ppm in moderately affected mid-canopy leaves, and 55 ppm in minimally affected young leaves still in the adequate range. Canopy position effects demonstrated boron toxicity quality relationship with transpiration, where south-facing exterior leaves showed 40% higher B accumulation and correspondingly more severe symptoms than shaded interior leaves at equivalent canopy positions. Yield and quality impact assessment showed boron toxicity quality causing 30% reduction in harvestable yield through early defoliation reducing photosynthetic capacity and direct fruit damage from B accumulation, with table grape pack-out reduced from 85% to 55% due to visible berry damage and reduced sugar accumulation.
Boron toxicity quality is a nutrient toxicity symptom quality that inheres in grapevine leaves and represents the characteristic damage pattern resulting from boron accumulation to levels exceeding the narrow range between adequacy and toxicity, with symptoms appearing as interveinal necrosis progressing from leaf margins inward, marginal and tip burn, and eventually extensive leaf necrosis and premature defoliation, reflecting boron's mobility in the transpiration stream that concentrates it at leaf margins and tips where water evaporates. This quality captures the grapevine response to boron excess, recognizing that grapevine is among the more boron-sensitive crop species with a narrow margin between the deficiency threshold (approximately 25-30 ppm in tissue) and toxicity threshold (approximately 80-100 ppm), making boron management challenging in regions where irrigation water or soil contains elevated boron. The ontological significance of boron toxicity quality lies in its indication of irrigation water quality problems or soil boron accumulation requiring management attention, and in its potential for substantial vine damage in arid and semi-arid regions where boron-containing groundwater or recycled water is used for irrigation. The biological basis of boron toxicity symptoms encompasses boron's mobility in the xylem where it moves with the transpiration stream and accumulates at sites of water loss, the direct cellular damage from high boron concentrations including membrane disruption and enzyme inhibition, and the interference with calcium metabolism that contributes to cell wall and membrane dysfunction. The symptom distribution pattern shows accumulation-driven damage at leaf margins and interveinal areas where transpiration concentrates boron, with older leaves showing more severe symptoms due to longer accumulation time, and with sun-exposed leaves experiencing more rapid symptom development due to higher transpiration rates. The threshold for toxicity varies with cultivar and rootstock, with some rootstocks providing partial exclusion of boron from xylem transport, offering a management option for sites with moderate boron excess.
Boron_Toxicity_Quality is_a Nutrient_Toxicity_Symptom_Quality, representing the characteristic marginal and interveinal necrosis from B accumulation via transpiration stream. This class addresses the specific B toxicity syndrome. Grapevine is B-sensitive with narrow adequacy-toxicity range (30-80 ppm tissue). Transpiration-driven accumulation produces marginal/interveinal concentration. Older leaves and sun-exposed positions show more severe symptoms. Tissue B threshold for toxicity ~80-100 ppm (varies by cultivar). Some rootstocks provide partial B exclusion. Arid regions with B-rich groundwater are primary risk areas.
B toxicity
B-Überschuss-Symptome
Bor-Exzess-Schaden
Bortoxizität
Borverbrennung
boron burn
boron excess symptoms
high boron damage
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication Nable RO et al. (1997) Boron toxicity. Plant and Soil 193:181-198
Boron Toxicity Quality
Boron uptake efficiency varies 2-3 fold among rootstocks, with 1103 Paulsen and 140 Ruggeri showing high efficiency (maintaining petiole B above 35 ppm) compared to 3309C and Riparia Gloire (petiole B 20-30 ppm) on low-B soils. Boron deficiency severely impairs reproductive development, reducing pollen germination by 40-70% and fruit set by 30-60% when petiole B falls below 25 ppm at bloom. Symptoms include shortened internodes, distorted leaves, cluster necrosis (hen and chickens from failed pollination), and surface corking on berries. Boron mobility in phloem is limited, requiring continuous uptake for developing tissues. The narrow range between deficiency (80 ppm) makes B nutrition management challenging in some regions.
A micronutrient uptake disposition that is the inherited capacity of a Vitis vinifera plant to absorb boric acid (H₃BO₃) and borate ions from the soil solution through aquaporin-type channels (NIP subfamily) and borate transporters (BOR family), with genotype-specific efficiency affecting cell wall structure, pollen development, and sugar translocation.
Boron uptake disposition is critical for reproductive success due to the essential role of B in pollen tube growth and ovule development. The element cross-links cell wall pectin through borate-diol ester bonds, affecting wall structure and extensibility. Boron deficiency is widespread in sandy, leached soils and in regions with low-B irrigation water. Unlike most micronutrients, B uptake occurs primarily as undissociated boric acid at typical soil pH values, through aquaporin channels rather than ion transporters. Foliar B application before bloom effectively prevents deficiency-induced coulure and millerandage.
B uptake capacity
B-Akquisition
Boraneignungsvermögen
Boraufnahme
boron acquisition
boron foraging trait
Keller M et al. (2015) Boron nutrition of grapevines. Acta Horticulturae 1115:79-84 Scott LE (1941) Effects of boron nutrition on grapevines. Botanical Gazette 102:726-736
Boron Uptake Disposition
Boron uptake efficiency assessment across vineyard blocks with varying soil characteristics reveals the site-specific acquisition patterns, with sandy loam block (low CEC, adequate drainage) showing marginal vine B status (leaf B 22 mg/kg, occasional shot berries) from leaching losses reducing soil B availability, clay block showing adequate B (leaf B 38 mg/kg, normal fruit set) from better B retention, and irrigated block with high-B water showing excess (leaf B 85 mg/kg, marginal leaf burn) from cumulative B accumulation, demonstrating the narrow optimal range requiring site-specific boron uptake efficiency assessment and management.
Boron uptake efficiency quality is a micronutrient uptake efficiency quality that represents the capacity of grapevine roots to acquire boron from soil, characterizing the effectiveness of borate absorption that determines vine boron status affecting cell wall structure, pollen tube growth, and sugar transport with critical implications for fruit set, berry development, and vine reproductive success. Within the BFO framework, this quality specifically addresses the boron component of micronutrient uptake, distinguishing boron acquisition from other trace elements while recognizing that boron has the narrowest range between deficiency and toxicity of any essential element, making precise management of boron uptake and status particularly important in viticulture. The ontological significance lies in boron uptake efficiency determining access to this essential element required for cell wall pectin cross-linking, membrane function, and reproductive development, where boron deficiency causes poor fruit set (pollen tube growth failure), shot berries (uneven berry development), and hen-and-chicken clusters that severely reduce yield and quality, while excess boron causes marginal leaf necrosis and reduced growth. The mechanistic basis for boron uptake involves both passive diffusion of undissociated boric acid (B(OH)₃) across membranes at adequate soil concentrations and active transport through BOR (Boron transporter) family proteins under deficiency, with aquaporin-like NIP channels also contributing to boric acid permeation, creating multiple pathways for boron entry depending on soil concentration. The soil factors affecting boron availability include soil texture (sandy soils prone to boron leaching), organic matter (providing mineralizable boron), rainfall and irrigation (leaching reducing available boron), and soil pH (availability declining at high pH though less dramatically than for iron or zinc), with boron deficiency more common in high-rainfall regions and sandy soils. The critical role of boron in reproduction makes pre-bloom boron status particularly important, as boron is required for pollen germination and pollen tube elongation such that deficiency at flowering causes fruit set failure even if vegetative symptoms are absent, emphasizing the timing importance of boron nutrition management. Understanding boron uptake efficiency quality enables assessment of boron deficiency and toxicity risk, timing of boron applications relative to flowering, and management of the narrow optimal boron range for grapevine productivity.
Boron_Uptake_Efficiency_Quality addresses the acquisition of this element with the narrowest deficiency-toxicity range among essential nutrients. This quality critically affects reproductive development through pollen tube function. The class supports boron nutrition management through uptake efficiency characterization balancing deficiency and toxicity risks.
B uptake efficiency
BOR activity
BOR-Aktivität
Borattransport
Boraufnahmeeffizienz
borate transport
Tanaka M, Fujiwara T (2008) Physiological roles and transport mechanisms of boron: perspectives from plants. Pflügers Archiv 456(4): 671-677 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 197-245
Boron Uptake Efficiency Quality
Multi-species evaluation of Botryosphaeria_Resistance_Disposition at Stellenbosch University, South Africa, compared host responses to four Botryosphaeriaceae species (Neofusicoccum australe, N. parvum, Diplodia seriata, Lasiodiplodia theobromae) across 12 cultivars using standardized green shoot inoculation assays. At 8 weeks post-inoculation, lesion lengths were measured and pathogens re-isolated to confirm Koch's postulates. Substantial cultivar × pathogen interactions were observed. 'Pinotage' showed broad Botryosphaeria_Resistance_Disposition with mean lesion lengths of 18-25 mm across all four species. 'Sauvignon Blanc' was broadly susceptible (lesions 65-95 mm). Interestingly, 'Cabernet Sauvignon' exhibited species-specific responses, showing strong resistance to D. seriata (22 mm) but high susceptibility to N. parvum (78 mm), demonstrating that Botryosphaeria_Resistance_Disposition is not uniform across the pathogen family. Field validation in a 15-year-old cultivar trial confirmed experimental rankings, with 'Pinotage' vines showing 70% lower incidence of black dead arm symptoms than 'Sauvignon Blanc'. Histological examination revealed that 'Pinotage' produces denser wound-associated tyloses and accumulates higher concentrations of antimicrobial phenolics in response to Botryosphaeriaceae colonization.
A Botryosphaeria_Resistance_Disposition is a Trunk_Disease_Resistance_Disposition that inheres in a Vitis vinifera plant and is realized through defense processes that limit infection, colonization, and pathogenesis by fungi in the family Botryosphaeriaceae, a diverse group causing Botryosphaeria dieback and related canker diseases of grapevine. This disposition represents the capacity to resist infection by multiple Botryosphaeriaceae species including Diplodia seriata, Lasiodiplodia theobromae, Neofusicoccum parvum, Botryosphaeria dothidea, and related taxa that cause wood necrosis, canker formation, graft union failure, and cordon dieback. Botryosphaeriaceae fungi are ubiquitous in vineyard environments, existing as endophytes in healthy tissue and transitioning to pathogenic behavior when host stress compromises defense responses. These opportunistic pathogens infect through pruning wounds and other injuries, producing wedge-shaped or sectorial necrosis in wood, black or brown vascular streaking, and occasionally shoot dieback and bud necrosis. Resistance mechanisms include efficient wound closure limiting infection opportunities, recognition and containment of endophytic populations before pathogenic transition, compartmentalization of necrotic tissues, and tolerance to botryosphaeriaceae phytotoxins (mellein, botryodiplodin). Botryosphaeriaceae diversity creates challenges for resistance breeding, as cultivars may differ in susceptibility to individual species. Generally, cultivars susceptible to esca also show heightened Botryosphaeria susceptibility, suggesting shared resistance mechanisms against trunk-colonizing fungi, but species-specific resistance components also exist.
Botryosphaeria_Resistance_Disposition is_a Trunk_Disease_Resistance_Disposition. The Botryosphaeriaceae family includes numerous species pathogenic on grapevine, with regional variation in predominant species and cultivar × species interactions creating complexity for resistance evaluation. Many Botryosphaeriaceae exist as latent endophytes, with disease expression triggered by host stress, complicating the distinction between resistance (preventing infection/colonization) and tolerance (limiting disease expression despite colonization). Shared resistance mechanisms with esca-associated fungi suggest common trunk disease resistance factors, but species-specific components also contribute to overall Botryosphaeria_Resistance_Disposition.
Botryosphaeria dieback tolerance
Botryosphaeria-Resistenz-Disposition
Botryosphaeria-Toleranz
Botryosphaeriaceae resistance
Diplodia-Resistenz
black dead arm resistance
Úrbez-Torres JR (2011) The status of Botryosphaeriaceae species infecting grapevines, Phytopathologia Mediterranea 50:S5-S45 Pitt WM et al. (2013) Evaluation of fungicides for the management of Botryosphaeria canker of grapevines, Plant Disease 97:1473-1482 Billones-Baaijens R, Savocchia S (2019) A review of Botryosphaeriaceae species associated with grapevine trunk diseases in Australia and New Zealand, Australasian Plant Pathology 48:3-18
Botryosphaeria Resistance Disposition
Comparative field evaluation of Botrytis_Resistance_Disposition at Geisenheim University assessed grey mold incidence across 47 cultivars over five harvest seasons with varying autumn precipitation. The susceptible cultivar 'Gewürztraminer' (compact clusters, thin skin, high sugar) exhibited mean cluster infection of 45-65% in wet years, with individual berries showing characteristic grey sporulation and entire clusters collapsing to "slip-skin" consistency. In contrast, 'Regent' demonstrated strong Botrytis_Resistance_Disposition with mean infection of 8-15% under identical conditions, attributable to its loose cluster architecture (OIV 204 rating 3), thick berry skin (mean 285 μm versus 180 μm in 'Gewürztraminer'), and elevated constitutive stilbene content. Intermediate cultivars such as 'Riesling' (mean infection 25-35%) showed characteristic partial resistance with infection concentrated at berry contact points and wound sites. No cultivar achieved complete immunity, reflecting the absence of major gene resistance and the pathogen's capacity to exploit any compromised tissue, but the 4-5 fold difference in infection severity between resistant and susceptible cultivars demonstrates the practical significance of selecting for enhanced Botrytis resistance dispositions.
A Botrytis_Resistance_Disposition is a Fungal_Resistance_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through defense processes upon encounter with the necrotrophic ascomycete Botrytis cinerea, the causal agent of grey mold (bunch rot) and, under specific conditions, noble rot. This disposition represents the capacity to limit B. cinerea infection, colonization, and spread through diverse mechanisms operating at pre-penetration, penetration, and post-penetration stages. Unlike biotrophic pathogens such as powdery and downy mildew that require living host tissue, B. cinerea kills host cells and feeds on dead tissue, fundamentally altering the host-pathogen interaction dynamics. Botrytis resistance dispositions encompass both constitutive and induced mechanisms, including physical barriers (cuticle thickness and composition, berry skin toughness), preformed and induced antimicrobial compounds (particularly stilbene phytoalexins such as resveratrol and pterostilbene), cluster architectural traits that reduce humidity and berry contact, and active defense responses including oxidative burst and cell wall reinforcement at wound sites. The complex, quantitative nature of Botrytis resistance reflects the pathogen's opportunistic infection strategy exploiting wounds, natural openings, and senescing tissue rather than specific recognition-based immunity. No major gene resistance to B. cinerea has been identified in Vitis, making breeding for Botrytis resistance dependent on accumulating multiple partial resistance factors. Botrytis resistance is economically critical as grey mold causes significant crop losses in humid climates and produces off-flavors that severely compromise wine quality.
Botrytis_Resistance_Disposition is_a Fungal_Resistance_Disposition. Unlike downy and powdery mildew resistances which involve major gene (R gene) mediated immunity, Botrytis resistance is exclusively quantitative with no identified major resistance loci. Child classes are organized by mechanism: Cluster_Architecture_Based_Resistance_Disposition (structural), Cuticle_Based_Resistance_Disposition (physical barrier), and Phytoalexin_Based_Botrytis_Resistance_Disposition (chemical defense). Noble_Rot_Susceptibility_Disposition represents the special case where controlled B. cinerea infection is desirable for certain wine styles.
Botrytis cinerea resistance
Botrytis-Resistenz-Disposition
Grauschimmel-Resistenz
Traubenfäule-Resistenz
bunch rot resistance
grey mold resistance disposition
Elmer PAG, Michailides TJ (2007) Epidemiology of Botrytis cinerea in orchard and vine crops, Botrytis: Biology, Pathology and Control, Springer Deytieux-Belleau C et al. (2009) Berry skin development in Vitis vinifera: morphological and biochemical events linked to Botrytis resistance, Journal of Experimental Botany 60:3919-3930 Bézier A et al. (2002) Carbon source-dependent expression of a resveratrol synthase gene of grapevine, Physiological and Molecular Plant Pathology 61:179-186
Botrytis Resistance Disposition
Botrytis symptom quality assessment in Sauternes Semillon illustrated the management challenge of distinguishing destructive bunch rot from desirable noble rot development. Early Botrytis symptom quality appeared post-véraison as isolated berry infections showing brown discoloration and softening at wound sites or where berries contacted each other within tight clusters. Under humid September conditions, aggressive Botrytis symptom quality developed with rapid expansion of gray sporulating mycelium across cluster surfaces, infected berries collapsing to soft rot, and the "slip-skin" symptom where gentle pressure separated infected skin from underlying pulp. Weather shift to warm, dry conditions in early October transformed the Botrytis symptom quality presentation, with existing infections ceasing sporulation, infected berries beginning to shrivel and concentrate rather than rot, and the characteristic golden-purple "pourri plein" appearance developing as noble rot established. Monitoring distinguished Botrytis symptom quality stages, as "pourri gris" (gray rot, destructive), "pourri plein" (full noble rot, optimal), and "pourri sec" (dried rot, over-developed), with selective harvest targeting the pourri plein stage where berry Brix reached 35-40° with characteristic noble rot aromatics. The Botrytis symptom quality assessment enabled tri-harvest strategy, with destructively rotted clusters removed first, optimal noble rot clusters harvested selectively over 3-4 passages, and over-dried clusters left for final late harvest, maximizing the proportion of production achieving the distinctive noble rot quality premium.
Botrytis symptom quality is a fungal infection indicator quality that inheres in grapevine organs infected by the necrotrophic fungus Botrytis cinerea and represents the characteristic disease manifestations including the gray fuzzy sporulating mycelium on infected tissues, the brown soft rot of infected berries and rachises, the "slip-skin" phenomenon where infected berry skin separates from pulp, and the distinctive symptoms of noble rot under specific conditions where infected berries develop the concentrated, honeyed characteristics valued for dessert wine production rather than the destructive bunch rot more commonly observed. This quality captures the diagnostic features of the most economically significant bunch rot pathogen worldwide, recognizing that B. cinerea can cause either devastating crop losses (gray mold, bunch rot) or contribute to premium wine production (noble rot, pourriture noble, Edelfäule) depending on environmental conditions and management, creating a uniquely dual relationship between this pathogen and viticulture. The ontological significance of Botrytis symptom quality lies in its role enabling disease identification, severity assessment, and critically the distinction between destructive rot and potentially beneficial noble rot, with symptom characteristics and progression determining whether infected fruit should be culled, harvested immediately, or allowed to develop for specialized sweet wine production. The biological basis of Botrytis symptoms encompasses the aggressive colonization of host tissues by B. cinerea through secretion of cell-wall-degrading enzymes and phytotoxins that kill host cells in advance of hyphal growth, the rapid development of gray conidiophores producing enormous quantities of airborne conidia under humid conditions, and the tissue maceration causing the soft rot characteristic of bunch rot. The noble rot syndrome develops when warm dry conditions following initial infection limit sporulation and allow slow dehydration of infected berries, concentrating sugars and producing the characteristic Botrytis-derived flavor compounds (sotolon, phenylacetaldehyde) valued in Sauternes, Tokaji, and Trockenbeerenauslese wines. The environmental switching between bunch rot and noble rot outcomes depends primarily on humidity during infection development, with high humidity favoring aggressive sporulation and rot spread while alternating humid-dry conditions favor the noble rot progression.
Botrytis_Symptom_Quality_(Botrytis_cinerea) is_a Fungal_Infection_Indicator_Quality, representing the diagnostic symptoms of Botrytis cinerea infection in both destructive (bunch rot) and beneficial (noble rot) forms. The gray sporulating mycelium is pathognomonic. Environmental switching between rot and noble rot outcomes depends on humidity during infection development. The pourri gris/pourri plein/pourri sec progression indicates disease stage for sweet wine harvest decisions. The "slip-skin" symptom aids field diagnosis. Unique dual relationship with viticulture as both devastating pathogen and quality enhancer.
Botrytis rot indicators
Botrytis-Symptome
Edelfäule-Symptome
Grauschimmel-Symptome
Traubenfäule-Zeichen
bunch rot symptoms
gray mold symptoms
noble rot symptoms
Elmer PAG, Michailides TJ (2007) Epidemiology of Botrytis cinerea in orchard and vine crops. In: Botrytis: Biology, Pathology and Control. Springer, pp. 243-272 Ribéreau-Gayon P et al. (2006) Handbook of Enology, Volume 1: The Microbiology of Wine and Vinifications. Wiley Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests, 2nd Edition. APS Press
Botrytis Symptom Quality (Botrytis cinerea)
In Irsai Olivér berries in Mátra, Hungary during BBCH 83-89, bound aroma accumulation increases glycosidically-bound monoterpenes from 2.2 to 9.8 mg/L over 42 days, with accumulation rates peaking at 0.25 mg/L per day during weeks 3-4 post-veraison, bound linalool reaching 3.2 mg/L, bound geraniol 2.1 mg/L, and bound nerol 1.4 mg/L, establishing aromatic potential 7.5-fold greater than free volatile content of 1.3 mg/L at harvest under continental conditions with warm days of 28°C and cool nights of 14°C.
Bound aroma accumulation describes the progressive increase in glycosidically-conjugated and otherwise bound aromatic compound concentrations within Vitis vinifera berry tissues during ripening, wherein continued glycosyltransferase activity converts newly synthesized free volatiles to their conjugated forms while previously synthesized glycosides remain stable in vacuolar storage, producing the monotonic increase in bound aromatic content that establishes the aromatic potential of harvested grapes for subsequent release during winemaking processes. This process reflects the net balance between ongoing glycosylation adding compounds to the bound pool and potential hydrolysis or degradation removing compounds, with successful accumulation requiring that conjugation rates substantially exceed any losses to maintain the positive accumulation trajectory characteristic of normal ripening progression in aromatic grape varieties. The quantitative aspects of bound aroma accumulation show variety-specific patterns, with aromatic varieties such as Muscat and Gewürztraminer accumulating bound monoterpene pools of 5-15 mg/L representing 85-95% of total terpenoid content, while neutral varieties such as Chardonnay and Pinot Noir accumulate substantially lower bound pools of 0.3-1.0 mg/L reflecting reduced glycosyltransferase expression and lower terpenoid biosynthetic capacity. Temporal dynamics of bound aroma accumulation demonstrate progressive increase throughout the post-veraison period, with accumulation rates typically highest during mid-ripening when both terpenoid biosynthesis and glycosyltransferase activities reach peak levels, followed by declining rates during late ripening as substrate availability decreases and enzyme activities wane approaching harvest maturity. The vacuolar sequestration mechanism supporting bound aroma accumulation involves active transport of glycosidic conjugates across the tonoplast membrane through ABC transporters and related systems, establishing concentration gradients with vacuolar glycoside levels of 2-10 mM representing 50-100 fold enrichment compared to cytosolic pools, ensuring stable storage protected from glycosidase activities maintaining precursor integrity throughout ripening and post-harvest handling. Environmental factors influence bound aroma accumulation through temperature effects on both terpenoid biosynthesis providing substrates and glycosyltransferase activities enabling conjugation, with moderate temperatures of 20-28°C generally supporting optimal accumulation while cool conditions may slow both processes and warm conditions may accelerate substrate volatilization reducing material available for glycosylation. The practical significance of bound aroma accumulation for winemaking reflects the requirement for precursor hydrolysis to release aromatic aglycones, with winemaker decisions regarding enzyme additions, yeast selection, and contact time determining the extent to which accumulated precursor pools are converted to sensory-active free volatiles in finished wines.
Bound_Aroma_Accumulation is subclass of Aroma_Precursor_Formation representing quantitative increase in conjugated aromatics. Distinguishes from Glycosidic_Precursor_Synthesis which describes reaction mechanism. Sibling to Glycosidic_Precursor_Synthesis and Precursor_Pool_Building. Bound forms typically 85-95% of total monoterpenoids. Accumulation continues throughout ripening. Determines aromatic potential for winemaking release.
Glycosid-Aroma-Anreicherung
Vorläufer-Aroma-Akkumulation
bound volatile increase
conjugated aroma build-up
gebundene Aroma-Akkumulation
glycoside aroma accumulation
konjugierte Aroma-Aufbau
precursor aroma accumulation
Wilson B et al (1984) Determination of free and bound monoterpenes in Muscat grapes. American Journal of Enology and Viticulture 35: 95-100 Gunata Z et al (1985) The aroma of grapes. Journal of Chromatography 331: 83-90 Keller M (2020) The Science of Grapevines 3rd Ed pp 295-305
Bound Aroma Accumulation
Boundary layer conductance assessment comparing exterior and interior canopy positions in dense Merlot reveals the microclimate decoupling in sheltered positions, with exterior leaves showing gb of 4.5 mol/m²/s (well-coupled to atmosphere, leaf temperature within 2°C of air), while interior leaves show gb of 0.8 mol/m²/s (thick boundary layer from calm conditions, leaf temperature 8°C above air, humidity elevated), with the reduced coupling in interior positions limiting effective gas exchange despite adequate stomatal conductance and creating heat stress conditions that differ markedly from atmospheric conditions, informing canopy management to improve air circulation.
Boundary layer conductance quality is a transpiration rate quality that represents the ease of gas diffusion through the layer of still air adjacent to leaf surfaces, characterizing the aerodynamic component of total leaf conductance that affects both water vapor loss and CO₂ uptake, particularly significant under low wind conditions when the boundary layer becomes a substantial resistance in series with stomatal conductance. Within the BFO framework, this quality specifically addresses the aerodynamic dimension of gas exchange, distinguishing boundary layer conductance from stomatal conductance while recognizing that these resistances operate in series, meaning that low boundary layer conductance can limit gas exchange even when stomata are fully open, decoupling leaves from atmospheric conditions and creating distinct microenvironments within dense canopies. The ontological significance lies in boundary layer conductance determining the coupling between leaf and atmosphere that affects both the effectiveness of stomatal regulation and the environmental conditions experienced at the leaf surface, where thick boundary layers insulate leaves from bulk atmospheric conditions, reducing transpiration and creating elevated leaf temperatures that affect photosynthesis, respiration, and heat stress vulnerability. The physical basis for boundary layer conductance involves the thickness of the unstirred air layer adjacent to leaf surfaces through which gases must diffuse, with boundary layer thickness determined by leaf size (larger leaves having thicker layers), wind speed (higher wind reducing thickness), and leaf surface characteristics (pubescence and roughness affecting air flow), and with conductance inversely proportional to thickness. The quantitative ranges for boundary layer conductance depend strongly on conditions, with calm conditions producing low gb (0.5-2 mol/m²/s) that substantially limits gas exchange, while windy conditions increase gb (5-10+ mol/m²/s) making stomatal conductance the dominant limitation, with intermediate conditions creating variable coupling between leaves and atmosphere. The canopy architecture implications of boundary layer conductance are significant, as dense canopies with limited air penetration create thick boundary layers on interior leaves, reducing their gas exchange efficiency and creating warm, humid microclimates that differ substantially from exterior canopy conditions. Understanding boundary layer conductance quality enables interpretation of canopy microclimate effects on gas exchange, assessment of heat stress risk under calm conditions, and canopy management optimization for appropriate aerodynamic coupling.
Boundary_Layer_Conductance_Quality addresses the aerodynamic resistance component affecting leaf-atmosphere coupling. This quality determines the effectiveness of stomatal regulation and microclimate at leaf surfaces. The class supports canopy management and heat stress assessment through boundary layer conductance characterization.
Blattgrenzschicht
Grenzschichtleitfähigkeit
aerodynamic conductance
aerodynamische Leitfähigkeit
air layer resistance
gb-Wert
leaf boundary layer
Jones HG (2014) Plants and Microclimate: A Quantitative Approach to Environmental Plant Physiology, 3rd Ed. Cambridge University Press, Cambridge, pp. 55-95 Daudet FA et al. (1999) Wind speed and leaf boundary layer conductance variation within tree crown. Agricultural and Forest Meteorology 97(3): 171-185
Boundary Layer Conductance Quality
Pinot Noir grafted on Riparia Gloire showing boundary transition zone on primary root tip, boundary visible at 0.28 mm from apex as distinct histological demarcation in longitudinal sections separating columella cap cells from corpus meristem initials, molecular marker analysis revealing sharp expression boundary with cap-specific WOX5 expression columella side and meristem-specific SCR expression corpus side, cell wall staining showing differential pectin versus cellulose content across boundary, boundary marking fundamental organizational separation between protective disposable cap and permanent growing meristem maintaining distinct developmental programs and cellular properties
Boundary transition zone is root cap transition zone emphasizing structural and organizational boundary marking separation between root cap and root meristem, characterized by distinct histological demarcation visible in microscopic sections separating columella and lateral cap cells from corpus meristem initials, showing differential gene expression patterns with cap-specific and meristem-specific genes showing expression boundaries at transition zone, demonstrating structural discontinuity where cap cells showing different wall properties, cytoskeletal organization, and cellular behaviors than meristem cells with transition occurring across narrow boundary, and representing fundamental organizational feature of closed-type root meristem where cap and corpus maintain separate identities and developmental programs despite cellular proximity and shared developmental origin, with boundary transition zone showing molecular markers differentially expressed across boundary, mechanical property differences between compliant sloughable cap and rigid persistent meristem, and functional separation where cap cells programmed for eventual death and loss contrast with meristem cells maintaining unlimited proliferative capacity, and demonstrating that boundary maintenance is essential for proper apex function preventing inappropriate fate mixing that would compromise both protective and growth functions. Boundary transition zone structural characteristics includes= showing histological demarcation (clear boundary visible in microscopic sections distinguishing cap from meristem tissues), exhibiting cell morphology change (cap cells showing different size, shape, and organization than meristem cells with transition at boundary), demonstrating wall property differences (cell walls showing different composition, thickness, or properties across boundary), revealing cytoskeletal organization (cortical microtubules and actin filaments potentially organized differently in cap versus meristem with transition at boundary), and displaying discontinuity (abrupt rather than gradual change in cellular characteristics marking distinct boundary). Boundary transition zone molecular markers includes= showing differential gene expression (cap-specific genes expressed one side of boundary, meristem-specific genes other side), exhibiting transcription factor boundaries (key regulators showing sharp expression boundaries at transition zone), demonstrating protein localization (specific proteins enriched on cap or meristem side with transition at boundary), revealing boundary-specific markers (genes or proteins specifically expressed at boundary marking interface), and displaying molecular asymmetry (different molecular environments on cap versus meristem sides of boundary). Boundary transition zone functional separation includes= showing distinct developmental programs (cap cells following differentiation and death program versus meristem cells maintaining division and differentiation), exhibiting different cell fates (cap cells destined for eventual sloughing versus meristem cells producing root body tissues), demonstrating independent regulation (cap and meristem responding to different signals and regulators), revealing functional specialization (cap optimized for protection and sensing versus meristem for growth and tissue production), and displaying fate restriction (boundary preventing cap cells from joining meristem or vice versa maintaining organizational integrity). Boundary transition zone mechanical properties includes= showing differential stiffness (cap potentially softer more compliant than meristem facilitating sloughing), exhibiting adhesion differences (cap cells showing weaker intercellular adhesion allowing eventual separation versus strong meristem adhesion), demonstrating structural differences (cap cells potentially with more pectin-rich walls versus meristem with different wall composition), revealing mechanical discontinuity (abrupt change in mechanical properties at boundary potentially important for cap sloughing), and displaying force transmission (boundary potentially serving as mechanical interface transmitting forces between cap and meristem). Boundary transition zone maintenance mechanisms includes= showing active enforcement (molecular mechanisms actively maintaining boundary preventing drift or mixing), exhibiting mutual inhibition (cap and meristem factors potentially inhibiting each other at boundary creating sharp interface), demonstrating positional memory (cells maintaining positional identity relative to boundary through epigenetic or other mechanisms), revealing signaling feedback (boundary signals potentially reinforcing cap and meristem identities preventing inappropriate fate changes), and displaying developmental stability (boundary maintained throughout apex life demonstrating robust organizational mechanism). Boundary transition zone in apex organization includes= determining apex architecture (boundary location and characteristics defining overall apex structure), enabling functional separation (boundary allowing cap and meristem to function independently optimizing both protection and growth), demonstrating organizational importance (proper boundary maintenance essential for normal apex development and function), revealing pattern formation (boundary participating in overall apex patterning coordinating cap, meristem, and other apex regions), and displaying evolutionary significance (boundary transition zone organization showing conservation reflecting fundamental importance for root apex function). Boundary transition zone disruption effects includes= showing organizational defects (boundary disruption potentially causing cap-meristem mixing compromising both functions), exhibiting developmental abnormalities (improper boundary potentially leading to aberrant apex structure and growth), demonstrating functional impairment (boundary defects potentially reducing protective capacity and growth rate), revealing regeneration involvement (boundary reorganization potentially important for apex recovery after damage), and displaying mutation phenotypes (genetic mutations affecting boundary potentially showing visible apex defects documenting boundary importance). Boundary transition zone visualization includes= showing histological visibility (boundary often visible in standard microscopic sections through morphological differences), exhibiting marker staining (molecular markers highlighting boundary through differential expression or protein localization), demonstrating vital dyes (fluorescent dyes potentially revealing boundary through differential uptake or retention), revealing imaging techniques (confocal microscopy or other advanced imaging clearly showing boundary structure), and displaying cell tracking (lineage tracing potentially revealing boundary maintenance over time documenting cellular dynamics).
Boundary_Transition_Zone emphasizes structural demarcation and organizational separation and is distinguished from meristematic activity emphasis by focus on boundary maintenance. Critical for understanding apex organization, structural discontinuity, molecular boundaries, functional separation, mechanical properties, and the sharp organizational boundary separating root cap and meristem maintaining distinct identities essential for proper apex structure and function.
Cap-meristem demarcation
Grenz-Übergangszone
Organisatorische Schnittstelle
Organizational interface
Structural boundary zone
Strukturelle Grenzregion
Dolan, L., et al. (1993). Cellular organisation of the Arabidopsis thaliana root. Development, 119(1), 71-84.
Boundary Transition Zone
A branch arm water shoot on a Gobelet-trained vine of Vitis vinifera cv. Grenache in the southern Rhône Valley, France, emerging from the junction point where three permanent arms diverge from the central head at 45 cm trunk height, developing in response to severe reduction pruning that removed two of the five original arms showing advanced Esca symptoms, growing 195 cm by September with 21 nodes, positioned at a favorable 45-degree angle relative to the remaining arms, leaves averaging 185 cm² area with good sun exposure, selected for retention and training as a replacement arm to restore the traditional five-arm Gobelet architecture over a three-season development period involving progressive crop load increase and lignification enhancement.
An arm water shoot originating from latent or adventitious buds on multi-branched permanent arm structures including secondary, tertiary, or higher-order framework branches in complex training systems such as Gobelet, bush vines, or multi-arm head-trained systems in Vitis vinifera, characterized by emergence from branch points, angles, or junction zones where multiple permanent structures meet, often showing enhanced vigor due to accumulated resources at branch intersections and vascular convergence zones. Branch arm water shoots develop preferentially at positions where pruning wounds, branch angles, or structural discontinuities create localized stress responses, where trunk diseases produce symptoms at branch junctions that compromise distal tissue function, or where severe reduction pruning removes entire branch sections and triggers compensatory growth from remaining framework. These shoots present both increased management complexity in multi-branched systems where framework geometry is already intricate, and strategic opportunities for framework reconstruction in aging bush vines or head-trained systems where progressive branch replacement maintains vine productivity while avoiding complete structural renewal. Branch arm water shoots in traditional training systems like Gobelet may be intentionally stimulated through selective branch removal to create new framework arms positioned for improved sun exposure, air circulation, or geometric balance in asymmetric vine structures.
Branch arm water shoots are distinguished from simple arm water shoots and cordon water shoots by origin at branch junction points or on multi-branched structures rather than on single linear cordons or arms. The complexity of multi-branched systems creates numerous potential positions for water shoot emergence, requiring systematic evaluation of each shoot's impact on overall framework geometry and balance. Branch arm water shoots at major junction points often show exceptional vigor due to convergence of vascular streams from multiple framework components, and may require more aggressive management through repeated removal or strategic retention with severe topping to moderate growth. In traditional bush vine or Gobelet systems, branch arm water shoots represent normal opportunities for progressive framework renewal without complete vine replacement.
Astgabelungstrieb
Verzweigungswassertrieb
Zweigarmwasserschoss
branch point sucker
framework branch sucker
junction water shoot
multi-arm water shoot
Based on Galet (2000) "General Viticulture" (English translation), Oenoplurimedia pp. 243-267 Champagnol (1984) "Éléments de physiologie de la vigne et de viticulture générale" Saint-Gely-du-Fesc, Imprimerie Déhan pp. 298-315 Carbonneau et al. (2015) "Traité de la Vigne= Physiologie, Terroir, Culture" 2nd Edition, Dunod pp. 267-289 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 366-388.
Branch Arm Water Shoot
A branch lateral petiole on an extremely deeply lobed mature leaf of an unusual Vitis vinifera Mourvèdre clone in Bandol, France, exhibiting distinct Y-shaped bifurcation at the junction where the main petiole meets the lamina base and divides into two lateral lobe supports, primary branch length of 2.4 cm before bifurcation with subsequent branches of 1.8 cm each supporting the two major lateral lobes, diameter of 2.2 mm at the primary section reducing to 1.5 mm in the branches, green coloration with slight red tinge, moderate pubescence, containing branched vascular strands visible as raised ridges showing actual petiolar vascular organization rather than simple vein divergence, providing meaningful mechanical support allowing the deeply dissected lateral lobes to function somewhat independently, occurring on approximately 25% of leaves on this extreme variant plant representing maximum lateral petiole expression, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing the most developed lateral petiolar structures observed in grapevine approaching compound leaf architecture.
A lateral petiole showing relatively well-developed branched or forked morphology creating distinct Y-shaped or T-shaped junction structures in Vitis vinifera, characterized by bifurcated or trifurcated stalk appearance at major vein junctions where main petiole transitions to lamina base, dimensions typically 1-3 cm length representing substantial development for lateral petiolar structures, occurrence in cultivars with extremely deep lobe dissection creating quasi-compound leaf architecture, anatomical structure including branched vascular traces and mechanical tissues creating actual forked petiolar support rather than simple vein divergence, and functional contribution providing meaningful additional mechanical support and vascular capacity for deeply dissected lobes approaching separation from main lamina. Branch lateral petioles represent maximum expression of lateral petiolar development in grapevine, approaching but not achieving true compound leaf architecture with independent leaflets on distinct petioles, occurring most commonly in cultivars or individual plants showing most extreme lobe dissection including some Mourvèdre clones, certain deeply dissected Muscat types, and occasional morphological variants in other cultivars, and serving as morphological indicators of extreme developmental variation within the simple leaf architectural constraints of Vitis vinifera.
Branch lateral petioles represent the most developed expression of lateral petiolar structures in Vitis vinifera, distinguished from simple lateral petioles by substantial branched morphology and from major veins by petiolar character. These structures remain rare even in cultivars showing tendencies toward deep lobe dissection, representing extreme developmental variants rather than typical features. The branched morphology creates quasi-compound leaf architecture with mechanically and somewhat vascularly independent lobe structures, approaching but not achieving true compound leaves with completely separate leaflets. Functional significance is greater than simple lateral petioles but still limited compared to main petiole, with enhanced mechanical support for deeply dissected lobes and improved vascular connection potentially enabling these lobes to function more independently. Evolutionary interpretation suggests these structures represent maximum expression of compound leaf developmental programs partially retained in Vitis vinifera genome but normally suppressed, with occasional breakthrough expression in extreme variants. Practical importance remains limited to morphological description, with these structures too inconsistent for ampelographic diagnostics but interesting for developmental biology studies.
bifurcated petiole
bifurkierter Nebenstängel
branched secondary petiole
forked lateral petiole
gegabelter sekundärer Petiolus
verzweigter lateraler Blattstiel
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 166-197 Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Tsukaya (2013) "Leaf development" The Arabidopsis Book 11=e0163 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 56-94.
Branch Lateral Petiole
Cabernet Franc showing branch second-order lateral shallow roots, excavation revealing second-order branches 0.9-2.4 cm diameter emerging from first-order lateral at 6-14 cm intervals, branching angles 40-75° from first-order axis, proximal second-order branches larger 1.8-2.4 cm than distal 0.9-1.4 cm, second-order branches extending 30-70 cm producing third-order branches every 4-9 cm, branch second-order laterals demonstrating characteristic branch morphology creating hierarchical architecture
Branch second order lateral shallow root is second-order lateral shallow root emphasizing branching relationship and role as branch from first-order parent, characterized by clear branch identity emerging laterally from first-order axis, variable size 0.5-3 cm diameter depending on position and age, demonstrating branch-typical characteristics including smaller diameter than parent, acute or obtuse branching angle, developmental subordination to parent lateral, and serving function of extending exploration and creating branching points for higher-order roots, with branch second-order laterals showing characteristics including endogenous origin from pericycle-like tissue in first-order parent, emergence through first-order cortex and bark creating visible branch point, predominantly lateral orientation spreading perpendicular or obliquely from first-order axis, variable development with proximal branches near first-order origin sometimes larger and more developed than distal branches, moderate growth potential showing limited to moderate secondary thickening, and architectural role creating hierarchical branching pattern where multiple second-order branches from single first-order parent collectively explore large soil volume. Branch second order lateral shallow root branching morphology includes= showing branch identity (clear morphological identity as branch from first-order parent axis), exhibiting branching angle (second-order branches emerging at characteristic angles 30-90° from first-order axis), demonstrating diameter contrast (branch diameter smaller than parent reflecting developmental hierarchy), revealing taper pattern (branches typically tapering from base to tip though less pronounced than parent), and displaying branch point (visible branch point where second-order emerges from first-order creating junction). Branch second order lateral shallow root positional variation includes= showing proximal-distal gradient (second-order branches near first-order base sometimes differing from those near tip), exhibiting size variation (proximal second-order branches potentially larger diameter 1.5-3 cm than distal 0.5-1.5 cm), demonstrating developmental differences (proximal branches forming earlier showing more secondary growth than later-forming distal branches), revealing functional gradients (proximal branches potentially emphasizing transport while distal emphasizing absorption), and displaying age structure (age of second-order branches decreasing with distance from first-order origin creating gradient). Branch second order lateral shallow root hierarchical relationship includes= showing parent dependence (branch second-order laterals physiologically dependent on first-order parent for water, nutrients, photosynthates), exhibiting developmental subordination (branch development potentially limited if parent vigor declines or parent dies), demonstrating resource competition (multiple second-order branches competing for resources from single first-order parent), revealing apical dominance effects (proximal second-order branches potentially showing suppression from first-order apical dominance), and displaying integration (branch second-order laterals fully integrated into hierarchical root system architecture). Branch second order lateral shallow root growth patterns includes= showing variable vigor (individual second-order branches showing widely varying growth rates and final sizes), exhibiting opportunistic growth (second-order branches potentially showing rapid growth when encountering favorable conditions), demonstrating arrested development (some second-order branches remaining small or dying if conditions unfavorable), revealing compensatory growth (remaining second-order branches potentially growing more vigorously if neighboring branches die), and displaying plasticity (second-order branch development showing high plasticity responding to heterogeneous conditions). Branch second order lateral shallow root third-order production includes= showing abundant branching (successful second-order branches producing numerous third-order branches), exhibiting branching frequency (third-order branches emerging every 3-10 cm along second-order length), demonstrating hierarchical progression (third-order branching creating fourth level progressing toward terminal fine roots), revealing multiplication effect (each second-order branch potentially supporting hundreds of higher-order roots), and displaying absorptive focus (third-order and higher derivatives from second-order branches providing majority of absorption). Branch second order lateral shallow root lifespan and fate includes= showing variable persistence (individual second-order branches potentially surviving months to years depending on development and conditions), exhibiting differential survival (larger more developed second-order branches showing higher survival than small suppressed branches), demonstrating mortality patterns (second-order branch death rates potentially higher in drought years or stressful conditions), revealing replacement (first-order parents continuously producing new second-order branches replacing losses), and displaying population turnover (second-order branch population showing continuous turnover through production and mortality). Branch second order lateral shallow root in root system function includes= providing architectural multiplication (second-order branching greatly expanding root system spatial extent and complexity), enabling thorough exploration (multiple second-order branches from each first-order creating intensive soil exploration), demonstrating resource capture (second-order branches and derivatives collectively providing substantial fraction of uptake), revealing structural distribution (second-order branches distributing root system mechanically reducing stress concentrations), and displaying functional importance (second-order branching pattern critically affecting overall root system performance).
Branch_Second_Order_Lateral_Shallow_Root emphasizes branching relationship and hierarchical identity and is distinguished from other aspects by focus on branch characteristics. Critical for understanding branching morphology, hierarchical dependence, positional variation, third-order support, and the second-order branches creating hierarchical architecture extending exploration and supporting absorptive network.
Ast-zweite Ordnung Lateralwurzel
Branch lateral root
Lateral branch second-order
Second-order branch root
Seitenast-Wurzel
Verzweigungs-Lateralwurzel
Smart, D. R., et al. (2006). Grapevine rooting patterns= A comprehensive analysis and a review. American Journal of Enology and Viticulture, 57(1), 89-104.
Branch Second Order Lateral Shallow Root
Pinot Noir grafted on Riparia Gloire rootstock at 12 years showing branch secondary root hierarchy, first-order laterals 2.2-4.5 cm diameter producing second-order branches 0.9-2.1 cm diameter at 8-14 cm intervals, second-order branches producing third-order branches 0.3-0.8 cm diameter at 4-9 cm intervals, third-order branches producing fourth-order fine roots <3 mm diameter at 2-5 cm intervals, fourth and fifth-order fine roots producing abundant root hairs and mycorrhizae, hierarchical branching creating exponential multiplication from approximately 12 primary and first-order roots to estimated 50,000+ finest branch roots, branch secondary root architecture enabling thorough soil exploration
Branch secondary root is secondary root emphasizing branching relationship as branch arising from parent secondary root creating hierarchical architecture, characterized by origin from another secondary root rather than from primary root distinguishing from first-order laterals, representing second or higher branching order in root system hierarchy, typically smaller diameter than parent root reflecting hierarchical size reduction with successive branching, and serving progressively more absorptive and less structural function with increasing branch order as root system transitions from large woody structural roots to fine ephemeral absorptive roots, with branch secondary root encompassing range from moderate-diameter second-order structural branches 1-3 cm diameter to fine high-order absorptive branches <2 mm diameter, showing typical branching pattern of parent root producing branch roots at intervals along length with branches being smaller and more numerous than parent, and demonstrating hierarchical organization principle where each branching level explores finer spatial scale creating nested architecture enabling thorough soil exploration. Branch secondary root hierarchical branching includes= arising from parent secondary root (branch roots originating from existing secondary root rather than from primary root), representing successive branching levels (second-order branches from first-order parents, third-order from second-order, continuing to higher orders), showing decreasing diameter (branch roots progressively smaller than parent roots with each successive branching level), exhibiting increasing number (more branch roots than parent roots at each level reflecting exponential branching), and demonstrating nested architecture (branching creating hierarchical nested structure with each level exploring finer spatial scale). Branch secondary root size hierarchy includes= ranging from moderate to fine (branch secondary roots spanning 0.1-3 cm diameter depending on branching order), showing inverse order-diameter relationship (lower-order branches larger diameter while higher-order branches progressively finer), exhibiting gradual taper within order (individual branch root diameter declining from origin toward tip as produces own branches), demonstrating exponential abundance (total number and length of branch roots increasing exponentially with each higher order), and revealing functional transition (larger branch roots more structural and transport while smaller branch roots more absorptive). Branch secondary root spacing and density includes= showing variable internode distance (spacing between successive branch roots along parent varying with order, vigor, and conditions), exhibiting typical ranges (second-order branches 5-15 cm intervals, third-order 3-8 cm, fourth and higher 1-5 cm reflecting finer branching scale), demonstrating acropetal gradient (branch density typically higher near parent root tip declining toward base), showing proliferation response (branch root density increasing in favorable soil zones and decreasing in poor zones), and creating exploration efficiency (branch spacing optimizing soil exploration without excessive overlap or gaps). Branch secondary root anatomy and structure includes= showing decreasing secondary growth (lower-order branches exhibiting secondary xylem development while higher-order branches showing minimal or no secondary growth), demonstrating anatomical transition (progressive change from woody structure in large branches to primary anatomy in fine branches), exhibiting variable bark (large branch roots developing protective periderm while fine branches maintaining thin epidermis or exodermis), displaying root hair presence (fine branch roots particularly tips producing abundant root hairs increasing absorption surface), and revealing mycorrhizal colonization (fine branch roots hosting arbuscular mycorrhizae enhancing nutrient and water uptake). Branch secondary root functional specialization includes= transitioning from transport to absorption (lower-order branches primarily conducting while higher-order branches increasingly absorptive), showing mixed functions (intermediate-order branches both conducting water from finer branches and absorbing directly), demonstrating storage capacity (larger branch roots accumulating carbohydrate reserves while fine branches showing minimal storage), exhibiting ephemeral versus persistent (fine high-order branches typically annual or multi-annual turnover while larger lower-order branches persisting years to decades), and revealing ecological roles (branch roots of different orders occupying different functional niches in resource acquisition strategy). Branch secondary root in hierarchical networks includes= creating exponential exploration (branching multiplication enabling vast total root length from moderate number of primary roots), enabling multi-scale exploitation (different branch orders exploring different spatial scales from coarse first-order to fine high-order), demonstrating fractal-like architecture (self-similar branching pattern repeated at decreasing scales creating fractal geometry), showing optimization principles (branching hierarchy potentially optimizing resource acquisition efficiency balancing construction costs and benefits), and revealing evolutionary adaptation (hierarchical branching representing fundamental plant architecture evolved for efficient soil exploration). Branch secondary root growth dynamics includes= showing continuous production (parent roots continuously producing new branch roots throughout growing season), exhibiting seasonal patterns (branch root initiation concentrated during active growth periods spring and fall), demonstrating turnover cycles (fine high-order branches showing rapid turnover with annual mortality and replacement while larger branches persistent), revealing responsive behavior (branch production rate and direction responding to soil resource heterogeneity), and enabling plasticity (branch root architecture adjusting to environmental conditions optimizing resource acquisition).
Branch_Secondary_Root emphasizes hierarchical branching relationship and is distinguished from lateral root (orientation emphasis) by focus on parent-branch hierarchy. Includes Major_Branch_Secondary_Root and Minor_Branch_Secondary_Root. Critical for understanding hierarchical organization, successive branching levels, size-order relationships, nested architecture, and the branching hierarchy creating exponential root multiplication enabling thorough multi-scale soil exploration.
Hierarchical branch root
Hierarchische Seitenwurzel
Nested lateral root
Successive order root
Sukzessive Wurzelordnung
Verzweigungs-Sekundärwurzel
Branch Secondary Root
A branched tendril at node 7 on a shoot of Vitis vinifera cv. Cabernet Sauvignon in Bordeaux, France, exhibiting bifurcated architecture with branch point at 6 cm from the shoot attachment, dividing into two arms of 11 cm and 12 cm length creating total tendril system span of 18 cm diameter when fully extended, with both branches showing active circumnutation in different planes expanding the explored volume to approximately 520 cubic centimeters, one branch making successful contact with a horizontal trellis wire and initiating coiling while the other branch continues searching movements, providing dual functionality of established support plus continued exploration for additional attachment opportunities, measured at BBCH stage 71 (fruit set) in early June, representing the species-typical bifurcated pattern common in this cultivar where 85-95% of tendrils show two-branch architecture.
A tendril exhibiting division of the main axis into two or more separate arms or branches in Vitis vinifera, characterized by presence of one or more branch points where the tendril shaft divides into distinct elongated segments, each branch capable of independent growth, circumnutation, contact detection, and coiling responses, with branching patterns determined by genetic programming specific to cultivar and modified by environmental conditions during development, representing an evolutionary adaptation that increases the probability of successful contact with support structures through expansion of the three-dimensional search space explored during circumnutation movements. Branched tendrils arise through differential activity of meristematic regions during early tendril development, with branch points established during the initial 1-2 weeks post-emergence when apical and lateral meristems determine architectural organization, followed by independent elongation of each branch segment creating the mature branched morphology. The number, position, and relative development of branches constitute important ampelographic characteristics used for cultivar identification, with branching patterns showing cultivar-specific preferences ranging from predominantly unbranched (simple) tendrils in some varieties to consistently multi-branched tendrils in others, though environmental factors including nutrient availability, temperature during development, and shoot vigor can modify expression of genetic branching tendencies. Branched tendrils provide functional advantages through increased contact probability as multiple independent arms explore different spatial zones, enhanced mechanical stability through multiple potential attachment points, and improved load distribution when multiple branches achieve successful attachment to different support structures.
Branched tendrils are distinguished from unbranched (simple) tendrils by presence of division points creating multiple independent elongated segments rather than a single continuous axis. Branch number, symmetry, and relative arm development serve as important cultivar identification features, with some varieties showing consistent branching patterns (e.g., Riesling predominantly bifurcated, Syrah showing higher trifurcated frequency) while others show more variable patterns. Branching should be distinguished from damage-induced splitting or pathological division, with true branching showing organized meristematic origin and symmetrical vascular development at branch points. Environmental stress during early development can reduce branching frequency or create asymmetric branch development even in cultivars with strong genetic branching tendencies. The functional advantage of branching increases with branch number up to approximately 3-4 branches, beyond which diminishing returns occur due to resource dilution across branches reducing individual branch vigor and coiling capacity.
divided tendril
gegabelte Ranke
mehrteilige Kletterorgan
multi-arm tendril
ramified tendril
verzweigte Ranke
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 58-73 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 79-94.
Branched Tendril
Syrah grafted on SO4 rootstock at 15 years showing branching lateral tertiary roots, representative third-order branching tertiary root 1.4 cm diameter at origin extending 11 cm length at depth 56 cm, branching root producing 14 fourth-order lateral branches 0.4-0.8 cm diameter at 6-9 cm intervals, fourth-order branches producing 8-12 fifth-order laterals 0.2-0.4 cm diameter at 3-5 cm intervals, fifth-order producing terminal sixth-order fine roots <2 mm diameter with abundant root hairs, branching lateral tertiary root network creating hierarchical architecture exploring approximately 500 cm³ local soil volume, branching root serving as structural framework and transport conduit for absorptive fine root network
Branching lateral tertiary root is lateral tertiary root showing active branching producing higher-order lateral roots, emphasizing branching capacity and creation of hierarchical network rather than terminal absorption, characterized by third or fourth-order position serving as parent to fourth fifth and sixth-order roots, moderate diameter typically 0.5-2 cm reflecting intermediate order between large secondary roots and finest absorptive roots, moderate secondary growth in larger specimens showing limited xylem development, and dual function as both structural support for higher-order branches and transport conduit moving water and nutrients from absorptive network to larger parent roots, with branching lateral tertiary roots showing branch production at 1-8 cm intervals depending on order and vigor creating dense hierarchical network, demonstrating lifespan typically 1-3 years intermediate between ephemeral fine roots and persistent woody roots, and representing critical architectural element connecting large persistent secondary root framework to ephemeral fine absorptive root network enabling extensive soil exploration through hierarchical branching multiplication. Branching lateral tertiary root branching activity includes= producing numerous higher-order laterals (branching tertiary roots generating 5-20+ fourth fifth and sixth-order branches depending on length and conditions), showing active branch initiation (continuous production of new branch roots during growing season), exhibiting variable branch spacing (higher-order branches typically emerging 1-8 cm intervals with closer spacing for finer branches), demonstrating acropetal branching gradient (branch density typically higher near growing tip declining toward base), and creating hierarchical exploration (branching pattern enabling systematic exploration of local soil volume through nested architecture). Branching lateral tertiary root diameter and structure includes= showing moderate diameter (branching tertiary roots typically 0.5-2 cm intermediate between large secondary roots and finest absorptive roots), exhibiting gradual taper (diameter declining along length as branches emerge and root extends), demonstrating limited secondary growth (third-order branching roots potentially showing 1-3 annual xylem rings while fourth-order minimal or absent secondary growth), displaying primary to transitional anatomy (smaller branching tertiary roots maintaining primary structure while larger showing initial secondary development), and revealing branching-order correlation (third-order branching roots thicker than fourth-order reflecting hierarchical size reduction). Branching lateral tertiary root hierarchical position includes= typically third or fourth-order (branching lateral tertiary roots most commonly being third-order from second-order parents or fourth-order from third-order parents), serving as intermediate generation (branching roots occupying middle position between large secondary framework roots and finest terminal absorptive roots), producing terminal branches (branching tertiary roots generating fourth through sixth-order roots which may be terminal in hierarchy), demonstrating multiplication function (branching roots creating exponential increase in root number and surface area through successive branching), and enabling nested architecture (branching pattern creating hierarchical nested structure exploring soil at progressively finer scales). Branching lateral tertiary root transport function includes= conducting water from branches (branching tertiary root xylem moving water absorbed by higher-order roots toward parent secondary roots), transporting nutrients basipetally (branching roots conveying minerals from absorptive network toward plant body), showing moderate hydraulic conductivity (branching root vessels providing adequate transport capacity for local network though less than larger roots), enabling phloem distribution (branching root phloem conducting photosynthates to finer absorptive roots supporting their metabolism), and demonstrating local hub function (branching roots serving as collection and distribution points for resources within local soil volume). Branching lateral tertiary root structural support includes= providing framework for fine roots (branching tertiary roots creating structural scaffold from which finest absorptive roots develop), bearing weight of branch network (branching roots supporting mechanical loads from higher-order branches and absorbed water), enabling spatial organization (branching root distribution determining architecture of local fine root network), demonstrating flexibility (branching roots bending and adjusting to soil structure accommodating obstacles), and showing limited persistence (branching roots functioning 1-3 years providing semi-permanent framework for dynamic fine root component). Branching lateral tertiary root absorption contribution includes= showing mixed function (branching tertiary roots both conducting from finer branches and absorbing directly particularly at growing tips), demonstrating direct uptake at tips (actively growing branching root tips producing root hairs and absorbing water and nutrients), hosting mycorrhizal colonization (branching roots showing AM fungal associations particularly in younger portions), exhibiting seasonal absorption variation (branching root direct absorption highest during active growth declining during dormancy), and revealing functional transition (branching roots transitioning from primarily absorptive when young to primarily transport when older and more branched). Branching lateral tertiary root lifespan and replacement includes= showing intermediate longevity (branching lateral tertiary roots typically functioning 1-3 years longer than finest roots but shorter than woody secondary roots), exhibiting eventual senescence (older branching roots declining in function and dying as newer branching roots produced), demonstrating continuous replacement (ongoing production of new branching roots replacing lost or senescing roots maintaining hierarchical network), revealing variable mortality (branching root death rate varying with environmental stress, age, and pathogen pressure), and contributing to turnover dynamics (branching root production and mortality representing intermediate component in overall root system turnover). Branching lateral tertiary root resource responsiveness includes= proliferating in favorable zones (branching tertiary roots producing more higher-order branches in resource-rich soil patches), showing enhanced branching density (branching intensity increasing when encountering high nutrient or moisture availability), demonstrating selective branching (preferential branch production in directions toward resources or away from barriers), revealing competitive plasticity (branching pattern adjusting in response to root competition from neighbors), and enabling efficient exploitation (branching architecture optimizing local resource acquisition while minimizing construction and maintenance costs).
Branching_Lateral_Tertiary_Root emphasizes active branching and hierarchical network creation and is distinguished from terminal non-branching fine roots by focus on branch production. Critical for understanding hierarchical architecture, branching multiplication, intermediate transport function, semi-permanent framework, nested exploration, and the branch-producing tertiary roots connecting persistent secondary framework to ephemeral absorptive periphery through successive branching levels.
Branch-producing tertiary root
Hierarchical lateral tertiary
Hierarchisch verzweigte Feinwurzel
Mehrfach-Ordnung Seitenwurzel
Multi-order lateral root
Verzweigende Laterale Tertiärwurzel
Branching Lateral Tertiary Root
Comprehensive characterization of Branching_Pattern_Disposition at INRAE Bordeaux quantified lateral shoot development across 55 cultivars grown under standardized conditions (VSP, moderate vigor, no lateral removal) throughout the growing season. Measurements included number of lateral shoots per primary shoot, total lateral length, lateral shoot positions (basal, mid, distal nodes), and the ratio of lateral to primary shoot biomass. Additionally, auxin concentration in shoot apices and cytokinin sensitivity of lateral buds were assessed to investigate mechanistic basis. Branching_Pattern_Disposition showed substantial cultivar variation. 'Carignan' exhibited prolific branching with 14.5 lateral shoots per primary shoot, total lateral length of 245 cm (exceeding primary shoot length of 210 cm), and lateral/primary biomass ratio of 1.15, indicating that more resources were allocated to lateral than primary growth. 'Merlot' showed strong apical dominance with only 4.2 lateral shoots per primary, total lateral length of 48 cm, and biomass ratio of 0.22. 'Cabernet Sauvignon' displayed intermediate branching (7.8 laterals, 95 cm total length, ratio 0.45). Mechanistic analysis revealed that apex auxin concentration was 35% higher in strong apical dominance cultivars, while lateral bud cytokinin sensitivity (measured by bud outgrowth response to exogenous cytokinin) was 2.5-fold higher in prolific branching types. The practical significance was confirmed by canopy management labor requirements—prolific branching cultivars required 85 hours/ha for lateral management versus 25 hours/ha for strong apical dominance types.
A Branching_Pattern_Disposition is a Growth_Habit_Disposition that inheres in a Vitis vinifera plant and is realized through the characteristic pattern of lateral shoot (axillary shoot) development from primary shoot nodes, determining the degree to which vines produce branched versus unbranched shoot structures during the growing season. This disposition represents the balance between apical dominance (hormonal suppression of lateral bud outgrowth by the dominant shoot apex) and the tendency for lateral bud release (outgrowth of axillary buds into lateral shoots), together determining canopy architecture complexity and density. Branching pattern profoundly affects canopy management requirements, as cultivars with prolific lateral production develop dense, multi-layered canopies requiring intensive shoot thinning and lateral removal, while cultivars with strong apical dominance produce simpler canopy structures with reduced management intervention. The disposition reflects genotypic differences in auxin production by shoot apices, cytokinin sensitivity of lateral buds, and the hormonal communication between apex and lateral buds that regulates correlative inhibition. Branching_Pattern_Disposition interacts with vigor—high-vigor cultivars express their branching tendency more fully than low-vigor cultivars where limited resources constrain lateral development regardless of hormonal predisposition. Understanding branching pattern enables prediction of canopy management needs and guides decisions about shoot thinning, lateral removal, hedging timing, and training system complexity appropriate for each cultivar.
Branching_Pattern_Disposition is_a Growth_Habit_Disposition addressing lateral shoot development patterns. Branching reflects the balance between apical dominance (suppression) and lateral bud release (outgrowth). Child classes include Strong_Apical_Dominance_Disposition, Weak_Apical_Dominance_Disposition, and Lateral_Shoot_Proliferation_Disposition. Auxin-cytokinin balance underlies branching pattern differences. Branching interacts with vigor—limited resources constrain lateral expression regardless of hormonal predisposition. Branching pattern determines lateral management requirements and canopy density.
Geiztrieb-Entwicklung
Triebarchitektur-Muster
Verzweigungsmuster-Disposition
axillary shoot development
lateral branching habit
shoot architecture pattern
Lebon E et al. (2006) Shoot development in grapevine (Vitis vinifera L.) is affected by the modular branching pattern of the stem and intra- and inter-shoot trophic competition, Annals of Botany 98:1045-1056 Cline MG (1991) Apical dominance, Botanical Review 57:318-358 Bennett T, Leyser O (2006) Something on the side: axillary meristems and plant development, Plant Molecular Biology 60:843-854
Branching Pattern Disposition
The brand name Regent is a registered trademark of the Julius Kühn-Institut for the PIWI cultivar bred from Diana (Silvaner x Müller-Thurgau) x Chambourcin, with the JKI controlling commercial use of the name Regent for planting material and requiring licensing agreements with nurseries propagating the variety — similarly, the brand name Cabernet Blanc is a registered trademark of Valentin Blattner for his interspecific hybrid breeding VB 91-26-29, and the brand name Piwi International is used collectively by a consortium of breeders to market disease-resistant cultivars under a shared quality mark.
A trade name that is a proprietary designation owned by a specific commercial entity — nursery, breeding company, or wine producer — used exclusively to market grapevine planting material or wine products under a distinctive brand identity that is legally controlled through trademark registration or equivalent intellectual property mechanisms.
Brand_Name represents the intersection of cultivar nomenclature with intellectual property law. Under UPOV (International Union for the Protection of New Varieties of Plants) conventions, Plant Variety Protection grants breeders exclusive rights to control propagation and sale of new cultivars, and brand names function as the commercial identity through which these rights are exercised: however, UPOV also requires that a registered variety denomination (distinct from the brand name) remains freely usable by all parties, creating a dual naming system where the brand name is proprietary but the variety denomination is public. This distinction is frequently confused in practice, particularly when the brand name and variety denomination are identical.
Markenname
Markensorten-Bezeichnung
branded cultivar designation
eingetragener Handelsname
proprietary name
trademarked variety name
UPOV (2020) International Convention for the Protection of New Varieties of Plants ICNCP (2016) International Code of Nomenclature for Cultivated Plants 9th Ed, ISHS Art. 11 Töpfer R et al. (2011) New horizons for grapevine breeding, Fruit, Vegetable and Cereal Science and Biotechnology 5: 79-100
Brand Name
Airén bears a Brandy_Production_Scion_Role as the primary variety for Brandy de Jerez production, where its neutral character, high yields (often exceeding 150 hL/ha in La Mancha), and abundant availability provide economical base wine for pot-still distillation and subsequent solera aging in Jerez sherry casks. Palomino Fino serves in a Brandy_Production_Scion_Role for premium Brandy de Jerez, where used sherry barrels (formerly containing Fino or Oloroso) contribute distinctive nutty, oxidative character to the aged spirit. Colombard bears a Brandy_Production_Scion_Role in California and South Africa, providing aromatic complexity superior to purely neutral varieties while maintaining the acidity and yield characteristics essential for distillation economics. Chenin Blanc (Steen) serves in a Brandy_Production_Scion_Role for South African brandy, where the variety's naturally high acidity supports quality spirit production from the warm Cape climate.
A distillation scion role that inheres in a grafted scion cultivar by virtue of its designation for general brandy production outside the geographically protected Cognac, Armagnac, and Pisco appellations, where wine is distilled into grape-based spirits for consumption neat, in cocktails, or for culinary applications. Brandy production scions share the fundamental distillation requirements—high acidity, neutral character, moderate alcohol potential, and economic yields—but are not constrained by the strict varietal regulations governing protected appellations. Brandy production occurs globally wherever wine grapes are cultivated, with major production in Spain (Brandy de Jerez), South Africa, California, Australia, Germany (Weinbrand), Greece (Metaxa), and numerous other regions. Variety selection varies by region and tradition, ranging from the Airén and Palomino of Spanish brandy through Thompson Seedless in California to Chenin Blanc (Steen) in South Africa. Production methods include pot still distillation (producing more flavorful spirits) and continuous column distillation (more efficient, lighter spirits), with aging in oak barrels creating the characteristic color and flavor of aged brandies.
Brandy_Production_Scion_Role captures the general category of brandy production distinct from the appellation-controlled Cognac, Armagnac, and Pisco categories with their strict varietal requirements. Brandy quality varies enormously—from industrial spirits distilled from surplus wine to artisanal pot-still brandies rivaling Cognac in complexity. The Spanish solera-aged Brandy de Jerez tradition represents a distinctive style where sherry cask aging contributes as much character as grape variety or distillation method. Global brandy consumption exceeds Cognac and Armagnac combined, with value-oriented production dominating volume while premium artisanal brandies serve quality-focused markets. Climate change is expanding brandy production into regions previously too cool for wine, as distillation can utilize high-acid, moderate-alcohol base wines unsuitable for table wine.
Brandytraube
Weinbrandsorte
allgemeine Brenntraube
brandy grape
general distillation grape
wine spirit variety
Piggott JR et al. (1989) Distilled Beverage Flavour, Ellis Horwood Faith N (2016) Cognac and Other Brandies, Infinite Ideas Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Consejo Regulador del Brandy de Jerez (2020) Production Regulations
Brandy Production Scion Role
The breeding code Gf.1993-22-6 assigned by the Julius Kühn-Institut at Geisenheim identifies the genotype that was subsequently released as the PIWI cultivar Calardis Blanc — the code encodes the institution (Gf for Geisenheim), the crossing year (1993), the cross combination number (22), and the individual seedling number (6) within that cross, providing a complete genealogical address within the institutional breeding records that remains the definitive identifier in all research publications and trial reports even after the commercial name Calardis Blanc was assigned at registration.
A cultivar designation consisting of an alphanumeric code assigned by a breeding institution to identify a grapevine genotype within the institutional selection and evaluation pipeline, serving as the primary identifier during the pre-release experimental phase before a commercial cultivar name is assigned upon official registration.
Breeding_Code is the standard identifier for grapevine genotypes during the 10-25 year period between initial crossing and potential cultivar release. The code structure varies by institution but typically encodes some combination of institution abbreviation, crossing year, cross number, and seedling number: examples include Gf (Geisenheim), Fr (Freiburg), We (Weinsberg), and JKI-Gm (Julius Kühn-Institut Geilweilerhof/Siebeldingen) in Germany, and INRA/INRAE numbering in France. Subclasses distinguish the principal code types within breeding programs — experimental codes, selection numbers, cross numbers, and seedling numbers — reflecting different stages and organizational levels of the breeding pipeline.
Selektionsnummer
Züchtungscode
breeder's code
institutional genotype identifier
institutionelle Genotypkennung
selection code
Eibach R and Töpfer R (2015) Traditional grapevine breeding techniques, in Reynolds AG (ed) Grapevine Breeding Programs for the Wine Industry, Woodhead Publishing pp: 3-22 Töpfer R et al. (2011) New horizons for grapevine breeding, Fruit, Vegetable and Cereal Science and Biotechnology 5: 79-100
Breeding Code
A downy mildew-resistant selection from the INRA-ResDur breeding program bears a Breeding_Line_Clone_Role, fifth-generation material carrying introgressed Muscadinia resistance genes under advanced evaluation for potential registration as a disease-resistant variety. A Pinot Noir × Gamay cross maintained at Geisenheim serves in a Breeding_Line_Clone_Role, early-ripening selections being evaluated for potential development into new varieties for cool-climate production. A white-fruited selection from Weinsberg crossing programs bears a Breeding_Line_Clone_Role, Müller-Thurgau × Riesling progeny under evaluation for quality and productivity before determining commercialization potential.
An experimental clone role that inheres in a grapevine genotype maintained as part of an active breeding program, representing intermediate steps in variety development through controlled hybridization, selection, and evaluation toward potential commercial release. Breeding line clones occupy the developmental pipeline between initial crosses (where thousands of seedlings may be generated) and final variety registration (where selected individuals become named, registered cultivars). Breeding lines undergo progressive selection—initial screening eliminating most seedlings, intermediate evaluation assessing promising selections, and advanced trials confirming commercial potential before registration. The breeding line designation indicates material under development rather than finished product, with genotypes potentially being discarded, further developed, or eventually released as new varieties depending on evaluation outcomes. Breeding lines may incorporate genetic material from wild Vitis species (particularly for disease resistance introgression), from interspecific hybridization programs, or from within-species controlled crosses to combine traits from distinct cultivated parents. Access to breeding lines is typically restricted during development, with intellectual property protection and competitive considerations limiting distribution.
Breeding_Line_Clone_Role captures material in the developmental continuum toward potential variety release, distinguished from Research_Clone which may be maintained indefinitely without commercialization intent. The breeding line designation reflects active evaluation toward commercial objectives—quality, productivity, disease resistance, or other market-relevant characteristics—with ongoing assessment determining whether material advances toward release or is discarded from further development. Contemporary breeding emphasizes disease resistance (downy mildew, powdery mildew through introgressed R-genes) and climate adaptation (heat tolerance, earlier ripening for cool climates), with breeding lines representing years of development investment before reaching registration. Intellectual property considerations increasingly influence breeding line management, with plant variety protection and licensing arrangements shaping distribution even after variety registration.
Entwicklungsklon
Zuchtlinie
Zuchtprogramm-Klon
Zuchtselektion
breeding line
breeding program clone
breeding selection
development clone
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, in Reynolds A (ed) Grapevine Breeding Programs for the Wine Industry, Elsevier pp.: 3-22 Schneider A et al. (2008) Clonal variation in grapevine, ISHS Acta Horticulturae 779: 397-404 Dry IB et al. (2010) Molecular strategies to enhance disease resistance in grapevine, Australian Journal of Grape and Wine Research 16: 94-105
Breeding Line Clone Role
Vitis vinifera cv. Riesling participates in a Breeding_Role as a quality parent in German resistance breeding programs, contributing wine quality traits to crosses with disease-resistant wild species derivatives. Muscadinia rotundifolia serves in a Breeding_Role as a resistance source in French breeding programs, its powdery mildew immunity targeted for introgression into V. vinifera backgrounds. A numbered breeding line from the Geisenheim program bears a Breeding_Role as both the product of previous crosses and potential parent for subsequent generations, illustrating the multi-generational nature of variety development.
A grapevine role that inheres in a plant or genotype by virtue of its participation in controlled breeding programs aimed at creating new varieties through sexual hybridization, combining genetic material from distinct parents to generate novel genotypes with desired trait combinations. Breeding roles encompass the functional positions plants occupy in crossing programs—as parents contributing genetic material (maternal, paternal), as sources of specific traits (resistance donors, quality donors), or as recipients of trait introgression (recurrent parents in backcross programs). Grapevine breeding differs fundamentally from clonal selection in creating genuinely new genetic combinations through meiosis and fertilization rather than selecting variants within existing clonal populations. Breeding programs pursue diverse objectives including disease resistance (introgression of R-genes from wild Vitis species), climate adaptation (earlier ripening, drought tolerance), and quality enhancement (improved fruit composition, wine character), with breeding cycles typically requiring 15-25 years from initial cross through variety evaluation and registration. The highly heterozygous nature of Vitis and strong inbreeding depression necessitate outcrossing strategies, with most breeding involving crosses between distinct genotypes rather than selfing.
Breeding_Role is the parent class for the hierarchy of breeding functions, subclassed by parental function (Parent_Role and its maternal/paternal subdivisions) and by trait contribution emphasis (Donor_Role for specific trait sources, Recurrent_Parent for elite variety improvement). The distinction between breeding roles and other grapevine roles lies in the sexual reproduction context—breeding involves meiosis, recombination, and novel genotype creation, while clonal propagation maintains existing genotypes. Grapevine breeding has intensified since phylloxera (necessitating resistant rootstock development) and accelerated dramatically since the 1990s with disease-resistant scion variety programs (PIWI varieties) seeking to reduce fungicide dependence. Contemporary breeding increasingly integrates molecular markers for trait selection, accelerating progress while building on traditional crossing and evaluation approaches.
Hybridisierungsrolle
Kreuzungsrolle
Sortenentwicklungsrolle
Züchtungsrolle
breeding role
crossing role
hybridization role
variety development role
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, in Reynolds A (ed) Grapevine Breeding Programs for the Wine Industry, Elsevier pp.: 3-22 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Hausmann L et al. (2019) Grapevine breeding for disease resistance, in Cantu D, Walker MA (eds) The Grape Genome, Springer pp.: 311-330
Breeding Role
Brett character evaluation comparing California cult Cabernet (zero tolerance, 4-EP below 50 μg/L detection limit) with traditional Châteauneuf-du-Pape (moderate Brett accepted, 4-EP 650 μg/L, 4-EG 85 μg/L creating leather-barnyard complexity) demonstrates the regional variation in Brett acceptance, with California example showing pristine fruit expression without animal complexity while Rhône example integrates Brett character with fruit and garrigue as traditional terroir expression, illustrating how identical compounds constitute fault or typicity depending on regional context and style expectation.
Brett character quality is an off-aroma quality that represents olfactory perception of volatile phenols produced by Brettanomyces and Dekkera yeast species, characterizing the distinctive barnyard, horse stable, medicinal, and leather aromatic impressions that indicate wine contamination by these spoilage organisms. Within the BFO framework, this quality specifically addresses the Brettanomyces fault category, distinguishing Brett-derived off-aromas from other fault types while acknowledging the controversial status of this character where low levels are accepted or even valued in certain regional traditions (Rhône, Bordeaux historically) while being strictly rejected in others. The ontological significance lies in Brett character representing perhaps the most philosophically complex wine fault, where the same aromatic compounds are considered essential terroir expression in some contexts, acceptable complexity in others, and unequivocal spoilage in still others, raising fundamental questions about the nature of quality and the role of tradition in fault definition. The compositional basis for Brett character includes volatile phenols produced through Brettanomyces metabolism of hydroxycinnamic acid precursors naturally present in grapes, primarily 4-ethylphenol (4-EP, providing barnyard, horse stable, Band-Aid character, threshold approximately 400-600 μg/L depending on wine matrix and taster sensitivity) and 4-ethylguaiacol (4-EG, contributing smoky, spicy, bacon notes, threshold approximately 50-100 μg/L), with the 4-EP/4-EG ratio and absolute concentrations determining the specific character expression. The infection dynamics of Brettanomyces involve slow-growing yeast capable of survival and metabolism under wine conditions (low pH, high alcohol, low nutrients) where Saccharomyces cannot compete, with contamination sources including barrel surfaces, winery equipment, and cross-contamination between wines, and risk factors including residual sugar, elevated pH, and warm storage providing conditions favorable for Brett metabolism. The remediation and prevention for Brett contamination include filtration (sterile filtration removing cells but not already-formed volatile phenols), SO₂ management (maintaining adequate free molecular SO₂ to inhibit Brett metabolism), and sanitation protocols (preventing winery establishment of Brett populations), with already-formed volatile phenols effectively permanent and only addressable through blending or off-label sale. Understanding Brett character quality enables fault identification from volatile phenol-derived character, contamination risk assessment based on winery conditions and wine parameters, and the nuanced quality evaluation that acknowledges regional variation in Brett acceptance.
Brett_Character_Quality addresses the Brettanomyces-derived volatile phenol fault category, which uniquely demonstrates the context-dependency of fault definition. This quality requires acknowledgment of regional and stylistic variation in acceptance thresholds, with the same concentrations considered spoilage in some contexts and terroir expression in others. The class supports both fault identification and nuanced quality evaluation that respects legitimate stylistic diversity.
Brett-Charakter
Brettanomyces character
Brettanomyces-Noten
Pferdeschweiß
Stallton
barnyard aroma
mouse character
volatile phenols
Loureiro V, Malfeito-Ferreira M (2003) Spoilage yeasts in the wine industry. International Journal of Food Microbiology 86(1-2): 23-50 Chatonnet P et al. (1992) The origin of ethylphenols in wines. Journal of the Science of Food and Agriculture 60(2): 165-178
Brett Character Quality
Cabernet Sauvignon on 3309C bridge callus tissue at 5 weeks post-grafting shows 12-22 mm continuous callus structure spanning graft interface, bridge comprising intermixed scion and rootstock callus cells (estimated 45-55% scion-derived, 55-45% rootstock-derived based on cell characteristics), callus cells 40-75 μm diameter showing active division and early differentiation, localized regions showing organized cambial-like tissue (3-6 cell layer bands extending 2-5 mm tangentially), scattered vessel element precursors (elongated cells 120-200 μm length with developing secondary walls), bridge providing continuous tissue connection enabling limited transport (water, photosynthate moving through apoplast and symplast), bridge serving as substrate for progressive vascular differentiation establishing functional xylem and phloem connections within subsequent 3-6 weeks enabling permanent union integration
Bridge callus tissue is callus tissue specifically forming the continuous connection between scion and rootstock at graft union, comprising intermixed undifferentiated parenchyma cells from both graft partners that meet, adhere, and create the physical and physiological link enabling subsequent vascular differentiation and functional integration. Bridge callus tissue formation process includes= independent proliferation (scion and rootstock each producing callus from cut surfaces within 1-2 weeks post-grafting), callus expansion toward junction (callus tissues growing from both partners advancing toward graft interface), callus contact (scion and rootstock callus meeting at junction typically 2-4 weeks post-grafting), cell-cell adhesion (callus cells from both partners adhering forming cohesive continuous tissue), callus intermixing (scion and rootstock cells interdigitating creating integrated tissue mass rather than discrete layers), and bridge establishment (continuous callus connection spanning junction enabling subsequent tissue differentiation and integration). Bridge callus tissue structural characteristics includes= continuous tissue mass (single cohesive callus structure rather than separate scion and rootstock portions), intermixed cell populations (scion and rootstock callus cells interspersed throughout bridge rather than maintaining distinct zones), variable thickness (bridge typically 5-20 mm vertical extent, 100-150% circumferential dimension encompassing original cut plane plus adjacent callus), filling irregularities (callus conforming to cut surface variations ensuring complete contact), and providing three-dimensional connection (bridge spanning radially from pith to bark and circumferentially around entire junction). Bridge callus tissue cellular composition includes= undifferentiated parenchyma (thin-walled cells 30-80 μm diameter lacking specialized structure comprising majority of bridge tissue initially), partially differentiated cells (cells showing early specialization toward vascular fates interspersed in callus matrix), scion-derived cells (callus cells originating from scion genotype contributing approximately half of bridge), rootstock-derived cells (callus from rootstock genotype forming other half of bridge), and intermixed populations (scion and rootstock cells scattered throughout bridge creating genetic mosaic). Bridge callus tissue functions includes= providing physical connection (callus bridge creating continuous tissue linking scion and rootstock), enabling initial transport (callus providing apoplastic and symplastic pathways for limited water and solute movement before organized vascular tissues form), serving as differentiation substrate (bridge callus cells being precursors for cambium, xylem, phloem that establish functional vascular continuity), facilitating integration (callus intermixing promoting cell-cell interactions necessary for subsequent tissue adhesion and function), and determining union quality (bridge callus formation adequacy and subsequent differentiation controlling union success or failure). Bridge callus tissue vascular differentiation within includes= cambium formation (localized zones of bridge callus differentiating into cambial initials typically 3-6 weeks post-grafting), cambial organization (individual cambial patches coalescing forming continuous cambium across bridge), xylem element formation (vessel elements differentiating from cambium establishing first water transport connections), phloem element formation (sieve tubes and companion cells arising enabling photosynthate transport), progressive elaboration (continued secondary growth adding vascular tissues strengthening connection improving transport capacity), and connection with partners (newly formed vascular tissues linking with scion and rootstock vascular systems establishing functional continuity). Bridge callus tissue in successful unions includes= adequate callus formation (sufficient bridge callus completely spanning junction without gaps or discontinuities), good cell-cell adhesion (strong bonding between scion and rootstock callus cells creating cohesive bridge), progressive organization (bridge callus showing increasing structure, early vascular strand formation within 3-6 weeks), timely differentiation (bridge transitioning from undifferentiated callus to organized vascular tissues within 2-4 months), and functional integration (bridge-derived vascular tissues establishing effective transport connections enabling normal vine function). Bridge callus tissue in problematic unions includes= inadequate bridge formation (insufficient callus leaving gaps, poor contact between partners), weak cellular adhesion (poor bonding causing bridge separation or mechanical weakness), persistent undifferentiated state (bridge remaining as disorganized callus beyond 3-4 months without vascular differentiation), necrotic zones (tissue death within bridge interrupting continuity), localized bridge formation (callus bridge forming only portions of union circumference leaving other sections unconnected), and failure to integrate (bridge callus present but failing to differentiate or connect with partners resulting in incompatibility). Bridge callus tissue assessment includes= visual examination (observing callus bridge formation, extent, continuity at graft interface), tactile assessment (palpating union detecting firm bridge indicating good callus versus soft zones or gaps suggesting problems), dissection (cutting through union revealing bridge structure, extent of scion-rootstock intermixing), microscopy (examining cellular organization, differentiation status, presence of vascular elements), and functional testing (measuring transport capacity, mechanical strength indicating bridge quality). Bridge callus tissue formation requirements includes= intimate tissue contact (cut surfaces properly aligned, pressed together enabling callus from both partners to meet), adequate callus proliferation from both partners (scion and rootstock each producing sufficient callus to span distance and create bridge), compatible cell recognition (scion and rootstock callus cells recognizing each other as compatible enabling adhesion), favorable environment (temperature 20-28°C, high humidity 85-95%, protection from desiccation and pathogens supporting callus survival and growth), and mechanical stability (immobilization during critical bridge formation period preventing disruption of developing connection). Bridge callus tissue hormonal signals includes= auxin promoting proliferation (elevated IAA at cut surfaces stimulating callus division and bridge formation), cytokinin synergy (cytokinins from both partners enhancing cell division), auxin gradient effects (auxin flow patterns potentially affecting bridge organization and vascular differentiation orientation), signal exchange (hormones and other compounds moving between scion and rootstock through developing bridge potentially coordinating differentiation), and compatibility factors (compatible partners showing normal hormone signaling and responses, incompatible pairs potentially showing abnormal hormone metabolism or sensitivity affecting bridge development). Bridge callus tissue genetic composition includes= chimeric structure (bridge containing both scion and rootstock genotypes creating genetic mosaic), random intermixing (scion and rootstock cells scattered throughout bridge without distinct boundaries), periclinal chimerism potential (bridge eventually becoming organized with scion and rootstock tissues in separate layers as vascular system develops), and genetic interactions (scion and rootstock gene products potentially interacting in bridge cells affecting compatibility, differentiation, function). Bridge callus tissue mechanical properties includes= initially weak (newly formed callus bridge showing limited mechanical strength requiring support), progressively strengthening (vascular differentiation and secondary growth increasing bridge integrity), mature strength (well-developed bridge in successful unions achieving 70-90% of non-grafted trunk strength), and failure mode (incompatible or poorly formed bridges remaining weak vulnerable to fracture at interface under stress). Bridge callus tissue significance includes= absolutely essential for graft success (union formation completely dependent on adequate bridge callus formation and differentiation), determining union quality (bridge characteristics controlling vascular continuity, mechanical strength, functional capacity), reflecting compatibility (successful bridge formation and differentiation indicating compatible partners, bridge failure suggesting incompatibility), enabling scion-rootstock integration (bridge providing physical and physiological connection linking genetically distinct plants), and foundational for grafted vine (all subsequent union structure and function deriving from initial bridge callus serving as template and substrate for permanent tissues).
Bridge_Callus_Tissue emphasizes scion-rootstock connection function and is distinguished from Callus_Tissue (general undifferentiated tissue), Healing_Callus_Tissue (wound repair emphasis), and Callus_Graft_Union_Zone (union context) by focus on specific junction bridging role. Critical for understanding graft union formation mechanism, scion-rootstock cellular intermixing, vascular differentiation substrate, integration enabling function, bridge formation requirements, and bridge callus as essential structure determining graft success providing physical and physiological connection foundation for all subsequent union development and function.
Brücken-Kallusgewebe
Connecting callus
Junction callus
Union bridge tissue
Verbindungs-Kallus
Verwachsungs-Brücke
Pina, A., & Errea, P. (2005). A review of new advances in mechanism of graft compatibility-incompatibility. Scientia Horticulturae, 106(1), 1-11. | Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press.
Bridge Callus Tissue
Brilliance assessment in fine Champagne reveals star-bright clarity (brilliance rating 9.5/10) where the wine actively sparkles under examination light, with fine persistent mousse contributing to overall luminosity, contrasting with commercial prosecco showing adequate limpidity (particle-free) but lacking brilliance (rating 6/10) due to residual colloidal matter creating slight dulling of light transmission, demonstrating how brilliance distinguishes exceptional from adequate clarity and correlates with production quality investment reflected in pricing differential between these sparkling wine categories.
Brilliance quality is a clarity quality that represents the active reflection and refraction of light creating sparkle, luminosity, and jewel-like visual impression, characterizing the highest grade of optical clarity where wine not only lacks turbidity but actively interacts with light to produce an aesthetically pleasing visual effect. Within the BFO framework, this quality specifically addresses the light-reflective dimension of clarity, distinguishing brilliance from mere limpidity by requiring active optical engagement rather than simple particle absence, with brilliance representing the positive visual quality beyond the negative criterion of haze absence. The ontological significance lies in brilliance serving as a quality marker indicating meticulous wine production, where brilliant wines suggest careful fining and filtration that removed not only visible particles but also sub-visible colloidal matter that would dull light transmission, with brilliance thus signifying the extra effort and quality focus associated with premium wine production. The optical basis for brilliance involves multiple light phenomena, with specular reflection from the wine surface creating highlights, refraction through the wine body producing depth and luminosity, and the absence of scattering particles allowing undimmed light transmission, together creating the sparkling, jewel-like appearance that distinguishes brilliant from merely limpid wines. The assessment of brilliance employs descriptive terminology ranging from dull (no light interaction) through clear (particle-free but not reflective) to bright (slight sparkle) to brilliant (intense light reflection, star-bright) to crystalline (exceptional, gem-like clarity), with these grades providing graduated clarity assessment beyond simple limpid/hazy distinction. The production factors favoring brilliance include fine filtration (removing colloidal particles that dull appearance), effective protein stabilization (eliminating haze-forming proteins), absence of dissolved gases (which scatter light at microscopic bubbles), and appropriate pH and temperature maintaining colloidal stability, with all these factors optimized in wines achieving brilliant clarity. Understanding brilliance quality enables premium quality recognition based on superior clarity, production quality inference from optical excellence, and appropriate assessment distinguishing adequate clarity from exceptional brilliance.
Brilliance_Quality addresses the actively light-reflective dimension of clarity beyond simple particle absence. This quality represents the positive visual criterion (active sparkle) complementing the negative criterion of limpidity (haze absence). The class supports premium quality recognition and production quality inference, with brilliance indicating the meticulous handling and treatment associated with fine wine production.
Brillanz
Glanz
Leuchtkraft
crystalline appearance
kristallklare Erscheinung
luminosity
sparkling clarity
star-bright clarity
Peynaud E, Blouin J (1996) The Taste of Wine: The Art and Science of Wine Appreciation, 2nd Ed. John Wiley & Sons, New York, pp. 63-91 Jackson RS (2014) Wine Science: Principles and Applications, 4th Ed. Academic Press, San Diego, pp. 831-887
Brilliance Quality
Brittleness assessment of Pinot Noir rachis tissue comparing hand-harvest and machine-harvest suitability reveals fracture energy of 0.45 J/cm² at véraison increasing to 0.28 J/cm² at optimal harvest maturity (22°Brix) and further declining to 0.15 J/cm² in overripe fruit, with the 67% reduction in fracture energy indicating progressive brittleness increase that causes shattering losses exceeding 8% when machine harvest is delayed beyond 24°Brix, establishing the harvest window for mechanical harvest based on rachis brittleness rather than fruit composition alone.
Brittleness quality is a physical quality that inheres in grapevine tissues and represents the tendency to fracture with minimal plastic deformation under applied stress, characterized by sudden failure without significant yielding or warning when stress exceeds the failure threshold. Within the BFO framework, this quality characterizes the failure mode where tissues break abruptly rather than deforming progressively, representing the opposite extreme from ductile behavior where extensive plastic flow precedes fracture. The ontological significance lies in brittleness determining how tissues fail under mechanical stress, with brittle materials showing catastrophic crack propagation once failure initiates while ductile materials absorb energy through plastic deformation, allowing graceful rather than sudden failure. Brittleness in grapevine structures varies with tissue composition, hydration status, and temperature, with lignified tissues (woody stems, seed coats) showing inherent brittleness due to cross-linked polymer networks that resist chain slippage, while parenchymatous tissues become brittle when dehydrated or frozen as water loss eliminates the plasticizing effect of cellular fluid. The rachis and pedicel structures show developmentally programmed brittleness changes, with the abscission zone at the pedicel-berry junction developing controlled brittleness facilitating berry detachment at maturity, while excessive brittleness in the rachis causes shattering losses during mechanical harvest. Temperature dramatically affects brittleness, with tissues becoming more brittle at lower temperatures as polymer chain mobility decreases and water crystallization eliminates plasticization, relevant to cold storage conditions and freezing damage assessment. Measurement approaches include fracture testing determining energy absorption before failure, impact testing assessing damage from dynamic loading, and acoustic emission monitoring detecting crack initiation and propagation during deformation. Understanding brittleness quality enables prediction of shattering losses during harvest and handling, optimization of harvest timing relative to rachis condition, and interpretation of mechanical damage patterns in terms of tissue failure mode.
Brittleness_Quality serves as the parent class for structure-specific brittleness qualities (Rachis, Pedicel), establishing the framework for fracture behavior characterization in vine structures. This quality represents failure mode rather than strength, as materials can be both strong and brittle (high stress required for failure but abrupt when it occurs). The class supports harvest timing optimization and mechanical damage prediction through brittleness characterization.
Brüchigkeit
Fragilität
Sprödigkeit
Zerbrechlichkeit
crack susceptibility
fracture tendency
fragility
friability
Niklas KJ (1992) Plant Biomechanics: An Engineering Approach to Plant Form and Function. University of Chicago Press, Chicago, pp. 241-287 Pezzi F, Caprara C (2009) Mechanical grape harvesting: investigation of the transmission of vibrations. Biosystems Engineering 103(3): 281-286
Brittleness Quality
Brix acid ratio tracking through ripening in Riesling for different wine styles reveals the style-dependent target application, with dry Riesling production targeting ratio of 3.2-3.5 (achieved at 21 Brix and TA 6.5 g/L, preserving acid backbone for ageability), off-dry production targeting 3.8-4.2 (achieved at 23 Brix and TA 5.8 g/L, balancing residual sugar potential with adequate acidity), and late harvest production accepting ratios >5.0 (28+ Brix with declining TA, sweetness-dominated balance), demonstrating how Brix/TA ratio targets vary with intended wine style and provide style-appropriate harvest timing guidance.
Brix acid ratio quality is a maturity index quality that represents the calculated relationship between sugar concentration (Brix or similar measure) and titratable acidity, characterizing the fundamental balance between sweetness potential and sourness that has historically served as the primary quantitative index for harvest timing decisions and continues to provide a simple, accessible indicator of basic must composition balance. Within the BFO framework, this quality specifically addresses the sugar-acid balance dimension of maturity indices, distinguishing this basic ratio from more complex multi-parameter indices while recognizing that the Brix/TA ratio captures the most fundamental compositional balance affecting wine style, with high ratios indicating ripe/soft character and low ratios indicating unripe/tart character that bracket the optimal range for different wine styles. The ontological significance lies in the Brix/TA ratio providing the most accessible and historically validated single index of compositional maturity, where the ratio directly predicts the perceived balance between alcohol-derived sweetness and acid-derived tartness in dry wines (or actual sugar-acid balance in wines with residual sugar), with target ratios varying by wine style (lower for sparkling and crisp whites, higher for rich reds) but providing consistent relative indication of ripening progress within any context. The calculation and interpretation of Brix/TA ratio involves dividing sugar concentration (typically °Brix) by titratable acidity (typically g/L as tartaric acid equivalents), yielding values that typically range from 2.5 (unripe, tart) through 3.0-4.0 (balanced, style-dependent target range) to 5.0+ (very ripe, potentially unbalanced), with the ratio increasing during ripening as sugar rises and acid declines, enabling tracking of progress toward target balance. The wine style associations with Brix/TA targets include sparkling wine production favoring lower ratios (2.5-3.0, preserving acidity for freshness and aging), crisp white wines targeting moderate ratios (3.0-3.5, balanced with acid emphasis), fuller whites and lighter reds at intermediate ratios (3.5-4.0), and rich red wines tolerating higher ratios (4.0-4.5, favoring roundness over tartness), with these style-specific targets providing the interpretive framework for ratio assessment. The limitations of Brix/TA ratio include its failure to capture phenolic maturity (potentially misleading for red wine quality assessment), its inability to distinguish among acids (high malic versus high tartaric acids have different sensory implications despite similar TA), and its variation with measurement methodology (different TA endpoints affecting calculated ratios), requiring complementation with other indices for comprehensive maturity assessment. Understanding Brix/TA ratio quality enables appropriate application of this fundamental index for basic balance assessment, target setting based on wine style objectives, and recognition of both its utility and limitations as a single-parameter maturity indicator.
Brix_Acid_Ratio_Quality addresses the fundamental sugar-acid balance index historically central to harvest decisions. This quality provides accessible indication of basic compositional balance varying by wine style. The class supports style-appropriate harvest timing through Brix/TA characterization with recognized limitations for phenolic maturity assessment.
Balanceindex
Brix-Säure-Verhältnis
Brix/TA
Reifeverhältnis
Zucker-Säure-Verhältnis
balance index
maturity ratio
sugar-acid ratio
Amerine MA, Winkler AJ (1944) Composition and quality of musts and wines of California grapes. Hilgardia 15(6): 493-675 Jackson RS (2014) Wine Science: Principles and Applications, 4th Ed. Academic Press, San Diego, pp. 232-298
Brix Acid Ratio Quality
Brix measurement for Cabernet Sauvignon at Rutherford, Napa Valley tracked from véraison through harvest (2023 vintage) progressed from 8.5° Brix at 50% véraison (August 5), through 18.0° Brix at the onset of rapid accumulation (August 22), to 24.8° Brix at harvest (September 28), with the target window of 24-26° Brix (predicting 14.0-15.2% potential alcohol) defining the optimal harvest date for full-bodied red wine production. Brix measurement for Sauvignon Blanc in Marlborough, New Zealand documents harvest at 21.5-23.0° Brix (March 2024), with the lower end of the range targeted for aromatic, crisp-style wines (12.5-13.5% alcohol) and the upper end for richer, barrel-fermented styles, demonstrating how Brix targets vary with intended wine style within a single cultivar and region.
Brix measurement is a sugar content data entity that records the total soluble solids concentration in grape must as determined by refractometry or density measurement, expressed in degrees Brix (°Bx), where one degree Brix equals one gram of sucrose per 100 grams of solution at 20°C, serving as the predominant sugar content scale in viticultural regions of the Americas, Australasia, and South Africa, and providing the basis for harvest decisions, grape purchasing contracts, and potential alcohol estimation in these regions.
The Brix scale measures total soluble solids rather than sugar specifically, meaning that non-sugar solutes (organic acids, minerals, phenolics, amino acids) contribute to the reading and create a systematic overestimation of true sugar content, typically by 1-3° Brix depending on must composition. This overestimation is most significant in early-season sampling (before véraison, when sugar-to-acid ratios are low) and in musts from heavily botrytized or raisined fruit where non-sugar dissolved solids are concentrated. Two primary measurement methods are employed — optical refractometry (handheld or digital instruments measuring refractive index, convenient for field use on individual berry or cluster samples) and density-based hydrometry (laboratory measurement of must density using pycnometry or digital density meters, more accurate but requiring larger expressed juice volumes). Temperature correction is essential for refractometric measurement, as refractive index varies with temperature — most instruments include automatic temperature compensation to the reference temperature of 20°C. The Brix scale was originally calibrated by Adolf Brix in the 19th century for the sugar beet industry and adopted by viticulture for its convenience, though the underlying sucrose calibration introduces minor theoretical discrepancies when applied to glucose-fructose solutions typical of grape must. In grape purchasing systems, Brix measurement frequently serves as the primary determinant of grape price, with premium bonuses for fruit exceeding minimum Brix thresholds and penalties for under-ripe deliveries, making measurement accuracy a matter of direct economic consequence.
Brix degree
Brix reading
Brix value
Brix-Bestimmung
Brix-Grad
Brix-Messung
Brix-Wert
Refraktometerwert
refractometer reading
soluble solids measurement
Iland P et al. (2011) Chemical Analysis of Grapes and Wine, 2nd Ed, Patrick Iland Wine Promotions, pp.: 45-52 OIV (2021) Compendium of International Methods of Analysis, Method OIV-MA-AS2-02 Boulton RB et al. (1999) Principles and Practices of Winemaking, Springer, pp.: 13-28
Brix Measurement
Brix quality tracking in Sonoma County Pinot Noir employs pre-dawn sampling to minimize temperature effects and diurnal sugar fluctuations, with measurements beginning at 18°Brix (typically 4 weeks pre-harvest) and continuing at 2-day intervals until harvest targets of 23.5-25.0°Brix are achieved the rate of Brix accumulation (typically 0.2-0.4°Brix per day during active ripening) enables harvest date prediction and crew scheduling, while deviations from expected accumulation curves signal irrigation adjustments or identify blocks requiring earlier or later picking than historical patterns suggest.
Brix_Quality represents the chemical quality characterizing the total soluble solids concentration in grape juice or must, expressed as grams of sucrose per 100 grams of solution (degrees Brix, °Brix). Although named for Adolf Brix who refined the measurement scale in the 19th century, the Brix scale has become the dominant sugar measurement standard in viticulture and winemaking throughout the Americas, Australia, and much of the international wine trade. Brix measurement employs refractometry, utilizing the relationship between dissolved solute concentration and refractive index, providing rapid, non-destructive estimation of sugar content with handheld field refractometers or laboratory instruments. In grape juice, approximately 90% of soluble solids are sugars, so Brix provides an excellent practical approximation of sugar concentration despite also including minor contributions from organic acids, phenolics, and other dissolved compounds. Typical Brix values at harvest range from 18-26°Brix for table wines, with sparkling wine base at the lower end (18-21°Brix) and full-bodied red wines at the higher end (24-27°Brix). The relationship between Brix and potential alcohol follows the approximate conversion of Brix × 0.55-0.60 = potential alcohol (% v/v), enabling harvest planning for target wine styles. Brix accumulation follows a characteristic sigmoidal pattern during ripening, with rapid increase during post-véraison cell expansion, gradual slowing as berries approach maximum volume, and potential plateau or decline in late ripening. Temperature-corrected Brix readings account for the effect of sample temperature on refractive index. The simplicity and speed of Brix measurement makes it the most frequently assessed chemical quality parameter in commercial viticulture.
Brix_Quality is positioned within Sugar_Content_Quality and represents the most widely used operational measure of grape sugar content in international viticulture. Brix is technically a measure of soluble solids rather than sugar specifically, but the dominance of sugars among grape juice solutes makes this distinction minor for practical purposes. The relationship between Brix and actual sugar concentration (g/L) follows the approximate conversion of °Brix × 10 = g/L sugar, though this varies slightly with juice composition. Brix measurement by refractometry can be affected by suspended solids and requires clear juice samples alternatively, digital density meters provide Brix readings based on the density-Brix relationship.
Brix degrees
Brixgrad
degrees Brix
lösliche Feststoffe
refractometric sugar
refraktometrischer Zuckergehalt
soluble solids content
Amerine MA and Ough CS (1980) Methods for Analysis of Musts and Wines, John Wiley and Sons pp.: 176-195 Iland P et al. (2004) Chemical Analysis of Grapes and Wine: Techniques and Concepts, Patrick Iland Wine Promotions pp.: 32-43
Brix Quality
In a Willamette Valley Pinot Noir vineyard, Brix stabilization was confirmed over 16 days in late September Year N+1 (BBCH 89) when twice-weekly refractometer measurements showed: 22.8°, 23.2°, 22.6°, 23.0°, 23.4°, 22.9° (mean 23.0° ± 0.3°, CV 1.3%), linear regression slope -0.008° per day (p = 0.62, not significant), confirming stability enabling harvest timing optimization for phenolic maturity (achieved day 12) and weather window (rain forecast day 18), under cool Oregon conditions maintaining stable plateau without degradation.
Brix_Stabilization represents the cessation of measurable refractometric changes during sugar plateau when soluble solids concentration remains constant within analytical precision creating the observable signal that harvest maturity has been achieved and providing practical criterion for harvest timing decisions in commercial viticulture. This stabilization occurs during plateau period (BBCH 88-89), typically days 55-75 post-veraison through harvest, and is characterized by sequential Brix measurements showing variation 0.05) confirming no systematic change, and refractive index stabilized at final value 1.3330-1.3430 (20°C reference) corresponding to Brix 20-30° depending on cultivar and conditions. Measurement methodology for confirming stabilization includes refractometer sampling 2-3 times per week during late ripening detecting plateau onset, statistical analysis calculating mean and standard deviation over measurement period with SD <0.3° typically indicating stability, temporal trend analysis using linear or polynomial regression testing for significant slope with p-value threshold 0.05 determining stability versus continued change, and comparison with historical data for cultivar and site establishing expected plateau Brix enabling recognition when stabilization achieved. Biological basis of stabilization includes underlying sugar equilibrium with import balancing consumption creating constant vacuolar content reflected in stable Brix measurement, water balance stabilization with berry expansion ceased and transpiration balanced by minor import maintaining constant fresh weight and thus constant concentration, solute composition reaching final proportions with hexoses, acids, phenolics, minerals all at mature levels producing constant refractive index, and metabolic homeostasis where all ripening processes completed or continuing at maintenance rates without directional compositional changes. Refractometric principles relevant to stabilization demonstrate refractive index n = (speed of light in vacuum) / (speed of light in medium) affected by all dissolved solutes with sugars contributing 85-95% of measured Brix at maturity, Brix scale calibrated for pure sucrose solutions approximating grape juice but imperfectly as glucose, fructose, and other solutes have different refractive indices, temperature dependence requiring standardization to 20°C with correction factor ~0.00025 per °C deviation, and measurement precision typically ±0.1-0.2° Brix for quality refractometers with sample preparation (crushing, filtering) and technique (temperature control, calibration) critical for accuracy. Temporal characteristics of stabilization show transition from declining accumulation rates 0.08-0.15% Brix per day to stabilization <0.03% per day occurring over 5-10 day period, plateau duration typically 10-30 days before harvest though potentially extending 30-60 days if harvest delayed, and occasional slight drift toward lower Brix 0.2-0.5° over 3-4 weeks potentially indicating senescence or dehydration though often within measurement variability. Environmental factors affecting stabilization include temperature stability with consistent conditions maintaining stable Brix while fluctuations may cause apparent variability through differential effects on berry water balance, water availability with adequate irrigation maintaining constant berry hydration and thus Brix stability while deficit may concentrate sugars through dehydration producing apparent increase, rainfall potentially diluting berry content causing Brix decrease though typically minor <0.5-1.5° for moderate precipitation, and disease or stress disrupting stability through metabolic perturbation, degradation, or water balance changes. Cultivar differences in stabilization characteristics show some varieties achieving very stable plateau with Brix variation <±0.2° over 2-4 weeks (Chardonnay, Merlot), others showing more variability ±0.4-0.8° potentially reflecting subtle ongoing processes or measurement challenges (Pinot Noir particularly heterogeneous clusters), and high-Brix varieties often showing stable elevated plateau 26-30° while lower-Brix types plateau 20-24° with similar relative stability. Practical viticulture applications include Brix stabilization as key harvest timing indicator signaling physiological maturity achieved, combination with other parameters (pH, TA, phenolics, flavor, seed color) providing integrated maturity assessment, flexible harvest window during stable plateau enabling optimization for quality factors beyond sugar, and documentation through sampling records demonstrating maturity for quality control and regulatory compliance. Quality assurance uses stabilization confirmation through multiple measurements over time verifying true plateau versus transient pause in accumulation, spatial sampling across vineyard zones ensuring uniform stabilization or identifying heterogeneity requiring selective harvest, comparison with target Brix for intended wine style confirming appropriate level achieved (table wines 20-24°, fuller styles 22-26°, high-alcohol or late-harvest 24-30°+), and integration with contract specifications or winery requirements ensuring delivered fruit meets sugar criteria. Challenges and considerations include measurement variability from sampling (different berries, cluster positions, canopy zones), temperature effects requiring consistent sample handling and measurement conditions, timing of sampling with diurnal variation potentially affecting measurements taken at different times of day, and distinguishing true stability from transient plateau during deceleration phase requiring extended monitoring period confirming sustained stability. Harvest logistics planning uses stabilization detection triggering harvest preparation 1-2 weeks advance, coordination with other vineyard blocks and winery capacity scheduling based on maturity timing, weather monitoring balancing quality optimization during stable plateau against harvest risks (rain, disease, wildlife), and flexibility to adjust timing within stable window based on developing conditions or market factors. Economic implications include achievement of target Brix commanding premium prices in some markets particularly minimum thresholds for appellations or quality designations, high-Brix grapes potentially receiving bonuses for high-alcohol wine production, moderate-Brix grapes preferred for balanced lower-alcohol styles increasingly popular, and documentation of Brix stabilization supporting claims of optimal harvest timing and quality. Research applications include statistical methods for detecting stabilization objectively using time-series analysis and trend detection, modeling approaches predicting stabilization timing based on accumulation trajectory and environmental conditions, comparative studies across cultivars and environments identifying factors controlling plateau Brix and stability, and sensor technology development enabling continuous automated monitoring detecting stabilization without manual sampling.
Brix_Stabilization is distinguished from declining accumulation by showing constant measurement (variation 0.05 for temporal trend. Duration typically 10-30 days plateau. Practical harvest timing indicator. Brix_Stabilization represents observable maturity signal enabling harvest optimization.
Plateau-Brix
Reife-Brix-Erreichung
constant soluble solids
konstante lösliche Feststoffe
maturity Brix achievement
plateau Brix
stabiler Brechungsindex
stable refractive index
Coombe BG (1992) Research on development and ripening of the grape berry Amer J Enol Vitic 43:101-110 Brix plateau Iland P, Bruer N, Edwards G, Weeks S, Wilkes E (2004) Chemical Analysis of Grapes and Wine:Techniques and Concepts Wine Promotions Adelaide pages 22-28 Brix measurement and interpretation Keller M (2020) The Science of Grapevines 3rd Ed pages 227-228 stabilization and harvest timing.
Brix Stabilization
The bronze coloration description of Gewürztraminer documents intense copper-red bronzing (OIV 051:4) persisting on the third through sixth unfolded leaves — the bronzing is uniformly distributed across the entire adaxial blade surface with no green sectors, creating the distinctive reddish-copper young canopy appearance that has historically been one of the primary field identification features for Gewürztraminer and led to its German synonym Roter Traminer (Red Traminer, distinguishing it from the non-bronze Grüner Traminer clone). The bronzing intensity in Gewürztraminer is among the strongest in V. vinifera white cultivars and correlates with the elevated flavonol and anthocyanin production capacity that also contributes to this cultivar's characteristically intense aromatic profile through shared phenylpropanoid pathway precursors.
A young leaf description component that specifically documents the presence, intensity, and distribution of the characteristic bronze coloration observed on the adaxial surface of young expanding grapevine leaves, resulting from the co-expression of anthocyanins, flavonols, and carotenoids in leaf tissue where chlorophyll has not yet accumulated to full photosynthetic levels, providing a highly distinctive cultivar identification character that is particularly useful for distinguishing cultivar groups during the spring growth period.
Bronze_Coloration_Description isolates a specific pigmentation phenomenon that deserves dedicated documentation because of its exceptional diagnostic utility and its complex biochemical basis. The bronze color of young grapevine leaves is not a simple anthocyanin effect (which would produce red or purple tones) but a composite visual phenotype arising from the simultaneous presence of multiple pigment classes in different cellular compartments — anthocyanins (in vacuoles, contributing red), carotenoids (in plastids, contributing yellow-orange), and flavonols (in vacuoles and cell walls, contributing yellow) — viewed against the background of incomplete chlorophyll accumulation (which would mask other pigments when fully developed): the ratio between these pigment pools varies between cultivars, producing the range of bronzing expressions from pale yellow-bronze (flavonol-dominated) through warm copper (balanced anthocyanin-carotenoid) to deep red-bronze (anthocyanin-dominated). The bronzing phenotype has attracted renewed research interest as a potential indicator of stress tolerance and UV radiation adaptation, as the photoprotective pigments underlying bronzing also serve as antioxidants and UV screens that may enhance stress resilience in young expanding tissues.
Bronze-Färbungs-Beschreibung
Jungblatt-Bronzierung
bronze pigmentation record
copper-red leaf coloration
kupferrote Blattfärbung
young leaf bronzing assessment
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Downey MO et al. (2006) Cultural practice and environmental impacts on the flavonoid composition of grapes and wine, American Journal of Enology and Viticulture 57: 257-268
Bronze Coloration Description
Cabernet Sauvignon brush after gentle hand harvest at maturity (BBCH 89) measures 1.5-2.5 mm length, 0.8-1.2 mm diameter, exhibits 3-4 visible vascular bundle traces with lignified xylem vessels 25-35 μm diameter, shows compact fibrous structure with clean separation at abscission plane, and tan to brown coloration indicating complete lignification
A brush is the dried remnant of vascular and supporting tissues that remains attached to a grape berry after separation from the pedicel, consisting of lignified vascular bundle traces, parenchyma cell remnants, and sometimes fragments of receptacle tissue, representing the berry's former connection point to the cluster. The brush develops as part of normal berry anatomy during fruit development, originating from the receptacle region where pedicel vascular bundles branch and enter the berry base, becoming visible as a distinct structure only after berry separation through abscission or mechanical detachment. Brush morphology varies significantly by cultivar and separation method= typical wine grape brushes measure 1-4 mm length and 0.5-2 mm diameter at berry attachment, exhibiting fibrous texture from exposed vascular bundle elements, brown to tan coloration from lignified tissues, and irregular surface topography reflecting vascular branching patterns. Brush anatomical composition includes= remnants of xylem vessels (tracheary elements with lignified secondary walls, typically 15-40 μm diameter), phloem tissue remnants (collapsed sieve tubes, companion cells, and phloem parenchyma), lignified sclerenchyma fibers providing structural support, parenchyma cell remnants from receptacle tissue, and sometimes fragments of exocarp tissue if separation occurred within berry rather than at abscission zone. Brush formation and characteristics depend on separation mechanism= abscission-mediated separation (natural or harvest-induced abscission at mature abscission zone) produces clean, compact brushes with separation occurring at defined abscission plane resulting in 1-3 mm brush length, while mechanical separation (forced berry removal before abscission zone maturation or tearing during harvest) produces larger, more irregular brushes with tissue tearing resulting in 2-5 mm length and potential berry flesh attachment. Brush significance in viticulture and enology includes= visual indicator of berry separation quality (compact uniform brushes indicate gentle separation at abscission zone, large irregular brushes suggest rough handling or premature harvest), potential source of phenolic compounds or off-flavors if brushes remain attached during crushing and fermentation (lignified tissues may contribute woody or astringent notes), relevance to mechanical harvest quality assessment (brush characteristics reflect harvest gentleness and berry maturity), and ampelographic character (brush morphology shows cultivar-specific variation useful for variety identification). Cultivar variation in brush characteristics includes= brush length (1-5 mm depending on variety and separation timing), brush diameter (0.5-2.5 mm), vascular bundle number visible in brush cross-section (2-8 bundles), lignification extent (highly lignified woody brushes versus partially lignified softer brushes), and ease of brush detachment from berry (some cultivars show brushes that separate cleanly leaving minimal remnant, others retain substantial brush tissue).
Brush emphasizes the complete vascular remnant structure and is distinguished from Brush_Attachment_Zone (connection interface) and Vascular_Bundle_Termination (vascular ending structures) by encompassing entire dried tissue complex remaining after berry separation. Critical for harvest quality assessment, understanding berry-pedicel separation mechanisms, ampelographic variety identification, and potential enological implications of brush tissue in crushed must.
Berry brush
Bürste
Gefäßbündelrest
Pedicel remnant
Stielansatz
Galet, P. (2000). A Practical Ampelography= Grapevine Identification. Comstock Publishing Associates. | Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press.
Brush
Pinot Noir brush attachment zone at harvest maturity (BBCH 89) shows receptacle region 0.8-1.5 mm diameter where 3 pedicel vascular bundles diverge into 8-12 berry traces, well-developed abscission zone positioned 0.5-0.8 mm below receptacle center producing clean separation with 0.9-1.3 N detachment force, and resulting compact brush 1.5-2.0 mm length after abscission
A brush attachment zone is the interface region where brush tissues connect to the berry base, encompassing the receptacle area where pedicel vascular bundles terminate and branch into berry vasculature, and where separation occurs during berry abscission or detachment. The brush attachment zone represents a complex anatomical region involving= vascular bundle divergence (pedicel bundles branching into multiple traces entering berry exocarp), receptacle tissue (expanded pedicel terminus with parenchyma and vascular elements forming transition between pedicel and berry), potential abscission zone location (if present, abscission layer forms within or near brush attachment region), and structural integration of pedicel vasculature with berry basal tissues. Brush attachment zone anatomy includes= receptacle parenchyma cells providing structural connection between pedicel and berry, branching vascular bundles transitioning from 2-4 main pedicel bundles to 6-15 berry vascular traces, zone of vascular bundle termination where xylem vessels and phloem sieve tubes end or merge with berry vasculature, supporting sclerenchyma or collenchyma tissues providing mechanical strength, and in mature berries, differentiated abscission layer (if abscission zone develops in this region). Brush attachment zone development proceeds through berry development= during early fruit set (BBCH 71-73) receptacle expands and vascular bundles differentiate establishing berry vascular supply, during berry growth phases (BBCH 73-81) vascular connections remain firm with active transport through attachment zone, during ripening (BBCH 81-89) progressive modifications including potential abscission zone differentiation, vascular activity changes, and tissue lignification. Brush attachment zone characteristics influencing brush formation include= abscission zone position relative to receptacle (abscission within receptacle produces minimal brush, abscission below receptacle produces larger brush including receptacle tissue remnants), vascular bundle architecture (number and branching pattern affecting brush vascular trace visibility), tissue lignification extent (highly lignified attachment zones produce woody persistent brushes), and structural integrity (weak attachment zones may fracture irregularly during separation producing variable brush morphology). Brush attachment zone separation patterns vary= clean abscission separation (occurs at defined abscission layer within attachment zone, produces compact uniform brush, typical of mature berries with developed abscission zones, BBCH 87-89), receptacle fracture (occurs through receptacle tissue above or below abscission zone, produces larger brush with receptacle remnants, may occur during mechanical harvest or forced separation), and irregular tearing (occurs when separation forced through unprepared tissues, produces variable brush with torn tissue fragments, typical of immature berries or rough handling). Cultivar variation in brush attachment zone includes= abscission zone development strength and timing (strong early-developing zones versus weak late-developing zones), receptacle size and vascular complexity (compact simple receptacles versus enlarged complex receptacles), tissue lignification patterns (uniform versus heterogeneous lignification), and attachment strength (firm attachment requiring high detachment force versus weak attachment with low force requirement).
Brush_Attachment_Zone emphasizes the connection interface and is distinguished from Brush (complete remnant structure) and specific attachment subtypes (Pedicel_Brush_Attachment, Seed_Brush_Attachment) by focus on anatomical transition region where separation occurs. Critical for understanding berry separation mechanisms, abscission zone positioning relative to receptacle, brush morphology variation, and harvest quality implications of separation location and pattern.
Beerenbasis-Übergangszone
Berry-pedicel junction
Brush connection zone
Bürstenansatzzone
Stielansatz-Verbindungsbereich
Galet, P. (2000). A Practical Ampelography= Grapevine Identification. Comstock Publishing Associates. | Pratt, C. (1971). Reproductive anatomy in cultivated grapes - a review. American Journal of Enology and Viticulture, 22(2), 92-109.
Brush Attachment Zone
Bubble persistence assessment in recently disgorged versus aged Champagne (same cuvée, different release timing) reveals disgorgement effect on persistence, with recently disgorged showing vigorous, persistent streams lasting 8+ minutes in glass (persistence rating 9/10), while 5-year post-disgorgement bottle displays gentler, shorter-lived streams (4 minutes, persistence 6/10) from gradual CO₂ loss through cork, demonstrating how bubble persistence indicates both initial quality and post-release freshness, guiding optimal consumption timing for aged sparkling wines.
Bubble persistence quality is an effervescence quality that represents the duration and sustainability of bubble release in sparkling wine, characterizing how long effervescence continues both in the glass (visual stream persistence) and in the mouth (sustained tactile stimulation) as an indicator of CO₂ retention, wine quality, and appropriate serving conditions. Within the BFO framework, this quality specifically addresses the temporal aspect of bubble behavior, distinguishing persistence from bubble size while recognizing that long-lasting effervescence indicates both adequate carbonation level and the presence of compounds that stabilize bubble formation over time. The ontological significance lies in bubble persistence serving as both a quality indicator and a practical freshness marker, where sustained effervescence demonstrates proper CO₂ dissolution and retention while quickly fading bubbles may indicate quality compromise from poor sealing, warm storage, or insufficient initial carbonation affecting the sparkling wine experience. The physical basis for bubble persistence involves the balance between CO₂ dissolution equilibrium and the kinetics of bubble nucleation and escape, with persistence enhanced by adequate initial CO₂ pressure (typically 5-6 atmospheres in Champagne), appropriate serving temperature (colder increasing CO₂ solubility), glass design (etched nucleation points sustaining bubble streams), and wine composition (proteins and polysaccharides stabilizing bubble formation). The production and storage factors affecting persistence include carbonation level achieved during secondary fermentation or carbonation process, crown cap or cork seal integrity during aging, storage temperature (warm storage accelerating CO₂ loss), and time since disgorgement or bottling (CO₂ slowly escaping even through quality closures). The assessment of bubble persistence involves observation of bubble streams in glass (timing until cessation or significant reduction) and palate evaluation of sustained fizzing character (duration of tactile effervescence during consumption), with quality sparkling wines showing persistence of several minutes in glass and sustained palate effervescence throughout consumption. Understanding bubble persistence quality enables freshness assessment, storage condition evaluation, and quality judgment incorporating appropriate persistence for sparkling wine style.
Bubble_Persistence_Quality addresses the temporal dimension of effervescence, indicating both quality and freshness in sparkling wine. This quality depends on initial carbonation level, seal integrity, and storage conditions, providing practical information for consumption timing. The class supports freshness assessment and quality evaluation through persistence observation in both glass and mouth.
Blasenpersistenz
Efferveszenz-Ausdauer
Perlage-Dauer
Schaumhaltbarkeit
bubble duration
effervescence longevity
fizz persistence
perlage endurance
Liger-Belair G (2005) The physics and chemistry behind the bubbling properties of Champagne and sparkling wines. Journal of Agricultural and Food Chemistry 53(8): 2788-2802 Kemp B et al. (2015) Effect of production phase on bottle-fermented sparkling wine quality. Journal of Agricultural and Food Chemistry 63(1): 19-38
Bubble Persistence Quality
Bubble size assessment comparing 30-month aged Champagne (bubble diameter approximately 0.4 mm, fineness rating 9/10) with entry-level Crémant (12-month aging, 0.8 mm diameter, fineness 6/10) and tank-method Sekt (1.2 mm diameter, fineness 4/10) demonstrates the aging and method effects on bubble size, with extended tirage releasing yeast mannoproteins that reduce surface tension and stabilize smaller bubble formation, creating the fine, elegant perlage that visually and texturally distinguishes premium from ordinary sparkling wines.
Bubble size quality is an effervescence quality that represents the diameter of CO₂ bubbles forming in sparkling wine, characterizing the visual and tactile dimension of effervescence where fine, small bubbles indicate quality and sophisticated production while coarse, large bubbles suggest simpler sparkling wine styles or carbonation methods. Within the BFO framework, this quality specifically addresses the dimensional aspect of bubble perception, distinguishing bubble size from persistence and mousse character while recognizing that size fundamentally affects both visual elegance in the glass and tactile fineness on the palate. The ontological significance lies in bubble size serving as a primary quality indicator in sparkling wine, where fine bubbles are universally preferred for their elegant visual display and delicate tactile impression, with bubble fineness correlating strongly with production method sophistication, aging duration, and overall quality tier across sparkling wine categories. The physical basis for bubble size involves nucleation dynamics and surface tension effects, with bubbles nucleating from surface imperfections and dissolved particles (nucleation sites), growing as CO₂ diffuses into them, and detaching when buoyancy overcomes surface tension, with smaller bubbles resulting from more nucleation sites producing more numerous smaller bubbles and from higher protein and polysaccharide content reducing surface tension and stabilizing smaller bubble formation. The production method associations of bubble size are strong, with méthode traditionnelle (secondary fermentation in bottle with extended lees aging) producing the finest bubbles due to elevated protein content and prolonged CO₂ saturation, while Charmat tank method and carbonation produce progressively larger bubbles due to less favorable formation conditions. The aging effect on bubble size shows improvement with extended tirage in traditional method wines as autolysis releases surfactant proteins that stabilize smaller bubbles, explaining the exceptional fineness achieved in aged Champagne and long-aged Cavas. Understanding bubble size quality enables sparkling wine quality assessment, production method inference from bubble character, and quality tier differentiation based on bubble fineness.
Bubble_Size_Quality addresses the dimensional aspect of effervescence, with fine bubbles universally indicating quality in sparkling wine. This quality correlates with production method sophistication and aging duration, providing a reliable quality indicator. The class supports sparkling wine assessment and production method inference through bubble size evaluation.
Blasengröße
Bläschendurchmesser
Perlage-Größe
Perlfeinheit
bead size
bubble diameter
bubble fineness
perlage size
Liger-Belair G et al. (2008) Study of effervescence in a glass of champagne. Journal of Physical Chemistry B 112(14): 4254-4265 Cilindre C et al. (2007) Evidence for the underlying proteins related to foam formation in Champagne base wines. Journal of Agricultural and Food Chemistry 55(5): 1595-1600
Bubble Size Quality
A compound bud at node position 5 on a mature cane of Vitis vinifera cv. Cabernet Sauvignon, containing visible primary and secondary primordia when dissected longitudinally, showing differentiated inflorescence initials in the primary bud and vegetative initials in the secondary positions.
A dormant or actively growing vegetative or reproductive structure in Vitis vinifera consisting of an embryonic shoot with compressed internodes, protected by modified leaves called bud scales, capable of developing into shoots, inflorescences, or both depending on primordial content and environmental conditions. Grapevine buds exhibit a compound structure unique among fruit crops, typically containing primary, secondary, and tertiary meristematic units within a single bud, each capable of independent development following damage or environmental stress to other components.
The grapevine bud differs fundamentally from simple buds in other plant species due to its compound nature, with multiple independent growing points housed within a single structure. Classification schemes distinguish buds by dormancy state (dormant vs. active), position (basal, lateral, apical), developmental potential (vegetative, reproductive, mixed), and hierarchical position within the compound structure (primary, secondary, tertiary).
Knospe
Rebknospe
grapevine bud
vine bud
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press Carmona et al. (2008) "Grapevine bud dormancy= environmental and molecular aspects" Revista Brasileira de Fruticultura 30=257-408.
Bud
Bud break synchrony assessment comparing cane-pruned versus spur-pruned Riesling reveals the pruning system effect on emergence uniformity, with cane pruning (long cane laid horizontally) showing moderate synchrony (bud break spanning 8 days from first to last emergence due to position effects along cane), while spur pruning (multiple short spurs at similar positions) shows higher synchrony (bud break spanning only 4 days due to positional uniformity), with the synchrony difference affecting subsequent canopy uniformity and the feasibility of single-pass shoot management operations.
Bud break synchrony quality is an intra-plant synchrony quality that represents the temporal coordination of bud break among different buds within a single grapevine, characterizing the uniformity of dormancy release and initial shoot emergence that establishes the foundation for subsequent developmental synchrony throughout the growing season and determines the homogeneity of shoot populations available for canopy management. Within the BFO framework, this quality specifically addresses the bud break dimension of intra-plant synchrony, distinguishing early-season emergence coordination from later phenological events while recognizing that bud break synchrony establishes the initial conditions affecting synchrony of all subsequent stages, as shoots emerging at different times will progress through later stages at different times unless differential growth rates compensate. The ontological significance lies in bud break synchrony establishing the baseline developmental coordination that propagates through the season, where synchronized bud break produces cohorts of shoots at similar developmental stages enabling uniform canopy management operations, while asynchronous bud break creates shoot populations spanning multiple developmental stages that complicate thinning, positioning, and other canopy operations that are optimally performed at specific shoot stages. The factors affecting bud break synchrony include chilling uniformity (all buds receiving consistent chilling enabling uniform dormancy release), bud position (distal buds sometimes breaking earlier due to reduced apical dominance, basal buds sometimes delayed), pruning wound effects (buds near pruning wounds potentially delayed by wound response), and microclimate uniformity (temperature variation across the vine potentially affecting local bud break timing). The acrotony pattern in grapevine, where distal buds break before basal buds in unpruned canes, requires pruning to redirect growth to basal buds, with the success of this redirection affecting bud break synchrony as compensatory growth of basal buds may lag the natural tendency for distal emergence. The management implications of bud break synchrony include shoot thinning timing (synchronized break enabling single-pass thinning at optimal stage), frost risk assessment (spread bud break potentially providing risk mitigation through developmental heterogeneity), and canopy uniformity establishment (synchronized break facilitating uniform shoot development and subsequent management). Understanding bud break synchrony quality enables assessment of early-season developmental uniformity, prediction of subsequent synchrony patterns based on initial conditions, and management planning that accounts for the degree of shoot population homogeneity.
Bud_Break_Synchrony_Quality addresses the coordination of emergence timing among buds within single vines. This quality establishes the foundation for subsequent seasonal synchrony through initial shoot population uniformity. The class supports early-season management planning through bud break synchrony characterization.
Austriebssynchronie
Frühjahrsentwicklungskoordination
Knospenbruchuniformität
Triebinitiierungsgleichmäßigkeit
budburst uniformity
emergence synchrony
shoot initiation uniformity
spring development coordination
Winkler AJ et al. (1974) General Viticulture. University of California Press, Berkeley, pp. 138-165 Friend AP, Trought MCT (2007) Delayed winter spur-pruning in New Zealand can alter yield components of Merlot grapevines. Australian Journal of Grape and Wine Research 13(3): 157-164
Bud Break Synchrony Quality
Cabernet Sauvignon in Napa Valley shows bud breaking at BBCH 07-09 in mid-April after 1050-1200 chilling hours and 320-380 growing degree days base 10°C, with green shoot tip emerging and extending 15-25 mm over 9-11 days under conditions of 16-20°C mean temperature, 50-60% relative humidity, and 14-hour photoperiod, representing typical warm-climate timing for this late-season cultivar
Bud breaking represents the critical transitional process where protective bud scales separate completely allowing green shoot tissue to emerge and initiate independent growth, marking the definitive exit from dormancy and beginning of vegetative development. This fundamental developmental process corresponds to BBCH stages 05-09, occurring when internal shoot primordium achieves sufficient size and turgor pressure to mechanically force scales apart and extend beyond protective structures. The process progresses from initial emergence of green shoot tip through scales at BBCH 07 to complete separation of first leaves from shoot axis at BBCH 09, typically requiring 7-12 days depending on temperature and cultivar. Breaking initiates when expanding shoot primordium reaches 5-8 mm length with first leaf primordia 3-5 mm, generating mechanical force of 0.5-1.2 MPa sufficient to overcome resistance of bud scale attachments. The green shoot tip, often called "green tip," first appears at bud apex where scales spread widest, initially showing pale yellow-green coloration due to limited chlorophyll in enclosed tissues. As emergence progresses, shoot extends 10-20 mm beyond scales with increasing chlorophyll synthesis producing characteristic green coloration and photosynthetic capacity beginning to develop. The protective bud scales separate progressively from tip toward base, eventually falling away or remaining attached at bud base as dry brown remnants while shoot continues elongating. Cell division rates remain high in shoot apical meristem producing new cells at 18-24 hour intervals, while cells behind meristem undergo rapid expansion through water uptake and wall loosening, creating visible shoot elongation of 2-5 mm per day. First true leaves unfold from their compressed primordia state, initially remaining appressed to shoot axis then progressively spreading away to establish characteristic palmate leaf configuration. Vascular tissues extend through elongating shoot establishing continuous xylem and phloem connections from permanent woody structures through new shoot tissues to emerging leaves, enabling water transport and nutrient distribution. Metabolic transition occurs from dependence on stored carbohydrate reserves to beginning photosynthetic carbon fixation as leaf area develops, though stored reserves remain primary carbon source until several leaves fully expand. Respiration rates remain elevated at 12-18 fold above dormancy levels supporting rapid growth, while photosynthetic capacity increases gradually as chloroplasts develop and leaf area expands. Hormonal regulation shows sustained high cytokinin and gibberellin concentrations promoting continued cell division and expansion, with auxin production in shoot apex establishing apical dominance suppressing lateral bud activation and coordinating leaf and shoot development. Temperature remains primary rate-determining factor with development doubling for each 10°C increase between 10-25°C base temperatures, making spring temperature patterns critically important for breaking progression rate. Water availability becomes increasingly important as transpiring leaf area develops, with water stress potentially slowing or arresting breaking process if root uptake cannot meet shoot demand. Light quality affects breaking progression with red: far-red ratios influencing elongation rate and leaf expansion, though light intensity effects are minimal until photosynthetic capacity develops. The process exhibits substantial cultivar variation in duration and environmental sensitivity, with early varieties like Chardonnay and Pinot Noir completing breaking in 7-9 days while late varieties like Riesling and Mourvèdre require 10-14 days under identical conditions. Regional timing follows late bud swelling by 3-5 days, occurring mid-to-late April in warm climates, late April to early May in temperate regions, and mid-May in cool continental areas. Viticultural management considerations include completing all dormant-season vineyard operations before breaking as mechanical damage risk increases substantially with fragile emerging tissues, deploying frost protection measures as breaking represents period of highest freeze vulnerability with tender tissues highly susceptible to cold damage, implementing early-season pest and disease management programs targeting newly available host tissues, and monitoring breaking uniformity within blocks to assess crop potential and identify areas requiring adjusted management. Frost damage during breaking causes immediate tissue death through ice formation in highly hydrated cells, killing primary shoots and forcing compensatory growth from secondary or basal buds with 2-3 week delays and substantially reduced fruitfulness. Research applications include phenological modeling incorporating breaking as key developmental milestone for growing season initiation, molecular analysis of genes regulating shoot emergence and organ differentiation, cellular and developmental biology studies of meristem activity and organ formation, biomechanical analysis of forces involved in scale separation and shoot emergence, and climate change impact assessment examining historical trends and future projections for breaking timing across viticultural regions.
Bud_Breaking follows Bud_Swelling and represents active emergence phase rather than expansion phase. Distinguished by green tissue extending beyond protective scales. Encompasses Initial_Bud_Breaking, Active_Bud_Breaking, and Complete_Bud_Breaking as progression subphases. Transitions to Shoot_Growth after BBCH 09 when leaves separate from axis. Most frost-vulnerable period of entire growing season.
Austriebsbeginn
Grünspitzenstadium
Knospenaufbruch
Triebaustrieb
bud break
budbreak
green tip emergence
shoot emergence
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 175-189 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 78-89 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Jones GV & Davis RE (2000) Climate influences on grapevine phenology, grape composition, and wine production and quality for Bordeaux, France, Am J Enol Vitic 51:249-261
Bud Breaking
Bud browning quality evaluation following a late February freeze event (-26°C) in Finger Lakes Riesling provided the basis for adjusted pruning decisions to optimize crop potential from surviving buds. Systematic sampling of 100 buds per block with longitudinal sectioning and visual browning assessment quantified the damage extent across bud positions and vineyard locations. Primary bud browning quality showed 55% mortality across all samples, with the brown, oxidized appearance of dead primordia clearly distinguishable from the green, turgid appearance of surviving tissue. Secondary bud browning quality showed only 12% mortality, confirming the expected 3-4°C hardiness advantage providing substantial backup survival. Tertiary buds showed less than 3% browning, indicating essentially complete survival of this backup tier. Positional analysis revealed bud browning quality gradient along canes, with nodes 1-4 showing 40% primary mortality while nodes 8-12 showed 70% mortality, reflecting the greater hardiness of basal buds and informing recommendations for shorter pruning to retain the more hardy basal positions. Topographic analysis showed bud browning quality of 75% primary mortality in valley-floor frost pockets versus 35% on upper slopes, guiding block-specific pruning adjustments. The bud browning quality assessment enabled yield projections accounting for primary bud losses (compensated partially by secondary bud fruitfulness at 60% of primary) and informed insurance claims with documented damage quantification.
Bud browning quality is a cold damage symptom quality that inheres in grapevine dormant buds and represents the extent and pattern of oxidative tissue discoloration visible in cross-sectioned buds following freezing injury, with the characteristic brown coloration of killed bud primordia providing the standard diagnostic indicator for assessing bud mortality and predicting spring shoot emergence potential following winter freeze events. This quality addresses the most widely used method for rapid cold damage assessment in vineyards, recognizing that bud survival directly determines spring growth capacity such that bud browning evaluation provides essential information for predicting yield potential and adjusting pruning strategies to compensate for bud losses. The ontological significance of bud browning quality lies in its practical utility for damage assessment, with the cross-section browning test providing a simple, rapid, and reliable method for quantifying bud mortality that can be performed in the field with minimal equipment, enabling extensive sampling to characterize damage patterns across vineyards and inform management decisions within days of freeze events. The biological basis of bud browning encompasses the oxidation of phenolic compounds and other cellular constituents released from damaged cells when membrane integrity is lost during freezing, with the enzymatic and non-enzymatic oxidation reactions producing the characteristic brown pigmentation that distinguishes dead from living tissue. The compound bud structure creates characteristic browning patterns where primary, secondary, and tertiary bud primordia may show differential damage reflecting their distinct hardiness levels, with primary bud browning commonly occurring while secondary buds remain green following marginal freeze events that exceed primary bud hardiness but not secondary bud tolerance. The timing of browning assessment affects accuracy, with evaluation recommended at least 48-72 hours after thawing to allow complete oxidation development, while assessment too long after damage may be complicated by tissue desiccation or secondary infection that modifies appearance. The spatial sampling protocol for bud browning assessment must account for position effects along the cane (basal buds typically more hardy than distal) and within the vineyard (frost pocket versus elevated positions), with representative sampling across these gradients required for accurate whole-vineyard damage estimation.
Bud_Browning_Quality is_a Cold_Damage_Symptom_Quality, representing the standard diagnostic technique for assessing bud mortality through cross-section browning evaluation. This class addresses the most important tissue for cold damage assessment given direct yield impact. The primary-secondary-tertiary differential provides backup system information. Basal-to-distal gradient along canes informs pruning position decisions. Assessment timing (48-72 hours post-thaw) critical for accurate browning development. Representative sampling protocol essential for whole-vineyard damage estimation.
Knospen-Frostschaden
Knospenmortalitäts-Bewertung
Knospennekrose
Knospenverbräunung
bud freeze damage
bud kill evaluation
bud mortality assessment
bud necrosis
Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection. Michigan State University Extension Bulletin E2930 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Pool RM et al. (1990) Assessing and predicting cold injury to grapevines. Proceedings of the New York Wine Industry Workshop, pp. 17-25
Bud Browning Quality
Pinot Noir in Burgundy initiates bud burst at BBCH 01-09 in early April after accumulating 850-950 chilling hours and 150-200 growing degree days base 10°C, with process completion requiring 15-21 days under typical spring conditions of 12-16°C mean temperature and increasing from 12 to 14 hours daylength, representing critical frost-risk period for this early-bursting cultivar in continental climate
Bud burst represents the fundamental phenological transition marking dormancy exit and active growth initiation in grapevine, encompassing the progressive morphological and physiological changes from winter bud to emerging shoot. This critical developmental process spans BBCH stages 00-09, typically occurring when sustained temperatures exceed 10°C and accumulated chilling requirements have been met. The process initiates with bud swelling as dormant meristematic tissues resume metabolic activity, progresses through scales separating and protective tissues breaking, and culminates with green tissue emergence and first leaf unfolding. Bud burst timing fundamentally determines growing season length, frost exposure risk, and harvest timing. The process involves coordinated activation of cell division and expansion in apical and axillary meristems, mobilization of stored carbohydrate reserves from permanent woody structures, resumption of vascular cambium activity connecting bud to cane, and establishment of bidirectional nutrient transport. Water uptake increases dramatically as cells expand, creating turgor pressure that mechanically forces protective bud scales apart. Hormonal regulation centers on declining abscisic acid levels from dormancy and rising cytokinin and gibberellin concentrations promoting growth. Environmental triggers include cumulative heat accumulation measured as growing degree days, photoperiod lengthening providing developmental cues, and soil temperature affecting root activity and water uptake. The process exhibits substantial cultivar variation, with early-bursting varieties like Chardonnay and Pinot Noir initiating growth 7-14 days before late-bursting varieties like Riesling and Cabernet Sauvignon under identical conditions. This variation reflects genetic differences in chilling requirements ranging from 200-1400 hours below 7°C and base temperature thresholds for heat accumulation. Regional timing varies from late March in warm climates to mid-May in cool regions, with spring frost risk inversely related to burst timing. Viticultural management critically depends on accurate burst prediction, as protective measures against spring frost, early-season pest and disease management, and canopy management operations must be timed relative to developmental stage. Bud burst uniformity within vineyards affects subsequent crop load management and harvest decisions. Climate change has advanced burst timing by 1-2 weeks in many regions over recent decades, increasing frost exposure in some areas while extending growing seasons in others. Research applications include phenological modeling for climate adaptation, molecular analysis of dormancy release mechanisms, and breeding for optimized burst timing relative to regional climate patterns.
Bud_Burst encompasses the entire dormancy-to-growth transition including Bud_Swelling and Bud_Breaking as constituent processes. Distinguished from Shoot_Growth which begins after BBCH 09 when leaves separate from shoot axis. Timing relative to other phenological events determines frost risk, pest pressure, and growing season duration. Regional terminology varies with some sources using "bud break" specifically for BBCH 07 while others apply it to the entire 00-09 period.
Austrieb
Frühjahrsaustrieb
Knospenaustrieb
bud sprouting
spring bud burst
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 157-189 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 63-89 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Fennell A & Hoover E (1991) Photoperiod influences growth, bud dormancy, and cold acclimation in Vitis species, J Am Soc Hort Sci 116:270-273
Bud Burst
Detailed characterization of Bud_Cold_Hardiness_Disposition at the Cool Climate Oenology and Viticulture Institute, Brock University, employed differential thermal analysis to separately determine LT50 values for primary, secondary, and tertiary bud components across 20 cultivars through the dormant season. At maximum hardiness in January, primary bud LT50 ranged from -17.8°C ('Merlot') to -36.2°C ('Frontenac'), while secondary buds were consistently 2.5-4.0°C more hardy and tertiary buds 4.5-6.5°C more hardy than primary buds within each cultivar. For 'Riesling' (primary LT50 -23.5°C, secondary -26.8°C, tertiary -28.9°C), a -26°C midwinter event would kill approximately 50% of primary buds while leaving secondary and tertiary buds largely intact, enabling 70-80% yield recovery through secondary shoot growth. Field validation over 6 winters confirmed that DTA-predicted Bud_Cold_Hardiness_Disposition accurately matched observed field survival (r = 0.91 between predicted and actual mortality). Economic modeling demonstrated that cultivars with primary bud LT50 below -25°C showed positive net returns in the Niagara Peninsula climate, while cultivars above -22°C required supplemental winter protection (hilling, geotextile covers) costing $800-1,200/ha annually. These findings establish Bud_Cold_Hardiness_Disposition thresholds for economically sustainable cold-climate viticulture.
A Bud_Cold_Hardiness_Disposition is a Midwinter_Hardiness_Disposition that inheres specifically in dormant grapevine buds and is realized through the capacity of bud tissues to survive freezing temperatures through deep supercooling of bud primordia and/or tolerance to ice formation in bud scales and adjacent tissues. This disposition represents the most economically critical component of grapevine cold hardiness, as bud survival determines the following season's shoot number, cluster number, and ultimately crop yield. Grapevine buds survive freezing primarily through supercooling—the maintenance of water in liquid state below its equilibrium freezing point—rather than through freeze tolerance, with bud primordia remaining ice-free while surrounding tissues freeze. The supercooling capacity of grapevine buds is limited by ice nucleation events that trigger rapid freezing, detectable as low-temperature exotherms (LTEs) in differential thermal analysis, with the LTE temperature correlating with bud lethal temperature. The compound bud structure of grapevine, containing primary, secondary, and tertiary bud primordia within a common bud scale envelope, creates a hierarchy of cold hardiness with important implications for crop recovery following winter injury. Primary buds are typically least hardy but most fruitful, secondary buds more hardy but less fruitful, and tertiary buds most hardy but minimally fruitful, enabling partial crop recovery when primary buds are killed but secondary or tertiary buds survive. Bud_Cold_Hardiness_Disposition is the primary selection target in cold-climate grape breeding and the key parameter for cultivar recommendations in regions with severe winters.
Bud_Cold_Hardiness_Disposition is_a Midwinter_Hardiness_Disposition. Bud cold hardiness is economically the most critical tissue-specific hardiness because bud survival determines next season's crop. Child classes address the three bud components within the compound bud structure: Primary_Bud_Hardiness_Disposition, Secondary_Bud_Hardiness_Disposition, and Tertiary_Bud_Hardiness_Disposition. Grapevine buds survive freezing primarily through supercooling rather than freeze tolerance, with low-temperature exotherms (LTEs) in differential thermal analysis marking lethal ice nucleation events. The differential hardiness among bud components enables partial crop recovery following intermediate freeze events.
Knospen-Frosthärte-Disposition
Knospen-Kältetoleranz
Schlafende-Knospen-Härte
bud LT50
bud freeze tolerance
dormant bud hardiness
Andrews PK, Sandidge CR, Toyama TK (1984) Deep supercooling of dormant and deacclimating Vitis buds, American Journal of Enology and Viticulture 35:175-177 Quamme HA (1986) Use of thermal analysis to measure freezing resistance of grape buds, Canadian Journal of Plant Science 66:945-952 Ferguson JC et al. (2011) Dynamic thermal profiles of grapevine cold hardiness, American Journal of Enology and Viticulture 62:407-413
Bud Cold Hardiness Disposition
In Chardonnay vines in Champagne, bud competence restoration is completed by mid-February following accumulation of approximately 1000 chilling hours, with restoration verified through forcing tests showing budbreak within 11 days, molecular analysis confirming restored CYCLIN D and CDK expression potential, and ultrastructural examination showing meristematic cells with characteristics of division-ready G1 phase including prominent nucleoli and dispersed chromatin.
Bud competence restoration in Vitis vinifera represents the comprehensive re-establishment of growth capability in dormant buds following chilling-dependent dormancy release, encompassing the restoration of all cellular, molecular, and physiological capacities required for meristematic activity and developmental growth upon receipt of appropriate environmental signals. This restoration process occurs through the final phases of endodormancy and early ecodormancy as chilling-dependent changes complete the dismantling of dormancy-maintaining mechanisms and prepare buds for growth response to spring warming. The cellular dimension of competence restoration involves the re-establishment of cell cycle readiness in meristematic cells, with restoration of cyclin expression potential, degradation of CDK inhibitors that blocked division during dormancy, and chromatin remodeling that makes cell cycle gene promoters accessible for transcription upon growth signal reception. The hormonal dimension encompasses the restoration of growth hormone signaling systems, including sensitization of cytokinin receptors that promote cell division, restoration of gibberellin response capacity that enables cell elongation, and re-establishment of auxin polar transport that organizes growth patterns, while ABA signaling capacity that inhibited growth during dormancy becomes reduced. The metabolic dimension involves the restoration of biosynthetic pathway capacity to support the rapid synthesis of proteins, nucleic acids, cell wall materials, and membranes required for growth, with enzyme systems prepared for rapid activation upon growth initiation even while remaining at low activity during ecodormant quiescence. The connectivity dimension addresses the restoration of symplastic communication through callose degradation at plasmodesmata, re-establishing the cell-to-cell signaling required for coordinated growth responses across meristematic tissues. The verification of competence restoration relies on forcing tests demonstrating rapid budbreak, with restored competence indicated by budbreak times below 14 days and high success rates, and on molecular markers confirming the completion of dormancy-releasing changes at the gene expression level. The completeness of competence restoration affects the quality of subsequent growth, with full restoration supporting vigorous uniform budbreak while incomplete restoration may result in weak or delayed growth.
Bud_Competence_Restoration represents the second component phase of Dormancy_Release addressing comprehensive re-establishment of growth capability. Distinguished from Competence_Achievement which marks the endpoint, while Bud_Competence_Restoration addresses the process encompassing cellular, hormonal, metabolic, and connectivity dimensions. Subdivides into Growth_Capacity_Restoration, Metabolic_Reactivation_Potential, and Budbreak_Readiness as component aspects. Verified through forcing tests and molecular markers. Completeness affects subsequent growth vigor and uniformity.
Knospen-Kapazitäts-Wiedererrichtung
Knospenkompetenz-Wiederherstellung
Knospenwachstumsfähigkeit-Restauration
Meristemkompetenz-Erholung
bud capacity re-establishment
bud growth competence restoration
dormant bud reactivation potential
meristem competence recovery
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Rohde A, Bhalerao RP (2007) Plant dormancy in the perennial context. Trends in Plant Science 12: 217-223 Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle
Bud Competence Restoration
Bud development rate comparison among cultivars following adequate chilling reveals the genetic variation affecting budbreak timing and frost vulnerability, with Chardonnay showing rapid bud development (150 growing degree-days base 10°C from February 1 to BBCH 09 budbreak), Merlot intermediate (195 GDD to budbreak), and Cabernet Sauvignon slow (240 GDD to budbreak), with the 60% difference in thermal time requirement translating to 10-18 days difference in budbreak date depending on spring temperatures, demonstrating how bud development rate variation provides natural frost avoidance in late-developing cultivars.
Bud development rate quality is a developmental rate quality that represents the speed of bud progression from dormancy through swelling, woolly bud, green tip, and leaf emergence stages, characterizing the rate of dormancy release and early shoot initiation that determines budbreak timing and the synchronization of early growth with spring conditions affecting frost risk and initial canopy establishment. Within the BFO framework, this quality specifically addresses the bud developmental dimension of phenological rate, distinguishing bud progression rate from subsequent shoot development while recognizing that bud development rate during the critical dormancy-to-budbreak transition establishes the timing of all subsequent phenological events through its determination of when active growth commences. The ontological significance lies in bud development rate determining the earliest vulnerable phenological stages, where rapid bud development following warm late-winter periods can advance budbreak into frost-risk windows, while slow development provides frost avoidance through delayed exposure of tender tissues, making bud development rate a primary determinant of spring frost vulnerability that varies with cultivar genetics and dormancy fulfillment status. The regulatory basis for bud development rate involves the interaction between chilling satisfaction (fulfilling endodormancy requirements) and subsequent heat accumulation (driving ecodormancy release and bud growth), with the degree of chilling affecting both the heat requirement for budbreak and the synchrony of bud development across the vine, such that well-chilled vines show uniform, predictable bud development while inadequately chilled vines show delayed, erratic budbreak. The cultivar variation in bud development rate distinguishes early-budding cultivars (Chardonnay, Gewürztraminer with rapid bud development requiring less heat accumulation) from late-budding cultivars (Mourvèdre, Cabernet Sauvignon with slower bud development requiring more heat), with this variation providing options for frost avoidance through cultivar selection in frost-prone sites. The management factors affecting bud development rate include pruning timing (later pruning delaying budbreak through wound response effects), cane position (horizontal positioning often accelerating bud development versus vertical), and growth regulator applications (hydrogen cyanamide accelerating dormancy release in warm climates with insufficient chilling). Understanding bud development rate quality enables prediction of budbreak timing, assessment of frost risk based on cultivar-specific development rates, and management strategies for manipulating bud development to optimize the frost-growth tradeoff.
Bud_Development_Rate_Quality addresses the speed of bud progression from dormancy through budbreak determining spring frost vulnerability. This quality is regulated by chilling satisfaction and subsequent heat accumulation. The class supports budbreak prediction and frost risk management through bud development rate characterization.
Austriebsrate
Dormanzfreisetzungsrate
Frühjahresentwicklungsrate
Knospenentwicklungsrate
bud growth rate
budbreak rate
dormancy release rate
spring development rate
Duchêne E et al. (2010) A grapevine (Vitis vinifera L.) deoxy-d-xylulose synthase gene colocates with a major quantitative trait loci for terpenol content. Theoretical and Applied Genetics 120(7): 1451-1461 Ferguson JC et al. (2014) Dynamic thermal time model of cold hardiness for dormant grapevine buds. Annals of Botany 113(5): 801-811
Bud Development Rate Quality
Bud freezing tolerance quality comparison across commercial Vitis vinifera cultivars established cultivar-specific hardiness rankings for cold-climate site selection guidance. Differential thermal analysis (DTA) measuring the low-temperature exotherm (LTE) indicating bud freezing provided precise supercooling point determination for buds collected at maximum midwinter hardiness. Riesling demonstrated superior bud freezing tolerance quality with mean LTE at -26.2°C and LT50 (50% bud mortality) at -25.5°C, establishing it as one of the most cold-hardy V. vinifera cultivars. Chardonnay showed intermediate bud freezing tolerance quality with LTE at -23.8°C and LT50 at -23.0°C. Merlot exhibited poor bud freezing tolerance quality with LTE at -20.5°C and LT50 at -19.8°C, restricting its successful cultivation to regions where winter minima rarely exceed -20°C. Within-cultivar variation showed that primary bud freezing tolerance was consistently 2.5-3.5°C lower than secondary bud tolerance, with tertiary buds showing an additional 1-2°C advantage, establishing the backup bud survival margins available for recovery following marginal freeze events. The bud freezing tolerance quality characterization enabled cultivar recommendations matched to site-specific winter minimum temperature probabilities, with Riesling recommended for sites with regular -25°C events while Merlot was restricted to sites with rare occurrence of temperatures below -18°C.
Bud freezing tolerance quality is a tissue freezing tolerance quality that inheres in grapevine dormant buds and represents the capacity to survive freezing temperatures through deep supercooling mechanisms that maintain bud primordia in an unfrozen state down to temperatures at which homogeneous ice nucleation occurs, with the supercooling point determining the survival threshold and varying among cultivars, bud positions, and acclimation states in ways that establish bud survival probability under specific winter minimum temperature exposures. This quality addresses the most economically critical tissue for cold hardiness assessment, recognizing that dormant compound buds contain the primordia for the following season's shoots, leaves, and inflorescences such that bud survival directly determines spring growth capacity and potential crop load, making bud freezing tolerance the primary determinant of cultivar suitability for cold-climate viticulture. The ontological significance of bud freezing tolerance quality lies in its direct relationship to crop security, with inadequate bud hardiness resulting in spring shoot deficits that reduce yield proportionally to bud mortality, while the differential hardiness of primary, secondary, and tertiary buds within compound buds provides a hierarchical recovery system where progressively hardier backup buds can partially compensate for primary bud loss. The biological basis of bud freezing tolerance encompasses the deep supercooling mechanism in which the meristematic tissues of bud primordia avoid ice formation through the combination of structural barriers limiting ice propagation from surrounding tissues, low water content reducing ice nucleation probability, and the accumulation of sugars and other compounds further depressing the freezing point of cellular water. The supercooling capacity of buds increases during autumn acclimation as dehydration reduces water content and cryoprotectant accumulation depresses the temperature at which unavoidable ice nucleation occurs, with maximum hardiness typically achieved in midwinter after sustained cold exposure has completed the acclimation process. The position of buds on the cane affects hardiness, with basal buds typically showing greater tolerance than distal buds reflecting both earlier acclimation and lower metabolic activity, creating the common pattern of basal bud survival accompanying distal bud loss following marginal freeze events.
Bud_Freezing_Tolerance_Quality is_a Tissue_Freezing_Tolerance_Quality, representing the cold hardiness specifically of dormant compound buds critical for spring growth capacity. This class addresses the economically most important tissue for cold hardiness assessment. Deep supercooling mechanism distinguishes bud hardiness from freezing tolerance of woody tissues. LTE (low-temperature exotherm) from DTA provides precise supercooling point measurement. Primary-secondary-tertiary bud hardiness differential provides hierarchical backup system. The -20 to -26°C range among V. vinifera cultivars enables climate-matched selection. Basal > distal bud hardiness affects pruning strategies for cold-damaged vines.
Knospen-Gefrier-Toleranz
Knospen-Kältehärte
Knospen-Unterkühlungskapazität
Winterknospen-Überlebensfähigkeit
bud cold hardiness
bud supercooling capacity
bud winter survival
dormant bud freezing resistance
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Mills LJ et al. (2006) Timing of low temperature exposure affects cold hardiness and cold deacclimation kinetics in grape. American Journal of Enology and Viticulture 57:20-26 Londo JP, Kovaleski AP (2019) Deconstructing cold hardiness: variation in supercooling ability and chilling requirements in the wild grapevine Vitis riparia. Australian Journal of Grape and Wine Research 25:276-285
Bud Freezing Tolerance Quality
Bud fruitfulness in Chardonnay increases from 0.8 clusters per bud at node 1-2 to 2.1 clusters at nodes 5-8, then decreases slightly to 1.8 at nodes 12-15. Riesling shows relatively uniform fruitfulness (1.4-1.8 clusters) across nodes 3-12, enabling flexible pruning strategies. Primary buds contain 1-3 inflorescence primordia in most cultivars, while secondary buds average 0.5-1.0 and tertiary buds rarely exceed 0.3 primordia. Microscopic dissection of dormant buds reveals primordium development stage, with fully differentiated primordia (stage 5-6) indicating high fruitfulness potential.
A fruitfulness disposition that is the inherited capacity of a Vitis vinifera plant to form inflorescence primordia within compound buds at specific positions along the cane, with characteristic patterns of primordium number and development across node positions from base to apex.
Bud_Fruitfulness_Disposition serves as the parent class for position-specific fruitfulness patterns. The compound bud structure in grapevine contains primary, secondary, and tertiary buds, each with different fruitfulness potential. Environmental conditions during primordium initiation (temperature, light, carbohydrate availability) in year N-1 strongly influence bud fruitfulness realized in year N. Pruning strategy must consider cultivar-specific fruitfulness patterns to optimize yield and quality.
Augenfertilität
Knospenfruchtbarkeit
bud fertility
node fruitfulness
positional fertility trait
positionelle Fruchtbarkeit
Buttrose MS (1974) Climatic factors and fruitfulness in grapevines. Horticultural Abstracts 44:319-326 Dunn GM, Martin SR (2000) Do temperature conditions at budburst affect flower number in Vitis vinifera L. cv. Cabernet Sauvignon? Australian Journal of Grape and Wine Research 6:116-124
Bud Fruitfulness Disposition
Bud fruitfulness assessment comparing canopy positions in Cabernet Sauvignon reveals the light exposure effect on cluster initiation, with well-exposed exterior buds showing high fruitfulness (2.2 clusters/bud, two inflorescence primordia visible in dissected buds), partially shaded interior buds showing moderate fruitfulness (1.4 clusters/bud), and heavily shaded basal interior buds showing low fruitfulness (0.6 clusters/bud, many buds with only leaf primordia), with the fruitfulness gradient explaining the 60% yield reduction when dense canopies shade developing buds, demonstrating how previous-season canopy management affects current-season yield potential through bud fruitfulness determination.
Bud fruitfulness quality is a yield potential quality that represents the number of inflorescence primordia (cluster initials) present within dormant buds, characterizing the reproductive capacity established during bud differentiation the previous season that determines the maximum cluster number achievable from each bud when it grows the following spring. Within the BFO framework, this quality specifically addresses the primordial cluster dimension of yield potential, distinguishing the reproductive structures formed within buds from subsequent flower development or fruit set while recognizing that bud fruitfulness sets the first and most fundamental limit on yield—a bud containing no inflorescence primordia cannot produce clusters regardless of how favorable subsequent conditions may be. The ontological significance lies in bud fruitfulness being determined a full season before the clusters will be harvested, with conditions during the critical differentiation period (typically 4-8 weeks post-flowering of the current season) establishing the reproductive potential that will only be expressed 12-14 months later, creating the temporal separation between determination and expression that makes fruitfulness both difficult to manage in the current season and potentially predictable from the previous season's conditions. The factors affecting bud fruitfulness during differentiation include light exposure (well-exposed buds differentiating more cluster primordia than shaded buds), temperature (moderate warmth favoring differentiation, extremes potentially limiting it), carbohydrate availability (adequate reserves supporting primordial development), and node position (basal buds often less fruitful than mid-cane positions), with these factors explaining the management connections between canopy architecture, vine balance, and subsequent yield potential. The measurement of bud fruitfulness involves dissection of dormant buds under magnification to count visible inflorescence primordia, enabling pre-season yield forecasting and assessment of the previous season's conditions for cluster initiation. Understanding bud fruitfulness quality enables yield prediction from bud dissection data, diagnosis of low fruitfulness causes, and management targeting improved light exposure and vine balance to enhance fruitfulness for future seasons.
Bud_Fruitfulness_Quality addresses the inflorescence primordia number within buds determining cluster potential. This quality is established during bud differentiation the previous season. The class supports yield forecasting and canopy management through fruitfulness characterization linking previous conditions to current potential.
Knopensfertilität
Knospenfruchbarkeit
Traubenanlage
cluster initiation
inflorescence primordia number
reproductive bud capacity
reproduktive Knospenkapazität
Srinivasan C, Mullins MG (1981) Physiology of flowering in the grapevine—a review. American Journal of Enology and Viticulture 32(1): 47-63 May P (2004) Flowering and Fruitset in Grapevines. Lythrum Press, Adelaide, pp. 1-119
Bud Fruitfulness Quality
The bud load specification for Pinot Noir in the Côte de Nuits prescribes 8 buds on the fruiting cane plus 2 buds on the renewal spur for a total of 10 count buds per vine at Guyot simple — at a planting density of 10,000 vines/ha this yields 100,000 buds/ha, of which approximately 85-90% will produce shoots (accounting for blind buds and winter mortality), generating 85,000-90,000 shoots/ha each bearing 1-2 clusters for a potential yield of 50-60 hl/ha before any green harvest reduction. In contrast, the bud load specification for Cabernet Sauvignon on bilateral cordon in Coonawarra prescribes 14-16 two-bud spurs (28-32 buds per vine) at 3,700 vines/ha yielding approximately 100,000-120,000 buds/ha for a target yield of 6-8 t/ha.
A pruning method description that documents the total number of retained buds per vine after pruning, specifying the primary determinant of potential shoot number, cluster number, and crop load that establishes the vine's yield ceiling for the upcoming growing season and constitutes the most direct mechanism for regulating the balance between vegetative growth and reproductive output.
Bud_Load_Specification is the single most important quantitative parameter in pruning because it establishes the potential shoot population from which all subsequent yield and canopy density outcomes derive. The optimal bud load for a given vine is determined by the interaction of vine capacity (the total amount of vegetative and reproductive growth the vine can ripen in the available season), target yield (set by appellation regulations, quality objectives, and economic requirements), and cultivar-specific fruitfulness patterns: excessive bud load relative to vine capacity produces overcropping symptoms (delayed ripening, poor fruit quality, reduced carbohydrate reserves), while insufficient bud load wastes vine capacity and reduces economic returns. The Ravaz index (yield/pruning weight ratio, optimal range 5-10 for most wine grapes) provides the retrospective assessment of whether bud load was appropriate, with values above 10 indicating overcropping and values below 3 indicating under-utilization of vine capacity. Appellation specifications frequently prescribe maximum bud loads (e.g., Burgundy Grand Cru limits typically 8-10 buds per vine for Guyot).
Anschnittmenge-Spezifikation
Augenzahl-Vorgabe
Knospenbelastungs-Spezifikation
bud number specification
retained bud count
vine bud load target
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 335-365 Ravaz L (1911) L'effeuillage de la vigne, Annales de l'École Nationale d'Agriculture de Montpellier 11: 216-244 Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles pp: 91-120
Bud Load Specification
Long-term evaluation of Bud_Mite_Resistance_Disposition at the Viticulture Research Center Haro, Spain, monitored bud mite damage across 25 cultivars over 8 seasons in replicated vineyard blocks without acaricide application. Spring shoot damage was scored on a 0-5 scale assessing internode shortening, leaf distortion, and scar tissue presence, while fruitfulness was measured as clusters per shoot. The susceptible cultivar 'Garnacha Tinta' exhibited chronic bud mite damage with mean spring scores of 3.2, shortened first internodes (4.2 cm vs 8.5 cm in healthy vines), and 25% fruitfulness reduction (0.9 vs 1.2 clusters/shoot in protected controls). Damage severity varied substantially among seasons (scores 1.8-4.5) correlating with winter minimum temperatures affecting mite survival. 'Tempranillo' demonstrated strong Bud_Mite_Resistance_Disposition with consistently low damage scores (mean 0.6, range 0.2-1.1 across seasons), normal internode development, and no detectable fruitfulness impact despite equivalent mite populations detected in dormant bud dissections. Anatomical examination revealed that 'Tempranillo' buds possessed tighter scale overlap and denser scale trichomes than 'Garnacha', potentially limiting mite access to meristematic tissues. Additionally, 'Tempranillo' showed more rapid spring bud development, reducing the duration of susceptibility to mite feeding on undifferentiated tissues. These combined structural and phenological factors likely contribute to its Bud_Mite_Resistance_Disposition.
A Bud_Mite_Resistance_Disposition is an Eriophyid_Mite_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that prevent or limit colonization, feeding, and damage within dormant and developing buds by bud-infesting eriophyid mites, particularly the bud mite strain of Colomerus vitis and the grape rust mite Calepitrimerus vitis. This disposition represents resistance to mites that overwinter within bud scales and feed on meristematic tissues during bud development, causing a spectrum of damage ranging from mild scarring and reduced leaf size to severe shoot stunting, shortened internodes, distorted leaves, and drastically reduced fruitfulness. Bud mite damage is often more economically significant than erineum damage, as severe infestations can reduce crop yields by 30-50% through destruction of developing inflorescences within buds. Resistance mechanisms potentially include bud architectural features limiting mite access to feeding sites, chemical factors within bud tissues toxic or deterrent to mites, rapid tissue differentiation reducing the window of susceptibility, and overall bud vigor enabling compensation for moderate mite damage. Bud mite damage is notoriously variable among vines and seasons, complicating resistance evaluation, as factors including vine stress, winter temperatures affecting mite survival, and spring weather affecting bud development interact with host genotype to determine damage severity. Nevertheless, consistent cultivar differences have been documented, with some cultivars showing chronic susceptibility while others rarely exhibit damage despite equivalent mite pressure, providing evidence for genetic variation in Bud_Mite_Resistance_Disposition amenable to selection.
Bud_Mite_Resistance_Disposition is_a Eriophyid_Mite_Resistance_Disposition. Bud mites include the bud strain of Colomerus vitis and Calepitrimerus vitis (grape rust mite), both of which overwinter in buds and damage developing shoots. Bud mite damage is more economically significant than erineum but more difficult to evaluate due to high environmental variability. Resistance appears to involve both structural (bud architecture) and phenological (rapid development) components. Bud mite biology differs sufficiently from the erineum strain that resistance may be independently controlled.
Colomerus bud strain resistance
Colomerus-Knospen-Resistenz
Knospengallmilben-Abwehr
Knospenmilben-Resistenz-Disposition
bud eriophyid resistance
grape bud mite resistance
Walton VM et al. (2007) Relationship between rust mites Calepitrimerus vitis (Nalepa) and bud damage on selected grape cultivars, Vitis 46:66-70 Bernard MB et al. (2005) Eriophyoid mite damage in Vitis vinifera (grapevine) in Australia: Calepitrimerus vitis and Colomerus vitis (Acari: Eriophyidae) as the causes of the damage, Experimental and Applied Acarology 35:207-223 Smith LM, Stafford EM (1948) The bud mite and the erineum mite of grapes, Hilgardia 18:317-334
Bud Mite Resistance Disposition
In Merlot vines in Saint-Émilion, Bordeaux, bud scale formation progresses during October and early November with outer scale primordia ceasing expansion and differentiating into protective scales by mid-October, dense trichome development on outer scale surfaces completing by late October, and full bud scale enclosure of dormant meristems achieved by early November when scales show characteristic brown coloration and tight overlapping arrangement typical of BBCH stage 93.
Bud scale formation in Vitis vinifera represents the developmental process during which the outermost leaf primordia of dormant buds differentiate into specialized protective structures that provide physical and physiological protection for enclosed meristematic tissues during winter dormancy. This process occurs during autumn as part of dormancy establishment, with bud scales developing from leaf primordia that would have formed foliage leaves under continued growth conditions but instead differentiate into the tough, often pubescent protective structures characteristic of dormant grapevine buds. The bud scales of Vitis vinifera are morphologically modified leaves that retain recognizable leaf-like features including a flattened blade-like structure and association with the node position, but exhibit distinctive specializations including thick cuticle development, dense trichome coverage particularly on outer scale surfaces, accumulation of phenolic compounds that provide antimicrobial protection, and the development of tight overlapping arrangement that seals the bud interior from environmental exposure. The developmental transition from leaf primordium to bud scale involves altered patterns of gene expression that suppress normal leaf blade expansion and promote the alternative developmental program leading to scale morphology, with this transition triggered by the same dormancy-inducing signals of short photoperiod and declining temperatures that drive overall dormancy establishment. The physical protection provided by bud scales includes reduction of water loss from enclosed tissues that would otherwise desiccate during winter, limitation of ice propagation into bud interior during freezing events, and exclusion of fungal pathogens and small invertebrates that might otherwise damage dormant meristems. The number of bud scales per dormant bud varies somewhat among cultivars and bud positions, typically ranging from 2-4 outer scales that provide the primary protective function, with the innermost protective structures transitioning toward leaf-like morphology representing an intermediate developmental state. Bud scale formation must be complete before severe winter conditions occur for adequate bud protection, with incomplete scale development leaving buds vulnerable to winter injury from desiccation, freezing, or pathogen invasion. The timing of bud scale formation correlates with other dormancy establishment processes, progressing in parallel with growth cessation and ABA accumulation during the final phase of dormancy induction.
Bud_Scale_Formation represents a morphological component of Dormancy_Establishment, specifically addressing the development of protective bud structures. Distinguished from normal leaf development by the alternative differentiation pathway producing protective rather than photosynthetic structures. Occurs in parallel with ABA_Accumulation and Deep_Dormancy_Entry during dormancy establishment. Typically involves 2-4 outer scales per bud. Provides physical protection against desiccation, freezing, and pathogens. Duration of active scale differentiation spans 3-4 weeks during autumn.
Knospenschuppen-Bildung
Knospenschutz-Entwicklung
Schuppenblatt-Entwicklung
Winterknospen-Beschuppung
bud scale development
dormant bud scale formation
protective scale development
scale leaf formation
Pratt C (1974) Vegetative anatomy of cultivated grapes - a review. American Journal of Enology and Viticulture 25: 131-150 Morrison JC (1991) Bud development in Vitis vinifera L. Botanical Gazette 152: 304-315 Mullins MG, Bouquet A, Williams LE (1992) Biology of the Grapevine Cambridge University Press
Bud Scale Formation
Riesling in Mosel initiates bud swelling at BBCH 01-03 in mid-April after 1100-1300 chilling hours and 180-220 growing degree days base 10°C, with swelling phase requiring 12-15 days under spring conditions averaging 11-14°C mean temperature and 60-75% relative humidity, representing later burst timing characteristic of this cool-climate cultivar
Bud swelling constitutes the initial visible phase of bud burst, characterized by volumetric expansion of dormant buds through water uptake and resumption of metabolic activity in meristematic tissues. This process corresponds to BBCH stages 01-03, marking the transition from endodormancy to active growth as winter buds increase 15-30% in volume over 7-14 days. The process initiates when sustained temperatures above 10°C trigger hormonal signals releasing growth inhibition, with abscisic acid concentrations declining in bud tissues while cytokinin levels increase through root-to-shoot transport. Water uptake drives initial expansion as osmotic potential increases in bud cells through starch hydrolysis to soluble sugars and protein synthesis from stored nitrogen compounds. Cell walls soften through expansin activity and loosening of cellulose-hemicellulose networks, allowing turgor-driven expansion without cell division at this stage. Vascular reconnection between bud and supporting cane tissue establishes functional xylem for water transport and phloem for nutrient delivery from storage reserves. The outer bud scales begin separating at margins as internal tissues expand, creating visible gaps between protective scales while green tissue remains concealed. Metabolic reactivation includes elevated respiration rates consuming stored carbohydrates, protein synthesis supporting cell structure and enzyme production, and lipid mobilization from storage membranes. Temperature sensitivity at this stage is high, with development rate approximately doubling for each 10°C increase between 10-25°C base temperatures. The process exhibits three distinguishable phases based on swelling progression and bud scale separation, progressing from initial barely-visible expansion through clearly enlarged buds to late swelling where scale tips separate revealing first hints of internal green tissue. Cultivar variation in swelling rate and duration reflects genetic differences in dormancy depth and temperature response thresholds. Early cultivars like Chardonnay may complete swelling in 7-10 days while later varieties like Cabernet Sauvignon require 12-16 days under identical temperature regimes. Regional timing varies from late March in Mediterranean climates to late April or early May in continental regions, with accumulated growing degree days providing more reliable prediction than calendar dates. Viticultural significance includes serving as first visible indicator for timing protective frost measures, determining optimal pruning completion deadlines, and providing developmental reference for early-season pest monitoring. Frost damage during swelling causes bud mortality through ice crystal formation in expanding cells with high water content, requiring replacement growth from secondary or basal buds with reduced crop potential. Research applications focus on modeling temperature responses for phenology prediction, analyzing molecular mechanisms of dormancy release and growth resumption, and evaluating climate change impacts on growing season initiation timing across viticultural regions.
Bud_Swelling precedes Bud_Breaking and represents expansion phase before protective scales actually separate fully. Distinguished by bud remaining closed with scales touching while overall bud volume increases measurably. Further subdivided into Early_Bud_Swelling, Advanced_Bud_Swelling, and Late_Bud_Swelling based on progression. Temperature and water availability are primary rate-limiting factors.
Knospenquellung
Knospenschwellung
Schwellphase
bud swell
swollen bud stage
winter bud swelling
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 157-172 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 63-78 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Bud Swelling
In Grüner Veltliner vines in the Wachau, Austria, bud swelling initiation is observed in late March when accumulated GDD reach approximately 120 units, with buds showing visible enlargement and scale separation at BBCH stage 05 within 3-4 days of threshold achievement, woolly trichomes becoming visible at scale margins within 5-6 days, and progression to green tip emergence at BBCH stage 07 occurring within 8-12 days depending on subsequent temperature conditions.
Bud swelling initiation in Vitis vinifera represents the first externally visible sign of dormancy break, characterized by the enlargement of dormant buds as the enclosed primordial shoot begins expansion that increases bud volume and produces visible separation of bud scales. This morphological change marks the transition from the externally static appearance of dormant buds to the dynamic developmental progression of budbreak, serving as the practical field indicator that dormancy has been broken and active growth has commenced. The cellular basis of bud swelling involves the initiation of cell expansion in the primordial shoot enclosed within bud scales, driven by turgor pressure generated through osmotic water uptake into enlarging cells, with cell wall loosening enabling the increase in cell volume that produces macroscopic bud enlargement. The anatomical changes during bud swelling initiation include the expansion of primordial leaves and internodes that were formed during the previous growing season and remained quiescent through dormancy, separation of bud scales as the expanding shoot pushes against them, and the appearance of woolly trichomes at scale margins as protective structures become visible through widening scale gaps. The temporal position of bud swelling initiation corresponds to BBCH stage 05 in the standardized grapevine phenological scale, representing the first stage of visible development following the dormant winter period at BBCH 00 and preceding the green tip emergence at BBCH 07 that follows as development progresses. The rate of bud swelling following initiation varies with temperature, with warm conditions accelerating expansion and producing rapid progression through subsequent developmental stages, while cool conditions slow expansion and may extend the bud swell phase over days to weeks. The synchrony of bud swelling initiation across a vine or vineyard provides important information about dormancy status and subsequent growth uniformity, with simultaneous initiation indicating uniform dormancy release while extended initiation periods suggest variable dormancy satisfaction or response heterogeneity. The vulnerability during bud swelling initiation relates to the increasing frost sensitivity as buds transition from the relatively hardy dormant state to the frost-sensitive swollen state, with lethal temperatures rising from approximately -15°C for dormant buds to -4 to -6°C for swelling buds.
Bud_Swelling_Initiation represents the first visible developmental event within Dormancy_Break. Distinguished from internal growth resumption by representing the externally observable morphological change. Corresponds to BBCH stage 05. Driven by cell expansion and turgor development in primordial shoot. Rate varies with temperature following initiation. Synchrony indicates uniformity of dormancy release. Marks transition to increased frost sensitivity requiring protection attention.
Knospenschwellungs-Initiierung
Knospenschwellungsbeginn
bud enlargement initiation
bud swell onset
early bud swelling
frühe Knospenvergrößerung
sichtbarer Austriebsbeginn
visible budbreak beginning
Lorenz DH et al (1995) Growth stages of the grapevine: Phenological growth stages of the grapevine - BBCH-scale. Australian Journal of Grape and Wine Research 1: 100-103 Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Pratt C (1974) Vegetative anatomy of cultivated grapes - a review. American Journal of Enology and Viticulture 25: 131-150
Bud Swelling Initiation
Bud viability assessment following severe winter in Cabernet Franc reveals the survival pattern guiding spring management, with dissection of 200 buds across the vineyard showing 62% primary bud viability (38% killed by mid-winter -24°C event), 85% secondary bud viability (more hardy secondary buds surviving better), and positional pattern of 45% primary viability at basal nodes (1-3) versus 72% at mid-cane positions (4-8), with this viability information enabling adjusted pruning (leaving additional buds to compensate for mortality) and realistic yield expectations (reduced crop from secondary bud contribution to surviving shoot population).
Bud viability quality is a phenological quality that represents the living status and growth potential of grapevine buds, characterizing whether buds contain living meristematic tissue capable of producing shoots when conditions become favorable, with viability being the fundamental prerequisite for spring growth that determines the productive potential available after winter and the recovery capacity following any damaging events. Within the BFO framework, this quality addresses the living status dimension of bud condition, distinguishing whether buds are alive from their dormancy status or hardiness level while recognizing that viability is binary at the individual bud level (alive or dead) but continuous at the population level (percentage of buds viable), with viability assessment providing the direct measure of bud survival that hardiness measurements only indirectly predict. The ontological significance lies in bud viability determining the productive capacity emerging from winter, where the percentage of viable buds directly establishes the maximum number of shoots possible (setting the ceiling for yield potential), the distribution of viability among bud types (primary versus secondary) affects the quality potential (primary buds being more fruitful), and viability patterns across the vine (basal versus distal positions) affect training and pruning decisions for the coming season. The compound bud structure of grapevine creates the hierarchy of primary, secondary, and tertiary buds within each compound bud, with differential viability among these components providing the backup system that enables some recovery even when primary buds are killed, though at the cost of reduced fruitfulness and altered shoot characteristics from secondary or tertiary bud origin. The assessment of bud viability involves either dissection methods (cutting buds to observe tissue color, with green indicating viable and brown indicating dead tissue) or forcing methods (placing bud samples under growth conditions and observing which buds produce shoots), with these methods providing direct viability data that can be compared to hardiness predictions and used for management decisions. Understanding bud viability quality enables assessment of winter survival outcomes, prediction of spring productive potential, and informed pruning decisions that account for the actual bud survival pattern revealed by viability testing.
Bud_Viability_Quality serves as the parent class for bud type-specific viability (Primary, Secondary, Basal). This quality addresses the living status determining productive potential after winter. The class supports post-winter assessment and management planning through bud viability characterization.
Knospenvitalität
Knospenüberlebensrate
Meristemlebensfähigkeit
bud liveness
bud survival
bud vitality
meristem viability
Wolpert JA, Howell GS (1986) Cold acclimation of Concord grapevines III. Relationship between cold hardiness, tissue water content, and shoot maturation. Vitis 25(3): 151-159 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 267-330
Bud Viability Quality
In Grüner Veltliner vines in the Wachau, Austria, budbreak readiness is achieved by mid-February following satisfaction of the approximately 1100 chilling hour requirement, with ready buds demonstrating forcing response within 9 days at 22°C, ultrastructural examination showing meristem cells with prominent nucleoli and dispersed chromatin characteristic of division readiness, and subsequent field budbreak occurring predictably within 5 days of accumulated growing degree days reaching 130 units above the 10°C base temperature in late March.
Budbreak readiness in Vitis vinifera represents the fully prepared state achieved following completion of dormancy release processes, describing buds that have restored all competencies required for growth and await only adequate heat accumulation to initiate the visible developmental changes of budbreak. This state of readiness is achieved when chilling requirements have been satisfied, growth capacity has been restored, and metabolic systems have been prepared for reactivation, establishing the complete preparation for growth that characterizes the ecodormant bud following endodormancy release. The cellular dimension of readiness involves meristematic cells poised at the G1/S boundary of the cell cycle prepared to initiate DNA replication and subsequent division upon appropriate signaling, with all cell cycle components present and functional awaiting activation. The molecular dimension encompasses the full complement of growth-related gene transcripts and proteins accumulated in preparation for growth, with transcription factors poised to activate growth programs and biosynthetic enzymes ready for rapid upregulation. The hormonal dimension involves the growth-permissive hormone balance established through dormancy release, with low ABA removing inhibition, sensitized cytokinin and gibberellin signaling prepared to transduce growth signals, and auxin transport systems ready to establish the polar gradients organizing growth. The structural dimension addresses the ultrastructural preparation of meristematic cells for division, including prominent nucleoli indicating ribosome biogenesis capacity, dispersed chromatin indicating transcriptional readiness, and organized cytoskeleton prepared for division and expansion. The practical significance of budbreak readiness relates to the predictability of subsequent development, with ready buds responding uniformly and predictably to heat accumulation according to growing degree day models, enabling accurate prediction of budbreak timing from temperature forecasts. The transition from readiness to active budbreak occurs when accumulated heat units satisfy the cultivar-specific requirement, typically 100-300 growing degree days above the base temperature, at which point visible bud swelling commences marking the initiation of the new growing season.
Budbreak_Readiness represents the complete preparedness state within Bud_Competence_Restoration describing fully prepared buds awaiting heat accumulation. Distinguished from earlier competence states by representing complete preparation for imminent growth. Encompasses cellular, molecular, hormonal, and structural readiness dimensions. Enables predictable budbreak response to heat accumulation. Transition to active growth when GDD requirement satisfied, typically 100-300 units. Corresponds to late BBCH 00 immediately preceding visible bud swelling.
Austriebsbereitschaft
Knospenaktivierungs-Bereitschaft
Knospenaustritts-Vorbereitung
Wachstumsinitiierungs-Bereitschaft
bud activation readiness
budbreak preparedness
growth initiation readiness
pre-budbreak competence
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Lorenz DH et al (1995) Growth stages of the grapevine: Phenological growth stages of the grapevine - BBCH-scale. Australian Journal of Grape and Wine Research 1: 100-103
Budbreak Readiness
The budbreak timing description of Chardonnay documents early budbreak (OIV 301:3) — typically reaching 50% green tip 3-5 days before the Chasselas reference, placing Chardonnay among the early-budding cultivars in European phenological rankings. At Burgundy latitudes (47°N), Chardonnay budbreak occurs in late March to early April (March 25 - April 10, depending on winter and spring temperatures), approximately 7-10 days before Pinot Noir and 12-18 days before the late-budding Cabernet Sauvignon. This early budbreak exposes Chardonnay to significant spring frost risk — the tender green shoots emerging at budbreak are killed at temperatures of -2 to -3°C, and late frost events in early April have caused catastrophic crop losses in Chablis (notably 2016, 2017, and 2021), motivating the widespread use of frost protection measures (bougies, aspersion, wind machines, and increasingly electrical resistance heating) in frost-prone Chardonnay vineyards.
A phenological timing description component that records the date at which 50% of buds on the vine have reached the green tip stage (E-L stage 4, Baggiolini stage C, BBCH 07) — the phenological event marking the end of winter dormancy and the beginning of the active growing season — scored according to OIV descriptor 301 on a scale from very early (state 1) to very late (state 9) relative to the Chasselas reference, with budbreak timing determining the cultivar's exposure risk to late spring frost and the length of the available growing season from first growth to harvest.
Budbreak_Timing_Description documents the first visible phenological event of the growing season — the moment when the protective bud scales separate and the green shoot tip emerges, signaling that the accumulated chilling requirement (typically 400-1,200 chilling hours below 7°C for V. vinifera) and the subsequent heat requirement for bud growth resumption have been satisfied. The timing of budbreak is controlled by the interaction of two sequential physiological processes — endodormancy release (requiring adequate chilling during winter to satisfy the cultivar-specific chilling requirement) and ecodormancy break (requiring adequate spring warmth to trigger growth resumption after endodormancy is satisfied): cultivars differ in both their chilling requirement (low-chill cultivars like Chasselas and Chardonnay satisfy dormancy earlier, enabling earlier growth resumption) and their base temperature for spring growth (some cultivars initiating growth at lower temperatures than others), with the combined effect determining the relative budbreak sequence that is consistent across sites. The practical significance of budbreak timing centers on frost risk management — each day of earlier budbreak increases the probability of frost damage because late frost events become less frequent as spring progresses, and the accumulated research from Burgundy, Champagne, Chablis, and other frost-prone regions has quantified the economic value of later budbreak at approximately 2-5% reduced frost damage probability per day of delay, making late budbreak a desirable trait for frost-prone sites even when the reduced growing season length constrains cultivar choice.
Austriebs-Zeitpunkt
Austriebszeitpunkt-Dokumentation
Knospendurchbruch-Beschreibung
budburst timing record
dormancy break date
spring growth initiation
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Duchêne E and Schneider C (2005) Grapevine and climatic changes, Agronomy for Sustainable Development 25: 93-99 Lavee S and May P (1997) Dormancy of grapevine buds, Australian Journal of Grape and Wine Research 3: 31-46 Jones GV and Davis RE (2000) Climate influences on grapevine phenology, American Journal of Enology and Viticulture 51: 249-261
Budbreak Timing Description
Budbreak timing spans a 4-6 week range across V. vinifera cultivars in any given location, with Chardonnay and Pinot Noir among the earliest (base 10°C GDD requirement of 50-80°C) and Mourvèdre and Carignan among the latest (GDD requirement of 120-160°C). In Bordeaux, Merlot reaches BBCH 09 (green tip visible) approximately 7-10 days before Cabernet Sauvignon in the same vineyard. Budbreak date shows year-to-year variation of ±10-18 days within cultivar depending on spring temperature patterns. The base temperature for bud development ranges from 7-12°C across cultivars, with lower base temperatures associated with earlier budbreak.
A phenological timing disposition that is the inherited capacity of a Vitis vinifera plant to initiate visible bud swelling and green tissue emergence (BBCH 07-09) at a characteristic time in spring, determined by the interaction of chilling fulfillment, heat accumulation requirements, and genotype-specific thermal sensitivity.
Budbreak_Timing_Disposition serves as the parent class for early, mid-season, and late budbreak categories. The trait represents the first visible phenological event of the growing season and determines the starting point for GDD accumulation toward subsequent stages. Budbreak timing interacts critically with spring frost risk, as earlier cultivars face greater frost exposure probability. Thermal time models using cultivar-specific base temperatures predict budbreak with ±3-5 day accuracy when dormancy release timing is correctly estimated.
Austriebs-Disposition
Austriebszeitpunkt
Knospentreiben
bud emergence timing
budburst timing
spring phenology trait
Duchêne E, Schneider C (2005) Grapevine and climatic changes: a glance at the situation in Alsace. Agronomy for Sustainable Development 25:93-99 Garcia de Cortázar-Atauri I et al. (2009) Performance of several models for predicting budburst date of grapevine (Vitis vinifera L.). International Journal of Biometeorology 53:317-326
Budbreak Timing Disposition
A dormant cane of Merlot clone 181 bears a Budwood_Role, harvested in February from a certified scion mother vine, stored at 2°C wrapped in moist newspaper, then transported to a California vineyard for T-budding onto established 110 Richter rootstocks in late May—each cane yielding 8-12 viable buds for field insertion. Budwood of Grüner Veltliner clone 17 serves in a Budwood_Role for chip budding operations at Austrian nurseries, with individual chip buds excised using specialized knives and immediately inserted into prepared rootstock notches. A budstick of Shiraz clone BVRC30 bears a Budwood_Role in Australian field budding programs, supporting T-bud insertion during the rootstock's active growth period when bark slips readily from cambium.
A scion wood role that inheres in stem material designated as the source of individual buds for budding propagation techniques (T-budding, chip budding), where single buds with associated bark shield or chip tissue are excised and inserted into rootstock tissue to establish the scion variety. Budwood provides the meristematic bud tissue that develops into the scion shoot system, with each bud potentially producing an independent grafted vine—enabling high multiplication rates from limited source material compared to grafting techniques requiring multi-node scion segments. Budwood quality assessment focuses on bud health (visual inspection for damage, disease symptoms, or developmental abnormalities), bud maturity (dormant buds for dormant budding, active buds for June budding), and bark condition (ease of peeling for T-budding, chip uniformity for chip budding). Budwood is typically harvested as whole canes (budsticks) from which individual buds are excised at the time of budding, maintaining bud viability through the brief period between harvest and insertion.
Budwood_Role captures scion material specifically destined for budding rather than grafting, emphasizing individual bud extraction over multi-node segment use. Budding techniques (T-budding, chip budding) differ from grafting in using minimal scion tissue—a single bud with bark shield rather than a multi-node cane segment—enabling maximum multiplication from limited budwood supplies and field propagation without the controlled-environment requirements of benchgrafting. Budwood storage duration is typically shorter than graftwood (days to weeks rather than months), as the small tissue mass is more susceptible to desiccation. The "budstick" designation refers to whole canes maintained as the bud source, with individual buds excised as needed during budding operations—maintaining bud viability better than pre-excision storage of isolated buds.
Augenstecklingsholz
Edelaugenholz
Okulations-Reiser
bud source material
budding wood
budstick
budwood
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 468-480 Whiting JR (2004) Grapevine rootstocks, Winetitles Dry N (2007) Grapevine rootstocks, Lythrum Press Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia
Budwood Role
Bulk density assessment of machine-harvested Chardonnay in transport bins shows values of 0.72±0.05 g/mL compared to theoretical maximum of 0.82 g/mL for perfectly packed berries, with the 12% void space providing essential cushioning that reduces crushing damage during transport, while overfilled bins with bulk density exceeding 0.78 g/mL show 35% higher juice loss indicating excessive compaction and quality compromise.
Bulk density quality is a density quality that inheres in aggregated grape structures at scales above individual berries and represents the mass per unit volume of clustered or canopy-level assemblages where interstitial air spaces between component elements significantly influence the measured value. Within the BFO framework, this quality addresses organizational scales where the spatial arrangement of discrete units creates a composite material whose bulk density differs substantially from the intrinsic density of constituent tissues, recognizing that packing geometry and void volume are determinative factors alongside component material properties. The ontological significance lies in bulk density capturing structural organization information not accessible through measurements of individual components, enabling characterization of cluster compactness, canopy porosity, and harvest bin fill efficiency that affect disease susceptibility, light penetration, spray coverage, and logistics operations. Bulk density measurement requires specification of the boundaries defining the measured volume, whether the convex hull of a cluster, the rectangular prism of a canopy measurement zone, or the container volume for harvested fruit, as these boundary definitions determine void space inclusion. The relationship between bulk density and component density depends on packing fraction, which for grape clusters typically ranges from 0.45-0.75 depending on berry size uniformity and cluster architecture, meaning cluster bulk density of 0.55-0.85 g/mL emerges from berries with intrinsic density of 1.05-1.10 g/mL. Understanding bulk density at cluster and canopy scales supports vineyard management decisions including cluster thinning for disease control, canopy manipulation for light exposure optimization, and harvest logistics planning for transport efficiency and fruit integrity during handling.
Bulk_Density_Quality serves as the parent class for specific aggregation scales including Cluster_Packing_Density_Quality and Canopy_Density_Quality, establishing the principle that interstitial void volume distinguishes bulk from intrinsic material density. This class bridges plant biology with agricultural engineering considerations of harvest logistics and processing efficiency. Measurement protocols must specify boundary definitions and compaction state for reproducibility.
Aggregatdichte
Füllungsdichte
Packungsdichte
Schüttdichte
aggregate density
apparent density of assemblage
packing density
volumetric fill density
Pezzi F, Caprara C (2009) Mechanical grape harvesting: investigation of the transmission of vibrations. Biosystems Engineering 103(3): 281-286 Intrieri C et al. (2008) Yield efficiency of high quality wine grape production in the Po Valley. Acta Horticulturae 785: 309-316
Bulk Density Quality
Bulk elastic modulus determination for Syrah leaves through pressure-volume analysis reveals values of 12.3 MPa under well-watered conditions increasing to 19.8 MPa following three weeks of progressive water deficit, with the 61% increase in modulus indicating cell wall stiffening that maintains leaf turgor at 0.15 MPa higher water content during drought, complementing the 0.35 MPa osmotic adjustment to provide compound improvement in drought tolerance through both osmotic and elastic mechanisms.
Bulk elastic modulus quality is an elastic modulus quality that represents the volumetric stiffness of plant tissue defined as the ratio of turgor pressure change to relative volume change, characterizing the tissue-level resistance to compression under hydrostatic pressure generated by osmotic water uptake into cells. Within the BFO framework, this quality specifically characterizes the mechanical property governing the relationship between turgor and cell volume at the tissue scale, integrating contributions from individual cell wall elasticity, cell geometry, and tissue architecture into a bulk parameter accessible through pressure-volume analysis. The ontological significance lies in bulk elastic modulus determining the water storage capacity of tissues (with lower modulus tissues storing more water per unit pressure change) and the sensitivity of turgor to water loss (with higher modulus tissues maintaining turgor with less volume change), representing a key parameter distinguishing drought response strategies among species and cultivars. Bulk elastic modulus in grapevine leaves typically ranges from 5-25 MPa depending on developmental stage, water status history, and measurement methodology, with higher values in mature, acclimated leaves compared to young expanding leaves. The distinction between bulk modulus and cell wall elastic modulus reflects the tissue-level measurement that includes cellular architecture effects, where cell geometry, intercellular spaces, and the distribution of cell types all influence the relationship between pressure and volume at tissue scale. Determination of bulk elastic modulus employs pressure-volume curve analysis, where the slope of the linear (turgor-positive) region relates pressure change to volume change, or may be calculated from paired measurements of water potential and relative water content during tissue dehydration. The acclimation plasticity of bulk elastic modulus enables adjustment of tissue mechanical properties in response to environmental conditions, with drought acclimation typically increasing modulus to maintain turgor despite water loss, representing an adaptive response complementing osmotic adjustment. Understanding bulk elastic modulus quality enables quantitative analysis of tissue water relations, prediction of turgor maintenance capacity under water deficit, and characterization of drought adaptation mechanisms.
Bulk_Elastic_Modulus_Quality specifically addresses the tissue-scale volumetric stiffness derived from pressure-volume analysis, distinguishing from cell wall elastic modulus measured at cellular scale and Young's modulus measured in uniaxial testing geometry. This quality appears in the pressure-volume relationship as the parameter governing the turgor-volume relationship, with higher values indicating more rigid tissues. The class supports quantitative water relations analysis and drought tolerance characterization.
Druck-Volumen-Elastizität
Gesamtelastizitätsmodul
Kompressionsmodul
Volumenelastizitätsmodul
pressure-volume elastic modulus
tissue bulk modulus
volumetric elastic modulus
Tyree MT, Jarvis PG (1982) Water in tissues and cells. In: Lange OL, Nobel PS, Osmond CB, Ziegler H (eds) Encyclopedia of Plant Physiology Vol 12B, Springer, Berlin, pp. 35-77 Schultz HR, Matthews MA (1993) Growth, osmotic adjustment, and cell-wall mechanics of expanding grape leaves during water deficits. Crop Science 33(2): 287-294
Bulk Elastic Modulus Quality
Tempranillo bears a Bulk_Red_Wine_Scion_Role in La Mancha, Spain, yielding 15-25 t/ha under irrigation on the high central plateau to produce everyday table wines at €2-5/bottle, contrasting with its Premium_Red_Wine_Scion_Role in Rioja and Ribera del Duero. Carignan (Mazuelo, Carignane) historically served in a Bulk_Red_Wine_Scion_Role across southern France and California, valued for productivity, color, and blending utility though now partially rehabilitated for quality production from old vines. Bonarda (Douce Noir) bears a Bulk_Red_Wine_Scion_Role in Argentina, planted extensively in Mendoza and San Juan for volume wine production at yields exceeding 20 t/ha. Nero d'Avola serves both bulk and premium roles in Sicily depending on yield level and site.
A red wine scion role that inheres in a grafted scion cultivar by virtue of its designation for high-volume, commodity-oriented red wine production, where viticultural and winemaking practices prioritize yield, consistency, and cost-efficiency to produce wines for everyday consumption at accessible price points. Bulk red wine scions are cultivated to achieve high yields (typically 12-25 t/ha, with some irrigated regions exceeding 30 t/ha) while maintaining acceptable color, basic varietal character, and commercial soundness. Bulk production favors vigorous, productive cultivars tolerant of mechanical harvesting, minimal hand labor, and industrial-scale vinification. Wines are typically released within months of harvest without extended aging, blended for consistency across vintages, and priced for volume markets (typically below €5-10/bottle retail). Major bulk red wine production occurs in Spain's La Mancha, southern France (Languedoc-Roussillon), Italy's Puglia and Sicily, California's Central Valley, Argentina's Mendoza, Chile's Central Valley, and Australia's Riverland and Murray-Darling regions.
Bulk_Red_Wine_Scion_Role represents the volume end of red wine production essential for supplying everyday wine consumption at affordable prices. The same cultivar frequently serves in both bulk and premium roles depending on viticulture practices—Tempranillo, Sangiovese, Grenache, and Shiraz span the quality spectrum from €2 supermarket wines to €200 classified growths. Bulk production increasingly involves internationally recognized varieties (Cabernet Sauvignon, Merlot, Syrah) grown at high yields for varietal-labeled wines, in addition to traditional volume varieties (Carignan, Monastrell, Bobal). Bulk wine often enters international trade as blending wine, adding color, body, or varietal character to wines finished and bottled in importing countries.
Großmengenrotweinsorte
Keltertraube für Massenwein
Massenrotweinsorte
bulk red wine variety
commodity red wine variety
high-yield red wine cultivar
volume red wine variety
Anderson K, Aryal NR (2020) Database of Regional, National and Global Winegrape Bearing Areas by Variety, 2000 and 2010, Wine Economics Research Centre, University of Adelaide Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane OIV (2019) Statistical Report on World Vitiviniculture, International Organisation of Vine and Wine
Bulk Red Wine Scion Role
Airén bears a Bulk_White_Wine_Scion_Role as the world's most planted white variety by area (though declining), yielding 15-30 t/ha on the arid central Spanish plateau to produce neutral wines for domestic consumption, blending, and brandy production at €1-3/bottle. Trebbiano Toscano (Ugni Blanc) serves in a Bulk_White_Wine_Scion_Role across central Italy and southwestern France, producing high-acid, neutral wines at yields exceeding 15 t/ha for Cognac distillation and everyday table wine. Pinot Grigio bears a Bulk_White_Wine_Scion_Role in the Veneto's high-yielding plains (Delle Venezie DOC), producing commercial wines at 15-20 t/ha for international markets, contrasting with premium Pinot Grigio from Friuli or Alto Adige at half those yields.
A white wine scion role that inheres in a grafted scion cultivar by virtue of its designation for high-volume, commodity-oriented white wine production, where viticultural and winemaking practices prioritize yield, consistency, and cost-efficiency to produce wines for everyday consumption at accessible price points. Bulk white wine scions are cultivated to achieve high yields (typically 12-25 t/ha, with some irrigated regions exceeding 30 t/ha) while maintaining commercial soundness, acceptable varietal character, and microbial stability. Bulk production favors vigorous, productive cultivars tolerant of warm climates (maintaining acidity despite heat), mechanical harvesting, and industrial-scale vinification with minimal hand labor. Wines are typically fermented in temperature-controlled stainless steel, released within months of harvest without extended aging, often sweetened (1-15 g/L residual sugar) for mass-market appeal, and priced for volume markets (typically below €5-8/bottle retail). Major bulk white wine production occurs in Spain (Airén in La Mancha), Italy (Trebbiano in central regions), southern France (Colombard, Ugni Blanc), California's Central Valley (Chardonnay, French Colombard), and Australia's inland regions.
Bulk_White_Wine_Scion_Role represents volume production essential for supplying everyday white wine consumption globally. Many white varieties span bulk and premium production depending on viticulture—Chardonnay ranges from €2 Central Valley wines (25 t/ha) to €500 Montrachet (5 t/ha) from the same varietal designation. Bulk white wine production increasingly focuses on internationally recognized varieties (Chardonnay, Sauvignon Blanc, Pinot Grigio) grown at high yields for varietal-labeled wines rather than traditional anonymous blending varieties. Climate is a critical factor warm-climate bulk production often requires acidification to compensate for acid loss during ripening at high yields.
Großmengenweißweinsorte
Keltertraube für weißen Massenwein
Massenweißweinsorte
bulk white wine variety
commodity white wine variety
high-yield white wine cultivar
volume white wine variety
Anderson K, Aryal NR (2020) Database of Regional, National and Global Winegrape Bearing Areas by Variety, Wine Economics Research Centre, University of Adelaide Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane OIV (2019) Statistical Report on World Vitiviniculture, International Organisation of Vine and Wine
Bulk White Wine Scion Role
Cinsault vines in the Languedoc-Roussillon bearing a bulk wine role are cultivated in mechanically managed bush-trained plantings at yields of 80-120 hl/ha exceeding AOC limits and therefore declassified to Vin de France or Vin sans IG status, harvested mechanically at 10-11° potential alcohol, sold in 24,000-liter flexitanks at 0.40-0.70 €/liter ex-cellar to négociants and bottling operations in northern Europe for private-label supermarket wines, with global bulk wine trade volumes estimated at 35-40 million hectoliters annually representing approximately 15% of total world wine production.
A quality tier role that inheres in a grapevine whose fruit is produced for the unbranded bulk wine market, sold in large-volume containers (flexitanks, tanker trucks, ISO containers) without consumer-facing labeling, destined for blending, private-label bottling, industrial processing, or distillation at the lowest price tier of the commercial wine chain.
Bulk_Wine_Role represents the commodity end of the wine market where the vine's fruit enters an undifferentiated supply chain with no traceability to specific vineyards, producers, or vintage in the final bottled product. The role is defined by the absence of provenance identity in the commercial chain rather than by inherently low fruit quality: viticultural conditions at bulk-producing sites may produce sound, clean fruit that is commercially competent but economically allocated to the lowest-value channel. Major bulk wine trading flows include southern France to northern Europe, Spain to multiple destinations, Chile and Argentina to export markets, and Australia's inland regions to domestic and Asian markets.
Fasswein-Rolle
Industriewein-Rolle
Schüttgutwein-Rolle
bulk wine vine role
commodity wine role
industrial wine role
OIV (2023) State of the World Vine and Wine Sector Ciatti Company (2023) Global Wine Market Update Anderson K (2004) The World's Wine Markets: Globalization at Work, Edward Elgar Publishing pp: 62-85
Bulk Wine Role
Bunch stem necrosis quality assessment for calcium deficiency in Australian Shiraz identified the disorder on a site with marginal calcium availability exacerbated by high evaporative demand limiting xylem calcium delivery to fruit clusters. Cluster evaluation at véraison revealed bunch stem necrosis quality affecting 18% of clusters, with symptoms showing the characteristic Ca-associated pattern of tip-to-base progression (acropetal to basipetal along the rachis) distinct from the branch-point concentration of Mg-associated grape stem necrosis. Affected clusters showed bunch stem necrosis quality with rachis tissue collapse beginning at the cluster apex, brown necrosis spreading basipetally over 2-3 weeks, and terminal berries shriveling and dropping while basal portions remained functional initially. Timing analysis confirmed the Ca-association with bunch stem necrosis quality appearing at 3 weeks post-fruit set, earlier than typical Mg-associated symptoms that develop post-véraison. Nutritional assessment showed petiole Ca at 1.0% (marginal) but importantly showed rachis Ca at only 0.3% versus 0.8% in unaffected clusters from the same vines, indicating that Ca delivery to rachis tissue rather than whole-vine Ca status was the limiting factor. Environmental analysis revealed that the bunch stem necrosis quality concentrated in positions with high cluster exposure and associated high berry transpiration, where xylem Ca preferentially followed the transpiration stream to leaves rather than the relatively low-transpiring rachis tissue. Corrective management with pre-véraison foliar CaCl₂ applications (0.5% solution, 3 applications at 7-day intervals) targeting cluster zone reduced bunch stem necrosis quality incidence to 5% the following season.
Bunch stem necrosis quality in calcium deficiency is a calcium deficiency symptom quality that inheres in grapevine cluster rachis tissue and represents the physiological disorder characterized by collapse and necrosis of rachis and pedicel tissue during fruit development resulting from calcium-limited cell wall integrity, with symptoms typically initiating at the cluster tip and progressing basipetally toward the peduncle, causing berry shrivel, detachment, and cluster desiccation that can result in significant crop loss, and distinguished from the magnesium-associated grape stem necrosis (Stiellähme) by its different distribution pattern, predisposing factors, and timing. This quality captures a distinct calcium-related disorder, recognizing that while both calcium and magnesium deficiencies can cause rachis necrosis, the Ca-associated form shows characteristic features enabling differential diagnosis and informing appropriate corrective management. The ontological significance of bunch stem necrosis quality in calcium deficiency lies in its economic impact through crop loss and its indication of calcium nutrition or transport limitations that may also affect other quality parameters, as well as the importance of distinguishing it from Mg-associated grape stem necrosis to enable correct diagnosis and treatment. The biological basis of calcium-deficiency bunch stem necrosis encompasses the role of calcium in pectin cross-linking that provides mechanical strength to cell walls, the vulnerability of rapidly expanding rachis tissue during early berry development when cell wall synthesis is most active, and the limited calcium supply to rachis tissue through xylem transport that may be inadequate during periods of high transpiration from leaves competing for xylem calcium delivery. The timing of Ca-associated bunch stem necrosis typically shows earlier development than Mg-associated grape stem necrosis, often appearing during the berry set to pre-véraison period when rachis cell expansion is most active and calcium demand highest, though symptoms may continue to develop through ripening if conditions remain unfavorable for calcium supply.
Bunch_Stem_Necrosis_Quality_(Ca) is_a Calcium_Deficiency_Symptom_Quality, representing the rachis collapse disorder from Ca-limited cell wall integrity. This class addresses a Ca-associated cluster disorder distinct from Mg-associated Stiellähme. Tip-to-base progression distinguishes from branch-point concentration of Mg disorder. Earlier timing (fruit set to véraison) than Mg-associated symptoms (véraison to harvest). Rachis Ca concentration more diagnostic than whole-vine Ca status. High evaporative demand diverts xylem Ca to leaves away from rachis. Foliar Ca application to clusters provides targeted correction.
BSN
Ca-induced rachis necrosis
Ca-induzierte Rappennekrose
Kalzium-Stiellähme
Traubenstiel-Kollaps
Traubenstiellähmung bei Ca-Mangel
calcium bunch stem disorder
cluster stem collapse
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Krasnow MN et al. (2010) Evidence for substantial maintenance of membrane integrity and cell viability in normally developing grape (Vitis vinifera L.) berries throughout development. Journal of Experimental Botany 59:849-859 White PJ, Broadley MR (2003) Calcium in plants. Annals of Botany 92:487-511
Bunch Stem Necrosis Quality (Ca)
A bush head arm on Vitis vinifera cv. Assyrtiko on volcanic soils of Santorini, Greece, one of six irregularly-positioned arms emerging from swollen head at ground level (head barely 15cm above soil), arms extending 35-65cm in various directions creating low spreading bush approximately 80cm diameter and 40cm height, traditional kouloura (basket) training protecting clusters within tangled arm and shoot structure from strong Aegean winds, total vine bearing 28 spurs distributed irregularly across six arms supporting approximately 48 shoots and 4.2kg crop, representing traditional bush head architecture adapted to extreme environmental conditions (strong winds, minimal rainfall, volcanic soils) where irregular low bush form provides protection and drought tolerance impossible with conventional upright training systems.
A head-trained arm in generalized bush or shrub configuration without specific gobelet architecture, characterized by multiple arms radiating from central head creating irregular bush-like form, variable arm lengths and angles creating informal structure, minimal or absent trellis support creating self-supporting bush vine, and flexible architecture adapting to vine vigor, terrain constraints, or traditional practices in diverse regions worldwide. Bush head arms represent broad category encompassing various head-training approaches beyond formal gobelet, including irregular bush vines in old Mediterranean vineyards, adapted head training in challenging terrain (steep slopes, rocky soils), traditional systems in diverse global regions (Greece, Portugal, South Africa, Australia, South America), and transitional or degraded gobelet structures that have lost formal architecture over time through irregular renewal or damage recovery.
Bush head arms encompass diverse informal head-training approaches unified by multi-arm radiating architecture and self-supporting bush habit but lacking standardized formal structure of gobelet or other named systems. The informal architecture often reflects adaptation to challenging conditions (extreme slopes preventing trellis installation, poor soils limiting vine vigor necessitating compact structure, harsh climates requiring protective low bush form), historical evolution in regions without strong training traditions, or degradation of formal systems over time through inconsistent renewal. Bush head training provides advantages in challenging situations including no trellis requirements (reducing establishment costs and infrastructure maintenance), adaptation to irregular terrain (individual vine positioning without row constraints), drought tolerance (limited canopy reducing water demand), and flexibility accommodating variable vine vigor. However, bush training severely limits mechanization, creates labor-intensive hand operations, produces typically low yields, and complicates disease management through dense tangled structures with poor air circulation.
Busch-Arm
free-form head arm
freie Buschform-Arm
informal bush arm
irregular head arm
unregelmäßige Kopfarm
Bush Head Arm
A 120-year-old Shiraz bush vine in Barossa Valley's Eden Valley subregion bears a Bush_Vine_Role, its gnarled trunk supporting an irregular crown of spurs producing <2 kg of intensely concentrated fruit per vine—the bush form representing pre-phylloxera planting practices and commanding premium pricing for "old vine" designation. A Chenin Blanc bush vine in Swartland's dryland conditions serves in a Bush_Vine_Role, the self-supporting form adapted to 350 mm annual rainfall without irrigation, producing low yields of concentrated fruit for the region's distinctive dry white wines. A Cinsault bush vine in the Cape's hot Stellenbosch foothills bears a Bush_Vine_Role, the spreading canopy providing self-shading for this heat-sensitive variety under conditions that would stress trellised vines.
A head-trained vine role that inheres in a grapevine grown as a free-standing bush without formal arm structure, with fruiting canes or spurs arising from a head in a less structured arrangement than formal gobelet, typically resulting in a spreading, informal canopy shape adapted to challenging growing conditions. Bush vine training represents the least infrastructure-intensive viticultural approach, requiring no trellis, minimal training in early years, and tolerance of vine-to-vine variation in form that would be unacceptable in more formal systems. Bush vines are particularly associated with old-vine vineyards in Australia (Barossa, McLaren Vale), South Africa (Swartland), California (Lodi, Sonoma), and Mediterranean regions where surviving pre-modern plantings retain traditional training approaches. The bush vine designation often carries quality connotations—low yields from mature, deep-rooted vines producing concentrated, terroir-expressive fruit—though the term may also describe any informal head-trained vine regardless of age. Bush vine management accepts canopy irregularity, with individual vines developing distinctive forms based on their specific growth patterns, unlike the uniformity sought in formal training systems.
Bush_Vine_Role captures the informal head-trained category, distinguished from formal gobelet by the absence of defined arm structure and tolerance of greater vine-to-vine variation. The "bush vine" term has acquired marketing significance, particularly in Australian and South African contexts where it signals old-vine character, dryland authenticity, and premium positioning. Bush vine vineyards often represent surviving remnants of historical planting practices, with the informal training reflecting labor patterns and viticultural knowledge of earlier eras. Contemporary bush vine management may impose more structure than historic practice—renewal pruning to maintain balance, selective shoot positioning, limited canopy management—while retaining the fundamental self-supporting architecture. The distinction between bush vine and gobelet is not always clear the same vineyard might be described as either depending on regional terminology and the degree of formality in training.
Buschform
Buschrebe
Sprawl-Erziehung
Trockenland-Busch
bush training
bush vine
dryland bush
sprawl training
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Dry PR (2000) Canopy management for fruitfulness, Australian Journal of Grape and Wine Research 6: 109-115 Iland P et al. (2011) The Grapevine: From the Science to the Practice of Growing Vines for Wine, Patrick Iland Wine Promotions
Bush Vine Role
Caftaric acid content quality in Riesling provides a key determinant of must oxidation trajectory, with typical concentrations of 80-150 mg/L creating the browning substrate that must be managed through protective handling German winemakers traditionally employ must sulfiting (50-80 mg/L SO₂) immediately after pressing to inhibit PPO and preserve caftaric acid in reduced form, maintaining the pale green-gold must color that indicates minimal oxidative damage and predicts superior wine freshness and aging potential.
Caftaric_Acid_Content_Quality represents the chemical quality characterizing the concentration of caftaric acid (2-caffeoyl-L-tartaric acid, trans-caffeoyltartaric acid) in grape berry tissue, capturing the dominant hydroxycinnamic acid derivative in Vitis vinifera grapes. Caftaric acid consists of caffeic acid (3,4-dihydroxycinnamic acid) esterified to the 2-position of L-tartaric acid, combining the phenylpropanoid moiety with grape's signature organic acid. This compound typically comprises 40-60% of total hydroxycinnamic acids in grape juice, with concentrations ranging from 50-200 mg/L depending on cultivar, vintage, and extraction conditions. Caftaric acid is the preferred substrate for grape polyphenol oxidase (PPO), undergoing rapid oxidation upon tissue disruption to form caftaric acid o-quinone. This quinone is highly reactive, capable of polymerizing to form brown pigments, reacting with glutathione to form grape reaction product (GRP, 2-S-glutathionyl caftaric acid), or oxidizing other phenolics through coupled oxidation reactions. The rate and extent of caftaric acid oxidation during must handling critically affects juice color and phenolic composition. Glutathione-rich musts convert caftaric acid quinone to the colorless, stable GRP rather than brown polymers, explaining why sulfur nutrition and grape glutathione content affect browning susceptibility. In wines that have not undergone extensive oxidation, caftaric acid persists as a major phenolic component, potentially undergoing slow hydrolysis to release free caffeic acid during aging. The caffeic acid moiety can serve as substrate for microbial reduction to volatile phenols, making caftaric acid an indirect precursor for vinylphenol formation.
Caftaric_Acid_Content_Quality specifically addresses the caffeic acid tartrate ester that dominates grape hydroxycinnamic acid composition. The oxidation chemistry of caftaric acid through quinone formation explains its central role in juice browning, making caftaric acid content a predictor of browning susceptibility rather than a direct quality attribute. The conversion to GRP through glutathione conjugation represents a detoxification pathway that protects other phenolics from coupled oxidation. Analysis employs HPLC with UV detection at 320 nm, with caftaric acid well-resolved from other hydroxycinnamic acid esters.
Caftarsäuregehalt
GRP precursor quality
GRP-Vorläuferqualität
Kaffeoyltartrat-Konzentration
Kaffeoylweinsäure-Gehalt
caffeoyl tartrate content
caffeoyltartaric acid concentration
caftaric acid level
Singleton VL et al. (1984) Caftaric Acid in Grapes and Conversion to a Reaction Product during Processing. Vitis 23(1): 27-40 Cheynier V et al. (1989) Characterization of 2-S-Glutathionyl Caftaric Acid and Its Hydrolysis in Relation to Grape Wines. Journal of Agricultural and Food Chemistry 37(4): 1069-1071
Caftaric Acid Content Quality
The calcium content of a Champagne vineyard on Campanian chalk documents very high exchangeable calcium of 35-45 cmol(+)/kg (7,000-9,000 mg/kg) — the calcium-saturated exchange complex reflects the calcareous parent material (>80% CaCO₃) that maintains calcium as the overwhelmingly dominant exchangeable cation (>90% of CEC), buffering the soil pH firmly at 8.0-8.3 and providing effectively unlimited calcium supply for vine uptake. This calcium abundance creates the distinctive nutritional environment of calcareous vineyard soils — the competition between calcium, potassium, and magnesium at root uptake sites (mediated by the Ca:K and Ca:Mg ratios on the exchange complex) restricts potassium uptake relative to calcium uptake, contributing to the lower must potassium and consequently lower must pH that characterizes wines from calcareous terroirs compared to wines from acid soils with lower calcium dominance. The very high calcium content of the Champagne chalk soils contrasts with the low exchangeable calcium (2-5 cmol(+)/kg) of acid granite-derived soils in parts of Galicia and the Minho (Portugal) where calcium leaching under high rainfall has depleted the exchange complex.
A nutrient analysis data component that records the concentration of exchangeable calcium in the vineyard soil, determined by ammonium acetate extraction (1M NH₄OAc at pH 7.0) and expressed in cmol(+)/kg or mg/kg, with calcium being the dominant exchangeable cation in most vineyard soils (typically occupying 60-85% of the exchange complex), serving as the primary soil structural stabilizer (promoting clay flocculation and aggregate stability), the major pH buffer in calcareous soils, and an essential nutrient for vine cell wall construction, membrane integrity, and fruit quality — though true calcium deficiency is rare in vineyard soils due to the abundance of calcium in most geological substrates.
Calcium_Content documents the base cation that dominates the exchange chemistry of most viticultural soils and whose abundance or scarcity fundamentally shapes the soil chemical environment. In calcareous soils (those containing free calcium carbonate, CaCO₃), the calcium content is effectively buffered at high levels by the dissolution equilibrium of the carbonate mineral — as plant uptake or leaching removes calcium from the exchange complex, CaCO₃ dissolution replenishes the exchangeable pool, maintaining calcium saturation until the carbonate reserve is exhausted (a process requiring millennia under normal weathering conditions): in non-calcareous soils, exchangeable calcium is maintained by the weathering of calcium-bearing primary minerals (plagioclase feldspar, pyroxenes, amphiboles) and by the return of calcium in organic matter cycling, with the calcium saturation declining progressively as weathering and leaching advance under humid conditions — the most intensely weathered tropical soils retaining only 10-20% calcium saturation compared to 80-95% in young calcareous soils. The viticultural significance of calcium extends beyond direct nutritional supply (vine calcium requirements are modest and easily met by most soils) to the indirect effects of calcium on potassium uptake — the Ca:K ratio on the exchange complex influences the relative rates of calcium and potassium uptake by vine roots, with high Ca:K ratios (>20:1) restricting potassium uptake and potentially reducing must pH, an effect exploited in the selection of calcareous sites and lime-tolerant rootstocks for quality-focused viticulture.
Boden-Calcium
Ca-Gehalt-Messung
Calciumgehalt
calcium availability assessment
exchangeable Ca measurement
soil calcium record
White RE (2003) Soils for Fine Wines, Oxford University Press Mpelasoka BS et al. (2003) A review of potassium nutrition in grapevines, Australian Journal of Grape and Wine Research 9: 154-168 Soil Survey Staff (2014) Keys to Soil Taxonomy, 12th edition, USDA-NRCS
Calcium Content
Calcium content quality in Champagne base wines reflects the region's famous chalk soils, with must calcium concentrations of 100-180 mg/L from Côte des Blancs vineyards on pure Cretaceous chalk versus 60-100 mg/L from Montagne de Reims sites on clay-limestone while the sensory contribution of this calcium difference remains debated, the correlation between chalk terroir, elevated calcium, and the perceived chalky, mineral texture of great Blanc de Blancs Champagne supports the terroir narrative linking soil composition through grape minerals to wine character.
Calcium_Content_Quality represents the chemical quality characterizing the concentration of calcium (Ca²⁺) in grape berry tissue, capturing the divalent cation essential for cell wall structure, membrane integrity, and fruit firmness. Calcium in grape berries ranges from 50-200 mg/L in juice, substantially lower than potassium but with distinct quality implications related to tissue integrity, tartrate stability, and potential sensory effects. Unlike potassium which accumulates via phloem, calcium enters berries predominantly through xylem transpirational flow during the pre-véraison period when berries are still transpiring actively. After véraison, the shift from xylem to phloem supply effectively stops calcium accumulation, meaning that berry calcium content is largely determined before ripening begins. This early-determined calcium pool binds extensively to cell wall pectins, contributing to tissue firmness and structural integrity that affects postharvest quality in table grapes and processing behavior in wine grapes. High calcium content in berries contributes to firmer texture valued in table grapes, while very high calcium can cause calcium tartrate precipitation in wine—a less common stability problem than potassium bitartrate but occurring in wines from calcium-rich soils. The calcium-to-magnesium ratio in soil affects uptake of both elements through competitive inhibition, with high-magnesium soils potentially limiting calcium nutrition. Limestone-derived soils provide abundant calcium, potentially contributing to the perceived mineral character associated with calcareous terroirs though direct sensory effects of calcium remain unproven. Calcium analysis employs atomic absorption spectroscopy or ICP-OES following appropriate sample preparation.
Calcium_Content_Quality addresses a mineral with distinct behavior from potassium due to its xylem-mediated accumulation that ceases at véraison. This early accumulation pattern means calcium content reflects pre-véraison conditions and cannot be modified by late-season management. Calcium's structural role in cell walls (binding pectin, providing rigidity) distinguishes its function from the osmotic and enzymatic roles of potassium. Calcium tartrate instability is less common than potassium bitartrate instability but can occur in wines from high-calcium terroirs, requiring cold stabilization or ion exchange treatment.
Beerencalciumqualität
Ca content
Ca-Gehalt
Calciumkonzentration
berry calcium quality
calcium concentration
calcium level
Rogiers SY et al. (2006) Mineral Sinks within Ripening Grape Berries (Vitis vinifera L.). Vitis 45(3): 115-123 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 341-385
Calcium Content Quality
Calcium deficiency symptom quality diagnosis in newly established Marlborough Sauvignon Blanc identified Ca limitation as the cause of tip burn symptoms and cluster disorders on an acidic site with soil pH 4.8 and exchangeable Ca at only 450 ppm. Visual calcium deficiency symptom quality assessment revealed characteristic tip burn on young leaves with brown necrosis at leaf tips and margins curling downward, affecting the apical 4-5 leaves on actively growing shoots while older leaves remained symptom-free, consistent with the immobile nutrient pattern affecting newest tissues. Cluster calcium deficiency symptom quality appeared as bunch stem necrosis with rachis tissue showing collapse and browning beginning at the cluster tip and progressing basipetally toward the peduncle, distinct from the Mg-associated grape stem necrosis pattern which typically shows concentration at branch points. Tissue analysis confirmed calcium deficiency symptom quality diagnosis with petiole Ca at 0.8% (deficient, threshold 1.2%) despite adequate soil calcium in total, with the low soil pH limiting Ca availability and high soil aluminum at 2.5 cmol/kg potentially interfering with Ca uptake. Root assessment revealed restricted root development attributed to Al toxicity at the low pH, compounding the calcium deficiency symptom quality by limiting the root system's capacity to acquire Ca from soil. Corrective lime application at 4 t/ha to raise pH above 5.5 and supply Ca, combined with foliar CaCl₂ application for immediate symptom relief, resolved calcium deficiency symptom quality within one season as pH correction improved both Ca availability and root function.
Calcium deficiency symptom quality is a macronutrient deficiency symptom quality that inheres in a grapevine organism and represents the characteristic visual symptoms and physiological disorders resulting from inadequate calcium availability, including necrosis of young leaf tips and margins (tip burn), bunch stem necrosis affecting cluster rachis integrity, reduced cell wall strength causing tissue collapse, and impaired membrane function affecting overall tissue quality, with calcium deficiency symptoms appearing on youngest tissues and growing points reflecting the phloem immobility of calcium that prevents remobilization from older organs. This quality captures the grapevine response to calcium limitation, recognizing that while calcium deficiency is relatively uncommon in field conditions due to adequate soil calcium in most viticultural regions, it can occur on highly acidic soils and creates characteristic symptoms affecting newest growth that distinguish it from mobile nutrient deficiencies. The ontological significance of calcium deficiency symptom quality lies in the critical structural and signaling roles of calcium in plant function, where deficiency affects cell wall integrity, membrane function, and signal transduction with consequences for tissue structure and stress tolerance, and in the distinctive symptom distribution on young tissues that provides diagnostic differentiation from deficiencies of mobile nutrients. The biological basis of calcium deficiency symptoms encompasses the structural role of calcium in cell wall pectin cross-linking providing mechanical strength, the function of calcium in membrane structure and stability, and the signaling role of calcium as a second messenger in numerous cellular responses, such that calcium deficiency produces symptoms reflecting compromised structural integrity and impaired cellular function. The phloem immobility of calcium results from its sequestration in cell walls and vacuoles in forms that cannot be remobilized, meaning that young tissues must acquire calcium from current xylem supply and cannot draw on reserves in older tissues, creating vulnerability to deficiency symptoms when root uptake or xylem transport is inadequate regardless of whole-plant calcium status.
Calcium_Deficiency_Symptom_Quality is_a Macronutrient_Deficiency_Symptom_Quality, serving as parent class for Bunch_Stem_Necrosis_Quality_(Ca) and Tip_Burn_Quality_(Ca) which address specific calcium deficiency manifestations. This class addresses the overall calcium deficiency syndrome while child classes capture particular symptom types. Immobile nutrient showing symptoms on youngest tissues first (opposite of N, K, Mg). Cell wall structure and membrane function affected. Petiole Ca threshold ~1.2% at bloom. Uncommon in field conditions but occurs on acidic soils where Ca availability is limited. Aluminum toxicity at low pH compounds Ca deficiency.
Ca deficiency symptoms
Ca-Mangel-Zeichen
Kalziummangel-Symptome
Kalziumstress-Indikatoren
Niedrig-Kalzium-Symptome
calcium limitation signs
calcium stress indicators
low calcium symptoms
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication White PJ, Broadley MR (2003) Calcium in plants. Annals of Botany 92:487-511
Calcium Deficiency Symptom Quality
Calcium uptake in grapevines ranges from 40-100 kg Ca/ha annually depending on soil availability and vine size, with relatively modest rootstock effects (15-30% variation) compared to potassium. Calcium moves primarily in the xylem with the transpiration stream, resulting in preferential accumulation in transpiring leaves (2-4% Ca dry weight) versus low-transpiring berries (0.02-0.05% juice). Petiole calcium concentration at bloom of 1.2-2.5% dry weight indicates adequate supply, with deficiency symptoms (marginal leaf necrosis) appearing below 0.8%. Calcicole rootstocks such as 41B and Fercal maintain Ca uptake efficiency on high-lime soils where calcifuge rootstocks show reduced absorption due to pH-impaired root function.
A macronutrient uptake disposition that is the inherited capacity of a Vitis vinifera plant to absorb calcium ions (Ca²⁺) from the soil solution through calcium channels and transporters, with uptake driven by transpiration stream transport and genotype-specific root membrane permeability.
Calcium uptake is less dependent on genotype than potassium uptake due to the predominantly passive nature of Ca transport in the transpiration stream. However, rootstock effects on root system health in calcareous soils indirectly affect Ca acquisition through overall root function. Calcium is immobile in phloem, making continuous uptake essential for developing tissues. Cell wall structural roles and signaling functions require adequate Ca supply throughout the growing season.
Ca uptake capacity
Ca-Akquisition
Calciumaneignungsvermögen
Calciumaufnahme
calcium acquisition
calcium foraging trait
Conradie WJ (1981) Seasonal uptake of nutrients by Chenin blanc in sand culture: II. Phosphorus, potassium, calcium and magnesium. South African Journal of Enology and Viticulture 2:7-13 White PJ, Broadley MR (2003) Calcium in plants. Annals of Botany 92:487-511
Calcium Uptake Disposition
Cabernet Franc cutting showing callus-derived adventitious roots, cutting base developing extensive white callus tissue 8-12 days after preparation, microscopic examination revealing root primordia organizing within callus tissue 14-16 days, adventitious roots emerging from callus surface 18-22 days, cutting producing 11 roots by day 28= 8 emerging from basal callus (callus-derived) and 3 from stem nodes (shoot-derived), callus-derived roots requiring vascular differentiation through callus tissue connecting to cutting vascular system
Callus derived adventitious root is adventitious root originating from callus tissue formed at wounds, graft unions, or cutting bases, characterized by development from undifferentiated or partially differentiated callus cells that organize into root primordia, showing indirect developmental pathway involving wound response, callus proliferation, and subsequent root organogenesis, demonstrating formation typically at interfaces between plant tissues and environment such as cutting bases or graft unions, and serving functions including wound healing through root formation, graft union stabilization through root development connecting scion and rootstock, and propagation through cutting base rooting, with callus-derived roots showing typical developmental sequence of wounding triggering dedifferentiation and cell proliferation creating callus mass, hormonal signals inducing subpopulations of callus cells to organize into root primordia, and developing roots emerging from callus surface creating less direct vascular connection to parent plant than shoot-derived roots requiring vascular differentiation within callus, and demonstrating that callus intermediary potentially provides advantage of creating numerous pluripotent cells from which many roots can form but disadvantage of indirect vascular connection potentially delaying functional integration. Callus derived adventitious root developmental pathway includes= showing wound response (initial injury or cutting triggering dedifferentiation and cell proliferation), exhibiting callus proliferation (undifferentiated parenchyma-like cells multiplying creating callus tissue mass), demonstrating hormonal induction (auxin and cytokinin signals inducing root organogenesis in callus subpopulations), revealing primordium organization (callus cells organizing into structured root primordia with meristems), and displaying root emergence (organized roots growing from callus surface). Callus derived adventitious root callus characteristics includes= showing parenchymatous structure (callus comprising relatively undifferentiated parenchyma-like cells), exhibiting meristematic capacity (callus cells retaining or regaining ability to divide and differentiate), demonstrating variable cell types (callus potentially containing cells with different developmental potentials), revealing vascular islands (callus sometimes developing isolated vascular elements or tracheary elements), and displaying pluripotency (callus cells capable of differentiating into various tissue types including roots under appropriate signals). Callus derived adventitious root at cutting bases includes= showing basal callus formation (callus developing at cut base of stem cuttings within days of preparation), exhibiting root emergence from callus (adventitious roots forming within and emerging from basal callus tissue), demonstrating callus-mediated rooting (some difficult-to-root cultivars showing predominantly callus-derived rather than direct stem-derived roots), revealing callus abundance correlation (extensive callus often associated with higher numbers of adventitious roots), and displaying vascular integration challenge (callus roots requiring establishment of vascular connection through callus to cutting vascular system). Callus derived adventitious root at graft unions includes= showing graft interface callus (callus forming between scion and rootstock at graft union), exhibiting bridging function (callus tissue potentially producing roots that bridge between scion and rootstock), demonstrating union stabilization (callus-derived roots strengthening graft union mechanical connection), revealing vascular continuity (callus roots potentially contributing to vascular connection between scion and rootstock), and displaying occasional occurrence (graft union roots not always present but occurring in some union types or with some compatibility issues). Callus derived adventitious root hormonal regulation includes= requiring auxin induction (high auxin levels promoting both callus formation and subsequent root organogenesis), showing cytokinin effects (moderate cytokinins potentially promoting callus proliferation while high levels inhibiting root formation), exhibiting auxin-to-cytokinin ratio importance (optimal ratio favoring root versus shoot organogenesis in callus), demonstrating hormone application effects (sequential hormone treatments sometimes optimizing callus production then root induction), and revealing endogenous hormone dynamics (callus cells showing changes in hormone synthesis and sensitivity during root organogenesis). Callus derived adventitious root vascular development includes= showing indirect connection (callus roots connecting to parent plant through callus tissue requiring vascular differentiation), exhibiting vascular island origin (roots potentially connecting to vascular elements within callus), demonstrating delayed integration (vascular connection establishment potentially slower than direct shoot-derived roots), revealing functional consequences (indirect vascular path potentially affecting early root function and plant establishment), and displaying eventual normalization (once established, callus-derived roots showing normal vascular function). Callus derived adventitious root advantages and disadvantages includes= showing higher number potential (callus providing large population of pluripotent cells from which many roots can form), exhibiting tissue repair (callus formation contributing to wound healing beyond just root production), demonstrating cultivation challenges (excessive callus without roots sometimes problematic in propagation), revealing delayed establishment (indirect vascular connection potentially slowing plant establishment compared to direct shoot roots), and displaying cultivar patterns (some cultivars showing predominantly callus-mediated rooting while others mainly direct). Callus derived adventitious root practical implications includes= affecting propagation protocols (callus formation and subsequent rooting requiring appropriate hormone concentrations and timing), influencing cutting management (basal callus development monitored as indicator of rooting potential), demonstrating graft compatibility indicators (graft union callus and root development potentially indicating compatibility or union quality), revealing rootstock evaluation (rootstock propagation sometimes assessed on callus formation capacity), and displaying commercial significance (understanding callus-mediated rooting important for optimizing propagation efficiency and success rates).
Callus_Derived_Adventitious_Root emphasizes callus tissue intermediary in root formation and is distinguished from direct shoot-derived roots by focus on indirect pathway. Includes Wound_Callus_Adventitious_Root and Graft_Callus_Adventitious_Root. Critical for understanding callus pathway, wound response, indirect vascular connection, graft unions, cutting base rooting, and the adventitious roots arising from callus tissue requiring differentiation through callus intermediary.
Callus-mediated root
Indirect adventitious root
Indirekte Adventivwurzel
Kallus-Adventivwurzel
Wound callus root
Wund-Kallus-Wurzel
Bellini, C., et al. (2014). Adventitious roots and lateral roots= Similarities and differences. Annual Review of Plant Biology, 65, 639-666.
Callus Derived Adventitious Root
Callus formation capacity varies among grapevine genotypes, with vigorous rootstocks such as 1103 Paulsen and SO4 showing rapid callus initiation (visible within 3-5 days at 25°C) and abundant proliferation (2-4 mm callus thickness by day 14), while less vigorous types like Riparia Gloire show slower response (5-7 days to visible callus, 1-2 mm by day 14). Scion callusing ability also varies, with Cabernet Sauvignon and Merlot showing strong callus response versus Pinot Noir with moderate callusing. Callus cells dedifferentiate from cambium, phloem parenchyma, and ray cells adjacent to the cut surface, with cell division rates of 1-2 divisions per day under optimal conditions. Callus quality (firm, white, uniform) correlates with subsequent vascular connection success (r = 0.70-0.85).
A graft union formation disposition that is the inherited capacity of a Vitis vinifera graft partner to produce callus tissue (parenchymatous wound tissue) at cut surfaces in response to grafting injury, with genotype-specific rates of callus initiation, proliferation, and the capacity for callus cells to intermingle with partner callus across the graft interface.
Callus formation represents the first visible step in graft union establishment, creating the cellular matrix within which vascular reconnection will occur. Adequate callus volume is necessary but not sufficient for compatibility, as callus cells must also show appropriate adhesion and competence for vascular redifferentiation. Callusing conditions (temperature, humidity, oxygen) strongly influence callus quantity and quality. Excessive callusing can impair union quality by displacing graft partners or creating excessive tissue mass without functional connections.
Kallusbildung
Kallusierungsvermögen
Wundgewebeproliferation
callus proliferation capacity
callusing ability
wound healing response
Cookson SJ, Ollat N (2013) Grafting with rootstocks induces extensive transcriptional re-programming in the shoot apical meristem of grapevine. BMC Plant Biology 13:147 Zarrouk O et al. (2010) Grapevine rootstocks differentially influence the rate of ripening and modulate auxin-related genes in Cabernet Sauvignon berries. Australian Journal of Grape and Wine Research 16:355-363
Callus Formation Disposition
Pinot Noir on Riparia Gloire rootstock callus graft union zone at 8 weeks post-grafting shows 18-28 mm zone dominated by undifferentiated callus tissue (cream-colored soft parenchyma comprising 70-82% of union zone volume), callus cells 35-75 μm diameter with thin walls and large vacuoles proliferating from cambial regions on both scion and rootstock, intermixed scion and rootstock callus forming continuous bridge, early vascular differentiation visible with scattered vessel elements 80-180 μm diameter and isolated sieve tubes beginning to form in callus matrix, localized cambial differentiation creating small patches of organized tissue, union showing limited transport capacity (30-50% of mature healed union) and mechanical strength requiring support, represents early healing stage with progressive differentiation expected over subsequent months
Callus graft union zone is graft union zone dominated by callus tissue comprising undifferentiated and partially differentiated parenchyma cells that proliferated from cambial regions and adjacent tissues forming the initial tissue bridge between scion and rootstock, representing the early-stage union or persistent callus-rich union characteristic of healing or poorly integrated grafts. Callus graft union zone tissue composition includes= undifferentiated callus parenchyma (thin-walled isodiametric cells 30-80 μm diameter lacking specialized structure, comprising 60-85% of union zone volume in young unions or incompatible grafts), partially differentiated tissues (cells showing initial specialization toward xylem, phloem, or cambial fates but not fully functional), scattered vascular elements (isolated or poorly connected xylem vessels and phloem sieve tubes embedded in callus matrix), discontinuous cambium (cambial cells present but not forming complete functional layer), and minimal secondary xylem and phloem (limited organized vascular tissues compared to healed unions with extensive secondary growth). Callus tissue origin and formation includes= arising from cambial region (vascular cambium and adjacent immature xylem and phloem cells dedifferentiating and proliferating), arising from mature tissues (mature xylem parenchyma, phloem parenchyma, cortex cells potentially dedifferentiating contributing to callus formation), rapid cell division (callus cells dividing 1-3 times per week during active formation creating rapidly expanding tissue mass), cell expansion (callus cells enlarging 2-4 fold after division), occurring on both partners (scion and rootstock independently producing callus from cut surfaces), and intermixing (scion and rootstock callus cells meeting, contacting, adhering forming continuous tissue bridge across junction). Callus graft union zone in early-stage healing includes= visible within 1-2 weeks (white to cream colored soft tissue appearing at graft interface), maximal proliferation weeks 2-6 (callus tissue rapidly expanding filling gaps between scion and rootstock), serving as initial bridge (callus providing first physical connection enabling some water and solute movement), providing substrate for differentiation (callus cells serving as precursors for vascular tissue and cambium formation), and progressive replacement (successful unions showing callus gradually replaced by differentiated tissues over months to years as secondary growth proceeds). Callus graft union zone cellular characteristics includes= thin primary cell walls (cellulose-pectin walls lacking secondary thickening enabling cell division and expansion), large central vacuoles (occupying 60-80% of cell volume in mature callus cells), dense cytoplasm initially (young callus cells showing high metabolic activity with numerous organelles), minimal intercellular spaces (callus cells tightly packed maximizing tissue continuity), random orientation (cells lacking organized files or patterns seen in cambial derivatives), and totipotency (callus cells retaining capacity to differentiate into various cell types given appropriate signals). Callus graft union zone vascular differentiation within includes= cambium formation (localized regions of callus differentiating into cambial initials typically 3-6 weeks post-grafting), xylem element formation (vessel elements differentiating from cambial cells or directly from callus providing initial water transport pathways), phloem element formation (sieve tube elements and companion cells differentiating enabling photosynthate transport), progressive organization (initially scattered vascular elements gradually becoming organized into continuous tissues with continued secondary growth), and integration with partners (newly formed vascular tissues connecting with scion and rootstock vascular systems establishing functional continuity). Callus graft union zone transport function includes= limited initial transport (callus tissue alone providing minimal conducting capacity, water and solutes moving slowly through apoplast and symplast), improving with differentiation (formation of vascular elements within callus progressively increasing transport capacity), tortuous pathways (water and photosynthate following indirect routes through callus before organized tissues form), hydraulic resistance (callus-dominated union creating 40-80% higher resistance than equivalent healed union or non-grafted trunk length), and functional limitation (persistent callus-rich unions showing reduced transport capacity 50-75% of healed union potentially limiting vine vigor and productivity). Callus graft union zone mechanical properties includes= initially weak (callus tissue providing limited mechanical strength, young unions requiring support), soft texture (callus parenchyma easily compressed or damaged compared to woody tissues), vulnerable to separation (newly formed callus bridge potentially rupturing under stress if not properly supported), gradually strengthening (secondary tissue formation within callus improving mechanical integrity), but persistently weaker (unions dominated by callus showing 40-70% of healed union strength potentially vulnerable to breakage under load). Callus graft union zone in successful versus problematic unions includes= successful unions (callus prominent first 6-12 months then progressively replaced by organized vascular tissues, callus comprising 35°C potentially inhibiting formation or causing cell death), moisture (adequate humidity supporting callus development, desiccation killing callus cells, excessive moisture promoting pathogen growth), oxygen (aerobic conditions necessary for callus respiration and cell division), light (darkness or low light typical for union zone, bright light potentially affecting callus metabolism), and mechanical stability (immobilization improving callus bridge formation, movement disrupting integration). Callus graft union zone hormonal regulation includes= auxin promoting proliferation (indole-3-acetic acid stimulating callus cell division and expansion), cytokinin enhancing division (synergistic effect with auxin promoting rapid callus formation), auxin, cytokinin ratio determining differentiation (high auxin favoring xylem formation, balanced ratio promoting cambium, high cytokinin favoring shoot formation), gibberellin effects (potentially modulating cell expansion and differentiation), and ethylene potentially inhibitory (excessive ethylene from stress potentially suppressing callus formation or promoting senescence). Callus graft union zone pathogen vulnerability includes= susceptible to infection (soft callus tissue providing favorable substrate for fungal and bacterial colonization), entry point for pathogens (exposed callus potentially allowing trunk disease organisms to invade), importance of protection (proper grafting technique, wound protection, avoiding contamination minimizing infection risk), and healing importance (rapid differentiation of callus into protective bark and organized tissues reducing vulnerability window). Callus graft union zone significance includes= essential for union formation (callus providing initial bridge enabling subsequent vascular differentiation and integration), indicating healing stage (callus prominence reflecting union maturity, declining with successful healing), suggesting compatibility status (persistent excessive callus potentially indicating incompatibility problems), affecting union function (callus-dominated unions showing reduced transport and strength compared to differentiated unions), and determining management (young callus-rich unions requiring support and protection, persistent callus suggesting need for evaluation and potential remedial action).
Callus_Graft_Union_Zone emphasizes callus tissue dominance and is distinguished from Graft_Union_Zone (general transitional tissue), Healed_Graft_Union_Zone (organized differentiated tissues), and Callus_Tissue (callus in general) by focus on callus-rich union bridge. Critical for understanding early union healing stages, callus as substrate for differentiation, callus-dominated union functional limitations, healing progression assessment, and callus persistence as potential incompatibility indicator requiring monitoring and evaluation in graft union development.
Callus-dominated union
Healing tissue zone
Kallus-Verwachsungszone
Parenchym-Brücke
Parenchyma bridge
Wundheilungs-Gewebe
Callus Graft Union Zone
Cabernet Sauvignon callus tissue at 4 weeks post-grafting shows cream-colored soft parenchymatous tissue 8-15 mm thickness covering graft interface, cells 40-70 μm diameter with thin primary walls, dense cytoplasm, large nuclei 10-14 μm, cells dividing actively (visible mitotic figures in 5-12% of cells indicating cell cycle duration 2-3 days), callus from scion and rootstock intermixing forming continuous bridge, localized regions showing early differentiation with elongated cambium-like cells and scattered vessel element precursors, callus tissue providing substrate for progressive vascular differentiation enabling functional transport pathways within subsequent 2-4 weeks
Callus tissue is undifferentiated and partially differentiated parenchyma tissue that proliferates from cambial regions and adjacent mature tissues in response to wounding, grafting, or other stimuli, comprising thin-walled actively dividing cells that serve as precursors for vascular tissue differentiation, wound healing, and graft union formation in Vitis vinifera. Callus tissue cellular characteristics includes= parenchyma cells (thin-walled isodiametric to slightly elongated cells 30-80 μm diameter lacking specialized structure), dense cytoplasm initially (young actively dividing callus cells showing high organelle density, numerous mitochondria, active metabolism), large central vacuoles in mature cells (occupying 60-80% of cell volume in non-dividing callus), prominent nuclei (nuclei 8-15 μm diameter reflecting active transcription and cell cycle progression), thin primary cell walls (cellulose-pectin walls enabling cell division, expansion, and differentiation without lignified barriers), and minimal intercellular spaces (cells tightly packed maximizing tissue continuity and potential for subsequent organization). Callus tissue formation triggers includes= wounding (mechanical injury triggering dedifferentiation of adjacent cells and proliferation), grafting (cut surfaces stimulating callus production bridging scion and rootstock), hormonal signals (elevated auxin and cytokinin levels promoting cell division and callus formation), cambial activity (vascular cambium and immature cambial derivatives most active in callus production), and tissue stress (various stimuli including pathogen infection, freeze damage, or other stresses potentially inducing callus-like proliferation). Callus tissue origin and dedifferentiation includes= arising from vascular cambium (cambial cells and recent derivatives most readily producing callus), arising from immature xylem and phloem (young incompletely differentiated vascular elements reversing differentiation becoming callus), arising from mature parenchyma (xylem parenchyma, phloem parenchyma, cortex cells potentially dedifferentiating contributing to callus especially in woody tissues), cell division activation (quiescent mature cells re-entering cell cycle initiating division), and loss of specialization (differentiated cells losing specialized structures, functions becoming generic parenchyma). Callus tissue growth and proliferation includes= rapid cell division (callus cells dividing every 1-4 days during active formation much faster than organized tissues), cell expansion (daughter cells enlarging 2-4 fold after division), continued proliferation (callus tissue potentially continuing division for weeks to months given adequate conditions), three-dimensional growth (callus expanding in all directions filling gaps, creating tissue masses), and regulation by hormones (auxin and cytokinin balance controlling proliferation rate, continued growth versus differentiation). Callus tissue functions in graft union includes= bridging scion and rootstock (callus from both partners meeting forming continuous tissue connection across junction), providing substrate for differentiation (callus cells serving as precursors for vascular cambium, xylem, phloem formation), enabling initial transport (callus providing some water and solute movement through apoplast and symplast before organized vascular tissues form), facilitating healing (callus filling gaps, excluding pathogens, creating stable structure enabling subsequent tissue organization), and determining union quality (callus formation adequacy, organization, differentiation determining union success or failure). Callus tissue differentiation potential includes= totipotency (callus cells retaining capacity to differentiate into various cell types given appropriate signals), vascular differentiation (callus forming cambium, xylem vessels, phloem sieve tubes under appropriate auxin, cytokinin ratios and conditions), organogenic potential (callus potentially forming shoot or root primordia under specific hormonal conditions though less common in graft unions), and reversion (callus cells potentially reverting to more differentiated states or maintaining undifferentiated state depending on signals and conditions). Callus tissue hormonal regulation includes= auxin requirement (indole-3-acetic acid necessary for callus cell division and growth), cytokinin synergy (cytokinins working with auxin promoting callus proliferation), auxin, cytokinin ratio effects (high auxin favoring root-like differentiation, balanced ratio promoting continued callus or cambium formation, high cytokinin favoring shoot-like development), gibberellin effects (potentially modulating cell expansion and differentiation patterns), and ethylene influences (excessive ethylene potentially inhibiting callus formation or promoting senescence). Callus tissue in wound healing includes= initial response to injury (callus forming within 1-2 weeks at wound sites covering exposed tissues), protecting wound surface (callus providing temporary covering excluding pathogens, preventing desiccation), filling cavities (callus tissue expanding filling pruning cuts, injury sites), differentiating into permanent tissues (callus eventually forming vascular tissues, protective periderm completing healing), and serving as healing substrate (callus providing material from which organized repair tissues arise). Callus tissue types and characteristics includes= compact callus (densely packed cells with minimal intercellular spaces typical of graft unions), friable callus (loosely organized easily fragmented tissue sometimes forming in tissue culture or certain wound conditions), organized callus (showing some tissue organization, vascular strand formation, partial differentiation), disorganized callus (lacking structure, random cell orientation, minimal differentiation potential), and necrotic callus (dead or dying callus showing brown discoloration from oxidized phenolics, cell death, common in incompatible grafts). Callus tissue in successful versus unsuccessful unions includes= successful unions (adequate callus formation bridging junction, progressive differentiation into organized vascular tissues, callus proportion declining over time as replaced by functional tissues), unsuccessful unions (inadequate callus formation leaving gaps, excessive persistent callus without differentiation, callus showing necrosis or disorganization preventing proper integration), and integration quality (callus formation quantity and quality determining union healing trajectory and ultimate success). Callus tissue assessment and observation includes= visual observation (white to cream colored soft tissue appearing at wounds or graft junctions), tactile assessment (soft compressible texture contrasting with firm woody tissues), microscopic examination (thin sections revealing parenchymatous cells, degree of organization, differentiation status), vital staining (dyes accumulating in living callus cells indicating viability), and growth monitoring (tracking callus expansion and differentiation over days to weeks). Callus tissue environmental and management influences includes= temperature effects (optimal 20-28°C promoting rapid callus formation, 35°C inhibiting formation), moisture requirements (high humidity supporting callus development, desiccation killing callus cells), oxygen availability (aerobic conditions necessary for callus respiration and growth), light typically not required (callus forming in darkness or low light typical of graft union or wound interior), and protection importance (maintaining favorable conditions around callus critical for successful healing or union formation). Callus tissue pathogen vulnerability includes= susceptibility to infection (soft callus tissue providing favorable substrate for fungal and bacterial growth), importance of aseptic technique (proper grafting, wound care minimizing pathogen introduction), rapid differentiation benefits (callus quickly forming protective periderm and organized tissues reducing vulnerability period), and contamination consequences (pathogen invasion of callus potentially preventing healing or causing union failure). Callus tissue significance includes= essential for wound healing (callus providing initial response covering injuries, substrate for permanent tissue formation), critical for graft union (callus forming bridge enabling scion-rootstock integration), indicating tissue viability and response (callus proliferation demonstrating living tissue capable of wound response), determining healing outcomes (callus formation quality affecting wound closure or union success), and reflecting plant vigor (vigorous healthy plants typically showing robust callus response, stressed or weak plants showing poor callus formation potentially affecting healing capacity).
Callus_Tissue emphasizes undifferentiated parenchyma and is distinguished from specific callus contexts (Callus_Graft_Union_Zone, Healing_Callus_Tissue, Bridge_Callus_Tissue) by focus on general callus characteristics. Critical for understanding wound response mechanisms, tissue dedifferentiation and proliferation, graft union formation substrate, differentiation potential, hormonal regulation, and callus as essential healing tissue determining wound closure, graft success, and plant capacity for injury response and tissue regeneration.
Kallusgewebe
Repair tissue
Undifferentiated parenchyma
Undifferenziertes Parenchym
Wound healing tissue
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Pina, A., & Errea, P. (2005). A review of new advances in mechanism of graft compatibility-incompatibility. Scientia Horticulturae, 106(1), 1-11.
Callus Tissue
Chardonnay flowers at BBCH 59 (flower buds swollen, calyptras green and tightly sealed) exhibit intact calyptras 2.8-3.2 mm length covering developing stamens and pistil, with calyptra shedding beginning at BBCH 60 when morning temperatures reach 18-20°C, proceeding over 8-12 days across the inflorescence to achieve full bloom (BBCH 65) with 50-70% of calyptras shed and exposing functional hermaphrodite flowers. Gewürztraminer at BBCH 60-65 shows calyptra shedding progression with basal flowers shedding calyptras 2-3 days before apical flowers, creating acropetal bloom pattern with freshly detached calyptras visible as yellowish-green caps fallen onto leaves and ground below inflorescences. Pinot Noir under cool spring conditions (12-15°C) exhibits delayed calyptra shedding with some flowers retaining calyptras 3-5 days beyond normal shedding time, resulting in irregular bloom (BBCH 61-63 extended period) and reduced fruit set (35-42% versus normal 48-55%) due to impaired pollination from delayed stamen and stigma exposure.
A calyptra is the fused petal structure characteristic of Vitis flowers comprising five petals united at their apical margins to form a continuous cap or hood that encloses the developing stamens and pistil during pre-anthesis stages and is shed as a single coherent unit at anthesis through basal abscission, representing a distinctive floral morphology that distinguishes Vitaceae from most other angiosperm families where petals separate individually during flower opening. The calyptra typically measures 2-4 mm in length and 1.5-3 mm in diameter in the intact pre-anthesis state, appearing as a green to yellowish-green ovoid or conical structure covering the flower bud, with five longitudinal ridges or ribs corresponding to the five individual petal components visible on the external surface. The calyptra structure comprises= (1) five distinct petal units fused along their lateral margins throughout most of their length but united most strongly at their apical tips, creating a cap-like structure (2) a basal attachment zone where petal bases insert on the receptacle rim, forming a continuous circular abscission zone that separates during anthesis (3) internal surface in contact with developing stamens and pistil, often showing imprints or impressions from enclosed organs (4) external surface bearing a cuticle and sometimes trichomes depending on genotype, providing protection to enclosed reproductive organs. The calyptra serves multiple critical functions during flower development= (1) physical protection of developing stamens and pistil from environmental stresses (desiccation, mechanical damage, pathogen entry) during the vulnerable pre-anthesis developmental period (2) maintenance of appropriate microclimate (humidity, temperature) within the flower bud supporting proper organ maturation (3) prevention of premature pollen release before stigma receptivity is achieved, ensuring developmental synchrony between male and female functions (4) possible chemical protection through cuticle barrier preventing pathogen penetration. The calyptra shedding process (calyptra dehiscence or calyptra abscission) represents the defining event of grapevine anthesis, with shedding triggered by= (1) elongation of stamen filaments that push upward against the calyptra apex creating mechanical pressure (2) formation of an abscission layer at the calyptra base through programmed cell wall dissolution and cell separation (3) environmental triggers particularly temperature and light that activate the biochemical cascades controlling abscission layer formation and filament elongation. Calyptra shedding typically occurs during morning hours (6-10 AM) when temperature rises above threshold levels (typically 15-20°C depending on cultivar) and proceeds rapidly once initiated, with the entire shedding process for an individual flower typically completing within 1-3 hours. The calyptra is lifted off by the elongating stamens and detaches at the basal abscission zone, falling away from the flower or being carried off by wind, leaving the exposed stamens and pistil to perform their reproductive functions. The timing of calyptra shedding shows progression patterns within inflorescences, typically proceeding acropetally (from base to apex) over 7-14 days for complete inflorescence bloom, with cultivar-specific variation in bloom duration and synchrony. Calyptra morphology and shedding dynamics vary somewhat among flower types= hermaphrodite flowers typically show robust calyptras with efficient shedding driven by vigorous stamen elongation functionally female flowers may show slightly delayed or more variable shedding due to reduced stamen vigor staminate and pistillate flowers in dioecious types show calyptras comparable to hermaphrodite flowers as the calyptra structure itself is not sex-differentiated. Failed or delayed calyptra shedding (calyptra retention) can occur under unfavorable environmental conditions (cool temperatures 90%, nutritional stress) and impairs pollination and fruit set by preventing pollen release and stigma exposure.
Calyptra represents the distinctive fused petal structure unique to Vitaceae and is the defining morphological feature of grapevine flower anthesis. The calyptra must be distinguished from individual petals (which separate individually in most angiosperms), from the calyx (the sepal whorl that remains at the receptacle base), and from the corolla (the general term for petal whorl, here specifically fused as calyptra). Calyptra shedding serves as the primary phenological marker for bloom timing and is the basis for BBCH growth stage codes 60-69 (flowering stages). Sub-classification into Intact, Separating, and Detached states reflects the critical developmental progression from pre-anthesis through anthesis to post-shedding stages, with each state having distinct functional and observational significance for viticulture, phenology monitoring, and reproductive biology studies.
Blütenhaube
Kelchhaube
Kronblattkappe
corolla cap
petal cap
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 98-102 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 92-95 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development. In= Roubelakis-Angelakis, K.A. (ed.) Molecular Biology & Biotechnology of the Grapevine, pp. 190-194 Lebon, G. et al. (2008). Sugars and flowering in the grapevine. Journal of Experimental Botany 59=2565-2578.
Calyptra
In Cabernet Sauvignon vines in the Médoc, Bordeaux, cambial activity decline progresses through late October and November with cambial zone width decreasing from 10-12 cell layers in early October to 4-5 cell layers by late November, periclinal division frequency declining by approximately 80% over this period, and complete cambial dormancy achieved by early December when cross-sections show the characteristic narrow dormant cambium configuration.
Cambial activity decline in Vitis vinifera represents the progressive reduction and eventual cessation of vascular cambium function during autumn, terminating the secondary growth that added new xylem and phloem tissues to stems and roots throughout the growing season. This process occurs during late autumn, typically from October through November in Northern Hemisphere viticulture, as the cambial meristem responds to the same photoperiod and temperature signals that drive overall dormancy induction by progressively reducing cell division rate until complete quiescence is achieved. The vascular cambium, a cylindrical meristem located between the secondary xylem and secondary phloem, produces xylem cells toward the interior and phloem cells toward the exterior through periclinal cell divisions, with the rate of cell production varying seasonally from maximum activity during spring and early summer to progressively reduced activity during late summer and autumn. During cambial activity decline, the rate of periclinal divisions in cambial initials progressively decreases, the width of the cambial zone narrows as fewer derivative cells are maintained in the division-competent state, and ultimately all cell division ceases leaving the cambium in a dormant condition with minimal cellular layers. The regulation of cambial activity decline involves declining auxin flux from shoots as shoot growth terminates, reduced cytokinin transport from roots as root activity decreases, and increased abscisic acid levels that directly inhibit cambial cell division, collectively creating the hormonal environment that favors cambial dormancy. Anatomically, the transition from active to dormant cambium is visible in cross-sections as a narrowing of the cambial zone from 8-12 cell layers during active growth to 3-5 cell layers in the dormant state, along with changes in cell morphology as cambial derivatives complete differentiation rather than continuing division. The timing of cambial activity decline relative to the onset of freezing temperatures affects both xylem vessel development and cold hardiness, as actively dividing cambial cells are highly susceptible to freezing injury while dormant cambium can survive temperatures that would kill active tissue. The cessation of cambial activity marks the end of the annual increment of secondary growth, with the final cells produced forming the outer boundary of that year's growth ring visible in cross-sections of woody stems.
Cambial_Activity_Decline represents the secondary growth component of Growth_Cessation, specifically addressing vascular cambium function termination. Distinguished from Shoot_Growth_Termination which addresses primary shoot growth, and from Metabolic_Downregulation which addresses overall metabolic reduction. Involves reduction in cambial zone width and cessation of periclinal divisions. Duration typically spans 4-6 weeks. Complete cessation necessary for proper cold hardiness of stem tissues. Marks end of annual secondary growth increment.
Kambium-Ruheeintritt
Kambiumaktivitäts-Rückgang
Sekundärwachstums-Ende
cambial dormancy onset
cambium cessation
secondary growth termination
vascular cambium decline
vaskuläre Kambium-Einstellung
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 1 Grapevine Structure and Function Esau K (1948) Phloem structure in the grapevine and its seasonal changes. Hilgardia 18: 217-296 Mullins MG, Bouquet A, Williams LE (1992) Biology of the Grapevine Cambridge University Press
Cambial Activity Decline
Cambium browning quality assessment in Niagara Peninsula Vidal following an extreme cold event (-33°C with 3-day duration) evaluated the extent of permanent structural damage requiring vine rehabilitation or replacement. The scratch test technique, removing a small bark section and evaluating cambium color, was applied systematically at trunk base, mid-trunk, cordon origin, and mid-cordon positions on 50 vines per block. Trunk-base cambium browning quality showed 22% of vines with brown, dead cambium at the critical crown region, indicating complete girdling that would prevent vascular function and require trunk replacement from suckers or new plantings. Mid-trunk cambium browning quality showed only 8% damage, indicating that many trunk-base injuries were localized to the soil-line zone where temperature dynamics differ from air temperatures. Cordon cambium browning quality showed 35% with partial to complete browning, indicating widespread structural damage that would require cordon replacement through retraining from trunk shoots even where trunk survival permitted vine retention. The cambium browning quality gradient from trunk base (most damaged) through aerial positions informed recovery strategies, with trunk-girdled vines marked for retraining from below-damage suckers, cordon-damaged vines scheduled for retraining to replacement cordons, and undamaged vines managed for normal production. The assessment established that cambium browning quality evaluation was essential for recovery planning beyond the immediate bud damage assessment that addressed only current-season crop potential.
Cambium browning quality is a cold damage symptom quality that inheres in grapevine woody tissues and represents the extent of oxidative discoloration visible in the cambial zone of canes, cordons, and trunks following freezing injury that has killed the meristematic cambium layer responsible for secondary growth, with cambium browning indicating potential loss of the regenerative capacity required for wound healing, diameter growth, and the replacement of damaged vascular tissues. This quality addresses damage to the growth-generating tissue layer that is essential for long-term vine structural development, recognizing that while cambium damage may not immediately affect current-season function if vascular tissues remain intact, it compromises the vine's capacity for repair and growth that determines long-term health and productivity. The ontological significance of cambium browning quality lies in its indication of structural damage severity beyond the bud losses that primarily affect current-season crop, with extensive cambium damage potentially requiring vine retraining or replacement depending on the extent and location of injury, and with localized cambium damage creating wound sites vulnerable to disease entry that may have long-term health consequences. The biological basis of cambium browning encompasses the death of the thin meristematic layer between xylem and phloem that contains the actively dividing cells producing new vascular tissues and bark, with the oxidation of killed cambial cells producing visible browning detectable through bark removal or cross-sectioning that exposes the cambial zone. The "scratch test" or bark scraping technique provides the standard field diagnostic for cambium viability, with green coloration indicating living cambium capable of continued function and brown coloration indicating cambium death that interrupts secondary growth at the damaged location. The distribution of cambium damage often shows characteristic patterns with the trunk-soil interface particularly vulnerable due to temperature dynamics at the snow-soil boundary and the potential for ice encasement, while aerial portions may show differential damage reflecting cold air drainage patterns and exposure differences.
Cambium_Browning_Quality is_a Cold_Damage_Symptom_Quality, representing freeze injury to the meristematic cambium layer critical for secondary growth and wound repair. This class addresses structural damage beyond bud/crop impacts. The scratch test (bark scraping to expose cambium) provides standard field diagnostic. Trunk-base (crown) region shows particular vulnerability to freeze damage. Complete girdling requires vine retraining or replacement. Partial damage creates disease entry points with long-term health consequences. Distinguished from Bud_Browning_Quality by focus on structural tissues rather than crop potential.
Kambium-Frosttötung
Kambiumschaden
Kambiumverbräunung
Meristemverbräunung
cambial injury
cambium damage
cambium freeze kill
meristem browning
Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection. Michigan State University Extension Bulletin E2930 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 508-545 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235
Cambium Browning Quality
Assessment of Cambium_Hardiness_Disposition at the Finger Lakes Grape Program examined cambium survival patterns and regeneration capacity across 12 cultivars following natural and controlled cold exposure. Controlled freezing of cane sections to temperatures from -25°C to -40°C was followed by histological examination of the cambial zone, scoring cambium viability on a 0-5 scale (0 = fully viable, 5 = completely dead) based on cell browning and structural integrity. Cambium_Hardiness_Disposition ranged from LT50 = -25°C ('Cabernet Franc') to LT50 = -38°C ('Frontenac'), with cambium hardiness intermediate between phloem and xylem within each cultivar. Histological detail revealed that cambium death initiated in localized patches at temperatures 2-3°C above LT50, progressing to coalescent and then circumferential death at and below LT50. Recovery experiments tracked cambial regeneration following partial damage. Canes with 30-50% cambium kill (score 2) showed successful regeneration from surviving sectors, producing continuous cambial ring within 6 weeks of spring growth initiation. Canes with 70-90% cambium kill (score 4) showed incomplete regeneration with persistent gaps, resulting in sectored growth and reduced cane diameter increment. Complete cambium death (score 5) eliminated secondary growth capacity entirely. These findings demonstrate that Cambium_Hardiness_Disposition determines not only immediate tissue survival but long-term regenerative capacity, with partial damage recoverable through cambial proliferation while complete circumferential loss is permanent.
A Cambium_Hardiness_Disposition is a Cane_Cold_Hardiness_Disposition that inheres specifically in the vascular cambium of dormant grapevine canes and is realized through the freezing tolerance of this meristematic tissue layer, determining the capacity for spring resumption of secondary growth and regeneration of functional vascular tissues. This disposition represents the cold hardiness of the single-cell-layer lateral meristem positioned between phloem and xylem, with cambium survival essential for producing new xylem and phloem to replace winter-damaged or aging vascular tissues and for wound healing following pruning cuts or mechanical injury. The vascular cambium, composed of fusiform initials (producing axial vascular elements) and ray initials (producing vascular rays), is a thin but critically important tissue whose loss eliminates the vine's capacity for radial growth and vascular regeneration. Cambium cells survive freezing through mechanisms similar to phloem (freeze tolerance with controlled extracellular ice formation), with hardiness typically intermediate between phloem and xylem. Cambium damage manifests as a dark brown or black line at the phloem-xylem interface when canes are cross-sectioned, and can occur as isolated patches or complete circumferential death depending on cold severity and cultivar hardiness. Partial cambium kill allows recovery through proliferation of surviving cambial sectors, while complete circumferential cambium death results in permanent loss of growth capacity in affected tissues. Cambium_Hardiness_Disposition is particularly important for vine longevity and recovery from severe winter damage, as cambial regeneration underlies all wound healing and structural repair processes.
Cambium_Hardiness_Disposition is_a Cane_Cold_Hardiness_Disposition. The vascular cambium is the lateral meristem responsible for secondary growth, positioned as a single-cell layer between phloem and xylem. Cambium hardiness is typically intermediate between the more sensitive phloem and hardier xylem. Cambium damage appears as browning at the phloem-xylem interface. Partial cambium damage can recover through proliferation from surviving sectors, but complete circumferential death eliminates growth capacity permanently. Cambium survival is critical for long-term vine recovery and wound healing.
Kambium-Härte-Disposition
Kambiumfrosthärte
Meristem-Kältetoleranz
cambium freezing tolerance
meristem cold hardiness
vascular cambium winter resistance
Ashworth EN, Wisniewski ME (1991) Response of fruit tree tissues to freezing temperatures, HortScience 26:501-504 Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235 Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection, Michigan State University Extension Bulletin E-2930
Cambium Hardiness Disposition
A cane on Vitis vinifera cv. Pinot Noir in a vertical shoot positioning system, retained during winter pruning at 12 nodes length from a bilateral cordon, measuring 11mm diameter at base tapering to 7mm at tip, exhibiting brown lignified bark with clearly visible nodes spaced 8-12cm apart, each node bearing a compound bud containing primary, secondary, and tertiary positions, this cane expected to produce 10-12 fruiting shoots in spring bearing an estimated 18-24 clusters totaling 4.5-6kg fruit depending on cluster thinning decisions and fruit set success.
A mature, lignified shoot of Vitis vinifera from the previous growing season, typically 8-15mm diameter and 1-3 meters in length, characterized by brown woody appearance, nodes bearing dormant compound buds at regular intervals, internodal regions with varying degrees of lignification, and retention during winter pruning to serve as fruiting wood or renewal wood for the coming season. Canes represent the fundamental structural unit for crop production in cane-pruned training systems, with bud positions along canes providing the source of fruiting shoots, while in spur-pruned systems canes are shortened to 2-3 bud spurs serving similar productive function in more compact architecture, making cane selection, retention number, and positioning critical management decisions affecting yield, fruit quality, and long-term vine balance.
Canes are distinguished from shoots (current season green growth), arms and cordons (multi-year permanent wood), and renewal canes (specifically retained for next season's fruiting wood). Cane selection criteria include appropriate diameter (8-12mm ideal, avoiding overly vigorous >15mm or weak <6mm), adequate length for desired bud count, good lignification without winter damage, and appropriate positioning for trellis integration. The choice between cane pruning and spur pruning systems depends on cultivar characteristics, climate, labor availability, and quality objectives, with cane pruning generally providing more precise crop load control but requiring more skilled labor.
Fruchtrute
Rute
einjähriges Holz
fruiting cane
mature shoot
one-year wood
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 89-112.
Cane
Systematic characterization of Cane_Cold_Hardiness_Disposition at Michigan State University employed controlled freezing of dormant cane sections with subsequent assessment of tissue-specific damage across 20 cultivars. Cane segments were frozen to temperatures from -20°C to -40°C in 2.5°C increments, held for 4 hours, then slowly thawed and evaluated for browning in phloem, cambium, xylem rays, and pith at 48 hours post-treatment. Additionally, electrical conductivity of freeze-thaw leachate provided integrated damage assessment. Cane_Cold_Hardiness_Disposition showed substantial cultivar variation, with V. vinifera 'Riesling' exhibiting phloem LT50 = -26°C, cambium LT50 = -27°C, and xylem LT50 = -29°C, while the cold-hardy hybrid 'Frontenac' achieved phloem LT50 = -35°C, cambium LT50 = -36°C, and xylem LT50 = -38°C. The differential between primary bud and cane phloem LT50 (the "bud-cane gap") ranged from 3°C ('La Crescent') to 8°C ('Marquette'), with larger gaps indicating greater likelihood that buds would be killed while canes remain functional. Field validation following the 2014 polar vortex (minimum -34°C) confirmed that cultivars with cane phloem LT50 above -34°C showed vascular dysfunction symptoms including delayed and weak budbreak, shoot dieback, and reduced vigor even from surviving buds, demonstrating that Cane_Cold_Hardiness_Disposition assessment is essential for predicting vine recovery capacity beyond simple bud survival.
A Cane_Cold_Hardiness_Disposition is a Midwinter_Hardiness_Disposition that inheres in the dormant canes (one-year-old wood) of a Vitis vinifera plant and is realized through the freezing tolerance of cane tissues including bark, phloem, cambium, xylem, and pith, determining the survival of shoot framework that supports bud positioning and vascular connectivity to the vine's permanent structure. This disposition represents the cold hardiness of the current season's mature shoots that will bear the following year's fruiting wood, with cane survival essential for maintaining trained vine architecture, providing vascular supply to retained buds, and avoiding the need for retraining from trunk or cordon tissues following winter damage. Cane tissues generally achieve greater cold hardiness than primary buds within the same cultivar, with the hardiness hierarchy typically being primary bud < secondary bud < tertiary bud < phloem < cambium ≤ xylem, creating a damage progression where buds are killed before cane function is compromised. However, severe cold events can damage cane tissues, disrupting vascular continuity even when some buds survive, and resulting in weak, poorly supplied shoots or complete failure of bud break despite apparent bud viability. Cane_Cold_Hardiness_Disposition encompasses multiple tissue types with distinct freezing behaviors, with phloem showing freeze tolerance through controlled extracellular ice formation, xylem exhibiting deep supercooling in ray parenchyma cells, and cambium requiring protection of the meristematic zone for spring growth resumption. Assessment of cane hardiness through controlled freezing with subsequent tissue browning evaluation or electrical conductivity measurements provides important complement to bud hardiness data for comprehensive winter damage prediction.
Cane_Cold_Hardiness_Disposition is_a Midwinter_Hardiness_Disposition. Cane hardiness encompasses multiple tissues with different freezing behaviors and damage thresholds, addressed through child classes Phloem_Hardiness_Disposition, Xylem_Hardiness_Disposition, and Cambium_Hardiness_Disposition. Cane damage can compromise shoot development even when buds survive, making cane hardiness assessment important for complete winter injury prediction. Canes are generally hardier than buds, so cane damage indicates severe cold events exceeding normal bud protection thresholds.
Einjähriges-Holz-Winterhärte
Rutenholz-Frosthärte
Trieb-Kältehärte-Disposition
cane winter hardiness
one-year wood freezing tolerance
shoot cold hardiness
Wolpert JA, Howell GS (1985) Cold acclimation of Concord grapevines II. Natural acclimation pattern and tissue moisture decline in canes and primary buds of bearing vines, American Journal of Enology and Viticulture 36:189-194 Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235 Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection, Michigan State University Extension Bulletin E-2930
Cane Cold Hardiness Disposition
The cane color description of Riesling documents pale yellowish-brown canes (OIV 103:2) — the lightest cane color among major wine grape cultivars, with the cork layer containing low concentrations of phenolic pigments that produce a straw-to-honey tone contrasting sharply with the dark reddish-brown canes of Cabernet Sauvignon (OIV 103:5) and the near-black canes of Alicante Bouschet (OIV 103:7). The pale cane color of Riesling is consistent across growing conditions and is one of the primary dormant-season identification features used by Mosel and Rheingau viticultural workers, who can distinguish Riesling from the darker-caned Müller-Thurgau (OIV 103:3-4) and the reddish Spätburgunder/Pinot Noir (OIV 103:4-5) at pruning distance in the vineyard.
A cane description component that records the bark color of the dormant cane according to OIV descriptor 103, scored on a scale from yellowish (state 1) through yellowish-brown, greyish-brown, reddish-brown, to dark brown-violet (state 7), documenting the pigmentation of the mature periderm that reflects the cultivar-specific composition and concentration of phenolic compounds, tannins, and melanin-like pigments deposited in the cork cell walls during the lignification process.
Cane_Color_Description documents the primary visual character for dormant-season cultivar identification — cane color is to winter identification what shoot tip color is to spring identification, providing the most immediately visible and diagnostically useful character when other organ-based descriptors are unavailable. The cane color results from the progressive deposition of phenolic compounds in the cell walls of the periderm (cork) cells during the lignification process from véraison through dormancy — these phenolic pigments include condensed tannins (proanthocyanidins), phenolic acids (hydroxycinnamic acids), and melanin-like oxidative polymerization products whose composition and concentration vary between cultivars: light-colored canes (Riesling, Chasselas, Sultana) contain lower concentrations of total phenolics and a higher proportion of unoxidized monomeric phenolics, while dark-colored canes (Cabernet Sauvignon, Mourvèdre, Alicante Bouschet) contain higher concentrations of polymeric oxidized tannins and melanin-like pigments. The genetic control of cane color is partially independent of berry and shoot anthocyanin production — white-berried cultivars can produce dark canes (e.g., Sémillon has moderately dark brown canes despite lacking berry anthocyanin) because cane pigmentation involves brown phenolic polymers rather than the red-purple anthocyanins that determine berry skin color.
Rutenfarbe-Beschreibung
Rutenrinden-Färbung
cane bark color record
dormant shoot pigmentation
lignified shoot color
verholzter Trieb-Pigmentierung
Cane Color Description
The cane cross-section of Cabernet Sauvignon documents a nearly circular profile (OIV 104:1) with the transverse and dorsiventral diameters differing by less than 10% — the round cross-section is maintained consistently from the mid-cane internodes through the basal internodes and is visible as a round cut surface when pruning cuts are made during winter. This circular profile contrasts with the distinctly elliptical cross-section of Sémillon (OIV 104:2, transverse diameter exceeding dorsiventral by 15-25%, creating a visibly flattened cane), the strongly flattened canes of some rootstock varieties (V. riparia selections showing dorsiventral compression of 30-40%), and the irregular cross-section of Mourvèdre (nominally round but with pronounced ridges creating a lobed outline). The cross-section profile is best observed at mid-cane positions (nodes 5-10) where the cane has achieved full diameter without the basal thickening that can modify the profile at the oldest internodes.
A cane description component that records the shape of the dormant cane in transverse cross-section, documenting the profile as circular, elliptical, or flattened (dorsiventrally compressed) according to OIV descriptor 104, with the cross-sectional shape reflecting the developmental geometry of cambial growth and pith development that produces cultivar-characteristic cane profiles visible when canes are cut during pruning operations.
Cane_Cross_Section documents a character that is observed directly during the most common viticultural operation — winter pruning — when every cut through a cane reveals its cross-sectional profile. The profile shape is determined by the geometry of secondary growth — the cambial cell division pattern that produces wood (xylem) centripetally and bark (phloem and periderm) centrifugally during the growing season — and by the shape and size of the central pith cavity: in cultivars with uniform radial cambial activity, the resulting cross-section is circular, while cultivars with differential cambial activity (stronger growth on the dorsal versus ventral face, or along the plane perpendicular to the leaf trace) produce elliptical or flattened profiles. The pith shape contributes independently — some cultivars have circular pith cavities centered in a circular cane (producing symmetrical round cross-sections), while others have eccentric or elliptical pith cavities that create asymmetric internal structure even within an externally round cane. The cross-section character is particularly useful for rootstock identification, where the strong interspecific variation in cane geometry (V. riparia producing markedly flattened canes, V. berlandieri producing round to slightly elliptical canes) aids the identification of rootstock varieties in established vineyards where above-ground cultivar identification of rootstocks is otherwise limited to the few visible suckers.
Ruten-Querschnittsform
Rutenquerschnitt
Triebquerschnitt-Geometrie
cane profile record
shoot cross-section shape
transverse cane geometry
Cane Cross Section
The cane description of Syrah documents medium-thick dormant canes (8-12 mm internode diameter) with dark brown bark showing a reddish-mahogany undertone (OIV 103:5), a smooth surface texture with fine longitudinal striations and minimal flaking, moderately prominent nodes swollen to approximately 120% of internode diameter, conspicuous dark brown diaphragms visible when the cane is split longitudinally, and medium-sized conical dormant buds with dense protective pubescence — this cane morphology is sufficiently distinctive that experienced Rhône Valley viticulturists can identify Syrah during winter pruning with high confidence, distinguishing it from the paler straw-brown canes of Viognier (OIV 103:2-3), the deeply furrowed dark canes of Mourvèdre (OIV 103:5-6 with rough bark texture), and the flexible, light brown canes of Grenache (OIV 103:3-4 with smooth texture and characteristic brittleness at the nodes).
An ampelographic description component that documents the morphological characteristics of the dormant cane — the mature, fully lignified shoot observed after leaf fall through the winter dormancy period — recording bark color, surface texture, cross-sectional profile, node prominence, bud morphology, and diaphragm structure that provide cultivar identification characters available exclusively during the 4-5 month dormant season when no leaf, flower, or fruit characters are observable.
Cane_Description addresses the identification challenge unique to the dormant season — the 4-5 month period from leaf fall (November-December in northern hemisphere vineyards) through budburst (March-April) during which the dormant cane is the only vegetative organ available for cultivar identification. This temporal niche makes cane characters practically important for viticultural operations that occur during dormancy — particularly pruning, where correct cultivar identification is essential in mixed-cultivar plantings, replacement plantings, and vineyard surveys, and propagation wood collection, where cane identification prevents the multiplication of incorrectly identified material: the practical importance of dormant-season identification has led to the development of specialized cane-based identification keys in all major viticultural regions, with experienced pruning crews developing remarkable proficiency in identifying cultivars by cane characteristics alone. The cane represents the final developmental stage of the shoot — the lignification process that converts green photosynthetic shoot tissue to brown woody cane tissue involves progressive periderm formation (replacing the green epidermis with protective cork cells), loss of chlorophyll and accumulation of brown phenolic pigments, and starch deposition in ray parenchyma cells (visible as white flecks when the cane is cross-sectioned) — these processes produce the cultivar-characteristic cane color, texture, and internal structure documented in the subclasses.
Ruten-Beschreibung
Winterruten-Charakterisierung
dormant cane characterization
lignified shoot morphology
mature shoot description
verholzter Trieb-Morphologie
Cane Description
Selection of fruiting canes for a head-trained, cane-pruned Riesling vineyard in the Mosel targets Cane_Diameter_Quality of 8-10 mm at the basal internode, rejecting both thin canes below 6 mm that indicate insufficient carbohydrate reserves for spring growth and bull canes exceeding 14 mm that typically show reduced bud fruitfulness at the 5-8 node positions critical for this training system's crop load distribution.
Cane_Diameter_Quality is a Vigor_Expression_Quality that inheres in a grapevine (Vitis vinifera L.) and represents the cross-sectional dimension of mature, lignified one-year-old shoots (canes) as measured during the dormant season, providing an indicator of individual shoot vigor and wood maturity independent of total vine biomass production. This quality captures the investment of resources into individual shoot structural development, reflecting both the growth vigor during shoot extension and the subsequent maturation processes that determine cane quality for various viticultural purposes including propagation material selection and fruiting cane retention. In BFO terms, Cane_Diameter_Quality exists as a two-dimensional spatial quality (cross-sectional area, typically approximated as circular diameter) measurable at standardized positions along the cane, most commonly at the basal internode or at a specified internode number. The physiological determinants of cane diameter include vascular cambium activity during the growing season, carbohydrate availability for secondary growth, and the duration of the growth period before dormancy onset, with water stress and excessive crop load both reducing diameter through competition for carbon resources. Optimal cane diameter for balanced vine management varies with cultivar and training system but typically falls between 7-12 mm at the basal internode, with smaller diameters indicating insufficient vigor or excessive crop load and larger diameters suggesting vegetative excess potentially at the expense of fruit quality. For cane-pruned systems where entire canes are retained as fruiting units, cane diameter uniformity and appropriate size directly influence bud fruitfulness and fruit set consistency along the retained cane, with excessively vigorous canes showing reduced fruitfulness at basal nodes and weak canes lacking the carbohydrate reserves to support strong early-season growth. Cane diameter also correlates with winter hardiness, as larger diameter canes with greater carbohydrate reserves typically demonstrate superior cold tolerance and more reliable bud viability following winter stress.
Cane_Diameter_Quality represents an individual shoot vigor measure complementing the whole-vine assessment provided by Pruning_Weight_Quality. This quality is particularly relevant for cane-pruned training systems where individual cane selection determines fruiting unit quality, and for propagation material assessment where cane diameter indicates cutting quality and establishment potential. The measurement standardization issue (which internode, caliper vs. digital measurement) affects comparability across studies and should be specified in experimental protocols.
Holzstärke
Rutendurchmesser
Triebdurchmesser
cane caliber
cane cross-section
cane thickness
shoot diameter
wood diameter
Wolf TK, Pool RM (1987) Factors affecting exotherm detection in the differential thermal analysis of grapevine dormant buds. Journal of the American Society for Horticultural Science 112:520-525 Howell GS, Shaulis N (1980) Factors influencing within-vine variation in the cold resistance of cane and primary bud tissues. American Journal of Enology and Viticulture 31:158-161
Cane Diameter Quality
Cane maturation progresses basipetally (from base to tip) during late summer and autumn, with fully matured canes showing brown periderm color, firm texture, and cold hardiness to -15 to -25°C depending on cultivar, while immature portions remain green, soft, and susceptible to freeze damage below -5°C. Maturation rate varies among cultivars, with early-maturing types such as Pinot Noir and Chardonnay completing lignification 3-4 weeks before late-maturing cultivars such as Carignan and Mourvèdre. Internode maturation percentage at first frost ranges from 95-100% in well-adapted cultivars to 60-80% in poorly matched cultivar-climate combinations or over-cropped vines. Carbohydrate reserve concentration in mature canes reaches 15-25% of dry weight (predominantly starch), providing energy for cold hardiness maintenance and spring growth resumption.
A longevity and resilience disposition that is the inherited capacity of a Vitis vinifera plant to complete the developmental transition of current-season shoots from green, herbaceous growth to lignified, cold-hardy canes capable of surviving winter dormancy and supporting following-season fruiting, encompassing periderm formation, reserve accumulation, and acquisition of freezing tolerance.
Cane_Maturation_Disposition serves as the parent class for periderm formation and reserve accumulation traits. Cane maturation is critical for vineyard survival in cold climates, as immature wood is highly susceptible to winter injury. The process integrates multiple developmental and physiological changes occurring in parallel, including secondary wall lignification, periderm development, starch accumulation, and cold acclimation gene expression. Viticultural practices (crop load management, shoot positioning, water management) significantly affect maturation rate and completeness.
Holzreife
Rutenreife
Triebausreifung
cane ripening
shoot maturation
wood maturation trait
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Keller M, Tarara JM (2010) Warm spring temperatures induce persistent season-long changes in shoot development in grapevines. Annals of Botany 106:131-141
Cane Maturation Disposition
A Pinot Noir vine in Burgundy's Côte d'Or trained to single guyot bears a Cane_Pruned_Vine_Role, with a single 8-bud cane arched and tied along the fruiting wire, plus a 2-bud renewal spur positioned to provide next year's cane—the annual cane replacement reflecting Pinot Noir's moderate basal bud fruitfulness. A Riesling vine in Rheingau guyot training serves in a Cane_Pruned_Vine_Role, the cane-based system suited to the variety's requirement for bud positions beyond the basal nodes for optimal fruitfulness. A Sauvignon Blanc vine on double guyot in Loire Valley bears a Cane_Pruned_Vine_Role with two canes extending in opposite directions, the cane-pruned approach standard for this variety's regional production.
A training system role that inheres in a grapevine managed through annual replacement of fruiting wood, where one-year-old canes (mature shoots from the previous season) are retained at pruning to bear the current season's crop, then removed entirely the following year and replaced with new canes selected from that season's growth. Cane pruning contrasts fundamentally with spur pruning, where permanent spur positions on cordons bear fruit year after year in cane systems, the entire fruiting structure is renewed annually, requiring selection of appropriately positioned, well-matured canes at each pruning. Cane pruning is preferred for varieties where basal buds (those closest to the cane origin) are less fruitful than buds further along the cane, as the retained cane length (typically 8-15 buds) positions fruiting buds at optimal cane positions. The guyot system and its variants (single guyot, double guyot) represent the dominant cane-pruning approaches, with the retained cane(s) tied horizontally along a fruiting wire while a short spur (2 buds) is retained to produce replacement canes for the following year. Cane pruning requires greater pruning skill than spur systems, as cane selection and positioning significantly influence vine balance and fruit quality.
Cane_Pruned_Vine_Role is subclassed by guyot variants (single and double) that specify cane number and arrangement. The choice between cane and spur pruning reflects variety physiology—varieties with fruitful basal buds (Cabernet Sauvignon, Merlot, Syrah) perform well under spur pruning, while those requiring mid-cane bud positions for fruitfulness (Pinot Noir, Riesling, Sauvignon Blanc, Sémillon) benefit from cane retention. Cane pruning demands pruning expertise the pruner must select well-positioned, adequately mature canes of appropriate diameter and internode length, then tie them without damage. The annual wood renewal in cane systems may reduce trunk disease accumulation compared to spur systems with permanent cordon wood, though this benefit remains debated. Cane-pruned systems are associated with premium production and traditional regions, though the labor requirements limit adoption in cost-sensitive contexts.
Bogrebenschnitt
Langschnitt
Streckerschnitt
Zapfenschnitt mit Strecker
cane pruning
cane-pruned vine
long pruning
replacement cane system
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles Galet P (2000) General Viticulture, Oenoplurimedia Winkler AJ et al. (1974) General Viticulture, University of California Press Jackson RS (2014) Wine Science: Principles and Applications, Academic Press
Cane Pruned Vine Role
The cane selection criteria for Riesling in the Rheingau specify retention of a cane with 7-10 mm diameter at the base (avoiding both thin whip canes below 6 mm that indicate poor vigor and thick bull canes above 12 mm that have wide internodes and potentially lower bud fruitfulness), regular internode spacing of 6-10 cm (indicating steady moderate growth throughout the previous season), fully mature brown periderm with no green or partially lignified sections, healthy buds with no visible mechanical or frost damage, and origin from the renewal spur zone within 5 cm of the head or cordon — canes meeting these criteria are bent to the fruiting wire and tied, while rejected canes are removed entirely, with the selection decision requiring experienced judgment honed over years of pruning practice.
A pruning method description that documents the qualitative and quantitative characteristics used to select the optimal fruiting cane during winter pruning in cane-pruned systems, specifying the assessment of cane diameter, internode length, bud health, wood maturity, position relative to the renewal spur, and overall cane vigor that determines which of the available one-year-old canes is retained as the fruiting unit for the upcoming growing season.
Cane_Selection_Criteria captures the qualitative assessment dimension of pruning that distinguishes skilled manual pruning from mechanical or rule-based approaches. The selection of the fruiting cane is the most consequential individual decision in cane-pruned systems because a single cane will produce 6-10 shoots bearing the vine's entire crop: selecting a cane of inappropriate diameter, maturity, or position can compromise an entire season's production from that vine. Research has established correlations between cane characteristics and subsequent performance — moderate-diameter canes (7-10 mm for most cultivars) with regular internodes produce shoots of more uniform fruitfulness and vigor than either thin or excessively thick canes, and well-positioned canes originating from the renewal spur zone maintain the vine's structural integrity over successive pruning cycles. The tacit knowledge embedded in expert cane selection is one of the most difficult viticultural skills to formalize and transfer, representing a significant challenge for the automation of precision pruning systems.
Bogenschnitt-Auswahlparameter
Fruchtrutenselektions-Standards
Rutenauswahl-Kriterien
cane evaluation parameters
cane quality assessment criteria
fruiting cane selection standards
Galet P (2000) General Viticulture, Oenoplurimedia pp: 255-275 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 340-360 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 100-115
Cane Selection Criteria
The cane surface description of Mourvèdre documents a distinctly rough, deeply furrowed bark surface with prominent longitudinal ridges and shallow bark flaking on mature cane sections — the bark develops a coarse, irregular texture from the second year onward, with raised lenticels clearly visible as pale dots against the dark brown background and longitudinal fissures 0.5-1.0 mm deep creating a rugose surface that is immediately distinguishable by touch from the smooth, glossy canes of Grenache (virtually no longitudinal furrowing, minimal lenticels, the bark surface appearing almost polished) and the moderately textured canes of Carignan (fine longitudinal striations without deep furrowing). The rough bark texture of Mourvèdre correlates with the thick periderm development characteristic of this cultivar, which also contributes to Mourvèdre's relatively high tolerance of trunk diseases by providing a more effective physical barrier against wood-colonizing pathogens.
A cane description component that records the textural characteristics of the dormant cane bark surface, documenting the degree of smoothness versus roughness, the presence and depth of longitudinal striations or ridges, the occurrence of bark flaking or exfoliation, lenticel prominence and density, and the overall surface regularity that provides cultivar-diagnostic tactile and visual characters complementing the cane color for dormant-season identification.
Cane_Surface_Description captures the tactile and microtopographic features of the bark surface that experienced viticulturists use alongside color for dormant-season identification — the combination of color and texture creates a multisensory identification impression (the experienced pruner simultaneously sees the color and feels the texture as the cane passes through the hand during pruning cuts) that is remarkably effective for cultivar discrimination. The bark surface texture is determined by the architecture of the periderm — the thickness and regularity of the cork cell layers, the frequency and prominence of lenticels (porous regions in the cork that permit gas exchange), and the occurrence of secondary periderm formation that produces bark flaking in some cultivars: smooth-barked cultivars (Grenache, Chasselas, Pinot Noir) develop thin, uniform periderm with inconspicuous lenticels and no secondary periderm, while rough-barked cultivars (Mourvèdre, Tannat, some rootstock species) develop thick, irregular periderm with prominent lenticels, deep longitudinal fissures formed by differential tangential growth of the bark layers, and occasional bark flaking from secondary periderm formation. The genetic control of bark texture has not been systematically mapped in Vitis, but the strong cultivar-specificity and environmental stability of the character suggest substantial genetic determination.
Rutenbark-Textur
Rutenoberflächen-Beschreibung
Rutenoberflächen-Morphologie
cane bark texture record
dormant cane texture characterization
shoot surface morphology
Cane Surface Description
Canopy-air temperature differential assessment in Central Valley Thompson Seedless vineyard at 14:00 shows values of -3.5°C in fully-irrigated rows and +2.1°C in deficit-irrigated rows, with calculation of CWSI values of 0.15 and 0.72 respectively using site-specific baseline equations, triggering irrigation of the deficit zone when CWSI exceeds the 0.5 threshold established for maintaining table grape berry size and appearance quality.
Canopy-air temperature differential quality is a temperature differential quality that represents the difference between grapevine canopy temperature and ambient air temperature measured at a standardized reference height, characterizing the net effect of energy balance processes determining whether the canopy is cooler than, warmer than, or in equilibrium with the surrounding atmosphere. Within the BFO framework, this quality specifically captures the relationship between vegetation thermal state and atmospheric conditions, providing a normalized indicator of plant physiological status that accounts for variation in ambient temperature across times, sites, and seasons. The ontological significance lies in this differential serving as the foundation for crop water stress indices, where the magnitude and sign of the canopy-air temperature difference reflects stomatal conductance and transpirational cooling capacity that directly indicate plant water status. Under well-watered conditions with open stomata, grapevine canopies typically maintain temperatures 2-6°C below air temperature due to latent heat dissipation through transpiration, while water-stressed vines with closed stomata lose cooling capacity and canopy temperature rises to or above air temperature. The crop water stress index (CWSI) normalizes the observed canopy-air differential against theoretical limits representing non-transpiring (upper limit, maximum stress) and fully-transpiring (lower limit, no stress) canopies, enabling comparison across varying atmospheric demand conditions. Measurement requires paired determination of canopy temperature (typically by infrared thermometry) and air temperature (by shaded, ventilated sensor at standard height), with timing standardized to periods of high atmospheric demand (typically early afternoon) when stress discrimination is greatest. This differential quality enables irrigation scheduling based on plant response rather than soil water depletion, identification of spatial variability in water stress for precision management, and evaluation of irrigation system performance.
Canopy_Air_Temperature_Differential_Quality provides the specific differential underlying crop water stress indices and thermal-based irrigation scheduling. This quality requires standardized measurement protocols including time of day, sensor height for air temperature, and canopy view angle for infrared measurement to ensure meaningful comparison across observations. The class distinguishes from Berry_Ambient_Temperature_Differential_Quality which addresses fruit rather than canopy thermal status.
Bestands-Luft-Temperaturdifferenz
CATD
Kronendach-Atmosphäre-Differenz
Laubwand-Luft-Temperaturdifferenz
Vegetations-Luft-Temperaturunterschied
canopy-air delta T
crop-air thermal differential
vegetation-atmosphere temperature difference
Idso SB et al. (1981) Normalizing the stress-degree-day parameter for environmental variability. Agricultural Meteorology 24: 45-55 Bellvert J et al. (2015) Vineyard irrigation scheduling based on airborne thermal imagery and water potential thresholds. Australian Journal of Grape and Wine Research 21(2): 195-204
Canopy Air Temperature Differential Quality
Canopy density quality comparison across three training systems in Finger Lakes Riesling demonstrated systematic differences in fruit zone light environment and disease pressure. Vertically shoot-positioned vines with bilateral cordons achieved moderate canopy density quality with leaf layer number 1.4 and 28% gaps at véraison, corresponding to Botrytis incidence of 8% at harvest. High-wire cordon trained vines exhibited excessive canopy density quality with leaf layer number 2.9 and only 6% gaps, suffering 34% Botrytis incidence despite equivalent fungicide programs. Geneva Double Curtain training distributing canopy across two curtains achieved lowest canopy density quality with leaf layer number 0.9 and 42% gaps per curtain, reducing Botrytis to 3% while maintaining equivalent yield through doubled canopy surface area.
Canopy density quality is a viticultural quality that inheres in a grapevine canopy system and represents the degree of compactness or openness of the foliar mass as determined by the spatial arrangement, layering, and overlap of leaves, shoots, and lateral growth within the canopy volume defined by the training system architecture. This quality fundamentally determines the light environment, air circulation patterns, and spray penetration characteristics within the canopy interior, thereby exerting pervasive influence on fruit composition, disease pressure, and management operation effectiveness. The ontological significance of canopy density quality lies in its role as an integrative expression of the interaction between vine vigor, training system geometry, and canopy management intensity that creates the microenvironmental conditions directly experienced by developing fruit clusters and interior leaves. Dense canopies arise from high shoot density, excessive shoot vigor producing long internodes and large leaves, abundant lateral development, and inadequate canopy management intervention, resulting in multiple overlapping leaf layers that shade interior tissues and restrict air movement. Conversely, open canopies with appropriate porosity result from balanced vigor, optimized shoot spacing, timely shoot positioning, and strategic leaf removal that maintains sufficient but not excessive leaf area in well-organized spatial arrangement. The measurement of canopy density quality employs various approaches including visual assessment using standardized scoring systems, point quadrat analysis quantifying leaf layer number and gap frequency, light interception measurements using ceptometers or quantum sensors, and image analysis techniques capturing canopy porosity from digital photographs. Optimal canopy density represents a cultivar-specific and climate-specific target wherein sufficient leaf area exists to ripen fruit and accumulate carbohydrate reserves while avoiding the negative consequences of excessive density including reduced photosynthetic efficiency of shaded interior leaves, elevated disease pressure from poor air circulation and extended wetness duration, herbaceous flavor development in shaded fruit, delayed and uneven ripening, and increased pesticide application requirements to achieve adequate coverage. The temporal dynamics of canopy density quality follow seasonal patterns of shoot growth, leaf expansion, and lateral development, with density typically increasing from budbreak through véraison before stabilizing or declining as late-season leaf senescence begins. Training system selection fundamentally influences achievable canopy density targets, with vertically shoot-positioned systems concentrating foliage in narrow planes that facilitate density management compared to sprawling systems where three-dimensional canopy expansion complicates density assessment and manipulation.
Canopy_Density_Quality serves as the parent class for Canopy_Porosity_Quality and Leaf_Layer_Number_Quality hierarchies, which represent complementary approaches to density characterization. This class addresses the overall density concept while child classes capture specific dimensions including light/air penetration and quantitative leaf layering. Strong functional relationships exist with Point_Quadrat_Index_Quality which provides standardized methodology for density assessment, and with Vigor_Quality classes that determine the growth characteristics creating density patterns.
Blattdichte
Kronendichte
Laubfülle
Laubwanddichte
canopy compactness
canopy fill
foliage density
leaf density
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 24-43 Dry PR (2000) Canopy management for fruitfulness. Australian Journal of Grape and Wine Research 6:109-115 Schultz HR (1995) Grape canopy structure, light microclimate and photosynthesis. I. A two-dimensional model of the spatial distribution of surface area densities and leaf ages in two canopy systems. Vitis 34:211-215
Canopy Density Quality
Canopy dimension quality comparison across training systems in a Mosel Riesling vineyard demonstrated systematic differences in light interception efficiency. Traditional single Guyot training produced canopy dimensions of 1.4 m height by 0.4 m width, yielding surface area of approximately 1.12 m² per meter of row and total light interception of 68% at 1.8 m row spacing. Conversion to vertical shoot positioning with bilateral cordons achieved dimensions of 1.2 m height by 0.5 m width with surface area of 1.7 m² per row meter through improved canopy fill, increasing light interception to 78%. Implementation of Scott Henry divided canopy created two curtains each 0.8 m height by 0.3 m width, achieving combined surface area of 2.2 m² per row meter and 85% light interception through increased exterior canopy surface despite equivalent total leaf area, demonstrating how canopy dimension quality manipulation through training system design can improve vineyard productivity without increasing vigor.
Canopy dimension quality is a viticultural quality that inheres in a grapevine canopy system and represents the spatial extent and geometric characteristics of the foliar mass as defined by the height, width, and derived surface area of the vegetative structure created through the interaction of training system architecture, vine vigor expression, and canopy management intensity. This quality fundamentally determines the light interception capacity of the vineyard, the ratio of exterior to interior canopy volume, and the compatibility of the canopy with mechanical operations including harvesting, hedging, and spray application that require specific dimensional tolerances for effective function. The ontological significance of canopy dimension quality lies in its role as the physical framework within which all other canopy characteristics manifest, wherein dimensions establish the boundaries containing leaf area, fruit clusters, and the microenvironmental gradients that influence grape quality development. Canopy dimensions result from the interplay between training system design specifying target architecture, vine vigor determining growth pressure toward dimensional expansion, and management interventions constraining growth within prescribed limits through shoot positioning, topping, and hedging operations. The three primary dimensional parameters of height, width, and depth interact to determine canopy volume, while the geometric arrangement of this volume relative to row spacing and vine spacing establishes the proportion of total vineyard area occupied by productive canopy versus the inter-row alleyways required for equipment passage and the inter-vine spaces determining canopy continuity along rows. Optimal canopy dimension quality represents a balance between maximizing light interception through adequate canopy extent and maintaining dimensional ratios that permit sufficient light penetration to interior positions where fruit develops. The measurement of canopy dimension quality employs direct physical measurement of height and width at standardized positions, photographic analysis capturing canopy profiles, and increasingly sophisticated approaches including LiDAR scanning that generates three-dimensional canopy models enabling precise volume and surface area computation. Training system selection fundamentally constrains achievable canopy dimensions, with vertical shoot positioned systems typically producing narrow, tall canopies while horizontally divided systems create wider, shorter profiles that distribute equivalent leaf area across greater horizontal extent to increase exterior surface area to volume ratios.
Canopy_Dimension_Quality serves as the parent class for specific dimensional parameters including Canopy_Height_Quality, Canopy_Width_Quality, and Canopy_Surface_Area_Quality. This class addresses the overall geometric framework of the canopy while child classes capture particular dimensional aspects. Distinguished from Canopy_Density_Quality which addresses foliage distribution within the dimensional framework rather than the framework dimensions themselves. Strong relationships exist with training system selection and mechanical operation compatibility.
Kronenausdehnung
Kronendimensionen
Laubwandgeometrie
Laubwandgröße
canopy extent
canopy geometry
canopy size
foliar dimensions
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 18-31 Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review. American Journal of Enology and Viticulture 60:251-268 Mabrouk H, Sinoquet H (1998) Indices of light microclimate and canopy structure of grapevines determined by 3D digitising and image analysis, and their relationship to grape quality. Australian Journal of Grape and Wine Research 4:2-13
Canopy Dimension Quality
A well-managed VSP Pinot Noir vine maintains nearly all leaf area in Canopy_Exterior_Role positions through shoot positioning and lateral removal, with the open canopy architecture ensuring >60% ambient light throughout the fruiting zone. A divided canopy lyre-trained vine doubles effective Canopy_Exterior_Role surface area by creating two separate exterior faces, with both curtain surfaces receiving the direct light that single-curtain systems provide only on one face. A sprawling, unmanaged vine has minimal Canopy_Exterior_Role area relative to total leaf area, with the dense, overlapping shoot growth creating extensive interior shading that limits both photosynthetic efficiency and fruit quality.
A canopy position role that inheres in the spatial region at the outer surface of the vine canopy that receives direct or minimally attenuated sunlight, experiencing full light intensity and the exposure conditions characteristic of unshaded positions. The canopy exterior represents the productive zone where leaf photosynthesis operates at maximum capacity, fruit receives the light exposure associated with optimal phenolic and flavor development, and the microclimate conditions (higher temperature, lower humidity, greater air movement) generally favor both vegetative function and fruit quality in temperate climates. Exterior-positioned leaves are the primary contributors to vine carbon balance, with their photosynthetic output supporting fruit ripening, reserve accumulation, and perennial structure maintenance. Exterior fruit exposure is associated with enhanced anthocyanin accumulation (color development), accelerated flavor compound synthesis, and more complete physiological ripeness compared to shaded interior fruit. Canopy management in quality viticulture fundamentally aims to maximize the proportion of canopy occupying exterior positions—through training system selection, shoot positioning, shoot and leaf removal—creating the open, well-exposed canopies associated with premium production.
Canopy_Exterior_Role captures the exposed canopy zone that modern viticulture seeks to maximize, recognizing that exterior positioning optimizes both photosynthetic function and fruit composition. The exterior/interior proportion is a key canopy management metric, with well-managed canopies achieving 80-90% exterior positioning while neglected or excessively vigorous canopies may have <50% exterior area. Training system selection fundamentally determines potential exterior area—divided canopy systems (lyre, GDC, Scott Henry) create more exterior surface than single-curtain systems at equivalent vine spacing, though with greater infrastructure and management requirements. The climate context matters in extremely hot regions, full exterior exposure may cause sunburn and heat damage, motivating retention of some protective interior canopy or use of shade cloth, in contrast to the full-exposure objectives of temperate-climate quality viticulture.
Außenbereich
Laubwand-Äußeres
canopy exterior
exponierte Zone
exposed canopy
exterior zone
outer canopy
äußere Laubwand
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Smart RE (1985) Canopy microclimate manipulation, American Journal of Enology and Viticulture 36: 230-239 Bergqvist J et al. (2001) Sunlight exposure and temperature effects on berry growth, American Journal of Enology and Viticulture 52: 1-7
Canopy Exterior Role
Canopy gaps quality manipulation in Casablanca Valley Sauvignon Blanc compared leaf removal strategies targeting different canopy positions. Control vines with no leaf removal exhibited canopy gaps quality of only 8% gap frequency with corresponding fruit zone PAR of 6% ambient and methoxypyrazine concentration of 32 ng/L. Basal leaf removal creating gaps specifically at fruit zone height achieved gap frequency of 28% with fruit zone PAR increasing to 19% and methoxypyrazines declining to 12 ng/L. Upper canopy leaf removal creating equivalent 28% gap frequency but positioned above the fruit zone improved overall light transmission but fruit zone PAR reached only 14% with methoxypyrazines at 18 ng/L, demonstrating that canopy gaps quality benefits depend not only on total gap percentage but critically on gap positioning relative to target tissues.
Canopy gaps quality is a viticultural quality that inheres in a grapevine canopy system and represents the presence, distribution, and characteristics of openings within the foliar mass where probe insertion or light transmission encounters no leaf obstruction, providing complementary characterization of canopy architecture that inversely relates to canopy density while specifically addressing the pathways available for light penetration, air circulation, and spray access to interior canopy positions including the fruit zone. This quality captures the functional importance of discontinuities in the leaf canopy that serve as essential conduits for environmental factors and management inputs that must reach interior tissues for effective fruit development and disease control. The ontological significance of canopy gaps quality lies in its recognition that optimal grapevine canopy architecture requires not merely appropriate total leaf area and canopy dimensions but also strategic distribution of openings that permit light exposure of fruit clusters, rapid canopy drying after wetting events, and effective pesticide deposition on interior surfaces that would otherwise remain protected by overlying foliage. Gaps in grapevine canopies arise from multiple sources including inherent discontinuities in shoot spacing along cordons or canes, incomplete leaf development in specific canopy positions, management-created openings through leaf removal or shoot thinning, and natural variation in shoot vigor and lateral development that creates irregular canopy fill. The measurement of canopy gaps quality through point quadrat analysis quantifies gap frequency as the percentage of probe insertions that traverse the canopy without contacting any leaves, providing standardized assessment enabling comparison across treatments, cultivars, and sites. Optimal canopy gaps quality targets generally recommend 20-40% gap frequency for wine grape production, balancing sufficient openings for fruit zone environmental optimization against the need to maintain adequate leaf area for photosynthesis and fruit protection from excessive sun exposure. The spatial distribution of gaps matters for their functional effectiveness, with gaps positioned at fruit zone height providing direct benefits for cluster exposure while gaps in upper canopy regions contribute primarily to reduced overall density rather than specific fruit zone improvement.
Canopy_Gaps_Quality serves as the parent class for Gap_Frequency_Quality, Gap_Size_Quality, and Cluster_Exposure_Quality which address specific aspects of canopy openings. This class addresses the overall presence and distribution of gaps as functional elements of canopy architecture complementary to density characterization. Distinguished from Canopy_Porosity_Quality by focusing specifically on complete openings rather than the general permeability concept. The percentage gaps metric from point quadrat analysis directly quantifies this quality, with 20-40% representing typical optimal target range.
Kronenlücken
Laubdiskontinuitäten
Laubwandlücken
Lückenverteilung
canopy discontinuities
canopy openings
foliage gaps
gap distribution
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 44-59 Dry PR (2000) Canopy management for fruitfulness. Australian Journal of Grape and Wine Research 6:109-115 Ryona I et al. (2008) Effects of cluster light exposure on 3-isobutyl-2-methoxypyrazine accumulation and degradation patterns in red wine grapes (Vitis vinifera L. cv. Cabernet Franc). Journal of Agricultural and Food Chemistry 56:10838-10846
Canopy Gaps Quality
Canopy height quality optimization in Mendoza Malbec evaluated effects of topping height on fruit composition and operational efficiency. Vines topped to maintain 1.0 m canopy height above the fruiting wire showed adequate but limited leaf area index of 1.8 and fruit Brix of 23.5° at target harvest date. Increased topping height to 1.3 m improved canopy height quality for photosynthetic capacity with LAI reaching 2.4 and Brix achieving 25.1° at equivalent hangtime. Further height increase to 1.6 m exceeded optimal canopy height quality, creating excessive shading at the fruit zone with light penetration declining from 18% to 9% of ambient, while also exceeding harvester head height requiring manual pre-harvest topping at additional cost. The 1.3 m canopy height quality target was established as optimal for this vigorous site balancing photosynthetic capacity, fruit zone environment, and mechanical harvest compatibility.
Canopy height quality is a canopy dimension quality that inheres in a grapevine canopy system and represents the vertical extent of the foliar mass measured from the lowest point of active foliage to the uppermost canopy boundary as determined by training system design parameters, shoot growth vigor, and the timing and intensity of topping operations that constrain upward canopy expansion. This quality fundamentally influences the total leaf area that can be accommodated within the canopy, the distribution of light through vertical canopy gradients, and the compatibility of the canopy with equipment designed for specific operational height windows. The ontological significance of canopy height quality lies in its role as a primary determinant of canopy surface area in vertically oriented training systems, wherein taller canopies present greater exterior surface for light interception while simultaneously creating longer path lengths for light penetration to interior and basal positions. The physiological basis of canopy height development reflects the cumulative extension growth of shoots from their point of origin on the vine permanent structure, with shoot length at any given time representing the integral of growth rate over the period since budbreak modified by any topping interventions removing apical portions. Optimal canopy height quality varies with training system objectives and practical constraints, with vertically shoot positioned systems typically targeting heights of 1.0-1.4 m above the fruiting wire to provide adequate leaf area for fruit ripening while maintaining compatibility with mechanical harvester head dimensions and avoiding excessive shading of basal fruit zone positions by upper canopy foliage. Excessively tall canopies create management challenges including difficulty achieving uniform spray coverage throughout the vertical extent, increased shading at lower canopy positions, potential for shoot breakage in wind events, and incompatibility with standard harvesting equipment, while insufficient canopy height may limit total photosynthetic capacity and yield potential. The measurement of canopy height quality employs direct measurement from established reference points, typically the fruiting wire height for vertically trained systems, using graduated measuring devices at multiple positions along the row to capture height variation. Temporal dynamics of canopy height quality follow shoot growth patterns, with rapid height increase during the spring growth flush followed by progressive slowing as shoots approach natural cessation or management-imposed height limits through topping, with target heights typically achieved 4-6 weeks post-budbreak in vigorous conditions.
Canopy_Height_Quality is_a Canopy_Dimension_Quality, representing the vertical extent parameter of canopy geometry. This class addresses height from lowest foliage to canopy top, distinct from cordon or fruiting wire heights which represent training system infrastructure positions. The typical VSP target of 1.0-1.4 m above fruiting wire provides quantitative reference, though optimal height varies with training system, vigor, and equipment specifications. Topping operations serve as primary management tool for canopy height quality control.
Kronenhöhe
Laubwandhöhe
Triebwandhöhe
canopy elevation
canopy vertical extent
foliage height
shoot wall height
vertikale Kronenausdehnung
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 18-24 Intrieri C, Poni S (1995) Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality of mechanized Italian vineyards. American Journal of Enology and Viticulture 46:116-127 Poni S et al. (2003) Effects of cluster thinning on yield and grape composition in field-grown Sangiovese. American Journal of Enology and Viticulture 54:102-108
Canopy Height Quality
The canopy height specification for a VSP-trained Sauvignon Blanc vineyard in Marlborough details a fruiting zone at 0.8-1.0 m above ground, a vegetative canopy extending from 1.0 m to 1.6 m (effective canopy wall height of 0.6 m after summer trimming), a total vine height of 1.6 m at a row spacing of 2.4 m yielding a canopy height-to-row-width ratio of 0.25, and a target leaf layer number (measured by point quadrat analysis) of 1.0-1.5 in the fruiting zone — the relatively low canopy height-to-row-width ratio ensures that the fruiting zone receives a minimum of 15% ambient photosynthetically active radiation (PAR), exceeding the 10% threshold below which Sauvignon Blanc develops excessive methoxypyrazine concentrations producing undesirable herbaceous character.
A training system description that documents the vertical dimensions of the vine canopy including the height of the fruiting zone above ground, the total canopy height from lowest foliage to trimming height, the height-to-row-width ratio, and the vertical distribution of leaf area and fruit within the canopy envelope, specifying the parameters that determine light interception efficiency, fruit zone microclimate, and spray penetration effectiveness.
Canopy_Height_Specification defines the vertical dimension of the canopy that, combined with row spacing and canopy density, determines the light environment experienced by both leaves and fruit. The canopy height-to-row-width ratio is a primary descriptor of vineyard light interception geometry: ratios above 1.0 indicate that canopy height exceeds row spacing (creating heavy shade in the lower canopy), ratios of 0.5-0.8 characterize most premium cool-climate VSP vineyards, and ratios below 0.3 indicate widely spaced rows with relatively short canopies typical of warm-climate irrigated viticulture. The Smart-Robinson canopy microclimate assessment framework uses canopy height measurements in combination with leaf layer number and percent interior clusters to evaluate whether a specific training system-canopy height combination is producing the desired fruit zone light and temperature environment for the target wine style.
Kronenhöhen-Parameter
Laubwandhöhen-Spezifikation
canopy dimension specification
canopy height documentation
vertical canopy parameters
vertikale Laubwand-Dokumentation
Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles pp: 91-120 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 331-380 Reynolds AG and Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition, American Journal of Enology and Viticulture 60: 251-268
Canopy Height Specification
A vigorous Merlot vine on fertile soil develops extensive Canopy_Interior_Role regions receiving <10% ambient light, with interior leaves becoming senescent by veraison and interior clusters showing green, unripe character at harvest—the excessive interior shading motivating shoot thinning or training system modification. A traditional gobelet-trained Grenache intentionally maintains Canopy_Interior_Role conditions around clusters, the self-shading architecture protecting fruit from Mediterranean sun intensity that would cause damage in more exposed positions. Lateral shoots developing in Canopy_Interior_Role positions contribute little to vine carbon balance due to light limitation, with their removal (lateral hedging) standard practice in quality-focused viticulture.
A canopy position role that inheres in the spatial region within the vine canopy that is shaded by surrounding foliage, receiving reduced light intensity (typically <30% of ambient) and experiencing modified microclimate conditions compared to the canopy exterior. The canopy interior develops in all but the most open vine architectures as shoot growth and leaf expansion progressively shade inner canopy regions, with interior conditions characterized by reduced photosynthetically active radiation (PAR), higher humidity, reduced wind movement, and more stable temperature compared to exposed exterior positions. Interior-positioned leaves have reduced photosynthetic capacity and may become net carbon consumers (respiration exceeding photosynthesis) under deep shade, while interior-positioned fruit typically shows delayed ripening, reduced color development, and altered flavor profiles compared to exposed fruit. Canopy interior management focuses on either maintaining interior openness (through shoot thinning, leaf removal, shoot positioning) or accepting interior conditions as appropriate for specific production objectives. The degree of interior shading correlates with canopy density, which itself reflects vine vigor, shoot number, and training system architecture.
Canopy_Interior_Role captures the shaded canopy volume that canopy management typically seeks to minimize or eliminate for quality wine production, recognizing that interior conditions are generally unfavorable for both leaf function (reduced photosynthesis) and fruit quality (delayed ripening, reduced phenolics). The interior/exterior distinction is fundamental to Smart's canopy microclimate framework, with the proportion of interior versus exterior canopy strongly predictive of fruit quality outcomes. Interior canopy quantification uses light porosity measurements (point quadrat analysis, light sensors) that document the fraction of canopy occupied by shaded interior positions. Climate change may shift the interior/exterior optimization, with some warming regions deliberately maintaining more interior shading to protect fruit from heat damage that exterior exposure would cause.
Innenbereich
Laubwand-Inneres
beschattete Zone
canopy interior
inner canopy
innere Laubwand
interior zone
shaded canopy
Smart RE (1985) Principles of grapevine canopy microclimate manipulation, American Journal of Enology and Viticulture 36: 230-239 Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Dokoozlian NK, Kliewer WM (1995) Light environment within grapevine canopies, American Journal of Enology and Viticulture 46: 209-218
Canopy Interior Role
The canopy management description for a Premier Cru Volnay vineyard documents a sequential program of six growing-season interventions — shoot thinning at E-L 12 (removing double shoots and non-fruitful water shoots to achieve 7-8 shoots per vine), first shoot positioning at E-L 15 (tucking shoots vertically between foliage wires), leaf removal on the east-facing side at E-L 29 (removing 4-5 basal leaves per shoot to expose the fruit zone to morning sun while retaining afternoon shade), second shoot positioning with lateral removal at E-L 31, hedging at E-L 33 (trimming shoot tips 15 cm above the top wire to redirect assimilate from vegetative growth to fruit ripening), and cluster thinning at véraison (removing second clusters, green-shouldered clusters, and lagging berries to achieve 6-7 clusters per vine at a target yield of 40 hl/ha).
A viticultural practice description that documents the suite of growing-season interventions applied to manipulate the spatial arrangement, density, and microclimate of the grapevine canopy, encompassing shoot positioning, leaf removal, shoot thinning, hedging, cluster thinning, and fruit zone management operations that collectively optimize the balance between light interception, air circulation, disease pressure, and fruit quality within the architectural framework established by the training system and pruning method.
Canopy_Management_Description documents the growing-season operations that refine and optimize the canopy architecture established by winter pruning and the training system infrastructure. The conceptual framework for systematic canopy management was developed principally by Richard Smart and colleagues from the 1980s onward, codified in the landmark handbook Sunlight into Wine (Smart and Robinson 1991) which established the principle that many wine quality deficiencies attributed to climate or terroir are actually consequences of suboptimal canopy microclimate that can be corrected through targeted canopy manipulation: this insight transformed viticulture from a predominantly winter pruning-focused discipline to one in which growing-season canopy management receives equal or greater attention and resource allocation. Subclasses capture the principal canopy management operations in their typical chronological sequence through the growing season.
Grünarbeit-Beschreibung
Laubarbeit-Dokumentation
Laubwandmanagement-Beschreibung
canopy manipulation record
green operations documentation
growing-season canopy operations
Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 331-380 Reynolds AG (ed) (2022) Managing Wine Quality Volume 1: Viticulture and Wine Quality, 2nd Ed, Woodhead Publishing
Canopy Management Description
Canopy microclimate data for a Scott Henry-trained Merlot vineyard in Hawke's Bay, New Zealand demonstrated that the upper (upward-positioned) curtain received 35% more PAR and maintained leaf temperatures 2-3°C higher than the lower (downward-positioned) curtain, resulting in 15% greater photosynthetic rates in upper shoots measured by portable gas exchange systems. Canopy microclimate sensors in a high-density (8000 vines/ha) Tempranillo plantation in Ribera del Duero recorded canopy interior temperatures exceeding ambient by 4-6°C during midday in July, with leaf-to-air vapor pressure deficit (VPD) reaching 4.5-5.0 kPa at the sun-exposed canopy exterior versus 2.0-2.5 kPa in shaded interior positions.
Canopy microclimate data is a microclimate characterization entity that records the environmental conditions within and surrounding the aerial vegetative structure of a grapevine, including temperature, humidity, radiation interception, and air movement parameters measured at the leaf surface, shoot zone, and canopy boundary layer, quantifying the photosynthetic environment and transpirational demand experienced by leaves and the light regime governing canopy carbon balance and vegetative growth allocation.
Canopy microclimate data provides the mechanistic link between vineyard management interventions and vine physiological responses, as training system design, shoot density, and leaf area index directly determine the radiation, temperature, and humidity environment in which photosynthesis and transpiration occur. The Smart-Dyson point quadrat method provides a standardized approach for characterizing canopy microclimate through estimation of leaf layer number (LLN), percentage of gaps, percentage of interior clusters, and percentage of interior leaves — metrics that correlate strongly with directly measured radiation and temperature parameters. Optimal canopy microclimates for quality wine production typically feature LLN of 1.0-1.5, gap frequencies of 20-40%, and interior cluster percentages below 40%. Dense canopies (LLN > 3) create conditions favoring vegetative flavor characters (methoxypyrazines), reduced fruit color, and elevated disease pressure, while excessively open canopies risk sunburn damage and excessive berry temperature. Wireless sensor networks enable continuous multi-point canopy microclimate monitoring that reveals spatial and temporal heterogeneity invisible to periodic manual assessment.
Bestandsklimatologie
Blattzonenmikroklima
Laubwandklima
Laubwandmikroklimadaten
canopy atmosphere data
canopy climate data
canopy environment data
leaf zone microclimate
vegetative zone microclimate data
Smart RE, Robinson MD (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles, pp.: 30-55 Smart RE (1985) Principles of grapevine canopy microclimate manipulation. American Journal of Enology and Viticulture 36(3): 230-239 Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review. American Journal of Enology and Viticulture 60(3): 251-268
Canopy Microclimate Data
Canopy microclimate quality characterization in a Côte de Beaune Pinot Noir vineyard compared environmental conditions between dense unmanaged canopies and shoot-thinned, leaf-removed treatments during the August véraison-to-harvest period. Dense canopies exhibited strongly modified canopy microclimate quality with fruit zone maximum temperatures averaging 3.2°C below ambient, minimum temperatures 1.8°C above ambient creating dampened diurnal amplitude of only 8.4°C versus 13.4°C ambient, relative humidity averaging 82% versus 65% ambient, and fruit zone PAR of only 45 μmol m⁻² s⁻¹ at solar noon. Managed canopies showed improved canopy microclimate quality with maximum temperatures only 0.8°C below ambient, minimums 0.5°C above, humidity of 71%, and fruit zone PAR of 185 μmol m⁻² s⁻¹, creating conditions more favorable for phenolic development while reducing Botrytis infection periods through faster drying after precipitation events.
Canopy microclimate quality is a viticultural quality that inheres in a grapevine canopy system and represents the ensemble of environmental conditions existing within the canopy interior and fruit zone that differ from ambient atmospheric conditions due to the modifying effects of the foliar mass on radiation penetration, air movement, humidity, and temperature through interception, absorption, reflection, transpiration, and boundary layer formation. This quality fundamentally determines the environmental conditions directly experienced by developing grape clusters and interior leaves, which may differ substantially from weather station measurements or ambient vineyard conditions and which ultimately drive the physiological processes of fruit ripening and quality development. The ontological significance of canopy microclimate quality lies in its role as the proximate environmental determinant of berry composition, wherein the temperature, light, and humidity regime at cluster positions during the critical ripening period shapes organic acid metabolism, phenolic accumulation, aromatic precursor development, and disease susceptibility independently of broader mesoclimate or macroclimate characteristics. The physical basis of canopy microclimate modification involves multiple interacting processes including attenuation of solar radiation as it passes through successive leaf layers, conversion of absorbed radiation to sensible and latent heat through leaf energy balance processes, transpirational water vapor release that elevates humidity within the canopy boundary layer, and reduction of wind speed and turbulent mixing in canopy interior positions that restricts exchange with ambient air. The relationship between canopy architecture and microclimate quality follows predictable patterns wherein dense canopies with high leaf layer numbers create strongly modified interior conditions with reduced light, elevated humidity, and dampened temperature extremes compared to well-ventilated open canopies where interior conditions more closely track ambient atmospheric states. Measurement of canopy microclimate quality requires deployment of sensors at specific canopy positions including the fruit zone, with temperature, humidity, and light intensity representing the primary parameters, supplemented by leaf wetness monitoring for disease-relevant applications. Management interventions affecting canopy microclimate quality operate primarily through canopy density and architecture manipulation, with shoot thinning, leaf removal, and shoot positioning modifying the radiation interception and air flow characteristics that drive microclimate formation. The temporal dynamics of canopy microclimate quality exhibit both seasonal patterns following canopy development and daily cycles reflecting solar angle progression and atmospheric boundary layer evolution.
Canopy_Microclimate_Quality serves as the parent class for specific environmental parameters including Fruit_Zone_Temperature_Quality, Fruit_Zone_Humidity_Quality, and Fruit_Zone_Light_Intensity_Quality. This class addresses the integrated environmental conditions within the canopy rather than individual parameters. Distinguished from Canopy_Density_Quality and Canopy_Gaps_Quality which address canopy structural characteristics that create microclimate conditions. Strong functional relationships exist with disease development, fruit composition, and vine physiological responses.
Bestandesklima
Fruchtzonenklima
Kronenmikroklima
Laubwandmikroklima
canopy environment
fruit zone microclimate
interior canopy climate
within-canopy conditions
Smart RE (1985) Principles of grapevine canopy microclimate manipulation with implications for yield and quality: a review. American Journal of Enology and Viticulture 36:230-239 Dry PR, Smart RE (1988) The grapegrowing regions of Australia. In: Coombe BG, Dry PR (eds) Viticulture Volume 1 Resources, Winetitles, Adelaide, pp. 37-60 Schultz HR (2000) Climate change and viticulture: a European perspective on climatology, carbon dioxide and UV-B effects. Australian Journal of Grape and Wine Research 6:2-12
Canopy Microclimate Quality
Canopy porosity quality manipulation in Hawke's Bay Merlot compared fruit zone outcomes across three porosity levels achieved through differential leaf removal intensity. Low porosity treatments with 12% gaps showed fruit methoxypyrazine concentrations of 28 ng/L and anthocyanin levels of 0.9 mg/g berry weight. Medium porosity with 30% gaps reduced methoxypyrazines to 14 ng/L while increasing anthocyanins to 1.4 mg/g. High porosity at 48% gaps further reduced methoxypyrazines to 6 ng/L with anthocyanins reaching 1.7 mg/g, though excessive fruit exposure in this treatment caused sunburn damage to 8% of berries on western canopy aspects, demonstrating the need to balance canopy porosity quality optimization against fruit protection requirements.
Canopy porosity quality is a canopy density quality that inheres in a grapevine canopy system and represents the proportion and distribution of open spaces or gaps within the foliar mass that permit penetration of light, air, and applied materials through the canopy volume to reach interior tissues and fruit clusters. This quality provides functional characterization of canopy openness that directly relates to the environmental conditions experienced by fruit and the effectiveness of management operations requiring canopy penetration. The ontological basis of canopy porosity quality rests on the recognition that gaps in the leaf canopy serve as conduits for energy and mass transfer that fundamentally influence fruit zone microclimate, disease development potential, and spray deposition efficiency. Porous canopies with well-distributed gaps permit solar radiation to reach fruit clusters promoting phenolic synthesis, anthocyanin accumulation, and flavor precursor development, while facilitating rapid drying after precipitation or dew that reduces pathogen infection periods. The spatial distribution of porosity matters as much as total gap percentage, with uniformly distributed small gaps providing better overall penetration than equivalent total gap area concentrated in few large openings. Measurement of canopy porosity quality employs several complementary approaches including visual gap estimation, point quadrat gap frequency determination through systematic probe insertion, transmitted light measurement using line quantum sensors, and digital image analysis calculating gap fraction from photographs taken through or of the canopy. The relationship between canopy porosity quality and fruit composition reflects the fundamental importance of light exposure for grape quality development, with research consistently demonstrating positive associations between fruit zone porosity and concentrations of anthocyanins, flavonols, and desirable aroma compounds, alongside negative associations with methoxypyrazine concentrations responsible for herbaceous character. Management interventions to improve canopy porosity quality include shoot positioning to minimize leaf overlap, leaf removal particularly in the fruit zone, shoot thinning to reduce canopy density at its source, and hedging to remove lateral growth that fills canopy gaps. The temporal dynamics of canopy porosity quality typically show declining porosity through the early growing season as shoots extend and leaves expand, with management interventions required to maintain or restore adequate porosity levels approaching véraison when fruit exposure becomes critical for quality development.
Canopy_Porosity_Quality is_a Canopy_Density_Quality, representing the functional openness dimension of density characterization. This class serves as parent for Light_Penetration_Quality, Air_Circulation_Quality, and Spray_Penetration_Quality which address specific consequences of canopy porosity. Distinguished from Leaf_Layer_Number_Quality which quantifies the layering of foliage rather than the gaps between leaves. The percentage gaps metric from point quadrat analysis directly quantifies this quality.
Kronenporosität
Laubdurchlässigkeit
Laubwandporosität
Lückenfraktion
canopy gap percentage
canopy openness
foliage porosity
gap fraction
Smart RE (1985) Principles of grapevine canopy microclimate manipulation with implications for yield and quality: a review. American Journal of Enology and Viticulture 36:230-239 Gladstone EA, Dokoozlian NK (2003) Influence of leaf area density and trellis/training system on the light microclimate within grapevine canopies. Vitis 42:123-131 Joscelyne VL et al. (2007) Determination of grape (Vitis vinifera L. cv. Shiraz) leaf phenolics using scanning spectroscopy: a comparison of canopy porosity treatments. Australian Journal of Grape and Wine Research 13:88-96
Canopy Porosity Quality
A cluster positioned in the exterior fruit zone bears a Canopy_Position_Role characterized by high light exposure, elevated temperature, and air movement—conditions favoring phenolic development but risking sunburn in hot climates. A shoot in the interior canopy position serves in a Canopy_Position_Role receiving <10% ambient light, with shaded conditions limiting photosynthetic contribution while the shoot competes for resources with more favorably positioned growth. Lateral shoots emerging from upper nodes bear Canopy_Position_Roles in the canopy exterior where their potential for secondary fruit production and photosynthetic contribution differs from basal lateral positions.
A vineyard role that inheres in a grapevine structure or organ by virtue of its spatial position within the vine canopy, with position determining light exposure, microclimate conditions, and functional significance for vine physiology and fruit development. Canopy position roles recognize that vine organs at different spatial locations experience fundamentally different environmental conditions—exterior positions receive direct light while interior positions are shaded upper canopy positions experience higher temperature and wind exposure than lower positions and fruit zone positioning determines the balance of exposure versus protection that influences fruit composition. Position-based role assignment enables precise description of where specific structures or developmental events occur within the three-dimensional canopy architecture, supporting targeted management interventions (selective leaf removal, differential harvest) and understanding of position-quality relationships. Canopy position concepts are fundamental to modern viticulture, where training system selection, shoot positioning, and canopy management all aim to optimize the spatial arrangement of vegetative and reproductive structures for quality outcomes.
Canopy_Position_Role provides the spatial framework for describing vine architecture, subclassed by specific zones (Fruiting_Zone and others) that define functionally significant canopy regions. The importance of canopy position reflects viticulture's transformation from empirical practice to science-based management—Smart's canopy microclimate research demonstrated that fruit quality correlates strongly with position-dependent exposure variables, making position management central to quality optimization. Position roles interact with training system—the same "fruiting zone" concept applies differently in VSP (narrow vertical band), GDC (pendant curtain perimeters), and bush vine (distributed throughout the canopy) architectures. Contemporary precision viticulture increasingly maps position-related variables (light interception, canopy density) to guide targeted interventions at sub-vineyard scale.
Architekturposition
Laubwandposition
Rebenposition
architectural position
canopy position
räumliche Position
spatial position
vine position
Smart RE (1985) Principles of grapevine canopy microclimate manipulation, American Journal of Enology and Viticulture 36: 230-239 Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Dokoozlian NK, Kliewer WM (1995) The light environment within grapevine canopies, American Journal of Enology and Viticulture 46: 209-218
Canopy Position Role
Canopy surface area quality comparison in a Riverland Shiraz trial evaluated training system effects on light interception efficiency and water use productivity. Single wire sprawl training produced nominal canopy surface area quality of 12,500 m² per hectare but achieved only 65% total light interception due to self-shading within the irregular canopy profile. Conversion to VSP increased effective canopy surface area quality through improved organization, achieving 78% light interception despite similar nominal surface area of 13,200 m² per hectare. Implementation of Shaw minimal pruning dramatically increased nominal canopy surface area quality to 22,000 m² per hectare through three-dimensional canopy expansion, achieving 88% light interception. Yield increased proportionally from 8 t/ha in sprawl to 11 t/ha in VSP to 18 t/ha in minimal pruning, with water use efficiency improving from 4.2 to 5.8 to 7.1 kg fruit per kiloliter of irrigation, demonstrating the productivity benefits of canopy surface area quality optimization through training system design.
Canopy surface area quality is a canopy dimension quality that inheres in a grapevine canopy system and represents the total exterior surface of the foliar mass available for light interception and gas exchange as determined by canopy height, width, continuity along the row, and the geometric complexity of the canopy profile that may increase actual surface area relative to simple rectangular approximations. This quality provides integrative characterization of canopy light capture potential that relates more directly to photosynthetic productivity than individual height or width dimensions alone, enabling comparison of vineyard systems with different canopy geometries on a common functional basis. The ontological significance of canopy surface area quality lies in its role as the primary determinant of total light interception when considered relative to vineyard ground area, wherein canopy surface area per hectare establishes the upper limit of radiation capture that drives photosynthesis, yield formation, and fruit quality development. The computation of canopy surface area employs various geometric approaches depending on canopy architecture complexity, ranging from simple rectangular approximation multiplying height by two (accounting for both canopy faces) by row length, to more sophisticated methods incorporating width variation with height, canopy curvature, and the additional surface created by gaps and canopy discontinuities that expose interior surfaces to direct light. Training systems designed to maximize canopy surface area quality achieve this objective through canopy division creating multiple separate curtains each presenting exterior surface, or through canopy geometry manipulation positioning foliage at angles to maximize exposed surface area to volume ratios. The relationship between canopy surface area quality and yield potential reflects the fundamental requirement for approximately 1.0-1.5 m² of well-exposed leaf area per kilogram of fruit for complete ripening, translating to canopy surface area requirements of 10,000-15,000 m² per hectare for typical yield targets, though actual requirements vary with climate, cultivar, and ripeness objectives. Measurement of canopy surface area quality extends beyond simple dimensional measurement to functional assessment using light interception as the ultimate indicator of effective surface area, recognizing that nominal geometric surface area may not translate fully to productive light capture if portions are self-shaded or oriented unfavorably relative to solar angles.
Canopy_Surface_Area_Quality is_a Canopy_Dimension_Quality, representing the integrative exterior surface parameter derived from height, width, and continuity. This class addresses the light interception surface that determines photosynthetic potential, distinct from total leaf area which includes interior leaves that may not contribute effectively to light capture. The target of 10,000-15,000 m² per hectare for typical production provides quantitative reference. Canopy division and geometry manipulation through training system design serve as primary approaches to canopy surface area quality optimization.
Kronenoberfläche
Laubwandoberfläche
Lichtfangfläche
canopy light interception surface
exponierte Kronenfläche
exposed canopy area
exterior canopy surface
foliar surface area
Smart RE (1973) Sunlight interception by vineyards. American Journal of Enology and Viticulture 24:141-147 Schultz HR (1995) Grape canopy structure, light microclimate and photosynthesis. I. A two-dimensional model of the spatial distribution of surface area densities and leaf ages in two canopy systems. Vitis 34:211-215 Poni S et al. (2003) Grapevine quality: a multiple choice issue. Scientia Horticulturae 234:445-462
Canopy Surface Area Quality
Canopy temperature mapping of a 40-hectare Cabernet Sauvignon vineyard in Paso Robles using drone-mounted thermal imaging at 14:00 reveals canopy temperatures ranging from 28°C in well-watered blocks with drip irrigation to 36°C in deficit-irrigated blocks, with the 8°C temperature range corresponding to predawn water potential differences from -0.3 to -1.1 MPa, enabling targeted irrigation to zones exceeding the 33°C threshold associated with photosynthetic decline and quality compromise.
Canopy temperature quality is a tissue temperature quality that inheres in the grapevine vegetation mass as an integrated system and represents the aggregate thermal state of the leaf and shoot population characterizing the spatially averaged temperature across the three-dimensional canopy structure where individual organ temperatures vary with position, exposure, and local microclimate. Within the BFO framework, this quality addresses the emergent thermal property of the canopy as a functional unit, integrating contributions from sunlit and shaded leaves, young and mature tissues, and varying degrees of transpirational cooling capacity across the heterogeneous vegetation mass. The ontological significance lies in canopy temperature serving as a practical metric for irrigation management and stress assessment at scales relevant to vineyard operations, where individual leaf measurements cannot capture the whole-vine thermal status but canopy-level assessment integrates multiple organ temperatures into a single manageable parameter. Canopy temperature is typically measured using infrared thermometry averaging radiation from a defined field of view encompassing multiple leaves and shoots, or through thermal imaging providing spatial temperature distributions that can be processed to extract mean, maximum, or percentile statistics characterizing canopy thermal heterogeneity. The relationship between canopy temperature and air temperature provides the basis for crop water stress indices (CWSI) where well-watered canopies maintain temperatures below air temperature through transpirational cooling while water-stressed canopies approach or exceed air temperature. Remote sensing approaches enable canopy temperature mapping across vineyard blocks using airborne or satellite thermal imagery, supporting precision irrigation management by identifying spatial variation in water stress. Understanding canopy temperature dynamics enables optimization of irrigation timing and volume, prediction of heat stress impacts on vine productivity, and interpretation of seasonal effects on fruit composition.
Canopy_Temperature_Quality addresses the aggregate canopy scale, complementing individual Leaf_Temperature_Quality measurements and enabling practical stress assessment at management-relevant scales. This quality forms the basis for thermal stress indices and remote sensing applications in precision viticulture. The class encompasses the spatially averaged temperature of vegetative structures, distinguishing from cluster or berry temperatures that may differ due to position and organ-specific properties.
Bestandstemperatur
Kronendachtemperatur
Laubwandtemperatur
Vegetationstemperatur
canopy thermal state
foliage temperature
integrated leaf temperature
vegetation surface temperature
Möller M et al. (2007) Use of thermal and visible imagery for estimating crop water status of irrigated grapevine. Journal of Experimental Botany 58(4): 827-838 Bellvert J et al. (2014) Mapping crop water stress index in a Pinot-noir vineyard: comparing ground measurements with thermal remote sensing imagery from an unmanned aerial vehicle. Precision Agriculture 15(4): 361-376
Canopy Temperature Quality
Canopy transpiration monitoring in Cabernet Sauvignon vineyard using sap flow sensors reveals the seasonal water demand pattern, with early season (small canopy, 1.2 m² leaf area/vine) transpiration of 1.8 L/vine/day, mid-season peak (full canopy, 4.5 m²) of 6.5 L/vine/day under high evaporative demand, and late season decline (partial senescence, declining stomatal conductance) to 3.2 L/vine/day, with seasonal total of approximately 750 L/vine (equivalent to 450 mm over the planted area) representing the water consumption requiring replacement through irrigation (520 mm applied) plus effective rainfall (180 mm) to maintain vine water status throughout the season.
Canopy transpiration quality is a transpiration rate quality that represents the total water vapor flux from the entire vine canopy to the atmosphere, characterizing the integrated water loss from all transpiring surfaces that determines whole-plant water demand and the water balance that must be maintained through irrigation or rainfall for sustained vine function. Within the BFO framework, this quality specifically addresses the whole-plant scale of transpiration, distinguishing integrated canopy flux from leaf-level rates while recognizing that canopy transpiration is the agronomically and hydrologically relevant measure that determines vineyard water requirements, irrigation scheduling needs, and the water footprint of grape production. The ontological significance lies in canopy transpiration determining the actual water consumption of vineyards that must be budgeted in water resource planning, where the integration of leaf-level transpiration across heterogeneous canopy environments, variable stomatal conductance, and different leaf types yields the total water flux that defines vineyard water demand and the sustainability of production in water-limited regions. The scaling from leaf to canopy transpiration involves integration across leaf area (total transpiring surface), canopy light environment (affecting stomatal conductance distribution), aerodynamic coupling (wind effects on boundary layer conductance), and canopy structure (affecting the microclimate experienced by individual leaves), with these factors creating complex relationships between leaf-level conductance and canopy-scale flux. The measurement approaches for canopy transpiration include sap flow sensors (measuring water flux through trunk or branches), lysimetry (weighing whole vines to detect water loss), eddy covariance (measuring water vapor flux above the canopy), and modeling approaches scaling leaf-level measurements to canopy using light distribution and leaf area models. The typical values for canopy transpiration in productive vineyards range from 2-8 mm/day (20-80 m³/ha/day) depending on climate, canopy size, and water availability, with seasonal totals of 300-700 mm representing a major component of vineyard water balance. Understanding canopy transpiration quality enables irrigation scheduling based on actual crop water use, water footprint assessment for sustainability certification, and evaluation of management practices affecting vineyard water demand.
Canopy_Transpiration_Quality addresses the whole-plant scale of water loss that determines vineyard water demand. This quality is the agronomically relevant measure for irrigation management and water resource planning. The class supports water budgeting and irrigation scheduling through canopy-scale transpiration assessment.
Gesamtpflanzenwasserverlust
Kronen-ET
Kronentranspiration
Rebenwasserverbrauch
canopy ET
vine transpiration
vineyard water use
whole-plant water loss
Williams LE, Ayars JE (2005) Grapevine water use and the crop coefficient are linear functions of the shaded area measured beneath the canopy. Agricultural and Forest Meteorology 132(3-4): 201-211 Intrigliolo DS, Castel JR (2006) Vine and soil-based measures of water status in a Tempranillo vineyard. Vitis 45(4): 157-163
Canopy Transpiration Quality
Canopy width quality manipulation in Carneros Chardonnay compared shoot positioning intensities and their effects on fruit zone environment. Minimal positioning allowing natural shoot sprawl produced canopy width quality of 0.85 m with leaf layer number of 2.6 and fruit zone PAR of only 8% ambient. Standard VSP positioning reducing width to 0.55 m achieved improved leaf layer number of 1.8 and fruit zone PAR of 16%. Intensive positioning with double catch wires maintaining canopy width quality of 0.35 m further reduced leaf layer number to 1.2 and increased fruit zone PAR to 24%, though requiring twice the positioning labor of standard practice. Economic analysis identified the standard 0.55 m canopy width quality target as optimal, achieving 75% of the quality improvement of intensive positioning at 40% of the labor cost, with fruit from this treatment producing wines with preferred balance of citrus character and reduced vegetal notes in sensory evaluation.
Canopy width quality is a canopy dimension quality that inheres in a grapevine canopy system and represents the horizontal extent of the foliar mass measured perpendicular to the row direction as determined by training system design, shoot orientation management, and the lateral growth characteristics of the cultivar under prevailing growing conditions. This quality fundamentally influences the proportion of canopy leaf area positioned in well-exposed exterior locations versus shaded interior positions, the penetration distance required for light, air, and spray materials to reach canopy center, and the clearance available in inter-row alleyways for equipment passage and worker access. The ontological significance of canopy width quality lies in its role as a primary determinant of canopy density when considered relative to leaf area, wherein narrow canopies concentrate equivalent foliage within reduced horizontal extent creating higher leaf area density compared to wider canopies that distribute leaves across greater cross-sectional area, reducing layering and improving interior exposure. The physiological basis of canopy width development involves both the natural spreading tendency of shoots growing under gravity and solar tracking influences and the management-imposed constraints of shoot positioning, catch wire placement, and hedging that confine lateral canopy expansion. Optimal canopy width quality varies substantially with training system philosophy, ranging from the narrow profiles of approximately 0.3-0.5 m targeted by vertically shoot positioned systems designed to minimize shading through canopy compaction, to the wider profiles of 0.6-1.0 m or greater characteristic of minimal pruning and sprawling systems that accept increased width as the tradeoff for reduced management input. The relationship between canopy width quality and row spacing determines the proportion of vineyard area occupied by canopy versus alleyway, with width-to-row-spacing ratios influencing total light interception, disease pressure through canopy isolation, and the economic efficiency of land utilization. Measurement of canopy width quality employs horizontal transect assessment at standardized heights, typically at mid-canopy and fruit zone levels, using graduated devices to record the distance from outer leaf on one canopy face to outer leaf on the opposite face, with multiple measurements along the row characterizing width uniformity.
Canopy_Width_Quality is_a Canopy_Dimension_Quality, representing the horizontal extent parameter of canopy geometry perpendicular to the row. This class addresses the cross-sectional width of the foliar mass that determines density characteristics for given leaf area. Distinguished from row spacing which represents vineyard infrastructure design. VSP systems typically target 0.3-0.5 m width, while sprawling systems may exceed 1.0 m. Shoot positioning and hedging serve as primary management tools for canopy width quality control.
Kronenbreite
Laubwandbreite
Laubwanddicke
canopy cross-section
canopy thickness
foliage width
horizontal canopy extent
horizontale Kronenausdehnung
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 18-24 Gladstone EA, Dokoozlian NK (2003) Influence of leaf area density and trellis/training system on the light microclimate within grapevine canopies. Vitis 42:123-131 Vanden Heuvel JE et al. (2004) Influence of training system and shoot density on the light microclimate and fruit composition of Vitis vinifera L. cv. Riesling. American Journal of Enology and Viticulture 55:87-97
Canopy Width Quality
Cabernet Sauvignon in Napa Valley shows cap fall during early-mid June Year N+1 at BBCH 60-65 over 7-9 days, progressing from first calyptra detachment June 4-5 through peak cap shedding June 7-10 with abundant fallen caps accumulating on clusters and ground, completing June 12-13, under warm climate conditions of 22-28°C temperature, 48-65% relative humidity supporting efficient detachment with minimal retention problems, typical of this late-season cultivar showing good cap shedding efficiency enabling 55-68% fruit set
Cap fall represents the distinctive grapevine flower opening mechanism where fused petal calyptra detaches as intact unit from receptacle base and falls away exposing reproductive organs, serving as definitive visual marker of anthesis onset and progression creating characteristic flowering appearance. This essential reproductive process occurs during flowering in early-mid June in temperate regions Year N+1, typically spanning 5-10 days from first cap detachment through mass cap fall completion as flowers progressively open throughout cluster, corresponding to BBCH 60-65 stages and serving as practical observational marker for anthesis timing. Cap structure represents unique grapevine floral morphology with five petals fused at tips forming coherent calyptra cap covering reproductive organs during bud development, cap remaining closed protecting developing stamens and pistil until anthesis initiation, and distinctive pentamerous form visible on detached caps showing five fused petal sections. Detachment mechanism involves specialized abscission zone development at junction between calyptra base and receptacle, enzymatic degradation of middle lamella in abscission zone cells creating separation plane through cellulase and pectinase activity, mechanical forces from expanding reproductive organs pushing upward against cap creating pressure, and combined enzymatic and mechanical processes driving physical cap separation and fall. Abscission zone anatomy shows specialized cell layers at calyptra-receptacle junction with smaller cells containing dense cytoplasm, thin cell walls particularly in middle lamella region facilitating enzymatic breakdown, vascular tissue disruption at separation plane, and coordinated cellular changes enabling clean cap detachment. Cap lifting sequence shows characteristic progression with initial loosening visible as slight gap appearing at base between cap and receptacle, progressive upward movement as abscission proceeds with cap elevating 0.5-2.0 mm above receptacle, continued lifting driven by expanding stamens and pistil pushing against enclosing cap, and final detachment when abscission completes and mechanical disturbance or gravity causes cap fall. Detached cap characteristics include intact fused petal structure maintaining pentamerous organization, attachment scar at base showing clean separation plane, occasional retention of stamens or pistil fragments if mechanical separation irregular, and accumulation of fallen caps on cluster surface and ground below creating visible flowering debris. Cap fall timing relative to anthesis shows detachment typically initiating anthesis process with reproductive organs exposing immediately upon cap removal, occasional delay of hours between cap loosening and actual fall creating brief transition period, and cap detachment representing definitive marker that flower has entered open reproductive state. Reproductive organ exposure following cap fall reveals functional flower structures with five stamens surrounding central pistil positioned for pollination, anthers beginning or completing dehiscence releasing pollen, stigma developing or showing receptivity with papillate surface and exudate production, and overall flower achieving functional reproductive state. Environmental effects on cap fall show strong humidity dependence with moderate 45-70% relative humidity optimal for normal cap detachment, high humidity above 75% interfering with abscission through maintained turgor and possibly affecting enzymatic activity delaying or preventing detachment, and low humidity below 40% potentially causing premature desiccation affecting tissues though generally facilitating mechanical separation. Temperature affects cap fall with optimal 20-28°C supporting normal abscission processes and flower function, cool conditions below 15°C substantially delaying detachment through slowed enzymatic activity, and warm conditions above 32°C potentially accelerating processes though risking tissue damage from heat stress. Rain during cap fall represents serious interference with water physically preventing cap detachment through elevated humidity and mechanical impedance, causing caps to remain attached or detach incompletely affecting reproductive organ exposure, and promoting fungal colonization on wet flower tissues particularly botrytis establishing infections. Wind effects show minor positive role with air movement potentially assisting final mechanical cap detachment once abscission completes, and removing detached caps from cluster vicinity reducing accumulated debris, though strong wind may cause damage to newly exposed reproductive structures. Diurnal pattern shows most cap fall occurring during morning hours as temperature rises and humidity declines creating favorable conditions, reduced activity during afternoon and evening when humidity may increase, and daily waves of cap fall creating characteristic temporal rhythm during flowering period. Cap retention problems occur when abscission fails or proceeds incompletely resulting in caps remaining partially or fully attached preventing normal reproductive organ exposure and function, retention associated with unfavorable environmental conditions particularly high humidity, cool temperatures, or developmental abnormalities, and causing reproductive failure through coulure creating incomplete fruit set. Coulure relationship shows cap retention as mechanical cause of poor fruit set with attached caps preventing anther dehiscence eliminating pollen availability, blocking stigma exposure preventing pollination, and causing flower abortion through failed reproduction, creating substantial yield losses particularly in susceptible cultivars or unfavorable flowering conditions. Observational utility of cap fall includes serving as definitive visual marker for anthesis onset enabling accurate flowering date determination, providing clear indication of flowering progression through cluster as cap fall advances spatially, and allowing assessment of flowering uniformity and potential problems through observation of retention patterns. Cap accumulation patterns create visible evidence of flowering with detached caps accumulating on cluster surfaces showing recent opening activity, fallen caps on ground below vines indicating flowering extent and timing, and progressive accumulation documenting flowering progression over time. Flowering intensity assessment uses cap fall as indicator with heavy cap fall indicating intense flowering activity, sparse cap fall suggesting limited or irregular flowering, and spatial patterns revealing developmental gradients within clusters or across vineyard. Developmental gradient visualization through cap fall shows temporal sequence with apical regions showing earlier cap fall from earliest-maturing flowers, middle regions showing progressive cap fall during peak activity, and basal regions showing latest cap fall reflecting developmental delays. The process shows two major phases progressing from initial cap fall marking anthesis onset with sporadic early detachments, through mass cap fall representing peak flowering with widespread simultaneous detachment. Duration typically spans 5-10 days from first cap detachment through completion depending on flowering synchrony and environmental conditions, with warm stable conditions promoting compressed 5-7 day period while variable conditions extend period to 10-14 days. Cultivar variation appears in cap detachment efficiency with some varieties showing clean complete detachment supporting high fruit set while others show frequent retention problems causing coulure, cap fall timing relative to overall phenology, and sensitivity to environmental conditions affecting detachment success. Regional timing shows cap fall during late May-early June in warm climates, early-mid June in temperate regions, and mid-late June in cool areas, coinciding with anthesis period and serving as practical flowering marker. Viticultural significance includes providing clear visual marker for flowering timing and progression enabling management scheduling, indicating anthesis onset and intensity supporting yield prediction, revealing retention problems indicating potential fruit set failures requiring attention, and serving as practical observational reference more easily assessed than detailed reproductive organ examination. Research applications include documenting precise flowering timing using cap fall as objective marker, analyzing environmental effects on abscission processes affecting flowering success, evaluating cultivar differences in cap detachment efficiency affecting reproductive reliability, quantifying relationships between cap fall patterns and fruit set outcomes, and assessing mechanisms coordinating cap abscission with reproductive organ maturation.
Cap_Fall represents distinctive grapevine flower opening mechanism serving as practical observable marker for anthesis. Occurs within Flowering corresponding to BBCH 60-65 parallel to Anthesis phases. Subdivided into Initial_Cap_Fall marking onset and Mass_Cap_Fall representing peak activity. Environmental conditions particularly humidity critically affect detachment efficiency. Retention problems cause coulure reducing fruit set. Serves as definitive visual reference for flowering timing and progression. Easier to observe than detailed reproductive organ status.
Blütenkappenabtrennung
Kalyptrafall
Kalyptralösung
Kappenabwurf
calyptra fall
cap shedding
petal cap detachment
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 360-375 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 275-285 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Cap Fall
Chardonnay in Champagne shows cap fall completion on approximately July 2 Year N+1 at BBCH 69, with all viable flowers 100% having completed cap detachment totaling 12-13 days from first to last fall, visible berry development 10-16 mm diameter representing 64-76% fruit set, no ongoing cap fall activity over multiple observations confirming cessation, under cool maritime conditions of 20-24°C temperature, typical of this early-bursting moderately-to-highly fruitful cultivar definitively completing flowering and transitioning to berry development focus
Cap fall completion represents the definitive cessation of calyptra detachment activity when all viable flowers have completed abscission establishing final cap shedding extent and marking observable flowering conclusion confirming reproductive phase termination and full transition to berry development. This conclusive observable milestone occurs during complete anthesis in late June-early July in temperate regions Year N+1 immediately following last caps falling, representing point when no further cap fall activity occurs or is possible, corresponding to BBCH 69 complete flowering definition. Completion confirmation requires verification through multiple indicators including cessation of cap fall activity with no ongoing detachment observable over observation period, all developmentally capable flowers showing detached caps with only aborted or aberrant buds remaining unopened, and cluster appearance showing predominant berry development with no significant residual flowering activity. Cap shedding cessation shows definitive end of detachment activity with no further viable flowers remaining capable of shedding caps, occasional persistent unopened structures representing non-viable aborted buds incapable of normal development rather than delayed flowers, and overall completion of flowering-related cap fall activity. Complete shedding achievement shows 100% of viable flowers having completed cap detachment creating maximum possible cap fall for cluster, total cap accumulation representing full flowering extent with no further additions forthcoming, and definitive establishment of final flowering scope. Cluster appearance at completion shows clear developmental state with visible developing berries 10-18 mm diameter throughout cluster creating young fruit cluster impression, senescing brown unfertilized flowers interspersed showing progressive abscission, minimal or no recently opened flowers with residual reproductive activity, and overall berry-dominated appearance confirming developmental transition. Berry development dominance at completion shows fruit structures as primary cluster feature with successful fruit set berries distributed throughout cluster providing visual confirmation of reproductive outcomes, berry number and spatial distribution clearly assessable enabling fruit set evaluation, and berry growth visibly progressing with daily size increases evident. Unfertilized flower abscission progression at completion shows active ongoing elimination of senescing flowers with brown failed reproductions developing completed abscission zones, daily flower drop creating accumulation though less conspicuous than earlier cap fall, and progressive cluster cleaning toward berry-only appearance over subsequent 1-2 weeks. Retained cap assessment at completion enables evaluation of shedding efficiency through identification of incompletely shed caps indicating abscission problems, occasional retention representing coulure causing reproductive failure, and overall retention frequency indicating flowering success and cultivar or environmental limitations. Observational characteristics enabling completion identification include sustained absence of cap fall activity over multiple observation periods confirming cessation, cluster appearance showing predominantly berries with minimal residual flowering, and overall visual evidence of flowering conclusion with developmental transition. Phenological confirmation at completion establishes definitive BBCH 69 complete flowering stage achievement, marks transition from reproductive BBCH 60-69 to developmental BBCH 71-79 stages, and provides clear observable reference easier to assess than detailed reproductive organ examination. Developmental transition confirmation at completion shows physiological commitment to berry development with all resources directing to expanding fruit, no residual reproductive investment or potential remaining, and complete metabolic reorientation from reproductive to developmental focus. Resource allocation at completion shows full reallocation to berry growth as dominant sink consuming 45-65% total photosynthate, vegetative growth continuing requiring 30-40% resources, and zero reproductive demands with flowering completely concluded. Berry growth acceleration following completion shows intensifying expansion through cell division phase, daily diameter increases of 0.5-0.9 mm during peak early growth creating rapid visible progression, and substantial resource demands requiring adequate photosynthetic capacity. Management implications at completion include confirming flowering conclusion enabling complete resumption of all normal operations without bloom restrictions, initiating comprehensive post-bloom management including fungicide applications for berry protection and canopy operations optimizing development, transitioning focus entirely to developmental management and quality optimization, and planning subsequent operations based on berry development progression. Fruit set assessment at completion enables accurate determination through berry counting establishing final berry number per cluster, calculation of fruit set percentage comparing berry number to total flowers opened, and yield prediction multiplying berry number by expected final berry weight providing season potential estimate. Yield prediction based on completion assessment combines berry number from fruit set evaluation with expected berry weight from cultivar norms and environmental conditions, enables comparison to historical norms and targets assessing season outcomes, and informs crop management decisions including possible thinning if load excessive or acceptance if appropriate. Flowering duration documentation at completion enables calculation of total flowering period from first cap detachment to completion typically 10-14 days, assessment of flowering compression versus extension affecting potential uniformity, and phenological characterization supporting management planning and climate studies. Flowering efficiency evaluation at completion considers proportion of flowers achieving successful cap shedding and fruit set, identifies retention problems indicating coulure issues, and assesses overall reproductive success determining yield outcomes. The process represents second and final aspect of final cap fall confirming definitive cessation, following last caps falling. Duration represents threshold point marking completion, with observation confirming sustained absence of activity. Cultivar variation in completion characteristics shows timing differences with early varieties completing late June while late types complete early-mid July, completion clarity with some showing definitive conclusion while others showing gradual extended completion, and retention problems with some varieties showing occasional persistent caps indicating incomplete flowering. Regional timing shows cap fall completion during late June in warm climates at approximately 1150-1200 growing degree days base 10°C, late June-early July in temperate regions at 1050-1100 GDD, and early-mid July in cool areas at 950-1000 GDD, marking flowering conclusion across regions. Viticultural significance includes marking definitive observable flowering completion enabling complete management transitions, confirming final fruit set and berry number establishing yield potential for season, indicating end of critical weather-vulnerable flowering period reducing risk, providing clear practical reference for phenological tracking and management planning, and establishing baseline for subsequent berry development progression and timing predictions. Research applications include documenting precise flowering completion timing for phenological studies, analyzing factors affecting complete flowering achievement and efficiency, evaluating observable markers for flowering conclusion enabling practical phenological assessment, quantifying relationships between cap fall patterns and fruit set outcomes, and assessing flowering duration and uniformity implications for berry development and quality.
Cap_Fall_Completion is second and final aspect of Final_Cap_Fall, distinguished by definitive cessation confirmation. Represents observable flowering conclusion marker. Corresponds to BBCH 69 complete flowering. Enables fruit set assessment and yield prediction. Marks full transition to developmental phase. Provides clear practical observable reference easier than detailed reproductive examination. Establishes baseline for subsequent development predictions. Critical phenological milestone in annual progression. Completes entire Cap_Fall hierarchical documentation from Initial through Mass, Progressive, to Final phases.
Abwurfabschluss
Blüteende-Marker
Kappenfall-Vollendung
calyptra fall conclusion
cap detachment termination
complete cap shedding
flowering completion marker
vollständige Kalyptralösung
Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 393-398
Cap Fall Completion
Carbon balance assessment in heavily cropped versus crop-thinned Cabernet Sauvignon reveals the crop load effect on vine carbon economy, with heavily cropped vines (12 kg/vine) showing late-season carbon balance approaching zero (photosynthetic gain barely exceeding respiratory losses plus fruit carbon demand) while thinned vines (6 kg/vine) maintain positive balance throughout ripening, with the carbon limitation in heavy-crop vines reflected in incomplete ripening (2.5 Brix lower), reduced reserve accumulation (35% lower starch in roots), and compromised following-season bud fruitfulness, demonstrating how carbon balance integrates the factors determining sustainable quality production.
Carbon balance quality is a physiological quality that represents the net result of carbon acquisition through photosynthesis minus carbon losses through respiration and other processes, characterizing the overall carbon economy of the grapevine that determines growth capacity, fruit development potential, and reserve accumulation for sustained productivity across seasons. Within the BFO framework, this quality addresses the integrative carbon accounting dimension of vine physiology, distinguishing the net balance from component fluxes while recognizing that carbon balance provides the ultimate measure of physiological success that determines whether vines can sustain growth, produce quality fruit, and maintain the reserves necessary for perennial productivity. The ontological significance lies in carbon balance determining the fundamental resource availability for all vine functions, where positive carbon balance enables growth and reproduction while negative balance requires mobilization of reserves that, if prolonged, leads to vine decline, making carbon balance the central integrating variable of vine physiology that all management practices ultimately affect. The components of carbon balance include gross photosynthetic carbon gain (total CO₂ fixation), respiratory carbon losses (maintenance, growth, and photorespiration), non-respiratory carbon losses (volatile emissions, root exudation), and the allocation of net carbon gain among competing sinks (shoots, roots, fruit, reserves), with the balance among these fluxes determining vine carbon status. The temporal dynamics of carbon balance show strong diurnal variation (positive during day, negative at night), seasonal patterns (peak positive balance at mid-season, declining toward harvest), and interannual variation depending on weather, crop load, and vine condition, with integration across these time scales determining cumulative carbon status. The measurement and modeling of carbon balance employs gas exchange integration over time, growth analysis calculating carbon increment, or whole-plant chamber measurements of net carbon exchange, with the approach depending on the scale and resolution required. Understanding carbon balance quality enables assessment of vine productive capacity, diagnosis of carbon limitation affecting fruit quality, and optimization of management practices to maintain the positive carbon balance required for sustainable quality production.
Carbon_Balance_Quality serves as the parent class for specific carbon balance components (Gross Gain, Net Balance, Use Efficiency). This quality provides the integrative measure of vine carbon economy that determines productive capacity. The class supports management optimization and sustainability assessment through comprehensive carbon balance evaluation.
C balance
C-Ökonomie
Kohlenstoffbilanz
Kohlenstoffhaushalt
Netto-C-Gewinn
carbon budget
carbon economy
net carbon gain
Lakso AN, Poni S (2005) "Viticulture-Enology 2020": linking carbohydrate supply and demand as a basis for grapevine yield and quality management. Acta Horticulturae 689: 331-340 Palliotti A et al. (2011) Changes in vineyard establishment and canopy management urged by earlier climate-related grape ripening. Australian Journal of Grape and Wine Research 17(2): 72-78
Carbon Balance Quality
Carbon fixation efficiency assessment in Cabernet Sauvignon comparing well-watered versus moderately stressed vines reveals the water status effect on carboxylation performance, with well-watered vines showing Vcmax of 105 μmol/m²/s and net assimilation of 14.5 μmol CO₂/m²/s, while stressed vines display reduced Vcmax of 72 μmol/m²/s and assimilation of 8.8 μmol CO₂/m²/s, with the 31% reduction in carboxylation capacity reflecting both reduced CO₂ supply (stomatal closure) and metabolic downregulation that together diminish carbon fixation efficiency under drought conditions.
Carbon fixation efficiency quality is a photosynthetic capacity quality that represents the effectiveness of converting absorbed light energy and atmospheric CO₂ into organic carbon compounds through the Calvin-Benson cycle, characterizing the biochemical performance of the carbon reduction pathway that determines the rate of carbohydrate synthesis available for vine growth, fruit development, and reserve accumulation. Within the BFO framework, this quality addresses the dark reaction component of photosynthesis, distinguishing carbon fixation from light capture while recognizing that efficient carbon fixation is essential for translating absorbed light energy into the chemical energy of sugars that supports all subsequent vine metabolism and grape quality development. The ontological significance lies in carbon fixation efficiency determining the net productivity of the grapevine, where the rate and efficiency of CO₂ assimilation establishes the carbohydrate supply that drives yield formation, sugar accumulation in berries, secondary metabolite synthesis, and reserve storage for subsequent season vigor, making carbon fixation efficiency a fundamental determinant of sustainable quality production. The biochemical basis for carbon fixation efficiency includes Rubisco enzyme kinetics (carboxylation rate and CO₂/O₂ specificity), Calvin cycle enzyme activities (regeneration of RuBP), CO₂ supply to chloroplasts (stomatal and mesophyll conductances), and the balance between carboxylation and oxygenation (photorespiratory losses), with all components contributing to the net rate of carbon gain per unit absorbed light or leaf area. The measurement approaches for carbon fixation efficiency include gas exchange analysis determining net CO₂ assimilation rate (A), photosynthesis-CO₂ response curves (A-Ci curves) characterizing carboxylation capacity (Vcmax) and electron transport capacity (Jmax), and isotope discrimination analysis indicating the balance between CO₂ supply and demand during photosynthesis. Understanding carbon fixation efficiency quality enables assessment of photosynthetic performance beyond light capture, diagnosis of biochemical limitations on carbon gain, and optimization of conditions that maximize the conversion of light into grape quality potential.
Carbon_Fixation_Efficiency_Quality serves as the parent class for specific carbon assimilation parameters (Net/Gross Photosynthesis Rate, Quantum Yield, Carboxylation Efficiency). This quality addresses the biochemical carbon reduction component distinct from light capture. The class supports productivity assessment and diagnosis of factors limiting the conversion of absorbed light into carbohydrate.
CO₂ assimilation efficiency
CO₂-Assimilationseffizienz
Calvin cycle efficiency
Calvin-Zyklus-Effizienz
Carboxylierungsleistung
Kohlenstofffixierungseffizienz
carbon reduction efficiency
carboxylation performance
Flexas J et al. (2012) Mesophyll conductance to CO₂ and Rubisco as targets for improving intrinsic water use efficiency in C₃ plants. Plant, Cell & Environment 35(2): 348-365 Sharkey TD et al. (2007) Fitting photosynthetic carbon dioxide response curves for C₃ leaves. Plant, Cell & Environment 30(9): 1035-1040
Carbon Fixation Efficiency Quality
Carbon isotope discrimination (Δ¹³C) in grapevine leaf dry matter ranges from 18-24‰ across cultivars and water regimes, with higher discrimination indicating lower water use efficiency. Well-watered vines with high stomatal conductance show Δ¹³C of 22-24‰, while water-stressed vines with restricted stomata show 18-21‰. Cultivar differences of 1-3‰ in Δ¹³C persist across environments, with Grenache showing consistently higher discrimination (lower WUE) than Tempranillo. Berry sugar Δ¹³C of 20-26‰ integrates vine water status over the ripening period and correlates with seasonal water deficit (r = -0.65 to -0.80). δ¹³C values in wine range from -22 to -28‰ (expressed relative to PDB standard), enabling retrospective assessment of vineyard water status.
A water use efficiency disposition that is the inherited capacity of a Vitis vinifera plant to discriminate against the heavier stable carbon isotope (¹³C) relative to the lighter isotope (¹²C) during photosynthetic CO₂ fixation, providing an integrated measure of the ratio of intercellular to atmospheric CO₂ concentration (Ci/Ca) and thus a time-integrated proxy for intrinsic water use efficiency.
Carbon isotope discrimination provides a powerful tool for phenotyping water use efficiency because the isotopic signature integrates gas exchange over the period of tissue formation, avoiding the instantaneous measurement challenges of leaf-level WUE. Lower discrimination (more negative δ¹³C, less positive Δ¹³C) indicates higher WUE through reduced Ci/Ca ratio resulting from stomatal limitation. The trait is used in breeding programs to screen for WUE, in production to assess irrigation adequacy, and in wine authentication to verify origin and vintage conditions.
C-Isotopenverhältnis
Kohlenstoff-Isotopendiskriminierung
carbon isotope ratio
delta 13C trait
Δ¹³C disposition
δ¹³C-Signatur
Gaudillère JP et al. (2002) Carbon isotope composition of sugars in grapevine, an integrated indicator of vineyard water status. Journal of Experimental Botany 53:757-763 Santesteban LG et al. (2015) Discrimination ability of leaf and stem water potential at different times of the day through a meta-analysis in grapevine (Vitis vinifera L.). Agricultural Water Management 158:52-59
Carbon Isotope Discrimination Disposition
Nerello Mascalese vines on the north slope of Mount Etna bearing a carbon sequestration role accumulate 1.5-3.0 t C/ha/year in permanent woody structures (trunk, cordons, root system) over a 50+ year lifespan, with associated soil organic carbon stocks of 40-70 t C/ha in the top 30 cm under traditional alberello training on volcanic soils, annual leaf and pruning residue inputs of 2-4 t dry matter/ha contributing 0.8-1.5 t C/ha/year to decomposition and humus formation, and total vineyard ecosystem carbon stocks estimated at 80-120 t C/ha representing significant carbon reservoirs comparable to some temperate forest systems.
An ecological role that inheres in a grapevine by virtue of its contribution to atmospheric carbon dioxide removal and long-term carbon storage through photosynthetic assimilation into woody biomass, root organic matter, and associated soil organic carbon pools within the vineyard ecosystem.
Carbon_Sequestration_Role addresses the increasingly recognized function of vineyards as carbon sinks within agricultural landscapes. Grapevines as perennial woody crops accumulate substantial carbon in permanent structures over their 25-100+ year lifespans, distinguishing them from annual crops that release sequestered carbon at harvest: however, net carbon balance depends critically on management practices including tillage (releasing soil C), fossil fuel inputs, and cover crop presence. Life cycle analyses suggest that well-managed vineyards with cover crops and minimal tillage can function as net carbon sinks, while intensively tilled, bare-soil vineyards may be net carbon sources.
CO₂-Senken-Rolle
Kohlenstoffbindungs-Rolle
Kohlenstoffspeicher-Rolle
carbon sink role
carbon storage role
vineyard carbon sequestration role
Brunori E et al. (2016) Carbon balance and life cycle assessment in an oak vineyard, Journal of Cleaner Production 112: 2569-2580 Wolff MW et al. (2018) Carbon sequestration in California vineyards, Soil Science Society of America Journal 82: 1422-1430 Morandé JA et al. (2017) Carbon dioxide, water, and energy fluxes of a vineyard, Agricultural and Forest Meteorology 240: 67-79
Carbon Sequestration Role
Carbon use efficiency comparison between cool-climate (Tasmania, mean growing season temperature 15°C) and warm-climate (Riverland, 22°C) Pinot Noir reveals the temperature effect on metabolic efficiency, with cool-climate vines achieving CUE of 0.58 (net retention of 58% of gross photosynthesis) while warm-climate vines show CUE of 0.44 (44% retention), with the 14 percentage point difference arising primarily from elevated respiratory losses at higher temperatures (Q₁₀ effect), partially offset by higher absolute gross photosynthesis in the warm climate, demonstrating how temperature affects not just absolute productivity but the efficiency of converting photosynthate into vine growth and fruit.
Carbon use efficiency quality is a carbon balance quality that represents the fraction of gross carbon gain that is retained as net carbon balance, characterizing the efficiency with which the grapevine converts photosynthetically fixed carbon into structural and storage compounds rather than losing it through respiration. Within the BFO framework, this quality specifically addresses the efficiency dimension of carbon metabolism, distinguishing the fraction retained from absolute carbon fluxes while recognizing that carbon use efficiency indicates how effectively the vine converts its photosynthetic production into useful structural and reproductive output. The ontological significance lies in carbon use efficiency providing a measure of metabolic efficiency that complements absolute productivity measures, where high CUE indicates that the vine is effectively converting fixed carbon into growth and reproduction with minimal respiratory loss, while low CUE indicates that a large fraction of fixed carbon is consumed by respiration, limiting the conversion of photosynthetic capacity into vine performance regardless of canopy size or photosynthetic rate. The calculation of carbon use efficiency divides net carbon balance by gross carbon gain, with CUE = Net Balance / Gross Gain = (Gross - Respiration) / Gross, yielding values typically ranging from 0.40-0.60 in grapevines depending on temperature, growth rate, and tissue composition. The factors affecting carbon use efficiency include temperature (higher temperatures increasing respiratory losses and reducing CUE), growth rate (actively growing tissues having higher biosynthetic efficiency than maintenance-dominated tissues), tissue composition (protein-rich tissues having higher construction and maintenance costs), and the balance between productive canopy and respiratory burden (excessive non-productive structure reducing CUE). The management implications of carbon use efficiency include the recognition that simply maximizing canopy size may not optimize productivity if additional leaf area adds more to respiratory burden than to photosynthetic gain, and that maintaining vine health and moderate growth rates may achieve higher CUE than pushing maximum vegetative growth. Understanding carbon use efficiency quality enables evaluation of the metabolic efficiency of vine carbon utilization, diagnosis of conditions where respiratory losses limit conversion of photosynthetic capacity into productivity, and optimization of management practices that maintain high CUE throughout the growing season.
Carbon_Use_Efficiency_Quality addresses the fractional retention of fixed carbon as a metabolic efficiency measure. This quality indicates how effectively photosynthetic production converts to vine performance. The class supports efficiency optimization and climate impact assessment through CUE evaluation revealing the respiratory burden on productivity.
C-Konversionseffizienz
CUE
Kohlenstoffnutzungseffizienz
Wachstumseffizienz
carbon conversion efficiency
growth efficiency
respiratory burden
DeLucia EH et al. (2007) Forest carbon use efficiency: is respiration a constant fraction of gross primary production? Global Change Biology 13(6): 1157-1167 Gifford RM (2003) Plant respiration in productivity models: conceptualisation, representation and issues for global terrestrial carbon-cycle research. Functional Plant Biology 30(2): 171-186
Carbon Use Efficiency Quality
Carboxylation efficiency assessment from A-Ci curves in nitrogen-sufficient versus deficient Pinot Noir reveals the nutrition effect on Rubisco capacity, with sufficient vines showing carboxylation efficiency of 0.095 mol/m²/s and Vcmax of 98 μmol/m²/s while deficient vines display reduced CE of 0.052 mol/m²/s and Vcmax of 54 μmol/m²/s, with the 45% reduction in carboxylation capacity reflecting lower Rubisco content from limited nitrogen availability, demonstrating how nitrogen nutrition directly affects the biochemical capacity for carbon fixation.
Carboxylation efficiency quality is a carbon fixation efficiency quality that represents the initial slope of the photosynthetic CO₂ response curve reflecting Rubisco carboxylation capacity at limiting CO₂ concentrations, characterizing the biochemical efficiency of the rate-limiting enzyme that determines carbon fixation rate under conditions where CO₂ supply rather than light limits photosynthesis. Within the BFO framework, this quality specifically addresses the Rubisco-limited phase of carbon fixation, distinguishing carboxylation efficiency from electron transport-limited capacity while recognizing that carboxylation efficiency determines photosynthetic rate under the stomatal closure conditions typical of water-stressed grapevines where internal CO₂ concentration becomes limiting. The ontological significance lies in carboxylation efficiency providing the key biochemical parameter for predicting photosynthesis under stress conditions, where stomatal closure reduces internal CO₂ and shifts photosynthesis into the Rubisco-limited regime where carboxylation efficiency determines carbon gain, making this parameter essential for understanding and modeling stressed vine productivity. The biochemical basis for carboxylation efficiency involves Rubisco kinetics, with the initial slope of A-Ci curves (dA/dCi at low Ci) reflecting Rubisco content, activation state, and kinetic properties (Vcmax, Km for CO₂, specificity factor), and with carboxylation efficiency (CE) typically expressed as the slope of A versus Ci below the inflection point (approximately 200 μmol/mol Ci). The quantitative ranges for carboxylation efficiency in Vitis vinifera typically span 0.05-0.12 mol/m²/s when expressed as dA/dCi, with variation reflecting Rubisco content (nitrogen nutrition dependent), activation state (regulated by Rubisco activase), and temperature effects on kinetic parameters. The stress responses of carboxylation efficiency include both short-term regulation (Rubisco activation state, carbamylation) and longer-term acclimation (Rubisco content adjustment), with drought and heat stress typically reducing carboxylation efficiency through multiple mechanisms. Understanding carboxylation efficiency quality enables assessment of Rubisco-limited photosynthetic capacity, prediction of carbon gain under stress conditions, and diagnosis of biochemical versus diffusional limitations on photosynthesis.
Carboxylation_Efficiency_Quality addresses the Rubisco-limited phase of photosynthesis determined from A-Ci curve analysis. This quality becomes the primary limitation under stress conditions causing stomatal closure. The class supports biochemical capacity assessment and stress response diagnosis through carboxylation efficiency measurement.
CE-Wert
Carboxylierungseffizienz
Rubisco efficiency
Rubisco-Effizienz
Vcmax proxy
initial slope A-Ci
initiale Steigung
Sharkey TD et al. (2007) Fitting photosynthetic carbon dioxide response curves for C₃ leaves. Plant, Cell & Environment 30(9): 1035-1040 Flexas J et al. (2012) Mesophyll conductance to CO₂ and Rubisco as targets for improving intrinsic water use efficiency in C₃ plants. Plant, Cell & Environment 35(2): 348-365
Carboxylation Efficiency Quality
Carotenoid content assessment in heat-stressed versus control Shiraz leaves reveals the photoprotective response, with control leaves showing carotenoid content of 120 μmol/m² with violaxanthin dominant in the xanthophyll pool, while heat-stressed leaves display elevated carotenoids (145 μmol/m²) with substantial zeaxanthin accumulation indicating xanthophyll cycle engagement for thermal energy dissipation, demonstrating how carotenoid content and composition indicate stress status and photoprotective capacity, with implications for both immediate photosynthetic function and downstream norisoprenoid precursor availability for wine aroma development.
Carotenoid content quality is a light capture efficiency quality that represents the concentration of carotenoid pigments in grapevine leaves, characterizing the accessory pigment pool that contributes to light harvesting in the blue-green spectral region, provides essential photoprotection through excess energy dissipation, and serves as precursor for important aroma compounds in grape berries. Within the BFO framework, this quality specifically addresses the carotenoid pigment dimension of light capture, distinguishing carotenoid function from chlorophyll while recognizing the dual roles of carotenoids in both productive light harvesting and protective energy dissipation depending on light conditions and plant stress status. The ontological significance lies in carotenoid content serving multiple functional roles beyond simple light capture, where carotenoids act as essential photoprotective agents preventing oxidative damage under excess light conditions through the xanthophyll cycle, while also serving as precursors for C₁₃-norisoprenoid aroma compounds (β-damascenone, β-ionone, TDN) that contribute to varietal character in wines, linking leaf physiology to fruit and wine quality. The biochemical basis for carotenoid function includes light harvesting (absorption in 400-500 nm blue region, energy transfer to chlorophyll), photoprotection (xanthophyll cycle converting violaxanthin to zeaxanthin under excess light, enabling thermal dissipation), antioxidant activity (quenching reactive oxygen species and triplet chlorophyll), and structural roles (stabilizing light-harvesting complexes), with the total carotenoid pool comprising both xanthophylls (oxygenated carotenoids) and carotenes (pure hydrocarbons). The environmental and developmental factors affecting carotenoid content include light intensity (high light increasing photoprotective carotenoids), temperature stress (affecting xanthophyll cycle activity), water status (drought enhancing carotenoid accumulation), nitrogen nutrition (affecting synthesis rates), and leaf age (carotenoids declining more slowly than chlorophylls during senescence, creating autumn colors). Understanding carotenoid content quality enables photoprotection status assessment, stress tolerance evaluation, and recognition of the leaf-to-fruit connection for norisoprenoid aroma precursors affecting wine quality.
Carotenoid_Content_Quality addresses the accessory pigment pool with dual light-harvesting and photoprotective functions. This quality links leaf physiology to wine aroma through norisoprenoid precursor roles. The class supports stress assessment, photoprotection evaluation, and understanding of the physiological basis for aroma precursor accumulation.
Carotinoidgehalt
Schutzpigmentpool
Xanthophyllgehalt
accessory pigment level
akzessorische Pigmente
carotenoid concentration
photoprotective pigment pool
xanthophyll content
Demmig-Adams B, Adams WW (1996) The role of xanthophyll cycle carotenoids in the protection of photosynthesis. Trends in Plant Science 1(1): 21-26 Mendes-Pinto MM (2009) Carotenoid breakdown products the—norisoprenoids—in wine aroma. Archives of Biochemistry and Biophysics 483(2): 236-245
Carotenoid Content Quality
Catechin content quality in Tannat seeds from Madiran reaches exceptionally high levels (35-55 mg/g seed dry weight) reflecting this variety's extreme tannic character, with the catechin serving as building block for the extensive proanthocyanidin polymers that produce Tannat's characteristic firm, gripping astringency traditional Madiran winemaking employs extended maceration (30-45 days) with periodic délestage to maximize catechin-based tannin extraction while the polymerization occurring during two years minimum oak aging softens the initially harsh mouthfeel to approachable roundness.
Catechin_Content_Quality represents the chemical quality characterizing the concentration of (+)-catechin in grape berry tissues, capturing the primary flavan-3-ol monomer with 2,3-trans stereochemistry. (+)-Catechin (2R,3S-flavan-3-ol) is distinguished from its stereoisomer (−)-epicatechin by the trans relationship between the C2 and C3 substituents, resulting in different three-dimensional geometry that affects both polymerization behavior and sensory properties. Catechin is the more abundant flavan-3-ol monomer in grape seeds, typically comprising 30-50% of total seed flavan-3-ols, and serves as a primary building block for condensed tannin polymers through C4-C6 and C4-C8 linkages. The concentration of free catechin in seeds decreases during berry development as it becomes incorporated into growing polymer chains, with monomeric catechin representing a diminishing proportion of total flavan-3-ol content as berries mature. Catechin contributes astringent and bitter taste qualities when present as free monomer, with sensory thresholds around 20-50 mg/L in wine matrices. The dihydroxylated B-ring of catechin (3',4'-dihydroxy pattern) reflects synthesis through the F3'H pathway, parallel to cyanidin among anthocyanins. Catechin serves as the terminal unit in many proanthocyanidin chains, occupying the lower position formed through electrophilic attack by the upper extension unit. The ratio of catechin to epicatechin in seeds and wines provides information about condensed tannin structure and potential sensory properties.
Catechin_Content_Quality specifically addresses the (+)-catechin stereoisomer with 2,3-trans configuration. The distinction from (−)-epicatechin (2,3-cis) is important because stereochemistry affects polymer structure, extraction kinetics, and sensory properties. Catechin serves primarily as a chain terminal unit in proanthocyanidins, while epicatechin more commonly functions as an extension unit. The ratio of terminal to extension units (catechin to epicatechin) correlates with polymer size and affects tannin sensory properties. Free catechin quantification employs normal-phase or reversed-phase HPLC with UV detection at 280 nm.
(+)-Catechin-Konzentration
(+)-catechin content
2,3-trans-Flavan-3-ol-Gehalt
2,3-trans-flavan-3-ol concentration
Catechingehalt
catechin monomer quality
free catechin content
freier Catechingehalt
Kennedy JA et al. (2000) Composition of Grape Skin Proanthocyanidins at Different Stages of Berry Development. Journal of Agricultural and Food Chemistry 48(12): 5348-5355 Prieur C et al. (1994) Oligomeric and Polymeric Procyanidins from Grape Seeds. Phytochemistry 36(3): 781-784
Catechin Content Quality
Cation exchange quality assessment guides cold stabilization protocols in German Riesling production, with cation analysis revealing the potassium-dominated (1200-1800 mg/L K⁺, 80-120 mg/L Ca²⁺) must composition that predicts rapid potassium bitartrate precipitation during cold treatment (−4°C for 7-14 days) the cation exchange approach enables calculation of KHT saturation temperature and optimization of stabilization duration, preventing both under-treatment (continued precipitation in bottle) and over-treatment (unnecessary quality loss from extended cold exposure) that waste energy and potentially strip wine of desirable complexity.
Cation_Exchange_Quality represents the chemical quality characterizing the overall balance and exchange relationships among cationic mineral species in grape must and wine, capturing the equilibrium chemistry that affects pH, tartrate stability, and protein interactions. The cation exchange concept recognizes that positively charged mineral ions (K⁺, Ca²⁺, Mg²⁺, Na⁺) compete for association with negatively charged organic acids (tartrate, malate) and proteins, with the exchange equilibria determining the proportion of free acid versus salt forms that fundamentally affect acidity expression. Potassium dominates cation content in grape must, but its effect on pH depends on competition with hydrogen ions for association with tartrate, a competition governed by pH, temperature, and the relative concentrations of all cationic species. The formation of potassium bitartrate (KHT) removes both potassium and tartrate from solution, with precipitation affected by temperature, ethanol concentration, and the presence of competing cations and nucleation inhibitors. Calcium competes with potassium for tartrate binding, forming the less soluble calcium tartrate that precipitates more slowly but over longer time periods. The cation exchange capacity of wines, measured through titration or conductivity methods, indicates the total exchangeable cation content available to participate in stability reactions and pH buffering. In wine stability management, understanding cation exchange enables prediction of tartrate precipitation, optimization of cold stabilization protocols, and application of ion exchange treatment for pH adjustment. The cation composition also affects protein stability, as cations interact with proteins and fining agents during clarification treatments.
Cation_Exchange_Quality addresses the equilibrium chemistry among cationic species rather than individual element concentrations. The concept derives from soil science (cation exchange capacity) but applies equally to juice and wine chemistry. The pH effects of cation exchange reflect the competition between hydrogen ions and metal cations for organic acid association, making cation balance a determinant of acidity expression beyond titratable acid concentration. Cold stability, protein stability, and pH adjustment treatments all involve cation exchange reactions, making this quality relevant to multiple winemaking interventions.
Ionengleichgewicht
Kationenaustauschqualität
austauschbarer Kationengehalt
cation balance quality
cationic composition quality
exchangeable cation content
ion exchange equilibrium
kationische Zusammensetzung
Boulton R (1980) The Relationship Between Total Acidity, Titratable Acidity and pH in Grape Tissue. Vitis 19(2): 113-120 Berg HW and Keefer RM (1958) Analytical Determination of Tartrate Stability in Wine. I. Potassium Bitartrate. American Journal of Enology and Viticulture 9(4): 180-193
Cation Exchange Quality
Systematic determination of Cavitation_Resistance_Disposition at the University of Bordeaux employed the Cavitron centrifuge technique to generate vulnerability curves for 25 cultivars, providing high-throughput, standardized assessment of P50 values. Stem segments were subjected to increasing centrifugal tension while conductivity was monitored, enabling rapid construction of complete vulnerability curves. Cavitation_Resistance_Disposition showed significant cultivar variation spanning 1.3 MPa. Most resistant cultivars included 'Grenache' (P50 = -2.85 MPa), 'Mourvèdre' (P50 = -2.72 MPa), and 'Carignan' (P50 = -2.65 MPa), all Mediterranean cultivars presumably selected under drought pressure. Most vulnerable cultivars included 'Merlot' (P50 = -1.58 MPa), 'Pinot Noir' (P50 = -1.65 MPa), and 'Gamay' (P50 = -1.72 MPa), originating from cooler, more humid climates. Anatomical analysis revealed that P50 correlated strongly with pit membrane thickness (r = -0.78) and weakly with vessel diameter (r = 0.35), confirming pit membrane properties as the primary determinant of Cavitation_Resistance_Disposition. Field observations during the severe 2003 European drought validated laboratory rankings, with vulnerable 'Merlot' vines showing 45-60% PLC and visible canopy dieback while resistant 'Grenache' maintained PLC below 20% despite similar exposure. The relationship between Cavitation_Resistance_Disposition and stomatal behavior was confirmed, with cavitation-resistant cultivars operating closer to their P50 (narrower safety margins) enabled by tight stomatal control, while vulnerable cultivars maintained wider margins through earlier stomatal closure.
A Cavitation_Resistance_Disposition is a Hydraulic_Architecture_Disposition that inheres in the xylem of a Vitis vinifera plant and is realized through structural properties that resist the formation and spread of embolism (air bubbles blocking water transport) under the tension developed in xylem during drought. This disposition represents the safety component of xylem hydraulic function, determining the water potential threshold at which catastrophic loss of hydraulic conductivity occurs. Cavitation (the phase change from liquid water to water vapor) occurs when xylem tension exceeds the capacity of water columns to withstand negative pressure, typically initiated by air-seeding through porous pit membranes connecting adjacent vessels. Resistance to cavitation depends primarily on pit membrane properties (thickness, porosity, flexibility) that determine the air-seeding pressure threshold, with thicker, less porous membranes providing greater resistance. Secondary factors include vessel diameter (larger vessels contain more water under tension and may have more vulnerable pit connections), vessel length (longer vessels have more pit area for potential air entry), and inter-vessel connectivity. Cavitation_Resistance_Disposition is quantified by vulnerability curves relating xylem water potential to percentage loss of conductivity (PLC), with P50 (water potential at 50% PLC) as the standard metric. Grapevine P50 values typically range from -1.5 to -3.0 MPa depending on genotype, with more negative values indicating greater cavitation resistance.
Cavitation_Resistance_Disposition is_a Hydraulic_Architecture_Disposition. Cavitation resistance determines the water potential threshold for hydraulic failure. P50 (water potential at 50% conductivity loss) is the standard metric. Pit membrane properties (thickness, porosity) are primary determinants. Vessel diameter has secondary influence. Mediterranean-origin cultivars generally show greater resistance than cool-climate cultivars. Cavitation resistance interacts with stomatal behavior to determine hydraulic safety margins.
Embolie-Resistenz
Hydraulische-Sicherheit
Kavitations-Resistenz-Disposition
embolism resistance
hydraulic safety
xylem cavitation resistance
Choat B et al. (2010) Measurement of vulnerability to water stress-induced cavitation in grapevine: a comparison of four techniques applied to a long-vesseled species, Plant Cell and Environment 33:1502-1512 Tyree MT, Sperry JS (1989) Vulnerability of xylem to cavitation and embolism, Annual Review of Plant Physiology and Plant Molecular Biology 40:19-38 Brodersen CR et al. (2010) The dynamics of embolism repair in xylem: in vivo visualizations using high-resolution computed tomography, Plant Physiology 154:1088-1095
Cavitation Resistance Disposition
Cavitation resistance comparison between own-rooted and grafted Tempranillo reveals rootstock effects on hydraulic safety, with own-rooted vines showing moderate resistance (P50 = -1.7 MPa, relatively vulnerable) while the same cultivar on 140Ru rootstock shows improved whole-plant resistance (effective P50 = -2.2 MPa from root system contribution), with the enhanced cavitation resistance from drought-adapted rootstock enabling operation at more negative water potentials without hydraulic damage, expanding the deficit irrigation range applicable without risking embolism-induced productivity loss.
Cavitation resistance quality is a vulnerability to cavitation quality that represents the capacity of xylem to maintain hydraulic function under water stress conditions, characterizing the robustness of the water transport system against embolism that determines the range of water potentials over which conductivity is preserved and the buffer against drought-induced hydraulic failure. Within the BFO framework, this quality specifically addresses the positive framing of hydraulic safety, distinguishing resistance to dysfunction from the vulnerability perspective while recognizing that cavitation resistance is the inverse of vulnerability and may be more intuitively related to drought tolerance capacity. The ontological significance lies in cavitation resistance determining the operational water potential range for effective water transport, where high resistance enables vines to function at more negative water potentials without conductivity loss, expanding the range of soil water conditions under which transpiration and photosynthesis can be maintained and providing greater tolerance of drought episodes. The anatomical basis for cavitation resistance involves pit membrane properties that determine the pressure required for air entry, with smaller pit membrane pores requiring more negative pressures to overcome surface tension and pull air through, creating the fundamental trade-off between pit conductivity (favoring large pores) and cavitation resistance (favoring small pores) that shapes xylem evolution. The additional factors affecting cavitation resistance include vessel wall strength (resisting implosion under tension), vessel network connectivity (affecting whether embolism spreads or is contained), and pit membrane composition (lignification and other chemical properties affecting porosity and stability). The plasticity of cavitation resistance through acclimation shows that xylem formed under drought conditions often has improved resistance compared to that formed under well-watered conditions, indicating that vines can partially adjust their hydraulic safety margins in response to experienced stress history. Understanding cavitation resistance quality enables assessment of drought tolerance capacity, evaluation of the efficiency-safety trade-off in xylem design, and prediction of the operational water potential range for sustained hydraulic function.
Cavitation_Resistance_Quality addresses the positive capacity to maintain hydraulic function under stress, inverse to vulnerability. This quality determines the operational water potential range for effective transport. The class supports drought tolerance assessment and safety margin evaluation through cavitation resistance characterization.
Embolieresistenz
Kavitationsresistenz
Xylemwiderstandsfähigkeit
drought resistance (hydraulic)
hydraulische Robustheit
xylem robustness
Hacke UG et al. (2006) Scaling of angiosperm xylem structure with safety and efficiency. Tree Physiology 26(6): 689-701 Alsina MM et al. (2007) Seasonal variation of xylem embolism in grapevine cultivars grafted on different rootstocks. Australian Journal of Grape and Wine Research 13(2): 106-114
Cavitation Resistance Quality
Merlot cavity locular tissue at maturity defining air-filled cavity structure= locular tissue lining 2 cavities each 4-5.5 mm diameter, with seeds (2-3 per berry) occupying 32-42% of cavity volume surrounded by air space occupying 58-68% of volume, creating air-filled chambers bounded by cavity locular tissue cellular lining, with 22-38 μm air gaps between seed surfaces and innermost cavity locular tissue cells demonstrating substantial cavity air space development during ripening creating distinct berry internal architecture with central air-filled chambers separated from flesh bulk by cavity locular tissue boundary. Cabernet Sauvignon cavity locular tissue air space development time-course= week 6 (lag phase) showing cavities 85-92% filled by seeds, 8-15% air space, cavity locular tissue closely apposed to seeds week 10 (veraison) showing 65-75% seed occupancy, 25-35% air space, 12-22 μm gaps developing week 14 (mid-ripening) showing 45-58% seed occupancy, 42-55% air space, 20-35 μm gaps week 18 (maturity) showing 35-48% seed occupancy, 52-65% air space, 25-45 μm gaps, demonstrating progressive cavity air space increase during ripening as berry expansion exceeds seed dimensions creating air-filled cavity structure bounded by cavity locular tissue. Syrah cavity locular tissue berry internal structure visualization by cross-sectioning= transverse section through berry middle showing 2 locular cavities positioned left and right of berry center (perpendicular to major dorsal vascular bundles running top-bottom), each cavity containing 1-2 seeds surrounded by air space, cavities bounded by continuous cavity locular tissue 160-240 μm thickness separating air-filled cavity interiors from surrounding mesocarp flesh, creating obvious visual distinction between dark-colored outer skin, light-colored flesh bulk, and internal air-filled chambers in fresh berry cross-section. Thompson Seedless stenospermocarpic cavity locular tissue in seedless cultivar= cavities containing small seed traces (arrested seed development, 2-4 mm length versus 5-7 mm in normal seeds) surrounded by substantial air space (65-80% of cavity volume, higher than seeded cultivars due to smaller seed traces), with cavity locular tissue structure similar to seeded types (140-190 μm thickness, 4-5 cell layers) demonstrating that cavity development and cavity locular tissue formation occur independently of successful seed development in seedless cultivars with arrested seeds still inducing normal locular cavity and cavity locular tissue differentiation. Pinot Noir cavity locular tissue potential role in berry splitting= rapid water uptake during rainfall creating berry volume increase potentially accommodated partially through cavity air space compression (estimated 5-12% volume reduction possible through air space compression before substantial cavity locular tissue or flesh compression required), potentially reducing splitting risk by providing internal volume buffer, though splitting typically initiates at external skin weak points (stylar scar, pedicel attachment, lenticels) rather than at internal cavity locular tissue suggesting cavity compression provides limited splitting protection with skin mechanical properties more critical for splitting resistance than internal cavity air space volume.
A cavity locular tissue is locular tissue emphasizing its role as boundary defining air-filled or partially air-filled cavity spaces within the berry, particularly relevant during ripening and maturity when locular cavities develop substantial air space (typically 50-70% of cavity volume) as berry expansion exceeds seed size increase, with cavity locular tissue providing structural boundary maintaining cavity integrity and creating distinct berry internal architecture with seed-containing air-filled chambers surrounded by fleshy mesocarp. Cavity locular tissue structural boundary function= (1) definition of cavity internal surfaces creating cellular boundary between soft mesocarp tissue and air-filled cavity space (2) structural support preventing cavity collapse during berry expansion and handling despite soft fleshy mesocarp providing minimal mechanical support (3) maintenance of cavity shape and volume during berry growth accommodating progressive air space development as seeds lag behind berry expansion (4) potential mechanical properties including wall strength and cell-cell adhesion maintaining structural integrity under compression and deformation stresses during berry packing in clusters and during harvest handling. Cavity locular tissue in berry internal architecture= (1) creation of distinct berry structure with outer protective skin zone (exocarp/outer pericarp), middle storage flesh zone (mesocarp bulk), inner cavity-defining zone (cavity locular tissue), and central air-filled chambers (locular cavities) rather than homogeneous flesh throughout berry (2) cavity position and size influencing overall berry geometry with large cavities potentially affecting berry shape and with cavity asymmetry potentially contributing to irregular berry shapes (3) cavity air space providing gas reservoir within berry potentially influencing internal atmosphere composition (O₂, CO₂, ethylene concentrations) affecting seed and tissue metabolism (4) cavity surfaces potentially showing different properties than external berry surface including different humidity, temperature buffering, and gas exchange characteristics. Cavity locular tissue air space development during ripening= early development and Phase I showing locular cavities filled primarily by seeds with minimal air space (<10-20% of cavity volume) lag phase showing beginning air space development (20-35% of volume) as seeds complete maturation while berry continues some expansion early ripening (weeks 9-12, early Phase II) showing increasing air space (35-50%) as berry expansion accelerates during ripening late ripening and maturity (weeks 13-18+) showing maximum air space development (50-70% of cavity volume) with seeds surrounded by air gaps 15-50 μm width creating substantial air-filled cavity space bounded by cavity locular tissue lining. Cavity locular tissue in berry texture and eating quality= cavity air space potentially contributing to berry textural perception during eating with air-filled cavities creating less resistance during compression than solid flesh cavity locular tissue potentially contributing to skin-flesh separation during chewing as cavity boundary may represent weak point where separation initiates cavity structure potentially affecting juice release pattern during eating with cavity compression potentially facilitating juice flow from ruptured mesocarp cells seeds within air-filled cavities potentially creating distinct textural element during eating as seeds surrounded by air rather than embedded in flesh allowing seeds to move independently when cavity compressed. Cavity locular tissue in berry splitting susceptibility= cavity air space potentially creating compressible zone absorbing volume changes during rapid water uptake reducing splitting risk by allowing berry expansion into cavity rather than requiring skin stretching alternatively, locular tissue as thinner-walled inner pericarp potentially representing weak zone where internal pressure could initiate cracks propagating outward cavity-locular tissue interface potentially representing location of internal crack initiation during water stress though most berry splitting initiates at external surface rather than internal cavities.
Cavity_Locular_Tissue emphasizes the cavity-boundary and air space aspects of locular tissue and is distinguished from Chamber_Locular_Tissue which may emphasize seed-chamber aspects. Understanding cavity locular tissue is relevant for comprehending berry internal architecture with air-filled cavities as distinct structural feature, for understanding ripening-associated cavity development creating increasing internal air space, and for investigating potential cavity roles in berry mechanical properties, gas exchange, and splitting susceptibility.
Hohlraum-Lokular-Gewebe
Luftkammer-Gewebe
Luftraum-Grenzgewebe
air cavity lining
chamber boundary tissue
Considine, J.A., Kriedemann, P.E. (1972). Fruit splitting in grapes. Australian Journal of Agricultural Research 23':17-27 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 180-185.
Cavity Locular Tissue
The CEC data for a Champagne vineyard on chalk (Cramant, Côte des Blancs) documents a high cation exchange capacity of 25-35 cmol(+)/kg in the upper 30 cm — dominated by exchangeable calcium (85-90% of CEC, reflecting the calcareous parent material), with exchangeable magnesium contributing 5-8%, potassium 2-4%, and sodium <1%, and the base saturation at 100% (typical of calcareous soils where the calcium carbonate buffer maintains complete base saturation at all times). This high CEC provides exceptional chemical buffering — the soil resists pH changes from acid inputs (acid rain, acidifying fertilizers, root-released organic acids) because the large reservoir of exchangeable calcium ions neutralizes proton additions, and the nutrient supply is well-buffered against depletion because the exchange complex releases cations to the soil solution as plant uptake removes them. The high CEC of the Champagne chalk soils contrasts with the low CEC (5-10 cmol(+)/kg) of the acidic granite-derived soils in Beaujolais where the exchange complex is dominated by aluminum and hydrogen ions rather than base cations.
A soil chemistry data component that records the cation exchange capacity (CEC) of the vineyard soil — the total quantity of exchangeable cations (positively charged ions) that the soil can retain per unit mass, expressed in centimoles of positive charge per kilogram of soil (cmol(+)/kg, equivalent to the older unit milliequivalents per 100 g) — measured by standard displacement methods (ammonium acetate at pH 7.0 for effective CEC, or barium chloride-triethanolamine for potential CEC), with CEC being the primary indicator of the soil's capacity to retain and supply essential base cations (calcium, magnesium, potassium, sodium) against leaching losses and to buffer the soil solution chemistry against rapid changes from fertilizer inputs or rainfall acidification.
CEC_Data documents the soil's ion exchange capacity that controls the retention and release dynamics of the essential base cation nutrients — calcium, magnesium, potassium, and (to a lesser extent) sodium — that together constitute the majority of the vine's mineral nutrient uptake by mass. The CEC is determined by two soil components — the clay mineral fraction (contributing CEC proportional to the clay content and the specific surface area of the clay minerals, ranging from 3-15 cmol(+)/kg for kaolinite through 25-40 for illite to 80-120 for smectite) and the organic matter fraction (contributing 150-300 cmol(+)/kg of organic matter, a CEC per unit mass far exceeding that of any clay mineral): the relative contribution of clay versus organic matter to the total CEC varies with depth and soil type — in surface horizons with 3-5% organic matter, the organic fraction may contribute 30-60% of the total CEC despite representing only a small proportion of the soil mass, while in subsoil horizons with <1% organic matter, the clay fraction dominates the CEC. The viticultural significance of CEC centers on the soil's capacity to supply potassium — a nutrient of particular importance in viticulture because potassium accumulation in berries affects must pH (potassium salts of tartaric acid raising pH), wine stability (potassium bitartrate precipitation), and fermentation kinetics — soils with low CEC and low exchangeable potassium may produce vines with potassium deficiency symptoms and low-potassium must (desirable for maintaining low pH in warm-climate wines), while soils with high CEC and high exchangeable potassium may produce excessive berry potassium uptake and elevated must pH (a quality concern in warm, fertile sites).
Basenaustausch-Kapazität
KAK-Daten
Kationenaustauschkapazität
base retention measurement
cation exchange capacity record
ion exchange characterization
White RE (2003) Soils for Fine Wines, Oxford University Press Mpelasoka BS et al. (2003) A review of potassium nutrition in grapevines with special emphasis on berry accumulation, Australian Journal of Grape and Wine Research 9: 154-168 Soil Survey Staff (2014) Keys to Soil Taxonomy, 12th edition, USDA-NRCS Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873
CEC Data
Cabernet Sauvignon grafted on 1103 Paulsen showing cell differentiation maturation zone on primary root, zone beginning 12 mm from apex, microscopic examination revealing epidermal cells differentiating into trichoblasts with emerging root hairs and atrichoblasts with thicker cuticle, cortex cells differentiating into parenchyma accumulating starch granules, endodermis cells developing casparian strips with suberin and lignin deposition in radial walls, procambial cells differentiating into xylem vessels undergoing secondary wall thickening and lignification, phloem precursors forming sieve elements and companion cells, transcriptional analysis showing tissue-specific gene expression patterns, cell differentiation creating functional specialized tissues from undifferentiated elongated cells
Cell differentiation maturation zone is maturation zone emphasizing cell specialization process and tissue-specific development, characterized by cells undergoing programmed changes converting from undifferentiated elongated cells into specialized cell types including trichoblasts and atrichoblasts in epidermis, parenchyma cells in cortex, endodermis cells with casparian strips, pericycle cells capable of lateral root initiation, xylem vessel elements and tracheids, phloem sieve elements and companion cells, showing sequential gene expression activating tissue-specific programs, progressive structural modifications including cell wall thickening, organelle reorganization, and metabolite accumulation, and demonstrating that differentiation represents fundamental developmental process determining cell fate and function enabling creation of complex multicellular root structure from simple meristematic cells, with cell differentiation involving position-dependent fate determination where cell location determines developmental trajectory, cell-cell communication through signaling molecules coordinating differentiation, and irreversible commitment where differentiated cells cannot revert to meristematic state creating permanent specialized tissues performing specific functions essential for root system operation. Cell differentiation maturation zone fate determination includes= showing position-dependent specification (cell location relative to root axis determining whether differentiating into dermal, ground, or vascular tissues), exhibiting radial patterning (concentric tissue layers epidermis, cortex, endodermis, pericycle, and vascular cylinder arising from positional information), demonstrating cell lineage (cells inheriting fate information from parent cells in meristem though position can override lineage), revealing signaling gradients (hormone and peptide gradients providing positional information), and displaying pattern formation (coordinate differentiation creating organized tissue architecture). Cell differentiation maturation zone epidermal differentiation includes= showing trichoblast-atrichoblast specification (epidermal cells differentiating into root hair-forming trichoblasts or non-hair atrichoblasts), exhibiting positional cues (cells overlying periclinal walls becoming atrichoblasts while those over anticlinal walls becoming trichoblasts), demonstrating root hair initiation (trichoblasts developing protrusion that elongates forming root hair), revealing transcriptional regulation (specific transcription factors activating trichoblast or atrichoblast programs), and displaying functional specialization (trichoblasts optimized for absorption while atrichoblasts for protection and cohesion). Cell differentiation maturation zone cortex differentiation includes= showing parenchyma development (cortex cells remaining relatively unspecialized maintaining parenchyma characteristics), exhibiting metabolic specialization (cortex cells developing storage capacity accumulating starch), demonstrating aerenchyma formation (some cortex cells potentially undergoing programmed cell death creating air spaces in waterlogged conditions), revealing mycorrhizal competence (cortex cells expressing genes enabling arbuscular mycorrhizal colonization), and displaying radial transport facilitation (cortex cells with appropriate plasmodesmatal connections enabling symplastic movement). Cell differentiation maturation zone endodermis differentiation includes= showing casparian strip synthesis (endodermis cells depositing suberin and lignin bands in radial and transverse walls), exhibiting selective permeability development (casparian strips forcing apoplastic flow into symplast creating selective barrier), demonstrating transport protein expression (endodermis cells expressing specific ion transporters controlling uptake), revealing secondary thickening (older endodermis cells potentially depositing U-shaped or O-shaped secondary walls), and displaying passage cell maintenance (some endodermis cells remaining thin-walled in older roots). Cell differentiation maturation zone vascular differentiation includes= showing procambium specification (provascular cells in meristem committing to vascular fate), exhibiting xylem-phloem distinction (procambium cells differentiating into either xylem or phloem depending on position), demonstrating xylem vessel formation (xylem precursors elongating, thickening walls, lignifying, and undergoing programmed cell death), revealing phloem sieve element formation (phloem precursors losing nuclei, forming sieve plates, developing specialized cytoplasm), and displaying companion cell development (phloem parenchyma cells differentiating into metabolically active companion cells). Cell differentiation maturation zone transcriptional control includes= showing master regulators (key transcription factors initiating tissue-specific differentiation programs), exhibiting gene cascades (sequential activation of downstream genes implementing differentiation), demonstrating epigenetic regulation (chromatin modifications controlling gene accessibility during differentiation), revealing hormonal regulation (auxin, cytokinins, and other hormones controlling differentiation through transcription factor activity), and displaying feedback loops (differentiation products potentially feeding back to regulate continued gene expression). Cell differentiation maturation zone structural changes includes= showing cell wall modifications (tissue-specific wall thickening, lignification, suberization, or other changes), exhibiting organelle reorganization (plastids differentiating into chloroplasts, amyloplasts, or remaining undifferentiated depending on cell type), demonstrating vacuole modification (vacuole composition and properties adjusting for cell function), revealing programmed cell death (vessel elements and some other cell types undergoing controlled death as final differentiation step), and displaying metabolite accumulation (cells accumulating specific compounds like starch, tannins, or other tissue-specific metabolites). Cell differentiation maturation zone timing includes= showing variable duration (differentiation requiring hours to days depending on cell type and tissue), exhibiting asynchrony (different tissues differentiating at different rates creating temporal sequence), demonstrating progressive maturation (sequential completion of differentiation features from initial commitment to full maturity), revealing wave patterns (differentiation proceeding as wave from tip basipetally continuously creating mature tissues), and displaying plasticity (differentiation timing potentially adjusting to environmental conditions).
Cell_Differentiation_Maturation_Zone emphasizes specialization process and developmental commitment and is distinguished from functional emphasis by focus on differentiation mechanism. Critical for understanding fate determination, tissue specification, transcriptional control, structural modifications, and the developmental process converting undifferentiated cells into specialized cell types creating functional root tissue organization.
Cell specialization zone
Developmental maturation zone
Gewebe-Spezifikationsbereich
Spezialisierungszone
Tissue specification region
Zelldifferenzierungs-Reifungszone
Cell Differentiation Maturation Zone
Syrah in Rhône Valley shows cell division cessation during early August Year N+1 at BBCH 75, with complete cell cycle exit by week 5-6, mitotic index declining to 0%, proliferation markers PCNA and Ki-67 becoming undetectable, all 2.2-2.8 million cells in G0/G1 expansion state, irreversible non-dividing status established, no further proliferation through subsequent 10-12 weeks development, under warm Mediterranean conditions of 25-30°C temperature, typical of this mid-to-late season cultivar completing division cessation
Cell division cessation represents the complete proliferation termination during division phase completion when all cells exit cell cycle, mitotic activity ceases entirely, proliferation markers disappear, cells establish non-dividing expansion state, and berry achieves truly non-proliferative status preventing any subsequent division throughout development, completing by weeks 5-6 as definitive proliferation end. This cessation establishment period occurs during late summer in early August in temperate regions Year N+1, typically completing week 5-6 representing complete division end, corresponding to BBCH 75 berry touch stage. Cessation mechanism involves final cell cycle exit of remaining cycling cells, complete loss of proliferative competence, cell cycle machinery downregulation to basal non-functional levels, establishment of expansion-committed non-dividing cell state, and irreversible cessation preventing division throughout subsequent 8-12 weeks until harvest. Complete cell cycle exit shows all cells leaving cycle to G0 quiescent or G1 expansion state, no cells remaining in S, G2, or M phases, cell populations uniformly non-dividing, and population synchrony in expansion state. Cell cycle phase distribution shows progression from mixed population with some cycling cells to uniform G0/G1 population, S/G2/M phase cells disappearing completely, all cells in growth phase preparing or undergoing expansion, and phase distribution documenting cessation. Mitotic activity complete cessation shows no observable mitoses after cessation completion, mitotic index 0% or <0.1% from rare measurement artifacts, absence of mitotic figures in histological examination, and truly zero division rate. Division marker disappearance shows PCNA proliferation marker becoming undetectable, cyclin proteins declining to basal non-functional levels, phospho-histone H3 mitotic marker absent, Ki-67 proliferation marker negative, and marker absence confirming non-dividing state. Cell cycle gene downregulation shows cyclin genes downregulated to minimal basal expression, cyclin-dependent kinase activity ceasing, cell cycle transcription factors E2F inactivated, and transcriptional program shift from proliferation to expansion genes. Proliferative competence loss shows cells unable to re-enter cell cycle under normal developmental conditions, division machinery dismantled preventing cycle re-entry, cells committed to expansion pathway unable to divide, and competence loss as irreversible developmental state. Expansion state establishment shows cells in growth mode through water uptake and vacuole expansion, expansion genes upregulated including expansins and transporters, cells enlarging through turgor-driven growth, and expansion as exclusive growth mode. Non-dividing cell population characteristics show population entirely in expansion state, cells progressively enlarging from 25-45 micrometers to eventual 80-200 micrometers, no further cell number increase throughout development, and population growth through cell volume increase exclusively. Cessation completion timing shows typical completion week 5 for majority of cultivars, occasional extension to week 6 in some late cultivars or cool conditions, completion timing relatively conserved across genotypes, and cessation by 4-6 weeks post-fertilization. Post-cessation development shows berry progressing through expansion, lag, veraison, ripening without any division, 8-12 weeks of development after cessation with zero proliferation, and division-free development characteristic of post-cessation berry. Irreversibility of cessation shows no division occurring after cessation under any normal conditions, environmental changes unable to reinitiate division, developmental program preventing division throughout berry development, and true irreversible cessation. Exception of wound healing shows localized cell division possible at wound sites for repair, wound-induced divisions limited to injury vicinity not affecting bulk berry, and exception not contradicting normal development cessation. Environmental independence shows cessation occurring across all environmental conditions within viable ranges, temperature variations unable to prevent or reverse cessation, and cessation as robust developmental transition. Developmental checkpoint at cessation shows likely checkpoint ensuring adequate cellularity before cessation, inadequate cell number potentially delaying cessation though evidence limited, and checkpoint passage enabling irreversible cessation. Molecular mechanisms of cessation show potential epigenetic modifications silencing cell cycle genes, chromatin remodeling preventing cell cycle gene transcription, developmental transcription factors directing expansion program, and integrated mechanisms ensuring stable cessation. Cessation as developmental commitment shows commitment to non-dividing state, establishment of expansion as exclusive growth mode, and commitment as critical developmental decision. Berry characteristics at cessation show diameter 15-25 mm depending on cultivar and achieved cellularity, fresh weight 4-10 g, established cell number 1-4 million cells, and berry in early expansion state. Cessation assessment methods include mitotic index measurement showing zero divisions, cell cycle marker immunohistochemistry showing absence, proliferation marker gene expression analysis showing downregulation, and cell cycle phase analysis showing uniform G0/G1 distribution. The process represents second aspect of division phase completion emphasizing complete cessation, following final mitotic events and preceding growth mode transition. Duration represents cessation completion typically week 5-6. Regional timing shows cell division cessation during early August across climates as proliferation ends completely. Viticultural significance includes representing complete irreversible end of cell proliferation, establishing permanent non-dividing berry state, demonstrating robust developmental transition, and marking definitive division phase conclusion. Research applications include analyzing cessation mechanisms and molecular regulation, investigating irreversibility establishment, evaluating checkpoint controls at cessation, quantifying cessation timing and completion, and assessing cessation as critical developmental transition point.
Cell_Division_Cessation is second aspect of Division_Phase_Completion, distinguished by complete proliferation termination. Represents truly zero division state by weeks 5-6. Mitotic index 0%. All cells in G0/G1 expansion state. Proliferation markers absent. Irreversible cessation preventing any subsequent division. Follows Final_Mitotic_Events. Precedes Growth_Mode_Transition. Definitive end of proliferative period. Critical developmental commitment to non-dividing state.
Zellteilungs-Beendigung
complete proliferation termination
division arrest
mitotic activity end
mitotische Aktivitäts-Ende
nicht-teilender Zustand-Etablierung
non-dividing state establishment
vollständige Proliferations-Terminierung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 441-443 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Cell Division Cessation
Merlot cell division stage berries progress from fruit set at BBCH 71 (1.8 mm diameter, 0.007 g, 0.8 million cells, 32 μm average cell diameter, 8% mitotic index) through peak division at week 3 (4.5 mm, 0.08 g, 1.6 million cells, 42 μm, 15% mitotic index) to end of cell division stage at BBCH 75 week 6 (9.5 mm, 1.1 g, 2.5 million cells, 58 μm, 3% mitotic index approaching cessation), with cell number increasing 313% while cell diameter increases 81% demonstrating cell division as primary growth mode, and with final cell number of 2.5 million determining maximum size potential for subsequent development phases. Chardonnay cell division stage berries examined at weekly intervals show mitotic activity progression= week 1 (10% mitotic index, 0.9 million cells), week 2 (14%, 1.2 million cells), week 3 (17% at peak, 1.7 million cells), week 4 (13%, 2.1 million cells), week 5 (7%, 2.3 million cells), week 6 (3%, 2.4 million cells at division cessation), demonstrating characteristic rise and fall of division activity during cell division stage with peak occurring at weeks 2-3. Syrah cell division stage berries under two water regimes show well-watered vines producing berries with final cell number of 2.7-3.1 million cells at BBCH 75 while water-deficit vines show reduced final cell number of 2-2.4 million cells (26-35% reduction), with this cell number difference established during cell division stage (weeks 0-6) persisting through maturity creating permanent size difference (well-watered mature berries 15.5-17 mm, 3-3.6 g versus deficit berries 13.5-15 mm, 2.2-2.8 g), demonstrating critical importance of water availability during cell division stage for determining ultimate berry size through effects on cell proliferation. Cabernet Sauvignon cell division stage berry mesocarp tissue sections stained for cell cycle markers show= week 2 berries with numerous cells positive for histone H3 phosphorylation (M phase marker) and cyclin B (G2/M transition marker) distributed throughout mesocarp indicating active division week 5 berries with declining marker-positive cells concentrated in inner mesocarp regions while outer mesocarp shows division cessation week 7 lag phase berries with <2% marker-positive cells indicating division has essentially ceased and cell population is fixed, confirming cell division stage completion by BBCH 75 week 6 and transition to cell expansion-dominated growth.
A cell division stage berry is a developing berry during the period when active cell division in berry mesocarp tissues is the dominant growth process (BBCH 71-75, approximately 0-6 weeks post-fruit set), corresponding to Phase I of berry development and early developing berry, characterized by rapid increase in cell number through mitotic divisions creating the total cell population that will determine ultimate berry size potential. Cell division stage berries are characterized by= (1) active mitosis in mesocarp (flesh) tissues with substantial proportions of cells in division, typically 5-18% mitotic index (percentage of cells in M phase) during peak division (2) cell number increasing from approximately 0.5-1.5 million cells at fruit set to final cell number of 1.5-3.5 million cells by end of cell division stage (BBCH 75, week 6), representing 200-350% increase in cell population (3) cells undergoing progressive cycles of division with individual cells dividing 2-5 times during the cell division stage creating exponential cell number increase (4) cells simultaneously undergoing modest expansion from 25-40 μm diameter at fruit set to 45-70 μm at end of cell division stage, with cell division being the predominant growth mode but accompanied by some cell enlargement (5) berry diameter increasing from 1.5-3 mm at fruit set to 6-11 mm at end of cell division stage (BBCH 75), with this 300-500% diameter increase driven primarily by cell number increase rather than cell size increase (6) berry fresh weight increasing from 0.005-0.015 g at fruit set to 0.6-1.5 g at BBCH 75, representing 60-150 fold mass increase achieved primarily through cell proliferation. Cell division stage berries show characteristic temporal dynamics of division activity= initial phase (weeks 0-2, fruit set establishment) with division initiating and accelerating as ovary-to-berry transition proceeds peak division phase (weeks 2-4) with maximum mitotic index (12-18% of cells dividing) and maximum cell number increase rate declining division phase (weeks 4-6) with mitotic index decreasing (5-10% then 2-5%) as cells progressively exit division and prepare for subsequent expansion-dominated growth. Cell division stage berries undergo critical size potential determination= final cell number achieved by end of cell division stage (BBCH 75) establishes the absolute maximum berry size potential, as subsequent growth occurs exclusively through expansion of this fixed cell population without additional division environmental factors during cell division stage affecting cell division (water availability, temperature, nutrition, crop load, vine vigor) have lasting effects on final berry size by determining cell number. Cell division stage berries show tissue-specific division patterns= mesocarp cells actively dividing throughout cell division stage creating the flesh tissue bulk exocarp (skin) cells dividing primarily during early cell division stage (weeks 0-3) then largely ceasing division before mesocarp division concludes seeds undergoing rapid cell proliferation in embryo and endosperm tissues overlapping with and extending beyond mesocarp cell division.
Cell_Division_Stage_Berry represents Phase I berry development (BBCH 71-75, weeks 0-6) emphasizing the cell proliferation growth mechanism, and is essentially synonymous with Early_Developing_Berry though emphasizing cellular process over developmental phase. Cell division stage represents the critical period when final cell number is determined, with lasting effects on berry size potential. Sub-classification into Rapid_Division and Peak_Division reflects temporal dynamics within the cell division period. Understanding cell division stage berry biology is essential for managing berry size through cultural practices affecting cell proliferation, and for understanding the cellular basis of berry size determination.
Proliferations-Stadium
Zellteilungs-Stadium-Beere
Zellvermehrungs-Beere
mitotic-stage berry
proliferative-phase berry
Coombe, B.G. (1976). The development of fleshy fruits. Annual Review of Plant Physiology 27=507-528 Ojeda, H. et al. (2002). Influence of water deficits on grape berry growth. Vitis 41=141-145 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 182-192.
Cell Division Stage Berry
Syrah cell enlargement stage berries progress from veraison at BBCH 81 (week 9, cells 68 μm diameter, berry 11 mm, 1.5 g, 2.4 million cells) through mid-enlargement at BBCH 85 (week 13, cells 118 μm, berry 14.5 mm, 2.8 g, 2.4 million cells) to maturity at BBCH 89 (week 17, cells 152 μm, berry 16.5 mm, 3.5 g, 2.4 million cells), with cell diameter increasing 124% (cell volume increasing 510%) while cell number remains constant at 2.4 million throughout cell enlargement stage, demonstrating that Phase II berry growth is driven exclusively by cell expansion without additional cell division. Cabernet Sauvignon cell enlargement stage berries examined at biweekly intervals show cell expansion progression= week 9 (72 μm), week 11 (95 μm, +32% from week 9), week 13 (122 μm, +69%), week 15 (142 μm, +97%), week 17 (158 μm, +119% at maturity), with expansion rate highest during early enlargement (11.5 μm per 2 weeks during weeks 9-11) and declining during late enlargement (8 μm during weeks 15-17), tracking the overall berry growth rate progression during Phase II ripening. Chardonnay cell enlargement stage berries under two irrigation regimes show well-watered vines producing cells expanding from 65 μm (veraison) to 148 μm (maturity, 128% increase, 485% volume increase) creating berries expanding from 11.5 mm to 14 mm (22% diameter increase, 83% volume increase), while deficit-irrigated vines show cells expanding only to 118 μm (82% increase, 303% volume increase) creating berries reaching 12.5 mm (9% diameter increase, 29% volume increase), demonstrating water limitation effects on cell expansion during cell enlargement stage with deficit reducing final cell size and berry size by limiting water availability for expansion. Merlot cell enlargement stage berry mesocarp cells show characteristic ultrastructural changes= veraison cells (BBCH 81) with vacuoles occupying 75-80% of cell volume, cytoplasm still substantial with numerous organelles, cell walls 0.8-1.2 μm thick mature cells (BBCH 89) with vacuoles occupying 92-95% of volume, thin cytoplasmic layer with reduced organelles, cell walls thinned to 0.4-0.7 μm through expansion and modification, demonstrating cellular reorganization during expansion creating mature berry cell structure dominated by sugar-filled vacuole.
A cell enlargement stage berry is a developing berry during the period when active cell expansion (increase in individual cell volume) in berry mesocarp tissues is the dominant growth process (BBCH 81-89, approximately 9-18+ weeks post-fruit set), corresponding to Phase II of berry development and late developing berry, characterized by dramatic increase in cell size through cell expansion driving berry enlargement, with cell number fixed at the population established during earlier cell division stage. Cell enlargement stage berries are characterized by= (1) essentially zero cell division with mitotic index <1-2% representing only occasional division events rather than systematic proliferation (2) cell number fixed at 1.5-3.5 million cells (established during Phase I) remaining constant throughout cell enlargement stage (3) individual cells undergoing dramatic expansion from veraison size (60-90 μm diameter) to mature size (100-180 μm diameter), representing 65-200% diameter increase and 280-800% volume increase per cell during cell enlargement stage (4) berry diameter increasing from veraison dimensions (9-13 mm) to mature dimensions (12-25 mm depending on cultivar), representing 30-90% diameter increase driven exclusively by cell expansion (5) berry fresh weight increasing from veraison weight (1-2 g) to mature weight (1.5-7 g), representing 50-300% weight increase achieved through water accumulation into expanding cells (6) cells accumulating water, sugars, and other solutes creating osmotic driving force for expansion, with vacuoles expanding to occupy 90-95% of cell volume in mature cells. Cell enlargement stage berries show characteristic progression= early enlargement (weeks 9-12, BBCH 81-85) with rapid cell expansion at 5-12 μm diameter increase per week driving rapid berry growth mid enlargement (weeks 12-15, BBCH 85-87) with continuing expansion at 3-8 μm per week late enlargement (weeks 15-18+, BBCH 87-89) with declining expansion rate approaching cessation at maturity. Cell enlargement stage berries undergo critical processes supporting expansion= cell wall loosening through expansins, xyloglucan endotransglucosylases, and pectinases allowing wall extension vacuole expansion through water accumulation driven by osmotic potential from sugar and solute accumulation membrane synthesis keeping pace with cell surface area increase cytoskeleton reorganization supporting cell shape changes during expansion. Cell enlargement stage berries show metabolic shift from biosynthesis (dominant during cell division stage) to accumulation (dominant during enlargement stage)= sugar accumulation in vacuoles reaching 18-28% at maturity organic acid changes with malate declining through respiration secondary metabolite accumulation including anthocyanins, tannins, flavor compounds declining respiration rates (from 25-35 nmol CO₂ g⁻¹ s⁻¹ at veraison to 8-15 nmol at maturity) reflecting shift from biosynthetic metabolism to storage metabolism.
Cell_Enlargement_Stage_Berry represents Phase II berry development (BBCH 81-89, weeks 9-18+) emphasizing the cell expansion growth mechanism, and is essentially synonymous with Late_Developing_Berry though emphasizing cellular process over developmental phase. Cell enlargement stage represents the period when water availability critically affects final berry size through effects on cell expansion. Sub-classification into Early_Enlargement and Maximum_Enlargement reflects temporal progression within the enlargement period. Understanding cell enlargement stage biology is essential for irrigation management affecting berry size and composition, and for understanding the cellular basis of Phase II berry growth and ripening.
Expansions-Beere
Phase II berry
Zellexpansions-Stadium
Zellvergrößerungs-Stadium-Beere
cell-expansion berry
Coombe, B.G., McCarthy, M.G. (2000). Dynamics of grape berry growth. Australian Journal of Grape and Wine Research 6=131-135 Matthews, M.A., Anderson, M.M. (1988). Fruit ripening in Vitis vinifera= responses to seasonal water deficits. American Journal of Enology and Viticulture 39=313-320.
Cell Enlargement Stage Berry
Pinot Noir grafted on 101-14 Millardet showing cell expansion elongation zone on third-order lateral root, cells entering zone at meristem-elongation boundary measuring 14 micrometers length exiting zone at maturation boundary measuring 380 micrometers length representing 27-fold length increase, cell volume increasing from 3500 cubic micrometers to 210,000 cubic micrometers (60-fold increase), central vacuole expanding from 40% to 88% cell volume, turgor pressure measured 0.54 MPa, osmotic potential -1.1 MPa, cell expansion through coordinated ion uptake, water influx, wall loosening by expansins, and wall synthesis producing irreversible turgor-driven growth
Cell expansion elongation zone is elongation zone emphasizing cell expansion mechanism and turgor-driven growth process, characterized by cells increasing volume 10-100 fold through water uptake into enlarging central vacuole, showing cell length increasing 10-50 fold from meristematic 10-20 micrometers to elongated 100-500 micrometers while diameter remaining relatively constant creating high aspect ratio, demonstrating turgor pressure 0.3-0.8 MPa generated by osmotic water influx providing mechanical force stretching cell walls, and revealing that cell expansion represents fundamental growth mechanism where osmotic potential difference between cell interior and soil solution drives water entry creating turgor that irreversibly extends cell walls through enzymatic loosening and continued synthesis producing permanent cell enlargement enabling root elongation, with cell expansion process requiring coordinated regulation of ion uptake generating osmotic potential, water transport providing volume increase, wall loosening allowing extension, and wall synthesis maintaining integrity creating complex integration of biochemical and biophysical processes. Cell expansion elongation zone volume increase includes= showing dramatic size change (cells increasing total volume 10-100 fold during passage through expansion zone), exhibiting vacuole dominance (central vacuole expanding from 30% to 80-90% of cell volume providing volume increase), demonstrating water influx (osmotic water uptake providing volume increase with minimal dry matter accumulation), revealing length preferentially (cell volume increase predominantly as length increase 10-50 fold versus diameter increase 1-2 fold), and displaying irreversible growth (cell expansion permanent creating stable enlarged cells unlike reversible elastic deformation). Cell expansion elongation zone osmotic mechanism includes= showing solute accumulation (active K+, Cl-, sugars, and organic acids uptake increasing cell osmotic potential), exhibiting membrane transport (H+-ATPases energizing ion uptake creating electrochemical gradients), demonstrating osmotic gradient (cell osmotic potential -0.5 to -1.5 MPa creating water potential gradient drawing water from soil), revealing aquaporin function (water channel proteins facilitating rapid water movement across membranes), and displaying continuous adjustment (ongoing solute uptake maintaining osmotic potential despite dilution from water influx). Cell expansion elongation zone turgor generation includes= showing water uptake (osmotic gradient drawing water into cell increasing volume), exhibiting pressure development (water influx constrained by cell wall creating turgor pressure 0.3-0.8 MPa), demonstrating mechanical force (turgor pressure exerting force on cell walls promoting extension), revealing pressure maintenance (continuous solute and water uptake maintaining turgor throughout expansion), and displaying yield threshold (turgor exceeding wall yield threshold causing irreversible extension). Cell expansion elongation zone wall loosening includes= showing expansin action (expansin proteins disrupting hydrogen bonds between cellulose and matrix polysaccharides increasing wall extensibility), exhibiting pH effects (apoplast acidification activating expansins and other wall-loosening processes), demonstrating enzymatic modification (xyloglucan endotransglucosylases, pectinases, and other enzymes modifying wall structure), revealing mechanical loosening (turgor-induced stress potentially accelerating loosening through stress-strain effects), and displaying regulated extensibility (controlled wall loosening balancing expansion with structural integrity). Cell expansion elongation zone wall synthesis includes= showing continuous deposition (new cellulose, hemicellulose, and pectin synthesized and incorporated during expansion), exhibiting plasma membrane synthesis (expanding cell surface requiring new membrane lipid and protein production), demonstrating balanced growth (wall synthesis rate matching expansion rate maintaining appropriate wall thickness 0.1-0.5 micrometers), revealing biosynthesis sites (Golgi apparatus producing matrix polysaccharides, plasma membrane cellulose synthase producing cellulose), and displaying resource requirement (wall synthesis consuming substantial carbohydrates and energy). Cell expansion elongation zone directional control includes= showing transverse microtubules (cortical microtubules oriented perpendicular to elongation axis directing cellulose deposition), exhibiting anisotropic expansion (preferential longitudinal expansion through constrained radial expansion), demonstrating cellulose orientation (newly deposited cellulose microfibrils aligned transversely restraining radial expansion), revealing mechanical anisotropy (wall structure creating differential extensibility longitudinal greater than radial), and displaying shape determination (microtubule-directed cellulose deposition determining final cell shape). Cell expansion elongation zone hormone integration includes= responding to auxin (promoting proton secretion, expansin expression, and wall loosening), affected by gibberellins (enhancing expansion through wall enzyme expression and metabolism), influenced by brassinosteroids (promoting expansion through multiple pathways), showing ethylene effects (potentially inhibiting expansion under stress), and demonstrating abscisic acid action (reducing expansion under water deficit protecting from turgor loss). Cell expansion elongation zone biophysics includes= showing Lockhart equation applicability (growth rate proportional to turgor exceeding yield threshold times wall extensibility), exhibiting stress-strain relationships (wall mechanical properties determining expansion response to turgor), demonstrating viscoelastic behavior (wall showing both elastic and plastic deformation components), revealing time-dependent extension (wall creep under sustained turgor producing continued expansion), and displaying mechanical-biological coupling (mechanical stress affecting gene expression and metabolism creating feedback). Cell expansion elongation zone completion includes= showing deceleration (expansion rate declining as cells approach maturation zone), exhibiting wall rigidification (progressive cross-linking reducing extensibility terminating expansion), demonstrating differentiation onset (cells beginning tissue-specific development incompatible with continued expansion), revealing final size (cells reaching mature dimensions 100-500 micrometers length typical for root cells), and displaying transition (completion of expansion marking entry into maturation zone for differentiation).
Cell_Expansion_Elongation_Zone emphasizes expansion mechanism and turgor-driven process and is distinguished from other elongation zone aspects by focus on biophysical growth mechanism. Critical for understanding osmotic water uptake, turgor generation, wall loosening, cell volume increase, anisotropic expansion, and the fundamental turgor-driven cell expansion mechanism enabling root elongation through irreversible cell enlargement.
Turgor-driven expansion zone
Turgor-getriebene Streckungszone
Vacuolation elongation region
Vakuolisierungs-Wachstumsbereich
Water-uptake growth zone
Zellexpansions-Elongationszone
Cosgrove, D. J. (2005). Growth of the plant cell wall. Nature Reviews Molecular Cell Biology, 6(11), 850-861.
Cell Expansion Elongation Zone
Pinot Noir in Burgundy shows cell number finalization during late July-early August Year N+1 at BBCH 75, with final late divisions completing week 4-5, total mesocarp cell count reaching 1.6-2.0 million cells representing final permanent value, cell number determining 74-82% mature berry size variation, no further proliferation occurring through subsequent development, under continental conditions of 22-28°C temperature, typical of this early-to-mid season cultivar establishing cellularity determining 9-11 mm final berry size potential
Cell number finalization represents the final cell count establishment phase during declining division activity when remaining late divisions complete, total mesocarp cell number reaches final permanent value typically 1-4 million cells per berry, no further cell proliferation occurs throughout subsequent berry development, and established cellularity determines berry size potential for remainder of development, occurring by weeks 4-5 post-fertilization as division phase concludes. This critical finalization phase occurs during late summer in late July-early August in temperate regions Year N+1, typically completing by week 4-5 representing division cessation, corresponding to BBCH 75 berry touch stage. Finalization mechanism involves completion of final late divisions from remaining cycling cells, cell cycle exit of last dividing cells establishing non-dividing cell population, final cell count representing cumulative proliferative output from entire division phase, irreversible cell number preventing any subsequent proliferation, and established cellularity as permanent berry characteristic. Final division completion shows last mitoses occurring week 4-5 in late-dividing cell populations, completion timing variable across cells creating extended finalization period, occasional rare divisions possibly extending to week 6 in small subpopulations, and eventual complete cessation establishing final count. Cell number accumulation to final value shows progressive increase during declining activity approaching plateau, typical progression to 85-95% final number by week 4 and 95-99% by week 5, remaining 1-5% cells from final late divisions, and final accumulation to complete count. Final cell number range shows cultivar variation with small-berried cultivars 0.8-1.5 million mesocarp cells, medium-berried 1.5-2.5 million cells, large-berried 2.5-4 million cells, and range reflecting genetic programming and environmental effects during division phase. Cell number determinants summary shows genetic programming as primary factor establishing cultivar-characteristic ranges, seed number affecting total through differential auxin dose creating 20-40% variation within cultivar, temperature during division phase affecting proliferation rate and final number with warm conditions enabling maximum genetic potential, water and nutrients supporting but usually not limiting division, and cumulative environmental effects during critical weeks 1-4 determining achieved cellularity. Seed number effects on final cellularity show positive correlation between seeds per berry and final cell number, typical 25-35% more cells in 3-seed versus 1-seed berries, auxin dose-response relationship mediating seed effects, and seed number as major within-cultivar variation source. Temperature integration effects show heat accumulation during division phase correlating with final cell number, growing degree days during weeks 1-4 affecting total divisions, cool conditions reducing final count 15-30% from genetic maximum, warm conditions enabling full genetic potential achievement, and temperature as major environmental determinant. Cell number measurement methods show enzymatic tissue dissociation followed by cell counting using hemocytometer or flow cytometry, DNA content analysis estimating cell number from total DNA, histological section analysis though technically challenging for large cell populations, and measurement enabling cellularity quantification. Cell number irreversibility shows established count permanent and unchangeable, no further cell division occurring during subsequent expansion, lag phase, ripening phases, rare exception of wound healing producing localized new cells though not affecting bulk berry, and irreversibility emphasizing critical nature of division phase. Cell number as size determinant shows strong positive correlation between final cell number and mature berry size, typical correlation coefficient r² 0.70-0.85 across cultivars and conditions, cell number accounting for 60-80% size variation with remainder from differential expansion, and cellularity as primary size determinant. Predictive relationship shows final cell number enabling early berry size prediction, berry diameter at maturity predictable from cell count with reasonable accuracy, and early finalization enabling yield estimation from cell number assessment. Cell size at finalization shows cells relatively small 25-45 micrometers diameter following division phase, cells retaining substantial expansion capacity for subsequent 5-20 fold volume increase, and initial small size characteristic of recently divided populations. Expansion potential establishment shows final cell number determining total expandable cell population, all cells capable of subsequent enlargement contributing to berry growth, and expansion phase exploiting established cell population. Tissue architecture finalization shows pericarp layer organization established with exocarp, mesocarp, endocarp differentiation complete, vascular network development matching cellularity, and structural organization from coordinated division patterns. Cell type proportions shows final proportions established with mesocarp 80-85% cells, exocarp 10-15%, endocarp 3-5%, and proportions reflecting tissue-specific proliferation patterns. Developmental commitment shows cell number finalization representing irreversible developmental commitment, established cellularity constraining subsequent berry size achievement, and finalization as critical developmental decision point. Environmental independence after finalization shows subsequent environmental conditions unable to affect established cell number, later favorable conditions unable to compensate for limited early division, and finalization emphasizing critical nature of division phase environment. The process represents third and final aspect of declining division activity emphasizing count establishment, following division rate reduction and transition to enlargement onset. Duration represents finalization completion by weeks 4-5. Regional timing shows cell number finalization during late July-early August across climates by division phase end. Viticultural significance includes establishing final permanent cell count determining berry size potential, demonstrating irreversible nature of division phase establishing size constraints, showing critical importance of optimal division phase conditions for size achievement, and marking definitive end of cell proliferation period. Research applications include analyzing cell number determinants and variation, investigating genetic and environmental effects on final cellularity, evaluating cell number as berry size predictor, quantifying cellularity-size relationships, and assessing cell number finalization as critical developmental determinant.
Cell_Number_Finalization is third and final aspect of Declining_Division_Activity, distinguished by final permanent cell count establishment. Represents completion by weeks 4-5. Final range 1-4 million cells depending on cultivar. Determined by genetics, seed number, temperature during weeks 1-4. Irreversible - no subsequent proliferation. Accounts for 60-80% size variation. Strong predictor of mature berry size. Follows Division_Rate_Reduction and Transition_to_Enlargement_Onset. Completes Declining_Division_Activity. Critical developmental commitment point.
Zellzahl-Etablierung
Zellzahl-Finalisierung
cell number establishment
cellularity finalization
final cell count
finale Zellzahl
permanent cell number
permanente Zellzahl
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 439-443 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Cell Number Finalization
In Mourvèdre vines in Bandol, Provence during November, cell separation process proceeds over 4 days following abscission zone activation, with polygalacturonase activity increasing from 0.8 to 28 nmol/min/g fresh weight, middle lamella dissolution visible histologically within 48 hours, cortical cell separation complete by day 3 while vascular strands persist until day 4, and final detachment occurring when wind gusts exceed 15 km/h providing sufficient mechanical force to sever remaining connections under cool Mediterranean autumn conditions.
Cell separation process describes the enzymatic dissolution of cell walls and middle lamellae in Vitis vinifera petiole abscission zone cells that physically separates leaf from shoot, wherein the coordinated action of cellulases, polygalacturonases, xylanases, and other cell wall degrading enzymes progressively weakens the structural connections between adjacent cells until mechanical forces from leaf weight and environmental factors complete physical detachment over a period of 3-5 days following abscission zone activation. This process involves the sequential degradation of cell wall components, with pectin hydrolysis by polygalacturonases dissolving the calcium-pectate-rich middle lamella that cements adjacent cells together, cellulase activity degrading cellulose microfibrils that provide structural strength to primary cell walls, and xylanase and other hemicellulase activities cleaving the matrix polysaccharides that cross-link cellulose within the cell wall architecture. The spatial progression of cell separation proceeds from the outer cortical regions of the abscission zone inward toward the vascular tissues, with peripheral cells separating first while vascular connections maintain leaf attachment until late stages when xylem and phloem elements are finally severed, explaining the observation that partially separated leaves may remain attached by vascular strands until mechanical stress completes detachment. The biochemical mechanisms involve secretion of cell wall degrading enzymes into the apoplastic space between abscission zone cells, with enzyme activities increasing 10-50 fold during the separation period compared to quiescent abscission zones, and optimal enzyme function at the slightly acidic apoplastic pH of 5.0-6.0 characteristic of this intercellular compartment. The role of expansins in cell separation involves loosening of cell wall structure through disruption of hydrogen bonds between cellulose microfibrils and matrix polysaccharides, creating access for hydrolytic enzymes to their substrates and facilitating the swelling and separation of loosened wall material under turgor pressure. Environmental factors influence cell separation rate, with adequate moisture maintaining turgor pressure that drives cell expansion and wall separation, cool temperatures enhancing enzyme activities in the temperature-sensitive polygalacturonase reaction, and wind or rain providing mechanical forces that accelerate separation of weakened tissues. The completion of cell separation creates a clean fracture surface at the leaf scar, with the shoot-side cells remaining attached to the stem and undergoing protective modifications while the leaf-side cells separate with the falling leaf.
Cell_Separation_Process is subclass of Petiole_Abscission representing enzymatic cell wall degradation. Distinguishes from activation by emphasizing physical dissolution. Sibling to Abscission_Zone_Formation, Abscission_Zone_Activation, and Petiole_Abscission_Completion. Duration 3-5 days. Involves cellulases, polygalacturonases, xylanases, expansins. Outer-to-inner progression with vascular persistence.
Abszissionszone-Zell-Separation
Zell-Trennungs-Prozess
Zellwand-Auflösung
abscission zone cell separation
cell detachment process
cell wall dissolution
intercellular separation
interzelluläre Trennung
Roberts JA et al (2002) Abscission, dehiscence and other cell separation processes. Annual Review of Plant Biology 53: 131-158 Taylor JE and Whitelaw CA (2001) Signals in abscission. New Phytologist 151: 323-340 Patterson SE (2001) Cutting loose. Plant Physiology 126: 494-500
Cell Separation Process
Merlot in Bordeaux shows cell size maximum during early-mid August Year N+1 at BBCH 77-79, with mesocarp cells reaching 92-95% final dimensions at 96-132 micrometers diameter representing 62-fold expansion from 28 micrometer post-division size, vacuoles occupying 89% cell volume, cytoplasm 2.8 micrometer peripheral layer, wall 0.16 micrometer thickness, osmotic potential -1.4 MPa, under oceanic conditions of 22-27°C temperature, typical of this mid-season cultivar approaching genetic cell size limits
Cell size maximum represents the genetic dimension limit approach phase during maximum berry enlargement when individual cells reach 90-95% final mature size, typical diameters 75-190 micrometers depending on cultivar and position, further expansion becomes increasingly constrained by genetic programming and physical limitations, cells achieve near-final volume representing culmination of 30-100 fold total expansion from post-division dimensions, occurring during late enlargement weeks 6-8 as cells approach size limits. This cell dimension culmination phase occurs during late summer in early-mid August in temperate regions Year N+1, representing maximum cell size achievement weeks 6-8 post-fertilization, corresponding to BBCH 77-79 bunch closure stages. Cell size maximum mechanism involves cells approaching genetically determined size limits programmed during development, physical constraints from cell wall properties and cellular architecture limiting further expansion, reduced wall plasticity preventing continued yielding, osmotic and turgor systems approaching equilibrium states, developmental signals directing expansion cessation, and integrated limitations establishing maximum cell dimensions. Cell dimensions at maximum show typical ranges 75-190 micrometers diameter depending on cultivar, small-berried cultivar cells 60-100 micrometers maximum, medium-berried 90-140 micrometers, large-berried 120-200 micrometers, and size ranges reflecting genetic variation. Cell volume at maximum shows volumes 0.18-3.6 nanoliters representing culmination of expansion, 30-100 fold increase from post-division 25-45 micrometer cells, substantial volume achieved through intensive water uptake and vacuole expansion, and maximum volume as endpoint of enlargement. Vacuole at maximum size shows central vacuole occupying 85-92% cell volume, vacuole diameter 65-185 micrometers in large cells, maximum vacuole representing genetic and physical limits, tonoplast membrane area 1200-6000 square micrometers, and vacuole as dominant compartment at maximum. Cytoplasm configuration at maximum shows thin peripheral layer 1.5-4 micrometers thickness, cytoplasm containing nucleus and organelles in compressed space, maintained metabolic function in minimal volume, and efficient organization in fully expanded cells. Genetic control of cell size shows developmental programming establishing cultivar-specific size ranges, cell cycle exit timing affecting expansion duration and final size, genetic variation in expansion capacity determining maximum dimensions, and intrinsic genetic regulation as primary size determinant. Cell size regulatory mechanisms show possible mechanosensing of cell wall tension signaling size achievement, developmental transcription factors directing expansion cessation, cell-cell communication coordinating population sizing, and integrated regulation ensuring appropriate maxima. Wall properties at maximum show increased wall thickness 0.12-0.25 micrometers from continued synthesis, reduced wall plasticity from cross-linking and structural modifications, increased yield threshold requiring higher turgor for further expansion, wall mechanical properties shifted toward growth-restrictive, and wall as physical constraint limiting expansion. Wall composition at maximum shows accumulated cellulose microfibrils creating structural framework, extensive hemicellulose and pectin cross-linking, possible calcium-mediated pectin cross-links strengthening walls, and mature wall composition limiting plasticity. Osmotic properties at maximum show osmotic potential -1.2 to -1.6 MPa at near-maximum from accumulated solutes, malate and tartrate at peak or declining concentrations 1.2-1.6% total, potassium 80-140 mM, sugars beginning accumulation 6-10%, and osmotic potential approaching steady state. Turgor at maximum shows pressure maintained 0.6-1.0 MPa, turgor sustained but expansion minimal from wall constraints, turgor-expansion relationship shifted with high pressure required for minimal growth, and pressure maintenance without substantial expansion. Cell expansion cessation approaching shows expansion rate declining to very low levels, eventual complete cessation establishing final cell dimensions within days to weeks, cells retaining minor expansion capacity adding final 5-10% during lag and ripening, and near-cessation at maximum enlargement. Remaining expansion capacity shows slight continued enlargement possible during lag and ripening phases, minor volume increases 5-10% from continued limited expansion and ripening processes, and residual capacity allowing final size adjustments. Cell size variation within berry shows mesocarp cells showing size range with inner cells typically larger than outer, exocarp epidermal cells smaller typically 40-80 micrometers, cell position affecting maximum size, and spatial heterogeneity in cell dimensions. Cell type size differences shows parenchyma mesocarp cells as largest cell type reaching maximum dimensions, vascular tissue cells smaller with specialized morphology, epidermal cells smallest with different expansion patterns, and cell type affecting size ranges. Environmental effects on cell size maximum include water availability during enlargement affecting achieved maximum with deficit potentially limiting final size 10-20%, temperature affecting expansion kinetics and timing of maximum, and genetic factors dominating over environmental within normal ranges. Water deficit effects on maximum show severe deficit during peak expansion permanently limiting cell size, deficit during maximum enlargement reducing final 5-15%, and water as major environmental factor affecting achieved maximum. Cultivar variation in cell size maximum shows genetic differences as primary variation source, small-berried cultivars genetically programmed for 60-100 micrometer cells, large-berried cultivars achieving 120-200 micrometers, and cultivar characteristic maxima determining berry size potential. Cell size as berry size determinant shows final berry size determined by cell number × cell size, cell size accounting for 20-40% berry size variation, cell size maximum establishing half of size equation, and maximum cell dimensions critical for final berry size. Cell size measurement shows microscopy-based diameter measurements, cell volume calculations from dimensions, population size distribution analysis, and measurement enabling cell size quantification. The process represents first aspect of maximum berry enlargement emphasizing cellular dimension limits, preceding berry size peak and expansion rate decline. Duration represents maximum size achievement during weeks 6-8. Regional timing shows cell size maximum during early-mid August across climates as cells approach genetic limits. Viticultural significance includes representing cellular size limit determining berry size potential, demonstrating genetic control of cell dimensions, establishing final cell size affecting berry size and quality, and marking cellular growth culmination. Research applications include analyzing genetic and developmental controls of cell size, investigating size limit mechanisms and constraints, evaluating environmental effects on maximum achievement, quantifying cell size variation and determinants, and assessing cell size maximum as berry size determinant.
Cell_Size_Maximum is first aspect of Maximum_Berry_Enlargement, distinguished by cellular dimension limits. Represents cells at 90-95% final size, 75-190 micrometers depending on cultivar. 30-100 fold total expansion from post-division. Vacuole 85-92% volume. Wall plasticity reduced. Osmotic potential -1.2 to -1.6 MPa plateau. Genetic programming primary determinant. Duration weeks 6-8. Preceded by Rapid_Cell_Expansion. Followed by Berry_Size_Peak. Establishes cellular component of final berry size.
Zellgröße-Maximum
Zellgrößen-Limit
cell size limit
cellular dimension peak
genetic cell maximum
genetisches Zell-Maximum
maximale Zelldimensionen
maximum cell dimensions
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 448-450 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Cell Size Maximum
Cell turgor comparison in Chardonnay leaf tissue reveals guard cell turgor of 2.8 MPa in open stomata during morning transpiration declining to 0.9 MPa at stomatal closure during afternoon water stress, while adjacent epidermal cells maintain relatively stable turgor of 0.45±0.12 MPa throughout the diurnal cycle, demonstrating the hydraulic isolation enabling guard cells to function as osmotic motors responding to environmental signals while bulk leaf turgor remains buffered.
Cell turgor quality is a turgor pressure quality that inheres in individual grapevine cells and represents the hydrostatic pressure specific to particular cell types that differ in their osmotic contents, wall properties, and hydraulic connectivity determining their characteristic turgor behavior under varying vine water status. Within the BFO framework, this quality enables ontological differentiation of turgor at the cellular level, recognizing that different cell types within the same tissue maintain distinct turgor values reflecting their specialized functions and structural properties. The ontological significance lies in cell-type-specific turgor determining functional capacity, where guard cells require turgor modulation for stomatal movement, mesocarp cells rely on turgor for fruit texture, and epidermal cells use turgor for surface integrity and pathogen resistance. Cell turgor varies dramatically among grapevine cell types, with actively growing cells maintaining turgor above the yield threshold for wall extension (typically 0.3-0.5 MPa), guard cells showing turgor fluctuations of 1-2 MPa driving stomatal aperture changes, and mature storage parenchyma in berries potentially operating near zero turgor post-véraison. The hydraulic isolation or connectivity of cells affects turgor dynamics, where symplastically connected cells equilibrate rapidly while apoplastically isolated cells may maintain turgor differentials relative to surrounding tissue. Measurement at cellular resolution requires pressure probe techniques capable of accessing individual cells, or may be inferred from cell-type-specific responses to osmotic manipulation. The developmental trajectory of cell turgor reflects both changing osmotic contents as cells mature and differentiate, and changing wall properties as primary walls transition to secondary walls in lignifying tissues. Understanding cell turgor quality at the resolution of individual cell types enables mechanistic interpretation of tissue-level properties emerging from the collective behavior of heterogeneous cell populations.
Cell_Turgor_Quality provides the cellular-resolution specification of turgor, serving as parent for cell-type-specific turgor classes (Mesocarp, Guard Cell, Epidermal). This quality emphasizes that tissue-level turgor represents an emergent property from heterogeneous cellular contributions. Direct measurement requires specialized microprobe techniques with limited throughput, making most turgor assessments rely on tissue-level methods that average across cell populations.
Einzelzellturgor
Zellturgor
cell-type turgor
cellular turgor pressure
individual cell pressure
single cell turgor
zelltypspezifischer Turgor
zellulärer Turgordruck
Franks PJ et al. (2001) Guard cell pressure/aperture characteristics measured with the pressure probe. Plant, Cell & Environment 24(2): 163-171 Wada H et al. (2008) Turgor-responsive starch phosphorylase in grape berry skin. Phytochemistry 69(6): 1298-1306
Cell Turgor Quality
In a Napa Valley Cabernet Sauvignon vineyard, cell wall breakdown maximum was measured during a 3-day period in mid-September Year N+1 (BBCH 84) when polygalacturonase activity peaked at 175 units, pectin molecular weight declined from 155 kDa to 92 kDa, soluble pectin increased from 56% to 73% of total, middle lamella adhesion strength dropped from 2.8 N to 1.2 N, and berry firmness decreased from 2.4 N to 1.6 N as temperature conditions averaged 25-27°C optimal for enzyme activity.
Cell_Wall_Breakdown_Maximum represents the interval during peak softening when enzymatic degradation of berry cell wall structural polysaccharides reaches maximum intensity, characterized by peak activities of multiple cell wall-modifying enzymes operating concurrently to maximize disassembly of the pectin-hemicellulose network that provides cellular mechanical strength. This critical phase typically occurs during a 2-4 day period within mid-veraison (BBCH 83-84) and is defined by convergence of maximum enzyme activities: polygalacturonase reaches 120-220 units (8-15 fold above pre-veraison baseline) cleaving homogalacturonan backbone α-1,4 linkages at maximum rate, pectin methylesterase achieves 110-200 units demethylating pectin polymers to create optimal PG substrates, pectate lyase shows 30-70 units eliminating methylated galacturonic acid residues through β-elimination, and β-galactosidase operates at 30-70 units removing galactose side chains from rhamnogalacturonan I regions facilitating backbone access by other enzymes. The maximum breakdown period produces profound structural modifications: pectin molecular weight undergoes steepest decline from approximately 130-180 kDa to 70-110 kDa representing extensive depolymerization with average chain length reduction from 700-1000 to 350-600 galacturonic acid residues, soluble pectin fraction increases maximally at 8-15% per day reaching 65-80% of total pectin (from 35-50% at breakdown initiation), and degree of methylesterification drops from 60-75% to 30-45% exposing carboxyl groups that create charge repulsion and pectin chain separation. Middle lamella degradation reaches maximum intensity during cell wall breakdown maximum: pectin-rich middle lamella between adjacent cells undergoes extensive solubilization with 60-80% of structural pectin depolymerized, intercellular adhesion strength declines by 50-70% from pre-breakdown values measured through cell separation force testing, and transmission electron microscopy reveals progressive middle lamella dissolution with electron-lucent zones replacing electron-dense pectin networks characteristic of firm berries. Hemicellulose modifications contribute substantially to maximum breakdown: xyloglucan endotransglycosylase activity peaks at 50-120 units catalyzing transglycosylation reactions that restructure xyloglucan chains reducing cell wall tensile strength by 40-60%, α-arabinosidase and β-xylosidase activities (15-40 units each) remove arabinose and xylose decorations from xyloglucans facilitating backbone modifications, and glucanase activity (20-50 units) contributes to general polysaccharide network loosening. Gene expression patterns underlying cell wall breakdown maximum show coordinated peak transcription: PG genes exhibit 100-200 fold upregulation above green berry levels with some cultivars showing multiple PG isoforms expressed sequentially or concurrently, expansin genes reach 70-140 fold elevation facilitating stress relaxation and irreversible cell wall extension, pectin methylesterase genes show 50-100 fold increases, and xyloglucan endotransglycosylase genes achieve 40-80 fold upregulation all orchestrated by sustained high abscisic acid signaling (1500-3500 ng/g fresh weight). Regulatory control of maximum breakdown involves transcriptional and post-translational mechanisms: ripening transcription factors including VvNAC, VvMYB, and VvMADS-box proteins reach peak expression coordinating cell wall enzyme gene transcription, protein synthesis machinery operates at maximum capacity producing 20-40 mg cell wall enzymes per kg berry fresh weight per day, and enzyme secretion through Golgi-mediated vesicular transport achieves maximum rates delivering enzymes to cell wall targets. Temperature effects on cell wall breakdown maximum are pronounced: optimal conditions (22-28°C) support maximum enzyme synthesis, stability, and activity producing complete breakdown patterns, cooler temperatures (16-22°C) reduce enzyme reaction rates by 35-55% extending the breakdown period and potentially producing incomplete degradation, while excessive heat (>30°C) can denature enzymes reducing maximum activities by 25-45% despite accelerated reaction kinetics at elevated temperatures. Subcellular consequences of maximum breakdown inclu cell wall thickness reduction from 2.5-4.0 μm to 1.5-2.5 μm through polymer degradation and water infiltration, porosity increases allowing enhanced diffusion of metabolites and water-soluble compounds between cells, and mechanical properties shift from rigid elastic behavior to compliant plastic deformation as structural integrity declines. Cultivar variation in cell wall breakdown maximum reflects genetic differences in enzyme complement and cell wall composition: high-PG cultivars like Gewürztraminer show very intense breakdown with 75-85% pectin solubilization and molecular weight dropping to 60-90 kDa, moderate cultivars like Chardonnay exhibit intermediate breakdown (65-75% solubilization, 80-120 kDa final MW), while low-enzyme varieties like Barbera display less extensive breakdown (55-70% solubilization, 100-140 kDa MW) often retaining firmer texture. Environmental stresses during maximum breakdown can substantially modify the process: water deficit reduces breakdown intensity through 30-50% lower enzyme gene expression and synthesis limiting cell wall modifications and potentially maintaining firmer berries, excess water can accelerate breakdown through enhanced turgor-driven cell expansion activating mechanosensitive cell wall loosening pathways, and nutrient deficiency particularly nitrogen limitation during ripening can reduce enzyme synthesis capacity limiting maximum breakdown achievement. Berry quality implications of cell wall breakdown maximum timing and completeness inclu complete breakdown coordinated with sugar and phenolic accumulation produces optimal berry texture for quality wine production, incomplete breakdown results in excessively firm berries with potential processing problems (difficult crushing, incomplete extraction), while premature or excessive breakdown creates overly soft fruit vulnerable to damage and with potential aromatic compound losses through increased volatilization from disrupted cellular compartmentation. Viticultural monitoring of cell wall breakdown maximum through enzyme assays and pectin analysis provides harvest timing guidance particularly for premium wine production where texture balance is critical for desired wine style and aging potential.
Cell_Wall_Breakdown_Maximum is distinguished from other softening phases by representing the convergence point of peak activities for multiple cell wall-degrading enzymes rather than firmness decline per se. Duration typically 2-4 days during peak softening. Enzyme activities at maximum: PG 120-220 units, PME 110-200 units, XET 50-120 units, β-Gal 30-70 units. Pectin solubilization contributes 55-70% to firmness loss, hemicellulose modification 20-30%, other factors 10-20%. Temperature optimum 22-28°C for maximum enzyme function. Cell_Wall_Breakdown_Maximum represents the fundamental mechanistic basis for berry softening and determines final berry texture characteristics.
Spitzen-Pektin-Depolymerisation
Spitzen-Zellwand-Enzym-Aktivität
maximale Polysaccharid-Degradation
maximaler struktureller Abbau
maximum polysaccharide degradation
maximum structural breakdown
peak cell wall enzyme activity
peak pectin depolymerization
Keller M (2020) The Science of Grapevines 3rd Ed pages 230-232 cell wall degrading enzymes in ripening Nunan KJ, Sims IM, Bacic A, Robinson SP, Fincher GB (1998) Changes in cell wall composition during ripening of grape berries Plant Physiol 118:783-792 pectin and hemicellulose breakdown Goulao LF, Santos J, de Sousa I, Oliveira CM (2007) Patterns of enzymatic activity of cell wall-modifying enzymes during growth and ripening of apples Postharvest Biol Technol 43:307-318 comparative enzyme dynamics.
Cell Wall Breakdown Maximum
Cell wall elasticity mapping in Pinot Noir berry mesocarp using atomic force microscopy reveals decline in wall elastic modulus from 125 MPa at pre-véraison to 45 MPa at harvest maturity, with the 64% reduction correlating temporally with polygalacturonase activity increase and pectin molecular weight decrease, demonstrating the causal relationship between cell wall biochemical modification and the mechanical changes underlying berry softening.
Cell wall elasticity quality is a tissue elasticity quality that inheres in the polysaccharide matrix surrounding plant cells and represents the capacity of the wall structure to undergo reversible deformation under turgor-generated or externally applied stress, determining the wall's contribution to cellular mechanical behavior independent of turgor pressure effects. Within the BFO framework, this quality specifically characterizes the elastic behavior of the extracellular structural component, distinguishing wall mechanical properties from the turgor-dependent elasticity of whole cells or tissues. The ontological significance lies in cell wall elasticity determining the relationship between turgor pressure and cell volume (and thus tissue texture), where rigid inelastic walls allow minimal volume change per unit turgor change while elastic walls permit substantial turgor-driven expansion, with fundamental implications for growth, water relations, and stress adaptation. Cell wall elasticity in grapevine tissues reflects the composition and architecture of the primary wall, primarily cellulose microfibrils embedded in a pectin-hemicellulose matrix, where the degree of cross-linking, pectin methylesterification status, and microfibril orientation determine directional elastic properties. The developmental changes in cell wall elasticity accompany differentiation, with walls in expanding cells maintaining elasticity through controlled loosening while maturing cells develop more rigid walls through increased cross-linking and in some tissues secondary wall deposition and lignification. During grape berry ripening, cell wall elasticity changes dramatically as pectin depolymerization and hemicellulose modification reduce wall integrity, contributing to the characteristic softening that defines véraison and subsequent ripening progression. Measurement approaches include atomic force microscopy providing nanoscale mapping of wall mechanical properties, cell pressure probe techniques measuring wall elastic modulus through pressure-volume relationships, and bulk tissue measurements from which wall contribution can be extracted through modeling. Understanding cell wall elasticity quality enables mechanistic interpretation of texture, prediction of growth potential, and analysis of the biochemical basis for developmental changes in tissue mechanical properties.
Cell_Wall_Elasticity_Quality specifically addresses the structural wall component's mechanical behavior, distinguishing from whole-cell or tissue elasticity that includes turgor contributions. This quality underlies the bulk elastic modulus measured in pressure-volume analysis and determines the biomechanical response to water deficit. The class supports interpretation of texture in terms of wall biochemistry and connects ripening-associated wall modification to mechanical property changes.
Wandelastizität
Wandmechanik
Zellwanddehnbarkeit
Zellwandelastizität
cell wall deformability
cell wall mechanical property
wall elastic modulus
wall extensibility
Cosgrove DJ (2005) Growth of the plant cell wall. Nature Reviews Molecular Cell Biology 6(11): 850-861 Ortega-Regules A et al. (2008) Anthocyanin fingerprint of grapes: environmental and genetic variations. Journal of the Science of Food and Agriculture 88(14): 2409-2421
Cell Wall Elasticity Quality
Cabernet Sauvignon in Napa Valley shows cell wall loosening during mid-late July to early August Year N+1 at BBCH 73-75, with expansin gene expression upregulated 6-12 fold, α-expansin protein accumulation in apoplast pH 4.8, xyloglucan endotransglucosylase activity increasing, wall yield threshold declining from 0.52 to 0.38 MPa, coordinated wall synthesis maintaining 0.18 micrometer thickness, enabling 3.4-fold cell volume increase, under warm climate conditions of 26-32°C temperature, typical of this late-season cultivar modifying walls for enlargement
Cell wall loosening represents the wall modification phase during initial cell enlargement when expansin proteins and other wall-modifying enzymes disrupt cell wall structure, wall plasticity increases enabling yielding under turgor pressure, coordinated loosening and synthesis maintain wall integrity while permitting expansion, and modified walls enable turgor-driven cell volume increase, proceeding over weeks 3-5 as essential enabling mechanism for enlargement. This wall modification phase occurs during mid-late summer in mid-late July to early August in temperate regions Year N+1, proceeding during initial enlargement weeks 3-5 post-fertilization, corresponding to BBCH 73-75 pea-sized to berry touch stages. Wall loosening mechanism involves expansin protein expression and secretion disrupting wall structure, xyloglucan endotransglucosylase rearranging wall polymers, pectin methylesterase modifying pectin affecting wall properties, wall plasticity increase enabling yielding under turgor, coordinated wall synthesis maintaining thickness preventing failure, and integrated loosening-synthesis enabling controlled expansion. Expansin function in wall loosening shows α-expansin family members expressed during berry enlargement, expansin proteins secreted to apoplast binding to cell wall, non-hydrolytic disruption of hydrogen bonds between cellulose microfibrils and matrix polysaccharides particularly xyloglucans, wall loosening enabling turgor-driven expansion, and expansins as key wall-loosening enzymes. Expansin mechanism shows proteins binding to cellulose-xyloglucan interface, disruption of non-covalent bonds without polymer hydrolysis, wall loosening through bond disruption enabling polymer slippage, reversible binding allowing processive action, and mechanism enabling controlled wall plasticity increase. Expansin gene expression shows upregulation during transition to enlargement phase, expression levels correlating with expansion rate, transcript accumulation in expanding berry tissues, and expression as marker of active expansion. Expansin pH dependence shows optimal activity at acidic pH 4.5-5.5 characteristic of apoplast, acidic environment from cell wall pH typical 4.5-5.0, pH dependence ensuring activity in appropriate compartment, and pH regulation potentially controlling expansin activity. Xyloglucan endotransglucosylase XET function shows enzyme cutting and religating xyloglucan chains, transglycosylation rearranging xyloglucan cross-links, wall restructuring enabling expansion, XET activity correlating with growth rate, and enzyme complementing expansin action. XET mechanism shows enzyme cleaving xyloglucan backbone, transferring cleaved segment to new acceptor chain, rearrangement enabling wall relaxation and expansion, and mechanism providing dynamic wall restructuring. Pectin methylesterase PME function shows enzyme de-esterifying pectin removing methyl groups, demethylated pectin enabling calcium cross-linking potentially strengthening walls, PME action regulating wall mechanical properties, and enzyme affecting wall plasticity through pectin modification. PME complexity shows enzyme creating demethylated pectin blocks, calcium binding to demethylated regions creating egg-box structures, cross-linking potentially stiffening walls though effects complex, and PME as regulator of wall properties. Cell wall structure during loosening shows primary wall consisting of cellulose microfibrils embedded in matrix of hemicelluloses and pectins, cellulose providing tensile strength, matrix polymers interconnecting microfibrils, and structure modified through loosening enabling expansion. Cellulose microfibril characteristics show semi-crystalline cellulose polymers 3-5 nm diameter, microfibrils providing structural framework resisting tensile stress, microfibrils relatively stable during loosening, and framework maintained while matrix loosens. Hemicellulose matrix shows xyloglucans as primary hemicelluloses connecting cellulose microfibrils, xyloglucan hydrogen bonding to cellulose, bonds disrupted by expansin enabling slippage, and matrix as loosening target. Pectin matrix shows pectin polymers including homogalacturonan and rhamnogalacturonan contributing to wall properties, pectin de-esterification and calcium binding affecting properties, and pectin as regulatory component. Wall plasticity increase shows loosening reducing wall mechanical resistance to expansion, yield threshold decreasing enabling expansion at lower turgor, wall capable of plastic yielding under sustained pressure, and plasticity increase as essential expansion enabler. Wall stress-strain relationships show elastic region with reversible deformation, plastic region with irreversible deformation enabling growth, yield point separating elastic and plastic behavior typically 0.3-0.5 MPa turgor, and loosening reducing yield point enabling expansion. Wall synthesis during loosening shows new cellulose deposition maintaining microfibril density despite area increase, new hemicellulose and pectin synthesis, coordinated synthesis preventing excessive wall thinning, typical wall thickness maintained 0.1-0.3 micrometers during expansion, and synthesis essential for structural integrity. Synthesis-loosening coordination shows balanced processes enabling controlled expansion, excessive loosening without synthesis causing wall failure, excessive synthesis without loosening preventing expansion, and coordination as critical regulation point. Wall thickness maintenance shows wall area increasing substantially during expansion, new material deposition maintaining thickness despite area increase, thickness relatively constant 0.1-0.3 micrometers, and maintenance preventing wall rupture. Environmental effects on wall loosening include temperature affecting enzyme activities and gene expression with optimal 22-28°C supporting normal loosening, acidic pH in apoplast supporting expansin activity, and overall conditions affecting loosening rate. Temperature effects show warm conditions promoting faster expansin expression and activity, cool conditions slowing but not preventing loosening, and temperature affecting loosening kinetics. Wall loosening assessment methods include expansin gene expression analysis, enzyme activity assays in isolated walls, wall mechanical testing measuring yielding properties, and atomic force microscopy examining wall structure. The process represents third and final aspect of initial cell enlargement enabling expansion through wall modification, following turgor-driven expansion onset and vacuole enlargement initiation. Duration weeks 3-5 representing progressive wall modification during initial enlargement. Regional timing shows cell wall loosening during mid-late July to early August across climates during initial enlargement. Viticultural significance includes representing essential enabling mechanism for expansion, demonstrating enzymatic regulation of growth, showing wall modification-synthesis coordination, and enabling turgor-driven cell volume increase. Research applications include analyzing wall loosening mechanisms and regulation, investigating expansin and XET function and control, evaluating wall synthesis-loosening coordination, quantifying wall mechanical properties during expansion, and assessing wall loosening as determinant of expansion rate and capacity.
Cell_Wall_Loosening is third and final aspect of Initial_Cell_Enlargement, distinguished by wall modification enabling expansion. Represents expansin-mediated disruption of cellulose-hemicellulose bonds. XET rearranges xyloglucans, PME modifies pectin. Wall plasticity increases, yield threshold decreases. Coordinated synthesis maintains 0.1-0.3 micrometer thickness. Apoplast pH 4.5-5.5 optimal for expansin. Duration weeks 3-5. Follows Turgor_Driven_Expansion_Onset and Vacuole_Enlargement_Initiation. Completes Initial_Cell_Enlargement. Essential enabling mechanism for turgor-driven growth.
Expansin-vermittelte Lockerung
Wandmodifikation
Wandplastizitäts-Erhöhung
Zellwand-Lockerung
expansin-mediated loosening
growth-enabling wall changes
wall modification
wall plasticity increase
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 445-448 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Cell Wall Loosening
Syrah in Rhône Valley shows cell wall modification onset during early September Year N+1 at BBCH 83, with polygalacturonase activity increasing 5.2 fold from 9 to 47 units over 2-3 days at veraison, pectin methylesterase increasing 3.8 fold from 16 to 61 units, PG cleaving homogalacturonan pectin backbone, PME demethylating pectin from 68% to 52% methylation degree, pectate lyase possibly increasing 2.4 fold, coordinated enzyme activities initiating pectin depolymerization with molecular weight declining from 240 to 165 kDa, middle lamella pectin as primary degradation target, under warm Mediterranean conditions of 24-29°C temperature, typical of this mid-to-late season cultivar activating softening enzymes at veraison
Cell wall modification onset represents the structural alteration initiation phase during initial berry softening when pectin-degrading enzymes activate beginning cell wall pectin breakdown, polygalacturonase activity increasing 3-8 fold from pre-veraison levels initiating pectin backbone cleavage, pectin methylesterase activity increasing 2-5 fold demethylating pectin creating enzyme substrate, pectate lyase possibly activating, enzyme actions initiating pectin depolymerization and structural weakening, and cell wall modification onset as primary softening mechanism initiating at veraison. This enzyme activation and structural change initiation phase occurs during late summer to early autumn in late August-early September in temperate regions Year N+1, representing cell wall modification beginning at veraison onset. Modification onset mechanism involves polygalacturonase gene expression upregulation and protein accumulation during late lag, enzyme activation at veraison onset initiating pectin cleavage, pectin methylesterase similarly upregulated and activated, coordinated enzyme activities attacking pectin structure, pectin backbone cleavage and demethylation initiating structural breakdown, middle lamella pectin as primary target, and enzymatic modification initiating firmness decline. Polygalacturonase activation shows PG activity increasing from pre-veraison low levels typically 5-15 units to initial onset 15-40 units representing 3-8 fold increase, continued increase to higher ripening levels 40-100 units, PG as endo-polygalacturonase cleaving internal pectin bonds, and PG as major softening enzyme. PG gene expression shows PG genes strongly upregulated during late lag and veraison, multiple PG gene family members possibly expressed, transcriptional activation preceding enzyme activity increase, and gene expression as activation mechanism. PG protein accumulation shows PG protein synthesized from upregulated genes, protein accumulating during late lag providing enzyme capacity, accumulated protein activated at veraison, and protein as functional enzyme pool. PG enzyme characteristics shows endo-PG cleaving homogalacturonan backbone randomly, optimal pH 4.5-5.5 in apoplast, calcium effects on activity variable, and enzymatic properties affecting function. PG substrate shows homogalacturonan smooth pectin region as primary substrate, PG attacking pectin backbone creating smaller fragments, cleavage reducing polymer size, and substrate accessibility as rate factor. Pectin methylesterase activation shows PME activity increasing from pre-veraison 10-25 units to onset 25-50 units representing 2-5 fold increase, PME demethylating pectin removing methyl ester groups, demethylation creating blocks of free carboxyl groups, and PME as pectin modification enzyme. PME gene expression shows PME genes upregulated during late lag preparation, multiple PME isoforms possibly expressed, expression coordinated with softening, and transcription as activation basis. PME enzyme characteristics shows PME removing methyl groups from galacturonic acid residues, PME creating demethylated pectin blocks, optimal pH 6.5-8.0 though functional across range, and enzymatic properties affecting activity. PME dual function shows initial demethylation possibly creating calcium cross-links temporarily increasing firmness in some systems, subsequent demethylation creating PG substrate enabling degradation, biphasic PME role possible though in grapevine primarily degradation facilitation, and complex PME effects. Demethylated pectin characteristics shows free carboxyl groups enabling calcium binding, demethylated regions accessible to PG cleavage, degree of methylation declining from 60-80% pre-veraison to 40-60% onset, and demethylation as structural change. Pectate lyase possible activation shows PL possibly increasing activity at veraison, PL cleaving pectin through β-elimination mechanism different from PG hydrolysis, PL creating unsaturated products, contribution varying by cultivar, and PL as additional enzyme. Coordinated enzyme activities shows PG, PME, and possibly PL working synergistically, PME creating PG substrate, PG cleaving demethylated pectin, coordinated action maximizing degradation, and enzyme cooperation as efficient mechanism. Pectin depolymerization initiation shows pectin chain length declining from enzyme cleavage, molecular weight beginning reduction from 150-300 kDa to 100-200 kDa during onset, depolymerization creating smaller pectin fragments, and chain shortening weakening structure. Pectin structural integrity loss shows pectin network in cell wall and middle lamella beginning breakdown, structural pectin framework fragmenting, integrity loss reducing wall strength, and degradation as softening basis. Middle lamella as primary target shows pectin-rich middle lamella particularly susceptible to enzyme attack, middle lamella between adjacent cells containing high pectin content, middle lamella degradation weakening cell-cell adhesion most effectively, and targeting as strategic degradation. Cell wall pectin also targeted shows pectin in primary cell walls also degrading, wall pectin contributing to firmness, wall and middle lamella degradation combined reducing firmness, and comprehensive pectin targeting. Pectin domain specificity shows enzymes possibly preferentially attacking specific pectin domains, smooth homogalacturonan regions primary PG target, rhamnogalacturonan regions possibly resistant, and domain selectivity affecting degradation pattern. Cell wall other polysaccharides shows hemicelluloses and cellulose largely unaffected during initial softening, cellulose framework maintaining structural integrity, pectin as specific softening target, and selective degradation as mechanism. Calcium dynamics shows calcium possibly released from demethylated pectin as degradation proceeds, calcium cross-links breaking, calcium redistribution possible, and calcium affecting pectin structure. Modification onset timing shows activation occurring at veraison onset coinciding with other transformation changes, enzyme activation following late lag preparation period, rapid activation over 1-3 days, and timing as transformation feature. Spatial modification pattern shows modification possibly initiating in specific berry regions, mesocarp cells experiencing softening, progression through berry tissues, and spatial pattern affecting firmness. Environmental effects on modification onset include temperature affecting enzyme activities with warm conditions promoting activation and faster modification, cool conditions slowing enzyme kinetics, and temperature as major rate factor. Temperature effects shows optimal enzyme activities at 20-30°C, suboptimal temperatures reducing rates, and temperature affecting modification progression. The process represents first aspect of initial berry softening establishing enzyme activation, preceding pectin solubilization start and turgor reduction initiation. Duration enzyme activation initiating over 1-3 days at veraison onset then continuing. Regional timing shows cell wall modification onset during late August-early September at veraison. Viticultural significance includes representing primary softening mechanism through pectin degradation, demonstrating enzyme regulation as developmental control, determining softening rate and progression, and establishing cell wall modification as softening basis. Research applications include analyzing pectin-degrading enzyme regulation and activation, investigating enzyme-substrate interactions and degradation mechanisms, evaluating enzyme gene expression and protein dynamics, quantifying pectin structural changes, and assessing cell wall modification as softening determinant.
Cell_Wall_Modification_Onset is first aspect of Initial_Berry_Softening, distinguished by enzyme activation. Represents 1-3 days veraison onset. Polygalacturonase 3-8 fold increase (5-15 → 15-40 units), pectin methylesterase 2-5 fold (10-25 → 25-50 units). PG cleaves homogalacturonan, PME demethylates creating PG substrate. Pectate lyase possible. Pectin depolymerization initiating (150-300 → 100-200 kDa). Middle lamella primary target. Temperature affects enzyme kinetics (optimal 20-30°C). Precedes Pectin_Solubilization_Start. Primary softening mechanism.
Enzym-Aktivierung
Pektin-Abbau-Beginn
Wand-Abbau-Initiierung
Zellwand-Modifikations-Beginn
enzyme activation
pectin degradation onset
structural modification start
wall breakdown initiation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 477-482
Cell Wall Modification Onset
Cabernet Sauvignon cell wall thickened epidermal layer showing characteristic thick wall structure= outer periclinal walls 4-5.5 μm thickness (8-11 fold thicker than middle pericarp cell walls at 0.5 μm), comprising outer cutin-impregnated zone (1.2-1.8 μm), middle suberin lamellae zone (2-2.8 μm showing 8-12 alternating light/dark bands in TEM indicating layered suberin structure), inner cellulose-hemicellulose zone (0.8-1.2 μm), creating substantial cellular barrier component complementing 4.5-6.5 μm acellular cuticle for total outer barrier 8.5-12 μm thickness providing robust protection. Syrah cell wall thickened epidermal layer wall composition analysis= outer walls containing 35-42% cutin, 28-38% suberin, 18-25% cellulose/hemicellulose, 3-8% lignin (dry weight basis), versus middle pericarp walls containing 78-85% cellulose/hemicellulose, 8-15% pectin, 2-5% lignin, essentially no cutin/suberin, demonstrating fundamental compositional differences between protective cell wall thickened epidermal walls and structural middle pericarp walls reflecting different functional requirements. Merlot cell wall thickened epidermal layer pathogen penetration resistance= Botrytis cinerea requiring 6-12 hours to penetrate thick epidermal outer walls (3.5-5 μm) after cuticle penetration through secretion of cell wall degrading enzymes (endopolygalacturonase, cellobiohydrolase), versus <2 hours to penetrate thin-walled middle pericarp cells (0.5-0.8 μm walls), demonstrating that cell wall thickened epidermal layer creates substantial time delay for pathogen penetration allowing berry defense responses (phytoalexin synthesis, cell wall reinforcement) to activate before pathogen reaches internal tissues. Chardonnay cell wall thickened epidermal layer showing moderate thickness= outer walls 2.5-4 μm (thinner than typical red cultivars 3.5-6 μm), providing moderate rather than strong barrier, potentially contributing to white cultivar vulnerability to Botrytis bunch rot as thinner epidermal walls penetrated more rapidly by fungi, requiring enhanced cultural disease management (canopy air circulation, cluster exposure) compensating for less robust cell wall thickened epidermal barrier. Pinot Noir cell wall thickened epidermal layer sun exposure effects= sun-exposed berries showing outer epidermal walls 4.5-5.8 μm thickness with enhanced suberin content (38-45% of wall versus 28-35% in shaded) shaded berries showing 3.2-4.5 μm walls with lower suberin, demonstrating UV/light induction of epidermal wall thickening and modification creating enhanced cell wall thickened epidermal layer in exposed berries providing better barrier properties potentially contributing to superior disease resistance and stress tolerance observed in well-exposed fruit.
A cell wall thickened epidermal layer is an epidermal layer emphasizing the characteristic thick heavily modified outer periclinal cell walls (typically 3-6 μm thickness) that distinguish epidermal cells from underlying pericarp cells, with outer wall modifications including cutinization, suberization, and wall reinforcement creating enhanced barrier properties contributing fundamentally to berry protection against water loss, pathogen penetration, and mechanical damage. Cell wall thickened epidermal layer wall structure= (1) outer periclinal walls 3-6 μm thick (versus 0.5-1.5 μm for inner periclinal and anticlinal walls, and 0.4-0.8 μm for middle pericarp walls), representing 5-10 fold thickening of outer wall relative to other cellular walls (2) outer wall multilayer organization comprising outer cutin-impregnated zone (0.8-2 μm depth where cutin from overlying cuticle penetrates into wall), middle suberin-containing zone (1-3 μm with characteristic suberin lamellae), and inner cellulose-hemicellulose framework zone (0.5-1.5 μm) transitioning to normal wall composition (3) anticlinal walls moderately thickened (1-2 μm) providing lateral strength and cell-cell adhesion (4) inner periclinal walls relatively thin (0.5-1.5 μm) as protection not required on interior-facing surface. Cell wall thickened epidermal layer chemical modifications= (1) cutinization involving cutin deposition in outer wall creating hydrophobic barrier, with cutin (polyester of C16-C18 hydroxy fatty acids) impregnating wall matrix and creating continuous hydrophobic pathway from cuticle into wall (2) suberization involving suberin deposition creating additional hydrophobic barrier, with suberin (complex polyester containing both aliphatic and phenolic domains) organized in characteristic lamellae visible by TEM as alternating electron-dense and electron-lucent bands (3) lignification in some cases with lignin deposition in outer walls creating additional rigidity and resistance to enzymatic degradation (4) enhanced cross-linking of wall polysaccharides creating stronger more resistant wall structure. Cell wall thickened epidermal layer barrier functions= (1) water barrier contributing to reduced cuticle-plus-epidermis transpiration (0.8-2 g H₂O m⁻² h⁻¹ total) with thick hydrophobic epidermal wall providing significant barrier in addition to cuticle (2) pathogen barrier requiring fungal and bacterial pathogens to produce cell wall degrading enzymes (cellulases, pectinases) in addition to cutinases to penetrate thick epidermal walls, with thicker walls providing more material requiring degradation creating longer penetration time allowing defense responses (3) mechanical protection with thick walls providing structural strength resisting compression during berry packing in clusters, abrasion during harvest and handling, and puncture from insect stylets or bird beaks (4) chemical barrier with hydrophobic wall modifications limiting diffusion of aqueous solutions into or out of berry creating barrier to leaching and uptake. Cell wall thickened epidermal layer development= outer wall thickening initiating during early berry development with progressive deposition of wall material and wall modifications continuing through ripening cutin and suberin deposition beginning during Phase I and continuing through Phase II with maximum outer wall thickness achieved at or near maturity wall thickness potentially showing cultivar variation with thick-skinned cultivars sometimes showing thicker epidermal outer walls than thin-skinned cultivars environmental modulation with sun exposure potentially enhancing wall thickening and modification as stress response. Cell wall thickened epidermal layer mechanical properties= tensile strength and elastic modulus higher than thin-walled parenchyma cells due to thick walls wall stiffness affecting berry firmness particularly skin toughness perception wall brittleness versus flexibility affecting berry splitting susceptibility with more flexible walls better accommodating turgor changes without rupture.
Cell_Wall_Thickened_Epidermal_Layer emphasizes the characteristic thick modified outer walls of epidermal cells and is distinguished from Outer_Epidermal_Layer which emphasizes surface and environmental interface aspects. Cell wall thickened epidermal layer represents the epidermis from the perspective of barrier properties and wall specializations. Understanding cell wall thickened epidermal layer structure and composition is essential for comprehending berry barrier mechanisms, pathogen resistance, mechanical protection, and for managing epidermal development through environmental manipulation (light exposure, water management) affecting wall thickness and modification enhancing berry protection.
Barriere-Epidermis
barrier epidermis
dickwandige Epidermis
thick-walled epidermis
zellwandverdickte Epidermis-Schicht
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 176-180 Harris, J.M. et al. (1968). The Vinifera grape berry= anatomical study. Botanical Gazette 129=1-23 Esau, K. (1977). Anatomy of Seed Plants, 2nd ed., pp. 216-245.
Cell Wall Thickened Epidermal Layer
A 130-year-old Shiraz vine planted in Barossa Valley circa 1890 bears a Centenarian_Vine_Role, its survival on sandy phylloxera-resistant soils creating a living link to pre-Federation Australian viticulture—registered in the Historic Vineyard Society's Old Vine Registry and producing fruit for single-vineyard wines commanding $100+/bottle. A 115-year-old Zinfandel vine in Lodi's sandy Mokelumne River soils serves in a Centenarian_Vine_Role, its Italian immigrant planting heritage (1880s) documented through historical records and producing concentrated fruit for premium "Ancient Vine" designations. A 100-year-old Tinta Negra vine in Madeira's steep terraces bears a Centenarian_Vine_Role, own-rooted survival on volcanic soils preserving genetics from pre-phylloxera European viticulture now lost on the continent.
An old vine role that inheres in a grapevine that has survived 100 or more years since planting, representing exceptional longevity that makes such vines biological monuments, viticultural heritage, and sources of irreplaceable genetic and historical value. Centenarian vines are rare survivors of multiple challenges—phylloxera (in non-grafted regions or on tolerant soils), world wars, economic pressures, disease epidemics, and climate fluctuations—with their survival reflecting either exceptional site conditions (sandy soils preventing phylloxera establishment, favorable microclimate, appropriate variety-site match) or deliberate preservation against replanting pressure. These vines often grow in regions where phylloxera never established (sandy soils in parts of Australia, Chile, Portugal) or on own roots where sandy or other unfavorable soils prevented phylloxera survival. Centenarian vines are increasingly recognized through heritage registries (Historic Vineyard Society, Barossa Old Vine Charter) that document their location, variety, planting date (when known), and cultural significance. Fruit from centenarian vines commands exceptional premiums when separately identified, with wines marketed as "100-year-old vine" or equivalent designations achieving prices reflecting both rarity and perceived quality.
Centenarian_Vine_Role captures the exceptional age category where chronological survival itself becomes the defining characteristic, transcending ordinary old vine status to represent viticultural and cultural heritage. Documentation of centenarian status often requires historical research—planting records, property transfers, tax documents, photographic evidence—as the vines themselves carry no age markers beyond their venerable appearance. The geographic distribution of centenarian vines reflects phylloxera history most are either ungrafted survivors in phylloxera-free regions or grafted vines in areas where early rootstock adoption (1890s-1900s) has allowed 100+ year survival on resistant roots. Centenarian vines are increasingly protected through heritage designation, though no regulatory framework prevents their removal their preservation depends on owner recognition of their value and the premium production economics that justify continued cultivation despite reduced yields.
100-year-old vine
Centennial-Rebe
Erbe-Rebe
Jahrhundertrebe
centenarian vine
century vine
heritage vine
hundertjährige Rebe
Historic Vineyard Society (2020) Old Vine Registry Barossa Grape and Wine Association (2009) Barossa Old Vine Charter Bonada M et al. (2020) Vine age and grape composition, Australian Journal of Grape and Wine Research 26: 139-152 Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane
Centenarian Vine Role
Merlot in Bordeaux shows central cell fertilization during mid-late June Year N+1 at BBCH 65-69, with second sperm cell fusing with central cell containing fused polar nuclei within 1-4 hours post-discharge occurring 28-42 hours post-pollination, nuclear fusion producing triploid 3n=57 endosperm nucleus with 2 per 1 maternal per paternal ratio, free nuclear division initiating rapidly producing multinucleate syncytium, under oceanic conditions of 22-27°C temperature, typical of this mid-season cultivar with 54-68% central cell fertilization success coordinated with egg fertilization enabling seed formation
Central cell fertilization represents the endosperm-forming fusion event in double fertilization where second sperm cell released from pollen tube fuses with large central cell occupying embryo sac center, combining haploid sperm nucleus with diploid central cell nucleus formed from polar nuclei fusion or fusing separately with two haploid polar nuclei, forming triploid 3n endosperm nucleus with 2 per 1 maternal per paternal genome ratio, and activating endosperm developmental program initiating nutritive tissue formation supporting embryo growth. This critical endosperm-forming fusion occurs during double fertilization in mid-late June in temperate regions Year N+1, typically completing within 1-4 hours following sperm release approximately 24-48 hours after initial pollination, corresponding to BBCH 65-69 fertilization stages. Fertilization mechanism involves second sperm cell migration from tube discharge site toward large central cell, sperm-central cell membrane fusion creating cytoplasmic continuity, sperm nucleus entry and migration toward central cell nucleus or polar nuclei, nuclear fusion combining sperm nucleus with fused diploid central cell nucleus or separate fusion with two haploid polar nuclei, triploid 3n endosperm nucleus formation containing two maternal and one paternal genome copies, and central cell activation triggering endosperm developmental program. Central cell structure at fertilization shows large cell occupying majority 60-80% of embryo sac volume, central cell nucleus formed from two polar nuclei fusion creating diploid 2n nucleus or polar nuclei remaining separate until sperm fusion, abundant cytoplasm providing resources for endosperm development, and plasma membrane receptive to sperm fusion. Polar nuclei characteristics show two haploid nuclei derived from megagametophyte development through mitotic divisions, nuclei migrating from chalazal and micropylar poles toward embryo sac center, fusion occurring before or during fertilization creating diploid central cell nucleus, and nuclei positioned centrally or remaining separated until sperm arrival. Sperm cell approach to central cell shows migration through embryo sac cytoplasm from synergid discharge site, longer travel distance 50-200 micrometers compared to egg fertilization reflecting central cell position, passive drift or directed movement reaching central cell, and arrival typically concurrent with or shortly following egg fertilization enabling coordinated double fertilization. Sperm-central cell fusion shows membrane contact and fusion similar to egg fertilization, cytoplasmic integration enabling sperm nucleus entry, and fusion possibly mediated by similar molecular recognition mechanisms as egg fertilization though poorly characterized. Nuclear fusion pattern shows two possible scenarios with sperm fusing with pre-fused diploid 2n central cell nucleus creating triploid 3n endosperm nucleus, or sperm fusing sequentially or simultaneously with two separate haploid 1n polar nuclei creating 3n nucleus, and fusion pattern variation possibly species-specific or individual-specific though outcome identical. Triploid endosperm nucleus formation shows combination of haploid sperm nucleus containing paternal genetic material with diploid central cell nucleus or two haploid polar nuclei containing maternal genetic material, resulting 3n nucleus with 38 chromosomes in grapevine comprising 19 paternal and 38 maternal chromosomes creating 2 per 1 maternal per paternal genomic ratio, and genomic imbalance affecting endosperm development and gene expression patterns. Genomic imbalance significance shows 2 per 1 ratio creating unique genomic composition with maternal genome overrepresentation, imbalance affecting gene dosage and expression particularly for imprinted genes, maternal genome dominance potentially influencing endosperm developmental characteristics and nutritive function, and evolutionary significance of endosperm genomic composition affecting seed development strategies. Genomic imprinting in endosperm shows parent-of-origin-specific gene expression with some genes expressed preferentially from maternal genomes and others from paternal genome, imprinting affecting endosperm growth and resource allocation, and imprinting phenomena important for reproductive isolation and hybrid seed development though poorly characterized in grapevine. Central cell activation following fusion shows developmental transition from quiescent to active proliferative state, metabolic activation with increased respiration and biosynthesis, and initiation of free nuclear division producing multinucleate endosperm. Free nuclear division in endosperm shows rapid mitotic divisions producing multiple nuclei without cell wall formation, syncytial tissue development with nuclei sharing common cytoplasm, division rate often exceeding embryo division creating abundant nuclei, typical progression to 32-128 nuclei before cellularization in some systems though timing variable, and free nuclear phase creating nutritive tissue supporting early embryo development. Cellularization process shows eventual cell wall formation partitioning endosperm into cellular tissue, cellularization timing varying among species occurring after free nuclear phase, cellular endosperm providing structured nutritive tissue, and endosperm cells accumulating storage reserves including proteins, lipids, and carbohydrates. Endosperm function shows nutritive role providing resources supporting embryo development throughout seed maturation, tissue serving as intermediary transferring maternal resources from integuments to developing embryo, and endosperm consumption during embryo growth reducing to thin layer in mature grapevine seed. Endosperm-embryo coordination shows developmental synchronization between tissues with coordinated cell division and growth rates, signaling between tissues regulating resource allocation and development, and balanced development ensuring adequate endosperm support throughout embryogenesis. Calcium signaling in central cell fertilization possibly shows activation mechanisms similar to egg with calcium transients triggering developmental programs, though central cell calcium dynamics less well characterized than egg, and calcium-dependent processes regulating endosperm development initiation. Environmental effects on central cell fertilization include temperature affecting fusion dynamics and activation with optimal 22-28°C supporting efficient processes, similar sensitivity to egg fertilization with coordinated temperature requirements, and overall favorable conditions essential for successful endosperm formation. Temperature sensitivity shows warm conditions promoting rapid fusion and activation, cool conditions potentially delaying endosperm formation, excessive heat impairing processes, and temperature effects coordinated with egg fertilization requirements. Failed central cell fertilization causes include sperm cell dysfunction preventing fusion, abnormal central cell or polar nuclei preventing fusion, failed nuclear fusion preventing endosperm formation, or environmental stress disrupting processes, and failure causing seed abortion even if egg fertilization succeeds due to lack of embryo nutritive support. Central cell fertilization success rate shows similar 50-70% success as egg fertilization in favorable conditions, coordinated success of both fertilizations required for viable seed development, and dual fertilization requirement emphasizing critical nature of double fertilization for seed formation. The process represents second component of double fertilization creating endosperm lineage, occurring in parallel with egg fertilization. Duration typically 1-4 hours from sperm release to completed fusion similar to egg fertilization. Cultivar variation may include fusion efficiency, endosperm development characteristics, and genomic imbalance effects on seed traits. Regional timing shows central cell fertilization during mid-late June across all climates within hours of tube discharge 24-48 hours post-pollination coordinated with egg fertilization. Viticultural significance includes representing endosperm-forming fusion essential for seed development, creating nutritive tissue supporting embryo growth, demonstrating coordinated dual fertilization requirement, establishing central cell fertilization success as determinant of seed viability, and contributing to seed size and berry development through endosperm function. Research applications include analyzing molecular mechanisms of central cell fusion and activation, investigating endosperm development and free nuclear division, evaluating genomic imbalance effects on endosperm characteristics, quantifying coordination between egg and central cell fertilization, and assessing endosperm formation as determinant of seed and berry development outcomes.
Central_Cell_Fertilization is second component of Double_Fertilization, distinguished by endosperm-forming fusion creating nutritive tissue lineage. Represents sperm fusion with central cell or polar nuclei producing triploid 3n nucleus. Duration 1-4 hours parallel to Egg_Fertilization. Creates 2 per 1 maternal per paternal genomic ratio. Initiates free nuclear division and endosperm development. Essential for embryo nutritive support. Success rate 50-70% coordinated with egg fertilization. Both fertilizations required for viable seed. Genomic imbalance affects endosperm function.
Endosperm-Fertilisation
Spermium-Zentralzell-Fusion
Zentralzell-Befruchtung
endosperm fertilization
polar nuclei fusion
sperm-central cell fusion
triploid nucleus formation
triploide Kernbildung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 396-398 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 300-302
Central Cell Fertilization
Chardonnay grafted on Teleki 5C rootstock showing central columella cells in primary root cap, central 3 longitudinal files out of total 7 columella files positioned directly over root axis, central columella cells 48 micrometers diameter containing 78 amyloplasts per cell versus 32 in peripheral files, central amyloplasts 12.4 micrometers diameter versus 7.8 micrometers peripheral, iodine staining showing most intense starch accumulation in central files, reorientation experiment showing central amyloplast sedimentation complete within 8 minutes versus 18 minutes peripheral, central columella cells providing primary gravity detection
Central columella cells are columella cells positioned in centermost files directly over root axis and quiescent center, typically comprising 2-4 central longitudinal files out of total 4-8 columella files, characterized by most direct alignment with root growth axis, showing largest amyloplasts with highest starch density reflecting most intense gravity sensing function, demonstrating most rapid and complete statolith sedimentation in response to gravity, and representing primary gravity perception zone within columella where gravitropic signal generation most intense, with central columella cells exhibiting maximum amyloplast density potentially 50-100+ statoliths per cell compared to 20-40 in peripheral columella cells, showing preferential starch accumulation creating amyloplasts with starch content 80-95% by weight versus 60-80% in peripheral files, and demonstrating that centrally positioned cells provide most accurate gravity vector detection due to axial alignment and optimal statolith characteristics making central columella critical for precise gravitropic control. Central columella cells positional characteristics includes= showing axial alignment (central cells positioned directly along root central axis providing optimal geometry for gravity detection), exhibiting proximity to quiescent center (central columella cells closest to quiescent center overlying organizing region), demonstrating innermost position (central cells forming core of columella structure surrounded by more peripheral columella files), revealing symmetry center (central files defining center of radial symmetry in columella organization), and displaying optimal orientation (central position providing direct alignment with root growth direction and gravity vector). Central columella cells amyloplast characteristics includes= showing maximum abundance (central cells containing 50-100+ amyloplasts highest density in root cap), exhibiting largest size (central columella amyloplasts often 10-15 micrometers larger than peripheral columella amyloplasts), demonstrating highest starch content (central amyloplasts with 80-95% starch creating maximum density and sedimentation capacity), revealing optimal sedimentation (central amyloplasts showing most rapid and complete settling in response to gravity), and displaying densest packing (central cells with amyloplasts occupying large proportion of cytoplasmic volume maximizing sensing capacity). Central columella cells sensing function includes= providing primary gravity detection (central cells contributing disproportionately to total gravitropic signal generation), exhibiting highest sensitivity (central cells detecting smaller gravity stimulus or orientation changes than peripheral cells), demonstrating rapid response (central cell amyloplasts sedimenting most rapidly upon reorientation initiating signal cascade), revealing signal intensity (central cells generating stronger gravitropic signal per cell than peripheral columella cells), and displaying critical importance (central columella function essential for normal gravitropic response while peripheral potentially compensatory). Central columella cells developmental features includes= showing preferential starch accumulation (central cells accumulating starch earlier and more extensively during maturation than peripheral cells), exhibiting maintained identity (central position maintained through development creating stable central sensing zone), demonstrating genetic specification (central versus peripheral columella identity potentially genetically determined through positional signals), revealing maturation priority (central cells potentially maturing earlier or more completely than peripheral cells), and displaying stability (central columella cells maintaining function and structure throughout root growth period). Central columella cells experimental observations includes= showing selective ablation effects (specific ablation of central columella cells producing greater gravitropic impairment than peripheral cell ablation), exhibiting starch localization (histological staining showing most intense starch accumulation in central files documenting specialization), demonstrating mutation effects (gravitropism mutations often showing most severe effects on central columella cells), revealing imaging studies (live imaging showing central amyloplasts sedimenting most rapidly and completely), and displaying quantitative measurements (measurements showing central cells having highest amyloplast density and largest amyloplast size). Central columella cells signal transduction includes= showing calcium signaling (central cells potentially showing strongest calcium transients in response to gravity), exhibiting auxin effects (central cell gravity sensing potentially most directly affecting auxin redistribution), demonstrating mechanotransduction (central cell amyloplast sedimentation creating strongest mechanical signals on ER and cytoskeleton), revealing downstream signaling (signals from central cells potentially having priority or dominance in gravitropic response), and displaying integration (central and peripheral columella signals integrated but central potentially weighted more heavily). Central columella cells functional importance includes= determining gravitropic sensitivity (central columella characteristics determining overall root gravitropic responsiveness), enabling precise orientation (central cells providing accurate gravity vector information enabling precise downward growth), demonstrating evolutionary selection (central columella specialization reflecting strong selection for accurate gravity sensing), revealing developmental priority (central columella development and maintenance receiving developmental priority), and displaying adaptive significance (optimal central columella function providing competitive advantage through efficient soil exploration and resource access).
Central_Columella_Cells emphasizes centermost axially-aligned gravity sensors and is distinguished from peripheral columella by focus on central position and maximal sensing function. Critical for understanding axial alignment, maximum statolith density, optimal sedimentation, primary signal generation, positional specialization, and the centrally-positioned columella cells providing most intense and accurate gravity sensing determining gravitropic precision.
Axial columella cells
Axiale Statolithen-Zellen
Core gravity-sensing cells
Innermost statocytes
Kern-Gravitationssensoren
Zentrale Columella-Zellen
Morita, M. T. (2010). Directional gravity sensing in gravitropism. Annual Review of Plant Biology, 61, 705-720.
Central Columella Cells
Mourvèdre central heartwood at 65 years post-planting shows 35-55 mm radius from trunk center comprising innermost 8-12 annual rings (wood formed years 1-12, now aged 53-65 years), dark brown coloration from 7-11% phenolic compounds (darker than surrounding peripheral heartwood 3-6% phenolics), vessels completely occluded with heavily aged tyloses and gum deposits, extensive colonization by white rot fungi causing 40-60% density reduction (sound zones 0.58 g/cm³, decayed zones 0.25-0.35 g/cm³), central cavity 15-25 mm diameter from complete decay, but vine maintaining vigor through intact 18-24 mm sapwood ring providing all hydraulic and storage functions
Central heartwood is the innermost heartwood zone comprising the oldest annual rings in the trunk core including the pith region, characterized by maximum age, extensive vessel occlusion, potential advanced decay, and representing the first-formed secondary xylem that has undergone longest duration of heartwood aging and potential degradation. Central heartwood anatomical position includes= location at trunk center surrounding or incorporating pith (central parenchymatous core tissue from primary growth), comprising innermost 5-15 annual rings in mature vines (rings formed during years 1-15 post-planting now aged 10-50+ years depending on current vine age), radial extent typically 8-30 mm from trunk center (in mature to old vines), position maximally distant from functional cambium and sapwood (completely isolated from living tissue and physiological activity), and gradual transition to peripheral heartwood (no sharp boundary, central zone distinguished by greater age and often more advanced decay). Central heartwood represents oldest wood with maximum aging effects= wood formed 15-50+ years previously (depending on current vine age and growth rate), subjected to longest duration of heartwood processes (phenolic deposition, parenchyma degradation, vessel occlusion occurring over decades), showing maximum accumulation of extractives and phenolic compounds (condensed tannins, stilbenes, other secondary metabolites concentrating over time potentially reaching 5-12% dry weight), exhibiting darkest coloration (progressive phenolic deposition producing brown, reddish-brown, or grey hues darker than peripheral heartwood), and showing most advanced structural changes (cell wall modifications, lumen degradation, tissue compaction from aging). Central heartwood vessel occlusion characteristics include= complete and irreversible occlusion (tyloses, gums, resins, and phenolic deposits thoroughly blocking all lumens), aged tyloses potentially degrading or collapsing (decades-old occlusions showing structural breakdown), vessel walls heavily impregnated with phenolics (lignin and tannin deposits in cell walls), and zero hydraulic conductance (absolutely no water transport capacity contrasting with conducting sapwood and transitional inner sapwood). Central heartwood decay susceptibility includes= high vulnerability to wood decay fungi (oldest wood providing longest exposure time for fungal colonization), white rot fungi degrading cellulose and lignin (Fomes, Stereum, other Basidiomycetes colonizing central core), brown rot fungi attacking cellulose (leaving modified lignin residue), progressive density reduction from decay (sound central heartwood 0.52-0.66 g/cm³ declining to 0.25-0.45 g/cm³ in extensively decayed wood), and potential hollow formation (advanced decay in central heartwood creating cavity while maintaining peripheral heartwood and sapwood ring integrity). Central heartwood in old vine trunks includes= central heartwood comprising 30-50% of total trunk radius in old vines (in 30-50 year vines with 70-120 mm total radius, central heartwood 20-60 mm radius), extensive central decay common in ancient vines (50-100+ year vines frequently showing hollow centers from complete central heartwood degradation), hollow trunk formation allowing continued survival (sapwood ring maintaining function while central heartwood decays without necessarily compromising vine viability), and structural impact variable (minor decay leaving adequate support, extensive decay potentially requiring external trunk support or causing collapse risk). Central heartwood mechanical properties include= contributing to overall trunk strength when sound (central heartwood providing 20-45% of trunk cross-sectional area in mature vines contributing to compression and bending resistance), reduced contribution when decayed (decay-softened central wood providing minimal mechanical function), stress concentration effects (hollow centers or weakened cores potentially creating stress points increasing breakage risk), and secondary support from peripheral heartwood and sapwood (outer wood zones compensating for central weakness in decayed trunks). Central heartwood diagnostic characteristics include= darkest wood coloration in trunk cross-section (concentric darkening pattern with darkest zone at center in non-decayed wood), softer texture if decayed (decay-weakened central wood easily penetrated compared to sound peripheral wood), absence of structural integrity in advanced decay (central void or crumbly brown residue in extensively decayed hollow trunks), and incorporation of pith remnants (central parenchymatous pith tissue often degraded or incorporated into innermost heartwood). Central heartwood significance includes= indicating vine age and longevity (presence and extent of central heartwood reflecting decades of growth), representing original vine structure (innermost rings formed during establishment years preserving record of early development), vulnerability to decay affecting old vine stability (central heartwood decay compromising structural integrity in ancient vines), and potential hollow formation characteristic of very old vines (extensive central heartwood decay creating iconic hollow trunks in century-old vines maintaining survival through sapwood ring).
Central_Heartwood emphasizes innermost oldest wood zone and is distinguished from Heartwood (complete non-functional wood), Inactive_Heartwood (physiological shutdown emphasis), and Dense_Heartwood (structural quality emphasis) by focus on central position, maximum age, and advanced decay susceptibility. Critical for understanding hollow trunk formation in ancient vines, age-related decay patterns, structural implications of central wood degradation, and survival of very old vines through peripheral sapwood maintenance.
Core heartwood
Inner core
Innermost wood
Innerstes Holz
Stammkern
Zentrales Kernholz
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Pratt, C. (1974). Vegetative anatomy of cultivated grapes - a review. American Journal of Enology and Viticulture, 25(3), 131-150.
Central Heartwood
A central main axis inflorescence at node 3 on a shoot of Vitis vinifera cv. Syrah in the Rhône Valley, France, measuring 16.5 cm length with rachis positioned along the geometric central axis of the structure, rachis diameter of 3.6 mm at midpoint representing the central core, exhibiting clear bilateral organization with 13 pairs of lateral branches arising from opposite or near-opposite positions along the rachis creating symmetric or near-symmetric architecture, left-side lateral branches showing similar sizes and branching patterns to corresponding right-side branches (e.g., branch pair at 4.5 cm from base= left 3.8 cm, right 3.6 cm pair at 8.2 cm= left 2.9 cm, right 3.1 cm), overall structure showing balanced conical profile with rachis as the central spine and lateral branches radiating to create roughly circular cross-sectional profiles at multiple levels, rachis serving as the central vascular pathway with symmetric lateral trace departure to left and right branches, architectural organization creating relatively even distribution of flowers throughout the three-dimensional space, measured at BBCH stage 65 (full bloom) in mid-June, representing typical central main axis architecture with rachis-centered symmetric organization characteristic of standard inflorescence morphology.
A main axis primary inflorescence emphasizing the central rachis as the geometric and functional center of the architectural organization in Vitis vinifera, characterized by rachis position along the central axis of the inflorescence with lateral branches radiating symmetrically or near-symmetrically from the central core, clear bilateral or radial organization about the rachis centerline, rachis extending through the geometric center of the structure from base to apex, equal or approximately equal development of left and right lateral branches creating balanced architecture, central rachis serving as the axis of symmetry organizing the spatial arrangement of all branches and flowers, and functional role as the central vascular and mechanical core from which all peripheral structures derive support and resources. Central main axis inflorescences represent the typical architectural pattern emphasizing the rachis as the organizing centerline creating symmetrical or near-symmetrical structure, exhibiting geometric organization facilitating even light distribution and air circulation through the inflorescence, and developing through coordinated lateral branch formation maintaining bilateral balance during inflorescence morphogenesis.
Central main axis inflorescences emphasize the geometric and organizational centrality of the rachis, distinguished from simply having a main axis by the emphasis on symmetric radial or bilateral arrangement about the central core. The central designation highlights the rachis position as the geometric center and organizational axis rather than just the dominant structural element. Bilateral symmetry is typical with lateral branches arising in opposite or near-opposite pairs, though perfect symmetry is rare with most inflorescences showing subtle asymmetries. Radial organization creates approximately circular cross-sections when viewing the inflorescence perpendicular to the rachis axis, indicating balanced distribution of branches around the central core. The symmetric architecture results from coordinated development with lateral branch primordia initiating in balanced patterns maintaining bilateral organization, though developmental perturbations can create asymmetries. Functional implications include relatively even light penetration and air circulation facilitating uniform flower development and pollination, and balanced mechanical load distribution on the central rachis. Vascular organization shows symmetric lateral trace departure creating balanced resource distribution to left and right branches. Transformation to cluster maintains central organization with rachis as central cluster stem and symmetric berry distribution. Central main axis architecture represents the typical condition in most well-developed inflorescences, with departures toward asymmetry resulting from developmental irregularities, damage, or resource limitation affecting balanced growth.
bilateral Rachis-Infloreszenz
bilateral main axis inflorescence
centrally organized inflorescence
symmetric rachis inflorescence
symmetrisch organisierter Blütenstand
zentrale Hauptachse-Infloreszenz
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 179-212 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 216-248 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 395-428.
Central Main Axis Inflorescence
A central main rachis on a primary inflorescence of Vitis vinifera cv. Syrah in the Rhône Valley, France, positioned along the geometric central axis of the inflorescence from base to apex, measuring 16.8 cm length, exhibiting clear bilateral organization with 13 pairs of primary branches arising from opposite or near-opposite positions creating symmetric architecture, left-side branches showing similar sizes and positions to corresponding right-side branches (e.g., branch pair at 4.2 cm from base= left 4.5 cm, right 4.3 cm pair at 9.5 cm= left 3.8 cm, right 4.0 cm), central rachis diameter of 3.5 mm representing the central structural core, position through the geometric center evident when viewing inflorescence from multiple angles showing approximately equal distribution of lateral branches around the central spine, vascular organization showing symmetric lateral bundle departure with equal numbers and sizes of vascular traces departing to left and right branches, mechanical load distribution balanced with approximately equal masses on left and right sides of the central axis, overall architecture creating roughly conical profile with central rachis as the spine and lateral branches radiating to form circular to oval cross-sections at multiple levels, measured at BBCH stage 65 (full bloom) in mid-June, representing typical central main rachis architecture with geometric centrality and bilateral symmetric organization.
The main rachis emphasizing its position as the geometric and organizational center of the inflorescence structure in Vitis vinifera, characterized by location along the central axis of the inflorescence with lateral branches radiating symmetrically or near-symmetrically from the central spine, role as the axis of bilateral or radial symmetry organizing spatial arrangement of all branches and flowers, position through the geometric center of the inflorescence volume from base to apex, equal or approximately equal lateral branch development on left and right sides creating balanced architecture about the central axis, centrality determining the focal point for vascular distribution and mechanical load bearing, and functional significance as the central organizing feature coordinating three-dimensional inflorescence architecture. Central main rachis emphasizes the geometric centrality and organizational role of the main rachis as the symmetry axis, representing the central reference line about which the inflorescence structure is organized, exhibiting spatial position through the geometric center of the three-dimensional inflorescence volume, and serving as the organizing centerline facilitating balanced architectural development.
Central main rachis emphasizes the geometric and spatial aspects of the main rachis position, distinguished from primary main rachis (hierarchical emphasis) and thick main rachis (dimensional emphasis) by focus on centrality and symmetry. The central designation highlights the position through the geometric center of the three-dimensional inflorescence volume and role as symmetry axis. Bilateral symmetry is typical with lateral branches arising in opposite or near-opposite pairs creating balanced left-right organization, though perfect symmetry is rare with most inflorescences showing subtle asymmetries. Radial organization creates approximately circular cross-sections when viewing perpendicular to the rachis axis, indicating balanced distribution of branches around the central core. Geometric centrality means the central main rachis occupies the spatial center of the inflorescence volume with approximately equal distances from rachis to inflorescence periphery in multiple radial directions. Functional implications include balanced mechanical load distribution preventing bending or twisting from asymmetric loading, symmetric vascular distribution enabling even resource delivery to all regions, and regular spatial organization potentially facilitating even light penetration and pollination. Developmental coordination maintains bilateral balance through coordinated lateral branch initiation and growth creating the symmetric architecture. Transformation to cluster maintains central organization with the central stem occupying the cluster geometric center and symmetric berry distribution creating balanced cluster architecture. Central main rachis architecture represents the typical well-developed condition, with departures toward asymmetry resulting from developmental irregularities, damage, or resource limitations affecting balanced growth.
Symmetrieachse-Rachis
centrally positioned main rachis
geometric center rachis
geometrisch zentrierte Spindel
symmetry axis rachis
zentrale Hauptrachis
Based on Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 219-254 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 219-258 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 399-438.
Central Main Rachis
Cabernet Franc grafted on 3309 Couderc rootstock at 11 years showing central main root axis positioned 6 cm from trunk vertical axis (within 10 cm threshold for central classification), central axis 9.4 cm diameter at crown growing vertically downward maintaining axial alignment to 135 cm depth, lateral roots emerging from central axis at relatively uniform radial intervals creating balanced pattern with similar lateral density in all directions, excavation revealing symmetrical root system architecture organized around central axis, central position enabling balanced water uptake and equal anchorage providing optimal geometric organization, central main root axis serving as structural and functional core of root system
Central main root axis is main root axis positioned at or very near trunk center showing axial alignment and central spatial position, emphasizing location and geometric organization role rather than size or dominance, characterized by vertical or near-vertical growth directly beneath trunk center typically within 5-10 cm radius of trunk axis, serving as structural and spatial reference axis around which root system organized creating radial symmetry, providing balanced framework from which lateral roots emerge in all directions enabling uniform soil exploration, and demonstrating optimal geometric organization minimizing transport distances, balancing anchorage forces, and facilitating symmetrical root system architecture, with central position most pronounced in seedling vines showing single radicle-derived central root and potentially less distinct but still identifiable in cutting-derived vines where central primary root or roots form functional center despite multiple root origin. Central main root axis positional characteristics includes= located directly beneath trunk (central axis positioned at trunk center vertical projection typically offset <10 cm from perfect alignment), showing vertical or near-vertical orientation (central axis growing downward maintaining alignment with trunk axis rather than oblique), demonstrating axial continuity (central axis appearing as downward extension of trunk axis creating continuous central stem from canopy through trunk through central root into soil), maintaining central position with depth (central axis remaining near trunk axis as penetrates deeper not deflecting substantially laterally), and serving as spatial reference (central axis defining center point for describing root system spatial organization and architecture). Central main root axis symmetry role includes= providing radial organization axis (central axis serving as center around which lateral roots distributed in balanced radial pattern), enabling balanced lateral distribution (central position allowing lateral roots to emerge in all directions creating uniform soil exploration without directional bias), creating geometric efficiency (central axis position minimizing average distance from any lateral root tip through branching network through central axis to trunk optimizing transport), facilitating symmetrical architecture (central axis serving as symmetry axis for entire root system creating aesthetically and functionally balanced organization), and demonstrating optimal design (central axis position representing geometrically optimal solution for supporting above-ground vertical trunk structure). Central main root axis in root system function includes= serving as main transport conduit (central axis position providing shortest pathway for water and nutrients from lateral roots to trunk and canopy), enabling balanced resource delivery (central axis symmetry allowing equal resource acquisition from all radial directions maintaining balanced nutrition and hydration), providing central anchorage foundation (central axis deep vertical penetration creating strong central anchor complemented by radial lateral anchorage), storing substantial reserves (central axis typically large diameter due to central position and balanced resource supply enabling significant carbohydrate storage), and organizing hydraulic architecture (central axis serving as main trunk of below-ground hydraulic system from which lateral pathways branch analogous to above-ground trunk and branch hydraulic organization). Central main root axis development includes= originating from radicle in seedlings (seedling central axis naturally forming from downward-growing radicle establishing central position from germination), developing from central cutting roots in propagated vines (adventitious roots at cutting center or dominant central roots becoming functional central axis), establishing centrality early (central axis position determined during establishment phase and maintained throughout vine development), showing reinforced centrality (central position possibly reinforcing through feedback with balanced resource supply supporting continued central axis growth), and potentially shifting if damaged (central axis loss potentially resulting in another primary root assuming functional central position though less ideal alignment). Central main root axis versus tap root includes= potentially being same structure (central main root axis and central tap root often referring to same root emphasizing different attributes), showing overlapping but distinct concepts (central main root axis emphasizing position and framework role while tap root emphasizing vertical orientation and depth), demonstrating compatibility (root can be both central main root axis by position and tap root by vertical orientation simultaneously), revealing analytical perspective (describing root as central main root axis highlights architectural organization role while tap root designation emphasizes penetration function), and enabling complementary description (comprehensive root system description potentially using both terms for same root in different contexts). Central main root axis anchorage advantages includes= providing axial load bearing (central axis position optimal for transmitting vertical forces from above-ground structure directly downward into soil), enabling balanced radial support (central axis with radial laterals creating anchorage equally resisting forces from all horizontal directions preventing directional instability), minimizing bending moments (central axis alignment with trunk reducing eccentric loading and bending stresses at crown junction), facilitating wind resistance (central axis deep penetration combined with radial lateral anchorage providing strong resistance to wind-induced overturning moments), and supporting large vine structure (central axis optimal geometric organization enabling strong anchorage for extensive canopy and heavy crops). Central main root axis spatial optimization includes= minimizing transport distances (central axis position reducing average path length for water and nutrients from soil to canopy compared to off-center roots), balancing hydraulic resistance (central axis symmetry creating equal resistance pathways in all radial directions optimizing water distribution), enabling uniform soil exploitation (central axis providing equally accessible framework for lateral root expansion in all directions maximizing soil volume utilization), facilitating management efficiency (central axis known position near trunk simplifying irrigation emitter placement and fertigation targeting), and demonstrating engineering optimality (central axis architecture resembling engineered support structures with central vertical column and radial horizontal members representing mechanically efficient design).
Central_Main_Root_Axis emphasizes central position and symmetry role and is distinguished from Dominant_Main_Root_Axis (size emphasis) by focus on geometric organization. Critical for understanding radial symmetry, spatial optimization, balanced architecture, transport efficiency, central anchorage, and the centrally positioned structural axis providing geometric framework organizing entire root system around trunk center.
Achsiale Primärwurzel
Axial main root
Central structural root
Centrally positioned primary axis
Zentral positionierte Strukturwurzel
Zentrale Hauptwurzelachse
Smart, D. R., et al. (2006). Root system architecture in Vitis= Influences of rootstock, cultivar, and environment. American Journal of Enology and Viticulture, 57(4), 408-418.
Central Main Root Axis
A central median vein on a mature leaf at node 6 of Vitis vinifera cv. Syrah in the Rhône Valley, France, positioned along the blade's axis of bilateral symmetry dividing the 15.8 cm × 14.2 cm blade into left half (53% of area) and right half (47% of area) showing typical slight asymmetry, extending centrally from petiolar junction through the blade for 13.5 cm to the apex of the terminal lobe, positioned equidistant (within 2-3 mm) from left and right lateral margins when measured perpendicular to the median vein axis at multiple points, serving as the central line about which the five-lobed palmate architecture is organized with two pairs of major lateral lobes arranged bilaterally, appearing as the most prominent central feature on both abaxial surface (raised ridge 0.5 mm above surrounding tissue) and adaxial surface (depressed groove 0.2 mm deep), left lateral veins branching at mean angle 58 degrees while right lateral veins branch at mean 62 degrees showing typical slight asymmetry in branch angles, serving as reference axis for all morphological measurements including lobe width, sinus depth, and bilateral comparison parameters, representing the fundamental central organizing feature establishing leaf symmetry and spatial organization, measured at BBCH stage 81 (beginning of berry ripening) in early August.
The median vein considered in its positional context as the central axis of the leaf blade in Vitis vinifera, characterized by position along the line of bilateral symmetry dividing the blade into equal or approximately equal left and right halves, extension from petiolar junction centrally through the blade to the apex of the terminal central lobe, equidistant positioning between left and right lateral margins in idealized symmetric leaves though showing slight asymmetries in most actual leaves, role as the geometric center line about which lobe and sinus patterns are organized creating overall leaf symmetry, visibility as the most prominent central structure on both upper and lower leaf surfaces, and functional significance as the central organizing axis coordinating left and right lateral vein development and establishing overall bilateral architecture that characterizes grapevine leaf morphology. Central median vein emphasizes positional and geometric aspects, highlighting the median vein's role as the bilateral symmetry axis and central structural feature organizing the spatial arrangement of all other blade components including lateral veins, lobes, sinuses, and overall outline shape.
Central median vein emphasizes positional and geometric roles of the median vein as the bilateral symmetry axis rather than hierarchical vascular organization or functional primacy. The central designation highlights the median vein's position as the geometric center line dividing the blade into left and right halves and serving as reference axis for spatial organization. Perfect bilateral symmetry is rare in actual grapevine leaves, with most showing subtle asymmetries in left versus right blade areas, lobe sizes, lateral vein angles, and other features, but the central median vein serves as the organizational axis approximating symmetry. Asymmetry patterns show some consistency within cultivars, with certain varieties showing characteristic directional asymmetry (e.g., consistently larger left lobes) though environmental factors also contribute creating individual variation. The central position makes the median vein the most visible and recognizable leaf feature, often used as reference for orientation and measurement in morphological studies and ampelographic descriptions. Damage or developmental abnormalities affecting the central median vein can create unusual leaf shapes including split or bilobed configurations lacking normal central organization. The term "central" derives from positional aspects while "median" emphasizes the middle or center location, with combined usage reinforcing the central axis concept.
Symmetrie-Achsen-Vene
axial central vein
axiale Mittelrippe
central midrib
symmetry axis vein
zentrale Hauptader
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 120-165 Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 137-172.
Central Median Vein
Cabernet Franc grafted on 110 Richter rootstock at 38 years post-grafting showing central rootstock heartwood 4.8 cm diameter occupying innermost trunk core immediately surrounding pith, central heartwood darkest reddish-brown color in trunk showing maximum extractive accumulation from 28-30 years since heartwood formation (central rings formed years 1-10 converted to heartwood years 10-12), central heartwood specific gravity 0.66 g/cm³ highest in trunk from extended aging and compaction, central heartwood moisture content 38% lowest in trunk, central heartwood comprising annual rings 1-10 representing earliest trunk growth, central heartwood showing some irregular grain near pith typical of juvenile wood, no decay detected in central heartwood core indicating good structural condition, central heartwood surrounded by progressively lighter outer heartwood showing aging gradient from dark center (oldest) to lighter periphery (younger) to light tan sapwood (active)
Central rootstock heartwood is rootstock heartwood occupying the innermost trunk core position immediately surrounding pith representing the oldest most aged heartwood formed earliest in trunk development, typically comprising innermost 2-5 cm diameter depending on trunk age and growth rate, characterized by longest duration since formation potentially 15-40+ years in mature to old trunk, greatest extractive accumulation and darkest coloration from extended aging, highest density from progressive compaction and extractive deposition, and potentially showing age-related decay particularly in very old damaged trunks where central heartwood may develop hollow areas while maintaining essential structural function in peripheral sound wood. Central rootstock heartwood positional definition includes= located at absolute trunk center (central heartwood occupying innermost core position closest to pith representing trunk axis), surrounding pith directly (central heartwood forming ring immediately around central pith marking original stem development), occupying centermost 2-5 cm diameter (central heartwood typically limited to inner core though extent varying with trunk age and diameter), showing circular or slightly elliptical cross-section (central heartwood typically regular shape following pith position and early trunk symmetry), and bounded by outer heartwood (central heartwood core surrounded by progressively younger outer heartwood layers transitioning eventually to sapwood at periphery). Central rootstock heartwood as oldest heartwood includes= formed earliest in trunk development (central heartwood representing first sapwood converted to heartwood typically beginning years 8-15 post-grafting), showing longest time since formation (central heartwood aged 15-40+ years since conversion in mature to old trunk compared to 5-15 years for outer heartwood), comprising oldest annual rings (central heartwood containing trunk's earliest growth rings from years 1-10 post-grafting recording initial establishment period), demonstrating maximum aging effects (central heartwood showing most pronounced age-related changes from decades of time since active sapwood status), and representing most transformed tissue (central heartwood furthest removed from functional sapwood state with maximum extractive accumulation and structural modification). Central rootstock heartwood extractive accumulation includes= showing highest extractive content (central heartwood accumulating maximum phenolics, tannins, and other compounds from extended aging period), exhibiting darkest coloration (central heartwood typically darkest brown or reddish-brown from greatest extractive deposition over decades), displaying progressive darkening with age (central heartwood color intensifying over years as additional extractives deposit in oldest wood), demonstrating cultivar-specific maximum colors (central heartwood showing rootstock variety's characteristic deepest color from maximum extractive expression), and revealing aging gradient (central heartwood darkest at center lightening progressively outward toward younger outer heartwood and sapwood showing visual record of progressive aging). Central rootstock heartwood density characteristics includes= showing highest wood density (central heartwood specific gravity 0.54-0.70 g/cm³ typically highest in trunk from extractive deposition and compaction), exhibiting maximum compaction (central heartwood cells most compressed from decades of radial pressure as trunk expanded around core), displaying reduced lumen volume (cell cavities in central heartwood compressed and filled with extractives reducing void space), demonstrating progressive density increase (wood density highest at center declining progressively outward through outer heartwood to sapwood showing aging gradient), and contributing to structural strength (dense central heartwood providing excellent compressive strength and load-bearing capacity at trunk center). Central rootstock heartwood pith proximity includes= directly surrounding pith (central heartwood forming immediate ring around central pith), sharing pith formation timing (central heartwood wood produced in same early years as pith development years 1-3 post-grafting), showing pith-related characteristics (central heartwood potentially showing irregular grain, knots, or structural features associated with young trunk and pith region), demonstrating pith effects on anatomy (central heartwood wood anatomy potentially influenced by proximity to juvenile pith tissue during formation), and revealing early trunk history (central heartwood rings recording earliest growth conditions and establishment stresses in vine development). Central rootstock heartwood in trunk cross-section includes= occupying innermost 2-5 cm diameter (central heartwood core size depending on trunk age and when heartwood formation began), showing darkest color at center (progressive darkening toward pith creating visual gradient across heartwood zone), displaying oldest annual rings (innermost rings visible in central heartwood representing years 1-10 or so post-grafting), exhibiting ring width patterns (central heartwood rings often relatively wide reflecting rapid juvenile growth versus narrower rings in outer heartwood and sapwood from mature slower growth), and revealing structural features (central heartwood potentially showing juvenile wood characteristics including irregular grain or growth anomalies from young trunk). Central rootstock heartwood age-related changes includes= showing maximum aging duration (central heartwood aged decades since formation allowing extended time for physical and chemical changes), exhibiting progressive extractive increase (extractive content in central heartwood continuously increasing over years as compounds deposit and concentrate), displaying potential compressive deformation (central heartwood cells potentially slightly deformed from decades of radial compression as trunk diameter expanded), demonstrating possible micro-crack development (very old central heartwood potentially developing fine cracks from age-related stress and drying), and potentially showing decay initiation (oldest central heartwood particularly around pith potentially first site of decay in very old or damaged trunk). Central rootstock heartwood decay vulnerability includes= showing potential center rot (central heartwood in old trunk potentially developing decay creating hollow center while outer wood remains sound), exhibiting increased decay risk with age (very old central heartwood particularly in damaged trunks vulnerable to fungal attack despite extractive protection), displaying preferential decay location (decay often initiating in central heartwood around pith progressing outward rather than starting at trunk periphery), demonstrating variable decay resistance (some rootstock varieties showing excellent central heartwood durability while others more susceptible to center rot), and requiring damage assessment (hollow central heartwood potentially acceptable if outer wood sound but extensive decay potentially compromising structural integrity). Central rootstock heartwood in different trunk ages includes= absent in young trunk (no heartwood in years 1-7 with no central heartwood core), minimal in establishing trunk (small central heartwood core 1-2 cm diameter in years 8-15 just beginning to form), moderate in mature trunk (central heartwood core 2-4 cm diameter in years 15-40 representing oldest inner heartwood), substantial in old trunk (central heartwood 3-6 cm diameter in years 40-80 potentially showing age-related decay), and dominant in ancient trunk (central heartwood 4-8+ cm diameter in 80-100+ year trunk potentially hollow in center but surrounded by sound outer heartwood). Central rootstock heartwood moisture content includes= showing lowest trunk moisture (central heartwood 25-50% moisture typically lowest in trunk due to maximum age and compression), exhibiting stable low levels (central heartwood moisture relatively constant over season and years due to complete isolation from transport system), demonstrating minimal seasonal variation (central heartwood moisture essentially invariant unlike dynamic sapwood moisture), displaying slow equilibration (moisture changes in central heartwood occurring very slowly over months to years due to tight dense structure), and contributing to dimensional stability (low dry stable moisture in central heartwood minimizing swelling, shrinking, and warping). Central rootstock heartwood structural role includes= bearing axial compressive loads (central heartwood at trunk axis bearing direct downward compression from vine weight), resisting bending at neutral axis (central heartwood at trunk center contributing less to bending resistance than peripheral wood but still significant), providing core structural stability (central heartwood forming solid core preventing trunk buckling or collapse), supporting trunk even if partially decayed (sound outer heartwood maintaining structural capacity even if central heartwood develops hollow from decay), and enabling trunk longevity (central heartwood structural capacity supporting trunk for decades even in very old vine with aged core). Central rootstock heartwood color gradient includes= showing darkest color at pith (immediate pith vicinity typically darkest due to longest aging), exhibiting progressive lightening outward (central heartwood color gradually lighter moving from pith toward outer heartwood showing aging gradient), displaying cultivar-specific dark colors (central heartwood deepest browns or reddish-browns characteristic of rootstock variety maximum extractive expression), revealing visual aging record (color gradient across heartwood showing progressive age-related extractive accumulation from center outward), and contrasting with peripheral wood (central heartwood dark core contrasting with progressively lighter outer heartwood and light tan sapwood creating concentric color zones). Central rootstock heartwood in rootstock cultivars includes= Riparia types showing light to medium brown central heartwood (2-4 cm diameter core in mature trunk with moderate extractive content), Rupestris types displaying dark reddish-brown central heartwood (dense high-tannin core), Berlandieri hybrids exhibiting medium to dark brown central heartwood (variable extractive content depending on specific hybrid), vigorous rootstocks like 1103P showing large central heartwood cores (potentially 4-6 cm diameter in old trunk from early heartwood formation), low-vigor rootstocks like Riparia showing smaller central cores (2-3 cm diameter from slower growth and delayed heartwood formation), and cultivar-specific decay resistance (some varieties showing durable central heartwood while others prone to center rot in old age). Central rootstock heartwood extractive chemistry includes= containing maximum phenolic compounds (highest concentrations of various phenolic substances in central aged heartwood), showing abundant tannins (condensed and hydrolyzable tannins particularly concentrated in central heartwood creating astringency and antimicrobial properties), exhibiting oxidized compounds (central heartwood containing quinones and other oxidation products from extended aging), displaying cultivar-specific chemistry (extractive composition characteristic of rootstock variety reaching maximum expression in central heartwood), and demonstrating protective function (antimicrobial and antifungal properties of central heartwood extractives providing decay resistance when tissue intact). Central rootstock heartwood assessment includes= identifying by central position (innermost heartwood core immediately surrounding pith), recognizing by darkest color (deepest brown color indicating central oldest heartwood), measuring radial extent from pith (determining central heartwood diameter from trunk center outward), assessing condition through increment cores (sampling central heartwood checking for decay or structural integrity), and evaluating through cross-sections (examining cut trunk sections revealing central heartwood size, color, and condition). Central rootstock heartwood in hollow trunk development includes= potentially initiating trunk hollowing (decay in central heartwood around pith potentially creating hollow center in very old trunk), showing acceptable hollow cores (limited central decay potentially not compromising trunk function if outer heartwood and sapwood sound), demonstrating progressive decay outward (center rot potentially progressing from pith outward through central heartwood toward outer zones), requiring structural assessment (hollow central heartwood necessitating evaluation of remaining sound wood adequacy for trunk support), and indicating advanced trunk age (hollow centers typically appearing only in very old trunk 60-100+ years confirming ancient vine status). Central rootstock heartwood significance includes= representing oldest trunk tissue (central heartwood comprising earliest-formed longest-aged wood in trunk preserving record of initial establishment), showing maximum aging effects (central heartwood exhibiting most pronounced age-related physical and chemical changes from decades since formation), demonstrating extractive accumulation endpoint (central heartwood showing cultivar's maximum extractive deposition and darkest color from extended aging), providing core structural support (central heartwood forming solid inner core bearing axial loads at trunk center), potentially developing age-related decay (central heartwood in very old trunk showing vulnerability to center rot despite extractive protection).
Central_Rootstock_Heartwood emphasizes innermost core position and oldest most-aged heartwood and is distinguished from Rootstock_Heartwood (all non-functional inner wood) and Dense_Rootstock_Heartwood (density emphasis) by focus on central location, maximum age since formation, darkest color, and potential age-related decay. Critical for understanding oldest trunk tissue, maximum aging effects, extractive accumulation endpoint, pith proximity, center rot vulnerability, core structural role, and the innermost heartwood representing earliest trunk development preserved in aged central core.
Inner core rootstock heartwood
Innermost rootstock heartwood
Innerster Wurzelstock-Kernholzkern
Marknahes Unterlagenkernholz
Pith-adjacent rootstock heartwood
Zentrales Unterlagen-Kernholz
Central Rootstock Heartwood
Pinot Noir grafted on SO4 rootstock at 12 years showing central tap root positioned 4 cm from trunk vertical axis growing directly downward maintaining axial alignment, central tap root diameter 8.6 cm at crown extending 142 cm depth, lateral roots emerging radially from central tap root at relatively uniform intervals creating symmetrical pattern, excavation revealing balanced root distribution in all directions with similar lateral root density north, south, east, and west indicating central tap root serving as symmetry axis, central position providing balanced anchorage and uniform soil resource exploitation, central tap root typical of SO4 moderate architecture
Central tap root is tap root positioned at or very near vine trunk center axis showing axial symmetry in root system, emphasizing central spatial position and axial alignment rather than depth or vigor, characterized by vertical growth directly downward from trunk center typically within 5-10 cm radius of trunk axis, providing balanced radial root system architecture with lateral roots emerging in all directions creating symmetrical pattern, serving as structural and functional axis around which root system organized, and most pronounced in seedling-derived vines showing single radicle-originated central root while cutting-derived vines potentially showing multiple primary roots with deepest or most vertical designated central tap root functionally, with central position providing balanced water and nutrient uptake, equal anchorage in all directions, and symmetrical structural framework minimizing asymmetric stress on trunk base and optimizing root system efficiency through geometric organization. Central tap root positional definition includes= located at trunk center axis (central tap root positioned directly beneath trunk center typically within 5-10 cm radius of vertical axis), showing axial alignment (central tap root growing vertically along vine's main axis rather than obliquely from center), demonstrating radial symmetry (central tap root serving as axis around which lateral roots distributed in balanced radial pattern), maintaining central position with depth (central tap root remaining aligned with trunk axis as penetrates deeper not deflecting substantially to one side), and providing geometric reference (central tap root defining center point for describing root system architecture and spatial organization). Central tap root in seedling vines includes= derived from radicle directly (seedling central tap root originating from germinating seed radicle growing downward from germination site), showing single dominant axis (seedling typically producing one central tap root rather than multiple primaries), maintaining perfect axial alignment (radicle-derived central tap root naturally positioned along seedling axis from origin), demonstrating strongest centrality (seedling central tap root showing most pronounced central position and axial symmetry), and providing natural architecture (seedling root system naturally organized around single central tap root without artificial influences from propagation). Central tap root in propagated vines includes= developing from cutting center (central tap root position determined by central adventitious roots at cutting base or callus), potentially less distinct (cutting may produce multiple primary roots with central position less clear than single seedling radicle), designated functionally (deepest or most vertical root from cutting center often considered central tap root by position and function), showing variable centrality (cutting central tap root potentially somewhat offset from perfect axial alignment depending on adventitious root emergence pattern), and influenced by planting (cutting planting angle and orientation affecting which primary root becomes functionally central in established vine). Central tap root symmetry effects includes= enabling balanced lateral root distribution (central tap root axis allowing lateral roots to emerge in all radial directions creating uniform soil exploration), providing equal directional anchorage (central tap root with radial laterals anchoring vine equally in all directions preventing directional bias), facilitating uniform water uptake (central tap root symmetry enabling balanced water acquisition from all sides reducing localized depletion), optimizing resource exploitation (radially symmetrical root system around central axis efficiently exploring available soil volume without gaps or overlap), and minimizing trunk stress (central tap root position directly beneath trunk reducing asymmetric bending moments or shear stresses at crown). Central tap root structural role includes= serving as main structural axis (central tap root providing primary structural framework from which entire root system organized), bearing vertical loads (central tap root position along trunk axis optimal for transmitting vine weight and forces downward into soil), providing anchorage foundation (central tap root deep penetration creating fundamental anchor with lateral roots providing additional stabilization), maintaining trunk alignment (central tap root position helping maintain vertical trunk orientation preventing lean or tilt), and distributing mechanical stress (central tap root symmetry allowing balanced stress distribution without directional asymmetry). Central tap root functional advantages includes= optimizing transport efficiency (central tap root position minimizing path length from any soil location through lateral roots to central tap root to trunk), balancing source-sink relationships (central tap root symmetry enabling equal resource delivery from all radial directions to shoot system), providing drought resilience (central tap root with radial laterals maintaining some functional roots even if parts of root zone dry), enabling flexible adaptation (central tap root architecture allowing root system to adjust growth in any direction responding to localized soil variation), and facilitating management (central tap root position near trunk simplifying irrigation, fertigation, and soil management by providing known root system center). Central tap root assessment and identification includes= located through excavation (exposing crown and upper root system revealing which primary root most centrally positioned), identified by trunk alignment (determining which root grows most directly along vertical trunk axis), confirmed by lateral pattern (observing balanced radial lateral emergence indicating central tap root axis), measured by offset distance (quantifying displacement of tap root from trunk center typically <10 cm for truly central root), and characterized by symmetry (assessing whether root system shows radial symmetry indicative of central tap root architecture).
Central_Tap_Root emphasizes axial position and symmetry and is distinguished from other tap root types by focus on central location rather than depth. Critical for understanding radial symmetry, balanced architecture, structural axis, anchorage optimization, uniform resource exploitation, and the centrally positioned vertical root providing geometric organization framework for entire root system.
Achsiale Hauptwurzel
Axial tap root
Centrally positioned primary root
Symmetrieachsen-Wurzel
Symmetry axis root
Zentrale Pfahlwurzel
Central Tap Root
The certification authority for French grapevine clones is ENTAV (Établissement National Technique pour l'Amélioration de la Viticulture), based at Le Grau-du-Roi, operating under delegation from FranceAgriMer (the national agricultural market authority), responsible for maintaining the Conservatoire National des Clones de Vigne and issuing the agrément that authorizes clonal multiplication. The certification authority for Austrian grapevine material is the Bundesamt für Wein- und Obstbau at Klosterneuburg, which operates the national clone evaluation program, conducts mandatory virus testing, maintains the Austrian clone register (Österreichisches Klonregister), and issues certification in compliance with the Austrian Rebenverkehrsgesetz and EU harmonized standards. The certification authority for German clonal material is the Julius Kühn-Institut (JKI), Federal Research Centre for Cultivated Plants, in coordination with the Bundessortenamt and regional Landesanstalten.
Certification authority is a certification status entity that records the identity of the governmental agency, accredited institution, or designated competent body authorized to conduct phytosanitary testing, verify varietal and clonal identity, issue certification documents, and maintain the official register of certified clonal material within a national or regional jurisdiction, establishing the institutional provenance and regulatory legitimacy of the certification.
Certification authority documentation is essential for international traceability because different national authorities apply different testing protocols, maintain different virus panels, and operate under different regulatory frameworks, meaning that "certified" status from one authority may not be equivalent to certification from another. The EU harmonization directives establish minimum standards but permit member states to impose additional requirements — France mandates clonal identity verification through the ENTAV agrément system (which includes oenological evaluation beyond health testing), while some countries certify material solely on phytosanitary grounds. The accreditation of testing laboratories within certification authorities follows international standards (ISO 17025 for laboratory competence, EPPO diagnostic protocols for specific pathogens), providing quality assurance for the analytical results underlying certification decisions. In federated countries (Germany, Australia, USA), the relationship between federal framework legislation and state/regional implementing authorities adds complexity — German Länder issue certification based on JKI reference standards, Australian states operate through their respective departments of agriculture with coordination through the Australian Vine Improvement Association (AVIA). The increasing internationalization of the nursery trade requires growers and importers to understand which certification authority's standards apply to specific plant material and whether mutual recognition agreements facilitate cross-border acceptance of certification.
Anerkennungsstelle
Prüfstelle
Zertifizierungsbehörde
Zertifizierungsinstitution
accredited testing institution
certification agency
certification organization
certifying body
competent authority
zuständige Behörde
European Commission (2005) Commission Directive 2005/43/EC Boidron R et al. (1995) Catalogue des variétés et clones de vigne cultivés en France, ENTAV-INRA, pp.: 5-15 Walter B, Martelli GP (1997) Clonal and sanitary selection of the grapevine. INRA Editions, pp.: 43-95
Certification Authority
Certification date for Merlot clone 181 (ENTAV) records initial agrément on March 15, 1983, with subsequent renewal confirmations in 1993, 2003, and 2013 following mandatory decennial re-testing of foundation mother vines at ENTAV Le Grau-du-Roi — a 40-year continuous certification history demonstrating sustained phytosanitary compliance and enduring commercial relevance. Certification date for Zweigelt clone Zw-70 documents initial Austrian certification granted January 1, 1998, by the Bundesamt für Wein- und Obstbau, with re-certification upon expanded virus panel testing (including GRSPaV and GPGV added to the mandatory panel in 2015) confirmed on July 1, 2016, and current certification valid through December 31, 2026 pending the next scheduled re-evaluation.
Certification date is a certification status entity that records the calendar date or year on which official certification was formally granted to a clonal line by the competent authority, including the date of initial certification, dates of certification renewal upon periodic re-testing, and where applicable, dates of certification suspension or revocation, establishing the temporal validity and currency of the clone's certified status within the regulatory framework.
Certification date documentation serves both regulatory compliance and practical decision-making functions. The initial certification date establishes when a clone first became legally available for commercial propagation, providing context for its distribution history and the vintage range over which commercial performance data may exist. The interval between selection origin date and certification date quantifies the development timeline — typically 15-25 years — reflecting the thoroughness of the evaluation process. Certification renewal dates confirm ongoing phytosanitary compliance, as periodic re-testing addresses the risk of virus reinfection in foundation plantings through vector transmission. Gaps between renewal dates may indicate testing delays, administrative issues, or periods of temporary decertification that should be investigated. The certification date also has economic significance for nurseries, as it determines when new clonal material enters the commercial supply chain and when older certifications may expire if not renewed. Under some regulatory frameworks, certification dates trigger time-limited propagation rights — for example, certain PVP (Plant Variety Protection) or plant patent provisions restrict commercial multiplication to defined periods from the certification date. The retroactive application of expanded virus testing panels (e.g., the addition of GPGV to mandatory screening in the 2010s) can require re-certification of clones originally certified under earlier, narrower testing protocols.
Genehmigungsdatum
Registrierungsdatum
Zertifizierungsdatum
certification grant date
certification validity date
date of certification
registration date
European Commission (2005) Commission Directive 2005/43/EC Boidron R et al. (1995) Catalogue des variétés et clones de vigne cultivés en France, ENTAV-INRA, pp.: 15-25 Mannini F (2004) Clonal selection in grapevines. Acta Horticulturae 652: 21-27
Certification Date
Certification level for Cabernet Sauvignon clone 337 mother block at ENTAV Domaine de l'Espiguette is classified as "Matériel initial" (Initial/Pre-basic), the highest category in the EU framework, maintained through annual ELISA/PCR testing and periodic biological indexing, from which all downstream multiplication is derived. Nursery-propagated vines of clone 337 available to growers carry certification level "Matériel certifié" (Certified), representing the third multiplication generation from Initial material and meeting the minimum health standard required under EU Directive 68/193/EEC for commercial sale. In Austria, Blaufränkisch clone Bl-2 mother vines at the Bundesamt für Weinbau hold "Vorstufenpflanzgut" (Pre-basic) status, while commercially available grafted vines are distributed as "Zertifiziertes Pflanzgut" (Certified material).
Certification level is a certification status entity that records the hierarchical category or grade assigned to a clonal line within the applicable certification framework, designating the stringency of health testing, the number of multiplication generations from the original nuclear stock, and the permissible end use of the material, ranging from the most rigorously tested nuclear or pre-basic material through intermediate multiplication stages to the final certified or standard category available for commercial vineyard planting.
Certification level reflects a deliberate trade-off between phytosanitary stringency and practical availability — the highest levels (Nuclear, Pre-basic/Initial) apply the most rigorous and costly testing protocols to a small number of conserved mother vines, while successive multiplication generations (Basic, Certified) apply progressively reduced testing frequency to accommodate the volume requirements of commercial nursery production. The EU certification framework established by Directive 68/193/EEC and amended by Directive 2005/43/EC defines four material categories: Initial (virus-tested foundation stock), Pre-basic (first multiplication from Initial), Basic (second multiplication), and Certified (third multiplication for commercial distribution), with an additional "Standard" category permitting sale of varietal-true but non-health-tested material. National implementations add complexity — France requires ENTAV agrément for all multiplied clones (effectively excluding Standard category material), while some member states permit Standard material for traditional local varieties. The certification level directly determines the frequency and scope of mandatory re-testing, the maximum number of permitted multiplication cycles, and the documentation chain required from nursery to grower. Understanding certification levels is essential for growers making planting decisions, as material from higher certification levels carries stronger health guarantees but may be limited in availability and higher in cost.
Anerkennungsstufe
Pflanzgutkategorie
Qualitätsstufe
Zertifizierungsklasse
Zertifizierungsstufe
certification category
certification class
certification grade
certification tier
plant material category
European Commission (2005) Commission Directive 2005/43/EC on marketing of vine propagation material Mannini F, Digiaro M (2017) The effects of viruses and viral diseases on grapes and wine. In Grapevine Viruses: Molecular Biology, Diagnostics and Management, Springer, pp.: 453-482 Walter B, Martelli GP (1997) Clonal and sanitary selection of the grapevine. In Sanitary Selection of the Grapevine, INRA Editions, pp.: 43-95
Certification Level
Certification number for Pinot Noir clone 115 in the French ENTAV system is registered as "PN 115" in the Catalogue Officiel des Variétés et Clones, cross-referenced internationally as ENTAV-INRA 115 to distinguish it from identically numbered clones in other national systems. Certification number for Riesling clone 239 in the German system is registered as "Gm 239" (Geisenheim selection 239) in the Bundessortenamt clone register, while the same cultivar's Neustadt selection carries certification number "N 90" — the prefix indicating the selecting institution and preventing confusion between genetically distinct clones bearing similar numeric identifiers. In Austria, certification number AT-BL-002-1998 for Blaufränkisch clone Bl-2 encodes country code (AT), cultivar abbreviation (BL), sequential clone number (002), and certification year (1998) in a structured format facilitating database management.
Certification number is a certification status entity that records the unique alphanumeric identifier assigned to a certified clonal line by the certifying authority, serving as the official registration code within the national or international clone register, enabling unambiguous identification of the specific clone across nursery catalogs, planting records, regulatory documents, and scientific publications, and functioning as the primary key linking all certification documentation, test results, and performance data to a single defined genetic entity.
Certification number systems vary substantially between national authorities, creating potential confusion in international plant material trade and scientific communication. The French ENTAV system uses cultivar abbreviation plus sequential number (e.g., CS 337 for Cabernet Sauvignon clone 337), the German system prefixes numbers with institutional codes (Gm for Geisenheim, Fr for Freiburg, Wü for Würzburg, N for Neustadt), the Italian Rauscedo system uses cultivar-specific sequential numbering (e.g., R24 for Sangiovese), and the Australian CSIRO/SARDI system uses institution-specific alphanumeric codes. This nomenclatural heterogeneity means that "clone 1" of a given cultivar designates entirely different genetic material depending on the certifying authority — a source of persistent mislabeling problems in international nursery supply chains. The VIVC (Vitis International Variety Catalogue) database at the Julius Kühn-Institut provides cross-referencing of certification numbers across national systems, though coverage remains incomplete for newer and less commercially distributed clones. Some certification numbers have become de facto variety descriptors in commercial contexts — "Dijon clones" for the ENTAV Burgundy selections, "CTPS clones" for French officially catalogued material — blurring the distinction between certification identifiers and marketing terminology. The integrity of certification number assignment requires maintenance of centralized clone registers with rigorous deduplication procedures, as numbering errors can propagate through decades of nursery multiplication.
Anerkennungsnummer
Klonnummer
Registrierungsnummer
Zertifizierungsnummer
amtliche Klonbezeichnung
certification code
certified clone ID
clone registration number
official clone number
registration identifier
Boidron R et al. (1995) Catalogue des variétés et clones de vigne cultivés en France, ENTAV-INRA, pp.: 10-20 Maul E et al. (2012) The European Vitis Database — a technical innovation through an online uploading and interactive modification system. Vitis 51(2): 79-85 Regner F et al. (2003) Clonal selection and clone identification of Austrian grapevine varieties. Mitteilungen Klosterneuburg 53: 172-180
Certification Number
Certification status for Pinot Noir clone 115 (ENTAV) records agrément (official approval) at the highest French certification level (clone agréé) granted by FranceAgriMer in 1978, based on complete virus-free status confirmed through ELISA, RT-PCR, and biological indexing on the mandatory indicator panel, with certification number FR-PN-115 and health declaration authorizing unrestricted propagation and distribution through the French national nursery system. Certification status for Grüner Veltliner clone 17 documents Austrian Anerkennungsstufe "Zertifiziertes Pflanzgut" (certified plant material) issued by the Bundesamt für Weinbau, Klosterneuburg, certification number AT-GV-017-2005, based on virus-free status confirmed by ELISA and RT-PCR panels meeting EU Directive 2005/43/EC requirements for Category Initial propagation material.
Certification status is a clonal selection record entity that documents the official phytosanitary and genetic identity verification standing of a clonal line within a national or international plant material certification framework, encompassing the certification level attained, the certifying authority, the date and registration number of certification, the virus testing results upon which health status was determined, and the formal health declaration issued, collectively establishing the legal and commercial standing of the clone for nursery propagation and vineyard planting.
Certification status represents the regulatory gateway between clone evaluation and commercial availability, translating scientific assessment results into the legal authorization required for nursery multiplication and sale. The certification framework operates hierarchically in most jurisdictions — the EU system defines categories of Nuclear, Pre-basic, Basic, Certified, and Standard material with progressively relaxed health requirements at each level, while the USDA-APHIS framework uses Foundation, Registered, and Certified tiers with similar graduated standards. Certification status is not permanent — mother blocks and foundation plantings require periodic re-testing (typically every 3-5 years) to maintain certified status, and positive virus detection triggers decertification procedures. The mutual recognition of certification between countries is governed by bilateral agreements and international protocols (EPPO standards, IPPC guidelines), facilitating or restricting the international movement of certified clonal material. The economic implications of certification status are substantial — certified plant material commands price premiums of 30-100% over standard (uncertified) material, reflecting both the health guarantee and the decades of evaluation underlying the certification. The trend toward molecular identity verification (SSR profiling) as a mandatory component of certification addresses the persistent problem of mislabeled plant material in nursery supply chains.
Anerkennungsstatus
Klonzertifizierung
Pflanzgutzertifizierung
Zertifizierungsnachweis
Zertifizierungsstatus
certification documentation
certification record
certified status
clone certification
plant material certification
Mannini F (2004) Clonal selection in grapevines: interactions between genetic and sanitary strategies. Acta Horticulturae 652: 21-27 European Commission (2005) Commission Directive 2005/43/EC amending Annexes to Council Directive 68/193/EEC on the marketing of material for the vegetative propagation of the vine Maree HJ et al. (2013) Grapevine leafroll-associated virus 3. Frontiers in Microbiology 4: 82
Certification Status
Pinot Noir clone 115 propagation material at the Certified grade bears a Certified_Clone_Role, derived from Basic-category mother plants tested negative for GFLV, GLRaV-1, GLRaV-3, GVA, and GVB—the "Certifié" label distinguishing it from Standard-grade clone 115 material that meets only visual inspection requirements. Chardonnay clone 96 Certified-grade vines serve in a Certified_Clone_Role for premium vineyard establishment, with laboratory testing documentation providing assurance of virus-free status beyond the visual freedom guaranteed by Standard-grade material. A Cabernet Sauvignon clone 337 shipment with Certified grade labeling bears a Certified_Clone_Role, the enhanced health assurance justifying the price premium for growers prioritizing long-term vineyard health over initial planting cost.
An official clone role that inheres in a grapevine clone that meets the "Certified" grade requirements within the EU certification framework or equivalent international standards, confirming both official registration and compliance with enhanced phytosanitary testing requirements beyond baseline (CAC/Standard) levels. Certified clones derive from mother plants that have been laboratory-tested and confirmed negative for regulated pathogens (viruses, phytoplasmas) according to prescribed testing protocols, rather than relying solely on visual inspection for disease symptoms. The "Certified" designation on plant labels indicates that the material originates from tested mother plant sources maintained within the certification hierarchy, providing growers with enhanced assurance of health status compared to Standard/CAC grade material. Certified clone material commands premium prices (typically 15-30% above Standard grade) reflecting the additional testing investment and the reduced risk of introducing virus-infected plants into vineyards. The Certified grade represents the intermediate quality tier in most certification systems, above Standard/CAC but below the highest tier (often termed "nuclear" or "pre-basic" stock maintained by research institutions).
Certified_Clone_Role captures the testing-enhanced tier of the certification system where laboratory verification supplements the visual inspection underlying Standard/CAC grade material. The distinction between Certified and Standard grades reflects the limitation of visual inspection for detecting virus infections—many economically important viruses (particularly leafroll viruses) can be present asymptomatically or with symptoms too subtle for reliable visual detection, particularly in the nursery environment where plants are young and symptoms may not have developed. Certified grade material offers documented health assurance but cannot guarantee absolute virus freedom testing detects known pathogens targeted by the testing panel, but novel or untargeted agents may escape detection. The economic calculus favoring Certified grade material considers both the immediate cost premium and the long-term costs of virus-infected vineyards—reduced yield (20-40% for leafroll), shortened vineyard lifespan, and fruit quality impacts that may justify the certification premium many times over.
certified clone
certified grade clone
getesteter Klon
tested clone
virus-tested clone
virusgetesteter Klon
zertifizierter Klon
European Union (2020) Commission Implementing Directive 2020/177 Annex IV EPPO (2008) Certification schemes for pathogen-tested material of grapevine, EPPO Bulletin 38: 422-429 Maliogka VI et al. (2015) Grapevine virus diseases, APS Press Golino DA et al. (2017) Foundation Plant Services, HortScience 52: 1190-1194
Certified Clone Role
A Syrah clone 470 vine in a commercial French nursery's scion block bears a Certified_Mother_Plant_Category_Role, derived from ENTAV Basic material two generations prior, maintained under INRAE certification protocols with annual visual inspection and isolation from non-certified plantings—its budwood used for benchgraft production meeting the "Certified" grade requirement of documented clean origin. A Kober 5BB rootstock plant in a German nursery increase block serves in a Certified_Mother_Plant_Category_Role at the CAC (Standard) level, providing hardwood cuttings for commercial rootstock production under visual inspection protocols without mandatory laboratory testing. A Tempranillo clone RJ-51 mother vine in a Spanish vivero bears a Certified_Mother_Plant_Category_Role, meeting EU certification requirements for documented traceability and freedom from visual symptoms while supplying the Spanish wine industry's substantial annual planting material demand.
A certified mother plant role that inheres in a grapevine maintained at the commercial multiplication tier (Certified/Standard) of the certification hierarchy, derived from Basic material and serving as the direct source of planting material for vineyard establishment. Certified category mother plants operate under less stringent testing requirements than Basic or Pre-Basic tiers, typically relying on visual inspection for virus symptoms, documented traceability to tested source material, and isolation requirements rather than routine laboratory testing. Certified material represents the final multiplication stage before vineyard planting, with generation limits (typically 2-3 from Basic source) preventing extended propagation chains that would increase genetic drift and pathogen accumulation risk. The Certified category includes both the higher "Certified" grade and the baseline "CAC" (Conformitas Agraria Communitatis/Standard) grade in EU terminology, with Certified grade requiring negative laboratory tests for specified pathogens while CAC requires only visual freedom from symptoms and traceable origin from the certification system.
Certified_Mother_Plant_Category_Role represents the commercial endpoint of the certification system where tested material meets nursery multiplication demand. The distinction between "Certified" grade (laboratory-tested negative) and "CAC/Standard" grade (visually free, documented origin) provides market segmentation—quality-focused growers specify Certified grade for premium plantings, while CAC material serves cost-sensitive applications. The Certified tier's reliance on visual inspection creates vulnerability to latent virus infections (asymptomatic but transmissible) and to low-titer infections below visual detection thresholds this recognized limitation drives periodic proposals to strengthen Certified-tier testing requirements. Commercial nursery economics favor maximizing multiplication from each mother plant, creating tension with certification principles favoring conservative generation limits regulatory compliance and market reputation provide counterbalancing incentives for nursery investment in certification program adherence.
CAC mother plant
CAC-Material
Handelsvermehrungsstock
Standardkategorie-Mutterstock
certified category mother vine
commercial mother plant
standard propagation plant
zertifiziertes Vermehrungsmaterial
European Union (2020) Commission Implementing Directive 2020/177 Annex IV EPPO (2008) Certification schemes for grapevine, EPPO Bulletin 38: 422-429 Gramaje D, Armengol J (2011) Fungal trunk pathogens in the propagation process, Phytopathologia Mediterranea 50: S217-S226 Maliogka VI et al. (2015) Grapevine virus diseases, APS Press
Certified Mother Plant Category Role
A Pinot Noir Dijon clone 115 vine in the ENTAV foundation vineyard at Montpellier bears a Certified_Mother_Plant_Role at the Basic category, having been derived from virus-tested Pre-Basic material, maintained under isolation protocols, and subjected to triennial ELISA screening for regulated pathogens—its budwood eligible for distribution to licensed nurseries for Certified category multiplication. A Chardonnay clone 96 plant at Foundation Plant Services (UC Davis) serves in a Certified_Mother_Plant_Role at the Pre-Basic (Foundation) category, representing nuclear stock subjected to comprehensive pathogen elimination (thermotherapy, tissue culture) and ongoing testing for all known grapevine viruses. A SO4 rootstock mother vine in a German VCR-affiliated nursery's multiplication block bears a Certified_Mother_Plant_Role at the Certified category, providing cuttings for commercial rootstock production under CAC (Conformitas Agraria Communitatis) or higher certification levels.
A mother plant role that inheres in a grapevine maintained within an official certification scheme, meeting specified requirements for genetic identity (trueness-to-type verification), health status (freedom from listed pathogens as confirmed by prescribed testing), and traceability (documentation linking the plant to its source within the certification hierarchy). Certified mother plants operate within the three-tier EU framework (Pre-Basic, Basic, Certified categories) or equivalent national certification schemes, with each tier defined by testing frequency, pathogen tolerance thresholds, generation limits from nuclear stock, and permissible uses of derived material. Certification schemes aim to provide growers with propagation material of verified identity and health status, preventing the economic losses caused by virus-infected planting material (yield reductions of 20-60%, shortened vineyard lifespan, reduced fruit quality) and the dissemination of graft-transmissible pathogens through nursery trade. Participation in certification is mandatory for commercial nursery production in EU member states and increasingly required or encouraged in other viticultural regions globally.
Certified_Mother_Plant_Role is subclassed into Pre_Basic, Basic, and Certified categories reflecting the EU certification hierarchy that has become the global reference framework. The terminology varies internationally—"Foundation" in the US corresponds roughly to "Basic" in EU terminology, while "Pre-Basic" parallels "Nuclear" or "Elite" stock—but the underlying principle of a health status pyramid with progressively more stringent requirements at higher tiers is universal. Certification requirements have expanded over time as new pathogens are characterized contemporary protocols test for 40+ virus species, phytoplasmas, and increasingly fungal trunk pathogens. The economic value of certified material is demonstrated by yield comparisons showing 20-60% advantages over uncertified plantings, particularly in regions with high virus pressure.
certified mother vine
certified stock plant
registered mother plant
registrierte Mutterrebe
zertifizierte Mutterpflanze
zertifizierter Mutterstock
European Union (2020) Commission Implementing Directive 2020/177 Mannini F (2000) Clonal selection in grapevine, in Proceedings of the Meeting on Grapevine Clonal Selection, ISHS Acta Horticulturae 528: 17-25 Golino DA et al. (2017) Foundation Plant Services, HortScience 52: 1190-1194 Rowhani A et al. (2018) Grapevine virus testing and certification, in Compendium of Grape Diseases 2nd Ed, APS Press
Certified Mother Plant Role
Grüner Veltliner vines at Nikolaihof in the Wachau bearing a certified organic role have maintained continuous organic certification under EU Regulation 2018/848 since 1971, with annual third-party audits by Austria Bio Garantie verifying compliance including copper usage below 4 kg Cu/ha/year averaged over 5 years, complete absence of synthetic inputs, and documented biodiversity management through permanent cover cropping with 12-15 native plant species.
An organic production role that inheres in a grapevine that has completed the full regulatory conversion period and is cultivated under an officially recognized organic certification program, entitling the resulting grapes and wine to bear the organic label of the certifying authority.
Certified_Organic_Role represents the terminal certification status following successful completion of the conversion period (typically 3 years under EU regulation, 36 months under USDA NOP). The role is contingent upon ongoing annual inspection and recertification: loss of certification returns the vine to conventional or transitional status. This role is distinguished from Transitional_Organic_Role by the regulatory authorization to market fruit and wine under the organic designation.
Bio-zertifizierte Produktionsrolle
anerkannte Öko-Rolle
certified organic vine role
fully certified organic role
organic-certified production role
zertifizierte Bio-Rolle
EU Regulation 2018/848 Art. 10-12 on organic certification USDA NOP 7 CFR 205.400-405 IFOAM Organic Guarantee System
Certified Organic Role
Cabernet Sauvignon chamber locular tissue defining 2-chambered seed organization= left chamber containing 1-2 seeds (4.8-6.2 mm length, 3.2-4.5 mm width) bounded by chamber locular tissue 155-225 μm thickness right chamber containing 1-2 seeds with similar chamber locular tissue boundary chambers separated by incomplete septum (partition) at berry center allowing some connection between chambers total 2-4 seeds per berry (most commonly 3 seeds) distributed between 2 chambers, with chamber locular tissue providing continuous cellular boundary around seed-containing spaces defining berry internal seed-chamber organization. Merlot chamber locular tissue seed number variation effects= berries with 2 seeds (1 per chamber) showing chamber locular tissue surface area 24-32 mm² per berry berries with 4 seeds (2 per chamber) showing 36-48 mm² surface area (45-65% increase) as chambers enlarge to accommodate multiple seeds, with chamber locular tissue cell number and thickness remaining similar suggesting chamber size adjusts to seed number through expansion of existing tissue rather than additional cell layers, demonstrating chamber accommodation of varying seed numbers within genetic and developmental constraints. Syrah chamber locular tissue seed-chamber relationship during development= week 4 (early Phase I) showing small seeds 1.5-2.5 mm filling small chambers, chamber locular tissue directly contacting seed coats week 6 (late Phase I) showing seeds 4-5.5 mm nearly filling expanded chambers week 8 (lag phase) showing seeds 5.5-6.5 mm at final size with chambers 6-7.5 mm allowing 0.5-1 mm clearance week 14 (mid-ripening) showing chambers expanded to 8-10 mm with 1-2 mm clearance around seeds week 18 (maturity) showing chambers 9-12 mm with 1.5-2.5 mm clearance creating substantial air space, demonstrating progressive chamber expansion during ripening creating transformation from seed-filled to seed-containing chambers. Thompson Seedless chamber locular tissue in stenospermocarpic seedless cultivar= 2 chambers containing small seed traces 2-3.5 mm length (arrested embryo development) versus normal seeds 5-7 mm in seeded cultivars, with chamber locular tissue structure similar to seeded types (4-5 cell layers, 145-195 μm thickness) suggesting chamber differentiation programmed independently of successful seed development, chambers showing large air space (75-85% of chamber volume) surrounding small seed traces reflecting proportion difference between chamber size and seed trace size. Pinot Noir chamber locular tissue in berry shape= symmetric 2-chamber organization with chambers balanced left-right contributing to round symmetric berry shape occasional asymmetric chamber development (one chamber larger or containing more seeds) potentially contributing to asymmetric berry shapes sometimes observed, with chamber size and position influencing overall berry geometry through internal structural organization affecting how berry expands during growth creating connection between internal chamber-seed architecture and external berry morphology.
A chamber locular tissue is locular tissue emphasizing its role defining seed-containing chambers within the berry, focusing on seed-chamber organization rather than air-space aspects, with chamber locular tissue providing structural boundary for typically 2 chambers (corresponding to bicarpellate ovary origin) each containing 1-2 seeds (total 2-4 seeds per berry typical range, occasionally 0-1 or 5-6), serving as interface between mesocarp flesh and seed-chamber interiors throughout development from seed formation through maturity. Chamber locular tissue in berry carpel organization= (1) typical presence of 2 chambers reflecting bicarpellate floral origin with 2 carpels fusing during ovary development creating 2-chambered ovary structure maintained in mature berry (2) chamber position typically showing left-right symmetry when viewing berry in longitudinal section or transverse section, with chambers separated by partition tissue (septum) though partition often incomplete or absent in mature berries (3) each chamber potentially containing 1-3 ovules developing into 0-3 seeds depending on fertilization success and developmental conditions, with 1-2 seeds per chamber most common creating 2-4 seeds per berry typical range (4) chamber orientation and position influencing overall berry symmetry and shape. Chamber locular tissue seed accommodation function= (1) provision of space for seed development during Phase I (weeks 0-6) when seeds undergo rapid growth through cell division and expansion (2) chamber size potentially limiting seed size in small-berried cultivars where chamber volume constrains maximum seed dimensions (3) chamber shape influencing seed shape with seeds developing conforming to available chamber space potentially creating flattened surfaces where seeds contact chamber locular tissue or adjacent seeds within chamber (4) multiple seeds within single chamber potentially showing growth competition with seed size inversely related to seed number per chamber when resources limiting. Chamber locular tissue developmental changes in chamber-seed relationships= early development (weeks 1-6, Phase I) showing seeds growing to fill available chamber space with chamber locular tissue directly contacting developing seed coats lag phase (weeks 6-8) showing seeds completing maturation and achieving final dimensions while chambers continue some expansion beginning seed-chamber separation ripening (weeks 9-18+, Phase II) showing progressive chamber enlargement exceeding seed dimensions creating air space between seeds and chamber locular tissue walls, transforming from seed-filled chambers to seed-containing air-filled chambers during berry maturation. Chamber locular tissue in seedless cultivars showing modified organization= stenospermocarpic seedless cultivars (Thompson Seedless, Flame Seedless) showing chamber development with small seed traces (arrested embryo development creating 2-4 mm rudimentary seeds) rather than normal 5-7 mm seeds chamber locular tissue differentiation occurring normally despite arrested seed development suggesting chamber formation programmed independently of successful seed maturation chambers containing seed traces surrounded by proportionally larger air space (70-85% of chamber volume versus 50-70% in seeded types) reflecting small seed trace size parthenocarpic truly seedless cultivars showing minimal or absent chamber structure reflecting absence of fertilization stimulus for chamber development. Chamber locular tissue in wine and table grape selection= seeded wine cultivars typically showing 2-4 seeds per berry with chambers well-developed seeded table cultivars sometimes showing larger seeds (4-8 mm length versus 5-6.5 mm in wine cultivars) in well-developed chambers seedless table cultivars valued for fresh consumption showing seed traces in chambers requiring no spitting during eating breeding programs targeting seedlessness modifying chamber development and seed presence affecting eating quality and potentially affecting berry size and shape through altered chamber-seed relationships.
Chamber_Locular_Tissue emphasizes the seed-chamber organization and boundary function and is distinguished from Cavity_Locular_Tissue which emphasizes air-space aspects. Understanding chamber locular tissue is relevant for comprehending berry carpel architecture and seed organization, for seedless breeding where chamber development occurs despite arrested seed maturation, and for understanding how seed number and chamber organization influence berry size, shape, and internal structure affecting both developmental biology and practical quality characteristics.
Kammer-Lokular-Gewebe
Samenkammer-Gewebe
Samenkammer-Grenzgewebe
seed chamber lining
Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22':92-109 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 172-185.
Chamber Locular Tissue
Pinot Noir bears a Champagne_Method_Scion_Role as the most planted variety in Champagne (38% of vineyard area), providing body, vinosity, and red fruit depth to assemblages, with blanc de noirs from Aÿ, Bouzy, and Ambonnay demonstrating the variety's capacity for structured, age-worthy Champagne aging 20+ years on lees. Chardonnay serves in a Champagne_Method_Scion_Role for blanc de blancs and assemblage contribution, with Côte des Blancs Grand Crus (Le Mesnil-sur-Oger, Cramant, Avize) producing Champagnes of exceptional finesse and longevity, developing toasted brioche character over 5-10+ years of lees aging. Pinot Meunier bears a Champagne_Method_Scion_Role providing fruitiness, suppleness, and early drinkability to non-vintage blends, dominating the Marne Valley where its late budding avoids frost damage.
A sparkling wine scion role that inheres in a grafted scion cultivar by virtue of its designation for traditional method (méthode champenoise/traditionnelle) sparkling wine production, where secondary fermentation occurs in the bottle that will be sold, followed by extended lees aging, riddling (remuage), and disgorgement (dégorgement). Champagne method scions must yield base wines with characteristics that develop complexity through autolysis during bottle aging (typically 15 months minimum, often 3-10+ years for prestige cuvées), including high acidity (8-11 g/L) for freshness and longevity, moderate alcohol (10-11% base wine), fine persistent mousse supported by appropriate protein and polysaccharide content, and aromatic profiles that evolve beneficially through lees contact (developing brioche, toast, hazelnut, and aged notes). The extended contact with yeast lees (18-72+ months) fundamentally transforms wine character through mannoproteins, amino acids, and nucleotides released during autolysis. Classic Champagne varieties (Chardonnay, Pinot Noir, Pinot Meunier) define the benchmark, with regional variations (Crémant varieties, Franciacorta grapes, English sparkling cultivars) following similar principles.
Champagne_Method_Scion_Role captures cultivars suited to the demanding requirements of traditional method production, particularly extended lees aging that distinguishes this technique from faster methods. The autolytic development that defines premium traditional method sparkling requires base wines capable of extended evolution without oxidation or aromatic degradation. Beyond Champagne, the same varieties serve in traditional method production in Crémant (Loire, Alsace, Burgundy, Limoux), Franciacorta (Italy), Cava (Spain), English sparkling wine, and New World regions (California, Tasmania, Marlborough). Regulatory frameworks often specify permitted varieties for traditional method appellations, institutionalizing cultivar selection.
Champagne method variety
Champagnersorte
Flaschengärungssorte
Méthode-traditionnelle-Sorte
bottle-fermented sparkling variety
méthode traditionnelle variety
traditional method variety
Stevenson T (1998) Christie's World Encyclopedia of Champagne and Sparkling Wine, Absolute Press Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Liger-Belair G (2005) The physics and chemistry behind the bubbling properties of champagne, Journal of Agricultural and Food Chemistry 53: 2788-2802 Alexandre H, Guilloux-Benatier M (2006) Yeast autolysis in sparkling wine, Australian Journal of Grape and Wine Research 12: 119-127
Champagne Method Scion Role
Glera bears a Charmat_Method_Scion_Role as the required variety (minimum 85%) for Prosecco DOC and DOCG, with tank fermentation (typically 30-60 days) preserving its delicate floral, apple, and pear aromatics that define the Prosecco style consumed within 1-2 years of production. Muscat Blanc à Petits Grains serves in a Charmat_Method_Scion_Role for Asti Spumante and Moscato d'Asti, where tank fermentation at low pressure (2.5-3.5 atm) and controlled temperature preserves the variety's intense terpene aromatics (5-10 mg/L) and residual sweetness (90-130 g/L). Müller-Thurgau bears a Charmat_Method_Scion_Role for German Sekt production, providing floral-grapey aromatics suited to fresh consumption. Lambrusco varieties serve in Charmat_Method_Scion_Roles for red sparkling production in Emilia-Romagna.
A sparkling wine scion role that inheres in a grafted scion cultivar by virtue of its designation for tank method (Charmat/Martinotti method) sparkling wine production, where secondary fermentation occurs in large pressurized tanks rather than individual bottles, enabling efficient production of fresh, fruit-forward sparkling wines without extended lees aging. Charmat method scions are selected for aromatic varieties whose primary fruit character should be preserved rather than transformed by autolysis, moderate-to-high acidity for freshness (7-10 g/L), and flavor profiles suited to the brief lees contact (typically 1-6 months) characteristic of tank fermentation. The Charmat method preserves primary grape aromatics that would be masked by extended autolysis, making it ideal for aromatic varieties (Muscat, Glera/Prosecco) and for producing everyday sparkling wines emphasizing fruit and freshness over complexity. Tank fermentation also enables production of sweeter styles (demi-sec, doux) more readily than traditional method. Major Charmat method production includes Prosecco (Italy), Sekt (Germany), and volume sparkling wines globally.
Charmat_Method_Scion_Role captures cultivars whose aromatic expression benefits from the fruit-preserving characteristics of tank fermentation rather than the transformative autolysis of traditional method. Prosecco's global success (over 500 million bottles annually) has elevated the Charmat method from a perceived inferior technique to a respected category with its own aesthetic criteria. The method's efficiency (weeks rather than years from base wine to finished product) enables lower-priced sparkling wines, democratizing sparkling consumption. Some varieties serve in both Charmat and traditional method production (Chardonnay for both premium tank sparklings and Champagne), with method choice reflecting stylistic intent rather than cultivar limitation.
Charmat method variety
Charmat-Sorte
Prosecco method variety
Prosecco-Sorte
Tankgärungssorte
cuve close variety
tank method variety
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Jackson RS (2014) Wine Science 4th Ed, Academic Press pp.: 430-480 Consorzio Tutela del Vino Prosecco DOC (2020) Production Regulations Stevenson T (1998) Christie's World Encyclopedia of Champagne and Sparkling Wine, Absolute Press
Charmat Method Scion Role
Comprehensive metabolomic analysis of Chemical_Defense_Disposition at the Edmund Mach Foundation characterized constitutive secondary metabolite profiles in healthy leaves of 35 cultivars using LC-MS-based untargeted metabolomics. Total phenolic content ranged from 12 mg gallic acid equivalents/g fresh weight ('Müller-Thurgau') to 58 mg/g ('Tannat'), with specific compound classes showing even wider variation. Constitutive flavonol content (quercetin and kaempferol glycosides) varied 8-fold (2.1-16.8 mg/g), hydroxycinnamic acids (caftaric, coutaric acids) 4-fold (3.2-12.8 mg/g), and proanthocyanidins 12-fold (0.8-9.6 mg/g) among cultivars. Correlation with pathogen resistance phenotypes revealed compound class-specific relationships. Constitutive stilbene levels (primarily piceid) correlated with downy mildew resistance (r = -0.52 with infection severity), while proanthocyanidin content predicted Botrytis resistance (r = -0.61 with berry infection). Flavonol content showed strongest association with powdery mildew resistance (r = -0.48). In vitro bioassays confirmed that purified grapevine phenolics inhibited pathogen growth in a dose-dependent manner, with 50% inhibition concentrations ranging from 25-150 μg/mL depending on compound and pathogen. These findings demonstrate that Chemical_Defense_Disposition comprises multiple semi-independent compound classes with differential effects on specific pathogens, enabling targeted breeding for enhanced chemical defense against particular disease threats.
A Chemical_Defense_Disposition is a Constitutive_Defense_Disposition that inheres in a Vitis vinifera plant and is realized through the preformed presence of secondary metabolites and proteins with antimicrobial, antifungal, insecticidal, or feeding-deterrent properties in plant tissues, providing chemical protection against pathogens and pests without requiring induced synthesis upon attack. This disposition represents the biochemical component of constitutive defense, encompassing the diverse array of specialized metabolites and defensive proteins that accumulate in healthy plant tissues and contribute to basal resistance. Grapevine chemical defenses include phenolic compounds (flavonoids, hydroxycinnamic acids, stilbenes at basal levels, proanthocyanidins/tannins), terpenoids (monoterpenes, sesquiterpenes), antimicrobial proteins (chitinases, glucanases, thaumatin-like proteins, defensins at constitutive levels), and cell wall-associated compounds (lignin, suberin precursors). The concentration, composition, and tissue distribution of constitutive chemical defenses vary substantially among grapevine cultivars and species, contributing to differences in basal resistance even before induced defenses are activated. Chemical defenses may function through direct toxicity to attackers, inhibition of pathogen enzymes (protease inhibitors, chitinase inhibitors), membrane disruption (saponins, certain phenolics), or feeding deterrence (tannins, bitter compounds). The relationship between Chemical_Defense_Disposition and resistance depends on the specific attacker and defense compounds involved, as pathogens and pests vary in their sensitivity to particular chemical classes.
Chemical_Defense_Disposition is_a Constitutive_Defense_Disposition. This class encompasses preformed chemical defenses present before attack, complementing Physical_Barrier_Disposition and contrasting with induced chemical defenses under Induced_Defense_Disposition. Child classes include Constitutive_Phenolic_Disposition and Preformed_Antimicrobial_Disposition addressing major compound categories. Chemical defenses interact with physical barriers, as phenolics may be incorporated into cuticle and cell walls, and with induced defenses, as basal compound pools may be augmented upon attack.
Chemische-Verteidigungs-Disposition
Konstitutive-Phytochemische-Verteidigung
Präformierte-chemische-Abwehr
basal antimicrobial capacity
constitutive phytochemical defense
preformed chemical defense
Lattanzio V et al. (2006) Role of phenolics in the resistance mechanisms of plants against fungal pathogens and insects, Phytochemistry: Advances in Research, Research Signpost Wittstock U, Gershenzon J (2002) Constitutive plant toxins and their role as defenses against herbivores and pathogens, Current Opinion in Plant Biology 5:300-307 Dixon RA (2001) Natural products and plant disease resistance, Nature 411:843-847
Chemical Defense Disposition
Chemical quality profiling of Sauvignon Blanc from Marlborough, New Zealand employs comprehensive analysis of varietal markers including methoxypyrazine concentrations (IBMP targeting 10-25 ng/L for characteristic herbaceous notes), thiol precursor levels (3MH-cysteine conjugates above 200 μg/L for tropical fruit potential), titratable acidity (8-10 g/L for signature freshness), and pH (3.1-3.3 for microbial stability), enabling winemakers to allocate fruit to different wine styles based on chemical composition rather than vineyard block alone, maximizing expression of regional typicity while managing vintage variation.
Chemical_Quality encompasses all qualities that characterize the biochemical composition of grapevine organs, capturing the presence, concentration, and proportional relationships of chemical compounds that determine nutritional value, sensory properties, processing behavior, and physiological function. This category represents the molecular dimension of grape quality, complementing the morphological qualities that address physical form and structure. Chemical composition is arguably the most important quality determinant for wine grapes because the compounds present in berries at harvest directly determine wine potential, while for table grapes, chemical composition governs taste, nutritional value, and consumer acceptance. The category subdivides into seven major domains: Sugar_Content_Quality addressing carbohydrate accumulation that determines sweetness and fermentation potential Acid_Content_Quality capturing the organic acids that provide tartness and pH stability Phenolic_Content_Quality encompassing the diverse phenolic compounds responsible for color, astringency, and health benefits Aromatic_Compound_Quality addressing the volatile and non-volatile aroma precursors that define varietal character Mineral_Content_Quality capturing the inorganic elements essential for plant nutrition and wine stability Nitrogen_Content_Quality addressing the nitrogen-containing compounds critical for fermentation and wine quality and Water_Content_Quality characterizing tissue hydration status. Chemical qualities result from the integrated activity of metabolic pathways that synthesize, transport, and accumulate compounds in specific organs and tissues. Environmental factors including light, temperature, water availability, and soil composition profoundly modulate chemical quality expression within genetically determined limits. The developmental trajectory of chemical composition follows characteristic patterns, with sugars and anthocyanins accumulating during ripening while acids decline and aroma precursors undergo complex transformations. Chemical quality assessment employs diverse analytical methods ranging from simple refractometry for sugar estimation through chromatographic separation of individual compounds to advanced mass spectrometry for comprehensive metabolomic profiling.
Chemical_Quality is positioned as a direct subclass of Grapevine_Quality, parallel to Morphological_Quality, Physical_Quality, Sensory_Quality, Physiological_Quality, Phenological_Quality, Viticultural_Quality, and Stress_Response_Quality. Chemical qualities describe the molecular composition of tissues and are distinct from the sensory perception of that composition (addressed in Sensory_Quality) and from the metabolic processes that generate the composition (addressed in process ontology classes). The boundary between Chemical_Quality and Physical_Quality can be subtle chemical composition determines physical properties such as density and osmotic potential. PATO_0000070 (amount) and PATO_0000033 (concentration of) provide related upper-ontology concepts for quantitative chemical qualities.
Inhaltsstoffqualität
biochemical quality
biochemische Zusammensetzung
chemical composition character
chemische Qualität
compositional quality
molecular quality
stoffliche Qualität
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed, John Wiley and Sons pp.: 1-140 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 267-340
Chemical Quality
In Merlot vines in Bordeaux, chilling accumulation progresses through winter with approximately 200 chilling hours accumulated by late December, 600 hours by mid-January, and the cultivar-specific requirement of approximately 900-1000 chilling hours satisfied by early February, as demonstrated by forcing tests showing budbreak time decreasing from more than 45 days in December to less than 20 days in February under identical 22°C forcing conditions, indicating progressive dormancy release through chilling accumulation.
Chilling accumulation in Vitis vinifera represents the progressive process during which exposure to low temperatures gradually satisfies the genetically determined chilling requirement, progressively reducing dormancy depth and restoring growth competence to endodormant buds. This process occurs throughout the winter period, typically from December through February in Northern Hemisphere viticulture, as cold temperatures induce the physiological changes that dismantle dormancy-maintaining mechanisms and prepare buds for eventual growth resumption. The quantification of chilling accumulation relies on models that integrate temperature exposure over time, with the simplest models counting hours below a threshold temperature, typically 7.2°C or 45°F based on research in deciduous fruit trees, while more sophisticated models apply different weights to different temperature ranges recognizing that moderately cold temperatures are most effective for dormancy release. The physiological changes occurring during chilling accumulation include progressive reduction in ABA concentration through increased catabolic enzyme activity, declining expression of dormancy-associated MADS-box genes, gradual removal of repressive chromatin marks from growth-related gene loci, and restoration of symplastic communication through callose degradation at plasmodesmata. The rate of chilling accumulation depends on temperature regime, with temperatures in the range of 0-7°C generally most effective for dormancy release, temperatures below -2°C being less effective or ineffective, and temperatures above 15°C potentially negating previously accumulated chilling in some models. The progress of chilling accumulation is not externally visible but can be tracked through periodic forcing tests that reveal progressively shorter times to budbreak as dormancy depth decreases with increasing chilling exposure. The completion of chilling accumulation, when the chilling requirement has been fully satisfied, marks the transition from endodormancy to ecodormancy, at which point buds have regained growth competence but remain quiescent due to unfavorable environmental temperatures rather than internal inhibition. Cultivar differences in chilling requirement significantly affect the duration of chilling accumulation, ranging from approximately 400 chilling hours for low-chill cultivars to more than 1500 hours for high-chill cultivars.
Chilling_Accumulation represents a major phase within Endodormancy addressing the progressive dormancy release through cold exposure. Distinguished from Deep_Endodormancy which precedes significant chilling accumulation, and from Dormancy_Release which marks completion of the process. Subdivides into Progressive_Chilling_Units, Dormancy_Depth_Reduction, and Cumulative_Cold_Exposure as component aspects. Duration spans most of winter from December through February typically. Effective temperature range approximately 0-7°C. Cultivar chilling requirements range from 400-1500+ hours.
Chilling-Akkumulation
Kälteeinheiten-Ansammlung
Kältestunden-Akkumulation
Kältesumme
chill accumulation
chilling hour accumulation
cold unit accumulation
winter chilling
Dokoozlian NK (1999) Chilling temperature and duration interact on the budbreak of Perlette grapevine cuttings. HortScience 34: 1054-1056 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Richardson EA, Seeley SD, Walker DR (1974) A model for estimating the completion of rest for Redhaven and Elberta peach trees. HortScience 9: 331-332
Chilling Accumulation
Chilling accumulation tracking through an anomalously warm winter in Merlot reveals the climate risk from insufficient cold exposure, with December accumulating only 180 chill hours (versus 320 typical), January adding 220 hours (versus 380 typical), and February adding 150 hours (versus 280 typical), yielding total accumulation of 550 chill hours against a requirement of approximately 600 hours, with the deficit resulting in delayed budbreak (12 days later than normal despite warm March), reduced budbreak percentage (78% versus typical 95%), and erratic timing (budbreak spread over 18 days versus typical 8 days), demonstrating how insufficient chilling accumulation relative to requirement produces the characteristic symptoms of incomplete endodormancy release.
Chilling accumulation quality is an endodormancy depth quality that represents the cumulative cold exposure actually received by grapevine buds during the dormant period, characterizing the progress toward satisfaction of the chilling requirement that determines how much of the endodormancy release threshold has been met and how much additional chilling is needed before buds achieve full growth competence. Within the BFO framework, this quality specifically addresses the accumulation dimension of chilling, distinguishing the cold actually experienced from the requirement threshold while recognizing that chilling accumulation relative to requirement determines endodormancy status, with complete accumulation (meeting or exceeding requirement) enabling transition to ecodormancy and incomplete accumulation leaving buds in partially released endodormancy with consequences for budbreak timing and uniformity. The ontological significance lies in chilling accumulation being the dynamic variable that tracks progress through winter toward dormancy release, where monitoring accumulation enables prediction of when endodormancy will be satisfied and subsequent heat accumulation will become relevant for budbreak timing, with this information critical for vineyard planning and for understanding the climate vulnerability of production systems dependent on adequate winter chilling. The temperature effectiveness for chilling accumulation varies with temperature according to the model used, with the chill hour model counting all hours below 7°C equally, the Utah model weighting temperatures optimally effective around 6°C with reduced effectiveness at lower temperatures and negative (chilling-negating) effects above 16°C, and the Dynamic model incorporating the two-step process where cold creates an intermediate that must be consolidated by moderate temperatures to create permanent chill portions, with model selection affecting accumulation calculations and predictions. The spatial and temporal variation in chilling accumulation reflects the microclimate differences within and among vineyards, with elevation, aspect, cold air drainage, and urban heat island effects creating variation in accumulated chilling that may cause differential dormancy release timing across short distances. Understanding chilling accumulation quality enables tracking of dormancy release progress, prediction of budbreak timing based on accumulation relative to requirement, and identification of sites or years where insufficient chilling may compromise dormancy release and subsequent vine performance.
Chilling_Accumulation_Quality addresses the cumulative cold exposure tracking progress toward endodormancy release. This quality varies with winter weather and determines dormancy status relative to requirements. The class supports dormancy monitoring and budbreak prediction through chilling accumulation characterization.
Chilling-Erfüllung
Kälteakkumulation
Kälteeinheitenakkumulation
Vernalisationsfortschritt
chill unit accumulation
chilling satisfaction
cold accumulation
vernalization progress
Luedeling E et al. (2009) Sensitivity of winter chill models for fruit and nut trees to climatic changes expected in California's Central Valley. Agriculture, Ecosystems & Environment 133(1-2): 23-31 Ferguson JC et al. (2014) Dynamic thermal time model of cold hardiness for dormant grapevine buds. Annals of Botany 113(5): 801-811
Chilling Accumulation Quality
In Riesling vines in the Rheingau, Germany, chilling completion occurs in late January to early February when accumulated chilling hours reach the cultivar requirement of approximately 1200-1400 hours, verified by forcing tests showing budbreak within 10 days at 22°C and greater than 95% bud break success rate, marking the definitive end of endodormancy and enabling subsequent response to spring warming for budbreak initiation.
Chilling completion in Vitis vinifera represents the achievement of full chilling requirement satisfaction marking the definitive end of endodormancy and the restoration of complete growth competence, occurring when cumulative cold exposure has fully dismantled all dormancy-maintaining mechanisms and enabled unrestricted growth response to favorable conditions. This milestone typically occurs during late January to February in Northern Hemisphere viticulture, representing the point at which the genetically determined chilling requirement has been met or exceeded through accumulated winter cold exposure. The physiological significance of chilling completion lies in the full restoration of growth capacity, with all internal barriers to growth having been removed through the chilling-dependent reduction of ABA, silencing of DAM genes, derepression of growth gene chromatin, and restoration of cellular communication pathways that were restricted during endodormancy. The verification of chilling completion relies on forcing tests demonstrating rapid and complete budbreak response, with completed chilling indicated by budbreak times of less than 10-14 days under optimal forcing conditions and high percentages of buds successfully breaking, contrasting with the extended times and incomplete response characteristic of insufficient chilling. The quantification of chilling completion uses accumulated chilling hours or units according to various models, with completion defined as achieving or exceeding the cultivar-specific requirement that typically ranges from 400-600 hours for low-chill cultivars to 1200-1500 hours for high-chill cultivars when using the simple hours-below-7.2°C model. The margin by which chilling exceeds the minimum requirement affects subsequent growth characteristics, with substantial excess chilling potentially advancing budbreak timing while barely adequate chilling may result in acceptable but suboptimal budbreak uniformity. The timing of chilling completion relative to subsequent warming patterns determines the duration of ecodormancy and influences budbreak timing, with early chilling completion in warm winters creating extended ecodormancy periods while late completion in cold winters may be followed rapidly by warming that triggers immediate budbreak. The consequences of incomplete chilling, where the requirement is not fully satisfied, include the delayed, prolonged, and uneven budbreak characteristic of inadequate dormancy release that can significantly impact vineyard productivity.
Chilling_Completion represents the first component phase of Dormancy_Release marking definitive endodormancy termination. Distinguished from Chilling_Accumulation which addresses the progressive process, while Chilling_Completion specifically marks full requirement satisfaction. Subdivides into Chilling_Requirement_Met, Full_Chill_Accumulation, and Competence_Achievement as component aspects. Verified by forcing test response less than 14 days with high success rate. Cultivar requirements range from approximately 400-1500 hours. Timing typically late January to February in Northern Hemisphere.
Chilling-Abschluss
Chilling-Befriedigung
Kältebedürfnis-Vollendung
chill requirement completion
chilling fulfillment
full chilling achievement
vollständige Kälteerfüllung
Dokoozlian NK (1999) Chilling temperature and duration interact on the budbreak of Perlette grapevine cuttings. HortScience 34: 1054-1056 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Lavee S, May P (1997) Dormancy of grapevine buds - facts and speculation. Australian Journal of Grape and Wine Research 3: 31-46
Chilling Completion
Chilling requirements in V. vinifera range from 150-400 chilling hours (hours below 7.2°C) for low-chill cultivars to 800-1400 hours for high-chill cultivars adapted to continental climates. Pinot Noir requires approximately 700-1000 chilling hours for complete dormancy release, while Thompson Seedless satisfies chilling requirement with only 200-400 hours. The Utah Model quantifies chilling as weighted units based on temperature effectiveness (0-2.4°C optimal, negative contributions above 16°C). Insufficient chilling causes delayed, erratic, or incomplete budbreak with budbreak percentages dropping from 95% to 40-60% when chilling is 50% below cultivar requirement.
A dormancy disposition that is the inherited capacity of a Vitis vinifera plant to require accumulation of a genotype-specific quantity of cold exposure (chilling units) during endodormancy before buds become competent to resume growth, determining the minimum winter cold necessary for uniform and complete budbreak.
Chilling_Requirement_Disposition serves as the parent class for high and low chilling categories. Multiple quantification models exist (chilling hours, Utah model, Dynamic model), with the Dynamic model showing best transferability across climates. Chilling requirement has a genetic basis but shows some environmental modification through prior-year temperature experience. Climate warming is reducing chilling accumulation in traditional wine regions by 10-30 hours per decade, threatening cultivar suitability in marginal areas.
Chilling-Anforderung
Kältebedarf
Kältesummen-Disposition
chilling hours requirement
cold accumulation need
vernalization requirement
Dokoozlian NK (1999) Chilling temperature and duration interact on the budbreak of Perlette grapevine cuttings. HortScience 34:1054-1056 Luedeling E et al. (2009) Sensitivity of grapevine phenology models to temperature: implications for climate change impacts across growing regions. Agricultural and Forest Meteorology 149:1115-1126
Chilling Requirement Disposition
In Merlot vines in Bordeaux, chilling requirement met is achieved in early February when accumulated chilling hours of approximately 950 reach the cultivar threshold, demonstrated by forcing test response decreasing from 25 days in mid-January to 12 days by February 10, indicating that the chilling requirement has been satisfied and buds have transitioned from residual endodormancy to fully released ecodormancy status.
Chilling requirement met in Vitis vinifera represents the specific point at which accumulated cold exposure equals the genetically determined threshold required for complete dormancy release, marking the transition from a state of partially satisfied chilling where dormancy constraints remain to a state of fully satisfied chilling where all internal growth barriers have been removed. This threshold achievement typically occurs during late January to February in Northern Hemisphere viticulture when cumulative chilling hours or units reach the cultivar-specific requirement through the integration of winter cold exposure over time. The physiological meaning of requirement satisfaction involves the completion of all chilling-dependent processes that progressively dismantle dormancy-maintaining mechanisms, with the requirement threshold representing the cumulative exposure needed to reduce ABA below inhibitory levels, silence DAM gene expression, remove chromatin repression from growth genes, and restore cellular communication pathways to degrees compatible with unrestricted growth response. The verification that the requirement has been met relies on forcing tests demonstrating the rapid budbreak response characteristic of fully released dormancy, with requirement satisfaction indicated by forcing times decreasing below approximately 14 days and approaching the minimum times characteristic of fully ecodormant material with no residual endodormancy constraints. The quantification of requirement satisfaction depends on the chilling model employed, with requirements typically expressed as hours below 7.2°C ranging from approximately 400-600 hours for low-chill cultivars adapted to mild winter climates to 1200-1500 hours for high-chill cultivars from continental regions, though more sophisticated models using temperature weighting or dynamic calculations may express requirements in different units. The practical determination of when requirements are met involves tracking chilling accumulation through winter and comparing accumulated units to known cultivar requirements, enabling prediction of when dormancy release will be complete and subsequent heat accumulation can effectively drive progress toward budbreak. The consequences of requirements not being met include the persistence of residual endodormancy that prolongs forcing response times, reduces budbreak uniformity, and may cause the delayed and erratic budbreak characteristic of inadequate chilling in warm winter climates.
Chilling_Requirement_Met represents the threshold achievement aspect of Chilling_Completion marking exact requirement satisfaction. Distinguished from progressive Chilling_Accumulation by representing the specific threshold crossing point. Defined by accumulated chilling equaling cultivar-specific requirement. Verified through forcing test response approaching minimum times. Requirements vary from approximately 400-1500 hours depending on cultivar origin. Critical transition point from partial to complete dormancy release.
Chilling-Schwelle-erreicht
Dormanz-Kältequote-erreicht
Kälteanforderung-befriedigt
Kältebedarf-Erfüllung
chill requirement satisfied
chilling quota achieved
chilling threshold reached
dormancy chilling fulfilled
Chilling Requirement Met
Chilling requirement comparison among cultivars reveals the genetic variation affecting climate adaptation, with low-chill cultivar (Sultana, requiring approximately 150 chill hours or 20 chill portions) suitable for warm Mediterranean climates but prone to erratic budbreak in continental climates from premature chilling satisfaction followed by warm spells, moderate-chill cultivar (Chardonnay, requiring approximately 400 chill hours or 35 chill portions) broadly adapted to temperate regions with typical winter cold, and high-chill cultivar (Riesling, requiring approximately 800 chill hours or 55 chill portions) requiring substantial winter cold and potentially experiencing delayed or erratic budbreak in mild winter climates where requirements are incompletely satisfied.
Chilling requirement quality is an endodormancy depth quality that represents the cumulative exposure to chilling temperatures needed to fully release endodormancy and restore complete growth competence to grapevine buds, characterizing the genetically determined threshold of cold exposure that must be satisfied before buds transition from endodormancy to ecodormancy and become fully responsive to subsequent warming. Within the BFO framework, this quality specifically addresses the threshold dimension of endodormancy release, distinguishing the required chilling amount from the actual accumulation while recognizing that chilling requirement represents a cultivar-specific characteristic that determines the minimum cold exposure needed for normal dormancy release, with consequences for cultivar adaptation to different climates and for budbreak behavior when requirements are incompletely satisfied. The ontological significance lies in chilling requirement determining the climate adaptation range of cultivars, where low-chilling cultivars (requiring less cold exposure) can succeed in mild winter climates but may break dormancy prematurely in cold climates, while high-chilling cultivars (requiring extensive cold exposure) are protected against premature budbreak in cold climates but may fail to release dormancy properly in mild winters, making chilling requirement a key factor in cultivar-climate matching. The genetic basis for chilling requirement variation shows substantial differences among Vitis vinifera cultivars (typically requiring 150-1500 chill units depending on cultivar and calculation method), with generally higher requirements in cultivars originating from continental climates and lower requirements in cultivars from Mediterranean origins, though substantial variation exists within any geographic grouping reflecting the complex genetic control of dormancy depth. The calculation methods for chilling requirement include simple chill hour accumulation (hours below 7°C), the Utah model (weighting different temperature ranges for chilling effectiveness, with some temperatures counted negatively), and the Dynamic model (incorporating the concept of chill portions that accumulate through a two-step process), with different models showing different predictive accuracy in different climates. Understanding chilling requirement quality enables selection of cultivars appropriate for local winter climate, prediction of dormancy release timing based on accumulated versus required chilling, and assessment of climate change vulnerability for high-chilling cultivars in warming regions.
Chilling_Requirement_Quality addresses the genetically determined threshold of cold exposure for complete endodormancy release. This quality varies among cultivars and determines climate adaptation range. The class supports cultivar-climate matching through chilling requirement characterization.
Chilling-Schwelle
Kälteeinheitenbedarf
Vernalisationsbedarf
chill unit requirement
chilling threshold
cold requirement
vernalization need
Dokoozlian NK (1999) Chilling temperature and duration interact on the budbreak of 'Perlette' grapevine cuttings. HortScience 34(6): 1054-1056 Ferguson JC et al. (2011) Modeling dormant bud cold hardiness and budbreak in twenty-three Vitis genotypes reveals variation by region of origin. American Journal of Enology and Viticulture 62(4): 516-527
Chilling Requirement Quality
In Merlot vines in Saint-Émilion, Bordeaux, chilling requirement satisfaction is achieved in early February when accumulated chilling hours reach the cultivar threshold of approximately 900-950 hours, confirmed by forcing tests showing 50% budbreak within 12 days at 22°C, ABA concentrations having declined to 60-70 ng per gram fresh weight, and subsequent field budbreak occurring uniformly within 10 days of sustained temperatures above 10°C in late March.
Chilling requirement satisfaction in Vitis vinifera represents the achievement of sufficient cumulative cold exposure to fully release endodormancy, restoring growth competence to previously dormant buds and enabling the transition from internally-controlled endodormancy to environmentally-controlled ecodormancy. This milestone typically occurs during late January to February in Northern Hemisphere viticulture, when accumulated chilling hours or units reach the cultivar-specific threshold that has been genetically determined through adaptation to the climate of origin. The physiological basis of chilling requirement satisfaction involves the cumulative dismantling of dormancy-maintaining mechanisms through chilling-dependent processes, with sufficient chilling reducing ABA levels below inhibitory thresholds, decreasing DAM gene expression to levels compatible with growth, removing repressive chromatin modifications from growth gene promoters, and restoring cellular communication pathways required for growth coordination. The concept of satisfaction implies a threshold phenomenon where a specific amount of chilling must be accumulated before dormancy release occurs, though in practice the transition from endodormancy to ecodormancy appears gradual with dormancy depth progressively decreasing until it crosses the threshold where maintaining mechanisms can no longer prevent growth response to favorable conditions. The quantification of chilling requirement satisfaction depends on the chilling model employed, with requirements typically expressed as hours below 7.2°C ranging from approximately 400-600 hours for low-chill cultivars adapted to Mediterranean climates to 1200-1500 hours for high-chill cultivars from continental regions. The practical importance of chilling requirement satisfaction relates to the timing and uniformity of budbreak in spring, with fully satisfied chilling requirements producing rapid, synchronous budbreak when temperatures permit, while inadequately satisfied requirements result in delayed, prolonged, and uneven budbreak that complicates vineyard management and may reduce productivity. Climate change poses significant challenges to chilling requirement satisfaction in many viticultural regions, as warming winters may fail to provide sufficient chilling for complete dormancy release, particularly for high-chill cultivars in regions near the warm margin of their adaptive range. The verification of chilling requirement satisfaction through forcing tests shows rapid budbreak within 10-14 days under optimal conditions, contrasting with the extended times characteristic of insufficiently chilled buds still in endodormancy.
Chilling_Requirement_Satisfaction represents the culminating phase of Endodormancy marking achievement of full dormancy release competence. Distinguished from Chilling_Accumulation which addresses the progressive process, while Chilling_Requirement_Satisfaction specifically marks threshold achievement. Subdivides into Adequate_Chilling_Achievement, Dormancy_Release_Competence, and Endodormancy_Completion as component aspects. Cultivar requirements range from approximately 400-1500 hours. Marks transition from endodormancy to ecodormancy. Critical for uniform spring budbreak.
Chilling-Requirement-Befriedigung
Kälteanforderungs-Erfüllung
Kältebedarf-Deckung
Kältebedürfnis-Erfüllung
chill satisfaction
chilling requirement completion
dormancy chilling achievement
Chilling Requirement Satisfaction
The chimera status of Pinot Gris documents a periclinal chimera in which the L1 (epidermal) layer carries a non-functional VvMYBA1 allele (homozygous for Gret1 insertion, producing no anthocyanin in the epidermis — similar to white-berried Pinot Blanc), while the L2 (subepidermal) layer retains the functional VvMYBA1 allele of the parental Pinot Noir genotype, producing anthocyanin in the subepidermal cell layer that shows through the unpigmented epidermis to create the characteristic greyish-pink berry color — this L1/L2 genotypic discordance was confirmed by laser microdissection of individual cell layers followed by SSR and VvMYBA1 genotyping (Hocquigny et al. 2004), demonstrating that Pinot Gris is not a simple intermediate between Pinot Noir and Pinot Blanc but a stable periclinal chimera whose distinctive phenotype depends on the maintenance of genetically distinct meristem layers through vegetative propagation.
A ploidy information record documenting the chimeric constitution of a specific Vitis vinifera accession in which genetically distinct cell populations coexist within the same plant body, recording the ploidy or genotypic composition of the meristematic cell layers (L1 epidermis, L2 subepidermis, L3 internal corpus) that differ from each other due to somatic mutation events followed by cell sorting within the structured shoot apical meristem, producing a stable periclinal chimera maintained through vegetative propagation.
Chimera_Status documents a phenomenon of profound importance for understanding grapevine clonal diversity — the coexistence of genetically different cell populations within a single plant, stabilized by the layered structure of the angiosperm shoot apical meristem. The grapevine shoot apex comprises three clonally independent cell layers (L1, L2, L3) that maintain their identity through predominantly anticlinal cell division, meaning that somatic mutations arising in one layer are propagated within that layer but do not normally cross into adjacent layers: this layered architecture enables the stable persistence of genotypic differences between layers — ploidy chimeras (different ploidy levels in different layers), genotypic chimeras (different alleles at specific loci in different layers), and sectorial chimeras (mutations affecting only part of a single layer). The berry color variants of the Pinot family provide the best-documented example — Pinot Noir (functional VvMYBA1 in all layers), Pinot Blanc (non-functional VvMYBA1 in all layers), and Pinot Gris (non-functional in L1, functional in L2) represent different chimeral states of the same base genotype, with the greyish-pink Pinot Gris phenotype being specifically chimera-dependent. Chimeral instability through occasional cell layer displacement can revert Pinot Gris to either Pinot Noir or Pinot Blanc, a phenomenon observed in vineyard practice as individual mutant shoots (reversion sports) on otherwise stable chimeral plants.
Chimären-Status
Meristemschicht-Genotyp-Nachweis
cytochimera status
meristem layer genotype documentation
periclinal chimera record
periklinal-Chimäre-Dokumentation
Hocquigny S et al. (2004) Diversification within grapevine cultivars goes through chimeric states, Genome 47: 579-589 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340 Walker AR et al. (2007) White grapes arose through the mutation of two similar and adjacent regulatory genes, Plant Journal 49: 772-785 Franks T et al. (2002) Regeneration of transgenic Vitis vinifera L. Sultana plants, Molecular Breeding 10: 135-145
Chimera Status
A chip bud of Pinot Noir clone 115 inserted into one-year-old field-rooted SO4 rootstock bears a Chip_Bud_Role in French nursery production, performed in August with the chip precisely matched to a corresponding excision in the rootstock, wrapped with degradable budding tape, and left to callus in place—the bud pushing growth the following spring. A chip bud of Shiraz clone BVRC30 serves in a Chip_Bud_Role for Australian field budding, where the technique's extended season window (March-May) provides flexibility compared to the brief T-budding window. A dormant chip bud of Riesling inserted in late winter bears a Chip_Bud_Role in German nursery practice, exploiting the technique's viability outside the active growth period when T-budding cannot be performed.
A field graft role that inheres in a grafted combination assembled using the chip budding technique, where a rectangular or shield-shaped chip of scion tissue containing a single bud is fitted into a matching excision on the rootstock, creating cambial contact along the chip margins. Chip budding involves removing a chip of bark and underlying tissue (typically 2-3 cm long, 1-1.5 cm wide) containing the scion bud, then cutting an identical-shaped notch in the rootstock into which the chip is fitted, secured with budding tape or elastic bands. The technique offers advantages over T-budding including an extended working season (chip budding can be performed from late dormancy through active growth, while T-budding requires bark slip), suitability for smaller-diameter rootstocks, and potentially higher success rates with difficult cultivars. Chip bud success depends on precise matching of chip and notch dimensions (ensuring cambial contact around the chip perimeter), secure wrapping to maintain contact during callus formation, and appropriate environmental conditions supporting callus development without excessive desiccation or pathogen infection.
Chip_Bud_Role captures a budding technique increasingly favored over traditional T-budding due to its extended season flexibility and suitability for smaller rootstock diameters. The chip format provides cambial contact around the entire chip perimeter (versus only the top and sides in T-budding), potentially improving callus formation and union strength. Chip budding can be mechanized to some degree—chip-cutting machines produce consistent chips from budwood—though insertion remains largely manual. The technique's viability during dormancy or early season (before bark slip permits T-budding) extends the nursery production calendar, enabling earlier field work or late-season grafting after T-budding windows close. Success rates of 80-90% are achievable with skilled operators and quality materials, comparable to T-budding under optimal conditions.
Chip-Okulation
Plattenokulation
Spanokulation
chip bud
chip budding graft
chip graft
shield chip bud
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 445-450 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Dry N (2007) Grapevine rootstocks, Lythrum Press Whiting JR (2004) Grapevine rootstocks, Winetitles pp.: 167-188
Chip Bud Role
Molecular characterization of Chitinase_Induction_Disposition at the University of Florence compared chitinase gene expression and enzyme activity across 15 cultivars following Botrytis cinerea berry inoculation. Expression of four chitinase genes (VvChit1b class I, VvChit4a class III, VvChit4c class IV, VvChi1 class I) was monitored by qRT-PCR at 0, 12, 24, 48, and 72 hpi, with parallel measurement of total chitinase enzyme activity using colloidal chitin substrate. Resistant 'Nebbiolo' exhibited strong Chitinase_Induction_Disposition with VvChit4c (class IV) showing 45-fold induction by 24 hpi, VvChit1b (class I) reaching 28-fold at 48 hpi, and total chitinase activity increasing 8-fold by 48 hpi. Susceptible 'Moscato Bianco' showed delayed, weak induction with VvChit4c only 6-fold at 48 hpi and enzyme activity increasing just 2-fold. Class IV chitinase (VvChit4c) showed the strongest correlation with resistance (r = -0.82 between expression and infection severity), suggesting this class is most important for B. cinerea defense. In vitro assays demonstrated that purified grapevine class IV chitinase inhibited B. cinerea hyphal growth by 70% at 25 μg/mL and reduced conidial germination by 55%, confirming direct antifungal function. Transgenic tobacco overexpressing VvChit4c showed enhanced B. cinerea resistance, providing functional validation of this gene's defensive role.
A Chitinase_Induction_Disposition is a Pathogenesis_Related_Protein_Disposition that inheres in a Vitis vinifera plant and is realized through rapid and strong transcriptional activation and accumulation of chitinase enzymes (PR-3 family) in response to pathogen attack, providing enhanced capacity to degrade chitin, a major structural component of fungal cell walls and insect exoskeletons. This disposition represents the induction of one of the most important PR protein families for antifungal defense, as chitinase activity directly attacks fungal hyphal integrity and can inhibit spore germination. Plant chitinases are classified into classes I-V based on structure, with grapevine expressing representatives of classes I, II, III, and IV, each with distinct biochemical properties and expression patterns. Chitinases function by hydrolyzing the β-1,4-glycosidic bonds between N-acetylglucosamine residues in chitin, releasing oligomers that themselves can act as elicitors triggering further defense responses in a positive feedback loop. The speed and magnitude of chitinase induction vary among genotypes, with resistant cultivars typically showing earlier induction onset (detectable within 6-12 hours post-inoculation), higher peak expression (20-50 fold above basal), and more sustained elevation compared to susceptible cultivars. Chitinase_Induction_Disposition is particularly relevant for resistance to fungal pathogens with chitin-containing cell walls, including Erysiphe necator (powdery mildew), Botrytis cinerea (grey mold), and various trunk disease fungi, but not for oomycetes like Plasmopara viticola whose cell walls lack chitin.
Chitinase_Induction_Disposition is_a Pathogenesis_Related_Protein_Disposition. Chitinases target chitin in fungal cell walls and insect exoskeletons, making them specifically effective against true fungi (Ascomycetes, Basidiomycetes) but not oomycetes (e.g., Plasmopara viticola) whose cell walls contain cellulose rather than chitin. Multiple chitinase classes exist with distinct properties and expression patterns. Chitinase activity releases chitin oligomers that can trigger additional defense responses, creating positive feedback amplification of defense signaling.
Antifungale-Chitinase-Antwort
Chitinase-Induktions-Disposition
PR-3 induction
PR-3-Induktion
antifungal chitinase response
chitinase gene activation
Collinge DB et al. (1993) Plant chitinases, Plant Journal 3:31-40 Robert N et al. (2002) Molecular characterization of the incompatible interaction of Vitis vinifera leaves with Pseudomonas syringae pv. pisi: expression of genes coding for stilbene synthase and class 10 PR protein, European Journal of Plant Pathology 108:837-847 Derckel JP et al. (1998) Differential expression of grapevine (Vitis vinifera L.) chitinase genes following challenge with fungal and bacterial pathogens, Vitis 37:165-170
Chitinase Induction Disposition
Ramsey (Salt Creek, V. champinii) is the benchmark chloride excluder, maintaining Sultana (Thompson Seedless) leaf Cl⁻ below 0.25% dry weight when irrigated at ECw 2.0 dS/m in the Murray-Darling Basin, Australia, while susceptible Schwarzmann allows leaf Cl⁻ accumulation to 1.5-2.0% causing severe necrosis under identical conditions. 1103 Paulsen restricts Cl⁻ translocation to Sangiovese foliage (leaf Cl⁻ 0.3-0.4% at ECw 1.5 dS/m) in saline-irrigated Tuscan coastal vineyards where SO4 permits leaf Cl⁻ exceeding 1.0%. Dog Ridge (V. champinii) demonstrates strong chloride exclusion (leaf Cl⁻/root zone Cl⁻ ratio of 0.015) supporting table grape production on saline soils of Israel's Negev Desert.
A salt-tolerant rootstock role that inheres in a grafted rootstock by virtue of its specific capacity to restrict chloride ion (Cl⁻) uptake by roots and translocation to the scion, thereby protecting scion foliage from chloride toxicity under saline conditions. Chloride exclusion is the primary determinant of grapevine salt tolerance, as Cl⁻ is more readily accumulated than Na⁺ in Vitis species and causes toxicity symptoms (marginal and interveinal leaf necrosis, premature leaf abscission) at lower tissue concentrations (toxicity threshold 0.5-1.0% Cl⁻ dry weight versus 0.25-0.5% Na⁺). Chloride-excluding rootstocks restrict Cl⁻ movement through mechanisms including reduced passive Cl⁻ permeability of root cell membranes, active Cl⁻ efflux from root cells back to the soil solution, compartmentalization of Cl⁻ in root cortex vacuoles, and restricted Cl⁻ loading into the xylem at the root stele. Exclusion efficiency is quantified by the ratio of leaf Cl⁻ concentration to external solution Cl⁻ concentration, with strong excluders maintaining ratios below 0.02 versus 0.05-0.10 for poor excluders.
Chloride_Excluding_Rootstock_Role captures the ion-specific exclusion mechanism that is quantitatively most important for grapevine salinity tolerance. V. champinii (Ramsey, Dog Ridge) and V. berlandieri × V. rupestris crosses (1103P, 140Ru, 110R) are generally strong Cl⁻ excluders, while V. riparia and V. berlandieri × V. riparia crosses (SO4, 5BB, 5C) are typically poor excluders. The genetic basis of Cl⁻ exclusion involves multiple QTLs controlling root membrane transport, with breeding programs at CSIRO and UC Davis actively selecting for enhanced exclusion. Chloride exclusion is largely independent of Na⁺ exclusion, and rootstocks may be strong excluders for one ion but not the other.
Chlorid-ausschließende Unterlage
Cl⁻ excluding rootstock
Unterlage mit geringer Chloridaufnahme
chloride excluding rootstock
chloride exclusion rootstock
chloridexkludierende Unterlage
low chloride uptake rootstock
Walker RR et al. (2002) Rootstock effects on salt tolerance of irrigated field-grown grapevines, Australian Journal of Grape and Wine Research 8: 146-155 Tregeagle JM et al. (2010) Chloride exclusion by grafted grapevines, Journal of Experimental Botany 61: 1043-1051 Gong H et al. (2011) Mechanisms of chloride exclusion in salt tolerant grapevines, Functional Plant Biology 38: 708-719 Fort KP et al. (2015) Chloride exclusion by roots of grapevine rootstocks, Scientia Horticulturae 192: 1-8
Chloride Excluding Rootstock Role
Mechanistic investigation of Chloride_Exclusion_Disposition at the University of Adelaide combined ion flux measurements, gene expression analysis, and grafting experiments to characterize chloride exclusion across 12 rootstocks. ²²Cl⁻ tracer studies quantified unidirectional root chloride uptake and xylem loading rates, while qRT-PCR assessed expression of candidate chloride transporter genes (VvCLC-c, VvSLAH, VvNPF). Chloride_Exclusion_Disposition showed strong genotypic variation with distinct mechanistic bases. '140 Ruggeri' exhibited strong exclusion through reduced xylem loading (only 8% of absorbed chloride translocated to shoot) combined with moderate uptake restriction. Gene expression analysis revealed that '140 Ruggeri' roots showed 4-fold lower expression of VvSLAH3 (a xylem-loading anion channel) under salinity compared to salt-sensitive 'Schwarzmann'. Reciprocal grafting experiments confirmed that Chloride_Exclusion_Disposition was rootstock-determined, with 'Shiraz' scions accumulating 22 mmol Cl⁻/kg DW on '140 Ruggeri' roots versus 385 mmol/kg on 'Schwarzmann' roots under identical salinity. Root chloride concentration analysis revealed that excluding rootstocks accumulated chloride in root cortex vacuoles (45 mM in root versus 18 mmol/kg in leaf), suggesting that root sequestration contributes to Chloride_Exclusion_Disposition alongside xylem loading restriction. These findings demonstrate that Chloride_Exclusion_Disposition involves coordinated reduction of xylem loading and enhanced root compartmentalization, with transporter gene expression differences underlying genotypic variation.
A Chloride_Exclusion_Disposition is an Ion_Exclusion_Disposition that inheres in a Vitis plant and is realized through physiological mechanisms that specifically prevent or limit the uptake of chloride ions from saline soil and their translocation to shoot tissues, maintaining leaf chloride concentrations below toxic thresholds. This disposition represents the most critical component of grapevine salt tolerance, as grapevine is considerably more sensitive to chloride toxicity than to sodium toxicity, distinguishing it from many other crop species. Chloride toxicity in grapevine manifests as leaf marginal and tip necrosis, premature leaf abscission, reduced photosynthesis, and ultimately vine decline, with symptoms typically appearing when leaf chloride exceeds 25-50 mmol/kg DW depending on cultivar and environmental conditions. Chloride exclusion involves both restriction of chloride uptake at root level and control of chloride loading into the xylem, with the latter appearing particularly important for grapevine rootstock differences. The molecular basis involves regulation of chloride channels (CLC family) and transporters in root cells, with effective excluders showing reduced expression or activity of xylem-loading transporters and potentially enhanced vacuolar sequestration in root tissues. Chloride_Exclusion_Disposition varies dramatically among rootstocks, with excluders maintaining leaf chloride below 30 mmol/kg DW at external concentrations that cause sensitive rootstocks to accumulate over 400 mmol/kg DW.
Chloride_Exclusion_Disposition is_a Ion_Exclusion_Disposition. Chloride exclusion is the dominant determinant of grapevine salt tolerance, as grapevine is more chloride-sensitive than sodium-sensitive. Child classes include Root_Chloride_Exclusion_Disposition (uptake restriction) and Xylem_Loading_Restriction_Disposition (translocation control). Xylem loading restriction appears more important than uptake restriction for rootstock differences. Key genes include anion channels (SLAH, CLC families) and transporters. Excluder rootstocks may compartmentalize chloride in root vacuoles as an additional tolerance mechanism.
Chlorid-Restriktion
Chloridausschluss-Disposition
Cl⁻ exclusion
Cl⁻-Ausschluss
chloride avoidance
chloride restriction
Tregeagle JM et al. (2010) Chloride and sodium accumulation in grapevine rootstocks, Australian Journal of Grape and Wine Research 16:14-26 Henderson SW et al. (2014) Shoot chloride exclusion and salt tolerance in grapevine is associated with differential ion transporter expression in roots, BMC Plant Biology 14:273 Gong H et al. (2011) Mechanisms of chloride transport in grapevines, Functional Plant Biology 38:1-9
Chloride Exclusion Disposition
Chloride toxicity quality assessment in Riverland Shiraz established the relationship between irrigation water chloride content, rootstock selection, and vine damage to guide management recommendations for this salinity-affected region. Visual chloride toxicity quality evaluation documented the characteristic marginal burn symptom with brown necrotic tissue extending 0.5-2.0 cm from leaf edges, symptom severity correlated with leaf age showing basal leaves with extensive marginal necrosis (mean 2.5 cm burn width) while apical leaves showed only trace symptoms (less than 0.3 cm). The progression of chloride toxicity quality through the season showed accumulation-driven intensification, with symptoms first visible at véraison when seasonal chloride accumulation exceeded tissue tolerance, and damage accelerating through harvest as continued transpiration further concentrated chloride in leaf margins. Tissue analysis quantified chloride toxicity quality with blade Cl at 1.2% in severely symptomatic leaves (toxic, threshold 0.5%), declining to 0.4% in mildly symptomatic young leaves, confirming the accumulation-driven symptom gradient. Rootstock comparison demonstrated dramatic chloride toxicity quality differences with own-rooted vines showing blade Cl of 1.4% and severe marginal burn, Ramsey-rooted vines showing 0.35% and minimal symptoms, and 1103P-rooted vines showing 0.28% and no visible damage, representing 75-80% reduction in scion Cl accumulation through rootstock exclusion from the same irrigation water (Cl 350 mg/L, approximately 10 mM). Yield impact assessment showed chloride toxicity quality causing 35% yield reduction in severely affected own-rooted vines through reduced photosynthetic leaf area and early defoliation, while Ramsey-rooted vines maintained full yield demonstrating the economic value of rootstock selection for chloride management.
Chloride toxicity quality is a nutrient toxicity symptom quality that inheres in grapevine leaves and represents the characteristic damage pattern resulting from chloride accumulation to toxic levels, with symptoms appearing as marginal leaf burn ("salt burn") where the leaf edge becomes brown and necrotic, progressing from older to younger leaves as chloride accumulates through the transpiration stream, and in severe cases leading to extensive leaf necrosis, premature defoliation, and reduced fruit quality. This quality captures the grapevine response to chloride excess, recognizing that chloride toxicity is often the most damaging component of salinity stress in viticulture due to grapevine's sensitivity to chloride and the mobility of chloride in the plant that efficiently transports it to leaves where damage occurs. The ontological significance of chloride toxicity quality lies in its prevalence as the primary cause of visible salt damage in irrigated vineyards, its direct relationship to irrigation water chloride content and rootstock chloride exclusion capacity, and its utility as an indicator of salinity management needs. The biological basis of chloride toxicity symptoms encompasses chloride's high mobility in both xylem and phloem that enables rapid transport to and accumulation in leaves, the direct cellular damage from chloride accumulation including interference with enzyme function and membrane integrity, and the characteristic marginal distribution resulting from chloride concentration at sites of transpiration water loss. The symptom progression from older to younger leaves reflects the cumulative accumulation over time, with older leaves having transpired more water and thus accumulated more chloride than recently expanded leaves, creating the basipetal-to-acropetal symptom gradient characteristic of mobile ion toxicity. The rootstock influence on chloride toxicity is well-documented, with chloride-excluding rootstocks providing substantial protection by limiting chloride loading into the xylem, enabling successful grape production with irrigation water chloride levels that would cause severe damage to vines on chloride-accumulating rootstocks.
Chloride_Toxicity_Quality is_a Nutrient_Toxicity_Symptom_Quality, representing the characteristic marginal leaf burn from Cl accumulation via transpiration stream. This class addresses the specific Cl toxicity syndrome, often the primary visible component of salinity damage. Blade Cl threshold for toxicity ~0.5% (varies by cultivar). Rootstock Cl exclusion is highly effective (Ramsey, 1103P, 140Ru >> own-rooted, Schwarzmann). High Cl mobility produces rapid transport to leaves and marginal accumulation. Older leaves show more severe symptoms due to longer accumulation time. Often co-occurs with Na toxicity from saline irrigation.
Chloridschaden
Chloridtoxizität
Cl toxicity
Cl-Überschuss-Symptome
Salzverbrennung
chloride damage
chloride excess symptoms
salt burn
Walker RR et al. (2010) Rootstock effects on salt tolerance of irrigated field-grown grapevines. Australian Journal of Grape and Wine Research 16:174-180 Tregeagle JM et al. (2006) Chloride analysis of grapevine tissues and the relationship with other ionic concentrations. Australian Journal of Grape and Wine Research 12:198-203 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345
Chloride Toxicity Quality
Chlorophyll a/b ratio assessment in vertically shoot-positioned Riesling canopy reveals light adaptation gradient, with exterior fully exposed leaves showing ratio of 3.4 indicating small antenna systems adapted for high light, mid-canopy partially shaded leaves showing ratio of 2.9 reflecting intermediate adaptation, and interior heavily shaded leaves showing ratio of 2.3 with expanded antenna systems compensating for low light, demonstrating how a/b ratio maps the light environment history of individual leaves and indicates their photosynthetic adaptation status.
Chlorophyll a/b ratio quality is a light capture efficiency quality that represents the molar proportion of chlorophyll a to chlorophyll b in grapevine leaves, characterizing the balance between reaction center-associated pigments (chlorophyll a dominant) and light-harvesting antenna pigments (chlorophyll b enriched) that indicates photosynthetic apparatus composition and adaptation to light environment. Within the BFO framework, this quality specifically addresses the chlorophyll type balance dimension of light capture, distinguishing the ratio between chlorophyll types from total chlorophyll content while recognizing that the a/b ratio reflects fundamental differences in thylakoid organization with implications for photosynthetic efficiency under different light conditions. The ontological significance lies in chlorophyll a/b ratio providing diagnostic information about photosynthetic apparatus structure that total chlorophyll content cannot reveal, where lower ratios (more chlorophyll b) indicate larger antenna systems adapted for efficient light capture under shade while higher ratios (more chlorophyll a) indicate smaller antennae adapted for high light where light harvesting is not limiting. The biochemical basis for chlorophyll a/b ratio reflects the differential distribution of these pigments in photosynthetic complexes, with chlorophyll a present in both reaction centers and antennae while chlorophyll b is restricted to light-harvesting complexes (LHCI and LHCII), such that the ratio indicates the proportion of total chlorophyll invested in antenna versus core complexes. The environmental and developmental factors affecting chlorophyll a/b ratio include light intensity during leaf development (primary determinant, with shade reducing ratio), leaf age (ratio typically declining slightly with age), and nitrogen status (affecting antenna protein expression), with grapevine sun leaves typically showing ratios of 3.0-3.5 while shade leaves show ratios of 2.2-2.8 reflecting their larger light-harvesting antennae. Understanding chlorophyll a/b ratio quality enables assessment of leaf light adaptation status, diagnosis of canopy light environment from leaf properties, and interpretation of photosynthetic capacity in context of antenna size and organization.
Chlorophyll_a_b_Ratio_Quality addresses the balance between chlorophyll types as an indicator of photosynthetic apparatus structure and light adaptation. This quality provides information about antenna size and leaf light history that complements total chlorophyll content. The class supports canopy light environment assessment and interpretation of photosynthetic adaptation across canopy positions.
Antennengröße-Indikator
Chl a/b ratio
Chl-a/b-Ratio
Chlorophyll-a/b-Verhältnis
LHCII content proxy
LHCII-Gehalt
antenna size indicator
chlorophyll ratio
Lichtenthaler HK et al. (2007) Chlorophylls and carotenoids: pigments of photosynthetic biomembranes. Methods in Enzymology 148: 350-382 Evans JR (1989) Photosynthesis and nitrogen relationships in leaves of C₃ plants. Oecologia 78(1): 9-19
Chlorophyll a b Ratio Quality
Chlorophyll content monitoring in deficit-irrigated Cabernet Sauvignon tracks seasonal dynamics from early season values of 520 μmol/m² (SPAD 48) indicating adequate nitrogen and full photosynthetic capacity, through mid-season slight decline to 480 μmol/m² (SPAD 45) as moderate water stress limits nitrogen uptake, to late-season senescence-driven decline to 320 μmol/m² (SPAD 35) coinciding with remobilization of nitrogen to developing fruit, with this seasonal chlorophyll trajectory indicating appropriate vine balance where moderate decline supports quality fruit composition without premature capacity loss.
Chlorophyll content quality is a light capture efficiency quality that represents the concentration of chlorophyll pigments in grapevine leaves, characterizing the primary photosynthetic pigment pool that determines light absorption capacity and serves as an indicator of leaf nitrogen status, photosynthetic potential, and overall plant health. Within the BFO framework, this quality specifically addresses the chlorophyll concentration dimension of light capture, distinguishing total chlorophyll content from the ratio between chlorophyll types while recognizing that chlorophyll concentration directly determines the light-absorbing capacity of leaves and correlates strongly with photosynthetic capacity and nitrogen nutrition. The ontological significance lies in chlorophyll content serving as both a determinant of photosynthetic capacity and an easily measurable indicator of leaf and vine physiological status, where chlorophyll concentration provides a non-destructive proxy for nitrogen status and photosynthetic potential enabling rapid assessment of large vineyard areas through spectral sensing technologies. The biochemical basis for chlorophyll function involves the tetrapyrrole structure with magnesium center that absorbs light primarily in blue (430 nm) and red (662 nm) regions, with absorbed energy transferred through antenna systems to reaction centers where photochemistry initiates, and with chlorophyll a serving as the primary photochemically active pigment while chlorophyll b functions primarily in light harvesting and energy transfer. The environmental and developmental factors affecting chlorophyll content include nitrogen availability (strong positive correlation as chlorophyll-binding proteins represent major leaf nitrogen investment), iron and magnesium nutrition (essential for chlorophyll synthesis), light environment (shade increasing chlorophyll per unit area), temperature (affecting synthesis and degradation rates), and leaf age (content increasing to full expansion then declining through senescence). The measurement approaches for chlorophyll content include destructive extraction and spectrophotometry (reference method), portable chlorophyll meters (SPAD, Dualex providing rapid non-destructive estimates), and remote sensing (vegetation indices like NDVI correlating with chlorophyll content across canopy scales). Understanding chlorophyll content quality enables nitrogen status assessment, photosynthetic capacity estimation, and early detection of stress or deficiency affecting vine productivity and fruit quality potential.
Chlorophyll_Content_Quality addresses the total chlorophyll pigment pool as a determinant of light absorption and indicator of nitrogen status. This quality is readily measured non-destructively, enabling broad application in precision viticulture. The class supports nitrogen management, stress detection, and photosynthetic capacity assessment through chlorophyll monitoring.
Blattgrün
Chlorophyllgehalt
Chlorophyllkonzentration
Pigmentgehalt
chlorophyll concentration
chlorophyll level
leaf greenness
Poni S et al. (2018) Grapevine quality: a multiple choice issue. Scientia Horticulturae 234: 445-462 Steele MR et al. (2008) Nondestructive estimation of leaf chlorophyll content in grapes. American Journal of Enology and Viticulture 59(3): 299-305
Chlorophyll Content Quality
In Chardonnay vines in Côte de Beaune, Burgundy during October-November, chlorophyll degradation reduces leaf chlorophyll content from 485 to 28 µg/g fresh weight over 25 days, with pheophorbide oxygenase activity peaking at 15 nmol/min/g fresh weight during mid-senescence, leaf color transitioning from green through yellow-green to uniform yellow, and carotenoid-to-chlorophyll ratio increasing from 0.25 to 4.8 as chlorophyll elimination reveals lutein and beta-carotene pigments under cool autumn conditions averaging 10°C.
Chlorophyll degradation describes the enzymatic catabolism of chlorophyll molecules in Vitis vinifera leaves during senescence, wherein the pheophytinase-pheophorbide oxygenase pathway systematically converts green chlorophyll pigments to colorless linear tetrapyrrole catabolites that are sequestered in vacuoles, producing the characteristic transition from green to yellow leaf coloration as chlorophyll loss reveals underlying carotenoid pigments over a period of 2-4 weeks during the senescence program. This process involves sequential enzymatic reactions beginning with chlorophyllase or pheophytinase removing the phytol tail from chlorophyll, followed by magnesium-dechelatase removing the central magnesium ion, pheophorbide a oxygenase opening the porphyrin macrocycle generating red chlorophyll catabolite, and red chlorophyll catabolite reductase reducing this intermediate to primary fluorescent chlorophyll catabolite that undergoes further modifications before vacuolar sequestration. The quantitative significance of chlorophyll degradation reflects the substantial chlorophyll content of mature grapevine leaves, typically 300-600 µg/g fresh weight comprising approximately 3-5% of total leaf dry weight, with complete degradation releasing this nitrogen-rich porphyrin pool for potential amino acid recovery while eliminating the phototoxic risk of free chlorophyll intermediates that could generate damaging reactive oxygen species under continued light exposure. The regulation of chlorophyll degradation involves transcriptional activation of catabolic enzyme genes by senescence-associated transcription factors, with stay-green mutants in other plant species demonstrating that chlorophyll retention can be achieved through reduced expression of specific pathway enzymes while other senescence processes continue, though such mutations have not been extensively characterized in grapevine. The temporal progression of chlorophyll degradation within individual leaves shows characteristic patterns, with degradation typically initiating in interveinal regions while areas adjacent to major veins retain chlorophyll longer due to continued photosynthate export demands, and complete chlorophyll loss achieved only in late senescence stages when phloem function declines and export requirements diminish. Environmental factors influence chlorophyll degradation rates, with cool temperatures of 5-15°C accelerating degradation through enhanced enzyme activities in the temperature-sensitive pheophorbide oxygenase reaction, while continued warm conditions may slow degradation maintaining green coloration longer into autumn. The visual progression of chlorophyll degradation produces the autumn color display characteristic of grapevine canopies, transitioning from uniform green through patchy yellow-green as degradation proceeds heterogeneously, to uniform yellow when chlorophyll is largely eliminated revealing carotenoid pigments, with some varieties additionally synthesizing anthocyanins producing red autumn coloration that serves photoprotective functions during the extended senescence period.
Chlorophyll_Degradation is subclass of Leaf_Senescence representing pigment catabolism specifically. Distinguishes from nutrient remobilization by focusing on chlorophyll pathway. Sibling to Leaf_Senescence_Initiation, Nutrient_Remobilization, and Progressive_Leaf_Senescence. Pheophytinase-pheophorbide oxygenase pathway. Duration 2-4 weeks. Produces yellow coloration through carotenoid exposure.
Blatt-Entgrünung
Chlorophyll-Abbau
Chlorophyll-Katabolismus
Chlorophyll-Verlust
chlorophyll breakdown
chlorophyll catabolism
chlorophyll loss
leaf degreening
Hörtensteiner S (2006) Chlorophyll degradation during senescence. Annual Review of Plant Biology 57: 55-77 Pruzinská A et al (2005) Chlorophyll breakdown in senescent Arabidopsis leaves. Plant Physiology 139: 52-63 Keller M (2020) The Science of Grapevines 3rd Ed pp 88-90
Chlorophyll Degradation
Chlorophyll fluorescence quality monitoring in Pinot Noir vineyard employs predawn Fv/Fm measurement to assess overnight recovery from previous day's stress, with healthy vines showing Fv/Fm of 0.82-0.84 indicating full PSII recovery, while vines in drought-stressed blocks display reduced Fv/Fm of 0.72-0.75 indicating chronic photoinhibition from incomplete overnight recovery, with this rapid non-destructive assessment enabling targeted irrigation intervention in affected blocks before visible stress symptoms or yield impact develop.
Chlorophyll fluorescence quality is a photosynthetic capacity quality that represents the characteristics of light re-emitted by chlorophyll molecules following photon absorption, characterizing the fate of absorbed light energy among photochemistry, heat dissipation, and fluorescence pathways that provides non-invasive diagnostic information about photosynthetic apparatus function and stress status. Within the BFO framework, this quality addresses the fluorescence dimension of photosynthetic performance, distinguishing fluorescence emission from productive photochemistry while recognizing that fluorescence yield inversely relates to photochemical efficiency, enabling inference about photosystem function from fluorescence measurements without directly measuring carbon fixation. The ontological significance lies in chlorophyll fluorescence providing a rapid, non-destructive probe of photosynthetic apparatus health that has revolutionized plant stress diagnostics, where fluorescence parameters reveal photosystem II efficiency, electron transport capacity, and photoprotective mechanisms in seconds rather than the minutes required for gas exchange measurement, enabling high-throughput screening and real-time stress monitoring in vineyard contexts. The photophysical basis for chlorophyll fluorescence involves the competitive de-excitation pathways for absorbed light energy, with excited chlorophyll molecules returning to ground state through photochemistry (productive electron transfer), heat dissipation (non-photochemical quenching), or fluorescence emission (radiative decay), and with the quantum yields of these pathways summing to unity such that changes in one pathway necessarily affect others. The measurement technology for chlorophyll fluorescence employs pulse-amplitude modulated (PAM) fluorometry that distinguishes variable fluorescence from background by applying saturating light pulses that transiently close all PSII reaction centers, revealing maximum fluorescence (Fm) from which parameters including Fv/Fm, ΦPSII, qP, and NPQ are calculated. The stress sensitivity of fluorescence parameters makes chlorophyll fluorescence particularly valuable for early stress detection, with changes in fluorescence yield and quenching patterns occurring before visible symptoms or growth effects, enabling proactive management intervention based on fluorescence-based stress diagnosis. Understanding chlorophyll fluorescence quality enables rapid photosynthetic health assessment, stress detection and diagnosis, and high-throughput phenotyping for vineyard monitoring and breeding programs.
Chlorophyll_Fluorescence_Quality serves as the parent class for specific fluorescence parameters (Fv/Fm, qP, NPQ, ETR). This quality enables rapid, non-destructive photosynthetic assessment through fluorescence measurement. The class supports stress diagnostics and high-throughput monitoring through fluorescence-based evaluation of photosystem function.
Chl fluorescence
Chlorophyllfluoreszenz
Fluoreszenzausbeute
Fluoreszenzemission
PAM fluorescence
PAM-Fluoreszenz
fluorescence emission
fluorescence yield
Maxwell K, Johnson GN (2000) Chlorophyll fluorescence — a practical guide. Journal of Experimental Botany 51(345): 659-668 Baker NR (2008) Chlorophyll fluorescence: a probe of photosynthesis in vivo. Annual Review of Plant Biology 59: 89-113
Chlorophyll Fluorescence Quality
The chloroplast DNA sequence analysis of 1,201 Vitis vinifera cultivars (Arroyo-García et al. 2006) identified four major chloroplast haplotypes (A, B, C, D) defined by polymorphisms at 5 cpSSR loci and 3 SNP positions in the trnL-trnF intergenic spacer — haplotype C predominates in Western European cultivars (Pinot Noir, Cabernet Sauvignon, Syrah, Chardonnay, 65% of cultivars surveyed), haplotype D characterizes many Eastern Mediterranean and Caucasian cultivars (Rkatsiteli, Saperavi, 20% of surveyed cultivars), haplotype A occurs primarily in Iberian cultivars (Tempranillo, Albariño, 10%), and haplotype B is found in Muscat-type cultivars and some Italian varieties (5%) — this maternal haplotype distribution provides evidence for multiple independent domestication events and maternal lineage contributions from distinct wild Vitis vinifera subsp. sylvestris populations across the species' natural range.
A DNA sequence derived from the chloroplast genome of Vitis vinifera, a circular double-stranded DNA molecule of approximately 160,928 base pairs located in the chloroplast organelle, maternally inherited in grapevine, encoding genes essential for photosynthesis, ribosomal RNA, and transfer RNA, and providing phylogeographic markers used to trace maternal lineage, domestication pathways, and the geographic origins of cultivar groups through chloroplast haplotype analysis.
Chloroplast_DNA_Sequence provides the maternally inherited genetic marker system that complements the biparentally inherited nuclear genome for evolutionary and phylogeographic studies of grapevine. The chloroplast genome is inherited exclusively through the maternal line in Vitis (as in most angiosperms), meaning that all cultivars sharing a chloroplast haplotype descend through an unbroken maternal lineage from a common female ancestor — this uniparental inheritance pattern, combined with the relatively slow mutation rate of cpDNA compared to nuclear DNA, makes chloroplast sequences particularly informative for deep evolutionary questions about grapevine domestication geography and timing: the distribution of chloroplast haplotypes across modern cultivars and wild V. vinifera subsp. sylvestris populations has provided key evidence supporting the current model of multiple domestication centers in the Near East and subsequent introgression from local wild populations during the westward spread of viticulture into Europe. The complete chloroplast genome of V. vinifera (Jansen et al. 2006, 160,928 bp) contains 130 genes arranged in the typical angiosperm quadripartite structure (large single copy, small single copy, and two inverted repeat regions).
Chloroplasten-DNA-Sequenz
Plastiden-Genomsequenz
chloroplast genomic data
cpDNA sequence
cpDNA-Daten
plastid genome sequence
Arroyo-García R et al. (2006) Multiple origins of cultivated grapevine (Vitis vinifera L. ssp. sativa) based on chloroplast DNA polymorphisms, Molecular Ecology 15: 3707-3714 Jansen RK et al. (2006) Analysis of 81 genes from 64 plastid genomes resolves relationships in angiosperms and identifies genome-scale evolutionary patterns, Proceedings of the National Academy of Sciences 103: 19369-19374 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519
Chloroplast DNA Sequence
Chlorosis pattern quality assessment for nitrogen deficiency in Central Valley Thompson Seedless employed systematic canopy evaluation to map nitrogen status variation across a 40-hectare vineyard with heterogeneous soil properties. The chlorosis pattern quality characteristic of nitrogen deficiency was identified by its uniform whole-leaf yellowing affecting all leaf tissues equally rather than the interveinal pattern with green veins characteristic of iron or magnesium deficiency, and by its initiation on basal leaves with progression toward shoot tips reflecting nitrogen remobilization from old to young tissues. Chlorosis pattern quality scoring using a chlorophyll meter (SPAD) quantified the deficiency gradient, with SPAD readings of 18-22 in severely chlorotic basal leaves (nitrogen deficient), 28-32 in mildly chlorotic mid-shoot leaves, and 35-40 in youngest leaves still receiving remobilized nitrogen (normal range greater than 35). The spatial distribution of chlorosis pattern quality mapped onto soil texture variation, with severe chlorosis concentrated on sandy areas with low organic matter and cation exchange capacity limiting nitrogen retention, while heavier-textured areas showed adequate canopy color. Tissue analysis confirmed chlorosis pattern quality interpretation, with petiole nitrogen at 0.35% in severely chlorotic zones versus 0.75% in green zones (sufficiency range 0.6-1.2%). The chlorosis pattern quality mapping enabled precision nitrogen management with variable-rate fertigation applying 45 kg N/ha to deficient sandy zones versus 15 kg N/ha maintenance rates on adequate heavier soils.
Chlorosis pattern quality in nitrogen deficiency is a nitrogen deficiency symptom quality that inheres in grapevine leaves and represents the characteristic distribution and progression of yellowing resulting from reduced chlorophyll synthesis when nitrogen availability limits the production of the nitrogen-containing porphyrin ring structure central to chlorophyll molecules, with nitrogen deficiency chlorosis showing a distinctive pattern of uniform whole-leaf yellowing beginning on oldest leaves and progressing acropetally as nitrogen is remobilized from mature tissues to support new growth. This quality captures the most visually diagnostic symptom of nitrogen deficiency, recognizing that the chlorosis pattern provides critical information distinguishing nitrogen deficiency from other causes of leaf yellowing including other nutrient deficiencies, viral infections, and environmental stresses that produce different chlorosis patterns. The ontological significance of chlorosis pattern quality in nitrogen deficiency lies in its diagnostic utility for field identification of nitrogen limitation, with the uniform interveinal and vein chlorosis producing a generally pale appearance distinct from the interveinal chlorosis with green veins characteristic of iron, manganese, or magnesium deficiency, enabling preliminary diagnosis before tissue analysis confirmation. The biological basis of nitrogen-deficiency chlorosis encompasses the nitrogen content of chlorophyll molecules where each chlorophyll contains four nitrogen atoms in its porphyrin ring, the dependence of chlorophyll synthesis on amino acid supply for porphyrin biosynthesis, and the degradation of existing chlorophyll in older leaves as nitrogen is remobilized to younger tissues, with the remobilization pattern producing the characteristic basal-to-apical symptom progression as oldest leaves are sacrificed to supply nitrogen to developing tissues. The intensity of chlorosis correlates with deficiency severity, ranging from subtle pale green coloration in mild deficiency through yellow-green in moderate deficiency to bright yellow or even white in severe deficiency where chlorophyll content approaches zero, with the progression providing information about deficiency duration and intensity.
Chlorosis_Pattern_Quality_(N) is_a Nitrogen_Deficiency_Symptom_Quality, representing the characteristic uniform yellowing pattern resulting from nitrogen-limited chlorophyll synthesis. This class addresses the visual chlorosis symptom that is most diagnostic for field identification. The uniform whole-leaf yellowing distinguishes N deficiency from interveinal chlorosis of Fe, Mn, or Mg deficiency. Basal-to-apical progression reflects nitrogen remobilization from old to young tissues. SPAD meter readings provide quantitative chlorosis assessment. Chlorosis intensity correlates with deficiency severity and duration.
Chlorose-Muster bei N-Mangel
N-deficiency yellowing
N-induzierte Gelbfärbung
Stickstoff-Chlorose
gleichmäßige Blattvergilbung
nitrogen chlorosis
nitrogen-induced yellowing
uniform leaf chlorosis
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Porro D et al. (2001) Use of SPAD meter in diagnosis of nutritional status in apple and grapevine. Acta Horticulturae 564:243-252 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication
Chlorosis Pattern Quality (N)
Citric acid content for Pinot Noir from Burgundy (2024 vintage) measures 0.22 g/L at harvest, within the typical range for Vitis vinifera must (0.1-0.5 g/L), establishing the natural baseline against which any post-fermentation citric acid addition for tartrate stabilization purposes can be analytically distinguished. Citric acid content for Sauvignon Blanc from Sancerre (2024 vintage) records 0.35 g/L, at the higher end of the natural range, contributing measurably to the total acidity value and providing a minor but perceptible contribution to the fresh, citrus-like acid impression characteristic of Loire Sauvignon. Citric acid content for Welschriesling from Neusiedlersee (2024 vintage) documents 0.15 g/L in the healthy must versus 0.08 g/L in the botrytized Beerenauslese lot, the reduction reflecting preferential metabolism of citric acid by Botrytis cinerea during noble rot development.
Citric acid content is an acidity data entity that records the concentration of citric acid (2-hydroxypropane-1,2,3-tricarboxylic acid) in grape must, determined by enzymatic assay (citrate lyase), HPLC, or FTIR estimation, expressed in grams per liter (g/L) or milligrams per liter (mg/L), representing a quantitatively minor but analytically significant organic acid component of grape berries whose low natural concentration in must (typically 0.1-0.5 g/L) provides baseline data for monitoring post-fermentation citric acid changes and for detecting exogenous citric acid additions regulated under various national winemaking codes.
Citric acid occupies a paradoxical position in grape and wine chemistry — quantitatively minor in must (typically an order of magnitude below tartaric and malic acid concentrations), yet analytically and regulatorily significant for several reasons. First, citric acid is almost completely consumed during malolactic fermentation by Oenococcus oeni and Lactobacillus species, meaning that post-MLF citric acid levels near zero are normal while elevated levels may indicate exogenous addition. Second, the metabolic products of bacterial citric acid degradation include diacetyl (2,3-butanedione), the compound responsible for the buttery flavor associated with MLF — making the initial must citric acid concentration a predictor of post-MLF diacetyl production potential. Third, citric acid addition to wine is a regulated winemaking practice in many jurisdictions — the EU permits addition up to 1 g/L for tartrate stabilization purposes, while some countries restrict or prohibit its use, making must baseline citric acid data important for compliance documentation. The biosynthesis of citric acid in grape berries occurs through the TCA cycle with accumulation primarily during early berry development, paralleling tartaric acid timing. Botrytis cinerea preferentially metabolizes citric acid during berry infection, and the citric acid concentration in botrytized must is characteristically reduced, providing an additional biochemical marker of noble rot intensity. Analytical determination by enzymatic assay (citrate lyase coupled with malate dehydrogenase and lactate dehydrogenase) offers high specificity, while HPLC provides simultaneous quantification within the organic acid profile.
Citratgehalt
Citronensäuregehalt
Zitronensäurebestimmung
Zitronensäuregehalt
Zitronensäurekonzentration
citrate content
citrate level
citric acid analysis
citric acid concentration
must citrate
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2, 2nd Ed, Wiley, pp.: 15-22 Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 405-410 Nielsen JC, Richelieu M (1999) Control of flavor development in wine during and after malolactic fermentation by Oenococcus oeni. Applied and Environmental Microbiology 65(2): 740-745
Citric Acid Content
Citric acid content quality monitoring in Languedoc rosé production targets the fresh, citrus-like character contributed by moderate citric levels (0.3-0.5 g/L), with winemakers balancing the desire for this brightness against the risk of diacetyl formation if citric acid is metabolized during unintended malolactic fermentation careful temperature control and sulfur dioxide management prevent bacterial activity that would convert citric acid to volatile compounds, preserving the clean, refreshing character that defines premium Provence-style rosé wines.
Citric_Acid_Content_Quality represents the chemical quality characterizing the concentration of citric acid (2-hydroxypropane-1,2,3-tricarboxylic acid, C₆H₈O₇) in grape berry tissue. Citric acid is a minor component of grape acidity, typically present at concentrations of 0.1-0.7 g/L at harvest, contributing less than 5% of total acidity but adding distinctive fresh, citrus-like flavor notes even at low concentrations. The citric acid content of grapes is substantially lower than many other fruits, reflecting the metabolic pathways favoring tartaric and malic acid accumulation in Vitis species. Citric acid is synthesized through the tricarboxylic acid cycle, where it represents the first product of acetyl-CoA condensation with oxaloacetate, but it does not accumulate to high levels in grape vacuoles. Unlike tartaric acid but like malic acid, citric acid can be metabolized during malolactic fermentation, with lactic acid bacteria converting citric acid to acetic acid and diacetyl, potentially increasing volatile acidity and adding buttery notes. This metabolism makes citric acid content relevant to malolactic fermentation management. In winemaking, citric acid may be added for acidification in some jurisdictions, though it is prohibited in the European Union for wines due to concerns about subsequent bacterial metabolism. Citric acid complexes effectively with iron and copper, contributing to metal stability in wine. The relatively low citric acid content of grapes compared to citrus fruits and many other tree fruits reflects the distinctive organic acid metabolism of grapevines. Measurement employs specific enzymatic assays using citrate lyase or HPLC separation from other organic acids.
Citric_Acid_Content_Quality addresses a minor but flavor-active acid component. The tricarboxylic structure provides three carboxyl groups capable of releasing hydrogen ions, contributing to buffering capacity. Citric acid's position in the TCA cycle as a central metabolic intermediate explains its presence in all plant tissues while its low accumulation in grapes reflects rapid cycling rather than vacuolar storage. The regulatory prohibition on citric acid addition in EU wines (with specified exceptions) reflects concerns about bacterial metabolism producing acetic acid and diacetyl rather than toxicological issues.
Citrat
Zitronensäure
citric level
lemon acid quality
tricarboxylic acid concentration
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed, John Wiley and Sons pp.: 3-25 Kliewer WM (1966) Sugars and Organic Acids of Vitis vinifera. Plant Physiology 41(6): 923-931
Citric Acid Content Quality
Citronellol content quality in aged Riesling increases relative to linalool and geraniol during bottle maturation, with citronellol rising from 30-60 μg/L in young wine to 80-150 μg/L in decade-old examples through reduction of geraniol this compositional shift contributes to the evolution of Riesling aroma from youthful floral-citrus toward the complex citronella-lime character of mature wines, demonstrating how citronellol's greater stability compared to unsaturated terpenes affects the aromatic trajectory of age-worthy aromatic whites.
Citronellol_Content_Quality represents the chemical quality characterizing the concentration of citronellol (3,7-dimethyl-6-octen-1-ol) in grape berries and wine, capturing the saturated monoterpene alcohol that contributes citrus-rose aroma notes distinct from the unsaturated terpene alcohols linalool, geraniol, and nerol. Citronellol differs from geraniol by reduction of the 2,3 double bond, removing the conjugation that affects the sensory character of unsaturated terpene alcohols. This saturation produces a somewhat different aroma quality, described as citronella-like, citrus-rose, with a fresh, green character distinct from the heavier floral notes of geraniol. Citronellol exists as two enantiomers, (+)-citronellol and (−)-citronellol, with different sensory properties and varying proportions in grape-derived material. In grape berries and wines, citronellol typically occurs at moderate concentrations in aromatic varieties, ranging from 10-50 μg/L in neutral varieties through 50-150 μg/L in semi-aromatic cultivars to 100-400 μg/L in intensely aromatic Muscat varieties. The sensory threshold of citronellol in wine is approximately 40-100 μg/L, placing it intermediate between the low-threshold linalool and high-threshold nerol. Citronellol can arise in grapes through direct biosynthesis or through chemical or enzymatic reduction of geraniol during winemaking, making its concentration in wine potentially higher than in grapes. During wine aging, citronellol is relatively stable compared to the unsaturated terpene alcohols, contributing to its increased relative importance in aged aromatic wines as linalool and geraniol decline through oxidation and rearrangement reactions.
Citronellol_Content_Quality addresses the saturated monoterpene alcohol whose stability distinguishes it from the unsaturated geraniol and nerol. The metabolic relationship between geraniol and citronellol (reduction) means citronellol can increase during winemaking and aging even as its precursor declines. The chiral nature of citronellol (two enantiomers with different odor qualities) affects sensory interpretation comprehensive analysis may include enantiomeric determination. Citronellol's intermediate sensory threshold places it between the highly odor-active linalool and the minimally perceptible nerol.
3,7-dimethyl-6-octen-1-ol concentration
Citronellol-Konzentration
Citronellolgehalt
Citronellylverbindungsgehalt
Zitrus-Terpenalkoholgehalt
citronellol level
citronellyl compound content
citrus terpene alcohol quality
Simpson RF (1979) Some Important Aroma Components of White Wine. Food Technology in Australia 31(12): 516-522 Rapp A and Mandery H (1986) Wine Aroma. Experientia 42(8): 873-884
Citronellol Content Quality
Citrus aroma profiling in Clare Valley Riesling identifies the signature lime character (intensity 7.2 on 10-point scale) from monoterpene composition including α-terpineol and citronellol, complemented by lemon zest notes (5.8) and subtle grapefruit (3.2), with the lime dominance distinguishing Clare Valley expression from Alsace Riesling where lemon and citrus blossom notes predominate, demonstrating how specific citrus subcategory balance creates regional aromatic identity within this varietal.
Citrus aroma quality is a fruity aroma quality that represents olfactory perception evoking citrus fruit-associated scents including lemon, lime, grapefruit, orange, and citrus zest notes, characterizing the fresh, bright, acidic-associated aromatic impression commonly found in white wines and contributing to perceived freshness and vibrancy. Within the BFO framework, this quality specifically addresses the citrus subcategory of fruity aroma distinguished by its association with perceived acidity and freshness, recognizing that citrus character often correlates with crisp wine styles and contributes to the impression of liveliness even when actual acidity levels are moderate. The ontological significance lies in citrus aroma providing key varietal markers for certain white varieties (Riesling's lime, Sauvignon Blanc's grapefruit, Albariño's lemon) and serving as a freshness indicator across wine styles, where citrus brightness suggests youth and vitality while its absence or transformation to candied citrus may indicate age or oxidation. The compositional basis for citrus aroma includes terpene compounds (limonene, citral contributing direct citrus character), thiols (3MH contributing grapefruit in Sauvignon Blanc), certain esters (ethyl citrate associated with citrus), and terpene-derived compounds that provide lemon-lime notes in Riesling and related varieties. The distinction among citrus subcategories reflects both perceptual differences and compositional variation, with lemon-lime notes typically associated with certain terpenes, grapefruit specifically linked to 3MH thiol, and orange-tangerine notes arising from different compound combinations. The interaction between citrus aroma and actual wine acidity demonstrates cross-modal perception, where citrus aromatic character enhances the impression of acidity through learned association, making citrus-scented wines seem more refreshing than equivalent acidity levels without citrus character. Understanding citrus aroma quality enables varietal identification where citrus is a signature note, freshness assessment based on citrus brightness, and wine style prediction from the specific citrus subcategory expressed.
Citrus_Aroma_Quality addresses the citrus subcategory of fruity aroma, encompassing lemon, lime, grapefruit, orange, and related citrus notes. This quality shows strong cross-modal interaction with acidity perception, enhancing freshness impression. The specific citrus notes expressed depend on volatile compound composition and provide varietal and regional markers particularly important in aromatic white wine evaluation.
Zitrusaroma
Zitruscharakter
Zitrusduft
Zitrusnoten
citrus character
citrus notes
citrus-like aroma
lemon-lime scent
Tominaga T et al. (1998) Identification of new volatile thiols in the aroma of Vitis vinifera L. var. Sauvignon blanc wines. Flavour and Fragrance Journal 13(3): 159-162 Rapp A (1998) Volatile flavour of wine: correlation between instrumental analysis and sensory perception. Nahrung 42(6): 351-363
Citrus Aroma Quality
Clarity description for Chablis Premier Cru (2023 vintage) records "brilliant" — star-bright with no visible haze or particulate when examined against a point light source, consistent with the cold-stabilized and cross-flow filtered processing specified in the winemaking protocol, and confirming complete protein and tartrate stability. Clarity description for unfiltered Barolo Riserva (2017 vintage) documents "clear with fine sediment" — transparent wine with a thin deposit of polymerized tannin-anthocyanin precipitate visible along the lower surface of the horizontally stored bottle, entirely expected in an unfiltered Nebbiolo of this age and considered a positive quality indicator reflecting minimal processing intervention. Clarity description for Grüner Veltliner (2024 vintage, bottled unfined and unfiltered) records "clear to bright" with subtle opalescence perceptible under direct lighting at certain angles, reflecting the colloidal mannoprotein content from 6 months of lees contact — intentionally preserved for textural contribution.
Clarity description is a visual description entity that documents the optical transparency and particulate status of wine as assessed by visual examination of light transmission through the tilted glass, characterized on a scale from brilliant (star-bright, no visible suspended matter) through clear (transparent, negligible haze) and slightly hazy to hazy and cloudy (significant light scattering or visible turbidity), with additional notation of specific particulate types where present (tartrate crystals, protein haze, sediment, microbial haze, colloidal precipitate), providing the visual stability assessment that indicates wine condition, filtration status, and potential faults requiring investigation.
Clarity description serves simultaneously as a quality assessment, a stability diagnostic, and an indicator of winemaking philosophy — brilliant clarity was historically considered an absolute quality requirement, but the growing natural wine and minimal-intervention movement has reframed slight haze in unfiltered wines as an acceptable or even positive quality attribute, creating a philosophical tension in clarity assessment. The physical basis of wine clarity involves the interaction of light with dissolved and suspended matter — truly clear wine transmits light without significant scattering, while haze arises from colloidal particles (typically 0.1-1.0 µm diameter) that scatter light through the Tyndall effect, and visible turbidity indicates larger particles (>1.0 µm) or microbiological contamination. The principal causes of clarity defects in wine include protein haze (heat-unstable grape proteins, predominantly thaumatin-like proteins and chitinases in white wines, precipitating upon temperature elevation), tartrate crystals (potassium bitartrate or calcium tartrate precipitating upon cooling), microbial haze (yeast or bacterial cells from refermentation or spoilage), and colloidal instability (polysaccharide-tannin complexes in aged red wines). The distinction between expected age-related sediment (polymerized pigment precipitate in aged red wines, tartrate crystals) and fault-indicating turbidity (protein haze in young whites, microbial haze, metallic casse) is essential for correct quality assessment. Objective turbidity measurement by nephelometry (expressed in Nephelometric Turbidity Units, NTU) provides quantitative confirmation of visual assessment, with brilliant wines typically measuring 4 NTU. Filtration technology (cross-flow, membrane, diatomaceous earth, pad filtration) provides the primary intervention for achieving clarity, with the level of filtration reflecting a deliberate quality trade-off — tight filtration (0.45 µm nominal) ensures microbiological stability and brilliant clarity but may remove colloidal components contributing to mouthfeel, while looser or no filtration preserves textural complexity at the risk of clarity defects and refermentation.
Klarheit
Klarheitsbeschreibung
Limpidität
Trübungsbewertung
brilliance
haze assessment
limpidity
turbidity description
wine clarity
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed, Wiley, pp.: 301-370 Waters EJ et al. (2005) Heat haze characteristics of fractionated wine proteins. Australian Journal of Grape and Wine Research 11(2): 185-196 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 462-468
Clarity Description
Clarity quality comparison between commercial Muscadet (sterile filtered, brilliant clarity rating 10/10) and natural wine Muscadet (unfiltered, visible haze rating 5/10) demonstrates style-dependent clarity standards, with conventional example meeting commercial clarity specifications required for major retail channels while natural example's deliberate turbidity is accepted and even valued by natural wine consumers as evidence of minimal intervention, illustrating how clarity assessment must incorporate stylistic context rather than applying universal clarity standards across all wine categories.
Clarity quality is a visual appearance quality that represents the degree to which wine transmits light without scattering from suspended particles, characterizing the transparency and optical purity that indicates successful fining, filtration, and stability in conventionally produced wines or deliberate turbidity retention in unfiltered and natural wine styles. Within the BFO framework, this quality addresses the particle-related dimension of visual appearance, distinguishing clarity from color and viscosity while recognizing that clarity assessment involves judgment of both technical quality (absence of unintended haze) and stylistic appropriateness (acceptance of intentional turbidity). The ontological significance lies in clarity serving as a primary quality indicator in conventional wine production where brilliant clarity signals careful winemaking, while in natural and minimal-intervention contexts, clarity standards relax to accept the turbidity inherent in unfiltered, unfined wines, creating style-dependent quality criteria that have evolved substantially in recent decades. The physical basis for clarity involves light scattering by suspended particles, with particles larger than approximately 0.1 μm causing visible turbidity through Mie scattering, while smaller particles and dissolved compounds do not significantly affect transparency, with clarity thus indicating the presence or absence of colloidal and larger particulate matter. The compositional determinants of clarity include residual yeast cells, bacterial cells, protein-tannin complexes, tartrate crystals, and various colloidal aggregates that may remain suspended or precipitate depending on wine treatment and storage conditions, with different particle types creating different haze characteristics and stability implications. The production factors affecting clarity include fining agent selection (bentonite for protein stabilization, gelatin for tannin adjustment), filtration level (sterile filtration for complete clarity versus coarse filtration or none for natural styles), cold stabilization (precipitating tartrates before bottling), and aging time allowing natural sedimentation. Understanding clarity quality enables quality control for clarity specifications, fault detection when unexpected haze develops, and appropriate assessment that respects stylistic variation in clarity expectations.
Clarity_Quality serves as the parent class for clarity subcategories (Limpidity, Brilliance, Haziness), representing the particle-related dimension of visual assessment. This quality is strongly style-dependent, with conventional wines requiring brilliant clarity while natural and unfiltered wines accept or embrace turbidity. The class supports both quality control and context-appropriate evaluation that respects stylistic diversity in clarity expectations.
Transparenz
Weinklarheit
optical clarity
optische Reinheit
transparency
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed. John Wiley & Sons, Chichester, pp. 1-45 Waters EJ et al. (2005) Preventing protein haze in bottled white wine. Australian Journal of Grape and Wine Research 11(2): 215-225
Clarity Quality
Classification status for Grüner Veltliner in Austria is "Qualitätsrebsorte" (quality grape variety) — the highest classification tier under the Austrian Weingesetz, conferring eligibility for all quality wine production levels from Qualitätswein through Kabinett to Prädikatswein, mandatory inclusion in the national variety catalogue, and eligibility for all Austrian DAC designations that specify the cultivar. Classification status for Cabernet Sauvignon in France varies by region — classified as a "cépage principal" (principal variety) in the Bordeaux appellations (Médoc, Graves, Pessac-Léognan, Saint-Émilion, Pomerol specifications), as "cépage accessoire" (accessory variety) in certain Languedoc appellations, as "cépage autorisé" at the national level for IGP production, and not authorized in Burgundy or Alsace appellation specifications — the regionally differentiated classification reflecting the French philosophy that cultivar suitability is site-specific rather than universal.
Classification status is a regulatory information entity that documents the formal regulatory ranking, categorization, and administrative standing of grape cultivars within national and regional varietal classification systems, encompassing the hierarchy of classification categories (recommended, authorized, provisionally authorized, experimental, heritage-designated, prohibited), the criteria applied for classification decisions (agronomic performance, wine quality potential, disease resistance, regional adaptation, historical significance), and the regulatory consequences of each classification level (eligibility for quality wine production, access to planting subsidies, inclusion in appellation specifications, research priority status), providing the administrative quality hierarchy that formally ranks cultivars according to their assessed suitability and importance for the viticultural sector of each jurisdiction.
Classification status documentation captures the formal administrative hierarchy through which viticultural jurisdictions rank their cultivar portfolios — a ranking that reflects the intersection of agronomic assessment, quality evaluation, economic importance, and cultural value judgment. The classification system concept originated in the French viticultural administrative tradition, where the distinction between "cépages recommandés" (recommended varieties, actively encouraged through subsidies and planting incentives), "cépages autorisés" (authorized varieties, permitted but not promoted), and "cépages tolérés" (tolerated varieties, permitted for existing plantings but discouraged for new planting) created a three-tier hierarchy encoding the state's assessment of each cultivar's value for the national viticultural sector. The post-2008 EU simplification reduced the formal classification to a binary authorized/not-authorized distinction at the national level, but the practical multi-tier hierarchy persists through appellation-level specifications where cultivars are ranked as principal, complementary, accessory, or excluded — effectively maintaining the quality hierarchy within the appellation framework rather than the national catalogue. The criteria for classification decisions vary across jurisdictions but typically encompass agronomic suitability (yield reliability, disease resistance, climate adaptation), wine quality potential (demonstrated capacity to produce wines of recognized quality in the jurisdiction), economic viability (market demand, commercial value of the resulting wines), and cultural-historical significance (the cultivar's role in regional viticultural tradition). The four child classes of Classification_Status — Recommended, Authorized, Provisional, and Heritage — capture the principal categories of the modern classification hierarchy, from actively promoted varieties through permitted varieties and trial varieties to historically significant varieties receiving conservation-oriented protection.
Klassifizierungsstatus
Sortenklassifizierung
Sortenstatus
administrative Klassifikation
administrative classification
cultivar ranking
regulatorische Einstufung
regulatory classification
varietal classification
varietal status
European Parliament (2013) Regulation (EU) No 1308/2013, Articles 81-85 Österreichisches Weingesetz (2009) §3 and Annex 1 INAO (2024) Classification des cépages par appellation Anderson K, Aryal NR (2013) Which winegrape varieties are grown where? University of Adelaide Press
Classification Status
The clay content of the Saint-Émilion plateau soils (Bordeaux) documents high clay proportions of 35-55% in the upper 60 cm — the clay fraction is dominated by smectitic (montmorillonite) and illitic clay minerals developed from the Oligocene-Miocene limestone and molasse parent materials, producing heavy, plastic soils with high water-holding capacity (180-250 mm plant-available water), high cation exchange capacity (25-35 cmol/kg, providing substantial nutrient retention), and pronounced shrink-swell behavior (the clay soils crack deeply during summer drought, creating preferential root channels and drainage pathways, then swell and seal when rewetted by autumn rainfall). The high clay content produces the cooler, later-ripening terroir of the Saint-Émilion plateau compared to the warmer, earlier-ripening gravel soils of the Graves and Médoc — clay soils have higher heat capacity and thermal conductivity than sandy soils, warming more slowly in spring (delaying budbreak by 3-7 days) and maintaining more stable temperatures during summer (moderating diurnal temperature amplitude by 2-4°C).
A soil texture description component that records the proportion of clay-sized particles (<0.002 mm diameter) in the fine earth fraction of the vineyard soil, expressed as a percentage by weight, with clay content being the single most influential textural parameter for vineyard soil behavior — determining water-holding capacity, cation exchange capacity (nutrient retention), shrink-swell behavior, drainage impedance, and the degree of soil compaction risk that together define the soil's capacity to regulate vine water and nutrient supply across the growing season.
Clay_Content records the finest particle fraction that exerts a disproportionate influence on soil physical and chemical properties relative to its mass fraction — soils with as little as 15-20% clay behave fundamentally differently from sand-dominated soils because the enormous surface area of clay particles (50-800 m² per gram, compared to 40% clay) retain abundant water but may become waterlogged and oxygen-depleted during wet periods, with the intermediate range (20-35% clay) generally providing the optimal balance for vine health and quality.
Ton-Fraktion
Tonanteil-Messung
Tongehalt
clay fraction record
clay percentage measurement
fine particle proportion
Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873 van Leeuwen C et al. (2004) Influence of climate, soil, and cultivar on terroir, American Journal of Enology and Viticulture 55: 207-217 White RE (2003) Soils for Fine Wines, Oxford University Press Soil Survey Staff (2014) Keys to Soil Taxonomy, 12th edition, USDA-NRCS
Clay Content
A cleft graft inserting Chardonnay scion wedge into a split 5-year-old 3309C rootstock trunk bears a Cleft_Graft_Role in vineyard top-working, with the mature root system supporting rapid scion development and return to production within 2-3 years. A cleft graft of Tempranillo onto established vineyard rootstock serves in a Cleft_Graft_Role for variety conversion in Rioja, where the technique's suitability for larger-diameter rootstocks enables grafting onto trunks too large for chip or T-budding. A double cleft graft (two scions inserted into opposite sides of the cleft) of Cabernet Sauvignon bears a Cleft_Graft_Role in vineyard renovation, with dual scions providing redundancy and the option to select the stronger for permanent training.
A graft role that inheres in a grafted combination assembled using the cleft grafting technique, where a wedge-shaped scion base is inserted into a split (cleft) cut in the rootstock, creating cambial contact along the angled wedge surfaces. Cleft grafting is a traditional technique particularly suited for field grafting onto established rootstocks or for top-working mature vines, where the rootstock diameter typically exceeds scion diameter—the split rootstock accommodating one or two wedge-shaped scions inserted at the cleft margins where cambial alignment is achieved. The technique involves making a vertical split (4-6 cm deep) in the rootstock, preparing the scion with a long wedge taper (matching the cleft depth), and inserting the scion so that its cambium aligns with the rootstock cambium along at least one side of the wedge. Cleft grafts are typically sealed with grafting wax or wound sealant to prevent desiccation, with the mechanical pressure of the split rootstock holding the scion in position during callus formation. The technique is less common in modern benchgraft production (where omega and whip-and-tongue dominate) but remains important for vineyard renovation, top-working, and field grafting where rootstock diameter substantially exceeds available scion material.
Cleft_Graft_Role captures a traditional technique optimized for situations where rootstock diameter exceeds scion diameter, making it complementary to omega and whip grafting (which require matched diameters) and chip/T-budding (which work best on smaller rootstock stems). The technique's persistence in vineyard top-working reflects this niche—established vine trunks (3-10 cm diameter) far exceed typical scion diameters (8-12 mm), with the cleft accommodating this mismatch by aligning cambium along the wedge margins rather than across the entire cut surface. Cleft graft success depends on precise wedge preparation (smooth cuts, appropriate taper), accurate cambial alignment (critical on at least one side), and effective sealing (preventing desiccation of the exposed surfaces). The technique produces strong unions once established but has lower initial success rates than omega or whip grafting due to the limited cambial contact and greater tissue damage inherent in the cleft cut.
Geißfuß
Keilpfropfung
Spaltpfropfung
V-graft
cleft graft
split graft
wedge graft
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 435-440 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Pongrácz DP (1983) Rootstocks for Grape-vines, David Philip Publishers Dry N (2007) Grapevine rootstocks, Lythrum Press
Cleft Graft Role
The climat name Clos de Bèze designates a 15.4-hectare Grand Cru parcel in Gevrey-Chambertin documented since 630 CE when it was donated to the Abbey of Bèze by Duke Amalgaire of Burgundy — making it one of the oldest continuously cultivated named vineyard sites in the world. The adjacent climat Chambertin (12.9 hectares) was historically considered its equal, leading to the authorized designation Chambertin-Clos de Bèze for wines from Clos de Bèze (but not the reverse — Chambertin wines may not use the Clos de Bèze name), illustrating the extraordinary parcel-level precision and historical depth embedded in the Burgundian climat naming system across approximately 1,250 officially delimited parcels.
A vineyard designation within the Burgundian viticultural tradition that identifies a wine as originating from a named climat — a precisely delimited vineyard parcel defined by a unique combination of soil, subsoil, microclimate, slope orientation, and elevation that has been recognized over centuries of viticultural observation as producing wines with consistent and distinctive sensory characteristics, officially inscribed on the UNESCO World Heritage List in 2015 as the Climats du vignoble de Bourgogne.
Climat_Name is the quintessential expression of terroir-based vineyard classification, representing over a millennium of accumulated viticultural observation in the Côte d'Or and related Burgundian vineyard areas. The climat concept embodies the proposition that a specific, bounded piece of land produces wine with consistent and identifiable character that distinguishes it from adjacent parcels — a proposition tested and refined through centuries of monastic and secular viticulture and codified in successive classification systems from Lavalle (1855) through the AOC delimitations (1936 onward): the 2015 UNESCO inscription recognized the climats as an outstanding example of a cultural landscape demonstrating the interaction between human viticultural practices and specific geographic conditions over an extended historical period. Each climat is defined by precisely surveyed boundaries recorded in both the INAO delimitation parcellaire and the national land cadaster, with the climat name constituting a legal designation whose use on wine labels is regulated by the applicable AOC cahier des charges.
Burgund-Lagenparzelle
Burgunder Klimaname
Burgundy vineyard parcel name
Climat-Bezeichnung
climat designation
named Burgundy climat
UNESCO (2015) The Climats, terroirs of Burgundy, World Heritage List No. 1425 Pitiot S and Servant JC (2010) Les Vins de Bourgogne 14th Ed, Collection Pierre Poupon INAO (2023) Délimitation parcellaire des appellations de Bourgogne Lavalle J (1855) Histoire et statistique de la vigne et des grands vins de la Côte d'Or
Climat Name
The climate classification for the Barossa Valley (South Australia, 34.5°S) records multiple classification results — Köppen-Geiger: Csb (warm-summer Mediterranean), transitioning toward Csa (hot-summer Mediterranean) in the warmest valley floor sites Winkler: Region III (1,670-1,940 GDD base 10°C, warm climate) Huglin index: HI+2 (warm temperate viticultural climate, 2,100-2,400 units) Smart-Dry aridity index: moderately dry (annual water deficit 200-350 mm requiring supplemental irrigation) MCC (Tonietto and Carbonneau 2004): warm, moderately dry, with warm nights (class IH+2, IS-1, IF-1). This convergent multi-system classification identifies the Barossa as a warm, Mediterranean-climate viticultural region suited to heat-tolerant, late-ripening cultivars (Shiraz, Grenache, Mourvèdre, Cabernet Sauvignon) but marginal for cool-climate cultivars (Riesling surviving only at the higher-elevation Eden Valley sites above 400 m, and Pinot Noir commercially unviable in the Barossa floor).
A macroclimate description component that records the formal climate type assigned to the viticultural region according to one or more standardized climate classification systems — both general-purpose systems (Köppen-Geiger, Thornthwaite) and viticulture-specific systems (Winkler regions, Huglin index classes, Multicriteria Climatic Classification MCC) — providing the categorical climate identity that enables systematic comparison of viticultural regions worldwide and the prediction of cultivar suitability, wine style potential, and viticultural management requirements from the climate category.
Climate_Classification provides the categorical framework that reduces the continuous multidimensional variation of climate parameters into discrete classes that communicate viticultural potential and enable regional comparison. The multiple classification systems used in viticulture differ in their input variables, calculation methods, and class boundaries, and no single system captures all aspects of the climate-vine interaction — the Winkler system (cumulative GDD above 10°C, April-October in the northern hemisphere) measures heat accumulation and predicts cultivar maturity potential but ignores rainfall, humidity, and diurnal temperature range the Huglin heliothermal index (cumulative adjusted mean temperature × daylength coefficient) improves on Winkler by incorporating the photoperiod effect on photosynthetic potential but still ignores moisture the Smart-Dry systems add aridity and humidity indices to address the water balance dimension: the Multicriteria Climatic Classification (MCC) of Tonietto and Carbonneau (2004) integrates thermal, hydric, and nocturnal thermal amplitude dimensions into a three-axis classification that more completely characterizes the viticultural climate space, identifying 36 possible climate types from the combination of six heliothermal classes (very cool to very warm), four hydric classes (very humid to very dry), and three nocturnal temperature classes (cool nights, temperate nights, warm nights). The VVO documents classification results from multiple systems to provide complementary perspectives, recognizing that each system illuminates different aspects of the climate-viticulture relationship and that regional comparison is most reliable when using the same classification system across regions.
Klimaklassifikation
Klimatyp-Zuordnung
classification system result
climate category record
climate type designation
klimatische Einordnung
Winkler AJ et al. (1974) General Viticulture, University of California Press Huglin P (1978) Nouveau mode d'évaluation des possibilités héliothermiques, Comptes Rendus de l'Académie d'Agriculture de France 64: 1117-1126 Tonietto J and Carbonneau A (2004) A multicriteria climatic classification system for grape-growing regions worldwide, Agricultural and Forest Meteorology 124: 81-97 Jones GV et al. (2005) Climate change and global wine quality, Climatic Change 73: 319-343 Peel MC et al. (2007) Updated world map of the Köppen-Geiger climate classification, Hydrology and Earth System Sciences 11: 1633-1644
Climate Classification
The climate data for the Rheingau wine region (Germany, 50.0°N) documents the multilayered climatic characterization — macroclimate classified as Cfb (Köppen-Geiger temperate oceanic) with mean annual temperature 10.2°C, growing season (April-October) mean 15.3°C, and 550 mm growing season precipitation mesoclimate of the south-facing Rhine gorge slopes showing significant modification from the macroclimate, with growing season mean temperature elevated to 16.0-16.5°C by slope-enhanced radiation and the thermal buffering of the Rhine river (reducing frost frequency by 40-60% compared to inland sites at the same elevation), and growing season precipitation reduced to 450-500 mm in the rain shadow of the Taunus hills to the north microclimate within the fruit zone of VSP-trained Riesling canopy showing further modification, with midday cluster-zone temperatures 3-5°C above ambient air temperature due to radiation trapping within the canopy, and relative humidity 10-20% higher than ambient due to transpiration from the surrounding leaf mass.
A terroir information component that documents the atmospheric conditions — temperature, precipitation, radiation, humidity, and wind — that characterize the vineyard's growing environment at multiple spatial scales, from the macroclimate (regional climate classification), through the mesoclimate (site-specific climate modified by topography, water bodies, and land use), to the microclimate (the atmospheric conditions within and immediately surrounding the vine canopy), providing the climatic framework that determines which cultivars can ripen successfully at a given site, the phenological timing and duration of the growing season, the disease pressure regime, and the vintage-to-vintage variation in fruit composition and wine quality.
Climate_Data is the root class for the atmospheric component of terroir characterization — the domain that has undergone the most profound reassessment in recent decades due to climate change, which is shifting the thermal boundaries of viticultural zones at approximately 10-17 km per decade poleward and 10-15 m per decade in elevation, fundamentally altering the cultivar-climate matching that has defined regional wine identities for centuries. The hierarchical organization of climate data at macro-, meso-, and micro-scales reflects the nested spatial structure of atmospheric conditions in viticultural landscapes — the macroclimate determines the broad viticultural potential of a region (which cultivar maturity groups can ripen, whether the growing season is sufficiently long and warm for viticulture at all), the mesoclimate determines the site-specific suitability within the region (which slope aspects, elevations, and landscape positions optimize the macroclimate for viticulture), and the microclimate determines the immediate environment experienced by the vine and its fruit (how canopy management modifies the atmospheric conditions within the fruit zone): viticultural management operates primarily at the microclimate level (through canopy architecture decisions that control fruit zone light, temperature, and humidity), while site selection operates at the mesoclimate level, and regional cultivar choice operates at the macroclimate level. The VVO documents climate at all three scales, recognizing that the complete climatic characterization of a terroir requires specification of the regional climate context, the site-specific topographic modification, and the canopy-level management-dependent modification that together define the atmospheric environment in which the vine grows and fruit ripens.
Klimadaten
Weinberg-Klimacharakterisierung
atmospheric environment documentation
vineyard climate characterization
viticultural climatology record
vitikultur-klimatologische Dokumentation
Jones GV et al. (2005) Climate change and global wine quality, Climatic Change 73: 319-343 Gladstones J (1992) Viticulture and Environment, Winetitles Smart RE and Robinson M (1991) Sunlight into Wine, Winetitles van Leeuwen C and Seguin G (2006) The concept of terroir in viticulture, Journal of Wine Research 17: 1-10
Climate Data
The climate zone for the Mosel wine region (Germany, 49.5-50.5°N) designates it as a cool viticultural zone — growing season mean temperature 14.5-15.5°C (measured at the Bernkastel-Kues weather station, 120 m elevation), placing the Mosel at the lower boundary of commercially viable viticulture where only early-to-medium-ripening cultivars achieve consistent maturity. The cool zone designation explains the cultivar spectrum — Riesling (occupying 61% of the planted area) succeeds because its long, slow ripening period at low temperatures preserves the high acidity and develops the complex terpene and norisoprenoid aroma profile that defines Mosel Riesling Müller-Thurgau (earlier-ripening, less demanding of heat) occupies a further 13% and Spätburgunder (Pinot Noir, the latest-ripening cultivar commercially grown in the region) succeeds only on the warmest south-facing slopes. Late-ripening cultivars such as Cabernet Sauvignon (requiring >17°C growing season mean) cannot achieve maturity in the Mosel's cool zone, though climate change projections suggest that the region may shift from cool to intermediate zone classification by 2040-2060.
A macroclimate description component that records the position of the viticultural region within the global hierarchy of viticultural climate zones — from cool (growing season mean temperature 13-15°C, suitable for early-ripening cultivars such as Müller-Thurgau, Pinot Noir, and Chardonnay) through intermediate (15-17°C, suitable for medium-ripening cultivars such as Riesling, Sauvignon Blanc, and Merlot), warm (17-19°C, suitable for late-ripening cultivars such as Cabernet Sauvignon, Syrah, and Grenache), to hot (>19°C, suitable for very late-ripening cultivars and fortified wine production) — providing the fundamental thermal categorization that determines the spectrum of cultivars achievable at the site and the characteristic wine styles producible from the regional climate.
Climate_Zone provides the broadest categorical framework for matching cultivar thermal requirements to regional growing conditions — the zone classification directly predicts which cultivars from the global repertoire can achieve physiological maturity (sufficient sugar accumulation, acid reduction, flavor development, and phenolic ripening) within the available growing season, making climate zone the primary filter in cultivar selection decisions for new vineyard plantings. The thermal boundaries between zones are not sharp thresholds but gradients along which cultivar success probability changes continuously — the 15°C growing season mean boundary between cool and intermediate zones, for example, represents the approximate temperature at which medium-ripening cultivars shift from marginal (high vintage-to-vintage variability, frequent failure to achieve target ripeness) to reliable (consistent maturity in most vintages) performance: the same cultivar may succeed in one vintage and fail in the next at the zone boundary, while performing reliably in the zone interior. The climate zone concept is increasingly dynamic due to climate change — Jones et al. (2005) documented that many traditional European wine regions have shifted one full zone warmer (cool to intermediate, or intermediate to warm) over the 1950-2000 period, with consequences for cultivar suitability that are already visible in the northward expansion of red wine production, the commercial viability of viticulture in England and southern Scandinavia, and the heat stress challenges emerging in traditionally warm-zone regions (southern Spain, southern Australia, inland California) where extreme heat events during ripening now threaten the quality paradigm that these regions built on reliable warm-but-not-hot growing seasons.
Anbauklima-Einstufung
Klimazone
growing season temperature class
heat zone designation
viticultural thermal zone
vitikulturde Wärmezone
Jones GV et al. (2005) Climate change and global wine quality, Climatic Change 73: 319-343 Gladstones J (1992) Viticulture and Environment, Winetitles Winkler AJ et al. (1974) General Viticulture, University of California Press Jones GV (2006) Climate and terroir: impacts of climate variability and change on wine, in Macqueen RW and Meinert LD (eds) Fine Wine and Terroir — The Geoscience Perspective, Geological Association of Canada
Climate Zone
A climbing functional tendril at node 10 on an actively extending shoot of Vitis vinifera cv. Tempranillo in Rioja, Spain, exhibiting circumnutation movements sweeping through approximately 160-degree arcs over 90-minute periods, achieving 18 cm extension length and positioned 8-12 cm from a vertical trellis wire, showing high probability of contact within 24-48 hours based on sweep amplitude and wire proximity, with two previously attached tendrils at nodes 6 and 8 providing stable support for the basal shoot section while this distal tendril seeks new attachment to enable continued vertical growth, measured at BBCH stage 73 (berries beginning to touch) in early July when shoot apex is extending at 4-5 cm per day and requiring progressive attachment point addition to maintain vertical trajectory and prevent shoot bending or collapse.
A functional tendril actively engaged in the climbing process in Vitis vinifera, characterized by either active searching behavior involving circumnutation movements to increase contact probability with potential support structures, or recent successful attachment contributing to vertical or directional shoot growth toward improved light conditions, representing the subset of tendrils directly facilitating the vine's lianescent growth strategy of using external supports to achieve canopy elevation and light capture without investing resources in self-supporting trunk architecture. Climbing functional tendrils include both pre-attachment tendrils executing exploratory movements and recently attached tendrils (typically within 2-4 weeks of attachment) that are actively pulling shoots toward supports and enabling vertical progression. These tendrils represent the active climbing apparatus distinguishing Vitis from self-supporting growth forms, enabling the characteristic rapid vertical extension growth of 1-3 meters per season that allows wild grapevines to climb into forest canopies and cultivated vines to fill trellis systems efficiently. The climbing function involves coordinated behaviors across multiple tendrils at different developmental stages, with proximal tendrils providing established support while distal tendrils extend and search for new attachment points, creating a progressive climbing system that advances the shoot apex vertically while maintaining stable support for developed shoot sections and fruit clusters.
Climbing functional tendrils are distinguished from non-climbing tendrils (established attachments no longer contributing to vertical progression, senescent tendrils, or failed tendrils) by their active role in current climbing behavior either through searching movements or recent attachment enabling shoot elevation. The climbing function is temporally bounded, with tendrils transitioning from pre-climbing (immature) to climbing (active searching or recent attachment) to post-climbing (established support) states over their developmental trajectory. Climbing efficiency depends on tendril density along shoots, with adequate tendril production (typically every 2-3 nodes in functional positions) required to maintain climbing progression. Training system design influences climbing tendril effectiveness, with support structures positioned to intercept searching tendrils optimizing attachment success and vertical growth rates.
active climbing tendril
attachment-seeking tendril
kletternde Ranke
searching tendril
suchende Ranke
vertical growth tendril
vertikalwachstum Kletterorgan
Based on Darwin (1875) "The Movements and Habits of Climbing Plants" John Murray, London Jaffe & Galston (1968) "The physiology of tendrils" Annual Review of Plant Physiology 19=417-434 Goriely & Tabor (2013) "Spontaneous helix hand reversal and tendril perversion in climbing plants" Physical Review Letters 80=1564-1567 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 237-258.
Climbing Functional Tendril
Pinot Noir clone 115 (Dijon) bears a Clonal_Selection_Clone_Role as an ENTAV-INRA selection, an individual vine identified in a Burgundy vineyard in the 1970s, evaluated through replicated trials across multiple French regions over 15 years, registered for its small-cluster character and concentrated wine quality, and propagated as a uniform population. Chardonnay clone 96 serves in a Clonal_Selection_Clone_Role as a UC Davis Foundation Plant Services selection, formally evaluated through California trials before release, with documented performance characteristics enabling growers to predict behavior in their vineyards. Sangiovese clone R10 bears a Clonal_Selection_Clone_Role from the Italian VCR program, selected at Rauscedo and evaluated through Tuscan trials before national registration.
A selection status clone role that inheres in a grapevine clone derived through formal clonal selection, where individual mother vines are identified, isolated, evaluated through replicated trials, and propagated as genetically uniform populations. Clonal selection isolates single genotypes from within-variety populations, evaluating them through systematic field trials (typically 5-15 years) assessing yield, vigor, phenology, disease resistance, and fruit composition, followed by winemaking trials evaluating wine quality and typicity. Selected genotypes demonstrating superior and consistent performance become registered clones, each identified by a unique code and propagated as a uniform population tracing to the single original mother vine. Clonal selection provides documented, predictable performance characteristics—growers selecting a registered clone can anticipate specific yield levels, ripening timing, cluster architecture, and wine style based on trial data. The approach emerged from scientific viticulture in the early-to-mid 20th century, with programs at Geisenheim (Germany, from 1926), Conegliano (Italy, from 1930s), and Montpellier (France, ENTAV from 1962) establishing systematic selection methodologies subsequently adopted worldwide.
Clonal_Selection_Clone_Role represents the systematic scientific approach to grapevine improvement that has produced the registered clones dominating contemporary vineyard plantings. The methodology's strength lies in documented, reproducible performance—trial data enables informed clone selection matching specific site conditions and production objectives. Potential limitations include genetic erosion (widespread planting of few clones reduces diversity), reduced complexity (some argue clonal uniformity simplifies wines), and fixation of virus infections present in original selections (addressed through sanitary selection integration). The proliferation of registered clones for major varieties (50+ for Pinot Noir, 40+ for Chardonnay) reflects both genuine genetic diversity and institutional interests debate continues about the appropriate number of clones and the optimal balance between clonal uniformity and genetic diversity.
Individualselektion
Klonenauslese
clonal selection
formal selection
individual vine selection
klonale Selektion
systematic selection
systematische Selektion
Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 Walter B, Martelli GP (1996) Clonal and sanitary selection of the grapevine, INRA Editions ENTAV-INRA (2007) Catalogue des Variétés et Clones de Vigne Regner F (2007) Genetic variation in grapevine clones, Blackwell pp.: 1-34
Clonal Selection Clone Role
The clonal selection record for Pinot Noir clone 115 (Dijon series) documents its selection from a single vine in Domaine Ponsot's Clos de la Roche vineyard by Raymond Bernard (ENTAV-INRA) in 1971, characterizing it as a small-cluster, millerandage-prone clone yielding 20-30% less than clone 114 but producing wines of exceptional concentration and aromatic complexity, with full ENTAV certification achieved in 1978 and subsequent global distribution through licensed nurseries. The clonal selection record for Grüner Veltliner clone 17 (Klosterneuburg) traces its origin to a mass selection at Weingut Bründlmayer in Langenlois, documenting virus-free status confirmed through ELISA and PCR testing, moderate vigor, and consistent production of structured wines with characteristic white pepper notes.
Clonal selection record is a grapevine information entity that documents the complete history, methodology, performance evaluation, certification status, and distribution of individually selected and propagated clonal lines of Vitis vinifera cultivars, serving as the primary information resource for tracing the origin, identity, and viticultural characteristics of certified plant material available to grape growers and nurseries worldwide.
Clonal selection records constitute the documentary backbone of grapevine plant material identity and quality assurance, bridging the gap between historical empirical selection and modern molecularly verified certification systems. The information entity encompasses the entire lifecycle of a clone from initial field identification through performance evaluation, virus testing, certification, and commercial distribution. National clonal selection programs differ substantially in their methodologies and documentation standards — France's ENTAV system, Germany's Geilweilerhof program, Italy's CRA-VIT selections, and Australia's CSIRO/SARDI program each maintain distinct record structures reflecting national regulatory frameworks. The economic significance of clonal selection records is substantial, as a single clone's characteristics can influence vineyard establishment decisions worth millions of euros over the 25-40 year productive life of a planting. Modern clonal records increasingly incorporate molecular characterization (SSR profiles, SNP genotyping, viral load assessment) alongside traditional ampelographic and oenological evaluation data.
Klonakte
Klondossier
Klonselektionsdokumentation
Klonselektionsnachweis
Klonselektionsprotokoll
clonal line record
clonal selection documentation
clone dossier
clone record
clone selection file
Mannini F (2004) Clonal selection in grapevines: interactions between genetic and sanitary strategies to improve propagation material. Acta Horticulturae 652: 21-27 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties. Heredity 104(4): 331-340 This P et al. (2006) Historical origins and genetic diversity of wine grapes. Trends in Genetics 22(9): 511-519
Clonal Selection Record
Pinot Noir clone 115 (Dijon clone, ENTAV-INRA selection) bears a Clone_Role as an officially registered French clone selected for small berries, loose clusters, and concentrated wine character—distinguished from clone 777 (also Dijon) selected for aromatic intensity and early ripening. Chardonnay clone 96 (FPS selection) serves in a Clone_Role as the most widely planted Chardonnay clone in California, selected for productivity, disease tolerance, and balanced wine character. Riesling clone 239 bears a Clone_Role as a German (Geisenheim) selection prized for aromatic intensity and site expressivity, contrasting with clone 21B selected for higher yield and neutral character.
A grapevine role that inheres in a plant or population of Vitis vinifera or Vitis hybrid by virtue of its status as a clone—a genetically uniform population derived by vegetative propagation from a single original mother vine, maintaining genetic identity across all individuals within the clonal lineage. Clone roles encompass both official recognition status (registered, certified, standard designations within regulatory frameworks) and selection methodology status (mass selection, clonal selection, sanitary selection, genetic selection approaches by which clones are identified and developed). Clonal identity in grapevines reflects the exclusively vegetative propagation of commercial cultivars, where each named clone traces to a specific original vine selected for desirable characteristics, with all derived plants theoretically genetically identical (though somatic mutations accumulate over propagation generations). Clone designation systems vary by country—France uses numeric codes with institutional prefix (ENTAV-INRA clones), Germany uses alphanumeric designations (Gm, Gf, Fr prefixes), Italy uses regional codes (VCR, ERSA)—but all serve to identify specific selections within cultivars that differ in agronomic, enological, or sanitary characteristics.
Clone_Role is the parent class for the complex classification systems distinguishing clones by official recognition (regulatory status within certification frameworks) and by selection methodology (how the clone was identified and developed). The biological reality of clonal identity is complicated by ongoing somatic mutation—over centuries of vegetative propagation, mutations accumulate creating genetic variation within nominally "clonal" populations—leading to debates about clone boundaries and the relationship between ancient clones and modern selections. Clone choice significantly impacts vineyard performance and wine character studies demonstrate yield differences of 20-50% and wine quality differences readily detected in blind tasting between clones of the same variety grown in identical conditions. The proliferation of clone designations (hundreds of registered clones for major varieties like Pinot Noir) reflects both genuine genetic diversity and the institutional interests of selection programs in registering distinct selections.
Klon
Klonauslese
Rebklon
clone
grapevine clone
vine clone
Mannini F (2000) Clonal selection in grapevine: interactions between genetic and sanitary strategies, ISHS Acta Horticulturae 528: 17-25 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 ENTAV-INRA (2007) Catalogue des Variétés et Clones de Vigne
Clone Role
A closed petiolar sinus on a mature leaf at node 6 of Vitis vinifera cv. Merlot in Pomerol, France, exhibiting characteristic nearly closed configuration with basal lobe margins separated by only 0.3 cm creating narrow slit-like opening, sinus depth of 1.8 cm creating moderate indentation, rounded U-shaped contour with smooth sinus base, basal lobe margins converging at approximately 10-degree angle nearly meeting at petiole attachment, lobe margin edges smooth without teeth in sinus region, moderate to dense pubescence on lobe surfaces particularly near sinus base, almost completely obscuring the petiole at its attachment point requiring careful leaf manipulation to observe petiole insertion, serving as primary diagnostic feature enabling clear distinction from Cabernet Sauvignon (open U-shaped sinus) and Cabernet Franc (open V-shaped sinus) in field identification, measured at BBCH stage 75 (berries pea-sized) in late June on well-developed exterior leaf representing reliable diagnostic expression, demonstrating the high identification value of closed petiolar sinus in this major cultivar.
A petiolar sinus exhibiting narrow separation or complete closure with basal lobe margins meeting or nearly meeting at the petiole-lamina junction in Vitis vinifera, characterized by minimal or no visible gap between lobe margins creating narrow slit-like opening or complete closure, lobe margins converging to meet or approach within 0-5 mm at the petiole attachment point, typically U-shaped configuration with lobes curving inward rather than diverging, obscured or completely hidden petiole at the attachment zone, occurrence in several important cultivars including Merlot (nearly closed to closed), Syrah (closed to slightly open), and Grenache (closed), high diagnostic value distinguishing these closed-sinus cultivars from open-sinus varieties like Cabernet Sauvignon or Sauvignon Blanc, and potential functional implications including more compact leaf base architecture and potentially increased mechanical support at the petiole-lamina junction though experimental evidence is limited. Closed petiolar sinuses develop when basal lobe primordia grow inward during leaf development with genetic programming directing convergent lobe trajectory rather than divergent growth, creating the characteristic narrow or absent gap that persists reliably across environmental conditions making closed sinus a highly stable diagnostic character.
Closed petiolar sinuses represent one of three primary sinus categories, distinguished from open sinuses by minimal or no gap between lobe margins and from overlapping sinuses by margins meeting rather than crossing. The closed configuration shows variation in completeness, with some cultivars showing complete closure (margins touching) while others show nearly closed configuration (margins separated by 1-5 mm), creating useful subcategories for finer discrimination. The distinction between closed and overlapping can be subtle in borderline cases, with classification sometimes depending on whether margins merely meet (closed) or actually cross creating occlusion (overlapping). Genetic control is strong with closed versus open showing relatively simple inheritance, though the distinction between closed and overlapping appears more complex suggesting multiple genes or modifying factors. Environmental stability is generally high, with closed configuration maintained across most conditions, though extreme vigor may produce slightly more open sinuses in some leaves requiring assessment of multiple leaves for confident classification. Functional implications remain largely speculative, with proposed advantages including more compact architecture or enhanced mechanical support at the petiole junction lacking strong experimental support. Ampelographic utility is very high, with closed petiolar sinus serving as definitive diagnostic feature distinguishing Merlot, Syrah, and other closed-sinus cultivars from related varieties.
closed basal sinus
convergent lobe sinus
enge Stielbucht
geschlossener Petiolarsinus
konvergente Basalbucht
narrow petiolar sinus
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 136-182 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 79-102.
Closed Petiolar Sinus
Clove aroma assessment comparing Rioja Reserva aged in American oak (clove intensity 6.8 on 10-point scale, eugenol 42 μg/L) versus French oak (clove 4.5, eugenol 28 μg/L) reveals the oak species effect on spice expression, with American oak's higher eugenol content and more aggressive extraction creating the pronounced clove character traditional in Rioja, while modern French oak-aged examples show more restrained spice allowing greater fruit expression, demonstrating how clove intensity indicates both oak type and regional style evolution.
Clove aroma quality is a spicy aroma quality that represents olfactory perception evoking Syzygium aromaticum (clove) bud-associated scents, characterizing the distinctive warm, sweet-spicy aromatic impression primarily derived from oak barrel aging that serves as a marker for new oak usage and toast level in wine production. Within the BFO framework, this quality specifically addresses the clove subcategory of spicy aroma, distinguished from pepper by its oak-derived rather than grape-derived origin and its role in indicating winemaking treatment rather than varietal character. The ontological significance lies in clove aroma serving as one of the primary sensory indicators of oak aging, where clove intensity and character reveal information about barrel newness, toast level, and the extent to which oak extraction has occurred, making clove assessment essential for evaluating oak integration and appropriateness in barrel-aged wines. The compositional basis for clove aroma centers on eugenol (4-allyl-2-methoxyphenol), a phenolic compound extracted from oak wood that provides clove-like aroma with detection threshold of approximately 6 μg/L in wine, with concentrations in oak-aged wines typically ranging from 5 to 50 μg/L depending on barrel age, toast level, and aging duration. Additional contributors include isoeugenol (providing nutmeg alongside clove character) and acetylated eugenol derivatives formed during toasting. The oak treatment factors affecting clove character include toast level (medium-plus toast maximizing eugenol extraction), barrel age (new oak providing highest eugenol, declining with reuse), and aging duration (longer contact increasing extraction until equilibrium). The interaction between clove and fruit in wine evaluation follows the principle that well-integrated clove complements and enhances fruit complexity while excessive or dominant clove indicates over-oaking that masks varietal character and creates unbalanced wine. Understanding clove aroma quality enables oak treatment assessment based on eugenol-derived spice expression, integration evaluation judging the balance between oak spice and fruit, and winemaking style characterization distinguishing heavily oaked from restrained or unoaked approaches.
Clove_Aroma_Quality addresses the specifically clove (eugenol-derived) subcategory of spicy aroma, representing an oak-derived rather than grape-derived spice character. This quality is essential for oak treatment evaluation and distinguishes from grape-derived pepper character. The perception of clove as positive or negative depends on integration with fruit and appropriateness for wine style, with excessive clove indicating potential over-oaking.
Eugenol-Charakter
Gewürznelkennote
Nelkenaroma
Nelkenduft
clove bud notes
clove spice aroma
eugenol character
sweet spice scent
Chatonnet P et al. (1999) Volatile and odourous compounds in barrel-aged wines: impact of cooperage techniques on the aromatic profile. American Journal of Enology and Viticulture 50(4): 470-476 Pollnitz AP et al. (1999) Determination of oak lactones in barrel-aged wines and in oak extracts by stable isotope dilution analysis. Journal of Chromatography A 857(1-2): 239-246
Clove Aroma Quality
Cluster accessibility quality comparison in Marlborough evaluated Sauvignon Blanc harvest efficiency under three training system configurations with equivalent canopy management intensity. Vertical shoot positioning with bilateral cordon demonstrated excellent cluster accessibility quality with 95% of clusters clearly visible at consistent 0.6-0.9 m height within the defined fruit zone, minimal foliage obscuration following standard basal leaf removal, and adequate spacing between clusters permitting individual access without interference, achieving harvest productivity of 195 kg per hour. Scott Henry divided canopy showed good cluster accessibility quality on the upper curtain comparable to VSP but reduced accessibility on the lower curtain where pendant clusters required upward reaching and competed with downward-hanging foliage, averaging 170 kg per hour combined. Sprawling single wire training exhibited poor cluster accessibility quality with clusters distributed unpredictably throughout the canopy volume from 0.3 to 1.4 m height, substantial foliage obscuration requiring leaf displacement during picking, and frequent cluster interference within dense zones, limiting productivity to 125 kg per hour and demonstrating how training system design critically influences cluster accessibility quality outcomes.
Cluster accessibility quality is a harvest logistics quality that inheres in a grapevine cultivar or clone within a specific training system configuration and represents the ease with which clusters can be visually located, physically reached, and operationally accessed for removal during harvest, determined by the consistency of cluster positioning relative to the canopy architecture, the degree of cluster obscuration by foliage, the clearance available for hand or tool access to the peduncle, and the spatial separation between clusters permitting individual removal without interference from adjacent fruit. This quality directly influences harvest efficiency by determining the search and access time required per cluster beyond the actual removal operation, with highly accessible cultivar-training combinations enabling rapid, systematic picking while poor accessibility creates time-consuming cluster location, awkward reaching, and interference between adjacent clusters that slows operations and increases damage risk. The ontological significance of cluster accessibility quality lies in its role as a primary determinant of harvest labor productivity independent of cluster removal difficulty, recognizing that pickers must locate and access clusters before removal can occur and that these preliminary operations may consume substantial time when accessibility is poor, particularly in dense canopies or training systems that position fruit in obscured or awkward locations. The biological basis of cluster accessibility quality encompasses the cultivar characteristics determining cluster positioning including peduncle length affecting cluster projection from the cordon or cane, the uniformity of cluster positioning creating predictable locations that facilitate systematic harvest, and the relationship between cluster development timing and canopy density that influences whether clusters emerge clearly from surrounding foliage or become buried within dense leaf growth. Training system factors strongly modulate accessibility through their influence on canopy architecture and fruit zone configuration, with well-designed systems positioning clusters at consistent, accessible heights within open fruit zones while poor training-cultivar matches may create variable positioning, excessive shading, or mechanical interference that compromises accessibility regardless of inherent cultivar characteristics. Assessment of cluster accessibility quality involves observation of cluster visibility and positioning uniformity, measurement of time allocation between location, access, and removal phases of harvest, and comparison of productivity rates achieved under standardized conditions across cultivar-training combinations.
Cluster_Accessibility_Quality is_a Harvest_Logistics_Quality, representing the ease of locating and physically accessing clusters for harvest. This class addresses positioning, visibility, and clearance characteristics rather than removal difficulty per se. Strong training system interaction distinguishes this from purely cultivar-determined qualities. Percentage of clusters in defined harvest zones and productivity time allocation provide assessment metrics. Related to Fruit_Zone_Leaf_Layers_Quality through the influence of canopy density on cluster visibility, though accessibility addresses operational rather than microclimate dimensions.
Erntezugang-Leichtigkeit
Fruchtpositionierungsqualität
Traubensichtbarkeit
Traubenzugänglichkeit
cluster positioning quality
cluster visibility
fruit accessibility
harvest access ease
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 64-85 Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review. American Journal of Enology and Viticulture 60:251-268 Morris JR (2007) Development and commercialization of a complete vineyard mechanization system. HortTechnology 17:411-420
Cluster Accessibility Quality
Quantitative analysis of Cluster_Architecture_Based_Resistance_Disposition at INRAE Bordeaux employed 3D imaging to characterize cluster compactness across 120 cultivars and correlate architectural parameters with grey mold susceptibility. Cluster compactness was quantified using the CI-12 index (berries per cm rachis) and OIV descriptor 204. The compact cultivar 'Pinot Noir' (CI-12 = 8.2, OIV 204 rating 7-9) exhibited mean porosity of only 12% (air space within cluster volume) and berry contact index of 4.3 (mean contact points per berry), creating a persistent humid microenvironment where relative humidity exceeded 90% for 14 hours following a 10mm rainfall event. In contrast, the loose cultivar 'Gewürztraminer clone 643' (CI-12 = 4.1, OIV 204 rating 3) showed porosity of 38% and contact index of 1.8, with cluster humidity returning to ambient within 4 hours post-rainfall. Under natural Botrytis pressure, 'Pinot Noir' developed 52% cluster infection while 'Gewürztraminer clone 643' showed only 18% infection despite 'Gewürztraminer' being generally considered susceptible, demonstrating how cluster architecture can override other susceptibility factors.
A Cluster_Architecture_Based_Resistance_Disposition is a Botrytis_Resistance_Disposition that inheres in a Vitis vinifera cluster and is realized through morphological and structural characteristics that create unfavorable microclimatic conditions for Botrytis cinerea infection and spread, independent of direct physiological or biochemical defense mechanisms. This disposition encompasses the genetically determined cluster structural traits that influence humidity retention, air circulation, berry contact surface area, and wound susceptibility within the cluster microenvironment. Key architectural parameters include cluster compactness (density of berry packing), cluster length and shape (cylindrical, conical, winged), berry size relative to rachis dimensions, pedicel length and flexibility, and rachis branching patterns. Compact clusters with tightly packed berries create persistent high humidity microenvironments following rain or dew events, maintain berry-to-berry contact surfaces that facilitate pathogen spread, and are prone to berry cracking when expanding berries compress against neighbors. In contrast, loose clusters with well-spaced berries dry rapidly after wetting, minimize contact-mediated pathogen transfer, and reduce mechanical stress on berry skins. Cluster architecture is under strong genetic control with moderate to high heritability, enabling effective selection in breeding programs. These structural traits represent constitutive, passive resistance mechanisms that function independently of active defense responses, providing baseline protection that complements inducible biochemical defenses.
Cluster_Architecture_Based_Resistance_Disposition is_a Botrytis_Resistance_Disposition. This class encompasses structural traits that reduce disease through environmental modification rather than direct host-pathogen interaction. Child classes include Loose_Cluster_Disposition (low compactness) and Thick_Skin_Disposition (mechanical barrier). Cluster architecture is distinct from but may interact with Cuticle_Based_Resistance_Disposition and Phytoalexin_Based_Botrytis_Resistance_Disposition which represent physiological rather than structural mechanisms.
Strukturelle Botrytis-Resistenz
Traubenarchitektur-basierte Resistenz-Disposition
Traubenmorphologie-Resistenz
architectural Botrytis avoidance
cluster morphology resistance
structural bunch rot resistance
Vail ME, Marois JJ (1991) Grape cluster architecture and the susceptibility of berries to Botrytis cinerea, Phytopathology 81:188-191 Molitor D et al. (2012) Impact of grape cluster division on cluster morphology and bunch rot epidemic, American Journal of Enology and Viticulture 63:508-514 Tello J, Ibáñez J (2018) What do we know about grapevine bunch compactness? A state-of-the-art review, Australian Journal of Grape and Wine Research 24:6-23
Cluster Architecture Based Resistance Disposition
Cluster architecture varies dramatically across V. vinifera cultivars, from the extremely compact cylindrical clusters of Pinot Noir (compactness index 8-12 berries/cm rachis) to the loose conical clusters of Sauvignon Blanc (3-5 berries/cm). Architecture components include rachis length (8-25 cm), primary branch number (4-12), pedicel length (4-12 mm), and berry count (50-350 per cluster). The OIV descriptor 204 classifies cluster density on a 1-9 scale, with 1 being very loose and 9 being very compact. Cluster architecture heritability is moderate to high (h² = 0.55-0.75) for most component traits.
A reproductive disposition that is the inherited capacity of a Vitis vinifera plant to develop grape clusters (bunches) with characteristic three-dimensional structure determined by rachis branching pattern, pedicel length, berry number, berry size, and the spatial arrangement of berries within the cluster framework.
Cluster_Architecture_Disposition serves as the parent class for traits determining bunch physical structure. Architecture profoundly affects disease susceptibility (particularly Botrytis), harvest method suitability, and wine quality through skin-contact dynamics. The trait complex results from the interaction of inflorescence development genes with berry growth patterns. Selection for loose cluster architecture has become a major breeding objective for disease resistance improvement.
Gescheinmorphologie
Traubenarchitektur
Traubenstruktur
bunch architecture trait
bunch structure capacity
cluster morphology disposition
Tello J, Ibáñez J (2018) What do we know about grapevine bunch compactness? A state-of-the-art review. Australian Journal of Grape and Wine Research 24:6-23 Shavrukov YN et al. (2004) Inflorescence and bunch architecture development in Vitis vinifera L. Australian Journal of Grape and Wine Research 10:116-124
Cluster Architecture Disposition
The cluster compactness description of Sémillon documents compact clusters resulting from a combination of medium-high fruit set (40-50% of flowers setting berries) and medium-large berry size (1.8-2.5 g) on a moderately short, relatively rigid rachis — this volumetric compactness produces the tightly packed, heavy clusters (200-350 g) that, combined with Sémillon's thin berry skin, create the ideal conditions for Botrytis cinerea infection that in the specific mesoclimate of Sauternes and Barsac develops as noble rot (pourriture noble) rather than destructive grey rot, concentrating sugars and producing the distinctive botrytized flavor compounds of Sauternes wines. The compact Sémillon cluster architecture contrasts with the structurally compact Pinot Noir (where compactness results primarily from high berry number on very short rachis internodes rather than from large berry volume) and the loose clusters of Sauvignon Blanc (where moderate berry number and elongated rachis internodes produce well-separated berries).
A cluster description component that records the overall structural tightness of the mature cluster as an integrated assessment of berry density, rachis flexibility, and the spatial arrangement of berries on the rachis framework, distinguishing between clusters where compactness results from high berry number on a short rigid rachis (structural compactness) versus those where compactness results from large berry size filling the available rachis space (volumetric compactness), with compactness evaluation providing a more holistic assessment of cluster architecture than the berry-level density descriptor alone.
Cluster_Compactness_Description provides a higher-level characterization that integrates the berry-level density assessment (OIV 204) with the structural properties of the rachis framework. The distinction between structural compactness and volumetric compactness is practically important because the two types respond differently to viticultural interventions — structural compactness (high berry number on short, rigid rachis) can be alleviated by reducing fruit set through gibberellic acid application at bloom or by mechanical cluster tipping (removing the distal third of the cluster before berry set), while volumetric compactness (large berries filling available space) can be reduced by water deficit strategies that limit berry size or by selecting clones with smaller berries: these different management approaches require correct diagnosis of the compactness type, which in turn requires understanding of the cultivar-specific cluster architecture. Some breeding programs have specifically targeted cluster looseness as a selection criterion — the Julius Kühn-Institut and other European breeding stations evaluate new PIWI selections for cluster architecture alongside disease resistance, recognizing that loose clusters reduce the need for Botrytis management even when chemical resistance is not available, and that the combination of loose cluster architecture with disease resistance gene pyramids creates cultivars requiring minimal to zero fungicide input.
Gesamtpackungs-Bewertung
Trauben-Strukturdichte
Traubenkompaktheits-Beschreibung
bunch compactness assessment
cluster structural tightness
overall packing evaluation
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Tello J and Ibáñez J (2018) What do we know about grapevine bunch compactness? A state-of-the-art review, Australian Journal of Grape and Wine Research 24: 6-23 Molitor D et al. (2012) Viticulture and wine quality under a changing climate, Journal of Agricultural Science 150: 137-156
Cluster Compactness Description
Cluster compactness is quantified by multiple metrics including berries per centimeter of rachis (3-15 range), the CI-12 compactness index, and visual OIV 204 scoring. Pinot Noir exhibits high compactness (10-14 berries/cm, OIV 7-9) while Gewürztraminer shows moderate-high values (8-11 berries/cm, OIV 5-7). Compactness increases Botrytis incidence exponentially, with infection rates of 5-10% in loose clusters versus 30-60% in compact clusters under identical conditions. Berry deformation in highly compact clusters can reach 15-25% of berry diameter, affecting skin integrity.
A cluster architecture disposition that is the inherited capacity of a Vitis vinifera plant to develop clusters with a characteristic degree of berry packing density, determined by the relationship between berry number, berry size, rachis length, and rachis branching pattern, affecting the spatial proximity of adjacent berries.
Cluster_Compactness_Disposition serves as the parent class for specific density categories. Compactness results from the interaction of multiple underlying traits and cannot be modified by single-gene manipulation. Management practices including gibberellin application, early leaf removal, and bloom-time irrigation stress can reduce compactness by 20-40% in susceptible cultivars. Compactness assessment timing is critical, as post-veraison berry softening increases apparent density.
Beerenpacking
Kompaktheits-Disposition
Traubendichte
berry packing disposition
bunch density trait
cluster tightness
Tello J et al. (2015) A new image-based tool for the high throughput phenotyping of pollen viability: evaluation of inter- and intra-cultivar diversity in grapevine. Plant Methods 11:16 Molitor D et al. (2012) Timing of cluster-zone leaf removal and its impact on canopy morphology, cluster structure and bunch rot susceptibility of grapes. Journal International des Sciences de la Vigne et du Vin 46:149-159
Cluster Compactness Disposition
The cluster density description of Pinot Noir documents very dense clusters (OIV 204:9) in which berries are so tightly packed that mutual compression produces flattened contact surfaces on adjacent berries, with the inner berries receiving no air circulation and the rachis branches completely obscured by the continuous berry mass — this extreme density is the primary factor driving Pinot Noir's notorious susceptibility to Botrytis cinerea bunch rot, as compressed berry skins crack under expansion pressure during ripening (particularly following rainfall events), creating entry points for fungal infection that spreads rapidly through the tightly packed interior where humidity remains near 100%. The very dense packing of Pinot Noir contrasts with the loose clusters of Gewürztraminer (OIV 204:3, berries separated by visible rachis segments, minimal berry-to-berry contact) and the very loose clusters of some Muscat clones (OIV 204:1, berries hanging freely on individual pedicels with complete air circulation around each berry).
A cluster description component that records the degree of berry packing within the mature cluster according to OIV descriptor 204, scored on a scale from very loose (state 1, berries clearly separated with visible rachis branches between them) through loose, medium, dense, to very dense (state 9, berries strongly compressed against each other with deformed contact surfaces and no rachis visible), with cluster density reflecting the ratio of total berry volume to the spatial volume defined by the cluster outline and having critical consequences for Botrytis cinerea susceptibility, berry skin integrity, and spray penetration effectiveness.
Cluster_Density_Description documents a character of exceptional viticultural importance because cluster density is the single strongest predictor of Botrytis bunch rot susceptibility among the morphological cluster characters — dense clusters trap moisture, prevent fungicide spray penetration, create berry skin damage through compression, and maintain the high-humidity microenvironment within the cluster interior that favors Botrytis infection and sporulation. The OIV scoring system assesses density visually and tactilely at harvest maturity, using the mobility of individual berries within the cluster as the practical criterion — in loose clusters (state 1-3), individual berries can be moved freely without displacing adjacent berries in medium clusters (state 5), berries move but contact adjacent berries in dense clusters (state 7-9), berries cannot be moved without deforming or displacing the entire cluster structure: this functional definition captures the practical consequence of density more effectively than geometric measurements alone. Cluster density is determined by the interaction of inflorescence branching architecture (rachis branch length and spacing, genetically determined) with fruit set percentage and berry size (environmentally modulated) — the same inflorescence architecture produces loose clusters under poor fruit set conditions and dense clusters under high fruit set conditions, explaining the season-to-season variation in density that can span 2-4 OIV scale points for a single cultivar.
Beerenpacking-Bewertung
Traubendichte-Beschreibung
Traubenkompaktheit-Einstufung
berry packing assessment
bunch density record
cluster tightness characterization
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Vail ME and Marois JJ (1991) Grape cluster architecture and the susceptibility of berries to Botrytis cinerea, Phytopathology 81: 188-191 UPOV (2011) Guidelines for the Conduct of Tests for Distinctness, Uniformity and Stability — Vine (Vitis L.), TG/50/9
Cluster Density Description
The cluster description of Pinot Noir documents small, compact, cylindrical clusters (OIV 202:3, OIV 204:7, OIV 208:1) weighing 80-150 g at harvest, with a short semi-lignified peduncle (15-25 mm), no wings or a single small wing, and tightly packed berries exhibiting mutual compression at the contact zones — this compact, small cluster architecture is one of Pinot Noir's most recognizable cultivar features (the name Pinot itself derives from the French pin, pine cone, reflecting the cluster's resemblance to a pine cone) and has direct viticultural consequences including high susceptibility to Botrytis cinerea infection in the compressed berry interior where air circulation is minimal and berry skin integrity is compromised by mutual pressure.
An ampelographic description component that documents the morphological characteristics of the mature grape cluster (bunch) at harvest — the compound fruit structure developed from the inflorescence — recording size, shape, compactness, wing development, peduncle length and lignification, and the overall architectural configuration that determines the cultivar-specific cluster phenotype with consequences for yield, disease susceptibility, harvest method suitability, and wine or table grape quality potential.
Cluster_Description addresses the harvested fruit structure that is the ultimate product of viticultural management and the starting material for winemaking or fresh market delivery — cluster morphology is therefore the ampelographic domain where cultivar identification intersects most directly with quality assessment and commercial value determination. The cluster develops from the inflorescence over approximately 14-16 weeks from bloom to harvest, with the final cluster architecture determined by the interaction of the genetically predetermined inflorescence branching pattern with fruit set percentage (the proportion of flowers that develop into berries), berry size (determined by seed number and cultivar-specific cell division and expansion patterns), and rachis elongation or contraction during berry development: the same inflorescence structure can produce either a compact or a loose cluster depending on fruit set — high fruit set with large berries produces compact clusters prone to berry compression and disease, while low fruit set or small berries produces loose clusters with better air circulation and disease resistance. The OIV descriptor system characterizes clusters at the harvest-ripe stage (E-L stage 38) when the cluster has achieved its final size and configuration, using descriptors for overall size (OIV 202), density/compactness (OIV 204), number of wings (OIV 209), and shape (OIV 208) that together define the cluster phenotype.
Ernte-Trauben-Charakterisierung
Trauben-Beschreibung
Traubenmorphologie
bunch morphology description
grape cluster characterization
harvest cluster documentation
Cluster Description
Cluster exposure quality optimization in Stellenbosch Cabernet Sauvignon compared leaf removal intensities in a warm climate context where balancing exposure benefits against sunburn risk presented management challenges. No leaf removal treatment produced cluster exposure quality of only 6% ambient PAR with 78% of clusters classified as interior clusters on point quadrat assessment, resulting in methoxypyrazine concentration of 24 ng/L and anthocyanins of 1.0 mg/g. Moderate morning-side-only leaf removal improved cluster exposure quality to 14% ambient PAR and 35% interior clusters, reducing methoxypyrazines to 11 ng/L while increasing anthocyanins to 1.6 mg/g with negligible sunburn. Aggressive both-sides leaf removal achieved cluster exposure quality of 28% ambient PAR and only 12% interior clusters, further reducing methoxypyrazines to 6 ng/L but with anthocyanins declining to 1.3 mg/g and 15% berry sunburn damage, demonstrating the cluster exposure quality optimum at intermediate exposure levels for this hot climate site.
Cluster exposure quality is a canopy gaps quality that inheres in the fruit zone of a grapevine canopy system and represents the degree to which developing grape clusters receive direct or minimally attenuated solar radiation, air circulation, and spray coverage as determined by the positioning of clusters relative to canopy gaps, the leaf layer number immediately surrounding cluster positions, and the management interventions specifically targeting fruit zone openness through basal leaf removal and fruit zone shoot management. This quality directly addresses the environmental conditions experienced by the primary target tissue for quality development, recognizing that overall canopy gap characteristics may not accurately reflect the specific conditions at cluster positions if gaps and clusters are not spatially aligned within the canopy architecture. The ontological significance of cluster exposure quality lies in its role as the proximate determinant of berry developmental environment, wherein light exposure intensity and duration drive phenolic biosynthesis, influence organic acid metabolism, affect aromatic precursor development, and determine the visual characteristics of berry pigmentation that contribute to wine color and quality perception. The functional consequences of cluster exposure quality operate through multiple interconnected pathways including direct photomorphogenic signaling in berry tissues responding to light quality and quantity, temperature effects wherein exposed clusters experience higher daytime temperatures promoting metabolic acceleration but also potentially heat stress in hot climates, and disease-related effects wherein exposed clusters dry rapidly and receive better spray coverage than shaded clusters surrounded by dense foliage. The measurement of cluster exposure quality employs various approaches including the percentage interior clusters metric from point quadrat analysis indicating clusters with obstructing foliage, direct measurement of photosynthetically active radiation at cluster positions using quantum sensors, visual scoring of cluster visibility from standardized observation positions, and image analysis quantifying the proportion of cluster surface visible in photographs taken perpendicular to the canopy face. Optimal cluster exposure quality represents a cultivar-specific and climate-specific balance between sufficient exposure for quality development, typically requiring 10-20% ambient PAR at cluster surfaces, and protection from excessive exposure that risks sunburn damage, with the threshold for damage varying from approximately 35°C berry temperature in thin-skinned white cultivars to over 40°C in thick-skinned reds adapted to hot climate conditions.
Cluster_Exposure_Quality is_a Canopy_Gaps_Quality, representing the specific fruit-zone-focused dimension of canopy openness assessment. This class addresses the environmental conditions at cluster positions specifically rather than general canopy gap characteristics. Distinguished from Light_Penetration_Quality by the explicit focus on cluster-level assessment. The percentage interior clusters metric from point quadrat analysis provides standardized quantification. The optimal PAR target of 10-20% ambient with cultivar and climate-specific upper limits for sunburn avoidance provides practical management guidance.
Fruchtzonenoffenheit
Traubenbelichtung
Traubenexposition
Traubenfreistellung
bunch exposure
fruit exposure
fruit zone openness
grape cluster illumination
Spayd SE et al. (2002) Separation of sunlight and temperature effects on the composition of Vitis vinifera cv. Merlot berries. American Journal of Enology and Viticulture 53:171-182 Downey MO et al. (2006) Cultural practice and environmental impacts on the flavonoid composition of grapes and wine: a review of recent research. American Journal of Enology and Viticulture 57:257-268 Chorti E et al. (2010) Effects of different cluster sunlight exposure levels on ripening and anthocyanin accumulation in Nebbiolo grapes. American Journal of Enology and Viticulture 61:23-30
Cluster Exposure Quality
The cluster length of Riesling documents short to medium clusters (OIV 202:3-5) measuring 100-160 mm from the proximal branch insertion to the terminal berries — the compact rachis with closely spaced lateral branches produces a relatively short cluster that is wider relative to its length (length-to-width ratio approximately 1.5-2.0) compared to the elongated clusters of Nebbiolo (200-280 mm, length-to-width ratio 3.0-4.0 for the characteristically long, loosely cylindrical Nebbiolo cluster). The cluster length measurement was standardized on 20 representative clusters per cultivar at harvest maturity, with the measurement axis defined as the straight-line distance along the rachis from the most proximal lateral branch to the cluster terminus, excluding the peduncle.
A cluster size description component that records the length of the mature cluster measured from the first (most proximal) branch insertion on the rachis to the tip of the terminal berry group, expressed in millimeters and scored according to OIV descriptor 202 on a scale from very short (state 1, 300 mm), providing the primary dimensional measurement of cluster extent along the rachis axis.
Cluster_Length documents the longitudinal dimension that is the most commonly measured cluster size parameter and the basis of the primary OIV cluster size descriptor (OIV 202). The measurement is straightforward but requires standardization of the measurement endpoints — the proximal boundary is defined at the first branch insertion (not the peduncle insertion, which would include the peduncle length as an additional variable), and the distal boundary at the tip of the terminal berry group (not the rachis tip, which may extend beyond the last berry in some cultivars): this standardization ensures comparability across observers and sites. Cluster length is determined by two developmental factors — the number of lateral branch insertions along the rachis (reflecting the inflorescence branching complexity established during bud organogenesis) and the internode length between successive branch insertions (reflecting rachis elongation during the bloom and post-bloom period) — and both factors are cultivar-specific with environmental modulation. The relationship between cluster length and cluster weight is non-linear because it depends on compactness — a long, loose cluster may weigh less than a short, compact cluster of the same cultivar due to lower berry density per unit of rachis length.
Rappen-Ausdehnung
Trauben-Längenmessung
Traubenlänge
bunch length measurement
cluster longitudinal dimension
rachis extent
Cluster Length
Cluster number assessment for yield estimation in Chardonnay reveals the pruning and fruitfulness interaction determining production, with pruning leaving 20 buds per vine, bud fruitfulness assessment showing 1.8 clusters per bud (36 potential clusters), actual budbreak at 90% (18 shoots from 20 buds), producing 32 clusters per vine (18 shoots × 1.8 clusters minus some weak distal clusters), which at target cluster weight of 150 g projects yield of 4.8 kg/vine (6.4 t/ha at 1333 vines/ha), demonstrating how cluster number integrates pruning decisions with vine fruitfulness to establish yield framework.
Cluster number quality is a quantity yield quality that represents the count of fruit clusters produced per vine, characterizing the first component of yield determination that is established primarily through bud fruitfulness (clusters initiated the previous season) and retention at pruning, with cluster number setting the framework within which cluster weight determines final yield. Within the BFO framework, this quality specifically addresses the cluster count dimension of yield components, distinguishing the number of clusters from their size or composition while recognizing that cluster number is the yield component most directly controlled by pruning decisions (how many buds are retained) and most predictable from bud fruitfulness assessment before budbreak. The ontological significance lies in cluster number being the primary determinant of yield potential that is established early and remains relatively fixed through the season, where the number of clusters emerging from retained buds sets the maximum possible cluster count (barring secondary or tertiary bud contribution), with subsequent cluster thinning providing the primary management tool for reducing cluster number if the natural count exceeds target levels. The determination of cluster number involves the interaction between bud fruitfulness (clusters per bud, determined by conditions during bud differentiation the previous season), bud retention at pruning (how many buds are left per vine), budbreak success (percentage of retained buds producing shoots), and cluster retention (whether all clusters on emerged shoots are retained or some are removed), with each factor contributing to final cluster count. The cultivar variation in typical cluster numbers reflects differences in fruitfulness, training system requirements, and quality optimization, with typical ranges from 15-25 clusters per vine for quality-focused vinifera production to 40-80 clusters for high-yielding juice grape or table grape systems, and with appropriate cluster number depending on variety, clone, training system, and production objectives. Understanding cluster number quality enables yield prediction from pre-season fruitfulness assessment, pruning decisions targeting appropriate bud numbers, and cluster thinning decisions adjusting production toward target levels.
Cluster_Number_Quality addresses the count of fruit clusters determining the first yield component. This quality is controlled by pruning decisions and bud fruitfulness from the previous season. The class supports yield prediction and crop load management through cluster number characterization.
Büschelzahl
Infrukteszenzzählung
Trauben pro Rebe
Traubenzahlqualität
bunch count
bunch number
clusters per vine
infructescence count
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 145-230 May P (2004) Flowering and Fruitset in Grapevines. Lythrum Press, Adelaide, pp. 1-119
Cluster Number Quality
Cluster packing density evaluation of Pinot Noir clones in Oregon reveals Dijon clone 777 producing compact clusters (OIV rating 7-9, packing density 0.78 g/mL) with 40% higher Botrytis incidence compared to Pommard clone producing loose clusters (OIV rating 3-5, packing density 0.58 g/mL), driving clonal selection decisions in humid vintage years where disease pressure outweighs the concentration advantages sometimes associated with compact cluster architecture.
Cluster packing density quality is a bulk density quality that inheres in grape clusters and represents the compactness of berry arrangement within the infructescence structure, quantifying the ratio of berry volume to total cluster volume including interstitial spaces between berries. Within the BFO framework, this quality characterizes the spatial organization emerging from the interaction of berry size, berry number, rachis architecture, and pedicel length that collectively determine how tightly berries are packed within the cluster envelope. The ontological significance extends to cluster compactness serving as a primary determinant of disease susceptibility, where tightly packed clusters with minimal air circulation create humid microenvironments favoring Botrytis cinerea development, while loose clusters with abundant interstitial space promote drying and resist fungal colonization. Cluster packing density is assessed through various methodologies including visual OIV descriptor scales (OIV 204) rating compactness from very loose to very dense, gravimetric approaches calculating density from cluster mass and measured or estimated volume, and image analysis quantifying the two-dimensional packing ratio in standardized photographs. The developmental determination of cluster packing involves both genetic factors controlling rachis length, branch angle, and berry size, and environmental influences including water status during cell division that affects berry number and post-véraison conditions affecting final berry volume. Cultivar differences in cluster packing range from extremely compact varieties like Pinot Noir where berry deformation occurs due to mutual pressure to loose-clustered varieties like Gewürztraminer where individual berries remain distinct. Understanding cluster packing density supports clonal selection for disease resistance, guides canopy management decisions affecting cluster microclimate, and informs sorting decisions where compact clusters may require different processing than loose clusters.
Cluster_Packing_Density_Quality addresses the infructescence organizational scale, distinguishing from Berry_Density_Quality of individual fruits and Canopy_Density_Quality of the vine vegetation mass. This quality aligns with OIV descriptor 204 (bunch density/compactness) but extends beyond ordinal rating to quantitative density measurement. The class encompasses both genetic determination (cluster architecture) and environmental modification (berry size variation) of packing density.
Beerenpachung
Traubenkompaktheit
bunch density
cluster compactness
infructescence packing
Tello J, Ibáñez J (2014) Evaluation of indexes for the quantitative and objective estimation of grapevine bunch compactness. Vitis 53(1): 9-16 Molitor D et al. (2012) Yield and quality of Pinot noir grapes as affected by early leaf removal. OENO One 46(1): 25-30
Cluster Packing Density Quality
The cluster shape description of Cabernet Sauvignon documents a conical shape (OIV 208:2) with one prominent wing — the cluster tapers from a broad shoulder region (where the single elongated primary lateral branch creates a wing extending 30-60 mm laterally from the rachis) to a narrow distal tip, producing the characteristic pyramidal silhouette that is one of Cabernet Sauvignon's most recognizable cluster features. This conical, single-winged shape distinguishes Cabernet Sauvignon from the cylindrical clusters of Pinot Noir (OIV 208:1, uniform width from top to bottom with no wing development), the cylindrical-conical clusters of Riesling (OIV 208:1-2, slight taper but no prominent wing), and the heavily winged clusters of Trebbiano Toscano (OIV 208:2 with 2-3 wings, creating a broad pyramidal cluster with multiple lateral extensions).
A cluster description component that records the overall outline form of the mature cluster according to OIV descriptor 208, classifying clusters as cylindrical (state 1), conical (state 2), funnel-shaped/inverted conical (state 3), or other forms including cylindrical-conical, winged, and branched types, with the shape determined by the spatial distribution of lateral branch lengths along the rachis axis and the number and prominence of wings (elongated primary lateral branches) that create the cultivar-characteristic cluster silhouette.
Cluster_Shape_Description documents the spatial architecture of the cluster that is one of the most visually distinctive and commonly cited cultivar identification characters — the cluster shape is frequently the first descriptor mentioned in popular cultivar descriptions (e.g., "Pinot Noir: small, compact, cylindrical cluster") and is recognizable at vineyard walking distance. The shape is determined by the pattern of lateral branch (wing) development along the rachis — in cylindrical clusters, the lateral branches are of similar length from top to bottom, producing uniform width in conical clusters, the proximal branches are longer than the distal branches, creating the characteristic taper and in winged clusters, one or more proximal branches are disproportionately elongated, producing prominent lateral extensions that break the simple geometric outline: the number and size of wings is a particularly informative character, ranging from no wings (Pinot Noir, yielding simple cylindrical or conical outlines) through one wing (Cabernet Sauvignon, Merlot) to two or more wings (some Trebbiano types, Carignan, many table grapes), with wing development reflecting the branching architecture of the underlying inflorescence primordium established during bud organogenesis. Cluster shape is among the more genetically stable cluster descriptors — while cluster size and weight vary substantially with yield and environment, the fundamental shape (cylindrical versus conical versus winged) is maintained across conditions because it reflects the structural branching pattern rather than the quantitative berry filling of that structure.
Trauben-Architekturtyp
Trauben-Umrissform
Traubenform-Beschreibung
bunch shape classification
cluster architecture type
cluster outline form
Cluster Shape Description
The cluster size description of Grenache documents large clusters (OIV 202:7) with typical dimensions of 200-280 mm length, 120-180 mm width, and 250-500 g weight at harvest — these large clusters, combined with Grenache's high fruitfulness (2-3 clusters per shoot), produce the substantial yield potential (10-15 t/ha without thinning) that makes Grenache one of the highest-yielding premium wine grape cultivars, but also necessitates aggressive cluster thinning for quality-focused production to achieve the 6-8 t/ha yields typical of high-quality Châteauneuf-du-Pape or Priorat wines. The large cluster size of Grenache contrasts with the very small clusters of Pinot Noir (OIV 202:3, 80-130 mm length, 80-150 g) and the medium clusters of Cabernet Sauvignon (OIV 202:5, 140-180 mm length, 100-200 g).
A cluster description component that records the dimensional characteristics of the mature cluster — length, width, and weight — scored according to OIV descriptor 202 (cluster length) on a scale from very small (state 1) to very large (state 9), with cluster size reflecting the cultivar-specific yield potential per cluster that is modulated by fruit set, berry size, and the number and length of rachis branches developed from the original inflorescence architecture.
Cluster_Size_Description integrates the three dimensional measurements — length, width, and weight — that together characterize the overall magnitude of the cluster as a yield-bearing unit. Cluster size is the most environmentally variable of the cluster descriptors, because it reflects the cumulative outcome of multiple developmental processes each subject to environmental modification — inflorescence primordium number (determined in the previous season), flower number per inflorescence, fruit set percentage (typically 20-50%, strongly affected by bloom-period weather), berry number per cluster, and individual berry weight (affected by water availability, crop load, and temperature during berry development): this compounded environmental sensitivity means that cluster size can vary by a factor of 2-3 between seasons for the same cultivar on the same site, making multi-year measurement essential for establishing the cultivar-characteristic size range. Despite this variability, the relative ranking of cultivars by cluster size is maintained across environments — large-clustered cultivars (Grenache, Mourvèdre, Cinsaut) consistently produce larger clusters than small-clustered cultivars (Pinot Noir, Riesling, Muscat à Petits Grains) under any given growing conditions, confirming the strong genetic component underlying the environmental variation.
Trauben-Dimensionserfassung
Trauben-Größenbestimmung
Traubengröße-Beschreibung
bunch dimension record
cluster size assessment
grape cluster measurement
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press
Cluster Size Description
Cluster size spans a 10-fold range across V. vinifera cultivars, from small-clustered Pinot Noir (80-150 g average) to large-clustered Trebbiano Toscano (400-800 g). Cluster weight comprises berry weight (60-75% of total) and rachis weight (2-5%), with berry number being the primary determinant of cluster size variation (r = 0.85). Size consistency within cultivar shows CV of 25-40% depending on vine uniformity and crop load. Cluster size interacts with cluster number to determine yield, with typical relationships of 20-30 clusters × 150 g = 3-4.5 kg/vine for premium wine production.
A cluster architecture disposition that is the inherited capacity of a Vitis vinifera plant to develop clusters of characteristic weight and dimensions, determined by rachis length, branch number, berry count, and individual berry weight, affecting yield potential and optimal crop load management.
Cluster_Size_Disposition serves as the parent class for large and small cluster categories. Size reflects both genetic potential and environmental expression, with significant plasticity in response to crop load, water status, and nutrition. Cluster thinning and shoot thinning modify effective cluster size at the vine level. Size assessment should specify developmental stage, as clusters gain 40-60% of final weight during the post-veraison ripening period.
Gescheingröße-Disposition
Traubengewicht
Traubengröße
bunch dimension capacity
bunch size trait
cluster weight disposition
Clingeleffer PR (2001) Crop development, crop estimation and crop control to secure quality and production of major wine grape varieties. Final Report to GWRDC, CSIRO Plant Industry Dunn GM, Martin SR (2007) A functional association in Vitis vinifera L. cv. Cabernet Sauvignon between the extent of primary branching and the number of flowers formed per inflorescence. Australian Journal of Grape and Wine Research 13:95-100
Cluster Size Disposition
Pinot Noir in Burgundy shows cluster spacing development during late May Year N+1 at BBCH 57 over 2-3 days, with rachis adding 3-5 mm final length distributing spacing to 2.5-3.5 mm between 16-22 primary branches, branch angles adjusting 8-12 degrees optimizing distribution, under continental conditions of 16-20°C temperature, typical of this moderately loose-clustered early-to-mid season cultivar completing architectural refinement before individual flower separation
Cluster spacing development represents the refinement phase where established branch separations enhance through continued differential elongation and positioning adjustments, optimizing spatial distribution of flower groups creating final cluster architecture before individual flower separation. This fine-tuning developmental phase occurs during late May in temperate regions Year N+1 following initial cluster separations, typically spanning 2-3 days as coordinated growth adjustments complete spatial organization preparing for individual flower separation and anthesis. Spacing enhancement mechanism involves continued rachis elongation between branch attachment points increasing inter-branch distances, progressive branch angle adjustment through differential growth refining spatial positioning, and localized branch elongation at specific positions correcting irregularities in flower distribution. Rachis growth during spacing development typically adds 2-5 mm total length distributing increment across inter-branch regions, with growth rate declining from earlier elongation phase as maturation approaches but sufficient to enhance established separations. Branch repositioning involves subtle angular adjustments typically 5-15 degrees refining three-dimensional cluster architecture, with adjustments creating more uniform spatial distribution and optimizing light penetration and air circulation within cluster structure. Localized elongation in specific branches showing compressed spacing provides corrective growth enhancing separation in tight regions, creating more uniform flower group distribution improving microenvironmental uniformity within cluster. Flower bud enlargement continues during spacing development with buds reaching 3-4 mm preparing for individual separation and anthesis, with flower organ maturation within enclosing calyptra approaching completion. Spatial optimization during this phase creates cluster architecture balancing competing requirements including adequate separation for air circulation reducing disease pressure, sufficient compactness for mechanical protection and harvest efficiency, and appropriate exposure for successful pollination and fruit development. Microenvironmental consequences of spacing include light penetration patterns with well-spaced clusters showing good light distribution to interior flowers supporting uniform development, air circulation characteristics with adequate spacing promoting rapid drying reducing infection risk, and temperature distribution with exposed regions showing higher temperatures potentially advancing development. Developmental uniformity within cluster improves during spacing development as flower groups achieve similar maturation states, with good spacing supporting synchronized flowering across cluster promoting uniform fruit set and ripening. Mechanical properties of cluster structure show progressive strengthening as rachis and branch tissues mature, with spacing pattern affecting load distribution affecting cluster mechanical performance under fruit weight and handling stresses. Vascular maturation continues during spacing development with vessels achieving functional capacity adequate for flowering water demands and prepared for substantial increases during berry expansion, with spacing pattern affecting vascular network geometry influencing transport efficiency. Metabolic activity during spacing development shows declining growth-related respiration as elongation slows approaching maturation, transitioning toward maintenance levels though remaining elevated relative to mature vegetative tissues. Resource allocation during spacing development shows continued inflorescence demand competing with vegetative growth and other reproductive structures, with balanced allocation supporting adequate spacing development without excessive vegetative competition. Hormonal regulation during spacing development involves declining gibberellins as elongation processes complete, sustained auxin from developing flowers coordinating final adjustments, and possibly increasing ethylene as maturation processes initiate. Environmental conditions during spacing development include temperature effects continuing with optimal 17-23°C supporting final adjustments, cool conditions potentially limiting refinements maintaining somewhat compressed architecture, and warm conditions accelerating maturation potentially truncating spacing development. Light exposure begins having more direct effects during spacing development as clusters emerge from leaf enclosure, with adequate light supporting robust final development while excessive shade may limit growth reducing spacing achievement. Water availability continues affecting development with adequate moisture supporting turgor-driven adjustments while stress limits growth maintaining established spacing without refinement. Disease management during spacing development becomes increasingly critical as susceptible flowering tissues develop with powdery mildew representing primary concern requiring protective fungicide coverage, with spacing pattern affecting infection risk through microenvironmental effects. The process represents third of four flower cluster separation aspects focusing on refinement rather than establishment, following primary and secondary separations and preceding individual cluster definition and individual flower separation. Duration typically spans 2-3 days representing final adjustment period before individual flower separation initiation, with warm conditions promoting rapid completion. Cultivar variation appears in spacing adjustment extent with some varieties showing substantial refinement creating notably improved uniformity while others show minimal adjustment maintaining initial separation pattern, reflecting genetic regulation of final architectural elaboration. Regional timing shows cluster spacing development during late May in warm climates, late May-early June in temperate regions, and early June in cool areas, occurring in final days before individual flower separation begins. Viticultural significance includes establishing final cluster microenvironment affecting disease pressure and fruit quality, completing architectural expression revealing previous season primordium formation pattern and current season environmental effects, preparing cluster for critical flowering period requiring adequate exposure and air circulation, and finalizing structural framework that will support fruit development through season. Research applications include quantifying spatial optimization processes during final cluster development, analyzing environmental effects on spacing refinement affecting final architecture, evaluating relationships between spacing patterns and disease susceptibility, and assessing functional consequences of architectural variation for fruit quality outcomes.
Cluster_Spacing_Development is third of four Flower_Cluster_Separation subphases, distinguished by refinement of established separations rather than initial establishment. Follows Primary and Secondary Cluster Separation. Transitions to Individual_Cluster_Definition completing group organization. Represents final architectural adjustments before individual flower separation. Optimizes cluster microenvironment for flowering and development.
Abstands-Entwicklung
Feinabstimmung der Gliederung
Gescheinsraumoptimierung
architektonische Verfeinerung
cluster architecture optimization
final spacing adjustment
spacing refinement
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 348-352 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Cluster Spacing Development
The cluster thinning protocol for Pinot Noir in the Côte de Nuits specifies vendange verte (green harvest) at E-L 33-35 (bunch closure to onset of véraison) retaining a maximum of 7-8 clusters per vine (1 cluster per shoot on the strongest shoots, removing second clusters entirely) — removal priorities are second clusters on each shoot (typically smaller and later-ripening), clusters with poor set showing extensive millerandage, and clusters on the weakest shoots with fewer than 10 leaves above the cluster, with the objective of reducing potential yield from 55-60 hl/ha to the target 38-42 hl/ha while ensuring that all retained clusters have a minimum of 12-15 cm² functional leaf area per gram of fruit, exceeding the 10 cm²/g threshold below which Pinot Noir fails to achieve adequate phenolic maturity in the Burgundian climate.
A canopy management description that documents the selective removal of entire grape clusters from the vine during the growing season to reduce crop load, improve the ratio of leaf area to fruit weight, and enhance the concentration and ripening uniformity of retained clusters, specifying the timing (phenological stage), target cluster number per vine or per shoot, cluster selection criteria (removal priorities), and target yield after thinning.
Cluster_Thinning_Protocol documents the most direct yield reduction intervention available during the growing season, widely practiced in premium viticulture where quality-oriented yield targets fall below the vine's natural productive capacity. The practice is conceptually simple — remove excess clusters to concentrate vine resources into fewer, better-ripened clusters — but the timing, severity, and selection criteria significantly influence the qualitative outcome: early thinning at fruit set (E-L 27) provides the longest period for compensatory berry enlargement in retained clusters (partially offsetting the yield reduction), while late thinning at véraison (E-L 35) minimizes compensatory growth but wastes vine resources already invested in the removed clusters. The optimal thinning timing for quality is generally considered to be around bunch closure to early véraison (E-L 33-35), when cluster quality assessment (set uniformity, health, ripening progression) is most reliable and sufficient season remains for the vine to redirect resources to retained fruit. The economic cost of cluster thinning is substantial — 40-80 labor hours/ha for manual removal, representing a direct input cost of 400-800 EUR/ha plus the opportunity cost of discarded crop.
Ertragsreduktions-Protokoll
Grünlese-Spezifikation
Traubenausdünnungs-Protokoll
crop thinning specification
green harvest protocol
vendange verte protocol
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 370-378 Galet P (2000) General Viticulture, Oenoplurimedia pp: 310-330 Reynolds AG et al. (2007) Magnitude of viticultural and enological effects I: Relative impacts of cluster thinning and yeast strain on composition of Chardonnay Musqué, American Journal of Enology and Viticulture 58: 25-37
Cluster Thinning Protocol
Cluster volume assessment in Sangiovese for Brunello di Montalcino production reveals the typical heterogeneity of this cultivar, with cluster volumes ranging from 80-180 mL within a single vine due to variable fruit set (60-120 berries per cluster), necessitating cluster-by-cluster sorting at harvest based on visual assessment of cluster architecture, compactness, and berry uniformity to select only well-formed, moderately loose clusters (volume 100-140 mL) for fermentation of the primary wine.
Cluster_Volume_Quality represents the three-dimensional spatial extent of the entire grape cluster (infructescence) including the rachis framework and all attached berries. This quality integrates individual berry volumes with cluster architecture to characterize the overall fruit unit that is the basis of harvest yield and the practical unit for many viticultural operations. Cluster volume varies enormously across cultivars and growing conditions, typically ranging from 50-500 mL for wine grapes and reaching 1000-2000 mL for some table grape cultivars with elongated clusters. Cluster volume is determined by the number of berries (typically 50-200 for wine grapes), individual berry volume, and the spatial arrangement imposed by rachis branching pattern. Loose clusters occupy greater volume relative to total berry volume than compact clusters, with important implications for air circulation, spray penetration, and disease susceptibility. Cluster volume measurement employs water displacement of entire clusters or volumetric estimation from cluster dimensions (length, width, depth) using shape-specific coefficients. The relationship between cluster volume and cluster weight depends on berry density and cluster compactness, with typical density values of 0.6-0.9 g/mL for intact clusters (reflecting interberry air spaces). Cluster volume assessment informs yield estimation when combined with cluster counts, and contributes to harvest logistics planning for bin and tank capacity allocation.
Cluster_Volume_Quality encompasses the entire infructescence as a functional unit, distinct from the summation of individual berry volumes which would exclude rachis volume and interberry spaces. Cluster compactness indices relate cluster volume to berry number or cluster weight, providing standardized descriptors for this important quality character. The OIV descriptor 204 assesses cluster compactness on a categorical scale (very loose to very compact) that can be related to volumetric measurements. Cluster volume changes through the season as berries develop, reaching maximum at harvest maturity. Post-véraison water deficit can reduce cluster volume through both berry shrinkage and berry abscission in extreme cases.
Gescheinvolumen
Traubenrauminhalt
Traubenvolumen
bunch volume
cluster spatial extent
grape bunch volumetric measurement
infructescence volume
Tello J and Ibáñez J (2018) What Do We Know about Grapevine Bunch Compactness? A State-of-the-Art Review. Australian Journal of Grape and Wine Research 24(1): 6-23 OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 87-91
Cluster Volume Quality
The cluster weight of Muscat of Alexandria documents heavy clusters (OIV state 7) typically weighing 400-800 g at table grape harvest maturity — reflecting the combination of large berries (5-8 g each), moderate to high berry number (80-150 berries per cluster), and extensive rachis development characteristic of this dual-purpose cultivar grown for both table grape and wine production. This high cluster weight contrasts with the low cluster weight of Pinot Noir (80-150 g, reflecting small berry size of 1.0-1.5 g and moderate berry number) and the very high cluster weight of Italia table grape (500-1,200 g, with individual berries reaching 8-12 g), illustrating the ten-fold range of cluster weight variation across V. vinifera cultivars from small-clustered wine grapes to large-clustered table grapes.
A cluster size description component that records the mass of the mature cluster at harvest, expressed in grams and scored on a scale from very low (state 1, 1000 g), providing the most directly yield-relevant cluster size measurement that integrates berry number, berry weight, and rachis weight into a single parameter reflecting the total fruit production per reproductive unit.
Cluster_Weight is the most integrative cluster size parameter — it reflects the multiplicative combination of berry number per cluster (determined by flower number and fruit set percentage) and mean berry weight (determined by seed number, cell division during the pre-véraison growth phase, and cell expansion during the post-véraison ripening phase), plus the rachis weight (typically 3-7% of total cluster weight in most wine grape cultivars). This multiplicative relationship means that cluster weight is the most variable of all cluster descriptors — combining the variation in berry number (coefficient of variation typically 20-30%) with the variation in berry weight (CV 15-25%) produces cluster weight CVs of 30-50% within a single cultivar-site-year combination: this high variability necessitates large sample sizes (typically 20-50 clusters per replicate) for reliable cultivar characterization and treatment comparison in viticultural research. Despite this variability, cluster weight is the most commercially relevant cluster parameter because it directly determines yield per vine (cluster weight × clusters per vine = yield) and is the basis for crop estimation models used to predict harvest volume and schedule processing capacity. The distinction between wine grape cluster weights (typically 80-300 g for premium cultivars) and table grape cluster weights (typically 300-1,200 g) reflects the fundamentally different quality criteria — wine grape quality generally benefits from smaller clusters and berries (higher skin-to-pulp ratio, lower yield, more concentrated flavors), while table grape quality requires large, visually attractive clusters with large, firm berries.
Ernte-Traubengewicht
Trauben-Massebestimmung
bunch weight measurement
cluster mass record
harvest cluster weight
Cluster Weight
Cluster weight assessment comparing irrigation treatments in Syrah reveals the water status effect on this yield component, with well-watered treatment producing clusters averaging 285 g (145 berries × 1.97 g/berry), moderate deficit producing 215 g (138 berries × 1.56 g/berry), and severe deficit producing only 145 g (125 berries × 1.16 g/berry), with the 50% cluster weight reduction under severe deficit arising from both reduced berry number (14% fewer) and reduced berry size (41% smaller), demonstrating how cluster weight integrates multiple developmental effects that accumulate through the season.
Cluster weight quality is a quantity yield quality that represents the mass of individual fruit clusters at harvest, characterizing the second major yield component that combines with cluster number to determine total production, with cluster weight resulting from the integration of berry number per cluster and berry weight that together determine the mass of each reproductive unit. Within the BFO framework, this quality specifically addresses the cluster mass dimension of yield components, distinguishing the weight of individual clusters from their number or internal composition while recognizing that cluster weight varies substantially with cultivar (genetic potential), vine condition (resource supply), and seasonal factors (weather during fruit development), with this variation affecting both total yield and cluster handling characteristics for harvest and processing. The ontological significance lies in cluster weight being the yield component that integrates fruit development success during the current season, where unlike cluster number (largely fixed early), cluster weight responds to growing conditions throughout fruit development, with favorable conditions (adequate water, nutrients, moderate temperatures) supporting larger clusters while stress conditions (drought, heat, nutrient deficiency) reduce cluster weight through effects on berry number and berry size. The determinants of cluster weight include berry number per cluster (set at fruit set, affected by flowering conditions and fruit set success), berry weight (determined through berry development, affected by water status and source-sink balance), and the mathematical relationship cluster weight = berry number × berry weight, with these components under somewhat independent control such that the same cluster weight can result from many small berries or fewer large berries with different implications for fruit surface-to-volume ratio and extraction characteristics. The cultivar variation in typical cluster weight ranges from small-clustered varieties (50-100 g, such as Pinot Noir) through moderate (150-300 g, such as Chardonnay, Merlot) to large-clustered varieties (400-800 g, such as Thompson Seedless, some Muscats), with this genetic variation establishing the baseline around which environmental and management factors cause additional variation. Understanding cluster weight quality enables yield assessment from cluster counts, identification of fruit development limitations affecting cluster mass, and harvest logistics planning based on cluster handling characteristics.
Cluster_Weight_Quality addresses the mass of individual fruit clusters as a yield component. This quality integrates berry number and berry weight reflecting development success. The class supports yield assessment and harvest planning through cluster weight characterization.
Büschelgewicht
Infrukteszenzmasse
Traubengewichtqualität
Traubenmasse
bunch mass
bunch weight
cluster mass
infructescence weight
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 145-230 Matthews MA, Nuzzo V (2007) Berry size and yield paradigms on grapes and wines quality. Acta Horticulturae 754: 423-435
Cluster Weight Quality
The cluster width of Cabernet Sauvignon documents medium width of 60-100 mm measured perpendicular to the rachis at the level of the first or second wing — combined with a cluster length of 140-180 mm, this produces a length-to-width ratio of approximately 1.8-2.5, reflecting the conical to cylindrical-conical cluster shape characteristic of Cabernet Sauvignon. The width measurement captures the lateral extension of the cluster determined by the length of the primary lateral branches (wings) and the number of berries borne on each branch — Cabernet Sauvignon's single prominent wing (shoulder) at the proximal end creates a wider upper cluster region tapering to a narrower distal end, producing the conical silhouette. This width profile contrasts with the uniformly narrow clusters of Nebbiolo (width 40-60 mm, producing the characteristically elongated cylinder with length-to-width ratio of 3.0-4.0) and the broad, shouldered clusters of Trebbiano Toscano (width 100-150 mm, with multiple wings creating a wide pyramidal shape).
A cluster size description component that records the maximum width of the mature cluster measured perpendicular to the rachis axis at the widest point, expressed in millimeters, providing the lateral dimensional measurement that together with cluster length characterizes the overall proportions and silhouette of the cluster and enables the calculation of the length-to-width ratio used in shape classification.
Cluster_Width captures the lateral dimension that is particularly informative for cluster shape classification — the distribution of width along the cluster length (uniform width producing cylindrical shapes, proximal maximum producing conical shapes, bimodal maxima producing winged shapes) provides shape information that complements the overall length and weight measurements. The width at any point along the rachis is determined by the length and berry load of the lateral branches at that position — positions with long, heavily-laden branches (wings or shoulders) produce width maxima, while positions with short or absent branches produce width minima: the cultivar-specific pattern of branch length variation along the rachis therefore directly determines the cluster silhouette when viewed laterally. Cluster width is more sensitive to fruit set variation than cluster length — high fruit set increases width (by filling lateral branches with more, larger berries) more than it increases length (which depends on rachis elongation rather than berry number), meaning that the length-to-width ratio decreases with increasing fruit set and compactness, and the environmental variation in ratio provides an indirect measure of fruit set conditions during the bloom period.
Trauben-Breitenmessung
Traubenbreite
bunch width measurement
cluster lateral dimension
maximale Traubenbreite
maximum cluster breadth
Cluster Width
Cluster zone microclimate data for east-exposed Cabernet Sauvignon in Oakville, Napa Valley showed berry surface temperatures of 32-36°C during morning sun exposure (09:00-12:00) versus 38-44°C on the west-exposed face during afternoon (14:00-17:00), with anthocyanin concentration 25-30% higher in morning-exposed berries due to reduced thermal degradation. Cluster zone monitoring in Burgundy Pinot Noir following targeted basal leaf removal (6 leaves stripped at fruit set) increased cluster zone PAR from 8% to 28% of ambient, reduced cluster zone humidity from 92% to 78% at dawn, and decreased Botrytis incidence from 35% to 8% of clusters at harvest. Riesling cluster zone microclimate in the Pfalz under traditional single-curtain training maintained berry temperatures below 32°C even during the 2019 and 2020 heat events, contrasting with exposed Dornfelder clusters on adjacent blocks reaching 42°C at the berry surface.
Cluster zone microclimate is a microclimate characterization entity that records the environmental conditions specifically within the fruiting zone of a grapevine canopy where clusters develop and ripen, including berry surface temperature, fruit zone radiation (PAR and UV), relative humidity at the cluster surface, and air circulation around clusters, directly governing sugar accumulation, organic acid metabolism, phenolic biosynthesis, aromatic compound development, and pathogen infection probability during the critical period from fruit set through harvest.
Cluster zone microclimate represents the most direct environmental determinant of berry composition and thus wine quality, making its characterization a central objective of canopy management research. The cluster zone differs from the general canopy microclimate in its proximity to the ground (lower in most training systems), its shading by surrounding leaf layers, and the localized humidity generated by berry transpiration. Temperature within the cluster zone influences virtually all aspects of berry metabolism: sugar accumulation rate, malic acid respiration (accelerated above 30°C), anthocyanin synthesis (optimal at 17-25°C, inhibited above 35°C), tannin polymerization, and volatile aroma compound retention. The distinction between berry skin surface temperature and ambient air temperature is critical — direct solar radiation can elevate berry surface temperature 8-15°C above ambient, creating conditions for sunburn (berry surface > 40°C) and anthocyanin degradation even when air temperature remains moderate. Leaf removal timing and intensity, training system height, row orientation, and cluster thinning collectively modulate cluster zone microclimate, making this data essential for optimizing the balance between sun exposure benefits (color, phenolic ripeness) and excessive heat risks.
Beerenzonen-Mikroklima
Traubenzonenmikroklima
Traubenzonenumgebung
berry zone microclimate
bunch zone environment
cluster environment data
fruiting zone climate
Bergqvist J et al. (2001) Sunlight exposure and temperature effects on berry growth and composition of Cabernet Sauvignon and Grenache in the Central San Joaquin Valley of California. American Journal of Enology and Viticulture 52(1): 1-7 Spayd SE et al. (2002) Separation of sunlight and temperature effects on the composition of Vitis vinifera cv. Merlot berries. American Journal of Enology and Viticulture 53(3): 171-182 Tardaguila J et al. (2010) Impact of early leaf removal on yield and fruit and wine composition of Vitis vinifera L. Graciano and Carignan. American Journal of Enology and Viticulture 61(3): 372-381
Cluster Zone Microclimate
Ugni Blanc (Saint-Émilion) bears a Cognac_Production_Scion_Role as the near-exclusive variety for modern Cognac, replacing the phylloxera-susceptible Folle Blanche and Colombard that dominated pre-crisis production—its neutral character, exceptional acidity (10-12 g/L even in warm vintages), late ripening (reducing frost risk), and disease tolerance making it ideally suited to Cognac's maritime climate and distillation requirements. Folle Blanche serves in a Cognac_Production_Scion_Role for prestige cuvées from producers (Château de Beaulon, Leopold Gourmel) seeking the delicate floral aromatics and finesse historically associated with pre-phylloxera Cognac, though its thin skin and rot susceptibility limit commercial viability. Colombard bears a Cognac_Production_Scion_Role contributing aromatic complexity and fruit character to blends, though it now represents less than 1% of Cognac plantings.
A distillation scion role that inheres in a grafted scion cultivar by virtue of its authorization and suitability for Cognac production within the delimited Cognac appellation of southwestern France, where base wine undergoes double distillation in traditional Charentais pot stills (alembics) followed by minimum two years of oak barrel aging. Cognac production scions must be approved varieties under AOC regulations, with Ugni Blanc (locally called Saint-Émilion) dominating current production at over 98% of plantings, supplemented by minor permitted varieties including Folle Blanche and Colombard. Cognac varieties must produce base wines of high acidity (9-12 g/L, essential for freshness through distillation), low alcohol (8-9%, minimizing harsh higher alcohols), neutral-to-delicate character (avoiding varietally assertive flavors that would concentrate unpleasantly), and sufficient disease resistance for the humid maritime climate. The double distillation (two passes through copper pot stills) and extended oak aging (VS minimum 2 years, VSOP 4 years, XO 10 years) develop the complex floral, fruity, and spicy character defining quality Cognac.
Cognac_Production_Scion_Role captures the strictly regulated varietal requirements of AOC Cognac, where variety authorization, viticulture practices, distillation methods, and aging protocols are all controlled. Ugni Blanc's dominance represents a post-phylloxera rationalization—the variety's disease resistance and consistent quality displaced the more aromatic but temperamental Folle Blanche and Colombard that created historic Cognac's reputation. Recent interest in terroir expression and stylistic diversity has prompted limited revival of minor varieties, with some producers championing single-variety Folle Blanche or Colombard Cognacs. Climate change threatens Cognac production by accelerating ripening and reducing acidity experimental plantings of heat-tolerant varieties and potential appellation modification of permitted varieties are under discussion.
Charentais variety
Charentais-Sorte
Cognac grape
Cognac variety
Cognac-Rebsorte
Cognac-Traube
Bureau National Interprofessionnel du Cognac (BNIC) (2020) Cognac Production Regulations Léauté R (1990) Distillation in alembic, American Journal of Enology and Viticulture 41: 90-103 Faith N (2016) Cognac, Infinite Ideas Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane pp.: 1101-1103
Cognac Production Scion Role
Cohesion quality maintenance in Italia table grapes during 90-day cold storage (0-2°C, 90-95% RH) requires controlled atmosphere conditions (5% CO₂, 3% O₂) that slow pectin solubilization and middle lamella degradation fruit maintaining high cohesion through storage (flesh separation force above 0.8 N) displays the firm, crunchy texture expected by consumers, while cohesion breakdown (force below 0.4 N) produces the mealy, disaggregating texture that renders fruit unmarketable regardless of sugar content, color, or external appearance.
Cohesion_Quality represents the structural integrity quality characterizing the internal binding forces that hold the tissues of a grapevine organ together as a unified structure. Cohesion reflects the strength of intercellular adhesion mediated primarily by the pectin-rich middle lamella that cements adjacent cells together, as well as the structural contribution of cell walls and the mechanical effect of turgor pressure pressing cells against their neighbors. High cohesion produces tissues that resist separation when forces are applied, maintaining structural unity under the stresses of handling, processing, and mastication. Low cohesion allows tissues to separate readily into individual cells or cell groups, creating the mealy, grainy, or disintegrating texture associated with overripe or improperly stored fruit. Cohesion changes during berry development and ripening as middle lamella composition is modified by pectin-degrading enzymes including polygalacturonase and pectin methylesterase that solubilize the cementing polysaccharides. Pre-véraison berries display high cohesion with cells firmly adhered, while ripening progressively reduces cohesion as part of the softening process. Excessive cohesion loss produces berries where the flesh falls apart upon chewing or pressing, releasing juice but providing poor textural sensation. Cultivars differ in the rate and extent of cohesion loss during ripening, distinguishing firm table grape varieties that maintain cohesion through extended storage from soft wine grape varieties that lose cohesion rapidly. Cohesion assessment employs mechanical tests measuring the force required to separate tissue layers, sensory evaluation of mealiness and cell separation during chewing, and microscopic observation of tissue structure after mechanical stress.
Cohesion_Quality addresses the internal binding of tissues within an organ, distinct from Attachment_Strength_Quality which addresses connections between organs. Cohesion is closely related to texture qualities, particularly the firmness and crispness aspects of Internal_Texture_Quality, but specifically focuses on intercellular adhesion rather than cellular mechanical properties. The middle lamella is the primary anatomical determinant of cohesion, with its pectin composition and degree of polymerization controlling adhesion strength. Cohesion assessment at the tissue level complements texture assessment at the organ level, providing insight into the cellular basis of textural properties.
Gewebezusammenhalt
Kohäsion
intercellular adhesion
interzelluläre Adhäsion
structural unity
strukturelle Bindung
tissue binding
tissue cohesiveness
Nunan KJ et al. (1998) Changes in Cell Wall Composition during Ripening of Grape Berries. Plant Physiology 118(3): 783-792 Brummell DA (2006) Cell Wall Disassembly in Ripening Fruit. Functional Plant Biology 33(2): 103-119
Cohesion Quality
A coiled functional tendril at node 11 on a shoot of Vitis vinifera cv. Pinot Noir in Burgundy, France, having made contact with a horizontal trellis wire three days previously, exhibiting four tight contact coils wrapped around the 2.5 mm diameter wire with coil diameters of 8-10 mm, connected to the shoot base through seven free coils in the 35 cm tendril shaft showing characteristic alternating clockwise-counterclockwise handedness that accommodates shoot movement, demonstrating structural reinforcement through increased lignification visible as brown coloration in coiled regions compared to green pre-contact tissue, supporting a 65 cm shoot section bearing four leaves and one fruit cluster with combined weight of approximately 320 grams, measured at BBCH stage 77 (berries beginning to touch) in mid-July.
A functional tendril that has undergone thigmotropic coiling response following sustained contact with a support structure in Vitis vinifera, characterized by formation of helical coils through differential cell elongation on opposite sides of the tendril axis, development of both contact coils directly gripping the support structure and free coils in unattached tendril sections between the support and the shoot base, achievement of mechanical anchoring function supporting vine weight and resisting wind-induced displacement forces, and structural modifications including enhanced lignification and cellulose reinforcement in coiled regions that increase tensile strength and elastic resilience. Coiled functional tendrils represent the successful culmination of tendril searching behavior and contact detection, executing the species' primary climbing strategy through rapid conversion of exploratory growth into permanent mechanical attachment. The coiling process involves sophisticated coordination of cellular responses including asymmetric cell wall loosening on the concave side, enhanced cell expansion on the convex side, differential lignin deposition, and formation of specialized gelatinous fibers in tension wood that generate the contractile forces driving coil formation and maintenance. Coiled tendrils create spring-like mechanical systems that absorb shock loading from wind gusts, accommodate shoot growth-induced position changes, and distribute mechanical stresses across multiple attachment points, with properly coiled tendrils capable of supporting loads exceeding 5-10 kilograms depending on tendril diameter, coil density, and tissue maturation state.
Coiled functional tendrils are distinguished from uncoiled searching tendrils by presence of helical structure and from attached but straight tendrils by formation of actual coils rather than simple wrapping or adhesion. The coiling pattern shows characteristic features including bidirectional coiling (reversal of handedness) in free coil sections that prevents twisting of the shoot, tighter coils in contact regions that maximize grip, and progressive coil tightening over 24-72 hours post-contact as lignification proceeds. Coiling degree varies with cultivar genetics, tendril position, support structure diameter, and environmental conditions during coiling. Failed coiling attempts where contact is broken before coil fixation results in permanently deformed but non-functional tendrils. Environmental stress during coiling can produce incomplete or weak coils with reduced load-bearing capacity.
coiled climbing organ
gewickelte Ranke
gewundene Kletterorgan
helical tendril
spiraled tendril
spiralisierte Ranke
wound tendril
Based on Bowling & Vaughn (2009) "Gelatinous fibers are widespread in coiling tendrils and twining vines" American Journal of Botany 96=719-727 Goriely & Tabor (2013) "Spontaneous helix hand reversal and tendril perversion in climbing plants" Physical Review Letters 80=1564-1567 Silk & Hubbard (1991) "Axial forces and normal stresses in coiling tendrils" Biomechanics and Modeling in Mechanobiology 24=599-606 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 95-118.
Coiled Functional Tendril
Cold acclimation quality monitoring in Finger Lakes Riesling tracked the seasonal development of freezing tolerance to establish the timeline of hardiness acquisition and identify the environmental factors driving acclimation progression. Weekly bud hardiness testing from September through January revealed that cold acclimation quality developed in distinct phases. Initial acclimation during September-October was triggered primarily by photoperiod reduction, with LT50 improving from the summer baseline of -5°C to -12°C independent of temperature fluctuations, representing the photoperiod-driven first stage of acclimation. Accelerated acclimation during November required exposure to temperatures below 5°C, with each week of cold exposure improving LT50 by approximately 2-3°C until reaching -20°C by late November. Maximum cold acclimation quality achievement in December-January required sustained near-freezing temperatures, with LT50 reaching the cultivar maximum of -26°C only after cumulative exposure to temperatures below 0°C. The cold acclimation quality assessment identified that warm autumn conditions delaying the onset of sustained cold exposure resulted in incomplete hardening by the time of first severe freeze events, explaining the greater winter injury observed in years with warm November weather compared to years with normal autumn cooling progression.
Cold acclimation quality is a cold stress indicator quality that inheres in a grapevine organism and represents the capacity to develop enhanced freezing tolerance through the progressive physiological and biochemical changes induced by exposure to low, non-freezing temperatures and shortening photoperiod during autumn, with the acclimation process transforming cold-sensitive summer tissues into cold-hardy winter tissues capable of surviving temperatures that would be lethal to non-acclimated vines. This quality captures the dynamic, inducible component of cold hardiness that supplements constitutive tolerance, recognizing that the substantial seasonal variation in grapevine freezing tolerance reflects an active acclimation process that must occur each autumn to prepare vines for winter survival, with the rate, extent, and stability of acclimation determining actual winter hardiness achieved. The ontological significance of cold acclimation quality lies in its role as the mechanism by which grapevines achieve the substantial increases in freezing tolerance required for winter survival in cold climates, with maximum hardiness representing a 20-30°C improvement over summer tolerance levels achievable only through successful completion of the acclimation process, such that factors compromising acclimation capacity directly affect winter survival probability. The biological basis of cold acclimation encompasses multiple coordinated changes including the cessation of growth removing metabolically active tissues most vulnerable to cold injury, the accumulation of cryoprotective compounds including sugars and specific proteins that stabilize cellular structures during freezing, the modification of membrane lipid composition to maintain fluidity at low temperatures, and the dehydration of tissues reducing the water available for damaging intracellular ice formation. The acclimation process proceeds in stages, with initial growth cessation and metabolic transition triggered by shortening photoperiod and mild temperature decline, followed by deeper hardening requiring sustained exposure to temperatures near or slightly below 0°C that trigger the full expression of cold-protective mechanisms. The genetic variation in cold acclimation quality among grapevine genotypes reflects differences in the sensitivity and responsiveness of the acclimation signaling pathways, the capacity for cryoprotectant accumulation, and the effectiveness of the protective mechanisms deployed, providing targets for selection and breeding to improve cold hardiness through enhanced acclimation capacity.
Cold_Acclimation_Quality is_a Cold_Stress_Indicator_Quality, serving as parent class for Acclimation_Rate_Quality, Maximum_Hardiness_Level_Quality, and Acclimation_Stability_Quality which address specific dimensions of the acclimation process. This class addresses the overall capacity for cold hardening while child classes capture particular aspects. The two-stage acclimation model (photoperiod-triggered, then cold-temperature-dependent) explains environmental requirements. The 20-30°C hardiness improvement from summer to winter demonstrates the magnitude of acclimation effects. Critical for understanding winter survival in cold-climate viticulture.
Frostabhärtung
Kälteakklimatisierung
Kältekonditionierung
Winterakklimatisierung
cold conditioning
cold hardening
frost hardening
winter acclimation
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Wake CMF, Fennell A (2000) Morphological, physiological and dormancy responses of three Vitis genotypes to short photoperiod. Physiologia Plantarum 109:203-210 Mills LJ et al. (2006) Timing of low temperature exposure affects cold hardiness and cold deacclimation kinetics in grape. American Journal of Enology and Viticulture 57:20-26
Cold Acclimation Quality
In Chardonnay vines in Champagne, cold acclimation start commences in early October when mean daily temperatures fall below 12°C, with measurable increases in CBF transcription factor expression within 48 hours of initial cold exposure, initial accumulation of raffinose detected within one week, and bud cold hardiness improving from -4°C to -10°C within the first two weeks of cool temperature exposure at the onset of BBCH stage 91.
Cold acclimation start in Vitis vinifera represents the initial phase of temperature-induced hardening during which exposure to cool but non-freezing autumn temperatures first activates the molecular and cellular processes that will progressively increase freezing tolerance over the following weeks. This phase typically commences when ambient temperatures consistently fall below approximately 10-15°C, triggering the first transcriptional and metabolic responses that mark the beginning of the acclimation continuum. The earliest molecular events during cold acclimation start include rapid activation of CBF/DREB transcription factors within hours of cold exposure, increased expression of immediate early cold-responsive genes encoding signaling components and transcriptional regulators, and initial changes in membrane lipid desaturase activity that begin the shift toward more unsaturated membrane composition. Calcium signaling plays important roles in cold perception and early signal transduction, with cold-induced cytosolic calcium increases activating calcium-dependent protein kinases and other signaling components that contribute to downstream cold responses. The initial phase of acclimation produces relatively modest increases in cold hardiness, typically improving freezing tolerance by 5-8°C within the first 1-2 weeks, establishing the foundation for more substantial hardiness gains during subsequent acclimation stages at lower temperatures. Cellular changes during cold acclimation start include initial accumulation of compatible solutes as genes encoding biosynthetic enzymes are upregulated, early dehydrin protein synthesis, and subtle changes in protein complement as cold-induced genes begin expression while some growth-related genes are repressed. The effectiveness of cold acclimation start is enhanced by prior short-day exposure, with photoperiod pre-conditioning improving both the speed and magnitude of cold hardiness development at any given temperature, illustrating the integration of photoperiod and temperature signals in dormancy induction. The timing of cold acclimation start relative to the first freezing temperatures significantly affects vine survival, with early-acclimating genotypes and well-conditioned vines showing greater resilience to early cold events than late-acclimating or poorly conditioned vines.
Cold_Acclimation_Start represents the initial phase within Temperature_Acclimation, specifically addressing the first responses to cool autumn temperatures. Distinguished from Freeze_Tolerance_Development which addresses the more advanced stages of hardening at lower temperatures. Occurs at temperatures of 10-15°C before freezing exposure. Duration typically spans 2-3 weeks. Produces initial hardiness gains of 5-8°C. Enhanced by prior short-day exposure. Key molecular events include CBF activation and early cold-responsive gene expression.
Kälteabhärtungs-Beginn
cold acclimation onset
early cold acclimation
erste Abhärtungsphase
first-stage hardening
frühe Frostabhärtung
initial cold hardening
initiale Kälteakklimatisierung
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Xin Z, Browse J (2000) Cold comfort farm: the acclimation of plants to freezing temperatures. Plant Cell and Environment 23: 893-902 Thomashow MF (1999) Plant cold acclimation: freezing tolerance genes and regulatory mechanisms. Annual Review of Plant Physiology 50: 571-599
Cold Acclimation Start
Cold damage symptom quality assessment following a January polar vortex event (-32°C) in Niagara Peninsula provided systematic evaluation of tissue injury patterns to guide vineyard recovery management. Bud cross-section analysis revealed cold damage symptom quality with clear browning in primary bud primordia in 75% of sampled buds, indicating lethal freezing injury to the meristematic tissue, while secondary buds showed browning in only 25% of samples, consistent with their 3-4°C greater hardiness providing partial protection. Cane sectioning showed cold damage symptom quality with phloem browning visible as a tan-to-brown discoloration in the inner bark layer in 40% of cane cross-sections, indicating vascular damage that would limit spring sap flow and nutrient transport. Cambium assessment through the scraping technique revealed cold damage symptom quality with brown rather than green coloration at the trunk base in 15% of vines, concentrated in young plantings and those with prior crown gall damage, indicating potential trunk mortality requiring retraining from surviving basal shoots. The systematic cold damage symptom quality evaluation enabled damage mapping across the vineyard, identifying blocks requiring aggressive remedial pruning to redirect growth to surviving secondary buds versus blocks with minimal damage permitting normal pruning approaches.
Cold damage symptom quality is a cold stress indicator quality that inheres in a grapevine organism and represents the visible and microscopically detectable signs of tissue injury resulting from freezing stress, providing the diagnostic basis for identifying cold damage occurrence, assessing damage severity, and distinguishing cold injury from other causes of tissue dysfunction through recognition of the characteristic symptom patterns associated with ice formation and freeze-induced cellular death in different vine tissues. This quality captures the observational dimension of cold stress assessment, recognizing that effective diagnosis of cold damage requires familiarity with the tissue-specific symptoms that distinguish freezing injury from drought stress, disease, or other causes of similar-appearing damage, enabling accurate damage assessment that informs recovery management decisions. The ontological significance of cold damage symptom quality lies in its role enabling rapid field-based damage assessment through visual inspection supplemented by simple diagnostic techniques including tissue sectioning and browning evaluation, providing the practical basis for post-freeze damage surveys that quantify injury extent and guide management response without requiring laboratory analysis or extended observation periods. The biological basis of cold damage symptoms encompasses the visible consequences of ice formation and cellular death including the oxidation of cellular contents producing characteristic browning, the loss of turgor in killed tissues, the disruption of vascular function preventing normal water and nutrient transport, and the secondary colonization of damaged tissues by saprophytic organisms that may modify symptom appearance. The timing of symptom development varies among symptom types, with some damage becoming visible within hours of thawing as oxidation progresses in killed tissues while other symptoms including vascular dysfunction may not become apparent until growth resumption reveals the functional consequences of structural damage. The tissue-specific nature of cold damage symptoms reflects the different responses of various tissue types to freezing injury, with buds, cambium, phloem, and xylem each showing characteristic symptom patterns that enable diagnosis of which tissues have been damaged and to what extent, informing predictions about vine recovery capacity and productivity consequences.
Cold_Damage_Symptom_Quality is_a Cold_Stress_Indicator_Quality, serving as parent class for Bud_Browning_Quality, Cambium_Browning_Quality, Phloem_Necrosis_Quality, and Xylem_Discoloration_Quality which address specific tissue symptoms. This class addresses the overall diagnostic approach while child classes capture tissue-specific symptom patterns. The cross-section browning method provides standard diagnostic technique. Timing of assessment (post-thaw to spring growth) affects symptom visibility. Critical for post-freeze damage surveys informing management response.
Frostschaden-Zeichen
Gefrierschaden-Indikatoren
Kälteschaden-Symptome
Winterverletzungs-Diagnose
cold damage signs
freeze injury symptoms
frost damage indicators
winter injury diagnosis
Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection. Michigan State University Extension Bulletin E2930 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 508-545
Cold Damage Symptom Quality
Cold hardiness assessment tracking Riesling through the dormant season reveals the dynamic nature of freeze tolerance, with October showing early acclimation (LT50 of -8°C, buds vulnerable to early freeze), December showing advanced hardiness (LT50 of -22°C, substantial protection developing), mid-January showing maximum hardiness (LT50 of -25°C, peak tolerance coinciding with typical coldest period), February showing maintained hardiness (LT50 of -23°C, slight decline but still protected), and late March showing rapid deacclimation (LT50 of -12°C, protection diminishing as spring approaches), demonstrating the seasonal trajectory that must align with local temperature patterns for successful winter survival.
Cold hardiness quality is a phenological quality that represents the ability of grapevine tissues to survive exposure to freezing temperatures without lethal damage, characterizing the dynamic cold tolerance that develops through autumn acclimation, reaches maximum during mid-winter, and declines through spring deacclimation, with hardiness levels determining the minimum temperatures that buds, canes, and other tissues can endure during the dormant period. Within the BFO framework, this quality addresses the freeze tolerance dimension of winter survival, distinguishing the capacity to survive cold from the dormancy status itself while recognizing that cold hardiness and dormancy are related but distinct—dormancy determines growth potential while hardiness determines survival capacity, and both must be appropriately coordinated with winter conditions for successful perennial cultivation. The ontological significance lies in cold hardiness determining the geographic limits of grape cultivation and the winter survival probability in regions with severe cold, where the match between tissue hardiness levels and minimum temperatures experienced determines whether vines survive winters intact, suffer partial damage requiring recovery, or experience lethal injury ending productive life. The physiological basis for cold hardiness involves cellular and biochemical changes during acclimation including membrane lipid modifications (increasing unsaturation for membrane fluidity at low temperatures), accumulation of cryoprotectant compounds (sugars, proline, dehydrins, and other protective molecules), controlled dehydration (reducing freezable water content), and gene expression changes (cold-responsive genes encoding protective proteins), with these changes progressively increasing freeze tolerance as acclimation proceeds. The tissue-specific variation in cold hardiness shows substantial differences within the vine, with dormant buds typically the most sensitive tissues (primary buds more sensitive than secondary buds), while phloem, xylem, and cambium show different hardiness levels that affect the type and severity of winter injury from cold events of different intensities. Understanding cold hardiness quality enables assessment of winter survival probability, selection of cultivars and sites appropriate for local winter severity, and management practices that promote hardiness development and retention through the vulnerable dormant period.
Cold_Hardiness_Quality serves as the parent class for hardiness dynamics (Maximum, Development Rate, Deacclimation, Reacclimation). This quality addresses freeze tolerance determining winter survival and geographic cultivation limits. The class supports site-cultivar matching and winter risk assessment through cold hardiness characterization.
Frosthärte
Frostresistenz
Kältetoleranz
Winterhärte
cold tolerance
freeze tolerance
frost resistance
winter hardiness
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10(1-2): 201-235 Mills LJ et al. (2006) Timing of low temperature exposure affects cold hardiness and survival of dormant grapevine buds. American Journal of Enology and Viticulture 57(2): 232-239
Cold Hardiness Quality
3309 Couderc (V. riparia × V. rupestris) demonstrates high cold hardiness with root LT₅₀ of -12°C to -14°C, widely used in continental climates of Burgundy, Champagne, and Germany where winter soil temperatures occasionally approach -10°C in shallow or snow-free conditions. Riparia Gloire de Montpellier (V. riparia) shows excellent root cold hardiness (LT₅₀ -13°C to -15°C) reflecting V. riparia's natural range extending to southern Canada, supporting Pinot Noir in the cold-winter regions of Oregon's Willamette Valley and New Zealand's Central Otago. 101-14 Millardet et de Grasset (V. riparia × V. rupestris) combines cold hardiness with low vigor, used in the continental climates of Austria, Hungary, and Switzerland where winter soil temperatures require hardy root systems.
An abiotic stress-tolerant rootstock role that inheres in a grafted rootstock by virtue of its capacity to survive severe winter cold and transmit cold hardiness characteristics that benefit the grafted vine composite. Cold-hardy rootstocks tolerate midwinter temperatures of -20°C to -30°C or below without lethal root or trunk damage, through mechanisms including timely cold acclimation in autumn (accumulation of cryoprotective sugars, dehydrins, and late embryogenesis abundant proteins), deep supercooling of xylem ray parenchyma, controlled extracellular ice formation that prevents intracellular freezing, and sufficient carbohydrate reserves to support spring recovery from partial freeze injury. Root cold hardiness is particularly critical because roots are less cold-hardy than above-ground tissues (root LT₅₀ typically -8°C to -15°C versus -20°C to -30°C for mature canes) and are protected only by soil insulation. Cold-hardy rootstocks may also enhance scion cold tolerance through effects on dormancy timing, carbohydrate partitioning, and hormonal signaling.
Cold_Hardy_Rootstock_Role addresses winter survival in continental and cool-climate viticulture where inadequate root cold hardiness can cause vine death or chronic decline. V. riparia parentage contributes the highest levels of cold hardiness, reflecting this species' natural distribution across cold-winter regions of northeastern North America. The V. berlandieri × V. rupestris rootstocks (110R, 1103P, 140Ru) dominant in Mediterranean climates have lower cold hardiness and are unsuitable for continental regions with severe winters. Climate change is paradoxically increasing cold damage risk in some regions through reduced snow cover insulation and more variable winter temperatures that disrupt cold acclimation.
cold hardy rootstock
cold tolerant rootstock
frost tolerant rootstock
frostharte Unterlage
kälteverträgliche Unterlage
winter hardy rootstock
winterharte Unterlage
Striegler RK, Howell GS (1991) The influence of rootstock on the cold hardiness of Seyval grapevines, Vitis 30: 1-11 Pierquet P et al. (1977) Grape hardiness and survival in Minnesota, Fruit Varieties Journal 31: 105-110 Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10: 201-235 Mills LJ et al. (2006) Timing of low temperature exposure affects cold hardiness of wine grape cultivars, American Journal of Enology and Viticulture 57: 229-235
Cold Hardy Rootstock Role
Long-term assessment of Cold_Recovery_Disposition at the University of Minnesota tracked vineyard recovery following the severe 2013-14 polar vortex winter (minimum -34°C) that caused extensive damage across cultivars. Annual measurements over 5 post-damage seasons documented bud survival, trunk condition, yield recovery, vine vigor, and mortality across 18 cultivars in replicated trials. Cold_Recovery_Disposition showed strong cultivar differentiation. 'Frontenac' exhibited exceptional recovery despite 85% primary bud kill, activating secondary buds to produce 45% of typical yield in year 1, achieving 90% yield recovery by year 2, and showing no detectable long-term vigor reduction or elevated mortality. 'La Crescent' showed slower recovery with 25% yield in year 1 (reflecting lower secondary bud fruitfulness), 70% by year 2, and 95% by year 3, with no trunk damage requiring structural recovery. V. vinifera 'Marquette' (grafted) suffered both bud and partial trunk damage (40% of vines with crown injury), requiring more complex recovery. Vines with crown injury produced only 15% yield in year 1 from basal buds, required trunk retraining from suckers, and did not achieve full yield recovery until year 4. Mortality over 5 years was 2% ('Frontenac'), 5% ('La Crescent'), and 18% ('Marquette', concentrated in crown-damaged vines). These findings demonstrate that Cold_Recovery_Disposition determines both short-term economic impact and long-term vineyard sustainability following severe winter events.
A Cold_Recovery_Disposition is a Cold_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to regenerate functional tissues, restore vine architecture, and resume productive growth following winter cold damage to buds, canes, trunks, or roots. This disposition represents the restorative component of cold adaptation, determining how rapidly and completely vines recover from cold injury events rather than preventing initial damage. Cold recovery encompasses multiple processes including activation of surviving backup buds (secondary, tertiary, latent, or basal buds) to replace killed shoots, regeneration of vascular tissues through cambial activity to repair damaged canes and trunks, wound compartmentalization to limit secondary infection of cold-damaged tissues, and restoration of carbohydrate reserves depleted by regrowth demands. The timeframe for recovery varies with damage severity, ranging from single-season crop recovery through secondary bud growth (moderate damage) to multi-year structural rebuilding when trunks are killed (severe damage). Cold_Recovery_Disposition varies among genotypes based on the abundance and fruitfulness of backup buds, the vigor of regenerative growth, the efficiency of wound healing, and the capacity to partition resources between regrowth and reserve replenishment. Strong recovery capacity reduces the long-term economic impact of winter damage events, enabling faster return to full production and avoiding cumulative decline from repeated partial damage.
Cold_Recovery_Disposition is_a Cold_Tolerance_Disposition. Recovery capacity determines long-term consequences of cold damage events, complementing hardiness traits that prevent initial damage. Child classes include Secondary_Bud_Activation_Disposition (backup bud mobilization) and Trunk_Regeneration_Disposition (structural recovery). Recovery timeframe ranges from single season (bud damage only) to multiple years (trunk/structural damage). Strong recovery capacity reduces cumulative impact of repeated cold events that individually cause incomplete damage.
Frostschadens-Wiederherstellung
Kälteschadens-Erholungs-Disposition
Winterschadens-Regeneration
cold damage restoration
freeze injury resilience
winter injury recovery
Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235 Dami IE et al. (2012) Evaluation of winter injury and subsequent performance of grapevine cultivars in Ohio, Acta Horticulturae 932:235-241 Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection, Michigan State University Extension Bulletin E-2930
Cold Recovery Disposition
Cold recovery quality comparison following equivalent freeze damage in Finger Lakes Riesling versus Chardonnay documented cultivar differences in recovery capacity with implications for rehabilitation management. Both cultivars experienced 65% primary bud mortality from a January freeze event, but subsequent cold recovery quality diverged substantially between the varieties. Riesling demonstrated excellent cold recovery quality with secondary bud activation occurring uniformly across surviving cane positions, secondary shoot fruitfulness averaging 75% of normal primary shoot fruitfulness, and yield recovery to 65% of normal by the first season post-freeze. Chardonnay showed inferior cold recovery quality with secondary bud activation delayed and less uniform, secondary shoot fruitfulness only 45% of normal primary shoots, and yield recovery reaching only 40% of normal despite equivalent initial damage. The cold recovery quality differential was attributed to Riesling's naturally higher secondary bud fruitfulness combined with more robust carbohydrate reserve status entering winter, providing greater resources to support regrowth. Trunk recovery assessment showed that both cultivars effectively compartmentalized freeze damage in canes and cordons, but Riesling showed faster wound healing and less subsequent canker development, contributing to better long-term cold recovery quality. The assessment established that cold recovery quality varies significantly among cultivars and should inform variety selection for cold-climate sites where periodic freeze damage is expected, favoring cultivars with strong recovery capacity over those with marginally greater hardiness but poor recovery following damage.
Cold recovery quality is a cold stress indicator quality that inheres in a grapevine organism and represents the capacity to restore productive function following freezing injury through the activation of surviving meristems, the regeneration of damaged tissues, and the physiological adaptation to the altered vine structure resulting from cold damage, with recovery capacity determining whether and how quickly freeze-damaged vines can return to productive status and the management interventions required to support effective recovery. This quality captures the resilience dimension of cold stress response, recognizing that freeze damage severity alone does not determine outcomes since vines with equivalent initial damage may show dramatically different recovery trajectories depending on the survival of backup meristems, the effectiveness of repair processes, and the management practices employed to support regrowth. The ontological significance of cold recovery quality lies in its role determining the long-term vineyard consequences of freeze events, with strong recovery capacity enabling damaged vines to return to productivity within one or two seasons while poor recovery may result in extended yield losses or vine mortality requiring replanting despite initial survival of the freeze event itself. The biological basis of cold recovery encompasses the activation of secondary and tertiary buds providing alternative shoot growth when primary buds are killed, the regenerative capacity of surviving cambium to produce new vascular tissues replacing damaged phloem and xylem, the mobilization of carbohydrate reserves from permanent vine structures to support regrowth, and the wound healing processes that isolate damaged tissue and prevent pathogen spread. The genetic variation in cold recovery quality among cultivars reflects differences in secondary bud fruitfulness affecting crop recovery, in cambial vigor affecting tissue regeneration, and in carbohydrate reserve accumulation patterns affecting the resources available to support regrowth. The management influences on cold recovery include pruning adjustments that direct growth to surviving buds and optimize the allocation of limited resources, irrigation and nutrition management that supports vigorous regrowth without exceeding the vine's capacity, and pest and disease management that prevents secondary damage to already-stressed vines.
Cold_Recovery_Quality is_a Cold_Stress_Indicator_Quality, serving as parent class for Secondary_Bud_Activation_Quality, Trunk_Recovery_Capacity_Quality, and Crown_Gall_Susceptibility_Quality which address specific recovery mechanisms and limitations. This class addresses the overall capacity for productive function restoration following freeze damage. The combination of backup bud availability, tissue regeneration, and reserve mobilization determines recovery success. Cultivar variation in recovery capacity (Riesling > Chardonnay) should inform variety selection for cold-climate sites. Management interventions (pruning, nutrition) significantly influence recovery outcomes.
Frostschaden-Erholung
Kälteerholungs-Qualität
Kälteschaden-Wiederherstellung
Winterschaden-Rehabilitation
cold damage rehabilitation
freeze recovery
frost recovery capacity
Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection. Michigan State University Extension Bulletin E2930 Pool RM et al. (1990) Assessing and predicting cold injury to grapevines. Proceedings of the New York Wine Industry Workshop, pp. 17-25 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235
Cold Recovery Quality
Cold stress indicator quality assessment following a late spring frost event (-4°C for 3 hours) in Burgundy evaluated the extent of damage to Pinot Noir vineyards at varying phenological stages and topographic positions. Visual cold stress indicator quality assessment 48 hours post-frost identified a gradient of damage severity correlated with both phenological advancement and cold air drainage patterns, with valley-floor sites showing 85% primary shoot damage while upper-slope sites showed only 25% damage despite equivalent air temperatures, reflecting cold air pooling effects on exposure duration. Physiological cold stress indicator quality measurements including electrolyte leakage from affected tissues confirmed visual assessments, with severely damaged shoots showing greater than 70% leakage indicating lethal membrane damage while mildly affected tissues showed 25-40% leakage consistent with recoverable injury. The cold stress indicator quality evaluation guided recovery management decisions, with severely damaged blocks requiring shoot thinning to redirect growth to secondary buds while lightly damaged blocks were managed to preserve partially injured primary shoots that would recover to produce reduced but viable crops.
Cold stress indicator quality is a stress response quality that inheres in a grapevine organism and represents the ensemble of physiological, biochemical, and morphological markers that signal the presence and severity of low temperature stress, encompassing both the damage symptoms resulting from freezing or chilling injury and the tolerance mechanisms that determine susceptibility to cold-induced damage across the range of low temperature exposures encountered in viticultural environments from mild chilling through severe winter freeze events. This quality captures the grapevine response to temperatures below the optimal range for growth and function, recognizing that cold stress manifests differently depending on the temperature regime, the phenological stage of the vine, the acclimation status of tissues, and the specific organs affected, with consequences ranging from temporary growth inhibition through tissue damage to complete vine mortality under the most severe conditions. The ontological significance of cold stress indicator quality lies in its role enabling assessment of vine cold hardiness status and damage extent following cold events, supporting decisions about site selection, cultivar choice, protective management practices, and post-damage recovery strategies that collectively determine the viability of grape production in cold-climate regions where winter survival and spring frost avoidance represent primary production constraints. The biological basis of cold stress indicators encompasses the cellular and tissue-level responses to low temperature including membrane phase transitions, ice crystal formation and propagation, cellular dehydration from extracellular freezing, and the metabolic disruptions that collectively cause cold injury, combined with the protective mechanisms including cold acclimation, cryoprotectant accumulation, and supercooling capacity that determine tissue survival at specific temperature exposures. The seasonal dynamics of cold hardiness create varying vulnerability windows, with dormant tissues showing maximum hardiness in midwinter following full acclimation, spring-deacclimated tissues showing high vulnerability to late frosts, and actively growing tissues lacking any freezing tolerance during the growing season. The interaction between cold stress and other environmental factors including prior temperature history affecting acclimation status, soil moisture affecting tissue hydration, and wind affecting heat loss rates creates complex determinants of cold injury outcomes that cold stress indicator quality assessment must integrate.
Cold_Stress_Indicator_Quality is_a Stress_Response_Quality, serving as parent class for Frost_Damage_Severity_Quality and Tissue_Freezing_Tolerance_Quality which address damage symptoms and tolerance mechanisms respectively. This class addresses the overall assessment of cold stress while child hierarchies capture particular dimensions. Distinguished from Heat_Stress_Indicator_Quality by the focus on low rather than high temperature stress. The seasonal dynamics of hardiness create phenology-dependent vulnerability patterns. Critical for cold-climate viticulture where winter survival and frost avoidance determine production viability.
Froststress-Reaktion
Kältehärte-Indikatoren
Kältestress-Indikatoren
Tieftemperaturstress-Marker
cold damage markers
cold hardiness indicators
freezing stress response
low temperature stress indicators
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 508-545 Mills LJ et al. (2006) Timing of low temperature exposure affects cold hardiness and cold deacclimation kinetics in grape. American Journal of Enology and Viticulture 57:20-26
Cold Stress Indicator Quality
Long-term monitoring of Cold_Tolerance_Disposition at the University of Minnesota characterized seasonal cold hardiness dynamics across 25 cultivars over 8 winters using differential thermal analysis (DTA) to determine bud LT50 values (temperature killing 50% of buds). Measurements at monthly intervals from November through March revealed dramatic cultivar differences in both maximum hardiness and seasonal patterns. V. vinifera 'Pinot Noir' showed marginal Cold_Tolerance_Disposition with maximum hardiness of -21°C reached in mid-January, rapid deacclimation beginning in late February, and frequent economic crop losses (5 of 8 winters with >50% primary bud mortality). The Minnesota-bred hybrid 'Frontenac' exhibited exceptional Cold_Tolerance_Disposition with maximum hardiness of -35°C, extended hardiness maintenance through February, and slower spring deacclimation, sustaining zero bud mortality across all 8 winters despite exposure to -32°C ambient temperature in the most severe winter. 'Marquette' showed intermediate but commercially adequate cold tolerance (LT50 -34°C) with an additional advantage of rapid reacclimation following midwinter thaws, recovering 4°C of hardiness within 72 hours after a January warming event that caused permanent hardiness loss in less adaptive cultivars. These findings demonstrate that Cold_Tolerance_Disposition encompasses not only absolute minimum survival temperature but also acclimation kinetics, hardiness maintenance, and reacclimation capacity.
A Cold_Tolerance_Disposition is an Abiotic_Stress_Tolerance_Disposition that inheres in a Vitis plant and is realized through physiological and biochemical mechanisms enabling survival of freezing temperatures that would otherwise cause lethal ice formation within tissues, membrane damage, and cell death. This disposition represents one of the most critical abiotic stress tolerances for grapevine, determining the geographic limits of cultivation and the economic sustainability of viticulture in continental and cold-climate regions. Cold tolerance in woody perennials involves a dynamic process of cold acclimation in autumn (triggered by shortening photoperiod and declining temperatures), maximum hardiness during midwinter dormancy, and deacclimation in spring as temperatures rise and growth resumes. The mechanisms underlying cold tolerance include controlled extracellular ice formation that draws water from cells (freeze tolerance), deep supercooling that avoids ice formation in critical tissues like xylem ray parenchyma and bud primordia, accumulation of cryoprotective compounds (sugars, proline, dehydrins, antifreeze proteins), membrane lipid remodeling increasing fluidity at low temperatures, and dormancy regulation preventing premature deacclimation. Cold tolerance varies dramatically among Vitis species, with V. vinifera generally least hardy (midwinter LT50 typically -15°C to -25°C), while V. riparia, V. amurensis, and cold-adapted hybrids tolerate -30°C to -40°C. Cold damage remains the primary limitation to V. vinifera cultivation in regions with severe winters, driving both rootstock selection and extensive breeding programs for cold-hardy cultivars.
Cold_Tolerance_Disposition is_a Abiotic_Stress_Tolerance_Disposition. This class encompasses all aspects of cold/freeze tolerance, with child classes addressing specific components including Midwinter_Hardiness_Disposition (deep dormancy cold tolerance), Spring_Frost_Tolerance_Disposition (post-budbreak frost survival), and Fall_Frost_Tolerance_Disposition (pre-dormancy cold events). Cold tolerance is the primary factor limiting V. vinifera cultivation in continental climates and drives extensive cold-hardy cultivar breeding programs. The dynamic nature of cold tolerance through the dormant season distinguishes it from static traits, requiring assessment of acclimation, maintenance, and deacclimation phases.
Frosttoleranz
Kältetoleranz-Disposition
Winterhärte-Kapazität
low temperature tolerance
Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235 Mills LJ et al. (2006) Timing of low temperature exposure affects cold hardiness and budbreak in Vitis vinifera, American Journal of Enology and Viticulture 57:528-535 Keller M (2020) The Science of Grapevines, 3rd Edition, Academic Press
Cold Tolerance Disposition
Color appearance assessment in blind tasting of red wine reveals deep ruby-purple hue with violet rim suggesting youth, high color intensity indicating thick-skinned variety and warm climate, and saturated appearance without browning confirming recent vintage, enabling preliminary identification as young Malbec or Syrah from warm region before aromatic evaluation, with color cues creating expectation of concentrated dark fruit and firm tannins that frames interpretation of subsequent sensory impressions.
Color appearance quality is a visual appearance quality that represents the chromatic properties of wine arising from light absorption and reflection by pigment compounds, encompassing hue, intensity, and saturation attributes that provide diagnostic information about grape variety, wine age, production method, and overall quality. Within the BFO framework, this quality addresses the chromatic dimension of visual assessment, distinguishing color-related properties from clarity and viscosity while recognizing that wine color provides immediate variety and age cues that frame subsequent sensory evaluation. The ontological significance lies in color serving as the most variety-specific and age-sensitive visual attribute, where red wine color immediately distinguishes thick-skinned from thin-skinned varieties and indicates developmental stage from youthful purple through mature garnet to aged tawny, while white wine color progression from pale straw through gold to amber signals oxidative evolution and age. The physical basis for wine color involves selective light absorption by chromophore compounds, with anthocyanins providing red-blue-purple colors in young red wines, tannin-anthocyanin polymers creating brick and tawny tones in aged reds, and carotenoid derivatives, flavonols, and oxidation products determining white wine coloration from pale green-yellow through deep gold. The compositional determinants of color include anthocyanin concentration and profile (variety-specific), pH (affecting anthocyanin equilibrium and color expression), copigmentation (enhancing color through molecular stacking), tannin-anthocyanin polymerization (shifting color toward orange with age), and oxidation state (browning in whites, tawny development in reds). The evaluation protocol for color assessment involves examination against white background with appropriate lighting, tilted glass observation for rim-to-core gradient, and standardized terminology for hue and intensity description, with color providing the first variety and age hypothesis before aromatic assessment begins. Understanding color appearance quality enables variety identification from color profile, age estimation from color evolution, production method inference from color characteristics, and quality assessment based on color appropriateness for wine type and age.
Color_Appearance_Quality serves as the parent class for color subcategories (Intensity, Hue, Saturation, Rim Variation), establishing the framework for chromatic assessment. This quality provides variety, age, and production information more immediately than any other visual attribute. The class addresses the pigment-related dimension of visual appearance, with color assessment providing essential diagnostic cues that guide subsequent evaluation.
Farbcharakter
Farberscheinung
Weinfarbe
chromatic quality
chromatische Qualität
color characteristics
hue and tint
wine color
Somers TC, Evans ME (1977) Spectral evaluation of young red wines: anthocyanin equilibria, total phenolics, free and molecular SO₂, chemical age. Journal of the Science of Food and Agriculture 28(3): 279-287 Boulton R (2001) The copigmentation of anthocyanins and its role in the color of red wine: a critical review. American Journal of Enology and Viticulture 52(2): 67-87
Color Appearance Quality
In a Chianti Classico Sangiovese vineyard, color break initiation was documented in mid-September Year N+1 (BBCH 83) when 25-35% of berry surface showed red-purple patches concentrated on sun-exposed southwestern aspects, skin anthocyanins measured 95 mg/kg (from 42 mg/kg at first pigment 3 days earlier), chlorophyll declined to 65 μg/g (from 145 μg/g green), berry firmness dropped 12% to 2.3 N, and sugar reached 11.5° Brix under typical Tuscan autumn conditions (daily high 25-28°C, night 14-17°C).
Color_Break_Initiation represents the developmental transition when initial scattered pigmented patches on berry surfaces begin expanding and coalescing creating the characteristic "breaking" of uniform green coloration into mixed green-red patterns that define early visible veraison. This process occurs 1-3 days following first pigment deposition (BBCH 83), when anthocyanin concentrations in pioneering pigmented regions reach 60-120 mg/kg skin fresh weight creating sufficient optical density to overcome chlorophyll masking, and marks the beginning of rapid visible color transformation from predominantly green to variegated appearance preceding complete berry pigmentation. The "break" in color uniformity results from spatial heterogeneity in anthocyanin synthesis and accumulation: initial pigmented patches typically measure 2-5 mm diameter representing clusters of 50-200 epidermal and sub-epidermal cells with coordinated anthocyanin accumulation, these patches appear preferentially on sun-exposed berry surfaces where light and temperature signals maximize anthocyanin induction creating asymmetric coloring with southwestern aspects (northern hemisphere) showing earliest and most intense patches, and within individual berries pigmentation often initiates near stylar end spreading toward pedicel attachment reflecting apical-to-basal developmental gradients in ripening signaling. Cellular basis of color break involves: expansion of initially pigmented cell clusters through recruitment of adjacent cells into anthocyanin synthesis with radial spreading patterns from initial foci, intercellular signaling possibly through diffusible factors or cell-to-cell contact enabling coordinated pigmentation responses creating coherent colored patches rather than isolated pigmented cells, and continued chlorophyll degradation in pigmented regions (chlorophyll declining from 100-200 μg/g to 40-80 μg/g) reducing green color masking allowing red-purple anthocyanins to dominate visual appearance. Anthocyanin accumulation during color break shows: total skin anthocyanins increasing from first deposition levels 30-60 mg/kg to color break detection 80-150 mg/kg representing 2-3 fold increase within 2-4 days, accumulation rates accelerating from initial synthesis 10-30 mg/kg per day to color break 25-60 mg/kg per day as biosynthetic machinery reaches 60-80% of maximum capacity, and malvidin-3-glucoside maintaining dominance at 55-70% of total while acylated derivatives increase to 10-20% enhancing color stability and intensity. Gene expression during color break shows: anthocyanin biosynthetic genes reach 70-90% of eventual maximum expression with UFGT transcripts 120-180 fold above green berry baseline, regulatory MYB genes (VvMYBA1/A2) achieve 80-120 fold induction coordinating pathway activation, chlorophyll degradation genes including pheophytinase and pheophorbide a oxygenase show 10-30 fold elevation facilitating green color removal, and carotenoid biosynthetic genes may increase slightly contributing yellow-orange background visible in white cultivars but masked by anthocyanins in reds. Optical properties during color break inclu pigmented patches show characteristic absorbance maxima 520-540 nm for anthocyanins with optical density 0.8-1.5 units at peak wavelength, reflectance spectra shift from predominantly green reflection (500-600 nm) in unpigmented regions to red-purple reflection (600-650 nm with secondary blue reflection 400-450 nm) in colored patches creating visually striking color contrast, and overall berry appearance transitions from solid green (pre-break) to mottled green-red (color break) to predominantly red with green remnants (progressive coloring) as pigmented area expands from 20-40% at break initiation to >80% at break completion. Environmental modulation of color break timing includes: optimal berry temperature 24-30°C accelerates color break producing visible patterns 2-4 days post-first pigment, cooler temperatures 18-24°C delay break by 1-3 days through slower anthocyanin accumulation despite potentially more intense final color, heat stress >33°C can severely delay or prevent normal color break patterns producing pale poorly-colored fruit through anthocyanin biosynthesis inhibition, and diurnal temperature variation >15°C day-night difference often enhances color break intensity creating more distinct pigmented patches through optimized synthesis-stabilization balance. Light effects during color break show: UV-B exposure enhances patch formation and intensity producing 30-50% higher anthocyanin levels in exposed versus shaded berry regions, visible light particularly blue wavelengths promotes color break progression through photoreceptor signaling, excessive direct sun combined with high temperature can inhibit break progression through heat stress overriding light benefits, and moderate canopy shading delaying color break 3-5 days while potentially improving ultimate color intensity through temperature moderation demonstrates complex light-temperature interactions. Water stress influences on color break inclu moderate deficit (stem ψ -0.9 to -1.3 MPa) can enhance color break intensity and advance timing 2-3 days through elevated ABA signaling and berry concentration effects, severe stress (<-1.5 MPa) disrupts normal break patterns producing irregular patchy coloring through metabolic impairment, while excess irrigation delays color break 2-4 days and reduces intensity through berry dilution and suppressed ABA-dependent anthocyanin gene activation. Cultivar variation in color break characteristics reflects genetic differences: rapid-breaking varieties like Pinot Noir show abrupt color break with 20-30% berry surface pigmented within 24-48 hours creating dramatic visual transformation, moderate cultivars like Merlot and Syrah display gradual break over 3-5 days with progressive patch expansion, while slow-breaking varieties like Cabernet Sauvignon and Nebbiolo show extended break period 5-8 days with initially subtle color changes requiring careful observation for detection. Within-cluster heterogeneity during color break creates: berries at different developmental stages showing simultaneous presence of completely green (lag phase), color breaking (early veraison), and substantially colored (mid-veraison) fruit reflecting inherent developmental asynchrony, exposed berries typically 2-4 days ahead of shaded berries in color break timing creating spatial patterns of coloring progression, and cluster shoulders often breaking color before cluster tips possibly reflecting vascular supply differences or developmental gradients. Viticultural significance of color break timing includes: visual assessment of color break provides practical veraison monitoring enabling estimation of harvest timing (typically 40-55 days post-color break depending on heat accumulation and cultivar), within-vineyard variation in color break timing reveals maturity heterogeneity informing decisions about selective harvest or acceptance of compositional variation, and irrigation management often adjusts at color break initiating deficit strategies or terminating water applications as berry growth slows. Physiological markers accompanying color break inclu berry firmness declining 10-20% from green berry values as softening initiates often concurrent with color break, sugar accumulation accelerating from <1.0% Brix per week to 1.5-3.0% per week, acid levels beginning decline particularly malate showing 5-15% reduction through respiratory consumption, and seed color transitioning from green to yellow-brown indicating coordinated ripening program activation across berry tissues. Monitoring color break progression informs management: photographic documentation enables quantitative tracking of color development through image analysis measuring colored versus green surface area, spectrophotometric measurement of skin extracts provides anthocyanin quantification correlating with visual break assessment, and regular cluster sampling across vineyard blocks identifies spatial variation in break timing revealing terroir effects or management impacts requiring attention.
Color_Break_Initiation is distinguished from First_Pigment_Deposition by representing visible pattern formation rather than initial sequestration, from Progressive_Color_Development by showing early mixed green-red rather than advancing pigmentation, and from Complete_Color_Change by exhibiting partial (20-40%) rather than full coverage. Anthocyanin range at break: 60-120 mg/kg skin. Duration from first pigment to break: fast cultivars 1-2 days, moderate 2-4 days, slow 3-6 days. Temperature optimum 24-30°C berry surface. Color_Break_Initiation provides practical visual marker for veraison monitoring and harvest forecasting.
Farb-Gleichförmigkeits-Bruch
Grün-Farb-Störung
Pigmentierungs-Muster-Entstehung
color uniformity breaking
gescheckter Färbungs-Beginn
green color disruption
pigmentation pattern emergence
variegated coloring onset
Keller M (2020) The Science of Grapevines 3rd Ed pages 233-234 color development patterns Mori K, Sugaya S, Gemma H (2005) Decreased anthocyanin biosynthesis in grape berries grown under elevated night temperature condition Sci Hort 105:319-330 environmental effects on color break Downey MO, Harvey JS, Robinson SP (2004) The effect of bunch shading on berry development and flavonoid accumulation in Shiraz grapes Australian J Grape Wine Res 10:55-73 spatial patterns of color break.
Color Break Initiation
In a Burgundian Pinot Noir vineyard near Gevrey-Chambertin, color change onset was observed in mid-August Year N+1 (BBCH 83, 9 weeks post-flowering) when first red patches appeared on sun-exposed berry surfaces coinciding with ABA levels rising from 120 ng/g to 850 ng/g, UFGT gene expression increasing 95-fold, and initial anthocyanin accumulation reaching 45 mg/kg skin from undetectable pre-veraison levels, under conditions of moderate temperature (daily high 24-27°C, night 14-16°C) and partial cluster exposure.
Color_Change_Onset represents the developmental transition during veraison when red grape cultivars initiate visible pigmentation through de novo anthocyanin biosynthesis, marking one of the most conspicuous phenotypic changes in berry ripening and serving as a universal marker for veraison timing in colored varieties. This process occurs during early-to-mid veraison (BBCH 83-84), typically 1-3 days before or concurrent with initial berry softening depending on cultivar and environmental conditions, and transforms berry skin color from uniform green (dominated by chlorophyll a and b absorbing red and blue light, reflecting green) to variegated red-green patches that progressively coalesce into complete berry pigmentation over 5-14 days cultivar-dependent. Color change onset is triggered by complex hormonal and developmental signals: abscisic acid levels spike from pre-veraison baseline 50-150 ng/g to veraison peak 500-2000 ng/g activating transcription factors VvMYBA1, VvMYBA2, and related regulators that coordinately upregulate anthocyanin biosynthetic genes 50-200 fold, sugar accumulation creates hexose signaling through hexokinase pathways that synergize with ABA to enhance anthocyanin gene expression, and declining gibberellin and auxin levels remove growth-related suppression of secondary metabolism allowing anthocyanin pathway activation. The onset process initiates anthocyanin biosynthesis through coordinated enzyme induction: phenylalanine ammonia lyase (PAL) activity increases 10-40 fold catalyzing the first committed step converting phenylalanine to cinnamic acid entering phenylpropanoid metabolism, chalcone synthase (CHS) shows 20-60 fold elevation producing naringenin chalcone the first flavonoid intermediate, flavanone 3-hydroxylase (F3H), dihydroflavonol 4-reductase (DFR), and leucoanthocyanidin dioxygenase (LDOX) activities increase sequentially producing leucoanthocyanidins, and UDP-glucose: flavonoid 3-O-glucosyltransferase (UFGT) achieves 50-150 fold induction catalyzing the final step producing stable anthocyanin-3-glucosides that accumulate in vacuoles creating visible red-blue pigmentation. Initial anthocyanin synthesis produces predominantly malvidin-3-glucoside (50-70% of total anthocyanins in most cultivars) along with cyanidin, peonidin, delphinidin, and petunidin derivatives whose relative proportions define cultivar-specific color characteristics: Pinot Noir shows high cyanidin content producing bright cherry-red color, Syrah exhibits elevated delphinidin creating purple-blue hues, Merlot displays balanced anthocyanin profiles producing deep red coloration, while Cabernet Sauvignon accumulates high malvidin with moderate delphinidin yielding characteristic purple-red color. Color change onset timing varies substantially among cultivars reflecting genetic differences in regulatory control and biosynthetic capacity: early-coloring varieties (Pinot Noir, Gamay, Grenache) initiate color change 8-10 weeks post-flowering with rapid progression to full coloring in 3-5 days, mid-season cultivars (Merlot, Syrah, Sangiovese) begin 9-12 weeks post-flowering completing coloring in 5-8 days, while late-coloring varieties (Cabernet Sauvignon, Nebbiolo, Mourvèdre) initiate 11-14 weeks post-flowering requiring 8-14 days for complete pigmentation reflecting slower anthocyanin accumulation rates. Environmental factors profoundly influence color change onset timing and intensity: temperature effects are complex with optimal ranges 20-28°C supporting maximum anthocyanin synthesis enzyme activities, cooler temperatures 15-20°C can enhance color through increased anthocyanin stability and reduced degradation despite slower synthesis rates, while heat stress >32-35°C inhibits anthocyanin biosynthesis through enzyme denaturation and gene expression suppression producing pale poorly-colored fruit, and dramatic diurnal temperature variation (>15°C difference day-night) often enhances coloring through combined warm-day synthesis and cool-night stabilization. Light quality and intensity significantly affect color development: ultraviolet-B radiation (280-315 nm) induces anthocyanin biosynthetic genes particularly CHS and UFGT increasing pigment accumulation 30-60% in exposed versus shaded fruit, visible red and blue wavelengths (400-500 nm, 600-700 nm) absorbed by photoreceptors enhance anthocyanin synthesis, while excessive direct sunlight combined with high temperature can inhibit coloring through heat stress effects, and moderate canopy shading reducing bunch temperature by 3-5°C while maintaining some light penetration often optimizes color development balancing synthesis induction with heat stress avoidance. Water relations during color change onset influence pigmentation intensity: moderate water deficit (stem water potential -0.9 to -1.3 MPa) can enhance anthocyanin accumulation 20-40% through stress signaling activating ABA pathways and concentrating berry solutes, severe deficit (35°C can delay onset through heat inhibition, while completely shaded clusters may show 30-50% reduced anthocyanin accumulation and delayed onset by 4-7 days demonstrating light requirement for optimal pigmentation despite potential temperature benefits of shade. Molecular regulation involves complex transcriptional control: VvMYBA1 and VvMYBA2 R2R3-MYB transcription factors act as master regulators directly activating UFGT and other late biosynthetic genes, these MYBs interact with bHLH and WD40 proteins forming MBW complexes that coordinately regulate anthocyanin pathway genes, ABA-responsive element binding factors (ABFs) mediate ABA signaling to anthocyanin genes, while other transcription factors including VvMYB5a, VvMYB5b, and WRKY proteins modulate pathway regulation creating cultivar-specific expression patterns. Physiological consequences of color change onset extend beyond visible pigmentation: anthocyanins function as antioxidants protecting berries from photo-oxidative damage particularly important as chlorophyll degrades, UV-screening properties protect cellular contents from radiation damage, and accumulation of phenolic intermediates and products creates chemical changes affecting berry flavor, astringency, and wine quality characteristics. White and rosé cultivars show distinct color change patterns: white varieties (Chardonnay, Sauvignon Blanc, Riesling) undergo minimal color change with green-to-golden transitions reflecting chlorophyll degradation and carotenoid exposure rather than anthocyanin synthesis (anthocyanins <5-10 mg/kg versus 500-3000 mg/kg in reds), while teinturier varieties with red flesh (Alicante Bouschet) accumulate anthocyanins in mesocarp as well as skin creating intense coloring useful for blending. Viticultural implications of color change onset inclu visible color provides easily monitored veraison marker for developmental tracking and harvest forecasting, uneven color development within clusters or vineyards indicates maturity heterogeneity requiring selective harvest or acceptance of variable grape composition, and color intensity at harvest substantially affects wine color, phenolic content, and aging potential making color development monitoring essential for quality prediction and optimization.
Color_Change_Onset is distinguished from later color development phases by representing initiation rather than progression, occurs typically 1-3 days before or with initial softening in red cultivars, and is absent or minimal in white varieties. Duration from first visible color to complete coloring: early cultivars 3-5 days, mid-season 5-8 days, late varieties 8-14 days. Temperature optimum 20-28°C for synthesis. UV-B and moderate light enhance coloring. Color_Change_Onset represents RED CULTIVAR-specific process white cultivars show only chlorophyll degradation without anthocyanin synthesis. Critical veraison marker for developmental tracking.
Anthocyan-Synthese-Beginn
Beeren-Färbungs-Anfang
Pigmentierungs-Start
Veraison-Farb-Initiierung
anthocyanin synthesis onset
berry coloring start
pigmentation commencement
veraison color initiation
Keller M (2020) The Science of Grapevines 3rd Ed pages 232-235 anthocyanin biosynthesis and color development Castellarin SD, Matthews MA, Di Gaspero G, Gambetta GA (2007) Water deficits accelerate ripening and induce changes in gene expression regulating flavonoid biosynthesis in grape berries Planta 227:101-112 environmental regulation Boss PK, Davies C, Robinson SP (1996) Analysis of the expression of anthocyanin pathway genes in developing Vitis vinifera L. cv Shiraz grape berries Plant Physiol 111:1059-1066 biosynthetic gene expression.
Color Change Onset
Color description for Cabernet Sauvignon from Coonawarra (2022 vintage, 2 years post-harvest) records deep purple-crimson core with a narrow violet-purple rim showing minimal evolution, the opaque depth requiring 10 mm path length for spectrophotometric measurement — consistent with the variety's characteristically high anthocyanin content (predominantly acylated malvidin glucosides providing blue-shifted stable color) and the youthful development stage. Color description for aged Riesling Spätlese from the Mosel (2016 vintage, 8 years) documents medium gold with developing amber-honey tones at the rim, a progressive deepening from the pale straw-green of youth through the golden-amber spectrum characteristic of Riesling oxidative aging and the gradual formation of melanoidin-like pigments from norisoprenoid degradation. Color description for Blaufränkisch from Mittelburgenland (2023 vintage) records deep ruby-purple core with vivid magenta-violet rim, reflecting the variety's exceptional anthocyanin concentration and the high proportion of delphinidin and petunidin glucosides that contribute blue-purple hue components.
Color description is a visual description entity that documents the hue, saturation, and intensity of wine color as perceived by human visual assessment, including the core color (the predominant hue observed at the center of the tilted glass), the rim color (the hue at the meniscus edge where the wine thins), the depth or intensity (the degree of color saturation from watery-pale through medium to deep-opaque), and the color evolution markers (purple-blue youthful indicators versus orange-brown aging indicators), described using standardized color vocabulary and optionally quantified by CIELab colorimetric coordinates (L*, a*, b*) or the traditional spectrophotometric color parameters (intensity, hue, modified color parameters), providing the visual chromatic fingerprint that immediately communicates wine type, cultivar character, age, and concentration.
Color description captures the most information-rich visual parameter, as wine color encodes cultivar identity, age, winemaking history, and quality indicators in a single integrated percept. The chromatic vocabulary used in wine assessment follows a standardized progression for red wines from purple (youngest) through ruby, garnet, tawny to brown (oldest), and for white wines from water-white through straw, gold, amber to brown, with rosé wines occupying a separate pale pink to salmon to copper-onion skin spectrum. The chemical basis of wine color is complex and changes continuously throughout the wine's lifetime — young red wines derive color primarily from monomeric anthocyanins (the five glucosides and their acylated derivatives), while aged red wines increasingly rely on polymeric pigments formed through anthocyanin-tannin condensation reactions mediated by acetaldehyde, pyruvic acid, and vinyl-phenol adducts (vitisins, pinotins, portisins). These polymeric pigments are more resistant to SO₂ bleaching and pH variation than monomeric anthocyanins, explaining why well-made red wines maintain color stability as they age despite the progressive loss of free anthocyanins. The rim-versus-core distinction provides a simple but effective age indicator — the thin meniscus at the glass rim reveals the wine's most evolved color (least protected from oxidation), while the deeper core preserves younger color characteristics. CIELab colorimetry (L* = lightness, a* = red-green axis, b* = yellow-blue axis) provides objective quantification that correlates with visual assessment and enables statistical analysis of color differences, with ΔE values >3 CIELab units generally perceptible by trained tasters. Cultivar-specific color signatures arise from genetically determined anthocyanin profiles — Pinot Noir's non-acylated, predominantly peonidin and malvidin monoglucosides produce the characteristically pale, rapidly evolving garnet color, while Cabernet Sauvignon's heavily acylated malvidin-dominated profile generates deep, slowly evolving purple-crimson color.
Farbbeschreibung
Farbbeurteilung
Farbcharakterisierung
chromatic description
chromatische Beschreibung
color assessment
color evaluation
hue description
Somers TC, Evans ME (1977) Spectral evaluation of young red wines. Journal of the Science of Food and Agriculture 28(3): 279-287 Glories Y (1984) La couleur des vins rouges. Connaissance de la Vigne et du Vin 18(4): 253-271 Pérez-Magariño S, González-San José ML (2003) Application of absorbance values used in wineries for estimating CIELab parameters in red wines. Food Chemistry 81(2): 301-306 He F et al. (2012) Anthocyanins and their variation in red wines. Molecules 17(2): 1571-1601
Color Description
In a Russian River Valley Pinot Noir vineyard, color development completion was documented 15 days post-color break in mid-September Year N+1 (BBCH 86) when anthocyanin accumulation plateaued at 1265 mg/kg (±35 mg/kg over 2 weeks), UFGT expression declined to 45% of peak (still 65-fold above green berry), CIE L* stabilized at 33.8 (±2.1 over 3 weeks), berry Brix 23.2° with accumulation <0.6% per week, titratable acidity 6.5 g/L, firmness 1.2 N, under optimal cool coastal conditions (daily high 20-23°C, night 10-13°C, morning fog) enabling extended hang time for flavor development.
Color_Development_Completion represents the developmental endpoint when all color-related processes including anthocyanin biosynthesis, vacuolar sequestration, copigmentation establishment, and chlorophyll degradation reach asymptotic termination transitioning from active developmental programming to stable mature state or beginning senescence depending on subsequent environmental conditions and harvest timing. This completion occurs during late veraison to early ripening (BBCH 85-89), typically 14-20 days post-color break depending on cultivar and conditions, and is characterized by cessation of net anthocyanin accumulation (25-40% weekly changes during active development. Molecular markers of completion inclu anthocyanin biosynthetic genes (UFGT, DFR, LDOX, F3'H, F3'5'H, CHS, PAL) showing coordinated downregulation from peak expression (100-200 fold) to sustained moderate levels (40-80 fold) providing maintenance synthesis without driving major accumulation, regulatory transcription factors (VvMYBA1, VvMYBA2, VvMYC1) declining from maximum induction (120-180 fold) to moderate sustained expression (50-90 fold) maintaining pathway activation without intense developmental signaling, and hormonal transition with ABA declining from veraison peak 2500-3500 ng/g to plateau 1200-2200 ng/g consolidating ripening program completion while other hormones (gibberellin, auxin) remain at low post-growth levels and ethylene showing cultivar-dependent patterns (slight elevation 0.2-0.8 nL g⁻¹ h⁻¹ in some varieties, baseline 70 mM providing exceptional color intensity, and membrane integrity generally maintained though beginning slight permeabilization in some cultivars marking transition toward senescence particularly if environmental stresses occur. Environmental factors affecting completion timing inclu cumulative heat during veraison with completion typically occurring 550-750 growing degree days (base 10°C) post-color break in most cultivars, temperature during late development with optimal 20-26°C enabling complete maturation in 14-18 days while cooler conditions 16-22°C extend completion to 18-24 days and heat stress >30°C can force premature apparent completion through biosynthetic shutdown at 10-14 days potentially preventing full color potential achievement, light exposure determining final anthocyanin levels achieved at completion with optimal exposure reaching 30-50% higher levels than shaded fruit, and water availability with moderate sustained deficit through completion producing 25-40% higher final anthocyanins compared to well-watered vines though severe stress can arrest development preventing completion. Cultivar variation in completion timing reflects: early-completing varieties (Pinot Noir, Gamay, Grenache) achieving completion 12-16 days post-color break under favorable conditions, moderate cultivars (Merlot, Syrah, Cabernet Franc, Tempranillo) completing 15-19 days post-break, and late-completing types (Cabernet Sauvignon, Nebbiolo, Mourvèdre, Tannat) requiring 18-24 days with genetic differences in biosynthetic capacity, regulatory control, and developmental programming determining completion kinetics. Coordination with other ripening processes at completion shows: berry sugar accumulation transitioning from rapid phase (1.5-3.5% Brix per week) to slow accumulation or plateau (<0.5-1.0% per week) often coinciding with color completion, acid decline slowing substantially as malate respiratory consumption approaches minimum sustainable levels with titratable acidity 4.0-7.5 g/L stabilizing, berry softening approaching final state with firmness 1.0-2.2 N representing 70-85% total softening completed, seed maturation achieving brown color and lignified testa synchronized with color completion, and flavor/aroma development reaching characteristic intensity and complexity suitable for harvest or continued maturation depending on wine style objectives. Physiological significance of completion includes: transition from sink-driven developmental processes to source-sink equilibrium or beginning source mobilization in extended ripening, shift from active biosynthetic programming to maintenance metabolism or senescence depending on vine status and environmental conditions, and establishment of final berry composition determining wine quality potential with further changes primarily degradative (over-ripening) or concentrative (dehydration) rather than developmental. Quality implications of color development completion timing inclu completion synchronized with chemical maturity (target sugar, acid, pH) and phenolic maturity (seed browning, tannin extractability) indicates optimal harvest window for balanced wine production, premature completion before adequate flavor development or tannin maturation requires difficult decisions between harvesting color-mature but compositionally immature fruit or extending hang time risking color degradation, delayed completion beyond chemical maturity creates opposite challenge of harvesting at peak composition before complete color or waiting for color completion risking over-ripeness and alcohol elevation, and completion timing variation within vineyards reveals maturity heterogeneity informing selective harvest decisions or requiring acceptance of compositional variation. Monitoring color development completion involves: tracking anthocyanin accumulation rates through regular sampling showing plateau (<5% increase over 2 weeks) indicating completion, gene expression analysis in research settings measuring UFGT and regulatory factor transcripts confirming downregulation to maintenance levels, colorimetric monitoring showing stabilization of L*a*b* values with minimal week-to-week variation, visual assessment noting stable fully developed color without further intensification, and integration with chemical and sensory evaluation ensuring color completion synchronized with overall maturity for optimal harvest timing. Post-completion berry status shows: stable color persisting 2-6 weeks under favorable conditions (moderate temperatures, no extremes) enabling flexible harvest scheduling particularly in stable cultivars, potential color degradation under stress conditions (heat, water deficit extremes, disease) reducing anthocyanins 15-40% and shifting hue toward brown-red indicating over-ripeness or senescence, continued flavor and aroma development post-color completion in some cultivars benefiting from extended hang time for complexity enhancement, and risk-benefit assessment of extended ripening weighing flavor/phenolic improvement against color stability, disease pressure, and weather forecasts informing harvest timing decisions. Viticultural management at completion includes: irrigation often reduced or terminated recognizing decreased berry water demand and growth, canopy adjusted to protect colored berries from excessive sun preventing degradation while maintaining adequate exposure for continued ripening in other parameters, disease prevention intensified as fully colored soft berries show maximum Botrytis vulnerability, and harvest logistics activated with completion indicating optimal window approach requiring readiness for rapid harvest if conditions deteriorate or quality peaks.
Color_Development_Completion is distinguished from Complete_Color_Change by representing process termination rather than state achievement, from Final_Color_Expression by focusing on developmental endpoint rather than phenotypic stability, and from Color_Change_Onset by marking completion versus initiation. Completion markers: anthocyanin plateau <3-5% over 2 weeks, gene expression 30-50% of peak, colorimetric stability ±8-10% week-to-week. Timing: fast cultivars 12-16 days, moderate 15-19 days, late 18-24 days post-color break. Color_Development_Completion indicates transition from active ripening to stable mature state suitable for harvest timing optimization based on integrated maturity assessment.
Anthocyan-Synthese-Vollendung
Farb-Entwicklungs-Endpunkt
Pigmentierungs-Programm-Terminierung
anthocyanin synthesis completion
chromatic maturation finalization
chromatische Reifungs-Finalisierung
color development endpoint
pigmentation program termination
Keller M (2020) The Science of Grapevines 3rd Ed pages 238-242 ripening completion and harvest timing Coombe BG (1992) Research on development and ripening of the grape berry Australian J Grape Wine Res 48:125-144 developmental completion Deluc LG, Grimplet J, Wheatley MD, Tillett RL, Quilici DR, Osborne C, Schooley DA, Schlauch KA, Cushman JC, Cramer GR (2007) Transcriptomic and metabolite analyses of Cabernet Sauvignon grape berry development Bmc Genomics 8:429 molecular markers of ripening completion.
Color Development Completion
Color developmental stage assessment in Shiraz vineyards employs standardized visual protocols where scouts evaluate the proportion of berries at each color stage (green, pink, red, dark purple, blue-black) across representative clusters, with the transition from predominantly green (less than 20% colored) through véraison (20-80% colored) to mature coloration (greater than 80% colored) occurring over approximately 14-21 days depending on seasonal temperature this color staging provides the basis for harvest date prediction models that estimate 35-50 days from 50% véraison to optimal harvest maturity for premium red wine production.
Color_Developmental_Stage_Quality encompasses the morphological qualities that characterize the coloration state of grapevine organs in relation to their developmental progression. This category recognizes that organ color changes systematically during development, with specific color states serving as markers of developmental stage. In grape berries, the most dramatic developmental color change occurs at véraison, the onset of ripening, when colored cultivars transition from green chlorophyll-dominated coloration to cultivar-specific anthocyanin-based colors. This transition provides a visually obvious phenological marker used for vineyard management decisions and research documentation. The progression of color through developmental stages reflects underlying biochemical and physiological transitions including chlorophyll degradation, carotenoid unmasking, anthocyanin biosynthesis activation, and changes in vacuolar pH that affect pigment expression. This category subdivides into stage-specific quality classes including Véraison_Color_Quality capturing the transitional coloration during the onset of ripening, and Mature_Color_Quality representing the final coloration achieved at harvest maturity. The developmental trajectory of color change varies among cultivars in timing, rate, and final state, providing cultivar identification characters and indicating adaptation to different growing conditions. Early-ripening cultivars complete color development in shorter time periods than late-ripening cultivars. Environmental factors including temperature, light, and water availability modulate the rate and completeness of developmental color change within genetically determined limits.
Color_Developmental_Stage_Quality connects color as a morphological quality to phenological staging, bridging the Quality and Process categories of the VVO. While the visual appearance at any moment is a quality, the progression through color stages reflects underlying developmental processes. The child classes Véraison_Color_Quality and Mature_Color_Quality represent specific stages in the developmental trajectory. For white (green-skinned) cultivars, the developmental color change is less dramatic but still occurs as chlorophyll degrades and yellow carotenoids become visible. Assessment protocols should standardize developmental stage reference points for reproducibility across studies and seasons.
Farbentwicklungsstadium
Reifestadiumfarbe
developmental color state
entwicklungsabhängige Färbung
maturation color character
ontogenetic color quality
ontogenetische Farbqualität
ripening stage coloration
Coombe BG and McCarthy MG (2000) Dynamics of Grape Berry Growth and Physiology of Ripening. Australian Journal of Grape and Wine Research 6(2): 131-135 Kuhn N et al. (2014) Berry Ripening: Recently Heard Through the Grapevine. Journal of Experimental Botany 65(16): 4543-4559
Color Developmental Stage Quality
Color distribution assessment in Flame Seedless table grapes determines pack-out grades, with premium fruit requiring uniform red coloration covering at least 90% of the berry surface without green shoulders or pale contact patches fruit exhibiting gradient distribution with green-to-red transitions covering more than 10% of surface area is downgraded to second tier, while variegated distribution with discrete green patches interspersed with red zones indicates delayed or incomplete ripening that relegates fruit to processing grades regardless of sugar content.
Color_Distribution_Quality encompasses all morphological qualities that characterize the spatial pattern of coloration across the surface or throughout the volume of grapevine organs. While Pigmentation_Intensity_Quality addresses how much color is present and Hue_Quality describes what type of color occurs, Color_Distribution_Quality captures where and how uniformly color appears within an organ. This spatial dimension of color is diagnostically important because many cultivars display characteristic distribution patterns that aid identification, and distribution uniformity affects commercial value particularly for table grapes where visual appearance commands premium prices. This category comprises three primary distribution patterns: Uniform_Coloration_Quality describing homogeneous color across the organ surface Variegated_Coloration_Quality capturing discrete patches or zones of contrasting color and Gradient_Coloration_Quality addressing continuous transitions from one color intensity or hue to another across the organ. In grape berries, color distribution results from the spatial pattern of anthocyanin accumulation in skin cells, which depends on the distribution of biosynthetic enzyme activity, light exposure patterns across the berry surface, and temperature gradients between exposed and shaded portions. Berries in compact clusters often display uneven coloration where contact zones between adjacent berries remain less colored than exposed surfaces. Environmental heterogeneity at the cluster scale, including variable light penetration through the canopy, creates distribution patterns that reflect microclimate conditions. In leaves, color distribution patterns during senescence reveal vascular architecture as green color persists longest near veins where chlorophyll degradation is delayed. Color distribution assessment employs visual evaluation of pattern type and extent, image analysis quantifying color variation across defined regions, or spectrophotometric mapping of pigment concentrations at multiple sample points.
Color_Distribution_Quality is positioned as a direct subclass of Color_Quality, parallel to Pigmentation_Intensity_Quality and Hue_Quality. Distribution patterns interact with both intensity and hue, as gradients may involve changes in intensity, hue, or both across the organ surface. The three child classes (Uniform, Variegated, Gradient) represent idealized distribution patterns actual organs commonly display combinations or intermediate patterns. Assessment of color distribution should specify the viewing angle and lighting conditions, as apparent distribution can vary with observation geometry. Digital image analysis enables objective quantification of distribution patterns through pixel-by-pixel color classification.
Farbmusterqualität
Farbverteilungsqualität
Pigmentverteilung
color pattern quality
color uniformity character
pigment distribution
räumliche Färbung
spatial coloration
Peppi MC et al. (2006) Abscisic Acid and Sucrose Regulate Berry Skin Color in Flame Seedless Grapes. Journal of the American Society for Horticultural Science 131(2): 241-248 Downey MO et al. (2006) Cultural Practice and Environmental Impacts on the Flavonoid Composition of Grapes and Wine: A Review of Recent Research. American Journal of Enology and Viticulture 57(3): 257-268
Color Distribution Quality
In a Napa Valley Cabernet Sauvignon vineyard, color distribution uniformity was assessed across 120 vines in late September Year N+1 (BBCH 86) showing within-berry CV 16%, within-cluster CV 24%, within-vine CV 28%, within-block CV 32%, mean skin anthocyanins 1420 mg/kg (SD ±455 mg/kg), berry sugar 24.2° Brix (SD ±1.8°), enabling single-pass mechanical harvest of uniformly ripe fruit under optimal management (uniform drip irrigation, standardized canopy, consistent crop load 8 kg/vine) on well-drained hillside benchland.
Color_Distribution_Uniformity represents the spatial consistency of pigmentation across berries within clusters, among clusters on vines, and throughout vineyard blocks reflecting synchronized ripening that enables optimal harvest timing and consistent wine quality by minimizing compositional variation in harvested fruit. This quality emerges during late veraison to early ripening (BBCH 85-87) when developmental heterogeneity diminishes through completion of ripening processes across the population, and is quantified by measuring color variation at multiple spatial scales: within-berry color uniformity (coefficient of variation 40 years) sometimes showing different ripening timing than prime-age vines (10-30 years), rootstock effects on ripening timing creating 2-4 day variation when mixed rootstocks exist within blocks, and management history including pruning severity, crop load, irrigation, and disease pressure creating spatial patterns of advancement or delay. Monitoring color distribution uniformity involves: stratified random sampling across vineyard blocks collecting berry samples from multiple vines, cluster positions, and berry locations within clusters enabling statistical analysis of variation, image analysis of cluster or vine photographs using color segmentation algorithms calculating percentages of fruit at target color stages, portable spectrophotometers or colorimeters measuring berry color at systematic sampling points across vineyard creating spatial variation maps, and visual assessment using trained observers rating uniformity on standard scales (1-10) providing rapid semi-quantitative evaluation. Environmental factors affecting distribution uniformity inclu weather stability during veraison with consistent moderate temperatures (20-26°C) and no extreme events promoting uniform development, while dramatic weather shifts (heat waves, cold snaps, heavy rain) create developmental disruption affecting berries differentially based on their developmental stage and location, irrigation uniformity with consistent water distribution across vineyard enabling similar vine water status and berry development versus heterogeneous irrigation creating spatial variation in ripening timing and intensity, and disease pressure with localized infections (Botrytis, powdery mildew) disrupting normal development in affected zones creating color variation even in otherwise uniform blocks. Viticultural practices influencing uniformity inclu shoot thinning and cluster thinning performed uniformly across vineyard promotes synchronized development by standardizing crop load and canopy density, selective thinning removing lagging clusters can enhance apparent uniformity by eliminating developmental outliers though reducing total yield, canopy management (hedging, leaf removal, shoot positioning) applied consistently optimizes light distribution improving within-vine and block-scale uniformity, and irrigation management maintaining consistent vine water status across spatial scales prevents drought or excess water from creating localized ripening heterogeneity. Harvest implications of color distribution uniformity inclu high uniformity (vineyard-scale CV 45%) necessitates difficult decisions between accepting compositional variation in single harvest or conducting multiple selective harvests increasing labor costs and complexity, and integration with sugar and acid uniformity determines overall harvest strategy since color uniformity alone may not indicate complete maturity synchrony. Wine quality implications inclu high color uniformity predicts consistent wine quality with low bottle-to-bottle variation (important for premium wines), moderate uniformity produces acceptable quality with some vintage variation requiring skillful blending or lot separation, poor uniformity creates quality challenges with unripe and overripe fruit components potentially requiring extensive sorting and selective processing, and uniformity correlates with other quality parameters (phenolic maturity, flavor development) suggesting color distribution uniformity serves as integrated maturity indicator. Temporal dynamics of uniformity show: color variation typically highest at veraison onset (CV 60-80% across berries), declining progressively during veraison as lagging berries catch up (CV reducing 5-10% per week), achieving minimum variation at complete color development (CV 20-35% depending on inherent block heterogeneity), and potentially increasing post-ripening through differential senescence or degradation rates creating renewed heterogeneity if harvest delayed beyond optimal window. Cultivar variation in uniformity characteristics: varieties with naturally synchronous development (Pinot Noir, Chardonnay) achieve high uniformity (CV 35-40%) even under optimal conditions requiring acceptance of heterogeneity or selective harvest approaches. Economic considerations of uniformity: high uniformity enables efficient single-pass machine harvest reducing labor costs substantially (savings 40-60% versus hand harvest), allows streamlined processing without extensive sorting reducing winery labor and equipment needs, and produces consistent wine reducing blending complexity and inventory management challenges, while low uniformity necessitates expensive selective hand harvest, extensive sorting and berry selection infrastructure, and complex lot management potentially justified only for premium wines commanding sufficient price premiums.
Color_Distribution_Uniformity is distinguished from Uniform_Color_Development by representing population-level variation rather than individual berry uniformity, from Even_Berry_Coloration by focusing on spatial distribution across multiple scales rather than single-berry consistency, and from Complete_Color_Change by emphasizing variation rather than completion. Uniformity metrics: within-berry CV <15-20%, within-cluster <20-30%, within-vine <25-35%, within-block <30-40%. High uniformity enables efficient harvest and consistent wine quality. Low uniformity requires selective harvest or acceptance of compositional variation.
homogene Pigmentierungs-Verteilung
homogeneous pigmentation distribution
räumliche Farb-Konsistenz
spatial color consistency
synchronisierte Farb-Entwicklung
synchronized color development
uniform berry coloring pattern
uniformes Beeren-Färbungs-Muster
Keller M (2020) The Science of Grapevines 3rd Ed pages 237-238, 266-268 ripening heterogeneity and harvest Bramley RGV, Hamilton RP (2004) Understanding variability in winegrape production systems Australian J Grape Wine Res 10:32-45 spatial variation in vineyards Kontoudakis N, Esteruelas M, Fort F, Canals JM, Zamora F (2011) Use of unripe grapes harvested during cluster thinning as a method for reducing alcohol content and pH of wine Aust J Grape Wine Res 17:230-238 maturity variation effects.
Color Distribution Uniformity
Color hue assessment in Bordeaux vertical tasting tracks the evolutionary trajectory from 2020 vintage (purple-ruby with violet reflections indicating youth, dominant free anthocyanins) through 2010 (ruby-garnet with orange hints at rim indicating mid-evolution, polymeric pigment development) to 2000 (garnet-tawny throughout indicating maturity, predominantly polymeric pigments), demonstrating how hue progression provides visual age confirmation and optimal drinking window assessment, with the 2000 hue suggesting consumption priority while 2020 hue indicates cellaring potential.
Color hue appearance quality is a color appearance quality that represents the dominant wavelength character of wine color determining its position on the color spectrum, characterizing whether wine appears purple, ruby, garnet, tawny, straw, gold, or amber as an indicator of variety, age, and oxidation state. Within the BFO framework, this quality specifically addresses the qualitative chromatic dimension of color, distinguishing hue (which color) from intensity (how much color) while recognizing that hue provides the most age-sensitive color information, with systematic hue shifts tracking wine evolution from youth through maturity to senescence. The ontological significance lies in hue serving as the primary age indicator in visual assessment, where red wine hue progression from purple through ruby through garnet to tawny-brown reflects anthocyanin-tannin polymerization and oxidation over time, while white wine progression from pale straw through gold to amber signals oxidative evolution, making hue the visual clock that indicates developmental stage. The physical basis for hue involves the specific wavelengths absorbed by wine chromophores, with free anthocyanins absorbing in the 520-540 nm range (producing red-purple), polymeric pigments absorbing at longer wavelengths (creating orange-brown), and the equilibrium between these forms shifting with age, pH, and oxidation to produce the characteristic hue evolution. The compositional determinants of hue include anthocyanin profile (malvidin-dominant varieties appearing more blue-purple, cyanidin-dominant appearing more red), pH (lower pH favoring red flavylium cation, higher pH shifting toward blue quinoidal forms), polymeric pigment proportion (increasing with age, shifting hue toward orange), and oxidation state (creating brown tones through quinone formation). The variety associations of hue are significant, with young Pinot Noir showing ruby-red hue distinct from young Malbec's purple-black, and aged wines of all varieties converging toward tawny-brown as polymeric pigments dominate over monomeric anthocyanins. Understanding color hue quality enables age estimation from hue position on the evolutionary trajectory, variety identification from characteristic hue expression, and condition assessment detecting premature oxidation or unusual hue development.
Color_Hue_Appearance_Quality addresses the dominant wavelength character of wine color, representing the most age-sensitive visual parameter. This quality follows predictable evolutionary trajectories that enable age estimation and maturity assessment. The class supports age evaluation, variety identification, and condition assessment based on hue position relative to expected evolutionary stage.
Farbnuance
Farbschattierung
Farbton
chromatic hue
chromatischer Ton
color shade
color tone
dominant wavelength
Somers TC, Evans ME (1977) Spectral evaluation of young red wines: anthocyanin equilibria, total phenolics, free and molecular SO₂, chemical age. Journal of the Science of Food and Agriculture 28(3): 279-287 Dallas C, Laureano O (1994) Effects of pH, sulphur dioxide, alcohol content, temperature and storage time on colour composition of a young Portuguese red table wine. Journal of the Science of Food and Agriculture 65(4): 477-485
Color Hue Appearance Quality
Color intensity for Shiraz must from McLaren Vale (2023 vintage) records CI 22.5 AU (A420 5.8, A520 12.2, A620 4.5, measured at 1 mm path length), representing exceptionally deep color indicative of the old-vine, low-yield source material and predicting a finished wine of opaque, inky-purple character. Color intensity for Pinot Noir must from Volnay, Burgundy (2024 vintage) measures CI 7.8 AU (A420 2.5, A520 4.2, A620 1.1), within the typical range for Pinot Noir (6-12 AU) that reflects the cultivar's moderate anthocyanin content and predominance of non-acylated monoglucosides producing the characteristically brilliant ruby-garnet hue. Color intensity for Zweigelt from Carnuntum (2024 vintage) documents CI 14.8 AU, confirming the variety's reputation as Austria's most deeply colored red cultivar and validating the chromatic density essential for the Carnuntum DAC quality designation.
Color intensity is a phenolic data entity that records the integrated spectrophotometric color expression of grape must or wine, determined by absorbance measurement at three wavelengths representing the yellow-brown (420 nm), red (520 nm), and blue-violet (620 nm) components of the visible spectrum, calculated as the sum CI = A420 + A520 + A620 (Glories method) and typically measured in absorbance units (AU) through a 1 mm or 10 mm path length cuvette, representing the net chromatic contribution of anthocyanins, copigmentation complexes, polymeric pigments, and oxidation products that collectively produce the visual color appearance of red and rosé wines.
Color intensity measurement provides the most direct and rapid assessment of red wine visual quality, correlating strongly with consumer perception of wine quality and price positioning — deeply colored red wines consistently command higher prices and quality ratings than pale examples of the same variety and region. The three-wavelength summation (A420 + A520 + A620) captures the full chromatic complexity of wine color, which arises from multiple chemical species in pH-dependent equilibrium: free anthocyanins in the flavylium cation form (red, dominant contributor to A520), anthocyanin-tannin copigmentation complexes (enhanced red with blue shift), polymeric pigments formed through acetaldehyde-mediated condensation (stable red-brown, contributing to A420 and A520), and oxidation products (yellow-brown, contributing to A420). The derived parameter "hue" (A420/A520) provides complementary information — young red wines exhibit hue values of 0.5-0.7 (red-dominant), while aged wines shift toward 0.8-1.2 (increasing brown-yellow contribution). Color intensity is measured in must (post-crushing, before fermentation), during fermentation (monitoring extraction kinetics), and in wine (assessing extraction efficiency and aging evolution), with each context providing different information. The path length specification (1 mm or 10 mm) must always be reported, as CI values differ 10-fold between these two common cuvette sizes — the 1 mm path length is standard for deeply colored wines to maintain absorbance within the linear range of the spectrophotometer. Must color intensity predicts but does not equal wine color intensity, as fermentation extraction, fining, and aging all modify the color profile. The OIV reference method and the Iland/Somers modified color assay provide internationally recognized standardized protocols for color intensity determination.
CI value
Farbdichte
Farbintensität
Farbintensitätswert
Farbkraft
chromatic intensity
chromatische Intensität
color density
spectral color intensity
wine color intensity
Glories Y (1984) La couleur des vins rouges. 2e partie: mesure, origine et interprétation. Connaissance de la Vigne et du Vin 18(4): 253-271 Somers TC, Evans ME (1977) Spectral evaluation of young red wines: anthocyanin equilibria, total phenolics, free and molecular SO₂, chemical age. Journal of the Science of Food and Agriculture 28(3): 279-287 Iland P et al. (2011) Chemical Analysis of Grapes and Wine, 2nd Ed, pp.: 105-115 OIV (2021) Compendium of International Methods of Analysis, Method OIV-MA-AS2-07B
Color Intensity
Color intensity data for Cabernet Sauvignon clone 337 versus clone 15 at Coonawarra across 8 vintages documents total anthocyanins of 1.8-2.4 mg/g berry fresh weight in clone 337 versus 1.4-1.8 mg/g in clone 15, with clone 337 additionally exhibiting a higher proportion of acylated anthocyanins (38% vs. 28% of total) conferring greater color stability during aging. Wine color intensity (CI = A420 + A520 + A620) from standardized micro-vinifications averaged 14.5 AU for clone 337 versus 10.8 AU for clone 15. Color intensity data for Blaufränkisch clone Bl-2 at Deutschkreutz records anthocyanin concentrations of 2.5-3.2 mg/g at 20° KMW harvest, dominated by malvidin-3-glucoside (55-60% of total), producing wines with CI values of 16-20 AU that rank among the highest in the Austrian clonal evaluation program.
Color intensity data is a quality performance data entity that records the concentration, composition, and spectrophotometric characteristics of pigment compounds — principally anthocyanins and their derivatives — in the berry skin, must, and wine of a red or rosé clonal line, expressed through optical density measurements (absorbance at 420, 520, and 620 nm), total anthocyanin concentration (mg/g berry weight or mg/L must/wine), individual anthocyanin profiling (malvidin, peonidin, delphinidin, cyanidin, petunidin glycosides and acylated forms), and derived color parameters (color intensity CI, hue, CIELab coordinates), characterizing the chromatic potential fundamental to red and rosé wine visual quality.
Color intensity data is a primary differentiator among red wine clones, as anthocyanin accumulation exhibits substantial clonal variation within cultivars — differences of 30-80% in total anthocyanin concentration between clones of the same variety are commonly documented. The genetic basis includes variation in the anthocyanin biosynthetic pathway genes (UFGT, MYBA1, MYBA2, F3'5'H) and their regulatory elements, with some clonal color differences traceable to specific mutations in the MYB transcription factor family. Environmental modulation of anthocyanin accumulation is pronounced — optimal synthesis occurs at berry temperatures of 17-25°C with adequate UV exposure, while temperatures above 35°C cause anthocyanin degradation, creating significant clone-by-environment interactions for color intensity. The anthocyanin profile (relative proportions of the five anthocyanidin types and their acylation and methylation patterns) provides clone-specific fingerprinting and influences both color hue (blue-shifted by delphinidin/petunidin, red-shifted by cyanidin/peonidin) and color stability (enhanced by acylation and copigmentation). Color intensity data should be assessed at standardized maturity rather than calendar date, as sugar-matched comparisons eliminate the confounding effect of harvest timing differences on color accumulation stage. The increasing importance of rosé wine production has expanded color intensity evaluation to include clonal assessment for moderate, controlled anthocyanin release during short maceration.
Anthocyandaten
Farbdaten
Farbintensitätsdaten
Pigmentgehaltsdaten
anthocyanin data
chromatic intensity data
color data
pigment concentration data
wine color record
Downey MO et al. (2006) Cultural practice and environmental impacts on the flavonoid composition of grapes and wine: a review. American Journal of Enology and Viticulture 57(3): 257-268 Boss PK et al. (1996) Expression of anthocyanin biosynthesis pathway genes in red and white grapes. Plant Molecular Biology 32(3): 565-569 Castellarin SD et al. (2007) Colour variation in red grapevines: the colour of berry skin and flesh is associated with differences in anthocyanin profile. Journal of Experimental Botany 58(15-16): 4257-4270
Color Intensity Data
In Petit Verdot berries in Margaret River, Australia during BBCH 81-89, color intensity development progresses from initial pink coloration with color density of 2.8 AU at veraison to deep purple-black with color density of 18.5 AU at harvest over 45 days, with CIELab lightness decreasing from 45 to 22, a* red coordinate increasing from 18 to 42, and b* blue coordinate shifting from +5 to -8 indicating blue hue development, paralleling anthocyanin increase from 0.25 to 2.85 mg/g and copigmentation enhancement contributing 42% of total color at optimal ripeness.
Color intensity development describes the progressive enhancement of visual pigmentation in Vitis vinifera berries during the post-veraison ripening period, wherein accumulating anthocyanin concentrations combined with copigmentation interactions and pH-dependent chromophore equilibria produce the increasing depth and saturation of red-purple-blue coloration that characterizes ripening progression from the initial pink flush at veraison to the deep color of fully mature fruit at harvest. This process reflects the relationship between absolute anthocyanin concentration and perceived color intensity, with color development following characteristic patterns as pigment levels increase from threshold visibility of 50-100 mg/kg through moderate intensity at 500-800 mg/kg to deeply saturated color above 1500 mg/kg, though the precise concentration-intensity relationship varies with anthocyanin composition, copigmentation status, and vacuolar pH conditions affecting chromophore expression. The spectroscopic basis of color intensity development involves the pH-dependent equilibrium among anthocyanin structural forms, with the red flavylium cation predominating at low vacuolar pH of 2.5-3.5, the colorless carbinol pseudobase forming at higher pH, and quinoidal base forms contributing blue-purple hues, with the distribution among forms determining the hue and intensity of color expression at any given anthocyanin concentration. Copigmentation interactions enhance color intensity development through molecular stacking of anthocyanins with colorless copigments including flavonols, hydroxycinnamic acids, and other flavonoids, producing hyperchromic shifts that increase apparent color intensity by 30-50% beyond the contribution of anthocyanins alone, with copigment-to-anthocyanin ratios influencing the magnitude of this enhancement throughout ripening. The temporal progression of color intensity development shows variety-specific patterns with rapid initial color development during early post-veraison as anthocyanins first exceed visual threshold, continued intensity increase during mid-ripening paralleling anthocyanin accumulation, and potential intensity stabilization or decline during late ripening when degradation may reduce anthocyanin content or structural changes modify chromophore expression. Environmental factors influence color intensity development through effects on both anthocyanin accumulation and color expression, with excessive temperature reducing intensity through inhibition of synthesis and accelerated degradation, while appropriate light exposure enhances intensity through promotion of anthocyanin and copigment accumulation. Assessment of color intensity development employs multiple methods including visual scoring systems rating berry color on structured scales, spectrophotometric measurements of color density and hue at characteristic wavelengths, and colorimetric analysis using CIELab coordinates that quantify lightness, red-green balance, and yellow-blue balance providing comprehensive color characterization.
Color_Intensity_Development is subclass of Anthocyanin_Accumulation representing visual color outcome. Distinguishes from Progressive_Anthocyanin_Synthesis which describes biosynthetic process. Sibling to Progressive_Anthocyanin_Synthesis and Anthocyanin_Profile_Evolution. Integrates anthocyanin concentration with copigmentation and pH effects. Spectrophotometric and colorimetric assessment. Threshold visibility at 50-100 mg/kg anthocyanins.
Beeren-Farb-Entwicklung
Farbintensitäts-Entwicklung
Farbsättigungs-Entwicklung
berry color development
color saturation development
pigment intensity enhancement
visual pigmentation increase
visuelle Pigmentierungs-Zunahme
Boulton R (2001) The copigmentation of anthocyanins and its role in the color of red wine. American Journal of Enology and Viticulture 52: 67-87 Ribéreau-Gayon P et al (2006) Handbook of Enology Vol 2 The Chemistry of Wine 2nd Ed Wiley pp 141-205 Keller M (2020) The Science of Grapevines 3rd Ed pp 315-322
Color Intensity Development
In a Mendoza Malbec vineyard in Argentina, color intensity increase was documented over 8 days in late February Year N+1 (BBCH 84-85) when CIE L* declined from 48 to 28 (darkening), chroma increased from 32 to 51 (greater saturation), skin anthocyanins rose from 225 mg/kg to 980 mg/kg, absorbance at 520 nm increased from 0.8 to 2.4 OD, berry Brix advanced 15.2 to 22.5°, under optimal high-altitude conditions (daily high 27-30°C, night 12-15°C, intense UV, moderate deficit stem ψ -1.15 MPa).
Color_Intensity_Increase represents the visually perceptible amplification of berry surface pigmentation during progressive color development when accumulated anthocyanins create progressively deeper, more saturated red-purple coloration transforming berry appearance from pale pink-red to intense deep color characteristic of fully pigmented fruit ready for harvest or further maturation. This process parallels increasing anthocyanin accumulation occurring during mid-to-late veraison (BBCH 84-85), typically 4-10 days post-color break, and is characterized by progressive shifts in color metrics: CIE L* (lightness) declining from 50-65 at color break to 25-40 at peak intensity indicating darkening, chroma (C*) increasing from 20-35 to 35-55 reflecting greater color saturation, and hue angle shifting from 20-35° (orange-red) toward 0-15° (pure red) or 340-355° (red-purple) as anthocyanins overwhelm residual chlorophyll and carotenoid contributions. Optical basis of intensity increase involves: anthocyanin concentration rising from 100-200 mg/kg to 500-1200 mg/kg creates proportional increase in light absorption at 520-540 nm wavelengths where anthocyanins show maximum absorbance, extinction coefficient effects where concentrated anthocyanin solutions show non-linear color intensity increases through molecular interactions and aggregation, copigmentation phenomena between anthocyanins and colorless flavonols or hydroxycinnamic acids producing 30-50% color intensification beyond that expected from anthocyanin concentration alone through bathochromic shifts and hyperchromic effects, and anthocyanin vacuolar inclusion formation in some cultivars creating localized pigment concentrations >60 mM generating extremely intense color visible even in small tissue volumes. Spectral characteristics during intensity increase show: absorbance spectra developing characteristic anthocyanin profile with maximum 520-540 nm and secondary peak 270-280 nm, visible absorbance at 520 nm increasing from optical density 0.3-0.6 at color break to 1.5-3.0 at peak intensity representing 5-10 fold amplification, reflectance spectra showing declining green reflection (500-600 nm) from 30-45% at color break to 32°C severely impairs intensity increase with final color 40-70% below optimal temperature treatment producing pale inadequately colored fruit, and diurnal temperature variation >12-15°C combining warm days facilitating synthesis with cool nights enhancing stability often optimizes final intensity. Light effects on intensity increase demonstrate: UV-B radiation enhances intensity 30-50% through increased anthocyanin synthesis and potential effects on copigment accumulation, moderate visible light exposure optimizes intensity while excessive radiation combined with heat can reduce intensity through anthocyanin degradation and synthesis inhibition, canopy position effects create 35-60% intensity variation between sun-exposed and shaded clusters, and cluster-level light distribution affects within-cluster intensity uniformity with exposed berries showing 2-4 fold higher intensity than completely shaded berries. Water stress influences show: moderate deficit (stem ψ -1.0 to -1.3 MPa) enhances intensity 25-40% through elevated anthocyanin concentration and potential berry size reduction concentrating pigments per unit surface area, severe deficit (<-1.5 MPa) reduces intensity despite high anthocyanin concentration through metabolic impairment limiting synthesis, while excess water dilutes berries and reduces intensity 30-50% through lower anthocyanin per unit fresh weight and potentially reduced synthesis through suppressed ABA signaling. Cultivar variation in intensity characteristics reflects: high-intensity varieties like Petit Verdot, Tannat, and Syrah achieve very deep color (L* 20-30, anthocyanins 1500-3000 mg/kg) with strong purple notes from high delphinidin, moderate-intensity cultivars like Cabernet Sauvignon and Merlot display deep red color (L* 25-35, 800-1500 mg/kg anthocyanins), while lower-intensity varieties like Pinot Noir and Grenache show bright-to-medium red (L* 30-40, 600-1200 mg/kg) with characteristic hue differences reflecting anthocyanin composition. Temporal progression of intensity increase follows: rapid initial increase during days 3-6 of progressive development when anthocyanin synthesis peaks, slower continued increase days 7-10 as accumulation rates decline approaching maximum, and plateau or slight decrease post-day 10 as synthesis stops and potential degradation begins in some conditions demonstrating importance of harvest timing for optimal color capture. Monitoring intensity increase provides quality assessment: visual comparison with color standards or photographic references enables semi-quantitative tracking, portable colorimeters measuring CIE L*a*b* values provide objective intensity quantification correlating with wine color potential (r² = 0.80-0.92), and spectrophotometric analysis of skin extracts measuring anthocyanin absorbance at 520 nm gives precise color intensity prediction for harvest timing optimization. Wine quality implications of intensity increase inclu final berry color intensity strongly predicts wine color density (r² = 0.85-0.95) with deeper berry color producing higher wine absorbance at 520 nm, intensity progression rate affects anthocyanin stability characteristics with rapid intense coloring sometimes associated with less stable anthocyanins versus moderate progression producing more stable pigments, and spatial uniformity of intensity within clusters or vineyards indicates ripening synchrony affecting harvest decisions and wine composition consistency. Physiological correlates of intensity increase inclu color deepening typically parallels berry softening with peak intensity coinciding with 50-70% total firmness reduction, synchronizes with maximum sugar accumulation rates (berry Brix rising 1.5-3.5% per week during peak intensity increase), and occurs during intense malate degradation creating flavor and acid balance changes accompanying color development demonstrating integrated ripening program controlling multiple quality parameters simultaneously.
Color_Intensity_Increase is distinguished from Increasing_Anthocyanin_Accumulation by representing visual/spectral manifestation rather than biochemical quantity, from Progressive_Color_Development by focusing on intensity rather than spatial progression, and from Complete_Color_Change by exhibiting ongoing increase rather than final state. CIE L* range: color break 50-65, peak intensity 25-40. Copigmentation contributes 30-50% intensity enhancement. Temperature optimum 22-28°C. Moderate deficit enhances 25-40%. Color_Intensity_Increase provides practical visual assessment of ripening progress and wine quality prediction.
amplifying pigmentation intensity
deepening color saturation
increasing chromatic strength
intensifying visual color
intensivierende visuelle Farbe
steigende chromatische Stärke
verstärkende Pigmentierungs-Intensität
zunehmende Farb-Sättigung
Keller M (2020) The Science of Grapevines 3rd Ed pages 234-236 color development and quality Rolle L, Torchio F, Giacosa S, Gerbi V (2009) Modifications of mechanical characteristics and phenolic composition in berry skins and seeds of Mondeuse winegrapes throughout the on-vine drying process Amer J Enol Vitic 60:418-425 color intensity dynamics Guidoni S, Ferrandino A, Novello V (2008) Effects of seasonal and agronomical practices on skin anthocyanin profile of Nebbiolo grapes Amer J Enol Vitic 59:22-29 environmental effects on color intensity.
Color Intensity Increase
Color intensity comparison between Argentinian Malbec (intensity rating 9/10, deep purple-black with barely visible light transmission at rim) and Burgundy Pinot Noir (intensity 5/10, translucent ruby with clear light transmission throughout) demonstrates variety-driven intensity difference where both wines represent quality benchmarks for their respective varieties despite dramatically different color depth, illustrating how intensity assessment must be calibrated to variety expectation rather than applying universal deep-color preference.
Color intensity quality is a color appearance quality that represents the depth or strength of wine coloration regardless of specific hue, characterizing how pale or deep the wine appears as an indicator of pigment concentration, extraction level, and production approach. Within the BFO framework, this quality specifically addresses the quantitative dimension of color, distinguishing intensity (how much color) from hue (which color) while recognizing that intensity provides information about grape variety, viticultural practices, and winemaking decisions affecting pigment extraction and retention. The ontological significance lies in color intensity serving as a concentration and quality indicator across wine types, where deep color in red wines suggests ripe fruit, effective extraction, and age-worthy structure, while pale color may indicate light variety, underripe fruit, or minimal extraction, with intensity assessment enabling quality inference before aromatic evaluation. The physical basis for color intensity involves the Beer-Lambert relationship between pigment concentration and light absorption, with deeper color reflecting higher chromophore concentration in the light path, though the relationship is complicated by copigmentation effects that can enhance apparent intensity beyond what pigment concentration alone would produce. The compositional determinants of intensity include anthocyanin concentration in red wines (variety-dependent, ranging from 200-2000 mg/L), extraction efficiency during maceration, copigmentation with colorless phenolics (potentially doubling apparent color), pH effects on anthocyanin equilibrium, and in white wines, flavonol and carotenoid derivative concentrations determining depth from water-white through deep gold. The variety associations of color intensity show characteristic ranges, with Malbec, Syrah, and Tannat producing deeply colored wines while Pinot Noir and Nebbiolo yield paler examples despite high quality, making variety-appropriate intensity assessment essential rather than universal preference for deep color. Understanding color intensity quality enables extraction assessment, variety identification based on expected intensity range, and quality evaluation incorporating appropriate depth for wine type and style.
Color_Intensity_Quality addresses the quantitative depth of wine coloration independent of hue. This quality is strongly variety-dependent, with different varieties showing characteristic intensity ranges unrelated to quality level. The class supports extraction assessment and variety identification while cautioning against universal preference for deep color that would unfairly penalize legitimately pale varieties like Pinot Noir and Nebbiolo.
Farbstärke
Farbtiefe
chromatic density
chromatische Dichte
color depth
color strength
pigment intensity
Glories Y (1984) La couleur des vins rouges. 2e partie: mesure, origine et interprétation. Connaissance de la Vigne et du Vin 18(4): 253-271 Harbertson JF et al. (2003) A rapid method for measuring red wine color. American Journal of Enology and Viticulture 54(4): 301-305
Color Intensity Quality
The color mutation data for the Muscat family documents a complete series of berry color variants derived from the ancestral red-berried Muscat Rouge de Madère genotype — Muscat Blanc à Petits Grains (white berries through homozygous VvMYBA1 inactivation by Gret1 retrotransposon insertion on both chromosomes, producing the world's most widely planted Muscat for winemaking), Muscat Rosé à Petits Grains (pink berries through partial VvMYBA1 restoration, likely chimeral with functional VvMYBA1 in L2 but not L1), and Muscat Rouge à Petits Grains (full red coloration with functional VvMYBA1 on at least one homologue) — all variants share identical SSR profiles at 9 OIV loci, confirming their origin from a single genotype through successive somatic color mutations. The same molecular mechanism — Gret1 retrotransposon insertion/excision at VvMYBA1 — has been documented independently in the Pinot, Grenache, Terret, Aramon, and Savagnin cultivar families.
A mutation information record specific to somatic mutations affecting berry skin color, berry flesh color, or shoot tip pigmentation in Vitis vinifera, documenting the molecular change in anthocyanin biosynthesis regulatory or structural genes — principally VvMYBA1 and VvMYBA2 on chromosome 2 — that produces the white-berry, grey-berry, pink-berry, or teinturier (red-fleshed) phenotypic variants of originally red/black-berried cultivars, with color mutations representing the most frequent, commercially significant, and molecularly best-characterized category of somatic variation in grapevine.
Color_Mutation_Data documents the most extensively studied category of grapevine somatic mutation, owing to the conspicuousness of berry color change (making color sports easily detectable in the vineyard), the commercial importance of color variants (many white cultivars including Chardonnay, Pinot Blanc, Grenache Blanc, and Savagnin are white-berry sports of red ancestors), and the relatively simple genetic architecture underlying berry color (a single regulatory locus on chromosome 2 controlling the entire anthocyanin pathway). The molecular mechanism centers on the VvMYBA1 and VvMYBA2 genes encoding R2R3-MYB transcription factors that activate the anthocyanin biosynthetic pathway in berry skin — red/black berry color requires at least one functional copy of VvMYBA1 or VvMYBA2, while white berries result from the inactivation of all functional copies: the Gret1 retrotransposon (a 10,422 bp Ty1-copia class LTR retrotransposon) is the primary mutational agent, with its insertion into the VvMYBA1 promoter abolishing gene expression and its occasional excision restoring function (producing red reversion sectors on white-berry cultivars). The ubiquity of Gret1-mediated color mutations across diverse cultivar families suggests ongoing retrotransposon activity in the grapevine genome, with the VvMYBA1 locus representing a mutational hotspot due to the element's preferential insertion target sequence.
Anthocyan-Varianten-Daten
Beerenhautfarbe-Mutations-Dokumentation
Farbmutations-Daten
anthocyanin mutation data
berry color mutation record
skin color variant documentation
Kobayashi S et al. (2004) Retrotransposon-induced mutations in grape skin color, Science 304: 982 Walker AR et al. (2007) White grapes arose through the mutation of two similar and adjacent regulatory genes, Plant Journal 49: 772-785 Fournier-Level A et al. (2009) Quantitative genetic bases of anthocyanin variation in grape, Molecular Genetics and Genomics 281: 383-395 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340
Color Mutation Data
Color quality assessment in Pinot Noir destined for premium Burgundy production employs both visual evaluation against standardized color charts at harvest and instrumental CIELAB colorimetry on pressed juice, with target values of L* (lightness) between 45-55, a* (red-green) between 35-50, and b* (yellow-blue) between 5-15 indicating optimal anthocyanin accumulation without excessive extraction of oxidized or polymerized pigments that would compromise the bright ruby hue characteristic of fine Pinot Noir wines.
Color_Quality encompasses all morphological qualities that characterize the chromatic appearance of grapevine organs resulting from the interaction between incident light and pigment molecules, surface structures, and tissue optical properties. Color represents one of the most visually prominent and economically significant quality categories in viticulture because grape and wine color directly influences consumer perception, market value, and quality classification. This category subdivides into three major domains: Pigmentation_Intensity_Quality addressing the concentration and visibility of color-producing compounds Hue_Quality capturing the spectral character that distinguishes different color types and Color_Distribution_Quality describing the spatial pattern of coloration across organ surfaces. Grapevine color results primarily from three pigment classes: chlorophylls producing green coloration in photosynthetic tissues carotenoids contributing yellow and orange hues and anthocyanins generating the red, blue, and purple colors characteristic of many grape cultivars. The balance among these pigments, their subcellular localization, and their chemical modification (particularly anthocyanin acylation and co-pigmentation) determine the perceived color of organs. Color assessment employs visual matching to standardized color charts, instrumental colorimetry using tristimulus values (L*a*b* color space) or spectrophotometry, and chemical analysis of individual pigment concentrations. Color qualities exhibit pronounced developmental dynamics, particularly in berries where the dramatic color change at véraison marks the transition from green (chlorophyll-dominated) to cultivar-specific mature coloration. Environmental factors including light exposure, temperature, and water status modulate color expression within genetically determined limits, with implications for canopy management practices aimed at optimizing fruit color development.
Color_Quality is positioned as a direct subclass of Morphological_Quality, parallel to Size_Quality, Shape_Quality, Texture_Quality, and Structural_Integrity_Quality. Color qualities describe the optical appearance of organs and are distinct from the chemical compounds (pigments) that produce color, which are addressed under Chemical_Quality. The relationship between pigment concentration and perceived color is non-linear and affected by co-pigmentation, pH, and matrix effects. PATO_0000014 (color) provides the upper-ontology alignment. OIV descriptors address color characters for multiple organs including young shoot tips, mature leaves, and berries.
Farbeigenschaft
Farbqualität
Färbung
coloration
optical color property
pigmentation quality
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 369-403 He F et al. (2012) Anthocyanins and Their Variation in Red Wines I. Monomeric Anthocyanins and Their Color Expression. Molecules 17(2): 1571-1601
Color Quality
Color rim variation assessment in aged Barolo reveals dramatic core-to-rim differentiation with garnet-brick core transitioning through orange at mid-depth to pale tawny-brown at the watery rim, indicating 15+ years evolution and approaching optimal drinking window, contrasting with young Barolo showing consistent ruby from core to rim with only slight intensity fading at edge and no hue shift, demonstrating how rim assessment provides decade-scale age differentiation not visible in core color alone.
Color rim variation quality is a color appearance quality that represents the gradient of color characteristics from the wine's center (core) to its edge (rim) when viewed in a tilted glass, characterizing the transition in hue and intensity that provides age, concentration, and condition information not apparent from core color alone. Within the BFO framework, this quality specifically addresses the spatial variation dimension of color, distinguishing rim-to-core gradient assessment from single-point color evaluation while recognizing that the rim, where the wine layer is thinnest, reveals subtle color characteristics masked by intensity at the core. The ontological significance lies in rim variation providing the most sensitive age and condition indicator in visual assessment, where the thin wine layer at the rim reveals evolved pigments (orange-brown polymeric forms) that are visually overwhelmed by dominant monomeric anthocyanins at the deeper core, making rim color the leading edge of visible color evolution. The physical basis for rim variation involves the Beer-Lambert relationship where shorter path length at the rim produces less total absorption, allowing minor pigment components (polymeric pigments, oxidation products) to contribute proportionally more to visible color than at the core where dominant pigments saturate perception. The compositional information from rim assessment includes polymeric pigment proportion (orange-brown rim indicating evolved wine), oxidation state (browning at rim suggesting oxidative exposure), and color stability (rapid rim-to-core transition versus gradual gradient indicating different pigment profiles). The assessment protocol for rim variation involves tilting the glass approximately 45 degrees against white background to create a visible gradient from thick core to thin rim, with terminology describing rim color (watery, pink, ruby, orange, brown) and transition character (sharp versus gradual). The variety and age associations of rim variation show young wines of any variety displaying similar hue at rim and core while aged wines show pronounced rim-core differentiation, with rim reaching tawny-brown stages years before core color changes become apparent. Understanding rim variation quality enables sensitive age assessment from rim evolution, condition evaluation detecting early oxidation visible at rim before core, and maturity prediction based on rim-core differentiation pattern.
Color_Rim_Variation_Quality addresses the spatial gradient of color from core to rim, representing the most sensitive visual age indicator. This quality exploits the thin wine layer at the rim to reveal evolved pigments masked at the deeper core, providing early detection of color evolution and condition changes. The class supports age assessment, maturity prediction, and condition evaluation through rim-specific observation that complements core color assessment.
Farbverlauf
Kern-Rand-Gradient
Meniskusfarbe
Randvariation
core-to-rim gradient
edge color
meniscus variation
rim color
Peynaud E, Blouin J (1996) The Taste of Wine: The Art and Science of Wine Appreciation, 2nd Ed. John Wiley & Sons, New York, pp. 63-91 Glories Y (1984) La couleur des vins rouges. 1re partie: les équilibres des anthocyanes et des tanins. Connaissance de la Vigne et du Vin 18(3): 195-217
Color Rim Variation Quality
Color saturation comparison between properly cellared and heat-damaged bottles of same Barossa Shiraz reveals dramatically different saturation despite similar intensity and hue, with cellared example showing vivid, saturated purple-ruby (saturation rating 8.5/10) while heat-damaged example displays desaturated, brownish-tinged color (saturation 4/10) from pigment degradation during improper storage, demonstrating how saturation serves as a condition indicator that can reveal storage abuse not immediately apparent from intensity or hue alone.
Color saturation quality is a color appearance quality that represents the purity and vividness of wine color relative to a neutral gray of equivalent lightness, characterizing how brilliant and pure versus dull and muted the chromatic impression appears as an indicator of pigment condition, wine freshness, and overall vitality. Within the BFO framework, this quality specifically addresses the chromatic purity dimension of color, distinguishing saturation (color vividness) from intensity (color depth) and hue (color type) while recognizing that saturation reflects the integrity and stability of wine pigments rather than simply their concentration. The ontological significance lies in color saturation serving as a vitality and condition indicator, where highly saturated colors suggest fresh, well-preserved pigments and overall wine vitality while desaturated, muddy colors may indicate oxidative damage, pigment degradation, or production faults that diminish visual appeal and often correlate with aromatic and gustatory deficiencies. The perceptual basis for saturation involves the degree to which perceived color departs from achromatic gray toward pure spectral color, with highly saturated wine colors appearing vivid, clean, and gem-like while desaturated colors appear brownish, grayish, or muddy regardless of their basic hue. The compositional determinants of saturation include pigment molecular integrity (degraded pigments losing chromatic purity), copigmentation status (enhancing saturation through molecular stacking), oxidation state (creating brown quinones that desaturate color), and pH stability (fluctuations potentially causing pigment precipitation or transformation that reduces saturation). The distinction between saturation and intensity is important, as wines may be deeply colored but desaturated (dark but muddy) or pale but highly saturated (light but vivid), with both dimensions contributing independently to color quality assessment. Understanding color saturation quality enables condition assessment based on chromatic purity, vitality evaluation from saturation level, and quality inference where high saturation indicates well-preserved wine while low saturation may signal problems requiring investigation.
Color_Saturation_Quality addresses the chromatic purity dimension of wine color independent of intensity and hue. This quality serves as a condition and vitality indicator, with desaturation often signaling oxidative damage or pigment degradation. The class supports condition assessment and quality inference based on chromatic purity, complementing intensity and hue evaluation for comprehensive color assessment.
Farbbrillanz
Farblebendigkeit
Farbsättigung
chroma
chromatic vividness
chromatische Reinheit
color brilliance
color purity
Fairchild MD (2013) Color Appearance Models, 3rd Ed. John Wiley & Sons, Chichester, pp. 89-134 Pérez-Magariño S, González-San José ML (2003) Application of absorbance values used in wineries for estimating CIELAB parameters in red wines. Food Chemistry 81(2): 301-306
Color Saturation Quality
Color stability varies substantially across red V. vinifera cultivars, with stable-color types such as Tannat, Syrah, and Cabernet Sauvignon retaining 60-80% of initial color density after 2 years of bottle aging, while less stable cultivars such as Pinot Noir and Grenache may retain only 30-50%. Color stability depends on multiple factors including initial anthocyanin concentration (higher = more stable), acylation proportion (acylated forms more stable), copigmentation capacity (enhancing color expression), and polymeric pigment formation rate (creating SO₂-resistant color). The color density (A520 + A420) to anthocyanin concentration ratio increases during aging from 0.04-0.06 in young wines to 0.10-0.20 in aged wines as monomeric anthocyanins convert to polymeric forms. Wine pH strongly affects stability, with lower pH (3.2-3.4) favoring the colored flavylium form over colorless hemiketal and chalcone equilibrium species.
A quality potential disposition that is the inherited capacity of a Vitis vinifera plant to produce grapes yielding wines with stable red-purple color that resists fading during winemaking and aging, determined by anthocyanin concentration and profile, copigmentation capacity, and the formation of polymeric pigments through anthocyanin-tannin reactions that create color forms resistant to SO₂ bleaching and pH-induced equilibrium shifts.
Color_Stability_Disposition serves as the parent class for copigmentation and polymeric pigment formation traits. Color stability represents the practical outcome of multiple underlying chemical processes, integrating grape composition with wine chemistry evolution. The trait is critical for red wine quality, as color loss is perceived as a quality defect and limits aging potential. Climate change effects on grape anthocyanin accumulation (temperature inhibition) indirectly affect color stability through reduced initial pigment concentration.
Farbbeständigkeit
Farbstabilität
Pigmentstabilitäts-Disposition
color durability disposition
color retention capacity
pigment stability trait
Boulton R (2001) The copigmentation of anthocyanins and its role in the color of red wine: a critical review. American Journal of Enology and Viticulture 52:67-87 He F et al. (2012) Anthocyanins and their variation in red wines. I. Monomeric anthocyanins and their color expression. Molecules 17:1571-1601
Color Stability Disposition
Merlot grafted on SO4 rootstock showing columella cells in primary root cap, columella comprising central core 0.58 mm length organized as 6 longitudinal files each with 4-5 cells, columella cells 42 micrometers average diameter, microscopic examination revealing abundant amyloplasts 8.2 micrometers average diameter, iodine staining showing dense starch in columella amyloplasts, root reorientation experiment demonstrating amyloplast sedimentation toward new gravity vector within 12 minutes, columella cells providing gravitropic sensing directing primary root downward growth
Columella cells are specialized root cap cells forming central core region directly covering quiescent center and apical meristem, typically organized in 4-8 longitudinal files of 3-6 cells each creating columnar structure 0.3-0.8 mm length, characterized by large cells 30-60 micrometers diameter containing numerous starch-filled amyloplasts serving as statoliths, showing radial symmetry around root axis, demonstrating gravitropic sensing function through statolith sedimentation, and representing primary gravity perception cells in root system enabling positive gravitropism directing root growth downward, with columella cells exhibiting distinctive features including exceptionally large amyloplasts 5-15 micrometers diameter containing dense starch, amyloplast sedimentation toward gravity vector within 5-30 minutes of reorientation, endoplasmic reticulum and actin cytoskeleton interacting with sedimenting amyloplasts potentially transducing mechanical signal, and auxin transport machinery responding to gravity signal redistributing hormone to create differential growth, and demonstrating that columella statolith sedimentation represents first step in gravitropic response cascade converting physical gravity stimulus into biochemical and growth responses. Columella cells structural organization includes= showing columnar arrangement (columella cells organized in longitudinal files creating column-like structure), exhibiting file organization (typically 4-8 files radiating from root center each containing 3-6 cells), demonstrating central position (columella occupying root apex center directly over quiescent center and meristem), displaying radial symmetry (columella files arranged symmetrically around root axis), and revealing tiered structure (cells within each file arranged in tiers progressing from young near meristem to older toward cap periphery). Columella cells cellular characteristics includes= showing large size (columella cells typically 30-60 micrometers substantially larger than meristem cells), exhibiting large central vacuole (mature columella cells with prominent central vacuole occupying 60-80% cell volume), demonstrating abundant amyloplasts (columella cells containing 20-100+ starch-filled plastids compared to few in other cap cells), revealing starch accumulation (amyloplasts packed with starch grains creating dense structures), and displaying cytoplasmic polarity (cytoplasm and organelles showing asymmetric distribution related to gravity sensing). Columella cells amyloplast characteristics includes= showing large size (columella amyloplasts typically 5-15 micrometers diameter among largest plant plastids), exhibiting high density (starch-filled amyloplasts with density 1.4-1.5 g/cm³ enabling sedimentation), demonstrating starch composition (amyloplasts containing 70-90% starch by weight creating gravitational mass), revealing mobility (amyloplasts capable of moving within cell in response to gravity or other forces), and displaying sedimentation capacity (amyloplasts settling toward gravity vector with velocity 0.1-1 micrometer per minute). Columella cells gravitropic sensing mechanism includes= showing statolith sedimentation (amyloplasts sedimenting toward bottom of cell in gravity field), exhibiting rapid response (amyloplast repositioning detectable within 5-10 minutes of root reorientation), demonstrating mechanical signal (sedimenting amyloplasts potentially generating tension on ER or cytoskeleton creating mechanotransduction), revealing calcium signaling (gravity stimulus triggering calcium influx or release potentially part of signal transduction), and displaying auxin redistribution (gravity signal ultimately causing asymmetric auxin transport creating differential growth). Columella cells in gravitropic response includes= providing gravity perception (columella cells serving as primary gravity sensors in root detecting direction of gravity vector), exhibiting signal generation (statolith sedimentation in columella initiating signal transduction cascade), demonstrating signal transmission (columella generating signal transmitted to elongation zone where growth response occurs), revealing auxin involvement (columella sensing gravity triggering asymmetric auxin distribution to lower versus upper root side), and displaying curvature production (auxin asymmetry causing differential elongation creating downward root curvature). Columella cells development and maintenance includes= showing origin from meristem (columella cells produced by root cap initials near quiescent center), exhibiting maturation gradient (youngest columella cells near meristem progressively maturing distally), demonstrating starch accumulation (amyloplasts accumulating starch as columella cells mature), revealing limited lifespan (individual columella cells functioning days to weeks before replaced), and displaying continuous renewal (columella population maintained through ongoing production despite turnover). Columella cells experimental evidence includes= showing starch staining (iodine staining revealing dense starch in columella amyloplasts documenting statolith presence), exhibiting sedimentation observation (microscopy documenting amyloplast movement in response to root reorientation), demonstrating ablation effects (laser or surgical ablation of columella eliminating gravitropism confirming necessity), revealing starch mutants (mutants lacking columella starch showing reduced or absent gravitropism supporting statolith role), and displaying clinostat experiments (roots on clinostat eliminating net gravity vector showing random growth confirming gravity dependence). Columella cells species variation includes= showing universal presence (columella cells present in essentially all seed plant roots reflecting fundamental importance), exhibiting size variation (columella cell and amyloplast size varying among species though general organization conserved), demonstrating number variation (columella file number and cell number per file varying with root diameter and species), revealing starch density variation (columella amyloplast starch content varying somewhat among species), and displaying functional conservation (despite morphological variation gravitropic function showing remarkable conservation across species).
Columella_Cells emphasizes statolith-containing central cap cells and is distinguished from lateral cap by focus on gravity sensing function. Includes Central_Columella_Cells and Gravity_Sensing_Columella_Cells. Critical for understanding gravitropic perception, statolith amyloplasts, columnar organization, sedimentation mechanism, signal transduction, and the specialized gravity-sensing cells in root cap center detecting gravity direction enabling positive gravitropism.
Central cap cells
Columella-Zellen
Gravitations-Sensorzellen
Gravity-sensing cells
Statolith-containing cells
Statolithen-Zellen
Barlow, P. W. (2003). The root cap= Cell dynamics, cell differentiation and cap function. Journal of Plant Growth Regulation, 21(4), 261-286. | Morita, M. T. (2010). Directional gravity sensing in gravitropism. Annual Review of Plant Biology, 61, 705-720.
Columella Cells
The cultivar with VIVC prime name Pinot Gris is marketed under the commercial synonym Pinot Grigio in export markets targeting the United States and United Kingdom where Italian wine terminology carries strong brand recognition, with the name Pinot Grigio implying a lighter, crisper wine style despite referring to the same genotype — similarly, Shiraz functions as a commercial synonym for Syrah in Australian and South African markets, and Fumé Blanc was coined by Robert Mondavi in 1968 as a commercial synonym for Sauvignon Blanc to evoke Pouilly-Fumé associations and differentiate his barrel-fermented style from standard Sauvignon Blanc.
A synonym that was created or adopted for commercial purposes to enhance marketability, comply with labeling regulations in specific export markets, or differentiate a wine product through the use of an alternative cultivar name perceived as more appealing, prestigious, or recognizable to target consumers.
Commercial_Synonym captures the market-driven dimension of cultivar naming where names are strategically selected or created to influence consumer perception and purchasing behavior. Unlike other synonym categories that arise from historical, geographic, or linguistic processes, commercial synonyms are deliberately chosen for their marketing value: the Syrah/Shiraz distinction exemplifies how the same genotype can be positioned differently in the marketplace through synonym selection — Syrah evoking French Rhône Valley prestige, Shiraz signaling Australian bold fruit-forward style. Some commercial synonyms have acquired regulatory recognition (Shiraz is an accepted synonym in Australian wine law) while others remain informal marketing conventions without official nomenclatural status.
Handelsname
Vermarktungsbezeichnung
commercial variety name
kommerzielles Synonym
marketing synonym
trade synonym
Robinson J et al. (2012) Wine Grapes, Allen Lane pp: 1-18 Anderson K and Aryal NR (2013) Which Winegrape Varieties are Grown Where?, University of Adelaide Press OIV (2013) OIV List of Vine Varieties and Their Synonyms
Commercial Synonym
Chardonnay vines in the Riverina region of New South Wales bearing a commercial wine role are mechanically pruned and harvested at yields of 15-25 t/ha, irrigated via drip systems delivering 4-6 ML/ha/season, managed with calendar-based spray programs minimizing per-hectare labor costs, and producing fruit at $200-400 AUD/tonne for brands retailing at $8-12 AUD/bottle in Australian supermarket channels, with annual production volumes of 50,000-200,000 cases per label ensuring consistent national distribution.
A quality tier role that inheres in a grapevine whose fruit is produced for wines targeting the high-volume everyday consumption segment, managed with cost-efficient viticultural practices emphasizing consistent yields and reliable quality at accessible price points, without the vineyard-specific selectivity or intensive management characteristic of premium and higher tiers.
Commercial_Wine_Role encompasses the volume segment that constitutes the majority of global wine consumption by volume, typically positioned at $5-10 USD retail in the US market. The tier is characterized by large-scale production, mechanical vineyard operations, warm-climate sourcing from high-yielding regions, and brand-driven rather than terroir-driven marketing strategies: viticultural decisions at this tier prioritize yield consistency, disease management efficiency, and cost minimization over the site-specific quality optimization pursued at higher tiers. Major commercial wine sourcing regions include Riverina and Riverland in Australia, the Central Valley in California, La Mancha in Spain, and the Languedoc in France.
Alltagswein-Rolle
Gebrauchswein-Rolle
Konsumwein-Rolle
commercial wine vine role
everyday wine role
popular premium role
Anderson K (2004) The World's Wine Markets: Globalization at Work, Edward Elgar Publishing pp: 38-62 McMillan R (2022) State of the US Wine Industry, Silicon Valley Bank OIV (2023) State of the World Vine and Wine Sector
Commercial Wine Role
Pinot Noir exemplifies compact cluster disposition with 10-15 berries/cm rachis, OIV 204 scores of 7-9, and characteristic spherical berry deformation creating polygonal cross-sections in cluster interiors. Riesling develops moderately compact clusters (8-11 berries/cm) that vary with clone selection, with loose clones achieving OIV scores of 5 versus compact clones at 7-8. Compact clusters in Pinot Noir show Botrytis infection rates 3-5× higher than loose-clustered cultivars under identical humidity conditions. Berry skin cracking at contact points occurs in 10-25% of berries in severely compact clusters during rapid post-veraison expansion.
A cluster compactness disposition that is the inherited capacity of a Vitis vinifera plant to develop tightly packed clusters where berries are in close contact and often deformed by mutual pressure, with high berry density per unit rachis length (typically greater than 8 berries/cm) and limited air circulation within the cluster interior.
Compact cluster disposition results from the combination of high fruit set, large berry size relative to rachis framework, and short pedicels. The trait increases bunch rot susceptibility through impaired spray penetration, elevated interior humidity, and berry skin damage at contact points. Clone selection within compact cultivars can identify loose variants, with Pinot Noir clones ranging from OIV 5 to OIV 9. Pre-bloom gibberellin application reduces compactness through rachis elongation and berry thinning.
dense bunch disposition
dichte Traube
eng gepackte Traubenstruktur
high-density cluster
kompakte Traube
tight cluster trait
Vail ME, Marois JJ (1991) Grape cluster architecture and the susceptibility of berries to Botrytis cinerea. Phytopathology 81:188-191 Hed B et al. (2009) Relationship between cluster compactness and bunch rot in Vignoles grapes. Plant Disease 93:1195-1201
Compact Cluster Disposition
Pinot Noir in Burgundy's Côte d'Or at BBCH 65 produces compact flower clusters of 8-11 cm length bearing 180-320 flowers with minimal secondary branching, exhibiting flower densities of 22-28 flowers/cm and creating tight aggregations (OIV code 7-8 at flowering, persisting to berry stage) requiring 6-8 fungicide applications for Botrytis management. Müller-Thurgau in Germany's Mosel Valley shows extremely compact flower clusters (OIV code 9) with 200-400 flowers packed into 7-10 cm length, necessitating aggressive cluster thinning at BBCH 71-73 to reduce final compactness and disease risk. Chardonnay in Champagne exhibits moderately compact flower clusters (OIV code 6-7) with controlled flower number (150-250) maintained through crop load management during previous-season bud differentiation.
A compact flower cluster is a flower cluster characterized by high flower density resulting from numerous flowers borne on abbreviated inflorescence axes with short internodal spacing, creating tight aggregations that restrict air circulation and promote conditions conducive to fungal disease development while facilitating complete pollination through close proximity of flowers. Compact flower clusters exhibit flower densities typically exceeding 18-25 flowers per cm of total inflorescence axis length (rachis plus all branch lengths combined), achieved through combinations of high flower initiation during primordium development, reduced rachis and branch elongation, and short pedicel length. These clusters typically measure 6-14 cm in length at BBCH 65 (full bloom) while bearing 150-600 flowers, creating length-normalized densities 2-4× those of loose flower clusters. The compact architecture results from genetic programming for reduced internode elongation during inflorescence development, often combined with reduced or absent secondary and tertiary branching that concentrates flowers on the main rachis and primary branches. Compact flower clusters characteristically show cylindrical or conical shapes with length=width ratios of 2=1-4=1, lacking the lateral lobes or wings characteristic of loose clusters with extensive branching. Flower spacing within compact clusters typically ranges from 1-3 mm between adjacent flower pedicels, compared to 4-12 mm in loose clusters, creating nearly continuous flower coverage along the axis. The tight flower arrangement creates a humid microclimate within the cluster even under open canopy conditions, with relative humidity often 10-20% higher within compact clusters than in ambient air, and drying times after rain or dew events extended by 2-6 hours compared to loose clusters. This microclimate profoundly affects disease dynamics, with compact flower clusters showing 3-8× higher Botrytis cinerea infection rates at bloom compared to loose clusters under the same environmental conditions and fungicide programs. Paradoxically, the close flower proximity in compact clusters can enhance pollination success by increasing probability of pollen capture, potentially offsetting disease disadvantages in dry climates where Botrytis pressure is minimal. Compact flower cluster morphology shows high heritability, with cultivars like Gewürztraminer, Müller-Thurgau, Pinot Noir, Chenin Blanc, and Chardonnay consistently producing compact clusters (OIV codes 7-9) across diverse environments, though water stress during the previous season's primordium differentiation can further exacerbate compactness. The transformation from compact flower cluster to compact berry cluster involves relatively minimal architectural change, as the inherent tight spacing persists through fruit set and development, with berry-berry contact occurring early (BBCH 73-75) and berry deformation from mutual compression evident by véraison.
Compact_Flower_Cluster is distinguished from Loose_Flower_Cluster by quantitative density metrics (>18-25 flowers/cm versus <8-12 flowers/cm), qualitative architectural features (reduced branching, short spacing), and functional consequences (elevated disease risk, enhanced pollination). The boundary between 'compact' and 'medium' density is necessarily somewhat arbitrary but conventionally corresponds to OIV codes 6-9 versus codes 4-5. Compact flower clusters represent a significant viticultural challenge in humid climates, driving substantial fungicide costs and quality losses, but may confer advantages in arid regions where disease pressure is minimal and pollination reliability is critical. Sub-classification into Densely_Packed and Tightly_Compact reflects gradations within the compact category. The degree of compactness at flowering serves as a critical decision point for cluster management interventions (thinning, leaf removal, berry removal) to modify harvest-stage architecture.
dense flower cluster
dichte Blütentraube
gedrängte Blütentraube
kompakte Blütentraube
tight flower bunch
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 95-98, 145-148 OIV (2009). Descriptor List for Grape Varieties and Vitis Species, OIV code 204 Vail, M.E., Marois, J.J. (1991). Grape cluster architecture and the susceptibility of berries to Botrytis cinerea. Phytopathology 81=188-191 Hed, B. et al. (2009). Evaluation of cultural practices for management of Botrytis bunch rot. Plant Disease 93=1148-1154.
Compact Flower Cluster
Compatibility determines the success of grafted vine production, with graft take rates ranging from 60-98% depending on scion-rootstock combinations under commercial bench grafting conditions. Compatible combinations such as Cabernet Sauvignon on SO4 achieve 90-95% take rates with strong vascular connections, while marginally compatible combinations may show 70-80% initial success with subsequent decline. Interspecific compatibility enables rootstock breeding, with V. vinifera × V. rupestris and V. berlandieri × V. riparia crosses showing high fertility (>80% viable seed) versus difficult crosses such as V. vinifera × V. rotundifolia (<5% viable seed). Long-term compatibility affects vineyard longevity, with some scion-rootstock combinations maintaining productivity for 50-100 years while incompatible unions decline within 10-20 years.
A grapevine disposition that is the inherited capacity of a Vitis vinifera plant to form successful biological unions with other grapevine genotypes through grafting or to produce viable offspring through interspecific hybridization, encompassing traits related to graft union formation, long-term graft functionality, and cross-compatibility with related Vitis species.
Compatibility_Disposition serves as the top-level parent class for traits related to graft union success and interspecific hybridization. The category addresses both the immediate success of graft union formation and the long-term physiological functionality of the grafted vine. Compatibility is fundamental to commercial viticulture, as phylloxera pressure requires grafting onto resistant rootstocks throughout most wine regions. Understanding compatibility mechanisms enables informed rootstock selection and improved nursery practices.
Kompatibilität
Vereinigungsfähigkeit
Verträglichkeit
biological integration disposition
grafting success capacity
union compatibility trait
Ollat N et al. (2016) Grapevine roots and soil environment. In: Gerós H et al. (eds) Grapevine in a Changing Environment, Wiley pp.: 175-202 Goldschmidt EE (2014) Plant grafting: new mechanisms, evolutionary implications. Frontiers in Plant Science 5:727
Compatibility Disposition
In Gewürztraminer vines in Alsace, competence achievement is confirmed in early February when forcing tests show budbreak within 10 days at 22°C with 98% success rate, ABA concentrations have declined to 55 ng per gram fresh weight, and transcript analysis confirms minimal VvDAM expression with restored CYCLIN D expression potential, indicating full restoration of growth capacity following completion of chilling requirement at approximately 1100 hours.
Competence achievement in Vitis vinifera represents the successful restoration of full growth capacity following completion of chilling-dependent dormancy release processes, describing the state in which buds have regained complete ability to respond to growth-promoting conditions after the internal constraints of endodormancy have been fully dismantled. This achievement marks the functional completion of dormancy release from the perspective of bud capability, establishing the growth-ready state that will enable rapid response to subsequent heat accumulation and eventual budbreak initiation. The physiological basis of competence achievement involves the completion of all cellular and molecular changes required for growth capacity, including restoration of cell cycle competence through cyclin and CDK expression potential, re-establishment of hormone signaling sensitivity enabling response to growth-promoting signals, restoration of metabolic enzyme systems capable of supporting biosynthesis, and recovery of symplastic communication enabling coordinated growth responses. The verification of competence achievement relies on forcing tests that demonstrate the capacity for rapid and complete budbreak, with achievement confirmed when forcing response times have decreased to minimum values of approximately 10-14 days and budbreak percentages have reached maximum levels of greater than 95% under optimal conditions. The molecular markers of competence achievement include ABA concentrations at baseline levels below approximately 60-80 ng per gram fresh weight, DAM gene expression at minimal or undetectable levels, and growth gene promoters showing full transcriptional responsiveness to activating signals. The timing of competence achievement coincides with chilling requirement satisfaction, occurring during late January to February for most cultivars in Northern Hemisphere viticulture, and marking the transition from the internally-constrained state of endodormancy to the environmentally-constrained state of ecodormancy. The uniformity of competence achievement across buds on a vine affects subsequent budbreak synchrony, with simultaneous achievement of competence at all bud positions promoting uniform budbreak while variable achievement may contribute to extended or uneven budbreak patterns.
Competence_Achievement represents the functional capacity outcome of Chilling_Completion describing restored growth ability. Distinguished from Dormancy_Release_Competence during endodormancy by representing verified complete competence. Confirmed by forcing response less than 14 days with greater than 95% success. Molecular markers include baseline ABA, minimal DAM, and restored growth gene responsiveness. Timing coincides with chilling requirement satisfaction. Uniformity across buds affects subsequent budbreak synchrony.
Kompetenz-Erreichung
Wachstumsfähigkeits-Erreichung
Wachstumskompetenz-Erlangung
competence restoration completion
full competence attainment
growth capacity achievement
growth competence achievement
vollständige Kompetenz-Wiederherstellung
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Rohde A, Bhalerao RP (2007) Plant dormancy in the perennial context. Trends in Plant Science 12: 217-223 Cooke JE, Eriksson ME, Junttila O (2012) The dynamic nature of bud dormancy in trees: environmental control and molecular mechanisms. Plant Cell and Environment 35: 1707-1728
Competence Achievement
Cabernet Sauvignon in Napa Valley shows complete anthesis during late June-early July Year N+1 at BBCH 69 on approximately June 16, with all viable flowers opened, visible berry development 5-8 mm diameter throughout cluster, unfertilized flowers showing brown senescence and progressive abscission, established fruit set of 55-68% representing 380-520 berries per cluster, under warm climate conditions of 24-29°C temperature transitioning from reproductive vulnerability to developmental stability, typical of this late-season cultivar definitively completing flowering 12-14 days after first flowers opened
Complete anthesis represents the definitive flowering completion state when all viable flowers have opened, reproductive processes have concluded, fruit set outcomes are established, and cluster transitions fully to developmental phase with visible berry growth and progressive unfertilized flower abscission. This conclusive reproductive milestone occurs during late June-early July in temperate regions Year N+1 following late anthesis, typically representing 1-2 day period marking transition from BBCH 68 to BBCH 69 as reproductive phase definitively concludes. Completion characteristics show all potentially viable flowers having completed opening with 100% of developmentally capable flowers showing exposed reproductive organs or developed berries, remaining unopened structures representing aberrant or aborted buds incapable of normal development, and overall cluster showing clear transition from reproductive to developmental state. Flower status distribution at completion shows successfully fertilized flowers as developing berries 4-8 mm diameter creating visible green fruit structures distributed throughout cluster, unfertilized flowers as senescing brown structures showing progressive abscission, occasional final-opening flowers showing minimal reproductive potential, and overall reproductive activity effectively ceased. Berry visibility at completion shows clear fruit structures distinguishable throughout cluster with small green berries easily identifiable on pedicels, berry number countable providing definitive fruit set assessment, spatial distribution revealing architectural pattern and potential density issues, and overall appearance of young fruit cluster rather than flower cluster. Abscission activity intensifies at completion with unfertilized flowers progressively detaching through completed abscission zone development, daily flower drop creating accumulation of fallen flowers on cluster surface and ground, and progressive cluster thinning as failed reproductions eliminate leaving only successful fruit set berries. Fruit set percentage determination at completion enables accurate assessment with berry count relative to total flowers opened calculating set percentage typically 30-75% depending on cultivar and conditions, comparison to potential flower number from previous season primordium formation evaluating reproductive success, and yield prediction based on berry number and expected final berry size. Cluster appearance at completion shows characteristic developmental state with small green berries dominating visual appearance, brown senescing unfertilized flowers interspersed though progressively abscising, cluster showing open architecture with berries spaced on pedicels and branches, and overall healthy developmental appearance contrasting with earlier reproductive vulnerability. Developmental phase transition at completion marks shift from reproductive biology to fruit development with metabolic priorities focusing on berry cell division and expansion, resource allocation directing photosynthate to rapidly growing berries as primary sink, and hormonal environment dominated by developmental signals from expanding berries rather than reproductive coordination. Berry development stage at completion shows early cell division phase with rapid cell proliferation establishing cell number potential typically 8-15 days duration, cells dividing rapidly 12-24 hour cycles creating berry cell population, and berry size increasing primarily through cell number increase during this early stage. Berry growth rate at completion shows rapid expansion 0.3-0.6 mm diameter per day driven primarily by cell division with modest cell expansion, water uptake supporting turgor maintaining expansion, and substantial resource demand requiring adequate photosynthetic capacity and favorable environmental conditions. Berry structure at completion includes developing pericarp with outer skin layers, inner flesh tissues undergoing cell proliferation, and seed development progressing with embryo and endosperm growth following successful fertilization, establishing anatomical framework for berry maturation. Seed development at completion shows early embryo development following fertilization with embryo cells dividing rapidly, endosperm development providing nutritive tissue supporting embryo growth, and integument layers developing into seed coat, with seed development coordinated with berry pericarp growth through hormonal signals. Resource demand at completion increases substantially with developing berries creating major sink competing with vegetative growth for available resources, requiring adequate photosynthetic production from expanding canopy, supplemented by stored reserves particularly during early rapid cell division, and balanced resource allocation supporting both fruit and vegetative development. Canopy development at completion shows concurrent vegetative growth with shoots typically having 15-20 leaves providing substantial photosynthetic capacity, continued shoot elongation though slowing from earlier rapid growth, tendril attachment supporting shoot positioning, and balanced vegetative-reproductive development supporting fruit growth without excessive vigor. Environmental effects at completion show reduced vulnerability compared to flowering with adverse weather having minimal effect on established berry development at this early stage, though extreme conditions like hail or severe wind potentially causing physical damage, and favorable conditions supporting optimal berry growth rates. Temperature at completion optimally 18-28°C supporting rapid berry cell division and growth, with cool conditions below 15°C slowing development and warm conditions above 32°C potentially causing stress effects, and overall temperature accumulation driving developmental rate toward subsequent berry stages. Water availability at completion becomes important for berry development with adequate moisture supporting cell division and expansion though berries remain relatively small with modest water demands, and beginning establishment of irrigation management supporting berry development through season. Nutrient requirements at completion emphasize nitrogen for protein synthesis supporting rapid cell division in berries and continued vegetative growth, potassium beginning to show importance for berry development affecting later quality outcomes, and balanced nutrition supporting healthy development. Disease management at completion transitions from flowering restrictions to active berry protection with fungicide applications resuming for botrytis and powdery mildew control, focus shifting to protecting developing berries from infection, and recognition that flowering-period infections may remain latent requiring continued vigilance. Canopy management at completion enables resumption of operations with shoot positioning optimizing berry exposure and air circulation, possible hedging or trimming managing excessive vegetative growth, leaf removal potentially implemented in some systems exposing fruit zone, and overall canopy optimization supporting berry development and disease management. Phenological significance of completion includes marking definitive end of BBCH 69 flowering completion stage, transitioning to BBCH 71-79 fruit development stages, establishing reference point for subsequent development predictions, and confirming reproductive phase conclusion enabling developmental focus. The process shows three distinguishable aspects progressing from all flowers opened establishing complete cluster flowering, through anthesis completion confirming reproductive conclusion, to post-anthesis transition marking shift toward fruit development. Duration represents brief 1-2 day milestone period marking definitive completion, with rapid progression toward fruit development phase. Cultivar variation in completion timing shows early varieties completing late June while late varieties complete early-mid July reflecting overall phenological differences, completion clarity with some showing clean definitive completion while others show extended gradual completion, and fruit set outcomes ranging from poor 30-45% in difficult-setting varieties to excellent 60-75% in good-setting types. Regional timing shows complete anthesis during late June in warm climates at approximately 1150-1200 growing degree days base 10°C, late June-early July in temperate regions at 1050-1100 GDD, and early-mid July in cool areas at 950-1000 GDD, marking transition to fruit development phase. Viticultural significance includes establishing definitive flowering completion enabling management transitions, confirming fruit set outcomes and berry number determining yield potential, marking end of critical weather-vulnerable flowering period reducing risk, providing reference for developmental phase management and timing, and establishing baseline for subsequent phenological predictions through season. Research applications include documenting flowering completion timing for phenological studies, analyzing fruit set determinants affecting yield outcomes, evaluating berry development initiation and early growth patterns, quantifying resource allocation transitions from reproduction to development, and assessing management effects on flowering outcomes and early berry development.
Complete_Anthesis represents definitive flowering completion milestone marking reproductive phase conclusion. Follows Late_Anthesis. Corresponds to BBCH 69. Subdivided into All_Flowers_Opened, Anthesis_Completion, and Post_Anthesis_Transition. Establishes final fruit set and berry number. Marks transition from reproductive vulnerability to developmental phase. Enables management transitions and establishes reference for subsequent phenological predictions. Critical milestone in annual phenological progression.
Anthesevollendung
Blüheende
Blütenabschluss
anthesis conclusion
bloom completion
end of flowering
flowering completion
vollständige Anthese
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 388-395 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 298-305 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Complete Anthesis
In a Bordeaux Merlot vineyard, complete berry softening was documented in early October Year N+1 (BBCH 85-87) when firmness stabilized at 1.5-1.7 N after declining from 5.2 N at green berry stage, soluble pectin reached 76-78% of total, sugar content plateaued at 23.5-24.0° Brix, titratable acidity stabilized at 5.2-5.4 g/L, skin anthocyanins achieved 1850-1950 mg/kg, and sensory evaluation indicated optimal flavor development suitable for harvest under typical autumn conditions (daily high 18-22°C, nighttime 10-14°C).
Complete_Berry_Softening represents the final stage of berry texture transformation during late veraison through early ripening when firmness decline approaches asymptotic minimum values and berries achieve their characteristic ripe texture suitable for harvest or further maturation. This terminal softening phase typically extends 1-2 weeks (BBCH 85-87) following peak softening and is characterized by berry firmness stabilizing at cultivar-specific final values: soft-fruited varieties like Pinot Noir and Gamay reach 0.8-1.5 N compression firmness representing 70-80% total decline from green berry values (3.0-5.5 N), medium-firmness cultivars like Merlot and Syrah stabilize at 1.2-2.0 N indicating 60-70% reduction from initial firmness (3.5-6.0 N), while firm-fruited varieties like Cabernet Sauvignon and Nebbiolo maintain 1.5-2.5 N firmness showing 50-65% decline from green berry baseline (4.0-6.5 N). The completion of softening involves establishment of final cell wall architecture: soluble pectin reaches stable maximum values of 70-85% total pectin (from 20-30% in green berries) with minimal further increase, pectin molecular weight stabilizes at cultivar-specific minima ranging 60-120 kDa (from 180-350 kDa pre-veraison), degree of methylesterification plateaus at 25-40% (from 65-80% initially), and hemicellulose modification reaches completion with xyloglucan molecular weight and structure stabilized at final degraded states. Enzyme activities during complete berry softening transition from peak to basal maintenance levels: polygalacturonase declines from maximum values (120-220 units) to sustained moderate activity (40-80 units) sufficient to maintain cell wall state but not drive further major degradation, pectin methylesterase similarly reduces from 110-200 units to 35-70 units, while other cell wall-modifying enzymes including xyloglucan endotransglycosylase, β-galactosidase, and expansins show parallel activity reductions to 30-50% of peak values maintaining cell wall plasticity without continued extensive breakdown. Cellular mechanical properties at complete softening reflect final berry texture: cell wall thickness stabilizes at 1.5-2.8 μm (reduced from 2.5-4.5 μm green berry values) with no further thinning, turgor pressure reaches final low-moderate levels 0.3-0.6 MPa (compared to 0.6-1.0 MPa in firm berries) contributing 15-25% to residual firmness while cell wall structural integrity provides 75-85% of remaining mechanical resistance, and skin tensile strength plateaus at 40-65% of green berry values creating the characteristic ripe berry vulnerability to compression and puncture. Complete berry softening coincides with other ripening process completions: sugar accumulation typically reaches plateau (20-28° Brix depending on cultivar and style goals) within 1-2 weeks of softening completion, acid levels stabilize at final values (4-8 g/L titratable acidity) as malate degradation slows substantially, anthocyanin accumulation in red cultivars approaches maximum (500-3000 mg/kg skin depending on variety and conditions), and aromatic compound development reaches characteristic ripe profiles suitable for harvest. Gene expression patterns at complete softening show transition from ripening peak to maintenance phase: cell wall enzyme genes decline from maximum expression (100-200 fold above green berry) to moderate sustained levels (20-50 fold), sugar transporter genes reduce from peak expression as accumulation slows, while genes involved in secondary metabolism including phenylpropanoid pathway components and volatile biosynthesis enzymes reach maximum expression supporting final quality compound synthesis. Hormonal regulation during complete softening reflects ripening program consolidation: abscisic acid levels remain elevated (800-2000 ng/g fresh weight) but decline from veraison peak values (1500-3500 ng/g) as major developmental transitions complete, ethylene production shows cultivar-dependent patterns with some varieties maintaining slight elevation (0.3-1.2 nL g⁻¹ h⁻¹) while others return to baseline, and auxin levels decline to minimal values reflecting completion of growth-related processes. Temperature effects on achieving complete softening show reduced sensitivity compared to earlier phases: once major structural transformations occur during peak softening, temperature primarily affects the timing of reaching final state rather than final firmness values themselves, with warmer conditions (25-30°C) accelerating approach to completion by 3-7 days compared to cooler temperatures (18-23°C), but final firmness values differing by only 10-20% across normal temperature ranges. Cultivar variation in complete softening characteristics determines varietal typicity: Pinot Noir achieves very soft final texture (0.9-1.4 N) with thin skins and delicate berry structure requiring careful handling, Cabernet Sauvignon maintains relatively firmer completion (1.6-2.4 N) with thicker skins and more robust cell walls tolerating mechanical harvest, and intermediate varieties like Merlot show balanced completion (1.3-1.9 N) suitable for varied viticultural and harvest approaches. Viticultural implications of complete berry softening inclu harvest timing decisions based on achieving desired firmness for intended wine style (sparkling wines may harvest before completion, while late-harvest wines extend beyond completion allowing slight over-ripening), mechanical harvest feasibility assessment since excessively soft berries (< 1.0 N) may suffer unacceptable damage requiring hand harvest, and disease management intensification as fully softened berries with minimal skin strength show maximum vulnerability to Botrytis infection and other pathogens requiring protective measures. Berry quality at complete softening represents the integration of all ripening processes: optimal harvest typically occurs when softening completion coincides with target sugar levels, desired acidity, complete flavor and aroma development, and phenolic maturity producing balanced grapes suitable for quality wine production, while discordant ripening with premature softening completion before chemical maturity or delayed softening past optimal chemical composition creates quality challenges requiring viticultural decisions balancing texture and composition priorities. Environmental stress effects on complete softening achievement inclu water deficit during late ripening can maintain slightly firmer final texture through mild turgor maintenance and reduced cell wall enzyme activity potentially beneficial for varieties prone to excessive softness, heat stress may accelerate softening completion but risks premature berry dehydration and shriveling before optimal flavor development, and cool conditions can delay completion requiring extended hang time that may improve complexity but increases disease pressure and harvest logistics challenges. Monitoring complete berry softening through regular firmness testing every 3-5 days during late veraison and early ripening combined with chemical maturity assessment (sugar, acid, pH) and sensory evaluation (flavor, tannin) enables precise harvest timing decisions optimizing berry quality for specific wine styles and market requirements.
Complete_Berry_Softening is distinguished from Peak_Softening by representing final state achievement rather than maximum transformation velocity, from Progressive_Softening by exhibiting stabilization rather than continued moderate decline, and from earlier ripening phases by indicating readiness for harvest assessment. Duration typically 1-2 weeks during late veraison to early ripening (BBCH 85-87). Final firmness values: soft cultivars 0.8-1.5 N, medium 1.2-2.0 N, firm varieties 1.5-2.5 N. Enzyme activities decline to 30-50% of peak maintaining cell wall state. Complete_Berry_Softening achievement typically signals optimal harvest window approach requiring integrated assessment of firmness, chemistry, and sensory characteristics.
Reifungs-Vollendungs-Textur
final berry texture achievement
finale Beeren-Textur-Erreichung
mature berry firmness
reife Beeren-Festigkeit
ripening completion texture
terminal softening state
terminaler Erweichungs-Zustand
Keller M (2020) The Science of Grapevines 3rd Ed pages 232-235 berry maturation and harvest criteria Coombe BG (1992) Research on development and ripening of the grape berry Australian J Grape Wine Res 48:125-144 berry ripening completion Rolle L, Torchio F, Zeppa G, Gerbi V (2009) Relationship between skin break force and anthocyanin extractability at different ripening stages Amer J Enol Vitic 60:93-97 firmness and maturity correlation.
Complete Berry Softening
Tempranillo in Rioja completes bud breaking at BBCH 09 in early May after 1100-1250 chilling hours and 420-480 growing degree days base 10°C, with shoot measuring 20-25 mm exposed length and first leaves spread 15-20 mm from axis over 3-4 days under conditions of 18-22°C mean temperature, 40-50% relative humidity, and 14.5-hour photoperiod, representing typical continental climate timing for this mid-season cultivar
Complete bud breaking represents the final emergence phase where first true leaves separate fully from shoot axis achieving independent configuration and establishing the transition from breaking to shoot growth, corresponding to BBCH 09 as the definitive conclusion of bud burst process. This culminating developmental stage occurs when shoot has extended sufficiently that first leaves no longer remain appressed to axis but instead spread perpendicular establishing characteristic grapevine leaf orientation and beginning substantial photosynthetic contribution. The shoot typically measures 15-30 mm exposed length with apical meristem continuing active cell production while elongation zone cells complete expansion creating visible daily growth increments of 3-7 mm under favorable conditions. First true leaves expand to 10-20 mm length with leaflets of palmate leaf structure clearly separated from each other, creating recognizable grapevine leaf form though still representing juvenile configuration smaller than mature leaf dimensions. Leaf surfaces show bright green coloration indicating substantial chlorophyll accumulation, with stomata becoming functional enabling gas exchange for photosynthesis and transpiration though stomatal density and control mechanisms are not yet fully mature. Protective bud scales have fallen away completely or remain only as dry brown remnants at shoot base, no longer providing any protective function as shoot and leaves are fully exposed to environmental conditions. The apical meristem maintains high cell division rates producing new cells for continued shoot elongation and initiating additional leaf and tendril primordia, with meristem structure showing characteristic layered organization of tunica and corpus zones producing organs in regular phyllotactic pattern. Vascular system extends throughout exposed shoot length with functional xylem providing water transport capacity of 1-3 mg per second supporting expanding tissues and increasing transpiration from developing leaf area, while phloem continues transporting stored reserves though beginning to carry photosynthate from expanding leaves. Chloroplasts in leaf cells achieve substantial though not complete development with thylakoid membranes organized for light capture and Calvin cycle enzymes assembled for carbon fixation, enabling photosynthetic rates reaching 40-50% of mature leaf capacity. The metabolic balance transitions toward net positive carbon fixation as photosynthetic rate in expanding leaves begins approaching and occasionally exceeding respiration rate, though shoots remain net carbon consumers for several more days until additional leaf area develops. Stored carbohydrate reserves continue supporting growth with starch concentrations in canes declining to 40-50% of winter maximum levels, while new photosynthate begins contributing measurably to growth carbon requirements. Protein synthesis continues at elevated rates producing structural and functional proteins for expanding tissues, with nitrogen demand high as amino acids support protein production for new cells and enzyme systems. Hormonal regulation shows sustained cytokinin and gibberellin promoting continued growth, with auxin concentration highest at shoot apex maintaining apical dominance and coordinating differentiation of leaves, tendrils, and beginning inflorescence primordia in nodes being produced by meristem. Temperature continues driving development rate with doubling per 10°C between 10-25°C, though increasing leaf area creates beginning canopy microclimate effects moderating temperature exposure of apical meristem. Water demand increases substantially with functional leaf area creating transpirational water loss of 10-30 mg per hour per shoot depending on environmental conditions, requiring adequate soil moisture and root uptake capacity to prevent water stress affecting development. Light intensity increasingly affects development as photosynthetic capacity develops, with high light promoting rapid leaf expansion and shoot growth while shade begins creating measurable growth rate reductions. The process typically requires 2-4 days from active breaking to complete breaking under favorable conditions with consistent temperatures above 16°C, with cooler conditions extending duration to 4-7 days. Cultivar variation appears in final shoot length and leaf size at complete breaking, with vigorous varieties showing 25-30 mm shoot length and 18-22 mm leaf length while less vigorous types achieve 15-20 mm shoot and 12-15 mm leaf dimensions. Regional timing follows active breaking by 2-4 days, occurring late April to early May in warm climates, early to mid-May in temperate areas, and mid-to-late May in cool continental regions. Viticultural management considerations include transitioning from frost protection focus to shoot training and canopy management planning, implementing pest management programs for insects and diseases colonizing expanding tissues, monitoring shoot growth uniformity to assess crop potential and identify management requirement variations, and planning irrigation scheduling as water demand increases with transpiring canopy development. Frost damage at complete breaking kills exposed shoot and leaves though damage severity may be slightly reduced compared to earlier stages as tissues begin hardening, still requiring compensatory growth from secondary or basal buds causing 7-14 day delays and fruitfulness reductions. Research applications include documenting transition from heterotrophic to autotrophic carbon balance in developing shoots, quantifying photosynthetic capacity development in expanding leaves, analyzing hormonal regulation of organogenesis and differentiation in active meristems, evaluating environmental effects on breaking completion timing across cultivars and regions, and assessing climate change impacts on growing season initiation and subsequent developmental timing.
Complete_Bud_Breaking is the third and final Bud_Breaking subphase, distinguished by full leaf separation from shoot axis marking bud burst completion. Transitions directly to Shoot_Growth as primary developmental process. Represents definitive end of dormancy period and beginning of vegetative growth phase. Frost vulnerability begins declining but remains elevated for next 7-10 days.
Blattentfaltung
abgeschlossener Knospenaustrieb
completed breaking
etablierter Trieb
full bud break
leaves unfolded
shoot established
vollständiger Austrieb
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 185-189 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 86-89 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Complete Bud Breaking
Chasselas demonstrates complete cane maturation disposition with 95-100% of internode length lignified by October in Swiss vineyards, including apical internodes within 5 cm of the shoot tip. In contrast, Carignan frequently shows incomplete maturation with 15-25% of distal shoot length remaining green at leaf fall in southern France. Complete maturation correlates with survival of 18-22 nodes per cane during winter, versus only 10-14 nodes in incompletely matured cultivars experiencing -15°C minimum temperatures.
A lignification disposition that is the inherited capacity of a Vitis vinifera plant to achieve thorough lignification along the entire length of current-season shoots, including distal internodes, resulting in fully mature canes with continuous periderm coverage and maximal cold hardiness potential.
Complete cane maturation is essential for spur-pruned training systems requiring viable buds at distal positions. The trait depends on timely cessation of vegetative growth, adequate carbohydrate supply, and appropriate environmental signals. Excessive nitrogen fertilization, late-season irrigation, and crop overcropping reduce maturation completeness. Assessment typically uses the percentage of brown versus green internode tissue at natural leaf fall.
complete cane ripening
durchgehende Verholzung
full shoot maturation
komplette Holzreife
thorough lignification
vollständige Triebausreifung
Galet P (2000) General Viticulture, Oenoplurimedia, Chaintre pp.: 185-192 Howell GS (2001) Sustainable grape productivity and the growth-yield relationship: a review. American Journal of Enology and Viticulture 52:165-174
Complete Cane Maturation Disposition
In a Pomerol Merlot vineyard, complete color change was documented 14 days post-color break in late September Year N+1 (BBCH 86) when skin anthocyanins plateaued at 1385 mg/kg (±60 mg/kg over subsequent 10 days), CIE L* stabilized at 30.2 (±1.8 over 2 weeks), berry Brix 24.6°, titratable acidity 5.4 g/L, firmness 1.5 N, seed tannins brown, under favorable Bordeaux autumn conditions (daily high 22-25°C, night 12-15°C, moderate deficit stem ψ -1.15 MPa maintained through veraison) with eastern canopy exposure providing morning sun.
Complete_Color_Change represents the developmental endpoint of berry pigmentation when anthocyanin accumulation reaches maximum cultivar-specific levels, spatial uniformity achieves >95% coverage, and color intensity stabilizes at final values indicating transition from active color development to stable mature state suitable for harvest or further ripening. This terminal phase occurs during late veraison to early ripening (BBCH 85-87), typically 12-18 days post-color break depending on cultivar and environmental conditions, and is characterized by skin anthocyanin concentrations plateauing at 600-3000 mg/kg fresh weight cultivar-dependent with 60 mM, and copigmentation with flavonols (120-280 mg/kg) and hydroxycinnamic acids (60-180 mg/kg) maximizing color intensity enhancement 40-60% beyond anthocyanin concentration alone. Optical characteristics at complete change inclu reflectance spectra showing stable profiles with anthocyanin absorbance maximum 520-540 nm at optical density 1.8-3.5 units cultivar-dependent, CIE L* values 22-38 with minimal week-to-week variation (32°C reducing final levels 40-70% through synthesis inhibition and degradation preventing complete change achievement, and light exposure with UV-B and moderate visible light enhancing final levels 30-55% while excessive heat-producing radiation reduces completeness. Water relations effects show: moderate sustained deficit (stem ψ -1.0 to -1.4 MPa) through veraison enhances complete change producing 25-40% higher final anthocyanins, severe deficit (<-1.5 MPa) impairs completion reducing final levels 35-65%, excess water delays and reduces complete change 25-45% through dilution and suppressed biosynthesis, and timing with pre-veraison deficit more beneficial than deficit imposed during active color development. Cultivar variation in complete change characteristics reflects: early-completing varieties (Pinot Noir, Gamay) achieve complete change 10-13 days post-color break, moderate cultivars (Merlot, Syrah, Cabernet Sauvignon) complete 12-16 days post-break, while late-completing varieties (Nebbiolo, Mourvèdre, Tannat) require 15-20 days, and final anthocyanin levels vary genetically from 600-1500 mg/kg in lower-accumulating types to 1500-3500 mg/kg in high-anthocyanin cultivars. Coordination with other ripening parameters at complete change: berry sugar typically 20-28° Brix (cultivar and style dependent) with accumulation rates declining from peak 1.5-3.5% per week to <0.5-1.0% per week, acid levels approaching harvest targets with titratable acidity 4.0-7.5 g/L and pH 3.2-3.8 as malate degradation slows substantially, berry firmness stabilized at 1.0-2.2 N representing 65-80% total softening completed, and seed maturation achieved with brown color, lignified testa, and extractable tannins indicating phenolic maturity synchronized with color completion. Molecular markers of complete change inclu anthocyanin biosynthetic gene transcripts declining to sustained moderate levels (40-70% of peak) providing maintenance expression, protein levels of biosynthetic enzymes similarly reduced through protein turnover exceeding synthesis, continued moderate ABA levels maintaining ripening program consolidation, and beginning elevation of senescence-associated genes in some cultivars indicating transition from active ripening to post-ripening phases. Temporal stability of complete color change varies: under favorable conditions (moderate temperatures 18-25°C, no environmental extremes) complete change persists 3-5 weeks enabling flexible harvest timing, heat stress can trigger anthocyanin degradation reducing levels 15-35% within 1-2 weeks through oxidative damage and enzyme activity, water stress or excess can destabilize color causing 10-25% losses, and cultivar differences with stable varieties (Cabernet Sauvignon, Syrah, Sangiovese) maintaining color 4-6 weeks while unstable types (Pinot Noir in warm climates, Grenache) may lose 20-40% through post-ripening degradation or senescence. Quality implications of complete color change inclu achievement indicates color development sufficient for harvest consideration with further hang time affecting other parameters (sugar, flavor, phenolics) more than color, final anthocyanin levels strongly predict wine color intensity and phenolic content affecting wine style and aging potential, color composition particularly acylated derivative proportions influences color stability during winemaking and aging, and timing of complete change relative to chemical and phenolic maturity determines optimal harvest window balancing color with other quality attributes. Monitoring complete change involves: visual assessment comparing berry color to cultivar-specific references indicating typical complete development, spectrophotometric measurement of skin anthocyanins confirming plateau in accumulation (<5-10% increase over 1-2 weeks), colorimetric tracking of L*a*b* values showing stabilization, and integration with sugar, acid, pH, and sensory evaluation ensuring color completion synchronized with overall maturity for optimal harvest timing. Viticultural management at complete change: irrigation often reduced or terminated recognizing reduced berry growth and water demand, canopy management adjusted to maintain cluster protection from excessive sun preventing color degradation while ensuring adequate exposure for continued ripening, disease prevention intensified as fully colored soft berries show maximum vulnerability to Botrytis and other pathogens, and harvest timing decisions integrate color completion with chemical maturity and wine style objectives balancing multiple quality parameters for optimal fruit composition.
Complete_Color_Change is distinguished from Complete_Skin_Pigmentation by representing the overall developmental endpoint rather than just pigment quantity, from Color_Development_Completion by focusing on achievement rather than process termination, and from earlier color phases by exhibiting plateau rather than continued progression. Anthocyanin plateau: <5-10% increase over 1-2 weeks post-completion. Timing: fast cultivars 10-13 days post-break, moderate 12-16 days, late 15-20 days. Final levels: 600-3000 mg/kg cultivar-dependent. Complete_Color_Change indicates color development sufficient for harvest consideration with optimal timing determined by integration with chemical and phenolic maturity.
Farb-Reifungs-Vollendung
color maturation completion
final color development achievement
finale Farb-Entwicklungs-Erreichung
maximale Pigmentierungs-Erlangung
maximum pigmentation attainment
terminal anthocyanin accumulation
terminale Anthocyan-Akkumulation
Keller M (2020) The Science of Grapevines 3rd Ed pages 235-240 color development completion and harvest maturity Coombe BG (1992) Research on development and ripening of the grape berry Australian J Grape Wine Res 48:125-144 ripening completion Kennedy JA, Matthews MA, Waterhouse AL (2002) Effect of maturity and vine water status on grape skin and wine flavonoids Amer J Enol Vitic 53:268-274 anthocyanin dynamics at maturity.
Complete Color Change
In Merlot vineyards of Saint-Émilion, Bordeaux, complete leaf detachment was achieved by December 8 following a frost event of -4°C that triggered separation of the final 8% of canopy leaves consisting primarily of apical shoot leaves and late-formed lateral leaves, resulting in fully dormant bare-cane vine appearance at BBCH stage 97 with all abscission scars showing characteristic suberized protective layer formation.
Complete leaf detachment in Vitis vinifera represents the terminal phase of the annual defoliation process, characterized by the separation and removal of all remaining leaves from the vine canopy, resulting in fully dormant bare-cane winter appearance. This phase encompasses the final 5-15% of leaf area removal, involving the last persistent leaves that either delayed senescence initiation due to favorable microclimate exposure or retained attachment despite completed abscission zone preparation. Complete leaf detachment typically occurs during December in Northern Hemisphere viticulture, finalizing the leaf fall process that began 4-8 weeks earlier with initial detachment of basal senesced leaves. The final detaching leaves are predominantly located at apical shoot positions that received optimal light exposure during the growing season and consequently maintained photosynthetic activity and chlorophyll retention longer than shaded canopy positions. Additionally, leaves on lateral shoots, water sprouts, and vigorous regrowth that initiated later in the growing season often represent the last leaves to complete detachment due to their delayed developmental timing relative to primary shoot leaves. Mechanistically, complete leaf detachment occurs through the same abscission zone separation processes as earlier phases, but may involve environmental forcing that overcomes incomplete abscission zone preparation in delayed leaves, particularly through severe frost damage that kills remaining leaf tissue and eliminates any physiological resistance to separation. The achievement of complete leaf detachment serves as an important phenological marker corresponding to BBCH stage 97, indicating that the vine has fully transitioned to winter dormancy status and has completed the annual cycle of deciduous leaf production and abscission. From a viticultural perspective, complete leaf detachment provides optimal conditions for winter pruning operations, dormant disease assessment, and vineyard structural maintenance, as the absence of foliage allows clear visualization of cane architecture and bud positioning. Delayed complete leaf detachment beyond typical timing may indicate cultivar characteristics, mild winter conditions, or nutritional factors that extended leaf functional lifespan. The completion of leaf detachment also marks the point at which vineyard floor organic matter inputs from the current season are finalized, influencing decisions regarding cover crop management and soil incorporation practices. The protective layer formed at abscission scars following complete leaf detachment consists of suberized cells that seal vascular tissues against winter desiccation and pathogen entry, completing the vine's preparation for dormant season survival.
Complete_Leaf_Detachment represents the terminal phase of Leaf_Detachment, marking achievement of 100% defoliation and full dormant canopy appearance. Distinguished from Mass_Leaf_Fall which represents rapid synchronous events affecting substantial leaf proportions, while Complete_Leaf_Detachment specifically addresses final residual leaf removal regardless of mechanism. Corresponds to BBCH stage 97 achievement. Final detaching leaves typically located at apical positions and on lateral shoots. Duration variable depending on environmental forcing, ranging from gradual final progression to event-triggered completion. Marks optimal timing for winter pruning commencement and dormant vine assessment.
abgeschlossener Blattfall
complete leaf loss
final leaf detachment
full defoliation
komplette Entblätterung
total leaf fall
totaler Laubfall
vollständiger Blattfall
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Coombe BG, Dry PR (1992) Viticulture Volume 1 Resources 2nd Ed Winetitles Adelaide Lorenz DH et al (1995) Growth stages of the grapevine: Phenological growth stages of the grapevine - BBCH-scale. Australian Journal of Grape and Wine Research 1: 100-103
Complete Leaf Detachment
Riesling in Mosel shows complete shoot lignification from late August through early October at BBCH 41-43 over 5-6 weeks, achieving lignification extending to apical regions with final 3-4 internodes showing lignin content of 14-20% dry weight and starch of 18-24% dry weight, creating fully mature brown woody shoots necessary for winter survival under cool climate conditions of 15-20°C declining to 10-15°C by maturation completion, declining precipitation, and minimal nitrogen availability typical of this moderate-vigor late-season cultivar requiring complete maturation for adequate cold hardiness in continental climate
Complete shoot lignification represents the final maturation phase where lignification extends to apical shoot regions achieving woody structure throughout entire shoot length from base to apex, establishing comprehensive dormancy preparation and maximum reserve accumulation before winter dormancy entry. This culminating developmental stage occurs during late summer-autumn corresponding to late berry development through harvest at BBCH 41-43, with apical regions completing maturation 8-14 weeks after basal initiation depending on cultivar, vigor, and environmental conditions. Complete lignification achievement requires successful progression of maturation through all shoot regions including apical internodes that formed latest and had shortest growing season, with apical maturation often most challenging particularly in vigorous cultivars or favorable growing conditions promoting extended late growth. Apical shoot lignification proceeds through same cellular processes as earlier regions with secondary wall deposition and lignin incorporation, though apical regions often achieve lower lignin content 12-20% dry weight compared to middle regions 20-30%, reflecting abbreviated maturation period before dormancy entry. Lignification extent along shoot shows characteristic gradient from thorough complete lignification in basal regions through maximum values in middle zones to progressively less complete lignification in apical regions, with most apical 2-4 internodes potentially showing incomplete lignification particularly in vigorous shoots or late-season growth. Mechanical properties in apically lignified regions show adequate rigidity for winter survival with tensile strength 30-60 MPa sufficient for supporting shoot weight and resisting snow loading, though lower than middle region values reflecting less extensive secondary wall development. Reserve accumulation in apical regions proceeds during final maturation period with starch concentrations reaching 15-25% dry weight in successfully matured apical internodes, representing important contribution to whole-shoot reserves though lower than middle region concentrations 25-35%. Complete shoot maturation assessment involves evaluating lignification extent typically requiring brown woody appearance throughout shoot with minimal green tissue remaining, adequate rigidity resisting bending with finger pressure, and periderm development complete or nearly complete over entire length. Incomplete apical lignification represents common maturation failure particularly in vigorous cultivars, excessive late-season growth, or unfavorable conditions, with unlignified or poorly lignified apical tissue showing green coloration, high flexibility, and susceptibility to winter freeze damage. Cold hardiness relationship with lignification shows critical importance of complete maturation for winter survival, with unlignified or poorly lignified apical tissues susceptible to freeze damage at temperatures as mild as -5 to -10°C while fully lignified tissues survive -15 to -25°C depending on cultivar and acclimation, making complete lignification essential in cold-climate viticulture. Reserve status at complete lignification represents maximum carbohydrate accumulation for season with whole-shoot starch content 18-28% dry weight in well-matured shoots, providing reserves essential for subsequent season bud burst, initial shoot growth before photosynthesis develops, and root growth resumption. Dormancy preparation accompanies complete lignification with metabolic downregulation toward maintenance levels, cold acclimation processes developing freeze tolerance, and water content reduction to 50-60% fresh weight approaching winter dormancy levels. Leaf senescence timing shows coordination with shoot lignification completion with leaves typically senescing and abscising during or shortly after final lignification in apical regions, representing coordinated transition to dormancy with nitrogen remobilization from senescing leaves supporting storage protein accumulation in woody tissues. Pruning wound healing capacity declines with complete lignification as cambial activity ceases and metabolic activity declines, with pruning ideally delayed until dormancy establishment after complete maturation to minimize disease infection risks. Cane quality for subsequent season pruning depends critically on complete lignification with well-matured canes showing brown color throughout length, adequate rigidity, plump appearance from good reserve status, and appropriate diameter 6-12 mm indicating balanced vigor. Maturation failure consequences include inadequate reserve accumulation reducing subsequent season bud burst vigor and early growth, increased winter freeze damage risk particularly in cold climates creating bud mortality and cane dieback, delayed spring development from limited reserve availability, and potentially reduced crop potential from winter damage. Environmental conditions during final maturation period critically affect completion with warm late summer-autumn promoting continued maturation progression, adequate soil moisture maintaining photosynthetic function supporting reserve accumulation, and moderate water stress promoting growth cessation and enhanced maturation. Temperature during complete lignification shows importance of warm conditions for final maturation progression with optimal 18-24°C during September-October in temperate regions, declining temperatures promoting growth cessation and dormancy transition, and absence of early freezes allowing maturation completion before cold damage occurs. Light intensity remains important through final maturation period with adequate late-season light supporting continued photosynthetic production for reserve accumulation, canopy management ensuring apical shoot exposure promoting thorough maturation. Water availability shows moderate stress beneficial for promoting growth cessation and final maturation without excessive stress limiting photosynthetic reserve production, with irrigation management in late season targeting moderate stress promotion. Nutrient availability particularly nitrogen shows declining levels appropriate for maturation with nitrogen management avoiding late-season application that would prolong growth and delay or prevent maturation completion. The process timing varies substantially with cultivar maturity characteristics, vigor levels, and environmental conditions, with early-maturing cultivars completing lignification by late August-early September while late-maturing types requiring September-October for completion. Cultivar variation appears in maturation timing with early-maturing varieties showing rapid progression to complete lignification while late-maturing types show extended maturation period, apical lignification completeness with some varieties consistently achieving thorough apical maturation while others frequently show incomplete apical lignification affecting winter hardiness, and reserve accumulation capacity affecting dormancy reserve status and subsequent season vigor potential. Regional timing shows complete shoot lignification occurring August-September in warm climates, September-October in temperate regions, and September-early October in cool areas with critical importance of completing maturation before significant freezes in cold regions. Viticultural management considerations include monitoring maturation progress through late season assessing completion likelihood before dormancy, implementing management promoting timely growth cessation and complete maturation including water stress management and nitrogen control, assessing maturation completion status at dormancy entry predicting winter survival and subsequent season development potential, selecting cultivars with maturation characteristics appropriate for climate ensuring reliable complete maturation, and understanding that incomplete maturation represents major limitation in cold-climate viticulture requiring management attention. Research applications include analyzing factors determining final maturation completeness including genetic and environmental controls, quantifying relationships between maturation extent and winter hardiness across cultivars and climates, evaluating management strategies ensuring reliable complete maturation, modeling maturation progression for predicting completion timing and dormancy preparation, and assessing climate change impacts on maturation patterns affecting winter survival and productivity across viticultural regions.
Complete_Shoot_Lignification is fourth and final Shoot_Lignification subphase, distinguished by maturation extending to apical regions achieving woody structure throughout shoot. Represents culmination of entire maturation process. Completeness critically determines winter survival, reserve status, and subsequent season vigor potential. Timing relative to dormancy entry and first freezes critical in cold climates. Assessment of completion status essential management activity at season end.
Ausreifungsabschluss
complete woodification
komplette Triebausreifung
shoot maturation completion
total shoot lignification
totale Holzbildung
vollständige Triebverholzung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 300-315 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 230-245 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Complete Shoot Lignification
In a Barolo Nebbiolo vineyard in Piedmont, complete skin pigmentation was achieved in early October Year N+1 (BBCH 87, 14 days post-color break) when skin anthocyanins reached 1180 mg/kg (malvidin-3-glucoside 625 mg/kg, delphinidin-3-glucoside 155 mg/kg, cyanidin-3-glucoside 195 mg/kg, acylated forms 28%), CIE L* 34.2, berry sugar 23.8° Brix, titratable acidity 6.8 g/L, seed tannins brown indicating phenolic maturity, under typical Langhe autumn conditions (daily high 18-22°C, night 8-12°C, moderate deficit stem ψ -1.2 MPa) with southeastern hillside exposure.
Complete_Skin_Pigmentation represents the final state of anthocyanin accumulation in berry epidermis and hypodermis when pigment concentrations reach cultivar-specific maxima throughout all skin cell layers creating the fully developed color potential that determines wine color intensity and phenolic composition. This state is achieved during late veraison to early ripening (BBCH 85-87), typically 10-16 days post-color break depending on cultivar and environmental conditions, and is characterized by skin anthocyanin concentrations reaching 500-3000 mg/kg fresh weight cultivar-dependent (Pinot Noir 600-1500 mg/kg, Merlot 800-1800 mg/kg, Cabernet Sauvignon 900-2000 mg/kg, Syrah 1200-2400 mg/kg, Petit Verdot 1500-3000 mg/kg) representing 95-100% of maximum achievable levels under given environmental and management conditions. Cellular and tissue distribution of complete pigmentation shows: epidermal cell layer (outermost 1-2 cell layers) contains highest anthocyanin concentrations 18-35 mM vacuolar concentration representing localized levels 2500-4500 mg/kg tissue equivalent, sub-epidermal layers (2-4 cells deep) show 60-85% of epidermal concentrations contributing 400-800 mg/kg to total skin anthocyanins, deeper hypodermal cells (if pigmented, cultivar-dependent) contain 30-60% of epidermal levels adding 200-500 mg/kg in extensively pigmented varieties, and spatial integration across 3-6 pigmented cell layers creates total skin anthocyanin pool determining extractability during winemaking and wine color potential. Anthocyanin composition at complete pigmentation establishes final profile: malvidin-3-glucoside dominates at 50-72% of total (absolute 300-2000 mg/kg cultivar-dependent), delphinidin-3-glucoside contributes 4-20% (20-400 mg/kg, higher in Syrah 12-18%, Petit Verdot 14-22%), cyanidin-3-glucoside ranges 6-22% (30-350 mg/kg, elevated in Pinot Noir 15-25%), peonidin and petunidin derivatives account for 5-15% each (25-300 mg/kg combined), and acylated forms (acetylated and coumaroylated anthocyanins) reach 18-32% of total (100-800 mg/kg) providing enhanced color stability through intramolecular copigmentation and resistance to pH-induced color changes. Biosynthetic status at complete pigmentation shows: anthocyanin biosynthetic gene expression declining from peak levels as accumulation approaches maximum with UFGT transcripts reducing to 60-80% of peak (still 80-130 fold above green berry baseline), enzyme activities similarly declining with UFGT at 60-75% maximum (20-35 units/kg) maintaining moderate synthesis balancing degradation, and regulatory factors (VvMYBA1/A2) showing 50-70% of peak expression indicating transition from active accumulation to maintenance phase. Vacuolar sequestration at complete pigmentation involves: maximum anthocyanin concentrations in cell vacuoles 15-35 mM approaching or exceeding solubility limits in some cells, vacuolar pH maintained at 2.8-3.5 through continued V-ATPase activity ensuring flavylium cation stability and color expression, formation of anthocyanin vacuolar inclusions (AVIs) in cultivars exhibiting this trait (Cabernet Sauvignon, Merlot) creating highly concentrated pigment domains >50 mM local concentration, and copigmentation with flavonols (100-250 mg/kg skin) and hydroxycinnamic acids (50-150 mg/kg) enhancing color intensity 35-55% beyond that expected from anthocyanin concentration alone through molecular stacking and bathochromic shifts. Environmental factors determining final pigmentation levels inclu temperature history during color development with optimal 22-28°C supporting maximum accumulation (1200-2500 mg/kg in high-anthocyanin cultivars), cool conditions 18-23°C potentially enhancing final levels 15-30% through improved stability despite slower synthesis, heat exposure >32°C reducing final pigmentation 40-70% through synthesis inhibition and degradation, and cumulative heat stress effects where multiple heat episodes create progressive reduction in achievable maximum. Light exposure effects show: UV-B radiation during veraison enhances final pigmentation 35-60% through sustained gene induction, optimal visible light exposure produces 25-45% higher final anthocyanins compared to shaded fruit, excessive direct sun combined with heat can reduce final levels 20-40% through heat stress overriding light benefits, and cluster position creating systematic variation with southwestern exposure (northern hemisphere) showing 30-50% higher final pigmentation than northeastern positions. Water deficit impacts on complete pigmentation demonstrate: moderate sustained deficit (stem ψ -1.0 to -1.4 MPa throughout veraison) enhances final pigmentation 25-45% through elevated ABA signaling and berry concentration, severe deficit (<-1.5 MPa) reduces final levels 30-60% through metabolic limitation despite high ABA, timing matters with pre-veraison deficit producing better results than deficit imposed during active accumulation, and excess water reduces final pigmentation 30-50% through berry dilution and suppressed ABA-responsive gene expression. Nutritional influences inclu moderate nitrogen limitation during ripening can enhance anthocyanin accumulation 15-30% through reduced vegetative competition and potential stress signaling, excessive nitrogen reduces pigmentation through vigorous vegetative growth creating shading and sink competition, potassium sufficiency appears important with deficiency potentially reducing final anthocyanins 20-35% through pH regulation effects or membrane transport impairment, and general nutrient balance supports optimal accumulation while severe deficiencies compromise biosynthetic capacity. Cultivar genetic variation determines: high-anthocyanin varieties (Petit Verdot, Tannat, Alicante Bouschet) achieve 1800-3500 mg/kg through high biosynthetic gene expression and extended accumulation duration, moderate cultivars (Cabernet Sauvignon, Syrah, Merlot) reach 900-1800 mg/kg with balanced synthesis rates and duration, while lower-anthocyanin types (Pinot Noir, Grenache, Sangiovese) attain 600-1400 mg/kg through lower synthesis capacity or shorter accumulation period reflecting genetic differences in VvMYBA regulation and pathway enzyme expression. Complete pigmentation timing relative to other maturity parameters: typically achieved when berry sugar reaches 20-26° Brix (cultivar and style dependent), coincides with 75-90% of total berry softening creating firmness 1.0-2.0 N, occurs during late acid decline with titratable acidity 4.5-7.5 g/L approaching harvest targets, and parallels seed maturation (brown color, lignified testa) and flavor/aroma development indicating integrated ripening completion. Wine quality implications inclu final skin anthocyanin levels strongly predict wine color intensity with correlation r² = 0.88-0.96, anthocyanin composition particularly acylated derivative proportions influences color stability and aging potential, extraction efficiency during maceration depends on total anthocyanin pool with higher skin levels enabling shorter extraction times or lower temperatures, and spatial uniformity of complete pigmentation affects extraction consistency with uniform berries producing more predictable wine color than heterogeneous fruit. Temporal stability of complete pigmentation varies: under favorable conditions (moderate temperatures 18-25°C, no water stress) complete pigmentation persists 3-5 weeks enabling flexible harvest timing, environmental stresses can trigger degradation within 1-2 weeks through oxidative damage, enzyme activation, or cellular disruption, and cultivar differences exist with stable varieties (Cabernet Sauvignon, Syrah) maintaining levels while unstable types (Pinot Noir in warm climates) may lose 20-40% anthocyanins through post-ripening degradation. Monitoring complete pigmentation involves: spectrophotometric analysis of skin extracts measuring total anthocyanins and composition through HPLC, portable spectrophotometers providing field measurements of skin color correlating with anthocyanin levels, visual assessment comparing berry color to cultivar-specific standards indicating typical complete pigmentation, and integration with sugar, acid, and sensory evaluation ensuring anthocyanin accumulation synchronized with overall maturity for optimal harvest timing.
Complete_Skin_Pigmentation is distinguished from Maximum_Color_Intensity by representing total anthocyanin accumulation rather than visual intensity per se, from Even_Berry_Coloration by focusing on pigment quantity rather than spatial distribution, and from Color_Development_Completion by emphasizing final state rather than process termination. Final levels: high cultivars 1800-3500 mg/kg, moderate 900-1800 mg/kg, lower 600-1400 mg/kg. Timing: typically 10-16 days post-color break. Temperature optimum 22-28°C. Moderate deficit enhances 25-45%. Complete_Skin_Pigmentation determines wine color potential and phenolic composition critical for quality and style.
complete anthocyanin deposition
final epidermal pigmentation
finale epidermale Pigmentierung
full pigment development
maximale Haut-Anthocyan-Akkumulation
maximum skin anthocyanin accumulation
volle Pigment-Entwicklung
vollständige Anthocyan-Ablagerung
Keller M (2020) The Science of Grapevines 3rd Ed pages 235-238 anthocyanin accumulation and wine quality Castellarin SD, Matthews MA, Di Gaspero G, Gambetta GA (2007) Water deficits accelerate ripening and induce changes in gene expression regulating flavonoid biosynthesis in grape berries Planta 227:101-112 environmental regulation of final levels Mattivi F, Guzzon R, Vrhovsek U, Stefanini M, Velasco R (2006) Metabolite profiling of grape:grape and wine standardization Anal Chim Acta 563:184-191 anthocyanin composition at maturity.
Complete Skin Pigmentation
Syrah in Rhône Valley shows completed pollen release during late June Year N+1 at BBCH 67-69 on approximately June 15-16, with all viable anthers having completed discharge, atmospheric pollen concentration declining to <25 grains per cubic centimeter, widespread empty withered anthers throughout cluster, visible berry development 14-22 mm diameter representing 52-64% established fruit set, under warm Mediterranean conditions of 25-29°C temperature, typical of this mid-to-late season cultivar completing male reproductive phase 11-13 days after first dehiscence and transitioning definitively to berry development
Completed pollen release represents the concluding male reproductive phase when all viable anthers have dehisced and exhausted pollen supplies, atmospheric pollen concentration declines to negligible levels, and male gamete availability effectively ceases, marking definitive end of male reproductive contribution and transition to exclusive reliance on established fertilizations for fruit set. This conclusive male phase occurs during late-complete flowering in late June-early July in temperate regions Year N+1 following peak release, typically spanning 2-3 days as final pollen discharge completes and atmospheric pool dissipates, corresponding to BBCH 67-69 late-complete anthesis stages. Completion characteristics show definitive cessation of male reproductive function with all developmentally capable anthers having completed dehiscence and pollen discharge, exhausted pollen supplies with anthers depleted of releasable grains, negligible atmospheric pollen concentration with remaining airborne pollen aged and non-viable, and overall termination of effective male gamete availability. Anther status at completion shows widespread empty dehisced anthers with longitudinal slits exposing depleted locules, withered anther structures showing brown discoloration indicating senescence, occasional late-dehiscing anthers in basal flowers completing final releases contributing minimal pollen, and overall cluster showing predominantly post-reproductive anther state. Pollen exhaustion progression shows characteristic depletion pattern with early-dehiscing anthers in apical regions completely exhausted having released all viable pollen over previous days, mid-cluster anthers mostly depleted with minimal residual non-releasable pollen remaining, late-dehiscing basal anthers completing discharge contributing final minor pollen additions, and overall progression toward complete cluster-level exhaustion. Atmospheric pollen decline shows rapid concentration reduction following peak maximum with levels dropping from 500-1500 grains per cubic centimeter at peak to 50-200 during late release to <20 at completion, continued atmospheric removal through settling and dispersal eliminating remaining airborne pollen, and cessation of fresh production preventing atmospheric pool replenishment. Residual atmospheric pollen characteristics include predominantly aged grains with extended atmospheric residence, severely reduced viability typically <10-20% germination capacity due to progressive aging and environmental exposure, insufficient concentration for effective pollination with encounter probabilities declining substantially, and overall non-functional male gamete pool unable to support significant additional fertilizations. Late pollen release from final anthers shows limited contribution with basal and delayed anthers completing dehiscence contributing minor pollen additions, fresh pollen from late release showing adequate viability 65-80% but insufficient quantity for meaningful pollination, and overall late release contributing <5-10% fruit set representing minor final additions. Male reproductive cessation at completion shows definitive termination with no further viable pollen production capacity, exhausted anther reserves preventing continued release, and physiological commitment to post-reproductive senescence eliminating male function. Berry development visibility at completion shows established fruit structures throughout cluster with early-set berries 12-20 mm diameter from peak flowering success dominating appearance, late-set berries 8-14 mm from late pollinations visible in some positions, and overall berry-dominated cluster confirming reproductive outcomes and male contribution completion. Unfertilized flower status at completion shows senescing unsuccessful reproductions with brown discoloration and tissue deterioration, active abscission eliminating failed flowers progressively over subsequent weeks, and visible flower drop creating accumulated debris documenting unsuccessful pollination attempts. Fruit set finalization at completion establishes definitive berry number with count of successfully fertilized flowers determining season yield, typical set percentages 30-75% depending on cultivar and conditions creating 100-500 berries per cluster, and recognition that male gamete contribution completed with established berry number representing maximum potential yield. Pollination opportunity cessation at completion shows terminated effective pollination capacity with negligible viable pollen availability, any remaining late receptive stigmas unable to achieve successful pollination, and final unsuccessful flowers destined for abscission due to failed reproduction. Developmental transition at completion marks shift from reproductive to exclusively developmental focus with all vine resources directing to established berry growth, no residual male reproductive investment or function, and physiological reorientation toward fruit development and maturation. Environmental effects at completion show reduced significance compared to earlier reproductive phases with outcomes largely determined by previous peak period conditions, though conditions during completion affecting final late flower success contributing minor proportion, and overall weather during completion having minimal impact on established fruit set. Anther senescence at completion shows progressive tissue deterioration with withered structures showing brown discoloration, cellular breakdown as anthers degenerate following exhaustion, and eventual abscission of senescent anthers from flowers over subsequent weeks though many persist. Pollen deposition evidence at completion includes accumulated pollen on horizontal surfaces within canopy showing yellow dust layers, ground deposition creating visible layer documenting flowering intensity, and persistent visual evidence of male reproductive activity despite completion. Observational characteristics enabling completion identification include absence of visible pollen clouds or fresh pollen release, widespread empty withered anthers throughout cluster, predominantly berry appearance with minimal residual flowering activity, and overall visual evidence of reproductive conclusion with developmental transition. Phenological significance at completion includes marking definitive end of male reproductive phase, establishing reference for developmental stage transitions with progression from reproductive BBCH 60-69 to developmental BBCH 71-79 stages, and confirming flowering completion enabling post-bloom management transitions. Management implications at completion include confirming male reproductive function completed enabling assessment of pollination outcomes, resuming full normal operations without reproductive period restrictions, implementing post-bloom berry development management, and transitioning focus entirely to fruit development and quality optimization. Fruit set assessment at completion enables accurate yield prediction through berry counting, evaluation of male reproductive success through comparison to flower number, and early season productivity estimates informing management decisions. The process shows two distinguishable aspects progressing from pollen exhaustion marking depletion of anther reserves, through empty anther formation establishing post-reproductive state. Duration typically spans 2-3 days representing completion period, with rapid transition to exclusively developmental phase. Cultivar variation in completion timing shows early varieties completing late June while late types complete early-mid July reflecting overall phenology, completion efficiency with some showing clean complete discharge while others showing residual retention, and transition clarity with some showing definitive completion while others showing gradual extended conclusion. Regional timing shows completed pollen release during late June in warm climates at approximately 1150-1200 growing degree days base 10°C, late June-early July in temperate regions at 1050-1100 GDD, and early-mid July in cool areas at 950-1000 GDD, marking male reproductive phase conclusion. Viticultural significance includes marking definitive end of male reproductive contribution confirming fruit set outcomes, indicating completion of critical weather-vulnerable flowering period reducing risk, enabling complete management transitions to developmental priorities, and establishing reference for subsequent phenological and developmental timing. Research applications include documenting precise male reproductive phase completion timing, analyzing factors affecting complete pollen discharge achievement, evaluating relationships between pollen release patterns and final fruit set outcomes, assessing post-reproductive anther senescence processes, and documenting phenological transitions marking major phase changes in reproductive biology.
Completed_Pollen_Release represents concluding male reproductive phase parallel to Complete_Anthesis within Flowering. Corresponds to BBCH 67-69 late-complete anthesis stages. Subdivided into Pollen_Exhaustion and Empty_Anther_Formation. Marks definitive end of male gamete availability and contribution. Establishes final fruit set from male perspective. Enables management transitions and developmental phase focus. Completes male reproductive cycle from dehiscence through peak release to exhaustion.
Antheren-Entleerung
Pollenproduktionsende
abgeschlossene Pollenfreisetzung
anther depletion
end of pollen production
male gamete exhaustion
männliche Gametenerschöpfung
pollen release completion
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 388-395 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 295-300
Completed Pollen Release
A complex compound inflorescence at node 3 on a vigorous shoot of Vitis vinifera cv. Muscat of Alexandria in a highly fertile vineyard in Crete, Greece, measuring 28.5 cm length and approximately 10.5 cm maximum width creating massive three-dimensional structure, exhibiting exceptional architectural complexity with branching extending through 6-7 hierarchical orders, central rachis diameter of 5.5 mm at base supporting the elaborate structure, bearing 28 major lateral branches densely distributed along the rachis, each major lateral showing extensive individual development with complex secondary, tertiary, and quaternary branching creating intricate architecture, some major laterals reaching 9-10 cm length themselves resembling complete moderate inflorescences, total flower number estimated at approximately 1150 flowers distributed throughout the elaborate branching network, architectural complexity including slight rachis forking in the apical region adding to structural intricacy, overall structure showing conical to cylindrical profile with very dense interior regions experiencing significant shading, measured at BBCH stage 59 (inflorescence fully developed, flowers separating) in late May, representing maximum complex compound inflorescence elaboration requiring aggressive cluster thinning to reduce crop load and prevent vine exhaustion and fruit quality degradation.
A compound primary inflorescence exhibiting maximum architectural elaboration combining multiple complexity features in Vitis vinifera, characterized by very high branching order typically 5-7 hierarchical levels creating exceptional depth, extensive lateral branch development with numerous major laterals showing complex individual architectures, very large overall dimensions often 24-32 cm length and 8-12 cm width creating substantial three-dimensional structures, very high flower number frequently exceeding 800-1200+ flowers, intricate branching patterns showing elaborate secondary, tertiary, quaternary, and higher-order branch systems, architectural irregularities potentially including forked rachis, co-dominant axes, or other developmental variations adding to complexity, and overall structural elaboration approaching the maximum expression observed in grapevine inflorescence architecture. Complex compound inflorescences represent the extreme architectural elaboration combining all complexity features including high branch order, extensive lateralization, large size, and high flower number, occurring in exceptionally fertile cultivars under optimal development conditions, exhibiting the maximum developmental investment creating the largest most elaborate flowering structures, and transforming into very large dense clusters requiring intensive management to maintain fruit quality and prevent overcropping damage.
Complex compound inflorescences represent the maximum architectural elaboration extreme, distinguished from multi-branched and other compound types by combination of all complexity features rather than emphasis on single attributes. The complex designation emphasizes the comprehensive elaboration including high branch order, extensive lateralization, large size, high flower number, and potential architectural irregularities. Branching order reaching 6-7 levels represents exceptional hierarchical depth approaching the maximum observed in grapevine. Size exceeding 25-30 cm length and 10+ cm width creates very large structures substantially exceeding typical inflorescence dimensions. Flower number >1000 represents exceptional fertility with some extreme examples potentially exceeding 1500+ flowers. Architectural irregularities including forked rachis or co-dominant axes occasionally occur in very complex inflorescences representing developmental variations adding to structural intricacy. Occurrence requires exceptional fertility combined with optimal development conditions during both primordium initiation and current-season growth, with most cultivars never producing this extreme elaboration even under favorable conditions. Management necessity involves aggressive intervention with cluster thinning often removing 30-50% of clusters and potentially additional berry thinning to achieve acceptable crop loads and fruit quality. Harvest challenges include very large heavy clusters requiring careful handling and potential non-uniform ripening from interior shading. Complex compound architecture represents fertility extreme requiring balancing through crop load management to maintain vine health and sustainable production.
extremely complex panicle
komplex zusammengesetzte Infloreszenz
maximal elaborierte Rispe
maximally elaborate inflorescence
ultimate compound structure
ultimative zusammengesetzte Struktur
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 419-458 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 466-512 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 659-708.
Complex Compound Inflorescence
Fercal (BC1 of V. berlandieri × V. vinifera 'Colombard' 31R crossed with Berlandieri Colombard 1B) represents a complex hybrid achieving extreme lime tolerance (60% active lime) through concentration of V. berlandieri and V. vinifera lime-tolerance genes, though with reduced phylloxera resistance requiring use in low-phylloxera regions. Freedom (1613 Couderc × Dog Ridge, where 1613C itself is V. solonis × Othello involving multiple species) combines V. champinii nematode resistance with improved graft compatibility and moderate cold tolerance from the complex 1613C parent. GRN-1 to GRN-5 (UC Davis breeding) represent complex hybrids pyramiding nematode resistance from V. champinii and V. rufotomentosa with phylloxera resistance and improved enological performance.
A parentage-based rootstock role that inheres in a grafted rootstock by virtue of its genetic derivation from hybridization involving three or more Vitis species, or from backcross or multi-generational breeding schemes that combine genetic contributions from diverse parental backgrounds beyond simple F1 interspecific crosses. Complex hybrid rootstocks are developed to stack multiple desirable traits that cannot be achieved through single crosses between two species, such as combining nematode resistance (from V. champinii) with lime tolerance (from V. berlandieri) and improved graft compatibility (from V. vinifera or early-generation hybrids). Complex hybrids include backcross derivatives designed to recover graft compatibility while retaining wild-species resistance traits, three-way and four-way crosses combining complementary resistances, and selections from segregating populations of complex pedigree. Examples include Fercal (BC1 involving V. berlandieri, V. vinifera, and V. riparia genetic background), Freedom and Harmony (involving V. champinii, V. riparia, V. rupestris, and V. vinifera), and Muscadinia rotundifolia-derived rootstocks under development for Xiphinema resistance.
Complex_Hybrid_Rootstock_Role captures rootstocks whose breeding history involves more than simple F1 crosses between two species, reflecting advanced breeding strategies to combine traits not co-occurring in any single species pair. Modern rootstock breeding increasingly focuses on complex hybrids to address multiple simultaneous challenges (e.g., phylloxera + nematodes + fanleaf virus vector + lime tolerance). Genomic selection and marker-assisted breeding are accelerating complex hybrid development by enabling trait stacking with reduced linkage drag. The Muscadinia rotundifolia introgressions for Xiphinema/GFLV resistance represent a particularly challenging complex hybrid development due to incompatibility barriers between Muscadinia and Vitis.
Mehrarten-Hybridunterlage
Rückkreuzungs-Unterlage
advanced generation rootstock
backcross rootstock
complex hybrid rootstock
komplexe Hybridunterlage
multi-species hybrid rootstock
Walker MA et al. (2014) Rootstock breeding for resistance to nematodes and phylloxera, Acta Horticulturae 1045: 89-97 Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22 Cousins P (2005) Evolution, genetics, and breeding: viticultural applications of the origins of our rootstocks, Grapevine Rootstocks: Current Use, Research, and Application, ASEV pp.: 1-7 Riaz S et al. (2019) Genetic analysis of nematode resistance in grape rootstocks, Theoretical and Applied Genetics 132: 1941-1955
Complex Hybrid Rootstock Role
Regent (Diana × Chambourcin) bears a Complex_Interspecific_Hybrid_Role with documented ancestry from V. vinifera, V. labrusca, V. rupestris, V. lincecumii, and V. riparia accumulated through French-American hybrid intermediates across multiple generations—its complex pedigree contributing multiple resistance mechanisms alongside wine quality approaching V. vinifera standards. Cabernet Blanc (complex hybrid from JKI Geilweilerhof) serves in a Complex_Interspecific_Hybrid_Role with contributions from Cabernet Sauvignon (V. vinifera quality) combined with resistance from complex hybrid resistance donors. VB91-26-4 (an advanced French breeding line) bears a Complex_Interspecific_Hybrid_Role incorporating M. rotundifolia-derived Run1 powdery mildew resistance alongside V. vinifera background through multiple backcross generations with interspecific complexity.
An interspecific hybrid role that inheres in grapevine genotypes with ancestry from multiple (typically three or more) Vitis species and/or Muscadinia rotundifolia, accumulated through multiple generations of interspecific crossing and selection. Complex interspecific hybrids carry genetic contributions from diverse wild species—typically resistance genes from American species (V. riparia, V. rupestris, V. lincecumii, V. labrusca, V. cinerea) combined with quality traits from V. vinifera and potentially additional species for specific characteristics (V. amurensis for cold hardiness, M. rotundifolia for powdery mildew immunity). The "complex" designation indicates pedigree depth and species diversity beyond simple two-species crosses, with multi-generational breeding progressively combining favorable alleles while selecting against deleterious contributions. Most contemporary PIWI (pilzwiderstandsfähig) varieties are complex interspecific hybrids, their pedigrees tracing through French-American hybrid intermediates and purpose-bred resistance donors to combine multiple R-genes ("pyramiding") with improved wine quality. Complex hybrids present both opportunities (combining traits from multiple sources) and challenges (genetic complexity complicating breeding predictions and potentially introducing unexpected trait interactions).
Complex_Interspecific_Hybrid_Role captures the multi-species pedigrees characteristic of advanced resistance breeding, where generations of crossing accumulate diverse genetic contributions. The complexity arises from breeding strategy—combining resistance from multiple sources (R-gene pyramiding) requires crossing resistance donors, each of which may carry different wild species ancestry, then combining these with V. vinifera quality parents. Pedigree complexity increases breeding program management challenges (tracking contributions from many sources) but enables trait combinations unachievable through simple crosses. Contemporary molecular tools (genome-wide markers, known R-gene sequences) help manage complexity by enabling precise selection within genetically diverse populations. The "complex hybrid" terminology acknowledges that species contributions may be uncertain for distant ancestors, with pedigree records incomplete for historic French-American hybrids that contribute to contemporary breeding.
Mehrarten-Hybride
advanced interspecific hybrid
complex hybrid
complex pedigree hybrid
komplexe Hybride
komplexe Interspezies-Hybride
multi-species hybrid
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330 Töpfer R et al. (2011) Molecular breeding in grapes, CRC Press pp.: 160-185 Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance, Molecular Genetics and Genomics 291: 1573-1594
Complex Interspecific Hybrid Role
A complex multi-branched permanent arm on heritage Vitis vinifera cv. Palomino vine in traditional Jerez region, featuring trunk dividing into three primary branches at 60cm height, each primary then bifurcating creating six secondary branches, several secondaries further subdividing creating total of nine terminal fruiting positions distributed across complex three-dimensional architecture extending 0.8-1.2m in various directions, total 67 spurs distributed across multiple branch levels, representing traditional complex branching approach developed over decades of incremental training without systematic design, now maintained as heritage architecture despite management challenges including difficult pruning access, uneven vigor distribution among branches, structural weakness at multiple junction points, and complete incompatibility with mechanical operations requiring hand management for all operations.
A multi-branched permanent arm featuring five or more distinct branches or multiple levels of branching creating highly complex architecture, characterized by extensive structural division beyond standard four-branch configurations, numerous branch points distributing fruiting capacity across many separate zones, and extreme architectural complexity providing maximum canopy distribution, specialized training system requirements, or adaptation to unusual site constraints, though at substantial cost in establishment difficulty, management complexity, and often reduced mechanical compatibility. Complex multi-branched permanent arms represent extreme expressions of branched architecture, rarely employed in commercial viticulture except in very specific contexts including experimental divided-canopy systems, adaptations to extreme terrain or spacing irregularities, traditional systems in limited regions employing complex branching philosophies, or artistic/heritage vineyards where architectural complexity serves aesthetic or historical preservation objectives despite practical challenges.
Complex multi-branched permanent arms are rare in modern commercial viticulture, with extreme structural complexity creating overwhelming practical challenges including very difficult establishment (achieving desired multi-level branching patterns requiring years of skilled training), complex ongoing management (pruning and canopy work at numerous branch levels and junction points), structural vulnerability (multiple branch points concentrating stress), severely unbalanced vigor (apical and better-positioned branches dominating weaker positions), and mechanical incompatibility (complex geometries impossible for standard equipment). The few contexts where complex multi-branching appears include heritage vineyards preserving historical training approaches, experimental systems exploring extreme canopy division concepts, adaptations to severe terrain or spacing irregularities where conventional architectures are impossible, and artistic or demonstration vineyards where architectural complexity serves educational or aesthetic purposes despite practical costs. The general trend in modern viticulture strongly favors simpler architectures (bilateral, three-branch, four-branch maximum) over complex multi-branched configurations.
elaborate branch architecture
extreme multi-fork arm
highly-branched arm
hochverzweigte Arme
komplex verzweigte Arme
Complex Multi Branched Permanent Arm
The compost application rate for a biodynamically managed Pinot Noir vineyard in the Willamette Valley specifies 4-6 t/ha of mature cow manure compost (C/N ratio 15-18, temperature-stabilized through 6 months of windrow composting with biodynamic preparations 502-507 inserted into the pile) applied annually in October after harvest, surface-broadcast across the full vineyard floor by side-throwing manure spreader, with the compost providing an estimated 20-30 kg N/ha (of which 10-15% is plant-available in the first year through mineralization), 10-15 kg P/ha, 25-35 kg K/ha, and approximately 1.5-2.0 t/ha of stable organic carbon — the application rate is calibrated to maintain soil organic matter above 3.5% (currently 4.1% after 15 years of annual composting, compared to 2.8% in adjacent non-composted blocks) and to sustain earthworm populations exceeding 300 individuals per m².
A soil management description that documents the quantity, composition, timing, and method of compost application to vineyard soils, specifying the compost source material, maturity (C/N ratio and stability indicators), application rate in tonnes per hectare, distribution method (surface broadcast, banded, or incorporated), application frequency, and the target soil organic matter level or nutrient supply objective that the composting program is designed to achieve.
Compost_Application_Rate documents the primary mechanism for organic matter replenishment and biological activation in vineyard soils. The application rate must balance multiple objectives — providing sufficient organic matter to maintain or increase soil carbon stocks, supplying plant-available nutrients (particularly nitrogen, phosphorus, and potassium) to meet vine nutritional requirements without excess, supporting soil biological communities (bacteria, fungi, earthworms, arthropods) that drive nutrient cycling and soil structural development, and avoiding excessive nutrient loading that could stimulate unwanted vegetative vigor or contribute to groundwater contamination: typical application rates of 3-8 t/ha of mature compost annually provide 1-3 t/ha of organic carbon input, comparable to the annual carbon loss from a cultivated vineyard soil of 1-2 t/ha through microbial respiration, maintaining a positive or neutral soil carbon balance. Compost maturity is critical — immature compost with C/N ratios above 25 causes temporary nitrogen immobilization as soil microorganisms consume available nitrogen to decompose the carbon-rich material, while mature compost with C/N ratios of 12-20 provides net nitrogen mineralization and stable humus formation.
Kompost-Aufwandmenge
Kompostausbringungsrate
Kompostierungs-Protokoll
compost rate specification
organic matter application rate
vineyard composting protocol
White RE (2015) Understanding Vineyard Soils 2nd Ed, Oxford University Press pp: 165-185 Morlat R and Chaussod R (2008) Long-term additions of organic amendments in a Loire Valley vineyard I: Effects on properties of a calcareous sandy soil, American Journal of Enology and Viticulture 59: 353-363 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 445-460
Compost Application Rate
A compound dormant bud at node 6 on a mature cane of Vitis vinifera cv. Merlot in January, with longitudinal dissection revealing a well-developed primary bud containing two inflorescence primordia with 180 and 165 flower initials respectively, a secondary bud with one small inflorescence primordium of 45 flowers, and a tertiary bud containing only vegetative primordia, all structures dormant and protected by seven overlapping brown bud scales.
A dormant bud exhibiting the characteristic compound architecture unique to Vitis species, containing multiple independent meristematic units (typically primary, secondary, and tertiary buds) enclosed within a single set of protective bud scales, each capable of independent development and serving as backup growing points in case of damage to more dominant primordia. Compound dormant buds represent a survival strategy providing developmental redundancy, with the primary bud typically containing both vegetative and reproductive primordia, secondary buds often predominantly vegetative with reduced or absent inflorescence initials, and tertiary buds usually solely vegetative, all maintained in dormant state until environmental cues trigger selective or sequential activation.
The compound bud structure is diagnostic for the genus Vitis and represents a key adaptation for survival in variable environments. Not all buds in compound structures are equally dormant or equally responsive to environmental cues primary buds typically break first, with secondary and tertiary activation occurring only under specific conditions (primary damage, strong forcing, hormonal shifts). Understanding compound bud architecture is critical for pruning decisions and yield prediction in viticulture.
Komplexknospe
compound bud
multi-primordium bud
zusammengesetzte Knospe
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Morrison (1991) "Bud development in Vitis= Anatomical study of formation and growth of the primary grapevine bud during one annual cycle" Australian Journal of Grape and Wine Research 72=296-301 Srinivasan & Mullins (1981) "Physiology of flowering in the grapevine - a review" American Journal of Enology and Viticulture 32=47-63.
Compound Dormant Bud
Cabernet Sauvignon pistils at BBCH 57-59 (flower buds swollen) exhibit characteristic compound pistil structure with two-carpel fusion evident in bilocular ovary organization visible in transverse sections showing two distinct chambers separated by central septum (formed from fused carpel margins), unified style connecting ovary to stigma, and stigma showing subtle bilobed organization reflecting bicarpellate origin though structurally fused into single receptive surface. Pinot Noir compound pistils show typical Vitis bicarpellate organization with 2-4 ovules distributed as 1-2 per locule attached to axile placentation on central septum, unified style transmitting tract supporting pollen tube growth from stigma to both locules, and stigma papillae showing bilateral distribution pattern corresponding to two-carpel organization. Syrah pistil anatomical sections reveal compound nature through bilocular ovary structure, though some flowers show asymmetric locule development with one locule slightly larger than the other while maintaining overall bicarpellate compound organization. Rare Merlot flowers (<0.05% frequency) exhibit abnormal compound pistils with three locules suggesting involvement of three carpels, producing enlarged ovaries and occasionally tricarpellate berries with unusual morphology and seed patterns.
A compound pistil is a pistil formed from two or more fused carpels creating a syncarpous gynoecium with carpel fusion evident in shared ovary chamber(s), unified style, and common stigma, representing the ancestral and universal pistil organization in Vitis where two carpels invariably fuse to create the characteristic bicarpellate compound structure. The compound pistil structure reflects fusion of originally separate carpel primordia during early flower development, with fusion evident in multiple anatomical features= (1) unified ovary formed from fusion of two carpel ovary walls, creating either a single shared chamber (unilocular) or two chambers (bilocular) depending on degree of fusion, with the bilocular condition (two locules separated by central septum formed from fused carpel margins) representing the typical Vitis state (2) unified style formed from fusion of two carpel styles into a single coherent structure, evident in mature pistils as a single style rather than two separate styles (3) unified or bilobed stigma formed from fusion of two carpel stigmas, often showing residual two-part organization (bilobed appearance, bilateral papillae distribution) reflecting the bicarpellate origin even when structurally fused. Compound pistil development involves carpel primordium initiation as separate structures (BBCH 51-52) followed by rapid fusion during early organogenesis (BBCH 52-53) creating a unified pistil structure by the time ovule initiation occurs (BBCH 54-55), with fusion so complete and occurring so early that individual carpel identity is largely obscured in mature structures except through anatomical details like locule number and stigma organization. The evolutionary and developmental significance of compound pistil organization relates to reproductive efficiency and fruit structure optimization= carpel fusion creates a unified ovary that develops into a unified berry rather than separate fruit units, concentrating resources into single-fruit development with enhanced seed protection and dispersal characteristics style fusion creates a single pathway for pollen tube growth rather than requiring separate pollination events for each carpel, potentially increasing fertilization efficiency stigma fusion (or close apposition) creates a unified receptive surface increasing probability of pollen capture. The compound pistil in Vitis invariably comprises two carpels (bicarpellate compound pistil), distinguishing grapevine from many other plant families where compound pistils may comprise three, four, five, or more carpels occasional developmental abnormalities produce pistils with more than two locules suggesting involvement of more than two carpels, but such variants are extremely rare (<0.1% of flowers). The bicarpellate nature is most clearly evident in= (1) bilocular ovary structure with two distinct chambers visible in transverse sections (2) two rows of ovules corresponding to the two carpel margins forming the central septum (3) often bilobed stigma appearance with two-part organization (4) developmental observations showing two carpel primordia initiating before fusion. Compound pistil functionality depends on complete fusion achieving unified tissue organization supporting coordinated stigma receptivity, pollen tube guidance through unified style transmitting tract to both locules, and synchronized ovule development and fertilization, with fusion failures or incomplete fusion potentially impairing pistil function and fruit set.
Compound_Pistil represents the syncarpous organization characteristic of all normal Vitis flowers and must be distinguished from apocarpous pistils (separate unfused carpels) which do not occur in Vitis, and from simple pistils (derived from single carpel) which also do not represent normal Vitis structure. The compound nature reflects two-carpel fusion and is the ancestral Vitaceae condition, with the bicarpellate organization universal across Vitis species and cultivars. Sub-classification into Two_Carpel_Compound_Pistil and Fused_Carpel_Compound_Pistil reflects aspects of carpel number and fusion degree, while Bicarpellate_Pistil represents the specific two-carpel condition that characterizes Vitis. Understanding compound pistil organization is essential for interpreting ovary anatomy, ovule distribution, and fruit development patterns.
fused-carpel pistil
syncarpous pistil
synkarpes Gynoeceum
verwachsenes Fruchtblatt
zusammengesetzter Stempel
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 92-95 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 186-189.
Compound Pistil
A compound primary inflorescence at node 3 on a shoot of Vitis vinifera cv. Muscat of Alexandria in Cyprus, measuring 24.5 cm length representing very large dimensions, exhibiting highly complex compound paniculate architecture with central rachis bearing 22 major lateral branches, branching extending through 5-6 hierarchical orders creating intricate structure, major lateral branches showing extensive secondary branching with some secondary branches reaching 6-8 cm length themselves resembling simple inflorescence structures, tertiary and quaternary branches abundant creating very dense architecture, total flower number estimated at approximately 850 flowers organized throughout the elaborate three-dimensional structure, rachis diameter of 4.8 mm at base supporting the massive complex architecture, architectural elaboration creating overall conical to cylindrical profile with substantial width (8-10 cm) complementing the length, branching density creating somewhat compacted architecture with limited light penetration to interior regions, measured at BBCH stage 65 (full bloom) in late May, representing maximum compound primary inflorescence elaboration characteristic of very high fertility Muscat types requiring cluster thinning to prevent overcropping.
A primary inflorescence exhibiting complex highly elaborate branching architecture in Vitis vinifera, characterized by multiple orders of branching typically 4-6 orders creating intricate hierarchical structure, extensive lateral branch development with numerous major lateral branches often 15-25+ branches along the rachis, secondary and tertiary branch elaboration creating dense complex architecture, large overall dimensions typically 18-28 cm length reflecting the elaborate structure, high flower number often exceeding 500-800+ flowers organized across the branching network, well-developed branch systems showing substantial secondary branch development comparable to simple primary rachis in less elaborate inflorescences, and architectural complexity creating three-dimensionally elaborate paniculate structures rather than simple elongated forms. Compound primary inflorescences represent maximum architectural elaboration, occurring in highly fertile cultivars producing very large complex flowering structures (some Muscat types, fertile clones of various cultivars), exhibiting developmental investment creating extensive branching hierarchy, and producing very large clusters following fruit set requiring management intervention to prevent overcropping and maintain fruit quality.
Compound primary inflorescences are distinguished from standard primary inflorescences by exceptional architectural elaboration including high branch order, extensive lateral development, and large overall size. The compound designation emphasizes the complex highly branched architecture rather than simple or moderately branched structures. Branch order elevation to 5-6 levels creates exceptional hierarchical depth compared to typical 3-4 orders in standard inflorescences. Lateral branch elaboration with major laterals developing extensive secondary and tertiary branching creates architecture where secondary branches approach the complexity of entire simple inflorescences. Size correlates with elaboration, with compound inflorescences typically exceeding 20 cm and sometimes reaching 30+ cm in extreme cases. Flower number elevation reflects the architectural elaboration with >500-800 flowers common and exceptional examples exceeding 1000+ flowers. Occurrence shows strong cultivar effects with Muscat varieties and some highly fertile clones consistently producing compound architecture while most cultivars show moderate elaboration. Developmental investment requires substantial resources during both primordium formation and current-season development, with compound architecture indicating favorable conditions and high fertility. Cluster compactness following fruit set creates management challenges with very tight berry packing increasing disease susceptibility and requiring thinning or other interventions. Harvest implications include very large heavy clusters potentially requiring specialized handling and occasionally showing non-uniform ripening due to interior shading.
complex panicle
elaborate primary inflorescence
highly branched inflorescence
hochverzweigter Blütenstand
komplexe Rispe
zusammengesetzte primäre Infloreszenz
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 343-382 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 383-425 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 573-618.
Compound Primary Inflorescence
Compression resistance quality specification for export table grapes from Chile targets minimum 300 g force at 20% deformation for varieties intended for North American and Asian markets with 4-6 week transit plus retail display fruit below this threshold experiences unacceptable damage rates (above 5% crushed berries) from the compressive stresses of stacking in shipping containers, while fruit meeting specification (300-400 g) arrives with less than 2% compression damage, demonstrating how instrumental compression measurement predicts commercial handling outcomes.
Compression_Resistance_Quality represents the physical quality characterizing the mechanical response of grape tissue to compressive loading, capturing the firmness dimension most relevant to handling and storage where berries experience squeezing forces between adjacent berries, container surfaces, and handling equipment. Compression testing applies force perpendicular to the sample surface using a flat plate or curved probe, generating force-deformation data that characterizes how tissue responds to pressing. The compression response of intact berries reflects the integrated behavior of skin containing pressurized pulp, with initial resistance dominated by skin tension and turgor pressure, followed by progressive yielding as cells deform and eventually rupture. Key compression parameters include maximum force at defined deformation (firmness), slope of the force-deformation curve (stiffness), work to defined deformation (toughness), and bioyield point (force at initial structural failure). Compression resistance decreases during ripening as cell wall degradation and turgor changes reduce structural integrity, with the softening trajectory affecting susceptibility to compression damage during handling. The geometry of compression testing (parallel plate versus convex probe, deformation extent, test speed) substantially affects results, requiring standardization for comparable measurements. Compression damage in berries appears as crushed zones where cell integrity is lost, releasing juice and initiating oxidation, microbial colonization, and quality deterioration. The relationship between instrumental compression resistance and actual damage susceptibility during handling depends on the correspondence between test geometry and real-world stress conditions, with empirical validation needed for predictive application. Compression testing is technically straightforward, requiring only a texture analyzer with compression platen, making it accessible for routine quality assessment.
Compression_Resistance_Quality addresses the specific loading mode of squeezing forces, complementing puncture resistance which addresses penetrating forces. The test geometry (compression versus puncture) determines which tissue properties dominate the measurement compression integrates skin and pulp response while puncture can isolate skin break force. The relevance to handling damage makes compression resistance particularly important for table grape quality assessment and for wine grapes subject to mechanical harvest. The class describes the quality parameter rather than the test method, though the two are intimately linked.
Druckfestigkeit
Kompressionswiderstand
Quetschresistenz
Zusammendrückwiderstand
compression hardness
compressive firmness
crush resistance
squeeze resistance quality
Letaief H et al. (2008) Mechanical Behavior of Winegrapes under Compression Tests. American Journal of Enology and Viticulture 59(3): 323-329 Rolle L et al. (2012) Instrumental Texture Analysis Parameters as Markers of Table-Grape and Winegrape Quality: A Review. American Journal of Enology and Viticulture 63(1): 11-28
Compression Resistance Quality
Thompson Seedless bears a Concentrate_Production_Scion_Role globally, with its neutral flavor, high sugar content (22-24°Brix), and abundant production in California, Argentina, and South Africa supporting bulk white grape juice concentrate manufacturing—the concentrate serving as a sweetening agent in juice blends, wine production, and food formulations. Pedro Giménez serves in a Concentrate_Production_Scion_Role as Argentina's most important concentrate variety, with high yields, neutral character, and concentration in Mendoza and San Juan producing white grape concentrate for export to North American and European beverage industries. Airén bears a Concentrate_Production_Scion_Role in Spanish production, where its abundant yields on La Mancha's central plateau support both wine and concentrate production, with juice concentrate exported for beverage blending throughout Europe.
A juice production scion role that inheres in a grafted scion cultivar by virtue of its designation for grape juice concentrate production, where juice is extracted and concentrated (typically to 68-72°Brix) through evaporation or reverse osmosis for use as an industrial ingredient in beverages, food products, and wine amelioration. Concentrate production scions prioritize juice yield, sugar content, and flavor neutrality over the distinctive varietal character valued in direct-consumption juice, enabling use as a sweetening, coloring, and flavoring agent in diverse applications. High sugar content at harvest (22-26°Brix) maximizes concentrate yield per tonne of grapes, while neutral flavor profiles ensure versatility in blending without imparting unwanted varietal character. Grape juice concentrate serves as an ingredient in fruit juice blends, soft drinks, sports beverages, wine (for sweetening or chapitalization where permitted), and food products requiring natural sweetness or grape flavor. Major concentrate production occurs in Argentina, Spain, Italy, South Africa, and the United States, with global trade exceeding 500,000 tonnes concentrate equivalent annually.
Concentrate_Production_Scion_Role captures the industrial end of grape juice utilization where varietal neutrality and sugar content outweigh distinctive flavor characteristics. Concentrate pricing follows commodity markets (tracked by reference indices), with production responding to relative returns versus wine or fresh markets. The concentrate industry enables utilization of grapes from high-yielding vineyards unsuited to premium wine production, providing an economic floor for grape prices in bulk-producing regions. Color concentrate (from deeply pigmented varieties like Rubired, Salvador) represents a specialized subcategory used for coloring beverages and foods. Rectified concentrated grape must (RCGM) is a highly purified, deionized concentrate used in wine enrichment where regulations permit.
Industrie-Safttraube
Konzentrattraube
Saftkonzentrat-Traube
bulk juice grape
concentrate grape
industrial juice grape
processing grape
Morris JR, Striegler RK (2005) Grape juice, in Processing Fruits 2nd Ed, CRC Press pp.: 585-616 Creasy GL, Creasy LL (2009) Grapes, CABI OIV (2019) International Code of Oenological Practices, Grape Juice Concentrate USDA Foreign Agricultural Service (2020) Global Grape Juice Trade Report
Concentrate Production Scion Role
In Monastrell berries in Jumilla, Spain during BBCH 83-89, condensation reactions generate B-type procyanidin linkages at rates sufficient to increase skin tannin mDP from 24 to 32 over 38 days, with C4-C8 bonds comprising 85% and C4-C6 bonds 15% of total interflavonoid linkages as determined by NMR analysis, while oxidative conditions promote A-type bond formation reaching 8% of total linkages by harvest under hot conditions averaging 32°C that enhance oxidative coupling.
Condensation reactions describe the chemical bond-forming processes in Vitis vinifera berry tissues wherein flavan-3-ol monomers and oligomers undergo interflavonoid coupling to generate proanthocyanidin polymers of increasing molecular size, encompassing both acid-catalyzed condensation forming B-type procyanidin linkages through C4-C8 and C4-C6 bonds and oxidative coupling generating A-type linkages through additional C2-O-C7 bonds that establish the diverse structural features characterizing grape tannin populations. This process involves multiple reaction mechanisms operating under the acidic vacuolar conditions of pH 2.5-3.5 where proanthocyanidins accumulate, with acid-catalyzed condensation proceeding through protonation of the C-ring hydroxyl generating carbocation intermediates at C4 that undergo electrophilic attack on electron-rich A-ring positions of other flavan-3-ol units to form new carbon-carbon bonds extending polymer chains. The stereochemistry of condensation reactions influences product structure, with the configuration at C4 determining whether 4-alpha or 4-beta linkages form, and the regioselectivity between C6 and C8 attack positions generating structural diversity within the polymer population that affects physical properties and biological activities. Oxidative condensation represents an alternative mechanism generating A-type proanthocyanidins through radical coupling processes potentially catalyzed by polyphenol oxidases or occurring spontaneously in the presence of oxygen, producing doubly-linked units with both C4-C8 and C2-O-C7 bonds that demonstrate enhanced resistance to acid-catalyzed cleavage and modified sensory properties compared to B-type polymers. The kinetics of condensation reactions depend on substrate concentration, pH, temperature, and potential enzymatic catalysis, with reaction rates increasing at lower pH where protonation equilibria favor carbocation formation, at elevated temperatures that provide activation energy for bond formation, and in the presence of enzymes that may facilitate electron transfer in oxidative coupling. Non-enzymatic condensation proceeds continuously in acidic vacuolar compartments throughout berry development and post-harvest handling, with reaction extent determined by the duration of contact between reactive intermediates and potential nucleophiles in the proanthocyanidin-rich storage pools. The practical significance of condensation reactions extends beyond the berry into winemaking and wine aging, where continued condensation modifies tannin structure and properties over time, contributing to the evolution of astringency and mouthfeel characteristics that distinguish young wines from aged productions.
Condensation_Reactions is subclass of Tannin_Polymerization representing the specific bond-forming chemistry. Distinguishes from Tannin_Chain_Elongation which describes polymer growth outcome. Sibling to Tannin_Chain_Elongation and Tannin_Structure_Maturation. Includes acid-catalyzed B-type and oxidative A-type mechanisms. pH, temperature, and enzyme dependent. Continues during winemaking and aging.
Interflavanoid-Kondensation
Kondensationsreaktionen
Procyanidin-Kopplung
Tannin-Bindungs-Bildung
flavanol condensation
interflavonoid condensation
procyanidin coupling
tannin bond formation
Haslam E (1980) In vino veritas: oligomeric procyanidins and the ageing of red wines. Phytochemistry 19: 2577-2582 Es-Safi NE et al (1999) Competition between (+)-catechin and (-)-epicatechin in acetaldehyde-induced polymerization. Journal of Agricultural and Food Chemistry 47: 2088-2095 Kennedy JA et al (2006) Grape and wine phenolics. In Chemistry and Biology of Ellagitannins World Scientific pp 220-255
Condensation Reactions
Condensed tannin quality in Tannat from Madiran defines this variety's reputation as among the world's most tannic red grapes, with seed tannin concentrations of 80-120 mg/g seed dry weight (versus 40-60 mg/g for Cabernet Sauvignon) and skin tannins of 3-6 mg/g contributing to finished wine tannin levels of 3500-5000 mg/L modern Madiran winemaking employs micro-oxygenation and extended oak aging to polymerize these condensed tannins, transforming the aggressive astringency of young Tannat into the plush, velvety texture of age-worthy wines while preserving the tannic backbone enabling 30+ years of cellaring.
Condensed_Tannin_Quality represents the chemical quality characterizing the concentration and structural properties of proanthocyanidins in grape tissues and wine, capturing the flavan-3-ol polymers that constitute the dominant tannin class in grape-derived products. Condensed tannins (proanthocyanidins) are polymers of flavan-3-ol monomers linked through C4-C6 or C4-C8 interflavonoid bonds, with polymer sizes ranging from dimers through oligomers to high molecular weight polymers exceeding 30 subunits. The designation "condensed" reflects their formation through condensation reactions during biosynthesis and their behavior upon heating with acid, where cleavage releases anthocyanidin pigments (hence proanthocyanidins). In grape berries, condensed tannins accumulate in seeds and skins with distinct compositional profiles: seed tannins contain primarily catechin and epicatechin with epicatechin-3-gallate, while skin tannins include epigallocatechin reflecting F3'5'H activity. Seed tannins typically show lower mean degree of polymerization (mDP 5-15) than skin tannins (mDP 20-80), affecting extraction kinetics and sensory properties. The concentration of condensed tannins in grape seeds ranges from 2-8% of seed dry weight, while skin tannins typically contribute 1-4 mg/g skin fresh weight. During winemaking, condensed tannin extraction depends on ethanol concentration, temperature, maceration duration, and cap management, with seed tannin extraction accelerating as ethanol increases beyond 8-10% volume. Condensed tannin concentration in wine correlates with astringency perception but the relationship is modified by tannin structure, wine matrix, and individual sensitivity.
Condensed_Tannin_Quality specifically addresses proanthocyanidins as distinct from hydrolyzable tannins. The structural complexity of condensed tannins (variable polymer size, subunit composition, branching) means that concentration measurements alone incompletely characterize tannin quality. Phloroglucinolysis enables determination of both concentration and structural parameters (mDP, subunit composition, proportion galloylated). The distinction between seed-derived and skin-derived condensed tannins is important because they differ in composition, polymer size, and sensory properties, with skin tannins generally perceived as softer and more velvety than seed tannins.
Flavan-3-ol-Polymer-Qualität
Kondensiertes-Tannin-Gehalt
PA content quality
Proanthocyanidingehalt
Traubentanninkonzentration
flavan-3-ol polymer quality
grape tannin concentration
proanthocyanidin content
Kennedy JA et al. (2001) Composition of Grape Skin Proanthocyanidins at Different Stages of Berry Development. Journal of Agricultural and Food Chemistry 49(11): 5348-5355 Cheynier V et al. (2006) Structure and Properties of Wine Pigments and Tannins. American Journal of Enology and Viticulture 57(3): 298-305
Condensed Tannin Quality
In Chenin Blanc vines in the Loire Valley, conditional dormancy persists from early February through late March, with the conditional nature demonstrated by partial bud swelling during mid-February warm periods with temperatures of 12-15°C followed by growth arrest when temperatures return to 5-8°C, and eventual budbreak occurring in late March when sustained warming provides sufficient heat accumulation for irreversible growth commitment.
Conditional dormancy in Vitis vinifera represents the environment-dependent quiescent state characteristic of ecodormancy, whereby bud growth status is conditionally determined by prevailing environmental factors rather than by fixed internal regulatory states, with buds capable of transitioning between quiescence and active growth depending on whether environmental conditions are favorable or unfavorable for development. This conditional nature fundamentally distinguishes ecodormancy from the unconditional growth inhibition of endodormancy, where internal factors prevent growth regardless of environmental conditions until chilling-dependent release processes have been completed. The physiological basis of conditional dormancy involves the absence of internal growth-inhibiting mechanisms that were active during endodormancy, leaving environmental factors as the sole determinants of bud behavior, with growth-competent buds responding dynamically to temperature fluctuations and other environmental variables throughout the ecodormancy period. The conditionality is manifest in the immediate reversibility of quiescence, with buds transitioning toward growth when conditions become favorable and reverting to quiescence when conditions become unfavorable, creating the dynamic relationship between bud status and environment that characterizes the late winter and early spring period. The practical implications of conditional dormancy for viticulture relate to the variable and weather-dependent nature of budbreak timing, with warm late winters potentially advancing budbreak into periods of frost risk while cold springs may delay growth beyond optimal timing, creating management challenges that are absent during the more predictable endodormancy period. The molecular state during conditional dormancy reflects the completion of dormancy release, with low ABA concentrations, minimal DAM gene expression, derepressed growth gene promoters, and restored hormone signaling systems that enable rapid response to growth-promoting conditions when they occur. The duration of conditional dormancy varies with climate, extending for longer periods in cold continental regions where spring warming is delayed and for shorter periods in Mediterranean climates where mild temperatures follow closely after chilling completion, with the transition from conditional dormancy to active growth occurring when sustained favorable conditions accumulate sufficient heat units to trigger budbreak initiation.
Conditional_Dormancy represents a characteristic state within Ecodormancy describing the environment-dependent nature of growth control. Distinguished from unconditional endodormancy by the reversible, environment-responsive nature of quiescence. Subdivides into Environment_Dependent_Quiescence, Reversible_Growth_Suppression, and Weather_Responsive_Dormancy as component aspects. Duration varies with climate from weeks to months. Creates weather-dependent budbreak timing variability. Characterized by absence of internal inhibitory mechanisms.
bedingte Dormanz
conditional quiescence
environment-dependent dormancy
facultative dormancy
fakultative Dormanz
konditionale Ruhe
umweltabhängige Dormanz
weather-dependent dormancy
Lang GA et al (1987) Endo-, para-, and ecodormancy: physiological terminology and classification for dormancy research. HortScience 22: 371-377 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle
Conditional Dormancy
In Sauvignon Blanc vines in Marlborough, New Zealand, conditional growth suppression maintains ecodormancy through August and early September, with the conditional nature demonstrated by forcing test response showing budbreak within 11 days at 20°C and by field observations of partial bud swelling during warm periods in late August followed by growth arrest when cool conditions return, illustrating the reversible environment-dependent nature of suppression during ecodormancy.
Conditional growth suppression in Vitis vinifera represents the environment-dependent nature of growth inhibition during ecodormancy, whereby bud quiescence is maintained conditionally upon unfavorable environmental factors rather than unconditionally as during endodormancy, with growth suppression being immediately reversible upon provision of favorable conditions. This conditional nature distinguishes ecodormancy fundamentally from endodormancy, where growth suppression persists regardless of environmental conditions until internal changes associated with chilling release the dormant state. The mechanistic basis of conditional suppression involves the absence of internal inhibitory mechanisms, with growth-competent buds poised for immediate response to favorable signals, and suppression depending entirely on environmental temperatures being insufficient to support the metabolic rates required for growth processes. The reversibility of conditional suppression is demonstrated through forcing tests where transfer of ecodormant buds to favorable temperature immediately initiates growth processes leading to budbreak within 10-14 days, in contrast to endodormant buds where forcing conditions fail to reverse internal inhibition and budbreak requires extended periods or does not occur. The practical implications of conditional growth suppression relate to frost vulnerability, as the reversible nature of suppression means that temporary warm periods during late winter can trigger partial or complete growth initiation that is then damaged when temperatures return to suppressive levels, representing a significant difference from the more robust suppression during endodormancy that protects against such false spring responses. The intensity of conditional suppression varies continuously with environmental conditions, with strong suppression during cold periods, weakening suppression during mild conditions, and no suppression when temperatures exceed the threshold for growth support, creating the dynamic response to weather variability characteristic of the ecodormancy period. The transition from conditional to active growth occurs when favorable conditions persist long enough to accumulate sufficient heat units for budbreak initiation, at which point the previously suppressed growth processes proceed to visible developmental changes marking the end of dormancy and beginning of the new growing season.
Conditional_Growth_Suppression represents the reversible nature of growth inhibition within Environmental_Constraint. Distinguished from unconditional suppression during endodormancy which persists regardless of conditions. Demonstrated by immediate growth response to favorable forcing conditions. Creates frost vulnerability due to potential premature growth during warm periods. Intensity varies continuously with temperature conditions. Transition to growth when favorable conditions persist sufficiently for heat unit accumulation.
bedingte Ruhe
bedingte Wachstumsunterdrückung
environment-dependent inhibition
reversible Wachstumshemmung
reversible dormancy
reversible growth suppression
umweltabhängige Inhibition
Conditional Growth Suppression
Syrah grafted on 3309C rootstock at 19 years post-grafting showing conducting rootstock sapwood 2.8 cm radial thickness measured as depth of functional water-conducting tissue, hydraulic conductivity measurements indicating outer 1.8 cm conducting sapwood providing 68% of total transport capacity while inner 1.0 cm conducting sapwood providing remaining 32% showing conductivity gradient, dye injection experiments confirming entire conducting sapwood depth showing dye uptake and transport demonstrating active water conduction, conducting sapwood vessel patency estimated 85% in outer zone declining to 55% in inner zone, total conducting sapwood area 38.4 square cm supporting estimated maximum transport capacity 75-95 liters daily, conducting sapwood vessels 85-115 micrometers diameter characteristic for 3309C, conducting sapwood maintaining adequate water supply for mature canopy under normal conditions
Conducting rootstock sapwood is rootstock sapwood defined by active water transport function, emphasizing the conducting role of functional xylem vessels moving water and dissolved minerals from root system through rootstock trunk to scion regardless of sapwood age or position, typically encompassing all sapwood zones where vessels remain patent and capable of water conduction though transport efficiency varying from peak capacity in outer active sapwood to reduced capacity in older inner sapwood approaching heartwood conversion, and representing the essential hydraulic tissue enabling transpiration stream and scion canopy water supply. Conducting rootstock sapwood functional emphasis includes= defined primarily by water transport function (conducting sapwood identified by capacity to move water through patent vessels rather than anatomical position or age), encompassing all water-conducting xylem in rootstock trunk (conducting sapwood including any wood tissue where vessels remain open and capable of water transport from outer active sapwood to inner older sapwood), emphasizing hydraulic transport role (conducting sapwood conceptualized in terms of water conduction function supporting transpiration and canopy water supply), showing variable efficiency (conducting capacity ranging from peak efficiency in recently formed outer sapwood to reduced efficiency in older inner sapwood with some vessel occlusion), and corresponding to functional sapwood (conducting sapwood essentially synonymous with sapwood as functional conducting tissue distinct from non-conducting heartwood). Conducting rootstock sapwood vessel patency spectrum includes= containing fully patent vessels (many vessels completely open with no occlusion providing maximum transport capacity particularly in outer conducting sapwood), including partially occluded vessels (some vessels with partial tyloses or gum deposits reducing but not eliminating conductivity particularly in inner conducting sapwood), showing variable embolism levels (some proportion of vessels embolized containing air rather than water temporarily or permanently non-functional varying with environmental history), exhibiting efficient vessels (majority of conducting sapwood vessels maintaining adequate patency for water transport supporting canopy needs), and transitioning to occluded vessels at heartwood boundary (inner edge of conducting sapwood marked by complete vessel occlusion through tyloses formation eliminating transport function). Conducting rootstock sapwood hydraulic conductivity gradient includes= showing highest conductivity in outer zone (outermost conducting sapwood typically 1.5-3 cm providing maximum transport efficiency per unit area), displaying declining conductivity moving inward (transport efficiency gradually decreasing in progressively older conducting sapwood as vessel occlusion increases and embolism accumulates), exhibiting variable inner boundary (depth where conducting capacity becomes negligible varying with trunk age, rootstock variety, and stress history typically 2-5 cm from cambium), maintaining adequate total conductivity (integrated transport capacity across all conducting sapwood zones sufficient to support canopy transpiration demands), and demonstrating functional redundancy (multiple conducting zones providing parallel transport pathways ensuring continued function despite localized vessel failure). Conducting rootstock sapwood water transport dynamics includes= moving water upward through transpiration-driven negative pressure (conducting sapwood vessels under tension pulling water from roots upward to evaporating leaf surfaces), transporting 50-100+ liters daily during growing season (total conducting sapwood capacity adequate for peak transpiration rates in mature productive vine), showing diurnal transport variation (water flow rates through conducting sapwood fluctuating with daily transpiration cycles maximum during midday minimum at night), exhibiting seasonal patterns (transport activity highest during active growing season declining during dormancy when transpiration minimal), and responding to environmental conditions (conducting sapwood flow rates adjusting to temperature, humidity, and canopy water demand maintaining plant water status). Conducting rootstock sapwood dissolved mineral transport includes= conducting nitrogen, phosphorus, and potassium (major macronutrients absorbed by roots transported in xylem sap through conducting sapwood to leaves and fruit), moving calcium and magnesium (secondary macronutrients moving exclusively in xylem through conducting sapwood since phloem-immobile), transporting micronutrients (iron, manganese, zinc, copper, boron, and molybdenum moving through conducting sapwood from root absorption sites to metabolically active tissues), showing concentration dynamics (mineral concentrations in xylem sap flowing through conducting sapwood varying with root uptake rates and transpiration dilution), and enabling tissue nutrition (conducting sapwood mineral transport providing essential nutrients for photosynthesis, growth, and fruit development throughout vine). Conducting rootstock sapwood in transpiration stream includes= serving as conduit for transpiration stream (conducting sapwood vessels providing continuous pathway from roots through trunk to leaves for water movement driven by leaf evaporation), maintaining negative pressure gradients (conducting sapwood water columns under tension typically -5 to -20 bar during active transpiration), exhibiting cohesion-tension mechanism (water molecules cohering through hydrogen bonding maintaining continuous columns through conducting sapwood under tension without breaking), showing vulnerability to cavitation (conducting sapwood water columns potentially breaking under extreme tension introducing air embolisms disrupting transport), and demonstrating refilling capacity (embolized vessels in conducting sapwood potentially refilling through positive root pressure or localized repair mechanisms restoring transport function). Conducting rootstock sapwood efficiency factors includes= showing vessel size effects (larger vessels in outer conducting sapwood providing disproportionately high transport capacity due to flow rate increasing with fourth power of radius), displaying vessel density influence (number of conducting vessels per unit area affecting total transport capacity independent of individual vessel size), exhibiting vessel connectivity importance (degree of connection between vessels affecting path length and resistance influencing overall conducting efficiency), demonstrating anatomical optimization (conducting sapwood vessel size distribution and arrangement generally optimized for efficient water transport), and varying with rootstock genetics (cultivar-specific conducting sapwood anatomy affecting inherent transport efficiency and maximum capacity). Conducting rootstock sapwood seasonal embolism and repair includes= accumulating winter embolism (freeze-thaw cycles during dormancy potentially introducing air emboli in conducting sapwood vessels reducing functional capacity), showing spring refilling (recovery of conducting capacity as embolized vessels refill during spring through root pressure and/or localized repair mechanisms), exhibiting drought-induced summer cavitation (severe water stress potentially causing vessel cavitation creating air-filled non-conducting vessels), demonstrating variable recovery (some embolized conducting sapwood vessels refilling and recovering function while others remaining permanently non-functional), and requiring functional redundancy (multiple parallel conducting pathways in sapwood ensuring adequate transport despite some vessel embolism). Conducting rootstock sapwood measurement approaches includes= determining hydraulic conductivity through flow measurements (quantifying water transport capacity under controlled pressure gradients identifying functional conducting tissue), using dye injection to trace pathways (introducing colored solutions showing which conducting sapwood vessels actively transporting water), applying staining techniques to assess vessel patency (histological examination revealing open versus occluded vessels in conducting tissue), measuring sap flow with sensors (thermal or other methods quantifying real-time water transport through conducting sapwood in field conditions), and estimating from sapwood area (calculating potential conducting capacity from functional sapwood cross-sectional area and vessel characteristics). Conducting rootstock sapwood in different trunk ages includes= young trunk (years 1-7) showing all wood as conducting (no heartwood formation yet with entire wood cross-section potentially conducting), establishing trunk (years 8-15) developing conducting-non-conducting differentiation (outer sapwood conducting while inner wood beginning heartwood conversion losing conductivity), mature trunk (years 15-40) displaying distinct conducting sapwood ring (clear zone of patent vessels surrounding central heartwood with occluded non-conducting vessels), old trunk (years 40+) potentially showing narrower conducting zone (limited conducting sapwood thickness surrounding large heartwood core in aged trunk), and very old trunk possibly showing minimal conducting capacity (very narrow conducting sapwood ring potentially limiting maximum water transport in ancient trunk). Conducting rootstock sapwood and trunk transport capacity includes= determining maximum vine water supply (conducting sapwood area and efficiency limiting peak transpiration rates supportable by trunk), influencing canopy size potential (conducting capacity constraining maximum leaf area that can be adequately supplied with water), affecting drought tolerance (conducting sapwood reserve capacity providing buffer allowing vine to maintain function during moderate water stress), showing relationship to productivity (adequate conducting capacity necessary for supporting large fruit loads with high berry water content), and indicating trunk functional status (conducting sapwood extent and condition reflecting overall trunk hydraulic health and capacity). Conducting rootstock sapwood stress responses includes= exhibiting vulnerability to severe drought (extreme water stress potentially causing extensive cavitation disrupting conducting sapwood function), showing freeze damage susceptibility (severe cold potentially causing conducting sapwood vessel damage or embolism reducing transport capacity), demonstrating disease impacts (vascular pathogens potentially blocking or damaging conducting sapwood vessels limiting water supply to canopy), responding to mechanical damage (injuries potentially disrupting conducting sapwood pathways though compartmentalization limiting damage spread), and recovering through compensation (remaining functional conducting sapwood vessels potentially increasing flow rates compensating for damaged non-functional tissue). Conducting rootstock sapwood in rootstock cultivars includes= Riparia types showing efficient narrow conducting sapwood (smaller but numerous vessels providing adequate conductivity in compact zone), Berlandieri hybrids exhibiting broad conducting sapwood (large vessels in thick sapwood ring providing high transport capacity), vigorous rootstocks like 1103P displaying wide conducting zones (broad conducting sapwood with large vessels supporting high-vigor vine water demands), low-vigor rootstocks like 101-14 showing narrower conducting zones (limited conducting sapwood appropriate for restricted vine size), and cultivar-specific anatomical traits (vessel size distribution, density, and arrangement characteristic of rootstock variety affecting conducting efficiency). Conducting rootstock sapwood and graft union includes= maintaining vascular continuity across graft union (conducting sapwood vessels in rootstock connecting with conducting sapwood vessels in scion enabling uninterrupted water flow), showing potential mismatch at graft interface (differences in conducting sapwood vessel size or density between rootstock and scion potentially creating resistance at graft union), exhibiting graft union effects on flow (callus tissue at graft interface potentially affecting conducting pathway efficiency requiring adequate integration for optimal function), demonstrating compatibility effects (poorly compatible rootstock-scion combinations potentially showing conducting tissue discontinuities reducing water transport efficiency), and requiring complete integration (successful graft union with well-connected conducting sapwood essential for adequate trunk transport capacity). Conducting rootstock sapwood and vine water relations includes= maintaining plant water potential (conducting sapwood transport capacity enabling vine to maintain adequate tissue hydration under transpirational demand), showing diurnal water potential patterns (conducting sapwood transport limitations potentially contributing to midday water potential declines under high transpiration), exhibiting water stress responses (conducting sapwood capacity determining vine sensitivity to drought and canopy water stress symptoms), demonstrating recovery dynamics (conducting sapwood refilling and restored transport enabling vine water status recovery following stress relief), and influencing stress tolerance (adequate conducting sapwood reserve capacity contributing to vine drought tolerance and resilience). Conducting rootstock sapwood significance includes= providing essential water transport pathway (conducting sapwood vessels enabling water delivery from roots to transpiring canopy essential for photosynthesis and growth), determining trunk hydraulic capacity (conducting sapwood cross-sectional area and efficiency limiting maximum water transport rate trunk can support), delivering dissolved minerals to canopy (conducting sapwood serving as conduit for nutrient transport from root absorption to leaves and fruit), maintaining transpiration stream continuity (conducting sapwood vessels preserving continuous water columns from soil to atmosphere), exhibiting functional definition emphasis (conducting sapwood concept emphasizing active transport function identifying tissue by capability rather than just anatomical position), and representing critical limitation (conducting sapwood capacity potentially limiting vine size, productivity, or stress tolerance when compromised).
Conducting_Rootstock_Sapwood emphasizes active water transport function and is distinguished from Active_Rootstock_Sapwood (peak-efficiency outer zone) and Rootstock_Sapwood (anatomical sapwood without transport function emphasis) by focusing on hydraulic conducting role across all functional zones. Critical for understanding water transport capacity, transpiration stream pathways, hydraulic conductivity, vessel function, trunk transport limitations, and the essential conducting tissue providing water supply enabling canopy function and productivity.
Hydraulically active rootstock sapwood
Hydraulisch aktives Unterlagenholz
Leitendes Unterlagen-Splintholz
Transporting rootstock sapwood
Wassertransportierendes Wurzelstockholz
Water-conducting rootstock sapwood
Conducting Rootstock Sapwood
Cabernet Sauvignon conducting sapwood at 12 years post-planting comprises 18-24 mm radial depth with outermost 6-8 annual rings, contains 85-95% patent vessels in outer rings declining to 40-60% patent in inner rings, large earlywood vessels 180-260 μm diameter providing 65-75% of total hydraulic conductance, specific hydraulic conductivity 4.2-6.8 kg m⁻¹ MPa⁻¹ s⁻¹, conducting sapwood cross-sectional area 520-780 mm² supporting 38-62 L/day water transport at -0.8 MPa xylem tension, and showing progressive summer embolism reducing conductance 25-40% by late season
Conducting sapwood is sapwood emphasizing active water transport function through patent xylem vessels, representing the wood zone responsible for hydraulic conductance from roots to canopy and determining vine water relations capacity, transpiration support, and drought stress susceptibility. Conducting sapwood comprises the portion of sapwood with functional vessel lumens= outermost 4-8 annual rings typically containing patent vessels (extent varying by vine age, species, environmental history), vessels free of air embolisms under normal conditions (water-filled and capable of transport), vessels without tyloses or gum occlusions (maintaining open lumens for water flow), and vessel walls with functional pit membranes (enabling vessel-to-vessel water movement). Conducting sapwood vessel characteristics include= large-diameter earlywood vessels (120-300 μm) providing majority of hydraulic conductance (single large vessel contributing more to flow than hundreds of small vessels due to fourth-power relationship between radius and flow in Hagen-Poiseuille equation), smaller latewood vessels (40-120 μm) providing additional capacity and safety (less prone to embolism, maintaining some function when large vessels cavitate), vessel density 20-60 vessels/mm² (varying by wood position and growth rate), vessel lumen occupying 25-45% of conducting sapwood cross-sectional area (remaining area comprising fibers and parenchyma), and vessel element length 200-600 μm with perforated end walls (creating continuous water columns). Conducting sapwood hydraulic conductance includes= specific hydraulic conductivity 2-8 kg m⁻¹ MPa⁻¹ s⁻¹ (varying by cultivar, growth conditions, and embolism extent), total hydraulic conductance proportional to conducting sapwood cross-sectional area (typical mature vine with 400-900 mm² conducting sapwood area providing 25-80 L/day capacity at 0.5-1.5 MPa tension), earlywood vessels contributing 60-80% of total conductance (despite being only 30-50% of vessel number), and conductance declining with distance from cambium (outer rings showing higher conductivity than inner conducting rings approaching heartwood transition). Conducting sapwood water transport mechanisms include= cohesion-tension theory (water columns under tension pulled by transpirational demand at leaves), xylem sap ascent from roots to canopy through continuous vessel lumens (water moving 0.1-5 m/hour depending on demand), lateral water movement through bordered pits (enabling water to bypass embolized or damaged vessels), and capacitance (stored water in parenchyma cells buffering rapid transpirational demands). Conducting sapwood vulnerability to embolism includes= air seeding through pit membranes when tension exceeds pit strength (typically at xylem water potentials -1.5 to -3.0 MPa depending on vessel size), large vessels more vulnerable to embolism (greater pit membrane area and higher tensions increasing cavitation probability), freeze-thaw embolism from ice crystal formation disrupting water columns (particularly damaging in late fall or early spring), and progressive embolism accumulation reducing conductance (30-60% conductance loss common during growing season, varying by water stress intensity and duration). Conducting sapwood embolism patterns include= preferential cavitation of large earlywood vessels (greatest hydraulic contribution but highest vulnerability), latewood vessels maintaining function longer (serving as backup conducting capacity), radial gradient with inner conducting sapwood showing higher embolism (older wood more vulnerable or previously embolized), and seasonal embolism cycling (accumulation during summer stress, potential winter refilling though limited in grapevine, new conducting wood formation each spring providing restored capacity). Conducting sapwood relationship to canopy includes= conducting sapwood area correlating with maximum supportable leaf area (Huber value, leaf area/sapwood area ratio typically 1.5-3.5 m² leaf per cm² conducting sapwood), hydraulic sufficiency determining canopy water status (adequate conducting capacity maintaining leaf water potential, insufficient capacity causing water stress), and conducting sapwood area limiting productivity potential (crop load and canopy size ultimately limited by hydraulic supply capacity). Conducting sapwood seasonal dynamics include= spring restoration of function (new ring production providing fresh conducting wood, potential limited refilling of previous year's embolism), summer peak transport (maximum sap flow rates 30-100 L/day supporting transpiration 2-6 L/m² leaf area/day), progressive summer embolism (declining conductance through growing season as stress accumulates), and fall-winter cessation (reduced water demand, potential freeze-induced embolism, minimal transport during dormancy). Conducting sapwood measurement and assessment includes= dye perfusion revealing functional vessel distribution (safranin or other dyes taken up by conducting vessels), hydraulic conductivity measurement using pressure-flow apparatus (quantifying transport capacity per unit pressure gradient), vulnerability curve analysis (measuring conductance loss across water potential gradient), and xylem sap flow measurement using heat pulse or thermal dissipation sensors (quantifying actual transport in field conditions). Conducting sapwood significance in viticulture includes= determining vine drought tolerance (adequate conducting sapwood maintaining function under water deficit, insufficient conducting capacity causing severe stress), limiting irrigation requirements (conducting sapwood capacity affecting water demand and irrigation scheduling), indicating trunk disease impacts (diseases reducing conducting sapwood causing water stress symptoms), and constraining vine size and productivity (conducting area ultimately limiting sustainable canopy and crop load).
Conducting_Sapwood emphasizes water transport function and is distinguished from Active_Sapwood (metabolic and storage emphasis) and Sapwood (complete functional zone including storage) by focus on hydraulic conductance through patent vessels. Critical for understanding vine water relations, drought stress physiology, canopy water status, embolism vulnerability, and hydraulic limitations on vine size and productivity.
Functional xylem vessels
Hydraulic wood
Hydraulisch aktives Xylem
Leitendes Splintholz
Wassertransport-Holz
Water-transporting sapwood
Tyree, M. T., & Zimmermann, M. H. (2002). Xylem Structure and the Ascent of Sap (2nd ed.). Springer. | Choat, B., Drayton, W. M., Brodersen, C., Matthews, M. A., Shackel, K. A., Wada, H., & McElrone, A. J. (2010). Measurement of vulnerability to water stress-induced cavitation in grapevine= A comparison of four techniques applied to a long-vesseled species. Plant, Cell & Environment, 33(9), 1502-1512.
Conducting Sapwood
Chardonnay conducting vascular connection zone at early véraison (BBCH 81-83) shows declining xylem conduction (0.15-0.30 mg water/berry/hour, 30-50% of vessels embolized) and increasing phloem conduction (1.0-1.6 mg sucrose/berry/hour supporting 0.8-1.2°Brix/day berry sugar increase), with total solute import providing osmotic drive for continued berry expansion despite reduced xylem water import
A conducting vascular connection zone is a vascular connection zone emphasizing active physiological transport function through xylem water and mineral conduction and phloem sugar and nutrient conduction, representing functional vascular pathways during periods of active berry growth and ripening. Conducting vascular connection zone characteristics vary dramatically by developmental phase reflecting shifting berry nutritional requirements= during Phase I berry growth (BBCH 73-77, 0-6 weeks post-fruit set) xylem conduction dominates with high water import rates supporting rapid berry cell division and expansion, during Phase II lag phase (BBCH 77-81, 6-8 weeks post-fruit set) both xylem and phloem conduct actively supporting slower berry growth and seed development, during Phase III ripening (BBCH 81-89, 8-16 weeks post-fruit set) phloem conduction increasingly dominates with high sugar import while xylem conduction declines or reverses. Xylem conduction in conducting vascular connection zone includes= water transport through vessel elements (hollow dead cells with lignified walls forming continuous water columns), mineral transport (calcium, potassium, nitrogen, other nutrients dissolved in xylem sap), hormone transport (cytokinin synthesis in roots transported to berries via xylem), and hydraulic signaling (water potential gradients affecting berry turgor and growth regulation). Xylem transport rates vary= Phase I peak rates 0.3-0.8 mg water/berry/hour supporting 50-100 μL/day berry volume increase, Phase II reduced rates 0.1-0.3 mg/berry/hour with slower growth, Phase III minimal rates or backflow as berry sugar accumulation creates osmotic gradient driving water export from berry. Phloem conduction in conducting vascular connection zone includes= sugar transport (primarily sucrose at 150-300 mM concentration in phloem sap), amino acid and nitrogen compound transport (supporting berry protein synthesis), hormone transport (auxin, gibberellin, abscisic acid in phloem sap), and mineral transport (particularly potassium accumulation during ripening). Phloem transport rates vary= Phase I moderate rates 0.2-0.5 mg sucrose/berry/hour, Phase II rates 0.3-0.8 mg/berry/hour with increasing importance, Phase III peak rates 0.8-2.5 mg/berry/hour supporting rapid sugar accumulation (0.5-1.5°Brix/day increase in berry soluble solids). Conducting vascular connection zone transport capacity is influenced by= vascular bundle number and size (greater vascular cross-section provides higher potential transport rates), environmental conditions (temperature affecting metabolic activity, water status affecting xylem transport), source-sink relationships (photosynthate availability, competing sinks affecting phloem transport), and developmental regulation (hormone and gene expression changes modulating transport activities). Conducting vascular connection zone dysfunction or modification includes= xylem embolism reducing hydraulic conductance (air bubbles breaking water columns particularly under drought stress or during rapid berry expansion), phloem callose deposition potentially restricting sieve tube transport, vascular blockage through tyloses or gums reducing conducting capacity, and abscission zone development disrupting vascular continuity at berry maturity. Measurement of conducting vascular connection zone function includes= xylem sap flow measurement through heat pulse or dye tracer techniques, phloem transport estimation through carbon isotope labeling or sugar accumulation rates, hydraulic conductance measurement through pressure-flow relationships, and microscopic observation of functional versus embolized xylem vessels.
Conducting_Vascular_Connection_Zone emphasizes active physiological transport and is distinguished from Primary_Vascular_Connection_Zone (structural pathway emphasis) and Vascular_Connection_Zone (complete architecture) by focus on functional conducting activity and phase-specific transport patterns. Critical for understanding berry growth and ripening physiology, water relations during berry development, sugar accumulation mechanisms and rates, and environmental or stress effects on berry development through vascular transport modulation.
Active vascular pathway
Aktive Transportbahnen
Funktionierende Leitbündel
Leitende Gefäßverbindungszone
Transporting vascular zone
Conducting Vascular Connection Zone
Cabernet Sauvignon clusters display archetypal conical form with pronounced shoulder development and progressive tapering to a relatively narrow tip, producing length-to-width ratios of 1.6-2.0 and shoulder-to-tip width ratios of 2.5-3.5 the small berry size (1.0-1.4 g) and thick skins characteristic of this cultivar interact with the conical architecture to create compact shoulders that require careful canopy management for disease prevention while the looser tips dry quickly after precipitation, exemplifying how cluster shape integrates with berry traits to determine cultivar-specific disease dynamics.
Conical_Quality represents the overall form characteristic of organs exhibiting geometry that tapers progressively from a broad base or shoulder region toward a narrower apex, creating a cone-like or pyramidal appearance. In grapevine contexts, conical form applies primarily to cluster architecture and represents one of the most common bunch shapes across Vitis vinifera cultivars. Conical clusters exhibit maximum width at or near the shoulder region (proximal end where the primary rachis branches emerge) and progressively decreasing width toward the cluster tip, with length-to-width ratios typically ranging from 1.5-2.5 depending on the degree of elongation. The tapering results from the developmental pattern where proximal branches set more fruit or develop larger berries than distal branches, creating the characteristic pyramid profile. Classic conical-clustered cultivars include Cabernet Sauvignon, Syrah, and Pinot Noir, though the sharpness of the taper varies among cultivars and with growing conditions. Conical cluster form affects compactness because the tapering concentrates berries in the shoulder region while the tip remains relatively loose. Winged conical form, where one or two pronounced lateral branches (wings) extend from the shoulder, represents a common variant with implications for cluster handling and disease management. Quantitative assessment of conical form employs the shoulder-to-tip width ratio combined with overall length-to-width ratio, with categorical boundaries distinguishing short conical, conical, and long conical variants.
Conical_Quality is the most frequent cluster shape category in Vitis vinifera and serves as the reference against which other shapes (cylindrical, funnel-shaped) are compared. OIV descriptor 208 category 2 (conical) provides the standardized ampelographic reference. The presence and size of wings (lateral shoulder branches) modifies the basic conical form and is captured in OIV descriptor 209. Conical berry form, while theoretically possible, is not recognized in standard ampelographic descriptions berries with pointed apices are classified under distinct shape categories. The sharpness of the taper (acute versus obtuse cone angle) provides additional shape information beyond the basic conical classification.
Kegelform
cone-shaped quality
funnel-form
konische Qualität
pyramidal shape
pyramidenförmige Gestalt
sich verjüngende Form
tapered form
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 87-88 Tello J and Ibáñez J (2018) What Do We Know about Grapevine Bunch Compactness? A State-of-the-Art Review. Australian Journal of Grape and Wine Research 24(1): 6-23
Conical Quality
Zierfandler (Spätrot) in the Thermenregion of Austria bearing a conservation priority role was identified through the Austrian National Action Plan for Plant Genetic Resources as a high-priority conservation target based on its unique parentage (Roter Veltliner × unknown), restriction to fewer than 85 hectares concentrated almost entirely within a 20 km radius of Gumpoldskirchen, documented decline from 200+ hectares in the 1960s, irreplaceable contribution to the distinctive Gumpoldskirchner Spätrot-Rotgipfler blend tradition, and assessed breeding value as a source of late-ripening and botrytis-tolerant genetic material, resulting in prioritized inclusion in the Bundesamt für Weinbau Klosterneuburg conservation program with 12 authenticated clones under long-term maintenance.
A conservation role that inheres in a grapevine cultivar that has been formally identified through systematic assessment as warranting prioritized conservation action based on evaluation criteria including genetic uniqueness, threat severity, cultural significance, potential breeding value, and the feasibility of effective conservation intervention.
Conservation_Priority_Role formalizes the triage process necessary when conservation resources are insufficient to protect all threatened cultivars simultaneously. Priority assessment integrates multiple criteria that cannot be reduced to a single endangerment metric: a moderately rare cultivar of high genetic uniqueness (no close relatives, unique alleles) and irreplaceable cultural significance may receive higher conservation priority than a critically endangered cultivar closely related to well-conserved varieties. Systematic prioritization frameworks have been developed by ECPGR (European Cooperative Programme for Plant Genetic Resources), national genebank programs, and heritage cultivar organizations such as ProSpecieRara and Slow Food Presidia.
Erhaltungsprioritäts-Rolle
Schutzprioritäten-Rolle
conservation target role
high-priority genetic resource role
priority conservation role
vorrangige Genressourcen-Rolle
ECPGR (2019) European Cooperative Programme for Plant Genetic Resources — Vitis Working Group Report FAO (2010) Second Report on the State of the World's Plant Genetic Resources for Food and Agriculture, Rome Maul E et al. (2012) Identification and characterization of grapevine genetic resources, Vitis 51: 53-63
Conservation Priority Role
Gouais Blanc vines maintained at the Conservatoire du Vignoble Charentais bearing a conservation role represent one of fewer than 50 surviving plantings worldwide of a cultivar identified as the maternal parent of Chardonnay, Gamay, Riesling, Aligoté, and at least 75 other modern cultivars through DNA parentage analysis, with conservation management including phytosanitary monitoring, annual propagation of backup material, cryopreservation of shoot tips at -196°C in liquid nitrogen, and SSR-verified identity confirmation on a 3-year cycle to prevent accidental loss of this genetically irreplaceable ancestor of European viticulture.
A grapevine role that inheres in a vine by virtue of its recognized significance for the preservation of grapevine genetic diversity, where the vine's conservation value — whether due to rarity, endangerment, historical importance, or unique genetic constitution — motivates deliberate protective management actions that would not be justified by commercial production considerations alone.
Conservation_Role classifies grapevines according to their value for genetic resource preservation, applying conservation biology principles to cultivated plant diversity. The role addresses the documented erosion of grapevine cultivar diversity over the past century: an estimated 50-70% of cultivars historically grown in European wine regions have disappeared from commercial cultivation since the phylloxera crisis and subsequent replanting with a narrow range of internationally marketable varieties. Subclasses capture distinct conservation status categories (endangered, rare), conservation strategies (ex situ, in situ), and recovery outcomes (recovered cultivars returned to cultivation from near-extinction).
Artenschutz-Rolle
Erhaltungs-Rolle
Sortenschutz-Rolle
conservation status role
cultivar conservation role
genetic preservation role
This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 FAO (2010) Second Report on the State of the World's Plant Genetic Resources for Food and Agriculture, Rome Lacombe T et al. (2018) Grapevine European catalogue: towards a comprehensive list, Vitis 57: 69-76
Conservation Role
Systematic quantification of Constitutive_Defense_Disposition at INRAE Colmar compared basal defense parameters in healthy, unchallenged leaves across 30 cultivars representing diverse genetic backgrounds. Physical parameters measured included cuticle thickness (transmission electron microscopy), adaxial and abaxial trichome density (light microscopy), stomatal density and aperture dimensions (SEM), and leaf toughness (penetrometer). Chemical parameters included total phenolics (Folin-Ciocalteu), flavonoid content (spectrophotometric), proanthocyanidin concentration (DMACA assay), and basal PR protein activity (chitinase, glucanase enzyme assays). V. vinifera cultivars showed wide variation in Constitutive_Defense_Disposition, with 'Gewürztraminer' exhibiting high physical investment (cuticle 4.8 μm, trichome density 42/mm²) but low chemical defense (total phenolics 18 mg/g), while 'Tannat' showed the inverse pattern (cuticle 2.1 μm, trichomes 8/mm², but phenolics 52 mg/g). The PIWI cultivar 'Solaris' demonstrated high constitutive chemical defense (phenolics 48 mg/g, proanthocyanidins 12 mg/g) combined with moderate physical barriers. Wild species V. riparia exhibited the highest overall Constitutive_Defense_Disposition with thick cuticle (5.2 μm), dense trichomes (65/mm²), and high phenolics (58 mg/g). Correlation analysis revealed that physical and chemical constitutive defenses were largely independent (r = 0.18), suggesting separate genetic control and potential for combining both defense modes through breeding.
A Constitutive_Defense_Disposition is a Defense_Mechanism_Disposition that inheres in a Vitis vinifera plant and is realized through preformed defensive structures, compounds, or properties that are present regardless of pathogen or pest attack, providing a first line of defense without requiring recognition of specific threats or activation of induced responses. This disposition represents the baseline defensive capacity present in healthy, unchallenged plants, encompassing both physical barriers that impede attacker access or establishment and chemical defenses that are toxic, deterrent, or antimicrobial without requiring de novo synthesis upon attack. Constitutive defenses in grapevine include structural features such as cuticle thickness and composition, trichome density and morphology, stomatal characteristics, cell wall composition and lignification, and tissue toughness, as well as preformed chemical defenses including phenolic compounds (flavonoids, hydroxycinnamic acids, tannins), antimicrobial proteins (chitinases, glucanases, thaumatin-like proteins at basal levels), and secondary metabolites with deterrent or toxic properties. The relative investment in constitutive versus induced defenses represents a fundamental trade-off in plant defense strategy, with constitutive defenses providing immediate protection but incurring ongoing metabolic costs, while induced defenses are metabolically economical but require time for deployment. Grapevine cultivars vary substantially in constitutive defense investment, with some relying primarily on preformed barriers while others emphasize rapid, strong induced responses, and the most resistant genotypes often combining both strategies effectively.
Constitutive_Defense_Disposition is_a Defense_Mechanism_Disposition. This class encompasses all preformed defenses present without pathogen/pest challenge, contrasting with Induced_Defense_Disposition requiring activation. Child classes include Physical_Barrier_Disposition (structural defenses) and Chemical_Defense_Disposition (preformed compounds). Constitutive and induced defenses may act synergistically, with constitutive barriers slowing initial infection while induced responses mount, or may represent alternative defense strategies varying among genotypes.
Basale-Abwehr-Kapazität
Konstitutive-Verteidigungs-Disposition
Präformierte-Abwehr
basal defense capacity
constitutive resistance
preformed defense
Wittstock U, Gershenzon J (2002) Constitutive plant toxins and their role as defenses against herbivores and pathogens, Current Opinion in Plant Biology 5:300-307 Underwood W (2012) The plant cell wall: a dynamic barrier against pathogen invasion, Frontiers in Plant Science 3:85 Lattanzio V et al. (2006) Role of phenolics in the resistance mechanisms of plants against fungal pathogens and insects, Phytochemistry: Advances in Research, Research Signpost
Constitutive Defense Disposition
Evaluation of Constitutive_Phenolic_Disposition effects on multiple pathogen resistance at UC Davis characterized phenolic profiles and disease susceptibility across 25 cultivars representing the phenolic content range in cultivated grapevine. Total phenolics, measured by Folin-Ciocalteu assay on healthy leaf tissue, ranged from 14 mg GAE/g ('Chasselas') to 62 mg/g ('Petit Verdot'). HPLC profiling quantified major compound classes including hydroxycinnamic acids, flavonols, flavan-3-ols, and stilbenes at basal levels. Controlled inoculation experiments tested resistance to P. viticola, E. necator, and B. cinerea. Multiple regression revealed that Constitutive_Phenolic_Disposition parameters explained 35-55% of resistance variation depending on pathogen. Flavan-3-ol content (catechin, epicatechin) was the strongest predictor of E. necator resistance (partial r² = 0.28), consistent with in vitro inhibition of conidial germination by these compounds. Hydroxycinnamic acid content predicted P. viticola resistance (partial r² = 0.21), while proanthocyanidin levels correlated most strongly with B. cinerea resistance (partial r² = 0.32), reflecting the importance of tannins in restricting necrotrophic pathogen colonization. The high-phenolic cultivar 'Petit Verdot' (62 mg/g total phenolics) showed 40-60% less infection across all three pathogens compared to low-phenolic 'Chasselas' (14 mg/g), demonstrating that strong Constitutive_Phenolic_Disposition provides broad-spectrum, though partial, resistance enhancement.
A Constitutive_Phenolic_Disposition is a Chemical_Defense_Disposition that inheres in a Vitis vinifera plant and is realized through high basal accumulation of phenolic compounds—secondary metabolites derived from the phenylpropanoid pathway—providing broad-spectrum antimicrobial, antioxidant, and anti-herbivore defense properties. This disposition represents one of the most important chemical defense systems in grapevine, as Vitis is notably rich in diverse phenolic compounds including flavonoids (flavonols, flavan-3-ols, anthocyanins), non-flavonoid phenolics (hydroxycinnamic acids, hydroxybenzoic acids, stilbenes), and polymeric phenolics (proanthocyanidins/condensed tannins, lignin). Constitutive phenolic defense operates through multiple mechanisms including direct antimicrobial activity (membrane disruption, enzyme inhibition), antioxidant scavenging of pathogen-generated reactive oxygen species, protein precipitation (tannin-protein complexes reducing digestibility), feeding deterrence through astringency and bitterness, and precursor availability for rapid induced defense responses. Phenolic content and composition vary dramatically among grapevine cultivars, tissues, and developmental stages, with red-fruited cultivars typically accumulating higher levels than white, and young tissues often showing higher concentrations than mature organs. The relationship between Constitutive_Phenolic_Disposition and resistance is generally positive but compound class-specific, as different phenolic subgroups show varying potency against particular pathogens and pests. Breeding for enhanced Constitutive_Phenolic_Disposition could improve broad-spectrum resistance while potentially affecting wine quality parameters that are strongly influenced by grape phenolic composition.
Constitutive_Phenolic_Disposition is_a Chemical_Defense_Disposition. Phenolics are the most abundant and diverse class of grapevine secondary metabolites, with established roles in defense. This class addresses basal phenolic content in healthy tissues, complementing induced stilbene synthesis (under Induced_Defense_Disposition) and specific phenolic subclasses addressed elsewhere. Phenolic content affects both disease resistance and wine quality, creating potential synergies or trade-offs requiring consideration in breeding applications.
Basaler-Phenolgehalt
Konstitutive-Phenol-Disposition
Präformierte-Phenylpropanoid-Abwehr
basal phenolic content
constitutive polyphenol defense
preformed phenylpropanoid defense
Lattanzio V et al. (2006) Role of phenolics in the resistance mechanisms of plants against fungal pathogens and insects, Phytochemistry: Advances in Research, Research Signpost Cheynier V et al. (2013) Plant phenolics: recent advances on their biosynthesis, genetics, and ecophysiology, Plant Physiology and Biochemistry 72:1-20 Dixon RA (2001) Natural products and plant disease resistance, Nature 411:843-847
Constitutive Phenolic Disposition
Syrah in Rhône Valley shows continued cell wall degradation during September-October Year N+1 at BBCH 85-89, with polygalacturonase activity sustained at 52-84 units through ripening representing maximum activity levels, pectin methylesterase 68-96 units continuing demethylation, pectate lyase possibly 18-32 units, sustained enzyme activities progressively degrading pectin with molecular weight declining from 165 to 68 kDa, pectin solubilization progressing from 48% to 72% soluble fraction, middle lamella extensively degraded with cells very loosely attached by maturity, cumulative degradation over 6-7 weeks creating substantially weakened wall structure, under warm Mediterranean conditions of 22-28°C temperature, typical of this mid-to-late season cultivar sustaining cell wall degradation through ripening
Continued cell wall degradation represents the sustained structural breakdown phase during progressive softening when pectin-degrading enzymes maintain elevated activities through ripening continuing pectin depolymerization and solubilization, polygalacturonase activity reaching maximum 40-100 units during ripening, pectin methylesterase activity 50-120 units, pectate lyase possibly contributing, sustained enzymatic modification progressively fragmenting cell wall and middle lamella pectin, and continued degradation as mechanism for progressive firmness decline through ripening creating soft ripe berry texture. This sustained breakdown phase occurs during late summer through autumn in late August-October in temperate regions Year N+1, representing cell wall modification continuation through ripening. Continued degradation mechanism involves sustained polygalacturonase gene expression and enzyme activity, continued pectin methylesterase activity creating enzyme substrate, coordinated enzyme activities maintaining pectin breakdown, progressive pectin depolymerization shortening chains, progressive solubilization converting structural to soluble pectin, cumulative degradation substantially weakening wall structure, and sustained enzymatic modification as softening mechanism. Polygalacturonase sustained activity shows PG activity increasing from initial veraison 15-40 units to ripening maximum 40-100 units, PG remaining highly active through ripening, enzyme activity sustained over weeks, PG as major continuing degradation enzyme, and high PG as ripening characteristic. PG activity pattern shows activity increasing during early ripening, reaching maximum mid-to-late ripening, possibly declining slightly late ripening in some cultivars, and activity pattern as temporal dynamics. PG gene expression shows PG genes remaining upregulated through ripening, continued transcription maintaining enzyme levels, expression as activity maintenance mechanism, and sustained expression through ripening. Pectin methylesterase sustained activity shows PME activity 50-120 units during ripening, PME continuing demethylation, sustained PME creating PG substrate continuously, and PME as continuing modification enzyme. PME activity pattern shows activity increasing during ripening, sustained at elevated levels, and pattern supporting continued degradation. Pectate lyase possible continued activity shows PL possibly contributing to continued degradation, PL activity variable among cultivars, and PL as additional continuing enzyme. Enzyme cooperation shows PG, PME, and possibly PL working synergistically throughout ripening, coordinated activities maximizing degradation, sustained cooperation as efficient mechanism, and enzyme synergy throughout. Progressive pectin depolymerization shows pectin chain length continuing decline, molecular weight declining from initial softening 100-200 kDa to ripening 50-150 kDa or lower, continued fragmentation creating smaller pieces, and depolymerization as progressive process. Pectin molecular weight trajectory shows progressive decline through ripening, eventual low molecular weight at maturity, and trajectory as degradation indicator. Chain length reduction shows degree of polymerization declining, shorter pectin chains from enzymatic cleavage, and reduction weakening structure. Progressive pectin solubilization shows soluble pectin continuing increase from initial 35-50% to ripening 60-80% total pectin, insoluble pectin declining to 20-40%, progressive conversion as continuing process, and solubilization throughout ripening. Solubilization kinetics shows progressive increase correlating with enzyme activities and firmness decline, rate depending on enzyme levels and substrate, and kinetics as degradation rate. Middle lamella progressive degradation shows middle lamella pectin extensively degraded during ripening, progressive loss of intercellular adhesive, substantial middle lamella loss by maturity, and middle lamella as primary continuing target. Cell-cell adhesion progressive loss shows intercellular bonding substantially reduced by ripening, cells very loosely attached in ripe berries, possible cell separation in very soft fruit, and adhesion loss as extensive degradation outcome. Cell wall pectin progressive loss shows wall pectin also extensively degraded, wall structure substantially weakened, walls very fragile in ripe berries, and wall degradation as softening mechanism. Cumulative degradation shows degradation accumulating over ripening weeks, cumulative effect creating substantial structural loss, total degradation much greater than initial, and cumulative as progressive feature. Structural integrity extensive loss shows cell wall and middle lamella structural networks extensively fragmented, integrity substantially lost creating very soft texture, and integrity loss as degradation outcome. Pectin domain degradation shows continued preferential attack on smooth homogalacturonan regions, progressive degradation of accessible domains, and domain-specific degradation. Calcium dynamics shows continued calcium release and redistribution as pectin degrades, calcium effects on residual pectin structure, and calcium as degradation factor. Other cell wall components shows hemicelluloses and cellulose remaining largely intact, selective pectin degradation continuing, and specificity as degradation characteristic. Degradation spatial pattern shows degradation throughout berry mesocarp, possible regional variation in extent, and spatial pattern affecting softening. Enzyme-substrate dynamics shows continuing enzyme-pectin interactions, substrate availability declining as degradation proceeds, and dynamics affecting rate. Environmental effects on continued degradation include temperature affecting enzyme activities with warm conditions promoting higher activities and faster degradation, cool conditions reducing enzyme kinetics, and temperature as major rate determinant. Temperature effects shows optimal enzyme activities at 20-30°C, temperature affecting cumulative degradation extent, and temperature as dominant factor. Cultivar variation shows enzyme activity levels varying among cultivars, some maintaining higher activities others lower, variation affecting degradation extent and softening, and genetic effects on degradation. Degradation regulation shows enzyme activities regulated at transcriptional and post-translational levels, developmental signals maintaining expression, and regulation as activity control. The process represents first aspect of progressive softening establishing sustained enzymatic modification, preceding increasing berry pliability and firmness decline. Duration continued degradation throughout 4-8 weeks ripening period. Regional timing shows continued cell wall degradation during late August-October through ripening. Viticultural significance includes representing sustained softening mechanism through ripening, determining progressive firmness decline and final texture, demonstrating enzyme activity regulation through ripening, and establishing cell wall degradation as ripening process. Research applications include analyzing continued enzyme activities and regulation through ripening, investigating progressive pectin degradation kinetics, evaluating enzyme-pectin interactions and degradation mechanisms, quantifying cumulative degradation extent, and assessing continued degradation as progressive softening mechanism.
Continued_Cell_Wall_Degradation is first aspect of Progressive_Softening, distinguished by sustained enzymatic modification. Represents 4-8 weeks ripening. Polygalacturonase 40-100 units maximum, pectin methylesterase 50-120 units. Sustained PG, PME activities throughout ripening. Progressive pectin depolymerization 100-200 → 50-150 kDa. Solubilization 35-50% → 60-80%. Middle lamella extensively degraded. Cumulative degradation over weeks. Temperature affects enzyme activities (optimal 20-30°C). Precedes Increasing_Berry_Pliability and Firmness_Decline. Primary progressive softening mechanism.
anhaltender Abbau
continuing wall breakdown
fortgesetzte Zellwand-Degradation
fortsetzender Wand-Abbau
progressive enzymatic modification
progressive enzymatische Modifikation
ripening degradation
sustained degradation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 482-488
Continued Cell Wall Degradation
In Chardonnay vines in Champagne, continued metabolic suppression maintains bud respiration rates at 0.3-0.5 μmol CO2 per gram dry weight per hour throughout the December through February endodormancy period, representing only 15% of rates measured in actively growing buds during June, with this suppression persisting even during late January warm periods when ambient temperatures of 10-12°C would support much higher metabolic rates in non-dormant tissues.
Continued metabolic suppression in Vitis vinifera represents the sustained reduction of overall metabolic activity in dormant tissues throughout the endodormancy period, maintaining the low rates of respiration, protein synthesis, and biosynthetic pathway activity that characterize the dormant state and conserve carbohydrate reserves for spring growth resumption. This metabolic state persists from dormancy establishment in late autumn until dormancy release in late winter, with metabolic rates remaining at levels far below those characteristic of active growth throughout the period regardless of temporary temperature fluctuations that might otherwise stimulate metabolism in non-dormant tissues. The respiratory component of continued metabolic suppression involves maintenance of low respiration rates that minimize carbohydrate consumption during the extended period without photosynthetic input, with dormant bud respiration rates typically only 10-20% of rates in actively growing buds at equivalent temperatures, reflecting both temperature-dependent rate reductions and active regulatory suppression of respiratory capacity. Protein metabolism during continued metabolic suppression is characterized by minimal protein turnover with very low rates of both synthesis and degradation, maintaining the dormancy-related protein complement including dehydrins and other protective proteins while avoiding the energetic costs of active protein synthesis that would deplete reserves. The biosynthetic pathway suppression encompasses reduced activity of pathways for amino acid, nucleotide, and lipid synthesis that would support growth, with metabolic flux directed toward maintenance functions rather than biosynthetic production of growth substrates. The regulatory basis of continued metabolic suppression involves both the direct temperature effects on enzyme reaction rates that reduce metabolism during cold exposure, and active regulatory mechanisms including ABA-mediated suppression of metabolic gene expression and the absence of growth hormone signals that normally stimulate metabolic activity in support of development. The persistence of metabolic suppression during temporary warm periods demonstrates that metabolic downregulation during endodormancy reflects regulatory programming rather than simply temperature limitation, with dormant tissues failing to increase metabolism substantially even when temperatures would permit higher activity in non-dormant tissues. The gradual relief of metabolic suppression occurs in parallel with dormancy release, as chilling accumulation reduces dormancy depth and allows progressive metabolic reactivation in preparation for spring growth.
Continued_Metabolic_Suppression represents the metabolic dimension of Dormancy_Maintenance addressing sustained low metabolic rates. Distinguished from Metabolic_Downregulation during dormancy induction which establishes the suppressed state, while Continued_Metabolic_Suppression addresses persistence of this state through endodormancy. Encompasses respiration, protein synthesis, and biosynthetic pathway suppression. Reflects both temperature effects and active regulatory suppression. Duration spans entire endodormancy period. Critical for reserve conservation during winter.
anhaltende metabolische Suppression
fortgesetzte Stoffwechselunterdrückung
kontinuierliche metabolische Reduktion
maintained metabolic dormancy
ongoing metabolic inhibition
persistent metabolic suppression
persistierende Stoffwechselhemmung
sustained metabolic reduction
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 5 Carbohydrate Supply and Reserve Storage Zapata C et al (2004) Partitioning and mobilization of starch and N reserves in grapevine. Journal of Experimental Botany 55: 2165-2175 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235
Continued Metabolic Suppression
In Sangiovese berries in Chianti Classico, Tuscany during BBCH 85-89, continuing acid decline proceeds consistently over 38 days from veraison to harvest, with titratable acidity decreasing from 16.8 to 5.9 g/L following a logarithmic decay curve with daily reductions averaging 0.29 g/L during weeks 1-3 and 0.12 g/L during weeks 4-5, correlating with 485 growing degree days accumulated above 10°C base temperature.
Continuing acid decline describes the sustained progressive decrease in berry organic acid concentration throughout the active ripening period in Vitis vinifera, characterized by persistent day-over-day reductions in titratable acidity of 0.1-0.3 g/L occurring consistently from early post-veraison through approach to harvest maturity over a period of 25-40 days. This process reflects the cumulative effect of ongoing malate catabolism through respiratory and enzymatic pathways, maintaining negative acid balance as consumption rates exceed any residual biosynthetic activity, resulting in monotonic decline curves when titratable acidity is plotted against thermal time accumulation during ripening. The biochemical basis involves continuous flux of vacuolar malate through cytosolic and mitochondrial metabolic pathways, with steady-state enzyme activities of NADP-malic enzyme, phosphoenolpyruvate carboxykinase, and mitochondrial malate dehydrogenase maintaining catabolism rates that collectively deplete 0.2-0.4 g/L malic acid equivalent per day throughout the active ripening window. Temperature integration determines the rate of continuing decline, with growing degree day accumulation correlating strongly with cumulative acid loss, enabling predictive modeling of harvest acidity based on thermal time calculations using base temperatures of 10°C and accumulated heat units of 300-500°C-days during the active reduction period. The process demonstrates asymptotic behavior as acid levels approach equilibrium values, with decline rates progressively decreasing from initial values of 0.4-0.5 g/L per day during early active reduction to 0.05-0.15 g/L per day as harvest approaches and substrate availability becomes limiting. Cultivar genetics influence the persistence of continuing decline, with varieties possessing high initial malic acid content such as Barbera and Sangiovese showing extended active decline periods of 35-45 days compared to low-acid varieties such as Viognier showing shorter periods of 20-28 days before reaching equilibrium. Diurnal patterns superimpose on the overall declining trend, with nighttime acid accumulation partially offsetting daytime respiratory losses, though net daily balance remains consistently negative throughout active ripening. Canopy management practices including leaf removal and shoot positioning influence continuing decline rates through modification of berry temperature exposure and resultant enzymatic activity levels. The consistency of continuing acid decline enables viticultural planning for harvest timing, with predictive equations incorporating initial acidity, temperature forecasts, and variety-specific decline coefficients achieving accuracy of plus-minus 0.5 g/L for harvest acidity prediction.
Continuing_Acid_Decline is subclass of Active_Acid_Reduction representing temporal persistence of acid loss. Distinguishes from discrete events by emphasizing sustained nature. Duration 25-40 days spanning active ripening. Sibling to Malate_Degradation and pH_Increase which represent mechanistic and chemical aspects respectively. Key characteristic is consistent day-over-day negative acid balance throughout active ripening period.
anhaltender Säure-Rückgang
continuous acid loss
fortlaufende Säure-Abnahme
kontinuierliche Aciditäts-Reduktion
ongoing titratable acidity reduction
persistent acidity decline
persistente Säure-Abnahme
sustained acid decrease
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 270-275 Coombe BG and McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6: 131-135
Continuing Acid Decline
In a Napa Cabernet Sauvignon vineyard, continuous sugar import was characterized over 48-hour intensive monitoring during late August Year N+1 (BBCH 85, mid-active accumulation) showing daytime peak 88 μmol hexose g⁻¹ h⁻¹ (12PM-4PM, berry temperature 31-34°C), evening 62 μmol g⁻¹ h⁻¹ (6PM-10PM, 26-28°C), nighttime sustained 42 μmol g⁻¹ h⁻¹ (12AM-4AM, 18-21°C, 48% of daytime rate), early morning 58 μmol g⁻¹ h⁻¹ (6AM-10AM, 22-26°C), demonstrating continuous 24-hour import without cessation, daily integrated 0.24% Brix per day.
Continuous_Sugar_Import represents the uninterrupted flux of hexoses from phloem unloading through cellular uptake into vacuolar storage throughout active ripening creating the sustained substrate flow that enables progressive sweetness development without diurnal cessation maintaining constant metabolic pressure for sugar accumulation. This continuous import occurs during active accumulation period (BBCH 84-86), typically days 15-38 post-veraison spanning 20-30 days, and is characterized by 24-hour hexose uptake without nocturnal cessation at rates 40-90 μmol combined glucose and fructose g⁻¹ h⁻¹ during daytime elevated to 50-110 μmol g⁻¹ h⁻¹ and reduced but maintained 25-60 μmol g⁻¹ h⁻¹ nighttime representing 40-70% of daytime flux, daily integrated import 0.20-0.32% Brix per day (1.4-2.2% per week) accumulating continuously rather than episodically, phloem delivery maintained throughout diel cycle providing substrate availability 24 hours though mass flow rates potentially declining 30-50% nighttime as transpiration reduces, and cumulative effect where continuous import over 25-35 day period produces 6-11° Brix gain representing major contribution to final sweetness without interruption that could limit accumulation efficiency. Physiological basis for continuity includes phloem sap sucrose concentration maintained 350-600 mM throughout diel cycle providing substrate pool, apoplastic sucrose turnover rapid with half-life 4-12 minutes ensuring continuous cleavage by cell wall invertase producing glucose and fructose substrate for uptake, hexose transporter proteins present continuously in plasma membrane with densities 8-14% of total membrane protein not subject to rapid degradation maintaining import capacity 24 hours, and tonoplast transport capacity similarly sustained enabling vacuolar sequestration preventing cytoplasmic hexose accumulation that could reduce plasma membrane uptake gradient. Diel pattern characteristics demonstrate daytime peak import 60-130 μmol hexose g⁻¹ h⁻¹ during hours 8AM-6PM when berry temperature optimal 24-32°C and phloem delivery maximum from active transpiration-driven mass flow, evening decline 45-85 μmol g⁻¹ h⁻¹ as temperature moderates and transpiration reduces but import continues, nighttime sustained flux 30-65 μmol g⁻¹ h⁻¹ despite reduced phloem delivery and cooler temperature 16-24°C slowing kinetics indicating metabolic capacity exceeds substrate availability during night, and early morning recovery 40-75 μmol g⁻¹ h⁻¹ as temperature increases and phloem delivery resumes demonstrating continuous 24-hour pattern without extended cessation periods. Molecular mechanisms enabling continuity include constitutive gene expression with VvHT transcript half-lives 8-14 hours requiring continuous transcription but steady-state levels maintained throughout diel cycle, protein stability with transporter half-lives 28-48 hours preventing rapid diurnal degradation maintaining functional capacity continuously, lack of strong diurnal transcriptional regulation unlike some genes showing circadian oscillation indicating ripening sugar transport prioritized over circadian control, and hormone signaling with ABA sustained 1600-2800 ng/g throughout 24-hour period maintaining transporter expression without nocturnal suppression. Substrate availability patterns show phloem sucrose delivery highest daytime 220-400 mg per berry per day declining nighttime to 120-240 mg per berry per day but remaining substantial supporting continued import, apoplastic hexose concentrations 180-350 mM daytime potentially declining to 120-250 mM nighttime but maintaining above threshold for transporter saturation (Km 1.5-10 mM depending on isoform), cytoplasmic hexose steady 60-100 mM throughout diel cycle through balance of plasma membrane import, metabolism, and tonoplast export maintaining homeostasis, and vacuolar accumulation progressive throughout 24 hours with concentrations rising 8-18 mM per day sustained day and night. Temperature effects on continuity demonstrate optimal import 26-30°C typically achieved midday enabling peak rates, cooler nighttime temperatures 16-24°C reducing rates 35-60% through kinetic effects on enzyme activities and membrane transport but not eliminating flux, and berry thermal mass plus nighttime atmospheric cooling creating diurnal temperature range 8-16°C modulating but not eliminating import maintaining continuity. Energetic considerations for continuous import include ATP requirements for membrane H⁺-pumps maintaining electrochemical gradients estimated 15-25 μmol ATP g⁻¹ h⁻¹ must be met continuously, respiratory ATP production from concurrent acid oxidation 30-55 μmol g⁻¹ h⁻¹ plus partial hexose respiration 15-30 μmol g⁻¹ h⁻¹ providing 45-85 μmol g⁻¹ h⁻¹ total adequate for transport and biosynthesis demands throughout diel cycle, and metabolic continuity where berry maintains active metabolism 24 hours supporting continuous import without nocturnal metabolic shutdown that would halt accumulation. Comparison with other fruits reveals grapes maintain continuous import unlike some climacteric fruits showing diurnal cessation, continuous pattern enabling efficient accumulation maximizing growing season utilization, and non-climacteric physiology with minimal ethylene involvement potentially contributing to sustained import without developmental interruption. Quality implications include continuous import enabling maximum total accumulation within available growing season achieving higher final sugars compared to hypothetical interrupted pattern, uniformity of import across 24-hour period reducing diurnal compositional variation potentially affecting berry heterogeneity, and efficiency where continuous process maximizes carbon use compared to episodic import requiring cyclic enzyme synthesis potentially wasting resources. Environmental factors affecting continuity include water status with adequate vine hydration essential for maintaining phloem delivery throughout diel cycle, severe deficit potentially disrupting continuity particularly nighttime when transpiration-driven flow minimal, temperature extremes with heat stress >35°C potentially disrupting daytime import or cool nights <12°C substantially reducing nighttime flux though rarely eliminating completely, and light operating primarily through source photosynthesis affecting substrate availability with adequate canopy function essential for sustained 24-hour sucrose supply. Viticultural management supporting continuity includes irrigation maintaining adequate vine water status throughout 24-hour period, canopy management ensuring sustained photosynthesis providing substrate, avoiding severe environmental stresses that could disrupt metabolic function, and recognizing continuous import as normal physiology not requiring intervention unless environmental conditions severely limiting.
Continuous_Sugar_Import is distinguished from intermittent or episodic import by showing 24-hour flux without nocturnal cessation, from peak import by representing sustained (40-90 μmol g⁻¹ h⁻¹ average) rather than maximum rates, and from initial import by showing established continuity rather than developing capacity. Nighttime flux 40-70% of daytime maintaining accumulation. Duration throughout active period BBCH 84-86. Essential for maximizing total accumulation. Continuous_Sugar_Import represents sustained metabolic process enabling efficient sweetness development.
24-Stunden-Zucker-Import
24-hour sugar import
anhaltender Substrat-Fluss
constant sweetening
konstante Süßung
sustained substrate flux
uninterrupted hexose uptake
ununterbrochene Hexose-Aufnahme
Keller M (2020) The Science of Grapevines 3rd Ed pages 223-225 continuous accumulation Dai ZW, Ollat N, Gomès E, Decroocq S, Tandonnet JP, Bordenave L, Pieri P, Hilbert G, Kappel C, van Leeuwen C, Vivin P, Delrot S (2011) Ecophysiological, genetic, and molecular causes of variation in grape berry weight and composition Plant Physiol 155:519-532 import dynamics Hayes MA, Davies C, Dry IB (2007) Isolation, functional characterization, and expression analysis of grapevine hexose transporters J Exp Bot 58:1985-1997 transporter function enabling continuity.
Continuous Sugar Import
The contrada name Contrada Rampante on the north slope of Mount Etna designates a specific volcanic terrace at approximately 750-850 meters elevation in the commune of Passopisciaro (Castiglione di Sicilia), planted to pre-phylloxera Nerello Mascalese vines on ungrafted rootstock in sandy volcanic soils derived from a specific historical lava flow — producers including Passopisciaro (Andrea Franchetti) and Girolamo Ferrara use the Contrada Rampante designation to communicate site specificity comparable to Burgundy's climat system, with the 20+ recognized contrade on Etna's north slope producing Nerello Mascalese wines of distinctly different character reflecting variations in lava flow composition, altitude, and exposure across distances of only a few hundred meters.
A vineyard designation within the Italian wine classification system that identifies a wine as originating from a named contrada — a traditional named district or locality within an Italian wine commune, used most prominently on Mount Etna in Sicily where contrada designations function as de facto single-vineyard indicators identifying specific volcanic slope parcels with distinctive terroir characteristics defined by elevation, lava flow geology, and microclimate exposure.
Contrada_Name documents the Italian tradition of sub-communal geographic naming that has gained particular prominence in the renaissance of Mount Etna viticulture since the early 2000s. While contrada designations exist throughout rural Italy as traditional administrative subdivisions of communes, their viticultural significance is most developed on Etna where the geological diversity created by successive volcanic eruptions over millennia produces extreme terroir variation within small areas: the Etna DOC disciplinare (amended 2011) formally recognizes contrada designations as Contrade dell'Etna, permitting their use on labels as geographic sub-units within the appellation. The broader Italian framework of Menzioni Geografiche Aggiuntive (MGA) and Unità Geografiche Aggiuntive (UGA) provides the regulatory structure for sub-appellation vineyard designation across Italian DOC and DOCG wines, with the Barolo MGA system (181 sites) and the Chianti Classico UGA system (11 zones) representing the most extensive implementations outside Etna.
Contrada-Bezeichnung
Contrada-Lagenbezeichnung
Etna contrada name
Italian district vineyard name
contrada designation
italienischer Bezirksname
Consorzio di Tutela dei Vini Etna DOC (2023) Contrade dell'Etna Italian Law 164/1992 on geographic sub-units Benjamin S (2019) Etna: Wine and People of the Volcano, Academos Press
Contrada Name
Cabernet Sauvignon vines in the Maipo Valley of Chile bearing a conventional production role receive 8-12 synthetic fungicide applications per season targeting Botrytis cinerea and downy mildew, utilize glyphosate-based herbicides for under-vine weed control, and may be treated with systemic insecticides such as imidacloprid at rates of 0.15-0.25 kg a.i./ha for mealybug management, achieving yields of 10-15 t/ha.
A production system role that inheres in a grapevine cultivated under standard agricultural management practices that permit the full range of legally authorized synthetic inputs including chemical fertilizers, synthetic fungicides, systemic insecticides, and herbicides as regulated by national plant protection legislation.
Conventional_Production_Role represents the globally predominant viticultural management approach accounting for approximately 90-93% of world vineyard area. The role permits all inputs authorized under national regulatory frameworks without the additional restrictions imposed by organic, biodynamic, or sustainable certification standards. This role does not preclude the adoption of integrated pest management (IPM) strategies: many conventional operations incorporate biological control agents alongside synthetic inputs.
Standardanbau-Rolle
conventional viticulture role
konventionelle Bewirtschaftungsrolle
konventionelle Produktionsrolle
non-organic production role
standard production role
OIV (2023) State of the World Vine and Wine Sector Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 521-548 Smart RE and Robinson M (1991) Sunlight into Wine, Winetitles pp: 1-12
Conventional Production Role
Cooling sensation assessment in Mosel Riesling served at 8°C reveals the style-appropriate thermal impression (cooling rating 8/10) that enhances the wine's racy acidity and delicate fruit, with the crisp coolness contributing to refreshment character that makes the wine perfect for summer afternoon consumption, while the same wine served at 14°C shows diminished cooling (4/10) and appears heavier and less refreshing despite identical composition, demonstrating how cooling sensation from serving temperature critically affects style expression and quality perception.
Cooling sensation quality is a temperature perception quality that represents the perception of coolness or cold character during wine consumption, arising from either actual low wine temperature or chemesthetic activation of cold-sensing receptors by specific compounds, characterizing the refreshing, crisp thermal impression particularly valued in wines emphasizing freshness and summer drinkability. Within the BFO framework, this quality specifically addresses the cool-cold dimension of thermal perception, distinguishing cooling sensation from warming while recognizing that cooling may derive from actual temperature (chilled wine) or chemical cooling effects (certain terpenes activating TRPM8 cold receptors). The ontological significance lies in cooling sensation contributing to the refreshment character essential for certain wine styles, where the crisp, cool impression enhances perception of freshness, acidity, and drinkability in wines intended for casual enjoyment and warm-weather consumption, while excessive coldness may mask flavor complexity and diminish appreciation of nuanced wines. The physiological basis for cooling perception involves TRPM8 ion channels in oral thermoreceptors that respond to both actual cold temperature and chemical agonists (menthol being the most potent), with wine cooling primarily deriving from serving temperature while certain terpenes (eucalyptol, menthol if present from herbal character) may contribute subtle chemical cooling enhancement. The compositional factors affecting cooling perception include actual wine temperature as served (primary determinant), certain monoterpenes potentially activating TRPM8 (contributing minor chemical cooling), high acidity (enhancing perception of freshness associated with coolness), and low alcohol (avoiding the warmth that would counteract cooling). The style associations of cooling sensation center on wines emphasizing freshness, including aromatic whites (Riesling, Vinho Verde) served well-chilled, rosé wines, sparkling wines, and light reds served cool, with the cooling impression contributing to the refreshing, thirst-quenching character that defines these styles' appeal. Understanding cooling sensation quality enables appropriate serving temperature selection, style-appropriate freshness assessment, and quality evaluation incorporating the cooling component of refreshing wine styles.
Cooling_Sensation_Quality addresses the cool-cold dimension of thermal perception, primarily arising from serving temperature with potential minor contribution from certain terpenes. This quality is essential for wine styles emphasizing freshness and refreshment. The class supports serving temperature guidance and style-appropriate assessment where cooling contributes to intended wine character.
Frische-Kälte
Kühlempfindung
chilled impression
cool sensation
crisp temperature
erfrischende Kühle
knackige Temperatur
refreshing coldness
McKemy DD et al. (2002) Identification of a cold receptor reveals a general role for TRP channels in thermosensation. Nature 416(6876): 52-58 Ross CF, Weller KM (2008) Effect of serving temperature on the sensory attributes of red and white wines. Journal of Sensory Studies 23(3): 398-416
Cooling Sensation Quality
Copigmentation contributes 30-50% of young red wine color through hyperchromic (increased absorbance) and bathochromic (red-shifted wavelength) effects on anthocyanin chromophores. Effective copigmentation requires molar ratios of copigment to anthocyanin of 5-20:1, with flavonols (quercetin, myricetin) and hydroxycinnamic acids (caffeic, p-coumaric) serving as primary cofactors. Cultivars with high flavonol accumulation (Pinot Noir 40-100 mg/kg, Cabernet Sauvignon 30-80 mg/kg) show enhanced copigmentation compared to low-flavonol types (Grenache 15-40 mg/kg). Self-association of anthocyanins (intramolecular copigmentation in acylated forms, intermolecular stacking at high concentration) provides additional stabilization. Copigmentation effects are maximal in young wines and decline as anthocyanins are incorporated into polymeric pigments, with the copigmentation contribution decreasing from 40-50% at bottling to 10-20% after 2 years.
A color stability disposition that is the inherited capacity of a Vitis vinifera plant to produce grapes with sufficient anthocyanin and copigment (flavonol, hydroxycinnamic acid, tannin) concentrations and appropriate molecular ratios to support copigmentation reactions that enhance and stabilize red wine color through non-covalent molecular stacking between anthocyanins and colorless phenolic cofactors.
Copigmentation represents the primary mechanism of color enhancement in young red wines, explaining why wine color intensity can exceed that predicted from anthocyanin concentration alone. The non-covalent nature of copigmentation complexes means they are in dynamic equilibrium, with complex stability affected by pH, temperature, and ethanol concentration. Copigmentation provides a "bridge" to polymeric pigment formation by maintaining anthocyanin in favorable conformations for subsequent covalent reactions. Viticultural practices affecting flavonol accumulation (cluster exposure) directly impact copigmentation capacity.
Copigment-Kapazität
Copigmentierung
Farbverstärkung
color enhancement trait
copigment capacity
molecular stacking disposition
Boulton R (2001) The copigmentation of anthocyanins and its role in the color of red wine: a critical review. American Journal of Enology and Viticulture 52:67-87 Hermosín Gutiérrez I et al. (2005) Influence of variety and aging on colour and anthocyanin composition of wines from Castilla-La Mancha. Food Chemistry 91:429-438
Copigmentation Disposition
Copper content quality management in organic Bordeaux production requires balancing Bordeaux mixture applications for downy mildew control against wine copper limits, with harvest timing targeted to allow 2-3 weeks after final copper spray for rainfall dilution and fruit growth to reduce surface residues must copper of 3-8 mg/L from well-managed organic vineyards requires copper removal during fermentation (yeast cell adsorption removes 30-50%) and potentially copper sulfate fining to achieve wine levels below the 1 mg/L EU limit, demonstrating the processing requirements imposed by organic disease management.
Copper_Content_Quality represents the chemical quality characterizing the concentration of copper (Cu) in grape berry tissue, capturing the essential micronutrient central to electron transport, oxidase enzymes, and lignin synthesis, while also addressing the quality implications of copper residues from vineyard fungicide applications. Copper in grape juice varies enormously depending on fungicide history, ranging from 0.1-1 mg/L in organically unfungicided fruit to 2-20+ mg/L where copper-based sprays (Bordeaux mixture, copper hydroxide, copper oxychloride) have been applied close to harvest. The essential functions of copper include electron transport in plastocyanin and cytochrome oxidase, catalytic activity in polyphenol oxidase and laccase, and cell wall lignification through laccase-mediated phenolic polymerization. Copper deficiency is relatively rare in viticulture due to the widespread use of copper fungicides, though it can occur in sandy soils with low organic matter. The primary copper quality concern relates to excessive residues from fungicide applications that affect wine production. Elevated must copper (above 5 mg/L) increases oxidation risk through copper-catalyzed Fenton reactions, can cause copper casse (copper sulfide haze) in reductive wines, and may inhibit fermentation at very high levels (above 20 mg/L) through yeast toxicity. Copper removal during winemaking occurs through precipitation with hydrogen sulfide (forming insoluble copper sulfide), adsorption to yeast cell walls, and fining treatments. Regulatory limits for copper in wine (typically 1 mg/L or less) require management when must copper is elevated. The balance between copper's essential nutritional role and its potential as a contaminant creates unique management challenges, particularly in organic viticulture where copper-based fungicides are primary disease control tools.
Copper_Content_Quality addresses a micronutrient with dual significance as essential element and potential contaminant. The widespread use of copper fungicides makes environmental contamination a greater concern than nutritional deficiency in most viticultural situations. Copper's role in wine oxidation parallels iron, with both metals participating in Fenton-type reactions that generate reactive oxygen species. The copper sulfide precipitation pathway provides natural copper removal in reductive winemaking but can cause problems if hydrogen sulfide is released from excess copper during storage. Regulatory limits on wine copper require monitoring and management when vineyard copper applications are heavy.
Beerenkupferqualität
Cu content
Cu-Gehalt
Kupfergehalt
Kupferkonzentration
berry copper quality
copper concentration
copper level
Danilewicz JC (2003) Review of Reaction Mechanisms of Oxygen and Proposed Intermediate Reduction Products in Wine: Central Role of Iron and Copper. American Journal of Enology and Viticulture 54(2): 73-85 Provenzano MR et al. (2010) Copper Contents in Grapes and Wines from a Mediterranean Organic Vineyard. Food Chemistry 122(4): 1338-1343
Copper Content Quality
A horizontal cordon on Vitis vinifera cv. Merlot in a bilateral cordon spur-pruned system, extending 88cm from trunk along trellis wire at 85cm height, 9 years old measuring 26mm diameter, bearing 14 two-bud spurs spaced approximately 12-13cm apart along upper surface, shoots from spurs trained vertically upward into 120cm of trellis space, supporting estimated 26-28 fruiting shoots bearing 42-48 clusters totaling 11-13kg crop, representing mature well-established cordon providing stable platform for annual spur renewal and consistent crop production with minimal structural modification required during annual dormant pruning operations.
A permanent arm trained and maintained in a specific linear configuration, typically horizontal or vertical, serving as the primary structural framework for spur-pruned training systems, characterized by multi-year woody structure (typically 3-20+ years old), consistent orientation along trellis wires or posts, regular distribution of fruiting spurs at intervals along its length, and function as permanent platform connecting trunk to annual fruiting positions while defining canopy architecture and mechanization compatibility. Cordons represent the defining structural element of cordon-trained systems (as opposed to cane-trained systems), with their orientation (horizontal, vertical, angled), height, length, and spur distribution fundamentally determining training system character, labor requirements, fruit exposure, and suitability for mechanical operations including pruning, hedging, and harvesting.
Cordons are distinguished from other permanent arms by their specific trained configuration and spur-pruning association, though terminology varies regionally with "cordon" and "arm" sometimes used interchangeably. The cordon concept is central to major training systems worldwide including bilateral cordon (most common in New World commercial viticulture), vertical cordon (common in some European regions), and various specialized configurations. Cordon establishment requires 2-4 years of deliberate training achieving desired architecture, with mature cordons then providing stable framework for decades of production, though eventual renewal becomes necessary as age-related decline, disease accumulation, or positioning problems reduce functionality, typically after 10-25 years depending on variety, region, and management quality.
Schenkelarm
Tragschenkel
permanent cordon
spur-bearing arm
structural cordon
trained arm
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Reynolds & Vanden Heuvel (2009) "Influence of grapevine training systems on vine growth and fruit composition= a review" American Journal of Enology and Viticulture 60=251-268 Intrieri et al. (2011) "Grapevine mechanization= Constraints and perspectives" Acta Horticulturae 978=471-478.
Cordon
A cordon arm water shoot on a bilateral horizontal cordon of Vitis vinifera cv. Pinot Noir in Oregon, USA, emerging from a latent bud at 55 cm from the trunk along the 90 cm east-facing cordon arm, developing in response to complete spur death in a 20 cm cordon section caused by Eutypa dieback, growing 145 cm vertically by August with 16 nodes positioned at the mechanized harvest head height of 120 cm where it interferes with harvester finger operation, leaves averaging 165 cm² area showing typical Pinot Noir morphology, requiring decision between immediate removal to preserve harvest efficiency or retention and training as a replacement spur to restore fruiting position density after two seasons of managed development.
An arm water shoot specifically originating from latent or adventitious buds on horizontal, vertical, or angled cordon structures in Vitis vinifera, characterized by emergence from the permanent cordon arm that extends from the trunk to support multiple fruiting spurs or canes, typically developing in response to severe spur damage, progressive trunk disease extension into cordon tissues, or management practices that create imbalanced vegetative-reproductive ratios along the cordon length. Cordon arm water shoots represent particular challenges in mechanized vineyard systems where cordons are positioned at precise heights and angles for machine harvest compatibility, with vigorous vertical shoots from horizontal cordons interfering with harvest head function, damaging harvester mechanisms, and creating fruit loss through cluster stripping or vine damage. These shoots commonly develop following severe winter freeze events that kill established spurs, creating extended bare cordon sections that accumulate carbohydrates and trigger latent bud activation, or in response to Eutypa dieback, Botryosphaeria dieback, or Esca symptoms that compromise vascular function in specific cordon sections. Management strategies for cordon arm water shoots include assessment of underlying cordon health through inspection for cankers, discoloration, or dieback symptoms, decisions about shoot retention for spur replacement in locally damaged zones, and removal timing that balances resource competition concerns against wound healing optimization.
Cordon arm water shoots are distinguished from other arm water shoots by specific origin on cordon structures, which in modern viticulture represent the most common permanent arm type in mechanized production systems. The horizontal orientation of most cordons creates specific management challenges as water shoots grow vertically and quickly extend above or below target fruiting zones. Timing of cordon water shoot removal affects wound healing, with early-season removal (May-June) allowing better wound closure but removing potential photosynthetic capacity, while late-season removal (August-September) preserves seasonal photosynthesis but creates larger wounds entering dormancy. Cordon water shoots in the same positions for multiple consecutive years indicate progressive trunk disease requiring diagnostic investigation.
Ersatzsporntrieb
Kordonwasserschoss
Spalierwassertrieb
horizontal cordon water shoot
spur-replacement shoot
vertical cordon water shoot
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles pp. 89-107 Sosnowski et al. (2007) "The influence of climate on foliar symptoms of Eutypa dieback in grapevines" Phytopathology 97=1284-1289 Úrbez-Torres & Gubler (2009) "Pathogenicity of Botryosphaeriaceae species isolated from grapevine cankers in California" Plant Disease 93=584-592 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 439-464.
Cordon Arm Water Shoot
Cordon suitability quality comparison in Barossa Valley evaluated long-term performance of Shiraz and Grenache under bilateral cordon spur-pruned training representing the regional standard. Shiraz demonstrated excellent cordon suitability quality with strong basal bud fruitfulness averaging 1.8 clusters per shoot from spur positions, vigorous but manageable shoot development, and maintained productivity over 40+ year cordon age with appropriate trunk disease management, supporting the cultivar's dominance in cordon-based regional production. Grenache showed good cordon suitability quality with adequate basal fruitfulness of 1.4 clusters per shoot and long cordon lifespan, though with inherent tendency toward large, loose clusters requiring careful crop load management to optimize fruit concentration. Mourvèdre exhibited marginal cordon suitability quality with inconsistent basal bud fruitfulness (CV 38% versus 22% for Shiraz), tendency toward blind basal buds following cold winters, and elevated susceptibility to Eutypa dieback causing progressive cordon deterioration, motivating some producers to employ cane-pruned Guyot training for this cultivar despite the increased labor requirements.
Cordon suitability quality is a trellis adaptation quality that inheres in a grapevine cultivar or clone and represents the degree of compatibility with cordon-trained spur-pruned systems characterized by permanent horizontal arms (cordons) established along fruiting wires from which short spurs bearing two to three buds are retained annually, with shoots developing upward into vertically positioned canopy structures, representing a widely adopted training approach that provides labor efficiency advantages, mechanization compatibility, and consistent vine structure while requiring cultivar characteristics suited to sustained productivity from fixed spur positions over the multi-decade lifespan of established cordons. This quality determines whether a cultivar possesses the basal bud fruitfulness, spur renewal capacity, and structural longevity characteristics necessary to achieve stable, productive performance under cordon training without the annual fruiting wood selection opportunity provided by cane-pruned systems. The ontological significance of cordon suitability quality lies in its role enabling efficient, mechanized viticulture through the permanence and uniformity of cordon structures that facilitate standardized management operations, mechanical pruning, and predictable production patterns across large vineyard areas, while requiring cultivar adaptation to the specific constraints of fixed spur positioning and multi-year cordon maintenance. The biological requirements for high cordon suitability quality include strong basal bud fruitfulness ensuring productive clusters from the proximal bud positions retained on short spurs, capacity for sustained spur renewal from the permanent cordon without progressive weakening or dead arm development, appropriate vigor expression creating balanced shoot development from the spur positions, structural cordon characteristics enabling stable horizontal growth along the training wire, and physiological resilience maintaining productivity despite the accumulation of trunk disease organisms in aging permanent wood. Cultivars with poor basal bud fruitfulness produce inadequate yields under spur pruning despite potentially high mid-cane fruitfulness suited to Guyot training, while those susceptible to trunk diseases may exhibit progressive cordon decline requiring costly remediation or replanting. Assessment of cordon suitability quality involves evaluation of basal bud fruitfulness patterns, long-term spur renewal sustainability, cordon structural stability, trunk disease susceptibility, and multi-year yield stability under spur-pruned management.
Cordon_Suitability_Quality is_a Trellis_Adaptation_Quality, representing compatibility with permanent cordon spur-pruned training systems. This class addresses the specific requirements of cordon training including basal bud fruitfulness, spur renewal capacity, and long-term cordon viability. Distinguished from Guyot_Suitability_Quality which addresses cane-pruned systems with annual fruiting wood renewal. The labor efficiency and mechanization compatibility of cordon systems favors their adoption where cultivar suitability permits. Trunk disease susceptibility represents key long-term consideration for cordon suitability assessment.
Kordon-Eignung
Kordonerziehungs-Kompatibilität
Spalierkordon-Eignung
Zapfenschnitt-Tauglichkeit
bilateral cordon compatibility
cordon training adaptation
permanent arm training fitness
spur pruning suitability
Winkler AJ et al. (1974) General Viticulture. University of California Press, Berkeley, pp. 287-324 Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 32-43 Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review. American Journal of Enology and Viticulture 60:251-268
Cordon Suitability Quality
A Merlot vine trained to bilateral cordon at 90 cm height bears a Cordon_Trained_Vine_Role, with two horizontal cordons extending 60 cm each direction from the trunk, each bearing 5-6 spur positions that produce the annual crop in a predictable fruit zone suitable for mechanical management. A Pinot Noir vine on low unilateral cordon in Oregon's Willamette Valley serves in a Cordon_Trained_Vine_Role, the single cordon at 75 cm height positioning fruit for optimal ripening and hand harvest access in the cool climate. A Sangiovese vine on high bilateral cordon in central Italian pergola-influenced systems bears a Cordon_Trained_Vine_Role, the elevated cordons at 180 cm enabling traditional agricultural practices beneath the vine canopy.
A training system role that inheres in a grapevine trained with one or more permanent horizontal extensions (cordons) from the trunk, typically supported by trellis wires, which bear the spur positions or cane replacement points from which annual fruiting shoots emerge. Cordon training establishes a permanent framework enabling consistent fruit zone positioning, predictable canopy architecture, and suitability for mechanical operations (pruning, shoot positioning, leaf removal, harvesting) that require uniform vine geometry. Cordons are typically established at 60-120 cm height depending on climate, mechanization requirements, and production objectives, with lower cordons providing frost protection and easier hand labor access while higher cordons facilitate mechanical harvesting and reduce back strain. Cordon development requires 2-3 years during vineyard establishment, with the horizontal structure grown along training wires and spur positions established at regular intervals (typically 10-20 cm) along the cordon length. Cordon longevity varies with management quality well-maintained cordons may remain productive for 30-50+ years, while poor pruning practices, disease, or mechanical damage may necessitate cordon renewal or replanting.
Cordon_Trained_Vine_Role is subclassed by cordon configuration—bilateral (two cordons extending opposite directions) versus unilateral (single cordon), and by cordon height—high versus low positioning. Cordon training represents the foundation of most modern commercial viticulture, enabling the mechanization and labor efficiency essential for economically viable production at scale. The permanent cordon structure creates management consistency but also potential problems—trunk diseases (Esca, Eutypa) can colonize the extensive permanent wood, while poorly positioned spurs may become unproductive over time. Cordon training interacts with pruning system most cordon-trained vines are spur-pruned (retaining short spurs on the cordon), though some systems combine cordons with cane replacement. Training system selection increasingly considers climate change adaptation, with cordon heights and orientations adjusted to manage fruit exposure and ripening timing under warming conditions.
Dauerkordonsystem
Horizontalkordon
Kordon-Erziehung
Kordonrebe
cordon training
cordon-trained vine
horizontal cordon
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems, American Journal of Enology and Viticulture 60: 251-268 Winkler AJ et al. (1974) General Viticulture, University of California Press
Cordon Trained Vine Role
Syrah cork cambium at 10 years post-planting shows active phellogen at 1.2-1.8 mm depth in secondary phloem producing phellem (cork) outward and phelloderm inward, cork cambium initial cells 25-35 μm dimensions dividing periclinally with 85% divisions producing suberized cork cells (accumulating 0.8-1.4 mm cork layer annually) and 15% producing living phelloderm cells (0.1-0.3 mm annual increment), complete periderm cylinder with scattered lenticels 2-3 mm diameter enabling gas exchange, bark exterior showing 2-3 successive periderm layers from current and previous cork cambium activity creating 3-5 mm total bark thickness
Cork cambium (phellogen) is the unidirectional lateral meristem in Vitis vinifera trunk producing cork cells (phellem) centrifugally toward trunk surface and phelloderm (secondary cortex) centripetally, forming protective periderm that replaces epidermis as trunk expands, representing the meristematic tissue responsible for bark formation and protection. Cork cambium origin and development includes= initial formation in sub-epidermal cortex (typically during year 1-2 as trunk diameter exceeds ~8-15 mm and epidermis ruptures), dedifferentiation of cortex parenchyma cells into meristematic cork cambium initials (transition from differentiated to dividing state), formation of complete or partial cambial cylinder (early cork cambium often forming lenticular patches, later forming continuous cylinders), successive cork cambium formation (first periderm in cortex replaced by deeper periderms developing in secondary phloem as trunk ages), and multiple active cork cambia potentially present (outer older periderms gradually sloughing while inner newer periderms form continuously renewing bark protection). Cork cambium cellular organization includes= rectangular to polygonal initial cells (relatively uniform cell dimensions 20-40 μm length and width), single cell layer of dividing initials (compared to 2-6 cell layer vascular cambium zone), cells with thin primary walls (enabling division and daughter cell differentiation), dense cytoplasm and prominent nuclei (reflecting meristematic status), and relatively simple organization (compared to complex vascular cambium with fusiform and ray initials, cork cambium comprises one cell type producing relatively uniform derivatives). Cork cambium position in trunk includes= initial periderm forming at 0.5-1.5 mm depth (in sub-epidermal cortex of young trunks), subsequent periderms forming progressively deeper (in secondary phloem at 1-4 mm depth as trunk ages), multiple periderm layers in old trunk bark (successive periderms at various depths producing layered bark structure), outermost cork cambium typically 0.5-2 mm from bark surface (producing actively accumulating phellem), and inner older cork cambia no longer active (embedded in outer bark having produced cork before ceasing activity and being replaced by deeper periderms). Cork cambium division pattern includes= periclinal divisions (parallel to surface producing cork cells outward and phelloderm cells inward), asymmetric production strongly favoring phellem (80-90% of divisions producing cork cells, 10-20% producing phelloderm), continuous activity during growing season (April-September producing new cork and phelloderm cells), intermittent division (less regularly active than vascular cambium with periodic bursts of activity), and limited anticlinal divisions (circumference increase accommodated primarily by cell expansion and some divisions). Cork cambium products include= phellem (cork cells) produced centrifugally (dead cells at maturity with suberized walls forming impermeable protective layer), typical cork accumulation 0.5-3 mm thickness per year depending on vine age and vigor, phelloderm produced centripetally (living parenchyma cells forming inner periderm layer), phelloderm typically thinner than phellem (0.1-0.5 mm annual increment), and complete periderm complex (phellem plus cork cambium plus phelloderm forming functional protective unit). Cork cambium phellem characteristics include= suberized cell walls (suberin deposition rendering walls impermeable to water and gases), dead cells at maturity (protoplasm degenerating leaving hollow suberized cell walls), rectangular brick-like cells (arranged in radial files reflecting cambial origin), air-filled lumens (providing insulation and flexibility), and protective functions (preventing water loss, excluding pathogens, providing thermal insulation, protecting cambium and phloem from mechanical damage). Cork cambium seasonal activity includes= reactivation in spring (April-May typically later than vascular cambium reactivation), active division during growing season (May-September producing new cork layers), gradual cessation in fall (September-October coinciding with overall growth slowdown), winter dormancy (November-February with minimal activity), and less synchronized than vascular cambium (individual cork cambium patches showing variable activity timing). Cork cambium successive formation includes= first periderm lifespan (initial cortical periderm functioning 2-5 years before being replaced), second periderm formation (new cork cambium arising in outer secondary phloem as first periderm becomes non-functional), continued periderm succession (new periderms forming every 3-8 years in progressively deeper phloem), ridged bark development (successive periderms creating layered structure, outer dead layers splitting and flaking as inner layers form), and continuous protection maintenance (successive periderm formation ensuring uninterrupted protective covering despite bark expansion and outer layer sloughing). Cork cambium variation by trunk age includes= young trunks (years 1-3) showing initial periderm formation in cortex (often incomplete cylinders or lenticular patches), maturing trunks (years 4-10) showing 1-2 successive periderms in outer phloem (developing more complete cylinders), mature trunks (years 10-25) showing 2-4 successive periderm layers (creating characteristic bark texture), and old trunks (years 25-50+) showing multiple overlapping periderms (producing thick furrowed bark 4-12 mm depth with complex layered structure). Cork cambium lenticel formation includes= specialized regions where cork cambium produces loosely arranged cells (creating porous structures in otherwise impermeable bark), lenticels enabling gas exchange (allowing oxygen influx and carbon dioxide efflux maintaining respiration in living phloem and cambium), lenticel density 2-8 per cm² bark surface, lenticel dimensions typically 1-4 mm diameter creating visible spots or lines on bark surface, and seasonal lenticel dynamics (expanding during growing season, closing somewhat during dormancy). Cork cambium significance in trunk protection includes= producing waterproof cork (preventing desiccation of underlying living tissues), creating pathogen barrier (suberized cork resisting fungal and bacterial penetration), providing mechanical protection (flexible cork cushioning impacts and abrasion), insulating cambium and phloem (cork layers buffering temperature extremes protecting meristematic tissues), and enabling bark expansion (successive periderm formation accommodating trunk diameter increase while maintaining protection).
Cork_Cambium emphasizes bark-forming meristem and is distinguished from Vascular_Cambium (wood and inner bark producer) and Trunk_Cambium (both cambial types) by specific function producing protective periderm. Critical for understanding bark formation, trunk protection mechanisms, successive periderm development, bark texture evolution, and maintenance of protective covering as trunk diameter increases.
Bark cambium
Cork-forming meristem
Korkkambium
Peridermkambium
Phellogen
Cork Cambium
Cabernet Sauvignon cork layer periderm at 17 years post-planting shows phellem 1.8-3.2 mm thick (representing 75-82% of total 2.4-3.9 mm periderm thickness) comprising 28-42 radial files of dead rectangular suberized cork cells 28-48 μm dimensions, cells with suberin-wax wall lamellae 0.2-0.4 μm thick providing waterproof barrier (<0.05% water permeability), air-filled lumens (contributing 45-58% void space), cork density 0.15-0.22 g/cm³, thermal conductivity 0.042-0.048 W/m·K providing insulation buffering 22°C external temperature swing to 9°C at phellogen depth, scattered lenticels 2-3 mm diameter interrupting otherwise impermeable cork, cork layer showing light tan fresh color weathering to grey-brown at surface
Cork layer periderm is periderm emphasizing the phellem (cork cells) component, representing the outermost dead suberized cell layers that provide the primary waterproof and insulating barrier, typically comprising 70-85% of total periderm thickness and determining protective capacity and bark surface characteristics. Cork layer anatomical characteristics includes= dead rectangular cells (20-50 μm dimensions arranged in radial files, cells lacking protoplasm at maturity), suberized cell walls (suberin polymer comprising aliphatic and aromatic domains creating waxy hydrophobic barrier impermeable to water and most gases), secondary wall with suberin lamellae (alternating suberin and wax layers 0.1-0.5 μm thick creating multilayered impermeable structure), thin primary wall (cellulose-pectin primary wall often compressed or degraded in mature cork cells), air-filled cell lumens (hollow cells providing lightweight structure, ~40-60% of cork volume is air contributing to insulation and compressibility), and radial orientation (cells stacked in files perpendicular to trunk surface reflecting cork cambium division pattern). Cork layer thickness variation includes= thin cork layers (0.3-0.8 mm in weak periderms or young trunk first periderm), moderate cork layers (0.8-2.0 mm typical in vigorous periderms on mature trunks), thick cork layers (2.0-4.0 mm in highly active periderms on old vigorous trunks or specific cork-producing cultivars), annual accumulation (0.4-3.0 mm radial increment per growing season depending on cork cambium activity and vine vigor), and cumulative depth (individual periderm cork layer representing 1-8 years accumulation before cork cambium becomes inactive and replaced by deeper formation). Cork layer formation process includes= cork cambium producing cork cell initials (periclinal divisions toward trunk surface), cork cells expanding radially (initial cells enlarging 2-4 fold achieving mature dimensions), secondary wall deposition (suberin synthesis and deposition creating lamellae, wax accumulation in wall), programmed cell death (protoplasm degenerating leaving hollow suberized cell wall), and continued outward accumulation (new cork cells added internally pushing older cells outward creating progressively thicker cork layer). Cork layer cellular organization includes= orderly radial files (reflecting organized cork cambium divisions, cells perfectly aligned radially), tangential rows (slight organization tangentially from cork cambium initial arrangement), lack of intercellular spaces (cells tightly packed maximizing barrier integrity), occasional tyloses or inclusions (some cork cells containing remains of protoplasm or mineral deposits), and growth rings (sometimes visible boundaries between growing seasons appearing as subtle lines in thick cork layers reflecting annual production variations). Cork layer physical properties includes= low density (0.12-0.25 g/cm³ for pure cork tissue, lower than most plant tissues due to air-filled structure), compressibility (cork cells collapsing under pressure then recovering providing cushioning function), elasticity (cork showing some elastic recovery after compression though with permanent deformation beyond certain threshold), low thermal conductivity (0.04-0.05 W/m·K, air-filled cells providing excellent insulation), low water permeability (suberized walls preventing water movement, <0.1% water absorption in intact cork), and resistance to microbial decay (suberin resisting enzymatic degradation, cork persisting for years without decomposition). Cork layer chemical composition includes= suberin (40-50% dry weight, complex polyester polymer with aliphatic C16-C24 fatty acids and aromatic phenolic compounds), lignin (15-25%, providing rigidity and chemical resistance), polysaccharides (10-20%, primarily cellulose and hemicelluloses in primary wall), extractives (10-20%, waxes, tannins, and other compounds), and ash (1-3%, mineral content primarily calcium, potassium, magnesium). Cork layer protective functions includes= primary waterproofing (preventing water loss from living tissues beneath, critical during drought and winter dormancy), pathogen exclusion (impermeable barrier preventing fungal spores and bacteria from reaching susceptible living phloem and cambium), mechanical protection (compressible cork absorbing impacts from equipment, hail, or other mechanical stress), thermal insulation (buffering daily and seasonal temperature fluctuations protecting sensitive cambium from heat and cold extremes), UV radiation barrier (cork absorbing damaging ultraviolet radiation preventing photodamage to living tissues), and moderate fire resistance (cork layers providing brief protection against flames or radiant heat). Cork layer in bark texture includes= smooth bark (thin uniform cork layers producing relatively even surface), furrowed bark (thick cork layers cracking longitudinally under expansion creating ridges and valleys), platy bark (cork layers separating in irregular plates), exfoliating bark (cork layers peeling in strips or sheets), cork layer thickness determining texture depth (thin cork creating shallow texture, thick cork creating pronounced 3-10 mm deep furrows or prominent plates), and weathering effects (exposed cork surface eroding to grey-brown from tan-brown fresh color). Cork layer lenticel interruptions includes= porous regions in otherwise impermeable cork (lenticels comprising loosely arranged cork cells with large intercellular spaces), lenticel formation by cork cambium (phellogen producing loosely organized cells in specific regions), gas exchange function (lenticels enabling oxygen influx and carbon dioxide efflux maintaining respiration in living inner bark), lenticel appearance (visible as spots or short lines 1-4 mm diameter on cork surface often lighter colored than surrounding cork), and lenticel density (2-8 per cm² cork surface providing adequate but not excessive gas exchange maintaining mostly impermeable barrier). Cork layer succession in outer bark includes= innermost functional cork layer (most recent periderm actively accumulating cork), middle cork layers (previous periderms no longer producing but maintaining protective function, embedded 2-6 mm depth), outermost cork layers (oldest periderms weathered and often sloughing, providing exterior surface), and crushed phloem between cork layers (thin bands of collapsed non-functional phloem separating successive periderm cork layers contributing to outer bark structure). Cork layer degradation and loss includes= surface weathering (UV radiation, freeze-thaw, moisture changes gradually degrading outermost cork), checking and cracking (brittle cork developing fissures particularly in dry conditions), fungal surface colonization (some fungi degrading outermost weathered cork though unable to penetrate intact suberized layers), mechanical erosion (wind, rain, physical abrasion removing exterior cork surfaces), and progressive exterior loss (oldest outermost cork layers eventually falling maintaining dynamic equilibrium as inner cork layers form). Cork layer cultivar characteristics includes= thick cork cultivars (producing 1.5-3.5 mm annual cork increment creating substantial protective layers), thin cork cultivars (producing 0.4-1.2 mm increment with limited cork accumulation), uniform cork (even production creating smooth consistent cork layers), irregular cork (variable production creating textured non-uniform layers), and cork color (cultivar-specific fresh cork hues from light tan to reddish-brown before weathering to grey-brown).
Cork_Layer_Periderm emphasizes phellem component and is distinguished from Periderm (complete three-layer complex), Protective_Periderm (function emphasis), and Cork_Cambium (meristem producing cork) by focus on dead suberized cell layers. Critical for understanding primary waterproofing and insulation mechanism, cork physical and chemical properties, bark texture development, protective barrier integrity, and cork layer as dominant protective component determining periderm effectiveness in trunk protection.
Cork cell zone
Korkzellschicht
Phellem layer
Phellemschicht
Suberinisierte Barriere
Suberized barrier
Evert, R. F. (2006). Esau's Plant Anatomy= Meristems, Cells, and Tissues of the Plant Body (3rd ed.). Wiley-Interscience. | Pereira, H. (2007). Cork= Biology, Production and Uses. Elsevier.
Cork Layer Periderm
Cabernet Sauvignon grafted on SO4 rootstock at 16 years showing cork layer rootstock periderm in outermost active periderm, cork layer 1.4 mm thick comprising radially aligned rectangular dead cells with suberized walls, microscopic examination confirming hollow air-filled lumens and absence of protoplasts indicating complete cellular death, suberin staining (Sudan IV) showing positive reaction confirming suberization, water impermeability testing demonstrating effective waterproofing, cork layer providing essential protective barrier, multiple deeper cork layers visible at 3.1 and 5.8 mm depths in older periderms embedded in inner bark
Cork layer rootstock periderm is the phellem (cork) component of rootstock periderm emphasizing the outermost dead protective layer produced by cork cambium, characterized by rectangular cells arranged in radial files with heavily suberized walls creating waterproof barrier, completely dead cells with hollow air-filled lumens lacking protoplasts, compact radial organization with thin radial walls and thicker tangential walls, typical thickness 0.5-3 mm in individual cork layer with multiple cork layers present in periderms of old bark, showing progressive accumulation as cork cambium continuously produces new cork cells outward during active periods, demonstrating essential waterproofing function preventing trunk water loss through suberin impermeability to water and gases except through lenticels, and representing the most superficial component of periderm providing immediate protective interface between living trunk tissues and external environment. Cork layer rootstock periderm cellular structure includes= showing rectangular cells (cork cells typically 15-40 micrometers in radial dimension and 20-60 micrometers tangential dimension), exhibiting radial files (cork cells arranged in ordered rows perpendicular to trunk surface reflecting origin from cambial divisions), displaying thin radial walls (walls between radially adjacent cells relatively thin allowing some compression), showing thick tangential walls (walls parallel to trunk surface heavier providing strength), and demonstrating hollow lumens (dead cork cells empty air-filled creating low-density structure). Cork layer rootstock periderm suberin characteristics includes= showing lamellae structure (suberin deposited in alternating waxy and phenolic layers within cell wall), exhibiting hydrophobic properties (suberin lipid-rich composition creating water-repellent barrier), demonstrating impermeability (suberized walls essentially impermeable to water, gases, and most solutes), providing chemical resistance (suberin resistant to degradation by most microorganisms), and showing thermal stability (suberin maintaining barrier properties across wide temperature range). Cork layer rootstock periderm protective functions includes= preventing water loss effectively (suberized cork eliminating transpiration from trunk surface estimated >95% reduction), blocking pathogen entry (cork physical and chemical barrier preventing fungal and bacterial penetration), insulating thermally (cork low conductivity 0.04-0.05 W/m°K protecting cambium from temperature extremes), resisting mechanical damage (cork compressibility and resilience protecting trunk from impacts and abrasion), and providing fire resistance (cork relatively non-flammable offering some protection from fire damage). Cork layer rootstock periderm formation and accumulation includes= produced by cork cambium outward (phellogen dividing periclinally generating cork cells toward trunk surface), showing continuous production (active cork cambium adding new cork cells throughout growing season), accumulating progressively (cork layers increasing in thickness as cambium produces more cells), demonstrating variable rate (cork production typically 0.3-2 mm annually depending on cambium activity and cultivar), and thickening with age (older trunk showing thicker cork accumulation from years of production). Cork layer rootstock periderm in multiple periderms includes= present in each periderm (every successive periderm system containing its own cork layer), showing increasing depth (cork layers from older deeper periderms progressively deeper in bark), contributing to total bark (all cork layers collectively comprising major component of outer bark thickness), demonstrating layered structure (multiple cork layers creating laminated appearance in thick old bark), and varying in condition (outermost cork fresh and functional, deeper cork aged and degraded). Cork layer rootstock periderm air-filled structure includes= creating low density (air-filled cork cells producing lightweight protective tissue 0.12-0.25 g/cm³), providing insulation (air spaces contributing to low thermal conductivity), enabling compression (hollow cells allowing cork to compress under pressure then recover), demonstrating elasticity (cork resilient behavior useful for accommodating trunk expansion and external forces), and showing acoustic properties (air-filled structure providing some sound dampening potentially relevant for hail impact).
Cork_Layer_Rootstock_Periderm emphasizes phellem component of periderm and is distinguished from complete Rootstock_Periderm (including cambium and phelloderm) by focus on cork tissue specifically. Critical for understanding waterproofing, suberization, cellular structure, protective function, and the essential dead cork tissue forming outermost periderm layer providing immediate trunk surface protection.
Cork tissue
Korkschicht-Periderm
Outer periderm layer
Phellern-Lage
Äußere Peridermschicht
Evert, R. F. (2006). Esau's Plant Anatomy= Meristems, Cells, and Tissues of the Plant Body (3rd ed.). Wiley-Interscience.
Cork Layer Rootstock Periderm
Cost of production for Pinot Noir in Burgundy's Côte d'Or represents the extreme upper end of viticultural cost structures — establishment costs exceeding €1.5-3.0 million/ha for Grand Cru vineyard acquisition (reflecting land value rather than planting cost, as the vineyard is typically already established), annual production costs of €15,000-25,000/ha reflecting intensive hand labor in steep, dense-planted vineyards (10,000+ vines/ha), low permitted yields (35-45 hL/ha Grand Cru), and meticulous manual management, and harvest costs amplified by mandatory hand-picking at €3,000-5,000/ha — costs that are commercially viable only because Grand Cru Burgundy wine prices (€100-25,000+/bottle) provide the revenue to support this cost structure. Cost of production for Grüner Veltliner in the Weinviertel represents a more typical Central European cost profile — establishment cost of approximately €25,000-35,000/ha (land acquisition €10,000-20,000/ha plus planting cost €15,000-20,000/ha), annual production cost of approximately €4,000-6,000/ha, and harvest cost of approximately €800-1,500/ha depending on mechanization — commercially viable at the Weinviertel's moderate grape price levels (€0.40-0.60/kg) given the cultivar's reliable yields of 8,000-10,000 kg/ha.
Cost of production is an economic data entity that documents the financial expenditures required for grape cultivation and wine production, encompassing establishment costs (the capital investment required to create a new vineyard from bare land to first commercial harvest), annual production costs (the recurring operational expenditures of vineyard management and wine production in a mature vineyard), and harvest costs (the specific expenditures associated with grape harvesting operations), differentiated by cultivar (reflecting cultivar-specific management requirements), region (reflecting local input costs, labor rates, and regulatory requirements), and production system (reflecting the cost implications of viticultural approach — conventional, organic, biodynamic, precision viticulture), providing the cost-side economic framework that determines the minimum viable grape and wine price for commercially sustainable cultivation and enables profitability assessment across cultivars and regions.
Cost of production analysis provides the financial foundation for assessing cultivar economic viability — the essential calculation of whether the expected revenue from grape or wine sales exceeds the total cost of production by a margin sufficient to provide acceptable return on investment, service debt obligations, and sustain the operation over the multi-decade productive life of a vineyard. The cost structure of viticultural production is characterized by several distinctive features — high establishment costs with long payback periods (typically 5-8 years from planting to positive cumulative cash flow), significant fixed cost components (land, infrastructure, permanent equipment) that create economies of scale, high labor intensity relative to most agricultural sectors (particularly in premium production systems with extensive manual canopy management and hand harvesting), and substantial vintage-to-vintage revenue variability (driven by yield fluctuation and quality-dependent pricing) against relatively stable annual costs. The cultivar-specific dimension of production costs reflects the management requirements inherent to each variety — vigorous cultivars require more canopy management labor, disease-susceptible cultivars incur higher spray costs, thick-skinned cultivars may tolerate machine harvest while thin-skinned cultivars may require hand-picking, and cultivars with irregular set or coulure susceptibility generate more variable yields increasing per-unit cost in poor set years. The organic and biodynamic production system dimension adds a cost layer — conversion costs during the 3-year certification transition period, typically higher per-hectare management costs (10-30% above conventional due to increased labor for mechanical weed control, more frequent spray applications with less effective biological products, and lower average yields), partially offset by premium pricing for certified organic/biodynamic wines and by reduced chemical input costs. The regional cost variation reflects the enormous diversity of viticultural economic conditions globally — labor costs ranging from €2-3/hour in North African viticulture to €20-35/hour in Northern European and Australian viticulture, land costs from €5,000-10,000/ha in bulk-producing regions to €1-3 million/ha in Grand Cru Burgundy or Napa Valley, and regulatory compliance costs varying from minimal in unregulated New World regions to substantial in heavily regulated European appellations.
Anbaukosten
Bewirtschaftungskosten
Herstellungskosten
Produktionskosten
Weinproduktionskosten
cultivation cost
farming cost
production cost
viticultural cost
wine production expenditure
Galet P (2000) General Viticulture, Oenoplurimédia, pp.: 350-400 Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles, pp.: 1-10 Wittwer G, Rothfield J (2005) The global wine statistical compendium. In Anderson K (ed.) The World's Wine Markets, Edward Elgar Eurostat (2023) Agricultural production costs — vineyards
Cost Of Production
A cotyledonary node first leaf on a seedling of Vitis vinifera cv. Chardonnay × Pinot Noir cross in a breeding program nursery, positioned at node 1 immediately above the cotyledonary node where two cotyledons remain attached and partially functional, measuring 5.2 cm length by 4.8 cm width representing typical small seedling leaf dimensions, exhibiting simplified three-lobed architecture with shallow sinuses and reduced marginal teeth compared to adult Chardonnay leaf morphology showing characteristic five-lobed deeply sinused pattern, emerging 14 days post-germination following cotyledon expansion and greening, developing while cotyledons still contribute photosynthate providing approximately 40% of seedling carbon requirements during first leaf expansion, showing juvenile characteristics including very thin cuticle of 1.8 μm (compared to 4-6 μm in mature vine leaves), sparse trichomes creating nearly glabrous appearance unlike pubescent adult leaves, simple vascular architecture with primary veins only partially developed, demonstrating modest photosynthesis rate of 4.5 μmol CO₂ m⁻² s⁻¹ (compared to 8-14 μmol CO₂ m⁻² s⁻¹ in mature vine basal leaves) reflecting immature photosynthetic apparatus, occurring at seedling stage approximately 3 weeks post-germination, representing the critical transition from cotyledon-dependent to autotrophic seedling growth.
A first basal leaf at node 1 on seedling shoots originating directly above cotyledon attachment in Vitis vinifera, characterized by position at the first true node above the cotyledonary node where seed leaves (cotyledons) attach, distinctive juvenile morphology often showing simplified leaf architecture with reduced or absent lobing compared to adult leaf forms typical of mature vines, smaller dimensions than subsequent leaves typically 4-8 cm length in seedlings compared to 12-20 cm for mature vine basal leaves, early emergence following cotyledon expansion typically 10-18 days post-germination, development under strong influence of seed reserves and cotyledon photosynthesis providing resources for initial true leaf development, anatomical characteristics reflecting seedling physiology including thinner cuticle, reduced trichome density, and simpler vascular architecture compared to leaves on mature vegetative or reproductive shoots, and functional role as the first autotrophic leaf enabling seedling independence from seed reserves and cotyledon support. Cotyledonary node first leaves occur exclusively on seedling shoots during the juvenile phase of grapevine development, representing a unique developmental context distinct from mature vine basal leaves emerging from dormant buds containing preformed primordia, with seedling leaves developing from newly initiated primordia without previous-season developmental history, showing juvenile phase characteristics including absence of reproductive structures and often simplified morphology that gradually transitions toward adult forms over subsequent growth.
Cotyledonary node first leaves are distinguished from first basal leaves on mature vines by seedling context, position above cotyledons, juvenile morphological characteristics, and developmental from newly initiated rather than preformed primordia. These leaves rarely occur in commercial viticulture focused on vegetatively propagated cultivars, appearing primarily in breeding programs, genetic studies, or rootstock seedling production. Morphological characteristics often differ substantially from adult forms, with progression toward adult morphology occurring over subsequent nodes and requiring months to years depending on species and environmental conditions. The juvenile-to-adult transition in grapevine involves progressive changes in leaf morphology, growth habit, thorniness, and reproductive competence, with cotyledonary node leaves representing the extreme juvenile end of this developmental spectrum. Functional capacity of cotyledonary node first leaves is limited compared to mature vine leaves, with reduced photosynthetic rates, simpler anatomy, and smaller size, but these leaves play critical roles in seedling establishment enabling transition from heterotrophic seed-dependent growth to autotrophic photosynthesis-based growth.
Keimblatt-Knoten erstes Blatt
Sämlingsblatt
juvenile first leaf
juveniles erstes Weinblatt
post-cotyledon leaf
seedling first leaf
Based on Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 109-134 This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Carmona et al. (2008) "Grapevine bud dormancy= environmental and molecular aspects" Revista Brasileira de Fruticultura 30=257-408 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 1-32.
Cotyledonary Node First Leaf
Grenache is notoriously susceptible to coulure, with fruit set dropping to 10-20% in cool or rainy flowering seasons compared to 30-40% under optimal conditions. Malbec shows severe coulure susceptibility with yield losses of 40-70% in years with temperatures below 15°C during bloom. Muscat cultivars including Muscat of Alexandria and Muscat Ottonel exhibit high coulure susceptibility linked to functional pistil abnormalities. Coulure severity correlates with flower abscission rates exceeding 60% within two weeks post-bloom.
A fruit set disposition that is the inherited capacity of a Vitis vinifera plant to exhibit flower or young berry abscission (coulure) under unfavorable conditions, resulting in reduced fruit set and loose clusters due to failure of fertilization, embryo abortion, or disruption of early berry development.
Coulure_Susceptibility_Disposition serves as the parent class for weather-induced and nutritional subtypes. The trait reflects both inherent sensitivity of reproductive processes and inadequate buffering capacity against environmental stress. Cultivars with long flowering periods are often more susceptible due to extended exposure to unfavorable conditions. Selection against coulure susceptibility has been a major breeding objective for cool climate viticulture.
Blütenabwurf-Disposition
Durchrieselungsanfälligkeit
Verrieselungsneigung
flower drop tendency
poor set disposition
shatter susceptibility
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 125-130 Galet P (2000) General Viticulture, Oenoplurimedia, Chaintre pp.: 136-145
Coulure Susceptibility Disposition
Coulure susceptibility comparison among cultivars under identical challenging flowering conditions (15°C average, frequent rain) reveals the genetic variation in abscission resistance, with highly susceptible Grenache showing severe coulure (only 12% fruit set, sparse clusters with <40 berries, 75% yield loss), moderately susceptible Merlot showing moderate coulure (25% set, 85 berries/cluster, 40% yield loss relative to potential), and resistant Cabernet Sauvignon showing minimal coulure (38% set, 125 berries/cluster, maintaining near-normal yield), demonstrating how coulure susceptibility determines yield reliability under stressful flowering conditions with implications for cultivar-climate matching.
Coulure susceptibility quality is a yield potential quality that represents the tendency of grape cultivars or vines to experience excessive flower and fruitlet abscission during and after flowering, characterizing the vulnerability to the physiological disorder where flowers fail to set or young berries drop shortly after set, substantially reducing cluster fill and yield potential. Within the BFO framework, this quality specifically addresses the abscission vulnerability dimension of yield potential, distinguishing susceptibility to this specific disorder from normal fruit set variation while recognizing that coulure represents pathological failure of fruit retention beyond normal set losses—affected clusters showing sparse berry populations with large gaps where flowers or fruitlets have abscised rather than the moderate spacing of normal set. The ontological significance lies in coulure susceptibility determining the risk of severe yield loss from conditions that trigger excessive abscission, where susceptible cultivars may lose 50-80% of potential berries under unfavorable flowering conditions while resistant cultivars maintain acceptable set under similar stress, with this differential susceptibility being a key factor in cultivar selection for climates with challenging flowering weather. The causes of coulure include carbohydrate deficiency during flowering (inadequate photosynthate for berry retention, often from cloudy, cool weather limiting carbon fixation during the high-demand flowering period), hormonal imbalance (insufficient auxin production in developing seeds failing to establish strong sink status), poor pollination (failed fertilization preventing seed development and the hormonal signals supporting berry retention), and genetic susceptibility (some cultivars inherently prone to abscission under stress). The cultivar variation in coulure susceptibility ranges from highly susceptible varieties (Grenache, Merlot, Gewürztraminer showing severe coulure under marginal conditions) to resistant varieties (Cabernet Sauvignon, Chardonnay maintaining reasonable set under similar stress), with this variation affecting cultivar suitability for different climates and the management intensity required to achieve reliable production. Understanding coulure susceptibility quality enables cultivar selection appropriate for local flowering conditions, management targeting reduced coulure risk (balanced vines, appropriate nitrogen), and realistic yield expectations for susceptible varieties in challenging climates.
Coulure_Susceptibility_Quality addresses the vulnerability to excessive flower and fruitlet abscission reducing yield. This quality varies genetically among cultivars and is triggered by flowering-period stress. The class supports cultivar selection and risk assessment through coulure susceptibility characterization.
Abszissionsempfindlichkeit
Blütenabwurfneigung
Coulure-Anfälligkeit
Verrieselungsanfälligkeit
abscission vulnerability
coulure proneness
Carbonneau A et al. (2003) Traité de viticulture. Dunod, Paris, pp. 1-400 May P (2004) Flowering and Fruitset in Grapevines. Lythrum Press, Adelaide, pp. 1-119
Coulure Susceptibility Quality
Coutaric acid content quality assessment in Barossa Valley Shiraz quantifies the volatile phenol precursor pool relevant to Brettanomyces spoilage risk, with typical concentrations of 40-80 mg/L providing substantial substrate for 4-ethylphenol formation winemakers managing Brett risk through SO₂ maintenance above 35 mg/L molecular, pH control below 3.6, and rigorous hygiene recognize that coutaric acid content establishes the ceiling for potential ethylphenol accumulation, making low-coutaric fruit preferable for age-worthy wines intended for extended barrel maturation.
Coutaric_Acid_Content_Quality represents the chemical quality characterizing the concentration of coutaric acid (2-p-coumaroyl-L-tartaric acid, trans-p-coumaroyltartaric acid) in grape berry tissue, capturing the second most abundant hydroxycinnamic acid derivative in Vitis vinifera grapes. Coutaric acid consists of p-coumaric acid (4-hydroxycinnamic acid) esterified to L-tartaric acid, differing from caftaric acid by the absence of the 3-hydroxyl group on the aromatic ring. This single hydroxyl group difference substantially affects chemical behavior, as coutaric acid is a poorer substrate for polyphenol oxidase than caftaric acid and does not form stable glutathione conjugates. Coutaric acid typically comprises 20-40% of total hydroxycinnamic acids, with concentrations ranging from 20-100 mg/L in grape juice. The coutaric-to-caftaric acid ratio varies among cultivars, providing fingerprinting information useful for varietal identification. In winemaking, coutaric acid gains particular significance as the precursor for p-coumaric acid release through hydrolysis, which can then serve as substrate for vinylphenol reductase in Brettanomyces and other microorganisms. The sequential conversion of coutaric acid to p-coumaric acid to 4-vinylphenol to 4-ethylphenol represents a major pathway for Brettanomyces spoilage compound formation. Higher coutaric acid content indicates greater precursor availability for volatile phenol production, making it relevant to spoilage risk assessment in red wines where Brettanomyces contamination poses quality threats. White wines show less volatile phenol development despite similar coutaric acid content because their production conditions disfavor Brettanomyces growth.
Coutaric_Acid_Content_Quality addresses the p-coumaric acid tartrate ester that serves as precursor for volatile phenol formation through microbial metabolism. The structural difference from caftaric acid (mono- versus di-hydroxylated ring) affects both oxidative behavior and downstream metabolism. The pathway coutaric acid → p-coumaric acid → 4-vinylphenol → 4-ethylphenol represents the major route to Brettanomyces spoilage compounds, making coutaric acid content a risk factor for wines susceptible to Brett contamination. Analysis parallels caftaric acid using HPLC with UV detection at 310-320 nm.
Coumaroyltartrat-Konzentration
Coumaroylweinsäure-Gehalt
Cutarsäuregehalt
Ethylphenol-Vorläuferqualität
coumaroyl tartrate content
coumaroyltartaric acid concentration
coutaric acid level
ethylphenol precursor quality
Chatonnet P et al. (1992) The Origin of Ethylphenols in Wines. Journal of the Science of Food and Agriculture 60(2): 165-178 Hernández T et al. (2006) Hydroxycinnamic Acids and Ferulic Acid Decarboxylase in the Determination of Guaiacol and 4-Vinylguaiacol in Beer. Food Research International 39(1): 66-72
Coutaric Acid Content Quality
Merlot vines in the Colchagua Valley of Chile bearing a cover crop integration role are cultivated with permanent inter-row cover of Trifolium subterraneum and Lolium perenne providing 60-120 kg N/ha/year through biological fixation, reducing irrigation requirements by 15-20% through improved soil water-holding capacity at 22-28% gravimetric moisture versus 14-18% in bare-soil controls, increasing soil organic matter from 1.2% to 2.1% over 8 years of cover cropping, and supporting predatory carabid beetle populations at 45-80 individuals per pitfall trap versus 12-25 in herbicide-maintained inter-rows.
An ecological role that inheres in a grapevine cultivated in association with deliberately established inter-row or under-vine cover crop vegetation, where the vine-cover crop system functions as an integrated agroecological unit providing nutrient cycling, soil structure improvement, water management, and habitat diversification beyond what either component achieves in isolation.
Cover_Crop_Integration_Role recognizes the grapevine as one component of a designed polyculture system rather than a monoculture stand. The ecological interactions between vine and cover crop are bidirectional and complex: cover crops compete with vines for water and nutrients (potentially beneficial in vigorous sites, detrimental in water-limited conditions), modify vine microclimate through altered albedo and humidity, suppress weeds through competitive exclusion, and fundamentally reshape the soil biome. This role intersects with multiple production system roles (organic, biodynamic, regenerative, sustainable) where cover cropping is a core management component.
Begrünungs-Integrations-Rolle
Deckfrucht-Rolle
Gründecken-Bewirtschaftungs-Rolle
cover-cropped vine role
green cover integration role
inter-row vegetation role
Celette F et al. (2009) Competition for nitrogen in an unfertilized intercropping system: grapevine and permanent cover, Environmental and Experimental Botany 65: 102-110 Guerra B and Steenwerth K (2012) Influence of floor management technique on grapevine growth, disease pressure, and juice and wine composition, American Journal of Enology and Viticulture 63: 244-251 Winter S et al. (2018) Effects of vegetation management intensity on biodiversity in vineyards, Agriculture Ecosystems and Environment 257: 172-181
Cover Crop Integration Role
The cover crop specification for a Cabernet Sauvignon vineyard in Coonawarra prescribes a managed alternate-row system — even-numbered inter-rows maintain permanent perennial ryegrass (Lolium perenne) and white clover (Trifolium repens) sown at 25 kg/ha and 3 kg/ha respectively, mowed to 8-10 cm height 6-8 times per season, providing year-round soil surface protection and nitrogen fixation estimated at 30-50 kg N/ha/year from the clover component, while odd-numbered inter-rows are maintained as bare soil by shallow cultivation (3 passes per season) to reduce total vineyard water demand by approximately 15-20% compared to full inter-row cover — this alternate-row strategy balances the soil health and erosion control benefits of permanent cover against the water competition reduction required in the semi-arid Coonawarra climate where growing-season rainfall of 200-250 mm provides only 40-50% of vine water requirements.
A soil management description that documents the species composition, establishment method, management regime, and functional objectives of vegetation intentionally grown or maintained in vineyard inter-rows and/or under-vine strips, specifying the cover crop type (permanent, annual, or self-reseeding), spatial distribution (every row, alternate rows, or under-vine), and management by mowing, rolling, or incorporation that collectively determine the cover crop's effects on soil health, water competition, nutrient cycling, and vineyard biodiversity.
Cover_Crop_Specification documents the most significant shift in vineyard floor management philosophy over the past 40 years — the transition from bare-soil management (herbicides or clean cultivation) to managed vegetation systems that treat the vineyard floor as a functional component of the vineyard ecosystem rather than merely a surface to be kept weed-free. Cover crop species selection involves deliberate trade-offs between functional objectives: grasses (Festuca, Lolium, Hordeum) provide rapid soil coverage, excellent erosion control, and competitive water uptake that can usefully devigorate excessively vigorous vines but may exacerbate water stress in dry conditions legumes (Trifolium, Vicia, Medicago) fix atmospheric nitrogen and reduce fertilizer requirements but provide less effective erosion control and diverse multi-species mixtures increasingly favored in biodiversity-oriented viticulture combine functional traits while supporting broader arthropod and microbial communities. The spatial management of cover crops — permanent versus seasonal, every-row versus alternate-row, inter-row versus under-vine — provides fine-tuned control over the balance between soil health benefits and resource competition.
Begrünungs-Spezifikation
Gründecke-Protokoll
Zwischenbegrünungs-Dokumentation
cover crop protocol
green cover management record
vineyard floor vegetation specification
Celette F et al. (2009) Dynamics of water and nitrogen stress along the grapevine cycle in a Mediterranean vineyard, European Journal of Agronomy 30: 38-46 White RE (2015) Understanding Vineyard Soils 2nd Ed, Oxford University Press pp: 210-240 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 440-455 Guerra B and Steenwerth KL (2012) Influence of floor management technique on grapevine growth, disease pressure, and juice and wine composition, American Journal of Enology and Viticulture 63: 244-255
Cover Crop Specification
Creaminess quality in aged Champagne (10 years sur lie, creaminess rating 8/10) demonstrates the textural transformation from young wine's fresh acidity to mature wine's luxurious, dairy-like richness arising from mannoprotein release during extended lees contact, creating the enveloping mouthfeel that complements the developed brioche-hazelnut flavors and justifies premium pricing for aged bottlings, while non-vintage Champagne with minimal lees time shows crisp, fresh mouthfeel (creaminess 3/10) appropriate for its immediate-consumption style.
Creaminess quality is a mouthfeel quality that represents the rich, smooth, dairy-like tactile impression arising from elevated viscosity combined with softness and roundness, characterizing the luxurious, enveloping mouthfeel particularly associated with barrel-fermented and lees-aged white wines and certain full-bodied wine styles. Within the BFO framework, this quality specifically addresses the rich-smooth dimension of mouthfeel, distinguishing creamy texture from simple smoothness or high viscosity by requiring the combination of lubricating smoothness with substantial body and soft, round character. The ontological significance lies in creaminess representing a specific textural quality highly valued in certain wine styles, where the dairy-like richness creates the luxurious mouthfeel associated with premium barrel-fermented Chardonnay, aged Champagne, and other wines where textural opulence complements flavor complexity, though the same character may be inappropriate in styles emphasizing freshness and crispness. The perceptual characteristics of creaminess include the soft, coating, enveloping sensation reminiscent of dairy products, with substantial but not heavy body, smooth movement without friction, and the round, cushioned quality that creates the impression of richness and luxury. The compositional determinants of creaminess include mannoproteins and polysaccharides released during lees contact (primary contributors to creamy perception), malolactic fermentation (converting sharp malic acid to softer lactic acid), barrel fermentation (contributing textural complexity), glycerol concentration (adding smoothness), and moderate alcohol (providing body without heat), with these factors combining to create the characteristic creamy profile. The production associations of creaminess include extended sur lie aging (particularly with bâtonnage), barrel fermentation, and full malolactic conversion, with these techniques deliberately employed to develop creamy texture in styles where it is valued. Understanding creaminess quality enables style recognition based on textural richness, production method inference from creamy character, and quality assessment incorporating appropriate creaminess for wine style.
Creaminess_Quality addresses the rich-smooth textural dimension particularly associated with lees-aged wines. This quality is stylistically specific, valued in certain wine categories while inappropriate in others emphasizing freshness. The class supports style recognition, production method inference, and appreciation of how winemaking techniques create textural opulence through mannoprotein and polysaccharide enrichment.
Cremigkeit
creamy texture
cremiger Charakter
dairy-like mouthfeel
lactic smoothness
milchiges Mundgefühl
sahnige Textur
unctuous richness
Vidal S et al. (2004) Use of an experimental design approach for evaluation of key wine components on mouth-feel perception. Food Quality and Preference 15(3): 209-217 Chalier P et al. (2007) Interactions between aroma compounds and whole mannoprotein extracts from Saccharomyces cerevisiae. Food Chemistry 100(1): 22-30
Creaminess Quality
Creep behavior assessment of Thompson Seedless berries under constant 0.5 N compression reveals instantaneous strain of 3.2% followed by creep strain accumulating to 8.7% over 60 minutes, with creep compliance of 11 × 10⁻³ MPa⁻¹ indicating significant time-dependent deformation that explains the progressive flattening observed in bottom layers of packed table grape clamshells during 4-week cold storage, motivating package redesign to reduce per-berry load below the 0.2 N threshold where creep becomes negligible.
Creep behavior quality is a viscoelastic property quality that represents the time-dependent increase in deformation occurring under constant applied stress, characterizing the progressive strain accumulation as polymer chains rearrange and cellular fluids redistribute in response to sustained mechanical loading. Within the BFO framework, this quality specifically addresses the temporal evolution of deformation under maintained stress, distinguishing creep from instantaneous elastic deformation and from stress relaxation which addresses force changes under constant strain. The ontological significance lies in creep behavior determining the long-term mechanical stability of tissues under sustained loads, where the magnitude and rate of creep affect storage deformation in stacked packaging, hanging cluster distortion under gravitational load, and progressive tissue failure under subliminal but sustained stress. Creep in grapevine tissues follows characteristic kinetics with an initial rapid elastic deformation, followed by primary creep (decreasing creep rate), secondary creep (constant creep rate), and potentially tertiary creep (accelerating rate leading to failure), with the transition between regimes depending on stress magnitude relative to tissue strength. The cellular basis of creep involves multiple mechanisms including cell wall polymer chain slippage and reorientation, pectin gel network rearrangement, water redistribution between cellular compartments, and turgor-driven flow through plasmodesmata, with each mechanism contributing different time constants to the overall creep response. Temperature acceleration of creep reflects the thermally activated nature of polymer chain motion, with the time-temperature superposition principle enabling prediction of long-term creep from short-term tests at elevated temperature. Measurement employs constant-stress loading (often using dead weights) with continuous strain recording over extended periods, with creep compliance (strain per unit stress) providing the standardized creep parameter enabling comparison across stress levels and materials. Understanding creep behavior quality enables prediction of time-dependent deformation during storage and handling, design of packaging to minimize creep-induced damage, and interpretation of texture changes during prolonged mechanical loading.
Creep_Behavior_Quality specifically addresses deformation increase under constant stress, complementing Stress_Relaxation_Quality which addresses stress decrease under constant strain. This quality is critical for storage and packaging applications where loads are sustained over extended periods. The class supports long-term deformation prediction and packaging optimization through characterization of time-dependent mechanical response to sustained loading.
Kriechdehnung
Kriechverhalten
Retardation
cold flow
creep compliance
retarded elasticity
time-dependent strain
zeitabhängige Verformung
Mohsenin NN (1986) Physical Properties of Plant and Animal Materials, 2nd Ed. Gordon and Breach, New York, pp. 556-619 Alamar MC et al. (2008) Micromechanics of grape berry tissue. Postharvest Biology and Technology 49(1): 50-60
Creep Behavior Quality
Critical temperature threshold analysis for Cabernet Sauvignon in Napa Valley identifies primary bud lethal temperature of -18°C during full dormancy declining to -2.5°C at budbreak, berry heat damage threshold of 40°C for anthocyanin degradation onset and 45°C for cell death, establishing the thermal envelope of 57°C (-12°C to +45°C) within which this variety maintains viability, enabling site selection and climate change scenario evaluation based on historical temperature extremes and projected future conditions.
Critical temperature threshold quality is a temperature quality that represents specific temperature values at which qualitative changes in physiological function, developmental trajectory, or tissue viability occur, marking the boundaries between normal function and stress response, reversible stress and irreversible damage, or survival and mortality. Within the BFO framework, this quality characterizes discrete transition points in the continuous temperature spectrum where biological responses show discontinuous or threshold-dependent changes rather than gradual proportional responses, enabling definition of thermal tolerance limits and optimal ranges. The ontological significance lies in threshold temperatures serving as critical reference values for stress assessment and management decisions, where temperatures below freezing thresholds cause ice formation and cellular damage, temperatures above heat damage thresholds cause protein denaturation and membrane disruption, and temperatures within the optimal range support maximum physiological performance. Grapevine critical temperature thresholds vary by tissue type, developmental stage, acclimation status, and the rate and duration of temperature change, with cold-hardened dormant buds surviving -25°C while actively growing shoot tips are damaged at -2°C, and photosynthetic optimum varying from 25-30°C for temperate-adapted varieties to 30-35°C for warm-climate varieties. The determination of critical thresholds employs controlled temperature exposure studies measuring physiological parameters, field observations correlating damage with temperature records, and modeling approaches predicting damage probability as functions of temperature and duration. Understanding critical temperature thresholds enables climate suitability assessment for variety selection, frost and heat warning system development, and evaluation of climate change impacts on viticulture viability in specific regions.
Critical_Temperature_Threshold_Quality serves as the parent class for specific threshold types (Freezing_Point, Heat_Damage_Threshold, Optimal_Photosynthesis_Temperature), establishing the ontological principle that certain temperatures represent qualitative rather than merely quantitative transitions in biological response. Threshold values are not absolute but vary with acclimation, tissue type, and exposure dynamics, requiring context specification for accurate application. The class supports risk assessment and climate suitability analysis.
Kardinaltemperatur
Temperaturgrenzwert
cardinal temperature
critical thermal point
kritische Temperaturschwelle
temperature limit
thermal tolerance threshold
thermischer Schwellenwert
Mullins MG et al. (1992) Biology of the Grapevine. Cambridge University Press, Cambridge, pp. 152-178 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed. Academic Press, San Diego, pp. 295-340
Critical Temperature Threshold Quality
Cross-compatibility between V. vinifera and other Vitis species ranges from highly compatible to essentially incompatible depending on the species pair. V. vinifera × V. rupestris crosses show high compatibility with 80-95% fruit set and 70-90% seed germination, enabling creation of rootstocks such as 3309C and 101-14 Mgt. V. vinifera × V. labrusca shows moderate compatibility (50-70% fruit set) with offspring exhibiting variable wine quality (foxy character inheritance). V. vinifera × V. rotundifolia crosses face severe post-zygotic barriers, with embryo abortion occurring 20-40 days post-fertilization in 90-98% of ovules, requiring embryo rescue for hybrid recovery. Pollen tube growth in incompatible crosses may proceed normally to fertilization, with barriers operating primarily at embryo development stages.
An interspecific hybridization disposition that is the inherited capacity of a Vitis vinifera plant to produce viable seeds when crossed with specific other Vitis species or genotypes, determined by pollen-pistil interactions, fertilization success, and embryo development through seed maturation without lethal incompatibility.
Cross-compatibility encompasses both pre-zygotic barriers (pollen germination, tube growth, fertilization) and post-zygotic barriers (embryo development, seed viability, seedling survival). Most Vitis species within the Vitis subgenus show adequate cross-compatibility for breeding, with the Muscadinia subgenus (V. rotundifolia) representing the major crossing barrier. Reciprocal cross differences occur in some combinations, with success depending on which species serves as maternal versus paternal parent. Marker-assisted selection enables efficient introgression of resistance loci while minimizing unfavorable hybrid characteristics.
Befruchtungsverträglichkeit
Kreuzungskompatibilität
crossing success
fertilization compatibility trait
sexual compatibility
sexuelle Kompatibilität
Bouquet A (1980) Vitis × Muscadinia hybridization: a new way in grape breeding for disease resistance in France. Proceedings of the 3rd International Symposium on Grape Breeding pp.: 42-61 Patel GI, Olmo HP (1955) Cytogenetics of Vitis: I. The hybrid V. vinifera × V. rotundifolia. American Journal of Botany 42:141-159
Cross Compatibility Disposition
Cross number Gf.1993-22 at the Julius Kühn-Institut designates the controlled pollination of Bacchus × Villard Blanc performed in 1993 as the 22nd cross combination of that year's breeding program, with 450 seeds obtained from this cross yielding 380 viable seedlings that were collectively identified by this family-level cross number through initial screening — individual seedlings within the family were then distinguished by appending a seedling number (e.g., Gf.1993-22-6 for the individual that became Calardis Blanc), but the cross number Gf.1993-22 remains the identifier linking all siblings from this parental combination in the breeding database.
A breeding code that identifies a specific controlled cross combination performed within a breeding program, encoding the parental combination and year of crossing to uniquely designate the family of seedlings generated from a defined maternal × paternal pollination event.
Cross_Number operates at the family level of breeding program organization, identifying the parental combination rather than the individual genotype. Each year a grapevine breeding program may perform 20-100 controlled crosses, each generating hundreds to thousands of seedlings, and the cross number provides the hierarchical grouping that organizes this material: the cross number is the primary unit for evaluating the value of specific parental combinations, calculating combining ability, and identifying promising cross combinations for repeat pollination. Cross numbers are essential for breeding program management but are rarely referenced in commercial or regulatory contexts, appearing primarily in breeding reports and research publications.
Kombinationsnummer
Kreuzungs-Kennung
Kreuzungsnummer
cross combination number
family identifier
hybridization code
Cross Number
Hydraulic architecture studies in drought-stressed Grenache reveal that xylem cross-sectional area in the shoot-to-leaf pathway decreases by 40-60% from basal internode (0.8-1.2 mm²) to distal petiole (0.15-0.25 mm²), creating a hydraulic bottleneck that promotes stomatal closure and protects the main stem vasculature from cavitation under water deficit, exemplifying the iso-to-anisohydric continuum of stomatal behavior in grapevine cultivars.
Cross_Sectional_Area_Quality represents the two-dimensional extent revealed when a planar cut is made through a grapevine organ perpendicular to a defined axis. This quality characterizes the internal spatial organization of organs and provides essential data for understanding vascular transport capacity, mechanical strength, and tissue proportions. Cross-sectional area of vascular tissues directly determines hydraulic conductance according to the Hagen-Poiseuille relationship, where flow capacity scales with the fourth power of vessel radius. Xylem cross-sectional area in shoots, petioles, and cluster rachises thus limits water supply to distal organs under high transpirational demand. Phloem cross-sectional area constrains photoassimilate transport to developing clusters, potentially limiting berry growth rate and final sugar accumulation in heavily cropped vines. Trunk cross-sectional area at standardized heights serves as a non-destructive proxy for whole-vine biomass and productive capacity, enabling yield estimation models and growth tracking. Berry cross-sectional area reveals the proportional contribution of skin, pulp, and seed compartments, with implications for wine composition. Cross-sectional measurements require either destructive sampling followed by image analysis of cut surfaces, or non-destructive imaging techniques including X-ray computed tomography, magnetic resonance imaging, or ultrasound. For approximately circular cross-sections, area can be calculated from diameter measurements irregular cross-sections require direct planimetric measurement or image analysis.
Cross_Sectional_Area_Quality requires specification of the cutting plane orientation (transverse, longitudinal, tangential) relative to organ axes, and the position along the organ where the section is taken. Variability in cross-sectional area along an organ's length is common and functionally significant. The class relates to but is distinct from the tissues and structures revealed in cross-section (material entities) and from the processes that generate cross-sectional dimensions (developmental processes). For xylem vessels, both individual vessel lumen area and total conductive area (sum of all vessels) are relevant measurements with different physiological interpretations.
Querschnittsbereich
Querschnittsfläche
Schnittflächenqualität
cut surface area
internal area extent
sectional area quality
transversale Flächenausdehnung
transverse section area
Tyree MT and Ewers FW (1991) The Hydraulic Architecture of Trees and Other Woody Plants. New Phytologist 119(3): 345-360 Schultz HR (2003) Differences in Hydraulic Architecture Account for Near-Isohydric and Anisohydric Behaviour of Two Field-Grown Vitis vinifera L. Cultivars during Drought. Plant, Cell and Environment 26(8): 1393-1405
Cross Sectional Area Quality
Systematic screening for Crown_Gall_Resistance_Disposition at Cornell University evaluated 120 Vitis accessions using standardized wound inoculation with tumorigenic Agrobacterium vitis strain CG49 on greenhouse-grown potted vines. Wounded stems were inoculated with bacterial suspension (10⁸ CFU/mL), and gall development was scored at 6, 12, and 20 weeks post-inoculation using a 0-5 scale based on gall size and coverage of the wound site. V. vinifera cultivars showed uniformly high susceptibility with mean scores of 4.2-4.8 across 40 tested cultivars. Wild species V. riparia (score 3.8), V. rupestris (4.1), and V. amurensis (4.5) showed equivalent susceptibility. Interspecific rootstock hybrids ranged from 3.5 to 4.6 with no commercially relevant resistance identified. The only accessions showing potentially useful Crown_Gall_Resistance_Disposition were certain V. labrusca × V. vinifera hybrids ('Niagara', score 2.1 'Catawba', score 2.4), though these remain susceptible and are unsuitable as wine grapes due to labrusca flavor characteristics. These findings confirm that breeding for crown gall resistance faces severe germplasm limitations, with no immunity and only moderate tolerance available in non-vinifera backgrounds with quality compromises.
A Crown_Gall_Resistance_Disposition is a Bacterial_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that limit infection, transformation, or tumor development following exposure to Agrobacterium vitis (syn. Allorhizobium vitis), the tumorigenic bacterium causing crown gall disease of grapevine. This disposition represents the capacity to resist the transformation of plant cells by bacterial T-DNA and/or the subsequent proliferation of transformed cells into characteristic galls at wound sites on trunks, cordons, and graft unions. Crown gall disease develops when wounded plant tissues are colonized by tumorigenic Agrobacterium strains carrying Ti (tumor-inducing) plasmids. The bacteria transfer T-DNA into plant cell nuclei, where it integrates into chromosomes and directs synthesis of auxins and cytokinins that cause uncontrolled cell proliferation (tumor formation) and opines that provide nutrients for bacterial growth. Crown gall resistance mechanisms potentially include interference with bacterial attachment to wound surfaces, inhibition of T-DNA transfer across plant cell membranes, prevention of T-DNA nuclear import or chromosomal integration, resistance to hormone-mediated transformation, and active elimination of transformed cells. Despite extensive screening, no commercially useful level of crown gall resistance has been identified in V. vinifera or most Vitis species. Crown gall is particularly damaging in cold climate viticulture where winter injury creates wound sites susceptible to infection, and in nurseries where grafting wounds provide entry points. Management relies primarily on prevention through use of non-infested planting material and protection of wound sites rather than genetic resistance.
Crown_Gall_Resistance_Disposition is_a Bacterial_Resistance_Disposition. Crown gall is caused by Agrobacterium vitis (grapevine-specific) rather than the broad-host-range A. tumefaciens. Child classes include Agrobacterium_Transformation_Resistance_Disposition (resistance to T-DNA transfer/integration) and Wound_Healing_Efficiency_Disposition (rapid wound closure preventing infection). The absence of effective resistance in V. vinifera and most breeding-relevant germplasm distinguishes crown gall from Pierce's disease where strong resistance sources exist. Cold climate viticulture faces greatest crown gall pressure due to winter injury creating infection sites.
Agrobacterium resistance disposition
Agrobakterien-Resistenz
Mauke-Resistenz-Disposition
Wurzelhalsgallen-Resistenz
crown gall tolerance
gall formation resistance
Burr TJ, Otten L (1999) Crown gall of grape: biology and disease management, Annual Review of Phytopathology 37:53-80 Szegedi E, Civerolo EL (2011) Bacterial diseases of grapevine, International Journal of Horticultural Science 17:45-49 Pulawska J (2010) Crown gall of stone fruits and nuts, economic significance and diversity of its causal agents: tumorigenic Agrobacterium spp., Journal of Plant Pathology 92:S1.87-S1.98
Crown Gall Resistance Disposition
Crown gall susceptibility quality assessment in Finger Lakes vineyards following repeated freeze events evaluated the relationship between cumulative cold injury and progressive crown gall development affecting vineyard productivity and longevity. Survey of 15-year-old Chardonnay and Riesling plantings with documented freeze damage history quantified the correlation between cold damage and gall incidence. Chardonnay showed high crown gall susceptibility quality with 65% of vines exhibiting trunk galls, gall incidence strongly correlated with cumulative freeze damage score (r² = 0.72), and average gall circumference of 8.5 cm in affected vines causing visible constriction of trunk vascular function. Riesling demonstrated lower crown gall susceptibility quality with only 28% gall incidence despite equivalent freeze exposure history, weaker correlation with damage score (r² = 0.45) suggesting partial resistance, and smaller average gall size (4.2 cm) indicating slower disease progression. The crown gall susceptibility quality differential translated to vineyard longevity, with severely galled Chardonnay vines showing 35% yield reduction compared to ungalled vines and 20% mortality rate over the 15-year period, while galled Riesling showed only 15% yield reduction and 8% mortality. The crown gall susceptibility quality assessment established that cultivar selection for cold-climate sites should consider gall resistance as well as cold hardiness, since highly susceptible cultivars may show poor long-term productivity even with adequate freeze survival due to progressive crown gall development at freeze wound sites.
Crown gall susceptibility quality is a cold recovery quality that inheres in a grapevine organism and represents the vulnerability to infection by Agrobacterium vitis (formerly Agrobacterium tumefaciens biovar 3) following freezing injury that creates wound sites where the bacterium can enter and initiate tumor formation, with crown gall susceptibility determining whether freeze wounds heal normally or become sites of gall development that progressively compromises vine structure and function. This quality addresses a critical interaction between cold damage and disease that particularly affects cold-climate viticulture, recognizing that freeze wounds provide the primary infection courts for crown gall such that the disease is largely a consequence of freezing injury in vineyards where A. vitis is present, with crown gall representing the major biotic factor limiting vine longevity and productivity in regions with frequent freeze damage. The ontological significance of crown gall susceptibility quality lies in its role converting temporary freeze injury into progressive structural disease, with gall formation at wound sites expanding over subsequent seasons to eventually girdle trunks and reduce vine productivity to the point of requiring removal, such that crown gall susceptibility transforms recoverable freeze damage into long-term vineyard decline. The biological basis of crown gall susceptibility encompasses the survival of A. vitis in vineyard soils and systemically within vine tissues, the wound-dependent nature of infection requiring freeze or mechanical injury to provide bacterial entry, the genetic transformation of plant cells by bacterial T-DNA that creates the undifferentiated tumor growth characteristic of crown gall, and the vine factors affecting susceptibility including genotype, tissue type, and physiological status that influence whether infection leads to gall formation or is contained by host defenses. The cultivar variation in crown gall susceptibility reflects genetic differences in resistance mechanisms, with some cultivars showing strong resistance to gall formation even when infected with A. vitis while others are highly susceptible, creating important differences in the long-term consequences of freeze damage in vineyards where the pathogen is present. The management of crown gall susceptibility involves reducing freeze injury to limit wound creation, selecting resistant cultivars and rootstocks where available, maintaining vine health to enhance wound healing capacity, and removing and destroying galled tissue to reduce inoculum pressure.
Crown_Gall_Susceptibility_Quality is_a Cold_Recovery_Quality, representing the vulnerability to Agrobacterium vitis infection at freeze wound sites leading to progressive structural disease. This class addresses the critical cold-disease interaction in cold-climate viticulture. Freeze wounds provide the primary infection courts for crown gall. Cultivar resistance varies substantially (Riesling > Chardonnay in examples). Cumulative freeze damage correlates with gall incidence where A. vitis is present. Long-term vineyard productivity and longevity affected by progressive gall development. Cultivar selection should consider gall resistance alongside cold hardiness.
Agrobacterium susceptibility
Agrobakterium-Anfälligkeit
Frostwunden-Infektionsanfälligkeit
Gallenbildungs-Vulnerabilität
Mauke-Anfälligkeit
crown gall disease risk
freeze-wound infection susceptibility
gall formation vulnerability
Burr TJ, Otten L (1999) Crown gall of grape: biology and disease management. Annual Review of Phytopathology 37:53-80 Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection. Michigan State University Extension Bulletin E2930 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 508-545
Crown Gall Susceptibility Quality
Crown gall symptom quality assessment following severe winter freeze damage (-30°C) in Niagara Peninsula Riesling documented the secondary disease impact compounding direct cold injury. Survey 18 months post-freeze identified crown gall symptom quality in 35% of vines that had shown trunk-level freeze damage, with galls developing at sites of freeze cracking and cambium injury where A. vitis had entered through wounds. Gall size ranged from small nodules 1-2 cm diameter on recently infected sites to large irregular masses 8-15 cm diameter on vines with presumed pre-existing infection exacerbated by new wound-site gall formation. Crown gall symptom quality distribution showed concentration at the trunk-soil interface (45% of galls) where freeze damage was most severe and temperature fluctuation at the snow line created repeated freeze-thaw injury, with additional galls at cordon renewal sites (30%) and scaffold positions (25%) reflecting the wound distribution pattern. Gall impact assessment showed that vines with galls exceeding 50% trunk circumference exhibited reduced vigor, chlorotic canopy, and 40% yield reduction compared to ungalled vines, while complete girdling by gall tissue resulted in vine death in 8% of affected vines. The crown gall symptom quality assessment established that freeze damage served as the primary predisposing factor, with A. vitis infection converting temporary freeze injury into progressive structural disease that would continue to impact vineyard productivity for years beyond the initial freeze event.
Crown gall symptom quality is a bacterial infection indicator quality that inheres in grapevine organisms infected by Agrobacterium vitis and represents the characteristic tumorous overgrowths (galls) developing at wound sites where the bacterium has transferred its tumor-inducing T-DNA into host plant cells, with galls typically forming at the crown region near the soil line, at graft unions, and at freeze-damaged sites along trunks and cordons, progressively enlarging over successive growing seasons and eventually girdling vascular tissue to cause vine decline and death. This quality captures the diagnostic features of the most prevalent bacterial disease of grapevine in cold-climate regions, recognizing that crown gall is intimately associated with freeze injury that creates the wound sites required for infection, establishing the disease as a secondary consequence of winter damage that compounds the direct effects of cold injury on vineyard productivity and longevity. The ontological significance of crown gall symptom quality lies in its role enabling diagnosis of a disease with no curative treatment, where symptom recognition informs decisions about vine retention versus removal and about preventive measures including wound protection and rootstock selection that may reduce future crown gall establishment. The biological basis of crown gall symptoms encompasses the unique pathogenesis mechanism of A. vitis in which bacterial cells at wound sites transfer a segment of DNA (T-DNA) from their tumor-inducing plasmid into plant cells, with the integrated T-DNA expressing genes for auxin and cytokinin biosynthesis that reprogram plant cells for uncontrolled proliferation creating the tumorous gall tissue. The systemic nature of A. vitis in grapevine means that the bacterium may be present throughout vine tissues including apparently healthy portions distant from visible galls, with propagation from infected mother vines representing a major pathway for disease spread to new plantings. The gall tissue itself is initially soft and fleshy, becoming corky and irregular as it ages, with secondary infection by wood-rotting fungi and insects often occurring in older galls, complicating the primary symptom picture while creating additional entry points for other pathogens.
Crown_Gall_Symptom_Quality_(Agrobacterium_vitis) is_a Bacterial_Infection_Indicator_Quality, representing the tumorous gall symptoms caused by Agrobacterium vitis T-DNA transfer to plant cells. The tumor formation at wound sites is pathognomonic. Strong association with freeze injury in cold climates creates disease clustering following cold winters. Systemic bacterial presence in vine tissues enables spread through propagation material. No curative treatment exists management focuses on prevention and vine removal. Crown/soil-line location most common, reflecting freeze damage and soil inoculum distribution.
Agrobacterium gall symptoms
Agrobakterium-Symptome
Mauke-Symptome
Rebmauke-Zeichen
bacterial gall indicators
bakterielle Gallen-Indikatoren
crown gall symptoms
grapevine gall disease
Burr TJ, Otten L (1999) Crown gall of grape: biology and disease management. Annual Review of Phytopathology 37:53-80 Eastwell KC et al. (2006) Agrobacterium vitis and crown gall of grape. In: Bentley S, Jackson RW (eds) Crown Gall of Stone Fruit, Walnut and Grape. APS Press Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests, 2nd Edition. APS Press
Crown Gall Symptom Quality (Agrobacterium vitis)
Crown regeneration enables vine survival of catastrophic events that destroy the entire above-ground structure, with regeneration source depending on damage extent and vine type. Severe spring frost (-8 to -12°C on green tissue) killing all primary shoots triggers regeneration from secondary and tertiary buds, restoring partial canopy within 3-6 weeks but typically reducing yield by 50-80% in the current season. Complete crown loss (trunk kill) initiates regeneration from basal latent buds (own-rooted vines) or necessitates regrafting (grafted vines), with full canopy restoration requiring 2-3 growing seasons. Fire-damaged vines show crown regeneration from surviving cambial tissue or basal buds if heat exposure duration was limited, with recovery success inversely related to trunk char depth. Cultivars with extensive latent bud populations persisting on old wood (Grenache, Mourvèdre, Carignan) show superior crown regeneration compared to types with limited latent bud longevity. The regenerated crown initially shows juvenile growth characteristics (high vigor, low fruitfulness) transitioning to mature production patterns over 2-4 seasons.
A recovery disposition that is the inherited capacity of a Vitis vinifera plant to regenerate a complete functional canopy (crown) from latent buds, adventitious meristems, or the root system following total or near-total loss of above-ground vegetative structures due to frost, fire, mechanical damage, or severe disease, re-establishing photosynthetic capacity and reproductive potential.
Crown regeneration represents the ultimate survival mechanism enabling grapevine persistence through catastrophic above-ground destruction. The trait depends on maintenance of viable meristematic tissue (latent buds on surviving trunk/root crown, root cambium capable of adventitious bud formation) and adequate carbohydrate reserves to fuel initial regrowth before photosynthetic independence. Crown regeneration capacity has historically enabled vineyard recovery from fire, extreme frost, and war damage, preserving genetic resources and established plantings. Climate change increasing frost and fire risk elevates the importance of crown regeneration capacity in cultivar and site resilience assessment.
Kronenregeneration
Laubwandwiederherstellung
above-ground restoration
canopy regeneration
oberirdische Regeneration
shoot system recovery
Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp.: 80-112 Keller M, Mills LJ (2007) Effect of pruning on recovery and productivity of cold-injured Merlot grapevines. American Journal of Enology and Viticulture 58:351-357
Crown Regeneration Disposition
The cru name Chambertin Grand Cru designates a 12.9-hectare vineyard in the commune of Gevrey-Chambertin classified since the original 1861 Lavalle classification and confirmed in the 1936 AOC decree — in contrast, the cru name Château Mouton Rothschild Premier Grand Cru Classé designates an estate (not a specific parcel) classified in the 1855 Bordeaux Classification (elevated from Deuxième to Premier Cru in 1973), while the cru name Morgon designates an entire commune of 1,100 hectares classified as one of Beaujolais's 10 cru villages — illustrating how the term cru operates at fundamentally different geographic scales (parcel, estate, village) and with different classificatory implications across French wine regions.
A vineyard designation within the French wine classification system that identifies a site, estate, or vineyard unit that has been formally classified or traditionally recognized as a cru — a term denoting a specific growth or production site of acknowledged quality, used across French wine regions with varying regulatory definitions ranging from classified vineyard parcels (Burgundy's Grand Cru and Premier Cru) through classified estates (Bordeaux's Cru Classé) to classified villages (Beaujolais Cru).
Cru_Name captures the most complex and historically layered vineyard classification concept in wine. The term cru (from the verb croître, to grow) carries distinct regulatory and cultural meanings in different French regions: in Burgundy it designates officially classified vineyard parcels organized in a two-tier hierarchy (Premier Cru and Grand Cru) based on terroir quality in Bordeaux it designates classified estates (châteaux) organized in historically fixed hierarchies (1855 Classification, Saint-Émilion Classification) in Beaujolais and the Loire it designates classified villages and in Champagne it historically classified entire communes on a percentage scale (échelle des crus, 80-100%). The common thread across these diverse applications is the recognition of differential quality associated with specific sites, estates, or villages — but the unit of classification, the criteria for inclusion, and the regulatory consequences vary fundamentally between regions.
Cru-Bezeichnung
Lage-Klassifikation
classified growth name
cru classé designation
cru designation
klassifiziertes Gewächs
Lavalle J (1855) Histoire et statistique de la vigne et des grands vins de la Côte d'Or INAO (2023) Classification des crus Pitiot S and Servant JC (2010) Les Vins de Bourgogne 14th Ed, Collection Pierre Poupon Markham D (1998) 1855: A History of the Bordeaux Classification, Wiley
Cru Name
Cryoprotection quality profiling during cold acclimation in Cabernet Franc characterized the accumulation of protective compounds underlying the seasonal development of freezing tolerance. Metabolomic and proteomic analysis at monthly intervals from September through January tracked the progression of cryoprotection quality development. Total soluble sugar concentration in bud tissues increased from 45 mg/g dry weight in September to 180 mg/g in January, with the 4-fold increase dominated by raffinose family oligosaccharides (RFOs) recognized for their membrane-stabilizing properties increasing from 5 to 65 mg/g. Proline concentration showed 8-fold increase from 2 to 16 mg/g, contributing both osmotic protection and protein stabilization functions. Dehydrin protein accumulation began in October and reached maximum in December with 15-fold increase over summer baseline levels, providing protein-protective and membrane-stabilizing functions. Membrane lipid analysis revealed the fatty acid desaturation index increasing from 1.2 to 1.8 through the acclimation period, reflecting increased unsaturated fatty acid content that maintains membrane fluidity at low temperatures. The temporal correlation between cryoprotection quality development and freezing tolerance improvement (r²=0.91) established that cryoprotectant accumulation directly underlies hardiness acquisition, with the metabolite and protein profiles providing molecular markers for acclimation status assessment as an alternative to destructive freezing tests.
Cryoprotection quality is a cold stress indicator quality that inheres in a grapevine organism and represents the capacity to accumulate and deploy cellular compounds and structural modifications that protect against freezing damage, including the synthesis of compatible solutes that stabilize membranes and proteins during freeze-induced dehydration, the expression of protective proteins that prevent ice crystal damage, and the modification of membrane lipid composition to maintain fluidity and function at low temperatures. This quality captures the molecular and biochemical mechanisms underlying cold hardiness, recognizing that the survival of plant cells during freezing ultimately depends on protective molecules that enable cellular structures to withstand the physical and chemical stresses of ice formation in or around them. The ontological significance of cryoprotection quality lies in its role as the mechanistic foundation for cold hardiness, with the accumulation of cryoprotective compounds directly enabling the survival of temperatures that would be lethal in their absence, and with variation in cryoprotection capacity explaining much of the genotypic differences in freezing tolerance observed at the whole-plant level. The biological basis of cryoprotection encompasses multiple protective mechanisms including osmotic adjustment through compatible solute accumulation that reduces the extent of cellular dehydration during extracellular ice formation, the stabilization of membrane structure by specific sugars and proteins that substitute for water in maintaining lipid bilayer organization, the prevention of protein denaturation by molecular chaperones and compatible solutes that preserve protein conformation during dehydration, and the physical modification of membrane lipid fatty acid composition to prevent damaging phase transitions at low temperatures. The coordination among cryoprotective mechanisms creates a comprehensive defense system where each component addresses specific aspects of freezing stress, with the integrated function of all components required for maximum protection and with deficiencies in any single mechanism potentially limiting overall hardiness. The seasonal dynamics of cryoprotection parallel cold acclimation, with cryoprotectant accumulation triggered by acclimation-inducing conditions and reaching maximum levels at full acclimation, followed by decline during deacclimation as protective compounds are catabolized or diluted by resumed growth.
Cryoprotection_Quality is_a Cold_Stress_Indicator_Quality, serving as parent class for Sugar_Cryoprotection_Quality, Protein_Cryoprotection_Quality, and Membrane_Lipid_Adjustment_Quality which address specific protective mechanisms. This class addresses the overall molecular basis of freezing protection while child classes capture particular mechanisms. The correlation between cryoprotectant accumulation and hardiness enables molecular markers for acclimation status. Multiple mechanisms provide complementary protection against different aspects of freezing stress. The seasonal dynamics parallel cold acclimation with accumulation during hardening and depletion during deacclimation.
Frostschutzkapazität
Gefrierschutzkapazität
Kryoprotektion
Kälteschutzmechanismen
antifreeze capacity
cold protection mechanisms
cryoprotective capacity
freezing protection
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Xin Z, Browse J (2000) Cold comfort farm: the acclimation of plants to freezing temperatures. Plant, Cell and Environment 23:893-902 Thomashow MF (1999) Plant cold acclimation: freezing tolerance genes and regulatory mechanisms. Annual Review of Plant Physiology and Plant Molecular Biology 50:571-599
Cryoprotection Quality
Shiraz CSIRO selection bears a CSIRO_Clone_Role, virus-eliminated material from traditional Australian Shiraz subjected to CSIRO thermotherapy and tissue culture protocols to produce certified clean stock for the national vine improvement scheme. Sultana (Thompson Seedless) CSIRO clone serves in a CSIRO_Clone_Role for Australian dried fruit and table grape production, with CSIRO sanitation ensuring pathogen-free material for the extensive Murray-Darling plantings. Chardonnay material imported through CSIRO quarantine bears a CSIRO_Clone_Role, international selections processed through CSIRO protocols to meet Australian phytosanitary requirements before release to the domestic industry.
An Australian clone role that inheres specifically in clones selected, evaluated, or processed through CSIRO (Commonwealth Scientific and Industrial Research Organisation) grapevine improvement programs, Australia's premier national research agency contributing virus elimination technology, clone evaluation, and variety development to Australian viticulture. CSIRO clones have typically undergone rigorous pathogen elimination (thermotherapy, tissue culture) and comprehensive testing, with the organization's scientific expertise ensuring high health standards for released material. CSIRO's viticultural research (historically based at Merbein, Victoria, now consolidated within broader programs) developed foundational technologies for Australian vine improvement, including early virus detection methods, elimination protocols, and clonal evaluation systems. CSIRO-processed clones may include both Australian selections (subjected to CSIRO sanitation) and international imports (processed through CSIRO quarantine protocols), with the CSIRO designation indicating institutional handling rather than necessarily Australian origin. The organization's research programs extend beyond clone selection to variety breeding, rootstock evaluation, and viticultural management systems.
CSIRO_Clone_Role identifies material processed through Australia's premier research institution, carrying quality assurance reflecting CSIRO's scientific standards and technical capabilities. CSIRO's role in Australian viticulture extends beyond clone selection to fundamental research underpinning the industry—irrigation science, pest management, wine chemistry—with clone programs representing one dimension of comprehensive viticultural research. The organization's evolution (consolidation of historically separate stations, changing research priorities, funding fluctuations) has affected program continuity, with some clone selection functions transferred to state agencies or industry bodies while CSIRO maintains research leadership. CSIRO-processed material commands confidence in health status, with the organization's virus elimination and testing protocols providing documented assurance valued by Australian nurseries and growers.
Australian research clone
CSIRO clone
CSIRO selection
CSIRO-Klon
CSIRO-Selektion
Merbein clone
Merbein-Klon
CSIRO (2020) Grapevine improvement research programs Dry PR, Coombe BG (2004) Viticulture Volume 1, Winetitles Constable F et al. (2012) Grapevine leafroll-associated virus 3 in Australia, Australasian Plant Pathology 41: 313-322 Habili N, Nutter FW (1997) Temporal and spatial analysis of grapevine leafroll disease, Phytopathology 87: 799-804
CSIRO Clone Role
The cultivar known internationally as Pinot Noir carries a complex web of cultivar designations including the OIV-registered prime name Pinot Noir (OIV variety code 08711), VIVC prime name Pinot Noir (VIVC number 09279), 285 documented synonyms including Spätburgunder (German), Blauburgunder (Austrian/Swiss), Pinot Nero (Italian), and Kisburgundi (Hungarian), historical names such as Morillon Noir (14th century) and Noirien (medieval Côte d'Or), and clone-specific codes such as ENTAV-INRA 115, 667, 777 and Geisenheim clones Gm 20-13 and Gm 2-47.
A grapevine information entity that serves as a naming or labeling expression assigned to a grapevine cultivar, clone, or selection for the purpose of identification, communication, regulatory registration, or commercial differentiation, encompassing all categories of names, codes, numbers, and trade designations used to refer to distinct grapevine genotypes.
Cultivar_Designation is the umbrella class for all naming and coding systems applied to grapevine genotypes. The complexity of grapevine nomenclature is exceptional among crop plants: the VIVC documents over 24,000 cultivar names referring to approximately 6,000-8,000 genetically distinct cultivars, with individual cultivars carrying anywhere from 1 to 300+ designations across languages, regions, historical periods, and regulatory jurisdictions. Subclasses organize this nomenclatural complexity into prime names, synonyms, historical names, regional names, breeding codes, and trade names.
Rebsortenname
Sortenbezeichnung
Sortenkennung
cultivar name
grapevine naming entity
variety designation
Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157 ICNCP (2016) International Code of Nomenclature for Cultivated Plants 9th Ed, ISHS OIV (2013) OIV List of Vine Varieties and Their Synonyms
Cultivar Designation
In Muscat Ottonel berries in Burgenland, Austria during BBCH 85-89, cultivar specific flavor develops through variety-characteristic accumulation of linalool to 1.8 mg/L and geraniol to 1.2 mg/L providing the distinctive Muscat floral character, with linalool-to-geraniol ratio of 1.5 typical for this variety, concurrent low methoxypyrazine of 3 ng/L distinguishing from Sauvignon-type varieties, and trained panel varietal typicity rating of 8.9 on 10-point scale confirming achievement of characteristic Muscat Ottonel expression under warm Pannonian conditions averaging 24°C.
Cultivar specific flavor describes the genetically determined aromatic signature characteristic of individual Vitis vinifera varieties, wherein variety-specific expression of flavor biosynthetic enzymes produces distinctive compound profiles that enable varietal identification and define the typicity expectations against which grape and wine quality are assessed throughout the 35-50 day post-veraison period culminating in harvest-maturity expression of characteristic varietal identity. This phenomenon reflects the genetic architecture of flavor biosynthesis with variety-specific alleles of terpene synthases, O-methyltransferases, carotenoid cleavage dioxygenases, and related enzymes determining which compound classes accumulate to sensory-relevant concentrations, producing the floral-Muscat character of aromatic varieties through high monoterpene synthase expression, the herbaceous-tropical profile of Sauvignon Blanc through methoxypyrazine and thiol pathway activities, and the subtle stone-fruit nuances of Chardonnay through specific norisoprenoid accumulation patterns. The molecular basis of cultivar specific flavor involves transcriptional regulation through variety-specific promoter sequences and transcription factor expression, structural variation in enzyme-coding genes affecting catalytic activities and substrate specificities, and copy number variation in gene families that influences total biosynthetic capacity for individual compound classes. Sensory recognition of cultivar specific flavor enables variety identification through trained panel assessment and supports appellation regulations requiring varietal labeling accuracy, with experienced tasters achieving 70-90% correct identification rates for major varieties based on aromatic character alone, demonstrating the distinctiveness and reproducibility of varietal flavor signatures. Environmental modulation of cultivar specific flavor occurs through terroir effects that enhance, diminish, or modify the genetically-determined baseline profile, with certain sites and vintages producing particularly intense or characteristic varietal expression while others may yield atypical profiles requiring blending or stylistic adjustment to meet consumer expectations. The preservation of cultivar specific flavor through winemaking represents a primary quality objective in varietal wine production, with gentle processing, appropriate yeast selection, and careful oxidation management employed to maintain the distinctive aromatic signatures established during grape development rather than introducing winemaking-derived flavors that mask varietal character.
Cultivar_Specific_Flavor is subclass of Flavor_Profile_Maturation representing genetically-determined varietal character. Distinguishes from Mature_Flavor_Expression which describes general maturity. Sibling to Mature_Flavor_Expression and Optimal_Flavor_Development. Determined by variety-specific enzyme expression. Enables varietal identification. Typicity criterion for quality assessment. Modified but not determined by environment.
Kultivar-aromatische-Identität
Rebsorten-Aroma-Charakter
cultivar aromatic identity
genetic flavor signature
genetische Aroma-Signatur
sortenspezifischer Geschmack
varietal flavor character
variety-specific aroma
Martin DM et al (2010) Functional annotation of the grapevine terpene synthase gene family. BMC Plant Biology 10: 226 Ebeler SE and Thorngate JH (2009) Wine chemistry and flavor. Journal of Agricultural and Food Chemistry 57: 8098-8108 Robinson AL et al (2014) Origins of grape and wine aroma. American Journal of Enology and Viticulture 65: 1-30
Cultivar Specific Flavor
Cultivar synchrony assessment across multiple Merlot blocks on an estate reveals the site modulation of varietal phenology, with four blocks spanning elevation range of 150-280 meters showing véraison dates spreading 12 days despite identical genetics (clone 181 on 101-14 Mgt), with lowland blocks (150-180m) reaching véraison August 8-10 and upland blocks (250-280m) reaching véraison August 18-20, demonstrating that cultivar synchrony is moderate (all blocks following expected Merlot timing relative to other varieties) but site effects create sufficient spread to require block-specific harvest timing rather than cultivar-based scheduling.
Cultivar synchrony quality is an inter-plant synchrony quality that represents the developmental uniformity among vines of the same cultivar across a vineyard or estate, characterizing the inherent phenological coherence of varietal plantings that determines whether the cultivar can be managed and harvested as a coordinated unit or whether site and management variation creates developmental heterogeneity requiring block-specific treatment timing. Within the BFO framework, this quality specifically addresses the cultivar-level dimension of inter-plant synchrony, distinguishing varietal phenological coherence from block-specific or vine-specific synchrony while recognizing that cultivar synchrony indicates the degree to which genetic uniformity translates to phenological uniformity when the same variety is planted across different vineyard conditions. The ontological significance lies in cultivar synchrony determining the feasibility of cultivar-based harvest scheduling and the predictability of varietal phenology across sites, where high cultivar synchrony enables estate-wide or regional harvest planning based on variety (scheduling Chardonnay before Cabernet Sauvignon regardless of block location), while low cultivar synchrony indicates that site effects overwhelm genetic tendency, requiring block-specific monitoring and potentially reordering harvest sequence based on actual rather than expected phenology. The factors affecting cultivar synchrony include the inherent phenological plasticity of the variety (some cultivars showing stable phenology across environments, others responding more variably to site differences), the range of environmental conditions across which the cultivar is planted (broader environmental range typically reducing synchrony), and the management uniformity applied to the cultivar across sites (consistent practices promoting more similar development). The clone and rootstock effects within cultivars can substantially affect synchrony, as different clones of the same variety may show significant phenological differences, and rootstock effects on vigor and phenology can modify cultivar timing, such that "cultivar synchrony" in practice may require consideration of specific clone-rootstock combinations rather than variety alone. The operational value of cultivar synchrony includes harvest logistics planning (sequencing varieties through the winery based on expected maturity order), labor allocation (anticipating harvest timing for different varieties), and winemaking preparation (timing of receiving capacity and processing equipment for expected varietal deliveries). Understanding cultivar synchrony quality enables realistic assessment of whether varietal generalizations apply across diverse plantings, appropriate harvest sequence planning based on cultivar phenology patterns, and recognition of when site effects require block-specific rather than variety-based scheduling.
Cultivar_Synchrony_Quality addresses the phenological uniformity among vines of the same variety across different sites and conditions. This quality indicates whether genetic identity translates to phenological uniformity or is modified by environmental factors. The class supports variety-based planning through cultivar synchrony characterization across diverse plantings.
Rebsortenkohärenz
Sortensynchronie
Sortenuniformität
cultivar coherence
genetic developmental uniformity
genetische Entwicklungsuniformität
varietal uniformity
variety-level synchrony
Jones GV, Davis RE (2000) Climate influences on grapevine phenology, grape composition, and wine production and quality for Bordeaux, France. American Journal of Enology and Viticulture 51(3): 249-261 Parker AK et al. (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17(2): 206-216
Cultivar Synchrony Quality
In Gewürztraminer berries in Tramin, Alto Adige during BBCH 85-89, cultivar typical aroma develops through characteristic accumulation of linalool to 2.4 mg/L and distinctive cis-rose oxide to 85 µg/L providing the variety-defining lychee-rose character, with geraniol at 0.9 mg/L and citronellol at 0.4 mg/L contributing secondary floral complexity, trained panel varietal typicity rating of 9.2 on 10-point scale confirming achievement of characteristic Gewürztraminer aromatic expression recognizable by experienced tasters despite the warm vintage conditions averaging 25°C.
Cultivar typical aroma describes the genetically determined olfactory signature characteristic of individual Vitis vinifera varieties, wherein variety-specific expression of aromatic compound biosynthetic enzymes produces distinctive volatile profiles that enable varietal identification through scent alone and define the typicity expectations against which grape and wine aromatic quality are assessed, establishing the fundamental aromatic identity that distinguishes each cultivar throughout the 35-50 day post-veraison period culminating in harvest-maturity expression of characteristic varietal scent. This phenomenon reflects the genetic architecture of aromatic biosynthesis with variety-specific alleles of terpene synthases determining monoterpene profiles, unique expression patterns of O-methyltransferases establishing methoxypyrazine signatures in Sauvignon and Cabernet family varieties, variety-specific thiol precursor synthesis capacity creating tropical fruit potential in Sauvignon Blanc, and cultivar-distinctive carotenoid cleavage patterns generating characteristic norisoprenoid contributions to varietal complexity. The molecular basis of cultivar typical aroma involves transcriptional regulation through variety-specific promoter sequences controlling biosynthetic gene expression, structural variation in enzyme-coding genes affecting substrate specificity and product profiles, gene copy number variation influencing total biosynthetic capacity, and post-transcriptional regulatory mechanisms fine-tuning pathway outputs to establish the quantitative compound ratios that define varietal aromatic signatures. Sensory recognition of cultivar typical aroma enables variety identification through trained panel assessment and blind tasting exercises, with experienced evaluators achieving 75-95% correct identification rates for major aromatic varieties based on olfactory character alone, demonstrating the distinctiveness and reproducibility of varietal scent signatures despite substantial environmental and vintage variation that modifies but does not fundamentally alter genetic aromatic identity. The specific compounds defining cultivar typical aroma vary among varieties, with Muscat characterized by elevated linalool and geraniol providing floral-grape character, Gewürztraminer showing high linalool with distinctive rose oxide contribution, Riesling demonstrating balanced terpene profile with characteristic TDN petrol note development potential, and Sauvignon Blanc defined by methoxypyrazine herbaceous character combined with thiol-derived tropical fruit potential upon fermentation. Environmental and viticultural factors modulate but do not fundamentally determine cultivar typical aroma, with terroir effects capable of enhancing, diminishing, or subtly shifting the genetically-established baseline profile, while the essential varietal character remains recognizable across diverse growing conditions and vintage years.
Cultivar_Typical_Aroma is subclass of Aroma_Profile_Maturation representing genetically-determined varietal olfactory character. Distinguishes from Mature_Aroma_Expression which describes general maturity. Sibling to Mature_Aroma_Expression and Optimal_Aroma_Development. Determined by variety-specific enzyme expression. Enables varietal identification. Typicity criterion for quality assessment. Modified but not determined by environment.
Kultivar-olfaktorische-Identität
Rebsorten-Duft-Charakter
cultivar olfactory identity
genetic aroma signature
sortentypisches Aroma
varietal aromatic character
variety-specific scent
Martin DM et al (2010) Functional annotation of the grapevine terpene synthase gene family. BMC Plant Biology 10: 226 Robinson AL et al (2014) Origins of grape and wine aroma. American Journal of Enology and Viticulture 65: 1-30 Ebeler SE and Thorngate JH (2009) Wine chemistry and flavor. Journal of Agricultural and Food Chemistry 57: 8098-8108
Cultivar Typical Aroma
In Grüner Veltliner vines in the Wachau, Austria, cumulative cold exposure builds through winter with total hours below 7.2°C reaching approximately 300 hours by late December, 700 hours by late January, and 1100 hours by late February, satisfying the cultivar requirement of approximately 1000-1100 hours, with this cumulative exposure achieved despite intermittent warm periods in January that temporarily interrupted but did not substantially reverse the dormancy release progress.
Cumulative cold exposure in Vitis vinifera represents the total physiologically effective low temperature experience accumulated by dormant buds during winter, encompassing all cold exposure that contributes to the progressive satisfaction of chilling requirements and dormancy release regardless of the specific quantification model employed. This concept emphasizes the integrative nature of dormancy release as a process responding to total cold exposure over time rather than to any single temperature event, with buds effectively summing their cold experience from dormancy establishment through eventual dormancy release. The physiological basis of cumulative cold exposure effects involves temperature-sensitive processes that gradually modify dormancy-maintaining mechanisms, with low temperatures activating specific enzymatic and regulatory processes that are largely inactive at warmer temperatures, allowing these dormancy-releasing reactions to proceed only during cold periods. The cellular processes activated by cold exposure include increased expression and activity of ABA catabolic enzymes, chromatin remodeling complexes that remove repressive modifications from growth-related genes, and signaling pathways that downregulate dormancy-associated MADS-box gene expression. The cumulative nature of cold exposure effects reflects the persistence of physiological changes induced during each cold period, with modifications to ABA levels, chromatin state, and gene expression being maintained during subsequent warmer periods rather than immediately reversing, allowing the effects of multiple cold exposures to accumulate toward dormancy release. The effectiveness of cold exposure for dormancy release varies with temperature, with moderately cold temperatures in the range of 0-7°C generally most effective, very cold temperatures below -5°C being less effective or ineffective, and the integration of variable temperatures into effective exposure being addressed by various chilling models. The relationship between cumulative cold exposure and dormancy status is fundamentally quantitative, with insufficient exposure resulting in incomplete dormancy release that manifests as delayed, prolonged, or uneven budbreak, while adequate exposure ensures synchronous and timely spring growth resumption. Climate variability creates year-to-year variation in cumulative cold exposure that affects dormancy release timing and completeness, with warm winters potentially providing insufficient exposure for adequate dormancy release in some cultivar-region combinations.
Cumulative_Cold_Exposure represents the physiological integration aspect of Chilling_Accumulation emphasizing total effective cold experience. Distinguished from Progressive_Chilling_Units which addresses model-based quantification, while Cumulative_Cold_Exposure emphasizes the underlying physiological integration. Reflects the persistence of cold-induced changes allowing accumulation across variable temperature periods. Effective temperature range approximately 0-7°C. Total exposure requirements vary from approximately 400-1500 hours depending on cultivar.
Gesamtkälteeinwirkung
Kältesummen-Exposition
accumulated cold
akkumulierte Kälte
integrated cold experience
kumulative Kälteexposition
total cold exposure
winter cold sum
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Dokoozlian NK (1999) Chilling temperature and duration interact on the budbreak of Perlette grapevine cuttings. HortScience 34: 1054-1056 Campoy JA et al (2011) High temperatures and time to budbreak in low chill apricot. Environmental and Experimental Botany 73: 134-142
Cumulative Cold Exposure
Black Corinth (Zante Currant, Korinthiaki, Staphis) bears a Currant_Production_Scion_Role as the definitive currant variety cultivated for over 3,000 years in Greece, with tiny parthenocarpic berries (0.5-0.8 g fresh weight) containing no seeds and drying rapidly (7-10 days sun-drying) to produce intensely flavored dried currants (0.15-0.25 g) essential for traditional baking—fruitcakes, hot cross buns, spotted dick, mincemeat. Carina serves in a Currant_Production_Scion_Role as an Australian-bred currant variety, developed from Black Corinth parentage with improved productivity and disease resistance while maintaining the small berry size and intense flavor required for currant production. White Corinth (historically) bore a Currant_Production_Scion_Role producing light-colored currants, though it is now rarely cultivated commercially.
A raisin production scion role that inheres in a grafted scion cultivar by virtue of its production of very small, seedless berries suitable for drying into currants—the smallest category of dried grapes, prized for baking applications where their diminutive size enables even distribution through doughs and batters. Currant production scions are defined by tiny berry size (typically 0.3-1.0 g fresh weight, drying to 0.1-0.3 g), complete seedlessness achieved through parthenocarpy (fruit development without fertilization), and intense flavor development during drying that compensates for minimal individual berry mass. The name "currant" derives from "Corinth" (Greece), reflecting the historical production center for Zante currants (Black Corinth variety) that defined the category. Dried currants differ botanically from the unrelated Ribes currants (blackcurrant, redcurrant) which are true berries of a different plant family. Currant production is concentrated in Greece (Corinth region, Peloponnese), Australia, South Africa, and California, with significantly smaller volumes than standard raisin production.
Currant_Production_Scion_Role captures a specialized niche within dried grape production defined by extremely small berry size and complete seedlessness through parthenocarpy—contrasting with the stenospermocarpic seedlessness of Thompson Seedless which produces larger berries with soft seed traces. The parthenocarpic nature of currant varieties means berry development occurs without pollination, resulting in inherently small size that breeding cannot easily overcome without losing the seedless character. Currant production volumes are minor compared to raisins (less than 5% of dried grape trade) but the product commands significant price premiums for specialty baking applications. Greek currant production faces challenges including aging vineyards, rural depopulation, and competition from Australian expansion.
Corinthian grape
Korinthen-Traube
Korinthtraube
Zante currant grape
Zante-Traube
currant grape
drying currant variety
Christensen LP (2000) Raisin Production Manual, UC ANR Publication 3393 Creasy GL, Creasy LL (2009) Grapes, CABI Ledbetter CA, Ramming DW (1989) Seedlessness in grapes, Horticultural Reviews 11: 159-184 Dry PR, Gregory GR (1988) Grapevine varieties, in Viticulture Vol 1 Resources, Winetitles pp.: 119-138
Currant Production Scion Role
A curved simple unbranched tendril at node 10 on a laterally oriented shoot of Vitis vinifera cv. Pinot Noir in Burgundy, France, measuring 17 cm arc length with approximately 45-degree total curvature creating a gentle C-shaped trajectory concave toward gravitational vertical, showing diameter variation from 3.2 mm near base to 2.6 mm near apex, asymmetric tissue development evident in cross-section with thicker cell walls on convex side indicating differential growth driving curvature, exhibiting modified circumnutation pattern where curved geometry creates three-dimensional spiral searching motion rather than planar sweeping, making contact with a horizontal trellis wire through curved approach facilitating rapid initial contact and coiling initiation, occurring on a shoot oriented 45 degrees from vertical where gravitropic and phototropic responses combine to induce tendril curvature, measured at BBCH stage 75 (berries pea-sized) in late June, representing the curved architecture common in Pinot Noir under non-vertical shoot orientations where 40-55% of unbranched tendrils show pronounced curvature.
A simple unbranched tendril exhibiting pronounced curvature in its relaxed non-coiled state in Vitis vinifera, characterized by tendril axis showing greater than 20-30 degrees deviation from a straight line between shoot attachment point and apex, often displaying S-shaped, C-shaped, or more complex curved trajectories resulting from differential growth patterns on opposite sides of the axis, variable diameter along length with potential tapering or localized thickening at curvature zones, and geometric properties creating three-dimensional exploration patterns during circumnutation that differ from straight tendril sweeping motions. Curved simple unbranched tendrils develop through differential growth rates on opposite sides of the developing axis driven by various factors including asymmetric light exposure creating phototropic curvature, gravitropic responses to shoot orientation inducing curvature toward or away from gravitational vector, mechanical perturbations from wind or touching creating thigmotropic responses, intrinsic developmental programs in some cultivars favoring curved rather than straight growth trajectories, or combinations of these factors operating during the elongation phase. The functional implications of curved architecture include modified spatial exploration patterns with curved geometry potentially enhancing probability of contact with three-dimensionally distributed supports in natural climbing contexts, reduced maximum reach distance compared to straight tendrils of equivalent length but potentially enhanced interaction with nearby supports through curved approach angles, possible increased flexibility facilitating coiling initiation and contact maintenance during wind-induced shoot movements, and potentially altered mechanical properties with curved geometry creating complex stress distributions under loading.
Curved simple unbranched tendrils are distinguished from straight simple unbranched tendrils by >20-30 degrees deviation from linear geometry in relaxed pre-coiling state, and from coiled tendrils by curvature representing growth trajectory rather than thigmotropic coiling response. Curvature degree, direction, and shape (C-curve, S-curve, complex) provide diagnostic information about causative factors, with consistent curvature direction across tendrils suggesting environmental drivers (light, gravity) while variable curvature patterns indicate intrinsic developmental variation or localized perturbations. Some cultivars show genetic tendencies toward curved rather than straight development, while others show facultative responses where curvature depends on shoot orientation, light conditions, and mechanical environment. Functionally, moderate curvature may enhance climbing in complex three-dimensional support structures characteristic of natural climbing contexts, while excessive curvature reduces reach distance and may compromise exploration efficiency in planar trellis systems typical of commercial viticulture.
bent unbranched tendril
curved-axis simple tendril
gebogene einfache Ranke
gekrümmte unverzweigte Ranke
nicht-lineare einzelne Kletterorgan
non-linear single tendril
Based on Darwin (1875) "The Movements and Habits of Climbing Plants" John Murray, London Jaffe & Galston (1968) "The physiology of tendrils" Annual Review of Plant Physiology 19=417-434 Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 136-150 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 173-194.
Curved Simple Unbranched Tendril
Detailed characterization of Cuticle_Based_Resistance_Disposition at INRA Montpellier examined cuticular properties across 12 cultivars using scanning electron microscopy of epicuticular wax morphology, gas chromatography-mass spectrometry of wax composition, and atomic force microscopy of surface nanomechanics. 'Grenache' exhibited dense epicuticular wax coverage (42 μg/cm²) with elaborate platelet crystals creating a rough, hydrophobic surface (water contact angle 142°) on which B. cinerea conidia showed 62% germination rate with delayed germ tube emergence (mean 18 hours). 'Chasselas' displayed sparse, amorphous wax (18 μg/cm²), smooth surface topography, and lower hydrophobicity (contact angle 98°), supporting 89% germination with rapid germ tube emergence (mean 8 hours). Controlled inoculation of surface-sterilized berries demonstrated that 'Grenache' developed infection in 24% of berries versus 58% in 'Chasselas' when inoculated with equivalent conidial suspensions onto intact surfaces. However, when cuticle was enzymatically removed with cutinase pre-treatment, infection rates equalized at 82-88%, confirming that differences in susceptibility were attributable to Cuticle_Based_Resistance_Disposition rather than post-penetration defense mechanisms.
A Cuticle_Based_Resistance_Disposition is a Botrytis_Resistance_Disposition that inheres in a Vitis vinifera berry and is realized through characteristics of the cuticular membrane that impede Botrytis cinerea conidial germination, germ tube penetration, and appressorium formation on the berry surface. The cuticle, comprising a cutin polymer matrix embedded with intracuticular waxes and covered by epicuticular wax deposits, serves as the outermost barrier between the berry and its environment, and its properties profoundly influence susceptibility to B. cinerea infection of intact fruit. Cuticle-based resistance mechanisms include cuticle thickness providing a physical barrier to direct penetration, wax crystal morphology and density affecting water retention and conidial adhesion, wax chemical composition influencing surface hydrophobicity and germination cue availability, and cuticle integrity (absence of microcracks and fissures) preventing pathogen access to underlying tissues. Epicuticular wax crystal structure varies among cultivars from amorphous films to elaborate crystalline plates, needles, or tubules, with potential effects on spore adhesion and germination microenvironment. Cuticle properties change during berry development, with cuticle deposition active during early development but often failing to keep pace with berry expansion during ripening, leading to cuticle thinning and microcracking that increases susceptibility. Cuticular characteristics are under genetic control with moderate heritability and represent selectable traits for breeding, though their contributions to field resistance are difficult to isolate from other resistance mechanisms.
Cuticle_Based_Resistance_Disposition is_a Botrytis_Resistance_Disposition. This class specifically addresses the outermost cuticular layer including epicuticular and intracuticular waxes, distinct from Thick_Skin_Disposition which encompasses overall exocarp mechanical properties. Cuticle properties affect the earliest infection stages including conidial adhesion, germination, and initial penetration. Cuticle integrity often declines during ripening as berry expansion outpaces cuticle deposition, explaining the increased susceptibility of mature berries to Botrytis infection.
Kutikula-basierte Resistenz-Disposition
Kutikuläre-Botrytis-Resistenz
Wachsschicht-Resistenz
cuticle barrier resistance
epicuticular defense
wax layer resistance
Comménil P et al. (1997) Surface waxes of grapevine grape berries and their role in pathogenicity of Botrytis cinerea, Journal of Experimental Botany 48:1599-1607 Gabler FM et al. (2003) Correlations of morphological, anatomical, and chemical features of grape berries with resistance to Botrytis cinerea, Phytopathology 93:1263-1273 Becker T, Knoche M (2012) Water movement through the surfaces of the grape berry and its stem, American Journal of Enology and Viticulture 63:100-108
Cuticle Based Resistance Disposition
Cabernet Sauvignon cuticle layer at maturity measures 4.5-6.5 μm total thickness comprising cuticular layer (0.8-1.2 μm interfacing with epidermal cell wall), cutin matrix layer (2.8-4.2 μm bulk of cuticle), and epicuticular wax layer (wax crystalloids 0.9-1.5 μm height projecting from surface), with total wax content 150-185 μg/cm² berry surface (composition= 42% alkanes predominantly C29 and C31, 28% primary alcohols predominantly C26 and C28, 16% esters, 14% other), providing water vapor permeability 1.1×10⁻⁹ m s⁻¹ limiting transpiration to 1.2 g H₂O m⁻² h⁻¹, contact angle 105-115° creating hydrophobic surface, and characteristic waxy bloom visible on dark purple-black berries. Chardonnay white cultivar cuticle layer showing thinner characteristics= 2.8-4.5 μm total thickness with 85-120 μg/cm² wax content (lower than typical red cultivars), providing water permeability 1.5×10⁻⁹ m s⁻¹ (higher than thicker red cultivar cuticles, creating 1.5-1.9 g H₂O m⁻² h⁻¹ transpiration), contact angle 98-108° (moderate hydrophobicity), with thinner cuticle potentially contributing to white wine cultivar sensitivity to bunch rots and requiring careful canopy management for disease prevention. Syrah cuticle layer wax composition developmental changes= veraison (BBCH 81, week 10) showing 95 μg/cm² wax with 38% alkanes (C27-C31), 32% alcohols (C24-C30), higher proportion of shorter-chain components maturity (BBCH 89, week 18) showing 165 μg/cm² wax with 45% alkanes (C29-C33), 26% alcohols (C26-C32), shift toward very-long-chain components, demonstrating progressive wax deposition and compositional maturation enhancing barrier properties during ripening phase. Pinot Noir cuticle layer light exposure effects= sun-exposed clusters showing 5.5-7.5 μm cuticle with 180-220 μg/cm² wax, contact angle 108-118°, transpiration 0.9-1.3 g H₂O m⁻² h⁻¹ shaded clusters showing 3.2-4.8 μm cuticle with 95-135 μg/cm² wax, contact angle 96-106°, transpiration 1.4-2.1 g H₂O m⁻² h⁻¹, demonstrating UV/sunlight induction of cuticle and wax deposition enhancing barrier properties in exposed berries important for stress tolerance. Merlot cuticle layer cracking during berry expansion= rapid Phase II expansion (weeks 10-14, BBCH 81-87) creating microcracks in cuticle as berry surface area increases 80-140% during this period potentially exceeding cuticle elastic capacity, with cracks 0.5-3 μm width concentrated at berry equator where expansion stress maximum, creating potential entry points for Botrytis and other pathogens particularly if wet conditions coincide with cracking period, demonstrating cuticle mechanical vulnerability during rapid berry expansion requiring disease management vigilance during this critical phase.
A cuticle layer is the outermost acellular covering of the berry epidermis comprising cutin matrix, embedded waxes, and epicuticular wax deposits, typically 2-8 μm total thickness at maturity, serving as primary barrier against water loss, pathogen penetration, and environmental stresses while determining berry surface properties including waxy bloom appearance, hydrophobicity, and gas exchange limitations. Cuticle layer structure comprises from interior to exterior= (1) cuticular layer (innermost cuticle zone) 0.5-1.5 μm thick interfacing with outer epidermal cell wall, consisting of cutin matrix with embedded polysaccharides from cell wall creating transition zone between wall and cuticle proper (2) cutin matrix layer (middle cuticle zone) 2-5 μm thick comprising the bulk of cuticle thickness, consisting of cross-linked polyester of C16 and C18 hydroxy and epoxy fatty acids (primarily 16-hydroxypalmitic acid, 9,10-epoxy-18-hydroxyoctadecanoic acid) creating hydrophobic three-dimensional polymer network providing structural framework and barrier properties (3) epicuticular wax layer (outermost cuticle zone) consisting of wax embedded in outer cutin matrix (intracuticular wax) and wax deposited on cuticle surface (epicuticular wax crystalloids projecting 0.5-2 μm from surface creating bloom visible as whitish coating on berry surface). Cuticle layer composition shows two major chemical components= (1) cutin (polymer component) comprising 40-60% of cuticle dry weight, providing structural integrity and continuous barrier (2) waxes (extractable lipid component) comprising 40-60% of cuticle dry weight, total 50-250 μg/cm² berry surface area, with wax composition including very-long-chain aliphatic compounds (alkanes C25-C33 typically 35-50% of total wax, primary alcohols C24-C32 typically 20-35%, fatty acid esters typically 10-25%, minor components including aldehydes, ketones, triterpenes, sterols). Cuticle layer barrier properties= (1) water permeability very low at 0.8-2×10⁻⁹ m s⁻¹ (cuticle alone) determining berry transpiration rate typically 0.8-2 g H₂O m⁻² h⁻¹ at 25°C and 50% RH under intact conditions (2) water permeability increasing dramatically (10-100 fold) when cuticle damaged through cracks, lenticel development, or pathogen penetration (3) gas permeability limited creating barrier to O₂, CO₂, and ethylene diffusion though lenticels provide gas exchange pathways (4) pathogen resistance providing physical barrier to fungal and bacterial penetration requiring pathogens to penetrate through cuticle enzymes (cutinases) or to enter through natural openings (stomata, lenticels) or wounds. Cuticle layer surface properties= (1) hydrophobicity with water contact angle typically 95-120° creating water-repellent surface causing water droplets to bead and roll off berry surface rather than forming water films (2) waxy bloom appearance from epicuticular wax crystalloids creating whitish coating on berry surface, particularly evident on dark-skinned cultivars where contrast with anthocyanin-pigmented skin creates visual bloom (3) tactile properties with intact cuticle creating smooth waxy feel versus damaged cuticle creating rougher sticky feel. Cuticle layer development= cuticle deposition beginning at fruit set with initial thickness 0.8-1.5 μm, continuing throughout berry development with progressive cutin polymerization and wax deposition, achieving mature thickness 2-8 μm at harvest wax composition changing during development with increasing proportion of very-long-chain components during ripening cuticle sometimes showing microcracks during rapid berry expansion phases creating stress points for potential pathogen entry or water loss.
Cuticle_Layer represents the outermost acellular barrier and is the primary component of the exocarp/berry skin providing water and pathogen barrier functions. Sub-classification into Waxy and Protective cuticle emphasizes different aspects of cuticle function. Understanding cuticle layer structure, composition, and properties is essential for comprehending berry water relations (transpiration, splitting susceptibility), pathogen resistance (barrier to infection), post-harvest handling (water loss rates), and environmental stress responses (drought tolerance). The cuticle is the first line of defense against environmental challenges.
Cuticula-Schicht
Wachsüberzug
cuticular layer
waxy covering
Rogiers, S.Y. et al. (2004). Grape berry epicuticular wax and transpiration during ripening. American Journal of Enology and Viticulture 55=121-127 Rosenquist, J.K., Morrison, J.C. (1989). Factors affecting cuticle and wax accumulation on grape berries. American Journal of Enology and Viticulture 40=241-244 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 176-180.
Cuticle Layer
Quantitative analysis of Cuticle_Thickness_Disposition at the University of Adelaide characterized cuticular properties across 18 wine grape cultivars on both leaves and berries using transmission electron microscopy and chemical extraction methods. Leaf cuticle thickness (measured on mature, fully expanded leaves at véraison) ranged from 1.9 μm ('Pinot Noir') to 4.8 μm ('Cabernet Sauvignon'), while berry cuticle thickness at harvest ranged from 2.4 μm ('Pinot Noir') to 6.2 μm ('Grenache'). Cuticle thickness was moderately heritable (h² = 0.52 for leaves, 0.61 for berries) and showed strong organ correlation (r = 0.78 between leaf and berry thickness within cultivars). Controlled inoculation experiments demonstrated that Cuticle_Thickness_Disposition significantly affected powdery mildew establishment, with infection success (appressorial penetration rate) negatively correlated with leaf cuticle thickness (r = -0.68). Cultivars with thick cuticles including 'Cabernet Sauvignon' (4.8 μm) showed 55% lower penetration success than thin-cuticle 'Pinot Noir' (1.9 μm) at equivalent inoculum loads. For Botrytis, berry cuticle thickness was inversely correlated with unwounded infection severity (r = -0.72), with 'Grenache' (6.2 μm) showing 75% lower infection than 'Pinot Noir' (2.4 μm) under controlled humidity chamber conditions. These findings validate Cuticle_Thickness_Disposition as a selectable trait contributing to disease resistance.
A Cuticle_Thickness_Disposition is a Physical_Barrier_Disposition that inheres in a Vitis vinifera plant and is realized through the development of a thick cuticular membrane covering aerial plant surfaces, providing enhanced resistance to pathogen penetration, reduced water loss, and protection against environmental stresses. This disposition represents the quantitative aspect of cuticle development, with thicker cuticles generally providing greater barrier function against direct-penetrating pathogens and environmental challenges. The plant cuticle is a continuous hydrophobic layer covering all aerial organs, composed of a cutin polyester matrix embedded with waxes (intracuticular) and overlaid by epicuticular wax crystals, together forming the primary interface between the plant and its environment. In grapevine, cuticle thickness varies substantially among organs (berries typically thicker than leaves), developmental stages (increasing during maturation), and genotypes (2-6 μm range among cultivars on mature leaves). Thick cuticles impede direct penetration by pathogens such as Erysiphe necator (powdery mildew), which must breach the cuticle to access epidermal cells, and Botrytis cinerea, which secretes cutinases to degrade the cuticle during infection. Additionally, thick cuticles reduce pathogen adhesion, limit diffusion of pathogen enzymes and toxins, and decrease the effectiveness of cutinase-mediated penetration. Cuticle_Thickness_Disposition is particularly relevant for berry resistance to bunch rots, where thick berry cuticles represent a primary defense against post-véraison infection.
Cuticle_Thickness_Disposition is_a Physical_Barrier_Disposition. Cuticle thickness affects resistance to direct-penetrating pathogens (powdery mildew, Botrytis) but not stomatal-penetrating pathogens (downy mildew). Thickness is one aspect of cuticular defense alongside wax composition, crystal morphology, and hydrophobicity. Berry cuticle thickness is particularly important for bunch rot resistance, relating to Cuticle_Based_Resistance_Disposition under Botrytis_Resistance_Disposition, though the present class addresses the mechanistic trait rather than resistance to a specific pathogen.
Kutikula-Entwicklung
Kutikuladicke-Disposition
Wachsschicht-Stärke
cuticle development
cuticular membrane thickness
epicuticular barrier
Yeats TH, Rose JKC (2013) The formation and function of plant cuticles, Plant Physiology 163:5-20 Comménil P et al. (1997) Host-pathogen interactions during infection of grape berries by Botrytis cinerea, VITIS Journal of Grapevine Research 36:193-198 Dry IB, Thomas MR (2015) Molecular genetics of grape berry ripening, Advances in Botanical Research 74:325-361
Cuticle Thickness Disposition
Cuticular transpiration in grapevine leaves accounts for 2-10% of total water loss under well-watered conditions but becomes the dominant pathway during prolonged stomatal closure, with minimum conductance (gmin) ranging from 3-15 mmol H₂O/m²/s across genotypes. Cultivars with thick cuticles and dense epicuticular wax such as Carignane and Monastrell show low gmin (3-6 mmol/m²/s), while thin-cuticled types like Pinot Noir show higher residual conductance (10-15 mmol/m²/s). Cuticular conductance increases with leaf age (senescent leaves showing 2-3× higher gmin than mature leaves) and following physical damage or pathogen infection. Berry cuticular conductance of 0.5-3.0 mmol/m²/s contributes to post-veraison berry water loss and concentration.
A transpiration disposition that is the inherited capacity of a Vitis vinifera plant to lose water vapor through the cuticle and associated epicuticular waxes when stomata are closed, with genotype-specific residual conductance determined by cuticle thickness, wax composition, and the presence of cuticular cracks or imperfections.
Cuticular transpiration represents the irreducible water loss that occurs even when stomata are fully closed, becoming critical during severe drought when stomatal closure is complete. Low cuticular conductance extends survival time during prolonged water deficit by slowing desiccation. The trait is determined by cuticle development during leaf expansion and shows limited plasticity in mature leaves. Berry cuticular properties affect fruit quality through water loss and concentration effects, particularly in late-harvest scenarios.
Mindestwasserverlust
Residualtranspiration
cuticular water loss trait
kutikuläre Transpiration
minimum conductance
residual transpiration
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 65-72 Blanke MM, Leyhe A (1987) Stomatal activity of the grape berry cv. Riesling, Müller-Thurgau and Ehrenfelser. Journal of Plant Physiology 127:451-460
Cuticular Transpiration Disposition
Cuticular transpiration assessment comparing drought-tolerant and sensitive grapevine selections reveals the contribution of cuticle properties to drought survival, with tolerant selection showing cuticular conductance of 4.5 mmol/m²/s (low residual water loss when stomata closed) while sensitive selection shows 12.8 mmol/m²/s, with the 2.8-fold difference in minimum conductance translating to proportional difference in water loss rate during prolonged stomatal closure, affecting survival time under severe drought by approximately 60% and demonstrating how cuticular properties contribute to drought tolerance beyond stomatal regulation.
Cuticular transpiration quality is a transpiration rate quality that represents the rate of water vapor loss through the cuticle when stomata are closed, characterizing the residual, unregulated water loss that continues regardless of stomatal aperture and determines minimum transpiration rate under severe stress when stomatal closure is complete. Within the BFO framework, this quality specifically addresses the non-stomatal component of transpiration, distinguishing cuticular loss from regulated stomatal transpiration while recognizing that cuticular transpiration, though typically representing only 5-10% of total transpiration when stomata are open, becomes the dominant pathway under severe stress when stomatal closure eliminates the regulated component. The ontological significance lies in cuticular transpiration determining the minimum water loss rate that the vine cannot reduce below regardless of stomatal regulation, where the magnitude of cuticular conductance establishes the water cost of surviving drought periods when stomata are fully closed and affects the time course of dehydration under severe stress conditions. The physical basis for cuticular transpiration involves water vapor diffusion through the lipophilic cuticular membrane that covers the epidermis, with cuticle permeability determined by wax composition, wax amount, and cuticle structure, creating a diffusion pathway that lacks the regulatory capacity of stomata but provides essential protection against excessive water loss. The quantitative ranges for cuticular conductance in Vitis vinifera typically span 3-20 mmol/m²/s (compared to stomatal conductance of 50-400 mmol/m²/s when open), with the variation reflecting both genetic differences in cuticle properties and environmental effects on cuticle development including light intensity, humidity during growth, and leaf age. The stress tolerance implications of cuticular transpiration are significant, as cultivars or conditions producing lower cuticular conductance enable longer survival under severe drought by reducing the rate of water loss when stomatal protection is maximized, while high cuticular conductance limits the effectiveness of stomatal closure in conserving water. Understanding cuticular transpiration quality enables assessment of residual water loss under stress, prediction of dehydration rates under severe drought, and evaluation of cultivar differences in water-conserving capacity beyond stomatal regulation.
Cuticular_Transpiration_Quality addresses the non-regulated water loss through the cuticle when stomata are closed. This quality determines minimum transpiration and affects drought survival capacity. The class supports drought tolerance assessment through cuticular conductance characterization revealing residual water loss potential.
Minimalleitfähigkeit
cuticular conductance
gmin
kutikulärer Wasserverlust
Kerstiens G (1996) Cuticular water permeability and its physiological significance. Journal of Experimental Botany 47(12): 1813-1832 Leide J et al. (2007) The developmental pattern of tomato fruit wax accumulation and its impact on cuticular transpiration barrier properties. Plant Physiology 144(3): 1667-1679
Cuticular Transpiration Quality
110 Richter rootstock hardwood cutting showing cutting adventitious roots, cutting 32 cm length with 4 nodes treated with 3500 ppm IBA, basal callus visible day 8, root primordia detected microscopically day 12, first roots emerging from nodes and basal callus day 18, cutting producing 11 adventitious roots by day 32 (7 nodal, 4 callus-derived), roots 3-8 cm length with lateral branching, cutting ready for nursery planting with functional root system enabling establishment, cutting adventitious roots enabling successful vegetative propagation maintaining 110R genetic identity
Cutting adventitious root is adventitious root formed on vegetative cutting comprising severed stem, cane, or shoot segment used for propagation, characterized by development in response to cutting preparation and favorable rooting conditions enabling severed shoot to establish independent root system, showing origin from nodes, internodes, or basal callus depending on cultivar and conditions, demonstrating dependence on endogenous carbohydrate reserves and hormones in cutting supporting root initiation and early development, and serving propagation function enabling clonal reproduction maintaining exact genetic identity of parent vine, with cutting adventitious roots in viticulture representing foundation of commercial propagation where hardwood or green cuttings treated with hormones and placed in appropriate media develop root systems allowing establishment as independent vines, exhibiting typical developmental sequence of cutting preparation triggering wound response and auxin accumulation, root primordia initiating 5-15 days after treatment, roots emerging 12-25 days, and functional root system developing 3-8 weeks enabling cutting transplanting, and demonstrating that cutting rooting capacity represents critical cultivar characteristic determining propagation ease, success rate, and commercial viability with easy-rooting cultivars producing abundant roots rapidly while difficult cultivars requiring special treatments and showing lower success. Cutting adventitious root in hardwood cuttings includes= showing dormant cane use (hardwood cuttings prepared from dormant one-year canes during winter pruning), exhibiting typical dimensions (cuttings 25-40 cm length with 3-5 nodes providing adequate reserves and rooting sites), demonstrating callusing period (cuttings often callused in warm humid conditions promoting basal callus before rooting), revealing auxin treatment (hardwood cuttings commonly treated with 2000-8000 ppm IBA or NAA promoting root initiation), and displaying spring planting (callused cuttings planted in nursery or propagation beds spring for root and shoot development). Cutting adventitious root in green cuttings includes= showing actively growing shoot use (green cuttings prepared from current season green shoots during growing season), exhibiting smaller size (green cuttings typically 10-20 cm length with 2-3 nodes due to tender tissue), demonstrating misting requirement (green cuttings needing high humidity preventing desiccation during rooting), revealing lower hormone needs (green cuttings sometimes rooting with lower auxin concentrations or without treatment due to endogenous auxin), and displaying faster rooting (green cuttings often producing roots more quickly than hardwood though requiring more care). Cutting adventitious root carbohydrate dependence includes= showing reserve utilization (cutting depending on stored carbohydrates for energy supporting root initiation and early development), exhibiting starch depletion (cutting carbohydrate reserves declining during rooting period as consumed for respiration and growth), demonstrating reserve importance (cuttings with adequate reserves showing higher rooting success than depleted cuttings), revealing cutting size effects (larger cuttings with more reserves generally showing better rooting than small cuttings), and displaying timing implications (cutting collection timing affecting reserve status influencing rooting success). Cutting adventitious root hormone dynamics includes= showing endogenous auxin (cuttings containing native auxin from shoot apex or synthesized at wound), exhibiting exogenous treatment (commercial rooting hormone application supplementing endogenous auxin improving initiation), demonstrating concentration effects (optimal auxin concentrations typically 1000-10,000 ppm depending on cultivar and cutting type), revealing formulation importance (IBA and NAA most common with IBA generally preferred for grapevine), and displaying application methods (quick-dip, long-soak, or powder formulations providing different auxin delivery). Cutting adventitious root environmental requirements includes= requiring adequate moisture (maintaining cutting hydration essential for rooting with misting, fog, or humidity tents), showing temperature optima (root zone temperatures 20-25°C promoting initiation while cooler temperatures delaying), exhibiting medium importance (rooting media providing aeration, moisture retention, and pathogen resistance), demonstrating light needs (moderate light supporting photosynthesis in green cuttings while hardwood rooting in dark or low light), and displaying sanitation necessity (pathogen-free conditions preventing cutting rot and rooting failure). Cutting adventitious root rooting stages includes= showing lag phase (initial period 5-15 days with no visible roots while primordia initiate), exhibiting emergence phase (roots breaking through cutting surface 10-25 days becoming visible), demonstrating elongation phase (emerged roots extending 1-5 cm creating functional absorption), revealing establishment phase (roots developing laterals and beginning active uptake 3-6 weeks supporting cutting), and displaying maturation phase (root system developing complete architecture 6-12 weeks enabling transplanting). Cutting adventitious root success factors includes= depending on cultivar capacity (intrinsic rooting ability varying widely among cultivars determining baseline success), requiring proper timing (cutting collection at appropriate season and physiological stage), showing treatment importance (hormone type, concentration, and application method affecting results), demonstrating environmental control (temperature, humidity, media affecting rooting success significantly), and displaying sanitation needs (pathogen management preventing losses from rot and disease). Cutting adventitious root in commercial propagation includes= enabling rootstock production (rootstock varieties propagated through hardwood cutting rooting producing millions of plants annually), providing scion propagation (some V. vinifera cultivars propagated directly through cuttings for own-rooted vineyards), demonstrating nursery foundation (cutting rooting representing first step in grafted vine production with rootstock cuttings rooted before grafting), revealing economic importance (propagation industry depending on reliable efficient cutting rooting), and displaying quality standards (propagation protocols designed to maximize rooting percentage, root system quality, and plant uniformity).
Cutting_Adventitious_Root emphasizes propagation cutting context and is distinguished from other adventitious roots by focus on severed cutting. Includes Propagation_Cutting_Adventitious_Root and Rooted_Cutting_Adventitious_Root. Critical for understanding cutting propagation, carbohydrate dependence, hormone treatment, rooting stages, success factors, and the adventitious roots enabling vegetative propagation through cutting rooting fundamental to viticulture.
Clonal root
Klonale Wurzel
Propagation cutting root
Stecklings-Adventivwurzel
Vegetative cutting root
Vermehrungs-Stecklingswurzel
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press.
Cutting Adventitious Root
A 30 cm segment of dormant SO4 rootstock cane bears a Cutting_Role, prepared for rooting in nursery beds or callusing boxes to produce either rooted rootstock liners or benchgraft rootstock components. A single-node green cutting of Chardonnay serves in a Cutting_Role for rapid multiplication of virus-tested material, rooted under mist with bottom heat to maximize propagation rate from limited nuclear stock. A three-node hardwood cutting of Riparia Gloire bears a Cutting_Role in traditional rootstock propagation, planted directly in nursery rows for field rooting without the controlled environment required for green cuttings.
A propagation material role that inheres in a severed stem segment (cane piece) of Vitis vinifera or a Vitis hybrid by virtue of its function in vegetative propagation through adventitious root formation, representing the primary multiplication method for grapevine rootstocks and the rootstock component of grafted vines. Cuttings exploit the grapevine's capacity for adventitious root initiation from cambial and parenchymal tissues, developing functional root systems when basal tissues are exposed to appropriate moisture, temperature, and hormonal conditions. Cutting propagation maintains clonal identity (all progeny genetically identical to the source plant) and represents the most cost-effective large-scale multiplication method for grapevines. Cutting roles are subclassified by physiological state at harvest (hardwood from dormant canes, softwood from actively growing shoots, semi-hardwood from partially matured tissue) and by node configuration (single-node, two-node, three-node), with these parameters influencing rooting success, multiplication efficiency, and handling requirements. Rootstock cutting propagation produces either standalone rooted rootstocks (for field budding) or serves as the root-forming component of benchgrafts.
Cutting_Role is the foundational propagation category for grapevines, with subclasses distinguishing physiological state (hardwood versus softwood versus semi-hardwood) and nodal configuration (single, two, three nodes). The choice among cutting types involves tradeoffs between multiplication rate (maximized with single-node cuttings that produce more propagules per unit of source material), rooting success (generally higher with larger cuttings having greater carbohydrate reserves), and production system requirements (hardwood cuttings require only dormant storage while green cuttings need mist facilities and climate control). Rooting capacity varies dramatically among grapevine genotypes—V. rupestris and V. champinii root easily (>80% success), while V. berlandieri roots with difficulty (<30% without treatment)—determining which rootstock genotypes are commercially viable for cutting propagation.
Rebsteckling
Steckholz
Steckling
cane cutting
stem cutting
vegetative cutting
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 344-380 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Pongrácz DP (1983) Rootstocks for Grape-vines, David Philip Publishers Smart DR et al. (2006) Grapevine rooting patterns, American Journal of Enology and Viticulture 57: 89-104
Cutting Role
Cyanidin-3-O-glucoside typically comprises 2-15% of total anthocyanins in most V. vinifera cultivars, but reaches elevated proportions (15-35%) in specific genotypes including Sangiovese, Nebbiolo, and Pinot Noir. The cyanidin pathway branches from dihydroquercetin via DFR and ANS without F3'5'H involvement, with F3'H converting dihydrokaempferol to dihydroquercetin. Cyanidin shows λmax of 506-513 nm, contributing orange-red hues distinct from the purple-blue of malvidin derivatives. The 3',4'-dihydroxylated B-ring of cyanidin lacks the methoxy groups that stabilize malvidin, resulting in faster color loss under oxidative conditions. Sangiovese wines owe their characteristic bright ruby-red color partly to elevated cyanidin proportions.
An anthocyanin accumulation disposition that is the inherited capacity of a Vitis vinifera plant to accumulate cyanidin-based anthocyanins (cyanidin-3-O-glucoside and derivatives) as a significant proportion of total pigments, resulting from elevated flavonoid 3'-hydroxylase (F3'H) activity relative to F3'5'H and producing red-orange hue contributions to wine color.
Cyanidin accumulation represents the alternative branch of the anthocyanin pathway competing with malvidin synthesis for common precursors. The F3'H/F3'5'H expression ratio largely determines the cyanidin/malvidin balance, with cultivar-specific regulatory patterns established early in berry development. Elevated cyanidin is characteristic of some traditional Italian cultivars (Sangiovese, Nebbiolo) and contributes to their distinctive color profiles. The lower stability of cyanidin-based color may contribute to the faster color evolution observed in wines from these cultivars.
Cyanidin-Akkumulation
Cyanidin-Einlagerung
Cyanidingehalt
cyanidin content trait
cyanidin-type pigmentation
di-hydroxylated anthocyanin
Mattivi F et al. (2006) Metabolite profiling of grape: flavonols and anthocyanins. Journal of Agricultural and Food Chemistry 54:7692-7702 Rinaldi A et al. (2020) Metabolic and RNA profiling elucidates proanthocyanidins accumulation in Aglianico grape. Food Chemistry 312:126045
Cyanidin Accumulation Disposition
Cyanidin content quality in Sangiovese from Montalcino distinguishes this Tuscan variety from Bordeaux cultivars, with cyanidin-based anthocyanins comprising 12-22% of total anthocyanins compared to less than 5% in Cabernet Sauvignon this elevated cyanidin proportion, combined with relatively low total anthocyanin concentration (300-600 mg/kg), produces the translucent ruby-garnet color and orange-red rim characteristic of Brunello di Montalcino that contrasts with the deeper, more purple-black hues of Bordeaux-style wines.
Cyanidin_Content_Quality represents the chemical quality characterizing the concentration of cyanidin-based anthocyanins in grape berry skins, capturing the simplest dihydroxylated anthocyanidin that produces orange-red coloration. Cyanidin (3',4',5,7-tetrahydroxyflavylium) bears hydroxyl groups at the 3' and 4' positions of the B ring without methoxylation, representing the direct product of flavonoid 3'-hydroxylase (F3'H) activity on dihydrokaempferol in the anthocyanin biosynthetic pathway. This structural simplicity, with only two B-ring substituents compared to three in delphinidin-type anthocyanidins, contributes to the characteristic orange-red rather than blue-red hue of cyanidin-based pigments. In most Vitis vinifera cultivars, cyanidin-3-glucoside comprises only 1-5% of total anthocyanins because F3'5'H activity preferentially directs flux toward trihydroxylated anthocyanidins (delphinidin, petunidin, malvidin). However, certain cultivars show elevated cyanidin content due to reduced F3'5'H expression or activity, creating redder, less blue-shifted color profiles. Sangiovese, the signature variety of Tuscany, notably produces relatively high proportions of cyanidin (10-25% of total anthocyanins) compared to Bordeaux varieties, contributing to its characteristically ruby-red rather than purple-black coloration. Cyanidin content provides a varietal marker and influences wine color hue, with cyanidin-rich wines displaying more orange-red tones particularly as they age and monomeric anthocyanins decline relative to polymeric pigments.
Cyanidin_Content_Quality addresses the dihydroxylated B-ring anthocyanidin that represents the most structurally basic colored anthocyanin. The F3'H/F3'5'H enzyme ratio controls the balance between dihydroxylated (cyanidin, peonidin) and trihydroxylated (delphinidin, petunidin, malvidin) anthocyanidins, making this ratio a key determinant of wine color hue. Cyanidin is less stable than methylated malvidin, contributing to color evolution during aging. The orange-red hue of cyanidin contrasts with the blue-red of malvidin, with intermediate colors resulting from mixtures.
Cy-3-glc content
Cyanidin-3-glucosid-Konzentration
Cyanidinpigmentqualität
cyanidin pigment quality
cyanidin-3-glucoside content
dihydroxylated anthocyanin concentration
dihydroxylierter Anthocyangehalt
Castellarin SD et al. (2007) Colour Variation in Red Grapevines (Vitis vinifera L.): Genomic Organisation, Expression of Flavonoid 3'-Hydroxylase, Flavonoid 3',5'-Hydroxylase Genes and Related Metabolite Profiling of Red Cyanidin-/Blue Delphinidin-Based Anthocyanins in Berry Skin. BMC Genomics 8(1): 428 Mattivi F et al. (2006) Metabolite Profiling of Grape: Flavonols and Anthocyanins. Journal of Agricultural and Food Chemistry 54(20): 7692-7702
Cyanidin Content Quality
Cluster architecture in Riesling exemplifies cylindrical form, with typical clusters displaying length-to-width ratios of 2.5-3.5 and relatively uniform width from the first lateral branches to the cluster tip this elongated cylindrical architecture with length of 12-18 cm and consistent width of 5-7 cm creates moderate compactness that balances disease resistance against the complete looseness that would impede harvesting, contributing to Riesling's adaptability across diverse climatic conditions from cool German valleys to warm Australian regions.
Cylindrical_Quality represents the overall form characteristic of organs exhibiting elongated geometry with relatively parallel sides along the major portion of their length, creating a tube-like or rod-like appearance. In grapevine contexts, cylindrical form applies primarily to clusters and to extremely elongated berries. Cylindrical cluster form describes bunches where the width remains relatively constant from shoulders to tip, contrasting with conical clusters that taper progressively. Cylindrical clusters typically exhibit length-to-width ratios exceeding 2.0 and are characteristic of cultivars including Riesling, Silvaner, and many Muscat varieties. The uniform width along the rachis results from consistent fruit set across successive branching orders and similar berry sizes throughout the cluster. Cylindrical cluster architecture affects harvest handling, with elongated cylindrical clusters requiring careful orientation in picking bins to avoid mechanical damage. Berry form approaching cylindrical occurs in highly elongated cultivars like Ladies Finger (Menindee Seedless), where length-to-width ratios may exceed 2.0-2.5 with nearly parallel sides along the berry midsection. Such extremely elongated berries are rare in wine grapes but valued in some table grape markets for their distinctive appearance. Cylindrical form assessment employs the length-to-width ratio combined with evaluation of side parallelism along the organ's length. OIV descriptor 208 (cluster shape) specifically identifies cylindrical as a primary cluster architecture category.
Cylindrical_Quality in the VVO applies primarily to cluster form rather than berry form, as truly cylindrical berries are rare. The distinction between cylindrical and conical cluster form can be subtle when clusters exhibit only slight tapering the cylindrical classification applies when the width at the cluster tip is at least 80% of the maximum width at the shoulders. OIV descriptor 208 category 1 (cylindrical) provides the standardized reference. For berries, extreme elongation with parallel sides grades into the Finger-shaped category in some classification systems. The cylindrical form of internodes and petioles is treated under Diameter_Quality rather than as a shape category.
Zylinderform
column-like shape
elongated parallel-sided form
rod-shaped quality
röhrenförmige Gestalt
tubular form
walzenförmige Form
zylindrische Qualität
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 87-88 Tello J and Ibáñez J (2014) Morphological and Microsatellite Characterization of Ancient Minority Grapevine Cultivars from Aragón. American Journal of Enology and Viticulture 65(4): 469-478
Cylindrical Quality
The DAC designation Weinviertel DAC (established 2003 as the first Austrian DAC) identifies wines from the Weinviertel region in Lower Austria that are produced exclusively from Grüner Veltliner displaying the characteristic peppery, spicy profile typical of the region — the DAC specification requires 100% Grüner Veltliner, dry style (maximum 6 g/L residual sugar), minimum 12% alcohol, and approval by a regional tasting commission assessing typicity and quality. Wines from the same region that do not meet DAC criteria (e.g., Riesling, Welschriesling, or atypical Grüner Veltliner styles) are classified under the broader Niederösterreich designation, creating a regulatory incentive for producers to align their production with the defined regional identity.
A regional wine designation within the Austrian wine classification system that identifies a wine as a Districtus Austriae Controllatus — a geographically specific, regionally typical wine from one of Austria's designated DAC appellations, produced according to regional production specifications that define authorized grape varieties, wine styles, and quality parameters designed to express the characteristic terroir identity of the designated region, formally classified as a Protected Designation of Origin under EU harmonization.
DAC_Designation documents Austria's distinctive appellation system introduced in 2003 as a typicity-focused complement to the traditional Qualitätswein system based primarily on must weight (Oechsle/KMW). The DAC concept explicitly prioritizes regional typicity — the distinctive character associated with a specific cultivar-terroir combination — over quality level defined by sugar concentration at harvest: this philosophical approach differs from both the French AOC model (which defines production rules without necessarily specifying a single dominant cultivar) and the traditional Austrian/German Prädikat model (which ranks wines by must weight). Austria currently has 18 DAC designations, each specifying the characteristic grape variety or varieties and wine style for its region — Weinviertel DAC for Grüner Veltliner, Mittelburgenland DAC for Blaufränkisch, Kamptal DAC and Kremstal DAC for Grüner Veltliner and Riesling, and Leithaberg DAC for both white and red wines from specified varieties.
Austrian controlled origin designation
DAC
DAC-Bezeichnung
Districtus Austriae Controllatus
Districtus-Austriae-Controllatus-Klassifikation
österreichische kontrollierte Herkunftsbezeichnung
Austrian Wine Law (Weingesetz 2009) §13-16 Austrian Wine Marketing Board (ÖWM) (2023) DAC Appellations EU Regulation 1308/2013 Art. 93
DAC Designation
Muscadinia rotundifolia and derived hybrids such as VR 043-43 and O39-16 show strong resistance to Xiphinema index with population reductions exceeding 90% compared to susceptible SO4 or 5BB Kober, critical for managing GFLV in replant situations in Champagne and Burgundy. Dog Ridge (V. champinii) demonstrates moderate resistance to X. americanum complex species (population reduction 50-70%), providing protection against Tomato ringspot virus transmission in northeastern United States vineyards. 1616C (Solonis × Riparia) shows resistance to X. diversicaudatum in German experimental trials, though this species is less commonly problematic in commercial viticulture than X. index.
A nematode-resistance rootstock role that inheres in a grafted rootstock by virtue of its capacity to resist feeding and population buildup by dagger nematodes of the genus Xiphinema, particularly species of viticultural importance including X. index (the Grapevine fanleaf virus vector), X. americanum sensu lato (vectors of Tomato ringspot virus and other nepoviruses), and X. diversicaudatum. Dagger nematodes are migratory ectoparasites that feed on root tips and elongation zones using a long odontostyle (80-150 µm depending on species), causing direct damage through root tip destruction and indirect damage as vectors of nepoviruses. Resistant rootstocks deter Xiphinema feeding through root anatomical features, production of repellent exudates, and reduced nutritional suitability of root tissue. Resistance is assessed by nematode population density per gram root or per 250 cm³ soil, with resistant genotypes supporting 70-90% lower populations than susceptible V. vinifera cultivars.
Dagger_Nematode_Resistant_Rootstock_Role encompasses resistance to the Xiphinema genus generally, with Xiphinema_Index_Resistant_Rootstock_Role as the most important specific subclass due to the critical role of X. index as the GFLV vector. Xiphinema species differ in their host associations, geographic distribution, and virus vector competence resistance to one species does not guarantee resistance to others. Muscadinia rotundifolia carries the strongest known Xiphinema resistance but presents graft compatibility challenges that ongoing breeding programs (INRAE Colmar, UC Davis) are working to overcome through backcrossing approaches.
Dolchnematodenresistente Unterlage
Xiphinema resistant rootstock
Xiphinema-resistente Unterlage
dagger nematode resistant rootstock
ectoparasitic nematode resistant rootstock
gegen Dolchälchen resistente Unterlage
Walker MA et al. (1994) Resistance to Xiphinema index in Vitis and Muscadinia species, Vitis 33: 75-78 Bouquet A et al. (2000) Grapevine rootstocks with resistance to Xiphinema index, Acta Horticulturae 528: 521-528 Brown DJF et al. (1993) Transmission of viruses by plant nematodes, Annual Review of Phytopathology 31: 55-80 Andret-Link P et al. (2004) Grapevine fanleaf virus, Journal of Plant Pathology 86: 183-195
Dagger Nematode Resistant Rootstock Role
Damage state quality assessment at arrival inspection for imported Flame Seedless follows USDA grade standards specifying maximum allowable defects of 4% total damage for US Fancy grade, including no more than 1% decay (any visible Botrytis, sour rot, or other microbial growth), 2% mechanical damage (cuts, punctures, bruises), and 1% other defects (sunburn, shatter) lots exceeding these thresholds are downgraded to US No. 1 (8% total damage tolerance) or US No. 2 (12% tolerance), with corresponding price reductions of 15-25% per grade level that determine whether shipments generate profit or loss for importers.
Damage_State_Quality encompasses the structural integrity qualities characterizing the presence, type, and extent of physical injury or deterioration in grapevine organs. This category addresses deviations from the intact, undamaged condition that compromise organ function, quality, or marketability. Damage represents the negative outcome of mechanical, environmental, physiological, or biological insults that disrupt tissue structure. The category subdivides into specific damage types including Mechanical_Damage_Quality addressing injury from physical forces, Splitting_Quality capturing rupture of the berry skin, and Shriveling_Quality describing water loss and tissue collapse. Additional damage types not captured in child classes include sunburn damage, frost injury, hail damage, insect feeding injury, and disease lesions. The damage state of a population is typically expressed as the percentage of organs showing each damage type and the severity distribution within damaged organs. Damage assessment follows standardized protocols that define damage categories, severity thresholds, and sampling procedures appropriate to the quality context. For table grapes, damage tolerance is very low, with damage exceeding 1-5% causing lot rejection or significant price penalties. For wine grapes, higher damage levels may be acceptable depending on intended use, though damage always increases processing requirements and can affect wine quality. Damage has cumulative effects, as initial injury creates entry points for secondary infection and initiates deterioration cascades that worsen during storage. Prevention of damage through careful handling, protective packaging, and rapid postharvest cooling provides greater quality preservation than any remediation strategy.
Damage_State_Quality serves as the parent class for specific damage type categories and addresses the compromised structural integrity that results from injurious events. Damage represents the current state of an organ resulting from past injurious processes the processes themselves (mechanical impact, water loss, splitting events) are addressed in process ontology classes while Damage_State_Quality captures the resulting condition. Assessment protocols should distinguish damage type, severity, and age (fresh versus healed), as these factors affect quality implications differently. The boundary between damage and normal variation can be subtle standardized reference materials and training are essential for consistent damage classification.
Beschädigungsgrad
Defektpräsenz
Schadenszustand
Verletzungszustand
damage condition
defect presence
deterioration state
injury state
Crisosto CH et al. (2001) Table Grapes Respond to Mechanical Damage. California Agriculture 55(5): 30-31 USDA (2021) United States Standards for Grades of Table Grapes (European or Vinifera Type). United States Department of Agriculture, Agricultural Marketing Service
Damage State Quality
Dark respiration rate measurement in Pinot Noir leaves following sunset quantifies the nighttime carbon loss, with mature leaves showing Rd of 1.4 μmol CO₂/m²/s at 20°C, translating to approximately 0.5 g CO₂/m²/night over 10-hour dark period, representing nighttime consumption of approximately 8% of the carbon fixed during the preceding 14-hour light period, with this respiratory loss reducing the daily net carbon gain from gross photosynthesis of 420 mmol CO₂/m²/day to net gain of approximately 350 mmol CO₂/m²/day after accounting for both dark respiration and daytime (maintenance) respiration.
Dark respiration rate quality is a respiration rate quality that represents the rate of mitochondrial CO₂ release measured in darkness or when photosynthesis is inhibited, characterizing the baseline oxidative metabolism that provides ATP and carbon skeletons for cellular processes when photosynthetic light reactions are not operating. Within the BFO framework, this quality specifically addresses the measurable dark respiratory flux, distinguishing dark respiration from photorespiration (which requires light) while recognizing that dark respiration measurement provides the practical assessment of mitochondrial respiratory capacity used in most physiological studies and carbon balance calculations. The ontological significance lies in dark respiration rate providing the operational measure of respiratory carbon loss used in photosynthesis research and carbon modeling, where the difference between gross photosynthesis and dark respiration gives an estimate of net carbon gain, and where dark respiration measurement enables calculation of daily carbon balance by accounting for nighttime respiratory losses when photosynthesis is zero. The physiological basis for dark respiration includes the same processes as total mitochondrial respiration (glycolysis, TCA cycle, oxidative phosphorylation), with dark measurement ensuring that photorespiratory CO₂ release is not confounded with mitochondrial respiration, providing a clean measure of the non-photorespiratory component that operates continuously regardless of illumination. The measurement considerations for dark respiration include dark adaptation period (allowing post-illumination metabolic transients to subside), temperature control (given strong temperature sensitivity), and tissue selection (leaf dark respiration being most commonly measured but root and fruit respiration also assessable in darkness). The relationship between dark respiration and light-adapted mitochondrial respiration is not straightforward, as evidence suggests that mitochondrial respiration may be partially inhibited in the light (Kok effect), making dark respiration potentially higher than daytime mitochondrial respiration and requiring careful interpretation when constructing complete carbon budgets. Understanding dark respiration rate quality enables carbon balance calculation, respiratory capacity assessment, and practical quantification of the respiratory CO₂ losses that reduce net productivity.
Dark_Respiration_Rate_Quality addresses the practically measurable dark respiratory flux used in most physiological studies. This quality excludes photorespiration by definition (measurement in darkness) and provides the operational respiration value for carbon balance calculations. The class supports carbon budget assessment through the standard respiratory measurement that enables daily carbon balance calculation.
Dunkel-CO₂-Abgabe
Dunkelatmung
Nachtrespiration
Rd-Wert
dark CO₂ release
night respiration
non-photorespiratory respiration
Atkin OK et al. (2000) Relationship between the inhibition of leaf respiration by light and enhancement of leaf dark respiration following light treatment. Australian Journal of Plant Physiology 27(7): 669-680 Griffin KL et al. (2002) Leaf dark respiration is related to leaf nitrogen content and leaf gas exchange. Plant Physiology 129(3): 1064-1074
Dark Respiration Rate Quality
Comprehensive analysis of Deacclimation_Disposition at Washington State University characterized hardiness loss kinetics across 22 cultivars by exposing dormant canes to forcing conditions (20°C, 16-hour photoperiod) at intervals from January through March and monitoring hardiness decline. Deacclimation rate (°C hardiness loss per day), lag time before deacclimation initiation, and final hardiness after 21 days forcing were quantified for each cultivar and date combination. Deacclimation_Disposition showed strong cultivar and timing effects. In January (deep endodormancy), all cultivars showed minimal deacclimation (< 2°C loss in 21 days) regardless of genetic background. By February 15 (post-chilling), dramatic cultivar differences emerged. 'Pinot Noir' exhibited rapid Deacclimation_Disposition with 2.1°C loss per day after 3-day lag, reaching -8°C (from -20°C initial) by day 21. 'Riesling' showed moderate deacclimation (1.4°C per day, 5-day lag, final -12°C from -23°C initial). 'Frontenac' demonstrated slow deacclimation (0.8°C per day, 8-day lag, final -22°C from -33°C initial). By March 1, even resistant cultivars showed increased deacclimation responsiveness. Field observations confirmed laboratory patterns, with rapid-deacclimating 'Pinot Noir' suffering 70% bud damage from a late February freeze following 10 days of warm weather, while slow-deacclimating 'Frontenac' showed 5% damage from the same event. These findings demonstrate that Deacclimation_Disposition critically determines late-winter and spring cold damage vulnerability.
A Deacclimation_Disposition is a Cold_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through the physiological processes by which acquired freezing tolerance is lost in response to warming temperatures and increasing photoperiod, transitioning tissues from the cold-hardy dormant state toward active growth and renewed cold sensitivity. This disposition represents the dynamic capacity for hardiness loss, encompassing the rate of deacclimation, the temperature threshold triggering deacclimation, and the responsiveness to warming cues. Deacclimation in grapevine involves reversal of many acclimation changes including catabolism of cryoprotective compounds, membrane lipid remodeling toward greater saturation, reactivation of growth-related gene expression, and hormonal shifts (declining ABA, increasing GA and cytokinins) preparing tissues for spring growth. The timing and rate of deacclimation reflect the release from endodormancy into ecodormancy and subsequently into active growth, with chilling requirement satisfaction determining the transition to deacclimation-competent state. Premature or excessively rapid deacclimation represents a significant source of winter and spring cold damage, as partially deacclimated tissues are vulnerable to temperature fluctuations that fully acclimated tissues would survive. Deacclimation_Disposition varies among genotypes in both the temperature sensitivity of hardiness loss and the rate of deacclimation once triggered, with some cultivars showing dangerous responsiveness to brief warm periods while others maintain hardiness tenaciously until sustained warming occurs.
Deacclimation_Disposition is_a Cold_Tolerance_Disposition. This class addresses the process of losing cold hardiness, complementing Acclimation_Disposition which addresses hardiness acquisition. Child classes include Deacclimation_Resistance_Disposition (capacity to resist hardiness loss) and Reacclimation_Capacity_Disposition (capacity to regain lost hardiness). Deacclimation timing is regulated by dormancy status, with endodormant tissues resistant to deacclimation until chilling requirement is satisfied. Premature deacclimation creates vulnerability to late-winter and spring cold events.
Deakklimatisierungs-Disposition
Enthärtungs-Kapazität
Frosthärte-Verlust
cold hardiness loss
deacclimation response
dehardening capacity
Kalberer SR et al. (2006) Deacclimation and reacclimation of cold-hardy plants: current understanding and emerging concepts, Plant Science 171:3-16 Ferguson JC et al. (2014) Modeling dormant bud cold hardiness and budbreak in twenty-three Vitis genotypes reveals variation by region of origin, American Journal of Enology and Viticulture 65:59-71 Pagter M, Arora R (2013) Winter survival and deacclimation of perennials under warming climate: physiological perspectives, Physiologia Plantarum 147:75-87
Deacclimation Disposition
Deacclimation rate assessment following late-winter warm spell reveals the cultivar variation in hardiness stability, with slow-deacclimating cultivar (Riesling) losing only 0.8°C hardiness per day during 10-day warm period (15°C average), declining from -23°C to -15°C while maintaining adequate protection, while fast-deacclimating cultivar (Chardonnay) lost 1.5°C per day (declining from -22°C to -7°C, severely compromised), with subsequent return to -10°C causing no damage to Riesling but killing 45% of Chardonnay primary buds, demonstrating how deacclimation rate determines vulnerability to the temperature fluctuations common in late winter.
Deacclimation rate quality is a cold hardiness quality that represents the speed at which freeze tolerance decreases in response to warming conditions during late winter and spring, characterizing how rapidly tissues lose their acquired cold hardiness when exposed to temperatures above the threshold for maintaining acclimation, with deacclimation rate determining the vulnerability to late-season cold events following warm periods. Within the BFO framework, this quality specifically addresses the hardiness loss dimension of cold hardiness, distinguishing how fast protection is lost from how it is gained or the level achieved while recognizing that deacclimation rate determines the persistence of winter protection into spring and the vulnerability created by mid-winter or early-spring warm spells that may trigger hardiness loss followed by damaging cold. The ontological significance lies in deacclimation rate determining the stability of cold protection during the variable conditions of late winter and early spring, where rapid deacclimation creates high vulnerability to temperature fluctuations (warm periods quickly eroding protection, leaving tissues vulnerable to subsequent cold) while slow deacclimation maintains protection more stably (requiring extended warmth before significant hardiness loss, providing buffering against brief warm spells). The physiological basis for deacclimation involves the reversal of acclimation changes—membrane lipid resaturation, cryoprotectant metabolism, rehydration, and gene expression shifts toward growth—that proceed at rates determined by both temperature and the underlying deacclimation tendency of the genotype. The factors affecting deacclimation rate include cultivar genetics (some cultivars inherently deacclimating slowly, others rapidly), the intensity and duration of warming (higher temperatures and longer warm periods accelerating deacclimation), and dormancy status (ecodormant buds potentially deacclimating more rapidly than endodormant buds still under internal growth inhibition). Understanding deacclimation rate quality enables assessment of late-season cold vulnerability, prediction of hardiness status following warm periods, and recognition of the climate patterns (frequent warm-cold fluctuations versus stable conditions) that create deacclimation-related risk.
Deacclimation_Rate_Quality addresses the velocity of freeze tolerance loss in response to warming. This quality determines vulnerability to late-season cold following warm periods. The class supports spring risk assessment through deacclimation rate characterization.
Deakklimatisierungsrate
Enthärtungsrate
Härterückgangsgeschwindigkeit
Kältetoleranzabbau
cold tolerance decline
deacclimation velocity
dehardening rate
hardiness loss rate
Keller M, Mills LJ (2007) Effect of pruning on recovery and productivity of cold-injured Merlot grapevines. American Journal of Enology and Viticulture 58(3): 351-357 Ferguson JC et al. (2014) Dynamic thermal time model of cold hardiness for dormant grapevine buds. Annals of Botany 113(5): 801-811
Deacclimation Rate Quality
Quantitative assessment of Deacclimation_Resistance_Disposition at Cornell University employed controlled warming experiments to characterize resistance parameters across 18 cultivars following endodormancy release. Canes collected in late February (post-chilling) were exposed to 18°C forcing conditions, with hardiness assessed by differential thermal analysis at 2-day intervals for 20 days. Deacclimation_Resistance_Disposition was quantified by three parameters: threshold warmth accumulation to initiate deacclimation (degree-days above 0°C before first measurable hardiness loss), deacclimation lag (days at forcing temperature before deacclimation onset), and half-life (days to lose 50% of initial cold hardiness). 'Frontenac' exhibited exceptional Deacclimation_Resistance_Disposition with threshold of 180 degree-days, 9-day lag, and 18-day half-life. 'Riesling' showed intermediate resistance (threshold 95 degree-days, 5-day lag, 11-day half-life). 'Pinot Noir' demonstrated low resistance (threshold 45 degree-days, 2-day lag, 6-day half-life). Plotting hardiness trajectories revealed that after 14 days at 18°C, 'Frontenac' retained 75% of initial hardiness while 'Pinot Noir' retained only 25%. Field validation during a variable March with alternating warm (15°C) and cold (-15°C) periods showed that 'Pinot Noir' lost hardiness during each warm phase and sustained cumulative damage, while 'Frontenac' maintained sufficient hardiness throughout. Strong Deacclimation_Resistance_Disposition thus provides resilience against the repeated warm-cold cycles characteristic of continental spring weather.
A Deacclimation_Resistance_Disposition is a Deacclimation_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to resist or delay loss of acquired freezing tolerance in response to warming temperatures, maintaining protective hardiness levels despite exposure to deacclimation-inducing conditions. This disposition represents resistance to premature hardiness loss, with high deacclimation resistance conferring extended protection during late-winter warm periods and variable spring weather. Deacclimation resistance reflects both dormancy depth (deeply dormant tissues are inherently resistant to warming responses) and intrinsic resistance of cellular cold tolerance mechanisms to disruption by warming. Genotypes with strong Deacclimation_Resistance_Disposition show minimal hardiness loss during brief warm periods, require extended or more intense warming to initiate deacclimation, and maintain protective hardiness later into spring, providing extended protection against late-season cold events. The temperature threshold for deacclimation initiation, the lag period before measurable hardiness loss begins, and the rate of hardiness decline once initiated all contribute to overall deacclimation resistance. Strong Deacclimation_Resistance_Disposition is particularly valuable in climates with highly variable late-winter and spring weather, where cycling between warm and cold conditions can repeatedly challenge deacclimation-susceptible cultivars while deacclimation-resistant cultivars maintain protective hardiness throughout the transition period.
Deacclimation_Resistance_Disposition is_a Deacclimation_Disposition. Deacclimation resistance determines vulnerability to late-winter and spring cold events following warm periods. Resistance is quantified by threshold warmth to trigger deacclimation, lag time, and rate of hardiness loss once initiated. Resistance is partially independent of maximum hardiness, representing a distinct adaptive trait. High deacclimation resistance is particularly valuable in climates with variable spring weather, providing extended protection during the dormancy-to-growth transition.
Deakklimatisierungs-Resistenz-Disposition
Enthärtungs-Resistenz
Härteverlust-Widerstand
deacclimation stability
dehardening resistance
hardiness loss resistance
Ferguson JC et al. (2011) Dynamic thermal profiles of grapevine cold hardiness, American Journal of Enology and Viticulture 62:407-413 Kalberer SR et al. (2006) Deacclimation and reacclimation of cold-hardy plants: current understanding and emerging concepts, Plant Science 171:3-16 Wolf TK, Cook MK (1992) Seasonal deacclimation patterns of three grape cultivars at constant warm temperature, American Journal of Enology and Viticulture 43:171-179
Deacclimation Resistance Disposition
Syrah grafted on SO4 rootstock at 16 years showing dead outer rootstock bark 2.9 mm thickness, microscopic examination confirming complete cellular death with no viable cells detected (no protoplasts, no intact membranes, no metabolic activity), dead bark comprising suberized cork cells, dead phelloderm cells, and isolated dead phloem cut off by periderm, cell viability staining showing zero staining in outer bark versus strong staining in inner living bark confirming complete death, dead outer bark providing purely passive physical and chemical protection through structural integrity and suberin waterproofing without any active physiological function
Dead outer rootstock bark is outer rootstock bark emphasizing complete absence of living cells and purely structural protective function, characterized by all tissues completely dead with no metabolic activity, respiration, or physiological processes, comprising entirely non-living cork cells with suberized walls, dead phelloderm cells, and isolated dead phloem tissues, showing no capacity for wound healing, pathogen defense, or adaptive responses beyond passive physical and chemical barriers, requiring continuous replacement through new periderm formation deeper in bark as dead outer layers age and slough, and representing terminal stage of bark tissue development where cells having completed suberization and death provide protection through structural integrity and chemical composition rather than active physiological processes. Dead outer rootstock bark complete cellular death includes= containing zero living cells (all cork, phelloderm, and isolated phloem cells completely dead with no viable protoplasts), showing absent membranes (plasma membranes and organellar membranes completely degraded during cell death), exhibiting degraded cell contents (cytoplasm, nucleus, mitochondria, and ribosomes disintegrated leaving only cell walls), demonstrating irreversible death (no possibility of revival or return to living state), and displaying only wall remnants (suberized and lignified cell walls remaining as structural framework without living contents). Dead outer rootstock bark absence of metabolic activity includes= showing zero respiration (no oxygen consumption or CO2 production indicating complete metabolic cessation), exhibiting zero synthesis (no production of proteins, carbohydrates, or any compounds lacking biosynthetic machinery), demonstrating zero enzyme activity (all enzymes degraded with no catalytic function remaining), having zero energy metabolism (no ATP production or energy processing), and displaying zero nutrient processing (no uptake, transformation, or mobilization of substances). Dead outer rootstock bark passive protection only includes= providing purely physical barrier (dead cork structure blocking water loss and pathogen entry without active processes), offering chemical deterrence (suberin and extractives deposited during life persisting after death providing antimicrobial properties), showing structural resistance (dead cell walls maintaining integrity resisting mechanical damage through passive strength), demonstrating no active responses (unable to produce antimicrobial compounds, form reaction zones, or adapt to threats), and requiring no maintenance (dead protective function not requiring energy or metabolic support). Dead outer rootstock bark progressive deterioration includes= showing weathering effects (outer surface oxidizing, darkening, and degrading from sun, rain, and atmospheric exposure), exhibiting crack formation (dead bark losing flexibility cracking under trunk expansion and environmental stress), displaying gradual breakdown (dead tissues slowly decomposing from oxidation and microbial activity), demonstrating sloughing tendency (outermost dead bark separating and falling from trunk as cohesion weakens), and revealing aging degradation (dead bark quality declining over years necessitating replacement through new periderm formation). Dead outer rootstock bark in protective function includes= blocking water loss effectively (suberized cork creating waterproof barrier despite being dead), preventing pathogen entry successfully (dead cork layers providing physical barrier blocking fungal and bacterial penetration), insulating adequately (dead cork low thermal conductivity protecting cambium despite lack of living function), resisting damage well (tough dead tissues protecting trunk from mechanical injury through structural properties), and functioning passively (dead bark providing all protective functions without requiring active physiology or energy expenditure).
Dead_Outer_Rootstock_Bark emphasizes complete cellular death and passive protection and is distinguished from Outer_Rootstock_Bark (general outer bark without death emphasis) by focus on zero living cells and purely structural function. Critical for understanding passive protection, absence of active processes, weathering vulnerability, replacement necessity, and the completely dead protective tissues providing trunk defense through physical and chemical properties without metabolic activity.
Abgestorbene Borke
Dead rhytidome
Inactive protective bark
Nicht-lebende Schutzrinde
Non-living outer bark
Tote äußere Unterlagenrinde
Dead Outer Rootstock Bark
Merlot dead outer trunk bark at 16 years post-planting shows 4.2-6.8 mm depth of completely non-living tissue comprising suberized cork cells (air-filled hollow cells with impermeable walls, zero protoplasm), crushed obliterated phloem (collapsed dead cell remnants 0.4-0.9 mm total thickness), and remnants of 3 successive non-functional cork cambia, all cells definitively dead contrasting sharply with living inner bark 2.8-3.6 mm beneath containing functional sieve tubes and active cambium, dead outer bark showing grey-brown weathered surface with longitudinal cracks 3-6 mm deep, providing waterproof insulation protecting living tissues despite complete lack of metabolic activity
Dead outer trunk bark is outer trunk bark emphasizing the complete absence of living cells throughout its depth, comprising entirely non-living tissue with dead cork cells, obliterated phloem, and remnants of dead cork cambia, representing the definitively non-metabolic protective covering external to living inner bark. Dead outer trunk bark cellular status includes= all cork cells dead at maturity (suberized cells lacking protoplasm having undergone programmed cell death following wall deposition), crushed phloem cells dead (old phloem compressed and killed by xylem expansion losing all metabolic function), dead phelloderm cells (inner periderm layers lacking living contents), remnants of non-functional cork cambia (previous cork cambial initials embedded in outer bark no longer dividing or living), and absolutely zero living cells (complete contrast with inner living bark containing functional phloem and active cambium). Dead outer trunk bark formation process includes= cork cell death during differentiation (phellem cells dying as suberin deposits rendering walls impermeable and incompatible with maintaining living protoplasm), phloem death through crushing (functional phloem becoming non-functional after 3-5 years then progressively crushed and killed as xylem expansion compresses tissue), phelloderm death with periderm aging (initially living phelloderm cells dying as periderm becomes non-functional and superseded by deeper periderms), cork cambium death after replacement (cork cambial initials dying when new deeper cork cambium forms rendering outer cambium obsolete), and progressive accumulation (dead tissues accumulating outward building multi-layered dead outer bark). Dead outer trunk bark structural characteristics includes= hollow suberized cork cells (dead cells with air-filled lumens and waxy impermeable walls), collapsed crushed phloem (flattened cell remnants forming thin bands), disorganized cellular structure (dead cells losing original organization through weathering and compression), progressive deterioration (older outer layers showing increased structural breakdown), and absence of cellular contents (no protoplasm, organelles, or metabolic machinery in any cells). Dead outer trunk bark distinguishing from living tissue includes= zero metabolic activity (no respiration, no enzyme function, no biosynthesis contrasting with active inner bark), no response to stimuli (dead cells incapable of responding to hormones, pathogens, or environmental signals), no wound healing capacity (dead outer bark unable to produce wound tissue, callus, or defense compounds), no starch storage (all carbohydrate reserves in living inner bark, dead outer bark containing zero stored compounds), and complete impermeability (dead suberized cork preventing water and solute movement contrasting with living phloem's active transport). Dead outer trunk bark protective functions despite death includes= waterproofing (dead suberized cork providing excellent water impermeability precisely because cells are dead and walls fully suberized), mechanical protection (dead cork compressibility absorbing impacts despite lack of living contents), thermal insulation (air-filled dead cork cells providing effective temperature buffering), pathogen barrier (impermeable dead cork excluding organisms despite inability to mount active defense), and UV shielding (dead outer bark absorbing radiation protecting living tissues beneath). Dead outer trunk bark thickness and composition includes= depth varying 1-12 mm depending on vine age and cultivar (young vines 1-3 mm, mature 3-7 mm, old 5-12 mm), proportion of cork versus crushed phloem varying (cork comprising 60-85% of dead outer bark volume, crushed phloem 15-40%), number of successive periderm layers (1 layer in young bark, 2-4 layers in mature, 4-8+ layers in old bark), and progressive accumulation (dead outer bark depth increasing with age until balanced by exterior sloughing). Dead outer trunk bark weathering and deterioration includes= UV degradation of lignin (solar radiation breaking down cell wall components causing grey coloration), oxidative breakdown (exposure to oxygen causing chemical degradation of cork suberin and other compounds), freeze-thaw damage (ice formation in outer bark causing physical disruption and cracking), microbial surface colonization (fungi and bacteria degrading outermost dead tissues though unable to penetrate intact suberized layers), and mechanical erosion (wind, rain, and physical abrasion gradually wearing away exterior surfaces). Dead outer trunk bark cracking and fragmentation includes= longitudinal fissures developing (dead inelastic bark cracking under trunk expansion creating vertical splits), deep furrows forming (repeated expansion causing pronounced ridges and valleys), outer layers separating (delamination between periderm layers as expansion exceeds dead bark's capacity), platy or scaly detachment (irregular bark plates separating from trunk), and progressive exterior sloughing (oldest outermost dead bark eventually falling revealing newer dead bark layers beneath maintaining protection). Dead outer trunk bark contrast with inner living bark includes= outer bark completely dead versus inner bark fully alive (sharp transition at innermost functional cork cambium), outer bark providing passive protection versus inner bark active transport (dead cork barrier versus living phloem conducting photosynthate), outer bark accumulating and sloughing versus inner bark continuously produced and functional (dead tissue progressive loss versus living tissue ongoing renewal), outer bark depth approximately equal to inner bark depth (2-6 mm dead outer bark, 2-4 mm living inner bark in typical mature trunk), and complementary functions (dead outer protection enabling living inner transport and storage). Dead outer trunk bark lenticel characteristics includes= lenticels comprising dead loosely arranged cork cells (maintaining porous structure despite cellular death), lenticel gas diffusion through dead tissue (oxygen and carbon dioxide moving through air spaces between dead cork cells reaching living inner bark), lenticel structural persistence (dead porous cork maintaining architecture enabling gas exchange), lenticel depth through entire dead outer bark (connecting atmosphere to living tissues), and lenticel function unaffected by cellular death (passive gas diffusion not requiring living cells). Dead outer trunk bark cultivar-specific characteristics includes= thick dead bark varieties (producing substantial cork accumulation creating 5-12 mm dead outer bark), thin dead bark varieties (limited cork formation creating 2-5 mm dead outer bark), smooth dead bark (uniform cork layers without pronounced cracking), deeply furrowed dead bark (extensive cracking creating 4-10 mm deep longitudinal furrows), platy dead bark (separating in irregular plates), and characteristic colors (cultivar-specific dead bark hues from tan to brown to grey before weathering). Dead outer trunk bark significance in protection includes= providing primary waterproof barrier (dead suberized cork preventing desiccation), creating pathogen exclusion (impermeable dead tissue preventing microbial entry), offering mechanical cushioning (compressible dead cork absorbing impacts), insulating living tissues (air-filled dead cork cells buffering temperature fluctuations), shielding from radiation (dead outer bark absorbing UV protecting sensitive living cambium), and enabling trunk expansion (dead outer bark cracking and sloughing accommodating growth without compromising living tissue). Dead outer trunk bark in wound response includes= wound periderm formation beneath dead bark injuries (living tissue beneath responding to damage by producing new protective cork), dead outer bark unable to heal (damaged dead tissue remaining permanently dead requiring replacement by living tissue beneath), callus growth covering dead bark wounds (living tissues proliferating to seal breaches in dead protective covering), and importance of maintaining integrity (intact dead outer bark critical for protection despite inability to self-repair requiring living inner bark to maintain covering).
Dead_Outer_Trunk_Bark emphasizes complete absence of living cells and is distinguished from Outer_Trunk_Bark (general exterior bark), Protective_Outer_Trunk_Bark (function emphasis), and Peeling_Outer_Trunk_Bark (sloughing behavior) by focus on definitively non-living status. Critical for understanding passive protection mechanisms, cellular death in bark formation, dead tissue structural persistence, contrast between living inner bark and dead outer bark, and importance of maintaining dead outer bark integrity for trunk protection despite inability to self-repair.
Abgestorbene Außenborke
Leblose Schutzrinde
Lifeless protective bark
Tote äußere Rinde
Evert, R. F. (2006). Esau's Plant Anatomy= Meristems, Cells, and Tissues of the Plant Body (3rd ed.). Wiley-Interscience. | Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press.
Dead Outer Trunk Bark
Decline resistance varies among cultivars, with Cabernet Sauvignon and Ugni Blanc showing relative tolerance (5-10% cumulative mortality over 20 years) while Sauvignon Blanc and Semillon are highly susceptible (15-30% mortality). Esca complex (caused by Phaeomoniella chlamydospora, Phaeoacremonium minimum, and Fomitiporia mediterranea) represents the primary decline syndrome in mature vineyards, with symptom expression varying from chronic (tiger-stripe leaf symptoms, progressive weakening) to apoplectic (sudden vine death). Decline-resistant genotypes show enhanced wound compartmentalization, production of antifungal stilbenes (resveratrol, ε-viniferin) in response to infection, and maintained hydraulic function despite partial vascular occlusion. Rootstock effects on scion decline resistance are significant, with 110 Richter and 140 Ruggeri conferring improved decline survival compared to SO4 in some studies.
A vine longevity disposition that is the inherited capacity of a Vitis vinifera plant to resist or tolerate the chronic progressive deterioration known as vine decline, maintaining productive function despite accumulating damage from trunk diseases (Esca complex, Botryosphaeria dieback, Eutypa dieback), vascular dysfunction, and age-related physiological changes.
Decline resistance is increasingly critical as trunk disease pressure intensifies globally and effective chemical controls remain limited. The trait encompasses both resistance (limiting pathogen establishment and spread) and tolerance (maintaining function despite infection). Understanding cultivar-specific decline susceptibility enables targeted prophylactic management and informed replanting decisions. The economic impact of vine decline includes direct replacement costs and indirect losses from reduced vineyard uniformity and quality.
Abbautoleranz
Dekline-Resistenz
Verfallsresistenz
chronic stress tolerance
decline tolerance
trunk disease resistance
Gramaje D et al. (2018) Managing grapevine trunk diseases with respect to etiology and epidemiology: current strategies and future prospects. Plant Disease 102:12-39 Murolo S, Romanazzi G (2014) Effects of grapevine cultivar, rootstock and clone on Esca disease. Australasian Plant Pathology 43:215-221
Decline Resistance Disposition
In a Chianti Classico Sangiovese vineyard, declining accumulation rate was quantified over 28 days in September-October Year N+1 (BBCH 86-89) when weekly Brix measurements showed progressive decline: Week 1:1.8° gain (0.26% per day), Week 2:1.4° (0.20% per day), Week 3:0.9° (0.13% per day), Week 4:0.5° (0.07% per day), demonstrating consistent 22-36% week-over-week reduction, exponential fit dBrix/dt = 0.082 × (26.2 - Brix_current) predicting Brix_max 26.2° (actual harvest 25.8°), enabling harvest decision when rate <0.10% per day threshold achieved week 4, under typical Tuscan autumn conditions.
Declining_Accumulation_Rate represents the progressive reduction in daily or weekly sugar gain velocity during late ripening creating the measurable deceleration that characterizes transition from active accumulation toward plateau defining the kinetic profile of ripening completion and providing quantitative metric for harvest timing optimization. This declining rate occurs throughout final accumulation period (BBCH 86-89), typically days 35-65 post-veraison spanning entire late ripening 25-35 days, and is characterized by daily Brix increase declining continuously from early final 0.25-0.40% per day to late final <0.05-0.12% per day representing 3-6 fold reduction over phase, weekly Brix gain decreasing from 1.8-2.8° per week to <0.35-0.85° per week showing clear trajectory toward cessation, sequential measurements showing negative second derivative d²Brix/dt² < 0 confirming deceleration rather than constant or accelerating rates, and rate constant k in exponential model dBrix/dt = k × (Brix_max - Brix_current) estimated 0.055-0.125 day⁻¹ from curve fitting indicating characteristic decay timescale. Quantitative characteristics of rate decline include percentage reduction week-over-week typically 15-35% per week showing consistent proportional decrease, absolute reduction in weekly gain declining 0.3-0.8° Brix per week sequentially from peak to final, correlation between current Brix and accumulation rate showing strong negative relationship (r² = 0.82-0.94) where higher current Brix predicts lower future rate consistent with approaching saturation, and threshold detection where rates <0.10% per day or <0.7° per week generally indicate approaching plateau within 1-2 weeks useful for harvest timing. Physiological basis for rate decline includes declining concentration gradients as vacuolar hexose rises from 1200-1700 mM early final to 1800-2500 mM late final reducing thermodynamic driving force proportionally, transporter downregulation with gene expression declining 20-40% over final phase reducing import capacity, substrate limitation as phloem delivery decreases 30-50% with canopy senescence and declining photoperiod limiting supply independent of demand, and increasing respiratory consumption proportion rising from 25-35% to 35-50% of import reducing net accumulation efficiency amplifying rate decline beyond import reduction alone. Kinetic analysis demonstrates hexose import flux declining from 55-95 μmol g⁻¹ h⁻¹ early final to 20-45 μmol g⁻¹ h⁻¹ late final correlating with Brix rate decline, net vacuolar accumulation (import minus respiration minus other metabolism) declining 2.5-4.0 fold from 40-70 μmol g⁻¹ h⁻¹ to 12-30 μmol g⁻¹ h⁻¹ directly determining Brix increase rate, and Michaelis-Menten-like saturation kinetics where accumulation rate = Vmax × (Brix_max - Brix_current) / (Km + (Brix_max - Brix_current)) with Km ~2-4° Brix indicating half-saturation occurs when 2-4° below maximum. Temporal pattern within declining phase shows early deceleration (days 35-45, BBCH 86) with rates declining from 0.30-0.45% to 0.18-0.30% Brix per day representing initial slowdown, mid-deceleration (days 45-55, BBCH 87-88) exhibiting further reduction to 0.10-0.20% per day approaching threshold, and late deceleration (days 55-65, BBCH 89) showing minimal rates <0.05-0.12% per day indicating imminent plateau with some cultivars achieving complete cessation while others maintain minimal ongoing accumulation. Environmental modulation of decline trajectory includes temperature affecting both absolute rates and decline slope with warm conditions maintaining higher rates longer then declining more abruptly while cool conditions showing lower rates throughout with more gradual decline, declining solar radiation through autumn reducing source photosynthesis accelerating rate decline independent of berry capacity, water deficit potentially accelerating decline through reduced phloem delivery and metabolic impairment, and disease or environmental stress disrupting normal trajectory potentially causing premature plateau or irregular decline pattern. Cultivar genetic differences demonstrate high-Brix varieties maintaining higher absolute rates longer with decline from 0.35-0.50% to 0.08-0.15% Brix per day over 25-30 days, moderate cultivars showing decline from 0.25-0.40% to 0.05-0.12% over 22-28 days, and lower-Brix types displaying decline from 0.18-0.30% to 0.03-0.08% over 18-25 days reflecting variation in maximum capacity and regulatory set-points governing decline kinetics. Predictive value of rate decline includes linear extrapolation of declining rates enabling harvest date estimation with accuracy improving as plateau approached and trajectory stabilized, recognition that non-linear decline common requiring exponential or polynomial fitting for accurate prediction, threshold-based forecasting where detecting rates <0.10% per day triggers 7-14 day harvest window prediction, and integration with other maturity indicators improving forecast reliability beyond sugar alone. Quality implications include rate decline trajectory revealing ripening health with smooth gradual decline indicating normal physiology while erratic or premature plateau suggesting stress or dysfunction, coordination between sugar rate decline and continuing acid catabolism affecting final balance with excessive acid loss during extended sugar plateau potentially problematic, and rate decline providing objective quantitative metric for harvest decisions supplementing subjective assessments. Harvest timing strategies using rate decline involve establishing target minimum accumulation rate (typically 0.08-0.15% per day) below which harvest initiated, monitoring week-over-week rate reduction detecting accelerating decline signaling approaching plateau, integrating rate decline with absolute Brix achieving both minimum sweetness and declining trajectory ensuring near-maximum sugar, and flexibility recognizing that environmental factors may disrupt ideal trajectory requiring adaptive decisions. Monitoring techniques include frequent Brix sampling 2-3 times per week during late ripening enabling accurate rate calculation, statistical analysis calculating moving average rates reducing measurement noise improving trend detection, derivative analysis computing d²Brix/dt² confirming negative second derivative diagnostic of decline, and curve fitting to exponential or polynomial models enabling extrapolation and maximum Brix estimation. Viticultural management considerations recognize rate decline as physiologically determined and largely unresponsive to intervention, disease control maintaining berry health preventing disruption of normal decline trajectory, irrigation management typically minimal late season allowing natural ripening completion, and harvest logistics coordination with predicted plateau timing based on rate decline analysis ensuring readiness when optimal maturity achieved. Research applications include mathematical modeling of rate decline revealing underlying mechanisms and constraints, comparative studies across cultivars and environments identifying genetic and environmental controls, and predictive algorithm development enabling precision viticulture decision support systems optimizing harvest timing based on quantitative accumulation kinetics.
Declining_Accumulation_Rate is distinguished from constant rates by showing negative second derivative d²Brix/dt² <0, from approaching maximum by focusing on rate measurement rather than distance-to-maximum, and from plateau by representing progressive decline rather than cessation. Quantitative metric: 0.25-0.40% → <0.05-0.12% Brix per day over 25-35 days. Week-over-week reduction typically 15-35%. Duration BBCH 86-89. Predictive value for harvest timing. Declining_Accumulation_Rate represents measurable kinetic deceleration enabling objective maturity assessment.
decelerating sugar gain
dezelerierende Zucker-Zunahme
progressive Raten-Reduktion
progressive rate reduction
reducing accumulation velocity
reduzierende Akkumulations-Geschwindigkeit
slowing Brix increase
verlangsamende Brix-Zunahme
Coombe BG, McCarthy MG (2000) Dynamics and environmental control of grape berry growth Australian J Grape Wine Res 6:73-83 rate decline dynamics Keller M (2020) The Science of Grapevines 3rd Ed pages 227-228 accumulation kinetics Dai ZW, Ollat N, Gomès E, Decroocq S, Tandonnet JP, Bordenave L, Pieri P, Hilbert G, Kappel C, van Leeuwen C, Vivin P, Delrot S (2011) Ecophysiological, genetic, and molecular causes of variation in grape berry weight and composition Plant Physiol 155:519-532 quantitative rate analysis.
Declining Accumulation Rate
Merlot in Bordeaux shows declining division activity during mid-late July Year N+1 at BBCH 73-75, with mitotic index declining from 11% week 3 to 5% week 4 to 2% week 5, cell cycle times extending from 11 hours to 15 hours, progressive cell cycle exit and enlargement onset, auxin levels declining from 220 ng/g to 80 ng/g, cell number approaching 2.1-2.4 million representing 92-96% final value, under oceanic conditions of 22-27°C temperature, typical of this mid-season cultivar transitioning from division to expansion phase
Declining division activity represents the progressive mitotic rate reduction phase during berry cell division when division intensity decreases from peak levels, cell cycle times extend from 10-12 hours to 14-18 hours, increasing proportion of cells exit cycle transitioning to expansion mode, and final cell number approaches completion, proceeding over weeks 3-5 post-fertilization as division phase concludes and expansion phase dominates. This gradual deceleration phase occurs during mid-late summer in mid-late July in temperate regions Year N+1, spanning approximately 1.5-2.5 weeks from peak division decline through completion, corresponding to BBCH 73-75 young fruit to pea-sized berry stages. Declining activity mechanism involves progressive division rate reduction from peak intensity, cell cycle time extension reflecting slowed progression through phases, transition to enlargement onset as cells increasingly commit to expansion rather than division, auxin levels declining from peak reducing proliferative stimulus, and cell number finalization approaching as final divisions complete. Division rate reduction shows mitotic index declining from peak 8-15% to 3-8% during decline phase then 1-3% approaching completion, absolute division rate decreasing from peak 40,000-120,000 cells per day to 10,000-40,000 cells per day during decline, progressive rate decrease correlating with declining auxin and developmental programming, and rate reduction characteristic of transition from exponential to stationary growth. Mitotic index decline dynamics show progressive decrease over weeks 3-5, typical reduction from 12% at peak week 3 to 6% week 4 to 2% week 5, declining index reflecting decreasing proportion of actively cycling cells, and index as convenient division activity measure. Cell cycle time extension shows cycles lengthening from minimal 10-12 hours during peak to 12-14 hours early decline to 14-18 hours late decline, extended times particularly in G1 and S phases, lengthening reflecting reduced proliferative stimulus and cellular transition, and extended cycles reducing division frequency contributing to rate decline. G1 phase extension shows progression time increasing from 3-4 hours at peak to 5-8 hours during decline, cells spending longer in growth phase before S entry commitment, G1 extension providing window for cell fate decisions, and prolonged G1 characteristic of declining proliferative state. S phase extension shows DNA replication slowing from 5-7 hours to 7-10 hours, extended replication reflecting reduced biosynthetic capacity or checkpoint activation, and S phase lengthening contributing to overall cycle extension. Cell cycle exit progression shows increasing proportion of cells completing final division and exiting to G0/G1 expansion state, exit beginning during peak activity week 3 with early-dividing populations, progressive wave of exit spreading through cell populations weeks 3-5, and exit completion establishing expansion-only cell populations. Cell fate transition shows cells deciding between continued division versus expansion commitment, transition correlating with auxin decline and developmental signals, expansion commitment establishing cells in growth rather than proliferative mode, and transition as critical developmental decision. Transition to enlargement onset shows cells increasingly entering expansion mode through water uptake and vacuole development, vacuole formation beginning in cells exiting cycle, progressive enlargement onset across cell populations, and enlargement starting while some cells still dividing creating overlap. Cell enlargement initiation characteristics show water uptake beginning in expansion-committed cells, vacuole development creating central storage compartment, turgor increase driving cell expansion, and enlargement accelerating as cells fully transition from division. Auxin decline effects show hormone levels decreasing from peak weeks 1-3 to declining levels weeks 3-5, reduced auxin weakening proliferative stimulus, declining hormone correlating with division rate reduction, and auxin decrease as major trigger for declining activity. Auxin level dynamics show typical peak 200-400 ng/g fresh weight weeks 2-3, decline to 100-200 ng/g week 4, further reduction to 50-100 ng/g week 5, and declining hormone shifting cells toward expansion over division. Cell number finalization approaching shows progressive accumulation to 85-95% final cell number by decline phase completion week 5, remaining 5-15% cells from final late divisions week 5-6, and cell number approaching permanent established value. Final cell number variation shows continued accumulation during decline adding 15-25% to peak numbers, late divisions from cells completing extended cycles, and cumulative number reflecting total proliferative output. Environmental effects on declining activity include temperature affecting decline rate and transition timing with warm conditions potentially accelerating decline through faster developmental progression, cool conditions extending decline phase through slowed division rate reduction, and temperature as factor affecting transition dynamics. Developmental programming shows genetically controlled division decline independent of environmental factors within normal ranges, cultivar-specific decline timing affecting total division duration, and intrinsic developmental program driving transition. Declining activity assessment methods include mitotic index measurement showing progressive decrease, cell cycle time analysis documenting extension, cell number counting showing slowing accumulation rate, and division marker expression decreasing correlating with activity decline. The process shows three distinguishable aspects including division rate reduction from peak intensity, transition to enlargement onset as expansion begins dominating, and cell number finalization approaching final value. Duration typically 1.5-2.5 weeks representing decline from peak through near-completion. Regional timing shows declining division activity during mid-late July across climates weeks 3-5 post-fertilization. Viticultural significance includes representing transition from division to expansion as growth mechanism, establishing final cell number determining berry size potential, demonstrating progressive developmental transition, and marking shift toward expansion-dominated growth phase. Research applications include analyzing division decline regulation and mechanisms, investigating cell fate transition controls, evaluating environmental effects on decline timing and dynamics, quantifying decline kinetics and cell number finalization, and assessing declining activity as transition to expansion phase.
Declining_Division_Activity represents progressive deceleration phase within Berry_Cell_Division_Phase. Subdivided into Division_Rate_Reduction from peak levels, Transition_to_Enlargement_Onset as expansion begins, and Cell_Number_Finalization approaching final value. Duration 1.5-2.5 weeks representing weeks 3-5. Mitotic index declining from 8-15% to 1-3%. Cell cycle times extending from 10-12h to 14-18h. Auxin decline triggering rate reduction. Approaches 85-95% final cell number. Precedes Division_Phase_Completion. Transition period to expansion-dominated growth.
Proliferationsreduktion
Teilungs-Dezeleration
abnehmende Teilungsaktivität
division deceleration
division slowdown
mitotic decline
mitotischer Rückgang
proliferation reduction
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 437-440 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Declining Division Activity
Merlot in Bordeaux shows declining pollen viability during late anthesis late June Year N+1 at BBCH 67-68 over 2-3 days, with fresh pollen from late-opening flowers showing 62-74% germination compared to peak 76-88%, aged pollen 48-72 hours old showing 12-24% viability, substantially constraining late pollination success reducing fruit set to 24-34% for late flowers compared to 54-64% during peak, under oceanic conditions of 21-26°C temperature, 58-72% relative humidity, typical of this mid-season cultivar showing moderate pollen longevity
Declining pollen viability represents the progressive deterioration of pollen functional capacity during late anthesis as aged pollen from earlier opening loses germination ability and fresh pollen from late-opening flowers shows reduced quality, substantially constraining late pollination success and reproductive opportunity. This critical male function decline occurs during late anthesis in mid-late June in temperate regions Year N+1, spanning 2-3 days as pollen viability drops from peak levels 70-90% germination to late levels 20-40% fresh pollen and 5-15% aged pollen, severely limiting final pollination success. Viability decline pattern shows characteristic temporal progression with fresh pollen from recently dehisced anthers maintaining moderate 60-80% viability though reduced from peak 70-90%, pollen aged 24-48 hours showing substantially reduced 20-40% viability, and pollen aged 48-72+ hours showing minimal 5-15% viability approaching complete loss of function. Pollen aging mechanisms involve progressive deterioration of cellular structures including membrane degradation compromising cellular integrity, protein denaturation particularly affecting enzymes required for germination and tube growth, DNA damage accumulating in vegetative and sperm cell nuclei potentially affecting fertilization, and depletion of stored energy reserves limiting capacity to support germination. Environmental effects accelerate viability decline with high temperature above 30°C causing rapid deterioration through protein denaturation and membrane damage, low humidity below 40% causing excessive desiccation destroying cellular function, high humidity above 75% promoting fungal colonization on pollen surfaces, and UV radiation potentially damaging DNA and proteins in exposed pollen. Pollen viability assessment shows multiple methods including in vitro germination tests placing pollen on artificial medium measuring germination percentage providing quantitative assessment, tetrazolium staining indicating metabolic activity through enzymatic reduction creating colored product, and fluorescent dyes distinguishing viable from nonviable pollen through membrane integrity assessment. Germination capacity decline shows that while peak-period pollen germinates readily within 1-2 hours producing vigorous tubes, late-period pollen shows delayed germination requiring 3-6 hours, reduced germination percentage with only 20-40% pollen producing tubes, and abnormal tube growth with twisted or stunted tubes indicating cellular dysfunction. Pollen tube growth from declining-viability pollen shows reduced vigor with slower elongation rates 0.5-1.5 mm per hour compared to 1-3 mm per hour with fresh viable pollen, increased probability of growth cessation before reaching ovary failing to deliver sperm cells, and potential for abnormal fertilization outcomes if defective pollen does reach embryo sac. Pollination success probability drops substantially with declining viability showing correlation between pollen germination percentage and fruit set rate, with 70-90% viable pollen supporting 50-70% fruit set during optimal window declining to 20-40% viable pollen supporting only 15-30% fruit set during late anthesis. Pollen availability interactions compound viability effects with reduced pollen production during late anthesis combined with declining viability creating severely constrained pollination opportunity, limiting reproductive success for late-opening or previously unpollinated flowers. Atmospheric pollen concentration declines during late anthesis through combined effects of reduced production from fewer anthers dehiscing, loss of viability in airborne pollen reducing functional pollen density, and dilution through continued air circulation dispersing remaining pollen, reducing pollination probability. Fresh pollen production during late anthesis continues from recently opened flowers providing limited renewed pollen source, though production levels may be reduced compared to peak period and pollen quality may show intrinsic reduction reflecting late developmental status. Cultivar differences in pollen viability characteristics show some varieties producing robust pollen maintaining 70-85% viability during late anthesis supporting adequate late pollination, while others show poor pollen with 40-60% viability during late period substantially limiting reproductive success. Boron nutrition effects on pollen function intensify during late period with adequate boron essential for maintaining germination and tube growth capacity in declining-viability pollen, while deficiency exacerbates viability problems potentially eliminating late pollination success. Temperature during late anthesis critically affects viability dynamics with optimal 22-28°C maintaining adequate though declining pollen function, cool conditions below 18°C further reducing viability and germination, and warm conditions above 30°C accelerating decline potentially eliminating late reproductive opportunity. Humidity during late anthesis affects pollen viability with moderate 45-65% maintaining adequate hydration without promoting fungal growth, while humidity extremes accelerate decline through desiccation or microbial colonization. Resource allocation shifts during late anthesis redirect vine resources from reproductive investment toward developing berries, potentially contributing to reduced pollen quality through decreased resource availability for late anther development and pollen maturation. Metabolic changes in declining-viability pollen show reduced respiration indicating cellular deterioration, depleted ATP levels limiting energy availability for germination, and accumulated cellular damage from oxidative stress or other aging processes. Hormonal regulation of pollen viability involves possible role for stress hormones including abscisic acid potentially affecting pollen longevity, though primary determinants are intrinsic aging processes and environmental stresses. Practical implications of declining viability include substantially reduced late fruit set from late-opening flowers unable to achieve adequate pollination, increased probability of incomplete fruit set in extended flowering conditions where late opening predominates, and possible fruit set failures in cultivars with inherently poor late pollen viability. Management considerations include recognizing that late anthesis conditions have diminishing impact on overall fruit set since majority of berries 75-85% already established during peak, understanding that cultivars with extended flowering may show reduced overall fruit set from prevalence of declining-viability pollen, and appreciating that boron nutrition adequacy critically affects late pollen function maintaining some reproductive capacity during viability decline. The process represents second of three late anthesis aspects focusing on male reproductive capacity decline, occurring concurrently with final flowers opening and contributing to pollination window closure. Duration spans 2-3 days of late anthesis representing progressive viability deterioration, with rapid decline under unfavorable conditions or slower decline under optimal conditions. Cultivar variation in viability decline shows some varieties maintaining adequate pollen function through late anthesis supporting reasonable late fruit set, while others show rapid severe decline substantially limiting late reproductive success and potentially creating yield limitations. Regional timing shows declining pollen viability during late anthesis period mid-late June in warm climates, late June in temperate regions, and late June-early July in cool areas, affecting final 2-3 days of flowering period. Viticultural significance includes representing major constraint on late pollination success and final fruit set contribution, emphasizing importance of compressed flowering period concentrating reproductive activity during optimal pollen viability window, illustrating vulnerability of extended flowering to poor fruit set from declining pollen quality, and demonstrating critical role of pollen function in determining reproductive success and yield. Research applications include analyzing pollen aging mechanisms including cellular and molecular deterioration processes, evaluating environmental effects on viability dynamics affecting pollination outcomes, investigating cultivar differences in pollen longevity affecting reproductive reliability, quantifying relationships between viability and fruit set success, and assessing management practices particularly boron nutrition affecting pollen function maintenance.
Declining_Pollen_Viability is second of three Late_Anthesis aspects, distinguished by progressive deterioration of male reproductive capacity through pollen aging. Occurs concurrently with Final_Flowers_Opening. Represents major constraint on late pollination success limiting final fruit set contribution. Environmental conditions particularly temperature and humidity accelerate decline. Boron nutrition critically affects maintenance of residual pollen function. Cultivar differences in pollen longevity significantly affect late reproductive success.
Pollenalterung
Pollenverschlechterung
abnehmende Pollenviabilität
declining pollen quality
pollen aging
pollen deterioration
reduced pollen viability
sinkende Pollenqualität
Declining Pollen Viability
Merlot in Bordeaux shows declining shoot elongation from early July through early August at BBCH 35-38 over 4-5 weeks, with shoots extending from 1800 mm following fruit set to 2200-2500 mm final length through elongation rate declining from 28-35 mm per day to 12-18 mm per day as berry development progresses, under oceanic climate conditions of 22-26°C mean temperature, decreasing precipitation requiring supplemental irrigation, and moderate fertility with nitrogen becoming redistributed to fruit typical of this moderately vigorous mid-season cultivar
Declining shoot elongation represents the decelerating growth phase where elongation rate progressively decreases from peak levels as environmental limitations, resource competition from developing fruit, or developmental programming engage growth reduction mechanisms, characterized by diminishing daily shoot extension transitioning toward eventual growth cessation. This developmental stage typically occurs during July in temperate regions corresponding approximately to BBCH 35-38 as fruit develops from pea-size through beginning of berry touch, with elongation rate declining from 30-50 mm per day to 10-25 mm per day over 3-5 week period creating shoot length increase of 400-800 mm before growth largely ceases. Declining elongation represents deceleration phase of sigmoid growth curve where growth rate progressively reduces as multiple factors including high fruit demand for assimilates, possible water or nutrient limitations, increasing temperature stress, or developmental signals engage processes reducing meristematic activity and cell expansion rates. The apical meristem continues producing new leaf primordia but at reduced frequency of 3-4 days between new primordia initiation, with cell division rates slowing and meristem size potentially reducing as growth regulatory signals shift developmental priorities. Internode lengths during declining elongation progressively shorten from 60-100 mm early in phase to 30-60 mm later reflecting reducing resource availability for elongation and changing hormonal environment, with final internodes before growth cessation typically quite short 15-30 mm. Cell expansion in subapical zones operates at progressively reducing rates as turgor pressure declines through increased tissue water stress or hormonal signals reducing cell wall extensibility, with expansion zone potentially shortening to 15-25 mm behind apex and cells achieving smaller final dimensions of 150-400 micrometers. Leaves continue emerging but at slower rate and expanding to progressively smaller final dimensions, with late-season leaves typically 50-70% size of mid-season leaves reflecting reducing resource availability and changing developmental state. Photosynthetic capacity maintains high levels in mature leaves though proportion of production allocated to shoot growth declines substantially as fruit demand dominates assimilate partitioning, with reproductive sink strength typically exceeding vegetative demand creating competition that contributes to growth decline. Carbon allocation during declining elongation shifts dramatically from approximately equal vegetative-reproductive partitioning to 70-85% fruit allocation as developing clusters exert dominant sink strength, with shoot growth consuming only 20-30% photosynthetic production and remainder supporting fruit development, root growth, and increasing reserve accumulation. Vascular system maintains functionality though new vessel and sieve element production slows, with water transport capacity of 3-5 mg per second adequate for reduced growth requirements and maintained transpiration from mature canopy. Water demand remains substantial at 400-600 mg per hour from mature canopy though growth component declines, with tissue water status often declining during this period in many regions as mid-summer heat and possible water limitation create moderate water stress contributing to growth reduction. Nutrient demand shifts predominantly to fruit with nitrogen particularly redirected from vegetative to reproductive tissues, with shoot growth nitrogen requirements declining while fruit accumulates nitrogen for protein synthesis and other developmental processes. Metabolic activity in shoot apical regions remains elevated though declining from peak levels, with respiration in expanding tissues 6-10 fold above mature levels declining as growth rate reduces, while mature leaf photosynthesis maintains high efficiency supporting fruit development. Protein synthesis continues in apical growing zones though at reduced rates, with declining production of cell cycle proteins and wall synthesis enzymes reflecting reduced growth while maintenance proteins and reserve storage proteins increase in mature tissues. Hormonal regulation shows declining auxin production from slowing shoot apex reducing growth-promoting signals, increasing abscisic acid from water-stressed tissues or fruit development promoting growth reduction, possible declining cytokinin supply from roots under stress or developmental change, and ethylene potentially increasing promoting growth cessation processes. Temperature during declining elongation typically includes warmest conditions of growing season with means often 24-30°C in many regions, with high temperatures contributing to growth reduction through increased respiration reducing carbon balance efficiency and exacerbating water stress effects. Light intensity remains high during mid-summer though increasing canopy density may reduce light penetration to lower leaves limiting whole-canopy photosynthetic efficiency, while high light on exposed tissues can create photo-inhibition stress. Water availability often becomes limiting during declining elongation in many regions as mid-late summer typically coincides with reduced precipitation and high evaporative demand, with moderate water stress contributing beneficially to growth reduction while severe stress risks premature leaf senescence and compromised fruit development. Nutrient availability particularly nitrogen becomes redistributed with mobile nutrients moving from vegetative tissues toward fruit, with continued excessive nitrogen availability potentially prolonging vegetative growth undesirably competing with fruit maturation. The process typically spans 3-5 weeks under normal conditions with progressive environmental stress and fruit sink development, with duration varying based on severity of limiting factors and cultivar growth habit characteristics. Cultivar variation appears in rate of decline from peak growth, final shoot length achieved before cessation, timing of growth cessation relative to fruit development stages, and sensitivity to environmental stresses affecting decline rate. Regional timing shows declining elongation occurring July in warm climates, July-August in temperate regions, and August in cool areas, typically spanning early berry development through beginning color change period when fruit demands are increasing substantially. Viticultural management considerations include using moderate water stress to promote timely growth decline improving fruit development conditions, avoiding excessive late nitrogen maintaining vegetative growth competing with fruit quality, implementing shoot trimming removing excessive length and lateral growth managing canopy density, monitoring canopy condition assessing stress levels and fruit development progress, and adjusting irrigation and nutrition management based on growth decline patterns and fruit development requirements. Premature severe growth decline from excessive stress risks insufficient photosynthetic capacity for fruit maturation and reserve accumulation, while prolonged excessive growth from over-supply of resources creates unfavorable fruit microclimate and delays maturation. Research applications include analyzing mechanisms of growth decline including hormonal regulation and source-sink dynamics, quantifying assimilate partitioning shifts during vegetative-reproductive competition, evaluating management strategies optimizing growth cessation timing for quality outcomes, modeling growth patterns for decision support systems, and assessing genetic variation in growth habit affecting management requirements and fruit quality potential.
Declining_Shoot_Elongation is fourth of five Shoot_Elongation subphases, distinguished by progressive reduction in growth rate from peak levels as limitations engage. Transitions to Terminal_Shoot_Elongation as growth approaches cessation. Represents critical phase where resource allocation shifts decidedly toward fruit development. Timing and management of decline substantially affect fruit quality outcomes through effects on canopy-fruit balance.
Verlangsamungsphase
Wachstumsrückgang
abnehmende Triebstreckung
decelerating growth
declining vegetative growth
growth reduction phase
slowing elongation phase
verlangsamte Wachstumsphase
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 248-258 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 165-175 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Declining Shoot Elongation
A deep basal lobe representing the left basal lobe on a mature leaf at node 7 of Vitis vinifera cv. Pinot Noir in Burgundy, France, exhibiting pronounced development with dimensions of 3.5 cm width and 2.8 cm length approaching lower lateral lobe size, deeply separated from lower lateral lobe by lower left lateral sinus of 2.2 cm depth creating clear lobe boundary, lobe extending 1.8 cm beyond the level of petiolar junction creating substantial basal extension, vascularized by prominent basal primary lateral vein 2.2 mm diameter at median vein junction, lobe oriented with strongly convergent inward direction bringing margin across the petiolar sinus, basal lobe margin crossing anterior to the right basal lobe margin by 0.6 cm creating the characteristic overlapping petiolar sinus diagnostic for Pinot varieties, lobe exhibiting rounded apex without sharp point, lobe area of 8.2 cm² representing 12% of total blade area approaching lower lateral lobe contribution (14%), prominent basal lobe development creating broad leaf base and distinctive overlapped petiolar sinus architecture, measured at BBCH stage 75 (berries pea-sized) in late June, representing maximum deep basal lobe expression characteristic of Pinot Noir and serving as definitive diagnostic feature.
A basal lobe exhibiting pronounced development with substantial size and depth of dissection in Vitis vinifera, characterized by well-developed lobe structure with dimensions approaching or occasionally exceeding lower lateral lobe size, deep separation from adjacent lateral lobes through lower lateral sinuses of substantial depth typically 1.5-3.0 cm, elongated or broad morphology extending substantially beyond the petiolar junction level, strong vascularization by prominent basal primary lateral veins, convergent orientation with lobe margins directed inward toward the petiolar sinus, and critical role in determining petiolar sinus configuration typically creating closed or overlapping sinus patterns through convergent or crossing basal lobe margins. Deep basal lobes represent maximum basal lobe expression, occurring in cultivars with broad leaf bases and pronounced lobe dissection (Pinot Noir, Gamay, some Grenache clones), creating distinctive leaf base architecture with prominent basal lobes flanking and often occluding the petiolar junction, and serving as important diagnostic features through effects on overall leaf base shape and particularly petiolar sinus configuration.
Deep basal lobes are distinguished from shallow basal lobes by substantial size, pronounced separation, and prominent development. The deep designation emphasizes degree of lobe expression rather than necessarily referring to sinus depth, though deep separation from lateral lobes typically accompanies deep basal lobe development. Size approaching or occasionally exceeding lower lateral lobes represents unusual subordination reversal, with most cultivars showing basal lobes as smallest structures but deep basal lobe cultivars showing enhanced basal development. Convergent orientation is characteristic, with deep basal lobes typically directed inward creating closed or overlapping petiolar sinuses, contrasting with divergent orientation of shallow basal lobes creating open sinuses. The overlapping petiolar sinus configuration in Pinot varieties results directly from deep convergent basal lobes crossing at the petiolar junction, representing the definitive example of deep basal lobe-petiolar sinus interaction. Occurrence is restricted primarily to cultivars with broad leaf bases and pronounced overall lobe dissection, with narrow-based cultivars universally showing shallow or absent basal lobes. Functional contribution involves meaningful area provision typically 20-30% combined area unusual for basal lobes, and critical mechanical integration at the petiole-blade junction potentially enhanced by robust basal lobe development. Ampelographic utility is exceptional through definitive effects on petiolar sinus configuration, with deep convergent basal lobes creating the overlapping sinus serving as primary diagnostic character.
ausgeprägte Petiolarlappen
prominent basal lobe
prominenter Basislappen
pronounced petiolar lobe
tiefer Basallappen
well-developed base lobe
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 58-119 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 79-118.
Deep Basal Lobe
In Zweigelt vines in Burgenland, Austria, deep dormancy entry is confirmed in mid-November when excised buds placed at 22°C with 16-hour photoperiod show no growth response after 4 weeks of observation, with parallel measurements showing maximum ABA concentrations of 200 ng per gram fresh weight, high VvDAM gene expression, and the beginning of chilling unit accumulation at field temperatures below 7.2°C marking the onset of the endodormancy period at BBCH stage 93.
Deep dormancy entry in Vitis vinifera represents the final phase of dormancy establishment during which buds transition into the fully endodormant state characterized by complete unresponsiveness to growth-promoting conditions and the requirement for specific chilling exposure before growth capacity can be restored. This transition typically occurs during November in Northern Hemisphere viticulture, marking the point at which buds have accumulated sufficient dormancy factors that even extended exposure to warm temperatures and optimal growing conditions fails to induce budbreak. The physiological basis of deep dormancy involves multiple reinforcing mechanisms including high concentrations of growth inhibitors particularly abscisic acid, low levels of growth promoters including cytokinins and gibberellins, chromatin modifications that silence growth-related genes, and the presence of dormancy-maintaining proteins that must be degraded before growth can resume. The molecular characteristics of deep dormancy include high expression of dormancy-associated MADS-box genes whose products actively maintain the dormant state, altered plasmodesmatal function that limits symplastic communication and signal transmission, and metabolic configurations that do not support the biosynthetic activities required for growth resumption. Testing for deep dormancy entry involves exposing buds to growth-promoting conditions and observing the failure of budbreak that confirms full endodormancy, with deeply dormant buds showing no growth response even after extended periods of favorable temperatures that would rapidly induce growth in non-dormant tissues. The depth of dormancy achieved during deep dormancy entry varies among cultivars, with some genotypes achieving very deep dormancy requiring 1000-1500 chilling hours for release while others achieve shallower dormancy requiring only 400-700 chilling hours, reflecting genetic variation in dormancy regulation. The transition into deep dormancy serves the ecological function of preventing premature growth resumption during temporary warm periods in autumn and early winter, which would expose sensitive growing tissues to subsequent freezing temperatures with potentially lethal consequences. Climate warming poses challenges to deep dormancy entry in some regions, as elevated autumn temperatures may delay or reduce dormancy depth, potentially leading to inadequate dormancy and erratic spring growth patterns.
Deep_Dormancy_Entry represents the culminating phase of Dormancy_Establishment marking achievement of full endodormancy. Distinguished from earlier dormancy phases by the complete lack of growth response to favorable conditions, confirming transition from dormancy induction to established endodormancy. Characterized by high ABA, high DAM gene expression, and growth unresponsiveness. Marks beginning of chilling requirement period. Dormancy depth varies among cultivars from 400-1500 chilling hours required for release. Corresponds to full achievement of BBCH stage 93.
Endodormanz-Eintritt
Tiefschlaf-Eintritt
deep rest entry
endodormancy entry
full dormancy achievement
tiefe Ruhe-Etablierung
true dormancy establishment
volle Dormanz-Erreichung
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Lang GA et al (1987) Endo-, para-, and ecodormancy: physiological terminology and classification for dormancy research. HortScience 22: 371-377 Rohde A, Bhalerao RP (2007) Plant dormancy in the perennial context. Trends in Plant Science 12: 217-223
Deep Dormancy Entry
In Cabernet Sauvignon vines in the Napa Valley, California, deep endodormancy occurs during December and early January when forcing tests show budbreak requiring more than 50 days at 22°C with many buds failing to break entirely, ABA concentrations in buds remain at maximum levels of 180-220 ng per gram fresh weight, and cold hardiness reaches seasonal maximum values of -18 to -20°C for primary buds, before chilling accumulation progressively begins reducing dormancy depth through January.
Deep endodormancy in Vitis vinifera represents the period of maximum dormancy intensity occurring during early to mid-winter when the internal factors inhibiting bud growth are at their strongest and buds exhibit complete unresponsiveness to any growth-promoting stimuli. This phase typically occurs during December and early January in Northern Hemisphere viticulture, following the completion of dormancy establishment and before significant chilling accumulation has begun to reduce dormancy depth. During deep endodormancy, buds represent the most dormant condition achieved in the annual cycle, with all growth-inhibiting mechanisms operating at maximum intensity including peak abscisic acid concentrations, highest expression levels of dormancy-associated MADS-box genes, and most repressive chromatin configurations at growth-related loci. The practical definition of deep endodormancy relies on forcing tests showing that buds require the longest time to budbreak or completely fail to break when exposed to growth-promoting conditions, in contrast to earlier or later dormancy phases when forcing response is faster or more complete. The duration of deep endodormancy varies among cultivars in relation to their overall dormancy depth, with deeply dormant cultivars adapted to continental climates maintaining deep endodormancy for longer periods than shallowly dormant cultivars from Mediterranean regions. Physiologically, deep endodormancy is maintained by the convergence of multiple inhibitory factors that must be progressively dismantled through chilling exposure before growth competence can be restored, with the requirement for multiple changes explaining why dormancy release is gradual rather than abrupt. The metabolic state during deep endodormancy is characterized by minimal activity, with respiration rates at annual minimum values, protein synthesis essentially ceased except for maintenance functions, and cellular processes largely restricted to survival functions rather than growth preparation. Cold hardiness typically reaches maximum levels during deep endodormancy, reflecting the full expression of cold acclimation processes and the absence of any growth activity that would reduce freezing tolerance. The transition from deep endodormancy to progressive dormancy release begins as chilling accumulation starts to reduce dormancy depth, though this transition may not be externally visible and is detected through changes in forcing test response times.
Deep_Endodormancy represents the initial phase of maximum dormancy intensity within Endodormancy. Distinguished from later phases where chilling accumulation has begun reducing dormancy depth. Subdivides into Maximum_Dormancy_Depth, Growth_Inhibition_Maximum, and ABA_Maximum_Level as component characteristics. Duration typically spans 4-8 weeks during early winter. Characterized by longest forcing times and lowest budbreak percentages in forcing tests. Cold hardiness at seasonal maximum during this phase.
Dormanz-Maximum
deepest dormancy
full endodormancy
maximale Dormanz
maximum dormancy
peak dormancy
tiefe Endodormanz
volle Endodormanz
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Lang GA et al (1987) Endo-, para-, and ecodormancy: physiological terminology and classification for dormancy research. HortScience 22: 371-377 Lavee S, May P (1997) Dormancy of grapevine buds - facts and speculation. Australian Journal of Grape and Wine Research 3: 31-46
Deep Endodormancy
Cabernet Sauvignon grafted on 110 Richter rootstock at 18 years in deep sandy loam showing deep lateral secondary roots emerging from primary roots at 90-140 cm depth, excavation revealing first-order deep laterals 2.4-3.8 cm diameter extending 60-110 cm horizontally at depths 105-145 cm, deep laterals accessing moist soil (14-18% volumetric water content) at depth during August drought when upper 90 cm soil severely dry (<6% VWC), soil core sampling indicating deep lateral fine root density 0.8-1.4 cm/cm³ soil at 110-140 cm depth providing water uptake, deep lateral network characteristic of 110R vigorous deep-rooting enabling drought tolerance
Deep lateral secondary root is lateral secondary root growing at substantial depth typically exceeding 80-100 cm depth and often reaching 100-150 cm or deeper, emphasizing depth distribution rather than branching order or structure, characterized by lateral orientation but at deep position in soil profile, emergence from deep primary roots or deep first-order laterals enabling deep lateral exploration, and access to deep soil resources including moisture, nutrients, and stable temperatures unavailable to shallow lateral roots, with deep lateral secondary roots serving specialized function of lateral soil exploration at depth complementing shallow lateral network near surface and deep vertical primary root penetration creating three-dimensional soil resource exploitation, particularly important during drought when surface soil dry and deep laterals accessing persistent moisture, and demonstrating rootstock genetic control with deep-rooting varieties producing extensive deep lateral networks while shallow-rooting types showing limited deep lateral development. Deep lateral secondary root depth definition includes= located below 80-100 cm (deep lateral secondary roots by definition positioned deeper than typical lateral root concentration zone), commonly at 100-150 cm depth (deep laterals frequently in this range in moderate to vigorous rootstocks in permeable soils), potentially exceeding 150 cm (deep laterals from very deep primary roots possibly reaching 150-200+ cm in favorable conditions), emerging from deep parent roots (deep lateral secondary roots originating from deep primary roots or deep first-order laterals enabling deep position), and exploring deep soil layers (deep laterals accessing C horizon or deeper B horizon unavailable to shallow root network). Deep lateral secondary root genetic determination includes= characteristic of deep-rooting rootstocks (Berlandieri-based varieties like 110R, 140Ru, 1103P showing well-developed deep lateral networks), typical of vigorous cultivars (high-vigor rootstocks producing both deep primary roots and extensive deep lateral branching), limited in shallow-rooting types (Riparia-based rootstocks showing few deep laterals due to shallow primary root systems), demonstrating breeding objectives (deep lateral capacity selected in drought-tolerant rootstock development programs), and showing hybrid variation (rootstock hybrids exhibiting intermediate deep lateral development reflecting parental genetic contributions). Deep lateral secondary root soil requirements includes= requiring deep permeable profile (deep lateral development needing soil without restrictive layers preventing deep root growth), favored by coarse textures (sandy or loamy soils facilitating deep lateral penetration and growth), inhibited by shallow limitations (bedrock, hardpans, water tables, or toxic layers preventing deep lateral development), depending on deep aeration (adequate oxygen at depth necessary for deep lateral root respiration and function), and enabled by deep moisture (persistent deep moisture supporting deep lateral growth and function even when surface dry). Deep lateral secondary root drought tolerance function includes= accessing deep persistent moisture (deep laterals potentially reaching water unavailable to shallow roots enabling continued function during surface drought), enabling drought avoidance (deep lateral network complementing deep tap root in accessing deep water allowing vine to avoid rather than tolerate drought stress), contributing summer water supply (deep laterals particularly important mid to late summer when surface soil exhausted), demonstrating seasonal importance (deep lateral contribution to total vine water increasing progressively through growing season as shallow soil dries), and providing survival advantage (deep lateral network critical for vine survival during extended drought periods in unirrigated systems). Deep lateral secondary root resource exploitation includes= exploring deep soil volume (deep laterals accessing three-dimensional soil space unavailable to shallow roots maximizing total resource acquisition), intercepting deep percolation (deep laterals positioned to capture water moving below shallow root zone during heavy rainfall or irrigation), accessing deep nutrients (deep laterals exploiting nutrients accumulated at depth through leaching or rock weathering), avoiding surface competition (deep laterals accessing resources unavailable to shallow-rooted weeds or cover crops), and showing complementary function (deep laterals accessing different resources than shallow laterals creating niche differentiation). Deep lateral secondary root temperature advantages includes= experiencing stable temperatures (deep soil showing minimal daily fluctuation providing consistent environment for deep lateral function), avoiding surface heat stress (deep laterals protected from extreme surface temperatures during heat waves), maintaining function during surface freeze (deep laterals below frost line continuing water uptake when shallow roots frozen), showing year-round activity (deep laterals potentially maintaining growth and absorption throughout winter in regions with surface soil freezing), and demonstrating thermal refugia (deep soil providing favorable temperature when surface too hot or cold). Deep lateral secondary root structural characteristics includes= showing moderate diameter (deep lateral secondary roots typically 1-4 cm reflecting first or second-order status), exhibiting secondary growth (deep laterals developing woody structure with xylem rings and bark), demonstrating sparse branching (deep laterals often showing lower branching density than shallow laterals possibly due to lower deep soil fertility), displaying longer internodes (spacing between higher-order branches potentially greater on deep laterals than shallow laterals), and revealing anatomical adaptations (deep lateral anatomy potentially showing larger vessels or other features reflecting deep environment). Deep lateral secondary root in vine water relations includes= buffering drought stress (deep lateral network moderating plant water potential during surface soil drought), enabling sustained photosynthesis (deep lateral water supply supporting continued stomatal opening and carbon assimilation under drought), improving crop quality (deep lateral contribution to vine water helping complete fruit ripening under stress), reducing irrigation needs (well-developed deep lateral network potentially reducing or eliminating irrigation requirements in some climates), and demonstrating cultivar water strategy (deep lateral extent reflecting rootstock drought adaptation strategy with deep-rooted types showing extensive deep lateral networks).
Deep_Lateral_Secondary_Root emphasizes deep soil position exceeding 80-100 cm depth and is distinguished from shallow laterals by focus on deep resource access. Critical for understanding deep soil exploration, drought avoidance, deep moisture access, three-dimensional resource exploitation, Berlandieri genetics, and the deep-positioned lateral roots complementing vertical tap root creating deep lateral soil exploration.
Deep-position lateral root
Lower profile lateral
Subsoil lateral root
Tiefe Lateralwurzel
Tieflagige Verzweigungswurzel
Unterboden-Seitenwurzel
Deep Lateral Secondary Root
A deep lower lateral sinus representing the right lower lateral sinus on a mature leaf at node 7 of Vitis vinifera cv. Syrah in the Rhône Valley, France, positioned between upper lateral and lower lateral lobes, exhibiting pronounced depth of 2.8 cm measured from sinus base to lobe edge connection line representing substantial dissection for lower blade region, penetration reaching approximately 48% of distance from margin toward median vein, well-defined U-shaped contour with clearly rounded sinus base, width of 1.3 cm at opening creating depth-to-width ratio of 2.15 indicating depth substantially exceeding width, sinus creating clear prominent separation between upper and lower lateral lobes contributing to well-defined five-lobed architecture, sinus base positioned 6.2 cm from petiolar junction in mid-blade region, sinus margins showing distinct boundaries with smooth curvature, deep lower sinus complementing deep upper lateral sinus (3.4 cm) creating overall deeply dissected blade characteristic of this cultivar, bilateral near-symmetry with left lower lateral sinus showing similarly deep configuration (2.7 cm), measured at BBCH stage 81 (beginning of berry ripening) in early August, representing pronounced deep lower lateral sinus development typical of deeply dissected Syrah morphology.
A lower lateral sinus exhibiting pronounced depth creating substantial separation between lower lateral lobes and adjacent lobes in Vitis vinifera, characterized by depth typically exceeding 2-2.5 cm representing substantial dissection for lower blade regions, well-defined configuration with distinguishable sinus base, penetration often reaching 40-60% of distance from margin toward median vein, clear visual prominence creating obvious lower lateral lobe separation, occurrence primarily in deeply dissected five-lobed or seven-lobed cultivars showing pronounced overall lobe development, and role in creating well-differentiated lower lateral lobes characteristic of deeply dissected cultivar morphology. Deep lower lateral sinuses represent maximum expression of lower sinus development, occurring in cultivars with strong overall lobe dissection including deeply lobed Syrah clones, Mourvèdre, and seven-lobed configurations in some cultivars, creating distinctive architecture with clearly separated lower lateral lobes contributing to overall deeply dissected appearance, and serving as diagnostic features indicating deeply lobed cultivar groups particularly when combined with deep upper lateral sinuses.
Deep lower lateral sinuses are distinguished from shallow lower lateral sinuses by depth typically >2-2.5 cm and well-defined configuration. The deep designation emphasizes pronounced dissection in lower blade regions contrasting with typical shallow or absent lower sinuses in most cultivars. Depth approaching or occasionally exceeding upper lateral sinus depth is unusual, with most cultivars showing decreasing dissection depth from upper to lower regions, making deep lower sinuses particularly distinctive. Penetration percentage assessment accounts for position-specific depth expectations, with >40-50% considered deep for lower regions compared to >50-60% for upper sinuses. Well-defined configuration with clear sinus base distinguishes deep from shallow poorly defined lower sinuses. Occurrence is restricted to deeply dissected cultivar groups, with moderately and weakly lobed varieties universally showing shallow or absent lower sinuses. Correlation with deep upper lateral sinuses is very strong, with cultivars showing deep lower sinuses virtually always also exhibiting deep upper sinuses creating overall pronounced dissection. Genetic control is strong with deep dissection patterns consistent across cultivars showing this character. Environmental sensitivity is moderate with deep lower sinuses potentially showing more modification than upper sinuses under extreme vigor variations. Functional implications are minimal given lower blade position. Ampelographic utility is moderate as secondary feature confirming deeply dissected cultivar group membership, though less diagnostic than upper sinus or petiolar sinus characteristics.
ausgeprägte untere Lateralbucht
deeply incised middle sinus
profound lower sinus
pronounced lower lateral sinus
tief eingeschnittene mittlere Sinus
tiefe untere Seitenbucht
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 90-135 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 79-118.
Deep Lower Lateral Sinus
Field investigation of Deep_Rooting_Disposition at INRAE Montpellier employed soil coring and root excavation across 12 rootstocks in a 15-year-old vineyard on deep alluvial soil to characterize maximum rooting depth and vertical root distribution. Root presence was assessed in 20 cm depth increments to 4 m depth, with root length density quantified in each layer. Deep_Rooting_Disposition varied dramatically among rootstocks. '140 Ruggeri' (V. berlandieri × V. rupestris) showed exceptional deep rooting with live roots detected to 3.8 m depth and 35% of total root length below 1.5 m, reflecting its breeding origin from drought-adapted parents on limestone soils of southern France. 'Riparia Gloire' (V. riparia selection) exhibited minimal Deep_Rooting_Disposition with maximum depth of 1.1 m and only 8% of roots below 0.5 m, consistent with its origin in riparian habitats with reliable surface water. Hybrid rootstocks showed intermediate patterns, with '110 Richter' reaching 2.6 m and 'SO4' limited to 1.6 m. Soil water extraction monitoring using neutron probe revealed that deep-rooting '140 Ruggeri' extracted water from 2-3 m depth during late-season drought while shallow '3309C' extraction was confined to the upper meter. This deep water access enabled '140 Ruggeri'-grafted vines to maintain 25% higher midday water potential during August drought (-1.2 versus -1.6 MPa), demonstrating the functional significance of Deep_Rooting_Disposition for drought adaptation.
A Deep_Rooting_Disposition is a Root_System_Drought_Adaptation_Disposition that inheres in a Vitis plant root system and is realized through the capacity to extend roots to substantial soil depths, accessing water reserves in subsoil layers that remain available when surface soils are depleted during drought. This disposition represents a drought avoidance mechanism operating through spatial exploitation of soil water resources, with deep-rooted vines able to maintain water uptake by shifting extraction to progressively deeper soil layers as surface moisture is exhausted. Deep rooting capacity depends on inherent root growth vigor, gravitropic response directing root extension downward, ability to penetrate compact subsoil layers, and tolerance of subsoil conditions (lower oxygen, higher bulk density, potentially different pH and mineral content). Grapevine rootstocks vary substantially in Deep_Rooting_Disposition, with maximum rooting depths ranging from less than 1 m in shallow-rooting genotypes to over 6 m in deep-rooting types under favorable soil conditions. Deep_Rooting_Disposition is particularly valuable in climates with winter rainfall recharging deep soil layers that are then progressively depleted during dry growing seasons, as deep roots access this stored water unavailable to shallow-rooted vines. However, deep rooting is only beneficial where soil conditions permit root penetration and where subsoil water is actually present, making site-soil characteristics critical determinants of deep rooting value.
Deep_Rooting_Disposition is_a Root_System_Drought_Adaptation_Disposition. Deep rooting provides drought avoidance through access to subsoil water reserves. V. berlandieri and V. rupestris parentage generally confers deeper rooting than V. riparia. Deep rooting value depends on soil conditions permitting root penetration and presence of subsoil water. Maximum rooting depth under field conditions ranges from ~1 m to >6 m depending on rootstock and soil. Deep rooting is particularly valuable in Mediterranean climates with winter-recharged deep soil water.
Tiefwurzelungs-Disposition
Unterboden-Durchwurzelung
Vertikale-Wurzelausdehnung
deep root penetration
subsoil rooting capacity
vertical root extension
Smart DR et al. (2006) Grapevine rooting patterns: a comprehensive analysis and review, American Journal of Enology and Viticulture 57:89-104 Southey JM, Archer E (1988) The effect of rootstock cultivar on grapevine root distribution and density, in The Grapevine Root and its Environment, Department of Agriculture and Water Supply Technical Communication 215, Pretoria Bauerle TL et al. (2008) Root foraging in response to heterogeneous soil moisture in two grapevines that differ in potential growth rate, New Phytologist 179:857-866
Deep Rooting Disposition
A deep shade mature leaf at node 9 on a basal lateral shoot of Vitis vinifera cv. Merlot in Pomerol, France, positioned in dense interior canopy receiving approximately 12% of full sunlight (daily PPFD averaging 185 μmol photons m⁻² s⁻¹ with rare brief peaks to 350 μmol photons m⁻² s⁻¹), measuring 18.5 cm length by 17.2 cm width representing 58% size increase compared to exterior leaves despite very thin fragile structure, lamina thickness of 185 μm with poorly developed single palisade layer of 35 μm and highly expanded spongy mesophyll of 130 μm with very large air spaces, very low chlorophyll content of 245 μg cm⁻² with high a=b ratio of 2.2 indicating maximum shade adaptation, very thin cuticle of 2.3 μm, low stomatal density of 125 stomata mm⁻² with large stomata, virtually absent trichomes, pale yellow-green coloration on both surfaces indicating chlorophyll deficiency, demonstrating maximum photosynthesis rate of only 4.8 μmol CO₂ m⁻² s⁻¹ even under experimental high light, very low light compensation point of 14 μmol photons m⁻² s⁻¹ but estimated daily carbon balance near zero or slightly negative considering 24-hour respiratory costs, showing signs of marginal function including slight chlorosis and reduced turgidity, measured at BBCH stage 79 (majority of berries touching) in early August, representing extreme shade adaptation with questionable carbon balance requiring canopy management intervention through shoot thinning or leaf removal to eliminate non-productive shaded zones.
A shade mature leaf exhibiting maximum structural and functional adaptations to severe light limitation receiving less than 15-20% of full sunlight in Vitis vinifera, characterized by extreme expression of shade syndrome including very large lamina size often 50-70% increased compared to sun leaves maximizing light-intercepting surface area despite marginal carbon economics, very thin lamina frequently 160-220 μm with single reduced palisade layer and highly expanded spongy mesophyll creating fragile structure, very low chlorophyll concentrations per unit area of 220-300 μg cm⁻² but very high per unit mass reflecting thin tissue attempting to maximize light capture with minimal structural investment, very thin cuticle often 2.0-3.0 μm providing minimal protection but reducing resource demand, low stomatal density of 110-140 stomata mm⁻² with very large individual stomata, minimal trichome presence, very pale green to yellow-green coloration reflecting chlorophyll deficiency and reduced photosynthetic capacity, high chlorophyll b proportion with a=b ratios often 2.0-2.5 indicating maximum investment in light-harvesting complexes attempting to compensate for severe light limitation, and physiological characteristics including very low light-saturated photosynthesis rates of 3-6 μmol CO₂ m⁻² s⁻¹, very low light compensation points of 10-18 μmol photons m⁻² s⁻¹ representing maximum efficiency adaptation but marginal or negative whole-leaf carbon balance when respiratory costs exceed photosynthetic gains, especially during periods of environmental stress increasing respiration or reducing already limited photosynthetic capacity.
Deep shade mature leaves represent the extreme shade end of the light adaptation spectrum, distinguished from moderate shade leaves by very pronounced structural modifications and severely compromised photosynthetic capacity approaching or below carbon balance threshold. These leaves develop in severely shaded interior canopy positions typical of dense unmanaged canopies, basal zones under heavy upper canopy coverage, or northern canopy faces (Northern Hemisphere) or southern faces (Southern Hemisphere) receiving minimal direct sunlight. Carbon economics of deep shade leaves are often marginal or negative, with photosynthetic gains barely exceeding or falling short of respiratory costs, making these leaves parasitic on the vine rather than contributors to carbon balance. Occurrence of substantial deep shade leaf populations (>20% of canopy) indicates excessive canopy density requiring management intervention, as these leaves consume resources without adequate return and create favorable environments for fungal diseases through poor air circulation and maintained humidity. Removal of deep shade leaves through strategic leaf removal or shoot thinning typically improves whole-vine carbon balance and fruit quality without yield penalty, as these leaves contribute minimally to carbohydrate production supporting fruit development.
extreme shade leaf
extremschatten Blatt
minimal light leaf
minimallicht Weinblatt
tiefschatten reifes Blatt
very low light-adapted leaf
Based on Smart (1985) "Principles of grapevine canopy microclimate manipulation with implications for yield and quality" American Journal of Enology and Viticulture 36=230-239 Poni et al. (2006) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 57=397-407 Hunter et al. (1995) "Partial defoliation of Vitis vinifera L. cv. Cabernet Sauvignon/99 Richter= Effect on root growth, canopy efficiency, grape composition, and wine quality" American Journal of Enology and Viticulture 46=306-314 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 329-352.
Deep Shade Mature Leaf
Syrah grafted on 110 Richter rootstock at 18 years in deep alluvial loam showing deep tap root extending 285 cm depth documented through excavation and augering, deep tap root diameter 11.8 cm at crown maintaining 5.2 cm diameter at 200 cm depth, deep tap root penetrating through multiple soil horizons including A horizon (0-30 cm), B horizon (30-120 cm), and deep C horizon (120-285 cm), soil moisture monitoring revealing deep tap root accessing moist soil (18-22% volumetric water content) at 240-285 cm depth during August drought when upper 180 cm soil severely dry (<8% VWC), deep tap root providing critical water supply enabling vine to maintain photosynthesis and complete fruit ripening despite prolonged surface drought, deep rooting characteristic of 110R Berlandieri parentage
Deep tap root is tap root extending to substantial depth typically exceeding 150 cm and often reaching 200-400+ cm in permeable soils, emphasizing depth penetration and access to deep soil moisture reserves, characterized by strong vertical orientation maintaining downward trajectory through multiple soil horizons, genetic determination by deep-rooting rootstock varieties particularly Vitis berlandieri-based cultivars, progressive deepening throughout vine life potentially adding 20-50 cm annually in young vigorous vine slowing to 5-15 cm in mature vine, and critical functional role in drought tolerance enabling vine to access deep permanent moisture or water table during surface soil drought providing continued water supply when shallow roots non-functional, with deep tap root development favored by deep well-drained soils, arid or Mediterranean climates selecting for deep rooting, and rootstock varieties bred specifically for drought tolerance through deep penetration capacity. Deep tap root depth achievement includes= exceeding 150 cm minimum (deep tap root by definition penetrating deeper than typical tap root depth threshold), commonly reaching 200-300 cm (deep tap roots in favorable conditions routinely extending to this depth range in mature vine), potentially exceeding 400 cm (extreme cases of deep tap root penetration documented particularly under drought stress or in very deep soils), demonstrating progressive deepening (deep tap root continuously growing deeper throughout vine life though rate declining with age), and showing environmental variation (same rootstock potentially producing deeper tap root under drought stress than with irrigation reflecting adaptive response). Deep tap root genetic determination includes= characteristic of Berlandieri types (Vitis berlandieri and hybrids showing inherent deep rooting capacity producing particularly deep tap roots), typical of vigorous rootstocks (varieties like 110 Richter, 140 Ruggeri, 1103 Paulsen producing deep tap roots as part of vigorous root system architecture), absent in shallow-rooting types (Riparia-based rootstocks showing limited tap root depth potential typically <100 cm reflecting genetic constraints), demonstrating breeding objective (deep tap root capacity specifically selected in rootstock development programs for drought tolerance), and showing hybrid variation (rootstock hybrids showing intermediate tap root depth potential reflecting genetic contributions from deep and shallow parents). Deep tap root soil requirements includes= requiring permeable substrates (deep tap root development needing deep soils without restrictive layers allowing continued penetration), favored by sandy or loamy textures (coarse-textured soils with large pores facilitating deep root penetration and providing adequate aeration at depth), inhibited by clay or compacted layers (dense soils with small pores restricting penetration limiting maximum tap root depth), prevented by hardpans or bedrock (impenetrable layers absolutely limiting depth regardless of genetic potential), and affected by water table (permanent shallow water table potentially limiting deep tap root development due to anaerobic conditions). Deep tap root drought tolerance function includes= accessing deep permanent moisture (deep tap root potentially reaching capillary fringe above water table or deep moist soil layers maintaining function during surface drought), enabling summer water supply (deep tap root contribution to vine water supply critical during peak summer when surface soil completely dry), demonstrating drought avoidance (deep tap root allowing vine to avoid rather than merely tolerate drought stress by maintaining adequate water access), showing survival advantage (vines with deep tap roots surviving drought periods that kill shallow-rooted vines), and providing sustained productivity (deep tap root enabling continued photosynthesis and growth under drought conditions improving yield and quality). Deep tap root hydraulic conductivity includes= showing large vessel diameters (deep tap root xylem containing large vessels optimizing water transport from depth), demonstrating high transport capacity (deep tap root capable of conducting 20-50+ liters daily from deep soil layers during peak transpiration), exhibiting efficient pathway (direct vertical deep tap root providing shortest distance from deep moisture to canopy compared to circuitous lateral root pathways), maintaining function year-round (deep tap root xylem remaining functional throughout growing season unlike some lateral roots subject to drying in surface soil), and contributing substantially to total vine water supply (deep tap root potentially providing 30-60% of total vine water during severe drought emphasizing critical importance). Deep tap root anchorage contribution includes= providing maximum vertical stability (deep tap root penetration creating strongest possible anchor against uprooting forces), resisting extreme wind loads (deep tap root essential for vine survival in exposed sites subject to severe winds), demonstrating pull-out resistance (deep tap root requiring very large forces to extract from soil exceeding that needed for shallow roots), showing safety factor (deep tap root providing anchorage reserve exceeding normal loads allowing vine to withstand extreme events), and enabling large vine structure (deep tap root anchorage supporting extensive canopy and heavy crop loads in high-vigor vines).
Deep_Tap_Root emphasizes substantial depth penetration exceeding 150 cm and is distinguished from Shallow_Tap_Root by focus on deep soil exploitation. Critical for understanding drought avoidance, deep moisture access, Berlandieri genetics, permanent water supply, extreme depth penetration, and the deeply penetrating vertical root enabling vine survival under severe drought through accessing deep permanent moisture.
Deeply penetrating tap root
Extended depth tap root
Profound tap root
Tiefe Pfahlwurzel
Tiefgreifende Hauptwurzel
Weitreichende Vertikalwurzel
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Smart, D. R., et al. (2006). Root system architecture in Vitis= Influences of rootstock, cultivar, and environment. American Journal of Enology and Viticulture, 57(4), 408-418.
Deep Tap Root
A deep upper lateral sinus representing the left upper lateral sinus on a mature leaf at node 7 of Vitis vinifera cv. Mourvèdre in Bandol, France, exhibiting pronounced depth of 4.5 cm measured from sinus base to the imaginary line connecting apical and upper lateral lobe edges, penetration reaching approximately 62% of the distance from blade margin toward median vein indicating very deep dissection, well-defined U-shaped contour with clearly rounded sinus base, width of 1.4 cm at opening creating depth-to-width ratio of 3.2 indicating substantial depth relative to width, sinus creating very clear prominent separation between the elongated terminal apical lobe and upper lateral lobe, sinus base positioned only 3.8 cm from petiolar junction due to the deep penetration, left margin formed by acutely tapering apical lobe edge and right margin by upper lateral lobe edge both showing smooth curvature without teeth in sinus region, deep sinus contributing to characteristic deeply dissected five-lobed Mourvèdre architecture with prominently isolated terminal lobe, bilateral near-symmetry with right upper lateral sinus showing similar deep dissection (4.3 cm), measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing maximum deep upper lateral sinus development diagnostic for deeply dissected cultivars.
An upper lateral sinus exhibiting pronounced depth creating substantial separation between apical and upper lateral lobes in Vitis vinifera, characterized by depth typically exceeding 3-4 cm measured from sinus base to lobe edge connection line, penetration often reaching 50-70% of the distance from blade margin toward median vein, well-defined U-shaped or V-shaped contour with clearly distinguished sinus base, substantial visual prominence creating obvious apical lobe isolation, occurrence primarily in deeply dissected cultivars showing pronounced overall lobe development, and critical role in creating prominent well-separated terminal apical lobes characteristic of deeply lobed cultivar morphology. Deep upper lateral sinuses represent maximum expression of upper sinus development, occurring in cultivars with strong lobe dissection including Syrah, Mourvèdre, deeply lobed Muscat types, and some Cabernet Franc clones, creating distinctive leaf architecture with prominently isolated terminal lobes, and serving as important diagnostic features indicating deeply dissected cultivar groups when combined with other morphological characteristics.
Deep upper lateral sinuses are distinguished from shallow upper lateral sinuses by depth typically >3-4 cm and penetration >50% toward median vein. The deep designation emphasizes pronounced dissection creating strong lobe isolation contrasting with weak shallow sinuses. Depth creates prominent visual separation making deeply dissected leaves immediately recognizable from considerable distance based on isolated apical lobe appearance. Penetration percentage (depth relative to distance toward median vein) provides normalized assessment accounting for overall blade size variation, with >50-60% indicating deep dissection. Shape is typically U-shaped though V-shaped deep sinuses occur in some cultivars. Occurrence is restricted primarily to deeply dissected cultivar groups, with weakly lobed varieties universally showing shallow upper lateral sinuses. Correlation with overall lobe dissection is strong, with cultivars showing deep upper lateral sinuses typically also exhibiting deep lower lateral sinuses and substantial overall lobe development. Genetic control is strong with deep dissection patterns showing high heritability and consistent expression. Environmental modification is relatively limited with deep sinus cultivars maintaining pronounced dissection across vigor conditions, though extreme low vigor may produce slightly shallower sinuses. Functional implications include maximum apical lobe isolation enabling independent movement and potentially enhanced blade flexibility. Ampelographic utility is substantial as secondary diagnostic feature, with deep upper lateral sinuses combined with other characters enabling identification of deeply dissected cultivar groups and specific varieties within those groups.
ausgeprägte obere Lateralbucht
deeply incised apical sinus
profound upper sinus
pronounced upper lateral sinus
tief eingeschnittene Apikalsinus
tiefe obere Seitenbucht
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 58-119 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 173-208.
Deep Upper Lateral Sinus
A deep winter dormant bud on Vitis vinifera cv. Cabernet Franc in mid-January in Chinon, France, showing no budbreak response when excised canes are placed in forcing conditions at 25°C with 16-hour photoperiod, having accumulated only 400 of the required 1100 chilling hours, with bud tissues exhibiting maximum concentrations of dehydrins and other cold-protective proteins.
A winter dormant bud in the most profound phase of endodormancy occurring during mid-winter (typically December-January in Northern Hemisphere), characterized by maximum chilling requirement for release, complete inability to resume growth even under optimal forcing conditions, minimal metabolic activity, and maximum cold hardiness. Deep winter dormant buds show no response to warm temperatures or long photoperiods until substantial chilling hour accumulation has occurred, representing the nadir of the annual growth-dormancy cycle where buds are most resistant to premature budbreak but also most vulnerable to catastrophic freeze damage if temperatures drop below cultivar-specific lethal thresholds.
Deep winter dormancy represents the transition point between dormancy induction (autumn) and dormancy release (late winter/early spring). Buds in deep dormancy often show maximum freezing tolerance, sometimes surviving temperatures as low as -25°C to -30°C in cold-hardy cultivars. Premature warm periods during deep dormancy generally do not trigger budbreak, protecting against false spring damage, though extended warm spells can accelerate chilling hour accumulation through negative chilling models.
deep dormancy bud
maximum endodormancy bud
mid-winter dormant bud
tiefe Winterruhe Knospe
Based on Schnabel & Wample (1987) "Dormancy and cold hardiness in Vitis vinifera L. cv. White Riesling as influenced by photoperiod and temperature" American Journal of Enology and Viticulture 38=265-272 Ferguson et al. (2011) "Dynamic changes in bud cold hardiness during dormancy and the influence of apple and pear budbreak" Journal of the American Society for Horticultural Science 136=313-323 Kovaleski & Londo (2019) "Tempo of gene regulation in wild and cultivated Vitis species shows coordination between cold hardiness and budbreak" Plant Science 287=110165.
Deep Winter Dormant Bud
Defect visibility assessment at grape sorting table identifies multiple defect categories requiring removal, including Botrytis-infected berries (gray fuzzy surface growth, immediately discarded), sunburned berries (brown necrotic patches, selectively removed), and insect-damaged clusters (visible feeding tracks and frass, entire clusters rejected), with defect visibility enabling rapid quality sorting that excludes compromised fruit from premium wine production while potentially directing affected material to lesser tiers, demonstrating defect detection as the practical quality gate protecting wine quality through visual screening.
Defect visibility quality is a surface appearance quality that represents the presence of visually detectable surface abnormalities on grape berries or in wine, characterizing visible damage, contamination, disease symptoms, or foreign matter that indicates quality compromise requiring assessment and potentially affecting processing or consumption suitability. Within the BFO framework, this quality specifically addresses the anomaly detection dimension of surface assessment, distinguishing visible defects from normal surface characteristics while recognizing that defect visibility enables quality screening before more detailed evaluation and prevents consumption of compromised product. The ontological significance lies in defect visibility serving as the primary visual quality gate in both grape reception and wine service, where visible surface defects trigger rejection, sorting, or investigation before proceeding with processing or consumption, making defect detection the first line of quality defense in practical grape and wine handling. The physical basis for defect visibility involves visual detection of surface abnormalities that contrast with expected normal appearance, including color anomalies (rot discoloration, oxidation browning), texture abnormalities (shriveling, cracking, disease lesions), foreign matter (insect damage, debris, sediment), and surface films (microbial growth, oil contamination). The defect categories in grape contexts include disease symptoms (Botrytis, powdery mildew, sour rot creating visible surface changes), physiological damage (sunburn, dehydration, cracking), pest damage (visible insect feeding, residues), and physical damage (crushing, abrasion from handling). The defect categories in wine contexts include surface contamination (floating particles, cork fragments), microbial films (aerobic spoilage creating visible surface growth), and foreign matter (any non-wine material visible at surface). The assessment protocol for defect detection involves systematic visual examination under appropriate lighting, with defect presence triggering either rejection, sorting (removing affected material), or further investigation to determine defect nature and quality implications. Understanding defect visibility quality enables quality screening at grape reception and wine service, appropriate response to detected defects, and consumer protection from visibly compromised product.
Defect_Visibility_Quality addresses the anomaly detection dimension of surface assessment, representing the primary visual quality screening function. This quality enables rapid identification of quality-compromising conditions in both grape and wine contexts, supporting rejection, sorting, or investigation decisions. The class provides the negative counterpart to positive surface qualities (bloom, gloss), with defect absence being the baseline expectation for quality product.
Defektsichtbarkeit
Fehlererscheinung
Oberflächenanomalien
anomaly visibility
quality defect appearance
sichtbare Mängel
surface abnormalities
visible defects
Ough CS, Amerine MA (1988) Methods for Analysis of Musts and Wines, 2nd Ed. John Wiley & Sons, New York, pp. 1-34 Steel CC et al. (2013) Grapevine bunch rots: impacts on wine composition, quality, and potential procedures for the removal of wine faults. Journal of Agricultural and Food Chemistry 61(22): 5189-5206
Defect Visibility Quality
Comprehensive characterization of Defense_Mechanism_Disposition at the University of Udine assessed multiple constitutive and induced defense parameters across 20 cultivars to evaluate the breadth and coordination of defense capacity. Constitutive measures included cuticle thickness, trichome density, constitutive phenolic content, and preformed PR protein levels. Induced responses were assessed following Plasmopara viticola inoculation, measuring stilbene accumulation, PR gene expression, and callose deposition at 24, 48, and 72 hours post-inoculation. Principal component analysis revealed that Defense_Mechanism_Disposition varied substantially among cultivars and comprised largely independent components. 'Cabernet Sauvignon' exhibited strong constitutive physical defenses (thick cuticle 4.2 μm, high trichome density 28/mm²) but moderate chemical and induced responses. Conversely, 'Regent' (PIWI cultivar) showed modest constitutive barriers but exceptionally strong induced stilbene production (45 μg/g fresh weight at 48h versus 8 μg/g in 'Cabernet Sauvignon'). 'Muscadinia rotundifolia' demonstrated uniformly high Defense_Mechanism_Disposition across all measured parameters, consistent with its broad-spectrum disease resistance. These findings demonstrate that overall resistance phenotypes emerge from combinations of defense mechanisms varying independently among genotypes, with implications for breeding strategies targeting multiple defense components.
A Defense_Mechanism_Disposition is a Biotic_Stress_Resistance_Disposition that inheres in a Vitis vinifera plant and represents the underlying capacity to deploy specific defensive structures, compounds, or processes that protect against herbivores and pathogens, abstracted from the particular pest or pathogen being resisted. This disposition captures the mechanistic basis of biotic stress resistance, complementing the pathogen-specific and pest-specific resistance classes that describe resistance outcomes against particular organisms. Defense mechanisms in plants are traditionally categorized as constitutive (preformed, always present) versus induced (activated upon attack), and as direct (affecting the attacker directly) versus indirect (recruiting natural enemies or signaling to neighboring plants). For grapevine, constitutive defenses include physical barriers such as cuticle, trichomes, and tough tissues, as well as preformed chemical defenses including phenolic compounds, antimicrobial proteins, and structural polymers. Induced defenses encompass the hypersensitive response, systemic acquired resistance, production of phytoalexins (particularly stilbenes in Vitis), pathogenesis-related proteins, and volatile emissions. Understanding defense mechanism dispositions enables prediction of resistance breadth across multiple attackers sharing susceptibility to particular mechanisms, identification of defense trade-offs and synergies, and targeted breeding for enhanced defense capacity rather than resistance to individual threats. The Defense_Mechanism_Disposition hierarchy provides a mechanistic framework complementing the taxonomically organized Pathogen_Resistance_Disposition and Pest_Resistance_Disposition hierarchies.
Defense_Mechanism_Disposition is_a Biotic_Stress_Resistance_Disposition. This class provides a mechanistic perspective on resistance, complementing the pathogen/pest-specific resistance classes elsewhere in VVO. Child classes include Constitutive_Defense_Disposition (preformed defenses) and Induced_Defense_Disposition (activated upon attack). Defense mechanisms may contribute to resistance against multiple pathogens and pests sharing susceptibility to particular barriers or compounds, enabling prediction of resistance breadth from mechanistic knowledge.
Abwehrmechanismus-Kapazität
Pflanzenabwehr-Disposition
Verteidigungsmechanismus-Disposition
defense trait disposition
plant defense capacity
resistance mechanism
Pieterse CMJ et al. (2012) Hormonal modulation of plant immunity, Annual Review of Cell and Developmental Biology 28:489-521 Dicke M, Baldwin IT (2010) The evolutionary context for herbivore-induced plant volatiles: beyond the "cry for help", Trends in Plant Science 15:167-175 Hammerschmidt R (1999) Phytoalexins: what have we learned after 60 years?, Annual Review of Phytopathology 37:285-306
Defense Mechanism Disposition
Defense response quality comparison between the downy mildew-resistant Vitis rupestris and susceptible V. vinifera Chardonnay characterized the differential immune activation underlying contrasting disease outcomes. Inoculation with Plasmopara viticola sporangia triggered rapid defense response quality in V. rupestris with reactive oxygen species burst detectable within 2 hours at infection sites, callose deposition visible by 6 hours forming papillae beneath attempted penetration sites, and hypersensitive cell death limiting pathogen spread by 24 hours. Transcriptomic analysis revealed defense response quality activation with greater than 500 defense-related genes upregulated greater than 4-fold within 12 hours in V. rupestris, including pathogenesis-related proteins, stilbene synthases, and signaling components. In contrast, V. vinifera Chardonnay showed delayed and attenuated defense response quality with ROS burst 60% lower amplitude and 4 hours delayed, callose deposition incomplete and occurring only after hyphal penetration, and no effective hypersensitive response allowing pathogen colonization to proceed. The defense response quality differential translated to disease outcomes, with V. rupestris showing less than 5% of inoculation sites developing sporulating lesions while Chardonnay showed greater than 80% successful infection and abundant sporulation. The defense response quality characterization identified the speed and amplitude of early defense activation as critical determinants of resistance, informing breeding strategies to introgress rapid-response capacity from resistant species into V. vinifera backgrounds.
Defense response quality is a pathogen infection indicator quality that inheres in a grapevine organism and represents the capacity to activate and deploy innate immune mechanisms in response to pathogen recognition, encompassing the localized rapid responses that limit pathogen spread at infection sites, the production of antimicrobial compounds that directly inhibit pathogen growth, the synthesis of pathogenesis-related proteins that contribute to defense, and the systemic signaling that primes distant tissues for enhanced resistance, with the effectiveness of defense responses determining whether attempted infections are successfully repelled, contained with minimal damage, or progress to cause significant disease. This quality captures the active component of disease resistance that complements constitutive barriers, recognizing that plants possess sophisticated immune systems capable of recognizing pathogen-associated molecular patterns and effector molecules, triggering defense cascades that can effectively limit pathogen colonization when appropriately activated. The ontological significance of defense response quality lies in its role as a primary determinant of disease resistance that varies among cultivars, can be induced by elicitor treatments, and provides targets for breeding programs seeking to enhance disease resistance through improved defense activation capacity. The biological basis of defense responses encompasses the recognition of pathogen molecules by plant receptor proteins triggering signal transduction cascades, the transcriptional reprogramming activating defense gene expression, the metabolic redirection producing antimicrobial compounds and structural reinforcements, and the systemic signaling through salicylic acid, jasmonic acid, and other hormones that coordinates whole-plant defense. The temporal dynamics of defense responses critically affect their effectiveness, with rapid responses more likely to limit pathogen establishment while delayed responses may occur after substantial colonization has already occurred, creating the kinetic competition between pathogen growth and defense activation that determines infection outcomes. The variation in defense response quality among grapevine cultivars reflects genetic differences in receptor repertoires, signaling pathway components, and metabolic capacity for defense compound production, providing the mechanistic basis for the differential disease susceptibility observed among cultivars and the targets for marker-assisted selection in resistance breeding.
Defense_Response_Quality is_a Pathogen_Infection_Indicator_Quality, serving as parent class for Hypersensitive_Response_Quality, Phytoalexin_Production_Quality, Pathogenesis_Related_Protein_Induction_Quality, and Systemic_Acquired_Resistance_Quality which address specific defense mechanisms. This class addresses the overall capacity for active defense while child classes capture particular response types. The kinetic competition between pathogen growth and defense activation determines infection outcomes. Cultivar variation in defense capacity provides breeding targets for resistance improvement. Distinguished from Disease_Tolerance_Quality which addresses performance maintenance rather than pathogen limitation.
Abwehrreaktion
Immunantwort
Pathogenabwehr
defense activation
immune response
induced resistance
induzierte Resistenz
pathogen defense
Jones JDG, Dangl JL (2006) The plant immune system. Nature 444:323-329 Armijo G et al. (2016) Grapevine pathogenic microorganisms: understanding infection strategies and host response scenarios. Frontiers in Plant Science 7:382 Dry IB et al. (2010) Molecular strategies to enhance the genetic resistance of grapevines to powdery mildew. Australian Journal of Grape and Wine Research 16:94-105
Defense Response Quality
Deficiency severity quality classification for nitrogen status in Oregon Pinot Noir provided a framework for calibrating vineyard nitrogen management to the range of conditions encountered across variable soil types and vine ages. The deficiency severity quality system established four categories based on integrated assessment of tissue nitrogen concentration, visual symptom expression, and functional indicators. Latent deficiency severity quality was diagnosed where petiole N fell in the 0.55-0.65% range with no visible chlorosis but reduced SPAD readings (32-35 versus normal greater than 38), indicating subclinical limitation warranting monitoring and potential preventive nitrogen application. Mild deficiency severity quality corresponded to petiole N of 0.45-0.55% with subtle generalized pallor visible primarily when comparing to N-sufficient reference vines, shoot length 10-20% below site norm, and SPAD readings of 28-32, indicating moderate limitation warranting corrective fertilization. Moderate deficiency severity quality corresponded to petiole N of 0.35-0.45% with obvious chlorosis visible without comparison (score 2-3 on 0-5 scale), shoot length 20-35% reduced, early senescence of basal leaves, and SPAD readings of 22-28, indicating significant limitation requiring prompt corrective action. Severe deficiency severity quality corresponded to petiole N below 0.35% with intense chlorosis (score 4-5), dramatic growth reduction (greater than 35%), extensive premature senescence, and SPAD readings below 22, indicating critical limitation requiring immediate intensive intervention. The deficiency severity quality framework enabled consistent communication among vineyard managers and consultants, with management recommendations calibrated to severity():) monitoring for latent, standard fertilization for mild, increased rates and foliar supplementation for moderate, and emergency intervention with split applications for severe.
Deficiency severity quality is a nutrient deficiency indicator quality that inheres in a grapevine organism and represents the degree or intensity of nutritional limitation as reflected in symptom expression, physiological impairment, and production impact, ranging from latent deficiency where tissue concentrations are suboptimal but visible symptoms have not yet developed, through mild, moderate, and severe categories that describe progressively greater symptom expression and functional impact, providing a framework for communicating deficiency status and calibrating management response intensity to deficiency magnitude. This quality captures the quantitative dimension of nutritional stress, recognizing that nutrient deficiency exists on a continuum from adequacy through subclinical limitation to overt deficiency, and that the severity classification provides critical information for management decision-making regarding the urgency and intensity of corrective action required. The ontological significance of deficiency severity quality lies in its role enabling calibrated management responses where latent deficiency might warrant monitoring and preventive adjustment while severe deficiency demands immediate intensive intervention, and in its utility for communicating nutritional status among researchers, advisors, and practitioners using standardized terminology. The biological basis of deficiency severity gradations encompasses the progressive decline in tissue nutrient concentration from luxury levels through adequate to deficient ranges, the threshold effects where visible symptoms appear only after tissue concentrations fall below critical levels, and the increasing functional impairment as deficiency intensifies affecting progressively more metabolic processes and larger proportions of the plant. The relationship between tissue nutrient concentration and severity classification shows characteristic ranges for each category, though precise thresholds vary by nutrient, tissue type, cultivar, and phenological stage, requiring calibration to local conditions and careful interpretation of tissue analysis results in the context of visual observations.
Deficiency_Severity_Quality is_a Nutrient_Deficiency_Indicator_Quality, serving as parent class for Latent_Deficiency_Quality, Mild_Deficiency_Quality, Moderate_Deficiency_Quality, and Severe_Deficiency_Quality which describe the severity continuum. This class addresses the overall concept of deficiency grading while child classes capture specific severity levels. Integrates tissue concentration, visual symptoms, and functional indicators for classification. Enables calibrated management response to deficiency magnitude. Thresholds vary by nutrient, tissue, cultivar, and phenology requiring local calibration. Communication tool for standardized terminology among practitioners.
Mangelgrad
Mangelschweregrad
Mangelstufe
Nährstoffstress-Intensität
deficiency grade
deficiency intensity
deficiency level
nutritional stress severity
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Robinson JB (1992) Grapevine nutrition. In: Coombe BG, Dry PR (eds) Viticulture, Volume 2. Winetitles, Adelaide, pp. 178-208 Bergmann W (1992) Nutritional Disorders of Plants. Gustav Fischer Verlag, Stuttgart
Deficiency Severity Quality
Cabernet Sauvignon vines in the Columbia Valley of Washington State bearing a deficit-irrigated role receive 70-80% of full ETc replacement pre-véraison and 30-50% post-véraison through regulated deficit irrigation (RDI), targeting midday stem water potentials of -1.0 to -1.2 MPa during fruit set and -1.2 to -1.6 MPa during ripening, resulting in 20-30% reductions in berry size, 15-25% increases in skin-to-pulp ratio, and measurably higher anthocyanin concentrations of 1.5-2.5 mg/g berry weight compared to fully irrigated controls.
An irrigated role that inheres in a grapevine managed under a deliberate deficit irrigation strategy that supplies less water than the vine's full evapotranspiration demand during defined phenological periods, inducing controlled water stress to regulate vegetative growth, enhance fruit composition, and improve water use efficiency.
Deficit_Irrigated_Role encompasses both regulated deficit irrigation (RDI) applying uniform deficit across the root zone and partial root zone drying (PRD) alternating irrigation between root sides to exploit ABA-mediated stomatal regulation. Both strategies exploit the differential sensitivity of vegetative and reproductive growth to water deficit: shoot elongation ceases at milder stress levels than berry ripening, enabling deficit irrigation to restrict canopy vigor while maintaining or enhancing fruit quality parameters. The optimal deficit level is cultivar-specific, site-specific, and phenological-stage-dependent, requiring ongoing monitoring of vine water status indicators.
Defizitbewässerungs-Rolle
controlled water stress role
deficit irrigation role
kontrollierte Trockenstress-Rolle
regulated deficit irrigation role
regulierte Defizitbewässerungs-Rolle
Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data, Annals of Botany 105: 661-676 Dry PR et al. (2001) Strategic irrigation management in Australian vineyards, Journal International des Sciences de la Vigne et du Vin 35: 129-139 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 510-520
Deficit Irrigated Role
The deficit irrigation strategy for Cabernet Sauvignon in Paso Robles specifies three phenological phases — Phase 1 (budburst to fruit set) at 80% ETc replacement to support canopy establishment without excessive vegetative vigor, Phase 2 (fruit set to véraison) at 40-50% ETc to impose moderate water deficit that reduces berry size by 15-25% (from 1.4 g to 1.0-1.2 g average berry weight) and increases skin-to-pulp ratio, enhancing anthocyanin and tannin concentration per unit volume of must, and Phase 3 (véraison to harvest) at 25-35% ETc to maintain controlled stress that promotes flavor concentration and tannin polymerization while avoiding severe stress that would arrest sugar accumulation or trigger premature leaf senescence.
An irrigation protocol description that documents the intentional application of water at rates below the vine's full evapotranspiratory demand during specific phenological periods, specifying the timing, severity, and duration of controlled water deficit designed to manipulate berry size, skin-to-pulp ratio, phenolic concentration, and flavor development in pursuit of defined wine quality objectives.
Deficit_Irrigation_Strategy represents the most important conceptual advance in irrigation management for premium wine production over the past three decades. The foundational insight — that moderate vine water stress during specific developmental windows improves wine quality parameters while excessive water supply produces dilute, vegetatively dominated wines — was formalized by McCarthy (1997) as Regulated Deficit Irrigation (RDI) and by Dry and Loveys (1998) as Partial Root-zone Drying (PRD): both approaches deliberately supply less water than the vine could use, but differ in implementation — RDI reduces total water volume uniformly across the root zone, while PRD alternates irrigation between root zone halves to exploit the ABA-mediated stomatal regulation triggered by drying roots while maintaining overall vine hydration. The quality benefits of deficit irrigation include reduced berry size (increasing skin-to-pulp ratio by 10-30%), enhanced phenolic concentration (anthocyanins, tannins), modified flavor profiles (reduced methoxypyrazine in Cabernet Sauvignon, enhanced rotundone in Shiraz), and improved wine structure and aging potential.
Defizitbewässerungs-Strategie
RDI protocol
controlled water stress strategy
kontrollierte Trockenstress-Strategie
regulated deficit irrigation strategy
regulierte Defizitbewässerung
McCarthy MG (1997) The effect of transient water deficit on berry development of cv. Shiraz, Australian Journal of Grape and Wine Research 3: 2-8 Dry PR and Loveys BR (1998) Factors influencing grapevine vigour and the potential for control with partial rootzone drying, Australian Journal of Grape and Wine Research 4: 140-148 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data, Annals of Botany 105: 661-676 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 400-420
Deficit Irrigation Strategy
Defoliation tolerance quality comparison across cultivars in Rheinhessen evaluated the threshold of leaf removal causing measurable fruit quality reduction using progressive basal defoliation at véraison. Riesling demonstrated high defoliation tolerance quality, sustaining removal of up to 40% of total leaf area without significant impact on fruit Brix, titratable acidity, or aromatic compound concentration, attributed to strong photosynthetic compensation with remaining leaves increasing assimilation by 35%. Müller-Thurgau showed moderate defoliation tolerance quality, with quality impacts becoming apparent at 30% leaf removal manifesting as 0.6°Brix reduction and altered acid balance. Ortega exhibited limited defoliation tolerance quality, with fruit quality compromise evident at only 20% leaf removal including 1.2°Brix reduction and premature leaf senescence indicating source limitation, informing cultivar-specific recommendations for conservative leaf removal intensity in this variety to preserve defoliation tolerance quality within sustainable limits.
Defoliation tolerance quality is a leaf removal response quality that inheres in a grapevine organism and represents the capacity to maintain essential physiological functions including fruit ripening, carbohydrate reserve accumulation, and bud development for subsequent season productivity despite reduction in total functional leaf area below the optimal level for unrestricted growth, through compensatory adjustments in remaining leaf photosynthetic efficiency, resource allocation priorities, and reserve mobilization from perennial storage tissues. This quality determines the threshold of leaf area loss that can be sustained without significant compromise to current season fruit quality or future vine productivity, enabling practical application of defoliation practices within safe limits while avoiding damage from excessive leaf removal. The ontological basis of defoliation tolerance quality rests on the principle that grapevines possess inherent redundancy and plasticity in their photosynthetic system that permits adjustment to partial capacity loss, with tolerance limits determined by the balance between reduced carbon supply from diminished leaf area and compensatory mechanisms that partially restore assimilate availability. The physiological mechanisms underlying defoliation tolerance include upregulation of photosynthetic rate in remaining leaves through increased stomatal aperture, enhanced mesophyll conductance, and elevated Rubisco activity that can increase per-leaf carbon gain by 20-40% following moderate defoliation, though this compensation rarely achieves full replacement of lost photosynthetic capacity. Additional tolerance mechanisms involve prioritization of reproductive over vegetative sinks that directs limited assimilates preferentially toward fruit ripening at the expense of continued shoot growth and reserve storage, and mobilization of stored carbohydrates from trunk, cordon, and root tissues that can buffer short-term source-sink imbalances though at cost to future vigor and productivity. Cultivar differences in defoliation tolerance quality reflect variation in photosynthetic compensation capacity, reserve accumulation patterns, and the inherent source-sink balance at typical crop loads, with high-vigor cultivars generally exhibiting greater tolerance than low-vigor types due to higher baseline leaf area relative to fruit load. The relationship between defoliation tolerance quality and fruit quality outcomes depends critically on timing, with pre-véraison defoliation affecting berry development during cell division and expansion phases while post-véraison defoliation impacts sugar accumulation, acid metabolism, and phenolic synthesis during the ripening phase.
Defoliation_Tolerance_Quality is_a Leaf_Removal_Response_Quality, representing the capacity to maintain function despite reduced leaf area. This class addresses tolerance mechanisms including photosynthetic compensation and reserve mobilization, distinct from Refoliation_Capacity_Quality which addresses replacement of lost leaf area through new growth. The threshold of ~30-40% leaf removal as general tolerance limit provides practical guidance, though cultivar-specific differences require consideration. Critical for determining safe limits of basal leaf removal practices.
Blattverlustoleranz
Defoliationsresistenz
Entblätterungstoleranz
Entlaubungsresilienz
canopy loss resilience
defoliation resistance
foliage reduction tolerance
leaf loss tolerance
Candolfi-Vasconcelos MC, Koblet W (1991) Influence of partial defoliation on gas exchange parameters and chlorophyll content of field-grown grapevines: mechanisms and limitations of the compensation capacity. Vitis 30:129-141 Petrie PR et al. (2000) Leaf removal effects on yield and fruit composition of Vitis vinifera L. cv. Pinot Noir. Annals of Botany 85:871-879 Poni S et al. (2008) Effects of pre-bloom leaf removal on growth of berry tissues and must composition in two red Vitis vinifera L. cultivars. Australian Journal of Grape and Wine Research 14:185-195
Defoliation Tolerance Quality
Deformation quality assessment in mechanical harvesting optimization balances the deformation tolerance of different cultivars, with Thompson Seedless tolerating 15-20% compression deformation with 80-90% recovery (primarily elastic), enabling relatively aggressive mechanical harvest, while Flame Seedless shows permanent damage (plastic deformation) above 12% compression, requiring gentler harvest parameters with lower shaking intensity and shorter duration to preserve marketable quality this deformation-guided mechanical harvest calibration reduces postharvest losses from 8-12% to 3-5% by matching equipment operation to variety-specific mechanical properties.
Deformation_Quality encompasses the physical qualities characterizing the nature and extent of shape change that grape tissue undergoes when subjected to mechanical stress, capturing the fundamental mechanical behavior that underlies both firmness response and damage susceptibility. Deformation describes how tissue geometry changes under load, providing information complementary to force measurement about the mechanical properties determining quality. This category subdivides into Elastic_Deformation_Quality addressing recoverable shape change where tissue returns to original dimensions upon unloading, and Plastic_Deformation_Quality capturing permanent shape change persisting after load removal. The proportion of elastic versus plastic deformation characterizes tissue mechanical behavior ranging from elastic (spring-like, fully recoverable) through viscoelastic (time-dependent recovery) to plastic (permanent deformation). Fresh, turgid grape tissue exhibits viscoelastic behavior with substantial but not complete recovery from moderate deformation, while damaged or overripe tissue shows predominantly plastic deformation reflecting structural failure. The deformation response at low stress levels indicates tissue resilience and ability to withstand transient handling forces without permanent damage, while deformation behavior at high stress reveals failure modes and damage thresholds. Deformation parameters extracted from texture testing include deformation at maximum force, deformation at failure (break strain), and recovery percentage after defined deformation. The rate of deformation affects measured properties due to tissue viscoelasticity, with fast deformation producing stiffer response than slow deformation. Understanding deformation quality enables prediction of handling damage patterns and optimization of mechanical operations (harvest, transport, packing) to minimize permanent tissue damage.
Deformation_Quality addresses the strain dimension of mechanical behavior complementing the force (stress) dimension captured by firmness. The elastic-plastic distinction captures fundamental differences in mechanical response determining whether tissue can recover from imposed stress or suffers permanent damage. The class provides the framework for understanding not just how much force tissue can resist but how it behaves under that force and what happens when the force is removed. Deformation parameters require complete force-deformation curve analysis rather than simple maximum force determination.
Deformierbarkeit
Formänderungsqualität
Verformungsqualität
deformability
deformation behavior
mechanical strain quality
mechanische Dehnungsqualität
shape change quality
Bourne MC (2002) Food Texture and Viscosity: Concept and Measurement, 2nd Ed, Academic Press pp.: 107-144 Rolle L et al. (2012) Instrumental Texture Analysis Parameters as Markers of Table-Grape and Winegrape Quality: A Review. American Journal of Enology and Viticulture 63(1): 11-28
Deformation Quality
Pinot Noir hermaphrodite flowers at BBCH 63-65 (30-50% of flowers in full bloom) exhibit dehiscing anthers with longitudinal slits fully opened along both lobes, bright yellow pollen actively releasing and coating anther surfaces, visible as yellow powder on stamens and pistils, creating pollen clouds when inflorescences are gently shaken, with individual flowers showing active dehiscence for 6-18 hours under favorable conditions (morning temperatures 20-26°C, humidity 65-75% declining to 45-55% by afternoon, light breezes 3-8 km/h) before transitioning to post-dehiscence state. Syrah at BBCH 61-63 shows dehiscing anthers with characteristic longitudinal slit openings, copious yellow pollen release creating visible yellow coloration on surrounding flower structures and upper leaf surfaces below inflorescences, with efficient pollen dispersal achieving 54-62% fruit set through combination of self-pollination and local cross-pollination among flowers. Chardonnay experiencing rain during bloom (BBCH 61-65) shows disrupted anther dehiscence with some anthers achieving partial slit opening but pollen release impaired by moisture, clumped wet pollen visible at dehiscence openings rather than dry dispersible powder, with reduced pollination efficiency contributing to decreased fruit set (38-46% versus normal 48-56% for this cultivar), demonstrating critical dependence of dehiscing anther function on favorable weather conditions.
A dehiscing anther is an anther actively releasing pollen through opened longitudinal slits (stomia) along the anther lobes, representing the functional anthesis state (BBCH 61-65) when pollen dispersal is occurring and when flowers achieve their primary male reproductive function of contributing pollen to the pollination process. Dehiscing anthers are characterized by= (1) visible longitudinal slits opened along one or both anther lobes, typically 0.4-0.9 mm in length running the length of each lobe, created by stomium cell breakdown and endothecium-driven tension (2) active or recent pollen release with yellow pollen grains visible at the dehiscence slit openings, coating anther surfaces, or forming visible clouds when flowers are disturbed by wind or mechanical movement (3) anther color remaining bright yellow to golden-yellow due to abundant pollen still contained in pollen sacs or coating external surfaces, though beginning to fade slightly compared to mature pre-dehiscence anthers as pollen depletes (4) anther lobes showing characteristic opening configuration with slit edges curled or spread apart creating gaps through which pollen escapes (5) pollen release dynamics varying from initial rapid discharge immediately upon dehiscence through gradual sustained release over subsequent hours as anther desiccation and mechanical movements continue to expel remaining pollen. The dehiscing anther state typically persists for 4-24 hours after dehiscence initiation, depending on environmental conditions (temperature, humidity, air movement) and pollen quantity, with progression to post-dehiscence state occurring as pollen reserves deplete and anther tissues undergo senescence. Dehiscence initiation occurs 1-3 hours after calyptra shedding in most flowers under favorable conditions, triggered by anther exposure to air causing moisture loss that activates the hygroscopic endothecium mechanism= differential cell wall thickening in endothecium cells creates asymmetric tensions as desiccation proceeds, generating mechanical forces that pull anther lobe walls apart along the pre-formed stomium weak zones, creating the characteristic longitudinal slits. Pollen release from dehiscing anthers follows characteristic kinetics= initial rapid release in the first 1-3 hours as dehiscence slits open and anther mechanical movements shake loose readily dispersible pollen sustained gradual release over subsequent 6-18 hours as anther desiccation continues and as air movement and anther positional changes dislodge additional pollen final slow release or retention of residual pollen that remains adhered to anther internal surfaces. The total pollen released from dehiscing hermaphrodite anthers typically represents 70-90% of total pollen content (10,500-27,000 grains per five-anther flower), with 10-30% remaining as residual pollen in post-dehiscence anthers. Environmental conditions strongly influence dehiscence dynamics and pollen release efficiency= optimal conditions (warm temperatures 22-28°C, moderate declining humidity 60-80% dropping to 40-50%, light air movement 2-6 km/h) promote complete dehiscence with efficient pollen release and dispersal suboptimal humidity (>85%) impairs dehiscence and causes pollen clumping reducing dispersal efficiency excessive wind (>20 km/h) may accelerate pollen dispersal but reduce self-pollination efficiency by removing pollen before local deposition rain disrupts dehiscence and washes released pollen away, severely impairing pollination. Dehiscing anthers represent the critical functional state for pollination, with the timing, duration, and efficiency of the dehiscing period directly determining pollen availability for both self-pollination (in hermaphrodite flowers) and cross-pollination (from staminate or hermaphrodite sources to functionally female or pistillate flowers), making dehiscing anther characteristics and environmental conditions during this phase critical determinants of fruit set success.
Dehiscing_Anther represents the active pollen release state and is distinguished from Mature_Anther (pre-dehiscence) by opened longitudinal slits and active pollen dispersal, and from Post_Dehiscence_Anther (pollen depleted) by retention of substantial pollen reserves and bright yellow color. The dehiscing state is the critical functional phase for male reproductive success and represents the peak period when environmental conditions most directly affect pollination success through effects on pollen release efficiency, pollen viability maintenance, and pollen dispersal patterns. Monitoring dehiscing anther prevalence and characteristics provides real-time assessment of bloom progression and pollination conditions, informing management decisions and enabling diagnosis of bloom-related fruit set problems.
Pollen freigebender Staubbeutel
opening anther
pollen-releasing anther
pollenstoßende Anthere
sich öffnender Staubbeutel
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 102-106 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 95-98 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109.
Dehiscing Anther
Delayed incompatibility in grapevine typically manifests 5-15 years after grafting, with vines showing initial normal growth followed by progressive vigor decline, premature autumn leaf coloration, reduced yield, and eventually vine death. Anatomical examination of delayed-incompatible unions reveals progressive necrotic zone expansion, discontinuous cambial bands, excessive starch accumulation in scion bark, and constricted vascular elements at the interface. The combination of Syrah on certain 110 Richter clones has shown delayed incompatibility in some California vineyards, with 15-30% of vines declining between years 8-15 despite initial take rates above 90%. Hydraulic conductance across delayed-incompatible unions declines progressively from 70-80% of normal at year 3 to 30-50% by year 10.
A graft incompatibility disposition that is the inherited capacity of certain Vitis vinifera scion-rootstock combinations to exhibit incompatibility symptoms that emerge years to decades after initial successful graft establishment, resulting from progressive vascular dysfunction, cumulative metabolic imbalances, or gradual failure of the graft union under production stress.
Delayed incompatibility represents a particularly challenging phenomenon because affected vines may perform normally for years before decline begins, complicating diagnosis and economic planning. The progressive nature suggests cumulative damage from subclinical dysfunction rather than acute rejection. Environmental stresses (drought, crop load, disease) may accelerate symptom expression in marginally compatible combinations. Distinguishing delayed incompatibility from other decline syndromes (trunk diseases, viral infections) requires careful union examination and exclusion of alternative causes.
Spätinkompatibilität
chronic incompatibility trait
late-onset incompatibility
progressive Veredlungsunverträglichkeit
progressive union failure
verzögerte Inkompatibilität
Goldschmidt EE (2014) Plant grafting: new mechanisms, evolutionary implications. Frontiers in Plant Science 5:727 Stamp JA (2001) The contribution of imperfections in nursery stock to the decline of young vines in California. Phytopathologia Mediterranea 40:S369-S375
Delayed Incompatibility Disposition
AXR#1 (Aramon × V. rupestris Ganzin No. 1) exhibited delayed incompatibility with some cultivars, showing union deterioration and vine decline after 15-25 years in addition to the phylloxera susceptibility that ultimately led to its abandonment in California. Certain Rupestris du Lot (St. George) combinations showed delayed incompatibility symptoms in old California vineyards, with graft union constriction and bark necrosis becoming evident in vines 20-30 years of age. Some V. champinii × V. riparia experimental hybrids developed at UC Davis demonstrated delayed incompatibility in long-term trials, with initially vigorous vines declining after 8-12 years, necessitating elimination from breeding programs despite promising early pest resistance data.
A graft affinity rootstock role that inheres in a grafted rootstock by virtue of its tendency to exhibit graft incompatibility symptoms that manifest years or decades after initially successful union formation, resulting in progressive vine decline, reduced productivity, and eventual graft failure. Delayed incompatibility (also termed slow incompatibility or translocated incompatibility) is characterized by an initial period of apparently normal graft function followed by gradual deterioration of the graft union and vine performance, typically becoming evident 5-20 years post-planting. Symptoms include progressive constriction or overgrowth at the graft junction, phloem dysfunction reducing assimilate transport (causing scion starch accumulation above and rootstock carbohydrate depletion below the union), xylem discontinuity restricting water flow (causing drought-like symptoms despite adequate soil moisture), bark necrosis at the union interface, reduced scion vigor and yield, premature autumn coloration, and increased susceptibility to trunk diseases. Delayed incompatibility is difficult to predict from early graft union appearance, often requiring decade-long field trials to identify. The condition may reflect progressive accumulation of incompatibility compounds, gradual vascular occlusion, or developmental divergence between grafting partners over time.
Delayed_Incompatibility_Prone_Rootstock_Role captures a particularly insidious form of graft incompatibility that cannot be detected through standard nursery evaluation, as affected combinations may perform normally for years before decline commences. This phenomenon necessitates long-term field testing (minimum 10-15 years) before declaring new rootstocks as universally compatible. The mechanisms underlying delayed incompatibility remain incompletely understood but may involve progressive phloem discontinuity, accumulation of phenolic oxidation products at the union, or immunological-like rejection responses that develop slowly over time. Historical instances of delayed incompatibility (AXR#1, some early French hybrids) have caused significant economic losses, motivating the conservative, multi-decade evaluation protocols applied to new rootstock candidates.
Spätunverträglichkeit-anfällige Unterlage
delayed incompatibility prone rootstock
late-onset incompatibility rootstock
slow incompatibility rootstock
translocated incompatibility rootstock
translokatische Inkompatibilität-Unterlage
verzögert inkompatible Unterlage
Hartmann HT et al. (2011) Hartmann and Kester's Plant Propagation 8th Ed, Pearson pp.: 415-460 Pina A, Errea P (2005) A review of new advances in mechanism of graft compatibility-incompatibility, Scientia Horticulturae 106: 1-11 Mosse B (1962) Graft incompatibility in fruit trees, Commonwealth Bureau of Horticulture Technical Communication No. 28 Cookson SJ, Ollat N (2013) Grafting with rootstocks induces extensive transcriptional re-programming in the shoot apical meristem of grapevine, BMC Plant Biology 13: 147
Delayed Incompatibility Prone Rootstock Role
Merlot on 140Ru rootstock delayed incompatibility zone at 11 years post-grafting shows union that healed apparently successfully years 1-5 (adequate callus, vascular differentiation, normal function) but progressively deteriorating years 6-11, current state showing 25-40 mm zone with developing necrotic regions (brown tissue 2-5 mm depth recently appeared), vascular connections showing 30-45% decline in conductance over past 3 years (water transport reduced from 85% to 55% of non-grafted capacity), progressive union swelling from 115% year 5 to 155% year 11, vine showing declining vigor (shoot growth 40-60% reduced), small chlorotic leaves, poor fruit development, symptoms worsening annually, representing delayed incompatibility expression after initially successful union
Delayed incompatibility zone is incompatibility zone that develops years after initially successful graft union healing, characterized by progressive deterioration of union quality including vascular dysfunction, tissue necrosis, and mechanical weakness appearing 3-15+ years post-grafting, representing late-expressing genetic incompatibility or cumulative stress-induced union failure. Delayed incompatibility development timeline includes= initial successful establishment (years 1-3, graft showing apparently normal healing, callus formation, vascular differentiation, adequate function), asymptomatic period (years 3-8, union appearing healthy, vine growing and producing normally without obvious problems), symptom onset (years 5-15+, vine beginning to show declining vigor, reduced growth, or other incompatibility indicators), progressive deterioration (symptoms worsening over subsequent years, union quality declining progressively), and eventual severe dysfunction or failure (union becoming severely compromised causing vine decline or death within 10-20 years post-grafting depending on severity and environmental stress). Delayed incompatibility manifestations includes= progressive necrosis development (dead tissue zones appearing at union interface years after grafting, expanding over time), vascular deterioration (initially functional xylem and phloem connections progressively failing or becoming occluded), cambial dysfunction (initially continuous cambium showing progressive gaps or reduced activity in zones), increasing union swelling (circumferential enlargement progressively increasing beyond 125-150% suggesting compensatory overgrowth), mechanical weakening (union breaking strength declining over years increasing fracture risk), and symptom intensification (decline symptoms appearing and progressively worsening rather than stable mild symptoms). Delayed incompatibility cellular and molecular processes includes= cumulative phenolic accumulation (phenolic compounds progressively depositing at interface creating barriers to transport and causing tissue death), progressive cell death (programmed cell death pathways activated years post-grafting causing necrosis at union), delayed phytotoxin effects (toxic compounds produced by one partner causing progressive damage to other over years), vascular occlusion (vessels and sieve tubes becoming plugged by tyloses, gums, or other substances reducing conductance), and cambial senescence (cambium at union losing division capacity or organization over time preventing maintenance and repair). Delayed incompatibility genetic and physiological basis includes= genetic incompatibility with delayed expression (molecular recognition problems not immediately causing failure but progressively disrupting function over years), stress-triggered incompatibility (environmental stress unmasking latent incompatibility or exacerbating marginal compatibility), cumulative biochemical effects (incompatibility-related compounds accumulating over years reaching toxic or disruptive levels), age-related changes (physiological aging processes affecting union differently than adjacent trunk potentially causing problems), and virus interactions (latent viruses potentially affecting union integrity or compatibility over time). Delayed incompatibility symptoms in vine includes= declining vigor (shoot growth progressively reducing over years despite unchanged management), reduced leaf size and chlorosis (leaves becoming smaller and yellower suggesting nutrient or water stress from union dysfunction), poor fruit development (declining yields, poor fruit quality, delayed ripening from inadequate photosynthate or water transport), water stress symptoms (midday wilting, poor water status despite adequate soil moisture suggesting hydraulic dysfunction at union), nutrient imbalances (deficiency or toxicity symptoms from disrupted mineral transport or distribution), and progressive worsening (symptoms intensifying year-over-year rather than stable chronic condition). Delayed incompatibility union morphology includes= progressive swelling increase (union circumference exceeding 130-160% and continuing to enlarge disproportionately), asymmetric overgrowth (excessive tissue formation on portions of union creating pronounced eccentricity), bark abnormalities developing (cracks, cankers, or necrotic zones appearing at union after years of apparently normal bark), and internal deterioration (dissection revealing necrotic zones, vascular discontinuities, tissue disorganization not present in younger unions or initially apparent). Delayed incompatibility distinction from immediate incompatibility includes= delayed showing initial success (union forming, healing, functioning adequately first few years) versus immediate showing failure from onset (poor callus, no integration within weeks-months), delayed showing asymptomatic period (years of normal function) versus immediate showing problems continuously, delayed showing progressive decline (symptoms appearing and worsening gradually) versus immediate showing consistent dysfunction from establishment, and delayed being harder to detect early (requiring long-term monitoring) versus immediate being obvious during nursery or first field season. Delayed incompatibility detection and diagnosis includes= long-term monitoring (tracking vine performance, union appearance over years detecting onset of decline), comparative assessment (comparing suspect unions with well-performing vines or different combinations), union examination (visual inspection and palpation detecting increasing swelling, bark problems, structural weakness), dissection analysis (cutting through declining vine unions revealing internal necrosis, vascular problems), and performance records (yield data, growth measurements documenting progressive decline characteristic of delayed incompatibility). Delayed incompatibility environmental and management interactions includes= stress exacerbation (drought, heat, nutrient deficiency, or disease potentially accelerating delayed incompatibility expression or worsening symptoms), age interaction (delayed incompatibility often becoming apparent as vines reach 8-15 years potentially coinciding with increased crop load stress), crop load effects (heavy cropping potentially unmasking marginal compatibility through increased transport demand), and management impacts (intensive production practices potentially stressing unions accelerating incompatibility expression versus lower-stress systems delaying symptom appearance). Delayed incompatibility in specific combinations includes= known problematic pairs (certain scion-rootstock combinations historically showing delayed incompatibility requiring long-term evaluation before recommendation), variable expression (delayed incompatibility showing incomplete penetrance, some vines affected severely while others acceptable within same combination), site and stress dependence (delayed incompatibility potentially expressing more severely in stressful environments or management systems), and identification challenges (delayed incompatibility difficult to detect during standard 2-3 year nursery evaluation requiring long-term field testing). Delayed incompatibility management implications includes= long-term evaluation necessity (new combinations requiring 10-15 year field trials adequately assessing delayed incompatibility potential), vineyard planning (avoiding combinations with known delayed incompatibility risk in long-term permanent plantings), replacement consideration (vines showing delayed incompatibility symptoms potentially requiring removal and replanting with compatible combinations), and breeding programs (delayed incompatibility being important consideration in rootstock development requiring extended compatibility testing). Delayed incompatibility compared with other decline causes includes= delayed incompatibility showing characteristic union symptoms (excessive swelling, necrosis, union-specific problems) versus trunk diseases showing cankers throughout trunk, delayed showing progressive decline related to union quality versus nutrient deficiency showing improvement with fertilization, delayed showing union structural weakness versus root diseases showing gradual vine weakening from below-ground problems, and delayed requiring union dissection for diagnosis versus other problems diagnosed through other tissues or methods. Delayed incompatibility significance includes= causing premature vine loss (delayed incompatibility leading to decline or death 8-20 years post-planting rather than normal 30-60+ year lifespan), limiting long-term compatibility (some apparently compatible combinations showing problems only after years requiring cautious long-term evaluation), affecting vineyard economics (premature replanting from delayed incompatibility increasing establishment costs and lost production), and emphasizing evaluation importance (delayed incompatibility highlighting need for comprehensive long-term compatibility testing before commercial rootstock recommendations, caution with novel scion-rootstock combinations).
Delayed_Incompatibility_Zone emphasizes late symptom development and is distinguished from Incompatibility_Zone (general failed integration), immediate incompatibility (early failure), and Visible_Incompatibility_Zone (observable symptoms without temporal emphasis) by focus on initially successful union showing progressive deterioration years later. Critical for understanding delayed compatibility expression, long-term union evaluation necessity, progressive decline characteristics, detection and diagnosis challenges, and delayed incompatibility as significant limitation requiring extended testing of new combinations and caution in long-term vineyard planning with marginal scion-rootstock pairings.
Late-developing incompatibility
Latent incompatibility
Progressive Verwachsungs-Störung
Progressive union failure
Spät-auftretende Abstoßung
Verzögerte Inkompatibilität
Delayed Incompatibility Zone
A delayed proleptic shoot on Vitis vinifera cv. Syrah, axillary bud formed at node 6 during previous growing season (May year 1), remained dormant through winter (December-March), remained dormant through normal spring budbreak period (April-May year 2), finally activated in early July year 2 approximately 14 months post-formation, emerged as lateral shoot from previous season's now-mature cane, developed to 38cm length by September with 6 leaves, representing very unusual delayed proleptic pattern attributed to severe hail damage in late June year 2 destroying most current-season shoots and triggering emergency activation of any available bud positions including this long-dormant axillary creating desperate compensatory growth, this delayed lateral providing minimal photosynthetic contribution and poor prospects for productive function but representing vine's survival response utilizing even very delayed dormant buds when catastrophic damage eliminates normal growth.
A proleptic lateral shoot emerging substantially later than normal spring budbreak timing, characterized by axillary bud dormancy through winter and into following growing season, delayed activation occurring mid-summer or later (June-August or beyond), previous-season origin with extended dormancy period before eventual activation, and unusual timing pattern suggesting specific triggers including severe damage creating delayed compensation response, environmental anomalies stimulating late-season bud activation, or variety-specific delayed proleptic tendencies creating unexpected mid-late season lateral emergence from previous-season positions. Delayed proleptic shoots represent extreme temporal delay in lateral development, with axillary buds remaining dormant 12-15+ months from formation through winter and spring into following summer before eventual activation, creating very late-emerging laterals from previous-season wood positions at unexpected times requiring careful assessment of origin and functional role.
Delayed proleptic shoots represent extreme expression of dormant axillary activation, with very extended dormancy periods (12-15+ months) and late-season emergence (summer or later) indicating unusual circumstances triggering activation of buds that would normally remain permanently dormant. The delayed pattern typically results from severe damage creating emergency compensation demands (activating all available bud positions regardless of origin or dormancy state), extreme environmental events (late-season frost, hail, drought stress ending triggering unusual bud responses), or rare variety-specific delayed tendencies. Delayed proleptic laterals emerging mid-late season have limited functional value given short remaining growing period, minimal development potential, and poor prospects for contributing meaningfully to current season or developing quality renewal wood, typically representing desperate vine survival responses rather than planned beneficial growth. Management generally involves removal of delayed proleptics recognizing minimal value, though occasional retention as last-resort renewal options if catastrophic damage leaves no alternatives, with recognition that such shoots indicate severe problems requiring broader vine assessment and recovery planning.
Spät-Achseltrieb
extended-dormancy lateral
late-emerging proleptic
summer-activated dormant lateral
verzögerter proleptischer Trieb
Based on Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press, pp. 85-105 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 113-145.
Delayed Proleptic Shoot
Syrah showing delicate fine feeder roots in irrigated vineyard, soil cores revealing fine roots 0.5-0.9 mm with extremely thin walls and fragile structure, roots collapsing immediately if allowed to dry during processing, mechanical testing showing breakage with <0.5 Newton force, delicate roots comprising 68% of fine feeder root population in well-watered upper soil but only 12% in drier deeper zones, delicate fine feeder roots demonstrating extreme fragility requiring moist conditions and gentle handling
Delicate fine feeder root is fine feeder root emphasizing extreme fragility and vulnerability to damage, characterized by thin-walled tissues easily damaged by mechanical stress, minimal structural reinforcement lacking secondary thickening or lignification, high water content creating turgidity-dependent structure, extreme sensitivity to desiccation and handling, and rapid deterioration when disturbed or stressed, with delicate fine feeder roots showing structural characteristics including thin primary cell walls 0.05-0.15 micrometers lacking reinforcement, minimal or absent lignification maintaining flexible unlignified structure, high proportion of thin-walled parenchyma cells, absence of protective features like suberization or thick cuticle, and fragile root hairs easily broken or collapsed, and demonstrating that delicacy represents functional trade-off where structural investment minimized to reduce costs and maximize absorption efficiency but creating extreme vulnerability requiring careful research methods and showing high sensitivity to environmental stresses or management disturbances. Delicate fine feeder root structural fragility includes= showing thin walls (cell walls extremely thin providing minimal mechanical strength), exhibiting low lignin (virtually absent lignification creating non-woody flexible tissues), demonstrating parenchyma dominance (thin-walled parenchyma cells comprising majority of tissue lacking mechanical cells), revealing minimal reinforcement (lacking sclerenchyma, thick-walled fibers, or other strengthening elements), and displaying turgidity dependence (structural integrity depending critically on cellular turgor collapsing when dehydrated). Delicate fine feeder root mechanical vulnerability includes= showing breakage susceptibility (delicate roots easily broken by minimal mechanical stress during growth or sampling), exhibiting compression damage (delicate tissues easily crushed by soil compaction or careful handling), demonstrating shearing sensitivity (delicate roots damaged by soil movement or cultivation), revealing penetration limits (delicate structure preventing penetration through dense or hard soil layers), and displaying handling requirements (studying delicate roots requiring exceptional care avoiding any mechanical disturbance). Delicate fine feeder root desiccation sensitivity includes= showing rapid water loss (delicate roots losing water quickly when soil dries due to thin cuticle and high surface area), exhibiting turgor dependence (delicate root function critically dependent on adequate turgor wilting rapidly under water stress), demonstrating quick death (delicate roots dying within hours to days if desiccated), revealing no drought tolerance (delicate roots completely intolerant of water deficit immediately losing function), and displaying moisture requirements (delicate root survival and function requiring continuously moist soil conditions). Delicate fine feeder root in soil sampling includes= requiring gentle extraction (delicate roots requiring very gentle soil washing avoiding mechanical damage), showing high loss rates (conventional sampling methods potentially losing 30-80% of delicate roots through breakage), exhibiting preservation challenges (delicate roots difficult to preserve for later study often degrading rapidly after collection), demonstrating immediate examination needs (delicate roots requiring immediate fresh examination before deterioration), and revealing underestimation (delicate root abundance often underestimated in studies due to sampling losses). Delicate fine feeder root stress vulnerability includes= showing drought sensitivity (delicate roots among first to die under water deficit given minimal reserves and high surface area), exhibiting temperature extremes intolerance (delicate roots damaged by freezing or excessive heat), demonstrating salinity sensitivity (delicate roots highly sensitive to salt stress given thin membranes), revealing toxicity susceptibility (delicate roots readily damaged by aluminum, heavy metals, or other soil toxins), and displaying pathogen vulnerability (delicate roots easily invaded by pathogens lacking structural or chemical defenses). Delicate fine feeder root functional implications includes= enabling maximum absorption efficiency (delicacy allowing minimal construction cost maximizing absorption per carbon investment), providing responsiveness (delicate roots rapidly adjusting to conditions through high production and mortality), demonstrating niche specialization (delicate roots functioning optimally only in favorable moist fertile conditions), revealing environmental indication (delicate root abundance potentially indicating favorable soil conditions), and displaying management sensitivity (delicate roots highly responsive to irrigation, mulching, and other practices affecting soil conditions). Delicate fine feeder root trade-offs includes= showing efficiency-persistence trade-off (delicacy providing efficiency but preventing persistence requiring continuous replacement), exhibiting cost-benefit optimization (delicate structure optimal when soil conditions favorable making structural investment unnecessary), demonstrating vulnerability-productivity balance (delicate roots highly productive in good conditions but vulnerable under stress), revealing allocation strategy (producing delicate expendable roots versus fewer robust roots representing different allocation strategies), and displaying ecological context (delicacy successful in stable favorable environments problematic in harsh or variable conditions). Delicate fine feeder root in vine performance includes= determining stress tolerance (delicate root abundance affecting vine drought and stress tolerance), influencing growth patterns (delicate root function supporting growth in favorable periods but limiting under stress), demonstrating irrigation importance (delicate roots benefiting from and requiring consistent irrigation), revealing mulch benefits (mulches protecting delicate roots from desiccation and temperature extremes), and displaying cultivation sensitivity (tillage readily damaging delicate roots reducing function).
Delicate_Fine_Feeder_Root emphasizes fragility and vulnerability to damage and is distinguished from robust roots by focus on extreme delicacy. Critical for understanding thin walls, mechanical fragility, desiccation sensitivity, sampling challenges, stress vulnerability, and the extremely delicate roots optimized for efficiency in favorable conditions but highly vulnerable to damage or stress.
Empfindliche Feinwurzel
Fragile Saugwurzel
Fragile fine root
Tender fine root
Vulnerable feeder root
Zarte Feinwurzel
Delicate Fine Feeder Root
Delphinidin content quality in Syrah from the Northern Rhône reaches notable levels (12-18% of total anthocyanins) reflecting the variety's active F3'5'H expression, with the trihydroxylated delphinidin plus petunidin plus malvidin comprising 85-92% of total anthocyanins this strong trihydroxylation pathway produces the characteristic deep purple-black color of young Hermitage and Côte-Rôtie that evolves toward garnet with age as the less stable delphinidin degrades while methylated forms persist.
Delphinidin_Content_Quality represents the chemical quality characterizing the concentration of delphinidin-based anthocyanins in grape berry skins, capturing the trihydroxylated anthocyanidin that produces blue-violet coloration and serves as the biosynthetic precursor to petunidin and malvidin. Delphinidin (3',4',5',5,7-pentahydroxyflavylium) bears three hydroxyl groups on the B ring at the 3', 4', and 5' positions, representing the most hydroxylated anthocyanidin structure and the direct product of flavonoid 3',5'-hydroxylase (F3'5'H) activity. This trihydroxylation creates the characteristic blue-shifted color compared to dihydroxylated cyanidin and contributes to the purple and violet hues of deeply colored grape varieties. In most V. vinifera cultivars, delphinidin-3-glucoside comprises 5-15% of total anthocyanins, with the majority of trihydroxylated flux proceeding through methylation to petunidin and ultimately malvidin. The delphinidin proportion relative to methylated derivatives (petunidin, malvidin) reflects O-methyltransferase activity, with high delphinidin indicating limited methylation capacity. Delphinidin is the least stable of the five V. vinifera anthocyanidins due to the three unprotected hydroxyl groups, making it susceptible to oxidation and degradation. This instability contributes to the color evolution of wines during aging, as delphinidin-based pigments decline more rapidly than methylated forms. Certain cultivars and species show elevated delphinidin content, producing notably blue-purple coloration distinct from malvidin-dominated varieties.
Delphinidin_Content_Quality addresses the trihydroxylated B-ring anthocyanidin that serves as both a colored pigment and biosynthetic intermediate. The F3'5'H enzyme controlling delphinidin synthesis is a key determinant of grape and wine color, distinguishing deeply colored varieties from those with primarily dihydroxylated anthocyanidins. Delphinidin instability relative to methylated forms affects color aging trajectories wines with high delphinidin proportion may show faster color evolution. The blue-violet hue of delphinidin contributes to the purple character of young red wines, with shifts toward red as delphinidin degrades and malvidin-tannin polymers dominate.
Delphinidin-3-glucosid-Konzentration
Delphinidingehalt
Delphinidinpigmentqualität
Dp-3-glc content
delphinidin pigment quality
delphinidin-3-glucoside content
trihydroxylated anthocyanin concentration
trihydroxylierter Anthocyangehalt
Castellarin SD et al. (2007) Colour Variation in Red Grapevines (Vitis vinifera L.): Genomic Organisation, Expression of Flavonoid 3'-Hydroxylase, Flavonoid 3',5'-Hydroxylase Genes and Related Metabolite Profiling of Red Cyanidin-/Blue Delphinidin-Based Anthocyanins in Berry Skin. BMC Genomics 8(1): 428 Boulton R (2001) The Copigmentation of Anthocyanins and Its Role in the Color of Red Wine: A Critical Review. American Journal of Enology and Viticulture 52(2): 67-87
Delphinidin Content Quality
Chenin Blanc vines at Domaine Huet in Vouvray bearing a Demeter-certified role since 1990 comply with Demeter International standards requiring a minimum of 10% of farm area dedicated to biodiversity reserves, copper usage limited to 3 kg Cu/ha/year, mandatory application of all nine biodynamic preparations (500-508), on-farm compost production, and annual on-site inspection by the Demeter national certification body verifying compliance with 140+ specific production standards.
A biodynamic production role that inheres in a grapevine cultivated on a farm holding valid Demeter International certification, confirming compliance with Demeter production standards including mandatory use of all biodynamic preparations, on-farm composting, biodiversity requirements, and processing restrictions that exceed both organic and general biodynamic practice thresholds.
Demeter_Certified_Role represents the most stringent commercially available viticultural certification standard. Demeter International, founded in 1928, is the oldest ecological certification organization worldwide. The certification imposes requirements beyond general biodynamic practice including mandatory minimum biodiversity set-aside, whole-farm conversion (no parallel conventional production), livestock integration where feasible, and specific processing standards for winemaking that restrict permitted additives more severely than organic wine regulations: for example, maximum SO₂ levels under Demeter are 110 mg/l for dry red wines versus 150 mg/l under EU organic wine rules.
Demeter vineyard role
Demeter-Weinbau-Rolle
Demeter-accredited production role
Demeter-anerkannte Produktionsrolle
Demeter-certified biodynamic role
Demeter-zertifizierte Rolle
Demeter International Production Standards (2022) Demeter International Processing Standards for Wine (2022) Reeve JR et al. (2005) Soil quality and biodynamic farming, Biodynamics 253: 18-25
Demeter Certified Role
Cabernet Sauvignon dense heartwood at 28 years post-planting shows 32-48 mm radius comprising rings 8-18 (wood aged 10-20 years), wood density 0.60-0.66 g/cm³ with 24-28% lignin content, 6-9% phenolic compounds (condensed tannins, resveratrol derivatives), dark reddish-brown coloration, compressive strength 42-52 MPa, intact cellular structure without decay symptoms, provides primary mechanical support for 85-105 mm diameter trunk, contrasts with adjacent sapwood (0.48-0.54 g/cm³) and shows no fungal colonization despite 10-20 year exposure period
Dense heartwood is heartwood characterized by high wood density, extensive lignification, substantial phenolic deposits, and maintained structural integrity without significant decay, representing high-quality structural wood providing maximum mechanical support and durability. Dense heartwood characteristics include= high wood density (0.56-0.70 g/cm³ compared to 0.45-0.58 g/cm³ for low-density or decayed heartwood), extensive lignification (22-30% lignin content compared to 18-24% in sapwood or low-density heartwood), substantial phenolic compound accumulation (5-12% condensed tannins, stilbenes, and other extractives contributing to density and decay resistance), intact cellular structure without decay-related degradation (cell walls and lumens maintaining structural integrity), and darker coloration (brown, reddish-brown, or tan hues from concentrated phenolic deposits). Dense heartwood formation factors include= cultivar genetics (some varieties producing denser wood with higher lignin and phenolic content), slow growth rates (narrow annual rings typically producing denser wood with thicker cell walls and smaller lumens), environmental stress (drought, low fertility producing wood with enhanced lignification and secondary compound deposition), absence of decay organisms (healthy wood maintaining structural integrity without fungal colonization), and appropriate heartwood aging (sufficient time for phenolic deposition and lignification without excessive aging leading to decay). Dense heartwood anatomical characteristics include= thick-walled fiber cells with extensive lignification (providing high mechanical strength), compact cellular structure with small cell lumens (contributing to density), substantial tyloses in vessels (well-developed occlusions filling vessel lumens adding mass), phenolic-impregnated cell walls (tannin and stilbene deposits in walls and lumens increasing density and providing color), and intact ray parenchyma structure (rays maintaining organization despite parenchyma cell death). Dense heartwood mechanical properties include= high compressive strength (35-55 MPa compared to 25-40 MPa in low-density or decayed heartwood), high tensile strength (70-110 MPa compared to 50-80 MPa in lower quality wood), excellent bending strength (supporting trunk flexure under wind load without failure), dimensional stability (resistant to warping, shrinking, or structural deformation), and durability (maintaining properties over decades without significant degradation). Dense heartwood decay resistance includes= enhanced antimicrobial properties (phenolic compounds inhibiting fungal growth and wood decay organism colonization), lignin providing recalcitrant substrate (highly lignified wood more resistant to enzymatic degradation), compact structure limiting fungal penetration (small lumens and dense cell wall matrix restricting hyphal growth), and extractives providing fungitoxic effects (stilbenes, tannins, and other compounds with antifungal activity), though not providing complete immunity (decay still possible but proceeding more slowly than in low-density heartwood). Dense heartwood significance in trunk structure includes= providing maximum mechanical support (dense heartwood contributing disproportionately to trunk strength relative to cross-sectional area), enabling trunk stability in old vines (high-quality heartwood maintaining structural integrity supporting heavy canopies), reducing decay-related structural compromise (decay resistance extending functional structural lifetime), and contributing to trunk durability (dense heartwood lasting decades or centuries without significant degradation in favorable conditions). Dense heartwood distribution in trunk varies= peripheral heartwood often denser (more recently formed heartwood retaining better structure), some radial positions showing higher density (annual rings formed during stress years producing denser wood), central heartwood potentially lower density (oldest wood subject to longest decay exposure, though genetic and environmental factors also important), and cultivar variation (some varieties characteristically producing dense heartwood throughout, others showing more variable wood quality). Dense heartwood contrast with low-density or decayed heartwood includes= 15-60% higher density (0.56-0.70 versus 0.40-0.55 g/cm³), 30-80% higher strength (compression, tension, bending all enhanced in dense wood), 50-300% slower decay rates (dense wood resisting fungal colonization and degradation), darker coloration (higher phenolic content producing richer brown hues), and intact structure (dense heartwood maintaining cellular organization versus collapsed degraded structure in advanced decay). Dense heartwood assessment includes= visual inspection (darker uniform coloration indicating phenolic deposition, firm texture suggesting maintained structure), density measurement (cores or samples showing 0.56-0.70 g/cm³), resistance to physical damage (intact wood resisting penetration, drilling, or crushing), and absence of decay symptoms (no fungal colonization, wood deterioration, or softening). Dense heartwood significance in old vine longevity includes= high-quality heartwood supporting extended productive lifespan (dense structural wood enabling vines to survive 50-100+ years), trunk stability in veteran vines (dense heartwood maintaining support despite extensive age), resistance to catastrophic failure (preventing trunk collapse or breakage), and enabling hollow trunk survival (dense peripheral heartwood maintaining structural integrity even with central cavity formation in extremely old vines).
Dense_Heartwood emphasizes high structural quality and is distinguished from Heartwood (general non-functional wood), Inactive_Heartwood (physiological shutdown emphasis), and Central_Heartwood (position emphasis) by focus on high density, intact structure, and maximum mechanical properties. Critical for understanding old vine structural support, trunk durability and longevity, wood quality variation affecting vine stability, and factors determining trunk mechanical integrity across decades.
Dichtes Kernholz
High-density heartwood
Hochdichtes Holz
Solid heartwood
Structural heartwood
Strukturell intaktes Kernholz
Dense Heartwood
Pinot Noir grafted on Riparia Gloire showing dense root hair zone on fifth-order fine root, zone located 18-32 mm from root apex comprising 14 mm length, hair density 245 per cm substantially higher than proximal zone (140 per cm) or distal zone (95 per cm), microscopic examination revealing closely spaced hairs creating nearly continuous coverage, hair age structure showing 82% of hairs 3-6 days old in peak functional state, isotope pulse-chase demonstrating dense zone providing 34% of total root system nitrogen uptake despite comprising only 3.2% of total root length
Dense root hair zone is root hair zone showing maximum hair density typically 150-300 hairs per centimeter root length, usually located in mid-portion of hair zone approximately 15-35 mm from root apex, characterized by closely spaced hairs creating particularly fuzzy or hairy appearance, showing optimal combination of hair abundance and hair functionality with most hairs in peak functional state, demonstrating maximum absorption capacity per unit root length representing zone of highest resource acquisition efficiency, and representing the focal region for root absorption where close hair spacing, high functional proportion, and optimal hair age combine to create peak uptake performance within root system, with dense root hair zone typically 10-20 mm length occupying central portion of 30-60 mm total hair zone, showing hair density 2-5 fold higher than proximal or distal hair zone portions, and demonstrating that despite being only small fraction of total root system, dense hair zone provides disproportionate contribution potentially 20-40% of total vine water and nutrient uptake from perhaps 2-5% of root system length. Dense root hair zone hair abundance includes= showing high numerical density (typically 150-300 hairs per centimeter substantially higher than proximal or distal zones), exhibiting close hair spacing (hairs spaced 30-70 micrometers apart along root creating nearly continuous coverage), demonstrating multiple hair emergence (potentially 10-25 hairs visible in single microscope field of view at low magnification), revealing epidermal coverage (high proportion of trichoblast cells successfully producing hairs maximizing hair abundance), and displaying uniform distribution (hairs relatively evenly spaced rather than patchy within dense zone). Dense root hair zone age composition includes= showing optimal age distribution (majority of hairs 3-6 days old in peak functional period), exhibiting high functional proportion (typically 70-90% of hairs functional and actively absorbing), demonstrating minimal senescent hairs (relatively few hairs in senescence due to optimal age structure), revealing few developing hairs (relatively few very young hairs still growing as zone downstream of initiation zone), and displaying peak performance composition (age structure optimized for maximum collective hair function). Dense root hair zone absorption performance includes= showing maximum uptake rates (dense zone exhibiting highest water and nutrient uptake per unit root length in entire root system), exhibiting disproportionate contribution (dense zone potentially providing 20-40% of total root uptake from 2-5% of root length), demonstrating synergistic effects (high hair density potentially creating mutually enhancing effects on uptake beyond simple addition), revealing efficiency peak (dense zone showing maximum uptake per unit construction or maintenance cost), and displaying critical importance (dense zone function particularly critical for vine resource status under limiting conditions). Dense root hair zone position includes= located mid-hair zone (dense zone typically 15-35 mm from root apex in central portion of total hair zone), exhibiting optimal distance from apex (position representing balance between sufficient hair age for function and insufficient age for senescence), demonstrating stability relative to apex (dense zone maintaining relatively constant position relative to moving root tip), revealing order-dependent location (dense zone closer to apex in fine fast-growing roots, farther in coarse slow-growing roots), and displaying dynamic movement (dense zone continuously moving basipetally as root grows maintaining position relative to advancing apex). Dense root hair zone surface area contribution includes= showing maximum surface area per length (dense zone creating highest absorption surface area per unit root length), exhibiting substantial area multiplication (dense hairs potentially increasing surface 10-25 fold compared to bare root cylinder), demonstrating critical surface provision (dense zone providing major fraction of functional absorption surface for entire root system), revealing efficiency advantage (high surface area achieved with minimal carbon investment in thin-walled hairs), and displaying absorption correlation (surface area strongly correlated with uptake rates confirming importance). Dense root hair zone environmental responsiveness includes= showing proliferation response (hair density increasing in nutrient-rich zones potentially creating or enhancing dense zones), exhibiting elongation response (individual hairs potentially growing longer in dense zones optimizing absorption), demonstrating stress sensitivity (dense zone vulnerable to drought, compaction, or other stresses reducing density and function), revealing recovery capacity (dense zone potentially recovering after stress relief through resumed hair production), and displaying management importance (practices protecting or enhancing dense zone directly benefiting vine resource acquisition). Dense root hair zone in root system architecture includes= contributing disproportionately to function (dense zones on multiple fine roots collectively providing large fraction of total uptake), exhibiting spatial distribution (dense zones distributed throughout fine root network creating distributed absorption capacity), demonstrating functional specialization (dense zones representing absorption hotspots within root system), revealing architectural optimization (root system architecture potentially evolved to maximize dense zone deployment and function), and displaying management target (optimizing dense zone abundance and function representing key management objective for improving vine resource status).
Dense_Root_Hair_Zone emphasizes maximum hair density and absorption hotspot and is distinguished from lower-density zones by focus on peak hair abundance. Critical for understanding maximum density, optimal age composition, absorption peak, disproportionate contribution, surface area multiplication, and the high-density region providing concentrated absorption capacity representing functional hotspot within root system.
Dichte Wurzelhaarzone
High-density hair zone
Hochdichte Haarregion
Maximale Absorptionszone
Maximum hair zone
Peak absorption zone
Dense Root Hair Zone
Syrah grafted on St. George (Rupestris) rootstock at 32 years post-grafting showing dense rootstock heartwood particularly in central 4 cm diameter core, dense heartwood specific gravity 0.68 g/cm³ significantly higher than typical heartwood 0.55-0.60 g/cm³, dense heartwood showing thick-walled fiber cells with reduced lumen volumes creating compact structure, abundant extractive deposition in cell cavities and walls contributing to high density, dense heartwood dark reddish-brown color from high tannin content characteristic of Rupestris dense wood, moisture content 36% relatively low due to reduced void space, dense heartwood providing excellent compressive strength bearing majority of trunk structural loads, cutting dense heartwood showing high resistance indicating compact tight structure, dense heartwood showing excellent decay resistance with no decay detected despite trunk age, dense wood enabling robust structural foundation supporting large vigorous vine
Dense rootstock heartwood is rootstock heartwood characterized by high wood density typically 0.58-0.70+ g/cm³ specific gravity, emphasizing physical density and structural compactness resulting from thick-walled cells, reduced lumen volumes, extractive deposition in cell cavities and walls, cellular compaction from radial pressure during trunk expansion, and potentially cultivar-specific dense wood anatomy, showing excellent compressive strength and load-bearing capacity providing maximum mechanical support, higher decay resistance from dense structure limiting fungal penetration, increased extractive content from more wood substance per unit volume, and representing either genetically determined dense wood characteristic of certain rootstock cultivars or age-related density increase in oldest central heartwood from progressive compaction and extractive accumulation. Dense rootstock heartwood density definition includes= showing high specific gravity (typically 0.58-0.70+ g/cm³ dry weight basis significantly higher than typical heartwood 0.52-0.62 g/cm³), exhibiting heavy wood (dense heartwood noticeably heavier than typical wood when handling samples or comparing pieces), displaying compacted structure (dense cellular arrangement with reduced void space and tight cell packing), demonstrating substance-rich composition (high proportion of cell wall material and extractives relative to volume), and representing upper range (dense heartwood at high end of wood density spectrum for grapevine tissues). Dense rootstock heartwood cellular basis includes= containing thick-walled cells (fiber cells and vessel elements with thick lignified secondary walls reducing lumen, wall ratio), showing reduced lumen volumes (cell cavities small relative to wall thickness creating compact structure with minimal void space), exhibiting extractive-filled cells (cavities partially or completely filled with deposited phenolics, tannins, and other extractive compounds), displaying compressed cells (cellular deformation from radial pressure during trunk expansion particularly in old central heartwood), and demonstrating dense fiber packing (tightly arranged fiber cells with minimal intercellular spaces creating compact wood structure). Dense rootstock heartwood extractive contribution includes= containing high extractive content (abundant phenolics, tannins, gums, and other compounds depositing in dense heartwood adding mass per unit volume), showing extractive deposition in lumens (compounds filling cell cavities reducing void space and increasing density), exhibiting extractive impregnation in walls (substances depositing within cell walls themselves adding mass to wall structure), displaying progressive extractive accumulation (density increasing over time as additional extractives deposit in aging heartwood), and demonstrating cultivar-specific extractive density (some rootstock varieties producing particularly high extractive levels creating characteristic dense heartwood). Dense rootstock heartwood mechanical properties includes= showing excellent compressive strength (dense heartwood highly resistant to crushing forces providing superior load-bearing capacity), exhibiting high modulus of elasticity (stiff dense wood resisting deformation under load maintaining trunk rigidity), displaying good tensile strength (dense heartwood resisting pulling forces maintaining structural integrity), demonstrating high modulus of rupture (dense wood requiring high bending forces to fail providing excellent flexural strength), and showing increased brittleness (very dense heartwood potentially more brittle and less flexible than lower-density wood cracking more readily under sudden shock). Dense rootstock heartwood structural advantages includes= providing maximum load-bearing capacity (dense heartwood supporting heaviest structural loads in trunk center bearing vine, canopy, and crop weight), enabling thin effective cores (dense heartwood allowing adequate structural support with smaller cross-sectional area than less-dense wood), resisting compression efficiently (high density optimal for bearing axial compressive loads at trunk center), maintaining dimensional stability (dense structure resisting warping, checking, and other dimensional changes), and supporting large old trunks (dense heartwood essential for structural integrity in massive ancient vine trunks requiring maximum strength). Dense rootstock heartwood decay resistance includes= showing improved durability (dense structure creating physical barrier limiting fungal hyphal penetration and spread), exhibiting high extractive protection (dense heartwood containing maximum antimicrobial compounds due to high extractive content per volume), displaying reduced permeability (tight dense structure limiting water and air movement reducing conditions favorable for decay organisms), demonstrating cultivar-related resistance (rootstock varieties producing dense heartwood often showing excellent decay durability), and requiring breach for infection (dense heartwood resisting decay well when intact but potentially vulnerable once damaged allowing decay entry). Dense rootstock heartwood formation mechanisms includes= resulting from genetic wood anatomy (some rootstock cultivars producing inherently dense wood with thick walls and small lumens as genetic trait), developing through aging compaction (progressive cellular compression from radial pressure during trunk expansion increasing density in older central heartwood), accumulating extractives over time (progressive extractive deposition in aging heartwood increasing density particularly in central oldest wood), showing cultivar-specific formation (vigorous rootstocks potentially producing denser heartwood from rapid growth and high extractive production), and demonstrating environmental influences (growing conditions affecting wood density through effects on cell wall thickness and extractive deposition). Dense rootstock heartwood in different rootstock cultivars includes= Rupestris types showing characteristically dense heartwood (thick-walled cells and high tannin content creating dense wood 0.62-0.70 g/cm³ as cultivar trait), Berlandieri hybrids exhibiting variable density (some hybrids showing dense heartwood while others moderate depending on genetic background), vigorous rootstocks like 1103P potentially showing dense heartwood (rapid growth potentially creating thick-walled cells and abundant extractives), SO4 displaying moderately dense heartwood (0.58-0.64 g/cm³ typical for this rootstock), Riparia types showing less dense heartwood (thinner-walled cells and lower extractive content creating lighter wood 0.52-0.58 g/cm³), and cultivar-specific density ranges (rootstock genetics determining characteristic heartwood density reflecting wood anatomy and extractive production). Dense rootstock heartwood positional variation includes= often concentrated in central heartwood (oldest innermost heartwood showing highest density from maximum compaction and extractive accumulation), showing density gradient (density often highest at center declining progressively toward sapwood showing age-related density increase), potentially uniform if genetic trait (cultivar-specific dense wood potentially showing consistent high density throughout heartwood if genetically determined), exhibiting position-dependent density in aged trunk (central heartwood denser than outer heartwood in old trunk reflecting aging effects), and demonstrating variable distribution (dense heartwood distribution varying with whether density primarily genetic or age-related). Dense rootstock heartwood measurement and assessment includes= determining through specific gravity (calculating density from dry weight and volume typically requiring oven-drying and displacement measurement), estimating from wood hardness (dense wood typically harder resisting penetration or indentation), inferring from appearance (dense heartwood often darker color from high extractive content and compact structure), assessing through increment cores (sampling wood and measuring density revealing distribution across trunk radius), and evaluating through cutting resistance (dense wood offering greater resistance to saw or knife providing qualitative density assessment). Dense rootstock heartwood moisture relations includes= showing lower moisture content (dense heartwood 30-55% moisture potentially lower than less-dense heartwood 35-60% due to reduced void space), exhibiting slower moisture equilibration (tight dense structure limiting moisture movement causing slow equilibration with environment), demonstrating reduced hygroscopic range (dense heartwood showing smaller moisture content swings with humidity changes), displaying minimal dimensional changes (dense structure resisting swelling and shrinking maintaining dimensional stability), and maintaining dry stable condition (dense heartwood typically dry and stable contributing to structural reliability). Dense rootstock heartwood in trunk structural function includes= bearing maximum loads efficiently (dense heartwood providing superior load-bearing capacity with minimum volume), enabling trunk strength with smaller diameter (dense heartwood allowing adequate structural support in smaller trunk than less-dense wood would require), resisting compression optimally (high density ideal for bearing axial compressive loads at trunk center), providing long-term structural reliability (dense heartwood maintaining mechanical properties over decades supporting sustained trunk function), and supporting heavy vine structures (dense heartwood adequate for bearing large cordons, extensive canopy, and heavy crop loads in productive mature vine). Dense rootstock heartwood color correlation includes= often showing darker coloration (dense heartwood typically darker brown from high extractive content creating color and density simultaneously), exhibiting color-density relationship (darker color generally indicating denser wood though correlation imperfect), displaying cultivar-specific colors (dense heartwood color ranging from medium brown to dark reddish-brown depending on rootstock extractive chemistry), revealing extractive-driven darkness (color intensity reflecting extractive concentration which also contributes to density), and showing visual density indicator (darker color potentially suggesting denser wood though not perfectly reliable without measurement). Dense rootstock heartwood age relationship includes= increasing density with age in central heartwood (oldest innermost heartwood showing progressive density increase from compaction and extractive accumulation over decades), showing higher density in old trunk (very old trunk central heartwood potentially reaching maximum density 0.65-0.70+ g/cm³ from extended aging), exhibiting age-density gradient (density increasing from outer heartwood toward central oldest heartwood in aged trunk), demonstrating genetic-age interaction (cultivar genetic dense wood trait amplified by age-related compaction creating maximum density in old central heartwood), and potentially stabilizing after maximum (density potentially reaching maximum and stabilizing rather than continuing to increase indefinitely in very old central heartwood). Dense rootstock heartwood advantages and considerations includes= providing superior structural strength (dense heartwood optimal for bearing structural loads maintaining trunk integrity), showing excellent decay resistance (density contributing to durability through physical and chemical protection), enabling efficient structural design (dense heartwood allowing adequate support with minimal volume), contributing to trunk longevity (durable dense heartwood supporting trunk for decades maintaining structural function), potentially limiting processing (very dense heartwood difficult to cut, saw, or work if removal needed), and indicating quality wood (dense heartwood suggesting high-quality durable trunk tissue with excellent structural properties). Dense rootstock heartwood in trunk capacity assessment includes= requiring consideration in structural evaluation (dense heartwood providing more strength per unit area than less-dense wood affecting capacity calculations), affecting heartwood proportion tolerance (trunk with dense heartwood potentially adequate with larger heartwood, sapwood ratio than trunk with low-density heartwood), contributing disproportionately to strength (dense heartwood core potentially providing majority of structural capacity despite comprising minority of trunk volume), influencing replacement decisions (vine with dense heartwood potentially maintaining structural adequacy longer than vine with lower-density heartwood), and indicating trunk quality (presence of dense heartwood suggesting robust structural foundation supporting continued production). Dense rootstock heartwood significance includes= providing maximum structural support per unit volume (dense heartwood offering superior load-bearing capacity making it most efficient structural tissue), showing enhanced decay resistance (density contributing to durability through reduced permeability and high extractive content), representing cultivar-specific trait (dense heartwood characteristic of certain rootstock varieties reflecting genetic wood anatomy and extractive production), demonstrating age-related compaction (density increase in central heartwood showing progressive aging effects over decades), enabling trunk structural longevity (dense durable heartwood supporting trunk integrity for decades or centuries), and indicating superior wood quality.
Dense_Rootstock_Heartwood emphasizes high wood density and structural compactness and is distinguished from Rootstock_Heartwood (all non-functional inner wood without density emphasis) and Central_Rootstock_Heartwood (position emphasis) by focus on high specific gravity, thick-walled cells, extractive content, compaction, and superior mechanical properties. Critical for understanding maximum structural strength, enhanced decay resistance, cultivar-specific dense wood traits, age-related density increase, optimal load-bearing capacity, and the high-density heartwood providing superior structural support and durability in trunk core.
Compact rootstock heartwood
Dichtes Unterlagen-Kernholz
Heavy rootstock heartwood
High-density rootstock heartwood
Hochdichtes Wurzelstockholz
Kompaktes Unterlagenkernholz
Dense Rootstock Heartwood
Gewürztraminer in Alsace Grand Cru vineyards at BBCH 65 exhibits densely packed compact flower clusters of 7-9 cm length bearing 300-450 flowers with flower densities of 35-50 flowers/cm, creating club-shaped masses (OIV code 9) that develop into severely compacted berry clusters with 40-60% rot losses in wet harvest seasons despite 10-12 fungicide applications. Müller-Thurgau (high-density clones) in cool German regions shows similarly extreme compaction with 280-420 flowers in 6-8 cm length, necessitating 50% cluster removal at BBCH 73 followed by aggressive berry thinning to achieve acceptable final compactness, reducing potential yield by 60-70% but enabling viable crop in humid climates.
A densely packed compact cluster is a compact flower cluster exhibiting extreme flower density with flowers so tightly aggregated that individual flowers are difficult to distinguish visually without manipulation, representing the maximum compactness observed in Vitis vinifera cultivars and creating severe disease management challenges while predictably developing into severely deformed berry clusters with high economic losses from rot and splitting. Densely packed compact clusters exhibit flower densities typically exceeding 28-35 flowers per cm of total inflorescence axis length, achieved through maximal flower initiation combined with minimal rachis and branch elongation and very short pedicel length (often <2 mm). These clusters typically measure only 6-10 cm in length at BBCH 65 while bearing 250-500+ flowers, creating nearly continuous flower mass with minimal visible axis. The extreme compactness results from genetic programming for severely reduced internode elongation, typically combined with complete absence of secondary and tertiary branching such that all flowers arise directly from the main rachis or very short primary branches (0.5-2 cm). Densely packed flower clusters characteristically show short cylindrical or bluntly conical shapes with length=width ratios of 1.5=1-2.5=1, creating a 'club-like' or 'pinecone-like' appearance. Flower spacing within densely packed clusters approaches zero, with adjacent flowers essentially touching and pedicels emerging at <1 mm intervals, creating overlapping flower coverage that obscures the underlying axis completely. The extreme flower crowding creates a highly humid, poorly ventilated microclimate with relative humidity often 15-25% above ambient, drying times after precipitation extended by 4-8 hours, and minimal air movement penetrating to interior flowers. This microclimate creates ideal conditions for Botrytis cinerea proliferation, with infection rates at bloom often 5-10× those of loose clusters, and secondary spread from infected flowers to adjacent flowers occurring readily due to physical contact. The transformation from densely packed flower cluster to densely packed berry cluster involves severe architectural consequences, as fruit set of even 30-40% (normal range) creates extreme berry crowding by BBCH 75-77, with berries severely compressed, deformed into polyhedral shapes, skin integrity compromised leading to splitting and secondary Botrytis infection, and bunch rots often destroying 20-50% of potential yield in humid climates. Densely packed flower cluster morphology characterizes cultivars like Gewürztraminer, Müller-Thurgau (extreme selections), and certain Pinot Noir clones, representing OIV code 9 (maximum compactness). Cultural interventions to mitigate densely packed clusters include severe crop load limitation during previous-season bud differentiation to reduce flower initiation, early cluster thinning (BBCH 71-73) to reduce berry number, aggressive berry thinning at véraison to open cluster architecture, and intensive fungicide programs (8-12 applications) focusing on the bloom-to-fruit-set critical period.
Densely_Packed_Compact_Cluster represents the extreme end of the compactness spectrum and is distinguished from Tightly_Compact_Cluster by quantitative density (>28-35 flowers/cm versus 18-28 flowers/cm), visual appearance (flowers obscure axis versus axis still partially visible), and severity of consequences (extreme versus moderate disease pressure and berry deformation). This class corresponds specifically to OIV code 9 (maximum compactness) and represents a serious viticultural liability in humid climates, often rendering affected cultivars economically unviable without intensive cultural and chemical interventions. Some viticulturists consider densely packed cluster architecture a disqualifying trait for organic or low-input production systems. The genetic basis involves multiple loci affecting flower initiation, internode elongation, and branching, with some modern breeding programs actively selecting against this extreme compactness.
extrem kompakte Blütentraube
extremely dense flower cluster
maximal gedrängte Blütentraube
maximally compact flower bunch
sehr dicht gepackte Blütentraube
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 96-98, 146-148 OIV (2009). Descriptor List for Grape Varieties and Vitis Species, OIV code 204 level 9 Vail, M.E., Marois, J.J. (1991). Grape cluster architecture and the susceptibility of berries to Botrytis cinerea. Phytopathology 81=188-191 Hed, B. et al. (2015). Managing Botrytis bunch rot of grapes. Plant Health Progress, doi=10.1094/PHP-2015-1024-01-RS.
Densely Packed Compact Cluster
Gewürztraminer functionally female stigmas exhibit densely papillate organization with 208-245 total papillae distributed across stigma surface (0.54-0.62 mm diameter, giving 240-285 papillae per mm²), with individual papillae 48-68 μm length and 22-32 μm diameter creating elaborate three-dimensional architecture, providing maximum pollen capture efficiency for cross-pollination from hermaphrodite sources with 78-84% of captured pollen germinating during the 6-8 day receptivity period. Wild V. vinifera ssp. sylvestris pistillate female stigmas show densely papillate organization with 185-230 papillae (0.5-0.58 mm diameter, 210-270 papillae per mm²), optimizing pollen capture from stochastic wind-borne pollen arrival from staminate males, with dense papillae creating high probability that incoming pollen grains contact multiple papillae regardless of landing position, achieving 30-42% fruit set when staminate males within 20-50 m provide adequate pollen during overlapping bloom. Thompson Seedless functionally female stigmas exhibit consistently densely papillate morphology with 195-238 papillae (0.52-0.6 mm diameter, 235-290 papillae per mm²) as genetically determined trait associated with functional female sexual expression, with dense papillae supporting cross-pollination-dependent reproduction achieving 25-35% fruit set when hermaphrodite pollinizers provide pollen. Comparative studies of densely papillate functionally female stigmas (220-280 papillae per mm²) versus moderately papillate hermaphrodite stigmas (150-210 papillae per mm²) reveal that dense papillae provide 25-40% greater total stigma surface area (including papillae), support 8-15% higher exudate production per stigma, and achieve 5-12% higher pollen retention after simulated wind exposure, demonstrating functional advantages of dense papillae organization for cross-pollination systems.
A densely papillate stigma is a papillate stigma bearing high papillae density typically exceeding 200 papillae per mm² stigma surface area, characteristic of functionally female and pistillate flowers with elaborated stigmas where dense papillae coverage provides maximum surface area for pollen capture and maximum secretory capacity for exudate production, representing stigmas optimized for cross-pollination systems requiring efficient capture of limited or stochastically arriving pollen. Densely papillate stigmas are characterized by= (1) high papillae number typically 150-250 total papillae on stigma surface (2) high papillae density typically 200-320 papillae per mm² stigma surface, creating closely spaced papillae with inter-papillae gaps often <15-30 μm (3) stigma diameter typically 0.45-0.65 mm providing substantial surface area for the dense papillae population (4) papillae dimensions often toward the larger end of the normal range (45-75 μm length, 20-35 μm diameter) creating particularly elaborate three-dimensional surface architecture (5) stigma surface area (including papillae contribution) typically 200-280% of equivalent flat surface area due to dense papillae coverage. Densely papillate stigmas develop in functionally female and pistillate flowers through enhanced papillae initiation and elaboration during stigma development, potentially reflecting genetic programs that upregulate epidermal specialization and papillae differentiation as part of overall female organ enhancement. The functional significance of dense papillae relates to cross-pollination efficiency= high papillae density creates maximum probability of pollen grain contact with papillae, as incoming pollen grains are likely to encounter multiple papillae regardless of landing position dense papillae provide maximum exudate secretion capacity with more numerous secretory cells producing more total exudate and creating more extensive wet surface dense papillae create maximum surface area increasing pollen retention probability by providing more physical barriers and adhesion points preventing pollen grain loss through wind or gravity dense papillae support simultaneous germination of multiple pollen grains providing statistical advantage in cross-pollination systems where pollen arrival is unpredictable and where genetic diversity in pollen sources may affect compatibility or tube growth success. Densely papillate stigmas in functionally female and pistillate flowers show correlation with extended stigma receptivity duration (typically 6-8 days versus 4-6 days for moderate papillae density hermaphrodite stigmas), suggesting that dense papillae organization may support prolonged exudate production or may reflect overall enhanced female reproductive investment including both papillae elaboration and extended receptivity. Densely papillate stigmas achieve high pollen germination efficiency (75-88% of compatible pollen germinating) comparable to or exceeding moderately papillate hermaphrodite stigmas (70-85% germinating), demonstrating that dense papillae enhance rather than impair pollen germination support despite increased structural complexity.
Densely_Papillate_Stigma represents high papillae density characteristic of functionally female and pistillate flowers and is distinguished from Sparsely_Papillate_Stigma (low density in functionally male flowers) by substantially higher papillae counts per unit area (>200 versus <180 papillae per mm²). Dense papillae correlate with enhanced female function and cross-pollination systems, providing maximum pollen capture and exudate secretion capacity. Understanding densely papillate stigma structure is relevant for identifying functionally female and pistillate flowers, for understanding adaptations to cross-pollination, and for assessing the functional significance of papillae elaboration in female reproductive success.
densely textured stigma
dicht besetzte Papillen-Narbe
dicht papillöse Narbe
heavily papillate stigma
reichlich papillentragendes Stigma
Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=94-109 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 96-101.
Densely Papillate Stigma
Density-based sorting of Cabernet Sauvignon berries using flotation in salt solutions of calibrated specific gravity (1.090-1.100 g/mL) separates optimally ripe berries that sink from underripe berries that float, enabling removal of 15-20% of harvested fruit that would otherwise dilute wine concentration, with sorted lots showing 2-3°Brix higher average sugar content.
Density quality is a physical quality that inheres in grape berry tissues and represents the mass per unit volume relationship characterizing the compactness of cellular organization and the relative proportions of structural, storage, and aqueous components within the tissue matrix. Within the BFO framework, this quality serves as a fundamental physical parameter integrating cellular architecture, biochemical composition, and developmental state into a single measurable attribute that reflects both genotypic determination and environmental modification during berry development. The ontological significance of density lies in its composite nature, where the measured value emerges from the interaction of cell wall material density, vacuolar content concentration, intercellular air space volume, and the relative proportions of distinct tissue types including skin, pulp, and seeds that differ substantially in their individual densities. Grapevine berry density follows a characteristic developmental trajectory, increasing during the pre-véraison phase as cells divide and accumulate organic acids, then showing complex dynamics post-véraison as sugar accumulation increases solute density while simultaneous cell expansion and softening may reduce structural density. This quality is measured through various methodologies including hydrostatic weighing, pycnometry, density gradient flotation, and calculated approaches using mass and volume measurements, with method selection depending on whether whole berry, tissue-specific, or must density is required. The practical significance extends to grape sorting technologies where density-based flotation separates berries by maturity level, to winemaking where must density predicts fermentation dynamics and potential alcohol, and to physiological research where tissue density changes indicate water status, ripening progression, and stress responses. Understanding density quality at multiple scales from cellular to whole berry enables integration of structural and compositional information for comprehensive quality assessment.
Density_Quality serves as the parent class for tissue-specific and measurement-context-specific density qualities, distinguishing the physical property from derived calculations such as must weight scales. This class encompasses true density (excluding air spaces) and apparent density (including air spaces) as child concepts. Measurement methodology significantly affects values and must be specified for meaningful comparison across studies.
Dichte
Gewebedichte
Massendichte
mass-volume ratio
material density
specific mass
spezifisches Gewicht
tissue compactness
Rolle L et al. (2015) Berry density and size as factors related to the physicochemical characteristics of Muscat Hamburg table grapes. Food Chemistry 173: 105-113 Torchio F et al. (2010) Influence of different withering conditions on the chemical composition of Nebbiolo grapes. Food Chemistry 119(4): 1298-1304
Density Quality
Leaf margin character in Müller-Thurgau displays predominantly dentate orientation of teeth, particularly along the lateral lobe margins where teeth project outward perpendicular to the lobe edge rather than curving toward the apex as in more clearly serrate varieties this dentate tendency, combined with medium tooth size (2-3 mm) and relatively sparse spacing, contributes to the distinctive appearance of Müller-Thurgau leaves that facilitated its ampelographic characterization when its parentage (Riesling × Madeleine Royale) remained uncertain prior to molecular confirmation.
Dentate_Margin_Quality represents the margin configuration where the leaf edge bears teeth that project perpendicular to the margin direction rather than pointing toward the leaf apex. Dentate margins are distinguished from serrate margins by this perpendicular orientation of teeth, creating a profile of regularly spaced projections extending outward from the blade edge. In Vitis vinifera, purely dentate margins are less common than serrate margins, but many cultivars exhibit teeth with intermediate orientation or mixed serrate-dentate patterns along different portions of the blade margin. The shape of individual teeth in dentate margins may vary from sharply pointed through blunt to rounded, the latter condition sometimes distinguished as crenate (scalloped) when teeth are very broad relative to their height. Dentate teeth typically display more symmetric form than serrate teeth, with similar curvature on the apical and basal sides of each tooth. The developmental processes producing dentate versus serrate margins involve subtle differences in the orientation and timing of marginal growth centers. Tooth size and density in dentate margins follow similar ranges to serrate margins, with cultivar-specific patterns that contribute to ampelographic identification. Assessment of dentate versus serrate margin type requires examination of tooth orientation relative to the blade margin, which may vary along the margin length with more pronounced directionality near the apex where the margin curves toward the leaf tip.
Dentate_Margin_Quality is distinguished from Serrate_Margin_Quality by tooth orientation, with perpendicular (dentate) versus apically-directed (serrate) teeth providing the diagnostic criterion. Many grape cultivars exhibit intermediate or variable tooth orientation, and the boundary between serrate and dentate involves judgment the classification applies when the predominant orientation is perpendicular. The term crenate describes margins with rounded, tooth-like projections and may be considered a subtype of dentate margin. OIV descriptors do not explicitly distinguish serrate from dentate orientation, focusing instead on tooth size and shape characters. PATO_0001809 (dentate) provides the upper-ontology reference for this margin type.
Zahnrand
dentate Qualität
denticular margin
gezahnte Blattkante
gezähnter Rand
outward-projecting teeth
perpendicular-toothed edge
toothed margin
Hickey LJ (1973) Classification of the Architecture of Dicotyledonous Leaves. American Journal of Botany 60(1): 17-33 Ellis B et al. (2009) Manual of Leaf Architecture, Cornell University Press pp.: 35-42
Dentate Margin Quality
Riesling bears a Dessert_Wine_Scion_Role producing some of the world's finest naturally sweet wines across the German Prädikat categories—Spätlese, Auslese, Beerenauslese, Trockenbeerenauslese, and Eiswein—where its exceptional acidity (10-15 g/L) balances residual sugars of 50-400+ g/L. Sémillon serves in a Dessert_Wine_Scion_Role for Sauternes and Barsac, where thin skin and loose clusters facilitate Botrytis cinerea infection, with concentrated musts of 300-450 g/L yielding wines of 120-200 g/L residual sugar balanced by 6-8 g/L acidity. Furmint bears a Dessert_Wine_Scion_Role for Tokaji Aszú in Hungary, with botrytised berries (aszú) added to base wine in proportion (puttonyos) creating wines of legendary complexity and longevity.
A wine production scion role that inheres in a grafted scion cultivar by virtue of its designation for sweet dessert wine production, where sugar concentration is achieved through natural dehydration, botrytis infection, or freezing rather than by spirit addition. Dessert wine scions must develop extraordinary sugar concentrations—typically 200-450 g/L in must—while retaining sufficient acidity (7-15 g/L) to balance sweetness and preserve freshness in the finished wine. These varieties achieve concentration through compatibility with one or more concentration methods: susceptibility to Botrytis cinerea noble rot (thin-skinned varieties with loose clusters), capacity for extended hang-time for late harvest concentration, tolerance of freezing for icewine production, or suitability for post-harvest drying (appassimento). Dessert wine production is inherently low-yielding (often 1-3 t/ha versus 8-15 t/ha for table wine from the same variety) due to water loss during concentration, making these wines necessarily expensive. The category excludes fortified sweet wines (which achieve concentration through arrested fermentation rather than grape concentration) while encompassing the world's most prized and long-lived sweet wines.
Dessert_Wine_Scion_Role is subclassed by concentration method (Late_Harvest, Noble_Rot, Ice_Wine, Passito), reflecting the distinct viticultural and winemaking requirements of each approach. A single variety may serve multiple dessert wine roles—Riesling produces late harvest, botrytis-affected, and icewine styles depending on vintage conditions and winemaker decisions. The category's shared characteristic is natural sugar concentration through water removal (evaporation, freeze concentration) rather than fermentation arrest, yielding wines typically of 8-14% alcohol where high sugar inhibits yeast activity. Dessert wine production requires specific climatic conditions and involves substantial risk (botrytis may become destructive grey rot, freeze events are unpredictable), contributing to limited supply and high prices for quality examples.
Dessertweinsorte
Süßweinsorte
concentrated wine variety
dessert wine variety
edelsüße Rebsorte
sweet wine variety
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Jackson RS (2014) Wine Science 4th Ed, Academic Press pp.: 530-580 Brook S (2002) Sauternes and Other Sweet Wines of Bordeaux, Mitchell Beazley Rippon S (2009) Tokaji, Mitchell Beazley
Dessert Wine Scion Role
Pinot Noir vineyards at BBCH 65 (full bloom, 50% of flowers open) exhibit abundant detached calyptras visible as yellowish-green caps temporarily suspended on extended stamens of newly opened flowers, accumulating on leaves as tan to light brown dried structures from earlier-opened flowers, and creating a visible layer of fallen calyptras on the ground beneath vine rows, providing clear visual evidence of active bloom in progress and allowing qualitative assessment that bloom is proceeding normally with good flower opening and calyptra shedding. Syrah at BBCH 67 (late bloom, 70-80% of flowers open) shows heavy accumulation of fallen detached calyptras on basal leaves and in cluster zones, with some clusters containing 15-25 trapped calyptras that may require removal through careful canopy management to reduce Botrytis inoculum sources. Chardonnay experiencing rain during bloom shows detached calyptras rapidly washed from leaf surfaces and diluted on ground, reducing visual accumulation but confirming through presence of wet, freshly fallen calyptras that bloom has proceeded despite rainfall. Research studies quantifying bloom timing and intensity sometimes employ systematic collection and counting of detached calyptras from standardized leaf surface areas or ground collection trays, providing objective measurements of bloom progression complementing visual phenological assessments.
A detached calyptra is a calyptra that has completely separated from the receptacle at the basal abscission zone and is no longer attached to the flower, either remaining temporarily suspended on the extended stamens, falling onto nearby vegetative structures (leaves, shoots, inflorescence rachis), or falling to the ground, representing the completion of the anthesis process and the transition to open flower status with fully exposed stamens and pistil available for pollination. Detached calyptras are characterized by= (1) complete basal separation around the entire circumference with no remaining connection to the receptacle (2) calyptra existing as a free structure independent of the flower, appearing as a small (2-4 mm) yellowish-green to brown cap-like or cup-like object with five-parted structure reflecting the fused five petals (3) in freshly detached state, calyptra may remain temporarily perched on top of the extended stamens for minutes to hours before falling or being dislodged by wind (4) in fallen state, calyptras accumulate on leaves, shoot surfaces, inflorescence structures below the flowers, or on the ground beneath vines, creating visible evidence of bloom progression (5) calyptra tissue showing senescence with color changes from yellowish-green through tan to brown as cellular degradation proceeds following detachment. The detached calyptra state represents the completion of the flower opening process and coincides with the functionally open flower capable of performing reproductive functions= stamens fully extended with anthers positioned for pollen release (erect and above pistil in hermaphrodite flowers, reflex and below pistil in functionally female or pistillate flowers) anthers dehiscing through longitudinal slits releasing pollen pistil fully exposed with stigma producing wet exudate and achieving peak or approaching peak receptivity for pollen capture. The timing of calyptra detachment relative to full reproductive maturity shows cultivar-specific patterns= in most hermaphrodite cultivars, calyptra detachment coincides closely with anther dehiscence and stigma receptivity, creating synchronized exposure and functionality in some functionally female or pistillate types, calyptra detachment may slightly precede or follow peak stamen (reflex) positioning, though pistil functionality correlates closely with detachment timing. Detached calyptras provide important visual and quantitative indicators of bloom progression and completion= observation of freshly detached calyptras confirms active anthesis is occurring and indicates flowers have achieved open status accumulation of fallen detached calyptras on leaves and ground provides visual evidence of bloom completion and can be used for qualitative or semi-quantitative assessment of bloom intensity counting or measuring detached calyptras can provide data for phenological studies tracking bloom timing, duration, and synchrony within and among cultivars. The fate of detached calyptras varies with environmental conditions and position= freshly detached calyptras temporarily suspended on stamens typically fall within minutes to hours depending on air movement, with wind facilitating rapid dislodgement while calm conditions may allow extended suspension fallen detached calyptras on leaf surfaces may remain visible for 1-3 days before desiccating and becoming less conspicuous or before being washed off by rain detached calyptras on the ground beneath vines decompose over days to weeks depending on moisture and temperature, contributing minimal organic matter to soil. Detached calyptras can serve as substrate for Botrytis cinerea and other fungal pathogens, with fallen calyptras on leaves or caught in developing clusters potentially providing inoculum sources for subsequent berry infections, making removal of excessive calyptra accumulations through canopy management (leaf removal, cluster zone air circulation improvement) a potential disease management strategy in high-risk situations. The detached calyptra stage encompasses the period from immediate post-separation (freshly detached, within minutes to hours of separation) through degradation and disappearance (fallen and decomposing, days to weeks post-separation), representing BBCH stages 61-69 (flowering progression from 10% through 80% of flowers open and declining) as successive waves of flowers shed calyptras creating continuous accumulation of detached structures throughout the bloom period.
Detached_Calyptra represents the post-anthesis state after complete separation and is distinguished from Separating_Calyptra (still partially attached) by complete independence from the flower structure. The detached state encompasses a continuum from fresh detachment through degradation, with sub-classification into Freshly_Detached (recently separated, still relatively intact and identifiable) versus Fallen_Detached (separated for extended period, degraded and less conspicuous). Detached calyptras serve as important phenological indicators and can provide objective quantitative data for bloom assessment, though their utility for quantification declines as degradation proceeds and as rain or wind removes accumulated structures. Management of detached calyptra accumulations may be relevant for disease control in high-risk situations, though in most cases detached calyptras pose minimal issues and simply represent normal bloom process completion.
abgefallene Kelchhaube
abgelöste Blütenhaube
abgeworfene Kronblattkappe
fallen petal cap
shed calyptra
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 102-104 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 95-97 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 194-196.
Detached Calyptra
Pinot Noir grafted on Riparia Gloire showing detaching border cells at cap periphery, microscopic examination revealing cells partially separated from cap surface with visible intercellular spaces, detaching cells showing transition from columnar to rounded morphology, enzyme assays detecting polygalacturonase and cellulase activity in separation zones, detaching cells showing increased mucilage secretion creating gelatinous coating, cells maintaining viability during separation with intact membranes and active metabolism, detaching process representing programmed developmental transition from attached cap cells to free-living border cells
Detaching border cells are border cells in process of separating from root cap surface, representing transition from attached peripheral cap cells to free-living border cells in rhizosphere, characterized by partial separation with cells loosely connected to cap through degrading middle lamella, active wall loosening through polygalacturonase and cellulase enzymes breaking down cell-cell adhesion, cytoplasmic changes preparing for independent existence including organelle reorganization and metabolic adjustments, and increased mucilage secretion lubricating separation and creating initial gelatinous coating, with detaching phase typically lasting hours to day representing critical transition period where cells complete attachment to root while beginning independent function, showing morphological changes including rounding of previously columnar shape, wall modifications reducing adhesion while maintaining integrity, and cytoplasmic reorganization transitioning from tissue-embedded to free-living state, and demonstrating that detachment represents programmed developmental process rather than passive sloughing creating organized border cell production. Detaching border cells separation mechanism includes= showing middle lamella degradation (enzymes breaking down pectin-rich middle lamella reducing cell-cell adhesion), exhibiting wall loosening (polygalacturonases, pectinases, and cellulases active in separation zones), demonstrating controlled process (cell separation following developmental program with regulated enzyme expression and activity), revealing intercellular spaces (spaces appearing between detaching cells and underlying cap as separation progresses), and displaying mechanical assistance (root growth and soil movement potentially providing mechanical forces completing separation). Detaching border cells morphological changes includes= showing shape transition (cells changing from columnar cap shape to rounded border cell shape during detachment), exhibiting wall modifications (cell walls potentially thickening or modifying composition preparing for independent existence), demonstrating cytoplasmic reorganization (organelles and cytoplasm redistributing during detachment preparing for free-living function), revealing turgor adjustment (cells potentially adjusting osmotic potential and turgor for new mechanical environment), and displaying surface changes (cell surface potentially modifying during detachment affecting interactions with soil and mucilage). Detaching border cells metabolic transitions includes= showing gene expression changes (detaching cells activating genes for independent function and survival), exhibiting protein synthesis (producing proteins for mucilage production, defense, and metabolism appropriate for border cell function), demonstrating enzyme production (synthesizing wall-degrading enzymes facilitating separation and mucilage components for secretion), revealing metabolic adjustments (metabolism potentially shifting from tissue-supported to independent creating self-sufficient cells), and displaying stress responses (detaching cells potentially activating stress pathways responding to separation process). Detaching border cells mucilage production increases includes= showing enhanced secretion (detaching cells increasing mucilage production creating coating), exhibiting Golgi activation (enhanced dictyosome activity producing mucilage vesicles), demonstrating vesicle fusion (increased vesicle fusion with plasma membrane releasing mucilage), revealing accumulation (mucilage accumulating around detaching cells creating initial gelatinous matrix), and displaying lubrication (mucilage facilitating cell separation reducing friction during detachment). Detaching border cells viability maintenance includes= showing continued viability (detaching cells maintaining intact membranes and metabolism during separation), exhibiting energy production (active respiration supporting separation process and continued function), demonstrating membrane integrity (plasma membrane remaining intact and functional during detachment), revealing responsive capacity (detaching cells capable of sensing and responding to environmental signals), and displaying survival preparation (cells preparing for independent survival through metabolic and structural adjustments). Detaching border cells positional characteristics includes= located at cap periphery (detaching cells at outermost cap surface where separation occurs), exhibiting transitional position (cells in intermediate state between attached cap and free border cells), demonstrating surface proximity (detaching cells at or very near root surface in contact with soil), revealing spatial heterogeneity (different cap regions potentially showing different detachment rates creating patterns), and displaying dynamic position (detaching cells continuously being replaced by newly arriving peripheral cells). Detaching border cells timing includes= showing variable duration (detachment process taking hours to day depending on conditions and mechanisms), exhibiting continuous occurrence (detachment continuously occurring as peripheral cells reach separation stage), demonstrating cell-specific timing (individual cells showing variable detachment timing creating asynchronous process), revealing environmental effects (soil conditions potentially affecting detachment rate and timing), and displaying developmental regulation (detachment timing controlled by developmental programs in cap cells). Detaching border cells functional transition includes= showing defensive preparation (detaching cells preparing for defensive function through mucilage and metabolite production), exhibiting sensing capacity (detaching cells potentially beginning to sense soil chemical and biological environment), demonstrating communication (detaching cells potentially receiving or sending signals coordinating with cap and other border cells), revealing adaptive responses (detaching cells potentially adjusting behavior based on soil conditions encountered), and displaying protective function initiation (detaching cells beginning defensive functions even before complete separation).
Detaching_Border_Cells emphasizes active separation process and transitional state and is distinguished from attached cap cells or fully separated border cells by focus on detachment mechanism. Critical for understanding separation process, wall degradation, morphological transition, metabolic changes, mucilage increase, programmed mechanism, and the transitional phase converting attached cap cells into free-living border cells through controlled developmental process.
Loosening cap cells
Separating border cells
Separierende Haubenzellen
Sich ablösende Grenzzellen
Transitional border cells
Übergangs-Grenzzellen
Detaching Border Cells
Syrah developed embryos at seed maturity (BBCH 87-89) exhibit complete structural organization with 2.8-3.2 mm cotyledons, 1.2-1.5 mm radicle, visible shoot apex, organized procambial tissues, and maintain viability for 80-85% germination after 100-day stratification at 4°C
A developed embryo is a fully differentiated embryo within a Vitis vinifera seed that has completed all structural development including cotyledon expansion, radicle formation, and meristem organization, representing an embryo that has achieved complete anatomical maturity regardless of dormancy status or germination capacity. Developed embryos are characterized by= complete cotyledon differentiation (paired, symmetric cotyledons typically 2-3 mm length each with organized storage parenchyma), well-formed radicle with root cap and procambial tissues (typically 1-1.5 mm length), distinct hypocotyl connecting cotyledons to radicle, clearly defined shoot apical meristem (plumule) positioned between cotyledons, and organized vascular tissue connecting embryo organs. Embryo development is complete when the torpedo stage transitions to the mature embryo configuration (approximately 35-50 days post-fertilization, corresponding to BBCH 79-83), after which the embryo undergoes desiccation and enters dormancy without further structural differentiation. Developed embryos occupy 40-60% of seed volume in mature Vitis vinifera seeds, with remaining space filled by endosperm tissue that provides nutritional reserves. Cellular organization in developed embryos includes= storage parenchyma in cotyledons containing accumulated lipids and proteins, organized meristematic cells in radicle root cap and shoot apex, differentiated procambial strands that will form vascular tissues upon germination, and intact cell membranes and organelles indicating cellular viability. The distinction between developed embryos and merely initiated embryos is critical for understanding seed abortion mechanisms= in stenospermocarpic seedless cultivars, embryo development typically arrests at globular or early heart stage (7-21 days post-fertilization) before achieving developed status, while in viable seeds embryo development proceeds through torpedo stage to complete development. Developed embryos in Vitis vinifera exhibit physiological dormancy requiring cold stratification= even fully developed viable embryos will not germinate without extended chilling treatment (90-120 days at 1-5°C), distinguishing grape seed dormancy as physiological rather than morphological. Embryo development status can be assessed through= seed dissection and microscopic examination, X-ray imaging to visualize internal embryo structure, tetrazolium staining to assess tissue viability, or embryo rescue techniques that excise and culture embryos to test developmental competence.
Developed_Embryo emphasizes complete structural differentiation and is distinguished from Embryo (encompassing all developmental stages) and Dormant_Embryo (emphasizing physiological dormancy state) by focus on achievement of full anatomical maturation. Critical for seed viability assessment, understanding abortion timing in seedless mechanisms, embryo rescue breeding techniques, and distinguishing structural development completion from physiological germination readiness.
Ausdifferenzierter Embryo
Fully differentiated embryo
Mature embryo structure
Reifer Keimling
Vollständig entwickelter Keim
Bouquet, A., & Davis, H. P. (1989). Culture in vitro d'embryons immatures de vigne (Vitis vinifera L.) appliquée à la sélection de variétés de cuve sans pépins. Agronomie, 9(6), 565-574. | Ebadi, A., Sedgley, M., May, P., & Coombe, B. G. (1996). Seed development and abortion in Vitis vinifera L., cv. Chardonnay. International Journal of Plant Sciences, 157(6), 703-712.
Developed Embryo
Merlot developing berries progress from fruit set at BBCH 71 (2-2.5 mm diameter, 0.005-0.01 g fresh weight, 8-12 days post-anthesis) through green growth Phase I reaching 9-11 mm diameter and 0.8-1.2 g at BBCH 75 (45-55 days post-anthesis), maintaining size during lag phase at BBCH 77 (60-70 days), undergoing veraison color change at BBCH 81-83 (75-85 days), and completing ripening to mature size 13-16 mm and 1.8-2.6 g at BBCH 89 (110-130 days post-anthesis depending on heat accumulation and vintage conditions). Syrah developing berries show characteristic compositional changes= sugars increasing from 3-5% at BBCH 73 (3 weeks post-fruit set) through 9-11% at veraison (BBCH 81, 10 weeks) to 24-27% at maturity (BBCH 89, 16-18 weeks) titratable acidity peaking at 20-24 g/L during green growth (BBCH 73-75, weeks 3-6), remaining at 18-22 g/L through lag phase (BBCH 77, week 8), then declining during ripening to 5.5-7 g/L at maturity anthocyanin accumulation beginning at veraison and continuing through ripening reaching 1.2-2.8 mg/g skin at maturity. Chardonnay developing berries examined at weekly intervals from fruit set through maturity reveal cell division phase during weeks 1-5 (cell number increasing from 0.8 million to 2.2 million per berry), lag phase during weeks 7-8 (cell number stable, cell size stable at 45-55 μm diameter), and cell expansion phase during weeks 9-15 (cell size increasing to 120-160 μm diameter, cell number constant) demonstrating that final berry size results from cell number established during early development multiplied by cell expansion during ripening. Pinot Noir developing berries show temperature-dependent development rate with harvest maturity achieved at 100-110 days post-anthesis in warm vintages (average growing season temperature 18-20°C) versus 115-130 days in cool vintages (15-17°C), demonstrating environmental modulation of developing berry progression through effects on metabolic rates and developmental processes.
A developing berry is a berry undergoing active growth, differentiation, and compositional changes from initial fruit set through the completion of ripening, representing berries in dynamic developmental states prior to achieving full maturity, encompassing the entire developmental trajectory from ovary retention and expansion (BBCH 71) through veraison and ripening (BBCH 81-89) until harvest-ready maturity is achieved. Developing berries are characterized by= (1) progressive size increases from initial 1.5-3 mm diameter at fruit set to final mature dimensions (6-28 mm depending on cultivar), achieved through two distinct growth phases separated by a lag phase (2) changing tissue composition including increasing dry matter accumulation, changing sugar=acid ratios, evolving secondary metabolite profiles, and shifting hormone balances (3) active cell division during early development (BBCH 71-75, weeks 0-6 post-fruit set) creating final cell number that determines size potential (4) active cell expansion during ripening (BBCH 81-89, weeks 8-18+) increasing cell size 5-20 fold driving final berry enlargement (5) dynamic gene expression patterns controlling developmental transitions including fruit set establishment, growth phase progression, veraison initiation, and ripening processes. Developing berry progression follows the characteristic double-sigmoid growth curve= Phase I rapid growth (BBCH 71-75) through cell division and initial expansion, creating green hard acidic berries reaching 60-80% of final diameter lag phase (BBCH 75-77) with minimal size change while seeds mature and metabolic reprogramming occurs preparing for ripening Phase II rapid growth (BBCH 81-89) through cell expansion and water/solute accumulation, creating softening berries with color development, sugar accumulation, and acid decline. Developing berries show dramatic compositional changes= sugars increase from <5% at fruit set through 8-12% at veraison to 18-28% at maturity titratable acidity peaks at 15-25 g/L during green growth, remains high through lag phase, then declines to 4-10 g/L during ripening pH increases from 2.8-3.2 during green growth to 3.2-4.2 at maturity secondary metabolites including anthocyanins (red cultivars), flavonols, tannins, and aromatic compounds accumulate primarily during ripening. Developing berries are metabolically highly active with= active photosynthesis in green berries contributing 10-15% of berry carbon during early development high respiration rates during growth phases supporting biosynthetic activities declining respiration during lag phase and ripening as metabolic activity shifts from growth to ripening processes active import of photoassimilates (sugars) from leaves through phloem particularly during ripening when berries transition from source (exporting) to sink (importing) organs. Environmental factors strongly influence developing berry dynamics= temperature effects on development rate (growing degree-day accumulation determining development timing), water availability effects on berry size and composition, light exposure effects on secondary metabolite accumulation, and nutritional status effects on growth vigor and composition.
Developing_Berry encompasses the entire progression from fruit set through ripening and represents berries in active developmental states prior to full maturity. Developing berries are distinguished from mature berries (which have completed ripening and achieved harvest-ready composition) and from senescing berries (which are over-ripe and beginning degradation). Sub-classification into Early_Developing_Berry (BBCH 71-75, weeks 0-6, growth Phase I) and Mid_Developing_Berry (BBCH 77-83, weeks 6-11, lag phase through veraison) captures major developmental transitions, with late developing berry (ripening Phase II, BBCH 83-89) representing the final pre-maturity phase. Understanding developing berry biology is essential for crop management, harvest timing prediction, and quality development.
growing grape
immature berry
sich entwickelnde Beere
unreife Traube
wachsende Weinbeere
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 172-210 Coombe, B.G., McCarthy, M.G. (2000). Dynamics of grape berry growth. Australian Journal of Grape and Wine Research 6=131-135 Ollat, N. et al. (2002). Grape berry development= a review. Journal International des Sciences de la Vigne et du Vin 36=109-131.
Developing Berry
Cabernet Sauvignon developing immature seeds at BBCH 81 (beginning of ripening) show yellow-green coloration, 5-6 mm length, 55-60% moisture content, tannin polymerization 45-50%, active tannin biosynthesis in outer integument layers, weekly size increases of 0.4-0.6 mm
A developing immature seed is an actively developing but not yet mature seed of Vitis vinifera that is progressing through normal developmental stages including embryogenesis, endosperm accumulation, and early seed coat differentiation, representing seeds on trajectory toward eventual maturity under favorable environmental conditions. Developing immature seeds are distinguished from incomplete or arrested seeds by evidence of ongoing developmental progression= active cell division and expansion in embryo and endosperm tissues, progressive lignification and tannin accumulation in developing seed coat, sequential transition through expected developmental stages (globular embryo → heart stage → torpedo stage → mature embryo), and increasing seed size and dry matter accumulation over time. These seeds typically occur in normally developing berries during mid-ripening phases (BBCH 75-83), before the final maturation period. Morphological characteristics of developing immature seeds include= progressive color transition from pale green through yellow-green to tan, gradual hardening of testa as lignification proceeds, increasing seed size from 3-4 mm at early stages to 5-7 mm at late immature stages, and developing definition of beak and chalazal structures. Physiological markers include= declining moisture content from 80% at early embryogenesis to 50-60% at late immature stages, increasing accumulation of storage lipids and proteins in endosperm, progressive polymerization of seed coat tannins (transitioning from 20-30% to 50-60% polymerization), and sequential expression of maturation-associated genes and proteins. Developing immature seeds are metabolically active and environmentally responsive= adverse conditions (severe water stress, extreme temperatures, nutrient deficiency) can arrest development and prevent achievement of full maturity, while favorable conditions support continued progression through remaining developmental stages. In viticulture practice, presence of developing immature seeds in berry samples indicates= harvest timing is premature for optimal wine quality, additional vine-ripening time may allow seed maturation to proceed, or environmental conditions may be limiting seed development progression. These seeds are particularly important in cool-climate viticulture where compressed growing seasons may result in harvest before full seed maturation is achieved.
Developing_Immature_Seed emphasizes active developmental progression and is distinguished from Incomplete_Immature_Seed (arrested development) by evidence of ongoing size increases, progressive lignification, and sequential developmental stage transitions. Critical for understanding harvest timing flexibility in cool climates, predicting potential for continued seed maturation with extended hang time, and assessing whether immature seeds reflect normal developmental timing versus arrested development.
Developing green seed
Fortschreitender grüner Kern
In Entwicklung befindlicher Traubenkern
Progressing immature seed
Sich entwickelnder unreifer Samen
Cadot, Y., Miñana-Castelló, M. T., & Chevalier, M. (2006). Anatomical, histological, and histochemical changes in grape seeds from Vitis vinifera L. cv Cabernet franc during fruit development. Journal of Agricultural and Food Chemistry, 54(24), 9206-9215. | Ristic, R., & Iland, P. G. (2005). Relationships between seed and berry development of Vitis vinifera L. cv Shiraz= Developmental changes in seed morphology and phenolic composition. Australian Journal of Grape and Wine Research, 11(1), 43-58.
Developing Immature Seed
Developmental completeness assessment in stressed versus balanced Cabernet Sauvignon reveals the management impact on phenological expression, with balanced vines achieving high completeness (full bud development with 95% bud fruitfulness, complete canopy with 12 m² leaf area/vine, 85% fruit set, harvest at 24.5 Brix with full phenolic maturity), while water-stressed vines show reduced completeness (compromised bud fruitfulness at 72%, incomplete canopy of 8 m²/vine, fruit set only 68%, ripening arrested at 22 Brix with green tannins), demonstrating how stress limits developmental completeness with cumulative effects on yield and quality.
Developmental completeness quality is a phenological quality that represents the degree to which developmental stages achieve full expression before transition to subsequent stages, characterizing whether organs complete all normal developmental processes within each stage or whether development is truncated, arrested, or incompletely expressed due to stress, resource limitation, or other factors disrupting normal phenological progression. Within the BFO framework, this quality addresses the expression dimension of developmental stages, distinguishing whether stages reach complete development from the timing or rate of progression while recognizing that incomplete development at any stage can have cascading effects on subsequent stages and ultimate fruit quality outcomes. The ontological significance lies in developmental completeness determining the quality of each phenological phase's outcomes, where full completion of each stage (complete bud development, full canopy establishment, complete fruit set, thorough ripening) provides the foundation for subsequent stages, while incomplete development creates deficits that may propagate through the season affecting yield and quality at harvest. The factors affecting developmental completeness include resource availability (water, nutrients supporting full development), environmental stress (temperature extremes, disease potentially truncating development), source-sink balance (competition potentially limiting resource supply for complete development), and genetic factors (some cultivars inherently achieving more complete stage expression than others under similar conditions). The stage-specific manifestations of incomplete development include partial bud development (weak buds with poor fruitfulness), incomplete canopy establishment (inadequate leaf area for fruit support), poor fruit set (incomplete fertilization leaving shot berries), and arrested ripening (failure to achieve full maturity before season end), with each manifestation having specific quality implications. The diagnostic significance of developmental completeness lies in its indication of vine condition and management effectiveness, where consistently incomplete development signals chronic stress or resource limitation requiring corrective management, while occasional incompleteness may indicate specific environmental events (frost, heat waves) disrupting normal progression. Understanding developmental completeness quality enables assessment of vine health and balance, diagnosis of factors limiting full phenological expression, and management interventions targeting complete development at critical stages.
Developmental_Completeness_Quality serves as the parent class for expression states (Full Expression, Arrested Development, Incomplete Transition). This quality addresses whether stages achieve normal completion before progression. The class supports vine health assessment and management optimization through completeness characterization.
Entwicklungsexpression
Entwicklungsvollständigkeit
Stadienabschluss
developmental expression
phenological completeness
phänologische Vollständigkeit
stage completion
stage fulfillment
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 59-144 Dry PR, Coombe BG (2004) Viticulture Volume 1: Resources, 2nd Ed. Winetitles, Adelaide, pp. 150-200
Developmental Completeness Quality
Developmental rate assessment comparing warm and cool vintages in Merlot reveals the temperature effect on phenological compression, with warm vintage showing rapid development (budbreak to harvest 155 days, average rate 0.65 BBCH units/day) compressing ripening into August heat, while cool vintage shows slower development (budbreak to harvest 178 days, average rate 0.56 BBCH units/day) extending ripening into cooler September conditions, with the 23-day difference arising from temperature-driven rate variation that affects fruit composition through the thermal conditions experienced during ripening.
Developmental rate quality is a phenological quality that represents the speed of progression through developmental stages, characterizing the temporal dynamics of vine development that determine how quickly organs mature and stages transition, with rate variation arising from genetic differences among cultivars and environmental effects (primarily temperature) that accelerate or slow developmental progress. Within the BFO framework, this quality specifically addresses the velocity dimension of phenological progression, distinguishing rate of development from the stages themselves while recognizing that developmental rate determines the duration of growth phases, the timing of phenological events, and the synchronization of development with seasonal climate patterns. The ontological significance lies in developmental rate determining the temporal compression or expansion of growth phases, where rapid development shortens the time between phenological events (potentially compressing ripening into hotter periods), while slow development extends phase durations (potentially improving flavor development but risking incomplete ripening in short seasons), making developmental rate a key determinant of fruit quality outcomes and climate adaptation. The temperature basis for developmental rate follows approximately linear relationships with temperature above base thresholds, commonly expressed through thermal time concepts where development progress is quantified as degree-day accumulation, with the slope of the rate-temperature relationship and the base temperature varying among developmental phases and cultivars. The genetic control of developmental rate creates the early-versus-late cultivar distinction, where early cultivars (Chardonnay, Pinot Noir) progress rapidly through development requiring fewer degree-days between stages, while late cultivars (Cabernet Sauvignon, Mourvèdre) progress slowly requiring more heat accumulation for equivalent development, with this genetic variation providing the basis for cultivar-climate matching. The phase-specific variation in developmental rate shows different rate sensitivities across the season, with budbreak to flowering showing high temperature sensitivity (rate varying strongly with temperature), flowering duration relatively compressed regardless of temperature, and ripening showing complex rate dynamics affected by both temperature and fruit composition feedbacks. Understanding developmental rate quality enables prediction of phenological timing from temperature forecasts, cultivar selection based on rate characteristics matching site thermal resources, and interpretation of vintage variation in phenological patterns arising from temperature differences affecting developmental rates.
Developmental_Rate_Quality serves as the parent class for organ-specific developmental rates (Bud, Shoot, Inflorescence, Berry). This quality addresses the velocity of phenological progression determined by genetics and temperature. The class supports phenological prediction and cultivar-climate matching through developmental rate characterization.
Entwicklungsgeschwindigkeit
Entwicklungsrate
Wachstumsgeschwindigkeit
development speed
growth velocity
phenological rate
phänologische Rate
stage progression rate
Parker AK et al. (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17(2): 206-216 Duchêne E, Schneider C (2005) Grapevine and climatic changes: a glance at the situation in Alsace. Agronomy for Sustainable Development 25(1): 93-99
Developmental Rate Quality
Developmental stage quality assessment for pesticide timing in Riesling demonstrates the practical importance of precise stage identification, with pre-bloom fungicide application optimally targeted at BBCH 57 (inflorescences fully developed, flowers separating) for Botrytis control, requiring accurate distinction from BBCH 55 (inflorescences clearly visible but still compact, application too early) and BBCH 60 (first flowers open, application potentially too late for preventive protection), with correct stage identification enabling the 2-3 day application window that maximizes spray efficacy for this critical disease control timing.
Developmental stage quality is a phenological quality that represents the current position of grapevine organs or the whole plant within the sequence of defined growth stages, characterizing the discrete developmental states that can be identified and communicated using standardized staging systems enabling precise specification of vine condition for research, management, and regulatory purposes. Within the BFO framework, this quality specifically addresses the categorical dimension of phenological status, distinguishing the identification of current developmental stage from the rate of progression between stages while recognizing that accurate stage identification provides the reference points for phenological modeling, management timing, and communication about vine developmental status. The ontological significance lies in developmental stage quality enabling unambiguous specification of vine condition at any point in the growing season, where standardized stage definitions (BBCH codes) provide the common language for describing whether buds are at woolly bud, green tip, or leaf emergence stages, whether flowering is at first flowers open, full bloom, or cap fall complete, and whether berries are at pea size, veraison beginning, or harvest maturity, with each stage having specific implications for management operations, pest vulnerability, and physiological processes. The hierarchical structure of stage identification employs principal growth stages (BBCH 0-9 representing major developmental phases from dormancy through senescence) and secondary stages within each principal stage (providing finer resolution of developmental progress), with grapevine-specific adaptations defining the observable markers for each stage based on bud, shoot, inflorescence, and berry characteristics. The practical applications of developmental stage identification include timing of pesticide applications (many products stage-specific in efficacy or safety), scheduling of canopy management operations (shoot thinning, leaf removal optimally timed to specific stages), yield estimation (berry counts meaningful only at specific stages), and regulatory compliance (harvest date restrictions based on stage in some appellations). The observer variation in stage assessment can affect precision, with some stages (full bloom, veraison onset) showing clearer identification criteria than transitional stages, making training and standardized assessment protocols important for consistent stage determination across observers and seasons. Understanding developmental stage quality enables precise communication of vine status, appropriate timing of stage-dependent management operations, and accurate phenological recording for modeling and research applications.
Developmental_Stage_Quality serves as the parent class for stage precision and clarity qualities. This quality addresses the categorical identification of current developmental position within standardized staging systems. The class supports management timing and phenological communication through accurate stage characterization.
BBCH stage
BBCH-Stadium
Entwicklungsstadium
Wachstumsstadium
developmental phase
growth stage
phenological stage
phänologische Phase
Lorenz DH et al. (1995) Growth stages of the grapevine: phenological growth stages of the grapevine (Vitis vinifera L. ssp. vinifera)—codes and descriptions according to the extended BBCH scale. Australian Journal of Grape and Wine Research 1(2): 100-103 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages. Australian Journal of Grape and Wine Research 1(2): 104-110
Developmental Stage Quality
Premium Tokaji Aszú production requires Furmint berries affected by Botrytis cinerea noble rot that have lost 40-60% of their water content through berry skin permeabilization, reducing berry equatorial diameter from the healthy 14-16 mm range to 9-12 mm while concentrating sugars to 450-700 g/L, organic acids, and glycerol to levels that define the distinctive Aszú wine style of the Tokaj-Hegyalja region.
Diameter_Quality represents the linear dimension measured across the widest extent of a circular or near-circular cross-section in grapevine organs. Diameter is the preferred measurement term for organs with approximately radial symmetry including shoots, canes, trunks, cordons, roots, tendrils, petioles, pedicels, and for the equatorial measurement of berries. This quality captures the transverse expansion that results from secondary growth in woody organs and primary cell enlargement in herbaceous tissues. Shoot diameter at basal internodes indicates early-season vigor and correlates with bud fruitfulness for the following season. Cane diameter at standardized positions (typically the fourth internode above the basal node) predicts winter hardiness and carbohydrate reserve status. Trunk diameter tracked annually quantifies vineyard establishment progress and long-term productivity trends. Berry equatorial diameter is the standard size measurement for table grapes, with market specifications requiring minimum values (typically 16-18 mm for premium grades). Root diameter distinguishes structural roots (greater than 2 mm diameter) from fine absorbing roots (less than 2 mm), with functional implications for water and nutrient uptake. Diameter measurements employ calipers positioned perpendicular to the organ axis at the point of maximum width. For non-circular cross-sections, two perpendicular diameter measurements enable calculation of a mean diameter and an indication of section ellipticity. Diameter increases during the growing season follow characteristic patterns that reflect carbon allocation, water availability, and sink-source relationships.
Diameter_Quality is the appropriate term for circular cross-sections where length and width would be equal or undefined. For berries, both polar diameter (length) and equatorial diameter (width/diameter) are measured Diameter_Quality specifically refers to the equatorial measurement unless otherwise specified. The class relates to but is distinct from circumference (calculable as π × diameter) and cross-sectional area (π × radius²). PATO_0001334 (diameter) provides the upper-ontology reference.
Durchmesser
Durchmessermaß
Organdicke
Querschnittsbreite
cross-sectional width
organ diameter
radial extent
transverse diameter
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 159-167 Coombe BG and McCarthy MG (2000) Dynamics of Grape Berry Growth and Physiology of Ripening. Australian Journal of Grape and Wine Research 6(2): 131-135
Diameter Quality
41B Millardet et de Grasset (V. vinifera Chasselas × V. berlandieri) demonstrates difficult rooting (30-50% without treatment) reflecting V. berlandieri's inherently poor rooting, requiring IBA treatment and extended callusing to achieve 60-70% success rates commercially, contributing to higher nursery costs and periodic supply limitations. Fercal (complex V. berlandieri hybrid) shows similarly difficult rooting (35-50% without treatment), limiting availability despite exceptional lime tolerance. Pure V. berlandieri selections (Berlandieri Resseguier No. 2, Berlandieri Colombard) are essentially non-propagable from cuttings (<20% rooting), explaining why this valuable species could only be commercially utilized through hybridization with easier-rooting partners.
A rooting capacity rootstock role that inheres in a grafted rootstock by virtue of its low capacity for adventitious root initiation from hardwood cuttings, achieving rooting success rates below 50% under standard nursery propagation conditions and typically requiring specialized treatments to achieve commercially acceptable propagation rates. Difficult-rooting rootstocks show delayed or sparse root primordium formation, high cutting mortality during callusing (often from desiccation or fungal infection before root emergence), and weak initial root system development. This poor rooting capacity reflects unfavorable endogenous hormone balances (low auxin sensitivity, high cytokinin), limited ray parenchyma, anatomical barriers to root primordium emergence, and/or insufficient wound-response cell division. Difficult-rooting rootstocks require auxin treatments (IBA dipping at 2000-4000 ppm), extended callusing periods, mist propagation, bottom heat, or in extreme cases tissue culture micropropagation to achieve commercial production. V. berlandieri parentage is the primary genetic source of difficult rooting.
Difficult_Rooting_Rootstock_Role captures rootstocks achieving below 50% rooting under standard conditions, representing genotypes requiring special propagation protocols or commanding premium nursery prices due to production challenges. V. berlandieri's notoriously poor rooting historically delayed adoption of lime-tolerant rootstocks until crosses with V. riparia and V. rupestris provided commercially viable propagation. Modern tissue culture and improved auxin formulations have partially overcome rooting limitations, but difficult-rooting genotypes remain more expensive and less reliably available than easy-rooting alternatives. The 50% threshold reflects the point below which standard propagation becomes economically marginal without intervention.
difficult rooting rootstock
hard to propagate rootstock
low rooting capacity rootstock
poor rooting rootstock
schlecht bewurzelbare Unterlage
schwer bewurzelbare Unterlage
vermehrungsschwierige Unterlage
Hartmann HT et al. (2011) Hartmann and Kester's Plant Propagation 8th Ed, Pearson Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Pongrácz DP (1983) Rootstocks for Grape-vines, David Philip Publishers Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22
Difficult Rooting Rootstock Role
The diploid status of Pinot Noir clone ENTAV 115 was confirmed by flow cytometry using propidium iodide staining of isolated leaf nuclei on a Partec CyFlow Space cytometer — the DNA content histogram showed a single G1 peak at channel 102 (relative fluorescence units) calibrated against the internal diploid standard Vitis vinifera cv. Seyval Blanc (2C DNA content = 0.95 pg), with a coefficient of variation of 2.1% and no secondary peaks indicative of tetraploid cell populations, confirming the uniform diploid (2n = 38) status throughout the sampled leaf tissue. This diploid confirmation is routinely documented for foundation plant material entering certification pipelines to exclude the possibility of ploidy chimeras or whole-plant tetraploids that could arise during in vitro propagation or through spontaneous somatic doubling.
A ploidy information record confirming the standard diploid chromosome complement (2n = 2x = 38) in a specific Vitis vinifera accession, documenting the normal condition of two complete sets of 19 chromosomes inherited one from each parent, representing the overwhelmingly predominant ploidy state in cultivated and wild grapevine and the baseline against which polyploid and aneuploid deviations are assessed.
Diploid_Status records the confirmation of the normal ploidy state that applies to virtually all commercially cultivated Vitis vinifera. While the diploid state might seem to require no explicit documentation (being the expected default), positive confirmation serves several practical purposes in germplasm management and breeding: first, it establishes the baseline DNA content value for a specific accession against which future ploidy tests can be compared, enabling detection of ploidy changes arising during propagation or tissue culture second, it distinguishes accessions that have been cytogenetically verified from those whose ploidy is merely assumed and third, it provides the reference for interpreting SSR and SNP genotyping results that expect diploid allele dosage patterns (maximum two alleles per locus). The 2C nuclear DNA content of diploid V. vinifera is approximately 0.95-1.00 pg (950-1,000 Mb total, of which approximately 487 Mb is the haploid nuclear genome), with minor cultivar-to-cultivar variation attributable to differences in repetitive sequence content rather than gene number.
2n = 38 status
Diploidie-Status
Normalploidie-Bestätigung
diploid confirmation
diploider Chromosomensatz
standard ploidy record
Jaillon O et al. (2007) The grapevine genome sequence suggests ancestral hexaploidization in major angiosperm phyla, Nature 449: 463-467 Loureiro J et al. (2007) Nuclear DNA content estimations in wild olive (Olea europaea L. ssp. europaea var. sylvestris) and Portuguese cultivars of O. europaea using flow cytometry, Genetic Resources and Crop Evolution 54: 21-25 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519
Diploid Status
Disease incidence data for Riesling clones in the Rheingau documents Botrytis bunch rot incidence at harvest averaging 12% for clone 198 (loose cluster architecture, OIV 204 score 3-5), 22% for clone 239 (moderate density, OIV 204 score 5-7), and 35% for clone 64 (compact clusters, OIV 204 score 7-9) across 10 vintages (2010-2019), establishing a clear clone-by-cluster morphology-by-Botrytis relationship. Disease incidence data for Merlot clones at Bordeaux records downy mildew (Plasmopara viticola) severity under reduced spray programs (50% of standard) ranging from 15% leaf area affected in clone 181 to 38% in clone 348, suggesting differential field tolerance potentially linked to leaf hair density and stomatal characteristics, though no clones exhibited true genetic resistance.
Disease incidence data is a disease performance data entity that records the frequency, severity, and temporal progression of fungal, bacterial, phytoplasmal, and physiological disease symptoms observed on a clonal line under field evaluation conditions, expressed as incidence (percentage of affected organs — leaves, clusters, shoots) and severity (percentage of organ area affected or standardized disease rating scales), measured across multiple vintages with and without chemical protection, characterizing the clone's relative susceptibility or field tolerance to the major grapevine pathosystem in the evaluation region.
Disease incidence data provides the field-level complement to laboratory virus status, capturing the clone's interaction with the full spectrum of regionally relevant pathogens under realistic growing conditions. The methodology of disease assessment must be carefully standardized — trained assessors using reference diagrams (e.g., the Horsfall-Barratt scale for foliar diseases, the OIV 459 scale for Botrytis bunch rot) provide more reproducible data than subjective estimation, though inter-assessor variability remains a recognized limitation. The distinction between disease incidence (proportion of affected units) and disease severity (degree of damage per affected unit) provides complementary information — a clone may show high incidence but low severity (many clusters lightly affected) or low incidence but high severity (few clusters devastated), with different implications for crop loss estimation. Environmental conditions during the evaluation period critically influence absolute disease levels, making clone ranking (relative performance) more reliable than absolute incidence values for cross-site and cross-vintage comparison. Clone-specific morphological traits that influence disease incidence include cluster compactness (the dominant factor for Botrytis susceptibility), leaf hair density (affecting downy mildew infection), cuticle thickness, berry skin thickness, and stomatal density. Unsprayed or reduced-spray trial blocks provide the most discriminating assessment of clonal disease tolerance but risk catastrophic crop loss in high-pressure vintages, leading most programs to employ split-plot designs comparing full and reduced protection within the same trial.
Anfälligkeitsbewertung
Befallshäufigkeitsdaten
Feldkrankheitsdaten
Krankheitsbefallsdaten
Krankheitsschweredaten
disease incidence record
disease severity data
disease susceptibility rating
field disease data
pathogen incidence record
Dry PR et al. (2010) Bunch architecture and the susceptibility of Vitis vinifera to Botrytis cinerea. In Proceedings of the 7th International Workshop on Grapevine Downy and Powdery Mildew, IOBC-WPRS Bulletin 67: 53-56 Vail ME, Marois JJ (1991) Grape cluster architecture and the susceptibility of berries to Botrytis cinerea. Phytopathology 81(2): 188-191 Kennelly MM et al. (2007) Primary infection, lesion productivity, and survival of sporangia in the grapevine downy mildew pathogen Plasmopara viticola. Phytopathology 97(4): 512-522
Disease Incidence Data
Disease performance data for Pinot Noir clone 115 (Dijon) documents virus-free certification status (negative for GFLV, GFkV, GLRaV-1, GLRaV-3, GVA, and GVB by ELISA and RT-PCR), moderate susceptibility to Botrytis cinerea (mean incidence 18% of clusters at harvest across 10 Burgundy vintages, compared with 28% for clone 114 under identical conditions), and average susceptibility to Plasmopara viticola requiring standard spray program adherence. Disease performance data for Grüner Veltliner clone 17 at Klosterneuburg records complete virus-free status confirmed by biological indexing on Vitis rupestris St. George and molecular testing, Botrytis incidence 35% below population mean attributable to its loose cluster architecture (OIV 204 score 5), and powdery mildew severity ratings consistently at or below regional clone averages.
Disease performance data is a performance data entity that documents the phytosanitary status, pathogen susceptibility, and disease tolerance characteristics of a clonal line, encompassing viral infection status determined through serological and molecular diagnostics, fungal and bacterial disease incidence and severity ratings under field conditions, and the clone's relative vulnerability or tolerance to major grapevine pathogens compared with population averages and reference clones, providing essential health information for certification decisions and grower risk assessment.
Disease performance data occupies a unique position within clone evaluation because phytosanitary status, particularly virus infection, simultaneously affects all other performance parameters — yield, quality, and agronomic behavior are all compromised by chronic viral infection, making virus status determination a prerequisite for meaningful interpretation of any other performance data. The distinction between virus status (presence/absence of specific viruses in the clonal material itself) and field disease incidence (susceptibility to airborne, soilborne, and vector-transmitted pathogens under growing conditions) represents two fundamentally different information categories within disease performance. Virus-free status achieved through sanitation protocols (heat therapy, meristem culture) requires ongoing monitoring because reinfection through vector transmission (nematodes for GFLV, mealybugs and scale insects for leafroll viruses) can compromise the sanitized material in commercial vineyards. Field disease tolerance shows significant clone-by-environment interactions — a clone rated as Botrytis-tolerant in a dry Mediterranean evaluation site may prove highly susceptible under humid oceanic conditions, necessitating regional validation of disease performance data. The increasing availability of disease-resistant interspecific cultivars carrying Rpv and Ren resistance loci is shifting disease performance evaluation toward molecular characterization of resistance gene content alongside traditional field observation.
Gesundheitsdaten
Krankheitsanfälligkeitsdaten
Krankheitsleistungsdaten
Pathogentoleranz
disease evaluation data
disease tolerance record
health performance data
pathogen susceptibility data
phytosanitary performance
phytosanitäre Leistungsdaten
Mannini F (2004) Clonal selection in grapevines: interactions between genetic and sanitary strategies. Acta Horticulturae 652: 21-27 Martelli GP (2014) Directory of virus and virus-like diseases of the grapevine and their agents. Journal of Plant Pathology 96(1S): 1-136 Maree HJ et al. (2013) Grapevine leafroll-associated virus 3. Frontiers in Microbiology 4: 82
Disease Performance Data
Pinot Noir clone 777 bears a Disease_Resistant_Clone_Role regarding botrytis, its loose cluster architecture reducing berry-to-berry contact and humidity retention that promotes bunch rot—enabling later harvest in maritime climates without the grey rot losses affecting tight-clustered clones. Riesling clone 198 serves in a Disease_Resistant_Clone_Role for reduced botrytis susceptibility in German production, selected for cluster looseness permitting dry-climate-like harvest windows in the humid Rhine and Mosel valleys. Chardonnay clone 96 bears a Disease_Resistant_Clone_Role regarding millerandage resistance, with more consistent fruit set than millerandage-prone clones—though some producers value millerandage for the complexity contributed by uneven berry sizes.
A performance clone role that inheres in a grapevine clone characterized by enhanced resistance or tolerance to specific diseases relative to other clones of the same variety, reducing disease pressure and fungicide requirements while maintaining varietal identity. Disease-resistant clones may exhibit reduced susceptibility to fungal diseases (powdery mildew, downy mildew, botrytis), viral diseases (through field tolerance though not genetic resistance), or physiological disorders (coulure, millerandage where undesirable), with resistance mechanisms including tighter cluster architecture reducing botrytis entry points, thicker cuticle limiting pathogen penetration, enhanced induced defense responses, or unknown genetic factors conferring reduced susceptibility. Disease resistance within V. vinifera clones is typically quantitative (degree of susceptibility) rather than qualitative (complete immunity), contrasting with the major resistance genes found in wild Vitis species. Disease-resistant clone selection supports sustainable viticulture by reducing fungicide requirements—particularly valuable for organic and integrated pest management production—while maintaining the varietal identity and wine character lost when resistant interspecific hybrids replace V. vinifera varieties.
Disease_Resistant_Clone_Role captures within-variety variation in disease susceptibility, a dimension increasingly valued as viticulture seeks sustainability through reduced chemical inputs. The resistance characterization is disease-specific a clone resistant to botrytis may show typical susceptibility to powdery mildew, requiring selection decisions considering the specific disease pressures of each site. Resistance within V. vinifera is generally modest compared to wild species, with "resistant" clones showing reduced rather than absent disease development. The category complements but does not replace the disease resistance breeding programs developing interspecific hybrids with major resistance genes V. vinifera clones offer familiar wine character while resistant varieties offer stronger protection at the cost of changing varietal expression. Climate change is increasing disease pressure in some regions, elevating the importance of disease-resistant clone selection for long-term vineyard sustainability.
disease-resistant clone
hardy clone
krankheitsresistenter Klon
low-susceptibility clone
tolerant clone
toleranter Klon
widerstandsfähiger Klon
Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 Vail ME, Marois JJ (1991) Grape cluster architecture and bunch rot susceptibility, American Journal of Enology and Viticulture 42: 57-64 Dry IB et al. (2010) Molecular strategies to enhance disease resistance in grapevine, Australian Journal of Grape and Wine Research 16: 94-105
Disease Resistant Clone Role
Disease severity quality assessment for powdery mildew (Erysiphe necator) in Napa Valley Cabernet Sauvignon employed standardized protocols combining incidence and intensity measures to guide fungicide program decisions and predict quality impacts. Pre-véraison assessment at BBCH 75 evaluated disease severity quality across 50 vines per block with systematic cluster and leaf sampling. Cluster disease severity quality showed 35% incidence (proportion of clusters with any visible infection) combined with mean intensity of 12% surface coverage on affected clusters, yielding an overall severity index of 4.2% (incidence × intensity). Leaf disease severity quality showed 65% incidence with 18% mean intensity on affected leaves, severity index 11.7%. The disease severity quality thresholds established for the region indicated that cluster severity index above 5% at véraison correlated with significant wine quality reduction (reduced varietal character, elevated volatile acidity risk), triggering intensified late-season spray coverage for blocks approaching this threshold. Post-harvest analysis confirmed the disease severity quality-quality loss relationship, with blocks exceeding 8% cluster severity index showing 15-point reduction in wine quality scores compared to blocks maintained below 3% through effective fungicide programs.
Disease severity quality is a pathogen infection indicator quality that inheres in a grapevine organism and represents the quantitative assessment of disease impact combining measures of how widespread infection is across the plant population or plant organs (incidence) with how intense or damaging infection is within affected units (intensity), providing the integrated evaluation of disease status required for management decisions, yield loss estimation, and efficacy assessment of control measures. This quality captures the multidimensional nature of disease assessment, recognizing that comprehensive evaluation requires both the proportion of plants or organs affected and the degree of damage within those affected units, since disease with high incidence but low intensity may have different management implications than disease with low incidence but high intensity in affected individuals. The ontological significance of disease severity quality lies in its role as the primary endpoint for disease management decision-making, with severity assessments determining whether disease levels warrant intervention, whether applied controls have been effective, and what yield or quality losses may be expected from observed disease levels. The biological basis of disease severity encompasses the epidemiological processes governing pathogen spread that determine incidence, the host-pathogen interaction dynamics that determine intensity within infected tissues, and the relationship between these disease parameters and the functional consequences for plant productivity and product quality. The standardization of severity assessment methods enables consistent evaluation across locations, seasons, and assessors, with established rating scales, reference diagrams, and assessment protocols providing the methodological framework for reproducible severity quantification. The relationship between severity measures and crop loss is often non-linear, with threshold effects where low severity causes minimal impact while severity above critical thresholds causes disproportionate damage, and with the timing of infection relative to crop development affecting the severity-loss relationship such that equivalent severity earlier versus later in the season may have very different yield and quality consequences.
Disease_Severity_Quality is_a Pathogen_Infection_Indicator_Quality, serving as parent class for Infection_Incidence_Quality, Infection_Intensity_Quality, and Lesion_Coverage_Quality which address specific severity components. This class addresses the integrated assessment combining incidence and intensity while child classes capture individual metrics. The severity index (incidence × intensity) provides composite assessment. Threshold severity levels triggering management decisions are pathogen- and crop-specific. The severity-loss relationship enables yield and quality impact prediction from disease assessment data.
Befallsstärke
Infektionsschwere
Krankheitsschwere
Pathogenschaden-Ausmaß
disease assessment
disease damage level
infection severity
pathogen impact severity
Seem RC (1984) Disease incidence and severity relationships. Annual Review of Phytopathology 22:133-150 Madden LV et al. (2007) The Study of Plant Disease Epidemics. APS Press Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests, 2nd Edition. APS Press
Disease Severity Quality
Disease tolerance quality comparison across V. vinifera cultivars evaluated the relationship between Botrytis cinerea infection level and yield loss to identify cultivars with superior tolerance maintaining productivity under equivalent disease pressure. Controlled inoculation experiments established uniform B. cinerea infection at 25% cluster severity across cultivars, then assessed the yield and quality impacts of this standardized disease level. Semillon demonstrated superior disease tolerance quality with 25% infection severity causing only 15% yield reduction and wines maintaining varietal character with only modest quality reduction, attributed to the ability of remaining healthy berries to compensate through enhanced individual berry growth and effective juice recovery from partially affected clusters. In contrast, Pinot Noir showed poor disease tolerance quality with 25% infection causing 40% yield reduction and substantial quality compromise including off-flavors and color loss, reflecting the thin-skinned cultivar's inability to limit damage progression and compensate for affected berries. The disease tolerance quality assessment established tolerance indices (yield retention at standardized infection level) ranging from 0.85 for Semillon to 0.60 for Pinot Noir, with the tolerance differential not explained by differences in defense response capacity but rather by physiological and morphological factors affecting damage compensation. The disease tolerance quality characterization informed cultivar recommendations for regions with high B. cinerea pressure, favoring tolerant cultivars where infection control is difficult and targeting limited fungicide resources to intolerant premium cultivars where protection provides greatest return.
Disease tolerance quality is a pathogen infection indicator quality that inheres in a grapevine organism and represents the capacity to maintain productivity and performance despite pathogen infection, achieving acceptable yield and quality outcomes even when pathogen colonization occurs at levels that would substantially impair susceptible cultivars, with tolerance representing a distinct resistance strategy from defense responses that limit pathogen growth by instead emphasizing the maintenance of plant function under disease pressure. This quality captures the performance-maintenance dimension of disease resistance, recognizing that from a practical viticulture perspective the critical outcome is crop production rather than pathogen population reduction per se, and that some cultivars achieve acceptable performance by tolerating infection rather than preventing it. The ontological significance of disease tolerance quality lies in its representation of an alternative resistance strategy that may provide more durable protection than defense-based resistance, since tolerance mechanisms do not impose selection pressure on pathogen populations to evolve virulence and thus avoid the resistance-breaking dynamics that can render defense-based resistance ineffective. The biological basis of disease tolerance encompasses the physiological capacity to compensate for pathogen-induced damage through enhanced photosynthetic rates in unaffected tissues, efficient resource reallocation from damaged to productive organs, maintenance of water and nutrient transport despite vascular colonization, and the suppression of disease-induced senescence that would otherwise accelerate tissue loss. The practical assessment of disease tolerance requires evaluation under equivalent pathogen pressure, comparing the relationship between infection level and performance impact among cultivars to identify those maintaining yield and quality at infection levels causing substantial losses in intolerant cultivars. The distinction between resistance (limiting infection) and tolerance (limiting impact of infection) is critical for disease management strategy, with tolerant cultivars potentially requiring different fungicide programs than resistant or susceptible types since the goal shifts from infection prevention to management of infection within tolerable levels.
Disease_Tolerance_Quality is_a Pathogen_Infection_Indicator_Quality, serving as parent class for Symptom_Suppression_Quality and Yield_Maintenance_Under_Infection_Quality which address specific tolerance mechanisms. This class addresses performance maintenance under infection distinct from pathogen limitation. Tolerance avoids resistance-breaking selection pressure on pathogens. Assessment requires evaluation under equivalent pathogen pressure to distinguish tolerance from resistance. Tolerance index (performance retention at standardized infection) provides quantitative metric. Different management implications than resistance-based protection.
Ertragstoleranz
Infektionstoleranz
Krankheitstoleranz
disease endurance
infection tolerance
pathogen tolerance
yield tolerance
Pagán I, García-Arenal F (2018) Tolerance to plant pathogens: theory and experimental evidence. International Journal of Molecular Sciences 19:810 Ney B et al. (2013) Crop tolerance to diseases: concepts, measurement, and application. Phytopathology 103:1163-1173 Roy BA, Kirchner JW (2000) Evolutionary dynamics of pathogen resistance and tolerance. Evolution 54:51-63
Disease Tolerance Quality
Thompson Seedless (Sultanina) exhibits pronounced distal fruitfulness disposition with peak fertility (2.0-2.8 clusters) at nodes 8-14 and minimal fruitfulness (0.3-0.6 clusters) at nodes 1-4, necessitating cane pruning for commercial production. Flame Seedless shows similar patterns with optimal fruitfulness at nodes 10-16. Guyot and other cane-pruned systems retain 8-15 node canes to capture distal fruitfulness, with typical cane-pruned Sultanina yields of 15-25 t/ha compared to only 4-8 t/ha under spur pruning. The fruitfulness gradient from base to apex may span a 4-6 fold difference in strongly distal cultivars.
A bud fruitfulness disposition that is the inherited capacity of a Vitis vinifera plant to develop high numbers of inflorescence primordia in buds at distal node positions (nodes 8-15 or beyond from the cane base), requiring cane pruning or long spur retention to achieve adequate crop levels.
Distal fruitfulness patterns evolved in wild Vitis as an adaptation to climbing growth where basal portions are shaded. Domestication selection for spur-trainable cultivars has reduced but not eliminated distal fruitfulness patterns. Distal fruitfulness complicates mechanized pruning and requires skilled manual cane selection for optimal production. Some cultivars show bimodal fruitfulness with peaks at both basal and distal positions.
Bogenfruchtbarkeit
apical bud fertility
cane fruitfulness
distal node fertility
distale Fruchtbarkeit
triebspitzige Knospenfertilität
Antcliff AJ et al. (1972) The relationship between bud position and fruitfulness in Sultana. Vitis 11:140-151 Huglin P, Schneider C (1998) Biologie et écologie de la vigne, 2nd Ed, Lavoisier pp.: 256-278
Distal Fruitfulness Disposition
Merlot on SO4 showing distal second-order lateral shallow roots, excavation 120-180 cm from trunk revealing second-order laterals 0.7-1.6 cm diameter originating from distal first-order portions, distal second-order extending 20-55 cm into inter-row alley, laterals showing 1-3 annual rings, producing third-order branches creating peripheral absorptive network, distal second-order lateral shallow roots demonstrating exploratory function accessing peripheral resources beyond central root system domain
Distal second order lateral shallow root is second-order lateral shallow root emphasizing distal position far from trunk originating from distal portions of first-order laterals, typically 80-200+ cm from trunk base, characterized by peripheral location at outer edge of lateral root system exploration zone, origin from mature or aging portions of first-order parents, smaller diameter 0.5-2 cm reflecting late formation or limited development, and serving exploratory function extending root system reach into previously unexplored or weakly explored soil zones potentially between rows or into alleys, with distal second-order laterals showing characteristics including origin from first-order lateral portions 80-200+ cm from trunk often near first-order tips, emergence from mature first-order tissues with well-developed bark, smaller size than proximal second-order branches reflecting developmental constraints or later formation timing, shorter length 15-60 cm compared to proximal second-order 30-100 cm, and pioneering role exploring peripheral zones accessing resources beyond central root system domain. Distal second order lateral shallow root positional characteristics includes= showing peripheral location (distal second-order laterals at outer edge of lateral root system 80-200+ cm from trunk), exhibiting inter-row extension (distal laterals potentially reaching into alleys or spaces between vine rows), demonstrating depth maintenance (distal laterals maintaining shallow depth 10-50 cm even at peripheral locations), revealing exploration frontier (distal laterals representing leading edge of lateral root system expansion), and displaying competition (distal laterals potentially encountering neighboring vine roots in peripheral zones creating competitive interactions). Distal second order lateral shallow root size limitations includes= showing smaller diameter (distal second-order laterals typically 0.5-2 cm smaller than proximal 1-3 cm), exhibiting limited secondary growth (distal laterals showing minimal secondary thickening potentially 1-3 annual rings), demonstrating shorter length (distal laterals often 15-60 cm shorter than proximal reflecting developmental constraints), revealing reduced vigor (distal laterals potentially showing lower growth rates than proximal due to distance from trunk and resource limitations), and displaying structural simplicity (distal laterals showing simpler structure with less extensive vascular development). Distal second order lateral shallow root developmental constraints includes= showing resource limitation (distal laterals potentially limited by long transport distance from trunk reducing photosynthate availability), exhibiting hydraulic limitation (water transport resistance from trunk to distal laterals potentially limiting turgor and growth), demonstrating competition effects (distal laterals competing with extensive proximal root network for limited resources), revealing parent aging (distal laterals emerging from aging distal first-order portions potentially showing reduced support from senescing parent tissues), and displaying late formation (distal laterals often forming later than proximal branches starting at developmental disadvantage). Distal second order lateral shallow root exploratory function includes= providing peripheral exploration (distal laterals extending root system into peripheral zones beyond proximal and mid-range exploration), exhibiting resource discovery (distal laterals potentially accessing resources unavailable to more proximal roots due to spatial separation), demonstrating competitive advantage (distal laterals potentially capturing resources before neighboring vine roots reach same zones), revealing opportunistic growth (distal laterals potentially showing rapid growth if encountering resource-rich patches in peripheral zones), and displaying expansion potential (distal lateral development potentially increasing total root system exploration volume 20-40%). Distal second order lateral shallow root third-order production includes= showing moderate branching (distal second-order laterals producing third-order branches though potentially less abundantly than proximal), exhibiting branching pattern (third-order branches emerging every 5-12 cm along distal second-order length), demonstrating absorptive support (third-order and higher derivatives from distal second-order providing peripheral absorption), revealing local network (distal laterals creating localized absorptive networks in peripheral zones), and displaying contribution (distal lateral-associated fine roots potentially providing 10-30% of total uptake accessing peripheral resources). Distal second order lateral shallow root in inter-row dynamics includes= showing alley exploration (distal laterals potentially extending into inter-row alleys accessing resources in these spaces), exhibiting cover crop interaction (distal laterals potentially competing with cover crop roots in alleys for water and nutrients), demonstrating fertilizer access (alley-applied fertilizers potentially accessed more efficiently by distal than proximal laterals), revealing irrigation capture (irrigation water in alleys potentially captured by distal lateral networks), and displaying management relevance (alley management practices directly affecting distal lateral function and distribution). Distal second order lateral shallow root vulnerability includes= showing exposure risk (distal laterals potentially more exposed to disturbance from alley traffic or operations), exhibiting drought sensitivity (distal laterals in peripheral zones potentially experiencing more severe drought than proximal roots in central zone), demonstrating competition pressure (distal laterals facing strong competition from neighboring vine roots in overlapping peripheral zones), revealing mortality risk (distal laterals potentially showing higher mortality than proximal due to peripheral challenges), and displaying replacement difficulty (distal lateral losses potentially difficult to replace given developmental constraints and distance from trunk). Distal second order lateral shallow root lifespan patterns includes= showing variable persistence (distal second-order laterals potentially surviving 2-8 years shorter than first-order but longer than fine roots), exhibiting turnover (distal lateral population showing moderate turnover as some laterals die and new ones form from advancing first-order tips), demonstrating progressive formation (distal laterals continuously forming as first-order laterals elongate creating continuous wave of peripheral exploration), revealing environmental sensitivity (distal lateral survival strongly affected by peripheral zone conditions including moisture, nutrients, competition), and displaying population dynamics (distal lateral abundance varying seasonally and annually responding to conditions and vine resource status).
Distal_Second_Order_Lateral_Shallow_Root emphasizes peripheral distal position far from trunk and is distinguished from proximal laterals by focus on outer exploration zone. Critical for understanding peripheral exploration, developmental constraints, inter-row extension, competitive dynamics, and the distal laterals extending root system into peripheral zones accessing resources beyond central domain.
Distal lateral root
Distale zweite Ordnung Lateralwurzel
Far-reaching lateral branch
Peripheral second-order root
Periphere Seitenwurzel
Weit-reichender Lateralast
Distal Second Order Lateral Shallow Root
Ugni Blanc (Trebbiano Toscano) bears a Distillation_Scion_Role as the dominant variety for Cognac production (over 98% of plantings), with its high acidity (9-11 g/L), neutral flavor, moderate sugar (17-19°Brix yielding 8-9% base wine), disease resistance, and high yields (100-120 hL/ha) providing ideal characteristics for double pot-still distillation. Folle Blanche serves in a Distillation_Scion_Role for Armagnac, contributing delicate floral aromatics that survive single continuous distillation to create distinctive spirit character. Quebranta bears a Distillation_Scion_Role as the primary non-aromatic variety for Peruvian Pisco, where its neutral character and Muscat-free heritage define the Pisco Puro Quebranta category.
A scion role that inheres in a grafted scion cultivar by virtue of its designation for distilled spirit production, where wine or fermented grape must is distilled to concentrate alcohol and volatile flavor compounds into brandy, eau-de-vie, or grape-based spirits. Distillation scions are selected for characteristics fundamentally different from table wine production, prioritizing high acidity (preserving freshness through distillation), neutral-to-delicate flavor profiles (avoiding coarse or overwhelming varietal character that would concentrate unpleasantly), moderate sugar content (target base wines of 8-10% alcohol minimize methanol formation), and high yields (economic viability requires substantial volume as distillation concentrates 7-10 liters of wine into 1 liter of spirit). The distillation process concentrates desirable volatiles (ethanol, esters, higher alcohols) while excluding water and non-volatile compounds, with spirit character developing through the interaction of grape-derived congeners, distillation technique (pot still versus column still, single versus multiple distillation), and aging (oak maturation for aged brandies). Major distillation traditions include Cognac (France), Armagnac (France), Pisco (Peru/Chile), brandy (global), and grappa (Italy).
Distillation_Scion_Role is subclassed by regional spirit tradition (Cognac, Armagnac, Pisco, Grappa, general Brandy), reflecting distinct regulatory frameworks, permitted varieties, distillation methods, and aging requirements. The economic logic of distillation favors high-yielding, disease-resistant varieties whose base wines would produce unremarkable table wine—Ugni Blanc exemplifies this, being the world's most planted white variety primarily for distillation rather than drinking. Distillation concentrates both desirable and undesirable compounds variety selection and careful viticulture minimize precursors of harsh or unpleasant volatiles (methanol from pectin, higher alcohols from nitrogen stress). Climate change affects distillation regions by increasing sugar accumulation and reducing acidity in base wines, challenging traditional spirit profiles.
Brandweintraube
Brenntraube
Destillationstraube
Spirituosen-Rebsorte
distillation grape
eau-de-vie variety
spirit grape
Léauté R (1990) Distillation in alembic, American Journal of Enology and Viticulture 41: 90-103 Piggott JR et al. (1989) Distilled Beverage Flavour, Ellis Horwood Galy B et al. (2008) Cognac and its Production, in Bentley R (ed) The Alcoholic Beverages, Kluwer Academic Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane
Distillation Scion Role
The distribution record for Pinot Noir clone 115 (ENTAV) documents global dissemination through 47 licensed nurseries across 12 countries, with propagation rights administered by ENTAV under the French catalogue system, current availability status of "widely available" in France, USA, New Zealand, and Australia, and documented commercial plantings spanning Burgundy, Champagne, Oregon, Marlborough, Yarra Valley, Central Otago, and Sonoma Coast — making it one of the most globally distributed premium Pinot Noir clones. The distribution record for Blaufränkisch clone Bl-2 documents distribution primarily through Austrian nurseries (Rebschule Tschida, Rebschule Schober, Rebveredlung Rittsteuer), with propagation rights held domestically under Austrian Rebenverkehrsgesetz provisions, current availability as "readily available" within Austria and "limited availability" internationally, and commercial plantings concentrated in Burgenland, Carnuntum, and emerging sites in the Czech Republic and Hungary.
Distribution record is a clonal selection record entity that documents the commercial dissemination pathway, legal propagation framework, material availability status, and geographic spread of a certified clonal line from the holding institution through nursery multiplication to end-user vineyard plantings, encompassing the nurseries authorized to propagate the clone, the intellectual property and licensing arrangements governing its multiplication, the current supply status in regional and international markets, and the geographic regions where the clone has been established in commercial production.
Distribution records track the transformation of a clonal line from a scientific achievement into a commercially available agricultural input, documenting the complex interplay of intellectual property law, nursery business relationships, regulatory authorization, and market demand that determines which clones reach growers and where. The distribution pathway from certification authority to vineyard involves multiple intermediaries — foundation block holders, nuclear stock custodians, licensed propagation nurseries, grafting operations, and regional distributors — each maintaining documentation that contributes to the complete distribution record. The international dimension of clone distribution is governed by phytosanitary import/export regulations, quarantine requirements, and material transfer agreements that vary substantially between jurisdictions, creating asymmetric availability patterns where clones readily available in one country may be difficult or impossible to obtain in another. The lag between certification and widespread availability can span 5-15 years, as nurseries require time to build stock from limited initial propagation material through successive multiplication cycles to volumes sufficient for commercial supply. Distribution records also capture the reverse flow of information — grower feedback on clone performance under diverse commercial conditions provides valuable post-certification data that complements the controlled trial results from evaluation sites.
Distributionsakte
Klonvertriebsdaten
Verbreitungsnachweis
Verteilungsdokumentation
Vertriebsnachweis
clone availability record
clone distribution record
dissemination record
distribution documentation
propagation and distribution data
Mannini F (2004) Clonal selection in grapevines: interactions between genetic and sanitary strategies. Acta Horticulturae 652: 21-27 Boidron R et al. (1995) Catalogue des variétés et clones de vigne cultivés en France, ENTAV-INRA, pp.: 25-35 Nicholas P (2004) Grapevine propagation. In Viticulture Volume 1: Resources, Winetitles, pp.: 213-235
Distribution Record
The diurnal range data for contrasting viticultural climates demonstrates the influence of maritime versus continental climate on thermal amplitude — Bordeaux (Médoc, maritime) shows a mean DTR of 8-10°C during the August-September ripening period (Tmax 25-28°C, Tmin 15-18°C), reflecting the thermal buffering of the Atlantic Ocean and Gironde estuary that moderates both day maxima and night minima Mendoza (Luján de Cuyo, continental) shows a mean DTR of 15-18°C during the corresponding February-March ripening period (Tmax 30-33°C, Tmin 14-17°C), reflecting the arid continental climate at 900 m elevation with unimpeded radiative cooling under clear skies. The wide diurnal range in Mendoza produces Malbec with intense color (the cool nights promoting anthocyanin stability), preserved acidity (reduced nocturnal malic acid respiration maintaining TA at 5-7 g/L at harvest sugars of 24-26°Brix), and the combination of ripe fruit flavor with structural freshness that distinguishes Argentine Malbec from warmer, narrow-amplitude Malbec production regions.
A temperature data component that records the daily temperature amplitude — the difference between the daily maximum temperature (Tmax) and the daily minimum temperature (Tmin) — averaged over the growing season or over specific phenological periods (particularly the ripening period from véraison to harvest), with diurnal temperature range (DTR) being a critical determinant of fruit acid retention (cool nights reducing nocturnal malic acid respiration), anthocyanin accumulation (the thermal amplitude promoting pigment synthesis and stability), and the overall balance between sugar maturation and aromatic/phenolic development that defines wine quality potential.
Diurnal_Range_Data documents the thermal parameter that most directly influences the balance between sugar accumulation and acid preservation during the ripening period — arguably the most important temperature characteristic for wine quality after the mean temperature level. The physiological basis of the diurnal range effect centers on the differential temperature sensitivity of sugar accumulation (driven by daytime photosynthesis, relatively insensitive to nighttime temperature) versus acid degradation (driven by respiratory consumption of malic acid in mitochondria, highly sensitive to nighttime temperature following the Q10 principle — the rate approximately doubling for each 10°C increase): warm days promote photosynthesis and sugar import to the berry, while cool nights slow the respiratory consumption of malic acid, producing the combination of high sugar and high acid that defines balanced must composition — conversely, warm nights accelerate acid respiration without contributing to sugar accumulation, producing the high-sugar, low-acid musts characteristic of hot continental and tropical viticultural climates. The Tonietto and Carbonneau (2004) MCC system formalized the diurnal range concept as the Cool Night Index (CI), defined as the mean minimum temperature of September (northern hemisphere), with CI 14°C as warm nights (unfavorable for balanced ripening) — this index captures the critical nighttime temperature conditions during the final month of ripening when acid preservation and aromatic compound accumulation are most sensitive to temperature.
Tag-Nacht-Schwankung
Tagestemperaturamplitude
day-night temperature range
diurnal temperature amplitude
diurnaler Temperaturbereich
thermal amplitude record
Tonietto J and Carbonneau A (2004) A multicriteria climatic classification system for grape-growing regions worldwide, Agricultural and Forest Meteorology 124: 81-97 Jackson DI and Lombard PB (1993) Environmental and management practices affecting grape composition and wine quality, American Journal of Enology and Viticulture 44: 409-430 Gladstones J (1992) Viticulture and Environment, Winetitles Kliewer WM (1973) Berry composition of Vitis vinifera cultivars as influenced by photo- and nycto-temperatures during maturation, Journal of the American Society for Horticultural Science 98: 153-159
Diurnal Range Data
Syrah grafted on 110 Richter rootstock showing dividing root apical meristem on primary root, microscopic examination of longitudinal sections revealing numerous mitotic figures, mitotic index calculated 21% of meristem cells in M phase, metaphase figures showing 38 condensed chromosomes (2n=38) aligned on metaphase plate with spindle fibers visible, anaphase figures showing sister chromatid separation, telophase figures showing chromosome decondensation and nuclear envelope reformation, cytokinesis evident with cell plates forming between daughter cells, dividing root apical meristem producing estimated 86 new cells per day supporting root elongation rate 7.2 mm per day, cell cycle duration estimated 16 hours during June active growth
Dividing root apical meristem is root apical meristem emphasizing cell division process and mitotic activity, characterized by cells actively progressing through mitosis with observable chromosome condensation, spindle formation, and cytokinesis, showing visible mitotic figures including prophase metaphase anaphase and telophase stages in microscopic sections, demonstrating cell plate formation creating new cell walls between daughter cells, continuous production of pairs of daughter cells from dividing mother cells doubling meristem cell population or producing derivatives, and quantifiable mitotic index typically 8-25% of cells in M phase during active growth reflecting division frequency, with dividing root apical meristem representing the fundamental growth process where single cells become two through nuclear division and cytoplasmic partitioning creating new cells that either remain meristematic maintaining meristem size or exit meristem becoming derivatives that elongate and differentiate into functional root tissues, and demonstrating that cell division in root apical meristem is ultimate cellular mechanism enabling root growth through continuous production of new cells throughout growing season. Dividing root apical meristem mitotic phases includes= showing prophase (chromosomes condensing becoming visible as distinct structures with nuclear envelope breaking down), exhibiting metaphase (condensed chromosomes aligning at cell equator on metaphase plate with spindle fibers attached to kinetochores), demonstrating anaphase (sister chromatids separating and moving toward opposite cell poles pulled by shortening spindle fibers), revealing telophase (chromosomes decondensing at poles with nuclear envelopes reforming around chromosome sets), and displaying cytokinesis (cell plate forming at equator growing outward creating new cell wall dividing cytoplasm producing two daughter cells). Dividing root apical meristem mitotic index includes= showing variable division frequency (mitotic index ranging from 5-10% in slowly growing meristems to 15-25% in rapidly growing meristems), exhibiting order-related patterns (primary root meristems often showing higher mitotic index than fine root meristems reflecting vigor differences), demonstrating seasonal variation (mitotic index highest spring and fall declining summer and approaching zero winter dormancy), revealing environmental effects (favorable conditions temperature moisture nutrients increasing mitotic index while stress reducing), and displaying measurement methods (mitotic index determined by counting mitotic cells as percentage of total cells in microscopic sections). Dividing root apical meristem spindle apparatus includes= forming microtubule spindle (tubulin polymerization creating spindle fibers connecting centrosomes to chromosome kinetochores), showing bipolar organization (spindle organized with two poles at opposite cell ends creating symmetric chromosome separation), demonstrating kinetochore attachment (spindle microtubules binding to protein complexes on centromeres ensuring accurate chromosome segregation), exhibiting dynamic instability (microtubules continuously growing and shrinking adjusting chromosome position), and revealing motor proteins (dynein and kinesin moving along microtubules generating forces for chromosome movement). Dividing root apical meristem cell plate formation includes= initiating at cell center (phragmoplast microtubules and vesicles organizing at former metaphase plate position), showing vesicle fusion (Golgi-derived vesicles carrying cell wall materials fusing to form cell plate), exhibiting centrifugal growth (cell plate expanding outward from center toward parent cell walls), demonstrating plasma membrane insertion (vesicle membranes contributing to plasma membranes of daughter cells), and completing at walls (cell plate reaching and fusing with parent walls completing cytokinesis producing two separate cells). Dividing root apical meristem chromosome segregation includes= ensuring equal distribution (mitotic mechanisms ensuring each daughter cell receives identical chromosome complement), showing diploid maintenance (grapevine 2n=38 chromosomes duplicated in S phase segregated equally in mitosis maintaining chromosome number), demonstrating accurate segregation (spindle checkpoint mechanisms preventing division until all chromosomes properly attached), revealing occasional errors (rare mis-segregation events potentially producing aneuploid cells though usually eliminated), and displaying fidelity importance (accurate chromosome segregation essential for maintaining genetic integrity in growing root). Dividing root apical meristem daughter cell fates includes= producing meristematic daughters (some divisions generating two daughter cells both remaining in meristem maintaining population), creating derivative cells (other divisions producing daughters that exit meristem entering elongation zone and differentiation), demonstrating asymmetric divisions (particularly near quiescent center divisions creating one stem cell and one derivative maintaining organization), revealing stochastic patterns (individual cell fates probabilistic with overall population dynamics regulated by position and signals), and showing tissue specification (daughter cells adopting specific fates dermatogen, periblem, or plerome based on position determining tissue contribution). Dividing root apical meristem cell cycle regulation includes= requiring cyclin-CDK complexes (cyclins and cyclin-dependent kinases driving cell cycle progression through checkpoints), showing G1/S transition (cyclin D-CDK4/6 and cyclin E-CDK2 promoting DNA synthesis initiation), exhibiting G2/M transition (cyclin B-CDK1 triggering mitosis entry), demonstrating checkpoint control (surveillance mechanisms ensuring proper completion of each phase before proceeding), and revealing growth factor regulation (auxin, cytokinins, nutrients affecting cyclin-CDK activity modulating division rate). Dividing root apical meristem energy requirements includes= demanding substantial ATP (mitosis requiring energy for spindle dynamics, chromosome movement, and cytokinesis), showing elevated respiration (dividing cells consuming oxygen and producing CO2 at high rates generating ATP), exhibiting biosynthesis needs (producing histones, tubulins, membrane lipids, and cell wall materials for daughter cells), demonstrating nucleotide requirements (DNA replication and chromosome duplication requiring dNTP pools), and revealing metabolic intensity (dividing meristem showing 2-5× higher metabolic rate than differentiated tissues reflecting growth demands). Dividing root apical meristem division timing includes= showing cell cycle duration (complete cycle from one division to next typically 10-24 hours in active meristem depending on conditions), exhibiting phase lengths (G1 3-8 hours, S 4-8 hours, G2 2-4 hours, M 1-2 hours with variation among cells and conditions), demonstrating temperature dependence (cycle duration decreasing with increasing temperature from threshold to optimum then increasing above), revealing resource effects (adequate nutrients and carbohydrates supporting rapid cycling while limitations extending cycle duration), and displaying developmental variation (meristem cells near quiescent center potentially showing longer cycles than peripheral cells). Dividing root apical meristem in root growth includes= producing cell number increase (continuous division creating expanding cell population providing material for root elongation), enabling tissue formation (divided cells exiting meristem differentiating into functional root tissues), demonstrating growth rate correlation (higher division frequency generally producing faster root elongation though elongation also depends on cell expansion), revealing cumulative effect (thousands to millions of divisions throughout growing season producing substantial root length increase), and showing indeterminate capacity (meristem maintaining division capacity throughout vine life enabling continuous root growth for years or decades).
Dividing_Root_Apical_Meristem emphasizes mitotic process and cell division mechanics and is distinguished from non-dividing meristem by focus on active mitosis. Critical for understanding cell division mechanism, mitotic phases, chromosome segregation, cell plate formation, daughter cell production, division frequency, and the fundamental mitotic process creating new cells enabling root meristem function and continuous root growth.
Cell-dividing apical meristem
Mitotic root meristem
Mitotisches Wurzelmeristem
Proliferating root meristem
Proliferierendes Wurzelmeristem
Teilendes Wurzel-Apikalmeristem
Dividing Root Apical Meristem
Chardonnay in Champagne shows division phase completion during late July-early August Year N+1 at BBCH 75, with final mitotic events completing week 4-5, mitotic index declining to <1%, complete division cessation establishing non-dividing 1.5-1.9 million cell population, growth mode transition to expansion exclusivity, berry diameter 16-19 mm at completion, under cool maritime conditions of 20-25°C temperature, typical of this early-bursting cultivar completing division phase enabling progression to expansion
Division phase completion represents the definitive cell proliferation cessation period during berry cell division when final mitotic events conclude, complete cell division cessation establishes non-dividing cell population, growth mode transition establishes expansion as exclusive mechanism, and division phase ends enabling berry progression to expansion-dominated growth phase, occurring by weeks 4-5 post-fertilization marking division phase conclusion. This phase termination period occurs during late summer in late July-early August in temperate regions Year N+1, typically completing weeks 4-5 representing division cessation, corresponding to BBCH 75 berry touch stage. Completion mechanism involves final mitotic events in last dividing cells producing terminal divisions, complete division cessation with all cells exiting cycle to expansion state, growth mode transition establishing expansion as sole growth mechanism replacing division, developmental checkpoint passage enabling progression to expansion phase, and completion marking definitive end of proliferative period. Final mitotic events characteristics show last observable divisions occurring week 4-5, mitotic index declining to very low <1-2% indicating rare remaining divisions, late-dividing cells completing final cycles with extended 16-18 hour cycle times, and eventual complete division cessation by week 5-6. Division cessation timing shows typical completion week 4-5 for majority of berry cell population, occasional rare divisions extending to week 6 in small subpopulations, cultivar variation in completion timing affecting total division duration, and completion establishing non-dividing state. Cell division cessation shows mitotic activity ceasing completely, cell cycle genes downregulating to basal levels, proliferation markers disappearing indicating non-cycling state, all cells in G0/G1 expansion state versus S/G2/M cycling states, and cessation as definitive proliferation end. Post-cessation cell state shows cells in G0 quiescent or G1 growth state committed to expansion, cells lacking division competence unable to re-enter cycle under normal development, expansion program activated with enlargement-associated genes upregulated, and cells fully committed to growth through enlargement rather than proliferation. Growth mode transition establishment shows division contribution to growth ceasing completely, expansion as exclusive growth mechanism throughout subsequent development, berry growth from week 5 onward entirely through cell enlargement not proliferation, and transition as fundamental growth mechanism shift. Expansion exclusivity shows no further cell number increase throughout expansion, lag, veraison, ripening phases, berry volumetric growth entirely from cell volume increase, and expansion as sole mechanism for 8-12 weeks until harvest. Division phase duration summary shows total division phase spanning 4-5 weeks from fruit set, active division weeks 0-3, declining division weeks 3-5, completion by week 4-5, and total duration relatively constant across cultivars with timing variation. Developmental progression shows division completion enabling berry advancement to expansion phase, sequential phase progression characteristic of berry development, completion as prerequisite for subsequent phases, and developmental program directing phase transitions. Checkpoint passage shows developmental checkpoints ensuring appropriate division completion before expansion phase progression, adequate cellularity achievement required for progression, checkpoint mechanisms likely involving auxin levels and cell number, and passage enabling phase transition. Molecular markers of completion show cell cycle gene downregulation to basal levels, proliferation marker PCNA expression loss, expansion gene upregulation indicating enlargement commitment, and transcriptional reprogramming marking completion. Morphological indicators show berry diameter 14-25 mm at completion depending on cultivar, berry transitioning from young green fruit to expanding berry appearance, and size at completion reflecting achieved cellularity. Berry composition at completion shows water content 80-85% fresh weight, sugars low 3-6% from metabolic consumption, organic acids 1-1.5% particularly malate and tartrate, pH 2.8-3.2, and green acidic composition characteristic. Irreversibility of completion shows division phase non-repeatable during berry development, established cell number unchangeable throughout berry life, and completion as permanent developmental state transition. Environmental independence after completion shows subsequent environmental conditions unable to reinitiate division, favorable later conditions unable to compensate for limited division phase, and completion establishing constraints for subsequent development. Berry development progression shows transition from division phase completion to expansion phase onset, berry entering rapid expansion period following completion, and sequential progression through expansion, lag, veraison, ripening phases. Cell population uniformity after completion shows all cells in expansion state, population synchrony in growth mode, and uniformity enabling coordinated subsequent expansion. The process shows three distinguishable aspects including final mitotic events representing terminal divisions, cell division cessation establishing non-dividing state, and growth mode transition to expansion exclusivity. Duration represents completion period weeks 4-5. Regional timing shows division phase completion during late July-early August across climates marking division end. Viticultural significance includes representing definitive end of cell proliferation establishing permanent cellularity, marking critical developmental transition to expansion-dominated growth, demonstrating irreversible completion preventing subsequent division, and establishing division phase as discrete developmental period. Research applications include analyzing completion mechanisms and regulation, investigating developmental checkpoint controls, evaluating completion timing variation and determinants, quantifying transcriptional reprogramming at completion, and assessing division phase completion as determinant of subsequent berry development.
Division_Phase_Completion represents definitive division cessation within Berry_Cell_Division_Phase. Subdivided into Final_Mitotic_Events with terminal divisions, Cell_Division_Cessation establishing non-dividing state, and Growth_Mode_Transition to expansion exclusivity. Completion weeks 4-5. Mitotic index declining to <1%. All cells in G0/G1 expansion state. Irreversible cessation. Marks transition from division to expansion phase. Completes Berry_Cell_Division_Phase. Enables progression to Berry_Expansion_Phase.
Proliferations-Terminierung
Teilungs-Beendigung
Teilungsphase-Vollendung
division cessation
division completion
mitotic phase end
mitotische Phasenende
proliferation termination
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 440-443 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Division Phase Completion
Cabernet Sauvignon in Napa Valley shows division rate reduction during mid-late July Year N+1 at BBCH 73-74, with peak division rate 95,000 cells per day week 2.5 declining to 48,000 cells per day week 3.5 and 22,000 cells per day week 4.5, mitotic index declining from 12% to 6%, cycling cell fraction reducing from 78% to 32%, cell cycle times extending from 11 hours to 15 hours, under warm climate conditions of 26-32°C temperature, typical of this late-season cultivar showing progressive rate reduction
Division_rate_reduction represents the progressive mitotic frequency decline phase during declining division activity when cell division rate decreases from peak maximum 40,000-120,000 cells per day to declining 10,000-40,000 cells per day, mitotic index falls from 8-15% to 3-8%, cell cycle times extend reducing division frequency, and overall proliferative output diminishes marking transition from exponential to declining growth, proceeding over weeks 3-4.5 as primary rate reduction period. This deceleration phase occurs during mid-late summer in mid-late July in temperate regions Year N+1, spanning approximately 1-2 weeks representing major rate decline, corresponding to BBCH 73-74 pea-sized berry stages. Rate reduction mechanism involves progressive decrease in cycling cell number as cells exit cycle, cell cycle time extension reducing division frequency in remaining cycling cells, combined effects producing substantial rate reduction, auxin decline weakening proliferative drive, and developmental programming directing transition from division to expansion. Division rate quantification shows peak rate 40,000-120,000 new cells per berry per day weeks 2-3, declining rate 20,000-60,000 cells per day week 3.5, further reduction to 10,000-40,000 cells per day week 4.5, and rate decline reflecting both reduced cycling fraction and slower cycles. Rate decline dynamics show approximately 50-70% rate reduction from peak to early decline week 3.5, additional 40-60% reduction from early to late decline week 4.5, and cumulative 70-85% total rate reduction during decline phase. Cycling cell fraction reduction shows proportion of actively dividing cells decreasing from 70-85% at peak to 40-60% early decline to 20-35% late decline, declining fraction as cells progressively exit cycle, and fraction reduction as major rate reduction contributor. Cell cycle exit kinetics show progressive asynchronous exit across cell populations, earlier-activating cells exiting first during week 3, later-activating cells continuing longer into week 4-5, and exit wave creating gradual rate decline rather than abrupt cessation. Mitotic index decline shows percentage of cells in M phase decreasing from peak 8-15% to 5-10% early decline week 3.5 to 3-8% late decline week 4.5, mitotic index directly reflecting division activity level, and index decline documenting rate reduction. Cell cycle time extension contribution shows cycles lengthening from 10-12 hours peak to 12-14 hours early decline to 14-16 hours late decline, extended times reducing division frequency even in continuously cycling cells, and cycle extension accounting for 30-50% rate reduction with remainder from cycling fraction decline. G1 phase extension effects show prolonged G1 from 3-4 hours to 5-7 hours reducing cycle frequency, extended G1 providing window for exit decision, and G1 lengthening as control point for rate reduction. S and G2 phase extension shows DNA replication and preparation phases lengthening contributing to overall cycle extension, extended phases reflecting reduced biosynthetic capacity or checkpoint stringency, and phase lengthening reducing cycle frequency. Cell number accumulation slowing shows rate of new cell addition declining correlating with division rate reduction, accumulation curve slope decreasing during decline phase, and slowing accumulation approaching plateau representing final cell number. Exponential to linear transition shows growth pattern shifting from exponential doubling during peak to linear addition during decline to plateau at completion, transition characteristic of declining proliferative capacity, and pattern shift marking developmental progression. Auxin decline correlation shows hormone levels decreasing from 200-400 ng/g peak to 150-250 ng/g early decline to 80-150 ng/g late decline, declining auxin reducing proliferative stimulus, hormone decrease correlating temporally with rate reduction, and auxin as primary rate reduction trigger. Auxin-division relationship shows dose-dependent proliferative response, declining hormone weakening cell cycle progression, reduced auxin enabling cell cycle exit and expansion transition, and hormone control as key rate regulation mechanism. Developmental program regulation shows intrinsic genetic programming driving rate reduction beyond auxin effects, cultivar-specific decline kinetics affecting total division duration, programmed transition as fundamental developmental control, and endogenous regulation ensuring appropriate phase progression. Cell population heterogeneity effects show asynchronous division across cell populations creating gradual rather than synchronous rate decline, population variation extending decline phase duration, and heterogeneity contributing to smooth transition. Environmental effects on rate reduction include temperature affecting decline kinetics through developmental rate effects with warm conditions potentially accelerating rate reduction through faster progression, cool conditions potentially extending decline through slower developmental rate, and temperature as factor affecting transition timing though rate reduction occurs across temperature ranges. Rate reduction assessment methods include serial cell counting quantifying division rate from accumulation slopes, mitotic index measurement documenting dividing cell proportion, cell cycle time analysis showing extension, and mathematical modeling fitting decline kinetics. The process represents first aspect of declining division activity emphasizing rate decrease, preceding transition to enlargement onset and cell number finalization. Duration typically 1-2 weeks representing major rate reduction period weeks 3-4.5. Regional timing shows division rate reduction during mid-late July across climates as peak activity declines. Viticultural significance includes representing major transition from maximum proliferation to declining division, demonstrating progressive developmental regulation, establishing deceleration toward division completion, and marking shift toward expansion as growth mechanism. Research applications include analyzing rate reduction mechanisms and regulation, investigating auxin and developmental control of decline, evaluating cell cycle exit kinetics and control, quantifying rate decline dynamics and kinetics, and assessing rate reduction as transition to expansion-dominated growth.
Division_Rate_Reduction is first aspect of Declining_Division_Activity, distinguished by progressive mitotic frequency decline. Represents 70-85% total rate reduction from peak. Duration 1-2 weeks representing weeks 3-4.5. Rate declining from 40,000-120,000 to 10,000-40,000 cells/day. Mitotic index 8-15% declining to 3-8%. Results from both cycling fraction reduction and cycle time extension. Auxin decline primary trigger. Followed by Transition_to_Enlargement_Onset. Major developmental transition marker.
Proliferations-Dezeleration
Teilungsfrequenz-Reduktion
Teilungsraten-Reduktion
declining mitotic activity
division frequency reduction
mitotic rate decline
mitotische Ratenabnahme
proliferation deceleration
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 437-439 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Division Rate Reduction
The DNA fingerprint for Pinot Noir clone 115 (ENTAV selection, France) integrates three layers of molecular identity — the standard 9-locus SSR fingerprint (shared with all Pinot Noir clones including Pinot Gris and Pinot Blanc, confirming cultivar identity), an extended 20-locus SSR profile (European Vitis Database panel, also shared among Pinot color variants), and a targeted SNP panel at 6 functional loci including VvMYBA1 (heterozygous G/T, functional allele present, confirming red-berry phenotype — distinguishing Pinot Noir from the VvMYBA1-inactive Pinot Blanc and the chimeric Pinot Gris), VvDXS (wild-type, non-muscat), and 4 clone-diagnostic SNPs identified through whole-genome comparison of ENTAV clones (Vondras et al. 2019) that distinguish clone 115 from clones 114, 667, 777, and 828 at specific somatic mutation positions — the multi-layer DNA fingerprint provides cultivar confirmation (SSR layer), color variant assignment (VvMYBA1 layer), and clone-level identification (somatic SNP layer) in a single integrated molecular identity document.
A genetic marker data record that provides the comprehensive molecular identity profile of a specific Vitis vinifera accession integrating data from multiple marker systems — SSR genotype, SNP genotype, and optionally functional gene variants — into a unified identity document that enables unambiguous cultivar identification, clonal discrimination where possible, parentage verification, and traceability through the grapevine propagation and production chain.
DNA_Fingerprint represents the most comprehensive molecular identity record in the VVO, integrating information from multiple marker classes into a unified profile that addresses the full spectrum of identification questions from species through cultivar to clone level. The concept extends beyond the SSR_Fingerprint (which provides cultivar-level identification using the standard 9-locus panel) by incorporating additional marker data that address the resolution limitations of SSR-only identification: most importantly, SSR fingerprints cannot distinguish clones within a cultivar because the somatic mutations that differentiate clones rarely occur at SSR loci — resolving clonal identity requires either extended SSR panels (50-100 loci with a small probability of capturing clone-diagnostic mutations) or SNP-based approaches (targeted panels at known somatic variant positions, or whole-genome resequencing to discover clone-specific mutations). The integration of functional gene variants (VvMYBA1 for berry color, VvDXS for muscat aroma, disease resistance gene markers) adds phenotype prediction capability to the identity function, enabling molecular confirmation that an accession not only belongs to the correct cultivar but carries the expected functional alleles for its designated clone or biotype.
DNA-Fingerabdruck
comprehensive DNA identity
molecular identity profile
molekulares Identitätsprofil
multi-marker fingerprint
umfassender genetischer Fingerabdruck
Vondras AM et al. (2019) The genomic diversification of grapevine clones, BMC Genomics 20: 972 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340 This P et al. (2004) Development of a standard set of microsatellite reference alleles, Theoretical and Applied Genetics 109: 1448-1458 Kobayashi S et al. (2004) Retrotransposon-induced mutations in grape skin color, Science 304: 982
DNA Fingerprint
Molecular assessment of DNA_Repair_Disposition at INRAE Montpellier quantified CPD accumulation and repair kinetics across 12 cultivars following acute UV-B exposure (15 kJ m⁻² over 2 hours) and subsequent recovery under photoreactivating light (white light) or dark conditions. CPD levels were measured by immunoassay at 0, 1, 2, 4, 8, and 24 hours post-UV, enabling calculation of repair rates. Photolyase gene expression (VvPHR1, VvUVR3) and protein levels were assessed in parallel. DNA_Repair_Disposition showed striking cultivar variation. 'Pinot Noir' exhibited exceptional repair capacity with CPD levels declining 78% within 4 hours under photoreactivating light (half-life 1.8 hours), associated with high constitutive VvPHR1 expression (3× mean across cultivars) and rapid UV-induced upregulation (4.5-fold within 2 hours). Dark repair in 'Pinot Noir' was also efficient, removing 45% of CPDs within 8 hours through NER pathway. 'Malbec' showed good repair (68% removal at 4 hours) complementing its strong screening capacity. 'Chasselas' displayed poor DNA_Repair_Disposition with only 35% CPD removal at 4 hours under photoreactivating light (half-life 4.8 hours) and minimal dark repair (18% at 8 hours), associated with low VvPHR1 expression and weak induction. Functional consequences were confirmed by measuring cell survival after UV exposure—'Pinot Noir' leaf discs showed 85% cell viability versus 52% for 'Chasselas' at 24 hours post-UV. Interestingly, repair capacity showed weak correlation with flavonoid accumulation (r = 0.32), indicating these represent independent components of UV tolerance and explaining why 'Pinot Noir' tolerates UV well despite only moderate screening capacity.
A DNA_Repair_Disposition is a UV_Tolerance_Disposition that inheres in Vitis vinifera cells and is realized through efficient enzymatic mechanisms that detect and repair UV-induced DNA damage, particularly cyclobutane pyrimidine dimers (CPDs) and 6-4 photoproducts that if unrepaired cause mutations, replication blocks, and cell death. This disposition represents the damage repair component of UV tolerance, essential for maintaining genome integrity when UV screening is insufficient to prevent all DNA damage. Plants possess multiple DNA repair pathways including photoreactivation (light-dependent direct repair by photolyases, the most efficient pathway for UV damage), nucleotide excision repair (NER, a dark repair pathway removing and replacing damaged segments), and base excision repair (for oxidative damage). Photoreactivation is particularly important for UV tolerance, with CPD photolyase and 6-4 photolyase using blue/UV-A light energy to directly reverse the respective lesion types, enabling rapid and error-free repair. DNA_Repair_Disposition varies among genotypes in constitutive repair enzyme levels, UV-inducibility of repair pathways, and the efficiency of damage detection and processing. Strong repair capacity enables plants to tolerate UV exposure levels that would cause cumulative genetic damage in repair-deficient genotypes, complementing screening and antioxidant protection to provide comprehensive UV defense.
DNA_Repair_Disposition is_a UV_Tolerance_Disposition. Photoreactivation (photolyase-mediated) is the most efficient repair pathway for UV-induced CPDs and 6-4 photoproducts. Requires blue/UV-A light for photolyase activation. NER provides dark repair backup but is slower. Repair capacity is genetically variable and partially independent of screening capacity. VvPHR1 and VvUVR3 encode key photolyases in grapevine. Strong repair capacity complements screening in providing comprehensive UV protection.
DNA-Reparatur-Disposition
Photolyase-Aktivität
UV damage repair
UV-Schadens-Reparatur
nucleotide excision repair capacity
photolyase activity
Britt AB (1996) DNA damage and repair in plants, Annual Review of Plant Physiology and Plant Molecular Biology 47:75-100 Kimura S, Sakaguchi K (2006) DNA repair in plants, Chemical Reviews 106:753-766 Waterworth WM et al. (2011) Repairing breaks in the plant genome: the importance of keeping it together, New Phytologist 192:805-822
DNA Repair Disposition
The DNA sequence database for Vitis vinifera encompasses multiple scales of resolution — from short tandem repeat (STR/SSR) sequences of 2-6 bp motifs repeated 5-30 times at specific genomic loci (e.g., the dinucleotide repeat (CT)n at the VVS2 locus on chromosome 11 used for cultivar identification), through gene-length sequences of 1-10 kb encoding individual proteins (e.g., the VvMYBA1 transcription factor gene on chromosome 2 controlling anthocyanin biosynthesis, whose inactivation by Gret1 retrotransposon insertion produces the white-berry phenotype in cultivars like Chardonnay and Sauvignon Blanc), to chromosome-scale and whole-genome sequences of 475-500 Mb that capture the complete nuclear genetic complement including intergenic regions, transposable elements (comprising approximately 40% of the genome), and structural variants.
A genetic information entity that records the nucleotide sequence of a specific DNA region from a Vitis vinifera genotype, specifying the ordered succession of adenine, guanine, cytosine, and thymine bases that encodes the genetic instructions for protein synthesis, gene regulation, and genome structure, with the sequence data serving as the fundamental molecular identifier for genotype authentication, phylogenetic analysis, and functional genomics research.
DNA_Sequence is the foundational molecular data class from which all other genetic information types derive. In Vitis vinifera genomics, DNA sequence data serves three primary functions — genotype identification (using polymorphic marker sequences such as SSRs and SNPs to create unique molecular fingerprints for cultivar authentication), evolutionary and phylogenetic reconstruction (using sequence variation to trace domestication history, parentage relationships, and geographic dispersal patterns), and functional genomics (linking specific sequence variants to phenotypic traits through QTL mapping, GWAS, and candidate gene analysis): the grapevine genome is characterized by high heterozygosity (approximately 11.2% of positions polymorphic in the reference Pinot Noir genome), extensive structural variation between cultivars (large-scale inversions, insertions, deletions, and copy number variants), and a paleohexaploid origin detectable in the organization of paralogous gene clusters across the 19 chromosomes. Subclasses distinguish the three compartmental genomes — nuclear, chloroplast, and mitochondrial — that together constitute the complete DNA complement of the grapevine cell.
DNA sequence data
DNA-Sequenz
Nukleotidsequenz
genomic sequence record
genomische Sequenzdaten
nucleotide sequence
Jaillon O et al. (2007) The grapevine genome sequence suggests ancestral hexaploidization in major angiosperm phyla, Nature 449: 463-467 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Liang Z et al. (2019) Whole-genome resequencing of 472 Vitis accessions for grapevine diversity and demographic history analyses, Nature Communications 10: 1190
DNA Sequence
The DO designation Ribera del Duero (established 1982, Consejo Regulador headquartered in Roa de Duero) delimits approximately 23,000 hectares of vineyards along the Duero river valley in Castilla y León at 750-900 meters elevation, with production regulations specifying Tempranillo (locally Tinto Fino or Tinta del País) as the dominant variety (minimum 75% in blends), maximum yields of 7,000 kg/ha, and mandatory aging classifications — Joven, Crianza (12 months oak minimum), Reserva (36 months total, 12 in oak), and Gran Reserva (60 months total, 24 in oak) — each Consejo Regulador back label bearing a unique control number verifying regulatory compliance.
A regional wine designation within the Spanish wine classification system that identifies a wine as a Denominación de Origen — a wine originating from a registered and delimited production zone with established quality characteristics, produced according to the regulations of the controlling Consejo Regulador, formally classified as a Protected Designation of Origin (DOP) under EU harmonization, representing the standard quality appellation tier between Vino de la Tierra (PGI equivalent) and the higher DOCa/VP tiers.
DO_Designation documents Spain's principal appellation tier, with approximately 70 registered DOs covering the major wine-producing regions. The Spanish DO system was established by the Estatuto del Vino of 1932 (making it one of the earliest formal appellation systems after France's AOC) and is administered by region-specific Consejos Reguladores that function as both regulatory authorities and collective marketing organizations: each Consejo Regulador maintains a register of authorized producers, vineyards, and bodegas, conducts analytical and sensory quality examinations, and issues the numbered back labels (contraetiquetas) that guarantee compliance with the DO production specification. The 2003 Spanish wine law (Ley 24/2003) introduced a hierarchical structure with Vino de Mesa, Vino de la Tierra (now IGP), DO (now DOP), DOCa (Denominación de Origen Calificada), and Vino de Pago (single-estate DO) tiers, subsequently harmonized with the EU PDO/PGI framework.
DO-Bezeichnung
Denominación de Origen
Denominación-de-Origen-Klassifikation
Spanish appellation designation
spanische Herkunftsbezeichnung
Spanish Wine Law (Ley 24/2003 de la Viña y del Vino) EU Regulation 1308/2013 Art. 93 Ministerio de Agricultura, Pesca y Alimentación (2023) Denominaciones de Origen Protegidas
DO Designation
The DOC designation Soave DOC (established 1968) delimits approximately 6,500 hectares of vineyards in the Veneto east of Verona, with the disciplinare specifying Garganega as the principal variety (minimum 70%), maximum yield of 14,000 kg/ha (reduced to 10,000 kg/ha for Soave Classico from the historical hillside zone), minimum total alcohol of 10.5%, and mandatory analytical examination — the DOC system operates through Consorzi di Tutela (protection consortia) that administer the disciplinare, conduct quality examinations, and manage collective brand promotion, with each DOC wine carrying a fascetta (government-issued neck band with serial number) certifying regulatory compliance.
A regional wine designation within the Italian wine classification system that identifies a wine as a Denominazione di Origine Controllata — a wine produced within a registered and delimited area according to a binding production specification (disciplinare di produzione) that defines authorized grape varieties, viticultural practices, yields, winemaking methods, aging requirements, and analytical parameters, formally classified as a Protected Designation of Origin (DOP) under EU harmonization.
DOC_Designation documents Italy's standard appellation tier, established by Presidential Decree 930/1963 and representing the foundation of the Italian quality wine classification. Italy has approximately 330 registered DOC designations — the largest number of any country — reflecting both the extraordinary diversity of Italian viticultural territory and the politically decentralized nature of Italian appellation creation: each DOC has its own disciplinare drafted by local producers and approved by the national Ministry of Agriculture, resulting in considerable variation in regulatory stringency between DOCs. The DOC tier sits between IGT (PGI) and DOCG (the higher quality tier requiring additional examination and bottle numbering) in the Italian quality pyramid, with the 2010 EU harmonization formally classifying both DOC and DOCG as DOP while permitting continued use of the national terminology.
DOC
DOC-Bezeichnung
Denominazione di Origine Controllata
Denominazione-di-Origine-Controllata-Klassifikation
Italian controlled origin designation
italienische kontrollierte Herkunftsbezeichnung
Italian Presidential Decree 930/1963 Italian Law 164/1992 on wine designations EU Regulation 1308/2013 Art. 93 Ministero delle Politiche Agricole Alimentari e Forestali (2023) Elenco DOC registrate
DOC Designation
The DOCG designation Barolo DOCG (elevated from DOC to DOCG in 1980) imposes the strictest production requirements in the Italian system for Nebbiolo wines from the Langhe — maximum yield of 8,000 kg/ha, minimum 38 months aging (18 in oak) for standard Barolo and 62 months (18 in oak) for Riserva, minimum 13% alcohol, mandatory chemical analysis and tasting commission approval, and government-issued numbered fascette affixed to each bottle that enable traceability from retail shelf to individual production lot, with the Consorzio di Tutela Barolo Barbaresco Alba Langhe e Dogliani administering quality control and promotional activities for approximately 11 million bottles annually.
A regional wine designation within the Italian wine classification system that identifies a wine as a Denominazione di Origine Controllata e Garantita — the highest tier of Italian wine classification, reserved for wines of particular prestige and established quality reputation that have been DOC for a minimum period (typically 5-10 years) and are subject to additional quality assurance measures including mandatory government-issued numbered neck bands (fascette) and more rigorous analytical and sensory examination, formally classified as a Protected Designation of Origin (DOP) under EU harmonization.
DOCG_Designation represents the apex of the Italian quality pyramid, with the Garantita (guaranteed) suffix signifying a higher level of state assurance than standard DOC. The DOCG tier was introduced in 1963 but the first designations were not awarded until 1980 (Barolo, Barbaresco, Brunello di Montalcino, Vino Nobile di Montepulciano, and Chianti): Italy now has approximately 77 DOCG designations, with the category reserved for wines that have demonstrated consistent quality over an extended period at DOC level. The distinctive feature of DOCG beyond stricter production parameters is the requirement for government-issued numbered fascette — serialized paper or foil bands applied to each bottle neck that create a chain of custody from production to point of sale, functioning as both an anti-fraud measure and a production volume control mechanism. The fascette system processes approximately 800 million individual bands annually across all DOCG wines.
DOCG
DOCG-Bezeichnung
Denominazione di Origine Controllata e Garantita
Denominazione-di-Origine-Controllata-e-Garantita-Klassifikation
Italian guaranteed origin designation
italienische garantierte Herkunftsbezeichnung
Italian Presidential Decree 930/1963 Italian Law 164/1992 EU Regulation 1308/2013 Art. 93 Consorzio di Tutela Barolo Barbaresco Alba Langhe e Dogliani (2023) Production regulations
DOCG Designation
Documentation source inventory for Furmint integrates three distinct evidentiary streams: historical document references (the 1571 Rákóczi estate inventory, the 1623 Latin document using the name "Furmint," the 1730 Tokaj classification decree — all housed in the Magyar Nemzeti Levéltár, Budapest), ampelographic sources (Trummer 1841 providing the first systematic morphological description, Galet 1962 updating the ampelographic characterization with modern terminology, VIVC database entry maintaining the current standard description), and scientific publication references (Bowers et al. 1999 establishing microsatellite fingerprint, Cipriani et al. 2010 confirming parentage relationships, Duchêne et al. 2009 characterizing aromatic profile genetics) — the three source categories collectively providing a 450-year evidentiary chain from first documentation to contemporary molecular characterization. Documentation source inventory for Grüner Veltliner relies more heavily on recent scientific literature than on historical documents, reflecting the cultivar's relatively shallow archival record — limited historical references (Burger 1837, Trummer 1841), key ampelographic sources (Austrian Bundesamt für Weinbau Klosterneuburg catalogues), and a substantial modern scientific corpus (Sefc et al. 1998 establishing the Traminer × St. Georgen parentage, subsequent Austrian and international research on clonal diversity, aromatic chemistry, and oenological potential).
Documentation source is a historical record entity that documents the bibliographic, archival, and scholarly references from which historical information about grape cultivars, viticultural practices, and wine production traditions has been derived, encompassing historical document references (primary archival and literary sources), ampelographic sources (systematic cultivar description treatises), and scientific publication references (peer-reviewed research contributing to cultivar knowledge), providing the evidentiary provenance chain that establishes the reliability, dating, and interpretive context of historical claims and enables the scholarly assessment of evidence quality that distinguishes well-documented historical knowledge from tradition, conjecture, and myth.
Documentation source as an ontological category serves a critical epistemic function within VVO — it makes explicit the evidentiary basis for historical claims about cultivars, enabling users to assess the reliability and provenance of information rather than accepting historical narratives as unquestioned fact. The quality and completeness of documentation sources varies enormously across cultivars — major international varieties (Cabernet Sauvignon, Pinot Noir, Riesling) are supported by extensive, multi-century archival records, comprehensive ampelographic descriptions from multiple traditions, and hundreds of peer-reviewed scientific publications, while minor indigenous cultivars may have only a single ampelographic catalogue entry and a handful of recent molecular studies. The tripartite classification of documentation sources into historical documents (primary archival sources), ampelographic sources (systematic cultivar description tradition), and scientific publications (peer-reviewed research) reflects three fundamentally different epistemological traditions — historical scholarship operates through source criticism and contextual interpretation, ampelography through systematic morphological observation and comparison, and modern science through hypothesis testing, experimental validation, and peer review — each with its own standards of evidence, methods of verification, and characteristic limitations. The integration of these three evidentiary streams is essential for robust cultivar knowledge — historical documents provide temporal depth but are subject to misidentification and interpretive ambiguity, ampelographic descriptions provide morphological precision but are limited by phenotypic plasticity and observer subjectivity, and scientific publications provide molecular certainty but typically lack historical depth. The VVO approach of documenting sources explicitly within the ontology structure (through definition_source fields in every class annotation) embodies the principle that knowledge claims should be traceable to their evidentiary foundations.
Belegquelle
Dokumentationsquelle
Quellendokumentation
Quellennachweis
Referenzdokumentation
bibliographic source
evidentiary source
reference documentation
scholarly reference
source documentation
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes: A Complete Guide to 1,368 Vine Varieties, Allen Lane, pp.: xv-xxxii This P et al. (2006) Historical origins and genetic diversity of wine grapes. Trends in Genetics 22(9): 511-519 VIVC (2024) Vitis International Variety Catalogue, Julius Kühn-Institut, Bundesforschungsinstitut für Kulturpflanzen, https://www.vivc.de
Documentation Source
The documented parentage of Pinotage records the deliberate cross performed by Abraham Izak Perold at the University of Stellenbosch in 1925 — Pinot Noir (seed parent) × Cinsaut (pollen parent, then known as Hermitage in South Africa, hence the portmanteau name Pino-tage) — documented in Perold's laboratory notebooks preserved in the Stellenbosch University archives, with the four original seedlings planted in his garden and subsequently transferred to the Elsenburg Agricultural College by CJ Theron in 1927 when Perold relocated. This documented parentage was confirmed by SSR analysis (Sefc et al. 1997) matching Pinotage's alleles to Pinot Noir and Cinsaut at all 9 OIV standard loci, but the documented record remains the primary parentage evidence because it provides information (crossing date, breeder identity, seed versus pollen parent assignment) that molecular analysis alone cannot determine.
A parentage information record based on contemporaneous written documentation of a deliberate hybridization event or controlled seedling selection, specifying the breeder identity, crossing date, recorded maternal seed parent and paternal pollen parent, breeding program context, and the archival source of the crossing record that provides direct historical evidence of a cultivar's biological origins independent of molecular confirmation.
Documented_Parentage captures parentage claims supported by primary historical evidence — breeding records, laboratory notebooks, institutional reports, or published accounts contemporaneous with the hybridization event. This evidence class applies primarily to cultivars produced in formal breeding programs (19th century onward) and to a small number of older cultivars whose origin from specific natural crosses was observed and recorded: examples include the deliberate crosses of the 19th-20th century breeding era (Müller-Thurgau 1882, Pinotage 1925, Seyval Blanc 1919, Chambourcin 1963) and the few historical cultivars with documented seedling selection (e.g., Cabernet Franc as a documented parent in Bordeaux nursery records). The reliability of documented parentage varies — some breeding records are meticulously maintained with pollen isolation, emasculation, and controlled pollination protocols ensuring parentage accuracy, while others (particularly 19th century records) may contain errors from inadequate pollen control, nursery mislabeling, or mistaken cultivar identification of parent material. Molecular confirmation of documented parentage is therefore standard practice, with several notable cases of documented parentage being contradicted by molecular evidence (Müller-Thurgau's Silvaner parent corrected to Madeleine Royale, Trebbiano Toscano excluded as a parent of Ugni Blanc despite historical claims).
Kreuzungsprotokoll-Abstammung
breeding record parentage
dokumentierte Abstammung
historical crossing documentation
historisch belegte Elternschaft
recorded parentage
Burger P et al. (2017) The heritage of two legendary grape cultivars, Pinotage and Kerner, American Journal of Enology and Viticulture 68: 176-183 Dettweiler E et al. (2000) Grapevine cultivar Müller-Thurgau and its true-to-type descent, Vitis 39: 63-65 Lacombe T et al. (2013) Large-scale parentage analysis in an extended set of grapevine cultivars, Theoretical and Applied Genetics 126: 401-414
Documented Parentage
A dominant apical shoot originating from a lateral bud at node 8 on a young trunk of Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, California, that emerged vigorously following hail damage to the original terminal bud in May, growing 140 cm vertically by August with 16 nodes and strong lateral bud suppression, achieving growth rates of 4-5 cm per day and leaf areas averaging 135 cm², effectively replacing the damaged terminal shoot and continuing trunk extension toward the target height of 180 cm, while the original trunk section above node 8 showed only weak lateral shoot development due to the dominant apical shoot's strong auxin production.
An apical shoot that originates from a lateral bud position rather than from the intended terminal bud of a woody structure, but has assumed apical dominance and terminal growth characteristics due to suppression, damage, or removal of the original main apical shoot in Vitis vinifera, characterized by vigorous upward growth, strong lateral bud suppression along its length, and functional replacement of the primary growth axis despite its lateral developmental origin. Dominant apical shoots represent significant architectural plasticity in grapevine growth, allowing compensation for damage to primary meristems and enabling training system adaptation when original framework plans require modification due to environmental pressures, disease damage, or management errors. These shoots typically emerge when apical dominance from the main shoot is released either through physical removal of the terminal meristem or through physiological signals indicating primary shoot dysfunction, and can achieve vigor levels comparable to or exceeding original main apical shoots depending on timing of release and resource availability. Dominant apical shoots may either be intentionally encouraged to create alternative framework structures or selectively removed to prevent competition with desired main shoots, representing a critical decision point in canopy architecture management.
Dominant apical shoots are distinguished from main apical shoots by their lateral rather than terminal developmental origin, and from ordinary lateral shoots by their assumption of apical dominance and terminal growth characteristics. The transition from lateral to dominant apical identity typically occurs rapidly following release from apical dominance suppression, with hormonal signaling shifting within 48-72 hours and morphological changes becoming apparent within one week. In training system management, decisions about whether to maintain or remove dominant apical shoots depend on timing (early season emergence more valuable), position (proximity to desired framework location), and vigor relative to remaining main shoots. Multiple dominant apical shoots creating competing growth axes require selection to re-establish architectural clarity.
Ersatztrieb mit apikaler Dominanz
dominanter Seitentrieb
emergent dominant shoot
lateral-derived terminal shoot
lateral-dominanter Trieb
lateral-origin apical shoot
substitute apical shoot
Based on Cline (1997) "Concepts and terminology of apical dominance" American Journal of Botany 84=1064-1069 Grose & Olmstead (1991) "Measurement of grapevine growth and fruit development by ocular estimation" American Journal of Enology and Viticulture 42=164-170 Bennett et al. (2005) "Morphological and environmental effects on compound bud development in Vitis vinifera L." Australian Journal of Grape and Wine Research 11=37-46 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 167-189.
Dominant Apical Shoot
A dominant main axis inflorescence at node 3 on a shoot of Vitis vinifera cv. Grenache in Châteauneuf-du-Pape, France, measuring 19.5 cm total length with rachis extending the full 19.5 cm representing complete dominant axis, rachis diameter of 5.2 mm at base compared to largest lateral branch diameter of 1.8 mm creating diameter ratio of 2.9=1 indicating substantial rachis dominance, rachis showing robust construction with thick-walled tissues and prominent vascular bundles, bearing 14 lateral branches distributed along its length with branches showing clear size subordination (largest laterals 4.5-5.0 cm, smallest 1.5-2.0 cm) all substantially smaller than the 19.5 cm rachis, lateral branches showing relatively simple secondary and tertiary branching contrasting with the massive primary rachis, rachis containing approximately 75% of total inflorescence vascular tissue providing dominant resource delivery capacity, architectural organization showing clear hierarchy with unequivocal rachis dominance and lateral branches as clearly subordinate appendages, robust rachis providing essentially all mechanical support bearing the complete inflorescence weight and wind loading, measured at BBCH stage 63 (30% flowers open) in early June, representing maximum dominant main axis architecture with pronounced rachis structural and functional superiority.
A main axis primary inflorescence emphasizing the structural and functional dominance of the central rachis over lateral branch systems in Vitis vinifera, characterized by rachis substantially larger and more robust than lateral branches with typical diameter ratios of 2=1 to 4=1 (rachis=largest lateral), rachis length representing 80-100% of total inflorescence length indicating full-length extension dominating the structure, lateral branches clearly subordinate showing progressive size reduction and simpler architecture compared to the massive central axis, rachis bearing the majority of vascular tissue and providing essentially all mechanical support, clear hierarchical dominance with rachis as the unequivocal primary structure and all laterals as secondary subordinate elements, and functional significance with the rachis serving as the dominant resource delivery pathway and structural framework upon which the lateral flowering system depends. Dominant main axis inflorescences represent maximum expression of rachis dominance, occurring in cultivars and conditions producing robust central axes with relatively reduced lateral elaboration, exhibiting clear architectural hierarchy facilitating efficient resource transport through the dominant central vascular system, and creating the structural framework supporting the complete flowering and subsequent fruiting structure.
Dominant main axis inflorescences emphasize the degree of rachis superiority over lateral branches, distinguished from simply having a central axis by the pronounced dominance relationship. The dominant designation highlights the substantial size, structural, and functional advantages of the rachis creating clear hierarchical relationships rather than just organizational centrality. Diameter ratios quantify dominance with >2=1 rachis=lateral ratios indicating substantial dominance, >3=1 indicating pronounced dominance, and approaching 4=1 representing extreme dominance. Full-length rachis extension representing essentially complete inflorescence length indicates maximum rachis development with laterals serving as appendages rather than co-equal branches. Vascular dominance with rachis containing majority of conducting tissue creates the dominant resource delivery pathway with laterals dependent on rachis supply. Mechanical dominance with robust rachis providing essentially complete structural support creates dependency relationships with laterals unable to support their own weight without rachis framework. Developmental priority establishes rachis dominance with apical meristem persistence creating continuous elongation while lateral branch development remains subordinate. Occurrence shows cultivar variation with some varieties consistently producing dominant rachis architecture while others show more balanced development, and environmental effects with favorable conditions supporting robust rachis construction while stress may limit dominance development. Transformation to cluster maintains dominance relationships with rachis becoming the dominant cluster stem providing primary support and vascular function.
Rachis-hierarchischer Blütenstand
dominant central axis inflorescence
dominante Hauptachse-Infloreszenz
rachis-dominant hierarchy inflorescence
subordinate lateral inflorescence
untergeordnete Lateral-Infloreszenz
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 213-248 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 249-285 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 429-468.
Dominant Main Axis Inflorescence
Merlot grafted on Teleki 5C rootstock at 14 years showing dominant main root axis 47% greater diameter than next-largest primary root (14.2 cm versus 9.7 cm at crown), dominant axis extending 178 cm depth while other primary roots maximum 125 cm depth, dominant axis producing lateral roots at 8-12 cm intervals with extensive branching while other roots showing sparser laterals at 15-25 cm intervals, hydraulic conductivity measurements indicating dominant axis conducting estimated 58% of total root system water transport, dominant axis biomass estimated 840 grams dry weight comprising 52% of total primary root mass, dominant axis demonstrating clear superiority in size, depth, branching, and function over other primary roots in system
Dominant main root axis is main root axis showing greatest size, extent, or functional importance in root system, characterized by largest diameter typically exceeding other primary roots by 20-50%, deepest penetration or greatest lateral extent exploring more soil volume than other roots, highest hydraulic conductivity potentially contributing 30-60% of total root system water transport, and most extensive lateral branching producing largest portion of absorptive root network, with dominance potentially reflecting developmental timing as earliest-formed root, favorable growth direction encountering permeable soil and optimal resources, or genetic determination in seedlings where radicle-derived tap root naturally dominant, and demonstrating that while grapevine root systems typically show multiple primary roots especially in propagated vines one root often establishes dominance through size, position, or function providing disproportionate contribution to vine support and resource supply. Dominant main root axis size superiority includes= showing largest diameter (dominant axis typically 20-50% greater diameter than next-largest root reflecting superior growth), exhibiting greatest length (dominant axis often extending deeper or farther laterally than other primary roots exploring more soil volume), demonstrating highest biomass (dominant axis containing more tissue mass than other individual primary roots potentially 30-60% of total primary root biomass), displaying most extensive branching (dominant axis producing more lateral roots than other primary roots creating largest branching network), and revealing disproportionate contribution (dominant axis providing greater contribution to total root system function than its numerical representation as single root among several). Dominant main root axis developmental origin includes= often earliest-formed root (dominant axis frequently being first primary root to develop from seedling radicle or cutting callus gaining temporal advantage), showing prolonged growth (dominant axis potentially maintaining growth longer each season than competing primary roots accumulating more tissue), receiving priority resources (dominant axis possibly receiving preferential allocation of photosynthates and resources from vine enhancing growth), encountering favorable conditions (dominant axis potentially growing in direction with most permeable soil and adequate moisture and nutrients enabling superior development), and demonstrating positive feedback (dominant axis size advantage potentially reinforcing through enhanced water and nutrient transport supporting continued superior growth). Dominant main root axis in seedling vines includes= typically being radicle-derived tap root (seedling dominant axis naturally being original radicle root growing first and maintaining growth advantage), showing single clear dominant root (seedling usually exhibiting one obviously dominant central tap root exceeding all other roots), maintaining vertical orientation (seedling dominant axis typically growing straight downward maximizing depth and stability), demonstrating natural dominance (seedling root system naturally organized with single dominant central axis without competition from multiple equal primary roots), and persisting throughout life (seedling dominant tap root maintaining superiority from germination through mature and old vine). Dominant main root axis in propagated vines includes= potentially less distinct (cutting producing multiple adventitious primary roots with dominance less pronounced than seedling single radicle), developing through competition (cutting primary roots initially similar size with one eventually establishing dominance through superior growth), showing variable which root (any of several cutting primary roots potentially becoming dominant depending on growth conditions and direction), demonstrating functional dominance (cutting dominant axis defined by size and function rather than developmental origin unlike seedling radicle natural dominance), and possibly changing over time (cutting root system dominant axis potentially shifting if initially dominant root damaged or encounters barrier allowing another root to surpass). Dominant main root axis functional importance includes= providing majority water transport (dominant axis potentially conducting 30-60% of total root system water supply due to large xylem cross-section), supplying greatest nutrient uptake (dominant axis lateral network providing largest soil exploration and absorption surface area), contributing most anchorage (dominant axis deep penetration or extensive lateral spread providing strongest anchoring component), storing largest reserves (dominant axis substantial biomass enabling greatest carbohydrate storage capacity potentially 40-70% of total root storage), and serving as main structural support (dominant axis providing primary structural foundation for entire root system). Dominant main root axis spatial position includes= potentially showing favorable orientation (dominant axis possibly positioned in direction with best soil conditions enabling superior growth), demonstrating maximum extent (dominant axis exploring furthest into soil profile either vertically, laterally, or both exceeding other primary roots), occupying optimal soil zone (dominant axis possibly accessing most favorable soil layers with adequate moisture, nutrients, and oxygen supporting superior function), showing unrestricted growth path (dominant axis potentially avoiding obstacles or barriers that limited other primary roots), and exploiting soil heterogeneity (dominant axis possibly growing preferentially toward soil zones with optimal resources while avoiding less favorable areas). Dominant main root axis lateral branching includes= producing most lateral roots (dominant axis generating more branch roots than other primary roots creating largest lateral network), showing highest branching density (lateral roots per unit length potentially higher on dominant axis than other roots), exhibiting most extensive lateral systems (dominant axis laterals extending farther and branching more extensively exploring largest soil volume), demonstrating greatest fine root production (dominant axis lateral network producing most absorptive fine roots providing largest uptake surface area), and contributing disproportionately to function (dominant axis branching network potentially providing 40-70% of total root system absorption despite being single axis among several). Dominant main root axis vulnerability includes= representing single point failure risk (damage or disease affecting dominant axis potentially severely compromising entire root system function), showing disproportionate loss impact (losing dominant axis more detrimental than losing smaller non-dominant root due to greater functional contribution), demonstrating limited redundancy (vine heavily dependent on single dominant axis potentially vulnerable if that root damaged), requiring protection priority (dominant axis representing most critical root requiring careful management to avoid damage), and potentially compensated over time (loss of dominant axis possibly eventually compensated by growth of remaining roots though requiring extended recovery period).
Dominant_Main_Root_Axis emphasizes superior size and function and is distinguished from other main root axes by focus on dominance. Critical for understanding root system hierarchy, functional inequality, developmental advantage, resource contribution, vulnerability concentration, and the single primary root providing disproportionate contribution to total root system structure and function.
Dominante Hauptwurzelachse
Führende Wurzelachse
Leading root axis
Main structural axis
Primary dominant root
Primäre Leitwurzel
Dominant Main Root Axis
Muscadinia rotundifolia serves in a Donor_Parent_Role as the source of powdery mildew resistance (Run1 locus) in French resistance breeding, its resistance genes the introgression target while its non-vinifera fruit character is selected against in subsequent generations. Vitis amurensis bears a Donor_Parent_Role for cold hardiness in continental climate breeding, contributing winter survival genes from its native range in northeastern Asia. Bouquet (a complex hybrid carrying multiple R-genes) functions as donor parent in second-generation PIWI breeding, providing "pyramided" resistance loci for introgression into V. vinifera backgrounds.
A parent role that inheres in a grapevine genotype by virtue of its function as the source of specific traits targeted for introgression into breeding populations, typically providing alleles absent from the recurrent parent or elite V. vinifera background. Donor parents contribute the genetic basis for desired traits—disease resistance genes, stress tolerance mechanisms, or quality attributes—that are the objectives of the breeding program, with the donor's broader genome considered neutral or potentially undesirable genetic background to be minimized through selection. In resistance breeding, donor parents are typically wild Vitis species (V. rotundifolia, V. amurensis, V. cinerea) or their derivatives carrying resistance (R) genes absent from susceptible V. vinifera. The donor concept frames breeding as trait transfer rather than novel variety creation, with the donor providing specific genetic factors for incorporation while the recurrent parent provides the genetic foundation. Donor identification requires genetic analysis to characterize the trait's inheritance and identify markers enabling selection, particularly for traits like disease resistance where phenotypic selection may be unreliable or require pathogen exposure.
Donor_Parent_Role is subclassed by the nature of the donated trait—Resistance_Donor_Role for pest/disease resistance sources and Quality_Donor_Role for fruit or wine quality trait sources. The donor concept explicitly frames the parent's contribution as targeted trait transfer rather than comprehensive genetic contribution donor parents may have many undesirable characteristics (non-vinifera fruit, vigorous growth, unfamiliar wine character) that breeding seeks to eliminate through backcrossing while retaining the specific valuable trait(s). Donor identification and characterization is increasingly molecular—mapping resistance loci, developing linked markers, identifying candidate genes—enabling precise trait tracking through breeding populations. The distinction between donor and recurrent parents clarifies breeding strategy, with donor genetics minimized except at target trait loci while recurrent parent genetics dominate the background genome.
Genspender
Merkmalsquelle
Resistenzquelle
Spenderelter
donor parent
gene donor
resistance source
trait source
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Dry IB et al. (2010) Molecular strategies for disease resistance, Australian Journal of Grape and Wine Research 16: 94-105 Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330
Donor Parent Role
In Riesling vineyards of the Rheingau, Germany, the dormancy period extends from late November through mid-March, with vines entering endodormancy following exposure to temperatures below 10°C and short photoperiods of less than 12 hours, accumulating approximately 1200 chilling hours below 7.2°C for complete dormancy satisfaction, and remaining in ecodormancy until growing degree day accumulation triggers budbreak in early to mid-April.
Dormancy in Vitis vinifera constitutes a fundamental phenological period characterized by the temporary suspension of visible growth and metabolic activity in meristematic tissues, representing an essential adaptive mechanism that enables perennial grapevines to survive unfavorable environmental conditions, particularly the freezing temperatures and desiccating conditions of winter in temperate viticulture regions. This complex physiological state encompasses a continuum of growth inhibition phases ranging from environmentally-induced dormancy induction through deep endodormancy to the environmentally-constrained ecodormancy that persists until conditions favor growth resumption in spring. The dormancy period in grapevine typically extends from late autumn through early spring, spanning approximately 4-6 months in Northern Hemisphere viticulture, during which buds remain in a quiescent state despite containing fully formed primordial shoots with leaf, tendril, and inflorescence primordia that were initiated during the previous growing season. Physiologically, dormancy involves profound changes in cellular metabolism, including accumulation of abscisic acid as a key regulatory hormone, decreased auxin and cytokinin activity, altered carbohydrate partitioning with starch accumulation in woody tissues, increased expression of dehydrin proteins that protect cellular structures during freezing, and modifications to membrane lipid composition that enhance freeze tolerance. The dormancy cycle is regulated by the interacting influences of photoperiod, temperature, and endogenous developmental programs, with short day length serving as the primary environmental signal initiating dormancy induction in autumn, chilling temperature accumulation required for dormancy release, and warming temperatures triggering growth resumption once dormancy has been satisfied. Cultivar differences in dormancy characteristics significantly influence viticultural adaptation, with cultivars originating from continental climates typically exhibiting deeper dormancy and higher chilling requirements than those from Mediterranean regions, affecting both winter hardiness and the timing of budbreak in spring. The dormancy period serves critical functions beyond winter survival, including synchronization of spring growth with favorable environmental conditions, prevention of premature budbreak during temporary warm periods, and provision of the rest period necessary for proper development of reproductive structures. Understanding dormancy physiology has become increasingly important in the context of climate change, as warming winters may provide insufficient chilling for complete dormancy satisfaction in some regions, potentially leading to erratic budbreak, reduced fruitfulness, and compromised vine performance.
Dormancy represents the highest-level process class within the dormancy hierarchy under Grapevine_Phenological_Process, encompassing all phases of growth suspension from induction through release. Distinguished from active growth phases by the absence of visible meristematic activity despite the presence of viable growing points. Subdivides into Dormancy_Induction, Endodormancy, Ecodormancy, and Dormancy_Release as sequential phases. Duration typically spans 4-6 months in temperate viticulture. Critical for winter survival and proper spring growth synchronization.
Dormanz
Ruheperiode
Vegetationsruhe
Winterruhe
growth cessation period
seasonal dormancy
winter dormancy
winter rest
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Lang GA et al (1987) Endo-, para-, and ecodormancy: physiological terminology and classification for dormancy research. HortScience 22: 371-377
Dormancy
In Cabernet Sauvignon vines in the Médoc, Bordeaux, dormancy break occurs in early to mid-April when accumulated growing degree days reach the cultivar threshold of approximately 160-180 GDD following complete chilling satisfaction in February, with visible bud swelling appearing at BBCH stage 05 within 3-5 days of threshold achievement and progression to green tip emergence at BBCH stage 07 within 7-10 days, marking the irreversible transition to active growth.
Dormancy break in Vitis vinifera represents the terminal event of the dormancy cycle at which accumulated heat exposure triggers the irreversible transition from quiescent ecodormancy to active growth, initiating the visible developmental changes that mark the beginning of the new growing season. This transition occurs when heat accumulation has satisfied the cultivar-specific requirement following complete chilling satisfaction, representing the culmination of the dormancy release process in the actual initiation of growth. The physiological basis of dormancy break involves the crossing of developmental thresholds where accumulated metabolic activity, biosynthetic production, and cellular preparation reach levels sufficient to support sustained growth, with the system transitioning from preparatory activity to active developmental progression. The irreversibility of dormancy break distinguishes it from the reversible quiescence of earlier ecodormancy, with buds that have passed the break point continuing developmental progression even if temperatures subsequently decline, in contrast to pre-break buds that return to quiescence during cool periods. The visible manifestation of dormancy break is bud swelling, where the enclosed primordial shoot begins expansion that separates bud scales and produces the characteristic appearance of swollen buds transitioning to green tip emergence, corresponding to BBCH stages 05 through 07. The timing of dormancy break is determined by the achievement of heat accumulation thresholds following chilling completion, with the specific timing varying among cultivars, seasons, and sites according to the interaction of genetically determined requirements with environmental temperature patterns. The synchrony of dormancy break across buds on a vine affects the uniformity of subsequent shoot development, with simultaneous break producing uniform shoot populations while extended break periods create the shoot heterogeneity that complicates canopy management. The vulnerability following dormancy break relates to the cold sensitivity of actively growing tissues, with broken buds and emerging shoots being highly susceptible to frost damage that can significantly impact vine productivity, making accurate prediction of dormancy break timing essential for frost protection management.
Dormancy_Break represents the fourth and final component phase of Dormancy_Release marking the actual transition to active growth. Distinguished from earlier phases by representing the irreversible growth initiation rather than preparation. Subdivides into Active_Growth_Resumption, Bud_Swelling_Initiation, and Dormancy_Exit_Completion as component aspects. Visible as bud swelling at BBCH 05-07. Timing determined by heat threshold achievement following chilling. Creates frost vulnerability requiring protection management. Synchrony affects subsequent shoot uniformity.
Dormanzbeendigung
Dormanzbruch
Wachstumsbeginn
budbreak initiation
dormancy ending
dormancy termination
growth commencement
Dormancy Break
Dormancy breaking potential assessment comparing agents in partially chilled Sultana (50% of chilling requirement satisfied) reveals the effectiveness variation among treatments, with hydrogen cyanamide (4% application) showing high breaking potential (advancing budbreak by 18 days, increasing budbreak percentage from 65% to 92%, improving uniformity), potassium nitrate (5%) showing moderate potential (advancing by 8 days, percentage to 78%), mineral oil (4%) showing limited potential (advancing by 5 days, percentage to 72%), and untreated control showing the baseline response to incomplete chilling (delayed, erratic budbreak at 65%), demonstrating how breaking potential varies among agents with implications for selecting appropriate dormancy management interventions in mild-winter climates.
Dormancy breaking potential quality is a dormancy transition quality that represents the capacity of various agents or conditions to overcome dormancy and stimulate bud growth, characterizing the effectiveness of chemical, physical, or environmental treatments in releasing buds from dormancy that may be used when natural dormancy release is inadequate or when manipulation of budbreak timing is desired for management purposes. Within the BFO framework, this quality specifically addresses the intervention dimension of dormancy transitions, distinguishing the potential for induced release from natural dormancy progression while recognizing that dormancy breaking agents work through various mechanisms (hormonal manipulation, metabolic disruption, oxidative stress) to overcome the physiological barriers to growth that constitute dormancy. The ontological significance lies in dormancy breaking potential providing management options for situations where natural dormancy release is problematic, including warm-winter climates with insufficient chilling (where agents substitute for inadequate natural conditioning), situations requiring advanced or synchronized budbreak (where agents enable timing manipulation), and research applications requiring controlled bud growth initiation (where agents provide experimental control over dormancy status). The agents with dormancy breaking potential include hydrogen cyanamide (the most widely used commercial agent, effective through oxidative stress and respiratory disruption), mineral oils (enhancing gas exchange and potentially affecting hormone balance), potassium nitrate (providing nitrogen stimulus and osmotic effects), thiourea (affecting sulfur metabolism), and physical treatments such as hot water (disrupting dormancy through controlled thermal damage), with different agents showing different effectiveness depending on dormancy depth, cultivar, and application conditions. The effectiveness of dormancy breaking agents depends on the current dormancy status, with agents generally more effective on buds that have received partial chilling (primed for release) than on deeply endodormant buds (still requiring substantial conditioning), creating the practical guideline that agents are most useful for completing marginal dormancy release rather than substituting entirely for chilling requirements. Understanding dormancy breaking potential quality enables selection of appropriate agents for specific situations, timing of applications relative to dormancy status for optimal effectiveness, and recognition of the limitations of chemical dormancy breaking as a partial rather than complete substitute for natural chilling.
Dormancy_Breaking_Potential_Quality addresses the capacity of treatments to overcome dormancy and induce bud growth. This quality enables management intervention when natural dormancy release is inadequate. The class supports dormancy management through breaking potential characterization of various agents and conditions.
Dormanzfreisetzungskapazität
Dormanzlösungspotential
Knospentreibvermögen
bud forcing potential
chemical dormancy release
chemische Dormanzlösung
dormancy breaking efficacy
dormancy release capacity
Or E et al. (2000) Changes in endogenous ABA level in relation to the dormancy cycle in grapevines grown in a hot climate. Journal of Horticultural Science & Biotechnology 75(2): 190-194 Dokoozlian NK et al. (1995) Chilling exposure and hydrogen cyanamide interact in breaking dormancy of grape buds. HortScience 30(6): 1244-1247
Dormancy Breaking Potential Quality
Dormancy completion assessment comparing regions with different winter climates in Merlot reveals the chilling-dependent outcomes affecting spring performance, with cold-winter region showing complete dormancy (chilling requirement fully satisfied by January 31, subsequent heat accumulation driving uniform budbreak with 96% of buds emerging within 8-day window, strong initial growth), while mild-winter region shows incomplete dormancy (chilling only 75% of requirement by February 28, subsequent budbreak showing only 72% emergence spread across 22 days, weak and erratic growth requiring compensatory management), demonstrating how dormancy completion quality determines the foundation for seasonal productivity.
Dormancy completion quality is a dormancy transition quality that represents the degree to which the full dormancy cycle has been successfully concluded, characterizing the satisfaction of all dormancy requirements (chilling for endodormancy release, heat for budbreak initiation) that enables normal spring growth resumption with the vigor, uniformity, and fruitfulness expected from properly conditioned buds. Within the BFO framework, this quality specifically addresses the completion dimension of dormancy transitions, distinguishing full cycle completion from partial or incomplete dormancy processing while recognizing that dormancy completion indicates whether buds have been properly conditioned for spring performance, with incomplete completion producing the characteristic symptoms of inadequate chilling (delayed budbreak, reduced budbreak percentage, erratic timing, poor fruitfulness) or inadequate subsequent conditioning. The ontological significance lies in dormancy completion determining the quality of spring growth initiation and subsequent vine performance, where complete dormancy cycling produces uniform budbreak with high percentage, synchronized timing, and normal fruitfulness from buds that have been fully released from all dormancy constraints, while incomplete cycling produces the problematic spring performance that can affect the entire season's productivity. The criteria for dormancy completion include high budbreak percentage (typically >90% of retained buds producing shoots), uniform timing (narrow window from first to last budbreak), rapid response to favorable conditions (short lag from consistent warmth to visible growth), and normal shoot development (adequate vigor without the weak, erratic growth characteristic of poorly conditioned buds), with these criteria providing the practical assessment of whether dormancy has been properly completed. The factors affecting dormancy completion include the adequacy of chilling relative to cultivar requirements (the primary determinant of endodormancy release), the consistency of chilling (interrupted chilling potentially less effective than continuous exposure), and the heat accumulation pattern following endodormancy release (adequate warmth to drive budbreak after chilling satisfaction), with deficiencies in any factor potentially compromising completion. Understanding dormancy completion quality enables assessment of spring readiness, prediction of budbreak characteristics from dormancy satisfaction status, and identification of incomplete dormancy as the cause of problematic spring performance in situations where chilling may have been inadequate.
Dormancy_Completion_Quality addresses the successful conclusion of the full dormancy cycle enabling normal spring performance. This quality depends on adequate chilling and subsequent heat accumulation. The class supports spring readiness assessment through dormancy completion characterization.
Dormanzabschluss
Dormanzerfüllung
Konditionierungsabschluss
Zyklusabschluss
conditioning completion
cycle completion
dormancy fulfillment
dormancy satisfaction
Dokoozlian NK (1999) Chilling temperature and duration interact on the budbreak of 'Perlette' grapevine cuttings. HortScience 34(6): 1054-1056 Lavee S, May P (1997) Dormancy of grapevine buds—facts and speculation. Australian Journal of Grape and Wine Research 3(1): 31-46
Dormancy Completion Quality
Dormancy depth assessment tracking Chardonnay buds through winter reveals the characteristic progression from deep to shallow dormancy, with early December showing maximum endodormancy depth (buds unresponsive to forcing conditions, 0% budbreak after 30 days at 20°C), mid-January showing intermediate depth after partial chilling (45% budbreak under forcing, delayed and erratic), and late February showing shallow dormancy after chilling completion (95% budbreak within 14 days under forcing, rapid and uniform), demonstrating how dormancy depth progressively decreases through winter as chilling accumulation releases endodormancy and transitions buds to the ecodormant state responsive to spring temperatures.
Dormancy depth quality is a phenological quality that represents the intensity of the dormant state in grapevine buds, characterizing the degree of developmental arrest and the magnitude of environmental signals required to release buds from dormancy and resume active growth, with dormancy depth varying through the winter period as buds transition through distinct physiological phases requiring different release mechanisms. Within the BFO framework, this quality addresses the dormancy intensity dimension of bud physiology, distinguishing the depth or strength of dormancy from the timing of dormancy transitions while recognizing that dormancy depth determines the responsiveness of buds to growth-promoting conditions, with deep dormancy requiring substantial environmental conditioning (chilling) before growth can resume regardless of favorable temperatures, and shallow dormancy permitting growth resumption with minimal conditioning. The ontological significance lies in dormancy depth providing the protective mechanism that prevents premature budbreak during winter warm spells, where deep endodormancy ensures that buds remain quiescent even if temperatures temporarily become favorable for growth, protecting against the frost damage that would follow growth initiation during unstable winter weather, while the progressive shallowing of dormancy through winter eventually permits spring growth resumption when seasonal risk has diminished. The two-phase model of grapevine dormancy distinguishes endodormancy (internally controlled dormancy requiring chilling accumulation for release, typically dominant from late autumn through mid-winter) from ecodormancy (environmentally imposed dormancy where buds are physiologically capable of growth but prevented by unfavorable temperatures, typically dominant from late winter through budbreak), with dormancy depth reflecting the current balance between these phases and the conditioning status that determines growth potential. The factors affecting dormancy depth include cultivar genetics (determining chilling requirements and dormancy intensity), autumn conditions (affecting dormancy induction depth), winter temperatures (providing chilling that releases endodormancy), and the interaction between accumulated chilling and subsequent warm exposure (determining the transition from endodormancy to ecodormancy). Understanding dormancy depth quality enables assessment of bud physiological status through winter, prediction of budbreak timing based on chilling satisfaction and subsequent heat accumulation, and recognition of the climate change implications for dormancy regulation in regions where warming winters may provide insufficient chilling for complete endodormancy release.
Dormancy_Depth_Quality serves as the parent class for the two dormancy phases (Endodormancy, Ecodormancy). This quality addresses the intensity of developmental arrest affecting bud responsiveness to growth conditions. The class supports winter bud status assessment and budbreak prediction through dormancy depth characterization.
Dormanzintensität
Dormanztiefe
Knospenruhetiefe
Winterruhestärke
bud quiescence depth
dormancy intensity
dormancy strength
winter rest intensity
Lang GA et al. (1987) Endo-, para-, and ecodormancy: physiological terminology and classification for dormancy research. HortScience 22(3): 371-377 Lavee S, May P (1997) Dormancy of grapevine buds—facts and speculation. Australian Journal of Grape and Wine Research 3(1): 31-46
Dormancy Depth Quality
In Pinot Gris vines in Oregon's Willamette Valley, dormancy depth reduction progresses through winter as demonstrated by forcing tests showing days to 50% budbreak decreasing from more than 55 days in early December before significant chilling accumulation, to 35 days by mid-January after 500 chilling hours, to 18 days by mid-February after 900 chilling hours, with this progressive reduction correlating with measured decreases in bud ABA concentration from 195 to 85 ng per gram fresh weight over the same period.
Dormancy depth reduction in Vitis vinifera represents the progressive decrease in dormancy intensity that occurs during chilling accumulation, reflecting the gradual dismantling of dormancy-maintaining mechanisms and the restoration of growth competence in endodormant buds. This process occurs continuously through the winter as effective chilling temperatures induce the physiological changes that reduce internal growth inhibition, eventually reaching the point where dormancy is fully released and buds transition to the ecodormant state. The measurement of dormancy depth reduction relies on forcing tests conducted at intervals through winter, with decreasing time to budbreak and increasing percentage of buds breaking indicating progressive reduction in dormancy depth as chilling accumulates. The physiological mechanisms underlying dormancy depth reduction include decreased ABA concentration through activation of catabolic pathways particularly ABA 8'-hydroxylase, declining expression of dormancy-associated MADS-box genes, progressive removal of repressive histone modifications from growth-related gene promoters, and gradual restoration of symplastic connectivity through callose degradation at plasmodesmata. The relationship between chilling accumulation and dormancy depth reduction is generally continuous but not necessarily linear, with some evidence suggesting that dormancy depth decreases slowly during early chilling accumulation when inhibitory mechanisms are strongest, more rapidly during mid-winter as multiple mechanisms are simultaneously affected, and more slowly again as dormancy approaches full release. The rate of dormancy depth reduction varies among cultivars in relation to their chilling requirements and the intensity of their maximum dormancy, with high-chill cultivars showing slower reduction per unit of chilling than low-chill cultivars. Environmental factors beyond temperature can modulate dormancy depth reduction, with some evidence that photoperiod may influence the rate of reduction at equivalent chilling accumulation, and water stress potentially affecting dormancy status. The completion of dormancy depth reduction to the point of full dormancy release restores bud growth competence, though environmental temperatures typically continue to constrain actual growth until spring warming occurs.
Dormancy_Depth_Reduction represents the functional outcome of Chilling_Accumulation describing the progressive decrease in dormancy intensity. Distinguished from Maximum_Dormancy_Depth which describes peak dormancy, and from Chilling_Requirement_Satisfaction which marks completion of reduction. Measured through forcing test response showing decreasing budbreak time. Rate varies among cultivars and through the chilling period. Associated with declining ABA, decreasing DAM expression, and chromatin derepression. Duration spans most of winter chilling period.
Dormanz-Abschwächung
Dormanztiefe-Reduktion
abnehmende Dormanzintensität
decreasing dormancy depth
dormancy intensity decline
dormancy weakening
fortschreitende Dormanzlösung
progressive dormancy release
Lavee S, May P (1997) Dormancy of grapevine buds - facts and speculation. Australian Journal of Grape and Wine Research 3: 31-46 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Zheng C et al (2015) Abscisic acid catabolism enhances dormancy release of grapevine buds. Plant Cell and Environment 38: 2490-2501
Dormancy Depth Reduction
Grapevine dormancy initiates in response to shortening photoperiod (<13 hours) and declining temperatures (<15°C) during autumn, with bud endodormancy fully established by 4-8 weeks post-veraison in temperate climates. Dormancy depth peaks in mid-winter, with forcing requirement (days to budbreak at 25°C) reaching 30-60 days in December-January for northern hemisphere cultivars. Tropical cultivars such as those grown in Thailand and Brazil show reduced dormancy capacity, enabling continuous growth or managed double-cropping. Dormancy duration from onset to natural budbreak spans 120-180 days depending on latitude and cultivar.
A phenological timing disposition that is the inherited capacity of a Vitis vinifera plant to enter, maintain, and exit the dormant state (endodormancy) characterized by arrested meristem activity, cold hardiness development, and metabolic quiescence during the winter period, with genotype-specific requirements for dormancy induction and release.
Dormancy_Disposition serves as the parent class for chilling requirement, dormancy release, and paradormancy traits. Endodormancy is physiologically distinct from ecodormancy (environmentally imposed) and paradormancy (correlatively imposed). The VvDAM and VvFT genes regulate dormancy induction and maintenance. Dormancy failure in warm climates causes erratic budbreak and reduced fruitfulness, necessitating chemical dormancy-breaking treatments (hydrogen cyanamide) or selection of low-chilling cultivars.
Vegetationsruhe-Disposition
endodormancy capacity
rest period disposition
winter dormancy trait
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Rubio S et al. (2019) Dormancy in grapevine buds: I. Physiological and genetic regulation. Acta Horticulturae 1248:29-36
Dormancy Disposition
In Grüner Veltliner vines in the Wachau, Austria, dormancy establishment is achieved by mid-November when bud scales have fully formed enclosing dormant meristems, shoot lignification has completed to within 3-4 nodes of tips, leaf abscission is complete, ABA concentrations in buds have reached maximum levels of 150-200 ng per gram fresh weight, and buds show no growth response to favorable temperatures indicating full endodormancy at BBCH stage 93.
Dormancy establishment in Vitis vinifera represents the culminating phase of dormancy induction during which the vine achieves the fully dormant state characterized by complete growth cessation, maximum physiological preparation for winter survival, and the internal conditions that will subsequently require specific environmental cues for dormancy release. This phase occurs during late autumn, typically from late October through November in Northern Hemisphere viticulture, as the accumulated effects of photoperiod response, temperature acclimation, and growth cessation converge to produce the stable dormant condition that will persist through winter. The establishment of dormancy involves the consolidation of multiple physiological changes including the complete arrest of all meristematic activity, the achievement of maximum cold hardiness appropriate to current acclimation conditions, the accumulation of abscisic acid to levels that actively maintain dormancy, and the developmental changes in bud structure that provide physical protection for dormant meristems. Morphological indicators of dormancy establishment include the completion of bud scale development providing physical protection for enclosed meristems, the full lignification of current-season shoots visible as complete browning of periderm to within a few nodes of shoot tips, the abscission or final senescence of remaining leaves, and the characteristic bare-cane winter appearance of fully dormant vines. At the molecular level, dormancy establishment involves the expression of dormancy-associated MADS-box genes including the DAM gene family that actively maintain the dormant state, chromatin modifications that establish dormancy-specific gene expression patterns, and the accumulation of dormancy markers including specific proteins and metabolites characteristic of the dormant condition. The transition from dormancy induction to established dormancy represents a qualitative shift from a process of becoming dormant to a state of being dormant, with the established dormancy being self-maintaining until environmental cues including chilling accumulation eventually lead to dormancy release. The timing and completeness of dormancy establishment significantly affects winter survival, with well-established dormancy providing maximum protection against mid-winter cold injury, while incomplete dormancy establishment leaves vines vulnerable to cold damage and may result in erratic budbreak in spring. Cultivar differences in dormancy establishment timing and depth contribute to regional adaptation, with cultivars showing earlier and deeper dormancy establishment being better suited to regions with severe winters.
Dormancy_Establishment represents the final phase of Dormancy_Induction, marking achievement of the fully dormant state. Distinguished from earlier induction phases by representing a stable state rather than transitional process, and from Endodormancy which represents the subsequent dormancy maintenance phase. Subdivides into Bud_Scale_Formation, ABA_Accumulation, and Deep_Dormancy_Entry as component processes. Corresponds to BBCH stage 93. Marks the beginning of the period requiring chilling for dormancy release. Critical for winter survival and proper spring growth resumption.
Dormanz-Etablierung
Dormanz-Konsolidierung
Ruhestandsetablierung
dormancy completion
dormancy consolidation
dormant state establishment
vollständige Dormanz-Erreichung
Dormancy Establishment
In Chenin Blanc vines in the Loire Valley, dormancy exit completion is achieved in mid-April approximately 8 days after bud swelling initiation, with green tip emergence at BBCH stage 07 confirming irreversible transition to active growth, accumulated GDD of approximately 180 units representing sufficient heat exposure for complete dormancy exit, and subsequent shoot development proceeding at rates of 3-5 cm per day as the vine enters the rapid growth phase of the new growing season.
Dormancy exit completion in Vitis vinifera represents the definitive termination of the dormancy cycle when the transition to active growth has been fully accomplished and the vine has irreversibly entered the new growing season, marked by the achievement of visible green tissue emergence that confirms successful progression beyond the dormancy break threshold. This completion represents the final stage of dormancy release, occurring when developmental progress has advanced from bud swelling through scale separation to the emergence of green shoot tissue that demonstrates committed growth progression. The morphological indicator of dormancy exit completion is green tip emergence corresponding to BBCH stage 07, where the expanding primordial shoot has pushed sufficiently through separating bud scales to expose the green tissue of emerging leaves, representing a clear visual confirmation that active growth is underway. The physiological state at dormancy exit completion involves fully active cell division in meristematic tissues, established patterns of cell expansion driving shoot elongation, mobilization of reserves from woody tissues providing resources for growth, and the hormonal and metabolic systems supporting sustained developmental activity. The irreversibility at dormancy exit completion distinguishes this stage from earlier phases where return to quiescence was possible, with green tip emergence representing commitment to growth that will continue regardless of subsequent temperature fluctuations, though growth rate will vary with environmental conditions. The timing of dormancy exit completion typically follows bud swelling initiation by 5-10 days depending on temperature, with warm conditions accelerating progression while cool conditions extend the interval, and the specific timing being predictable from continued heat accumulation following the initial budbreak threshold. The significance of dormancy exit completion for vineyard management relates to the confirmation of successful dormancy release and the establishment of the developmental trajectory for the new season, with uniform completion across a vineyard indicating consistent dormancy satisfaction and predicting uniform subsequent development. The frost sensitivity at dormancy exit completion is substantially elevated relative to earlier stages, with green tip tissue being damaged at temperatures of -2 to -3°C compared to -4 to -6°C for swelling buds and -15°C or below for dormant buds, making frost protection particularly critical during this vulnerable transition period.
Dormancy_Exit_Completion represents the terminal event of Dormancy_Break and the entire Dormancy cycle marking definitive transition to active growth. Distinguished from Bud_Swelling_Initiation by representing complete rather than initial transition. Corresponds to BBCH stage 07 green tip emergence. Represents irreversible commitment to growth. Timing typically 5-10 days after bud swelling initiation. Maximum frost sensitivity at this stage requiring vigilant protection. Marks beginning of new annual growth cycle.
Dormanzaustritts-Vollendung
Dormanzzyklus-Abschluss
complete dormancy ending
dormancy cycle completion
dormancy termination completion
full dormancy exit
komplette Dormanzbeendigung
vollständiger Dormanzaustritt
Lorenz DH et al (1995) Growth stages of the grapevine: Phenological growth stages of the grapevine - BBCH-scale. Australian Journal of Grape and Wine Research 1: 100-103 Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235
Dormancy Exit Completion
In Cabernet Sauvignon vineyards of Napa Valley, California, dormancy induction commences in late September when photoperiod decreases below 12.5 hours, with visible signs including cessation of shoot elongation, periderm browning advancing acropetally along shoots, and initial bud scale formation, progressing through October as night temperatures consistently fall below 10°C to achieve full dormancy establishment by early November.
Dormancy induction in Vitis vinifera represents the transitional phase during which actively growing vines perceive and respond to environmental signals that trigger the cessation of growth and the initiation of physiological preparations for winter dormancy. This critical period typically occurs during late summer through autumn, from August through October in Northern Hemisphere viticulture, as declining photoperiod and decreasing temperatures interact to activate the genetic and hormonal programs that progressively inhibit meristematic activity and redirect metabolism toward cold acclimation and reserve accumulation. The process is initiated primarily by photoperiod sensing, with the shortening day length below a cultivar-specific threshold serving as the most reliable environmental cue for upcoming winter conditions, detected through phytochrome-mediated pathways in leaf tissues that generate systemic signals affecting bud and cambial meristems. Temperature serves as a secondary but important modulating factor, with cool temperatures enhancing the dormancy induction response and warm temperatures potentially delaying the process, creating the risk of inadequate cold hardiness development when unseasonably warm autumn conditions occur. Physiologically, dormancy induction involves coordinated changes including cessation of shoot apical meristem activity, formation of terminal buds or shoot tip abscission, development of bud scales from modified leaves that provide physical protection, accumulation of abscisic acid which inhibits growth and promotes stress tolerance gene expression, and initiation of cold acclimation processes that progressively increase freezing tolerance. The hormonal transition during dormancy induction involves declining auxin and gibberellin levels that normally promote growth, increasing abscisic acid concentrations that inhibit cell division and elongation, and altered cytokinin transport from roots that affects bud activity. Carbohydrate metabolism shifts during induction from supporting active growth toward starch accumulation in woody tissues and synthesis of cryoprotective sugars including raffinose family oligosaccharides. The timing and completeness of dormancy induction significantly affects winter survival, with vines that fail to achieve adequate dormancy before severe cold exposure suffering increased cold injury, while premature or excessive dormancy induction may reduce photosynthetic capacity and reserve accumulation during favorable late-season conditions.
Dormancy_Induction represents the first major phase within the Dormancy hierarchy, encompassing the transition from active growth to dormant state. Distinguished from Endodormancy which represents the established dormant condition, and from active growth phases by the progressive inhibition rather than absence of growth activity. Subdivides into Photoperiod_Response, Temperature_Acclimation, Growth_Cessation, and Dormancy_Establishment as component processes. Duration typically spans 6-10 weeks depending on cultivar and environmental conditions. Critical for establishing adequate cold hardiness before winter.
Dormanzeinleitung
Dormanzinduktion
Ruheinduktion
Vegetationsruhebeginn
autumn dormancy entry
dormancy initiation
dormancy onset
growth cessation induction
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Wake CMF, Fennell A (2000) Morphological, physiological and dormancy responses of three Vitis genotypes to short photoperiod. Physiologia Plantarum 109: 203-210
Dormancy Induction
Dormancy induction assessment comparing autumn management effects in Chardonnay reveals the factors affecting transition completeness, with balanced vines (moderate crop, adequate water) showing timely induction (growth cessation by October 15, measurable endodormancy by November 1, cold hardiness developing on schedule), while over-cropped, drought-stressed vines show delayed induction (growth continuation until November 5 from delayed sugar translocation, incomplete endodormancy establishment by November 15, cold hardiness lagging), with the induction delay creating vulnerability to early November freeze that damaged 25% of buds on stressed vines versus 5% on balanced vines, demonstrating how induction quality affects winter survival outcomes.
Dormancy induction quality is a dormancy transition quality that represents the initiation and establishment of the dormant state in grapevine buds during autumn, characterizing the transition from active growth competence to endodormancy that involves coordinated physiological changes preparing buds for winter survival including growth cessation, cold acclimation initiation, and metabolic shifts toward storage and protection functions. Within the BFO framework, this quality specifically addresses the induction phase dimension of dormancy transitions, distinguishing the entry into dormancy from subsequent deepening or release while recognizing that successful induction establishes the dormancy framework within which winter cold hardiness develops, with inadequate induction potentially leaving buds vulnerable to cold damage or to premature growth during autumn warm spells. The ontological significance lies in dormancy induction establishing the protective state essential for perennial survival in temperate climates, where the coordinated changes during induction (ABA accumulation, gibberellin sensitivity reduction, growth inhibitor accumulation, storage compound deposition, cold hardiness gene expression) together create the physiological condition that enables survival of winter conditions that would kill actively growing tissue. The environmental triggers for dormancy induction include shortening photoperiod (the primary signal in many woody plants including grapevine), declining temperatures (reinforcing the photoperiod signal and contributing to cold acclimation), and potentially water and nutrient limitation (accelerating senescence and dormancy entry), with these signals integrated through hormonal and transcriptional networks that coordinate the induction response. The timing and completeness of dormancy induction vary among cultivars and with autumn conditions, where early induction in response to marginal signals may sacrifice late-season photosynthate accumulation while delayed induction may leave buds inadequately prepared for early cold events, creating the adaptive tradeoff between maximizing growing season productivity and ensuring adequate winter protection. Understanding dormancy induction quality enables assessment of autumn bud status, prediction of cold hardiness development timing, and recognition of conditions that may compromise induction with consequences for winter survival and spring performance.
Dormancy_Induction_Quality addresses the establishment of endodormancy during autumn transition from active growth. This quality is triggered by photoperiod and temperature affecting cold hardiness development. The class supports winter preparation assessment through dormancy induction characterization.
Dormanzeintritt
Herbsttransition
autumn transition
dormancy entry
dormancy establishment
endodormancy initiation
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10(1-2): 201-235 Wake CMF, Fennell A (2000) Morphological, physiological and dormancy responses of three Vitis genotypes to short photoperiod. Physiologia Plantarum 109(2): 203-210
Dormancy Induction Quality
In Cabernet Sauvignon vines in Napa Valley, dormancy maintenance persists through December and January as demonstrated by forcing tests showing continued growth inhibition with budbreak requiring more than 40 days at 22°C despite exposure to warm periods with temperatures exceeding 15°C for several consecutive days in early January, with the maintaining mechanisms only progressively weakening as chilling accumulates toward the cultivar requirement of approximately 900-1000 hours.
Dormancy maintenance in Vitis vinifera represents the sustained preservation of the endodormant state during the winter period, encompassing the active physiological mechanisms that prevent premature growth resumption and maintain bud quiescence until sufficient chilling has been accumulated to release dormancy. This process operates throughout the endodormancy period, from dormancy establishment in late autumn until chilling-dependent dormancy release in late winter, ensuring that buds remain growth-incompetent during the period when environmental conditions would make growth resumption potentially lethal due to subsequent freezing exposure. The maintenance of dormancy is not a passive state but involves active regulatory processes that sustain growth inhibition, including continued expression of dormancy-associated MADS-box genes that repress growth-related transcription, ongoing ABA signaling that maintains cell cycle arrest, and the preservation of chromatin modifications that silence genes required for meristematic activity. The ecological function of dormancy maintenance is to prevent the vine from responding to temporary favorable conditions during winter, such as warm spells that might otherwise trigger growth in non-dormant buds, thereby protecting the vine from the catastrophic consequences of freeze damage to actively growing tissues. The effectiveness of dormancy maintenance varies through the endodormancy period, being strongest during deep endodormancy when all maintaining mechanisms operate at maximum intensity, and progressively weakening as chilling accumulation gradually dismantles inhibitory mechanisms and reduces dormancy depth. The robustness of dormancy maintenance against premature release has been selected through evolution in grapevine populations exposed to variable winter climates where temporary warm periods are common, with cultivars from continental climates typically showing more robust maintenance than those from Mediterranean regions where winters are milder and more predictable. Factors that can compromise dormancy maintenance include exposure to dormancy-breaking chemicals such as hydrogen cyanamide that artificially override maintaining mechanisms, severe dehydration stress that may disrupt normal hormonal and metabolic regulation, and potentially inadequate dormancy establishment that fails to fully activate maintaining mechanisms. The gradual transition from active dormancy maintenance to dormancy release competence represents the physiological basis for the chilling requirement, with each increment of chilling exposure reducing the strength of maintaining mechanisms until they can no longer prevent growth in response to favorable conditions.
Dormancy_Maintenance represents a functional phase within Endodormancy addressing the active preservation of dormant state. Distinguished from Deep_Endodormancy which represents maximum dormancy intensity, and from Chilling_Accumulation which addresses progressive dormancy release. Subdivides into Sustained_Growth_Inhibition, Continued_Metabolic_Suppression, and Endodormancy_Persistence as component aspects. Duration spans most of endodormancy period from establishment through progressive release. Involves active regulatory mechanisms rather than passive quiescence.
Dormanz-Aufrechterhaltung
Dormanz-Persistenz
Ruheerhaltung
anhaltende Dormanz
dormancy persistence
dormancy preservation
dormant state maintenance
sustained dormancy
Lang GA et al (1987) Endo-, para-, and ecodormancy: physiological terminology and classification for dormancy research. HortScience 22: 371-377 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Rohde A, Bhalerao RP (2007) Plant dormancy in the perennial context. Trends in Plant Science 12: 217-223
Dormancy Maintenance
In Pinot Noir vines of the Côte d'Or, Burgundy, dormancy release proceeds through February and March with chilling completion achieved by early February after approximately 1100 chilling hours, followed by progressive heat accumulation through February and March reaching approximately 150 growing degree days by late March, with visible budbreak commencing in early April as the final stage of dormancy release marks the transition to active growth at BBCH stage 05.
Dormancy release in Vitis vinifera represents the terminal phase of the dormancy cycle during which all remaining constraints on growth are removed, enabling the transition from dormant winter status to active spring growth through the coordinated completion of chilling requirements, restoration of bud competence, and response to warming temperatures that triggers budbreak initiation. This process encompasses the final events of dormancy termination occurring during late winter and early spring, integrating the completion of endodormancy release through chilling satisfaction with the subsequent ecodormancy release through heat accumulation to achieve the fully growth-competent and growth-active state. The physiological basis of dormancy release involves the completion of all chilling-dependent changes that dismantle endodormancy maintaining mechanisms, followed by the heat-dependent activation of growth processes that overcomes the environmental constraints of ecodormancy, culminating in the visible developmental changes of budbreak. The molecular markers of dormancy release completion include ABA concentrations reduced to levels characteristic of growing tissues, DAM gene expression at minimal or undetectable levels, growth gene promoters fully derepressed and transcriptionally active, and cell cycle machinery engaged in active progression through division cycles. The timing of dormancy release is determined by the sequential satisfaction of chilling and heat requirements, with chilling completion typically occurring during late January to February and heat accumulation proceeding through late winter and early spring until the threshold for budbreak is reached, typically after 100-300 growing degree days above the base temperature. The practical importance of dormancy release timing relates to the synchronization of growth with favorable environmental conditions, with optimal release timing balancing the benefits of early growth initiation against the risks of frost damage to emerging shoots, and with release timing variability among cultivars contributing to their adaptation to different climatic regions. The consequences of incomplete or abnormal dormancy release include delayed, prolonged, or uneven budbreak that complicates vineyard management, reduced shoot vigor from inadequate chilling, and potential yield impacts from poorly developed shoots, highlighting the importance of adequate dormancy release for vineyard productivity. Climate change poses challenges to dormancy release in many regions through effects on both chilling accumulation and heat accumulation timing, potentially disrupting the evolved synchronization between dormancy release and optimal growth conditions.
Dormancy_Release represents the fourth and final major phase within the Dormancy hierarchy, encompassing all processes leading to growth resumption. Distinguished from Ecodormancy which represents maintained quiescence, while Dormancy_Release specifically addresses the termination of dormancy and transition to growth. Subdivides into Chilling_Completion, Bud_Competence_Restoration, Heat_Accumulation_Response, and Dormancy_Break as component phases. Timing determined by sequential chilling and heat requirement satisfaction. Corresponds to transition from BBCH 00 through BBCH 09.
Dormanzlösung
Wachstumswiederaufnahme
dormancy break
growth resumption
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Lang GA et al (1987) Endo-, para-, and ecodormancy: physiological terminology and classification for dormancy research. HortScience 22: 371-377 Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle
Dormancy Release
In Sauvignon Blanc vines in Marlborough, New Zealand, dormancy release competence is achieved by early August following accumulation of approximately 850 chilling hours, demonstrated by forcing tests showing complete budbreak within 11 days at 20°C, with competent buds subsequently responding rapidly to field warming in September, initiating visible bud swelling within 5-7 days of mean temperatures exceeding 10°C.
Dormancy release competence in Vitis vinifera represents the restored capacity for growth following sufficient chilling accumulation, describing the physiological state in which buds have regained the ability to respond to growth-promoting conditions after the internal constraints of endodormancy have been dismantled through chilling-dependent processes. This competence represents a qualitative transition from the endodormant state where growth is impossible regardless of conditions to a growth-ready state where only environmental factors constrain actual growth initiation. The physiological basis of dormancy release competence involves the successful completion of chilling-dependent changes including reduction of ABA to non-inhibitory levels, silencing of DAM gene expression, removal of repressive chromatin marks from growth-related genes restoring their transcriptional responsiveness, and re-establishment of symplastic communication pathways that enable coordination of growth responses. The cellular characteristics of growth-competent buds include the presence of cell cycle regulators poised for activation, metabolic enzyme systems capable of rapid upregulation to support biosynthesis, and hormone receptor systems sensitized to respond to growth-promoting signals. The concept of competence distinguishes between the potential for growth and actual growth, with competent buds remaining quiescent during ecodormancy when environmental temperatures are insufficient for growth but capable of immediate response when conditions become favorable. Testing for dormancy release competence involves forcing assays that expose buds to optimal growth conditions and assess the speed and completeness of budbreak response, with competent buds showing rapid budbreak within 10-14 days while buds lacking competence due to incomplete dormancy release require extended periods or fail to break entirely. The achievement of dormancy release competence marks the transition from endodormancy to ecodormancy in the dormancy classification system, representing a fundamental change in the nature of growth control from internal inhibition to environmental limitation. The uniformity of competence achievement across buds on a vine affects the synchrony of subsequent budbreak, with simultaneous competence acquisition producing uniform budbreak while variable competence results in extended budbreak periods.
Dormancy_Release_Competence represents the functional capacity aspect of Chilling_Requirement_Satisfaction describing restored growth potential. Distinguished from actual growth which requires favorable environmental conditions, while competence represents the potential for growth when conditions permit. Marks the endodormancy to ecodormancy transition. Verified through rapid forcing test response within 10-14 days. Involves completion of all chilling-dependent physiological changes. Uniformity of competence affects budbreak synchrony.
Dormanzlösungs-Kompetenz
Wachstumskompetenz-Wiederherstellung
Wachstumswiederaufnahme-Fähigkeit
budbreak readiness
growth competence restoration
growth resumption competence
Lang GA et al (1987) Endo-, para-, and ecodormancy: physiological terminology and classification for dormancy research. HortScience 22: 371-377 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Lavee S, May P (1997) Dormancy of grapevine buds - facts and speculation. Australian Journal of Grape and Wine Research 3: 31-46
Dormancy Release Competence
Dormancy release in Chardonnay occurs progressively during January-February in northern temperate zones, with forcing time at 25°C decreasing from 45-60 days in early December to 10-15 days by late February as chilling accumulates. Riesling demonstrates abrupt dormancy release once chilling threshold is met, with forcing time dropping from 50 to 12 days within a 2-week window. Biological markers of dormancy release include increased bud respiration rate (2-3× dormant levels), resumed cell division in meristems detectable by BrdU incorporation, and upregulation of VvFT expression. Field budbreak follows dormancy release by 2-6 weeks depending on spring temperature progression.
A dormancy disposition that is the inherited capacity of a Vitis vinifera plant to transition from endodormancy to ecodormancy once chilling requirements are satisfied, characterized by acquisition of growth competence in bud meristems and responsiveness to forcing temperatures for budbreak initiation.
Dormancy release is distinct from budbreak itself, representing the internal transition from endodormancy (inhibition from within the bud) to ecodormancy (inhibition from environmental conditions). Once dormancy is released, buds will grow whenever temperatures exceed the threshold (approximately 10°C base temperature). The timing of dormancy release relative to field temperatures determines frost risk, with early release in warm winter regions creating vulnerability to late cold events. Hormonal changes during release include declining ABA and increasing cytokinin levels in bud tissues.
Endodormanz-Aufhebung
Ruhebeendigung
dormancy exit capacity
endodormancy termination trait
rest completion
Lavee S, May P (1997) Dormancy of grapevine buds: facts and speculation. Australian Journal of Grape and Wine Research 3:31-46 Pérez FJ et al. (2008) Hydrogen peroxide and glucanase activity during budbreak in grapevine. Vitis 47:17-22
Dormancy Release Disposition
Dormancy transition tracking through the annual cycle in Pinot Noir reveals the sequential phases requiring distinct environmental conditions, with October-November showing induction transition (shortening photoperiod triggering metabolic shifts toward dormancy despite still-favorable temperatures), December-January showing deepening transition (maximum endodormancy depth coinciding with accumulating chilling), February showing release transition (chilling satisfaction enabling shift to ecodormancy), and March showing completion transition (heat accumulation driving budbreak), with successful vineyard performance requiring appropriate conditions for each transition phase in sequence.
Dormancy transition quality is a phenological quality that represents the progression of grapevine buds through the changes that establish, deepen, maintain, and ultimately release dormancy, characterizing the dynamic trajectory of dormancy status through the seasonal cycle rather than the static depth at any moment, with transitions including the induction of dormancy in autumn, the progressive changes through winter, and the release processes enabling spring growth. Within the BFO framework, this quality addresses the temporal progression dimension of dormancy, distinguishing the transitions between dormancy states from the dormancy depth itself while recognizing that dormancy transitions involve coordinated physiological changes (hormonal shifts, metabolic reprogramming, gene expression changes, chromatin modifications) that must proceed in proper sequence for successful seasonal cycling. The ontological significance lies in dormancy transitions determining the successful completion of the annual dormancy cycle, where proper induction establishes winter protection, proper progression maintains cold hardiness while accumulating chilling, and proper release enables timely spring growth, with disruption of any transition potentially compromising vine survival, productivity, or the coordination of development with seasonal opportunities. The environmental and endogenous regulation of dormancy transitions involves the integration of photoperiod signals (triggering autumn induction), temperature patterns (driving both chilling accumulation and spring warming responses), and internal developmental programs (determining the cultivar-specific patterns and timing of transitions), with this integration enabling adaptive responses to the variable conditions encountered across different climates and years. The climate change implications for dormancy transitions include potential disruption of induction timing from altered autumn temperature and photoperiod relationships, modification of winter progression from changed chilling patterns, and shifting of spring release from warming trends, with these effects potentially desynchronizing dormancy transitions from the seasonal patterns to which cultivars are historically adapted. Understanding dormancy transition quality enables tracking of bud progression through dormancy phases, prediction of vulnerability windows during transition periods, and recognition of the environmental factors driving successful transition completion.
Dormancy_Transition_Quality serves as the parent class for specific transition phases (Induction, Completion, Breaking Potential). This quality addresses the dynamic progression through dormancy states rather than static depth. The class supports seasonal dormancy tracking through transition characterization.
Dormanzphasenwechsel
Dormanzprogression
Dormanztransition
dormancy cycling
dormancy phase change
dormancy progression
saisonale Dormanzdynamik
seasonal dormancy dynamics
Horvath DP et al. (2003) Knowing when to grow: signals regulating bud dormancy. Trends in Plant Science 8(11): 534-540 Cooke JEK et al. (2012) The dynamic nature of bud dormancy in trees: environmental control and molecular mechanisms. Plant, Cell & Environment 35(10): 1707-1728
Dormancy Transition Quality
A dormant bud on a pruned cane of Vitis vinifera cv. Chardonnay in mid-January in Burgundy, France, showing no external signs of growth activity despite mild ambient temperatures, with internal examination revealing arrested primordial development and accumulated starch reserves in bud tissues.
A bud in a state of temporarily suspended growth and development, characterized by reduced metabolic activity, protective scale coverage, and inability to resume growth despite potentially favorable environmental conditions, requiring specific environmental cues or passage of time for dormancy release. Dormant buds in Vitis vinifera serve as survival structures during unfavorable seasons and represent the primary mechanism for perennial growth habit, with dormancy regulated by complex interactions between photoperiod, temperature, hormonal signals (particularly abscisic acid and gibberellins), and cultivar-specific genetic programming.
Distinguish between endodormancy (internally regulated, requiring chilling accumulation for release), ecodormancy (environmentally imposed, released when conditions improve), and paradormancy (controlled by signals from other plant parts). Dormant buds differ from latent buds, which are suppressed by apical dominance but not truly dormant. Summer and winter dormancy represent fundamentally different physiological states despite similar external appearance.
Ruheknospe
quiescent bud
resting bud
schlafende Knospe
Based on Lang et al. (1987) "Endo-, para-, and ecodormancy= physiological terminology and classification for dormancy research" HortScience 22=371-377 Fennell & Hoover (1991) "Photoperiod influences growth, bud dormancy, and cold acclimation in Vitis labruscana and V. riparia" Journal of the American Society for Horticultural Science 116=270-273 Considine & Considine (2016) "On the language and physiology of dormancy and quiescence in plants" Journal of Experimental Botany 67=3189-3203.
Dormant Bud
Pinot Noir dormant embryos in freshly harvested seeds (BBCH 89, October harvest) show 0-5% germination without stratification despite full embryo development, but achieve 75-85% germination after 105-115 days moist stratification at 3-5°C with progressive ABA decline and GA increase during chilling period
A dormant embryo is a fully developed embryo within a mature Vitis vinifera seed that is metabolically quiescent and unable to germinate despite favorable environmental conditions (moisture, temperature, oxygen availability) due to endogenous physiological dormancy mechanisms requiring specific dormancy-breaking treatments. Grape seed dormancy is classified as physiological dormancy (PD) rather than morphological dormancy= embryos are anatomically complete at seed maturity but physiologically prevented from germinating by endogenous factors including hormone balance (high abscisic acid relative to gibberellins), metabolic inhibitors in seed coat or endosperm tissues, and embryo tissue characteristics requiring maturation. Dormancy in Vitis vinifera seeds serves important ecological functions= preventing germination during unfavorable seasons (preventing autumn germination that would expose seedlings to winter cold), ensuring germination timing aligns with favorable spring conditions after winter chilling, and promoting seed dispersal by preventing immediate germination. The dormancy-breaking requirement for grape seeds is cold stratification= extended exposure to moist chilling conditions (90-120 days at 1-5°C) that progressively reduces dormancy through= increased embryo gibberellin synthesis and sensitivity, decreased abscisic acid levels, breakdown of metabolic inhibitors, and increased embryo metabolic activity and enzyme production. Stratification duration requirements vary by= cultivar genetics (wild Vitis species typically require longer stratification 120-150 days compared to V. vinifera cultivars 90-120 days), seed maturity at harvest (over-mature seeds may have reduced dormancy), and storage conditions (warm dry storage can intensify dormancy while moist chilling progressively breaks it). Dormant embryos exhibit characteristic physiological states= low respiration rates (5-10% of germinating embryo rates), minimal protein synthesis, accumulated storage reserves (lipids, proteins) not yet mobilized, intact but quiescent meristematic tissues, and hormone profiles favoring dormancy maintenance (ABA=/GA ratio typically 5-10= during dormancy decreasing to 0.5-1= after stratification). Commercial viticulture and breeding programs manage dormancy through= controlled stratification protocols for breeding programs, embryo rescue techniques that bypass dormancy by culturing excised immature embryos, and gibberellin treatments that can partially substitute for stratification requirements in some applications.
Dormant_Embryo emphasizes physiological dormancy state and is distinguished from Developed_Embryo (structural maturation) by focus on metabolic quiescence and germination inhibition requiring specific dormancy-breaking treatments. Critical for understanding breeding program stratification requirements, embryo rescue timing decisions, seed storage and germination protocols, and ecological adaptations preventing off-season germination in wild populations.
Dormanter Embryo
Keimruhender Samenkeimling
Physiologically dormant embryo
Quiescent embryo
Ruhender Keim
Bouquet, A., & Davis, H. P. (1989). Culture in vitro d'embryons immatures de vigne (Vitis vinifera L.) appliquée à la sélection de variétés de cuve sans pépins. Agronomie, 9(6), 565-574. | Nikolaeva, M. G. (1977). Factors controlling the seed dormancy pattern. In The Physiology and Biochemistry of Seed Dormancy and Germination (pp. 51-74). North-Holland Publishing Company.
Dormant Embryo
A dormant vine of Cabernet Sauvignon/110R harvested in January after 1,200 chilling hours bears a Dormant_Vine_Role, buds fully endodormant, stored at 2°C and 95% RH until March shipment—the dormant state enabling 8-week storage without quality deterioration. A dormant bare-root vine of Pinot Noir/3309C serves in a Dormant_Vine_Role for spring planting, the buds beginning ecodormancy release as soil temperatures rise, with root growth initiating 2-3 weeks before visible budbreak. A dormant grafted vine held in cold storage until late planting serves in a Dormant_Vine_Role, extended storage maintaining dormancy until site preparation completes and planting can proceed.
A nursery plant role that inheres in a grapevine in the dormant physiological state, with buds in endodormancy or ecodormancy, no active growth, and metabolism reduced to maintenance levels—the standard condition for bare-root vine harvest, storage, shipping, and winter/early spring planting. Dormant vines have completed the annual growth cycle, accumulated cold exposure satisfying chilling requirements (typically 400-1500 hours below 7°C depending on variety), and entered the resting state that permits handling without the wilting, desiccation, and growth disruption that would affect actively growing plants. The dormant condition enables extended storage (4-6 months at 0-4°C with adequate humidity), long-distance shipping without specialized life-support systems, and transplanting with root systems temporarily inactive—allowing root pruning, soil removal, and replanting without the immediate moisture demands of actively transpiring plants. Dormant vine planting is timed before budbreak, when soil moisture and temperature support root initiation before shoot emergence creates water demand exceeding the newly planted root system's capacity.
Dormant_Vine_Role captures the physiological state that enables traditional bare-root vine commerce, where the temporary suspension of active growth permits handling and transplanting that would severely stress or kill actively growing plants. Dormancy is not merely the absence of growth but a regulated physiological condition involving hormonal controls (ABA accumulation, auxin and cytokinin decline), metabolic adjustments (reduced respiration, mobilization of starch reserves), and protective mechanisms (cold hardiness development, tissue dehydration tolerance). The commercial importance of dormancy reflects the logistical requirements of grapevine distribution—plants must survive weeks between nursery harvest and vineyard planting, with dormancy providing the physiological basis for this tolerance. Proper dormant vine handling maintains the dormant state through cold storage while preventing desiccation premature dormancy release (from warm storage or delayed planting) produces shoots before root establishment, often causing plant failure.
Rebe in Winterruhe
Winterrebe
dormant plant
resting vine
ruhende Rebe
schlafende Rebe
winter vine
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 75-95 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson Lavee S, May P (1997) Dormancy of grapevine buds, Australian Journal of Grape and Wine Research 3: 31-46
Dormant Vine Role
Syrah in Rhône Valley shows double fertilization during mid-late June Year N+1 at BBCH 65-69, with two sperm cells from discharged tube fusing independently within 2-6 hours post-discharge occurring 26-40 hours post-pollination, one sperm creating diploid 2n zygote with egg and other creating triploid 3n endosperm nucleus with central cell, coordinated embryo and endosperm development initiating seed formation, under warm Mediterranean conditions of 25-29°C temperature, typical of this mid-to-late season cultivar achieving 52-66% double fertilization success producing 1-3 seeds per berry
Double fertilization represents the characteristic angiosperm reproductive process where two independent sperm-gamete fusion events occur simultaneously or sequentially following pollen tube discharge, with one sperm fusing with egg cell forming diploid zygote initiating embryo development and other sperm fusing with central cell forming triploid endosperm nucleus initiating nutritive tissue development, establishing successful sexual reproduction unique to flowering plants. This distinctive dual fusion process occurs during fertilization in mid-late June in temperate regions Year N+1, typically completing within hours following tube discharge approximately 24-48 hours after initial pollination, corresponding to BBCH 65-69 fertilization completion stages. Double fertilization mechanism involves coordinated processes including simultaneous or near-simultaneous release of two sperm cells from discharged pollen tube, independent sperm cell migration to respective fusion targets with one approaching egg and other approaching central cell, membrane fusion events integrating sperm cells into target cell cytoplasm, nuclear fusion combining sperm nuclei with egg nucleus forming 2n zygote and with central cell nucleus or polar nuclei forming 3n endosperm nucleus, and coordinated activation of embryo and endosperm development establishing viable seed. Evolutionary significance shows double fertilization as synapomorphy defining angiosperms distinguishing from gymnosperms having single fertilization, adaptive advantage through coordinated embryo-endosperm development with endosperm nutritive support matched to embryo requirements, and reproductive efficiency through utilizing both sperm cells from single pollen tube maximizing reproductive output. Two sperm cells characteristics show genetically identical haploid cells derived from generative cell mitosis during pollen development or tube growth, cells containing minimal cytoplasm optimized for fusion, non-motile cells requiring passive transport after discharge, and functionally equivalent cells though ultimately fusing with different targets in embryo sac. Sperm delivery timing shows two cells releasing simultaneously from ruptured tube apex into synergid and embryo sac cytoplasm, immediate availability for migration to fusion targets given close proximity within embryo sac, and rapid positioning within minutes to hours enabling prompt fusion events. Egg fertilization as first fusion event shows one sperm cell migrating short distance 20-100 micrometers from discharge site to egg cell at micropylar pole, sperm-egg membrane fusion integrating sperm into egg cytoplasm, nuclear fusion combining haploid sperm nucleus with haploid egg nucleus, formation of diploid 2n zygote nucleus containing equal genetic contributions from maternal and paternal parents, and zygote activation initiating embryo developmental program. Central cell fertilization as second fusion event shows other sperm cell approaching large central cell occupying majority of embryo sac volume, sperm fusion with central cell containing fused diploid nucleus from two polar nuclei or fusing separately with individual polar nuclei, formation of triploid 3n endosperm nucleus with 2 per 1 maternal per paternal genome ratio reflecting two maternal polar nuclei and one paternal sperm nucleus, and endosperm activation initiating nutritive tissue development supporting embryo growth. Fusion timing coordination shows two events typically occurring synchronously or within hours of each other, coordinated timing ensuring embryo and endosperm development initiate together, temporal synchronization critical for normal seed development with embryo requiring endosperm nutritive support from earliest stages, and coordination possibly regulated through common calcium signaling or developmental triggers. Membrane fusion mechanisms show sperm and target cell plasma membranes coming into close contact, lipid bilayer reorganization enabling membrane merger, fusion pore formation creating cytoplasmic continuity, and membrane proteins possibly mediating fusion specificity and efficiency though molecular details in grapevine poorly characterized. Nuclear fusion dynamics show sperm nucleus migrating through target cell cytoplasm toward female nucleus, nuclear envelope breakdown in both gamete nuclei, chromatin mixing and nuclear membrane reformation around combined genetic material, and establishment of diploid or triploid nucleus with integrated parental genomes. Calcium signaling in fertilization shows calcium waves propagating through egg cytoplasm following sperm fusion, localized calcium elevations triggering developmental activation, calcium-dependent processes regulating membrane fusion and nuclear migration, and calcium as key second messenger coordinating fertilization events though specific roles incompletely understood. Egg activation following fertilization shows cortical reaction preventing polyspermy through membrane modifications blocking additional sperm, metabolic activation with respiratory rate increasing and biosynthesis initiating, cell cycle reentry with transition from meiotic arrest to mitotic division, and developmental commitment to embryogenesis with first division within 24-48 hours. Endosperm activation shows free nuclear division initiating rapidly producing multinucleate coenocyte within hours to days, absence of cell wall formation during initial divisions creating syncytial tissue, eventual cellularization forming cellular endosperm, and nutritive function providing resources supporting embryo growth throughout seed development. Genomic imbalance in endosperm shows 2 per 1 maternal per paternal ratio creating unique genomic composition, imbalance affecting gene expression patterns with maternal genome overrepresentation, and imprinting phenomena possibly influencing endosperm development and function though poorly characterized in grapevine. Polyspermy prevention shows mechanisms ensuring single sperm fusion per target with egg showing rapid cortical reaction blocking additional sperm, central cell possibly showing similar prevention though mechanisms less characterized, and ovule-level prevention through single tube reception per ovule eliminating multiple sperm delivery. Fertilization success indicators include formation of functional zygote capable of embryo development, viable endosperm providing nutritive support, coordinated development of both products ensuring seed formation, and subsequent ovary expansion confirming successful fertilization through berry growth initiation. Failed double fertilization scenarios include single fertilization with either egg or central cell fusion succeeding but not both causing developmental abortion, failed membrane fusion preventing nuclear combination despite sperm delivery, abnormal nuclear fusion producing aneuploid or polyploid nuclei causing developmental failure, or complete fertilization failure with neither fusion occurring causing flower abortion. Environmental effects on double fertilization include temperature critically affecting membrane fusion dynamics and developmental activation with optimal 22-28°C supporting efficient processes, calcium availability affecting signaling cascades regulating fusion and activation, and overall favorable conditions during brief fertilization window critical for success. Temperature sensitivity shows optimal conditions supporting rapid efficient fusion and activation within 1-4 hours post-discharge, cool conditions potentially delaying or impairing fusion processes reducing success rate, excessive heat above 33°C causing membrane dysfunction or signaling disruption preventing normal fusion, and critical temperature dependence during fertilization period affecting reproductive outcomes. Developmental coordination following double fertilization shows synchronized embryo and endosperm development with coordinated cell division timing, integrated signaling between embryo and endosperm coordinating growth, and matched developmental rates ensuring appropriate nutritive support throughout embryogenesis. Seed development outcomes show successful double fertilization leading to viable seed formation with embryo developing into seedling and endosperm consumed during maturation, seed containing genetic contributions from both parents through zygote, and functional propagule enabling sexual reproduction and genetic recombination. Berry development initiation following successful double fertilization shows auxin production from developing seeds preventing flower abscission, hormone signaling triggering ovary wall expansion creating berry flesh, and visible berry growth within 1-2 weeks providing evidence of fertilization success. The process shows two distinguishable aspects occurring in parallel with egg fertilization creating embryo lineage and central cell fertilization creating endosperm support tissue. Duration represents brief period with fusion events completing within hours of sperm discharge approximately 24-48 hours post-pollination. Cultivar variation in double fertilization may include fusion efficiency affecting success rates, developmental coordination affecting seed viability, and endosperm-embryo balance affecting seed and berry characteristics. Regional timing shows double fertilization during mid-late June in warm climates, late June in temperate regions, and late June-early July in cool areas, completing 24-48 hours after pollination as culminating reproductive events. Viticultural significance includes representing unique angiosperm reproductive mechanism enabling successful seed formation, requiring coordinated dual fusion events for viable seed development, demonstrating critical environmental sensitivity during fusion period, establishing double fertilization success as direct determinant of seed number and berry development, and marking definitive completion of sexual reproductive sequence initiating developmental phase. Research applications include analyzing molecular mechanisms of membrane and nuclear fusion, investigating coordination between egg and central cell fertilization events, evaluating calcium signaling and other regulatory pathways, quantifying fusion success rates and seed formation outcomes, assessing genomic imbalance effects on endosperm function, and developing strategies for improving fertilization efficiency and fruit set.
Double_Fertilization represents characteristic angiosperm reproductive process distinguishing flowering plants. Subdivided into Egg_Fertilization producing diploid zygote and Central_Cell_Fertilization producing triploid endosperm. Duration hours post-discharge occurring 24-48h post-pollination. Involves coordinated dual fusion events. Creates both embryo and nutritive tissue lineages. Critical temperature sensitivity. Success rate 50-70% determining seed formation. Evolutionary synapomorphy of angiosperms. Enables coordinated embryo-endosperm development. Final reproductive process completing Flowering sequence.
Angiospermen-Befruchtung
Doppelbefruchtung
Zwei-Spermien-Fusion
angiosperm fertilization
coordinated gamete fusion
dual fertilization
duale Fertilisation
two-sperm fusion
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 393-398 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 298-302 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Double Fertilization
A Pinot Noir vine in Nuits-Saint-Georges premier cru trained to guyot double bears a Double_Guyot_Role with two 6-bud canes extending in opposite directions, the double structure managing the vigorous sites while maintaining cane-pruning benefits—a configuration permitted for premier cru parcels with adequate vine age and balance. A Sancerre Sauvignon Blanc vine serves in a Double_Guyot_Role with bilateral cane arrangement, the double-cane approach standard for Loire Sauvignon Blanc's moderate vigor and regional yield expectations. A Riesling vine in Pfalz double guyot training bears a Double_Guyot_Role adapted to the region's fertile soils and higher-vigor conditions compared to steeper Mosel sites where single guyot predominates.
A guyot-trained vine role that inheres in a grapevine with two fruiting canes retained at pruning, extending in opposite directions from the vine head along the fruiting wire, creating bilateral cane structure analogous to bilateral cordon but with annual cane replacement rather than permanent cordons. Double guyot (guyot double) doubles the fruiting capacity compared to single guyot while maintaining the cane-pruning approach suited to varieties with limited basal bud fruitfulness. Each cane typically carries 6-10 buds (12-20 buds total per vine), with cane length adjusted based on vine vigor, variety characteristics, and yield targets. Two renewal spurs are positioned below the fruiting canes, one on each side, providing replacement cane candidates for both directions. Double guyot is preferred for moderate-to-high vigor vines where single-cane crop load would be insufficient to balance vegetative growth, and in regions or appellations permitting higher bud numbers per vine. The symmetrical structure facilitates canopy management and maintains balanced vine appearance, though the increased bud number may reduce per-cluster concentration compared to more restrictive single guyot.
Double_Guyot_Role represents the higher-capacity variant of guyot training, selected when vine vigor or yield targets exceed single-cane capacity. The transition from single to double guyot may reflect vine age (young vines trained to single guyot until established, then converted to double), site fertility (richer soils supporting double-cane vigor), or appellation requirements (some French appellations mandate single guyot for grand cru, permit double for premier cru or village wines). Double guyot pruning is more complex than single, requiring selection and positioning of two canes plus two spurs, with attention to balance between sides. The system creates vineyard appearance similar to bilateral cordon from a distance, but close inspection reveals the cane origin points at the vine head rather than the continuous cordon structure of spur-pruned systems.
Doppelstrecker
Guyot-Doppel
Zweibogen-Erziehung
beidarmiger Guyot
bilateral guyot
double guyot
guyot double
two-cane guyot
Galet P (2000) General Viticulture, Oenoplurimedia Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles INAO (2020) Cahiers des charges des appellations d'origine contrôlées Huglin P, Schneider C (1998) Biologie et écologie de la vigne, Lavoisier
Double Guyot Role
A double vertical cordon on Vitis vinifera cv. Aglianico in tendone system, featuring two vertical cordons extending 205cm upward from bilateral trunk branches spaced 35cm apart at trunk level, each vertical cordon bearing 16 spurs distributed along length, paired vertical structures supporting overhead canopy framework with combined 32 spur positions producing approximately 60 fruiting shoots and 10-12kg total crop distributed between cordons, representing robust double vertical architecture providing structural redundancy (if one cordon damaged or declining, second maintains partial production), balanced capacity distribution, and substantial overhead canopy support from paired vertical columns typical of productive tendone systems in vigorous southern Italian vineyards.
A vertical cordon configuration featuring two upward-extending cordons per vine, characterized by paired vertical axes typically arising from bilateral trunk or head branching, fruiting spurs distributed along both vertical cordons, and architectural redundancy providing capacity distribution between paired vertical elements similar to bilateral horizontal cordons but with vertical rather than horizontal orientation. Double vertical cordons represent enhanced vertical cordon architecture, providing advantages of bilateral capacity distribution (vigor split between paired structures, production insurance if one cordon fails) within vertical orientation framework, suitable for overhead trellis systems requiring multiple vertical support columns, wider-spaced high-density plantings accommodating paired verticals, or situations where vertical architecture is preferred but adequate vine vigor and space permit paired rather than single vertical structure.
Double vertical cordons provide benefits analogous to bilateral horizontal cordons (capacity distribution, redundancy, balanced growth) but within vertical architecture suited to overhead systems or other applications requiring vertical permanent structure. The paired vertical configuration allows substantial total fruiting capacity (combining spur counts from both verticals) while maintaining manageable individual cordon dimensions, provides insurance against single-cordon loss (damage, disease, decline), and creates balanced architectural framework supporting symmetrical canopy development. However, double vertical cordons require adequate vine vigor to support two vertical structures (combined length often 3-4+ meters total wood), appropriate spacing to accommodate paired verticals (typically 0.8-1.2m between verticals), and management attention to maintaining balance between cordons preventing dominance of one vertical over the other through differential vigor or positioning effects.
Doppelhochkordon
doppelte vertikale Kordon
dual vertical cordon
paired vertical cordon
twin-column cordon
Based on Intrieri & Poni (1995) "Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality of mechanized Italian vineyards" American Journal of Enology and Viticulture 46=116-127 Reynolds & Vanden Heuvel (2009) "Influence of grapevine training systems on vine growth and fruit composition= a review" American Journal of Enology and Viticulture 60=251-268.
Double Vertical Cordon
Field evaluation of Downy_Mildew_Resistance_Disposition at INRAE Colmar, France, involves planting replicated plots of test genotypes alongside susceptible controls ('Müller-Thurgau', OIV resistance rating 1) and resistant references ('Regent', OIV rating 7) in a vineyard with naturally high P. viticola pressure. Disease severity is assessed at multiple timepoints using the OIV 452 descriptor, recording percentage leaf area with sporulating lesions and percentage of infected clusters. The resistant breeding selection Gf.Ga-52-42, carrying Rpv3-1 from 'Villard blanc' and Rpv10 from V. amurensis, demonstrates strong Downy_Mildew_Resistance_Disposition with mean leaf infection below 5% and cluster infection below 2% across five seasons, compared to 60-85% leaf infection in 'Müller-Thurgau' controls. This resistance is realized through rapid hypersensitive response visible as small necrotic flecks within 48 hours post-inoculation, preventing sporangiophore emergence and limiting secondary spread.
A Downy_Mildew_Resistance_Disposition is a Fungal_Resistance_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through defense processes upon encounter with the oomycete Plasmopara viticola, the causal agent of grapevine downy mildew. This disposition represents the capacity to limit one or more stages of the P. viticola infection cycle, including zoospore encystment and germination on the leaf surface, stomatal penetration, intercellular hyphal growth, haustorial formation in mesophyll cells, and sporangiophore emergence through stomata. Downy mildew resistance dispositions in Vitis species operate through diverse mechanisms including pre-formed physical barriers (stomatal density and morphology, cuticle composition), constitutive chemical defenses, rapid recognition of pathogen-associated molecular patterns triggering PAMP-triggered immunity, and effector-triggered immunity mediated by resistance genes of the NLR family. The latter mechanism underlies the major resistance loci designated Rpv (Resistance to Plasmopara viticola), with over 30 loci identified to date in various wild Vitis species including V. rupestris, V. riparia, V. amurensis, V. cinerea, and Muscadinia rotundifolia. Downy mildew resistance is of paramount importance for viticulture in humid climates where P. viticola can cause complete crop loss in susceptible cultivars, driving intensive fungicide programs that represent significant economic and environmental costs.
Downy_Mildew_Resistance_Disposition is_a Fungal_Resistance_Disposition. Although Plasmopara viticola is an oomycete rather than a true fungus, resistance to this pathogen is classified under Fungal_Resistance_Disposition following viticultural convention. Child classes distinguish resistance mechanisms: Rpv_Mediated_Resistance_Disposition for major gene resistance, Quantitative_Downy_Mildew_Resistance_Disposition for polygenic resistance, Ontogenic_Downy_Mildew_Resistance_Disposition for age-related resistance, and Induced_Downy_Mildew_Resistance_Disposition for elicitor-triggered enhanced resistance.
DM resistance potential
Falscher Mehltau-Resistenz
Peronospora-Resistenz-Disposition
Plasmopara resistance disposition
Rebenperonospora-Widerstandsfähigkeit
peronospora resistance
Gessler C, Pertot I, Perazzolli M (2011) Plasmopara viticola: a review of knowledge on downy mildew of grapevine and effective disease management, Phytopathologia Mediterranea 50:3-44 Buonassisi D et al. (2017) Breeding for grapevine downy mildew resistance: a review of "omics" approaches, Euphytica 213:103 VIVC (2024) Vitis International Variety Catalogue - Resistance loci database
Downy Mildew Resistance Disposition
Downy mildew symptom quality monitoring in Bordeaux Merlot during a wet spring documented disease progression informing intensive fungicide management. Initial downy mildew symptom quality appeared in late May as scattered oil-spot lesions 1-2 cm diameter showing the characteristic yellow-green translucent appearance on upper leaf surfaces, with morning inspection during high humidity revealing white sporulation on corresponding lower leaf positions confirming P. viticola diagnosis. Disease progression over 2 weeks showed downy mildew symptom quality increasing from 5% to 45% leaf incidence despite calendar-based fungicide applications, with lesions expanding to 3-5 cm and coalescing, and older lesions developing the brown "mosaic" necrotic appearance indicating tissue death following initial biotrophic colonization. Inflorescence downy mildew symptom quality appeared at BBCH 57-61 as gray-brown discoloration and curvature of infected flower clusters, with affected inflorescences failing to set fruit and desiccating to characteristic "shepherd's crook" appearance. Young berry infection showed downy mildew symptom quality as gray-brown soft rot with white sporulation under humid conditions, distinct from the leather berry stage of older infected fruit that becomes resistant to further sporulation. The downy mildew symptom quality progression triggered shift to shortened spray intervals with systemic fungicides, ultimately limiting yield loss to 15% in the most affected blocks.
Downy mildew symptom quality is a fungal infection indicator quality that inheres in grapevine organs infected by the oomycete Plasmopara viticola and represents the characteristic disease manifestations including the yellow-green "oil spot" lesions on upper leaf surfaces corresponding to white cottony sporulation on lower surfaces, the brown "mosaic" or necrotic symptoms on older infections, and the gray-brown shriveling of infected inflorescences and young berries, with symptom expression dependent on tissue age, environmental conditions, and the timing of infection relative to organ development. This quality captures the diagnostic features of one of the most destructive grapevine diseases worldwide, recognizing that downy mildew can cause complete crop loss under favorable conditions and that early symptom recognition is essential for timely fungicide intervention before explosive sporulation enables rapid epidemic development. The ontological significance of downy mildew symptom quality lies in its role enabling disease identification and timing assessment for management decisions, with the distinctive oil-spot appearance providing reliable field diagnosis while symptom progression from initial chlorosis through sporulation to necrosis indicates disease stage and management urgency. The biological basis of downy mildew symptoms encompasses the intercellular growth of P. viticola hyphae within leaf mesophyll causing the initial chlorotic response, the emergence of sporangiophores through stomata on the lower leaf surface producing the characteristic white sporulation visible during humid conditions, and the eventual host cell death causing the brown necrotic appearance of older lesions. The tissue-specific symptoms reflect differential susceptibility, with young actively growing tissues most vulnerable to infection and showing the most dramatic symptom development, while mature tissues show increasing ontogenic resistance limiting both infection success and symptom severity. The environmental dependence of symptom expression creates diagnostic challenges, with sporulation requiring high humidity (greater than 95%) such that the white downy growth diagnostic for the disease may be absent during dry conditions even on infected leaves showing oil-spot chlorosis.
Downy_Mildew_Symptom_Quality_(Plasmopara_viticola) is_a Fungal_Infection_Indicator_Quality, representing the diagnostic symptoms of Plasmopara viticola infection. Note that P. viticola is technically an oomycete rather than true fungus but is traditionally managed as a fungal disease. The oil-spot lesion with corresponding abaxial sporulation is pathognomonic. Young tissues show greatest susceptibility (ontogenic resistance develops). Sporulation requires >95% humidity creating diagnostic challenges in dry conditions. One of the most destructive grapevine diseases requiring intensive management in humid climates.
Falscher Mehltau-Symptome
Peronospora-Symptome
Plasmopara symptoms
Plasmopara-Symptome
downy mildew symptoms
oil spot disease
peronospora symptoms
Ölflecken-Krankheit
Gessler C et al. (2011) Plasmopara viticola: a review of knowledge on downy mildew of grapevine and effective disease management. Phytopathologia Mediterranea 50:3-44 Kennelly MM et al. (2007) Primary infection, lesion productivity, and survival of sporangia in the grapevine downy mildew pathogen Plasmopara viticola. Phytopathology 97:512-522 Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests, 2nd Edition. APS Press
Downy Mildew Symptom Quality (Plasmopara viticola)
The drainage description for the Haut-Médoc classified growth vineyards documents well drained to somewhat excessively drained conditions (drainage class 5-6) — the deep Quaternary gravel deposits provide rapid gravitational drainage through the abundant macropore network between the 10-60 mm gravel clasts, with the water table typically at 2-4 m depth during the growing season and rising to 1-2 m only during winter rainfall. This free-draining condition is the pedological common denominator identified by Seguin across the Médoc First Growths — Châteaux Latour, Lafite, Margaux, and Mouton all occupy the highest, best-drained gravel mounds (croupes), while lesser-quality vineyards in the same communes occupy lower-lying positions with less effective drainage and periodically elevated water tables. The well-drained Médoc gravel soils contrast with the somewhat poorly drained clay soils of parts of the Saint-Émilion plateau (where artificial drainage tiles were installed beginning in the 18th century to improve vine conditions) and the poorly drained alluvial soils of the Palus zone along the Gironde estuary (excluded from quality wine production due to waterlogging risk).
A soil description component that records the soil's capacity to remove excess water from the rooting zone following rainfall or irrigation, classified according to the standard drainage classes from very poorly drained (state 1, water removed so slowly that the soil remains saturated for most of the growing season) through poorly drained, somewhat poorly drained, moderately well drained, well drained, somewhat excessively drained, to excessively drained (state 7, water removed very rapidly with essentially no retention), with drainage being the soil physical property most consistently correlated with vineyard wine quality across all terroir studies — well-drained soils producing higher-quality wines than poorly drained soils in virtually every documented comparison.
Drainage_Description documents the soil hydrological property that Seguin identified as the most reliable predictor of vineyard wine quality — his comprehensive survey of Bordeaux appellations demonstrated that the classified First Growth vineyards consistently occupied the best-drained positions in the landscape, regardless of whether the drainage was achieved naturally (deep gravel deposits in the Médoc, sloping limestone in Saint-Émilion) or artificially (drainage tile installation in heavy clay soils). The mechanism linking drainage to wine quality operates through the soil-vine water balance — well-drained soils remove excess water rapidly after rainfall, preventing waterlogging and root oxygen deficiency, but retain limited plant-available water in the mesoporosity, creating the moderate water deficit during the ripening period that restricts vegetative growth, concentrates berry flavor compounds, and promotes phenolic maturation: poorly drained soils retain excess water for prolonged periods, saturating the root zone with water that displaces soil air (creating anaerobic conditions that inhibit root function and promote root diseases), maintaining high soil moisture availability that sustains vigorous vegetative growth through the ripening period (competing with fruit for assimilates and shading the fruit zone), and delaying soil warming in spring (because water has high heat capacity). The seven standard drainage classes are assessed in the field by observing soil morphological indicators — the depth to grey (gleyed) mottles or horizons indicating the seasonal high water table position, the prominence of oxidized (rust-brown) and reduced (blue-grey) color patterns reflecting alternating aerobic and anaerobic conditions, and the depth to free water observed in augered boreholes during the wet season.
Bodendrainage-Bewertung
Entwässerungsbeschreibung
hydrological soil characterization
hydrologische Boden-Charakterisierung
soil drainage assessment
water removal capacity
Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873 van Leeuwen C et al. (2004) Influence of climate, soil, and cultivar on terroir, American Journal of Enology and Viticulture 55: 207-217 White RE (2003) Soils for Fine Wines, Oxford University Press Soil Science Division Staff (2017) Soil Survey Manual, USDA Handbook 18 van Leeuwen C and Seguin G (2006) The concept of terroir in viticulture, Journal of Wine Research 17: 1-10
Drainage Description
Dried fruit aroma profiling in Amarone della Valpolicella from traditional 120-day appassimento reveals signature raisin character (8.2 on 10-point scale), complemented by fig (6.5), prune (5.8), and dried cherry (5.2), correlating with 42% berry weight loss during drying and sotolon concentration of 85 μg/L, with the dried fruit complexity distinguishing Amarone from Ripasso (shorter drying, dried fruit 4.5) and fresh Valpolicella (minimal dried fruit 2.1), demonstrating how the dried fruit profile directly reflects production methodology.
Dried fruit aroma quality is a fruity aroma quality that represents olfactory perception evoking dehydrated and concentrated fruit-associated scents including raisin, prune, fig, date, dried apricot, and fruitcake notes, characterizing the rich, concentrated, often sweet-associated aromatic impression found in fortified wines, late-harvest wines, wines from dried grapes, and aged wines where fresh fruit has evolved toward preserved fruit character. Within the BFO framework, this quality addresses a distinctive subcategory of fruity aroma associated with concentration processes (grape drying, botrytis infection, extended aging), distinguishing dried fruit character from fresh fruit while recognizing its positive contribution in appropriate wine styles and potential indication of over-ripeness or oxidation in styles where fresh fruit is expected. The ontological significance lies in dried fruit aroma serving as both a quality marker in specific wine styles (raisin in Amarone, fig in vintage Port, dried apricot in Sauternes) and a developmental indicator where fresh fruit transitions toward dried fruit character during aging, providing information about wine evolution and optimal drinking window. The compositional basis for dried fruit aroma includes furaneol and sotolon (contributing caramelized, dried fruit character), specific lactones associated with dried fruit, oxidation-derived aldehydes (methional, phenylacetaldehyde), and concentration of glycosidically-bound aroma precursors released during extended aging. The production pathways to dried fruit character include grape drying (appassimento for Amarone, passerillage for vin de paille), botrytis concentration (noble rot creating dried apricot character), fortification (arrested fermentation preserving grape sugar in dried fruit-noted form), and oxidative aging (sherry, tawny Port developing raisin and fig notes). The context-dependency of dried fruit quality assessment is critical, as these notes indicate quality in appropriate styles but may signal defect (raisining, premature oxidation) in wines intended to show fresh fruit. Understanding dried fruit aroma quality enables style identification based on characteristic dried fruit profile, aging assessment from dried fruit development, and quality evaluation where appropriate dried fruit expression indicates successful production of concentrated wine styles.
Dried_Fruit_Aroma_Quality addresses the concentrated and preserved fruit subcategory of fruity aroma, encompassing raisin, prune, fig, date, and dried apricot notes. This quality is style-dependent, indicating quality in appropriate wine types (fortified, late-harvest, dried grape wines) while potentially signaling defect in fresh wine styles. The class supports style identification, aging assessment, and production methodology evaluation based on dried fruit expression.
Dörrfruchtcharakter
Rosinen-Duft
Trockenfruchtnoten
Trockenobstaroma
concentrated fruit scent
dried fruit notes
preserved fruit character
raisin aroma
Ferreira V et al. (2003) Quantitative determination of important odorants in prunes. Journal of Agricultural and Food Chemistry 51(8): 2463-2469 Urlaub R, Coulon D (2004) The aroma compounds of dried vine fruit Recioto della Valpolicella Amarone. In: Ferreira V, López R (eds) State-of-the-Art in Flavour Chemistry and Biology. Deutsche Forschungsanstalt für Lebensmittelchemie, Munich, pp. 301-305
Dried Fruit Aroma Quality
Cabernet Sauvignon vines in the Napa Valley bearing a drip-irrigated role receive water through pressure-compensating emitters at 2-4 L/hour spaced at 60-90 cm intervals along polyethylene laterals, delivering 0.8-1.5 ML/ha/season in 2-4 hour irrigation sets scheduled 1-3 times weekly based on ET₀ calculations from on-site weather stations, achieving application uniformity coefficients above 90% with negligible deep percolation losses compared to 30-50% losses typical of flood systems.
An irrigated role that inheres in a grapevine receiving supplemental water through a pressurized drip irrigation system that delivers water at low flow rates directly to the root zone via emitters positioned along or below the vine row, enabling precise control of water volume, timing, and spatial distribution.
Drip_Irrigated_Role represents the dominant precision irrigation technology in modern viticulture, adopted across approximately 70-80% of irrigated vineyard area in major New World producing regions. The technology enables fertigation (fertilizer delivery through the irrigation system), partial root zone drying protocols, and highly targeted deficit irrigation strategies not achievable with flood systems: drip systems also minimize leaf wetness and associated fungal disease pressure compared to overhead sprinkler methods. Subsurface drip irrigation (SDI) with emitter lines buried 30-50 cm below the surface represents a further refinement reducing surface evaporation losses.
Mikrobewässerungs-Produktionsrolle
Tröpfchenbewässerungs-Rolle
drip-irrigated vine role
micro-irrigated production role
trickle-irrigated role
tropfbewässerte Rolle
Netzer Y et al. (2014) Drip irrigation scheduling in grapevines, Irrigation Science 32: 41-53 Intrigliolo DS and Castel JR (2010) Response of grapevine to irrigation, Agricultural Water Management 97: 1-6 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 502-510
Drip Irrigated Role
The Dryness Index (DI) for Barossa Valley typically ranges from -150 to -200 mm, classifying it as "very dry" (DI < -100) under the Géoviticulture MCC system, confirming the necessity of deficit irrigation strategies for Shiraz production. The Palmer Drought Severity Index (PDSI) for the Douro Valley during the exceptional 2017 vintage reached -4.5 (extreme drought), contributing to historically low yields but extraordinary Port wine concentration. Standardized Precipitation Index (SPI) values of -1.5 to -2.0 for Mendoza's 2020-2021 growing season indicated severe precipitation deficit, mitigated by the snowmelt-fed irrigation infrastructure.
Drought index is a precipitation data entity that records a calculated numerical indicator quantifying the severity, duration, and frequency of water deficit conditions at a viticultural site, derived from precipitation, temperature, evapotranspiration, and/or soil moisture parameters through standardized meteorological formulae, enabling comparative assessment of water stress potential across sites, vintages, and climate scenarios.
Multiple drought indices exist, each capturing different aspects of water deficit relevant to viticulture. The Géoviticulture MCC system employs the Dryness Index (DI = P - ETP for April-September) as one of three bioclimatic indices for global viticultural zoning, with thresholds at DI > 150 (humid), 150 to -100 (sub-humid), -100 to -200 (moderately dry), and < -200 (very dry). The Standardized Precipitation Index (SPI) and Palmer Drought Severity Index (PDSI) are more widely used in general climatology but require adaptation for viticultural interpretation. Emerging vine-specific drought indices incorporate predawn leaf water potential thresholds (Ψ_pd), soil water balance modeling, and remote sensing-derived crop water stress indices (CWSI) to provide more direct assessment of vine water status. Under climate change scenarios, drought index projections suggest significant increases in water deficit for many established viticultural regions, particularly in southern Europe and Australia.
Ariditätsindex
Dürreindex
Trockenheitsindex
Trockenheitskenngröße
Wassermangelindex
aridity index
drought severity index
dryness index
vineyard drought metric
water deficit index
Tonietto J, Carbonneau A (2004) A multicriteria climatic classification system for grape-growing regions worldwide. Agricultural and Forest Meteorology 124(1-2): 81-97 Van Leeuwen C et al. (2009) Why climate change will not dramatically decrease viticultural suitability in main wine-producing areas by 2050. Proceedings of the National Academy of Sciences 106(38): 16713-16718 Fraga H et al. (2016) Modelling climate change impacts on viticultural yield, phenology, and grape composition in Europe. Environmental Research Letters 11(4): 044007
Drought Index
Systematic evaluation of Drought_Recovery_Disposition at INRAE Bordeaux imposed standardized drought stress (predawn water potential of -1.0 MPa for 14 days) followed by rewatering on 20 cultivars, with recovery tracked over 21 days through measurements of water potential, stomatal conductance, photosynthesis, stem hydraulic conductivity, and shoot growth. Recovery was quantified as percentage of pre-stress function restored at each timepoint. Drought_Recovery_Disposition varied substantially among cultivars. 'Grenache' (isohydric) showed rapid, complete recovery with predawn water potential restored within 24 hours, stomatal conductance reaching 90% of pre-stress levels by day 3, and photosynthesis fully recovered by day 5, with no persistent hydraulic damage (PLC < 5% throughout). 'Syrah' (anisohydric) displayed slower, less complete recovery with water potential restored by 48 hours, but stomatal conductance reaching only 75% by day 7 and photosynthesis 80% by day 14, associated with 25% PLC requiring gradual embolism repair. 'Merlot' showed intermediate recovery but with persistent effects, achieving 85% stomatal recovery by day 7 but maintaining elevated ABA levels suggesting continued stress memory. Growth resumption analysis revealed that shoot elongation recommenced within 5 days in 'Grenache' versus 12 days in 'Syrah', with cumulative 21-day recovery growth 40% higher in 'Grenache'. These findings demonstrate that Drought_Recovery_Disposition reflects the integration of multiple recovery processes with different kinetics and potential for incomplete restoration.
A Drought_Recovery_Disposition is a Drought_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through physiological mechanisms that enable rapid restoration of normal function following relief of water stress, encompassing rehydration of tissues, repair of drought-induced damage, and resumption of growth and productive processes. This disposition represents the restorative component of drought adaptation, determining how quickly and completely vines return to pre-stress function after water availability improves through rainfall, irrigation, or reduced evaporative demand. Drought recovery involves multiple processes operating at different timescales, from rapid stomatal reopening (minutes to hours), through tissue rehydration and turgor restoration (hours to days), repair of cavitation damage and cellular injury (days to weeks), and growth resumption and reserve replenishment (weeks to months). The completeness and speed of recovery depend on drought severity and duration (more severe stress causing more damage requiring more recovery time), cultivar physiology (stomatal behavior, hydraulic properties, cellular protection mechanisms), and environmental conditions during recovery (temperature, light, water availability). Drought_Recovery_Disposition is economically important because incomplete recovery can reduce productivity in the current season (delayed canopy development, reduced fruit quality) and subsequent seasons (depleted reserves, persistent hydraulic damage), making recovery capacity a key component of sustainable viticulture in drought-prone regions.
Drought_Recovery_Disposition is_a Drought_Tolerance_Disposition. Recovery encompasses multiple processes at different timescales, from rapid stomatal reopening to slow hydraulic repair. Child classes include Rehydration_Capacity_Disposition (tissue water restoration) and Growth_Resumption_Disposition (developmental recovery). Recovery completeness depends on drought severity and duration as well as genotype. Isohydric cultivars typically show faster, more complete recovery due to less hydraulic damage during stress. Incomplete recovery can have persistent effects on current and subsequent season productivity.
Post-Stress-Wiederherstellung
Trockenheitserholungs-Disposition
Wiederbewässerungs-Antwort
post-drought restoration
rewatering response
stress recovery capacity
Flexas J et al. (2006) Drought-inhibition of photosynthesis in C3 plants: stomatal and non-stomatal limitations revisited, Annals of Botany 98:871-892 Galmes J et al. (2007) Water relations and stomatal characteristics of Mediterranean plants with different growth forms and leaf habits: responses to water stress and recovery, Plant and Soil 290:139-155 Pou A et al. (2012) Anisohydric behaviour in grapevines results in better performance under moderate water stress and recovery than isohydric behaviour, Plant and Soil 359:335-349
Drought Recovery Disposition
Drought recovery quality evaluation in Napa Valley Cabernet Sauvignon compared the post-stress restoration of function following controlled moderate versus severe stress episodes imposed by irrigation withholding. Following moderate stress (midday stem water potential -1.2 MPa), vines demonstrated excellent drought recovery quality with predawn water potential returning to pre-stress levels within 24 hours of irrigation, stomatal conductance recovering to 80% of unstressed values by 48 hours and 95% by 72 hours, and photosynthesis reaching 90% of maximum within 3 days. In contrast, following severe stress (-1.6 MPa), the same vines showed compromised drought recovery quality with incomplete hydraulic recovery evidenced by predawn potential remaining 0.15 MPa below pre-stress baseline even after 7 days, stomatal conductance recovering to only 70% of unstressed values suggesting persistent non-stomatal limitation, and visible tip desiccation and basal leaf senescence representing permanent tissue loss not recoverable through rehydration. The drought recovery quality differential established -1.4 MPa as the approximate threshold separating fully recoverable moderate stress from partially recoverable severe stress in this cultivar-rootstock combination, informing irrigation management targets ensuring stress episodes remained within the fully recoverable range.
Drought recovery quality is a drought stress indicator quality that inheres in a grapevine organism and represents the capacity to restore normal physiological function, resume growth, and repair stress-induced damage following relief of water deficit through irrigation or precipitation, encompassing the speed of rehydration, the completeness of recovery of photosynthetic and hydraulic function, the restoration of growth processes, and the repair of any stress-induced cellular or tissue damage that may persist beyond the immediate stress period. This quality captures the resilience dimension of drought response that complements tolerance during stress, recognizing that the cumulative impact of drought episodes depends not only on performance during stress but also on the speed and completeness of recovery that determines how rapidly normal function returns and whether lasting damage accumulates across repeated stress cycles. The ontological significance of drought recovery quality lies in its role determining the net productivity and health consequences of drought episodes under the fluctuating water availability conditions typical of many viticultural environments, where recovery between stress episodes may be as important as tolerance during stress for overall seasonal performance. The biological basis of drought recovery encompasses the restoration of tissue water content and turgor as soil water becomes available, the reopening of stomata and recovery of photosynthetic function, the repair of any stress-induced damage to the hydraulic pathway including refilling of embolized xylem vessels, and the resumption of growth processes that were suspended during the stress period. The rate and completeness of recovery vary substantially among grapevine genotypes, with some cultivars showing rapid, complete recovery within days while others exhibit slow recovery with persistent impairment that may require weeks or new tissue development to fully overcome. The interaction between stress severity and recovery capacity creates a threshold phenomenon wherein moderate stress permits rapid, complete recovery while severe stress exceeding critical thresholds causes lasting damage that recovery processes cannot fully repair.
Drought_Recovery_Quality is_a Drought_Stress_Indicator_Quality, serving as parent class for Rehydration_Rate_Quality, Post_Drought_Growth_Resumption_Quality, and Hydraulic_Recovery_Quality which address specific dimensions of the recovery process. This class addresses the overall recovery capacity while child classes capture particular aspects. The stress severity threshold separating full from partial recovery represents critical management information. Recovery may be complete, partial, or negligible depending on stress intensity and duration. Critical for understanding cumulative drought impacts under fluctuating water availability.
Dürre-Erholungsqualität
Nach-Dürre-Wiederherstellung
Rehydratationsantwort
Stresserholungskapazität
drought resilience
rehydration response
Flexas J et al. (2006) Keeping a positive carbon balance under adverse conditions: responses of photosynthesis and respiration to water stress. Physiologia Plantarum 127:343-352 Lovisolo C et al. (2010) Drought-induced changes in development and function of grapevine (Vitis spp.) organs and in their hydraulic and non-hydraulic interactions at the whole-plant level: a physiological and molecular update. Functional Plant Biology 37:98-116 Hochberg U et al. (2017) Stomatal closure, basal leaf embolism, and shedding protect the hydraulic integrity of grape stems. Plant Physiology 174:764-775
Drought Recovery Quality
Drought stress indicator quality evaluation in Mendoza established the diagnostic value of multiple water status assessment approaches for Malbec deficit irrigation management. Predawn leaf water potential demonstrated excellent drought stress indicator quality as an integrative measure of vine water status equilibrated with soil, with values correlating strongly (r²=0.89) with soil moisture content and providing reliable differentiation of stress categories with distinct physiological and quality implications. Midday stem water potential showed superior drought stress indicator quality for detecting current-day stress intensity with values responding within hours to irrigation or stress onset, though requiring greater measurement effort. Trunk diameter variation measured by dendrometers provided continuous automated monitoring with good indicator quality for tracking stress development, though with calibration requirements specific to trunk size and growth stage. Visual assessment using standardized symptom scoring showed moderate drought stress indicator quality adequate for identifying severe stress requiring intervention but insufficient for the precise management of mild-to-moderate stress targeting quality optimization, establishing multi-indicator protocols combining practical visual assessment with periodic water potential measurement as the regional standard for deficit irrigation management.
Drought stress indicator quality is a stress response quality that inheres in a grapevine organism and represents the ensemble of physiological, morphological, and biochemical markers that signal the presence and severity of water deficit conditions, providing diagnostic information for assessing vine water status and guiding irrigation management decisions to optimize the balance between water conservation, vine health maintenance, and fruit quality enhancement achievable through controlled water stress application. This quality encompasses both the sensitivity and reliability of various stress indicators as diagnostic tools, recognizing that effective drought stress management requires accurate assessment of vine water status that different indicators provide with varying precision, timing, and practical accessibility. The ontological significance of drought stress indicator quality lies in its role enabling evidence-based irrigation management through objective assessment of vine water status, replacing calendar-based or intuitive watering decisions with responsive protocols that achieve target stress levels supporting specific production objectives from maximum yield through premium quality-focused deficit strategies. The biological basis of drought stress indicators encompasses the progressive physiological responses to declining soil and plant water status, beginning with subtle changes in stomatal conductance and shoot extension rate at incipient stress, progressing through measurable changes in leaf water potential, stem water potential, and trunk diameter as stress intensifies, and ultimately manifesting as visible symptoms including leaf wilting, tendril desiccation, and basal leaf senescence under severe deficit. The temporal sequence of indicator response provides diagnostic information, with highly sensitive indicators including stem water potential and stomatal conductance detecting stress onset before visible symptoms appear, while lagging indicators including growth cessation and leaf yellowing confirm sustained stress that may already have impacted vine function and fruit development. The selection of appropriate drought stress indicators depends on production objectives, available monitoring technology, and the precision required for management decisions, with research-grade measurements including pressure chamber water potential providing high accuracy while practical field indicators including visual scoring and dendrometer trunk shrinkage offer accessibility advantages for routine monitoring.
Drought_Stress_Indicator_Quality is_a Stress_Response_Quality, serving as parent class for Water_Deficit_Severity_Quality which addresses the categorization of stress intensity levels. This class addresses the diagnostic markers and assessment approaches for water stress rather than the severity classification per se. Distinguished from irrigation management qualities by focusing on stress assessment rather than water application strategies. The hierarchy of indicator sensitivity and practical accessibility informs monitoring protocol design. Critical for implementing regulated deficit irrigation and other water management strategies targeting specific stress levels for quality optimization.
Dürresymptome
Rebenwasserstatus-Indikatoren
Trockenstress-Indikatoren
Wasserstress-Marker
drought symptoms
vine water status indicators
water deficit markers
water stress indicators
Choné X et al. (2001) Stem water potential is a sensitive indicator of grapevine water status. Annals of Botany 87:477-483 Williams LE, Araujo FJ (2002) Correlations among predawn leaf, midday leaf, and midday stem water potential and their correlations with other measures of soil and plant water status in Vitis vinifera. Journal of the American Society for Horticultural Science 127:448-454 Intrigliolo DS, Castel JR (2006) Vine and soil-based measures of water status in a Tempranillo vineyard. Vitis 45:157-163
Drought Stress Indicator Quality
Drought symptom expression quality assessment in Mendoza established visual scoring protocols for Malbec water status monitoring across 200 hectares of vineyard blocks with variable irrigation infrastructure. A five-point symptom severity scale was developed integrating tendril turgor, shoot tip condition, leaf angle, and basal leaf status into composite scores ranging from 1 (no visible stress) through 3 (moderate stress with wilting and tendril drooping) to 5 (severe stress with extensive leaf damage). Weekly scoring by trained vineyard staff achieved strong correlation (r²=0.78) with periodic predawn water potential measurements, validating drought symptom expression quality assessment as a practical monitoring tool. Spatial mapping of symptom scores identified irrigation system deficiencies causing localized severe stress and guided targeted infrastructure repairs. Management thresholds were established with symptom score 2.5 triggering irrigation scheduling review and score 3.5 requiring immediate irrigation intervention regardless of scheduled timing, demonstrating practical application of drought symptom expression quality assessment for large-scale vineyard management.
Drought symptom expression quality is a drought stress indicator quality that inheres in a grapevine organism and represents the visible morphological manifestations of water deficit stress that develop as declining plant water status exceeds the capacity for physiological compensation, providing field-observable indicators of stress severity that complement instrumental measurements and enable rapid assessment of vine water status across vineyard blocks without requiring specialized equipment or laboratory analysis. This quality encompasses the progression of visual symptoms from subtle early indicators detectable only by trained observers through pronounced symptoms visible to any observer, providing a practical diagnostic framework for stress assessment that integrates multiple observable changes into coherent severity interpretation. The ontological significance of drought symptom expression quality lies in its practical value for vineyard monitoring where the accessibility of visual assessment enables frequent, spatially comprehensive stress evaluation that instrumental methods cannot economically achieve, while recognizing that visual symptoms represent lagging indicators appearing after physiological stress responses have already altered vine function and potentially influenced fruit quality development. The biological basis of drought symptom expression encompasses the progressive failure of turgor maintenance in various plant organs as water potential declines, with different tissues exhibiting symptoms at characteristic stress thresholds based on their osmotic adjustment capacity, cell wall properties, and position within the plant hydraulic architecture. The sequence of symptom appearance provides diagnostic information, with highly sensitive indicators including tendril drooping and shoot tip desiccation appearing at mild-to-moderate stress levels, followed by leaf wilting and rolling at moderate stress, and culminating in leaf chlorosis, necrosis, and abscission under severe prolonged stress. Cultivar differences in drought symptom expression reflect variation in osmotic adjustment capacity, cuticular properties, and hydraulic architecture that determine how readily water deficit translates to visible turgor loss, with some cultivars maintaining turgid appearance despite significant physiological stress while others display pronounced symptoms at relatively mild water deficits. Assessment of drought symptom expression quality employs standardized visual scoring systems rating symptom severity on defined scales, enabling semi-quantitative stress assessment suitable for management decision support.
Drought_Symptom_Expression_Quality is_a Drought_Stress_Indicator_Quality, serving as parent class for specific symptom types including Leaf_Wilting_Quality, Leaf_Rolling_Quality, Tendril_Drooping_Quality, and Shoot_Tip_Desiccation_Quality. This class addresses the visible manifestations of water stress while child classes capture particular symptom characteristics. Distinguished from physiological stress indicators by the focus on observable morphological changes. The lagging nature of visual symptoms relative to physiological responses represents important limitation for time-sensitive management decisions. Practical accessibility advantages visual assessment for routine monitoring across large areas.
Trockenstress-Symptomausprägung
Wasserstress-Symptome
drought morphological indicators
sichtbare Stressexpression
visible stress expression
visual drought symptoms
visuelle Dürresymptome
water stress symptoms
Flexas J et al. (2002) Effects of drought on photosynthesis in grapevines under field conditions: an evaluation of stomatal and mesophyll limitations. Functional Plant Biology 29:461-471 Lovisolo C et al. (2010) Drought-induced changes in development and function of grapevine (Vitis spp.) organs and in their hydraulic and non-hydraulic interactions at the whole-plant level: a physiological and molecular update. Functional Plant Biology 37:98-116 Cifre J et al. (2005) Physiological tools for irrigation scheduling in grapevine (Vitis vinifera L.): an open gate to improve water-use efficiency? Agriculture, Ecosystems and Environment 106:159-170
Drought Symptom Expression Quality
Comprehensive characterization of Drought_Tolerance_Disposition at INRAE Montpellier compared physiological responses to progressive soil drying across 25 cultivars in a lysimeter experiment with precise water deficit control. Measurements included predawn and midday leaf water potential, stomatal conductance, photosynthesis, and osmotic potential at full turgor across a range of soil water availability from well-watered to severe stress (predawn water potential -0.2 to -1.2 MPa). Cultivars showed distinct Drought_Tolerance_Disposition strategies. 'Grenache' exhibited isohydric drought tolerance with early stomatal closure (conductance reduced 50% at predawn potential of -0.4 MPa), strict midday water potential regulation (maintained above -1.4 MPa), and minimal osmotic adjustment (0.15 MPa). 'Syrah' displayed anisohydric behavior with sustained stomatal opening (50% reduction only at -0.8 MPa predawn), progressive midday water potential decline to -1.9 MPa under severe stress, and moderate osmotic adjustment (0.35 MPa). 'Sangiovese' showed intermediate characteristics. Under severe stress, 'Grenache' maintained higher leaf relative water content (78% versus 62%) but accumulated less carbon (30% photosynthesis reduction versus 15% in 'Syrah' at moderate stress). Recovery following rewatering was faster in 'Grenache' (full stomatal recovery within 24 hours) compared to 'Syrah' (72 hours to full recovery, with some persistent embolism). These findings demonstrate that Drought_Tolerance_Disposition involves trade-offs between water conservation and carbon gain, with optimal strategy depending on drought severity and duration.
A Drought_Tolerance_Disposition is an Abiotic_Stress_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through physiological, biochemical, and morphological mechanisms that enable maintenance of cellular function, growth, and productivity under soil water deficit conditions that would otherwise cause wilting, metabolic disruption, and yield loss. This disposition represents the capacity to tolerate water stress, critically important for viticulture as grapevines are cultivated across diverse climates ranging from humid to semi-arid, and increasingly relevant under climate change scenarios projecting reduced precipitation and increased drought frequency in many wine regions. Drought tolerance in grapevine involves multiple mechanisms including drought avoidance (deep rooting accessing subsoil water, early phenology escaping late-season drought), dehydration postponement (stomatal regulation conserving water, osmotic adjustment maintaining turgor), and dehydration tolerance (cellular protection mechanisms enabling function at low water potential). The strategic balance between water conservation and carbon gain varies among cultivars, with isohydric types prioritizing water conservation through early stomatal closure while anisohydric types maintain gas exchange longer under stress at the cost of greater dehydration risk. Drought_Tolerance_Disposition encompasses both constitutive traits (root architecture, stomatal density, cuticular properties) and inducible responses (stomatal closure, osmotic adjustment, protective protein accumulation). Understanding genotypic variation in drought tolerance informs cultivar selection for dry climates, rootstock choice for water-limited sites, and irrigation management strategies balancing vine stress with fruit quality optimization.
Drought_Tolerance_Disposition is_a Abiotic_Stress_Tolerance_Disposition. Drought tolerance encompasses multiple mechanisms with different costs and benefits, addressed through child classes including Stomatal_Regulation_Disposition and Osmotic_Adjustment_Disposition. The isohydric-anisohydric spectrum represents a fundamental strategic axis in drought response, with neither extreme optimal for all conditions. Drought tolerance interacts with heat tolerance through shared mechanisms (stomatal behavior, membrane stability) and co-occurrence of stresses. Rootstock genotype strongly influences scion drought tolerance through root system properties and hydraulic conductance.
Dürreresistenz
Trockentoleranz-Disposition
Wassermangel-Toleranz
drought resistance
water deficit tolerance
water stress tolerance
Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data, Annals of Botany 105:661-676 Lovisolo C et al. (2010) Drought-induced changes in development and function of grapevine (Vitis spp.) organs and in their hydraulic and non-hydraulic interactions at the whole-plant level: a physiological and molecular update, Functional Plant Biology 37:98-116 Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought, Plant Cell and Environment 26:1393-1405
Drought Tolerance Disposition
110 Richter (V. berlandieri × V. rupestris) demonstrates high drought tolerance with root penetration exceeding 4 m depth on permeable substrates, maintaining Tempranillo yields of 5-7 t/ha under 300-400 mm annual precipitation in non-irrigated Ribera del Duero vineyards where shallow-rooted rootstocks fail. 140 Ruggeri (V. berlandieri × V. rupestris) shows exceptional drought tolerance in Sicily, supporting Nero d'Avola production on hillside sites receiving less than 350 mm rainfall with summer drought periods exceeding 4 months. 1103 Paulsen (V. berlandieri × V. rupestris) provides drought tolerance to Aglianico in Campania's volcanic tuff soils, maintaining predawn leaf water potentials above -0.4 MPa when own-rooted vines decline below -0.8 MPa.
An abiotic stress-tolerant rootstock role that inheres in a grafted rootstock by virtue of its capacity to maintain scion productivity and survival under water-limited conditions. Drought-tolerant rootstocks sustain vine function during soil water deficit through multiple physiological mechanisms including deep and extensive root system architecture (rooting depth exceeding 3-4 m, root density of 1.5-3.0 km/m³ in deep soil horizons), high root hydraulic conductance under declining soil water potential, efficient stomatal regulation coordinated with root-sourced ABA signaling, osmotic adjustment in root cells maintaining turgor and water uptake at soil water potentials below -1.5 MPa, and xylem anatomy featuring smaller vessel diameters (typically 30-50 µm) that resist cavitation under high tension. Drought tolerance is quantified by the soil water potential threshold at which stomatal closure occurs (drought-tolerant rootstocks maintaining stomatal conductance to -1.5 to -2.0 MPa versus -0.8 to -1.2 MPa for drought-sensitive genotypes), by predawn leaf water potential maintenance, and by yield stability across wet and dry seasons.
Drought_Tolerant_Rootstock_Role is subclassed into High_Drought_Tolerance_Rootstock_Role and Moderate_Drought_Tolerance_Rootstock_Role, reflecting the continuum of drought adaptation among rootstock genotypes. The V. berlandieri × V. rupestris parentage combination (110R, 140Ru, 1103P, 99R) consistently produces drought-tolerant progeny, as both parental species evolved in semi-arid limestone habitats of central Texas (V. berlandieri) and the Ozark Plateau (V. rupestris). Climate change projections of increasing drought frequency and severity are driving renewed breeding interest in drought tolerance as a primary rootstock selection criterion.
Trockenlagen-Unterlage
drought resistant rootstock
drought tolerant rootstock
dry-farming rootstock
trockenheitsresistente Unterlage
trockenheitstolerante Unterlage
wasserstresstolerante Unterlage
water stress tolerant rootstock
Carbonneau A (1985) The early selection of grapevine rootstocks for resistance to drought conditions, American Journal of Enology and Viticulture 36: 195-198 Serra I et al. (2014) The interaction between rootstocks and cultivars to enhance drought tolerance in grapevine, Australian Journal of Grape and Wine Research 20: 1-14 Alsina MM et al. (2011) Seasonal changes of whole root system conductance by a drought-tolerant grape root system, Journal of Experimental Botany 62: 99-109
Drought Tolerant Rootstock Role
Zinfandel vines in Sonoma County's Dry Creek Valley bearing a dry-farmed role develop root systems reaching 5-8 m depth in deep alluvial soils, maintain predawn leaf water potentials of -0.3 to -0.6 MPa during July-August without supplemental water, and produce concentrated fruit at 4-7 t/ha with berry weights of 1.0-1.4 g compared to 1.6-2.0 g for irrigated counterparts, yielding wines with characteristically higher phenolic concentration and flavor intensity valued at premium price points.
A production system role that inheres in a grapevine cultivated exclusively with natural precipitation and stored soil moisture without any supplemental irrigation, relying on deep root development, adapted cultivar selection, and soil moisture conservation techniques to sustain vine productivity through seasonal drought periods.
Dry_Farmed_Role is the historical default for European viticulture where irrigation was traditionally prohibited in many appellations (and remains restricted in most French AOC and many Italian DOC regulations). The role is increasingly recognized as a quality-differentiation strategy in New World regions where irrigation is otherwise standard practice: organizations such as the Dry Farming Institute actively promote dry farming as a climate-resilient and terroir-expressive approach. The role is mutually exclusive with Irrigated_Role and its subclasses. Minimum annual precipitation thresholds for viable dry farming vary from 250-400 mm depending on soil water-holding capacity, evapotranspiration rates, and cultivar drought tolerance.
Regenfeldbau-Rolle
Trockenanbau-Rolle
dry-farmed vine role
non-irrigated production role
rainfed viticulture role
unbewässerte Produktionsrolle
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 469-498 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data, Annals of Botany 105: 661-676 Smart RE and Robinson M (1991) Sunlight into Wine, Winetitles pp: 98-112
Dry Farmed Role
Growth analysis in virus-infected versus healthy Cabernet Franc vines utilizes seasonal dry mass accumulation to quantify the impact of Grapevine Leafroll-associated Virus 3 (GLRaV-3) on vine productivity, revealing reductions of 25-40% in total vine dry mass accumulation in infected vines due to impaired photosynthesis, reduced leaf area development, and altered carbon partitioning, with berry dry mass at harvest reduced by 15-30% contributing to lower yield and delayed ripening.
Dry_Mass_Quality represents the mass of a grapevine organ after removal of all free and bound water through controlled drying, reflecting the accumulated structural compounds, storage reserves, and metabolites exclusive of water. This quality provides a stable measurement basis unaffected by hydration status, enabling rigorous comparison of growth, productivity, and composition across varying environmental conditions and measurement times. Dry mass is the preferred basis for expressing concentrations in plant physiology and biochemistry because it normalizes for water content variation. Organ dry mass accumulates through the growing season as photosynthetically fixed carbon is incorporated into structural tissues (cellulose, lignin, proteins) and storage compounds (starch, sugars, lipids, minerals). Berry dry mass increases continuously from fruit set through harvest, even during late ripening when fresh mass may plateau or decline due to water loss. Cluster dry mass relative to vegetative dry mass indicates the partitioning of photoassimilates between reproductive and vegetative sinks. Dry mass determination requires drying organs to constant mass, typically achieved at 60-80°C for 48-96 hours in forced-air ovens, followed by weighing on analytical balances. Higher temperatures accelerate drying but may volatilize some organic compounds. Freeze-drying (lyophilization) preserves volatile compounds for subsequent chemical analysis but requires longer processing times. The ratio of dry mass to fresh mass, expressed as dry matter percentage or dry matter content, typically ranges from 15-25% for ripe berries depending on sugar concentration.
Dry_Mass_Quality provides the stable mass measurement unaffected by organ water status. Drying protocols must be standardized and reported (temperature, duration, apparatus) because residual moisture content depends on drying conditions. Some dry mass determination protocols distinguish structural dry mass (residue after extraction of soluble compounds) from total dry mass, relevant for carbon balance studies. Dry mass is the denominator for specific leaf area (SLA = leaf area / leaf dry mass), a key ecophysiological trait. For chemical analysis, concentration data expressed per unit dry mass (mg/g DM) enables comparison across samples with varying moisture content.
Trockengewicht
Trockenmasse
Trockensubstanz
dry matter mass
dry weight
oven-dried mass
wasserfreie Masse
water-free mass
Keller M et al. (2015) Warm Spring Temperatures Induce Persistent Season-Long Changes in Shoot Development in Grapevines. Annals of Botany 116(3): 399-408 Ollat N et al. (2002) Organic Acid Metabolism in Roots of Various Grapevine (Vitis) Rootstocks Submitted to Iron Deficiency and Bicarbonate Nutrition. Journal of Plant Nutrition 25(8): 1649-1665
Dry Mass Quality
Chardonnay hermaphrodite flowers at BBCH 57-59 (flower buds swollen, pre-anthesis) exhibit pre-receptive dry stigmas with matte appearance, no visible exudate, and papillae in non-secretory state, achieving <12% pollen germination if pollen is artificially applied at this stage, demonstrating that dry pre-receptive stigmas cannot support effective pollination and that calyptra shedding and receptivity activation must occur before successful pollination is possible. Merlot flowers at 7-9 days post-calyptra shedding show post-receptive dry stigmas with dry matte appearance, no wet exudate (compared to glistening wet appearance during days 1-6), and often brownish discoloration as papillae undergo senescence, achieving <8% pollen germination and <5% fruit set when pollen is applied at this late stage, demonstrating that stigmas have exited the receptivity window and pollination is no longer effective. Syrah rare functionally male flowers exhibit non-receptive dry stigmas throughout the bloom period with consistently dry matte appearance, minimal or absent exudate production even at expected peak receptivity timing (days 1-3 post-calyptra shedding when hermaphrodite stigmas are maximally receptive), achieving <15% pollen germination and <8% fruit set despite adequate pollen availability, with reproductive failure primarily due to stigma non-receptivity rather than to stamen defects or ovule problems. Wild V. vinifera ssp. sylvestris staminate male flowers show completely non-receptive dry vestigial stigmas with dry appearance throughout bloom, no exudate secretion, and <5% pollen germination capability, representing non-functional stigmas where developmental suppression prevents any receptivity achievement.
A dry stigma is a stigma lacking wet stigmatic exudate on its surface, appearing dry or matte rather than glistening, representing either the pre-receptive state before exudate secretion activates, the post-receptive state after exudate production has ceased and surface exudate has dried, or the non-receptive state in functionally male and staminate flowers where exudate secretion is absent or minimal, with dry stigmas generally unable to support pollen germination and fertilization. Dry stigmas are characterized by= (1) absence of visible wet exudate coating papillae, with stigma surface appearing dry, matte, or dull rather than glistening (2) papillae in non-secretory state with secretory apparatus inactive (pre-receptive) or degraded (post-receptive), or secretory apparatus absent/non-functional (non-receptive functionally male/staminate types) (3) papillae often showing reduced turgor in post-receptive dry stigmas with cells partially collapsed or desiccated (4) surface sometimes showing remnants of dried crystallized exudate components in post-receptive stigmas, appearing as deposits or residues (5) inability to support pollen germination, with 6-8 days post-calyptra shedding) where exudate secretion has ceased and existing exudate has dried, representing normal developmental state after receptivity completion (3) non-receptive dry stigmas in functionally male and staminate flowers where exudate secretion is absent or minimal throughout the bloom period, representing genetic/developmental condition preventing normal receptivity achievement. The transition from dry (pre-receptive) to wet (receptive) marks the onset of stigma receptivity and defines when pollination becomes possible, typically occurring at or slightly before calyptra shedding (BBCH 60-61) in hermaphrodite flowers through activation of exudate secretion triggered by developmental signals and environmental conditions. The transition from wet (receptive) to dry (post-receptive) marks the end of the receptivity window and defines when pollination is no longer effective, typically occurring 5-8 days post-calyptra shedding as exudate secretion declines and ceases, with existing exudate drying through evaporation creating the dry surface. Dry stigmas show drastically reduced pollen germination support= controlled pollination experiments applying viable pollen to dry stigmas achieve <5-20% germination and <8-15% fruit set versus 70-90% germination and 45-65% fruit set on receptive wet stigmas, demonstrating that dry stigma condition is incompatible with successful pollination regardless of pollen availability or viability.
Dry_Stigma represents absence of wet exudate and must be distinguished from Receptive_Stigma (wet exudate present, pollen germination supported). The dry condition can represent normal developmental states (pre-receptive or post-receptive) or genetic/developmental conditions preventing receptivity (non-receptive functionally male/staminate types). Sub-classification into Non_Receptive_Dry and Post_Receptive_Dry distinguishes between stigmas that fail to achieve receptivity versus stigmas that achieved receptivity but have completed the receptivity period. Understanding dry stigma conditions is essential for interpreting stigma developmental state, for timing pollination operations, and for diagnosing reproductive failures related to stigma dysfunction or asynchrony between stigma state and pollen availability.
exudatlose Narbe
matte stigma surface
nicht-feuchtes Stigma
non-secreting stigma
trockene Narbe
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 102-108 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 95-101 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=94-106.
Dry Stigma
Drying sensation comparison between young Tannat (severe drying 8/10, chalky-parching character persisting 30+ seconds) and young Pinot Noir (mild drying 3/10, subtle textural change clearing within 10 seconds) demonstrates variety-driven tannin differences where Tannat's high tannin concentration creates dramatic mouth drying requiring substantial aging for softening, while Pinot Noir's naturally fine tannins produce gentle drying appropriate for early consumption, illustrating how drying intensity indicates both variety character and aging requirements.
Drying sensation quality is a tactile astringency quality that represents the perception of moisture removal and reduced lubrication in the oral cavity, characterizing the fundamental tactile component of astringency where the mouth feels progressively drier with continued wine contact as salivary proteins are precipitated by polyphenolic compounds. Within the BFO framework, this quality specifically addresses the dehydration-like dimension of astringent tactile perception, distinguishing the drying sensation from puckering while recognizing that drying provides the primary sensory evidence of tannin-saliva interaction that defines astringent character. The ontological significance lies in drying sensation providing the quantitative assessment of astringency intensity, where the degree of perceived dryness correlates with tannin concentration and reactivity, enabling evaluation of both current astringency level and aging trajectory through the relationship between drying intensity and tannin structural parameters. The physiological basis for drying sensation involves progressive depletion of salivary lubrication as proline-rich proteins bind to tannins and precipitate, with the resulting reduction in oral lubricant creating friction between tongue, palate, and cheek surfaces that mechanoreceptors interpret as dryness, a sensation that intensifies with repeated exposure as protein reserves are depleted faster than saliva can replenish them. The compositional factors affecting drying intensity include tannin concentration (primary determinant), tannin molecular weight distribution (smaller tannins creating more aggressive drying), tannin galloylation degree (increasing protein-binding affinity), and wine matrix factors (polysaccharides and proteins reducing drying, lower pH increasing it). The descriptive vocabulary for drying ranges from barely perceptible (slight textural change) through moderate (obvious dryness requiring palate refreshment) to severe (chalky, parching sensation dominating mouthfeel), with appropriate drying level depending on wine style and consumption context. Understanding drying sensation quality enables astringency intensity assessment, tannin quality evaluation from drying character, and quality judgment incorporating appropriate drying for wine style and intended aging trajectory.
Drying_Sensation_Quality addresses the moisture-reduction dimension of tactile astringency, representing the primary quantitative component of astringent perception. This quality correlates with tannin concentration and reactivity, enabling intensity assessment and aging prediction. The class supports astringency evaluation through the specific drying component distinct from puckering or roughening sensations.
Austrocknungsgefühl
Mundaustrocknung
Trocknungsempfindung
adstringierende Trockenheit
astringent dryness
mouth drying
oral dehydration
parching sensation
Prinz JF, Lucas PW (2000) Saliva tannin interactions. Journal of Oral Rehabilitation 27(11): 991-994 Condelli N et al. (2006) Prediction of perceived astringency induced by phenolic compounds. Food Quality and Preference 17(7-8): 555-564
Drying Sensation Quality
Riparia Gloire de Montpellier on shallow chalky soils in Champagne acts as a dwarfing rootstock for Chardonnay, with pruning weights falling below 0.15 kg/m in high-density plantings at 8,000-10,000 vines/ha, where the reduced canopy facilitates mechanical harvesting and promotes the high-acid, low-yield profile essential for base wine production. In experimental high-density trials at INRAE Montpellier, certain V. riparia × V. rupestris selections reduced Syrah scion growth to under 0.1 kg/m pruning weight at densities exceeding 12,000 vines/ha.
A vigor-modifying rootstock role that inheres in a grafted rootstock by virtue of its capacity to severely restrict scion vegetative growth, reducing pruning weight by 40-60% or more below moderate-vigor references. Dwarfing rootstocks produce scion pruning weights typically below 0.2 kg/m, shoot lengths often under 0.6 m per season, and markedly reduced canopy volume with leaf area indices below 1.5. The extreme growth limitation results from very restricted root system development, low root hydraulic conductance, minimal xylem vessel diameter (often below 40 µm), and strongly reduced cytokinin-to-ABA ratios in xylem sap. Dwarfing rootstocks are used in specialized contexts such as high-density planting systems (8,000-15,000 vines/ha), mechanized viticultural operations, and experimental settings investigating minimum canopy requirements for fruit maturation.
Dwarfing_Rootstock_Role is distinguished from Low_Vigor_Rootstock_Role by the severity of growth restriction, which may compromise economic viability at conventional planting densities (1,500-4,000 vines/ha). True dwarfing effects in grapevine rootstocks are less pronounced and less precisely controlled than in apple rootstocks (e.g., M.9, M.27), and the dwarfing classification in viticulture often reflects an interaction between inherently low-vigor genotypes and restricting site conditions rather than genotype alone. The concept is gaining renewed interest in the context of climate adaptation strategies seeking smaller canopies with reduced water demand.
Zwergunterlage
dwarfing rootstock
sehr schwache Unterlage
severely devigorating rootstock
stark schwachwüchsige Unterlage
very low vigor rootstock
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 285-310 Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review, American Journal of Enology and Viticulture 60(3): 251-268 Dry N (2007) Grapevine rootstocks, Lythrum Press
Dwarfing Rootstock Role
Cabernet Sauvignon showing dying senescent feeder roots in November, soil core examination revealing feeder roots with dark brown coloration, completely collapsed morphology, extensive tissue disintegration, microscopic examination showing cytoplasmic liquefaction with no recognizable organelles, FDA staining completely negative confirming membrane death, isotope tracing showing zero uptake, roots 9-13 months post-initiation in terminal dying phase comprising estimated 8% of feeder root population, dying feeder roots representing final stage before complete death and decomposition
Dying senescent feeder root is senescent feeder root in final stages of programmed cell death showing advanced degradation approaching complete loss of cellular organization, typically 6-14 months post-initiation representing terminal days to weeks of feeder root existence, characterized by complete loss of turgor with fully collapsed structure, extensive cytoplasmic disintegration with few or no recognizable organelles, severely compromised membranes allowing unrestricted leakage, dark brown coloration from accumulated phenolics and oxidation products, and imminent or occurring cell death marking transition from senescing living tissue to dead organic matter before decomposition, with dying senescent feeder roots showing terminal autophagy consuming remaining cellular contents, walls potentially beginning to fragment or separate from parent root, cortex potentially sloughing leaving only vascular cylinder remnant, and demonstrating that dying phase represents final irreversible stage where cellular machinery ceased function and feeder root transitioning to substrate for soil microorganisms contributing to decomposition and nutrient cycling. Dying senescent feeder root structural deterioration includes= showing complete collapse (total turgor loss causing feeder root to flatten or shrivel beyond recognition of original form), exhibiting extensive wall damage (cell walls potentially fragmenting, cracking, or separating), demonstrating cortical sloughing (cortex tissues potentially detaching from stele leaving exposed vascular cylinder), revealing tissue liquefaction (advanced degradation reducing tissues to amorphous material), and displaying detachment tendency (dying feeder roots potentially separating from parent root through continued degradation). Dying senescent feeder root membrane failure includes= showing complete permeability loss (membranes so degraded allowing free passage of all solutes eliminating compartmentation), exhibiting plasma membrane rupture (plasma membrane potentially completely broken down losing all barrier function), demonstrating organellar collapse (mitochondrial, ER, and other membranes completely degraded), revealing ion leakage (cellular ions freely leaking from dying root into soil solution), and displaying lipid degradation (membrane lipid breakdown producing free fatty acids and other decomposition products). Dying senescent feeder root biochemical endgame includes= showing terminal autophagy (final cellular self-digestion consuming last recyclable components), exhibiting maximum protease activity (complete protein degradation hydrolyzing remaining cellular proteins), demonstrating nucleic acid breakdown (complete DNA and RNA degradation by activated nucleases), revealing oxidative burst (potentially showing burst of reactive oxygen species marking terminal death), and displaying metabolite release (remaining cellular contents leaking or actively released during cell death). Dying senescent feeder root functional cessation includes= showing zero absorption (complete inability to uptake water or nutrients confirming functional death), exhibiting no respiration (complete cessation of oxygen consumption and CO2 production indicating metabolic death), demonstrating absent membrane potential (complete depolarization confirming transport cessation), revealing total unresponsiveness (complete inability to respond to any stimuli confirming cellular death), and displaying irreversible state (dying phase completely irreversible with no possibility of recovery). Dying senescent feeder root visible characteristics includes= showing dark brown to black coloration (extensive phenolic accumulation and oxidation creating dark appearance), exhibiting completely collapsed morphology (feeder root flattened or extremely shriveled unrecognizable from original cylindrical form), demonstrating fragmentation (dying root potentially breaking into pieces as walls fail), revealing surface disintegration (root surface potentially showing breaks, lesions, or complete tissue breakdown), and displaying microbial colonization (dying and dead roots often showing fungal hyphae or bacterial colonies accelerating decomposition). Dying senescent feeder root timing includes= showing brief duration (dying phase typically lasting only days to few weeks representing final stage before complete death), exhibiting rapid progression (once dying phase begins progressing quickly to complete death and decomposition), demonstrating variable onset (dying beginning 6-14 months after initiation depending on conditions and root characteristics), revealing position correlation (dying roots often in oldest most distal portions of root system), and displaying continuous occurrence (some feeder roots always in dying phase as part of continuous turnover). Dying senescent feeder root population context includes= showing small proportion (dying roots typically representing 5-15% of total feeder root population at any time due to brief dying phase), exhibiting spatial concentration (dying roots concentrated in zones with oldest roots or most stressful conditions), demonstrating turnover flux (dying root numbers reflecting rate at which feeder roots exiting functional pool), revealing replacement need (dying roots requiring replacement through new root production maintaining functional population), and displaying decomposition initiation (dying roots marking beginning of organic matter input from feeder root turnover). Dying senescent feeder root decomposition and fate includes= showing imminent breakdown (dying roots about to or beginning to decompose), exhibiting microbial attack (soil microorganisms colonizing and decomposing dying root tissues), demonstrating rapid decomposition (thin-walled nitrogen-rich feeder root tissues decomposing quickly once dead), revealing nutrient release (nitrogen, phosphorus, and other nutrients released during decomposition potentially available for plant uptake), and displaying carbon cycling (dying feeder root carbon contributing to soil respiration and organic matter pools). Dying senescent feeder root in carbon budget includes= representing production cost loss (dying feeder roots representing complete loss of carbon investment in their construction), showing opportunity cost (resources allocated to dying roots representing foregone opportunity for other allocations), exhibiting turnover component (dying feeder roots contributing to total annual root turnover carbon flux), demonstrating soil input (dying roots providing carbon input to soil though rapidly decomposing), and revealing system efficiency (high proportion of dying versus active roots potentially indicating inefficient root system requiring excessive production).
Dying_Senescent_Feeder_Root emphasizes terminal death stage and imminent decomposition and is distinguished from earlier senescence by focus on complete functional loss and structural collapse. Critical for understanding final degradation, membrane failure, biochemical endgame, decomposition initiation, and the terminal dying phase marking feeder root transition from senescing tissue to dead organic matter entering decomposition.
Final-stage senescent root
Moribund fine root
Moribunde Feinwurzel
Sterbende seneszente Feinwurzel
Terminal-seneszente Saugwurzel
Terminally dying feeder root
Dying Senescent Feeder Root
Chardonnay grafted on Teleki 5C rootstock showing dying senescent root hairs 55-65 mm from root tip, hairs 9-12 days post-initiation in terminal dying phase, microscopic examination revealing completely collapsed morphology with dark brown coloration, extensive cytoplasmic disintegration with no recognizable organelles, FDA viability staining completely negative indicating membrane death, some hairs partially detached from root surface, dying hairs representing <8% of total hair population due to brief dying phase preceding complete death and sloughing
Dying senescent root hair is senescent root hair in final stages of programmed cell death showing advanced degradation approaching complete loss of cellular organization, typically 7-12 days post-initiation representing terminal 1-2 days of hair existence, characterized by complete loss of turgor with fully collapsed structure, extensive cytoplasmic disintegration with few or no recognizable organelles, severely compromised membranes allowing unrestricted leakage, brown or dark coloration from accumulated phenolics and oxidation products, and imminent or occurring cell death marking transition from living senescing cell to dead cellular remains, with dying senescent root hair showing terminal autophagy consuming remaining cellular contents, wall potentially beginning to fragment or separate from root surface, and demonstrating that dying phase represents final irreversible stage of hair life where cellular machinery has ceased function and hair is transitioning to dead organic matter before sloughing or decomposition. Dying senescent root hair structural deterioration includes= showing complete collapse (total loss of turgor causing hair to flatten or shrivel beyond recognition of original cylindrical form), exhibiting extensive wall damage (cell wall potentially fragmenting, cracking, or separating from parent cell), demonstrating cytoplasmic liquefaction (cytoplasm degrading to amorphous material lacking organization), revealing organelle remnants only (no intact functional organelles remaining only debris from degradation), and displaying detachment tendency (dying hairs often partially or completely separating from root surface). Dying senescent root hair membrane failure includes= showing complete loss of selective permeability (membranes so damaged allowing free passage of all solutes losing all compartmentation), exhibiting plasma membrane rupture (plasma membrane potentially breaking down completely losing barrier function), demonstrating organellar membrane collapse (mitochondrial, ER, and other organellar membranes completely degraded), revealing ion leakage (cellular ions freely leaking from dying hair into apoplast or soil), and displaying lipid degradation products (membrane lipid breakdown producing fatty acids and other degradation compounds). Dying senescent root hair biochemical endgame includes= showing terminal autophagy (final stage of self-digestion consuming remaining recyclable cellular components), exhibiting protease activity peak (maximum protein degradation as cellular proteins completely hydrolyzed), demonstrating nucleic acid degradation (DNA and RNA completely broken down by nucleases activated during death process), revealing oxidative burst (potentially showing burst of reactive oxygen species marking terminal cell death), and displaying metabolite release (cellular contents leaking or actively released from dying cell). Dying senescent root hair functional cessation includes= showing zero absorption capacity (complete inability to uptake water or nutrients confirming functional death), exhibiting no respiratory activity (complete cessation of oxygen consumption and CO2 production indicating metabolic death), demonstrating absence of membrane potential (complete depolarization confirming membrane death and transport cessation), revealing no responsiveness (complete inability to respond to any stimuli confirming cellular death), and displaying irreversible state (dying phase irreversible with no possibility of recovery or resurrection). Dying senescent root hair visible indicators includes= showing dark brown to black coloration (extensive phenolic accumulation and oxidation creating characteristic dark appearance), exhibiting collapsed morphology (completely flattened or shriveled appearance markedly different from turgid living hairs), demonstrating fragmentation (hair potentially breaking into pieces as walls weaken and degrade), revealing detachment (dying hairs often separating partially or completely from parent trichoblast cell or root surface), and displaying fungal or bacterial colonization (dead and dying hairs often showing microbial growth accelerating decomposition). Dying senescent root hair timing includes= showing brief duration (dying phase typically lasting only 12-48 hours representing final stage of senescence), exhibiting rapid progression (once dying phase begins proceeding quickly to complete death), demonstrating variable onset (dying beginning 7-12 days after hair initiation depending on conditions and hair characteristics), revealing position correlation (dying hairs farthest from root tip marking oldest hair zone), and displaying continuous occurrence (some hairs always in dying phase as part of continuous turnover). Dying senescent root hair population context includes= showing small proportion (dying hairs typically representing <5-10% of total hair population at any time due to brief dying phase), exhibiting spatial pattern (dying hairs concentrated in oldest distal portion of hair zone), demonstrating continuous replacement (dying hairs continuously replaced by new hairs at proximal end maintaining functional population), revealing turnover flux (dying hair numbers reflecting turnover rate of entire hair population), and displaying decomposition initiation (dying hairs marking beginning of organic matter input from hair turnover to rhizosphere). Dying senescent root hair fate includes= showing imminent sloughing (dying hairs often completely separating from root falling into soil), exhibiting in situ decomposition (some dying hairs remaining attached decomposing in place), demonstrating microbial substrate (dying and dead hairs providing readily degradable substrate for rhizosphere microbes), revealing organic matter contribution (dying hairs contributing to soil organic matter particularly in hair-bearing root zones), and displaying nutrient recycling (cellular contents from dying hairs potentially recovered by microbes or plant released into rhizosphere).
Dying_Senescent_Root_Hair emphasizes terminal death stage and advanced degradation and is distinguished from earlier senescence by focus on imminent complete cell death. Critical for understanding final degradation, membrane failure, functional cessation, biochemical endgame, visible indicators of death, and the terminal dying phase marking transition from living senescing cell to dead organic matter before sloughing.
Final-stage senescent hair
Moribund root hair
Moribunde Wurzelhaare
Sterbende seneszente Wurzelhaare
Terminal-alternde Haare
Terminally dying hair
Dying Senescent Root Hair
Thompson Seedless exhibits characteristic early seed abortion at 7-12 days post-fertilization, producing 0.5-1.0 mm brown seed traces and requiring GA3 treatment (10-15 ppm at BBCH 71) to achieve 15-17 mm berry diameter, with embryo abortion occurring during globular stage before significant endosperm accumulation
An early aborted seed is an aborted seed of Vitis vinifera that undergoes developmental failure and tissue degeneration during early post-fertilization stages (1-3 weeks post-fertilization, corresponding to BBCH 71-73), resulting in minimal seed trace remnants (typically 0.2-1.5 mm) and significant reduction in final berry size in cultivars that normally produce seeds. Early seed abortion represents the most severe form of seed developmental failure, occurring before substantial embryo or endosperm development and before seeds have exerted significant hormonal influence on berry expansion. This abortion timing is characteristic of stenospermocarpic seedless cultivars where genetic programs trigger early embryo or endosperm collapse. Developmental characteristics of early aborted seeds include= abortion occurring during embryo globular or early heart stage (7-21 days post-fertilization), minimal tissue differentiation before abortion initiation, rapid tissue browning and desiccation following abortion, and integration of small brown remnants into developing berry locular cavity. The early timing of abortion has critical consequences for berry development= because developing seeds are primary sources of auxins and gibberellins that promote berry cell division and expansion during Phase I growth (0-4 weeks post-fruit set), early seed abortion results in dramatically reduced berry size. Seedless table grape cultivars with early abortion typically produce berries 6-12 mm diameter without gibberellin treatment compared to 12-18 mm for seeded cultivars. Commercial seedless table grape production employs exogenous gibberellin applications (typically GA3 at 5-20 ppm during fruit set and cell division phases) to compensate for loss of seed-derived hormones and achieve marketable berry sizes (14-20 mm diameter). Early aborted seeds in normally seeded cultivars indicate severe problems during fruit set= extreme environmental stress (temperatures 35°C during flowering and immediate post-fertilization), nutritional deficiencies (particularly boron deficiency affecting pollen tube growth and fertilization), severe water stress during critical 1-3 week post-fertilization period, or pathological factors affecting embryo or endosperm initiation.
Early_Aborted_Seed emphasizes very early developmental failure timing and is distinguished from Late_Aborted_Seed (abortion after substantial embryo/endosperm development) by minimal seed trace size and maximal impact on berry size reduction. Critical for understanding stenospermocarpic seedless mechanisms, gibberellin application requirements in seedless table grape production, and severe stress indicators during fruit set in seeded cultivars.
Frühabortierter Samen
Minimal seed trace
Minimaler Kernrest
Stenospermocarpic abortion
Stenospermokarpe Samenanlage
Early Aborted Seed
An early aborted tendril at node 3 on a severely nitrogen-deficient shoot of Vitis vinifera cv. Pinot Noir in Burgundy, France, appearing as a 1.2 cm brownish protrusion at the node position where a tendril would normally develop, showing no recognizable tendril morphology or differentiation beyond a simple bump with necrotic tip, occurring in a vineyard with soil nitrogen levels below 15 ppm nitrate-nitrogen during critical shoot development in May, with 75% of expected tendril positions on affected vines showing similar early abortion, while leaves showed characteristic nitrogen deficiency symptoms including pale yellow-green coloration and reduced size, and shoot extension was severely limited to 45 cm total length compared to normal 120-150 cm, measured at nominal BBCH stage 55 (inflorescence clearly visible) though inflorescence development was also severely compromised.
An aborted tendril that ceased development during the initial phases of organogenesis in Vitis vinifera, characterized by developmental arrest occurring before tendril primordia have completed basic structural differentiation into recognizable tendril form, typically manifesting as small protrusions, bumps, or rudimentary structures of 0.5-3 cm length at node positions where tendrils would normally develop, showing minimal tissue differentiation with largely undifferentiated parenchyma cells, absence of functional vascular tissue, and failure to initiate the elongation growth phase that would produce characteristic tendril morphology. Early aborted tendrils result from severe developmental constraints present at the earliest organogenic stages including extreme resource scarcity during shoot apex development when meristematic activity establishes organ primordia, critical environmental stresses including severe water deficit or temperature extremes during the 1-2 week period of initial tendril initiation, fundamental hormonal imbalances affecting organ identity specification or meristematic competence, severe pathogen infection or pest damage affecting shoot apex tissues, or genetic mutations disrupting early developmental programs. These abortion events represent the most severe developmental failures, occurring before significant metabolic investment in tendril structures, and indicate critical stress conditions or fundamental physiological problems affecting the entire shoot apex rather than individual organ development, often correlating with simultaneous failures in leaf development, internode elongation, or inflorescence formation.
Early aborted tendrils are distinguished from late aborted tendrils by the developmental stage at which arrest occurs, with early abortion preventing achievement of recognizable tendril morphology while late abortion occurs after basic structural differentiation is complete. Early abortion is diagnostically significant because it indicates severe stress during critical shoot apex development when organ primordia are being established, representing more serious physiological problems than late abortion which affects individual organs after apex function is established. The distinction between early aborted tendrils and complete tendril absence (no primordium formation) can be subtle, requiring careful inspection of node positions during early shoot development. Early abortion patterns correlating with leaf and inflorescence problems indicate systemic shoot apex dysfunction requiring urgent intervention, while isolated early tendril abortion may reflect normal developmental variation or localized damage.
anfangsstadium entwicklungsgestörte Kletterorgan
early developmental arrest tendril
früh-abortierte Ranke
initial stage failed tendril
primordial abortion tendril
primordial-stadium fehlgeschlagene Ranke
Based on Srinivasan & Mullins (1981) "Physiology of flowering in the grapevine - a review" American Journal of Enology and Viticulture 32=47-63 Boss & Thomas (2002) "Tendrils of grapevine Vitis vinifera L. are inflorescence homologues" Australian Journal of Grape and Wine Research 8=37-39 Carmona et al. (2008) "Grapevine bud dormancy= environmental and molecular aspects" Revista Brasileira de Fruticultura 30=257-408 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 329-352.
Early Aborted Tendril
Pinot Noir in Burgundy shows early anthesis during early June Year N+1 at BBCH 61-62 over 2-3 days June 3-5, with 10-20% flowers opening initially in apical cluster regions, caps shedding exposing reproductive organs, stigmas developing receptivity with exudate production, under continental climate conditions of 19-24°C temperature, 55-70% relative humidity, typical of this early-to-mid season cultivar initiating flowering before mid-June
Early anthesis represents the initial flowering phase when first flowers begin opening through cap shedding, establishing onset of reproductive function within cluster and marking critical transition from developmental to reproductive processes, corresponding to BBCH 61-62 stages with 10-20% flowers open. This critical initiation stage occurs during early June in temperate regions Year N+1, typically spanning 2-3 days as earliest-maturing flowers begin opening creating foundation for subsequent mass flowering, with timing representing major phenological marker for vineyard management and climate studies. Initial flower opening shows sporadic pattern with earliest flowers typically in apical cluster regions reflecting most advanced development from earliest primordium formation, with 5-10% flowers opening on first day increasing to 15-25% by end of early anthesis period, establishing progressive opening pattern. Cap shedding initiation in earliest flowers shows characteristic sequence with calyptra loosening visible as slight gap appearing at base between cap and receptacle, progressive lifting as abscission zone cells separate, upward movement driven by expanding reproductive organs and turgor forces, and complete detachment with cap falling away exposing reproductive structures. Reproductive organ exposure in early-opening flowers reveals functional structures with five stamens positioned around central pistil, anthers beginning to show color change from green to yellow-brown indicating approaching maturity, stigma surface developing papillate cells and exudate production, and overall flower structure delicate and vulnerable. Anther maturation in early-opening flowers shows final preparation for dehiscence with pollen completing maturation to tricellular state, anther walls undergoing structural changes preparing stomium zones for opening, and anthers positioned for effective pollen release and transfer to stigma. Stigma development in early-opening flowers shows receptivity onset with papillate surface cells elongating creating increased surface area, exudate secretion initiating producing adhesive proteinaceous coating capturing pollen, and stigma surface showing characteristic glistening appearance indicating functional receptivity. Pollen release timing shows variable pattern with some early-opening flowers releasing pollen within hours of cap shedding under favorable dry conditions, while others may delay release 12-24 hours if humidity is elevated, with initial pollen release establishing first opportunity for pollination within cluster. Pollination opportunity in early anthesis remains limited as relatively few flowers are open simultaneously, with successful pollination depending on overlap between pollen availability from early-opening flowers and stigma receptivity in same or neighboring flowers, creating first window for fruit set though majority of pollination occurs during subsequent peak anthesis. Flower appearance at early anthesis shows opened flowers with exposed stamens and pistil contrasting with majority of unopened buds still enclosed in calyptra caps, creating mixed developmental appearance within cluster with clear visual distinction between opened and closed flowers. Cluster-level progression shows characteristic basipetal-acropetal pattern with apical flowers often opening earliest, followed by middle region flowers, and basal flowers typically opening latest, though pattern shows variation among cultivars and environmental conditions affecting developmental uniformity. Environmental sensitivity intensifies during early anthesis with opened flowers highly vulnerable to adverse conditions including rain causing cap retention or washing pollen, excessive heat or cold affecting pollen viability or stigma function, and high humidity delaying anther dehiscence. Weather monitoring becomes critical during early anthesis as conditions during this period affect not only early-opening flowers but indicate likely conditions during subsequent peak anthesis, with unfavorable early anthesis weather potentially portending problems for entire flowering period. Temperature requirements for early anthesis show optimal 20-28°C supporting normal cap shedding and organ function, with cool conditions below 15°C substantially delaying opening and reducing organ vigor, and excessive heat above 32°C potentially causing organ dysfunction or accelerated senescence. Humidity effects on early anthesis show optimal 40-70% relative humidity supporting cap shedding and anther function, high humidity above 75% potentially preventing cap detachment or delaying dehiscence, and low humidity below 35% causing rapid tissue desiccation potentially affecting function. Metabolic activity in early-opening flowers shows elevated respiration supporting reproductive processes, active carbohydrate metabolism providing energy for organ function, and specialized metabolism in anthers supporting final pollen maturation and release. Hormonal status during early anthesis involves ethylene potentially involved in triggering cap abscission, declining growth hormones as reproductive transition completes, and preparing auxin system in ovaries ready for fertilization signal. Developmental coordination shows early anthesis occurring when shoot vegetative growth remains active with shoots typically having 12-18 expanded leaves, with concurrent vegetative-reproductive development requiring balanced resource allocation supporting both processes. Canopy microclimate effects become evident during early anthesis with well-exposed clusters showing earlier warmer conditions promoting earlier opening, while shaded interior clusters may show delayed opening creating within-vineyard heterogeneity. Management implications of early anthesis include providing indicator for subsequent peak anthesis timing enabling preparation of labor and operations, establishing critical period requiring weather monitoring and possible protective interventions, and representing final opportunity for pre-bloom disease management as fungicide applications become restricted once significant flowering begins. The process shows three distinguishable aspects progressing from first flowers opening establishing reproductive onset, through initial cap shedding revealing organ function, to stigma receptivity onset creating first pollination opportunity. Duration typically spans 2-3 days representing initiation period before peak flowering, with warm favorable conditions promoting rapid progression while cool conditions may extend period. Cultivar variation appears in early anthesis timing with early-flowering varieties showing onset late May-early June while late types begin mid-late June, proportion of flowers opening during early phase with some showing gradual 10-15% early opening while others show more abrupt 20-30% opening, and sensitivity to environmental conditions affecting initiation reliability. Regional timing shows early anthesis during late May-early June in warm climates corresponding to heat accumulation of 900-1100 growing degree days base 10°C, early-mid June in temperate regions at 800-1000 GDD, and mid-late June in cool areas at 700-900 GDD, with timing critical for frost avoidance and favorable weather window. Viticultural significance includes representing definitive flowering onset enabling precise phenological tracking, providing early indicator of flowering conditions affecting subsequent peak bloom, establishing critical weather-sensitive period requiring monitoring and possible intervention, and marking transition point from pre-bloom to bloom management requiring operational adjustments. Research applications include documenting precise anthesis initiation timing for phenology studies, analyzing environmental controls of flowering onset affecting adaptation, evaluating developmental uniformity affecting anthesis duration and fruit set patterns, quantifying early pollination success contributing to overall fruit set, and assessing climate change impacts on flowering timing across regions.
Early_Anthesis is first of two major Anthesis subphases, distinguished by initial flower opening establishing reproductive onset. Corresponds to BBCH 61-62 with 10-20% flowers open. Subdivided into First_Flowers_Opening, Initial_Cap_Shedding, and Stigma_Receptivity_Onset. Transitions to Peak_Anthesis as flowering intensity increases. Critical period for weather sensitivity and management preparation. Timing represents major phenological marker.
Blühbeginn
beginnende Blüte
beginning anthesis
early flower opening
flowering onset
frühe Anthese
initial bloom
initiale Blütenöffnung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 360-368 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Early Anthesis
Pinot Noir in Burgundy shows early berry formation during early-mid July Year N+1 at BBCH 71, with pericarp differentiating into 8-10 cell layer exocarp skin, abundant mesocarp flesh cells, and 2-3 cell layer endocarp, vascular network ramifying through tissues, chlorophyll content reaching 0.8-1.2 mg/g fresh weight creating green photosynthetic berry, berry diameter 14-20 mm, under continental conditions of 22-28°C temperature, typical of this early-to-mid season cultivar establishing functional berry structure
Early berry formation represents the structural organization phase during berry initiation when rapidly expanding ovary differentiates into recognizable berry architecture with distinct pericarp layers including exocarp skin, mesocarp flesh, and endocarp seed cavity lining, establishes vascular network for nutrient delivery, develops green photosynthetically active tissues, and creates functional berry structure capable of supporting continued development through ripening. This essential organization phase occurs during early fruit development in late June-early July in temperate regions Year N+1, proceeding over 2-3 weeks as berry structure establishes, corresponding to BBCH 71 young fruit stages. Berry formation encompasses coordinated processes including pericarp tissue differentiation creating specialized layers, vascular system development establishing transport infrastructure, cellular organization producing functional tissue architecture, chlorophyll accumulation creating green photosynthetic berry, and structural maturation preparing berry for subsequent developmental phases. Pericarp differentiation shows subdivision into three distinct tissue layers with specific functions, differentiation proceeding during early expansion phase, and layer specialization establishing by 3-4 weeks post-fertilization. Exocarp outer layer development shows formation of future berry skin from outer pericarp cells, cell division producing epidermal and subepidermal layers, chloroplast development creating green coloration, cuticle formation beginning waxy surface protection, and specialized cell types differentiating including future stomata though functionality declining later. Exocarp characteristics show typically 6-12 cell layers thick in young berry, actively photosynthetic during early development contributing 5-15% berry carbon requirements, protective barrier function against pathogens and water loss, and eventual transformation during ripening to colored skin. Mesocarp middle layer development shows formation of bulk berry flesh tissue from middle pericarp region, extensive cell division producing abundant flesh cells, cell enlargement through water uptake creating juicy tissue, vascular tissue ramification throughout mesocarp, and structural organization creating functional flesh capable of storage and expansion. Mesocarp characteristics show representing 80-90% berry volume in mature fruit, cells containing vacuoles storing water and eventually sugars acids and other compounds, tissue providing major storage capacity for ripening compounds, and architecture determining berry texture and quality. Endocarp inner layer development shows formation of seed cavity lining from inner pericarp adjacent to developing seeds, thinner layer compared to mesocarp typically 2-4 cell layers, specialized cells possibly providing seed-pericarp communication, and protective function surrounding developing seeds. Vascular system development shows differentiation of xylem and phloem bundles within berry, primary vasculature entering through pedicel and receptacle, branching vascular network ramifying through mesocarp reaching exocarp, xylem providing water and mineral delivery during growth, and phloem preparing for sugar import during ripening though initially less active. Vascular architecture shows major bundles in receptacle region distributing to berry tissues, radial branches extending through mesocarp toward skin, anastomosing network creating redundant pathways, and terminal endings providing local resource delivery. Xylem function during early formation shows active water import supporting cell expansion, mineral nutrient delivery providing essential elements, and xylem-mediated import dominating during growth phase before phloem activation. Chlorophyll accumulation shows chloroplast development in exocarp and outer mesocarp cells, chlorophyll synthesis creating green coloration characteristic of young berries, photosynthetic capacity developing contributing to berry carbon economy, maximum chlorophyll content typically 2-4 weeks post-fertilization, and eventual chlorophyll degradation during ripening revealing anthocyanin colors in red cultivars or carotenoid yellows in white cultivars. Berry photosynthesis characteristics show young green berries photosynthetically active with measurable CO2 fixation, photosynthesis contributing 5-15% berry carbon requirements supplementing vine import, activity highest in young berries declining as ripening approaches, and eventual cessation as chlorophyll degrades. Cellular organization shows coordinated cell arrangement within tissues, cell-cell communication through plasmodesmata enabling signaling, tissue cohesion maintaining structural integrity, and organized architecture supporting berry function. Cell wall characteristics show primary walls in young dividing cells enabling continued division, progressive wall thickening as cells mature, pectin-rich middle lamella between cells maintaining tissue cohesion, and wall properties affecting berry texture. Tissue mechanical properties show firm texture in young berry from active turgor and cell wall integrity, mechanical strength from organized cellular architecture, and structural capacity to support continued expansion and development. Berry appearance characteristics show green coloration from chlorophyll, smooth surface from active epidermal cell division, firm texture from turgor, spherical to ovoid shape, and overall young berry morphology distinct from flowers. Berry size at formation completion shows typical diameter 12-25 mm by 3-4 weeks post-fertilization depending on cultivar and conditions, size range reflecting seed number effects and environmental variation, and size at this stage indicating developmental progress. Environmental effects on berry formation include temperature affecting differentiation and development rate with optimal 22-28°C supporting normal progression, nutrient availability affecting tissue synthesis particularly nitrogen for protein, light possibly influencing chlorophyll content and photosynthesis, and overall favorable conditions supporting organized development. Temperature effects show warm conditions promoting faster differentiation and structure establishment, cool conditions slowing but generally not preventing formation, and temperature as factor affecting formation timing. Nutrient effects show nitrogen essential for protein synthesis in dividing and differentiating cells, phosphorus and potassium supporting metabolism, and micronutrients enabling enzymatic functions. Failed formation causes include inadequate cell division limiting tissue development, environmental stress disrupting differentiation, developmental abnormalities preventing proper organization, though failures rare after successful expansion initiation. The process represents third phase of berry initiation establishing functional structure, following ovary expansion initiation and preceding berry commitment. Duration typically 2-3 weeks though overlapping with expansion. Cultivar variation includes pericarp layer thickness ratios, vascular architecture patterns, and structural characteristics affecting berry quality. Regional timing shows early berry formation during early-mid July across climates as berry structure establishes. Viticultural significance includes establishing berry functional architecture determining quality potential, creating photosynthetic capacity contributing to carbon economy, developing vascular system supporting continued development, and producing recognizable berry structure distinguishing from flowers. Research applications include analyzing pericarp differentiation mechanisms and tissue specialization, investigating vascular development and architecture, evaluating environmental effects on structural formation, quantifying berry photosynthesis contributions, and assessing berry formation as quality determinant.
Early_Berry_Formation is third phase of Berry_Initiation, distinguished by structural organization and tissue differentiation. Represents pericarp differentiation into exocarp, mesocarp, endocarp layers. Vascular development establishes transport network. Chlorophyll accumulation creates photosynthetic green berry. Duration 2-3 weeks overlapping with expansion. Establishes functional berry architecture. Follows Ovary_Expansion_Initiation. Precedes Berry_Commitment. Determines berry structural foundation affecting quality.
Beeren-Architektur-Etablierung
Beerenstruktur-Entwicklung
Perikarp-Differenzierung
berry architecture establishment
berry structure development
frühe Beerenbildung
pericarp differentiation
young berry organization
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 420-425 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Early Berry Formation
Chardonnay in Champagne shows early bud swelling at BBCH 01 in late March after 800-900 chilling hours and 120-150 growing degree days base 10°C, with buds increasing 5-8% in volume over 3-4 days under conditions of 11-13°C mean temperature and 13-hour photoperiod, representing initial growth resumption in this early-bursting cultivar typical of cool maritime climate
Early bud swelling represents the initial detectable phase of bud expansion following dormancy release, characterized by subtle volumetric increase of 5-10% as dormant buds transition from quiescent to actively growing state. This developmental stage corresponds to BBCH 01, occurring when winter buds first show visible enlargement though scales remain tightly appressed and green tissue is completely concealed. The process initiates when accumulated growing degree days trigger metabolic reactivation in meristematic tissues, with cells resuming protein synthesis and starch mobilization providing soluble sugars for osmotic water uptake. At this earliest phase, expansion results primarily from cell expansion rather than division, with turgor pressure increasing through osmotic potential changes as stored reserves mobilize. The bud profile changes from winter's compressed ellipsoid to slightly rounded form, with the most noticeable change occurring at bud tip where apical meristem drives initial expansion. Bud scales remain brown and appear dry with tight overlap between adjacent scales showing no visible gaps, distinguishing this stage from more advanced swelling where scale margins begin separating. Hormonal changes include declining abscisic acid from winter dormancy levels and initial cytokinin accumulation transported from root systems resuming activity with soil warming. Respiration rates increase 3-5 fold from dormancy baselines as mitochondrial activity escalates to support growth, creating measurable temperature elevation in bud tissues through metabolic heat production. Vascular tissues between bud and supporting cane begin differentiating to establish functional xylem for water transport, critical for sustaining turgor-driven expansion. Water content in bud tissues increases from winter's 35-40% to 45-50% fresh weight as hydration supports cellular metabolism and physical expansion. The process requires sustained temperatures above 10°C for 3-5 consecutive days to maintain development, with cooler periods temporarily slowing but not reversing expansion. Cultivar differences appear in minimum temperature thresholds triggering this initial swelling, ranging from 8-9°C for early varieties like Chardonnay to 11-12°C for later varieties like Cabernet Sauvignon. Duration of early swelling phase varies from 2-4 days in warm spring conditions to 5-8 days when temperatures fluctuate near threshold values. Regional timing follows chilling requirement completion and heat accumulation patterns, occurring late March in warm climates, mid-April in temperate regions, and late April to early May in cool continental areas. Viticultural significance includes providing first visible confirmation of dormancy release for timing completion of dormant-season vineyard operations like pruning, serving as developmental marker for initiating early-season pest monitoring programs, and indicating need to prepare frost protection measures as increasingly vulnerable tissues develop. Frost damage at this stage causes cell death through ice crystal formation in hydrating tissues, resulting in bud mortality requiring compensatory growth from secondary buds. Research applications include phenological modeling incorporating early swelling as first observable response to heat accumulation, microscopic analysis of cellular changes accompanying dormancy exit, and molecular profiling of gene expression changes initiating growth resumption in perennial woody plants.
Early_Bud_Swelling is the first of three Bud_Swelling subphases, distinguished by subtle expansion with fully closed scales and no visible gaps. Transitions to Advanced_Bud_Swelling when scale margins begin separating. Most difficult stage to identify consistently requiring close observation. Temperature fluctuations commonly cause progression-regression cycles at this developmental point.
beginnende Knospenquellung
beginning bud expansion
erste Knospenschwellung
first bud enlargement
initial bud swelling
initiale Schwellung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 157-165 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Jones GV & Davis RE (2000) Climate influences on grapevine phenology, grape composition, and wine production and quality for Bordeaux, France, Am J Enol Vitic 51:249-261
Early Bud Swelling
Chardonnay exhibits early budbreak disposition, reaching BBCH 09 approximately 7-14 days before the cultivar median in multi-variety trials, with heat sum requirement of only 50-70 GDD (base 10°C) post-dormancy release. Chasselas and Gewürztraminer show similarly early budbreak, emerging within the first 20% of V. vinifera cultivars across European phenology networks. Early budbreak in Pinot Noir at Burgundy reference sites (Clos de Vougeot) occurs at mean date of April 8 versus April 22 for late cultivars. Spring frost damage frequency in early cultivars exceeds late cultivars by 2-4× in continental climates with April frost probability above 30%.
A budbreak timing disposition that is the inherited capacity of a Vitis vinifera plant to initiate budbreak earlier than average for the species under given environmental conditions, typically within the first tercile of cultivar budbreak dates, requiring lower heat accumulation following dormancy release.
Early budbreak disposition confers advantages in short-season climates by maximizing the available growing period, but increases frost risk and may advance flowering into suboptimal pollination conditions. The trait results from lower thermal thresholds for meristem reactivation and/or reduced heat accumulation requirement following endodormancy release. Site selection (elevated, air-drained) partially mitigates frost risk for early cultivars. Climate warming is differentially advancing early cultivars, potentially disrupting traditional cultivar phenological sequences.
Frühaustrieb
early budburst
early spring phenology
früher Austrieb
frühes Knospentreiben
precocious budbreak
Jones GV, Davis RE (2000) Climate influences on grapevine phenology, grape composition, and wine production and quality for Bordeaux, France. American Journal of Enology and Viticulture 51:249-261 Leolini L et al. (2018) Late spring frost impacts on future grapevine distribution in Europe. Field Crops Research 222:197-208
Early Budbreak Disposition
Syrah in Rhône Valley shows early cap shedding during early June Year N+1 at BBCH 61-62 over 1-2 days June 5-6, with cap fall expanding from initial 6-8% to 18-24% flowers showing detachment predominantly in apical and middle cluster regions, accelerating toward mass fall, under warm Mediterranean conditions of 23-27°C temperature, 52-66% relative humidity supporting efficient progression, typical of this mid-to-late season cultivar showing rapid flowering expansion
Early cap shedding represents the progressive calyptra detachment phase following first detachment when increasing numbers of flowers complete abscission expanding cap fall activity from sporadic initial event to broader cluster involvement, establishing accelerating pattern toward mass cap fall during peak anthesis. This expansion phase occurs during early flowering in early June in temperate regions Year N+1, typically spanning 1-2 days as cap fall frequency increases from initial 5-10% to 15-25% flowers showing detachment, corresponding to BBCH 61-62 progressive stages. Shedding expansion pattern shows characteristic acceleration with increasing daily detachment frequency from sporadic first event to regular progressive cap fall, spatial extension from initial apical location toward middle cluster regions as additional flowers reach maturity, and temporal intensification preparing transition toward mass cap fall period. Cap fall frequency progression shows daily increase with first day showing 5-10% total flowers with detached caps, second day showing 12-18% accumulated detachment representing 5-10% additional daily increment, and accelerating pattern indicating approaching peak flowering. Spatial progression shows basipetal-acropetal pattern with cap fall extending from apical initiation site toward middle cluster regions reflecting developmental gradient, occasional scattered detachment in middle regions from locally advanced flowers, and overall expansion creating broadening cap fall zone within cluster. Detached cap accumulation creates increasingly visible evidence of flowering with fresh caps showing recent separation accumulating on cluster surfaces, fallen caps on ground below clusters indicating flowering extent, and progressive accumulation documenting expansion of flowering activity. Flower exposure expansion reveals increasing numbers of opened flowers throughout cluster with newly exposed reproductive organs showing fresh pristine appearance, spatial distribution reflecting cap fall progression pattern, and overall transition from predominantly closed-bud cluster toward mixed open-closed appearance. Environmental effects on shedding progression show continued humidity sensitivity with favorable moderate 50-70% levels supporting efficient detachment expansion, high humidity above 75% potentially slowing progression through interference with abscission processes, and optimal temperature 20-26°C maintaining enzymatic activity supporting accelerating cap fall. Temperature during early shedding affects progression rate with warm conditions 24-28°C promoting rapid acceleration toward mass fall potentially compressing early phase to 1 day, cool conditions 18-22°C slowing progression potentially extending phase to 2-3 days, and overall thermal accumulation driving developmental rate. Humidity during early shedding critically affects efficiency with moderate levels enabling normal progression, elevated humidity potentially causing retention problems in susceptible flowers delaying progression, and optimal conditions minimizing retention supporting smooth acceleration toward peak. Weather stability during early shedding affects uniformity with consistent favorable conditions promoting coordinated progression creating compressed flowering period, variable conditions causing irregular progression potentially extending flowering and reducing synchrony. Cap retention problems may become evident during early shedding with some flowers showing incomplete abscission or persistent caps, retention frequency indicating likely problems during peak flowering, and early identification enabling assessment of potential fruit set difficulties. Abscission zone activity intensifies during early shedding with enzymatic processes accelerating in progressively more flowers, coordinated activation across multiple flowers creating expanding detachment zone, and overall abscission activity transitioning from sporadic individual events toward mass synchronized processes. Developmental readiness indicators show flowers completing maturation preparing for cap shedding through calyptra loosening visible as slight gaps at base, organ expansion creating mechanical pressure against caps, and progressive physiological preparation for reproductive function. Observational characteristics enable recognition of early shedding through increasing cap fall frequency distinguishing from single first event, spatial expansion showing broader cluster involvement, visible cap accumulation documenting flowering progression, and overall transition from sparse to moderate cap fall activity. Phenological progression shows early shedding representing intermediate stage between flowering initiation BBCH 61 and peak flowering BBCH 63-65, with rapid progression typically requiring 1-2 days, and clear trajectory toward mass cap fall representing peak reproductive activity. Management implications include confirming flowering expansion indicating approaching peak requiring readiness for critical bloom period, continued weather monitoring as sensitivity intensifies with expanding reproductive activity, and final preparations for bloom-period management including disease protection strategies. Flowering intensity assessment uses early shedding pattern as indicator with rapid acceleration suggesting compressed intensive flowering supporting high fruit set potential, slow irregular progression indicating extended flowering potentially affecting fruit set uniformity. Cultivar differences in early shedding show variation in acceleration rate with some varieties showing rapid 1-day progression while others show gradual 2-3 day expansion, spatial pattern reflecting cultivar-specific developmental uniformity, and overall progression characteristics affecting flowering duration and synchrony. The process represents second aspect of initial cap fall focusing on expansion from first event toward mass activity, following first calyptra detachment. Duration typically spans 1-2 days representing acceleration phase, with warm favorable conditions promoting rapid progression while variable conditions may extend period. Regional timing shows early cap shedding during early June in warm climates 1-2 days after first detachment, early-mid June in temperate regions, and mid-late June in cool areas, occurring during initial flowering expansion. Viticultural significance includes documenting flowering expansion confirming progression toward peak bloom, providing indication of flowering intensity and potential uniformity affecting fruit set predictions, revealing early retention problems indicating potential coulure issues, and establishing temporal pattern for subsequent mass cap fall. Research applications include analyzing progression dynamics from initiation to mass flowering, evaluating environmental effects on cap fall acceleration affecting flowering patterns, quantifying cultivar differences in shedding progression affecting flowering characteristics, assessing relationships between early shedding patterns and ultimate fruit set outcomes, and developing models predicting peak flowering timing from early shedding observations.
Early_Cap_Shedding is second aspect of Initial_Cap_Fall, distinguished by progressive expansion from first detachment toward broader cluster involvement. Follows First_Calyptra_Detachment. Represents acceleration phase preparing for mass fall. Duration 1-2 days showing increasing frequency. Environmental conditions particularly humidity critically affect progression efficiency. Retention problems may become evident indicating potential fruit set issues. Provides indication of flowering intensity and coordination.
Kappenfall-Ausbreitung
early calyptra detachment phase
expandierender Kalyptrafall
expanding cap shedding
frühe Abwurfphase
progressive Kappenabtrennung
progressive cap fall
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 365-370 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Early Cap Shedding
Merlot in Bordeaux shows early cell division growth during early-mid July Year N+1 at BBCH 71-73, with peak mitotic activity weeks 1-3 post-fertilization, cell cycle times 11-14 hours, progressive accumulation reaching 1.8-2.6 million mesocarp cells by week 4, under oceanic conditions of 20-26°C temperature, 1.6-2.2 seeds per berry supporting division through auxin, typical of this mid-season cultivar establishing cell number determining 10-13mm final berry size
Early cell division growth represents the initial intensive mitotic phase during berry development when pericarp mesocarp cells undergo peak division activity producing berry flesh cellularity, establishing final cell number determining berry size potential, and creating cellular foundation for subsequent expansion, proceeding over first 3-4 weeks post-fertilization as primary growth mechanism. This critical cell proliferation phase occurs during early summer in late June-July in temperate regions Year N+1, spanning approximately 2-4 weeks from berry commitment, corresponding to BBCH 71-73 young fruit stages. Cell division growth mechanism involves auxin-stimulated cell cycle activation in mesocarp tissue, peak mitotic activity with rapid cell proliferation, progressive cell number accumulation throughout pericarp layers, and division completion establishing final cell population determining berry flesh cellularity and size potential. Mitotic activity characteristics show maximum division intensity during weeks 1-3 post-fertilization, cell cycle times 10-15 hours during peak activity representing very rapid division, multiple rounds of division per initial cell producing clonal populations, and progressive division rate decline toward week 4 as cells transition to expansion mode. Cell cycle regulation shows cyclin-dependent kinases driving cell cycle progression, checkpoint controls ensuring DNA replication and chromosome segregation fidelity, auxin and gibberellin signaling promoting division, and eventual cell cycle exit as cells commit to expansion. Cell division spatial patterns show divisions occurring throughout mesocarp tissue creating flesh cells, more intensive division in outer mesocarp regions compared to inner areas near seeds in some studies, coordinated division across pericarp maintaining tissue proportions, and exocarp skin cells also dividing though less intensively than mesocarp. Cell number accumulation shows exponential increase during peak division weeks 1-3, typical accumulation from initial ~50,000-100,000 cells at commitment to 1-4 million cells by division completion, final cell number range reflecting cultivar genetic variation and environmental effects, and cell number as critical determinant of berry size. Cell number measurement methods include cell counting through tissue dissociation and microscopy, DNA content analysis estimating cell number, and histological section analysis though technically challenging. Final cell number determinants show genetic programming as primary factor with cultivar characteristic ranges, seed number effects through auxin dose influencing division intensity and duration, temperature affecting division rate with warm conditions promoting more divisions, and overall favorable conditions during critical 3-4 week period maximizing cell number. Seed number effects on cell division show positive correlation between seed number and final berry cell number, multiple seeds producing greater total auxin stimulating more intensive division, typical 20-40% more cells in 3-seed versus 1-seed berries, and auxin dose-response relationship explaining seed number effects on berry size. Temperature effects on division show optimal 24-28°C supporting maximum division rate and cell number achievement, cool conditions 18-22°C slowing division potentially reducing final cell number 15-30%, warm conditions 28-32°C potentially accelerating division though excessive heat above 35°C may impair division, and temperature during critical weeks 1-4 as major cell number determinant. Nutrient effects show nitrogen essential for protein synthesis supporting rapid division, phosphorus for DNA synthesis and energy metabolism, and adequate mineral nutrition supporting optimal division though usually not limiting in vineyard conditions. Division completion timing shows progressive transition from division to expansion beginning around week 3, division largely complete by week 4 in most cells, occasional continued division in some cell populations extending to week 5-6, and division cessation marking transition to expansion-dominated growth phase. Cell number as size determinant shows strong correlation between final cell number and mature berry size, more cells providing greater expansion capacity, typical 60-80% berry size variation attributable to cell number differences, and cell number established during early weeks determining size potential regardless of subsequent expansion. Cell size at division completion shows cells relatively small typically 20-40 micrometers diameter at division exit, cells capable of 10-100 fold volume expansion during subsequent enlargement, and initial small cell size enabling efficient packing in growing berry. Division phase versus expansion phase contribution shows cell number (division product) accounting for 60-80% final size variation, cell expansion accounting for 20-40% final size variation, and division phase as more critical determinant emphasizing importance of early favorable conditions. Cultivar variation in division capacity shows genetic differences in maximum cell number potential, small-berried cultivars typically 0.8-1.5 million cells achieving 6-8mm final diameter, large-berried cultivars 2.5-4 million cells achieving 14-18mm diameter, and genetic programming as primary cell number determinant. Environmental limitations on division show suboptimal temperature or water stress during critical early weeks permanently reducing cell number, lost division potential irreversible with later favorable conditions unable to compensate, and early environment as critical determinant of size potential. The process represents first phase of initial berry growth emphasizing cell proliferation, preceding rapid initial expansion and berry size establishment. Duration typically 2-4 weeks representing peak division though declining activity continues to week 4-5. Regional timing shows early cell division growth during late June-July in warm climates, early-mid July in temperate regions, and mid-late July in cool areas. Viticultural significance includes establishing final cell number as major berry size determinant, demonstrating critical importance of favorable conditions during early weeks for size potential, showing irreversible nature of early development affecting final outcomes, and emphasizing early environment as quality and yield determinant. Research applications include analyzing cell division regulation and controls, investigating auxin and hormone effects on division, evaluating temperature and environmental effects on cell number, quantifying cell number as size determinant, and assessing division phase as berry quality determinant.
Early_Cell_Division_Growth is first phase of Initial_Berry_Growth, distinguished by intensive mitotic activity establishing final cell number. Duration 2-4 weeks with peak weeks 1-3. Cell cycle times 10-15 hours during maximum activity. Final cell number 1-4 million determining 60-80% size variation. Temperature and seed number major determinants. Division completion by week 4. Irreversible cell number establishment. Precedes Rapid_Initial_Expansion. Critical phase for berry size potential.
Beeren-Zellproliferation
berry cell proliferation
early division phase
frühe Zellteilungs-Wachstum
intensive mitotic phase
intensive mitotische Phase
Early Cell Division Growth
Cabernet Sauvignon early developing berries progress from fruit set at BBCH 71 (2-2.3 mm diameter, 0.006-0.01 g fresh weight, containing 0.6-1 million cells, 10-14 days post-anthesis) through rapid growth achieving 8.5-10.5 mm diameter, 0.9-1.3 g, containing 2-2.8 million cells at BBCH 75 (end of Phase I, 42-52 days post-anthesis), with green chlorophyll-containing skin, hard texture, acidity increasing from 10 g/L at fruit set to 22-24 g/L at BBCH 75, and sugars remaining at 5-7% throughout early development phase. Chardonnay early developing berries examined at weekly intervals show cell division phase spanning weeks 0-5 post-fruit set with cell number increasing from 0.7 million (fruit set) to 1.2 million (week 2) to 1.9 million (week 4) to 2.3 million (week 5-6, BBCH 75), with cell division rate peaking during weeks 2-4 then declining, and with berry diameter increasing from 2 mm to 4.5 mm (week 2) to 7.5 mm (week 4) to 10.5 mm (week 6) tracking cell number increases and initial cell expansion. Pinot Noir early developing berries show photosynthetic activity in green skin and outer flesh tissues contributing 10-14% of berry carbon requirements during weeks 2-5 post-fruit set (BBCH 73-75), with net photosynthesis measured at 2-4 μmol CO₂ m⁻² s⁻¹ under full sun, declining as berries approach end of Phase I and transition to lag phase when photosynthetic contribution diminishes. Merlot experiencing water deficit during early berry development (0-4 weeks post-fruit set) shows reduced cell division resulting in final cell number of 1.8-2.2 million per berry versus 2.4-2.9 million under adequate water, creating smaller berries at maturity (12-14 mm diameter, 1.5-2 g) versus normal size (14-17 mm, 2.2-3 g), demonstrating that early development phase cell division determines ultimate berry size potential independent of later environmental conditions.
An early developing berry is a developing berry in the initial post-fertilization growth phase (BBCH 71-75) spanning approximately 0-6 weeks post-fruit set, characterized by rapid growth through active cell division and initial cell expansion, green coloration, hard texture, high acidity, low sugar content, and active establishment of the berry developmental program, representing the first phase of the double-sigmoid growth curve (Phase I or growth stage I). Early developing berries are characterized by= (1) rapid size increase from initial 1.5-3 mm diameter at fruit set to 6-11 mm diameter by end of early development phase, achieving 50-80% of final berry diameter through this first growth phase (2) fresh weight increase from 0.005-0.015 g at fruit set to 0.6-1.5 g by BBCH 75 (end of Phase I), representing approximately 40-70% of final berry fresh weight (3) active cell division throughout early development creating the final cell number that determines berry size potential, with cell number increasing from approximately 0.5-1.5 million cells at fruit set to 1.5-3.5 million cells by BBCH 75 depending on cultivar and environmental conditions (4) green coloration due to chlorophyll in skin and outer flesh tissues, with berries photosynthetically active contributing 8-15% of their carbon requirement through photosynthesis (5) hard firm texture due to small cell size (30-60 μm diameter), high turgor pressure, and rigid cell walls not yet modified by ripening processes (6) high acidity with titratable acidity increasing from 8-12 g/L at fruit set to peak levels of 15-25 g/L by mid-to-late Phase I (BBCH 73-75), with malic acid accumulating to maximum concentrations while tartaric acid remains relatively stable (7) low sugar content typically 3-8% during most of Phase I, with sugars primarily imported from maternal tissues to support growth rather than accumulating for storage. Early developing berries undergo critical developmental processes= ovary-to-berry transition establishing fruit identity and commitment to development pedicel vascular system differentiation creating the transport pathways supporting berry growth seed development and embryo formation (in seeded types) or seed initiation and abortion (in stenospermocarpic seedless types) establishment of berry size potential through determination of final cell number. Environmental factors during early berry development have lasting effects on final berry characteristics= water availability affects cell division and final cell number, influencing ultimate berry size potential temperature influences development rate and cell division duration light exposure affects photosynthesis in green berries and may influence cell division through effects on carbohydrate availability nutritional status particularly nitrogen availability affects vigor and cell division rates. Early developing berries are highly sensitive to environmental stresses that can cause fruit set failure, coulure (excessive berry drop), or millerandage (shot berries remaining small), with the 0-3 week post-fruit set period representing maximum vulnerability to these disorders.
Early_Developing_Berry represents the first growth phase (Phase I, BBCH 71-75, weeks 0-6 post-fruit set) and is distinguished from Mid_Developing_Berry (lag phase and veraison, BBCH 77-83, weeks 6-11) by active cell division versus minimal growth and metabolic transition. Early developing berries are characterized by rapid growth, green coloration, high acidity, and establishment of size potential through cell division. Sub-classification into Post_Fertilization_Early and Initial_Growth_Early reflects progression within the early development phase. Understanding early berry development is critical for managing fruit set, predicting final berry size, and optimizing early-season cultural practices affecting berry characteristics.
Phase I berry
Phase-I-Traube
früh entwickelnde Beere
junge wachsende Beere
young green berry
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 175-192 Coombe, B.G. (1992). Research on development and ripening of the grape berry. American Journal of Enology and Viticulture 43=101-110 Ojeda, H. et al. (2002). Influence of water deficits on grape berry growth. Vitis 41=141-145.
Early Developing Berry
Pinot Noir early enlargement stage berries progress from veraison at BBCH 81 (week 9, cells 70 μm diameter, berry 10.5 mm, 1.3 g, 14% sugars, 18 g/L acidity, 0.9 mg/g anthocyanins) through early enlargement achieving BBCH 85 (week 13, cells 115 μm, berry 13.5 mm, 2.1 g, 20% sugars, 12 g/L acidity, 2.2 mg/g anthocyanins), with cell expansion at 11.25 μm per week driving berry diameter increase at 0.75 mm per week and weight increase at 0.2 g per week, sugar accumulation at 1.5% per week, acid decline at 1.5 g/L per week, and anthocyanin accumulation at 0.33 mg/g per week, demonstrating rapid coordinated changes during early enlargement creating substantial ripening progression over 4-week period. Cabernet Sauvignon early enlargement stage berries under three irrigation treatments show full irrigation producing maximum expansion (cells from 72 μm at veraison to 122 μm at week 13, berries from 11 mm to 15 mm, 1.5 g to 2.9 g), moderate deficit producing intermediate expansion (cells to 108 μm, berries to 13.5 mm, 2.3 g), and severe deficit producing minimal expansion (cells to 92 μm, berries to 12 mm, 1.7 g), demonstrating water availability effects on cell expansion rate during early enlargement with deficit irrigation reducing expansion and creating smaller berries with more concentrated composition (severe deficit berries showing 23% sugars at week 13 versus full irrigation 19% through concentration effects). Syrah early enlargement stage berries examined for anthocyanin biosynthesis show maximum expression of anthocyanin biosynthesis genes (VvMYBA1 transcription factor, VvUFGT anthocyanin-3-O-glucosyltransferase) during weeks 9-12 (early enlargement) with 8-15 fold higher expression than at veraison or at maturity, and maximum anthocyanin accumulation rate of 0.4-0.7 mg/g skin per week during early enlargement versus 0.15-0.3 mg/g per week during late enlargement (weeks 14-17), demonstrating that color development is most active during early enlargement phase with subsequent late enlargement showing continued but reduced accumulation. Merlot early enlargement stage berries show maximum turgor-driven expansion with pressure probe measurements revealing cell turgor increasing from 0.4-0.6 MPa at veraison to 0.6-0.9 MPa during early enlargement as sugar and solute accumulation in vacuoles creates osmotic driving force for water influx and cell expansion, with turgor subsequently declining to 0.3-0.5 MPa at maturity as cell walls yield and cells achieve final expanded dimensions.
An early enlargement stage berry is a cell enlargement stage berry during the initial portion of Phase II (BBCH 81-85, approximately 9-13 weeks post-fruit set) characterized by rapid cell expansion following veraison, high growth rates, active sugar accumulation, and active color development, representing berries in the early ripening phase when expansion rate is maximum. Early enlargement stage berries are characterized by= (1) rapid cell expansion from veraison size (60-90 μm diameter) to mid-ripening size (100-130 μm diameter), representing 40-80% diameter increase and 175-380% volume increase during early enlargement (2) cell expansion rate at maximum of 6-14 μm diameter increase per week during early enlargement, declining from this peak as berries progress through mid-to-late enlargement (3) berry diameter increasing rapidly from veraison dimensions (9-13 mm) to mid-ripening dimensions (12-17 mm), representing 25-45% diameter increase during early enlargement (4) berry fresh weight increasing rapidly from veraison weight (1-2 g) to mid-ripening weight (1.8-3.5 g depending on cultivar), representing 60-120% weight increase during early enlargement (5) growth rate at maximum with diameter increasing 0.25-0.6 mm per week and fresh weight increasing 0.15-0.4 g per week during early enlargement phase (6) sugar accumulation rate high at 0.8-1.5% increase per week, progressing from veraison levels (12-17%) to mid-ripening levels (18-23%) (7) anthocyanin accumulation rate (red cultivars) high at 0.3-0.8 mg/g skin per week, progressing from early accumulation (0.5-1.5 mg/g) to substantial levels (1.5-2.8 mg/g). Early enlargement stage berries show active ripening processes= cell wall modification continuing with progressive softening vacuole expansion driving cell enlargement through water accumulation sugar import and accumulation at high rates supporting osmotic potential for water uptake anthocyanin biosynthesis highly active (red cultivars) with rapid color development malic acid respiration active causing rapid acid decline flavor and aroma precursor synthesis beginning. Early enlargement stage berries represent the period of maximum growth vigor during Phase II= high phloem import rates delivering sugars and water high membrane transport activity moving sugars into vacuoles active metabolism supporting ripening processes high sensitivity to water availability with deficit reducing expansion rate and growth. Early enlargement stage berries are highly responsive to irrigation management= adequate water during early enlargement supports maximum cell expansion creating larger final berry size moderate deficit during early enlargement reduces expansion creating smaller more concentrated berries potentially beneficial for wine quality severe deficit during early enlargement can create excessive size reduction and composition imbalances. Environmental factors during early enlargement critically affect berry development trajectory= temperature influences sugar accumulation rate and anthocyanin synthesis with moderate temperatures optimal water availability determines expansion rate and final size potential light exposure affects anthocyanin accumulation in exposed clusters.
Early_Enlargement_Stage_Berry represents rapid expansion phase (weeks 9-13, BBCH 81-85) and is distinguished from Maximum_Enlargement_Stage_Berry (weeks 14-18+, approaching maximum size) by active rapid expansion versus declining expansion approaching completion. Early enlargement represents the period of maximum Phase II growth rate and maximum irrigation effects on berry size. Understanding early enlargement stage biology is critical for irrigation management, for harvest timing prediction based on ripening rate, and for understanding the cellular and biochemical mechanisms of rapid berry expansion and sugar accumulation during early ripening.
Früh-Expansions-Beere
Schnell-Expansions-Stadium
early-ripening berry
frühe Vergrößerungs-Stadium-Beere
rapid-expansion berry
Matthews, M.A., Anderson, M.M. (1988). Fruit ripening in Vitis vinifera= responses to seasonal water deficits. American Journal of Enology and Viticulture 39=313-320 Castellarin, S.D. et al. (2007). Transcriptional regulation of anthocyanin biosynthesis. Plant Physiology 143=1347-1363.
Early Enlargement Stage Berry
Chardonnay exhibits early flowering disposition, reaching 50% capfall approximately 5-10 days before the cultivar median in comparative trials, with budbreak-to-flowering thermal requirement of 320-380 GDD (base 10°C). Pinot Noir and Müller-Thurgau show similarly early flowering, completing anthesis within the first 25% of V. vinifera cultivars. Early flowering in Chardonnay at Burgundy reference sites occurs at mean date of June 5-10 versus June 18-25 for late-flowering cultivars. The budbreak-to-flowering interval in early flowering cultivars averages 55-65 days compared to 70-85 days for late flowering types under identical thermal conditions.
A flowering timing disposition that is the inherited capacity of a Vitis vinifera plant to initiate and complete flowering earlier than average for the species, typically within the first tercile of cultivar bloom dates, reaching anthesis with lower thermal time accumulation from budbreak.
Early flowering disposition may expose vines to suboptimal pollination conditions if bloom coincides with cool, wet spring weather, increasing coulure risk in some climates. However, early flowering extends the post-flowering period available for berry development and ripening, potentially improving fruit quality in short-season regions. Early flowering cultivars show tighter correlation between budbreak and flowering dates (r = 0.85-0.92) than late cultivars, suggesting more deterministic developmental programming.
Frühblüte
early anthesis trait
early bloom
frühe Blüte
früher Blühtermin
precocious flowering
Duchêne E et al. (2010) A grapevine (Vitis vinifera L.) genetic map based on candidate genes. Theoretical and Applied Genetics 120:145-163 Cola G et al. (2014) Description and testing of a weather-based model for predicting phenology, canopy development and source–sink balance in Vitis vinifera L. cv. Barbera. Agricultural and Forest Meteorology 184:117-136
Early Flowering Disposition
Malbec in Mendoza shows early fruit drop during late June-early July Year N+1 at BBCH 69-73, with 18-32% berries aborting and abscising within 2-3 weeks post-flowering from fertilization failures and early seed abortion, peak drop week 2-3, abscission zones forming at pedicels, under variable spring conditions including occasional rain during flowering, typical of this moderately coulure-susceptible cultivar requiring favorable flowering for acceptable fruit set
Early fruit drop represents berry abscission during first 2-4 weeks post-fertilization when young developing berries fail developmentally and abscise from clusters due to inadequate fertilization, seed abortion, insufficient hormonal rescue, or physiological dysfunction, contributing to coulure poor fruit set and reducing berry number from initial flowering, occurring during critical early development period. This developmental failure-induced drop occurs during early summer in late June-July in temperate regions Year N+1, typically within 1-4 weeks following flowering as fertilization outcomes manifest, corresponding to BBCH 69-73 fertilization completion to young fruit stages. Early drop mechanism involves fertilization failure or early seed abortion eliminating auxin source, inadequate hormone production failing to prevent abscission, abscission zone activation at berry pedicel, enzymatic cell wall degradation creating separation layer, and berry detachment and fall reducing cluster berry number. Fertilization failure causes show inadequate pollination preventing fertilization, pollen tube growth failure from boron deficiency or environmental stress, fusion abnormalities preventing zygote or endosperm formation, and complete fertilization failure in all ovules causing inevitable berry abortion. Seed abortion causes show early post-fertilization developmental failure, embryo or endosperm abortion within days to weeks of fertilization, genetic abnormalities preventing normal seed development, environmental stress during critical early period disrupting development, and seed failure eliminating hormonal rescue enabling abscission. Inadequate hormonal rescue shows insufficient auxin production from failed or aborting seeds, hormone levels below threshold required to suppress abscission, auxin-ethylene balance shifting toward abscission promotion, and hormonal failure permitting abscission zone activation. Abscission zone development shows specialized separation cells at pedicel-berry junction activating following hormone signal loss, cell wall degrading enzymes including cellulase and polygalacturonase synthesizing, middle lamella dissolution between cells creating separation plane, vascular tissue severing eliminating nutrient flow, and mechanical weakness enabling berry detachment. Abscission timing shows progressive abscission over 1-4 weeks post-flowering as fertilization failures manifest, peak drop typically 2-3 weeks post-flowering, early failures dropping within days and late failures within weeks, and timing reflecting developmental failure progression. Drop rate dynamics show initial lower drop rate during first week as berries still supported by residual hormones, acceleration during weeks 2-3 as failures accumulate, peak drop rate week 2-3 with daily berry loss visible, and eventual decline as remaining berries stabilize representing successful fruit set. Berry characteristics before drop show small size 4-8 mm diameter indicating limited growth, yellowing or browning suggesting physiological failure, failure to expand distinguishing from growing berries, and pedicel abscission zone visible as swelling or discoloration. Dropped berry examination shows small arrested berries on ground below clusters, internal examination revealing absent or aborted seeds, lack of normal berry development structures, and pathological features distinguishing from normal berries. Cluster appearance after early drop shows gaps from missing berries, pedicel scars indicating abscission sites, remaining berry spacing increased, and overall sparse cluster from high drop typical of poor set. Drop incidence variation shows cultivar differences with coulure-prone cultivars experiencing 30-50% early drop, reliable cultivars maintaining <10-20% drop, and genetic susceptibility as major determinant. Environmental effects include unfavorable flowering weather increasing fertilization failure and drop, rain preventing pollination and fertilization increasing early drop 20-40%, temperature extremes impairing fertilization and seed development, boron deficiency preventing pollen tube growth causing severe drop, and overall adverse conditions during flowering and early fruit set maximizing drop severity. Temperature effects show cool conditions during fertilization potentially increasing failure and drop, excessive heat impairing reproductive function, and optimal 22-28°C minimizing drop risk. Boron deficiency effects show critical importance with inadequate boron causing pollen tube failure, fertilization failure despite adequate flowering, severe early drop 40-60% in deficient conditions, and boron supplementation essential for reducing drop. Rain effects show rain during flowering preventing pollination and washing pollen, poor fertilization success leading to early drop, and rain as major early drop risk factor. Coulure relationship shows early fruit drop as primary coulure mechanism, excessive drop creating poor fruit set <30% set percentage, coulure-prone cultivars and seasons showing severe early drop, and coulure representing extreme early drop scenario. Drop prevention strategies include cultivar selection avoiding coulure-prone varieties, boron fertilization ensuring adequate levels 10-20 ppm during flowering, weather protection during flowering period where feasible, and overall management supporting successful fertilization reducing drop. Early drop versus later drop distinction shows early drop within first month from developmental failure, later drop from different mechanisms including shatter or stress, and timing distinguishing early developmental losses from other drop types. Drop assessment shows berry number counts comparing flowering to 4 weeks post-flowering, drop percentage calculation quantifying loss, cluster examination documenting remaining berry number, and early assessment enabling yield prediction adjustment. The process represents first type of fruit drop caused by developmental failure, distinguished from weather-induced shatter and other drop mechanisms. Duration typically 1-4 weeks post-flowering representing critical failure manifestation period. Regional timing shows early fruit drop during late June-July across climates following flowering. Viticultural significance includes representing primary coulure mechanism reducing fruit set, demonstrating critical importance of successful fertilization and early development, showing cultivar and environmental susceptibility requiring management, and determining final berry number affecting yield. Research applications include analyzing fertilization failure and seed abortion mechanisms, investigating hormonal regulation of early drop, evaluating environmental and nutritional effects on drop incidence, quantifying genetic variation in drop susceptibility, and developing strategies for reducing early drop and improving fruit set.
Early_Fruit_Drop is first type of Fruit_Drop, distinguished by developmental failure within 1-4 weeks post-flowering. Primary coulure mechanism. Results from fertilization failure or seed abortion eliminating hormonal rescue. Timing weeks 1-4 post-flowering. Incidence 10-50% depending on cultivar and conditions. Boron deficiency and weather major causes. Abscission zone activation at pedicel. Reduces berry number determining yield. Prevention through nutrition, cultivar selection, favorable conditions.
Coulure-Beerenverlust
Entwicklungsversagen-Abfall
Post-Blüten-Fall
coulure berry loss
developmental failure drop
early berry abortion
früher Fruchtfall
post-flowering drop
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 410-415 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Early Fruit Drop
An early juvenile unbranched tendril at node 12 on an actively extending shoot of Vitis vinifera cv. Merlot in Pomerol, France, measuring 3.2 cm length at 4 days post-emergence, showing entirely bright green uniform coloration without any tissue differentiation visible, 1.2 mm diameter composed of turgid parenchymatous cells, growing at 11 mm per day based on daily measurements, positioned 5 cm behind the shoot apex in the region of maximum cell division and expansion, showing no response to experimental touch stimulation indicating absent mechanosensory differentiation, procambial strands barely visible as slightly darker green lines within the translucent tissue indicating vascular development not yet begun, occurring on a vigorous shoot (current total length 95 cm) with excellent resource access in well-irrigated and fertilized conditions supporting rapid tendril development, measured at nominal BBCH stage 71 for mature shoot positions though this distal tendril has only recently emerged, representing the universal early juvenile stage through which all tendrils pass during the first week following emergence from apical meristem protection.
A juvenile unbranched tendril in the initial phase of post-emergence development in Vitis vinifera, characterized by very small size typically 2-5 cm length representing 15-30% of eventual mature size, recently emerged from shoot apical meristem region typically within 3-7 days of becoming externally visible, very rapid extension growth at maximum daily elongation rates of 8-15 mm per day consuming substantial resources relative to current biomass, entirely undifferentiated tissue composed predominantly of meristematic and rapidly expanding parenchyma cells without specialized cell types, complete absence of lignification with uniformly bright green coloration and maximum tissue flexibility, no mechanosensory function with cells not yet differentiated into touch-responsive types, minimal vascular development showing only procambial strands without functional xylem or phloem elements, and representing the most vulnerable developmental stage with high sensitivity to stress, damage, or resource limitation that frequently causes abortion if conditions are unfavorable. Early juvenile unbranched tendrils occur universally in tendril development regardless of cultivar or final architecture, representing the initial rapid growth phase following emergence, and require continuous high resource availability particularly carbohydrates for cell division and expansion, nitrogen for protein synthesis and cell proliferation, and water for turgor-driven cell expansion, with stress during this critical period commonly resulting in early aborted tendril outcomes rather than progression to functional maturity.
Early juvenile unbranched tendrils are distinguished from late juvenile unbranched tendrils by very small size (typically <5 cm), very recent emergence (typically <7 days), complete absence of tissue differentiation, and maximum growth rates with no signs of maturation processes beginning. This developmental stage shows highest abortion rates (15-30% in stressed conditions) due to extreme vulnerability of undifferentiated rapidly growing tissues to water deficit, temperature extremes, pest damage, or resource limitation. All tendrils regardless of cultivar or final architecture pass through early juvenile unbranched stage, making this a universal rather than architecturally informative classification. Growth rate, tissue quality, and survival through early juvenile stage serve as sensitive indicators of shoot vigor and resource status, with robust early juvenile development predicting successful maturation while weak or slow early growth often precedes abortion. Environmental protection during this vulnerable stage through appropriate irrigation, fertility, and pest management supports successful tendril development and ultimately climbing function.
früh-juvenile Ranke
initial juvenile tendril
neu-emergierte Ranke
newly emerged tendril
primordial unbranched tendril
primordiale unverzweigte Kletterorgan
Based on Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 60-92 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 219-236.
Early Juvenile Unbranched Tendril
Pinot Noir exhibits early lignification disposition with visible browning of basal internodes by mid-July in Burgundy (approximately 60 days post-budbreak), completing 90% lignification 3-4 weeks before Grenache under identical conditions. Riesling shows early lignification completing by late August in German wine regions, contributing to its exceptional cold hardiness reaching -22°C LT50. Early-lignifying cultivars typically allocate 25-35% more starch reserves to perennial tissues before leaf fall.
A lignification disposition that is the inherited capacity of a Vitis vinifera plant to initiate and complete shoot lignification earlier in the growing season relative to other cultivars, characterized by rapid color change from green to brown, accelerated periderm formation, and early cessation of shoot elongation.
Early lignification correlates strongly with cold hardiness (r = 0.78) and is a key selection criterion for cultivars in continental climates. The trait shows moderate to high heritability (h² = 0.55-0.70) and is influenced by the VvMYB5a and VvMYB5b transcription factors regulating phenylpropanoid biosynthesis. Rootstock influence on scion lignification timing is moderate, with vigor-reducing rootstocks promoting earlier maturation.
early cane maturation
early shoot hardening
frühe Triebausreifung
frühe Verholzung
precocious lignification
vorzeitige Lignifizierung
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Schnabel BJ, Wample RL (1987) Dormancy and cold hardiness in Vitis vinifera L. cv. White Riesling as influenced by photoperiod and temperature. American Journal of Enology and Viticulture 38:265-272
Early Lignification Disposition
Merlot early post-veraison berries at weeks 11-13 post-fruit set (0-2 weeks post-color change, BBCH 81-83) show progression= week 11 (10.5 mm, 1.4 g, cells 72 μm, 15% sugars, 17 g/L acidity, 1.1 mg/g anthocyanins, recently colored red-purple), week 12 (12 mm, 1.9 g, cells 95 μm, 17.5% sugars, 14.5 g/L acidity, 1.8 mg/g anthocyanins), week 13 (13 mm, 2.4 g, cells 110 μm, 19.5% sugars, 12.5 g/L acidity, 2.4 mg/g anthocyanins), demonstrating rapid compositional and size changes during early post-veraison period with sugar increasing 1.25% per week, acids declining 2.25 g/L per week, anthocyanins increasing 0.65 mg/g per week at maximum rates. Syrah early post-veraison berries under two irrigation treatments showing water effects= full irrigation producing rapid expansion (cells from 68 to 115 μm in 3 weeks, berries from 11 to 14.5 mm), moderate sugar concentration (14.5% to 19%) deficit irrigation limiting expansion (cells from 68 to 95 μm, berries from 11 to 12.8 mm), enhanced sugar concentration (14.5% to 20.5%), demonstrating early post-veraison period as critical for irrigation effects on final berry size through cell expansion effects. Pinot Noir early post-veraison berry color development tracking showing anthocyanin accumulation kinetics= week 0 post-veraison (0.9 mg/g, 50% berries colored), week 1 (1.4 mg/g, 80% colored), week 2 (2.1 mg/g, 95% colored), week 3 (2.7 mg/g, 100% colored), with maximum color development rate (0.6-0.7 mg/g per week) during weeks 1-2 of early post-veraison period. Chardonnay white cultivar early post-veraison berries showing flavor precursor development= GC-MS analysis revealing monoterpene precursors (geraniol, linalool glycosides) increasing 3-fold during weeks 0-3 post-veraison, establishing aromatic potential that will be expressed during fermentation, demonstrating early post-veraison as critical period for flavor compound synthesis. Cabernet Sauvignon early post-veraison berry monitoring for harvest prediction= sugar accumulation rate week 11-13 measured at 1.2% per week, predicting maturity at 24% sugars in 6-7 weeks (week 17-18) assuming rate maintenance, allowing early harvest planning and resource allocation, though actual maturity timing at week 18.5 (rate declined to 0.8% per week during late ripening) showing that early predictions require adjustment for late-season rate changes.
An early post-veraison berry is a post-veraison berry in the initial portion of the post-veraison ripening period (BBCH 81-85, approximately 0-4 weeks post-color change), characterized by recent color change completion, active rapid ripening, rapid sugar accumulation, active cell expansion, and progression from veraison composition toward mid-ripening composition, representing berries in early Phase II actively establishing ripening changes. Early post-veraison berries are characterized by= (1) recently completed or recently completing color change with anthocyanins (red cultivars) at 0.8-2.5 mg/g representing early accumulation, or golden-yellow color recently established (white cultivars) (2) sugars at early ripening levels typically 13-20% progressing rapidly from veraison levels (13-17%) toward mid-ripening levels (18-22%) (3) acids at moderately high levels typically 12-18 g/L declining from veraison peak (17-20 g/L) through malic acid respiration (4) cells undergoing rapid expansion from veraison size (60-90 μm) to mid-expansion size (90-120 μm), driving berry enlargement (5) berry diameter increasing from veraison dimensions (9-13 mm) to mid-ripening dimensions (11-16 mm) (6) berry fresh weight increasing from veraison weight (1-2 g) to mid-ripening weight (2-3.5 g) (7) texture actively softening from firm veraison state (2.5-4 N) toward softer ripening state (1.5-2.5 N). Early post-veraison berries show active ripening processes at high rates= sugar accumulation at 0.8-1.5% per week (maximum Phase II rate) cell expansion at 6-12 μm diameter increase per week (maximum expansion rate) acid decline at 0.8-1.5 g/L per week (rapid malic acid respiration) anthocyanin accumulation at 0.3-0.8 mg/g per week (maximum color development rate) flavor compound synthesis initiating with primary fruit aromatics developing. Early post-veraison berries represent critical quality development period= color foundation established through initial anthocyanin synthesis determining final color potential sugar accumulation trajectory established influencing final sugar levels and harvest timing flavor precursors accumulating determining final wine aromatic profile cell expansion determining final berry size with water availability during early post-veraison critically affecting final dimensions. Early post-veraison berries are highly responsive to environmental and management factors= irrigation affecting expansion rate and berry size temperature influencing ripening rate and anthocyanin synthesis (optimal 22-28°C for balanced ripening) light exposure enhancing anthocyanin accumulation in exposed berries crop load affecting resource availability and ripening rate. Early post-veraison berry monitoring provides harvest timing predictions= sugar accumulation rate during early post-veraison (if maintained) predicts maturity timing color development rate indicates anthocyanin synthesis activity and final color potential acid decline rate indicates acid levels at maturity these early indicators allow harvest planning 4-6 weeks in advance though late-season conditions may alter predictions.
Early_Post_Veraison_Berry represents initial ripening phase (0-4 weeks post-color change, BBCH 81-85) and is distinguished from Late_Post_Veraison_Berry (4+ weeks, approaching maturity) by active rapid ripening versus declining ripening rates and approaching maturity. Early post-veraison represents the period of maximum ripening activity and maximum irrigation effects on berry size. Understanding early post-veraison berry development is essential for irrigation management, harvest prediction, and quality development management during the critical early ripening period.
Anfangs-Reife-Traube
Früh-Reifephase-Traube
early Phase II berry
frühe nach-Véraison Beere
initial ripening berry
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 196-215 Castellarin, S.D. et al. (2007). Transcriptional regulation of anthocyanin biosynthesis. Plant Physiology 143=1347-1363.
Early Post Veraison Berry
Pinot Noir exhibits early ripening disposition, reaching 22-24°Brix approximately 40-50 days post-veraison with total season heat requirement of 1100-1300 GDD (base 10°C) from budbreak. Chardonnay and Gamay show similar early ripening, completing harvest in the first third of the regional vintage window. Early ripening in Pinot Noir at Burgundy reference sites occurs at mean date of September 15-25, approximately 3-4 weeks before Cabernet Sauvignon in Bordeaux. The extended post-harvest period (6-10 weeks before leaf fall) in early cultivars facilitates carbohydrate reserve accumulation for following-season fruitfulness.
A ripening timing disposition that is the inherited capacity of a Vitis vinifera plant to reach harvest maturity early in the season, typically within the second quartile of cultivar harvest dates for a region, requiring moderate heat accumulation and providing extended post-harvest vine recovery period before dormancy.
Early ripening disposition confers advantages in cool and moderate climates by ensuring complete maturation before autumn temperature decline and rain events that can compromise fruit quality. The trait correlates moderately with early budbreak (r = 0.50-0.65) and early flowering (r = 0.55-0.70), but genetic independence allows selection for late budbreak with early ripening to minimize frost risk while maximizing ripening reliability. Climate change is shifting early cultivars toward even earlier harvest, potentially advancing into warmer August conditions that may alter wine style.
Frühreife-Disposition
early harvest trait
early maturation
frühe Reife
früher Erntetermin
precocious ripening
Van Leeuwen C et al. (2008) Influence of climate, soil, and cultivar on terroir. American Journal of Enology and Viticulture 55:207-217 Duchêne E, Schneider C (2005) Grapevine and climatic changes: a glance at the situation in Alsace. Agronomy for Sustainable Development 25:93-99
Early Ripening Disposition
An early season prompt lateral shoot on Vitis vinifera cv. Gewürztraminer arising from node 4 axillary bud in late May, only 3 weeks after primary shoot reached this node during rapid spring elongation, developed to 78cm length by late August with 11 leaves totaling approximately 1265 cm² leaf area, representing vigorous early prompt lateral development characteristic of Gewürztraminer's tendency toward lateral production combined with high-vigor site conditions, this lateral emerged during primary shoot active growth (primary still elongating when lateral initiated), extended growing period enabling substantial development creating significant additional photosynthetic capacity but also canopy density concerns, lateral removed during July shoot thinning operation despite photosynthetic potential due to excessive canopy density creating shading problems and disease pressure in already-vigorous canopy, with removal decision reflecting priority for canopy architecture and air circulation over maximum leaf area in this humid climate vineyard.
A prompt lateral shoot emerging early in the growing season shortly after primary shoot node formation, characterized by activation within 2-4 weeks of node development, emergence during active primary shoot elongation phase (typically May-June in Northern Hemisphere), extended growing period enabling substantial lateral development (often 50-90cm length by season end), and potential for significant canopy contribution or density problems depending on vigor, positioning, and management approach. Early season prompt laterals represent rapid axillary activation, arising during peak vine growth when resources are abundant and apical dominance potentially reduced by vigorous overall growth, with their early emergence providing maximum development opportunity creating either valuable photosynthetic supplementation or problematic excessive growth requiring management intervention.
Early season prompt laterals develop during most active vine growth period, with favorable early timing enabling extended development creating both opportunities and challenges. The extended growing period (3-4 months from May-June emergence to September maturity) allows substantial lateral growth potentially exceeding 80cm length and 1000+ cm² leaf area, providing significant photosynthetic supplementation if retained. However, early vigorous laterals also create substantial density increases, potentially severe shading (blocking light to interior canopy and fruit zones), disease pressure (reduced air circulation in dense canopies), and management complexity (requiring positioning, tying, or removal interventions). Early lateral emergence often indicates high vigor conditions (young vines, fertile soils, generous irrigation/nutrition), variety tendencies (certain cultivars prone to early lateral development), or management factors (hedging timing, shoot density). Management typically involves removal in vigorous situations preventing excessive density, possible selective retention in moderate-vigor balanced canopies where lateral benefits outweigh costs, with decisions site-specific based on vigor assessment and canopy objectives.
Frühjahrsnebentrieb
early-emerging lateral
früher prompter Seitentrieb
initial prompt lateral
spring lateral shoot
Based on Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 113-145 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide.
Early Season Prompt Lateral Shoot
Flame Seedless bears an Early_Season_Table_Grape_Role as the benchmark early red seedless, reaching commercial maturity (16-17°Brix, full red color) in late May to early June in California's Coachella Valley, commanding FOB prices of $18-28/box versus $10-14 for mid-season Crimson Seedless, with the early window lasting approximately 3-4 weeks before mid-season volume depresses prices. Sugraone (Superior Seedless) serves in an Early_Season_Table_Grape_Role for premium early white seedless, harvested late May to mid-June in Chile (for Northern Hemisphere winter markets) and Mediterranean regions, with large berries and muscat flavor commanding premium early-market positioning. Perlette bears an Early_Season_Table_Grape_Role as the earliest commercial white seedless in California, reaching 16°Brix by mid-May in the desert production regions, though small berry size limits premium positioning compared to later Sugraone.
A table grape scion role that inheres in a grafted scion cultivar by virtue of its early maturation timing, reaching commercial harvest before the main-season varieties and capturing premium early-market prices when fresh grape supply is limited. Early season table grape scions mature 10-30 days before mid-season benchmarks (typically Thompson Seedless in California, referenced to August 1), reaching harvest-ready condition in May-July in Northern Hemisphere warm production regions (California, Chile counter-season, Mediterranean). Early maturity results from early budbreak, rapid flowering-to-veraison interval, short veraison-to-harvest period, or combinations thereof, determined by genetically controlled heat accumulation requirements (typically 1400-1800 GDD base 10°C to harvest versus 1800-2200 GDD for mid-season varieties). Early season varieties command price premiums of 50-200% above mid-season prices due to limited supply and consumer anticipation of fresh grape availability after winter. Major early season varieties include Flame Seedless, Sugraone (Superior Seedless), Perlette, and numerous proprietary early selections from breeding programs.
Early_Season_Table_Grape_Role captures the market-timing dimension of table grape production where earliness commands significant economic premiums. Production regions with suitable climates (hot deserts, Mediterranean) compete to deliver the earliest fruit, with Coachella Valley (California), Atacama (Chile), and Murcia (Spain) among the earliest suppliers to their respective markets. Early varieties face quality trade-offs—achieving ripeness before full flavor development, color challenges from insufficient heat accumulation, and smaller berry size from compressed growing periods. Breeding programs actively target earliness combined with improved size and flavor, though extending earliness below approximately 1400 GDD approaches physiological limits. Counter-seasonal production (Chile, South Africa, Peru supplying Northern Hemisphere winter) has reduced the premium for early varieties by providing year-round fresh grape availability.
Frühsorte
early harvest grape
early maturing grape
early season grape
early table grape
früh reifende Tafeltraube
frühe Tafeltraube
Creasy GL, Creasy LL (2009) Grapes, CABI Christensen LP (2000) Raisin Production Manual, UC ANR California Table Grape Commission (2020) Grape Variety Harvest Dates Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press
Early Season Table Grape Role
Thompson Seedless exemplifies early seed abortion disposition with embryo collapse occurring 14-21 days post-anthesis, producing seed traces averaging 10-15 mg and 3-4 mm length. Perlette aborts even earlier (10-18 days post-anthesis) with seed traces of only 5-10 mg. Early aborting cultivars produce the smallest seed traces but often have inherently smaller berries, with Thompson Seedless averaging 2.5-3.5 g without gibberellin treatment compared to 4-5 g for later-aborting Crimson Seedless. Consumer acceptance studies show seed traces below 20 mg are imperceptible to 90% of tasters.
A stenospermocarpy disposition that is the inherited capacity of a Vitis vinifera plant to abort developing seeds at an early post-fertilization stage (typically 1-3 weeks after anthesis), resulting in very small, soft seed traces that are minimally perceptible during consumption.
Early seed abortion correlates with reduced berry size due to the loss of seed-derived hormones (auxin, gibberellin) that normally promote pericarp growth. The timing of abortion appears to be controlled by modifier genes in addition to the major sdI locus. Embryo rescue for breeding purposes is more challenging with early-aborting genotypes due to the smaller developmental window for viable embryo recovery.
early embryo abortion
early stenospermocarpy
frühe Embryoabtötung
frühe Samenabortion
kleine Samenrudimente
small seed trace formation
Ramming DW et al. (2000) Seedless grapes. In: Janick J (ed) Fruit Breeding Vol. II: Vine and Small Fruits, Wiley pp.: 197-239 Perl A et al. (2000) Establishment of an Agrobacterium-mediated transformation system for grape (Vitis vinifera L.): the role of antioxidants during grape-Agrobacterium interactions. Nature Biotechnology 14:624-628
Early Seed Abortion Disposition
An early semi-lignified cane on Vitis vinifera cv. Chardonnay examined in early September, showing predominantly green appearance throughout 140cm length with light brown coloration visible at basal 3-4 nodes extending approximately 25cm from base, soft flexible texture when gently bent but showing slightly increased firmness at brown basal section, cross-sectional examination revealing early periderm formation at base with cork cambium activity initiated, secondary xylem development in progress but primary tissues still dominant, representing normal early-autumn developmental status with expectation of progressive lignification advancing apically over subsequent 10-12 weeks reaching full maturation by late November suitable for December pruning retention.
A semi-lignified one-year cane in initial phases of lignification typically occurring during late summer to early autumn (August-September in Northern Hemisphere), characterized by predominantly green appearance with brown coloration just beginning at basal nodes, soft to moderately firm texture throughout most of length, active secondary cell wall development detectable histologically but not yet producing obvious external woody appearance, and normal developmental status for this seasonal timing representing expected progression that will continue toward full maturation over subsequent 8-12 weeks under favorable conditions. Early semi-lignified canes indicate healthy ongoing physiological transition from vegetative growth to dormancy preparation, with lignification initiation triggered by declining photoperiod, moderating temperatures, and internal hormonal shifts redirecting resources from shoot elongation to wood maturation and reserve accumulation.
Early semi-lignified canes are normal and expected during late summer, with their presence indicating appropriate physiological progression toward dormancy. Concerns arise only if environmental conditions (early cold snaps, severe drought, defoliation from disease or pests) interrupt the lignification process before completion. Management during this period focuses on maintaining conditions favorable for continued lignification including adequate but not excessive water (severe stress can halt development), disease and pest control preserving leaf function for carbohydrate production, and avoiding practices that might stimulate renewed vegetative growth (late nitrogen applications, excessive irrigation). The rate of lignification progression from early semi-lignified to fully lignified status depends on thermal conditions, with warm autumns promoting rapid completion and cool autumns slowing progress potentially leading to incomplete maturation.
beginnend ausreifende Rute
beginning ripening cane
early maturation cane
früh-teilverholzte Rute
initial lignification cane
Based on Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 245-267 Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Fennell (2004) "Freezing tolerance and injury in grapevines" Journal of Crop Improvement 10=201-235.
Early Semi Lignified Cane
Early senescence quality assessment for nitrogen deficiency in Walla Walla Valley Merlot documented the impact of premature canopy decline on fruit ripening and wine quality in a nitrogen-limited vineyard. Canopy monitoring through the ripening period tracked early senescence quality progression, with basal leaf senescence (greater than 50% yellow or brown) occurring at 65 days post-véraison in nitrogen-deficient blocks versus 95 days in adequate-nitrogen areas, representing 30-day acceleration of senescence onset. The early senescence quality spatial pattern showed 45% of the canopy showing advanced senescence symptoms at harvest in deficient areas versus 15% in adequate areas, with the reduced green leaf area limiting late-season photosynthesis during the critical sugar accumulation period. Functional impact assessment showed that vines with severe early senescence quality achieved only 23.5°Brix at harvest versus 25.2°Brix in vines maintaining functional canopy, a 1.7°Brix deficit attributed to the approximately 40% reduction in photosynthetically active leaf area during the final 4 weeks of ripening. Wine quality evaluation revealed that early senescence quality affected more than sugar accumulation, with wines from prematurely senesced blocks showing reduced color intensity (12% lower anthocyanins), higher titratable acidity (reflecting delayed acid degradation), and less developed fruit character in sensory evaluation. The early senescence quality assessment established that maintaining adequate nitrogen to prevent premature canopy decline was essential for achieving target ripeness, even though modest nitrogen limitation was acceptable earlier in the season for vigor management purposes.
Early senescence quality in nitrogen deficiency is a nitrogen deficiency symptom quality that inheres in grapevine leaves and represents the premature initiation and acceleration of leaf aging processes when nitrogen remobilization from mature leaves to support new growth depletes these source leaves below the threshold for maintained function, with nitrogen-deficiency-induced senescence producing earlier-than-normal leaf yellowing, necrosis, and abscission that reduces the functional canopy duration and photosynthetic capacity during the critical fruit ripening period. This quality captures the accelerated aging dimension of nitrogen deficiency, recognizing that the remobilization of nitrogen from older leaves represents both a survival mechanism allowing continued growth under nutrient limitation and a cost in terms of reduced functional leaf area that may compromise fruit development and reserve accumulation. The ontological significance of early senescence quality in nitrogen deficiency lies in its impact on late-season vine function when sustained photosynthesis is critical for fruit ripening and carbohydrate storage, with premature senescence potentially causing incomplete ripening, reduced sugar accumulation, and inadequate reserve storage affecting subsequent season performance. The biological basis of nitrogen-deficiency senescence encompasses the programmed cell death pathways activated when nitrogen remobilization reduces cellular nitrogen below critical thresholds, the degradation of chloroplasts and their protein content including the abundant Rubisco enzyme that constitutes a major nitrogen reservoir, and the export of remobilized nitrogen as amino acids transported to developing tissues through the phloem. The visual progression of nitrogen-deficiency senescence shows characteristic patterns with initial chlorosis as chlorophyll degrades, followed by yellowing intensification as remaining pigments are catabolized, then necrosis as cell death occurs, and finally abscission leaving the vine with reduced functional canopy during late ripening. The timing of senescence onset relative to fruit development stage determines the functional impact, with early senescence during véraison potentially limiting ripening capacity while senescence delayed until post-harvest has minimal production impact.
Early_Senescence_Quality_(N) is_a Nitrogen_Deficiency_Symptom_Quality, representing the premature leaf aging and abscission resulting from nitrogen remobilization to support new growth. This class addresses the canopy-duration dimension of nitrogen deficiency. Senescence timing relative to fruit development determines functional impact. The progression from chlorosis through yellowing to necrosis and abscission follows nitrogen remobilization. Late-season photosynthetic capacity affects sugar accumulation and complete ripening. Distinguishing seasonal management needs (moderate early deficiency acceptable vs. late-season adequacy critical) informs nitrogen timing decisions.
Frühe Seneszenz bei N-Mangel
N-deficiency senescence
accelerated leaf aging
beschleunigte Blattseneszenz
early leaf drop
früher Blattfall
premature leaf aging
vorzeitige Blattalterung
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Lim PO et al. (2007) Leaf senescence. Annual Review of Plant Biology 58:115-136 Bell SJ, Henschke PA (2005) Implications of nitrogen nutrition for grapes, fermentation and wine. Australian Journal of Grape and Wine Research 11:242-295
Early Senescence Quality (N)
Chardonnay in Champagne shows early shoot elongation from late April through mid-May at BBCH 15-19 over 3-4 weeks, with shoots extending from 180 mm at 5 leaves to 550-650 mm at 9 leaves through elongation rates progressing from 10-14 mm per day initially to 16-20 mm per day as photosynthetic capacity develops, under cool maritime climate conditions of 14-18°C mean temperature, adequate spring rainfall, and moderate fertility typical of this moderately vigorous early-bursting cultivar
Early shoot elongation represents the initial growth phase where shoots begin extending beyond leaf unfolding stage, characterized by relatively slow elongation rates as photosynthetic capacity develops and shoots transition from complete dependence on stored reserves toward autotrophic growth. This developmental stage occurs during late April through May in temperate regions corresponding approximately to BBCH 15-19 as shoots develop 5-9 unfolded leaves, with elongation rate typically 8-18 mm per day creating total shoot length increase from 150-200 mm to 400-700 mm over 2-4 week period. Early elongation shows progressive acceleration as each additional expanding leaf contributes photosynthetic capacity, with shoots transitioning from net carbon consumers during initial phase to approaching carbon neutrality and eventually positive carbon balance when 4-5 leaves achieve substantial expansion and photosynthetic function. The apical meristem maintains high activity producing new leaf primordia every 2-3 days and contributing cells to shoot axis, with meristem structure showing organized tunica-corpus organization and cell division rates of 20-24 hour cycles characteristic of active growth. Internode lengths during early elongation typically range 25-45 mm reflecting moderate growth rate and developing resource availability, with internodes progressively lengthening as growth accelerates toward rapid elongation phase. Cell expansion in subapical zones proceeds through water uptake and wall loosening, with expansion zone extending 15-25 mm behind apex containing cells increasing from meristematic dimensions to final lengths of 100-300 micrometers depending on internode position and growth conditions. Leaves continue unfolding and expanding sequentially with new leaves separating from axis every 3-5 days, with leaf sizes progressively increasing from basal leaves toward middle shoot positions reflecting improving resource status as photosynthetic area accumulates. Photosynthetic capacity increases progressively during early elongation as leaf area expands and chloroplasts mature, with whole-shoot net photosynthesis transitioning from negative to positive typically around 4-5 leaf stage when photosynthetic carbon fixation begins exceeding respiratory carbon consumption. Carbon sources during early elongation include primarily stored carbohydrates mobilized from woody tissues in declining amounts as reserves deplete, and increasingly photosynthate from expanding leaves as photosynthetic capacity develops, with transition from stored to current assimilate occurring progressively over 2-3 week period. Vascular system extends through elongating shoot with xylem differentiation maintaining functional water transport capacity of 1-3 mg per second supporting expansion and transpiration, and phloem development enabling import of stored reserves initially and bidirectional transport as photosynthetic production begins. Water demand increases substantially during early elongation as transpiring leaf area expands from 100-200 square mm with 4-5 leaves to 500-1200 square mm with 8-9 leaves, requiring shoot water uptake rates increasing from 30-60 mg per hour to 100-200 mg per hour depending on environmental conditions. Nutrient requirements intensify particularly for nitrogen supporting protein synthesis in rapidly expanding tissues, with nitrogen concentration in young leaves 3-5% dry weight and substantial nitrogen import from storage reserves and root uptake supporting growth demands. Metabolic activity remains elevated with respiration rates in expanding tissues 10-15 fold above mature levels supporting growth processes, with respiratory carbon loss declining as proportion of photosynthetic gain as carbon balance improves. Protein synthesis continues at high rates producing structural proteins for expanding cells, photosynthetic proteins accumulating in developing leaves reaching 40-50% total leaf protein as Rubisco and other Calvin cycle enzymes concentrate in chloroplasts, and enzymes supporting rapid metabolism. Hormonal regulation maintains elevated auxin from shoot apex coordinating elongation and apical dominance, gibberellins supporting cell expansion and internode elongation, and cytokinins from roots promoting cell division in meristematic tissues and leaf expansion. Temperature strongly affects early elongation rate with development approximately doubling per 10°C increase between 15-28°C optimal range, with cool spring conditions often limiting early growth while warm conditions accelerate development. Light intensity affects early elongation through effects on photosynthetic capacity development, with high light promoting rapid photosynthetic maturation supporting accelerating growth while shade reduces photosynthetic gain potentially prolonging early elongation phase. Water availability must be adequate to support turgor maintaining cell expansion, with spring water stress potentially limiting early elongation rate and delaying transition to rapid growth impacting final canopy development. Nutrient availability particularly nitrogen affects early elongation rate through effects on protein synthesis capacity and photosynthetic development, with adequate nutrition supporting rapid acceleration toward peak growth while deficiency prolongs slow early phase. The process typically spans 2-4 weeks under favorable conditions with consistent temperatures above 16°C and adequate moisture and nutrition, with suboptimal conditions extending duration or limiting acceleration toward rapid elongation. Cultivar variation appears in early elongation rate reflecting inherent vigor differences, with vigorous cultivars showing more rapid acceleration toward peak growth while less vigorous types maintain modest rates longer. Regional timing shows early elongation occurring late April through May in warm climates, May through early June in temperate regions, and late May through mid-June in cool areas, with duration depending on spring temperature patterns and resource availability. Viticultural management considerations include ensuring adequate water availability during early elongation supporting transition to rapid growth, providing sufficient nitrogen supporting protein synthesis and photosynthetic capacity development without excessive promotion of vigor, monitoring shoot development uniformity assessing vine health and resource distribution, and planning canopy management operations based on elongation progression. Limitations during early elongation including water stress, nutrient deficiency, or pest damage can permanently reduce final canopy size affecting productivity. Research applications include analyzing transition from heterotrophic to autotrophic growth documenting carbon balance dynamics, quantifying photosynthetic capacity development relative to growth requirements, evaluating environmental effects on early growth patterns affecting final canopy development, and assessing hormonal regulation coordinating transition from establishment to rapid growth phase.
Early_Shoot_Elongation is first of five Shoot_Elongation subphases, distinguished by modest elongation rates and transition from stored reserves to photosynthetic carbon sources. Transitions to Rapid_Shoot_Elongation as photosynthetic capacity becomes established and growth accelerates. Represents critical establishment phase determining subsequent growth potential. Carbon balance transition during this phase fundamentally affects growth capacity.
Etablierungsphase
beginnende Längenwachstum
beginning elongation phase
early vegetative growth
establishment growth
frühe Triebstreckung
initial shoot extension
initiale Triebverlängerung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 205-220 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 128-140 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Early Shoot Elongation
An early stage reproductive active bud on Vitis vinifera cv. Riesling at BBCH stage 15 in early May in the Mosel Valley, Germany, showing 3-5 leaves unfolding, two compact inflorescences visible as green elongated structures 12-15mm in length enclosed within developing shoot, individual flowers not yet separated, highly vulnerable to forecast frost event of -2.5°C requiring active frost protection through wind machines, heaters, or overhead sprinklers to prevent catastrophic yield loss.
A reproductive active bud in initial phases of development corresponding approximately to BBCH stages 09-53, characterized by bud swelling and scale separation revealing both vegetative and reproductive primordia, emergence of shoot with woolly appearance, first leaves unfolding, and inflorescences visible as compact clusters tightly enclosed within shoot apex, not yet showing individual flower separation. Early stage reproductive active buds are extremely vulnerable to frost damage which can result in complete crop loss, with critical temperatures typically -2°C to -3°C for developing inflorescences, slightly more cold-tolerant than vegetative tissues at similar developmental stages due to higher sugar concentrations in floral tissues providing some cryoprotection.
Early stage reproductive active buds represent the period of maximum economic risk in viticulture, as frost damage at this stage results in near-complete crop loss for the affected shoots with minimal compensation from secondary buds which typically have reduced or no inflorescence content. The short window between budbreak and flower separation (typically 3-4 weeks) coincides with late spring frost season in many cool-climate regions, making frost protection a critical management requirement. Inflorescence primordia at this stage are already fully formed, having developed during the previous growing season, with current growth primarily involving cell expansion rather than new flower initiation.
early flowering bud
frühe reproduktive aktive Knospe
initial stage fruiting bud
Based on Molitor et al. (2014) "Late frost damage risk for viticulture under future climate conditions= a case study for the Luxembourgish winegrowing region" Australian Journal of Grape and Wine Research 20=160-168 Rodrigo (2000) "Spring frosts in deciduous fruit trees - morphological damage and flower hardiness" Scientia Horticulturae 85=155-173 Mosedale et al. (2015) "Climate change impacts and adaptive strategies= lessons from the grapevine" Global Change Biology 21=2562-2579.
Early Stage Reproductive Active Bud
An early stage vegetative active bud on Vitis vinifera cv. Chardonnay at BBCH stage 05 in early April in Chablis, France, showing bud scales separated revealing green woolly shoot tip 4mm in length, first leaf scales visible but not yet unfolded, occurring 5 days after initial bud swell, with frost protection measures activated when nighttime temperatures forecast below -1°C due to high vulnerability of tissues at this developmental stage.
A vegetative active bud in the initial phase of growth activation and shoot emergence, corresponding approximately to BBCH stages 01-09, characterized by bud swelling, scale separation, emergence of green shoot tip with woolly appearance due to dense trichomes, and first leaf beginning to unfold but not yet fully separated from shoot apex. Early stage vegetative active buds are maximally vulnerable to spring frost damage due to tender, water-rich tissues with minimal lignification and limited cold hardening capacity, with lethal temperatures often as high as -2°C to -4°C depending on development stage, cultivar cold tolerance, and acclimation status following previous temperature fluctuations.
The early stage represents the period of maximum frost risk in viticulture, as buds transition from dormant cold-hardy state (tolerating -15°C to -25°C) to actively growing frost-sensitive state (damaged at -2°C to -4°C) over a period of only 2-3 weeks. Cultural practices including delayed pruning, double pruning, and application of dormancy-extending compounds aim to delay early stage budbreak past the typical frost season. Once damage occurs at early stage, secondary buds may break to compensate, though with delayed development and typically reduced fruitfulness.
early breaking vegetative bud
frühe vegetative aktive Knospe
initial stage vegetative bud
Based on Trought et al. (1999) "The impact of differences in soil texture within a vineyard on vine vigour, vine earliness and juice composition" Journal of Wine Research 10=77-84 Ferguson et al. (2011) "Dynamic changes in bud cold hardiness during dormancy and the influence of apple and pear budbreak" Journal of the American Society for Horticultural Science 136=313-323 Molitor & Junk (2019) "Climate change is implicating a two-fold impact on air temperature increase in the ripening period under the conditions of the Luxembourgish grapegrowing region" OENO One 53=409-422.
Early Stage Vegetative Active Bud
Chasselas exhibits early veraison disposition, reaching 50% color change approximately 45-50 days post-flowering with thermal requirement of 680-780 GDD (base 10°C) from bloom. Pinot Noir and Chardonnay show similarly early veraison, initiating ripening within the first 25% of V. vinifera cultivars in multi-variety trials. Early veraison in Chasselas at Swiss reference sites (Pully) occurs at mean date of July 25-30 versus August 20-28 for late cultivars. The extended post-veraison period in early cultivars (55-70 days versus 40-50 days for late cultivars in the same climate) allows gradual flavor development and tannin maturation under cooler autumn conditions.
A veraison timing disposition that is the inherited capacity of a Vitis vinifera plant to initiate ripening earlier than average for the species, typically within the first tercile of cultivar veraison dates, reaching the onset of color change and softening with lower thermal time accumulation from flowering.
Early veraison disposition provides advantages in cool or short-season climates by maximizing the ripening period before autumn temperature decline and frost risk. The trait correlates moderately with early flowering (r = 0.65-0.75) but shows independent genetic control, enabling selection for early veraison without necessarily early budbreak vulnerability. Early veraison cultivars benefit from extended hang time for phenolic maturity while achieving sugar targets, though excessively early veraison in warm climates may accelerate ripening during peak summer heat, reducing aromatic complexity.
Frühfärbung
early color change trait
early ripening onset
frühe Véraison
früher Reifebeginn
precocious veraison
Parker AK et al. (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17:206-216 Duchêne E et al. (2012) Phenology and grape ripening in relation to climate change. Climate Research 53:145-156
Early Veraison Disposition
Chardonnay grafted on Teleki 5C rootstock ring 8 showing early wood rootstock xylem comprising inner 0.38 cm of 0.64 cm total ring width (59% of ring), earlywood appearing light tan contrasting with darker latewood, earlywood vessels 110-145 micrometers diameter significantly larger than latewood vessels 55-75 micrometers, earlywood showing thin cell walls and large lumens optimized for water transport, earlywood density estimated 0.42 g/cm³ versus latewood 0.61 g/cm³, earlywood zone containing approximately 65-70% of ring's total water transport capacity despite occupying only 59% of width due to large vessel flow advantage
Early wood rootstock xylem is the xylem tissue produced during early growing season typically April-June in Northern Hemisphere forming inner lighter-colored portion of annual ring, characterized by large-diameter vessels 80-180 micrometers optimized for maximum water transport capacity, thin-walled cells maximizing lumen volume for conductivity, lower wood density than latewood due to large void spaces, rapid formation during peak cambial activity supporting spring shoot growth and leaf expansion, comprising typically 40-60% of annual ring width, and providing majority of annual water transport capacity despite occupying only portion of ring due to water flow rate increasing with fourth power of vessel radius making large earlywood vessels disproportionately important for hydraulic function. Early wood rootstock xylem formation timing includes= initiated in spring (cambium reactivating April-May as temperatures rise and photoperiod lengthens), produced during early season (majority of earlywood formed May-June during most rapid cambial activity), corresponding to shoot growth (earlywood formation coinciding with bud break, shoot elongation, and leaf expansion requiring maximum water supply), showing peak production rate (cambium dividing most rapidly during earlywood phase producing cells at maximum rate), and transitioning to latewood (earlywood formation declining July-August as cambium shifts to producing smaller-vessel latewood completing annual ring). Early wood rootstock xylem structural characteristics includes= showing large vessel diameter (vessels 80-180 micrometers significantly larger than latewood vessels 40-80 micrometers optimizing water transport), exhibiting thin cell walls (vessel elements and surrounding cells with minimal secondary wall deposition maximizing lumen, wall ratio), displaying low wood density (earlywood 0.35-0.50 g/cm³ lower than latewood 0.55-0.75 g/cm³ due to large lumens and thin walls), appearing light colored (earlywood light tan or cream in cross-section due to large void spaces and minimal wall material), and showing gradual or abrupt transition (earlywood-latewood boundary potentially gradual with progressive vessel size reduction or sharp with sudden change depending on growing conditions). Early wood rootstock xylem vessel characteristics includes= containing largest vessels in ring (earlywood vessels reaching maximum diameter for rootstock cultivar enabling peak water flow rates), showing high vessel density (numerous large vessels per unit area maximizing total conducting area), exhibiting optimal perforation plates (large openings in vessel element end walls minimizing resistance to water flow), displaying efficient pit connections (abundant lateral pits connecting vessels to adjacent cells facilitating water distribution), and demonstrating vulnerability to cavitation (large vessels potentially more susceptible to embolism formation under water stress compared to smaller latewood vessels due to larger diameter). Early wood rootstock xylem hydraulic function includes= providing majority of water transport (earlywood conducting 60-80% of total annual water flow despite comprising only 40-60% of ring width due to large vessel advantage), enabling peak transpiration (earlywood capacity adequate for maximum canopy water loss during hot periods supporting photosynthesis and growth), showing highest conductivity per area (earlywood hydraulic conductivity per unit cross-sectional area exceeding latewood due to large vessels), supporting spring growth (earlywood formation timing coinciding with water demand peak from expanding shoots and leaves), and demonstrating flow efficiency (large earlywood vessels minimizing resistance enabling high flow rates with minimal pressure gradient). Early wood rootstock xylem in annual ring position includes= occupying inner portion (earlywood forming first part of annual ring immediately outside previous year's latewood), showing distinct inner boundary (sharp line marking transition from previous year dark latewood to current year light earlywood creating visible annual ring boundary), exhibiting variable width (earlywood radial extent 0.2-0.6 cm varying with spring growing conditions and cambial activity duration), transitioning to latewood (earlywood giving way to latewood at mid-ring position marking shift from spring to late summer wood production), and comprising ring fraction (earlywood typically 40-60% of total ring width with proportion varying by year and conditions). Early wood rootstock xylem seasonal water demand includes= supporting bud break (earlywood formation and function coinciding with bud expansion requiring increased water supply), enabling shoot growth (earlywood conducting water for rapidly expanding shoots during peak elongation phase May-June), supplying leaf expansion (earlywood transport supporting developing leaves as leaf area increases exponentially early season), providing flowering water (earlywood delivering water for inflorescence development and bloom), and facilitating early fruit development (earlywood conducting water for initial berry cell division and growth phases). Early wood rootstock xylem formation conditions includes= reflecting spring moisture (earlywood width and vessel size indicating water availability during early season with adequate moisture producing wide earlywood), showing temperature effects (favorable spring temperatures supporting vigorous cambial activity producing abundant earlywood), exhibiting nutrient influences (adequate spring nitrogen availability supporting rapid cell division creating extensive earlywood), demonstrating vigor effects (vigorous trunk producing broader earlywood zone with larger vessels than low-vigor trunk), and recording stress impacts (spring frost, early drought, or disease reducing earlywood production creating detectable anomalies). Early wood rootstock xylem cultivar variation includes= showing variety-specific vessel sizes (Berlandieri hybrids with large earlywood vessels 120-180 micrometers, Riparia with moderate vessels 80-110 micrometers, cultivar genetics determining maximum earlywood vessel diameter), exhibiting different earlywood proportions (some rootstocks showing broad earlywood zones 50-60% of ring while others showing narrower zones 40-50%), and demonstrating vigor-related patterns (vigorous rootstocks like 1103P producing extensive earlywood with very large vessels supporting high water transport capacity).
Early_Wood_Rootstock_Xylem emphasizes spring-formed large-vessel portion of annual ring and is distinguished from Late_Wood_Rootstock_Xylem (summer-formed small-vessel portion) by focus on inner ring position, large vessels, thin walls, and primary water transport function. Critical for understanding seasonal xylem production, hydraulic conductivity, spring water supply, vessel size effects on transport, and the light-colored inner ring zone providing majority of annual water conduction capacity.
Earlywood
Frühholz-Xylem
Frühjahrsholz
Großlumiges Unterlagen-Xylem
Large-vessel rootstock xylem
Spring wood rootstock xylem
Early Wood Rootstock Xylem
Syrah early wood xylem vessel in ring 10 shows 210-280 μm diameter, thin secondary wall 2.5-3.8 μm thickness, formed April-May during spring canopy expansion, arranged in 2-3 tangential rows forming conspicuous earlywood band occupying 38-46% of ring width, individual vessels providing hydraulic conductivity 15-35 times greater than average latewood vessel, earlywood zone providing 68-76% of total ring water transport capacity, vessels showing 45-65% embolism by late summer but restored by next spring's new earlywood formation
An early wood xylem vessel (earlywood vessel) is a xylem vessel formed during the early portion of the growing season (typically April-June in Northern Hemisphere temperate climates), characterized by large diameter (120-300 μm), thin walls, and high water transport capacity, forming the distinctive inner band of large vessels in ring-porous wood that provides majority of annual ring hydraulic conductance. Early wood vessel formation timing includes= differentiation beginning with spring cambial reactivation (typically late March-April corresponding to budbreak timing), production during initial 6-10 weeks of growing season (April-June primary earlywood formation period), formation synchronized with leaf expansion (large vessels supporting water demand from rapidly growing canopy), cessation as latewood formation begins (transition to smaller vessels typically June-July), and earlywood zone completion representing 25-45% of total annual ring width despite being only 35-45% of formation duration. Early wood vessel anatomical characteristics include= large diameter (150-300 μm typical, compared to 40-120 μm latewood vessels), thin secondary walls (wall thickness 2-4 μm compared to 3-6 μm in latewood vessels), large lumens (lumen occupying 140-290 μm of 150-300 μm diameter), simple perforation plates (large single openings in end walls providing minimal resistance to axial flow), abundant bordered pits (50-80 pits per 100 μm² wall area enabling extensive lateral connections), and arrangement in tangential bands (1-3 rows of large vessels forming conspicuous earlywood zone). Early wood vessel hydraulic function includes= providing 60-80% of annual ring water transport capacity (despite being 35-50% of ring area, large vessels dominate conductance), maximum specific conductivity (individual large vessels transporting 10-100 times more water than small vessels due to radius⁴ relationship), supporting spring canopy development (earlywood vessels providing water for rapid shoot and leaf expansion April-June), enabling peak summer transpiration (earlywood conductance supporting maximum water use 3-6 L/m² leaf area/day), and critical for vine water status (earlywood vessel functionality determining vine hydraulic capacity). Early wood vessel embolism vulnerability includes= high susceptibility to cavitation (large diameter vessels more vulnerable due to increased pit membrane area and higher tensions), preferential embolism during drought (water deficit-induced cavitation affecting earlywood vessels first and most extensively), freeze-thaw embolism risk (large vessels particularly vulnerable to ice formation disrupting water columns), progressive seasonal loss (earlywood vessels often showing 40-70% embolism by late summer compared to 20-40% in latewood), and catastrophic failure in extreme stress (severe drought potentially causing 80-95% earlywood vessel embolism causing hydraulic crisis). Early wood vessel structural characteristics include= relatively weak mechanical support (thin walls providing less bending and compression resistance than thick-walled fibers or latewood vessels), vulnerability to crushing (large thin-walled vessels potentially collapsing under mechanical stress or negative pressure), dependence on surrounding tissues (fibers and thick-walled cells providing structural support preventing vessel collapse), and optimized for transport not support (vessel evolution favoring maximum conductance even at cost of reduced mechanical strength). Early wood vessel development and differentiation includes= differentiation from early cambial divisions (spring cambial reactivation producing large fusiform daughter cells becoming earlywood vessel precursors), rapid cell expansion (initial fusiform derivatives undergoing dramatic radial expansion achieving large diameters), secondary wall deposition (lignified wall formation with relatively thin walls optimized for conductance), programmed cell death (protoplasm breakdown creating hollow lumen), and rapid maturation (vessels completing differentiation and becoming functional within 2-4 weeks of cambial production). Early wood vessel significance in ring-porous pattern includes= creating visible earlywood band (concentration of large vessels producing distinctive inner ring zone lighter in color and coarse in texture), defining ring-porous classification (Vitis exhibiting classic ring-porous pattern with earlywood-latewood vessel size distinction), enabling ring boundary identification (transition from small latewood vessels to large earlywood vessels marking annual ring boundaries), and affecting wood appearance (earlywood vessels creating characteristic porous texture on transverse surfaces). Early wood vessel numbers and density include= 15-35 earlywood vessels per mm² (lower density than latewood but much larger individual size), 1-4 tangential rows of large vessels (rows forming continuous or interrupted bands across earlywood zone), total earlywood vessels comprising 20-35% of total ring vessel number (but providing 60-80% of conductance), and earlywood zone occupying 35-50% of ring area (high vessel lumen fraction 45-65% of earlywood area). Early wood vessel implications for viticulture includes= earlywood conductance determining canopy potential (adequate large vessel production enabling full canopy development, reduced earlywood formation limiting leaf area), earlywood embolism affecting vine water stress (loss of large vessel function causing severe impact disproportionate to vessel number), annual earlywood renewal restoring capacity (new earlywood formation each spring providing fresh conducting vessels replacing previous year's embolized vessels), and earlywood characteristics varying by vigor (vigorous growth producing wider earlywood zones with more/larger vessels, stress reducing earlywood vessel size and number).
Early_Wood_Xylem_Vessel emphasizes spring-formed large vessels and is distinguished from Late_Wood_Xylem_Vessel (summer-formed smaller vessels), Large_Diameter_Xylem_Vessel (size emphasis without seasonal context), and Xylem_Vessel (all vessels regardless of position). Critical for understanding ring-porous wood pattern, seasonal hydraulic capacity dynamics, spring water supply for canopy development, embolism vulnerability patterns, and annual hydraulic capacity renewal through earlywood formation.
Earlywood vessel
Frühholzgefäß
Frühjahrsgefäß
Großes Leitgefäß
Large vessel
Spring wood vessel
Tyree, M. T., & Zimmermann, M. H. (2002). Xylem Structure and the Ascent of Sap (2nd ed.). Springer. | Carlquist, S. (2001). Comparative Wood Anatomy (2nd ed.). Springer.
Early Wood Xylem Vessel
Earthy aroma profiling in Gevrey-Chambertin Pinot Noir identifies the characteristic sous-bois expression comprising forest floor (6.8 on 10-point scale), truffle (4.5), and subtle wet stone (4.2), creating the terroir signature that distinguishes Burgundy from New World Pinot Noir where fruit typically dominates over earth, with these earthy notes valued as evidence of site expression that justifies premium pricing despite scientific uncertainty about their direct geological origin.
Earthy aroma quality is an aroma character quality that represents olfactory perception evoking soil, mineral, and geological substrate-associated scents, characterizing the complex, often savory aromatic impression that contributes to perceived terroir expression and distinguishes site-specific wines from generic fruit-driven examples. Within the BFO framework, this quality addresses a major aroma category distinguished by its association with the perception of place rather than fruit or winemaking, recognizing that earthy notes create the impression of vineyard specificity and mineral substrate expression valued in terroir-focused wine evaluation. The ontological significance lies in earthy aroma serving as the primary sensory evidence for terroir perception, where earthy character creates the impression that wine expresses its geological origin regardless of whether direct mineral transport from soil to wine occurs, making earthy descriptors central to the communication of site-specificity that defines fine wine identity. The compositional basis for earthy aroma remains incompletely characterized despite extensive research, with potential contributors including geosmin (petrichor, beetroot character at sub-ng/L threshold), 2-methylisoborneol (musty-earthy), and various sulfur compounds and thiols that may contribute mineral or flinty perception, though the exact chemical basis for minerality perception remains scientifically debated. The perceptual psychology of earthy character involves complex cognitive associations, where terms like minerality may reflect non-olfactory attributes (acidity, texture, saltiness) or learned expectations about prestigious wine origins rather than specific volatile compound perception, creating challenges for objective earthy aroma assessment. The regional associations of earthy character are strong, with Burgundy Pinot Noir expected to show forest floor and sous-bois notes, Chablis Chardonnay prized for flinty minerality, and Loire Valley Chenin Blanc valued for chalky or wet stone character, creating style templates that define quality expectations for these origins. Understanding earthy aroma quality enables terroir expression assessment based on perceived geological character, regional typicity evaluation where earthy notes indicate site influence, and quality judgment incorporating the complexity that earthy character adds to wine aromatic profile.
Earthy_Aroma_Quality serves as the parent class for earth-related subcategories (Mineral, Flinty, Petrichor), representing an aroma character category central to terroir perception and fine wine evaluation. This quality encompasses perceptions whose chemical basis remains debated, requiring acknowledgment of the gap between sensory experience and scientific understanding. The class addresses the geological association dimension of wine character distinct from fruit, floral, and spicy categories.
Bodennote
Erdige Aromen
Terroir-Duft
geological scent
mineral character
mineralischer Charakter
soil-like notes
terroir aroma
Heymann H et al. (2014) A study of the perceptions of minerality in Sauvignon blanc wines. Journal of Wine Research 25(4): 259-273 Ballester J et al. (2013) Mapping perceived minerality in wines. Food Quality and Preference 28(1): 340-350
Earthy Aroma Quality
A Riesling vine in east-west rows on steep Rheingau slopes bears an East_West_Row_Role with south-facing exposure essential for ripening at 50°N latitude, the orientation maximizing solar interception during Germany's limited growing season. A Grüner Veltliner vine in east-west terraces of the Wachau serves in an East_West_Row_Role dictated by the Danube Valley's east-west trend, with careful canopy management moderating the exposure asymmetry between south and north canopy faces. A Pinot Noir vine in east-west rows in cool Burgundy sites bears an East_West_Row_Role traditional to the region, where south-facing exposure has historically been valued for ripening assistance in marginal conditions.
A row orientation role that inheres in a grapevine planted in rows aligned along the east-west axis (or approximately east-west within ±30°), creating north-facing and south-facing canopy exposures with substantially different sunlight reception depending on hemisphere. In the Northern Hemisphere, east-west rows position one canopy side facing south (receiving substantially more solar radiation throughout the day) while the north-facing side remains more shaded the asymmetry is reversed in the Southern Hemisphere. East-west orientation is often preferred at high latitudes (>45°) where the low sun angle means south-facing exposure is essential for adequate heat accumulation and ripening, or on east-west trending slopes where contour alignment prevents erosion. The asymmetric exposure creates predictable differences in fruit composition between canopy sides—the sun-facing side typically shows earlier ripening, higher sugar accumulation, more color development, and greater heat stress risk, while the shaded side ripens later with higher acidity retention. East-west orientation may be deliberately chosen where the differentiated fruit provides blending options, or may result from constraints (slope direction, property geometry) that prevent preferred north-south alignment.
East_West_Row_Role represents the orientation favored for cool-climate viticulture and slope-constrained situations, creating the south-facing exposure (Northern Hemisphere) that maximizes heat accumulation where ripening is challenging. The exposure asymmetry inherent to east-west rows requires management consideration—the sun-facing side may require different leaf removal, irrigation, or harvest timing than the shaded side—and the fruit differentiation may be exploited (separate harvest of sides for different programs) or averaged (combined harvest accepting the blend of ripeness levels). East-west orientation on slopes is often mandated by erosion control requirements contour planting with rows following elevation lines prevents row-channeled runoff regardless of the resulting compass orientation. Climate change is shifting east-west optimization, as warming reduces the need for maximum sun exposure at sites where ripening was previously marginal, potentially favoring north-south conversion where practical.
E-W orientation
O-W-Ausrichtung
Ost-West-Zeilen
east-west alignment
east-west rows
latitudinal orientation
latitudinale Orientierung
Smart RE (1973) Sunlight interception, American Journal of Enology and Viticulture 24: 141-147 Hunter JJ et al. (2016) Row orientation effects, South African Journal of Enology and Viticulture 37: 107-119 Intrieri C et al. (1998) Row orientation and grape composition, Vitis 37: 83-88
East West Row Role
St. George (Rupestris du Lot, V. rupestris) demonstrates exceptional rooting capacity (90-95% success) reflecting V. rupestris's inherent easy-rooting nature, producing abundant roots from single-node cuttings and enabling economical propagation despite the cultivar's otherwise limited commercial use. 3309 Couderc (V. riparia × V. rupestris) shows easy rooting (85-90%) inherited from V. rupestris, with nurseries routinely achieving 85%+ success rates using standard hardwood cutting protocols. Ramsey (Salt Creek, V. champinii) roots easily (85-90%), facilitating its widespread commercial adoption for nematode-resistant rootstock production in California and Australia. Freedom (1613C × Dog Ridge) inherits easy rooting (85-90%) from Dog Ridge, contributing to its commercial success.
A rooting capacity rootstock role that inheres in a grafted rootstock by virtue of its high capacity for adventitious root initiation from hardwood cuttings, achieving rooting success rates exceeding 80% under standard nursery propagation conditions without specialized treatments. Easy-rooting rootstocks develop abundant root primordia rapidly (visible root emergence within 3-4 weeks under optimal conditions), show low cutting mortality during callusing, and produce vigorous, well-branched root systems supporting strong first-year growth. This high rooting capacity reflects favorable endogenous hormone balances (high auxin sensitivity, low cytokinin), abundant ray parenchyma for dedifferentiation, optimal carbohydrate reserves, and efficient wound-induced cell division. Easy-rooting rootstocks offer economic advantages through reduced propagation costs, reliable nursery supply, and the ability to use shorter cuttings (single-node or two-node) that multiply planting material more efficiently. V. rupestris parentage is the primary genetic source of easy rooting.
Easy_Rooting_Rootstock_Role captures rootstocks achieving greater than 80% rooting under standard conditions, representing the economically optimal propagation category. V. rupestris and V. champinii parentage consistently contribute easy rooting, while V. riparia × V. rupestris hybrids typically root easier than pure V. riparia selections. Easy rooting enables single-node cutting propagation (maximizing multiplication rate), reduces losses during callusing and nursery establishment, and ensures reliable commercial supply. The 80% threshold reflects industry standards where losses below 20% are economically acceptable without special interventions.
easy rooting rootstock
easy to propagate rootstock
gut bewurzelbare Unterlage
high rooting capacity rootstock
leicht bewurzelbare Unterlage
vermehrungsfreudige Unterlage
Hartmann HT et al. (2011) Hartmann and Kester's Plant Propagation 8th Ed, Pearson Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Whiting JR (2004) Grapevine rootstocks, in Viticulture Vol 1 Resources, Winetitles pp.: 167-188 Dry N (2007) Grapevine rootstocks: selection and management for South Australian vineyards, Lythrum Press
Easy Rooting Rootstock Role
In Pinot Noir vines of the Côte d'Or, Burgundy, ecodormancy extends from early February when chilling requirements are satisfied through late March when spring warming triggers budbreak, with buds demonstrating growth competence through rapid budbreak within 10 days under forcing conditions at 22°C while remaining quiescent in the field where mean temperatures of 5-8°C during February and early March are insufficient to satisfy heat unit requirements for growth initiation.
Ecodormancy in Vitis vinifera represents the phase of winter dormancy during which growth inhibition is controlled by environmental factors external to the bud, with buds having regained growth competence through completed chilling but remaining quiescent due to unfavorable environmental conditions, particularly low temperatures. This dormancy phase follows endodormancy and typically extends from late winter through early spring in Northern Hemisphere viticulture, persisting until warming temperatures satisfy the heat requirements for growth initiation and budbreak commences. The defining characteristic of ecodormancy is the external nature of growth control, distinguishing it from endodormancy where internal factors prevent growth regardless of environmental conditions, with ecodormant buds capable of immediate growth response when transferred to favorable conditions while endodormant buds remain inhibited even under optimal forcing. The physiological state during ecodormancy differs fundamentally from endodormancy in that all dormancy-maintaining mechanisms have been dismantled through chilling accumulation, growth-related genes have been derepressed and are competent for transcription, hormone balances have shifted toward growth-permissive configurations with low ABA and responsive cytokinin and gibberellin signaling, and cellular communication pathways have been restored enabling coordinated growth responses. The ecological function of ecodormancy is to delay growth until environmental conditions are reliably favorable, with temperature serving as the primary constraint that prevents budbreak during late winter when frost risk remains high despite the restoration of growth competence. The duration of ecodormancy varies substantially depending on climate, with longer ecodormancy periods in cold continental climates where spring warming is delayed, and shorter periods in Mediterranean climates where mild temperatures may follow closely after chilling completion. During ecodormancy, buds undergo preparatory changes for growth including gradual metabolic activation, accumulation of growth-related transcripts, and ultrastructural changes in meristematic cells that prepare for rapid division upon warming. The concept of heat unit or growing degree day accumulation during ecodormancy quantifies the temperature exposure required for budbreak initiation, with cultivar-specific heat requirements determining the timing of growth resumption relative to spring temperature patterns. The transition from ecodormancy to active growth is triggered when accumulated heat units satisfy the requirement for budbreak, initiating the visible swelling and eventual opening of buds that marks the beginning of the new growing season.
Ecodormancy represents the third major phase within the Dormancy hierarchy following Endodormancy, characterized by environmentally-controlled rather than internally-controlled growth inhibition. Distinguished from Endodormancy by the external nature of growth control and immediate growth response under forcing conditions. Subdivides into Ecodormancy_Transition, Environmental_Constraint, Conditional_Dormancy, and Quiescence_Maintenance as component phases. Duration varies from weeks to months depending on climate. Growth competent buds await heat accumulation for budbreak initiation.
Ekodormanz
Umweltdormanz
environmental dormancy
erzwungene Ruhe
imposed dormancy
quiescence
temperaturbedingte Dormanz
temperature-limited dormancy
Ecodormancy
Ecodormancy duration assessment comparing contrasting springs in Riesling reveals the temperature-dependent variation in vulnerability period length, with cold spring (endodormancy released February 1, temperatures remaining below 10°C until March 25, budbreak April 8) showing long ecodormancy duration of 66 days with extended frost vulnerability but ultimately successful budbreak, while warm spring (endodormancy released January 28, temperatures exceeding 10°C consistently from February 20, budbreak March 15) showing short ecodormancy of only 46 days but with frost damage on March 22 when late freeze followed early budbreak, demonstrating how ecodormancy duration affects the nature and timing of frost risk rather than eliminating it.
Ecodormancy duration quality is an ecodormancy status quality that represents the length of time between endodormancy release and visible budbreak, characterizing the period during which buds are physiologically ready for growth but environmentally prevented from growing by temperatures below the threshold for active development, with duration depending on both the timing of endodormancy release and the subsequent temperature pattern through late winter and early spring. Within the BFO framework, this quality specifically addresses the temporal extent dimension of ecodormancy, distinguishing how long this phase lasts from the depth of dormancy or the thresholds for transition while recognizing that ecodormancy duration represents the waiting period between achieving growth competence and actually initiating growth, a period of vulnerability during which warm spells can trigger growth that subsequent cold may damage. The ontological significance lies in ecodormancy duration determining the frost risk exposure window, where long ecodormancy (from early chilling completion or late spring) creates extended vulnerability to warm spell-induced premature budbreak, while short ecodormancy (from late chilling completion or early consistent spring) minimizes the period of risk by reducing the time between growth competence and appropriate conditions for sustained growth. The factors affecting ecodormancy duration include the timing of endodormancy release (earlier release creating potential for longer ecodormancy), the temperature pattern of late winter and spring (consistent cold extending ecodormancy, early warming shortening it), and cultivar heat requirements (high heat requirements extending ecodormancy even after temperatures become marginally favorable by requiring more accumulation before budbreak). The year-to-year variation in ecodormancy duration can be substantial, with cold springs maintaining ecodormancy for months after endodormancy release while warm springs may compress ecodormancy to mere weeks, creating the vintage-specific budbreak patterns that contribute to growing season timing variation and frost risk differences among years. Understanding ecodormancy duration quality enables assessment of the frost vulnerability window, prediction of when the risk period will end with budbreak, and recognition of the factors that extend or compress ecodormancy with implications for spring frost management strategies.
Ecodormancy_Duration_Quality addresses the time period between endodormancy release and budbreak representing the frost vulnerability window. This quality varies with release timing and subsequent temperatures. The class supports frost risk assessment through ecodormancy duration characterization.
Nachruheperiode
Umweltdormanzlänge
Vor-Austrieb-Periode
ecodormancy length
environmental dormancy duration
post-rest period
pre-budbreak period
Ökodormanzdauer
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 59-144 Ferguson JC et al. (2014) Dynamic thermal time model of cold hardiness for dormant grapevine buds. Annals of Botany 113(5): 801-811
Ecodormancy Duration Quality
Ecodormancy status assessment in late winter Cabernet Sauvignon reveals the growth-ready state of post-chilling buds, with forcing tests showing rapid, uniform budbreak (95% within 10 days at 20°C) confirming complete endodormancy release and ecodormancy establishment, and field observations showing immediate response to warm spell (5 consecutive days above 15°C triggering visible bud swell within 7 days), followed by frost damage when temperatures returned to -4°C (30% of swelling buds killed), demonstrating how ecodormancy status indicates full growth competence creating vulnerability to the temperature fluctuations common in late winter and early spring.
Ecodormancy status quality is a dormancy depth quality that represents the environmentally imposed quiescence of grapevine buds that have completed endodormancy release and are physiologically competent for growth but remain dormant due to unfavorable temperatures, characterizing the post-chilling dormancy phase during which buds await adequate warmth to initiate visible growth. Within the BFO framework, this quality specifically addresses the environmental control dimension of late dormancy, distinguishing ecodormancy from the internally controlled endodormancy while recognizing that ecodormant buds are fundamentally different from endodormant buds in their responsiveness—ecodormant buds will grow immediately when exposed to favorable temperatures, while endodormant buds remain inhibited regardless of temperature until chilling is satisfied. The ontological significance lies in ecodormancy representing the final barrier to growth that is entirely under environmental control, where the duration of ecodormancy depends solely on when temperatures become favorable for growth, with this period providing protection against early-season frost only through the environmental constraint of cold temperatures rather than through any internal growth inhibition, making ecodormant buds vulnerable to growth initiation during any sufficiently warm period regardless of calendar date. The physiological state during ecodormancy involves buds that have regained full growth competence, with hormonal balances, chromatin states, and metabolic readiness restored to growth-permissive conditions, but with the external temperature constraint preventing the actual initiation of cell division and expansion that would produce visible budbreak. The management implications of ecodormancy status include the recognition that late-winter warm spells may trigger growth of ecodormant buds creating frost vulnerability, that practices delaying the onset of ecodormancy responsiveness (such as late pruning) may provide frost protection by slowing the transition to full growth competence, and that heat accumulation models become relevant for budbreak prediction only after ecodormancy is established. Understanding ecodormancy status quality enables assessment of bud readiness for spring growth, prediction of budbreak timing from temperature forecasts once ecodormancy is established, and recognition of the frost vulnerability created by the fully responsive state of ecodormant buds awaiting only adequate warmth to initiate growth.
Ecodormancy_Status_Quality serves as the parent class for heat-related qualities (Requirement, Duration). This quality addresses the environmentally controlled dormancy phase following endodormancy release. The class supports frost risk assessment and budbreak prediction through ecodormancy status characterization.
aufgezwungene Dormanz
environmental quiescence
post-rest dormancy
temperaturlimitierte Dormanz
Ökodormanzstatus
Lang GA et al. (1987) Endo-, para-, and ecodormancy: physiological terminology and classification for dormancy research. HortScience 22(3): 371-377 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 59-144
Ecodormancy Status Quality
In Chardonnay vines in Champagne, ecodormancy transition occurs during the first two weeks of February when accumulated chilling hours reach approximately 1000, demonstrated by forcing tests showing budbreak time decreasing from 35 days in late January to 12 days by mid-February, with this transition marking the shift from internal dormancy control to environmental temperature limitation that will persist until March warming initiates budbreak.
Ecodormancy transition in Vitis vinifera represents the physiological shift from internally-controlled endodormancy to environmentally-controlled ecodormancy, occurring when sufficient chilling has been accumulated to satisfy the chilling requirement and dismantle dormancy-maintaining mechanisms while environmental temperatures remain too low to permit growth initiation. This transition typically occurs during late January to February in Northern Hemisphere viticulture, marking a fundamental change in the nature of dormancy from a state where growth is impossible regardless of conditions to a state where growth is merely delayed by unfavorable environment. The transition involves the completion of all chilling-dependent physiological changes including reduction of ABA to non-inhibitory levels, cessation of DAM gene expression, removal of repressive chromatin modifications from growth gene promoters, and restoration of symplastic communication through callose degradation at plasmodesmata, collectively restoring the capacity for growth that was suspended during endodormancy. The practical identification of ecodormancy transition relies on forcing tests that demonstrate rapid budbreak response indicating growth competence, with the transition point defined as the time when forcing response times decrease to less than approximately 14 days under optimal conditions, contrasting with the extended times characteristic of endodormant buds. The transition is not instantaneous but represents the crossing of a threshold where dormancy-maintaining mechanisms have been sufficiently weakened that they can no longer prevent growth under favorable conditions, with individual buds potentially crossing this threshold at slightly different times creating a brief period of transition heterogeneity across the vine. Environmental conditions during ecodormancy transition affect subsequent budbreak characteristics, with continued cold exposure during early ecodormancy potentially improving budbreak uniformity while premature warming might advance budbreak into periods of elevated frost risk. The metabolic state following ecodormancy transition involves gradual reactivation of biosynthetic pathways in preparation for growth, though full metabolic activation awaits the warming temperatures that will trigger budbreak, with ecodormant buds maintaining intermediate metabolic rates between the minimal activity of deep endodormancy and the high activity of actively growing tissues.
Ecodormancy_Transition represents the phase shift from endodormancy to ecodormancy within the dormancy cycle. Distinguished from Chilling_Requirement_Satisfaction which marks the quantitative chilling achievement, while Ecodormancy_Transition emphasizes the qualitative change in dormancy control mechanism. Subdivides into Endodormancy_to_Ecodormancy_Shift, Environmental_Control_Onset, and Growth_Competence_Restoration as component aspects. Identified through forcing test response less than 14 days. Duration of transition phase typically spans 1-2 weeks.
Dormanzphasen-Übergang
Ekodormanz-Übergang
Endodormanz-Ekodormanz-Wechsel
Ruhetyp-Transition
dormancy phase shift
dormancy type transition
endodormancy to ecodormancy transition
quiescence onset
Ecodormancy Transition
Old-vine Carignan in the Corbières appellation of southern France fulfills an ecological role through its extensive root system stabilizing steep schist slopes against erosion, its permanent woody structure providing nesting habitat for passerines including Sylvia melanocephala and Serinus serinus, its unpruned canopy margins supporting overwintering populations of predatory mites (Typhlodromus pyri) and parasitoid wasps (Anagrus atomus), and its deep litter layer beneath bush-trained vines contributing 2-4 t/ha/year of organic matter to soil carbon pools.
A grapevine role that inheres in a vine by virtue of its functional contribution to the ecological processes, biodiversity dynamics, and ecosystem services operating within and surrounding the vineyard landscape, capturing the vine's participation in habitat provision, nutrient cycling, carbon dynamics, and soil conservation beyond its primary agricultural production function.
Ecological_Role recognizes that grapevines function as components of complex agroecosystems extending beyond their primary viticultural purpose. This role class is increasingly relevant in the context of biodiversity-integrated viticulture, ecosystem service valuation, and agroecological transition frameworks: the EU Biodiversity Strategy 2030 and Common Agricultural Policy reform explicitly recognize the ecological functions of permanent crop systems including vineyards. Subclasses capture specific ecosystem service categories — biodiversity support, cover crop integration, agroforestry, carbon sequestration, and soil conservation — reflecting the multifunctional ecological contributions of vineyard systems.
Weinberg-Ökologie-Rolle
ecological function role
ecosystem service role
vineyard ecology role
Ökosystemfunktions-Rolle
ökologische Rolle
Viers JH et al. (2013) Vinecology: pairing wine with nature, Conservation Letters 6: 287-299 Winkler KJ et al. (2017) Biodiversity in vineyards, Agriculture Ecosystems and Environment 239: 44-57 Paiola A et al. (2020) Exploring the potential of vineyards for biodiversity conservation, Conservation Biology 34: 1092-1104
Ecological Role
Economic data for Grüner Veltliner in Austria integrates production statistics (approximately 13,500 ha planted, yielding approximately 850,000 hL annually, representing roughly 30% of Austrian wine production by volume), market positioning data (price spectrum from €3-5/bottle for entry-level Qualitätswein through €15-30 for DAC and Federspiel/Smaragd to €50-150+ for single-vineyard Smaragd crus from top Wachau estates), export data (key export markets Germany, Switzerland, United States, with growing presence in Japan and Scandinavia), and comparative economic performance (Grüner Veltliner generating the highest aggregate revenue of any Austrian cultivar due to the combination of substantial volume and increasingly premium price positioning). Economic data for Pinot Noir globally documents the cultivar's exceptional economic polarization — global acreage approximately 115,000 ha producing wines spanning from €5 commodity Pinot Noir to the €5,000-25,000+ per bottle commanded by Grand Cru Burgundy, with the price variance ratio between entry and apex exceeding that of any other cultivar.
Economic data is a grapevine information entity that documents the quantitative economic dimensions of grape cultivar cultivation and wine production, encompassing production statistics (acreage, yield, volume), market data (pricing, trade flows, consumption patterns), and economic performance indicators (profitability, cost structures, return on investment), derived from governmental statistical agencies, international organizations (OIV, FAO, Eurostat), national wine industry bodies, and commercial market research, providing the economic-quantitative foundation for understanding the commercial significance, market positioning, and financial viability of individual cultivars and the viticultural sectors that produce them.
Economic data documentation provides the quantitative commercial framework that contextualizes the biological, viticultural, and oenological information documented elsewhere in VVO — transforming descriptive cultivar knowledge into commercially actionable information by establishing the economic dimensions of cultivar significance. The economic data domain intersects with virtually every other VVO information entity hierarchy — production statistics reflect the biological productivity characteristics documented in yield data and phenological records, market pricing reflects the quality assessments documented in sensory descriptions and quality ratings, trade flows reflect the regulatory frameworks documented in trade regulations and label requirements, and economic trends reflect the historical trajectories documented in historical importance records. The principal data sources for viticultural economic data operate at different scales and with different methodologies — the OIV provides global aggregates and country-level statistics based on member country submissions, Eurostat provides harmonized EU-level data, national statistical agencies provide detailed domestic production and trade data, and commercial market research firms (IWSR, Wine Intelligence, Rabobank Wine Research) provide proprietary market analysis and consumer insight data. The challenge of economic data comparability across jurisdictions is substantial — national statistical classifications, reporting methodologies, vintage-year versus calendar-year accounting, currency effects, and varying levels of data disaggregation (some countries report cultivar-specific production data while others report only aggregate wine production) limit the precision of cross-national economic comparisons. The temporal dimension of economic data is essential for understanding cultivar market dynamics — point-in-time statistics provide snapshots but may be misleading without trend context, as a cultivar's economic significance may be rapidly expanding, stable, or declining depending on market dynamics, climate adaptation pressures, and regulatory changes.
Marktdaten
Weinwirtschaftsdaten
Wirtschaftsdaten
commercial data
economic indicators
economic information
market data
viticultural economics
wirtschaftliche Kennzahlen
ökonomische Daten
OIV (2023) State of the World Vine and Wine Sector, International Organisation of Vine and Wine Anderson K, Nelgen S (2020) Which Winegrape Varieties are Grown Where? Revised edition, University of Adelaide Press Eurostat (2023) Vineyard statistics, European Statistical Office IWSR (2023) Global Wine Market Analysis
Economic Data
Effervescence quality comparison between méthode traditionnelle Champagne and Charmat-method Prosecco reveals production-driven differences across all bubble dimensions, with Champagne showing fine bubbles (2/10 size, where 1 is finest), exceptional persistence (9/10, streams lasting minutes in glass), and creamy mousse (8/10 quality), while Prosecco displays larger bubbles (5/10), moderate persistence (5/10, diminishing within minutes), and frothier mousse (5/10), demonstrating how effervescence character reflects production method and enables quality differentiation within sparkling wine categories.
Effervescence quality is a tactile quality that represents the sensory perception of dissolved carbon dioxide releasing as bubbles during wine consumption, characterizing the fizzing, tingling, and foaming sensations that define sparkling wine texture and contribute to freshness perception in slightly effervescent still wines. Within the BFO framework, this quality addresses the carbonation dimension of tactile perception, distinguishing effervescence from other mouthfeel qualities while recognizing that bubble-related sensations involve both mechanical stimulation (bubbles bursting on oral surfaces) and chemesthetic components (CO₂ activating sour receptors and trigeminal neurons). The ontological significance lies in effervescence defining the fundamental character distinction between sparkling and still wines, where bubble presence, quality, and behavior create the textural experience that distinguishes Champagne from still wine more decisively than any flavor difference, making effervescence assessment essential for sparkling wine quality evaluation. The physiological basis for effervescence perception involves multiple sensory mechanisms, with mechanical stimulation from bubble formation and bursting on oral surfaces, chemesthetic activation of trigeminal neurons by dissolved CO₂, and stimulation of acid-sensing receptors contributing to the overall fizzing, tingling sensation that characterizes effervescent wines. The compositional determinants of effervescence include dissolved CO₂ concentration (determining bubble vigor), wine surface tension (affecting bubble formation and stability), nucleation site availability (controlling where bubbles form), and temperature (affecting CO₂ solubility and release rate). The quality dimensions of effervescence span intensity (from pétillant through frizzante to fully sparkling), fineness (bubble size from coarse to fine), persistence (duration of effervescence), and integration (how bubbles contribute to overall mouthfeel), with different sparkling wine styles showing characteristic profiles across these dimensions. Understanding effervescence quality enables sparkling wine assessment based on bubble character, production method inference from effervescence properties, and quality evaluation incorporating appropriate bubble quality for wine style.
Effervescence_Quality serves as the parent class for specific bubble qualities (Size, Persistence, Mousse), establishing the carbonation dimension of tactile assessment. This quality involves both mechanical (bubble bursting) and chemesthetic (CO₂ receptor activation) components. The class supports sparkling wine assessment and production method inference through comprehensive effervescence evaluation.
Kohlensäureempfindung
Mousseux-Qualität
Perlage
bubble sensation
carbonation quality
fizzing character
sparkling texture
sprudelnde Textur
Liger-Belair G (2005) The physics and chemistry behind the bubbling properties of Champagne and sparkling wines. Journal of Agricultural and Food Chemistry 53(8): 2788-2802 Pozo-Bayón MA et al. (2009) Influence of carbonation on aroma release from liquid systems. Journal of Agricultural and Food Chemistry 57(5): 1663-1668
Effervescence Quality
Cabernet Franc in Loire Valley shows egg cell approach during late June Year N+1 at BBCH 65-69, with sperm cells released from tube apex in synergid migrating 30-80 micrometers to egg cell at micropylar pole within 15-90 minutes, establishing membrane contact preparing for fusion, under temperate maritime conditions of 22-27°C temperature, typical of this mid-season cultivar completing approach 28-42 hours post-pollination enabling zygote formation
Egg cell approach represents the sperm cell positioning phase following tube discharge when one sperm cell migrates or drifts through embryo sac cytoplasm from discharge site in penetrated synergid toward egg cell at micropylar pole, achieving proximity and contact preparing for membrane fusion and zygote formation. This critical sperm positioning phase occurs during ovule penetration in mid-late June in temperate regions Year N+1, typically proceeding within minutes to hours following tube discharge, corresponding to BBCH 65-69 fertilization stages. Approach mechanism involves sperm cell release from ruptured tube apex into synergid and embryo sac cytoplasm, passive drift or active migration through cytoplasm toward egg cell positioned adjacent to discharge site, short-distance movement typically <100 micrometers given egg proximity to synergid, and arrival at egg cell surface establishing contact for fusion. Sperm cell characteristics show haploid nucleus containing paternal genetic material, minimal cytoplasm optimized for fusion with cytoplasmic material largely excluded from gamete, non-motile cell lacking flagella requiring passive transport or cytoplasmic streaming for movement, and maintained integrity during discharge and approach maintaining fusion competence. Egg cell position shows location at micropylar pole of embryo sac immediately adjacent to synergid cells, proximity to tube discharge site facilitating rapid sperm arrival, and positioning optimizing fertilization efficiency through minimal required sperm travel. Distance and timing of approach show short travel distance typically 20-100 micrometers from synergid discharge site to egg cell surface, rapid approach completed within minutes to hours depending on cytoplasmic dynamics, and brief interval minimizing time between discharge and fusion reducing risk of sperm degradation. Cytoplasmic environment for sperm movement includes embryo sac cytoplasm providing aqueous medium, possible cytoplasmic streaming or flow affecting sperm transport, and chemical or physical gradients possibly guiding sperm toward egg though mechanisms poorly characterized. Sperm guidance to egg possibly involves chemotactic signals from egg cell attracting sperm, calcium gradients directing movement, physical constraints in embryo sac architecture channeling sperm toward egg, or passive drift enabled by close proximity, though specific mechanisms in grapevine largely unknown. Egg cell surface contact shows sperm arrival at egg cell plasma membrane, initial contact establishing opportunity for membrane fusion, and positioning of sperm nucleus adjacent to egg preparing for fusion. Egg cell structure at approach includes large haploid nucleus containing maternal genetic contribution, abundant cytoplasm with organelles and stored reserves, plasma membrane receptive to sperm fusion, and developmental arrest state awaiting fertilization activation. Sperm-egg recognition possibly involves surface molecules mediating species-specific recognition, compatibility assessment though self-compatibility in grapevine eliminates rejection, and molecular interactions preparing for membrane fusion. Membrane interaction preparation shows sperm and egg plasma membranes in close contact, possible adhesion molecules mediating attachment, and membrane modifications preparing for fusion including lipid reorganization and protein recruitment. Calcium signaling during approach possibly includes calcium waves in egg cytoplasm triggered by sperm contact, calcium-dependent processes regulating fusion, and calcium as second messenger coordinating fertilization events though specific roles incompletely understood. Approach verification methods include microscopic observation of embryo sac following pollination though technically challenging, fluorescent labeling of sperm cells enabling tracking though rarely applied in grapevine, and indirect inference from successful fertilization outcomes. Sperm fate determination shows one sperm cell destined for egg fusion while sibling targets central cell, differentiation possibly occurring during discharge or through stochastic selection upon release, and specific mechanisms determining which sperm fuses with which target unknown. Second sperm simultaneous approach to central cell shows parallel process with other discharged sperm migrating toward large central cell, coordinated double fertilization requiring both sperm cells reaching appropriate targets, and synchronization ensuring embryo and endosperm development initiate together. Environmental effects on approach include temperature affecting cytoplasmic dynamics and membrane properties, appropriate hydration maintaining cellular integrity and function, and overall favorable conditions 22-28°C supporting efficient approach and fusion. Failed approach causes include sperm cell damage during discharge compromising integrity or viability, abnormal cytoplasmic environment preventing movement or causing degradation, egg cell dysfunction preventing normal reception, or structural abnormalities in embryo sac disrupting approach pathway. Approach success rate shows high efficiency given close proximity between discharge site and egg, majority of successfully discharged sperm cells reaching egg under normal conditions, and approach success contributing to overall 50-70% fertilization rate in successful penetration events. The process represents first aspect of ovule penetration positioning sperm for fusion, followed by pollen tube discharge discussing release mechanism and fusion. Duration typically brief minutes to hours representing sperm travel and positioning phase. Cultivar variation in egg cell approach may include embryo sac structure affecting sperm transit, cytoplasmic properties affecting movement dynamics, and egg cell characteristics affecting reception. Regional timing shows egg cell approach occurring during mid-late June across all climates within hours of tube discharge 24-48 hours after pollination. Viticultural significance includes representing essential sperm positioning enabling zygote formation, demonstrating efficiency of fertilization mechanism through minimal required sperm travel, and establishing approach success as prerequisite for fusion and embryo development. Research applications include analyzing sperm movement mechanisms and guidance, investigating egg cell signals and sperm-egg recognition, evaluating factors affecting approach efficiency, developing visualization techniques for observing approach in vivo, and assessing approach success as determinant of fertilization outcomes.
Egg_Cell_Approach is first aspect of Ovule_Penetration, distinguished by sperm cell positioning for zygote formation. Represents short-distance migration <100 micrometers from discharge site to egg. Duration minutes to hours post-discharge. High efficiency given close proximity. Enables subsequent sperm-egg fusion creating diploid zygote. Parallel process with second sperm approaching central cell. Essential prerequisite for embryo development. Followed by Pollen_Tube_Discharge discussing release and fusion mechanisms.
Eizell-Annäherung
Gameten-Begegnung
Präfusions-Positionierung
Spermium-Eizell-Heranführung
gamete encounter
pre-fusion positioning
sperm cell positioning
sperm-egg approach
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 393-395 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 298-300
Egg Cell Approach
Chardonnay in Champagne shows egg fertilization during late June-early July Year N+1 at BBCH 65-69, with sperm cell fusing with egg cell within 1-3 hours post-discharge occurring 28-42 hours post-pollination, membrane fusion and nuclear combination producing diploid 2n=38 zygote, calcium signaling activating developmental program, under cool maritime conditions of 20-25°C temperature, typical of this early-bursting cultivar with 64-76% egg fertilization success initiating embryo development
Egg fertilization represents the zygote-forming fusion event in double fertilization where one sperm cell released from pollen tube fuses with egg cell at embryo sac micropylar pole, combining haploid sperm nucleus with haploid egg nucleus forming diploid 2n zygote nucleus containing equal maternal and paternal genetic contributions, and activating egg developmental program initiating embryogenesis. This critical embryo-forming fusion occurs during double fertilization in mid-late June in temperate regions Year N+1, typically completing within 1-4 hours following sperm release approximately 24-48 hours after initial pollination, corresponding to BBCH 65-69 fertilization stages. Fertilization mechanism involves sperm cell migration from tube discharge site in penetrated synergid to adjacent egg cell positioned at micropylar pole, sperm-egg membrane fusion creating cytoplasmic continuity, sperm nucleus entry into egg cytoplasm and migration toward egg nucleus, nuclear envelope breakdown in both gamete nuclei, chromatin mixing and nuclear fusion combining genetic material, diploid 2n zygote nucleus formation containing complete chromosome sets from both parents, and egg activation triggering developmental transition from arrest to active embryogenesis. Sperm cell approach to egg shows short-distance migration typically 20-100 micrometers from synergid discharge site, passive drift or directed movement through embryo sac cytoplasm, and rapid arrival within minutes to hours given close proximity facilitating efficient fertilization. Egg cell structure at fertilization includes large haploid nucleus containing maternal genetic contribution, abundant cytoplasm with organelles and stored reserves providing resources for early embryo development, plasma membrane receptive to sperm fusion showing molecular recognition capacity, and developmental arrest in metaphase of second meiotic division awaiting fertilization activation. Sperm-egg recognition shows surface molecule interactions mediating species-specific recognition, molecular complementarity between sperm coat proteins and egg membrane receptors, compatibility assessment though self-compatibility in grapevine eliminates rejection mechanisms, and recognition signals preparing for membrane fusion. Membrane fusion initiation shows sperm cell contacting egg plasma membrane, lipid bilayer reorganization at contact site enabling membrane merger, fusion pore formation creating small cytoplasmic bridge, and pore expansion integrating sperm completely into egg cytoplasm. Fusion pore dynamics show initial small opening 20-50 nanometers diameter, progressive expansion through membrane flow and cytoskeletal forces, complete membrane integration within seconds to minutes, and cytoplasmic continuity establishing allowing sperm nucleus entry. Sperm nucleus entry shows nucleus entering egg cytoplasm through fusion site, minimal sperm cytoplasm transferred with nucleus entering leaving most paternal cytoplasm and organelles excluded, and sperm nucleus integrity maintained during entry enabling subsequent nuclear fusion. Sperm nucleus migration shows movement through egg cytoplasm toward centrally positioned egg nucleus, migration possibly driven by cytoskeletal forces and cytoplasmic streaming, travel distance typically 10-50 micrometers depending on egg size, and arrival at egg nucleus within minutes to hour following membrane fusion. Nuclear fusion process shows sperm and egg nuclei coming into close contact, nuclear envelope breakdown in both nuclei releasing chromatin into common cytoplasmic space, chromatin mixing and condensation combining maternal and paternal chromosomes, nuclear envelope reformation around combined genetic material, and diploid 2n nucleus formation containing complete haploid chromosome sets from both parents. Diploid zygote nucleus characteristics show 2n chromosome number typically 38 chromosomes in Vitis vinifera combining 19 maternal and 19 paternal chromosomes, equal genetic contribution from both parents enabling Mendelian inheritance, and functional diploid nucleus capable of directing embryo development. Genetic recombination through fertilization shows meiotic recombination during gamete formation creating genetic variation, random fertilization combining different parental gametes, and offspring genetic diversity supporting evolutionary adaptation and breeding applications. Egg activation following fusion shows calcium signaling waves propagating through cytoplasm triggered by sperm entry, calcium-dependent processes activating developmental programs, cortical granule exocytosis modifying membrane preventing polyspermy, metabolic activation with respiratory rate increasing 2-3 fold, protein synthesis initiation producing developmental regulators, and cell cycle reentry transitioning from meiotic arrest to mitotic division. Calcium signaling dynamics show initial localized elevation at sperm entry point, propagating waves spreading throughout egg cytoplasm at 5-20 micrometers per second, multiple calcium transients possibly occurring over minutes to hours, and calcium-dependent kinases activating downstream signaling cascades triggering development. Cortical reaction shows cortical granules beneath egg plasma membrane undergoing exocytosis releasing contents modifying membrane properties, block to polyspermy preventing additional sperm fusion, and rapid reaction within seconds to minutes of first sperm entry protecting against abnormal polyploid development. Metabolic activation shows transition from quiescent to active state with oxygen consumption increasing substantially, ATP production intensifying supporting biosynthesis and division, protein synthesis initiating from maternal mRNAs accumulated during oogenesis, and metabolic reorientation preparing for embryo development. Cell cycle activation shows exit from meiotic metaphase II arrest completing meiosis, second polar body extrusion removing excess chromosomes, transition to first mitotic division preparing zygote cleavage, and S-phase entry initiating DNA replication for division. Zygote polarity establishment shows asymmetric cytoplasmic organization following fertilization, molecular gradients establishing developmental axes, and polarity determining embryo proper versus suspensor fate in first division. Environmental effects on egg fertilization include temperature affecting membrane fusion kinetics and activation processes with optimal 22-28°C supporting efficient fertilization within 1-2 hours, calcium availability affecting signaling cascades critical for activation, and overall favorable conditions essential during brief fertilization window. Temperature sensitivity shows warm conditions promoting rapid fusion and activation, cool conditions delaying processes potentially reducing success, excessive heat impairing membrane dynamics or signaling, and critical temperature dependence affecting fertilization outcomes. Failed egg fertilization causes include sperm cell damage preventing membrane fusion, abnormal egg preventing fusion or activation, failed nuclear fusion preventing zygote formation, environmental stress disrupting fusion or activation processes, or genetic incompatibility though rare in self-compatible grapevine. Egg fertilization success rate shows 50-70% eggs successfully fertilizing in favorable conditions given successful tube discharge, cumulative effect of all reproductive processes determining final success, and success directly determining embryo formation potential and seed development. The process represents first component of double fertilization creating embryo lineage, occurring in parallel with central cell fertilization. Duration typically 1-4 hours from sperm release to completed fusion and activation. Cultivar variation in egg fertilization may include fusion efficiency, activation dynamics, and developmental competence affecting embryo viability. Regional timing shows egg fertilization during mid-late June across all climates within hours of tube discharge 24-48 hours post-pollination. Viticultural significance includes representing embryo-forming fusion event essential for seed development, requiring successful sperm-egg fusion and activation, demonstrating critical environmental sensitivity, establishing fertilization success as determinant of embryo formation and seed viability, and initiating embryo developmental lineage. Research applications include analyzing molecular mechanisms of sperm-egg recognition and fusion, investigating calcium signaling and activation processes, evaluating factors affecting fusion success and zygote viability, quantifying fertilization timing and efficiency, and assessing egg fertilization as determinant of seed formation outcomes.
Egg_Fertilization is first component of Double_Fertilization, distinguished by zygote-forming fusion creating embryo lineage. Represents sperm-egg fusion combining haploid gametes producing diploid 2n nucleus. Duration 1-4 hours from sperm release to activation. Involves membrane fusion, nuclear fusion, calcium signaling, egg activation. Establishes embryo developmental program. Success rate 50-70% in favorable conditions. Parallel process with Central_Cell_Fertilization. Essential for seed formation. Creates genetic diversity through recombination.
Eizell-Befruchtung
Embryo-Fertilisation
Spermium-Eizell-Fusion
Zygoten-Bildung
egg cell fusion
embryo fertilization
sperm-egg fusion
zygote formation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 393-396 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 298-300 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Egg Fertilization
An Australian viticulturist recording E-L 23 for Shiraz in the Barossa Valley on October 15 communicates that 17-20 leaves are separated and inflorescences are fully developed with flower caps still intact — the E-L system's finer resolution compared to Baggiolini is evident in its provision of 7 distinct stages (E-L 12 through E-L 19) for the period between leaf separation and flowering that Baggiolini covers with only 3 stages (E-F-G), enabling more precise scheduling of pre-bloom canopy management and disease protection interventions in the extended Australian growing season.
A phenological code assigned according to the Eichhorn-Lorenz (E-L) modified staging system for grapevine development, consisting of a numerical code (1-47) that designates one of forty-seven visually defined developmental stages from winter dormancy through berry overripeness, representing an influential Australian-adapted phenological classification that expanded and refined the original Baggiolini system with additional intermediate stages and quantitative descriptors.
Eichhorn_Lorenz_Code was originally developed by Eichhorn and Lorenz (1977) and subsequently modified by Coombe (1995) to create the system widely known as the modified E-L or Coombe scale. The system uses 47 numbered stages (E-L 1 through E-L 47) that provide finer resolution than Baggiolini's 16 alphabetic stages while preceding the BBCH system chronologically: the E-L system was particularly influential in Australian, New Zealand, and South African viticulture where it became the standard phenological reference before BBCH adoption. Cross-reference tables mapping E-L stages to both Baggiolini letters and BBCH codes are available (Coombe 1995), facilitating data interconversion across phenological recording traditions. Subclasses represent key E-L staging milestones used as management decision points.
E-L growth stage
E-L-Stadium
Eichhorn-Lorenz stage number
Eichhorn-Lorenz-Code
Eichhorn-Lorenz-Entwicklungsstadium
modified E-L code
Eichhorn KW and Lorenz DH (1977) Phänologische Entwicklungsstadien der Rebe, Nachrichtenblatt des Deutschen Pflanzenschutzdienstes 29: 119-120 Coombe BG (1995) Growth stages of the grapevine: adoption of a system for identifying grapevine growth stages, Australian Journal of Grape and Wine Research 1: 104-110 Dry PR et al. (2004) Viticulture Volume 1: Resources, 2nd Ed, Winetitles
Eichhorn Lorenz Code
A phenological record of E-L 12 for Semillon in the Hunter Valley on October 5 documents five fully separated leaves on the primary shoot with the shoot measuring 10-15 cm in length, inflorescence primordia clearly visible as 1-2 cm green protuberances at nodes 3-5, and lateral shoots beginning to emerge from basal node axils — E-L 12 serves as the reference stage for the first post-budburst shoot thinning pass in premium viticulture, where excess shoots are removed from the cordon or cane to achieve the target shoot density of 12-18 shoots per meter, and marks the developmental threshold beyond which compensatory regrowth from latent buds is unlikely to produce fruitful replacement shoots.
An Eichhorn-Lorenz code designating stage E-L 12 that describes the developmental point where five leaves have fully unfolded and separated from the shoot tip, with the shoot actively elongating and inflorescence primordia becoming visible as distinct lateral structures along the shoot axis, corresponding approximately to Baggiolini E and BBCH 15.
EL_12_Five_Leaves_Separated marks the transition from early post-budburst establishment to active shoot elongation and canopy development. The E-L system provides individual staged increments for each additional leaf separated (E-L 5 through E-L 12 corresponding to 1 through approximately 5+ leaves), offering finer resolution than the BBCH system which also counts individual leaves (BBCH 11-19) but assigns stage boundaries at each single leaf: the practical value of E-L 12 as a management milestone lies in its coincidence with the point where shoot growth rate accelerates to 3-8 cm per day, inflorescence primordia are reliably assessable, and canopy management interventions (shoot thinning, early positioning) become both necessary and effective. The stage also coincides with the conventional timing of first fungicide applications in many integrated pest management programs.
E-L Fünf-Blatt-Stadium
E-L early shoot development
E-L five-leaf stage
E-L frühes Triebentwicklungsstadium
E-L stage 12
E-L-Stadium 12
Coombe BG (1995) Growth stages of the grapevine: adoption of a system for identifying grapevine growth stages, Australian Journal of Grape and Wine Research 1: 104-110 Eichhorn KW and Lorenz DH (1977) Phänologische Entwicklungsstadien der Rebe, Nachrichtenblatt des Deutschen Pflanzenschutzdienstes 29: 119-120 Dry PR et al. (2004) Viticulture Volume 1: Resources, 2nd Ed, Winetitles pp: 141-167
EL 12 Five Leaves Separated
A phenological record of E-L 17 for Grenache in McLaren Vale on November 1 documents twelve fully separated leaves on the primary shoot with the shoot measuring 40-60 cm, inflorescences 5-8 cm in length with individual flower buds clearly distinguishable and beginning to separate, and a functional leaf area of approximately 600-800 cm² per shoot — E-L 17 represents the standard reference stage for pre-bloom canopy management interventions including shoot positioning (tucking growing shoots into the trellis foliage wires), lateral shoot removal in the cluster zone, and assessment of canopy density to determine whether leaf removal in the fruiting zone is required to improve cluster-zone light exposure and air circulation prior to flowering.
An Eichhorn-Lorenz code designating stage E-L 17 that describes the developmental point where twelve leaves have fully separated from the shoot tip, the shoot is in its rapid elongation phase, and inflorescences are well developed with individual flower buds distinguishable, corresponding approximately to Baggiolini F and BBCH 55-57.
EL_17_Twelve_Leaves_Separated positions the vine at the threshold of the critical pre-bloom period when the canopy is sufficiently developed to support reproductive growth while still requiring active management to optimize the microenvironment around developing inflorescences. The E-L system distinguishes stages 12 through 19 with progressively increasing leaf counts and inflorescence development descriptors, providing a continuous developmental narrative through the pre-bloom period that bridges the Baggiolini E-F-G transition with greater numerical precision: this resolution is particularly valued in warm-climate viticulture (Australia, California, South Africa) where the pre-bloom period is compressed and management windows are correspondingly narrow. At E-L 17 the vine's photosynthetic capacity is sufficient to support both continued shoot growth and the carbohydrate demands of the approaching flowering and fruit set period.
E-L Vorblüte-Laubstadium
E-L Zwölf-Blatt-Stadium
E-L pre-bloom canopy stage
E-L stage 17
E-L twelve-leaf stage
E-L-Stadium 17
Coombe BG (1995) Growth stages of the grapevine: adoption of a system for identifying grapevine growth stages, Australian Journal of Grape and Wine Research 1: 104-110 Eichhorn KW and Lorenz DH (1977) Phänologische Entwicklungsstadien der Rebe, Nachrichtenblatt des Deutschen Pflanzenschutzdienstes 29: 119-120 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 125-145
EL 17 Twelve Leaves Separated
A phenological record of E-L 19 for Pinot Noir in the Adelaide Hills on November 15 documents the first detachment of flowerhoods from approximately 5-10% of flowers within the most advanced inflorescences, with reflexed stamens and pollen visible on the earliest-opening flowers while the majority of buds remain closed — E-L 19 triggers the critical pre-bloom to early-bloom fungicide application for Botrytis cinerea protection, as the period from E-L 19 through E-L 25 (full flowering, 80% caps fallen) represents the window of maximum floral susceptibility to grey mould infection that can cause latent berry infection manifesting as bunch rot during the ripening phase 8-12 weeks later.
An Eichhorn-Lorenz code designating stage E-L 19 that describes the onset of flowering where the first flowerhoods (calyptrae) begin to detach and fall from the receptacle, exposing stamens and stigmas and initiating the period of pollination and fertilization, corresponding to Baggiolini H (beginning) and approximately BBCH 61.
EL_19_Flowering_Begins marks the transition from vegetative to reproductive phenology in the E-L system. The original Eichhorn-Lorenz system and Coombe's modification provide stages E-L 19 (flowering begins) through E-L 26 (cap fall complete) to describe the flowering period with greater resolution than Baggiolini's single stage H: this subdivision is operationally important because flowering-period management decisions (fungicide applications, growth regulator treatments in table grapes, pollination assessments) require precise timing within the flowering window rather than merely knowing that flowering is occurring. The E-L 19 stage corresponds to approximately 10% cap fall, providing an earlier trigger point than BBCH 61 (also nominally 10% cap fall) because E-L 19 is defined by the first observable cap detachment rather than by a percentage threshold applied to the entire inflorescence.
E-L Anthese-Anfang
E-L Blühbeginn
E-L anthesis onset
E-L early flowering
E-L stage 19
E-L-Stadium 19
Coombe BG (1995) Growth stages of the grapevine: adoption of a system for identifying grapevine growth stages, Australian Journal of Grape and Wine Research 1: 104-110 Eichhorn KW and Lorenz DH (1977) Phänologische Entwicklungsstadien der Rebe, Nachrichtenblatt des Deutschen Pflanzenschutzdienstes 29: 119-120 May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press pp: 30-80
EL 19 Flowering Begins
A phenological observer recording E-L 1 for Cabernet Sauvignon in Coonawarra on July 15 documents the vine in mid-winter dormancy with buds tightly appressed to the cane, brown, pointed, and approximately 3-4 mm in length with no detectable change from the previous month's observation — the vine has accumulated approximately 900 chilling hours (base 7.2°C) since leaf fall, endodormancy has been largely satisfied, but soil temperatures of 8-9°C and ambient maxima of 12-14°C remain below the threshold for growth resumption, with E-L 1 persisting until cumulative heat accumulation triggers the transition to E-L 2 (bud scales separating) in late August.
An Eichhorn-Lorenz code designating stage E-L 1 that describes the grapevine in winter dormancy with buds in their dormant state — brown, tightly closed by overlapping bud scales, and showing no visible sign of swelling or metabolic activation, corresponding to Baggiolini A and BBCH 00.
EL_1_Winter_Bud defines the baseline dormant state in the E-L system, equivalent to Baggiolini A and BBCH 00. The E-L system begins its numbering at 1 rather than 0, and proceeds through stages 2 (bud scales opening) and 3 (woolly bud visible) before reaching E-L 4 (budburst) — providing three intermediate stages for the pre-budburst period where Baggiolini uses only two (A and B): this additional resolution reflects the importance of early spring phenological precision in Australian viticulture where frost risk management and early-season spray timing decisions require finer temporal discrimination during the critical dormancy-to-budburst transition. The E-L 1 stage encompasses the entire winter rest period during which no externally visible change occurs despite internal physiological processes of chill accumulation and starch-sugar interconversion.
E-L Ruhestadium
E-L Winterknospe
E-L dormant bud stage
E-L stage 1
E-L winter dormancy
E-L-Stadium 1
Coombe BG (1995) Growth stages of the grapevine: adoption of a system for identifying grapevine growth stages, Australian Journal of Grape and Wine Research 1: 104-110 Eichhorn KW and Lorenz DH (1977) Phänologische Entwicklungsstadien der Rebe, Nachrichtenblatt des Deutschen Pflanzenschutzdienstes 29: 119-120
EL 1 Winter Bud
A phenological record of E-L 23 for Shiraz in the Barossa Valley on November 20 documents 50% of flowerhoods fallen across observed inflorescences, with a sweet floral fragrance perceptible in the vineyard during morning hours and active bee and hoverfly visitation — E-L 23 serves as the principal phenological reference point for predicting harvest date in Australian viticulture, with the conventional estimate of 8-10 weeks from E-L 23 to E-L 35 (véraison) and a further 5-7 weeks from véraison to harvest maturity, yielding a total of approximately 100-120 days from full flowering to harvest for Shiraz in warm inland regions and 120-140 days in cooler maritime-influenced sites such as the Adelaide Hills and Yarra Valley.
An Eichhorn-Lorenz code designating stage E-L 23 that describes full flowering where approximately 50% of flowerhoods have detached and fallen, stamens are fully reflexed on opened flowers, and active pollination is occurring across the majority of inflorescences on the vine, corresponding to Baggiolini H (mid-point) and approximately BBCH 65.
EL_23_Full_Flowering is the most consistently recorded reproductive phenological milestone in the E-L system and serves as the anchor point for phenological models predicting subsequent developmental timing. The 50% cap fall criterion provides an objective threshold that is relatively consistent across observers, though the asynchronous progression of flowering within and between clusters introduces observer variability of 1-3 days: standardized observation protocols specify assessment of 50 randomly selected inflorescences per observation plot to minimize this variability. The E-L system subdivides the flowering period into stages 19-26, providing percentage-based resolution (E-L 19 at onset, E-L 20 at 10%, E-L 21 at 30%, E-L 23 at 50%, E-L 25 at 80%, E-L 26 at 100% cap fall) that is directly comparable to the BBCH flowering subdivisions (BBCH 61-69).
E-L 50% cap fall
E-L 50%-Abblühe-Stadium
E-L Vollblüte
E-L full bloom
E-L stage 23
E-L-Stadium 23
EL 23 Full Flowering
A phenological observation of E-L 27 for Cabernet Sauvignon in Margaret River on December 10 documents young berries of 2-4 mm diameter clearly formed on the cluster rachis with abscised floral remnants visible beneath the trellis, fruit set estimated at 35-45% of total flowers — the attainment of E-L 27 triggers the first post-set yield estimation where berry count per cluster (typically 80-180 for Cabernet Sauvignon depending on set rate) is multiplied by target berry weight at harvest (1.2-1.5 g) and cluster number per vine (20-40) to generate preliminary yield projections of 6-12 t/ha that guide crop thinning decisions and winery intake planning.
An Eichhorn-Lorenz code designating stage E-L 27 that describes the completion of fruit set where young berries have formed from fertilized ovaries and are clearly visible at 2-4 mm diameter, unfertilized flowers have abscised, and the cluster has transitioned from a floral to a fruiting structure with berry number per cluster definitively established, corresponding to Baggiolini I and approximately BBCH 71.
EL_27_Fruit_Set resolves the most significant yield uncertainty by converting flower number (potential yield) into berry number (actual yield). The E-L system provides stages 26 (cap fall complete) and 27 (setting, young berries enlarging) to mark this transition, while Coombe's modification adds qualitative descriptors for set quality including coulure severity assessment: a well-set cluster at E-L 27 shows 30-60% berry retention with relatively uniform berry size, while a poorly-set cluster displays extensive gaps in the rachis, variable berry size (millerandage), and berry retention below 20%. The observation of E-L 27 is the first point at which the oenological implications of vintage conditions during flowering become visible — poor weather during E-L 19-26 manifests as reduced and uneven fruit set at E-L 27, with consequences for yield, cluster compactness, and ultimately wine style.
E-L Beerenansatz-Stadium
E-L Fruchtansatz
E-L fruit set
E-L setting stage
E-L stage 27
E-L-Stadium 27
Coombe BG (1995) Growth stages of the grapevine: adoption of a system for identifying grapevine growth stages, Australian Journal of Grape and Wine Research 1: 104-110 Eichhorn KW and Lorenz DH (1977) Phänologische Entwicklungsstadien der Rebe, Nachrichtenblatt des Deutschen Pflanzenschutzdienstes 29: 119-120 May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press pp: 80-120
EL 27 Fruit Set
A phenological record of E-L 31 for Riesling in the Eden Valley on January 5 documents berries at 7-9 mm diameter, uniformly green and hard, with the cluster hanging 12-15 cm below the cordon wire — berry chemistry at E-L 31 shows maximum titratable acidity of 30-40 g/L (as tartaric acid) with malic acid contributing 60-70% of total acidity, sugar content of 3-6° Brix, and negligible varietal aroma development, the berry being physiologically optimized for seed protection through unpalatability. E-L 31 is the standard reference stage for post-set cluster thinning in premium Australian viticulture, with target yields of 5-8 t/ha for premium Riesling requiring removal of 30-50% of clusters at this stage.
An Eichhorn-Lorenz code designating stage E-L 31 that describes the berry development stage where berries have reached approximately pea size (7-10 mm diameter), clusters are hanging under their own weight, and berries are hard, green, and at or near maximum organic acid concentration, corresponding to Baggiolini J and approximately BBCH 75.
EL_31_Pea_Size marks the approximate mid-point of Phase I (cell division phase) of berry development in the E-L system. The E-L numbering provides stages 27 through 33 for the fruit development period between set and bunch closure, with finer resolution than the BBCH 71-79 range: E-L 29 (berries pepper-corn size, 4 mm), E-L 31 (pea size, 7 mm), and E-L 33 (berries still hard and green, beginning of bunch closure) define three distinct management reference points within this growth phase. At E-L 31 the seed endosperm is solidifying and the seed coat (testa) is beginning to lignify, marking the approach of seed physiological maturity that will trigger the transition to the lag phase (Phase II) and subsequently to véraison — the developmental trajectory from E-L 31 to E-L 35 (véraison) typically requires 4-6 weeks depending on temperature accumulation.
E-L Beerenwachstum-Mitte
E-L Erbsengröße
E-L mid-berry-growth stage
E-L pea size
E-L stage 31
E-L-Stadium 31
Coombe BG (1995) Growth stages of the grapevine: adoption of a system for identifying grapevine growth stages, Australian Journal of Grape and Wine Research 1: 104-110 Eichhorn KW and Lorenz DH (1977) Phänologische Entwicklungsstadien der Rebe, Nachrichtenblatt des Deutschen Pflanzenschutzdienstes 29: 119-120 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 195-225
EL 31 Pea Size
A phenological observation of E-L 35 for Cabernet Sauvignon in Coonawarra on February 10 documents the first berries in observed clusters transitioning from hard green to soft purple-red, with approximately 10-20% of berries showing color change — E-L 35 triggers a cascade of management responses in Australian viticulture including the shift from standard to deficit irrigation (reducing water application to 30-50% of ETc to promote flavor concentration), installation of bird exclusion netting, commencement of twice-weekly berry sampling for Brix, pH, and titratable acidity monitoring, and the scheduling of post-véraison botryticide applications in Botrytis-susceptible cultivars and seasons.
An Eichhorn-Lorenz code designating stage E-L 35 that describes véraison — the onset of berry ripening characterized by berry softening (detectable by gentle compression), color change (green to yellow-translucent in white cultivars, green through red to blue-black in red cultivars), and the metabolic switch from organic acid accumulation to sugar importation, corresponding to Baggiolini L and approximately BBCH 81.
EL_35_Veraison is arguably the single most important phenological stage in the E-L system for oenological decision-making. The stage marks the developmental switch triggered by abscisic acid (ABA) signaling that simultaneously initiates sugar importation via hexose transporters, anthocyanin biosynthesis in red cultivars, cell wall disassembly and berry softening, malic acid catabolism, and the onset of flavor and aroma precursor accumulation: these coordinated changes transform the hard, green, acidic pre-véraison berry into the soft, colored, sugar-accumulating ripening berry within 24-48 hours at the individual berry level, though the asynchronous progression across berries within a cluster typically spans 5-12 days. The date of E-L 35 is the strongest single predictor of harvest date, with the E-L 35 to E-L 38 (harvest ripe) interval averaging 35-55 days depending on cultivar and regional temperature conditions.
E-L Reifebeginn
E-L Véraison
E-L ripening onset
E-L stage 35
E-L véraison
E-L-Stadium 35
Coombe BG (1995) Growth stages of the grapevine: adoption of a system for identifying grapevine growth stages, Australian Journal of Grape and Wine Research 1: 104-110 Eichhorn KW and Lorenz DH (1977) Phänologische Entwicklungsstadien der Rebe, Nachrichtenblatt des Deutschen Pflanzenschutzdienstes 29: 119-120 Coombe BG and McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening, Australian Journal of Grape and Wine Research 6: 131-135
EL 35 Veraison
A phenological record of E-L 38 for Shiraz in the Barossa Valley on March 15 documents berries at optimal harvest maturity with 25.5° Brix, pH 3.55, titratable acidity 5.8 g/L, and fully developed dark fruit and spice aroma profile assessed by sensory evaluation of crushed berry samples — the E-L system provides stages 35-38 for the ripening period (E-L 35 véraison, E-L 36 berries with intermediate Brix values, E-L 37 berries not quite ripe, E-L 38 harvest ripe) and stages 39-47 for the post-optimal period (berry shrivel, overripeness, and noble rot development), with the E-L 38 harvest decision integrating analytical chemistry, sensory assessment, weather forecasts, disease pressure evaluation, and logistical constraints into a single operational judgment.
An Eichhorn-Lorenz code designating stage E-L 38 that describes the attainment of harvest maturity where berry sugar concentration, acid balance, flavor development, and phenolic maturity have reached levels appropriate for the intended wine style, representing the oenologically defined endpoint of the ripening period, corresponding to Baggiolini M and approximately BBCH 89.
EL_38_Harvest_Ripe shares with Baggiolini M and BBCH 87-89 the inherent subjectivity of defining maturity as a phenological stage. The E-L system's provision of stages 35-38 for the active ripening period and stages 39-47 for the post-harvest and overripeness period provides more graduated resolution than either Baggiolini (L and M) or BBCH (81-89) for the oenologically critical final weeks of fruit development: E-L 36 (intermediate sugar at 18-22° Brix), E-L 37 (sugar still increasing, not yet at target), and E-L 38 (harvest ripe) enable tracking of ripening progression with management-relevant precision. The post-harvest stages E-L 39-47 document berry shrivel, raisining, and botrytized concentration processes relevant to dessert wine production — a domain where the E-L system provides uniquely detailed staging that neither Baggiolini nor BBCH addresses with comparable resolution.
E-L Erntereife-Stadium
E-L Lesereife
E-L harvest maturity
E-L ripe for harvest
E-L stage 38
E-L-Stadium 38
Coombe BG (1995) Growth stages of the grapevine: adoption of a system for identifying grapevine growth stages, Australian Journal of Grape and Wine Research 1: 104-110 Eichhorn KW and Lorenz DH (1977) Phänologische Entwicklungsstadien der Rebe, Nachrichtenblatt des Deutschen Pflanzenschutzdienstes 29: 119-120 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 260-295
EL 38 Harvest Ripe
A phenological record of E-L 4 for Chardonnay in the Yarra Valley on September 10 documents the first visible emergence of green shoot tissue (1-3 mm) through the separating brown bud scales, with the green tip just protruding beyond the surrounding protective wool — this observation triggers activation of frost protection systems (wind machines, overhead sprinklers) for the following nights when forecast minimum temperatures fall below -1°C, as the exposed green tissue at E-L 4 is killed at -2 to -3°C compared to the -15 to -20°C tolerance of the enclosed dormant bud at E-L 1, representing a 15-18°C reduction in frost hardiness within the span of the E-L 1 to E-L 4 transition.
An Eichhorn-Lorenz code designating stage E-L 4 that describes budburst — the emergence of green shoot tissue through the bud scales marking the irreversible transition from dormancy to active vegetative growth and the onset of acute spring frost vulnerability, corresponding to Baggiolini C (green point) and approximately BBCH 07.
EL_4_Budburst is the most operationally critical spring phenological stage because it defines the threshold between frost-tolerant dormancy and frost-vulnerable active growth. The E-L system provides three stages between winter dormancy and budburst (E-L 2: bud scales opening E-L 3: woolly bud with brown wool clearly visible E-L 4: green tip emergence) that refine the monitoring of this transition with greater precision than the two-stage Baggiolini progression (A to B to C). The timing of E-L 4 is cultivar-specific and temperature-driven — early-bursting cultivars (Chardonnay, Pinot Noir) typically reach E-L 4 two to three weeks before late-bursting cultivars (Cabernet Sauvignon, Mourvèdre) at the same site, a phenological spread that is exploited in site-cultivar matching to minimize frost risk in frost-prone locations.
E-L Austrieb
E-L Knospendurchbruch
E-L budburst stage
E-L green tip
E-L stage 4
E-L-Stadium 4
EL 4 Budburst
Elastic deformation quality in Red Globe table grapes provides the handling resilience enabling sea freight export from Chile to China (28-35 days transit), with fresh-harvest elastic recovery of 85-92% after 10% compression maintaining berry integrity through multiple handling transfers monitoring elastic recovery during storage (declining from 90% to 70-75% over 8 weeks at 0°C) enables quality-based inventory management that prioritizes shipment of fruit before elastic capacity declines below the threshold (approximately 70% recovery) associated with unacceptable damage rates during subsequent distribution.
Elastic_Deformation_Quality represents the physical quality characterizing the recoverable shape change of grape tissue, where deformation imposed by applied stress disappears upon unloading and tissue returns to its original dimensions. Elastic deformation reflects the reversible stretching of molecular bonds and cellular structures without permanent rearrangement, analogous to spring behavior where stored energy drives recovery. In grape berries, elastic deformation arises primarily from the reversible response of turgid cells where applied force compresses the vacuole and stretches the cell wall, with turgor pressure providing the restoring force that drives recovery upon unloading. The extent of elastic deformation that tissue can sustain (elastic limit) indicates resilience—the ability to absorb mechanical energy without permanent damage. Fresh, turgid grape tissue at optimal harvest maturity exhibits substantial elastic deformation capacity, recovering from 5-15% compression without permanent distortion. As ripening progresses and turgor declines or cell walls weaken, elastic capacity decreases and plastic deformation becomes dominant. The modulus of elasticity (Young's modulus, ratio of stress to strain in the elastic region) characterizes tissue stiffness, with higher modulus indicating stiffer, less easily deformed tissue. Elastic deformation during normal handling represents the safe operational range where berries experience stress without damage, making elastic capacity an indicator of handling tolerance. Measurement of elastic deformation requires controlled deformation to sub-damage levels followed by unloading to assess recovery, typically expressed as percentage recovery of initial dimensions after defined deformation and recovery time.
Elastic_Deformation_Quality specifically addresses the recoverable component of total deformation, reflecting fundamentally different mechanical behavior from plastic (permanent) deformation. The elastic response correlates with turgor pressure, making it sensitive to water status and cellular integrity. Elastic parameters (modulus, elastic limit, recovery percentage) provide information about tissue mechanical state distinct from firmness alone. The time-dependent viscoelastic behavior of biological tissue means that recovery assessment requires standardized recovery time for comparable results.
Elastische Verformung
elastic strain
elastische Dehnung
recoverable deformation
reversible Verformungsqualität
reversible deformation quality
rückfederbare Verformung
spring-like deformation
Bourne MC (2002) Food Texture and Viscosity: Concept and Measurement, 2nd Ed, Academic Press pp.: 107-144 Nicolaï BM et al. (2014) Texture and Quality. In: Florkowski WJ et al. (eds) Postharvest Handling: A Systems Approach, 3rd Ed, Academic Press pp.: 435-457
Elastic Deformation Quality
Elastic modulus comparison of grape berry tissues using compression testing reveals mesocarp modulus of 0.28±0.05 MPa, skin modulus of 3.2±0.6 MPa, and seed coat modulus of 185±32 MPa at harvest maturity, with the three orders of magnitude range demonstrating the mechanical heterogeneity requiring tissue-specific analysis, and the 11-fold skin-to-pulp stiffness ratio explaining why berry compression force measurements primarily reflect skin properties despite pulp comprising 85% of berry volume.
Elastic modulus quality is an elasticity quality that represents the quantitative measure of tissue stiffness defined as the ratio of stress to strain within the elastic deformation region, characterizing the resistance to deformation under applied force with higher values indicating stiffer materials requiring greater stress to produce a given strain. Within the BFO framework, this quality provides the numerical parameterization of elastic behavior, enabling quantitative comparison among tissues, developmental stages, varieties, and environmental conditions where qualitative elasticity descriptions are insufficient for rigorous analysis. The ontological significance lies in elastic modulus serving as the material property connecting applied forces to resulting deformations, where knowledge of modulus enables prediction of mechanical behavior under specified loading conditions and interpretation of observed deformations in terms of the forces that caused them. Elastic modulus in grapevine tissues spans several orders of magnitude, from approximately 0.1 MPa for highly hydrated, turgor-pressurized parenchyma to over 1000 MPa for lignified woody tissues, with the range reflecting the diversity of mechanical functions from compliant growing tissues to rigid structural support elements. The determination of elastic modulus requires measurement of both stress (force per unit area) and strain (relative deformation), with testing geometry affecting whether tension, compression, or shear modulus is obtained, and these directional moduli may differ substantially in anisotropic tissues. The distinction between different elastic modulus definitions (bulk, Young's, shear) relates to the mechanical situation being characterized, with each providing specific information about tissue mechanical behavior relevant to particular applications. Understanding elastic modulus quality enables engineering analysis of tissue mechanics, comparison of mechanical properties across biological and materials science contexts, and quantitative prediction of deformation behavior under specified loading scenarios.
Elastic_Modulus_Quality provides the quantitative specification of elastic behavior, serving as parent for specific modulus types (Bulk, Young's) that differ in their mechanical interpretation. This quality requires specification of measurement geometry and direction for anisotropic tissues. The class enables comparison with materials science data and engineering analysis of plant mechanical systems using standardized mechanical parameters.
E-Modul
Elastizitätsmodul
Steifigkeitskoeffizient
elastic constant
elastische Konstante
modulus of elasticity
rigidity modulus
stiffness coefficient
Vincent JFV (1990) Structural Biomaterials, Revised Ed. Princeton University Press, Princeton, pp. 28-57 Geitmann A (2006) Experimental approaches used to quantify physical parameters at cellular and subcellular levels. American Journal of Botany 93(10): 1380-1390
Elastic Modulus Quality
Elasticity assessment of Merlot berry skin during ripening reveals elastic recovery of 85% at véraison declining to 45% at harvest maturity, with the progressive loss of elastic behavior correlating with cell wall pectin solubilization and the transition from firm to soft texture, while the residual elasticity at harvest determines the skin's capacity to stretch during pressing without rupture, affecting juice yield and phenolic extraction during winemaking.
Elasticity quality is a physical quality that inheres in grapevine tissues and structures and represents the capacity to undergo reversible deformation in response to applied mechanical stress, returning to original dimensions upon stress removal without permanent structural change. Within the BFO framework, this quality characterizes the fundamental mechanical property distinguishing elastic from plastic deformation behavior, where the stress-strain relationship within the elastic region follows Hooke's law with deformation proportional to applied force. The ontological significance lies in elasticity determining how tissues respond to mechanical loads encountered during growth, wind exposure, handling, and processing, where elastic behavior enables reversible volume changes maintaining cellular integrity while plastic deformation indicates permanent damage or developmental modification. Grapevine tissue elasticity varies dramatically among organ types and developmental stages, with young growing tissues showing high elasticity enabling turgor-driven cell expansion, mature woody tissues exhibiting reduced elasticity due to lignification, and ripening berry tissues transitioning from elastic to increasingly plastic behavior as cell wall modification accompanies softening. The elastic properties of cell walls determine the relationship between turgor pressure and cell volume, where rigid inelastic walls maintain turgor with minimal volume change while elastic walls allow substantial volume fluctuation, with implications for water storage capacity and drought response. Measurement approaches include mechanical testing (compression, tension, puncture) recording force-deformation curves from which elastic parameters are extracted, atomic force microscopy for cellular-scale elasticity mapping, and pressure-volume analysis providing bulk tissue elastic modulus from turgor-volume relationships. The temperature dependence of tissue elasticity affects mechanical behavior under field conditions, with elasticity generally decreasing at lower temperatures as polymers stiffen. Understanding elasticity quality enables prediction of mechanical damage susceptibility, interpretation of texture measurements, and analysis of the biomechanical basis for tissue function and dysfunction.
Elasticity_Quality serves as the parent class for tissue-specific elasticity qualities and quantitative elastic modulus measurements, establishing the ontological framework for mechanical characterization of plant tissues. This quality represents the reversible component of deformation, complementing Plastic_Deformation_Quality which addresses permanent shape change. The class supports texture analysis, mechanical damage prediction, and biomechanical interpretation of tissue properties.
Elastizität
Verformbarkeit
elastic property
elastische Eigenschaft
mechanical elasticity
mechanische Elastizität
reversible deformation capacity
spring-like behavior
Bourne MC (2002) Food Texture and Viscosity: Concept and Measurement, 2nd Ed. Academic Press, San Diego, pp. 107-144 Rolle L et al. (2012) Instrumental texture analysis parameters as markers of table-grape and winegrape quality. American Journal of Enology and Viticulture 63(1): 11-20
Elasticity Quality
Electron transport rate assessment in water-stressed Shiraz comparing ETR with concurrent gas exchange reveals the stress effect on electron allocation, with well-watered vines showing ETR of 145 μmol electrons/m²/s and net assimilation of 14.2 μmol CO₂/m²/s (ETR/A ratio of 10.2, near theoretical minimum indicating efficient electron use), while stressed vines display ETR of 95 μmol/m²/s and assimilation of 6.8 μmol/m²/s (ETR/A ratio of 14.0, elevated ratio indicating substantial electron flow to photorespiration and Mehler reaction), demonstrating how stress increases the proportion of electron transport supporting alternative sinks rather than carbon fixation.
Electron transport rate quality is a chlorophyll fluorescence quality that represents the rate of linear electron flow through the photosynthetic electron transport chain estimated from chlorophyll fluorescence measurements, characterizing the flux of electrons from water oxidation through PSII and PSI to NADP⁺ reduction that drives ATP and NADPH production for carbon fixation. Within the BFO framework, this quality specifically addresses the electron flow dimension of photosynthetic function, distinguishing electron transport (light reactions) from carbon fixation (dark reactions) while recognizing that ETR provides a fluorescence-based estimate of the light reaction capacity that determines the ATP and NADPH supply for the Calvin cycle. The ontological significance lies in electron transport rate bridging light absorption and carbon fixation, where ETR indicates the capacity of the light reactions to provide reducing power and energy for CO₂ assimilation, with the relationship between ETR and actual carbon fixation depending on the allocation of electrons between carboxylation and alternative sinks including photorespiration and the Mehler reaction. The calculation of ETR from fluorescence employs the equation ETR = ΦPSII × PAR × leaf absorptance × 0.5, where ΦPSII is the operational quantum yield of PSII (Fm' - Fs)/Fm', PAR is photosynthetically active radiation, absorptance is the fraction of PAR absorbed (typically assumed 0.84), and 0.5 assumes equal distribution of absorbed photons between PSI and PSII. The relationship between ETR and carbon assimilation shows close correlation under normal conditions with approximately 4 electrons required per CO₂ fixed, but divergence occurs under stress conditions when electrons flow to alternative sinks, with the ETR/A ratio providing diagnostic information about electron allocation. The light response of ETR shows saturation behavior paralleling but typically exceeding carbon fixation saturation, as alternative electron sinks can utilize electrons beyond those required for CO₂ fixation at saturating light. Understanding electron transport rate quality enables assessment of light reaction capacity, diagnosis of electron transport limitations, and evaluation of the coupling between electron flow and carbon fixation under diverse environmental conditions.
Electron_Transport_Rate_Quality addresses the linear electron flow through the photosynthetic electron transport chain. This quality bridges light absorption and carbon fixation through fluorescence-based electron flux estimation. The class supports assessment of light reaction capacity and diagnosis of the coupling between electron transport and carbon assimilation.
ETR
ETR-Wert
Elektronentransportrate
J (electron flux)
PSII electron transport
PSII-Elektronentransport
linear electron flow
linearer Elektronenfluss
Genty B et al. (1989) The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence. Biochimica et Biophysica Acta 990(1): 87-92 Baker NR (2008) Chlorophyll fluorescence: a probe of photosynthesis in vivo. Annual Review of Plant Biology 59: 89-113
Electron Transport Rate Quality
Gewürztraminer functionally female flowers at BBCH 59-65 exhibit pronounced elevated superior ovaries positioned 0.6-0.8 mm above the receptacle surface on a distinct raised platform, with the prominent ovary (1.1-1.3 mm diameter) clearly elevated above the level of reflex stamen attachment points (stamens curving downward from attachments 0.6-0.8 mm below ovary base), creating striking visual and functional dominance of the female pistil over the reduced male organs. Chardonnay hermaphrodite flowers show moderate elevated superior ovaries with ovary base positioned 0.4-0.6 mm above receptacle surface, providing clear vertical separation between ovary and the erect stamens attached below, with elevation evident in longitudinal floral sections as an extended receptacle region supporting the pistil. Pinot Noir exhibits variable elevation among flowers, with some flowers showing minimal elevation (0.2-0.3 mm) creating subtly superior ovary positioning while other flowers on the same inflorescence show moderate elevation (0.5-0.7 mm) creating more pronounced pistil prominence, demonstrating within-cultivar developmental variation in elevation degree. Wild V. vinifera ssp. sylvestris pistillate female flowers exhibit particularly pronounced elevated superior ovaries (0.7-0.9 mm elevation) positioning the robust ovary (1.3-1.5 mm diameter) prominently above the vestigial stamen attachment points, maximizing pistil visibility and accessibility in the obligate cross-pollination system where elevated stigma positioning may enhance pollen capture from wind-borne or insect-borne pollen sources.
An elevated superior ovary is a superior ovary that is positioned notably above the receptacle surface on a distinct raised region (gynophore or carpel base extension), creating clear vertical separation between the ovary base and the attachment points of sepals, petals, and stamens, representing a common but variable expression of superior ovary positioning in Vitis where pistil elevation enhances the prominence of female organs. Elevated superior ovaries are characterized by= (1) ovary positioned 0.3-0.8 mm above the receptacle surface where other floral organs attach, with elevation evident as a distinct raised platform or stalk-like base supporting the ovary (2) the elevated region (sometimes termed gynophore when well-developed, or simply as an extended receptacle region supporting the pistil) consisting of receptacle-derived tissue that has undergone localized elongation beneath the ovary (3) ovary itself retaining superior position and free structure (no fusion with receptacle or perianth), with elevation representing enhanced positional superiority rather than a different ovary type (4) variable elevation among flowers depending on cultivar genetics, developmental conditions, and flower sexual type, with functionally female and pistillate flowers often showing greater elevation than hermaphrodite flowers, and staminate flowers showing minimal elevation. Elevated superior ovaries develop through differential growth of the receptacle region immediately beneath the developing pistil primordium during early flower development (BBCH 51-53), with localized cell division and expansion in this receptacle zone creating an elevated platform that raises the pistil above the level of other floral organ attachment points. The degree of elevation varies= minimal elevation (0.2-0.4 mm) creates subtle pistil prominence moderate elevation (0.4-0.6 mm) creates clearly visible pistil positioning above the receptacle surface pronounced elevation (0.6-0.8 mm in some functionally female or pistillate flowers) creates striking pistil prominence with the ovary positioned well above other floral structures. The functional significance of ovary elevation relates to several aspects of flower biology= elevated positioning may enhance stigma accessibility for pollen deposition by positioning the pistil prominently in the flower center elevation may facilitate post-fertilization ovary expansion by providing the developing fruit with advantageous positioning relative to other floral structures in functionally female and pistillate flowers, pronounced ovary elevation combined with reduced or reflex stamens creates clear visual and functional dominance of female over male organs elevation may provide mechanical advantage during fruit set by positioning the young berry where it experiences minimal interference from senescing floral organs. Elevated superior ovaries show cultivar-specific variation in degree of elevation, with some genotypes consistently producing more elevated ovaries than others under equivalent growing conditions, suggesting genetic control of receptacle elongation beneath the pistil. Environmental factors during early flower development can influence elevation degree, with favorable growing conditions (adequate nutrition, optimal temperatures, good water status during BBCH 51-54) potentially promoting greater elevation, while stress conditions may reduce elevation through reduced receptacle growth. The elevated condition should not be confused with inferior ovary positioning (where ovary is below other floral organs), as elevated superior ovaries remain superior (above other floral organs) with elevation simply enhancing the degree of superiority.
Elevated_Superior_Ovary emphasizes the raised positioning aspect of superior ovary organization and is distinguished from Free_Superior_Ovary (which emphasizes the non-fused aspect) though these are overlapping rather than mutually exclusive categories. Elevation represents variation in the degree of superiority, with all elevated ovaries being superior but not all superior ovaries showing pronounced elevation. Elevation degree varies with flower sexual type (greater in functionally female/pistillate types) and shows genetic and environmental variation. Understanding elevation as a variable feature of superior ovaries is relevant for interpreting flower sexual type differences, for assessing pistil development and prominence, and for comparative morphology studies within and among cultivars.
erhobenes oberständiges Ovar
erhöhter oberständiger Fruchtknoten
prominent free ovary
prominenter freier Fruchtknoten
raised superior ovary
Elevated Superior Ovary
The elevation data for vineyards in the Alto Adige/Südtirol wine region (northern Italy, 46.5°N) documents the extraordinary elevation range of viticulture in this alpine region — from 200 m on the Adige valley floor at Salurn (where Merlot and Lagrein are grown in a warm sub-Mediterranean mesoclimate of 16-17°C mean growing season temperature) through 500-700 m on the mid-slope terraces of Terlan and Tramin (where Pinot Bianco, Gewürztraminer, and Sauvignon Blanc thrive at 14-16°C) to the extreme-altitude vineyards at 800-1,000 m in Stava and above Naturns (where only early-ripening cultivars like Müller-Thurgau and Kerner achieve maturity at 12-14°C mean growing season temperature). This 800 m elevation range produces a temperature gradient of approximately 5-6°C across the region, equivalent to spanning from a warm Mediterranean climate to a cool continental climate within a horizontal distance of only 5-10 km — a mesoclimatic diversity that enables the successful cultivation of cultivars ranging from the warm-climate Lagrein to the cool-climate Riesling within a single appellation system.
A site description component that records the height above sea level of the vineyard site, expressed in meters and determined from GPS altitude measurements, digital elevation models (DEMs), or topographic map contour interpolation, with elevation being a primary determinant of temperature (the adiabatic lapse rate of approximately 0.6-1.0°C per 100 m altitude producing measurable temperature differences between vineyard elevations), growing season length, frost risk distribution, and the overall mesoclimate that positions the vineyard along the temperature gradient from warm valley floors to cool hilltops.
Elevation_Data documents the topographic parameter that exerts the strongest single influence on vineyard mesoclimate after latitude. The adiabatic lapse rate — the decrease in air temperature with increasing altitude — produces a systematic cooling effect that is remarkably consistent across viticultural regions worldwide at approximately 0.6°C per 100 m for the environmental lapse rate (the average temperature decrease in a free atmosphere) and 0.8-1.0°C per 100 m for the dry adiabatic lapse rate applicable to daytime conditions on sun-exposed slopes: this temperature gradient means that a 300 m difference in vineyard elevation produces a growing season temperature difference of 1.8-3.0°C — equivalent to the temperature difference between Bordeaux and Burgundy, or between Napa Valley and the Sonoma Coast — with proportional effects on phenological timing (budbreak delayed 5-8 days per 100 m elevation, harvest delayed 7-12 days per 100 m), fruit composition (sugar accumulation rate reduced, acid retention increased, aromatic compound profiles shifted toward cooler-climate expression), and wine style. Elevation also influences frost risk through two opposing mechanisms — higher elevations are generally warmer during radiation frost events (because cold air drains downslope, pooling in valley floors and depressions) but experience colder absolute minimum temperatures during advective frost events (where cold air masses are not stratified by density) — making the frost risk assessment site-specific and dependent on the landscape position within the elevation gradient.
Höhendaten
Seehöhe-Messung
Weinberg-Elevation
altitude record
height above sea level
vineyard elevation measurement
van Leeuwen C and Seguin G (2006) The concept of terroir in viticulture, Journal of Wine Research 17: 1-10 Jones GV et al. (2004) Spatial analysis of climate in winegrape growing regions, American Journal of Enology and Viticulture 55: 304-316 Gladstones J (1992) Viticulture and Environment, Winetitles Jackson DI and Schuster DF (2001) The Production of Grapes and Wine in Cool Climates, Gypsum Press
Elevation Data
ELISA test results for Cabernet Sauvignon clone 337 foundation mother vine record DAS-ELISA screening for six viruses conducted on phloem scrapings collected November 2023: GFLV OD405 = 0.085 (negative threshold < 0.300), GLRaV-1 OD405 = 0.062 (negative threshold < 0.250), GLRaV-3 OD405 = 0.071 (negative threshold < 0.250), GFkV OD405 = 0.092 (negative threshold < 0.300), GVA OD405 = 0.055 (negative threshold < 0.200), ArMV OD405 = 0.078 (negative threshold < 0.250) — all targets negative, positive controls confirmed reactive, maintaining virus-free certification status. ELISA test results for a Zweigelt candidate clone during pre-certification screening detected GLRaV-3 positive (OD405 = 1.85, threshold 0.250), triggering exclusion from the certification pathway pending virus elimination through heat therapy and meristem culture.
ELISA test result is a virus testing status entity that records the outcome of enzyme-linked immunosorbent assay testing performed on tissue samples from a clonal line, including the specific ELISA format employed (DAS-ELISA, TAS-ELISA, or PTA-ELISA), the target virus or viruses assayed, the optical density (OD) absorbance values obtained, the positive and negative control readings, the cut-off threshold applied, and the resulting qualitative determination (positive, negative, or inconclusive), providing the serological component of virus status assessment within grapevine certification programs.
ELISA remains the most widely used first-line diagnostic assay in grapevine certification programs due to its relative simplicity, high throughput capacity (96-well microplate format processing hundreds of samples per day), low cost per sample, and the commercial availability of validated antisera for all major grapevine viruses. The DAS-ELISA (double antibody sandwich) format is standard for most grapevine virus testing, employing virus-specific polyclonal or monoclonal antibodies that capture viral coat proteins from tissue homogenates. The cut-off threshold for positive/negative determination is typically set at 2-3 times the mean absorbance of negative control wells, though some laboratories apply fixed OD thresholds derived from historical validation data. The principal limitation of ELISA is its relatively modest sensitivity compared with molecular methods — detection limits of approximately 1-10 ng/mL coat protein mean that low-titer infections, early-stage infections, and unevenly distributed viruses may yield false-negative results. Seasonal variation in viral titer creates diagnostic windows for optimal ELISA sensitivity — GLRaV testing is most reliable from phloem tissue collected August-November, while GFLV detection in root or dormant cane tissue is less seasonally constrained. The interpretation of borderline ELISA results (OD values near the cut-off threshold) requires confirmation through repeat testing, alternative tissue sampling, or molecular assay (RT-PCR) to resolve ambiguous serological findings.
ELISA result
ELISA screening result
ELISA-Befund
ELISA-Testergebnis
ELISA-Untersuchungsergebnis
Immunoassay-Ergebnis
antibody test result
immunoassay result
serological test result
serologisches Testergebnis
Clark MF, Adams AN (1977) Characteristics of the microplate method of enzyme-linked immunosorbent assay for the detection of plant viruses. Journal of General Virology 34(3): 475-483 Martelli GP (2014) Directory of virus and virus-like diseases of the grapevine. Journal of Plant Pathology 96(1S): 1-136 Rowhani A et al. (2005) Development of a sensitive colorimetric-PCR assay for detection of viruses in woody plants. Plant Disease 89(12): 1347-1352
ELISA Test Result
Riesling berries display characteristic ellipsoid form with length-to-width ratios of 1.15-1.25 that remains remarkably consistent across the diverse growing regions from Germany's Mosel to Australia's Clare Valley, though absolute berry dimensions vary considerably with climate and cultural practices this stable shape character facilitates ampelographic identification and contributes to the moderate cluster compactness that supports Riesling's relative resistance to bunch rot compared to tighter-clustered cultivars.
Ellipsoid_Quality represents the overall form characteristic of organs exhibiting elongated geometry with rounded ends, where one principal axis (typically length) exceeds the others while maintaining smooth, continuously curved surfaces without pointed termini. Ellipsoid form in berries is defined by length-to-width ratios typically ranging from 1.10 to 1.40, producing the characteristic oval appearance common to many cultivars. Classic ellipsoid-berried varieties include Cabernet Sauvignon, Merlot, Riesling, and Sauvignon Blanc, representing some of the world's most important wine grapes. The elongated form results from differential expansion rates along the polar versus equatorial axes during berry development, with greater extension in the polar direction. Ellipsoid berries present slightly greater surface area relative to volume compared to globose berries of equal volume, potentially enhancing the skin-to-juice ratio. The reduced width in the equatorial plane can create slightly looser cluster packing compared to globose berries of similar size, potentially improving air circulation. Ellipsoid cluster form, distinct from berry form, describes bunches where length substantially exceeds width with relatively parallel sides along the cluster body. Quantitative characterization employs the length-to-width ratio, with categorical boundaries distinguishing short ellipsoid (ratio 1.10-1.20), ellipsoid (1.20-1.35), and long ellipsoid (1.35-1.50) forms in some classification systems.
Ellipsoid_Quality describes the form intermediate between globose (ratio near 1.0) and more extremely elongated shapes. The class should be distinguished from Ovoid_Quality which implies asymmetry between the two ends (broader at one end). True ellipsoid form exhibits similar curvature at both poles, producing bilateral symmetry along the long axis. OIV descriptor 223 category 2 (ellipsoid) provides the ampelographic standard. The boundary between ellipsoid and adjacent categories (globose, ovoid, narrow ellipsoid) involves judgment reporting the actual length-to-width ratio provides more precise characterization than categorical assignment alone.
Ellipsoidform
elliptic form
elliptische Gestalt
elongated rounded shape
länglich-runde Form
oval form
ovale Form
ovate-ellipsoid quality
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 83-84 Galet P (1979) A Practical Ampelography, Cornell University Press pp.: 64-67
Ellipsoid Quality
Wild V. vinifera ssp. sylvestris pistillate female flowers in Mediterranean riparian populations exhibit elongated long styles 0.88-0.98 mm connecting robust ovaries 1.2-1.35 mm to elaborate stigmas 0.55-0.62 mm, with elongated long styles representing 33-37% of total pistil length (2.65-2.85 mm), positioned prominently above vestigial stamens (0.3-0.5 mm), requiring pollen tube growth times of 82-105 hours for transit from stigma to ovules when cross-pollinated from staminate males at 20-24°C, and achieving 32-42% fruit set when males within 20-40 m provide adequate wind-borne pollen during overlapping bloom periods. Rare extreme functionally female cultivar selections show elongated long styles 0.86-0.96 mm as part of maximal pistil elaboration (total pistil 2.7-2.9 mm), with elongated styles containing extensively developed transmitting tract visible in anatomical sections as 35-50 cell layers of specialized elongated cells surrounding central canal 0.1-0.14 mm diameter filled with abundant extracellular matrix, representing maximum transmitting tract elaboration supporting effective pollen tube growth despite extreme style length. Breeding populations from wild pistillate × cultivated hermaphrodite crosses produce F1 pistillate offspring with elongated long styles 0.85-0.95 mm versus hermaphrodite offspring with medium styles 0.5-0.66 mm, with the 40-55% style length difference representing maximum sexual dimorphism in pistil morphology and allowing unambiguous flower type identification, while backcross populations show continuous style length variation from medium through extended long to elongated long ranges demonstrating quantitative genetic control of style elongation. Pollen tube growth kinetics studies in elongated long styles reveal that pollen tubes maintain consistent growth rates (14-22 μm per hour at 22-24°C) throughout the extended transmitting tract length, requiring 48-68 hours to traverse the 0.85-0.98 mm style proper plus additional time in stigma (10-18 hours) and ovary (18-28 hours) totaling 76-114 hours for complete stigma-to-ovule transit, demonstrating that elongated styles maintain functional transmitting tract supporting pollen tube growth across the extreme length.
An elongated long style is a long style at the extreme upper end of the length range, typically exceeding 0.85 mm at maturity and occasionally reaching 1 mm, representing styles showing maximal elongation characteristic of some pistillate flowers in wild dioecious populations and rare extreme functionally female cultivars, creating the longest pollen tube pathways in Vitis and representing maximum pistil elaboration. Elongated long styles are characterized by= (1) length typically 0.85-1 mm at anthesis, exceeding typical hermaphrodite styles by 30-50% and representing the upper limit of normal style length variation in Vitis (2) diameter typically 0.3-0.4 mm proportional to extreme length (3) well-developed transmitting tract tissue extending throughout the elongated length with elaborate extracellular matrix and abundant specialized cells maintaining functionality across the extended pathway (4) connection between robust ovary (typically 1.1-1.35 mm) and elaborate stigma (typically 0.52-0.65 mm) creating maximally elaborated pistils (total length typically 2.6-2.9 mm) with elongated long styles representing 32-38% of pistil length, sometimes showing slightly disproportionate style elongation contributing more than proportionally to total pistil length increase. Elongated long styles represent maximal pistil elaboration characteristic of wild dioecious V. vinifera ssp. sylvestris pistillate females and rare cultivated functionally female types showing extreme female organ investment= elongated styles develop through maximum expression of style elongation developmental programs, potentially involving enhanced cell division and expansion in style primordia (BBCH 52-56), extended duration of style elongation phase, or enhanced response to hormonal signals promoting style growth. Elongated long styles in pistillate flowers maintain proportional or slightly enhanced representation within maximally elaborated pistils (32-38% of total pistil length versus 20-35% in hermaphrodite), suggesting that extreme style elongation is part of overall maximal pistil elaboration with possible additional style-specific enhancement contributing to the extreme length. The functional implications of elongated long styles relate to maximum cross-pollination adaptation= elongated transmitting tract length (0.85-1 mm plus additional stigma and ovary tissues totaling 2-2.8 mm pollen tube pathway) requires longest pollen tube growth times in Vitis (75-110 hours at typical bloom temperatures 20-24°C), potentially providing maximum opportunity for pollen tube selection, competition, or quality assessment through the extended passage elongated styles in pistillate flowers coincide with extended stigma receptivity (often 6-8 days) creating maximum temporal window for successful cross-pollination from distant or asynchronous staminate sources the extreme style length combined with vestigial/rudimentary stamens creates maximum spatial separation between female organs and non-functional male remnants, absolutely preventing any possibility of self-pollination and enforcing complete cross-pollination dependency pistillate flowers with elongated long styles achieve 30-45% fruit set when staminate males providing pollen are within 15-50 m, with the extended style pathway not impairing fertilization success when adequate cross-pollination occurs, demonstrating that elongated styles remain fully functional despite extreme dimensions.
Elongated_Long_Style represents the extreme upper end of style length variation and is distinguished from Extended_Long_Style (0.7-0.85 mm, moderate long style) by maximal dimensions. Elongated long styles are characteristic of wild pistillate flowers and rare extreme functionally female cultivars, representing maximum pistil elaboration and maximum pollen tube pathway length in Vitis. Understanding elongated long styles is relevant for characterizing wild dioecious reproductive morphology, for understanding the upper limits of pistil elaboration, and for genetic studies of the developmental and evolutionary mechanisms producing extreme style elongation in female-functioning flower types. Elongated long styles demonstrate that Vitis transmitting tract can support effective pollen tube growth across pathway lengths up to 2.8 mm total, nearly double the hermaphrodite pathway length.
extrem verlängerter Stempelgriffel
extremely long style
maximal style
maximal verlängerter Stylus
stark verlängerter Griffel
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 95-101 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=94-109 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644.
Elongated Long Style
Merlot grafted on SO4 rootstock showing elongation zone on second-order lateral root, zone extending 4.2 mm from 0.6 mm behind meristem apex to 4.8 mm from apex, elongation zone cells 180-420 micrometers length 18-24 micrometers diameter showing high length-to-width ratio, microscopic examination revealing large central vacuoles occupying 85% cell volume and thin primary walls, cells in mid-elongation zone expanding at peak rate contributing estimated 68% of total root elongation rate 3.8 mm per day, elongation zone demonstrating active turgor-driven cell expansion converting meristematic cell production into root length increase
Elongation zone is the root tip region immediately basal to apical meristem where recently divided cells undergo rapid elongation increasing cell length 10-50 fold, typically 1-8 mm length depending on root order and growth rate positioned 0.3-1 mm behind meristem apex, characterized by cells actively expanding through vacuolation and turgor-driven wall extension, showing longitudinal cell expansion with minimal radial expansion creating elongated cylindrical cells, demonstrating rapid tissue elongation contributing 70-90% of total root length increase while meristem contributes 10-30%, and serving as primary growth zone where cell expansion converts meristematic cell production into actual root elongation enabling root penetration through soil, with elongation zone cells showing large central vacuoles occupying 80-90% cell volume, thin primary cell walls allowing expansion, high turgor pressure 0.3-0.8 MPa driving expansion, and active metabolism supporting wall synthesis, ion uptake, and osmotic adjustment, and representing critical transition region where undifferentiated meristematic cells transform into elongating cells that subsequently differentiate into functional root tissues in maturation zone. Elongation zone cellular characteristics includes= showing elongated cells (cells in elongation zone typically 50-500 micrometers length 10-30 micrometers diameter creating high length-to-width ratio), exhibiting large central vacuoles (single large vacuole occupying 80-90% cell volume providing osmotic potential for expansion), demonstrating thin primary walls (cell walls remaining thin and flexible allowing extension without secondary thickening), displaying dense cytoplasm peripherally (cytoplasm compressed into thin layer between vacuole and cell wall), and revealing active nuclei (nuclei still functional directing protein synthesis and metabolism though cells no longer dividing). Elongation zone cell expansion mechanism includes= showing turgor-driven growth (osmotic water influx creating turgor pressure stretching cell walls), exhibiting wall loosening (expansin proteins and pH changes loosening wall structure allowing extension), demonstrating wall synthesis (new wall material deposited maintaining wall thickness despite expansion), revealing directional expansion (preferential longitudinal extension through transverse microtubule orientation directing cellulose deposition), and displaying irreversible growth (cell expansion permanent and irreversible unlike elastic deformation creating stable length increase). Elongation zone position and extent includes= located behind meristem (elongation zone beginning 0.3-1 mm from root apex where cells exit meristem ceasing division), showing variable length (zone typically 1-8 mm depending on root growth rate with faster-growing roots showing longer zones), exhibiting gradual transition (smooth gradient from minimal elongation near meristem to maximum elongation mid-zone declining toward maturation zone), demonstrating order-related variation (primary roots showing longer elongation zones than fine tertiary roots reflecting vigor differences), and revealing growth rate correlation (elongation zone length positively correlated with root growth rate). Elongation zone growth contribution includes= providing majority of elongation (elongation zone cell expansion contributing 70-90% of root length increase), showing rapid expansion rate (cells in mid-elongation zone potentially increasing length 1-5 micrometers per hour), exhibiting collective effect (thousands of cells simultaneously elongating creating measurable root tip advancement), demonstrating growth velocity gradient (elongation rate varying along zone with maximum mid-zone declining toward extremes), and revealing total contribution (elongation zone producing 0.7-12 mm daily root elongation depending on conditions and root type). Elongation zone metabolism and transport includes= showing active respiration (elongation requiring substantial ATP for ion uptake, wall synthesis, and metabolism), exhibiting ion uptake (active transport accumulating K+, Cl-, and other osmolytes increasing osmotic potential), demonstrating water influx (osmotic gradient drawing water into expanding cells creating turgor), revealing protein synthesis (producing wall proteins, enzymes, and structural components supporting expansion), and displaying hormone responsiveness (auxin, gibberellins, and brassinosteroids promoting elongation while abscisic acid and ethylene potentially inhibiting). Elongation zone differentiation initiation includes= showing early differentiation signals (cells in elongation zone beginning to receive tissue-specific signals), exhibiting position-dependent fate (cell position determining whether differentiating into epidermis, cortex, endodermis, or vascular tissues), demonstrating pattern formation (coordinate differentiation creating organized tissue patterns), revealing transcriptional changes (differentiation-specific genes beginning expression in elongation zone), and displaying structural preparation (cells preparing for differentiation through organelle development and biochemical changes). Elongation zone environmental responsiveness includes= demonstrating mechanical sensitivity (elongation zone sensing soil resistance potentially modulating expansion rate), showing moisture dependence (adequate turgor requiring soil water availability with drought reducing elongation), exhibiting temperature effects (elongation rate increasing with temperature from threshold to optimum), revealing nutrient influence (adequate nutrients supporting metabolism and wall synthesis necessary for elongation), and displaying hormone integration (environmental signals transduced through hormonal changes affecting elongation zone activity).
Elongation_Zone emphasizes cell expansion region behind meristem and is distinguished from meristem (cell division) and maturation zone (differentiation) by focus on cell elongation. Includes Rapid_Elongation_Zone and Cell_Expansion_Elongation_Zone. Critical for understanding root elongation mechanism, turgor-driven growth, cell expansion, length increase contribution, and the expansion region providing majority of root elongation converting meristematic cell division into actual root length.
Cell elongation region
Cell expansion zone
Elongationszone
Streckungswachstumsbereich
Stretching zone
Zellstreckungszone
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Cosgrove, D. J. (2005). Growth of the plant cell wall. Nature Reviews Molecular Cell Biology, 6(11), 850-861.
Elongation Zone
Embolism recovery capacity assessment in Shiraz following a severe drought event (stem water potential reaching -2.4 MPa, 70% conductivity loss) reveals partial but incomplete recovery, with conductivity recovering to 55% of pre-stress levels within 48 hours of irrigation (rapid refilling component), further recovering to 75% over two weeks (slower refilling or early new xylem), and achieving 90% recovery only after six weeks (new vessel differentiation contributing), demonstrating the multi-phase recovery process and the timeline implications for sustained hydraulic function following severe stress.
Embolism recovery capacity quality is a vulnerability to cavitation quality that represents the ability of grapevine xylem to restore hydraulic conductivity following embolism events, characterizing the potential for repair of air-filled conduits that determines whether drought-induced hydraulic damage is permanent or reversible and affects the vine's resilience to repeated stress episodes. Within the BFO framework, this quality addresses the recovery dimension of hydraulic vulnerability, distinguishing the capacity to regain function from the resistance to initial dysfunction while recognizing that recovery capacity may be as important as initial vulnerability in determining hydraulic performance across seasons with multiple drought events and recovery opportunities. The ontological significance lies in embolism recovery determining whether vines can restore hydraulic function after stress without requiring new xylem construction, where effective recovery enables rapid restoration of water transport capacity following rain or irrigation, while limited recovery capacity means that drought damage accumulates through the season, progressively reducing the conductivity available to support transpiration. The mechanistic basis for embolism recovery remains an active area of research, with proposed mechanisms including positive pressure refilling (root pressure or stem pressure dissolving air and refilling vessels), osmotic refilling (solute secretion into embolized vessels creating osmotic water influx), and new xylem production (replacing rather than repairing damaged conduits), with evidence for different mechanisms operating in different species, organs, and conditions. The temporal dynamics of recovery show potential for rapid restoration (hours to days) through refilling mechanisms if conditions permit, versus slower restoration (weeks to months) through new xylem differentiation, with the relative importance depending on the severity of embolism, the season, and the vine's physiological status. The practical significance of recovery capacity for viticulture includes the potential for overnight or post-irrigation hydraulic recovery that affects next-day gas exchange capacity, the cumulative effects of incomplete recovery creating progressive seasonal decline in hydraulic function, and the implications for drought recovery following stress events. Understanding embolism recovery capacity quality enables assessment of vine resilience to drought episodes, prediction of recovery timelines following stress, and evaluation of the long-term hydraulic health implications of repeated drought exposure.
Embolism_Recovery_Capacity_Quality serves as the parent class for specific recovery parameters (Refilling Capacity, Safety Margin). This quality determines vine resilience to drought through the ability to restore hydraulic function. The class supports drought recovery assessment and long-term hydraulic health evaluation through recovery capacity characterization.
Embolieerholungskapazität
Leitfähigkeitswiederherstellung
Xylemerholung
conductivity restoration
embolism repair
hydraulic recovery
hydraulische Erholung
xylem recovery
Brodersen CR, McElrone AJ (2013) Maintenance of xylem network transport capacity: a review of embolism repair in vascular plants. Frontiers in Plant Science 4: 108 Hochberg U et al. (2017) Grapevine petioles are more sensitive to drought induced embolism than stems: evidence from in vivo MRI and microCT observations of hydraulic vulnerability segmentation. Plant, Cell & Environment 40(9): 1831-1842
Embolism Recovery Capacity Quality
Investigation of Embolism_Repair_Disposition at UC Davis employed X-ray computed microtomography (microCT) to visualize embolism formation and potential refilling in living grapevine stems across 8 cultivars. Potted vines were droughted to induce 50-70% PLC, then rewatered, with microCT imaging at daily intervals tracking the status of individual vessels (water-filled versus air-filled). Embolism_Repair_Disposition showed substantial variation among cultivars. 'Chardonnay' exhibited rapid repair with 75% of embolized vessels refilling within 7 days of rewatering, coinciding with sustained positive xylem pressure (up to +0.08 MPa) measured at night, suggesting root pressure-driven refilling. 'Cabernet Sauvignon' showed slower, less complete repair (45% refilling at 7 days) with weaker root pressure development (+0.02 MPa). Interestingly, repair was vessel-size dependent, with narrow vessels ( 200 μm), consistent with the physics of dissolving air bubbles under positive pressure. Season also affected Embolism_Repair_Disposition, with spring drought events showing more complete recovery (85% vessel refilling) than late-season drought (55% refilling), potentially reflecting root starch reserves available for osmotic work and/or active vessel development in spring. Cumulative effects were documented over 3 seasons of repeated drought cycles, with vines experiencing chronic stress showing progressive decline in maximum conductivity (15% reduction from year 1 to year 3), indicating that Embolism_Repair_Disposition has limits and severe repeated stress causes lasting hydraulic damage.
An Embolism_Repair_Disposition is a Hydraulic_Architecture_Disposition that inheres in a Vitis vinifera plant and is realized through physiological mechanisms that restore hydraulic conductivity in embolized xylem vessels, either by refilling cavitated vessels with water or by growing new functional xylem to bypass blocked conduits. This disposition represents the recovery component of hydraulic function, determining how effectively vines can restore water transport capacity following drought-induced cavitation and whether hydraulic damage has persistent effects on vine function. Embolism repair mechanisms remain debated but may include root pressure-driven refilling (positive xylem pressure generated in roots dissolving gas and forcing water into embolized vessels, typically occurring at night or following rewetting), osmotic refilling (localized solute secretion lowering water potential and drawing water into embolized vessels from adjacent living cells), and new xylem production (vessel formation bypassing irreparably blocked conduits). The relative importance of these mechanisms likely varies with species, timing, and drought severity, with root pressure potentially effective for moderate embolism while severe damage may require new vessel development. Embolism_Repair_Disposition in grapevine appears substantial, as vines commonly recover from significant conductivity loss following rewetting, but repair capacity has limits and repeated severe cavitation may cause cumulative hydraulic damage reducing vine productivity over time.
Embolism_Repair_Disposition is_a Hydraulic_Architecture_Disposition. Embolism repair restores conductivity after cavitation through vessel refilling and/or new xylem growth. Root pressure is implicated in refilling, operating at night or after rewetting. Narrow vessels refill more readily than wide vessels. Repair capacity varies among genotypes and with season. Repeated severe cavitation can exceed repair capacity, causing cumulative hydraulic damage. The mechanisms and extent of embolism repair remain active research areas.
Embolie-Reparatur-Disposition
Hydraulische-Erholung
Xylem-Wiederherstellung
embolism reversal
hydraulic restoration
Brodersen CR et al. (2010) The dynamics of embolism repair in xylem: in vivo visualizations using high-resolution computed tomography, Plant Physiology 154:1088-1095 Sperry JS et al. (1987) Spring filling of xylem vessels in wild grapevine, Plant Physiology 83:414-417 Zwieniecki MA, Holbrook NM (2009) Confronting Maxwell's demon: biophysics of xylem embolism repair, Trends in Plant Science 14:530-534
Embolism Repair Disposition
Cabernet Sauvignon mature seed embryos at BBCH 89 measure 4-5 mm length with well-differentiated cotyledons (2.5-3 mm length each), distinct radicle (1-1.5 mm), and require 100-110 days cold stratification at 3-5°C for 80-90% germination success
An embryo is the rudimentary plant structure within a Vitis vinifera seed that develops from the fertilized egg cell and contains the genetic material and organized tissues necessary for germination and seedling establishment. The grape seed embryo develops through characteristic stages= proembryo (immediate post-fertilization, 0-7 days), globular embryo (7-14 days post-fertilization during early cell division), heart-stage embryo (14-21 days with initial cotyledon differentiation), torpedo-stage embryo (21-35 days with elongated cotyledons and radicle formation), and mature embryo (35+ days post-fertilization with fully differentiated structures). The mature grape embryo consists of= two cotyledons that serve as storage organs and initial photosynthetic structures upon germination, a radicle that develops into the primary root system, a hypocotyl connecting cotyledons to radicle, and an apical meristem (plumule) that will form the shoot system. Embryo size in Vitis vinifera mature seeds typically ranges 3-5 mm length, occupying approximately 40-60% of total seed volume with the remainder filled by endosperm tissue. Embryo development is tightly coordinated with endosperm development= the endosperm provides nutritional support during embryogenesis and continues to serve as nutrient reserve during germination, while the embryo produces hormones that regulate both seed and berry development. Embryo viability is critical for seed functionality= viable embryos show organized cellular structure, intact cell membranes, and metabolic competence for germination, while non-viable embryos exhibit cellular degeneration, membrane breakdown, or developmental abnormalities. Embryo dormancy in Vitis vinifera seeds is physiological rather than morphological= embryos are fully formed at seed maturity but require extended cold stratification (90-120 days at 1-5°C) to overcome dormancy and initiate germination. Environmental stresses during embryogenesis can cause embryo abortion= temperature extremes (below 10°C or above 35°C), severe water deficit, or nutritional limitations during the 2-5 week post-fertilization period can arrest embryo development, leading to seed abortion in seedless cultivars or non-viable seeds in seeded cultivars.
Embryo emphasizes the complete rudimentary plant structure and is distinguished from Developed_Embryo (fully differentiated mature structure) and Dormant_Embryo (physiological state) by encompassing all embryonic developmental stages from proembryo through mature dormant embryo. Critical for understanding seed viability, germination requirements, breeding program success rates, and mechanisms of seed abortion in seedless cultivars where embryo development fails.
Embryonale Pflanzenanlage
Keim
Plant embryo
Samenkeimling
Seed embryo
Ebadi, A., Sedgley, M., May, P., & Coombe, B. G. (1996). Seed development and abortion in Vitis vinifera L., cv. Chardonnay. International Journal of Plant Sciences, 157(6), 703-712. | Bouquet, A., & Davis, H. P. (1989). Culture in vitro d'embryons immatures de vigne (Vitis vinifera L.) appliquée à la sélection de variétés de cuve sans pépins. Agronomie, 9(6), 565-574.
Embryo
Merlot in Bordeaux shows embryo desiccation tolerance development during September-early October Year N+1, with LEA proteins accumulating to 8-16% total embryo protein including LEA 1, 2, 3 families, protective sugars accumulating with sucrose 7-12% and raffinose 2.8-4.6% dry weight, possible HSP70 and HSP90 heat shock proteins accumulating, embryo water content declining from 54% to 28% during tolerance acquisition weeks 12-15, full tolerance enabling survival at 12-16% final moisture, tolerance tested by artificial desiccation showing >85% germination after rehydration, under oceanic conditions of 20-26°C temperature, typical of this mid-season cultivar developing desiccation tolerance enabling seed viability and storage
Embryo desiccation tolerance represents the dehydration survival capacity acquisition phase during seed maturation when embryo develops ability to survive water loss from 50-70% moisture to 10-20% in mature seed, late embryogenesis abundant LEA proteins accumulate providing cellular protection, protective sugars including sucrose and raffinose oligosaccharides accumulate stabilizing membranes and proteins, heat shock proteins possibly accumulate as chaperones, integrated protective mechanisms enable embryo survival at low moisture, and desiccation tolerance essential for seed viability and storage, developing during late maturation weeks 12-16 as critical seed survival mechanism. This dehydration tolerance acquisition phase occurs during early autumn in September-early October in temperate regions Year N+1, representing embryo protective capacity development. Tolerance acquisition mechanism involves late embryogenesis abundant protein synthesis and accumulation, protective sugar biosynthesis and accumulation, possible heat shock protein accumulation, membrane compositional changes possibly enhancing stability, cellular structural adaptations enabling low-moisture survival, and integrated protective systems enabling desiccation tolerance. Desiccation tolerance definition shows ability to survive water content reduction to very low levels 10-20% moisture, tolerance enabling seed survival during maturation desiccation and storage, tolerance reversible through imbibition during germination, and tolerance as seed-specific adaptation. Moisture reduction shows embryo water content declining from 50-70% during mid-development to 30-40% during early desiccation to final 10-20% in mature dry seed, substantial water loss requiring protective adaptations, desiccation occurring progressively during maturation, and low moisture as mature characteristic. Desiccation tolerance timing shows tolerance developing during late maturation weeks 12-16, tolerance acquisition preceding or accompanying desiccation, fully developed tolerance by maturation completion, and timing as developmental requirement. Late embryogenesis abundant protein accumulation shows LEA proteins as major desiccation protectants, multiple LEA protein families expressed including LEA 1, 2, 3, 4, 5, LEA proteins accumulating substantially during late embryogenesis, typical LEA protein content 5-20% total embryo protein in mature seed, and LEA proteins as characteristic seed proteins. LEA protein families shows diverse LEA protein types with different structures and functions, all LEA proteins hydrophilic with high proportion of charged and polar amino acids, LEA proteins lack ordered structure in solution but may fold during desiccation, and diversity providing comprehensive protection. LEA protein functions shows proteins protecting membranes during dehydration, proteins preventing protein aggregation, proteins acting as molecular chaperones, proteins stabilizing cellular structures, proteins binding water molecules, and multiple protective functions supporting desiccation survival. LEA protein gene expression shows LEA genes strongly upregulated during late maturation, abscisic acid inducing LEA gene expression, transcription factors regulating LEA expression, and expression as LEA accumulation mechanism. LEA protein localization shows LEA proteins distributed in cytoplasm, nucleus, possibly associated with membranes, and cellular distribution enabling protection. Protective sugar accumulation shows sucrose accumulating to high levels 5-15% embryo dry weight, raffinose family oligosaccharides including raffinose, stachyose possibly accumulating 2-6% dry weight, total soluble sugars 10-25% providing substantial protection, and sugars as essential desiccation protectants. Sucrose characteristics shows sucrose as non-reducing disaccharide, sucrose highly soluble and stable, sucrose as major transported and storage sugar, and sucrose as primary protective sugar. Raffinose family oligosaccharides shows raffinose as trisaccharide, stachyose as tetrasaccharide, RFOs synthesized from sucrose, RFOs particularly effective desiccation protectants, and RFOs as specialized protective sugars. Sugar protective mechanisms shows sugars replacing water in hydrogen bonding with membranes, sugars maintaining membrane spacing and fluidity, sugars preventing membrane fusion during dehydration, sugars stabilizing proteins preventing denaturation, sugars forming glasses at low moisture vitrifying cytoplasm, glass transition preventing crystallization and molecular mobility, and multiple mechanisms providing comprehensive protection. Water replacement hypothesis shows sugars substituting for water in hydrogen bonds with phospholipid headgroups, substitution maintaining membrane integrity, and replacement as membrane protection mechanism. Glass formation shows concentrated sugars forming highly viscous glassy state at low moisture, glass limiting molecular motion preventing damage, glass stabilizing cellular structures, and glass formation as critical protection mechanism. Heat shock protein possible accumulation shows HSPs possibly accumulating during late maturation, HSP70 and HSP90 families as molecular chaperones, HSPs protecting proteins from denaturation, HSPs assisting protein refolding, and HSPs as additional protective proteins. Membrane compositional changes shows possible phospholipid composition changes increasing membrane stability, possible increases in saturated fatty acids reducing fluidity, membrane adaptations enhancing desiccation tolerance, and membrane changes as structural protection. Cellular structural adaptations shows cells becoming highly concentrated during desiccation, organelles remaining intact through protective mechanisms, cytoplasm vitrifying in glassy state, cellular architecture preserved enabling rehydration, and structural maintenance essential for survival. Desiccation damage prevention shows protective mechanisms preventing protein denaturation, membrane fusion, DNA damage, oxidative damage, and structural disruption, prevention enabling embryo survival and germination capacity retention. Oxidative stress management shows antioxidant systems possibly upregulated, reactive oxygen species scavenging preventing damage, oxidative protection as component of desiccation tolerance, and antioxidants as protective molecules. Desiccation tolerance testing shows tolerance assayed by artificial desiccation and rehydration testing viability, tolerant embryos surviving desiccation to 10-20% moisture and recovering upon imbibition, tolerance quantified by germination or growth after desiccation, and testing enabling tolerance characterization. Tolerance development dynamics shows progressive tolerance acquisition during maturation, tolerance increasing as protective molecules accumulate, full tolerance achieved by maturation completion, and dynamics as developmental progression. Tolerance as seed characteristic shows desiccation tolerance distinguishing orthodox seeds like grapevine from recalcitrant seeds lacking tolerance, tolerance enabling dry seed storage, tolerance as adaptation for seed dispersal and survival, and tolerance as fundamental seed trait. Tolerance reversibility shows desiccation tolerance maintained in dry seed, imbibition reversing desiccation, rehydration enabling resumption of metabolism and germination, and reversibility enabling seed function. Storage stability shows desiccation-tolerant seeds storable long-term under appropriate conditions, dry cool storage maintaining viability, tolerance enabling seed banks and propagation, and stability as tolerance benefit. Environmental effects on tolerance acquisition include temperature affecting development rate with optimal 20-26°C supporting tolerance development, environmental stresses possibly enhancing tolerance, and conditions affecting protective molecule accumulation. Temperature effects shows warm conditions supporting normal tolerance development, temperature affecting accumulation kinetics, and temperature as rate factor. The process represents first aspect of seed maturation establishing desiccation survival capacity, preceding seed dormancy acquisition and ripening completion. Duration tolerance acquisition during weeks 12-16 representing protective capacity development. Regional timing shows embryo desiccation tolerance development during September-early October as protection establishes. Viticultural significance includes representing critical seed survival mechanism enabling maturation and storage, determining seed storability and longevity, demonstrating seed adaptation to dispersal and dormancy, and establishing seed viability maintenance capacity. Research applications include analyzing desiccation tolerance mechanisms and regulation, investigating protective molecule functions and interactions, evaluating tolerance development dynamics, quantifying protective compound accumulation, and assessing desiccation tolerance as seed viability and quality determinant.
Embryo_Desiccation_Tolerance is first aspect of Seed_Maturation, distinguished by survival capacity acquisition. Represents weeks 12-16. LEA proteins 5-20% total protein (multiple families) protecting membranes and proteins. Protective sugars: sucrose 5-15%, raffinose family 2-6% DW. Water replacement and glass formation mechanisms. Possible HSP accumulation. Embryo moisture 50-70% → 10-20%. Enables survival at low moisture, seed storage. Precedes Seed_Dormancy_Acquisition. Critical seed survival adaptation. Temperature 20-26°C optimal.
Austrocknungs-Überlebenskapazität
Dehydratations-Toleranz
Embryo-Austrocknungstoleranz
Wasserverlust-Toleranz
dehydration tolerance
desiccation survival capacity
drought tolerance acquisition
water loss tolerance
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 441-442
Embryo Desiccation Tolerance
Merlot in Bordeaux shows embryo development during July-October Year N+1, with zygote dividing 36-48 hours post-fertilization creating two-cell proembryo, globular embryo 60-85 micrometers weeks 1-3, heart embryo 180-260 micrometers weeks 3-7 with cotyledon primordia, torpedo embryo 600-1000 micrometers weeks 7-12 with elongated axis, mature embryo 3.6-4.8 mm length weeks 12-16 with fully developed radicle, hypocotyl, cotyledons, and shoot apex, storage reserves accumulating 31-37% lipids and 27-33% proteins in cotyledons, under oceanic conditions of 20-26°C temperature, typical of this mid-season cultivar completing embryogenesis coordinating with seed and berry maturation
Embryo development represents the embryonic structure formation process during seed development when diploid zygote 2n=38 undergoes cell division, morphogenesis, and maturation creating organized seedling comprising radicle, hypocotyl, cotyledons, and shoot apex with accumulated storage reserves, proceeding through sequential developmental stages from first zygotic division through globular, heart, and torpedo embryo stages to mature embryo ready for germination, spanning approximately 12-18 weeks from fertilization. This embryo formation process occurs during summer to early autumn in July-October in temperate regions Year N+1, representing complete embryogenesis from single-cell zygote to multicellular organized embryo. Embryo development mechanism involves zygote division with first asymmetric division producing two-cell proembryo establishing apical-basal axis, embryo morphogenesis with progressive cell divisions and differentiation creating globular undifferentiated embryo weeks 1-3, heart stage embryo weeks 3-7 with cotyledon primordia emergence establishing bilateral symmetry, torpedo stage embryo weeks 7-12 with elongated embryonic axis and developing cotyledons, and embryo maturation weeks 12-18 with structure completion and storage reserve accumulation creating viable embryo. Zygote characteristics shows single diploid cell 2n=38 from sperm-egg fusion, zygote polarized with apical-basal axis, cell preparing for division through DNA replication and mitotic apparatus formation, and zygote as embryo developmental starting point. First zygotic division shows asymmetric division 24-72 hours post-fertilization depending on temperature, division producing smaller apical cell and larger basal cell, apical cell giving rise to embryo proper, basal cell producing suspensor support structure, and asymmetric division establishing body plan. Two-cell proembryo shows apical cell 20-40 micrometers diameter destined for embryo, basal cell 40-60 micrometers forming suspensor, suspensor anchoring embryo in endosperm and possibly transferring nutrients, and two-cell stage as developmental transition from zygote to embryo. Embryo morphogenesis progression shows globular stage embryo weeks 1-3 with spherical cell mass 50-100 micrometers diameter, undifferentiated cells undergoing rapid division, no visible organ primordia, heart stage weeks 3-7 with 150-300 micrometers size showing cotyledon bulges creating heart shape, bilateral symmetry established, torpedo stage weeks 7-12 with 500-1200 micrometers length showing elongated embryonic axis with distinct radicle, hypocotyl, and enlarging cotyledons, and progressive morphogenesis creating seedling structures. Globular embryo characteristics shows spherical to ovoid cell mass, cells small and densely cytoplasmic, protoderm differentiating at surface, ground meristem and procambium beginning specification, and globular as undifferentiated embryo stage. Heart embryo characteristics shows cotyledon primordia emerging as two lateral bulges, bilateral symmetry from cotyledon positioning, embryonic axis visible between cotyledons, shoot apex initiating at axis apex, radicle region differentiating at basal end, and heart stage as organ initiation stage. Torpedo embryo characteristics shows elongated embryonic axis 3-8 fold length to width ratio, cotyledons enlarging and beginning to curve, radicle clearly differentiated with root cap initiation, hypocotyl region between radicle and cotyledons, shoot apical meristem with leaf primordia possibly initiating, and torpedo as major growth and differentiation stage. Embryonic axis formation shows radial organization with protoderm epidermis, ground meristem cortex and pith, procambium vascular tissue precursor, and axial organization establishing seedling body plan. Cotyledon development shows cotyledons enlarging during torpedo and maturation stages, cotyledons as embryonic leaves storing reserves, cotyledons eventually large accounting for 70-85% embryo volume, and cotyledons as major embryo component. Radicle differentiation shows radicle as embryonic root, root cap forming at radicle tip, root apical meristem establishing behind cap, procambium extending through radicle, and radicle as first structure emerging during germination. Hypocotyl formation shows hypocotyl as embryonic stem between radicle and cotyledons, hypocotyl elongating during germination, vascular connection through hypocotyl, and hypocotyl as axis component. Shoot apex development shows shoot apical meristem initiating between cotyledon bases, meristem as source of post-germination shoot growth, possible first leaf primordia initiating in some species though unclear in grapevine, and shoot apex completing embryo structures. Embryo maturation shows storage reserve accumulation primarily in cotyledons weeks 12-18, lipid bodies forming accumulating triacylglycerols reaching 30-40% cotyledon dry weight, protein bodies forming accumulating storage proteins 25-35% dry weight, embryo growth slowing during maturation, metabolic activity declining, and maturation preparing embryo for germination. Lipid accumulation in embryo shows oleic acid 18 per 1 and linoleic acid 18 per 2 as major fatty acids, triacylglycerols in lipid bodies, lipid synthesis from photosynthate imported from parent plant, and lipids as major energy reserves. Protein accumulation shows albumins and globulins as storage proteins, proteins in protein bodies, amino acid composition balanced for seedling nutrition, and proteins as nitrogen reserves. Desiccation tolerance development shows embryo acquiring tolerance to dehydration during maturation, late embryogenesis abundant LEA proteins accumulating, sugars particularly sucrose accumulating providing cellular protection, and desiccation tolerance enabling seed survival. Embryo size shows mature embryo typically 3-5 mm length and 1.5-3 mm width, size variation from seed development conditions, embryo occupying 40-60% seed volume with remainder endosperm and coat, and mature size adequate for germination success. Embryo viability shows mature embryo capable of germination if provided appropriate conditions, viability requiring complete development including reserve accumulation and desiccation tolerance, and viable embryo as seed development outcome. Environmental effects on embryo development include temperature affecting development rate with optimal 22-28°C for normal progression, nutrient availability affecting reserve accumulation, water stress potentially limiting development, and favorable conditions ensuring complete embryogenesis. Temperature effects shows warm conditions accelerating development, cool conditions slowing progression, and temperature determining development duration within 12-18 week range. Embryo abortion shows some embryos failing to complete development from genetic abnormalities, inadequate resources, or developmental disruptions, abortion creating seeds without viable embryos or empty ovules, and abortion as developmental failure. The process encompasses three sequential developmental phases including zygote division initiating embryogenesis, embryo morphogenesis creating structures through globular, heart, and torpedo stages, and embryo maturation completing development with reserve accumulation. Duration typically 12-18 weeks representing complete embryogenesis. Regional timing shows embryo development during July-October coordinating with seed and berry development. Viticultural significance includes representing formation of next generation plant, storage reserve accumulation determining germination vigor, embryo viability determining seed quality, and embryo development as fundamental reproductive process. Research applications include analyzing embryogenesis regulation and morphogenesis, investigating storage reserve biosynthesis and accumulation, evaluating environmental effects on embryo development, quantifying developmental stages and timing, and assessing embryo development as seed viability determinant.
Embryo_Development represents embryonic structure formation within Seed_Development. Subdivided into Zygote_Division with first asymmetric division establishing axis, Embryo_Morphogenesis with globular (weeks 1-3), heart (weeks 3-7), torpedo (weeks 7-12) stages, and Embryo_Maturation (weeks 12-18) with storage reserve accumulation. Duration 12-18 weeks. Mature embryo 3-5 mm with radicle, hypocotyl, cotyledons, shoot apex. Storage reserves 30-40% lipids, 25-35% proteins. Creates viable seedling structures. Critical reproductive development.
Embryo-Bildung
Embryo-Entwicklung
Embryogenese
embryo formation
embryogenesis
embryonale Entwicklung
embryonic development
seedling structure development
Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 438-442
Embryo Development
Syrah in Rhône Valley shows embryo genesis start during early July Year N+1 at BBCH 69-71, with first asymmetric zygotic division occurring 28-48 hours post-fertilization producing smaller apical cell developing into embryo proper and larger basal cell forming suspensor, under warm Mediterranean conditions of 25-30°C temperature, typical of this mid-to-late season cultivar completing first division 60-84 hours post-pollination initiating embryogenesis and seed development supporting berry growth
Embryo genesis start represents the definitive embryogenesis initiation marked by first zygotic division when unicellular diploid zygote undergoes asymmetric mitotic cleavage producing two-cell embryo with distinct apical and basal daughter cells establishing embryo proper and suspensor lineages, marking transition from unicellular zygote to multicellular embryo beginning developmental progression toward mature seed. This critical first division occurs during zygote formation in late June-early July in temperate regions Year N+1, typically 24-72 hours following fertilization depending on temperature, corresponding to BBCH 69 fertilization completion transitioning to BBCH 71 early berry development stages. Genesis initiation involves first mitotic division with chromosome condensation, spindle formation, and segregation, asymmetric cytokinesis creating unequal daughter cells with smaller apical cell and larger basal cell, cell fate determination with apical cell developing into embryo proper and basal cell forming suspensor, and establishment of two-cell embryo marking definitive embryogenesis beginning. First mitotic division characteristics show chromosome condensation making chromatin visible as distinct structures, 38 chromosomes in Vitis vinifera aligning at metaphase plate, spindle microtubule attachment to kinetochores, anaphase segregation distributing sister chromatids to opposite poles, and telophase nuclear envelope reformation around separated chromosome sets. Division asymmetry shows unequal cytokinesis creating daughter cells of different sizes, smaller apical cell typically 20-40 micrometers diameter positioned toward micropyle, larger basal cell 40-60 micrometers diameter positioned toward chalaza, and size asymmetry reflecting differential developmental fates. Cell division plane shows transverse orientation perpendicular to apical-basal axis established during zygote development, plane positioning determining apical versus basal daughter identity, and plane regulated by polarity cues and cytoskeletal organization. Cytokinesis mechanism shows phragmoplast microtubule array forming at division plane, cell plate synthesis initiated at center extending outward, vesicle fusion depositing new cell wall material creating partition, and cell plate maturation completing separation into two distinct cells. Daughter cell characteristics show apical cell as smaller dense cytoplasm-rich cell with high nucleus-to-cytoplasm ratio, basal cell as larger vacuolated cell with lower nucleus-to-cytoplasm ratio, cytoplasmic differences reflecting distinct developmental programs, and asymmetric distribution of developmental determinants establishing cell fates. Apical cell fate shows development into embryo proper through subsequent symmetric divisions, cell proliferation producing globular embryo, progressive differentiation establishing cotyledons, shoot apical meristem, hypocotyl, and radicle, and ultimate development into mature embryo within seed. Basal cell fate shows development into suspensor through elongation and limited division, suspensor connecting embryo proper to surrounding tissues, function providing nutritive and possibly hormonal support for embryo development, and suspensor degeneration during later embryogenesis as embryo proper becomes self-sufficient. Cell fate determination shows asymmetric distribution of developmental regulators during division, apical cell inheriting factors promoting meristematic identity and embryo development, basal cell inheriting factors promoting suspensor differentiation, and molecular differences establishing distinct lineages from first division. Molecular asymmetry includes differential expression of transcription factors regulating embryo versus suspensor identity, signaling molecules affecting cell fate decisions, and epigenetic modifications distinguishing cell lineages. First division timing shows temperature-dependent variation with warm conditions 25-28°C promoting division at 24-36 hours post-fertilization, cool conditions 18-22°C delaying division to 48-72 hours or more, and timing as critical determinant of overall embryogenesis rate. Division synchrony among embryos shows variable timing with some embryos dividing earlier and others later even within same seed, asynchrony creating developmental heterogeneity, and timing differences potentially affecting seed development uniformity. Two-cell embryo characteristics show definitive multicellular state with distinct apical and basal cells, cell-cell communication through plasmodesmata enabling signaling, coordinated development of both cells supporting embryogenesis, and two-cell stage representing committed embryo having initiated developmental program. Embryo proper subsequent development shows progressive divisions producing four-cell then octant-stage embryos, globular stage with radial symmetry, transition stage initiating bilateral symmetry, heart stage with cotyledon primordia emergence, torpedo stage with elongating cotyledons and axis, and mature embryo ready for seed desiccation and dormancy. Suspensor subsequent development shows basal cell elongation creating filamentous structure, limited cell division producing small suspensor cell number typically 6-10 cells, suspensor connecting embryo to seed tissues, possible nutritive and signaling functions, and eventual degeneration as embryo proper develops. Environmental requirements during genesis start include optimal temperature 22-28°C supporting normal division timing and progression, adequate nutrient supply from endosperm providing resources, appropriate osmotic and hydration conditions, and overall favorable environment during critical first division period. Temperature as major regulator shows warm conditions accelerating division enabling rapid embryogenesis, cool conditions substantially delaying division potentially extending seed development period, and temperature affecting not only timing but possibly division symmetry and developmental outcomes. Endosperm coordination shows endosperm development proceeding parallel to early embryogenesis, endosperm providing nutritive support for embryo divisions, signaling between tissues possibly coordinating development, and balanced embryo-endosperm development essential for normal seed formation. Failed genesis start causes include cell cycle defects preventing mitosis, abnormal division producing equal or inverted asymmetry disrupting proper patterning, cell fate determination failure preventing normal embryo-suspensor establishment, genetic abnormalities causing developmental arrest, or environmental stress disrupting division processes. Genesis success rate shows high efficiency in normal activated zygotes with majority successfully completing first division, genetic or environmental problems causing minority failures, and successful genesis essential for seed development. The process represents second and final aspect of zygote formation marking definitive embryogenesis initiation, following zygote development initiation. Duration represents culminating event with division occurring 24-72 hours post-fertilization establishing two-cell embryo. Cultivar variation may include division timing, asymmetry characteristics, and subsequent embryo developmental patterns. Regional timing shows embryo genesis start during late June-early July across all climates 1-3 days following fertilization as embryogenesis begins. Viticultural significance includes representing definitive embryogenesis initiation marking seed development beginning, requiring successful zygote activation and division, demonstrating temperature sensitivity affecting timing and potentially outcomes, establishing genesis success as determinant of seed and berry development, and marking transition from fertilization to embryogenesis and berry growth phases. Research applications include analyzing first division mechanisms and asymmetry, investigating cell fate determination and molecular asymmetry, evaluating environmental effects on division timing and characteristics, quantifying embryogenesis initiation dynamics, and assessing genesis success as determinant of seed development outcomes.
Embryo_Genesis_Start is second and final aspect of Zygote_Formation, distinguished by first division event initiating multicellular embryogenesis. Represents asymmetric mitotic cleavage producing apical and basal cells. Occurs 24-72 hours post-fertilization. Establishes embryo proper and suspensor lineages. Marks definitive transition from unicellular zygote to multicellular embryo. Temperature strongly affects timing. Success essential for seed development. Completes Fertilization sequence. Initiates Berry_Development phase with seed formation. Final documented process in Flowering hierarchy.
Embryo-Genesis-Start
Embryogenese-Initiierung
Zygoten-Spaltung
embryogenesis initiation
erste embryonale Teilung
first embryonic division
two-cell embryo formation
zygote cleavage
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 400-405 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 304-308
Embryo Genesis Start
Chardonnay in Champagne shows embryo maturation during September-October Year N+1, with structurally complete torpedo embryo beginning reserve accumulation week 12, lipids accumulating in cotyledons reaching 33-39% dry weight with oleic acid 18-24% and linoleic acid 48-58% total fatty acids, proteins accumulating 27-34% dry weight, lipid bodies 0.8-1.6 micrometers and protein bodies 1.2-2.4 micrometers forming, LEA proteins and protective sugars accumulating conferring desiccation tolerance, embryo growth slowing and ceasing, mature embryo 3.8-4.6 mm length with full reserves by weeks 16-18, under cool maritime conditions of 16-22°C temperature, typical of this early-bursting cultivar completing embryogenesis
Embryo maturation represents the final embryonic development phase during embryo development when structurally complete embryo accumulates storage reserves, acquires desiccation tolerance, reduces metabolic activity, and prepares for dormancy and germination, with cotyledons accumulating lipids 30-40% and proteins 25-35% dry weight, embryo growth slowing and eventually ceasing, desiccation tolerance proteins accumulating, and mature embryo ready for germination when conditions appropriate, occurring weeks 12-18 post-fertilization completing embryogenesis. This reserve accumulation and preparation phase occurs during late summer to early autumn in September-October in temperate regions Year N+1, representing final embryo development completing seed maturation. Maturation mechanism involves storage reserve biosynthesis and accumulation primarily in cotyledons, lipid body and protein body formation, desiccation tolerance acquisition through protective molecule accumulation, metabolic activity decline toward quiescence, possible dormancy imposition preventing premature germination, and integrated maturation preparing embryo for dispersal and germination. Embryo structure at maturation onset shows torpedo stage embryo 800-1200 micrometers with fully formed organs including radicle, hypocotyl, cotyledons, and shoot apex, all seedling structures present requiring reserve accumulation and physiological maturation, and structural foundation for maturation. Storage reserve accumulation shows lipids and proteins accumulating primarily in cotyledon cells, reserve biosynthesis from photosynthate imported from parent plant through seed coat and funiculus, massive accumulation over weeks 12-18 creating energy and nutrient stores, and reserves supporting germination and seedling establishment. Lipid accumulation in cotyledons shows triacylglycerols as major storage lipids reaching 30-40% cotyledon dry weight by maturation completion, oleic acid C18 per 1 and linoleic acid C18 per 2 as predominant fatty acids 70-85% total fatty acids, lipid biosynthesis in endoplasmic reticulum, lipid bodies 0.5-2 micrometers diameter forming in cytoplasm storing lipids, and lipids as major energy reserves providing 9 kcal/g. Fatty acid composition shows oleic acid typically 15-25% total fatty acids, linoleic acid 40-60%, linolenic acid 5-15%, palmitic acid 5-10%, stearic acid 2-5%, and fatty acid profile affecting seed quality and germination. Lipid body characteristics shows lipid bodies as organelles surrounded by phospholipid monolayer, triacylglycerols in core, oleosins and other proteins at surface, and lipid bodies as lipid storage compartments. Protein accumulation in cotyledons shows storage proteins reaching 25-35% cotyledon dry weight, albumins as water-soluble proteins, globulins as salt-soluble proteins, proteins providing amino acids for seedling growth during germination, protein biosynthesis and accumulation in protein bodies 1-3 micrometers diameter, and proteins as nitrogen and sulfur reserves. Protein body characteristics shows protein bodies as specialized vacuoles storing proteins, surrounded by tonoplast membrane, proteins crystalline or amorphous in bodies, and protein bodies as protein storage compartments. Amino acid composition shows balanced amino acid profile for seedling nutrition, essential amino acids present, arginine often abundant amino acid in storage proteins, and composition supporting germination requirements. Carbohydrate accumulation shows minor carbohydrate reserves typically <10% dry weight, sucrose and oligosaccharides as major carbohydrates, raffinose family oligosaccharides possibly present, and carbohydrates having storage and protective functions. Reserve accumulation timing shows accumulation initiating during late torpedo stage week 10-12, intensive accumulation weeks 12-16, completion by week 16-18, and accumulation as major maturation process. Embryo growth during maturation shows continued growth during early maturation increasing size and mass, growth slowing during reserve accumulation, eventual growth cessation by maturation completion, final mature embryo size 3-5 mm length and 1.5-3 mm width, and growth pattern characteristic of maturation. Fresh weight and dry weight dynamics shows embryo fresh weight increasing from water and reserve accumulation, dry weight increasing substantially from reserve deposition, eventual water loss during desiccation, and dry weight as reserve measure. Desiccation tolerance acquisition shows embryo acquiring ability to survive water loss during maturation, late embryogenesis abundant LEA proteins accumulating providing cellular protection during desiccation, sugars particularly sucrose and raffinose family oligosaccharides accumulating stabilizing membranes and proteins, heat shock proteins possibly accumulating, and desiccation tolerance enabling seed survival and storage. Late embryogenesis abundant proteins shows hydrophilic proteins accumulating during late maturation, LEA proteins protecting cellular structures during desiccation, proteins preventing protein aggregation and membrane fusion, and LEA proteins as desiccation protection. Sugar protective function shows sugars replacing water in hydrogen bonding stabilizing membranes, sugars forming glasses vitrifying cytoplasm preventing damage, and sugars as desiccation protectants. Metabolic activity decline shows respiratory rate decreasing during maturation, metabolic flux declining toward quiescence, gene expression shifting toward dormancy-associated genes, and metabolism declining to very low levels in mature desiccated embryo. Dormancy possible imposition shows embryos possibly acquiring dormancy preventing immediate germination, dormancy requiring stratification cold treatment or time for release, dormancy mechanisms including abscisic acid accumulation and germination inhibitors, and dormancy as germination control though variable in grapevine. Abscisic acid in embryo shows ABA possibly accumulating during maturation, ABA imposing or maintaining dormancy, ABA levels declining during stratification or after-ripening enabling germination, and ABA as dormancy hormone. Embryo color changes shows embryo progressing from white or pale during early maturation to yellowish or greenish at maturation from reserve accumulation and chlorophyll, color as maturation indicator, and color changing with desiccation. Maturation completion shows embryo achieving mature state with full reserve complement, desiccation tolerance acquired, metabolic quiescence established, embryo ready for dispersal and eventual germination, and completion by weeks 16-18 post-fertilization. Mature embryo characteristics shows embryo 3-5 mm length with fully formed structures, reserves stored in cotyledons, desiccation tolerant able to survive low moisture, dormant or germination-competent depending on regulation, and viable ready for germination. Germination competence shows mature embryo capable of germination if provided appropriate conditions including moisture, temperature, possible stratification, and germination competence as maturation outcome. Environmental effects on maturation include temperature affecting maturation rate with warm conditions accelerating reserve accumulation and cool conditions slowing, nutrient availability affecting reserve accumulation quality and quantity, and favorable conditions ensuring complete maturation. Temperature effects shows optimal range 22-26°C supporting normal maturation, temperature affecting maturation duration, and temperature as rate determinant. The process represents third and final phase of embryo development completing embryogenesis, following embryo morphogenesis. Duration approximately 6 weeks representing reserve accumulation and maturation weeks 12-18. Regional timing shows embryo maturation during September-October completing seed development. Viticultural significance includes representing reserve accumulation determining germination vigor, desiccation tolerance enabling seed survival and storage, dormancy potentially regulating germination timing, and maturation completing reproductive development creating viable seed. Research applications include analyzing storage reserve biosynthesis and regulation, investigating desiccation tolerance mechanisms and protective molecules, evaluating dormancy imposition and regulation, quantifying reserve accumulation and composition, and assessing embryo maturation as seed viability and quality determinant.
Embryo_Maturation is third and final phase of Embryo_Development, distinguished by reserve accumulation. Represents weeks 12-18. Storage reserves: lipids 30-40% (mainly oleic, linoleic acids in triacylglycerols), proteins 25-35% (albumins, globulins). Lipid bodies 0.5-2 μm, protein bodies 1-3 μm in cotyledons. Desiccation tolerance (LEA proteins, protective sugars). Metabolic decline. Possible dormancy. Mature embryo 3-5 mm. Follows Embryo_Morphogenesis. Completes Embryo_Development. Determines germination vigor.
Embryo-Reifung
Embryo-Reserven-Akkumulation
Reifungsphase
embryo preparation
embryo reserve accumulation
final embryo development
finale Embryo-Entwicklung
maturation phase
Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 441-442
Embryo Maturation
Syrah in Rhône Valley shows embryo morphogenesis during July-September Year N+1, with globular embryo 65-90 micrometers weeks 1-3, heart embryo 200-280 micrometers weeks 3-7 showing cotyledon primordia emergence and bilateral symmetry, torpedo embryo 650-1100 micrometers weeks 7-12 with elongated axis showing radicle, hypocotyl, enlarging cotyledons, and developing shoot apex, progressive tissue differentiation with protoderm, ground meristem, and procambium, under warm Mediterranean conditions of 24-29°C temperature, typical of this mid-to-late season cultivar completing morphogenesis with organized seedling structures ready for maturation
Embryo morphogenesis represents the embryonic structure formation phase during embryo development when proembryo undergoes progressive cell divisions and differentiation creating organized embryo with recognizable organs, proceeding through sequential morphological stages including globular undifferentiated embryo weeks 1-3, heart embryo with emerging cotyledon primordia weeks 3-7, and torpedo embryo with elongated axis and developing organs weeks 7-12, establishing seedling body plan with radicle, hypocotyl, cotyledons, and shoot apex. This structure formation phase occurs during summer to early autumn in July-September in temperate regions Year N+1, spanning approximately 12 weeks representing major morphogenesis period. Morphogenesis mechanism involves coordinated cell divisions increasing cell number, progressive cell differentiation establishing cell types, pattern formation creating spatial organization, organ initiation and development forming seedling structures, and developmental progression through morphological stages creating recognizable embryo architecture. Globular embryo stage shows spherical to ovoid embryo 50-100 micrometers diameter weeks 1-3 post-fertilization, undifferentiated cell mass with no visible organ primordia, cells small densely cytoplasmic actively dividing, protoderm surface layer differentiating, ground meristem and procambium beginning internal specification, and globular as early undifferentiated stage. Globular embryo development shows rapid cell divisions increasing cell number from 2-cell to hundreds of cells, cells relatively uniform without clear differentiation, embryo growth from 50 to 100 micrometers, and transition to heart stage with organ initiation. Cell division patterns during globular stage shows irregular division planes creating three-dimensional cell mass, divisions establishing radial symmetry, cell lineages not strictly determined enabling regulation, and division patterns creating embryo foundation. Protoderm formation shows outer cell layer differentiating as protoderm epidermis precursor, protoderm cells acquiring surface identity, periclinal divisions maintaining surface layer, and protoderm as first differentiated tissue. Ground meristem specification shows internal cells beginning ground tissue specification, ground meristem giving rise to cortex and pith, specification preparing for axis formation, and ground meristem as internal tissue precursor. Procambium initiation shows vascular tissue precursor cells beginning specification, procambium as centrally located cell files, procambium forming vascular connection through axis, and procambium as vascular precursor. Heart embryo stage shows bilateral symmetrical embryo 150-300 micrometers size weeks 3-7, cotyledon primordia emerging as two lateral bulges creating heart shape, embryonic axis visible between cotyledons, organ initiation marking developmental transition, and heart as organ formation stage. Cotyledon primordium emergence shows two lateral outgrowths from embryo body, cotyledon primordia as embryonic leaf structures, primordia establishing bilateral symmetry, and cotyledon initiation as defining heart stage feature. Bilateral symmetry establishment shows two cotyledons creating symmetry plane, symmetry from developmental patterning mechanisms, symmetry characteristic of dicotyledonous plants, and bilateral pattern distinguishing from radial globular symmetry. Embryonic axis formation shows axis region between cotyledons becoming apparent, axis containing shoot apical meristem at apex, radicle region differentiating at base, and axis as future seedling stem. Shoot apex initiation shows shoot apical meristem beginning formation at axis apex between cotyledon bases, meristem as source of post-embryonic shoot growth, meristem dome with organizing center, and shoot apex as critical structure. Radicle region differentiation shows basal axis region differentiating as radicle embryonic root, radicle region with root organization, future root cap initiating, and radicle as basal embryo structure. Torpedo embryo stage shows elongated embryo 500-1200 micrometers length weeks 7-12, embryonic axis elongating with 3-8 fold length to width ratio, cotyledons enlarging and beginning to curve, organs clearly differentiated and growing, and torpedo as major growth and differentiation stage. Embryonic axis elongation shows hypocotyl region between radicle and cotyledons elongating, axis length increasing substantially, axis showing clear radial organization, and elongation creating torpedo shape. Cotyledon development shows cotyledons enlarging from small primordia to substantial organs, cotyledons beginning to curve accommodating axis, cotyledons preparing for storage reserve accumulation, and cotyledon growth as major feature. Radicle differentiation completion shows radicle clearly defined with root cap forming at tip, root apical meristem establishing behind cap, procambium extending through radicle, and radicle ready for germination emergence. Hypocotyl formation shows hypocotyl as axis region between radicle and cotyledons, hypocotyl with characteristic tissue organization, vascular procambium continuous through hypocotyl, and hypocotyl as embryonic stem. Shoot apical meristem development shows meristem becoming clearly organized, meristem with tunica corpus organization typical of shoot apices, possible leaf primordia initiating though unclear in grapevine, and meristem as shoot growth source. Tissue differentiation during morphogenesis shows protoderm developing into epidermis, ground meristem differentiating into cortex and pith, procambium developing into vascular tissue, and progressive differentiation creating tissue types. Vascular tissue development shows procambium forming continuous strand through axis, xylem and phloem precursors differentiating, vascular connection from radicle through hypocotyl to cotyledons, and vascular system as transport pathway. Pattern formation shows spatial gene expression patterns directing morphogenesis, transcription factors regulating organ identity and polarity, hormonal signals including auxin affecting pattern, and coordinated patterning creating organized structure. Organ identity specification shows genetic programs determining organ fate, cotyledon identity genes expressed in cotyledons, root identity genes in radicle, shoot identity genes in shoot apex, and identity specification as organ determination. Morphogenesis completion shows embryo achieving characteristic seedling structure by week 12, organs differentiated ready for maturation, embryo transitioning to maturation phase with reserve accumulation, and morphogenesis establishing structure for subsequent maturation. Environmental effects on morphogenesis include temperature affecting development rate with optimal 22-28°C supporting normal progression through stages, nutrient availability supporting cell division and growth, and favorable conditions ensuring complete morphogenesis. Temperature effects shows warm conditions accelerating stage progression, cool conditions slowing development, and temperature determining morphogenesis duration. The process represents second phase of embryo development creating structures, following zygote division and preceding embryo maturation. Duration approximately 12 weeks representing globular through torpedo stages. Regional timing shows embryo morphogenesis during July-September as structures form. Viticultural significance includes representing organ formation creating seedling structures, establishing body plan for germination, demonstrating progressive developmental program, and completing structural organization preparing for maturation. Research applications include analyzing morphogenesis regulation and pattern formation, investigating organ identity specification and development, evaluating developmental stage progression and timing, quantifying morphological changes and growth, and assessing morphogenesis as seedling structure determinant.
Embryo_Morphogenesis is second phase of Embryo_Development, distinguished by structure formation. Represents progression through globular (50-100 μm, weeks 1-3), heart (150-300 μm, weeks 3-7), torpedo (500-1200 μm, weeks 7-12) stages. Creates radicle, hypocotyl, cotyledons, shoot apex. Tissue differentiation: protoderm, ground meristem, procambium. Duration ~12 weeks. Follows Zygote_Division. Precedes Embryo_Maturation. Temperature 22-28°C optimal. Establishes seedling body plan.
Embryo-Morphogenese
Embryo-Organogenese
Organ-Bildung
embryo organogenesis
organ formation
seedling structure creation
structural development
strukturelle Entwicklung
Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 440-442
Embryo Morphogenesis
An emergency reserve renewal cane on Vitis vinifera cv. Riesling in frost-prone German vineyard, retained at 7 nodes positioned 35cm from cordon (primary at 9cm, secondary at 18cm, backup reserve at 25cm), measuring 7mm diameter at minimum acceptable threshold, complete lignification but with slightly irregular node spacing, representing last-resort option rarely used (utilized in only 3 of past 20 seasons, all following severe spring frost damage exceeding -4°C killing 75%+ of primary renewal shoots), with typical outcome being complete removal during normal pruning but providing critical capacity during catastrophic damage events preventing total renewal system failure requiring multi-year recovery through latent bud development.
A reserve renewal cane retained specifically for catastrophic failure scenarios, characterized by low expectation of actual utilization (perhaps 5-10% selection probability in typical years), marginal positioning or quality making it last-resort rather than routine option, and function as ultimate backup when all preferred renewal options (primaries, secondaries, backup reserves) fail due to severe winter damage, spring frost, disease, or other major problems requiring utilization of any available wood to maintain training system continuity. Emergency reserve renewal canes represent extreme risk management acknowledging that occasional severe events exceed normal renewal system capacity, with emergency reserves providing final insurance against complete renewal system collapse requiring latent bud activation or drastic retraining, though their retention cost (bud capacity, pruning labor, canopy management) approaches economic viability limits given low utilization probability.
Emergency reserve renewal canes represent the extreme limit of renewal planning, with their retention reflecting either exceptional risk aversion (premium vineyards where training system preservation justifies high insurance costs), documented vulnerability to catastrophic events (frequent severe frost, hail, or disease pressure), or conservative management philosophy accepting significant opportunity costs for maximum system resilience. The low utilization probability (typically under 10% annually) raises questions about economic viability, with retention consuming bud capacity that could support productive fruiting positions or being reallocated to preferred renewal tiers. However, the catastrophic consequences of complete renewal failure (multi-year productivity loss, training system breakdown, potential vine removal) may justify emergency reserve retention in high-value situations where catastrophic risk cannot be otherwise managed.
Katastrophen-Vorratsrute
Notfall-Reserverute
catastrophic backup renewal
extreme contingency wood
last-resort renewal cane
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Molitor et al. (2014) "Late frost damage risk for viticulture under future climate conditions= a case study for the Luxembourgish winegrowing region" Australian Journal of Grape and Wine Research 20=160-168.
Emergency Reserve Renewal Cane
An emergency tertiary vegetative bud at node 8 on Vitis vinifera cv. Syrah following catastrophic spring frost (-4.2°C) combined with subsequent hail that destroyed 96% of primary buds and 84% of secondary buds across an entire vineyard block, this tertiary successfully breaking 19 days post-damage, developing into a weak but persistent shoot reaching 52cm with 9 leaves totaling 285 cm² photosynthetic area by season end, producing moderately well-differentiated buds during summer including one primary position showing possible inflorescence initials, enabling partial crop recovery (0.3 kg per vine, 18% of normal) in second season and progressive improvement to 1.4 kg (82% of normal) by fourth season post-damage.
A tertiary vegetative bud that successfully activates and develops following catastrophic destruction of primary and secondary positions, representing functional rather than merely anatomical backup capacity, characterized by confirmed developmental competence through actual budbreak and shoot production, weak but sustained vegetative growth providing minimal but non-zero contribution to vine survival and canopy recovery. Emergency tertiary vegetative buds demonstrate the practical value of compound bud architecture under extreme damage scenarios, with their successful activation enabling vine survival and establishing foundation for multi-year recovery through progressive canopy rebuilding, reserve accumulation, and bud differentiation leading to eventual return to productive status, though typically requiring 2-4 seasons for substantial yield recovery depending on damage extent and vine vigor.
Emergency tertiary vegetative buds represent the difference between vine survival and vine death in catastrophic damage scenarios, with their successful activation determining whether damaged blocks can recover or require replanting. The multi-year recovery timeline following tertiary activation reflects the cumulative nature of grapevine productivity - first year focuses on survival and minimal canopy (tertiary shoots), second year on improved canopy from better-developed buds formed on tertiary shoots, third year on establishing productive fruiting wood, and fourth year on approaching normal production. Economic analysis of tertiary-dependent recovery must account for 3-4 years of reduced or zero income against replanting costs and similar time-to-production for new vines, with decisions influenced by vine age, variety value, and vineyard site characteristics.
Notfall-Tertiärknospe
activated tertiary bud
aktivierte dritte Knospe
functional emergency bud
successfully breaking tertiary bud
Based on Molitor et al. (2014) "Late frost damage risk for viticulture under future climate conditions= a case study for the Luxembourgish winegrowing region" Australian Journal of Grape and Wine Research 20=160-168 Ferguson et al. (2011) "Dynamic changes in bud cold hardiness during dormancy and the influence of apple and pear budbreak" Journal of the American Society for Horticultural Science 136=313-323 Trought & Bramley (2011) "Vineyard variability in Marlborough, New Zealand= characterising spatial and temporal changes in fruit composition and juice quality in the vineyard" Australian Journal of Grape and Wine Research 17=79-89.
Emergency Tertiary Vegetative Bud
An emerging young leaf at the shoot apex of Vitis vinifera cv. Chardonnay in Champagne, France, measuring 1.8 cm length in tightly folded configuration representing approximately 12% of expected mature size, positioned 2 cm behind the apical meristem dome still partially protected by surrounding bud scale remnants and older emerging leaves, exhibiting very pale yellow-green coloration with pronounced bronze-red tinge from high anthocyanin concentration (4.2 mg g⁻¹ fresh weight) providing protection against photooxidative damage, extremely soft fragile texture with cells showing active division particularly in marginal meristematic zones establishing final lobe pattern, virtually no cuticle evident from completely dull non-reflective surface highly susceptible to water loss, microscopic examination revealing undifferentiated parenchymatous tissue without distinct mesophyll layers, non-functional stomata present but with incompletely developed guard cells, demonstrating minimal photosynthetic activity with CO₂ uptake barely detectable representing pure resource sink consuming approximately 12 mg carbohydrate per day for cell division and expansion, occurring at 4 days post-primordium initiation, measured at shoot BBCH stage 53 (inflorescence clearly visible) in early May, representing the earliest recognizable leaf developmental stage following meristematic initiation.
A young leaf in the earliest post-initiation developmental stages from initial visibility as a primordium through initial unfolding in Vitis vinifera, characterized by very small size typically 0.5-3 cm length representing 5-20% of final mature dimensions, position within or just emerging from protective bud scales or surrounding older leaf primordia, tightly folded or rolled architecture with unexpanded lamina, very pale green to reddish-bronze coloration from minimal chlorophyll and high anthocyanin concentrations providing photoprotection, extremely soft delicate texture with cells composed primarily of actively dividing meristematic tissue and rapidly expanding parenchyma, virtually no cuticle development leaving surfaces vulnerable to desiccation, completely undifferentiated internal anatomy without distinct palisade or spongy mesophyll layers, non-functional or minimally functional stomata, and negligible photosynthetic capacity contributing essentially no carbon gain while representing major resource sink for cell division and initial expansion. Emerging young leaves develop within the shoot apical meristem dome where new leaf primordia are initiated at regular intervals creating the phyllotactic spiral, progressing through initial cell division phases establishing basic leaf architecture including lobe patterns determined during primordial stages, followed by initial expansion beginning to unfold the compressed lamina, with the entire emerging phase typically spanning 7-12 days from primordium visibility through initial unfolding, representing the most vulnerable developmental period with extreme susceptibility to frost damage, desiccation, mechanical injury, and pest attack by mites and thrips that preferentially feed on tender meristematic tissues.
Emerging young leaves represent the transition from primordial initiation to recognizable leaf structures, distinguished from mature or expanding leaves by extremely small size, tight folding, minimal differentiation, and pure sink status without photosynthetic contribution. The emerging phase encompasses two distinct substages= primordium stage where leaf primordia are barely visible as small bumps on the apical meristem, and unfolding stage where initial expansion begins revealing basic leaf outline. Developmental timing from initiation through emergence completion varies with shoot vigor, temperature, and resource availability, typically 7-12 days under optimal conditions but extending to 15-20 days under stress. Emerging leaves show maximum vulnerability to environmental stresses, with frost temperatures below -2°C typically causing complete tissue death, desiccation rapidly occurring if relative humidity drops below 60%, and pest damage from mites or thrips often causing permanent deformations that persist through maturity. Protection of emerging leaves is critical for maintaining shoot extension and achieving target leaf area, requiring frost protection strategies in cool climates, careful pesticide application timing to prevent young tissue damage while controlling pests, and adequate water availability to prevent desiccation stress.
anfangsstadium Weinblatt
initial emergence leaf
neu-emergierendes junges Blatt
newly initiated leaf
primordial leaf
primordiales Blatt
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 43-59 Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 33-55.
Emerging Young Leaf
Field evaluation of Empoasca_Resistance_Disposition at the Edmund Mach Foundation, San Michele all'Adige, Italy, monitored Empoasca vitis population development and damage across 18 cultivars in replicated vineyard plots under natural infestation over three seasons without insecticide application. Nymphal counts (per 25 leaves) and adult sticky trap catches provided population metrics, while damage was assessed using a 0-5 visual scale for stippling severity and chlorophyll content measurements (SPAD readings) as physiological indicators. The susceptible cultivar 'Müller-Thurgau' supported high populations (mean 85 nymphs per 25 leaves at peak), severe stippling (score 4.2), and 35% reduction in chlorophyll content. 'Riesling' demonstrated enhanced Empoasca_Resistance_Disposition with 60% lower nymph counts (34 per 25 leaves), moderate stippling (score 1.8), and only 12% chlorophyll reduction. Laboratory choice tests confirmed that E. vitis adults showed reduced settling (45% fewer) and oviposition (62% fewer eggs) on 'Riesling' versus 'Müller-Thurgau' leaf discs. Microscopic examination revealed that 'Riesling' possessed 2.3-fold higher trichome density on abaxial leaf surfaces, potentially impeding leafhopper movement and feeding. The correlation between trichome density and Empoasca_Resistance_Disposition across all cultivars was significant (r = -0.68), suggesting that this morphological trait contributes substantially to leafhopper deterrence and could be targeted in breeding programs.
An Empoasca_Resistance_Disposition is a Leafhopper_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that limit feeding, oviposition, nymphal development, or damage by grape leafhoppers of the genus Empoasca and related typhlocybine leafhoppers (Erythroneura, Arboridia) that cause direct feeding injury through extraction of mesophyll and phloem cell contents. This disposition represents resistance to small (3-4mm) phloem-feeding leafhoppers that produce characteristic stippling damage visible as white to yellow speckling on leaves from emptied cells, progressing to leaf bronzing, marginal necrosis, and premature defoliation under heavy infestation. Unlike xylem-feeding sharpshooters, Empoasca and related genera are not known vectors of major grapevine pathogens, making their impact purely through direct feeding damage affecting photosynthetic capacity and, in severe cases, fruit ripening and vine vigor. Multiple Empoasca species affect grapevine in different regions: E. vitis (European grape leafhopper) in Europe, E. fabae (potato leafhopper) and Erythroneura elegantula (western grape leafhopper) in North America. Resistance mechanisms include leaf physical properties (trichome density, thickness, toughness) impeding settling and feeding, constitutive or induced chemical factors reducing palatability or nymphal survival, and rapid wound responses limiting damage from individual feeding events. Variation in leafhopper susceptibility among V. vinifera cultivars has been documented, with some traditional cultivars showing enhanced tolerance potentially reflecting historical selection in leafhopper-prone environments. Breeding for Empoasca_Resistance_Disposition could reduce insecticide requirements in regions with chronic leafhopper pressure.
Empoasca_Resistance_Disposition is_a Leafhopper_Resistance_Disposition. This class addresses resistance to phloem-feeding grape leafhoppers (Empoasca, Erythroneura, and related typhlocybine genera) that cause direct stippling damage, distinct from Glassy_Winged_Sharpshooter_Resistance_Disposition addressing xylem-feeding Pierce's disease vectors. Multiple Empoasca and Erythroneura species affect grapevine in different regions with similar damage symptoms. These leafhoppers are not known vectors of major grapevine pathogens, making their importance purely economic through direct damage. Resistance mechanisms often involve leaf surface properties, particularly trichome density, which can be readily assessed and selected in breeding programs.
Empoasca-Resistenz-Disposition
Erythroneura resistance
Kleinzikaden-Abwehr
Rebzikaden-Resistenz
grape leafhopper resistance
typhlocybine leafhopper resistance
Pavan F et al. (2000) Evaluation of damage caused by Empoasca vitis on grapevine, Integrated Control in Viticulture, IOBC/WPRS Bulletin 23:67-71 Daane KM, Williams LE (2003) Manipulating vineyard irrigation amounts to reduce insect pest damage, Ecological Applications 13:1650-1666 Fornasiero D et al. (2012) Empoasca vitis management in organic and integrated vineyards, IOBC/WPRS Bulletin 79:213-219
Empoasca Resistance Disposition
Pinot Noir in Oregon shows empty anther formation during early July Year N+1 at BBCH 68-69 over 3-4 days July 3-6, with withered brown anthers showing obvious empty locules visible through dehiscence slits, shrunken structures reflecting tissue dehydration and senescence, contrasting with visible berry development 14-20 mm diameter, under cool climate conditions of 20-25°C temperature, typical of this early-to-mid season cultivar transitioning definitively to post-reproductive state confirming completed male reproductive function 13-15 days after first dehiscence
Empty anther formation represents the post-exhaustion structural state when depleted anthers transition to withered senescent structures lacking functional pollen, creating characteristic empty appearance confirming completed male reproductive function and marking definitive post-reproductive anther condition. This concluding structural state occurs during completed pollen release in late June-early July in temperate regions Year N+1 following pollen exhaustion, developing over several days as exhausted anthers undergo senescence changes, corresponding to BBCH 68-69 complete anthesis stages. Empty anther characteristics show depleted locules completely lacking yellow pollen masses creating hollow appearance visible through dehiscence slits, withered shrunken anther structure reflecting progressive tissue dehydration and senescence, brown to gray-brown coloration contrasting with earlier yellow-brown fresh pollen-containing state, and overall visual evidence of completed reproductive function and post-exhaustion senescence. Structural changes creating empty appearance include tissue dehydration causing volumetric shrinkage with anthers becoming smaller and more compact, cell wall collapse eliminating turgid structure creating wrinkled shriveled appearance, progressive desiccation removing tissue moisture content, and overall transformation from functional reproductive organ to senescent depleted structure. Anther wall deterioration shows progressive breakdown with outer epidermis maintaining basic structure though becoming brittle and fragile, fibrous layer losing hygroscopic function following exhaustion with no further mechanical role, endothecium remaining structurally intact though non-functional, and overall tissue persisting in degraded state. Dehiscence slit persistence shows maintained longitudinal openings exposing empty locule interiors enabling visual confirmation of depletion, slit edges potentially showing deterioration as tissue senesces, and permanent structural modification from dehiscence process creating lasting evidence of reproductive function. Locular emptiness appearance shows hollow chambers visible through dehiscence slits, light-colored or translucent locule walls contrasting with earlier pollen-filled opaque appearance, occasional residual pollen remnants adhered to walls representing discharge failures, and overall obvious visual evidence of pollen depletion. Color transformation progression shows characteristic change from yellow-brown fresh pollen-containing anthers through intermediate tan brown during exhaustion to final brown or gray-brown empty senescent state, reflecting tissue aging and oxidative changes, with color providing visual indicator of reproductive status. Tissue senescence processes include declining metabolic activity as respiratory function reduces in post-reproductive anthers, progressive cellular breakdown with organelle degradation and membrane deterioration, enzymatic activity potentially contributing to tissue degradation, and overall physiological transition from active reproductive function to senescent degeneration. Dehydration dynamics show continued water loss following exhaustion with tissue moisture content declining substantially, hygroscopic responses potentially causing dimensional changes with atmospheric humidity fluctuations, and progressive desiccation creating characteristic withered shrunken appearance. Persistence on flower shows empty anthers typically remaining attached to filaments for extended periods despite senescence, occasional anther abscission removing senescent structures from flowers though many persist, and overall tendency for empty anthers to remain visible throughout berry development period providing lasting evidence of flowering. Observational value of empty anthers includes providing definitive visual confirmation of completed pollen discharge and male reproductive function, enabling post-flowering assessment of reproductive status, and serving as obvious marker that flowering concluded with developmental phase established. Empty anther distribution pattern at formation shows spatial gradient with earliest-emptied anthers in apical cluster regions showing most advanced senescence, progressive extension to middle and basal regions as sequential anthers exhaust and senesce, and overall cluster-level transition creating predominantly empty anther appearance by formation completion. Temporal development of empty state shows progressive appearance over 2-4 days following exhaustion as tissue changes proceed, accelerated formation under warm dry conditions promoting rapid senescence, and delayed formation under cool humid conditions extending fresh post-exhaustion state before obvious emptiness appears. Environmental effects on empty anther formation include temperature affecting senescence rate with warm conditions accelerating tissue deterioration, humidity affecting desiccation rate with dry conditions promoting rapid dehydration creating shrunken appearance, and overall conditions influencing appearance timing and characteristics. Functional significance of empty anther formation includes marking definitive end of male reproductive capacity confirming no further pollen production possible, providing visual evidence enabling reproductive phase assessment without detailed examination, and indicating developmental transition with established fruit set representing final reproductive outcomes. Relationship to fruit set outcomes shows empty anther formation coinciding with fruit set finalization, confirming that male gamete contribution completed with berry number established, and enabling assessment that observed fruit set represents complete potential from successfully pollinated flowers. Berry development relationship shows empty anther formation occurring while established berries undergo rapid early growth 12-20 mm diameter, contrasting appearance between senescing reproductive structures and vigorous developing fruit, and overall visual evidence of developmental phase dominance replacing reproductive function. Cluster appearance at empty anther formation shows mixed state with visible developing berries dominating appearance, brown senescing unfertilized flowers showing progressive abscission, empty withered anthers on successfully fertilized flowers providing brown accent, and overall transitional appearance documenting reproductive conclusion and developmental establishment. Management implications include confirming flowering completion through empty anther observation, recognizing that reproductive phase concluded enabling full operational resumption, and understanding that observed cluster state reflects final reproductive outcomes requiring no further pollination management. The process represents second and final aspect of completed pollen release emphasizing post-exhaustion structural state, following pollen exhaustion. Duration represents gradual development over several days as tissue senescence proceeds creating characteristic empty appearance. Cultivar variation in empty anther characteristics may include structural differences affecting appearance, senescence rate affecting formation timing, and persistence affecting duration that empty anthers remain visible on flowers. Regional timing shows empty anther formation during late June-early July in warm climates, early July in temperate regions, and early-mid July in cool areas, developing during complete anthesis as male reproductive structures transition to post-reproductive state. Viticultural significance includes providing obvious visual confirmation of completed male reproductive function, enabling practical assessment of flowering completion without detailed examination, and serving as definitive marker that developmental phase established with reproductive phase concluded. Research applications include documenting anther senescence processes and structural changes, analyzing environmental effects on post-reproductive anther characteristics, evaluating empty anther formation as practical marker for reproductive phase completion, and assessing relationships between anther senescence patterns and reproductive success outcomes.
Empty_Anther_Formation is second and final aspect of Completed_Pollen_Release, distinguished by post-exhaustion structural state confirming completed function. Represents gradual senescence process developing over several days. Provides obvious visual confirmation of reproductive conclusion. Enables practical assessment without detailed examination. Contrasts with developing berries showing developmental phase dominance. Completes entire Pollen_Release hierarchical documentation from dehiscence through peak release to exhaustion and final empty state. Definitively concludes male reproductive cycle.
depleted anther state
erschöpfte Antherenzustand
leere Antheren-Formation
post-exhaustion anther condition
senescent empty anthers
seneszente Staubbeutel
verwelkte Antheren-Bildung
withered anther formation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 393-398 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 298-300
Empty Anther Formation
Merlot flowers at BBCH 65-67 (full to late bloom, 50-75% of flowers past peak anthesis) exhibit empty pollen sacs visible as collapsed anther locules with wide-open longitudinal slits (0.2-0.3 mm), cream to light tan anther coloration indicating pollen depletion, and residual pollen adhered to locule walls representing approximately 15-25% of original pollen content (2,200-4,500 residual grains per anther versus 14,000-28,000 original content per five-anther flower), with empty sac appearance confirming that dehiscence occurred 1-4 days previously and that pollination window has passed. Cabernet Sauvignon at BBCH 69 (end of flowering, >80% flowers past anthesis) shows advanced empty pollen sacs with tan to brown coloration, extensively collapsed locule structures, and minimal residual pollen (<5% of original), providing visual evidence that bloom completed successfully and that fruit set determination phase is underway. Pinot Noir at 5-7 days post-peak bloom exhibits empty pollen sacs in various stages of senescence ranging from cream (recent dehiscence) to dark brown (earliest dehiscence), creating heterogeneous anther appearance reflecting acropetal bloom progression, with the range of empty sac colors providing retrospective information about bloom duration and timing patterns across the inflorescence.
An empty pollen sac is a pollen sac that has released its pollen content through anther dehiscence with locule cavity substantially or completely depleted of pollen grains, representing the post-dehiscence state (BBCH 61-69) when the pollen storage and release function has been fulfilled and when the pollen sac transitions from reproductive function to senescence and eventual degradation. Empty pollen sacs are characterized by= (1) locule cavity containing <30% of original pollen content (substantially empty) or <5% (completely empty), with most pollen having escaped through opened longitudinal dehiscence slits (2) collapsed or partially collapsed locule walls creating reduced cavity volume compared to the swollen pollen filled state, as internal pollen mass no longer provides outward pressure maintaining expanded configuration (3) residual pollen (when present) consisting of grains adhered to locule wall surfaces or trapped in cavity recesses, often showing degraded quality (reduced viability, desiccated appearance, or fungal colonization) (4) anther showing color change from bright yellow (pollen filled) to cream, tan, or brown (empty) as pollen depletion removes the dominant yellow pigmentation (5) wide-opened dehiscence slits (often 0.15-0.4 mm wide) with slit edges curled back or collapsed inward reflecting loss of internal support from pollen mass. The empty pollen sac state develops progressively following dehiscence initiation, with time course varying with environmental conditions and initial pollen quantity= under favorable dispersal conditions (moderate temperatures 22-28°C, declining humidity, light air movement), transition from pollen filled to substantially empty occurs within 6-18 hours as rapid pollen release proceeds under less favorable conditions (cool temperatures, high humidity, calm air), transition may require 12-36 hours as pollen release proceeds slowly. The empty state typically persists for 3-7 days (BBCH 65-69) before anther abscission occurs, representing the period when anthers have completed their reproductive function but structures remain attached to senescing filaments. Empty pollen sacs contain residual pollen typically representing 10-30% of original pollen content, with residual pollen showing characteristics distinct from released pollen= stronger adhesion to locule wall surfaces through hygroscopic or electrostatic forces tendency toward pollen grain aggregation or clumping rather than existing as free individual grains reduced viability (typically 40-65% viable compared to 70-85% in fresh released pollen) due to aging, desiccation, or environmental exposure potential for fungal colonization by Botrytis or other saprophytic fungi attracted to nutrient-rich residual pollen. The biological significance of empty pollen sacs is minimal as pollen release has completed, though several minor functions or effects include= residual pollen may contribute minimally to late pollination of late-receptive stigmas or late-opening flowers empty sac structures may serve as minor substrates for saprophytic fungi though small tissue mass limits epidemiological significance empty sac presence provides diagnostic evidence that normal dehiscence occurred and that bloom proceeded successfully. Empty pollen sac visual characteristics (cream to brown collapsed locules with wide-open slits) serve as useful retrospective indicators of bloom completion and pollination history, allowing observers to determine that flowers achieved anthesis and pollen release even when observations occur days after peak bloom when direct observation of active dehiscence is no longer possible. The progression from pollen filled through partially empty to completely empty sacs creates a temporal gradient of anther appearance (bright yellow → fading yellow → cream → tan → brown) that provides information about time since dehiscence and relative flower age, useful for assessing bloom progression patterns and for diagnosing bloom timing in relation to environmental events or management operations.
Empty_Pollen_Sac represents the post-dehiscence pollen-depleted state and is distinguished from Pollen_Filled_Sac (pre-release state with maximum pollen content) by substantial pollen depletion, collapsed locule structures, and color change from yellow to cream/tan/brown. The empty state indicates completed pollen release and fulfilled male reproductive function, with limited ongoing biological significance but with diagnostic value for retrospective assessment of bloom occurrence and timing. Understanding the progression from filled to empty pollen sacs and the visual indicators of this transition (color change, structural collapse, slit opening width) provides practical tools for field assessment of bloom status and for diagnosis of pollination history in relation to fruit set outcomes.
discharged pollen chamber
entleerter Mikrosporangium
erschöpfte Pollenkammer
leerer Pollensack
pollen-depleted sac
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 103-106 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 96-98.
Empty Pollen Sac
In Chardonnay vineyards of Burgundy following early-season primary shoot topping at node 12-14 to limit vegetative vigor, vigorous lateral shoots can develop 15-20 nodes by mid-summer, with 5-15% of lateral shoots developing end branch terminal clusters at lateral shoot apices during late July-August (BBCH 75-79 for primary crop), exhibiting small clusters of 5-8 cm length with 100-200 flowers that bloom 2-3 weeks after primary crop completion, typically achieving only 15-30% fruit set and producing small, immature berries that fail to ripen before harvest and are removed during harvest sorting. Gewürztraminer in Alsace under vigorous growing conditions occasionally exhibits end branch terminal clusters on lateral shoots, creating harvest complications when 8-12% of crop weight consists of immature secondary crop requiring selective harvest or mechanical removal.
An end branch terminal cluster is a terminal flower cluster positioned at the terminus of a lateral branch rather than at the primary shoot apex, representing flowers borne on inflorescences that develop at lateral shoot tips following lateral branch outgrowth, with this positioning affecting exposure, resource access, and developmental timing in ways distinct from both primary shoot lateral clusters and apex terminal clusters. End branch terminal clusters occur on inflorescences that develop at the apices of lateral shoots (secondary shoots arising from axillary buds on the primary shoot), representing conversion of lateral shoot apical meristems to reproductive development or development of inflorescences at lateral shoot tips. In standard Vitis vinifera viticulture, lateral shoots (also called secondary shoots or suckers) typically exhibit vegetative development, producing leaves and occasionally tendrils but rarely producing inflorescences under normal cultural management. However, end branch terminal clusters can develop under several circumstances= (1) on vigorous lateral shoots allowed to develop substantially before shoot positioning or removal, particularly if lateral shoot tips are exposed to high light conditions that may trigger floral conversion (2) following shoot tipping or topping of primary shoots that stimulates lateral shoot development, with subsequent lateral shoot tip conversion to reproductive development, particularly late in the season or under specific hormonal conditions (3) in second or third crop scenarios where lateral shoots developing late in the season undergo floral conversion and produce small terminal inflorescences that bloom substantially after the primary crop (4) under cultural practices that intentionally promote lateral shoot development and retention (e.g., certain sprawling or low-intervention training systems) where lateral shoots achieve sufficient development to support terminal cluster formation. End branch terminal clusters typically exhibit modified characteristics compared to primary shoot lateral clusters= cluster size is usually reduced (often 40-70% of primary lateral cluster size) reflecting abbreviated development time and competition with primary crop for resources, flower number is correspondingly reduced (often 80-300 flowers versus 200-800 in primary clusters depending on cultivar), and developmental timing is often delayed by 1-4 weeks or more, creating asynchronous flowering that complicates disease management and fruit maturation. The positional characteristics of end branch terminal clusters create distinctive environmental exposure= lateral shoot positions are often shaded within the canopy interior, potentially affecting flower development and fruit set through reduced light availability, or conversely, if lateral shoots extend beyond the primary canopy, end branch terminal clusters may experience enhanced light exposure but also greater wind and temperature fluctuation stress. Resource allocation to end branch terminal clusters typically involves competition with the primary crop, with late-developing lateral shoot clusters often exhibiting poor fruit set (20-40% versus 40-60% for primary clusters) and slow berry development, frequently failing to achieve maturity before harvest or winter dormancy. Viticultural management typically involves removal of lateral shoots and any associated end branch terminal clusters to eliminate resource competition with primary crop and prevent development of immature secondary crop that complicates harvest decisions and potentially harbors disease inoculum.
End_Branch_Terminal_Cluster is distinguished from Apex_Terminal_Flower_Cluster by position (lateral shoot apex versus primary shoot apex) and from standard lateral clusters by position (terminal on lateral shoot versus lateral on primary shoot) and developmental context (late-developing secondary structures versus normally programmed primary structures). End branch terminal clusters represent a generally undesirable developmental outcome in quality wine production, as they compete with primary crop for resources, complicate disease management through extended flowering period, and produce immature fruit that dilutes quality if included in harvest or requires costly selective removal. Cultural practices typically aim to prevent end branch terminal cluster development through timely lateral shoot removal, appropriate crop load management to limit excessive vegetative vigor that drives lateral shoot development, and shoot positioning to maintain lateral shoots in shaded positions that discourage floral conversion. Understanding the environmental and hormonal triggers for end branch terminal cluster development could inform precision management strategies for preventing or managing secondary crop development.
Blütentraube an Seitentriebspitze
Lateraltrieb-Blütentraube
lateral-shoot terminal cluster
secondary-shoot flower cluster
seitentrieb-endständige Blütentraube
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 102-106 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 83-86 Poni, S. et al. (1994). Effects of shoot trimming on leaf area, leaf efficiency, and yield components in Cabernet Franc. American Journal of Enology and Viticulture 45=193-200.
End Branch Terminal Cluster
Pinot Noir end point vascular bundle in basal mesocarp at berry maturity (BBCH 89) consists of single xylem vessel 12-15 μm diameter (embolized, non-functional) and 2 phloem sieve tubes 8-12 μm diameter (callose-blocked, minimal transport), embedded in parenchyma 2-3 mm from brush center, representing absolute termination of vascular trace with no further organized vascular elements distally
An end point vascular bundle is the ultimate terminal segment of a vascular bundle representing the final conducting elements where xylem and phloem transport pathways definitively conclude, characterized by minimal organized vascular structure, direct connection to surrounding parenchyma or specialized cells, and cessation of long-distance transport function. End point vascular bundles represent the finest ramifications of berry vascular architecture where= individual xylem vessel elements terminate through blind endings or connections to surrounding cells, individual phloem sieve tube elements conclude their transport pathway through unloading to adjacent tissues, organized vascular bundle structure transitions to scattered individual conducting cells or complete dissolution into parenchyma, and vascular supply transitions from organized bundle transport to local cell-to-cell diffusion. End point vascular bundle anatomical characteristics include= minimal vascular organization (1-2 xylem vessels, 1-3 phloem sieve tubes, often lacking distinct bundle sheath or organized arrangement), very small conducting element diameter (terminal xylem vessels 8-18 μm, terminal phloem sieve tubes 6-15 μm, both smaller than typical functional elements), high proportion of parenchyma cells (70-90% of bundle cross-section versus 30-50% in organized bundles), lack of cambial tissue (no potential for secondary growth), and gradual or abrupt transition to non-vascular tissue. End point vascular bundle xylem elements show distinctive features= single or few vessel elements (1-3 vessels representing absolute terminal xylem), reduced vessel diameter (8-18 μm, approaching tracheary element minimum functional size), blind vessel endings (vessels terminating without continuation), increased pit density in terminal vessel walls (facilitating water release to surrounding tissues), and high susceptibility to embolism (small diameter vessels readily cavitate under water stress or during ripening). End point vascular bundle phloem elements include= minimal sieve tube number (1-3 sieve tubes at absolute endpoint), reduced sieve tube diameter, few or single companion cells, high callose deposition potential (terminal phloem readily accumulates callose blocking transport), and direct connection to surrounding parenchyma for solute unloading. End point vascular bundle distribution in berry includes= primary end points in receptacle/brush region where major vascular traces conclude (4-8 per berry), secondary end points distributed through basal berry mesocarp (10-20 per berry), peripheral end points in exocarp or distal mesocarp regions (potentially 30-50+ per berry representing finest vascular ramifications), and potential end points near seed attachment sites. End point vascular bundle function emphasizes= final delivery point for xylem-transported water and minerals (end point xylem releases water to surrounding berry tissues through pit connections), final delivery point for phloem-transported sugars and nutrients (end point phloem unloads solutes to berry parenchyma), transition from vascular to apoplastic or symplastic transport (from organized long-distance transport to local cellular movement), and sites of transport regulation (end point unloading may be regulated by sugar concentrations, hormones, or developmental signals). End point vascular bundle developmental changes include= establishment during early berry development (BBCH 71-77) as vascular architecture differentiates, functional maturity during berry growth (BBCH 77-85) with active xylem and phloem transport, and potential dysfunction during late ripening (BBCH 85-89) through xylem embolism, phloem callose blockage, or complete vascular disconnection.
End_Point_Vascular_Bundle emphasizes ultimate conducting segment and is distinguished from Terminal_Vascular_Bundle (distal bundle region) and Vascular_Bundle_Termination (complete ending zone) by focus on absolute final elements where organized vascular transport definitively ceases. Critical for understanding limits of vascular supply distribution, solute unloading sites, berry tissue regions dependent on cell-to-cell transport, and vascular network extent in berry anatomy.
Endpunkt-Leitbündel
Final conducting element
Letzte Leitelemente
Ultimate vascular termination
Ultimatives Gefäßbündel
Chatelet, D. S., Rost, T. L., Shackel, K. A., & Matthews, M. A. (2008). The peripheral xylem of grapevine (Vitis vinifera). 1. Structural integrity in post-veraison berries. Journal of Experimental Botany, 59(8), 1987-1996. | Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press.
End Point Vascular Bundle
Räuschling Rot in Switzerland bearing an endangered cultivar role survives as fewer than 30 documented vines in a single historic parcel near Zürich identified through systematic cantonal vineyard surveys in 2015, with the cultivar confirmed as genetically distinct from white Räuschling through SSR profiling at 20 loci, emergency conservation measures including collection of 50 hardwood cuttings for propagation at Agroscope Wädenswil, establishment of 0.05 ha backup planting from grafted material, and cryopreservation of meristem tips to prevent total loss of this unique red-berried genotype documented in Swiss ampelographic records since 1654.
A conservation role that inheres in a grapevine cultivar whose total surviving population has declined to critically low levels — typically fewer than 50-100 individual vines or fewer than 1-2 hectares of commercial planting — placing the cultivar at imminent risk of irreversible genetic extinction without active conservation intervention.
Endangered_Cultivar_Role applies IUCN-analogous threat assessment criteria to cultivated grapevine genetic diversity, though no formal Red List system exists for Vitis vinifera cultivars. The role addresses the critical conservation gap where cultivars decline below viable population thresholds without triggering the regulatory protections available for wild species: cultivated plant diversity falls outside the scope of most endangered species legislation, making conservation dependent on voluntary institutional efforts. Quantitative endangerment thresholds are necessarily approximate — for clonally propagated grapevines, even a single surviving vine theoretically contains the complete genotype, but effective conservation requires multiple individuals to guard against accidental loss, disease, or extreme weather events.
akut bedrohte Sorten-Rolle
at-risk cultivar role
critically endangered cultivar role
extinction-threatened variety role
gefährdete Sorten-Rolle
vom Aussterben bedrohte Rebsorten-Rolle
Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157 Spring JL et al. (2019) Prospection et sauvegarde du patrimoine viticole en Suisse, Revue suisse de viticulture arboriculture horticulture 51: 236-243 FAO (2010) Second Report on the State of the World's Plant Genetic Resources for Food and Agriculture, Rome
Endangered Cultivar Role
Cabernet Sauvignon endocarp at maturity comprises 5-7 cell layers (total 150-220 μm thickness) lining locular cavities containing 2-3 seeds per berry, with cells showing systematic size decrease from outermost (75-92 μm diameter) through middle (62-78 μm) to innermost (52-68 μm) adjacent to seeds, walls 0.9-1.3 μm thick (intermediate between middle mesocarp 0.5-0.7 μm and outer mesocarp 1-1.4 μm), containing tannins 4.5-6.5 mg/g dry weight (substantially elevated above middle mesocarp 0.4-0.8 mg/g), contributing approximately 18-24% of wine skin tannins during standard 14-18 day maceration with endocarp extraction occurring primarily during second week as cells break down releasing phenolic contents. Syrah endocarp tannin gradient demonstrating systematic increase toward seed proximity= outermost endocarp (200-280 μm from seed surface) with 3.2-4.5 mg/g tannins middle endocarp (100-200 μm) with 4.8-6.2 mg/g innermost endocarp (0-100 μm from seeds) with 5.8-8 mg/g, showing progressive tannin concentration increase approaching seeds potentially reflecting seed influence on pericarp phenolic metabolism or accumulation, with tannin enrichment gradient extending 200-300 μm into pericarp from seed-adjacent position creating distinct innermost tissue zone. Merlot endocarp developmental seed contact changes= early development (BBCH 73-75, weeks 3-6 post-fruit set) showing developing seeds filling locular cavities with seed coats directly contacting endocarp cells creating intimate tissue interface mid development (BBCH 77-81, weeks 6-10) showing seeds completing maturation while pericarp continues expansion beginning tissue separation maturity (BBCH 89, week 18) showing 20-45 μm air-filled locular cavity gaps between mature seed surfaces and innermost endocarp cells as berry expanded beyond seed dimensions, demonstrating temporal progression from direct seed contact to seed proximity affecting potential for seed-pericarp biochemical interactions during berry development. Chardonnay white cultivar endocarp showing presence in white grapes= 4-5 cell layers, 135-195 μm total thickness, cells 54-88 μm diameter, tannins 2.8-4.5 mg/g (elevated above middle mesocarp 0.4-0.9 mg/g but lower than red cultivar endocarp 4-7 mg/g), demonstrating endocarp as distinct tannin-enriched innermost pericarp zone present in both red and white cultivars though typically not extracted during white wine production where immediate pressing separates juice from solids avoiding endocarp phenolic extraction. Pinot Noir endocarp contribution to wine tannins during maceration time-course= day 5 maceration extracting primarily outer pericarp tannins with minimal endocarp extraction day 10 showing beginning endocarp cell breakdown contributing 8-12% of wine skin tannins day 15 showing substantial endocarp extraction contributing 20-28% of wine skin tannins, demonstrating that endocarp represents later-extracting tannin reservoir requiring extended maceration for full extraction with extraction timing allowing winemakers to control wine tannin levels and structure through maceration duration management balancing desired extraction depth against potential over-extraction risks.
An endocarp is the innermost layer of the fruit pericarp in direct contact with or immediately adjacent to seeds, essentially synonymous with inner pericarp in grape berry anatomy, typically 100-300 μm thickness comprising 3-8 cell layers, characterized by cells smaller than middle mesocarp (40-90 μm diameter versus 80-180 μm), moderately thick walls (0.8-1.5 μm versus 0.4-0.8 μm in middle mesocarp), and elevated tannin content (3-7 mg/g versus 0.3-1 mg/g in middle mesocarp), serving as boundary tissue between fleshy mesocarp and seed-containing locular cavities while contributing to wine tannin structure during extended maceration. Endocarp anatomical organization comprises= (1) innermost mesocarp cell layers arranged parallel to locular cavity surfaces creating cellular boundary between flesh and seed chambers (2) progressive cell size decrease from outermost endocarp near middle mesocarp boundary (70-90 μm diameter) through middle endocarp (55-75 μm) to innermost endocarp adjacent to seeds or locular cavities (40-60 μm) (3) progressive wall thickness increase from 0.8-1 μm approaching middle mesocarp to 1.2-1.5 μm in innermost layers adjacent to seeds (4) total endocarp representing 4-7% of pericarp volume, small proportion but compositionally distinct from middle mesocarp bulk (75-85% of volume) (5) variable layer number (3-8 layers) depending on berry size, cultivar characteristics, and developmental conditions affecting pericarp thickness. Endocarp compositional characteristics= (1) tannin content 3-7 mg/g dry weight substantially elevated above middle mesocarp (0.3-1 mg/g) and intermediate between middle mesocarp minimum and outer pericarp maximum (8-15 mg/g), with tannins predominantly procyanidins showing polymerization degree 10-16 subunits similar to skin tannins (2) sugar and acid content similar to middle mesocarp (18-26% sugars, 0.5-0.9% organic acids) showing minimal radial compositional gradients in primary metabolites despite significant gradients in cell morphology and secondary metabolites (3) anthocyanins essentially absent (10-15 days) when deeper tissue extraction occurs (4) transition zone creating compositional and structural gradient from soft large-celled middle mesocarp to smaller firmer endocarp cells approaching seeds, preventing abrupt tissue property changes. Endocarp development follows berry growth= early development (Phase I, BBCH 71-75) with endocarp cells undergoing modest division establishing cell population expansion during ripening (Phase II, BBCH 81-89) with cells showing 80-140% diameter increase (less than middle mesocarp 180-250% increase) creating smaller final size despite similar initial size tannin synthesis and accumulation during ripening creating elevated tannin levels at maturity locular cavity air space development during late ripening as berry expansion exceeds seed size creating 10-50 μm gaps separating innermost endocarp from seed surfaces in mature berries though direct contact maintained during early-to-mid development. Endocarp in winemaking= endocarp tannins extracting during extended maceration typically after 10-15 days as endocarp cells require more time and vigorous conditions to break down than readily-extracting middle mesocarp endocarp contributing 15-25% of total wine skin tannins during extended maceration beyond outer pericarp contribution (45-60%) and middle mesocarp contribution (2-5%) endocarp tannins showing properties similar to outer pericarp skin tannins (higher polymerization degree, less astringent than seed tannins with degree of polymerization 4-10) contributing to wine structure and aging potential through phenolic complexity.
Endocarp is essentially synonymous with Inner_Pericarp and Inner_Mesocarp in grape berry anatomy, representing the innermost pericarp layer in contact with or adjacent to seeds. Sub-classification into Seed_Coat_Adjacent_Tissue (emphasizing seed proximity) and Locular_Tissue (emphasizing cavity boundary function) reflects different functional aspects. Understanding endocarp structure and composition is relevant for detailed berry anatomical characterization, for comprehending spatial tannin distribution affecting wine phenolic profiles, and for winemaking extraction strategies where endocarp represents intermediate-to-difficult extraction requiring extended maceration protocols.
Endokarp
Samenschicht
inner fruit wall
innere Fruchtwand
seed-adjacent layer
Cadot, Y. et al. (2006). Tannin composition and localization in grape skins and seeds. Journal of Agricultural and Food Chemistry 54':2973-2987 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 180-185 Downey, M.O. et al. (2003). The relationship between skin tannin concentration and berry size. American Journal of Enology and Viticulture 54':114-119.
Endocarp
In Pinot Noir vines of the Côte d'Or, Burgundy, endodormancy extends from mid-November through late January, with buds showing no growth response when excised and placed under forcing conditions of 22°C and 16-hour photoperiod during December, requiring approximately 1000-1200 chilling hours below 7.2°C accumulated during this period before growth competence is restored and transition to ecodormancy occurs in late January to early February.
Endodormancy in Vitis vinifera represents the phase of winter dormancy during which growth inhibition is controlled by factors internal to the bud itself, rendering buds incapable of growth even when exposed to environmental conditions that would otherwise be favorable for development. This dormancy phase, also termed true dormancy or rest in older literature, typically extends from late autumn through mid-winter in Northern Hemisphere viticulture, persisting until sufficient chilling exposure has satisfied the genetically determined chilling requirement and restored growth competence. The defining characteristic of endodormancy is the internal nature of growth inhibition, distinguishing it from ecodormancy where external environmental constraints prevent growth of otherwise competent buds, and from paradormancy where inhibitory signals from other plant parts suppress growth. The physiological basis of endodormancy involves multiple reinforcing mechanisms including elevated abscisic acid concentrations that directly inhibit cell cycle progression and growth-related gene expression, high expression of dormancy-associated MADS-box genes whose protein products actively maintain the dormant state, chromatin modifications that establish repressive configurations at growth-promoting gene loci, and altered cellular communication through modified plasmodesmatal function. During endodormancy, buds contain fully formed primordial shoots with leaf, tendril, and inflorescence primordia that developed during the previous growing season, but these structures remain arrested in development regardless of temperature, photoperiod, or nutrient availability until endodormancy is released through chilling accumulation. The ecological function of endodormancy is to prevent premature growth during temporary warm periods in autumn and winter that would expose sensitive growing tissues to subsequent lethal freezing, with the chilling requirement ensuring that growth does not resume until winter has sufficiently progressed that the probability of subsequent severe cold is reduced. The depth and duration of endodormancy vary substantially among cultivars, reflecting adaptation to different climatic regions, with cultivars from continental climates typically exhibiting deeper endodormancy requiring more chilling for release than cultivars from Mediterranean regions. Climate change poses significant challenges to endodormancy in many viticultural regions, as warming winters may provide insufficient chilling for complete dormancy release, potentially leading to delayed, erratic, or incomplete budbreak with consequences for vine productivity and fruit quality.
Endodormancy represents the second major phase within the Dormancy hierarchy following Dormancy_Induction, characterized by internally-controlled growth inhibition. Distinguished from Ecodormancy where growth-competent buds are environmentally constrained, and from Dormancy_Induction which represents the transition into dormancy. Subdivides into Deep_Endodormancy, Chilling_Accumulation, Dormancy_Maintenance, and Chilling_Requirement_Satisfaction as component phases. Duration typically spans 2-4 months depending on cultivar and chilling accumulation rate. Release requires cultivar-specific chilling accumulation typically ranging from 400-1500 hours below 7.2°C.
Endodormanz
echte Dormanz
innere Dormanz
internal dormancy
rest
true dormancy
Endodormancy
In Grüner Veltliner vines in the Wachau, Austria, endodormancy completion occurs in early February when accumulated chilling of approximately 1100 hours has satisfied the cultivar requirement, confirmed by forcing tests showing budbreak within 12 days at 22°C, with subsequent field observations showing buds remaining in ecodormancy until late March warming triggers the transition through BBCH stage 01 to visible bud swelling at BBCH 05.
Endodormancy completion in Vitis vinifera represents the termination of the endodormant phase when sufficient chilling has been accumulated to fully dismantle dormancy-maintaining mechanisms, marking the end of internally-controlled dormancy and the beginning of the ecodormant phase where only environmental factors constrain growth. This transition typically occurs during late January to February in Northern Hemisphere viticulture, when cumulative chilling exposure has satisfied the genetically determined requirement and all physiological changes associated with dormancy release have been accomplished. The molecular markers of endodormancy completion include the reduction of ABA concentrations to levels characteristic of ecodormant or growing buds, cessation of DAM gene expression, removal of repressive chromatin modifications from growth gene promoters, and restoration of cellular communication pathways that were restricted during endodormancy. The physiological state following endodormancy completion is characterized by buds that are metabolically prepared for growth resumption, with cell cycle machinery poised for activation, biosynthetic enzyme systems ready for rapid upregulation, and hormone signaling systems sensitized to respond to the warming temperatures of spring. The verification of endodormancy completion relies on forcing tests demonstrating rapid budbreak response comparable to that of ecodormant reference material, with completion indicated by budbreak times of less than 14 days under optimal forcing conditions contrasting with the extended times characteristic of endodormant buds. The timing of endodormancy completion relative to the onset of spring warming significantly affects budbreak date and potential frost exposure, with early completion in warm winters potentially advancing budbreak into periods of frost risk, while late completion in cold winters may delay budbreak and shorten the growing season. The concept of completion emphasizes the discrete transition between dormancy phases, though in physiological reality the transition is gradual with dormancy depth continuously declining until it crosses the threshold defining endodormancy release. Following endodormancy completion, buds enter ecodormancy where they remain quiescent due to environmental temperature limitation rather than internal inhibition, awaiting the warming of spring to initiate visible growth.
Endodormancy_Completion represents the terminal transition of Chilling_Requirement_Satisfaction marking the end of endodormancy phase. Distinguished from Dormancy_Release_Competence which describes the achieved capacity, while Endodormancy_Completion emphasizes the phase transition. Marks beginning of ecodormancy phase. Verified by forcing response less than 14 days. Timing affects budbreak date and frost risk. Molecular markers include low ABA, minimal DAM expression, and derepressed chromatin state.
Endodormanz-Abschluss
Endodormanz-Beendigung
echte Dormanz-Ende
endodormancy ending
endodormancy termination
innere Dormanz-Abschluss
internal dormancy completion
true dormancy completion
Endodormancy Completion
Endodormancy depth assessment comparing chilling accumulation effects in Riesling reveals the progressive release of internal dormancy, with 200 chill units accumulated showing deep endodormancy (forcing test: 15% budbreak after 45 days, highly erratic timing), 500 chill units showing moderate depth (55% budbreak after 28 days, improving uniformity), 800 chill units showing shallow endodormancy (85% budbreak after 18 days, relatively uniform), and 1100 chill units showing complete endodormancy release (98% budbreak after 12 days, highly synchronous), demonstrating the quantitative relationship between chilling accumulation and endodormancy depth reduction that underlies budbreak prediction models.
Endodormancy depth quality is a dormancy depth quality that represents the intensity of internally controlled dormancy in grapevine buds, characterizing the physiological block to growth that cannot be overcome by favorable environmental conditions alone but requires accumulation of chilling temperatures to progressively release the internal inhibition and restore growth competence. Within the BFO framework, this quality specifically addresses the internal regulation dimension of dormancy, distinguishing endodormancy from environmentally imposed quiescence while recognizing that endodormancy depth reflects the magnitude of the internal growth inhibition that must be overcome through chilling exposure before buds regain the capacity to respond to growth-promoting conditions. The ontological significance lies in endodormancy providing the robust protection against premature growth that is essential for perennial survival in temperate climates, where the internal nature of growth inhibition ensures that even extended warm periods during winter cannot trigger growth as long as endodormancy remains deep, with this protection gradually released only after sufficient chilling confirms that winter has progressed far enough to reduce the probability of subsequent damaging cold. The physiological basis for endodormancy involves hormonal changes (elevated ABA levels inhibiting growth, declining gibberellin sensitivity), chromatin modifications (epigenetic changes affecting gene expression), and metabolic shifts (altered carbohydrate and lipid metabolism) that together create the internal barrier to growth, with chilling exposure progressively reversing these changes to restore growth competence through mechanisms that remain incompletely understood but involve both hormonal rebalancing and chromatin remodeling. The measurement of endodormancy depth involves forcing tests where buds are exposed to growth-promoting conditions (typically 20-25°C with adequate moisture) and the time to budbreak, percentage of buds breaking, and uniformity of response indicate the depth of remaining dormancy, with deep endodormancy producing no response or very delayed, erratic budbreak, and shallow or released endodormancy producing rapid, uniform budbreak. Understanding endodormancy depth quality enables assessment of chilling progress through winter, prediction of when buds will become responsive to spring temperatures, and recognition of the climate change risks from insufficient chilling in warming winters that may leave buds in incompletely released endodormancy with consequences for budbreak uniformity and vine productivity.
Endodormancy_Depth_Quality serves as the parent class for chilling-related qualities (Requirement, Accumulation, Release Timing). This quality addresses the internally controlled dormancy requiring chilling for release. The class supports winter dormancy assessment through endodormancy depth characterization.
Endodormanztiefe
echte Dormanztiefe
innere Dormanzintensität
internal dormancy intensity
physiological dormancy
physiologische Dormanz
rest intensity
true dormancy depth
Dokoozlian NK (1999) Chilling temperature and duration interact on the budbreak of 'Perlette' grapevine cuttings. HortScience 34(6): 1054-1056 Pérez FJ et al. (2008) The expression of VvPHYA and VvPHYB transcripts is differently regulated by photoperiod in leaves and buds of grapevines. Plant Science 174(3): 259-263
Endodormancy Depth Quality
In Cabernet Franc vines in the Loire Valley, endodormancy persistence is demonstrated through January when buds excised on January 15 after accumulation of only 600 chilling hours require 38 days for 50% budbreak under forcing conditions, compared to buds excised February 15 after 1000 chilling hours requiring only 15 days, illustrating the continued persistence of endodormancy at insufficient chilling levels and its eventual release as chilling accumulation satisfies the cultivar requirement of approximately 950 hours.
Endodormancy persistence in Vitis vinifera represents the continued maintenance of the true dormant state characterized by internal growth inhibition that resists release by favorable environmental conditions, persisting until sufficient chilling has been accumulated to satisfy the genetically determined chilling requirement. This persistence reflects the robustness of dormancy-maintaining mechanisms that have evolved to prevent premature growth during the unpredictable temperature fluctuations characteristic of temperate winters, ensuring that growth does not resume until winter has sufficiently progressed that subsequent severe cold is unlikely. The molecular basis of endodormancy persistence involves the stable maintenance of dormancy-related gene expression patterns including sustained DAM gene activity, persistent chromatin modifications that silence growth genes, and continued production of growth-inhibiting proteins and metabolites that accumulate faster than they are degraded under conditions of insufficient chilling. The concept of persistence emphasizes the resistance of endodormancy to release, distinguishing this phase from ecodormancy where growth is prevented by environmental limitation and would occur immediately upon exposure to favorable conditions, whereas endodormant buds fail to grow even under optimal forcing conditions until internal changes associated with chilling have released the dormant state. Testing for endodormancy persistence involves exposing buds to growth-promoting conditions and confirming continued growth inhibition, with persistent endodormancy indicated by extended times to budbreak or complete failure to break that demonstrates internal control of dormancy status. The duration of endodormancy persistence is determined by the interaction between chilling requirement genetics and environmental chilling exposure, with persistence continuing until accumulated chilling has sufficiently reduced dormancy depth that maintaining mechanisms can no longer prevent growth, at which point the transition to ecodormancy occurs. Factors affecting endodormancy persistence include cultivar genetics that determine both dormancy depth and chilling requirement, environmental conditions during dormancy induction that affect the completeness of dormancy establishment, and the rate of chilling accumulation during winter that determines how rapidly maintaining mechanisms are dismantled. The ecological significance of endodormancy persistence lies in protecting the vine from responding to false spring conditions, with more persistent dormancy providing greater protection against freeze damage following premature growth but also requiring more chilling for release and potentially delaying spring growth in regions with limited winter cold.
Endodormancy_Persistence represents the temporal continuity aspect of Dormancy_Maintenance emphasizing resistance to premature release. Distinguished from Deep_Endodormancy which represents maximum dormancy intensity, while Endodormancy_Persistence addresses the duration and robustness of dormancy through the winter period. Measured through forcing test response showing continued growth inhibition. Duration determined by interaction of chilling requirement and chilling exposure. Provides ecological protection against false spring growth.
Endodormanz-Persistenz
anhaltende Endodormanz
continued true dormancy
endodormancy continuation
fortgesetzte echte Dormanz
persistent internal dormancy
persistierende innere Dormanz
sustained endodormancy
Lang GA et al (1987) Endo-, para-, and ecodormancy: physiological terminology and classification for dormancy research. HortScience 22: 371-377 Lavee S, May P (1997) Dormancy of grapevine buds - facts and speculation. Australian Journal of Grape and Wine Research 3: 31-46 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235
Endodormancy Persistence
Endodormancy release timing assessment comparing vintages in Pinot Noir reveals the year-to-year variation affecting growing season establishment, with cold winter showing early release (chilling requirement satisfied by January 15, long ecodormancy period, budbreak ultimately determined by spring temperatures arriving April 5), mild winter showing delayed release (requirement not satisfied until February 28, shortened ecodormancy, budbreak April 12 despite warmer March from late responsiveness onset), and average winter showing intermediate release (February 5, typical ecodormancy duration, budbreak April 2), demonstrating how endodormancy release timing interacts with subsequent temperatures to determine budbreak date through the combined effects of chilling satisfaction and heat accumulation.
Endodormancy release timing quality is an endodormancy depth quality that represents the temporal point during winter when chilling accumulation satisfies the endodormancy requirement and buds transition from internally controlled dormancy to environmentally controlled ecodormancy, characterizing the timing of this critical physiological transition that determines when buds become responsive to subsequent heat accumulation and thus establishes the framework for budbreak timing prediction. Within the BFO framework, this quality specifically addresses the transition timing dimension of endodormancy, distinguishing when release occurs from the depth of dormancy or the amount of chilling while recognizing that endodormancy release timing establishes the starting point from which heat accumulation drives budbreak, with earlier release enabling earlier budbreak (potentially increasing frost risk) and later release delaying budbreak (potentially providing frost protection but also delaying the growing season). The ontological significance lies in endodormancy release timing establishing the phenological calendar for the remainder of the season, where the date of release determines how much of winter and early spring heat accumulation contributes to budbreak timing versus being irrelevant because it occurs before buds become responsive, with this timing varying among cultivars (different requirements satisfied at different times) and among years (different chilling patterns reaching thresholds at different dates). The determination of endodormancy release timing involves either direct measurement through serial forcing tests (identifying when buds become highly responsive to warm temperatures) or model-based estimation (calculating when accumulated chilling reaches the requirement threshold), with the transition often gradual rather than abrupt as different buds on the vine may release at slightly different times depending on their individual exposure and sensitivity. The climate change implications for endodormancy release timing include the potential for delayed release in warming climates where chilling accumulation slows (extending the period of non-responsiveness into spring), for earlier release in climates that remain adequately cold (chilling completed earlier due to warmer autumn temperatures enabling earlier accumulation start), and for increased year-to-year variability as marginal chilling conditions create greater sensitivity to annual temperature patterns. Understanding endodormancy release timing quality enables prediction of when heat accumulation begins driving budbreak, assessment of frost risk based on the combination of release timing and subsequent temperatures, and recognition of climate change effects on the dormancy-to-growth transition that affects growing season timing.
Endodormancy_Release_Timing_Quality addresses the temporal point when chilling requirements are satisfied and ecodormancy begins. This quality determines when heat accumulation becomes relevant for budbreak prediction. The class supports phenological modeling through release timing characterization.
Dormanzbruchzeitpunkt
Endodormanzlösungszeitpunkt
Kältebedarfserfüllungsdatum
Ruhebeendigungsdatum
chilling completion date
dormancy break timing
rest release date
Caffarra A, Donnelly A (2011) The ecological significance of phenology in four different tree species: effects of light and temperature on bud burst. International Journal of Biometeorology 55(5): 711-721 Ferguson JC et al. (2014) Dynamic thermal time model of cold hardiness for dormant grapevine buds. Annals of Botany 113(5): 801-811
Endodormancy Release Timing Quality
In Riesling vines in the Mosel Valley, the endodormancy to ecodormancy shift occurs in late January to early February when accumulated chilling of approximately 1200 hours has satisfied the cultivar requirement, verified by forcing test response decreasing below 14 days and ABA concentrations declining to 70-80 ng per gram fresh weight, with buds subsequently remaining in ecodormancy until late March warming triggers budbreak initiation at BBCH stage 01.
Endodormancy to ecodormancy shift in Vitis vinifera represents the specific transition point at which the nature of dormancy control changes from internal physiological inhibition to external environmental limitation, occurring when chilling-dependent dormancy release processes have sufficiently advanced that growth competence is restored while environmental temperatures remain below the threshold required for growth initiation. This shift constitutes a discrete change in dormancy classification despite the gradual nature of the underlying physiological changes, defining the boundary between two fundamentally different dormancy states based on the source of growth inhibition. The physiological basis of the shift involves the cumulative effects of chilling exposure having reduced all dormancy-maintaining mechanisms below the threshold required to prevent growth, with ABA concentrations, DAM gene expression, and chromatin repression all declining to levels compatible with growth response to favorable conditions. The timing of the shift is determined by the interaction between cultivar-specific chilling requirements and environmental chilling accumulation, occurring earlier in warm winters with rapid chilling accumulation for low-chill cultivars and later in cold winters for high-chill cultivars, with typical timing ranging from mid-January to late February in Northern Hemisphere viticulture depending on these factors. The verification of shift completion relies on forcing tests demonstrating that buds have acquired the capacity for rapid growth response, with shifted buds breaking within 10-14 days under optimal forcing conditions compared to the extended times or failure to break characteristic of buds that remain in endodormancy. The practical significance of shift timing relates to frost risk management, with earlier shifts in warm winters creating longer ecodormancy periods during which premature warming could trigger vulnerable early growth, while later shifts in cold winters reduce this risk but may delay spring growth initiation. The molecular markers of shift completion include reduced ABA to less than approximately 80 ng per gram fresh weight, DAM gene expression declined to less than 20% of maximum endodormancy levels, and restoration of growth gene transcriptional competence verified through expression analysis of cell cycle and meristem function genes.
Endodormancy_to_Ecodormancy_Shift represents the discrete transition point within Ecodormancy_Transition marking the change in dormancy control source. Distinguished from gradual Dormancy_Depth_Reduction by representing the threshold crossing rather than continuous change. Defined by forcing response less than 14 days and molecular markers including low ABA and minimal DAM expression. Timing determined by chilling requirement satisfaction. Critical boundary in dormancy classification system.
Dormanzphasen-Transition
Ekodormanz-Eintritt
Endodormanz-Austritt
dormancy control shift
dormancy phase transition
ecodormancy entry
endodormancy exit
Lang GA et al (1987) Endo-, para-, and ecodormancy: physiological terminology and classification for dormancy research. HortScience 22: 371-377 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Zheng C et al (2015) Abscisic acid catabolism enhances dormancy release of grapevine buds. Plant Cell and Environment 38: 2490-2501
Endodormancy to Ecodormancy Shift
Cabernet Sauvignon mature seed endosperm at BBCH 89 comprises 45-50% of seed volume, contains 12-15% lipids and 18-22% proteins (dry weight basis), shows cellular structure with 20-40 μm diameter cells with thickened walls, and provides nutrient reserves supporting seedling growth for 7-10 days post-germination until photosynthetic capacity develops
An endosperm is the nutritive tissue within a Vitis vinifera seed that develops from the triple fusion of one sperm nucleus with two polar nuclei during double fertilization, providing nutrient reserves that support embryo development during embryogenesis and early seedling growth during germination. Grape seed endosperm is classified as nuclear endosperm that becomes cellular through progressive cell wall formation during mid-development. Endosperm development in Vitis vinifera proceeds through characteristic phases= initial free nuclear division without cell wall formation (0-14 days post-fertilization), cellularization through centripetal cell wall formation (14-28 days), cell expansion and accumulation of storage reserves (28-56 days), and final maturation with partial desiccation (56+ days post-fertilization corresponding to BBCH 83-89). Mature endosperm in grape seeds occupies 40-60% of total seed volume, surrounding the embryo and filling the space between embryo and seed coat. Endosperm tissue composition includes= storage lipids (10-20% dry weight, primarily triglycerides with oleic and linoleic acids), storage proteins (15-25% dry weight, primarily globulins and albumins), starch (minimal, 2-5% in Vitis compared to high-starch endosperms of cereals), cell wall polysaccharides (hemicellulose, cellulose), and various secondary metabolites. Endosperm functions in seed biology include= nutritional support for embryo during embryogenesis (providing sugars, amino acids, lipids through maternal vascular connections), storage reserves mobilized during germination (lipids and proteins are hydrolyzed and transported to growing embryo tissues), mechanical support maintaining embryo position within seed, and potential synthesis of hormones regulating seed development and dormancy. Endosperm development is critical for seed viability= inadequate endosperm development (due to fertilization failure, early abortion, or environmental stress) results in seed abortion or non-viable seeds, while complete endosperm development supports successful embryogenesis and germination capacity. In seedless grape cultivars, endosperm development failure is one mechanism underlying seed abortion= stenospermocarpic seedless varieties may show early endosperm degeneration (2-3 weeks post-fertilization) leading to embryo starvation and subsequent seed abortion. Endosperm tissue shows characteristic cellular anatomy= cells with thick cell walls rich in hemicelluloses, dense cytoplasm packed with protein bodies and lipid droplets, and progressive desiccation during final seed maturation reducing moisture content from 70-80% during development to 30-40% at seed maturity.
Endosperm emphasizes the complete nutritive tissue and is distinguished from Nutritive_Endosperm (functional support role) and Storage_Endosperm (accumulated reserve compounds) by encompassing both developmental processes and mature storage functions. Critical for understanding seed development coordination, seed abortion mechanisms in seedless cultivars, germination nutrition support, and seed storage compound composition for breeding or industrial applications.
Endosperm
Nutritive tissue
Nährgewebe
Samennährgewebe
Seed endosperm
Ebadi, A., Sedgley, M., May, P., & Coombe, B. G. (1996). Seed development and abortion in Vitis vinifera L., cv. Chardonnay. International Journal of Plant Sciences, 157(6), 703-712. | Pratt, C. (1971). Reproductive anatomy in cultivated grapes - a review. American Journal of Enology and Viticulture, 22(2), 92-109.
Merlot in Bordeaux shows endosperm development during July-October Year N+1, with triploid endosperm 3n=57 initiating free nuclear divisions immediately post-fertilization producing 24-48 nuclei in multinucleate syncytium by week 2, cellularization proceeding weeks 2-4 creating cellular endosperm, continued cell divisions and nutrient accumulation weeks 4-12 with proteins reaching 18-28% and lipids 12-22% dry weight, endosperm maturation weeks 12-16 completing development, mature endosperm surrounding 3.8-4.6 mm embryo accounting for 45-55% seed volume providing germination reserves, under oceanic conditions of 20-26°C temperature, typical of this mid-season cultivar completing endosperm development coordinating with embryo maturation
Endosperm development represents the triploid nutritive tissue formation process during seed development when triploid endosperm 3n=57 from central cell fertilization proliferates, accumulates nutrients, and matures creating tissue supporting embryo development and providing germination reserves, proceeding through free nuclear phase with rapid nuclear divisions producing multinucleate syncytium, cellular phase with cellularization creating cellular endosperm, nutrient accumulation establishing storage reserves, and endosperm maturation completing development, spanning approximately 8-14 weeks coordinating with embryo development. This nutritive tissue development process occurs during summer to early autumn in July-October in temperate regions Year N+1, representing endosperm formation parallel to embryo development. Endosperm development mechanism involves free nuclear proliferation with rapid mitotic divisions without cytokinesis creating multinucleate syncytium weeks 0-2, cellularization with walls forming around nuclei creating cellular tissue weeks 2-4, nutrient accumulation with storage compounds depositing in endosperm cells weeks 4-12, possible partial endosperm consumption by embryo though extent unclear in grapevine, and endosperm maturation completing development providing nutritive tissue. Triploid endosperm characteristics shows endosperm with 3n=57 chromosome content from sperm fusion with diploid central cell or two haploid polar nuclei, triploid genome with 2 maternal to 1 paternal genomic ratio affecting gene expression, endosperm as angiosperm-characteristic nutritive tissue, and triploidy distinguishing endosperm from diploid embryo. Central cell fertilization shows sperm from pollen tube fusing with diploid central cell creating triploid primary endosperm nucleus, fusion as second fertilization event in double fertilization, fertilization immediately initiating endosperm development, and central cell fertilization coordinating with egg fertilization. Free nuclear phase shows endosperm initiating rapid nuclear divisions without accompanying cytokinesis, nuclear divisions producing multinucleate syncytium with shared cytoplasm, typical 8-64 free nuclei forming during weeks 0-2, nuclei distributed throughout endosperm cavity, and free nuclear phase as initial proliferation mode. Nuclear division characteristics shows mitotic divisions with typical cell cycle phases, divisions initially synchronous later becoming asynchronous, division rate very rapid with cycle times possibly 6-12 hours, and rapid divisions creating numerous nuclei. Multinucleate syncytium shows multiple nuclei sharing common cytoplasm, cytoplasm filling developing seed, nuclei possibly migrating within syncytium, and syncytium as unique tissue organization. Cellularization phase shows cell walls forming around free nuclei creating cellular endosperm weeks 2-4, cellularization proceeding centripetally from periphery toward center, alveolar or cellular endosperm resulting from cellularization, and cellularization as transition to cellular tissue. Cell wall formation shows walls synthesizing between adjacent nuclei, phragmoplast-like structures possibly mediating wall formation, progressive cellularization creating multicellular tissue, and cell wall deposition completing cellularization. Cellular endosperm characteristics shows endosperm as cellular tissue after cellularization, cells containing single nucleus, cells connected through plasmodesmata possibly, and cellular organization enabling storage and transfer functions. Endosperm proliferation shows continued cell divisions after cellularization increasing endosperm cell number, divisions less rapid than free nuclear phase, eventual proliferation cessation as endosperm matures, and proliferation establishing endosperm mass. Nutrient accumulation phase shows endosperm cells accumulating storage reserves weeks 4-12, proteins accumulating as major storage compounds, lipids accumulating though less than in embryo cotyledons, carbohydrates including starch possibly accumulating, and nutrients providing embryo support and germination reserves. Protein accumulation in endosperm shows storage proteins accumulating in protein bodies, proteins providing amino acid reserves, protein content variable possibly 15-30% endosperm dry weight, and proteins supporting embryo nutrition. Lipid accumulation shows lipids in endosperm possibly 10-25% dry weight less than cotyledon levels, lipids providing energy reserves, and lipid accumulation supplementing protein storage. Carbohydrate storage shows possible starch accumulation though content variable and often low in grapevine, other carbohydrates including cell wall polysaccharides, and carbohydrates as potential reserves. Endosperm function shows endosperm supporting embryo development through nutrient transfer, endosperm providing germination reserves mobilized during seedling establishment, and endosperm as nutritive tissue essential for reproduction. Nutrient transfer to embryo shows endosperm possibly transferring nutrients to developing embryo, transfer mechanisms including plasmodesmatal connections or secretion and uptake, and transfer supporting embryo growth. Endosperm consumption shows embryo possibly consuming portion of endosperm during development, extent of consumption variable and unclear in grapevine, some mature seeds retaining substantial endosperm while others have minimal, and consumption pattern varying. Endosperm persistence shows mature seed typically retaining endosperm tissue surrounding embryo, endosperm accounting for 40-60% seed volume depending on consumption degree, persistent endosperm providing germination reserves, and persistence characteristic of grapevine seeds. Endosperm maturation shows endosperm completing development weeks 12-16, reserves accumulated to final levels, endosperm desiccating along with embryo during seed maturation, mature endosperm as storage tissue ready for germination mobilization, and maturation coordinating with embryo maturation. Desiccation shows endosperm water content declining during seed maturation, desiccation tolerance acquired, mature endosperm dry with low moisture, and desiccation enabling seed storage. Endosperm in mature seed shows endosperm as layer surrounding embryo, tissue firm and hard in dry seed, endosperm rehydrating during germination, and endosperm as persistent reserve tissue. Germination mobilization shows endosperm reserves mobilized during germination, hydrolytic enzymes degrading storage compounds, nutrients transported to growing seedling, and mobilization supporting seedling establishment. Environmental effects on endosperm development include temperature affecting development rate, nutrient availability affecting accumulation, and conditions affecting development coordinating with embryo. The process encompasses three sequential phases including endosperm proliferation with free nuclear and cellular phases, nutrient accumulation establishing storage, and endosperm maturation completing development. Duration approximately 8-14 weeks representing major endosperm development. Regional timing shows endosperm development during July-October parallel to embryo development. Viticultural significance includes representing nutritive tissue supporting embryo, providing germination reserves, demonstrating coordinated seed development, and contributing to seed quality. Research applications include analyzing endosperm developmental regulation, investigating nutrient accumulation and storage, evaluating endosperm-embryo interactions, quantifying endosperm composition and reserves, and assessing endosperm development as seed quality component.
Endosperm_Development represents nutritive tissue formation within Seed_Development. Subdivided into Endosperm_Proliferation with free nuclear (weeks 0-2, multinucleate syncytium) and cellular (weeks 2-4, cellularization) phases, Nutrient_Accumulation (weeks 4-12) with storage reserves, and Endosperm_Maturation (weeks 12-16). Triploid 3n=57. Duration ~8-14 weeks. Storage: proteins 15-30%, lipids 10-25%. Surrounds embryo 40-60% seed volume. Supports embryo development and germination. Parallel to Embryo_Development.
Endosperm-Bildung
Endosperm-Entwicklung
endosperm formation
nutritive tissue development
nutritives Gewebe-Entwicklung
storage tissue development
triploid tissue development
triploide Gewebe-Entwicklung
Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 439-442
Endosperm Development
Pinot Noir in Burgundy shows endosperm maturation during September-October Year N+1, with storage reserves completing accumulation reaching 22-27% proteins and 15-21% lipids dry weight, desiccation tolerance acquired through LEA proteins and protective sugars, metabolic activity declining toward quiescence, water content declining from 58% to 14% during weeks 12-16, mature endosperm firm and pale surrounding 3.6-4.4 mm embryo accounting for 48-56% seed volume, endosperm ready for germination mobilization providing nutrient support, under continental conditions of 18-24°C temperature, typical of this early-to-mid season cultivar completing endosperm maturation coordinating with seed maturation
Endosperm maturation represents the final endosperm development phase during endosperm development when endosperm completes storage reserve accumulation, acquires desiccation tolerance, reduces metabolic activity approaching quiescence, and desiccates along with embryo preparing for seed dispersal and storage, with endosperm achieving mature state by weeks 12-16 post-fertilization ready to support germination when seed imbibes water, completing endosperm development coordinating with embryo maturation and overall seed maturation. This final endosperm development phase occurs during early-mid autumn in September-October in temperate regions Year N+1, representing endosperm maturation completion within seed maturation. Maturation mechanism involves storage reserve accumulation completion, desiccation tolerance acquisition through protective molecule synthesis, metabolic activity decline toward dormant state, water loss during seed desiccation, structural changes preparing for desiccation and storage, and mature endosperm as stable tissue providing germination reserves. Reserve accumulation completion shows storage compounds reaching final levels weeks 12-16, protein content typically 18-28% dry weight at maturity, lipid content 12-22%, carbohydrates including structural and storage components, and completed reserves representing maximum accumulation. Final reserve composition shows balanced protein, lipid, carbohydrate composition determining nutritional value, composition varying with genotype and development conditions, reserves in stable storage forms in protein bodies and lipid bodies, and composition as seed quality characteristic. Reserve stability shows storage compounds stable during desiccation and storage, proteins and lipids in organelles protected, reserves mobilizable during germination, and stability essential for seed viability. Desiccation tolerance acquisition shows endosperm acquiring ability to survive water loss during seed maturation, late embryogenesis abundant LEA proteins possibly accumulating providing cellular protection, sugars including sucrose and raffinose oligosaccharides accumulating stabilizing cellular structures, and desiccation tolerance mechanisms enabling seed survival. Protective proteins shows LEA proteins and heat shock proteins possibly accumulating, proteins preventing cellular damage during dehydration, proteins stabilizing membranes and other proteins, and protective proteins as desiccation tolerance mechanisms. Protective sugars shows sugars replacing water in hydrogen bonding with membranes and proteins, sugars forming glasses vitrifying cytoplasm preventing crystallization damage, and sugars as important protectants during desiccation. Metabolic activity decline shows respiratory rate declining during maturation, biosynthetic activity decreasing as accumulation completes, gene expression shifting toward dormancy-associated patterns, and metabolism approaching quiescent dormant state. Respiratory decline shows oxygen consumption decreasing substantially, metabolic rate minimal in mature desiccated endosperm, quiescent metabolism enabling long-term storage, and respiration as metabolic activity indicator. Gene expression changes shows storage protein and lipid biosynthesis genes downregulating as accumulation completes, maturation and desiccation-associated genes upregulating, dormancy-related genes possibly expressed, and expression patterns reflecting maturation. Water content decline shows endosperm progressively desiccating during seed maturation weeks 12-18, water content declining from 50-70% during accumulation to 10-20% in mature dry seed, desiccation proceeding along with embryo water loss, and low moisture enabling seed storage and dormancy. Desiccation process shows water loss through seed coat and funiculus to parent plant and atmosphere, progressive dehydration over final maturation weeks, osmotic and metabolic changes accommodating water loss, and desiccation as controlled developmental process. Desiccation tolerance mechanisms shows cellular adaptations enabling survival at low moisture, membrane stabilization preventing fusion, protein protection preventing denaturation, and tolerance mechanisms essential for seed viability. Structural changes during maturation shows endosperm cells becoming compressed and dense as water lost, cell contents concentrated, protein and lipid bodies maintaining integrity, and structural changes characteristic of mature dry endosperm. Cell wall modifications shows possible cell wall changes during maturation, walls maintaining integrity during desiccation, walls affecting water relations, and wall properties important for mature tissue. Endosperm texture shows mature endosperm firm and hard in dry seed, tissue surrounding embryo as protective layer, texture affecting seed handling, and hardness from desiccation. Endosperm color shows endosperm pale to yellowish in mature seed, color from accumulated reserves and pigments, color as maturity indicator, and appearance characteristic of mature tissue. Endosperm in mature seed shows endosperm as persistent tissue surrounding embryo, endosperm accounting for 40-60% seed volume depending on embryo size and endosperm consumption, mature endosperm ready for mobilization during germination, and endosperm as stable reserve tissue. Maturation completion timing shows endosperm maturation largely completing by weeks 14-16 coordinating with embryo maturation and seed desiccation, timing varying with cultivar and environment, and completion as maturation endpoint. Mature endosperm characteristics shows tissue with completed reserves, desiccation tolerant able to survive low moisture, metabolically quiescent in dormant state, structurally stable for storage, and viable ready to support germination. Germination preparation shows mature endosperm reserves positioned for mobilization, hydrolytic enzymes synthesized or activated during germination degrading storage compounds, nutrients transported to growing seedling, and endosperm depletion during seedling establishment. Reserve mobilization shows proteins hydrolyzed to amino acids by proteases, lipids hydrolyzed to fatty acids and glycerol by lipases, fatty acids converted to sugars through β-oxidation and gluconeogenesis, carbohydrates hydrolyzed to sugars by carbohydrases, and mobilized nutrients supporting seedling growth. Endosperm function during germination shows endosperm as nutrient source for seedling until photosynthesis established, mobilization coordinated with seedling development, eventual endosperm depletion as reserves consumed, and endosperm role as temporary nutritive tissue. Environmental effects on maturation include temperature affecting maturation rate and completion timing, water availability affecting desiccation timing though process is developmentally controlled, and conditions affecting maturation coordinating with overall seed maturation. Temperature effects shows optimal range 20-26°C supporting normal maturation, temperature affecting maturation duration, and temperature as rate determinant. Maturation as seed completion shows endosperm maturation as component of overall seed maturation, coordinated completion with embryo maturation and seed coat maturation, and integrated maturation creating viable mature seed ready for dispersal. The process represents third and final phase of endosperm development completing nutritive tissue maturation, following nutrient accumulation. Duration approximately 4 weeks representing maturation completion weeks 12-16. Regional timing shows endosperm maturation during September-October completing seed development. Viticultural significance includes representing endosperm maturation completion, establishing viable nutritive tissue for germination, coordinating with embryo and seed maturation, and determining seed storage stability and germination capacity. Research applications include analyzing endosperm maturation regulation and mechanisms, investigating desiccation tolerance and protective mechanisms, evaluating metabolic decline and quiescence establishment, quantifying reserve stability and mobilization capacity, and assessing endosperm maturation as seed viability and germination vigor determinant.
Endosperm_Maturation is third and final phase of Endosperm_Development, distinguished by maturation completion. Represents weeks 12-16. Reserve completion: proteins 18-28%, lipids 12-22%. Desiccation tolerance (LEA proteins, protective sugars). Water content 50-70% → 10-20%. Metabolic quiescence. Mature endosperm 40-60% seed volume surrounding embryo. Ready for germination mobilization. Duration ~4 weeks. Follows Nutrient_Accumulation. Completes Endosperm_Development. Coordinates with Embryo_Maturation and seed maturation.
Endosperm-Dehydratation
Endosperm-Reifung
Endosperm-Vollendung
endosperm completion
endosperm desiccation
final endosperm development
finale Endosperm-Entwicklung
Endosperm Maturation
Cabernet Sauvignon in Napa Valley shows endosperm proliferation during July-early August Year N+1, with triploid endosperm 3n=57 initiating free nuclear divisions hours post-fertilization, rapid synchronous divisions with 8-12 hour cycle times producing 32-56 free nuclei in multinucleate syncytium by week 2, cellularization proceeding weeks 2-3.5 with centripetal wall formation creating cellular endosperm, continued cellular divisions weeks 3.5-5 with 16-20 hour cycle times, final endosperm cellularity supporting 45-55% seed volume, under warm climate conditions of 25-30°C temperature, typical of this late-season cultivar completing endosperm proliferation establishing tissue foundation
Endosperm proliferation represents the tissue expansion phase during endosperm development when triploid endosperm 3n=57 undergoes rapid nuclear divisions and cellularization creating multinucleate syncytium then cellular tissue, proceeding through free nuclear phase weeks 0-2 with mitotic divisions producing typically 8-64 free nuclei in shared cytoplasm, followed by cellularization phase weeks 2-4 with cell walls forming around nuclei creating cellular endosperm, and continued cell divisions increasing endosperm mass, establishing tissue foundation for subsequent nutrient accumulation. This proliferative tissue expansion phase occurs during early-mid summer in July-early August in temperate regions Year N+1, representing initial endosperm development weeks 0-4 post-fertilization. Proliferation mechanism involves free nuclear division phase with rapid mitosis without cytokinesis creating multinucleate syncytium, nuclear divisions initially synchronous becoming asynchronous, cellularization phase with wall formation around nuclei, cellular division phase with continued mitotic divisions in cellular endosperm, and proliferation establishing endosperm cellularity and mass. Free nuclear division initiation shows endosperm initiating rapid nuclear divisions immediately following central cell fertilization, first division occurring within hours to 1 day post-fertilization, divisions producing free nuclei in common cytoplasm, and free nuclear phase as initial proliferation mode. Nuclear division characteristics shows mitotic divisions with normal cell cycle phases G1, S, G2, M, divisions without accompanying cytokinesis producing free nuclei, cell cycle times very rapid possibly 6-12 hours during peak proliferation, and rapid cycles enabling exponential nuclear increase. Division synchrony shows early divisions often synchronous with nuclei dividing simultaneously, synchrony creating waves of division, later divisions becoming asynchronous with individual nuclear cycles, and synchrony pattern characteristic of free nuclear development. Multinucleate syncytium formation shows multiple free nuclei sharing common cytoplasm, typical nuclear number 8-64 by cellularization onset depending on genotype and conditions, nuclei distributed throughout endosperm cavity, cytoplasm filling developing seed, and syncytium as unique tissue organization. Nuclear number dynamics shows exponential increase from single primary endosperm nucleus to typically 16-64 free nuclei over weeks 0-2, doubling time initially 6-12 hours, eventual nuclear number depending on proliferation rate and cellularization timing, and nuclear accumulation establishing endosperm foundation. Nuclear positioning shows nuclei distributed throughout syncytium, possible nuclear migration within cytoplasm, nuclei possibly positioned near periphery in some species though pattern unclear in grapevine, and nuclear distribution affecting cellularization. Cytoplasm characteristics shows shared cytoplasm connecting all nuclei, cytoplasm with ribosomes, endoplasmic reticulum, organelles, active metabolism supporting rapid divisions, and cytoplasm as common cellular environment. Free nuclear phase duration shows phase typically lasting 1-2 weeks, timing varying with temperature and genotype, phase ending with cellularization initiation, and duration determining nuclear number at cellularization. Cellularization initiation shows cell walls beginning to form around free nuclei weeks 2-4, cellularization as transition from syncytial to cellular organization, walls separating nuclei into individual cells, and cellularization as critical developmental transition. Cell wall formation mechanism shows phragmoplast-like structures possibly forming around nuclei, cell plate synthesis creating walls, wall deposition proceeding centripetally from periphery inward in many species, progressive wall formation creating cellular tissue, and cellularization completing over days to week. Centripetal cellularization pattern shows walls forming first at endosperm periphery adjacent to seed coat, cellularization progressing inward toward center, central region last to cellularize, and centripetal pattern common in angiosperms though pattern not well-documented in grapevine. Cellular endosperm formation shows cellularization creating multicellular endosperm tissue, individual cells each containing one nucleus, cells separated by walls but possibly connected through plasmodesmata, and cellular organization replacing syncytial structure. Cell characteristics shows endosperm cells typically 20-50 micrometers diameter after cellularization, cells with dense cytoplasm and central nucleus, cells capable of continued division and growth, and cellular state enabling storage function. Cellular division phase shows continued mitotic divisions in cellular endosperm after cellularization, divisions less rapid than free nuclear phase, divisions with cytokinesis creating daughter cells, divisions increasing endosperm cell number and mass, and cellular proliferation completing tissue expansion. Division rate in cellular phase shows cell cycle times slower than free nuclear phase possibly 12-24 hours, reduced rate characteristic of cellular versus syncytial division, division rate declining as proliferation concludes, and rate reduction marking transition toward maturation. Proliferation completion shows cell divisions eventually ceasing weeks 4-6, endosperm achieving final cellularity, transition to non-dividing tissue focused on accumulation, and completion establishing mature tissue architecture. Final endosperm cellularity shows total endosperm cell number from combined free nuclear and cellular proliferation, typical endosperm mass sufficient to surround embryo occupying 40-60% seed volume, cellularity determining accumulation capacity, and final cellularity as proliferation outcome. Endosperm growth during proliferation shows endosperm expanding to fill available space in developing seed, endosperm mass increasing from cell number increase and cell growth, and growth coordinating with seed enlargement. Environmental effects on proliferation include temperature affecting division rate with warm conditions 24-28°C supporting rapid proliferation and cool conditions slowing divisions, nutrient availability supporting division biosynthetic demands, and conditions affecting proliferation rate and duration. Temperature effects shows optimal range supporting rapid free nuclear and cellular divisions, suboptimal temperatures extending proliferation phase, and temperature as major rate determinant. The process represents first phase of endosperm development establishing tissue mass, preceding nutrient accumulation and endosperm maturation. Duration approximately 4 weeks representing free nuclear and cellular proliferation weeks 0-4. Regional timing shows endosperm proliferation during July-early August as endosperm tissue forms. Viticultural significance includes representing endosperm tissue establishment, determining endosperm cellularity and mass, providing foundation for storage reserve accumulation, and establishing nutritive tissue supporting embryo. Research applications include analyzing endosperm proliferation regulation and mechanisms, investigating free nuclear division and cellularization processes, evaluating environmental effects on proliferation dynamics, quantifying nuclear and cell number accumulation, and assessing endosperm proliferation as determinant of endosperm mass and storage capacity.
Endosperm_Proliferation is first phase of Endosperm_Development, distinguished by tissue expansion. Represents free nuclear phase (weeks 0-2) with rapid divisions producing 8-64 nuclei in syncytium, followed by cellularization (weeks 2-4) creating cellular tissue, and cellular division (weeks 4-6). Cycle times 6-12h free nuclear, 12-24h cellular. Triploid 3n=57. Duration ~4 weeks. Establishes endosperm cellularity 40-60% seed volume. Precedes Nutrient_Accumulation. Temperature 24-28°C optimal.
Endosperm-Gewebe-Expansion
Endosperm-Proliferation
Endosperm-Wachstum
endosperm growth
endosperm tissue expansion
free nuclear and cellular phases
nuclear and cellular proliferation
nukleare und zelluläre Proliferation
Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Endosperm Proliferation
Cabernet Sauvignon in Napa Valley shows enlargement phase completion during mid August-early September Year N+1 at BBCH 79-81, with final cell expansion adding 3.2% volume completing 98% final size, berry growth rate declining from 48 to 8 mg per day entering lag phase, metabolic transition with respiratory rate declining from 2.2 to 1.6 μmol O₂ g⁻¹ h⁻¹, growth genes downregulated, seeds mature with brown hard coats, berry achieving 16.8-19.4 mm diameter and 12.4-18.6 g, under warm climate conditions of 24-30°C temperature, typical of this late-season cultivar completing enlargement
Enlargement phase completion represents the growth phase termination period during berry cell enlargement when final minimal cell expansion concludes, pre-lag phase transition establishes metabolic reorganization, growth rate decelerates to near-zero entering lag phase quiescence, and berry completes size establishment achieving 95-100% final dimensions ready for ripening transformation, occurring weeks 7-10 post-fertilization as enlargement phase ends and lag phase begins. This phase termination period occurs during late summer to early autumn in mid August-early September in temperate regions Year N+1, spanning approximately 1-2 weeks representing enlargement conclusion, corresponding to BBCH 79-81 majority berries touching to beginning of ripening stages. Completion mechanism involves final cell expansion producing last 2-8% size increase, pre-lag metabolic transition with transcriptional reprogramming from growth to lag phase metabolism, growth deceleration to very low rates <10-20 mg per day establishing quiescence, seed maturation completion enabling berry developmental progression, and phase completion enabling transition to lag phase and subsequent veraison. Final cell expansion characteristics show continued minimal cell enlargement adding last 2-8% cell volume, cell expansion rate very low completing final dimensions, cells reaching 98-100% final mature size, and final expansion establishing ultimate cell dimensions. Cell volume finalization shows cells achieving final mature volumes, typical dimensions 80-200 micrometers representing complete 30-100 fold expansion from post-division size, vacuoles occupying 88-93% final cell volume, and finalized cell architecture persisting through ripening. Cell expansion cessation shows expansion rate declining to near-zero, eventual complete growth cessation in most cells though minor expansion possible during ripening, cells in mature expanded state, and cessation marking end of cell enlargement phase. Pre-lag phase metabolic transition shows metabolic reorganization preparing lag phase characteristics, respiratory rate declining from growth phase elevated levels toward lag phase lower metabolism, transcriptional reprogramming downregulating growth genes and upregulating lag phase-associated genes, enzyme activities shifting from growth-supporting to maintenance metabolism, and metabolic transition as key completion process. Respiratory changes show oxygen consumption declining from peak 2-4 μmol O₂ g⁻¹ h⁻¹ to declining 1.5-2.5 μmol O₂ g⁻¹ h⁻¹ during completion, reduced respiration reflecting declining metabolic activity, transition toward lag phase lower respiration typical 1.0-1.8 μmol O₂ g⁻¹ h⁻¹, and respiratory decline marking metabolic transition. Gene expression reprogramming shows growth-related genes including expansins, cell cycle regulators downregulated to basal levels, lag phase-associated genes including stress response, secondary metabolism genes upregulated, metabolic enzyme gene expression shifting toward maintenance and preparation for ripening, and transcriptional changes establishing lag phase program. Protein expression changes show growth-related proteins declining, lag phase-specific proteins accumulating, enzyme complements shifting toward lag metabolism, and protein changes implementing metabolic transition. Growth deceleration characteristics show berry fresh weight gain declining to very low rates <10-20 mg per day, eventual near-zero growth establishing lag phase quiescence typical <5 mg per day, growth effectively ceasing entering lag phase, and deceleration marking phase transition. Berry growth rate trajectory shows decline from 30-80 mg per day during late maximum enlargement to 10-30 mg per day early completion to <10 mg per day late completion entering lag, progressive deceleration to quiescence, and rate trajectory characteristic of growth cessation. Berry size at completion shows berry achieving 95-100% final mature dimensions, typical ranges 6-10 mm small-berried, 10-15 mm medium-berried, 15-23 mm large-berried cultivars at completion, fresh weight 2-6 g small, 5-13 g medium, 10-28 g large cultivars, and completion size strongly predicting harvest size. Berry composition at completion shows water content 78-83% fresh weight, organic acids declining from peak 1.0-1.4%, sugars beginning accumulation 8-12%, pH rising slightly 2.9-3.1 from acid decline, and composition transitioning toward lag phase characteristics. Organic acid dynamics show peak acids 1.2-1.6% during late enlargement declining to 1.0-1.4% by completion, malate declining through respiration, tartrate relatively stable, and acid decline as metabolic shift indicator. Sugar dynamics show sugars increasing from 6-10% during late enlargement to 8-12% by completion, limited sugar accumulation during enlargement phase, beginning sugar import through phloem, and sugar increase initiating during completion. Berry morphology at completion shows berry at near-final size, green color persisting though chlorophyll beginning decline, texture softening slightly from peak firmness, and appearance as mature-sized green berry approaching lag phase. Berry surface characteristics show smooth skin maintained, cuticle development continuing, epidermis intact, and surface properties at maturity for growth phase. Seed maturation during completion shows seeds completing development by completion phase, seed color change from green to brown indicating maturation, seed coat lignification and hardening, embryo and endosperm development complete, and seed maturation prerequisite for berry progression. Seed characteristics at completion shows fully developed seeds, brown colored coat, hard texture, viable embryo, and mature seed ready for dispersal if separated. Cluster architecture at completion shows berries at maximum compactness, bunch closure complete, cluster at mature pre-ripening configuration, and architecture affecting disease risk and ripening access. Environmental effects on completion include temperature affecting completion timing with warm conditions accelerating transition to lag, cool conditions extending completion period, water availability less critical during completion than earlier expansion, and environmental factors having reduced impact as developmental programming dominates completion. Temperature effects show warm conditions promoting faster metabolic transition and earlier lag phase entry, cool conditions delaying completion extending enlargement, and temperature affecting transition timing. Completion timing shows typical occurrence weeks 7-10 post-fertilization, cultivar variation in timing with early cultivars completing earlier week 7-8 and late cultivars week 9-10, and timing variation affecting total development duration. Completion indicators show growth rate declining to <10 mg per day, berry size at 95-100% final dimensions, metabolic markers showing lag phase characteristics, seed maturation complete with brown hard seeds, and indicators documenting completion. The process shows three distinguishable aspects including final cell expansion producing last volume increase, pre-lag phase transition establishing metabolic reorganization, and growth deceleration to quiescence. Duration typically 1-2 weeks representing completion period weeks 7-10. Regional timing shows enlargement phase completion during mid August-early September across climates as growth phase ends. Viticultural significance includes representing definitive growth phase termination, marking transition to lag phase and progression toward ripening, establishing final berry size determining yield, and completing berry structural development preparing for ripening transformation. Research applications include analyzing completion mechanisms and regulation, investigating metabolic transition and transcriptional reprogramming, evaluating seed maturation as completion prerequisite, quantifying completion timing and indicators, and assessing enlargement completion as determinant of subsequent lag and ripening phases.
Enlargement_Phase_Completion represents growth phase termination within Berry_Cell_Enlargement_Phase. Subdivided into Final_Cell_Expansion producing last 2-8% increase, Pre_Lag_Phase_Transition with metabolic reorganization, and Growth_Deceleration to quiescence <10 mg/day. Duration 1-2 weeks representing weeks 7-10. Berry achieves 95-100% final size. Metabolic and transcriptional reprogramming. Seed maturation complete. Completes Berry_Cell_Enlargement_Phase. Enables transition to Lag_Phase. Critical developmental checkpoint.
Expansionsphase-Ende
Vergrößerungs-Abschluss
Vergrößerungsphase-Vollendung
Wachstumsphase-Terminierung
enlargement conclusion
expansion phase end
growth phase termination
pre-lag completion
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 451-455 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Enlargement Phase Completion
Pinot Noir ENTAV-INRA clone 777 bears an ENTAV_INRA_Clone_Role, selected in the Côte d'Or during the 1970s-1980s, evaluated through replicated trials across Burgundy and other regions, registered in the Catalogue after demonstrating consistent quality character (aromatic intensity, early ripening), and distributed globally through the ENTAV licensing network. Merlot clone 181 serves in an ENTAV_INRA_Clone_Role as the most widely planted Merlot clone in Bordeaux, its ENTAV registration documenting balanced performance suitable for diverse site conditions and wine styles. Syrah clone 470 bears an ENTAV_INRA_Clone_Role as a recent registration (1990s) selected for northern Rhône production, with documented resistance to coulure and performance data supporting adoption for quality Hermitage and Côte-Rôtie production.
A French clone role that inheres specifically in clones selected, evaluated, and registered through the ENTAV-INRA partnership program, representing the official French national clone selection system established in 1962 and producing the authoritative Catalogue des Variétés et Clones de Vigne. ENTAV-INRA clones undergo standardized evaluation protocols including 10-20 year field trials at multiple sites across French viticultural regions, standardized winemaking and sensory evaluation, comprehensive virus testing and elimination where necessary, and administrative registration conferring official status. The ENTAV-INRA designation identifies clones within the formal French certification hierarchy (Pre-Basic, Basic, Certified/Standard categories), distinguishing them from informal selections, foreign imports, or material outside the certification system. ENTAV-INRA maintains mother plant collections at the Domaine de l'Espiguette (Mediterranean coast) and regional stations, supporting certified propagation through licensed French nurseries and international distribution through material transfer agreements. The ENTAV-INRA catalog includes over 400 scion variety clones and 40+ rootstock clones, representing the world's most comprehensive single-institution clone program.
ENTAV_INRA_Clone_Role specifies the official institutional pathway for French clone registration, distinguishing these selections from historic French material predating ENTAV, from selections by other French institutions (universities, regional stations), and from foreign clones naturalized in France. The ENTAV-INRA partnership divided responsibilities—ENTAV managing practical selection, mother plant maintenance, and nursery relations while INRA contributed research expertise, virus detection methodology, and scientific evaluation protocols. This partnership created the systematic approach that became the model for clone programs worldwide. ENTAV-INRA clones carry institutional validation that market participants increasingly demand the "Dijon clones" designation specifically refers to ENTAV-INRA selections made from Burgundy source material, not to all Pinot Noir from Dijon.
Catalogue-registered clone
ENTAV-INRA clone
ENTAV-INRA-Klon
French certified clone
Katalog-Klon
official French clone
offizieller französischer Klon
ENTAV-INRA (2007) Catalogue des Variétés et Clones de Vigne Cultivés en France Audeguin L et al. (1999) 30 years of clonal selection in France, ISHS Acta Horticulturae 473: 69-76 Boidron R (1995) Clonal selection in France, in Bentley R (ed) The Handbook of Grape Processing, Blackwell Walter B, Martelli GP (1996) Clonal and sanitary selection of the grapevine, INRA Editions
ENTAV INRA Clone Role
The Central Asian table grape cultivar Nimrang approaches entire margin condition more closely than most Vitis vinifera varieties, displaying only very shallow, broadly rounded marginal undulations that lack the distinct tooth points characteristic of European wine grapes this reduced marginal complexity, combined with large berry size and thick skin, reflects adaptation to the continental climate of the Fergana Valley where the variety originated, and distinguishes Nimrang from European varieties in ampelographic assessments of Central Asian germplasm collections.
Entire_Margin_Quality represents the margin configuration where the leaf edge is smooth and continuous without teeth, serrations, or other indentations along its length. An entire margin forms a smooth curve or line from one point to another without the projecting or receding features that characterize toothed margins. Among Vitis vinifera cultivars, truly entire margins are rare because the vast majority of grape varieties possess some degree of marginal serration. However, the degree of serration varies substantially, and some cultivars approach entire margins with only very shallow, indistinct teeth that may be described as subentire or obsoletely toothed. Entire margins are more common in some Vitis species outside the vinifera complex, particularly certain North American species, and can appear in interspecific hybrids. Within V. vinifera, the closest approaches to entire margins occur in certain table grape varieties and in some local varieties from arid regions where reduced marginal complexity may relate to water conservation. The entire margin condition results from developmental patterns where marginal meristematic activity produces uniform expansion without the localized growth peaks that generate teeth. Assessment of entire versus toothed margins involves examining the blade edge under magnification if necessary to detect subtle teeth, and may employ categorical scales ranging from entire through crenate (scalloped) to toothed. The functional implications of entire versus toothed margins remain debated, with hypotheses involving gas exchange, water relations, and herbivore deterrence.
Entire_Margin_Quality represents the limiting case of Margin_Shape_Quality where teeth are absent or indistinct. True entire margins meeting strict botanical definition are rare in Vitis vinifera the term may be applied relatively to indicate much-reduced serration compared to typical cultivars. Assessment should distinguish between genuinely entire margins and margins where teeth are present but small or worn away through abrasion or senescence. OIV descriptor 077 does not include an entire category, implicitly acknowledging that most V. vinifera cultivars possess teeth comparison to interspecific hybrids or wild Vitis species provides context for margin type variation. PATO_0002012 (entire) provides the upper-ontology reference for this margin type.
continuous margin
edentate margin
ganzrandige Qualität
glatter Rand
smooth margin
ungezähnter Rand
untoothed edge
zahnloser Blattrand
Galet P (1979) A Practical Ampelography, Cornell University Press pp.: 89-94 Hickey LJ (1973) Classification of the Architecture of Dicotyledonous Leaves. American Journal of Botany 60(1): 17-33
Entire Margin Quality
Airen vines in La Mancha bearing an entry-level wine role are cultivated in low-density bush-trained plantings at 1,600 vines/ha on arid meseta soils without irrigation, mechanically harvested at yields of 8-12 t/ha, with fruit purchased by large cooperatives at 0.15-0.25 €/kg for wines retailing at 1.50-3.00 €/bottle in Spanish supermarkets under private-label or cooperative brands, representing the accessible baseline of Spanish wine consumption at over 150 million liters annual production from La Mancha alone.
A quality tier role that inheres in a grapevine whose fruit is designated for the lowest-priced branded wines within a producer's portfolio or market segment, positioned at the threshold between branded commercial wine and unbranded bulk wine, offering basic varietal or regional identity at the most accessible price point that still carries a consumer-facing label.
Entry_Level_Wine_Role describes the price-quality position immediately above bulk wine, distinguished from the bulk tier by the presence of branded packaging, consumer-facing labeling, and retail shelf placement. The boundary between entry-level and commercial wine is fluid and market-dependent: in high-cost markets such as Scandinavia or the UK, entry-level branded wines may retail at $5-8 USD equivalent, while in producing countries such as Spain, Italy, or Argentina the same tier may begin below $2 USD. Viticultural practices at this tier are typically extensive, low-input, and yield-maximizing within the constraints of appellation regulations where applicable.
Basiswein-Rolle
Einstiegspreislage-Rolle
Einstiegswein-Rolle
entry-level wine vine role
opening price point role
value wine role
Anderson K and Nelgen S (2020) Which Winegrape Varieties are Grown Where? 2nd Ed, University of Adelaide Press OIV (2023) State of the World Vine and Wine Sector Rabobank (2019) Wine Quarterly Report: Global Wine Market Segmentation
Entry Level Wine Role
In Pinot Noir vines of the Willamette Valley, Oregon, environment dependent quiescence maintains bud inactivity from early February through mid-March, with quiescence entirely dependent on temperatures remaining below 10°C as demonstrated by immediate growth initiation when single-node cuttings are transferred to 22°C forcing conditions, achieving visible bud swelling within 5 days and full budbreak within 10 days, illustrating the absence of internal barriers to growth.
Environment dependent quiescence in Vitis vinifera represents the state of bud inactivity during ecodormancy that is entirely determined by external environmental conditions, particularly temperature, with the quiescent state being maintained only as long as environmental factors remain unfavorable for growth and transitioning immediately to active development when favorable conditions occur. This quiescence differs fundamentally from the dormancy of endodormancy in being externally imposed rather than internally generated, with no physiological barriers to growth existing within the bud itself and quiescence depending entirely on the environmental context. The temperature-dependence of quiescence reflects the fundamental requirement for adequate warmth to support the metabolic processes of growth, with temperatures below approximately 10°C being insufficient to sustain the enzyme activities, biosynthetic reactions, and cellular processes required for cell division and expansion regardless of the growth competence of bud tissues. The immediate responsiveness of environment dependent quiescence to changing conditions is demonstrated by forcing experiments, where transfer of ecodormant buds to warm conditions immediately initiates growth processes without the delay characteristic of endodormant material, and by field observations showing partial growth responses during warm spells followed by return to quiescence when temperatures decline. The ecological function of environment dependent quiescence is to synchronize growth resumption with reliably favorable conditions, with temperature serving as an indicator of seasonal progression that correlates with reduced frost risk and adequate conditions for sustained growth, though this synchronization is imperfect and vulnerable to unseasonable weather patterns. The metabolic state during quiescence involves minimal but not absent activity, with some respiratory metabolism continuing at rates determined by temperature, gradual accumulation of growth-related transcripts and proteins in preparation for growth, and maintenance of cellular structures in a state ready for rapid activation when conditions permit. The transition from quiescence to active growth occurs when environmental conditions become and remain favorable long enough to trigger irreversible commitment to budbreak, typically requiring accumulation of 100-300 growing degree days above the base temperature threshold.
Environment_Dependent_Quiescence represents the externally-determined inactive state within Conditional_Dormancy. Distinguished from internally-determined endodormancy by complete dependence on environmental factors. Demonstrated by immediate forcing response without delay. Temperature is the primary determining factor with base threshold approximately 10°C. Metabolic activity continues at low levels during quiescence. Transition to growth requires sustained favorable conditions and heat unit accumulation.
environment-controlled inactivity
environmentally-imposed quiescence
exogene Ruhe
exogenous quiescence
extern bedingte Quieszenz
external condition dependent dormancy
umweltabhängige Ruhe
umweltkontrollierte Inaktivität
Lang GA et al (1987) Endo-, para-, and ecodormancy: physiological terminology and classification for dormancy research. HortScience 22: 371-377 Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Lavee S, May P (1997) Dormancy of grapevine buds - facts and speculation. Australian Journal of Grape and Wine Research 3: 31-46
Environment Dependent Quiescence
In Cabernet Sauvignon vines in the Médoc, Bordeaux, environmental constraint maintains bud quiescence from early February when chilling requirements are satisfied through late March when spring warming releases constraint, with mean February temperatures of 7-9°C remaining below the 10°C base temperature threshold and preventing budbreak despite fully restored growth competence, until sustained March warming provides sufficient growing degree day accumulation for growth initiation.
Environmental constraint in Vitis vinifera represents the external environmental factors that prevent growth initiation in ecodormant buds despite their restored growth competence, encompassing primarily temperature limitation and secondarily moisture limitation that maintain bud quiescence during late winter and early spring until conditions become favorable for growth. This constraint operates from the completion of dormancy release through budbreak initiation, representing the interval during which growth-competent buds await environmental conditions permissive for development. The temperature component of environmental constraint is quantified through the concept of base temperature below which metabolic and developmental processes cannot proceed at rates sufficient for growth, typically 10°C for grapevine, with temperatures below this threshold effectively preventing growth regardless of bud competence. The molecular basis of temperature constraint involves the temperature-dependence of enzyme activities required for biosynthesis and energy metabolism, the requirement for adequate ATP production to support anabolic processes, and the temperature sensitivity of hormone signaling pathways that transduce environmental signals into developmental responses. The practical effect of temperature constraint is to delay budbreak until accumulated warmth has satisfied the heat requirement for growth initiation, with this delay providing protection against frost damage by preventing premature growth during the period when late frosts remain likely. The intensity of environmental constraint varies with ambient conditions, with continuous cold maintaining strong constraint while warming periods reduce constraint and may trigger partial growth responses if sustained, creating the characteristic fluctuating bud development often observed during transitional spring weather. The moisture constraint component relates to the requirement for adequate soil water availability to support the cellular expansion and transpiration associated with active growth, though moisture limitation during early spring is uncommon in most viticultural regions where winter precipitation has recharged soil water reserves. The release of environmental constraint occurs progressively as spring warming provides increasing heat unit accumulation, eventually crossing the threshold for budbreak initiation when accumulated growing degree days satisfy the cultivar-specific heat requirement.
Environmental_Constraint represents the external limitation phase within Ecodormancy addressing factors that prevent growth in competent buds. Distinguished from internal inhibition characteristic of endodormancy. Subdivides into Temperature_Limitation, Moisture_Limitation, and Conditional_Growth_Suppression as component factors. Primary constraint is temperature below approximately 10°C base. Duration from dormancy release through budbreak initiation. Progressive release through spring warming and heat accumulation.
Umweltbeschränkung
climatic constraint
environmental growth inhibition
environmental limitation
external growth constraint
externe Wachstumsbeschränkung
klimatische Beschränkung
umweltbedingte Wachstumshemmung
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Parker AK et al (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17: 206-216
Environmental Constraint
In Gewürztraminer vines in Alsace, environmental control onset occurs in early February following chilling requirement satisfaction, after which budbreak timing becomes predictable from temperature accumulation models, with growing degree day accumulation above 10°C base temperature tracked from this point and budbreak occurring when approximately 150-180 GDD have accumulated, typically in late March to early April depending on spring temperature patterns.
Environmental control onset in Vitis vinifera represents the establishment of external environmental factors as the primary determinants of growth status following the release of internal dormancy constraints, marking the beginning of the period during which bud behavior is governed by temperature, moisture, and other environmental variables rather than by intrinsic physiological inhibition. This transition occurs coincident with the endodormancy to ecodormancy shift, as the dismantling of internal growth inhibition transfers control of bud quiescence from endogenous regulatory mechanisms to exogenous environmental conditions. The practical consequence of environmental control onset is that bud growth status becomes predictable from environmental monitoring, with temperature-based models able to forecast budbreak timing based on heat unit accumulation, in contrast to the endodormancy period when internal factors override environmental influences and make growth unpredictable from environmental data alone. The temperature component of environmental control involves the requirement for heat unit accumulation above a base temperature threshold, typically 10°C for grapevine, with budbreak initiation requiring cultivar-specific accumulations typically ranging from 100-300 growing degree days following the onset of environmental control. The responsiveness to environmental signals during this period reflects the restored sensitivity of growth-regulatory pathways to external cues, with temperature affecting the activity of growth-promoting hormones, the rate of metabolic processes supporting growth, and the expression of genes required for cell division and expansion in meristematic tissues. The moisture component of environmental control relates to the requirement for adequate water availability to support the cellular expansion and metabolic activity associated with growth resumption, though moisture limitation is typically less important than temperature limitation in most viticultural environments during early spring. The onset of environmental control represents a vulnerable period for the vine, as growth-competent buds may respond prematurely to temporary warm conditions, initiating growth that is subsequently damaged by frost when temperatures return to winter levels, highlighting the ecological risk associated with the transition from robust internal control to more responsive environmental control.
Environmental_Control_Onset represents the regulatory transition aspect of Ecodormancy_Transition marking the shift to external growth control. Distinguished from internal control characteristic of endodormancy. Enables temperature-based budbreak prediction through heat unit models. Primary controlling factor is temperature with secondary moisture effects. Creates vulnerability to premature growth from early warming. Coincides with chilling requirement satisfaction timing.
Umweltfaktoren-Dominanz
Umweltkontroll-Beginn
environmental growth control
environmental regulation onset
exogene Kontroll-Etablierung
exogenous control establishment
external factor dominance
externe Wachstumskontrolle
Environmental Control Onset
Environmental synchronization assessment comparing traditional and warming climate scenarios for Cabernet Sauvignon reveals the disruption of historical phenology-climate alignment, with traditional climate showing well-synchronized development (budbreak after last frost, flowering during optimal temperature window, ripening completing before autumn rains), while warming scenario shows degraded synchronization (budbreak advancing into frost-risk period from accumulated heat despite unchanged last-frost date, flowering shifted into heat-stress temperatures, ripening compressed into hottest period before historical autumn cooling), demonstrating how climate change disrupts the environmental synchronization that historical cultivar-region matching had optimized.
Environmental synchronization quality is a synchronization quality that represents the degree of alignment between grapevine phenological development and the environmental signals that regulate developmental timing, characterizing how effectively vine developmental programs respond to and integrate photoperiod, temperature, and other environmental cues to coordinate growth phases with seasonal climate patterns that determine the success of each phenological stage. Within the BFO framework, this quality addresses the environment-development interface dimension of synchronization, distinguishing the alignment between external signals and internal developmental responses from synchrony among organs or plants while recognizing that environmental synchronization determines whether vine development is appropriately phased with the seasonal opportunities and constraints that affect productivity and fruit quality. The ontological significance lies in environmental synchronization determining the adaptive fit between vine phenology and climate, where well-synchronized vines initiate growth when frost risk has passed, flower when conditions favor fruit set, ripen when temperatures promote quality development, and enter dormancy before winter cold threatens survival, while poorly synchronized vines may experience frost damage from premature budbreak, heat stress during flowering, or incomplete ripening before autumn deterioration. The environmental signals coordinating vine development include photoperiod (day length affecting dormancy induction and potentially other responses), temperature (both as threshold triggers and as cumulative heat sums driving development rate), and their interaction with endogenous developmental programs that determine the cultivar-specific phenological patterns characterizing different grape varieties. The climate change implications for environmental synchronization are substantial, as warming temperatures alter the thermal environment relative to the unchanged photoperiod signal, potentially disrupting the historical coordination between temperature-driven and photoperiod-driven developmental processes, with consequences for dormancy, budbreak timing, and the overall alignment of phenology with seasonal patterns. The cultivar adaptation dimension of environmental synchronization reflects the selection history that has matched cultivar thermal requirements and photoperiod sensitivities to the climate patterns of traditional growing regions, with this historical alignment potentially disrupted by climate change or by planting cultivars outside their regions of adaptation. Understanding environmental synchronization quality enables assessment of cultivar-climate fit, prediction of climate change impacts on phenological alignment, and recognition of the environmental factors coordinating vine development with seasonal patterns.
Environmental_Synchronization_Quality serves as the parent class for specific environmental signal alignments (Photoperiod, Temperature Sum). This quality addresses the coordination between vine development and environmental cues determining adaptive fit. The class supports climate adaptation assessment through environmental synchronization characterization.
Klima-Phänologie-Abstimmung
Umweltkoordination
Umweltsynchronisation
climate adaptation fit
climate-phenology alignment
environmental coordination
saisonale Synchronisation
seasonal synchronization
Environmental Synchronization Quality
Enzyme thermostability quality investigation in Shiraz characterized the molecular basis of metabolic heat sensitivity during late-season ripening when heat events most frequently affect this cultivar. In vitro thermostability assessment of key enzymes extracted from berry tissue identified differential heat sensitivities relevant to fruit composition. Vacuolar invertase, essential for sugar accumulation, showed good enzyme thermostability quality with 50% activity retained after 10 minutes at 50°C and only gradual inactivation at temperatures below 45°C, suggesting sugar accumulation capacity is relatively heat-stable. Malate dehydrogenase, crucial for organic acid metabolism, exhibited moderate enzyme thermostability quality with substantial activity loss above 42°C, potentially contributing to the accelerated malate respiration and elevated pH observed in heat-affected fruit. Phenylalanine ammonia-lyase, the entry point to phenolic biosynthesis, showed poor enzyme thermostability quality with significant inactivation above 38°C, providing a molecular explanation for the reduced anthocyanin accumulation characteristic of berries exposed to high temperatures during véraison. The enzyme thermostability quality hierarchy established that phenolic synthesis limitation represents the most heat-sensitive metabolic process affecting Shiraz fruit quality, followed by acid metabolism, with sugar accumulation relatively robust, explaining the compositional signature of heat-affected fruit showing normal Brix but elevated pH and reduced color.
Enzyme thermostability quality is a thermal tolerance quality that inheres in a grapevine cultivar or clone and represents the capacity of metabolic enzymes to maintain catalytic activity and structural integrity under elevated temperature conditions, with thermostability varying among enzymes according to their intrinsic structural properties and among genotypes reflecting genetic variation in protein sequences and the expression of protective mechanisms that maintain enzyme function at high temperatures. This quality addresses the metabolic foundation for continued physiological function under heat stress, recognizing that enzyme inactivation or denaturation disrupts the metabolic pathways supporting all aspects of vine growth, development, and stress response, making enzyme thermostability a fundamental determinant of overall heat tolerance. The ontological significance of enzyme thermostability quality lies in its role as the molecular basis for temperature sensitivity of metabolism, with different enzymes showing characteristic inactivation temperatures that create a hierarchy of metabolic vulnerabilities determining which processes fail first as temperatures rise and which pathways continue functioning to support stress survival and recovery. The biological basis of enzyme thermostability encompasses the inherent thermal stability of protein structure determined by amino acid sequence and post-translational modifications, the protection conferred by molecular chaperones including heat shock proteins that prevent aggregation and assist refolding of partially denatured enzymes, and the stabilizing effects of compatible solutes and other factors that modulate the thermal denaturation behavior of proteins in the cellular environment. Key enzymes limiting heat tolerance in grapevine include Rubisco activase controlling photosynthetic carbon fixation, membrane-associated ATPases maintaining ionic gradients, and enzymes of sugar and organic acid metabolism affecting fruit composition, with the thermostability of these and other critical enzymes collectively determining the metabolic heat tolerance of the whole plant. Assessment of enzyme thermostability quality employs in vitro activity measurements across temperature ranges, thermal inactivation kinetics, and differential scanning calorimetry to characterize denaturation temperatures, providing detailed mechanistic understanding of heat tolerance limitations.
Enzyme_Thermostability_Quality is_a Thermal_Tolerance_Quality, representing the heat resistance of metabolic enzymes as a molecular basis for metabolic function under stress. This class addresses the protein-level basis of heat sensitivity. Temperature for 50% inactivation (T₅₀) provides standard quantification for individual enzymes. The hierarchy of enzyme sensitivities explains differential metabolic impacts of heat. Rubisco activase, invertase, malate dehydrogenase, and PAL represent key enzymes with different stabilities affecting photosynthesis and fruit composition. Heat shock proteins play critical protective role. Distinguished from Membrane_Thermostability_Quality by focus on soluble enzymes rather than membrane systems.
Enzym-Hitzestabilität
Enzym-Thermostabilität
Protein-Thermostabilität
enzyme heat stability
metabolic heat tolerance
metabolische Hitzetoleranz
protein thermal stability
protein thermostability
Salvucci ME, Crafts-Brandner SJ (2004) Inhibition of photosynthesis by heat stress: the activation state of Rubisco as a limiting factor in photosynthesis. Physiologia Plantarum 120:179-186 Wahid A et al. (2007) Heat tolerance in plants: an overview. Environmental and Experimental Botany 61:199-223 Greer DH, Weedon MM (2013) The impact of high temperatures on Vitis vinifera cv. Semillon grapevine performance and berry ripening. Frontiers in Plant Science 4:491
Enzyme Thermostability Quality
Epicatechin content quality analysis in Cabernet Sauvignon seeds employs phloroglucinolysis to cleave proanthocyanidin polymers and release epicatechin extension units, revealing mean degree of polymerization of 8-12 units (epicatechin/catechin molar ratio of 7-11) that indicates moderate polymer length associated with balanced astringency comparison with skin tannins (mDP 25-40, epicatechin/catechin ratio 24-39) demonstrates the structural differences between seed and skin tannin pools that winemakers exploit through differential extraction timing to craft wines with optimal tannin structure.
Epicatechin_Content_Quality represents the chemical quality characterizing the concentration of (−)-epicatechin in grape berry tissues, capturing the flavan-3-ol stereoisomer with 2,3-cis configuration that serves as the primary extension unit in condensed tannin polymers. (−)-Epicatechin (2R,3R-flavan-3-ol) differs from (+)-catechin in the cis relationship between C2 and C3 substituents, a stereochemical difference that profoundly affects its role in tannin structure. Epicatechin functions predominantly as an extension unit in proanthocyanidins, contributing the upper positions in polymer chains where its C4 carbon links to C6 or C8 of the next unit. The proportion of epicatechin relative to catechin in condensed tannins increases with polymer chain length, as each internal position adds an epicatechin unit while only the terminal position contains catechin. In grape seeds, epicatechin and its 3-O-gallate ester together typically comprise 40-60% of total flavan-3-ols, with the gallate ester contributing enhanced astringency and bitterness. Epicatechin is perceived as slightly more bitter and astringent than catechin at equivalent concentrations, with differences attributed to the stereochemical effects on receptor binding. The mean degree of polymerization of seed tannins, calculable from the epicatechin-to-catechin ratio after depolymerization, typically ranges from 5-15 units for grape seeds. Skin tannins show higher mean degree of polymerization (10-80 units) with corresponding elevated epicatechin proportion. Epicatechin content and its ratio to catechin provide structural information about condensed tannins relevant to understanding astringency potential and mouthfeel evolution.
Epicatechin_Content_Quality addresses the 2,3-cis stereoisomer that dominates the extension unit positions in proanthocyanidin polymers. The analytical distinction between free epicatechin monomers and epicatechin released by polymer cleavage is important phloroglucinolysis or thiolysis enables quantification of extension unit epicatechin separate from terminal unit catechin. The epicatechin-3-gallate ester, while derived from epicatechin, contributes distinct sensory properties and is sometimes quantified separately. The predominance of epicatechin as extension unit versus catechin as terminal unit reflects the reaction mechanism of proanthocyanidin biosynthesis and polymerization.
(−)-Epicatechin-Konzentration
(−)-epicatechin content
2,3-cis-Flavan-3-ol-Gehalt
2,3-cis-flavan-3-ol concentration
Epicatechingehalt
Extensionseinheitengehalt
epicatechin monomer quality
extension unit content
Kennedy JA and Jones GP (2001) Analysis of Proanthocyanidin Cleavage Products Following Acid-Catalysis in the Presence of Excess Phloroglucinol. Journal of Agricultural and Food Chemistry 49(4): 1740-1746 Downey MO et al. (2003) Analysis of Tannins in Seeds and Skins of Shiraz Grapes Throughout Berry Development. Australian Journal of Grape and Wine Research 9(1): 15-27
Epicatechin Content Quality
Cabernet Sauvignon epidermal layer at maturity comprises single cell layer 24-32 μm radial height with cells 35-55 μm tangential width arranged in irregular polygonal mosaic pattern, outer periclinal walls 4-5.5 μm thick heavily cutinized and suberized creating hydrophobic barrier continuous with overlying 4.5-6.5 μm cuticle, bearing approximately 95 stomata per mm² (mostly non-functional at maturity, guard cells collapsed) and 12 lenticels per mm², containing moderate flavonol levels (0.6-1.2 mg/g in epidermal cells, higher 1.2-2.2 mg/g in sun-exposed berries) providing UV protection. Chardonnay white cultivar epidermal layer showing white wine characteristics= cells 18-28 μm radial height, 28-52 μm tangential width, outer walls 2.5-4 μm thick (thinner than typical red cultivars), higher stomatal density 130-180 per mm² than Cabernet Sauvignon, containing flavonols and carotenoids in epidermal cells creating golden-yellow coloration contribution, with thinner walls and higher stomatal density potentially contributing to white cultivar susceptibility to Botrytis requiring careful disease management. Syrah epidermal layer ultrastructure examined by transmission electron microscopy showing outer wall fine structure= outer cuticle-wall interface region with cutin penetrating 0.8-1.5 μm into outer wall creating cutin-impregnated zone middle outer wall region 2-3.5 μm with suberin lamellae (alternating light and dark bands in TEM) creating suberin-impregnated zone inner outer wall region 0.5-1 μm transitioning to cellulose-hemicellulose framework total outer wall 4-6 μm creating substantial barrier through multiple hydrophobic layer system. Pinot Noir epidermal layer sun exposure effects= sun-exposed berries showing epidermal cells with higher flavonol content (1.5-2.5 mg/g quercetin derivatives in epidermal layer versus 0.5-1 mg/g in shaded), demonstrating UV-B induction of flavonol biosynthesis in epidermal cells providing enhanced UV protection in exposed berries, with epidermal flavonols contributing to wine phenolic profile particularly in skin-contact white wines or lightly extracted red wines where epidermal extraction occurs. Merlot epidermal layer stomatal development and function timeline= stomata forming during weeks 2-4 post-fruit set (BBCH 71-73) as epidermal cells differentiate functional stomata present during green berry phase (weeks 4-9, BBCH 73-79) with stomatal conductance 80-150 mmol H₂O m⁻² s⁻¹ supporting berry transpiration and photosynthesis stomatal closure beginning at veraison (week 10, BBCH 81) as guard cells begin losing turgor non-functional stomata at maturity (week 18, BBCH 89) with guard cells collapsed and pores closed or partially open but non-responsive to environmental signals, demonstrating temporal pattern of epidermal layer stomatal function supporting early development but not ripening.
An epidermal layer is the outermost cellular layer of the berry comprising a single layer of specialized epidermal cells covered externally by cuticle, typically 15-35 μm radial thickness and 20-60 μm tangential cell width at maturity, serving as primary cellular boundary between berry internal tissues and external environment, providing structural support for cuticle attachment, bearing specialized surface structures (stomata, lenticels, trichome scars), and contributing to berry protection through thick modified outer cell walls and barrier functions. Epidermal layer cellular characteristics= (1) single continuous cell layer completely covering berry surface, with cells arranged in irregular polygonal pattern creating mosaic appearance when viewed from surface (2) small cell dimensions radially (15-35 μm height) relative to underlying pericarp cells (80-180 μm in middle pericarp), with tangential dimensions (20-60 μm width, 25-80 μm length) creating roughly isodiametric to slightly elongated cells (3) thick heavily modified outer periclinal walls (wall facing exterior) typically 3-6 μm thick (versus 0.5-1.5 μm for inner periclinal and anticlinal walls), with outer wall heavily cutinized (cutin impregnation) and suberized (suberin impregnation) creating hydrophobic barrier integrated with overlying cuticle (4) thin inner periclinal walls (0.5-1.5 μm) interfacing with underlying hypodermis (5) anticlinal walls (lateral walls between adjacent epidermal cells) moderately thick (1-2 μm) creating cellular boundaries (6) reduced vacuolar space (vacuole occupying 40-60% of cell volume versus 90-95% in middle pericarp cells) with denser cytoplasm containing nucleus, plastids, mitochondria supporting metabolic activity. Epidermal layer specialized structures= (1) stomata (pores for gas exchange) consisting of pairs of guard cells surrounding stomatal pore, typically 50-200 stomata per mm² berry surface with higher density on stylar (blossom) end versus pedicel (stem) end, functional primarily during green berry development providing CO₂ for photosynthesis and transpiration pathway, largely non-functional during ripening as guard cells lose turgor and stomata remain closed or partially closed (2) lenticels (raised multicellular structures) typically 5-20 per mm² providing gas exchange particularly during ripening when stomatal function declines, appearing as small protrusions 30-80 μm diameter on berry surface (3) trichome scars (remnants of hair-like structures) occasionally present particularly near stylar end, representing trichomes shed during berry expansion (4) anticlinal wall patterns sometimes showing specific cultivar-characteristic configurations visible in epidermal peels. Epidermal layer composition and properties= (1) cutin content high in outer wall creating cutin-impregnated wall layer continuous with overlying cuticle proper (2) suberin content in outer wall providing additional hydrophobic barrier (3) cellulose and hemicellulose in wall framework providing structural strength (4) phenolic compounds sometimes present in epidermal cells including flavonols (quercetin glycosides particularly in sun-exposed berries) and occasionally anthocyanins though anthocyanins primarily concentrated in underlying hypodermis (5) low sugar and acid content relative to pericarp bulk as epidermal cells not primary storage tissue. Epidermal layer functions= (1) structural support for cuticle attachment providing cellular foundation for acellular cuticle layer (2) contribution to berry barrier through thick hydrophobic outer walls creating cellular component of protective barrier complementing acellular cuticle (3) mechanical strength through thick walls and cell arrangement providing resistance to compression and abrasion (4) gas exchange regulation through stomatal and lenticel structures (5) defense responses through ability to synthesize antimicrobial compounds (phytoalexins, phenolics) in response to pathogen attack or wounding.
Epidermal_Layer represents the outermost cellular layer of the berry and is the cellular component of the exocarp/outer pericarp, distinguished from the acellular Cuticle_Layer covering it and from the underlying Hypodermal_Layer. Sub-classification into Outer_Epidermal (emphasizing surface properties) and Cell_Wall_Thickened_Epidermal (emphasizing barrier properties) reflects different aspects of epidermal structure and function. Understanding epidermal layer structure is essential for comprehending berry barrier properties, cuticle-cell integration, stomatal function, and cellular contributions to berry protection and quality.
Epidermis-Schicht
Oberflächenzellschicht
epidermal cell layer
epidermale Zellschicht
surface cell layer
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 176-180 Harris, J.M. et al. (1968). The Vinifera grape berry= anatomical study. Botanical Gazette 129=1-23 Rosenquist, J.K., Morrison, J.C. (1989). Some factors affecting cuticle and wax accumulation. American Journal of Enology and Viticulture 40=241-244.
Epidermal Layer
Cabernet Sauvignon epidermal outer pericarp at maturity comprises cuticle (4.5-6 μm thick with epicuticular wax crystalloids 0.8-1.5 μm height creating bloom, total wax content 140-190 μg/cm²) and epidermis (single cell layer 24-32 μm radial height, cells 35-55 μm tangential width, outer wall 4-5.5 μm thick heavily cutinized), bearing approximately 95 stomata per mm² (mostly closed/non-functional at maturity) and 12 lenticels per mm², with total epidermal outer pericarp thickness 28-38 μm providing water barrier limiting transpiration to 0.8-1.5 g H₂O m⁻² h⁻¹ at 25°C and 50% RH. Chardonnay white cultivar epidermal outer pericarp showing cultivar characteristics= thinner cuticle (2.5-4 μm) with lower wax content (80-120 μg/cm²) than typical red cultivars, epidermis (18-28 μm height) with moderate outer wall thickness (2.5-4 μm), higher stomatal density (130-180 per mm²) than Cabernet Sauvignon, creating epidermal outer pericarp 20-32 μm thickness with higher water permeability (1.2-2 g H₂O m⁻² h⁻¹) than thicker red cultivar epidermis. Syrah epidermal outer pericarp wax analysis by GC-MS showing wax composition= alkanes (C25-C33, primarily C29 and C31) comprising 42% of total wax, primary alcohols (C24-C32, primarily C26 and C28) 28%, fatty acid esters 16%, other components 14%, with total epicuticular wax 165 μg/cm² providing characteristic waxy bloom appearance and hydrophobic surface (water contact angle 95-110°). Pinot Noir epidermal outer pericarp light exposure effects= sun-exposed berries developing thicker cuticle (5-7 μm versus 3-4.5 μm shaded), higher wax content (180-220 versus 90-130 μg/cm²), and higher flavonol concentration in epidermal cells (quercetin glycosides 1.5-2.5 mg/g versus 0.5-1 mg/g), demonstrating sunlight induction of protective compounds and barrier enhancement in epidermal outer pericarp. Merlot epidermal outer pericarp stomatal development and function= stomata forming during weeks 2-4 post-fruit set reaching density 110-140 per mm² at BBCH 73-75, functional during green berry phase (weeks 4-8) with stomatal conductance 80-150 mmol H₂O m⁻² s⁻¹ supporting berry transpiration and photosynthesis, declining function during veraison (weeks 9-10) as guard cells lose turgor and stomata progressively close, non-functional at maturity (weeks 16-18) with stomatal conductance <5 mmol m⁻² s⁻¹, demonstrating temporal stomatal activity pattern supporting early development but not ripening berry function.
An epidermal outer pericarp is the outermost portion of the outer pericarp comprising the epidermis and cuticle layers (with or without hypodermis depending on classification), representing the absolute surface of the berry in direct contact with external environment, typically 15-45 μm total thickness (epidermis plus cuticle), serving as primary protective barrier against water loss, UV radiation, pathogen invasion, and mechanical damage while bearing specialized structures (stomata, lenticels, trichome scars) important for gas exchange and berry surface characteristics. Epidermal outer pericarp comprises= (1) cuticle (outermost acellular layer) 2-8 μm thick at maturity, composed of cutin (cross-linked polyester polymer of C16 and C18 hydroxy fatty acids), waxes (aliphatic compounds including alkanes, alcohols, aldehydes, ketones, esters), and epicuticular wax crystalloids projecting from surface creating bloom (waxy appearance) cuticle functions include water barrier reducing transpiration (permeability 10⁻⁸ to 10⁻⁹ m s⁻¹), pathogen barrier resisting penetration, UV protection through light scattering, and hydrophobic surface reducing water film formation (2) epidermis (single outermost cell layer) consisting of small tightly-packed cells typically 15-35 μm radial thickness and 20-60 μm tangential width, with thick outer periclinal walls (3-6 μm thick, heavily cutinized and suberized) providing structural support and barrier function, thin inner periclinal walls (0.5-1.5 μm) interfacing with underlying tissues, and lateral anticlinal walls creating cellular boundaries. Epidermal cells show specialized characteristics= small cell size relative to pericarp bulk (15-35 μm height versus 80-180 μm in middle pericarp cells) thick heavily modified outer walls with cuticle integration dense cytoplasm with limited vacuolar space (vacuole occupying 40-60% of cell volume versus 90-95% in middle pericarp) moderate-to-high phenolic content including flavonols and sometimes anthocyanins. Epidermal outer pericarp bears specialized structures= stomata (pores for gas exchange) typically 50-200 per mm² berry surface with higher density on stylar (blossom) end versus pedicel (stem) end, consisting of two guard cells surrounding stomatal pore, functional primarily during green berry development (Phase I) providing CO₂ for photosynthesis and transpiration for cooling, largely non-functional during ripening as guard cells lose turgor and stomata remain closed or partially closed lenticels (raised multicellular structures) 5-20 per mm² providing gas exchange particularly during ripening when stomata less functional, appearing as small protrusions 30-80 μm diameter on berry surface trichome scars (remnants of hair-like structures) occasionally present particularly near stylar end, representing trichomes that were present during early development but shed during berry expansion. Epidermal outer pericarp cuticle composition shows cultivar and environmental variation= total wax content 50-250 μg/cm² berry surface with sun-exposed berries showing higher wax content than shaded berries wax composition including alkanes (30-50% of total wax), primary alcohols (20-35%), esters (10-25%), and other components cutin=wax ratio affecting barrier properties with higher cutin providing stronger barrier. Epidermal outer pericarp development follows berry development= cuticle deposition beginning during Phase I and continuing through ripening with thickness increasing from 1-2 μm at fruit set to 2-8 μm at maturity epidermal cells undergoing limited expansion (15-25% diameter increase) compared to middle pericarp cells (200-300% increase) creating progressive thinning of epidermis relative to expanding berry stomata forming during early development (BBCH 71-73) and becoming less functional during ripening (BBCH 81-89).
Epidermal_Outer_Pericarp represents the absolute outermost pericarp layer and is distinguished from Surface_Outer_Pericarp which may include hypodermal layers beneath epidermis. Epidermal outer pericarp provides the primary protective barrier and bears specialized surface structures. Understanding epidermal outer pericarp structure and function is important for comprehending water relations (transpiration, splitting susceptibility), pathogen resistance (barrier to infection), and quality protection (UV screening, mechanical protection). The cuticle component is particularly important for postharvest quality and storage.
Beerenoberfläche
Epidermis
berry surface
cuticle layer
epidermales äußeres Perikarp
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 176-180 Rogiers, S.Y. et al. (2004). Grape berry cv. Shiraz epicuticular wax and transpiration. American Journal of Enology and Viticulture 55=121-127 Rosenquist, J.K., Morrison, J.C. (1989). Some factors affecting cuticle and wax accumulation. American Journal of Enology and Viticulture 40=241-244.
Epidermal Outer Pericarp
Epidermal turgor assessment in Riesling grape berries reveals values of 0.35±0.06 MPa at véraison providing the skin tautness resisting crack initiation, declining to 0.12±0.04 MPa at harvest maturity when the reduced turgor combined with continued berry expansion creates the stretch conditions predisposing to rain-induced splitting, with the threshold turgor of 0.15 MPa identified as the critical value below which splitting risk increases exponentially during precipitation events.
Epidermal turgor quality is a cell turgor quality that inheres in the outermost cell layer of grapevine organs and represents the hydrostatic pressure within epidermal cells that contributes to surface mechanical integrity, provides the counter-pressure against which guard cells operate for stomatal function, and determines the resistance of plant surfaces to physical damage and pathogen penetration. Within the BFO framework, this quality specifically characterizes the turgor of the tissue layer forming the interface between plant and environment, distinguishing epidermal cell behavior from underlying parenchyma and the specialized guard cells embedded within the epidermis. The ontological significance lies in epidermal turgor maintaining surface tautness that affects cuticular wax orientation, stomatal mechanics, and the physical barrier function of the plant surface against water loss and biotic invasion. Epidermal turgor in grapevine leaves typically ranges from 0.3-0.6 MPa under well-watered conditions, declining with water stress but generally showing smaller fluctuations than guard cell turgor due to the absence of active osmotic regulation mechanisms present in guard cells. The interaction between epidermal and guard cell turgor determines stomatal mechanics, where the aperture reflects the pressure differential rather than absolute values, and epidermal backpressure influences the sensitivity of stomatal response to guard cell turgor changes. In grape berries, epidermal turgor contributes to skin mechanical properties affecting texture perception, splitting resistance, and the force required for berry detachment, with declining turgor during ripening increasing susceptibility to physical damage and Botrytis infection. Measurement of epidermal turgor requires pressure probe access to individual epidermal cells or may be inferred from tissue-level measurements combined with modeling of cellular contributions. Understanding epidermal turgor quality enables interpretation of surface mechanical properties, prediction of damage susceptibility, and analysis of stomatal mechanics in the context of the epidermal pressure environment.
Epidermal_Turgor_Quality addresses the surface cell layer contributing to boundary properties and stomatal mechanics. This quality interacts with Guard_Cell_Turgor_Quality in determining stomatal aperture through the mechanical model where aperture = f(Pguard - Pepidermal). In berry context, epidermal turgor relates to but is not identical with Skin_Firmness_Quality which integrates turgor with cell wall mechanical properties.
Epidermisturgor
Hautzellenturgor
Oberflächenzellturgor
dermal cell pressure
epidermaler Zellturgor
outer layer turgor
skin cell turgor
surface cell turgor
Franks PJ et al. (1998) A study of stomatal mechanics using the cell pressure probe. Plant, Cell & Environment 21(1): 94-100 Becker T, Knoche M (2012) Deposition, strain, and microcracking of the cuticle in developing 'Riesling' grape berries. Vitis 51(1): 1-6
Epidermal Turgor Quality
An equal trifurcated tendril at node 8 on a highly vigorous shoot of Vitis vinifera cv. Syrah in the northern Rhône Valley, France, exhibiting a true three-way branch point at 6 cm from shoot attachment, dividing into three arms of 10.5 cm, 11 cm, and 11.5 cm length (pairwise ratios 0.95=1, 0.96=1, and 0.91=1), with branch angles of 115, 120, and 125 degrees creating nearly perfect radial symmetry, all three branches showing 2.9-3.1 mm diameter at mid-length indicating balanced vascular development, all exhibiting comparable circumnutation amplitudes of 140-155 degrees in complementary spatial zones, all demonstrating rapid coiling response (8-12 minutes) when experimentally contacted, occurring on a shoot with exceptional vigor (total length 185 cm) in optimally fertilized and irrigated conditions supporting robust three-way resource partitioning, measured at BBCH stage 71 (fruit set) in mid-June, representing the rare achievement of truly balanced trifurcation that maximizes the architectural potential of three-arm design.
A trifurcated tendril exhibiting balanced development across all three branches in Vitis vinifera, characterized by arm length ratios among all pairwise comparisons greater than 0.85=1 indicating nearly equal resource allocation and developmental timing across the three arms, branch angles approximating 120 degrees separation creating geometrically balanced radial architecture, equal or nearly equal diameter across all three branches reflecting similar vascular development and structural investment, and comparable functional capabilities in all arms including similar circumnutation amplitude, coiling response vigor, and load-bearing capacity. Equal trifurcated tendrils develop when three lateral meristems at a single branch point, or meristems at sequential branch points in pseudo-trifurcation, are activated simultaneously or in rapid succession, receive balanced resource allocation throughout development without preferential flows favoring particular arms, and experience uniform environmental conditions without differential stress creating developmental asymmetries. This branching pattern represents optimal morphological expression of trifurcated architecture, maximizing functional benefits of triple searching arms through equivalent exploration capacity in all branches, providing maximum redundancy in attachment opportunities with any combination of arms capable of establishing functional support, and creating mechanically balanced structures that can distribute loads optimally across multiple attachment points when two or three arms achieve contact. Equal trifurcation is relatively rare compared to unequal patterns, typically representing 20-35% of trifurcated tendrils in most cultivars, and requires high vigor and optimal resource availability to support balanced three-way development without resource limitation creating dominance hierarchies.
Equal trifurcated tendrils are distinguished from unequal trifurcated tendrils by arm length ratios >0.85=1 across all pairwise comparisons, balanced branch angles approximating 120-degree separation, and equivalent structural and functional development across all three arms. Achievement of equal trifurcation requires substantially higher resource availability than equal bifurcation due to three-way rather than two-way resource partitioning, making this pattern sensitive to vigor variation and stress conditions. Equal trifurcation frequency serves as an indicator of vine resource status, with high frequencies (>30% of trifurcated tendrils) suggesting excellent vigor and resource availability while low frequencies (<15%) indicate moderate resource limitation. Some cultivars show genetic tendencies toward equal trifurcation when resources permit, while others rarely achieve balanced three-arm development even under optimal conditions reflecting different developmental regulation strategies. Functionally, equal trifurcation provides maximum exploration efficiency and attachment redundancy among trifurcated architectures.
ausgewogene dreifach-verzweigte Ranke
balanced trifurcated tendril
even tri-arm tendril
gleichmäßig dreigabelige Ranke
symmetrical three-branch tendril
symmetrische dreiarmige Kletterorgan
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 136-150 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 173-194.
Equal Trifurcated Tendril
Pinot Noir hermaphrodite flowers at BBCH 63-65 (active bloom, 30-60% flowers open) exhibit five erect filaments maintaining vertical orientation (5-15° from true vertical), with filaments 3-3.6 mm positioning anthers 0.8-1.4 mm above pistil stigma (pistil 2-2.3 mm), creating optimal self-pollination geometry with pollen falling from elevated dehiscing anthers directly onto receptive stigma below, achieving 48-56% fruit set through efficient autonomous self-pollination. Wild V. vinifera ssp. sylvestris staminate male flowers show particularly robust erect filaments (3.8-4.5 mm) maintaining strictly vertical orientation (0-10° from vertical), elevating large anthers (0.8-1 mm) to 4.6-5.5 mm above receptacle and 4-5 mm above vestigial pistil, maximizing pollen release height for wind dispersal to distant pistillate females. Cabernet Sauvignon under strong wind conditions (15-25 km/h) during bloom shows erect filaments experiencing transient displacement to 20-40° from vertical during wind gusts but recovering to 5-15° vertical orientation between gusts, maintaining adequate anther elevation for self-pollination despite mechanical perturbation and achieving 50-58% fruit set comparable to calm-weather blooms.
An erect filament is a filament that maintains upward vertical or near-vertical orientation (within 0-30° from vertical) following the rapid elongation phase at anthesis, positioning the anther at or above the level of the pistil stigma, characteristic of hermaphrodite and staminate flowers where erect positioning facilitates self-pollination (hermaphrodites) or maximizes pollen dispersal height (staminates), representing the ancestral and functionally optimal orientation for male reproductive function in self-compatible reproductive systems. Erect filaments are characterized by= (1) vertical or near-vertical orientation with filament axis aligned 0-30° from vertical, achieved through inherent growth directionality and tissue structural properties maintaining upright positioning (2) anther positioned at or above stigma level, creating spatial geometry where pollen released from elevated anthers can fall or be blown downward onto lower stigmas (self-pollination in hermaphrodites) or be dispersed into air currents for cross-pollination (3) straight or slightly curved filament morphology without the pronounced downward curvature characteristic of reflex filaments (4) maintenance of erect positioning throughout anthesis and post-anthesis periods (BBCH 61-69) until senescence causes general structural collapse. Erect filaments in hermaphrodite flowers position anthers typically 0.5-1.5 mm above the pistil stigma (with 2.5-4 mm filaments supporting 0.6-0.9 mm anthers above 1.8-2.4 mm pistils), creating optimal geometry for autonomous self-pollination= pollen released from anthers falls under gravity or is carried by slight air movements downward to contact the stigma surface positioned directly below, with the elevated anther position also allowing pollen dispersal to adjacent flowers for local cross-pollination. Erect filaments in staminate flowers achieve greater anther elevation (typically 3-4.5 mm above receptacle base with 3-4.5 mm filaments supporting 0.7-1.1 mm anthers well above vestigial <0.5 mm pistils), maximizing pollen release height for wind-borne cross-pollination with pollen entering air currents and being transported 20-200 m to receptive pistillate flowers. The developmental basis of erect filament orientation involves= (1) inherent growth polarity directing cell expansion in apical direction during rapid elongation (2) structural organization of cortex parenchyma cells and vascular tissues creating mechanical rigidity maintaining upright positioning (3) absence of the differential growth patterns or hormonal influences that create the pronounced curvature in reflex filaments. Erect filament maintenance requires adequate structural rigidity to resist gravitational forces and wind pressure that might cause drooping or displacement, with rigidity provided by= cell wall reinforcement in cortex cells turgor pressure maintaining cell and tissue firmness vascular strand providing internal structural support. Environmental factors can influence erect filament positioning= wind forces can cause transient displacement but filaments generally recover erect positioning when wind subsides water stress reducing turgor can cause some drooping though severe stress affecting flower function generally structural damage from hail or mechanical injury can disrupt erect positioning. The functional significance of erect filament orientation is fundamental for hermaphrodite flower reproductive strategy, as the erect positioning creates the essential spatial arrangement enabling efficient self-pollination (pollen source elevated above pollen destination), with erect positioning distinguishing self-fertile hermaphrodite flowers from cross-pollination-requiring functionally female flowers (reflex filaments positioning pollen source below pollen destination preventing self-pollination). Erect filaments represent the dominant filament orientation in cultivated V. vinifera as hermaphrodite flowers with erect stamens were strongly selected during domestication for reliable self-fertility enabling consistent fruit production, contrasting with the ancestral wild dioecious condition where only staminate males exhibit erect filaments (pistillate females having vestigial/rudimentary stamens of variable orientation).
Erect_Filament represents the upright orientation characteristic of hermaphrodite and staminate flowers, distinguished from reflex filaments (strongly curved downward >60-120° from vertical in functionally female/pistillate flowers) by vertical positioning and from horizontal or variable orientations in vestigial/rudimentary stamens. The erect orientation is functionally essential for self-pollination capability in hermaphrodite flowers by positioning pollen source above pollen destination, and is the defining morphological feature distinguishing hermaphrodite from functionally female flowers where reflex orientation prevents self-pollination. Erect filament maintenance represents a stable developmental and structural outcome of normal stamen development in self-compatible flower types, contrasting with the specific developmental modifications creating reflex curvature in functionally female types. Understanding erect versus reflex filament determination is central to Vitis sex determination biology and reproductive strategy evolution.
aufrechter Staubfaden
gerader Staubblattfaden
upright stamen stalk
vertical filament
vertikales Filament
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 99-106 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 93-101 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644.
Erect Filament
Mechanistic investigation of Erineum_Formation_Resistance_Disposition at INRAE Montpellier compared cellular responses to Colomerus vitis feeding in susceptible V. vinifera 'Carignan' versus resistant V. rupestris accessions using microscopy and gene expression analysis. Leaf pieces were infested with standardized mite numbers, and cellular responses were monitored at 24, 48, 72, and 168 hours post-infestation. On susceptible 'Carignan', mite feeding induced visible trichome proliferation at 48 hours, with recognizable erineum structures by 72 hours and fully developed felt-like galls by 168 hours. Trichome length increased 8-fold (from 50μm to 400μm) within developing erinea, and expression of trichome-associated genes (MYB transcription factors, structural genes) was strongly upregulated. On resistant V. rupestris, initial mite probing occurred normally, but by 24 hours, cells at and adjacent to feeding sites showed browning and collapse characteristic of hypersensitive response. No trichome proliferation was induced, and by 72 hours, mites had died or emigrated from the necrotic areas. Gene expression analysis revealed rapid upregulation of defense-associated genes (PR-1, chitinase, phenylalanine ammonia-lyase) in V. rupestris but not in 'Carignan' following mite feeding. These findings demonstrate that Erineum_Formation_Resistance_Disposition in V. rupestris involves active recognition of mite feeding and deployment of defense responses that sacrifice attacked cells rather than supporting gall development, mechanistically parallel to effector-triggered immunity against pathogens.
An Erineum_Formation_Resistance_Disposition is an Eriophyid_Mite_Resistance_Disposition that inheres in a Vitis plant leaf and is realized through defense processes that prevent or limit the induction and development of erinea—the characteristic felt-like galls formed on leaf surfaces in response to feeding by the erineum strain of the grape erineum mite (Colomerus vitis). This disposition represents resistance to a specific host-parasite interaction in which mite feeding on the abaxial (lower) leaf epidermis triggers abnormal proliferation of epidermal trichomes, creating convex blisters visible on the adaxial (upper) surface with corresponding concave, felt-lined pockets on the lower surface where mites feed and reproduce within the protective trichome mass. Erineum development requires coordinated plant responses to mite feeding including localized suppression of normal developmental programs, induction of trichome initiation and hyper-elongation, and maintenance of the modified tissue as a favorable mite habitat. Erineum_Formation_Resistance_Disposition disrupts this sequence through alternative host responses including hypersensitive cell death at feeding sites preventing the coordinated tissue response, failure of epidermal cells to respond to mite-secreted effectors or growth regulators, constitutive trichome density that may physically impede mite access to preferred feeding sites, or rapid wound healing responses that abort incipient gall development. Substantial variation in erineum susceptibility exists among V. vinifera cultivars and wild Vitis species, with some genotypes essentially immune to erineum formation, providing genetic resources for understanding resistance mechanisms and potential breeding applications where erineum is economically or aesthetically problematic.
Erineum_Formation_Resistance_Disposition is_a Eriophyid_Mite_Resistance_Disposition. Erinea are leaf galls specifically induced by the erineum strain of Colomerus vitis through manipulation of plant trichome development. Resistance operates by preventing the coordinated plant response required for gall formation, analogous to Gall_Formation_Resistance_Disposition against foliar phylloxera. Erineum damage is primarily cosmetic in commercial viticulture but may be problematic for fresh grape sales or vineyard tourism. Wild Vitis species, particularly V. rupestris and V. amurensis, provide potential resistance sources.
Colomerus vitis gall resistance
Erineum-Bildungs-Resistenz-Disposition
Filzgallen-Resistenz
Pockenmilben-Gallen-Resistenz
erineum resistance
leaf gall mite resistance
Duso C, de Lillo E (1996) Grape, Eriophyoid Mites: Their Biology, Natural Enemies and Control, Elsevier Westphal E et al. (1990) Histological observations on grapevine leaves showing the erineum strain of Colomerus vitis (Acari: Eriophyidae), Entomologia Generalis 15:261-270 Smith LM, Stafford EM (1948) The bud mite and the erineum mite of grapes, Hilgardia 18:317-334
Erineum Formation Resistance Disposition
Systematic screening for Eriophyid_Mite_Resistance_Disposition at Hochschule Geisenheim University evaluated Colomerus vitis erineum strain colonization across 50 cultivars using controlled mite transfer and natural field infestation. Detached leaf assays involved placing mite-infested leaf pieces on test cultivar leaves and scoring erineum development at 14 and 28 days post-transfer on a 0-5 scale based on gall coverage. Field assessment scored natural erineum development at mid-season. V. vinifera 'Silvaner' showed extreme susceptibility with rapid erineum development (score 4.5 at 14 days, 5.0 at 28 days in detached leaves 4.2 field score) covering over 60% of adaxial leaf surface with corresponding felt-like galls on abaxial surfaces. 'Riesling' demonstrated moderate Eriophyid_Mite_Resistance_Disposition (detached: 2.2 at 14 days, 2.8 at 28 days field: 1.8), with erineum development delayed and limited in extent. Remarkably, certain V. amurensis accessions exhibited apparent immunity with no erineum development (scores 0.0) despite successful mite transfer, suggesting complete resistance to gall induction. Microscopic examination revealed that attempted feeding on resistant V. amurensis leaves triggered rapid browning of contacted cells within 24 hours, preventing the coordinated epidermal proliferation required for erineum formation. This hypersensitive-like response parallels mechanisms observed in pathogen resistance, suggesting that Eriophyid_Mite_Resistance_Disposition may involve active recognition and defense rather than passive incompatibility.
An Eriophyid_Mite_Resistance_Disposition is a Mite_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that limit colonization, feeding, gall induction, or population development by eriophyid mites (Eriophyoidea), microscopic, worm-shaped mites that are highly specialized parasites of grapevine causing distinctive erineum galls on leaves and damaging buds. This disposition represents resistance to the grape erineum mite (Colomerus vitis), which in its erineum strain induces characteristic felt-like galls (erinea) on leaf undersides, and in its bud mite strain causes stunted, distorted shoot growth, as well as the grape rust mite (Calepitrimerus vitis), which damages buds and young foliage. Eriophyid mites are among the smallest arthropods (100-300 μm), possess only two pairs of legs, and feed by penetrating individual epidermal cells with their stylet-like mouthparts, either triggering abnormal cell proliferation forming galls or causing cell death and tissue scarring. Resistance mechanisms may include leaf anatomical features (stomatal density, trichome presence, epidermal cell properties) affecting mite access and feeding success, biochemical factors preventing or limiting gall induction, rapid cell death responses sacrificing attacked cells before gall development proceeds, and overall plant vigor enabling tolerance to mite damage. Eriophyid mite damage is generally considered of limited economic importance in commercial viticulture except where severe bud mite infestation reduces fruitfulness, but can be cosmetically problematic in direct-market operations and may serve as reservoirs for virus transmission in some systems.
Eriophyid_Mite_Resistance_Disposition is_a Mite_Resistance_Disposition. Eriophyid mites include both gall-inducing (erineum strain of C. vitis) and non-gall-inducing (bud mite strain of C. vitis, rust mite C. vitis) forms, with different damage symptoms requiring separate assessment. Child classes include Erineum_Formation_Resistance_Disposition (resistance to leaf galling) and Bud_Mite_Resistance_Disposition (resistance to bud damage). Eriophyid mites are highly host-specific, with C. vitis found only on Vitis, limiting their ecological impact but ensuring consistent pest pressure in vineyards.
Eriophyiden-Resistenz
Eriophyoidea resistance
Gallmilben-Resistenz-Disposition
Pockenmilben-Abwehr
gall mite resistance
rust mite resistance
Duso C, de Lillo E (1996) Grape, Eriophyoid Mites: Their Biology, Natural Enemies and Control, Elsevier Smith LM, Stafford EM (1948) The bud mite and the erineum mite of grapes, Hilgardia 18:317-334 Walton VM et al. (2007) Relationship between rust mites Calepitrimerus vitis and bud damage on selected grape cultivars, Vitis 46:66-70
Eriophyid Mite Resistance Disposition
Touriga Nacional vines on the steep terraced socalcos of the Douro Valley bearing an erosion control role anchor schist soils on 30-50° gradients through root penetration into fractured bedrock at 3-6 m depth, with traditional stone terrace walls reducing slope length to 3-8 m limiting rill formation, vine canopy intercepting 20-30% of rainfall energy during summer convective storms, and managed inter-row cover of Medicago polymorpha reducing surface runoff velocity by 60-80% compared to bare tilled soil, collectively reducing measured erosion rates from 30-45 t/ha/year (bare soil) to 3-8 t/ha/year under integrated vine-terrace-cover crop management.
A soil conservation role that inheres in a grapevine whose root system, canopy cover, and associated ground management practices actively prevent or reduce the detachment and transport of soil particles by water, wind, or tillage forces, maintaining soil depth and structural integrity on the vineyard site.
Erosion_Control_Role addresses the preventive dimension of soil conservation — maintaining existing soil resources against degradation. Vineyard erosion is driven by the interaction of slope gradient, rainfall intensity, soil erodibility, and management practices: the vine's contribution to erosion control is mediated through multiple mechanisms including root reinforcement of soil shear strength (increasing soil cohesion by 5-20 kPa), canopy interception reducing raindrop kinetic energy, and organic matter inputs improving soil aggregate stability. The role is amplified when combined with structural measures (terraces, contour planting) and vegetative measures (cover crops, mulching).
Bodenverlust-Schutz-Rolle
Erosionskontroll-Rolle
Erosionsverhütungs-Rolle
anti-erosion role
erosion prevention role
soil loss control role
Rodrigo-Comino J (2018) Five decades of soil erosion research in Mediterranean vineyards, Geomorphology 307: 204-217 Prosdocimi M et al. (2016) Soil water erosion on Mediterranean vineyards, Catena 141: 1-21 Novara A et al. (2019) Cover crop impact on soil loss in a vineyard, Catena 172: 145-153
Erosion Control Role
The name Sauvignonasse was erroneously applied to Sauvignon Blanc plantings in Chile for decades until DNA analysis in the 1990s revealed that approximately 70% of Chilean vineyards labeled Sauvignon Blanc were actually Sauvignonasse (VIVC prime name Sauvignon Vert, synonym Friulano/Tocai Friulano), a genetically distinct cultivar — similarly, Malbec plantings in Argentina long included material misidentified as Malbec that was subsequently revealed through SSR profiling to be genetically distinct, and Petite Sirah in California was shown to comprise a mixture of true Durif and Peloursin, both erroneously designated under a single synonym.
A synonym that was incorrectly applied to a grapevine cultivar through misidentification, taxonomic confusion, or propagation error, creating a false nomenclatural link between a name properly belonging to one cultivar and a genetically distinct cultivar to which it was mistakenly assigned.
Erroneous_Synonym documents nomenclatural errors that propagated through the viticultural system before molecular identification tools became available. These errors arose through multiple mechanisms including visual misidentification of morphologically similar cultivars, confusion of seedlings with their parents, accidental mixing of propagation material in nurseries, and perpetuation of historical misattributions in ampelographic literature: some erroneous synonyms persisted for over a century before molecular resolution. The documentation of erroneous synonymy is critical for correcting historical records, understanding the true genetic composition of old vineyard plantings, and preventing recurring misidentification. Erroneous synonyms are flagged distinctly from valid synonyms in the VIVC and OIV databases.
Fehlbezeichnung
erroneous identification name
false synonym
fehlerhafte Sortenbezeichnung
irrtümliches Synonym
misapplied name
This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Meredith CP (2003) Science as a window into wine history, Proceedings of the 3rd International Symposium on Molecular Markers pp: 71-78 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157
Erroneous Synonym
Comparative analysis of Esca_Resistance_Disposition at the University of Florence employed controlled trunk inoculation of 15 cultivars with Phaeomoniella chlamydospora to characterize early resistance responses. Six months post-inoculation, wood cores were extracted and analyzed for extent of vascular discoloration, pathogen DNA quantity (qPCR), and host defense gene expression. 'Sangiovese' (tolerant) exhibited 65% shorter wood streaking (mean 4.2 cm versus 12.1 cm in susceptible 'Trebbiano'), 78% lower Pch DNA concentration, and 3.5-fold higher expression of genes encoding stilbene synthase, phenylalanine ammonia-lyase, and chitinase in tissues adjacent to inoculation sites. Histological examination revealed that 'Sangiovese' produced tyloses in 45% of vessels within 5mm of infection versus only 12% in 'Trebbiano', and accumulated dense phenolic deposits visible as dark reaction zones compartmentalizing the infection. Phytotoxin sensitivity assays showed that 'Sangiovese' callus tissue maintained 70% viability when exposed to 50 μg/mL scytalone, while 'Trebbiano' callus viability dropped to 30% at equivalent concentrations, demonstrating both structural and biochemical components of enhanced Esca_Resistance_Disposition.
An Esca_Resistance_Disposition is a Trunk_Disease_Resistance_Disposition that inheres in a Vitis vinifera plant and is realized through defense processes that limit infection, colonization, and pathogenesis by the fungal complex causing esca disease, primarily Phaeomoniella chlamydospora (Pch), Phaeoacremonium minimum (Pm), and the basidiomycete Fomitiporia mediterranea (Fmed). This disposition represents the capacity to resist the progressive wood degradation and vascular dysfunction caused by these pathogens acting in succession or combination, manifesting as various syndromes including brown wood streaking (young vine decline), Petri disease, and classical esca with its characteristic white rot and tiger-stripe foliar symptoms. Esca resistance mechanisms include efficient wound healing limiting initial Pch colonization, compartmentalization of vascular infections through tylosis formation and accumulation of phenolic compounds in vessel-occluding gels, tolerance to the naphthoquinone phytotoxins (scytalone, isosclerone) produced by Pch and Pm, and resistance to the lignin-degrading enzymes of Fmed that cause white rot of heartwood. The chronic nature of esca, with symptoms often appearing years or decades after initial infection, complicates resistance evaluation. No cultivar is immune, but substantial variation exists, with 'Cabernet Sauvignon' and 'Merlot' generally showing better tolerance than 'Sauvignon Blanc', 'Ugni Blanc', and 'Grenache'. Understanding esca resistance dispositions is critical for cultivar and rootstock selection to maximize productive vineyard lifespan, as esca has become the most economically significant trunk disease in European viticulture.
Esca_Resistance_Disposition is_a Trunk_Disease_Resistance_Disposition. Esca is a disease complex involving multiple fungal species acting in succession: Phaeomoniella chlamydospora and Phaeoacremonium species cause vascular symptoms, while Fomitiporia mediterranea causes white rot of mature wood. Child classes include Wound_Compartmentalization_Disposition and Tylosis_Formation_Disposition representing specific resistance mechanisms. Esca resistance evaluation requires long-term studies due to the chronic disease progression, with controlled inoculation studies providing early indicators of potential field tolerance.
Esca-Resistenz-Disposition
Esca-Toleranz
Phaeomoniella resistance
Phaeomoniella-Resistenz
black measles resistance
esca complex tolerance
Gramaje D et al. (2018) Managing grapevine trunk diseases with respect to etiology and epidemiology, Plant Disease 102:12-39 Magnin-Robert M et al. (2011) Changes in plant metabolism and accumulation of fungal metabolites in response to esca proper and apoplexy expression in the whole grapevine, Phytopathology 101:424-436 Murolo S, Romanazzi G (2014) Effects of grapevine cultivar, rootstock and clone on esca disease, Australasian Plant Pathology 43:215-221
Esca Resistance Disposition
Esca symptom quality assessment in 25-year-old Barossa Valley Shiraz documented the prevalence and impact of esca complex in an aging vineyard approaching economic replanting threshold. Annual surveys scored esca symptom quality using the standard foliar symptom scale as 0 = no symptoms, 1 = mild interveinal chlorosis affecting 1-3 leaves, 2 = moderate tiger-stripe affecting 4-10 leaves, 3 = severe symptoms on more than 10 leaves or partial defoliation, 4 = apoplexy (complete sudden collapse). Cumulative esca symptom quality assessment showed 35% of vines expressing foliar symptoms at least once over a 5-year monitoring period, with year-to-year symptom expression highly erratic such that only 40% of previously symptomatic vines showed symptoms in consecutive years. Apoplexy events (esca symptom quality score 4) occurred in 3-5% of vines annually during hot January weather, with affected vines showing complete desiccation of leaves and fruit within days of initial wilting symptoms. Cross-sectional examination of symptomatic trunks revealed characteristic esca internal symptoms including white soft rot in the heartwood, dark vascular streaking radiating from pruning wounds, and the brown-black necrosis associated with Phaeomoniella colonization of xylem vessels. The esca symptom quality prevalence informed replanting decision, with 35% cumulative incidence combined with 3-5% annual mortality establishing that vineyard productivity was declining faster than acceptable, triggering block-by-block replacement planning prioritizing the most severely affected sections.
Esca symptom quality is a fungal infection indicator quality that inheres in grapevine organisms infected by the complex of wood-colonizing fungi causing esca disease and represents the characteristic disease manifestations including the dramatic "tiger-stripe" or "leopard-spot" interveinal chlorosis and necrosis on leaves, the black measles spotting on berries, the sudden vine death ("apoplexy") occurring without warning during hot weather, and the internal wood symptoms including the white soft rot and dark streaking visible in cross-sectioned trunks and cordons, with the esca complex caused primarily by Phaeomoniella chlamydospora, Phaeoacremonium minimum, and various basidiomycete wood-decay fungi including Fomitiporia mediterranea. This quality captures the diagnostic features of one of the most economically devastating trunk disease complexes affecting mature vineyards worldwide, recognizing that esca causes progressive vine decline reducing productivity over multiple seasons while also causing sudden catastrophic mortality that can eliminate individual vines without warning. The ontological significance of esca symptom quality lies in its role enabling diagnosis of this chronic, incurable disease affecting vine longevity and vineyard sustainability, with symptom recognition essential for identifying affected vines, estimating disease prevalence, and implementing management strategies aimed at limiting disease spread and maintaining productivity in affected vineyards. The biological basis of esca symptoms encompasses the vascular colonization by pioneer pathogens (Phaeomoniella, Phaeoacremonium) entering through pruning wounds, the subsequent wood decay by basidiomycete fungi creating the soft white rot, the toxin production by fungal colonizers causing the characteristic foliar symptoms, and the progressive vascular dysfunction leading to water stress that triggers the dramatic apoplexy symptoms during high evaporative demand conditions. The symptom expression in esca is notably erratic, with affected vines often showing severe foliar symptoms one season and appearing healthy the next, creating diagnostic challenges and complicating prevalence assessment. The "young esca" or Petri disease syndrome affecting young vines represents early infection by the vascular pathogens before wood decay fungi become established, showing stunted growth and mild leaf symptoms distinct from the dramatic tiger-stripe of mature-vine esca.
Esca_Symptom_Quality is_a Fungal_Infection_Indicator_Quality, representing the diagnostic symptoms of the esca disease complex caused by multiple wood-colonizing fungi. The tiger-stripe foliar pattern is pathognomonic but expression is erratic between seasons. Apoplexy (sudden vine death) typically occurs during hot weather with high evaporative demand. Internal wood symptoms (white rot, vascular streaking) visible only in cross-section. Disease complex involves multiple fungal species with different roles. Chronic nature affecting vineyard longevity makes this a primary economic concern in mature vineyards worldwide.
Esca-Symptome
Rebholzkrankheit-Symptome
Schwarze Masern-Symptome
Tigerstreifen-Symptome
black measles symptoms
esca symptoms
grapevine trunk disease symptoms
tiger stripe symptoms
Mugnai L et al. (1999) Esca (black measles) and brown wood-streaking: two old and elusive diseases of grapevines. Plant Disease 83:404-418 Bertsch C et al. (2013) Grapevine trunk diseases: complex and still poorly understood. Plant Pathology 62:243-265 Gramaje D et al. (2018) Managing grapevine trunk diseases with respect to etiology and epidemiology: current strategies and future prospects. Plant Disease 102:12-39
Esca Symptom Quality
A 7-year-old Syrah vine bears an Established_Vine_Role with full cordon structure and stable spur positions, producing 80% of expected mature yield while root system continues developing toward the depth and distribution characteristic of mature vines. An 8-year-old Chardonnay vine serves in an Established_Vine_Role contributing to commercial wine production, with fruit quality now approaching mature character though vintage-to-vintage variation remains higher than in fully mature blocks. A 6-year-old grafted vine recovered from young vine decline bears an Established_Vine_Role, its delayed development reaching productive stability later than optimally established vines.
A vine age role that inheres in a grapevine that has completed establishment and initial training but has not yet reached full mature productivity, typically spanning years 5-10 after planting when root system expansion continues, vine architecture stabilizes, and production approaches mature levels. Established vines have completed the major structural training (trunk, cordons or head, initial spur positions) and can support commercial crop loads, though yields and fruit character may continue evolving as root systems mature and vine balance optimizes. The established phase represents the transition from investment (establishment costs, restricted production) toward return (commercial yields, characteristic fruit quality), with vineyard economics improving as production increases while per-vine management intensity decreases. Established vines typically show vigorous growth capacity that must be managed through appropriate canopy practices, as the developing root system can supply more resources than the maturing vine optimally utilizes. Fruit quality from established vines may approach but not fully equal mature vine quality, with the continued physiological development reflected in evolving wine character across vintages.
Established_Vine_Role captures the intermediate phase between the intensive management of young vines and the stable productivity of mature vines. The category recognizes that vine development is continuous rather than abrupt—there is no single moment when a vine transitions from "establishing" to "mature"—with the established phase representing the period of continued development between clear youth and full maturity. Fruit from established vines is increasingly incorporated into premium programs as quality approaches mature standards, though some producers maintain distinction between established and mature vine sources for critical bottlings. The established phase duration varies substantially vigorous varieties on fertile sites may reach mature status by year 8, while low-vigor combinations on challenging sites may remain in extended establishment for 12+ years.
developing vine
entwickelnde Rebe
established vine
etablierte Rebe
intermediate-age vine
mittelaltrige Rebe
transitional vine
Übergangsrebe
Keller M (2015) The Science of Grapevines, Academic Press Dry PR, Coombe BG (2004) Viticulture Volume 1 - Resources, Winetitles Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles
Established Vine Role
Cabernet Sauvignon grafted on 3309C rootstock at 3.5 years post-grafting showing establishing grafted rootstock trunk 4.6 cm diameter, trunk extending 24 cm from root crown to graft union positioned 19 cm above soil, graft union displaying stable 1.8 cm diameter bulge with complete integration, trunk bark 1.4 mm thickness showing characteristic 3309C fine longitudinal fissures and grey-brown coloration, trunk self-supporting without stake, diameter growth 0.5 cm current season, no disease symptoms, supporting full canopy 3.8 m total shoot length and 2.2 kg fruit load indicating transition to production phase
Establishing grafted rootstock trunk is young grafted rootstock trunk in late young stage spanning approximately third year post-grafting and transition into fourth year, characterized by near-complete graft union integration, moderating diameter growth transitioning from rapid juvenile expansion toward stable mature growth rate, well-developed bark showing characteristic rootstock cultivar texture, diameter typically 3.5-5.5 cm approaching mature trunk dimensions, substantially reduced vulnerability with self-supporting capacity, and representing transition phase from young vulnerable trunk toward fully established mature rootstock trunk. Establishing grafted rootstock trunk temporal and developmental definition includes= spanning approximately year 3 post-grafting and transition into year 4 (typically months 24-48 post-grafting), representing late young trunk stage (final phase of young trunk period before achieving mature status), showing transition characteristics (combining some young trunk features with emerging mature trunk properties), approaching but not yet achieving full maturity (developing toward but not yet meeting all criteria for mature trunk classification), and marking establishment completion phase (consolidating foundation established in years 1-2 into robust sustainable trunk structure). Establishing grafted rootstock trunk graft union completion includes= showing largely complete callus integration (first 2 years' active callus formation finished with mature callus tissue fully incorporated into trunk structure), displaying well-established vascular continuity (xylem and phloem connections across graft union essentially complete providing efficient transport comparable to non-grafted trunk), exhibiting stable graft union bulge (swelling at graft union 1.2-2.5 cm diameter increase stabilized with minimal additional callus accumulation), showing increasingly obscure graft line (thickening bark and mature callus integration making original grafting point less visually prominent), and achieving mature mechanical integration (graft union strength approaching continuous trunk with minimal risk of mechanical failure under normal conditions). Establishing grafted rootstock trunk physical characteristics includes= exhibiting near-mature diameter (3.5-5.5 cm typical range approaching mature trunk dimensions of 4-8 cm), showing well-developed bark (periderm 1.0-2.5 mm thickness displaying characteristic rootstock cultivar texture), displaying established fissure patterns (longitudinal furrows or ridges characteristic of specific rootstock clearly visible), revealing mature bark coloration (grey-brown, reddish-brown, or tan tones typical of rootstock variety fully expressed), and showing robust appearance (trunk looking substantial and well-established rather than fragile or obviously young). Establishing grafted rootstock trunk bark maturation includes= forming mature periderm structure (multiple cork cambium layers creating protective bark with adult characteristics), showing rootstock-specific patterns (bark texture, fissure depth, and ridge patterns characteristic of rootstock cultivar clearly expressed), displaying weathered coloration (bark surface aged and weathered to mature color rather than fresh light colors of young trunk), developing pronounced lenticels (gas exchange pores visible as prominent horizontal lines or oval structures in maturing bark), and approaching final bark characteristics (bark approaching but not yet achieving maximum thickness and texture of old mature trunk). Establishing grafted rootstock trunk reduced vulnerability includes= substantially resistant to mechanical damage (well-developed bark and robust diameter providing good protection against equipment injury, wind damage, or animal browsing), showing improved environmental tolerance (better resistance to freeze damage and sunburn compared to young trunk though not yet maximum mature trunk hardiness), achieving self-supporting capacity (trunk diameter and lignification adequate for structural independence eliminating stake requirement in most cases), demonstrating moderate disease resistance (improved resistance to trunk cankers and crown gall compared to young trunk approaching mature trunk resistance), and requiring minimal protection measures (trunk protection generally unnecessary except in extremely challenging environments). Establishing grafted rootstock trunk growth transition includes= showing moderating diameter increase (annual growth 0.4-0.7 cm in establishing phase compared to 0.5-1.2 cm in earlier young trunk years), displaying transition from juvenile to mature wood properties (wood density, vessel size, and mechanical properties approaching mature trunk characteristics), exhibiting stabilizing growth patterns (year-to-year diameter increase becoming more consistent and predictable), forming distinct annual rings (third and fourth growth rings clearly visible in cross-section showing regular seasonal growth), and approaching mature growth rate (diameter increase declining toward stable 0.3-0.5 cm annual increment typical of mature trunk). Establishing grafted rootstock trunk vascular capacity includes= providing adequate water transport (xylem capacity 30-60 liters daily supporting full productive canopy), enabling complete nutrient distribution (phloem and xylem systems fully functional distributing resources throughout vine), showing efficient graft union transport (vascular connections at graft interface essentially eliminating transport resistance), supporting full crop load (trunk capacity adequate for commercial fruit production without overload stress), and building carbohydrate reserves (increasing trunk ray parenchyma accumulating significant starch reserves supporting vine resilience). Establishing grafted rootstock trunk structural capability includes= achieving full mechanical strength (wood properties and diameter providing complete structural support for canopy and crop), showing mature load-bearing capacity (trunk supporting full scion weight, canopy mass, and fruit load without stress or failure risk), displaying rigid wood properties (mature lignification creating stiff strong trunk resisting bending or breakage), maintaining upright position independently (self-supporting without staking in normal vineyard conditions), and providing stable foundation (robust trunk serving as reliable base for permanent vine structure). Establishing grafted rootstock trunk management transition includes= eliminating stake support (trunk achieving adequate strength and diameter for independence from staking), reducing intensive monitoring (less frequent inspection needed compared to young trunk though periodic checking still valuable), implementing standard mature vine practices (transitioning from establishment-focused management to production-focused practices), maintaining sucker removal (continuing to eliminate rootstock shoots below graft union though typically less vigorous than in young trunk), and beginning full cropping (allowing complete crop load as trunk capacity reaches adequate level for commercial production). Establishing grafted rootstock trunk in production transition includes= supporting first significant crop (enabling first commercial crop load in year 3 or 4 depending on scion cultivar and vineyard goals), providing adequate resources for fruit development (trunk capacity sufficient for both canopy maintenance and fruit maturation), showing balance capability (trunk able to support balanced vine with appropriate shoot growth and fruit load relationship), enabling sustainable production (trunk achieving capacity for continued annual cropping without excessive stress), and marking production phase entry (establishing trunk indicating vine ready for transition from establishment to production management). Establishing grafted rootstock trunk cultivar-specific characteristics includes= vigorous rootstocks (1103P, 110R) potentially reaching 5-6 cm diameter by end of year 3, moderate vigor rootstocks (SO4, 3309C, Teleki 5C) typically achieving 4-5 cm diameter establishing phase, low vigor rootstocks (Riparia Gloire, 101-14) reaching 3.5-4.5 cm diameter, thick-bark rootstocks showing pronounced fissuring and texture by establishing stage, and smooth-bark rootstocks maintaining relatively smooth appearance longer into establishing phase. Establishing grafted rootstock trunk transport efficiency includes= showing near-mature xylem area (cross-sectional xylem approaching mature trunk dimensions supporting full water transport needs), displaying complete phloem integration (bidirectional photosynthate and nutrient transport fully functional across graft union), exhibiting efficient vascular architecture (vessel size distribution and connectivity approaching optimal mature trunk patterns), conducting peak season water demands (meeting maximum transpiration requirements during hot periods without stress), and distributing resources to roots (adequate phloem capacity supporting root system maintenance and carbohydrate storage). Establishing grafted rootstock trunk as foundation indicator includes= predicting long-term vine success (healthy establishing trunk indicating excellent foundation for decades of productive life), showing integration quality (graft union condition in establishing phase revealing long-term compatibility and stability), demonstrating rootstock performance (establishing trunk characteristics reflecting whether rootstock choice appropriate for site and objectives), indicating vine balance (establishing trunk size and condition showing whether vine achieving appropriate vigor for training system and production goals), and marking maturity approach (establishing characteristics confirming vine progressing normally toward fully mature status). Establishing grafted rootstock trunk completion indicators includes= achieving 4+ cm diameter threshold (meeting minimum size criteria for mature trunk classification in most cultivars), displaying fully characteristic bark (rootstock-specific bark texture completely expressed), showing stable graft union (no additional callus formation and complete integration), demonstrating consistent growth (annual diameter increase stabilized at mature trunk rate), supporting full production (adequate capacity for commercial crop without stress), and requiring standard management (no longer needing establishment-specific protection or intensive monitoring). Establishing grafted rootstock trunk risk factors includes= residual vulnerability to severe stress (extreme drought, deep freeze, or mechanical damage still potentially causing more harm than fully mature trunk), potential trunk disease development (cankers or crown gall infections still possible though risk lower than young trunk), equipment damage risk (careless vineyard operations potentially injuring establishing trunk though damage less severe than in young trunk), overload potential (excessive crop or vigorous growth potentially stressing trunk that has not yet achieved maximum capacity), and management transition errors (inappropriate switch to mature vine practices before trunk truly ready potentially compromising development). Establishing grafted rootstock trunk significance includes= representing transition to maturity (establishing trunk marking critical shift from vulnerable young vine to robust productive vine), confirming successful establishment (healthy establishing trunk indicating first 2-3 years' management succeeded in building solid foundation), enabling commercial production (establishing trunk achieving capacity necessary for economic fruit production), requiring final establishment attention (last phase needing some establishment-focused management before full transition to mature practices), and predicting long-term success (establishing trunk characteristics strongly indicating whether vine will achieve desired productive lifespan and performance).
Establishing_Grafted_Rootstock_Trunk emphasizes transition from young to mature status (year 3-4) and is distinguished from Young_Grafted_Rootstock_Trunk (earlier establishment) and Mature_Grafted_Rootstock_Trunk (fully established) by focus on late young stage approaching maturity. Critical for understanding establishment completion, reduced vulnerability, transition to self-supporting capacity, production phase entry, and the final phase before achieving fully mature trunk status.
Late young grafted rootstock trunk
Near-mature grafted rootstock trunk
Reifender Pfropfstamm
Sich etablierender verpfropfter Unterlagenstamm
Transitioning grafted rootstock trunk
Übergangs-Wurzelstockstamm
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Pongrácz, D. P. (1983). Rootstocks for Grape-vines. David Philip Publisher.
Establishing Grafted Rootstock Trunk
Cabernet Sauvignon establishing grafted scion trunk on 110R vigorous rootstock in Napa Valley during years 2-3 develops bilateral cordons at 75 cm above union (85 cm above soil) with 14-18 spur positions by year 3 dormancy, achieves 36-48 mm scion diameter with 3 annual rings (year 1= 3.0 mm, year 2= 5.2 mm, year 3= 4.5 mm), rootstock diameter 42-54 mm below union, accumulates 85-110 g combined starch (scion 30-42 g, rootstock 55-68 g), supports year 3 crop of 22-28 clusters (2.8-4.2 kg fruit), and demonstrates vigorous rootstock-enhanced canopy requiring intensive shoot management
An establishing grafted scion trunk is a young grafted scion trunk during the critical architectural development period (typically years 2-3 post-planting) when permanent vine structure is defined, rootstock-scion interaction patterns are established, and transition from establishment to regular productivity occurs, representing the formative phase when rootstock vigor effects, graft union stability, and long-term scion architecture are confirmed. Establishing grafted scion trunk development during years 2-3 involves= selection and positioning of permanent cordons or head positions (accounting for union position 5-15 cm above soil), development of spur or cane positions for fruiting, progressive scion diameter increase (from 16-28 mm at year 2 start to 28-50 mm by year 3 end depending on rootstock), rootstock-mediated carbohydrate reserve accumulation (from 30-55 g combined at year 2 start to 60-105 g by year 3 end= scion 22-38 g, rootstock 38-67 g), and transition from minimal cropping to approaching commercial loads (0-12 clusters year 2, 12-28 clusters year 3 depending on rootstock vigor). Establishing grafted scion trunk architectural development varies by training system and rootstock= vigorous rootstock vertical shoot positioning (establishing 1-2 bilateral cordons at 60-90 cm height, rapid cordon extension 80-120 cm length by year 3 on SO4, 1103P, 110R), moderate rootstock systems (balanced cordon development 60-100 cm length by year 3 on 5BB, 420A, 101-14Mgt), low-vigor rootstock systems (slower cordon establishment 50-80 cm by year 3 on 3309C, Riparia Gloire), and head-trained systems (establishing head 35-65 cm above union with 3-8 positions, faster on vigorous rootstocks). Establishing grafted scion trunk anatomical characteristics include= scion annual ring formation (2.5-6 mm radial increments reflecting rootstock vigor, compared to 2.5-5 mm in own-rooted), scion bark thickening (2-3 mm year 2 to 3-5 mm year 3), graft union swelling stabilized (typically 108-120% equilibrium diameter ratio maintained years 2-3), expanding scion sapwood (years 2-3 rings providing majority of water transport with rootstock-influenced vessel characteristics), and beginning scion heartwood formation (central pith and year 1 wood may show early discoloration by year 3 end). Establishing grafted scion trunk rootstock influence on architecture includes= vigorous rootstocks promoting rapid architecture establishment (achieving full cordon length and spur positions by year 3, enabling year 4 commercial production), moderate rootstocks providing balanced development (achieving architecture by year 3-4, standard productivity timeline), low-vigor rootstocks slowing establishment (full architecture potentially delayed to year 4-5, particularly on low-fertility sites), and rootstock soil adaptation affecting timeline (well-adapted rootstock-soil combinations accelerating establishment, poorly adapted combinations delaying architecture development). Establishing grafted scion trunk management priorities include= architecture finalization accounting for union position (ensuring permanent cordons or head positions provide adequate clearance above graft union), continued vigilant rootstock sucker removal (preventing rootstock canopy development that would bypass scion), crop load progression managed for rootstock vigor (vigorous rootstock combinations may support 18-28 clusters year 3, low-vigor combinations limited to 8-15 clusters), graft union protection from damage and disease (union remains vulnerable to breakage and pathogen entry during architecture establishment), and canopy management establishing shoot density appropriate for rootstock vigor (vigorous rootstocks requiring more intensive shoot thinning, positioning). Establishing grafted scion trunk physiological development includes= rootstock-mediated photosynthetic capacity (total leaf area 2.0-4.5 m² by year 3 varying with rootstock vigor, compared to 1.8-3.5 m² own-rooted), rootstock effects on water relations (drought-tolerant rootstocks providing scion stress resistance, water-demanding rootstocks requiring irrigation), rootstock nutrient acquisition patterns (efficient rootstocks supporting vigorous scion development on low-fertility soils, lime-tolerant rootstocks preventing chlorosis on calcareous soils), and rootstock phenology effects (early rootstocks advancing scion budbreak and véraison 5-10 days, late rootstocks delaying phenology). Establishing grafted scion trunk compatibility confirmation includes= successful combinations showing balanced vigorous development (scion and rootstock diameters proportional, union swelling moderate, architecture developing on schedule), incompatible combinations requiring replanting (excessive union swelling >130%, severe scion stunting, premature decline symptoms), and marginal combinations with adapted management (acceptable but suboptimal growth requiring modified pruning, nutrition, or other interventions). Establishing grafted scion trunk variation by rootstock class includes= phylloxera-resistant Vitis riparia-based rootstocks (Riparia Gloire, 3309C producing low-vigor scions requiring longer establishment), drought-tolerant Vitis berlandieri-based rootstocks (140Ru, 1103P, SO4 producing vigorous scions with rapid establishment and early productivity), lime-tolerant rootstocks (41B, Fercal enabling scion establishment on challenging calcareous soils), and nematode-resistant rootstocks (Harmony, Freedom providing protection while influencing scion vigor patterns).
Establishing_Grafted_Scion_Trunk emphasizes formative architecture phase with rootstock influence and is distinguished from Young_Grafted_Scion_Trunk (broader establishment including year 1), specific year trunks (First/Second_Year), and Mature_Grafted_Scion_Trunk (completed architecture) by training system implementation with rootstock-mediated development. Critical for rootstock selection impacts on establishment timeline, architecture development accounting for union position, rootstock vigor effects on training decisions, and compatibility assessment during formative period.
Architecture establishment grafted trunk
Architekturentwicklung Veredelungsstamm
Erziehungsphase Aufpropfung
Etablierungsphase gepfropfter Stamm
Training phase grafted scion
Establishing Grafted Scion Trunk
Cabernet Sauvignon establishing young own-rooted trunk in Napa Valley during years 2-3 develops bilateral cordons at 75 cm height with 12-16 spur positions by year 3 dormancy, achieves 32-42 mm trunk diameter with 3 annual rings (year 1= 2.8 mm, year 2= 4.5 mm, year 3= 3.8 mm), accumulates 65-85 g starch reserves, supports year 3 crop of 18-24 clusters (2.0-3.5 kg fruit), and establishes mature canopy architecture ready for year 4 commercial production
An establishing young own-rooted trunk is a young own-rooted trunk during the critical architectural development period (typically years 2-3 post-planting) when permanent vine structure is defined, training systems are implemented, and transition from establishment to regular productivity occurs, representing the formative phase determining long-term vine architecture and productivity potential. Establishing young own-rooted trunk development during years 2-3 involves= selection and positioning of permanent cordons or head positions, development of spur or cane positions for annual fruiting, progressive trunk diameter increase (from 15-25 mm at year 2 start to 25-45 mm by year 3 end), increasing carbohydrate reserve accumulation (from 25-45 g at year 2 start to 50-90 g by year 3 end), and transition from minimal/no cropping to approaching commercial crop loads (0-10 clusters year 2, 10-25 clusters year 3 depending on vigor). Establishing young own-rooted trunk architectural development varies by training system= vertical shoot positioning systems (establishing 1-2 bilateral cordons at 60-90 cm height with spur positions every 10-20 cm), head-trained systems (establishing head at 30-60 cm height with 3-8 spur or cane positions), high-wire systems (establishing cordon at 120-180 cm height for pergola or overhead systems), and specialized systems (lyre, Scott Henry, Geneva Double Curtain requiring specific architectural development). Establishing young own-rooted trunk anatomical characteristics during years 2-3 include= annual ring formation producing 2.5-5 mm radial increments (compared to 1-3 mm in mature trunks and 5-8 mm in year 1), progressive bark thickening (increasing from 2-3 mm to 3-5 mm), expanding sapwood cross-section (years 2-3 rings providing majority of water transport), beginning heartwood formation (central pith and year 1 wood may show early discoloration or reduced function), and cordon or arm development showing similar anatomical progression. Establishing young own-rooted trunk management priorities include= architecture finalization (positioning permanent wood that will persist for vine lifetime), pruning decisions establishing fruiting positions (spur versus cane pruning implementation, shoot positioning), crop load progression (year 2 typically 0-1 kg fruit per vine, year 3 typically 1-3 kg per vine, approaching year 4+ target of 3-6 kg depending on training system and spacing), disease prevention at pruning wounds (critical period for trunk disease infection), and canopy management establishing desired shoot density and positioning. Establishing young own-rooted trunk physiological development includes= progressive increase in photosynthetic capacity (total leaf area expanding from 1.5-3 m² in year 2 to 2.5-4.5 m² in year 3), increasing carbohydrate production and storage capacity (starch reserves supporting larger crops and more vigorous growth), improving water transport capacity (expanding xylem cross-section providing 0.3-1.5 L/day capacity by year 3), and developing consistent annual productivity rhythm (bud fruitfulness becoming more uniform and predictable). Establishing young own-rooted trunk environmental response shows= increased stress tolerance compared to year 1 (larger root systems, greater reserves), but continued vulnerability to severe stress (extreme drought, winter injury in marginal climates, severe pest or disease pressure), relatively rapid recovery from moderate stress (adequate reserves supporting regrowth), and expression of site adaptation (beginning to show terroir influences on vigor, fruit quality, phenology). Establishing young own-rooted trunk variation by site and management includes= fertile sites or high water availability promoting rapid establishment (achieving commercial crop by year 3-4), low-fertility or water-limited sites showing slower establishment (commercial crop delayed to year 4-5), intensive management accelerating establishment (irrigation, nutrition, canopy management), and minimal intervention extending establishment period (dry-farmed, low-input systems).
Establishing_Young_Own_Rooted_Trunk emphasizes formative architectural development and is distinguished from First_Year_Own_Rooted_Trunk (initial establishment), Second_Year_Own_Rooted_Trunk (single year acceleration), and Mature_Own_Rooted_Trunk (established productive phase) by focus on training system implementation and architecture definition. Critical for long-term productivity foundation, optimal training decisions, crop load progression, and establishing disease-free trunk structure.
Architecture development phase
Architekturentwicklungs-Stamm
Erziehungsphase Rebstamm
Etablierungsphase-Stamm
Training establishment trunk
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Smart, R., & Robinson, M. (1991). Sunlight into Wine= A Handbook for Winegrape Canopy Management. Winetitles.
Establishing Young Own Rooted Trunk
Establishment cost for a new Blaufränkisch vineyard in Mittelburgenland (2023 cost estimates) documents a total investment of approximately €30,000-45,000/ha over the 4-year establishment period: land acquisition €8,000-15,000/ha (reflecting Mittelburgenland's moderate land values compared to more prestigious Austrian regions), site preparation €2,000-4,000/ha (soil analysis, subsoil preparation, drainage where needed), trellis system installation €8,000-12,000/ha (vertical shoot positioning system with galvanized steel end posts, intermediate posts, and 3-4 foliage wires), plant material €5,000-8,000/ha (certified Blaufränkisch grafted on appropriate rootstock, 4,500-5,500 vines/ha at €1.20-1.50 per grafted vine), planting operation €2,000-3,000/ha, and young vine maintenance €5,000-8,000/ha cumulative over 3 years of non-productive establishment — with first partial commercial harvest in year 3 and full production reached by year 5-6. Establishment cost for a Cabernet Sauvignon vineyard in Napa Valley (2023 estimates) documents a dramatically higher investment — total €250,000-500,000+/ha including land acquisition at €200,000-400,000+/ha reflecting Napa Valley's exceptional land values, with physical development costs of approximately €50,000-80,000/ha comparable in magnitude to other premium regions.
Establishment cost is a cost of production entity that documents the capital expenditure required to create a productive vineyard from the point of land acquisition or site preparation through to the first commercial harvest, encompassing land acquisition or lease costs (the price of vineyard-suitable land reflecting location, soil quality, water availability, and appellation prestige), site preparation costs (clearing, drainage, soil amendment, erosion control infrastructure), planting infrastructure costs (trellis system materials and installation, irrigation system where applicable, headland and access road construction), plant material costs (rootstock, scion wood, bench grafts, or potted vines, reflecting cultivar, clone, and certification status), planting operation costs (labor and machinery for vine installation, initial training), and young vine maintenance costs during the 2-4 year non-productive establishment period (canopy management, crop thinning or removal, weed and pest control, irrigation management until vines reach commercial production maturity), expressed per hectare with specification of the production system, region, and cost year.
Establishment cost documentation captures the capital investment dimension of vineyard economics — the substantial upfront expenditure that must be amortized over the productive life of the vineyard (typically 25-40 years for modern plantings, though some traditional vineyards produce commercially for 60-100+ years) and that represents the primary financial barrier to entry for new vineyard development. The cost structure of vineyard establishment is dominated by land acquisition in premium regions — in Burgundy Grand Cru, Napa Valley, Barossa Valley premium sites, and other prestige appellations, land cost may represent 80-90% of total establishment investment, making vineyard economics primarily a function of real estate dynamics rather than agricultural economics in these locations. In regions with moderate land values (most of Central Europe, southern France, much of the New World outside prestige zones), physical development costs (trellis, irrigation, plant material, labor) typically represent the majority of establishment investment. The cultivar-specific dimension of establishment cost primarily manifests through plant material costs — certified virus-free grafted vines of popular cultivars and clones are produced at scale by commercial nurseries at relatively low unit costs (€1.00-2.00/vine in Europe), while rare cultivars, heritage varieties, or recently released clones may command substantial premiums (€3-5+/vine) due to limited nursery production and high demand, and PIWI varieties may carry additional premiums reflecting smaller production volumes and patent or licensing fees for protected varieties. The trellis system selection, which significantly affects establishment cost, is influenced by cultivar characteristics — vigorous cultivars may require more elaborate trellis systems with higher infrastructure costs, while compact cultivars may be suited to simpler, less expensive training systems. The time value of capital during the establishment period represents a significant hidden cost — the 3-5 years between planting and first commercial harvest represent not only the direct maintenance costs but also the opportunity cost of capital tied up in a non-productive asset, which at typical agricultural discount rates (5-8%) adds substantially to the effective establishment cost.
Anpflanzungskosten
Aufbaukosten
Errichtungskosten
Investitionskosten
Pflanzkosten
capital investment
development cost
planting cost
setup cost
vineyard establishment cost
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Archer E, Van Schalkwyk D (2007) The effect of alternative pruning methods on the viticultural and oenological performance of some wine grape varieties. South African Journal of Enology and Viticulture 28(2): 107-139 LWK Burgenland (2023) Kosten der Weingartenanlage — Kalkulationsgrundlagen UC Davis (2023) Vineyard establishment cost studies, Department of Viticulture and Enology
Establishment Cost
Establishment date for Cabernet Sauvignon in Coonawarra, South Australia distinguishes four chronological milestones: first introduction (1891, John Riddoch's plantings at Katnook Estate, though cultivar identity is uncertain from early records), first confirmed commercial planting (1950s, replanting by Wynns and Mildara following the district's post-war revival), first significant vintage attracting critical recognition (1962, Wynns Coonawarra Estate Cabernet Sauvignon), and period of major area expansion establishing Coonawarra as Australia's premier Cabernet region (1970s-1990s, area increasing from approximately 200 ha to over 2,000 ha of Cabernet Sauvignon). Establishment date for Grüner Veltliner in the Wachau documents pre-20th-century presence in mixed plantings (no precise date, inferred from vineyard surveys), first documented varietal vineyard plantings (1950s-1960s, as part of the post-war Austrian viticultural restructuring), critical recognition milestone (1985, founding of Vinea Wachau and the subsequent quality revolution that positioned Grüner Veltliner as the region's flagship), and major area expansion confirming dominance (1990s-2000s, Grüner Veltliner reaching approximately 50% of Wachau vineyard area).
Establishment date is a spread history entity that documents the chronological milestones marking the permanent, commercially significant cultivation of a grape cultivar in a specific viticultural region, distinguished from the initial introduction date (which records the first arrival of plant material) by requiring evidence of sustained cultivation beyond experimental or trial status, encompassing the date or date range of first commercial vineyard planting, the date of first commercial vintage production, the date of official varietal recognition or registration in the receiving region, and the date of significant area expansion indicating industry-level adoption, expressed with appropriate precision (exact date, year, decade, or approximate period) and confidence level, and providing the temporal framework for assessing a cultivar's depth of establishment in each region of its global distribution.
Establishment date documentation addresses the important distinction between a cultivar's first arrival in a region and its permanent integration into the regional viticultural economy — a distinction that is often obscured in simplified historical narratives that conflate introduction with establishment. Many cultivar introductions fail to achieve permanent establishment — trial plantings may be abandoned due to climatic unsuitability, disease susceptibility, market indifference, or disruption by historical events (wars, phylloxera, economic crises), and even successful introductions may require decades of gradual adoption before achieving commercial significance. The multi-milestone framework adopted here (introduction, first commercial planting, first commercial vintage, official recognition, significant expansion) captures this temporal depth, enabling differentiation between cultivars with deep establishment histories (centuries of continuous cultivation, as with Riesling in the Rheingau) and those with shallow but rapidly expanding establishment (recent introduction followed by swift commercial adoption, as with Grüner Veltliner in New Zealand). The precision of establishment dates varies dramatically by region and historical period — European regions with rich archival traditions may document establishment dates to the exact year or even month, while New World regions often have precise dates for documented introductions but imprecise timelines for the transition from trial to commercial cultivation. The phylloxera crisis created a widespread establishment discontinuity in European viticulture — many regions effectively re-established their cultivar portfolios during the post-phylloxera replanting period (1880s-1920s), meaning that the functional establishment date of many European cultivars in their traditional regions is substantially more recent than the historical first mention might suggest. The contemporary wave of cultivar introduction driven by climate change adaptation — warm regions trialing heat-tolerant Mediterranean cultivars, cool regions exploring previously marginal varieties — is creating a new generation of establishment events whose long-term success remains uncertain.
Anbaubeginn
Anbauzeitpunkt
Etablierungsdatum
Etablierungszeitpunkt
Kultivierungsbeginn
commercial planting date
cultivation commencement
cultivation milestone
establishment date
permanent establishment
Halliday J (1994) A History of the Australian Wine Industry 1949-1994, Australian Wine and Brandy Corporation Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes: A Complete Guide to 1,368 Vine Varieties, Allen Lane Unwin T (1991) Wine and the Vine, Routledge, pp.: 300-400 Anderson K, Aryal NR (2013) Which winegrape varieties are grown where? A global empirical picture. University of Adelaide Press, pp.: 1-700
Establishment Date
In Malvasia Bianca berries in Friuli, Italy during BBCH 85-89, ester formation generates total volatile esters increasing from 45 to 185 µg/L over 35 days, with ethyl acetate rising from 18 to 65 µg/L, isoamyl acetate from 8 to 42 µg/L providing banana-fruity notes, and ethyl hexanoate from 5 to 28 µg/L contributing apple-fruity character, establishing the ester profile that will serve as template for fermentation-enhanced ester development under warm conditions averaging 25°C with significant diurnal temperature variation.
Ester formation describes the biosynthesis of volatile ester compounds in Vitis vinifera berries during ripening, wherein alcohol acetyltransferases and related acyltransferase enzymes catalyze the condensation of alcohol and acyl-CoA substrates to produce the fruity, sweet-smelling esters that contribute distinctive aromatic notes to ripe grape character and serve as precursors for fermentation-derived ester accumulation during subsequent winemaking. This process involves the enzymatic transfer of acyl groups from activated acyl-CoA donors to hydroxyl-bearing alcohol acceptors, with acetyl-CoA serving as the predominant acyl donor producing acetate esters such as ethyl acetate, hexyl acetate, and isoamyl acetate, while longer-chain acyl-CoAs generate ethyl esters of fatty acids including ethyl butanoate, ethyl hexanoate, and ethyl octanoate contributing fruity-tropical aromatic character. The substrate specificity of grape alcohol acyltransferases determines the ester profile produced during berry ripening, with these enzymes accepting diverse alcohol substrates including ethanol, higher alcohols derived from amino acid catabolism, and C6 alcohols from lipid oxidation, generating the complex mixture of esters characteristic of ripe grape aroma. The quantitative contribution of berry-derived esters to total wine ester content is relatively modest compared to fermentation-derived esters produced by yeast metabolism, with grape esters typically contributing 5-20% of final wine ester concentrations while providing important baseline aromatic character and serving as indicators of ripeness and fruit quality during harvest assessment. Temporal dynamics of ester formation show progressive increase during ripening coincident with alcohol substrate accumulation and acyltransferase enzyme expression, with ester concentrations typically rising 3-8 fold between veraison and harvest as both substrates and enzyme activities increase during berry maturation. Environmental factors influence ester formation through temperature effects on enzyme activities and substrate availability, with moderate temperatures of 20-26°C generally supporting optimal ester synthesis while temperature extremes may reduce enzyme function or alter substrate pools affecting ester production capacity. Cultivar variation in ester formation reflects genetic differences in acyltransferase expression and alcohol precursor accumulation, with some varieties producing more prominent ester-derived fruity character than others under equivalent ripening conditions.
Ester_Formation is subclass of Volatile_Compound_Synthesis representing acyltransferase-mediated ester biosynthesis. Distinguishes from fermentation-derived ester production by yeast. Sibling to Free_Volatile_Production and Aldehyde_Synthesis. Grape esters contribute 5-20% of final wine ester content. Provides fruity-sweet aromatic character. Progressive increase during ripening.
Acyltransferase-vermittelte-Veresterung
Ester-Bildung
Frucht-Ester-Formation
acyltransferase-mediated esterification
ester aroma synthesis
flüchtige Ester-Biosynthese
fruit ester formation
volatile ester biosynthesis
Kalua CM and Boss PK (2009) Evolution of volatile compounds during development of Cabernet Sauvignon grapes. Journal of Agricultural and Food Chemistry 57: 3818-3830 Dunlevy JD et al (2013) The production of flavour and aroma compounds in grape berries. In The Biochemistry of the Grape Berry Bentham Science pp 293-340 Keller M (2020) The Science of Grapevines 3rd Ed pp 288-292
Ester Formation
Etymology record for Riesling documents a contested derivation with multiple competing hypotheses: (1) from Middle High German "rîzen" (to tear, to crack), possibly referring to the small, tightly packed berries that split easily, supported by the early orthographic form "Rießling" (2) from "Rußling," referring to the dark (sooty) coloration of the mature wood (3) from a toponym, possibly the Riesling creek near the village of Rissling in Alsace (4) from "Rissling" as a diminutive related to "Riß" (crack or gap) scholarly consensus is lacking, with the geographic derivation (hypothesis 3) currently considered least likely and the morphological derivations (hypotheses 1 and 2) most plausible but not definitively established. Etymology record for Grüner Veltliner combines a transparent color descriptor ("Grüner" — green, referring to the berry color at maturity) with the more opaque "Veltliner," derived from the town of Veltlin (Valtellina) in Lombardy, northern Italy — though genetic analysis has not confirmed any Italian origin for the cultivar, suggesting that the name may reflect a perceived similarity to Italian wines or a false historical attribution rather than actual geographic provenance. Etymology record for Blaufränkisch combines "Blau" (blue, referring to the dark berry color) with "Fränkisch" (Frankish), historically used in Central European wine culture to distinguish higher-quality varieties (fränkisch/Frankish) from ordinary varieties (heunisch/Hunnish) — a quality-marker etymology rather than a geographic derivation, paralleling the Hungarian synonym "Kékfrankos" (kék = blue, frankos = Frankish).
Etymology record is an origin record entity that documents the linguistic history, derivation, and semantic evolution of grape cultivar names, encompassing the proposed etymological root (the word, morpheme, or phrase from which the name derives), the source language or languages, the semantic motivation for the naming (whether the name refers to a geographic origin, a morphological characteristic, a person, a flavor attribute, or another referent), alternative etymological hypotheses where the derivation is disputed, the chronological development of the name through successive orthographic and phonetic forms, and the relationship between the primary name and its regional synonyms, providing the linguistic dimension of cultivar identity that often encodes historical information about origin, migration, and cultural perception unavailable from other sources.
Etymology record documentation illuminates one of the most linguistically rich and historiographically underexploited dimensions of cultivar identity, as grape cultivar names frequently encode historical information about origin, migration, morphology, and cultural valuation that supplements or even contradicts evidence from other sources. The naming conventions for grape cultivars fall into several broad etymological categories — geographic names (Riesling possibly from a place name, Trebbiano from the Trebbia River, Aglianico from "Hellenikon"/Greek), morphological descriptions (Cabernet referring to small berries, Gros Plant referring to vigorous growth, Blaufränkisch referencing blue berries), personal names (Müller-Thurgau from its breeder Hermann Müller, Pinotage from its parental portmanteau Pinot × Hermitage-Cinsault), quality indicators (the fränkisch/heunisch distinction encoding a quality hierarchy), and sensory attributes (Muscat from "musk" referring to the aroma, Gewürztraminer from "Gewürz" = spice plus the town of Tramin). The fränkisch/heunisch dichotomy documented in the Blaufränkisch etymology is particularly revealing of medieval Central European viticultural culture — the Frankish designation (associated with Carolingian-introduced quality viticulture) versus the Hunnish designation (associated with inferior, possibly pre-Carolingian varieties) encodes a cultural hierarchy that influenced cultivar naming across the German-speaking viticultural world for centuries. The relationship between cultivar names and their synonyms across languages and regions provides a linguistic map of cultivar migration — the same variety acquiring different names as it crossed linguistic and political boundaries, with the pattern of naming conventions encoding the direction and cultural context of dissemination. Etymological claims must be evaluated with the same critical rigor as other historical evidence — folk etymologies (plausible-sounding but linguistically unfounded derivations) are common in viticultural tradition, and even scholarly etymological proposals often remain hypothetical due to the limited and ambiguous nature of the available linguistic evidence.
Namensdeutung
Namensherkunft
Sortennamenetymologie
cultivar name history
etymologische Aufzeichnung
linguistic origin
name derivation
name etymology
onomastic record
onomastische Dokumentation
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes: A Complete Guide to 1,368 Vine Varieties, Allen Lane Kramer M (2004) New California Wine: Making Sense of Napa Valley, Sonoma, Central Coast, and Beyond, Running Press, pp.: 30-55 Staab J (2015) Die Erstnennung des Riesling von 1435. Schriften zur Weingeschichte 188: 1-20 Vouillamoz J, Grando MS (2006) Genealogy of wine grape cultivars: "Pinot" is related to "Syrah." Heredity 97(2): 102-110
Etymology Record
Eutypa dieback symptom quality assessment in Napa Valley Cabernet Sauvignon documented the progressive structural disease requiring systematic surgical management across a 30-year-old planting. Spring scouting at BBCH 15-19 identified vines showing Eutypa dieback symptom quality with characteristic stunted shoots emerging from specific spurs, affected shoots measuring 5-10 cm versus 25-40 cm for healthy shoots on the same vine and showing the distinctive small, chlorotic, cupped leaves with necrotic margins. Mapping of Eutypa dieback symptom quality expression over 3 seasons identified 28% of vines as affected, with spatial analysis showing concentration along row ends and headland areas where pruning debris accumulated, suggesting inoculum concentration driving localized disease pressure. Surgical remediation followed identification, with cross-sectional cutting to locate the Eutypa characteristic wedge-shaped canker, then removal of affected wood with 10 cm margin beyond visible necrosis and wound protection with fungicidal paste. The Eutypa dieback symptom quality assessment and remediation program reduced annual symptomatic vine proportion from 12% to 4% over 5 years, demonstrating that systematic monitoring combined with surgical removal could maintain vineyard productivity despite endemic disease presence, extending productive vineyard life compared to blocks where disease was allowed to progress unmanaged.
Eutypa dieback symptom quality is a fungal infection indicator quality that inheres in grapevine organisms infected by the fungus Eutypa lata and represents the characteristic disease manifestations including the stunted spring shoots with small, chlorotic, cupped leaves showing necrotic margins emerging from spurs adjacent to old pruning wounds, the progressive dieback of cordons and arms as the canker expands, and the wedge-shaped necrotic sector visible in cross-sectioned trunks and cordons that is pathognomonic for this disease, with symptom expression typically delayed 3-5 years after initial infection through pruning wounds during the prolonged latent colonization period. This quality captures the diagnostic features of a major trunk disease affecting grapevine longevity and vineyard productivity, recognizing that Eutypa dieback causes progressive structural decline requiring surgical removal of affected wood or vine replacement, with the characteristic foliar symptoms providing diagnostic opportunity only during spring when affected shoots emerge from spurs near the infection site. The ontological significance of Eutypa dieback symptom quality lies in its role enabling diagnosis during the brief window when foliar symptoms are visible, identification of affected vine structures for surgical remediation, and assessment of disease prevalence for vineyard management and longevity planning decisions. The biological basis of Eutypa dieback symptoms encompasses the wound infection by E. lata ascospores released from perithecia on dead wood during rainfall, the slow colonization of xylem vessels and wood parenchyma extending from the infection site over multiple years, the production of phytotoxins (eutypine and derivatives) translocated to developing shoots causing the characteristic stunting and chlorosis, and the eventual girdling of vascular tissue causing dieback of distal vine portions. The seasonal timing of symptom expression reflects the translocation of toxins to actively growing shoots, with symptoms most apparent in spring when affected shoots emerge stunted and chlorotic from spurs positioned near the canker margin, then often obscured later in the season as healthy shoots dominate the canopy. The diagnostic confirmation through cross-sectional examination reveals the characteristic hard, dark, wedge-shaped necrotic sector extending from the pruning wound into the wood, distinct from the soft white rot of esca and providing definitive diagnosis when foliar symptoms are ambiguous.
Eutypa_Dieback_Symptom_Quality is_a Fungal_Infection_Indicator_Quality, representing the diagnostic symptoms of Eutypa lata infection causing progressive trunk disease and dieback. The stunted, chlorotic spring shoots with cupped leaves are pathognomonic during the brief spring visibility window. Internal wedge-shaped canker in cross-section provides definitive diagnosis. The 3-5 year latent period between infection and symptom expression complicates management timing. Pruning wound protection during wet weather is primary prevention strategy. Surgical removal of affected wood can extend vineyard life when disease is managed systematically.
Eutypa canker
Eutypa dieback signs
Eutypa symptoms
Eutypa-Absterben-Symptome
Eutypa-Krebs
Eutyplose-Zeichen
dead arm symptoms
toter Arm-Symptome
Moller WJ, Kasimatis AN (1978) Dieback of grapevines caused by Eutypa armeniacae. Plant Disease Reporter 62:254-258 Gramaje D et al. (2018) Managing grapevine trunk diseases with respect to etiology and epidemiology: current strategies and future prospects. Plant Disease 102:12-39 Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests, 2nd Edition. APS Press
Eutypa Dieback Symptom Quality
Controlled inoculation studies of Eutypa_Resistance_Disposition at CSIRO Adelaide compared host responses across 18 cultivars following E. lata trunk inoculation. Vines were inoculated with mycelial plugs inserted into drill wounds on the trunk, and disease development was assessed at 12 and 24 months through destructive sampling of inoculated trunks. Canker dimensions (length, width, depth) and pathogen extent (visualized by wood discoloration and confirmed by re-isolation) varied dramatically among cultivars. 'Grenache' exhibited strong Eutypa_Resistance_Disposition with mean canker length of 28 mm and pathogen reisolation limited to 35 mm from inoculation point at 24 months. Susceptible 'Shiraz' developed cankers averaging 95 mm length with pathogen recovery to 120 mm. 'Grenache' also showed enhanced tolerance to eutypine in detached leaf bioassays, with 50% less necrosis than 'Shiraz' at equivalent toxin concentrations (50 μg/mL), suggesting both structural resistance (compartmentalization) and biochemical tolerance contribute to its Eutypa_Resistance_Disposition. Field surveys in South Australian vineyards corroborated experimental findings, with 'Grenache' blocks showing 60% lower incidence of Eutypa symptoms than adjacent 'Shiraz' of equivalent age.
A Eutypa_Resistance_Disposition is a Trunk_Disease_Resistance_Disposition that inheres in a Vitis vinifera plant and is realized through defense processes that limit infection, colonization, and pathogenesis by Eutypa lata and related diatrypaceous fungi causing Eutypa dieback (also known as dying arm disease). This disposition represents the capacity to resist the progressive wood necrosis, canker formation, and stunted shoot growth caused by E. lata colonization of pruning wounds and subsequent spread through vascular and structural wood tissues. Eutypa dieback symptoms include wedge-shaped cankers visible in trunk cross-sections, stunted spring shoots with small, chlorotic, cup-shaped leaves, and eventual death of affected cordons or entire vines. Resistance mechanisms operating against E. lata include efficient wound healing that limits the infection window (ascospore germination and penetration primarily occur within 2-4 weeks of pruning), compartmentalization restricting mycelial spread from initial infection sites, and tolerance to eutypine and other phytotoxic secondary metabolites produced by the fungus that cause the characteristic stunted growth symptoms. Substantial variation in Eutypa susceptibility exists among cultivars, with 'Cabernet Sauvignon', 'Shiraz', and 'Chenin Blanc' considered highly susceptible, while 'Merlot', 'Semillon', and 'Clairette' show greater tolerance. No cultivar is immune, but selection for enhanced Eutypa_Resistance_Disposition can significantly extend productive vineyard life in regions where this disease is prevalent, particularly in Mediterranean and warm-temperate wine regions with winter rainfall patterns.
Eutypa_Resistance_Disposition is_a Trunk_Disease_Resistance_Disposition. Eutypa dieback is distinct from esca in etiology (Eutypa lata versus Phaeomoniella/Phaeoacremonium/Fomitiporia complex) and symptomatology (stunted shoots with small leaves versus tiger-stripe foliar symptoms), though both diseases may co-occur in the same vine. Eutypa resistance involves both wound healing/compartmentalization and tolerance to the phytotoxin eutypine. Disease evaluation requires multi-year studies due to slow pathogen progression, though controlled inoculation accelerates screening compared to natural infection studies.
Eutypa dieback tolerance
Eutypa lata resistance
Eutypa-Resistenz-Disposition
Eutypa-lata-Toleranz
Eutypiose-Resistenz
dying arm resistance
Sosnowski MR et al. (2007) Influence of grapevine cultivar and isolate of Eutypa lata on wood and foliar symptoms, Plant Disease 91:924-931 Peros JP, Berger G (1994) A rapid method to assess the aggressiveness of Eutypa lata isolates and the susceptibility of grapevine cultivars to Eutypa dieback, Agronomie 14:515-523 Trouillas FP et al. (2010) Taxonomy and DNA phylogeny of Diatrypaceae associated with Vitis vinifera and other woody plants in California, Mycologia 102:319-336
Eutypa Resistance Disposition
PIWI selection Calardis Blanc (Gf.1993-22-6) at the Julius Kühn-Institut bearing an evaluation cultivar role is planted in replicated trials across 6 German sites (Geisenheim, Weinsberg, Freiburg, Siebeldingen, Neustadt, Veitshöchheim) in randomized complete block designs with 4 replications × 48 vines per plot, evaluated over 8 consecutive vintages for yield stability (CV 80° Oechsle, TA 6-9 g/L across sites), and wine sensory acceptability (DLG score > 3.5/5.0 across 25+ trained panelists), culminating in application for Sortenschutz and entry into the Bundessortenamt descriptive variety list.
An experimental cultivar role that inheres in a grapevine genotype that has passed initial screening trials and is undergoing advanced multi-site, multi-year evaluation in replicated field trials designed to comprehensively characterize agronomic performance, fruit quality, disease resistance stability, and wine sensory properties under diverse environmental conditions prior to potential cultivar release and registration.
Evaluation_Cultivar_Role represents the penultimate stage before cultivar release, typically spanning 5-12 years of intensive multi-environment testing. At this stage, experimental resources per genotype are substantial — replicated multi-site trials, annual micro-vinifications, formal sensory evaluation panels, and comprehensive disease resistance monitoring — reflecting the significant investment required to validate a candidate's commercial potential: only 1-5% of genotypes entering the evaluation stage ultimately achieve cultivar release and registration. The role terminates upon either official cultivar registration (the genotype transitions to Registered_Clone_Role or equivalent) or rejection from the evaluation pipeline.
Evaluierungssorten-Rolle
Leistungsprüfungs-Rolle
Zulassungskandidaten-Rolle
advanced evaluation role
elite selection role
pre-release candidate role
Eibach R and Töpfer R (2015) Traditional grapevine breeding techniques, in Reynolds AG (ed) Grapevine Breeding Programs for the Wine Industry, Woodhead Publishing pp: 3-22 Bundessortenamt (2023) Richtlinien für die Durchführung von Prüfungen auf Unterscheidbarkeit, Homogenität und Beständigkeit — Rebe Schneider C et al. (2019) Grapevine breeding programs in France, in Ferrão MAG et al. (eds) Genetic Improvement of Tropical Crops, Springer pp: 301-344
Evaluation Cultivar Role
Field investigation of Evaporative_Cooling_Disposition at UC Davis Oakville Research Station compared canopy temperature dynamics across 12 cultivars under hot conditions (> 38°C air temperature) using infrared thermography and concurrent gas exchange measurements. Canopy-to-air temperature differential (Tc-Ta), stomatal conductance, and vine water status (stem water potential) were monitored hourly from 10:00 to 16:00 on heat event days. Cultivars displayed contrasting Evaporative_Cooling_Disposition reflecting their stomatal regulation strategies. Anisohydric 'Syrah' maintained high stomatal conductance (0.25 mol m⁻² s⁻¹ at midday) and strong evaporative cooling (Tc-Ta = -8.5°C at 14:00), keeping canopy temperature at 32°C when air reached 40.5°C. Isohydric 'Grenache' showed progressive stomatal closure (conductance declining to 0.08 mol m⁻² s⁻¹ by 14:00), reduced cooling (Tc-Ta = -3.2°C), and canopy temperature reaching 37.3°C. Leaf damage (marginal necrosis) was observed on 'Grenache' following consecutive heat days, while 'Syrah' showed no visible damage despite identical air temperature exposure, demonstrating the protective value of Evaporative_Cooling_Disposition. However, stem water potential measurements revealed that 'Syrah' reached -1.5 MPa by late afternoon versus -1.0 MPa for 'Grenache', highlighting the water cost of aggressive cooling. Under limited irrigation, 'Grenache' maintained better water status and avoided late-season water stress, illustrating the environment-dependent value of different Evaporative_Cooling_Disposition strategies.
An Evaporative_Cooling_Disposition is a Heat_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to dissipate heat through transpirational water loss, maintaining leaf and canopy temperatures below ambient air temperature during hot conditions through the latent heat of evaporation. This disposition represents the hydraulic component of heat tolerance, with transpirational cooling potentially reducing leaf temperatures by 5-15°C below air temperature depending on evaporative demand and water availability. Effective evaporative cooling requires adequate soil water supply, hydraulic conductance sufficient to replace transpired water, stomatal aperture maintained despite high evaporative demand, and canopy architecture facilitating air movement and heat dissipation. Cultivars differ in evaporative cooling capacity based on stomatal behavior (isohydric cultivars close stomata conserving water but sacrificing cooling, anisohydric cultivars maintain stomatal opening and cooling under moderate stress), hydraulic conductance (root and shoot xylem properties), and canopy characteristics affecting boundary layer conductance. Evaporative_Cooling_Disposition represents a trade-off between heat stress avoidance through cooling and drought stress risk through water expenditure, with optimal strategy depending on the relative severity and timing of heat versus water stress in specific environments. In irrigated viticulture with adequate water supply, high evaporative cooling capacity provides significant heat stress mitigation, while in rainfed or deficit irrigation systems, conservative stomatal behavior may be preferable despite reduced cooling.
Evaporative_Cooling_Disposition is_a Heat_Tolerance_Disposition. Evaporative cooling links heat and water relations, as transpiration provides cooling but consumes water. Isohydric versus anisohydric stomatal behavior determines the heat/water stress trade-off strategy. Cooling capacity depends on water availability, hydraulic conductance, stomatal behavior, and boundary layer conditions. In well-watered conditions, high cooling capacity protects against heat in water-limited conditions, conservative stomatal behavior may be preferable. Canopy temperature monitoring (thermal imaging) directly assesses evaporative cooling effectiveness.
Hydraulische-Thermoregulation
Transpirative-Kühlung
Verdunstungskühlungs-Disposition
evaporative heat dissipation
hydraulic thermoregulation
transpirational cooling
Jones HG (1999) Use of infrared thermometry for estimation of stomatal conductance as a possible aid to irrigation scheduling, Agricultural and Forest Meteorology 95:139-149 Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought, Plant, Cell and Environment 26:1393-1405 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data, Annals of Botany 105:661-676
Evaporative Cooling Disposition
Evaluation of Evasion_Based_Resistance_Disposition at INRAE Bordeaux compared powdery mildew development on 'Sauvignon Blanc' (compact clusters, dense canopy) versus 'Merlot' (looser clusters, more open canopy) under identical management and natural disease pressure. Despite similar genetic susceptibility to E. necator in controlled inoculation tests (OIV rating 3 for both), field disease severity differed significantly. 'Merlot' clusters exhibited 40% lower berry infection severity (mean 18% versus 30% in 'Sauvignon Blanc') correlated with its looser cluster architecture (OIV 204 rating 5 versus 7) promoting faster drying after rainfall and dew events. Microclimate monitoring revealed that cluster-zone relative humidity exceeded 80% for 2.3 fewer hours per day in 'Merlot' canopy compared to 'Sauvignon Blanc'. Similarly, training system comparisons showed that VSP (Vertical Shoot Positioning) trained 'Chardonnay' with optimal leaf removal exhibited 35% lower cluster infection than minimally pruned 'Chardonnay', demonstrating how viticultural management can enhance evasion-based resistance through canopy architecture modification even in genetically identical material.
An Evasion_Based_Resistance_Disposition is a Powdery_Mildew_Resistance_Disposition that inheres in a Vitis vinifera plant and is realized through architectural, phenological, or morphological characteristics that reduce pathogen encounter, establishment, or epidemic development without direct physiological or genetic resistance mechanisms. This disposition encompasses traits that enable plants to "escape" disease through spatial or temporal avoidance of optimal infection conditions rather than through active defense responses. Architectural evasion mechanisms include open canopy structure promoting air circulation and rapid tissue drying, upright shoot growth reducing humidity in the fruiting zone, and loose cluster architecture minimizing berry-to-berry contact surfaces favorable for fungal spread. Phenological evasion involves flowering and fruit development timing that positions susceptible stages outside periods of peak inoculum availability or environmental conditions favorable for infection. Morphological evasion includes traits such as reduced leaf hair density that may impede conidial adhesion, glossy cuticle surfaces that affect spore germination microenvironments, and sparse cluster compactness reducing microhabitat humidity. While not providing true resistance, evasion-based mechanisms can substantially reduce disease severity in field conditions, contributing to integrated disease management strategies and reducing reliance on chemical control. These traits are often under quantitative genetic control and can be selected in breeding programs alongside true resistance genes.
Evasion_Based_Resistance_Disposition is_a Powdery_Mildew_Resistance_Disposition. This class represents disease avoidance through architectural, phenological, or morphological traits rather than physiological resistance mechanisms. Evasion operates independently of true resistance genes and can be enhanced through viticultural management practices. The distinction from true resistance is important: evasion reduces disease through avoiding favorable conditions for the pathogen rather than limiting pathogen development after contact. Traits contributing to evasion are often quantitatively inherited and amenable to selection.
Architektonische Mehltau-Resistenz
Ausweich-basierte Resistenz-Disposition
Krankheitsvermeidungs-Disposition
architectural PM avoidance
disease escape disposition
phenological disease evasion
Austin CN, Wilcox WF (2011) Effects of fruit-zone leaf removal, training systems, and irrigation on the development of grapevine powdery mildew, American Journal of Enology and Viticulture 62:193-198 Molitor D et al. (2012) Impact of grape cluster division on cluster morphology and bunch rot epidemic, American Journal of Enology and Viticulture 63:508-514 Dry IB et al. (2010) Molecular strategies to enhance the genetic resistance of grapevines to powdery mildew, Australian Journal of Grape and Wine Research 16:94-105
Evasion Based Resistance Disposition
In a Rioja Tempranillo vineyard in Spain, even berry coloration was documented in late September Year N+1 (BBCH 86) when colorimetric measurements at 8 berry positions showed CIE L* 32.4 (SD ±2.8), a* 35.2 (SD ±3.6), hue angle 8.4° (SD ±4.2°), anthocyanin spatial variation CV 16%, skin anthocyanins averaging 945 mg/kg across positions (range 835-1055 mg/kg), berry sugar 23.1° Brix, firmness 1.6 N, under typical continental conditions (daily high 24-27°C, night 11-14°C) with balanced canopy providing moderate cluster exposure.
Even_Berry_Coloration represents the state of consistent pigmentation across the entire berry surface when color intensity, hue, and saturation show minimal spatial variation creating visually uniform appearance without patches, gradients, or discontinuities that characterize earlier developmental stages. This condition emerges during late veraison to early ripening (BBCH 85-87), achieved 8-12 days post-color break in most cultivars, and is quantified by colorimetric measurements showing spatial coefficients of variation 50-70% during color break), regulatory factors (VvMYBA1/A2, VvMYC1, ABFs) showing consistent expression and activity across berry surface driven by uniform developmental signals including ABA (1800-3000 ng/g with 10-15°C create synthesis rate differences maintaining color variation. Cultivar variation in even coloration characteristics shows: varieties with naturally uniform development (Pinot Noir, Merlot) achieve even coloration showing 92-96% for even coloration), and anthocyanin extraction from berry surface sections (usually 6-8 sections) revealing quantitative spatial variation. Quality implications of even coloration inclu even color indicates complete ripening across entire berry enabling predictable and efficient extraction during winemaking, uneven coloration suggests maturity heterogeneity potentially requiring extended maceration or enzymatic treatment to achieve adequate extraction from less pigmented zones, color evenness correlates with phenolic maturity uniformity (r² = 0.70-0.85) suggesting berries with even anthocyanin distribution also show consistent tannin and other phenolic development, and visual evenness assessment provides rapid quality screening enabling harvest lot selection based on apparent maturity uniformity. Physiological basis of even coloration achievement includes: synchronized berry development across all surface regions through coordinated hormonal signaling, adequate carbon supply enabling anthocyanin synthesis in all zones without source limitation favoring particular regions, and completion of developmental programs throughout berry surface rather than spatial progression from leading to lagging zones. Viticultural factors affecting evenness inclu canopy management creating uniform cluster light exposure (leaf removal timing, shoot positioning, cluster thinning) improves evenness 30-50%, irrigation management maintaining consistent vine water status avoids stress-induced spatial variation in anthocyanin synthesis, crop load optimization ensures adequate resource availability for uniform berry development across all fruit, and disease prevention avoids localized metabolic disruption from pathogen infection that creates color heterogeneity. Temporal stability of even coloration shows: once achieved, even coloration typically persists 2-4 weeks under favorable conditions enabling flexible harvest timing, environmental stresses (heat waves, water deficit, disease pressure) can disrupt evenness within days through spatially variable stress responses requiring careful monitoring, and cultivar variation exists with stable varieties (Cabernet Sauvignon, Syrah) maintaining evenness 3-5 weeks while unstable types (Pinot Noir in warm climates) may lose uniformity within 1-2 weeks through differential senescence or degradation. Integration with other maturity parameters: even coloration typically achieved when berry sugar uniformity also improves (within-berry Brix CV declining from 15-25% to <8-12%), firmness becomes consistent across berry surface, and flavor/aroma development shows reduced spatial variation all indicating coordinated ripening completion approaching optimal harvest.
Even_Berry_Coloration is distinguished from Uniform_Color_Development by representing the achieved state rather than the developmental process, from Color_Distribution_Uniformity by focusing on individual berry evenness rather than cluster/vineyard uniformity, and from Maximum_Color_Intensity by emphasizing spatial consistency rather than peak pigmentation. Evenness criteria: anthocyanin CV <12-18%, CIE L* CV <8-12%, hue variation <6-10°. Typical timing: 8-12 days post-color break. Temperature uniformity and light distribution critical. Even_Berry_Coloration indicates maturity suitable for harvest quality assessment and efficient winemaking extraction.
consistent color distribution
homogeneous berry appearance
homogenes Beeren-Aussehen
konsistente Farb-Verteilung
räumlich konsistente Beeren-Farbe
spatially consistent berry color
uniform surface pigmentation
uniforme Oberflächen-Pigmentierung
Keller M (2020) The Science of Grapevines 3rd Ed pages 235-237 berry color uniformity Rolle L, Torchio F, Giacosa S, Gerbi V (2009) Modifications of mechanical characteristics and phenolic composition in berry skins and seeds of Mondeuse winegrapes throughout the on-vine drying process Amer J Enol Vitic 60:418-425 spatial color distribution Guidoni S, Ferrandino A, Novello V (2008) Effects of seasonal and agronomical practices on skin anthocyanin profile of Nebbiolo grapes Amer J Enol Vitic 59:22-29 cultivar variation in color evenness.
Even Berry Coloration
Juhfark accession Vas 1847Mtp3 at INRAE Domaine de Vassal in Marseillan, France bearing an ex situ conservation role preserves this rare Hungarian cultivar — originally from Somló in western Hungary with fewer than 60 hectares remaining in situ — as a living field accession of 5 vines maintained 2,000 km from its homeland, complemented by cryopreserved shoot tips stored at -196°C at INRAE Montpellier providing redundant backup, with the accession authenticated through SSR comparison against Hungarian reference material at NAIK Budapest confirming genetic identity despite 40+ years of cultivation under Mediterranean rather than continental Pannonian conditions.
A conservation role that inheres in a grapevine maintained outside its original or traditional growing area in a dedicated conservation facility such as a germplasm repository, botanical garden, cryopreservation bank, or institutional collection vineyard, where the primary purpose of the planting is long-term genetic preservation under controlled curatorial conditions rather than commercial production or in-field conservation.
Ex_Situ_Conservation_Role represents the institutional approach to genetic resource preservation that removes material from its environment of origin to secure it under professional curatorial management. The strategy provides protection against catastrophic in-field losses (disease epidemics, land-use change, abandonment) but sacrifices the ongoing adaptive evolution that occurs under in situ conservation: ex situ accessions are genetically frozen at the moment of collection, while in situ populations continue to accumulate adaptive mutations and respond to selection pressures. Modern best practice therefore advocates complementary ex situ and in situ conservation strategies (the "complementary conservation" model) to capture both genetic security and ongoing evolutionary potential.
Ex-situ-Erhaltungs-Rolle
Genbank-Erhaltungs-Rolle
collection-based conservation role
genebank conservation role
off-site conservation role
standortferne Konservierungs-Rolle
Lacombe T et al. (2011) Inventory and characterization of Vitis vinifera ssp. vinifera at INRA Domaine de Vassal, Vitis 50: 56-68 Engelmann F (2011) Cryopreservation of embryos: an overview, Methods in Molecular Biology 710: 155-184 FAO (2010) Second Report on the State of the World's Plant Genetic Resources for Food and Agriculture, Rome
Ex Situ Conservation Role
Cabernet Sauvignon exocarp (strict definition= cuticle plus epidermis) at maturity comprises cuticle 4.5-6.5 μm thick with 165 μg/cm² wax content (42% alkanes, 28% primary alcohols, 16% esters) and epidermis 24-32 μm thick with cells 35-55 μm tangential width bearing 95 stomata and 12 lenticels per mm², creating total exocarp 28-38 μm thickness providing water vapor permeability 0.9×10⁻⁹ m s⁻¹ limiting berry transpiration broader definition including hypodermis (2-3 layers, 42-55 μm) creates total exocarp 70-93 μm containing 3.2-4.2 mg/g anthocyanins providing deep purple-black color. Chardonnay white cultivar exocarp showing white wine characteristics= cuticle 2.8-4.2 μm with 95 μg/cm² wax (lower than typical red cultivars), epidermis 18-28 μm, hypodermis (when included in exocarp definition) 24-38 μm lacking anthocyanins but containing 1.1-1.8 mg/g flavonols and carotenoids creating golden-yellow color, demonstrating exocarp structure in white cultivars with different pigment composition than red cultivars. Syrah exocarp cuticle ultrastructure examined by transmission electron microscopy showing three-layer organization= epicuticular wax layer (outer surface) with crystalloid projections 0.8-1.5 μm height creating bloom cutin matrix layer (middle) 3-4.5 μm thick electron-dense polyester network cuticular layer (inner) 0.6-1.2 μm interfacing with epidermal cell wall, with total cuticle 4.5-6.8 μm providing primary barrier to water loss and pathogen entry. Pinot Noir exocarp light exposure effects= sun-exposed clusters showing exocarp with thicker cuticle (5.5-7.5 μm versus 3.2-4.8 μm shaded), higher wax content (190 μg/cm² versus 105 μg/cm²), higher flavonols in epidermis/hypodermis (1.8-2.6 mg/g versus 0.6-1.1 mg/g), demonstrating sunlight induction of exocarp protective barriers and UV-screening compounds important for berry protection and potentially affecting wine phenolic composition. Merlot exocarp developmental changes tracked from fruit set through maturity= fruit set (BBCH 71) with cuticle 1.2 μm, epidermis 18 μm, green chlorophyll-containing mid-development (BBCH 75, week 6) with cuticle 2.8 μm, epidermis 22 μm, still green veraison (BBCH 81, week 10) with cuticle 4.2 μm, epidermis 26 μm, hypodermis showing beginning anthocyanin synthesis (0.3 mg/g) maturity (BBCH 89, week 18) with cuticle 5.5 μm, epidermis 28 μm, hypodermis with 3.2 mg/g anthocyanins, demonstrating progressive exocarp development and maturation creating final protective and pigmented outer berry layer.
An exocarp is the outermost tissue layer of the fruit pericarp comprising the epidermis and cuticle (and sometimes including hypodermis in broader definitions), representing the external protective boundary of the berry in direct contact with the environment, synonymous with or closely related to outer pericarp in grape berry anatomy where exocarp typically refers to the 15-60 μm thick epidermal-cuticular layer or the 40-180 μm thick skin region including hypodermal layers, serving critical barrier functions against water loss, pathogen invasion, mechanical damage, and UV radiation while bearing specialized surface structures and containing phenolic compounds important for berry protection and wine quality. Exocarp in strict anatomical definition comprises= (1) cuticle (outermost acellular layer) 2-8 μm thick at maturity, composed of cutin matrix (polyester of C16-C18 hydroxy fatty acids providing structural framework), embedded and surface waxes (aliphatic compounds including alkanes, alcohols, esters creating water barrier), and epicuticular wax crystalloids (projecting 0.5-2 μm from surface creating bloom) (2) epidermis (single outermost cell layer) 15-35 μm radial thickness, consisting of small tightly-packed cells with heavily modified outer periclinal walls (3-6 μm thick, cutinized and suberized), bearing stomata (50-200 per mm²) and lenticels (5-20 per mm²). Exocarp in broader viticultural usage often includes hypodermis (1-3 sub-epidermal cell layers, 20-60 μm total thickness) creating total exocarp thickness of 40-90 μm, corresponding to outer pericarp or berry skin, with hypodermis containing anthocyanins in red/black cultivars providing berry color and wine color potential. Exocarp functional roles include= (1) water barrier reducing transpiration through low cuticle permeability (10⁻⁸ to 10⁻⁹ m s⁻¹) critical for preventing berry dehydration during ripening and post-harvest (2) pathogen barrier providing physical resistance to fungal and bacterial penetration through intact cuticle and epidermal walls, with cuticle waxes showing antimicrobial properties and wounded exocarp inducing defense responses including phytoalexin synthesis (resveratrol, pterostilbene) (3) UV protection through flavonols and anthocyanins in epidermal and hypodermal cells absorbing UV-B radiation (280-315 nm) preventing photodamage to internal tissues (4) mechanical protection through structural integrity of thick cuticle and epidermal walls resisting compression, puncture, and abrasion during berry development, harvest, and handling (5) gas exchange through stomata (primarily during green berry phase) and lenticels supporting respiration and, during early development, photosynthesis in green chlorophyll-containing exocarp. Exocarp composition shows cultivar and environmental variation= cuticle wax content 50-250 μg/cm² berry surface with sun-exposed berries accumulating more wax than shaded berries epidermal and hypodermal phenolic content including anthocyanins (0-6+ mg/g in red/black cultivars, essentially zero in white cultivars), flavonols (0.5-2.5 mg/g higher in sun-exposed), hydroxycinnamic acids, and tannins (6-12 mg/g in outer layers) cuticle permeability varying with wax composition and thickness affecting berry susceptibility to splitting, shriveling, and post-harvest water loss. Exocarp development follows berry development= cuticle deposition beginning at fruit set and continuing through ripening with thickness increasing from 1-2 μm to 2-8 μm epidermal cells undergoing limited expansion (15-25% diameter increase) compared to pericarp bulk creating progressive thinning relative to berry stomata forming during early development and becoming largely non-functional during ripening anthocyanin synthesis (red cultivars) initiating at veraison and continuing through ripening wax composition changing during development with increasing proportion of very-long-chain aliphatic compounds during ripening.
Exocarp represents the outermost fruit layer and is essentially synonymous with Outer_Pericarp (when broadly defined to include hypodermis) or Epidermal_Outer_Pericarp (when strictly defined as cuticle plus epidermis only). The term exocarp emphasizes the protective barrier function and external position. Sub-classification into Berry_Skin (functional/viticultural term) and Cuticle_Layer (anatomical component) reflects different perspectives on the outermost tissues. Understanding exocarp structure and function is essential for berry protection, post-harvest handling, pathogen resistance, and for managing phenolic development in outer tissues affecting wine quality.
Außenschicht
Exokarp
berry surface layer
outer fruit wall
äußerste Fruchtwand
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 176-182 Rosenquist, J.K., Morrison, J.C. (1989). Factors affecting cuticle and wax accumulation. American Journal of Enology and Viticulture 40=241-244 Rogiers, S.Y. et al. (2004). Grape berry epicuticular wax and transpiration. American Journal of Enology and Viticulture 55=121-127.
Exocarp
Pinot Noir expanding mid berries undergoing veraison (BBCH 81-83, weeks 9-11 post-fruit set) show progression from green berries at 9.5 mm diameter, 1.1 g fresh weight, firm texture (4.2 N), 8% sugars, 23 g/L acidity, no anthocyanins (end of lag phase BBCH 77) through color change initiation with chlorophyll decline and beginning anthocyanin synthesis (week 9, 0.15 mg/g skin anthocyanins, berries showing pink coloration at stylar end) to mid-veraison with 50% color change (week 10, 10.5 mm, 1.35 g, 2.8 N, 14% sugars, 18 g/L acidity, 1.2 mg/g anthocyanins, berries red-purple at stylar end transitioning to pink toward pedicel end) to late veraison (week 11, 11.5 mm, 1.5 g, 2.2 N, 16% sugars, 16 g/L acidity, 1.8 mg/g anthocyanins, berries uniformly colored purple). Syrah expanding mid berries examined during veraison show cell expansion kinetics= lag phase cell diameter 52 μm (BBCH 77, week 8), beginning expansion 62 μm (early veraison BBCH 81, week 9), rapid expansion 85 μm (mid-veraison BBCH 83, week 10.5), with cell expansion from 52 to 85 μm representing 64% diameter increase (135% volume increase) driving berry diameter increase from 10 mm to 11.8 mm (18% diameter increase, 64% volume increase), demonstrating that veraison berry expansion results from cell enlargement of the fixed cell population established during Phase I. Chardonnay white cultivar expanding mid berries show veraison marked by color change from green to golden-yellow through chlorophyll degradation (from 0.6 mg/g to <0.1 mg/g during weeks 9-11) and carotenoid/flavonol accumulation creating yellow pigmentation, softening from 4.5 N to 2.5 N, sugar increase from 9% to 16%, acid decline from 19 g/L to 14 g/L, demonstrating veraison transitions comparable to red cultivars despite subtle color change. Merlot experiencing veraison at different temperatures shows 22-25°C optimal conditions producing rapid uniform color development (90% colored berries by BBCH 83, 1.9-2.3 mg/g skin anthocyanins) versus 30-34°C heat stress showing delayed irregular color development (60% colored, 1.1-1.6 mg/g anthocyanins) and slower sugar accumulation (13% versus 15% at comparable developmental time), demonstrating critical temperature sensitivity of veraison processes affecting color and composition development.
An expanding mid berry is a mid developing berry undergoing veraison and early ripening (BBCH 81-83, approximately 8-11 weeks post-fruit set) with onset of berry expansion through cell enlargement, color change, softening, and beginning of sugar accumulation and acid decline, representing the critical developmental transition from vegetative to reproductive berry physiology marking the start of Phase II ripening growth. Expanding mid berries are characterized by= (1) beginning berry expansion as cells initiate enlargement from lag phase size (45-70 μm diameter) toward mature size (100-180 μm), with berry diameter increasing 8-20% during the veraison period (BBCH 81-83) from lag phase dimensions (2) fresh weight increasing 15-35% during veraison as cells expand and water/solute accumulation begins (3) color change as the defining visible veraison characteristic= chlorophyll degradation eliminating green color anthocyanin synthesis in red/black cultivars creating progressive color development from green to pink to red to purple carotenoid and flavonol changes in white cultivars creating golden-yellow coloration color change typically proceeding from stylar (blossom) end toward pedicel (stem) end creating transient bicolored appearance during mid-veraison (4) beginning softening as cell walls undergo modifications including pectin solubilization and hemicellulose depolymerization, reducing firmness from lag phase values of 4-6 N to veraison values of 2.5-4 N (measured by compression) (5) sugar accumulation accelerating from lag phase rates of 0.1-0.3% per week to veraison rates of 0.8-1.5% per week, increasing from lag phase levels of 7-10% to veraison levels of 12-17% by BBCH 83 (6) titratable acidity beginning decline from peak levels of 20-25 g/L to 15-20 g/L by BBCH 83 through malic acid respiration and dilution effects from berry expansion. Expanding mid berries undergo coordinated developmental changes= cell wall modifications initiating softening and allowing cell expansion osmotic potential declining through sugar and solute accumulation creating driving force for water import and cell expansion vascular function shifting from predominant xylem to predominant phloem import as berry transitions to strong sink secondary metabolite biosynthesis activating with anthocyanin, tannin, and aromatic compound synthesis beginning acid metabolism shifting from synthesis to degradation with malic acid respiration activated hormone balance showing ABA increases, auxin decreases, and ethylene beginning to increase. Expanding mid berries show rapidly changing composition= sugars increasing at approximately 1% per week during active veraison (versus 0.1-0.3% per week during lag phase) titratable acidity declining at 0.5-1.5 g/L per week through malic acid respiration (malate decarboxylation) and dilution anthocyanins (red cultivars) accumulating from essentially zero (32°C inhibiting anthocyanin accumulation light exposure enhances anthocyanin synthesis and secondary metabolite accumulation in exposed berries versus shaded berries water availability affects berry expansion with adequate water supporting cell enlargement versus deficit reducing expansion and concentrating composition.
Expanding_Mid_Berry represents veraison and early ripening (BBCH 81-83) and is distinguished from Active_Growth_Mid_Berry (lag phase, minimal expansion) by beginning expansion, color change, and ripening versus static dimensions and green color. Expanding mid berries undergo the most visible and most dramatic berry developmental transition (veraison) marking the shift from vegetative to reproductive berry physiology. Understanding expanding mid berry biology is essential for harvest timing prediction, quality management through cultural practices affecting veraison conditions, and for interpreting compositional development determining wine/grape quality.
Farbumschlags-Stadium
Véraison-Beere
color-change berry
expandierende Mittel-Beere
veraison berry
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 196-212 Coombe, B.G., McCarthy, M.G. (2000). Dynamics of grape berry growth. Australian Journal of Grape and Wine Research 6=131-135 Castellarin, S.D. et al. (2007). Transcriptional regulation of anthocyanin biosynthesis. Plant Physiology 143=1347-1363.
Expanding Mid Berry
An expanding young leaf at node 10 on a shoot of Vitis vinifera cv. Merlot in Bordeaux, France, measuring 8.5 cm length by 7.8 cm width representing approximately 55% of expected mature dimensions based on position and cultivar norms, exhibiting flat expanded lamina with clearly defined five-lobed architecture and moderate-depth sinuses, medium green coloration on adaxial surface with reduced bronze tinge compared to younger stages indicating increasing chlorophyll content measured at 340 μg cm⁻², residual anthocyanin particularly along veins and margins at 1.8 mg g⁻¹ fresh weight, moderately soft texture with cells averaging 28 μm diameter in palisade approaching but not achieving mature size of 35 μm, two distinct palisade cell layers totaling 65 μm thickness (compared to mature 85 μm) and developing spongy mesophyll of 95 μm (compared to mature 120 μm), functional stomata with density of 165 stomata mm⁻² showing appropriate closure response to vapor pressure deficit increase though with reduced sensitivity, partial cuticle development creating moderate surface sheen, demonstrating photosynthesis rate of 8.7 μmol CO₂ m⁻² s⁻¹ representing 68% of mature leaf capacity and approaching carbon balance with daily photosynthetic gain of 42 mg carbohydrate approximately equaling growth respiration costs, occurring at 16 days post-emergence during active expansion phase, measured at shoot BBCH stage 71 (fruit set) in mid-June, representing mid-expansion stage with continued growth expected over subsequent 10-14 days before achieving mature dimensions.
A young leaf in active growth phases after initial unfoldment dominated by cell expansion processes in Vitis vinifera, characterized by intermediate size typically 3-12 cm length representing 20-70% of final mature dimensions, essentially flat lamina with completed basic unfoldment though continued area increase through cell enlargement, light to medium green coloration as chlorophyll concentrations increase though still below mature levels typically 250-400 μg cm⁻² depending on expansion stage, reduced but still significant anthocyanin content providing partial photoprotection, soft to moderately firm texture as cell walls thicken but without complete lignification, ongoing tissue differentiation with palisade and spongy mesophyll layers becoming distinct though not fully mature, developing vascular system with progressive secondary wall deposition in xylem and phloem maturation, functional but incompletely developed stomata showing responses to environmental signals though with reduced sensitivity compared to mature stomata, incomplete cuticle development with partial wax deposition, and increasing photosynthetic capacity typically 40-80% of mature leaf rates transitioning from net carbon sink early in expansion to approaching carbon balance or slight net source late in expansion depending on environmental conditions and resource availability. Expanding young leaves represent the major growth phase accounting for most final size achievement through cell expansion, with cell division largely complete by expansion initiation and final leaf area determined primarily by the extent of cell enlargement driven by water uptake, vacuole expansion, and wall loosening, requiring high resource availability particularly water for turgor-driven expansion and nitrogen for continued protein synthesis supporting membrane and enzyme production, with the expansion phase typically spanning 14-21 days under optimal conditions but extending to 25-35 days under stress creating smaller final leaf size.
Expanding young leaves represent the primary growth phase following initial unfoldment, distinguished from emerging leaves by substantially larger size and flat architecture, and from mature leaves by incomplete size achievement and tissue development with continued active growth. Cell expansion driving leaf enlargement depends critically on water availability, with turgor pressure providing the physical force for cell wall extension, making this phase extremely sensitive to water deficit that reduces final leaf size proportionally to stress severity and duration. Light environment during expansion determines final anatomical characteristics, with high light promoting development of multiple palisade layers and dense tissue creating sun leaf characteristics, while low light produces single palisade layers and thin tissue creating shade leaf characteristics. Temperature affects expansion rate but not typically final size, with optimal temperatures of 22-28°C supporting rapid expansion completing in 14-21 days while cooler temperatures extend expansion duration without necessarily reducing final dimensions. Nutrient availability particularly nitrogen affects both expansion rate and final size, with deficiency reducing cell expansion and creating smaller mature leaves.
actively enlarging leaf
aktiv vergrößerndes Weinblatt
expanding leaf
growth phase leaf
sich ausdehnendes junges Blatt
wachstumsphase Blatt
Based on Poni et al. (2009) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 60=397-407 Palliotti et al. (2012) "Morpho-anatomical and physiological response of Pinot Noir and Montepulciano grapevine leaves to mild and severe water stress" Vitis 51=109-116 Tsukaya (2013) "Leaf development" The Arabidopsis Book 11=e0163 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 103-136.
Expanding Young Leaf
Chardonnay in Champagne shows expansion rate decline during early-mid August Year N+1 at BBCH 77-79, with berry growth rate declining from peak 128 mg per day to 84 mg per day early decline to 38 mg per day late decline representing 70% total reduction, wall yield threshold increasing from 0.48 to 0.74 MPa, osmotic potential plateau at -1.3 MPa, expansin expression downregulated 65%, under cool maritime conditions of 20-26°C temperature, typical of this early-bursting cultivar transitioning to lag phase
Expansion rate decline represents the growth deceleration phase during maximum berry enlargement when berry fresh weight gain decreases from peak 100-250 mg per day to declining 30-80 mg per day, expansion velocity progressively reduces as cells approach size limits, growth rate deceleration results from reduced wall yielding and osmotic plateau, and declining rate marks transition from intensive enlargement to lag phase minimal growth, occurring over weeks 6-8 as growth phase concludes. This deceleration phase occurs during late summer in early-mid August in temperate regions Year N+1, proceeding over 1-2 weeks representing rate decline period, corresponding to BBCH 77-79 bunch closure stages. Rate decline mechanism involves reduced wall plasticity limiting turgor-driven expansion, osmotic potential plateau reducing water uptake driving force, cells approaching genetic size limits constraining further growth, developmental programming directing growth cessation, and integrated factors producing progressive rate deceleration. Growth rate quantification shows decline from peak 100-250 mg per day to 60-120 mg per day early decline to 30-80 mg per day late decline, progressive rate reduction over 1-2 weeks, eventual very low rate <20-30 mg per day entering lag phase, and rate trajectory characteristic of growth cessation. Rate decline kinetics show approximately 40-60% rate reduction from peak to early decline, additional 30-50% reduction from early to late decline, cumulative 70-85% total rate reduction during decline phase, and substantial deceleration marking growth phase end. Wall plasticity reduction shows wall extensibility declining as structural modifications accumulate, reduced expansin expression and activity limiting wall loosening, increased wall cross-linking strengthening structure, yield threshold increasing from 0.4-0.6 MPa to 0.6-0.9 MPa, and wall properties shifted toward growth-restrictive state. Expansin downregulation shows gene expression declining during rate decline, protein levels decreasing in expanding tissues, enzyme activity reduction limiting wall modification, and expansin decline as growth cessation mechanism. Wall cross-linking increase shows enhanced pectin cross-linking through calcium-mediated interactions, increased hemicellulose-cellulose associations, structural modifications strengthening walls, and cross-linking limiting wall yielding under turgor. Osmotic potential plateau shows osmotic potential approaching steady state at -1.2 to -1.6 MPa, minimal further solute accumulation, organic acids at peak or beginning decline 1.2-1.6%, potassium approaching maximum 80-140 mM, and plateau limiting additional water uptake driving force. Solute accumulation deceleration shows organic acid synthesis slowing or reversing, malate and tartrate at peak levels with minimal additional accumulation, potassium import approaching maximum, sugar accumulation still limited 6-10%, and overall solute dynamics approaching equilibrium. Water uptake reduction shows water influx declining from peak 1.5-2.5 g per day to 0.8-1.5 g per day during decline, reduced uptake from osmotic plateau and reduced expansion, water import still significant but decreasing, and uptake decline contributing to rate reduction. Turgor maintenance shows pressure sustained 0.6-1.0 MPa during decline, turgor maintaining adequate levels, but expansion minimal from wall constraints, turgor-expansion coupling reduced with high pressure producing minimal growth, and pressure alone insufficient for expansion without wall yielding. Cell size constraints show cells approaching genetic and physical size limits, further volume increase increasingly difficult, possible mechanosensing signaling size achievement, and intrinsic constraints limiting expansion. Developmental signals show possible growth cessation signals from developmental programming, transcriptional changes directing metabolic shift from growth to lag phase, hormonal changes possibly affecting growth rate, and integrated developmental control directing decline. Berry growth rate decline shows berry fresh weight gain following cell expansion rate decline, berry volumetric increase slowing proportionally, berry diameter increase declining from 1-2 mm per week to 0.3-0.7 mm per week, and berry growth deceleration mirroring cellular rate decline. Berry composition during decline shows water content maintained 80-85%, organic acids at peak or slightly declining 1.2-1.6%, sugars beginning accumulation 6-10% still relatively low, pH 2.8-3.0, and composition transitioning toward lag phase characteristics. Metabolic changes during decline show respiratory rate beginning decrease from peak levels, metabolic shift from growth-supporting to maintenance metabolism, early metabolic reorganization toward lag phase, and declining metabolism correlating with growth rate. Gene expression changes show growth-related genes downregulating, lag phase-associated genes beginning upregulation, transcriptional reprogramming initiating metabolic transition, and expression changes accompanying rate decline. Environmental effects on rate decline include temperature affecting decline kinetics with warm conditions potentially accelerating transition, water availability less critical during decline than peak expansion though continued supply beneficial, and environmental factors having reduced impact as genetic programming dominates. Temperature effects show warm conditions promoting faster transition through decline phase, cool conditions extending decline period, and temperature affecting transition timing. Water effects show adequate water beneficial for maximizing final size, deficit during decline reducing final 5-10%, and water less limiting than during peak expansion. Rate decline assessment shows serial growth measurements quantifying deceleration, growth curve analysis documenting rate trajectory, molecular marker changes showing metabolic transition, and measurements enabling decline characterization. The process represents third and final aspect of maximum berry enlargement emphasizing growth deceleration, following cell size maximum and berry size peak. Duration typically 1-2 weeks representing progressive rate decline weeks 6-8. Regional timing shows expansion rate decline during early-mid August across climates as growth phase concludes. Viticultural significance includes representing transition from intensive growth to lag phase, marking growth phase completion, demonstrating developmental control of growth cessation, and establishing rate decline as indicator of approaching lag phase. Research applications include analyzing rate decline mechanisms and regulation, investigating wall modification and osmotic plateau effects, evaluating developmental signals directing cessation, quantifying decline kinetics and trajectories, and assessing rate decline as transition to lag phase.
Expansion_Rate_Decline is third and final aspect of Maximum_Berry_Enlargement, distinguished by progressive growth deceleration. Represents 70-85% total rate reduction from peak 100-250 to 30-80 mg/day. Mechanisms: reduced wall plasticity (yield threshold 0.4-0.6 → 0.6-0.9 MPa), osmotic plateau (-1.2 to -1.6 MPa), size constraints. Expansin downregulation. Duration 1-2 weeks. Follows Cell_Size_Maximum and Berry_Size_Peak. Completes Maximum_Berry_Enlargement. Marks transition to Enlargement_Phase_Completion and lag phase.
Expansions-Verlangsamung
Expansionsraten-Abnahme
Ratenreduktion
Wachstums-Dezeleration
declining growth velocity
expansion slowdown
growth deceleration
rate reduction
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 450-453 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Expansion Rate Decline
A Pinot Noir selection identified in an Oregon research vineyard bears an Experimental_Clone_Role, undergoing initial evaluation trials to assess whether its distinctive characteristics warrant advancement through the 15-year process toward potential registration. A virus-resistant breeding line at INRA serves in an Experimental_Clone_Role, with introgressed resistance genes from wild Vitis species under evaluation for commercial potential but not yet approved for market release. An ancient variety accession in an ENTAV conservatory collection bears an Experimental_Clone_Role for germplasm preservation, maintained for genetic conservation research rather than commercial propagation.
A clone role that inheres in a grapevine clone maintained in experimental or research status, not yet registered for commercial propagation but under evaluation, in development, or retained for research purposes without commercialization intent. Experimental clones occupy the pipeline between initial selection and official registration, undergoing the multi-year evaluation trials (typically 10-20 years) required before registration approval, or may be maintained indefinitely for research without advancement toward commercial release. The experimental designation encompasses diverse situations—promising new selections in active evaluation, material of scientific interest without commercial potential, breeding lines under development, reference collections for research comparison, and selections withdrawn from commercial consideration but retained for genetic conservation. Experimental clones typically exist in limited quantities at research institutions, unavailable to commercial nurseries, with access restricted to research collaborators or requiring special authorization. The extended timeline from initial selection through registration means that experimental status may persist for decades before resolution into either registered commercial release or permanent research-only status.
Experimental_Clone_Role is subclassed by specific experimental function—Breeding_Line (in active development toward commercialization) and Research_Clone (maintained primarily for research purposes)—recognizing the diverse situations encompassed by non-commercial clone status. The experimental pipeline represents substantial investment advancing a selection through registration typically requires 10-20 years of trials and evaluation before commercial availability. Many experimental selections never achieve registration, either failing evaluation criteria or serving research purposes that don't require commercial release. Access to experimental material typically requires research collaboration agreements, with institutions protecting unreleased selections from unauthorized distribution. The experimental category also includes material under intellectual property protection awaiting licensing arrangements before commercial release.
Evaluierungsklon
Prüfungsklon
Versuchsklon
evaluation clone
experimental clone
experimenteller Klon
pre-registration clone
trial clone
Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Lacombe T et al. (2011) Grapevine genetic diversity, in Bentley R (ed) The Handbook of Grape Processing, Blackwell
Experimental Clone Role
The experimental code VB 91-26-17 assigned by INRAE Colmar identifies a downy-mildew-resistant genotype in advanced evaluation trials across 4 French sites since 2005, with the code appearing in all trial plot maps, harvest data sheets, micro-vinification records, and sensory evaluation panels — upon successful completion of DUS (Distinctness, Uniformity, Stability) testing and CTPS evaluation, the genotype received the cultivar name Floreal in 2018 for official registration, but the experimental code VB 91-26-17 remains the primary identifier in all research publications and breeding database records linking the released cultivar to its complete experimental history.
A breeding code assigned to a grapevine genotype that has advanced beyond initial seedling screening into replicated experimental field trials, serving as the working identifier in trial design documentation, data collection systems, and research communications throughout the multi-year evaluation phase.
Experimental_Code identifies genotypes that have survived initial screening (typically eliminating 85-95% of seedlings) and entered formal replicated trials. The code serves as the stable identifier across multiple years, sites, and evaluation parameters — maintaining data integrity through the decade-long evaluation process during which a genotype may be assessed for 50+ agronomic, quality, and resistance traits: the experimental code therefore functions as the primary key in breeding databases linking phenotypic, genotypic, and oenological data accumulated over the genotype's evaluation history. The code persists in the scientific literature even after cultivar naming, enabling traceability from published cultivar to its complete pre-release experimental record.
Experimentalkennung
Feldversuchs-Nummer
Versuchscode
experimental genotype code
field trial identifier
trial code
Eibach R and Töpfer R (2015) Traditional grapevine breeding techniques, in Reynolds AG (ed) Grapevine Breeding Programs for the Wine Industry, Woodhead Publishing pp: 3-22 Schneider C et al. (2019) Grapevine breeding programs in France, in Ferrão MAG et al. (eds) Genetic Improvement of Tropical Crops, Springer pp: 301-344
Experimental Code
PIWI breeding selection Gf.2001-050-024 at the Julius Kühn-Institut in Siebeldingen bearing an experimental cultivar role is planted in replicated randomized complete block trials with 4 blocks × 20 vines per plot, undergoing evaluation in its 8th leaf for downy mildew resistance (scored weekly on OIV 452 scale), powdery mildew resistance (OIV 455), yield components (cluster number, cluster weight, berry weight), must quality parameters (Brix, pH, titratable acidity, YAN), and wine sensory quality through micro-vinification of 50 kg lots evaluated by a trained 12-member panel using the DLG 5-point scoring system.
A research and education role that inheres in a grapevine that is the active subject of scientific investigation, planted within formally designed experimental trials for the purpose of generating new knowledge about cultivar performance, trait expression, management responses, or environmental adaptation under controlled or semi-controlled conditions.
Experimental_Cultivar_Role captures the epistemic status of vines serving as research subjects within hypothesis-driven scientific investigation. The role implies formal experimental design — replication, randomization, and systematic data collection — distinguishing experimental vines from informally observed plantings: this experimental framework is essential for generating statistically valid conclusions about genotype performance and trait expression. Subclasses distinguish between trial cultivars (undergoing initial screening and elimination) and evaluation cultivars (in advanced multi-site assessment approaching potential release).
Forschungsobjekt-Rolle
Versuchssorten-Rolle
experimental variety role
experimentelle Rebsorten-Rolle
research subject role
trial genotype role
Eibach R and Töpfer R (2015) Traditional grapevine breeding techniques, in Reynolds AG (ed) Grapevine Breeding Programs for the Wine Industry, Woodhead Publishing pp: 3-22 Töpfer R et al. (2011) New horizons for grapevine breeding, Fruit, Vegetable and Cereal Science and Biotechnology 5: 79-100 OIV (2009) Guidelines for the conduct of tests for distinctness, uniformity and stability of table and wine grape varieties
Experimental Cultivar Role
Export certification for Austrian wine shipments to Japan requires: a VI-1 document (the EU wine attestation form confirming analytical composition and compliance with EU production regulations, issued by the Austrian Bundeskellereiinspektion after chemical analysis verification), a certificate of origin (for PDO/PGI wines, confirming geographic authenticity), Japanese import permit documentation (including Japanese-language label compliance verification), and analytical certificates confirming compliance with Japanese maximum residue limits for pesticides and maximum limits for specific additives — the certification package reflecting both EU export requirements and Japanese import specifications. Export certification for grapevine cuttings shipped from the ENTAV-INRA nursery in France to a Chilean vineyard requires a phytosanitary certificate issued by the French DRAAF (Direction Régionale de l'Alimentation, de l'Agriculture et de la Forêt) confirming the material is free from regulated quarantine pests, supplemented by laboratory test results for specified viruses (ELISA certificates for GLRaV-1, -3, GFkV, GFLV, ArMV), and a varietal authenticity certificate confirming cultivar and clone identity through microsatellite verification.
Export certification is a trade regulation entity that documents the official attestation requirements that must be fulfilled before grape plant material or wine products may be legally exported from their country of origin, encompassing phytosanitary certificates for plant material (issued by the National Plant Protection Organization confirming compliance with the importing country's pest and disease requirements), wine export certificates (issued by designated inspection bodies confirming compliance with production standards and analytical specifications), certificates of origin (authenticating the geographic provenance of wine for appellation-protected products), and varietal authenticity documentation (confirming cultivar identity through molecular or ampelographic verification where required), specifying the issuing authority, required documentation, analytical testing, inspection procedures, and the validity period and geographic scope of each certificate type.
Export certification documentation captures the administrative gatekeeping function that ensures viticultural exports meet both the originating country's export standards and the destination country's import requirements — a dual compliance obligation that creates substantial documentation burdens for exporters but provides essential quality assurance and phytosanitary protection. The phytosanitary certificate for plant material exports follows the IPPC Model Certificate format (ISPM 12), requiring the exporting country's NPPO to inspect the material, verify compliance with the importing country's specified phytosanitary requirements, and issue a standardized certificate that accompanies the consignment — a system that relies on mutual trust between NPPOs and is periodically tested by interceptions of non-compliant material that may trigger enhanced inspection protocols or temporary trade suspensions. The wine export certification system varies by exporting country — the EU employs the VI-1 and VI-2 document system (a standardized attestation form confirming analytical composition and production compliance), Australia uses the Wine Australia Export Label Directory system (requiring pre-export label approval and analytical verification), and the US employs TTB export certificates — each system reflecting the exporting country's regulatory structure and the bilateral agreements governing wine trade with specific destination markets. The cost and administrative burden of export certification represent a significant non-tariff trade barrier, particularly for small producers — obtaining the full documentation package for a single wine shipment may involve multiple government agencies, laboratory analyses, and administrative fees, creating economies of scale that favor large exporters and potentially excluding artisanal producers from international markets. The electronic certification trend (e-phyto for plant health certificates, electronic VI-1 documents) is progressively reducing the administrative burden through digitization of the certification process, though full electronic interoperability between national systems remains incomplete.
Ausfuhrbescheinigung
Exportgenehmigung
Exportzertifizierung
Weinausfuhrdokumentation
export attestation
export compliance certificate
export permit
phytosanitary export certificate
phytosanitäres Exportzertifikat
wine export documentation
FAO (2023) ISPM 12 — Phytosanitary certificates, International Plant Protection Convention European Parliament (2019) Commission Delegated Regulation (EU) 2019/934, Article 28 (VI-1 documents) Wine Australia (2023) Export Label Directory requirements ENTAV-INRA (2023) Certification procedures for grapevine plant material
Export Certification
Crimson Seedless bears an Export_Table_Grape_Role as the dominant red seedless in international trade, with its firm texture surviving 4-6 week sea freight from Chile/Peru to Europe and Asia, excellent storage life (8-12 weeks) providing supply chain flexibility, and globally recognized quality commanding consistent demand across North American, European, and Asian markets. Red Globe serves in an Export_Table_Grape_Role as the preferred variety for Asian export, with exceptional shipping durability, 12-16 week storage capability enabling sea freight from South America and South Africa, and giant berry size (12-20 g) matching Asian market preferences that justify premium pricing despite seeds. Thompson Seedless bears an Export_Table_Grape_Role for volume trade in white seedless, with moderate storage capability (6-10 weeks), global brand recognition, and competitive pricing for markets prioritizing affordability over premium characteristics.
A table grape scion role that inheres in a grafted scion cultivar by virtue of its suitability for international export trade, possessing the combination of shipping durability, storage stability, visual uniformity, and market acceptance required for successful long-distance transport and foreign market penetration. Export table grape scions must satisfy multiple simultaneous requirements including mechanical durability (firm berry attachment, resistant rachis, shatter-resistant clusters) for handling and transit, storage longevity (minimum 4-8 weeks for sea freight, longer for distant markets), phytosanitary compliance (freedom from quarantine pests, tolerance of required treatments such as cold treatment, fumigation, or irradiation), visual quality standards (uniform size, color, shape meeting import market specifications), and eating quality meeting destination market preferences (sweetness, texture, flavor profiles varying by market). Export varieties dominate international trade flows, with a small number of globally accepted cultivars (Thompson Seedless, Crimson Seedless, Red Globe, Flame Seedless, Sugraone) constituting the majority of the approximately 4 million tonnes of table grapes traded internationally.
Export_Table_Grape_Role captures the demanding requirements of international trade where varieties must satisfy shipping logistics, regulatory compliance, and diverse market expectations simultaneously. The concentration of global trade in a few varieties reflects the difficulty of meeting all export requirements—many excellent local varieties lack shipping durability, storage life, or international market recognition for viable export. Counter-seasonal trade (Southern Hemisphere supplying Northern Hemisphere winter) has grown dramatically since the 1980s, now exceeding 2 million tonnes annually from Chile, Peru, South Africa, and Australia. Proprietary varieties (Sweet Celebration, Allison, Sable Seedless) are increasingly important in export, with licensed grower networks in multiple production countries enabling year-round branded supply. Phytosanitary requirements vary by destination—Japan and Korea require cold treatment or methyl bromide fumigation, creating variety-specific trade access limitations.
Exporttafeltraube
Handelstafeltraube
Versandtraube
export grape
international market grape
shipping grape
trade grape
Creasy GL, Creasy LL (2009) Grapes, CABI Crisosto CH, Mitchell FG (2002) Postharvest handling systems: table grapes, UC ANR OIV (2019) Statistical Report on World Vitiviniculture USDA Foreign Agricultural Service (2020) Fresh Deciduous Fruit Annual Reports Fidelibus MW et al. (2017) California Table Grape Production Guidelines
Export Table Grape Role
The exposure description for the Clos Sainte Hune vineyard (Hunawihr, Alsace, 48.18°N) documents a favorably sheltered, south-southeast exposed mid-slope position on the eastern flank of the Vosges foothills — the vineyard benefits from the pronounced Vosges rain shadow effect (Colmar, 15 km south, receives only 550 mm annual rainfall, one of the driest stations in France) that reduces growing season rainfall and disease pressure, while the Vosges ridge to the west (rising to 1,000-1,200 m) blocks the prevailing westerly winds and Atlantic-sourced precipitation systems, and the south-southeast aspect captures full morning and midday sun with the Vosges ridge providing afternoon shade that moderates maximum temperatures. This combined exposure — sheltered from wind and rain by the Vosges, facing the morning sun on a moderate slope with limited horizon obstruction to the east — creates the dry, warm, well-lit mesoclimate that enables Riesling at this northerly latitude to achieve full aromatic maturity while preserving the high acidity that defines the Clos Sainte Hune as one of the world's greatest dry Riesling vineyards.
A site description component that records the overall radiation and atmospheric exposure conditions of the vineyard — integrating the effects of slope aspect and angle, horizon obstruction (topographic shading by surrounding hills, ridgelines, or structures), wind exposure (sheltered versus exposed positions), and proximity to reflective or thermally moderating surfaces (water bodies, rock faces, urban areas) — providing a holistic assessment of the energy and atmospheric environment that determines the radiation budget, temperature extremes, evaporative demand, and disease pressure at the vineyard site.
Exposure_Description synthesizes the multiple environmental factors that determine the energy balance and atmospheric conditions at the vineyard site into an integrated assessment that captures effects not fully described by the individual slope parameters alone. The concept of exposure extends beyond the geometric relationship between slope and sun (captured by aspect and angle) to include the dynamic atmospheric environment — wind exposure affects evapotranspiration, canopy drying speed (influencing disease pressure), mechanical stress on canopy structures, and convective heat transfer that modifies the radiation-driven temperature regime horizon obstruction from surrounding topography reduces total radiation by blocking the sun during early morning and late afternoon hours (significant in narrow valleys where the surrounding ridgelines may obstruct 1-3 hours of potential direct radiation per day) and the proximity of thermally significant landscape features (large water bodies providing thermal buffering, rock walls providing radiation reflection and thermal mass, forests providing wind shelter and humidity modification) creates exposure-specific microclimates that deviate from the regional norm: the interaction of these factors produces the site-specific radiation and atmospheric environment that is the proximate determinant of vine photosynthetic capacity, evaporative demand, disease pressure, and fruit ripening conditions. The quantification of exposure has been advanced by solar radiation modeling tools (hemispherical photography for horizon mapping, GIS-based solar radiation calculators, on-site pyranometer measurements) that enable the total growing season radiation budget to be calculated for any vineyard position, revealing that exposure-related radiation differences of 20-40% between sites within the same commune are common and contribute substantially to the within-appellation terroir variation recognized by vineyard classification systems.
Expositions-Beschreibung
Standort-Exposition
Weinberg-Strahlungsumgebung
radiation environment characterization
site exposure assessment
vineyard exposure record
van Leeuwen C and Seguin G (2006) The concept of terroir in viticulture, Journal of Wine Research 17: 1-10 Gladstones J (1992) Viticulture and Environment, Winetitles Jones GV et al. (2004) Spatial analysis of climate in winegrape growing regions, American Journal of Enology and Viticulture 55: 304-316 Riou C et al. (1994) Le déterminisme climatique de la maturation du raisin: application au zonage de la teneur en sucre dans la Communauté Européenne, Office des Publications Officielles des Communautés Européennes
Exposure Description
Merlot in Bordeaux shows extended cap shedding during mid-late June Year N+1 at BBCH 67-68 over 4-5 days June 12-16, with total flowering period extending to 12-13 days from first to last cap detachment, continuous low-intensity shedding 4-7% per day creating 24-32% sequential tail, resulting in 8-11 day developmental spread between earliest and latest-set berries, under variable oceanic conditions averaging 20-26°C temperature, 58-75% relative humidity, typical of this mid-season cultivar showing moderate developmental coordination in variable maritime climate
Extended cap shedding represents the prolonged calyptra detachment period resulting from developmental heterogeneity when sequential individual flower maturation creates sustained low-intensity cap fall activity over several days, contrasting with compressed synchronized flowering and affecting overall flowering duration and uniformity. This temporal flowering characteristic occurs during progressive cap fall in mid-late June in temperate regions Year N+1, spanning 3-5 days or more in cultivars or conditions showing pronounced developmental spread, corresponding to BBCH 67-68 extended stages. Temporal extension characteristics show prolonged flowering period from first to last cap detachment spanning 10-14 days in extended scenarios compared to 7-10 days in compressed flowering, continuous low-intensity cap fall activity sustained over extended period rather than concentrated mass fall, and overall duration reflecting developmental heterogeneity magnitude. Duration factors contributing to extension include developmental gradient magnitude established during previous season primordium formation with greater spread producing more extended flowering, current season environmental variation affecting differential flower progression rates, and cultivar-specific developmental coordination capacity with some varieties showing inherently more extended flowering. Developmental heterogeneity underlying extension reflects variable flower maturation resulting from heterogeneous inflorescence primordium initiation timing previous season Year N, differential development during current season emergence and pre-anthesis stages, and accumulated variation creating substantial maturation spread by flowering. Sequential progression creating extension shows continuous low-rate cap detachment with daily additions of 3-7% flowers over extended period, cumulative effect producing substantial total completion though daily intensity low, and sustained activity creating prolonged flowering period. Cap accumulation pattern during extended shedding shows continued modest additions over many days creating ultimately substantial total accumulation, less visually dramatic than mass fall but cumulatively significant, and extended debris accumulation period documenting prolonged flowering. Flower exposure during extended shedding shows progressive gradual increase in opened flower proportion over many days, extended period creating wide developmental spread within cluster from earliest to latest opening, and maximum within-cluster heterogeneity at extended shedding completion. Reproductive implications of extended shedding include prolonged pollination window potentially beneficial for overcoming brief adverse conditions affecting any single day, though overall extended period creates greater cumulative weather vulnerability, and late flowers in extended tail showing minimal reproductive success due to pollen limitation. Fruit set outcomes with extended shedding show potential for adequate overall set if conditions favorable during extended period enabling cumulative success, but risk of reduced set if extended period experiences predominantly unfavorable conditions, and increased developmental heterogeneity among set berries from extended pollination timing spread. Berry development heterogeneity resulting from extended shedding creates substantial within-cluster variation with earliest-set berries potentially 8-12 days ahead of latest-set berries by extended shedding completion, affecting ripening uniformity and harvest timing considerations, and potentially creating quality variations within clusters. Environmental effects on extended shedding include variable conditions over prolonged period affecting different portions of flowering differently, cumulative weather exposure increasing probability of encountering some adverse conditions, and overall environmental impact more complex than compressed flowering. Temperature during extended shedding shows averaged effect over many days with warm conditions promoting faster progression shortening extension, cool conditions slowing progression lengthening extension, and variable temperatures creating irregular progression patterns. Humidity during extended shedding affects efficiency throughout period with sustained moderate levels supporting continued detachment, episodes of elevated humidity potentially causing temporary retention or delays, and overall humidity pattern affecting completion timing. Weather vulnerability during extended shedding shows increased cumulative exposure risk with prolonged sensitive period increasing probability of encountering rain or other adverse conditions, though any single adverse event affects smaller proportion of flowering compared to compressed scenario, creating trade-off between acute and chronic vulnerability. Cultivar variation in extension tendency shows some varieties inherently showing extended 12-16 day flowering reflecting poor developmental coordination, others showing compressed 7-9 day flowering from good synchronization, and intermediate varieties showing moderate 9-12 day duration, affecting variety characteristics and management requirements. Environmental effects on extension include unfavorable conditions during inflorescence emergence or pre-anthesis stages potentially increasing heterogeneity extending flowering, while optimal conditions promoting uniformity compressing period, and year-to-year variation in extension within same cultivar reflecting environmental effects. Management implications of extended shedding include prolonged critical weather-sensitive period requiring sustained vigilance and management restrictions, increased within-cluster heterogeneity affecting harvest timing and quality uniformity, and extended flowering indicating possible developmental or environmental problems affecting synchronization. Fruit set assessment complications from extended shedding include difficulty determining appropriate timing for evaluation with wide spread in flower developmental states, mixed developmental appearance complicating visual assessment, and need for extended observation period confirming completion before definitive assessment. Observational characteristics indicating extended shedding include sustained low-intensity cap fall over many days distinguishable from compressed mass fall pattern, wide developmental spread visible within cluster with early berries and late flowers coexisting, and prolonged visible flowering period documented through extended cap accumulation. The process represents second aspect of progressive cap fall emphasizing temporal extension implications, following sequential cap detachment. Duration represents extended 3-5+ day period though individual flowers shed sequentially, with overall flowering period potentially reaching 12-16 days in extreme extensions. Cultivar variation in extension shows some varieties prone to extended shedding reflecting inherent poor developmental coordination while others reliably show compressed flowering, affecting variety selection for regions where flowering conditions critical. Regional expression shows extended shedding most problematic in marginal climates where unfavorable conditions during development may exacerbate heterogeneity, while favorable stable conditions generally promote compressed flowering even in extension-prone varieties. Viticultural significance includes identifying potential quality limitations from excessive within-cluster heterogeneity, complicating harvest timing and uniformity management, increasing cumulative weather vulnerability through prolonged sensitive period, and affecting variety suitability assessment for particular sites or regions. Research applications include analyzing developmental and environmental factors producing flowering extension, evaluating genetic variation in developmental coordination affecting flowering duration, quantifying relationships between flowering extension and berry development heterogeneity, assessing impacts of extension on fruit set outcomes and quality characteristics, and developing management strategies for varieties or situations prone to extended flowering.
Extended_Cap_Shedding is second aspect of Progressive_Cap_Fall, distinguished by temporal extension emphasis and uniformity implications. Follows Sequential_Cap_Detachment. Results from developmental heterogeneity creating prolonged flowering period. Increases within-cluster berry development variation affecting ripening uniformity. Creates prolonged weather vulnerability. Reflects cultivar coordination capacity and environmental effects. Management implications for quality uniformity and harvest timing. Problem characteristic in some cultivars or unfavorable conditions.
Langzeit-Kappenfall
ausgedehnte Kappenablösung
extended calyptra shedding
prolonged cap fall
protracted cap detachment period
protrahierte Abwurfperiode
verlängerte Kalyptralösung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 385-390 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Extended Cap Shedding
An extended long fruiting cane on Vitis vinifera cv. Thompson Seedless in a table grape production system, retained at 18 nodes spanning 195cm along overhead trellis wires, arising from exceptionally vigorous vine showing 1450g pruning weight in highly fertile irrigated soil, positioned to allow extensive lateral shoot development across broad overhead canopy space, expected to produce 17-18 shoots bearing 32-36 clusters totaling 11-14kg fruit, requiring multiple gibberellic acid applications for berry sizing, extensive shoot positioning and tying, cluster thinning to commercial standards, and regular disease management in dense canopy environment, representing maximum productivity approach typical of table grape economics.
A long fruiting cane retained at maximum practical length, typically 14-20 nodes, representing upper limit of cane retention length in high-productivity systems, characterized by extensive linear extent (160-220cm), very high bud count maximizing crop potential, suitability only for exceptionally vigorous vines with documented capacity for high bud support, and requirement for expansive training systems with substantial trellis space and intensive canopy management. Extended long fruiting canes represent aggressive crop load strategies emphasizing yield maximization in fertile sites with vigorous cultivars, accepting increased labor requirements for shoot and crop management, and typically associated with bulk wine production or table grape systems rather than premium wine production where shorter canes and lower crop loads prevail.
Extended long fruiting canes are rare in wine grape production, limited to specific high-yield systems in fertile regions with cultivars combining high fruitfulness with strong vigor (e.g., certain Riesling or Müller-Thurgau clones in Germany, vigorous hybrids in some New World regions). The extended length creates significant canopy management challenges including shoot crowding, shading, disease pressure, uneven ripening, and delayed maturation, making this approach economically viable only where labor costs are low relative to crop value or mechanization can partially offset hand labor requirements. Premium wine production universally avoids extended long canes, with consensus maximum around 12-14 nodes even in vigorous vineyards, recognizing quality compromise beyond this threshold despite potential yield gains.
Maximallänge-Rute
maximum length fruiting cane
sehr lange Fruchtrute
ultra-extended cane
very long cane
Based on Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Bravdo et al. (1985) "Effect of crop level and crop load on growth, yield, must and wine composition, and quality of Cabernet Sauvignon" American Journal of Enology and Viticulture 36=125-131.
Extended Long Fruiting Cane
Gewürztraminer (many clones) exhibits extended long styles 0.72-0.82 mm connecting enhanced ovaries 1.05-1.18 mm to elaborate stigmas 0.52-0.58 mm, with extended long styles representing 29-33% of total pistil length (2.45-2.65 mm), requiring cross-pollination from hermaphrodite pollinizers with pollen tubes traversing the extended style and additional pistil tissues in 65-88 hours at 21-24°C, achieving 30-38% fruit set when adequate hermaphrodite pollinizers provide synchronous pollen availability. Thompson Seedless functionally female flowers show extended long styles 0.74-0.84 mm as part of elaborated pistil morphology (total pistil 2.4-2.7 mm), with extended styles containing well-developed transmitting tract supporting normal pollen tube growth from co-planted hermaphrodite pollinizers, achieving 25-35% fruit set through cross-pollination with subsequent seed abortion during early berry development producing seedless berries despite initial successful fertilization. Orange Muscat constitutively functionally female cultivar exhibits consistent extended long styles 0.72-0.8 mm (averaging 0.76 mm) across diverse sites and vintages, demonstrating genetic stability of extended long style phenotype and representing the typical functionally female style morphology, with styles supporting cross-pollination-dependent reproduction achieving 28-36% fruit set when hermaphrodite pollinizers are present. Breeding populations segregating for flower sexual type show pistillate and functionally female offspring with extended long styles 0.7-0.84 mm versus hermaphrodite offspring with medium styles 0.52-0.68 mm, with the 20-40% style length difference serving as reliable morphological marker for identifying female-functioning flower types during seedling evaluation before flowering and fruit set assessment.
An extended long style is a long style in the moderate-to-upper portion of the long length range, typically 0.7-0.85 mm at maturity, representing styles that clearly exceed hermaphrodite medium length but remain in the more moderate portion of the long style spectrum, characteristic of many functionally female cultivars and some pistillate flowers showing substantial but not extreme style elongation. Extended long styles are characterized by= (1) length typically 0.7-0.85 mm at anthesis, exceeding hermaphrodite medium styles (0.45-0.7 mm) by 10-30% but remaining below elongated long styles (>0.85 mm) (2) diameter typically 0.25-0.35 mm proportional to extended length (3) well-developed transmitting tract tissue with elaborate extracellular matrix and specialized cells supporting effective pollen tube growth through the extended length (4) connection between enhanced ovary (typically 0.95-1.2 mm) and elaborate stigma (typically 0.48-0.6 mm) creating pistils (total length typically 2.3-2.7 mm) with extended long styles representing 28-34% of pistil length. Extended long styles represent moderate pistil elaboration characteristic of functionally female flowers where female organ enhancement is substantial but not maximal, allowing clear distinction from hermaphrodite morphology while avoiding the extreme dimensions seen in some pistillate flowers. Extended long styles develop through= (1) genetic determination in functionally female cultivars producing moderate pistil elaboration including extended style length (2) developmental processes enhancing style elongation beyond hermaphrodite norms but not to extreme levels (3) potentially favorable growing conditions allowing fuller expression of style elongation genetic potential in functionally female genotypes. Extended long styles in functionally female flowers maintain typical proportions within elaborated pistils, representing approximately 28-34% of total pistil length similar to proportions in hermaphrodite pistils, suggesting coordinated pistil enlargement rather than disproportionate style-specific elongation. The functional implications of extended long styles relate to enhanced cross-pollination capability= extended transmitting tract length (0.7-0.85 mm plus additional stigma and ovary tissues totaling 1.7-2.4 mm pollen tube pathway) requires pollen tube growth times of 62-95 hours at typical bloom temperatures (20-24°C), intermediate between hermaphrodite (48-90 hours) and elongated long style flowers (75-110 hours) extended styles provide substantial pollen tube pathway potentially allowing pollen tube selection or quality assessment while avoiding excessive delays that might increase environmental vulnerability functionally female flowers with extended long styles achieve 28-40% fruit set through cross-pollination, comparable to or slightly exceeding elongated long style types, suggesting that extended style length represents functional optimum for cross-pollinating systems without disadvantages from excessive elongation.
Extended_Long_Style represents moderate expression within the long style category and is distinguished from Elongated_Long_Style (>0.85 mm, extreme elongation) by more moderate dimensions while both exceed hermaphrodite medium length. Extended long styles are characteristic of many functionally female cultivars and represent pistil elaboration sufficient for clear sexual type distinction without extreme morphology. Understanding extended long styles is relevant for identifying functionally female flowers, for understanding the range of style length variation in female-functioning types, and for morphometric characterization of pistil elaboration in cross-pollinating flower types.
ausgedehnter langer Griffel
beträchtlich verlängerter Stempelgriffel
moderately long style
mäßig verlängerter Stylus
substantial style
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 95-101 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=94-109.
Extended Long Style
An extended unilateral permanent arm on Vitis vinifera cv. Cabernet Sauvignon at row end, extending 165cm from trunk toward row interior, bearing 26 two-bud spurs distributed along length at average 12.5cm spacing, 35mm diameter at trunk junction tapering to 24mm at terminus indicating substantial structure supporting extended length, total fruiting capacity approximately 50 shoots supporting 8.5-9.5kg crop from single arm, representing compensatory architecture achieving yields comparable to interior bilateral vines (typically two 85cm arms, 26 combined spurs, similar total crop) through extended single-arm length utilizing full available trellis space in permitted direction, with structural integrity and vigor management critical given concentration of entire vine capacity into single elongated element.
A unilateral permanent arm developed to exceptional length compensating for absence of opposite arm, characterized by extended reach typically exceeding 100cm and often approaching 150-200cm, high spur count distributed along extended length, substantial diameter reflecting vigor requirements supporting elongated structure, and function as single-arm equivalent to bilateral pair requiring extended length to achieve comparable total fruiting capacity within unilateral architecture constraint. Extended unilateral permanent arms represent maximum expression of unilateral training, pushing length limits to achieve production targets normally distributed across bilateral pairs, common at row ends where full inter-vine spacing (2-3 meters typical) is available for single-direction extension unlike bilateral configurations dividing this space between paired arms.
Extended unilateral permanent arms approach the practical limits of single-arm training, with lengths beyond 150-180cm encountering problems including structural weakness (excessive cantilever stress at trunk junction), declining spur vigor toward distal positions (vascular pathway length effects), difficult harvest and canopy management (extended reach requirements), and increased winter damage risk (terminal portions exposure). The extended length is typically necessary only at row ends where bilateral development is impossible and where spacing between vines (often 2-3 meters) permits extended single-arm reach, with interior unilateral configurations rarely requiring or supporting such extreme lengths. Management focuses on structural support (avoiding arm breakage under crop load or mechanical stress), maintaining distal spur vigor (potentially through renewal spur positioning or fertilization), and eventual renewal planning (extended arms often showing earlier decline than moderate-length configurations).
Langschenkel einseitig
compensatory unilateral arm
long unilateral arm
maximum-length single arm
verlängerte unilaterale Arm
Extended Unilateral Permanent Arm
Gewürztraminer exhibits extended veraison disposition with the ripening transition spanning 18-28 days from first to last berry within clusters, creating harvest timing dilemmas between under-ripe green berries and over-ripe pink berries. Zinfandel notoriously shows extended veraison with individual cluster sugar ranges of 18-28°Brix at any single harvest date, necessitating multiple harvest passes or acceptance of heterogeneous fruit. Extended veraison in Grenache contributes to the characteristic raisined berry proportion (5-15%) observed at harvest in warm vintages. The coefficient of variation for berry sugar concentration exceeds 20% at harvest in extended veraison cultivars compared to 8-12% in uniform types.
A veraison timing disposition that is the inherited capacity of a Vitis vinifera plant to undergo the ripening transition asynchronously across berries, with substantial temporal dispersion between first and last berries reaching color change and softening, resulting in pronounced within-cluster and within-vine maturity heterogeneity.
Extended veraison disposition complicates harvest decisions and may reduce wine quality consistency through variable fruit maturity. The trait is exacerbated by compact cluster architecture (limited light to interior berries), high crop loads, and water stress during lag phase. Some winemakers exploit extended veraison for stylistic purposes, as the mixture of maturity levels contributes complexity in certain wine styles. Management interventions including cluster thinning, leaf removal, and irrigation management can reduce but not eliminate veraison dispersion in genetically predisposed cultivars.
asynchronous veraison
gestaffelter Farbumschlag
protracted color change
uneven ripening onset
ungleichmäßiger Reifebeginn
verlängerte Véraison
Kasimatis AN et al. (1975) Conversion of cane-pruned Zinfandel vineyards to bilateral cordon training and machine harvest. American Journal of Enology and Viticulture 26:157-160 Friend AP et al. (2009) Delayed winter spur-pruning causes a shift in the source of fruit-zone leaf area in Vitis vinifera Merlot. Australian Journal of Grape and Wine Research 15:254-260
Extended Veraison Disposition
Detailed documentation of Extensive_Root_System_Disposition at Stellenbosch University characterized root systems of rootstocks classified as extensive based on industry experience and research trials. Reference rootstocks '110 Richter', '140 Ruggeri', '1103 Paulsen', and 'SO4' were excavated at 8 years of age from deep granite-derived soils, with complete root recovery and morphological analysis. Extensive_Root_System_Disposition rootstocks showed consistent characteristics. '110 Richter' exhibited archetypal extensive root development with total excavated root length of 3,150 m, root dry weight of 4.8 kg, maximum rooting depth of 2.9 m with 35% of root length below 1 m depth, and horizontal spread of 2.4 m radius from the trunk. Root architecture showed strong vertical orientation with primary framework roots descending steeply before branching. '140 Ruggeri' displayed even more extensive characteristics (3,420 m, 5.2 kg, 3.4 m depth) reflecting its V. berlandieri × V. rupestris parentage combining deep-rooting tendencies of both parents. '1103 Paulsen' showed extensive but shallower distribution (2,880 m, 4.1 kg, 2.2 m depth, 45% in upper 50 cm). Functional assessment revealed that Extensive_Root_System_Disposition rootstocks maintained higher scion water potential during drought (Ψpd -0.35 MPa versus -0.65 MPa for limited root systems at equivalent surface soil moisture), demonstrating the stress-buffering advantage of extensive root development. Total soil volume explored by extensive root systems (estimated from spread and depth) reached 15-25 m³ per vine, compared to 5-10 m³ for limited systems.
An Extensive_Root_System_Disposition is a Root_Vigor_Disposition that inheres in a Vitis rootstock and is realized through vigorous root growth producing a large, well-distributed root system with high total root length, deep penetration, and thorough exploration of the available soil volume, providing substantial capacity for water and nutrient acquisition. This disposition represents the upper range of root vigor, characteristic of rootstocks that develop dominant below-ground structures capable of supporting vigorous scion growth and buffering against environmental stresses through access to large resource pools. Extensive root systems typically show total root lengths exceeding 2,000 m per mature vine, maximum rooting depths of 2-4 m (soil permitting), root length densities above 2 cm/cm³ in the primary rooting zone, and root dry weights of 3-5 kg. Such root systems are advantageous for drought tolerance (accessing deep soil water), nutrient acquisition (exploring large soil volumes), vine stability (extensive anchoring), and reserve storage (large root mass for carbohydrate accumulation). However, Extensive_Root_System_Disposition may confer excessive vigor to scions on fertile, well-watered sites, creating canopy management challenges. Rootstocks with extensive root systems are typically recommended for deep soils, sites with periodic water limitation, and situations requiring vigor enhancement of weak-growing scions.
Extensive_Root_System_Disposition is_a Root_Vigor_Disposition representing high root vigor. Typical characteristics include total root length > 2,000 m, dry weight > 3 kg, rooting depth > 2 m (soil permitting), root length density > 2 cm/cm³. Examples include '110 Richter', '140 Ruggeri', '1103 Paulsen', 'SO4', 'Ramsey'. Advantages include drought tolerance, nutrient acquisition, stress buffering. V. berlandieri and V. rupestris parentage typically confers extensive rooting. Extensive root systems may confer excessive scion vigor on favorable sites.
Ausgedehntes-Wurzelsystem-Disposition
Extensive-Bewurzelung
Kräftiges-Wurzelsystem
extensive rooting
large root system
vigorous root system
Southey JM, Archer E (1988) The effect of rootstock cultivar on grapevine root distribution and density, in The Grapevine Root and its Environment, Department of Agriculture and Water Supply Technical Communication 215, Pretoria Smart DR et al. (2006) Grapevine rooting patterns: a comprehensive analysis and review, American Journal of Enology and Viticulture 57:89-104 Paranychianakis NV et al. (2017) The effect of soil type and irrigation regime on vine performance and root system development, Australian Journal of Grape and Wine Research 23:426-436
Extensive Root System Disposition
External leaf layer quality comparison across training systems in Pfalz Riesling evaluated light interception efficiency and interior canopy illumination. Vertical shoot positioning produced continuous external leaf layer quality with 94% of point quadrat insertions contacting leaves within the first 10 cm of canopy penetration, resulting in 85% total light interception but only 12% transmission to interior fruit zone. Scott Henry divided canopy training positioned two separate external layers, each with 88% continuity, achieving 92% total light interception while providing 22% transmission to interior positions through the canopy division gap. Minimal pruning produced discontinuous external leaf layer quality with only 71% exterior contact rate, reducing total interception to 74% but permitting 35% interior transmission through the irregular canopy structure, demonstrating tradeoffs between exterior continuity and interior accessibility inherent in different canopy architectures.
External leaf layer quality is a leaf layer number quality that inheres in a grapevine canopy system and represents the density and continuity of the outermost layer of foliage forming the canopy exterior surface that intercepts incident solar radiation, serves as the primary interface for atmospheric gas exchange, and presents the initial barrier to spray penetration attempting to reach interior canopy positions. This quality characterizes the photosynthetically most active and environmentally most exposed portion of the canopy where leaves experience full sunlight, maximum temperature fluctuation, and direct environmental exposure that drives the highest rates of carbon assimilation but also the greatest water loss and potential stress susceptibility. The ontological significance of external leaf layer quality lies in its dual role as both the primary productive surface capturing light energy for photosynthesis and the outer barrier whose completeness determines the proportion of incident light reaching interior canopy layers where it could support additional photosynthesis or provide beneficial fruit exposure. A continuous external leaf layer with minimal gaps maximizes light interception by the canopy system as a whole but creates dense shading of interior positions, while a discontinuous exterior with substantial gaps permits light penetration to inner layers but may underutilize total light interception capacity in sparse canopies. The measurement of external leaf layer quality through point quadrat analysis focuses on the first leaf contact per insertion, with the percentage of insertions contacting at least one leaf characterizing exterior continuity and the mean depth to first contact indicating how prominently outer leaves project into the canopy airspace. Optimal external leaf layer quality depends on production objectives and canopy architecture, with dense exterior layers appropriate for systems where interior exposure is managed through canopy division or targeted leaf removal, while more permeable exteriors suit single-plane canopies requiring direct light transmission to interior positions. The relationship between external leaf layer quality and overall canopy function involves consideration of the proportion of total canopy leaf area positioned in well-exposed exterior positions capable of operating near maximum photosynthetic rates, with training system designs differing substantially in the fraction of leaves they position in favorable exterior versus suboptimal interior locations.
External_Leaf_Layer_Quality is_a Leaf_Layer_Number_Quality, representing the outermost canopy surface characterization. This class addresses the first-contact dimension of point quadrat analysis capturing exterior density and continuity. Distinguished from Interior_Leaf_Layer_Quality which addresses deeper canopy positions and from Fruit_Zone_Leaf_Layers_Quality which focuses on cluster-height region regardless of depth. The balance between exterior continuity for light interception and gaps for interior penetration represents key management consideration.
Außenlaubdichte
canopy surface continuity
exterior leaf density
externe Blattschichtqualität
outer canopy density
peripheral foliage layer
äußere Laubschicht
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 44-59 Schultz HR (1995) Grape canopy structure, light microclimate and photosynthesis. I. A two-dimensional model of the spatial distribution of surface area densities and leaf ages in two canopy systems. Vitis 34:211-215 Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review. American Journal of Enology and Viticulture 60:251-268
External Leaf Layer Quality
Extractability maturity assessment comparing analytical phenolic indices with extraction trial wines in Merlot reveals the predictive value of extractability measurement, with pre-optimal harvest showing total anthocyanins 2.0 mg/g berry and extractability index 42% (cell walls relatively intact) yielding wine with 320 mg/L anthocyanins, optimal harvest showing total 1.7 mg/g but extractability 68% (cell wall modification advanced) yielding 450 mg/L in wine, and late harvest showing total 1.4 mg/g with extractability 75% but yielding only 380 mg/L (declining content limiting extraction despite high accessibility), demonstrating how extractability maturity assessment identifies the harvest point where the product of content and accessibility is maximized for wine phenolic concentration.
Extractability maturity quality is a phenolic maturity quality that represents the ease with which phenolic compounds can be released from grape tissues during winemaking, characterizing the cell wall and membrane modifications that affect the transfer of anthocyanins, tannins, and other phenolics from grape solids into must and wine, recognizing that phenolic extractability may be as important as total phenolic content in determining wine phenolic composition. Within the BFO framework, this quality specifically addresses the extraction potential dimension of phenolic maturity, distinguishing the accessibility of phenolic compounds from their accumulation while recognizing that cell wall barriers mediate between grape phenolic content and wine phenolic composition, such that grapes with lower total phenolics but higher extractability may yield wines with greater phenolic richness than grapes with high content but resistant cell walls. The ontological significance lies in extractability maturity determining how much of the grape's phenolic investment is realized in the wine, where cell wall pectin degradation during ripening increases cell permeability, middle lamella breakdown facilitates tissue maceration, and membrane integrity changes affect the release of vacuolar contents, together establishing the extraction efficiency that determines how effectively winemaking can capture phenolic potential. The cell wall changes affecting extractability include pectin solubilization (reducing cell wall rigidity and enabling tissue breakdown), changes in cross-linking among cell wall polymers (affecting porosity), and modifications to the cuticle and epidermis (affecting the initial penetration of extraction medium), with these changes occurring progressively during ripening and varying among tissues (skin cell walls becoming more permeable while seed coats maintain barriers). The measurement approaches for extractability include model extraction systems (ITV method using buffer solutions to simulate maceration), mechanical properties assessment (berry skin hardness correlating with cell wall integrity), and specific cell wall analysis (pectin, hemicellulose quantification), with these methods enabling prediction of extraction behavior beyond what total phenolic analysis can provide. The tissue-specific extractability patterns show skin anthocyanins becoming more extractable during ripening (reaching maximum extractability at intermediate ripeness then potentially declining), skin tannins showing similar patterns, and seed tannins becoming progressively less extractable as lignification proceeds, with these tissue-specific trajectories affecting optimal harvest timing for different phenolic components. Understanding extractability maturity quality enables prediction of winemaking extraction efficiency, harvest timing that optimizes the ratio of extractable to total phenolics, and recognition of the cell wall factors that mediate between grape composition and wine composition.
Extractability_Maturity_Quality addresses the cell wall-mediated accessibility of phenolic compounds during winemaking. This quality recognizes that wine phenolics depend on both grape content and extraction efficiency. The class supports winemaking outcome prediction through extractability characterization beyond total phenolic analysis.
Extrahierbarkeitsreife
Extraktionspotential
Zellwandreife
cell wall maturity
extraction potential
maceration efficiency
phenolic accessibility
phenolische Zugänglichkeit
Saint-Cricq N et al. (1998) Méthode de caractérisation de la maturité phénolique des raisins à partir de l'évolution de l'extractibilité des anthocyanes. Journal International des Sciences de la Vigne et du Vin 32(1): 1-10 Ortega-Regules A et al. (2006) A first approach towards the relationship between grape skin cell-wall composition and anthocyanin extractability. Analytica Chimica Acta 563(1-2): 26-32
Extractability Maturity Quality
The extreme temperature data for the Barossa Valley (South Australia) documents the heat wave profile — mean annual frequency of days with Tmax >35°C is 18-25 days (concentrated in January-February), with multi-day heat wave events (≥3 consecutive days >38°C) occurring approximately 2-3 times per growing season. The January 2014 heat wave (5 consecutive days of 41-46°C at Nuriootpa, with the 46°C reading on January 14 being the highest temperature ever recorded in an Australian wine region) caused catastrophic damage to the 2014 vintage — berry sunburn affecting 20-40% of exposed clusters, berry desiccation (shrivel) reducing yield by 15-25%, premature véraison triggered by heat stress, anthocyanin degradation in red cultivars producing atypical brick-brown coloration, and elevated volatile acidity from accelerated microbial activity in heat-damaged berries. This extreme event intensity is projected to increase under climate change, with the frequency of >40°C days in the Barossa expected to increase from the current 3-5 per year to 8-15 per year by 2050, potentially threatening the viability of traditional warm-climate viticulture without adaptive management (shade cloth, overhead misting, modified canopy architecture, heat-tolerant rootstocks).
A temperature data component that records the frequency, magnitude, and timing of extreme temperature events — both extreme heat (daily maximum temperatures exceeding 35°C that cause photosynthetic shutdown, sunburn damage, accelerated berry dehydration, and inhibition of anthocyanin synthesis) and extreme cold (minimum temperatures below -15°C during dormancy that risk trunk and cordon freeze damage, and below -2°C during the growing season causing frost injury) — providing the assessment of temperature hazard probability that determines the viticultural risk profile of the site and the management interventions required to mitigate extreme temperature impacts.
Extreme_Temperature_Data documents the tail risks of the temperature distribution that have disproportionate impacts on vine health and wine quality relative to their frequency — extreme heat and cold events may occur only a few days per year but can cause more damage than the entire remainder of the growing season. The physiological basis of heat damage involves multiple simultaneous mechanisms — above approximately 35°C, stomatal closure reduces CO₂ assimilation to near zero while respiration continues at accelerated rates, producing net carbon loss above 40°C, protein denaturation begins affecting enzyme function and membrane integrity direct sunburn on berry surfaces facing the afternoon sun produces localized tissue necrosis (brown, desiccated patches) through the combined effects of extreme temperature and UV radiation and anthocyanin degradation occurs through both enzymatic (increased peroxidase activity) and non-enzymatic (thermal decomposition) pathways, producing the characteristic loss of red-purple color and shift toward brown hues in heat-affected red wine vintages: the vine's adaptive response to heat stress includes osmotic adjustment (accumulation of proline and other compatible solutes), heat shock protein synthesis (providing temporary thermotolerance), and emergency transpirational cooling (if water is available), but these defenses are overwhelmed during prolonged multi-day heat events when soil water reserves are depleted and nighttime temperatures remain above 25°C, preventing recovery. The management of extreme heat risk involves both long-term design decisions (row orientation, canopy architecture, shade provision) and short-term emergency responses (overhead misting to provide evaporative cooling, kaolin clay application to reflect radiation, emergency irrigation to maintain transpirational cooling capacity).
Extremtemperatur-Daten
Hitze- und Kälteereignisse
heat wave and cold event data
temperature extreme record
thermal hazard assessment
thermische Gefahrenbewertung
Webb LB et al. (2009) Modelled impact of future climate change on the phenology of winegrapes in Australia, Australian Journal of Grape and Wine Research 15: 165-175 Greer DH and Weston C (2010) Heat stress response of photosynthesis and berry composition of different grapevine species and cultivars, Functional Plant Biology 37: 91-105 Gladstones J (1992) Viticulture and Environment, Winetitles Molitor D et al. (2014) Late frost damage risk for viticulture under climate change, International Journal of Climatology 34: 1127-1144
Extreme Temperature Data
An extremely vigorous water shoot on Vitis vinifera cv. Tempranillo grafted on Richter 110 rootstock in Rioja Alta, Spain, emerging from a trunk latent bud following complete vineyard defoliation by hailstorm in May, achieving 520 cm total extension growth by October with 42 nodes, individual internodes in the 22-28 cm range throughout the mid-shoot region, leaves consistently exceeding 350 cm² area with some approaching 450 cm², stem diameter reaching 19 mm at node 20, total shoot fresh weight of 3.8 kg at season end, continuing active growth into late September preventing proper periderm formation and lignification, creating severe winter freeze vulnerability and requiring removal despite potential framework renewal value due to poor tissue maturation and excessive vigor indicating underlying nutritional imbalance requiring corrective management.
A vigorous water shoot representing the maximum extreme of vegetative growth capacity in Vitis vinifera, characterized by daily elongation rates exceeding 10-15 cm during peak growth phases, internodes routinely reaching 20-30 cm or greater length, leaves frequently surpassing 300-400 cm² area with very pale green coloration and thin cuticle development indicating extremely rapid cell expansion, stem diameters of 15-20 mm or more despite single-season growth, and total seasonal extension potentially reaching 4-6 meters with 35-45 nodes or more under optimal growth conditions. Extremely vigorous water shoots develop under combinations of severe triggering events (90-100% canopy loss), highly fertile soils with abundant nitrogen availability, ample soil moisture throughout the growing season, very vigorous rootstock genotypes, warm growing season temperatures, and cultivars with inherently high vegetative growth potential, creating growth responses that can exceed normal shoot growth by factors of 5-10 or more. These shoots create severe management challenges including massive resource diversion that can prevent fruit ripening in remaining crop, continued late-season growth that compromises dormancy preparation and winter hardiness, mechanical interference with vineyard operations, and increased disease pressure through dense succulent growth that maintains humidity and reduces air circulation. Extremely vigorous water shoots also pose structural hazards through wind breakage of the heavy, poorly lignified stems, and can create permanent framework distortions if their weight bends or breaks underlying support structures.
Extremely vigorous water shoots represent the upper tail of the vigor distribution and are distinguished from merely vigorous water shoots by growth parameters exceeding cultivar-typical values by factors of 2-3 or more. These shoots often indicate serious underlying problems beyond the triggering damage event, including excessive nitrogen fertilization, over-irrigation creating waterlogged root zones, extremely vigorous rootstock incompatibilities, or root system damage that paradoxically triggers excessive shoot growth. Management requires not only shoot removal but diagnostic investigation of causative factors and corrective actions addressing nitrogen reduction, irrigation adjustment, or evaluation of root health. Extremely vigorous water shoots retained for any reason require multi-year vigor reduction programs including severe crop loading, controlled water stress, and potential root pruning to achieve manageable growth characteristics.
Maximalwüchsiger Stammtrieb
extrem kräftiger Wassertrieb
extreme epicormic shoot
hyper-vigorous water shoot
hypervigorer Wasserschoss
maximum vigor sucker
Based on Winkler et al. (1974) "General Viticulture" University of California Press pp. 268-289 Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Naor et al. (2002) "Responses of apple fruit size to tree water status and crop load" Tree Physiology 22=43-50 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 465-492.
Extremely Vigorous Water Shoot
An F1 population from Riesling × VB91-26-4 (powdery mildew resistant) bears F1_Hybrid_Roles, with each seedling carrying 50% Riesling and 50% VB91-26-4 genetics—the population segregating for Run1 resistance, quality traits, and other characteristics for phenotypic selection. F1 progeny from SO4 × V. cinerea crosses serve in F1_Hybrid_Roles in rootstock breeding, evaluated for nematode resistance inherited from the V. cinerea parent and vigor characteristics from SO4. A single selected F1 individual from Cabernet Sauvignon × disease-resistant donor bears an F1_Hybrid_Role as a candidate for variety release if it combines adequate resistance with acceptable wine quality, or as a parent for backcrossing if further quality improvement is needed.
A hybrid role that inheres in a grapevine genotype representing the first filial (F1) generation from a controlled cross, the direct progeny of two parental genotypes combining one haploid chromosome set from each parent without additional generations of crossing or selfing. F1 hybrids exhibit maximum heterozygosity (every locus carrying one allele from each parent), often displaying hybrid vigor (heterosis) manifested as enhanced vegetative growth, stress tolerance, or productivity exceeding either parent. In grapevine breeding, F1 populations represent the initial output of crossing programs, with population sizes ranging from dozens to thousands of individuals depending on breeding objectives and resources. F1 individuals are evaluated for target traits (disease resistance, fruit quality, agronomic performance), with selected individuals advanced either as varieties (if evaluation criteria are met) or as parents for subsequent crossing generations (F2, backcross, or complex crosses). The F1 designation indicates a specific position in breeding program structure—first-generation products of a defined cross—without specifying whether parents were from the same or different species.
F1_Hybrid_Role specifies the generational position that determines genetic expectations—F1 individuals are obligate heterozygotes at any locus where parents differ, enabling prediction of trait segregation based on parental alleles. The F1 generation is particularly important in resistance breeding, where dominant R-genes from heterozygous donors segregate 1:1 in F1 progeny (50% resistant, 50% susceptible), enabling efficient phenotypic or marker-based selection. Heterosis (hybrid vigor) is commonly observed in grapevine F1 populations, potentially complicating breeding by producing exceptional F1 individuals whose performance is not reproducible in subsequent generations due to heterosis loss upon further crossing. The F1 designation does not specify parental taxonomic relationships F1 populations may arise from intraspecific or interspecific crosses, simple or complex parental genotypes.
Erstkreuzung
F1 hybrid
F1-Hybride
Primärhybride
erste Filialgeneration
first cross generation
first filial generation
primary hybrid
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Töpfer R et al. (2011) Molecular breeding in grapes, CRC Press pp.: 160-185 Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330
F1 Hybrid Role
Grenache in Rhône Valley shows failed ovule fertilization during late June-early July Year N+1 at BBCH 69, with 35-55% ovules experiencing fertilization failure from combination of environmental stress and cultivar susceptibility, failed ovules aborting within 2 weeks showing brown collapsed tissue, flowers with complete ovule failure aborting causing coulure, partial failure creating 1-2 seeds instead of potential 3-4, under variable Mediterranean conditions, typical of this coulure-prone cultivar requiring favorable flowering for acceptable fruit set
Failed ovule fertilization represents the reproductive failure outcome when ovule experiences fertilization failure from pollination inadequacy, pollen tube growth problems, fusion abnormalities, or early post-fertilization developmental arrest, resulting in ovule abortion, resource reabsorption, and no seed contribution, potentially causing flower abortion unless other ovules successfully fertilize providing hormonal rescue. This unfavorable reproductive outcome occurs during fertilization and early post-fertilization in late June-early July in temperate regions Year N+1, determined within days of pollination attempts with abortion manifesting over 1-2 weeks, corresponding to BBCH 69 stages. Failed fertilization causes include pollen absence or scarcity preventing ovule pollination, inadequate pollen tube growth failing to reach ovule micropyle from tube dysfunction or environmental limitations, pollen tube penetration failure preventing sperm delivery to embryo sac, failed sperm-egg or sperm-central cell fusion preventing zygote or endosperm formation despite tube discharge, and early post-fertilization developmental failure causing embryo or endosperm abortion. Pollination failure as cause shows insufficient pollen delivery from poor weather during flowering, inadequate pollen production or release, ineffective pollen dispersal, or pollen viability loss preventing successful pollination. Pollen tube failure shows tube growth cessation before reaching ovule from boron deficiency, environmental stress, pollen vigor limitations, or guidance problems, preventing sperm delivery and fertilization. Fusion failure shows sperm cells reaching embryo sac but failing membrane or nuclear fusion, abnormal gamete function preventing successful union, or cellular incompatibility though rare in self-compatible grapevine. Early developmental failure shows initial fertilization success but subsequent embryo or endosperm abortion within days to weeks, developmental arrest from genetic abnormalities or physiological dysfunction, and seed development cessation leaving aborted ovule. Ovule abortion manifestation shows failed ovule tissue beginning degeneration within days of fertilization failure, browning and collapse distinguishing from viable green developing seeds, tissue breakdown and resource mobilization recovering invested nutrients, and aborted remnant visible as dark collapsed structure or seed trace in developing berry. Abortion timing shows rapid progression within 1-2 weeks of fertilization failure, early detection possible 2-3 weeks post-flowering through ovule examination, and complete abortion with tissue reabsorption by berry development phase. Flower fate following complete ovule failure shows all ovules failing eliminating seed development and auxin production, flower lacking hormonal rescue signal failing to prevent abscission, abscission zone formation at pedicel creating separation layer, and flower abortion and drop contributing to coulure poor fruit set. Partial ovule failure within flower shows some ovules failing while others succeed, successful ovule(s) producing seed and auxin rescuing flower from abscission, berry development proceeding with reduced seed number 1-2 instead of potential 3-4, and failed ovules contributing only through reabsorbed resources. Reduced seed number effects show lower total auxin production potentially limiting berry size, berry developing smaller than multi-seed counterparts, and within-cluster size variation from seed number diversity. Auxin absence effects from complete failure show lack of anti-abscission signal permitting programmed flower senescence, abscission progression unchecked by hormonal suppression, and flower drop typically 1-3 weeks post-flowering. Abscission zone development shows specialized cells at pedicel-receptacle junction activating separation program, cell wall degradation creating abscission layer, vascular tissue severing eliminating nutrient flow, and mechanical weakness enabling flower detachment. Coulure relationship to failed fertilization shows poor fruit set resulting from excessive ovule and flower failure, environmental causes including rain during flowering washing pollen and preventing pollination, temperature extremes during fertilization affecting fusion and development, boron deficiency preventing pollen tube growth and fertilization, and genetic susceptibility in certain cultivars prone to fertilization failure. Environmental effects on ovule failure include unfavorable flowering weather increasing failure rate, rain preventing pollination and pollen tube growth, temperature extremes impairing fertilization and early development, boron deficiency as major cause preventing tube growth, and overall conditions during critical 2-3 week period determining failure extent. Temperature effects show cool conditions increasing failure through delayed development and reduced fusion efficiency, excessive heat causing reproductive dysfunction and developmental failure, and optimal 22-28°C range minimizing failure rate. Boron deficiency effects show inadequate boron preventing normal pollen tube wall synthesis, tube bursting or growth cessation before reaching ovules, fertilization failure even with adequate flowering, and boron supplementation critical for reducing failure in deficient conditions. Cultivar variation in failure susceptibility shows genetic differences in fertilization reliability, coulure-prone cultivars showing high 40-70% failure rates even under favorable conditions, reliable cultivars maintaining low 20-40% failure rates across conditions, and variation affecting fruit set predictability. Failed ovule proportion shows typical 30-60% individual ovules failing across flowers depending on conditions, high redundancy from multiple ovules per flower buffering flower-level failure, and population-level patterns determining overall fruit set percentage. Assessment methods include microscopic ovule examination distinguishing viable green from aborted brown ovules, dissection of young berries counting seeds versus aborted remnants, and fruit set percentage calculation comparing berry number to initial flower number. The process represents unfavorable ovule fertilization outcome within Ovule_Fertilization category. Duration represents failure determination and abortion within days to weeks post-pollination. Regional timing shows failed ovule abortion during late June-early July across climates following unsuccessful fertilization attempts. Viticultural significance includes contributing to coulure poor fruit set when excessive, determining final seed number through partial failure patterns, responding to environmental and nutritional conditions affecting management outcomes, and serving as yield limitation in unfavorable seasons or cultivars. Research applications include analyzing fertilization failure mechanisms and causes, evaluating environmental and nutritional effects on failure rates, quantifying genetic variation in failure susceptibility, and developing strategies for reducing failure and improving fruit set reliability.
Failed_Ovule_Fertilization represents unfavorable reproductive outcome within Ovule_Fertilization causing ovule abortion. Results from pollination failure, tube growth problems, fusion abnormalities, or developmental arrest. Manifests within days to weeks as ovule degeneration. Complete failure causes flower abortion contributing to coulure. Partial failure reduces seed number affecting berry size. Environmental factors particularly boron deficiency and weather major causes. Cultivar variation in susceptibility. Determines fruit set loss and yield reduction.
Befruchtungsversagen
Samenanlage-Fehlschlag
Samenentwicklungsversagen
fertilization failure
ovule abortion
reproductive failure
reproduktives Versagen
seed development failure
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 410-415 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 312-315
Failed Ovule Fertilization
Cabernet Sauvignon vineyards at BBCH 69 (end of flowering, >80% of calyptras shed) exhibit heavy accumulation of fallen detached calyptras on basal leaves appearing as brown to dark brown dried brittle structures, often fragmented into individual petal components, with concentrations of 25-60 degraded calyptras per 100 cm² leaf surface area on leaves positioned below inflorescences, providing visual evidence of completed bloom though individual calyptras are difficult to count accurately due to advanced degradation and fragmentation. Pinot Noir vineyards one week post-bloom show fallen detached calyptras as dark brown papery remnants accumulated on leaves, caught in cluster zones, and scattered on ground, with rain events having removed many fallen calyptras from leaf surfaces creating patchy distribution patterns, though sufficient accumulation remains to confirm that vigorous bloom occurred. Riesling in high humidity conditions during bloom shows fallen detached calyptras with visible fungal colonization appearing as grayish or whitish mycelial growth on degraded tan-brown calyptra tissue, with potential for these colonized structures to serve as Botrytis inoculum sources requiring enhanced air circulation through leaf removal to reduce infection risk to developing berries.
A fallen detached calyptra is a detached calyptra that has been separated from the flower for an extended period (typically >1-2 days) and has undergone substantial desiccation, degradation, or fragmentation, appearing as tan, brown, or dark brown dried structures that have lost the fresh yellowish-green coloration and intact morphology of freshly detached calyptras, representing the terminal stage of calyptra existence as recognizable structures before complete decomposition or disappearance through weathering and microbial degradation. Fallen detached calyptras are characterized by= (1) advanced desiccation with tissue appearing dry, brittle, and papery rather than pliable or soft (2) color change to tan, brown, dark brown, or even blackish as chlorophyll degradation completes, polyphenol oxidation proceeds, and tissue necrosis advances (3) structural degradation with partial or complete fragmentation common, creating incomplete cap-like remnants or individual petal fragments rather than intact five-parted structures (4) accumulation on leaves, shoot surfaces, inflorescence structures, or ground beneath vines, often in clusters or layers representing multiple days of calyptra shedding (5) reduced conspicuousness compared to fresh calyptras due to color matching with senescing leaf tissues or soil, making visual detection more difficult. The fallen detached state represents calyptras from flowers that opened days to weeks previously, with the oldest fallen calyptras corresponding to the earliest-opening flowers at the basal inflorescence positions while more recent fallen calyptras represent later-opening apical flowers, creating a mixed population of varying degradation states accumulating throughout the bloom period and persisting for days to weeks post-bloom. The degradation kinetics of fallen detached calyptras depend on environmental conditions= dry conditions (low humidity, high temperatures, direct sun exposure) accelerate desiccation and promote brittleness, leading to rapid fragmentation but slower microbial decomposition moist conditions (high humidity, rain, shade) slow desiccation but accelerate microbial colonization and decomposition intermediate conditions create the longest persistence as recognizable though degraded structures. Fallen detached calyptras accumulate preferentially on surfaces below inflorescences, particularly on basal leaves that intercept falling calyptras, creating characteristic concentration patterns with higher densities on leaves directly beneath flower clusters. Ground accumulation beneath vine rows can create visible layers of brown calyptra debris during and after bloom, though individual calyptras become difficult to distinguish once advanced degradation occurs. The biological significance of fallen detached calyptras relates primarily to their potential role as disease inoculum sources= Botrytis cinerea and other fungal pathogens can colonize fallen calyptras, particularly under moist conditions, with sporulation on degraded calyptra tissue potentially contributing to inoculum loads for subsequent infections of young berries, leaves, or shoots management of excessive calyptra accumulations in cluster zones through leaf removal or air circulation improvement may reduce localized inoculum concentrations in high-risk situations. The utility of fallen detached calyptras for bloom assessment is limited compared to fresh calyptras, as= (1) advanced degradation makes identification and counting difficult and unreliable (2) accumulation represents the summation of multiple days of bloom rather than providing real-time information on current bloom status (3) weathering and removal by rain or wind create stochastic variation in accumulation density unrelated to bloom intensity. However, the presence and abundance of fallen detached calyptras can provide retrospective evidence that bloom has occurred and can give qualitative indications of bloom intensity, with heavy accumulations suggesting vigorous bloom and minimal accumulations suggesting light bloom or significant weather-related removal. Research applications rarely employ fallen detached calyptras for quantitative bloom assessment due to reliability limitations, though they may be sampled for studies of litter decomposition dynamics, microbial succession on senescing floral tissues, or pathogen ecology in vineyard canopy environments.
Fallen_Detached_Calyptra represents the degraded terminal state of detached calyptras and is distinguished from Freshly_Detached_Calyptra by advanced desiccation, color change to brown tones, structural fragmentation, and extended time since separation (>1-2 days versus hours to 1-2 days). The fallen detached state has limited utility for precise bloom assessment due to degradation and accumulation dynamics but provides qualitative retrospective evidence of bloom occurrence and intensity. Management considerations for fallen detached calyptras are primarily related to their potential role as disease inoculum sources in high-risk situations, though in most viticultural contexts fallen calyptras pose minimal issues and decompose naturally within weeks of bloom completion.
abgefallene Blütenhaube
degraded shed calyptra
dried fallen petal cap
verwitterte Kelchhaube
zerfallene Kronblattkappe
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 103-104 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 96-97.
Fallen Detached Calyptra
The false homonym pair Riesling and Welschriesling (syn. Riesling Italico, Olaszrizling, Laški Rizling) creates persistent confusion despite the two names being orthographically distinguishable — Welschriesling (VIVC 13096) is a genetically distinct cultivar unrelated to Riesling (VIVC 10077) with no confirmed parentage connection, yet the shared root Riesling causes consumer and trade confusion across Central European markets where both cultivars are widely planted, leading to regulatory measures in Germany prohibiting the use of the bare name Riesling for Welschriesling and EU labeling rules requiring qualifying modifiers to prevent consumer deception.
A homonym where two or more cultivar names are sufficiently similar in spelling or pronunciation to be confused as identical designations but are in fact orthographically distinct names that can be discriminated through careful attention to spelling, diacritical marks, or qualifying modifiers, creating apparent rather than true nomenclatural ambiguity.
False_Homonym addresses the penumbra of nomenclatural confusion where names are similar enough to generate misidentification or consumer deception but technically distinct upon careful inspection. The category is practically important because false homonyms drive regulatory intervention and labeling disputes more frequently than true homonyms: the Riesling/Welschriesling distinction has generated decades of legal and regulatory conflict across EU wine law, and the Pinot/Pinotage pair (genetically unrelated despite the shared root — Pinotage being a Pinot Noir x Cinsaut cross) similarly requires regulatory clarity. False homonymy also affects database operations where fuzzy matching algorithms may erroneously link similar but distinct names.
Pseudo-Homonym
confusable name pair
falsches Homonym
near-homonym
pseudo-homonym
verwechselbare Sortenbezeichnung
OIV (2013) OIV List of Vine Varieties and Their Synonyms EU Regulation 1308/2013 Art. 100 on protection of designations Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157
False Homonym
Famous wine association for Nebbiolo documents an exceptionally concentrated prestige network — the cultivar is inseparably identified with Barolo DOCG (the "wine of kings, king of wines," classified since 1980) and Barbaresco DOCG, with specific legendary estates defining its quality ceiling (Giacomo Conterno's Monfortino Riserva, Gaja's single-vineyard Barbaresco crus, Bruno Giacosa's Le Rocche del Falletto), its reputation reinforced by the wines' extraordinary aging capacity (documented examples from the 1930s-1960s still vibrant), and yet the cultivar's global planting remains remarkably limited (approximately 5,500 ha worldwide), illustrating how famous wine association can sustain cultivar prestige independently of commercial ubiquity. Famous wine association for Grüner Veltliner documents a more recent but transformative prestige construction — the cultivar's international reputation was essentially created by three celebrated Wachau estates (F.X. Pichler, Emmerich Knoll, Prager) whose Smaragd bottlings demonstrated world-class quality from the 1980s onward, reinforced by the 2002 London tasting organized by Jancis Robinson where Grüner Veltliner outperformed white Burgundy in blind assessment, and consolidated through the DAC appellation system (Weinviertel 2002, Mittelburgenland 2005, Kamptal 2008, Kremstal 2007, Wachau 2020) that codified Grüner Veltliner's role as the defining Austrian white grape.
Famous wine association is a historical importance entity that documents the relationship between grape cultivars and the celebrated wines, estates, appellations, and wine personalities with which they are most prominently identified in wine culture, encompassing the association with classified or iconic estates (Bordeaux classified growths, Burgundy Grand Crus, Barolo crus), the identification with specific appellations whose identity is inseparable from the cultivar (Chablis-Chardonnay, Barolo-Nebbiolo, Sancerre-Sauvignon Blanc, Wachau-Grüner Veltliner), the connection to influential winemakers or viticulturists who championed the cultivar, and the role of specific wines in defining or transforming the cultivar's global reputation, providing the cultural-prestige dimension that explains why certain cultivars command premium prices and cultural reverence beyond what their intrinsic agronomic or oenological characteristics alone would justify.
Famous wine association documentation addresses the cultural construction of cultivar prestige — the process by which specific cultivar-place-producer combinations become embedded in wine culture as reference points of quality and authenticity that subsequently influence the perceived value of all wines made from that cultivar. This process operates through multiple reinforcing mechanisms — critical acclaim (high scores from influential critics creating reputation feedback loops), auction performance (record prices validating and publicizing quality claims), classification systems (formal hierarchies like the 1855 Bordeaux classification encoding prestige into institutional structures that persist for generations), and narrative construction (the stories, myths, and personalities associated with famous wines becoming part of the cultivar's cultural identity). The relationship between cultivar and famous wine is bidirectional — Pinot Noir derives prestige from its association with Romanée-Conti, but Romanée-Conti also validates Pinot Noir as a vehicle for transcendent wine quality, creating a self-reinforcing prestige cycle. The geographic concentration of famous wine associations is notably uneven — a small number of European regions (Bordeaux, Burgundy, Champagne, Piedmont, the Mosel, Tokaj) account for a disproportionate share of the world's most famous wines, reflecting both genuine quality achievement and the accumulated cultural capital of centuries of documented excellence. The "halo effect" of famous wine associations extends beyond the specific estates and regions to influence the perception of the cultivar globally — Pinot Noir commands premium prices in Oregon, New Zealand, and Tasmania partly because of its Burgundian associations, even though these New World expressions are stylistically distinct. The contemporary democratization of wine criticism through digital platforms and social media is gradually diversifying the famous wine association landscape beyond the traditional European prestige centers, creating new cultivar-region associations (Argentine Malbec, New Zealand Sauvignon Blanc, Austrian Grüner Veltliner) that challenge the historical European monopoly on viticultural prestige.
Prestigeassoziation
berühmte Weinassoziation
berühmte Weingutverbindung
celebrated wine link
cultural wine association
famous estate connection
iconic wine association
ikonische Weinverbindung
kulturelle Weinassoziation
prestige association
Robinson J, Harding J (2015) The Oxford Companion to Wine, 4th Ed, Oxford University Press Johnson H, Robinson J (2019) The World Atlas of Wine, 8th Ed, Mitchell Beazley, pp.: 1-416 Veseth M (2011) Wine Wars: The Curse of the Blue Nun, the Miracle of Two Buck Chuck, and the Revenge of the Terroirists, Rowman and Littlefield, pp.: 1-250 Lukacs P (2012) Inventing Wine: A New History of One of the World's Most Ancient Pleasures, W.W. Norton, pp.: 1-350
Famous Wine Association
Long-term evaluation of Fanleaf_Resistance_Disposition at the University of California, Davis, compared disease development in 'Cabernet Sauvignon' grafted onto susceptible ('AxR#1', V. vinifera × V. rupestris) versus resistant ('O39-16', V. vinifera × M. rotundifolia) rootstocks planted in a vineyard naturally infested with GFLV-viruliferous Xiphinema index. After 12 years, 95% of vines on 'AxR#1' tested GFLV-positive by ELISA, exhibiting severe fanleaf symptoms with mean yield of 4.2 kg/vine compared to 11.8 kg/vine for healthy controls (64% reduction). In contrast, only 8% of vines on 'O39-16' tested positive, with infected individuals showing low viral titers and minimal symptoms, and plot mean yield of 10.5 kg/vine (89% of healthy controls). Nematode population monitoring revealed X. index densities of 850 individuals per 250g soil around 'AxR#1' roots versus only 45 per 250g soil around 'O39-16', demonstrating that Fanleaf_Resistance_Disposition of the rootstock operated primarily through suppression of vector populations rather than direct viral resistance, effectively interrupting the transmission cycle and protecting the susceptible scion.
A Fanleaf_Resistance_Disposition is a Viral_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that limit infection, replication, systemic spread, or symptom expression by Grapevine fanleaf virus (GFLV) and the closely related Arabis mosaic virus (ArMV), nepoviruses causing fanleaf degeneration disease. This disposition represents the capacity to resist one of the most ancient and economically important viral diseases of grapevine, endemic to European and Mediterranean viticulture for centuries and now distributed worldwide. GFLV causes characteristic symptoms including fan-shaped leaves with abnormally acute sinuses, yellow mosaic or vein-banding patterns, shortened internodes, abnormal branching, poor fruit set (millerandage), and progressive vine decline with yield losses of 50-80% in severely affected vines. The virus is transmitted persistently by ectoparasitic dagger nematodes of the genus Xiphinema, primarily X. index and X. italiae, which acquire virus particles during root feeding and retain infectivity for months to years. Fanleaf resistance dispositions can operate at multiple levels including resistance to nematode feeding preventing virus acquisition from soil inoculum, resistance to virus replication and systemic movement following nematode-mediated inoculation, and tolerance enabling maintenance of productivity despite infection. True resistance to GFLV itself is rare in Vitis, but strong resistance to X. index exists in certain North American species and Muscadinia rotundifolia, providing the basis for resistant rootstock breeding that represents the most effective long-term management strategy for fanleaf-affected vineyards.
Fanleaf_Resistance_Disposition is_a Viral_Resistance_Disposition. Fanleaf disease is caused by GFLV and related nepoviruses transmitted by Xiphinema dagger nematodes. Child classes distinguish GFLV_Resistance_Disposition (direct resistance to virus replication/movement) from Xiphinema_Transmission_Resistance_Disposition (resistance to nematode vector). Practical fanleaf management relies primarily on X. index-resistant rootstocks, as true resistance to GFLV itself is limited in available germplasm. The disease has been present in European viticulture for centuries, likely since Roman times.
Fanleaf-Resistenz
GFLV resistance disposition
GFLV-Resistenz
Reisigkrankheit-Resistenz-Disposition
fanleaf degeneration resistance
nepovirus resistance
Andret-Link P et al. (2004) Grapevine fanleaf virus: still a major threat to the grapevine industry, Journal of Plant Pathology 86:183-195 Walker MA et al. (1994) Resistance to Xiphinema index in Vitis and its hybrids, Vitis 33:21-26 Fuchs M (2017) Grapevine fanleaf virus, Grapevine Viruses: Molecular Biology, Diagnostics and Management, Springer
Fanleaf Resistance Disposition
Fanleaf symptom quality assessment in Burgundy Pinot Noir documented the characteristic symptom complex enabling field diagnosis of this historically significant disease affecting some of the world's most valuable vineyard land. Spring shoot emergence revealed fanleaf symptom quality with the diagnostic leaf malformation showing abnormally wide petiolar sinus creating the fan-shaped appearance, acute angles between primary veins deviating from the normal obtuse venation, and asymmetric blade development with unequal lobe sizes. By midsummer, fanleaf symptom quality included the yellow mosaic pattern with bright chrome yellow patches distributed irregularly across the leaf blade, shortened internodes creating bunchy growth habit with shoot length reduced by 40% compared to healthy vines, and the zigzag shoot pattern resulting from asymmetric internode development. Cluster fanleaf symptom quality showed severe millerandage with 60% of berries remaining small, seedless, and unripe at harvest while the remainder developed normally, combined with reduced cluster number per shoot from coulure, resulting in yield reduction of 75% compared to healthy adjacent vines. Soil sampling confirmed high populations of Xiphinema index (150-300 nematodes per 500cc soil) in the root zone of symptomatic vines, establishing the vector presence sustaining endemic infection. The fanleaf symptom quality assessment informed the replanting protocol requiring vine removal followed by 7-year fallow with nematode monitoring before replanting with GFLV-free certified stock.
Fanleaf symptom quality is a viral infection indicator quality that inheres in grapevine organisms infected by grapevine fanleaf virus (GFLV) and represents the characteristic disease manifestations including the diagnostic fan-shaped leaves with abnormally open petiolar sinus and acute vein angles, the yellow mosaic or chrome yellow discoloration patterns on leaves, the shortened internodes creating a bunchy or bushy growth habit, the zigzag growth pattern of affected shoots, and the severe fruit symptoms including millerandage (shot berries), coulure (poor fruit set), and reduced cluster size that can decrease yields by 80% or more in severely affected vines. This quality captures the diagnostic features of one of the oldest known and most damaging viral diseases of grapevine, recognizing that fanleaf degeneration has been described in European vineyards for centuries and remains a major production constraint wherever the nematode vector is present in soils. The ontological significance of fanleaf symptom quality lies in its role enabling identification of a disease causing severe yield reduction that cannot be cured in established vines, where diagnosis informs replanting decisions including the necessity of extended fallow periods or soil fumigation to reduce nematode vector populations before establishing new plantings. The biological basis of fanleaf symptoms encompasses the systemic infection and high viral titers achieved by GFLV in all vine tissues, the interference with auxin signaling that causes the characteristic developmental abnormalities including leaf malformation and growth habit changes, and the disruption of reproductive development causing the flower abortion and berry development failures that produce the severe yield losses. The nematode transmission by Xiphinema index feeding on roots creates soil-borne disease pressure that persists in infested sites for decades, with nematodes retaining transmissible virus for months to years and virus surviving in root fragments remaining after vine removal, necessitating extended fallow periods or nematicide treatment before replanting. The strain variation in GFLV creates diverse symptom expressions from severe fanleaf malformation through yellow mosaic to vein banding patterns, with mixed infections potentially causing more severe symptoms than single strains.
Fanleaf_Symptom_Quality is_a Viral_Infection_Indicator_Quality, representing the symptoms of grapevine fanleaf virus (GFLV) infection. The fan-shaped leaf malformation is pathognomonic yellow mosaic pattern also highly diagnostic. Nematode (Xiphinema index) transmission creates persistent soil-borne disease pressure. Severe yield impacts (up to 80% loss) from millerandage and coulure. Extended fallow (5-7 years) required before replanting infested sites. One of the oldest described viral diseases of grapevine with centuries of European documentation.
Fanleaf-Virose
Fächerblatt-Symptome
GFLV symptoms
GFLV-Symptome
Reisigkrankheit-Symptome
fanleaf degeneration symptoms
fanleaf virus disease
grapevine fanleaf symptoms
Andret-Link P et al. (2004) Grapevine fanleaf virus: still a major threat to the grapevine industry. Journal of Plant Pathology 86:183-195 Fuchs M et al. (2017) Grapevine fanleaf virus. In: Meng B et al. (eds) Grapevine Viruses: Molecular Biology, Diagnostics and Management. Springer, pp. 319-330 Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests, 2nd Edition. APS Press
Fanleaf Symptom Quality
Pinot Noir vineyard showing feeder roots in upper 30 cm soil, minirhizotron imaging revealing feeder roots 0.4-1.8 mm diameter third through fifth-order laterals, feeder roots showing white coloration with abundant root hairs, primary anatomical structure lacking secondary xylem, arbuscular mycorrhizal colonization 58% of feeder root length, median feeder root lifespan 4.2 months with continuous production and mortality, feeder roots comprising 8% of total root biomass but providing estimated 84% of vine nitrogen and water uptake, feeder roots representing ephemeral absorptive specialist root class
Feeder root is fine absorptive root typically 2 mm), showing different functions (feeder specialized for absorption versus structural for transport, storage, anchorage), exhibiting lifespan differences (feeder ephemeral weeks-months versus structural persistent years-decades), demonstrating anatomical distinctions (feeder primary structure versus structural secondary growth and wood), and revealing biomass proportions (feeder roots small biomass fraction providing majority of function). Feeder root research and measurement includes= requiring specialized methods (minirhizotrons, soil cores, root washing for observing and quantifying ephemeral feeder roots), showing isotope applications (tracers demonstrating feeder root uptake contribution), exhibiting imaging techniques (scanners or cameras quantifying feeder root length, diameter, and turnover), demonstrating molecular approaches (DNA/RNA analysis revealing feeder root gene expression and function), and displaying modeling applications (feeder root dynamics incorporated into models predicting vine water and nutrient status).
Feeder_Root emphasizes fine absorptive specialization and ephemeral nature and is distinguished from structural roots by focus on absorption function. Includes Active_Feeder_Root and Senescent_Feeder_Root hierarchies. Critical for understanding absorption specialization, high turnover, disproportionate uptake, root hair production, mycorrhizal associations, and the fine ephemeral roots providing majority of vine resource acquisition despite small biomass fraction.
Absorptive Terminalwurzel
Ephemeral root
Feinwurzel
Fine absorptive root
Saugwurzel
Terminal root
Feeder Root
Female flower disposition characterizes wild V. vinifera ssp. sylvestris populations (approximately 40% of individuals) and rare cultivated varieties. Stamens in female flowers are reflexed (bent downward and away from the pistil) and produce shriveled, non-viable pollen or no pollen at all. Female vines require pollinizer rows within 15-30 m for adequate fruit set, with pollen transfer primarily by wind. Some table grape cultivars including Almeria (Ohanez) and certain Muscats possess female flowers, requiring interplanting with hermaphrodite pollinizers at ratios of 1:8 to 1:10.
A flower type disposition that is the inherited capacity of a Vitis vinifera plant to develop pistillate (functionally female) flowers containing a fully developed pistil with receptive stigma but only reflexed, non-functional stamens producing sterile or absent pollen, requiring cross-pollination from hermaphrodite or male vines for fruit set.
Female flower disposition results from homozygosity for the female allele (ff) at the Sex locus. The reflexed stamen phenotype results from failure of filament elongation during flower development. Female-flowered cultivars were more common in historical viticulture before the advantages of hermaphroditism were understood, and persist in some traditional growing regions and wild populations.
Weiblichkeit
female vine disposition
functionally female flower
pistillate Blühdisposition
pistillate flower trait
weibliche Blütenanlage
Antcliff AJ (1980) Inheritance of sex in Vitis. Annales de l'Amélioration des Plantes 30:113-122 Negi SS, Olmo HP (1971) Conversion and determination of sex in Vitis vinifera L. Vitis 9:265-279
Female Flower Disposition
Wild V. vinifera ssp. sylvestris female plants in riparian populations of southern Europe occasionally exhibit female only flowers completely lacking any stamen structures, with well-developed pistils (2.4-2.7 mm length, ovary 1.4-1.6 mm diameter containing 3-4 ovules, elaborate wet stigma with 200-240 papillae persisting 6-8 days receptive), no recognizable stamens in the five stamen positions surrounding the pistil, achieving 32-44% fruit set when staminate male plants providing wind-borne pollen are within 20-40 m and bloom synchrony is good, but only 6-10% fruit set when males exceed 80 m distance or bloom asynchrony creates temporal mismatch between stigma receptivity and pollen availability. Wild Vitis riparia female plants in North American riparian forests characteristically bear female only flowers with complete stamen absence, enhanced pistil development, and absolute cross-pollination dependency, producing clusters of 20-50 small berries (6-10 mm diameter) when pollinated by nearby male plants. Breeding populations from wild dioecious species crosses occasionally produce female only individuals serving as female parents in controlled crossing programs but unsuitable for self-fertile cultivar development.
A female only flower is a pistillate flower in which stamens are completely absent or reduced to such an extreme degree that no recognizable stamen structures can be identified even under magnification, representing the most complete expression of morphological female unisexuality with exclusive investment in female reproductive function through a well-developed pistil capable of fertilization and fruit set when cross-pollinated. Female only flowers are characterized by= (1) complete absence of any stamen structures in the positions where stamens would occur in hermaphrodite flowers, with the pistil surrounded by empty space or minimal undifferentiated tissue rather than by recognizable filaments and anthers, or (2) extremely minute, completely non-differentiated tissue remnants 100 m, wet or humid weather preventing pollen dispersal). The extended stigma receptivity period (5-8 days) and enhanced stigmatic surface area in female only flowers represent adaptations increasing probability of successful cross-pollination by maximizing the temporal and spatial windows for pollen capture from stochastic wind-borne pollen arrival. Female only flowers are characteristic of some female individuals in wild dioecious V. vinifera ssp. sylvestris populations, particularly in populations with strong sex determination, where approximately 30-50% of female individuals may bear female only flowers while remaining females bear pistillate flowers with vestigial or rudimentary stamen remnants. Female only flowers also characterize female plants in several dioecious North American wild Vitis species (V. riparia, V. rupestris) where stamen suppression is complete. The developmental genetics underlying female only flowers involves sex-determining genes that completely suppress stamen primordium specification or arrest stamen development at extremely early stages, possibly through direct transcriptional repression of stamen identity genes (B-class MADS-box genes) or through developmental pathway disruption preventing stamen organ formation.
Female_Only_Flower represents maximum stamen suppression among pistillate flowers and is distinguished from Reduced_Stamen_Flower (which retains vestigial or rudimentary stamen structures) by complete or near-complete absence of recognizable stamen morphology. The boundary between 'female only' (no stamens) and 'vestigial stamen' (minimal stamen remnants) can be subtle, requiring careful dissection and magnification, as extremely reduced vestigial stamens <0.2 mm can be overlooked in casual examination. Female only flowers represent the endpoint of stamen suppression and likely involve the strongest expression of female-determining genetic factors. The complete absence of male structures enables maximum resource allocation to female function (enhanced pistil development, increased ovule number, prolonged stigma receptivity), potentially providing fitness advantages in dioecious populations through enhanced seed production per successfully pollinated flower. Sub-classification into Fully_Developed and Mature types reflects developmental state distinctions within the female only category, paralleling the staminate flower classification.
purely pistillate flower
rein weibliche Blüte
stamen-absent flower
staubblattlose Blüte
vollständig weibliche Blüte
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 101-105 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Caporali, E. et al. (2003). The arrest of development of abortive reproductive organs in the unisexual flower of Vitis vinifera. Sexual Plant Reproduction 15=291-300 Badouin, H. et al. (2020). The wild grape genome sequence provides insights into the transition from dioecy to hermaphroditism. Genome Biology 21=223.
Female Only Flower
Fermentation protocol for Sauvignon Blanc from Marlborough specifies cold settling at 6°C for 24 hours, inoculation with VL3 yeast (selected for thiol-releasing carbon-sulfur lyase activity) at 20 g/hL, fermentation temperature controlled at 12-14°C in jacketed stainless steel tanks (15,000 L) to preserve volatile thiol stability, DAP addition of 0.20 g/L at 15° Brix based on YAN assessment of 180 mg/L, fermentation duration 18-22 days, endpoint at <2 g/L residual sugar, malolactic fermentation blocked by early SO₂ addition (50 mg/L) and cold storage at 4°C to preserve the high-acid, thiol-dominant aromatic profile. Fermentation protocol for Blaufränkisch Mittelburgenland DAC documents destemmed must inoculated with BDX yeast at 25 g/hL, fermentation initiated at 22°C rising to a peak of 28-30°C through the cap heating effect during maceration, nutrient supplementation with 0.30 g/L Fermaid-K at one-third sugar depletion, fermentation completed to dryness (<1 g/L RS) after 10-12 days, followed by co-inoculation of Oenococcus oeni VP41 for malolactic fermentation completing within 3 weeks at 20°C, converting 4.5 g/L malic acid to approximately 3.0 g/L lactic acid.
Fermentation protocol is a winemaking protocol entity that documents the parameters, conditions, and management practices governing the primary alcoholic fermentation of grape must by Saccharomyces cerevisiae and, where applicable, the secondary malolactic fermentation by Oenococcus oeni or Lactobacillus species, encompassing yeast selection (commercial strain or indigenous), inoculation rate and timing, temperature profile (target range, cooling capacity, peak temperature), nutrient management (YAN assessment and supplementation strategy), fermentation vessel type and volume, fermentation kinetics monitoring (density curve, CO₂ evolution), endpoint criteria (residual sugar target), and malolactic fermentation management (inoculated or spontaneous, timing relative to alcoholic fermentation), providing the microbiological process control framework for converting grape sugar to ethanol while optimizing the production of desirable fermentation-derived aroma compounds.
Fermentation protocol documentation captures the most microbiologically complex and kinetically dynamic phase of winemaking, where the interaction between yeast metabolism, must composition, and environmental conditions determines not only the basic conversion of sugar to alcohol but also the production of hundreds of fermentation-derived volatile compounds that constitute the fermentative bouquet. The yeast strain selection decision has become one of the most consequential protocol elements, as commercial Saccharomyces cerevisiae strains vary dramatically in their metabolic phenotypes — β-glucosidase activity (releasing bound terpenes in aromatic varieties), carbon-sulfur lyase activity (releasing volatile thiols from cysteinylated precursors), ester production profiles, glycerol yield, sulfur compound generation, and temperature tolerance range. The indigenous versus inoculated fermentation debate represents a fundamental philosophical divide — indigenous fermentation (relying on vineyard and cellar microflora, typically involving successive dominance of non-Saccharomyces species followed by S. cerevisiae) is favored by terroir-oriented producers for contributing site-specific complexity, while inoculated fermentation provides predictability and targeted metabolic outcomes. Temperature management during fermentation represents the primary control variable — white and rosé wines are typically fermented at 12-18°C to preserve primary fruit aromas and minimize higher alcohol production, while red wines ferment at 25-32°C to optimize phenolic extraction during concurrent maceration. The malolactic fermentation decision (encourage, prevent, or partial) is among the most stylistically consequential protocol choices — MLF softens acidity, enhances mouthfeel, and adds diacetyl-derived buttery complexity, but sacrifices primary fruit freshness and malic acid-derived crispness, making it desirable for most red wines and barrel-fermented Chardonnay but typically avoided in aromatic whites and high-acid styles. Fermentation kinetics monitoring through daily density measurement (hydrometer, digital densitometer, or in-line sensors) provides real-time process control, enabling early detection of sluggish or stuck fermentation before remedial intervention becomes difficult.
Fermentationsprotokoll
Gärführung
Gärprotokoll
Gärungssteuerung
alcoholic fermentation protocol
alkoholische Gärung Protokoll
fermentation management
fermentation parameters
fermentation process specification
vinification fermentation data
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1, 2nd Ed, Wiley, pp.: 53-200 Ugliano M, Henschke PA (2009) Yeasts and wine flavour. In Wine Chemistry and Biochemistry, Springer, pp.: 313-392 Boulton RB et al. (1999) Principles and Practices of Winemaking, Springer, pp.: 102-190
Fermentation Protocol
The fertigation program for a Shiraz vineyard in the Barossa Valley specifies nutrient injection through the subsurface drip irrigation system using a Dosatron proportional injector at 1:200 dilution ratio — nitrogen delivered as calcium ammonium nitrate solution (CAN-17, 8.5% NO3-N + 8.5% NH4-N) at 15 kg N/ha split across 6 fertigation events from E-L 9 to E-L 27 (2.5 kg N/ha per event), with nitrogen front-loaded during the pre-flowering period to support canopy development and progressively reduced after fruit set to avoid stimulating late-season vegetative growth. Potassium delivered as potassium nitrate (KNO3, 13% N + 46% K2O) at 20 kg K/ha in 4 post-véraison fertigations to support sugar transport and replace harvest potassium removal. Phosphorus and micronutrients (chelated Fe, Zn, Mn) applied in a single early-season fertigation at E-L 12 through phosphoric acid and chelate solution injection — total annual fertigation nutrient cost of 180-250 AUD/ha compared to 300-400 AUD/ha for equivalent broadcast granular fertilization.
A nutrition management description that documents the delivery of dissolved mineral nutrients through the irrigation system, specifying the nutrient sources (soluble fertilizer formulations), injection rates, application timing synchronized with vine phenological demand, nutrient concentrations in the irrigation solution, and the integration of fertigation scheduling with the irrigation protocol to achieve targeted nutrient delivery directly to the active root zone with high spatial precision and temporal flexibility.
Fertigation_Program documents the integration of nutrition management with irrigation infrastructure that has become the standard nutrient delivery method in irrigated vineyards worldwide. Fertigation exploits the spatial precision of drip irrigation (delivering nutrients directly to the wetted root zone volume rather than broadcasting across the entire vineyard floor) and the temporal flexibility of soluble nutrient injection (enabling nutrient supply to be adjusted at each irrigation event in response to vine demand and tissue analysis feedback) to achieve nutrient use efficiency 20-40% higher than conventional broadcast fertilization: this efficiency gain results from reduced nutrient losses through leaching below the root zone, surface runoff, volatilization, and fixation in non-root-zone soil, with the economic benefit of lower total nutrient application rates combined with the environmental benefit of reduced nutrient loading to groundwater and surface water. The principal technical requirements are water-soluble fertilizer sources (excluding standard granular formulations), injection equipment (Venturi, diaphragm pump, or proportional dosing systems), and irrigation system filtration adequate to prevent emitter clogging from precipitated salts — the risk of calcium phosphate or iron oxide precipitation in the irrigation lines requires careful attention to solution pH and nutrient compatibility when multiple fertilizer sources are co-injected.
Bewässerungsdüngung-Plan
Fertigation-Programm
Tropfbewässerungs-Düngung
drip-applied nutrition program
fertigation schedule
irrigation-delivered fertilization
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 180-195 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 168-195 Bravdo BA (2005) Nutrient management in table and wine grapes by means of fertigation, in Christensen LP and Smart DR (eds) Proceedings of the Soil Environment and Vine Mineral Nutrition Symposium, ASEV pp: 91-98
Fertigation Program
A fertile reproductive shoot on Vitis vinifera cv. Sangiovese from node 7 on bilateral cordon, 148cm length with 17 leaves totaling approximately 2380 cm² leaf area, bearing 2 well-formed clusters with 56 and 52 berries showing normal development progression, good fruit set estimated at 42% (typical for this variety), clusters developing normally through cell division and early expansion phases (currently BBCH 75, pea-sized berries), adequate leaf-to-fruit ratio (14.8 cm² per gram projected fruit assuming 325g per cluster) supporting complete maturation, no visible disease, stress, or developmental problems, representing successful fertile shoot contributing 650g to projected whole-vine crop of 8.2kg in balanced mature vineyard, with fertile performance reflecting appropriate winter pruning (balanced bud retention), successful spring pollination (favorable bloom weather, adequate bee activity), good fruit set, and ongoing favorable conditions supporting normal development toward harvest maturity.
A reproductive shoot demonstrating high reproductive capacity and successful fruit development, characterized by presence of well-developed inflorescences with good fruit set (typically 35-65% of flowers setting fruit), healthy cluster development progressing normally through phenological stages, adequate shoot vigor and leaf area supporting fruit maturation (12+ cm² leaf area per gram fruit minimum), and absence of significant reproductive failures or developmental problems, representing successful productive shoots contributing reliably to commercial yields with good prospects for fruit quality outcomes. Fertile reproductive shoots represent the productive ideal in viticulture, exhibiting successful progression from bud differentiation through flower development, pollination, fruit set, and berry development without significant losses or developmental abnormalities, typical of well-managed balanced vineyards under favorable conditions, with fertile shoot performance indicating appropriate vine capacity, adequate resource availability, suitable environmental conditions, and effective management practices supporting consistent productive outcomes.
Fertile reproductive shoots represent successful productive outcomes, with their consistent performance across vineyards and seasons being fundamental to viable viticulture. The fertile designation indicates successful navigation of multiple critical developmental stages including previous-season inflorescence differentiation (requiring adequate light exposure and balanced vine conditions June-August), current-season flower development (requiring adequate spring growth and resource availability), pollination and fertilization (requiring favorable bloom weather and pollinator activity), fruit set (requiring appropriate environmental conditions and vine capacity), and berry development (requiring continued favorable conditions and adequate photosynthetic support). Factors supporting fertile shoot prevalence include balanced vine conditions (appropriate crop load, adequate vigor, good health), favorable environmental conditions (suitable temperatures, adequate moisture, disease-free), effective management (proper pruning, canopy management, disease control), and variety characteristics (some cultivars showing consistently high fruitfulness and good fruit set). Understanding fertile shoot requirements and optimizing conditions promoting fertile performance is central to consistent commercial production.
fertiler Fruchttrieb
highly productive shoot
hochfruchtbarer Trieb
successful fruiting shoot
well-developed reproductive shoot
Based on Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181.
Fertile Reproductive Shoot
Fertility data for Pinot Noir clone 828 (Dijon) documents bud fruitfulness of 1.9-2.2 inflorescence primordia per bud at positions 3-8 on one-year canes, with only 3-5% blind bud incidence, supporting its suitability for short spur pruning (2-bud spurs on bilateral cordon) while maintaining adequate crop levels of 40-50 clusters per vine. Fertility data for Syrah clone 174 (ENTAV) reveals a pronounced positional fruitfulness gradient — basal buds (positions 1-3) average only 0.8 inflorescences per bud while mid-cane positions (5-10) reach 1.6-1.8, necessitating longer cane pruning (8-12 bud canes on Guyot training) to capture the productive bud zone and avoid the significant crop loss associated with short spur pruning for this clone.
Fertility data is an agronomic performance data entity that records the reproductive capacity of a clonal line, quantified through bud fruitfulness (number of inflorescence primordia per dormant bud), shoot fruitfulness (number of clusters per shoot in the current season), proportion of blind buds (non-bursting buds), proportion of fruitful versus vegetative shoots, inflorescence number per vine, and inflorescence size (flower number per inflorescence), measured across vintages and bud positions along the cane or cordon, characterizing the clone's inherent reproductive potential and its compatibility with specific pruning strategies and crop load management approaches.
Fertility data is essential for matching clone selection to training system and pruning strategy, as the positional distribution of fruitful buds along the cane directly determines which pruning approaches can achieve target crop levels. Bud fruitfulness is established during the previous growing season's inflorescence initiation period (approximately BBCH 53-65 of the preceding year), meaning that environmental conditions 12-15 months before harvest influence the current season's fertility — a physiological lag that connects consecutive vintages through carry-over effects. High-fertility clones facilitate short spur pruning systems (cordon-trained) that offer labor efficiency advantages but may require cluster thinning to prevent overcropping, while low-basal-fertility clones demand cane pruning systems (Guyot variants) that are more labor-intensive but access the productive bud zone. The fruitfulness of basal buds (positions 1-3) versus distal buds (positions 8-12) shows substantial clonal variation within cultivars — this positional profile is a key clone selection criterion in regions transitioning from traditional cane pruning to mechanization-compatible cordon systems. Environmental factors modulating fertility include light exposure during inflorescence initiation (shaded buds produce fewer primordia), temperature during the critical 4-6 week initiation window, and carbohydrate reserve status of the vine. Fertility data should always be reported alongside vigor data, as the two parameters together determine the vine's inherent vegetative-reproductive balance and the management interventions required to optimize it.
Fertilitätsdaten
Fruchtbarkeitsdaten
Gescheinsansatzdaten
bud fertility record
bud fruitfulness data
clone fertility assessment
fruitfulness data
reproductive capacity data
reproduktive Kapazitätsdaten
Sánchez LA, Dokoozlian NK (2005) Bud microclimate and fruitfulness in Vitis vinifera L. American Journal of Enology and Viticulture 56(4): 319-329 Dry PR (2000) Canopy management for fruitfulness. Australian Journal of Grape and Wine Research 6(2): 109-115 Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 145-175
Fertility Data
Cabernet Sauvignon in Napa Valley shows fertilization during mid-late June Year N+1 at BBCH 65-69, with pollen tubes penetrating micropyle and discharging sperm cells 26-38 hours after pollination, double fertilization producing diploid zygotes and triploid endosperm nuclei, 1-3 seeds developing per berry, under warm climate conditions of 24-29°C temperature, adequate boron supporting tube penetration and fusion, typical of this late-season cultivar achieving 58-68% fertilization success and fruit set
Fertilization represents the culminating sexual reproductive process where pollen tube penetrates embryo sac through micropyle, discharges two sperm cells enabling double fertilization with one sperm fusing with egg cell forming diploid zygote initiating embryo development and other sperm fusing with central cell forming triploid endosperm nucleus supporting seed development, establishing successful reproduction and fruit set. This essential fertilization process occurs during late flowering in mid-late June in temperate regions Year N+1, typically occurring 24-48 hours after initial pollination as pollen tube completes growth journey, corresponding to BBCH 65-69 flowering completion stages. Fertilization mechanism involves coordinated processes including pollen tube penetration through micropyle accessing embryo sac, tube discharge releasing two sperm cells into female gametophyte, sperm cell fusion with egg forming diploid 2n zygote, second sperm fusion with central cell containing two polar nuclei forming triploid 3n endosperm nucleus, and initiation of embryo and endosperm development establishing successful seed and berry formation. Pollen tube arrival at micropyle shows culmination of 12-36 hour tube growth journey from stigma through style to ovary, tube navigating to micropyle outer opening guided by ovule-derived attractant signals, and positioning for penetration through micropyle canal to embryo sac requiring final 100-300 micrometer growth. Micropyle structure providing access pathway includes opening in integument layers surrounding ovule, canal 100-300 micrometers length extending from outer to inner integument surface, inner micropyle opening adjacent to nucellus surface providing embryo sac access, and overall architecture creating controlled entry pathway to female gametophyte. Embryo sac structure at fertilization shows mature seven-celled eight-nucleate megagametophyte organization including egg cell positioned at micropylar pole flanked by two synergid cells providing tube reception, large central cell occupying majority of sac volume containing two polar nuclei fusing or separate, and three antipodal cells at chalazal pole opposite micropyle though degenerating by fertilization. Synergid cell function shows specialized reception cells adjacent to egg providing attractant signal guiding tube to micropyle, developing filiform apparatus creating elaborate cell wall ingrowths facing micropyle, and preparing for tube penetration and discharge serving as initial contact point for arriving tube. Pollen tube penetration through micropyle involves tube apex growing through micropyle canal between integument layers, navigating tortuous 100-300 micrometer passage requiring 1-3 hours, reaching nucellus surface adjacent to embryo sac, and penetrating through nucellus wall entering synergid cell completing access to female gametophyte. Tube discharge mechanism shows tube apex penetrating synergid cell and rupturing, releasing two sperm cells into embryo sac cytoplasm, and delivering male gametes positioned for fusion with egg and central cell. Sperm cell characteristics include non-motile cells lacking flagella transported passively in tube cytoplasm, cells containing haploid nucleus and minimal cytoplasm optimized for fusion, and two genetically identical sperm from generative cell mitosis though functioning independently in double fertilization. Double fertilization process shows one sperm cell migrating to and fusing with egg cell creating diploid 2n zygote containing genetic material from both parents, second sperm cell fusing with central cell or polar nuclei forming triploid 3n endosperm nucleus with two maternal and one paternal genome copies, and synchronized fusion events typically occurring within hours of tube discharge. Egg cell fusion creating zygote involves sperm nucleus fusing with egg nucleus combining genetic material, activation of egg development transitioning from arrest to active division, and first zygotic division initiating embryo formation typically within 24-48 hours of fertilization. Central cell fusion forming endosperm shows sperm nucleus fusing with central cell containing two polar nuclei or fused diploid nucleus, creating triploid nucleus with unbalanced genomic ratio affecting endosperm development, and endosperm development initiating providing nutritive support for embryo growth. Fertilization timing shows tube arrival at micropyle typically 18-40 hours after initial pollination depending on style length and temperature, tube penetration and discharge proceeding over 2-6 hours following arrival, and sperm-egg fusion completing within hours of discharge creating successful fertilization typically 24-48 hours post-pollination. Successful fertilization outcomes include viable zygote initiating embryo development, functional endosperm supporting embryo growth, ovule developing into seed, ovary wall developing into fruit flesh, and overall flower transitioning from reproductive to developmental state producing berry. Berry development initiation following fertilization shows rapid ovary expansion from 4-8 mm pre-fertilization to 10-18 mm within weeks, cell division intensity maximizing during early berry growth, resource allocation shifting to developing berries consuming 40-60% photosynthate, and visible fruit cluster formation confirming reproductive success. Auxin production from developing seeds drives berry growth with auxin from embryo and endosperm preventing flower abscission, triggering cell division and expansion in ovary wall, and coordinating berry development with seed formation. Failed fertilization causes include pollen tube failure to reach ovule from inadequate growth vigor or guidance problems, abnormal embryo sac development preventing proper gamete formation, sperm cell dysfunction from pollen quality problems, egg or central cell abnormalities preventing fusion, environmental stress during fertilization period disrupting delicate processes, or genetic incompatibility though rare in self-compatible grapevine. Coulure relationship to fertilization shows poor fruit set from fertilization failure with unfertilized flowers aborting and abscising, environmental conditions during flowering particularly rain or temperature extremes affecting fertilization success, and fruit set percentage 30-75% reflecting fertilization success under varying conditions. Environmental effects on fertilization include temperature affecting tube growth rate and fusion processes with optimal 22-28°C supporting efficient fertilization, boron nutrition essential for tube integrity enabling micropyle penetration, appropriate humidity maintaining tissue function, and overall favorable conditions during 24-48 hour fertilization window critical for success. Temperature sensitivity during fertilization shows optimal 22-28°C supporting normal tube penetration and fusion processes, cool conditions 15-20°C potentially delaying or impairing fertilization reducing success rate, excessive heat above 33°C causing reproductive dysfunction preventing normal fusion, and critical temperature dependence during fertilization period affecting outcomes. Boron absolute requirement emphasizes continued critical importance with boron essential for tube wall integrity during micropyle penetration, deficiency causing tube bursting or penetration failure preventing sperm delivery, and adequate boron 10-20 ppm in reproductive tissues supporting successful fertilization. Multiple tube arrivals per flower show typically 1-4 tubes reaching ovary region, competitive dynamics with usually only 1-2 tubes achieving fertilization, excess tubes unable to fertilize already-fertilized ovules, and single successful fertilization per ovule establishing seed number and berry size. Fertilization competition among ovules shows 2-4 ovules per flower potentially receiving tubes, typically 1-3 ovules successfully fertilizing contributing to final seed number 1-3 per berry, and seed number affecting berry size and development with more seeds generally producing larger berries. The process shows two major phases progressing from pollen tube penetration accessing embryo sac through micropyle and synergid, through ovule penetration culminating in egg approach and tube discharge enabling sperm-egg fusion. Duration typically spans 2-8 hours from micropyle arrival to completed fusion, as final phase of 24-48 hour total reproductive timeline from pollination to fertilization. Cultivar variation in fertilization may include embryo sac development affecting fusion success, ovule number affecting seed production, and fertilization efficiency affecting fruit set under varying conditions. Regional timing shows fertilization during mid-late June in warm climates, late June in temperate regions, and late June-early July in cool areas, occurring 24-48 hours after pollination as culmination of flowering period. Viticultural significance includes representing essential reproductive culmination determining fruit set success, requiring favorable environmental conditions throughout flowering period particularly during final fertilization phase, demonstrating critical dependence on coordinated pollination-germination-growth-fertilization sequence, establishing fertilization success as direct determinant of berry number and yield, and marking transition from reproductive to developmental phase initiating berry growth. Research applications include analyzing fertilization mechanisms including tube-embryo sac interactions and sperm-egg fusion, investigating environmental and genetic factors affecting fertilization success, evaluating double fertilization coordination and timing, quantifying fertilization success rates under varying conditions, assessing seed number effects on berry development, and developing models predicting fruit set from fertilization dynamics and environmental factors.
Fertilization represents culminating sexual reproductive process within Flowering determining fruit set success. Corresponds to BBCH 65-69 stages occurring 24-48 hours post-pollination. Subdivided into Pollen_Tube_Penetration accessing embryo sac and Ovule_Penetration culminating in discharge. Involves double fertilization creating zygote and endosperm. Critical environmental sensitivity particularly temperature and boron. Determines berry number and yield. Marks transition to berry development phase. Final reproductive process completing flowering.
Befruchtung
Gametenfusion
double fertilization
gamete fusion
sexual reproduction
sexuelle Reproduktion
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 388-395 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 293-300 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Fertilization
The fertilization program for a Tempranillo vineyard in Ribera del Duero specifies autumn soil-applied potassium as potassium sulfate (K2SO4) at 120 kg/ha (50 kg K/ha) broadcast and shallowly incorporated by disc cultivator in November to replenish potassium exported in the 7 t/ha grape harvest (estimated K removal of 2.0 kg K per tonne of fruit = 14 kg K/ha/year) and to maintain exchangeable soil K above 150 mg/kg — no nitrogen fertilizer is applied, with vine nitrogen nutrition managed through the biennial compost application (3-4 t/ha) and the crimson clover (Trifolium incarnatum) winter cover crop contributing an estimated 30-40 kg N/ha through biological fixation. Phosphorus is applied only when soil Olsen-P falls below 15 mg/kg, currently at 22 mg/kg and not requiring supplementation. Magnesium supplementation at 40 kg MgO/ha as kieserite (MgSO4·H2O) is applied every third year based on soil analysis indicating marginal exchangeable Mg levels.
A nutrition management description that documents the schedule, composition, rate, placement, and method of soil-applied mineral or organic fertilizer materials, specifying the nutrient sources, application timing relative to vine phenology, target nutrient delivery rates, and the soil-test-based diagnostic rationale that governs routine fertilizer applications designed to maintain adequate soil nutrient reserves for grapevine root uptake throughout the growing season.
Fertilization_Program documents the baseline soil-applied nutrient supply strategy that maintains soil fertility at levels adequate for grapevine requirements without excess. The program is fundamentally informed by two complementary diagnostic tools — soil analysis (providing information about nutrient reserves and availability in the root zone) and tissue analysis (providing information about actual vine nutritional status): discrepancies between soil nutrient levels and vine tissue concentrations indicate uptake limitations caused by soil pH, antagonistic ion interactions (e.g., K-Mg competition, Ca-induced Fe chlorosis), root health problems, or waterlogging that must be addressed through soil amendment or foliar supplementation rather than additional soil fertilization. The nutrient budget approach — calculating the balance between nutrient inputs (fertilizer, compost, atmospheric deposition, irrigation water) and outputs (grape removal, pruning wood removal, leaching, erosion) — provides the quantitative framework for fertilization rate determination, with the objective of maintaining a neutral or slightly positive nutrient balance that sustains soil fertility without accumulation of excess nutrients that could contaminate groundwater or stimulate unwanted vegetative vigor.
Bodendüngung-Plan
Düngungsprogramm
Nährstoffversorgung-Protokoll
nutrient application program
soil fertilization schedule
vineyard fertilizer protocol
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 125-195 White RE (2015) Understanding Vineyard Soils 2nd Ed, Oxford University Press pp: 95-135 Christensen LP and Smart DR (eds) (2005) Proceedings of the Soil Environment and Vine Mineral Nutrition Symposium, ASEV
Fertilization Program
A T-bud of Cabernet Sauvignon inserted into the trunk of an established 110 Richter rootstock in a Napa Valley vineyard bears a Field_Graft_Role, performed in June when bark slips readily, wrapped with budding tape, and allowed to callus in place—the bud pushing growth the following spring to establish the scion variety on the mature root system. A chip bud of Grüner Veltliner serves in a Field_Graft_Role for Austrian nursery production, inserted into one-year-old rooted rootstocks growing in nursery rows during August, with successful takes producing saleable grafted vines the following dormant season. A cleft graft of Merlot onto an established Riparia Gloire trunk bears a Field_Graft_Role in a French vineyard topworking operation, converting an unproductive variety to a higher-value cultivar while retaining the established root system.
A propagation material role that inheres in a grafted combination created in situ on established rootstock plants growing in field conditions, rather than under controlled nursery bench conditions. Field grafts (including T-budding, chip budding, and various field grafting techniques) utilize rootstocks that have been established as rooted plants in vineyard or nursery settings, with scion buds or segments inserted into the actively growing rootstock to establish the fruiting variety. Field grafting enables variety change on established root systems (topworking), vineyard establishment without the expense of nursery-grown bench grafts, and propagation in regions lacking specialized grafting nursery infrastructure. Field graft success depends on timing (coinciding with rootstock cambial activity and bark condition), environmental conditions (temperature, humidity affecting callus formation), scion material quality (bud viability, tissue freshness), and operator skill in making and securing the graft union. Field budding (T-bud or chip bud insertion) represents the most common field grafting approach, performed during the growing season when rootstock bark slips readily.
Field_Graft_Role captures grafting operations performed on established plants rather than dormant bench material, encompassing budding (T-bud, chip bud) and grafting (cleft, bark, whip) techniques applied in field conditions. Field grafting predates bench grafting historically and remains important for topworking (variety conversion on established vines), for regions without nursery infrastructure, and where direct field establishment of rootstocks followed by budding is economically preferred to purchasing bench grafts. Success rates are generally lower than bench grafting (60-80% versus 80-95%) due to environmental variability and the single-attempt nature of most field operations, but the approach enables propagation without specialized facilities. Field graft timing is season-dependent—dormant grafting in late winter, T-budding during active growth when bark slips (typically June-August), chip budding over an extended season—requiring coordination with rootstock phenology and environmental conditions.
Feldveredelung
Freiland-Okulation
Standortveredelung
Umveredelung
field bud
field graft
in-situ graft
topwork graft
vineyard graft
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 468-490 Whiting JR (2004) Grapevine rootstocks, Winetitles Dry N (2007) Grapevine rootstocks, Lythrum Press Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia
Field Graft Role
A breeding line established in replicated field trials at three German sites bears a Field_Selection_Role, with five years of yield data, disease ratings, and wine evaluations determining advancement toward variety registration or elimination from the program. Selected genotypes from marker-assisted seedling screening serve in Field_Selection_Roles following vineyard establishment, with field performance data validating or contradicting molecular predictions. An advanced PIWI breeding line bears a Field_Selection_Role in multi-environment trials across European sites, with broad adaptation confirmed through consistent performance ranking across diverse growing conditions.
A selection role that inheres in a grapevine genotype during vineyard-based evaluation, where established vines are assessed over multiple growing seasons for agronomic performance, disease response, fruit characteristics, and wine quality under field conditions. Field selection represents the extended evaluation phase where breeding material demonstrates performance under realistic cultivation conditions, with assessment spanning the 3-5+ years required for vines to reach mature, stable production. Field selection criteria encompass vegetative characteristics (vigor, growth habit, winter hardiness), reproductive performance (yield, fruit set, cluster architecture), disease response (resistance expression under natural infection pressure), fruit composition (sugar, acidity, phenolics, aromatics), and wine quality (varietal character, balance, aging potential). Multi-site field evaluation enables assessment of genotype × environment interactions, identifying genotypes with broad adaptation versus site-specific performance. Field selection intensity increases progressively—initial field populations may include hundreds of genotypes, reduced through successive evaluation cycles to the elite selections (typically <5% of original population) advanced as variety candidates.
Field_Selection_Role captures the extended evaluation phase that consumes the majority of breeding program timeline and resources, reflecting the perennial nature of grapevines and the multi-year data required for reliable performance assessment. Field selection integrates information unavailable at earlier stages—disease response under natural pressure, adaptation to local conditions, fruit and wine quality from mature vines—providing the comprehensive evaluation necessary for variety release decisions. The field selection phase is rate-limiting for breeding progress even with accelerated molecular selection at seedling stage, field evaluation requires years of vineyard performance data. Strategies to accelerate field selection include early fruit forcing (inducing fruit production on young vines), multi-environment parallel trials (testing across sites simultaneously rather than sequentially), and genomic prediction (using genome-wide markers to predict field performance from molecular data).
Feldselektion
Freilandauslese
Standortselektion
Weinbergselektion
field evaluation
field selection
mature vine selection
vineyard selection
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Töpfer R et al. (2011) Molecular breeding in grapes, CRC Press pp.: 160-185 Reynolds A (ed) (2015) Grapevine Breeding Programs for the Wine Industry, Elsevier Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330
Field Selection Role
Cabernet Sauvignon hermaphrodite flowers at BBCH 60-61 (beginning of flowering) exhibit filaments undergoing rapid elongation from pre-anthesis length of 1.2-1.6 mm to mature erect length of 3.2-3.8 mm within 2-3 hours of calyptra shedding (under morning conditions with temperatures rising from 18°C to 24°C), positioning anthers 0.8-1.3 mm above the pistil (2-2.4 mm) and creating optimal geometry for self-pollination. Gewürztraminer functionally female flowers show strongly reflex filaments 2-2.8 mm that curve sharply downward through 120-160° arc positioning anthers 1.5-2.2 mm below the prominent erect pistil (2.4-2.7 mm), preventing any possibility of self-pollination even if the reduced pollen production were viable. Wild V. vinifera ssp. sylvestris staminate flowers exhibit particularly vigorous erect filaments 3.8-4.5 mm supporting large anthers (0.8-1 mm) positioned well above the vestigial pistil remnant (<0.4 mm), maximizing pollen dispersal height for wind-borne cross-pollination to distant pistillate females. Pinot Noir experiencing water stress during bloom shows reduced filament elongation with mature lengths of 2.6-3.2 mm (versus normal 3.2-3.8 mm for well-watered vines), reducing anther-pistil separation to 0.3-0.9 mm (versus normal 0.7-1.4 mm) but still maintaining adequate positioning for self-pollination and achieving 44-52% fruit set (modestly reduced from normal 48-56%).
A filament is the stalk-like basal portion of the stamen that supports and elevates the anther, extending from the point of attachment to the receptacle to the base of the anther connective, with filament length, orientation (erect versus reflex), and elongation dynamics critically determining anther positioning relative to the pistil and consequently determining flower sexual function and pollination efficiency. The filament structure consists of= (1) a slender cylindrical or slightly flattened stalk typically 0.5-4.5 mm in length and 0.1-0.3 mm in diameter at maturity, though dimensions vary dramatically among flower types (2) a central vascular strand running longitudinally providing water and nutrient transport from receptacle vasculature to the anther (3) parenchymatous cortex surrounding the vascular strand, with cortex cell expansion driving filament elongation (4) epidermal cell layer often bearing trichomes particularly on functionally female and pistillate flower filaments. Filament development proceeds through distinct phases= (1) primordium initiation during early flower bud development (BBCH 51-53) as part of stamen primordium differentiation (2) early elongation (BBCH 53-57) creating distinct separation between developing anther and receptacle, with filaments reaching 0.3-1 mm by late pre-anthesis (3) slow growth phase (BBCH 57-59) with minimal elongation while enclosed by calyptra and while anther pollen maturation proceeds (4) rapid elongation phase (BBCH 60-61) coinciding with calyptra shedding, involving rapid cortex cell expansion that doubles or triples filament length within 1-3 hours, creating the characteristic upward pressure that lifts and sheds the calyptra while positioning anthers for dehiscence (5) post-elongation stabilization (BBCH 61-65) with filaments maintaining erect (hermaphrodite, staminate) or reflex (functionally female, pistillate) positions during active pollination (6) senescence (BBCH 65-69) with browning, loss of turgor, and eventual abscission. Filament length and orientation represent the primary morphological determinants of flower sexual function in Vitis= hermaphrodite flowers exhibit erect filaments 2.5-4 mm at maturity positioning anthers 0.5-1.5 mm above pistil stigma, facilitating efficient self-pollination as pollen falls or is blown from elevated anthers onto the lower stigma functionally female flowers show strongly reflex filaments 1.5-3 mm curved sharply downward positioning anthers 1-2.5 mm below pistil, preventing self-pollination and creating obligate cross-pollination dependency staminate flowers exhibit enhanced erect filaments 3-4.5 mm positioning enlarged anthers well above the vestigial or rudimentary pistil, maximizing pollen dispersal for cross-pollination pistillate flowers show vestigial or rudimentary filaments 0.2-1.5 mm (straight, reflex, or erect) supporting non-functional vestigial anthers. The rapid elongation phase at anthesis involves filament length increases of 100-300% within 1-3 hours, achieved through= (1) rapid cell expansion in cortex parenchyma driven by turgor pressure and cell wall loosening mediated by expansin proteins and auxin signaling (2) water uptake into expanding cells creating the turgor pressure driving expansion (3) oriented cell wall deposition directing expansion longitudinally rather than radially. Environmental conditions during filament rapid elongation affect elongation efficiency and final positioning= optimal conditions (temperatures 20-28°C, adequate water status) support full elongation achieving typical cultivar-specific lengths heat stress >35°C can impair elongation reducing final length by 15-30% water stress can reduce turgor limiting elongation and reducing final length cool temperatures <15°C slow elongation kinetics extending the elongation phase duration and sometimes resulting in incomplete elongation.
Filament represents the stalk component of the stamen and must be distinguished from Anther (terminal pollen-producing structure) and from the complete Stamen (filament plus anther). The filament serves critical mechanical functions (anther elevation and positioning) rather than direct reproductive functions (pollen production), but filament characteristics (length, orientation, elongation dynamics) are primary determinants of flower sexual function through their control of anther-pistil spatial relationships. Sub-classification into Long_Filament, Short_Filament, and Erect_Filament reflects morphological and functional variations related to flower sexual types and pollination strategies. Filament characteristics provide key diagnostic features for identifying flower types and for understanding reproductive biology across the spectrum of Vitis sexual expression.
Filament
Staubblattfaden
Staubfaden
anther support
stamen stalk
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 99-104 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 93-98 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644.
In Viognier berries in Condrieu, Rhône Valley during BBCH 89, final acid balance stabilizes at titratable acidity of 5.4 g/L and pH 3.52 during the final 10 days before harvest, with malic acid equilibrating at 1.6 g/L and tartaric acid at 5.1 g/L, representing daily acid change of only 0.04 g/L compared to mid-ripening rates of 0.35 g/L, under late-season temperatures averaging 26°C.
Final acid balance constitutes the terminal phase of organic acid metabolism in Vitis vinifera berries wherein acid reduction processes approach equilibrium and titratable acidity stabilizes at harvest-appropriate levels of 5-9 g/L during the final 7-14 days before picking, representing the culmination of ripening-phase acid dynamics and establishing the definitive acid composition that will characterize the resulting wine following vinification. This process emerges as malic acid degradation rates decline asymptotically due to substrate depletion, with daily acid loss decreasing from active-phase rates of 0.2-0.5 g/L per day to equilibrium rates of 0.02-0.08 g/L per day as vacuolar malate pools approach minimum sustainable concentrations of 1-3 g/L in most cultivars. The attainment of final acid balance integrates the outcomes of malate catabolism and tartrate stability, producing characteristic harvest acid profiles comprising residual malate of 1-4 g/L and stable tartrate of 5-8 g/L that together constitute the 6-10 g/L titratable acidity typical of mature wine grapes across diverse growing regions. Temperature during the final ripening phase continues to influence the precise endpoint of acid balance, with warm conditions of 28-35°C driving malate to lower residual levels of 1-2 g/L compared to cool conditions of 15-20°C where final malate may stabilize at 3-5 g/L, producing the characteristic acidity differences between warm and cool climate wines. The pH at final acid balance typically ranges from 3.2-3.6 for white varieties and 3.3-3.8 for red varieties, reflecting the combined effects of organic acid composition, potassium content, and cultivar-specific buffering capacity, with pH providing critical information for winemaking decisions regarding acidification requirements, malolactic fermentation management, and sulfur dioxide additions. Cultivar variation in final acid balance reflects genetic differences in malate degradation endpoint, tartrate biosynthesis capacity, and potassium accumulation patterns, with high-acid varieties such as Riesling maintaining titratable acidity of 8-12 g/L at full maturity compared to low-acid varieties such as Viognier achieving balance at 5-7 g/L under equivalent conditions. The stability of final acid balance once achieved enables flexibility in harvest timing decisions, as the plateau phase allows picking windows of 7-14 days during which acid composition changes minimally while other quality parameters continue developing.
Final_Acid_Balance is parent class under Acid_Reduction representing the terminal equilibrium phase. Distinguishes from Active_Acid_Reduction which represents dynamic catabolism phase. Contains subclasses Mature_Berry_Acidity, Sugar_Acid_Ratio_Optimization, and Harvest_Acidity_Level. Duration 7-14 days at end of ripening. Key characteristic is asymptotic approach to stable acid levels with minimal daily change.
Ernte-Säure-Balance
finale Säure-Balance
harvest acid balance
mature acid composition
reife Säure-Zusammensetzung
ripening acid endpoint
terminal acid equilibrium
terminales Säure-Gleichgewicht
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 275-280 Coombe BG (1992) Research on development and ripening of the grape berry. American Journal of Enology and Viticulture 43: 101-110 Conde C et al (2007) Biochemical changes throughout grape berry development. Food Chemistry 1: 1-22
Final Acid Balance
Syrah in Rhône Valley shows final cap fall during late June Year N+1 at BBCH 69 on approximately June 15-16, with last 4-7% flowers completing cap detachment predominantly in basal positions, visible berry development 10-16 mm diameter throughout cluster representing 52-64% fruit set, unfertilized flowers showing active abscission, under warm Mediterranean conditions of 24-28°C temperature, typical of this mid-to-late season cultivar completing cap fall 11-12 days after first detachment and transitioning definitively to berry development phase
Final cap fall represents the concluding calyptra detachment phase when last remaining flowers complete abscission establishing complete cap shedding within cluster and marking definitive flowering completion with no further reproductive structures to contribute, transitioning fully to berry development focus. This conclusive shedding phase occurs during complete anthesis in late June-early July in temperate regions Year N+1 following progressive cap fall, typically spanning 1-2 days as final flowers complete detachment, corresponding to BBCH 69 approaching complete flowering stage. Completion characteristics show all viable flowers having completed cap shedding with 100% developmentally capable flowers showing detached caps or unopened structures representing aborted buds, minimal ongoing cap fall activity with only final 3-8% flowers completing detachment, and definitive cessation of calyptra shedding activity across cluster. Final flower cap shedding shows last individual flowers completing abscission typically in basal cluster positions or scattered locations representing most delayed development, cap detachment occurring through same abscission mechanisms as earlier flowers though under potentially suboptimal late conditions, and completion representing end of flowering-related cap fall with no further flowers capable of contributing. Cap accumulation at completion reaches maximum density with total fallen cap debris representing full flowering extent, accumulated caps on cluster surfaces and ground below documenting complete flowering period, and cap debris beginning decomposition processes creating organic matter return. Cluster appearance at final cap fall shows definitive developmental state with small developing berries 8-15 mm diameter dominating visual appearance from successful early-to-mid fruit set, brown senescing unfertilized flowers interspersed showing varying abscission progress, occasional final-opening flowers with minimal reproductive potential, and overall appearance of developmental phase with reproductive activity concluded. Berry development visibility at completion shows clear green fruit structures distributed throughout cluster creating recognizable young fruit cluster appearance, berry number countable enabling definitive fruit set assessment, spatial distribution revealing architectural pattern and final cluster density, and overall berry-dominated appearance contrasting with earlier flower-dominated state. Unfertilized flower abscission at completion shows active progressive detachment with senescing flowers developing completed abscission zones enabling separation, daily flower drop creating accumulation of fallen brown flower debris, and progressive cluster cleaning eliminating failed reproductions over subsequent 1-2 weeks leaving only productive berries. Abscission completion timing shows most unfertilized flower drop occurring during 1-2 weeks following final cap fall, though some dried flower remnants may persist longer before eventual removal, and overall cluster achieving predominantly berry-only appearance by post-completion period. Retained cap problems at completion may show occasional flowers with incompletely shed caps representing failed abscission from developmental abnormalities or unfavorable conditions, retained caps potentially interfering with berry development if fruit set succeeded despite retention, and overall retention assessment indicating flowering efficiency and potential coulure extent. Flowering completion confirmation requires verification that no further viable unopened flowers remain capable of contributing to fruit set, assessment that reproductive activity has definitively ceased across cluster, and recognition that final cap fall represents true flowering conclusion rather than temporary lull. Developmental transition at completion shows cluster fully committed to berry development phase with all metabolic resources directing to expanding berries, no residual reproductive investment or potential, and physiological state completely transitioned from reproductive to developmental focus. Berry growth acceleration following completion shows rapidly increasing expansion rates as cell division intensifies, daily diameter increase of 0.4-0.8 mm during early rapid growth, and visible berry size progression creating obvious fruit development. Resource allocation at completion shows complete reorientation to berry development as dominant sink consuming 40-60% total photosynthate, vegetative growth continuing requiring 30-40% resources, and no reproductive demands with flowering completely concluded. Phenological significance at completion includes marking definitive end of BBCH 69 cap fall and flowering stages, establishing transition to BBCH 71-79 fruit development stages, and providing clear observable reference for flowering conclusion enabling developmental phase focus. Management implications at completion include confirming flowering conclusion enabling complete resumption of normal operations without bloom-period restrictions, initiating post-bloom fruit development management including disease protection and crop adjustment, and transitioning focus entirely to developmental and quality optimization priorities. Fruit set determination at completion enables accurate assessment through berry count establishing final berry number, calculation of fruit set percentage from berry number relative to total flowers opened, and yield prediction based on berry number and expected final berry weight. Observational characteristics enable definitive completion identification through cessation of cap fall activity with no ongoing detachment visible, cluster appearance showing predominantly berries with minimal remaining flowers, and overall transition to developmental state clearly evident. The process shows two distinguishable aspects progressing from last caps falling completing final detachments, through cap fall completion confirming definitive cessation. Duration represents brief 1-2 day final period marking completion, with rapid transition to fully developmental state. Cultivar variation in final cap fall shows differences in completion timing with early varieties completing late June while late types complete early-mid July, retention problems with some varieties showing occasional persistent caps while others show complete clean shedding, and completion clarity with some showing definitive conclusion while others showing gradual extended completion. Regional timing shows final cap fall during late June in warm climates at approximately 1150-1200 growing degree days base 10°C, late June-early July in temperate regions at 1050-1100 GDD, and early-mid July in cool areas at 950-1000 GDD, marking flowering completion across regions. Viticultural significance includes marking definitive flowering completion enabling complete management transitions, confirming final fruit set and berry number establishing yield potential, indicating end of critical flowering period reducing vulnerability and restrictions, and providing clear observable reference for developmental phase transition. Research applications include documenting precise flowering completion timing for phenological studies, analyzing factors affecting complete cap shedding achievement, evaluating relationships between cap fall patterns and fruit set outcomes, assessing retention problems affecting flowering completion and fruit set, and documenting phenological transitions marking major phase changes in reproductive biology.
Final_Cap_Fall is fourth and conclusive major Cap_Fall phase marking definitive calyptra shedding completion. Corresponds to BBCH 69 parallel to Complete_Anthesis. Subdivided into Last_Caps_Falling and Cap_Fall_Completion. Establishes flowering conclusion through observable cap shedding cessation. Enables management transitions and fruit set assessment. Marks transition to developmental phase. Provides clear practical observable marker for flowering completion easier than detailed reproductive organ assessment.
Blüteende-Kappenabwurf
Kappenfall-Abschluss
abschließende Kalyptralösung
cap fall completion
concluding cap shedding
final calyptra detachment
finaler Kappenfall
flowering completion caps
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 388-395 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Final Cap Fall
Pinot Noir in Burgundy shows final cell expansion during mid-late August Year N+1 at BBCH 79, with cells completing expansion adding final 4.2% volume reaching 100% final dimensions at 90-106 micrometers diameter, vacuoles 91% cell volume, water uptake declining from 0.5 to 0.2 g per berry per day, expansion rate <1.5% per week approaching cessation, under continental conditions of 22-28°C temperature, typical of this early-to-mid season cultivar completing cell sizing
Final cell expansion represents the terminal enlargement phase during enlargement phase completion when cells undergo last minimal volume increases adding final 2-8% to achieve 98-100% mature dimensions, expansion rate very low completing cell sizing, cells reach ultimate volumes 80-200 micrometers diameter depending on cultivar, and final expansion establishes definitive cell dimensions persisting through ripening and harvest, occurring during late enlargement weeks 7-9 as cell growth concludes. This terminal expansion phase occurs during late summer in mid-late August in temperate regions Year N+1, proceeding over 1-2 weeks representing final enlargement, corresponding to BBCH 79 majority berries touching stage. Final expansion mechanism involves continued minimal water uptake adding last volume increment, vacuole reaching final 88-93% cell volume maximum, sustained minimal turgor enabling limited expansion, reduced wall yielding allowing minimal growth, and eventual expansion cessation establishing final cell dimensions. Cell volume increase during final expansion shows typical 2-8% volume gain from late maximum enlargement to completion, small absolute increase adding final micrometers to cell diameter, cumulative 30-100 fold total expansion from post-division cells, and final increase completing expansion program. Cell diameter progression shows expansion from 92-96% final size during late maximum enlargement to 98-100% at final expansion completion, typical progression from 75-185 micrometers to final 80-200 micrometers, final diameter establishing permanent cell dimensions, and completion sizes showing cultivar ranges. Cell volume finalization shows cells achieving final mature volumes 0.2-4.2 nanoliters depending on size, volumes representing culmination of expansion program, finalized volumes determining cell contribution to berry size, and final volumes unchanging through subsequent development. Vacuole final dimensions show central vacuole reaching 88-93% final cell volume maximum, vacuole diameter 70-190 micrometers in large cells, vacuole volume representing major cell volume component, tonoplast membrane area 1500-7000 square micrometers, and vacuole at maximum development. Cytoplasm final configuration shows thin peripheral layer 1.5-3 micrometers at completion, cytoplasm volume minimized in fully expanded cells, maintained metabolic function in compressed space, nucleus and organelles in peripheral cytoplasm, and efficient final organization. Water uptake during final expansion shows continued minimal water influx completing cell hydration, typical uptake rates 0.3-0.8 g per berry per day during final expansion declining to <0.3 g per day at completion, water adding final volume increment, and minimal uptake reflecting expansion cessation. Osmotic potential during final expansion shows values -1.2 to -1.6 MPa maintained near steady state, minimal additional solute accumulation, osmotic gradient approaching equilibrium, and plateau characteristic of expansion cessation. Turgor pressure during final expansion shows sustained pressure 0.5-0.9 MPa enabling minimal expansion, turgor maintained but insufficient for substantial growth from wall constraints, pressure-expansion coupling minimal with low yielding, and turgor alone inadequate for expansion without wall plasticity. Wall properties during final expansion shows walls largely non-yielding with high yield threshold 0.7-1.0 MPa, minimal wall plasticity limiting expansion, wall thickness 0.12-0.28 micrometers at completion from continued synthesis, wall mechanical properties growth-restrictive, and wall as physical constraint ending expansion. Wall composition at completion shows mature wall with extensive cellulose microfibril network, cross-linked hemicellulose and pectin matrix, possible calcium-mediated pectin associations, and mature composition limiting plasticity. Expansion rate during final expansion shows very low rate producing minimal volume gain, typical expansion velocity <2% volume per week, declining rate approaching zero, and low rate characteristic of expansion cessation. Cell expansion cessation timing shows expansion effectively ceasing by week 8-10, occasional minor expansion possible during lag and ripening adding <2% volume, essential cessation establishing final dimensions, and timing varying slightly by cultivar and environment. Final cell dimensions show cultivar-specific size ranges achieved at completion, small-berried cultivar cells 60-105 micrometers final diameter, medium-berried 90-145 micrometers, large-berried 120-200 micrometers, and final sizes determining cell contribution to berry volume. Cell size variation shows spatial variation with inner mesocarp cells typically larger than outer, position-dependent sizes reflecting growth gradients, cell type affecting final dimensions with parenchyma largest, and within-berry heterogeneity in final cell sizes. Environmental effects on final expansion include water availability during final phase affecting last volume gain though impact modest <5%, temperature affecting completion timing, and environmental factors having minimal impact on final dimensions as genetic programming dominates. Water effects show adequate water beneficial for completing final expansion, severe deficit potentially limiting final 2-5%, and water less critical than during peak expansion. Temperature effects show warm conditions promoting faster completion, cool conditions extending final expansion period, and temperature affecting timing rather than achieved final size. Final expansion completion indicators show expansion rate declining to near-zero <0.5% volume per week, cell dimensions stable over time, and completion establishing final cell size. Cell size permanence shows finalized dimensions unchanging through lag phase and ripening, minor volume changes <2% possible from osmotic fluctuations, and essential stability of achieved dimensions. The process represents first aspect of enlargement phase completion establishing final cell dimensions, preceding pre-lag phase transition and growth deceleration. Duration typically 1-2 weeks representing terminal expansion weeks 7-9. Regional timing shows final cell expansion during mid-late August across climates completing cell sizing. Viticultural significance includes establishing final cell dimensions determining cellular contribution to berry size, completing 30-100 fold cell expansion program, demonstrating expansion cessation as developmental endpoint, and determining final cell architecture affecting berry quality. Research applications include analyzing final expansion mechanisms and cessation, investigating cell size determination and limits, evaluating environmental effects on final dimensions, quantifying final cell size ranges and determinants, and assessing final expansion as determinant of ultimate berry size.
Final_Cell_Expansion is first aspect of Enlargement_Phase_Completion, distinguished by terminal cell sizing. Represents last 2-8% volume increase reaching 98-100% final dimensions. Very low expansion rate declining to near-zero. Final cell diameters 80-200 micrometers. Vacuole 88-93% volume maximum. Wall largely non-yielding (threshold 0.7-1.0 MPa). Water uptake <0.3 g/berry/day at completion. Duration 1-2 weeks. Followed by Pre_Lag_Phase_Transition. Establishes permanent cell dimensions (±2% through ripening).
Ultimate Zellexpansion
completion expansion
final cell sizing
finale Zellexpansion
finale Zellgröße
terminal cell growth
terminales Zellwachstum
ultimate cell expansion
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 451-453 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Final Cell Expansion
In a Brunello di Montalcino Sangiovese vineyard, final color expression was established 16 days post-color break in early October Year N+1 (BBCH 87) showing CIE L* 33.5 (±1.2 over 3 weeks), hue angle 6.8° (±2.1° variation) indicating characteristic Sangiovese red-orange color, chroma 42.8, skin anthocyanins 1085 mg/kg (malvidin-3-glucoside 58%, cyanidin-3-glucoside 19%, delphinidin-3-glucoside 7%), berry Brix 24.3°, titratable acidity 6.4 g/L, under typical Tuscan autumn conditions (daily high 21-24°C, night 10-13°C, moderate deficit) with southern hillside exposure.
Final_Color_Expression represents the stable endpoint of berry pigmentation when anthocyanin composition, copigmentation, and cellular architecture establish the characteristic cultivar-specific color that persists through harvest providing the definitive visual quality marker for varietal identification, ripeness assessment, and wine color potential prediction. This final expression emerges during late veraison to ripening (BBCH 85-89), typically 12-18 days post-color break, and is characterized by stable colorimetric values maintained ±8-12% over 2-4 weeks: CIE L* (lightness) stabilizing at cultivar-typical values (Petit Verdot 22-26, Syrah 25-29, Cabernet Sauvignon 27-31, Merlot 29-33, Pinot Noir 32-37), hue angle defining cultivar color signature (pure reds 0-12°, red-purple 345-0°, purple-red 335-345°), and chroma representing saturation level (28-56 depending on variety and conditions) creating the distinctive color profile enabling varietal recognition and quality assessment. Anthocyanin compositional basis of final expression includes: malvidin-3-glucoside proportion (48-72% total anthocyanins) establishing base color intensity and hue, delphinidin-3-glucoside content (4-22%) contributing purple-blue notes (elevated in Syrah 14-20%, Petit Verdot 16-24% creating characteristic purple color, lower in Merlot 6-10%, Pinot Noir 5-8% producing redder hues), cyanidin-3-glucoside levels (6-25%) adding bright red character (high in Pinot Noir 18-28% creating cherry-red color, moderate in others 8-15%), peonidin and petunidin derivatives (5-15% each) contributing to hue complexity and color stability, and acylated anthocyanins (20-35% of total) providing enhanced color persistence through intramolecular copigmentation and pH-buffering effects creating more stable final expression. Copigmentation phenomena stabilizing final expression involve: flavonol copigments (quercetin, kaempferol, myricetin glycosides 150-350 mg/kg skin) creating molecular stacking with anthocyanins producing 40-55% color intensity enhancement through bathochromic shifts (absorbance maximum moving from 520 nm to 530-540 nm) and hyperchromic effects (increased extinction coefficient), hydroxycinnamic acid copigments (caffeic, coumaric, ferulic acids 80-220 mg/kg) contributing additional 15-25% intensity enhancement, and self-association of anthocyanins at high concentrations (>15 mM vacuolar) creating aggregates with modified spectral properties intensifying color expression beyond dilute solution predictions. Cellular architecture effects on final expression inclu multilayered pigmented epidermis and hypodermis (3-6 cell layers deep cultivar-dependent) creating optical depth amplifying color intensity through extended light path length, vacuolar pH stabilization at 2.8-3.3 maintaining optimal flavylium cation to carbinol ratio (85-95% colored form), anthocyanin vacuolar inclusions in some cultivars (Cabernet Sauvignon, Merlot) concentrating pigments >60 mM locally creating regions of exceptional intensity contributing to overall color depth, and cell wall-bound phenolics providing additional light absorption and color background supporting anthocyanin-based primary coloration. Optical characteristics of final expression show: reflectance spectra displaying cultivar-specific profiles with characteristic red reflection peak 620-650 nm (intensity 30-55% depending on anthocyanin level), purple-blue secondary reflection 400-450 nm (15-45% in delphinidin-rich varieties, 32°C reducing final expression quality through anthocyanin degradation and compositional shifts potentially producing atypical pale or brownish colors, UV-B exposure enhancing expression through increased anthocyanin synthesis (30-50% higher levels) and potentially enhanced copigment accumulation intensifying final color, and water deficit history with moderate sustained stress producing more intense expression (30-45% deeper color) through berry concentration and elevated biosynthesis while severe deficit may compromise expression through metabolic limitation. Cultivar genetic variation creates distinctive final expressions: Cabernet Sauvignon shows deep purple-red (L* 27-31, hue 352-358°) from balanced delphinidin and malvidin, Merlot displays intense red-purple (L* 29-33, hue 355-2°) with moderate delphinidin creating slight purple note, Pinot Noir exhibits bright cherry-red (L* 32-37, hue 5-12°) from elevated cyanidin and lower delphinidin producing pure red without purple, Syrah shows deep purple-red (L* 25-29, hue 340-348°) from high delphinidin creating pronounced purple character, Grenache displays medium red (L* 33-38, hue 8-15°) from moderate anthocyanins with red-dominant hue, and Petit Verdot achieves very deep purple-black (L* 22-26, hue 335-345°) from extremely high anthocyanins with elevated delphinidin. Temporal stability of final expression varies: under favorable conditions (moderate temperatures 18-26°C, no environmental extremes) final expression persists 3-6 weeks enabling flexible harvest timing particularly in stable cultivars (Cabernet Sauvignon, Syrah, Sangiovese), moderate stability exists in varieties like Merlot and Tempranillo maintaining expression 2-4 weeks, while unstable cultivars (Pinot Noir in warm climates, Grenache) may deteriorate 1-2 weeks post-final expression through anthocyanin degradation, copigment losses, or cellular senescence creating color shifts toward brown-red or pink-red indicating over-ripeness. Wine quality prediction from final expression: berry color intensity strongly correlates with wine color density (r² = 0.88-0.95) enabling harvest quality forecasting, hue angle predicts wine color character with low angles producing red-dominant wines suitable for young consumption while purple hues indicate age-worthy wines with anthocyanin-tannin complexation potential, color uniformity across berries predicts wine color consistency and stability important for premium production, and comparison of actual versus expected final expression for cultivar and vintage reveals environmental or management effects requiring attention. Harvest timing optimization using final expression involves: visual assessment comparing berry color to cultivar-specific references indicating typical ripe expression suitable for harvest, colorimetric monitoring tracking stabilization of L*a*b* values indicating color development completion, integration with chemical maturity (sugar, acid, pH) and sensory evaluation (flavor, tannin) ensuring color expression synchronized with overall ripeness, and style considerations with some wines requiring harvest before complete final expression (lighter styles, rosé from red varieties) while others benefit from full expression development (premium reds, age-worthy wines). Viticultural factors affecting final expression achievement inclu canopy management optimizing cluster light exposure without excessive heat enabling full expression development, irrigation strategy imposing moderate deficit during veraison enhancing expression intensity 25-40%, crop load adjustment to 6-10 kg/vine depending on variety improving expression through adequate resource allocation, disease prevention maintaining berry health enabling complete biosynthetic programs, and harvest logistics planning based on expression stability characteristics with stable varieties allowing extended decision windows while unstable types requiring prompt harvest upon expression achievement. Marketing and quality designation implications: final expression provides visual quality assessment enabling premium lot selection, color intensity and typicity influence wine classification and pricing with deeper more characteristic expression commanding premiums, and varietal authenticity verification using color characteristics detecting potential mislabeling or blending issues important for appellation compliance and brand integrity.
Final_Color_Expression is distinguished from Maximum_Color_Intensity by representing stable characteristic color rather than peak intensity, from Complete_Color_Change by emphasizing cultivar-specific expression rather than completion per se, and from Color_Development_Completion by focusing on phenotypic endpoint rather than process termination. Stability criterion: colorimetric values ±8-12% over 2-4 weeks. Timing: typically 12-18 days post-color break. Persistence: stable cultivars 3-6 weeks, moderate 2-4 weeks, unstable 1-2 weeks. Final_Color_Expression provides definitive quality marker for harvest timing and wine style prediction.
characteristic cultivar pigmentation
charakteristische Kultivar-Pigmentierung
definitive Beeren-Farbe
definitive berry color
stabile Sorten-Farbe
stable varietal color
terminal color phenotype
terminaler Farb-Phänotyp
Keller M (2020) The Science of Grapevines 3rd Ed pages 236-240 cultivar color characteristics and harvest González-Neves G, Gil G, Favre G, Ferrer M (2015) Influence of grape composition and winemaking on the anthocyanin composition of red wines J Food Comp Anal 41:124-132 anthocyanin composition effects on color Mazzuca P, Ferranti P, Picariello G, Chianese L, Addeo F (2005) Mass spectrometry in the study of anthocyanins and their derivatives J Mass Spectrom 40:83-90 anthocyanin characterization.
Final Color Expression
Pinot Noir in Burgundy shows final flowers opening during late June Year N+1 at late BBCH 68 over 1-2 days June 14-15, with remaining 12-18% flowers predominantly in basal cluster positions completing cap shedding, occurring while 3-5 mm berries visible from earliest fruit set, limited pollen availability constraining late pollination success to 22-32%, under continental conditions of 21-26°C temperature, typical of this early-to-mid season cultivar completing flowering with final opening contributing 10-14% berry number
Final flowers opening represents the completion of cluster flowering activity when remaining 10-20% unopened buds complete cap shedding and expose reproductive organs, establishing last opportunities for pollination and fruit set before anthesis conclusion. This concluding opening phase occurs during late anthesis in mid-late June in temperate regions Year N+1, typically spanning 1-2 days as final flowers predominantly in basal cluster positions achieve open state, corresponding to late BBCH 67-68 stages. Opening pattern shows final flowers concentrated in basal cluster regions reflecting latest primordium formation and most delayed development, with 10-20% flowers remaining unopened at late anthesis onset completing opening during this final phase, and very occasional flowers showing failure to open remaining as unopened buds that will eventually abscise. Cap shedding in final flowers proceeds through same abscission mechanisms as earlier flowers but often under less favorable conditions with declining flower vigor, potentially suboptimal environmental conditions after peak period, and reduced developmental resources as vine prioritizes developing berries, creating somewhat less efficient opening in some cases. Reproductive organ exposure in final-opening flowers reveals functional structures though vigor may be reduced compared to earlier flowers, with anthers showing appropriate maturation and dehiscence capacity though pollen production may be slightly reduced, and stigmas developing receptivity though exudate production may be somewhat diminished. Flower appearance at final opening shows opened structures similar to earlier flowers but opening timing indicates late developmental status, with opened final flowers distinguishable from earlier-opened flowers through spatial position predominantly basal, timing relative to fruit development elsewhere in cluster, and occasionally slightly smaller organ dimensions. Timing relationships show final opening occurring simultaneously with advanced development elsewhere in cluster including visible small berries 3-6 mm from earliest fruit set, senescing unfertilized flowers from mid-flowering showing brown discoloration, and declining reproductive activity in earlier-opened flowers, creating maximum developmental heterogeneity. Pollen availability for final-opening flowers shows suboptimal conditions with limited fresh pollen from own anthers providing immediate source, minimal viable pollen remaining from earlier-opened flowers with most aged pollen expired, and reduced atmospheric pollen concentration as cluster-level pollen production declining substantially. Stigma function in final-opening flowers develops normally with receptivity initiating within 6-24 hours after opening and reaching moderate peak levels, though overall pollination opportunity is constrained by limited pollen availability rather than stigma function, with receptive period potentially abbreviated compared to earlier flowers. Pollination probability for final-opening flowers shows reduced success with estimated 15-35% achieving successful pollination and fruit set compared to 50-70% during optimal window, resulting primarily from limited pollen availability rather than stigma dysfunction, affecting final fruit set contribution from these latest flowers. Self-pollination remains primary mechanism for final flowers with pollen from own anthers providing main opportunity, limited cross-pollination from neighboring late-opening flowers supplementing, and minimal contribution from aged pollen from earlier flowers, creating dependence on immediate pollen production. Developmental outcomes from final opening show variable fruit set with some late-opening flowers achieving successful pollination and contributing to final berry number, while others fail to set becoming part of abscising unfertilized flower population, with late-set berries potentially showing delayed development affecting ripening uniformity. Berry development from final pollinations proceeds normally but with temporal offset showing berries from final opening developing 5-10 days behind earliest-set berries, creating developmental gradient within cluster affecting ripening progression and harvest timing considerations. Environmental effects on final opening continue with temperature affecting cap shedding efficiency and organ function, humidity influencing anther dehiscence and pollen release, and overall conditions often less optimal than during peak period potentially contributing to reduced opening vigor. Weather impacts show that adverse conditions during final opening have limited overall effect on fruit set since majority 75-85% berry number already established, though conditions do affect final opening success and contribution to ultimate yield. Cluster architecture effects show that basal flowers opening latest may experience more shaded microclimate with reduced light and air circulation potentially affecting opening efficiency and subsequent reproductive success compared to earlier-opening better-exposed apical flowers. Resource competition increases during final opening with developing berries from earlier fruit set creating substantial sink demand competing with reproductive processes in final-opening flowers, potentially contributing to reduced late flower success through resource limitation. Metabolic status in final-opening flowers shows adequate function but possibly reduced vigor with respiration supporting reproductive processes though potentially at lower levels than earlier flowers, and carbohydrate availability adequate though preferentially allocated to developing berries creating possible resource constraint. Hormonal environment during final opening shows declining reproductive hormone levels as flowering culminates with increasing dominance of developmental hormones from successfully fertilized ovaries, creating hormonal context less favorable for sustained reproductive investment. Developmental coordination shows final opening occurring while vegetative growth continues with shoot elongation ongoing though slowing, leaf expansion proceeding, and tendrils developing, maintaining concurrent vegetative-reproductive processes though shifting toward vegetative-developmental balance. Management implications of final opening include indicating approaching flowering completion enabling planning for post-bloom operations, confirming extent of cluster flowering activity affecting yield predictions, and providing final assessment of flowering uniformity and potential developmental heterogeneity. Observational characteristics enable identification of final opening through spatial concentration in basal positions, timing relative to berry development elsewhere in cluster, and progression toward complete cluster flowering creating all-flowers-opened state. The process represents first of three late anthesis aspects focusing on completion of opening activity, followed by declining pollen viability and pollination window closure affecting reproductive opportunity. Duration typically spans 1-2 days representing brief concluding opening period, with warm conditions promoting rapid completion while cool conditions may extend period. Cultivar variation appears in proportion of final opening with some varieties showing minimal 8-12% late opening reflecting compressed flowering while others show substantial 18-25% late opening indicating extended flowering, final flower vigor with some maintaining good reproductive capacity while others show substantially reduced capacity, and timing relative to fruit development affecting developmental heterogeneity. Regional timing shows final flowers opening during mid-late June in warm climates, late June in temperate regions, and late June-early July in cool areas, occurring 6-9 days after first flowers opening depending on overall flowering duration. Viticultural significance includes completing cluster flowering activity establishing all potential fruit set sites, contributing final 8-15% berry number from late successful pollinations, marking transition toward complete anthesis and flowering conclusion, and establishing maximum within-cluster developmental heterogeneity affecting subsequent ripening patterns. Research applications include analyzing factors determining final flower opening success, evaluating developmental constraints on late flower reproductive capacity, quantifying fruit set contributions from final opening affecting yield components, assessing resource allocation conflicts between late reproduction and early berry development, and documenting within-cluster developmental gradients affecting ripening dynamics.
Final_Flowers_Opening is first of three Late_Anthesis aspects, distinguished by completion of cluster opening activity with remaining unopened buds. Occurs predominantly in basal cluster regions. Represents final pollination opportunity contributing 8-15% berry number. Constrained by limited pollen availability rather than stigma dysfunction. Establishes maximum within-cluster developmental heterogeneity. Marks transition toward complete anthesis.
Endblüten
abschließende Blütenöffnung
concluding bloom
final bud opening
finale Knospenöffnung
last flowers opening
letzte Blüten öffnen
terminal flowering
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 382-386 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Final Flowers Opening
Merlot in Bordeaux shows final mitotic events during late July-early August Year N+1 at BBCH 75, with last dividing cells representing 8-14% population week 4 declining to 2-4% week 5, terminal divisions proceeding with 16-18 hour extended cycle times, mitotic index declining from 2.8% to 0.6%, final divisions adding 180,000-240,000 cells representing 9-12% total cellularity, completing permanent 2.0-2.3 million cell count, under oceanic conditions of 22-27°C temperature
Final mitotic events represent the terminal division occurrences during division phase completion when last remaining cycling cells complete final divisions, mitotic index declines to very low <1-2% indicating rare divisions, late-dividing cells undergo extended cycle times 16-18 hours reflecting reduced proliferative capacity, and terminal divisions produce final cells completing total berry cellularity, occurring during weeks 4-5 as last proliferative activity. This terminal division period occurs during late summer in late July-early August in temperate regions Year N+1, representing final division activity weeks 4-5, corresponding to BBCH 75 berry touch stage. Final events mechanism involves completion of ongoing cycles in remaining late-dividing cells, extended cell cycle times reflecting declining proliferative stimulus, terminal divisions producing last cells exiting to expansion state, mitotic index decline to minimal levels <1-2%, and eventual complete cessation establishing non-dividing population. Last dividing cell populations show remaining cycling cells representing 10-20% total cells during early final events week 4, progressive cycle exit reducing cycling fraction to 5-10% late week 4 and <5% week 5, and final dividing cells representing late-activated or slow-cycling populations. Extended cycle times show late divisions proceeding with 16-18 hour cycles versus 10-12 hours during peak, G1 phase extension to 7-10 hours indicating reduced progression stimulus, S phase extension to 8-10 hours showing slowed DNA replication, and overall cycle lengthening characteristic of declining proliferative state. Cycle extension mechanisms show reduced auxin levels weakening proliferative drive, checkpoint stringency possibly increasing delaying phase transitions, reduced biosynthetic capacity slowing S phase progression, and developmental signals directing cell cycle exit over continued division. Terminal division characteristics show divisions morphologically normal with proper chromosome segregation, cytokinesis completing producing viable daughter cells, daughter cells entering expansion state rather than cycling, and terminal divisions functional though infrequent. Mitotic index during final events shows percentage of cells in M phase declining to very low levels, typical progression from 3-5% early final events to 1-2% late final events to <1% at complete cessation, mitotic index accurately reflecting rare remaining divisions, and minimal index indicating near-complete division cessation. Division rate during final events shows new cell generation rate declining to very low 2,000-10,000 cells per berry per day versus 40,000-120,000 at peak, rate reduction reflecting both minimal cycling fraction and extended cycle times, and minimal rate representing final proliferative output. Cell number addition from final events shows terminal divisions adding 5-15% total cells to earlier accumulated populations, contribution relatively small compared to peak activity but completing final cellularity, and final additions establishing permanent cell count. Spatial distribution of final events shows remaining divisions possibly concentrated in specific regions though evidence limited, possible later division in peripheral mesocarp regions versus earlier completion near seeds, and spatial heterogeneity reflecting asynchronous division across berry. Temporal spread of final events shows extended period weeks 4-5 from asynchronous division completion, early-activated cells completing first during week 4, late-activated cells continuing into week 5, and extended completion period creating gradual cessation. Developmental heterogeneity shows variation in completion timing across cells creating population asynchrony, heterogeneity enabling smooth transition rather than abrupt cessation, and variation characteristic of developmental populations. Final events completion shows last observable divisions typically week 5, occasional rare divisions possibly extending to week 6 in small subpopulations, and eventual complete cessation establishing truly non-dividing state. Post-final event cell state shows cells exiting cycle after terminal division, cells entering G0 quiescence or G1 expansion state, cells initiating or continuing expansion program through water uptake and vacuole development, and cells committed to enlargement rather than further division. Division marker disappearance shows proliferation markers like PCNA declining during final events, cyclin proteins decreasing to basal levels, mitotic marker phospho-histone H3 becoming undetectable, and marker loss documenting division cessation. Environmental effects on final events include temperature affecting completion timing through developmental rate effects with warm conditions potentially accelerating final event completion, cool conditions extending final division period, and temperature as factor affecting cessation timing. Auxin effects show very low hormone levels during final events correlating with minimal division, auxin decline below proliferative threshold enabling completion, and hormone as proliferation signal loss marking division end. Final events assessment methods include mitotic index measurement showing rare divisions, time-lapse imaging capturing terminal division events, division marker immunohistochemistry detecting last dividing cells, and temporal analysis documenting completion timing. The process represents first aspect of division phase completion emphasizing terminal divisions, preceding complete division cessation and growth mode transition. Duration represents final division period weeks 4-5. Regional timing shows final mitotic events during late July-early August across climates as division concludes. Viticultural significance includes representing last proliferative activity completing berry cellularity, demonstrating progressive division cessation through extended low-frequency divisions, marking transition to complete non-dividing state, and completing cell number establishment. Research applications include analyzing terminal division characteristics and regulation, investigating cell cycle exit mechanisms during final events, evaluating final event timing and spatial patterns, quantifying final proliferative contribution, and assessing final events as transition to division cessation.
Final_Mitotic_Events is first aspect of Division_Phase_Completion, distinguished by terminal division occurrences. Represents last proliferative activity weeks 4-5. Mitotic index declining to <1-2%. Extended cycle times 16-18 hours. Contributes 5-15% final cells. Asynchronous completion creating gradual cessation. Followed by Cell_Division_Cessation as divisions end completely. Last observable divisions typically week 5. Completes berry cellularity establishment.
Schluss-Zellteilungen
final cell divisions
finale mitotische Ereignisse
last mitotic activity
letzte mitotische Aktivität
terminal divisions
terminale Teilungen
ultimate proliferation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 440-442 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Final Mitotic Events
In a Margaret River Cabernet Sauvignon vineyard in Western Australia, final softening state was achieved in late March Year N+1 (BBCH 87) when firmness stabilized at 1.8-2.0 N over 10 days, soluble pectin plateaued at 68-70% of total, sugar content reached 24.2-24.5° Brix, skin anthocyanins stabilized at 2150-2250 mg/kg, seed tannins showed brown coloration indicating phenolic maturity, and sensory evaluation confirmed balanced flavor development under warm dry autumn conditions (daily maximum 26-29°C, minimal precipitation).
Final_Softening_State represents the stable terminal berry texture established at complete softening when all major softening processes have asymptotically approached their endpoints and berry mechanical properties reach cultivar-characteristic equilibrium values that define ripe berry handling characteristics and processing behavior. This state is achieved during late veraison through early ripening (BBCH 85-89) and persists through harvest unless environmental extremes or senescence processes cause further texture degradation. The final softening state is quantitatively defined by stable mechanical property measurements: compression firmness stabilizing within narrow ranges specific to each cultivar (±0.1-0.3 N variation over 1-2 week periods), puncture resistance reaching plateau values reflecting equilibrated skin strength and cell wall integrity, elastic modulus stabilizing at final values indicating completed transition from elastic to plastic deformation behavior characteristic of ripe fruit, and compression deformation curves showing consistent shape indicating reproducible cellular mechanical responses. Cell wall architecture at final softening state exhibits completed structural reorganization: pectin composition stabilized with soluble fraction 70-85% of total (compared to 20-30% green berries), pectin molecular weight distributions plateaued at cultivar-specific values ranging 60-120 kDa with polydispersity indices stabilized, degree of methylesterification equilibrated at 25-40% creating electrostatic repulsion maintaining pectin chain separation, and middle lamella extensively solubilized (60-80% degraded) establishing final intercellular adhesion characteristics that determine berry integrity under compression and cell separation behavior during crushing. Hemicellulose composition at final state shows completed modifications: xyloglucan molecular weight reduced to stable minimum values through transglycosylation reactions, arabinose and xylose decorations removed to final extent affecting xyloglucan-cellulose interactions and cell wall mechanical properties, and cellulose microfibrils maintained largely intact (80-95% of original cellulose preserved) providing residual structural framework supporting remaining firmness. Enzyme activities at final softening state transition to maintenance levels: polygalacturonase activity stabilizes at 30-60% of peak values (40-80 units) sufficient to maintain pectin solubilization equilibrium without driving further extensive degradation, pectin methylesterase operates at 30-50% peak levels (35-70 units) balancing methylation-demethylation dynamics, and other cell wall enzymes including xyloglucan endotransglycosylase, expansins, and glycosidases maintain low-moderate activities (20-40% of peak) preserving cell wall plasticity without continued breakdown. Cellular turgor at final softening state reaches equilibrium reflecting water relations and osmotic adjustments: turgor pressure stabilizes at 0.3-0.6 MPa (compared to 0.6-1.0 MPa green berries) contributing 15-25% to residual berry firmness, osmotic potential reaches final values -1.8 to -2.6 MPa driven by accumulated sugars (20-28° Brix) and organic acids creating characteristic ripe berry water relations, and cell membrane permeability stabilizes at elevated levels compared to green berries facilitating metabolite exchange but maintained below catastrophic leakage thresholds. Final softening state mechanical properties determine berry and wine quality characteristics: skin tensile strength at 40-65% of green berry values affects berry splitting resistance, bunch compression, and ease of crushing during processing, berry compression firmness correlates with must yield during pressing (softer berries r² = 0.75-0.88 with higher free-run fraction), and cell wall porosity influences extraction efficiency of phenolic compounds, flavors, and aromatics during maceration with more degraded cell walls facilitating enhanced extraction. Cultivar-specific final softening states reflect genetic variation in cell wall composition and degradation extent: very soft cultivars (Pinot Noir, Gamay) achieve 0.8-1.5 N firmness with 75-85% pectin solubilization and extensive middle lamella degradation creating delicate berries requiring gentle handling, moderately soft varieties (Merlot, Syrah, Chardonnay) reach 1.2-2.0 N with 65-75% solubilization and intermediate cell wall breakdown producing balanced texture for varied applications, while firmer cultivars (Cabernet Sauvignon, Nebbiolo, Sangiovese) maintain 1.5-2.5 N through 55-70% solubilization and more limited structural degradation allowing mechanical harvest tolerance. Environmental factors influencing final softening state characteristics inclu water availability during ripening with moderate deficit potentially maintaining slightly firmer final states through 10-20% higher turgor and reduced enzyme activity while extreme deficit risks desiccation and abnormal texture, temperature during late ripening affecting enzyme activity levels with cooler conditions (18-22°C) potentially preserving slightly firmer texture compared to warmer ripening (25-30°C) though final differences typically 30% berry rupture), and may indicate over-ripening with potential flavor degradation, insufficient softening (>2.5 N at nominal sugar maturity) suggests incomplete ripening potentially producing astringent wines with harsh tannins and limited aromatic expression, while optimal final states (1.2-2.2 N for most premium wine cultivars) balanced with chemical and phenolic maturity indicate ideal harvest readiness producing wines with optimal extraction, balanced structure, and characteristic varietal expression. Viticultural management targeting optimal final softening state involves: irrigation scheduling during late ripening to avoid water stress that might maintain excessive firmness or excess that accelerates over-softening, canopy management to moderate bunch temperatures preventing accelerated softening while allowing adequate warmth for complete ripening, and disease control particularly Botrytis prevention since soft berries with compromised skin integrity show maximum vulnerability requiring fungicide protection or harvest timing minimizing disease window exposure.
Final_Softening_State is distinguished from Complete_Berry_Softening by representing the stable equilibrium condition rather than the achievement process, from Peak_Softening by exhibiting stability rather than maximum transformation velocity, and from Progressive_Softening by showing plateau rather than continued decline. Stability typically maintained 2-4 weeks under optimal conditions. Firmness values: very soft cultivars 0.8-1.5 N, moderate 1.2-2.0 N, firm varieties 1.5-2.5 N. Pectin solubilization 70-85%, turgor contribution to firmness 15-25%, cell wall structural integrity 75-85%. Final_Softening_State achievement indicates berry readiness for integrated maturity assessment guiding harvest timing decisions.
Gleichgewichts-Beeren-Textur
equilibrium berry texture
erntereife Textur
harvest-ready texture
stabile reife Festigkeit
stable ripe firmness
terminal mechanical state
terminaler mechanischer Zustand
Keller M (2020) The Science of Grapevines 3rd Ed pages 232-238 berry maturation and harvest assessment Rolle L, Giacosa S, Gerbi V, Novello V (2011) Comparative study of texture properties, color characteristics, and chemical composition of ten white table-grape varieties Amer J Enol Vitic 62:49-56 final berry texture characteristics Letaief H, Rolle L, Zeppa G, Gerbi V (2008) Assessment of grape skin hardness by a puncture test J Sci Food Agric 88:1567-1575 final state mechanical properties.
Final Softening State
In a Bordeaux Cabernet Sauvignon vineyard, final sugar accumulation was tracked over 32 days from mid-September through mid-October Year N+1 (BBCH 86-89) when berry Brix increased from 22.8° to 25.6° (2.8° gain, average 0.88% per week declining from 1.2% early to 0.6% late final phase), hexose import estimated declining from 58 to 28 μmol g⁻¹ h⁻¹, berry fresh weight stable 1.92-1.98 g (minimal expansion), phenolic maturity achieved week 3 enabling harvest at optimal balance despite minimal continued sugar accumulation final week, under typical Bordeaux autumn conditions (daily high declining 24° to 18°C, canopy beginning senescence).
Final_Sugar_Accumulation represents the decelerating phase of hexose import and storage during late ripening when berries approach cultivar-specific maximum sugar capacity causing progressive reduction in accumulation rates creating the asymptotic approach toward sugar equilibrium that defines final harvest sweetness and signals physiological maturity. This final accumulation occurs during late ripening approaching harvest (BBCH 86-89), typically days 35-65 post-veraison onset spanning 25-35 days, and is characterized by berry Brix increasing from 21-26° at phase onset to final harvest levels 23-30° representing 2-6° gain during final phase substantially less than peak accumulation 5-9° over similar duration, daily Brix increase declining progressively from 0.18-0.32% per day early final phase to 0.05-0.15% per day late phase representing 2-4 fold deceleration compared to peak rates 0.50-0.85% per day, hexose import rates declining from 50-90 μmol combined glucose and fructose g⁻¹ h⁻¹ to 20-50 μmol g⁻¹ h⁻¹ as vacuolar concentrations approach saturation, and cumulative sugar gain 80-140 g/L over final phase accounting for 15-35% of total ripening accumulation with declining marginal contribution as limits approached. Physiological basis for deceleration includes vacuolar hexose concentrations approaching maximum sustainable levels 1600-2400 mM combined glucose and fructose (cultivar-dependent) creating reduced concentration gradients diminishing thermodynamic driving force for continued import, osmotic potential reaching extreme values -2.8 to -3.8 MPa approaching cell tolerance limits potentially constraining further accumulation, cytoplasmic hexose potentially accumulating slightly 70-120 mM as tonoplast transport capacity saturates reducing plasma membrane uptake gradient, and phloem delivery potentially declining 20-40% as source photosynthesis decreases with advancing season and canopy senescence beginning reducing substrate availability. Molecular mechanisms during final phase show hexose transporter gene expression declining gradually from peak levels 100-150 fold to final 60-100 fold above green berry as ripening program begins downregulation, protein turnover potentially exceeding synthesis causing transporter density to decline 15-30% from peak reducing import capacity, post-translational modifications possibly reducing transporter activity though mechanisms in grape incompletely characterized, and potential feedback inhibition where high vacuolar hexose signals through unknown mechanisms to reduce transport gene expression or activity preventing excessive accumulation. Metabolic changes during final phase include increased proportion of imported hexoses respired rising from 20-30% during active phase to 30-45% during final phase as vacuolar storage becomes less favored, respiratory substrate shifting toward greater hexose contribution 45-65% of CO₂ production as organic acids deplete reducing alternative substrate availability, and energetic efficiency declining as higher proportion of import oxidized rather than stored representing less efficient carbon use but necessary for maintaining cellular ATP demands as acid catabolism wanes. Temporal pattern within final phase demonstrates early final (days 35-45, BBCH 86) showing moderate deceleration with Brix increase 0.15-0.28% per day and hexose import 40-75 μmol g⁻¹ h⁻¹ representing transition from peak, mid-final (days 45-55, BBCH 87-88) exhibiting progressive slowing 0.08-0.18% Brix per day and 25-50 μmol g⁻¹ h⁻¹ import as limits approached, and late final approaching harvest (days 55-65, BBCH 89) showing minimal accumulation 0.03-0.10% per day and 15-35 μmol g⁻¹ h⁻¹ import asymptotically approaching plateau. Environmental influences on final accumulation include temperature effects persisting with warm conditions 26-32°C maintaining higher rates while cooler temperatures 18-24°C further reducing already declining rates potentially extending final phase duration, declining photoperiod and solar angle reducing source photosynthesis 20-40% compared to peak ripening limiting substrate availability accelerating rate decline, autumn cooling in late-ripening varieties potentially limiting accumulation through temperature effects on metabolism, water stress effects with late-season deficit potentially hastening approach to plateau through reduced berry expansion and phloem delivery, and canopy senescence particularly nitrogen-deficient vines showing earlier photosynthesis decline limiting late accumulation. Cultivar variation in final accumulation demonstrates high-Brix varieties maintaining higher rates longer with final phase rates 0.12-0.25% Brix per day achieving maximum 26-30° Brix, moderate cultivars showing 0.08-0.18% per day reaching 22-26° typical commercial levels, and lower-Brix types displaying 0.05-0.12% per day approaching 20-24° final levels reflecting genetic differences in maximum capacity and regulatory set-points determining when deceleration occurs and plateau achieved. Coordination with other late ripening processes shows acid decline continuing during final sugar accumulation maintaining 0.10-0.30 g/L TA per day loss creating developing sugar-acid balance, berry softening completed with firmness stabilized at ripe levels 0.4-1.2 N, color development in reds reaching maximum anthocyanin 600-1800 mg/kg with no further increase, flavor-aroma synthesis continuing with volatiles accumulating to final 150-750 μg/kg total creating characteristic varietal profiles, and seed maturation progressing with browning and hardening indicating physiological maturity independent of sugar levels. Quality implications of final accumulation include rate and duration determining final achievable Brix within growing season with early plateau potentially limiting maximum sugar in cool vintages, extended final phase enabling very high sugars >26-28° Brix beneficial for full-bodied or late-harvest styles but risking over-ripeness, coordination between final sugar accumulation and phenolic maturity particularly reds critical as tannin development may continue after sugar plateau requiring harvest timing decisions balancing parameters, and vintage variation substantial with warm years showing briefer final phase reaching plateau earlier while cool years exhibiting extended gradual accumulation potentially beneficial for flavor development. Harvest timing considerations during final phase recognize declining accumulation rates signal approaching maturity requiring increased monitoring frequency, target sugar achievement timing becomes more uncertain as rates slow necessitating flexibility in harvest scheduling, integration with acid levels particularly important as continued acid decline during final phase can produce unbalanced high-sugar low-acid fruit if harvest delayed excessively, phenolic and flavor assessment essential particularly reds where extended hang time during final phase may improve quality despite minimal sugar gain, and weather monitoring critical as late-season risks (rain, frost, disease) increase requiring harvest decisions balancing quality optimization against crop protection. Viticultural management during final phase includes maintaining canopy health supporting continued photosynthesis though senescence natural and inevitable, disease control particularly critical as high-sugar berries vulnerable to Botrytis and sour rot with infection risk increasing with autumn moisture, irrigation typically reduced or ceased allowing slight deficit potentially enhancing final concentration and triggering ripening completion, bird protection essential as peak sugar attractiveness to wildlife, and harvest logistics advancing as approaching plateau indicates imminent optimal harvest window requiring readiness. Physiological limits constraining final accumulation include maximum vacuolar solute concentration ~2200-2600 mM approaching saturation beyond which further accumulation thermodynamically unfavorable or physically impossible, cell wall and membrane mechanical limits at extreme osmotic potentials <-3.5 to -4.0 MPa potentially compromising integrity, metabolic feedback regulation where high sugar signals ripening completion downregulating transport machinery, and declining source capacity as photosynthesis wanes with advancing season fundamentally limiting substrate availability regardless of sink capacity. Research applications include transporter downregulation studies revealing molecular control of accumulation cessation, osmotic potential measurements characterizing thermodynamic limits, flux analysis quantifying changing import-respiration partitioning, and modeling approaches predicting plateau timing enabling harvest optimization based on accumulation trajectory analysis.
Final_Sugar_Accumulation is distinguished from Peak_Sugar_Accumulation by showing declining rates (0.05-0.32% Brix per day progressive deceleration) rather than maximum (0.50-0.85%), from Active_Sugar_Accumulation by representing asymptotic approach (2-6° gain) rather than sustained accumulation (7-13° gain), and from Sugar_Plateau by showing continued though slowing accumulation rather than equilibrium. Duration 25-35 days BBCH 86-89. Accounts for 15-35% total ripening gain. Signals approaching maturity. Final_Sugar_Accumulation represents deceleration phase completing sugar accumulation trajectory.
Spätphasen-Zucker-Import
approaching sugar maximum
decelerating accumulation
dezelerierende Akkumulation
late-stage sugar import
sich näherndes Zucker-Maximum
Keller M (2020) The Science of Grapevines 3rd Ed pages 226-228 final accumulation Coombe BG (1992) Research on development and ripening of the grape berry Amer J Enol Vitic 43:101-110 deceleration phase Matthews MA, Anderson MM (1988) Fruit ripening in Vitis vinifera L.:responses to seasonal water deficits Amer J Enol Vitic 39:313-320 late ripening dynamics.
Final Sugar Accumulation
Pinot Noir showing fine feeder roots in May, soil cores 15-30 cm revealing roots 0.4-0.7 mm diameter fourth and fifth-order terminal branches, fine feeder roots showing white coloration indicating youth, primary anatomical structure with 2-3 xylem vessels, abundant root hairs 380 per cm, arbuscular mycorrhizal colonization 64%, median lifespan 3.1 months, fine feeder roots comprising 3.2% total root biomass but providing estimated 52% of vine nitrogen and phosphorus uptake, fine feeder roots demonstrating extreme absorption specialization
Fine feeder root is feeder root at finest end of diameter spectrum typically <1 mm often 0.3-0.8 mm representing highest-order terminal branches in root system hierarchy, characterized by extremely small diameter approaching root hair dimensions, minimal vascular development with only 1-4 xylem vessels, very short lifespan typically 1-6 months creating highest turnover rates, maximal surface-to-volume ratio optimizing absorption per biomass unit, and serving specialized absorption function providing disproportionate contribution to uptake despite minuscule individual biomass, with fine feeder roots in grapevine showing typical characteristics including fourth-order or higher branching representing terminal tips in hierarchical architecture, primary anatomical structure completely lacking secondary growth maintaining ultimate thinness, abundant root hair production potentially 300-500+ hairs per centimeter creating extensive absorption surface, dense arbuscular mycorrhizal colonization often 50-80% of length enhancing function, extremely ephemeral nature with median lifespan 2-4 months shortest among root classes, and demonstrating functional extreme where fine feeder roots represent maximum absorption specialization trading longevity for efficiency creating roots optimized purely for resource capture with minimal investment in structure or persistence. Fine feeder root diameter characteristics includes= showing very small diameter (fine feeder roots typically 0.3-0.8 mm substantially smaller than general feeder root 0.5-2 mm range), exhibiting hair-like thinness (finest specimens approaching root hair dimensions 0.2-0.4 mm creating extremely thin structures), demonstrating uniform caliber (fine feeder roots maintaining consistent diameter along short length), revealing fragility (extreme thinness creating delicate structures easily damaged by handling or soil disturbance), and displaying measurement challenges (fine diameter requiring careful measurement techniques avoiding compression or breakage). Fine feeder root anatomical minimalism includes= showing minimal vascular tissue (fine feeder root stele containing only 1-4 xylem vessels versus 5-15+ in coarser roots), exhibiting thin cortex (cortex only 2-4 cell layers thick minimizing radial transport distance), demonstrating simple endodermis (single-layer endodermis with casparian strips though sometimes irregular or incomplete), revealing small cell size (cells in fine feeder roots smaller than in coarser roots creating fine tissue texture), and displaying primary structure exclusively (complete absence of any secondary growth maintaining pure primary anatomical organization). Fine feeder root extremely short lifespan includes= showing median longevity (fine feeder roots living median 2-4 months substantially shorter than coarser feeder roots 4-8 months), exhibiting highest turnover (fine feeder root population potentially replacing 2-4+ times per year highest turnover rate), demonstrating ephemeral extreme (fine feeder roots representing most ephemeral root class in grapevine system), revealing rapid mortality (individual fine feeder roots quickly senescing after brief functional period), and displaying production intensity (maintaining fine feeder root population requiring continuous intensive production). Fine feeder root functional specialization includes= providing maximum specific uptake (uptake per unit fine feeder root surface area or biomass highest among all root types), exhibiting extreme efficiency (fine feeder roots showing best ratio of absorption to construction and maintenance costs), demonstrating exclusive absorption (fine feeder roots performing purely absorptive function with no structural, transport, or storage roles), revealing vulnerability (fine feeder roots highly vulnerable to any stress rapidly losing function or dying), and displaying expendable design (fine feeder roots designed as expendable absorption organs readily sacrificed under stress). Fine feeder root distribution patterns includes= concentrating in resource-rich zones (fine feeder roots proliferating intensively where nutrients and water abundant), showing extreme patchiness (fine feeder root distribution highly heterogeneous varying 100-1000 fold spatially), exhibiting seasonal variation (fine feeder root abundance varying dramatically seasonally with peaks spring and fall), demonstrating shallow preference (fine feeder roots predominantly in upper 20-40 cm soil), and revealing responsive proliferation (fine feeder roots rapidly increasing in favorable patches declining in poor zones). Fine feeder root absorption contribution includes= providing disproportionate uptake (fine feeder roots potentially contributing 40-60% of total vine uptake despite comprising only 2-5% biomass), exhibiting nutrient specialization (fine feeder roots particularly important for immobile nutrients like phosphorus requiring close soil contact), demonstrating water acquisition (fine feeder roots contributing substantially to water uptake particularly from small pores), revealing mycorrhizal synergy (fine feeder roots and arbuscular mycorrhizae working synergistically creating enhanced absorption system), and displaying critical importance (fine feeder root abundance and activity critically affecting vine resource status). Fine feeder root versus coarser feeder roots includes= contrasting in diameter (fine <1 mm versus coarse 1-2 mm), showing different longevity (fine 2-4 months versus coarse 4-8 months median), exhibiting turnover differences (fine showing 2-4+ versus coarse 1-2 replacements per year), demonstrating anatomical distinctions (fine with 1-4 xylem vessels versus coarse 5-15+), and revealing functional extremes (fine representing ultimate absorption specialization versus coarse showing some structural development). Fine feeder root research and measurement includes= requiring careful methods (fine feeder roots easily damaged during sampling requiring gentle techniques), showing imaging applications (minirhizotrons or root scanners documenting fine feeder root dynamics), exhibiting isotope utility (tracers demonstrating fine feeder root uptake contribution), demonstrating molecular approaches (gene expression analysis revealing fine feeder root specialization), and revealing modeling challenges (fine feeder root extreme turnover and patchiness creating modeling difficulties).
Fine_Feeder_Root emphasizes finest diameter end of feeder root spectrum and is distinguished from coarser feeder roots by focus on ultimate thinness and ephemeral nature. Includes Hair_Like_Fine_Feeder_Root, Delicate_Fine_Feeder_Root, and White_Fine_Feeder_Root. Critical for understanding extreme thinness, minimal anatomy, shortest lifespan, highest turnover, maximum efficiency, and the finest feeder roots representing ultimate absorption specialization with disproportionate uptake contribution.
Sehr feine Feinwurzel
Terminal fine root
Terminale Feinwurzel
Ultrafeine Saugwurzel
Ultrafine absorptive root
Very fine root
Fine Feeder Root
Pinot Noir grafted on Riparia Gloire rootstock at 11 years showing fine lateral tertiary roots sampled at 38 cm depth, fifth-order fine lateral tertiary roots 0.6 mm diameter 2.4 cm length branching from fourth-order parent, sixth-order fine laterals 0.3 mm diameter 1.2 cm length terminal in hierarchy, fine lateral tertiary roots bearing abundant root hairs estimated 180 hairs per cm root length increasing surface area 12-fold, primary anatomical structure lacking any secondary growth, mycorrhizal colonization 68% of fine root length, fine lateral tertiary roots providing estimated 82% of total vine water uptake and 76% of nitrogen uptake, minirhizotron observations indicating fine root lifespan 4-9 months average
Fine lateral tertiary root is lateral tertiary root showing very small diameter typically <2 mm and ephemeral nature, emphasizing fine size and high surface area-to-volume ratio optimizing absorption, characterized by fourth-order or higher branching position representing finest lateral branches in root hierarchy, primary anatomical structure lacking secondary growth maintaining delicate organization, abundant root hair production creating extensive absorption surface, short lifespan typically 2-12 months reflecting high turnover strategy, and serving almost exclusively absorptive function providing majority of vine water and nutrient uptake from soil micro-sites, with fine lateral tertiary roots showing diameter typically 0.2-2 mm length 1-5 cm creating vast numbers and enormous collective surface area, producing minimal or no further branching being terminal or near-terminal in hierarchy, responding rapidly to soil resource heterogeneity proliferating in favorable patches and senescing in poor zones, and representing ephemeral dynamic root component with continuous production and mortality contrasting with persistent woody root framework demonstrating high-turnover resource acquisition strategy. Fine lateral tertiary root diameter and morphology includes= showing very small diameter (<2 mm by definition distinguishing from larger tertiary roots), exhibiting typical range 0.2-2 mm (finest fine lateral tertiary roots 0.2-0.5 mm while larger 1-2 mm reflecting fourth through sixth or higher orders), demonstrating high surface area-to-volume ratio (small diameter maximizing surface relative to volume optimizing absorption efficiency), displaying delicate structure (fine lateral tertiary roots fragile and easily damaged during excavation or handling), and revealing white to tan coloration (young fine roots typically light-colored darkening with age or mycorrhizal colonization). Fine lateral tertiary root anatomy includes= showing primary structure exclusively (fine lateral tertiary roots lacking any secondary growth maintaining primary anatomical organization throughout life), exhibiting thin epidermis (outer cell layer with or without root hairs depending on age and position), demonstrating cortex dominance (cortical parenchyma comprising large proportion 50-70% of root cross-section), containing endodermis (innermost cortical layer with casparian strip regulating ion uptake), showing small vascular cylinder (limited xylem typically 2-8 vessels and small phloem reflecting fine root size), and lacking protective bark (fine roots maintaining epidermis or forming thin exodermis rather than developing periderm). Fine lateral tertiary root hair production includes= producing abundant root hairs (fine lateral tertiary roots bearing numerous epidermal extensions particularly on youngest portions), showing hair density (potentially 50-300+ root hairs per centimeter root length depending on conditions), exhibiting hair dimensions (individual root hairs typically 0.1-1 mm length 10-20 micrometers diameter), demonstrating surface area multiplication (root hairs increasing effective absorption surface 5-20 fold compared to root alone), and revealing responsive production (root hair abundance varying with soil moisture and nutrient availability). Fine lateral tertiary root absorptive specialization includes= optimizing water uptake (small diameter and abundant root hairs maximizing water absorption per unit root biomass), demonstrating nutrient absorption efficiency (fine roots with high surface area-to-volume ratio efficiently acquiring dilute soil nutrients), showing direct uptake capacity (epidermis and root hairs directly absorbing water and minerals without intervening tissues), exhibiting high metabolic activity (fine roots showing elevated respiration supporting active absorption), and revealing selective permeability (endodermis casparian strip enabling selective ion uptake and regulation). Fine lateral tertiary root mycorrhizal colonization includes= hosting AM fungi extensively (fine lateral tertiary roots showing high arbuscular mycorrhizal colonization rates typically 40-80% root length), demonstrating arbuscule formation (fungal structures in cortical cells facilitating nutrient exchange), showing vesicle development (fungal storage structures in root cortex containing lipids), exhibiting hyphal networks (external mycorrhizal hyphae extending from fine roots exploring soil beyond root surface), and revealing phosphorus uptake enhancement (mycorrhizal association particularly important for phosphorus acquisition from low-availability soils). Fine lateral tertiary root lifespan and turnover includes= showing short lifespan (fine lateral tertiary roots typically living 2-12 months representing shortest-lived root component), exhibiting high turnover rate (fine roots potentially replacing themselves 1-3+ times per year), demonstrating seasonal mortality peaks (fine root death concentrated late summer-fall and winter though occurring year-round), revealing stress sensitivity (fine roots dying readily under drought, temperature extremes, or pathogen attack), and contributing substantially to carbon cycling (fine root turnover providing 30-50% of total root system carbon input to soil annually). Fine lateral tertiary root distribution patterns includes= concentrating in upper soil (fine lateral tertiary roots most abundant 20-60 cm depth in nutrient and moisture-rich topsoil), showing extremely patchy distribution (fine root density varying 10-100 fold over short distances reflecting resource heterogeneity), proliferating in favorable zones (fine roots rapidly increasing in resource-rich patches around organic matter, nutrient accumulations, or moist zones), demonstrating avoidance of poor zones (fine roots absent or sparse in dry, nutrient-poor, compacted, or toxic soil areas), and revealing competition zones (fine root density potentially reduced in regions with high root density from neighboring plants). Fine lateral tertiary root functional importance includes= providing majority of absorption (fine lateral tertiary roots despite small individual size providing 70-90% of total vine water and nutrient uptake), demonstrating cost-effectiveness (fine root construction and maintenance costs low relative to absorption benefits), showing responsive flexibility (fine root architecture rapidly adjusting to changing soil conditions), enabling resource foraging (fine roots exploring soil at millimeter scale detecting and exploiting micro-sites), and representing critical vine function (vine survival and productivity ultimately depending on fine root absorption despite dominance of woody roots in biomass).
Fine_Lateral_Tertiary_Root emphasizes very small diameter <2 mm and ephemeral absorptive function and is distinguished from larger tertiary roots by focus on finest size and highest turnover. Critical for understanding absorption dominance, root hair function, mycorrhizal dependence, high turnover strategy, resource foraging, and the finest lateral roots providing majority of vine resource uptake despite small individual size and short lifespan.
Ephemeral lateral tertiary
Ephemere tertiäre Seitenwurzel
Haartragende Feinwurzel
Hair-bearing fine root
Sehr feine Lateralwurzel
Very fine lateral root
Fine Lateral Tertiary Root
Cabernet Sauvignon grafted on 1103 Paulsen rootstock at 17 years showing fine tertiary root system, sequential soil coring at 45 cm depth revealing fine roots fourth through sixth branching orders, fifth-order fine tertiary roots 0.8 mm diameter 2.8 cm length with 220 root hairs per cm length, sixth-order terminal fine roots 0.4 mm diameter 1.4 cm length, fine tertiary root length density 32 cm per cm³ soil providing estimated 0.18 square meters absorption surface per liter soil, mycorrhizal colonization 64% of fine root length with arbuscules and external hyphae, minirhizotron observations indicating fine root median lifespan 7.2 months, fine tertiary roots providing estimated 84% of vine water uptake and 78% of nutrient uptake
Fine tertiary root is tertiary root emphasizing fine diameter typically <2 mm and specialized absorptive function, characterized by fourth-order or higher branching position representing finest roots in hierarchy, primary anatomical structure lacking secondary growth throughout lifespan, abundant root hair production maximizing absorption surface area, short ephemeral lifespan typically 2-12 months showing high turnover, and serving almost exclusively for water and nutrient absorption providing 70-90% of total vine resource uptake despite comprising only 5-15% of root system biomass, with fine tertiary roots showing typical dimensions 0.2-2 mm diameter and 1-5 cm length creating enormous numbers estimated thousands to millions per vine and vast collective surface area potentially hundreds of square meters, responding rapidly to soil resource heterogeneity through proliferation and senescence, hosting dense arbuscular mycorrhizal colonization enhancing uptake capacity, and representing the absorptive organ analogous to leaves as photosynthetic organs demonstrating that fine roots are functional endpoint of hierarchical root architecture where actual resource acquisition occurs. Fine tertiary root size and abundance includes= showing very small diameter (<2 mm by definition distinguishing from larger coarse roots), exhibiting typical range 0.2-2 mm (finest fine roots 0.2-0.5 mm common while 1-2 mm representing lower size threshold), demonstrating short length (individual fine roots typically 1-5 cm rarely exceeding 10 cm), revealing enormous numbers (mature vine potentially producing tens of thousands to millions of fine roots), and creating vast surface area (fine root collective surface area potentially 50-200+ square meters per vine despite small individual size). Fine tertiary root anatomical specialization includes= showing primary structure exclusively (fine roots maintaining primary anatomical organization lacking any secondary growth or wood formation), exhibiting thin epidermis (single-cell outer layer with or without root hairs and often forming thin exodermis), demonstrating cortex dominance (cortical parenchyma tissue comprising 50-70% of fine root cross-sectional area providing storage and transport), containing well-developed endodermis (casparian strip and potentially suberin lamellae regulating radial solute movement), showing small vascular cylinder (limited xylem typically 2-12 vessels and small phloem adequate for fine root size), and lacking protective periderm (fine roots never developing bark maintaining delicate primary structure). Fine tertiary root hair morphology includes= producing prolific root hairs (fine roots bearing numerous epidermal extensions particularly on youngest root portions 1-5 cm from tip), showing high hair density (typically 50-300+ root hairs per centimeter root length depending on species and conditions), exhibiting hair dimensions (individual root hairs 0.1-1 mm length 10-20 micrometers diameter creating extensive surface), demonstrating rapid turnover (individual root hairs living only days to weeks with continuous production and death), and revealing surface multiplication (root hairs increasing effective absorption surface area 5-20 fold compared to root cylinder alone). Fine tertiary root absorptive dominance includes= providing majority of water uptake (fine roots contributing 70-90% of total vine water absorption despite small biomass proportion), demonstrating nutrient acquisition dominance (fine roots providing 70-90% of nitrogen, phosphorus, potassium, and micronutrient uptake), showing high uptake efficiency (fine roots absorbing 10-100+ times more water and nutrients per gram biomass than coarse roots), exhibiting active absorption capacity (fine root epidermis and root hairs with abundant transporters enabling active ion uptake), and revealing critical functional importance (vine survival and productivity ultimately dependent on fine root absorptive function). Fine tertiary root mycorrhizal symbiosis includes= hosting arbuscular mycorrhizae extensively (fine roots showing AM fungal colonization typically 40-80% of root length), forming arbuscules (tree-like fungal structures in cortical cells serving as nutrient exchange sites), developing vesicles (spherical fungal storage structures in root cortex containing lipid reserves), supporting external hyphae (mycorrhizal hyphae extending 1-10+ cm from fine root surface greatly expanding exploration volume), and demonstrating mutual dependency (grapevine strongly mycorrhiza-dependent showing reduced growth without fungal symbionts while fungi depending on plant carbohydrates). Fine tertiary root lifespan and dynamics includes= showing very short lifespan (fine roots typically living 2-12 months representing most ephemeral root component), exhibiting continuous turnover (ongoing fine root production and mortality throughout year with peaks in spring and fall), demonstrating high replacement rate (fine root population potentially replacing itself 1-3+ times annually), revealing stress sensitivity (fine roots dying readily under drought, temperature extremes, flooding, or pathogen attack), and contributing substantially to carbon cycling (fine root turnover providing 30-60% of total root system annual carbon input to soil). Fine tertiary root distribution and responsiveness includes= showing extremely heterogeneous distribution (fine root density varying 10-1000 fold spatially depending on resource availability), concentrating in upper soil horizons (majority of fine roots 20-60 cm depth in nutrient-rich topsoil), proliferating rapidly in resource patches (fine root density increasing 5-50 fold in localized zones of high nutrients or moisture), demonstrating avoidance of poor conditions (fine roots absent from dry, compacted, toxic, or severely nutrient-poor zones), and revealing dynamic architecture (fine root spatial distribution continuously changing through production and mortality responding to resource heterogeneity). Fine tertiary root functional ecology includes= representing resource acquisition organs (fine roots functionally analogous to leaves but for below-ground resources), demonstrating construction-maintenance trade-offs (short fine root lifespan potentially optimal balancing construction costs against benefits in variable environments), showing foraging behavior (fine root proliferation and senescence enabling resource foraging at fine spatial scales), exhibiting competitive interactions (fine roots competing with neighboring plants for limiting resources), and revealing evolutionary adaptation (fine root traits including diameter, lifespan, and branching reflecting adaptation to soil environments and resource strategies).
Fine_Tertiary_Root emphasizes fine diameter <2 mm and absorptive specialization and is distinguished from larger coarse roots by focus on absorption dominance. Includes Hair_Like_Fine_Tertiary_Root and Absorptive_Fine_Tertiary_Root. Critical for understanding absorption organs, resource acquisition, root hair function, mycorrhizal dependence, high turnover, dynamic responsiveness, and the finest ephemeral roots providing majority of vine water and nutrient uptake representing functional endpoint of root architecture.
Absorptive Wurzel
Absorptive root
Ephemeral fine root
Ephemere Feinwurzel
Feine Tertiärwurzel
Terminal absorptive root
Eissenstat, D. M., et al. (2000). Building roots in a changing environment= Implications for root longevity. New Phytologist, 147(1), 33-42. | Pregitzer, K. S., et al. (2002). Fine root architecture of nine North American trees. Ecological Monographs, 72(2), 293-309.
Fine Tertiary Root
A fine veinlet network in a sun leaf at node 6 of Vitis vinifera cv. Cabernet Sauvignon in Bordeaux, France, developing under high light exposure (average 85% full sun during expansion), individual veinlet segments measuring 0.18-0.35 mm length with diameter of approximately 0.07 mm visible only under 15-20× magnification, containing single small tracheid (8-10 μm diameter) without clearly distinguishable phloem, creating very dense reticulate mesh with areoles averaging 0.12 mm² area containing 10-12 mesophyll cells, veinlets extending to within 45-55 μm of all intercostal regions representing maximum 2 cell layers diffusion distance, total fine veinlet density of 13.8 mm per mm² representing approximately 80% of total venation length and substantially exceeding shade leaf veinlet density (8.5 mm per mm²) from the same cultivar, creating approximately 1150 areoles per mm² blade area, enabling light-saturated photosynthesis rate of 14.2 μmol CO₂ m⁻² s⁻¹ and maximum transpiration rate of 4.5 mmol H₂O m⁻² s⁻¹ without hydraulic limitation, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing the fine-scale high-density terminal vascular network characteristic of sun leaf adaptation.
A veinlet exhibiting particularly small diameter and high density creating very fine reticulate mesh in Vitis vinifera, characterized by diameter at the lower end of the veinlet range typically 0.05-0.10 mm requiring magnification for clear visibility, minimal vascular anatomy often consisting of single tracheid without distinct phloem elements, very high density with individual veinlet segments spaced 0.15-0.30 mm apart creating very fine mesh, small areole sizes typically 0.08-0.20 mm² containing 8-15 mesophyll cells per areole, very short mesophyll-to-vein diffusion distances typically 40-60 μm representing maximum 2-3 cell layers, total fine veinlet length often exceeding 12-15 mm per mm² blade area, occurrence primarily in sun leaves where high photosynthetic capacity requires enhanced hydraulic conductivity, and functional implications including enabling maximum transpiration and photosynthesis rates through minimized hydraulic resistance and diffusion distances, but requiring substantial resource investment in vascular tissue potentially reducing photosynthetic tissue proportion. Fine veinlets represent the extreme elaboration of terminal vascular networks, developing under high light conditions that provide resources and create functional demands for enhanced vascular density, creating the anatomical basis for high photosynthetic capacity characteristic of sun-adapted leaves through superior hydraulic efficiency.
Fine veinlets represent the high-density extreme of terminal venation, distinguished from typical veinlets by smaller diameter, denser spacing, and smaller areoles. Development occurs primarily in sun leaves where high light during expansion provides resources and creates functional demands for enhanced vascular capacity, with shade leaves typically showing lower veinlet densities creating larger areoles and longer diffusion distances. The functional advantage involves reduced hydraulic resistance through shorter pathways and reduced diffusion limitations through shorter mesophyll-to-vein distances, enabling higher maximum photosynthetic and transpiration rates before hydraulic constraints limit function. The resource cost includes substantial investment in vascular tissues, with fine veinlet networks potentially occupying 8-12% of blade cross-sectional area compared to 5-7% in lower-density shade leaves, reducing the proportion available for photosynthetic mesophyll. Anatomical simplification to single tracheids represents minimum functional capacity, with some fine veinlets possibly functioning purely for xylem delivery without photosynthate collection capacity. Cultivar variation exists in fine veinlet development capacity, with some varieties showing enhanced veinlet elaboration under high light while others show more limited plasticity. Fine veinlet density correlates positively with maximum photosynthetic capacity across sun-shade spectrums and among cultivars, supporting the functional importance of terminal vascular architecture in determining leaf performance.
dense terminal vein
dichte Terminalvene
sehr feines Äderchen
ultimate veinlet
ultrafeine Ader
very fine vein
Based on Sack & Holbrook (2006) "Leaf hydraulics" Annual Review of Plant Biology 57=361-381 Brodribb et al. (2007) "Leaf hydraulic capacity in ferns, conifers and angiosperms= impacts on photosynthetic maxima" New Phytologist 165=839-846 Schultz (1996) "Leaf absorptance of visible radiation in Vitis vinifera L.= estimates of age and shade effects with a simple field method" Scientia Horticulturae 66=93-102 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 219-242.
Fine Veinlet
Finish description for Grand Cru Burgundy (Chambertin, 2015 vintage) records an exceptionally long finish of 30+ seconds with progressive evolution from concentrated red fruit through mineral-saline complexity to a final ethereal dried-rose and iron-mineral persistence, the aftertaste maintaining aromatic complexity and evolving continuously rather than simply fading — the hallmark of great Burgundy where the finish reveals new dimensions not perceived during the mid-palate assessment. Finish description for commercial Pinot Grigio from Veneto (2024 vintage) documents a short finish of 4-5 seconds with simple almond-pear fade to a clean but unremarkable conclusion, consistent with the wine's positioning as a fresh, uncomplicated everyday drinking wine. Finish description for Blaufränkisch Reserve from Eisenberg DAC (2021 vintage) assesses at "long" (18-22 seconds) with persistent dark cherry, mineral-slate, and fine peppery-tannin persistence, the iron-mineral aftertaste reflecting the distinctive Eisenberg terroir signature derived from the region's iron-rich soils.
Finish description is a taste description entity that records the persistence, quality, and evolution of gustatory, olfactory, and tactile sensations remaining on the palate after the wine has been swallowed or expectorated, assessed through multiple dimensions including length (duration in seconds from swallowing to the extinction of detectable flavor, classified as short [25 s]), quality of persistence (whether the lingering flavors are pleasant, complex, and evolving or simple, harsh, or bitter), dominant aftertaste character (the specific descriptors persisting longest), and the final impression (clean, integrated, lingering warmth, drying, or bitter), providing the concluding quality assessment that experienced tasters weight most heavily in overall quality judgment.
Finish description captures the quality dimension that professional tasters and wine critics consistently identify as the single most reliable indicator of overall wine quality, with the correlation between finish length and expert quality scores stronger than for any other individual sensory parameter. The physiological basis of finish perception involves the sustained activation of gustatory, trigeminal, and olfactory receptors by taste-active compounds remaining on the oral mucosa, in saliva, and in the pharyngeal space after swallowing — high-extract wines leave more residual flavor-active material than dilute wines, tannin compounds continue to interact with salivary proteins over extended periods, and volatile compounds trapped in mucus layers and saliva continue to reach olfactory receptors retronasally for many seconds after swallowing. The quality of the finish — not merely its duration — is the critical evaluation criterion, as a long but unpleasant finish (dominated by bitterness, harsh astringency, or hot alcohol) indicates a flawed or unbalanced wine despite technically impressive persistence. The evolution of the finish — whether the persistent flavors remain static, develop new dimensions, or simply fade — provides additional quality information, with the most highly rated wines exhibiting "evolving" finishes where new aromatic and flavor facets emerge during the persistence phase. The distinction between "length" (the total duration of perceptible flavor) and "persistence" (the sustained intensity during the finish) adds nuance — some wines maintain high intensity that cuts off abruptly, while others diminish gradually but extend far longer. Temperature influences finish perception — cooler wines generally show shorter perceived finishes due to reduced vapor pressure and slower dissolution of residual compounds, while warmer wines may show artificially extended finishes through increased alcohol volatility. The concept of "caudalies" (from the French "caudale," a unit proposed for finish measurement where 1 caudalie equals 1 second of persistence) has not been widely adopted in formal tasting protocols but reflects the recognition that finish length is a quantifiable and quality-relevant parameter. Great wines from recognized terroirs often express site-specific mineral or textural signatures most clearly in the finish — the saline persistence of great Chablis, the iron-mineral aftertaste of Eisenberg Blaufränkisch, the graphite-tobacco persistence of Pauillac Cabernet — suggesting that the finish reveals terroir expression most transparently.
Abgang
Finish
Nachhall
Persistenz
aftertaste
length
lingering impression
palate finish
persistence
Peynaud E, Blouin J (2006) The Taste of Wine: The Art and Science of Wine Appreciation, 4th Ed, Wiley, pp.: 240-280 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 540-545 Gladstones J (2011) Wine, Terroir and Climate Change, Wakefield Press, pp.: 200-220 Saenz-Navajas MP et al. (2012) Modelling wine astringency from its chemical composition using machine learning approaches. Food Quality and Preference 25(2): 175-184
Finish Description
Finish length measurement in vertical tasting of Château Margaux reveals duration of 28 caudalies for the 2010 vintage (exceptional year, fully mature fruit), 19 caudalies for 2007 (challenging year, less concentration), and 32 caudalies for 2009 (outstanding year, optimal conditions), with the finish length ranking corresponding exactly to critic scores and release prices, and providing quantitative support for the quality tier assignments based on this single objective parameter.
Finish length quality is a taste persistence quality that represents the measured duration of perceptible taste sensation following swallowing, quantifying the temporal extent of the finish in units of time (typically seconds or caudalies) that serves as an objective persistence parameter in wine quality assessment. Within the BFO framework, this quality specifically addresses the quantitative duration dimension of finish, distinguishing measured length from the qualitative character of aftertaste and providing the numerical parameter most directly comparable across wines and assessment contexts. The ontological significance lies in finish length serving as one of the most reliable quality discriminators in wine evaluation, with the strong correlation between finish duration and quality tier reflecting the concentration and complexity required to sustain taste perception, and the relative objectivity of duration measurement reducing assessor variability compared to hedonic quality judgments. The measurement of finish length employs standardized protocols timing from swallow to return to baseline perception, with the caudalie unit (1 caudalie = 1 second) providing the traditional measure and typical values ranging from 3-5 seconds for simple wines through 8-12 seconds for quality wines to 15-25+ seconds for exceptional wines. The compositional determinants of finish length include extract concentration providing the material basis for sustained perception, phenolic compounds with slow receptor dissociation, and the overall complexity of the taste-active compound mixture where multiple components with different persistence characteristics create extended total duration. The relationship between finish length and other quality parameters shows strong positive correlation with critic scores, auction prices, and aging potential, supporting finish length as a primary quality indicator applicable across wine styles. Understanding finish length quality enables objective quality assessment complementing subjective evaluation, prediction of finish duration from compositional analysis, and quality tier classification based on quantitative persistence measurement.
Finish_Length_Quality provides the quantitative duration measurement of finish, complementing Aftertaste_Quality which addresses the qualitative character of post-swallow perception. This quality enables objective comparison across wines using standardized timing protocols. The caudalie unit, though traditional, simply equals seconds the choice of terminology depends on regional wine culture conventions.
Abgangslänge
Caudalie-Zahl
Finishlänge
Persistenzdauer
caudalie count
finish persistence
length of finish
persistence duration
Amerine MA, Roessler EB (1983) Wines: Their Sensory Evaluation, 2nd Ed. W.H. Freeman, New York, pp. 146-172 Robinson J, Harding J (2015) The Oxford Companion to Wine, 4th Ed. Oxford University Press, Oxford, pp. 274-275
Finish Length Quality
Thompson Seedless table grape firm outer mesocarp providing crisp texture= 4-6 cell layers, cells 35-58 μm diameter, walls 1.1-1.6 μm thick, creating firm zone 52-88 μm thickness beneath skin contributing to characteristic crisp firm texture desirable for table grape consumption, with firm outer mesocarp texture maintained during 3-4 week cold storage (0-2°C) essential for shipping durability, while softer middle mesocarp (cells 145-185 μm, walls 0.5-0.7 μm) provides juicy texture creating textural contrast between crisp skin/outer mesocarp and juicy soft flesh. Cabernet Sauvignon firm outer mesocarp in thick-skinned wine cultivar= cells 42-75 μm diameter (small), walls 1-1.4 μm (moderately thick), creating firm texture contributing to overall thick firm skin (total skin plus outer mesocarp 98-145 μm) requiring vigorous maceration during winemaking, with firm outer mesocarp cells breaking down during days 7-14 of extended 18-21 day maceration releasing tannins (3.5-5 mg/g) and contributing to full-bodied wine structure. Pinot Noir firm outer mesocarp in thin-skinned cultivar= cells 48-72 μm, walls 0.9-1.2 μm (thinner than Cabernet), creating moderately firm texture contributing to overall thin delicate skin (total 58-95 μm) requiring gentle maceration (5-10 days, minimal punchdown vigor) avoiding excessive extraction from limited firm outer mesocarp tissue, with firm outer mesocarp representing significant proportion of total skin thickness in thin-skinned cultivars (outer mesocarp 35-58 μm representing 40-65% of skin versus 30-45% in thick-skinned cultivars). Merlot firm outer mesocarp ripening changes= veraison (BBCH 81) showing cells 38-55 μm, walls 1.2-1.5 μm, firmness 4.5-6 N penetration force maturity (BBCH 89) showing cells 48-72 μm (27-45% expansion), walls 1-1.3 μm (slight thinning through expansion), firmness 2.8-4.2 N (38-47% reduction), demonstrating ripening-associated softening through cell expansion and wall thinning but firm outer mesocarp maintaining firmer texture than middle mesocarp (1.2-2 N at maturity) throughout ripening. Flame Seedless table grape firm outer mesocarp calcium effects= standard calcium nutrition showing firm outer mesocarp cells with walls 1.1-1.5 μm, firmness 5.2-7 N enhanced calcium (foliar sprays during development) showing walls 1.3-1.7 μm, firmness 6.5-8.5 N (18-25% increase), demonstrating calcium enhancement of cell wall properties and firm outer mesocarp texture important for table grape shipping quality through improved firmness and reduced decay susceptibility during post-harvest storage.
A firm outer mesocarp is an outer mesocarp emphasizing relatively firm texture compared to soft middle mesocarp, characterized by smaller cells (30-70 μm diameter versus 80-180 μm in middle mesocarp), thicker walls (0.9-1.5 μm versus 0.4-0.8 μm), reduced vacuolar volume (85-90% of cell volume versus 90-95%), and higher solid content creating firmer texture that contributes to skin firmness perception and provides structural support in transition from protective skin to soft flesh. Firm outer mesocarp textural characteristics= (1) firmness intermediate between firm skin (hypodermis, epidermis with thick walls and dense packing) and soft flesh (middle mesocarp with thin walls and large vacuoles) (2) cells maintaining higher turgor pressure relative to cell size creating firmer feel when compressed (3) thicker walls providing resistance to compression and deformation (4) smaller cell size requiring more cell wall material per unit volume creating higher structural density (5) reduced intercellular spaces (1-3% versus 2-5% in middle mesocarp) creating more compact tissue. Firm outer mesocarp cellular basis of firmness= (1) wall thickness-to-cell size ratio higher than middle mesocarp (wall=diameter ratio approximately 1=50 to 1=70 in firm outer mesocarp versus 1=150 to 1=400 in middle mesocarp) creating proportionally stronger cell walls relative to cell volume (2) turgor pressure potentially higher in smaller cells through osmotic effects creating resistance to deformation (3) wall composition potentially showing higher cellulose crystallinity or different pectin methylesterification affecting mechanical properties (4) cell-cell adhesion strength potentially higher through middle lamella properties affecting tissue cohesion. Firm outer mesocarp in berry texture perception= (1) contribution to skin firmness and toughness perceived during eating, with firm outer mesocarp adding to total skin thickness and firmness (2) influence on skin-flesh textural contrast creating transition from firm skin to soft flesh rather than abrupt boundary (3) contribution to overall berry firmness measured by penetrometry or compression testing (4) potential influence on berry splitting resistance through contribution to skin structural integrity. Firm outer mesocarp in different cultivar types= table grape cultivars sometimes showing particularly firm outer mesocarp (cells 30-55 μm, walls 1.1-1.6 μm) creating crisp firm texture desirable for fresh consumption and providing shipping durability wine grape cultivars showing variable firmness (some thick-skinned wine cultivars with firm outer mesocarp similar to table grapes, thin-skinned cultivars with less firm outer mesocarp) firmness potentially correlating with skin thickness and cell wall properties affecting cultivar suitability for different uses. Firm outer mesocarp developmental changes= early development showing firm outer mesocarp with small cells and thick walls ripening showing some softening through cell wall modifications (pectin depolymerization, hemicellulose breakdown) but outer mesocarp maintaining firmer texture than middle mesocarp throughout ripening over-ripening potentially showing excessive softening degrading firm texture and creating mealy or mushy texture undesirable for quality. Firm outer mesocarp in winemaking= firmer texture requiring more vigorous maceration conditions (higher temperature, more frequent punchdowns, enzymatic treatment) to break down cells and extract cellular contents compared to soft middle mesocarp firm outer mesocarp potentially releasing contents more slowly during maceration creating temporal extraction gradient with middle mesocarp extracting before firm outer mesocarp during gentle maceration.
Firm_Outer_Mesocarp emphasizes the textural properties of outer mesocarp and is distinguished from Structural_Outer_Mesocarp which emphasizes mechanical support functions. Firm outer mesocarp represents the outer mesocarp from the perspective of texture and eating quality. Understanding firm outer mesocarp is relevant for table grape breeding and production (selecting for desirable crisp texture), for berry texture assessment, and for winemaking extraction protocol design accounting for firmness differences between outer and middle mesocarp affecting extraction kinetics.
Festes hautnahes Fleisch
crisp sub-skin zone
feste äußere Fruchtfleisch-Schicht
festes äußeres Mesokarp
firm outer flesh
Thomas, T.R. et al. (2008). Changes in the physical properties of grape berries during development. American Journal of Enology and Viticulture 59=398-409 Considine, J.A., Kriedemann, P.E. (1972). Fruit splitting in grapes. Australian Journal of Agricultural Research 23=17-27.
Firm Outer Mesocarp
A firmly attached functional tendril at node 9 on a one-year cane of Vitis vinifera cv. Nebbiolo in Barolo, Italy, exhibiting seven extremely tight contact coils around a 4 mm diameter wire with coil diameters of 12-14 mm, complete dark brown lignification throughout all coiled regions visible six weeks after initial attachment, resisting 2.8 kilograms force in mechanical pull tests without detachment or coil deformation, supporting a 95 cm shoot section bearing nine leaves and three fruit clusters with combined weight of 720 grams through two severe thunderstorms with wind gusts exceeding 80 km/h without any displacement or loosening, requiring wire cutters and careful sectioning during dormant pruning because simple pulling would damage the cane bark, measured in January following the previous season's attachment at BBCH stage 71 (fruit set) in June.
An attached functional tendril exhibiting maximum grip strength and mechanical stability in Vitis vinifera, characterized by very tight coiling with contact coil diameters minimally exceeding support structure diameter, extensive lignification creating rigid tissue throughout coiled regions, multiple contact points generating high friction forces resisting displacement, and demonstrated load-bearing capacity substantially exceeding normal shoot and fruit weights typically supporting 1-3 kilograms or more without detachment or visible strain. Firmly attached functional tendrils develop through optimal contact and coiling conditions including sustained mechanosensory stimulation, appropriate support diameter matching tendril grasping range (2-8 mm), favorable temperature and moisture during the critical 24-72 hour coiling period enabling complete cell expansion and wall deposition, absence of mechanical disturbances during attachment formation, and sufficient time for thorough lignification to fix coil geometry permanently. These tendrils provide the most reliable mechanical support in vineyard systems, capable of maintaining shoot position through severe weather events including storms, heavy rain, and strong winds that may dislodge weakly attached or non-lignified tendrils. Firmly attached tendrils show remarkable longevity, with well-formed attachments potentially persisting for multiple seasons on perennial wood, and may require forceful removal during pruning operations, sometimes pulling bark or wood fragments from support structures when detached.
Firmly attached functional tendrils are distinguished from weakly attached tendrils through quantitative assessment of grip strength (resistance to measured detachment forces), qualitative evaluation of coiling quality (tightness, lignification, tissue integrity), and demonstrated performance under loading conditions (resistance to shoot weight, wind forces, or applied test loads). Firm attachment represents the ideal outcome of tendril development and is most common in mid-season tendrils on vigorous shoots with access to adequate resources. Environmental conditions strongly influence attachment strength, with water stress, nutrient deficiency, or temperature extremes during the critical coiling period potentially preventing achievement of firm attachment despite apparently normal morphology. Firmly attached tendrils may become so integrated with support structures that removal damages both tendril and support, creating pruning challenges in some training systems.
fest verankerte Ranke
high-strength attachment tendril
hochfeste Greifranke
secure grip tendril
stark befestigte Ranke
strongly attached tendril
Based on Silk & Hubbard (1991) "Axial forces and normal stresses in coiling tendrils" Biomechanics and Modeling in Mechanobiology 24=599-606 Bowling & Vaughn (2009) "Gelatinous fibers are widespread in coiling tendrils and twining vines" American Journal of Botany 96=719-727 Isnard & Silk (2009) "Moving with climbing plants from Charles Darwin's time into the 21st century" American Journal of Botany 96=1205-1221 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 195-218.
Firmly Attached Functional Tendril
Cabernet Sauvignon in Napa Valley shows firmness decline during September-October Year N+1 at BBCH 85-89, with penetrometer firmness declining from pre-veraison maximum 3.8 N through initial softening 3.0 N to progressive ripening reaching mature 1.8 N over 7-8 weeks, total firmness reduction 53% from maximum, progressive decline rate 0.8% per day during ripening, firmness strongly inversely correlated with water-soluble pectin (r²=-0.91) and polygalacturonase activity (r²=-0.88), firmness decline driven primarily by cell wall pectin degradation 78% contribution with minor turgor 22%, firmness used to guide harvest timing for wine production, under warm climate conditions of 22-30°C temperature, typical of this late-season cultivar achieving target soft texture at harvest maturity
Firmness decline represents the progressive mechanical resistance reduction phase during progressive softening when berry penetrometer firmness declines from initial softening 2.0-3.5 N through ripening to mature 1.0-2.5 N, total firmness reduction 40-70% from pre-veraison maximum 2.5-4.5 N, firmness declining progressively at rate 0.5-1.5% per day over 4-8 weeks ripening, and firmness decline as quantitative measure of progressive softening establishing soft ripe berry texture. This mechanical resistance reduction phase occurs during late summer through autumn in late August-October in temperate regions Year N+1, representing firmness decrease through ripening. Firmness decline mechanism involves progressive cell wall structural weakening from sustained pectin degradation reducing mechanical resistance, middle lamella loss reducing tissue cohesion, possible minor turgor contribution though structural changes dominant, cumulative softening effect accumulating over ripening weeks, and firmness decline as quantitative softening measure. Firmness measurement shows penetrometer or texture analyzer measuring force required to penetrate berry, measurement quantifying firmness objectively, force in Newtons N, standard assessment method, and measurement as firmness quantification. Penetrometer technique shows probe typically 2-5 mm diameter penetrating berry, force recorded at standard depth or at berry failure, measurement on individual berries, and technique as standard method. Compression testing shows alternative firmness measurement through berry compression, force at percentage deformation, compression resistance correlating with penetrometer firmness, and compression as firmness measure. Firmness trajectory shows firmness declining from pre-veraison maximum 2.5-4.5 N through initial softening 2.0-3.5 N to progressive ripening decline reaching mature 1.0-2.5 N, trajectory showing rapid initial decline then slower progressive decline, eventual plateau at ripe levels, and trajectory as softening pattern. Pre-veraison firmness maximum shows berry at peak firmness during lag phase, firmness from turgid cells with rigid pectin-rich walls, maximum as baseline for decline measurement, and peak as starting point. Initial softening firmness shows firmness declining 10-25% at veraison to 2.0-3.5 N, initial decline establishing softening, and initial state as progressive baseline. Progressive decline shows firmness continuing decline through ripening 4-8 weeks, typical decline 0.5-1.5% firmness per day representing slower rate than initial 2-4% per day, cumulative decline substantial, and progressive reduction as ripening feature. Mature ripe firmness shows firmness stabilizing at ripe levels 1.0-2.5 N, final firmness varying by cultivar and maturity, ripe firmness as endpoint, and mature state as firmness target. Total firmness reduction shows cumulative decline 40-70% from pre-veraison to ripe, total reduction representing substantial softening, total as overall change measure, and reduction magnitude as softening extent. Firmness decline rate shows progressive decline rate 0.5-1.5% per day during ripening, rate slower than initial rapid decline, rate relatively constant through ripening, and rate as decline kinetics. Decline rate variation shows rate varying with temperature, cultivar, and conditions, warm conditions promoting faster decline, cool slowing decline, and variation as environmental response. Firmness-softening correlation shows firmness inversely correlated with softening, declining firmness indicating increasing softness, strong correlation enabling firmness as softening measure, and relationship as measurement basis. Firmness-cell wall relationship shows firmness strongly correlated with cell wall pectin content and structure, declining firmness correlating with pectin degradation and solubilization, cell wall changes as firmness determinant, and correlation supporting structural basis. Pectin-firmness correlation shows water-insoluble pectin content positively correlated with firmness, soluble pectin negatively correlated, pectin molecular weight positively correlated, and correlations showing pectin as firmness factor. Enzyme activity-firmness relationship shows polygalacturonase and pectin methylesterase activities negatively correlated with firmness, high enzyme activities correlating with low firmness, and enzymes as decline drivers. Firmness-turgor relationship shows firmness weakly correlated with turgor, turgor contributing minor 15-30% variance in firmness, structural factors dominant 70-85%, and relationship showing turgor as secondary factor. Firmness variation shows firmness varying among berries in cluster from asynchronous ripening, within-cluster variation creating firmness distribution, variation affecting harvest decisions, and heterogeneity as practical issue. Cultivar firmness differences shows ripe firmness varying among cultivars, range 1.0-2.5 N representing genetic variation, some cultivars maintaining firmer texture others very soft, and differences as genetic trait. Temperature effects on decline shows temperature strongly affecting firmness decline rate, warm conditions accelerating decline, cool conditions slowing decline, temperature as major environmental factor, and rate variation as temperature response. Water effects shows adequate water supporting normal firmness decline, water deficit possibly affecting decline through turgor effects, and water as secondary environmental factor. Firmness as maturity indicator shows declining firmness indicating ripening progression, firmness used for maturity assessment, target firmness varying by use, and firmness as practical maturity measure. Harvest timing shows firmness guiding harvest decisions, table grapes harvested at firmer texture 1.8-2.5 N, wine grapes at softer 1.0-1.8 N, and firmness as harvest criterion. Firmness-quality relationships shows firmness affecting quality depending on use, optimal firmness varying by application, firmness critical for table grape quality, less critical for wine grapes, and quality as firmness context. Sensory-instrumental correlation shows instrumental firmness correlating with sensory texture assessment, correlation enabling instrumental prediction of sensory, and correlation as practical tool. Firmness preservation shows cool storage slowing firmness decline post-harvest, temperature management preserving texture, and preservation as post-harvest practice. The process represents third aspect of progressive softening establishing quantitative softening measure, following continued cell wall degradation and concurrent with increasing berry pliability. Duration firmness declining progressively over 4-8 weeks ripening. Regional timing shows firmness decline during late August-October through ripening to harvest. Viticultural significance includes representing quantitative softening measure enabling objective assessment, determining harvest maturity and berry quality, demonstrating progressive cell wall degradation outcome, and establishing firmness as ripeness indicator. Research applications include analyzing firmness decline kinetics and mechanisms, investigating firmness-structural relationships and correlations, evaluating environmental effects on decline rate, quantifying firmness changes and variation, and assessing firmness as quality and maturity determinant.
Firmness_Decline is third aspect of Progressive_Softening, distinguished by quantitative resistance reduction. Represents 4-8 weeks ripening. Firmness 2.0-3.5 → 1.0-2.5 N (40-70% total decline from pre-veraison 2.5-4.5 N). Rate 0.5-1.5%/day (slower than initial 2-4%/day). Mechanism: cell wall degradation 70-85%, turgor 15-30%. Strong correlation with pectin solubilization (r²>0.85). Cultivar variation 1.0-2.5 N ripe. Temperature major rate factor. Follows Continued_Cell_Wall_Degradation. Concurrent with Increasing_Berry_Pliability. Completes Progressive_Softening and Berry_Softening. Maturity and quality indicator.
Textur-Erweichungs-Messung
firmness decrease
mechanical resistance decline
mechanische Widerstands-Abnahme
progressive Festigkeits-Reduktion
progressive firmness reduction
texture softening measurement
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 485-490 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Firmness Decline
Firmness quality evolution in Crimson Seedless table grapes guides harvest timing for export markets, with optimal firmness of 280-350 g (8mm probe puncture test) providing the crisp texture and handling durability required for 6-8 week cold storage plus retail display harvest below 250 g produces soft, perishable fruit unsuitable for long-distance markets, while premature harvest above 400 g yields immature flavor despite acceptable texture, requiring growers to balance firmness maintenance against flavor development in this slow-softening variety.
Firmness_Quality encompasses the physical qualities characterizing the resistance of grape tissue to deformation under applied force, capturing the textural property most directly relevant to handling durability, processing behavior, and eating quality perception. Firmness represents the integrated mechanical response of grape tissue structure, reflecting cell turgor pressure, cell wall rigidity, intercellular adhesion, and tissue architecture. This category subdivides into Tissue_Firmness_Quality addressing firmness of specific anatomical structures (berry, pulp, skin) Compression_Resistance_Quality capturing response to compressive loading Puncture_Resistance_Quality describing resistance to penetrating probes and Deformation_Quality characterizing the nature of shape change under stress. Firmness changes substantially during grape development, with berries softening dramatically at véraison as cell walls are modified and turgor dynamics shift, then potentially continuing to soften during late ripening as cell wall degradation progresses. The rate and extent of softening vary among cultivars, with crisp-textured table grape varieties maintaining firmness longer than soft wine grape varieties. Environmental factors including temperature, water status, and disease pressure modulate firmness trajectories within cultivar-determined limits. Firmness affects quality throughout the production chain: firm berries resist mechanical damage during harvest and transport appropriate firmness enables efficient juice release during crushing while preventing excessive solid extraction and textural firmness provides the crispness valued in table grapes. Firmness measurement employs instrumental texture analysis with compression, puncture, or tensile test geometries, generating force-deformation curves from which firmness parameters (maximum force, slope, work) are extracted. The correlation between instrumental firmness and sensory firmness perception requires validation for specific cultivar and measurement conditions, as the relationship depends on test geometry, sample preparation, and the specific sensory attribute assessed.
Firmness_Quality serves as the parent class for the firmness-related quality hierarchy, positioned within Physical_Quality. The class addresses firmness as a general concept encompassing multiple measurement approaches and tissue types. Firmness is technically distinct from hardness (resistance to permanent indentation) though the terms are often used interchangeably in food texture literature. The instrumental measurement of firmness provides objectivity but requires standardization of test parameters (probe size, speed, sample preparation) for comparable results across studies and applications.
Festigkeit
Gewebefestigkeit
Textur-Festigkeit
hardness quality
mechanical resistance
mechanischer Widerstand
texture firmness
tissue rigidity
Rolle L et al. (2012) Instrumental Texture Analysis Parameters as Markers of Table-Grape and Winegrape Quality: A Review. American Journal of Enology and Viticulture 63(1): 11-28 Letaief H et al. (2008) Assessment of Grape Skin Hardness by a Puncture Test. Journal of the Science of Food and Agriculture 88(9): 1567-1575
Firmness Quality
A first basal leaf at node 1 on a spur-pruned shoot of Vitis vinifera cv. Pinot Noir in Burgundy, France, emerging 5 days post-budbreak as the first visible leaf on the developing shoot, measuring 18.5 cm length by 17.2 cm width at maturity representing 35% larger area than leaves at nodes 7-9, exhibiting somewhat simplified five-lobed architecture with moderate rather than deep sinuses characteristic of this cultivar at node 1 position, achieving full expansion at 24 days post-budbreak (compared to 32-38 days for median leaves), demonstrating peak photosynthesis rate of 14.2 μmol CO₂ m⁻² s⁻¹ in early May when it represented the only mature leaf on the shoot contributing 100% of photosynthate production supporting shoot extension from 15 cm to 45 cm and early inflorescence development, positioned opposite the first inflorescence (if present) or tendril following species-typical phyllotactic pattern, experiencing progressive shading as 12 additional leaves developed above creating light reduction from 90% full sun to 25% by mid-July, showing senescence initiation in early August with yellowing and reduced function, measured at shoot BBCH stage 77 (berries beginning to touch), representing the unique first leaf position with maximum early-season contribution and typical progressive decline.
A basal leaf at node 1 representing the first true foliage leaf on a developing shoot in Vitis vinifera, characterized by position immediately above the point of shoot origin from the previous season's cane or spur, earliest emergence among all shoot leaves typically visible 3-7 days post-budbreak, development from preformed primordium created during previous growing season and overwintered within dormant bud, often the largest leaf on the shoot with dimensions frequently 20-40% greater than median position leaves reflecting developmental priority as first photosynthetic organ, distinctive morphological characteristics sometimes including reduced lobe development or simplified architecture compared to median leaves depending on cultivar, very early achievement of maturity typically 3-4 weeks post-budbreak providing critical early-season photosynthetic capacity, disproportionately high contribution to early shoot carbon balance often providing 30-50% of total shoot photosynthate during the first 4-6 weeks following budbreak when it may represent the only or dominant mature photosynthetic surface, and eventual decline through shading, senescence, or intentional removal during canopy management. First basal leaves play unique developmental and functional roles, emerging from the most developed primordium within dormant buds positioned closest to vascular connections with permanent vine structures enabling rapid resource mobilization, experiencing optimal early-season environmental conditions before canopy development creates competition and shading, and providing the foundational photosynthetic capacity that supports initial shoot extension, early inflorescence development, and mobilization of stored reserves for current season growth.
First basal leaves are distinguished from all other leaves by node 1 position and earliest emergence timing, representing the developmental pioneer establishing initial shoot photosynthetic capacity. Morphological characteristics at node 1 sometimes differ from cultivar-typical patterns, with some varieties showing reduced lobe development, altered sinus depth, or modified overall shape at this position compared to nodes 3-8 where characteristic morphology is most reliably expressed. The first basal leaf often shows the largest dimensions on the shoot, though this varies with shoot vigor and environmental conditions, with vigorous shoots sometimes producing even larger leaves at nodes 2-4. Functional importance of first basal leaves is greatest during the first 3-5 weeks post-budbreak when they may represent 50-100% of shoot photosynthetic capacity, declining progressively as upper canopy develops. Management decisions about first basal leaf removal must consider timing relative to shoot development stage, with early removal (pre-bloom) significantly impacting carbohydrate availability for fruit set while late removal (post-fruit set) has reduced impact on reproductive development but affects fruit exposure and microclimate.
Knoten-eins Blatt
erstes basales Blatt
initial shoot leaf
node one leaf
primary basal leaf
primäres Spross-Weinblatt
Based on Poni et al. (2006) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 57=397-407 Palliotti et al. (2011) "Vine performance and grape composition as affected by early-season source limitation induced with anti-transpirant and shading in two red Vitis vinifera L. cultivars" Australian Journal of Grape and Wine Research 17=368-375 Intrieri et al. (2008) "Grapevine growth and development" in Biology of the Grapevine, Cambridge University Press pp. 44-81 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 95-118.
First Basal Leaf
Pinot Noir in Burgundy shows first calyptra detachment on June 3 Year N+1 at BBCH 61, with single flower in apical cluster position completing cap shedding during morning hours after accumulating 890-920 growing degree days base 10°C, exposing fresh reproductive organs under continental conditions of 19-23°C temperature, 58-68% relative humidity, marking definitive flowering onset for this early-to-mid season cultivar approximately 12 days before complete anthesis
First calyptra detachment represents the threshold flowering event when earliest-maturing flower completes abscission zone activation and sheds petal cap exposing reproductive organs, establishing definitive anthesis onset and serving as precise phenological marker for flowering date determination. This critical milestone event occurs during early flowering in early June in temperate regions Year N+1, typically representing single moment or brief hours when first individual flower achieves cap detachment, corresponding to BBCH 61 defining characteristic. Detachment completion in first flower shows characteristic final sequence with abscission zone fully degraded creating complete separation plane, calyptra lifted 1-2 mm above receptacle by expanding reproductive organs, mechanical disturbance from wind, turgor change, or natural movement triggering final detachment, and cap falling away from flower exposing stamens and pistil. First flower location typically in apical cluster position reflecting earliest primordium formation previous season and most advanced development, occasionally in middle regions if developmental variation creates alternative earliest-maturing flower, and spatial positioning providing indicator of developmental patterns within cluster. Observational requirements for detecting first detachment include careful cluster inspection as single cap fall may be inconspicuous, timing of observations with morning assessment most likely to detect fresh overnight detachment, and distinction between true first detachment versus cap loosening without complete separation. Detached cap characteristics include fresh appearance with green or yellow-green color indicating recent separation, intact fused petal structure showing pentamerous organization, moist attachment scar suggesting recent abscission completion, and positioning on cluster surface or recently fallen to ground below. Exposed flower appearance at first detachment shows newly revealed reproductive organs with pristine fresh condition, stamens positioned around central pistil showing beginning maturity, anthers showing early color transition from green toward yellow indicating approaching dehiscence, and stigma showing developing receptivity with papillate surface beginning exudate production. Environmental conditions at first detachment typically favorable for abscission with temperature 18-26°C supporting enzymatic activity and organ function, humidity 50-70% facilitating cap loosening without retention, and overall weather enabling normal flowering initiation establishing pattern for subsequent detachments. Timing predictability shows first detachment occurring at relatively consistent thermal time accumulation with typical range 900-1100 growing degree days base 10°C depending on cultivar and regional climate, though exact timing affected by short-term weather affecting final abscission completion. Phenological significance includes defining flowering date with first calyptra detachment establishing BBCH 61 stage, providing precise reference for phenological records and climate studies, and establishing baseline for heat summation calculations and subsequent stage predictions. Developmental implications show first detachment confirming successful completion of previous season primordium formation and current season inflorescence development, indicating reproductive readiness in earliest-maturing flowers, and marking transition from pre-bloom to bloom state requiring management adjustments. Observational challenges include detection difficulty with single inconspicuous event requiring vigilant monitoring, potential confusion between cap loosening and actual detachment requiring verification of complete separation, and timing uncertainty as exact moment may occur between observations requiring retrospective assessment. Documentation importance emphasizes recording precise first detachment date as critical phenological marker for vineyard records, comparison across seasons revealing phenological trends and climate effects, and standardization of observation protocols ensuring data consistency. Weather window implications show that conditions at first detachment indicate likely conditions during subsequent mass flowering, with favorable weather at onset suggesting good flowering potential while unfavorable conditions portending potential problems. Management implications include confirming flowering onset requiring final preparations for bloom period, cessation of disruptive canopy operations to avoid flower damage, and intensification of weather monitoring as critical reproductive period begins. Subsequent progression from first detachment shows rapid acceleration with additional flowers detaching within hours to days, progressive spatial expansion from initial location toward broader cluster involvement, and transition toward mass cap fall during peak anthesis typically 2-4 days after first detachment. Cultivar differences in first detachment timing show early varieties achieving onset late May-early June while late types show mid-late June timing reflecting overall phenological variation, with timing affected by accumulated thermal units, chilling satisfaction, and cultivar-specific heat requirements. Regional variation shows first calyptra detachment during late May-early June in warm climates with heat accumulation reaching thresholds earlier, early-mid June in temperate regions with moderate heat accumulation, and mid-late June in cool areas requiring longer periods for thermal unit accumulation. Climate effects on timing include warmer years showing earlier first detachment reflecting accelerated heat accumulation, cooler years showing delayed timing, and interannual variation in timing reflecting climate variability affecting viticultural calendars and management scheduling. Research applications include documenting precise flowering phenology using first detachment as objective marker, analyzing environmental controls of flowering onset including thermal time requirements and short-term weather effects, evaluating genetic variation in flowering timing affecting adaptation and regional suitability, developing phenological models predicting flowering dates from climate variables, and assessing climate change impacts on flowering timing across regions and seasons. The process represents first aspect of initial cap fall marking definitive flowering onset, followed by early cap shedding expanding activity. Duration represents threshold moment occurring over hours to single day as first flower completes detachment. Cultivar variation in first detachment characteristics includes timing differences reflecting overall phenology, flower position showing cultivar-specific developmental patterns, and detachment efficiency indicating genetic differences in abscission processes. Viticultural significance includes establishing definitive flowering date for phenological records, providing practical observable marker for bloom onset enabling management transitions, indicating beginning of weather-sensitive period requiring monitoring, and serving as reference for predicting subsequent flowering progression and developmental timing through season. Research applications include documenting flowering phenology with precision, analyzing mechanisms triggering abscission onset, evaluating cultivar and environmental effects on flowering timing, developing predictive models for flowering dates supporting management planning, and assessing climate change effects on phenological timing affecting viticultural adaptation.
First_Calyptra_Detachment is first aspect of Initial_Cap_Fall, distinguished by threshold flowering event with earliest cap detachment. Corresponds to BBCH 61 defining characteristic. Represents precise phenological marker for flowering date. Brief threshold moment occurring over hours to single day. Critical for phenological records and management transitions. Detection requires careful observation. Establishes reference for subsequent flowering progression.
Blühzeitpunkt
Erstblüte
erste Kalyptralösung
first bloom
first cap fall
flowering onset moment
initial cap detachment
initiale Kappentrennung
Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 360-365
First Calyptra Detachment
In a Willamette Valley Pinot Noir vineyard, first color change was documented in early September Year N+1 (BBCH 83, 63 days post-flowering) when 15% of berries showed initial red patches on sun-exposed surfaces, anthocyanin concentration measured 42 mg/kg skin (from <5 mg/kg pre-veraison), UFGT expression was 78-fold elevated, berry sugar reached 10.2° Brix, and ABA levels were 920 ng/g under conditions of warm days (25-28°C) and cool nights (11-14°C) with partial cluster exposure to morning sun.
First_Color_Change represents the earliest visible manifestation of anthocyanin accumulation during veraison onset when initial red-purple pigmentation becomes detectable on berry surfaces, typically appearing as small discrete patches or sectors on individual berries within the cluster marking the beginning of the dramatic color transformation from green to fully pigmented fruit. This initial change occurs during early veraison (BBCH 83), usually 1-3 days following molecular and biochemical initiation of anthocyanin biosynthesis, and becomes visible when anthocyanin concentration in epidermal and sub-epidermal cell vacuoles reaches 30-60 mg/kg skin fresh weight (compared to 35°C inhibits anthocyanin synthesis causing delayed or suppressed first color change even under high light exposure. Cultivar variation in first color change characteristics reflects genetic differences in anthocyanin biosynthetic regulation: early-coloring varieties like Pinot Noir show first color 8-9 weeks post-flowering with rapid appearance across berry surface (1-2 days from first patches to 50% colored), Merlot and Syrah display intermediate timing (9-11 weeks) with moderate progression (2-4 days to 50% coverage), while late cultivars like Cabernet Sauvignon initiate first color 11-13 weeks post-flowering with slower spread (3-6 days to 50%) reflecting lower initial anthocyanin synthesis rates and potentially different regulatory control. First color change serves as practical viticultural marker: clusters showing 10-30% berries with visible color patches define veraison onset for vineyard monitoring, the transition from 0% to 50% berries showing color typically requires 2-5 days in uniform blocks providing harvest timing forecasts (harvest typically 40-60 days post-first color depending on cultivar and heat accumulation), and within-vineyard spatial variation in first color timing reveals maturity heterogeneity requiring management decisions about selective harvest or acceptance of compositional variation. Cluster architecture affects first color patterns: compact clusters with berry-to-berry contact may show color first on exposed exterior berries while interior berries remain green for extended periods, loose clusters with better air circulation and light penetration show more uniform first color appearance across all berries, and cluster position on the canopy creates systematic variation with exterior clusters coloring 2-4 days before interior positions in dense canopies. Anthocyanin composition at first color change establishes cultivar-specific profiles: malvidin-3-glucoside typically comprises 50-70% of total anthocyanins even at first color reflecting preferential UFGT substrate specificity and methyltransferase activities, relative proportions of delphinidin, cyanidin, petunidin, and peonidin derivatives are established early and remain relatively constant through complete coloring, and acylated anthocyanins (acetyl, coumaroyl derivatives) begin accumulating providing enhanced color stability and modified hue characteristics. Water stress effects on first color are complex: moderate deficit (stem ψ -0.9 to -1.2 MPa) can advance first color by 2-4 days through enhanced ABA signaling and potentially reduced berry size concentrating pigments, severe stress (<-1.4 MPa) delays first color through metabolic impairment despite elevated ABA levels, while excess water can delay color 3-5 days through berry dilution and reduced ABA signaling suppressing anthocyanin gene induction. Physiological significance of first color change includes: anthocyanins provide UV protection as chlorophyll content declines reducing photo-oxidative damage risk, antioxidant properties of anthocyanins and phenolic intermediates protect cellular components during metabolic transitions, and visible color signals birds and other animals potentially affecting berry predation patterns though commercial viticulture typically prevents animal access. Monitoring first color change informs management decisions: irrigation can be reduced or terminated at first color in deficit irrigation strategies since berry growth slows substantially post-veraison, disease management intensifies as berry sugar increases and skin integrity decreases creating favorable Botrytis conditions, and canopy management to improve cluster exposure may be adjusted recognizing that excessive exposure during active color development risks heat inhibition while moderate exposure enhances coloring through light signals.
First_Color_Change is distinguished from Color_Change_Onset by representing visible detection rather than molecular initiation, from Progressive_Color_Development by showing initial patches rather than spreading coloration, and from Complete_Color_Change by exhibiting 10-30% berry coverage rather than full pigmentation. Anthocyanin threshold for visibility: 30-60 mg/kg skin. Timing: typically 1-3 days after molecular onset. Spatial pattern: sun-exposed surfaces and stylar end first. Temperature optimum 24-30°C berry surface. First_Color_Change serves as practical veraison marker for vineyard monitoring and harvest forecasting.
Farbbruch-Beginn
color break onset
earliest anthocyanin detection
erste Rotflecken-Erscheinung
first red patch appearance
früheste Anthocyan-Detektion
initial visible pigmentation
initiale sichtbare Pigmentierung
Keller M (2020) The Science of Grapevines 3rd Ed pages 233-234 color development patterns Downey MO, Dokoozlian NK, Krstic MP (2006) Cultural practice and environmental impacts on the flavonoid composition of grapes and wine Plant Physiol Biochem 44:170-181 anthocyanin accumulation dynamics Mori K, Goto-Yamamoto N, Kitayama M, Hashizume K (2007) Loss of anthocyanins in red-wine grape under high temperature J Exp Bot 58:1935-1945 temperature effects on initial coloring.
First Color Change
Chardonnay in Champagne shows first flowers opening on June 4 Year N+1 at BBCH 61, with 5-8% flowers in apical cluster positions completing cap shedding exposing reproductive organs during morning hours, under cool maritime climate conditions of 18-22°C temperature, 60-75% relative humidity, after accumulating 880-920 growing degree days base 10°C, typical of this early-bursting cultivar establishing flowering onset for subsequent bloom progression
First flowers opening represents the definitive flowering initiation milestone when earliest-maturing flowers complete cap shedding and expose functional reproductive organs, marking unambiguous transition from pre-bloom to bloom state and establishing reproductive phase onset within cluster. This critical threshold event occurs during early June in temperate regions Year N+1 as part of early anthesis, typically representing single day or 1-2 day period when first 5-10% flowers achieve open state, corresponding to BBCH 61 phenological stage. Initial opening characteristics show first opened flowers typically in apical cluster positions reflecting earliest primordium formation and most advanced development, with flowers exhibiting freshly shed caps, newly exposed stamens and pistil showing delicate pristine appearance, and beginning anther color development from green toward yellow-brown indicating approaching pollen maturity. Flower identification at opening shows opened flowers clearly distinguishable from unopened buds through absence of calyptra cap exposing reproductive structures, visible stamens arranged around central pistil, and flower diameter expanded to 4-5 mm compared to 3-4 mm unopened buds. Cap detachment completion in first-opening flowers shows calyptra completely separated and fallen away, typically found on cluster surface or ground below, with detached caps showing characteristic intact fused petal structure and attachment scar where abscission occurred. Reproductive organ exposure reveals functional anatomy with five stamens showing anthers on short filaments positioned around pistil, anthers 0.8-1.2 mm length with bilocular structure containing pollen in two compartments separated by connective tissue, pistil centrally positioned with superior ovary, short style, and bilobed stigma surface. Anther status at flower opening shows approaching dehiscence readiness with anthers beginning color transition indicating pollen maturation, anther wall structure showing preparation for stomium opening, though actual pollen release typically occurs hours to days after flower opening depending on environmental conditions particularly humidity. Stigma appearance at opening shows developing receptivity with papillate surface cells visible under magnification, beginning exudate production creating slight glistening appearance, and stigma lobes positioned above anthers facilitating pollen capture from wind-dispersed pollen or direct contact with nearby anthers. Nectary visibility in opened flowers shows disc structure at flower base surrounding ovary, with some flowers showing active nectar secretion visible as small droplets though production is variable among flowers and cultivars. Flower timing shows first opening typically occurs during morning hours when temperature rises and humidity declines creating favorable conditions for cap shedding, with subsequent flowers throughout cluster following similar diurnal pattern with most opening occurring morning to mid-day. Developmental gradient within cluster shows earliest flowers opening 12-24 hours before neighboring flowers, creating progressive opening wave that will extend over subsequent 5-10 days encompassing entire cluster, with gradient reflecting primordium formation sequence from previous season. Environmental conditions at first opening indicate likely conditions during subsequent peak flowering, with favorable warm dry conditions at first opening suggesting good flowering potential, while unfavorable cool wet conditions may portend problems for bloom period. Temperature at first opening typically ranges 18-26°C supporting normal cap shedding and organ function, with opening delayed if temperatures remain below 15°C and potentially accelerated if temperatures exceed 28°C. Humidity at first opening preferably 40-70% relative humidity supporting cap detachment and organ function without excessive drying, with high humidity above 75% potentially preventing some caps from shedding cleanly and very low humidity below 35% causing rapid tissue desiccation. Weather sensitivity at first opening is extreme with opened flowers highly vulnerable to rain causing physical damage, preventing normal organ function, and promoting fungal infection particularly botrytis, making weather monitoring critical at this stage. Metabolic status at first opening shows elevated respiration in opened flowers supporting reproductive processes, active sugar metabolism providing energy for organ function and potential nectar production, and specialized anther metabolism completing final pollen maturation. Resource availability at opening requires adequate carbohydrate reserves supporting energy-intensive reproductive processes, with flowering success depending partially on vine reserve status affected by previous season crop load and current season photosynthetic production. Phenological significance of first flowers opening includes defining definitive flowering date used for heat summation calculations and phenological tracking, providing baseline for predicting subsequent peak anthesis timing typically 2-4 days later, and establishing critical management period requiring operational adjustments. Observational requirements for identifying first flowers opening include careful cluster inspection as initial opening may involve only few flowers at apical positions, appropriate timing with morning observations most likely to detect fresh opening, and distinguishing truly opened flowers with exposed organs from buds showing cap loosening but not yet detached. Documentation importance emphasizes recording precise first flower opening date as critical phenological marker for vineyard records, climate studies, and management planning, with date providing key reference for subsequent developmental predictions. The process represents first of three early anthesis aspects marking definitive flowering onset, followed by progressive cap shedding in additional flowers and stigma receptivity development. Duration represents threshold event occurring over 1-2 days as first 5-10% flowers open, with rapid transition toward increasing opening frequency during subsequent early anthesis. Cultivar variation appears in first opening timing with early varieties showing late May-early June opening while late types show mid-late June opening reflecting overall phenological differences, cluster position of first opening with some varieties showing consistent apical opening while others show more variable patterns, and environmental sensitivity affecting opening under marginal conditions. Regional timing shows first flowers opening during late May-early June in warm climates at approximately 950-1050 growing degree days base 10°C, early June in temperate regions at 850-950 GDD, and mid-June in cool areas at 750-850 GDD, with timing critical for frost avoidance and favorable weather. Viticultural significance includes defining flowering date for phenological records and management scheduling, providing early warning of flowering conditions affecting subsequent bloom success, establishing critical observation point requiring careful monitoring, and marking transition requiring operational adjustments including restricting fungicide applications and canopy operations. Research applications include documenting precise flowering onset for phenology and climate studies, analyzing environmental triggers of first opening affecting phenological variation, evaluating developmental mechanisms determining first flower identity and timing, quantifying relationships between first opening and subsequent bloom progression, and assessing climate change impacts on flowering dates across regions.
First_Flowers_Opening is first of three Early_Anthesis aspects, distinguished by definitive flowering onset with first flowers achieving open state. Represents threshold phenological event corresponding to BBCH 61. Followed by progressive Initial_Cap_Shedding affecting more flowers and Stigma_Receptivity_Onset. Critical phenological marker for records and management. Timing highly sensitive to temperature and accumulated heat units.
Anthesestart
Blütenbeginn
anthesis initiation
erste Blüten öffnen
flowering start
initial flower opening
First Flowers Opening
Seyval Blanc (Seyve-Villard 5-276, 1920s origin, commercial from 1950s) bears a First_Generation_PIWI_Role as an early disease-tolerant hybrid predating formal PIWI classification, with neutral wine character suited to blending and sparkling production in England and eastern North America. Léon Millot (Kuhlmann 194-2, 1911) serves in a First_Generation_PIWI_Role as an early cold-hardy resistant hybrid, still planted in continental climates for its reliable production despite wine quality limitations compared to contemporary PIWIs. Rondo (released 1999, Geisenheim, but from 1970s breeding) bears a First_Generation_PIWI_Role as an early German resistant red variety, offering good disease resistance and cold hardiness with wine character acceptable for regional production but not approaching V. vinifera standards.
A PIWI cultivar role that inheres in early-generation disease-resistant varieties developed from the 1970s through 1990s, representing initial efforts to combine interspecific disease resistance with commercially acceptable wine quality. First-generation PIWIs emerged from breeding programs at German, French, and Swiss institutions that crossed complex French-American hybrids (themselves carrying resistance from earlier interspecific work) with V. vinifera varieties, producing varieties with effective field resistance but often retaining "hybrid character" in wine (atypical aromas, excessive acidity, unfamiliar phenolic profiles). These varieties represented proof-of-concept that resistance breeding could produce wines approaching commercial viability, though quality limitations restricted adoption primarily to niche markets and sustainability-focused producers. First-generation PIWIs remain commercially significant in challenging climates where V. vinifera cultivation is difficult (northern Europe, continental regions), and several serve as parents in continued breeding programs seeking to improve wine quality in subsequent generations. The designation "first generation" is approximate, reflecting the iterative nature of breeding where generation boundaries are not precisely defined.
First_Generation_PIWI_Role captures the foundational resistant varieties that demonstrated resistance breeding viability while revealing the quality challenges requiring subsequent improvement. The generation designation is necessarily imprecise—some French-American hybrids predate formal resistance breeding yet function as first-generation PIWIs, while varieties released in the 1990s-2000s but derived from 1970s crosses straddle generational boundaries. First-generation PIWIs typically carry resistance from complex hybrid ancestors (Seibel, Seyve-Villard, Kuhlmann selections) without the targeted R-gene introgression characteristic of later generations, resulting in multigenic resistance that may be durable but is accompanied by substantial non-vinifera genome contribution. Commercial success of first-generation PIWIs varies regionally—widely adopted where V. vinifera is not viable (England, Scandinavia, upper Midwest USA), limited penetration in traditional wine regions with established V. vinifera production.
PIWI erste Generation
Pionier-Resistenzsorte
early PIWI
first-generation PIWI
first-wave resistant variety
frühe PIWI-Sorte
pioneering resistant variety
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Pedneault K, Provost C (2016) Fungus resistant grape varieties, Applied Microbiology and Biotechnology 100: 3499-3519 Reynolds A (ed) (2015) Grapevine Breeding Programs for the Wine Industry, Elsevier PIWI International (2020) PIWI Variety Database
First Generation PIWI Role
Pinot Noir in Burgundy shows first green tip visible at early BBCH 07 in early April after 850-950 chilling hours and 280-320 growing degree days base 10°C, with 1-2 mm pale green tissue faintly visible through 2-3 mm scale opening over 1-2 days under conditions of 13-15°C mean temperature and variable spring weather typical of continental climate, representing early-bursting pattern for this cultivar
First green tip visible represents the initial detection phase where green shoot tissue becomes barely perceptible through maximally separated bud scales, marking the earliest observable confirmation of dormancy exit and growth initiation. This subtle developmental stage occurs at the onset of BBCH 07 when expanding shoot primordium first breaches protective scale barrier, with green coloration faintly visible at bud apex where scale separation creates narrow viewing aperture. The visible green tissue measures 1-3 mm in observable length, appearing pale yellow-green due to limited chlorophyll accumulation in tissues recently emerged from enclosed environment within protective scales. At this initial visibility stage, green coloration may be intermittently visible depending on viewing angle and lighting conditions, requiring careful observation to detect particularly in overcast conditions or when viewing from non-optimal angles relative to scale opening. The shoot apex has reached 5-7 mm total length beneath scales with apical meristem actively producing cells through divisions every 20-24 hours, while subapical cells expand creating turgor pressure of 0.4-0.6 MPa sufficient to maintain position at scale opening but not yet generating force for rapid extension beyond scales. Bud scales maintain separation of 2-3 mm at apex creating viewing gap, with scales remaining structurally intact and providing continued protection to majority of developing shoot tissues beneath. Chlorophyll synthesis has recently initiated in shoot apex cells with concentrations reaching 0.1-0.2 mg per gram fresh weight, sufficient to create visible green coloration but representing early stage of chloroplast development. Photosynthetic capacity remains negligible at less than 10% of mature tissue rates, with respiration greatly exceeding any carbon fixation making shoots entirely dependent on stored carbohydrate reserves mobilized from woody tissues. Vascular connection between shoot and permanent structures has established functional xylem vessels conducting water to support turgor maintenance and beginning cellular metabolism, with water transport rate of 0.3-0.8 mg per second supporting current developmental stage. Cell division continues actively in apical meristem initiating new cells for shoot axis elongation and organ primordia, with cell production rate maintaining consistent 20-24 hour cycle characteristic of early emergence phase. Metabolic activity operates at elevated levels with respiration 12-15 fold above winter dormancy baseline, consuming stored sugars and mobilizing stored proteins to support rapid growth processes. Hormonal regulation maintains high cytokinin and gibberellin concentrations promoting continued meristem activity and cell expansion, with auxin synthesis in apex establishing developmental polarity. Temperature sensitivity remains high with development rate doubling per 10°C increase between 10-25°C, making progression highly dependent on spring temperature stability. Water availability must be adequate to support turgor maintenance in expanding tissues, with water stress potentially slowing visibility stage if uptake cannot meet demand. Light quality affects chlorophyll synthesis rate with red light promoting chloroplast development, though visibility stage occurs regardless of light conditions once developmental threshold is reached. The process typically requires 1-2 days from first faint visibility to progression toward extended green tip under favorable spring conditions above 14°C mean temperature, with cooler conditions extending duration to 2-4 days. Cultivar variation in first visibility appears in shoot apex size and chlorophyll synthesis rate at emergence, with some varieties showing brighter green at first detection while others remain pale yellow-green. Regional timing follows late bud swelling by 2-3 days, occurring mid-April in Mediterranean climates, late April in temperate areas, and early May in cool regions. Viticultural significance includes providing earliest possible observation point for confirming dormancy exit and growth initiation, enabling maximum lead time for frost protection deployment, serving as initial reference for beginning phenological monitoring programs, and establishing earliest marker for regulatory restrictions on vineyard operations in some regions. Frost damage at first green tip visible stage causes complete primary bud death through ice formation in tender emerging tissues, requiring compensatory growth from secondary buds with substantial delays and fruitfulness reduction. Field observation challenges include subtle visibility requiring careful inspection, variable detectability under different lighting conditions, and potential confusion with bud scale coloration changes that can mimic green tissue appearance. Research applications include precise determination of dormancy exit timing for phenological models, establishing earliest observation point for climate change monitoring studies, documenting cultivar variation in emergence characteristics, and evaluating prediction accuracy of thermal time models for green tip timing.
First_Green_Tip_Visible is the first of three Green_Tip_Emergence subphases, distinguished by subtle initial visibility of green tissue with minimal extension. Transitions to Green_Tip_Extended as shoot elongates and visibility becomes unambiguous. Represents earliest possible field observation point but requires careful inspection. Most challenging stage for consistent identification across observers.
barely visible green
erste Grünspitze sichtbar
faint green tip
first green visible
initial green tip
initiale Grünspitze
kaum sichtbare Grünfärbung
schwache Grünfärbung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 175-177 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
First Green Tip Visible
Chardonnay in Champagne shows first internode formation from late April to mid-May at BBCH 09-11 over 12-14 days, developing internode of 18-25 mm length between cotyledonary node and first true leaf, with elongation rate of 1.2-2.0 mm per day under cool maritime climate conditions of 13-17°C mean temperature, adequate spring moisture, and complete dependence on stored carbohydrate reserves typical of this early-bursting moderate-vigor cultivar
First internode formation represents the initial development of the stem segment between cotyledonary node and first true leaf node, establishing foundational shoot structure following bud burst and marking transition from embryonic to vegetative axis development. This developmental stage occurs during late April to May in temperate regions corresponding to BBCH 09-11 as first leaves unfold, with first internode initiating at shoot apex immediately behind apical meristem once first leaf primordium establishes nodal position defining proximal boundary. First internode formation proceeds through intercalary meristem establishment in axis tissue between cotyledonary position and first leaf attachment point, with meristematic cells undergoing divisions adding cells to developing internode segment over 7-14 days until cell production phase completes. Cell division in first internode intercalary meristem produces cells through organized divisions in longitudinal files, with division rates initially high at 20-24 hour cycles declining as meristem activity transitions from proliferation to expansion, typically producing 500-2000 cells along internode length depending on cultivar and final dimensions. The developing first internode shows procambial strand differentiation establishing vascular connection between permanent structures and developing shoot, with xylem vessels beginning to differentiate providing water transport capacity and phloem sieve elements forming enabling assimilate movement both upward from reserves and eventually downward from photosynthesizing leaves. Elongation of first internode begins while cell division continues, with cells in central zones initiating expansion through water uptake and wall loosening while marginal zones near nodes continue divisions, creating developmental gradient along internode length. First internode elongation typically produces final length of 15-35 mm substantially shorter than subsequent internodes, reflecting limited resource availability during early shoot development when photosynthetic capacity is minimal and growth depends entirely on stored reserves. The first internode shows characteristic anatomical structure including epidermis with developing cuticle providing protection, cortex with chloroplast-containing parenchyma contributing photosynthetic function, vascular bundles arranged in ring pattern characteristic of dicot stems, and central pith providing structural support and potential storage function. Mechanical properties of first internode develop progressively from soft herbaceous tissue during expansion to increasing rigidity as primary walls thicken and lignification initiates earlier than more apical internodes, providing stable base for developing shoot. Vascular connection through first internode is critical for shoot establishment, with xylem capacity of 0.5-1.5 mg per second adequate for supporting first leaf transpiration and growth water requirements, and phloem enabling initial upward movement of stored reserves and later bidirectional transport as photosynthesis initiates. Metabolic activity in developing first internode remains high with respiration 10-15 fold above mature tissue levels supporting growth processes, consuming stored carbohydrates transported through developing phloem until first leaf photosynthetic production begins supplementing energy supply. Cortical chloroplasts in first internode contribute modestly to carbon fixation once tissue is exposed to light, with stem photosynthesis potentially contributing 5-10% total shoot carbon gain though respiration in growing tissues typically exceeds local photosynthetic production. Hormonal regulation involves auxin from shoot apex promoting elongation and coordinating acropetal vascular differentiation pattern, gibberellins supporting cell expansion influencing final internode length, and cytokinins from roots promoting cell division in intercalary meristem. Temperature affects first internode development rate with elongation approximately doubling per 10°C increase between 12-25°C, influencing both cell production duration in meristem and subsequent expansion rate and duration. Light intensity and quality begin affecting development once internode emerges from bud scales, with high light promoting chloroplast development in cortex and potentially affecting elongation through photomorphogenic responses, while shade might promote slight elongation through etiolation response. Water availability must be adequate to support turgor maintaining cell expansion, with water stress during first internode development potentially permanently limiting length affecting subsequent shoot architecture through reduced basal internode stability. Nutrient availability particularly nitrogen affects development through effects on protein synthesis supporting cell division and expansion, with adequate nutrition supporting normal development while deficiency can reduce cell production and expansion limiting internode formation. The process typically requires 10-16 days from initiation to completion of elongation under favorable spring conditions, with cool temperatures or resource limitations extending duration. Cultivar variation appears in first internode length ranging from 12-20 mm in compact varieties to 25-40 mm in vigorous types, affecting basal shoot architecture and mechanical stability. Regional timing shows first internode formation occurring late April to early May in warm climates, May in temperate regions, and late May to early June in cool areas, following initial leaf unfolding and during continued bud burst progression. Viticultural significance includes establishing structural foundation for shoot development with adequate first internode length providing mechanical stability for shoot positioning operations, while insufficient development may create weak basal region susceptible to breakage during training. Damage to first internode during formation from mechanical injury, pests, or diseases can compromise shoot stability and vascular function affecting entire shoot development. Research applications include analyzing earliest stages of vegetative axis development documenting transition from embryonic to adult structure, evaluating cellular and molecular mechanisms of internode initiation and elongation, quantifying vascular differentiation patterns establishing shoot hydraulic architecture, and assessing environmental effects on early development affecting subsequent shoot growth patterns.
First_Internode_Formation is first of four Internode_Development subphases, distinguished by representing initial axis segment following bud burst. Shorter than subsequent internodes reflecting limited early resource availability. Transitions to Sequential_Internode_Formation as additional internodes develop. Critical for establishing stable shoot base. Timing coordinated with First_Leaf_Unfolding processes.
Grundinternodium
basal internode formation
basale Internodienbildung
erstes Internodium
first stem segment
initial internode development
primary internode
primäres Stängelglied
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 205-215 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 128-138 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
First Internode Formation
Tempranillo in Rioja shows first leaf expansion following BBCH 09 in early May after 1150-1300 chilling hours and 520-580 growing degree days base 10°C, with first leaf expanding from 18 mm to 35-45 mm over 4-5 days under conditions of 18-22°C mean temperature, 50-60% relative humidity, adequate soil moisture, and 14.5-hour photoperiod, representing typical continental climate pattern for this mid-season cultivar
First leaf expansion represents the progressive enlargement phase where first true leaf continues growing from initial separation configuration toward mature dimensions, characterized by rapid cell expansion and photosynthetic capacity development establishing functional leaf contribution to shoot carbon assimilation. This developmental stage follows initial separation with leaf blade expanding from 15-20 mm to 30-50 mm length over 3-5 days, developing characteristic palmate grapevine leaf morphology with clearly separated leaflets spreading from central rachis establishing recognizable cultivar-specific leaf form. Leaf expansion proceeds through coordinated processes including continued cell division in marginal and plate meristems producing new cells particularly at leaf margins, and rapid cell expansion throughout blade through turgor-driven cell wall extension increasing individual cell volumes 20-100 fold from meristematic dimensions. The expanding leaf displays intensifying green coloration as chlorophyll concentration increases to 1.5-2.5 mg per gram fresh weight approaching mature leaf levels, with upper leaf surface typically darker green than lower surface reflecting differential chloroplast distribution between palisade and spongy mesophyll tissues. Leaflets separate progressively from each other with sinuses deepening between lobes creating characteristic leaf outline, with final sinus depth varying substantially among cultivars from shallow creating nearly entire leaves to deep producing highly dissected configurations. Leaf surface texture develops characteristic features including trichome density varying from glabrous to densely tomentose depending on cultivar genetic programming, with trichome development affecting leaf optical properties including light reflectance and absorption. Stomata achieve full functional capacity with stomatal apertures responding dynamically to environmental conditions including light intensity providing photosynthetic substrate CO2, atmospheric vapor pressure deficit affecting transpirational water loss, and internal CO2 concentration reflecting photosynthetic activity. Chloroplast development approaches completion with thylakoid membranes fully organized into grana and stroma lamellae, photosystem complexes assembled and functional for light capture and electron transport, and Calvin cycle enzymes including Rubisco accumulated to concentrations supporting substantial carbon fixation. Photosynthetic capacity increases rapidly during expansion reaching 60-80% of mature rates when leaf achieves 70-80% final size, with light saturation point increasing and photosynthetic efficiency approaching mature tissue values. The expanding leaf transitions from net carbon consumer to approaching carbon neutrality as photosynthetic rate increases with area expansion and chloroplast maturation, with crossover to net production typically occurring when leaf reaches 80-90% final dimensions. Vascular system extends throughout expanding blade with primary veins providing structural framework and transport pathways, secondary veins forming reticulated network connecting primary veins, and tertiary veins developing fine network ensuring all photosynthetic cells have access to water and export pathway for photosynthate. Water content maintains 80-85% fresh weight typical of actively expanding tissues, requiring continuous water uptake through shoot xylem with transpirational water loss from expanding leaf area increasing to 15-40 mg per hour depending on environmental conditions and leaf size. Cell wall composition changes during expansion with initial walls rich in pectin allowing plastic deformation, transitioning toward walls with increased cellulose and hemicellulose content providing structural rigidity as expansion completes, determining final cell and leaf mechanical properties. Metabolic activity remains elevated with respiration 6-10 fold above mature leaf levels supporting growth processes and photosynthetic infrastructure development, gradually declining as expansion completes and maintenance requirements decrease. Protein synthesis continues at high rates producing structural proteins for expanding cells, photosynthetic proteins approaching mature leaf concentrations with Rubisco accumulating to 40-50% total leaf protein, and regulatory proteins coordinating development and establishing metabolic capacity. Hormonal regulation maintains cytokinin and gibberellin supporting continued expansion, with auxin concentration declining from separation levels as expansion proceeds and polarity patterns become established. Temperature strongly affects expansion rate with optimal range 20-25°C for maximum rate, temperatures above 28-30°C reducing expansion through increased respiration and water stress while temperatures below 15°C slowing all processes substantially. Water availability critically affects expansion rate and final size with adequate soil moisture essential for maintaining turgor driving cell expansion, while water stress arrests expansion causing permanent size reduction even after recovery affecting photosynthetic capacity. Light intensity influences expansion rate and photosynthetic capacity development with high light promoting rapid chloroplast maturation to high capacity while shade reduces capacity though may increase specific leaf area through thinner blades. Nutrient availability particularly nitrogen affects expansion rate and final photosynthetic capacity, with adequate nitrogen supporting protein synthesis for both structural and photosynthetic components while deficiency reduces expansion and limits photosynthetic potential. The process typically requires 3-5 days under favorable conditions with consistent temperatures 18-22°C and adequate resources, with suboptimal conditions extending duration to 5-8 days or causing incomplete expansion to reduced final dimensions. Cultivar variation appears in expansion rate, final leaf dimensions ranging from small 50-80 mm leaves to large 150-200 mm leaves, leaflet number and sinus depth creating diverse leaf shapes, and trichome density affecting leaf appearance and microclimate. Regional timing follows first leaf separation by 2-3 days, occurring early May in warm climates, mid-May in temperate regions, and late May in cool areas. Viticultural management considerations include ensuring adequate water availability particularly during rapid expansion phase when stress severely impacts final size, providing sufficient nitrogen supporting protein synthesis for photosynthetic capacity, monitoring for pest colonization including leafhoppers and spider mites preferring young expanding leaves, and assessing expansion uniformity indicating shoot vigor and resource availability affecting canopy development. Damage during expansion from pests, diseases, or environmental stress permanently reduces photosynthetic capacity affecting shoot growth potential and subsequent fruit development. Research applications include analyzing cellular mechanisms of coordinated division and expansion, quantifying photosynthetic capacity development relative to leaf age and size, evaluating environmental factor effects on expansion rates and final characteristics including size and photosynthetic capacity, documenting hormonal and genetic regulation of leaf form development, and assessing implications of early leaf development for subsequent shoot growth and crop potential.
First_Leaf_Expansion is the second of three First_Leaf_Unfolding subphases, distinguished by rapid enlargement of separated first leaf approaching functional dimensions. Transitions to Multiple_Leaf_Unfolding as additional leaves begin separating and expanding. Represents critical phase establishing shoot photosynthetic capacity. Final leaf size at this stage strongly affects subsequent shoot growth potential.
Blattwachstum
enlarging leaf
erste Blattexpansion
expandierendes Blatt
expanding first leaf
growing leaf
leaf expansion phase
wachsendes Erstblatt
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 195-205 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 95-105 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
First Leaf Expansion
Chardonnay in Champagne shows first leaf separation at BBCH 09 in late April after 900-1000 chilling hours and 420-480 growing degree days base 10°C, with first leaf petiole angling 30-45 degrees from axis and blade spreading to 12-16 mm over 2-3 days under conditions of 15-18°C mean temperature, 65-75% relative humidity, and 14-hour photoperiod, representing typical cool maritime climate progression for this early-bursting cultivar
First leaf separation represents the initial unfolding phase where first true leaf begins spreading away from compressed position along shoot axis, marking the beginning of functional leaf development and photosynthetic capacity establishment. This developmental stage occurs at BBCH 09 when shoot elongation has progressed sufficiently that mechanical constraints maintaining leaf compression are released, allowing leaf petiole to angle away from axis and leaf blade to begin expanding perpendicular to shoot. The separating first leaf measures 10-15 mm length at separation initiation, having developed from 5-8 mm compressed primordium through continued cell production in marginal meristems and beginning cell expansion in blade tissues. Leaf blade shows beginning spread of palmate leaflet configuration with leaflets starting to separate from each other though remaining partially folded, creating transitional form between compressed primordium and fully expanded leaf architecture. The leaf petiole angles 20-40 degrees from shoot axis at initial separation, progressively increasing to 60-90 degrees as separation advances and cell expansion in petiole abaxial side creates upward bending establishing characteristic leaf orientation. Leaf coloration transitions from pale yellow-green of recently uncompressed tissue to brighter green as chlorophyll synthesis accelerates with light exposure, with chlorophyll concentration increasing from 0.4-0.6 mg per gram to 0.8-1.2 mg per gram fresh weight during separation phase. Chloroplast development progresses rapidly in expanding blade cells with thylakoid membranes organizing into grana stacks, photosystem complexes assembling for light capture, and Calvin cycle enzymes accumulating for carbon fixation capability. Photosynthetic capacity begins developing as chloroplasts mature though rates remain below 30% of mature leaf capacity during initial separation, with light saturation point low and photosynthetic efficiency suboptimal compared to fully developed leaves. Stomata complete development and become functional during separation, with stomatal apertures responding to environmental conditions including light intensity, CO2 concentration, and leaf water status to regulate gas exchange for photosynthesis and water loss through transpiration. Leaf surface features including trichomes and epicuticular waxes develop during separation creating characteristic cultivar-specific leaf appearance and affecting optical properties including reflectance and light penetration to photosynthetic tissues. Cell division continues in marginal and plate meristems at leaf edges and throughout blade respectively, producing new cells that will expand to create final leaf dimensions, with cell production rates highest during early separation phase. Cell expansion begins throughout leaf blade through water uptake creating turgor pressure that drives cell wall extension, with expansion rate initially limited by wall properties and increasing as wall-loosening enzymes accumulate and allow plastic deformation. Vascular system extends throughout separating leaf establishing primary veins forming structural framework and beginning development of secondary vein network for resource distribution, with xylem vessels differentiating to transport water and phloem sieve elements forming to export beginning photosynthetic products. Water content in separating leaf increases to 75-85% fresh weight typical of actively growing tissues, requiring continuous water uptake through shoot xylem to maintain turgor supporting expansion and prevent wilting. The shoot continues elongating at 3-5 mm per day with apical meristem producing additional leaf primordia while first leaf separates, creating sequential pattern of leaf development along shoot axis. Metabolic activity remains elevated in separating leaf with respiration 8-12 fold above mature leaf levels supporting rapid growth processes, consuming both stored carbohydrates transported from woody tissues and beginning to use photosynthate produced by developing photosynthetic capacity. Protein synthesis operates at high rates producing structural proteins for cell walls and membranes of expanding cells, photosynthetic proteins including Rubisco and other Calvin cycle enzymes, and regulatory proteins coordinating development. Hormonal regulation involves cytokinin promoting cell division in meristematic zones, gibberellin supporting cell expansion throughout blade, and auxin establishing polarity patterns affecting leaf shape and orientation relative to axis. Temperature affects separation rate and subsequent expansion with optimal range 18-24°C for maximum development rate, higher temperatures potentially reducing efficiency through increased respiration and water stress while lower temperatures slow all processes. Water availability critically affects separation and expansion with adequate soil moisture essential to maintain turgor driving cell expansion, while water stress arrests development and can cause permanent size reduction even after stress relief. Light intensity influences separation rate and subsequent photosynthetic development with high light promoting rapid chloroplast maturation and full photosynthetic capacity while shade reduces development rate and ultimate capacity. Nutrient availability particularly nitrogen affects protein synthesis capacity supporting both structural and photosynthetic protein production required for full leaf development. The process typically requires 1-3 days from initial separation to obvious spreading under favorable conditions above 18°C, with cooler temperatures extending duration to 3-5 days. Cultivar variation appears in separation timing relative to shoot elongation and in separation angle and rate, with some varieties showing rapid perpendicular spreading while others separate more gradually to acute angles. Regional timing follows green shoot tip visible by 2-4 days, occurring late April to early May in warm climates, early to mid-May in temperate areas, and mid-to-late May in cool regions. Viticultural significance includes marking beginning of rapid leaf area development requiring adequate water and nutrient availability, establishing point where photosynthetic capacity begins contributing to carbon balance affecting subsequent growth potential, providing developmental reference for monitoring shoot growth uniformity across vineyard blocks, and serving as marker for intensifying pest monitoring as leaf area increases colonization opportunity. Damage during separation from mechanical injury, pests, or diseases reduces photosynthetic capacity affecting shoot growth potential and fruit development. Research applications include analyzing cellular and molecular mechanisms of leaf separation and unfolding, quantifying photosynthetic capacity development during early leaf expansion, evaluating environmental effects on leaf development rates and final characteristics, and assessing hormonal regulation coordinating leaf positioning and expansion processes.
First_Leaf_Separation is the first of three First_Leaf_Unfolding subphases, distinguished by initial spreading of first leaf from compressed position along axis. Transitions to First_Leaf_Expansion as blade continues enlarging and spreading. Represents beginning of significant photosynthetic contribution to shoot carbon balance. Standard BBCH 09 observation point.
Blattablösung vom Trieb
beginnende Blattentfaltung
beginning leaf unfolding
erste Blattspreizung
first leaf spreading
initial leaf separation
initiale Blattseparation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 189-195 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 89-95 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
First Leaf Separation
Merlot in Bordeaux shows first leaf unfolding at BBCH 09-11 in early May after 1000-1100 chilling hours and 480-550 growing degree days base 10°C, with first true leaves separating from axis and expanding to 18-25 mm over 6-8 days under conditions of 17-20°C mean temperature, 60-70% relative humidity, adequate soil moisture, and 14.5-hour photoperiod, representing typical oceanic climate pattern for this mid-season cultivar
First leaf unfolding represents the critical developmental transition where initial true leaves separate from shoot axis and expand toward functional photosynthetic configuration, marking the shift from bud burst completion to active vegetative growth phase. This fundamental process corresponds to BBCH 09-11, occurring when shoot has elongated sufficiently that first leaves no longer remain compressed against axis but instead spread perpendicular establishing characteristic grapevine leaf positioning and beginning substantial contribution to carbon assimilation. The process encompasses progression from initial leaf separation through partial expansion to establishment of multiple unfolding leaves, typically requiring 5-10 days depending on environmental conditions and cultivar growth characteristics. First leaf unfolding marks definitive transition from heterotrophic growth depending entirely on stored reserves to beginning autotrophic capacity as expanding leaves develop photosynthetic function, though shoots remain net carbon consumers until 3-5 leaves fully expand. The unfolding leaves expand from compressed primordia configuration measuring 5-10 mm to partially expanded leaves of 15-30 mm length, with characteristic palmate grapevine leaf form becoming clearly recognizable as leaflets separate and spread from central rachis. Leaf surfaces transition from pale yellow-green of enclosed tissues to bright green as chlorophyll accumulates and chloroplasts mature, with upper leaf surface typically developing darker green than lower surface reflecting chloroplast distribution differences. Leaf expansion proceeds through coordinated cell division in marginal and plate meristem zones producing new cells, and cell expansion throughout leaf blade through turgor-driven growth increasing individual cell volumes 10-50 fold. Stomata develop and become functional enabling gas exchange for photosynthesis and transpirational cooling, with stomatal density on lower leaf surface reaching 150-250 per square mm compared to 50-100 per square mm on upper surface. Trichomes characteristic of grapevine leaves emerge from epidermal cells creating varying degrees of leaf hairiness depending on cultivar, with density ranging from sparse to dense coverage affecting leaf reflectance and transpiration rates. Vascular system extends throughout expanding leaf blade establishing network for water delivery and photosynthate export, with primary veins forming structural framework and secondary and tertiary veins creating reticulated pattern for resource distribution. Chloroplast development accelerates in leaf cells with chlorophyll concentration increasing to 1.2-2.0 mg per gram fresh weight, thylakoid membranes organizing into grana stacks for light capture, and Calvin cycle enzymes assembling for carbon fixation. Photosynthetic capacity increases progressively as leaves expand, reaching 50-70% of mature rates when leaves achieve 40-50% final size, with full photosynthetic capacity developing as expansion completes and chloroplasts mature. The metabolic balance transitions from net carbon consumption to approaching neutral and eventually positive as photosynthetic rate increases with leaf area expansion, with crossover point typically occurring when 3-4 leaves have substantially expanded. Shoot elongation continues with apical meristem producing new leaf primordia every 2-3 days while older leaves unfold and expand sequentially, creating characteristic pattern of progressively larger leaves from shoot base toward apex. Stored carbohydrate reserves continue supporting growth during unfolding phase with cane starch concentrations declining to 30-40% of winter maximum, while new photosynthate increasingly supplements stored reserves as leaf area develops. Protein synthesis continues at elevated rates supporting leaf tissue development, with particular emphasis on photosynthetic proteins including Rubisco comprising up to 50% of total leaf protein in mature tissues. Hormonal regulation maintains elevated cytokinin and gibberellin promoting continued leaf expansion, with auxin from shoot apex coordinating leaf positioning and establishing patterns for additional organ differentiation including tendrils and inflorescence primordia. Temperature continues affecting development rate with leaf expansion approximately doubling per 10°C increase between 10-25°C, though optimal temperature for maximum expansion rate is 20-25°C with higher temperatures potentially reducing expansion through increased respiration and water stress. Water availability becomes critically important as transpiring leaf area increases exponentially, with adequate soil moisture and root function essential to maintain turgor supporting expansion and preventing water stress that arrests development. Light intensity strongly affects leaf expansion rate and final size with high light promoting rapid expansion and larger final dimensions while shade reduces expansion rate and ultimate leaf size, with adaptation beginning whereby shaded leaves develop thinner blades with higher specific leaf area. Nutrient availability particularly nitrogen affects expansion rate and photosynthetic capacity development, with adequate nitrogen supporting protein synthesis for both structural and photosynthetic components. The process shows three distinguishable phases based on leaf separation and expansion progression, advancing from initial first leaf separation through first leaf expansion to multiple leaves unfolding creating developing shoot canopy. Cultivar variation appears in leaf unfolding rate, final leaf size, and timing of photosynthetic capacity development, with vigorous cultivars showing rapid expansion to large final dimensions while less vigorous types develop more slowly to smaller leaf sizes. Regional timing follows complete bud breaking by 2-4 days, occurring late April to early May in warm climates, early to mid-May in temperate regions, and mid-to-late May in cool areas. Viticultural management considerations include transitioning from frost protection focus to shoot positioning and canopy management, implementing nutrient management particularly nitrogen to support rapid leaf development, establishing irrigation scheduling as water demand increases dramatically, monitoring for pest colonization of expanding leaf area including mites and leafhoppers, and assessing shoot growth uniformity for crop load and canopy management decisions. Frost damage during leaf unfolding kills expanded leaf tissue though growing point may survive if sufficiently protected, causing growth delays and reduced photosynthetic capacity affecting subsequent development. Research applications include quantifying transition from heterotrophic to autotrophic carbon balance, analyzing photosynthetic capacity development in expanding leaves, evaluating environmental effects on leaf expansion rates and final sizes, documenting hormonal regulation of leaf development, and assessing cultivar differences in early canopy development affecting light interception and photosynthetic capacity.
First_Leaf_Unfolding follows Complete_Bud_Breaking and represents transition from bud burst to active shoot growth. Distinguished by leaf separation from axis and progressive expansion. Subdivided into First_Leaf_Separation, First_Leaf_Expansion, and Multiple_Leaf_Unfolding based on progression. Marks beginning of significant photosynthetic contribution and transition toward autotrophic growth. Critical phase for establishing shoot photosynthetic capacity.
Blattexpansion
Blattöffnung
erste Blattentfaltung
leaf expansion initiation
opening leaves
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 189-205 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 89-105 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
First Leaf Unfolding
Merlot grafted on 3309 Couderc rootstock at 14 years showing first-order lateral secondary roots branching directly from primary roots, largest first-order lateral 5.8 cm diameter at origin from primary root extending 142 cm horizontally at average depth 52 cm, first-order laterals emerging approximately 18 cm intervals along primary root length, first-order lateral showing 14 annual xylem rings indicating formation in establishment year, second-order branches arising from first-order lateral at 8-12 cm intervals creating hierarchical network, first-order lateral serving as structural framework and transport conduit from second and higher-order absorptive roots to primary root and trunk
First order lateral secondary root is lateral secondary root branching directly from primary root representing first hierarchical level in root system branching architecture, characterized by largest diameter among lateral roots typically 2-8 cm at origin from primary root tapering along length, earliest branching position in hierarchy being first generation of lateral branches, moderate to substantial secondary growth with secondary xylem development and bark formation, and serving dual functions as structural framework supporting higher-order branches and transport conduit moving water and nutrients from absorptive network to primary root, with first-order laterals typically emerging 10-30 cm intervals along primary root length creating regular branching pattern, showing predominantly horizontal or oblique orientation exploring soil laterally, producing second-order branches that generate third and higher-order roots creating complete hierarchical network, and representing critical architectural element connecting deep primary root axis to extensive fine absorptive root system in upper soil layers. First order lateral secondary root hierarchical position includes= branching directly from primary root (first-order lateral arising from main primary root or tap root as first branching level), representing lowest lateral order (first-order being first generation of lateral branches in root order classification), serving as parent to higher orders (first-order laterals producing second-order branches which produce third-order and so on), showing order-1 designation (in root order numbering system where finest roots are order-1 and orders increase toward primary root, or alternatively first-order laterals are order-1 in decreasing system), and demonstrating foundational role (first-order laterals providing structural foundation from which entire lateral root network develops). First order lateral secondary root diameter and size includes= showing large diameter (first-order laterals typically 2-8 cm diameter at origin from primary root largest among secondary roots), exhibiting substantial size (first-order lateral length potentially 50-200 cm with significant biomass), demonstrating gradual taper (diameter decreasing progressively along length from origin toward tip), displaying secondary growth (first-order laterals developing secondary xylem rings and protective bark like smaller primary roots), and revealing age correlation (older first-order laterals showing greater diameter from years of secondary thickening compared to young laterals). First order lateral secondary root origin pattern includes= emerging at intervals along primary root (first-order laterals typically arising 10-30 cm apart along primary root length depending on vigor and conditions), showing acropetal development (first-order laterals more abundant near primary root growing tip declining toward base), arising from primary root pericycle (lateral origin from internal pericycle tissue differentiating into root primordium), demonstrating variable number (primary roots typically producing 5-15+ first-order laterals depending on length and environmental conditions), and exhibiting branching angle (first-order laterals emerging 45-90° from primary root axis typically tending toward horizontal). First order lateral secondary root orientation and extent includes= growing predominantly horizontally (first-order laterals showing horizontal or oblique trajectory typically 0-45° from horizontal), extending laterally from primary root (first-order lateral exploration primarily sideways rather than vertical creating lateral spread), reaching substantial distance (individual first-order laterals potentially extending 80-150 cm from primary root origin), occupying upper to middle soil depths (first-order laterals typically 30-90 cm depth when emerging from upper primary root or deeper when from deep primary root), and creating radial pattern (first-order laterals from vertical primary root growing in multiple radial directions creating disk-like soil exploration). First order lateral secondary root secondary branching includes= producing second-order branches (first-order laterals generating numerous second-order lateral roots creating next hierarchical level), showing branch spacing (second-order roots typically emerging 5-15 cm intervals along first-order lateral), exhibiting branching density (branching potentially higher near first-order lateral tip declining toward base), demonstrating progressive branching (first-order lateral continuing to produce new second-order branches as elongates), and creating hierarchical network (first-order lateral with second and higher-order branches forming complete absorptive network exploring local soil volume). First order lateral secondary root structural function includes= providing mechanical framework (first-order laterals creating structural skeleton from which finer absorptive roots branch), contributing to anchorage (first-order laterals helping stabilize vine through horizontal spread and branching network), bearing mechanical loads (first-order laterals supporting weight and tension from higher-order branches and absorptive roots), enabling root system architecture (first-order lateral distribution and orientation determining overall root system spatial organization), and demonstrating persistence (first-order laterals typically persistent structures functioning for many years unlike ephemeral fine roots). First order lateral secondary root transport function includes= conducting water from branches (first-order lateral xylem moving water absorbed by second and higher-order roots toward primary root), moving nutrients to primary root (first-order laterals transporting minerals from absorptive network basipetally), showing high hydraulic conductivity (large-diameter xylem vessels in first-order laterals providing efficient transport pathway), enabling bidirectional phloem transport (first-order lateral phloem conducting photosynthates to finer roots and storing excess in parenchyma), and demonstrating substantial capacity (first-order lateral potentially conducting 10-30+ liters daily from branch network depending on lateral size and absorption intensity). First order lateral secondary root storage function includes= accumulating carbohydrate reserves (first-order lateral parenchyma and ray cells storing starch particularly during dormancy), showing seasonal storage cycles (reserves highest fall-winter declining spring as mobilized for growth), providing substantial storage capacity (first-order lateral moderate to large size enabling significant storage contribution to total root reserves), supporting spring growth (first-order lateral reserves mobilized for renewed root and shoot growth before leaves functional), and enabling stress recovery (first-order lateral carbohydrates supporting vine recovery from damage or environmental stress).
First_Order_Lateral_Secondary_Root emphasizes direct branching from primary root and hierarchical position and is distinguished from Second_Order_Lateral_Secondary_Root (branching from first-order) by focus on first branching level. Critical for understanding root order hierarchy, structural framework, primary-to-lateral connection, branching architecture, and the first-level lateral roots providing foundational framework from which entire secondary root network develops.
Direct lateral from primary
Erste Verzweigungsebene
First branch lateral
Lateralwurzel erster Ordnung
Primary lateral root
Primäre Seitenwurzel
First Order Lateral Secondary Root
Merlot on 3309C showing first-order lateral shallow roots, excavation revealing laterals 4.2-6.8 cm diameter branching from primary root at 22-34 cm depth, first-order laterals extending 120-165 cm horizontal distance, predominantly horizontal orientation 72-88° from vertical, microscopic sections showing 8-11 annual rings indicating 8-11 years age, first-order laterals producing second-order branches every 15-25 cm, first-order lateral shallow roots providing primary lateral structural framework supporting higher-order absorptive network
First order lateral shallow root is lateral shallow root representing first branching level originating directly from primary taproot or main cutting-derived root, typically 2-8 cm diameter at origin tapering distally, characterized by direct connection to central root axis, substantial structural development with moderate to extensive secondary growth, important transport function conveying resources between central axis and higher-order lateral branches, and serving as primary lateral framework supporting second-order and higher lateral branches that create complete hierarchical network, with first-order lateral shallow roots in grapevine showing typical characteristics including origin from primary root at 10-40 cm depth, horizontal to oblique orientation 45-90° from vertical, lengths 50-200+ cm extending substantial distance from trunk, moderate to extensive secondary growth developing woody tissue with 5-15+ annual rings in older roots, production of numerous second-order lateral branches every 10-30 cm along length creating hierarchical structure, and dual functions as structural framework providing mechanical support and anchorage plus transport conduit connecting higher-order absorptive network to main root axis and vine vascular system. First order lateral shallow root origin and initiation includes= showing primary root connection (first-order laterals branching directly from primary taproot or cutting-derived primary roots), exhibiting endogenous origin (laterals initiating from pericycle tissue inside primary root emerging through cortex), demonstrating depth range (first-order laterals typically originating 10-40 cm depth though variable with soil and rootstock), revealing spacing patterns (first-order laterals emerging at irregular intervals typically 10-30 cm apart along primary root), and displaying developmental timing (first-order laterals forming progressively as primary root elongates creating age gradient). First order lateral shallow root diameter and structure includes= showing substantial diameter (first-order laterals 2-8 cm diameter at origin tapering to 1-4 cm distally depending on age and vigor), exhibiting secondary growth (first-order laterals undergoing extensive secondary thickening developing woody tissue), demonstrating annual rings (woody first-order laterals showing 5-15+ annual rings in older roots indicating years to decades of growth), revealing bark development (older first-order laterals developing protective bark as secondary growth proceeds), and displaying structural robustness (first-order laterals showing rigid woody structure providing mechanical strength). First order lateral shallow root length and extent includes= showing substantial reach (first-order laterals extending 50-200+ cm from trunk accessing large soil volume), exhibiting variable length (individual first-order lateral length varying widely depending on direction, soil conditions, competition), demonstrating radial distribution (first-order laterals extending in multiple directions creating roughly radial pattern), revealing asymmetries (first-order lateral extent potentially greater in favorable directions or away from competing vines), and displaying exploration (first-order laterals providing primary mechanism for lateral soil exploration in shallow zone). First order lateral shallow root branching pattern includes= showing second-order production (first-order laterals producing numerous second-order lateral branches along their length), exhibiting regular spacing (second-order branches emerging at roughly regular intervals 10-30 cm though variable), demonstrating hierarchy creation (first-order lateral branching creating second level in root system hierarchical architecture), revealing progressive branching (branching density potentially increasing distally as first-order lateral ages and develops), and displaying network formation (first-order lateral branching creating complete network exploring soil volume). First order lateral shallow root transport function includes= providing vascular connection (first-order laterals connecting higher-order branches to main root axis and vine), exhibiting xylem transport (first-order lateral xylem conducting water and nutrients from absorptive roots toward shoot), demonstrating phloem transport (first-order lateral phloem transporting photosynthates from shoot toward fine roots), revealing bidirectional flow (simultaneous xylem ascent and phloem descent in first-order laterals), and displaying transport capacity (substantial vascular tissue in first-order laterals providing high transport conductance). First order lateral shallow root structural function includes= providing framework (first-order laterals creating primary lateral structural framework in shallow soil), exhibiting mechanical support (first-order laterals bearing weight and forces from extensive higher-order branch network), demonstrating anchorage contribution (first-order laterals contributing to vine anchorage particularly resisting lateral forces), revealing load distribution (first-order laterals distributing mechanical stresses from fine root network across multiple branches), and displaying persistence (first-order lateral structural function maintained years to decades). First order lateral shallow root absorptive support includes= supporting higher-order network (first-order laterals serving as parent axes for extensive second-order and higher branching), enabling fine root production (second-order and higher branches from first-order laterals ultimately supporting thousands of fine absorptive roots), demonstrating indirect absorption (first-order laterals providing absorption primarily through higher-order derivatives rather than direct uptake), revealing resource delivery (first-order laterals transporting absorbed resources from higher-order network toward vine), and displaying critical role (first-order lateral extent and branching determining total root system absorption capacity). First order lateral shallow root lifespan and turnover includes= showing long persistence (first-order laterals potentially functioning years to decades once established), exhibiting low turnover (first-order lateral mortality and replacement rates very low compared to fine roots), demonstrating cumulative growth (first-order laterals showing continued diameter growth and secondary development over years), revealing investment (first-order laterals representing substantial long-term carbon investment), and displaying permanence (first-order laterals forming relatively permanent structural framework).
First_Order_Lateral_Shallow_Root emphasizes first branching level from primary root and is distinguished from higher orders by focus on direct primary root connection. Includes Primary_First_Order_Lateral_Shallow_Root and Proximal_First_Order_Lateral_Shallow_Root. Critical for understanding primary lateral framework, substantial structure, transport function, second-order support, and the first-level lateral branches creating foundation for hierarchical lateral root network.
Erste Ordnung laterale flache Wurzel
First-level horizontal root
Haupt-Seitenast
Main lateral branch
Primäre Lateralwurzel
First Order Lateral Shallow Root
A first-order primary branch rachis representing the seventh primary branch on a primary inflorescence of Vitis vinifera cv. Cabernet Sauvignon in Bordeaux, France, emphasizing its hierarchical position as first-order lateral branching from the central main rachis, originating directly from the main rachis at 10.2 cm from inflorescence base without any intermediate branching structures, representing second-order position in the complete hierarchy (following first-order main rachis), measuring 4.8 cm length, diameter of 2.2 mm at main rachis junction, containing 4 vascular bundles representing first-order lateral vascular capacity, serving as the source for 6 secondary branch rachides (third-order structures, second-order laterals) which subsequently elaborate into tertiary branches and flowers, first-order lateral position creating the initial three-dimensional expansion with this branch extending 4.8 cm perpendicular to the main rachis axis significantly increasing inflorescence volume beyond the linear main rachis alone, bearing approximately 38 flowers distributed across its secondary and tertiary branch system, measured at BBCH stage 65 (full bloom) in early June, representing typical first-order primary branch rachis in its role as initial lateral elaboration from main rachis.
A primary branch rachis emphasizing the first-order lateral position in the branching hierarchy in Vitis vinifera, characterized by hierarchical position as the first lateral branching order from the central main rachis, direct origin from the main rachis without intermediate structures, second-order position in the complete rachis hierarchy (first-order being main rachis), role as the source from which all higher-order lateral branches derive through sequential branching, dimensions and organization reflecting the first lateral order typically 3-7 cm length with 3-6 vascular bundles, and functional significance as the primary lateral elaboration creating the initial three-dimensional expansion from the linear main rachis. First-order primary branch rachides emphasize the hierarchical position as the initial lateral branching level, representing the first departure from the simple linear main rachis architecture, developing through the first lateral meristem activity establishing lateral axes, and serving as the foundation for higher-order branch elaboration creating the complete paniculate structure.
First-order primary branch rachides emphasize the hierarchical position as the initial lateral branching level, distinguished from simple primary branch designation by explicit focus on first-order lateral status. The first-order designation clarifies the branching hierarchy with main rachis as zero-order or first-order depending on numbering scheme, these primary branches as first-order laterals (or second-order absolute), and secondary branches as second-order laterals (third-order absolute). Direct origin from main rachis without intermediates defines the first-order lateral status, contrasting with higher-order branches originating from other lateral branches. Architectural role involves creating the initial three-dimensional expansion from the linear main rachis, with first-order branches extending perpendicular or oblique to the main axis significantly increasing inflorescence volume and distributing flowers spatially. Source function for higher orders means first-order primary branches serve as the foundation for second-order secondary branches and third-order tertiary branches, creating the hierarchical elaboration. Developmental sequence shows first-order primary branch primordia as the initial lateral structures initiating along the developing main rachis, preceding all higher-order elaboration. Vascular organization shows first-order branches receiving vascular traces directly from main rachis bundles, subsequently distributing to higher-order branches through progressive branching. Functional capacity reflects the first-order position with dimensions and vascular capacity intermediate between main rachis and higher-order branches. Transformation to cluster maintains first-order lateral identity with these structures becoming the major cluster arms bearing the secondary and tertiary berry-bearing branches.
direct main rachis branch
direkte Hauptrachis-Verzweigung
erste Ordnung primäre Zweigrachis
initial lateral branch rachis
initiale laterale Zweigachse
second-order hierarchical rachis
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 327-362 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 519-558.
First Order Primary Branch Rachis
In a Barossa Valley Shiraz vineyard, first pigment deposition was observed by microscopy in late January Year N+1 (BBCH 83) showing red-purple vacuolar pigmentation in epidermal cells at anthocyanin concentration 48 mg/kg skin when berries appeared still green to unaided eye, with malvidin-3-glucoside comprising 68% of deposited anthocyanins, delphinidin-3-glucoside at 14%, vacuolar pH measured 3.4, and flavonol co-pigments at 85 mg/kg under warm conditions (berry temperature 26-29°C) providing optimal deposition environment.
First_Pigment_Deposition represents the initial accumulation of synthesized anthocyanins in berry skin cell vacuoles at concentrations sufficient to create microscopically detectable pigmentation, occurring during the earliest phase of color development 1-2 days following biosynthetic pathway activation and immediately preceding macroscopically visible color change. This process involves vacuolar sequestration of anthocyanin-3-glucosides at 30-80 mg/kg skin fresh weight (compared to 32°C inhibits deposition through reduced synthesis and potential membrane disruption affecting vacuolar transport efficiency. Light influences first pigment deposition beyond biosynthetic effects: UV-B radiation may directly affect vacuolar transport mechanisms enhancing anthocyanin sequestration efficiency, visible light particularly blue wavelengths could modulate tonoplast transporter activity, and light-induced changes in vacuolar pH through effects on proton pump activities might optimize conditions for anthocyanin stabilization in flavylium form. Metabolic requirements for first pigment deposition inclu ATP consumption for active transport across tonoplast estimated 2-4 molecules ATP per anthocyanin molecule sequestered representing significant energy demand during rapid deposition, proton gradient maintenance through V-ATPase activity consuming additional ATP to sustain low vacuolar pH essential for color stability, and potential synthesis of vacuolar inclusion membrane components in cultivars forming AVIs requiring lipid and protein synthesis. Cultivar variation in first pigment deposition patterns includes: rapid-depositing varieties like Pinot Noir show visible pigment within 24-36 hours of synthesis initiation with abrupt color appearance, moderate cultivars like Merlot require 36-60 hours for detectable deposition with more gradual onset, while slow-depositing varieties like Cabernet Sauvignon need 60-96 hours reflecting either slower synthesis or less efficient transport and sequestration mechanisms. Coordination with berry softening varies: in most cultivars first pigment deposition occurs 0-2 days before or concurrent with initial berry softening onset, some early-coloring varieties show pigment deposition 2-3 days preceding softening, while certain cultivars may soften 1-2 days before visible pigment suggesting relatively independent regulation though ultimately coordinated through common ABA signaling. Physiological consequences of pigment deposition inclu anthocyanin-containing vacuoles gain antioxidant capacity protecting cellular contents from reactive oxygen species particularly as photosynthetic apparatus degrades, UV-screening properties reduce DNA damage and photoinhibition in pigmented cells, and potential osmotic effects of high vacuolar anthocyanin concentrations (5-15 mM) might influence water relations though likely minor compared to sugar and acid effects. Quality implications inclu early and intense pigment deposition correlates with eventual high color intensity at harvest important for red wine quality, spatial uniformity of first deposition within berries and clusters predicts maturity synchrony affecting harvest decisions and wine composition consistency, and anthocyanin composition established during first deposition largely determines final wine color characteristics including hue angle, color stability, and aging behavior.
First_Pigment_Deposition is distinguished from Initial_Anthocyanin_Synthesis by representing vacuolar sequestration rather than biosynthesis, from First_Color_Change by occurring at microscopic rather than macroscopic detection scale, and from Progressive_Deposition by showing initial rather than expanding accumulation. Anthocyanin threshold for microscopic detection: 30-80 mg/kg skin. Vacuolar pH 3.0-3.8 critical for color stability. Epidermal and sub-epidermal layers primary deposition sites. Temperature optimum 22-28°C for transport efficiency. First_Pigment_Deposition represents the critical transition from biosynthesis to visible color creation.
early anthocyanin deposition
erste Pigment-Sequestrierung
first pigment sequestration
frühe Anthocyan-Ablagerung
initial vacuolar anthocyanin accumulation
initiale vakuoläre Anthocyan-Akkumulation
primary color establishment
primäre Farb-Etablierung
Conn S, Curtin C, Bezier A, Franco C, Zhang W (2008) Purification, molecular cloning, and characterization of glutathione S-transferases involved in anthocyanin sequestration J Biol Chem 283:26484-26491 anthocyanin transport mechanisms Conn S, Zhang W, Franco C (2003) Anthocyanic vacuolar inclusions in cell suspension cultures of Vitis vinifera Plant Cell Rep 22:227-229 vacuolar sequestration Braidot E, Zancani M, Petrussa E, Peresson C, Bertolini A, Patui S, Macrì F, Vianello A (2008) Transport and accumulation of flavonoids in grapevine Plant Signal Behav 3:626-632 pigment deposition mechanisms.
First Pigment Deposition
Merlot grafted on 101-14 rootstock in first growing season (5 months post-grafting, late September) showing first year grafted rootstock trunk 1.4 cm diameter, trunk extending 18 cm from root crown to graft union positioned 15 cm above soil, graft union displaying prominent 1.8 cm diameter callus bulge with active white callus tissue visible, trunk bark absent (thin greenish-tan epidermis covering trunk surface with visible lenticels), trunk requiring stake support, diameter growth 0.4 cm since grafting, no disease symptoms, successful scion shoot development to 45 cm height indicating functional graft union
First year grafted rootstock trunk is young grafted rootstock trunk in initial growing season following grafting, representing the most critical early developmental stage characterized by initial graft union callus formation, establishment of first vascular connections across graft interface, very small diameter typically 0.8-2.5 cm, smooth ungrafted appearance with minimal or absent periderm, extreme vulnerability requiring intensive protection and support, and marking the transition from independent rootstock cutting to integrated grafted vine. First year grafted rootstock trunk temporal definition includes= spanning first complete growing season post-grafting (from grafting date through first dormancy typically May-November in Northern Hemisphere), representing initial 4-8 months of grafted existence (shorter period than full calendar year, focused on first active growing season), beginning at successful graft union formation (within 2-4 weeks post-grafting when callus bridge initiates), concluding at first dormancy (end of first growing season when young trunk enters winter dormancy), and marking year 1 developmental stage (first of 2-3 year young trunk period before transition to mature status). First year grafted rootstock trunk graft union establishment includes= forming initial callus tissue (wound healing cells accumulating at graft interface within first 2-3 weeks creating bridge between rootstock and scion), establishing first vascular connections (initial xylem and phloem continuity across graft union developing by 4-6 weeks post-grafting enabling limited transport), showing prominent callus bulge (swelling at graft union 1.2-2.5 cm diameter increase very visible in first year due to active callus proliferation), displaying fresh graft union scar (grafting tape or wax potentially still present or recently removed revealing union line), and beginning integration process (rootstock and scion tissues starting to unite though full integration requiring additional 1-2 years). First year grafted rootstock trunk physical characteristics includes= exhibiting very small diameter (0.8-2.5 cm typical range reflecting limited growth in first season), showing minimal bark development (absent or very thin initial periderm 0.1-0.4 mm thickness, often still showing greenish color), displaying smooth surface (no longitudinal fissures or furrows characteristic of older trunks, surface largely intact epidermis or initial thin cork), revealing visible lenticels (gas exchange pores conspicuous as light-colored dots on smooth young trunk surface), and showing green undertones (chlorophyll visible through thin bark or epidermis giving trunk greenish cast especially in shade-protected areas). First year grafted rootstock trunk extreme vulnerability includes= highly susceptible to mechanical damage (very thin bark and small diameter providing minimal protection against equipment injury, handling damage, or animal browsing), extremely sensitive to environmental stress (freeze damage, sunburn, wind damage, or desiccation potentially fatal to first year trunk), very vulnerable to graft union failure (mechanical disruption of developing graft union or poor initial callus formation potentially leading to graft failure and vine death), requiring absolute support (staking essential preventing wind breakage of weak first year trunk and protecting developing graft union), and showing high disease susceptibility (young tissues vulnerable to crown gall infection at graft union, trunk cankers, or systemic diseases). First year grafted rootstock trunk vascular development includes= establishing initial water transport (limited functional xylem providing minimal water conduction capacity 2-10 liters daily insufficient for large canopy), forming first phloem connections (limited photosynthate transport from scion to rootstock restricting carbohydrate storage), showing incomplete vascular integration (partial rather than complete continuity across graft union limiting transport efficiency), requiring time for vessel maturation (xylem elements at graft union initially narrow and irregular maturing over subsequent years), and developing secondary vascular tissue (vascular cambium activity beginning to add xylem and phloem thickness building transport capacity). First year grafted rootstock trunk growth patterns includes= showing limited diameter increase (typically 0.3-0.8 cm growth increment first season reflecting establishment rather than expansion focus), displaying callus-driven growth (graft union bulge increasing more rapidly than trunk diameter as callus tissue accumulates), exhibiting minimal height increase (2-5 cm vertical growth primarily from graft union callus rather than intercalary trunk elongation), focusing energy on integration (limited diameter growth reflecting resource allocation to graft union formation rather than general trunk expansion), and showing variable growth (depending on grafting timing, technique, rootstock vigor, growing conditions, and post-graft care). First year grafted rootstock trunk management requirements includes= providing essential staking (sturdy stake supporting weak first year trunk preventing wind damage and maintaining graft union alignment), ensuring graft union protection (wrapping graft union with tape, grafting compound, or protective covering preventing desiccation and mechanical damage), implementing trunk protection measures (painting trunk white or wrapping with protective material in hot climates preventing sunburn damage), maintaining optimal irrigation (regular water supply essential supporting establishment without waterlogging which could promote disease), removing competing growth (eliminating any rootstock shoots below graft union that compete with scion establishment), and intensive monitoring (frequent inspection detecting problems early when intervention most effective). First year grafted rootstock trunk in field grafting includes= grafted in situ (rootstock already planted in vineyard when grafting occurs typically on 1-3 year old rootstock trunk), showing shorter below-graft trunk (10-20 cm from soil surface to graft union in field grafts compared to bench grafts), requiring careful post-graft protection (exposed graft union vulnerable to weather and damage requiring immediate protection), potentially showing more variable success (field conditions more challenging than controlled nursery environment affecting first year establishment), and needing season-long support (field-grafted first year trunk requiring continued stake support through entire first season and winter). First year grafted rootstock trunk in bench grafting includes= grafted before planting (rootstock cutting grafted to scion cutting in controlled nursery conditions typically winter/early spring), developing in nursery (first few months of first year occurring in nursery callusing room and nursery beds), showing longer below-graft trunk (15-30 cm rootstock portion typical in bench-grafted vines), planted as established graft (moved to vineyard after initial callus formation giving some advantage over field grafts), and potentially showing better uniformity (controlled conditions supporting more consistent first year development). First year grafted rootstock trunk survival factors includes= successful callus formation (adequate callus bridge development essential for graft union survival), adequate water supply (preventing desiccation stress while avoiding waterlogging maintaining developing trunk), protection from extremes (avoiding freeze damage, sunburn, or severe wind damage during vulnerable first year), disease avoidance (preventing crown gall, fungal cankers, or systemic infections in susceptible young trunk), and appropriate scion-rootstock compatibility (compatible combination supporting successful first year integration rather than early rejection). First year grafted rootstock trunk failure risks includes= graft union failure (insufficient callus formation or mechanical disruption leading to separation typically within first 2-4 months), freeze damage (severe cold killing young trunk cambium and developing graft union particularly in late spring frosts), sunburn damage (intense sun exposure causing bark necrosis and cambium death on southwest-facing young trunk), mechanical breakage (wind damage or handling injury breaking weak first year trunk or disrupting graft union), crown gall infection (Agrobacterium tumefaciens entering through graft union wounds potentially fatal to first year trunk), and desiccation (inadequate water supply during critical establishment period leading to trunk death). First year grafted rootstock trunk monitoring priorities includes= inspecting graft union (checking for callus formation, absence of separation, no disease symptoms), assessing trunk condition (evaluating bark integrity, absence of sunburn or freeze damage, healthy green/tan coloration), checking stake security (ensuring support maintains trunk upright and protects graft union from stress), monitoring water status (avoiding both drought stress and excessive moisture preventing disease), removing suckers promptly (eliminating rootstock shoots below graft union preventing resource competition), and evaluating scion growth (assessing whether scion developing appropriately indicating successful graft and adequate trunk function). First year grafted rootstock trunk significance includes= representing most critical developmental period (first year determining whether graft succeeds or fails establishing foundation for entire vine lifespan), requiring maximum attention and care (first year trunk needs most intensive management and protection of any vine developmental stage), establishing graft union integration (first year callus formation and vascular connection critical for all subsequent vine function), marking beginning of grafted vine existence (first year trunk representing transition from separate rootstock and scion to unified grafted organism), and predicting long-term success (first year trunk development quality indicating probability of establishing productive long-lived vine).
First_Year_Grafted_Rootstock_Trunk emphasizes initial growing season post-grafting and is distinguished from Second_Year_Grafted_Rootstock_Trunk (second growing season) by focus on most critical early establishment period. Critical for understanding graft union formation, extreme vulnerability, intensive management needs, establishment success factors, and the foundational first season determining grafted vine survival and development.
Anfangsstadium Pfropfstamm
Erstes Jahr verpfropfter Unterlagenstamm
Initial season grafted rootstock trunk
Neuverpfropfter Wurzelstockstamm
Newly grafted rootstock trunk
Year 1 post-graft rootstock trunk
First Year Grafted Rootstock Trunk
Chardonnay first year grafted scion trunk on 420A rootstock planted April in Burgundy shows by November dormancy 7-14 mm scion diameter at 20 cm above graft union, single scion annual ring 2.0-3.5 mm width, graft union at 9 cm above soil with 108-115% diameter swelling, 75-120 cm scion shoot length, rootstock diameter 9-16 mm at 10 cm below union, combined starch reserves 12-20 g (scion 4-7 g, rootstock 8-13 g), and 80% survival rate with adequate management
A first year grafted scion trunk is a young grafted scion trunk during its initial growing season following vineyard planting of grafted vine, characterized by graft union stabilization in field conditions, establishment of scion-rootstock vascular connection, and initial scion shoot development from nursery-grafted plant, representing the most vulnerable establishment phase requiring intensive management to ensure graft success and vine survival. First year grafted scion trunk development from planted grafted vine proceeds= at planting (typically dormant season) vine has established graft union from nursery with 1-2 scion buds above union and rooted rootstock below, during spring budbreak (BBCH 07-09) scion shoot emerges from union bud, during early growth (BBCH 11-15, weeks 1-6) scion shoot elongates with primary growth while rootstock roots expand, during mid-season (BBCH 17-31, weeks 6-12) vascular cambium initiates in scion shoot base beginning secondary growth, during late season (BBCH 33-39, weeks 12-20) progressive secondary growth extends through developing scion trunk, and during senescence (BBCH 91-97) first dormancy with 5-15 mm scion trunk diameter achieved above union. First year grafted scion trunk graft union field establishment includes= union exposure to field environmental conditions (temperature fluctuations, moisture variation, physical stresses different from nursery), union vascular connection strengthening under transpirational demand (field-grown scion transpiration testing hydraulic continuity across graft interface), potential union incompatibility early manifestation (poor scion growth, wilting, or death indicating failed or incompatible graft), rootstock sucker emergence requiring removal (rootstock buds below union attempting growth competing with scion), and union position maintenance (ensuring union remains above soil preventing scion rooting that would bypass rootstock resistance benefits). First year grafted scion trunk anatomical development includes= scion initial primary growth with primary xylem and phloem (weeks 1-6), scion cambium initiation beginning secondary growth (weeks 6-10 in shoot base), first secondary xylem production forming initial scion annual ring (typically 1.5-4 mm radial increment), scion periderm initiation replacing primary epidermis (beginning scion bark formation), and graft union callus maturation (nursery callus lignifying and strengthening under field conditions). First year grafted scion trunk physiological priorities include= graft union hydraulic function (establishing adequate water transport from rootstock through union to scion supporting 8-15 scion leaves), rootstock root system expansion (roots growing 20-60 cm depth providing water and nutrient supply), scion photosynthetic capacity development (producing carbohydrates supporting both scion and rootstock growth), minimal to no fruit production (flowers removed prioritizing establishment), and initial carbohydrate storage (accumulating 3-8 g starch in scion trunk and 8-18 g in rootstock by season end). First year grafted scion trunk rootstock influence on establishment includes= vigorous rootstocks (1103P, SO4) promoting faster scion growth (shoot length 80-140 cm, trunk diameter 8-18 mm by dormancy), moderate rootstocks (5BB, 101-14Mgt) producing balanced establishment (shoot length 60-100 cm, diameter 6-14 mm), low-vigor rootstocks (3309C, Riparia Gloire) resulting in slower scion development (shoot length 50-90 cm, diameter 5-12 mm), and rootstock adaptation to soil affecting scion performance (lime-tolerant rootstocks performing well on calcareous soils supporting vigorous scion growth, while non-adapted rootstocks showing chlorosis and poor scion development). First year grafted scion trunk management is critical= adequate irrigation maintaining root zone moisture (particularly first 8-12 weeks critical for graft union function and survival), prompt rootstock sucker removal (removing rootstock shoots within 1-2 weeks of emergence preventing competition), scion shoot training to stake (supporting vertical growth, preventing union breakage from wind or equipment), protection from environmental extremes (frost protection, heat stress mitigation, particularly protecting vulnerable union), and pest and disease prevention (young tissue vulnerable, union representing potential pathogen entry point). First year grafted scion trunk challenges include= higher establishment mortality than own-rooted (15-25% potential loss from graft failure, union breakage, incompatibility), graft union vulnerability to mechanical damage (equipment contact, animal damage, wind stress at union particularly hazardous), union freeze damage risk (graft interface more cold-sensitive than uniform trunk tissue), rootstock suckering vigor (some rootstocks producing vigorous suckers requiring frequent removal), and delayed growth compared to vigorous own-rooted vines (graft union representing physiological resistance potentially slowing initial establishment).
First_Year_Grafted_Scion_Trunk emphasizes initial field establishment with graft union and is distinguished from First_Year_Own_Rooted_Trunk (no union, simpler establishment) and Second_Year_Grafted_Scion_Trunk (union stabilized, accelerated growth) by graft union field establishment challenges. Critical for understanding grafted vine establishment mortality factors, union vulnerability in first year, rootstock influence on initial scion development, and intensive management requirements for grafted vine success.
Erster Vegetationsjahr-Aufpropfung
Erstes Jahr gepfropfter Stamm
First season grafted scion
Initial year grafted trunk
Pflanzjahr Veredelungsstamm
First Year Grafted Scion Trunk
Chardonnay first year own-rooted trunk planted March in Burgundy shows by November dormancy 8-12 mm diameter at soil level, single annual growth ring 2.0-3.5 mm width, 85-110 cm shoot length reaching first trellis wire, 12-18 leaves produced during season supporting 8-12 g starch accumulation, and 85% survival rate with adequate irrigation and frost protection
A first year own-rooted trunk is a young own-rooted trunk during its initial growing season following planting of rooted cutting or seedling, characterized by establishment of root-shoot system, initiation of secondary growth, and development from herbaceous shoot to beginning woody stem, representing the most vulnerable establishment phase requiring intensive care and protection. First year own-rooted trunk development from rooted cutting proceeds= at planting (typically dormant season) cutting has established adventitious roots and dormant buds, during spring budbreak (BBCH 07-09) shoot emergence occurs from cutting buds, during early shoot growth (BBCH 11-15, weeks 1-6) herbaceous shoot elongates with primary growth only, during mid-season (BBCH 17-31, weeks 6-12) vascular cambium initiates in shoot base beginning secondary growth, during late season (BBCH 33-39, weeks 12-20) progressive secondary growth extends upward through developing trunk, and during senescence (BBCH 91-97) first dormancy occurs with 5-15 mm trunk diameter achieved. First year own-rooted trunk anatomical development includes= initial primary growth with primary xylem and phloem in vascular bundles, vascular cambium initiation between primary xylem and phloem (typically 6-10 weeks after budbreak in shoot base), first secondary xylem production forming initial annual ring (typically 1.5-4 mm radial increment), periderm initiation replacing primary epidermis (beginning of bark formation), and central pith remaining prominent with no heartwood formation. First year own-rooted trunk physiological priorities include= root system establishment and expansion (roots growing 20-60 cm depth and 20-40 cm radius during first season), adequate photosynthetic capacity for carbohydrate production (developing 8-15 leaves on primary shoot), water transport capacity supporting leaf area (initial xylem development providing 0.05-0.2 L/day transport capacity), minimal to no fruit production (flowers typically removed to prioritize vegetative establishment), and initial carbohydrate storage (accumulating 5-15 g starch reserves in roots and trunk base by season end). First year own-rooted trunk management is critical= adequate irrigation maintaining root zone moisture (particularly first 8-12 weeks preventing establishment failure), shoot training to stake or trellis (supporting vertical growth, preventing wind damage), protection from environmental extremes (frost protection in cool climates, shade or irrigation in hot climates), pest and disease prevention (young tissue vulnerable to many pests and pathogens), and shoot management (typically maintaining 1-2 primary shoots, removing others to focus growth). First year own-rooted trunk challenges include= high mortality risk (10-30% establishment failure in difficult conditions), vulnerability to drought stress (limited root system and high shoot transpiration), susceptibility to frost damage (tender shoot tissue easily damaged by late spring or early fall frosts), pest pressure (young vigorous growth attractive to aphids, mites, leafhoppers), disease susceptibility (powdery mildew, downy mildew on young foliage), and competition from weeds (vigorous weed growth can suppress young vine establishment). First year own-rooted trunk success indicators include= achieving 5-15 mm trunk diameter by dormancy, developing 10-20 leaves on primary shoot during growing season, shoot length reaching 60-120 cm (sufficient to reach first trellis wire), successful overwintering survival, and adequate root development (visible through gentle excavation showing 20-40 cm root radius).
First_Year_Own_Rooted_Trunk emphasizes initial establishment season and is distinguished from Second_Year_Own_Rooted_Trunk (second season with accelerated growth) and Establishing_Young_Own_Rooted_Trunk (years 2-3 combined) by focus on most vulnerable establishment phase. Critical for understanding establishment success factors, intensive management requirements, mortality risk factors, and foundation for subsequent vine development.
Erster Vegetationsjahr-Stamm
Erstes Jahr Stamm
First growing season trunk
Initial year trunk
Pflanzjahr-Rebstamm
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Winkler, A. J., Cook, J. A., Kliewer, W. M., & Lider, L. A. (1974). General Viticulture. University of California Press.
First Year Own Rooted Trunk
A five-plus arm head on Vitis vinifera cv. Carignan in Languedoc-Roussillon, featuring six arms emerging from swollen head at 52cm height, arms positioned irregularly (not symmetrically spaced) at various angles reflecting incremental addition over vine's 35-year history, arms extending 35-55cm bearing 4-7 spurs each, total 32 spurs supporting approximately 58 shoots and 6.8kg crop, representing evolved head training adapted to vigorous site through progressive arm addition as vine matured, now providing enhanced productivity (approximately 51 hl/ha) while maintaining head-trained architecture avoiding trellis conversion, common pattern in Languedoc where economic pressure drives moderate intensification of traditional systems.
A multi-arm head featuring five or more arms radiating from central head, representing enhanced-capacity head training beyond traditional four-arm standard, characterized by complex multi-arm architecture (5, 6, 7, or occasionally 8+ arms), increased fruiting capacity through additional structural elements, irregular or symmetrical spacing depending on deliberate design versus organic development, and suitability for vigorous vines, fertile sites, or productivity-enhanced adaptations of traditional head training. Five-plus arm heads represent high-capacity head training, employed where traditional three-four arm configurations inadequately utilize vine vigor (vigorous varieties, fertile soils, generous water availability), adapted traditional systems modified for improved productivity while maintaining head-trained character, incremental arm addition over time as vines mature and vigor increases, or complex irregular structures resulting from opportunistic renewal and arm replacement creating accumulated multiple arms.
Five-plus arm heads represent the upper range of head-trained capacity, with additional arms increasing productivity beyond traditional three-four arm standards while maintaining head-trained architecture. The five-plus configuration can be deliberate (intentional five-six arm symmetric design) or organic (accumulated arms through incremental addition over decades), symmetrical (regular spacing) or irregular (opportunistic positioning), and stable (established five-six arm architecture maintained through renewal) or transitional (continuing addition toward seven-eight arms as vigor permits). Benefits include increased productivity (25-60% yield improvement versus four-arm standard), better vigor utilization (adequate capacity for vigorous vines), and production insurance (redundancy across numerous arms). Challenges include management complexity (coordinating pruning and renewal across many arms), potential structural congestion (many arms competing for space at head), irregular architecture (especially in organic development patterns), and uncertain optimal limits (unclear whether seven-eight+ arms provide benefits or create excessive complexity).
Mehrarmen-Kopf
enhanced capacity head
high-arm-count head
many-armed head
verstärkte Kopferziehung
Five Plus Arm Head
A five-plus-branch tendril at node 5 on an exceptionally vigorous shoot of Vitis vinifera cv. Muscat of Hamburg in a heavily fertilized experimental vineyard in Italy, exhibiting complex architecture with primary trifurcation at 4.5 cm from shoot attachment, secondary bifurcation of two primary arms and tertiary bifurcation of one secondary arm, creating total of six terminal branches ranging 6.5-9.5 cm individual length with combined extension of 48 cm across all branches, individual arm diameters 2.0-2.6 mm showing resource dilution across the numerous branches, highly complex three-dimensional architecture with branches exploring complementary spatial zones collectively covering approximately 1150 cubic centimeters, four branches making successful attachment to various trellis wires and one stake creating exceptional multi-point support system, occurring on a shoot with extreme vigor (total length 245 cm, 22 nodes, 14 mm basal diameter) receiving luxury nitrogen fertilization (180 kg N/ha) and full irrigation throughout development, measured at BBCH stage 71 (fruit set) in June, representing the rare achievement of functional extreme multi-branching requiring exceptional resource availability rarely encountered in commercial viticulture.
A multi-branched tendril exhibiting five or more terminal branches in Vitis vinifera, characterized by complex hierarchical architecture with multiple branching levels creating numerous independent arms, combined total tendril length across all branches often exceeding 40-60 cm, substantial total resource investment rivaling or exceeding resource allocation to small leaves, representing the extreme upper range of branching complexity occurring in less than 1-3% of branched tendrils in most cultivars with higher frequencies only in specific varieties showing strong genetic predisposition toward maximal branching under exceptional vigor conditions. Five-plus-branch tendrils develop through extensive meristematic activity during early tendril development creating multiple branching events, either through very rare five-way or six-way single branch points, through multiple sequential bifurcations creating deep hierarchical structures with three or four branching levels, or through mixed patterns combining various branching configurations across multiple levels, requiring sustained high resource availability throughout the extended development period necessary to support numerous growing points and extensive total growth. The functional implications include maximum possible spatial exploration volumes potentially exceeding 1000 cubic centimeters, very high probability of achieving 3-5 or more successful attachments creating exceptionally robust and redundant mechanical support systems, excellent load distribution capacity across numerous attachment points enabling support of very heavy shoot sections or extreme fruit loads, but resource costs approaching those of moderate-sized leaves raising questions about cost-benefit optimization and suggesting five-plus branching may represent overinvestment in climbing function except under specific conditions where maximum mechanical security provides critical survival advantage.
Five-plus-branch tendrils represent the absolute upper extreme of branching complexity in Vitis vinifera and are distinguished from four-branch tendrils by five or more terminal arms. These structures are extremely rare in commercial viticulture, with occurrence typically limited to unusually vigorous shoots in highly fertilized or irrigated situations, occasionally in cultivars with genetic tendencies toward maximum branching (certain Muscats, some table grape varieties), or in experimental conditions with luxury resource availability. The resource cost of five-plus-branch tendrils raises functional questions, as the metabolic investment approaches or exceeds that required for moderate leaves which would provide photosynthetic return versus the purely mechanical function of climbing support. Ecological contexts where extreme branching provides selective advantage likely involve climbing through dense vegetation where maximum contact probability and mechanical redundancy justify high resource costs. In commercial viticulture, frequent five-plus-branch tendrils may indicate excessive vegetative vigor requiring management intervention through reduced fertilization, crop load increase, or vigor control strategies.
extreme multi-branch tendril
fünf-plus-armige Ranke
highly complex tendril
hochkomplex verzweigte Ranke
maximal ramifizierte Kletterorgan
maximally branched tendril
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 198-215 This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 259-282.
Five Plus Branch Tendril
Flavan-3-ol content quality assessment in Napa Valley Cabernet Sauvignon guides extraction management, with seed flavan-3-ol concentration (8-15 mg/g seed dry weight) indicating the astringent potential that must be carefully managed through maceration timing winemakers monitor seed color change from green to brown as a visual proxy for flavan-3-ol polymerization and reduced extractability, pressing red wines when seed taste shifts from harsh astringency to nutty character, typically at 18-24 days post-crush for premium Cabernet Sauvignon.
Flavan_3_ol_Content_Quality encompasses the chemical qualities characterizing the concentration and composition of flavan-3-ol compounds in grape berry tissues, capturing the flavonoid subclass that serves as both monomeric taste-active compounds and the building blocks for polymeric condensed tannins. Flavan-3-ols are characterized by a saturated C ring with hydroxyl groups at the C3 position, distinguishing them from the unsaturated flavonols. The primary grape flavan-3-ols are (+)-catechin, (−)-epicatechin, and (−)-epicatechin-3-gallate, with (+)-gallocatechin and (−)-epigallocatechin present in lesser amounts in some cultivars. Flavan-3-ols accumulate primarily in grape seeds, where they comprise the dominant phenolic class, with smaller amounts in skins where they contribute to skin tannin composition. The localization differs between tissues: seeds contain primarily catechin and epicatechin with their gallate esters, while skins contain higher proportions of epigallocatechin reflecting F3'5'H activity in skin tissues. Flavan-3-ol monomers contribute to taste through direct astringency and bitterness perception, with epicatechin and particularly epicatechin-gallate perceived as more bitter and astringent than catechin. More importantly, flavan-3-ols polymerize through C4-C6 and C4-C8 linkages to form proanthocyanidins (condensed tannins), the major astringent compounds in red wine. The concentration of flavan-3-ols in grape tissues provides an indicator of tannin potential, though the polymerization state critically affects sensory properties. Flavan-3-ol accumulation occurs primarily during early berry development, with seed flavan-3-ols largely established before véraison while skin flavan-3-ols continue accumulating through ripening.
Flavan_3_ol_Content_Quality serves as the parent class for individual flavan-3-ol monomers including Catechin_Content_Quality and Epicatechin_Content_Quality. The class addresses monomeric flavan-3-ols distinct from their polymeric forms (proanthocyanidins/condensed tannins) treated under Tannin_Quality. The stereochemistry of flavan-3-ols (catechin versus epicatechin series, cis versus trans) affects both their polymerization behavior and sensory properties. Seed and skin flavan-3-ol pools differ in composition and must be considered separately for complete characterization. Analysis employs HPLC with UV or fluorescence detection, or phloroglucinolysis to characterize polymer subunit composition.
Catechinverbindungsgehalt
Flavan-3-ol-Gehalt
Flavanolgehalt
Proanthocyanidin-Vorläuferkonzentration
catechin-type compound content
flavan-3-ol monomer quality
flavanol content
proanthocyanidin precursor concentration
Kennedy JA et al. (2000) Composition of Grape Skin Proanthocyanidins at Different Stages of Berry Development. Journal of Agricultural and Food Chemistry 48(12): 5348-5355 Downey MO et al. (2003) Analysis of Tannins in Seeds and Skins of Shiraz Grapes Throughout Berry Development. Australian Journal of Grape and Wine Research 9(1): 15-27
Flavan 3 ol Content Quality
Systematic screening for Flavescence_Doree_Resistance_Disposition at INRAE Bordeaux evaluated FD responses across 120 V. vinifera cultivars and 30 interspecific genotypes using controlled transmission via infectious Scaphoideus titanus collected from FD-infected vineyards. Test plants were exposed to viruliferous leafhoppers (5 per plant) for a 3-week acquisition access period, then maintained in insect-proof conditions for 24 months with periodic PCR testing and symptom assessment. Among V. vinifera cultivars, infection rates ranged from 35% ('Cabernet Sauvignon') to 95% ('Baco Blanc'), with symptom severity scores of 1.5 to 4.8 on a 0-5 scale. Recovery rates (symptom remission while remaining PCR-positive) ranged from 0% ('Chardonnay') to 45% ('Ugni Blanc'). The most resistant V. vinifera cultivar identified was 'Cabernet Sauvignon' with low infection rate, mild symptoms (score 1.5), and 25% recovery. Among interspecific genotypes, several rootstock varieties including '3309 Couderc' (V. riparia × V. rupestris) showed reduced infection rates (15-25%) and minimal symptom expression, suggesting enhanced Flavescence_Doree_Resistance_Disposition in non-vinifera germplasm. Importantly, cultivar differences in vector feeding preference did not correlate with infection rates, indicating that resistance operates primarily at the plant-phytoplasma interaction rather than plant-vector level.
A Flavescence_Doree_Resistance_Disposition is a Phytoplasma_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that limit infection, multiplication, systemic spread, symptom development, or vector transmission of 'Candidatus Phytoplasma vitis', the causal agent of Flavescence dorée (FD), the most severe and regulated grapevine yellows disease in Europe. This disposition represents the capacity to resist a quarantine pathogen that has caused devastating epidemics across European wine regions since its emergence in southwestern France in the 1950s, necessitating mandatory control measures including insecticide treatments against the vector Scaphoideus titanus and removal of infected vines. Flavescence dorée produces characteristic symptoms including downward leaf rolling with golden yellowing (white cultivars) or reddening (red cultivars) beginning from basal leaves and progressing acropetally, rubber-like flexibility of incompletely lignified canes, flower abortion or berry shriveling, and rapid vine decline often leading to death within 2-3 years. Resistance mechanisms may include restriction of phytoplasma multiplication in phloem, limitation of systemic colonization, resistance to phytoplasma-induced metabolic reprogramming, and recovery capacity enabling symptom remission. Additionally, resistance to the leafhopper vector S. titanus can provide indirect protection by reducing transmission. No V. vinifera cultivar shows immunity to FD, but substantial variation in susceptibility, symptom severity, and recovery rates provides opportunities for cultivar selection in endemic regions. Some interspecific hybrids and rootstocks show enhanced tolerance or resistance, potentially contributing to reduced disease pressure in mixed plantings.
Flavescence_Doree_Resistance_Disposition is_a Phytoplasma_Resistance_Disposition. Flavescence dorée is a quarantine disease in the EU (EPPO A2 list) with mandatory control measures including vector treatment and infected vine removal. The child class Scaphoideus_Titanus_Resistance_Disposition addresses vector resistance as an indirect protection mechanism. True immunity to FD has not been identified in Vitis, making tolerance (reduced symptoms despite infection) and recovery (symptom remission) the relevant resistance phenotypes. Cultivar selection can meaningfully reduce disease impact but cannot eliminate the need for vector control in endemic areas.
Candidatus Phytoplasma vitis resistance
FD resistance disposition
FD-Resistenz
Flavescence-dorée-Resistenz
Goldgelbe-Vergilbung-Resistenz-Disposition
golden flavescence resistance
Chuche J, Thiéry D (2014) Biology and ecology of the Flavescence dorée vector Scaphoideus titanus, Journal of Pest Science 87:381-393 Eveillard S et al. (2016) Contrasting susceptibilities to Flavescence dorée in Vitis vinifera, rootstocks and wild Vitis species, Frontiers in Plant Science 7:1762 EFSA Panel on Plant Health (2014) Scientific opinion on pest categorisation of Grapevine flavescence dorée, EFSA Journal 12:3851
Flavescence Doree Resistance Disposition
In Sangiovese berries in Chianti Classico, Tuscany during BBCH 75-89, flavonoid accumulation establishes flavonol content reaching 145 mg/kg in sun-exposed clusters compared to 28 mg/kg in shaded fruit, with quercetin-3-glucoside at 85 mg/kg, quercetin-3-glucuronide at 35 mg/kg, and kaempferol-3-glucoside at 18 mg/kg, concurrent with skin proanthocyanidin accumulation to 3.2 mg/g and anthocyanin accumulation to 1.15 mg/g fresh weight by harvest under warm Tuscan conditions averaging 26°C.
Flavonoid accumulation encompasses the progressive increase in diverse flavonoid compound concentrations within Vitis vinifera berry exocarp tissues during development and ripening, wherein the coordinated expression of flavonoid biosynthetic pathway genes enables synthesis and deposition of flavonols, flavan-3-ols, and in red varieties anthocyanins that collectively establish the flavonoid profile determining wine color, copigmentation capacity, antioxidant status, and potential health-related bioactive properties in wines produced from these grapes. This process integrates multiple branches of flavonoid metabolism, with early pathway enzymes including chalcone synthase, chalcone isomerase, and flavanone 3-hydroxylase providing common intermediates that branch to flavonol synthase generating quercetin, kaempferol, and myricetin glycosides accumulating primarily in epidermal cells, to leucoanthocyanidin reductase and anthocyanidin reductase producing flavan-3-ols for proanthocyanidin synthesis, and to anthocyanidin synthase and subsequent enzymes generating anthocyanins in red variety hypodermal cells. The temporal dynamics of flavonoid accumulation show compound-class-specific patterns, with flavonol synthesis occurring primarily during early berry development before veraison in response to UV radiation exposure serving photoprotective functions, proanthocyanidin accumulation in skin tissues continuing through veraison and into post-veraison stages, and anthocyanin synthesis initiating at veraison and continuing throughout ripening as previously described. The quantitative aspects of flavonoid accumulation demonstrate substantial environmental responsiveness, with light exposure essential for flavonol induction through UV-B responsive regulatory elements in biosynthetic gene promoters, moderate water stress enhancing overall flavonoid accumulation through metabolic activation and berry size effects, and temperature influencing the balance among pathway branches with cool conditions favoring certain compound classes. Flavonol accumulation reaches characteristic levels of 50-200 mg/kg berry fresh weight in sun-exposed fruit compared to 10-30 mg/kg in heavily shaded clusters, with composition dominated by quercetin derivatives including quercetin-3-glucoside and quercetin-3-glucuronide in most varieties, while kaempferol and myricetin glycosides contribute varying proportions depending on hydroxylase expression patterns. The significance of flavonoid accumulation for wine quality extends beyond direct sensory contributions to include copigmentation effects where flavonols and other colorless flavonoids stack with anthocyanins enhancing color intensity by 30-50%, antioxidant capacity protecting both grape and wine from oxidative deterioration, and potential health benefits associated with flavonoid consumption that contribute to wine's position in dietary recommendations when consumed moderately.
Flavonoid_Accumulation is subclass of Skin_Phenolic_Development representing comprehensive flavonoid compound increase in exocarp. Distinguishes from specific compound classes by integrating multiple flavonoid types. Sibling to Skin_Tannin_Maturation and Skin_Phenolic_Profile. Includes flavonols, flavan-3-ols, and anthocyanins. Light-responsive with UV-B induction of flavonols. Copigmentation function for color enhancement.
Exokarp-Flavonoid-Synthese
Flavonoid-Akkumulation
Flavonol-und-Flavan-3-ol-Anreicherung
Haut-Flavonoid-Entwicklung
berry flavonoid accumulation
exocarp flavonoid synthesis
flavonol and flavan-3-ol accumulation
skin flavonoid development
Downey MO et al (2004) The effect of bunch shading on berry development and flavonoid accumulation in Shiraz grapes. Australian Journal of Grape and Wine Research 10: 55-73 Price SF et al (1995) Cluster sun exposure and quercetin in Pinot noir. American Journal of Enology and Viticulture 46: 187-194 Keller M (2020) The Science of Grapevines 3rd Ed pp 310-325
Flavonoid Accumulation
Detailed investigation of Flavonoid_Accumulation_Disposition at Stellenbosch University characterized flavonoid profiles and UV-screening capacity across 15 cultivars grown under ambient and supplemental UV-B (12 kJ m⁻² d⁻¹) for 28 days. Epidermal flavonoids were extracted and quantified by HPLC, UV transmittance was measured spectrophotometrically on epidermal peels, and expression of phenylpropanoid pathway genes (VvPAL, VvCHS, VvFLS1) was assessed by qRT-PCR. Flavonoid_Accumulation_Disposition showed substantial cultivar variation in both constitutive levels and UV inducibility. 'Malbec' exhibited exceptional flavonoid accumulation with constitutive epidermal flavonol content of 1.85 mg/g FW increasing to 3.42 mg/g under UV supplementation (1.85× induction), dominated by quercetin-3-glucoside (45%) and quercetin-3-glucuronide (28%). UV-B transmittance through 'Malbec' epidermis was only 6% at 310 nm, providing effective screening. VvFLS1 expression increased 12-fold under UV in 'Malbec'. 'Sauvignon Blanc' showed strong Flavonoid_Accumulation_Disposition (2.95 mg/g under UV, 8% transmittance) with distinct flavonol profile (kaempferol-dominant). 'Chasselas' displayed weak accumulation (1.25 mg/g under UV, only 1.4× induction, 18% transmittance) with correspondingly low VvFLS1 induction (3.5-fold). Correlation analysis revealed that UV-B transmittance was strongly predicted by total flavonol content (r = -0.92) and that flavonol accumulation correlated with photosynthetic protection under UV stress (r = 0.85 with Fv/Fm maintenance). Anthocyanins provided additional UV protection in red cultivar berries, with 'Malbec' berry skin anthocyanins absorbing 40% of incident UV-B.
A Flavonoid_Accumulation_Disposition is a UV_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through enhanced capacity to synthesize and accumulate UV-absorbing flavonoid compounds in epidermal tissues, creating a chemical sunscreen that prevents UV penetration to underlying sensitive mesophyll cells, chloroplasts, and nuclei. This disposition represents the primary protective mechanism against UV damage in plants, with flavonoids absorbing strongly in the UV-B and UV-A wavelengths while transmitting visible light required for photosynthesis. Key UV-protective flavonoids in grapevine include flavonols (quercetin, kaempferol, and their glycosides), which absorb maximally at 250-270 nm and 350-380 nm, and hydroxycinnamic acids (caffeic, ferulic, coumaric acids and derivatives), which absorb at 290-330 nm. These compounds accumulate predominantly in the vacuoles of epidermal cells, creating a UV-absorbing layer protecting underlying tissues. Flavonoid_Accumulation_Disposition encompasses both constitutive flavonoid levels (providing baseline protection) and UV-inducible accumulation (enhanced synthesis in response to UV exposure), with the phenylpropanoid pathway enzymes (PAL, CHS, CHI, F3H, FLS) upregulated by UV-B through the UVR8 photoreceptor signaling pathway. Genotypic variation in Flavonoid_Accumulation_Disposition reflects differences in constitutive pathway flux, UV-responsiveness of pathway genes, and potentially the specific flavonoid profile affecting UV absorption spectrum and cellular localization.
Flavonoid_Accumulation_Disposition is_a UV_Tolerance_Disposition. Flavonoids (particularly flavonols) are the primary UV-screening compounds in leaves. Accumulation occurs predominantly in epidermal vacuoles, creating a UV-absorbing layer. Both constitutive and UV-inducible accumulation contribute to protection. UVR8 photoreceptor mediates UV-induced pathway activation. Flavonol profiles (quercetin vs kaempferol dominance) vary among cultivars. Anthocyanins provide additional UV protection in pigmented tissues.
Epidermale-Flavonoid-Synthese
Flavonoid-Akkumulations-Disposition
UV-Schutzverbindungs-Akkumulation
UV-screening compound accumulation
epidermal flavonoid synthesis
phenylpropanoid UV protection
Agati G, Tattini M (2010) Multiple functional roles of flavonoids in photoprotection, New Phytologist 186:786-793 Jenkins GI (2009) Signal transduction in responses to UV-B radiation, Annual Review of Plant Biology 60:407-431 Kolb CA et al. (2001) UV-A screening in epidermis of grapevine leaves, Australian Journal of Plant Physiology 28:515-522
Flavonoid Accumulation Disposition
Flavonoid quality assessment in Willamette Valley Pinot Noir integrates measurements of skin anthocyanins (200-500 mg/kg berry fresh weight), flavonols (15-40 mg/kg), and skin plus seed tannins (1.5-3.5 mg/g berry fresh weight) to characterize the phenolic potential of fruit from different vineyard blocks the relatively low anthocyanin content typical of Pinot Noir compared to Cabernet Sauvignon or Syrah requires careful extraction management to achieve sufficient color intensity without over-extracting seed tannins, making the flavonoid profile a critical guide for winemaking decisions in this thin-skinned, challenging variety.
Flavonoid_Quality encompasses the chemical qualities characterizing the concentration and composition of flavonoid compounds in grape tissue, capturing the major phenolic subclass defined by the C6-C3-C6 carbon skeleton consisting of two aromatic rings (A and B) connected by a three-carbon bridge that typically forms a heterocyclic C ring. Flavonoids represent the dominant phenolic class in grape skins and seeds, comprising anthocyanins (responsible for red and blue colors), flavonols (yellow pigments and co-pigmentation factors), and flavan-3-ols (the building blocks of condensed tannins). These three flavonoid subclasses share common biosynthetic origins through the phenylpropanoid and flavonoid pathways, with branch points controlled by specific enzymes that determine the proportional allocation among end products. Flavonoid accumulation in berries follows distinct spatial and temporal patterns: anthocyanins accumulate exclusively in the skin of colored cultivars from véraison onward flavonols concentrate in the skin in response to UV light exposure and flavan-3-ols accumulate primarily in seeds during early berry development, with skin flavan-3-ols synthesized throughout ripening. The flavonoid profile varies dramatically among cultivars, providing chemical signatures that distinguish varieties and contribute to wine typicity. Environmental factors strongly influence flavonoid content, with light exposure promoting flavonol and anthocyanin synthesis while temperature affects accumulation through both biosynthetic and degradative effects. The extractability of flavonoids during winemaking varies with compound class and tissue localization, making grape flavonoid content a potential indicator rather than direct predictor of wine composition.
Flavonoid_Quality serves as the parent class for major flavonoid subclasses including Anthocyanin_Content_Quality, Flavonol_Content_Quality, and Flavan_3_ol_Content_Quality. The C6-C3-C6 skeleton distinguishes flavonoids from simpler C6-C3 phenylpropanoids (hydroxycinnamic acids) and C6-C2-C6 stilbenoids. Flavonoid analysis typically employs HPLC separation with UV-Vis or mass spectrometric detection, enabling both quantification of compound classes and identification of individual species. The co-occurrence of flavonoid classes in the same biosynthetic pathway creates potential for metabolic trade-offs where enhanced flux toward one class may reduce accumulation of others.
C6-C3-C6 phenolic quality
C6-C3-C6-Phenolgehalt
Flavonoidgehalt
Flavonoidqualität
Flavonoidzusammensetzung
flavonoid composition
flavonoid compound quality
flavonoid content
Teixeira A et al. (2013) Berry Phenolics of Grapevine under Challenging Environments. International Journal of Molecular Sciences 14(9): 18711-18739 Boss PK et al. (1996) Expression of Anthocyanin Biosynthesis Pathway Genes in Red and White Grapes. Plant Molecular Biology 32(3): 565-569
Flavonoid Quality
Flavonol concentration in grape skins ranges from 2-100 mg/kg berries across cultivars and light exposure conditions, with quercetin-3-O-glucoside and quercetin-3-O-glucuronide typically predominating. High-flavonol cultivars such as Pinot Noir (40-100 mg/kg) and Cabernet Sauvignon (30-80 mg/kg) contrast with lower-flavonol types such as Grenache (15-40 mg/kg). Light exposure is the dominant environmental modifier, with sun-exposed clusters showing 3-10 fold higher flavonol concentration than shaded clusters through VvFLS1 induction. The flavonol profile mirrors the anthocyanin profile regarding B-ring hydroxylation, with quercetin (3',4'-diOH) and myricetin (3',4',5'-triOH) reflecting F3'H and F3'5'H activity respectively. Flavonols serve as UV-B protectants in berry skins and copigments enhancing anthocyanin color expression in wine.
A phenolic compound disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate flavonol compounds (quercetin, myricetin, kaempferol, isorhamnetin, laricitrin, syringetin glycosides) in grape berry skins through flavonol synthase (VvFLS) activity, with genotype-specific total concentration and profile affecting wine color through copigmentation, bitterness contribution, and antioxidant capacity.
Flavonol accumulation shows the strongest light response among grape phenolics, making cluster exposure management a primary tool for modulating wine flavonol content. The copigmentation function of flavonols enhances and stabilizes red wine color by forming molecular stacks with anthocyanins, shifting the color equilibrium toward the colored flavylium form. Quercetin can form insoluble precipitates in wine under specific pH and temperature conditions, creating instability concerns in some contexts. The genetic basis involves VvFLS expression regulated by light-responsive VvMYB transcription factors.
FLS activity disposition
Flavonol-Disposition
Flavonolakkumulation
Flavonolbildung
flavonol glycoside trait
flavonol synthesis capacity
Downey MO et al. (2004) The effect of bunch shading on berry development and flavonoid accumulation in Shiraz grapes. Australian Journal of Grape and Wine Research 10:55-73 Martínez-Lüscher J et al. (2014) Sensitivity of grapevine phenology to water availability, temperature and CO₂ concentration. Frontiers in Environmental Science 2:48
Flavonol Accumulation Disposition
Flavonol content quality assessment in Marlborough Sauvignon Blanc guides canopy management decisions, with sun-exposed fruit developing quercetin-3-glucoside concentrations of 25-50 mg/kg compared to less than 5 mg/kg in heavily shaded clusters this 5-10 fold difference enables quality differentiation, as elevated flavonol content correlates with enhanced varietal thiol release during fermentation, linking the UV-protective phenolic response to the tropical fruit aromatics that define premium Marlborough Sauvignon Blanc style.
Flavonol_Content_Quality encompasses the chemical qualities characterizing the concentration and composition of flavonol compounds in grape berry skins, capturing the yellow-pigmented flavonoid subclass that plays critical roles in UV protection, co-pigmentation, and wine color stability. Flavonols are characterized by a 3-hydroxyflavone backbone with a double bond between C2 and C3 and a hydroxyl group at C3, distinguishing them structurally from the saturated flavan-3-ols. The primary grape flavonols are quercetin, kaempferol, and myricetin, present predominantly as glycosides with glucose, galactose, or glucuronic acid attached at the 3-position. Flavonol accumulation in grape skins is strongly light-dependent, with synthesis dramatically upregulated by UV-B radiation exposure, making flavonol content a sensitive indicator of fruit zone light environment. Sun-exposed berries may contain 5-20 times higher flavonol concentrations than shaded berries on the same vine, representing one of the largest environmental modulation effects among grape phenolics. This light responsiveness reflects the photoprotective function of flavonols, which absorb UV radiation and scavenge reactive oxygen species generated by light stress. In winemaking, flavonols contribute to color through co-pigmentation with anthocyanins, where non-covalent stacking interactions between flavonols and anthocyanins enhance and stabilize red-purple coloration. Flavonol glycosides are extracted during maceration and may undergo hydrolysis during aging, releasing aglycones that can precipitate and contribute to sediment. Total flavonol concentration in grape skins typically ranges from 10-100 mg/kg fresh weight depending on cultivar, light exposure, and developmental stage, with accumulation beginning before véraison and continuing through ripening.
Flavonol_Content_Quality serves as the parent class for individual flavonol species including Quercetin_Content_Quality, Kaempferol_Content_Quality, and Myricetin_Content_Quality. The strong light dependence of flavonol synthesis distinguishes this class from anthocyanins and tannins, which show more moderate light responses. Flavonol analysis typically employs HPLC with UV detection at 360 nm, with individual glycosides and aglycones quantified against authentic standards. The B-ring hydroxylation pattern (mono-, di-, or tri-hydroxylated) parallels the anthocyanidin pattern and reflects the same F3'H and F3'5'H enzyme activities, creating correlations between flavonol and anthocyanin profiles.
3-Hydroxyflavon-Konzentration
3-hydroxyflavone concentration
Flavonolgehalt
Gelbflavonoidgehalt
UV-Schutzphenolqualität
UV-protective phenolic quality
flavonol compound content
yellow flavonoid quality
Price SF et al. (1995) Cluster Sun Exposure and Quercetin in Pinot Noir Grapes and Wine. American Journal of Enology and Viticulture 46(2): 187-194 Downey MO et al. (2006) Cultural Practice and Environmental Impacts on the Flavonoid Composition of Grapes and Wine: A Review of Recent Research. American Journal of Enology and Viticulture 57(3): 257-268
Flavonol Content Quality
In Riesling berries in Wachau, Austria during BBCH 83-89, flavor complexity development establishes concurrent accumulation of free monoterpenes reaching 1.8 mg/L with 8 individual compounds above threshold, norisoprenoid precursors at 125 µg/L contributing 4 distinct aroma notes, and volatile thiol precursors at 185 µg/L providing 3 additional sensory dimensions, with trained panel complexity ratings increasing from 4.5 to 8.2 on 10-point scale as the number of identifiable aromatic notes increases from 6 to 14 over 48 days of ripening under cool conditions averaging 17°C.
Flavor complexity development describes the progressive diversification and integration of multiple flavor compound classes within Vitis vinifera berries during ripening, wherein the concurrent accumulation of terpenoids, norisoprenoids, thiols, esters, and other volatile classes creates increasingly multifaceted aromatic profiles that transcend simple varietal character to express the nuanced complexity distinguishing premium wine grapes from ordinary productions at harvest maturity. This process involves the parallel operation of multiple biosynthetic pathways each contributing distinct compound classes with characteristic sensory attributes, including floral notes from monoterpene alcohols, fruity-tropical character from volatile thiols and esters, spicy nuances from sesquiterpenes and phenolic derivatives, and complex background aromatics from norisoprenoids and lipid-derived compounds that collectively produce the layered sensory experience associated with quality wines. The temporal dynamics of flavor complexity development show compound-class-specific accumulation patterns that create evolving aromatic profiles throughout ripening, with early development of green-herbaceous character from C6 compounds and methoxypyrazines transitioning to floral-fruity dominance as monoterpenes and norisoprenoids accumulate during mid-to-late ripening, and final integration phases during the last weeks before harvest establishing the balanced complexity of optimally ripe fruit. Sensory evaluation of flavor complexity development employs multidimensional assessment methods including descriptive analysis mapping individual aromatic attributes, complexity indices integrating the number and balance of perceived notes, and trained panel ratings of overall aromatic complexity that correlate with wine quality potential and consumer preference. Cultivar genetics establish baseline complexity potential through variety-specific expression of multiple biosynthetic pathway genes, with certain varieties such as Riesling demonstrating inherent complexity through balanced accumulation of terpenoids, norisoprenoids, and volatile thiols, while other varieties may require specific terroir or viticultural conditions to achieve comparable aromatic multidimensionality. Environmental factors profoundly influence complexity development through differential effects on individual biosynthetic pathways, with the combination of adequate light exposure for terpenoid synthesis, moderate temperatures for norisoprenoid accumulation, and appropriate vine stress for thiol precursor formation creating the conditions necessary for optimal complexity development in premium vineyard sites.
Flavor_Complexity_Development is subclass of Flavor_Compound_Accumulation representing diversification of aromatic profile. Distinguishes from Flavor_Intensity_Increase which emphasizes potency. Sibling to Progressive_Flavor_Build_Up and Flavor_Intensity_Increase. Key characteristic is parallel accumulation of multiple compound classes. Measured by number of distinct aromatic notes and complexity indices. Quality differentiator for premium wines.
Aroma-Komplexitäts-Entwicklung
Geschmacks-Vielschichtigkeit-Entwicklung
aroma diversification
aromatic complexity increase
aromatische Diversifizierung
complex flavor formation
flavor multidimensionality development
komplexe Aroma-Bildung
Robinson AL et al (2014) Origins of grape and wine aroma. American Journal of Enology and Viticulture 65: 1-30 Ferreira V (2010) Volatile aroma compounds and wine sensory attributes. In Managing Wine Quality Vol 1 Woodhead Publishing pp 3-28 Keller M (2020) The Science of Grapevines 3rd Ed pp 300-310
Flavor Complexity Development
In Torrontés berries in Cafayate, Argentina during BBCH 83-89, flavor compound accumulation increases total free monoterpenes from 0.6 to 4.8 mg/L and total bound monoterpenes from 2.1 to 12.5 mg/L over 42 days, with linalool accumulating from 0.25 to 1.85 mg/L free and 0.8 to 4.2 mg/L bound, representing net accumulation rates of 0.29 mg/L total monoterpenes per day during peak mid-ripening phase under high-altitude conditions at 1700 m elevation with intense solar radiation and cool nights of 12°C.
Flavor compound accumulation describes the progressive increase in concentration of flavor-active metabolites within Vitis vinifera berries during the ripening period, wherein the outputs of primary and secondary biosynthetic pathways are retained and concentrated within berry tissues to establish the quantitative flavor compound pools that determine aromatic intensity and complexity at harvest maturity over the 35-50 day post-veraison period. This process integrates the net balance between ongoing biosynthesis generating new flavor compounds and potential losses through volatilization, oxidative degradation, and enzymatic transformation, with successful accumulation requiring that synthesis rates exceed loss rates to produce the monotonic increase in total flavor compound content characteristic of normal ripening progression. The cellular mechanisms supporting flavor compound accumulation involve vacuolar sequestration of both free volatiles and glycosidic precursors through tonoplast transport systems that actively import synthesized compounds against concentration gradients, establishing intracellular reservoirs protected from degradative processes occurring in cytosolic compartments where oxidases and other catabolic enzymes maintain activity throughout ripening. The temporal dynamics of flavor compound accumulation show compound-class-specific patterns with monoterpenes typically accumulating continuously throughout ripening in aromatic varieties, norisoprenoids showing accelerated accumulation during mid-to-late ripening coincident with carotenoid degradation, and methoxypyrazines uniquely demonstrating net decline rather than accumulation as degradation exceeds residual synthesis during the post-veraison period. Cultivar genetics profoundly influence flavor compound accumulation through effects on both biosynthetic capacity determining synthesis rates and transport efficiency affecting retention, with aromatic varieties such as Muscat achieving total monoterpene accumulation of 5-15 mg/L compared to neutral varieties such as Chardonnay accumulating less than 0.5 mg/L under equivalent conditions. Environmental factors modulate flavor compound accumulation through temperature effects on enzyme activities and compound volatility, light exposure influencing biosynthetic pathway activity, and water status affecting berry size and therefore concentration effects, with moderate conditions generally favoring optimal accumulation compared to environmental extremes that may either limit synthesis or accelerate degradation. The relationship between flavor compound accumulation and wine quality reflects the fundamental importance of adequate aromatic intensity for sensory impact, with research demonstrating threshold concentrations required for perception of individual compounds and synergistic interactions among accumulated compounds that enhance overall aromatic complexity beyond the sum of individual contributions. Viticultural practices targeting flavor compound accumulation include canopy management optimizing cluster microclimate, irrigation strategies balancing vine stress with metabolic function, and harvest timing decisions capturing optimal accumulation before potential late-season degradation compromises aromatic quality.
Flavor_Compound_Accumulation is parent class under Flavor_Development representing quantitative increase in flavor metabolites. Distinguishes from Flavor_Compound_Synthesis which describes biosynthetic processes. Contains subclasses Progressive_Flavor_Build_Up, Flavor_Intensity_Increase, and Flavor_Complexity_Development. Duration 35-50 days. Key characteristic is net positive balance between synthesis and degradation producing concentration increase.
Aroma-Metabolit-Anreicherung
Aromaverbindungs-Akkumulation
Geschmacksstoff-Konzentrations-Zunahme
Volatil-Akkumulation
aroma compound build-up
flavor concentration increase
flavor metabolite accumulation
volatile accumulation
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 285-310 Robinson AL et al (2014) Origins of grape and wine aroma. American Journal of Enology and Viticulture 65: 1-30 Günata Z et al (1985) The aroma of grapes. Journal of Chromatography 331: 83-90
Flavor Compound Accumulation
In Muscat Blanc à Petits Grains berries in Frontignan, France during BBCH 83-89, flavor compound synthesis generates free monoterpenes increasing from 0.4 to 2.8 mg/L over 40 days, with free linalool reaching 1.2 mg/L, free geraniol 0.65 mg/L, and free nerol 0.35 mg/L, while concurrent glycosylation sequesters 85% of synthesized terpenoids as bound precursors, under Mediterranean conditions with mean temperature of 25°C and substantial diurnal variation.
Flavor compound synthesis encompasses the enzymatic biosynthesis of free, volatile flavor-active compounds in Vitis vinifera berries during ripening, wherein primary and secondary metabolic pathways generate the diverse array of volatiles including monoterpenoids, sesquiterpenoids, C13-norisoprenoids, methoxypyrazines, volatile thiols, esters, and lipid-derived C6 compounds that collectively constitute the characteristic flavor profile of grape varieties and contribute directly to wine sensory quality without requiring precursor transformation for flavor expression. This process involves multiple biosynthetic pathways operating in distinct cellular compartments, with the methylerythritol phosphate pathway generating monoterpene precursors in plastids followed by modification reactions producing linalool, geraniol, nerol, and citronellol in cytosolic compartments, the carotenoid cleavage pathway generating norisoprenoid volatiles including beta-damascenone and beta-ionone through dioxygenase-mediated oxidation in chloroplasts and chromoplasts, and amino acid-derived pathways producing methoxypyrazines through O-methyltransferase activity in multiple cellular locations. The temporal progression of flavor compound synthesis shows compound-class-specific patterns with methoxypyrazines reaching maximum concentrations before veraison followed by progressive degradation during ripening, monoterpenes increasing throughout the post-veraison period with accumulation continuing until harvest in aromatic varieties, and C6 compounds showing complex dynamics reflecting the balance between lipoxygenase-mediated synthesis and oxidative degradation processes. Cultivar genetics fundamentally determine flavor compound synthesis capacity through variety-specific expression of biosynthetic enzymes, with aromatic varieties such as Muscat showing 10-100 fold higher monoterpene synthase expression than neutral varieties, Sauvignon Blanc demonstrating unique methoxypyrazine and thiol precursor synthesis capacity, and Cabernet Sauvignon expressing characteristic methoxypyrazine accumulation patterns distinct from aromatic cultivars. Environmental modulation of flavor compound synthesis occurs through temperature effects on enzyme activities and compound stability, light-mediated regulation of carotenoid and derived norisoprenoid synthesis, and water stress effects on secondary metabolism activation, with these factors interacting to produce the vintage variation and terroir expression characteristic of quality wine production. The partitioning between free volatile synthesis and conjugation to precursor forms represents a critical regulatory point determining flavor compound availability at harvest, with aromatic varieties typically maintaining 5-20% of terpenoids as free volatiles while neutral varieties may show less than 5% in free form.
Flavor_Compound_Synthesis is parent class under Flavor_Development representing biosynthesis of free volatile compounds. Distinguishes from Precursor_Formation which generates conjugated forms. Contains subclasses Primary_Flavor_Formation, Secondary_Metabolite_Production, and Flavor_Pathway_Activation. Multiple pathways including MEP terpenoid, carotenoid cleavage, and amino acid-derived routes. Cultivar-specific expression determines synthesis capacity.
Aromaverbindungs-Synthese
Flavor-Volatil-Synthese
aroma compound biosynthesis
flavor volatile synthesis
flüchtige Aroma-Biosynthese
free flavor compound formation
freie Aromaverbindungs-Bildung
volatile flavor biosynthesis
Martin DM et al (2010) Functional annotation, genome organization and phylogeny of the grapevine terpene synthase gene family. BMC Plant Biology 10: 226 Dunlevy JD et al (2009) A methyltransferase essential for methoxypyrazine-derived flavour. Plant Journal 59: 404-412 Keller M (2020) The Science of Grapevines 3rd Ed pp 285-295
Flavor Compound Synthesis
In Sauvignon Blanc berries in Marlborough, New Zealand during BBCH 81-89, flavor development proceeds over 45 days with methoxypyrazine concentrations declining from 35 to 8 ng/L, volatile thiol precursors increasing from 50 to 280 µg/L, and total free volatiles accumulating to 850 µg/L at harvest, establishing the characteristic tropical and herbaceous flavor profile under cool maritime conditions averaging 18°C.
Flavor development constitutes the comprehensive ripening-phase process in Vitis vinifera berries wherein the characteristic taste and aromatic compounds that define wine sensory quality are synthesized, accumulated, and transformed during the 35-50 day post-veraison maturation period, encompassing the formation of flavor precursors, synthesis of primary flavor compounds, and production of secondary metabolites that collectively establish the cultivar-typical and terroir-expressive flavor profile at harvest maturity. This process integrates multiple biosynthetic pathways including amino acid metabolism generating flavor-active compounds and precursors, carbohydrate-derived pathways producing furanones and related sweetness-associated volatiles, lipid oxidation yielding C6 aldehydes and alcohols contributing green and herbaceous notes, and specialized secondary metabolism generating terpenoids, norisoprenoids, and phenolic derivatives that constitute the aromatic signature of individual grape varieties. The temporal progression of flavor development follows characteristic patterns with early post-veraison accumulation of primary metabolites serving as biosynthetic precursors, mid-ripening activation of secondary metabolic pathways producing variety-specific flavor compounds, and late-ripening transformations including glycosylation, esterification, and oxidative modifications that modulate flavor compound availability and sensory impact in the mature berry. Cellular localization of flavor development processes varies among compound classes, with amino acid and organic acid metabolism occurring predominantly in mesocarp parenchyma cells, terpenoid and norisoprenoid synthesis concentrated in exocarp cells particularly in aromatic varieties, and phenolic compound accumulation distributed across skin and seed tissues with distinct compositional profiles reflecting tissue-specific enzyme expression. Temperature during ripening profoundly influences flavor development through effects on enzyme activities, precursor availability, and compound stability, with moderate temperatures of 20-25°C generally favoring balanced flavor accumulation while temperature extremes either accelerate degradation of heat-labile compounds or slow biosynthetic rates below optimal levels. Cultivar genetics establish the fundamental flavor development potential through variety-specific expression of biosynthetic enzymes, with aromatic varieties such as Muscat, Gewürztraminer, and Riesling showing elevated terpenoid pathway activity compared to neutral varieties such as Chardonnay and Pinot Noir where flavor complexity derives from different compound classes. Viticultural practices influence flavor development through effects on vine balance, canopy microclimate, and berry composition, with moderate water stress, appropriate crop load, and optimized light exposure generally enhancing flavor compound accumulation compared to excessive vigor or overcropping conditions. The relationship between flavor development and other ripening processes including sugar accumulation, acid reduction, and phenolic maturation reflects the integrated nature of berry metabolism, with carbon and nitrogen partitioning among competing biosynthetic pathways determining the final balance of flavor-active compounds at harvest. Research methodologies for assessing flavor development include gas chromatography-mass spectrometry quantification of volatile compounds, HPLC analysis of non-volatile precursors, and sensory evaluation techniques that correlate chemical composition with perceived flavor attributes in grapes and wines.
Flavor_Development is parent class under Ripening representing all flavor-related biosynthetic processes. Distinguishes from Aroma_Development which focuses specifically on volatile aromatic compounds. Contains subclasses Precursor_Formation, Flavor_Compound_Synthesis, Flavor_Compound_Accumulation, and Flavor_Profile_Maturation. Duration 35-50 days post-veraison. Integrates amino acid, carbohydrate, lipid, and secondary metabolic pathways.
Beeren-Aroma-Bildung
Geschmacks-Entwicklung
Reife-Aromasynthese
berry flavor formation
flavor compound accumulation
grape flavor development
ripening flavor synthesis
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 285-310 Robinson AL et al (2014) Origins of grape and wine aroma. American Journal of Enology and Viticulture 65: 1-30 Dunlevy JD et al (2009) A methyltransferase essential for the methoxypyrazine-derived flavour of wine. Plant Journal 59: 404-412
Flavor Development
In Gewürztraminer berries in Alsace, France during BBCH 85-89, flavor intensity increase elevates free linalool from 0.4 mg/L at 1.6-fold perception threshold to 2.2 mg/L at 8.8-fold threshold over 35 days, with total odor activity value of monoterpene fraction increasing from 12 to 85 threshold units, and trained panel intensity ratings progressing from 3.2 to 7.8 on 10-point scale for floral-rose character, demonstrating the sensory impact of continued accumulation under cool continental conditions averaging 19°C.
Flavor intensity increase describes the progressive enhancement of aromatic impact and sensory potency in Vitis vinifera berries during ripening, wherein accumulating flavor compound concentrations progressively exceed perception thresholds and reach levels capable of producing pronounced aromatic impressions in harvested grapes and resulting wines, transitioning from subtle or imperceptible aromatics in unripe berries to the intense varietal character defining quality wine grapes at optimal maturity. This process reflects the relationship between absolute flavor compound concentrations and human sensory perception thresholds, with intensity increase occurring as compounds accumulate from sub-threshold levels of 10-50% of perception threshold at veraison to supra-threshold concentrations of 2-10 fold above threshold at harvest that generate clear sensory impact and varietal recognition. The sensory significance of flavor intensity increase varies among compound classes based on their respective perception thresholds, with potent odorants such as methoxypyrazines perceptible at nanogram-per-liter concentrations requiring only modest absolute accumulation for intensity impact, while less potent compounds such as certain monoterpene alcohols require milligram-per-liter accumulation to achieve equivalent sensory prominence. The measurement of flavor intensity increase integrates analytical quantification of compound concentrations with sensory evaluation methods including odor activity value calculations expressing concentration as multiples of perception threshold, trained panel assessments rating aromatic intensity on structured scales, and consumer studies correlating chemical composition with perceived quality attributes. Cultivar genetics determine the potential for flavor intensity increase through variety-specific biosynthetic capacities that establish maximum achievable compound concentrations, with aromatic varieties capable of producing intense floral, fruity, or spicy aromatics while neutral varieties develop more subtle aromatic profiles even at full physiological ripeness. Environmental and viticultural factors modulate flavor intensity increase through effects on both absolute accumulation and perception threshold modification, with berry composition factors including sugar content and pH influencing volatile release and perception in complex matrix effects that extend beyond simple concentration relationships. The optimization of flavor intensity increase represents a primary quality objective in premium wine grape production, with harvest timing decisions targeting concentration levels that maximize sensory impact while avoiding potential overripeness that may shift aromatic profiles toward less desirable dried fruit or oxidized characters.
Flavor_Intensity_Increase is subclass of Flavor_Compound_Accumulation representing sensory potency enhancement. Distinguishes from Progressive_Flavor_Build_Up which emphasizes quantity. Sibling to Progressive_Flavor_Build_Up and Flavor_Complexity_Development. Key parameter is odor activity value increase from sub-threshold to 2-10 fold supra-threshold. Relates concentration to perception threshold. Quality determinant for harvest timing.
Aroma-Intensitäts-Zunahme
Aroma-Wirkung-Steigerung
Geschmacks-Stärke-Erhöhung
aroma impact increase
aromatic potency increase
flavor strength enhancement
sensorische Intensitäts-Entwicklung
sensory intensity development
Ferreira V et al (2000) Quantitative determination of the odorants of young red wines. Journal of the Science of Food and Agriculture 80: 1659-1667 Guth H (1997) Quantitation and sensory studies of character impact odorants of different white wine varieties. Journal of Agricultural and Food Chemistry 45: 3027-3032 Keller M (2020) The Science of Grapevines 3rd Ed pp 300-310
Flavor Intensity Increase
The flavor mutation data for Chardonnay Musqué documents a gain-of-function point mutation in the VvDXS gene (1-deoxy-D-xylulose 5-phosphate synthase) on chromosome 5 — a single nucleotide substitution (c.1117A>C) producing a threonine-to-proline amino acid change (p.Asn373Thr) in the active site of this key enzyme in the methylerythritol phosphate (MEP) terpenoid precursor pathway that increases monoterpene (linalool, geraniol, nerol) production by 3-5 fold compared to standard Chardonnay, generating the characteristic muscat-floral aroma that distinguishes the Musqué clone. The same VvDXS mutation has been identified independently in muscat-aromatic sports of other normally non-muscat cultivars (Savagnin → Gewürztraminer, Chasselas → Chasselas Musqué), representing a remarkable case of convergent somatic mutation at the same nucleotide position in different cultivar lineages.
A mutation information record specific to somatic mutations affecting aroma and flavor compound biosynthesis in Vitis vinifera berries, documenting molecular changes in genes controlling terpenoid, norisoprenoid, methoxypyrazine, thiol precursor, or other flavor-active metabolite pathways that produce variant aroma profiles distinguishing specific clones or sports from the standard phenotype of their parent cultivar.
Flavor_Mutation_Data documents somatic mutations affecting the metabolic pathways that produce volatile and non-volatile flavor compounds in grape berries — a category of mutation with significant commercial consequences because aroma profile is one of the primary characteristics defining cultivar and wine identity. The best-characterized flavor mutation is the VvDXS gain-of-function mutation producing the muscat aroma phenotype, which has arisen independently in multiple cultivar lineages (Emanuelli et al. 2010), but other flavor-relevant somatic mutations include those affecting: carotenoid cleavage dioxygenase (VvCCD) genes involved in norisoprenoid (C13) aroma production (β-damascenone, β-ionone), O-methyltransferase genes controlling methoxypyrazine levels (the green pepper aroma of Cabernet Sauvignon family cultivars), and cysteine conjugate lyase genes involved in volatile thiol release (the grapefruit and passionfruit aromas of Sauvignon Blanc). The molecular characterization of flavor mutations is less advanced than color mutation research because flavor phenotypes are quantitative (influenced by environment and winemaking), chemically complex (involving dozens of compounds contributing to perceived aroma), and analytically demanding (requiring GC-MS or GC-olfactometry for compound identification and quantification), making the linkage between specific DNA mutations and flavor outcomes more difficult to establish definitively.
Aromamutations-Daten
Aromastoffweg-Mutations-Daten
Geschmacksvarianten-Mutations-Dokumentation
aroma mutation record
flavor variant mutation data
volatile compound pathway mutation
Emanuelli F et al. (2010) A candidate gene association study on muscat flavor in grapevine (Vitis vinifera L.), BMC Plant Biology 10: 241 Battilana J et al. (2009) The 1-deoxy-D-xylulose 5-phosphate synthase gene co-localizes with a major QTL affecting monoterpene content in grapevine, Theoretical and Applied Genetics 118: 653-669 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340
Flavor Mutation Data
In Moscato Giallo berries in Trentino, Italy during BBCH 81-85, flavor pathway activation induces linalool synthase gene expression 8.5-fold between pre-veraison and 14 days post-veraison, with corresponding enzyme activity increasing from 2.5 to 18.5 nmol/min/g fresh weight in exocarp tissue, enabling subsequent monoterpene accumulation that reaches 4.2 mg/L total terpenoids at harvest, with activation coinciding with ABA increase from 0.2 to 1.8 µg/g fresh weight during the same period.
Flavor pathway activation describes the developmental and environmental induction of biosynthetic pathway gene expression and enzyme activities in Vitis vinifera berries during ripening that establishes the metabolic capacity for flavor compound synthesis, encompassing the transcriptional upregulation of terpene synthases, carotenoid cleavage dioxygenases, O-methyltransferases, and related biosynthetic enzymes that convert primary metabolic precursors into the flavor-active secondary metabolites characterizing mature grape berries. This process involves signal transduction cascades triggered by developmental cues at veraison including abscisic acid accumulation, ethylene signaling, and sugar sensing, with downstream transcription factor activation inducing coordinated expression of pathway genes within the 7-14 day period following veraison onset when biosynthetic capacity is established for subsequent flavor compound accumulation. The molecular mechanisms underlying flavor pathway activation include MYB and bHLH transcription factor binding to promoter elements of terpenoid pathway genes, hormone-responsive element activation linking ABA signaling to secondary metabolism induction, and chromatin remodeling events that transition flavor biosynthesis genes from repressed to transcriptionally active states during the developmental transition at veraison. Cultivar variation in flavor pathway activation reflects genetic differences in transcription factor expression, promoter element sequences, and signal transduction pathway sensitivity that collectively determine the magnitude and timing of biosynthetic gene induction, with aromatic varieties showing 5-20 fold higher induction of monoterpene synthase genes compared to neutral varieties during the veraison transition. Environmental modulation of flavor pathway activation occurs through light-responsive elements in terpenoid pathway gene promoters that enhance expression under appropriate irradiance conditions, temperature-sensitive regulatory mechanisms that may accelerate or delay pathway induction, and stress-responsive signaling pathways that activate secondary metabolism under conditions of water deficit, pathogen challenge, or oxidative stress. The relationship between pathway activation timing and final flavor compound concentrations reflects the integration of biosynthetic capacity with substrate availability and compound stability, with early activation in the ripening season potentially maximizing accumulation time while late activation may limit total synthesis despite equivalent enzyme activities due to shortened biosynthetic windows before harvest.
Flavor_Pathway_Activation is subclass of Flavor_Compound_Synthesis representing the regulatory initiation of biosynthetic capacity. Distinguishes from Primary_Flavor_Formation and Secondary_Metabolite_Production which describe compound synthesis. Sibling to those classes. Occurs during 7-14 days post-veraison. Involves transcriptional induction of terpene synthases, CCDs, and related enzymes. Cultivar-specific activation magnitude determines aromatic potential.
Aroma-Enzym-Induktion
Aroma-Stoffwechselweg-Aktivierung
Flavor-Biosynthese-Induktion
aroma pathway upregulation
flavor biosynthesis induction
flavor enzyme induction
secondary pathway activation
sekundärer Stoffwechselweg-Aktivierung
Martin DM et al (2010) Functional annotation of the grapevine terpene synthase gene family. BMC Plant Biology 10: 226 Castellarin SD et al (2011) Transcriptional regulation of anthocyanin biosynthesis in ripening fruits of grapevine. Plant Physiology 156: 1232-1246 Keller M (2020) The Science of Grapevines 3rd Ed pp 285-295
Flavor Pathway Activation
In Muscat of Alexandria berries in Rutherglen, Australia during BBCH 85-89, flavor precursor synthesis generates glycosidically-bound monoterpenes at rates of 0.15 mg/L per day during peak activity at 25 days post-veraison, with UDP-glucose monoterpenol glucosyltransferase activity reaching 45 nmol/min/g fresh weight in exocarp tissue, producing final bound linalool concentrations of 3.2 mg/L and bound geraniol of 2.4 mg/L under mean ripening temperatures of 26°C.
Flavor precursor synthesis describes the specific enzymatic reactions in Vitis vinifera berry cells during ripening wherein free flavor compounds are conjugated with carrier molecules to form stable, non-volatile precursor compounds, involving glycosyltransferase-mediated attachment of sugar moieties to terpenols and other aglycones, and glutathione S-transferase catalyzed conjugation of cysteine or glutathione to volatile thiol precursors that will release flavor-active compounds during subsequent winemaking processes. This process encompasses the activity of UDP-glucose-dependent glycosyltransferases that transfer glucose residues to monoterpene alcohols including linalool, geraniol, nerol, and citronellol, producing beta-glucoside conjugates that accumulate in vacuolar compartments where the acidic environment and absence of hydrolytic enzymes ensure precursor stability throughout ripening and harvest handling. The enzymatic mechanisms involve recognition of hydroxyl-bearing aglycone substrates by glycosyltransferase active sites, nucleophilic attack on UDP-glucose to form enzyme-glucoside intermediates, and transfer of the glucose moiety to the acceptor molecule with release of UDP as the leaving group, with reaction rates determined by enzyme expression levels, substrate availability, and cofactor concentrations that vary developmentally during berry maturation. Thiol precursor synthesis proceeds through glutathione S-transferase mediated conjugation of reduced glutathione to alpha-beta-unsaturated carbonyl compounds including mesityl oxide and related precursor aldehydes, followed by peptidase cleavage removing glutamate and glycine residues to generate cysteine conjugates that serve as the direct substrates for yeast carbon-sulfur lyase activity during fermentation. The cellular compartmentalization of flavor precursor synthesis involves cytosolic localization of most conjugating enzymes with subsequent transport of synthesized conjugates into vacuolar storage compartments through tonoplast-localized transporters, establishing the sequestered precursor pools that represent the dominant form of potential flavor compounds in ripe berries. Developmental regulation of precursor synthesis involves transcriptional activation of glycosyltransferase and glutathione S-transferase genes during the veraison-to-harvest period, with expression profiles showing variety-specific patterns that correlate with final precursor concentrations and distinguish aromatic from neutral cultivars at the molecular level. Temperature modulation of precursor synthesis reflects the thermal sensitivity of conjugating enzyme activities, with optimal synthesis rates at moderate temperatures of 20-28°C and reduced activity at temperature extremes that may limit precursor accumulation in climatically challenging vintages.
Flavor_Precursor_Synthesis is subclass of Precursor_Formation representing the specific enzymatic conjugation reactions. Distinguishes from Amino_Acid_Accumulation which provides nitrogen-containing precursors. Sibling to Amino_Acid_Accumulation and Precursor_Pool_Development. Key enzymes are glycosyltransferases and glutathione S-transferases. Produces stable conjugates for subsequent flavor release.
Aroma-Glycosylierung
Aromavorläufer-Synthese
Glycosid-Vorläufer-Bildung
bound volatile formation
conjugate biosynthesis
flavor glycosylation
gebundene Volatil-Biosynthese
glycoside precursor synthesis
Bönisch F et al (2014) A UDP-glucose monoterpenol glucosyltransferase adds to the chemical diversity of the grapevine metabolome. Plant Physiology 165: 561-581 Tominaga T et al (1998) Identification of cysteinylated aroma precursors. Journal of Agricultural and Food Chemistry 46: 5215-5219 Keller M (2020) The Science of Grapevines 3rd Ed pp 298-302
Flavor Precursor Synthesis
In Pinot Noir berries in Côte de Nuits, Burgundy during BBCH 87-89, flavor profile maturation refines the aromatic profile over the final 18 days before harvest, with methoxypyrazine declining from 12 to 4 ng/L eliminating green character, beta-damascenone increasing from 2.8 to 4.5 µg/L enhancing fruity complexity, and trained panel ratings for varietal typicity increasing from 6.2 to 8.5 while green-herbaceous scores decline from 4.8 to 1.5 on 10-point scales, establishing the characteristic red-fruit and earthy profile under mean temperatures of 22°C.
Flavor profile maturation constitutes the terminal phase of flavor development in Vitis vinifera berries wherein the accumulated flavor compounds undergo final qualitative refinements and the overall aromatic profile achieves the balanced, harmonious character associated with optimal ripeness during the final 10-21 days before harvest, representing the culmination of biosynthetic processes and the establishment of definitive varietal and terroir expression that will carry through vinification into the finished wine. This process involves subtle transformations in flavor compound composition including shifts in terpenoid profiles through interconversion reactions, modification of norisoprenoid balances through continued carotenoid degradation, and evolution of precursor-to-free-volatile ratios that determine immediate aromatic expression versus aromatic potential requiring winemaking intervention for release. The qualitative aspects of flavor profile maturation extend beyond simple concentration changes to encompass integration of disparate aromatic elements into cohesive sensory impressions, reduction of green-unripe character through continued methoxypyrazine degradation and C6 compound metabolism, and enhancement of ripe-fruit character through late-stage accumulation of ester and norisoprenoid compounds associated with optimal maturity. Sensory evaluation of flavor profile maturation employs holistic assessment methods including overall quality ratings, balance and harmony scores, and varietal typicity judgments that integrate multiple aromatic attributes into unified quality determinations, complementing analytical measurements of individual compound concentrations. The timing of flavor profile maturation relative to sugar accumulation and acid reduction varies among cultivars and growing conditions, with certain varieties achieving optimal flavor maturity coincident with target sugar levels while others may require extended hang-time for complete flavor development potentially at the cost of elevated sugar or reduced acidity. Environmental conditions during the final ripening phase critically influence flavor profile maturation, with moderate temperatures of 20-26°C favoring continued flavor refinement while excessive heat may accelerate degradation of heat-labile compounds or shift aromatic profiles toward overripe character before optimal balance is achieved. The assessment of flavor profile maturation status represents a primary criterion for harvest timing decisions in quality-focused viticulture, with sensory evaluation of berry samples often taking precedence over analytical measurements in determining optimal picking dates for premium wine production.
Flavor_Profile_Maturation is parent class under Flavor_Development representing the terminal qualitative refinement phase. Distinguishes from Flavor_Compound_Accumulation which emphasizes quantity. Contains subclasses Mature_Flavor_Expression, Cultivar_Specific_Flavor, and Optimal_Flavor_Development. Duration 10-21 days before harvest. Emphasizes balance, harmony, and typicity rather than concentration. Primary harvest timing criterion.
Aroma-Charakter-Finalisierung
Geschmacksprofil-Vollendung
aroma profile ripening
aromatic maturation
aromatische Reifung
flavor character finalization
flavor profile completion
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 305-315 Coombe BG (1992) Research on development and ripening of the grape berry. American Journal of Enology and Viticulture 43: 101-110 Robinson AL et al (2014) Origins of grape and wine aroma. American Journal of Enology and Viticulture 65: 1-30
Flavor Profile Maturation
Fleck symptom quality assessment using indicator plant indexing evaluated the virus status of candidate mother vines for a foundation vineyard establishment program. Chip budding from candidate selections to the fleck-sensitive indicator Vitis rupestris St. George revealed fleck symptom quality in 78% of tested selections, with affected indicators showing the diagnostic translucent vein flecking appearing as lighter-colored fine veins visible against the green leaf blade when viewed with transmitted light, along with subtle leaf margin distortion and mild stunting compared to virus-free control indicators. The fleck symptom quality expression required 6-8 weeks post-grafting for clear development, with symptom intensity varying among selections reflecting differences in viral titer in the source material. Candidate selections showing fleck symptom quality on indicators were subsequently tested by PCR for GFkV presence, confirming infection in all symptom-positive materials. The fleck symptom quality-based screening eliminated 78% of candidates from consideration for the foundation program, with only the 22% testing negative for GFkV by both indicator indexing and PCR proceeding to additional testing for other viruses. The comprehensive fleck symptom quality evaluation established the foundation vineyard using only GFkV-free material as the starting point for the certified propagation program, ensuring that resulting propagation material would meet the most stringent certification standards requiring absence of this latent but ubiquitous virus.
Fleck symptom quality is a viral infection indicator quality that inheres in grapevine organisms infected by grapevine fleck virus (GFkV) and represents the characteristic disease manifestations visible primarily on sensitive indicator hosts where infection produces the diagnostic clearing or translucency of minor leaf veins (vein flecking) and localized leaf distortion, while remaining latent and symptomless in most Vitis vinifera cultivars where infection is detectable only through laboratory testing or grafting to indicator plants, with the primary significance of GFkV lying in its role as a component of mixed infections that may exacerbate symptoms of other viruses and its universal presence making it a benchmark for certification program stringency. This quality captures the diagnostic features of one of the most prevalent grapevine viruses worldwide, recognizing that GFkV infection approaches 100% incidence in many traditional wine regions where uncertified propagation material has been used historically, with the latent nature in commercial cultivars meaning that most infected vines show no obvious symptoms despite harboring the virus at high titers. The ontological significance of fleck symptom quality lies primarily in certification and clean-stock program contexts, where GFkV detection indicates either the use of uncertified material or the breakdown of virus-exclusion protocols, serving as a marker for overall viral status rather than causing economically significant direct disease impact in most commercial situations. The biological basis of fleck symptoms encompasses the maculavirus replication of GFkV in phloem tissues with accumulation causing the vein clearing visible in sensitive indicators, though the mechanism of symptom production and the basis for the cultivar-specific symptom expression remain incompletely characterized. The transmission of GFkV occurs primarily through vegetative propagation, with no confirmed vector transmission, making the virus a marker for propagation history where presence indicates derivation from infected source material. The role of GFkV in mixed infections has been documented to increase symptom severity when present together with other viruses including GFLV and rupestris stem pitting-associated virus, though the mechanism of this synergism is not fully understood.
Fleck_Symptom_Quality is_a Viral_Infection_Indicator_Quality, representing the symptoms of grapevine fleck virus (GFkV) infection visible on sensitive indicator hosts. Latent (symptomless) in most V. vinifera cultivars despite high viral titers. The vein flecking on indicator plants (V. rupestris St. George) provides diagnostic biological indexing. Very high prevalence (~100% in traditional regions) makes detection a marker for propagation history. No vector transmission confirmed spread only through propagation. Role as synergist in mixed infections may increase severity of other viral diseases. Primary significance in certification programs rather than direct disease impact.
Aderaufhellung
Fleck-Symptome
GFkV symptoms
GFkV-Symptome
Rebenfleck-Virose
fleck disease symptoms
grapevine fleck symptoms
vein fleck indicators
Martelli GP (2014) Directory of virus and virus-like diseases of the grapevine and their agents. Journal of Plant Pathology 96(1 Supplement):1-136 Sabanadzovic S et al. (2017) Grapevine fleck and similar viruses. In: Meng B et al. (eds) Grapevine Viruses: Molecular Biology, Diagnostics and Management. Springer, pp. 331-349 Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests, 2nd Edition. APS Press
Fleck Symptom Quality
Evaluation of Fleck_Tolerance_Disposition at ENTAV-INRA Grau du Roi compared responses to GFkV across V. vinifera cultivars and Vitis rootstock genotypes following graft inoculation. V. vinifera cultivars (n=30) uniformly demonstrated Fleck_Tolerance_Disposition, with 100% infection rates as confirmed by RT-PCR but complete absence of symptoms and no detectable impacts on growth rate, yield, or fruit quality parameters over 5-year evaluation. The sensitive indicator V. rupestris 'St. George' developed characteristic symptoms within 60 days of graft inoculation: chlorotic flecking on third-fourth leaves expanding basipetally, translucent vein appearance, and 25% reduction in shoot elongation. Rootstock genotypes with V. rupestris parentage showed variable responses, with 'SO4' (V. berlandieri × V. riparia) remaining asymptomatic while 'Schwarzmann' (V. riparia × V. rupestris) exhibited mild transient flecking. Importantly, when GFkV-infected vines were co-infected with GLRaV-3, leafroll symptom severity increased 40% compared to GLRaV-3 single infection, demonstrating synergistic interactions that justify pursuing virus-free status even for generally tolerated pathogens. These findings confirm that V. vinifera cultivars inherently possess Fleck_Tolerance_Disposition but that rootstock sensitivity and mixed infection dynamics complicate management decisions.
A Fleck_Tolerance_Disposition is a Viral_Resistance_Disposition that inheres in a Vitis plant and is realized through the capacity to maintain normal growth, productivity, and vine health despite infection by Grapevine fleck virus (GFkV), a maculavirus that is symptomless in most V. vinifera cultivars but causes disease in sensitive Vitis species and rootstock hybrids. This disposition represents tolerance to a virus that is nearly ubiquitous in grapevine germplasm worldwide, with infection rates exceeding 80% in many surveyed populations, yet causes no apparent harm in the vast majority of commercial V. vinifera cultivars. GFkV, a member of the family Tymoviridae, has isometric particles containing a positive-sense RNA genome and is transmitted through vegetative propagation but has no known insect vector for field spread. The virus induces characteristic symptoms only in sensitive indicator hosts including V. rupestris (particularly clone 'St. George'), where it causes chlorotic flecking, leaf distortion, and vein clearing, hence its use in biological indexing. The near-universal tolerance of V. vinifera to GFkV represents an unusual case where most cultivated material inherently possesses the Fleck_Tolerance_Disposition, in contrast to diseases like leafroll where tolerance is the exception. However, certain rootstock hybrids with V. rupestris parentage may exhibit sensitivity, and GFkV presence can exacerbate symptoms of other viral infections in mixed infections, making virus-free status still desirable for high-quality planting material even for this generally benign pathogen.
Fleck_Tolerance_Disposition is_a Viral_Resistance_Disposition. Fleck disease is unusual among grapevine viral diseases in that commercial V. vinifera cultivars are inherently tolerant, with sensitivity restricted to certain wild species (V. rupestris) and some rootstock hybrids. The term "tolerance" rather than "resistance" is used because V. vinifera supports viral replication without symptoms rather than restricting infection. GFkV has no known vector for field transmission, spreading only through propagation. Despite its generally benign nature, GFkV-free status is preferred for mother plant material due to potential synergistic interactions in mixed viral infections.
Fleck-Virus-Toleranz
GFkV tolerance
GFkV-Toleranz
Marmorierungs-Toleranz-Disposition
fleck virus asymptomatic carriage
maculavirus tolerance
Sabanadzovic S et al. (2017) Grapevine fleck virus, Grapevine Viruses: Molecular Biology, Diagnostics and Management, Springer Martelli GP (2017) An overview on grapevine viruses, viroids, and the diseases they cause, Grapevine Viruses: Molecular Biology, Diagnostics and Management, Springer Fuchs M et al. (2007) Serological and molecular characterization of Grapevine fleck virus isolates from different grapevine-growing regions, Archives of Virology 152:1191-1197
Fleck Tolerance Disposition
The flesh color description of Alicante Bouschet documents intensely colored dark red flesh (OIV 230:2) — the mesocarp tissue is deeply pigmented with anthocyanin throughout, producing dark red juice immediately upon crushing without any skin contact maceration, with free-run juice color density exceeding that of many red wines made by conventional skin-contact fermentation of non-teinturier cultivars. This teinturier phenotype results from a gain-of-function mutation in the VvMYBA gene regulatory region that extends anthocyanin pathway activation from the skin (where it occurs in all red cultivars) into the flesh tissue (where it is normally suppressed), producing the commercially valuable ability to generate deeply colored wines from direct-press juice. The colored flesh of Alicante Bouschet (total flesh anthocyanin 200-600 mg/kg) contrasts with the colorless flesh of Cabernet Sauvignon (flesh anthocyanin <5 mg/kg despite the intensely pigmented skin), Pinot Noir, Merlot, and virtually all other standard red wine cultivars.
A berry color description component that records the color of the berry flesh (mesocarp) revealed when the berry is cut transversely at the equatorial plane, scored according to OIV descriptor 230 as not colored (state 1, colorless to pale green translucent flesh, the condition of virtually all standard V. vinifera cultivars) or colored (state 2, red-pigmented flesh indicating anthocyanin biosynthesis in mesocarp tissue, the teinturier condition found in a small number of cultivars), with flesh color being the diagnostic criterion for identifying teinturier cultivars whose colored juice enables direct-press red winemaking without skin maceration.
Flesh_Color_Description documents a binary character (colored versus not colored) that identifies a small but commercially significant group of cultivars — the teinturiers (from French teint, dyed) — whose anthocyanin production extends beyond the normal skin-specific expression into the flesh tissue. In standard V. vinifera cultivars, the VvMYBA regulatory genes are expressed exclusively in the skin hypodermis (2-10 cell layers beneath the epidermis), with the underlying mesocarp (flesh) cells lacking VvMYBA expression and consequently producing no anthocyanin — this tissue-specific expression pattern means that the juice of even deeply pigmented cultivars like Cabernet Sauvignon is colorless when gently pressed without skin contact: the teinturier condition, in which VvMYBA expression extends into the mesocarp, is found in only a handful of V. vinifera cultivars, most notably Alicante Bouschet (a 19th-century cross of Petit Bouschet × Grenache, where Petit Bouschet inherited the teinturier trait from Teinturier du Cher), Saperavi (a Georgian cultivar with independently evolved flesh pigmentation), and a few others. Teinturier cultivars have historically been used as blending components to intensify the color of wines from weakly pigmented cultivars (a practice now restricted or prohibited in many appellations), but are increasingly valued in their own right for deeply colored single-varietal wines and as genetic donors of the teinturier trait in modern breeding programs seeking cultivars that produce highly colored wines with minimal skin maceration.
Beerenfleisch-Färbung
Fruchtfleischfarbe-Beschreibung
Mesokarp-Pigmentierung
flesh pigmentation assessment
mesocarp color record
pulp coloration description
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Falginella L et al. (2010) Expansion and subfunctionalisation of flavonoid 3',5'-hydroxylases in the grapevine lineage, BMC Genomics 11: 562 Boursiquot JM et al. (2009) Parentage of Merlot and related winegrape cultivars of southwestern France, Australian Journal of Grape and Wine Research 15: 144-150
Flesh Color Description
The flesh consistency of Autumn Royal documents very firm, crunchy flesh (OIV 235:4) — the berry resists substantial finger pressure before rupturing, and when bitten produces a clean, audible crunch as the flesh fractures along cell walls rather than deforming plastically, with the flesh fragments maintaining their structure during chewing rather than dissolving into juice. This crisp, shattering texture is the result of intensive table grape breeding selection for the firm-fleshed phenotype driven by consumer preference and commercial handling requirements (firm berries resist transit damage) — the flesh cells in Autumn Royal maintain high turgor pressure and thick, rigid cell walls with elevated cellulose content that produce brittle fracture behavior rather than the plastic deformation of thin-walled, high-turgor soft-fleshed cells. The very firm flesh of Autumn Royal contrasts with the soft, melting flesh of Chasselas (OIV 235:1, flesh deforms under gentle pressure and dissolves into juice during chewing) and the moderately firm flesh of Italia (OIV 235:2-3, intermediate between the soft wine grape and crisp modern table grape textures).
A berry flesh description component that records the firmness and textural behavior of the berry flesh when compressed and bitten, scored according to OIV descriptor 235 on a scale from soft (state 1, flesh deforms easily under finger pressure and releases juice freely) through slightly firm (state 2, flesh resists light compression but yields under moderate pressure), firm (state 3, flesh maintains shape under moderate pressure and ruptures crisply), to very firm (state 4, flesh resists strong compression and shatters with a crunchy snap), with flesh consistency being the primary textural quality attribute for table grape evaluation and a determinant of juice extraction efficiency in winemaking.
Flesh_Consistency documents the mechanical property that is the single most important quality criterion in modern table grape breeding and marketing — consumer preference surveys consistently rank berry firmness and crunchiness as the top textural attributes, and postharvest quality assessment protocols use firmness (measured instrumentally as penetration force or compression resistance) as the primary indicator of freshness and shelf life. The cellular basis of flesh firmness involves three components — cell turgor (the internal hydrostatic pressure that inflates each cell, maintaining firmness when high), cell wall rigidity (the structural resistance of the wall to deformation, determined by cellulose, hemicellulose, and pectin composition), and intercellular adhesion (the strength of the middle lamella bonds between adjacent cells, determining whether stress causes cell rupture or cell separation): firm-fleshed cultivars combine high turgor with rigid walls and strong adhesion, producing the crisp fracture behavior in which the flesh breaks cleanly, while soft-fleshed cultivars have lower turgor or weaker adhesion, producing the melting, juice-releasing behavior characteristic of traditional wine grape and eating grape cultivars. The flesh consistency also affects winemaking — soft-fleshed cultivars release juice more readily during crushing and pressing (higher free-run yield), while firm-fleshed cultivars require more aggressive pressing to extract equivalent juice volumes, potentially increasing phenolic extraction from skin and seed contact.
Beerenfleisch-Festigkeit
Fruchtfleisch-Konsistenz
Pulpa-Textur-Bewertung
berry flesh rigidity
flesh firmness record
pulp texture assessment
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Segade SR et al. (2011) Phenolic ripeness assessment of grape skin by texture analysis, Journal of Food Composition and Analysis 24: 644-649
Flesh Consistency
Flesh consistency quality evaluation in Crimson Seedless identifies the physiological disorder 'berry collapse' through non-uniform flesh consistency where affected berries display localized softening at the stylar end (penetration force below 0.5 N) while the shoulder region maintains normal firmness (force above 1.5 N), creating the characteristic inconsistent texture that renders fruit unmarketable vineyard management protocols targeting consistent flesh quality include moderate crop loads, gradual irrigation deficit imposition avoiding sudden water stress, and calcium applications to maintain cell wall integrity and middle lamella cohesion throughout the berry volume.
Flesh_Consistency_Quality represents the internal texture quality characterizing the uniformity, cohesion, and structural integrity of grape berry flesh throughout its volume. While Pulp_Texture_Quality addresses the mechanical properties of mesocarp tissue, Flesh_Consistency_Quality captures the homogeneity of these properties and the overall structural organization of the flesh. Consistent flesh displays uniform texture properties throughout the berry without localized soft spots, hollow regions, or textural discontinuities, while inconsistent flesh may show zones of different firmness, separation between tissue regions, or breakdown around seeds and vascular bundles. Flesh consistency depends on uniform cellular development throughout the mesocarp, consistent ripening progression across the berry volume, and absence of damage or disorder that creates localized texture changes. Water deficit during ripening can create uneven flesh consistency as peripheral cells dehydrate while interior cells maintain turgor. Certain physiological disorders including berry shrivel and sunburn produce characteristic consistency defects with damaged peripheral zones surrounding relatively normal interior tissue. In some cultivars, the flesh separates readily from the skin (slip-skin character) or from seeds, creating inherent consistency discontinuities that may be desirable (easy seed removal) or undesirable (poor structural integrity). Flesh consistency assessment employs careful sectioning and visual examination of cut surfaces, palpation detecting localized soft zones, and instrumental analysis at multiple points within individual berries to quantify spatial variation in mechanical properties.
Flesh_Consistency_Quality complements Pulp_Texture_Quality by addressing spatial uniformity and structural organization rather than absolute mechanical properties. Consistency assessment requires sampling multiple locations within berries or examining cross-sections to detect heterogeneity. Perfect consistency (identical properties throughout) is an ideal practical assessment evaluates whether variation falls within acceptable limits. The distinction between consistency (uniformity) and firmness (absolute resistance) is important flesh can be uniformly soft or inconsistently firm. Physiological disorders and handling damage often manifest as consistency defects before affecting overall texture measurements.
Fleischkonsistenz
Fruchtfleischgleichmäßigkeit
Gewebehomogenität
flesh integrity quality
flesh uniformity
innere Beschaffenheit
internal consistency
tissue homogeneity
Krasnow MN et al. (2008) Evidence for Substantial Maintenance of Membrane Integrity and Cell Viability in Normally Developing Grape (Vitis vinifera L.) Berries Throughout Development. Journal of Experimental Botany 59(4): 849-859 Rolle L et al. (2015) Berry Density and Size as Factors Related to the Physicochemical Characteristics of Muscat Hamburg Table Grapes (Vitis vinifera L.). Food Chemistry 173: 105-113
Flesh Consistency Quality
The flesh flavor of Gewürztraminer documents an intensely aromatic muscat-type flavor (OIV 236:2) dominated by linalool (150-400 μg/L free terpene in juice, 5-10× threshold concentration), geraniol, and rose oxide monoterpenes that produce the characteristically exotic, lychee-rose-spice aroma that is one of the most distinctive and immediately recognizable flavor profiles in the entire V. vinifera cultivar spectrum. The aromatic intensity of Gewürztraminer is controlled by a gain-of-function mutation in the VvDXS gene (1-deoxy-D-xylulose 5-phosphate synthase, the rate-limiting enzyme of the MEP terpenoid biosynthesis pathway) — the same c.1117A>C mutation independently present in all aromatic V. vinifera cultivars including Muscat à Petits Grains and its relatives (Emanuelli et al. 2010). This muscat flavor contrasts with the neutral flavor of Chardonnay (OIV 236:1, no distinctive terpene aroma, producing wines whose character derives from winemaking technique rather than varietal aroma), the herbaceous flavor of Sauvignon Blanc (OIV 236:4, 3-isobutyl-2-methoxypyrazine at 10-40 ng/L contributing the distinctive green capsicum character), and the foxy flavor of Concord (OIV 236:3, methyl anthranilate at 2-5 mg/L producing the characteristic American grape candy aroma).
A berry flesh description component that records the aromatic flavor profile of the berry flesh perceived during chewing at harvest maturity, scored according to OIV descriptor 236 as neutral/non-aromatic (state 1, no distinctive aroma beyond generic grape sweetness), muscat/floral (state 2, characteristic monoterpene-derived floral aroma of linalool, geraniol, and nerol), foxy/labrusca (state 3, methyl anthranilate-dominated aroma characteristic of American Vitis species), herbaceous (state 4, methoxypyrazine-derived green bell pepper aroma), or other distinctive flavor (state 5), with flesh flavor providing one of the most immediately perceptible cultivar-diagnostic characters and a primary determinant of wine aromatic style.
Flesh_Flavor documents the aromatic character that is arguably the most commercially significant cultivar attribute for wine identity — the concept of varietal character (the recognizable aroma profile associated with a specific cultivar's wines) is primarily determined by the flesh flavor compounds that are extracted into the must during crushing and fermentation. The OIV classification into five flavor categories captures the major aromatic classes but vastly simplifies the true diversity of flesh flavor chemistry — the neutral category alone encompasses cultivars with profoundly different wine aromatic profiles (e.g., Chardonnay, Pinot Noir, Tempranillo, Nebbiolo) whose wines are distinguished by subtle differences in norisoprenoid, ester, thiol, and other volatile compound profiles not perceptible in the fresh berry tasting that the OIV descriptor assesses: the muscat category similarly encompasses the full spectrum of terpene-rich cultivars from the intensely aromatic Muscat à Petits Grains (the archetype, with total free monoterpenes of 3-6 mg/L) through the moderately aromatic Gewürztraminer and Torrontés to the subtly aromatic Viognier and Albariño. The herbaceous flavor class, associated with methoxypyrazine accumulation in the flesh and skin, is cultivar-specific (high in Sauvignon Blanc, Cabernet Sauvignon, Cabernet Franc, Carmenère) and environmentally modulated (reduced by sun exposure and later harvest), providing a flavor character that can be either a defining varietal attribute (Sauvignon Blanc) or a defect (over-herbaceous Cabernet Sauvignon from cool, shaded conditions).
Beeren-Aromaprofil
Fruchtaroma-Charakterisierung
Fruchtfleisch-Geschmack
berry aroma profile
flesh taste description
fruit flavor characterization
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Emanuelli F et al. (2010) A candidate gene association study on muscat flavor in grapevine, BMC Plant Biology 10: 241 Dunlevy JD et al. (2009) A methyltransferase essential for the methoxypyrazine-derived flavour of wine, Plant Journal 59: 404-412
Flesh Flavor
A flexible green one-year cane on Vitis vinifera cv. Chardonnay assessed during early November canopy evaluation, demonstrating ability to bend through 180-degree arc without structural resistance or cracking, returning to approximately original position when released, showing green color throughout length with no brown woody appearance, mechanical testing revealing essentially zero resistance to bending (elastic modulus 5 GPa for fully lignified canes), representing immature developmental status unsuitable for retention as fruiting wood, requiring complete removal during dormant pruning and triggering management assessment to identify causes of incomplete ripening (late vigor flush following September rains, excessive crop load depleting reserves, or insufficient thermal time in cool growing season).
A green one-year cane characterized by high flexibility and elasticity, readily bending without breaking or showing structural resistance, capable of significant deformation and return to original position, possessing mechanical properties dominated by turgor pressure and primary cell wall elasticity rather than lignified secondary walls, and representing developmental status where structural support derives from internal hydraulic pressure rather than rigid woody skeleton. Flexible green one-year canes demonstrate the biomechanical properties of unlignified plant tissues, suitable for active growing shoots that must bend in wind and accommodate trellising but unsuitable for mature fruiting wood requiring structural stability to support crop loads, making flexibility assessment a useful quick diagnostic for maturation status with flexible green appearance in late autumn indicating problematic incomplete lignification requiring pruning removal.
Flexible green one-year canes provide simple mechanical diagnostic for wood maturity, with flexibility assessment offering quick field evaluation without requiring destructive sampling or laboratory analysis. The test involves attempting to bend cane sections - flexible green canes bend easily forming tight curves or loops, while properly lignified canes show stiff resistance and crack or snap if forced beyond moderate curvature. This flexibility difference reflects fundamental structural changes during lignification as extensible primary cell walls are reinforced with rigid lignin polymers creating composite materials with dramatically increased stiffness. The persistence of high flexibility into late autumn unambiguously indicates incomplete maturation, while development of stiffness and brittleness signals progression toward appropriate woody character for fruiting wood retention.
bendable unlignified cane
biegsame grüne Rute
elastic green cane
flexible unverholzte Rute
pliable herbaceous cane
Based on Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 245-267 Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Niklas (1992) "Plant Biomechanics= An Engineering Approach to Plant Form and Function" University of Chicago Press, pp. 337-389.
Flexible Green One Year Cane
Flinty aroma evaluation comparing Chablis Premier Cru with and without malolactic fermentation reveals flinty intensity of 7.2 versus 4.5 (10-point scale), with the higher flinty expression in non-MLF wine attributed to benzenemethanethiol preservation at 1.2 ng/L versus 0.4 ng/L in MLF wine where oxidative handling during secondary fermentation reduces volatile sulfur compounds, demonstrating that winemaking decisively influences flinty character even on identical Kimmeridgian terroir, while both expressions rate higher than equivalent Mâconnais Chardonnay (2.8) where flinty character is not part of regional style expectations.
Flinty aroma quality is a mineral aroma quality that represents olfactory perception specifically evoking struck flint (silex) or gun flint-associated scents, characterizing the distinctive sharp, smoky-mineral aromatic impression particularly associated with certain Chardonnay and Sauvignon Blanc wines from specific terroirs where flint character is considered a marker of quality and site expression. Within the BFO framework, this quality specifically addresses the flinty subcategory of mineral aroma, distinguished by its reference to the specific struck-flint smell and its strong association with particular appellations (Chablis, Pouilly-Fumé) where flinty character defines regional typicity. The ontological significance lies in flinty aroma serving as the archetypal minerality descriptor for certain prestigious white wine regions, where gun flint or pierre à fusil character creates the impression of chalky, flinty terroir expression that distinguishes site-specific wines from generic examples and justifies premium pricing based on perceived terroir authenticity. The compositional basis for flinty perception has been investigated extensively, with proposed contributors including benzenemethanethiol (gun flint, smoky character with detection threshold of 0.3 ng/L), certain volatile sulfur compounds formed during reductive winemaking (dimethyl sulfide, methanethiol), and compounds formed through sur lie aging, though the exact chemistry remains debated and flinty character shows stronger correlation with winemaking reduction than with vineyard geology. The winemaking influence on flinty character is substantial, with reductive handling (minimal oxygen exposure, sur lie aging, no malolactic fermentation) enhancing flinty perception regardless of soil type, suggesting that production method contributes more to flinty character than direct geological influence from flint-containing soils. The regional expectations for flinty character concentrate in Chablis (where Kimmeridgian limestone and oyster shell fossils create the terroir narrative) and Pouilly-Fumé (named for the smoky-flinty character), with flinty notes valued as typicity markers in these regions while the same character might indicate reduction fault in regions without flinty terroir reputation. Understanding flinty aroma quality enables regional typicity assessment for flint-associated appellations, reductive winemaking evaluation based on flinty expression, and quality judgment incorporating flinty character as valued complexity in appropriate contexts.
Flinty_Aroma_Quality addresses the specifically struck-flint subcategory of mineral aroma, strongly associated with Chablis and Pouilly-Fumé regional typicity. This quality shows clearer chemical associations (benzenemethanethiol, volatile sulfur compounds) than general minerality, though the connection to actual flint in soil remains unestablished. Reductive winemaking influence on flinty character is well-documented, creating the apparent paradox that flinty terroir expression depends substantially on production method.
Feuersteinaroma
Flintcharakter
Schießpulvernote
Silexduft
gun flint aroma
pierre à fusil notes
silex character
struck flint scent
Tominaga T et al. (2003) Identification of new volatile thiols in the aroma of Vitis vinifera L. var. Sauvignon blanc wines. Flavour and Fragrance Journal 18(2): 97-103 Blanchard L et al. (2004) Reactivity of 3-mercaptohexanol in red wine. American Journal of Enology and Viticulture 55(2): 115-120
Flinty Aroma Quality
Malbec vines in Mendoza's traditional Maipú district bearing a flood-irrigated role receive water diverted from the Río Mendoza through an acequia canal network dating to colonial-era infrastructure, with irrigation turns allocated every 14-21 days during summer delivering 80-120 mm per application through inter-row furrows, consuming 6-10 ML/ha/season at application efficiencies of 40-60% compared to 85-95% for drip systems, supporting yields of 15-25 t/ha on deep alluvial soils.
An irrigated role that inheres in a grapevine receiving supplemental water through gravity-driven surface irrigation methods that distribute water across the vineyard floor via furrows, basin flooding, or border strips, representing the oldest and least water-efficient form of viticultural irrigation.
Flood_Irrigated_Role describes the traditional irrigation method still practiced in many established viticultural regions with historical canal infrastructure, particularly in Argentina, Chile, parts of Australia, and Central Asian wine regions. While substantially less water-efficient than drip or sprinkler systems, flood irrigation can provide ancillary benefits including groundwater recharge, salt leaching in saline soils, and maintenance of historically significant communal water distribution networks: however, increasing water scarcity and water pricing reforms are driving progressive conversion from flood to drip systems in most wine regions where flood irrigation persists.
Furchenbewässerungs-Produktionsrolle
flood-irrigated vine role
flutbewässerte Rolle
furrow-irrigated role
surface-irrigated production role
Überflutungsbewässerungs-Rolle
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 499-502 Williams LE and Matthews MA (1990) Grapevine, in Stewart BA and Nielsen DR (eds) Irrigation of Agricultural Crops, ASA Monograph 30 pp: 1019-1055 Deis L et al. (2015) Water and radiation use efficiency in grapevine cultivars in Mendoza, Revista de la Facultad de Ciencias Agrarias 47: 83-97
Flood Irrigated Role
Floral aroma profiling of Alsace Gewürztraminer identifies the varietal signature comprising intense rose (8.5 on 10-point scale) from cis-rose oxide at 85 μg/L, lychee-floral character (7.8) from the same compound's sensory interaction with geraniol, and subtle violet undertones (4.2) from β-ionone, with the rose dominance distinguishing Gewürztraminer from Muscat where orange blossom and jasmine notes predominate, demonstrating how floral subcategory balance creates varietal identity within the broader floral character category.
Floral aroma quality is an aroma character quality that represents olfactory perception evoking flower-associated scents including blossom, perfume, and petal notes, characterizing the delicate, often fragrant aromatic impression that contributes elegance and complexity to wine while serving as a key varietal marker for aromatic grape varieties. Within the BFO framework, this quality addresses a major aroma category distinguished by its association with perceived elegance, femininity, and aromatic intensity, recognizing that floral character derives from volatile compounds shared between wines and actual flowers through common biosynthetic pathways. The ontological significance lies in floral aroma serving as the defining varietal marker for several commercially important aromatic varieties, most notably Gewürztraminer (rose, lychee-floral), Muscat (orange blossom, jasmine), Torrontés (white flowers), and Viognier (violet, honeysuckle), where floral intensity and specific floral note identity constitute the varietal fingerprint. The compositional basis for floral aroma includes monoterpenes (linalool providing lavender and floral lift, geraniol contributing rose character, nerol adding orange blossom), rose oxides (creating the distinctive rose-lychee note in Gewürztraminer), β-ionone (violet aroma with extremely low threshold), and certain esters enhancing overall floral impression. The developmental aspects of floral aroma show grape-derived floral compounds (terpenes) present as glycosidically-bound precursors released during fermentation and aging, with some varieties showing more pronounced floral character with bottle age as slow hydrolysis continues releasing aromatic terpenes. The distinction among floral subcategories (white flowers, rose, violet) reflects both perceptual groupings and underlying compositional differences, with different varieties showing characteristic profiles across these subcategories that enable variety and quality identification. Understanding floral aroma quality enables varietal authentication based on floral profile, quality assessment where appropriate floral expression indicates varietal typicity, and style prediction where floral intensity distinguishes aromatic from neutral wine styles.
Floral_Aroma_Quality serves as the parent class for floral subcategories (White_Flower, Rose, Violet), representing a major aroma character category particularly important for aromatic variety evaluation. This quality is strongly associated with monoterpene compounds that provide both direct compositional correspondence to actual flower volatiles and perceptual similarity through shared receptor activation. The class distinguishes aromatic from neutral wine styles and provides key varietal markers.
Blumenaroma
Blütenduft
blossom character
blumiger Charakter
floral notes
florale Note
flower aroma
perfumed scent
Mateo JJ, Jiménez M (2000) Monoterpenes in grape juice and wines. Journal of Chromatography A 881(1-2): 557-567 Ribéreau-Gayon P et al. (1975) Aroma of Muscat grape varieties. Journal of Agricultural and Food Chemistry 23(6): 1042-1047
Floral Aroma Quality
Cabernet Sauvignon in Bordeaux vineyards at BBCH 65 (full bloom, mid-June) exhibits hermaphrodite flowers of 3-3.5 mm diameter with erect stamens extending 1-2 mm beyond the pistil, producing abundant viable pollen and achieving 45-55% fruit set under favorable conditions (warm, dry weather during bloom). Gewürztraminer flowers show functionally female morphology with strongly reflex stamens curved downward below the pistil, relying on cross-pollination from co-planted pollinizers and typically achieving only 25-35% fruit set. Pinot Noir flowers exhibit temperature-sensitive sex expression, with cool bloom conditions (15-18°C) inducing partial stamen reflexing and reduced pollen viability, decreasing fruit set from typical 48-55% in warm blooms to 30-40% in cool blooms, contributing to the cultivar's reputation for erratic yields.
A flower is the reproductive structure of Vitis vinifera comprising a complete set of floral organs (calyx, corolla, androecium, gynoecium) arranged in concentric whorls on a receptacle, functioning as the site of sexual reproduction through pollination and fertilization, with successful flower development determining fruit set and ultimate berry production. The grapevine flower is a perfect (bisexual) or functionally unisexual structure typically 2-4 mm in diameter at anthesis, characterized by a distinctive calyptra (fused petals forming a cap) that is shed at flower opening rather than the individual petal separation seen in most angiosperms. Flower structure comprises from outside to inside= (1) a minute calyx (five fused sepals) forming a shallow cup at the receptacle (2) five petals fused at their tips forming the calyptra that detaches as a unit during anthesis (3) five stamens positioned opposite the petals (unusual among angiosperms where stamens typically alternate with petals), each consisting of filament and anther containing pollen (4) a superior gynoecium of two fused carpels forming a two-locular ovary with typically two ovules per locule, a short style, and a wet stigma receptive to pollen. The flower is borne on a pedicel 2-8 mm long that connects to the inflorescence axis (rachis or branches) at a receptacle bearing a nectary disk surrounding the ovary base. Grapevine flowers exhibit three functional sex types despite morphological bisexuality= hermaphrodite flowers with fully functional stamens and pistil, functionally female flowers with reflex stamens or non-viable pollen but functional pistil, and functionally male flowers with functional stamens but non-receptive or aborted pistil. Flower development proceeds through distinct stages defined in the BBCH scale= BBCH 51-54 (inflorescence emergence and separation), BBCH 55-59 (individual flower buds visible and separating), BBCH 60 (first calyptra lifting), BBCH 61-65 (beginning through full bloom with progressive calyptra shedding), BBCH 67-69 (end of flowering with calyptra fallen and petal remnants drying). Anthesis (flower opening) occurs when the calyptra detaches from the receptacle base and is lifted off by the elongating stamens, typically occurring over 7-14 days for a complete inflorescence, with individual flowers opening over 6-24 hours depending on temperature (optimum 20-30°C). Pollination in cultivated Vitis vinifera is primarily anemophilous (wind-pollinated) with some secondary entomophilous (insect-pollinated) contribution, with pollen released from anthers through longitudinal dehiscence slits 1-2 hours after calyptra shedding and remaining viable for 24-72 hours depending on humidity (optimal at 60-80% RH). Stigma receptivity typically persists for 3-6 days post-anthesis under favorable conditions. Successful pollination and pollen tube growth through the style leads to fertilization of ovules typically 48-96 hours post-pollination at 20-25°C, with subsequent fruit set (transition from flower to developing berry) visible as ovary enlargement beginning 5-10 days post-fertilization. Flower-to-fruit conversion efficiency (fruit set percentage) typically ranges from 30-60% in hermaphrodite cultivars under optimal conditions, with 40-70% of flowers typically abscising as unfertilized or poorly fertilized structures during and after bloom.
Flower in Vitis represents the fundamental reproductive unit and should be distinguished from Inflorescence (the complete flower-bearing structure), Flower_Cluster (the aggregate of flowers), and Berry (the post-fertilization fruit structure). The grapevine flower exhibits several unusual botanical features= calyptra (fused petal cap) rather than separate petals, stamens opposite rather than alternate to petals, wet stigma unusual among wind-pollinated plants, and diversity of functional sex types despite morphological similarity. Flower structure, particularly sex expression and fruit set capability, is among the most economically critical traits in viticulture, as fruit set percentage directly determines yield potential and can vary 2-3 fold between cultivars and environmental conditions. Sub-classification into Perfect_Flower with hermaphrodite versus functionally unisexual subtypes reflects the critical functional diversity underlying reproductive success and crop production in different cultivar types.
Blüte
Rebblüte
Weinblüte
grapevine flower
vine flower
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 98-108 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 91-106 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 185-198.
Flower
Cabernet Sauvignon in Napa Valley shows flower bud separation during late May Year N+1 at BBCH 57 over 1-2 days, with tightly packed 3.2-3.6 mm flower buds beginning to separate as pedicels elongate from 0.6-0.8 mm to 1.2-1.8 mm creating initial gaps between buds under warm climate conditions of 19-24°C temperature, approaching optimal flowering conditions typical of this late-season cultivar initiating pre-anthesis flower individuation
Flower bud separation represents the initial individualization phase where tightly clustered flower buds begin differentiating into distinguishable units through beginning pedicel elongation and bud expansion, marking transition from compact undifferentiated flower masses to recognizable individual flower structures. This critical initiation stage occurs during late May-early June in temperate regions Year N+1 as part of individual flower separation process, typically requiring 1-2 days as initial pedicel growth and bud enlargement create perceptible flower individuation. Separation initiation shows flower buds loosening from extremely tight packing where buds physically contact neighbors creating compressed undifferentiated appearance, toward beginning separation with small gaps appearing between adjacent buds as pedicels start extending. Initial pedicel elongation during bud separation extends flower attachment stalks from very short 0.5-1.0 mm initial length to 1.0-2.0 mm as first separation increment, with elongation creating minimal but perceptible spatial projection beginning to isolate flowers. Flower bud enlargement from 3.0-3.5 mm to 3.5-4.0 mm diameter contributes to separation as expanding buds require more space, with expansion resulting from continued flower organ growth within enclosing calyptra approaching final pre-anthesis dimensions. Bud surface characteristics during separation show smooth green calyptra tightly enclosing flower organs with seam between fused petal tips potentially becoming slightly more apparent as cap loosening begins, though cap remains firmly attached during separation with detachment occurring during subsequent anthesis. Spatial positioning of separating buds shows transition from chaotic compressed arrangement with random bud orientations to beginning organized array with buds showing more consistent upward or lateral orientation as pedicels establish positioning. Flower organ status within separating buds shows continued maturation with anthers completing pollen development, pistils achieving final size with ovary expansion and stigma development, and nectaries becoming functional, all occurring while enclosed within protective calyptra. Mechanical properties of separating flower structures show delicate fragile nature with thin-walled tissues easily damaged by physical contact, rain impact, or pest feeding, requiring gentle handling and protective conditions. Vascular function in beginning-to-elongate pedicels shows developing transport capacity with xylem vessels differentiating to enable water delivery and phloem elements forming to support nutrient import, though capacity remains limited at this early separation stage. Metabolic activity during bud separation maintains elevated levels supporting continued growth and maturation, with respiration providing energy for pedicel elongation, organ expansion, and approaching reproductive function. Developmental uniformity within cluster during bud separation shows flowers at similar stages with synchronized separation initiation across cluster supporting coordinated anthesis, though slight developmental gradient may persist with distal flowers sometimes slightly advanced. Resource requirements during separation emphasize carbohydrate demand for continued growth and energy storage for anthesis, nitrogen for final protein synthesis particularly in anthers and pollen, and boron critical for pollen viability and approaching germination and tube growth. Hormonal regulation during bud separation involves auxin affecting pedicel initiation and early elongation, possible declining gibberellins as reproductive maturation proceeds, and beginning ethylene involvement in cap loosening processes. Environmental conditions during bud separation include temperature nearing flowering optimum with 18-24°C supporting separation and continued maturation, adequate moisture maintaining turgor supporting elongation, and favorable weather protecting delicate developing structures. Disease vulnerability during bud separation emphasizes powdery mildew with susceptible tissues fully exposed and infection potential high, requiring protective fungicide coverage particularly in conducive humid conditions or susceptible varieties. Developmental progression from bud separation proceeds rapidly toward continued pedicel elongation and full flower individuation, with separation representing brief initial phase in 3-5 day individual flower separation process. Within-bud organ positioning shows anthers positioned centrally within calyptra cap preparing for cap fall and pollen release, pistil extending upward with stigma approaching cap apex positioning for early exposure at anthesis, and nectaries at flower base preparing for nectar secretion. The process represents first of four individual flower separation aspects marking beginning of flower individuation, transitioning rapidly toward pedicel elongation and complete flower separation. Duration typically spans 1-2 days representing brief initiation phase as separation begins, with warm conditions promoting rapid progression toward elongation. Cultivar variation appears in separation initiation timing with some varieties showing early coordinated bud separation while others show more gradual progressive separation, affecting synchrony of subsequent flowering. Regional timing shows flower bud separation during late May-early June in warm climates, early June in temperate regions, and early-mid June in cool areas, occurring 2-4 days before anthesis initiation. Viticultural significance includes marking beginning of final pre-anthesis development, establishing vulnerability period requiring disease protection and favorable conditions, indicating imminent flowering requiring preparation for bloom management, and representing point where flower number becomes definitively countable for yield prediction. Research applications include documenting earliest stages of flower individuation, analyzing coordination of separation initiation across cluster, evaluating environmental effects on separation affecting flowering uniformity, and assessing developmental readiness markers for anthesis prediction.
Flower_Bud_Separation is first of four Individual_Flower_Separation subphases, distinguished by initial flower individuation before substantial pedicel elongation. Represents beginning of transition from compact cluster to separated flowers. Transitions rapidly to Pedicel_Elongation as separation proceeds. Brief phase marking initiation of critical pre-anthesis development. Indicates imminent flowering requiring preparation and protection.
Blütenknospentrennung
Knospenlockerung
beginnende Einzelblütenisolierung
beginning flower individuation
bud loosening
initial flower separation
initiale Blütenseparation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 352-355 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Flower Bud Separation
Gewürztraminer in Alsace vineyards at BBCH 65 (full bloom) exhibits very compact flower clusters of 8-12 cm length bearing 200-400 flowers with minimal branching and tight spacing, creating dense aggregations that predictably develop into severely compact berry clusters (OIV code 9) with 150-300 deformed berries requiring aggressive disease management. Conversely, Grenache in Southern Rhône displays loose flower clusters of 15-25 cm length with 800-1500 flowers distributed across extensive primary, secondary, and tertiary branching, creating open, well-aerated structures (OIV code 2-3) that develop into large, loose berry clusters of 400-1000 berries with minimal disease pressure but increased bird damage risk. Cabernet Sauvignon in Napa Valley shows intermediate flower cluster compactness (OIV code 4-5) with 12-16 cm clusters bearing 400-600 flowers on moderate branching.
A flower cluster is the entire aggregate of flowers borne on a single grapevine inflorescence, representing the complete reproductive unit that will develop into a berry cluster (bunch) following pollination, fertilization, and fruit set, with cluster morphology serving as a critical ampelographic descriptor and determinant of disease susceptibility, harvest logistics, and wine quality potential. The flower cluster encompasses all flowers borne on the inflorescence rachis and its hierarchical branching system (primary, secondary, and tertiary branches where present), typically comprising 50-2000+ individual flowers depending on cultivar genetics, vine vigor, and previous-season developmental conditions. Flower cluster architecture reflects the underlying inflorescence branching pattern established during primordium differentiation within the dormant bud 8-12 months before anthesis, with cluster compactness determined by the ratio of flower number to total inflorescence axis length (rachis plus all branch lengths). Compact flower clusters exhibit high flower density (>15-25 flowers per cm of total axis length) resulting from either high flower number on abbreviated branching systems or from short internodal spacing, while loose flower clusters show low density (3=1) through conical (ratio 2=1-3=1) to broadly winged or compound (ratio <2=1 with pronounced lateral lobes). The spatial arrangement of flowers within the cluster profoundly influences subsequent berry development, disease dynamics, and harvest quality, as compact clusters promote humidity retention, restrict air circulation, and create conditions favoring Botrytis cinerea infection, while loose clusters facilitate drying, spray penetration, and disease management but may suffer bird predation and uneven ripening. Flower cluster morphology exhibits high heritability as a cultivar-specific trait, with compact types (Gewürztraminer, Pinot Noir, Müller-Thurgau) consistently producing tight clusters and loose types (Grenache, Mourvèdre, Syrah) producing open clusters under standard cultural conditions, though environmental modulation during primordium differentiation introduces significant phenotypic plasticity. Cluster compactness assessment follows standardized OIV descriptor code 204, classifying clusters from very loose (code 1= berries on distinct lateral branches, cluster appears lacy) through medium (code 5= berries touch but not compressed) to very compact (code 9= berries severely deformed by mutual compression). The transformation from flower cluster to berry cluster involves substantial architectural modification through differential fruit set, berry abscission (shot berries), and differential berry growth, but fundamental compactness characteristics established at flowering largely persist through maturation.
Flower_Cluster represents the entire reproductive aggregate at anthesis and must be distinguished from Berry_Cluster (the mature fruit aggregate), Inflorescence (which includes the complete floral structure with all supporting axes), and individual Flower (the single reproductive unit). The flower cluster is the critical architectural unit for ampelographic classification and disease risk assessment, as compactness at flowering strongly predicts berry cluster compactness at harvest despite fruit set losses of 30-70%. Cluster compactness is among the most economically significant grapevine traits, directly affecting fungicide requirements, harvest labor costs, and wine quality potential through impacts on disease susceptibility and berry microclimate. Sub-classification into Compact, Loose, and Terminal types reflects major morphological and functional distinctions. The term 'cluster' in viticulture conventionally refers to the berry stage, but botanical accuracy requires 'flower cluster' specification when discussing the anthesis-stage structure.
Blütenbüschel
Blütenstand
Blütentraube
flower bunch
inflorescence cluster
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 94-98 OIV (2009). Descriptor List for Grape Varieties and Vitis Species, 2nd ed., OIV code 204 (bunch density) Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 178-185 Vasconcelos, M.C., Castagnoli, S. (2000). Leaf canopy structure and vine performance. American Journal of Enology and Viticulture 51=390-396.
Flower Cluster
Merlot in Bordeaux shows flower cluster separation during mid-late May Year N+1 at late BBCH 55-early 57 over 6-7 days, with 22-30 primary branches separating along rachis creating 2-4 mm spacing between branch attachment points, secondary branches where present separating 1-2 mm on primary branches, establishing compound cluster architecture with distinguishable flower groups under oceanic spring conditions of 16-20°C mean temperature, adequate moisture, developed photosynthetic capacity, typical of this moderately compound mid-season cultivar approaching flowering readiness
Flower cluster separation represents the developmental phase where compact undifferentiated inflorescence regions differentiate into distinct flower groups through localized spacing development, establishing recognizable cluster organization with separated groups of flowers that will further individualize during subsequent development, corresponding to late BBCH 55 through early BBCH 57 stages. This important organizational phase occurs during mid-May in temperate regions Year N+1 following initial inflorescence elongation, typically spanning 5-7 days as coordinated differential growth creates spatial separation between flower groups revealing underlying architectural pattern established during previous season primordium formation. Cluster separation mechanism involves differential branch elongation with primary branches extending at varying rates creating unequal spacing, localized rachis elongation between branch attachment points generating gaps, and beginning secondary branch spreading where present establishing hierarchical spatial organization. Primary cluster separation proceeds through primary branch differentiation from main rachis with branches elongating and angling away from axis creating first-order spatial organization, with branch spacing along rachis typically 2-5 mm reflecting primordium formation pattern from previous season. Secondary cluster separation occurs in compound architectures where secondary branches separate from primary branches creating second-order spatial organization, with secondary branch spacing typically 1-3 mm establishing fine-scale architectural pattern. Developmental progression shows basipetal-acropetal gradient with apical regions often showing earlier separation reflecting advanced development in earlier-formed structures, while basal regions show progressive separation as development proceeds, creating temporal heterogeneity within inflorescence. Cluster definition emerges during separation as grouped flowers on branch tips become visually distinguishable from neighboring groups, with separation creating perceptual organization of continuous inflorescence into discrete clusters facilitating observation and assessment. Spacing development creates microenvironmental variation within inflorescence with separated regions showing improved air circulation reducing disease pressure potential, enhanced light penetration affecting flower development uniformity, and differential temperature exposure potentially affecting developmental timing within cluster. Architectural pattern expression during separation reveals branching structure established during previous season primordium formation, with highly branched compound architectures showing complex hierarchical separation pattern while simply branched types show straightforward primary branch separation. Branch angle development accompanies separation with branches moving from acute compressed angles toward more obtuse spreading positions, creating three-dimensional spatial organization enhancing resource distribution and later fruit exposure. Flower bud development continues during cluster separation with buds enlarging from 1.5-2.5 mm to 2.5-3.5 mm as flower organs mature within enclosing calyptra, preparing for subsequent individual flower separation and anthesis. Vascular differentiation proceeds during separation with branch vascular traces differentiating from main rachis establishing independent transport pathways, enabling localized resource delivery to separated flower groups supporting development and subsequent fruit growth. Metabolic activity during separation maintains elevated levels supporting continued growth processes, with respiration 8-12 fold above mature tissue levels providing energy for differential expansion, cell division, and organ differentiation. Photosynthetic contribution increases during separation as shoot canopy develops providing substantial assimilate production, transitioning from early emergence dependence on stored reserves toward current photosynthetic support, though inflorescences remain net carbon consumers requiring import. Hormonal regulation involves auxin from developing flowers affecting rachis and branch development through polar transport establishing growth gradients, gibberellins maintaining elongation promotion though concentrations may begin declining, and cytokinins supporting continued cell division in active growth zones. Environmental conditions during separation affect development with temperature continuing as primary factor with optimal 16-24°C supporting rapid uniform separation, cool conditions slowing progression potentially causing compressed cluster development, and warm conditions accelerating separation though excessive heat may cause stress effects. Light intensity affects separation through photosynthetic effects on whole-plant vigor with adequate light supporting robust development, though direct light effects on inflorescence are modest at this stage with structures still partially shaded by developing leaves. Water availability affects separation through effects on cell turgor supporting expansion and overall vine function, with adequate moisture essential for normal spacing development while water stress reduces growth limiting separation and potentially creating compact cluster architecture. Nutrient availability affects separation with adequate nitrogen supporting vigorous development, phosphorus supporting energy metabolism, and balanced nutrition enabling optimal architectural expression without excessive vegetative growth competition. Disease pressure during separation includes powdery mildew colonization risk with developing inflorescence tissues susceptible to infection, requiring protective fungicide applications particularly in favorable disease conditions or susceptible varieties. Pest monitoring during separation includes grape berry moth larval presence potentially affecting flower development, leafhoppers feeding on tender tissues, and thrips colonization affecting flower quality. Within-inflorescence developmental uniformity improves during separation as flower groups individualize at similar rates preparing for coordinated flowering, with well-separated clusters showing good flowering synchrony supporting uniform fruit set. The process shows four distinguishable developmental aspects progressing from primary cluster separation establishing first-order organization, through secondary cluster separation creating hierarchical structure, cluster spacing development enhancing separation, to individual cluster definition completing group organization before individual flower separation. Duration typically spans 5-7 days depending on temperature and cultivar characteristics, with warm conditions promoting rapid separation while cool conditions extend period. Cultivar variation appears in separation extent with loose-clustered varieties showing substantial spacing creating well-separated flower groups while tight-clustered types show minimal separation maintaining compact architecture, branching complexity affecting hierarchical separation pattern, and separation rate affecting developmental timing toward flowering. Regional timing shows flower cluster separation during mid-May in warm climates, late May in temperate regions, and late May-early June in cool areas, occurring during late inflorescence emergence preparing for flowering. Viticultural management considerations include recognizing separation as indicator of approaching flowering enabling preparation for critical bloom period, understanding that separation pattern affects cluster microenvironment influencing disease pressure and fruit quality, monitoring separation uniformity assessing development synchrony affecting flowering timing predictions, implementing pre-bloom disease management protecting susceptible tissues during separation, and appreciating that architectural expression during separation reflects previous season primordium formation quality. Research applications include analyzing mechanisms regulating differential growth creating spatial separation, investigating environmental effects on separation affecting cluster architecture and microenvironment, quantifying relationships between separation patterns and disease susceptibility, evaluating architectural diversity among cultivars and its functional implications, and assessing developmental coordination between separation processes and approaching anthesis.
Flower_Cluster_Separation represents organizational phase within Inflorescence_Development distinguishing flower groups before individual flower separation. Follows Inflorescence_Expansion. Subdivided into Primary_Cluster_Separation, Secondary_Cluster_Separation, Cluster_Spacing_Development, and Individual_Cluster_Definition. Transitions to Individual_Flower_Separation as flowers individualize. Important for establishing cluster microenvironment affecting disease and quality. Architectural expression reveals previous season primordium formation pattern.
Blütengruppenseparation
Blütenstandskompartimentierung
Gescheinsgliederung
Teilblütenstandsseparation
cluster group separation
flower group definition
inflorescence compartmentalization
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 340-350 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Flower Cluster Separation
The flower description of Cabernet Sauvignon documents perfect (hermaphrodite) flowers (OIV 151:3) with five erect stamens bearing fertile anthers producing abundant viable pollen (>90% stainability with Alexander's stain), a well-developed pistil with a receptive bifid stigma on a short style above a bicarpellate ovary containing 2-4 ovules, and a green calyptra that detaches cleanly at the base during anthesis (capfall) — this fully functional hermaphrodite flower enables self-pollination and reliable fruit set (typically 30-50% of flowers setting berries) without requiring cross-pollination from other cultivars, the standard condition for the vast majority of commercially grown V. vinifera wine and table grape cultivars.
An ampelographic description component that documents the morphological characteristics of the grapevine flower, recording the flower type (hermaphrodite, functionally female, or male), the structure and size of stamens and pistil, the calyptra (fused petal cap) morphology, and the functional relationships between reproductive organs that determine pollination biology, fruit set efficiency, and the requirement for cross-pollination in cultivars with imperfect flowers.
Flower_Description addresses the brief but critically important reproductive phase of the grapevine annual cycle — flowering (anthesis) occupies only 7-14 days (typically late May to mid-June in northern hemisphere temperate viticulture) but determines the number and distribution of berries that constitute the harvestable crop. The grapevine flower is structurally distinctive among angiosperms — the five petals are fused at the apex to form a cap-like calyptra (operculum) that detaches at the base and is shed as a single unit during anthesis (a process termed capfall or calyptra abscission), exposing the stamens and pistil simultaneously and enabling both self-pollination and wind-mediated cross-pollination within the 2-4 day window of stigma receptivity: this unusual floral mechanism, combined with the small size of individual flowers (3-5 mm diameter), makes grapevine flower morphology difficult to observe without magnification, limiting the practical utility of floral descriptors for routine cultivar identification compared to vegetative and fruit characters. Nevertheless, flower type (hermaphrodite versus functionally female) is a critically important cultivar attribute because functionally female cultivars (e.g., some table grape varieties and wild V. vinifera subsp. sylvestris) require nearby planting of hermaphrodite pollinizer cultivars for fruit set.
Blüten-Beschreibung
Blütenmorphologie
floral morphology description
flower characterization
reproductive organ documentation
reproduktive Organ-Dokumentation
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press Carmona MJ et al. (2008) A molecular genetic perspective of reproductive development in grapevine, Journal of Experimental Botany 59: 2579-2596
Flower Description
Cabernet Franc in Loire Valley shows flower spacing completion during early June Year N+1 at BBCH 59 over 1-2 days, with 300-550 individual flowers per cluster achieving final spacing of 4-7 mm on pedicels of 3.0-4.5 mm, flower buds 4.5-5.0 mm showing advanced cap loosening preparing for anthesis initiation within 24-36 hours, under temperate maritime conditions of 19-23°C temperature, 50-65% relative humidity, favorable for imminent flowering in this moderately loose-clustered vigorous mid-season cultivar
Flower spacing completion represents the final pre-anthesis arrangement phase where individual flowers achieve definitive spatial positioning establishing optimal inter-flower distances for pollination, air circulation, and accommodation of developing berries, marking culmination of inflorescence development and readiness for flowering. This culminating organizational stage occurs during early June in temperate regions Year N+1 as individual flower separation concludes, typically representing final 1-2 days of development before anthesis initiation at BBCH 59 immediately preceding flowering. Completion achievement involves cessation of pedicel elongation at final lengths of 2-5 mm establishing permanent flower attachment dimensions, finalization of flower positioning with stable spatial orientations, and cluster architecture stabilization with growth processes declining as structure transitions from development to reproductive function. Final spacing characteristics show inter-flower distances typically 3-8 mm depending on cultivar architecture and pedicel length, with spacing adequate for air circulation around individual flowers promoting rapid drying and reducing disease pressure, sufficient for pollen transfer between flowers during wind-assisted self-pollination, and appropriate for accommodating berry expansion after fruit set without excessive crowding. Spatial optimization during completion may involve minor positioning adjustments through differential pedicel growth or branch repositioning, creating more uniform flower distribution and improved microenvironmental conditions within cluster. Flower arrangement pattern at completion reveals complete architectural expression reflecting previous season primordium formation with branching hierarchy fully realized, current season environmental effects on elongation and spacing development integrated, and genetic architectural program completely manifested in final cluster structure. Microenvironmental characteristics in completed flower arrays include favorable air circulation with adequate spacing enabling rapid evaporation of surface moisture reducing powdery mildew and botrytis infection risks, appropriate light exposure with flowers positioned for moderate illumination supporting function without excessive heat stress, and temperature distribution with exposed flowers experiencing conditions near ambient promoting normal development. Flower organ final maturation during spacing completion shows anthers fully developed with mature pollen grains containing bicellular or tricellular male gametophytes, pistils with receptive stigmas showing papillate surface cells and active exudate production, ovaries containing mature ovules with developed megagametophytes, and nectaries producing nectar in 30-50% flowers attracting potential insect pollinators. Calyptra cap loosening advances during completion with connection between fused petals and receptacle weakening through abscission zone development, preparing for cap detachment during imminent anthesis with cap fall typically initiating within 12-24 hours of completion. Pedicel structural maturation at completion includes cessation of elongation growth with cells achieving final dimensions, progressive wall thickening particularly in vascular tissues preparing for berry support function, and beginning subtle lignification in basal pedicel regions establishing permanent structural properties. Vascular function at completion shows fully developed transport capacity with xylem conducting water at rates adequate for flower function and prepared for substantial increases during berry expansion, phloem enabling bidirectional transport with current import supporting flower metabolism and prepared for future export during late berry ripening. Mechanical properties at completion show flexible yet adequate strength for supporting flowers during anthesis and small developing berries immediately after fruit set, with major mechanical development awaiting berry expansion when loads increase substantially requiring pedicel strengthening. Resource status at completion emphasizes stored energy reserves in flowers providing carbohydrates for anthesis processes particularly anther dehiscence and pollen tube growth, nitrogen concentrated in reproductive organs supporting protein synthesis for reproductive function, and critical minerals especially boron essential for pollen germination positioned for utilization. Metabolic transition at completion shows declining growth-related respiration as elongation ceases, increasing reproductive metabolism preparing for anthesis including elevated respiration in anthers supporting pollen maturation and dehiscence, and active sugar metabolism in nectaries producing nectar. Hormonal environment at completion involves low growth promoters as developmental processes complete, sustained reproductive hormones supporting flower function, increasing ethylene triggering or coordinating cap abscission, and integrated signals preparing flowering initiation. Environmental conditions at completion should approach flowering optimum with temperature 20-28°C ideal for anthesis supporting cap fall and pollen release, relative humidity 40-70% adequate for pollen viability without excessive moisture causing issues, and calm to light wind conditions facilitating pollination without physical damage. Weather sensitivity increases at completion as flowers become maximally vulnerable with rain or high humidity potentially delaying anthesis or reducing pollen viability, cool temperatures slowing or preventing normal flowering progression, and excessive heat potentially causing pollen sterility or stigma dysfunction. Disease pressure reaches critical stage at completion with powdery mildew infection potential highest on fully exposed susceptible flower tissues, requiring that protective fungicide coverage be established before completion as applications during flowering are restricted, and botrytis potentially establishing latent infections that will remain dormant through development expressing during ripening. Pest presence at completion may include thrips reaching maximum abundance affecting flower quality and potentially interfering with pollination, grape berry moth adults present potentially ovipositing during or immediately after flowering, and leafhoppers at varying populations depending on region and management. Developmental uniformity at completion shows high synchrony with most flowers at similar maturation state supporting coordinated anthesis across cluster, with synchronized flowering promoting effective pollination through overlapping pollen availability and receptive stigmas, and uniform fruit set creating consistent berry development within cluster. The process represents fourth and final individual flower separation aspect completing all pre-anthesis spatial organization, following bud separation, pedicel elongation, and flower visibility achievement. Duration represents brief 1-2 day culminating period as final adjustments complete and structure stabilizes before flowering, with rapid transition to anthesis initiation. Cultivar variation appears in final spacing characteristics with loose-clustered types achieving 6-10 mm inter-flower distances creating very open architecture while tight-clustered varieties maintaining 3-5 mm spacing producing compact structure, affecting disease pressure, pollination efficiency, and fruit quality potential. Regional timing shows flower spacing completion during early June in warm climates, early-mid June in temperate regions, and mid-June in cool areas, occurring immediately before anthesis with flowering typically initiating within 24-48 hours of completion. Viticultural significance includes marking definitive flowering readiness enabling precise bloom timing predictions, establishing final cluster architecture determining disease pressure, pollination conditions, and fruit development potential throughout season, representing culmination of two-season inflorescence development from previous season primordium formation through current season emergence and separation, and indicating critical transition point requiring optimal conditions and management for successful flowering and fruit set. Research applications include documenting precise spatial relationships in completed flower arrays, analyzing environmental effects on final spacing affecting functional outcomes, quantifying relationships between spacing characteristics and pollination efficiency and fruit set uniformity, evaluating microenvironmental consequences of architectural variation for disease susceptibility and fruit quality, and assessing coordination mechanisms synchronizing spacing completion with anthesis initiation across diverse genotypes and environments.
Flower_Spacing_Completion is fourth and final Individual_Flower_Separation subphase, distinguished by achievement of final pre-anthesis spatial arrangement. Follows Flower_Bud_Separation, Pedicel_Elongation, and Individual_Flower_Visibility. Represents culmination of entire Inflorescence_Development spanning two growing seasons. Transitions immediately to Flowering phase as anthesis initiates. Critical milestone indicating flowering readiness and requiring optimal conditions for successful bloom.
Blütenabstandsabschluss
Vorblüten-Positionierung
abgeschlossene Blütenverteilung
completed flower arrangement
final flower spacing
finale Blütenanordnung
pre-anthesis positioning
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 358-365 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Flower Spacing Completion
The flower type description of Corinto Nero (syn. Korinthiaki, the traditional seedless raisin grape) documents functionally female flowers (OIV 151:4) with well-developed pistils bearing receptive stigmas but reflexed stamens that release sterile, non-viable pollen — this functionally female condition means Corinto Nero cannot self-pollinate and requires wind-borne or insect-transferred pollen from nearby hermaphrodite cultivars for fertilization and seeded berry development, although in practice Corinto Nero is cultivated specifically for its capacity to produce parthenocarpic (seedless, unfertilized) berries that develop without pollination into the small, seedless Zante currants of commerce. The functionally female flower type contrasts with the hermaphrodite flowers of nearly all modern wine grape cultivars (OIV 151:3), which represent the derived condition selected during domestication from the ancestral dioecious (separate male and female plants) state of wild V. vinifera subsp. sylvestris.
A flower description component that records the sexual type of the grapevine flower according to OIV descriptor 151, classifying flowers as hermaphrodite (type 3, fully developed functional stamens and pistil, capable of self-pollination), functionally female (type 4, developed pistil with reflexed non-functional stamens producing sterile pollen, requiring cross-pollination), or male (type 1, developed stamens with rudimentary non-functional pistil, producing pollen but no fruit), with flower type being a fundamental cultivar attribute that determines pollination requirements, fruit set biology, and vineyard planting design.
Flower_Type_Description documents the fundamental reproductive biology attribute that connects grapevine breeding history, domestication genetics, and vineyard management. Wild V. vinifera subsp. sylvestris is dioecious — individual plants are either male (producing pollen but no fruit) or female (producing fruit but requiring cross-pollination) — a condition controlled by a sex-determining locus on chromosome 2 containing linked genes for male sterility and female sterility. The transition from dioecy to hermaphroditism was one of the key events in grapevine domestication, occurring through a mutation that inactivated the female sterility gene while maintaining pollen fertility, producing the self-fertile hermaphrodite flower type that characterizes virtually all cultivated V. vinifera: this hermaphroditic condition eliminated the need for dedicated pollinizer plants and enabled the vegetative propagation of individual productive genotypes without concern for pollination requirements. The small number of cultivated varieties retaining functionally female flowers (Corinto Nero, some Sultana-type cultivars, rare wine grapes such as Madeleine Angevine) represent either incomplete domestication selection or secondary loss of pollen function, and their cultivation requires either intentional cross-pollination management or exploitation of parthenocarpic fruit development.
Blütengeschlecht-Klassifizierung
Blütentyp-Beschreibung
Reproduktionstyp-Charakterisierung
floral gender classification
reproductive type characterization
sexual flower type record
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Fechter I et al. (2012) Candidate genes within a 143 kb region of the flower sex locus in Vitis, Molecular Genetics and Genomics 287: 247-259 May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press
Flower Type Description
Wild Vitis species are predominantly dioecious with separate male and female vines, while domesticated V. vinifera cultivars are 99% hermaphroditic due to selection during domestication. The sex-determining region on chromosome 2 spans approximately 150 kb and includes the VviFSEX gene. Flower sex ratios in wild V. vinifera ssp. sylvestris populations show approximately 50% male, 40% female, and 10% hermaphrodite individuals.
A flowering disposition that is the inherited capacity of a Vitis vinifera plant to develop flowers of a specific sexual morphology, determined by the functional presence or absence of male (androecium) and female (gynoecium) reproductive organs within individual flowers.
Flower_Type_Disposition serves as the parent class for specific sex morphologies. Grapevine flower sex determination involves a two-locus genetic model with male-sterility and female-sterility factors. Hermaphroditism arose through a mutation suppressing female-sterility in male flowers. Some cultivars show partial reversion to ancestral forms under stress conditions.
Blütengeschlecht
Blütenmorphologie-Disposition
Blütentyp
floral morphology trait
flower sex type
sexual type disposition
Picq S et al. (2014) A small XY chromosomal region explains sex determination in wild dioecious V. vinifera and the reversal to hermaphroditism in domesticated grapevines. BMC Plant Biology 14:229 This P et al. (2006) Historical origins and genetic diversity of wine grapes. Trends in Genetics 22:511-519
Flower Type Disposition
Merlot in Bordeaux shows flowering during early-mid June Year N+1 at BBCH 60-69 over 8-10 days, progressing from first flowers opening June 5-7 through peak anthesis June 8-12 with 60-80% flowers open simultaneously to late anthesis completing June 14-16, achieving 45-60% fruit set under oceanic climate conditions of 18-25°C temperature, 50-70% relative humidity, light winds, and no significant rain during bloom, typical of this mid-season moderately productive cultivar producing 300-450 berries per cluster
Flowering represents the critical reproductive phase encompassing anthesis, pollination, and fertilization processes where flowers open through cap shedding, pollen releases and transfers to receptive stigmas, and fertilization occurs establishing zygotes that will develop into seeds and berries, determining fruit set and yield for the season. This essential reproductive period occurs during early-mid June in temperate regions Year N+1 following inflorescence development, typically spanning 7-14 days from first flower opening through completion of anthesis across cluster depending on environmental conditions and developmental uniformity. Flowering encompasses cap shedding where fused petal calyptra detaches exposing reproductive organs, pollen release through anther dehiscence liberating mature pollen grains, pollination through wind-assisted self-pollination transferring pollen to stigmas, pollen germination on receptive stigma surfaces, pollen tube growth through style tissues delivering sperm cells to ovules, and fertilization with sperm-egg fusion initiating embryo development and endosperm formation. The process shows characteristic progression beginning with sporadic early flower opening, accelerating through peak anthesis when maximum flowers open simultaneously, and declining during late anthesis as final flowers complete opening, with entire cluster typically requiring 5-10 days to complete flowering depending on developmental synchrony. Grapevine flowers show distinctive morphology with pentamerous organization having five sepals fused into calyptra cap, five petals fused and coherent with calyptra forming cap that sheds as unit, five stamens with anthers producing pollen, and superior bicarpellate ovary containing typically four ovules though only 0-4 develop into seeds. Cap shedding mechanism involves abscission zone development at junction between fused petals and receptacle, with zone cells degrading middle lamella through enzymatic activity creating separation plane, and mechanical stresses from expanding reproductive organs and potentially turgor changes driving physical detachment. Anther dehiscence proceeds through specialized stomium zones in anther walls where cells undergo programmed breakdown creating longitudinal slits, with dehiscence triggered by humidity decrease causing anther wall dehydration and mechanical opening releasing pollen through slits. Pollen characteristics include small spheroidal grains 25-35 micrometers diameter with three-colporate aperture pattern, tricellular maturity state at release containing vegetative cell and two sperm cells, and limited longevity remaining viable only hours to few days depending on environmental conditions. Pollination mechanism in grapevine involves predominantly self-pollination with wind assistance transferring pollen from anthers to stigmas within same flower or neighboring flowers on same vine, with occasional insect visitation attracted by nectar but playing minor role compared to wind. Stigma receptivity develops progressively showing papillate surface cells producing adhesive exudate capturing pollen grains, with receptivity peaking 1-3 days after flower opening and persisting 4-7 days depending on conditions. Pollen germination on receptive stigma occurs within 1-4 hours under favorable conditions with pollen grain hydrating from stigma exudate, producing pollen tube that penetrates stigma surface growing through transmitting tract tissues in style toward ovary. Pollen tube growth proceeds at rate of 1-3 mm per hour requiring 12-36 hours to traverse 15-30 mm style length reaching ovary, with tube guided by chemotactic signals from ovules growing through micropyle into embryo sac. Double fertilization characteristic of angiosperms occurs with one sperm cell fusing with egg forming diploid zygote initiating embryo development, and other sperm cell fusing with two polar nuclei forming triploid endosperm nucleus initiating nutritive tissue development. Fruit set following successful fertilization shows developing seeds producing hormones particularly auxin and gibberellins triggering berry development, with successful fertilization preventing flower abscission and initiating berry expansion from initial 2-3 mm ovary to mature berry dimensions. Environmental conditions critically affect flowering success with temperature representing primary factor with optimal 20-30°C supporting rapid coordinated anthesis and effective pollination, cool conditions below 15°C delaying flowering and reducing pollen viability, and excessive heat above 35°C potentially causing pollen sterility or stigma dysfunction. Water status affects flowering with moderate water stress potentially beneficial for concentrating carbohydrate reserves supporting reproductive processes, while severe stress impairs function potentially causing poor fruit set. Humidity affects pollen function with moderate 40-70% relative humidity optimal for pollen viability and dehiscence, while high humidity above 80% delays dehiscence and promotes fungal infection, and very low humidity below 30% causes rapid pollen desiccation reducing viability. Rain during flowering represents major concern with water washing pollen from anthers and stigmas, preventing dehiscence, promoting fungal infection particularly botrytis, and physically damaging delicate reproductive structures, potentially severely reducing fruit set. Wind during flowering shows dual effects with light-moderate wind facilitating pollination through pollen dispersal, while strong wind may damage flowers or excessively dry tissues reducing function. Light intensity during flowering affects overall vine vigor and photosynthetic support for reproductive processes, with adequate light supporting robust flowering though direct effects on individual flowers are modest. Nutrient requirements during flowering emphasize boron critical for pollen germination and tube growth with deficiency causing poor fruit set, adequate nitrogen supporting protein synthesis in reproductive tissues, and balanced nutrition supporting overall vine function. Carbohydrate availability affects flowering success with adequate reserves supporting energy-demanding reproductive processes, while limitation from excessive crop load, disease, or previous season stress may reduce flowering vigor affecting fruit set. Hormonal regulation involves auxin from developing seeds triggering berry development and preventing abscission, gibberellins affecting flowering and early berry development with applications sometimes used to manipulate berry size and seedlessness, cytokinins supporting cell division in developing berries, and ethylene potentially involved in cap abscission and flower senescence. Fruit set percentage varies substantially among cultivars and conditions ranging from 10-30% in varieties with poor set or unfavorable conditions creating loose clusters, to 40-70% in good-setting varieties under favorable conditions producing compact clusters, with fruit set determined by successful fertilization frequency among total flowers. Developmental coordination shows flowering synchronized with vegetative growth with shoots typically having 15-20 leaves at flowering providing photosynthetic capacity supporting reproductive metabolism, and balanced vegetative-reproductive allocation supporting both canopy development and successful flowering. Disease pressure during flowering includes botrytis bunch rot with flowers highly susceptible to Botrytis cinerea infection establishing latent infections that remain dormant through development expressing during ripening, and powdery mildew continuing as threat requiring protective management though fungicide applications are restricted during flowering. Pest management during flowering includes monitoring grape berry moth oviposition on clusters potentially affecting berry development, and managing other pests that might interfere with flowering processes or damage developing structures. The process shows two major phases progressing from anthesis encompassing flower opening and pollination, through fertilization establishing seed development and fruit set, with successful flowering determining yield and berry number for the season. Duration typically spans 7-14 days from first to last flower opening depending on temperature, developmental uniformity, and environmental conditions, with warm favorable conditions promoting rapid coordinated flowering within 7-10 days while cool variable conditions may extend period to 12-16 days. Cultivar variation appears in flowering timing with early-flowering varieties showing anthesis in late May-early June while late types flower mid-late June, flowering duration with some varieties showing compressed 5-7 day bloom period while others show extended 10-14 day period, fruit set percentage affecting cluster density and yield, and sensitivity to environmental conditions affecting flowering success reliability. Regional timing shows flowering during late May-early June in warm climates, early-mid June in temperate regions, and mid-late June in cool areas, with timing critical for frost avoidance and favorable weather window. Viticultural management considerations include monitoring flowering progress for timing management operations, protecting flowering period from adverse weather through site selection and possible protective measures, managing canopy optimizing light exposure and air circulation for flower function, ensuring adequate vine nutrition particularly boron supporting successful pollination and fruit set, avoiding stressful conditions during flowering supporting reproductive success, and recognizing flowering as critical determinant of yield and quality requiring optimal conditions and management. Research applications include analyzing mechanisms regulating flowering including hormonal and environmental controls, investigating pollination biology including pollen-stigma interactions and fertilization processes, evaluating environmental effects on flowering success affecting fruit set and yield, quantifying genetic variation in flowering characteristics affecting productivity and adaptation, and assessing climate change impacts on flowering phenology and success across viticultural regions.
Flowering represents critical reproductive phase within Grapevine_Phenological_Process determining yield and quality. Distinguished from Inflorescence_Development which establishes structure and from Fruit_Set which follows successful fertilization. Subdivided into Anthesis covering flower opening and pollination, and subsequent fertilization and early berry development phases. Environmental conditions during flowering critically affect success. Timing represents major phenological marker. Corresponds to BBCH 60-69.
Blütezeit
anthesis period
bloom
flowering period
reproductive phase
reproduktive Phase
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 360-385 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 275-295 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Flowering
Grapevine flowering (anthesis) occurs over 5-15 days per vine, with Sauvignon Blanc completing flowering in 6-8 days under warm conditions (daily mean >18°C) while Cabernet Franc may require 12-15 days during cool springs. Individual flower opening involves calyptra abscission triggered by turgor pressure, typically occurring between 06:00-10:00 hours. A single inflorescence contains 100-500 individual flowers depending on cultivar, with Grenache averaging 350-450 flowers and Pinot Noir 150-250 flowers per cluster.
A reproductive disposition that is the inherited capacity of a Vitis vinifera plant to develop functional inflorescences and individual flowers, encompassing traits related to flower organogenesis, calyptra formation, anthesis timing, and the coordination of male and female reproductive structures within flowers.
Flowering_Disposition encompasses both the structural aspects of flower development and the temporal dynamics of anthesis. The grapevine flower is small (2-3 mm diameter), hypogynous, and pentamerous with a distinctive calyptra (fused petals) rather than separate petals. Flowering duration and synchrony significantly impact fruit set success and cluster uniformity.
Blühdisposition
Blühfähigkeit
Blühvermögen
anthesis disposition
bloom trait
flowering capacity
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 95-115 Vasconcelos MC et al. (2009) The flowering process of Vitis vinifera: a review. American Journal of Enology and Viticulture 60:411-434
Flowering Disposition
Flowering synchrony assessment in Pinot Noir comparing shoots of different vigor reveals the vigor effect on bloom coordination, with moderate-vigor shoots (balanced carbohydrate status) showing high flowering synchrony (90% of flowers within individual clusters opening within 3-day window, minimal between-cluster variation), while excessively vigorous shoots (vegetative sink competition) show reduced synchrony (only 65% of flowers within 3-day window, basal clusters delayed relative to apical), with the synchrony reduction in vigorous shoots correlating with increased shot berry incidence from asynchronous pollination, demonstrating how vine balance affects flowering synchrony with fruit set consequences.
Flowering synchrony quality is an intra-plant synchrony quality that represents the temporal coordination of flowering among different inflorescences and flowers within a single grapevine, characterizing the uniformity of bloom timing that affects pollination success, fruit set uniformity, and the subsequent homogeneity of berry populations through their common fertilization timing. Within the BFO framework, this quality specifically addresses the flowering dimension of intra-plant synchrony, distinguishing bloom coordination from earlier bud break or later ripening synchrony while recognizing that flowering synchrony critically affects fruit set success (synchronized flowering enabling effective self-pollination among flowers opening simultaneously) and establishes the developmental age uniformity of the resulting berry population. The ontological significance lies in flowering synchrony determining both immediate fruit set success and subsequent berry homogeneity, where synchronized flowering enables effective pollen transfer among simultaneously receptive flowers, produces berry populations with common fertilization dates that will develop and ripen together, and compresses the flowering period reducing exposure to adverse weather during this vulnerable stage. The factors affecting flowering synchrony include shoot developmental uniformity (established at bud break and propagating to flowering), cluster position (basal clusters often flowering before apical clusters within shoots), environmental conditions during flowering (temperature affecting flowering rate and thus the spread of bloom across organs), and the inherent synchrony characteristics of different cultivars (some varieties showing naturally more compressed flowering than others). The fruit set implications of flowering synchrony involve the coincidence of pollen release and stigma receptivity necessary for successful fertilization, where synchronized flowering ensures that flowers throughout the vine are at appropriate stages simultaneously, maximizing the opportunity for pollen transfer and minimizing the proportion of flowers that miss optimal pollination timing. The berry homogeneity consequences of flowering synchrony extend throughout the season, as berries fertilized within a narrow window will have similar developmental ages, progress through growth and ripening stages together, and achieve maturity simultaneously, while berries from extended flowering will represent a range of developmental ages at any subsequent time point. Understanding flowering synchrony quality enables prediction of fruit set uniformity, assessment of subsequent berry population homogeneity, and appreciation of the compound effects of early-season synchrony on harvest-time fruit quality.
Flowering_Synchrony_Quality addresses the coordination of bloom timing affecting pollination success and berry population uniformity. This quality determines both immediate fruit set and subsequent berry homogeneity through common fertilization timing. The class supports fruit set prediction and quality assessment through flowering synchrony characterization.
Anthesensynchronie
Bestäubungsfensteruniformität
Blühsynchronie
Blütenuniformität
anthesis synchrony
bloom uniformity
flowering coordination
pollination window uniformity
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 59-144 May P (2004) Flowering and Fruitset in Grapevines. Lythrum Press, Adelaide, pp. 1-119
Flowering Synchrony Quality
The flowering timing description of Merlot documents early-medium flowering (OIV 302:3-5) — reaching 50% capfall approximately 2-3 days before Chasselas in Bordeaux, typically during the first week of June (June 1-10 at Saint-Émilion), approximately 5-7 days before Cabernet Sauvignon at the same site. The flowering-to-harvest interval for Merlot at Bordeaux averages 110-115 days, placing Merlot harvest in mid-to-late September — the earlier flowering and consequently earlier harvest of Merlot compared to Cabernet Sauvignon (harvest mid-October) is the fundamental phenological reason why Merlot dominates the cooler, earlier-harvesting Right Bank appellations (Saint-Émilion, Pomerol) while Cabernet Sauvignon dominates the warmer, later-harvesting Left Bank (Médoc, Graves), with the 10-15 day harvest date difference determining which cultivar achieves optimal ripeness before autumn rainfall and temperature decline compromise fruit quality.
A phenological timing description component that records the date at which 50% of flowers have shed their calyptras (capfall, E-L stage 23, Baggiolini stage I, BBCH 65) — the phenological event marking full bloom and the critical pollination/fertilization period — scored according to OIV descriptor 302 on a scale from very early (state 1) to very late (state 9) relative to Chasselas, with flowering timing determining the expected harvest date (approximately 100-120 days later for most cultivars), the vulnerability to poor fruit set from unfavorable bloom-period weather, and the first temporal anchor for vintage quality prediction.
Flowering_Timing_Description documents the phenological event that is the most critical determinant of vintage success — flowering represents the irreversible commitment of the vine's reproductive effort, and the weather conditions during the 7-14 day bloom period have a disproportionate impact on fruit set, cluster compactness, and ultimately yield and quality. The flowering date is the product of two accumulated thermal intervals — the budbreak-to-flowering period (typically 55-75 days, requiring 350-500 GDD base 10°C depending on cultivar) and the budbreak date itself — with the budbreak-to-flowering interval being more consistent across years than the absolute flowering date because this interval is primarily temperature-driven with minimal photoperiod effect: the consistency of the flowering-to-harvest interval (approximately 100-120 days, or 1,100-1,500 GDD base 10°C, depending on cultivar) makes flowering date the earliest reliable predictor of harvest date, enabling advance planning of harvest logistics, winery scheduling, and labor requirements. The critical sensitivity of fruit set to bloom-period weather — temperatures below 15°C or above 35°C during anthesis reduce pollen viability and stigma receptivity, while rainfall during bloom washes pollen from stigmatic surfaces and promotes calyptra retention (failure of capfall) — means that flowering timing also determines the probability of poor fruit set, with early-flowering cultivars at greater risk from cool, wet conditions (early June being statistically cooler and wetter than mid-June in many European regions) and late-flowering cultivars benefiting from the generally warmer, drier conditions of later June.
Anthese-Datum
Blütezeitpunkt-Beschreibung
Vollblüte-Dokumentation
anthesis date description
bloom timing record
flowering period documentation
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Jones GV and Davis RE (2000) Climate influences on grapevine phenology, American Journal of Enology and Viticulture 51: 249-261 May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press Duchêne E and Schneider C (2005) Grapevine and climatic changes, Agronomy for Sustainable Development 25: 93-99
Flowering Timing Description
Flowering occurs 50-80 days after budbreak across V. vinifera cultivars, with the interval showing less variability (CV 8-12%) than budbreak timing itself (CV 12-22%) due to the stabilizing effect of accumulated thermal time. Chardonnay reaches 50% capfall (BBCH 65) after accumulating 350-400 GDD (base 10°C) from budbreak, while Grenache requires 450-550 GDD to the same stage. The flowering window per cultivar spans 8-15 days, with individual vine flowering duration of 5-10 days under favorable conditions. Optimal flowering temperatures of 20-25°C promote rapid, synchronized anthesis temperatures below 15°C extend flowering duration by 50-100% and reduce fruit set.
A phenological timing disposition that is the inherited capacity of a Vitis vinifera plant to initiate and complete anthesis (BBCH 60-69) at a characteristic time relative to budbreak and environmental conditions, determining the calendar period of pollination and fruit set and influencing growing season partitioning between vegetative and reproductive development.
Flowering_Timing_Disposition serves as the parent class for early and late flowering categories. The timing of flowering critically affects fruit set success through temperature effects on pollen germination and tube growth. Flowering timing also determines the interval available for berry development and ripening, with later flowering compressing the post-veraison period. Climate change is advancing flowering by 2-3 days per decade in many regions, potentially shifting bloom into warmer or cooler periods depending on latitude.
Anthese-Timing
Blühtermin
anthesis timing
bloom timing
flowering phenology trait
Zapata D et al. (2017) Predicting key phenological stages for 17 grapevine cultivars (Vitis vinifera L.). American Journal of Enology and Viticulture 68:60-72 Caffarra A, Eccel E (2011) Projecting the impacts of climate change on the phenology of grapevine in a mountain area. Australian Journal of Grape and Wine Research 17:52-61
Flowering Timing Disposition
The foliar application specification for a Riesling vineyard on calcareous limestone soils in the Rheingau addresses two deficiency conditions — iron chlorosis corrected by 3 applications of chelated iron (Fe-EDDHA at 2 kg/ha in 400 L water) at E-L 9, E-L 15, and E-L 19 to maintain photosynthetic function in young leaves that develop interveinal chlorosis characteristic of lime-induced iron deficiency (soil pH 7.8, active CaCO3 12%, Olsen-P adequate but iron sequestered by carbonate), and magnesium deficiency addressed by 4 applications of magnesium sulfate (MgSO4·7H2O at 10 kg/ha in 400 L water with 0.1% non-ionic surfactant) at véraison and post-véraison intervals to prevent the interveinal reddening of basal leaves that reduces photosynthetic capacity during the critical sugar accumulation period — foliar applications are scheduled for early morning (before 09:00) or late afternoon (after 17:00) when stomata are open and leaf surface temperature is below 25°C to maximize absorption and minimize phytotoxicity risk.
A nutrition management description that documents the direct application of nutrient solutions to grapevine leaf surfaces for absorption through stomata and cuticle, specifying the nutrient element, chemical form, concentration, spray volume, surfactant or adjuvant inclusion, application timing, and repeat schedule designed to correct specific nutritional deficiencies or provide supplemental nutrition during critical developmental windows when soil uptake is insufficient.
Foliar_Application_Specification documents the targeted nutritional intervention used when soil-based nutrient supply is inadequate due to soil chemistry constraints (high pH-induced micronutrient unavailability, antagonistic ion competition), root function limitations (drought, waterlogging, root disease), or the need for rapid correction of acute deficiency during critical developmental windows. Foliar absorption provides a pathway that bypasses soil chemistry limitations — iron, zinc, manganese, and boron deficiencies caused by high soil pH or carbonate content can often be corrected only through foliar application because soil-applied forms of these elements are rapidly immobilized in calcareous soils: foliar nutrient uptake occurs through both stomatal and cuticular pathways, with stomatal absorption providing faster uptake of ionic forms while cuticular penetration of chelated forms provides sustained delivery even when stomata are partially closed. The limitations of foliar nutrition include the small quantities absorbable per application (typically 5-15% of soil-applied rates), the need for multiple applications to sustain tissue concentrations, and the risk of leaf burn from excessive concentration or inappropriate application timing.
Blattapplikations-Protokoll
Blattdüngungs-Spezifikation
foliar feeding specification
foliar nutrition documentation
foliare Nährstoffversorgung-Dokumentation
leaf nutrient spray protocol
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 175-195 Christensen LP (2005) Foliar fertilization in vine mineral nutrition management programs, in Christensen LP and Smart DR (eds) Proceedings of the Soil Environment and Vine Mineral Nutrition Symposium, ASEV pp: 83-90 White RE (2015) Understanding Vineyard Soils 2nd Ed, Oxford University Press pp: 120-135
Foliar Application Specification
Comparative analysis of Foliar_Phylloxera_Resistance_Disposition at Cornell University evaluated leaf galling responses across 35 Vitis accessions following controlled exposure to phylloxera crawlers applied to expanding leaves. Gall density (galls per 100 cm² leaf area), gall size, and phylloxera reproduction within galls were assessed at 21 days post-infestation. V. vinifera cultivars ('Chardonnay', 'Cabernet Sauvignon', 'Riesling') demonstrated strong Foliar_Phylloxera_Resistance_Disposition with gall densities of 0-0.8 galls/100cm², minimal successful gall formation, and no sustained phylloxera reproduction. In contrast, V. riparia accessions—though highly resistant to root phylloxera—showed extreme leaf susceptibility with gall densities of 45-120 galls/100cm², large functional galls, and active reproduction averaging 35 offspring per gall. V. labrusca and V. labrusca hybrids ('Concord', 'Catawba') showed intermediate responses (8-25 galls/100cm²). Histological examination revealed that phylloxera probing on V. vinifera leaves triggered localized cell death within 24 hours, preventing the coordinated cell proliferation required for gall formation. On susceptible V. riparia, no cell death response occurred, and gall development proceeded normally. This inverse pattern of foliar versus root resistance demonstrates that Foliar_Phylloxera_Resistance_Disposition is genetically and mechanistically distinct from Root_Phylloxera_Resistance_Disposition.
A Foliar_Phylloxera_Resistance_Disposition is a Phylloxera_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that prevent or limit feeding, gall formation, and population development by the gallicole (leaf-feeding) form of grape phylloxera (Daktulosphaira vitifoliae) on foliage. This disposition represents resistance to a different phase of the phylloxera life cycle than root resistance, addressing the aerial galls that form on leaf undersides when winged or crawling phylloxera colonize leaves. Foliar phylloxera galls are pouch-like structures opening on the lower leaf surface, each containing a fundatrix female that reproduces parthenogenetically, with subsequent generations either remaining on leaves or migrating to roots. While foliar phylloxera infestation is less economically damaging than root infestation—rarely causing significant vine decline—heavy galling can reduce photosynthetic area, stress young vines, and serves as a population reservoir for root colonization. Interestingly, susceptibility patterns to foliar versus root phylloxera are often inversely related among Vitis species: American species (V. riparia, V. rupestris) that show strong root resistance often support extensive leaf galling, while V. vinifera with catastrophic root susceptibility shows resistance to leaf gall formation. This inverse relationship suggests different genetic control of resistance in different tissues. Mechanisms of foliar resistance include leaf anatomical features preventing gall initiation, hypersensitive response killing cells at probing sites, and failure of leaves to undergo the cell proliferation required for gall formation.
Foliar_Phylloxera_Resistance_Disposition is_a Phylloxera_Resistance_Disposition. This class addresses resistance to the gallicole (leaf-feeding) form, which is economically less important than root resistance but biologically fascinating due to the inverse susceptibility pattern between V. vinifera (leaf resistant, root susceptible) and American species (leaf susceptible, root resistant). The child class Gall_Formation_Resistance_Disposition specifically addresses the cellular mechanisms preventing leaf gall development. Foliar phylloxera populations can serve as inoculum source for root colonization, making leaf resistance potentially relevant for integrated phylloxera management even where root resistance is provided by rootstocks.
Blattgallen-Resistenz
Blattreblaus-Resistenz-Disposition
Gallikolenresistenz
foliar galling immunity
gallicole resistance
leaf gall resistance
Granett J et al. (2001) Biology and management of grape phylloxera, Annual Review of Entomology 46:387-412 Forneck A, Huber L (2009) (A)sexual reproduction - a review of life cycles of grape phylloxera, Daktulosphaira vitifoliae, Entomologia Experimentalis et Applicata 131:1-10 Sterling C (1952) Ontogeny of the phylloxera gall of grape leaf, American Journal of Botany 39:6-15
Foliar Phylloxera Resistance Disposition
V. riparia-derived rootstocks such as Riparia Gloire de Montpellier show high foliar phylloxera resistance with fewer than 3 galls per leaf under heavy gallicole pressure, essential for maintaining healthy mother-vine canopies in rootstock multiplication blocks in Champagne and Burgundy. V. rupestris-based rootstocks including St. George exhibit moderate foliar resistance (10-20 galls per leaf) in exposed nursery conditions. V. berlandieri × V. riparia crosses such as SO4 and 5BB Kober demonstrate intermediate foliar resistance, with gall formation reduced by 60-80% compared to susceptible V. vinifera cultivars like Chasselas or Riesling in Austrian and German mother-vine plantations.
A phylloxera-resistance rootstock role that inheres in a grafted rootstock by virtue of its specific capacity to resist or suppress leaf-galling (gallicole) forms of grape phylloxera (Daktulosphaira vitifoliae). Although rootstock foliage is typically not exposed in commercial grafted vineyards, foliar phylloxera resistance is relevant in rootstock mother-vine blocks where rootstock foliage is directly exposed to gallicole attack, in ungrafted rootstock nurseries, and in breeding programs evaluating overall phylloxera resistance profiles. Foliar resistance mechanisms include leaf epidermal thickness, trichome density that inhibits crawler settling, rapid localized necrosis at oviposition sites, production of volatile terpenoids that deter egg-laying, and reduced nutritional suitability of mesophyll tissue for gall development. Resistance is quantified by gall density per leaf (resistant genotypes typically showing fewer than 5 galls per leaf versus 50-200+ on susceptible V. vinifera foliage) and by egg production per gall (fewer than 20 eggs in resistant genotypes versus 200-500 in susceptible hosts).
Foliar_Phylloxera_Resistant_Rootstock_Role is ontologically specific to the gallicole (leaf-galling) phylloxera life cycle form. In standard commercial viticulture with grafted vines, the rootstock foliage is underground or absent foliar resistance becomes operationally relevant primarily in propagation systems (mother-vine blocks, nurseries) and in regions where ungrafted rootstock plantings or rootstock suckers provide canopy for gallicole colonization. Certain V. vinifera cultivars (e.g., Frontenac, some Muscadinia × Vitis hybrids) show natural foliar resistance, but these are classified under scion-related roles rather than rootstock roles.
blattreblausresistente Unterlage
foliar phylloxera resistant rootstock
gallicol-resistente Unterlage
gallicole phylloxera resistant rootstock
gegen Blattreblaus resistente Unterlage
leaf phylloxera resistant rootstock
Forneck A, Huber L (2009) (A)sexual reproduction – a review of life cycles of grape phylloxera, Journal of Insect Science 9: 1-10 Powell KS (2012) A holistic approach to future management of grapevine phylloxera, Springer pp.: 219-251 Granett J et al. (2001) Biology and management of grape phylloxera, Annual Review of Entomology 46: 387-412 Nabity PD et al. (2013) Leaf-galling phylloxera on grapes reprograms host metabolism and morphology, PNAS 110(41): 16663-16668
Foliar Phylloxera Resistant Rootstock Role
A following-season proleptic shoot on Vitis vinifera cv. Pinot Noir, axillary bud formed at node 8 during previous growing season (June year 1), entered and maintained dormancy through winter (November-March), activated during normal spring budbreak (late April year 2) concurrent with primary bud breakage from winter positions, emerged as lateral shoot from previous season's cane (now mature one-year wood) at similar timing to primary shoots, developed to 62cm length by August with 9 leaves, representing standard proleptic timing (spring emergence following dormancy) contrasting with delayed variants showing late-season activation, this following-season lateral arose following partial winter freeze damage reducing primary bud survival and potentially stimulating compensatory axillary activation creating both primary shoot emergence from surviving winter buds and proleptic lateral emergence from previous-season axillary positions as combined recovery response.
A proleptic lateral shoot emerging during normal spring budbreak of the season following axillary bud formation, characterized by axillary bud formation during primary shoot development one season, winter dormancy period, and activation at normal spring timing following season creating lateral shoots from previous-season positions concurrent with primary shoot emergence from winter buds, and representation of standard proleptic timing pattern when proleptic laterals occur contrasting with delayed proleptic variants showing abnormal late activation. Following-season proleptic shoots demonstrate typical dormancy-mediated lateral development when proleptic pattern occurs, with axillary buds breaking dormancy at normal spring timing though from previous-season origin positions creating laterals concurrent with normal primary shoot development, uncommon overall pattern in viticulture but when proleptic laterals do occur, following-season spring timing is standard rather than exceptional.
Following-season proleptic shoots represent typical timing for proleptic lateral development when this dormancy-mediated pattern occurs, with spring emergence during normal budbreak period being standard proleptic timing contrasting with delayed variants showing unusual late activation. The following-season spring pattern creates laterals emerging from previous-season cane positions concurrent with primary shoots from winter bud positions, potentially creating mixed-origin canopy with some shoots from current fruiting wood (normal) and others from previous-season lateral positions (unusual), requiring assessment to distinguish origins and understand canopy composition. Following-season proleptic activation typically results from triggers including winter or spring damage reducing primary bud availability (stimulating compensatory axillary activation), variety tendencies toward delayed lateral development (rare but documented in some cultivars), or specific environmental conditions favoring proleptic over sylleptic pattern. Management involves recognizing previous-season origin of proleptic laterals (distinguishing from normal primary shoots based on emergence positions and wood age), assessing functional value (typically limited compared to primary shoots), and making retention versus removal decisions based on canopy needs and shoot quality.
Folgesaison-proleptischer Trieb
Frühjahrs-Achseltrieb
normal-timing proleptic lateral
spring-emerging proleptic
standard proleptic shoot
Based on Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press, pp. 85-105 Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150.
Following Season Proleptic Shoot
The formal appellation name Châteauneuf-du-Pape is registered in the French INAO appellation registry as an AOC/AOP designation covering 3,200 hectares across the communes of Châteauneuf-du-Pape, Courthézon, Orange, Bédarrides, and Sorgues in the southern Rhône Valley — the formal name must appear on all wine labels claiming this origin and is legally protected against misuse, imitation, or evocation within the EU and in countries recognizing the Lisbon Agreement on Appellations of Origin, with the precise orthography (including hyphenation, capitalization, and diacritical marks) legally defined in the cahier des charges registered with INAO.
An appellation designation consisting of the officially registered name of a delimited viticultural area as recorded in the relevant national or supranational appellation registry, constituting the legally protected designation that must appear on wine labels and commercial documents when claiming origin from that geographic area.
Formal_Appellation_Name is the specific textual string that constitutes the legally recognized designation of a geographic wine origin. The formal name may differ from colloquial or historical usage — for example, the formal appellation name registered with INAO is Chablis rather than the historical Grand Chablis or Chablis Grand Cru (which is a separate registered appellation name for the top tier within the Chablis system): precision in formal appellation naming is legally consequential because labeling regulations require exact correspondence between the name on the label and the registered designation, and variations in spelling, punctuation, or capitalization can constitute regulatory violations. The formal appellation name is the primary key linking geographic indications to their associated regulatory specifications (cahier des charges, disciplinare di produzione, Produktionsvorschriften).
amtliche Ursprungsbezeichnung
eingetragener Herkunftsname
official origin name
offizieller Appellationsname
protected appellation designation
registered appellation name
INAO (2023) Les signes officiels de la qualité et de l'origine EU Regulation 1308/2013 Art. 96-97 on registration of designations EU eAmbrosia database of registered geographic indications
Formal Appellation Name
Touriga Nacional bears a Fortified_Wine_Scion_Role as the most prized variety for Vintage and Vintage Port, contributing intense color (anthocyanins exceeding 2500 mg/kg), floral aromatics (violets, roses), and concentrated tannins that define top Douro Port houses' flagship wines. Palomino Fino serves in a Fortified_Wine_Scion_Role as the exclusive variety for dry Sherry styles (Fino, Manzanilla, Amontillado, Oloroso), providing neutral character that allows flor yeast and oxidative aging to define the wine. Muscat Blanc à Petits Grains bears a Fortified_Wine_Scion_Role for Vins Doux Naturels (Beaumes-de-Venise, Frontignan, Rivesaltes) and Rutherglen Muscat, where fortification preserves intense terpene aromatics and residual sweetness.
A wine production scion role that inheres in a grafted scion cultivar by virtue of its designation for fortified wine production, where grape-derived or neutral spirit is added to wine before, during, or after fermentation to elevate alcohol content (typically to 15-22% v/v) and stabilize the wine against microbial spoilage. Fortified wine scions are selected for characteristics that complement the specific fortification style, including capacity for high sugar accumulation (supporting residual sweetness in styles where fermentation is arrested by spirit addition), appropriate aromatic profiles that integrate with elevated alcohol and oxidative aging, and phenolic structures suited to extended cask maturation. Fortification fundamentally alters wine chemistry and aging trajectory, with the timing of spirit addition determining style—early addition (during fermentation) yields sweet wines with residual grape sugar (Port, Banyuls), while post-fermentation addition yields dry wines (fino Sherry, Sercial Madeira) that may be sweetened subsequently. The category encompasses diverse regional traditions including Port (Portugal), Sherry (Spain), Madeira (Portugal), Marsala (Italy), and Vins Doux Naturels (France).
Fortified_Wine_Scion_Role is subclassed by regional style (Port, Sherry, Madeira, Muscat Fortified), reflecting distinct cultivar requirements, fortification protocols, and aging regimes. The fortification timing distinction is fundamental: Port and most Muscats are mutéd (spirit-arrested) during fermentation preserving grape sugar, while Sherry undergoes complete fermentation before fortification. Fortified wines historically developed as solutions to wine stability during long sea voyages, with spirit addition preventing refermentation and spoilage. Global fortified wine production has declined relative to table wine but maintains strong regional traditions and collector markets for aged examples.
Likörweinsorte
Süßweinrebe für Likörwein
fortified grape variety
fortified wine variety
liqueur wine variety
verstärkte Weinsorte
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Jackson RS (2014) Wine Science 4th Ed, Academic Press pp.: 480-530 Jefford A (2004) The New France, Mitchell Beazley Mayson R (2016) Port and the Douro 3rd Ed, Infinite Ideas
Fortified Wine Scion Role
Fortified wine specification for Touriga Nacional Port from the Douro Valley documents harvest at 12-13° Baumé (lower than table wine to prevent excessive final alcohol), fermentation arrested by addition of 77% vol. grape spirit (aguardente vínica) at approximately 6-8° Baumé remaining (retaining 90-110 g/L residual sugar), fortification to 19-22% vol. alcohol, and subsequent classification pathway — Vintage Port (single-year declaration, 2 years cask aging, reductive bottle aging for 10-40+ years) or Tawny Port (multi-year blending, extended oxidative cask aging in 550 L pipes for 10-40+ years indicated on label). Fortified wine specification for Muscat à Petits Grains from Rutherglen documents progressive classification through the Rutherglen Muscat quality ladder — Rutherglen Muscat (3-5 years average age, 180-250 g/L sugar, fresh raisin character), Classic (6-10 years, 200-280 g/L, dried fruit complexity), Grand (11-19 years, 250-350 g/L, toffee-rancio development), and Rare (20+ years, 300-400+ g/L, extraordinary concentration with mahogany color and limitless complexity).
Fortified wine specification is a wine style description entity that documents the compositional, sensory, and production-method parameters defining wines whose alcohol content has been increased through the addition of grape spirit (aguardente, brandy, or neutral spirit) during or after fermentation, encompassing the timing of fortification (during fermentation to retain residual sugar, or post-fermentation for dry styles), target final alcohol content (typically 15-22% vol.), aging regime (oxidative, reductive, or combined), classification within regional systems (Vintage, LBV, Tawny, Colheita for Port Fino, Amontillado, Oloroso, Palo Cortado for Sherry Rutherglen Muscat classifications), and the specific grape composition and cultivar requirements that distinguish fortified wine production from table wine applications.
Fortified wine specification documents a winemaking category that represents some of the most historically significant and geographically defined wine traditions, yet one that occupies a diminishing share of global wine consumption — a tension that makes comprehensive ontological documentation particularly valuable for preserving knowledge of these specialized production systems. The fundamental distinction within fortified wines is the timing of spirit addition — fortification during active fermentation (as in Port, Banyuls, Muscat de Beaumes-de-Venise, and Rutherglen Muscat) arrests yeast activity and preserves unfermented grape sugar as natural residual sweetness, while post-fermentation fortification (as in dry Sherry, dry Madeira, and some Marsala styles) produces wines that are dry despite elevated alcohol. The grape spirit used for fortification varies by tradition — Portuguese aguardente at 77% vol. for Port, neutral spirit at 96% vol. for French Vins Doux Naturels, and brandy spirit at 57-60% vol. for Australian fortified wines — each contributing differently to the aromatic profile of the finished product. The aging systems for fortified wines are among the most complex in oenology — Sherry's solera fractional blending system, Port's dichotomy between reductive (Vintage, LBV) and oxidative (Tawny) aging pathways, Madeira's estufagem heat treatment, and Rutherglen Muscat's outdoor solera-like blending all create unique maturation environments that transform base wine over decades. Cultivar requirements for fortified wine production differ substantially from table wine — Port regulations specify recommended varieties (Touriga Nacional, Touriga Franca, Tinta Roriz, Tinta Barroca, Tinto Cão), Sherry is produced exclusively from Palomino (with Pedro Ximénez and Muscat for sweet styles), and Rutherglen Muscat requires Muscat à Petits Grains (Brown Muscat clone).
Aufspritung-Weinspezifikation
Likörweinparameter
Likörweinspezifikation
Port specification
Sherry specification
Süßweinspezifikation
dessert wine specification
fortified wine parameters
liqueur wine specification
verstärkter Wein Spezifikation
Bakker J, Clarke RJ (2012) Wine Flavour Chemistry, 2nd Ed, Wiley-Blackwell, pp.: 312-355 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 776-828 Jefford A (2004) Port and the Douro Valley, Mitchell Beazley, pp.: 30-65 Chambers E, Hemstead J (2004) Muscat: The Classification of Australian Liqueur Muscat and Tokay, Rutherglen Winemakers Association
Fortified Wine Specification
A Cabernet Sauvignon clone 08 vine in the Foundation Plant Services vineyard at UC Davis bears a Foundation_Plant_Role, derived from virus-eliminated nuclear stock, maintained in an isolated foundation block with annual visual inspection and periodic ELISA/PCR testing, available for budwood acquisition by California-licensed nurseries under CDFA protocols. A Chardonnay clone 04 plant in the New York State foundation vineyard (Cornell) serves in a Foundation_Plant_Role, providing tested material for northeastern US nursery production adapted to the region's climate and pest pressures. A Riparia Gloire rootstock maintained at the Virginia Tech foundation collection bears a Foundation_Plant_Role, supporting East Coast nursery access to certified rootstock material without dependence on West Coast sources.
A certified mother plant role that inheres in a grapevine maintained as foundation stock within certification schemes using US-derived terminology, corresponding functionally to the Basic category of EU certification frameworks. Foundation plants represent tested material derived from nuclear (Pre-Basic) stock and maintained under controlled conditions for distribution to commercial nurseries, occupying the intermediate tier between elite nuclear collections and commercial multiplication. Foundation plant programs, exemplified by UC Davis Foundation Plant Services (FPS) and similar state programs, maintain virus-tested clones of commercially important varieties available for acquisition by licensed nurseries, with material undergoing specified testing protocols (ELISA panels, PCR assays) and maintained under isolation to prevent reinfection. The "foundation" designation emphasizes the role as the base (foundation) from which the commercial propagation pyramid is built, with foundation material periodically renewed from nuclear stock to prevent generation drift and accumulated pathogen risk.
Foundation_Plant_Role captures US certification terminology where "foundation" corresponds to EU "Basic" category, representing tested material one tier below nuclear/Pre-Basic stock. The US system developed independently from EU frameworks but serves equivalent functions international harmonization efforts increasingly align terminology, though legacy usage persists. Foundation Plant Services (established 1958) pioneered systematic grapevine virus testing and clean stock programs, with its protocols influential globally. Foundation material distribution typically requires licensing agreements and phytosanitary certification for interstate or international movement, with receiving nurseries accepting responsibility for maintaining health status through the final multiplication stages.
Basisstock
Foundation-Mutterpflanze
foundation mother vine
foundation stock plant
registered stock plant
Golino DA et al. (2017) Foundation Plant Services and clean plant programs, HortScience 52: 1190-1194 Rowhani A et al. (2018) Grapevine virus testing protocols, UC Davis Foundation Plant Services USDA-APHIS (2020) Grapevine certification program standards Weber E et al. (2002) The California grapevine certification program, American Journal of Enology and Viticulture 53: 130-134
Foundation Plant Role
A four-arm head on Vitis vinifera cv. Grenache in Châteauneuf-du-Pape, featuring four arms emerging from head at 38cm height positioned at 88°, 91°, 89°, and 92° intervals (nearly perfect 90° spacing), each arm extending 28-32cm bearing 4 two-bud spurs, total 16 spurs supporting approximately 29 shoots and 3.1kg crop in classic four-arm gobelet architecture, representing traditional Châteauneuf standard maintained across most estates and protected under AOC regulations, four-arm configuration providing balanced structure, adequate yield for economic viability, and proven quality outcomes supporting premium pricing justifying lower productivity compared to modern cordon alternatives.
A multi-arm head featuring exactly four arms radiating from central head, representing traditional gobelet standard and most common head-trained configuration, characterized by four-way branching typically at approximately 90° intervals creating balanced square or cross pattern when viewed from above, proven architecture balancing productivity and quality across diverse varieties and regions, and widespread employment in traditional Mediterranean viticulture as archetypal head-training form. Four-arm heads represent classic head-training architecture, established as standard in many traditional regions (Southern Rhône, Languedoc, Priorat, parts of Rioja), providing balanced capacity distribution (each arm carrying one-quarter vine capacity), architectural stability (square geometry providing structural balance), adequate productivity for economic viability (12-20 spurs typical), and proven performance across centuries of traditional viticulture in Mediterranean climates and beyond.
Four-arm heads represent the gold standard of head training, with four-arm architecture dominating traditional viticulture in Mediterranean regions and beyond. The four-arm configuration provides optimal balance among competing objectives including adequate productivity (16-24 spurs typical providing economically viable yields), balanced structure (90° spacing creating stable square geometry), quality outcomes (proven performance in prestigious regions), architectural simplicity (four arms easier to maintain than more complex configurations), and traditional authenticity (four-arm gobelet being archetypal form recognized across regions and markets). The ubiquity of four-arm configurations reflects centuries of empirical optimization, with four arms emerging as sweet spot balancing all relevant factors. Alternative arm counts (three, five, six) appear in specific contexts but four-arm remains dominant standard in traditional head-trained viticulture worldwide.
Vierarm-Kopf
four-way head
klassische Gobelet-Form
quadruple-arm head
standard gobelet head
Four Arm Head
A four-branch permanent arm on Vitis vinifera cv. Catawba in Geneva Double Curtain system, featuring trunk dividing bilaterally at 90cm height creating two primary arms extending left and right 35cm each, each primary arm then bifurcating upward and downward at 45° angles creating four secondary arms total (two ascending to 140cm height, two descending to 70cm height), each secondary arm extending 60cm bearing 10-11 spurs, total 42 spurs distributed across four distinct curtain positions creating maximum canopy division, shoots from upper arms growing upward, shoots from lower arms growing downward, resulting in four separate fruiting zones with enhanced light penetration and air circulation, representing classic four-branch Geneva Double Curtain architecture designed for disease management and quality improvement through extreme canopy division.
A multi-branched permanent arm featuring exactly four distinct branches, characterized by quadruple division creating four separate fruiting zones, typical configurations including bilateral primary division with secondary bifurcation creating four terminals (Geneva Double Curtain pattern), four-way symmetrical branching at 90° intervals, or paired bilateral arms at different heights, and suitability for divided-canopy systems requiring maximum canopy distribution within compact footprint or specialized training approaches emphasizing extreme canopy division for light and air penetration. Four-branch permanent arms represent common multi-branched configuration in specialized training systems, particularly Geneva Double Curtain and variants, providing substantial canopy division benefits through multiple distinct fruiting zones while maintaining manageable structural complexity compared to higher-order multi-branched configurations.
Four-branch permanent arms are most closely associated with Geneva Double Curtain training system, developed specifically to create divided canopy structure improving light penetration and reducing disease pressure in humid regions. The four-branch configuration achieved through two-stage branching (primary bilateral split, secondary bifurcation of each primary) creates geometrically complex but functionally effective architecture. Establishment requires 3-4 years of careful training achieving proper branch angles and positions, with particular attention to secondary bifurcation points which concentrate substantial mechanical stress. Management challenges include maintaining balance among four branches (preventing dominance), managing complex pruning at multiple branch levels, and ensuring structural integrity at junction points. Benefits include maximum canopy division within reasonable structural complexity, proven disease pressure reduction in humid climates, improved fruit quality through enhanced light penetration, and adaptability to mechanical operations despite geometric complexity.
Viergabelarm
four-way arm
quad arm configuration
quadruple-branch arm
vierfach verzweigte Arm
Based on Shaulis et al. (1966) "Response of Concord grapes to light, exposure and Geneva Double Curtain training" Proceedings of the American Society for Horticultural Science 89=268-280 Reynolds & Vanden Heuvel (2009) "Influence of grapevine training systems on vine growth and fruit composition= a review" American Journal of Enology and Viticulture 60=251-268 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide.
Four Branch Permanent Arm
A four-branch tendril at node 7 on a vigorous shoot of Vitis vinifera cv. Carignan in Languedoc, France, exhibiting symmetric double bifurcation architecture with primary bifurcation at 5 cm from shoot attachment dividing into two arms that each bifurcate again at 9-10 cm creating four terminal branches of 8-11 cm individual length (total combined length 35 cm), all four arms showing 2.4-3.0 mm diameter indicating good vascular development with slight advantage to the two earlier-bifurcating arms, branch angles creating roughly radial architecture covering 280-degree arc when viewed from above, circumnutation movements across all arms collectively exploring approximately 740 cubic centimeters, three arms making successful contact with horizontal and vertical trellis wires creating very stable triangulated support system, one arm continuing search movements, occurring on a shoot with strong vigor (total length 165 cm, 16 nodes) in well-managed vineyard with balanced fertilization and irrigation, measured at BBCH stage 73 (berries beginning to touch) in early July, representing typical four-branch development through symmetric double bifurcation pattern.
A multi-branched tendril exhibiting exactly four terminal branches in Vitis vinifera, characterized by architecture producing four independent arms through either a single four-way branch point (rare), two sequential bifurcations creating balanced four-arm architecture, or asymmetric branching patterns with combinations of bifurcation and trifurcation producing four total terminals, representing the most common multi-branched configuration accounting for 60-75% of tendrils with four or more branches, and providing substantial enhancement in contact probability and mechanical redundancy compared to simpler branching patterns while maintaining manageable resource demands achievable under favorable but not exceptional vigor conditions. Four-branch tendrils develop through various developmental pathways including simultaneous activation of four meristems at a single branch point creating true four-way division (very rare, <5% of four-branch tendrils), double bifurcation where initial bifurcation is followed by bifurcation of both arms creating symmetric four-branch architecture (most common, 60-70%), or mixed patterns where bifurcation is followed by trifurcation of one arm creating asymmetric four-branch layout (20-30%). The functional capabilities of four-branch tendrils include exploration volumes of 650-850 cubic centimeters depending on arm lengths and spreading angles, high probability of 2-3 successful attachments creating robust multi-point support, and effective load distribution across multiple attachment points supporting heavy fruit loads or extended shoot sections, achievable on moderately vigorous to vigorous shoots without requiring the exceptional vigor necessary for five or more branch development.
Four-branch tendrils are distinguished from trifurcated tendrils by four rather than three terminal branches, and from five-plus-branch tendrils by exactly four terminals. Developmental pathway identification (true four-way versus double bifurcation versus mixed patterns) provides information about branching regulation mechanisms and can serve as cultivar-specific diagnostic features. Symmetric four-branch architecture through double bifurcation represents balanced resource allocation and creates geometrically optimal exploration patterns, while asymmetric patterns indicate differential meristem activation or resource availability. Four-branch frequency correlates with shoot vigor across cultivars, with vigorous varieties and optimal growing conditions supporting higher frequencies (8-15% of branched tendrils) while moderate vigor reduces four-branch occurrence (2-5%). Functional performance of four-branch tendrils depends critically on individual arm development, with robust arms providing excellent climbing capacity while weak arms compromise the architectural investment despite increased branch number.
four-arm tendril
quadrupel-gabelige Kletterorgan
quadruple-branched tendril
tetra-furcated tendril
vierarmige Ranke
vierfach verzweigte Ranke
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 183-197 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 237-258.
Four Branch Tendril
Cabernet Sauvignon hermaphrodite flowers at BBCH 59-61 show four ovule ovaries in 68-76% of flowers, with ovules distributed as two per locule (paired ovules attached to septum surfaces in each of the two locules), achieving 50-58% fruit set with successful flowers producing berries containing 2.4-3.4 seeds on average reflecting 60-85% fertilization success of the 4 available ovules. Syrah exhibits constitutive high four ovule ovary frequency (72-82%) across diverse sites and vintages, with four ovule ovaries distributed symmetrically (2 ovules per locule in bilocular organization), producing berries with mean seed numbers of 2.6-3.5 seeds when environmental conditions during bloom support high fertilization success. Chardonnay under optimal growing conditions (adequate irrigation, balanced nutrition, moderate temperatures during BBCH 52-55) produces 74-83% four ovule ovaries versus 62-68% under moderate stress conditions, demonstrating environmental modulation of ovule initiation with favorable conditions maximizing ovule number. Merlot berry size analysis shows berries from four ovule ovaries averaging 12.2-13.8 mm diameter versus 11.2-12.6 mm from two ovule ovaries (8-10% size increase), with the correlation persisting across different fruit set levels and environmental conditions, suggesting that ovule/seed number positively influences berry growth potential through seed-derived hormone effects on pericarp cell division and expansion during the critical BBCH 71-77 berry development phase.
A four ovule ovary is an ovule containing ovary that contains exactly four ovules, typically distributed as two ovules per locule in the bilocular Vitis ovary organization, representing the upper end of the normal ovule number range and characteristic of many cultivars that consistently produce maximum ovule numbers or representing a substantial proportion of flowers in cultivars showing variable ovule numbers. Four ovule ovaries are characterized by= (1) exactly four ovules present, typically two per locule in symmetric distribution in the standard bilocular organization (2) ovules positioned on septum surfaces in axile placentation with two attachment points per locule corresponding to positions along the fused carpel margins forming the septum (3) ovary showing normal bilocular structure with two locules separated by septum, with paired ovules in each locule (4) ovules showing normal mature structure at anthesis with completed megasporogenesis and viable embryo sacs when ovule development is successful. Four ovule ovaries represent the maximum typical ovule number in Vitis, with rare exceptions showing 5-6 ovules through supernumerary ovule development representing <2% of flowers in most cultivars. Four ovule ovaries occur through= (1) genetic determination in cultivars with constitutively high ovule number, where 70-90% of flowers consistently produce 4 ovules (2) favorable developmental conditions promoting complete ovule initiation in cultivars with moderate ovule numbers, increasing four ovule ovary frequency from baseline 40-60% to 60-80% (3) enhanced resource availability during ovule development (BBCH 54-55) supporting full ovule complement development. Four ovule ovaries have functional implications for reproduction and berry characteristics= maximum ovule number (4) provides maximum seed number potential, with successful fertilization producing berries with up to 4 seeds (typical 2-4 seeds depending on fertilization success, averaging 2.5-3.5 seeds in hermaphrodite cultivars under favorable conditions) maximum ovule number may slightly increase fruit set probability by providing more fertilization targets, though differences versus three ovule ovaries are typically modest berries from four ovule ovaries tend to be slightly larger on average than berries from two ovule ovaries when other factors are equal (8-12% diameter increase), possibly due to enhanced embryo/endosperm-derived growth hormone production from more developing seeds stimulating berry cell division and expansion. Many economically important wine cultivars show high four ovule ovary frequencies= Cabernet Sauvignon produces 65-78% of flowers with 4 ovules Merlot shows 58-72% four ovule ovaries Chardonnay exhibits 62-75% four ovule ovaries Syrah produces 68-80% four ovule ovaries. The four ovule condition represents maximum reproductive investment in ovule production within the normal range, potentially reflecting genetic programs optimizing seed number for fruit dispersal success or reflecting cultivar selection for larger berries through enhanced seed number. Environmental conditions influence four ovule ovary frequency= favorable growing conditions during flower development (adequate water, nutrition, moderate temperatures during BBCH 52-55) maximize four ovule ovary frequency stress conditions (water deficit, heat stress, nutritional limitations) reduce four ovule frequency by impairing ovule initiation or causing ovule abortion, shifting distributions toward three and two ovule ovaries. Post-fertilization berry development from four ovule ovaries involves potential fertilization of all four ovules, though typical fertilization success rates (50-70% of ovules successfully fertilized) mean that four ovule ovaries typically produce berries with 2-3 seeds rather than 4 seeds, with 4-seed berries representing upper range outcomes requiring successful fertilization of all available ovules.
Four_Ovule_Ovary represents the upper end of normal ovule number range and is distinguished from Two_Ovule_Ovary (lower end of range) and from three ovule ovaries (intermediate, not specifically classified in current hierarchy). The four ovule condition is common in many cultivars and represents maximum typical seed number potential. Understanding four ovule versus lower ovule number ovaries is relevant for predicting maximum seed number, for understanding cultivar differences in reproductive investment, and for assessing berry size determinants. Four ovule ovaries are compatible with high fruit set and produce berries with maximum seed number potential when fertilization is successful, potentially contributing to larger berry size through seed-hormone effects on berry growth.
Vier-Eizellen-Ovar
Vier-Ovulum-Fruchtknoten
four-seed-potential ovary
maximum-ovule ovary
viereiiger Fruchtknoten
Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 94-98 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 104-108.
Four Ovule Ovary
Pinot Noir "FPMS 2A" (now FPS 02A) bears an FPMS_Clone_Role designation in older California wine literature, with the historic terminology identifying material distributed during the 1970s-1990s period when FPMS established certified clone availability for the expanding premium wine industry. Chardonnay selections described in 1980s research as "FPMS clones" serve in an FPMS_Clone_Role context, the historic designation linking to UC Davis's continuous clean plant program despite subsequent renaming. Cabernet Sauvignon material planted in 1990s Napa vineyards and documented as "FPMS certified" bears an FPMS_Clone_Role reflecting the terminology in use at planting, though the program continues today as FPS.
An American clone role that inheres in clones historically associated with the Foundation Plant Materials Service (FPMS), the predecessor designation for UC Davis's grapevine clean plant program before its reorganization and renaming as Foundation Plant Services (FPS) in the late 1990s. FPMS clones were processed through the same institutional infrastructure that continues as FPS, with the FPMS designation appearing in older literature, wine labels, and industry references from the program's earlier decades (1958-1990s). The FPMS terminology remains historically significant for identifying material distributed during the formative period of California's modern wine industry, when FPMS protocols established the foundations for certified clean plant distribution that continues under the FPS name. Material identified as "FPMS clone" or "FPMS selection" traces to the same program and physical location (UC Davis) as current FPS material, with the name change reflecting administrative reorganization rather than programmatic discontinuity. Understanding FPMS as the historical FPS designation enables interpretation of older clone references in the viticultural literature.
FPMS_Clone_Role captures the historical designation for what is now FPS, addressing the nomenclature continuity important for interpreting older literature and understanding vineyard history. The distinction between FPMS and FPS is administrative rather than substantive—the same institution, location, and programmatic mission continue under the updated name—but recognition of both designations enables proper interpretation of clone references across publication periods. Older California wineries may reference "FPMS clones" in their vineyard descriptions, with these references indicating FPS-equivalent material from earlier decades. The transition from FPMS to FPS coincided with program expansion and modernization (adoption of PCR testing, expanded screening panels), though the core mission of providing tested clean material remained constant.
FPMS clone
FPMS-Klon
Foundation Plant Materials Service clone
Foundation-Plant-Materials-Service-Klon
historic FPS clone
historischer FPS-Klon
Goheen AC, Cook JA (1959) Leafroll (red-leaf or rougeau) and its effects on vine growth, American Journal of Enology and Viticulture 10: 78-84 Weber E et al. (2002) The California grapevine certification program, American Journal of Enology and Viticulture 53: 130-134 Golino DA et al. (2017) Foundation Plant Services, HortScience 52: 1190-1194
FPMS Clone Role
Pinot Noir FPS 02A bears an FPS_Clone_Role as a tested selection maintained in the Foundation Vineyard, processed through comprehensive FPS protocols including HTS screening and available to licensed nurseries for certified propagation. Cabernet Sauvignon FPS 08 serves in an FPS_Clone_Role as one of the most widely distributed California Cabernet selections, its FPS processing ensuring virus-free status for growers prioritizing long-term vineyard health. Chardonnay FPS 04 (Old Wente) bears an FPS_Clone_Role representing the historically important California Chardonnay selection, maintained in tested condition through FPS protocols despite its 19th-century origins predating modern certification.
An American clone role that inheres specifically in clones processed, maintained, and distributed through Foundation Plant Services (FPS) at the University of California, Davis, the primary national grapevine clean plant center providing virus-tested propagation material to the American nursery industry. FPS clones have undergone comprehensive pathogen testing (biological indexing, ELISA, PCR, and increasingly high-throughput sequencing), with infected material subjected to elimination protocols (thermotherapy, meristem culture) before release. FPS maintains the Russell Ranch Foundation Vineyard and screenhouse collections housing tested mother plants of over 500 grape cultivars and selections, serving as the national repository for clean grapevine genetics. FPS clone designations typically use variety name followed by numeric code (e.g., Pinot Noir FPS 02A, Merlot FPS 03), with the code indicating the specific FPS accession. FPS operates within the National Clean Plant Network (NCPN) framework, coordinating with regional centers while maintaining its primary role for grapevine improvement. Material distributed through FPS carries documented traceability, testing history, and health certification valued by quality-focused nurseries and growers.
FPS_Clone_Role identifies the specific institutional pathway that represents the gold standard for clean grapevine material in the United States. FPS's combination of comprehensive testing (including adoption of HTS for broad-spectrum pathogen detection), pathogen elimination expertise, secure mother plant maintenance, and documented distribution creates a certification system comparable to European programs while serving a geographically dispersed industry. The FPS designation has become a market signal "FPS-certified" or "Davis clone" carries quality connotations commanding premium positioning. FPS's public mission (as a University of California service program) distinguishes it from commercial nurseries, with its focus on providing tested material rather than maximizing propagation sales. Recent FPS investments in HTS technology have positioned the program at the forefront of pathogen detection, identifying novel viruses that conventional testing would miss.
Davis clone
FPS clone
FPS-Klon
Foundation Plant Services clone
Foundation-Plant-Services-Klon
UC Davis clone
UC-Davis-Klon
Golino DA et al. (2017) Foundation Plant Services, HortScience 52: 1190-1194 Al Rwahnih M et al. (2015) High-throughput sequencing for grapevine pathogens, Phytopathology 105: 758-763 Rowhani A et al. (2018) Grapevine virus testing protocols, UC Davis FPS Foundation Plant Services (2020) FPS Grape Program Annual Report
FPS Clone Role
Free SO₂ for Sauvignon Blanc from Marlborough (2024 vintage, at bottling) measures 35 mg/L by aspiration-oxidation, corresponding to 0.82 mg/L molecular SO₂ at the wine pH of 3.15 — well above the 0.6 mg/L molecular SO₂ target considered adequate for microbial stability in low-pH white wines, and providing sufficient antioxidant protection for the 3-5 year cellaring window intended for this style. Free SO₂ for Cabernet Sauvignon from Napa Valley (2022 vintage, at bottling) records 28 mg/L, yielding only 0.38 mg/L molecular SO₂ at the wine pH of 3.65 — below the 0.6 mg/L target, prompting the winemaker to increase the SO₂ addition by 8 mg/L at bottling to achieve adequate microbial protection at the higher pH. Free SO₂ for Grüner Veltliner from Kamptal (2024 vintage) documents 30 mg/L at bottling, calculated to maintain >25 mg/L free SO₂ through 18 months of bottle storage given the expected binding rate of approximately 2-3 mg/L per month.
Free sulfur dioxide is a wine analysis data entity that records the concentration of sulfur dioxide present in wine in its unbound, antimicrobially and antioxidantly active forms — comprising molecular SO₂ (the biocidal species), bisulfite ion (HSO₃⁻, the antioxidant and carbonyl-binding species), and sulfite ion (SO₃²⁻) in pH-dependent equilibrium — determined by aspiration-oxidation (Rankine method), iodometric titration (Ripper method), or flow injection analysis, expressed in milligrams per liter (mg/L), and representing the immediately available protective capacity of sulfur dioxide against both microbial spoilage and oxidative degradation in the finished wine.
Free sulfur dioxide management is one of the most technically demanding aspects of wine quality control because the protective efficacy of SO₂ is entirely governed by pH through the molecular SO₂ equilibrium — at wine pH, free SO₂ exists predominantly as bisulfite ion (>90%) with only a small fraction in the biocidally active molecular SO₂ form, and this fraction decreases dramatically with increasing pH. The relationship is quantified by the Henderson-Hasselbalch equation applied to the SO₂/bisulfite equilibrium (pKa₁ = 1.77 at 20°C): at pH 3.0, approximately 6% of free SO₂ is molecular, at pH 3.3 approximately 3.5%, at pH 3.5 approximately 2%, and at pH 3.8 approximately 1% — meaning that a wine at pH 3.8 requires approximately 6× the free SO₂ concentration of a wine at pH 3.0 to achieve the same molecular SO₂ level and equivalent microbial protection. The target molecular SO₂ concentration of 0.5-0.8 mg/L is considered sufficient for microbial stability in most wines, with higher targets (0.8-1.0 mg/L) recommended for wines with residual sugar or elevated pH. The aspiration-oxidation method (aerating wine sample into hydrogen peroxide and titrating the resulting sulfuric acid) is considered the most accurate reference method, as the Ripper iodometric titration overestimates free SO₂ by 5-15 mg/L in red wines due to anthocyanin bleaching interference. Free SO₂ decreases continuously during wine storage through oxidative consumption and progressive binding to acetaldehyde, glucose, and other carbonyl compounds, necessitating periodic monitoring and supplementation to maintain protective levels. The natural and minimal-intervention winemaking movement's rejection or reduction of SO₂ use creates wines with minimal oxidative and microbial protection, demanding impeccable cellar hygiene and rapid consumption as compensating measures.
active SO2
aktives SO₂
available SO2
free SO2
free sulfite
free sulfur dioxide content
freie schweflige Säure
freies SO₂
freies Schwefeldioxid
verfügbares SO₂
OIV (2021) Compendium of International Methods of Analysis, Method OIV-MA-AS323-04B (free sulfur dioxide by aspiration) Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2, 2nd Ed, Wiley, pp.: 223-250 Waterhouse AL et al. (2016) Understanding Wine Chemistry, Wiley, pp.: 178-210 Rankine BRC, Pocock KF (1969) Alkalimetric determination of sulphur dioxide in wine. Australian Wine, Brewing and Spirit Review 88(4): 40-44
Free SO2
Merlot ovaries at BBCH 59-71 (flower buds swollen through fruit set) exhibit characteristic free superior organization with ovary wall composed entirely of carpel-derived tissues visible in anatomical sections as distinct epidermis, mesocarp precursor parenchyma, and inner epidermis, clearly demarcated from receptacle tissues at the ovary base, with no fusion zones or intermediate tissues indicating ovary-receptacle integration. Cabernet Sauvignon berry development from BBCH 71 through BBCH 89 demonstrates the functional advantages of free superior ovary, with berry (transformed ovary) expanding from 1 mm to 12-15 mm diameter over 90-110 days proceeding independently as other floral organs (sepals, petals, stamens) senesce and abscise by BBCH 73-75, leaving the free berry attached only by the pedicel without constraints from persisting perianth or receptacle tissues. Syrah ovary-receptacle boundary anatomical studies reveal free superior condition through abrupt transitions in cellular organization at the ovary base, with carpel epidermis and parenchyma showing distinct morphology from receptacle epidermis and vascular tissues, and with vascular bundles from receptacle entering ovary base as discrete strands without extensive lateral fusion creating intermediate vascular zones. Comparative morphology studies demonstrate that Vitis free superior ovaries contrast with inferior ovaries in Rosaceae (apple, pear) where receptacle tissues fuse with and enclose ovary tissues contributing to fruit flesh, and with Cucurbitaceae (melon, cucumber) where ovary fuses with floral tube and receptacle creating inferior ovaries that develop into fruits with complex tissue origins, highlighting the distinctive free superior condition as a defining and functionally significant Vitaceae characteristic.
A free superior ovary is a superior ovary that is completely unfused with receptacle, floral tube, or perianth (sepal, petal) tissues, with ovary wall consisting entirely of carpel-derived tissues and with no fusion or adnation to other floral structures, representing the universal condition in Vitis where ovary development and post-fertilization expansion proceed completely independently of other floral organs. Free superior ovaries are characterized by= (1) ovary wall comprising exclusively carpel-derived tissues without any fusion or integration with receptacle tissues, evident in anatomical sections showing clear cellular and vascular boundaries between ovary tissues and receptacle tissues (2) complete absence of floral tube or hypanthium (cup-like structure formed from fusion of perianth and/or receptacle tissues enclosing or fused with ovary) which characterizes inferior or partially inferior ovary conditions in other plant families (3) ovary attached to receptacle only at a discrete basal insertion point, with lateral and apical ovary surfaces free and not fused to any other structures (4) post-fertilization ovary expansion occurring independently without requiring coordinated growth of fused receptacle or perianth tissues, allowing rapid and unconstrained berry development. The free superior condition represents the ancestral angiosperm ovary organization and is retained in Vitis without evolutionary modifications toward fusion with other floral structures. This free condition contrasts with= (1) inferior ovaries in families like Rosaceae, Cucurbitaceae, or Asteraceae where ovary is fused with receptacle and/or floral tube tissues with perianth appearing to arise from ovary apex (2) partially inferior or half-inferior ovaries where ovary is partially fused with receptacle or floral tube (3) superior but adnate ovaries in some taxa where ovary remains above other organs but shows partial fusion with floral tube or receptacle tissues. The free superior condition in Vitis creates several functional and developmental advantages= ovary development proceeds autonomously from floral bud stages through anthesis to fruit maturation without developmental coordination requirements with receptacle or perianth tissues post-fertilization ovary expansion is mechanically unconstrained by fusion with other tissues, allowing rapid berry growth (diameter increasing 10-25 fold from anthesis to maturity over 60-120 days depending on cultivar and conditions) berry tissue composition reflects ovary tissue organization exclusively, with all berry structures (skin, flesh, seeds) derived from carpel tissues without receptacle contribution. The free superior ovary allows independent timing of ovary versus other organ development, evident in= early ovary development (BBCH 51-54) proceeding while other organs are still initiating ovary achieving developmental maturity (BBCH 59) coordinated with stigma receptivity independent of stamen or perianth maturation timing post-fertilization ovary expansion (BBCH 71-89) proceeding while other floral organs senesce and abscise, with berry development continuing long after sepals, petals, and stamens have disappeared. Anatomical studies of ovary-receptacle boundaries reveal the free condition through= distinct cellular organization changes at the ovary base where carpel tissues transition to receptacle tissues vascular tissue organization showing carpel vascular bundles connecting to but not fusing extensively with receptacle vascular system absence of intermediate tissue zones showing mixed carpel-receptacle cellular characteristics that would indicate fusion. The free superior ovary is so universal in Vitis that it represents the standard condition against which rare developmental abnormalities can be assessed, with any deviation from free superior organization (such as abnormal fusion with receptacle or retention of perianth tissues fused to ovary) representing significant developmental disruption rather than normal variation.
Free_Superior_Ovary emphasizes the unfused nature of superior ovary organization and is distinguished from Elevated_Superior_Ovary (which emphasizes the raised positioning) though these are overlapping rather than mutually exclusive categories. The free condition is universal in normal Vitis flowers and represents the ancestral ovary organization. Understanding the free superior condition is important for comparative floral morphology, for interpreting the exclusively ovary-derived nature of berry tissues, and for appreciating the developmental independence of ovary from other floral organs that allows unconstrained post-fertilization berry expansion.
freier oberständiger Fruchtknoten
independent ovary
unfused superior ovary
unfusioniertes Ovar
unverschmolzener Fruchtknoten
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 92-95 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109.
Free Superior Ovary
In Traminer berries in Südtirol, Italy during BBCH 85-89, free volatile production maintains unconjugated monoterpene content increasing from 0.15 to 0.95 mg/L over 38 days, representing 12% of total monoterpenoid content at harvest, with free linalool at 0.42 mg/L and free geraniol at 0.28 mg/L providing immediate floral-rose impact detectable in berry tasting, while the free-to-bound ratio remains relatively stable at 0.11-0.14 throughout ripening indicating balanced biosynthesis and conjugation rates under alpine conditions averaging 21°C.
Free volatile production describes the generation and maintenance of unconjugated, immediately aromatic volatile compounds in Vitis vinifera berries during ripening, wherein biosynthetic pathway outputs remain in their free, non-glycosylated state contributing directly to the perceptible aroma of intact berries and freshly crushed grape juice without requiring enzymatic or chemical transformation for aromatic expression. This process represents the fraction of total volatile biosynthesis that escapes conjugation to precursor forms, typically constituting 5-15% of total monoterpenoid content in aromatic varieties and varying proportions of other volatile classes depending on compound-specific glycosylation efficiencies and biosynthetic rates relative to conjugation capacity. The biochemical basis of free volatile production involves the kinetic competition between continued synthesis adding volatiles to the free pool and glycosyltransferase-mediated conjugation removing them, with free volatile concentrations representing the steady-state balance between these opposing processes rather than absolute accumulation without loss to conjugation pathways. Cellular mechanisms influencing free volatile production include subcellular compartmentalization separating sites of volatile synthesis from glycosyltransferase localization, membrane permeability determining volatile retention versus volatilization losses, and the relative expression levels of biosynthetic versus conjugating enzymes that establish the partitioning ratio between free and bound pools. The sensory significance of free volatile production reflects the immediate aromatic impact of these compounds in berry tasting and early-stage juice evaluation, with free volatiles providing the first olfactory impression encountered during harvest assessment and initial winemaking stages before precursor hydrolysis contributes additional aromatic complexity. Cultivar differences in free volatile production reflect genetic variation in both biosynthetic capacity and conjugation efficiency, with some aromatic varieties maintaining relatively high free-to-bound ratios of 0.15-0.25 while others show lower ratios of 0.05-0.10 despite equivalent total volatile content, suggesting variety-specific regulation of the partitioning between free and conjugated pools. Environmental factors including temperature and humidity influence free volatile production through effects on volatilization rates that deplete the free pool, with high temperatures and low humidity accelerating volatile loss and potentially reducing free volatile concentrations even when biosynthetic rates remain constant, necessitating consideration of environmental conditions when interpreting free volatile measurements during ripening assessment.
Free_Volatile_Production is subclass of Volatile_Compound_Synthesis representing unconjugated aromatic volatile generation. Distinguishes from Bound_Aroma_Accumulation which involves conjugated forms. Sibling to Ester_Formation and Aldehyde_Synthesis. Typically 5-15% of total monoterpenoids remain free. Provides immediate aromatic impact. Balance between synthesis and conjugation determines free pool size.
free aroma generation
freie Aroma-Erzeugung
freie Volatil-Produktion
immediate volatile production
non-glycosylated volatile formation
unconjugated volatile synthesis
unkonjugierte Volatil-Synthese
unmittelbare Volatil-Bildung
Wilson B et al (1984) Determination of free and bound monoterpenes in Muscat grapes. American Journal of Enology and Viticulture 35: 95-100 Marais J (1983) Terpenes in the aroma of grapes and wines. South African Journal of Enology and Viticulture 4: 49-58 Keller M (2020) The Science of Grapevines 3rd Ed pp 290-295
Free Volatile Production
In Cabernet Franc vines in the Loire Valley, freeze tolerance development progresses through November and December with bud hardiness increasing from -10°C following initial acclimation to -15°C after exposure to temperatures of 5-8°C during early November, reaching -20°C following late November temperatures of 0-3°C, and achieving maximum hardiness of -23°C by late December following exposure to minimum temperatures of -5 to -8°C during early winter.
Freeze tolerance development in Vitis vinifera represents the progressive phase of cold acclimation during which grapevine tissues acquire substantial resistance to freezing stress through accumulated changes in cellular composition and structure that enable survival of ice formation in extracellular spaces. This phase follows initial cold acclimation and proceeds through autumn and early winter as temperatures progressively decline, with each exposure to lower temperatures potentially inducing additional hardiness gains until maximum cold tolerance is achieved. The progressive nature of freeze tolerance development reflects the requirement for multiple rounds of gene expression and metabolic adjustment at successively lower temperatures, with each stage building upon the foundation established during previous acclimation. Cellular mechanisms underlying freeze tolerance include controlled ice formation in extracellular spaces rather than within cells, facilitated by ice nucleation active proteins and cell wall properties that favor extracellular ice crystal growth, combined with cellular dehydration tolerance as water moves osmotically to growing extracellular ice masses. The accumulation of cryoprotective molecules reaches maximum levels during this phase, with proline concentrations increasing 5-10 fold in cold-hardy tissues, raffinose family oligosaccharides accumulating to several percent of dry weight, and dehydrin proteins becoming major components of the soluble protein fraction. Membrane stabilization through lipid remodeling continues during freeze tolerance development, with the ratio of unsaturated to saturated fatty acids in membrane phospholipids increasing to maintain appropriate fluidity at sub-zero temperatures and prevent phase transitions that would damage cellular membranes. The extent of freeze tolerance development varies among tissue types within the vine, with primary bud tissues typically achieving hardiness of -20 to -25°C in well-adapted cultivars, secondary buds slightly hardier at -25 to -27°C, and cane phloem and xylem parenchyma achieving -25 to -30°C depending on cultivar genetics and acclimation conditions. The rate of freeze tolerance development depends on both genetic factors determining the capacity for hardiness and environmental factors including temperature regime, vine water status, and carbohydrate availability.
Freeze_Tolerance_Development represents the progressive hardening phase within Temperature_Acclimation following Cold_Acclimation_Start. Distinguished from initial acclimation by the requirement for exposure to progressively lower temperatures and the substantially greater hardiness gains achieved. Duration spans 6-10 weeks through autumn into early winter. Produces major hardiness gains from approximately -10°C to -20 to -25°C depending on cultivar. Key mechanisms include cryoprotectant accumulation, membrane remodeling, and controlled extracellular ice formation tolerance.
Frosthärte-Aufbau
Frosttoleranz-Entwicklung
deep cold acclimation
freeze hardiness acquisition
frost tolerance development
progressive Kälteabhärtung
progressive cold hardening
tiefe Kälteakklimatisierung
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Mills LJ, Ferguson JC, Keller M (2006) Cold-hardiness evaluation of grapevine buds and cane tissues. American Journal of Enology and Viticulture 57: 194-200 Wisniewski M et al (2003) Observations of ice nucleation and propagation in plants using infrared thermography. Plant Physiology 133: 852-858
Freeze Tolerance Development
Freezing point determination for Riesling dormant buds in Rheinhessen using differential thermal analysis reveals high-temperature exotherm at -5.2°C representing extracellular ice formation in bud scales and low-temperature exotherm at -21.3°C representing loss of supercooling in primordial tissues, with the 16°C separation between these events defining the temperature range where buds survive through supercooling avoidance before lethal intracellular freezing occurs at the low-temperature exotherm.
Freezing point quality is a critical temperature threshold quality that represents the temperature at which ice crystal formation initiates within grapevine tissues, marking the transition from supercooled liquid state to ice-containing state that damages cells through mechanical disruption, osmotic stress from extracellular ice formation, and intracellular ice crystal damage. Within the BFO framework, this quality characterizes the specific temperature threshold where phase transition from liquid to solid water occurs within plant tissue, with the realized freezing point depending on tissue solute concentration providing freezing point depression, the presence of ice-nucleating agents, and the rate of cooling affecting supercooling capacity. The ontological significance lies in freezing point defining the absolute lower limit of tissue survival without ice formation, distinguishing from frost damage thresholds that may occur at warmer temperatures when extracellular ice formation causes osmotic damage even without intracellular freezing. Grapevine tissue freezing points vary with solute concentration, ranging from approximately -2°C for dilute xylem sap to -5°C for sugar-concentrated berry mesocarp, with cold-acclimated dormant tissues accumulating cryoprotective solutes that depress freezing point by 2-4°C compared to non-acclimated tissues. The distinction between supercooling and equilibrium freezing is critical, as grapevine buds can supercool to -15 to -25°C before ice nucleation occurs, with the lethal temperature determined by the low-temperature exotherm marking loss of supercooling rather than the equilibrium freezing point. Measurement approaches include differential thermal analysis detecting ice formation exotherms, direct thermocouple observation of tissue temperature arrest during freezing, and calculated depression from solute concentration using the cryoscopic constant.
Freezing_Point_Quality specifically addresses the ice formation threshold, distinguishing from broader frost damage thresholds that may occur at warmer temperatures through osmotic or other mechanisms. This quality varies with tissue composition and acclimation status, requiring seasonal context for accurate specification. The class encompasses both equilibrium freezing point depressed by solutes and nucleation temperature determined by supercooling capacity, with the physiologically relevant threshold depending on cooling rate and tissue freeze-avoidance mechanisms.
Eiskristallisationstemperatur
Frostpunkt
Gefrierpunkt
Gefriertemperatur
crystallization point
frost point
ice nucleation temperature
tissue freezing temperature
Mills LJ et al. (2006) Timing of low temperature exposure affects cold hardiness and bud dormancy of Vitis vinifera L. cv. Cabernet Sauvignon. American Journal of Enology and Viticulture 57(4): 412-421 Pierquet P, Stushnoff C (1980) Relationship of low temperature exotherms to cold injury in Vitis riparia Michx. American Journal of Enology and Viticulture 31(1): 1-6
Freezing Point Quality
Spätburgunder (Pinot Noir) clone 1 Fr bears a Freiburg_Clone_Role as the historically dominant Pinot Noir selection in Baden, selected before French Dijon clones became internationally available, with adaptation to Baden's continental climate and production traditions. Grauburgunder (Pinot Gris) clone 2 Fr serves in a Freiburg_Clone_Role as a quality-focused selection for Baden's important Grauburgunder production, with documented wine quality supporting the variety's premium positioning in German markets. Weißburgunder (Pinot Blanc) clone 2 Fr bears a Freiburg_Clone_Role, supporting Baden's significant Pinot Blanc production with selections adapted to local conditions and market requirements.
A German clone role that inheres specifically in clones selected and registered by the Staatliches Weinbauinstitut Freiburg (WBI, State Viticulture Institute Freiburg), located in Baden (southwestern Germany), with selections designated by the "Fr" suffix (e.g., Spätburgunder 1 Fr, Grauburgunder 2 Fr). Freiburg clones reflect the institution's focus on Baden's warmer viticultural conditions (Germany's sunniest wine region), with particular strength in Burgundy variety selections (Spätburgunder/Pinot Noir, Grauburgunder/Pinot Gris, Weißburgunder/Pinot Blanc) suited to the region's climate. WBI Freiburg operates comprehensive clone evaluation programs including long-term field trials, wine quality assessment, and virus-free mother plant maintenance, serving the substantial Baden wine industry as well as contributing to national clone resources. Freiburg's Burgundy variety expertise has positioned its selections for adoption beyond Germany in regions sharing similar climatic conditions—Austria, northern Italy, and other warm-margin cool-climate zones. The institution also conducts resistance breeding programs developing disease-resistant varieties complementing its clone selection work.
Freiburg_Clone_Role identifies WBI Freiburg selections within the German institutional framework, with particular significance for Burgundy varieties and warmer German production conditions. Baden's position as Germany's warmest major wine region oriented Freiburg's programs toward varieties and clone characteristics suited to more generous growing conditions—contrasting with Geisenheim's cool-climate Rheingau focus. Freiburg's Burgundy variety strength reflects Baden's historical connection to Alsace and the broader Upper Rhine viticultural tradition, with Spätburgunder particularly important to regional identity. The institution's resistance breeding programs (developing PIWI varieties—pilzwiderstandsfähige Rebsorten) increasingly complement clone selection, addressing sustainability concerns through genetic improvement beyond V. vinifera limitations.
Baden clone
Baden-Klon
Fr clone
Fr-Klon
Freiburg clone
Freiburg selection
Freiburg-Klon
Freiburger Selektion
WBI clone
WBI-Klon
Staatliches Weinbauinstitut Freiburg (2020) Klonenzüchtung Rebsorten Bundessortenamt (2020) Beschreibende Sortenliste Reben Becker H (1985) German viticulture and breeding, American Journal of Enology and Viticulture 36: 58-63 Regner F (2007) Genetic variation in grapevine clones, Blackwell
Freiburg Clone Role
Seyval Blanc (Seyve-Villard 5-276) bears a French_American_Hybrid_Role as one of the most successful white French hybrids, widely planted in England, eastern North America, and other challenging climates for its cold hardiness, disease tolerance, and neutral wine character suited to blending and sparkling production. Baco Noir (Baco 1) serves in a French_American_Hybrid_Role as a historically important red hybrid, its Folle Blanche × V. riparia parentage unique in combining a female V. vinifera parent with American pollen. Chambourcin (Joannes-Seyve 26-205) bears a French_American_Hybrid_Role as a complex hybrid with strong color and increasingly recognized wine quality, important both commercially and as a parent in contemporary resistance breeding.
An interspecific hybrid role that inheres in grapevine varieties developed primarily by French breeders in the late 19th and early 20th centuries, crossing American Vitis species (V. labrusca, V. riparia, V. rupestris, V. lincecumii, V. aestivalis) with V. vinifera to create disease-resistant "direct producer" varieties capable of producing wine without grafting to resistant rootstocks. French-American hybrids (also termed French hybrids or producteurs directs) emerged as a response to phylloxera, downy mildew, and powdery mildew epidemics, with French breeders (Seibel, Seyve-Villard, Couderc, Baco, Landot, Ravat, among others) creating thousands of numbered selections from interspecific crosses. These hybrids achieved widespread planting in France (peaking at over 400,000 hectares in the 1950s) and internationally, offering viticultural simplicity (own-rooted cultivation without grafting) and disease tolerance. French policy subsequently discouraged hybrid cultivation to protect traditional V. vinifera appellations, reducing French hybrid plantings dramatically from the 1960s onward. Legacy French-American hybrids remain significant in regions with challenging disease pressure (eastern North America, humid climates) and as germplasm sources for contemporary resistance breeding.
French_American_Hybrid_Role captures the historically significant breeding effort that produced direct producer alternatives to grafted V. vinifera. The French hybrid era (roughly 1880-1960) generated enormous genetic diversity—over 10,000 named selections from major breeders—most of which were abandoned but some of which persist commercially or in breeding programs. French hybrids' commercial decline reflects both quality limitations (many early hybrids produced wines with foxy or herbaceous characters) and regulatory discrimination (French appellation law excluded hybrids from quality wine designations, subsequently influencing EU-wide policy). Contemporary reassessment recognizes that select French-American hybrids produce acceptable wines while offering disease resistance, with varieties like Chambourcin serving as parents in second-generation PIWI breeding. The "French" designation reflects developer nationality rather than geographic origin of all genetic components, which typically include substantial American species contributions.
Direktträger
French hybrid
French-American hybrid
Hybride producteur
direct producer hybrid
französisch-amerikanische Hybride
französische Hybride
producteur direct
Reisch BI et al. (2012) Grape breeding in the United States, CRC Press pp.: 171-203 Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Bouquet A (2011) Grapevine breeding in France, in Adam-Blondon AF et al. (eds) Genetics, Genomics, and Breeding of Grapes, CRC Press
French American Hybrid Role
Pinot Noir clone 115 bears a French_Clone_Role as the most widely planted of the ENTAV-INRA "Dijon clones," selected in Burgundy for small berries, loose clusters, and concentrated wine character—now distributed globally as the benchmark quality Pinot Noir selection. Chardonnay clone 76 serves in a French_Clone_Role with ENTAV registration documenting its selection for small-cluster character and concentrated must composition, widely adopted for premium Burgundy, Champagne, and New World Chardonnay production. Cabernet Sauvignon clone 337 bears a French_Clone_Role as one of the most planted Cabernet clones internationally, ENTAV evaluation documenting its balance of productivity and quality for Bordeaux and global Cabernet production.
A regional clone role that inheres in a grapevine clone registered within the French national clone certification system, administered by ENTAV-INRA (Établissement National Technique pour l'Amélioration de la Viticulture - Institut National de la Recherche Agronomique) and documented in the official Catalogue des Variétés et Clones de Vigne Cultivés en France. French clones are identified by variety name followed by numeric code (e.g., Pinot Noir 115, Chardonnay 76, Cabernet Sauvignon 337), with codes assigned sequentially within each variety as selections complete the registration process. The French clone system, established formally in 1962 with ENTAV's creation, pioneered systematic clonal selection integrating sanitary (virus elimination) and genetic (performance evaluation) dimensions, producing the world's most extensive registered clone catalog with over 1,000 approved scion and rootstock clones. French clones dominate international planting of major varieties, with ENTAV selections of Pinot Noir (Dijon clones), Chardonnay, Cabernet Sauvignon, Merlot, and Syrah planted across all major wine-producing countries. French certification distinguishes Certifié (laboratory-tested virus-free) and Standard grades within the registered clone framework.
French_Clone_Role captures the dominant international clone system, reflecting France's pioneering role in systematic clonal selection and the global distribution of French viticultural material. The ENTAV-INRA partnership (ENTAV for practical selection and distribution, INRA for research) created an integrated program combining virus elimination techniques with extended field and wine trials, producing clones with documented performance characteristics and certified health status. French clone dominance creates both opportunities (access to thoroughly evaluated selections) and concerns (genetic narrowing from widespread planting of few clones) contemporary French programs emphasize conservatory collections alongside commercial registration to preserve diversity. The numeric code system lacks inherent meaning—clone 115 was simply the 115th Pinot Noir registered—though wine culture has invested certain numbers with quality associations (the "Dijon clones" 113, 114, 115, 667, 777 became prestige selections despite similar origins to less famous registrations).
Catalogue clone
ENTAV clone
ENTAV-Klon
French clone
French registered clone
französisch registrierter Klon
französischer Klon
ENTAV-INRA (2007) Catalogue des Variétés et Clones de Vigne Cultivés en France Audeguin L et al. (1999) The results of 30 years of clonal selection in France, ISHS Acta Horticulturae 473: 69-76 Walter B, Martelli GP (1996) Clonal and sanitary selection of the grapevine, INRA Editions Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25
French Clone Role
Sugraone (Superior Seedless) bears a Fresh_Consumption_Table_Grape_Role as a premium early-season white seedless, with large berries (6-8 g), crisp muscat-influenced flavor, and attractive yellow-green color commanding premium early-market prices in Mediterranean and Chilean production for European retail. Flame Seedless serves in a Fresh_Consumption_Table_Grape_Role as the dominant early red seedless, with 4-5 g berries, bright red color, and firm crunchy texture providing the benchmark for supermarket red grape expectations. Cotton Candy (IFG Seven) bears a Fresh_Consumption_Table_Grape_Role as a proprietary flavor variety, where distinctive spun-sugar flavor and crisp texture command retail premiums of 2-3× standard seedless grapes, exemplifying the premium flavor-branded table grape category.
A table grape scion role that inheres in a grafted scion cultivar by virtue of its primary designation for immediate fresh consumption without processing, drying, or extended storage. Fresh consumption table grape scions are optimized for eating quality at the point of sale, including optimal sugar-acid balance for immediate palatability (typically 16-20°Brix, 0.4-0.7% titratable acidity, creating Brix/acid ratios of 25-35), crisp or crunchy texture (firm cell walls, high turgor), attractive visual appearance (bright color, intact bloom, well-filled clusters), and pleasant mouthfeel without astringency or excessive seediness. Fresh consumption varieties are distinguished from storage varieties (bred for extended shelf life) and from dual-purpose varieties used for both fresh consumption and raisin or juice production. The category encompasses seeded and seedless varieties across all color classes (white/green, red/pink, black/blue), with market preferences varying regionally. Quality assessment standards (USDA, EU, Codex) grade fresh table grapes by berry size, color uniformity, cluster fill, and defect tolerance.
Fresh_Consumption_Table_Grape_Role captures the primary utilization for most table grape cultivars, distinguishing immediate-consumption optimization from varieties bred for storage longevity or multi-purpose use. The category is subclassed by berry color (Red, White, Black), reflecting both genetic and market organization. Modern breeding has created an expanding array of proprietary "flavor grapes" (Cotton Candy, Moon Drop, Witch Fingers/Tear Drops, Gum Drop) protected by plant patents and marketed under trademarked names, transforming table grapes from commodity to branded product category. Organic and "residue-free" production has grown substantially, requiring varieties tolerant of reduced fungicide programs while maintaining cosmetic quality standards.
Esstraube
Frischtraube zum Direktverzehr
Frischverzehr-Tafeltraube
eating grape
fresh consumption grape
fresh table grape
Creasy GL, Creasy LL (2009) Grapes, CABI Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press Crisosto CH, Mitchell FG (2002) Postharvest handling systems: table grapes, in Postharvest Technology of Horticultural Crops 3rd Ed, UC ANR pp.: 357-363 Fidelibus MW et al. (2017) California Table Grape Commission Annual Report
Fresh Consumption Table Grape Role
Table grape quality standards for Red Globe specify minimum berry fresh mass of 10-12 g per berry and minimum cluster fresh mass of 500-700 g for premium grade fruit destined for export markets, achieved through aggressive crop load management (one cluster per shoot), gibberellic acid application (10 ppm at fruit set, 20-40 ppm at 8-12 mm berry diameter), and optimal irrigation maintaining soil moisture at 80-100% of field capacity throughout the cell expansion period from fruit set through véraison.
Fresh_Mass_Quality represents the total mass of a grapevine organ at the time of measurement including all contained water. This quality captures the organ in its living or recently harvested state before any drying or water loss has occurred, providing the practically relevant mass for yield determination, harvest logistics, and commercial transactions. Fresh mass is the standard measurement for grape yield, expressed as kg per vine, tonnes per hectare, or tons per acre, and forms the basis of most grape purchase contracts. Berry fresh mass typically ranges from 0.8-4.0 g depending on cultivar and environmental conditions, while cluster fresh mass ranges from 50-500 g for wine grapes. Fresh mass combines the contributions of structural dry matter and water content, with water typically constituting 70-85% of berry fresh mass at harvest. Because water content varies with vine water status, time of day, and weather conditions, fresh mass measurements exhibit greater variability than dry mass for the same underlying biological material. Fresh mass decreases under water deficit conditions as organs lose turgor and may lose water to the atmosphere or redistribute water to other plant parts. Conversely, fresh mass can increase transiently following irrigation or precipitation as tissues rehydrate. Fresh mass measurement requires prompt weighing after harvest to minimize evaporative losses, with analytical balances providing precision of 0.01-0.001 g for small samples.
Fresh_Mass_Quality is distinguished from Dry_Mass_Quality by including water content in the measurement. The relationship between fresh mass and dry mass is mediated by water content or its inverse, dry matter percentage. Fresh mass measurements should specify the time elapsed since harvest and storage conditions to account for potential water loss. For comparison across studies or conditions with varying water status, expressing data on a dry mass basis may be more appropriate. Berry fresh mass correlates highly with berry volume (r² greater than 0.95) but the relationship slope (density) varies with sugar concentration during ripening.
Erntemasse
Feuchtmasse
Frischgewicht
Frischmasse
fresh weight
harvest mass
undried mass
wet mass
Coombe BG and McCarthy MG (2000) Dynamics of Grape Berry Growth and Physiology of Ripening. Australian Journal of Grape and Wine Research 6(2): 131-135 Dai ZW et al. (2011) Ecophysiological, Genetic, and Molecular Causes of Variation in Grape Berry Weight and Composition: A Review. American Journal of Enology and Viticulture 62(4): 413-425
Fresh Mass Quality
Merlot vines at BBCH 63-65 (30-50% of flowers open, active bloom progressing) exhibit numerous freshly detached calyptras visible as yellowish-green caps suspended on stamens of recently opened flowers (10-20% of open flowers showing suspended calyptras at any given time), creating characteristic 'capped' appearance, with dislodgement typically occurring 30-90 minutes post-detachment under moderate morning air movement (5-8 km/h winds, rising temperatures from 18°C to 24°C). Sauvignon Blanc leaves during mid-bloom show accumulation of 8-15 freshly detached calyptras per 100 cm² leaf surface area on basal leaves positioned below inflorescences, with fresh yellowish-green coloration and intact cup-like structure allowing easy visual identification and providing qualitative confirmation of vigorous bloom progression. Cabernet Franc at BBCH 61-63 shows freshly detached calyptras suspended on stamens, recently fallen on rachis and pedicels within inflorescences, and accumulating on upper leaf surfaces, with fresh calyptra distribution patterns providing visual evidence of the acropetal bloom progression from basal toward apical inflorescence flowers. Research plots employing standardized bloom quantification collect freshly detached calyptras from 50 cm × 50 cm collection trays positioned 15 cm below inflorescences, counting accumulated fresh calyptras daily during bloom period to generate quantitative bloom intensity curves showing peak bloom timing and duration across cultivars and environmental treatments.
A freshly detached calyptra is a detached calyptra that has recently separated from the flower (typically within the past few minutes to several hours) and retains relatively intact structure, recognizable morphology, and characteristic yellowish-green to light green coloration, often still suspended temporarily on the extended stamens or recently fallen onto nearby surfaces, representing the immediate post-separation state when calyptra identity and origin are most clearly evident and when calyptras are most useful as visual indicators of active bloom. Freshly detached calyptras are characterized by= (1) intact cap-like or cup-like structure with the five-parted petal fusion pattern clearly visible, showing minimal physical degradation or fragmentation (2) coloration of yellowish-green, light green, or light tan, reflecting recent senescence but not advanced desiccation or decomposition (3) tissue texture that remains relatively pliable or soft rather than brittle, though beginning to lose turgor and moisture (4) in many cases, temporary suspension on top of the extended stamen filaments where the calyptra has been lifted off but not yet dislodged, creating the characteristic appearance of flowers with cap-like structures perched on stamens (5) when fallen, calyptras appearing fresh and conspicuous on leaf surfaces or inflorescence structures rather than degraded or difficult to distinguish. The freshly detached state typically persists for several hours to 1-2 days depending on environmental conditions, with duration influenced by= temperature effects on desiccation rate (higher temperatures accelerating tissue drying and transition to degraded state) humidity effects on tissue moisture retention (higher humidity prolonging fresh appearance, lower humidity accelerating desiccation) wind and rain effects on physical integrity (wind and rain accelerating fragmentation, removal, or degradation). Freshly detached calyptras suspended on stamens represent a transient but visually striking phase, with suspension duration varying from minutes (in windy conditions or with vigorous air movement) to several hours (in calm conditions), before wind, gravity, or stamen movement causes dislodgement and fall. The suspended freshly detached calyptra creates a characteristic appearance of flowers with yellowish-green caps sitting atop the radiating stamens, a distinctive visual signature of recently opened flowers at peak anthesis that is commonly noted in phenological descriptions and photographic documentation of grapevine bloom. Freshly detached calyptras are the most useful detached calyptra stage for bloom assessment and documentation, as= (1) their intact structure and fresh appearance make them easily recognizable and distinguishable from other floral debris (2) their presence on stamens or as recent fallen structures directly indicates that flowers have just opened, providing real-time evidence of active anthesis (3) their conspicuous appearance on leaf surfaces allows rapid visual assessment of bloom progression without requiring close flower examination (4) their concentration and distribution can provide qualitative or semi-quantitative indicators of bloom intensity and spatial patterns within the canopy. Research applications sometimes employ systematic collection of freshly detached calyptras from defined leaf surface areas or from collection trays positioned beneath inflorescences, with counts of fresh calyptras per unit area or per unit time providing quantitative metrics of bloom timing, rate, and total flower number, complementing visual phenological stage assessments. Freshly detached calyptras can also serve as sampling material for molecular or biochemical studies of petal senescence, abscission processes, or pathogen colonization dynamics, as the recently detached tissue retains RNA, proteins, and cellular organization better than degraded fallen calyptras.
Freshly_Detached_Calyptra represents the immediate post-separation state with intact structure and is distinguished from Fallen_Detached_Calyptra (degraded, desiccated, or fragmented structures from earlier-separated calyptras) by fresh appearance, intact morphology, and recent separation timing. The freshly detached stage is the most valuable detached calyptra state for practical bloom assessment and for research quantification, as the fresh appearance and intact structure facilitate reliable identification and counting. The characteristic suspended calyptra appearance (caps perched on stamens) is a widely recognized visual indicator of active grapevine bloom and is frequently featured in viticultural photography and educational materials illustrating anthesis. The duration of the freshly detached state (hours to 1-2 days) is sufficient for practical field observation but short enough that accumulation of fresh calyptras provides meaningful temporal resolution for tracking bloom progression.
frisch abgelöste Blütenhaube
frisch abgeworfene Kronblattkappe
kürzlich abgefallene Kelchhaube
newly detached petal cap
recently shed calyptra
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 102-103 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 95-96.
Freshly Detached Calyptra
Pinot Noir freshly harvested berries hand-picked in early morning (6-9 AM, berry temperature 14-18°C), transported 2 km to winery in small 20 kg bins, sorted and destemmed within 2.5 hours of first pick, show composition identical to on-vine assessment (24.2% sugars measured on vine, 24.1% in must, 6.8 g/L acidity both, pH 3.68 both), no fermentation signs, no oxidation, optimal aromatic profile with fresh red fruit notes, representing ideal freshly harvested state allowing maximum quality wine production. Sauvignon Blanc freshly harvested berries night-harvested (11 PM-3 AM, berry temperature 10-12°C maintaining cool state) for aromatic white production, whole-cluster pressed within 3 hours of harvest beginning, show fresh characteristic grapefruit and passionfruit aromatics at peak intensity, no oxidation (juice pale greenish-white), sugars and acidity exactly matching on-vine targets (21.5% sugars, 8.2 g/L acidity), demonstrating premium aromatic white production benefits from freshly harvested state maintaining volatile aromatic compounds at peak levels. Chardonnay freshly harvested berry cooling protocol= harvest at 18-22°C berry temperature, immediate placement in refrigerated transport maintaining 10-12°C during 1 hour transport, processing at 4 hours post-harvest while maintaining cool temperature throughout, showing <0.05% sugar loss, no acid change, no oxidation, no fermentation, optimal quality retention demonstrating that freshly harvested status can be maintained for several hours through temperature control. Thompson Seedless table grapes freshly harvested for fresh market= hand-harvested in cool morning into specialized table grape containers, field-packed into shipping boxes, immediately placed in cold storage at 0-2°C within 2 hours of harvest, shipped via refrigerated transport, arriving at retail within 48 hours while maintaining freshly harvested quality characteristics (firm texture, green stems, fresh appearance, no fungal growth), demonstrating table grape cold chain maintaining fresh-harvested quality through rapid cooling and temperature maintenance. Cabernet Sauvignon freshly harvested berry quality comparison= Block A processed at 2.5 hours post-harvest (freshly harvested state) versus Block B delayed to 10 hours post-harvest due to processing capacity limits, showing Block A wine with better color (15% higher anthocyanin extraction), fresher fruit character (sensory evaluation 89 versus 85 points), and lower volatile acidity (0.32 versus 0.45 g/L), demonstrating quality advantages of freshly harvested state versus delayed processing even with moderate delays.
A freshly harvested berry is a harvested berry in the immediate post-harvest period (typically 0-6 hours post-detachment), characterized by minimal post-harvest quality changes, composition very similar to on-vine state, and optimal condition for processing or fresh market delivery, representing the preferred state for high-quality wine production or fresh consumption. Freshly harvested berries are characterized by= (1) harvest timing typically 0-6 hours since vine detachment with minimal opportunity for post-harvest changes (2) composition essentially unchanged from on-vine state with sugars, acids, pH, color, and flavor compounds at levels achieved on vine (3) minimal respiration effects with sugar loss typically <0.1% and acid loss minimal (4) no spontaneous fermentation with berries showing no microbial activity if harvested cleanly (5) optimal color retention with anthocyanins stable and no oxidative browning (6) optimal aromatic profile with volatile compounds at peak and no off-odor development (7) physical condition optimal with minimal crushing or damage if handled appropriately. Freshly harvested berries maintain quality through= prompt harvest-to-processing timeline minimizing time for degradative changes cool berry temperatures (ideally <15-18°C) slowing metabolic activity gentle handling preventing physical damage and juice release clean harvest excluding diseased or damaged fruit appropriate containers preventing crushing during transport. Freshly harvested berries for wine production show ideal processing conditions= sugars at target levels achieved on vine without respiration losses acids at optimal levels without post-harvest degradation no oxidation requiring minimal sulfur dioxide additions no spontaneous fermentation allowing controlled inoculation optimal color extraction potential with anthocyanins in native form without degradation optimal aromatic potential with volatile compounds at peak concentration. Freshly harvested berries for table grape market show ideal fresh condition= firm texture with no softening beyond on-vine state fresh appearance without shriveling or browning stems green and firmly attached bloom (waxy cuticle) intact creating attractive appearance no fungal growth or spoilage signs optimal eating quality with sugar and acid at desired levels. Freshly harvested berries require rapid processing or market delivery to maintain fresh state= wine grapes ideally processed within 2-6 hours of harvest table grapes ideally packed and cooled within 2-4 hours cooling to appropriate storage temperature (wine grapes 10-15°C, table grapes 0-2°C) if immediate processing not possible careful handling throughout maintaining berry integrity. Harvest logistics influence freshly harvested berry window= proximity of vineyard to winery affects achievable timeline with close vineyards allowing 1-3 hour harvest-to-crush versus distant vineyards requiring 4-8+ hours harvest method affects damage and timeline with hand harvest allowing selective gentle picking versus machine harvest producing immediate damage requiring prompt processing harvest capacity affects freshly harvested berry proportion with high-capacity harvest and processing maintaining fresh state for larger volumes versus limited capacity creating processing delays.
Freshly_Harvested_Berry represents optimal post-harvest state (0-6 hours) and is distinguished from Post_Harvest_Berry (extended post-harvest period >6 hours) by minimal versus progressive post-harvest changes. Freshly harvested berries maintain quality closest to on-vine condition, representing the target state for high-quality production. Understanding freshly harvested berry characteristics and maintaining this state is essential for quality wine production, particularly for aromatic whites and premium reds, and for table grape fresh market quality.
Frisch-Lese-Traube
frisch geerntete Beere
immediately harvested berry
just-picked berry
unmittelbar geerntete Traube
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 265-275 Zoecklein, B.W. et al. (1995). Wine Analysis and Production, pp. 95-115.
Freshly Harvested Berry
Frost damage severity quality assessment following a radiation frost event (-6°C minimum) in Marlborough evaluated Sauvignon Blanc vineyards across topographic positions to quantify damage patterns and inform recovery management. Low-lying blocks in frost-prone valley floors showed severe frost damage severity quality with 90% of primary shoots exhibiting complete necrosis, secondary bud damage estimated at 60%, and projected yield loss exceeding 70% from the combined effects of primary shoot loss and reduced secondary shoot fruitfulness. Mid-slope blocks showed moderate frost damage severity quality with 45% primary shoot damage, minimal secondary bud injury, and projected yield loss of 30-40% recoverable through adjusted canopy management favoring surviving shoots. Upper-slope blocks with effective cold air drainage showed mild frost damage severity quality with only 15% primary shoot damage limited to the most advanced shoots in early-burst positions, with projected yield loss below 10% and no special management required. The frost damage severity quality gradient informed triage decisions allocating protective resources including portable wind machines and helicopter contracts to the highest-value blocks most vulnerable to subsequent frost events forecast for the following week.
Frost damage severity quality is a cold stress indicator quality that inheres in a grapevine organism and represents the extent and intensity of tissue injury resulting from exposure to freezing temperatures, encompassing the range of damage manifestations from minor injury affecting only the most cold-sensitive tissues through moderate damage affecting substantial portions of vegetative or reproductive organs to severe damage causing widespread tissue death with major consequences for current season productivity and potentially long-term vine health. This quality provides the diagnostic framework for assessing actual damage from frost events, recognizing that damage severity depends on the interaction between minimum temperature reached, duration of freezing exposure, rate of temperature decline, tissue type and developmental stage, and the acclimation status of affected organs at the time of frost occurrence. The ontological significance of frost damage severity quality lies in its role enabling quantification of frost event impacts for production planning, insurance assessment, and recovery management decisions, while also providing feedback for evaluating the effectiveness of frost protection measures and identifying sites or cultivars with inadequate frost tolerance for reliable production. The biological basis of frost damage severity encompasses the cellular injury mechanisms triggered by ice formation including direct mechanical damage from ice crystals, cellular dehydration as water moves to extracellular ice, and membrane disruption from the combined effects of dehydration and low temperature, with damage severity reflecting the proportion of cells experiencing lethal injury and their functional importance for vine productivity. The visual manifestations of frost damage vary with tissue type and injury severity, including water-soaked appearance immediately following thaw, subsequent browning and necrosis of affected tissues, and the characteristic patterns of damage reflecting differential tissue sensitivity within complex organs such as compound buds where primary buds may be killed while secondary buds survive. The spatial distribution of frost damage within vineyards reflects microclimate variation in minimum temperatures reached, with low-lying frost pockets showing more severe damage than elevated or sloping sites with better cold air drainage.
Frost_Damage_Severity_Quality is_a Cold_Stress_Indicator_Quality, serving as parent class for Spring_Frost_Damage_Quality, Autumn_Frost_Damage_Quality, and Winter_Freeze_Damage_Quality which address seasonally-specific damage patterns. This class addresses the overall extent of frost injury while child classes capture particular seasonal contexts. Scoring systems (percentage damage, yield loss projection) enable quantitative assessment. The spatial pattern of damage reflects microclimate variation useful for site characterization. Distinguished from Tissue_Freezing_Tolerance_Quality which addresses resistance capacity rather than actual damage.
Frostschaden-Schweregrad
Frostverletztungs-Schwere
Gefrierschaden-Intensität
Kälteschaden-Ausmaß
cold damage level
freeze damage severity
freezing injury intensity
frost injury extent
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Trought MCT et al. (1999) Vineyard variability in Marlborough, New Zealand: characterising variation in vineyard performance and options for management. Australian Journal of Grape and Wine Research 5:84-89 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 508-545
Frost Damage Severity Quality
The frost risk data for the Chablis wine region (Burgundy, 47.8°N) documents the acute spring frost vulnerability that defines this region's viticultural identity — the historical record (1950-2020) shows late frost events (minimum temperature below -2°C after April 1) occurring in 60-70% of years, with severe frost events (minimum below -4°C, sufficient to kill all emerged green tissue) in 25-30% of years. The frost damage incidence has increased since the 1990s despite overall warming, because the earlier budburst driven by warmer March temperatures (budburst advancing by approximately 13 days between 1970 and 2020) exposes the vulnerable green shoots to the late frost events that continue to occur in April with unchanged frequency — a climate change paradox where warming increases frost damage by decoupling the advancing phenology from the unchanged frost probability. The 2016, 2017, and 2021 frost events demonstrated the devastating economic impact — the April 2016 frost destroyed an estimated 50% of the Chablis crop, and the April 2021 event (three consecutive frost nights of -5 to -8°C following 10 days of >20°C temperatures that had advanced growth to the 3-4 leaf stage) caused the most widespread frost damage in French viticulture since 1991, affecting Chablis, Burgundy, the Loire, Bordeaux, and the Rhône.
A temperature data component that records the frequency, severity, timing, and duration of freezing temperature events that pose damage risk to grapevine tissues — documenting both spring frost risk (temperatures below -2 to -3°C during the post-budburst period when young green shoots are freeze-sensitive, potentially destroying the current season's crop) and winter frost risk (temperatures below -15 to -20°C during dormancy that can damage or kill woody trunk and cordon tissues, potentially destroying the vine itself) — with frost risk assessment based on historical temperature records, landscape frost mapping, and the relationship between frost probability and phenological timing at the specific site.
Frost_Risk_Data documents the temperature hazard that represents the single largest source of catastrophic crop loss in cool-climate viticulture — frost damage differs from other viticultural weather risks (drought, hail, disease) in that a single night of subfreezing temperatures can destroy the entire crop potential of a vineyard in hours, with no possibility of recovery (unlike hail or disease, which may damage only a portion of the crop or allow compensatory regrowth). The two frost types — radiation frost and advective frost — require different risk assessment approaches and different protection strategies: radiation frost (occurring on clear, calm nights when the ground surface radiates heat to the sky, cooling the surface air layer below freezing while upper air remains warmer) is the more common type in viticultural regions, is strongly influenced by topography (cold air drainage creating warm thermal belts on mid-slopes and cold frost pockets in valley floors), and can be combated by disrupting the temperature inversion (wind machines, helicopters) or by adding thermal energy to the vine zone (bougies/smudge pots, aspersion/sprinkler systems, electrical heating) advective frost (occurring when a large cold air mass moves through the region, producing subfreezing temperatures regardless of topographic position) is less frequent but more damaging because it affects entire regions uniformly and cannot be effectively mitigated by site-specific protection measures. The frost protection economic calculus is significant — aspersion systems cost €5,000-15,000/ha to install and €500-2,000/ha to operate per frost event, bougies cost €2,000-5,000/ha per night, and wind machines cost €30,000-50,000 per unit covering 3-5 ha — investments justified only by the high value of the crop in premium appellations.
Frostgefährdung
Frostrisiko-Daten
Kälteschaden-Risikoabschätzung
cold damage risk assessment
freeze event characterization
frost frequency record
Gladstones J (1992) Viticulture and Environment, Winetitles Poling EB (2008) Spring cold injury to winegrapes and protection strategies and methods, HortScience 43: 1652-1662 Molitor D et al. (2014) Late frost damage risk for viticulture under climate change, International Journal of Climatology 34: 1127-1144 Leolini L et al. (2018) Late spring frost impacts on future grapevine distribution in Europe, Field Crops Research 222: 197-208
Frost Risk Data
Fructose content quality determines residual sweetness perception in off-dry Vouvray from the Loire Valley, where controlled fermentation arrest leaves 15-30 g/L residual sugar typically comprising 60-70% fructose this fructose dominance (resulting from preferential glucose fermentation) amplifies perceived sweetness beyond what equi-concentrated glucose would provide, enabling the characteristic demi-sec style with balanced sweetness at relatively moderate residual sugar levels, while the accompanying malic and tartaric acids provide the freshness that prevents cloying character.
Fructose_Content_Quality represents the chemical quality characterizing the concentration of fructose (levulose, D-fructose, fruit sugar) in grape berry tissue. Fructose is a six-carbon ketose monosaccharide that constitutes approximately half of the total reducing sugar in ripe grape berries, typically matching or slightly exceeding glucose concentration at harvest maturity with values of 75-130 g/L for wine grapes. Fructose is the sweetest naturally occurring sugar, approximately 1.2-1.8 times sweeter than sucrose and 1.4-1.7 times sweeter than glucose on a weight basis, making it the primary contributor to perceived berry sweetness despite equimolar concentration with glucose. Like glucose, fructose accumulates through invertase-mediated cleavage of transported sucrose and is stored in mesocarp cell vacuoles. Fructose is fermented by Saccharomyces cerevisiae, but at a slower rate than glucose due to glucose-mediated repression of fructose transport systems. This differential fermentation rate results in preferential glucose consumption during the early and middle stages of fermentation, with fructose becoming the dominant residual sugar in stuck or arrested fermentations. Wines with high residual fructose taste sweeter than wines with equivalent residual glucose due to fructose's higher sweetness intensity. The fructose-to-glucose ratio increases during over-ripening and berry dehydration because fructose is more resistant to respiratory consumption than glucose. Fructose measurement employs specific enzymatic assays using hexokinase, phosphoglucose isomerase, and glucose-6-phosphate dehydrogenase, or chromatographic separation and quantification. The ratio of fructose to glucose provides a maturity index and can indicate metabolic abnormalities when outside normal ranges.
Fructose_Content_Quality specifically addresses the fructose monosaccharide, parallel to Glucose_Content_Quality within Reducing_Sugar_Quality. The higher sweetness intensity of fructose relative to glucose has practical implications for residual sugar perception in finished wines. Fructose undergoes Maillard browning reactions more readily than glucose, contributing to color development during heating and aging. The anomeric forms of fructose (α and β pyranose, α and β furanose) have different sweetness intensities, with the β-D-fructopyranose form dominant at lower temperatures being the sweetest, which is why chilled wines taste sweeter than warm wines at identical sugar concentrations.
D-fructose level
Fruchtzuckergehalt
Fructoseanteil
Fructosegehalt
Levulosekonzentration
fruit sugar concentration
ketohexose quality
levulose content
Kliewer WM (1967) The Glucose-Fructose Ratio of Vitis vinifera Grapes. American Journal of Enology and Viticulture 18(1): 33-41 Margalit Y (2012) Concepts in Wine Chemistry, 3rd Ed, Board and Bench Publishing pp.: 23-47
Fructose Content Quality
Grenache in Rhône Valley shows fruit drop during July-August Year N+1 at BBCH 71-81, with early fruit drop 12-28% from fertilization failure weeks 1-3, occasional shatter 5-15% from summer storms, millerandage formation 8-18% creating shot berries, cumulative berry loss 25-45% in poor years, under variable Mediterranean conditions, typical of this coulure and drop-prone cultivar requiring favorable conditions for acceptable berry retention
Fruit drop represents berry loss phenomena during post-fertilization development when berries abort and abscise from clusters through various mechanisms including early developmental failure, weather-induced shattering, millerandage shot berry formation, or late abscission, reducing berry number and yield from initial fruit set, occurring variably during fruit development depending on specific drop mechanism. This yield-reducing phenomenon occurs during summer development from early fruit set through ripening in late June-September in temperate regions Year N+1, with timing and severity varying by drop type, corresponding to BBCH 71-85 stages from young fruit through ripening. Fruit drop encompasses multiple distinct mechanisms including early fruit drop from failed berry development shortly after fertilization, shatter from weather-induced mechanical berry detachment during vulnerable periods, millerandage formation creating small arrested berries with variable persistence, post-set berry abscission from late developmental failure or stress responses, and overall berry loss reducing cluster berry number below initial fruit set. Drop incidence and timing show variable patterns with early drop typically 1-4 weeks post-flowering from developmental failure, shatter during flowering or early fruit set from rain or wind damage, millerandage formation during fruit set creating persistent shot berries, late drop occasionally during veraison or ripening from stress, and cumulative losses potentially reducing berry number 5-50% depending on severity. Early fruit drop characteristics show young berry abortion and abscission within first 2-4 weeks post-fertilization, failure from inadequate fertilization, seed abortion, or developmental dysfunction, abscission zone formation at pedicel enabling berry detachment, and early losses contributing to coulure poor fruit set if severe. Shatter characteristics show mechanical berry detachment from pedicel through weather damage, rain impact during flowering or early fruit set causing physical separation, wind damage breaking fragile pedicel connections, fallen berries lost from cluster reducing berry number, and shatter losses potentially severe 20-40% in extreme weather events. Millerandage characteristics show formation of small seedless or poorly developed shot berries during fruit set, arrested development creating persistently small 3-6 mm berries, variable persistence with some shot berries remaining through harvest and others abscising, and millerandage berries reducing cluster uniformity and potentially wine quality. Post-set abscission shows occasional late berry drop during veraison or ripening, abscission from developmental stress, water deficit, or disease, late losses typically limited <5% except under severe stress, and late drop distinguished from early developmental failures. Berry loss mechanisms show abscission requiring abscission zone formation and activation, abscission zone at pedicel-berry junction containing specialized separation cells, enzymatic cell wall degradation creating separation layer, vascular tissue severing enabling berry detachment, and mechanical weakness allowing berry fall. Abscission regulation shows hormone control with auxin from seeds preventing abscission, ethylene promoting abscission zone activation, auxin-ethylene balance determining abscission versus retention, and hormonal rescue failure enabling abscission in berries with inadequate seed development. Environmental effects on fruit drop include rain during flowering causing shatter and poor set increasing drop, temperature extremes affecting fertilization and development increasing early drop risk, water stress potentially triggering late drop, wind damage causing mechanical shatter, and overall adverse conditions increasing drop severity across mechanisms. Cultivar variation shows genetic differences in drop susceptibility, coulure-prone cultivars showing high early drop 30-50%, shatter-resistant cultivars maintaining berry retention despite weather, and cultivar characteristics affecting drop patterns. Drop assessment methods include berry number counts comparing flowering to harvest, cluster examination documenting berry scars from abscised berries, pedicel remnants indicating drop timing, and quantification of drop percentage relative to initial set. Drop impacts on yield show direct berry number reduction decreasing cluster and vine yield, severe drop 30-50% substantially affecting production economics, drop variability creating year-to-year yield fluctuations, and drop as major yield loss factor in susceptible cultivars or unfavorable years. Drop effects on cluster architecture show berry spacing increasing with drop, cluster looseness from missing berries, and architecture changes affecting subsequent disease risk and harvest characteristics. Drop prevention strategies include cultivar selection avoiding drop-prone varieties, weather protection during vulnerable periods reducing shatter risk, adequate boron nutrition supporting fertilization reducing early drop, water management preventing late drop from deficit stress, and overall cultural practices minimizing drop causes. The process encompasses four distinguishable drop mechanisms including early fruit drop from developmental failure, shatter from weather damage, millerandage formation creating arrested berries, and post-set berry abscission from late failures. Duration and timing vary by mechanism from early post-flowering through ripening. Regional timing shows fruit drop during late June-September across climates depending on drop type. Viticultural significance includes representing major yield loss factor potentially reducing production 5-50%, demonstrating cultivar and environmental susceptibility patterns, requiring management strategies to minimize drop, and affecting cluster characteristics and wine quality through berry loss. Research applications include analyzing drop mechanisms and abscission regulation, investigating environmental and genetic factors affecting drop susceptibility, evaluating prevention strategies and interventions, quantifying drop impacts on yield and quality, and assessing drop as production limitation.
Fruit_Drop represents berry loss phenomena within Fruit_Set reducing berry number and yield. Subdivided into Early_Fruit_Drop from developmental failure, Shatter from weather damage, Millerandage_Formation creating shot berries, and Post_Set_Berry_Abscission from late failures. Timing and severity vary by mechanism. Cumulative losses 5-50% depending on cultivar and conditions. Major yield-limiting factor. Environmental and genetic susceptibility. Management strategies can reduce drop. Affects cluster architecture and quality.
Beerenabszission
Beerenfall
Fruchtfall
Fruchtverlust
berry abscission
berry drop
cluster berry loss
fruit loss
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 410-415, 450-455 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Fruit Drop
Fruit integrity at harvest quality assessment in Carneros compared damage incidence and juice release across Pinot Noir and Chardonnay under standardized hand harvest protocols with matched handling intensity. Pinot Noir demonstrated poor fruit integrity at harvest quality consistent with its thin-skinned reputation, showing 4.2% visible berry damage, 2.8% berry detachment from pedicels during handling, and juice release of 85 mL per 20 kg harvest bin attributable to harvest damage, requiring enhanced care protocols including smaller picking containers and rapid transport to minimize quality impact. Chardonnay exhibited superior fruit integrity at harvest quality with thicker, more resilient skin, showing only 1.1% visible damage, 0.8% berry detachment, and juice release of only 22 mL per equivalent bin weight, permitting standard handling protocols without quality compromise. The fruit integrity quality differential translated directly to must quality, with Pinot Noir juice showing 35% higher polyphenol oxidase activity indicating advanced oxidation from harvest damage compared to Chardonnay, establishing cultivar-specific harvest handling requirements based on fruit integrity characteristics.
Fruit integrity at harvest quality is a harvest logistics quality that inheres in a grapevine cultivar or clone and represents the capacity of grape clusters and individual berries to withstand the physical stresses of the harvest operation including handling, placement in collection containers, and initial transport within the vineyard without suffering structural damage, juice release, berry detachment, or other forms of deterioration that would compromise quality of the harvested material delivered to the winery. This quality captures the resilience of fruit to the inevitable physical impacts and compression forces encountered during harvest, recognizing that even careful handling subjects fruit to stresses that may exceed damage thresholds for fragile cultivars while remaining well within tolerance for robust types. The ontological significance of fruit integrity at harvest quality lies in its role as a determinant of achievable quality from the vineyard gate forward, wherein fruit that arrives at the winery damaged, leaking juice, or partially crushed has already begun oxidation and microbial processes that limit the quality potential of resulting wines regardless of subsequent processing excellence, making harvest integrity a critical quality control point that varies substantially among cultivars. The biological basis of fruit integrity at harvest quality encompasses the structural characteristics determining damage resistance including berry skin thickness and tensile strength, the flexibility versus brittleness of skin tissue under impact, the firmness of flesh tissue providing support against compression, the strength of berry attachment to pedicels resisting detachment under handling stress, and the cluster architecture determining whether individual berries are protected within compact structures or exposed in loose arrangements more vulnerable to contact damage. Cultivar differences in fruit integrity at harvest span a wide range, from robust thick-skinned varieties tolerating substantial handling without visible damage to thin-skinned types where even gentle handling causes skin cracking, juice release, and accelerated oxidation onset. The maturity stage at harvest influences fruit integrity through the softening of flesh tissue and weakening of skin structure that accompanies ripening, creating vintage and timing interactions wherein the same cultivar may exhibit different integrity characteristics depending on ripeness level and pre-harvest conditions affecting berry turgor and structural integrity.
Fruit_Integrity_At_Harvest_Quality is_a Harvest_Logistics_Quality, representing the damage resistance of fruit during harvest operations. This class addresses structural resilience to handling stresses distinct from transport tolerance which extends to post-harvest logistics. Berry skin thickness, firmness, and pedicel attachment strength are key determinants. Quantitative assessment through damage incidence percentage and juice release volume provides objective metrics. Distinguished from Mechanical_Harvesting_Suitability_Quality by addressing hand harvest handling as well as machine harvest impacts. Critical for quality-sensitive production where harvest damage initiates quality-limiting oxidation and microbial processes.
Beerenstrukturstabilität
Ernte-Schadensresistenz
Fruchtintegrität bei der Ernte
Traubenrobustheit
berry resilience
cluster robustness
fruit structural integrity
harvest damage resistance
Rolle L et al. (2012) Influence of grape density and harvest date on changes in phenolic composition, phenol extractability indices, and instrumental texture properties during ripening. Journal of Agricultural and Food Chemistry 60:7571-7581 Letaief H et al. (2008) Assessment of grape skin hardness by a puncture test. Journal of the Science of Food and Agriculture 88:1567-1575 Coombe BG, Iland P (2004) Grape berry development and winegrape quality. In: Dry PR, Coombe BG (eds) Viticulture Volume 1 Resources, 2nd Edition, Winetitles, Adelaide, pp. 210-248
Fruit Integrity At Harvest Quality
Fruit maintenance respiration assessment tracking berry development in Shiraz reveals the respiratory losses affecting final composition, with post-véraison berries respiring 8.5 mg CO₂/kg/hour at 25°C (combined maintenance and ripening metabolism), translating to daily sugar loss of approximately 0.15 g/berry that competes with daily sugar import of approximately 0.45 g/berry, resulting in net sugar accumulation of 0.30 g/berry/day, with elevated temperatures (35°C) increasing respiration to 15.2 mg CO₂/kg/hour and reducing net accumulation to 0.18 g/berry/day, demonstrating how fruit respiration determines the efficiency of sugar accumulation and the heat stress sensitivity of final Brix.
Fruit maintenance respiration quality is a maintenance respiration quality that represents the rate of respiratory metabolism required to maintain grape berry cellular structures and ripening processes without net growth, characterizing the carbon cost of sustaining membrane integrity, enzyme systems, and metabolic activity in developing and ripening fruit that represents an increasing respiratory sink as clusters develop toward harvest. Within the BFO framework, this quality specifically addresses the fruit component of maintenance respiration, distinguishing berry maintenance costs from growth respiration while recognizing that the boundary between maintenance and active ripening metabolism is less distinct in fruit than in vegetative tissues, as the ripening process involves continuous metabolic transformation rather than simple homeostatic maintenance. The ontological significance lies in fruit maintenance respiration representing the respiratory cost of sustaining berry viability and ripening capacity, where maintenance respiration continues throughout berry development and ripening, consuming sugars that would otherwise contribute to final Brix, and where the balance between maintenance cost and sugar accumulation affects both yield (berry mass) and quality (final sugar concentration). The physiological basis for fruit maintenance respiration includes membrane integrity (maintaining compartmentalization essential for ripening), enzyme turnover (sustaining the enzymatic machinery for sugar accumulation, acid degradation, and secondary metabolite synthesis), protein synthesis (continued expression of ripening-related genes), and cellular homeostasis (ion balance, pH regulation, and oxidative protection). The developmental trajectory of fruit respiration shows high rates during cell division (combined maintenance and growth), declining rates during cell expansion (lower growth component), and continued maintenance respiration through ripening with gradual decline as metabolism slows toward harvest, with total fruit respiratory losses representing 15-25% of sugars translocated to clusters. The temperature sensitivity of fruit respiration is particularly significant for wine grape quality, as elevated temperatures increase respiratory losses of sugar while simultaneously reducing acidity through enhanced malic acid catabolism, making fruit respiration a key determinant of the heat stress impact on fruit composition. Understanding fruit maintenance respiration quality enables prediction of sugar losses to respiration, heat stress impact assessment, and appreciation of the metabolic costs affecting final fruit composition and wine quality potential.
Fruit_Maintenance_Respiration_Quality addresses the specific maintenance costs of developing and ripening grape berries. This quality determines respiratory sugar losses that affect final fruit composition and quality. The class supports ripening prediction and heat stress assessment through characterization of the fruit respiratory component affecting sugar accumulation efficiency.
Beeren-Grundumsatz
Frucht-Erhaltungsatmung
Fruchtmetabolismuskosten
Trauben-Instandhaltungsatmung
berry basal respiration
cluster respiration
fruit metabolic cost
grape maintenance respiration
Ollat N, Gaudillère JP (1998) The effect of limiting leaf area during stage I of berry growth on development and composition of berries of Vitis vinifera L. cv. Cabernet Sauvignon. American Journal of Enology and Viticulture 49(3): 251-258 Sweetman C et al. (2009) Regulation of malate metabolism in grape berry and other developing fruits. Phytochemistry 70(11-12): 1329-1344
Fruit Maintenance Respiration Quality
Merlot in Bordeaux shows fruit set during early July Year N+1 at BBCH 69-71, with 54-64% flowers successfully producing berries containing 1-3 seeds, ovary expansion from 6 mm to 14-18 mm over 2-3 weeks post-fertilization, auxin from developing seeds preventing abscission and triggering growth, unfertilized flowers aborting, under oceanic conditions following favorable flowering period, typical of this mid-season cultivar with moderate-to-good set reliability determining season yield
Fruit set represents the critical reproductive outcome phase when successful fertilization leads to berry development with fertilized ovules producing auxin preventing flower abscission and triggering ovary expansion, while unfertilized flowers lacking seed development abort and abscise, establishing final berry number determining yield and marking definitive transition from flowering reproductive phase to berry developmental phase. This essential outcome phase occurs during late flowering and early berry development in late June-early July in temperate regions Year N+1, typically proceeding over 1-3 weeks following fertilization as fertilization outcomes manifest, corresponding to BBCH 69-71 stages transitioning from flowering completion to young fruit development. Fruit set mechanism involves fertilization success determining ovule and seed development, auxin and other hormone production from developing seeds preventing programmed flower abscission, hormone signaling triggering ovary wall cell division and expansion initiating berry formation, unfertilized flowers lacking hormonal rescue undergoing abscission, and final berry number establishment representing successful fruit set percentage. Fertilization outcomes show variable success with 30-75% flowers achieving successful fertilization and fruit set depending on cultivar and environmental conditions, successful fertilization producing 1-3 seeds per berry from fertilized ovules in 2-4 ovule flowers, and fertilization failure causing flower abortion eliminating potential berries from yield. Successful ovule fertilization shows double fertilization completion producing viable zygote and endosperm, seed development initiation with embryo and endosperm growth, integument development forming seed coat, and overall seed maturation proceeding coordinated with berry development. Failed ovule fertilization shows fertilization failure from inadequate pollination, pollen tube growth failure, or fusion problems, ovule abortion lacking seed development, and flower abortion unless other ovules in same flower fertilize successfully providing hormonal rescue. Partial ovule fertilization shows some ovules fertilizing successfully while others fail within multi-ovule flower, variable seed number 1-3 per berry depending on partial success, and berry development proceeding with reduced seed number potentially affecting berry size. Auxin production from developing seeds shows critical hormone synthesis in embryo and endosperm beginning within days of fertilization, auxin export from seed to surrounding ovary tissues, and hormone accumulation preventing abscission and triggering growth responses. Auxin prevention of abscission shows hormone blocking formation of abscission zone at flower-pedicel junction, maintaining vascular connection supporting developing berry, and rescuing flower from programmed senescence and abscission pathway. Auxin induction of berry growth shows hormone triggering cell division in ovary wall creating berry flesh, promoting cell expansion increasing berry size, and coordinating berry development with seed maturation. Post-fertilization signaling shows complex hormone interactions including auxin as primary signal, gibberellins possibly contributing to growth promotion, cytokinins potentially involved in cell division stimulation, and integrated signaling establishing berry developmental program. Ovary expansion initiation shows rapid growth beginning 1-2 weeks post-fertilization, ovary diameter increasing from 4-8 mm pre-fertilization to 10-20 mm within 2-3 weeks, cell division intensity maximizing with 10-15 hour cycle times during early expansion, and visible berry formation confirming successful fruit set. Cell division characteristics during early berry show peak mitotic activity in ovary wall pericarp tissue, rapid cell proliferation producing berry flesh cells, division rate declining after initial 3-4 weeks as cell expansion phase begins, and final cell number established during early development determining potential berry size. Early berry formation shows ovary wall thickening creating berry flesh, pericarp tissue differentiation into exocarp skin, mesocarp flesh, and endocarp near seeds, vascular development establishing nutrient transport to growing berry, and overall berry structure establishment during first weeks. Berry commitment shows developmental commitment to berry formation becoming irreversible after initial growth, continued development independent of ongoing seed development in some cases though seed presence beneficial, and berry reaching point of assured maturation barring severe environmental stress or disease. Unfertilized flower abscission shows flowers lacking seed development failing to produce adequate auxin rescue, abscission zone formation at pedicel creating separation layer, vascular connection severing and flower dropping, and coulure poor fruit set from excessive abscission reducing yield. Coulure characteristics show environmental or physiological poor fruit set with high proportion flowers aborting, typical occurrence during unfavorable flowering conditions particularly rain or extreme temperatures, variable cultivar susceptibility with some showing consistent good set and others prone to coulure, and severe coulure potentially reducing yield by 50-80% in affected vintages. Millerandage relationship to fruit set shows shot berries resulting from partial fertilization or early developmental disruption, small seedless berries persisting but failing to develop normally, and millerandage creating berry size heterogeneity affecting harvest and quality. Fruit set percentage calculation shows proportion of flowers producing berries, typical range 30-75% depending on conditions with good set 60-75% under favorable flowering weather and poor set 30-50% under adverse conditions, and set percentage directly determining cluster density and yield. Environmental effects on fruit set include flowering period weather critically affecting pollination and fertilization success, rain during flowering causing poor set through pollination disruption and pollen damage, temperature extremes reducing fertilization efficiency and seed development, boron deficiency preventing normal pollen tube growth and fertilization, and overall flowering conditions as primary fruit set determinant. Temperature effects show optimal 22-28°C during flowering supporting high fruit set 60-75%, cool conditions 15-20°C reducing set to 40-60% through delayed development and reduced fertilization, excessive heat above 33°C severely reducing set to 20-40% through reproductive dysfunction, and critical temperature sensitivity during 2-3 day peak flowering window. Boron nutrition critical importance shows adequate boron 10-20 ppm essential for pollen tube integrity and fertilization success, deficiency causing poor fruit set even with adequate flowering through fertilization failure, and boron as frequent limiting factor for set in deficient soils. Cultivar variation in fruit set shows genetic differences in setting ability with some cultivars showing reliable 65-75% set across conditions, others showing variable 40-70% set depending on weather, and problematic cultivars prone to coulure showing poor 30-50% set even under favorable conditions. Cluster architecture effects on set show compact clusters potentially showing reduced set from crowding and competition, loose clusters generally showing better set from improved flower exposure and pollination, and cluster density reflecting both genetic architecture and fruit set percentage. Resource allocation following fruit set shows dramatic shift to developing berries consuming 40-60% total photosynthate, vegetative growth continuing requiring 30-40% resources, and remaining allocation to root growth and reserves, establishing berry development as dominant sink. Yield determination through fruit set shows final berry number per cluster established during set period, cluster number per vine determined by previous season bud formation, and total yield calculated from berry number, berry weight, and cluster number. Fruit set assessment timing shows preliminary evaluation 2-3 weeks post-flowering observing berry versus abscised flower proportion, definitive assessment 4-6 weeks post-flowering confirming established berry number, and early yield prediction based on set percentage and cluster counts. The process encompasses two major components including ovule fertilization showing variable success creating seed number diversity, and berry initiation showing hormone-mediated transition to berry development. Duration typically spans 2-4 weeks from fertilization through definitive berry establishment. Cultivar variation includes fruit set reliability, coulure susceptibility, and seed number characteristics. Regional timing shows fruit set during late June-early July in warm climates, early-mid July in temperate regions, and mid-late July in cool areas, occurring 1-3 weeks post-fertilization. Viticultural significance includes representing yield determination phase establishing final berry number, requiring favorable fertilization outcomes and environmental conditions, demonstrating critical importance of flowering period weather for set success, and marking definitive transition from reproductive flowering to developmental berry growth phase. Research applications include analyzing fertilization outcomes and fruit set determinants, investigating hormone signaling coordinating seed and berry development, evaluating environmental and nutritional effects on set success, quantifying fruit set percentage and yield relationships, and developing strategies for improving set reliability and reducing coulure.
Fruit_Set represents critical reproductive outcome phase determining berry number and yield. Corresponds to BBCH 69-71 stages 1-3 weeks post-fertilization. Subdivided into Ovule_Fertilization showing variable success outcomes and Berry_Initiation showing hormone-mediated berry development. Success rate 30-75% depending on conditions. Auxin from seeds critical for preventing abscission and triggering growth. Coulure represents failed fruit set. Marks definitive transition from Flowering to Berry_Development. Yield-determining phase.
Beerenbildung
Fruchtansatz
Fruchtbildung
berry establishment
berry set
fruit formation
reproductive success
reproduktiver Erfolg
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 405-415 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 308-315 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Fruit Set
Fruit set percentage in grapevine typically ranges from 15-50% of total flowers, with Chardonnay achieving 35-45% under favorable conditions and Merlot averaging 20-30%. A cluster with 300 flowers at bloom producing 90 berries represents 30% fruit set. Optimal fruit set requires temperatures of 20-25°C during flowering, adequate carbohydrate supply, and successful pollination of at least 2 ovules per flower. Fruit set below 20% often results in excessively loose clusters, while set above 50% may cause compact clusters prone to disease.
A reproductive disposition that is the inherited capacity of a Vitis vinifera plant to successfully convert pollinated flowers into developing berries through fertilization, initial cell division, and retention on the rachis, determining the proportion of flowers that become harvestable fruit.
Fruit_Set_Disposition serves as the parent class for traits affecting the flower-to-berry transition. The process involves multiple sequential steps including pollen germination, tube growth, fertilization, and hormone-triggered berry development. Unfertilized flowers abscise within 7-14 days post-bloom. Both genetic factors (cultivar) and environmental conditions (temperature, nutrition, water status) strongly influence fruit set success.
Beerenansatz
Befruchtungserfolg
berry set capacity
fruit retention trait
set percentage disposition
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 115-135 Vasconcelos MC et al. (2009) The flowering process of Vitis vinifera: a review. American Journal of Enology and Viticulture 60:411-434
Fruit Set Disposition
Fruit set percentage assessment comparing flowering conditions in Merlot reveals the weather effect on flower-to-berry conversion, with favorable conditions (25°C, dry, calm during flowering) achieving 48% fruit set (144 berries from 300 flowers per cluster, compact clusters), marginal conditions (18°C, intermittent rain) achieving 32% set (96 berries, moderately loose clusters), and poor conditions (14°C, persistent rain during bloom) achieving only 18% set (54 berries, very loose clusters), with the fruit set variation creating 2.7-fold difference in berry number per cluster from identical flower populations, demonstrating how fruit set percentage determines yield realization from flowering potential.
Fruit set percentage quality is a yield potential quality that represents the proportion of flowers that successfully develop into berries following pollination and fertilization, characterizing the conversion efficiency from flowers to fruit that determines how much of the flowering potential is realized as berry production and thus establishing the berry number component of final yield. Within the BFO framework, this quality specifically addresses the flower-to-berry conversion dimension of yield potential, distinguishing fruit set success from prior flower number or subsequent berry development while recognizing that fruit set is the critical determination point where pollination success, fertilization completion, and post-fertilization retention together establish whether each flower becomes a berry or abscises as a failed reproductive attempt. The ontological significance lies in fruit set percentage determining the berry population that will develop and ripen, with typical wine grape fruit set of 25-50% meaning that the majority of flowers fail to produce berries under normal conditions, and with variation in this percentage substantially affecting final yield—a shift from 30% to 45% fruit set representing a 50% increase in berry number and potentially similar yield increase from the same flower population. The factors affecting fruit set include weather during flowering (warm, dry conditions favoring set cool, wet, or windy conditions reducing it), carbohydrate availability (adequate current photosynthate and reserves supporting berry retention), hormone balance (appropriate auxin, gibberellin, and cytokinin levels promoting fruit development), and cultivar genetics (some varieties inherently setting more reliably than others). The assessment of fruit set involves counting flowers per inflorescence before bloom and berries per cluster after set, with the ratio (berries/flowers × 100) expressing set percentage, though practical assessment often uses cluster compactness or berry count as indicators given the difficulty of precise flower counting. Understanding fruit set percentage quality enables diagnosis of poor set causes, yield forecasting based on flower counts and expected set percentages, and management targeting improved set conditions during the critical flowering period.
Fruit_Set_Percentage_Quality addresses the conversion efficiency from flowers to developing berries. This quality is determined during flowering by weather, nutrition, and cultivar factors. The class supports yield realization assessment through fruit set percentage characterization.
Ansatzeffizienz
Beerenansatzrate
Blüte-Frucht-Konversion
Fruchtansatzprozentsatz
berry set percentage
flower-to-fruit conversion
fruit set rate
set efficiency
Fruit Set Percentage Quality
Fruit zone humidity quality manipulation in Loire Valley Chenin Blanc evaluated disease management implications across canopy density treatments during a Botrytis-favorable autumn with frequent precipitation events. Dense unthinned canopies exhibited poor fruit zone humidity quality with relative humidity exceeding 90% for mean 16.2 hours following 5mm precipitation events, surpassing the Botrytis germination threshold and resulting in 38% bunch rot incidence at harvest. Shoot-thinned canopies achieving 25% gap frequency showed improved fruit zone humidity quality with >90% RH duration reduced to 9.4 hours, corresponding to 22% rot incidence. Aggressively managed canopies with 40% gaps and basal leaf removal achieved optimal fruit zone humidity quality with >90% RH duration of only 5.8 hours, below the germination threshold, resulting in negligible 4% rot incidence without additional fungicide applications beyond the standard program, demonstrating that canopy-based fruit zone humidity quality optimization can substitute for chemical intervention in disease management.
Fruit zone humidity quality is a canopy microclimate quality that inheres in the fruit-bearing region of a grapevine canopy and represents the water vapor concentration in the air immediately surrounding grape clusters as characterized by relative humidity, vapor pressure deficit, and the temporal patterns of humidity including duration of near-saturation conditions following precipitation, dew formation, or irrigation events that determine the wetness periods enabling fungal pathogen germination and infection. This quality fundamentally influences disease development potential by controlling the moisture conditions required for spore germination and hyphal penetration while also affecting berry transpiration rates, cuticle development, and potentially berry splitting susceptibility under high humidity conditions preceding precipitation. The ontological significance of fruit zone humidity quality lies in its role as the primary environmental determinant of fungal disease pressure independent of fungicide applications, with the duration of high humidity periods and surface wetness serving as key epidemiological parameters for diseases including Botrytis bunch rot, downy mildew, and powdery mildew that collectively represent major threats to grape production worldwide. The physical basis of fruit zone humidity modification within grapevine canopies involves transpirational water vapor release from leaves that elevates humidity in the canopy interior, restricted air exchange with drier ambient atmosphere due to reduced wind penetration and turbulent mixing in dense canopies, and dew formation and persistence patterns that depend on radiative cooling rates influenced by canopy openness. Dense canopies with multiple leaf layers create humid microclimates where relative humidity may remain above 80% for extended periods even when ambient humidity is moderate, while open canopies with good ventilation permit rapid equilibration with ambient conditions and faster drying of wetted surfaces. The relationship between fruit zone humidity quality and disease incidence follows established epidemiological principles wherein Botrytis cinerea requires approximately 15 hours of relative humidity above 90% for spore germination followed by additional wetness for infection establishment, with canopy management directly influencing whether these thresholds are reached following precipitation events. Management approaches to optimize fruit zone humidity quality include shoot thinning and leaf removal to improve air circulation, row orientation considering prevailing wind direction for cross-row ventilation, and training system selection affecting the vertical distribution of foliage relative to fruit positioning.
Fruit_Zone_Humidity_Quality is_a Canopy_Microclimate_Quality, representing the moisture dimension of fruit zone environmental conditions. This class addresses humidity at cluster positions with particular emphasis on disease-relevant wetness duration. Key thresholds include ~15 hours >90% RH for Botrytis germination, ~6 hours leaf wetness for downy mildew infection. Distinguished from Air_Circulation_Quality which addresses ventilation as a structural characteristic, while this class addresses the humidity outcomes of ventilation patterns. Critical for integrated pest management and sustainable viticulture approaches.
Beerenzonen-Luftfeuchtigkeit
Fruchtzonenfeuchte
Traubenbereichsfeuchte
Traubenzonenluftfeuchtigkeit
bunch zone relative humidity
cluster zone humidity
fruit zone moisture
fruit zone vapor pressure
English JT et al. (1989) Influence of grapevine canopy management on evaporative potential in the fruit zone. American Journal of Enology and Viticulture 40:123-129 Percival DC et al. (1994) Canopy microclimate and Botrytis bunch rot development in Riesling grapevines. Vitis 33:17-22 Valdés-Gómez H et al. (2008) Modelling of grapevine bunch rot susceptibility to Botrytis cinerea. Australian Journal of Grape and Wine Research 14:142-151
Fruit Zone Humidity Quality
Comparison of fruit zone leaf layers quality across basal leaf removal timings in Napa Valley Cabernet Sauvignon evaluated effects on fruit composition and sunburn incidence. Pre-bloom leaf removal achieving fruit zone LLN of 0.6 by véraison produced highest anthocyanin concentration at 2.1 mg/g but suffered 12% berry sunburn on western exposures. Berry set timing leaf removal achieving fruit zone LLN of 0.9 produced 1.8 mg/g anthocyanins with only 4% sunburn. Véraison timing leaf removal achieving fruit zone LLN of 1.1 showed 1.5 mg/g anthocyanins with minimal 1% sunburn. No leaf removal controls with fruit zone leaf layers quality of 2.3 produced lowest anthocyanins at 1.1 mg/g with no sunburn but elevated Botrytis incidence of 8% versus less than 2% in all leaf-removed treatments.
Fruit zone leaf layers quality is a leaf layer number quality that inheres in a grapevine canopy system and represents the mean number of leaf contacts encountered when probing horizontally through the canopy at the height corresponding to cluster position, providing specific characterization of the foliage density immediately surrounding developing fruit that directly determines cluster microclimate conditions for ripening, disease development potential, and spray coverage effectiveness. This quality receives particular management attention because the fruit zone represents the target tissue for quality development, and density conditions at cluster height may differ substantially from overall canopy density depending on training system architecture and shoot positioning effectiveness. The ontological significance of fruit zone leaf layers quality lies in its direct relationship to the environmental conditions experienced by ripening clusters, wherein excessive leaf layering at fruit height creates the shading, humidity retention, and spray interception problems that compromise fruit quality and increase disease incidence regardless of conditions elsewhere in the canopy. The specific microclimate consequences of fruit zone leaf layer number include attenuation of photosynthetically active radiation reaching berry surfaces that influences phenolic biosynthesis and chlorophyll degradation, restriction of convective air exchange that extends wetness duration after precipitation, and physical interception of spray droplets before they reach cluster surfaces. Optimal fruit zone leaf layers quality for wine grape production typically targets values between 0.8 and 1.2, somewhat lower than whole-canopy targets to ensure adequate fruit exposure given the critical importance of cluster microclimate for quality outcomes. Management interventions specifically targeting fruit zone leaf layers quality include basal leaf removal eliminating leaves in the cluster zone while retaining upper canopy foliage for photosynthesis, shoot positioning that separates fruiting zone from the main foliar wall, and lateral shoot removal in the fruit zone that can contribute substantially to local density without appearing in upper canopy assessments. The timing of fruit zone leaf layers quality optimization involves balancing early intervention benefits against the protective value of initial cluster shading during early development when berries are most susceptible to sunburn damage, with typical recommendations suggesting leaf removal initiation at berry set through véraison depending on climate and cultivar sun sensitivity.
Fruit_Zone_Leaf_Layers_Quality is_a Leaf_Layer_Number_Quality, representing the specific cluster-height assessment of leaf layering. This class addresses the critical fruit zone region where density most directly impacts quality outcomes. Distinguished from External_Leaf_Layer_Quality and Interior_Leaf_Layer_Quality which address outer canopy and deep interior positions respectively. The lower optimal target of 0.8-1.2 compared to overall canopy LLN reflects the importance of fruit exposure. Basal leaf removal serves as the primary management tool specifically addressing this quality.
Fruchtzonenblattdichte
Traubenbereich-Belaubung
Traubenzone-Laubschichtzahl
bunch zone LLN
cluster zone density
fruit zone foliage density
fruiting zone leaf layers
Intrieri C et al. (2008) Early defoliation (hand vs mechanical) for improved crop control and grape composition in Sangiovese (Vitis vinifera L.). Australian Journal of Grape and Wine Research 14:25-32 Tardaguila J et al. (2010) Impact of early leaf removal on yield and fruit and wine composition of Vitis vinifera L. Graciano and Carignan. American Journal of Enology and Viticulture 61:372-381 Poni S et al. (2006) Effects of early defoliation on shoot photosynthesis, yield components, and grape composition. American Journal of Enology and Viticulture 57:397-407
Fruit Zone Leaf Layers Quality
Fruit zone light intensity quality optimization in Waipara Pinot Noir established dose-response relationships between cluster irradiance and fruit composition parameters. Shaded clusters receiving fruit zone light intensity quality of only 4% ambient PAR (approximately 80 μmol m⁻² s⁻¹ at solar noon) produced fruit with anthocyanin concentration of 0.6 mg/g, total phenolics of 1.8 mg/g, and methoxypyrazine concentration of 22 ng/L. Moderate exposure achieving 15% ambient PAR (300 μmol m⁻² s⁻¹) improved fruit zone light intensity quality to optimal range, with anthocyanins increasing to 1.4 mg/g, total phenolics to 3.2 mg/g, and methoxypyrazines declining to 8 ng/L. High exposure at 35% ambient PAR (700 μmol m⁻² s⁻¹) further increased phenolics to 3.8 mg/g with methoxypyrazines at 4 ng/L, but anthocyanins declined to 1.1 mg/g attributed to temperature-induced degradation with berry temperatures exceeding 38°C, establishing 15-20% ambient PAR as the optimal fruit zone light intensity quality target for this cool climate site.
Fruit zone light intensity quality is a canopy microclimate quality that inheres in the fruit-bearing region of a grapevine canopy and represents the quantity of photosynthetically active radiation reaching grape cluster surfaces as characterized by instantaneous irradiance values, daily light integral accumulation, and the proportion of incident above-canopy radiation transmitted to fruit positions through canopy gaps and attenuated through overlying leaf layers. This quality directly drives the photomorphogenic and photochemical processes in berry tissues that determine phenolic composition, color development, and flavor precursor synthesis, with light exposure serving as the primary environmental signal activating flavonoid biosynthetic pathways while simultaneously influencing berry temperature through radiative heating. The ontological significance of fruit zone light intensity quality lies in its role as the fundamental energy input controlling quality-determining biochemical pathways in grape berries, wherein light-responsive genes encoding enzymes of the phenylpropanoid and flavonoid pathways exhibit expression patterns directly correlated with fruit zone irradiance levels. The physics of light transmission to the fruit zone involves exponential attenuation through successive leaf layers following Beer-Lambert principles, with each leaf layer absorbing approximately 85-90% of incident photosynthetically active radiation such that fruit positions behind two leaf layers typically receive less than 5% of above-canopy irradiance. The relationship between fruit zone light intensity quality and berry composition reflects established light dependencies of phenolic biosynthesis, with anthocyanin concentration, flavonol accumulation, and tannin polymerization all positively correlated with fruit exposure up to saturation thresholds, while methoxypyrazine degradation accelerates under high light conditions reducing herbaceous character in ripening fruit. Optimal fruit zone light intensity quality for wine grape production typically targets 10-20% of above-canopy PAR, corresponding to approximately 150-400 μmol m⁻² s⁻¹ at solar noon under clear sky conditions, providing sufficient irradiance for phenolic synthesis without excessive radiative heating that risks sunburn damage or anthocyanin degradation at temperatures above 35°C. The spectral quality of fruit zone light changes with canopy depth as chlorophyll preferentially absorbs red and blue wavelengths, enriching transmitted light in far-red and green wavelengths that alter the phytochrome signaling environment potentially influencing berry developmental responses independently of total irradiance effects.
Fruit_Zone_Light_Intensity_Quality is_a Canopy_Microclimate_Quality, representing the radiation dimension of fruit zone environmental conditions. This class addresses PAR specifically at cluster positions, distinct from Light_Penetration_Quality which addresses canopy-level transmission characteristics. The optimal target of 10-20% ambient PAR (~150-400 μmol m⁻² s⁻¹) provides quantitative benchmark, though climate-specific adjustments may be required to balance phenolic benefits against temperature risks. Spectral quality changes with canopy depth provide additional dimension beyond total irradiance.
Beerenbelichtungsintensität
Fruchtzonenbelichtung
Traubenstrahlung
Traubenzonenlicht
bunch zone irradiance
cluster light exposure
fruit zone PAR
fruit zone solar radiation
Dokoozlian NK, Kliewer WM (1995) The light environment within grapevine canopies. I. Description and seasonal changes during fruit development. American Journal of Enology and Viticulture 46:209-218 Downey MO et al. (2006) Cultural practice and environmental impacts on the flavonoid composition of grapes and wine: a review of recent research. American Journal of Enology and Viticulture 57:257-268 Matus JT et al. (2009) Post-veraison sunlight exposure induces MYB-mediated transcriptional regulation of anthocyanin and flavonol synthesis in berry skins of Vitis vinifera. Journal of Experimental Botany 60:853-867
Fruit Zone Light Intensity Quality
The fruit zone management strategy for Riesling in the Wachau integrates three interventions targeting the cluster zone between 0.6-1.0 m on the VSP trellis — selective leaf removal of 3-4 basal leaves on the east-facing side at E-L 27-29 (exposing clusters to morning sun while retaining afternoon shade from the western leaf wall), lateral shoot removal within the fruit zone at E-L 31 (eliminating the secondary vegetative growth that recolonizes the opened fruit zone within 2-3 weeks of leaf removal), and cluster wing removal on excessively compact clusters at E-L 33 (removing the upper shoulder of tight Riesling clusters to reduce internal berry contact, lower humidity, and improve spray penetration for pre-véraison Botrytis protection) — the combined effect is a fruit zone receiving 25-35% ambient PAR with an average temperature 1-2°C above ambient during the day and 1-2°C below ambient at night due to radiative cooling in the exposed zone.
A canopy management description that documents the integrated set of interventions applied specifically to the cluster zone of the canopy — the vertical band between the fruiting wire and the first foliage wire — to optimize the microenvironment experienced by developing clusters with respect to light exposure, temperature, humidity, air circulation, and spray penetration, coordinating leaf removal, lateral shoot management, and cluster positioning into a coherent fruit zone management strategy.
Fruit_Zone_Management represents the integrative concept that coordinates individual canopy management operations (leaf removal, lateral management, cluster manipulation) into a coherent strategy focused specifically on the microenvironment of developing fruit. The fruit zone microclimate — defined by light intensity, spectral quality, temperature, humidity, and wind speed within the 20-40 cm band occupied by clusters — directly determines berry composition parameters including anthocyanin and tannin concentration (enhanced by moderate light exposure), methoxypyrazine degradation (accelerated by light and temperature), organic acid metabolism (malic acid respiration accelerated by temperature), and disease incidence (Botrytis favored by >95% relative humidity and poor air circulation): managing these microclimate parameters through coordinated fruit zone interventions is the most direct mechanism available to the viticulturist for influencing wine quality during the growing season. The Smart-Robinson canopy microclimate assessment framework (point quadrat analysis measuring leaf layer number, percent gaps, and percent interior clusters) provides the diagnostic tool for evaluating fruit zone management effectiveness and identifying sites requiring intervention.
Fruchtzone-Bewirtschaftung
Traubenbereichs-Mikroklima-Management
Traubenzone-Management
bunch zone microclimate management
cluster zone management
fruiting zone protocol
Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles pp: 91-120 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 365-378 Reynolds AG (ed) (2022) Managing Wine Quality Volume 1: Viticulture and Wine Quality, 2nd Ed, Woodhead Publishing
Fruit Zone Management
Fruit zone temperature quality monitoring in San Joaquin Valley Cabernet Sauvignon compared thermal regimes across canopy management treatments during a heat wave event with ambient maxima exceeding 40°C for five consecutive days. Heavily shaded clusters within dense canopies exhibited moderated fruit zone temperature quality with maximum berry surface temperatures of 36°C, maintaining temperatures below the anthocyanin synthesis inhibition threshold but with persistently low light limiting color development. Partially exposed clusters from moderate leaf removal reached fruit zone temperatures of 42°C, approaching but not exceeding the sunburn damage threshold while achieving beneficial light exposure. Fully exposed clusters on the western canopy face of aggressively defoliated vines experienced fruit zone temperature quality extremes of 48°C, suffering 23% visible sunburn damage and anthocyanin degradation evident as 35% lower color density compared to optimally managed treatments, demonstrating the critical importance of fruit zone temperature quality management in hot climate viticulture.
Fruit zone temperature quality is a canopy microclimate quality that inheres in the fruit-bearing region of a grapevine canopy and represents the thermal regime experienced by developing grape clusters as characterized by mean temperature, maximum and minimum temperature extremes, diurnal temperature amplitude, and the duration of exposure to temperature ranges relevant for specific metabolic processes including organic acid respiration, anthocyanin synthesis, and aromatic compound development. This quality directly influences the rate and outcome of berry ripening processes, with temperature serving as a primary driver of metabolic reaction rates and a modulator of gene expression patterns controlling quality-determining biosynthetic pathways. The ontological significance of fruit zone temperature quality lies in its role as the proximate thermal signal received by berry tissues, which may differ substantially from ambient air temperature due to radiative heating of exposed berries, convective coupling with surrounding foliage, and the buffering effects of canopy boundary layers that restrict heat exchange with the atmosphere. The physics of fruit zone temperature involves complex energy balance interactions wherein berry surface temperature represents the equilibrium between absorbed solar radiation, longwave radiation exchange with sky and surroundings, convective heat transfer with adjacent air, and evaporative cooling when berry surfaces are wet. Exposed berries in direct sunlight can exceed ambient air temperature by 10-15°C during peak radiation periods, potentially reaching temperatures of 40-50°C that induce heat stress, enzyme denaturation, and sunburn damage, while shaded berries within dense canopies may remain several degrees below ambient due to restricted radiative heating and evaporative cooling from adjacent transpiring leaves. The relationship between fruit zone temperature quality and fruit composition reflects established temperature dependencies of ripening biochemistry, with malic acid respiration accelerating exponentially above 25°C leading to rapid acid decline in hot conditions, anthocyanin biosynthesis showing optimum activity between 20-30°C with inhibition at higher temperatures, and aromatic compound accumulation and retention favoring cool conditions with high diurnal amplitude that preserves volatile compounds against thermal degradation. Management manipulation of fruit zone temperature quality operates primarily through canopy architecture modification, with leaf removal increasing radiative exposure and maximum temperatures while potentially improving nocturnal cooling, and canopy shading reducing temperature extremes but limiting light-dependent phenolic synthesis.
Fruit_Zone_Temperature_Quality is_a Canopy_Microclimate_Quality, representing the thermal dimension of fruit zone environmental conditions. This class addresses temperature at cluster positions specifically, recognizing that berry temperatures can substantially exceed ambient air temperature under direct solar exposure. Key thresholds include ~25°C for accelerated malic acid respiration, 30-35°C for anthocyanin synthesis inhibition, and 40-45°C for sunburn damage onset depending on cultivar. The balance between beneficial warming for ripening acceleration and damaging heat stress represents core management challenge.
Fruchtzonentemperatur
Traubenzonenklima
berry temperature
bunch zone temperature
cluster temperature
fruit zone thermal regime
Spayd SE et al. (2002) Separation of sunlight and temperature effects on the composition of Vitis vinifera cv. Merlot berries. American Journal of Enology and Viticulture 53:171-182 Tarara JM et al. (2008) Berry temperature and solar radiation alter acylation, proportion, and concentration of anthocyanin in Merlot grapes. American Journal of Enology and Viticulture 59:235-247 Greer DH, Weston C (2010) Heat stress affects flowering, berry growth, sugar accumulation and photosynthesis of Vitis vinifera cv. Semillon grapevines grown in a controlled environment. Functional Plant Biology 37:206-214
Fruit Zone Temperature Quality
Fruitfulness varies widely across V. vinifera cultivars, with Trebbiano Toscano averaging 2.5-3.2 clusters per shoot compared to 1.2-1.8 for Cabernet Sauvignon under similar management. The fruitfulness coefficient (clusters per shoot) shows cultivar-specific responses to environmental conditions, with a coefficient of variation of 15-35% across years. Total potential yield correlates strongly with bud fruitfulness (r = 0.75-0.85) when fruit set and berry weight are consistent. Fruitfulness assessment during dormancy through bud dissection enables crop forecasting with 80-90% accuracy.
A reproductive disposition that is the inherited capacity of a Vitis vinifera plant to initiate and develop inflorescence primordia in latent buds that will produce fruit-bearing shoots in the following season, determining the potential yield through the number and distribution of clusters per vine.
Fruitfulness_Disposition serves as the parent class for traits related to inflorescence primordia formation in buds. Fruitfulness is determined during the previous growing season (year N-1) when primordia differentiate in developing buds, making it responsive to environmental conditions 12-14 months before harvest. The trait is distinct from fruit set, which determines what proportion of flowers become berries after fruitfulness has established cluster number potential.
Ertragspotential
Fruchtbarkeit
Fruchtbarkeitsdisposition
bud fertility disposition
fertility trait
reproductive potential
Sánchez LA, Dokoozlian NK (2005) Bud microclimate and fruitfulness in Vitis vinifera L. American Journal of Enology and Viticulture 56:319-329 Dry PR (2000) Canopy management for fruitfulness. Australian Journal of Grape and Wine Research 6:109-115
Fruitfulness Disposition
A fruiting cane on Vitis vinifera cv. Pinot Noir in a Scott Henry training system, selected during January pruning from well-positioned shoots originating from previous season's renewal cane, measuring 115cm length with 10 nodes retained after tip removal, 9mm average diameter showing complete brown lignification, positioned along lower trellis wire with buds facing upward for optimal shoot orientation, expected to produce 9-10 fruiting shoots (90% budbreak typical for well-lignified mid-cane positions) bearing 16-18 clusters totaling 4.2-4.8kg fruit assuming balanced vine capacity and appropriate environmental conditions during flowering and fruit set.
A one-year cane retained during winter pruning specifically to serve as current season crop production wood, characterized by appropriate length for desired bud retention (typically 6-15 nodes), adequate lignification for winter survival and structural support, positioning suitable for trellis integration and shoot management, and expectation of producing multiple fruiting shoots bearing harvestable grape clusters from its compound buds. Fruiting canes represent the primary production unit in cane-pruned training systems and the shortened equivalent (spurs) in spur-pruned systems, with their selection, number, length, and positioning determining potential crop load, canopy architecture, fruit exposure, and ultimately yield quantity and quality, making fruiting cane management the central decision point in annual winter pruning protocols that balance productivity with vine sustainability.
Fruiting canes are distinguished from renewal canes (retained for following season's fruiting wood production rather than current crop), water shoots or suckers (unwanted vegetative growth), and permanent structure (multi-year arms, cordons, trunks). Selection criteria include appropriate diameter (8-12mm ideal, avoiding overly vigorous or weak wood), complete lignification, node spacing indicating balanced growth, positioning allowing shoot management without crowding, and location maintaining desired vine architecture. The number and length of fruiting canes retained depends on vine size and vigor (balanced pruning formulas), training system requirements, labor availability for shoot management, and quality versus quantity objectives, with longer canes and higher bud counts favoring yield while shorter canes and lower counts favor fruit quality.
Ertragsrute
Tragrute
bearing cane
crop cane
production cane
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 89-112 Bravdo et al. (1985) "Effect of crop level and crop load on growth, yield, must and wine composition, and quality of Cabernet Sauvignon" American Journal of Enology and Viticulture 36=125-131.
Fruiting Cane
A VSP-trained Pinot Noir vine has a Fruiting_Zone_Role positioned at 70-100 cm height, the narrow vertical band between cordon and the first catch wire receiving targeted leaf removal to achieve 40-60% cluster exposure for optimal color and flavor development in cool-climate conditions. A GDC-trained Concord vine bears Fruiting_Zone_Roles on the exterior surfaces of both pendant curtains, with fruit positioned for mechanical harvester access and sufficient exposure for juice quality despite the high-yield production system. A traditional gobelet-trained Grenache has a Fruiting_Zone_Role distributed throughout the interior of the bush form, the self-shading architecture protecting fruit from the intense Mediterranean sun that would damage exposed clusters.
A canopy position role that inheres in the spatial region of a grapevine canopy where clusters develop and mature, defined by the architectural positioning of fruiting structures within the vine's three-dimensional form. The fruiting zone represents the most management-critical canopy region, as conditions within this zone—light exposure, temperature, humidity, air movement—directly influence fruit composition and wine quality through effects on sugar accumulation, acid metabolism, phenolic development, and aromatic compound synthesis. Fruiting zone characteristics vary dramatically with training system: VSP systems create narrow, defined fruiting zones at cordon height sprawling or bush systems distribute fruit throughout the canopy volume divided canopy systems create multiple distinct fruiting zones with different exposure characteristics. Fruiting zone management encompasses practices specifically targeting this zone—leaf removal to enhance exposure, shoot positioning to optimize fruit placement, selective harvest of zone-differentiated fruit—all recognizing that fruiting zone conditions are manipulable independently of overall vine management. The concept of defined fruiting zones, with quantifiable exposure metrics and targeted management interventions, represents a core principle of modern canopy management.
Fruiting_Zone_Role captures the spatial concept fundamental to targeted canopy management, recognizing that fruit-bearing canopy regions require distinct management attention. Fruiting zone definition varies with training system architecture and management philosophy—some systems create highly defined zones with uniform exposure, while others accept or intentionally create exposure variation within the fruiting zone. The relationship between fruiting zone microclimate and wine quality has been extensively researched, with exposure metrics (percent ambient light, sunfleck patterns, temperature accumulation) correlated with fruit composition parameters across varieties and regions. Climate change is shifting optimal fruiting zone management—practices that enhanced exposure in cool climates may cause damage in warming conditions, prompting reconsideration of leaf removal intensity, row orientation, and other fruiting zone management decisions.
Beerenzone
Fruchtzone
Traubenbereich
Traubenzone
bunch zone
cluster zone
fruit zone
fruiting zone
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Dokoozlian NK, Kliewer WM (1995) Light environment within grapevine canopies, American Journal of Enology and Viticulture 46: 209-218 Bergqvist J et al. (2001) Sunlight exposure and temperature effects on berry growth and composition, American Journal of Enology and Viticulture 52: 1-7 Spayd SE et al. (2002) Separation of sunlight and temperature effects, American Journal of Enology and Viticulture 53: 171-182
Fruiting Zone Role
Fruity aroma assessment in Viognier reveals characteristic profile dominated by stone fruit (apricot, peach from C13-norisoprenoids and lactones), complemented by tropical notes (mango, lychee from terpenes and thiols) and citrus undertones (orange blossom), with the stone fruit dominance distinguishing Viognier from Gewürztraminer where tropical and lychee notes predominate, demonstrating how the specific fruit subcategory balance creates varietal aromatic identity within the broader fruity character category.
Fruity aroma quality is an aroma character quality that represents olfactory perception evoking fruit-associated scents, encompassing the broad category of aromatic descriptors ranging from fresh citrus through tropical and stone fruits to berry and dried fruit notes that collectively constitute the most prominent and positively valued aroma category in wine evaluation. Within the BFO framework, this quality addresses the major aromatic category most associated with wine quality and varietal expression, distinguishing fruit-derived and fruit-reminiscent aromas from other character categories while recognizing that fruity character arises from diverse volatile compounds through both direct compositional correspondence and perceptual similarity. The ontological significance lies in fruity aroma serving as the primary positive aromatic attribute across wine styles, where fruit character expression indicates varietal typicity, appropriate ripeness, successful fermentation, and overall quality, with the specific fruit profile distinguishing varieties, regions, and vintages more than any other aroma category. The compositional basis for fruity aroma includes multiple compound classes, with esters (ethyl and acetate esters from fermentation) providing generic fruity background, terpenes (linalool, geraniol) contributing specific fruit notes in aromatic varieties, thiols (3MH, 4MMP) creating tropical fruit character in Sauvignon Blanc and related varieties, and norisoprenoids (β-damascenone, β-ionone) adding stone fruit and berry complexity. The developmental trajectory of fruity aroma proceeds from grape-derived varietal fruit character (primary aroma) through fermentation-enhanced ester fruitiness (secondary aroma) to evolved fruit notes in aged wines where fresh fruit transforms toward dried and compote character. The subcategories of fruity aroma (citrus, tropical, stone fruit, pome fruit, berry, dried fruit) reflect both perceived similarity groupings and underlying compositional relationships, with different varieties and styles showing characteristic profiles across these subcategories. Understanding fruity aroma quality enables varietal identification based on fruit profile, ripeness assessment from fruit character development, and quality evaluation where appropriate fruity expression indicates successful viticulture and winemaking.
Fruity_Aroma_Quality serves as the parent class for fruit subcategories (Citrus, Tropical, Stone Fruit, Pome Fruit, Berry, Dried Fruit), representing the most prominent and positively valued aroma category in wine. This quality encompasses both grape-derived varietal fruit character and fermentation-enhanced fruitiness. The distinction among fruit subcategories reflects both perceptual grouping and underlying compositional differences, with characteristic profiles distinguishing varieties and styles.
Fruchtaroma
Fruchtcharakter
Obstduft
fruchtiger Duft
fruit aroma
fruit character
fruit-like scent
fruity nose
Escudero A et al. (2007) Characterization of the aroma of a wine from Maccabeo. Key role played by compounds with low odor activity values. Journal of Agricultural and Food Chemistry 55(11): 4501-4510 Francis IL, Newton JL (2005) Determining wine aroma from compositional data. Australian Journal of Grape and Wine Research 11(2): 114-126
Fruity Aroma Quality
The full BBCH code 81 recorded for Tempranillo in Rioja on August 10 encodes the precise developmental moment of véraison onset — principal stage 8 (ripening) combined with secondary stage 1 (beginning of ripening, berries begin to develop color) — and this two-digit code serves as the standard entry in phenological databases, the trigger point for irrigation regime changes (shifting from moderate to severe deficit irrigation at BBCH 81), the reference stage for predicting harvest date (typically 40-55 days after BBCH 81 for Tempranillo in Rioja), and the mandatory staging reference for EU-registered plant protection product application windows specifying BBCH 81-85 for botryticide application in ripening grape clusters.
A BBCH code consisting of the complete two-digit numerical expression formed by combining a principal growth stage code (first digit) with a secondary growth stage code (second digit), constituting the standard unit of phenological recording that unambiguously identifies a specific grapevine developmental stage within the BBCH classification system.
Full_BBCH_Code is the operational unit of the BBCH staging system — the complete two-digit expression that appears in phenological records, regulatory documents, precision viticulture decision systems, and scientific publications. The code's power lies in its combination of compactness (two digits), universality (applicable across cultivars, regions, and management systems), and standardization (identical coding convention worldwide): a researcher in New Zealand reading BBCH 65 in a German publication instantly recognizes full flowering without requiring conversion between incompatible local staging systems. Approximately 47 full BBCH codes are formally defined for Vitis, spanning from BBCH 00 (dormancy) through BBCH 97 (end of leaf fall), with the defined codes providing sufficient resolution for most viticultural management decisions and phenological research applications.
BBCH stage identifier
BBCH-Gesamtcode
complete BBCH code
two-digit BBCH stage
vollständiger BBCH-Code
zweistelliger BBCH-Wert
Full BBCH Code
Full body quality in Barossa Valley Shiraz (15% alcohol, substantial extract, body rating 9/10) provides the powerful structure supporting the style's dark fruit, chocolate, and spice intensity, creating the rich, enveloping mouthfeel appropriate for winter evening contemplation or pairing with char-grilled beef, with the substantial weight demanding focused attention and limiting consumption to moderate quantities while rewarding the concentration with complex, layered sensory experience.
Full body quality is a body quality that represents substantial perceived weight and powerful palate presence, characterizing the rich, dense mouthfeel typical of high-alcohol wines, warm-climate expressions, concentrated styles, and wines intended for extended aging where structural weight supports complex flavor development and food pairing with substantial dishes. Within the BFO framework, this quality specifically addresses the heavy end of the body spectrum, distinguishing powerful weight from medium and light body while recognizing that full body represents a stylistic choice appropriate for specific wine categories rather than a universal quality ideal. The ontological significance lies in full body defining wines of power and concentration valued for special occasions, rich cuisine pairing, and meditative consumption, where substantial weight creates the sensory impact appropriate for serious wine appreciation and matches the intensity of hearty dishes that would overwhelm lighter wines. The perceptual characteristics of full body include substantial coating of oral surfaces, significant resistance to oral manipulation, persistent palate impression after swallowing, and the sense of richness and density that demands attention and rewards focused appreciation. The compositional profile associated with full body typically includes elevated alcohol (13.5-16%+), often residual sugar in off-dry to sweet styles, high extract from extended maceration or concentration techniques, and often lower acidity contributing to the impression of weight without counterbalancing lift. The variety and regional associations of full body include Barossa and McLaren Vale Shiraz, Napa Valley Cabernet Sauvignon, Châteauneuf-du-Pape, Amarone, vintage Port, Sauternes and other botrytized wines, and oaked Chardonnay from warm regions, with these wines valued specifically for their powerful presence and concentration. Understanding full body quality enables appropriate style recognition, quality assessment valuing power and concentration in suitable contexts, and food pairing recommendations matching full-bodied wines with substantial cuisine.
Full_Body_Quality addresses the powerful end of the body spectrum, representing the wines of concentration and impact valued for special occasions and substantial cuisine pairing. This quality is stylistically appropriate for specific wine categories rather than representing universal quality ideal. The class supports recognition of power and concentration as positive attributes in suitable contexts while acknowledging that full body limits versatility compared to lighter styles.
concentrated mouthfeel
full weight
heavy body
konzentriertes Mundgefühl
kraftvolle Fülle
powerful texture
schwerer Körper
voller Körper
Jackson RS (2014) Wine Science: Principles and Applications, 4th Ed. Academic Press, San Diego, pp. 831-887 Nurgel C, Pickering GJ (2005) Contribution of glycerol, ethanol and sugar to the perception of viscosity and density elicited by model white wines. Journal of Texture Studies 36(3): 303-323
Full Body Quality
In Cabernet Sauvignon vines in the Médoc, Bordeaux, full chill accumulation is achieved by late January when integrated cold exposure reaches approximately 1000 chilling hours, with accumulation proceeding from approximately 200 hours by late November through 600 hours by early January to full accumulation by January 25, tracked using the hours-below-7.2°C model and verified by forcing tests showing budbreak within 11 days indicating complete dormancy release.
Full chill accumulation in Vitis vinifera represents the complete integration of effective cold exposure over the winter period resulting in total satisfaction of the chilling requirement, encompassing the cumulative physiological effects of all chilling temperatures experienced from dormancy establishment through requirement completion. This comprehensive accumulation reflects the summation of effective cold exposure according to temperature-dependent effectiveness functions, with the full accumulation representing the total chilling credit acquired through winter that equals or exceeds the genetically determined requirement for complete dormancy release. The physiological basis of full accumulation involves the progressive and persistent effects of cold exposure on dormancy-maintaining mechanisms, with each increment of effective chilling contributing to the cumulative reduction of ABA through catabolic activation, the progressive silencing of DAM gene expression, and the gradual removal of repressive chromatin modifications, with full accumulation representing the completion of these cumulative changes. The quantification of full accumulation depends on the chilling model employed, with simple models summing hours below a threshold temperature to yield totals typically ranging from 400-1500 hours depending on cultivar, while more sophisticated models apply temperature-dependent weighting functions or dynamic calculations that account for the varying effectiveness of different temperatures and potential negation by warm exposure. The temperature range most effective for chill accumulation in grapevine spans approximately 0-7°C, with temperatures in this range providing maximum contribution to accumulated chilling, temperatures below freezing being less effective or ineffective, and temperatures above approximately 15°C potentially negating previously accumulated chilling in some model formulations. The seasonal pattern of chill accumulation typically shows slow initial accumulation during autumn as temperatures fluctuate around effectiveness thresholds, accelerating accumulation during consistent winter cold, and eventual completion when cumulative effective exposure reaches the requirement threshold. The verification of full accumulation relies on forcing tests confirming rapid budbreak response, with full accumulation demonstrated by the minimum forcing times and maximum budbreak percentages characteristic of completely released dormancy.
Full_Chill_Accumulation represents the quantitative completeness aspect of Chilling_Completion addressing total integrated cold exposure. Distinguished from partial accumulation during endodormancy by representing requirement satisfaction. Quantified through various models from simple hour counting to dynamic calculations. Effective temperature range approximately 0-7°C. Seasonal progression from slow autumn accumulation through winter completion. Verified by forcing test minimum response times.
complete chilling accumulation
cumulative chilling
integrated cold exposure
integrierte Kälteexposition
komplette Chilling-Ansammlung
kumulative Kälteeinwirkung
total chill accumulation
vollständige Kälteakkumulation
Richardson EA, Seeley SD, Walker DR (1974) A model for estimating the completion of rest for Redhaven and Elberta peach trees. HortScience 9: 331-332 Fishman S, Erez A, Couvillon GA (1987) The temperature dependence of dormancy breaking in plants: mathematical analysis of a two-step model. Journal of Theoretical Biology 126: 309-321 Dokoozlian NK (1999) Chilling temperature and duration interact on the budbreak of Perlette grapevine cuttings. HortScience 34: 1054-1056
Full Chill Accumulation
A full cluster secondary bud at node 6 on Vitis vinifera cv. Riesling, activated following 91% primary bud mortality from severe spring frost at -4.1°C, dissection revealing one well-formed inflorescence primordium containing 118 flower initials plus vegetative shoot primordia with 9 leaf initials, developing into a shoot bearing one cluster of 56 berries (47% fruit set typical for Riesling) weighing 142g at harvest, plus 14 leaves providing 795 cm² photosynthetic area, achieving 14.2 cm²/g leaf area-to-fruit ratio within acceptable range for quality production, enabling partial vineyard economic viability (estimated 48% normal revenue) despite catastrophic primary loss.
A secondary reproductive bud containing well-developed inflorescence primordia approaching primary bud quality in terms of flower number and potential cluster size, representing the upper range of secondary fruitfulness found in cultivars with strong genetic predisposition toward reproductive capacity in secondary positions, characterized by inflorescence initials with 80-150+ flower primordia (50-75% of typical primary counts), development into shoots bearing substantial harvestable clusters comparable to smaller primary clusters, and occurrence primarily in high-secondary-fruitfulness cultivars (Chardonnay, Riesling, Gewürztraminer) at favorable node positions. Full cluster secondary buds enable significant yield recovery following primary bud damage, potentially achieving 40-60% of normal production levels when widely activated, making their prevalence a critical cultivar characteristic for risk management in frost-prone viticultural regions.
Full cluster secondary buds exemplify the value of cultivar selection for frost-prone regions, where varieties like Chardonnay and Riesling showing 45-60% secondary positions with full clusters can maintain partial economic viability following severe frost damage, while low-secondary-fruitfulness cultivars may experience near-total crop loss from equivalent damage. The mechanisms underlying cultivar variation in secondary fruitfulness remain incompletely understood but likely involve genetic differences in meristem size, resource allocation priorities during differentiation, and hormonal regulation of inflorescence versus vegetative fate determination. Plant breeders increasingly recognize secondary fruitfulness as a valuable trait for climate adaptation, with selection programs incorporating secondary bud dissection assessments alongside traditional primary fruitfulness evaluation to develop cultivars combining productivity with resilience.
high-fruitfulness secondary bud
productive secondary bud
produktive Nebenknospe
vollständige Trauben-Sekundärknospe
well-developed secondary reproductive bud
Based on Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Molitor et al. (2014) "Late frost damage risk for viticulture under future climate conditions= a case study for the Luxembourgish winegrowing region" Australian Journal of Grape and Wine Research 20=160-168 Trought & Bramley (2011) "Vineyard variability in Marlborough, New Zealand= characterising spatial and temporal changes in fruit composition and juice quality in the vineyard" Australian Journal of Grape and Wine Research 17=79-89 Vasconcelos et al. (2009) "The flowering process of Vitis vinifera= a review" American Journal of Enology and Viticulture 60=411-434.
Full Cluster Secondary Bud
Full stage expression assessment for berry development in optimally managed Shiraz reveals the complete developmental achievement possible with appropriate management, with Phase I showing full expression (berry volume reaching 1.2 mL by lag phase, complete cell division producing 250,000 cells/berry, seed development complete with viable embryos), veraison showing full expression (complete color development in 12 days, softening uniform across cluster, acid decline and sugar accumulation initiating synchronously), and ripening showing full expression (sugars reaching 24 Brix, anthocyanins 2.1 mg/g, tannin polymerization complete, flavor volatiles fully developed), demonstrating the complete developmental potential achievable under optimal conditions.
Full stage expression quality is a developmental completeness quality that represents the achievement of complete developmental potential within a phenological stage, characterizing the ideal condition where all normal processes within a stage proceed to completion, organs achieve their full developmental potential, and the stage provides an optimal foundation for subsequent phenological progression. Within the BFO framework, this quality specifically addresses the complete achievement dimension of developmental stages, distinguishing full expression from partial or arrested development while recognizing that full stage expression represents the benchmark against which incomplete development is assessed and the goal that management seeks to achieve at each phenological phase. The ontological significance lies in full stage expression establishing the quality potential that each phenological phase can contribute to ultimate fruit quality, where complete bud development produces fruitful buds, complete canopy development provides adequate photosynthetic capacity, complete fruit set establishes appropriate crop load, and complete ripening achieves target fruit composition, with full expression at each stage providing the cumulative foundation for quality outcomes. The criteria for full stage expression vary among stages but include complete structural development (all normal organ components present and properly formed), complete physiological function (all normal processes occurring at appropriate rates), and complete preparation for subsequent stages (all prerequisites for next phase development accomplished). The environmental and management requirements for full stage expression include adequate water supply (supporting cell expansion and physiological processes), adequate nutrition (providing building blocks for structural and metabolic development), appropriate temperature range (enabling normal developmental processes without stress), and balanced vine condition (source-sink relationships supporting development without excessive competition). The cultivar and site interactions affecting full expression show that some combinations consistently achieve complete development (well-adapted cultivars on favorable sites) while others struggle (marginal cultivar-climate matches, challenging soil conditions), with management intensity requirements varying accordingly. Understanding full stage expression quality enables definition of developmental targets, assessment of achievement relative to potential, and management guidance for supporting complete development at each phenological stage.
Full_Stage_Expression_Quality addresses the complete achievement of developmental potential within each phenological stage. This quality represents the benchmark for assessing developmental success at each phase. The class supports management target setting and achievement assessment through full expression characterization.
Entwicklungserreichung
complete development
developmental achievement
full phenological expression
optimal stage completion
optimaler Stadienabschluss
vollständige Entwicklung
vollständige Stadienexpression
Coombe BG, McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6(2): 131-135 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 59-144
Full Stage Expression Quality
Wild V. vinifera ssp. sylvestris female plants in Mediterranean riparian populations during peak bloom (BBCH 65, late May-early June) exhibit fully developed female only flowers with calyptra shed, pistil fully erect (2.5-2.7 mm length, ovary 1.5-1.6 mm diameter), stigma glistening with abundant wet exudate and elaborate papillae (220-245 papillae), achieving peak receptivity persisting 3-4 days with successful fruit set (38-44%) when wind-borne pollen from nearby staminate males (within 15-35 m) arrives during this receptivity window, but reduced fruit set (15-22%) when temporal asynchrony causes pollen arrival before or after peak receptivity period. Individual female plants bearing 250-500 fully developed flowers simultaneously (progressing acropetally within inflorescences creating 6-10 day total bloom period) provide extended opportunity for successful pollination across variable male bloom timing and stochastic pollen dispersal events.
A fully developed female only flower is a female only flower that has completed all developmental stages and achieved full anthesis with mature pistil exhibiting maximum stigma receptivity and optimal conditions for successful cross-pollination and fertilization, representing the functionally active reproductive state of female only flowers at peak receptivity for pollen capture. Fully developed female only flowers are characterized by= (1) complete calyptra shedding exposing the well-developed pistil and the empty stamen positions (2) pistil fully elongated to maximum length (typically 2.3-2.8 mm) with erect, prominent positioning optimizing stigma exposure to wind-borne pollen (3) stigma at peak receptivity with maximum wet exudate production creating glistening stigmatic surface, elaborate papillae fully extended (200-250 papillae in optimal development), and biochemical conditions optimal for pollen grain hydration, germination, and pollen tube initiation (4) ovary fully developed containing 3-4 viable ovules at developmental stage ready for fertilization, with embryo sacs mature and egg cells prepared for sperm cell fusion (5) style fully elongated with transmitting tract tissue mature and ready to support pollen tube growth from stigma to ovules (6) flower receptacle and nectary disk fully developed, though nectary function may be reduced compared to hermaphrodite flowers as insect pollination is secondary to wind pollination in Vitis. The 'fully developed' designation indicates completion of the developmental program and achievement of peak functional receptivity for cross-pollination, contrasting with pre-anthesis developmental stages or post-receptivity senescent stages. Fully developed female only flowers represent the critical window for successful cross-pollination, with stigma receptivity persisting at peak levels for 2-4 days post-calyptra shedding under optimal conditions (moderate temperature 18-26°C, moderate humidity 55-75%), then declining gradually over subsequent 3-5 days as stigmatic exudate production decreases and papillae begin senescence. The timing of transition to fully developed state follows temperature-dependent kinetics similar to hermaphrodite flowers, with calyptra shedding occurring in early morning (typically 6-10 AM) and stigma achieving peak receptivity by mid-day (11 AM-2 PM) when stigmatic exudate production maximizes. Fully developed female only flowers exhibit optimal pollen capture efficiency during the peak receptivity window, with the elaborate stigmatic papillae and abundant wet exudate creating effective pollen adhesion surfaces that can capture wind-borne pollen grains arriving at low flux rates (as few as 5-15 pollen grains landing on stigma surface may be sufficient for successful fertilization of multiple ovules given high pollen viability from staminate sources). The probability of successful cross-pollination and fruit set depends critically on temporal overlap between female only flower full development and pollen availability from nearby staminate or hermaphrodite sources, with fruit set maximized (35-45%) when fully developed female flowers coincide with peak pollen dispersal from males, but reduced substantially (12-25%) when bloom asynchrony creates temporal mismatch. Fully developed female only flowers on wild dioecious V. vinifera ssp. sylvestris female plants provide a limited receptivity window (2-4 day peak plus 3-5 day declining receptivity) during which successful pollination must occur, creating selection pressure for= (1) extended stigma receptivity maximizing temporal window (2) enhanced stigmatic surface area maximizing spatial pollen capture (3) synchronization mechanisms aligning female bloom timing with male pollen dispersal periods in the local population.
Fully_Developed_Female_Only_Flower represents the functionally active anthesis stage at peak receptivity and is distinguished from Mature_Female_Only_Flower by emphasis on active pollen reception (fully developed, peak stigma receptivity) versus achieved developmental maturity that may extend beyond peak receptivity (mature, including pre-peak and post-peak stages). The distinction parallels that made for staminate flowers between fully developed (peak pollen release) and mature (broader developmental window). Fully developed female only flowers represent the critical reproductive window when successful cross-pollination probability is maximized, making this developmental stage of particular importance for understanding reproductive success in dioecious populations and for managing controlled pollination in breeding programs using female only parents.
anthetic female flower
pollen-receptive female flower
reife rezeptive Blüte
voll ausgebildete Pistillatblüte
voll entwickelte weibliche Blüte
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 103-108 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 99-108 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 190-198.
Fully Developed Female Only Flower
Wild V. vinifera ssp. sylvestris male plants in southern France during peak bloom (BBCH 65, late May-early June) exhibit fully developed male only flowers with calyptra shed, stamens fully erect (filaments 3.8-4.4 mm), anthers dehisced and releasing clouds of yellow pollen (40,000-47,000 grains per flower, 84-90% viability), creating visible pollen dispersal plumes extending 80-150 m downwind and successfully fertilizing pistillate female plants within that range, with individual male plants bearing 800-1500 fully developed male only flowers simultaneously providing sufficient pollen for 25-40 female plants. Staminate reversion shoots on cultivated Pinot Noir occasionally produce 15-35 fully developed male only flowers creating localized pollen clouds visible during bloom period, potentially contributing to cross-pollination of nearby functionally female cultivars if present.
A fully developed male only flower is a male only flower that has completed all developmental stages and achieved full anthesis with mature, dehisced anthers actively releasing pollen, representing the functionally active reproductive state of male only flowers at peak pollen production and dispersal capability. Fully developed male only flowers are characterized by= (1) complete calyptra shedding exposing the five stamens and empty flower center (2) stamens fully elongated to maximum length (typically 3.5-4.5 mm filaments) with erect, spread positioning optimizing pollen dispersal (3) anthers fully mature, dehisced through longitudinal slits, and actively releasing pollen grains (4) pollen at peak viability (typically 78-92% viable grains immediately post-dehiscence) and maximum abundance (35,000-50,000 grains per flower fully released or readily releasable) (5) flower receptacle and nectary disk fully developed and functional, though nectary function in staminate flowers is reduced compared to hermaphrodite flowers as pollinator attraction is less critical for wind-pollinated Vitis. The 'fully developed' designation indicates completion of the developmental program and achievement of functional reproductive maturity, contrasting with pre-anthesis developmental stages or post-anthesis senescent stages. Fully developed male only flowers represent the critical window for pollen dispersal, typically persisting in peak functional state for 24-48 hours post-calyptra shedding under optimal conditions (moderate temperature 20-28°C, moderate humidity 50-70%), with pollen viability declining thereafter as released pollen ages and anthers deplete their pollen reserves. The timing of transition to fully developed state follows temperature-dependent kinetics similar to hermaphrodite flowers, with calyptra shedding and anther dehiscence occurring in early morning (typically 6-10 AM) on days when temperature and humidity conditions are favorable, and pollen release peaking in late morning to early afternoon (10 AM-2 PM) when declining humidity and increasing temperature create optimal pollen dispersal conditions. Fully developed male only flowers produce visible pollen clouds when disturbed by wind or mechanical agitation, with the abundant pollen appearing as yellow dust coating nearby surfaces and creating conspicuous evidence of male flower activity. The pollen dispersal range from fully developed male only flowers can extend 50-200 m downwind under moderate wind conditions (3-8 m/s), substantially exceeding the 10-30 m effective pollination range from hermaphrodite flowers due to the 2-3× greater pollen production per flower and the presence of numerous male only flowers on dedicated male plants (dioecious male plants may bear 500-2000+ male only flowers versus 100-300 hermaphrodite flowers on typical hermaphrodite cultivar shoots). Fully developed male only flowers on wild dioecious V. vinifera ssp. sylvestris male plants or on staminate reversion shoots in cultivated vineyards serve as highly effective pollen sources, capable of fertilizing pistillate or functionally female flowers over considerable distances and potentially contributing to unintended cross-pollination in cultivar identity maintenance scenarios. The anthesis phenology of fully developed male only flowers shows acropetal progression within inflorescences (basal flowers achieving fully developed state 1-3 days before apical flowers) and typically proceeds 12-36 hours faster than hermaphrodite flower anthesis on the same site due to absence of pistil development that may slow the calyptra shedding process in hermaphrodite flowers where both male and female organs must coordinate maturation.
Fully_Developed_Male_Only_Flower represents the functionally active anthesis stage and is distinguished from Mature_Male_Only_Flower by emphasis on active pollen release (fully developed) versus achieved developmental maturity that may extend beyond peak pollen dispersal (mature). The distinction is subtle but reflects the difference between peak functional state (pollen actively dispersing) versus completed development (which includes post-dispersal senescent stages). Fully developed male only flowers represent the critical reproductive window when male plants maximize their pollen contribution to population reproduction in dioecious systems. The timing, duration, and environmental conditions affecting the fully developed state are critical for understanding pollination dynamics in wild dioecious Vitis populations and for managing unintended cross-pollination in cultivated vineyards containing staminate reversions or wild male plants in proximity.
anthetic male flower
pollen-releasing male flower
reife pollenproduzierende Blüte
voll ausgebildete Staminatblüte
voll entwickelte männliche Blüte
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 102-106 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 98-108 Lebon, G. et al. (2008). Sugars and flowering in the grapevine. Journal of Experimental Botany 59=2565-2578.
Fully Developed Male Only Flower
Pinot Noir fully developed mature own-rooted trunk in Burgundy at 11 years post-planting shows 64-78 mm diameter with 11 annual rings (recent 3 years averaging 2.8-3.2 mm width), sapwood 22-28 mm radial depth with excellent water conductance providing 42-58 L/day capacity, heartwood 28-36 mm diameter, 135-165 g starch reserves, and peak productivity of 3.8-5.2 kg per vine (72-88 hL/ha) with excellent fruit quality and vintage consistency
A fully developed mature own-rooted trunk is a mature own-rooted trunk at peak developmental status (typically years 8-15 post-planting) characterized by optimal trunk diameter for training system, maximum functional sapwood cross-section, peak carbohydrate storage capacity, and highest productivity potential, representing the culmination of trunk development before gradual senescence processes begin. Fully developed mature own-rooted trunk characteristics include= trunk diameter approaching cultivar and site maximum for sustained health (typically 70-110 mm for moderate-vigor wine grape cultivars at year 10-12), sapwood cross-section at maximum radial extent (20-35 mm radial depth providing optimal water transport capacity), heartwood occupying 30-50% of trunk cross-section (providing mechanical support while maintaining adequate sapwood conducting area), annual ring width stabilizing at mature values (2-4 mm radial increment compared to 3-6 mm in young trunks), and bark achieving mature thickness and protective capacity (5-8 mm providing thermal insulation and pathogen barrier). Fully developed mature own-rooted trunk anatomical optimization includes= maximum number of functional xylem vessels in sapwood (providing 40-100 L/day water transport capacity depending on canopy size), optimal phloem cross-section in inner bark (2-3 mm active phloem supporting efficient sugar and nutrient distribution), well-developed ray parenchyma system (radial parenchyma cells providing lateral transport and storage), mature periderm structure (cork layers providing protection without excessive thickness restricting trunk expansion), and stable cambial activity (consistent secondary growth without excessive vigor or premature decline). Fully developed mature own-rooted trunk physiological optimization includes= peak carbohydrate storage capacity (120-180 g starch reserves providing optimal spring growth support and stress buffering), maximum photosynthetic efficiency per unit crop (established vine architecture and physiology supporting efficient carbon assimilation), optimal hydraulic conductance per unit leaf area (sapwood providing adequate water supply without excessive oversizing), balanced hormone production and distribution (appropriate levels of auxins, cytokinins, gibberellins supporting healthy development), and established stress response mechanisms (drought tolerance, cold hardiness, disease resistance at peak levels). Fully developed mature own-rooted trunk productivity characteristics include= maximum sustainable yield potential (typically 4-8 kg per vine, 60-100 hL/ha at typical densities, varying by cultivar, training system, and quality goals), optimal fruit quality potential (established vine physiology supporting consistent berry composition and ripening), uniform cluster development (consistent flower initiation, fruit set, and berry development across vine), minimal year-to-year variation in absence of extreme weather (stable productivity reflecting mature vine homeostasis), and excellent recovery capacity from moderate stress or crop adjustment. Fully developed mature own-rooted trunk represents optimal commercial vineyard status= peak economic return on vineyard investment (maximum yield with optimal quality), lowest per-unit production costs (established vines requiring minimal remedial intervention), greatest flexibility for crop management (adequate reserves supporting various crop load strategies), optimal terroir expression (mature vine physiology fully expressing site characteristics), and foundation for sustained productivity (healthy trunk supporting 10-20+ additional years production before significant senescence). Fully developed mature own-rooted trunk management considerations include= maintaining optimal crop load preventing overcropping stress (balancing yield and quality while preserving vine health), protecting trunk from disease infection (especially important preserving healthy mature trunks with maximum productive potential), appropriate nutrition and irrigation (maintaining productivity without promoting excessive vigor), and monitoring for early senescence signs (trunk disease symptoms, declining vigor, reduced fruitfulness indicating potential issues). Fully developed mature own-rooted trunk duration varies= optimal sites may maintain fully developed status 8-20 years (years 8-28 post-planting), stressed sites may show briefer peak period 5-12 years (years 8-20), and exceptional sites with excellent management potentially extending peak 15-30 years (years 8-38).
Fully_Developed_Mature_Own_Rooted_Trunk emphasizes peak developmental status and is distinguished from Mature_Own_Rooted_Trunk (broader productive phase) and Productive_Mature_Own_Rooted_Trunk (sustained production emphasis) by focus on optimal anatomical and physiological state. Critical for understanding peak vineyard value period, optimal quality potential timing, trunk disease prevention importance, and economic analysis of vineyard productive lifespan.
Höchstreifer Rebstamm
Optimal entwickelter Eigenwurzelstamm
Optimal maturity trunk
Peak development trunk
Voll entwickelter reifer Stamm
Fully Developed Mature Own Rooted Trunk
Cabernet Sauvignon fully developed mature seeds at BBCH 89 measure 7.5-8.5 mm length with uniform black coloration, lignification index >95%, tannin polymerization 75-80%, moisture content 28-32%, and germination rates of 85-90% after 90-day cold stratification
A fully developed mature seed is a mature seed of Vitis vinifera that has completed all developmental stages including full embryo differentiation, maximum endosperm accumulation, complete testa lignification, and optimal desiccation, representing the highest quality seed state for both viticultural assessment and germplasm preservation. Fully developed mature seeds are characterized by= complete morphological development (uniform dark brown to black coloration, maximum size for genotype typically 6-9 mm length, well-defined beak and chalazal structures, symmetrical pyriform shape), complete anatomical maturation (fully lignified multi-layered testa with continuous sclereid layers, viable embryo with well-differentiated cotyledons and radicle, fully developed endosperm with accumulated lipid and protein reserves), and optimal biochemical composition (tannin polymerization index >70%, moisture content 25-35%, lipid content 10-20% dry weight, complete anthocyanin degradation in seed coat). These seeds represent successful completion of the entire developmental program from fertilization through final maturation, contrasting with seeds that achieve partial maturity but retain some immature characteristics. In wine grape production, fully developed mature seeds indicate optimal or slightly overripe harvest timing, as seed maturation typically precedes optimal berry sugar/acid balance by 1-2 weeks. The distinction between fully developed and merely viable mature seeds is important for breeding programs and germplasm conservation= fully developed seeds show maximum germination rates (80-95% after stratification), optimal seedling vigor, and best long-term storage viability. Seed coat characteristics of fully developed seeds include complete lignification without green or tan patches, uniform thickness of sclerified layers, and maximum accumulation of polymerized condensed tannins that are less extractable during crushing operations. Winemaking implications include minimal risk of harsh seed tannin extraction during fermentation, optimal ease of separation during pressing, and seed coat fragments that contribute less astringency to finished wines compared to immature seed material.
Fully_Developed_Mature_Seed emphasizes optimal complete developmental maturation and is distinguished from Viable_Mature_Seed (functional but potentially incomplete maturation) by achievement of maximal developmental potential across all morphological, anatomical, and biochemical parameters. Critical for breeding program seed quality assessment, germplasm conservation standards, and understanding harvest timing implications where seed over-maturity may indicate delayed harvest.
Completely mature seed
Optimal ausgereifter Kern
Optimal maturity seed
Vollreifer Traubenkern
Vollständig entwickelter Samen
Fully Developed Mature Seed
A fully expanded mature leaf at node 6 on a shoot of Vitis vinifera cv. Syrah in the Rhône Valley, France, measuring 15.2 cm length by 14.8 cm width representing maximum dimensions for this cultivar and mid-shoot position, with dimensions unchanged over four weeks of weekly measurements confirming expansion completion, exhibiting complete architectural development with five lobes showing final depth and configuration, 9.5 cm petiole length providing optimal orientation for light interception, all cells in lamina showing mature dimensions averaging 35 μm in palisade layers and 45 μm in spongy mesophyll without evidence of continued expansion, cell walls fully thickened and lignified in vascular and support tissues, occurring on a vigorous shoot (total length 175 cm) with excellent resource access supporting maximal leaf development, measured at BBCH stage 77 (berries beginning to touch) in mid-July at 5 weeks post-emergence, representing optimal achievement of maximum size potential with complete structural development enabling peak photosynthetic surface area for this node position.
A mature leaf that has achieved maximum lamina dimensions and complete structural development without further expansion in Vitis vinifera, characterized by attainment of cultivar-specific and position-specific final size typically 8-20 cm length and width depending on variety and environmental conditions, cessation of cell expansion with all cells reaching mature dimensions, stabilization of leaf architecture including fixed lobe depths, sinus configurations, and marginal characteristics, complete intercalary growth producing fully extended petiole length optimizing light capture positioning, and maintenance of stable dimensions over extended periods of weeks to months indicating developmental completion. Fully expanded mature leaves represent leaves that have completed the growth phase involving cell division in early development and cell expansion in subsequent weeks, achieving the architectural and dimensional characteristics that determine light interception capacity, photosynthetic surface area, and transpirational water loss, with final size determined by complex interactions between genetic programming, shoot vigor, resource availability during development, environmental conditions particularly temperature and water availability, and position effects with basal leaves typically larger than apical leaves on the same shoot. The transition to fully expanded status typically occurs 3-5 weeks post-emergence depending on environmental conditions and shoot vigor, marking achievement of maximum photosynthetic surface area and beginning of the extended period of functional photosynthetic contribution that may continue for 8-16 weeks until senescence initiation in autumn or stress-induced earlier degradation under adverse conditions.
Fully expanded mature leaves are distinguished from expanding juvenile leaves by cessation of dimensional increase and complete cell expansion, and from mature leaves that achieved less than maximum potential size due to stress or resource limitation during development. Assessment requires temporal monitoring showing dimensional stability over 1-2 weeks, indicating growth cessation rather than temporary pause, combined with anatomical examination confirming complete cell expansion without continued enlargement capacity. Final size achieved varies substantially with environmental conditions during development, with water stress, nutrient deficiency, or temperature extremes commonly limiting expansion resulting in smaller mature dimensions compared to optimal conditions. Position effects create systematic size gradients along shoots, with basal leaves (nodes 1-4) typically achieving largest dimensions, mid-shoot leaves (nodes 5-10) showing intermediate sizes, and apical leaves (nodes 11+) exhibiting smallest mature dimensions reflecting declining resource availability and shortened development periods in distal positions.
dimensionally complete leaf
full-size mature leaf
größenmäßig vollständiges Weinblatt
maximal ausgedehnte Blatt
maximum expansion leaf
voll expandiertes reifes Blatt
Based on Poni et al. (2009) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 60=397-407 Patakas et al. (1997) "Changes in water relations, photosynthetic capacity, and leaf anatomy induced by intermittent drought in two grapevine cultivars" Vitis 36=167-171 Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 95-118.
Fully Expanded Mature Leaf
Cabernet Sauvignon in premium Napa Valley sites under balanced crop load (1.2-1.5 kg fruit/m canopy) consistently exhibits fully functional hermaphrodite flowers at BBCH 65 with stamens extending 1-1.4 mm beyond pistil, producing 22,000-28,000 pollen grains per flower with 75-82% viability, and achieving 56-63% fruit set under favorable bloom conditions (late May-early June with daytime temperatures 25-28°C, overnight lows 12-15°C, relative humidity 60-75% at dawn declining to 35-45% mid-afternoon), yielding clusters of 90-140 berries from initial flower complements of 160-220 flowers per inflorescence. Syrah in warm, dry sites of Northern Rhône under moderate crop loads exhibits fully functional hermaphrodite flowers with particularly pronounced stamen extension (1.2-1.6 mm beyond pistil), 24,000-30,000 pollen grains per flower, 78-85% viability, and 58-65% fruit set in optimal vintage years.
A fully functional hermaphrodite flower is a hermaphrodite flower exhibiting optimal development and function of both male and female organs, with stamens extending substantially beyond the pistil, maximum pollen production and viability, optimal stigma receptivity and ovule viability, and consequently achieving the highest fruit set percentages observed in Vitis vinifera, representing the reproductive ideal for commercial fruit production. Fully functional hermaphrodite flowers are characterized by= (1) stamens extending 0.8-1.5 mm or more beyond pistil length, creating maximum spatial separation between anther dehiscence position and stigma, optimizing pollen shedding geometry for efficient self-pollination (2) anthers producing maximum pollen quantities (20,000-30,000+ pollen grains per flower) with high viability (70-85% viable pollen under optimal conditions) (3) pistil with fully developed ovary containing four viable ovules, functional style supporting rapid pollen tube growth (typical growth rate 3-5 mm/hour at 25°C), and highly receptive wet stigma with abundant exudate facilitating pollen grain hydration and germination (4) optimal developmental synchrony between pollen maturation and stigma receptivity, with stigma achieving peak receptivity coincident with or slightly before anther dehiscence, maximizing probability of successful pollination. The 'fully functional' designation indicates maximum expression of hermaphrodite characteristics rather than merely adequate functionality, with these flowers representing the upper range of reproductive performance. Fully functional hermaphrodite flowers typically achieve fruit set percentages of 50-65% under optimal environmental conditions (warm, dry bloom period with temperatures 23-30°C, relative humidity 50-70%, calm to light air movement), representing near-maximum fertilization success rates given that even under perfect conditions some flowers fail to set fruit due to developmental abnormalities, resource limitations, or stochastic failures in pollen-stigma interaction. The superior fruit set of fully functional hermaphrodite flowers results from multiple complementary advantages= abundant pollen production ensures saturation of stigmatic surfaces with viable pollen, extended stamen length ensures efficient pollen delivery to stigma through gravity even in still air, high pollen viability ensures adequate functional pollen despite environmental losses, and optimal pistil function ensures high fertilization success of delivered pollen. Fully functional hermaphrodite flowers characterize cultivars and individual vines exhibiting vigorous reproductive development, typically associated with moderate crop loads (avoiding over-cropping that diverts resources from flower development), favorable environmental conditions during previous-season flower primordium differentiation (adequate water and nutrients during June-July bud development), and optimal bloom-period weather. The fully functional condition can exhibit some phenotypic plasticity, with individual cultivars or vines showing year-to-year variation in degree of functionality depending on pre-bloom and bloom-period environmental conditions, nutritional status, and crop load, though some cultivars consistently exhibit fully functional hermaphrodite expression across diverse conditions while others show more variable expression.
Fully_Functional_Hermaphrodite_Flower represents the upper range of hermaphrodite functionality and is distinguished from Balanced_Hermaphrodite_Flower by quantitative measures of organ development and function (greater stamen extension, higher pollen production, higher fruit set) rather than by fundamental morphological differences. The boundary between 'fully functional' and 'balanced' is necessarily somewhat arbitrary but reflects the distinction between maximum and adequate hermaphrodite performance. Fully functional expression is desirable for commercial production but not always achievable or necessary, as balanced hermaphrodite flowers still provide adequate fruit set for economic yields. Cultural practices influencing fully functional expression include crop load management (limiting yields to 8-12 clusters per shoot to optimize flower development), canopy management (ensuring adequate light exposure to flower primordia during previous-season differentiation), and nutritional management (avoiding nitrogen excess that promotes vegetative over reproductive vigor). Understanding factors promoting fully functional versus merely adequate hermaphrodite expression could inform precision viticulture strategies for optimizing reproductive performance.
highly fertile hermaphrodite flower
maximally functional bisexual flower
optimal funktionelle Zwitterblüte
optimal hermaphroditisch
voll funktionelle Zwitterblüte
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 102-108 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 96-103 Lebon, G. et al. (2008). Sugars and flowering in the grapevine. Journal of Experimental Botany 59=2565-2578 Vasconcelos, M.C., Castagnoli, S. (2000). Leaf canopy structure and vine performance. American Journal of Enology and Viticulture 51=390-396.
Fully Functional Hermaphrodite Flower
Merlot grafted on Teleki 5C rootstock at 24 years post-grafting showing fully integrated mature grafted rootstock trunk 8.6 cm diameter, trunk extending 27 cm from root crown to graft union positioned 21 cm above soil, graft union displaying decades-old stable 1.1 cm diameter bulge completely invisible beneath thick weathered bark, trunk bark 4.2 mm thickness showing pronounced Teleki 5C longitudinal fissures with dark grey-brown heavily weathered coloration, bark showing some lichen growth and minor weathering after decades of exposure, trunk fully self-supporting, diameter growth 0.3 cm current season, 24 distinct annual rings in cross-section, no significant disease or damage, supporting mature canopy 3.6 m total shoot length and 7.4 kg fruit load, vine maintaining consistent production after decades
Fully integrated mature grafted rootstock trunk is mature grafted rootstock trunk in mid to late mature phase typically years 18-30 post-grafting, characterized by decades of stable graft union integration with complete callus incorporation and vascular continuity, extensively developed bark showing maximum characteristic rootstock texture, diameter typically 7-10 cm, stable annual growth of 0.3-0.4 cm, complete long-term structural stability, and representing late optimal productive phase where trunk has provided reliable foundation for decades and continues to support high-quality production though approaching eventual transition toward old trunk status. Fully integrated mature grafted rootstock trunk age and maturity includes= spanning years 18-30 post-grafting typically (mid to late mature trunk period), representing long-term mature status (trunk having maintained mature characteristics for 10-22 years), showing late prime productive phase (continuing optimal production while approaching eventual transition to old trunk), displaying fully aged mature characteristics (trunk exhibiting mature features for many years with first hints of aging beginning to appear), and marking sustained maturity (trunk demonstrating long-term stability and consistent performance over decades). Fully integrated mature grafted rootstock trunk decades of integration includes= showing completely consolidated decades-old graft union (callus tissue formed 18-30 years prior fully incorporated into trunk structure), displaying perfect long-term vascular continuity (xylem and phloem connections across graft union functionally flawless after decades of integration), exhibiting stable time-tested graft union morphology (bulge morphology unchanged for decade+ with no ongoing structural changes), showing extensively obscured graft line (very thick mature bark and complete tissue integration making original grafting point essentially invisible), and demonstrating proven mechanical reliability (graft union having withstood decades of stress confirming long-term structural integrity). Fully integrated mature grafted rootstock trunk physical characteristics includes= exhibiting larger mature diameter (7-10 cm typical range for fully integrated phase reflecting decades of annual growth), showing extensively developed bark (periderm 3-5 mm thickness with maximum expression of rootstock cultivar texture after decades of development), displaying deeply fissured bark patterns (longitudinal furrows reaching maximum depth characteristic of rootstock variety after years of cork accumulation), revealing heavily weathered coloration (dark grey-brown bark colors showing decades of environmental exposure and weathering), and showing substantial robust appearance (trunk appearing very solid and well-established reflecting decades of growth and stability). Fully integrated mature grafted rootstock trunk growth stabilization includes= showing consistent but moderating annual increase (0.3-0.4 cm diameter growth in fully integrated phase slightly reduced from younger mature trunk), displaying very regular predictable patterns (decades of consistent growth creating highly predictable annual increments), exhibiting continued cambial activity (vascular cambium still productive though perhaps slightly less vigorous than younger mature trunk), forming 18-30 distinct annual rings (cross-section showing decades of growth rings with clear regular patterns), and maintaining stable wood properties (wood characteristics consistent after decades though perhaps showing slight increase in density). Fully integrated mature grafted rootstock trunk decades of vascular function includes= providing proven water transport capacity (xylem area adequate for 60-100+ liters daily after decades of supporting productive vine), enabling long-term nutrient distribution (phloem and xylem systems having distributed resources effectively for decades), showing sustained transport efficiency (vascular architecture maintaining efficient transport despite decades of use), supporting consistent decades of crop production (trunk having provided adequate capacity for annual fruit loads of 5-12+ kg over many years), and maintaining substantial long-term reserves (extensive ray parenchyma having accumulated and mobilized carbohydrates over decades of annual cycles). Fully integrated mature grafted rootstock trunk decades of production includes= supporting consistent fruit quality over many years (stable trunk function enabling high-quality grape production year after year for decades), enabling reliable sustained yields (adequate trunk capacity for commercial crop loads annually over long productive life), showing stable long-term vigor balance (trunk size having proven well matched to training system and production goals over decades), maintaining productive canopy year after year (trunk supporting healthy leaf area and photosynthesis over many growing seasons), demonstrating proven vine balance (trunk having supported equilibrium between growth and production for many years), and providing long-term foundation for quality (robust trunk having enabled superior production over decades validating rootstock-scion choice). Fully integrated mature grafted rootstock trunk weathering and exposure includes= showing extensively weathered bark surface (outer bark displaying decades of sun, rain, wind, and temperature exposure), displaying accumulated environmental impacts (bark potentially showing lichen growth, moss accumulation, or minor weathering damage after decades), exhibiting deep fissure weathering (longitudinal furrows showing erosion and rounding from years of environmental exposure), revealing darkened coloration (bark colors darkened through decades of oxidation and weathering from initial lighter mature colors), and potentially showing minor damage accumulation (small scars, cracks, or abrasions accumulated over decades though overall trunk remaining healthy). Fully integrated mature grafted rootstock trunk long-term management history includes= reflecting decades of vineyard practices (trunk characteristics showing effects of long-term irrigation, nutrition, and pest management), showing accumulated minor injuries (equipment scars, pruning wounds, or other minor damage from decades of vineyard operations gradually accumulating), displaying disease exposure history (trunk potentially showing old canker scars, healed wounds, or other evidence of past disease events successfully managed), revealing training system effects (trunk form and bark wear patterns reflecting decades of trellis support and canopy management), and indicating management quality (overall healthy fully integrated trunk condition demonstrating decades of appropriate vineyard care). Fully integrated mature grafted rootstock trunk diameter characteristics includes= typical range 7-10 cm for years 18-30 (common dimensions after decades of annual diameter increase), vigorous rootstocks showing 9-12 cm (1103P, 110R showing upper range approaching large old trunk dimensions), moderate vigor rootstocks showing 6.5-9 cm (SO4, 3309C typical for fully integrated phase), low vigor rootstocks showing 6-8 cm (Riparia types showing smaller fully integrated trunk dimensions), and continuing slow steady increase (trunk still expanding though at reduced rate compared to younger mature trunk). Fully integrated mature grafted rootstock trunk cultivar characteristics includes= SO4 showing 6.5-9 cm diameter with deeply furrowed weathered grey-brown bark after decades, 3309C displaying 6-8.5 cm diameter with well-developed fissures and grey bark showing weathering, Teleki 5C exhibiting 7-9 cm diameter with pronounced bark texture after decades of development, 1103P showing 9-12 cm diameter with very thick deeply furrowed bark extensively weathered, Riparia Gloire displaying 6-7.5 cm diameter with moderately furrowed bark for Riparia, 110 Richter exhibiting 9-11 cm diameter with extremely thick rough bark, and 101-14 showing 6-7.5 cm diameter with thin but well-developed fissured bark. Fully integrated mature grafted rootstock trunk heartwood formation includes= potentially beginning heartwood development (innermost xylem in very center of trunk starting to lose function and convert to heartwood in some fully integrated trunks), showing functional sapwood ring (outer xylem layers remaining functional for water transport), displaying slight sapwood reduction (functional sapwood potentially narrowing slightly as trunk ages and central wood converts to heartwood), exhibiting beginning of age-related xylem changes (oldest annual rings beginning to show heartwood conversion in preparation for old trunk status), and maintaining adequate transport capacity (despite any initial heartwood formation, adequate functional sapwood remaining for continued production). Fully integrated mature grafted rootstock trunk as vineyard foundation includes= representing decades of reliable service (fully integrated trunk having supported productive vine through many growing seasons and vintages), demonstrating long-term rootstock-scion compatibility (decades of healthy function confirming successful pairing and integration), showing proven disease resistance (trunk having withstood decades of pathogen exposure with maintained health), exhibiting long-term structural stability (trunk having provided reliable support through decades of wind, weight, and stress), and indicating continued productive potential (healthy fully integrated trunk capable of supporting production for additional years though approaching eventual old trunk transition). Fully integrated mature grafted rootstock trunk late mature characteristics includes= beginning subtle transition toward old trunk (earliest signs of age-related changes appearing though trunk still clearly in mature rather than old category), showing first hints of growth slowing (annual diameter increase perhaps slightly reduced compared to early mature phase), displaying decades of bark accumulation (very thick mature bark from years of cork cambium activity), exhibiting extensive weathering (bark surface showing substantial environmental exposure effects over many years), and maintaining overall mature status (despite hints of aging, trunk still exhibiting primarily mature rather than old trunk characteristics). Fully integrated mature grafted rootstock trunk management in late mature phase includes= continuing standard production practices (fully integrated trunk still requiring only normal vineyard maintenance), monitoring for age-related decline (watching for signs of transition from mature to old trunk status including reduced growth or declining vigor), assessing trunk structural integrity (evaluating bark condition, absence of significant decay, and continued good health), maintaining disease management (inspecting for trunk diseases particularly in aging trunk potentially becoming more susceptible), and evaluating replacement timing (considering whether vine approaching end of productive life or capable of many more years of quality production). Fully integrated mature grafted rootstock trunk production in late phase includes= generally maintaining production capacity (most fully integrated trunks continuing to support adequate crop loads), potentially showing subtle vigor reduction (some older mature trunks perhaps showing slightly reduced vegetative vigor compared to early mature phase), continuing fruit quality support (trunk typically still providing stable resource supply for high-quality fruit), exhibiting stable consistent performance (year-to-year production generally consistent though perhaps with slightly more variability than younger mature trunk), and approaching transition consideration (some fully integrated trunks nearing point where vineyard operator might consider replacement while others capable of decades more production). Fully integrated mature grafted rootstock trunk in old vineyard context includes= characterizing established old vineyards (fully integrated mature trunks typical of vineyards planted 20-30 years prior), representing valuable mature plantings (older vineyards with fully integrated trunks often valued for consistent quality production), showing vintage vineyard characteristics (fully integrated trunks contributing to old vineyard character and reputation), demonstrating long-term site adaptation (decades of growth showing how rootstock-scion combination suited to specific vineyard site), and marking significant vineyard asset (mature vineyard with healthy fully integrated trunks representing substantial value). Fully integrated mature grafted rootstock trunk significance includes= representing proven long-term success (decades of healthy function demonstrating successful grafting, appropriate rootstock selection, and good management), providing late optimal production phase (fully integrated trunk continuing to support quality fruit production while approaching eventual aging transition), demonstrating rootstock-scion compatibility (decades of integration confirming successful partnership between rootstock and scion genotypes), exemplifying viticulture sustainability (fully integrated trunk showing grapevines can remain productive for decades with appropriate care), and marking transition period (fully integrated trunk in late mature phase representing period between peak prime and eventual old trunk status).
Fully_Integrated_Mature_Grafted_Rootstock_Trunk emphasizes mid to late mature phase (years 18-30) and is distinguished from Well_Established_Mature_Grafted_Rootstock_Trunk (early mature phase) and Old_Grafted_Rootstock_Trunk (declining phase beyond mature) by focus on long-term mature status approaching eventual aging transition. Critical for understanding decades of proven integration, late optimal production, extensive weathering, subtle aging onset, and the late mature phase representing proven long-term success before eventual old trunk transition.
Decades-integrated grafted rootstock trunk
Jahrzehnte-integrierter Pfropfstamm
Langfristig etablierter Wurzelstockstamm
Late mature grafted rootstock trunk
Long-established grafted rootstock trunk
Vollständig integrierter reifer verpfropfter Unterlagenstamm
Fully Integrated Mature Grafted Rootstock Trunk
A fully lignified one-year cane on Vitis vinifera cv. Riesling in a cool-climate vineyard, examined during December pruning showing uniform brown color from basal node to apical tip, no green coloration visible even at nodes and bud positions, consistent 8-9mm diameter throughout length with smooth bark texture, firm woody resistance to bending across entire 120cm length, cross-sectional examination at base, middle, and tip positions all revealing well-developed periderm layers, continuous vascular cambium, dense secondary xylem, and minimal pith, indicating complete physiological maturity suitable for retention as 8-node fruiting cane with confidence in winter survival and spring budbreak uniformity.
A lignified one-year cane exhibiting complete secondary cell wall development throughout its entire length from base to tip, characterized by uniform brown woody appearance with no trace of green tissue, consistent firmness and structural rigidity across all internodal regions, well-developed periderm providing protective bark layer, and maximum cold hardiness potential for the cultivar and growing season conditions. Fully lignified one-year canes represent optimal maturation status for fruiting wood, indicating successful completion of the annual growth cycle with adequate growing degree day accumulation, appropriate carbohydrate production and storage, balanced vine crop load allowing complete wood maturation, and favorable environmental conditions during late season ripening period, making such canes preferred selections during winter pruning for maximizing yield potential and fruit quality while ensuring winter survival.
Fully lignified one-year canes are the gold standard for fruiting wood selection, providing optimal balance of structural support, cold hardiness, vascular function, and bud development for maximum productivity. Achievement of full lignification depends on adequate heat accumulation during growing season (typically 1400-1800 growing degree days base 10°C for most Vitis vinifera cultivars), appropriate crop load allowing carbohydrate availability for wood maturation (leaf area-to-fruit ratios above 10-12 cm²/g), and favorable late-season conditions without premature cold snaps forcing early growth cessation. In marginal climates or short growing seasons, achieving full lignification can be challenging, requiring management strategies including early crop thinning, canopy management for light penetration, and cultivar selection favoring early maturation characteristics.
completely lignified cane
komplett ausgereifte Rute
totally mature cane
vollständig verholzte Rute
well-ripened cane
Based on Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 245-267 Fennell (2004) "Freezing tolerance and injury in grapevines" Journal of Crop Improvement 10=201-235 Howell (2001) "Sustainable grape productivity and the growth-yield relationship= A review" American Journal of Enology and Viticulture 52=165-174.
Fully Lignified One Year Cane
Pinot Noir hermaphrodite flowers at days 2-3 post-calyptra shedding (BBCH 63-65, active bloom) exhibit fully receptive stigmas with abundant wet exudate creating glistening appearance, exudate sugar concentration 12-14%, supporting 82-88% germination of self-pollen applied in controlled pollinations, with the fully receptive state persisting for 3-4 days before gradually transitioning to declining receptivity at days 4-5 post-shedding, and with natural self-pollination during the fully receptive period achieving 48-56% fruit set. Chardonnay stigma exudate collected from fully receptive stigmas (days 1-3 post-calyptra shedding) shows optimal biochemical composition= sucrose 7.2-8.8%, glucose 2.1-3.2%, fructose 0.8-1.4% (total sugars 10.1-13.4%), protein 1.1-1.4 mg/ml, invertase activity 2.8-4.2 units/ml, pH 6.1-6.4, providing optimal environment for pollen hydration and germination with 78-86% of applied pollen grains germinating within 3-5 hours. Gewürztraminer functionally female flowers maintain fully receptive stigmas for extended duration of 4-5 days post-calyptra shedding, with exudate production remaining robust (wet glistening stigmas, 11-14% sugar concentration) and pollen germination support high (75-84% of applied hermaphrodite cross-pollen germinating) throughout this extended period, providing prolonged window for capturing asynchronous hermaphrodite pollinizer pollen and achieving 30-38% fruit set through successful cross-pollination. Cabernet Franc controlled pollination experiments applying pollen to stigmas at day 2 (fully receptive) versus day 5 (declining receptive) versus day 7 (post-receptive dry) achieve 68%, 42%, and 6% fruit set respectively, demonstrating that fully receptive stigma state provides optimal conditions for fertilization success with fruit set declining substantially as stigmas transition beyond full receptivity even though some residual receptivity persists.
A fully receptive stigma is a receptive stigma exhibiting complete functional maturity with robust wet exudate production and high pollen germination support efficiency, representing stigmas at or near peak receptivity achieving maximum pollen germination rates typical for the flower type, characteristic of the optimal receptivity period when stigma biochemical and cellular conditions are most favorable for successful pollination. Fully receptive stigmas are characterized by= (1) abundant wet exudate visibly coating papillae creating glistening appearance, with exudate volume and distribution optimal for pollen grain capture and hydration (2) exudate biochemical composition at optimal levels with sugar concentration 10-15%, protein concentration 0.9-1.5 mg/ml, appropriate pH 5.8-6.6, and full complement of enzymes and signaling molecules active (3) papillae in fully turgid state with maximum surface area exposed for pollen capture (4) pollen germination support at high efficiency typical for the flower type= 75-90% germination for hermaphrodite stigmas, 70-88% for functionally female/pistillate stigmas, with germination occurring rapidly (within 2-6 hours of pollen landing) (5) pollen tube initiation and guidance functioning optimally with tubes readily entering transmitting tract. Fully receptive stigmas represent the typical receptivity state during the main receptivity window= in hermaphrodite flowers, fully receptive state persists approximately 2-4 days post-calyptra shedding (days 1-4 of typical 5-6 day total receptivity window) in functionally female and pistillate flowers, fully receptive state persists 3-5 days (days 1-5 of typical 6-8 day window) representing the majority of the receptivity period when conditions are most favorable for pollination. Fully receptive stigmas develop through complete maturation of secretory apparatus and activation of exudate production systems, achieving steady-state exudate secretion that maintains wet stigma surface and optimal biochemical environment despite some exudate loss through evaporation or pollen uptake. Fully receptive stigmas show resilience to moderate environmental variation= moderate temperature fluctuations (15-30°C), moderate humidity variation (45-85%), and typical diurnal cycles do not substantially impair fully receptive stigma function, with pollen germination remaining at 70-90% efficiency across these ranges more extreme conditions (heat >32°C, excessive humidity >90%, severe water stress) can reduce fully receptive stigma performance creating transition toward marginal receptivity. Fully receptive stigmas represent the state targeted for controlled crossing operations in breeding programs, with anthers typically collected and pollen applied when stigmas are 1-3 days post-calyptra shedding to maximize fertilization success, and represent the state when natural pollination (self-pollination in hermaphrodites, cross-pollination in functionally female/pistillate types) is most likely to succeed, determining the critical bloom period when environmental conditions most directly affect fruit set through effects on stigma receptivity maintenance and pollen availability/viability.
Fully_Receptive_Stigma represents optimal receptivity state and is distinguished from Maximally_Receptive_Stigma (which emphasizes absolute peak rather than sustained optimal state) though these categories overlap substantially, with fully receptive encompassing the main high-quality receptivity period while maximally receptive emphasizes the very peak. Fully receptive stigmas represent the normal target state for both natural pollination and controlled crossing operations, providing high-efficiency pollen germination support and fertilization success. Understanding fully receptive stigma characteristics and timing is essential for optimizing pollination success, for timing controlled crosses, and for predicting fruit set potential based on bloom conditions during the fully receptive period.
maximal empfangsfähige Narbe
optimal empfangsbereites Stigma
optimally receptive surface
peak-receptive stigma
voll rezeptive Narbe
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 104-108 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 96-101.
Fully Receptive Stigma
Merlot fully ripe berries at optimal harvest timing (week 17-18, BBCH 89) achieve 24-25.5% sugars, 5.8-6.8 g/L titratable acidity, pH 3.7-3.85, 2.8-3.5 mg/g skin anthocyanins, creating optimal composition for premium Merlot wine with balanced structure, deep color, characteristic plum and black cherry fruit flavors, soft velvety tannins from ripe seeds and skins, and pH supporting microbial stability and aging potential. Riesling fully ripe berries for dry wine production achieve 21-23% sugars, 7.5-9 g/L acidity, pH 3.1-3.3, creating the high-acid balanced style characteristic of quality dry Riesling with crisp acidity supporting aging and food pairing, moderate alcohol potential, and expression of characteristic peach, citrus, and mineral notes. Syrah fully ripe berries show convergence of multiple maturity indicators= analytical maturity (25% sugars, 6.2 g/L acidity, pH 3.75), sensory maturity (tasting shows ripe blackberry and black pepper notes, no green flavors, appropriate tannin levels), phenolic maturity (3.8 mg/g anthocyanins providing deep purple-black color, tannins showing soft ripe character rather than green astringency), and seed maturity (seeds brown, crunchy, showing nutty flavor rather than green bitter notes), with all indicators aligning at week 18 defining optimal harvest window. Cabernet Sauvignon fully ripe berry assessment across three vineyard sites shows site variation in timing= cool site achieving full ripeness at week 19 (23.5% sugars, 6.8 g/L acidity), moderate site at week 18 (24.5% sugars, 6.2 g/L acidity), warm site at week 17 (25% sugars, 5.8 g/L acidity), demonstrating that full ripeness timing varies with mesoclimate while compositional targets remain similar, requiring site-specific harvest timing to achieve full ripeness. Pinot Noir fully ripe berries showing narrow optimal window= week 17 produces near-ripe berries (23% sugars, 7.5 g/L acidity, 2.2 mg/g anthocyanins, still developing flavor), week 17.5-18 achieves full ripeness (24-24.5% sugars, 6.5-7 g/L acidity, 2.8-3.2 mg/g anthocyanins, optimal red fruit character), week 19 shows beginning overripeness (26% sugars, 5.5 g/L acidity, declining anthocyanins), demonstrating 7-10 day optimal harvest window requiring precise timing for this sensitive cultivar.
A fully ripe berry is a mature berry that has achieved complete physiological ripeness with optimal or near-optimal compositional balance for the intended use, characterized by target sugar levels, appropriate acidity, full color development, mature flavor profile, and optimal harvest timing, representing berries at peak quality before overripening begins. Fully ripe berries are characterized by= (1) sugars at target levels for intended use, typically 22-26% for premium red wine, 20-24% for white wine, 18-22% for table grapes, representing optimal fermentable sugar content or eating quality (2) titratable acidity at balanced levels typically 5-8 g/L for wine grapes providing structure and freshness without excessive tartness, 4-7 g/L for table grapes (3) pH in optimal range typically 3.4-3.9 for red wines, 3.2-3.6 for white wines, reflecting appropriate acid buffering capacity (4) full color development with anthocyanins at cultivar-typical mature levels (2-4 mg/g skin in most red cultivars, 3-6+ mg/g in deeply colored cultivars like Syrah or Tannat), creating characteristic varietal color intensity (5) flavor and aroma compounds at optimal levels creating mature varietal character with primary fruit aromas well developed, herbaceous notes diminished, and optimal balance of fruit, floral, and spice notes depending on cultivar (6) tannins (in red cultivars) at optimal ripeness with reduced astringency through polymerization and galloylation changes, appropriate extractability for wine style (7) seeds fully mature showing brown coloration and characteristic flavor (nutty, toasted notes rather than green, bitter notes of unripe seeds). Fully ripe berries represent the optimal harvest window= compositional maturity with sugar=acid ratio balanced for quality phenolic maturity (red cultivars) with color, tannin composition, and extractability optimal flavor maturity with characteristic varietal profile fully expressed physiological maturity with seeds mature and berry metabolism transitioning from ripening to senescence. Fully ripe berries show multiple maturity indicators aligning= analytical maturity (measured composition meeting targets) sensory maturity (tasting assessment showing optimal flavor development) phenolic maturity (skin and seed tannins at optimal ripeness) physiological maturity (seeds mature, berry detachment force declining). Fully ripe berries for different wine styles show varying target compositions= early-harvest fresh styles may achieve full ripeness at 20-22% sugars with higher acidity 7-9 g/L emphasizing freshness standard-harvest balanced wines achieve full ripeness at 23-25% sugars, 5.5-7 g/L acidity concentrated styles may target 25-27% sugars with moderate acidity 5-6 g/L. Fully ripe berries require precise timing= harvest too early produces under-ripe fruit lacking flavor development and phenolic maturity harvest too late produces overripe fruit with excessive sugar, insufficient acidity, and potential flavor degradation optimal harvest window may be brief (3-10 days) requiring careful monitoring and timely harvest execution. Environmental conditions during final maturation affect achievement of full ripeness= temperature influences sugar accumulation rate and flavor development with moderate temperatures 20-28°C optimal water availability affects composition concentration versus dilution sunlight exposure affects anthocyanin development and flavor compound synthesis.
Fully_Ripe_Berry represents optimal harvest-ready state and is distinguished from Overripe_Berry (past optimal timing) by optimal versus declining quality. Fully ripe berries show convergence of multiple maturity indicators defining the harvest decision point. Sub-classification into Optimally_Ripe and Peak_Ripeness reflects subtle distinctions within the fully ripe state. Understanding fully ripe berry characteristics is essential for harvest timing decisions, for quality optimization, and for achieving target wine or table grape quality through harvesting at appropriate maturity.
Vollreife Ernte-Traube
harvest-ready berry
optimal reife Traube
optimally ripe berry
vollreife Beere
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 230-250 Jackson, D.I., Lombard, P.B. (1993). Environmental and management practices affecting grape composition. American Journal of Enology and Viticulture 44=409-430.
Fully Ripe Berry
Merlot on 3309C showing functional active feeder roots in May, minirhizotron observation revealing feeder roots 0.8-1.6 mm with white appearance indicating active status, FDA staining showing 68% viable, isotope pulse-chase (15N-nitrate applied to soil, detected in leaves 2 hours later) demonstrating functional feeder roots actively absorbing and transporting nutrients to shoot, pressure probe measurements showing turgor 0.52 MPa, functional active feeder root population estimated 720 meters per vine providing current absorption capacity supporting active vegetative growth
Functional active feeder root is active feeder root emphasizing operational capacity and demonstrated physiological performance, characterized by measurable uptake of water and nutrients confirming functional status, intact cellular organization maintaining all physiological systems, adequate turgor and membrane integrity enabling normal function, integration into vine physiology with absorbed resources transported to shoot, and representation of feeder roots currently performing absorption supporting vine resource needs, with functional active feeder roots showing complete operational characteristics including functional root hairs extending absorption zone, active arbuscular mycorrhizal associations with developing arbuscules and vesicles, close soil particle contact maximizing exchange interface, demonstrated ion fluxes and water uptake measurable through physiological or isotopic methods, and responsive capacity adjusting absorption to soil conditions and plant demands, and demonstrating that functional status represents operational state where all cellular and physiological systems working normally enabling feeder root to fulfill absorption role contributing measurably to current vine water and nutrient acquisition. Functional active feeder root operational criteria includes= showing demonstrated absorption (measurable water and nutrient uptake confirming functional rather than merely viable status), exhibiting intact physiology (all cellular systems including respiration, transport, biosynthesis operating normally), demonstrating structural integrity (tissues and membranes intact maintaining proper cellular organization and compartmentation), revealing adequate resources (feeder root receiving sufficient photosynthate supporting metabolism and function), and displaying integration (absorbed resources transported from feeder root to vine vascular system reaching shoot). Functional active feeder root performance metrics includes= showing uptake rates (quantified absorption of water and nutrients per unit feeder root length or surface area), exhibiting specific activity (uptake per unit feeder root biomass or surface area indicating efficiency), demonstrating responsive capacity (functional feeder roots adjusting uptake in response to changing soil or plant conditions), revealing diurnal patterns (uptake potentially varying daily correlating with transpiration and metabolism), and displaying population effects (total functional feeder root absorption determining vine resource status). Functional active feeder root in resource acquisition includes= providing current supply (functional feeder roots supplying vine water and nutrients meeting immediate demands), enabling growth support (adequate functional feeder root activity supporting shoot elongation, leaf expansion, fruit development), demonstrating stress buffering (robust functional feeder root population buffering against temporary resource limitations), revealing demand responsiveness (functional feeder root activity responding to increased demands during rapid growth), and displaying seasonal importance (functional feeder root population particularly critical during spring and summer active growth). Functional active feeder root anatomical basis includes= showing primary structure maintenance (functional feeder roots retaining primary anatomical organization facilitating absorption), exhibiting thin cortex (cortex thickness minimized reducing radial transport distance from epidermis to stele), demonstrating functional endodermis (endodermis with casparian strips regulating ion movement while maintaining adequate permeability), revealing appropriate vasculature (stele with sufficient xylem and phloem elements for transport though not extensive secondary growth), and displaying root hair production (abundant functional root hairs on youngest functional feeder root portions). Functional active feeder root mycorrhizal function includes= showing active colonization (functional arbuscular mycorrhizae with developing arbuscules indicating active symbiosis), exhibiting nutrient exchange (fungal-plant nutrient transfer occurring through arbuscules in functional association), demonstrating external hyphae (functional mycorrhizal networks extending from feeder roots exploring soil), revealing phosphorus enhancement (functional mycorrhizae substantially increasing feeder root phosphorus acquisition), and displaying carbon cost (vine allocating 5-20% photosynthate to mycorrhizal fungi in exchange for enhanced uptake). Functional active feeder root environmental responsiveness includes= responding to moisture (functional feeder root activity depending on adequate soil moisture declining under drought), affected by temperature (function optimal 15-25°C declining at temperature extremes), influenced by nutrients (functional feeder root uptake showing kinetic response to soil nutrient concentrations), showing oxygen requirement (function requiring adequate soil aeration declining under waterlogging), and demonstrating pH sensitivity (functional feeder root activity affected by soil pH influencing nutrient availability and transport). Functional active feeder root assessment includes= requiring uptake measurement (isotope tracers or direct measurements confirming functional absorption capacity), showing viability staining (vital dyes indicating membrane integrity necessary for function), exhibiting microscopic examination (anatomy and cytology assessment confirming structural basis for function), demonstrating physiological testing (ion flux measurements or other tests quantifying functional capacity), and revealing integration confirmation (demonstrating absorbed resources reaching vine confirming functional connection). Functional active feeder root in vine performance includes= determining uptake capacity (functional feeder root population limiting vine resource acquisition and growth), enabling yield potential (adequate functional feeder roots supporting fruit production through resource supply), demonstrating quality effects (functional feeder root activity affecting berry composition through nutrient supply), revealing stress tolerance (robust functional feeder root population improving drought and nutrient stress tolerance), and displaying management importance (practices maintaining functional feeder root population critical for vine performance).
Functional_Active_Feeder_Root emphasizes operational performance and demonstrated capacity and is distinguished from merely viable roots by focus on measurable function. Critical for understanding operational status, performance metrics, resource acquisition, anatomical basis, mycorrhizal function, assessment methods, and the demonstrably functional feeder roots currently performing absorption supporting vine resource needs.
Demonstrably functional root
Funktionelle aktive Feinwurzel
Nachweislich funktionsfähige Wurzel
Operational feeder root
Operationale Saugwurzel
Performing absorptive root
Functional Active Feeder Root
Cabernet Sauvignon grafted on 1103 Paulsen showing functional active root hairs on fifth-order fine root, hairs 0.62 mm average length showing complete structural development, microscopic examination confirming intact cytoplasm with abundant mitochondria, FDA viability staining showing bright fluorescence indicating membrane integrity and metabolic activity, pressure probe measurements showing turgor 0.48 MPa, isotope pulse-chase demonstrating hairs actively absorbing and transporting 15N-nitrate to root vascular system within 30 minutes, functional active root hairs 3-5 days post-initiation in peak operational state providing estimated 73% of root uptake
Functional active root hair is active root hair emphasizing operational capacity and physiological performance, characterized by fully developed morphology at final length 0.2-1.2 mm with complete cytoplasmic organization, intact functional membranes maintaining selective permeability and transport capacity, adequate turgor pressure 0.3-0.6 MPa reflecting osmotic function, demonstrated absorption performance through measurable uptake of water and nutrients, and integration into root physiological system with absorbed resources transported through parent cell to root vascular system, with functional active root hair representing optimal operational state typically 2-5 days after hair initiation when growth completed and absorption capacity maximized before senescence begins, showing all cellular systems operating normally including respiration, membrane transport, biosynthesis, and cytoplasmic streaming, and demonstrating that functional state distinguishes operational hairs actively contributing to vine resource acquisition from developing hairs still growing or senescing hairs with declining function. Functional active root hair operational criteria includes= showing structural completion (hair reached final length and ceased tip growth indicating developmental maturity), exhibiting intact cytoplasm (protoplast present and functional with organelles maintaining activity), demonstrating membrane integrity (plasma membrane and organellar membranes intact and functional maintaining compartmentation), revealing adequate turgor (hair turgid reflecting osmotic potential and water status adequate for function), and displaying absorption capacity (demonstrated uptake of water and ions confirming operational status). Functional active root hair physiological integration includes= showing vascular connection (absorbed resources transported from hair through parent cell to root stele and vine vascular system), exhibiting source-sink relationship (hair absorption responding to shoot demand creating coordinate function), demonstrating hormone responsiveness (hair function modulated by auxin, abscisic acid, and other hormonal signals reflecting integration), revealing signal transduction (environmental stimuli affecting hair function transduced through biochemical pathways), and displaying system-level coordination (hair population function integrated into whole-plant resource acquisition strategy). Functional active root hair performance assessment includes= showing uptake measurements (isotope tracers or other methods quantifying hair absorption rates), exhibiting turgor measurements (pressure probe or plasmolysis methods determining hair osmotic function), demonstrating viability staining (fluorescent dyes indicating membrane integrity and metabolic activity), revealing microscopic examination (light or electron microscopy assessing hair structure and cytoplasmic condition), and displaying physiological indicators (respiration, ion fluxes, or other measurements confirming functional state). Functional active root hair peak performance includes= showing maximum absorption rate (functional hairs in peak state showing highest water and nutrient uptake per surface area), exhibiting optimal efficiency (functional hairs showing best ratio of uptake to maintenance cost during peak period), demonstrating full capacity (all cellular systems operating at or near maximum capacity during functional period), revealing brief duration (peak functional state typically lasting 2-4 days representing narrow window in hair lifespan), and displaying subsequent decline (functional capacity gradually declining after peak as hair ages toward senescence). Functional active root hair in root system function includes= providing absorption surface (functional hair population creating majority of current root absorption interface), enabling resource acquisition (functional hairs performing bulk of water and nutrient uptake supporting vine needs), demonstrating dynamic population (functional hairs continuously produced near root tips and senescing basally maintaining relatively stable functional population), revealing turnover dynamics (functional hair lifetime brief requiring continuous production maintaining functional surface), and displaying management sensitivity (practices affecting functional hair production or lifespan directly impacting vine resource status). Functional active root hair environmental effects includes= responding to moisture availability (functional hair performance depending on adequate soil moisture declining under drought), affected by soil strength (functional hair abundance and function reduced in compacted soil limiting hair development), influenced by nutrients (functional hair uptake rates varying with soil nutrient availability showing uptake kinetics), showing temperature effects (functional hair metabolism and transport increasing with temperature within optimal range 15-25°C), and demonstrating pH sensitivity (functional hair performance affected by soil pH influencing membrane transport and nutrient availability). Functional active root hair stress impacts includes= showing drought sensitivity (functional hairs losing turgor and function rapidly under water deficit due to unfavorable surface-to-volume ratio), exhibiting toxicity effects (functional hair performance impaired by aluminum, sodium, or other toxic ions damaging membranes), demonstrating pathogen vulnerability (functional hairs potentially susceptible to root pathogens though rapid turnover may limit disease impact), revealing physical damage (functional hairs easily damaged by soil fauna, cultivation, or other mechanical disturbance), and displaying recovery capacity (root system compensating for functional hair loss through rapid new hair production at growing tips). Functional active root hair contribution quantification includes= showing uptake proportion (functional hairs providing 60-80% of root system water and nutrient uptake based on isotope studies), exhibiting surface area proportion (functional hairs comprising 10-20% of total root surface area though providing disproportionate uptake), demonstrating efficiency advantage (functional hairs showing 10-50 fold higher uptake per unit surface area than root cylinder), revealing cost-effectiveness (functional hairs providing uptake at low carbon cost due to thin walls and single-cell structure), and displaying critical importance (functional hair population essential for adequate vine resource acquisition particularly under stress or limiting conditions).
Functional_Active_Root_Hair emphasizes operational capacity and performance and is distinguished from developing or senescing hairs by focus on fully functional state. Critical for understanding operational criteria, physiological integration, performance assessment, peak capacity, environmental responsiveness, and the fully functional state representing optimal hair contribution to vine resource acquisition.
Fully functional hair
Funktionelles aktives Wurzelhaar
Operational root hair
Operationales Haar
Performing active hair
Voll-funktionsfähiges Absorptionshaar
Functional Active Root Hair
Merlot grafted on 3309 Couderc rootstock showing functional maturation zone on third-order lateral root extending 12-85 mm from root apex, zone exhibiting fully differentiated tissues, abundant mature root hairs 180 per cm creating 12-fold surface area increase, functional endodermis with complete casparian strips regulating uptake, mature lignified xylem vessels conducting water, functional phloem with active sieve elements, cortex cells containing starch reserves, isotope tracing demonstrating functional zone providing 84% of root water uptake and 78% of nitrogen uptake, pericycle showing lateral root initiation, functional maturation zone performing essential absorption, transport, and storage functions
Functional maturation zone is maturation zone emphasizing physiological capacity and performance of mature functions, characterized by fully differentiated tissues actively performing root physiological functions including water and nutrient absorption through epidermis and root hairs, radial transport regulation through endodermis casparian strips, axial water transport through mature lignified xylem vessels, photosynthate transport through functional phloem sieve elements, carbohydrate storage in cortex and pericycle parenchyma, and lateral root initiation capacity in pericycle cells, showing mature tissue organization with all cell types complete and functional, demonstrable physiological activity with measurable uptake, transport, and metabolic rates, and representing root region performing essential vine functions distinguishing functional mature root from immature growing tip or senescent dying root, with functional maturation zone providing majority of root system absorption capacity through extensive root hair surface area, enabling long-distance transport connecting root uptake to shoot demand, demonstrating storage and buffer capacity moderating resource availability, and showing developmental plasticity through lateral root formation enabling root system architectural adjustment. Functional maturation zone absorption capacity includes= showing high uptake rates (functional zone performing 70-90% of root system water and nutrient absorption), exhibiting root hair function (abundant mature root hairs creating 5-20 fold surface area increase optimizing absorption), demonstrating active transport (epidermal and cortical cells with functional ion transporters enabling selective nutrient uptake), revealing mycorrhizal contribution (arbuscular mycorrhizae in cortex enhancing phosphorus and water uptake 3-10 fold), and displaying responsive uptake (absorption rates adjusting to soil resource availability and plant demand). Functional maturation zone transport capacity includes= showing xylem function (mature lignified vessels conducting water and minerals from uptake sites to shoot system), exhibiting phloem function (functional sieve elements transporting photosynthates from shoots to roots), demonstrating bidirectional transport (simultaneous upward xylem flow and downward phloem flow), revealing transport rates (functional zone potentially conducting 10-50+ liters water daily per vine depending on transpiration), and displaying hydraulic connectivity (functional zone vessels integrated into continuous hydraulic pathway from soil to leaves). Functional maturation zone endodermis regulation includes= showing selective permeability (casparian strips forcing radial transport through symplast enabling ion selectivity), exhibiting uptake control (endodermis transport proteins regulating mineral entry into vascular cylinder), demonstrating backflow prevention (casparian barrier preventing mineral leakage from stele to cortex), revealing passage cells (thin-walled endodermis cells in older functional zone maintaining flexibility in transport regulation), and displaying adaptive regulation (endodermis transport activity adjusting to nutrient status and demand). Functional maturation zone storage function includes= showing carbohydrate storage (cortex and pericycle cells accumulating starch particularly during dormancy), exhibiting seasonal cycles (reserves accumulating fall-winter mobilizing spring supporting growth), demonstrating buffer capacity (stored reserves buffering short-term supply-demand imbalances), revealing nutrient storage (functional zone accumulating nitrogen, phosphorus, and other nutrients in organic forms), and displaying remobilization capacity (stored compounds mobilizing when needed supporting metabolism and growth). Functional maturation zone lateral root formation includes= showing pericycle competence (pericycle cells in functional zone capable of dedifferentiating and initiating lateral root primordia), exhibiting lateral emergence (lateral roots forming in functional zone growing through cortex and emerging from parent root), demonstrating architectural plasticity (lateral root production enabling root system branching and adaptation), revealing responsive initiation (lateral root formation responding to environmental signals and resource availability), and displaying continued capacity (functional zone maintaining lateral root initiation capacity throughout zone lifespan). Functional maturation zone metabolic activity includes= showing active respiration (functional zone cells consuming oxygen supporting uptake, transport, and biosynthesis), exhibiting ATP production (mitochondria generating energy for active transport and metabolism), demonstrating biosynthesis (producing cell wall materials, membrane lipids, proteins, and other compounds), revealing defensive metabolism (synthesizing antimicrobial compounds and mounting responses to pathogen attack), and displaying stress responses (adjusting metabolism in response to drought, temperature, or other stresses). Functional maturation zone lifespan includes= showing extended function (functional maturation zone potentially remaining active weeks to months depending on root order and conditions), exhibiting order-dependent duration (primary roots showing functional zones persisting years while fine roots showing weeks to months), demonstrating eventual senescence (functional zone eventually declining as older root tissues age and function declines), revealing turnover (fine root functional zones showing rapid turnover while structural root zones persisting), and displaying replacement (senescing functional zones continuously replaced by new maturation zone formation at elongating root tips). Functional maturation zone in root system performance includes= determining uptake capacity (total functional maturation zone surface area and activity limiting vine water and nutrient acquisition), enabling hydraulic function (functional zone xylem conductivity contributing to total root system hydraulic conductance), demonstrating storage reservoir (functional zone carbohydrate reserves supporting vine through dormancy and stress), showing architectural control (functional zone lateral root production determining root system branching patterns), and revealing management significance (practices affecting functional maturation zone directly impacting vine resource status and performance).
Functional_Maturation_Zone emphasizes physiological performance and active functions and is distinguished from differentiation emphasis by focus on functional capacity. Critical for understanding absorption performance, transport capacity, storage function, lateral root formation, metabolic activity, and the physiologically active mature root region performing essential root system functions enabling vine resource acquisition and physiological integration.
Functional root area
Funktionelle Reifungszone
Leistungsfähige Wurzelregion
Performing maturation region
Physiologically active zone
Physiologisch aktiver Bereich
Functional Maturation Zone
A functional tendril at node 9 on a vigorous shoot of Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, California, having achieved 22 cm extension length over three weeks of development, exhibiting active circumnutation movements sweeping through approximately 180-degree arcs over 2-3 hour periods seeking contact opportunities, demonstrating rapid coiling response within 8 minutes when brought into contact with an experimental support rod, achieving complete mechanical attachment with both contact and free coils within 18 hours post-contact, and supporting approximately 400 grams of shoot and leaf weight when tested mechanically at BBCH stage 75 (berries pea-sized), representing a fully functional climbing organ contributing to vertical shoot positioning and canopy architecture.
A tendril that has achieved full physiological and structural maturation sufficient to execute its climbing function in Vitis vinifera, characterized by completed elongation growth reaching species-typical and cultivar-specific length ranges of 10-30 cm, development of mechanosensory capabilities with responsive epidermal cells capable of detecting contact and initiating coiling responses, adequate structural integrity with appropriate lignification and cellulose deposition to withstand mechanical loading from vine weight and wind forces, and active metabolic status maintaining the capacity for thigmotropic movements and coiling reactions. Functional tendrils represent the subset of all tendrils that contribute meaningfully to vine anchorage and climbing behavior, excluding aborted tendrils that cease development prematurely, senescing tendrils that have lost physiological activity, damaged tendrils with compromised structural integrity, and immature tendrils still undergoing developmental processes. The transition to functional status typically occurs 2-3 weeks after tendril emergence from the apical meristem, requiring completion of cell expansion, differentiation of mechanosensory apparatus, and achievement of minimal structural strength thresholds. Functional tendrils exhibit diverse behavioral states including active searching movements involving slow circumnutation to increase probability of contact, rapid coiling responses upon successful contact, and maintained tension generation in established attachments that support vine architecture against gravitational and wind loading.
Functional tendrils are distinguished from non-functional tendrils through multiple criteria including structural competence, physiological responsiveness, and actual or potential contribution to vine support. Assessment of functionality requires evaluation of elongation completion, mechanosensory capacity (tested through contact response), structural strength (evaluated through loading tests or visual inspection for adequate lignification), and metabolic activity (indicated by tissue color, turgor, and growth capacity). Tendrils may lose functional status through senescence, desiccation, pathogen infection, or mechanical damage, transitioning from functional to non-functional categories. Environmental conditions strongly influence functional tendril development, with water stress, nutrient deficiency, or extreme temperatures potentially preventing achievement of full functionality despite apparently normal morphology.
active tendril
aktive Ranke
functional climbing organ
funktionsfähige Ranke
mature tendril
reife Kletterorgan
responsive tendril
Based on Bowling & Vaughn (2009) "Gelatinous fibers are widespread in coiling tendrils and twining vines" American Journal of Botany 96=719-727 Jaffe & Galston (1968) "The physiology of tendrils" Annual Review of Plant Physiology 19=417-434 Darwin (1875) "The Movements and Habits of Climbing Plants" John Murray, London Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 79-94.
Functional Tendril
A functional tertiary reproductive bud at node 6 on Vitis vinifera cv. Riesling under exceptional growing conditions (optimal light, temperature, water, nutrition during previous season bud differentiation), dissection revealing small but unambiguous inflorescence primordium containing 32 clearly identifiable flower initials with visible ovary and stamen primordia, this bud activating following combined frost-hail damage eliminating primaries and secondaries, developing into weak shoot bearing one tiny cluster of 11 berries (34% fruit set) weighing 18g at harvest, representing minimal but non-zero crop contribution that slightly improved economic recovery from catastrophic damage event, though contributing only 1.2% of normal node yield and requiring same multi-year recovery timeline as purely vegetative tertiary positions.
A tertiary reproductive bud containing clearly identifiable inflorescence primordia of sufficient size and organization to represent legitimate reproductive potential, albeit minimal, characterized by unambiguous flower initials numbering 20-40 under microscopic examination, demonstrated capacity to produce small but harvestable clusters when activated, and documented occurrence in a few high-fruitfulness cultivars under optimal bud development conditions. Functional tertiary reproductive buds represent the upper extreme of tertiary reproductive capacity, extraordinarily rare but occasionally documented, providing theoretical possibility of minimal crop contribution even under catastrophic damage scenarios requiring tertiary activation, though practical significance remains negligible given the rarity of functional tertiary reproductive buds (perhaps 0.5-2% of tertiary positions in most fruitful cultivars, zero in most others) and the extreme damage conditions required for their activation.
Functional tertiary reproductive buds, while academically interesting as demonstrations of extreme reproductive capacity in certain cultivars, have minimal practical significance in viticulture due to their extreme rarity and the catastrophic damage scenarios required for activation. Even in cultivars like Riesling and Gewürztraminer that occasionally show functional tertiary reproductive buds, their prevalence is so low (typically under 2% of positions) that they contribute negligibly to damage recovery calculations. The existence of functional tertiary reproductive buds does demonstrate that reproductive pathway suppression in tertiary positions is not absolute but rather represents resource allocation priorities during compound bud development, with some cultivars maintaining minimal reproductive capacity even in tertiary positions while most concentrate reproductive investment entirely in primary and to lesser extent secondary positions.
developed tertiary inflorescence bud
entwickelte Tertiärknospe
funktionale reproduktive Tertiärknospe
productive tertiary bud
viable tertiary reproductive bud
Based on Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Srinivasan & Mullins (1981) "Physiology of flowering in the grapevine - a review" American Journal of Enology and Viticulture 32=47-63 Vasconcelos et al. (2009) "The flowering process of Vitis vinifera= a review" American Journal of Enology and Viticulture 60=411-434.
Functional Tertiary Reproductive Bud
Thompson Seedless (Sultana) exhibits strongly functionally female flowers with stamens reflex 1-2 mm below pistil level, producing minimal pollen (<3,000 grains per flower) with <15% viability, requiring co-planting with hermaphrodite pollinizers such as Calmeria or Perlette at 1=10 ratios to achieve 25-35% fruit set, with inadequate pollination resulting in <10% fruit set and uneconomical yields. Gewürztraminer in Alsace shows variable sex expression, with some clones and cool bloom conditions inducing functionally female expression with strongly reflex stamens and reduced pollen viability, requiring cross-pollination from co-planted Riesling or other hermaphrodite cultivars to achieve acceptable 30-40% fruit set, while warm bloom conditions may induce more hermaphrodite-like expression with partially erect stamens and improved self-fertility. Orange Muscat exhibits constitutive functionally female flowers with reflex stamens and sterile pollen, dependent on pollinizer co-planting throughout its cultivation range.
A functionally female flower is a perfect flower possessing both stamens and pistil morphologically but with male function impaired through stamen reflexing (downward curvature positioning anthers below pistil) or production of non-viable or severely reduced-viability pollen, while female organs (pistil with receptive stigma and viable ovules) remain fully functional, requiring cross-pollination from hermaphrodite flowers for successful fruit set and characterizing certain cultivars that depend on co-planted pollinizers for fruit production. Functionally female flowers exhibit the morphological components of both sexes but functional unisexuality due to male sterility, with the critical diagnostic feature being strongly reflex stamens that curve sharply downward, positioning anthers 0.5-2 mm below the stigma rather than at or above stigma level as in hermaphrodite flowers, effectively preventing self-pollination through spatial separation of pollen source and pollen destination. In some functionally female types, stamens may show less pronounced reflexing but produce pollen with severely reduced viability (<10-20% viable grains) or drastically reduced pollen quantity (<5,000 grains per flower), achieving functional male sterility through gametic rather than purely morphological mechanisms. The pistil in functionally female flowers is fully developed and functional, with normal ovary, style, and stigma morphology, high stigma receptivity to cross-pollination, and viable ovules capable of fertilization by pollen from hermaphrodite sources. Functionally female flowers absolutely require cross-pollination for fruit set, relying on wind-borne or insect-borne pollen from co-planted hermaphrodite cultivars (pollinizers) to achieve fertilization. Fruit set in functionally female flowers is typically lower (20-40%) and more variable than in hermaphrodite flowers (40-60%), depending critically on= (1) proximity and abundance of hermaphrodite pollinizer plants providing adequate pollen supply (2) favorable weather during bloom facilitating pollen dispersal and transfer (moderate wind, low humidity reducing pollen clumping, dry conditions preventing pollen washout) (3) bloom synchrony between functionally female and pollinizer cultivars ensuring temporal overlap in stigma receptivity and pollen availability. The functionally female condition characterizes several commercially important table grape cultivars (many seedless types including Thompson Seedless/Sultana, Flame Seedless, and numerous others) and a few wine cultivars (Gewürztraminer exhibits functionally female expression in some clones and environmental conditions, Picolit, Orange Muscat). The evolutionary and genetic basis of functionally female flowers involves mutations affecting stamen development and pollen formation, with the reflex stamen condition controlled by specific genetic loci and showing Mendelian inheritance in breeding populations. Viticultural management of functionally female cultivars requires= (1) pollinizer co-planting at ratios typically 1=8 to 1=12 (pollinizer=functionally female) ensuring adequate pollen supply (2) selection of pollinizer cultivars with overlapping bloom periods, abundant pollen production, and compatible pollen (3) row orientation and spacing facilitating wind-borne pollen movement (4) avoidance of practices disrupting bloom synchrony (differential irrigation or nutrition between pollinizer and primary cultivar).
Functionally_Female_Flower represents morphological bisexuality with functional unisexuality due to male sterility, and must be distinguished from Hermaphrodite_Flower (both organs functional), Functionally_Male_Flower (female organs non-functional), and true female flowers lacking stamens entirely (rare in Vitis vinifera cultivars but present in wild V. vinifera ssp. sylvestris). The term 'functionally female' specifically denotes the functional state rather than morphology, as stamens are morphologically present but functionally impaired. Functionally female cultivars present significant viticultural challenges through pollinizer dependency, variable fruit set, and breeding complications, but are economically important particularly in seedless table grape production where the functionally female condition is often associated with seedlessness genes. Sub-classification into Reflex_Stamen and Reduced_Pollen types reflects the two primary mechanisms of male sterility in functionally female flowers.
female-sterile flower
funktionell pistillate Blüte
funktionell weibliche Blüte
pistillate functional flower
weiblichblütig
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 104-106 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 98-101 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Antcliff, A.J. (1980). Inheritance of sex in Vitis. Annales de l'Amélioration des Plantes 30=113-122 Battilana, J. et al. (2013). The genetic basis of the conversion of hermaphrodite flowers to male flowers in grapevine. Nature Communications 4=2615.
Functionally Female Flower
Occasional functionally male vines appear in Pinot Noir vineyards as somatic mutations or seedlings from uncontrolled pollination, exhibiting flowers with erect stamens extending 1-1.5 mm beyond the abbreviated pistil (pistil only 60-70% of normal length), producing abundant pollen (20,000-28,000 grains per flower, 70-80% viability) but achieving <2% fruit set due to non-receptive stigma and aborted ovules, with affected vines producing normal vegetative growth but essentially no crop and typically removed upon identification during first bearing season. Wild Vitis vinifera ssp. sylvestris male plants in riparian populations of Mediterranean regions bear exclusively functionally male flowers with vigorous stamens producing copious pollen that fertilizes nearby female plants, but setting <1% fruit themselves due to complete pistil abortion with stigma absent and ovary rudimentary. Certain breeding population seedlings segregate for sex determination genes, producing 10-25% functionally male individuals with hermaphrodite-appearing stamens but sterile pistils, requiring early identification and removal to avoid allocating vineyard space to non-productive genotypes.
A functionally male flower is a perfect flower possessing both stamens and pistil morphologically but with female function impaired through pistil abortion, stigma non-receptivity, or ovule sterility, while male organs (stamens with viable pollen) remain functional, producing abundant pollen that can fertilize hermaphrodite or functionally female flowers but incapable of setting fruit on the plant bearing these flowers, characteristic of staminate (male-functional) vines that serve as pollen sources but not fruit producers. Functionally male flowers exhibit morphological bisexuality but functional unisexuality due to female sterility, with stamens showing normal hermaphrodite morphology (erect position, extending beyond pistil) and producing abundant viable pollen (15,000-30,000+ grains per flower with 60-85% viability), but pistil showing one or more defects preventing fruit set= (1) severely abbreviated pistil length (often only 50-70% of normal hermaphrodite pistil length), positioning stigma well below anther level (2) non-receptive or absent stigma lacking the characteristic wet exudate that facilitates pollen hydration and germination (3) aborted or sterile ovules incapable of fertilization and embryo development even if pollination occurs (4) style dysfunction preventing pollen tube growth to ovules. The functionally male condition results in complete or near-complete failure of fruit set (<1-5%) on plants bearing these flowers, despite normal or enhanced pollen production, creating plants that function as pollen donors but not fruit producers. Functionally male flowers characterize certain individual vines within otherwise hermaphrodite cultivars (somatic mutations or genetic segregation), occur as the male phase in dioecious wild Vitis vinifera ssp. sylvestris populations, and appear in some breeding populations segregating for sex determination genes. The stamens in functionally male flowers are often more vigorous than in hermaphrodite flowers, with greater stamen extension, larger anthers, and enhanced pollen production, potentially representing resource reallocation from non-functional female structures to enhanced male function. The pollen produced by functionally male flowers is fully viable and functional, capable of successfully fertilizing hermaphrodite or functionally female flowers when transferred by wind or insects, making functionally male vines potentially valuable as pollinizer sources despite their inability to produce fruit. However, the functionally male condition is generally undesirable in commercial viticulture, as vines bearing exclusively functionally male flowers produce no harvestable crop and occupy vineyard space unproductively, typically leading to roguing (removal) of such vines when identified. The genetic basis of functionally male expression involves sex determination loci that segregate in breeding populations, with functionally male types arising from specific allelic combinations that suppress female function while maintaining or enhancing male function. The evolutionary significance relates to the ancestral dioecious condition of wild Vitis, with functionally male flowers representing the male phase of sexual reproduction that was largely eliminated during domestication through selection for hermaphrodite self-fertile types, but which occasionally resurfaces through genetic segregation or reversion.
Functionally_Male_Flower represents morphological bisexuality with functional unisexuality due to female sterility, and must be distinguished from Hermaphrodite_Flower (both organs functional), Functionally_Female_Flower (male organs non-functional), and true male flowers completely lacking pistils (characteristic of some wild Vitis species and dioecious V. vinifera ssp. sylvestris). The term 'functionally male' specifically denotes the functional state rather than morphology, as pistils are morphologically present but functionally impaired. Functionally male vines are generally undesirable in commercial viticulture as they produce no crop, though they could theoretically serve as dedicated pollinizer sources for functionally female cultivars if pollen characteristics are favorable. The functionally male condition appears at low frequency in commercial vineyards through somatic mutation or seed propagation, requiring vigilant early identification and removal. Sub-classification into Short_Pistil and Non_Receptive types reflects the two primary mechanisms of female sterility in functionally male flowers.
funktionell männliche Blüte
männlich-fertile Blüte
pollen-producing sterile flower
pollenproduzierend steril
staminate functional flower
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 100-103 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Antcliff, A.J. (1980). Inheritance of sex in Vitis. Annales de l'Amélioration des Plantes 30=113-122 Battilana, J. et al. (2013). The genetic basis of the conversion of hermaphrodite flowers to male flowers in grapevine. Nature Communications 4=2615 Picq, S. et al. (2014). A small XY chromosomal region explains sex determination in wild dioecious V. vinifera and the reversal to hermaphroditism in domesticated grapevines. BMC Plant Biology 14=229.
Functionally Male Flower
3309 Couderc (V. riparia × V. rupestris) demonstrates moderate Armillaria resistance in replant situations in Burgundy, with vine survival rates of 70-85% over 15 years compared to 40-60% on susceptible SO4 in Armillaria-infested sites. Freedom (1613C × Dog Ridge) shows enhanced Phytophthora tolerance in poorly drained clay soils of the Napa Valley floor, maintaining productivity where susceptible rootstocks succumb to crown and root rot. Dog Ridge (V. champinii) combines moderate fungal disease resistance with nematode resistance, providing multi-stress protection in replant vineyards of the San Joaquin Valley.
A stress-resistance rootstock role that inheres in a grafted rootstock by virtue of its genetically determined capacity to resist infection and colonization by soil-borne fungal pathogens affecting grapevine root systems. Fungal disease-resistant rootstocks prevent or limit pathogen establishment through constitutive defense mechanisms (thick-walled cortical cells, high baseline phenolic content, preformed antimicrobial compounds), induced defense responses (phytoalexin synthesis, pathogenesis-related protein accumulation, lignification at infection sites), and anatomical barriers that restrict pathogen spread within root tissues. Resistance is assessed by lesion size, pathogen colonization extent (measured as DNA copy number per gram root tissue), and vine survival rates following controlled inoculation. The most economically important fungal root pathogens in viticulture are Armillaria mellea sensu lato (oak root fungus), Phytophthora spp. (particularly P. cinnamomi and P. megasperma), and various Cylindrocarpon/Ilyonectria species causing black foot disease.
Fungal_Disease_Resistant_Rootstock_Role is subclassed by specific pathogen targets, currently Armillaria_Resistant_Rootstock_Role and Phytophthora_Resistant_Rootstock_Role. Resistance to fungal root pathogens is generally less well characterized genetically than phylloxera or nematode resistance, and field screening remains the primary evaluation method. Fungal resistance often shows quantitative inheritance and strong environmental interaction, making categorical resistance classification more challenging than for insect pests. Black foot disease (Cylindrocarpon/Ilyonectria) resistance is an emerging breeding target not yet represented by a specific subclass.
bodenbürtige Pilzkrankheiten resistente Unterlage
fungal disease resistant rootstock
gegen Wurzelpilze resistente Unterlage
pilzresistente Unterlage
root rot resistant rootstock
soil-borne fungal pathogen resistant rootstock
Aguín O et al. (2008) Armillaria species infesting vineyards in northwestern Spain, European Journal of Plant Pathology 120: 261-272 Agusti-Brisach C, Armengol J (2013) Black-foot disease of grapevine: an update on taxonomy, epidemiology and management strategies, Phytopathologia Mediterranea 52: 245-261 Marais PG (1979) Fungi associated with root rot in vineyards in the Western Cape, Phytophylactica 11: 65-68
Fungal Disease Resistant Rootstock Role
Fungal infection indicator quality assessment during a disease diagnostic survey in Burgundy identified multiple concurrent fungal infections requiring differentiated management responses. Leaf symptoms showing fungal infection indicator quality included the chlorotic oil-spot lesions with white sporulation characteristic of downy mildew (Plasmopara viticola), the powdery white colonies of powdery mildew (Erysiphe necator), and the angular necrotic lesions with black pycnidia of black rot (Guignardia bidwellii), each pathogen producing distinctive fungal infection indicator quality patterns enabling confident field diagnosis. Trunk symptoms showing fungal infection indicator quality included the wedge-shaped necrotic sectors in cross-section characteristic of Eutypa dieback and the tiger-stripe leaf patterns associated with esca complex, both representing chronic wood-colonizing fungi requiring long-term management strategies distinct from the foliar and fruit diseases. The comprehensive fungal infection indicator quality assessment enabled prioritized management response, with immediate fungicide application targeting the actively sporulating powdery and downy mildew, cultural practices to reduce Botrytis risk on the approaching harvest, and trunk disease management deferred to the dormant season pruning period.
Fungal infection indicator quality is a pathogen infection indicator quality that inheres in a grapevine organism and represents the ensemble of symptoms and diagnostic signs specifically associated with fungal pathogen colonization, including the characteristic lesions, sporulating structures, mycelial growth, and tissue alterations produced by the diverse fungal pathogens that constitute the primary biotic disease agents affecting grapevine in most viticultural regions worldwide. This quality captures the plant responses and pathogen manifestations that enable recognition and diagnosis of fungal diseases, recognizing that fungi cause the majority of economically significant grapevine diseases and that effective disease management depends on accurate identification of the specific fungal pathogen responsible for observed symptoms. The ontological significance of fungal infection indicator quality lies in its role enabling differential diagnosis among the numerous fungal pathogens affecting grapevine, with pathogen-specific symptom patterns, host tissue preferences, environmental associations, and microscopic features providing the diagnostic basis for targeted management responses that differ substantially among fungal disease types. The biological basis of fungal infection indicators encompasses the visible consequences of fungal colonization including hyphal penetration and growth within host tissues, the production of cell-wall-degrading enzymes causing tissue maceration, the synthesis of toxins causing cell death and discoloration, and the formation of reproductive structures including conidiophores, sporangiophores, and fruiting bodies that produce the spores enabling pathogen spread. The diversity of fungal pathogens affecting grapevine creates a corresponding diversity of symptom types, from the biotrophic powdery mildews producing superficial colonies without killing host cells through the hemibiotrophic downy mildew initially growing intercellularly before causing tissue death to the necrotrophic Botrytis and black rot fungi that rapidly kill and colonize host tissues. The seasonal timing and environmental associations of fungal disease development create characteristic epidemiological patterns that aid diagnosis, with spring conditions favoring certain pathogens, summer conditions others, and the progression of symptoms through the growing season reflecting pathogen biology and disease cycle characteristics.
Fungal_Infection_Indicator_Quality is_a Pathogen_Infection_Indicator_Quality, serving as parent class for Downy_Mildew_Symptom_Quality, Powdery_Mildew_Symptom_Quality, Botrytis_Symptom_Quality, Black_Rot_Symptom_Quality, Esca_Symptom_Quality, and Eutypa_Dieback_Symptom_Quality which address specific fungal diseases. This class addresses the general features of fungal disease symptoms while child classes capture pathogen-specific diagnostic criteria. The biotrophic-hemibiotrophic-necrotrophic spectrum affects symptom types and management approaches. Fungi cause the majority of economically significant grapevine diseases worldwide.
Pilzinfektions-Indikatoren
Pilzkrankheits-Symptome
Pilzpathogen-Marker
fungal disease markers
fungal disease symptoms
fungal pathogen signs
mycotic infection indicators
mykotische Infektionszeichen
Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests, 2nd Edition. APS Press Pearson RC, Goheen AC (1988) Compendium of Grape Diseases. APS Press Armijo G et al. (2016) Grapevine pathogenic microorganisms: understanding infection strategies and host response scenarios. Frontiers in Plant Science 7:382
Fungal Infection Indicator Quality
The fungal resistance breeding program at the Julius Kühn-Institut (JKI) in Siebeldingen, Germany, systematically evaluates Fungal_Resistance_Disposition in segregating populations derived from crosses between Vitis vinifera and resistant wild species. Seedlings are screened for resistance to Plasmopara viticola using leaf disc assays with standardized sporangial suspensions (50,000 sporangia/mL), scoring sporulation intensity on a 1-9 OIV scale after 7 days incubation. Individuals rated OIV 1-3 (no to weak sporulation) possess strong Fungal_Resistance_Disposition against downy mildew, while those rated OIV 7-9 (strong sporulation covering >50% leaf area) lack effective resistance. Selected resistant genotypes undergo further evaluation for Erysiphe necator resistance in greenhouse trials and Botrytis cinerea resistance in field clusters, as pyramiding multiple Fungal_Resistance_Dispositions is essential for durable protection under diverse environmental conditions and evolving pathogen populations.
A Fungal_Resistance_Disposition is a Pathogen_Resistance_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through defense processes upon encounter with fungal pathogens and oomycetes, the most economically significant group of grapevine disease-causing organisms. This disposition encompasses resistance mechanisms effective against obligate biotrophic pathogens such as Plasmopara viticola (downy mildew) and Erysiphe necator (powdery mildew), necrotrophic pathogens including Botrytis cinerea (grey mold), hemibiotrophic pathogens such as Guignardia bidwellii (black rot) and Elsinoë ampelina (anthracnose), and wood-inhabiting fungi causing trunk diseases including Phaeomoniella chlamydospora and various Botryosphaeriaceae species. The molecular basis of fungal resistance dispositions includes recognition of fungal cell wall components (chitin, β-glucans) by pattern recognition receptors, detection of fungal effector proteins by NLR receptors, and activation of defense responses including chitinase and glucanase production, phytoalexin biosynthesis (particularly stilbenes such as resveratrol and viniferins), lignification, and callose deposition. Fungal resistance dispositions may operate at multiple infection stages including spore germination inhibition, penetration resistance, haustorial encasement, and limitation of mycelial spread. Given that fungal and oomycete diseases account for over 80% of pesticide use in viticulture globally, enhancing fungal resistance dispositions through breeding represents a critical objective for sustainable grape production.
Fungal_Resistance_Disposition is_a Pathogen_Resistance_Disposition. By convention in plant pathology and viticulture, this class includes resistance to oomycetes (e.g., Plasmopara viticola) despite their phylogenetic distinction from true fungi (Kingdom Stramenopila vs. Kingdom Fungi). Child classes are organized by specific pathogen: Downy_Mildew_Resistance_Disposition, Powdery_Mildew_Resistance_Disposition, Botrytis_Resistance_Disposition, Black_Rot_Resistance_Disposition, Anthracnose_Resistance_Disposition, and Trunk_Disease_Resistance_Disposition.
Mykose-Resistenz
Pilzresistenz-Disposition
Pilzwiderstandsfähigkeit
fungal disease resistance
mycotic resistance disposition
oomycete resistance potential
Gadoury DM et al. (2012) Grapevine powdery mildew (Erysiphe necator): a fascinating system for the study of the biology, ecology and epidemiology of an obligate biotroph, Molecular Plant Pathology 13:1-16 Kassemeyer HH (2017) Fungi of grapes, Biology of Microorganisms on Grapes, in Must and in Wine, Springer Töpfer R, Hausmann L (2010) Breeding for fungal disease resistant grapevines, Genetics, Genomics and Breeding of Grapes, Science Publishers
Fungal Resistance Disposition
Cabernet Sauvignon pistils at BBCH 61-65 (flowering to full bloom) exhibit characteristic fused carpel organization with completely unified ovary showing no external evidence of carpel boundaries, single coherent style with no trace of separate carpel styles, and unified stigma with only subtle bilobing as evidence of bicarpellate origin, with fusion so complete that carpel identity is evident only through internal bilocular ovary organization and bilateral stigma papillae distribution. Riesling pistil developmental studies show carpel fusion initiating at BBCH 52 when carpel primordia are only 0.08-0.12 mm, with fusion proceeding rapidly (completing by BBCH 53) creating unified structure by the time significant organ elaboration begins, resulting in mature pistils (BBCH 59) with essentially no evidence of original separate carpel identity except bilocular ovary. Gewürztraminer pistils show typical complete fusion with integrated vascular tissues, unified style transmitting tract supporting pollen tube growth to both ovary locules, and stigma showing only minimal bilobing (two slight lateral extensions) as residual evidence of two-carpel origin. Rare Syrah flowers (<0.02% frequency) exhibit fusion failures producing partially separated carpel pistils with partially divided styles or deeply cleft stigmas showing incomplete fusion, typically associated with impaired fruit set (<15% versus normal 54-62%) demonstrating functional importance of complete carpel fusion.
A fused carpel compound pistil is a compound pistil in which the constituent carpels (invariably two in Vitis) show complete or nearly complete fusion creating a unified structure with integrated ovary, style, and stigma tissues, representing the normal developmental outcome in Vitis flowers where early and extensive carpel fusion during organogenesis (BBCH 52-53) produces pistils in which individual carpel identity is largely obscured except through anatomical details like locule number and bilateral stigma organization. Fused carpel compound pistils are characterized by= (1) unified ovary structure with carpel boundaries not externally visible and with internal organization (typically bilocular with central septum) representing the only clear evidence of two-carpel origin (2) completely fused style forming a single coherent cylinder with no evidence of separate carpel styles, though internal transmitting tract tissue may show subtle bilateral organization (3) unified or subtly bilobed stigma with carpel fusion creating either a completely unified receptive surface or a stigma with shallow bilateral lobing as the only evidence of two-carpel origin (4) integrated vascular tissues with carpel vascular strands fused into unified vascular supply to the compound pistil (5) early developmental fusion occurring during primordium stages (BBCH 52-53) before significant organ elaboration, resulting in mature structures showing minimal evidence of separate carpel identity. The fused carpel condition represents the universal state in normal Vitis flowers, with complete fusion being the ancestral and developmentally stable outcome of pistil organogenesis across all species and cultivars. The degree of fusion in Vitis is extensive compared to many other plant families where compound pistils may show partially free carpel walls, separate styles, or clearly separated stigma lobes in Vitis, only the internal septum dividing the ovary into two locules and the sometimes-subtle stigma bilobing provide clear evidence of the bicarpellate origin. The developmental timing and completeness of fusion critically affect pistil function= early fusion (during primordium stages) allows integrated development of all pistil tissues including unified style transmitting tract supporting efficient pollen tube growth to both locules, integrated stigma providing unified receptive surface for pollen capture, and coordinated ovule development in both locules incomplete fusion or fusion failures (rare developmental abnormalities) can produce pistils with partially or completely separate carpel elements (separate styles, deeply divided stigmas, partially separated ovary chambers) potentially impairing function through disrupted tissue coordination. The fused carpel organization creates several functional advantages= unified stigma provides larger, more continuous receptive surface potentially increasing pollen capture efficiency compared to separate stigmatic surfaces unified style creates single transmitting tract potentially increasing efficiency of pollen tube guidance and support compared to separate pathways unified ovary develops as single integrated fruit rather than aggregate of separate carpel fruits, potentially optimizing resource allocation and fruit dispersal characteristics. The molecular and cellular mechanisms underlying carpel fusion in Vitis involve= expression of carpel identity genes (C-class and D-class MADS-box genes) specifying carpel fate expression of boundary genes preventing separation and promoting fusion at carpel margins coordinated cell proliferation and differentiation creating unified tissues programmed cell death potentially removing boundary cells to facilitate fusion hormone signaling (particularly auxin) coordinating growth patterns promoting fusion.
Fused_Carpel_Compound_Pistil emphasizes the completeness of carpel fusion characteristic of normal Vitis pistil development and is essentially synonymous with typical pistil structure in grapevines. The fused carpel specification highlights that carpel fusion is extensive and early, creating highly integrated structures with minimal evidence of separate carpel identity. This contrasts with some other plant families where compound pistils may retain more obvious evidence of constituent carpels (partially free walls, separate styles, divided stigmas). Understanding fused carpel organization is important for interpreting pistil anatomy, for understanding developmental processes underlying pistil formation, and for recognizing rare fusion failures or abnormalities that may affect fruit set and berry development.
completely fused pistil
integrated carpel pistil
verwachsenes Karpell-Gynoeceum
vollständig fusionierter Stempel
vollständig verschmolzener Stempel
Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development. In= Roubelakis-Angelakis, K.A. (ed.) Molecular Biology & Biotechnology of the Grapevine, pp. 186-189 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 93-95.
Fused Carpel Compound Pistil
Fv/Fm ratio assessment tracking heat stress and recovery in Chardonnay reveals the time course of photoinhibition, with pre-stress Fv/Fm of 0.83 declining to 0.68 immediately after heat event (40°C, 4 hours), recovering to 0.74 after one night (partial PSII repair), 0.79 after two nights (substantial recovery), and 0.82 after three nights (full recovery), demonstrating how Fv/Fm monitoring enables both damage assessment and recovery tracking, with the recovery trajectory indicating that stress severity did not exceed the repair capacity of the D1 protein turnover system.
Fv/Fm ratio quality is a chlorophyll fluorescence quality that represents the maximum quantum efficiency of photosystem II photochemistry measured in dark-adapted leaves, characterizing the proportion of absorbed light energy that can potentially drive electron transport when all PSII reaction centers are open and photoprotective mechanisms are relaxed. Within the BFO framework, this quality specifically addresses the maximum PSII efficiency dimension of fluorescence, distinguishing potential (dark-adapted) from operational (light-adapted) quantum yield while recognizing that Fv/Fm provides the most widely used and robust indicator of photosynthetic apparatus integrity and stress status. The ontological significance lies in Fv/Fm serving as the universal stress indicator in plant physiology, where the ratio of variable to maximum fluorescence (Fv/Fm = (Fm-Fo)/Fm) provides a dimensionless efficiency parameter that is remarkably consistent across unstressed C₃ plants (0.80-0.84) regardless of species, making deviation below this range a reliable indicator of photoinhibition, damage, or stress affecting PSII function. The photophysical basis for Fv/Fm involves the measurement of minimal fluorescence (Fo) when all PSII reaction centers are open (dark-adapted state, QA fully oxidized) and maximum fluorescence (Fm) when all centers are closed (saturating pulse, QA fully reduced), with the difference (Fv = Fm-Fo) representing the variable fluorescence that reflects PSII photochemical capacity. The measurement protocol for Fv/Fm requires dark adaptation (typically 20-30 minutes for full relaxation of NPQ) followed by application of a saturating light pulse (>3000 μmol/m²/s for 0.8-1.0 s) that transiently closes all reaction centers, with Fo measured immediately before and Fm at the peak of the saturating pulse. The stress responses of Fv/Fm show characteristic reduction patterns, with mild stress causing small reductions (0.75-0.80) from which overnight recovery is possible, while severe stress causes larger reductions (< 0.70) indicating damage requiring days for recovery or indicating permanent PSII damage. Understanding Fv/Fm ratio quality enables rapid stress screening, assessment of photoinhibition severity, and monitoring of recovery from stress events through this universally applicable efficiency parameter.
Fv_Fm_Ratio_Quality addresses the maximum PSII quantum efficiency in dark-adapted state, providing the universal stress indicator in plant physiology. This quality requires proper dark adaptation for valid measurement. The class supports stress screening and photoinhibition assessment through this remarkably consistent parameter.
Fv/Fm-Verhältnis
PSII efficiency
PSII-Effizienz
dark-adapted quantum yield
maximale Quantenausbeute
maximale photochemische Effizienz
maximum photochemical efficiency
maximum quantum yield
Björkman O, Demmig B (1987) Photon yield of O₂ evolution and chlorophyll fluorescence characteristics at 77 K among vascular plants of diverse origins. Planta 170(4): 489-504 Maxwell K, Johnson GN (2000) Chlorophyll fluorescence — a practical guide. Journal of Experimental Botany 51(345): 659-668
Fv Fm Ratio Quality
Delaware grape shows strong GA-responsive seedlessness disposition, with GA₃ application (25-50 ppm) at 50% capfall reducing seed development to trace levels while increasing berry size from 1.5 g to 2.5-3.0 g. Kyoho responds to GA treatment at bloom with 80-95% reduction in normal seed formation. Application timing is critical, with treatment 10-14 days pre-bloom for berry thinning and at full bloom for seedlessness induction being standard protocols in Japanese viticulture. GA responsiveness varies from complete seed suppression (Delaware) to partial reduction (Muscat Bailey A, 40-60% seed reduction).
A seedlessness disposition that is the inherited capacity of a Vitis vinifera plant to produce functionally seedless berries following exogenous gibberellic acid (GA₃) application during bloom or early fruit development, through GA-induced inhibition of seed development in otherwise seeded cultivars.
GA-responsive seedlessness differs mechanistically from genetic stenospermocapy, involving hormone-mediated suppression of ovule development or fertilization rather than post-fertilization embryo abortion. The trait is exploited commercially primarily in East Asian table grape production. GA treatment also affects cluster loosening, berry sizing, and skin texture. Cultivar-specific protocols are essential, as excessive GA concentration causes berry shatter and cluster deformation.
GA-induced seed suppression
GA-induzierte Kernlosigkeit
Gibberellin-responsives Apyrenie
gibberellin-induced seedlessness
hormone-induced apyreny
hormonell induzierte Samenlosigkeit
Iwahori S et al. (1968) Gibberellin-induced parthenocarpy in grape. Plant Physiology 43:333-337 Lu J et al. (1997) Effects of gibberellic acid on berry development and cluster morphology of several grape cultivars. Fruit Varieties Journal 51:219-227
GA Responsive Seedlessness Disposition
GABA content quality in Kyoho table grapes from Japan has become a marketed quality attribute, with postharvest treatments (controlled atmosphere, temperature manipulation) employed to enhance GABA accumulation from baseline 50-100 mg/kg to 200-400 mg/kg, enabling health-positioned marketing of GABA-enriched grapes while such deliberate GABA enhancement has limited application in wine production, understanding natural GABA accumulation (100-300 mg/L in stressed fruit) provides winemakers with a potential stress indicator complementing visual and conventional ripening assessments.
GABA_Content_Quality represents the chemical quality characterizing the concentration of gamma-aminobutyric acid (GABA, 4-aminobutanoic acid) in grape tissue and wine, capturing the non-protein amino acid that accumulates under stress conditions and has emerged as a bioactive compound of interest for potential health effects. GABA is synthesized from glutamate through the action of glutamate decarboxylase (GAD), a stress-responsive enzyme activated by low pH, anoxia, mechanical damage, and various biotic and abiotic stresses. In grape berries, GABA accumulation increases substantially during ripening and in response to stress, with concentrations ranging from 50-500 mg/L in must depending on cultivar, ripening conditions, and stress exposure. The stress-responsive nature of GABA makes it a potential biomarker for berry stress history, with elevated GABA suggesting challenging ripening conditions. As a non-protein amino acid, GABA is not incorporated into proteins but participates in nitrogen metabolism and may have signaling functions in plant stress response. During fermentation, Saccharomyces cerevisiae can utilize GABA as a nitrogen source, though it is not among the preferred amino acids, meaning GABA may persist into finished wine at levels reflecting must content modified by yeast uptake. Interest in wine GABA content has increased with research suggesting health effects including blood pressure reduction and anxiety modulation, though concentrations in wine are substantially below therapeutic doses. Some winemakers and producers have explored production methods to enhance GABA content, including extended prefermentation handling and selection of GABA-accumulating cultivars or clones.
GABA_Content_Quality addresses a non-protein amino acid distinct from the standard amino acids incorporated into proteins. The stress-responsive accumulation of GABA links its content to ripening conditions and berry physiology. The emerging health interest in GABA creates potential marketing relevance beyond traditional quality parameters. GABA analysis requires methods capable of detecting non-standard amino acids, with HPLC methods generally capable of GABA quantification though derivatization conditions may need optimization. The regulatory role of GABA in plant stress signaling remains an active research area with potential implications for understanding quality formation in stressed grapevines.
4-Aminobutyratgehalt
4-aminobutyrate content
GABA level
GABA-Gehalt
Gamma-Aminobuttersäure-Konzentration
gamma-aminobutyric acid concentration
stress-responsive non-protein amino acid quality
stressresponsive Nichtprotein-Aminosäurequalität
Bouché N and Fromm H (2004) GABA in Plants: Just a Metabolite? Trends in Plant Science 9(3): 110-115 Shelp BJ et al. (1999) Metabolism and Functions of Gamma-Aminobutyric Acid. Trends in Plant Science 4(11): 446-452
GABA Content Quality
Mechanistic investigation of Gall_Formation_Resistance_Disposition at the University of Natural Resources and Life Sciences, Vienna, employed time-course microscopy comparing cellular responses to phylloxera probing in resistant V. vinifera 'Grüner Veltliner' versus susceptible V. riparia 'Gloire de Montpellier'. Detached leaves were infested with synchronized first-instar crawlers, and feeding sites were sectioned at 6, 12, 24, 48, and 96 hours for histological analysis and cell death visualization (trypan blue staining). On susceptible V. riparia, phylloxera stylet insertion at 6 hours was followed by localized cell division visible at 24 hours, coordinated tissue bulging at 48 hours, and recognizable gall primordia at 96 hours, with no cell death in adjacent tissues. On resistant 'Grüner Veltliner', equivalent stylet insertion occurred at 6 hours, but by 12 hours, cells within 200μm of the feeding site showed intense trypan blue staining indicating cell death. By 24 hours, a zone of collapsed, dead cells surrounded the probing site, and no cell division was observed. Attempted gall formation was aborted, and phylloxera either died or abandoned the leaf. Gene expression analysis revealed that 'Grüner Veltliner' showed rapid (6-12 hour) upregulation of defense genes including PR-1, chitinase, and phenylalanine ammonia-lyase at feeding sites, while V. riparia showed no defense gene induction. These findings demonstrate that Gall_Formation_Resistance_Disposition in V. vinifera involves active recognition and hypersensitive response rather than passive incompatibility.
A Gall_Formation_Resistance_Disposition is a Foliar_Phylloxera_Resistance_Disposition that inheres in a Vitis plant leaf and is realized through cellular mechanisms that prevent the coordinated cell proliferation and tissue reorganization required for functional phylloxera gall development following insect probing and feeding attempts. This disposition represents the specific mechanism underlying foliar phylloxera resistance, focusing on the host tissue response that determines whether phylloxera feeding can induce the characteristic pouch galls or is aborted before gall formation proceeds. Successful phylloxera leaf gall formation requires a precise sequence of host responses including localized suppression of cell death, induction of cell division in adaxial mesophyll, coordinated cell expansion creating the gall chamber, differentiation of nutritive tissue lining the gall, and maintenance of a favorable environment for the enclosed insect. Gall_Formation_Resistance_Disposition disrupts this sequence through alternative host responses including hypersensitive cell death at feeding sites sacrificing potential gall tissues, failure of cells to respond to phylloxera-secreted growth regulators, production of toxic or deterrent compounds at probing sites, and anatomical features (thick cuticle, dense trichomes) preventing initial feeding establishment. V. vinifera exhibits strong Gall_Formation_Resistance_Disposition characterized by rapid cell death in response to phylloxera probing, explaining the near-absence of functional leaf galls on European cultivars despite their catastrophic root susceptibility. This tissue-specific resistance provides insight into the distinct genetic control of phylloxera defense in different plant organs.
Gall_Formation_Resistance_Disposition is_a Foliar_Phylloxera_Resistance_Disposition. This class addresses the specific cellular mechanism preventing leaf gall development, mechanistically analogous to hypersensitive response against pathogens. The presence of strong gall formation resistance in V. vinifera, which lacks root resistance, demonstrates that phylloxera defense is controlled by tissue-specific genetic systems. Understanding these mechanisms could potentially inform strategies for enhancing root resistance, though the distinct biology of root versus leaf tissues may limit direct translation.
Gallenbildungs-Resistenz-Disposition
Galleninitiations-Resistenz
Gallenmorphogenese-Blockade
gall initiation resistance
gall morphogenesis block
leaf gall suppression
Sterling C (1952) Ontogeny of the phylloxera gall of grape leaf, American Journal of Botany 39:6-15 Forneck A et al. (2002) Histochemistry and anatomy of phylloxera galls on Vitis spp., Vitis 41:93-97 Nabity PD et al. (2013) Leaf-galling phylloxera on grapes reprograms host metabolism and morphology, Proceedings of the National Academy of Sciences 110:16663-16668
Gall Formation Resistance Disposition
Seasonal monitoring of gap frequency quality in Wachau Grüner Veltliner tracked canopy development and management effects from budbreak through harvest. Early season gap frequency quality at BBCH 15 showed 85% gaps reflecting sparse emerging canopy. By bloom at BBCH 65, gap frequency declined to 42% as rapid shoot growth filled canopy volume. Pre-véraison at BBCH 79, gap frequency quality had decreased to only 12% indicating need for intervention, prompting basal leaf removal that immediately improved gap frequency to 31%. Post-véraison stability maintained gap frequency quality between 28-32% through harvest, meeting the 20-40% optimal target throughout the critical ripening period. Fruit from optimized canopies showed 18% higher concentration of characteristic pepper-spice aromatics compared to non-managed comparison blocks with gap frequency remaining below 15% at harvest.
Gap frequency quality is a canopy gaps quality that inheres in a grapevine canopy system and represents the proportion of point quadrat probe insertions or equivalent sampling positions that traverse the canopy without encountering any leaf contact, providing standardized quantification of canopy openness that directly relates to the probability that light rays, air currents, or spray droplets will successfully penetrate to interior canopy positions without interception by foliage. This quality serves as the primary metric for canopy gaps characterization, enabling objective comparison of canopy management treatments and training system effects on canopy openness using repeatable measurement protocols. The ontological basis of gap frequency quality rests on the statistical sampling principle that systematic probe insertions through the canopy at regular intervals provide representative characterization of the overall gap distribution, with the proportion of gap encounters to total insertions estimating the true fraction of the canopy cross-section permitting unobstructed passage. The measurement protocol involves inserting a thin rigid probe horizontally through the canopy at predetermined height intervals and horizontal positions along the row, recording whether each insertion contacts zero leaves (gap), one leaf, or multiple leaves (layered foliage), with gap frequency computed as the percentage of zero-contact insertions from the total sample size. Optimal gap frequency quality for wine grape production generally targets values between 20-40%, representing canopy architecture where one-fifth to two-fifths of probe insertions encounter no foliage, corresponding to light transmission characteristics suitable for fruit zone illumination while maintaining sufficient canopy density for adequate total photosynthesis and fruit protection. Gap frequency quality below 10% indicates dense canopies with severely restricted light penetration likely to compromise fruit composition and elevate disease pressure, while values exceeding 50% may indicate sparse canopies with insufficient leaf area for optimal fruit ripening or excessive exposure risking sunburn damage in hot climates. The relationship between gap frequency quality and other canopy metrics follows predictable patterns, with gap frequency inversely related to leaf layer number such that higher LLN values mathematically require lower gap frequencies, though the specific relationship depends on the spatial distribution of leaves within the canopy volume.
Gap_Frequency_Quality is_a Canopy_Gaps_Quality, representing the quantitative proportion of openings in the canopy as measured through point quadrat analysis. This class addresses the percentage gaps metric specifically, distinct from Gap_Size_Quality which addresses the dimensions of individual openings. The optimal target of 20-40% provides key quantitative benchmark. Gap frequency inversely correlates with leaf layer number, with both metrics derived from point quadrat sampling providing complementary density characterization.
Lückenanteil
Lückenfrequenz
Prozent Lücken
canopy gap ratio
gap percentage
opening frequency
percent gaps
Öffnungshäufigkeit
Smart RE (1985) Principles of grapevine canopy microclimate manipulation with implications for yield and quality: a review. American Journal of Enology and Viticulture 36:230-239 Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 44-59 Meyers JM, Vanden Heuvel JE (2008) Enhancing the precision and spatial acuity of point quadrat analyses via calibrated exposure mapping. American Journal of Enology and Viticulture 59:425-431
Gap Frequency Quality
Gap size quality evaluation in Adelaide Hills Shiraz compared canopy management approaches achieving equivalent 30% gap frequency through different gap size distributions. Precision shoot positioning at 10 cm spacing created uniform gap size quality with openings averaging 6 cm width occurring regularly between each shoot pair, producing homogeneous fruit zone illumination with coefficient of variation in fruit surface PAR of 18%. Conventional positioning with less precise shoot placement produced irregular gap size quality with openings ranging from 2 cm to 25 cm, creating heterogeneous exposure with fruit PAR CV of 42%. The uniform small-gap treatment produced more consistent fruit composition with berry-to-berry anthocyanin CV of 24% versus 38% in the variable gap treatment, while also reducing sunburn incidence from 7% to 2% by eliminating the large gaps that permitted excessive direct solar exposure during afternoon periods.
Gap size quality is a canopy gaps quality that inheres in a grapevine canopy system and represents the spatial dimensions of individual openings within the foliar mass characterized by their horizontal extent, vertical extent, and overall area as determined by the spacing between adjacent shoots, the arrangement of leaves along shoots, and the effectiveness of shoot positioning in creating organized canopy structure with regular gap distribution. This quality provides geometric characterization of canopy openings that complements gap frequency assessment by capturing the nature of gaps that exist, recognizing that equivalent total gap area can manifest as either numerous small openings distributed throughout the canopy or fewer large openings that create localized exposure zones with different functional consequences. The ontological significance of gap size quality lies in its influence on the spatial patterns of light penetration, wherein many small well-distributed gaps create relatively uniform diffuse illumination of interior canopy positions while few large gaps produce heterogeneous exposure with some interior regions receiving direct beam radiation and others remaining heavily shaded. The physics of light transmission through canopy gaps depends critically on gap dimensions relative to solar angle, with small gaps permitting penetration primarily of diffuse radiation while larger gaps allow direct beam passage during periods when solar position aligns with the gap geometry, creating sunfleck patterns that may provide beneficial or excessive exposure depending on timing and intensity. Management factors influencing gap size quality include shoot spacing along cordons or canes, which establishes the fundamental between-shoot gap potential lateral shoot development, which can fill gaps between primary shoots and the precision of shoot positioning, which affects whether gaps occur in organized positions or randomly throughout the canopy volume. The measurement of gap size quality extends beyond simple point quadrat assessment to approaches capturing the spatial extent of individual openings, including systematic gap width measurement at fruit zone height, image analysis of canopy photographs quantifying gap area distribution, and light transmission mapping using sensor grids that characterize the spatial pattern of penetrating radiation. Optimal gap size quality depends on climate and production objectives, with hot climate viticulture potentially benefiting from numerous small gaps providing ventilation without excessive direct exposure, while cool climate production may tolerate or benefit from larger gaps that maximize fruit zone illumination for phenolic development.
Gap_Size_Quality is_a Canopy_Gaps_Quality, representing the dimensional characterization of individual openings complementary to Gap_Frequency_Quality which addresses the proportion of openings. This class recognizes that gap distribution patterns matter beyond total gap percentage, with uniform small gaps generally preferable to irregular large gaps for consistent fruit exposure. Image analysis and systematic gap measurement provide assessment approaches beyond standard point quadrat methodology. Related to the spatial uniformity of fruit exposure and ripening.
Lückenausdehnung
Lückengröße
canopy aperture size
gap area distribution
gap dimensions
opening size
Öffnungsdimensionen
Öffnungsflächenverteilung
Dokoozlian NK, Kliewer WM (1995) The light environment within grapevine canopies. II. Influence of leaf area density on fruit zone light environment and some canopy assessment parameters. American Journal of Enology and Viticulture 46:219-226 Schultz HR (1995) Grape canopy structure, light microclimate and photosynthesis. I. A two-dimensional model of the spatial distribution of surface area densities and leaf ages in two canopy systems. Vitis 34:211-215 Meyers JM, Vanden Heuvel JE (2014) Row orientation effects on sunlight quality in vineyards of the eastern United States. Australian Journal of Grape and Wine Research 20:435-446
Gap Size Quality
Cabernet Sauvignon gas exchange lenticel at 14 years post-planting shows 2.8-3.6 mm diameter structure with loose cork cells and 58-65% air space volume enabling oxygen influx 0.7-1.1 μmol O₂/cm² lenticel/hour and carbon dioxide efflux 0.8-1.3 μmol CO₂/cm²/hour (RQ 1.14), lenticel density 5-7 per cm² collectively providing 3.2-4.1% bark area sufficient for supporting 0.8-1.4 μmol O₂/g bark/hour respiration in living inner bark (phloem parenchyma, companion cells, cambium consuming oxygen for ATP production maintaining transport, storage, and growth functions), lenticels maintaining gas exchange year-round including winter when supporting 0.3-0.5 μmol O₂/g/hour basal metabolism in dormant tissues
Gas exchange lenticel is lenticel emphasizing its functional role in facilitating oxygen influx and carbon dioxide efflux through otherwise impermeable cork bark, enabling aerobic respiration in living inner bark and cambium tissues, representing the critical gas exchange pathway maintaining metabolic activity in bark tissues surrounded by waterproof covering. Gas exchange lenticel oxygen supply function includes= atmospheric oxygen diffusion inward (O₂ moving down concentration gradient from 21% atmospheric through lenticel air spaces to oxygen-depleted inner bark), diffusion rate (typically 0.4-1.2 μmol O₂/cm² lenticel area/hour depending on lenticel structure, air space configuration, and respiratory demand), oxygen delivery to living tissues (lenticel-supplied oxygen reaching living phloem parenchyma, companion cells, cambial cells, and phelloderm supporting aerobic respiration), supporting ATP production (oxygen enabling mitochondrial oxidative phosphorylation producing energy for cellular processes), and preventing anaerobic conditions (adequate oxygen supply through lenticels preventing potentially damaging fermentation and anaerobic metabolism in bark tissues). Gas exchange lenticel carbon dioxide removal function includes= CO₂ efflux from respiring tissues (carbon dioxide produced by inner bark respiration diffusing outward through lenticel pathways), concentration gradient (CO₂ partial pressure 1-5% in inner bark declining to 0.04% atmospheric creating driving force for outward diffusion), diffusion rate (0.5-1.5 μmol CO₂/cm² lenticel/hour, slightly exceeding O₂ influx reflecting respiratory quotient near 1.0-1.2), preventing CO₂ accumulation (lenticel gas exchange removing respiratory CO₂ preventing buildup that could inhibit metabolism or acidify tissues), and maintaining pH balance (CO₂ removal preventing carbonic acid accumulation helping maintain appropriate cellular pH for enzyme function). Gas exchange lenticel structural features enabling function includes= loose complementary tissue (cork cells with minimal contact creating tortuous air pathways enabling gas diffusion), large intercellular spaces (air channels occupying 50-70% of lenticel volume providing high permeability to gases), depth through cork layer (lenticels extending 0.5-3 mm from surface to living phelloderm or phloem ensuring complete gas exchange pathway through waterproof barrier), sufficient cross-sectional area (lenticel dimensions 1-4 mm diameter providing adequate surface area for diffusion), and appropriate density (2-8 lenticels per cm² bark surface collectively providing sufficient gas exchange capacity). Gas exchange lenticel quantitative gas flux includes= lenticel oxygen influx (0.4-1.2 μmol O₂/cm² lenticel/hour typical rates depending on respiratory demand and lenticel structure), total bark oxygen demand (living inner bark consuming 0.5-1.5 μmol O₂/g fresh weight/hour requiring adequate lenticel supply), lenticel area requirements (lenticel coverage 2-5% of bark surface area typically sufficient for meeting respiratory oxygen demand), carbon dioxide production (inner bark respiration producing 0.6-1.8 μmol CO₂/g/hour requiring lenticel removal pathway), and respiratory quotient (CO₂ produced / O₂ consumed ratio typically 1.0-1.2 reflecting primarily carbohydrate respiration in bark tissues). Gas exchange lenticel seasonal variation in function includes= peak growing season activity (April-September, maximum gas exchange rates 0.8-1.4 μmol O₂/cm²/hour supporting high inner bark metabolic activity), dormant season reduced flux (November-February, decreased rates 0.2-0.6 μmol O₂/cm²/hour reflecting lower respiratory demand in quiescent tissues), closing layer effects (some cultivars forming compacted cork layer at lenticel base during dormancy reducing gas exchange 30-60% conserving water while maintaining minimal oxygen supply), spring reactivation (March-April, increasing gas exchange as metabolism resumes and closing layers disrupted), and year-round critical function (even minimal winter gas exchange essential for maintaining basal metabolism in dormant but living bark tissues). Gas exchange lenticel enabling inner bark metabolism includes= supporting phloem transport (aerobic respiration in companion cells and sieve tubes enabling active sugar loading and phloem maintenance), maintaining storage tissue viability (parenchyma cells requiring oxygen for sustaining cellular integrity and regulating starch synthesis/hydrolysis), enabling cambial division (vascular cambium and cork cambium requiring aerobic metabolism for cell division and secondary growth), supporting wound healing (callus tissue respiration depending on lenticel oxygen supply for proliferation and differentiation), and maintaining dormancy metabolism (even dormant tissues requiring minimal respiration for cellular maintenance, cryoprotectant synthesis, and preparation for spring reactivation). Gas exchange lenticel vulnerability and limitations includes= blocking by water (rain or high humidity potentially filling lenticel air spaces temporarily reducing gas exchange), obstruction by debris (dust, fungi, lichens potentially coating lenticels reducing permeability), freeze damage (ice formation in lenticel tissues potentially disrupting structure), pathogen entry potential (porous lenticels representing possible infection route though typically minor compared to wounds), and insufficient lenticel density (inadequate lenticel coverage potentially causing oxygen limitation reducing bark vigor). Gas exchange lenticel balance with impermeability includes= small proportional area (lenticels typically <3-5% of bark surface minimizing water loss while enabling gas exchange), tortuous air pathways (intercellular spaces providing high gas permeability but low water vapor transmission maintaining mostly waterproof covering), closing layers seasonal formation (some cultivars forming barriers during dormancy improving water conservation while maintaining minimal gas exchange), optimal lenticel density (2-8 per cm² providing adequate respiration support without compromising bark protective functions), and critical compromise (lenticels solving challenge of enabling aerobic metabolism through waterproof pathogen-resistant bark). Gas exchange lenticel assessment and measurement includes= visual lenticel identification (observing lighter colored spots on bark surface, counting lenticel density), oxygen electrode measurements (quantifying oxygen concentration gradients across bark and lenticel structures), gas flux chambers (measuring actual O₂ influx and CO₂ efflux rates through bark sections with lenticels), respiratory quotient determination (calculating CO₂/O₂ ratio indicating metabolic substrate), and lenticel permeability assays (comparing gas diffusion through lenticel tissue versus intact cork quantifying facilitation). Gas exchange lenticel significance in bark function includes= absolutely essential for inner bark survival (living tissues requiring continuous oxygen supply, lenticels providing only pathway through impermeable cork), determining metabolic capacity (adequate lenticel function enabling normal inner bark respiration and metabolism, inadequate lenticels potentially causing oxygen limitation and reduced activity), enabling trunk longevity (functional gas exchange maintaining cambial viability supporting decades of continued growth and bark renewal), and balancing protection with permeability (lenticels exemplifying elegant solution to maintaining waterproof protective covering while enabling necessary gas exchange for tissue survival).
Gas_Exchange_Lenticel emphasizes respiratory function and is distinguished from Lenticel (anatomical structure), Surface_Lenticel (location emphasis), and general periderm by focus on oxygen supply and CO₂ removal. Critical for understanding aerobic metabolism support in bark, gas flux quantification, seasonal variation in gas exchange, oxygen limitation prevention, balance between impermeability and permeability, and essential role as sole gas exchange pathway through waterproof cork enabling inner bark and cambium survival and function.
Aerating lenticel
Aerationsstruktur
Atmungspore
Breathing structure
Gasaustausch-Lenticelle
Respiratory pore
Gas Exchange Lenticel
The GBS data for a biparental mapping population of 300 F1 individuals from a Riesling × Cabernet Sauvignon cross were generated using ApeKI restriction enzyme digestion and Illumina HiSeq 2500 sequencing at 5× average coverage per sample — after SNP discovery and filtering using the TASSEL-GBS pipeline aligned to the PN40024 v2 reference genome, 22,450 high-quality biallelic SNP markers (minimum read depth 5, maximum missing data 20%, minor allele frequency above 5%) distributed across all 19 chromosomes at an average spacing of 21 kb provided the marker framework for construction of a high-density genetic linkage map totaling 1,245 cM, QTL mapping for véraison date, cluster compactness, and powdery mildew resistance, and genomic prediction models for 12 quantitative traits with prediction accuracies of 0.35-0.68 depending on trait heritability.
A genetic information entity that records the genotype data generated through genotyping-by-sequencing (GBS) — a reduced-representation sequencing approach that uses restriction enzyme digestion to sample a reproducible subset of the genome, producing tens of thousands of SNP genotypes per accession at reduced cost compared to whole-genome sequencing, applicable to large population studies, genetic mapping, genomic selection, and diversity analysis in Vitis vinifera and interspecific Vitis populations.
GBS_Data documents the sequencing-based genotyping approach that occupies the cost-performance space between fixed-content SNP arrays (lower cost per sample but limited to predefined SNP positions) and whole-genome resequencing (comprehensive but expensive for large populations). GBS exploits restriction enzyme digestion to reduce genome complexity by 10-100 fold, sequencing only the fragments adjacent to restriction sites and thereby sampling a reproducible, evenly distributed subset of the genome at sufficient depth for reliable SNP calling: the approach was adapted for Vitis from the original GBS protocol (Elshire et al. 2011) using enzymes such as ApeKI, PstI-MspI, and EcoT22I that produce fragment libraries of appropriate complexity for the 475 Mb grape genome. The principal advantages of GBS over array genotyping are the discovery of novel variants in each dataset (not limited to predefined SNP positions), applicability to any Vitis species or interspecific hybrid without ascertainment bias, and scalability to thousands of samples through multiplexed library preparation. The principal limitations are variable missing data rates (5-30% per position depending on sequencing depth and restriction site polymorphism), computational intensity of SNP calling and imputation, and reduced reproducibility compared to array-based genotyping for database cross-referencing across studies.
GBS-Daten
Genotypisierung-durch-Sequenzierung-Daten
RAD-seq genotype data
genotyping-by-sequencing data
reduced representation sequencing data
reduzierte Genomrepräsentations-Daten
Elshire RJ et al. (2011) A robust, simple genotyping-by-sequencing (GBS) approach for high diversity species, PLoS ONE 6: e19379 Barba P et al. (2014) Grapevine powdery mildew resistance and target site analysis of Ren3 gene family, Molecular Plant Pathology 15: 350-361 Hyma KE et al. (2015) Heterozygous mapping strategy (HetMappS) for high resolution genotyping-by-sequencing markers: a case study in grapevine, PLoS ONE 10: e0134880
GBS Data
GDC suitability quality evaluation in Finger Lakes compared cultivar performance under Geneva Double Curtain training designed for the region's vigorous growing conditions. Concord demonstrated excellent GDC suitability quality with high vigor filling both curtains to target 0.9 m depth, strongly pendant growth habit maintaining curtain separation with minimal intervention, and yield increase of 45% compared to umbrella Kniffin training with equivalent fruit quality for juice and jelly production. Cayuga White showed good GDC suitability quality for hybrid wine production with adequate vigor and moderate trailing habit, though requiring quarterly curtain separation to prevent bridging. Riesling exhibited poor GDC suitability quality with insufficient vigor producing only 60% curtain fill despite fertile site conditions, combined with semi-upright growth habit causing persistent curtain convergence, confirming that this moderate-vigor, upright-growing Vitis vinifera cultivar is better suited to single-canopy vertical systems regardless of site vigor potential.
GDC suitability quality is a trellis adaptation quality that inheres in a grapevine cultivar or clone and represents the degree of compatibility with Geneva Double Curtain training systems characterized by bilateral cordon division onto two parallel horizontal wires separated by 1.0-1.2 m spacing with shoots trained downward to create two distinct pendant canopy curtains that increase total exposed leaf area while improving light penetration to both curtain surfaces and the central inter-curtain zone. This quality determines whether a cultivar possesses the vigor capacity, trailing growth habit, and physiological characteristics necessary to fill and maintain two separated canopy curtains achieving the yield and quality benefits that justify the increased infrastructure complexity of divided canopy systems. The ontological significance of GDC suitability quality lies in its role enabling productive management of high-vigor cultivars and sites where single-canopy systems fail to achieve adequate vine balance, providing a vigor management tool through canopy division that increases sink capacity through doubled exposed leaf area and improved light distribution compared to dense single canopies. The biological requirements for high GDC suitability quality include sufficient vigor to fill two canopy curtains with adequate shoot density and length, naturally pendant or trailing growth habit enabling shoots to hang downward without persistent repositioning requirements, moderate lateral development that fills curtains without causing inter-curtain bridging, appropriate internode length for curtain depth dimensions, and cluster characteristics suitable for the exposed environment of pendant fruit positioning. Cultivars with insufficient vigor produce sparse curtains failing to achieve the light interception benefits of divided canopy systems, while excessively vigorous cultivars may overwhelm even divided configurations requiring additional management interventions. The trailing growth habit requirement fundamentally constrains GDC suitability to cultivars exhibiting this characteristic, with strongly upright cultivars proving poorly adapted despite adequate vigor. Assessment of GDC suitability quality involves evaluation of curtain fill adequacy, shoot orientation consistency, inter-curtain separation maintenance, fruit zone exposure characteristics, and comparison of yield-quality outcomes to alternative training approaches for the cultivar-site combination.
GDC_Suitability_Quality_(Geneva_Double_Curtain) is_a Trellis_Adaptation_Quality, representing compatibility with the classic divided canopy system developed for high-vigor conditions. This class addresses the specific requirements of GDC including high vigor, trailing growth habit, and appropriate lateral development. Distinguished from Lyre_Suitability_Quality which addresses the related but geometrically different lyre configuration. Most relevant for vigorous sites and cultivars where single-canopy systems prove inadequate for vine balance management. Originally developed for Concord in New York but applicable to diverse vigorous cultivar-site combinations.
Doppelvorhang-Tauglichkeit
GDC compatibility
GDC-Eignung
Geneva Double Curtain adaptation
Geneva-System-Kompatibilität
divided canopy suitability
double curtain training fitness
geteilte Laubwand-Eignung
Shaulis NJ et al. (1966) Response of Concord grapes to light, exposure and Geneva Double Curtain training. Proceedings of the American Society for Horticultural Science 89:268-280 Smart RE (1988) Shoot spacing and canopy light microclimate. American Journal of Enology and Viticulture 39:325-333 Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review. American Journal of Enology and Viticulture 60:251-268
GDC Suitability Quality (Geneva Double Curtain)
The GDD base 10°C for the Napa Valley (Oakville, California, 38.43°N) documents a mean annual accumulation of 1,850-2,050 GDD over the April 1 - October 31 period (1991-2020 normal) — this thermal resource places Napa Valley in Winkler Region III (1,670-1,940 GDD) to Region IV (1,940-2,220 GDD), confirming its warm viticultural climate suited to the full range of premium V. vinifera wine cultivars from Chardonnay and Pinot Noir (on cooler Carneros sites with 1,300-1,500 GDD) through Cabernet Sauvignon and Merlot (on the Oakville and Rutherford benches with 1,700-1,900 GDD) to Zinfandel and Petite Sirah (on the warmest valley floor and eastern hillside sites with 1,900-2,100 GDD). The GDD base 10°C value for Napa Valley (1,850-2,050) can be directly compared with Bordeaux (1,400-1,600 GDD, cooler by approximately 400 GDD), Barossa Valley (1,800-2,100 GDD, similar thermal resource), and the Douro Valley (2,200-2,600 GDD, substantially warmer), demonstrating the international comparability of the index.
A growing degree-day component that records the cumulative heat accumulation calculated using a base temperature of 10°C — the approximate threshold below which Vitis vinifera developmental processes cease — summed over the conventional growing season period (April 1 to October 31 in the northern hemisphere, October 1 to April 30 in the southern hemisphere), using the simple daily calculation method of GDD = max(0, Tmean - 10), where Tmean is the daily mean temperature calculated as (Tmax + Tmin)/2, providing the most widely used and internationally comparable thermal index for viticultural climate characterization.
GDD_Base_10 documents the simplest and most universally applied heat accumulation index, using the 10°C base temperature that approximates the biological zero for V. vinifera development. The 10°C base temperature was established empirically by Amerine and Winkler's observations that grapevine growth cessation occurs at approximately 10°C and that the cumulative temperature above this threshold correlates well with cultivar maturity timing and grape composition across the California viticultural landscape — this base temperature has been adopted internationally as the standard for viticultural GDD calculations despite evidence that the actual base temperature varies between cultivars (ranging from approximately 8°C for early-phenology cultivars to 12°C for late-phenology cultivars) and between developmental stages (lower base temperature for vegetative growth, higher for reproductive development): the standardization at 10°C sacrifices precision for comparability, enabling direct comparison of thermal resources across regions worldwide using a consistent methodology. The April 1 - October 31 calculation period (northern hemisphere) is a convention that approximates the growing season for most viticultural regions but may not capture the full growing season in very warm climates (where vine growth begins in March and continues into November) or may overestimate the effective growing season in very cool climates (where vine growth does not begin until late April and ceases in September) — the Winkler system uses this fixed-period convention, while other systems use phenologically defined or temperature-threshold-defined periods.
10°C threshold heat units
Gradtage Basis 10°C
Standard-Gradtag-Berechnung
Wärmesumme ab 10°C
base-10 degree-day accumulation
standard GDD calculation
Winkler AJ et al. (1974) General Viticulture, University of California Press Amerine MA and Winkler AJ (1944) Composition and quality of musts and wines of California grapes, Hilgardia 15: 493-675 Jones GV et al. (2005) Climate change and global wine quality, Climatic Change 73: 319-343
GDD Base 10
The Winkler Index for the major Bordeaux appellations documents the thermal differentiation within this celebrated wine region — the coolest areas (Pessac-Léognan, northern Médoc) accumulate approximately 1,350-1,450 GDD, placing them at the warm end of Winkler Region I the mid-range appellations (Saint-Julien, Pauillac, Saint-Estèphe) accumulate 1,400-1,550 GDD, squarely within Region II and the warmest areas (Saint-Émilion plateau, Pomerol) accumulate 1,500-1,600 GDD, approaching the Region II-III boundary. This internal thermal gradient explains the cultivar distribution across Bordeaux — Merlot (earlier-ripening, lower heat requirement) dominates the cooler Left Bank appellations' second variety position and the warmer Right Bank appellations' primary variety position, while Cabernet Sauvignon (later-ripening, higher heat requirement) achieves its fullest expression in the warmest Left Bank appellations (Pauillac, Saint-Julien) where the additional 100-200 GDD compared to cooler sites enables consistent phenolic maturity.
A growing degree-day component that records the Winkler Index — the specific implementation of GDD base 10°C developed by Amerine and Winkler (1944) at the University of California Davis using the fixed April 1 to October 31 accumulation period and the classification into five viticultural Regions (Region I: 2,220 GDD) — providing the historically most influential viticultural climate classification that has guided cultivar selection and regional viticultural planning in California and globally since its publication.
GDD_Winkler documents the specific implementation of the degree-day concept that became the founding framework of modern viticultural climatology. Amerine and Winkler's 1944 Hilgardia publication established the five-Region classification based on their analysis of grape composition and wine quality across California's diverse viticultural landscape — Region I (the coolest, typified by Napa-Carneros and Santa Cruz Mountains) was recommended for premium table wines from early-ripening cultivars, Region II (typified by Napa-St. Helena and Sonoma Valley) for premium table wines from the full range of cultivars, Region III (typified by Livermore and northern San Joaquin) for standard table wines, Region IV (typified by Lodi and Sacramento) for standard wines and dessert wines, and Region V (the hottest, typified by Fresno and Bakersfield) for dessert wines and raisin production: this classification has been refined and challenged over eight decades but remains the most widely recognized viticultural climate framework worldwide. The limitations of the Winkler system have been extensively documented — the fixed April-October period does not accommodate southern hemisphere seasons (requiring the October-April substitution), the lack of an upper temperature cutoff means that extreme heat days (>35°C, which suppress rather than promote vine development) contribute the same GDD as moderately warm days, and the single-variable thermal summation ignores radiation, moisture, and diurnal amplitude effects that independently influence wine quality — leading to the development of improved indices (Huglin, BEDD, MCC) that address specific limitations while building on the fundamental degree-day principle.
Amerine-Winkler classification
UC Davis degree-day regions
Winkler heat summation index
Winkler-Index
Winkler-Regionen
Winkler-Wärmesumme
Amerine MA and Winkler AJ (1944) Composition and quality of musts and wines of California grapes, Hilgardia 15: 493-675 Winkler AJ et al. (1974) General Viticulture, University of California Press Jones GV et al. (2005) Climate change and global wine quality, Climatic Change 73: 319-343
GDD Winkler
Riesling clone 239 Gm bears a Geisenheim_Clone_Role as perhaps the world's most celebrated quality Riesling selection, prized for aromatic complexity, mineral expression, and elegant wine character—adopted for premium production in Germany, Alsace, Austria, Australia, and New Zealand. Riesling clone 198 Gm serves in a Geisenheim_Clone_Role for reduced botrytis susceptibility combined with quality character, the loosened cluster architecture addressing one of Riesling's viticultural challenges while maintaining wine quality. Spätburgunder (Pinot Noir) clone 20 Gm bears a Geisenheim_Clone_Role as a quality-focused Pinot selection for German conditions, though French Dijon clones have increasingly supplemented German Spätburgunder selections internationally.
A German clone role that inheres specifically in clones selected and registered by Hochschule Geisenheim University (formerly Forschungsanstalt Geisenheim), Germany's premier viticultural research institution located in the Rheingau region, with selections designated by the "Gm" suffix (e.g., Riesling 198 Gm, Riesling 239 Gm). Geisenheim clones reflect the institution's 150+ year history of viticultural research (founded 1872), with clone selection programs emphasizing wine quality, aromatic intensity, and site expression—characteristics valued in premium German wine production. Geisenheim's location in the Rheingau (one of Germany's finest Riesling regions) has focused selection efforts particularly on Riesling, producing numerous registered clones that have become international benchmarks for quality-oriented cool-climate viticulture. Geisenheim maintains extensive clone collections (Klonarchiv), conducts multi-decade evaluation trials, and provides certified mother plant material for German nursery production. The institution's global reputation in viticultural education extends its clone influence through generations of trained winemakers who adopt Geisenheim selections worldwide.
Geisenheim_Clone_Role identifies the specific institutional origin within the German system, carrying quality associations reflecting Geisenheim's prestige in international viticulture. The institution's Rheingau location embedded its programs in Germany's premium Riesling culture, orienting selection toward quality, site expression, and characteristics valued for fine wine production rather than purely productivity. Geisenheim's educational mission has amplified clone influence students trained at Geisenheim carry familiarity with Gm selections to positions worldwide, facilitating adoption of Geisenheim material in diverse wine regions. The clone 239 Gm has achieved iconic status for quality Riesling, commanding specification in vineyard planning and premium positioning analogous to the Dijon clones for Pinot Noir—demonstrating how institutional reputation combines with documented performance to create market demand for specific selections.
Geisenheim clone
Geisenheim selection
Geisenheim-Klon
Geisenheimer Selektion
Gm clone
Gm-Klon
Rheingau clone
Rheingau-Klon
Hochschule Geisenheim (2020) Klonenzüchtung und Rebenveredlung Becker H (1985) German viticulture and breeding, American Journal of Enology and Viticulture 36: 58-63 Bundessortenamt (2020) Beschreibende Sortenliste Reben Regner F (2007) Genetic variation in grapevine clones, Blackwell
Geisenheim Clone Role
The genetic information compiled for the reference cultivar Pinot Noir (clone PN40024) encompasses the complete nuclear genome sequence (487 Mb across 19 chromosomes, published by Jaillon et al. 2007 as the first fruit crop genome), 29,971 predicted protein-coding genes, SSR marker profiles at 9 standard OIV-recommended microsatellite loci (VVS2, VVMD5, VVMD7, VVMD25, VVMD27, VVMD28, VVMD32, VrZAG62, VrZAG79) providing the genotype fingerprint registered in the VIVC database, chloroplast genome sequence (160,928 bp, haplotype C common to most Western European cultivars), parentage determination by SSR analysis confirming Pinot as a parent of Chardonnay, Gamay, Aligoté, Auxerrois, and Melon (Bowers et al. 1999), and QTL associations linking specific genomic regions to disease resistance, phenology, and berry composition traits.
A grapevine information entity that documents the heritable molecular data characterizing Vitis vinifera genotypes, encompassing DNA sequence data, molecular marker profiles, genetic map positions, quantitative trait loci, gene expression patterns, and epigenetic modifications that collectively define the genomic identity, phylogenetic relationships, trait associations, and breeding potential of individual grapevine cultivars, clones, and populations.
Genetic_Information is the umbrella class for all molecular-level heritable data in the VVO, positioned under Grapevine_Information_Entity because genetic data constitutes recorded information about the organism rather than a physical component of the organism itself (the physical DNA molecule is a material entity the sequence information it encodes is an information entity). The genomic era in grapevine research began with the publication of the Pinot Noir PN40024 reference genome (Jaillon et al. 2007, French-Italian Public Consortium) and the parallel Pinot Noir ENTAV115 genome (Velasco et al. 2007), providing the foundation for subsequent resequencing of hundreds of cultivars, genome-wide association studies (GWAS), and marker-assisted selection in breeding programs: the current Vitis genomic landscape includes reference-quality genome assemblies for multiple cultivars (Cabernet Sauvignon, Chardonnay, Zinfandel, Carménère, Riesling), pan-genome resources capturing structural variation across the genus, and extensive transcriptomic and epigenomic datasets. Subclasses are organized by data type, beginning with DNA sequence data.
genetic characterization data
genetische Information
genomic data
genomische Daten
grapevine molecular information
molekulargenetische Charakterisierung
Jaillon O et al. (2007) The grapevine genome sequence suggests ancestral hexaploidization in major angiosperm phyla, Nature 449: 463-467 Velasco R et al. (2007) A high quality draft consensus sequence of the genome of a heterozygous grapevine variety, PLoS ONE 2: e1326 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519
Genetic Information
The genetic marker data compiled for the OIV-recommended grapevine identification panel encompasses 9 microsatellite markers (VVS2, VVMD5, VVMD7, VVMD25, VVMD27, VVMD28, VVMD32, VrZAG62, VrZAG79) characterized across more than 5,000 cultivars, with each marker documented by its chromosomal location (mapped to specific positions on 8 of the 19 Vitis chromosomes), repeat motif structure, primer sequences, allele size range (spanning 30-80 bp per locus), number of detected alleles (8-25 per locus in V. vinifera), observed and expected heterozygosity (0.72-0.89), probability of identity (0.03-0.12 per locus, cumulative <10⁻⁸ across 9 loci), and null allele frequency estimates — this comprehensive marker characterization enables informed selection of marker subsets for specific applications ranging from routine cultivar verification (minimum 6 loci) through parentage analysis (9+ loci) to population structure assessment (20+ loci recommended).
A genetic information entity that records the characterization of specific DNA polymorphisms used as landmarks for genome navigation, cultivar identification, genetic mapping, and marker-assisted selection in Vitis vinifera, encompassing the marker identity, genomic position, polymorphism type, allele frequency distribution, and informativeness metrics that collectively define the utility of each marker system for specific applications in grapevine genetics and breeding.
Genetic_Marker_Data is the class for documented properties of DNA markers as tools, distinguished from Genotype_Data (which records the allelic state of a specific accession at marker loci) by its focus on the marker itself rather than on the individual genotyped. The development of genetic markers for grapevine progressed through four technological generations — isozymes (1980s, 5-10 polymorphic loci, limited discrimination), RFLPs (late 1980s, technically demanding, largely superseded), SSRs/microsatellites (1990s-present, the current standard for cultivar identification with >300 published Vitis SSR markers), and SNPs (2010s-present, millions of positions available, dominant for research applications): each generation increased marker density, throughput, and automation potential while building on the foundational genetic map framework established by earlier technologies. Subclasses organize marker data by marker type, beginning with microsatellite markers that remain the backbone of applied grapevine identification.
DNA marker characterization
DNA-Marker-Spezifikation
genetic marker specification
genetische Marker-Daten
molecular marker record
molekulare Marker-Charakterisierung
This P et al. (2004) Development of a standard set of microsatellite reference alleles for identification of grape cultivars, Theoretical and Applied Genetics 109: 1448-1458 Adam-Blondon AF et al. (2004) Genetic mapping of grapevine, in Bentham Science Publishers, Functional and Integrative Genomics 4: 196-211 Sefc KM et al. (2001) Microsatellite markers for grapevine, in Roubelakis-Angelakis KA (ed) Molecular Biology and Biotechnology of the Grapevine, Kluwer pp: 433-463
Genetic Marker Data
Pinot Noir accession Vas 3082Mtp2 at INRAE Domaine de Vassal bearing a genetic reference role provides the authenticated DNA source for the international SSR reference profile (VVS2: 131-151, VVMD5: 236-238, VVMD7: 239-243, VVMD25: 239-249, VVMD27: 179-189, VVMD28: 233-259, VVMD32: 252-272, VrZAG62: 187-193, VrZAG79: 245-247) deposited in the VIVC database and used by 80+ genotyping laboratories worldwide for cultivar identity verification, with whole-genome sequence data (PN40024 reference genome, 475 Mb assembled) serving as the primary reference assembly for Vitis vinifera genomic research.
A research and education role that inheres in a grapevine maintained as an authenticated source of DNA for molecular identification, parentage analysis, genetic mapping, and genomic research, with its genotype profile deposited in reference databases to enable standardized cultivar identification and inter-laboratory calibration of molecular marker systems.
Genetic_Reference_Role addresses the molecular dimension of cultivar authentication that complements classical ampelographic identification. The role became critical following the development of microsatellite (SSR) marker systems in the 1990s-2000s that enabled unambiguous cultivar identification independent of environmental and developmental variation affecting morphological descriptors: the establishment of standardized SSR reference profiles and inter-laboratory calibration protocols (notably the GENRES CT96 No. 81 EU project) transformed grapevine identification from a subjective morphological art to a reproducible molecular science. The role requires rigorous chain-of-custody documentation linking the reference vine to its deposited genetic profile.
DNA reference role
DNA-Referenz-Rolle
genetic standard role
genetische Referenz-Rolle
molecular identification reference role
molekulare Identifikations-Rolle
This P et al. (2004) Development of a standard set of microsatellite reference alleles for identification of grape cultivars, Theoretical and Applied Genetics 109: 1448-1458 Jaillon O et al. (2007) The grapevine genome sequence suggests ancestral hexaploidization, Nature 449: 463-467 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157
Genetic Reference Role
Vitis rotundifolia (Muscadine) accessions in the USDA National Clonal Germplasm Repository bear Genetic_Resource_Roles as sources of powdery mildew immunity (Run1, Ren1 loci) unavailable in V. vinifera, accessed by breeding programs worldwide for resistance introgression. Ancient Georgian varieties preserved in the Kakheti region serve in Genetic_Resource_Roles as representatives of 8,000 years of viticultural selection, potentially carrying unique alleles for adaptation, quality, and disease tolerance. A heritage Zinfandel selection maintained in the UC Davis germplasm collection bears a Genetic_Resource_Role as a representative of 19th-century California viticulture, preserved for historical significance and potential breeding value.
A breeding role that inheres in a grapevine genotype by virtue of its value as a source of genetic diversity for breeding programs, variety conservation, or scientific research—encompassing wild species, landraces, historic cultivars, and germplasm collection accessions maintained for their potential contribution to current or future genetic improvement. Genetic resources represent the raw material of plant breeding, providing alleles for disease resistance, stress tolerance, quality attributes, and adaptation that may be absent from elite cultivated material but essential for addressing emerging challenges (new pathogens, climate change, market evolution). Grapevine genetic resources span enormous diversity—approximately 60-70 Vitis species distributed across North America, Asia, and Europe, plus thousands of cultivated varieties representing centuries of human selection, plus wild populations and landraces preserving locally adapted genetic variation. Conservation of genetic resources balances ex situ preservation (germplasm collections, repositories) with in situ maintenance (wild populations, traditional vineyards), recognizing that both approaches have complementary strengths and limitations. Access to genetic resources is increasingly governed by international frameworks (Nagoya Protocol, International Treaty on Plant Genetic Resources) balancing conservation, equitable benefit sharing, and breeding program access.
Genetic_Resource_Role is subclassed by resource category—Wild_Vitis for uncultivated species diversity, Landrace for traditional locally-adapted populations, and Germplasm_Collection_Member for formally accessioned and maintained materials. The genetic resource concept emphasizes potential future value alongside current utility material may be conserved without immediate breeding application, preserved against future needs that cannot be anticipated. Climate change has intensified genetic resource importance, with traits for heat tolerance, drought resistance, and phenological flexibility becoming priorities that may require accessing variation not present in current elite cultivars. The tension between genetic resource conservation (maintaining diversity) and variety improvement (concentrating favorable alleles) reflects the broader challenge of balancing short-term breeding gains against long-term adaptive capacity.
Genressource
Keimplasma
breeding resource
genetic diversity source
genetic resource
genetische Diversitätsquelle
genetische Ressource
germplasm
This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Riaz S et al. (2019) Genetic resources for grape improvement, in Cantu D, Walker MA (eds) The Grape Genome, Springer pp.: 25-51 FAO (2010) Second Report on the State of the World's Plant Genetic Resources for Food and Agriculture Lacombe T et al. (2011) Grapevine European catalogue and worldwide genetic diversity, Vitis 50: 65-68
Genetic Resource Role
Pinot Noir Dijon clones (113, 114, 115, 667, 777, etc.) bear a Genetic_Selection_Clone_Role from ENTAV-INRA programs prioritizing wine quality characteristics, with selections made based on replicated field trials and winemaking evaluations identifying genotypes producing superior wines in Burgundian conditions. Chardonnay clone 76 serves in a Genetic_Selection_Clone_Role, selected specifically for small-cluster character and concentrated must composition—genetic evaluation identifying this phenotype among Burgundy candidate genotypes. Sangiovese clone BBS11 bears a Genetic_Selection_Clone_Role from Italian selection programs evaluating genotypes for Brunello di Montalcino production quality, with extended winemaking trials demonstrating superior aging potential compared to other candidates.
A selection status clone role that inheres in a grapevine clone whose development prioritized agronomic and enological performance characteristics, selecting among genotypes based on field performance, fruit composition, and wine quality rather than primarily health status considerations. Genetic selection focuses on identifying genotypes with superior production characteristics—yield level and stability, cluster architecture, disease resistance, fruit composition (sugar, acid, phenolics), and wine quality attributes—through systematic evaluation programs comparing candidate genotypes under replicated trial conditions. The "genetic" designation distinguishes this selection emphasis from sanitary selection (prioritizing pathogen elimination) though contemporary programs typically integrate both dimensions. Genetic selection programs evaluate candidate genotypes through 10-20 year cycles including establishment, multi-year field data collection, wine production and evaluation, and eventual registration of superior performers. The approach has produced the diversity of registered clones with documented performance differences—enabling growers to select clones matching specific production objectives, from high-yield bulk wine production through premium low-yield quality-focused clones.
Genetic_Selection_Clone_Role captures the performance-focused dimension of clone development where agronomic and enological characteristics drive selection decisions. The term "genetic" reflects selection among genotypes (genetic variants within the cultivar) rather than genetic engineering or marker-assisted breeding (though molecular markers increasingly support selection by enabling genotype identification and diversity assessment). Genetic selection has produced the documented performance differences among registered clones—yield variations of 30-50%, ripening date differences of 5-15 days, and wine style differences readily detected in blind tasting. The diversity of registered clones enables site-specific and objective-specific clone selection, from high-productivity clones for value-tier production through low-yield quality-focused clones for premium wines. Contemporary genetic selection integrates molecular tools for genotype identification, pathogen testing, and increasingly for trait-associated marker selection, though phenotypic evaluation through field and wine trials remains the foundation of clone evaluation.
Leistungsselektion
Qualitätsselektion
agronomic selection
agronomische Selektion
genetic selection
genetische Selektion
performance selection
quality selection
Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 ENTAV-INRA (2007) Catalogue des Variétés et Clones de Vigne This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340
Genetic Selection Clone Role
A Concord vine in New York's Finger Lakes trained to GDC bears a Geneva_Double_Curtain_Role with two parallel cordons at 170 cm height spaced 120 cm apart, the pendant shoot growth managing the variety's notorious vigor while improving light exposure for the juice-quality fruit—yields of 15-20 tonnes/hectare achievable with acceptable quality. A Niagara table grape vine serves in a Geneva_Double_Curtain_Role, the divided high-cordon system providing the vigor management essential for this productive variety while enabling mechanical harvesting. A Chambourcin wine grape vine in Virginia bears a Geneva_Double_Curtain_Role, the system adapted from juice grape origins to manage this vigorous French-American hybrid in humid eastern conditions.
A training system role that inheres in a grapevine trained to the Geneva Double Curtain (GDC) system, a high-cordon divided canopy architecture with two parallel cordons at elevated height (150-180 cm), each bearing downward-growing shoots that create pendant curtains separated by the inter-cordon space. GDC was developed by Nelson Shaulis at Cornell University's Geneva Experiment Station in the 1960s specifically for managing the excessive vigor of Concord and other American hybrid varieties in New York's Finger Lakes region, where conventional training produced dense, shaded canopies with poor fruit quality and disease pressure. The system positions two cordons approximately 120 cm apart at elevated height, with shoots growing downward (pendant) from spur positions on each cordon, disrupting apical dominance and naturally limiting shoot elongation. GDC dramatically improved light interception and fruit exposure for vigorous varieties, increasing yields while improving fruit quality compared to single-curtain systems—an achievement that stimulated subsequent research into canopy management principles applicable across viticulture.
Geneva_Double_Curtain_Role captures Shaulis's pioneering divided canopy system that established the principles underlying subsequent canopy management development. The system's pendant (downward) shoot orientation distinguishes it from lyre's upward-growing curtains, with pendant growth naturally limiting shoot vigor through disrupted apical dominance. GDC's success for vigorous American varieties demonstrated that training system selection profoundly affects fruit quality—a principle now fundamental to viticulture but revolutionary when Shaulis's work was published. The system remains important for juice and jelly grape production in eastern North America, though wine grape applications are limited by the pendant canopy's association with volume rather than premium production. GDC's historical significance exceeds its current acreage the research it stimulated informed Smart's canopy management principles, Carbonneau's lyre development, and the broader understanding of light interception, canopy microclimate, and fruit quality relationships.
Doppelvorhang-System
GDC
GDC-Erziehung
Geneva Double Curtain
Genfer Doppelvorhang
Shaulis system
double curtain
Shaulis N et al. (1966) Response of Concord grapes to light exposure and Geneva Double Curtain training, Proceedings of the American Society for Horticultural Science 89: 268-280 Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Reynolds AG, Vanden Heuvel JE (2009) Training systems, American Journal of Enology and Viticulture 60: 251-268
Geneva Double Curtain Role
The genotype data for Cabernet Sauvignon compiled in the VIVC database and cross-referenced with the UC Davis Foundation Plant Services collection comprises SSR genotypes at 9 standard microsatellite loci (providing the primary cultivar fingerprint), SNP genotypes at 10,000+ positions from the Vitis18K or Vitis50K arrays (providing high-resolution genomic characterization for GWAS and genomic selection), and targeted gene-level genotypes at functional loci including VvMYBA1 (functional allele, encoding the anthocyanin transcription factor responsible for red berry color), VvDXS (wild-type allele, associated with non-muscat aroma profile), and Rpv/Ren resistance loci (absent, confirming full susceptibility to downy and powdery mildew) — collectively these genotype data layers provide unambiguous molecular identification, trait prediction, and breeding value assessment for the cultivar.
A genetic information entity that records the specific allelic composition of a Vitis vinifera individual at one or more polymorphic loci, providing the molecular identity data used for cultivar authentication, parentage verification, clonal distinction, and population genetic analysis through the documentation of which genetic variants are present in a specific grapevine accession at defined marker positions in the genome.
Genotype_Data is the class for recorded allelic states at specific genomic positions, distinguished from DNA_Sequence (which records the nucleotide succession itself) by its focus on the identity of variant alleles at polymorphic loci rather than the continuous nucleotide string. The transition from morphological to molecular genotyping transformed grapevine cultivar identification from an expert-dependent, phenotype-based practice subject to environmental variation and observer subjectivity into a reproducible, laboratory-based system capable of unambiguous identification from any tissue sample at any developmental stage: this transformation, beginning with isozyme markers in the 1980s and accelerating through SSR (microsatellite) markers in the 1990s and SNP arrays from the 2010s onward, has resolved thousands of synonymy and homonymy cases that confounded ampelographic identification for centuries. Subclasses capture the dominant marker systems used in grapevine genotyping, beginning with SSR markers that remain the international standard for cultivar identification.
Allelprofil-Daten
Genotyp-Daten
allelic profile data
marker genotype information
molecular genotype record
molekulare Genotypisierung
This P et al. (2004) Development of a standard set of microsatellite reference alleles for identification of grape cultivars, Theoretical and Applied Genetics 109: 1448-1458 Laucou V et al. (2018) An extended haplotype reference panel and high-density SNP genotyping data for grapevine, Scientific Data 5: 180259 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database referenced by genetic profiles and morphology, Vitis 54: 155-157
Genotype Data
Geographic distribution of Chardonnay clone 76 (ENTAV) spans five continents with documented commercial plantings in Burgundy and Champagne (France, origin region, estimated 2500+ ha), Napa and Sonoma Valleys (California, introduced 1984, approximately 800 ha), Marlborough and Hawke's Bay (New Zealand, introduced 1991, approximately 400 ha), Yarra Valley and Adelaide Hills (Australia, introduced 1988, approximately 300 ha), and Casablanca Valley (Chile, introduced 1995, approximately 150 ha) — establishing it across a latitudinal range of 34-49°N/S and elevation range of sea level to 450 m in climates spanning cool maritime (Champagne) through warm Mediterranean (Napa). Geographic distribution of Blaufränkisch clone Bl-2 documents concentrated plantings in Mittelburgenland (Austria, primary region, approximately 450 ha), Sopron (Hungary, introduced 2005, approximately 30 ha), Moravia (Czech Republic, introduced 2008, approximately 15 ha), and trial plantings in Finger Lakes (New York, introduced 2012, <5 ha), reflecting a distribution pattern closely tied to the cultivar's Central European climatic adaptation zone.
Geographic distribution is a distribution record entity that records the spatial extent of commercial vineyard plantings established with a specific certified clonal line, including the countries, wine regions, appellations, and climatic zones where the clone has been planted, the approximate planted area or vine count in each region where data is available, the date of first introduction to each region, and the range of growing conditions (latitude, elevation, climate type) under which the clone is cultivated, characterizing the global footprint and environmental adaptation range of the clonal line.
Geographic distribution data provides empirical evidence of a clone's environmental adaptation range, complementing the controlled trial data from evaluation sites with real-world performance information across diverse terroirs. The distinction between documented plantings (verified through nursery sales records, grower surveys, or regional planting statistics) and estimated distribution (extrapolated from nursery production volumes or industry sources) should be clearly maintained, as the latter introduces substantial uncertainty. The geographic spread of a clone is influenced by multiple factors beyond agronomic merit — nursery supply logistics, regulatory import requirements, historical trade relationships between wine regions, and the promotional activities of selection programs and nursery cooperatives all shape distribution patterns. Some clones achieve global ubiquity (the ENTAV Dijon Pinot Noir series is planted on every wine-producing continent) while others remain regionally confined despite excellent performance, limited by nursery availability, lack of international promotion, or regulatory barriers to cross-border plant material movement. Geographic distribution data carries important climate adaptation information — the latitudinal, altitudinal, and climatic range across which a clone produces acceptable results defines its practical adaptation envelope, informing planting decisions for new vineyard establishments, particularly under climate change scenarios where producers seek clones with demonstrated performance across a wide environmental range. The progressive development of national and international vineyard census databases incorporating clone-level planting data is improving the accuracy of geographic distribution documentation, though comprehensive clone-level statistics remain unavailable in many regions.
Anbauverbreitung
Verbreitungsgebiet
clone distribution map
clone geographic range
geographic spread
geographische Distribution
geographische Verbreitung
planting distribution
regional distribution
regionale Verbreitung
This P et al. (2006) Historical origins and genetic diversity of wine grapes. Trends in Genetics 22(9): 511-519 Anderson K, Aryal NR (2013) Which Winegrape Varieties are Grown Where? A Global Empirical Picture, University of Adelaide Press Maul E et al. (2012) The European Vitis Database — a technical innovation through an online uploading and interactive modification system. Vitis 51(2): 79-85
Geographic Distribution
The geographic indication system governing Bordeaux wines operates across multiple hierarchical levels — from the broad regional indication Bordeaux AOC (covering approximately 111,000 hectares) through sub-regional appellations (Haut-Médoc, Graves, Saint-Émilion) to communal appellations (Pauillac, Margaux, Pomerol) and individual classified estates (Château Lafite Rothschild as Premier Grand Cru Classé 1855) — each level adding specificity to the geographic origin claim and imposing progressively stricter requirements for grape varieties, viticultural practices, yields, and winemaking methods as the geographic scope narrows.
A grapevine information entity that identifies a wine, grape product, or viticultural practice as originating from a defined geographic area where a specific quality, reputation, or characteristic is essentially attributable to that geographic origin, encompassing all forms of place-based designations used in viticulture and wine production from informal regional references through legally protected appellations and designations of origin.
Geographic_Indication is the umbrella class for all place-based designation systems in viticulture and wine production. The concept operates at the intersection of intellectual property law, trade regulation, terroir science, and cultural heritage protection: the WTO TRIPS Agreement (1994) provides the international legal framework defining geographic indications as identifiers linking product qualities to geographic origin, while the EU PDO/PGI system (Regulation 1308/2013) and analogous national systems implement this framework through specific registration, specification, and enforcement mechanisms. Subclasses distinguish between appellation designations (the naming and hierarchical classification of geographic units) and protected designations of origin (the legal registration and regulatory documentation of geographic indications).
Herkunftsbezeichnung
Ursprungsangabe
geografische Angabe
geographic designation
origin indication
place-based wine designation
WTO (1994) Agreement on Trade-Related Aspects of Intellectual Property Rights (TRIPS), Art. 22-24 EU Regulation 1308/2013 Art. 93-108 on wine designations of origin and geographical indications OIV (2015) OIV Guidelines for Geographic Indications for Wines
Geographic Indication
Geographic origin for Riesling is documented as the Rhine Valley of Germany with high confidence — molecular kinship analysis reveals maximum genetic diversity and closest parental relationships in the Upper Rhine-Alsace-Pfalz region, the earliest documented references (1435 Rheingau inventory, 1477 Alsatian document) cluster in this area, archaeobotanical evidence of Vitis vinifera cultivation in the Rhine Valley dates to Roman times (though not specifically Riesling), and the ecological niche of cool continental climate with steep river-valley mesoclimates corresponds precisely to the cultivar's known climatic preferences, collectively establishing the Middle-Upper Rhine corridor as the near-certain geographic origin. Geographic origin for Furmint is documented as the Carpathian Basin (present-day Hungary-eastern Austria-Slovakia-northern Serbia) with moderate confidence — genetic analysis reveals relationships with multiple Pannonian cultivars, the Tokaj-Hegyalja region provides the earliest documentation, but contested linguistic evidence suggests possible earlier Adriatic or Balkan provenance.
Geographic origin is an origin record entity that documents the specific geographic location or region identified as the primary center of origin, domestication, or first cultivation of a grape cultivar, integrating evidence from molecular population genetics (genetic diversity and kinship patterns indicating center of origin), archaeobotanical findings (archaeological grape seed and wood remains in stratigraphic context), archival documentation (earliest geographic references in historical texts), and ecological niche analysis (climatic and edaphic compatibility with proposed origin regions), expressed as a geographic region with associated coordinates, confidence level, and evidentiary basis, and providing the spatial anchor for the cultivar's historical narrative.
Geographic origin determination for grape cultivars involves a characteristic epistemological challenge — the distinction between the geographic location where a cultivar first arose (center of origin, often unknown or unknowable for cultivars arising from spontaneous crosses centuries ago), the location where it was first identified and propagated (center of domestication or selection), and the location where it was first documented in writing (center of earliest attestation). These three locations may coincide but frequently diverge — a cultivar that arose as a chance seedling in one vineyard may have been recognized and propagated in a neighboring region, and may appear in written records only from a third location where literate record-keeping was practiced. The molecular genetic approach to geographic origin determination relies on the principle that the center of origin of a species or cultivar typically harbors the greatest genetic diversity — applied to grape cultivars, this means analyzing microsatellite diversity across regions and identifying the population with the highest allelic richness and heterozygosity as the probable center of origin. However, for cultivars that are clonally propagated (genetically identical), this approach applies not to the cultivar itself but to its closest relatives and parental lineages. The archaeobotanical dimension has been transformed by advances in morphometric analysis of grape seeds (which can distinguish wild from domesticated Vitis vinifera and, with decreasing confidence, cultivar groups) and ancient DNA extraction from waterlogged or charred archaeological material, though the identification of specific cultivars from archaeological remains remains technically challenging and results are often contested. The political dimension of geographic origin claims should not be underestimated — cultivar origin narratives carry cultural prestige and can influence appellation regulations, marketing claims, and national identity, creating incentives for origin claims that may exceed the available evidence.
Entstehungsort
Herkunftsgebiet
Ursprungsregion
center of origin
cultivar homeland
geographic provenance
geographische Provenienz
geographischer Ursprung
origin region
place of origin
Myles S et al. (2011) Genetic structure and domestication history of the grape. Proceedings of the National Academy of Sciences 108(9): 3530-3535 Bacilieri R et al. (2013) Genetic structure in cultivated grapevines is linked to geography and human selection. BMC Plant Biology 13: 25 Terral JF et al. (2010) Evolution and history of grapevine under domestication: new morphometric perspectives to understand seed domestication syndrome and reveal origins of ancient European cultivars. Annals of Botany 105(3): 443-455 McGovern PE (2003) Ancient Wine, Princeton University Press, pp.: 40-80
Geographic Origin
Geraniol concentration in grape berries ranges from 5-30 µg/L in non-aromatic cultivars to 200-1200 µg/L in aromatic types, with sensory threshold of 20-40 µg/L for the characteristic rose-geranium aroma. Muscat of Alexandria and Muscat Ottonel show elevated geraniol (600-1200 µg/L), while Gewürztraminer typically shows moderate levels (200-500 µg/L) with lychee-rose character. VvTPS-ger (geraniol synthase) produces geraniol directly from GPP, while geraniol reductase can convert geraniol to citronellol (citrus character). Geraniol serves as the immediate precursor to nerol (geometric isomer) and can be converted to linalool through enzymatic or acid-catalyzed rearrangement. The geraniol/nerol ratio typically ranges from 3-6 in fresh grapes, with higher nerol proportions developing during wine aging. Geraniol oxidation products (rose oxide, citral) contribute additional complexity to aged aromatic wines.
A monoterpene accumulation disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate geraniol (3,7-dimethyl-2,6-octadien-1-ol) as a significant monoterpene alcohol in grape berries, with genotype-specific concentrations contributing rose-like floral aroma and serving as a biosynthetic precursor for other monoterpenes including citronellol, nerol, and linalool through enzymatic interconversion.
Geraniol occupies a central position in monoterpene metabolism, serving both as an aroma-active compound and as a precursor for structural diversity through enzymatic modification. The rose-like character of geraniol complements the lily/lavender notes of linalool, with the relative proportions contributing to cultivar-specific aromatic profiles. Geraniol shows somewhat greater stability than linalool during winemaking but undergoes oxidation and esterification reactions that modulate its sensory contribution during aging.
Geraniol-Disposition
Geraniolakkumulation
Geraniolgehalt
geraniol content disposition
geraniol synthesis capacity
rose terpene trait
Luan F et al. (2005) Effects of different substrates on monoterpene production in Vitis vinifera cv. Gewürztraminer berry exocarp. Journal of Agricultural and Food Chemistry 53:4520-4525 Ilc T et al. (2016) A grapevine cytochrome P450 generates the precursor of wine lactone, a key odorant in wine. New Phytologist 211:864-874
Geraniol Accumulation Disposition
Geraniol content quality in Gewürztraminer from Alto Adige reaches 400-700 μg/L, substantially higher relative to linalool than in Muscat varieties, producing the variety's signature rose-petal, lychee character distinct from the citrus-floral Muscat profile the geraniol-dominant terpene profile combines with elevated terpenoid acids and bound glycosides to create the intensely perfumed, exotic aromatic character that makes Gewürztraminer among the most instantly recognizable and aromatically distinctive wine varieties, with South Tyrolean examples considered benchmarks for aromatic intensity and varietal expression.
Geraniol_Content_Quality represents the chemical quality characterizing the concentration of geraniol (3,7-dimethyl-trans-2,6-octadien-1-ol) in grape berries and wine, capturing the acyclic monoterpene alcohol responsible for distinctive rose-like floral aroma notes in aromatic cultivars. Geraniol, the trans-isomer of the geranyl alcohol structure, is distinguished from its cis-isomer nerol by the configuration of the 2,3 double bond, with both isomers contributing rose character but geraniol typically present at higher concentrations in grapes. As the second most abundant monoterpene alcohol in many aromatic varieties after linalool, geraniol typically comprises 15-30% of total free monoterpenes, with concentrations ranging from 5-50 μg/L in neutral varieties through 100-300 μg/L in semi-aromatic cultivars to 300-800 μg/L in intensely aromatic Muscat and Gewürztraminer. The sensory threshold of geraniol in wine is approximately 20-40 μg/L, ensuring perceptible contribution in aromatic varieties. Geraniol is biosynthetically related to linalool through geranyl pyrophosphate, with the relative proportions reflecting cultivar-specific differences in terpene synthase activities. Gewürztraminer characteristically shows elevated geraniol relative to linalool compared to Muscat varieties, contributing to its distinctive rose-petal rather than citrus-floral character. Geraniol accumulates in berry skins as both free alcohol and glycosylated conjugates, with the bound-to-free ratio affecting aroma potential during aging. During wine maturation, geraniol can undergo reduction to citronellol or acid-catalyzed cyclization, participating in the complex terpene transformations that contribute to bouquet development in aromatic white wines.
Geraniol_Content_Quality addresses a key monoterpene whose relative abundance versus linalool contributes to cultivar-specific aromatic signatures. The trans configuration of geraniol distinguishes it from nerol (cis-isomer) both contribute rose notes but at different intensities and with subtle character differences. Geraniol serves as precursor for citronellol through enzymatic or chemical reduction, linking these terpene compounds metabolically. The geraniol/linalool ratio provides cultivar fingerprinting information, with Gewürztraminer showing characteristically higher ratios than Muscat varieties.
Geraniol-Konzentration
Geranylverbindungsgehalt
Rosenduft-Terpenalkoholgehalt
geraniol level
geranyl compound content
rose terpene alcohol quality
trans-3,7-dimethyl-2,6-octadien-1-ol concentration
Marais J (1983) Terpenes in the Aroma of Grapes and Wines: A Review. South African Journal of Enology and Viticulture 4(2): 49-58 Guth H (1997) Quantitation and Sensory Studies of Character Impact Odorants of Different White Wine Varieties. Journal of Agricultural and Food Chemistry 45(8): 3027-3032
Geraniol Content Quality
Riesling clone 239 Gm bears a German_Clone_Role as a Geisenheim selection prized for aromatic intensity and site expressivity, widely adopted for premium German and cool-climate international Riesling production. Spätburgunder (Pinot Noir) clone 1 Fr serves in a German_Clone_Role as the Freiburg selection historically dominant in Baden, selected for German growing conditions before French Dijon clones expanded internationally. Müller-Thurgau clone 1 We bears a German_Clone_Role from Weinsberg, supporting the variety's continued importance in German bulk and regional wine production despite declining prestige.
A regional clone role that inheres in a grapevine clone registered within the German national clone certification system, selected and evaluated by authorized research institutions including Geisenheim University (Hochschule Geisenheim), the Julius Kühn-Institut (JKI, formerly Bundesanstalt für Züchtungsforschung), the Staatliches Weinbauinstitut Freiburg (WBI), and the Staatliche Lehr- und Versuchsanstalt Weinsberg (LVWO). German clones are identified by variety name, numeric code, and institutional suffix indicating selecting institution (Gm for Geisenheim, Fr for Freiburg, We for Weinsberg, N for Neustadt, Wü for Würzburg—e.g., Riesling 239 Gm, Spätburgunder 1 Fr). The German clone system reflects the country's cool-climate viticulture focus, with particular strength in Riesling, Silvaner, and Spätburgunder (Pinot Noir) selections optimized for northern European growing conditions. German clone evaluation emphasizes adaptation to marginal climates, winter hardiness, and disease resistance alongside yield and quality parameters. The multi-institutional structure creates healthy competition and diverse selection approaches, with different institutions developing distinct clone portfolios.
German_Clone_Role captures the distinctive German institutional structure where multiple competing research stations conduct clone selection within a coordinated national framework. The institutional suffix system enables immediate identification of selecting institution—information valued by German producers who may prefer material from their regional station or from institutions with specific selection emphasis. German clone programs pioneered cold-climate variety selection, developing Riesling clones optimized for the marginal growing conditions of Mosel, Rheingau, and Pfalz that have proven valuable for cool-climate production globally (Alsace, Austria, New Zealand, Oregon). The German system's fragmentation (multiple institutions with separate programs) contrasts with the centralized French ENTAV-INRA approach, creating diverse selection philosophies but also nomenclature complexity.
Bundessortenamt clone
Bundessortenklon
German clone
German institutional clone
German registered clone
deutsch registrierter Klon
deutscher Klon
Bundessortenamt (2020) Beschreibende Sortenliste Reben Hochschule Geisenheim (2020) Klonenzüchtung und Rebenveredlung Regner F (2007) Genetic variation in grapevine clones, Blackwell Becker H (1985) German viticulture and breeding, American Journal of Enology and Viticulture 36: 58-63
German Clone Role
Merlot in Bordeaux shows germination pore activation during early-mid June Year N+1 at BBCH 60-65, with one of three colporate apertures per grain undergoing wall loosening and cytoplasmic polarization over 25-55 minutes following hydration, enzymatic wall modification at selected pore creating breakthrough site, turgor pressure 0.8-1.3 MPa accumulating, under oceanic conditions of 21-26°C temperature, adequate boron nutrition 12-18 ppm supporting normal wall synthesis, typical of this mid-season cultivar enabling tube emergence 2-4 hours post-adhesion
Germination pore activation represents the aperture preparation process where one of three colporate pores in hydrated pollen grain undergoes structural modification, wall loosening, and functional preparation enabling pollen tube emergence through selected aperture establishing growth pathway from grain. This critical pore preparation process occurs during pollen germination in early-mid June in temperate regions Year N+1, typically proceeding over 15-60 minutes following pollen hydration as germination approaches, corresponding to BBCH 60-65 flowering stages. Activation mechanism involves aperture selection from three available colporate pores distributed around grain equator, localized cell wall modification at selected pore through enzymatic loosening, intine layer preparation for tube extension with structural changes enabling growth, and turgor pressure accumulation creating mechanical force driving tube emergence. Pollen grain aperture structure shows three colporate apertures in typical grapevine pollen grain providing potential germination sites, each aperture representing thinned region in exine outer wall with modified intine inner wall, and aperture positioning distributed approximately equidistantly around grain equator creating multiple germination options. Aperture selection mechanism shows typically single pore activating for tube emergence though occasionally multiple tubes may initiate from different pores, selection possibly influenced by contact geometry with stigma surface, orientation at adhesion affecting accessibility, intrinsic polarity possibly favoring particular pore, or stochastic factors with first-activated pore proceeding while others suppress, though mechanisms poorly understood. Localized wall loosening at selected aperture involves enzymatic degradation of cell wall polysaccharides with pectinase and cellulase activity concentrated at pore region, wall component dissolution creating flexibility enabling tube breakthrough, and controlled degradation maintaining structural integrity while allowing expansion. Cell wall modification shows intine inner layer at aperture undergoing structural changes, pectin and hemicellulose components partially degrading creating loosened matrix, cellulose microfibrils reorganizing enabling expansion, and overall wall modification transitioning from rigid barrier to flexible growth-permissive structure. Enzymatic activity driving activation includes pectinase enzymes degrading pectin polysaccharides in middle lamella and wall matrix, cellulase activity breaking down cellulose components, polygalacturonase specifically targeting pectin creating wall loosening, and coordinated enzyme action producing controlled wall modification enabling tube emergence. Cytoplasmic polarization at aperture shows vegetative cell cytoplasm concentrating at selected pore region, organelles including mitochondria, rough ER, and Golgi apparatus migrating toward aperture supporting active secretion and metabolism, secretory vesicles accumulating delivering wall precursors for tube formation, and overall cellular polarization establishing growth axis. Turgor pressure generation shows water uptake creating osmotic swelling, cell membrane pushing against wall creating mechanical pressure, concentrated pressure at weakened aperture region creating driving force for tube emergence, and regulated turgor typically 0.5-1.5 MPa sufficient for breakthrough without grain rupture. Intine preparation at aperture involves inner wall layer at pore undergoing reorganization, structural proteins and polysaccharides modifying creating growth-competent state, wall synthesis machinery activating preparing for tube wall construction, and transition from static grain wall to dynamic growing tube wall. Exine outer wall role shows rigid sculptured outer layer maintaining structural integrity preventing grain collapse, aperture regions having naturally thinned exine providing predetermined emergence sites, and exine remaining intact during tube emergence with tube extending through aperture rather than breaching solid wall. Pore activation timing shows process initiating 30-90 minutes after hydration as metabolic activation proceeds, progressing over 15-60 minutes as wall loosening and polarization develop, and completing immediately before tube emergence establishing ready state for breakthrough. Activation coordination with overall germination shows pore activation proceeding concurrently with metabolic activation and reserve mobilization, timing synchronized so tube emergence occurs when cellular resources and organization prepared, and integrated developmental program coordinating multiple preparation processes. Multiple pore potential shows grain capable of activating any of three apertures, first-activated pore typically proceeding to emergence while others remain inactive, occasional simultaneous activation producing multiple tubes from single grain though usually only one succeeds in fertilization, and selection mechanism ensuring directed coordinated growth. Calcium signaling in activation possibly involves calcium gradients establishing polarity, localized calcium accumulation at prospective germination site, and calcium-dependent enzyme activation though specific roles in grapevine poorly characterized. Cytoskeleton role in activation includes microtubules and actin filaments reorganizing establishing polarity, cytoskeleton directing organelle positioning and vesicle trafficking to aperture, and structural framework supporting polarized cellular organization. Environmental effects on pore activation include temperature affecting enzymatic activity and metabolic processes with optimal 22-28°C supporting efficient activation, boron essential for proper wall modification and synthesis with deficiency preventing normal activation and tube formation, and adequate hydration providing turgor for mechanical breakthrough. Temperature effects show warm conditions 25-28°C promoting rapid activation within 15-30 minutes, cool conditions 18-22°C slowing activation requiring 45-90 minutes, and temperature sensitivity reflecting enzymatic basis of wall modification processes. Boron critical role shows absolute requirement for pore activation with boron essential for pectin cross-linking and wall structure, deficiency causing abnormal wall modification preventing proper tube emergence or causing tube bursting, and optimal boron concentration 10-20 ppm in tissues supporting normal activation and germination. Failed pore activation causes include inadequate hydration preventing turgor generation, boron deficiency disrupting wall modification, metabolic dysfunction from pollen damage or aging preventing enzyme production, extreme temperature stress disrupting activation processes, or developmental abnormalities affecting pore structure or cellular organization. The process represents second aspect of pollen germination preparing emergence pathway, following pollen hydration and preceding pollen tube emergence. Duration typically spans 15-60 minutes as wall modification and polarization proceed. Cultivar variation in pore activation may include aperture structure affecting modification requirements, enzymatic capacity affecting activation speed, and activation efficiency affecting germination success rate. Regional timing shows germination pore activation occurring during flowering period across all climates as part of germination sequence following hydration within hours of pollination. Viticultural significance includes representing critical preparation enabling tube emergence, requiring adequate boron nutrition for normal wall modification, demonstrating coordination of structural, enzymatic, and mechanical processes, and establishing activation success as prerequisite for tube emergence and successful fertilization. Research applications include analyzing aperture selection mechanisms and determinants, investigating wall modification biochemistry and enzymology, evaluating boron effects on activation and wall synthesis, quantifying activation timing and coordination with germination, and assessing activation success as determinant of tube emergence and fertilization outcomes.
Germination_Pore_Activation is second aspect of Pollen_Germination, distinguished by aperture preparation for tube emergence. Represents localized wall modification and cytoplasmic polarization at selected pore. Duration 15-60 minutes. Critical boron dependence for wall synthesis. Temperature affects enzymatic activity. Transitions to Pollen_Tube_Emergence as preparation completes. Essential structural and organizational preparation enabling breakthrough.
Apertur-Aktivierung
Emergenz-Stellenaktivierung
Keimporus-Aktivierung
Porenöffnungsprozess
aperture activation
emergence site activation
germination pore preparation
pore opening process
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 383-385 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 289-290
Germination Pore Activation
Accession DVIT 2082 (Vitis rupestris 'St. George') at the USDA Davis repository bears a Germplasm_Collection_Member_Role with documented passport data, morphological characterization, disease resistance evaluation, and molecular marker genotype—available to researchers through formal request and material transfer agreement. The Domaine de Vassal accession of Pinot Noir (France's reference collection) serves in a Germplasm_Collection_Member_Role with comprehensive ampelographic description, genotypic data, and historical documentation supporting its function as the variety standard. A wild V. amurensis accession at VIR (Vavilov Institute, Russia) bears a Germplasm_Collection_Member_Role preserving cold-hardy germplasm from Siberian populations, with characterization data enabling breeder access for cold-climate breeding programs.
A genetic resource role that inheres in a grapevine genotype formally accessioned into a germplasm repository or collection, maintained under documented curation protocols with assigned accession identifiers enabling tracking, characterization, and distribution to researchers and breeders. Germplasm collection members are conserved ex situ (outside their natural or traditional growing environments) in living collections (field plantings, screen houses), tissue culture (in vitro maintenance), or cryopreservation (long-term frozen storage), with each accession assigned a unique identifier linking to passport data (origin, taxonomy, collection history) and characterization records (morphological descriptions, genotypic data, evaluation results). Major grapevine germplasm collections include USDA National Clonal Germplasm Repository (Davis, California ~5,000 accessions), INRAE Domaine de Vassal (France ~2,700 varieties, 7,800 accessions), JKI Geilweilerhof (Germany ~4,000 accessions), and national collections in Italy, Spain, Portugal, and other viticultural countries. Germplasm collections serve multiple functions—conservation (preventing genetic erosion), characterization (documenting diversity), distribution (providing material to researchers and breeders), and research (supporting genetic and genomic studies). Access to collection material is typically governed by institutional policies and international agreements, with material transfer agreements documenting terms of use.
Germplasm_Collection_Member_Role captures the institutional conservation framework distinguishing formally accessioned material from informal heritage or research collections. The accession structure provides the documentation, traceability, and accessibility essential for effective genetic resource utilization—material can be located, characterized, and obtained through established channels. Collection management faces perpetual challenges: maintaining living collections is resource-intensive (field space, labor, plant health management), while cryopreservation protocols for grapevine are still developing. Molecular characterization increasingly supplements morphological description, with SSR and SNP genotyping enabling duplicate identification, parentage verification, and diversity analysis across collections. International coordination (ECPGR, ITPGRFA) promotes harmonized characterization descriptors, data sharing, and distributed conservation strategies reducing redundancy while ensuring comprehensive coverage.
Genbank-Akzession
Keimplasmasammlung-Mitglied
Repository-Akzession
Sammlungsakzession
collection accession
genebank accession
germplasm accession
repository accession
Maul E et al. (2012) Vitis International Variety Catalogue, JKI This P et al. (2006) Historical origins of wine grapes, Trends in Genetics 22: 511-519 Lacombe T et al. (2011) Grapevine European catalogue, Vitis 50: 65-68 USDA-ARS National Clonal Germplasm Repository (2020) Vitis Germplasm Collection INRAE (2020) Domaine de Vassal Grape Germplasm Repository
Germplasm Collection Member Role
Furmint accession De-Gm-0847-Furmint at the Institut für Rebenzüchtung Geisenheim bearing a germplasm repository role is one of approximately 2,200 accessions maintained in the Geisenheim living collection, authenticated through SSR profiling at 9 standard loci, documented with complete passport data (provenance, collection date, donor institution), maintained under standardized phytosanitary protocols with annual visual inspection for virus symptoms and ELISA testing on a 5-year rotation for GLRaV-1, GLRaV-3, GVA, and GFLV, and available for distribution to qualified researchers under Standard Material Transfer Agreement (SMTA) terms compliant with the International Treaty on Plant Genetic Resources (ITPGRFA).
A research and education role that inheres in a grapevine maintained within a formally designated germplasm collection whose institutional mission encompasses the systematic acquisition, authentication, preservation, documentation, and distribution of grapevine genetic diversity for current and future research, breeding, and conservation purposes.
Germplasm_Repository_Role captures the institutional conservation function of grapevine collections maintained specifically for long-term genetic resource preservation. The role implies formal curation standards including accession numbering, passport data documentation, identity verification, phytosanitary management, and access protocols: these standards distinguish germplasm repositories from commercial nurseries or informal varietal collections. The global grapevine germplasm conservation network encompasses approximately 35,000-40,000 accessions across 100+ institutions, with major collections at INRAE Domaine de Vassal (~2,800 accessions), USDA-ARS Geneva (~3,200), JKI Siebeldingen (~3,200), and Geisenheim (~2,200). Subclasses distinguish the geographic scope of the repository's collection mandate.
Genbank-Rolle
Genressourcen-Repositorium-Rolle
Sortimenterhaltungs-Rolle
genebank accession role
genetic resource repository role
germplasm conservation role
Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157 Lacombe T et al. (2011) Inventory and characterization of Vitis vinifera ssp. vinifera at INRA Domaine de Vassal, Vitis 50: 56-68 FAO (2010) Second Report on the State of the World's Plant Genetic Resources for Food and Agriculture, Rome
Germplasm Repository Role
Systematic evaluation of GFLV_Resistance_Disposition at INRAE Colmar utilized mechanical inoculation to bypass vector-mediated transmission and directly assess host responses to viral infection. Carborundum-assisted rub inoculation of young leaves with purified GFLV particles was performed on 60 Vitis accessions, with systemic infection assessed by ELISA and qRT-PCR at 6 weeks post-inoculation. V. vinifera cultivars showed uniformly high susceptibility, with 95-100% infection rates and high viral titers (10⁶-10⁸ RNA copies per μg total RNA). V. rupestris and V. riparia accessions exhibited variable responses, with infection rates of 60-90% but generally lower viral titers (10⁴-10⁶ copies). Remarkably, Muscadinia rotundifolia accession 'Carlos' showed apparent immunity with 0% systemic infection across 40 inoculated plants, and localized necrosis at inoculation sites suggesting hypersensitive response. When 'Carlos' was used as rootstock with V. vinifera 'Chardonnay' scion, 100% of scions became infected following scion inoculation, demonstrating that M. rotundifolia GFLV_Resistance_Disposition is cell-autonomous and does not confer systemic protection to grafted susceptible scions.
A GFLV_Resistance_Disposition is a Fanleaf_Resistance_Disposition that inheres in a Vitis plant and is realized through host defense processes that directly limit Grapevine fanleaf virus replication, cell-to-cell movement, or systemic spread within plant tissues, independent of resistance to the Xiphinema nematode vector. This disposition represents true antiviral resistance mechanisms operating after the virus has entered the plant, in contrast to vector resistance that prevents initial inoculation. GFLV, a member of the genus Nepovirus (family Secoviridae), has a bipartite positive-sense RNA genome encoding replication proteins on RNA1 and movement and coat proteins on RNA2. Host resistance mechanisms targeting GFLV may include RNA silencing (RNAi) pathways that recognize and degrade viral RNA, translation inhibition preventing viral polyprotein synthesis, interference with viral replicase complex assembly, restriction of cell-to-cell movement through modification of plasmodesmatal transport, and limitation of systemic spread through the vascular system. True genetic resistance to GFLV replication is rare in the Vitis genus, with most accessions supporting viral multiplication following mechanical or nematode-mediated inoculation. Muscadinia rotundifolia exhibits the strongest GFLV resistance phenotype, with some accessions apparently immune to infection. Introgression of M. rotundifolia resistance into V. vinifera backgrounds has proven challenging due to crossing barriers and extensive linkage drag, but represents a long-term breeding objective for comprehensive fanleaf resistance combining both viral and vector resistance components.
GFLV_Resistance_Disposition is_a Fanleaf_Resistance_Disposition. This class represents direct antiviral resistance operating after inoculation, distinguished from Xiphinema_Transmission_Resistance_Disposition which prevents initial virus delivery. True resistance to GFLV replication is rare, with M. rotundifolia providing the strongest known source. The cell-autonomous nature of M. rotundifolia resistance limits its utility in rootstock applications for protecting V. vinifera scions, though resistant rootstocks still provide benefit through vector resistance. Mechanical inoculation studies bypass vector complications but may not fully reflect natural infection dynamics.
Direkte GFLV-Resistenz-Disposition
GFLV-Vermehrungs-Resistenz
Virusreplikations-Resistenz
direct GFLV resistance
nepovirus multiplication resistance
viral replication resistance
Spielmann A et al. (2000) Analysis of transgenic grapevine (Vitis rupestris) and Nicotiana benthamiana plants expressing an Arabis mosaic virus coat protein gene, Plant Science 156:235-244 Vigne E et al. (2004) Evaluation of transgenic Vitis vinifera expressing the coat protein or movement protein genes of Grapevine fanleaf virus, Vitis 43:79-89 Oliver JE, Fuchs M (2011) Tolerance and resistance to viruses and their vectors in Vitis sp., American Journal of Enology and Viticulture 62:438-451
GFLV Resistance Disposition
The GI Australia designation Barossa Valley (registered as a region-level GI within the Barossa zone of the South Australia super zone) delimits approximately 13,000 hectares of vineyards recognized for warm-climate Shiraz, Grenache, and Mourvèdre production — a wine labeled Barossa Valley must contain at least 85% of grapes from the registered GI area, but unlike European PDO appellations the GI imposes no restrictions on grape varieties, maximum yields, or winemaking techniques, with the Australian system relying on Label Integrity Program (LIP) auditing rather than production specification enforcement to ensure geographic origin claims are accurate.
A regional wine designation within the Australian wine regulatory framework that identifies a wine as originating from a registered Geographic Indication (GI) under the Australian Grape and Wine Authority Act 2013, administered by the Geographic Indications Committee (GIC), encompassing a three-tier hierarchy of zones, regions, and sub-regions that delimit Australian wine-producing areas based on geographic and viticultural distinctiveness without prescribing grape varieties, yields, or winemaking methods.
GI_Australia_Designation captures the Australian approach to geographic indication that shares the American AVA philosophy of geographic delimitation without production regulation but implements it within a more structured hierarchical framework. The Australian GI system operates at three levels — zones (broad areas such as South Australia or South Eastern Australia), regions (recognized viticultural areas such as Barossa Valley, McLaren Vale, Margaret River), and sub-regions (smaller distinctive areas within regions such as Eden Valley within the Barossa zone): this hierarchy provides geographic nesting comparable to the European appellation pyramid but without the associated production rule escalation. The super zone South Eastern Australia encompasses approximately 95% of Australian wine production and appears on labels of multi-regional blends, functioning as the broadest commercially used GI. Australia has bilateral wine trade agreements with the EU recognizing mutual protection of geographic indications.
Australian GI
Australian geographic indication
Australian wine region designation
australische GI-Bezeichnung
australische Herkunftsangabe
australische Weinregion-Kennung
Australian Grape and Wine Authority Act 2013 Wine Australia (2023) Register of Protected GIs and Other Terms Australia-EU Wine Trade Agreement (2008)
GI Australia Designation
Glabrous surface quality on Chasselas shoot tips and young leaves (OIV descriptor 004 rating 1, none or very sparse) distinguishes this cultivar from most other major white wine varieties and contributes to the bright green, shiny appearance of young Chasselas shoots valued by Swiss vignerons in the Lavaux region the glabrous condition persists on mature leaf upper surfaces (OIV 051 rating 1), creating the smooth, glossy canopy appearance characteristic of Chasselas vineyards in contrast to the more matte, pubescent canopies of Chardonnay or Pinot Blanc growing in adjacent blocks.
Glabrous_Surface_Quality represents the surface texture condition where the external boundary of a grapevine organ lacks trichomes, presenting a hairless surface regardless of other texture properties such as smoothness or roughness. Glabrous surfaces occur on organs or organ regions where trichomes either never develop or have been lost through maturation processes. In grapevine, berry surfaces are essentially glabrous in all Vitis vinifera cultivars, lacking the trichomes found on vegetative organs. Upper (adaxial) leaf surfaces range from glabrous to sparsely pubescent depending on cultivar, while lower (abaxial) surfaces display the full range from glabrous through densely tomentose. Young shoot tips may be glabrous in some cultivars, contrasting with the densely pubescent tips of others, providing a primary ampelographic identification character. The glabrous condition exposes the underlying cuticular surface, making other texture qualities (smooth versus rough) readily perceptible. Glabrous surfaces may present higher boundary layer conductance than pubescent surfaces, potentially increasing transpiration rates but also facilitating gas exchange for photosynthesis. The absence of trichomes removes a physical barrier to small arthropod herbivores and may affect pathogen spore adhesion and germination. Assessment of glabrous versus pubescent condition employs visual observation for absence of visible hairs, tactile evaluation detecting only the underlying surface texture without fuzzy or bristly sensation, and microscopic confirmation of trichome absence.
Glabrous_Surface_Quality is the complement of Pubescent_Surface_Quality, specifically addressing the absence of trichomes. A glabrous surface may be either smooth or rough the glabrous classification addresses only trichome presence, not other texture properties. OIV descriptors rate pubescence density, with category 1 (none or very sparse) corresponding to glabrous condition. The glabrous surface of grape berries is universal in Vitis vinifera and distinguishes grapes from many other fruit crops with pubescent fruit surfaces. Assessment should distinguish truly glabrous surfaces from surfaces where sparse trichomes are present but inconspicuous.
bald texture
glabröse Qualität
haarlose Textur
hairless surface
kahle Oberfläche
non-pubescent surface
smooth-surfaced quality
unbehaarte Oberflächenbeschaffenheit
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 47-54 Konlechner C and Sauer U (2016) Ultrastructural Leaf Features of Grapevine Cultivars (Vitis vinifera L. ssp. vinifera). OENO One 50(4): 195-207
Glabrous Surface Quality
Investigation of Glassy_Winged_Sharpshooter_Resistance_Disposition at UC Riverside employed video tracking and electrical penetration graph (EPG) analysis to characterize H. vitripennis feeding behavior on resistant versus susceptible Vitis genotypes. Adult sharpshooters were confined on test plants for 24-hour sessions with continuous EPG monitoring of feeding waveforms. On susceptible V. vinifera 'Chardonnay', sharpshooters achieved sustained xylem ingestion (X-wave) for mean cumulative duration of 14.2 hours per 24-hour session, with ingestion bouts averaging 85 minutes. On the PdR1-carrying resistant selection 94-27-008 (V. arizonica × V. vinifera), xylem ingestion duration was reduced to 8.5 hours with significantly more frequent stylet withdrawal events, though sustained feeding still occurred. Notably, 'Chardonnay' and 94-27-008 showed equivalent X. fastidiosa transmission rates when sharpshooters fed first on infected source plants and then on test plants (75% vs 72% transmission), indicating that the Glassy_Winged_Sharpshooter_Resistance_Disposition of 94-27-008 was insufficient to prevent inoculation. This finding confirms that PdR1-mediated Pierce's disease resistance operates through bacterial resistance mechanisms (limiting X. fastidiosa multiplication) rather than through vector deterrence, and that GWSS-specific resistance remains an unmet breeding objective that could enhance durability of existing bacterial resistance.
A Glassy_Winged_Sharpshooter_Resistance_Disposition is a Leafhopper_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that limit feeding, reproduction, or Xylella fastidiosa transmission by the glassy-winged sharpshooter (Homalodisca vitripennis), a large xylem-feeding leafhopper that has emerged as the most important Pierce's disease vector in California and poses ongoing invasion threats to other viticultural regions. This disposition represents resistance to an insect whose primary economic impact derives not from direct feeding damage—which is minimal despite prodigious xylem consumption rates of up to 100× body weight per day—but from its exceptional efficiency as a Xylella vector, capable of transmitting the bacterium within minutes of feeding on infected plants and retaining infectivity through adult molts. H. vitripennis is native to the southeastern United States but invaded California in the 1990s, where it has dramatically altered Pierce's disease epidemiology due to its large size enabling long-distance dispersal, broad host range supporting populations on numerous ornamental and crop plants, and overwintering adults that transmit X. fastidiosa in early spring when native vectors are inactive. Resistance mechanisms affecting H. vitripennis may include xylem sap composition deterring sustained feeding, physical properties of xylem vessels impeding stylet penetration, induced defenses triggered by probing that cause stylet withdrawal, and antixenosis reducing settling preference. While direct resistance to H. vitripennis feeding would complement Xylella resistance for comprehensive Pierce's disease protection, research has focused primarily on bacterial resistance (PdR1 locus) rather than vector resistance, as the vector's broad host range limits the impact of grapevine-specific deterrence.
Glassy_Winged_Sharpshooter_Resistance_Disposition is_a Leafhopper_Resistance_Disposition. H. vitripennis is the primary Pierce's disease vector in California, with vector efficiency and ecological characteristics that have transformed disease epidemiology since its 1990s invasion. This class addresses resistance to the insect vector, distinct from Pierces_Disease_Resistance_Disposition which addresses resistance to X. fastidiosa itself. Current Pierce's disease resistant cultivars rely on bacterial resistance (PdR1) rather than vector resistance. True GWSS-specific resistance could complement bacterial resistance for enhanced durability but has not been a primary breeding target due to the vector's polyphagous nature limiting impact of grapevine resistance alone.
GWSS resistance
GWSS-Resistenz
Glasflügelzikaden-Resistenz-Disposition
Homalodisca resistance
Homalodisca-vitripennis-Abwehr
sharpshooter vector resistance
Almeida RPP, Purcell AH (2003) Homalodisca vitripennis (Germar) (Hemiptera, Cicadellidae) transmission of Xylella fastidiosa to almond, Plant Disease 87:1255-1259 Backus EA et al. (2012) EPG comparisons of feeding behavior and X. fastidiosa transmission rates by glassy-winged sharpshooter on Vitis vinifera and Nicotiana tabacum, Journal of Economic Entomology 105:629-637 Blua MJ et al. (2001) The Glassy-winged Sharpshooter in California, Proceedings of the Pierce's Disease Research Symposium, California Department of Food and Agriculture
Glassy Winged Sharpshooter Resistance Disposition
Global acreage for the top five wine grape cultivars by planted area (Anderson and Nelgen 2020, approximately 2016 data year): Cabernet Sauvignon approximately 340,000 ha (the world's most widely planted wine grape since overtaking Merlot circa 2015), Merlot approximately 266,000 ha, Tempranillo approximately 231,000 ha, Airén approximately 218,000 ha (the world's most widely planted white grape, almost entirely in Spain's La Mancha), and Chardonnay approximately 210,000 ha — the top 5 cultivars collectively occupying approximately 1.27 million ha, or approximately 18% of the world's approximately 7.3 million ha of vineyard. Global acreage for Grüner Veltliner approximately 18,500 ha, of which Austria accounts for approximately 73% (13,500 ha), the Czech Republic approximately 8%, Slovakia approximately 5%, and the remainder scattered across Germany, Hungary, Italy, the United States, New Zealand, and Australia — illustrating a cultivar with modest global acreage but strong national dominance in its primary country.
Global acreage is a production statistics entity that documents the total worldwide vineyard area planted to a specific grape cultivar, aggregated from national and regional acreage data, expressed in hectares (ha), and referencing the data year, compilation source, and methodological notes regarding data completeness and reliability, providing the single most widely used quantitative indicator of a cultivar's global commercial importance and enabling cross-cultivar comparison of worldwide cultivation significance.
Global acreage provides the broadest quantitative measure of cultivar commercial importance — the single number that positions each cultivar within the worldwide viticultural landscape and enables the most basic ranking of cultivar significance by cultivation extent. The interpretive value of global acreage requires contextual qualification on several dimensions — acreage does not directly indicate quality importance (Airén's 218,000 ha produces primarily bulk distillation material while Nebbiolo's 5,500 ha produces some of the world's most celebrated wines), acreage does not indicate price or revenue (a cultivar's economic value depends on yield, wine quality, and price per bottle as well as on planted area), and global aggregates mask the geographic concentration patterns that are often more informative than the total (Grüner Veltliner's 18,500 ha global acreage is dominated by Austria, making it essentially a national rather than global cultivar by distribution). The temporal stability of global acreage statistics also varies — some cultivars show remarkable acreage stability over decades (Riesling has maintained approximately 50,000-55,000 ha globally for over 30 years), while others show dramatic expansion (Sauvignon Blanc more than doubled from approximately 65,000 ha in 2000 to approximately 125,000 ha by 2020) or contraction (Carignan declined from over 200,000 ha in the 1980s to approximately 80,000 ha by 2020 through systematic EU-subsidized grubbing). The data reliability caveat is essential — global acreage figures represent the best available compilation from sources of varying quality, with confidence intervals that may span ±10-20% for cultivars where major producing countries (notably China) provide imprecise data, meaning that global acreage statistics should be understood as informed estimates rather than precise measurements.
Gesamtweltanbaufläche
global planting area
global vineyard area
globale Anbauausdehnung
globale Rebfläche
total world acreage
weltweite Anbaufläche
weltweite Bestockung
worldwide cultivation extent
worldwide planted area
Anderson K, Nelgen S (2020) Which Winegrape Varieties are Grown Where? Revised edition, University of Adelaide Press OIV (2023) State of the World Vine and Wine Sector OIV (2017) Distribution of the World's Grapevine Varieties, Focus OIV
Global Acreage
Pinot Noir berries exhibit characteristic globose form with length-to-width ratios of 0.97-1.03 across diverse growing regions from Burgundy to Oregon to New Zealand, demonstrating the strong genetic control of this shape character the compact clustering of these uniformly round berries contributes to the cultivar's susceptibility to Botrytis bunch rot by creating microenvironments with high humidity and limited air circulation between tightly packed spherical fruit.
Globose_Quality represents the overall form characteristic of organs exhibiting approximately spherical geometry with essentially equal dimensions along all three principal axes. Globose form is defined by a length-to-width ratio approaching unity (typically 0.95-1.05) and uniform curvature of the bounding surface in all directions. Among grape berries, globose form is one of the most common configurations, characteristic of many widely cultivated wine and table grape varieties including Pinot Noir, Chardonnay, Chasselas, and Muscat of Alexandria. Globose berries exhibit maximum volume relative to surface area among common berry shapes, which has implications for the ratio of skin-derived compounds to juice volume. The spherical geometry results from isotropic cell expansion during berry development, where growth occurs at similar rates in all directions. Deviation from perfect sphericity commonly occurs due to mutual compression of berries in compact clusters, seed number and position effects on internal expansion patterns, and asymmetric vascular supply. Globose form in other organs is less common but occurs in some cultivar-specific structures. The OIV descriptor 223 (berry shape) includes globose as the first category, recognizing its frequency and distinctiveness. Quantitative assessment of globose form employs the length-to-width ratio measured on representative berry samples, with values between 0.95 and 1.05 indicating globose classification.
Globose_Quality represents the limiting case of Overall_Form_Quality where all principal axes are equal. The class aligns with PATO_0000411 (spheroid) as an upper-ontology reference. Practical measurement typically assesses only two dimensions (polar and equatorial diameter) and infers three-dimensional form true sphericity would require confirmation that the third axis is also equal. Minor deviations from perfect sphericity are expected in biological organs the globose classification applies when the overall impression and measured ratios indicate approximately equal dimensions. OIV descriptor 223 category 1 (globose) provides the standardized ampelographic reference.
Kugelform
globular quality
globuläre Gestalt
kugelige Qualität
round shape
runde Form
spherical form
spheroid form
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 83-84 Boursiquot JM et al. (2009) Parentage of Merlot and Related Winegrape Cultivars of Southwestern France: Discovery of the Missing Link. Australian Journal of Grape and Wine Research 15(2): 144-155
Globose Quality
Glories index monitoring through Merlot ripening reveals the cell wall maturation affecting extraction potential, with véraison +3 weeks showing total anthocyanins 1.95 mg/g but EA only 38% (cell walls intact, only 0.74 mg/g extractable), seed maturity index 62% (high seed tannin contribution expected), véraison +5 weeks showing total anthocyanins 1.72 mg/g with EA 58% (improving extractability, 1.00 mg/g extractable), SM 48%, and véraison +7 weeks showing total 1.45 mg/g with EA 72% (advanced cell wall degradation, 1.04 mg/g extractable despite lower total), SM 32% (seed contribution minimized), demonstrating how Glories indices identify optimal phenolic maturity where extractable quality exceeds that achievable from higher-content but lower-extractability fruit.
Glories index quality is a maturity index quality that represents the phenolic extractability indices developed by Yves Glories and colleagues at the University of Bordeaux, characterizing the relationship between total phenolic content and extractable phenolic content that indicates the cell wall maturity state affecting how much of the grape's phenolic potential will be realized during winemaking. Within the BFO framework, this quality specifically addresses the extractability dimension of phenolic maturity indices, distinguishing Glories-type indices from simple concentration measures while recognizing that these indices capture the critical mediation of cell walls between grape phenolic content and wine phenolic composition, providing information about extraction potential that total phenolic analysis alone cannot reveal. The ontological significance lies in Glories indices distinguishing between phenolic accumulation and phenolic availability, where the cellular extractability index (EA) for anthocyanins and the seed maturity index (SM) for tannins indicate whether cell wall and seed coat barriers will permit effective extraction during maceration, with high extractability (EA >60%) indicating advanced cell wall degradation enabling efficient anthocyanin release, and low seed contribution (SM <40%) indicating that seed tannin release will not dominate wine tannin character. The calculation methodology for Glories indices involves comparison of phenolic extraction under different conditions, with the anthocyanin extractability index (EA) calculated from the ratio of anthocyanins extracted at wine pH versus total anthocyanins released by more complete extraction (typically at pH 1 or with pectinase treatment), yielding values from 0% (no extraction at wine pH, completely locked in cells) to 100% (complete extraction at wine pH, cell walls fully permeable). The seed maturity index (SM) calculation involves comparing tannin extracted at wine pH (skin-dominated) versus total tannin (skin plus seed), with high SM indicating substantial seed tannin contribution (immature seeds with extractable, harsh tannins) and low SM indicating skin tannin dominance (mature seeds with reduced extractability, softer wine tannin profile). The interpretive framework for Glories indices establishes that optimal phenolic maturity typically corresponds to EA values above 55-60% (indicating cell wall modification enabling efficient color extraction) combined with SM values below 40-50% (indicating reduced seed tannin contribution enabling softer wine structure), with indices outside these ranges suggesting either premature harvest (low EA, high SM) or potentially overripe conditions (very high EA with declining total anthocyanins). Understanding Glories index quality enables assessment of phenolic extraction potential beyond simple concentration measures, harvest timing that optimizes extractable rather than total phenolics, and recognition of the cell wall maturity factors that determine winemaking success for red wines.
Glories_Index_Quality addresses the phenolic extractability indices distinguishing total from extractable phenolic content. This quality captures cell wall maturity affecting winemaking extraction efficiency. The class supports phenolic quality optimization through extractability index characterization beyond simple concentration analysis.
EA index
EA-Index
Extrahierbarkeitsindex
Glories-Index
cellular maturity index
extractability index
phenolic maturity index
phenolischer Reifeindex
Saint-Cricq N et al. (1998) Méthode de caractérisation de la maturité phénolique des raisins à partir de l'évolution de l'extractibilité des anthocyanes. Journal International des Sciences de la Vigne et du Vin 32(1): 1-10 Glories Y (2001) Caractérisation du potentiel phénolique: adaptation de la vinification. Progrès Agricole et Viticole 118(15-16): 347-350
Glories Index Quality
Gloss quality assessment tracking berry development in Merlot vineyard shows increasing gloss from early véraison (matte appearance, gloss rating 3/10, heavy bloom obscuring cuticle) through mid-ripening (developing sheen, 5/10, cuticle expansion creating smoother surface) to harvest maturity (pronounced gloss, 8/10, taut cuticle producing strong specular reflection visible alongside bloom), with gloss development serving as a visual ripening indicator complementing Brix and flavor sampling for harvest timing decision.
Gloss quality is a surface appearance quality that represents the specular reflectivity of grape berry or wine surfaces, characterizing the shiny, lustrous appearance arising from smooth surface structure that reflects light directionally rather than scattering it diffusely. Within the BFO framework, this quality specifically addresses the specular reflection dimension of surface appearance, distinguishing gloss from diffuse reflection (which produces matte appearance) while recognizing that gloss in grape and wine contexts provides different but complementary quality information depending on assessment target. The ontological significance lies in gloss serving as a maturity and condition indicator in grapes and a clarity indicator in wine, where increasing grape berry gloss during ripening signals cuticle development and approaching harvest readiness, while wine surface gloss indicates clarity and absence of surface contamination, making gloss relevant across viticultural and enological assessment contexts. The physical basis for gloss involves specular reflection from smooth surfaces where reflected light maintains directional coherence rather than scattering, with surface smoothness at the wavelength scale determining gloss intensity, and factors disrupting smoothness (wax bloom on grapes, particles or films on wine) reducing gloss through diffuse scattering. The compositional and developmental determinants of gloss differ between grape and wine contexts, with grape gloss reflecting cuticle thickness and smoothness beneath the wax bloom layer (gloss increasing as berry expansion smooths the cuticle surface during ripening), while wine gloss reflects clarity (brilliant wines showing higher surface gloss) and absence of surface-active contaminants (films or particles reducing gloss). The assessment methodology for gloss involves observation of light reflection characteristics, with glossy surfaces showing distinct highlight reflections while matte surfaces show even illumination without highlights. Understanding gloss quality enables grape maturity assessment from increasing gloss during ripening, wine clarity confirmation from surface gloss observation, and quality evaluation incorporating gloss as a condition indicator in both viticultural and enological contexts.
Gloss_Quality addresses the specular reflection dimension of surface appearance in both grape berry and wine contexts. This quality provides maturity indication in grapes (gloss increasing with ripening) and clarity indication in wine (gloss reflecting brilliance). The class supports quality assessment across the grape-to-wine continuum through surface reflectivity observation.
Oberflächenglanz
Schimmer
Spiegelglanz
shine
specular reflectivity
surface luster
surface sheen
Rosenquist JK, Morrison JC (1989) Some factors affecting cuticle and wax accumulation on grape berries. American Journal of Enology and Viticulture 40(4): 241-244 Pensec F et al. (2014) Changes in the epicuticular wax of grape berries during development and at ripening. Journal of Agricultural and Food Chemistry 62(22): 5024-5029
Gloss Quality
Comprehensive screening for GLRaV_Replication_Resistance_Disposition at Cornell University evaluated 180 Vitis accessions including wild species, interspecific hybrids, and V. vinifera cultivars using chip-bud graft inoculation with GLRaV-3-infected buds. Systemic infection was assessed by ELISA and RT-PCR at 6 and 12 months post-grafting. Results revealed universal susceptibility with no accession showing immunity or high-level resistance. V. vinifera cultivars exhibited 95-100% transmission rates with high viral titers (Ct values 18-22 in RT-qPCR). North American species (V. riparia, V. rupestris, V. labrusca) showed equivalent susceptibility (90-100% transmission) despite evolutionary separation from GLRaV-endemic regions. Even M. rotundifolia, which provides strong resistance to GFLV and many fungal pathogens, supported GLRaV-3 replication with 85% transmission rates. The only statistically significant variation detected was in symptom expression and yield impact, with 'Chambourcin' and 'Norton' showing tolerance phenotypes maintaining 90% of healthy vine productivity despite infection, compared to 65% productivity retention in symptomatic 'Cabernet Franc'. These findings confirm that breeding for GLRaV_Replication_Resistance_Disposition faces fundamental limitations in available germplasm diversity.
A GLRaV_Replication_Resistance_Disposition is a Leafroll_Resistance_Disposition that inheres in a Vitis plant and is realized through host defense processes that directly limit Grapevine leafroll-associated virus replication, phloem cell colonization, or systemic spread within the plant vascular system, independent of resistance to mealybug or soft scale vectors. This disposition represents true antiviral resistance mechanisms operating after viral particles have been introduced into phloem tissues by vector feeding. GLRaVs are phloem-limited, positive-sense RNA viruses belonging to the families Closteroviridae (GLRaV-1, -3) and Botourmiaviridae (GLRaV-2), with complex genomes encoding replicases, movement proteins, and multiple coat protein variants. Host resistance mechanisms potentially targeting GLRaVs include RNA silencing pathways operative in phloem companion cells, restriction factors limiting viral replicase function, impaired viral movement through plasmodesmata connecting sieve elements and companion cells, and phloem-specific defense responses. True genetic resistance to GLRaV replication has proven extremely difficult to identify in Vitis germplasm, with screening of hundreds of accessions failing to reveal immune or highly resistant genotypes. All tested V. vinifera cultivars support GLRaV replication when graft-inoculated, though viral titers and symptom severity vary. Wild Vitis species show similarly broad susceptibility, with no strong resistance source identified for breeding programs. This absence of natural resistance has motivated research into transgenic approaches expressing viral coat proteins or RNA interference constructs, though these remain experimental.
GLRaV_Replication_Resistance_Disposition is_a Leafroll_Resistance_Disposition. This class represents direct antiviral resistance operating after inoculation, distinguished from Vector_Transmission_Resistance_Disposition which prevents initial virus delivery. Despite extensive germplasm screening, no strong natural resistance to GLRaV replication has been identified in Vitis or Muscadinia. This absence of resistance sources distinguishes leafroll from fanleaf disease, where M. rotundifolia provides strong GFLV resistance. Current management relies on clean planting material and vector control rather than genetic resistance. Tolerance variation among cultivars provides the only genetic tool currently available.
Ampelovirus-Vermehrungs-Resistenz
Direkte GLRaV-Resistenz-Disposition
GLRaV-Replikations-Resistenz
ampelovirus multiplication resistance
direct GLRaV resistance
leafroll replication resistance
Oliver JE, Fuchs M (2011) Tolerance and resistance to viruses and their vectors in Vitis sp., American Journal of Enology and Viticulture 62:438-451 Maree HJ et al. (2013) Grapevine leafroll-associated virus 3, Frontiers in Microbiology 4:82 Burger JT et al. (2017) Grapevine leafroll-associated viruses, Grapevine Viruses: Molecular Biology, Diagnostics and Management, Springer
GLRaV Replication Resistance Disposition
Comparative analysis of Glucanase_Induction_Disposition at INRAE Bordeaux examined β-1,3-glucanase gene expression and activity across 18 cultivars following Plasmopara viticola inoculation, specifically targeting the oomycete-relevant glucanase response. Expression of three glucanase genes (VvGluc1 class I, VvGluc2 class II, VvGluc3 class II) was monitored at 0, 12, 24, 48, and 72 hpi alongside enzyme activity assays using laminarin substrate. Resistant 'Regent' (Rpv3-positive) showed strong Glucanase_Induction_Disposition with VvGluc3 reaching 35-fold induction by 18 hpi and total glucanase activity increasing 6-fold by 24 hpi. Susceptible 'Chasselas' exhibited delayed, weak induction with VvGluc3 only 5-fold at 48 hpi and activity increasing 1.8-fold. Importantly, class II extracellular glucanase (VvGluc3) showed strongest correlation with resistance (r = -0.79 with sporulation intensity), suggesting that apoplastic deployment is critical for defense against the intercellularly colonizing P. viticola. Combined glucanase-chitinase activity assays demonstrated synergistic effects, with the enzyme combination inhibiting in vitro germination of Botrytis cinerea conidia by 85% compared to 45% (glucanase alone) and 55% (chitinase alone) at equivalent concentrations. For P. viticola zoospores lacking chitin, glucanase alone provided 60% germination inhibition, confirming the importance of Glucanase_Induction_Disposition specifically for oomycete resistance where chitinase has no direct target.
A Glucanase_Induction_Disposition is a Pathogenesis_Related_Protein_Disposition that inheres in a Vitis vinifera plant and is realized through rapid and strong transcriptional activation and accumulation of β-1,3-glucanase enzymes (PR-2 family) in response to pathogen attack, providing enhanced capacity to degrade β-1,3-glucans, major structural polysaccharides in the cell walls of fungi and oomycetes. This disposition represents the induction of another critical PR protein family for antimicrobial defense, as β-1,3-glucanase activity directly compromises pathogen cell wall integrity and can synergize with chitinases for enhanced antifungal effect. Unlike chitinases, β-1,3-glucanases are effective against both true fungi (which contain chitin and β-glucans) and oomycetes like Plasmopara viticola (which lack chitin but contain abundant β-1,3-glucans), making Glucanase_Induction_Disposition particularly important for downy mildew resistance. Grapevine β-1,3-glucanases are classified into class I (vacuolar, with N-terminal extension) and class II (extracellular, without extension) based on structure and localization, with both classes contributing to defense through complementary spatial deployment. The hydrolysis products of glucanase activity (β-glucan oligosaccharides) can function as elicitors triggering additional defense responses, similar to chitin oligomers released by chitinases. Glucanase and chitinase induction typically occur coordinately, with the combined activity providing synergistic antifungal effects exceeding either enzyme alone through simultaneous attack on multiple cell wall components.
Glucanase_Induction_Disposition is_a Pathogenesis_Related_Protein_Disposition. β-1,3-Glucanases target β-glucans present in both fungal and oomycete cell walls, providing broader pathogen coverage than chitinases which are ineffective against chitin-lacking oomycetes. Class II (extracellular) glucanases may be particularly important for defense against pathogens like P. viticola that colonize apoplastic spaces. Glucanase and chitinase typically show coordinated induction and synergistic antifungal effects, representing a coherent defense module targeting pathogen cell wall integrity.
Glucanase-Induktions-Disposition
PR-2 induction
PR-2-Induktion
glucan hydrolase response
β-1,3-Glucanase-Aktivierung
β-1,3-glucanase activation
Mauch F et al. (1988) Antifungal hydrolases in pea tissue I. Purification and characterization of two chitinases and two β-1,3-glucanases differentially regulated during development and in response to fungal infection, Plant Physiology 87:325-333 Renault AS et al. (1996) β-1,3-Glucanase gene expression in grapevine leaves as a response to infection with Botrytis cinerea, American Journal of Enology and Viticulture 47:57-62 van Loon LC et al. (2006) Significance of inducible defense-related proteins in infected plants, Annual Review of Phytopathology 44:135-162
Glucanase Induction Disposition
Glucose content quality monitoring during fermentation of Château Margaux Grand Vin employs daily enzymatic analysis tracking the decline from initial 110-115 g/L glucose through progressive depletion, with the steep glucose consumption phase (days 2-7, dropping to below 10 g/L) indicating healthy yeast metabolism, while persistently elevated glucose levels beyond day 10 signal potential fermentation problems requiring intervention the transition from glucose to fructose fermentation is marked by declining fermentation rate as Saccharomyces shifts to the less-preferred substrate, a normal progression that winemakers monitor to anticipate stuck fermentation risk.
Glucose_Content_Quality represents the chemical quality characterizing the concentration of glucose (dextrose, D-glucose, grape sugar) in grape berry tissue. Glucose is a six-carbon aldose monosaccharide that constitutes approximately half of the total reducing sugar in ripe grape berries, typically reaching concentrations of 75-125 g/L at harvest maturity for wine grapes. Together with fructose, glucose forms the primary fermentable substrate for alcoholic fermentation, being readily metabolized by Saccharomyces cerevisiae through glycolysis to produce ethanol and carbon dioxide. Glucose accumulates in berry vacuoles primarily through cleavage of phloem-transported sucrose by cell wall and vacuolar invertases, with the resulting glucose and fructose compartmentalized in the large central vacuoles of mesocarp cells. The concentration of glucose relative to fructose (glucose-fructose ratio, G/F ratio) provides diagnostic information about berry maturation status and metabolic condition. Early in ripening, the G/F ratio may slightly favor glucose, transitioning toward equality (G/F approximately 1.0) at maturity, and potentially shifting toward fructose dominance in over-ripe or dehydrated fruit because glucose is more readily metabolized. Glucose is less sweet than fructose by approximately 25-30%, so berries with higher glucose proportion taste less sweet than equi-concentrated berries with higher fructose. In winemaking, Saccharomyces preferentially ferments glucose over fructose due to glucose repression of fructose transport, resulting in rising G/F ratio during fermentation until glucose is nearly depleted, whereupon fructose fermentation accelerates. Glucose-specific measurement employs enzymatic assays using hexokinase and glucose-6-phosphate dehydrogenase, or high-performance liquid chromatography.
Glucose_Content_Quality specifically addresses the glucose monosaccharide, distinguished from Fructose_Content_Quality and from aggregate sugar measures. Glucose is the reference compound for the °Brix scale (grams sucrose per 100 g solution, with sucrose hydrolysis yielding glucose and fructose), creating a conceptual link between glucose and practical sugar measurement. The metabolic centrality of glucose in cellular energy production makes it significant beyond its role as fermentation substrate. Glucose can form glycosidic bonds with aroma compounds and anthocyanins, affecting the extractability and stability of these quality-relevant molecules.
D-glucose level
Dextrosekonzentration
Glucoseanteil
Glucosegehalt
Traubenzuckergehalt
blood sugar homolog quality
dextrose content
grape sugar concentration
Kliewer WM (1967) The Glucose-Fructose Ratio of Vitis vinifera Grapes. American Journal of Enology and Viticulture 18(1): 33-41 Bisson LF (1999) Stuck and Sluggish Fermentations. American Journal of Enology and Viticulture 50(1): 107-119
Glucose Content Quality
Glutamine content quality in Sonoma Coast Chardonnay from high-nitrogen vineyard blocks reaches 100-180 mg/L, indicating vigorous nitrogen translocation to fruit that supports rapid, complete fermentation but may contribute to the reduced complexity sometimes associated with excessive vine nitrogen site selection and nitrogen management targeting moderate glutamine levels (40-80 mg/L) enable winemakers to balance fermentation assurance with the restrained fruit expression valued in premium cool-climate Chardonnay styles emphasizing mineral and textural character over primary fruit.
Glutamine_Content_Quality represents the chemical quality characterizing the concentration of the amino acid glutamine in grape must, capturing a major transportable nitrogen form that reflects vine nitrogen status and provides rapidly assimilable nitrogen for yeast metabolism. Glutamine is an amide amino acid (glutamate with an amide group on the gamma-carboxyl) that serves as a primary nitrogen transport compound in plants, shuttling nitrogen from roots and senescing tissues to growing organs and developing fruit. In grape must, glutamine concentration typically ranges from 20-200 mg/L, with higher levels indicating strong vine nitrogen status and active nitrogen translocation to berries. Glutamine is among the preferred nitrogen sources for Saccharomyces cerevisiae, taken up rapidly through general amino acid permeases and assimilated directly into central nitrogen metabolism through glutamate formation. The preferential uptake of glutamine (alongside glutamate, asparagine, and aspartate) means it is depleted early in fermentation, contributing to initial fermentation vigor. The nitrogen-sensing pathways in yeast (NCR—nitrogen catabolite repression) respond strongly to glutamine availability, making glutamine content influential beyond its direct nitrogen contribution by affecting the regulation of other nitrogen metabolism genes. Glutamine-rich musts may show different nitrogen metabolism patterns than musts with equivalent YAN from other amino acid profiles. The lability of glutamine (spontaneous deamidation to glutamate under acidic conditions and elevated temperature) means that glutamine content can decrease during must handling and storage, requiring timely analysis for accurate measurement. Glutamine analysis employs HPLC amino acid methods with appropriate sample handling to prevent degradation.
Glutamine_Content_Quality addresses an amino acid with significance as a nitrogen transport form linking vine status to must composition. The regulatory role of glutamine in yeast nitrogen metabolism (NCR signaling) means its effects extend beyond simple nitrogen contribution. The chemical lability of glutamine (deamidation to glutamate) distinguishes it from stable amino acids and requires analytical attention. The glutamine/glutamate ratio in must reflects both vine nitrogen metabolism and any degradation during handling, providing information about sample history alongside vine status.
Amidaminosäuregehalt
Glutamingehalt
L-Glutamin-Konzentration
L-glutamine concentration
Stickstofftransport-Aminosäurequalität
amide amino acid content
glutamine level
nitrogen transport amino acid quality
Bell SJ and Henschke PA (2005) Implications of Nitrogen Nutrition for Grapes, Fermentation and Wine. Australian Journal of Grape and Wine Research 11(3): 242-295 ter Schure EG et al. (2000) The Role of Ammonia Metabolism in Nitrogen Catabolite Repression in Saccharomyces cerevisiae. FEMS Microbiology Reviews 24(1): 67-83
Glutamine Content Quality
In Moscato Bianco berries in Asti, Piedmont during BBCH 85-89, glycosidic precursor synthesis generates monoterpene glucosides at rates of 0.12 mg/L per day during peak activity, with UDP-glucose linalool glucosyltransferase activity reaching 52 nmol/min/g fresh weight in exocarp tissue at 22 days post-veraison, producing bound linalool of 2.8 mg/L as monoglucoside and 0.9 mg/L as disaccharide forms, representing 91% of total linalool in glycosidic conjugates under warm Piedmont conditions averaging 25°C.
Glycosidic precursor synthesis describes the enzymatic conjugation of volatile aroma compounds with sugar moieties in Vitis vinifera berry cells during ripening, wherein UDP-glucose-dependent glycosyltransferases catalyze the transfer of glucose residues from activated sugar donors to hydroxyl groups of monoterpenols, sesquiterpenols, and other volatile alcohols, producing beta-glucoside conjugates that represent the predominant storage form of aromatic compounds in grape berries and constitute 85-95% of total terpenoid content at harvest maturity. This process involves the recognition of diverse aglycone substrates by glycosyltransferase active sites, with broad-specificity enzymes accepting multiple terpenol substrates including linalool, geraniol, nerol, citronellol, and alpha-terpineol, enabling the glycosylation of the complete spectrum of monoterpene alcohols produced through terpenoid biosynthetic pathways in berry tissues. The biochemical mechanism proceeds through nucleophilic attack on UDP-glucose by the hydroxyl group of the acceptor terpenol, forming an enzyme-bound glucose ester intermediate followed by transfer to the aglycone with UDP release, with reaction rates determined by enzyme expression levels, substrate concentrations, and cofactor availability that vary developmentally and show cultivar-specific regulation patterns. Secondary glycosylation reactions further modify monoterpene glucosides through attachment of additional sugar residues including rhamnose, arabinose, and apiose by specific rhamnosyltransferases and related enzymes, generating disaccharide and trisaccharide glycosides that demonstrate enhanced stability and modified release characteristics compared to simple monoglucosides during subsequent hydrolysis. The cellular compartmentalization of glycosidic precursor synthesis involves cytosolic localization of most glycosyltransferase activities with subsequent transport of synthesized conjugates into vacuolar storage through tonoplast transporters, establishing protected precursor pools sequestered from cytosolic glycosidase activities that would prematurely hydrolyze glycosidic bonds releasing volatile aglycones. Cultivar genetics profoundly influence glycosidic precursor synthesis capacity through differential expression of glycosyltransferase genes, with aromatic varieties such as Muscat showing 5-15 fold higher glucosyltransferase expression and activity compared to neutral varieties such as Chardonnay, directly determining the accumulation capacity for glycosidically-bound terpenoids characteristic of aromatic grape varieties. Temperature during ripening modulates glycosidic precursor synthesis through effects on glycosyltransferase enzyme activities, with optimal synthesis rates at moderate temperatures of 22-28°C and reduced activity at temperature extremes that may limit precursor accumulation or shift the balance between free and bound volatile pools.
Glycosidic_Precursor_Synthesis is subclass of Aroma_Precursor_Formation representing specific glycosyltransferase-mediated reactions. Distinguishes from thiol precursor formation using glutathione conjugation. Sibling to Bound_Aroma_Accumulation and Precursor_Pool_Building. Key enzymes are UDP-glucose-dependent glycosyltransferases. Produces mono-, di-, and tri-glycoside conjugates. Cultivar-specific expression determines capacity.
Glycosyltransferase-vermittelte-Konjugation
Monoterpenol-Glycosylierung
Terpen-Glucosid-Bildung
glycosidische Vorläufer-Synthese
glycosyltransferase-mediated conjugation
monoterpenol glycosylation
sugar-bound volatile synthesis
terpene glucoside formation
Bönisch F et al (2014) A UDP-glucose monoterpenol glucosyltransferase adds to the chemical diversity of the grapevine metabolome. Plant Physiology 165: 561-581 Gunata Z et al (1985) The aroma of grapes. Journal of Chromatography 331: 83-90 Williams PJ et al (1982) Use of C13 norisoprenoids as natural wine flavour precursors. ACS Symposium Series 388: 35-48
Glycosidic Precursor Synthesis
A gobelet head arm on Vitis vinifera cv. Mourvèdre in Bandol, France, one of four arms emerging from head at 40cm trunk height, extending 32cm outward at 55° angle from vertical, bearing 5 two-bud spurs, arms arranged radially creating characteristic gobelet profile with open center allowing light penetration and air circulation, total vine structure resembling inverted cone or goblet approximately 70cm diameter at top tapering to 15cm diameter head, combined four arms with 20 total spurs supporting 36-38 shoots and 3.4kg crop in classic gobelet architecture, representing traditional Bandol training maintained for centuries and now protected under AOC regulations requiring gobelet or similar traditional low-yielding systems.
A head-trained arm specifically configured in traditional gobelet (goblet or goblet-shaped) architecture, characterized by multiple short arms (typically 3-5) radiating upward and outward from compact central head at angles creating cup or goblet profile, spur positions distributed along arms creating compact fruiting zone, and classical training form employed in many Mediterranean wine regions representing centuries of viticultural tradition emphasizing low yields, hand operations, and terroir expression. Gobelet head arms represent the archetypal head-training configuration, with the gobelet name (French for goblet, referring to the cup-like shape when viewed from above or side) identifying this specific architectural pattern among various head-training approaches, particularly associated with Southern Rhône, Languedoc-Roussillon, Spain, and other warm Mediterranean regions where traditional low-intervention viticulture persists in prestigious appellations and heritage vineyards.
Gobelet head arms exemplify traditional Mediterranean viticulture, with the gobelet form optimized for hand operations (compact accessible structure), drought tolerance (low leaf area and crop reducing water demand), heat management (open center allowing air circulation, reducing heat stress), and quality focus (low yields from limited spur numbers). The gobelet architecture requires no trellis support (vines are self-supporting), minimal equipment (hand tools for pruning and cultivation), and relatively simple establishment (developing head and radiating arms over 3-4 years). However, gobelet training creates significant modern challenges including incompatibility with mechanical harvest (compact irregular structure unsuitable for equipment), labor-intensive hand operations (all pruning, shoot management, harvest requiring manual work), typically low yields (2-4 kg per vine versus 6-12 kg for trellised cordons), and difficult weed management (equipment access limited by irregular spacing and low canopy). The persistence of gobelet in prestigious regions reflects cultural heritage, regulatory protection, and market recognition of gobelet-trained wines as traditional authentic expressions.
Becherform-Arm
Gobelet-Arm
Pokalform-Arm
cup-trained arm
goblet-trained arm
vase-shaped head arm
Gobelet Head Arm
Gobelet suitability quality comparison in Languedoc-Roussillon evaluated traditional bush vine performance for Carignan and Grenache under the dry, windy Mediterranean conditions characteristic of the region. Old-vine Carignan demonstrated excellent gobelet suitability quality with moderate vigor well-matched to the dryland conditions, stiff upright shoots creating stable self-supporting structure, excellent spur fruitfulness maintaining productivity from the short arm positions, and concentrated fruit with characteristic savory complexity attributed to the stress conditions and old vine age. Grenache showed good gobelet suitability quality with similar structural adaptation and spur productivity, though with tendency toward more sprawling architecture requiring occasional positioning in vigorous years. Mourvèdre exhibited marginal gobelet suitability quality with excessive vigor under any but the most limiting conditions creating unstable, lodging-prone structures and requiring conversion to trellised Cordon de Royat training on sites with adequate moisture, demonstrating that gobelet suitability quality depends critically on the moderate vigor expression achievable under limiting conditions.
Gobelet suitability quality is a trellis adaptation quality that inheres in a grapevine cultivar or clone and represents the degree of compatibility with gobelet (bush vine, head-trained) systems characterized by free-standing vines without trellis support trained to a short trunk with multiple short arms bearing spur positions from which shoots grow upward and outward without mechanical guidance, creating compact, self-supporting canopy structures traditionally employed in Mediterranean climates where low vigor conditions and wind exposure favor the protected, low-profile architecture of bush vine training. This quality determines whether a cultivar possesses the structural characteristics, moderate vigor expression, and fruiting habit suitable for the unique constraints and opportunities of unsupported vine training where shoots must develop self-supporting architecture without catch wires and where fruit positioning within the compact canopy influences quality development. The ontological significance of gobelet suitability quality lies in its role enabling continuation of traditional viticultural practices in regions where bush vine training represents cultural heritage with specific quality associations, and in supporting low-input viticulture approaches where trellis infrastructure costs and maintenance requirements are minimized through self-supporting vine architecture. The biological requirements for high gobelet suitability quality include moderate vigor avoiding excessive shoot extension that would create unstable, lodging-prone canopy structures, appropriate shoot stiffness providing mechanical self-support without trellis guidance, internode lengths creating compact rather than sprawling architecture, spur fruitfulness enabling productive pruning to the short arm structures characteristic of gobelet training, and cluster characteristics suited to the interior positioning and variable exposure conditions within bush vine canopies. Cultivars with excessive vigor produce unstable gobelet structures requiring conversion to trellised systems, while those with trailing or weak shoot habit fail to create the upright-outward architecture essential for functional bush vine canopy organization. The gobelet environment creates distinct fruit zone conditions with clusters positioned among the upward-radiating shoots experiencing variable exposure depending on position, with exterior clusters receiving substantial direct radiation while interior clusters remain shaded by surrounding foliage, requiring cultivar adaptation to this heterogeneous microenvironment. Assessment of gobelet suitability quality involves evaluation of vine structural stability, canopy architecture development, fruit positioning and exposure patterns, and yield-quality outcomes achieved without trellis support.
Gobelet_Suitability_Quality is_a Trellis_Adaptation_Quality, representing compatibility with unsupported bush vine training systems. This class addresses the specific requirements of free-standing vine architecture including moderate vigor, shoot stiffness, compact growth habit, and spur fruitfulness. Distinguished from all trellised systems by the absence of support infrastructure. Most relevant for Mediterranean climates with dry, windy conditions favoring the protected bush vine architecture, and for heritage vineyard preservation. The moderate vigor requirement under limiting conditions represents key suitability determinant.
Buscherziehungs-Tauglichkeit
Gobelet-Eignung
Kopferziehungs-Kompatibilität
bush vine suitability
free-standing vine compatibility
freistehende Rebstock-Eignung
goblet training fitness
head training adaptation
Carbonneau A (2002) Grapevine training systems. In: Roubelakis-Angelakis KA (ed) Molecular Biology and Biotechnology of the Grapevine, Kluwer Academic Publishers, pp. 1-34 Huglin P, Schneider C (1998) Biologie et écologie de la vigne. Lavoisier, Paris, pp. 253-278 van Leeuwen C, Seguin G (2006) The concept of terroir in viticulture. Journal of Wine Research 17:1-10
Gobelet Suitability Quality
A Grenache vine in Châteauneuf-du-Pape's galets roulés (rolled stones) terroir bears a Gobelet_Trained_Role, its five arms rising 40 cm from a twisted old trunk, each arm carrying two spurs—the traditional form adapted to Mistral winds exceeding 100 km/h and summer temperatures regularly exceeding 35°C. A Monastrell (Mourvèdre) vine in Spanish Jumilla serves in a Gobelet_Trained_Role for dryland viticulture receiving <300 mm annual rainfall, the compact form concentrating the vine's limited resources into low yields of intensely flavored fruit. A Nero d'Avola vine in Sicilian alberello (Italian equivalent to gobelet) training bears a Gobelet_Trained_Role reflecting centuries of Mediterranean viticultural tradition, the bush form enabling cultivation on steep hillsides without trellis infrastructure.
A head-trained vine role that inheres in a grapevine trained to the formal gobelet (French: "goblet" or "cup") system, with a short trunk (30-60 cm) supporting 3-8 short arms radiating outward and upward in a cup or vase shape, each arm terminating in spur positions bearing the annual fruiting shoots. Gobelet training represents the classical Mediterranean training system, developed over millennia for hot, dry, windy conditions where the compact, self-shading canopy protects fruit from sunburn and desiccation while the sturdy, low structure resists wind damage without trellis support. Spur pruning on permanent arms is standard, with 1-2 buds retained per spur position to limit crop load and maintain vine balance under dryland conditions. Gobelet-trained vineyards typically feature wide spacing (1.5-2.5 m between vines, similar or wider between rows) to accommodate the spreading canopy and enable inter-row cultivation for weed control in traditional systems. The gobelet form maximizes vine self-sufficiency—no trellis investment, natural canopy ventilation, and adaptation to limited water availability—but requires hand labor for all operations, limiting adoption in high-labor-cost environments.
Gobelet_Trained_Role captures the formal, structured version of head training with defined arm and spur positions, distinguishing it from the less formal bush vine category. Regional terminology varies—gobelet (France), alberello (Italy), en vaso (Spain), bush vine (Australia, South Africa)—with slight differences in execution but similar fundamental architecture. Gobelet training creates a distinctive vineyard aesthetic, with individual vine "bushes" creating a landscape of separated plants rather than the continuous hedgerows of trellised systems. The system is particularly associated with old-vine designations, as many gobelet vineyards date from pre-mechanization eras and have survived because their traditional training and low yields produce valued fruit character. Conversion pressures favor trellising for new plantings, but quality-focused producers increasingly recognize gobelet's contributions to fruit concentration, terroir expression, and sustainable dryland production, motivating maintenance of traditional gobelet vineyards.
Becherform
Gobelet-Erziehung
Kelchform
Vasenform
cup training
gobelet
goblet training
vase training
Huglin P, Schneider C (1998) Biologie et écologie de la vigne, Lavoisier Galet P (2000) General Viticulture, Oenoplurimedia Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Dry PR (2000) Canopy management for fruitfulness, Australian Journal of Grape and Wine Research 6: 109-115
Gobelet Trained Role
Thompson Seedless bears a Golden_Raisin_Scion_Role when processed with SO₂ treatment during controlled dehydration, producing "Golden Seedless Raisins" with characteristic light amber-gold color, softer texture (typically dried to 14-16% moisture versus 12-14% for natural), and sweeter, less caramelized flavor than natural raisins—commanding retail premiums of 20-50% above natural raisin prices. Sultana in Australian production serves in a Golden_Raisin_Scion_Role for export markets preferring light-colored dried fruit, with sulfur treatment and mechanical dehydration producing consistent golden product for European and Asian markets. Muscat Gordo Blanco (Muscat of Alexandria) bears a Golden_Raisin_Scion_Role in Australian production, where its distinctive muscat flavor combines with golden color processing to create specialty raisins for premium applications.
A raisin production scion role that inheres in a grafted scion cultivar by virtue of its suitability for golden (sulfured) raisin production, where grapes are treated with sulfur dioxide (SO₂) before or during dehydration to prevent enzymatic browning and preserve the original light golden-amber color of the fresh grape. Golden raisin scions produce berries whose flesh and skin develop attractive golden-to-amber tones when dried under sulfur treatment, rather than the dark brown of oxidized natural raisins. Sulfur dioxide application (typically 1500-3000 ppm exposure during drying) inactivates polyphenol oxidase, preventing melanin formation and preserving the light color valued in certain culinary applications and markets. Golden raisins are typically dehydrated in controlled-temperature facilities (tunnel dryers, dehydrators) rather than sun-dried, enabling precise moisture control and consistent color development. The golden raisin category represents a smaller but premium-priced market segment compared to natural raisins, valued for visual appeal in baked goods, salads, and applications where light color is preferred.
Golden_Raisin_Scion_Role captures a processing-defined category where the same varieties (primarily Thompson Seedless/Sultana) serve in different product roles depending on drying treatment. Sulfur dioxide use is essential for golden color preservation but raises health concerns for sensitive individuals (particularly asthmatics), requiring labeling and limiting use in organic production where golden-style products are achieved through rapid mechanical dehydration minimizing oxidation. The distinction between Natural and Golden scion roles reflects market segmentation rather than genetic differentiation, with production economics (labor costs for sun-drying versus capital costs for dehydration facilities) and market preferences determining variety allocation between categories.
Goldrosinen-Traube
bleached raisin variety
geschwefelte Rosinentraube
golden raisin grape
helle Rosinentraube
light raisin grape
sulfured raisin grape
Christensen LP (2000) Raisin Production Manual, UC ANR Publication 3393 Creasy GL, Creasy LL (2009) Grapes, CABI Karabulut OA et al. (2002) Control of postharvest diseases of grapes with sulfur dioxide, Postharvest Biology and Technology 26: 219-225 Food and Drug Administration (2020) Code of Federal Regulations, Title 21, Sulfites in Dried Fruit
Golden Raisin Scion Role
The GPS coordinates for the centroid of the Clos de Vougeot Grand Cru vineyard (Burgundy) are recorded as 47.1747°N, 4.9561°E (WGS84 datum, ±5 m precision from differential GPS survey) — these coordinates enable the vineyard to be precisely located within the 1:25,000 topographic map series (IGN carte 3024 OT), the 1:50,000 geological map (BRGM carte géologique Nuits-Saint-Georges), the Burgundy high-resolution digital elevation model (0.5 m LiDAR-derived DEM providing slope and aspect at sub-parcel resolution), and the Landsat/Sentinel-2 satellite imagery time series used for monitoring vine phenological development, canopy vigor, and water stress across the Côte de Nuits appellations. The GPS boundary survey of Clos de Vougeot (approximately 120 waypoints defining the 50.6 ha perimeter) provides the definitive digital delineation of this historically important vineyard, replacing the traditional cadastral plans that were the previous spatial reference.
A site description component that records the precise geographic position of the vineyard or specific sampling points within the vineyard, expressed as latitude and longitude coordinates in the WGS84 (World Geodetic System 1984) reference datum with a specified precision level (typically ±1-5 m for differential GPS, ±3-10 m for consumer-grade GPS), providing the georeferenced spatial anchor that enables integration of vineyard-specific data with satellite imagery, digital terrain models, soil maps, climate grids, and other geospatial datasets used in terroir characterization and precision viticulture.
GPS_Coordinates provides the fundamental spatial reference that enables all modern geospatial analysis of vineyard terroir. The transition from traditional paper-based vineyard mapping to GPS-referenced digital mapping has transformed terroir research by enabling the precise co-registration of multiple environmental data layers — soil chemistry, soil depth, slope gradient, solar radiation, temperature, humidity, and vine performance data can all be spatially registered to the same coordinate system, enabling multivariate spatial analysis that reveals the environmental drivers of within-vineyard variation in vine behavior and wine quality: this capability is the technological foundation of precision viticulture, which uses GPS-referenced data to create management zones within vineyards that receive tailored irrigation, nutrition, and harvest timing based on the spatial pattern of vine needs rather than uniform whole-vineyard management. The precision of GPS coordinates is important for different applications — vineyard boundary surveys require sub-meter accuracy (achievable with RTK-GPS or differential GPS), precision viticulture soil and yield mapping requires 1-3 m accuracy (achievable with DGPS), and regional terroir mapping requires only 10-30 m accuracy (achievable with consumer-grade GPS) — and the VVO documents the coordinate precision alongside the coordinates themselves to enable appropriate use of the spatial data.
GPS-Koordinaten
Weinberg-Georeferenzierung
geographic coordinates
geographische Koordinaten
spatial position record
vineyard georeferencing
Bramley RGV (2005) Understanding variability in winegrape production systems, Australian Journal of Grape and Wine Research 11: 33-42 Jones GV et al. (2004) Spatial analysis of climate in winegrape growing regions, American Journal of Enology and Viticulture 55: 304-316 Arnó J et al. (2009) Review — precision viticulture, Australian Journal of Grape and Wine Research 15: 89-108
GPS Coordinates
Gradient coloration development in Cabernet Franc during véraison follows a characteristic pattern where anthocyanin accumulation initiates at the berry shoulder (pedicel attachment region) and progresses toward the stylar end over 5-8 days, creating a diagnostic red-to-green gradient that allows vineyard managers to track ripening progression without destructive sampling the gradient typically spans 3-4 mm in the transition zone during mid-véraison, narrowing to less than 1 mm as coloration approaches uniformity, with persistent strong gradients at normal harvest date indicating delayed or problematic ripening requiring investigation.
Gradient_Coloration_Quality represents the color distribution condition where coloration changes continuously and progressively across the surface of a grapevine organ, creating smooth transitions from one color state to another without discrete boundaries. Gradients may involve changes in pigmentation intensity (light to dark), hue (e.g., green to red), or both dimensions simultaneously. In grape berries, gradient coloration commonly appears as the shoulder-to-tip transition where the exposed berry shoulder displays more intense coloration than the shaded stylar end, a pattern reflecting differential light exposure and temperature across the berry surface. This shoulder gradient is particularly evident during véraison when coloration progresses from the pedicel attachment region toward the berry apex. Radial gradients occur from exposed to shaded faces of individual berries, with the sun-exposed hemisphere developing stronger pigmentation. In leaves, gradient coloration appears during senescence as color change progresses from margins toward the midrib or from interveinal regions toward veins, reflecting the spatial pattern of chlorophyll degradation and anthocyanin synthesis. Nutrient deficiency symptoms often display gradient patterns, as with potassium deficiency where marginal necrosis grades into yellow chlorosis and then green tissue toward the midrib. Gradient coloration assessment requires characterization of the gradient direction (the axis along which color changes), gradient magnitude (the difference between extremes), and gradient linearity (whether the transition is uniform or shows inflection points). Image analysis enables quantitative gradient characterization through color profiles along defined transects.
Gradient_Coloration_Quality is distinguished from Variegated_Coloration_Quality by the continuous nature of color transitions versus discrete patches. Gradients may be linear (uniform rate of change along the gradient axis) or non-linear (varying rate of change), and may be unidirectional or radial. The presence of gradients implies spatial heterogeneity in the conditions controlling pigmentation, whether light exposure, temperature, developmental timing, or resource supply. Some degree of gradient is normal during developmental transitions assessment should consider developmental stage when evaluating whether observed gradients indicate normal or problematic conditions.
Gradientenfärbung
abgestufte Pigmentierung
color gradient quality
graduated color
progressive pigmentation
transitional coloration
verlaufende Farbverteilung
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 369-403 Mori K et al. (2007) Loss of Anthocyanins in Red-Wine Grape under High Temperature. Journal of Experimental Botany 58(8): 1935-1945
Gradient Coloration Quality
SO4 (V. berlandieri × V. riparia) demonstrates excellent graft affinity with virtually all V. vinifera cultivars, forming strong unions with Riesling, Chardonnay, Cabernet Sauvignon, and hundreds of other scions without compatibility issues. In contrast, Börner (V. riparia × V. cinerea) shows limited graft affinity with certain cultivars, requiring careful scion selection to avoid weak unions. Dog Ridge (V. champinii) displays variable compatibility, forming excellent unions with table grape cultivars like Thompson Seedless but showing reduced affinity with some wine grape scions. The Muscadinia rotundifolia-derived rootstocks under development for Xiphinema resistance face severe graft incompatibility barriers with V. vinifera due to chromosomal (2n=40 vs 2n=38) and biochemical differences between Muscadinia and Vitis.
A rootstock role that inheres in a grafted rootstock by virtue of its genetically determined capacity to form successful, long-lasting graft unions with scion cultivars, enabling functional vascular connectivity, mechanical strength, and sustained physiological integration between rootstock and scion partners. Graft affinity (also termed graft compatibility) encompasses the complex processes of callus bridge formation, vascular cambium reconnection, xylem and phloem differentiation across the graft interface, and long-term maintenance of functional transport without decline. Compatibility is determined by taxonomic relatedness (intra-generic grafts generally succeeding, inter-generic grafts often failing), cellular recognition mechanisms, wound response coordination, and absence of biochemical incompatibility factors (phenolic oxidation products, peroxidases, incompatible proteins). Graft affinity varies from universal compatibility (successful unions with virtually all V. vinifera cultivars and most Vitis interspecific hybrids) through limited compatibility (successful with some cultivars, problematic with others) to incompatibility (union failure or delayed decline). Incompatibility may manifest as immediate failure (poor callus formation, no vascular connection) or delayed incompatibility (initial success followed by decline after years or decades).
Graft_Affinity_Rootstock_Role is subclassed into Universal_Compatibility, Limited_Compatibility, and Delayed_Incompatibility_Prone categories reflecting the practical importance of compatibility for vineyard establishment and longevity. Graft compatibility in Vitis is generally high compared to other fruit crops (e.g., Prunus, Malus) because all Vitis species share the same chromosome number (2n=38) and are readily interfertile. The major exception is Muscadinia (2n=40), where graft incompatibility with Vitis remains a significant barrier to deploying Muscadinia-derived pest resistance. Compatibility assessment requires long-term field evaluation, as delayed incompatibility may not manifest until 10-20 years after planting.
Veredelungsaffinität-Unterlage
Veredlungsverträglichkeit-Unterlage
graft affinity rootstock
graft compatibility rootstock
graft union rootstock
pfropfkompatible Unterlage
scion compatibility rootstock
Hartmann HT et al. (2011) Hartmann and Kester's Plant Propagation: Principles and Practices 8th Ed, Pearson pp.: 415-460 Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22 Cookson SJ et al. (2013) Graft union formation in grapevine induces transcriptional changes related to cell wall modification, wounding, hormone signalling, and secondary metabolism, Journal of Experimental Botany 64: 2997-3008 Pina A, Errea P (2005) A review of new advances in mechanism of graft compatibility-incompatibility, Scientia Horticulturae 106: 1-11
Graft Affinity Rootstock Role
Chardonnay grafted on 3309 Couderc showing graft callus adventitious root, bench graft with union 5 cm above soil line, field inspection revealing root emerging from graft union callus, excavation exposing union root 4.2 cm length originating from callus at scion-rootstock interface, molecular analysis using microsatellite markers identifying root as scion-derived (Chardonnay genotype not 3309C), root removed to maintain rootstock function and prevent phylloxera risk, graft union root demonstrating need for proper union height and monitoring
Graft callus adventitious root is callus-derived adventitious root arising from callus tissue at graft union between scion and rootstock, characterized by formation from callus proliferating between grafted partners as they heal and unite, showing origin from either scion callus, rootstock callus, or intermixed callus at graft interface, demonstrating variable occurrence with some grafts developing union roots while others showing none, and serving functions including graft union mechanical reinforcement, potential vascular bridging between partners, and occasionally complicating graft relationships by allowing scion to root independently, with graft callus roots showing typical development from parenchymatous callus cells at union organizing into root primordia and emerging from union periphery, exhibiting anatomical pathway requiring penetration through callus and potentially both scion and rootstock tissues to emerge, and demonstrating practical significance in viticulture where graft union roots occurrence affects vine management particularly if scion roots compromise rootstock functions such as phylloxera resistance or vigor control. Graft callus adventitious root formation context includes= showing graft union callus (callus forming at interface between scion and rootstock as part of graft healing process), exhibiting intermingled tissue (scion and rootstock callus cells mixing at union creating composite tissue), demonstrating variable occurrence (graft union roots forming in some but not all grafts depending on multiple factors), revealing rootstock effects (some rootstock varieties or graft types more prone to union root formation than others), and displaying environmental influences (union root formation potentially affected by soil moisture, depth, and other environmental factors at union). Graft callus adventitious root origin ambiguity includes= showing scion-derived possibility (union roots potentially originating from scion-side callus cells), exhibiting rootstock-derived possibility (roots potentially from rootstock-side callus cells), demonstrating mixed origin (some union roots potentially from intermixed scion-rootstock callus), revealing identification difficulty (determining root origin requiring molecular markers or careful anatomical study), and displaying practical ambiguity (uncertain origin affecting management decisions particularly regarding removal). Graft callus adventitious root in union stability includes= providing mechanical reinforcement (union roots potentially strengthening graft union mechanical connection), exhibiting anchorage contribution (roots from union potentially providing additional vine anchorage), demonstrating union bridging (roots growing through union potentially physically connecting scion and rootstock), revealing vascular enhancement (union roots potentially contributing to vascular continuity between partners), and displaying stabilization function (union roots helping secure graft partners particularly in young or weak unions). Graft callus adventitious root problematic aspects includes= enabling scion rooting (if union roots originate from scion callus allowing scion to develop own root system), compromising rootstock function (scion roots bypassing rootstock potentially eliminating intended rootstock benefits), demonstrating phylloxera risk (scion roots from susceptible V. vinifera providing pathway for phylloxera attack), revealing vigor effects (scion roots potentially changing vine vigor if rootstock selected for vigor control), and displaying management necessity (union roots requiring monitoring and potentially removal maintaining graft relationships). Graft callus adventitious root detection and identification includes= showing visual inspection (union roots sometimes visible at graft union or emerging nearby), exhibiting excavation (careful soil removal around union revealing union roots if present), demonstrating anatomical examination (microscopic study potentially determining root origin based on anatomy), revealing molecular approaches (DNA markers potentially identifying whether roots originated from scion or rootstock), and displaying field indicators (vine behavior changes potentially suggesting scion rooting through union). Graft callus adventitious root prevention and management includes= showing proper graft height (grafting above soil line reducing union burial and root formation risk), exhibiting soil management (avoiding excessive soil near union preventing burial stimulating root formation), demonstrating suckering attention (removing suckers potentially preventing conditions favoring union roots), revealing inspection protocols (regular union examination detecting and allowing removal of union roots), and displaying removal techniques (carefully removing union roots if detected maintaining rootstock function). Graft callus adventitious root in different graft types includes= showing bench graft occurrence (callused bench grafts potentially developing union roots during callusing or nursery growth), exhibiting field graft patterns (field grafts showing different union root frequency than bench grafts), demonstrating cleft versus whip differences (graft type potentially affecting union callus production and root formation), revealing compatibility effects (poor compatibility sometimes associated with excessive callus and union root formation), and displaying rootstock variation (some rootstock varieties showing higher union root frequency than others). Graft callus adventitious root research and monitoring includes= requiring systematic surveys (documenting union root frequency across cultivar-rootstock combinations), exhibiting molecular identification (using genetic markers determining root origin in union root populations), demonstrating functional assessment (testing whether union roots compromise rootstock functions), revealing management evaluation (assessing effectiveness of practices aimed at preventing or removing union roots), and displaying breeding implications (considering union root tendency in rootstock variety selection and breeding).
Graft_Callus_Adventitious_Root emphasizes graft union callus origin and is distinguished from cutting roots by focus on graft interface context. Critical for understanding union callus, scion-rootstock interface, mechanical reinforcement, problematic scion rooting, rootstock function compromise, and the adventitious roots arising from graft union callus potentially affecting graft relationships and rootstock benefits.
Edelreis-Unterlage-Wurzel
Graft interface root
Pfropf-Kallus-Adventivwurzel
Scion-rootstock callus root
Union callus root
Unions-Kallus-Wurzel
Graft Callus Adventitious Root
Graft compatibility in commercial viticulture is generally high among V. vinifera scions and standard rootstocks, with take rates of 85-98% for most combinations under optimal nursery conditions. Compatibility is assessed by callus bridge formation (visible within 14-21 days), vascular differentiation (28-42 days), and mechanical strength of the union (3-6 months). Compatible unions show continuous cambial activity, aligned vascular elements, and absence of necrotic tissue at the interface. Incompatibility symptoms include poor callus development, discontinuous cambium, excessive starch accumulation above the union, and delayed or absent vascular connections, with affected vines showing reduced vigor, leaf discoloration, and premature decline.
A compatibility disposition that is the inherited capacity of a Vitis vinifera scion to form and maintain a functional graft union with rootstock genotypes, determined by the ability to establish cellular continuity, vascular connections, and physiological integration across the graft interface without adverse incompatibility reactions.
Graft_Compatibility_Disposition serves as the parent class for graft union formation and graft incompatibility traits. Compatibility operates at multiple levels including cellular (callus intermingling), vascular (xylem and phloem connection), and physiological (hormone and metabolite exchange). Most V. vinifera cultivars show broad compatibility with standard rootstocks, though specific combinations may exhibit marginal or delayed incompatibility. Temperature, humidity, and grafting technique significantly influence compatibility expression.
Pfropfaffinität
Pfropfkompatibilität
Veredlungsverträglichkeit
graft success trait
grafting compatibility
scion-rootstock compatibility
Pina A, Errea P (2005) A review of new advances in mechanism of graft compatibility-incompatibility. Scientia Horticulturae 106:1-11 Cookson SJ et al. (2013) Graft union formation in grapevine induces transcriptional changes related to cell wall modification, wounding, hormone signalling, and secondary metabolism. Journal of Experimental Botany 64:2997-3008
Graft Compatibility Disposition
Graft incompatibility in grapevine ranges from immediate failure (0-20% graft take) in severe cases to delayed decline emerging 5-15 years after grafting in marginal combinations. Incompatibility symptoms include necrotic tissue at the graft interface, discontinuous cambial zones, bark involution, excessive starch accumulation in scion tissue above the union, reduced hydraulic conductance (30-60% of compatible unions), and vine decline with premature senescence. The combination of V. vinifera with V. rotundifolia (Muscadinia) shows severe incompatibility with <30% initial survival and universal long-term failure. Within V. vinifera × standard rootstock combinations, incompatibility is rare but reported for specific clonal combinations, with incidence of 2-8% showing measurable impairment.
A graft compatibility disposition that is the inherited capacity of certain Vitis vinifera scion-rootstock combinations to exhibit impaired graft union function despite initial callus contact, resulting from adverse cellular, biochemical, or physiological interactions at the graft interface that prevent normal union development or cause progressive functional decline.
Graft_Incompatibility_Disposition serves as the parent class for translocated and localized incompatibility mechanisms. Incompatibility represents failure of the complex biological processes required for successful grafting, potentially involving any stage from initial cell recognition through long-term vascular function. The phenomenon is less prevalent in grapevine than in Prunus and other genera but remains a consideration in rootstock selection and nursery quality control. Climate stress may exacerbate marginal incompatibility that would remain subclinical under favorable conditions.
Abstoßungsreaktion
Pfropfinkompatibilität
Veredlungsunverträglichkeit
graft rejection
incompatibility trait
union dysfunction
Pina A, Errea P (2005) A review of new advances in mechanism of graft compatibility-incompatibility. Scientia Horticulturae 106:1-11 Mudge K et al. (2009) A history of grafting. Horticultural Reviews 35:437-493
Graft Incompatibility Disposition
Cabernet Sauvignon on 3309C rootstock graft union at 8 years post-grafting shows 12-18 mm vertical zone positioned 10-14 cm above soil line, circumferential swelling 112-118% of adjacent trunk diameter creating slight bulge, healed union comprising callus bridge tissue with differentiated continuous cambium producing annual xylem and phloem rings connecting scion and rootstock vascular systems, functional xylem vessels enabling 15-35 L/day water transport and phloem sieve tubes enabling 8-18 g/day photosynthate basipetal flow, union covered by protective periderm 4-6 mm bark thickness, mechanical strength 75-85% of non-grafted trunk, no visible incompatibility symptoms
Graft union is the junction zone in grafted Vitis vinifera trunks where scion and rootstock tissues have been joined through grafting, comprising the original cut surfaces, proliferated callus tissue, and differentiated vascular connections, representing the permanent anatomical discontinuity and functional bridge enabling scion-rootstock integration in grafted vines. Graft union formation process includes= initial tissue contact (cut scion and rootstock surfaces placed together typically in dormant season February-March), callus proliferation (undifferentiated parenchyma cells arising from cambial region and adjacent tissues at cut surfaces within 1-3 weeks), callus bridge formation (scion and rootstock callus tissues meeting and intermixing creating continuous tissue bridge across junction), vascular differentiation (new cambium differentiating within callus bridge connecting scion and rootstock vascular systems), and functional integration (xylem and phloem elements establishing continuity enabling water, mineral, and photosynthate transport across union within 4-8 weeks). Graft union location and dimensions includes= position typically 5-15 cm above soil line (bench grafts, field grafts, or top-worked grafts positioning union above ground preventing scion rooting), union zone typically 10-30 mm vertical extent (encompassing original cut surfaces plus callus tissue above and below), circumferential swelling 105-125% of adjacent trunk diameter (graft union often appearing as bulge from extensive callus formation and differential cambial activity), asymmetric development common (union showing uneven cambial activity around circumference creating eccentric appearance), and permanent visibility (graft union remaining identifiable throughout vine life even in successfully healed mature unions). Graft union anatomical components includes= original scion-rootstock interface (plane where tissues first contacted, often visible as line or discontinuity even in old unions), callus tissue (proliferated undifferentiated and partially differentiated parenchyma filling gaps and creating bulk of union tissue), differentiated vascular connections (xylem vessels and phloem sieve tubes bridging across union enabling transport), continuous cambium (vascular cambium established across union producing secondary xylem and phloem maintaining connection), periderm covering (bark tissues forming over union surface protecting internal tissues), and potential incompatibility zones (regions of poor cell adhesion, necrotic tissue, or vascular discontinuity in unsuccessful or delayed unions). Graft union vascular continuity includes= xylem connections (secondary xylem vessels differentiating in callus bridge establishing water transport pathways from rootstock to scion), phloem connections (sieve tubes and companion cells forming in callus enabling photosynthate transport from scion to rootstock), ray continuity (radial parenchyma potentially connecting across union facilitating lateral transport), cambial continuity (functioning vascular cambium spanning union producing annual xylem and phloem increments maintaining connection strength), and functional integration (successful union showing normal bidirectional transport rates approaching non-grafted trunk transport capacity). Graft union cambial activity and growth includes= differential cambial vigor (scion and rootstock often showing different inherent growth rates), union swelling development (more vigorous partner producing excess callus and wood creating characteristic bulge 110-130% of trunk diameter at union), asymmetric secondary growth (cambium more active on certain sides creating eccentric annual rings and irregular circumference), gradual moderation (initial pronounced swelling often becoming less dramatic with age as growth rates equilibrate), and persistent enlargement (union typically maintaining 105-120% swelling throughout vine life though occasionally approaching uniform diameter in very old successfully integrated unions). Graft union strength and mechanical properties includes= initial weakness (newly formed union with only callus tissue showing limited strength requiring support during first growing season), developing strength (vascular differentiation and secondary growth increasing union strength over 2-4 years), mature union strength (successful 5+ year unions typically achieving 70-90% of non-grafted trunk breaking strength), failure modes (union fractures typically occurring at scion-rootstock interface or through incompatibility zones under excessive bending or torsional stress), and vulnerability factors (incompatibility, inadequate callus formation, pathogen invasion, or freeze damage potentially compromising union strength creating permanent weak points). Graft union healing assessment includes= early-stage evaluation (6-12 weeks post-grafting, assessing callus proliferation, tissue contact, and absence of necrosis or separation), intermediate evaluation (6-18 months, examining vascular differentiation, cambial continuity, absence of incompatibility symptoms), mature evaluation (3-5+ years, assessing functional transport capacity, mechanical strength, absence of delayed incompatibility), visual indicators (smooth healed appearance, minimal swelling, continuous bark, uniform growth above and below union indicating successful integration), and failure indicators (excessive swelling >130%, necrotic zones, bark separation, growth disparities, decline symptoms suggesting incompatibility or inadequate healing). Graft union types by grafting method includes= whip-and-tongue unions (interlocking tongue configuration promoting alignment and stability, common in bench grafting), omega unions (machine-cut interlocking profile providing precise fit, standard in commercial propagation), cleft unions (split rootstock receiving wedge-shaped scion, used in field grafting mature vines), approach unions (growing plants grafted together then separated after union forms, occasional in vine grafting), and top-working unions (established vine trunk receiving new scion variety, positioned higher 30-100 cm above ground). Graft union position effects includes= above-ground positioning (5-15 cm above soil preventing scion adventitious rooting maintaining rootstock influence on vigor, disease resistance, soil adaptation), height variation (bench grafts 8-15 cm, field grafts 10-20 cm, top-worked grafts 30-100+ cm affecting accessibility and management), soil coverage risk (union positioned too low potentially becoming buried allowing scion rooting negating rootstock benefits), frost exposure (above-ground union vulnerable to freeze damage in severe winters particularly in young vines with thin bark), and management accessibility (visible union enabling inspection, assessment, and intervention if problems develop). Graft union incompatibility manifestations includes= immediate incompatibility (union failure to form adequate callus bridge causing graft death within weeks to months), delayed incompatibility (initially successful union developing problems years later showing declining vigor, vascular dysfunction, or mechanical failure), localized incompatibility (portions of union showing poor integration while other sections function normally creating weak zones), and compatibility (successful unions showing complete integration, normal growth, and absence of symptoms maintaining function throughout vine life potentially 30-60+ years). Graft union pathogen vulnerability includes= union as infection site (wounds and tissue discontinuities at union providing entry points for trunk disease pathogens), Eutypa dieback (fungal pathogen potentially entering through union wounds or cracks), Esca complex (union injuries or incompatibility zones potentially facilitating fungal colonization), Phomopsis (canker pathogen potentially infecting union particularly in first years before bark fully protective), crown gall (Agrobacterium potentially infecting union wounds causing gall formation), and protection importance (maintaining union integrity, proper healing, protective covering minimizing pathogen entry risk). Graft union environmental stress sensitivity includes= freeze damage vulnerability (union tissue heterogeneity creating differential cold hardiness, severe cold potentially killing cambium at union), sunscald risk (southwest exposure on young unions potentially causing tissue death from radiant heating and freezing), drought stress effects (water deficit potentially exacerbating incompatibility symptoms or slowing union healing), heat stress (excessive temperature potentially damaging developing union in first growing season), and protection strategies (wrapping, shading, or painting young unions improving survival particularly in extreme climates). Graft union significance in viticulture includes= enabling rootstock benefits (phylloxera resistance, nematode tolerance, soil adaptation, vigor control achieved through grafted vine with functional union), determining vine longevity (successful union supporting 30-60+ year vine life, poor union causing premature decline or death within 5-15 years), affecting vine performance (union quality influencing vigor, yield, fruit quality through impact on transport efficiency and stress tolerance), requiring skilled propagation (union success depending on precise grafting technique, timing, aftercare determining establishment rates 70-95%), and influencing vineyard management (union position, condition affecting cultivation practices, trunk disease management, vine replacement decisions).
Graft_Union emphasizes complete scion-rootstock junction and is distinguished from specific union components (Graft_Union_Zone, Callus_Tissue) and union problems (Incompatibility_Zone) by encompassing entire junction structure. Critical for understanding grafted vine anatomy, scion-rootstock integration, vascular continuity establishment, union healing process, incompatibility detection, mechanical strength development, and graft union as permanent anatomical feature determining grafted vine function, longevity, and performance.
Graft junction
Grafting zone
Pfropfstelle
Scion-rootstock union
Veredelungszone
Veredlungsstelle
Graft Union
Graft union formation in grapevine omega grafts proceeds through defined stages over 21-35 days under optimal conditions (25-28°C, 85-95% RH). Callus initiation occurs within 3-7 days, with callus cells from scion and rootstock intermingling by day 10-14 to form a callus bridge. Vascular differentiation begins at day 14-21, with first continuous xylem vessels detectable by day 21-28 and phloem connections by day 28-35. Mechanical union strength increases from near zero at grafting to 50-100 N pull-apart force by 6 weeks, reaching 200-400 N by 12 weeks in compatible unions. Bench graft callusing success rates of 90-98% are achieved in climate-controlled callusing rooms versus 70-85% under field conditions.
A graft compatibility disposition that is the inherited capacity of a Vitis vinifera scion and rootstock combination to establish the initial graft union through coordinated wound response, callus proliferation, cellular adhesion, and de novo vascular differentiation connecting the previously separate vascular systems of the graft partners.
Graft_Union_Formation_Disposition serves as the parent class for callus formation and vascular connection traits. Union formation represents the critical establishment phase where compatibility or incompatibility first manifests. The process requires coordinated wound responses from both graft partners, with auxin and cytokinin signaling driving callus proliferation and vascular redifferentiation. Successful union formation does not guarantee long-term compatibility, as delayed incompatibility may emerge years after initial graft take.
Kallusbrückenbildung
Pfropfvereinigung
Veredlungsstellen-Bildung
graft healing capacity
initial compatibility trait
union establishment
Pina A et al. (2012) Differential induction of phenylalanine ammonia-lyase gene expression in response to in vitro callus unions of Prunus spp. Journal of Plant Physiology 169:1724-1730 Melnyk CW et al. (2015) A developmental framework for graft formation and vascular reconnection in Arabidopsis thaliana. Current Biology 25:1306-1318
Graft Union Formation Disposition
Merlot on SO4 rootstock graft union zone at 5 years post-grafting shows 14-22 mm vertical extent positioned 11-15 cm above soil, union zone comprising callus bridge tissue (gradually being replaced by secondary growth), continuous vascular cambium producing coordinated annual rings in scion and rootstock, functional xylem connections enabling 18-32 L/day water transport (85-92% of non-grafted equivalent capacity), phloem continuity supporting 10-16 g/day photosynthate basipetal flow, circumferential swelling 108-116% indicating well-balanced integration, microscopic examination revealing organized vascular tissues with vessel elements 140-240 μm diameter crossing interface, no necrotic zones or discontinuities
Graft union zone is the specific tissue region at graft union where scion and rootstock tissues directly contact and integrate, comprising the callus bridge, differentiated vascular connections, and transitional tissues spanning the original scion-rootstock interface, representing the functional bridge enabling physiological continuity between genetically distinct plant parts. Graft union zone boundaries and extent includes= vertical extent typically 8-25 mm (encompassing tissue 3-12 mm above and below original cut plane), inner boundary at pith or early secondary xylem (union zone affecting all tissues from trunk center to bark surface), outer boundary at bark surface (periderm covering union protecting internal tissues), circumferential extent (complete cylinder around trunk circumference though integration quality potentially varying around periphery), and temporal changes (union zone initially comprising primarily callus gradually incorporating more differentiated tissues as secondary growth proceeds over years). Graft union zone tissue composition includes= callus bridge tissue (undifferentiated and partially differentiated parenchyma cells forming initial connection, comprising 40-70% of union zone volume in first 2-3 years), differentiated xylem (secondary xylem vessels, fibers, and parenchyma establishing water transport continuity across interface), differentiated phloem (sieve tubes, companion cells, and phloem parenchyma enabling photosynthate transport), continuous cambium (vascular cambium spanning union producing additional xylem and phloem strengthening connection), rays potentially continuous (radial parenchyma connecting across union facilitating lateral transport), and bark covering (periderm protecting union zone surface). Graft union zone vascular differentiation process includes= initial callus proliferation (undifferentiated cells arising from cambial region at cut surfaces within 1-2 weeks post-grafting), callus intermixing (scion and rootstock callus cells meeting and integrating forming continuous tissue bridge), cambial differentiation (new cambium forming within callus tissue typically 3-6 weeks post-grafting), xylem element formation (vessel elements, fibers differentiating from cambium establishing first water transport pathways 4-8 weeks post-grafting), phloem element formation (sieve tubes and companion cells differentiating enabling photosynthate transport), secondary growth (established cambium producing additional xylem and phloem annually strengthening union and improving transport capacity), and progressive integration (continued growth gradually replacing callus tissue with organized vascular tissues improving union quality over 2-5 years). Graft union zone vascular continuity characteristics includes= xylem connections (vessel elements bridging union typically showing 70-95% connectivity compared to non-grafted trunk depending on healing quality), phloem connections (sieve tube pathways across union enabling functional transport though potentially tortuous in young unions), transport capacity (successful union zone enabling water flow 80-95% and photosynthate transport 75-90% of non-grafted capacity), resistance to flow (union zone creating increased hydraulic resistance 10-40% compared to equivalent non-grafted trunk length depending on connection quality), and improvement with age (vascular continuity increasing over years as secondary growth adds connections and improves organization). Graft union zone cambial activity patterns includes= differential cambial vigor (scion and rootstock cambium often showing different division rates), cambial continuity establishment (continuous cambium forming across union within 4-12 weeks enabling coordinated secondary growth), asymmetric growth (union zone cambium often more active on certain sides creating eccentric rings), swelling development (differential cambial activity producing union bulge 105-125% adjacent diameter), and annual ring formation (cambium in union zone producing annual xylem and phloem rings though potentially discontinuous or irregular in early years gradually improving with age). Graft union zone healing stages includes= initial contact (cut surfaces placed together, stage 0-1 week), callus proliferation (undifferentiated cells arising from both partners, week 1-4), callus fusion (callus tissues meeting and integrating, week 2-6), vascular differentiation (cambium and conducting elements forming, week 4-12), functional integration (established transport pathways, month 3-6), secondary strengthening (continued growth adding tissue, year 1-3), and mature union (well-integrated zone with organized tissues, year 3-10+). Graft union zone in successful unions includes= complete tissue continuity (scion and rootstock tissues fully connected without gaps or necrotic zones), functional vascular connections (xylem and phloem enabling normal transport rates), continuous cambium (producing coordinated secondary growth), strong mechanical integration (union approaching non-grafted trunk strength 70-90%), minimal swelling (circumference 105-115% of adjacent trunk indicating balanced growth), healthy bark covering (protective periderm without cracks or necrosis), and absence of incompatibility symptoms (normal growth, vigor, productivity above and below union). Graft union zone in problematic unions includes= incomplete tissue contact (gaps or discontinuities preventing full integration), necrotic zones (dead tissue regions impeding vascular connections), vascular discontinuities (poor xylem or phloem connectivity reducing transport capacity), cambial discontinuities (non-functional or absent cambium sections preventing strengthening), excessive swelling (>125-150% diameter suggesting severe incompatibility or overgrowth), bark problems (cracks, cankers, or separation indicating union stress), and decline symptoms (reduced growth, vigor, or productivity suggesting inadequate union function). Graft union zone assessment methods includes= visual external examination (observing swelling magnitude, bark condition, symmetry), dissection (cutting through union revealing internal tissue organization, continuity, necrosis), dye perfusion (injecting dyes showing vascular connectivity patterns), microscopy (examining cellular organization, differentiation, tissue adhesion), mechanical testing (measuring union breaking strength, flexibility), physiological measurements (assessing water potential, sap flow rates across union), and long-term monitoring (tracking growth, vigor, symptoms over years detecting delayed incompatibility). Graft union zone in different rootstock combinations includes= highly compatible combinations (e.g., Vitis vinifera scions on V. riparia or V. berlandieri × V. riparia hybrids showing excellent integration, minimal swelling 105-115%, normal function), moderately compatible (e.g., some V. vinifera on V. rupestris hybrids showing adequate but imperfect integration, moderate swelling 115-125%, acceptable function), problematic combinations (incompatible scion-rootstock pairs showing poor tissue adhesion, extensive swelling >130%, vascular discontinuities, eventual decline), and genetic factors (molecular incompatibility mechanisms affecting cell recognition, adhesion, vascular differentiation determining union success). Graft union zone environmental influences includes= temperature during healing (optimal 20-28°C promoting rapid callus formation, 35°C slowing healing), moisture conditions (adequate humidity supporting callus proliferation, excessive drying inhibiting healing), pathogen pressure (wound infections potentially disrupting union formation), mechanical stress (movement at interface preventing callus fusion, support improving success), and seasonal timing (dormant season grafting typically most successful, growing season grafts requiring careful management). Graft union zone significance includes= determining graft success (union zone quality controlling vine establishment, survival 70-95% depending on technique and compatibility), enabling scion-rootstock integration (union zone providing physical and physiological connection), affecting vine performance (union quality influencing transport efficiency, stress tolerance, vigor, productivity), requiring time to develop (union zone progressively improving over 2-5 years reaching mature function), and permanent vine feature (union zone persisting throughout vine life potentially 30-60+ years as site of scion-rootstock connection and potential weak point or incompatibility expression).
Graft_Union_Zone emphasizes specific tissue bridge region and is distinguished from Graft_Union (complete junction), Healed_Graft_Union_Zone (successful integration emphasis), Callus_Graft_Union_Zone (callus tissue emphasis), and Incompatibility_Zone (failed integration) by focus on transitional tissue connecting scion and rootstock. Critical for understanding vascular continuity establishment, tissue differentiation process, healing progression, transport capacity development, union quality assessment, and graft union zone as functional bridge determining grafted vine physiological integration and performance.
Integration zone
Integrations-Gewebe
Junction tissue
Union bridge
Vereinigungszone
Verwachsungsstelle
Graft Union Zone
Pinot Noir grafted on 3309C rootstock at 12 years post-grafting showing grafted rootstock trunk 6.8 cm diameter extending 24 cm from root crown to graft union, graft union positioned 19 cm above soil surface displaying characteristic 2.1 cm diameter bulge with complete callus integration, grafted rootstock trunk bark light grey-brown with fine longitudinal fissures typical of 3309C, no signs of incompatibility (absence of necrosis, cracks, or excessive swelling), scion trunk above graft union 5.9 cm diameter showing normal Pinot Noir bark characteristics
Grafted rootstock trunk is rootstock trunk that has been successfully joined with scion cultivar at graft union, representing the integrated rootstock aerial portion of grafted vine where rootstock tissue supports and connects with scion above, showing complete or progressive vascular and structural integration across graft union, typical of commercial viticulture where nearly all Vitis vinifera wine grapes grow on grafted rootstock for phylloxera resistance and site adaptation. Grafted rootstock trunk integration status includes= having established graft union (successful connection between rootstock and scion tissues at grafting point marked by callus bridge and vascular continuity), showing functional vascular connections (xylem and phloem continuity across graft union enabling bidirectional transport between root system and scion canopy), maintaining structural integrity (mechanical stability of graft union capable of supporting scion weight and stress without separation), potentially displaying graft union bulge (characteristic swelling at grafting point 1.2-3.5 cm diameter increase due to callus tissue and differential growth rates), and expressing compatibility (absence of severe incompatibility symptoms indicating successful rootstock-scion partnership). Grafted rootstock trunk as component of complete vine includes= extending from root system to graft union (spanning the critical 15-40 cm section between root crown and scion connection), providing rootstock contributions to whole vine (phylloxera resistance, vigor control, drought tolerance, lime tolerance, and other rootstock traits affecting complete vine performance), supporting scion tissues above (bearing weight of scion trunk, canopy, and fruit load while conducting resources from roots), enabling specialized viticulture (grafted combination allowing superior scion cultivar to grow on appropriate rootstock in challenging soil or pest conditions), and representing below-graft component (distinct from scion trunk above while forming integrated functional whole). Grafted rootstock trunk developmental stages includes= young grafted stage (years 1-3 post-grafting) showing initial callus formation and establishing vascular connections, establishing grafted stage (years 4-7) demonstrating complete integration and stable graft union, mature grafted stage (years 8-30) exhibiting well-established integration with full rootstock contributions, old grafted stage (years 30-60+) showing long-term stable integration with minimal additional callus formation, and ancient grafted stage (70-100+ years) displaying decades of successful rootstock-scion partnership with thick bark and large diameter. Grafted rootstock trunk graft union characteristics includes= located at grafting height (typically 10-30 cm above soil surface in field grafting, 15-40 cm in bench grafting, positioning graft union above wet soil preventing scion rooting), showing callus tissue (wound healing tissue forming bridge between rootstock and scion at graft union interface), displaying characteristic bulge (swelling at graft union 0.5-3 cm diameter increase from combined callus accumulation and differential growth rates between rootstock and scion), potentially showing graft union scar (visible line or ridge marking original grafting point even decades after grafting), and maintaining distinct tissue zones (rootstock tissues below showing rootstock characteristics while scion above shows scion traits). Grafted rootstock trunk in compatible combinations includes= showing smooth vascular integration (well-connected xylem and phloem across graft union without tissue discontinuities), exhibiting normal growth patterns (rootstock trunk diameter increasing steadily without excessive bulging or constriction), maintaining healthy appearance (absence of necrotic tissue, cracks, or separations at graft union), enabling vigorous vine growth (rootstock providing optimal support and resource transport to scion canopy), and supporting long vine lifespan (stable integration sustaining productive vine for 30-80+ years). Grafted rootstock trunk in incompatible combinations includes= potentially showing excessive bulging (severe swelling at graft union >3.5 cm diameter increase indicating poor integration), displaying tissue discontinuities (gaps or separations in vascular connections reducing transport efficiency), exhibiting constriction above or below union (growth restriction creating weak point in trunk), showing necrotic areas (dead tissue zones at graft interface indicating rejection response), developing cracks or splits (mechanical failures at poorly integrated graft union), and potentially leading to vine decline (incompatibility symptoms progressing to reduced vigor, poor productivity, or vine death). Grafted rootstock trunk transport functions includes= conducting water from root system to scion (xylem vessels moving 10-100+ liters daily during growing season across graft union to canopy), transporting mineral nutrients upward (nitrogen, phosphorus, potassium, calcium, magnesium, and micronutrients absorbed by roots delivered to scion leaves and fruit), moving photosynthates downward (phloem transporting carbohydrates from scion leaves through graft union to rootstock trunk and root system for storage), enabling hormone transport (signaling molecules moving bidirectionally between roots and shoots coordinating growth and development), and maintaining continuous circulation (uninterrupted flow despite graft union interface connecting two distinct genotypes). Grafted rootstock trunk rootstock effects includes= conferring phylloxera resistance (rootstock genetics providing immunity to root form of Daktulosphaira vitifoliae protecting root system while susceptible scion cultivar grows above), controlling vine vigor (rootstock variety determining growth intensity with low vigor rootstocks restricting growth and high vigor rootstocks promoting growth), providing drought adaptation (deep-rooted or drought-tolerant rootstocks improving water stress resistance), enabling lime tolerance (rootstock genetics allowing vine growth in high pH calcareous soils where susceptible scions would develop chlorosis), and contributing nematode resistance (some rootstocks tolerant to root-knot and dagger nematodes protecting vine). Grafted rootstock trunk management considerations includes= maintaining graft union above soil surface (preventing scion rooting which would bypass rootstock effects), protecting from mechanical damage (avoiding equipment injury to graft union which could compromise integration), monitoring for disease (inspecting graft union for crown gall, cankers, or decay), managing rootstock suckers (removing shoots emerging from grafted rootstock trunk below graft union), and assessing compatibility (evaluating graft union health, trunk symmetry, and absence of incompatibility symptoms). Grafted rootstock trunk in vineyard establishment includes= planted at appropriate depth (positioning graft union 10-20 cm above final soil surface), initially supported (staking or trellising protecting vulnerable young grafted trunk and graft union from wind damage), gradually acclimating (young grafted rootstock trunk establishing root system and trunk stability over first 2-4 years), eventually self-supporting (mature grafted rootstock trunk achieving structural independence though still trellised for crop support), and representing standard commercial practice (>95% commercial wine grape vineyards worldwide using grafted vines rather than own-rooted vines due to phylloxera and other benefits). Grafted rootstock trunk cultivar-specific characteristics includes= varying by rootstock variety (SO4, 3309C, 1103P, Riparia Gloire, 101-14, Teleki 5C, Fercal, and dozens of others each showing characteristic bark texture, vigor, and growth patterns), differing in graft compatibility (some rootstocks grafting easily with wide scion range while others showing selective compatibility), showing distinct diameter patterns (vigorous rootstocks like 1103P producing larger diameter grafted trunks than low-vigor rootstocks like Riparia), and displaying recognizable bark (rootstock-specific bark textures potentially identifiable even decades post-grafting). Grafted rootstock trunk significance includes= enabling modern viticulture (grafted rootstock trunks representing solution to phylloxera devastation allowing Vitis vinifera cultivation in infested soils since 1880s), providing site-specific adaptation (appropriate rootstock-scion grafted combinations optimizing performance in diverse soil, climate, and pest conditions), maintaining long-term integration (successful grafted rootstock trunks supporting vines for 50-100+ years demonstrating durability of grafting technology), and representing human intervention (grafted rootstock trunk as artifact of viticulture joining two genotypes creating vine that would not exist naturally).
Grafted_Rootstock_Trunk emphasizes successful integration with scion via graft union and is distinguished from Rootstock_Trunk (which could theoretically refer to ungrafted rootstock) by focus on the grafted state. Critical for understanding rootstock-scion relationships, graft union integrity, rootstock contributions to complete vine, compatibility issues, and the fundamental architecture of commercial grafted grapevines.
Aufgepfropfter Unterlagenstamm
Integrated rootstock trunk
Joined rootstock trunk
United rootstock trunk
Veredelter Wurzelstockstamm
Verpfropfter Unterlagenstamm
Grafted Rootstock Trunk
Cabernet Sauvignon grafted scion trunk on 3309C rootstock at 12 years post-planting shows 62-78 mm diameter at 30 cm above graft union (compared to 68-82 mm rootstock diameter at 10 cm below union), graft union at 8 cm above soil with 110-118% diameter swelling, 12 annual rings above union with scion-specific ring-porous anatomy (earlywood vessels 180-260 μm diameter), and rootstock-mediated moderate vigor producing balanced 4.8-6.2 kg annual yield
A grafted scion trunk is a scion trunk that develops above a graft union where the desired Vitis vinifera cultivar (scion) has been grafted onto a rootstock (typically Vitis species hybrid or Vitis vinifera × non-vinifera cross), representing the standard modern viticulture configuration in phylloxera-infested regions and enabling rootstock-mediated adaptation to soil conditions, pests, and diseases. Grafted scion trunk develops from successful graft union formation= dormant scion budwood (typically 1-2 buds) grafted onto rooted rootstock cutting during nursery propagation, callus formation and vascular connection establishment at graft interface (2-6 weeks post-grafting), shoot development from scion bud creating above-union trunk, and progressive trunk thickening through secondary growth while maintaining graft union as permanent anatomical feature. Grafted scion trunk anatomical characteristics include= genetic identity matching desired cultivar variety (all tissues above graft union are scion genotype), vascular connection through graft union (xylem and phloem continuity established during graft healing connecting rootstock and scion vascular systems), graft union swelling (typical diameter enlargement 5-25% at union due to differential growth rates or anatomical incompatibility between scion and rootstock), secondary xylem and phloem development above union reflecting scion genotype characteristics (wood anatomy, vessel diameter, ray parenchyma, phenolic content specific to cultivar), and bark characteristics above union matching scion variety. Grafted scion trunk diameter development varies by rootstock influence= vigorous rootstocks (SO4, 1103P, 140Ru) promoting larger scion trunk diameter (year 10 typically 60-100 mm), moderate rootstocks (5BB, 5C, 420A) producing intermediate diameter (year 10 typically 50-80 mm), low-vigor rootstocks (101-14Mgt, Riparia Gloire, 3309C) resulting in smaller trunk (year 10 typically 40-70 mm), with scion diameter typically 80-110% of rootstock diameter at comparable trunk positions. Grafted scion trunk graft union characteristics include= union position typically 5-15 cm above soil surface (industry standard preventing scion rooting while protecting union from soil-borne pathogens), union diameter typically 105-125% of adjacent trunk diameter (swelling from differential cambial activity or anatomical mismatch), union anatomy showing vascular reconnection (xylem vessels and phloem sieve tubes bridging graft interface through callus tissue), potential union incompatibility symptoms (excessive swelling >150% indicating poor compatibility, union necrosis or separation indicating graft failure), and union as potential weakness point (physical breakage risk, pathogen entry site, physiological bottleneck). Grafted scion trunk advantages over own-rooted include= phylloxera resistance (rootstock genetics providing immunity or tolerance to phylloxera root feeding), adaptation to difficult soils (rootstock tolerance to lime-induced chlorosis, salinity, drought, waterlogging, nematodes), vigor control (rootstock influence on scion growth allowing precise canopy management), early maturity in some combinations (certain rootstocks advancing phenology 5-10 days), and improved stress tolerance (enhanced drought resistance, cold hardiness depending on rootstock selection). Grafted scion trunk disadvantages compared to own-rooted include= graft union as structural weakness (breakage risk from wind, equipment contact, freeze damage at union), potential graft incompatibility (some scion-rootstock combinations showing poor growth, reduced longevity, physiological disorders), additional propagation costs (grafting labor, nursery time, technical expertise required), rootstock suckering management (removal of rootstock shoots emerging below union), and potential modification of terroir expression (rootstock effects on scion mineral uptake, water relations, vine balance potentially altering fruit composition). Grafted scion trunk global prevalence includes= dominant in phylloxera-infested regions (Europe post-1880s, California, Eastern USA, New Zealand), utilized for soil adaptation regardless of phylloxera (calcareous soils requiring lime-tolerant rootstocks, saline soils requiring salt-tolerant rootstocks), employed for vigor control in fertile sites (dwarfing rootstocks managing excessive vegetative growth), and representing 70-90% of global commercial vineyard area (with own-rooted vines limited to phylloxera-free regions or resistant cultivars).
Grafted_Scion_Trunk emphasizes scion portion above graft union and is distinguished from Own_Rooted_Trunk (no graft union, genetic continuity root to canopy) and Rootstock_Trunk (portion below graft union) by scion genotype identity and graft union dependency. Critical for understanding modern viticulture standard configuration, rootstock selection impacts on scion development, graft union as structural and physiological feature, and phylloxera management in commercial viticulture.
Aufgepfropfter Sortensstamm
Gepfropfter Edelstamm
Grafted trunk
Scion above union
Veredelte Rebe
Grafted Scion Trunk
A grafted vine combining Chardonnay clone 96 scion with SO4 rootstock bears a Grafted_Vine_Role, produced by omega benchgrafting, grown one season in nursery rows, harvested as bare-root dormant vine, and graded by root count (minimum 4), trunk diameter (minimum 5 mm), and graft union integrity before sale to wineries for vineyard planting. A field-budded grafted vine (Merlot on 110 Richter) serves in a Grafted_Vine_Role after two nursery seasons—first season rooting rootstock cuttings, second season budding and growth—the longer production cycle offset by lower callusing infrastructure requirements. A benchgraft-origin Pinot Noir/3309C vine bears a Grafted_Vine_Role meeting French certification standards (catégorie certifiée), with documented traceability from certified mother plant sources.
A nursery plant role that inheres in a grapevine consisting of a scion cultivar united by grafting to a rootstock genotype, representing the standard commercial planting material for vineyard establishment in phylloxera-affected regions worldwide. Grafted vines combine the desirable fruiting characteristics of the scion (variety identity, fruit quality, yield potential) with the pest resistance, stress tolerance, and vigor modification provided by the rootstock, creating a composite plant optimized for specific site conditions. Grafted vine quality assessment encompasses three components—root system (development, distribution, health), shoot system (bud viability, trunk caliper, absence of damage), and graft union (callus completeness, mechanical strength, vascular continuity)—with all three contributing to establishment success and long-term vine performance. Grafted vines are produced through benchgrafting (dormant scion and rootstock joined in nursery, callused, and field-planted) or field budding (scion bud inserted into established rooted rootstock), with benchgrafting dominant in modern commercial production due to controlled conditions, higher success rates, and single-season production cycles.
Grafted_Vine_Role represents the dominant commercial product of the global grapevine nursery industry, with annual production of hundreds of millions of grafted plants supplying vineyard establishment worldwide. The grafted vine format became universal following the phylloxera epidemic (1860s-1900s), when European V. vinifera vines proved susceptible to the root-feeding insect while American species and their hybrids demonstrated resistance—leading to the grafting solution that enabled continued V. vinifera cultivation on resistant rootstocks. Grafted vine quality directly impacts vineyard economics through establishment success rates, early productivity, vine uniformity, and long-term health quality failures (poor graft unions, infected material, rootstock misidentification) impose multi-decade costs on growers. The ongoing young vine decline problem has focused attention on nursery practices, particularly trunk disease pathogen management through the propagation chain.
Edelrebe
budded vine
compound vine
gepfropfte Rebe
grafted plant
grafted vine
veredelte Rebe
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 415-450 Gramaje D, Armengol J (2011) Fungal trunk pathogens, Phytopathologia Mediterranea 50: S217-S226 Waite H et al. (2015) Hot water treatment, Phytopathologia Mediterranea 54: 325-339 European Union (2020) Commission Implementing Directive 2020/177
Grafted Vine Role
A 6 cm two-node segment of Cabernet Sauvignon clone 337 prepared with an omega cut at its base bears a Graftwood_Role, matched to a disbudded SO4 rootstock cutting of equivalent diameter (10 mm ± 1 mm) for mechanical omega grafting in a French nursery. Graftwood of Sangiovese clone VCR5 serves in a Graftwood_Role for whip-and-tongue grafting onto 420A rootstock, with the diagonal scion cut and interlocking tongue providing maximum cambial contact for union formation. A scion segment of Riesling clone 239 bears a Graftwood_Role in German benchgraft production, prepared on automated grafting machines achieving 3,000-5,000 grafts per hour with consistent cut geometry ensuring reliable union formation.
A scion wood role that inheres in stem material designated for preparation into grafting segments (typically one to three nodes with associated internode tissue) used in grafting techniques where scion and rootstock cane pieces are joined to create unified grafted plants. Graftwood provides multi-node scion segments (typically 5-10 cm length with one to two intact buds) that are joined to rootstock cuttings through various grafting techniques (omega, whip-and-tongue, cleft, machine grafting), with the larger tissue mass compared to budwood providing greater carbohydrate reserves supporting callus formation and union development. Graftwood quality parameters include cane diameter (matching rootstock diameter within 1-2 mm for optimal cambial alignment), internode length (accommodating the grafting cut while preserving at least one viable bud), bud health (both buds viable in two-bud scions for redundancy), tissue maturity (adequate lignification for graft union strength), and freedom from trunk disease pathogens that could colonize the graft interface. Graftwood preparation involves cutting canes to standard scion lengths, making the appropriate grafting cut (omega, whip, or wedge depending on technique), and matching with compatible rootstock material.
Graftwood_Role captures scion material destined for segment-based grafting techniques rather than single-bud budding, with the multi-node format providing greater reserves and redundancy but lower multiplication rate than budwood approaches. The graftwood diameter-matching requirement is critical for grafting success—cambial alignment between scion and rootstock enables vascular connection and nutrient/water transport across the union. Machine grafting systems (omega, whip-and-tongue) have standardized scion dimensions, typically two buds in 5-8 cm length with diameter matching the rootstock component within tight tolerances. Graftwood storage requirements parallel hardwood cutting storage (0-4°C, 95%+ RH) with attention to preventing both desiccation (compromising bud viability) and excess moisture (promoting mold development). The transition from hand grafting to machine grafting has increased graftwood dimensional specifications while improving production efficiency and union consistency.
Pfropfreiser
Propfholz
Veredelungsreiser
grafting segment
grafting wood
graftwood
scion piece
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 450-468 Gramaje D, Armengol J (2011) Fungal trunk pathogens, Phytopathologia Mediterranea 50: S217-S226 Waite H et al. (2015) Hot water treatment of grapevine propagation material, Phytopathologia Mediterranea 54: 325-339
Graftwood Role
Graininess quality assessment comparing filtered versus unfiltered bottlings of same Côtes du Rhône reveals the stylistic choice affecting texture, with filtered version showing smooth, polished mouthfeel (graininess 2/10) appropriate for mainstream commercial expectations, while unfiltered version displays perceptible fine grain (6/10) from residual fine lees that natural wine enthusiasts value as authenticity marker and complexity contributor, demonstrating how identical graininess level may indicate defect or quality depending on wine category and consumer expectation.
Graininess quality is a texture perception quality that represents the presence of fine particulate or gritty tactile sensation during wine consumption, characterizing the perception of discrete textural elements rather than continuous smooth surface that may indicate colloidal particles, certain tannin structures, or residual solids affecting mouthfeel refinement. Within the BFO framework, this quality specifically addresses the particulate dimension of texture perception, distinguishing grainy character from smooth or silky texture while recognizing that graininess may arise from diverse compositional sources with different quality implications depending on wine style and production method. The ontological significance lies in graininess serving as both a potential quality indicator and a stylistic marker, where unwanted graininess suggests quality deficiency from coarse tannins or inadequate finishing, while deliberate graininess in certain styles (sur lie wines with fine lees, unfiltered wines) may represent authenticity and complexity valued by specific consumer segments. The perceptual basis for graininess involves mechanoreceptor detection of discrete particles or textural discontinuities as wine contacts oral surfaces, with the sensation ranging from barely perceptible fine grain through obvious particulate texture to frankly gritty character depending on particle size, concentration, and hardness. The compositional sources of graininess include colloidal tannin aggregates (creating fine grainy texture in tannic red wines), residual fine lees (contributing particulate character in sur lie styles), micro-crystalline tartrate (producing gritty sensation if unstabilized), and protein-tannin complexes (forming grainy precipitates in inadequately fined wines). The quality interpretation of graininess requires context, with grainy texture representing defect in wines expected to be polished (premium filtered wines) while acceptable or valued in styles embracing textural complexity from minimal intervention (natural wines, extended sur lie wines). The developmental aspects of graininess show some sources diminishing with age (tannin aggregates precipitating, settling removing particles) while others persist, affecting aging recommendations for grainy wines. Understanding graininess quality enables texture defect detection in polished styles, authenticity recognition in minimal-intervention wines, and quality assessment incorporating context-appropriate graininess evaluation.
Graininess_Quality addresses the particulate dimension of texture perception, representing textural discontinuity from discrete elements. This quality is context-dependent, indicating defect in polished styles but potentially valued in minimal-intervention wines. The class supports texture assessment and quality evaluation that incorporates style-appropriate graininess expectations.
Körnigkeit
Partikeltextur
fine grain
granular sensation
grießige Textur
gritty mouthfeel
körniges Mundgefühl
particulate texture
Gawel R et al. (2000) A 'Mouth-feel Wheel': terminology for communicating the mouth-feel characteristics of red wine. Australian Journal of Grape and Wine Research 6(3): 203-207 Vidal S et al. (2003) The mouth-feel properties of grape and apple proanthocyanidins in a wine-like medium. Journal of the Science of Food and Agriculture 83(6): 564-573
Graininess Quality
A Riesling vine in the Scharzhofberger vineyard in the Saar bearing a Grand Cru equivalent role under the VDP.Große Lage classification is subject to maximum yields of 50 hl/ha (versus 75 hl/ha for Gutswein), mandatory hand harvesting, minimum must weights of 80-85° Oechsle for dry wines, and origin labeling restricted to the individual vineyard name, with resulting wines commanding 50-200 €/bottle compared to 10-20 €/bottle for regional Mosel Riesling from the same producer.
An appellation-specific role that inheres in a grapevine situated within a vineyard parcel classified at the highest tier of a hierarchical appellation system, representing the most prestigious geographic designation attainable and imposing the most stringent viticultural requirements including lowest maximum yields, highest minimum ripeness levels, and most restrictive production practices.
Grand_Cru_Role represents the apex of vineyard classification systems, most formally codified in Burgundy (33 Grand Cru vineyards for red and white) and Alsace (51 Grand Cru vineyards), with analogous top-tier classifications in Germany (VDP.Große Lage), Austria (ÖTW Erste Lagen), and the Bordeaux 1855 Classification. The ontological status of this role merits attention: it inheres in the vine by virtue of its geographic position within historically and geologically defined parcels, not by any intrinsic biological property. In Burgundy, Grand Cru vineyards carry their own appellation status independent of the commune, with the vineyard name alone appearing on the label.
Grand-Cru-Rolle
Große-Lage-Rolle
Spitzenlagen-Rolle
grand cru vine role
highest classification role
top-tier cru role
INAO (2023) Cahiers des charges Grands Crus de Bourgogne VDP (2022) VDP.Klassifikation Statuten Pitiot S and Servant JC (2010) The Wines of Burgundy, Collection Pierre Poupon pp: 44-68 Johnson H and Robinson J (2019) The World Atlas of Wine 8th Ed, Mitchell Beazley pp: 54-57
Grand Cru Role
Field evaluation of Grape_Berry_Moth_Resistance_Disposition at INRAE Bordeaux monitored Lobesia botrana infestation and damage across 25 cultivars in replicated plots without insecticide application over four seasons. Oviposition density (eggs per 50 clusters), larval penetration success (percent of eggs producing feeding larvae), and berry damage (percent of berries with entry holes or frass) were assessed at first and second generation peaks. 'Merlot' showed high susceptibility with 28 eggs per 50 clusters, 72% penetration success, and 18% berry damage at second generation. 'Cabernet Sauvignon' demonstrated enhanced Grape_Berry_Moth_Resistance_Disposition with equivalent oviposition (25 eggs per 50 clusters) but significantly lower penetration success (41%) and berry damage (8%). Laboratory bioassays revealed that neonate larvae offered 'Cabernet Sauvignon' berries showed 35% mortality during the critical first 24 hours of establishment versus 12% on 'Merlot', with surviving larvae exhibiting 25% longer development time. Berry skin puncture force was significantly higher for 'Cabernet Sauvignon' (0.85 N) than 'Merlot' (0.52 N), and correlation analysis across all cultivars showed significant relationship between skin mechanical resistance and larval establishment failure (r = 0.72). These findings indicate that Grape_Berry_Moth_Resistance_Disposition in 'Cabernet Sauvignon' operates primarily through berry physical properties impeding larval penetration, a trait potentially amenable to selection in breeding programs.
A Grape_Berry_Moth_Resistance_Disposition is an Insect_Resistance_Disposition that inheres in a Vitis plant cluster and is realized through defense processes that limit oviposition, larval establishment, feeding damage, or successful development by grape berry moths (Lepidoptera: Tortricidae), economically important pests whose larvae feed within developing flowers and ripening berries, causing direct crop loss and facilitating secondary infections. This disposition represents resistance to a complex of tortricid moths affecting grapevine globally, including the European grapevine moth (Lobesia botrana) dominant in European, Middle Eastern, and recently South American viticulture, and the grape berry moth (Paralobesia viteana) important in eastern North American viticulture. Female moths oviposit on flower clusters or developing berries, with emerging larvae burrowing into reproductive tissues, destroying florets before bloom, hollowing developing berries, and creating wounds that serve as entry points for bunch rot pathogens including Botrytis cinerea, Aspergillus, and sour rot complexes. Resistance mechanisms may include cluster and berry physical properties (tough skin, compact cuticle) limiting larval penetration, chemical factors deterring oviposition or reducing larval survival, volatile profiles affecting host location by ovipositing females, and cluster architecture (loose clusters with exposed berries) reducing favorable oviposition sites and larval microhabitats. The intimate relationship between berry moth damage and secondary fungal infection means that Grape_Berry_Moth_Resistance_Disposition contributes to bunch rot management beyond direct pest control. Variation in berry moth susceptibility among cultivars has been documented, providing opportunities for breeding and cultivar selection in regions with chronic berry moth pressure.
Grape_Berry_Moth_Resistance_Disposition is_a Insect_Resistance_Disposition. Grape berry moths comprise multiple tortricid species in different regions (L. botrana in Europe, P. viteana in eastern North America), with similar biology and damage. Berry moth damage creates entry points for bunch rot fungi, linking this class to Botrytis_Resistance_Disposition through pest-disease interactions. Resistance mechanisms involving berry skin properties overlap with Thick_Skin_Disposition for Botrytis resistance, suggesting potential for combined selection. L. botrana is a quarantine pest in some regions including the United States following eradication of a 2009 California introduction.
Beeren-Motten-Resistenz
Lobesia resistance
Lobesia-botrana-Abwehr
Traubenwickler-Resistenz-Disposition
grape berry moth tolerance
tortricid berry pest resistance
Ioriatti C et al. (2011) Lobesia botrana (Lepidoptera: Tortricidae), a new pest of grapevine in the Americas, Journal of Economic Entomology 104:1037-1044 Thiéry D, Moreau J (2005) Relative performance of European grapevine moth (Lobesia botrana) on grapes and other hosts, Oecologia 143:548-557 Roehrich R, Boller E (1991) Tortricids in vineyards, Tortricid Pests: Their Biology, Natural Enemies and Control, Elsevier
Grape Berry Moth Resistance Disposition
Concord bears a Grape_Juice_Scion_Role as the definitive variety for American grape juice consumption, with Welch's, Kedem, and other brands processing over 250,000 tonnes annually into the purple, intensely "grapey" juice that defines the category—its methyl anthranilate content (3-5 mg/L) creating the aroma Americans recognize as "grape." Niagara serves in a Grape_Juice_Scion_Role for white grape juice production, with its labrusca aromatic character, balanced sweetness (17-19°Brix), and pale color providing the white grape juice standard in North American markets. Muscadine (Vitis rotundifolia) bears a Grape_Juice_Scion_Role in southeastern United States production, with its distinctive bronze or purple color and unique muscadine flavor supporting regional juice products with claimed health benefits from high polyphenol content.
A juice production scion role that inheres in a grafted scion cultivar by virtue of its designation for direct-consumption grape juice production, where juice is processed, pasteurized, and packaged for drinking as single-strength (100%) grape juice or grape juice beverages. Grape juice scions for direct consumption require distinctive, appealing flavor profiles that consumers recognize as "grape juice" character—in North American markets this means the intense labrusca aromatics of Concord (methyl anthranilate concentrations of 2-5 mg/L creating the characteristic "grapey" flavor), while in other markets V. vinifera-based juices with milder, fruitier profiles may be preferred. Direct-consumption juice production demands consistent quality across seasons, appropriate sugar-acid balance for palatability (typically 16-18°Brix, 0.6-0.8% titratable acidity), attractive color (deep purple for Concord-type, pale gold for white varieties), and absence of off-flavors or excessive astringency. Production involves crushing, enzymatic treatment for yield and clarity, pasteurization (typically 82-88°C), and packaging in glass, PET bottles, or aseptic cartons for retail distribution.
Grape_Juice_Scion_Role captures direct-consumption juice production where varietal flavor character is essential to consumer acceptance—contrasting with Concentrate_Production_Scion_Role where neutral flavor enables blending versatility. The North American grape juice market's dependence on labrusca varieties (Concord, Niagara) reflects historical development and consumer preference formation, creating a market where V. vinifera grapes taste "wrong" as juice despite their dominance in wine. Kosher certification is important in US grape juice production (Kedem, Manischewitz brands), requiring specific production protocols. Health marketing emphasizing polyphenol content and cardiovascular benefits has supported grape juice consumption, though sugar content concerns have moderated growth.
100% juice grape
100%-Saft-Traube
Direktsaft-Traubensorte
Trinksafttraube
drinking juice grape
single-strength juice variety
table juice grape
Morris JR, Striegler RK (2005) Grape juice, in Processing Fruits 2nd Ed, CRC Press pp.: 585-616 Iyer MM et al. (2010) Methyl anthranilate in grape juice, American Journal of Enology and Viticulture 61: 1-7 Fuleki T, Silva JMR (1997) Catechin and procyanidin composition of grape juice, Journal of Agricultural and Food Chemistry 45: 1156-1160 Welch's Company (2020) Annual Production Report
Grape Juice Scion Role
Grape price data for Cabernet Sauvignon in Napa Valley (2023 vintage, Napa Valley Vintners Grape Price Survey) documents the extreme premium commanded by the region's flagship cultivar — average price approximately USD 8,500/ton for Napa Valley AVA fruit, with sub-appellation premiums reaching USD 15,000-30,000+/ton for Oakville, Rutherford, and Stags Leap District sources, and individual vineyard contract prices for the most celebrated sites (To Kalon, Beckstoffer Dr. Crane, Beckstoffer Missouri Hopper) reportedly exceeding USD 35,000-50,000/ton — compared to California-wide Cabernet Sauvignon averages of approximately USD 1,200/ton, demonstrating the 7-40× price multiplier that geographic prestige commands at the grape level. Grape price data for Grüner Veltliner in Austria (2023 vintage, Österreichischer Weinbauverband survey) ranges from approximately €0.40-0.60/kg for Weinviertel Qualitätswein-grade fruit to €1.00-1.50/kg for Kamptal and Kremstal DAC-grade fruit to €2.50-5.00+/kg for top-classified Wachau sites, with cooperative settlement prices typically at the lower end of each range and estate-grown transfer prices imputed at the higher end.
Grape price data is a market data entity that documents the prices paid for harvested grapes at the vineyard gate, winery intake, or on the bulk grape market, expressed per unit of weight (€/kg, USD/ton, AUD/tonne) or per unit of area (€/ha for contracted vineyard production), differentiated by cultivar, quality tier, geographic origin, and vintage year, derived from regional grape price surveys, grower cooperative settlement statements, contract price disclosures, and industry average calculations, and specifying the pricing mechanism (spot market, long-term contract, cooperative pooling, estate-grown transfer price), quality parameters influencing price (sugar content, acidity, health status, yield level), and the geographic and temporal scope of the reported prices, providing the primary indicator of a cultivar's agricultural commodity value at the production level.
Grape price data captures the agricultural commodity valuation that forms the foundation of the wine value chain — the price at which the viticultural product transitions from grower to winemaker, establishing the baseline cost of wine production and the primary revenue stream for grape-growing operations. The pricing mechanisms for grapes vary substantially across wine regions and industry structures — spot market pricing (grapes sold on the open market at harvest, prices determined by current supply-demand conditions, prevalent in Australia's warm-climate regions and California's Central Valley), long-term contracts (multi-year agreements specifying price formulas, quality criteria, and volume commitments, common in premium regions where winery-grower relationships are sustained over decades), cooperative pooling (member growers deliver grapes to a cooperative winery that processes, markets, and distributes revenue based on quality-weighted volume contributions, dominant in many European regions), and estate-grown production (vertically integrated operations where grapes are grown and vinified by the same entity, with the "price" being an internal transfer value estimated for accounting purposes). The quality-price relationship in grape markets is non-linear and regionally specific — premium prices are typically associated with lower yields (the inverse yield-quality relationship), specific geographic origins (site prestige), optimal ripeness parameters (sugar-acid balance), excellent sanitary condition (absence of rot, disease), and cultivar reputation in the specific region. The transparency of grape pricing varies enormously — some regions publish detailed annual grape price surveys (the California Grape Crush Report, the Australian Wine Research Institute average grape price data), while others treat grape pricing as commercially sensitive information disclosed only between contracting parties. The seasonal timing of grape price formation also varies — some markets establish indicative prices before harvest based on crop forecasts, while others determine prices at or after delivery based on actual quality assessment, creating different risk-sharing dynamics between growers and wineries.
Lesepreis
Traubenkaufpreis
Traubenpreis
Traubenpreisdaten
Traubenvergütung
fruit price
grape commodity price
grape pricing
grape purchase price
vineyard gate price
CDFA (2023) California Grape Crush Report, California Department of Food and Agriculture Wine Australia (2023) National Vintage Report — average grape prices Napa Valley Vintners (2023) Napa Valley Grape Price Report Österreichischer Weinbauverband (2023) Traubenpreiserhebung
Grape Price Data
Grape stem necrosis quality assessment in Franken Müller-Thurgau documented the relationship between magnesium status, K:Mg ratio, and Stiellähme incidence to inform corrective nutritional management. Pre-harvest cluster evaluation scored grape stem necrosis quality as the percentage of clusters showing rachis lesions, with severity categories ranging from trace (less than 5% of rachis affected, no berry loss) through moderate (20-40% rachis affected, some berry drop) to severe (greater than 50% rachis affected, significant cluster desiccation). Vineyard-wide grape stem necrosis quality incidence reached 28% of clusters affected, with severity distribution of 15% trace, 9% moderate, and 4% severe, representing estimated crop loss of 8% through direct berry drop plus additional quality compromise from uneven ripening. Nutritional analysis revealed grape stem necrosis quality correlation with tissue nutrient ratios, with blocks showing greater than 20% incidence having mean petiole K:Mg ratio of 14.2 versus 6.8 in blocks with less than 5% incidence, establishing the K:Mg imbalance as the primary predisposing factor. Corrective management integrated immediate foliar MgSO₄ application at 3% concentration (two applications 10 days apart starting at véraison), soil magnesium supplementation with kieserite at 100 kg/ha, and reduced potassium fertilization for two seasons to rebalance the K:Mg ratio. Follow-up assessment showed grape stem necrosis quality incidence reduced from 28% to 8% by the second season post-correction, with the persistent low incidence attributed to complete rebalancing of the K:Mg ratio below 8:1.
Grape stem necrosis quality is a magnesium deficiency symptom quality that inheres in grapevine cluster rachis tissue and represents the physiological disorder (Stiellähme in German) characterized by necrotic lesions developing on the rachis (cluster stem) and pedicels during the ripening period, with affected tissue showing brown-to-black discoloration, tissue collapse, and eventual desiccation that interrupts vascular connection to berries causing berry shrivel, premature drop, or cluster portion loss, with grape stem necrosis strongly associated with magnesium deficiency and elevated K:Mg ratio though other factors including calcium status and environmental conditions contribute to susceptibility. This quality captures a physiological disorder with significant crop loss potential, recognizing that grape stem necrosis can affect up to 30-50% of clusters in severe cases, causing direct yield loss through berry drop and indirect quality loss through uneven ripening of partially affected clusters. The ontological significance of grape stem necrosis quality lies in its economic impact through crop loss and its diagnostic association with magnesium nutrition status, providing an indicator of nutritional imbalance requiring management attention. The biological basis of grape stem necrosis encompasses the role of magnesium in maintaining rachis tissue integrity through its functions in cell membrane stability and enzyme activation, the susceptibility of rachis tissue during the rapid expansion and metabolic activity of the ripening period, and the interaction between magnesium status and other predisposing factors including low calcium, high potassium, water stress, and high crop load. The timing of grape stem necrosis development typically shows symptoms appearing between véraison and harvest when rachis metabolic demand peaks, with symptom severity increasing under conditions that intensify nutrient competition including high crop load, vigorous vegetative growth, and environmental stress. The distribution of grape stem necrosis within clusters often shows concentration at the rachis branch points where vascular restriction may limit nutrient supply, and at the cluster shoulder where weight stress combines with nutritional factors.
Grape_Stem_Necrosis_Quality is_a Magnesium_Deficiency_Symptom_Quality, representing the physiological disorder (Stiellähme) of rachis necrosis associated with Mg deficiency and K-Mg imbalance. This class addresses a significant crop-loss disorder strongly linked to Mg nutrition. The K-Mg ratio (threshold ~8 to 1) is more predictive than absolute Mg alone. Véraison-to-harvest timing reflects peak rachis metabolic demand. Distribution at rachis branch points and cluster shoulders reflects vascular restriction. Correction requires both Mg supplementation and K reduction to rebalance. Note, different from Ca-associated Bunch_Stem_Necrosis (BSN).
Rappen-Nekrose
Stiellähme
Stiellähmung
Traubenstielkrankheit
cluster stem necrosis
pedicel necrosis
rachis necrosis
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Currle O et al. (1983) Biologie der Rebe. Meininger Verlag, Neustadt Cline J, Bradt O (2007) Grape stem necrosis. Ontario Ministry of Agriculture, Food and Rural Affairs Factsheet
Grape Stem Necrosis Quality
A Cabernet Sauvignon vine planted in the Médoc possesses multiple Grapevine_Dispositions that may or may not be realized depending on conditions encountered during each growing season. Its Downy_Mildew_Resistance_Disposition remains unrealized in dry years when Plasmopara viticola inoculum is absent, but becomes realized through defense responses when humid conditions promote infection pressure. Similarly, its Cold_Hardiness_Disposition exists as a potentiality throughout the year but is only realized when acclimation processes are triggered by shortening photoperiod and declining temperatures in autumn, ultimately determining whether primary buds survive winter minima of -15°C. The vine's Quality_Potential_Disposition for producing wines with characteristic blackcurrant aromatics and firm tannins requires appropriate terroir conditions, vine age, crop load management, and harvest timing to be fully realized in the resulting wine.
A Grapevine_Disposition is a disposition (BFO_0000016 (https://www.ebi.ac.uk/ols4/ontologies/bfo/classes/http%253A%252F%252Fpurl.obolibrary.org%252Fobo%252FBFO_0000016)) that is an internally-grounded, realized entity inhering in a Vitis vinifera plant or plant part, representing a potentiality that may be realized under specific triggering conditions through characteristic processes. As a realizable entity, a grapevine disposition exists as a genuine property of its bearer even when not being manifested, distinguishing it from qualities which are always fully present. Grapevine dispositions encompass the full spectrum of potentialities relevant to viticulture and oenology, including resistance and tolerance dispositions that may be realized upon encounter with biotic or abiotic stressors, developmental dispositions governing growth patterns and phenological timing, reproductive dispositions affecting flowering and fruit set, physiological dispositions relating to water relations and nutrient acquisition, and quality-determining dispositions that influence grape and wine characteristics. The realization of a grapevine disposition depends on the presence of appropriate triggers (such as pathogen contact for resistance dispositions or environmental stress for tolerance dispositions) and may be modulated by genetic background, developmental stage, environmental conditions, and viticultural management practices. Understanding grapevine dispositions is fundamental to cultivar selection, rootstock-scion matching, and vineyard site assessment.
This is the root class for all dispositional properties specific to Vitis vinifera in VVO. Dispositions are distinguished from Grapevine_Quality in that qualities are fully present whenever they exist, while dispositions exist as potentialities that require triggering conditions to be realized through processes. Dispositions are also distinguished from Grapevine_Role in that dispositions are internally grounded in the physical structure of the bearer, while roles are externally grounded in social or relational contexts.
Rebendisposition
Rebenpotential
Vitis disposition
Weinrebendisposition
grapevine potentiality
vine disposition
Röhl J, Jansen L (2011) Why functions are not special dispositions, Monist 94(3):379-402 Keller M (2020) The Science of Grapevines, 3rd Edition, Academic Press
Grapevine Disposition
The VIVC database entry for Riesling Weiss (variety number 10077) constitutes a grapevine information entity comprising the prime name, 147 documented synonyms across 38 languages, the SSR reference profile at 9 standard loci, 68 OIV morphological descriptor scores, passport data linking the cultivar to 85 germplasm collection accessions worldwide, 312 bibliographic references, and parentage data identifying Gouais Blanc x Savagnin as the most probable parent combination.
A generically dependent continuant that is about a grapevine or grapevine-related entity, encompassing all forms of documented knowledge, recorded data, classificatory designations, regulatory identifiers, and scientific measurements that represent, describe, or reference Vitis vinifera and its cultivation, processing, and evaluation.
Grapevine_Information_Entity is aligned with BFO_0000031 (https://www.ebi.ac.uk/ols4/ontologies/bfo/classes/http%253A%252F%252Fpurl.obolibrary.org%252Fobo%252FBFO_0000031) and follows IAO (Information Artifact Ontology) conventions for classifying information content entities. The class captures the complete information landscape surrounding Vitis vinifera — from cultivar names and regulatory codes through scientific measurements and ampelographic descriptions to viticulture management records and wine quality assessments. Subclasses are organized by information domain: cultivar designation, identification systems, regulatory documentation, ampelographic description, genetic and genomic data, phenological records, viticulture management information, oenological data, and environmental characterization.
Reben-Informationseinheit
Weinbau-Informationsentität
Weinbau-Wissensobjekt
grapevine data entity
grapevine information content entity
viticulture information entity
IAO (2020) Information Artifact Ontology Smith B et al. (2007) The OBO Foundry: coordinated evolution of ontologies, Nature Biotechnology 25: 1251-1255 OIV (2009) 2nd Edition of the OIV Descriptor List for Grape Varieties and Vitis Species
Grapevine Information Entity
The perennial trunk and cordon system of a spur-pruned Cabernet Sauvignon vine represents a set of Grapevine_Morphological_Structures that persist across growing seasons, while the annual shoot with its nodes, internodes, leaves, lateral shoots, tendrils, and grape clusters exemplifies the seasonally renewed morphological structures that arise each year from dormant buds.
A Grapevine_Morphological_Structure is a material entity (BFO_0000040 (https://www.ebi.ac.uk/ols4/ontologies/bfo/classes/http%253A%252F%252Fpurl.obolibrary.org%252Fobo%252FBFO_0000040)) that is a spatially extended part of Vitis vinifera L., bearing a distinct anatomical identity and developmental origin, encompassing all vegetative and reproductive organ systems, tissues, and cellular structures of the grapevine from the root system through the permanent woody framework to the annual shoot system and its appendages including leaves, tendrils, inflorescences, and berries.
Grapevine_Morphological_Structure serves as the root class for all 772 morphology classes in VVO. It is aligned with BFO_0000040 (material entity) and imports from the Plant Ontology (PO) where applicable. Subclasses are organized by organ system (root, trunk, shoot, leaf, inflorescence, berry) and by tissue and cellular level. Each subclass carries bilingual EN/DE terminology and genus-differentia definitions referencing established ampelographic literature.
Morphologische Struktur der Weinrebe
Rebenmorphologische Struktur
Rebkörperstruktur
Weinrebenorgan
grapevine anatomical structure
grapevine organ system
grapevine plant structure
vine morphological entity
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 1-57 Mullins MG, Bouquet A, Williams LE (1992) Biology of the Grapevine, Cambridge University Press pp.: 17-80 Galet P (2000) General Viticulture, Oenoplurimedia, Chaintre pp.: 51-130 OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species 2nd Ed
Grapevine Morphological Structure
The progression of Riesling through BBCH principal growth stages 0 (sprouting/bud development) to 9 (senescence/dormancy) in the Rheingau over a single vintage year represents a complete annual sequence of Grapevine_Phenological_Processes, where each principal stage comprises subordinate phenological processes such as bud swelling (BBCH 01), wool stage (BBCH 03), green tip (BBCH 05), full bloom (BBCH 65), veraison onset (BBCH 81), and harvest maturity (BBCH 89).
A Grapevine_Phenological_Process is a process (BFO_0000015 (https://www.ebi.ac.uk/ols4/ontologies/bfo/classes/http%253A%252F%252Fpurl.obolibrary.org%252Fobo%252FBFO_0000015)) that unfolds in time through an ordered sequence of developmental stages in Vitis vinifera L., driven by genotype-environment interactions involving thermal accumulation, photoperiod, and endogenous hormonal regulation, and that is systematically classifiable according to the BBCH extended scale for grapevine (Lorenz et al. 1995), spanning the full annual cycle from winter dormancy through bud burst, shoot growth, flowering, fruit set, berry development, veraison, ripening, leaf senescence, and return to dormancy.
Grapevine_Phenological_Process is the root class for all 463 phenology classes in VVO. It aligns with BFO_0000015 (process) and references BBCH staging (Lorenz et al. 1995) as the primary temporal framework. Subclasses model individual phenophases and their sub-stages using n-ary causal relations that capture the interaction of environmental drivers (temperature, photoperiod, water status) with genotype-specific response thresholds. Each subclass is cross-referenced to BBCH codes and, where applicable, to Plant Ontology growth stage terms.
Entwicklungsprozess der Rebe
Phänologischer Prozess der Weinrebe
Phänophase der Weinrebe
Rebenphänologischer Prozess
grapevine developmental process
grapevine growth stage process
grapevine phenophase
vine phenological event
Lorenz DH, Eichhorn KW, Bleiholder H, Klose R, Meier U, Weber E (1995) Growth Stages of the Grapevine: Phenological growth stages of the grapevine (Vitis vinifera L. ssp. vinifera) - Codes and descriptions according to the extended BBCH scale, Australian Journal of Grape and Wine Research 1(2): 100-103 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 59-155 Coombe BG (1995) Growth Stages of the Grapevine, Australian Journal of Grape and Wine Research 1(1): 100-110 Mullins MG, Bouquet A, Williams LE (1992) Biology of the Grapevine, Cambridge University Press pp.: 81-146
Grapevine Phenological Process
Riesling cultivated in the Mosel Valley expresses distinctive grapevine qualities including high titratable acidity (7-9 g/L), moderate sugar accumulation (80-90° Oechsle), pronounced floral and citrus aroma precursor concentrations, and characteristic elongated cluster morphology that collectively define the regional wine style and distinguish it from Riesling grown in warmer climates such as Alsace or Australia.
Grapevine_Quality represents the fundamental ontological category encompassing all specifically dependent continuants that inhere in grapevine entities and determine their characteristics without constituting separable parts. Within the BFO 2020 framework, qualities are dependent entities that exist only by virtue of their bearer and cannot migrate between bearers. For Vitis vinifera, this category captures the complete spectrum of measurable and observable attributes that define the phenotypic expression of grapevine organs, tissues, and the whole plant throughout its lifecycle. Grapevine_Quality serves as the taxonomic root for eight major subcategories: (1) Morphological_Quality addressing physical form and structure, (2) Chemical_Quality encompassing biochemical composition, (3) Physical_Quality covering mechanical and thermodynamic properties, (4) Sensory_Quality capturing human-perceptible attributes, (5) Physiological_Quality representing functional capacities, (6) Phenological_Quality addressing developmental timing characteristics, (7) Viticultural_Quality encompassing agronomic performance metrics, and (8) Stress_Response_Quality capturing plant responses to environmental challenges. Each quality instance is fully present at any time point of its existence and determines how its bearer is at that moment. The systematic organization of grapevine qualities enables precise characterization of cultivar differences, terroir expression, vintage variation, and viticultural management effects. Understanding the quality structure is essential for grape and wine production because qualities ultimately determine fruit composition, wine potential, and vineyard economic value. The VVO quality hierarchy integrates ampelographic descriptors, oenological parameters, and ecophysiological measurements into a coherent ontological framework that supports data interoperability across viticulture research, precision agriculture applications, and wine quality assessment systems.
Grapevine_Quality aligns directly with BFO_0000019 (https://www.ebi.ac.uk/ols4/ontologies/bfo/classes/http%253A%252F%252Fpurl.obolibrary.org%252Fobo%252FBFO_0000019) as its parent class. This category excludes dispositions (tendencies to respond), roles (context-dependent classifications), and functions (selected effects), which are treated as separate BFO categories in the VVO. The boundary between qualities and dispositions can be subtle in plant biology: a quality describes current state (e.g., current sugar content) while a disposition describes potential behavior (e.g., tendency toward high sugar accumulation). All subclasses inherit the BFO quality axioms including specific dependence on a single bearer and full presence at each time of existence.
Qualitätsmerkmal der Weinrebe
Rebqualität
Traubenqualität
Vitis vinifera quality
Weinrebenqualitätseigenschaft
grape quality attribute
grapevine characteristic
vine quality
Grenon P and Smith B (2004) SNAP and SPAN: Towards Dynamic Spatial Ontology. Spatial Cognition and Computation 4(1): 69-104 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 1-15 Arp R, Smith B, and Spear AD (2015) Building Ontologies with Basic Formal Ontology, MIT Press pp.: 71-98
Grapevine Quality
A Pinot Noir vine planted in the Domaine de la Romanée-Conti simultaneously bears a Wine_Production_Scion_Role, a Premium_Red_Wine_Scion_Role, and a Grand_Cru_Role, while the same clone maintained at ENTAV-INRA Montpellier bears a Certified_Mother_Plant_Role and a Reference_Cultivar_Role. A Cabernet Sauvignon vine in the UC Davis Foundation Plant Services collection bears both a Germplasm_Repository_Role and a Virus_Tested_Mother_Plant_Role.
A role is a realizable entity that inheres in a Vitis vinifera plant or plant part by virtue of its position, function, or designation within viticultural production systems, breeding programs, propagation schemes, conservation efforts, or ecological contexts. Grapevine roles are not intrinsic to the organism's biological nature but are externally conferred through human management decisions, institutional frameworks, or environmental circumstances. A single grapevine individual may simultaneously bear multiple roles, such as serving as both a scion mother plant and an experimental cultivar in a breeding trial. Roles are distinguished from dispositions in that they depend on relational context rather than on inherent physical or genetic properties of the bearer.
Grapevine_Role is the top-level class for all roles in VVO, directly aligned with BFO_0000023 (https://www.ebi.ac.uk/ols4/ontologies/bfo/classes/http%253A%252F%252Fpurl.obolibrary.org%252Fobo%252FBFO_0000023). Roles are context-dependent and institutionally conferred, contrasting with BFO_0000016 (disposition) which captures intrinsic biological capacities. The subclass hierarchy organizes roles by functional domain: rootstock function, scion utilization, propagation status, clonal identity, breeding function, vineyard management, production system, regional designation, quality classification, ecological function, research use, and conservation status.
Funktionsrolle der Rebe
Rebenrolle
grapevine functional role
vine role
viticultural role
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press Mullins MG, Bouquet A, Williams LE (1992) Biology of the Grapevine, Cambridge University Press OBI Ontology - role definition pattern (http://purl.obolibrary.org/obo/OBI_0000316)
Grapevine Role
Moscato Bianco bears a Grappa_Base_Scion_Role producing Italy's most celebrated aromatic grappas, where pomace from Moscato d'Asti production retains sufficient terpenes (linalool, geraniol) to create intensely floral spirits prized by connoisseurs—Distilleria Berta, Nonino, and Marolo producing benchmark Grappa di Moscato expressions. Nebbiolo serves in a Grappa_Base_Scion_Role for prestigious Piedmontese grappas (Grappa di Barolo, Grappa di Barbaresco), where pomace from extended maceration wines contains concentrated tannins and aromatic compounds creating structured, age-worthy spirits distinct from lighter grappas. Prosecco (Glera) bears a Grappa_Base_Scion_Role for Veneto grappa production, with pomace from the enormous Prosecco industry supporting substantial grappa volume, though the brief skin contact of Prosecco vinification yields less characterful pomace than extended-maceration red wine production.
A distillation scion role that inheres in a grafted scion cultivar by virtue of its production of grape pomace (vinacce) suitable for grappa distillation, where the skins, seeds, and stems remaining after wine pressing are fermented (if from white wine production) or directly distilled (if from red wine production with prior fermentation) to produce Italy's distinctive pomace brandy. Grappa base scions contribute character through the aromatic and phenolic compounds retained in pomace after primary juice extraction, with variety selection influencing grappa style—aromatic varieties (Moscato, Gewürztraminer, Prosecco/Glera) yield perfumed, floral grappas, while tannic red varieties (Nebbiolo, Amarone pomace from Corvina) produce more structured, complex spirits. Unlike wine distillation where grapes are crushed specifically for spirit production, grappa utilizes winemaking byproducts, linking grappa character inextricably to regional wine production and elevating what was historically a farmhouse utilization of waste into a protected Italian specialty. EU regulations restrict "grappa" designation to Italian production from Italian grapes, with parallel pomace brandies produced elsewhere under different names (marc in France, orujo in Spain, tsipouro in Greece).
Grappa_Base_Scion_Role captures the unique position of grappa production as a byproduct utilization rather than a dedicated grape-to-spirit pathway. Variety influence on grappa derives from compounds remaining in pomace after winemaking—aromatic varieties retain terpenes in skins, while phenolic-rich red varieties contribute tannins and color precursors. The quality revolution in Italian grappa since the 1970s (pioneered by distillers like Nonino) transformed the category from crude firewater into refined spirits, with single-variety grappas (Grappa di Moscato, Grappa di Nebbiolo) and vintage-dated bottlings commanding premium prices. Fresh pomace handling is critical—delays between pressing and distillation cause oxidation and off-flavor development, driving integration between wineries and distilleries or on-site distillation at quality-focused producers. The sustainability narrative of utilizing winemaking waste adds appeal in environmentally conscious markets.
Grappa-Traube
Tresterbrand-Sorte
Vinaccia-Rebsorte
grappa grape
marc variety
pomace brandy grape
vinaccia variety
Ministero delle Politiche Agricole (2020) Disciplinare Grappa Da Porto C (2002) Grappa: a review of production, composition and quality, Journal of Wine Research 13: 179-194 Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Piggott JR et al. (1989) Distilled Beverage Flavour, Ellis Horwood
Grappa Base Scion Role
Grassy aroma comparison between whole-cluster pressed and destemmed Sauvignon Blanc reveals cut grass intensity of 3.5 versus 5.8 (10-point scale) respectively, with cis-3-hexen-1-ol concentrations of 85 versus 180 μg/L, demonstrating how processing decisions (stem contact releasing additional C6 compounds) affect grassy character intensity and enable winemakers to modulate this herbaceous dimension according to style target, with Loire producers often preferring higher grassy intensity while New Zealand style minimizes it in favor of tropical thiols.
Grassy aroma quality is an herbaceous aroma quality that represents olfactory perception evoking freshly cut grass and green leaf-associated scents, characterizing the fresh, sharp, green aromatic impression arising from C6 aldehydes and alcohols that contributes to varietal character in certain white wines and indicates freshness in young wines generally. Within the BFO framework, this quality specifically addresses the grass and green leaf subcategory of herbaceous aroma, distinguished from green pepper by its derivation from different compound classes (C6 compounds rather than methoxypyrazines) and its more universal presence across grape varieties compared to the variety-specific methoxypyrazine character. The ontological significance lies in grassy aroma representing the immediate, fresh-cut vegetative character that indicates wine youth, careful handling, and appropriate harvest timing, where grassy notes suggest vibrant, just-crushed grape character while their absence in young wines may indicate oxidation or processing losses. The compositional basis for grassy aroma includes C6 aldehydes (hexanal contributing green, grassy, fatty notes cis-3-hexenal providing intense cut grass character) and C6 alcohols (cis-3-hexen-1-ol, the dominant leaf alcohol with characteristic freshly cut grass aroma trans-2-hexen-1-ol adding green, leafy notes), formed through enzymatic lipoxygenase activity when grape tissue is disrupted during harvest and processing. The processing sensitivity of grassy character makes it an indicator of handling practices, with rapid processing and inert atmosphere limiting C6 aldehyde oxidation and preserving grassy freshness, while extended skin contact or oxidative handling reduces grassy character through aldehyde-to-alcohol reduction or further oxidation. The fermentation dynamics show most C6 aldehydes reduced to less odor-active alcohols by yeast, with residual grassy character depending on initial concentration and fermentation conditions. Understanding grassy aroma quality enables freshness assessment based on grassy note preservation, harvest and processing evaluation from C6 compound profiles, and wine style characterization where grassy intensity contributes to varietal expression in Sauvignon Blanc and similar varieties.
Grassy_Aroma_Quality addresses the cut grass and green leaf subcategory of herbaceous aroma, derived from C6 aldehydes and alcohols formed during grape tissue disruption. This quality distinguishes from Green_Pepper_Aroma_Quality by its different chemical origin (lipoxygenase products versus methoxypyrazines) and more universal presence across varieties. The processing sensitivity of C6 compounds makes grassy character an indicator of handling practices and wine freshness.
Grasaroma
Wiesenduft
cut grass aroma
fresh grass character
frischer Grasduft
green leaf notes
grüne Blattcharakter
lawn scent
Joslin WS, Ough CS (1978) Cause and fate of certain C6 compounds formed enzymatically in macerated grape leaves. American Journal of Enology and Viticulture 29(1): 11-17 Ferreira V et al. (2000) Chemical characterization of the aroma of Grenache rosé wines. Aroma extract dilution analysis, quantitative determination, and sensory reconstitution studies. Journal of Agricultural and Food Chemistry 48(10): 4563-4571
Grassy Aroma Quality
Pinot Noir grafted on 101-14 Millardet showing gravity sensing columella cells, root reorientation from vertical to horizontal triggering amyloplast sedimentation detectable within 9 minutes, calcium imaging revealing transient calcium increase in columella 18 minutes post-reorientation, auxin reporter showing auxin accumulation on lower root side 75 minutes post-reorientation, differential elongation producing visible downward curvature 3.2 hours post-reorientation, gravity sensing columella cells converting gravity stimulus into growth response through statolith sedimentation, mechanotransduction, calcium and auxin signaling
Gravity sensing columella cells are columella cells emphasizing gravitropic perception function and signal transduction mechanism, characterized by statolith amyloplasts sedimenting in response to gravity creating mechanotransduction signal, endoplasmic reticulum networks interacting with sedimenting amyloplasts potentially detecting mechanical stimulus, cortical actin cytoskeleton connecting amyloplasts to plasma membrane and signaling machinery, calcium signaling cascades triggered by statolith sedimentation transducing mechanical to biochemical signals, and auxin transport machinery responding to gravity signal redistributing hormone asymmetrically, with gravity sensing process showing temporal sequence including rapid amyloplast sedimentation 5-30 minutes, early signaling events calcium transients and protein phosphorylation 10-60 minutes, auxin redistribution 30-120 minutes, and growth response differential elongation visible 1-6 hours, and demonstrating that gravity sensing represents complex multi-step process converting simple physical stimulus of gravity into coordinated biochemical and developmental response. Gravity sensing columella cells statolith mechanism includes= showing amyloplast sedimentation (dense starch-filled plastids settling toward cell bottom in gravity field), exhibiting density-driven movement (amyloplast density 1.4-1.5 g/cm³ versus cytoplasm 1.05-1.1 g/cm³ creating net downward force), demonstrating velocity characteristics (amyloplasts sedimenting 0.1-1 micrometer per minute depending on viscosity and amyloplast size), revealing position sensing (amyloplast position relative to cell reflecting gravity vector direction), and displaying equilibrium establishment (amyloplasts reaching steady-state position representing balance of gravitational and cytoplasmic resistance forces). Gravity sensing columella cells mechanotransduction includes= showing ER interaction (endoplasmic reticulum networks interacting with sedimenting amyloplasts potentially detecting mechanical displacement), exhibiting actin involvement (cortical actin filaments connecting amyloplasts to cell periphery potentially transducing mechanical signal), demonstrating mechanosensitive channels (potential stretch-activated ion channels in membranes responding to amyloplast-induced tension), revealing cytoskeletal tension (amyloplast sedimentation creating tension on cytoskeletal elements potentially triggering signaling), and displaying mechanical-to-biochemical conversion (mechanical stimulus from statolith position converted to biochemical signals initiating response cascade). Gravity sensing columella cells calcium signaling includes= showing calcium transients (gravity stimulus triggering transient increases in cytoplasmic calcium concentration), exhibiting spatial patterns (calcium changes potentially showing asymmetry correlated with gravity direction), demonstrating temporal dynamics (calcium signals occurring 10-30 minutes after reorientation early in response cascade), revealing calcium sources (calcium potentially released from internal stores or entering from apoplast through channels), and displaying downstream effects (calcium changes activating calcium-dependent protein kinases and other signaling components). Gravity sensing columella cells signal transduction cascade includes= showing receptor activation (mechanical signal potentially activating mechanoreceptors or associated proteins), exhibiting protein phosphorylation (signal triggering phosphorylation cascades through kinases and phosphatases), demonstrating second messengers (calcium, pH changes, reactive oxygen species potentially serving as second messengers), revealing gene expression changes (early response genes activated within 30-60 minutes of gravity stimulus), and displaying auxin effects (signal cascade ultimately affecting auxin transport and distribution). Gravity sensing columella cells auxin redistribution includes= showing asymmetric transport (gravity signal causing auxin preferential movement to lower root side), exhibiting PIN protein involvement (PIN auxin efflux carriers potentially relocated or activated asymmetrically in response to gravity), demonstrating lateral auxin movement (auxin moving laterally from upper to lower root side in columella and elongation zone), revealing concentration gradient (auxin accumulating higher concentration on lower side creating asymmetry), and displaying growth effects (auxin asymmetry causing differential cell elongation lower side inhibited creating downward curvature). Gravity sensing columella cells temporal dynamics includes= showing rapid perception (amyloplast sedimentation detectable within 5-10 minutes providing rapid gravity detection), exhibiting intermediate signaling (calcium, kinase cascades, gene expression occurring 10-60 minutes), demonstrating auxin effects (auxin redistribution occurring 30-120 minutes after stimulus), revealing growth response (differential elongation producing visible curvature 1-6 hours after reorientation), and displaying sustained response (continued growth adjustment maintaining downward trajectory). Gravity sensing columella cells molecular components includes= showing starch biosynthesis genes (genes encoding starch synthases and branching enzymes required for statolith formation), exhibiting cytoskeletal proteins (actins, myosins, microtubule components involved in mechanotransduction), demonstrating calcium signaling proteins (calcium channels, calcium-binding proteins, calcium-dependent kinases in signal cascade), revealing auxin transport components (PIN proteins, AUX1 influx carrier, and other auxin transport machinery), and displaying transcription factors (early-responsive transcription factors mediating gene expression changes). Gravity sensing columella cells experimental approaches includes= showing clinostat studies (rotation eliminating net gravity vector abolishing gravitropic response confirming gravity dependence), exhibiting starch mutants (mutants defective in amyloplast starch showing impaired gravitropism documenting statolith requirement), demonstrating calcium imaging (fluorescent calcium indicators revealing gravity-induced calcium changes in columella), revealing auxin reporters (fluorescent auxin sensors showing asymmetric auxin distribution after reorientation), and displaying molecular genetics (mutants in signaling components revealing genes required for gravity response).
Gravity_Sensing_Columella_Cells emphasizes perception mechanism and signal transduction and is distinguished from structural aspects by focus on sensing function. Critical for understanding statolith sedimentation, mechanotransduction, calcium signaling, auxin redistribution, temporal dynamics, molecular components, and the complex multi-step process converting gravity stimulus into gravitropic growth response.
Graviperception cell
Gravitations-wahrnehmende Columella-Zellen
Gravitropism sensor
Gravitropismus-Sensor
Statocyt
Statocyte
Gravity Sensing Columella Cells
A green graft of Chardonnay clone 96 scion cleft-grafted onto actively growing SO4 rootstock shoot bears a Green_Graft_Role, performed in a California greenhouse in July using 10-15 cm green shoots from both partners, wrapped with parafilm, and maintained under 80% shade with intermittent mist—the union callusing within 7-10 days and producing a growing grafted plant within 4-6 weeks. A green graft of Cabernet Franc serves in a Green_Graft_Role for rapid virus-tested stock multiplication at FPS, with meristem-derived mother plant shoots providing scion material year-round for continuous production independent of the dormant season. A green graft of Nebbiolo onto Kober 5BB bears a Green_Graft_Role in Italian nursery acceleration programs, enabling same-year production of rooted grafted vines without the overwintering required for dormant bench grafts.
A propagation material role that inheres in a grafted combination created from actively growing (green/herbaceous) scion and rootstock tissue during the growing season, exploiting the rapid callus formation and vascular connection characteristic of juvenile, metabolically active tissues. Green grafts (also termed herbaceous grafts or summer grafts) utilize non-lignified shoot tissue from both scion and rootstock, joined through cleft, whip, or specialized green grafting techniques under protected conditions (typically greenhouse or high humidity chambers) that maintain tissue turgor during the union formation period. Green grafting offers advantages including rapid union formation (callus visible within days rather than weeks), high success rates with proper technique (85-95%), ability to propagate throughout the growing season independent of dormant material availability, and compatibility with virus-tested mother plant multiplication programs where year-round propagation accelerates clean stock increase. The technique requires controlled environment facilities (mist systems, shade structures, humidity chambers) to prevent desiccation of the delicate herbaceous tissues before vascular connection establishes water transport across the union.
Green_Graft_Role captures the herbaceous grafting approach that complements dormant bench grafting by enabling growing-season propagation with rapid union formation. The technique's advantages (speed, year-round capability, high success with proper conditions) are balanced against requirements for controlled environment facilities and the more delicate handling required for non-lignified tissues. Green grafting has gained importance in clean stock programs where accelerated multiplication of virus-tested material is prioritized—the ability to propagate continuously throughout the growing season substantially increases annual output from limited nuclear stock. The physiological basis for rapid green graft success involves the high meristematic activity and cellular plasticity of juvenile tissues, enabling callus formation and vascular differentiation more rapidly than in lignified dormant tissues. Commercial application of green grafting varies regionally, with extensive use in table grape production (particularly in warm climates) and increasing adoption for accelerated wine grape nursery production.
Grünveredelung
Herbstveredelung
Sommerveredelung
green graft
growing-season graft
herbaceous graft
krautige Veredelung
soft graft
summer graft
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 460-465 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press Golino DA et al. (2017) Foundation Plant Services, HortScience 52: 1190-1194
Green Graft Role
A green-growing grafted vine (Chardonnay/SO4) with 30 cm of active shoot growth and established leaf canopy bears a Green_Growing_Vine_Role, produced by February greenhouse forcing of callused benchgrafts, hardened in April, and marketed in May for immediate vineyard planting with continued growth throughout the first season. A green-growing potted vine of Sauvignon Blanc/101-14 serves in a Green_Growing_Vine_Role for accelerated vineyard establishment, the active growth status enabling significant first-year trunk and root development exceeding that achievable from dormant planting. A green-growing vine planted in June bears a Green_Growing_Vine_Role in late-establishment situations where dormant planting windows were missed, with the active plant tolerating summer planting when provided adequate irrigation.
A nursery plant role that inheres in a grapevine in an actively growing physiological state, with expanding leaves, elongating shoots, and functional root activity, produced and sold during the growing season for immediate vineyard establishment. Green-growing vines (also termed hot plants, growing plants, or accelerated vines) are produced by initiating growth of benchgrafts or rooted cuttings under controlled conditions (greenhouse, heated propagation house) before the natural outdoor growing season, then marketing plants with established leaf area and active root systems for spring through summer planting. The green-growing format offers advantages including immediate continued growth after transplanting (no dormancy-break delay), first-year shoot and root development supporting earlier vineyard productivity, extended planting season (into summer with irrigation), and potential for year-one fruit in favorable conditions. Green-growing vine production requires heated facilities for growth initiation, container systems (plants cannot be bare-root while actively growing), and careful hardening-off to transition from protected greenhouse conditions to outdoor vineyard environments.
Green_Growing_Vine_Role captures the premium nursery product category that has expanded significantly as growers seek accelerated vineyard establishment and extended planting flexibility. The format represents an inversion of traditional dormant-vine commerce—rather than exploiting dormancy for handling tolerance, green-growing production exploits active growth for establishment speed. Production economics differ substantially from bare-root systems heated greenhouse space, container systems, extended growing time, and careful handling increase per-plant costs but enable price premiums reflecting faster vineyard productivity. The green-growing format has particular value for high-value plantings (premium wine grapes, licensed varieties) where earlier production offsets higher plant costs, and for vineyard renovation where active plants compete effectively with adjacent mature vines. Hardening-off (gradual acclimation to outdoor conditions) is essential plants moved directly from greenhouse to vineyard suffer shock from temperature fluctuation, wind, and full sun exposure.
Grünpflanze
accelerated vine
actively growing vine
aktiv wachsende Rebe
beschleunigte Rebe
green plant
green-growing vine
grünwachsende Rebe
hot plant
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 380-390 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press Smart DR et al. (2006) Grapevine rooting patterns, American Journal of Enology and Viticulture 57: 89-104 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press
Green Growing Vine Role
Green hue assessment in Sauvignon Blanc berries at harvest provides quality indicators, with optimal fruit displaying pale green to yellow-green hue (CIELAB h° 100-115°) indicating appropriate maturity without overripening berries retaining intense green hue (h° above 120°) suggest underripeness with excessive methoxypyrazine concentrations that would produce overly herbaceous wines, while berries shifted toward pure yellow (h° below 95°) indicate possible overripeness with degraded acidity and diminished varietal character.
Green_Hue_Quality represents the chromatic character of grapevine tissues displaying coloration in the green portion of the visible spectrum, corresponding to wavelengths approximately 495-570 nm and CIELAB hue angles of approximately 90-150°. Green hue results primarily from chlorophyll pigments that absorb red and blue light while reflecting green wavelengths, making this the dominant hue of photosynthetically active tissues. In grapevine, green hue characterizes leaves throughout the growing season, young shoots, tendrils, immature berries, and the flesh of all grape cultivars regardless of skin color. The specific shade of green varies with chlorophyll concentration, the ratio of chlorophyll a to chlorophyll b, and the presence of modifying pigments particularly carotenoids. Young expanding leaves often display yellow-green hues due to lower chlorophyll content, transitioning to deeper blue-green as chlorophyll accumulates in mature leaves. Green hue in berries prior to véraison reflects their photosynthetic activity and serves as a developmental marker, with the loss of green hue signaling the onset of ripening. Some cultivars retain green skin coloration through maturity, including most white grape varieties where the term white actually describes pale green to yellow-green berries. Environmental factors affecting chlorophyll content, including nitrogen availability, light environment, and temperature, modulate green hue expression. Stress conditions causing chlorophyll degradation shift green hue toward yellow as the underlying carotenoid pigments become visible.
Green_Hue_Quality is positioned as a subclass of Hue_Quality representing the green region of the color spectrum. This hue dominates in chlorophyll-containing tissues and serves as the baseline against which véraison color changes are assessed in ripening berries. The distinction between green-skinned (white) cultivars and colored cultivars represents a fundamental division in grape classification. Green hue in leaves can indicate physiological status, with yellow-green shifts signaling nitrogen deficiency, water stress, or disease. OIV descriptor 006 (color of upper surface of blade) includes green categories for young leaf assessment.
Chlorophyllfärbung
Grünton
chlorophyll coloration
green color
green spectrum quality
grüne Farbqualität
grüne Färbung
verdant hue
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 119-135 Lichtenhaler HK and Buschmann C (2001) Chlorophylls and Carotenoids: Measurement and Characterization by UV-VIS Spectroscopy. Current Protocols in Food Analytical Chemistry 1(1): F4.3.1-F4.3.8
Green Hue Quality
A green one-year cane on Vitis vinifera cv. Gewürztraminer examined in late October in a cool-climate vineyard, retaining green appearance throughout 165cm length with no brown coloration visible, soft flexible texture when bent showing no woody rigidity, cross-sectional examination revealing minimal periderm development with epidermis still primary protective tissue, limited secondary xylem formation, and large central pith occupying majority of stem diameter, representing problematic incomplete maturation resulting from excessive late-season vigor (heavy nitrogen application in August, continued irrigation through September) that prevented normal growth cessation and lignification initiation, requiring complete removal during pruning due to zero cold hardiness and unsuitability as fruiting wood.
A one-year cane that has not yet initiated or completed lignification, characterized by retention of green chlorophyll-containing tissues throughout its length, soft herbaceous texture lacking woody firmness, active or recently ceased primary growth with minimal secondary cell wall development, and developmental status representing either early-season immaturity (spring through mid-summer) when continued growth is expected or problematic late-season immaturity (late autumn) indicating incomplete wood ripening that threatens winter survival. Green one-year canes represent normal developmental status during active growing season when shoots are elongating and maturing, but indicate serious maturation failure if green appearance persists into late autumn approaching dormant season, requiring management intervention to promote lignification completion or acceptance of reduced fruiting wood availability and potential cold damage.
Green one-year canes are normal and expected during spring through early autumn when shoots are actively growing, with the critical distinction being timing of assessment. Green appearance in May-August indicates healthy growing shoots that will progressively lignify during autumn maturation. Green appearance persisting in October-November indicates problematic incomplete ripening requiring corrective action. Causes of late-season green canes include excessive vigor from over-fertilization or irrigation, heavy crop loads depleting carbohydrates needed for lignification, shade limiting photosynthesis, late-season growth flushes from mistimed cultural practices, and insufficient thermal time in short growing seasons. Management responses include immediate cessation of water and nitrogen inputs, crop reduction in following season, canopy management improvements, and consideration of earlier-maturing cultivars if problem is chronic.
grüne Rute
herbaceous cane
immature cane
krautige Rute
non-woody cane
unlignified cane
unverholzte Rute
Based on Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 245-267 Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Howell (2001) "Sustainable grape productivity and the growth-yield relationship= A review" American Journal of Enology and Viticulture 52=165-174.
Green One Year Cane
Green pepper aroma quantification in Cabernet Franc from Loire versus California reveals 28 ng/L IBMP and green pepper intensity of 6.5 (10-point scale) in Chinon versus 6 ng/L IBMP and intensity of 2.8 in Napa Valley, with the 4.7-fold concentration difference creating the stylistic distinction between Loire's herbaceous-accented expression and California's fruit-dominant style, demonstrating how green pepper intensity reflects both climate (sun exposure degrading IBMP) and regional style expectations for this variety.
Green pepper aroma quality is an herbaceous aroma quality that represents olfactory perception specifically evoking green bell pepper (Capsicum) scent, characterizing the distinctive sharp, vegetable aromatic impression that serves as a defining varietal marker for Sauvignon Blanc family varieties and may indicate ripeness level across grape varieties containing methoxypyrazine compounds. Within the BFO framework, this quality specifically addresses the green pepper subcategory of herbaceous aroma, distinguished by its derivation from a single compound class (methoxypyrazines) and its strong variety-specific associations and ripeness implications. The ontological significance lies in green pepper aroma providing one of the most chemically precise aroma-compound relationships in wine, where the perception is almost entirely attributable to 3-isobutyl-2-methoxypyrazine (IBMP) and related pyrazines, creating clear compositional basis for this specific aromatic quality. The compositional basis centers on 3-isobutyl-2-methoxypyrazine (IBMP), with detection threshold of approximately 2 ng/L in red wine and 1 ng/L in white wine, making it one of the most potent aroma compounds in wine, with additional contributions from 3-isopropyl-2-methoxypyrazine (IPMP, asparagus, pea) and 3-sec-butyl-2-methoxypyrazine (SBMP, bell pepper, galbanum). The ripeness dynamics of green pepper character show strong decline with grape maturation as IBMP degrades through photolytic breakdown in sun-exposed fruit, with concentrations potentially declining 10-fold from véraison to full ripeness in well-exposed clusters, creating the clear inverse relationship between green pepper intensity and perceived ripeness. The variety distribution of green pepper capacity is genetically determined, with Sauvignon Blanc, Cabernet Sauvignon, Cabernet Franc, Carmenère, and related varieties showing significant IBMP accumulation while varieties lacking the relevant biosynthetic pathway (Chardonnay, Pinot Noir, Merlot) show minimal green pepper potential regardless of ripeness. Understanding green pepper aroma quality enables varietal identification for methoxypyrazine-producing varieties, ripeness assessment from green pepper intensity, and quality evaluation incorporating style-appropriate pyrazine expression.
Green_Pepper_Aroma_Quality addresses the specifically green pepper (Capsicum) subcategory of herbaceous aroma, with an unusually direct relationship to a single compound class (methoxypyrazines, particularly IBMP). This quality provides clear ripeness indication through the well-characterized photolytic degradation of IBMP during grape maturation. The variety-specific presence of methoxypyrazines makes green pepper character relevant only for genetically capable varieties.
Capsicum-Charakter
Gemüsepaprika-Note
Paprikaaroma
bell pepper notes
capsicum aroma
green capsicum scent
grüner Paprikaduft
pyrazine character
Roujou de Boubée D et al. (2000) Identification of 3-isobutyl-2-methoxypyrazine in wine and estimation of its content. Journal of Agricultural and Food Chemistry 48(10): 4830-4834 Lacey MJ et al. (1991) Methoxypyrazines in Sauvignon blanc grapes and wines. American Journal of Enology and Viticulture 42(2): 103-108
Green Pepper Aroma Quality
Syrah in Barossa Valley shows green shoot tip visible at late BBCH 07 in late April after 800-900 chilling hours and 450-500 growing degree days base 10°C, with 12-15 mm shoot visible including compressed first leaves along axis over 2-3 days under conditions of 18-21°C mean temperature, 45-55% relative humidity, and 13-hour photoperiod, representing typical warm-climate pattern for this mid-season cultivar
Green shoot tip visible represents the advanced emergence phase where elongated shoot extends prominently beyond bud scales with beginning evidence of shoot structure including compressed first leaves, marking transition from simple tip visibility to recognizable shoot configuration. This developmental stage occurs during late BBCH 07 progression toward BBCH 09 when shoot has extended 10-20 mm visible length beyond protective scales, with shoot structure showing not only apical tip but also beginning differentiation of shoot axis and enclosed leaf primordia. The visible shoot displays bright green coloration throughout with characteristic grapevine shoot morphology becoming apparent, including slightly angular cross-section reflecting vascular bundle arrangement and subtle nodes marking positions where leaf primordia attach to developing axis. Shoot apex remains conical or rounded at tip where apical meristem continues active growth enclosed by youngest primordia, with older leaf primordia visible as compressed structures along shoot axis though not yet unfolded or separated from axis. Scale separation has progressed extensively with gaps of 5-8 mm at widest point and scales beginning to fall away from shoot base, transitioning from functional protective role to vestigial remnants that will completely detach as shoot continues elongating. The shoot primordium beneath visible portion has reached 15-25 mm total length including sections still enclosed by loosened scales, with apical meristem producing new cells every 18-20 hours and elongation zone expanding at 2-4 mm per day. First leaf primordia along visible shoot axis measure 5-10 mm length remaining tightly appressed against shoot but showing beginning separation preparation, with leaf surfaces displaying characteristic grapevine leaf texture including developing trichomes and beginning venation patterns. Chlorophyll concentration in shoot and leaf tissues reaches 0.6-0.9 mg per gram fresh weight creating intense green coloration, with chloroplasts achieving 50-60% structural development and beginning substantial photosynthetic activity. Photosynthetic capacity increases to 30-40% of mature rates as functional chloroplasts accumulate and leaf area begins expanding, though respiration still exceeds carbon fixation keeping shoots as net carbon consumers for several more days. Vascular system extends through entire visible shoot length with xylem conducting water at 1.5-2.5 mg per second supporting expansion and increasing transpiration, while phloem continues primary role of transporting stored reserves though beginning to carry limited photosynthate from developing leaves. Cell division continues actively in apical meristem and in young leaf primordia, with mitotic zones producing cells for continued shoot elongation and leaf expansion while older cells behind meristem complete expansion and begin differentiation into specialized tissue types. Metabolic activity maintains elevated levels with respiration 12-16 fold above dormancy baseline supporting rapid growth, drawing on stored carbohydrates though beginning transition toward photosynthetic carbon contribution. Protein synthesis continues at high rates producing components for expanding tissues, with particular emphasis on photosynthetic proteins as chloroplast development accelerates in expanding leaf primordia. Hormonal regulation maintains elevated cytokinin and gibberellin supporting continued growth, with auxin concentration highest at apex and declining basipetally establishing polarity for shoot orientation and coordinating leaf and tendril positioning. Temperature continues as primary rate factor with development doubling per 10°C between 10-25°C, though increasing shoot size begins creating microclimate effects that moderate temperature exposure. Water demand increases substantially as transpiring surface area expands to 20-40 square mm per shoot, requiring adequate soil moisture and functional root systems to prevent water stress slowing development. Light intensity increasingly affects development with photosynthetic capacity enabling response to light availability, with high light promoting rapid shoot and leaf expansion while shade begins measurably reducing growth rates. The process typically requires 2-3 days progression from extended green tip to visible shoot tip under favorable conditions above 16°C mean temperature, with cooler conditions extending duration to 3-5 days. Cultivar variation appears in shoot structure visibility and leaf primordium development at this stage, with some varieties showing earlier leaf structure definition while others maintain compressed configuration longer. Regional timing follows extended green tip by 2-3 days, occurring late April in warm climates, early May in temperate regions, and mid-May in cool areas. Viticultural significance includes marking transition point where shoot structure becomes apparent enabling assessment of shoot density and uniformity, providing developmental reference for beginning canopy management planning, serving as observation point for detecting shoot abnormalities or damage, and establishing marker for transitioning pest monitoring focus from buds to shoots and developing leaves. Frost damage at this stage kills exposed shoot and developing leaves, forcing secondary bud growth with 1-2 week delays and reduced fruitfulness. Research applications include documenting shoot morphological development during emergence, analyzing leaf initiation and early expansion processes, evaluating photosynthetic capacity development in young tissues, and assessing cultivar differences in early shoot architecture affecting subsequent canopy development.
Green_Shoot_Tip_Visible is the third of three Green_Tip_Emergence subphases, distinguished by substantial shoot extension revealing shoot structure including compressed leaves. Transitions toward First_Leaf_Unfolding as leaves begin separating from axis. Represents advanced emergence stage approaching BBCH 09. Marks point where shoot architecture becomes assessable for uniformity evaluation.
Triebaufbau sichtbar
Triebspitze mit Blättern
emerging shoot visible
erkennbarer Trieb
recognizable shoot
shoot tip with leaves
sichtbare Triebstruktur
visible shoot structure
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 180-185 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 82-86 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Green Shoot Tip Visible
Riesling in Mosel shows green tip emergence at BBCH 07 in mid-April after 1200-1350 chilling hours and 320-380 growing degree days base 10°C, with 3-5 mm green shoot tip visible through separated scales over 4-5 days under conditions of 12-15°C mean temperature, 65-75% relative humidity, and 13.5-hour photoperiod, representing typical cool-climate timing for this late-bursting cultivar
Green tip emergence represents the specific phenological event marking first visibility of green shoot tissue through separating bud scales, serving as standardized reference point for dormancy exit observation and regional phenological monitoring. This critical developmental milestone corresponds primarily to BBCH 07, occurring when expanding shoot primordium breaches protective scale barrier and green tissue becomes visible at bud apex. The process encompasses the progression from barely detectable green coloration through scales to clearly visible green shoot tip extending beyond scale margins, typically requiring 3-6 days depending on temperature regime and cultivar characteristics. Green tip emergence serves as universally recognized phenological marker across viticultural regions worldwide, providing standardized observation point for comparing seasonal timing, assessing frost risk, and tracking long-term climate impacts on growing season initiation. The visible green tissue represents the shoot apical meristem and immediately surrounding cells that have accumulated sufficient chlorophyll to create characteristic coloration, transitioning from pale yellow-green of newly emerged tissues to brighter green as chloroplast development proceeds and light exposure increases. Scale separation at bud apex creates viewing opportunity for enclosed tissues, with gaps of 2-4 mm allowing observation of green shoot tip that measures 2-8 mm visible length depending on progression stage within emergence process. The shoot primordium beneath visible tip has reached 5-10 mm total length with actively dividing apical meristem producing new cells every 18-24 hours, while cells in subapical elongation zone expand through turgor pressure generating mechanical force pushing shoot through scale opening. Emergence timing depends on accumulated heat units from dormancy release, with most cultivars requiring 250-400 growing degree days base 10°C from initial bud swelling to green tip visibility. Temperature remains primary controlling factor with emergence rate doubling approximately every 10°C increase between 10-25°C, creating strong temperature dependence in timing and progression rate. The process shows three distinguishable phases based on visibility and extension of green tissue, progressing from first barely visible green coloration through clearly extended green tip to prominent green shoot tip with beginning leaf tissue visibility. Chlorophyll synthesis accelerates in light-exposed tissues creating increasingly intense green coloration, with chlorophyll concentrations reaching 0.3-0.5 mg per gram fresh weight in emerged tissues compared to negligible levels in enclosed portions. Photosynthetic capacity begins developing as chloroplasts assemble light-harvesting complexes and Calvin cycle enzymes, though photosynthetic rate remains below 20% of mature leaf capacity and respiration continues exceeding carbon fixation making shoots net carbon consumers. Vascular connection between emerging shoot and permanent structures achieves full functionality with xylem conducting water at 0.5-2.0 mg per second supporting turgor-driven growth and beginning transpiration from exposed surfaces. Metabolic activity maintains peak levels with respiration 15-20 fold above winter baseline supporting rapid cell division and expansion, consuming stored carbohydrates mobilized from woody tissue reserves through phloem transport. Protein synthesis operates at maximum capacity producing structural proteins for new cells, enzymes supporting metabolism, and regulatory proteins coordinating development. Hormonal regulation shows elevated cytokinin and gibberellin promoting cell production and expansion, with auxin concentration highest at shoot apex establishing apical dominance. Cultivar variation in emergence timing reflects genetic differences in chilling requirements, base temperature thresholds, and developmental rates, with early cultivars like Chardonnay and Pinot Noir showing green tip 7-14 days before late cultivars like Riesling and Mourvèdre under identical conditions. Regional timing varies from late March in warm Mediterranean climates through mid-April in temperate regions to early May in cool continental areas, with accumulated heat units providing better prediction than calendar dates across diverse climatic zones. Viticultural significance includes providing standardized observation point for regional phenology monitoring and historical comparisons, marking critical reference for deploying frost protection as tissue vulnerability reaches maximum, establishing developmental marker for initiating pest and disease management programs, and serving as regulatory milestone for vineyard operation restrictions in some appellations. Green tip stage represents period of maximum frost vulnerability with tender emerging tissues highly susceptible to freeze damage, requiring protection when temperatures approach -2°C compared to -5°C or lower tolerance during dormancy. Frost damage at green tip causes complete primary bud mortality through ice formation in hydrated tissues, necessitating compensatory growth from secondary or basal buds with 2-3 week delays and 40-60% fruitfulness reduction. Research applications include long-term phenological monitoring documenting climate change impacts on growing season timing, thermal time modeling for predicting emergence across regions and climate scenarios, molecular analysis of genes regulating shoot emergence and meristem activity, and standardized observations supporting viticulture decision support systems and climate adaptation strategies.
Green_Tip_Emergence focuses specifically on visibility milestone of green tissue appearance, overlapping temporally with Initial_Bud_Breaking through Active_Bud_Breaking but emphasizing observation-based phenological marker rather than developmental process. Subdivided into First_Green_Tip_Visible, Green_Tip_Extended, and Green_Shoot_Tip_Visible based on visibility and extension. Serves as universally recognized phenological reference point across viticultural regions.
Grünspitze
Spitzenaustrieb
green tip
green tip visible
sichtbare Grünfärbung
tip emergence
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 175-180 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Jones GV & Davis RE (2000) Climate influences on grapevine phenology, grape composition, and wine production and quality for Bordeaux, France, Am J Enol Vitic 51:249-261
Green Tip Emergence
Cabernet Sauvignon in Napa Valley shows green tip extended at mid-BBCH 07 in mid-April after 1100-1250 chilling hours and 380-430 growing degree days base 10°C, with 5-7 mm bright green shoot tip clearly visible through 4-5 mm scale opening over 2-3 days under conditions of 16-19°C mean temperature, 50-60% relative humidity, and 14-hour photoperiod, representing typical warm-climate timing for this late-season cultivar
Green tip extended represents the intermediate visibility phase where green shoot tissue extends noticeably beyond bud scales, characterized by unambiguous green tip observation and progressive shoot elongation creating clear phenological marker. This developmental stage occurs during mid-progression of BBCH 07 when shoot primordium has achieved sufficient growth to extend 4-8 mm visible length beyond protective scales, with bright green coloration clearly distinguishable under all normal viewing conditions. The extended green tip shows characteristic conical or cylindrical shape tapering toward apex where apical meristem remains enclosed by youngest leaf primordia, with shoot tissue displaying smooth surface texture and uniform green coloration indicating active chlorophyll synthesis and chloroplast development. Scale separation has progressed to 3-5 mm at bud apex with scales beginning to separate further down bud structure as internal pressure from expanding tissues continues forcing mechanical separation, though scales maintain structural connection at base and continue providing protection to enclosed developing structures. The shoot primordium beneath visible tip has reached 8-12 mm total length with apical meristem maintaining active cell production every 18-22 hours, while elongation zone cells expand rapidly through turgor pressure of 0.6-0.9 MPa driving observable shoot extension of 1-3 mm per day. Chlorophyll concentration in extended tip tissues reaches 0.4-0.7 mg per gram fresh weight creating bright green coloration characteristic of actively growing vegetative tissues, with chloroplasts achieving 30-40% of structural development toward mature configuration. Photosynthetic capacity increases to 15-25% of mature leaf rates as chloroplasts become functional, though respiration continues exceeding carbon fixation meaning shoots remain net carbon consumers dependent on stored reserves. Vascular tissues extend through elongating shoot with functional xylem conducting water at 1.0-1.8 mg per second supporting expansion and increasing transpiration from exposed surfaces, while phloem continues transporting stored carbohydrates mobilized from woody tissue reserves. Cell division maintains high rates in apical meristem producing cells for continued shoot elongation and initiating additional organ primordia including leaves and tendrils, with mitotic activity creating zone of actively dividing cells 100-200 micrometers behind apex. Metabolic activity continues at elevated levels with respiration 15-18 fold above winter baseline, consuming stored sugars at rates creating measurable starch depletion in supporting cane tissues. Protein synthesis operates at high capacity producing structural proteins for new cell walls and membranes, enzymes supporting rapid metabolism, and regulatory proteins coordinating developmental progression. Hormonal concentrations maintain elevated cytokinin and gibberellin promoting continued growth, with auxin gradients from shoot apex coordinating upward growth orientation and beginning to establish patterns for lateral organ differentiation. Temperature remains primary rate-determining factor with development doubling per 10°C between 10-25°C, creating strong thermal dependence in extension rate and subsequent developmental timing. Water demand increases substantially as exposed tissue area expands and transpiration from green surfaces accelerates, requiring adequate soil moisture and functional root uptake to maintain turgor supporting extension. Light intensity begins affecting development as photosynthetic capacity increases, with higher light promoting chlorophyll synthesis and shoot extension though development continues in reduced light. The process typically requires 2-3 days progression from first visibility to extended tip under favorable spring conditions with consistent temperatures above 15°C, with cooler conditions extending duration to 3-5 days. Cultivar variation appears in extension rate and final tip length at this stage, with vigorous varieties showing 2-3 mm per day extension achieving 7-9 mm total visible length while less vigorous types extend 1-2 mm per day reaching 4-6 mm length. Regional timing follows first green tip visible by 1-3 days, occurring mid-April in warm climates, late April in temperate areas, and early May in cool regions. Viticultural significance includes providing clear unambiguous observation point for phenological monitoring with minimal observer variability, serving as reliable reference for frost risk assessment and protection deployment decisions, marking developmental stage for beginning early-season pest monitoring programs, and establishing standardized phenological marker for research studies and climate monitoring. Frost damage at extended green tip stage causes primary bud mortality through ice formation in exposed tissues, requiring secondary bud compensation with 2-3 week delays and 40-60% fruitfulness reduction. Research applications include standardized phenological observations for climate studies, validation of thermal time models for development prediction, analysis of cultivar variation in emergence patterns, and documentation of environmental effects on shoot extension rates during early development.
Green_Tip_Extended is the second of three Green_Tip_Emergence subphases, distinguished by clear unambiguous visibility and measurable shoot extension beyond scales. Transitions to Green_Shoot_Tip_Visible when shoot elongation reveals additional structures beyond simple tip. Represents most reliable observation stage with minimal inter-observer variability. Standard reference point for many phenological studies.
ausgeprägte Grünfärbung
clear green tip
deutliche Grünspitze
elongated shoot tip
extended green tip
gestreckte Triebspitze
prominent green tip
verlängerte Grünspitze
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 177-180 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Buttrose MS (1974) Climatic factors and fruitfulness in grapevines, Hort Abstr 44:319-325
Green Tip Extended
Gross carbon gain assessment comparing VSP-trained versus minimally pruned Chardonnay reveals the canopy architecture effect on total carbon acquisition, with VSP (leaf area 3.2 m²/vine) achieving gross carbon gain of 1850 g C/vine/season through efficient light distribution to high-capacity leaves, while minimally pruned (leaf area 5.8 m²/vine) shows gross gain of 2150 g C/vine/season with the 16% increase substantially less than the 81% increase in leaf area due to mutual shading limiting interior leaf contribution, demonstrating diminishing returns from excessive leaf area and the importance of canopy efficiency rather than simply canopy size.
Gross carbon gain quality is a carbon balance quality that represents the total rate of carbon acquisition through photosynthesis before accounting for any respiratory losses, characterizing the maximum carbon input to the vine system that sets the upper limit on carbon available for growth, fruit development, and reserve storage. Within the BFO framework, this quality specifically addresses the carbon acquisition dimension of carbon balance, distinguishing total photosynthetic fixation from net retention while recognizing that gross carbon gain represents the fundamental productivity potential that respiratory losses subsequently reduce. The ontological significance lies in gross carbon gain establishing the ceiling on vine productivity, where no management practice can increase net carbon gain above the gross photosynthetic input, making optimization of gross carbon gain through canopy management, water supply, and nutrition the first priority for maximizing productive potential before considering how to minimize respiratory losses that erode gross gain. The calculation of gross carbon gain integrates photosynthetic rate across all photosynthetically active leaf area and throughout the photoperiod, with gross gain = Σ(A × leaf area) integrated over time, where A represents leaf-level photosynthetic rate that varies with canopy position, time of day, and environmental conditions. The factors affecting gross carbon gain include total leaf area (canopy size determining light interception capacity), leaf photosynthetic capacity (affected by nitrogen nutrition, water status, and age), canopy architecture (affecting light distribution among leaves), environmental conditions (light, temperature, humidity affecting instantaneous rates), and the duration of the growing season determining integration time. The typical values for gross carbon gain in productive vineyards range from 15-30 tonnes dry matter/hectare/year depending on climate, cultivar, and management, with this gross gain reduced by 40-60% through respiratory losses to yield net primary productivity of 6-15 tonnes/hectare/year. Understanding gross carbon gain quality enables assessment of photosynthetic productivity potential, evaluation of canopy efficiency in capturing available light, and appreciation of the maximum carbon supply that subsequent physiological processes allocate among vine functions.
Gross_Carbon_Gain_Quality addresses the total photosynthetic carbon input before respiratory deductions. This quality establishes the ceiling on vine productivity that determines maximum potential for growth and fruit production. The class supports canopy optimization and productivity potential assessment through gross carbon gain characterization.
Bruttokohlenstoffgewinn
Bruttophotosynthese
GPP
Gesamt-C-Fixierung
Kronenassimilation
canopy photosynthesis
gross photosynthesis
total carbon fixation
Poni S et al. (2006) Effects of early defoliation on shoot photosynthesis, yield components, and grape composition. American Journal of Enology and Viticulture 57(4): 397-407 Schultz HR (1995) Grape canopy structure, light microclimate and photosynthesis. Australian Journal of Grape and Wine Research 1(1): 2-17
Gross Carbon Gain Quality
Gross photosynthesis rate assessment in heat-stressed Merlot reveals the temperature effect on respiratory losses, with leaves at 25°C showing net photosynthesis of 14 μmol CO₂/m²/s and gross photosynthesis of 17 μmol CO₂/m²/s (respiration 3 μmol/m²/s), while leaves at 38°C display reduced net rate of 6 μmol/m²/s but gross rate of 12 μmol/m²/s (enhanced respiration 6 μmol/m²/s), demonstrating how heat stress reduces net carbon gain primarily through increased respiratory losses rather than carboxylation limitation, informing heat mitigation strategies that reduce respiratory carbon costs.
Gross photosynthesis rate quality is a carbon fixation efficiency quality that represents the total rate of CO₂ fixation by Rubisco before accounting for respiratory losses, characterizing the maximum carboxylation activity of the photosynthetic apparatus that determines the upper limit of carbon acquisition from which respiration and photorespiration subtract. Within the BFO framework, this quality specifically addresses the total carboxylation flux, distinguishing gross photosynthesis from net photosynthesis while recognizing that gross rate reveals the true capacity of the photosynthetic apparatus that may be substantially higher than observable net CO₂ uptake due to concurrent respiratory processes. The ontological significance lies in gross photosynthesis providing insight into true photosynthetic capacity independent of respiratory metabolism, where the difference between gross and net rates indicates the respiratory cost of maintaining photosynthetic tissue and the photorespiratory losses that reduce carbon gain efficiency, with gross rate assessment enabling diagnosis of whether limitations reside in carbon fixation capacity or respiratory losses. The physiological basis for gross photosynthesis involves the sum of all Rubisco carboxylation events, with gross rate (Ag) equal to net photosynthesis (An) plus mitochondrial respiration (Rd) plus photorespiration (Rp), and typically estimated as Ag = An + Rd (where Rd is measured in darkness) or calculated from oxygen evolution which is not reduced by respiration during measurement. The quantitative relationships between gross and net photosynthesis show gross rates typically 15-30% higher than net rates in C₃ plants like grapevine under normal conditions, with the difference increasing at higher temperatures (enhanced photorespiration) and representing a substantial carbon cost that C₄ plants and CAM plants largely avoid through CO₂-concentrating mechanisms. The measurement approaches for gross photosynthesis include calculation from net rate plus measured dark respiration, oxygen evolution measurement (not affected by respiratory O₂ consumption), and isotope methods distinguishing CO₂ fluxes in opposite directions. Understanding gross photosynthesis rate quality enables assessment of total carboxylation capacity, quantification of respiratory losses limiting net productivity, and appreciation of the carbon costs associated with C₃ photosynthesis in grapevine.
Gross_Photosynthesis_Rate_Quality addresses total carboxylation before respiratory losses, providing insight into true photosynthetic capacity. This quality requires calculation from net rate plus respiration or specialized measurement techniques. The class supports diagnosis of whether productivity limitations arise from carboxylation capacity or respiratory costs.
Bruttoassimilation
Bruttophotosyntheseraten
Gesamt-CO₂-Fixierung
gross assimilation
total CO₂ fixation
true photosynthesis
wahre Photosynthese
Yin X et al. (2011) Using combined measurements of gas exchange and chlorophyll fluorescence to estimate parameters of a biochemical C₃ photosynthesis model. Plant, Cell & Environment 34(11): 1843-1858 Valentini R et al. (1995) In situ estimation of net CO₂ assimilation, photosynthetic electron flow and photorespiration in Turkey oak leaves. Plant, Cell & Environment 18(6): 631-640
Gross Photosynthesis Rate Quality
A ground-level basal shoot on grafted Vitis vinifera cv. Chardonnay (on 101-14 rootstock) emerging exactly at soil surface level, discovered during cultivation operations, vigorous shoot reaching 172cm length with 21 leaves by mid-July, origin verification required excavating 8cm around shoot base revealing emergence from trunk 3cm above visible graft union confirming scion origin (not rootstock sucker), activated following trunk disease (Esca symptoms in upper portions) and previous winter freeze damage creating combination stress triggering low basal activation, shoot position at soil surface creating management challenges including mechanical damage risk during cultivation, disease exposure from soil proximity, and difficult training access, this shoot tentatively retained as potential renewal option pending further vine assessment determining whether recovery viable or vine removal necessary, with scion origin verification critical for retention decision (rootstock shoots would be removed immediately as unwanted and potentially detrimental).
A basal shoot originating at or very near soil surface level, characterized by emergence from trunk base, crown region, or underground portions at depths within 0-5cm of surface, intermediate position between clearly above-ground trunk shoots and deeply underground rootstock shoots, potential ambiguity regarding scion versus rootstock origin requiring verification particularly in grafted vines, and function as low-level recovery mechanism, potential rootstock sucker requiring removal, or ground-level renewal option with careful origin verification essential before utilization decisions. Ground-level basal shoots represent transition zone between trunk and underground origins, creating classification and management challenges through positional ambiguity, potential rootstock involvement (particularly in grafted vines where graft unions often positioned 5-15cm above soil surface), and difficult origin determination requiring careful excavation or other verification methods before appropriate management decisions possible.
Ground-level basal shoots require careful assessment addressing multiple critical questions including origin verification (scion above graft versus rootstock below graft), activation triggers (damage recovery, stress response, or spontaneous problematic activation), utilization potential (viable renewal source versus removal candidate), and management implications (training feasibility, disease risks, long-term prospects). Origin verification is paramount in grafted vines, requiring excavation exposing shoot emergence point relative to graft union, with scion-origin confirmation necessary before retention consideration while rootstock shoots warrant immediate removal preventing rootstock overgrowth and scion suppression. Ground-level position creates inherent challenges including soil splash contamination, mechanical damage vulnerability, disease exposure, difficult access, and extended training requirements achieving normal canopy heights. Management decisions balance potential renewal value (particularly following severe damage leaving few alternatives) against complications and risks, often favoring removal unless clear need and scion origin confirmed, with retention requiring vigilant disease management, mechanical protection, and patient multi-year training investment.
Kronenausschlag
bodenniveau-Trieb
crown shoot
near-surface basal shoot
soil-level shoot
surface-origin shoot
Ground Level Basal Shoot
In Pinot Noir vines of the Côte d'Or, Burgundy, growing degree day accumulation proceeds through March with daily contributions varying from 0-2 GDD during cool periods with mean temperatures of 8-12°C to 5-8 GDD during warm periods with mean temperatures of 15-18°C, with cumulative GDD reaching the cultivar threshold of approximately 150 units by late March to early April depending on seasonal temperature patterns, triggering budbreak initiation at BBCH stage 05.
Growing degree day accumulation in Vitis vinifera represents the progressive integration of effective heat exposure above the base temperature threshold, quantifying the thermal progress toward budbreak through summation of daily temperature contributions according to established calculation methods. This accumulation process operates continuously during the ecodormancy period following chilling completion, with each day's temperature regime contributing to the cumulative heat total that drives progress toward growth initiation. The calculation of growing degree days employs various methods that integrate temperature above the base threshold, with the simplest approach calculating the difference between daily mean temperature and base temperature when this difference is positive, while more sophisticated methods use hourly temperatures or apply corrections for temperature fluctuation patterns. The base temperature for growing degree day calculation in grapevine budbreak models is typically set at 10°C, representing the threshold below which metabolic activity is insufficient to support meaningful progress toward growth, though some models employ slightly different base temperatures based on regional calibration or cultivar-specific optimization. The daily accumulation rate varies with temperature regime, with warm spring days contributing substantially to the growing total while cool days near the base temperature contribute minimally, creating the weather-dependent variability in accumulation rate that affects budbreak timing prediction accuracy. The cumulative nature of growing degree day accumulation reflects the integration of temperature effects over time, with the developmental progress achieved during warm periods being retained during subsequent cool periods rather than being lost, allowing gradual progress toward budbreak even when temperatures fluctuate around the base threshold. The relationship between accumulated growing degree days and developmental stage is approximately linear for budbreak prediction, with cultivar-specific thresholds typically falling in the range of 100-300 GDD for the transition from dormant bud to visible bud swelling at BBCH stage 05. The practical application of growing degree day accumulation involves tracking cumulative heat units through spring and comparing to cultivar thresholds to predict budbreak timing, enabling proactive management decisions regarding frost protection deployment and early season cultural operations.
Growing_Degree_Day_Accumulation represents the quantitative tracking method within Heat_Accumulation_Response. Distinguished from Temperature_Sum_Response which addresses the physiological integration, while Growing_Degree_Day_Accumulation specifically addresses the calculation methodology. Base temperature typically 10°C. Daily accumulation rate varies with temperature from 0 to 10+ GDD. Cumulative thresholds for budbreak typically 100-300 GDD depending on cultivar. Enables budbreak prediction from temperature monitoring.
GDD accumulation
GDD-Akkumulation
Gradtag-Summierung
Wachstumsgradtag-Akkumulation
Wärmesummen-Akkumulation
degree day summation
heat sum accumulation
thermal time accumulation
Moncur MW et al (1989) Base temperatures for budbreak and leaf appearance of grapevines. American Journal of Enology and Viticulture 40: 21-26 Parker AK et al (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17: 206-216 McMaster GS, Wilhelm WW (1997) Growing degree-days: one equation, two interpretations. Agricultural and Forest Meteorology 87: 291-300
Growing Degree Day Accumulation
The growing degree-day accumulation for the 2022 vintage at Geisenheim (Rheingau, Germany, 49.98°N, 86 m elevation) documents a total of 1,520 GDD base 10°C over the April 1 - October 31 period — this was 15% above the 1991-2020 normal of 1,320 GDD, reflecting the exceptionally warm growing season with prolonged heat events in June and July that accelerated phenological development by 10-14 days compared to the long-term average. The elevated GDD accumulation enabled Riesling to reach optimal harvest maturity (85-90°Oechsle, equivalent to 20-22°Brix) by mid-October with no difficulty, and even the traditionally marginal late-ripening Cabernet Sauvignon experimental plantings achieved acceptable maturity (80°Oechsle) for the first time in the Rheingau — illustrating how a single warm vintage can temporarily shift the effective climate zone and expand the range of ripening cultivars.
A growing season data component that records the cumulative heat unit accumulation above a defined base temperature over a specified time period, calculated as the sum of daily contributions where each day's contribution is the daily mean temperature minus the base temperature (with negative values set to zero), providing the standardized thermal summation index that quantifies the total heat energy available for vine development and enables comparison of viticultural thermal resources across sites, regions, and vintages.
Growing_Degree_Days is the parent class for the specific GDD calculation methods (base 10°C, Winkler, Huglin, BEDD) that differ in their base temperature, calculation period, daily temperature treatment, and daylength adjustment but share the fundamental principle of cumulative heat summation as a predictor of biological development rate. The physiological basis of the GDD concept is the approximately linear relationship between temperature and the rate of plant developmental processes (cell division, cell expansion, enzymatic reactions) within the range of 10-30°C — below the base temperature (~10°C for V. vinifera), development effectively ceases, while above the base temperature, the rate of development increases approximately proportionally with temperature: the cumulative sum of temperature exceedances above the base temperature therefore estimates the total developmental progress achieved over the growing season. The practical utility of GDD lies in its predictive power — the thermal requirements for specific phenological events are relatively consistent across years for a given cultivar (e.g., Cabernet Sauvignon typically requires 1,200-1,400 GDD base 10°C from budburst to harvest), enabling the prediction of phenological timing from temperature forecasts and the assessment of cultivar maturation potential from historical climate data. The limitations of GDD include the assumption of linearity (the actual temperature-development relationship is curvilinear, with reduced developmental progress above approximately 30°C due to enzyme denaturation and stomatal closure), the insensitivity to temperature distribution (the same GDD total can be achieved by moderate temperatures over a long season or by high temperatures over a short season, with different biological outcomes), and the neglect of other climatic variables (radiation, moisture, humidity) that influence vine development independently of temperature.
Gradtage
Temperatursummen-Index
Wärmesumme
degree-day total
heat unit accumulation
thermal summation index
Winkler AJ et al. (1974) General Viticulture, University of California Press Gladstones J (1992) Viticulture and Environment, Winetitles Jones GV et al. (2005) Climate change and global wine quality, Climatic Change 73: 319-343
Growing Degree Days
Merlot grafted on 3309 Couderc rootstock monitored via minirhizotron showing growing primary root tip, sequential images documenting primary root tip position at 48-hour intervals, elongation rate calculated 4.8 mm per day during early June observation period, growing primary root tip advancing through loam soil at 82 cm depth, root tip maintaining downward gravitropic trajectory, growing tip producing continuous trail of newly formed root tissue behind advancing meristem, elongation continuing over 6-week observation period adding 201 mm total length to primary root, growing primary root tip demonstrating successful soil penetration and resource exploration
Growing primary root tip is primary root tip showing measurable increase in length over time, emphasizing elongation process and growth dynamics, characterized by active meristem producing new cells through division, elongation zone where cells increase length through expansion, demonstrable length increment typically 2-15 mm per day depending on root order temperature and conditions, and visible advancement of root tip position through soil over observation periods of hours to days, with growing primary root tip representing functional growing state distinguished from structurally present but dormant or non-growing tip, showing characteristic white coloration of young growing tissue, producing continuous trail of newly formed root tissues behind advancing tip including epidermis cortex endodermis and vascular cylinder, responding to environmental gradients including gravity moisture and nutrients affecting growth rate and direction, and demonstrating that growing primary root tips enable primary root length increase which accumulates over weeks and months producing deep extensive primary root framework essential for vine anchorage water access and soil resource exploitation. Growing primary root tip elongation rates includes= showing rapid growth in young vine (primary root tips potentially elongating 8-15 mm per day in first year during optimal spring conditions), exhibiting moderate growth in mature vine (established vine primary root tips showing 3-8 mm daily elongation during active periods), demonstrating temperature effects (elongation rate increasing with soil temperature from threshold 8-10°C to optimum 20-25°C declining above 28-30°C), revealing moisture influence (adequate soil moisture supporting maximum elongation while deficit reducing rates and severe drought halting growth), and displaying seasonal variation (highest elongation rates spring and fall declining summer and ceasing winter). Growing primary root tip growth measurement includes= using sequential observation (marking root tip position and measuring displacement over 24-48 hours documenting growth rate), employing minirhizotron imaging (video cameras in soil tubes capturing root tip advancement providing non-destructive growth documentation), demonstrating excavation methods (carefully exposing roots at intervals measuring length increase though destructive), revealing tracer techniques (pulse-labeling with isotopes documenting incorporation of new material into growing tip), and showing model calculations (growth rates estimated from root length distributions and turnover data though indirect). Growing primary root tip growth zones includes= comprising meristem cell production (apical meristem generating new cells through division contributing to length increase), including elongation zone expansion (recently divided cells increasing length 10-50 fold through vacuolation contributing majority of elongation), demonstrating differential contribution (elongation zone typically contributing 70-90% of length increase while meristem 10-30%), revealing zone lengths (meristem 0.5-0.8 mm elongation zone 3-10 mm determining growing tip total active length), and showing zone dynamics (zones maintaining relatively constant length as tip advances with cells continuously entering elongation from meristem and exiting into differentiation). Growing primary root tip directional control includes= demonstrating gravitropic growth (growing primary root tips maintaining downward trajectory through positive gravitropic response), showing hydrotropic response (growing tips potentially deflecting toward moisture when water gradient present), exhibiting thigmotropic avoidance (growing tips potentially growing around obstacles when encountering mechanical impedance), revealing chemotropic responses (growing tips possibly responding to nutrient or oxygen gradients affecting direction), and displaying combined responses (multiple environmental cues integrated determining actual growth trajectory through soil). Growing primary root tip carbon economy includes= requiring substantial photosynthate (growing primary root tips consuming 15-30% of total root carbon allocation supporting division and elongation), competing with shoots (primary root tip growth potentially reduced during peak shoot and fruit growth periods when competing for limited carbohydrates), demonstrating investment return (carbon invested in growing primary root tips producing permanent structural framework amortized over years), showing seasonal allocation (primary root tip receiving high allocation spring before shoot flush and fall after harvest when shoot demand reduced), and revealing efficiency variation (faster-growing tips more carbon-efficient producing more root length per gram carbon invested than slower tips). Growing primary root tip and soil conditions includes= responding to soil strength (growing primary root tips showing reduced elongation in compacted soil unable to penetrate when strength exceeds root pressure capacity), affected by soil structure (well-aggregated soil with continuous pores facilitating rapid primary root tip advancement), influenced by moisture (adequate but not saturated moisture optimal for growing tip elongation with both deficit and excess reducing rates), showing pH sensitivity (growing primary root tips potentially reduced in strongly acid pH 8 soils), and demonstrating nutrient effects (adequate nutrients supporting vigorous growing tip elongation while deficiencies or toxicities reducing growth). Growing primary root tip lifespan includes= maintaining growth for extended periods (individual primary root tip potentially growing continuously for weeks to months during favorable season), eventually slowing or ceasing (growing primary root tip elongation declining as root ages or conditions deteriorate), demonstrating seasonal limits (growing period for primary root tips typically 4-8 months per year in temperate climates with dormancy interrupting), showing stress termination (growing primary root tip halting elongation under severe drought, temperature extremes, or damage), and revealing persistence (primary root once established continuing growth from tip over multiple years adding length annually). Growing primary root tip in root system development includes= producing primary root length (cumulative elongation of growing primary root tips determining total primary root length and depth), enabling soil exploration (growing tips advancing into unexplored soil bringing root system into contact with new resources), demonstrating establishment importance (vigorous growing primary root tips during establishment year critical for developing adequate root system), showing continued expansion (growing primary root tips maintaining root system growth throughout vine life adapting to changing conditions), and revealing management impact (practices affecting growing primary root tip function directly influencing root system development and vine performance).
Growing_Primary_Root_Tip emphasizes measurable elongation and length increase and is distinguished from static or dormant primary root tips by focus on demonstrable growth over time. Critical for understanding elongation dynamics, growth measurement, soil advancement, directional control, seasonal patterns, and the elongation process enabling primary root length accumulation producing deep extensive structural root framework.
Advancing primary root apex
Length-increasing primary tip
Längen-zunehmende Primärwurzel
Sich verlängernde Hauptwurzel-Spitze
Wachsende Primäre Wurzelspitze
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Malamy, J. E. (2005). Intrinsic and environmental response pathways that regulate root system architecture. Plant, Cell & Environment, 28(1), 67-77.
Growing Primary Root Tip
The growing season data for Burgundy (Beaune, 47.02°N, 215 m elevation) documents a growing season extending from approximately April 5 (mean budburst date for Pinot Noir) through September 20 (mean harvest date), a duration of approximately 168 days, with mean growing season temperature of 16.2°C (1991-2020 normal), accumulated GDD base 10°C of 1,250-1,400 (Winkler Region I-II boundary), Huglin index of 1,750-1,950 (cool-temperate viticultural climate), and a mean diurnal temperature range of 10-12°C during the July-September ripening period. This thermal profile places Burgundy at the cool margin of reliable Pinot Noir maturation — the growing season provides just sufficient heat accumulation for Pinot Noir to achieve 12.0-13.5% potential alcohol in average vintages, with cool vintages (1991-2020 frequency approximately 20%) producing underripe fruit requiring chaptalization, and warm vintages (frequency approximately 30%) producing fully ripe fruit at natural potential alcohol of 13.5-14.5% — the climate change trend has shifted this balance, with underripe vintages becoming progressively rarer and warm vintages more frequent since the 1980s.
A climate data component that documents the temporal extent and thermal characteristics of the period during which grapevine vegetative and reproductive development occurs — from budburst through harvest — recording the start and end dates of the growing season, its total duration in days, the accumulated heat units (growing degree-days by various calculation methods), and the temperature distribution parameters (mean, maximum, minimum, diurnal amplitude) that together quantify the thermal resources available for vine development and fruit maturation at a specific site.
Growing_Season_Data provides the temporal and thermal characterization of the period that determines all vine developmental outcomes — the growing season is the fundamental time unit of viticulture within which budburst, shoot growth, flowering, fruit set, véraison, ripening, and harvest must be completed, and the thermal resources accumulated during this period determine whether each developmental stage can proceed at the rate required for the subsequent stages to complete before autumn temperature decline terminates the season. The definition of growing season boundaries varies between classification systems — the meteorological growing season is commonly defined as the period with daily mean temperatures consistently above 10°C (the approximate base temperature for V. vinifera growth), while the phenological growing season is defined by actual vine developmental events (budburst to harvest or budburst to leaf fall), and the frost-free growing season is defined by the last spring frost to first autumn frost dates: these definitions can differ by 2-4 weeks, and the VVO documents the definition used alongside the recorded dates to ensure interpretive clarity. The growing season length is a critical viticultural parameter independent of heat accumulation — even if total GDD is adequate, an excessively compressed growing season (short duration with high daily temperatures) produces qualitatively different fruit than an extended growing season (long duration with moderate daily temperatures), because the rate of sugar accumulation relative to the rate of flavor compound development differs between rapid-hot and slow-cool ripening trajectories.
Vegetationsperioden-Daten
Wachstumsperioden-Charakterisierung
growing season thermal record
phenological season documentation
phänologische Saisondaten
vegetation period characterization
Jones GV et al. (2005) Climate change and global wine quality, Climatic Change 73: 319-343 Gladstones J (1992) Viticulture and Environment, Winetitles Winkler AJ et al. (1974) General Viticulture, University of California Press Duchêne E and Schneider C (2005) Grapevine and climatic changes, Agronomy for Sustainable Development 25: 93-99
Growing Season Data
Growing season precipitation in Napa Valley averages only 50-80 mm (April-October), with the Mediterranean climate concentrating 85-90% of annual rainfall in winter months, making supplemental irrigation essential for vine health. Burgundy receives 350-400 mm during the April-September growing season, generally sufficient for unirrigated viticulture on limestone-derived soils with moderate water-holding capacity. The Rheingau records 250-300 mm of growing season precipitation (April-October), typically adequate for Riesling on the deep loess soils overlying phyllite bedrock, though increasingly variable summers create vintage-dependent irrigation needs.
Growing season precipitation is a precipitation data entity that records the cumulative atmospheric water deposition received at a viticultural site during the active vegetative and reproductive growth period of the grapevine, typically measured from budburst through harvest (approximately April-October in the Northern Hemisphere, October-April in the Southern Hemisphere), expressed in millimeters (mm), providing a direct indicator of vine water supply during the physiologically critical period.
Growing season precipitation is arguably more viticulturally meaningful than annual precipitation, as it directly quantifies water availability during the period of active vine physiological demand. The critical thresholds vary substantially by soil type and rootstock — sandy soils with low water-holding capacity may require 500+ mm of growing season rainfall for unirrigated viticulture, while deep clay-loam soils may sustain vines on 250 mm. The timing within the growing season further modulates the impact: rainfall during flowering (BBCH 60-69) reduces fruit set, rainfall at véraison through harvest (BBCH 81-89) dilutes sugars and promotes Botrytis, while moderate rainfall during the lag phase (BBCH 71-79) supports cell division in developing berries. Climate change projections suggest significant redistribution of growing season precipitation patterns even where annual totals remain stable.
Niederschlag in der Vegetationszeit
Niederschlag während der Wachstumsperiode
Vegetationsperioden-Niederschlag
Wachstumszeitniederschlag
growing period rainfall
in-season precipitation
seasonal precipitation total
vegetative season rainfall
Van Leeuwen C et al. (2004) Influence of climate, soil, and cultivar on terroir. American Journal of Enology and Viticulture 55(3): 207-217 Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 327-350 Coombe BG, Dry PR (2004) Viticulture Volume 1: Resources, 2nd Ed, Winetitles, pp.: 64-82
Growing Season Precipitation
Chardonnay grafted on Teleki 5C rootstock growing season (April 28-October 15) showing growing season rootstock vascular cambium, cambium reactivation detected late April with first periclinal divisions resuming after winter dormancy, peak cambial activity observed late May-early June producing widest earlywood vessels, sustained activity through July-August producing transition and latewood zones, declining activity September with latewood completion, cambium entering dormancy mid-October as temperatures declined below 10°C, complete growing season producing 0.44 cm xylem ring and 1.1 mm phloem increment documenting 5.5 month functional period
Growing season rootstock vascular cambium is rootstock vascular cambium during annual active growth period typically April-October in Northern Hemisphere, emphasizing temporal phase rather than instantaneous activity state, characterized by capacity for cell division and derivative production throughout growing season even if division rate varying from peak in spring to minimal in late summer, showing reactivation from winter dormancy in spring triggered by warming temperatures and hormonal signals, maintaining division capacity through summer producing complete annual xylem ring and corresponding phloem increment, declining toward dormancy in fall as temperatures cool and photoperiod shortens, and representing functional cambial state enabling annual diameter increase distinguishing growing season from dormant winter period when cambium quiescent. Growing season rootstock vascular cambium temporal extent includes= beginning in spring (cambium exiting dormancy and resuming division capacity April-May as temperatures rise above threshold), spanning active growth months (cambial function continuing through spring and summer April-September producing bulk of annual increment), extending into early fall (cambium potentially maintaining some division capacity into October completing annual ring), concluding before winter (cambial activity ceasing October-November as dormancy induced), and comprising approximately 6-7 months (growing season cambium functional roughly half year in temperate climates with longer seasons in warm regions and shorter in cool regions). Growing season rootstock vascular cambium reactivation includes= triggered by temperature (cambium resuming division when temperatures exceed threshold typically 10-15°C for several consecutive days), influenced by photoperiod (lengthening days in spring contributing to dormancy break and activity resumption), mediated by hormones (auxin from expanding buds signaling cambium to resume division), showing progressive activation (cambium reactivating first near buds then basipetally toward roots), and requiring metabolic preparation (cambium cells accumulating energy reserves and synthesizing division machinery before first mitosis). Growing season rootstock vascular cambium activity phases includes= showing early-season peak (maximum division rate May-June producing earlywood with large vessels), exhibiting mid-season sustained activity (continued division June-July producing transition wood between earlywood and latewood), demonstrating late-season decline (reduced division August-September producing latewood with small vessels and thick walls), approaching dormancy (minimal division September-October as cambium prepares for winter quiescence), and maintaining readiness (cambium retaining capacity for division throughout growing season able to respond to favorable conditions or stress relief). Growing season rootstock vascular cambium complete ring formation includes= producing earlywood phase (spring cambial activity creating inner light-colored portion of annual ring with large vessels), generating latewood phase (late summer cambial activity creating outer dark-colored portion of annual ring with small vessels and thick fibers), completing annual increment (growing season cambium producing full xylem ring from first spring vessels to final fall fibers), forming ring boundary (cessation of cambial activity at season end creating distinct line between current year latewood and next year earlywood), and generating phloem layer (growing season cambium producing corresponding phloem increment outward accompanying xylem ring formation). Growing season rootstock vascular cambium environmental sensitivity includes= responding to water availability (adequate moisture supporting sustained cambial activity while drought potentially halting division mid-season), affected by temperature patterns (optimal temperatures 20-25°C promoting maximum activity while extremes reducing division), influenced by nutrient supply (adequate nitrogen, phosphorus, and other nutrients supporting cambial metabolism and division), modulated by canopy development (expanding leaves producing auxin stimulating cambial activity and supplying photosynthates), and showing stress responses (severe stress potentially forcing premature dormancy ending growing season cambial activity early). Growing season rootstock vascular cambium in different climates includes= showing long season in warm climates (Mediterranean or mild temperate regions with 7-9 month growing seasons enabling extended cambial activity), exhibiting moderate season in temperate climates (continental regions with 5-7 month growing seasons typical for commercial viticulture), demonstrating short season in cool climates (northern or high-elevation sites with 4-6 month growing seasons limiting annual growth), varying with elevation (higher elevations showing later start and earlier end reducing growing season cambium duration), and responding to local conditions (microclimate, slope aspect, and site factors affecting growing season length and cambial activity period). Growing season rootstock vascular cambium and vine phenology includes= correlating with bud break (cambial reactivation coinciding with bud expansion and shoot emergence), aligning with flowering (peak cambial activity occurring during bloom period May-June), overlapping with fruit development (sustained cambial activity through berry growth and early ripening phases), declining during ripening (cambial activity decreasing as berries complete maturation and canopy senesces), and ending before leaf fall (cambium entering dormancy typically before complete leaf abscission).
Growing_Season_Rootstock_Vascular_Cambium emphasizes temporal active phase April-October and is distinguished from dormant winter cambium by focus on growth period with division capacity. Critical for understanding seasonal growth dynamics, annual ring formation timing, phenological correlation, climate effects on growth duration, and the functionally active phase producing annual diameter increment.
Functional period cambium
Saisonales Wurzelstock-Kambium
Seasonal rootstock vascular cambium
Spring-fall rootstock cambium
Vegetationsperiode Unterlagen-Gefäßkambium
Wachstumszeit-Kambium
Growing Season Rootstock Vascular Cambium
The growing season temperature for the world's major Pinot Noir regions illustrates the cultivar-specific thermal niche that Jones identified — Burgundy (Beaune) GST 16.0-16.5°C, Willamette Valley (McMinnville) GST 15.5-16.5°C, Central Otago (Cromwell) GST 14.5-15.5°C, Sonoma Coast (Occidental) GST 14.5-15.5°C, Marlborough (Blenheim) GST 15.5-16.5°C, Tasmania (Launceston) GST 14.5-15.5°C — all clustering within the 14-17°C range that Jones defined as the optimal growing season temperature band for premium Pinot Noir production. Regions with GST below 14°C (southern England at 13.5°C) produce Pinot Noir only in warm vintages and predominantly for sparkling wine base, while regions with GST above 17.5°C (Barossa Valley at 19°C) produce Pinot Noir that lacks the aromatic finesse and structural elegance associated with cool-climate Pinot Noir — demonstrating the predictive power of the GST metric for cultivar-climate matching.
A temperature data component that records the mean temperature over the grapevine growing season — conventionally April through October in the northern hemisphere and October through April in the southern hemisphere — providing the most directly relevant single thermal index for predicting cultivar maturity potential, wine style, and vintage quality, with Jones (2006) identifying the 12-22°C growing season mean temperature range as the global envelope of quality wine production and establishing the cultivar-specific optimal temperature bands within this range.
Growing_Season_Temperature is arguably the single most useful climatic metric for viticultural site characterization because it captures the thermal resources available during the biologically active period while excluding the dormant winter months that contribute to MAT but have minimal relevance to vine development and fruit composition. Jones's (2006) comprehensive analysis of wine quality ratings, cultivar distribution, and growing season temperatures across the world's wine regions established GST as the preferred single-variable predictor of viticultural climate suitability, with the 12-22°C range encompassing virtually all quality wine production and the cultivar-specific optimal bands providing actionable guidance for site-cultivar matching: the analysis identified optimal GST ranges for 21 major cultivars — Müller-Thurgau 13-15°C, Riesling 13-16°C, Pinot Noir 14-17°C, Chardonnay 14.5-17.5°C, Sauvignon Blanc 13.5-16.5°C, Cabernet Sauvignon 16-19°C, Syrah 16-19°C, Grenache 17-20°C, Zinfandel 17-20°C — ranges that correspond closely with the observed global distribution of these cultivars and with the regions producing the most consistently highly-rated wines from each cultivar. The GST metric also serves as the primary indicator for climate change impact assessment — the observed warming trend of approximately 0.3°C per decade in growing season temperature across European wine regions since 1980 translates directly into predictions of cultivar suitability shift, with each 1°C increase in GST equivalent to a poleward shift of approximately 150-200 km or an elevation shift of approximately 150 m.
GST record
Vegetationsperioden-Temperatur
Wachstumszeittemperatur
growing season mean temperature
mittlere Vegetationstemperatur
vegetation period temperature
Jones GV (2006) Climate and terroir, in Macqueen RW and Meinert LD (eds) Fine Wine and Terroir, Geological Association of Canada Jones GV et al. (2005) Climate change and global wine quality, Climatic Change 73: 319-343 Gladstones J (1992) Viticulture and Environment, Winetitles Hall A and Jones GV (2009) Effect of potential atmospheric warming on temperature-based indices describing Australian winegrape growing conditions, Australian Journal of Grape and Wine Research 15: 97-119
Growing Season Temperature
Cabernet Sauvignon growing season vascular cambium in Napa Valley shows reactivation late March (synchronized with budbreak), peak activity June-July producing 1.8-2.6 mm xylem increment (60-65% of annual 3.2-4.0 mm ring), declining activity August-September producing latewood with smaller vessels and higher fiber content (1.2-1.6 mm representing 35-40% of ring), cessation early October as temperatures decline and day length shortens, total growing season duration 27-29 weeks producing complete annual ring plus 0.3-0.5 mm phloem increment, season ending with dormancy entry and cold hardiness development
Growing season vascular cambium is vascular cambium during the annual growth period from spring reactivation through fall cessation, encompassing the complete cycle of cambial activity from dormancy break to dormancy re-entry, representing the temporally defined active phase when secondary xylem and phloem production occurs. Growing season vascular cambium temporal extent includes= spring reactivation (late March-April in Northern Hemisphere temperate climates, triggered by rising temperatures and hormonal signals from budbreak), active growth phase (May-August with peak activity June-July during maximum shoot growth), late season activity (September with declining division rates as growth slows), fall cessation (late September-October as cambium enters dormancy), and total growing season duration typically 24-28 weeks (6-7 months) varying by climate (shorter 18-22 weeks in cool climates, longer 28-32 weeks in warm climates). Growing season vascular cambium reactivation triggers include= temperature accumulation (sustained daily mean temperatures >10-15°C for 1-2 weeks triggering reactivation), hormonal signals (auxin produced in expanding buds transported basipetally activating cambium 0-2 weeks before or after budbreak), photoperiod lengthening (increasing day length in spring contributing to dormancy release), carbohydrate availability (stored reserves mobilized from trunk and root providing energy for cambial reactivation), and water availability (adequate soil moisture necessary for cell turgor and division). Growing season vascular cambium activity phases include= early season phase (April-May, reactivation and earlywood formation, large vessel production supporting spring canopy development), peak season phase (June-July, maximum division rates producing wide xylem increments, transition from earlywood to latewood), mid-late season phase (August-September, continued latewood formation with declining division rates, smaller vessels and more fibers), and cessation phase (September-October, progressive slow-down and eventual dormancy entry preparing for winter). Growing season vascular cambium xylem production timeline includes= earlywood formation (April-June producing 35-50% of annual ring width with large vessels 150-300 μm diameter), latewood formation (June-September producing 50-65% of annual ring width with smaller vessels 40-120 μm and higher fiber content), total annual ring width 1.5-5 mm depending on vigor and conditions (young vigorous vines 3-6 mm, mature moderate vines 2-4 mm, old declining vines 0.8-2.5 mm), and ring completion (late September-October when cambium ceases producing final latewood cells marking ring boundary). Growing season vascular cambium phloem production includes= continuous phloem formation throughout season (cambium producing new sieve tubes, companion cells, parenchyma, and fibers), annual phloem increment 0.2-0.8 mm radial width (much less than xylem due to asymmetric cambial division), functional phloem zone (most recent 1-2 years production maintaining active transport 0.4-1.5 mm depth), older phloem crushed (phloem produced 3-5+ years previously compressed and non-functional from xylem expansion pressure), and total living phloem depth 2-4 mm (comprising recent 3-5 years production still containing living cells). Growing season vascular cambium environmental modulation includes= water availability strongly affecting season length and intensity (drought causing premature cessation or intermittent activity, irrigation extending season and increasing production), temperature effects (cool springs delaying reactivation, warm autumns extending season, heat waves potentially causing mid-season pauses), nutrient availability (adequate nitrogen supporting vigorous cambial activity throughout season, deficiency reducing production rates), crop load effects (heavy crop potentially reducing cambial activity through photosynthate competition, light crop allowing enhanced vegetative growth), and stress events (severe stress causing temporary cessation, recovery allowing resumed activity if season permits). Growing season vascular cambium variation by vine age includes= young vines (years 1-5) showing extended vigorous season (26-30 weeks active period producing 4-7 mm annual rings), mature vines (years 8-25) showing moderate season (24-28 weeks producing 2-4 mm rings), old vines (years 30-50+) showing shortened season (20-26 weeks producing 1-2.5 mm rings reflecting declining vigor), and very old vines (60-100+ years) showing brief season (18-24 weeks producing 0.5-1.8 mm rings with intermittent activity). Growing season vascular cambium geographic variation includes= cool temperate regions (northern Europe, cool California regions) showing 20-26 week seasons (late April-September), moderate temperate regions (Bordeaux, Burgundy, Oregon) showing 24-28 week seasons (late March-October), warm temperate regions (southern France, warm California, Argentina) showing 26-32 week seasons (mid-March-November), and Mediterranean climates sometimes showing bimodal activity (spring peak April-June, reduced summer activity during drought stress, potential fall resumption September-October with rain). Growing season vascular cambium cessation process includes= progressive division rate decline (from peak 1-2 divisions/week June to 0.5-1 division/week September to 0 divisions October), final latewood cell production (last small-diameter vessels and fibers formed late September-October), cessation of cell expansion (newly formed cells completing expansion and wall deposition), transition to dormant state (cambial cells entering quiescence with reduced metabolism), and dormancy establishment (cambium becoming cold-hardy and insensitive to short-term warming periods requiring extended chilling for spring reactivation). Growing season vascular cambium significance includes= determining annual trunk growth (growing season length and intensity controlling annual ring width and total diameter increase), defining wood characteristics (seasonal cambial activity dynamics creating earlywood-latewood patterns in ring-porous structure), responding to management (cultural practices during growing season affecting cambial productivity), enabling dendrochronological reconstruction (ring patterns recording growing season conditions across years), and reflecting climate impacts (growing season characteristics varying with temperature, precipitation, and other climatic factors).
Growing_Season_Vascular_Cambium emphasizes temporal growth period and is distinguished from Active_Vascular_Cambium (cellular activity emphasis) and Vascular_Cambium (cambium in all states) by focus on annual cycle from reactivation through cessation. Critical for understanding seasonal growth dynamics, annual ring formation timeline, environmental effects on growth season length, climate-growth relationships, and dendrochronological interpretation of ring patterns.
Active season cambium
Growth period cambium
Saisonales vaskuläres Kambium
Seasonal cambium
Vegetationsperiode-Kambium
Wachstumssaison-Kambium
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Larson, P. R. (1994). The Vascular Cambium= Development and Structure. Springer.
Growing Season Vascular Cambium
A growing tip leaf at node 14 on a vigorously extending shoot of Vitis vinifera cv. Syrah in the Rhône Valley, France, positioned 6 cm behind the shoot apex within the active growth zone, measuring 5.8 cm length by 5.2 cm width representing mid-expansion stage at approximately 35% of expected mature dimensions, exhibiting active cell expansion with cells increasing from average 18 μm to 24 μm diameter over 3-day observation period, light green coloration with chlorophyll content of 245 μg cm⁻² still accumulating, soft flexible texture easily deformed by handling, developing tissue organization with emerging palisade layer of 38 μm and loosely organized spongy mesophyll, procambial strands differentiating into recognizable veins with beginning secondary wall deposition, stomata beginning to function showing partial closure response to VPD increase though with reduced sensitivity, thin developing cuticle creating slight sheen on adaxial surface, demonstrating photosynthesis rate of 3.8 μmol CO₂ m⁻² s⁻¹ representing 29% of mature capacity and remaining strongly net sink consuming approximately 15 mg carbohydrate per day while producing only 5 mg through limited photosynthesis, occurring at 9 days post-emergence during rapid expansion phase, measured during active shoot extension at 7-9 cm per week growth rate at shoot BBCH stage 73 (berries beginning to touch) in early July, representing typical growing tip leaf in active development within the shoot extension zone.
An apical leaf within the active growth zone at the shoot terminus in Vitis vinifera, characterized by position within 5-10 cm of the shoot apex representing the zone of active internode elongation and leaf expansion, very recent emergence within previous 1-2 weeks, small to intermediate dimensions typically 2-8 cm length representing early to mid-expansion stages, active growth processes including ongoing cell division in marginal meristems and rapid cell expansion throughout lamina, pale to medium green coloration reflecting developing chlorophyll systems, soft flexible texture from thin cell walls and high water content, incomplete tissue differentiation with developing mesophyll layers showing progressive organization, immature vascular development with procambial differentiation proceeding, beginning stomatal function with partial environmental responsiveness, minimal cuticle development, low to moderate photosynthetic capacity typically 20-50% of mature leaf rates, net carbon sink status consuming resources for continued expansion and development, and functional integration with the growing tip as part of the active extension zone. Growing tip leaves represent the developmental transition zone between primordial emergence and approaching maturity, showing active progression through expansion phases, experiencing the protected favorable microenvironment of the shoot tip region including elevated humidity, moderated temperature, and mechanical protection, consuming substantial resources supporting the rapid growth processes that will determine final leaf size and characteristics.
Growing tip leaves occupy the active developmental zone immediately behind terminal positions, distinguished from terminal leaves by somewhat advanced development and from upper canopy leaves by very recent emergence and active expansion. The growing tip zone extends 5-10 cm from apex depending on internode lengths and shoot vigor, encompassing 2-4 leaves in various expansion stages from early unfolding through mid-expansion. These leaves experience the protected shoot tip microenvironment favoring rapid development, with gradually increasing exposure to ambient conditions as internodes elongate displacing leaves from the immediate tip zone. Active growth processes including cell division in marginal zones and rapid cell expansion throughout lamina consume substantial resources, with daily carbohydrate consumption often exceeding 10-20 mg per leaf despite small current size. Developmental progression depends critically on favorable conditions including adequate water for turgor-driven expansion, sufficient nitrogen for protein synthesis, and moderate temperatures supporting metabolic activity. Stress during growing tip leaf expansion affects final size and characteristics, with water deficit reducing cell expansion creating smaller mature leaves, nutrient limitation affecting tissue quality, and temperature extremes potentially arresting development.
Expansionsphase apikales Weinblatt
Wachstumsspitzen Blatt
active growth zone leaf
aktive Wachstumszone Blatt
developing tip leaf
expansion phase apical leaf
Based on Palliotti & Cartechini (2000) "Developmental changes in gas exchange activity in flowers, berries, and tendrils of field-grown Cabernet Sauvignon" American Journal of Enology and Viticulture 51=317-323 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 86-108 Tsukaya (2013) "Leaf development" The Arabidopsis Book 11=e0163 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 103-154.
Growing Tip Leaf
Comprehensive characterization of Growth_And_Development_Disposition at UC Davis compared growth parameters across 50 cultivars grown under standardized conditions (own-rooted, 2.4 × 1.5 m spacing, drip irrigation at 80% ETc) over four growing seasons. Measurements included shoot elongation rate, final shoot length, pruning weight, leaf area development, internode characteristics, and canopy architecture metrics. Growth_And_Development_Disposition showed substantial cultivar variation across multiple parameters. High-vigor cultivars like 'Barbera' and 'Carignane' produced mean pruning weights of 2.8-3.2 kg/vine, shoot lengths exceeding 250 cm, and dense canopies requiring multiple hedging passes. Moderate-vigor cultivars including 'Cabernet Sauvignon' and 'Merlot' showed pruning weights of 1.5-2.0 kg/vine and shoot lengths of 150-200 cm, balancing vegetative and reproductive development effectively. Low-vigor cultivars such as 'Pinot Noir' and 'Gewürztraminer' produced pruning weights of 0.8-1.2 kg/vine with shoots of 100-140 cm, often requiring vigor-enhancing rootstocks to achieve adequate canopy development. The stability of Growth_And_Development_Disposition rankings across years (r = 0.92 between years for pruning weight) confirmed strong genetic determination. Importantly, cultivars also differed in growth habit (erect vs. trailing), lateral shoot production, and the distribution of growth over the season, demonstrating that Growth_And_Development_Disposition encompasses multiple partially independent growth characteristics beyond simple vigor classification.
A Growth_And_Development_Disposition is a Grapevine_Disposition that inheres in a Vitis vinifera plant and is realized through the inherent patterns, rates, and characteristics of vegetative and reproductive growth that determine vine architecture, canopy development, and the temporal progression through developmental stages from dormancy through harvest. This disposition represents the fundamental growth program of the grapevine genotype, encompassing vigor (the intensity of vegetative growth), growth habit (the spatial organization of shoots and canopy), developmental timing (phenological progression), and the balance between vegetative and reproductive development. Growth_And_Development_Disposition integrates genetic determinants of meristem activity, hormone signaling, carbon allocation priorities, and environmental responsiveness that together produce the characteristic growth phenotype of each cultivar. Understanding Growth_And_Development_Disposition is essential for vineyard design (training system selection, vine spacing), canopy management (shoot positioning, hedging, leaf removal requirements), and predicting cultivar performance across different environments. The disposition is modified by environmental factors (climate, soil, water availability), management practices (pruning severity, irrigation, fertilization), and in grafted vines by rootstock influence on scion vigor. However, the underlying genetic disposition establishes the baseline growth characteristics that management can modulate but not fundamentally transform—a high-vigor cultivar remains inherently vigorous even under restrictive conditions, while a low-vigor cultivar cannot be induced to match high-vigor growth even with optimal resources.
Growth_And_Development_Disposition is_a Grapevine_Disposition and represents a major category encompassing vegetative growth characteristics. Child classes include Vigor_Disposition (growth intensity), Growth_Habit_Disposition (spatial organization), Phenological_Timing_Disposition (developmental timing), and potentially others addressing specific growth aspects. Growth disposition is genetically determined but environmentally modulated. In grafted vines, rootstock significantly influences scion growth expression. Growth characteristics have major implications for training system selection and canopy management requirements.
Entwicklungs-Charakteristik
Vegetatives-Wachstumsprogramm
Wachstums-und-Entwicklungs-Disposition
developmental program
growth characteristics
vegetative growth disposition
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Edition, Academic Press Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles, Adelaide
Growth And Development Disposition
In Pinot Gris vines in Oregon, growth capacity restoration is completed by early February following accumulation of 1000 chilling hours, demonstrated by forcing tests showing visible bud swelling within 5 days and full budbreak within 10 days at 22°C, with capacity restoration confirmed through transcript analysis showing restored expression potential of CYCLIN D, EXPANSIN, and cell wall biosynthesis genes in response to growth-promoting conditions.
Growth capacity restoration in Vitis vinifera represents the re-establishment of the fundamental ability of meristematic cells to undergo division and expansion following chilling-dependent dormancy release, encompassing the restoration of cell cycle machinery, growth-related gene expression potential, and cellular structures required for developmental growth. This restoration occurs progressively during the final phases of chilling accumulation as dormancy-maintaining mechanisms are dismantled, reaching completion when all components required for growth have been restored to functional status. The cell cycle dimension of capacity restoration involves the re-establishment of division competence through restoration of cyclin expression potential enabling cell cycle progression, degradation of CDK inhibitor proteins that maintained G0/G1 arrest during dormancy, and chromatin derepression of cell cycle genes allowing their transcription upon appropriate signaling. The gene expression dimension encompasses the broader restoration of growth-related transcriptional programs, with derepression of genes encoding biosynthetic enzymes, structural proteins, and regulatory factors required for cell division and expansion, enabling rapid gene expression responses to growth-promoting signals. The structural dimension involves restoration of cellular infrastructure supporting growth, including re-organization of cytoskeleton for division and expansion, preparation of endomembrane systems for secretory activity supporting cell wall deposition, and maintenance of mitochondria and plastids in states ready for the increased energy and biosynthetic demands of growth. The hormonal signaling dimension addresses the restoration of growth hormone response capacity, with sensitization of receptor systems for cytokinins, gibberellins, and auxin that transduce growth-promoting signals into cellular responses. The verification of capacity restoration uses forcing tests demonstrating rapid growth response, with restored capacity indicated by short times to visible bud swelling and subsequent budbreak progression, and molecular markers confirming the restoration of growth gene expression potential through transcript analysis or chromatin accessibility assays. The completeness of capacity restoration determines the vigor of subsequent growth, with full restoration supporting rapid synchronous budbreak while partial restoration may limit growth potential.
Growth_Capacity_Restoration represents the fundamental cellular capacity aspect of Bud_Competence_Restoration. Distinguished from Metabolic_Reactivation_Potential which addresses metabolic preparation, while Growth_Capacity_Restoration focuses on cell cycle and growth machinery restoration. Encompasses cell cycle, gene expression, structural, and hormonal signaling dimensions. Verified through forcing response and molecular markers. Completeness determines subsequent growth vigor.
Entwicklungskapazitäts-Erholung
Meristem-Wachstumsfähigkeits-Wiedererrichtung
Wachstumskapazitäts-Wiederherstellung
Wachstumspotential-Restauration
developmental capacity recovery
growth competence recovery
growth potential restoration
meristem growth capacity re-establishment
Rohde A, Bhalerao RP (2007) Plant dormancy in the perennial context. Trends in Plant Science 12: 217-223 Cooke JE, Eriksson ME, Junttila O (2012) The dynamic nature of bud dormancy in trees: environmental control and molecular mechanisms. Plant Cell and Environment 35: 1707-1728 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235
Growth Capacity Restoration
In Pinot Noir vineyards of the Côte de Beaune, Burgundy, growth cessation progresses through October with shoot elongation terminating by mid-October when day length falls below 11 hours, cambial activity ceasing by late October as mean temperatures drop below 10°C, and complete growth cessation achieved by early November at BBCH stage 91 when all meristematic activity has terminated and periderm browning has advanced to within 5 nodes of shoot tips.
Growth cessation in Vitis vinifera represents the terminal phase of active vegetative development during which all meristematic growth activities progressively decline to complete inactivity, marking the functional transition from the growth period to the dormant period in the annual phenological cycle. This process occurs during autumn, typically from September through November in Northern Hemisphere viticulture, as the integrated effects of photoperiod response and temperature acclimation converge to terminate shoot elongation, arrest cambial activity, and reduce metabolic rates to winter maintenance levels. Growth cessation encompasses the coordinated shutdown of multiple growth centers including the shoot apical meristem which ceases producing new leaf and tendril primordia, the vascular cambium which terminates secondary xylem and phloem production, and the root system which reduces or ceases extension growth as soil temperatures decline. The process is regulated by the shifting hormonal balance characteristic of dormancy induction, with declining auxin and gibberellin levels removing growth promotion while increasing abscisic acid concentrations actively inhibit cell division and elongation in meristematic tissues. Morphological indicators of growth cessation include the termination of shoot elongation visible as static shoot tip position over successive observations, browning and hardening of shoot tips as they undergo lignification rather than continued extension, formation of terminal buds or abscission of immature shoot tips depending on cultivar and conditions, and the distinctive color change from green to brown as periderm development advances acropetally along shoots. The timing of growth cessation exhibits substantial variation among cultivars, with early-maturing varieties typically ceasing growth 2-4 weeks before late-maturing cultivars under equivalent environmental conditions, reflecting genetic differences in sensitivity to dormancy-inducing signals and the balance between vegetative vigor and dormancy tendency. Complete growth cessation is essential for proper cold hardiness development, as actively growing tissues are highly susceptible to freezing injury while dormant tissues with completed growth cessation achieve maximum cold tolerance. The relationship between harvest timing and growth cessation influences vine balance and subsequent season performance, with growth cessation before harvest allowing carbohydrate accumulation in reserves, while delayed cessation after harvest may compromise reserve status if photosynthetic capacity is reduced before growth terminates.
Growth_Cessation represents a major component process within Dormancy_Induction, specifically addressing the termination of all growth activities. Distinguished from Growth_Inhibition_Initiation which marks the beginning of growth decline, while Growth_Cessation represents complete termination. Subdivides into Shoot_Growth_Termination, Cambial_Activity_Decline, and Metabolic_Downregulation as component processes. Duration typically spans 4-6 weeks. Prerequisite for proper cold hardiness development and reserve accumulation. Corresponds to BBCH stages 91-93.
Vegetationsende
Wachstumsbeendigung
Wachstumseinstellung
Wachstumsstopp
end of growth
growth arrest
growth termination
vegetative growth cessation
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Mullins MG, Bouquet A, Williams LE (1992) Biology of the Grapevine Cambridge University Press
Growth Cessation
In Merlot vines in Bordeaux, growth competence restoration is completed by early February following accumulation of approximately 950 chilling hours, demonstrated by excised buds achieving visible bud swelling within 8 days and full budbreak within 12 days under forcing conditions at 22°C, with restored competence verified through detection of cell cycle gene transcripts including CYCLIN D and CDK expression potential in meristematic tissues.
Growth competence restoration in Vitis vinifera represents the re-establishment of the capacity for meristematic activity and developmental growth following the completion of chilling-dependent dormancy release processes, describing the physiological state in which all cellular and molecular prerequisites for growth have been restored after their suspension during endodormancy. This restoration is achieved through the cumulative effects of chilling exposure that progressively dismantle dormancy-maintaining mechanisms and re-establish growth-permissive conditions within bud tissues. The cellular basis of growth competence restoration involves the re-establishment of cell cycle competence in meristematic cells, with restoration of cyclin and CDK expression potential, degradation of CDK inhibitor proteins that blocked cell cycle progression during dormancy, and chromatin remodeling that makes cell cycle gene promoters accessible for transcription upon growth signal reception. The hormonal dimension of competence restoration involves the re-establishment of hormone signaling systems that transduce environmental cues into growth responses, including sensitization of cytokinin and gibberellin receptors, restoration of auxin polar transport capacity, and reduction of ABA signaling that inhibited growth during dormancy. The metabolic preparedness component of growth competence restoration encompasses the restoration of biosynthetic pathway capacity to support the rapid synthesis of proteins, nucleic acids, cell wall materials, and membranes required for cell division and expansion, with metabolic enzyme systems poised for rapid activation upon growth initiation. The symplastic connectivity restoration through callose degradation at plasmodesmata re-establishes the cell-to-cell communication required for coordinated growth responses across meristematic tissues. The verification of growth competence restoration relies on forcing tests demonstrating rapid and complete budbreak under optimal conditions, with fully competent buds achieving visible growth within 10-14 days at 22°C in contrast to the extended times characteristic of incompletely restored buds. The timing of competence restoration varies among buds on a vine, potentially creating heterogeneity that affects subsequent budbreak uniformity, with factors including bud position, light exposure history, and local microenvironment influencing the rate of restoration at individual bud positions.
Growth_Competence_Restoration represents the functional capacity aspect of Ecodormancy_Transition describing the re-establishment of growth potential. Distinguished from Dormancy_Release_Competence which describes the achieved state, while Growth_Competence_Restoration emphasizes the process of capacity recovery. Encompasses cell cycle, hormonal, metabolic, and connectivity restoration. Verified by forcing test response within 10-14 days. Heterogeneity among buds affects subsequent budbreak uniformity.
Entwicklungskompetenz-Restauration
Meristem-Kompetenz-Erholung
Wachstumspotential-Wiedererrichtung
developmental competence restoration
growth capacity restoration
growth potential re-establishment
Growth Competence Restoration
Syrah in Rhône Valley shows growth deceleration during late August-early September Year N+1 at BBCH 79-81, with berry growth rate declining from 38 mg per day early completion to 14 mg per day mid-deceleration to 4 mg per day entering lag phase representing 89% total reduction, water uptake declining from 0.4 to <0.1 g per berry per day, berry achieving 99% final growth phase size at 14.2-16.0 mm diameter and 12.8-17.4 g, under warm Mediterranean conditions of 24-30°C temperature, typical of this mid-to-late season cultivar completing deceleration
Growth deceleration represents the progressive growth rate reduction phase during enlargement phase completion when berry fresh weight gain declines from late enlargement 30-80 mg per day to transitional 10-30 mg per day to near-zero <5-10 mg per day entering lag phase, volumetric increase effectively ceases establishing growth quiescence, berry achieves final growth phase dimensions 98-100% mature size, and deceleration marks definitive transition from enlargement to lag phase minimal growth, occurring weeks 8-10 as growth phase concludes. This deceleration culmination phase occurs during late summer to early autumn in late August-early September in temperate regions Year N+1, spanning 1-2 weeks representing final deceleration, corresponding to BBCH 79-81 majority berries touching to beginning of ripening stages. Deceleration mechanism involves progressive growth rate reduction from declining cell expansion, water uptake declining to minimal levels approaching cessation, berry approaching final growth phase dimensions constraining further enlargement, metabolic transition reducing growth-supporting processes, developmental programming directing growth cessation, and integrated factors producing deceleration to quiescence. Growth rate progression shows decline from 30-80 mg per day late maximum enlargement to 15-40 mg per day early completion to <10 mg per day late completion entering lag phase, typical lag phase growth <5 mg per day representing essential quiescence, and rate trajectory documenting deceleration to near-zero. Growth rate quantification shows approximately 50-70% reduction from early to late completion phase, additional 60-80% reduction entering lag phase from completion levels, cumulative 85-95% total reduction from peak expansion to lag phase, and substantial deceleration establishing quiescence. Berry volumetric growth shows volumetric increase declining proportionally to fresh weight gain, berry volume approaching final growth phase dimensions, minimal volume increase <1-2% during deceleration, and effective volumetric cessation establishing size stability. Berry diameter dynamics shows diameter increase declining from 0.3-0.7 mm per week during late enlargement to <0.1-0.2 mm per week during deceleration, minimal diameter gain during final deceleration, and diameter stabilization at 98-100% final size. Berry fresh weight stabilization shows fresh weight achieving near-final growth phase values, typical ranges 2-6 g small, 5-13 g medium, 10-28 g large cultivars at deceleration completion, weight remaining relatively stable through lag phase with <3-5% change, and weight stability characteristic of quiescence. Water uptake decline shows water influx reducing from 0.3-0.8 g per berry per day during final expansion to <0.2 g per day during deceleration to <0.1 g per day entering lag, minimal water import reflecting growth cessation, and water uptake decline as deceleration mechanism. Water balance during deceleration shows reduced water import with minimal uptake, possible water loss through transpiration beginning to balance import, net water gain minimal <1-2% during deceleration, and approaching water balance characteristic of lag phase. Cell expansion contribution shows minimal cell enlargement <2% volume during deceleration, cell expansion effectively ceased contributing to deceleration, cells at final dimensions with stable size, and cellular quiescence establishing growth cessation. Berry composition during deceleration shows water content 78-82% relatively stable, organic acids declining 0.9-1.3%, sugars increasing 10-14%, pH rising 3.0-3.2, and composition transitioning toward lag phase characteristics. Organic acid decline shows malate continuing respiration declining from 0.8-1.0% to 0.6-0.9%, tartrate relatively stable 0.3-0.6%, total acids declining contributing to pH increase and softening, and acid dynamics as metabolic indicator. Sugar accumulation shows sugars increasing from 10-12% to 12-14% during deceleration, beginning sugar import preparing for ripening accumulation, and sugar increase as early ripening preparation. pH increase shows pH rising from 2.9-3.0 to 3.0-3.2 from acid decline, pH increase correlating with acid respiration, and pH shift toward lag phase values. Berry texture during deceleration shows slight softening from peak firmness, texture changes from acid decline and cell wall modifications, berry transitioning from hard to firm texture, and softening as deceleration indicator. Berry morphology shows berry at final growth phase size, green color maintained or beginning chlorophyll decline, surface smooth, and appearance as mature-sized green berry entering lag. Metabolic activity during deceleration shows respiratory rate at transitional low levels 1.2-2.0 μmol O₂ g⁻¹ h⁻¹ approaching lag phase 1.0-1.5 μmol O₂ g⁻¹ h⁻¹, metabolic activity declining toward maintenance levels, and metabolism characteristic of approaching quiescence. Developmental signals shows developmental programming directing deceleration and quiescence, possible hormonal changes regulating growth cessation, mature seeds signaling developmental readiness for progression, and integrated signals implementing deceleration. Environmental effects on deceleration include temperature affecting deceleration kinetics with warm conditions accelerating transition to lag quiescence, cool conditions extending deceleration period, water availability having minimal impact during deceleration as uptake minimal, and environmental factors having reduced influence as developmental programming dominates. Temperature effects show warm conditions promoting faster deceleration and earlier lag phase entry, cool conditions extending deceleration period, and temperature affecting timing of quiescence achievement. Deceleration completion shows growth rate declining to <5 mg per day establishing lag phase quiescence, berry dimensions stable at final growth phase size, metabolic and compositional markers indicating lag phase, and completion marking definitive enlargement phase end. Lag phase entry shows deceleration culminating in lag phase quiescence, characteristic low growth rate <5 mg per day, berry in quiescent state, and lag phase as distinct developmental period before veraison. Deceleration timing shows typical occurrence weeks 8-10 post-fertilization, cultivar variation with early cultivars entering lag earlier week 8-9 and late cultivars week 9-11, and timing determining total growth phase duration. Deceleration assessment shows serial growth measurements quantifying rate decline, growth curve analysis documenting deceleration trajectory, compositional analysis showing lag phase characteristics, and measurements enabling deceleration characterization. The process represents third and final aspect of enlargement phase completion establishing growth quiescence, following final cell expansion and pre-lag phase transition. Duration typically 1-2 weeks representing final deceleration weeks 8-10. Regional timing shows growth deceleration during late August-early September across climates entering lag phase. Viticultural significance includes representing definitive growth cessation establishing lag phase entry, marking completion of berry size establishment, demonstrating developmental transition to quiescence, and establishing stable berry state before ripening transformation. Research applications include analyzing deceleration mechanisms and regulation, investigating growth cessation dynamics, evaluating developmental signals directing quiescence, quantifying deceleration kinetics and lag phase entry, and assessing growth deceleration as determinant of lag phase and subsequent veraison timing.
Growth_Deceleration is third and final aspect of Enlargement_Phase_Completion, distinguished by progressive rate reduction to quiescence. Represents 85-95% cumulative reduction from peak to lag phase. Growth rate: 30-80 → 10-30 → <5 mg/day. Water uptake <0.1 g/berry/day. Berry 98-100% final growth phase size. Composition transitioning: acids declining, sugars 10-14%, pH 3.0-3.2. Duration 1-2 weeks. Follows Final_Cell_Expansion and Pre_Lag_Phase_Transition. Completes Enlargement_Phase_Completion and Berry_Cell_Enlargement_Phase. Establishes lag phase entry.
Dezeleration zur Ruhephase
Wachstums-Beendigungs-Annäherung
Wachstumsraten-Abnahme
deceleration to quiescence
final growth reduction
growth cessation approach
growth rate decline
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 453-458 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Growth Deceleration
Growth habit compatibility quality comparison in Carneros demonstrated contrasting cultivar responses to vertical shoot positioning requirements. Chardonnay exhibited excellent growth habit compatibility quality with its moderately upright natural habit, requiring only two positioning passes per season to achieve proper shoot placement within catch wires with 92% of shoots maintaining vertical orientation through harvest. Sauvignon Blanc showed intermediate growth habit compatibility quality, with semi-erect to spreading habit necessitating four positioning passes and achieving only 78% vertical shoot compliance with persistent tendency toward sprawling in the upper canopy. Muscat of Alexandria displayed poor growth habit compatibility quality for vertical systems, with strongly trailing growth habit requiring six positioning passes yet achieving only 61% vertical orientation as shoots continuously escaped catch wires and reverted toward horizontal growth, ultimately justifying conversion to lyre training where the cultivar's trailing habit became an asset rather than a liability, demonstrating how growth habit compatibility quality assessment can guide optimal training system selection.
Growth habit compatibility quality is a training system suitability quality that inheres in a grapevine cultivar or clone and represents the degree of alignment between the natural orientation, directional tendency, and architectural development pattern of growing shoots and the spatial configuration requirements of specific training systems that depend on particular shoot orientations to achieve functional canopy organization, effective light distribution, and operational compatibility with management and harvest operations. This quality addresses the fundamental geometric interaction between genetically determined shoot growth direction and the physical framework of trellis structures that must guide, contain, or accommodate shoot development to create productive and manageable canopy architectures. The ontological significance of growth habit compatibility quality lies in its role as a primary determinant of canopy management labor requirements and achievable canopy quality outcomes, wherein cultivars with growth habits naturally aligned with training system geometry require minimal intervention to achieve proper shoot placement, while mismatched combinations demand continuous corrective positioning or accept compromised canopy organization. The biological basis of shoot growth habit involves the gravitropic and phototropic responses governing shoot tip orientation, the mechanical properties of shoot tissues determining their flexibility and tendency to maintain versus change orientation, and the branching patterns affecting lateral development that may reinforce or counteract primary shoot direction. Upright growth habit, wherein shoots naturally orient vertically regardless of their point of origin, confers high compatibility with vertical shoot positioned systems but may create management challenges in sprawling or downward-trained configurations that require the shoot to grow against its natural tendency. Conversely, trailing growth habit with shoots tending toward horizontal or pendant orientation suits lyre, Geneva Double Curtain, and pergola systems but proves poorly compatible with vertical systems requiring substantial positioning labor to achieve and maintain upright shoot arrangement. The strength and consistency of growth habit expression varies among cultivars from strong deterministic patterns that resist repositioning to more plastic habits that readily adapt to imposed orientations, with this plasticity itself representing a compatibility characteristic favoring flexible cultivars for diverse training applications. Assessment of growth habit compatibility quality involves observation of shoot behavior under controlled conditions and practical evaluation of positioning requirements across training system trials.
Growth_Habit_Compatibility_Quality is_a Training_System_Suitability_Quality, representing the shoot orientation dimension of cultivar-training compatibility. This class serves as parent for Upright_Growth_Suitability_Quality, Trailing_Growth_Suitability_Quality, and Sprawling_Tendency_Quality which address specific habit types. Distinguished from Vigor_Quality by the focus on directional characteristics rather than growth intensity. The positioning labor requirement provides practical metric for compatibility assessment. Critical for matching cultivars to appropriate training systems.
Triebhaltungseignung
Wuchsformkompatibilität
Wuchshabitus-Erziehungs-Passung
Wuchsrichtungsverträglichkeit
architectural compatibility
growth direction suitability
habit-training match
shoot orientation compatibility
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 32-43 Reynolds AG et al. (2004) Influence of training system and vine spacing on vine performance of Vitis vinifera L. cv. Riesling. American Journal of Enology and Viticulture 55:79-86 Intrieri C, Poni S (1995) Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality of mechanized Italian vineyards. American Journal of Enology and Viticulture 46:116-127
Growth Habit Compatibility Quality
The growth habit description of Grenache documents vigorous, semi-erect to sprawling growth with long internodes, strong apical dominance producing dominant primary shoots with limited lateral branching, a tendency toward an open, spreading canopy architecture, and moderate to high fruitfulness from basal buds enabling spur pruning on gobelet (bush vine) or bilateral cordon training — this growth habit is ideally suited to the traditional Mediterranean bush vine training of southern Rhône, Spain, and southern Australia, where the spreading canopy provides self-shading in high-radiation environments and the vine's vigor is naturally constrained by summer drought stress on the shallow, stony soils of Châteauneuf-du-Pape, Priorat, and Barossa Valley. The vigorous, spreading Grenache growth habit contrasts with the compact, upright growth of Pinot Noir (short internodes, semi-erect shoot orientation, dense canopy requiring aggressive leaf removal) and the strongly trailing, pendulous growth of Syrah (shoot tips drooping under their own weight, requiring vertical shoot positioning to prevent canopy collapse).
An ampelographic description component that documents the vegetative growth characteristics of the grapevine cultivar — the overall vigor, shoot orientation and architecture, canopy density, and the spatial pattern of reproductive versus vegetative growth — that collectively define the cultivar-specific growth phenotype determining training system compatibility, canopy management requirements, yield potential, and the balance between vegetative and reproductive development that is the central management objective in quality-focused viticulture.
Growth_Habit_Description integrates the individual vegetative characters (internode length, shoot orientation, lateral branching, vigor) into a holistic characterization of the cultivar's growth phenotype — the emergent canopy architecture that results from the interaction of these individual traits with the specific training system and site conditions. The growth habit is the primary determinant of training system compatibility — erect cultivars with short internodes are suited to vertical shoot positioning (VSP) on bilateral cordons or cane-pruned systems, spreading cultivars with long internodes are suited to gobelet, pergola, or sprawl systems, and pendulous cultivars require trellising with catch wires to support the drooping shoots: the mismatch between cultivar growth habit and training system is one of the most common sources of canopy management problems in commercial viticulture, producing either excessively dense, shaded canopies (vigorous cultivars on restrictive training systems) or insufficiently filled trellis space (low-vigor cultivars on expansive training systems). The growth habit also determines the cultivar's response to site vigor — vigorous cultivars planted on fertile, well-watered sites with vigorous rootstocks produce excessive vegetative growth that shades the fruit zone, delays ripening, and promotes disease, while the same cultivars on low-vigor sites achieve the balanced growth that produces optimal fruit quality, explaining the viticultural principle that high-quality wine production generally requires matching high-vigor cultivars to low-vigor sites and vice versa.
Erziehungsform-Phänotyp
Wuchsform-Beschreibung
canopy architecture description
vegetative Wachstumscharakterisierung
vegetative growth characterization
vine growth phenotype
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Smart RE and Robinson M (1991) Sunlight into Wine — A Handbook for Winegrape Canopy Management, Winetitles Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Dry PR (2000) Canopy management for fruitfulness, Australian Journal of Grape and Wine Research 6: 109-115
Growth Habit Description
Comprehensive characterization of Growth_Habit_Disposition at INRAE Montpellier assessed canopy architecture across 60 cultivars grown on a standardized trellis allowing free shoot expression (wide T-trellis with catch wires but no forced positioning). Measurements included shoot angle from vertical at multiple positions (base, mid, tip), percentage of shoots achieving vertical positioning without intervention, natural canopy width and density distribution, and the proportion of shoot length requiring positioning to achieve VSP geometry. Growth_Habit_Disposition showed wide cultivar variation. Strongly upright cultivars like 'Cabernet Sauvignon' and 'Merlot' showed mean shoot angles of 12-18° from vertical, with 85-92% of shoots naturally achieving VSP-compatible orientation and minimal positioning requirement ( 50% of shoot length). The disposition was stable across years (r = 0.94 for cultivar rankings) and only weakly correlated with vigor (r = 0.28), confirming that Growth_Habit_Disposition represents an independent architectural characteristic. Canopy light interception modeling demonstrated that cultivars with Growth_Habit_Disposition matched to training system achieved 15-25% higher light interception efficiency than mismatched combinations.
A Growth_Habit_Disposition is a Growth_And_Development_Disposition that inheres in a Vitis vinifera plant and is realized through the characteristic spatial organization, architecture, and form of vegetative growth, determining how shoots are oriented in space, how the canopy develops its three-dimensional structure, and how the vine naturally occupies and fills its allotted growing space. This disposition represents the architectural program of the genotype, distinct from but interacting with vigor (growth intensity) to determine the overall canopy phenotype. Growth habit encompasses shoot orientation (the angle at which shoots grow relative to vertical), branching pattern (apical dominance versus lateral proliferation), internode characteristics affecting compactness, tendril development and climbing behavior, and the natural tendency toward particular canopy forms. Understanding Growth_Habit_Disposition is essential for training system selection, as cultivars with strongly upright growth are well-suited to vertical shoot positioning (VSP) systems, while trailing cultivars may perform better on sprawl or divided canopy systems that accommodate their natural growth direction. Growth habit also affects canopy management requirements—upright cultivars may need minimal shoot positioning while trailing types require intensive positioning or acceptance of non-vertical canopy architecture. The disposition is genetically determined and relatively stable across environments, though the degree to which inherent growth habit is expressed depends on vigor level (low-vigor vines may not fully express trailing tendency due to limited shoot length) and training system constraints.
Growth_Habit_Disposition is_a Growth_And_Development_Disposition representing spatial/architectural characteristics. Growth habit is distinct from vigor (intensity) though they interact in canopy expression. Primary child class is Shoot_Orientation_Disposition addressing the angle/direction of shoot growth. Growth habit determines training system suitability and canopy management requirements. The disposition is genetically stable but expression depends on vigor and training constraints. Matching growth habit to training system optimizes canopy light interception and management efficiency.
Rebarchitektur
Wuchsform-Disposition
Wuchshabitus
canopy form
growth form
vine architecture
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles, Adelaide Carbonneau A (1995) La surface foliaire exposée potentielle—guide pour sa mesure, Progrès Agricole et Viticole 112:204-212 Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review, American Journal of Enology and Viticulture 60:251-268
Growth Habit Disposition
In Syrah vines in the Rhône Valley, growth inhibition initiation becomes measurable in early October approximately 3 weeks after critical photoperiod exposure, with shoot elongation rate declining from 2-3 cm per day to less than 0.5 cm per day, plastochron extending from 3-4 days to more than 7 days, and apical meristem cell division rate decreasing by approximately 60% as determined by mitotic index measurements.
Growth inhibition initiation in Vitis vinifera represents the developmental phase during which photoperiod-derived signals achieve sufficient strength to begin actively suppressing meristematic activity, marking the functional transition from growing to pre-dormant physiological status. This phase occurs when the accumulated signals from short day perception and subsequent transduction pathways reach threshold levels that overcome the growth-promoting activities maintained during the growing season, initiating measurable reductions in cell division and elongation in shoot apical meristems and cambium. The earliest manifestations of growth inhibition include slowing of shoot elongation rate, reduced plastochron as new leaf initiation declines, decreased internode length in newly forming stem segments, and subtle changes in apical meristem geometry as cell proliferation slows. At the cellular level, growth inhibition involves declining cyclin and cyclin-dependent kinase activity that reduces cell cycle progression, altered auxin transport and signaling that affects cell elongation, and increasing expression of growth-inhibiting transcription factors that suppress genes required for meristematic function. The hormonal basis of growth inhibition centers on the shifting balance between growth promoters and inhibitors, with rising abscisic acid levels directly inhibiting cell cycle progression and cell elongation, declining gibberellin activity reducing its growth-promoting effects, and altered cytokinin import from roots affecting bud meristem activity. The initiation of growth inhibition exhibits spatial patterns within the vine, typically affecting apical meristems before lateral positions and young leaves before mature ones, reflecting differential sensitivity of tissues at different developmental stages. Growth inhibition initiation does not represent complete cessation but rather the beginning of progressive reduction, with full growth cessation achieved through continued signaling and reinforcement during subsequent dormancy establishment. The timing of growth inhibition initiation relative to environmental conditions affects both the adequacy of cold hardiness development and the completeness of reserve accumulation for the following season.
Growth_Inhibition_Initiation represents the functional outcome phase within Photoperiod_Response, marking the point where signaling produces measurable developmental effects. Distinguished from Photoperiod_Signal_Transduction which addresses upstream signaling, and from Growth_Cessation which represents complete growth stoppage. Represents beginning rather than completion of growth inhibition. Affects different meristems at different rates. Typically occurs 2-4 weeks after critical photoperiod exposure. Key indicators include declining elongation rate and extended plastochron.
Meristem-Inhibierung-Start
Wachstumseinstellung-Beginn
Wachstumshemmung-Initiierung
growth cessation onset
growth suppression initiation
initial growth arrest
initiale Wachstumsunterdrückung
meristem inhibition start
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Rohde A, Bhalerao RP (2007) Plant dormancy in the perennial context. Trends in Plant Science 12: 217-223
Growth Inhibition Initiation
In Gewürztraminer vines in Alsace, growth inhibition maximum is evident in mid-December when molecular analysis shows VvDAM1 and VvDAM2 expression at 8-10 fold higher levels than during active growth, CDK inhibitor ICK1 homolog expression elevated 5-fold, and complete absence of CYCLIN D expression in bud meristems, with these molecular markers of maximum inhibition coinciding with forcing test results showing zero budbreak after 30 days under optimal growth conditions.
Growth inhibition maximum in Vitis vinifera represents the peak expression of all cellular and molecular mechanisms that prevent meristematic activity during deep endodormancy, encompassing the convergent action of hormonal, transcriptional, and structural factors that collectively render buds completely unresponsive to growth-promoting signals. This condition occurs during early winter coincident with maximum dormancy depth, representing the state in which all growth-inhibiting systems operate at their highest intensity before chilling accumulation begins the progressive dismantling of inhibitory mechanisms. At the molecular level, growth inhibition maximum involves peak expression of growth-repressing transcription factors including dormancy-associated MADS-box proteins that directly suppress growth-related gene transcription, maximum activity of growth-inhibiting hormonal pathways particularly ABA signaling, and the most extensive chromatin modifications that establish repressive configurations at loci encoding cell cycle regulators, hormone biosynthesis enzymes, and other growth-essential genes. The cell cycle arrest during growth inhibition maximum is maintained by multiple mechanisms including CDK inhibitor protein accumulation, absence of cyclins required for cell cycle progression, and chromatin-based silencing of genes encoding cell cycle machinery components. The hormonal dimension of growth inhibition maximum involves not only peak ABA concentrations and signaling but also minimum levels of growth-promoting hormones including cytokinins, gibberellins, and auxin, creating the most growth-suppressive hormonal environment of the annual cycle. Plasmodesmatal modifications contribute to growth inhibition maximum by restricting symplastic communication and limiting the movement of signaling molecules that might otherwise coordinate growth responses, effectively isolating dormant meristems from potential growth-promoting signals. The structural barriers to growth during this phase include the presence of callose deposits at plasmodesmata and potentially within phloem elements that restrict both symplastic and long-distance transport. The release from growth inhibition maximum requires the coordinated reversal of multiple inhibitory mechanisms through chilling-dependent processes, explaining why dormancy release is gradual and requires extended cold exposure rather than occurring as an abrupt transition.
Growth_Inhibition_Maximum represents the functional dimension of Deep_Endodormancy describing peak inhibitory mechanism activity. Distinguished from Growth_Inhibition_Initiation which marks beginning of inhibition during dormancy induction. Encompasses transcriptional, hormonal, cell cycle, and structural inhibitory mechanisms. Characterized molecularly by peak DAM expression, CDK inhibitor accumulation, and growth gene silencing. Duration corresponds to deep endodormancy phase of 4-8 weeks. Release requires coordinated reversal through chilling-dependent processes.
Wachstumshemmungs-Maximum
complete growth inhibition
full meristem arrest
maximale Wachstumsunterdrückung
maximum growth arrest
peak growth suppression
voller Meristem-Arrest
vollständige Wachstumsinhibition
Rohde A, Bhalerao RP (2007) Plant dormancy in the perennial context. Trends in Plant Science 12: 217-223 Cooke JE, Eriksson ME, Junttila O (2012) The dynamic nature of bud dormancy in trees: environmental control and molecular mechanisms. Plant Cell and Environment 35: 1707-1728 Vergara R et al (2017) VvDAM1 gene expression regulation during sweet cherry bud dormancy. Tree Genetics and Genomes 13: 106
Growth Inhibition Maximum
Cabernet Sauvignon in Napa Valley shows growth mode transition during early August Year N+1 at BBCH 75-77, with division contribution ceasing completely week 5-6, expansion becoming 100% growth mechanism, berry growth rate maintaining 110-180 mg per day entirely from cell enlargement, water uptake accelerating to 1.2-1.8 g per berry per day, cell volume increase from 35 micrometers to progressive enlargement, under warm climate conditions of 26-32°C temperature, typical of this late-season cultivar establishing expansion exclusivity
Growth mode transition represents the fundamental growth mechanism shift during division phase completion when cell division ceases as growth contributor, cell expansion becomes exclusive growth mechanism, berry volumetric increase proceeds entirely through cell enlargement not proliferation, and developmental program establishes expansion as sole growth mode throughout subsequent expansion, lag, veraison, and ripening phases spanning 8-12 weeks until harvest. This critical mechanism shift occurs during late summer in early August in temperate regions Year N+1, typically establishing by weeks 5-6 as division completes, corresponding to BBCH 75 berry touch stage transitioning to BBCH 77 beginning bunch closure. Transition mechanism involves division contribution to growth ceasing completely with zero cell number increase, expansion contribution to growth becoming 100% of volumetric increase, growth rate maintained or increasing despite division loss through compensating expansion acceleration, developmental reprogramming establishing expansion gene expression, and transition establishing permanent growth mode for remainder of berry development. Division contribution cessation shows cell proliferation no longer contributing to berry growth, volumetric increase from division ceasing completely at transition, division phase having contributed cells now growing exclusively through enlargement, and zero ongoing division contribution post-transition. Expansion contribution establishment shows cell enlargement as sole growth mechanism after transition, 100% berry volumetric increase from water uptake and cell expansion, expansion phase growth rates 0.5-2.0 g fresh weight per day entirely from enlargement, and expansion exclusivity throughout subsequent 8-12 weeks. Growth rate dynamics during transition show berry growth rate maintained or increased despite division cessation, expansion acceleration compensating for division loss, typical fresh weight gain 50-150 mg per day during transition period, and growth continuity through mechanism shift. Compensatory expansion acceleration shows enlargement rate increasing as division declines, water uptake accelerating from 0.5 g to 1.0-2.0 g per berry per day, osmotic gradient intensification driving increased water influx, and expansion compensation maintaining growth. Berry growth pattern shows continuation of volumetric increase through transition, brief plateau possible during mode shift in some cases though often growth continuous, and overall growth trajectory relatively smooth through transition. Cell volume increase post-transition shows cells enlarging from 25-45 micrometer diameter at division cessation to progressive increase toward 80-200 micrometers at maturity, 5-20 fold total volume increase during expansion phase, and cell enlargement as exclusive growth mechanism. Water uptake as growth driver shows water import as primary expansion mechanism, xylem-mediated delivery during early expansion, osmotic gradient from solute accumulation driving influx, and water comprising 80-85% berry fresh weight. Vacuole expansion as enlargement mechanism shows central vacuole progressively enlarging occupying increasing cell volume fraction, eventual vacuole dominance at 85-95% cell volume in mature cells, and vacuole as major storage and osmotic compartment. Cell wall yielding enabling expansion shows wall modification through expansins, xyloglucan endotransglucosylase, pectin methylesterase, wall plasticity enabling turgor-driven expansion, and coordinated wall loosening with synthesis maintaining integrity. Turgor pressure driving expansion shows osmotic water uptake generating 0.5-1.2 MPa turgor, pressure force driving cell expansion against yielding walls, and sustained turgor through continued solute accumulation and water import. Developmental gene expression reprogramming shows downregulation of cell cycle and division genes, upregulation of expansion genes including expansins, aquaporins, transporters, metabolic enzyme shift toward expansion metabolism, and transcriptional program establishing expansion mode. Expansion gene upregulation shows expansin expression increasing enabling wall loosening, aquaporin water channel expression facilitating water transport, sugar and acid transporter expression supporting accumulation, and coordinated expression supporting expansion. Metabolic reprogramming shows shift from division-associated high nucleotide synthesis to expansion metabolism, carbohydrate import for osmotic gradient and wall synthesis, organic acid synthesis for osmotic potential, and metabolic reorganization supporting enlargement. Berry developmental progression shows transition enabling progression from division phase through expansion phase, sequential phase progression characteristic of berry development, expansion phase duration 3-5 weeks following transition, and eventual progression to lag phase as expansion slows. Growth phase I to II transition shows division phase representing growth phase I in double sigmoid pattern, transition period representing inflection between phases, expansion phase representing continued phase I growth, and eventual lag phase representing plateau between growth phases I and II. Post-transition berry characteristics show diameter 16-28 mm at transition depending on cultivar, fresh weight 5-12 g, established cell number 1-4 million cells, and berry composition remaining green acidic during expansion. Environmental effects on transition include temperature affecting transition timing through developmental rate with warm conditions accelerating transition, water availability critical for expansion phase post-transition with adequate water essential, and overall favorable conditions supporting smooth transition and subsequent expansion. Temperature independence of mode shows transition occurring across temperature ranges, mechanism shift robust to environmental variation, and developmental programming ensuring transition completion. Transition as developmental checkpoint shows transition representing committed developmental state change, checkpoint ensuring adequate preparation for expansion phase, and passage enabling berry progression through subsequent phases. The process represents third and final aspect of division phase completion emphasizing growth mechanism shift, following final mitotic events and cell division cessation. Duration represents transition establishment weeks 5-6. Regional timing shows growth mode transition during early August across climates as expansion becomes exclusive mechanism. Viticultural significance includes representing fundamental growth mechanism change determining subsequent berry development, establishing expansion as exclusive growth mode throughout remainder of development, demonstrating developmental robustness maintaining growth through mechanism shift, and marking critical transition from division-driven to expansion-driven berry growth. Research applications include analyzing transition mechanisms and regulation, investigating compensation maintaining growth rate, evaluating developmental reprogramming at transition, quantifying growth dynamics through mode shift, and assessing growth mode transition as critical developmental determinant.
Growth_Mode_Transition is third and final aspect of Division_Phase_Completion, distinguished by fundamental growth mechanism shift. Represents establishment of expansion as 100% exclusive growth mechanism. Division contribution ceasing, expansion compensating maintaining growth rate. Duration weeks 5-6. Expansion phase 8-12 weeks post-transition. Follows Final_Mitotic_Events and Cell_Division_Cessation. Completes Division_Phase_Completion and Berry_Cell_Division_Phase. Enables progression to Berry_Expansion_Phase. Critical developmental transition establishing expansion dominance.
Expansions-Exklusivitäts-Etablierung
Teilung-zu-Expansion-Wechsel
Wachstumsmechanismus-Änderung
Wachstumsmodus-Übergang
division to expansion shift
expansion exclusivity establishment
growth mechanism change
proliferation to enlargement transition
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 442-445 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Growth Mode Transition
The growth mutation data for Pinot Fin de Bourgogne documents a clonal variant of Pinot Noir characterized by shorter internodes (4-6 cm versus 6-9 cm in standard Pinot Noir), smaller leaves (blade area 90-120 cm² versus 130-170 cm²), smaller and more compact clusters (80-120 g versus 120-180 g), and smaller berries (0.9-1.1 g versus 1.2-1.5 g) that collectively produce the compact vine phenotype historically associated with the highest quality Pinot Noir production in the Côte d'Or — the genetic basis of the Pinot Fin phenotype has not been fully characterized at the molecular level, but whole-genome comparison with standard Pinot Noir clones (Vondras et al. 2019) identified candidate somatic mutations in genes affecting gibberellin signaling (potentially explaining the reduced internode length), auxin transport (potentially affecting apical dominance and branching), and cell cycle regulation (potentially reducing cell expansion rates) that are being evaluated as causal variants through functional studies.
A mutation information record specific to somatic mutations affecting vegetative growth characteristics in Vitis vinifera, documenting molecular changes in genes controlling shoot architecture, internode length, branching pattern, leaf morphology, growth vigor, maturity timing, cluster compactness, and other developmental traits that distinguish growth-habit variants among clones of a single cultivar, typically identified through clonal selection programs as phenotypically divergent individuals within commercial vineyards.
Growth_Mutation_Data documents the least molecularly characterized but most commercially exploited category of clonal variation in grapevine. Clonal selection programs have traditionally focused on growth and yield characteristics — selecting clones with optimal vigor, cluster architecture, yield potential, and disease tolerance for specific production contexts — long before the molecular basis of clonal differences was understood: the result is a rich collection of phenotypically defined clonal variants (e.g., the ENTAV-INRA French clonal collection of over 1,300 certified clones across 250+ cultivars) whose growth differences are empirically documented through multi-year field trials but whose underlying mutations are largely uncharacterized at the DNA level. The molecular characterization of growth mutations is inherently more challenging than color or flavor mutations because vegetative growth traits are polygenic (controlled by many genes each with small effect), strongly influenced by environment (masking genetic differences in variable field conditions), and difficult to phenotype precisely (growth measurements have high within-vine and between-vine variability): consequently, while whole-genome resequencing can identify hundreds of candidate somatic mutations distinguishing clones, linking specific mutations to specific growth phenotypes requires extensive functional validation through gene editing, expression analysis, and replicated field trials that remain in early stages for most growth-related clonal variants.
Entwicklungsvarianten-Dokumentation
Wuchsmutations-Daten
developmental variant data
growth habit mutation record
morphological mutation documentation
morphologische Mutations-Daten
Vondras AM et al. (2019) The genomic diversification of grapevine clones, BMC Genomics 20: 972 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340 ENTAV-INRA (2007) Catalogue des variétés et clones de vigne cultivés en France, 2nd Ed, Institut Français de la Vigne et du Vin Torregrosa L et al. (2019) The microvine, a biological model, very versatile and efficient to boost grapevine research, OENO One 53: 373-391
Growth Mutation Data
Merlot in Bordeaux shows growth rate decline start during late August-early September Year N+1 at BBCH 81, with berry growth rate declining from 14 mg per day to 4 mg per day over 4-6 days, water uptake reducing from 0.2 to <0.08 g per berry per day, berry dimensions stable at 12.2-12.6 mm diameter and 8.4-9.2 g fresh weight, volumetric growth <0.8% during period, under oceanic conditions of 20-26°C temperature, typical of this mid-season cultivar initiating lag growth cessation
Growth rate decline start represents the growth deceleration initiation phase during lag phase initiation when berry fresh weight gain begins final reduction from enlargement completion levels 10-30 mg per day toward lag phase minimal rates <5 mg per day, volumetric growth effectively ceases, berry enters dimensional stability characteristic of quiescence, and growth decline establishes lag phase defining minimal growth rate, occurring over days at lag initiation. This growth cessation onset phase occurs during late summer to early autumn in late August-early September in temperate regions Year N+1, proceeding over several days representing decline initiation, corresponding to BBCH 81 beginning of ripening stage early transition. Decline start mechanism involves final growth rate reduction continuing from enlargement completion deceleration, cellular processes supporting growth declining to minimal activity, water uptake reducing to near-equilibrium levels, berry approaching stable mass and volume, developmental programming directing final growth cessation, and integrated changes initiating lag minimal growth. Growth rate at decline start shows typical rates 10-30 mg fresh weight per day at initiation onset declining during initiation, progressive deceleration toward lag levels, eventual stabilization at <5 mg per day by decline completion, and rate trajectory establishing quiescence. Growth rate reduction kinetics show approximately 50-70% reduction during decline start phase from onset to early lag levels, additional reduction to final lag minimal rates, cumulative 70-85% total reduction from late completion to established lag, and substantial deceleration initiating quiescence. Berry fresh weight dynamics shows minimal weight gain <10-20 mg total during decline start period, weight stabilization approaching lag equilibrium, eventual stable weight within 2-5% variation characteristic of lag, and weight constancy as lag feature. Berry volumetric growth shows volumetric increase declining to minimal levels, volume change <0.5-1% during decline start, effective volumetric cessation establishing dimensional stability, and volume constancy defining lag phase. Berry diameter stability shows diameter remaining essentially constant during decline start and lag phase, variation <1-2% from minor fluctuations and measurement variability, stable dimensions characteristic of quiescent state, and diameter constancy contrasting with enlargement growth. Water uptake during decline start shows water influx declining from completion minimal 0.1-0.3 g per berry per day to lag very low <0.05-0.1 g per day, water balance approaching equilibrium with uptake approximately balancing transpiration loss, net water flux minimal establishing stable berry hydration, and minimal uptake characteristic of growth cessation. Water balance characteristics shows uptake-transpiration balance approaching equilibrium during decline start, possible slight net water loss in some conditions creating minor weight decrease, overall water balance stability, and equilibrium as quiescence feature. Cell expansion cessation shows cell enlargement effectively ceased during decline start, cells at final mature dimensions, no volumetric growth at cellular level, and cellular stability establishing berry dimensional constancy. Cell volume stability shows cells maintaining 98-100% final dimensions achieved during enlargement, minor volume fluctuations <1-2% from osmotic changes, overall cell size constancy, and stability persisting through lag and into ripening. Turgor maintenance shows turgor pressure sustained 0.4-0.8 MPa during decline start and lag, turgor adequate for cell integrity, minimal expansion despite sustained pressure from non-yielding walls, and turgor-stability relationship characteristic of quiescent cells. Wall properties during decline start shows walls non-yielding with high yield threshold 0.8-1.2 MPa, wall plasticity minimal preventing expansion, wall mechanical properties growth-restrictive, and walls as physical constraint maintaining dimensional stability. Osmotic potential during decline start shows values -1.2 to -1.6 MPa maintained relatively stable, minimal osmotic potential changes from slow compositional shifts, osmotic stability characteristic of quiescence, and potential adequate for turgor maintenance. Berry composition during decline start shows water 78-82% stable, organic acids 0.8-1.2% declining slowly, sugars 11-15% increasing slowly, pH 3.0-3.2 rising gradually, and composition showing slow changes characteristic of lag initiation. Metabolic activity during decline start shows respiratory rate at transitional low levels 1.0-1.5 μmol O₂ g⁻¹ h⁻¹, metabolic activity declining to lag maintenance levels, minimal biosynthetic activity, and metabolism supporting maintenance not growth. Growth-supporting processes decline shows biosynthetic pathways for structural components at minimal activity, protein synthesis reduced to maintenance levels, cell wall synthesis ceased or minimal, and growth processes effectively silenced. Environmental effects on decline start include temperature affecting decline kinetics with warm conditions accelerating growth cessation, cool conditions potentially extending decline period, water availability affecting final berry hydration state, and environmental factors influencing transition dynamics. Temperature effects show warm conditions promoting faster decline and earlier lag establishment, cool conditions extending decline period, and temperature affecting growth cessation timing. Decline start assessment shows serial growth measurements quantifying rate reduction, volumetric analysis documenting growth cessation, water flux measurements showing uptake decline, and measurements enabling decline characterization. The process represents first aspect of lag phase initiation establishing minimal growth, preceding metabolic transition onset. Duration typically several days representing decline initiation period. Regional timing shows growth rate decline start during late August-early September across climates at lag initiation. Viticultural significance includes representing final growth cessation establishing lag quiescence, determining berry dimensional stability, demonstrating developmental control of growth termination, and establishing minimal growth as lag defining characteristic. Research applications include analyzing growth cessation mechanisms and regulation, investigating cellular and metabolic bases of quiescence, evaluating water relations during growth termination, quantifying decline kinetics and final growth rates, and assessing growth cessation as determinant of lag phase characteristics.
Growth_Rate_Decline_Start is first aspect of Lag_Phase_Initiation, distinguished by final growth deceleration. Represents 70-85% total reduction from 10-30 mg/day to <5 mg/day. Water uptake declining to <0.05-0.1 g/berry/day approaching equilibrium. Volumetric growth <0.5-1%. Cell expansion ceased, walls non-yielding (threshold 0.8-1.2 MPa). Berry dimensions stable within 1-2%. Duration several days. Precedes Metabolic_Transition_Onset. Establishes lag minimal growth defining characteristic.
Dezeleration-Initiierung
Wachstums-Beendigungs-Beginn
Wachstumsraten-Abnahme-Start
deceleration initiation
final growth decline
finale Wachstums-Abnahme
quiescence growth start
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 455-458 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Growth Rate Decline Start
Growth reduction quality assessment for nitrogen deficiency in Burgundy Pinot Noir compared vegetative development across nitrogen management zones to establish the optimal nitrogen status balancing vine health with fruit quality objectives. Shoot growth measurements at véraison quantified growth reduction quality, with severely nitrogen-deficient vines showing mean shoot length of 55 cm and 12 nodes versus 95 cm and 18 nodes in adequate-nitrogen vines, representing 42% length reduction and 33% node number reduction. Leaf area measurements revealed growth reduction quality in individual organ size, with mature leaf blade area averaging 85 cm² in deficient vines versus 135 cm² in adequate vines, combined with the reduced leaf number to produce total shoot leaf area of 850 cm² versus 2,100 cm² representing 60% reduction in photosynthetic capacity. Internode length provided the most sensitive growth reduction quality indicator, with mean internode of 4.2 cm in deficient versus 6.8 cm in adequate vines, a parameter easily assessed in the field without destructive sampling. Shoot diameter measurements showed growth reduction quality with cane caliper of 5.2 mm versus 7.8 mm at the fifth internode, with thin canes potentially affecting bud fruitfulness for the following season. The growth reduction quality assessment established that moderate nitrogen limitation (shoot length 75-85 cm) optimized the balance between vine health and Pinot Noir fruit quality, while severe deficiency (less than 60 cm) compromised both current yield through inadequate photosynthetic capacity and future productivity through reduced bud fruitfulness and carbohydrate reserves.
Growth reduction quality in nitrogen deficiency is a nitrogen deficiency symptom quality that inheres in a grapevine organism and represents the diminished vegetative growth resulting from inadequate nitrogen availability for protein synthesis and cell division, manifesting as shortened internodes, reduced shoot length, smaller leaf size, thinner shoots, and overall decreased vine vigor that limits canopy development and photosynthetic capacity with downstream effects on fruit production and carbohydrate reserve accumulation. This quality captures the growth-limiting dimension of nitrogen deficiency, recognizing that as a component of all proteins including the enzymes catalyzing growth processes and the structural proteins of new tissues, nitrogen deficiency fundamentally constrains the rate and extent of vegetative development. The ontological significance of growth reduction quality in nitrogen deficiency lies in its dual nature as both a deficiency symptom indicating nutritional stress and a potential viticultural objective where controlled nitrogen limitation is used to manage vigor and optimize fruit quality, creating the need for diagnostic assessment that distinguishes problematic deficiency from intentional vigor management. The biological basis of growth-reduction under nitrogen deficiency encompasses the dependence of cell division on nitrogen supply for DNA and histone synthesis, the requirement for amino acids and proteins in all aspects of cell expansion and differentiation, and the competition among organs for limited nitrogen that prioritizes developing tissues while constraining overall growth rate. The quantitative relationship between nitrogen status and growth rate shows characteristic response curves, with growth increasing with nitrogen supply up to sufficiency levels then plateauing or even declining at excess nitrogen that promotes excessive vegetative growth, creating the optimal nitrogen range that balances productivity with fruit quality. The morphological manifestations of nitrogen-limited growth include not only reduced dimensions of individual organs but altered architecture with reduced lateral branching, decreased tendril development, and modified root-shoot allocation that may enhance root exploration of soil nitrogen but cannot fully compensate for inadequate supply.
Growth_Reduction_Quality_(N) is_a Nitrogen_Deficiency_Symptom_Quality, representing the diminished vegetative development resulting from nitrogen-limited protein synthesis and cell division. This class addresses the vigor dimension of nitrogen deficiency. Shoot length, internode length, leaf size, and shoot diameter provide quantitative assessment. The dual nature as deficiency symptom versus vigor management objective creates diagnostic complexity. Optimal nitrogen range balances productivity with fruit quality (moderate status often desirable for quality). Severe deficiency compromises both current and future productivity.
N deficiency vigor reduction
N-Mangel-Kümmerwuchs
Stickstoff-limitiertes Wachstum
Wachstumsreduktion bei N-Mangel
nitrogen-limited growth
reduced vegetative development
reduzierte vegetative Entwicklung
stunted growth from N deficiency
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Treeby MT et al. (2000) Irrigation management and rootstock effects on Shiraz yield and must composition. Australian Journal of Grape and Wine Research 6:206-214 Bell SJ, Henschke PA (2005) Implications of nitrogen nutrition for grapes, fermentation and wine. Australian Journal of Grape and Wine Research 11:242-295
Growth Reduction Quality (N)
Growth respiration assessment during rapid shoot elongation in Cabernet Sauvignon quantifies the biosynthetic cost of new tissue production, with shoot tips growing at 2.5 g dry weight/day showing growth respiration of 0.85 g CO₂/day (growth respiration coefficient of 0.34 g CO₂/g new biomass), requiring total carbon allocation of approximately 3.4 g glucose/day to support both the structural increment and associated biosynthetic costs, demonstrating how growth respiration substantially increases the photosynthate demand of growing tissues beyond the carbon appearing in new structure.
Growth respiration quality is a respiration rate quality that represents the rate of respiratory metabolism associated with the biosynthesis of new structural and storage compounds, characterizing the energetic and carbon skeleton costs of converting photosynthate into the proteins, lipids, cell walls, and other compounds that constitute new tissue growth. Within the BFO framework, this quality specifically addresses the growth-associated component of total respiration, distinguishing the biosynthetic costs of creating new biomass from the maintenance costs of sustaining existing tissue while recognizing that growth respiration scales with growth rate and represents a fixed cost per unit new biomass synthesized that determines the carbon conversion efficiency of growth processes. The ontological significance lies in growth respiration determining the efficiency of converting photosynthate into new vine tissue, where the growth respiration coefficient (g CO₂ released per g new biomass) establishes the carbon cost of growth that reduces the fraction of photosynthate appearing as structural increment, with growth respiration typically consuming 25-35% of the carbon allocated to growth processes. The biochemical basis for growth respiration includes the ATP costs of biosynthesis (polymerization of monomers into macromolecules), the reducing power requirements of anabolic reactions (NADPH for fatty acid and amino acid synthesis), the transport costs of moving substrates to growth sites, and the carbon skeleton requirements obtained from respiratory pathway intermediates (organic acids from TCA cycle for amino acid synthesis). The composition-dependent nature of growth respiration reflects the varying biosynthetic costs of different compounds, with protein synthesis having highest respiratory cost (approximately 0.4 g CO₂/g protein), lipid synthesis intermediate (0.3 g CO₂/g lipid), and carbohydrate polymerization lowest (0.1 g CO₂/g cell wall), such that tissue composition determines the overall growth respiration coefficient. The growth rate relationship with total respiration shows growth respiration dominating in rapidly growing tissues (young leaves, shoot tips, developing berries) while maintenance dominates in mature tissues, with the shift from growth to maintenance respiration marking the transition from expanding to steady-state tissue function. Understanding growth respiration quality enables prediction of the carbon cost of growth, efficiency assessment of biomass production, and appreciation of how tissue composition affects the respiratory cost of growth across vine organs.
Growth_Respiration_Quality addresses the biosynthetic component of respiration associated with new tissue production. This quality is proportional to growth rate and depends on the composition of new tissue being synthesized. The class supports growth efficiency assessment and carbon allocation modeling through characterization of the respiratory cost of biomass production.
Konstruktionskosten
Rg-Wert
Wachstumsatmung
anabolic respiration
biosynthetic respiration
biosynthetische Respiration
construction cost
Penning de Vries FWT et al. (1974) Products, requirements and efficiency of biosynthesis a quantitative approach. Journal of Theoretical Biology 45(2): 339-377 Thornley JHM, Cannell MGR (2000) Modelling the components of plant respiration: representation and realism. Annals of Botany 85(1): 55-67
Growth Respiration Quality
Detailed tracking of Growth_Resumption_Disposition at Cornell University monitored shoot elongation, leaf emergence, and lateral shoot development following relief of a 21-day drought stress across 18 cultivars. Daily measurements of primary shoot length, leaf number, and lateral bud activation were recorded for 28 days post-rewatering, enabling characterization of growth resumption kinetics. Growth_Resumption_Disposition varied substantially among cultivars. 'Cayuga White' (cold-hardy hybrid) showed rapid resumption with shoot elongation detectable within 4 days of rewatering, reaching elongation rate of 2.8 cm day⁻¹ by day 10 (exceeding pre-stress rate of 2.4 cm day⁻¹), and new leaf emergence beginning day 6. 'Cabernet Franc' displayed delayed Growth_Resumption_Disposition with elongation not detectable until day 9, reaching only 1.5 cm day⁻¹ by day 14 (60% of pre-stress), and new leaf emergence delayed until day 12. Analysis of growth components revealed that resumption timing correlated strongly with pre-stress carbohydrate reserves in shoot and root tissues (r = 0.72), as stressed vines with depleted starch showed delayed regrowth. Hormonal analysis showed that rapid-resuming 'Cayuga White' achieved pre-stress cytokinin/ABA ratios by day 5, while slow-resuming 'Cabernet Franc' maintained elevated ABA until day 12. Meristem viability assessment revealed that severe-stress treatments (predawn -1.2 MPa) killed 15% of 'Cabernet Franc' shoot apical meristems versus 0% in 'Cayuga White', indicating that Growth_Resumption_Disposition involves both meristem stress tolerance and metabolic recovery capacity.
A Growth_Resumption_Disposition is a Drought_Recovery_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to rapidly reinitiate shoot elongation, leaf expansion, and developmental processes following relief of water stress, restoring the growth trajectory interrupted by drought. This disposition represents the developmental component of drought recovery, determining how quickly productive growth resumes after the immediate physiological crisis of dehydration is resolved. Growth resumption requires not only restored water status but also reactivation of meristematic activity, remobilization of carbohydrate reserves, resumption of cell division and expansion, and restoration of hormonal balances favoring growth over stress responses. The timing and vigor of growth resumption depend on the extent of meristem damage during stress (severe dehydration can injure or kill shoot apical and lateral meristems), carbohydrate reserve status (stressed vines may have depleted reserves limiting growth fuel), and the speed of transitioning from stress-protective to growth-promoting hormonal states (particularly declining ABA and increasing cytokinins and gibberellins). Growth_Resumption_Disposition is economically important because delayed regrowth reduces canopy development, shortens the effective growing season, and may compromise fruit development and ripening, particularly when drought occurs during critical phenological stages.
Growth_Resumption_Disposition is_a Drought_Recovery_Disposition. Growth resumption requires restoration of meristematic activity, carbohydrate supply, and growth-promoting hormonal balance. Resumption timing reflects both stress damage extent and recovery capacity. Carbohydrate reserves and meristem viability are key determinants. Delayed growth resumption reduces seasonal productivity and may compromise fruit development. Growth resumption can be measured by shoot elongation rate, leaf emergence, and lateral activation.
Entwicklungs-Wiederherstellung
Post-Stress-Wachstumsstart
Wachstumswiederaufnahme-Disposition
developmental recovery
post-drought growth restart
Flexas J et al. (2006) Drought-inhibition of photosynthesis in C3 plants: stomatal and non-stomatal limitations revisited, Annals of Botany 98:871-892 Chaves MM et al. (2003) Understanding plant responses to drought—from genes to the whole plant, Functional Plant Biology 30:239-264 Patakas A et al. (2002) Leaf age effects on solute accumulation in water-stressed grapevines, Journal of Plant Physiology 159:969-976
Growth Resumption Disposition
The growth type description of Syrah documents a distinctly trailing, pendulous growth habit — primary shoots grow initially semi-erect for 40-60 cm then progressively curve downward as the shoot tip mass exceeds the structural capacity of the internode tissue to support it, with the shoot apex eventually hanging vertically if not supported by trellis catch wires. This pendulous tendency is one of Syrah's most recognized viticultural characteristics, requiring vertical shoot positioning (VSP) with 2-3 pairs of movable catch wires that are progressively raised during the growing season to redirect the drooping shoots into an upright canopy wall — without this support, Syrah shoots collapse into a dense, tangled mass that shades the fruit zone, promotes disease, and prevents mechanized canopy management. The trailing Syrah growth type contrasts with the erect growth of Cabernet Sauvignon (shoots maintaining upright orientation with minimal catch wire support, growing between fixed foliage wires with limited manual positioning needed), the semi-erect growth of Merlot (shoots moderately arching but maintaining general upward orientation), and the spreading growth of Grenache (shoots growing laterally in all directions from the vine head, suited to the unsupported bush vine form).
A growth habit description component that records the characteristic shoot orientation, branching pattern, and canopy architecture type of the cultivar — classifying the growth as erect (shoots growing vertically with minimal deviation), semi-erect (shoots initially vertical then arching outward), horizontal/spreading (shoots growing laterally from the vine head or cordon), trailing/pendulous (shoot tips drooping under gravitational influence), or prostrate — with growth type determining the natural canopy form that the training system must accommodate or redirect for optimal fruit zone exposure and vineyard management access.
Growth_Type_Description captures the spatial orientation of vegetative growth that, together with vigor, defines the cultivar's canopy architecture potential and training system requirements. The shoot orientation is determined by the interplay of gravitropism (the tendency of shoots to grow away from gravity), phototropism (the tendency to grow toward light), the structural rigidity of the shoot (determined by internode diameter, cell wall thickness, and lignification rate), and the weight of the growing shoot tip (determined by internode length, leaf size, and growth rate): erect cultivars have rigid shoots with strong negative gravitropic response that maintains vertical growth even as shoot weight increases, while trailing cultivars have flexible shoots whose structural capacity is exceeded by the increasing tip weight as the shoot elongates beyond 40-60 cm, producing the progressive downward curvature characteristic of this growth type. The training system implications of growth type are direct and constraining — erect cultivars are suited to minimal trellising (gobelet, vertical cordons, free-standing vines), trailing cultivars require structured trellising with catch wires for shoot support (VSP, Scott Henry, Smart-Dyson), and spreading cultivars are suited to horizontal or overhead training (pergola, tendone, GDC) that accommodates the lateral growth orientation. The choice of training system is therefore not arbitrary but must respect the cultivar's natural growth type to minimize the canopy management labor required to maintain optimal canopy architecture.
Triebrichtungs-Charakterisierung
Wuchs-Architekturtyp
Wuchstyp-Beschreibung
canopy form classification
growth architecture type
shoot orientation record
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Smart RE and Robinson M (1991) Sunlight into Wine, Winetitles Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia
Growth Type Description
Cabernet Sauvignon growth year annual ring from 2015 growing season shows 1.4-2.2 mm width (narrow ring indicating severe California drought documented that year with <40% normal precipitation), can be precisely dated by counting from outermost 2025 ring inward (10 rings = 2015), shows characteristic narrow width matching 2015 rings across multiple vineyard sites confirming synchronous drought stress response, and correlates with reduced yields and concentrated fruit composition documented in 2015 vintage records
A growth year annual ring is an annual ring explicitly associated with a specific calendar year of formation, enabling precise dating of wood tissues, correlation of ring characteristics with known environmental conditions or management practices, and temporal reconstruction of vine growth history through dendrochronological analysis. Growth year annual ring identification requires= ring counting from known reference (outermost ring under bark representing most recent growing season with known year, counting inward to date all rings), establishment date determination (planted year for own-rooted vines, grafting year for grafted vines providing starting point for ring accumulation), cross-dating validation (matching ring width patterns across multiple vines confirming year assignments), and accounting for potential anomalies (missing rings in severe stress years, false rings from mid-season events requiring careful interpretation). Growth year annual ring dating methodology includes= outermost complete ring representing previous dormant season (ring under bark in March 2026 represents 2025 growing season April-September 2025), sequential inward counting (each ring inward representing one year earlier= 2024, 2023, 2022, etc.), correlation with planting or grafting date (pith or innermost ring corresponding to establishment year), and verification through cross-dating (comparing patterns across vines and sites confirming date assignments especially in older sections where counting errors accumulate). Growth year annual ring width patterns correlated with known conditions= year-specific environmental records (matching narrow rings with documented drought years, wide rings with wet years), historical management events (ring width changes following irrigation installation, rootstock change, or training system modification), disease outbreak timing (narrow rings corresponding to documented Eutypa, Esca, or phylloxera impacts), and vintage quality correlation (ring characteristics in grape years relating to wine quality records). Growth year annual ring examples include= 2012 ring showing reduced width (narrow 1.2-2.5 mm) in California vines reflecting documented severe drought conditions, 2003 ring showing narrow width (0.8-1.8 mm) in European vines reflecting historic heat wave and drought, 1976 ring showing extreme narrowness (0.4-1.2 mm) marking famous European drought year, young vine wide rings (5-8 mm) in establishment years 2020-2022 reflecting pre-cropping vigor, and progressive narrowing in rings 2015-2025 indicating old vine senescence in 40-50 year vines. Growth year annual ring enabling historical reconstruction= climate history (ring width series providing precipitation and temperature proxies for decades or centuries), management history (ring responses revealing irrigation changes, pruning modifications, crop load adjustments), disease history (ring patterns indicating disease onset, progression, and potential treatment or recovery), and vigor trends (ring width trajectories showing youth vigor, mature stability, or senescent decline phases). Growth year annual ring cross-dating validation= matching ring width patterns across vines (synchronous narrow rings in stress years confirming dates), comparing with master chronologies (reference ring width sequences from long-lived trees or well-dated vine collections), statistical correlation (quantifying pattern matching between samples using dendrochronological techniques), and verification with known events (confirming narrow rings correspond to documented drought years, wide rings follow crop failures). Growth year annual ring applications in viticulture research= climate-growth relationships (quantifying temperature, precipitation, or drought impacts on vine growth across decades), rootstock performance evaluation (comparing ring width patterns between scion-rootstock combinations on identical sites and years), disease impact assessment (documenting growth reduction following disease infection or progression), management practice effects (evaluating irrigation, nutrition, or canopy management impacts on trunk growth), and old vine age determination (establishing precise ages for heritage vines through ring counting and cross-dating). Growth year annual ring limitations and uncertainties= missing rings (severe stress years potentially producing no detectable xylem requiring careful analysis and cross-dating), false rings (mid-season stress events creating intra-annual boundaries resembling true ring boundaries), eccentric growth (non-centered pith and variable ring widths on different radii requiring multiple measurements), pith absence (oldest wood decayed or sample missing center requiring minimum age estimation), and cross-dating requirement (single vine ring counts requiring validation through comparison with other vines or chronologies). Growth year annual ring data representation= ring width series (sequence of measurements for each year typically in 0.01 mm precision), standardized indices (removing age-related trends to emphasize year-to-year climatic variation), chronologies (averaged series across multiple vines producing robust site or regional records), and correlation with environmental variables (statistical relationships between ring widths and temperature, precipitation, drought indices).
Growth_Year_Annual_Ring emphasizes temporal dating and is distinguished from Annual_Ring (any yearly increment), Narrow_Annual_Ring (width emphasis), and Wide_Annual_Ring (favorable growth) by explicit calendar year assignment. Critical for dendrochronological dating, correlating ring characteristics with known historical events, climate reconstruction, management impact assessment, and precise aging of old vines through ring counting and cross-dating validation.
Calendar year ring
Chronological ring
Chronologischer Wachstumsring
Dated ring
Datierter Jahresring
Kalenderjahr-Ring
Schweingruber, F. H. (1996). Tree Rings and Environment= Dendroecology. Paul Haupt Publishers. | Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press.
Growth Year Annual Ring
Cabernet Sauvignon grafted on SO4 rootstock grafted in spring 2008 showing growth year rootstock annual ring for year 2019 (ring 12 in trunk sequence counting from pith) measuring 0.32 cm width, ring year 2019 showing moderate width intermediate between narrow ring year 2018 (0.22 cm from documented drought) and wider ring year 2020 (0.41 cm from optimal irrigation), ring 2019 characteristics correlated with weather records showing normal precipitation and moderate temperatures during that growing season, ring year designation enabling precise temporal positioning within 12-year trunk development history, increment core extracted 2020 allowing designation of all rings from ring 1 (2009 partial season after spring grafting) through ring 12 (2019 most recent complete season) to ring 13 (2020 current forming season incomplete), ring year sequence revealing complete chronological growth record from establishment through current year
Growth year rootstock annual ring is rootstock annual ring identified by its formation year and associated with specific calendar year or years post-grafting, emphasizing the temporal designation and chronological position of ring within trunk developmental sequence, enabling precise dating of ring formation for dendrochronological analysis, correlation with known environmental events or management practices during formation year, documentation of trunk age and growth history, and interpretation of ring characteristics in context of specific year's conditions, with ring potentially designated by calendar year (e.g. ring 2018) or relative year from grafting (e.g. ring year 12 meaning 12th year post-grafting) depending on analysis purpose and available information. Growth year rootstock annual ring temporal designation includes= identified by specific formation year (each ring associated with one calendar year or one year post-grafting designating when ring produced), enabling precise chronological positioning (ring placement in exact temporal sequence within trunk developmental history), allowing historical correlation (ring linked to known events, conditions, or management practices during formation year), supporting dendrochronological dating (ring year designation enabling cross-dating across samples and correlation with environmental records), and facilitating age determination (counting rings from pith outward with year designation revealing trunk age and annual growth sequence). Growth year rootstock annual ring designation methods includes= using calendar year (ring designated by formation year e.g. "ring 2015" or "ring 1998-1999" linking ring to specific calendar date), employing years post-grafting (ring designated by position in trunk sequence e.g. "ring 5" meaning 5th year after grafting regardless of calendar year), combining both systems (ring designated as "ring 12 (formed 2020)" providing both relative and absolute temporal information), starting from graft year (year 1 ring representing first complete growing season after grafting with subsequent rings numbered sequentially), and requiring establishment date (precise ring year designation requiring known grafting or planting date providing temporal anchor point). Growth year rootstock annual ring in dendrochronological analysis includes= enabling cross-dating (comparing ring year sequences across multiple trunks synchronizing chronologies by matching patterns of wide and narrow rings to specific years), supporting climate reconstruction (correlating ring characteristics for specific years with climate data revealing temperature, precipitation, or other environmental patterns), documenting historical events (linking narrow or wide rings for specific years to known events such as droughts, freezes, or management changes), validating age assessment (confirming age determination by matching ring patterns to known reference chronologies ensuring accurate year assignments), and preserving long-term records (year-designated rings maintaining temporal information allowing reconstruction of conditions and events throughout trunk life). Growth year rootstock annual ring formation year correlation includes= linking to weather data (associating ring characteristics with meteorological records for formation year revealing climate-growth relationships), connecting to management records (correlating ring features with known vineyard practices during formation year assessing management effects), relating to known events (matching rings to documented events such as frost years, pest outbreaks, or irrigation installation during formation year), enabling cause-effect analysis (determining whether narrow ring in year X explained by drought or whether wide ring in year Y corresponds to optimal irrigation), and supporting interpretation (ring characteristics understood in context of specific year's documented conditions rather than just relative patterns). Growth year rootstock annual ring in trunk age context includes= early rings (years 1-5 representing initial establishment period typically showing wide rings from juvenile vigor), establishing rings (years 6-15 representing transition period showing progressively narrowing rings as growth rate stabilizes), mature rings (years 16-40 representing prime period showing relatively stable moderate ring widths with variation reflecting annual conditions), old rings (years 40-80+ representing aging period showing narrow rings from age-related growth decline), and ancient rings (years 80-100+ representing extreme age showing very narrow rings near growth cessation in very old trunk). Growth year rootstock annual ring ring year recording includes= showing visible annual sequence (rings forming chronological sequence from center outward with each ring representing consecutive year), enabling complete history (ring sequence preserving complete trunk developmental history from establishment through current year), maintaining temporal continuity (no gaps in ring sequence with every year represented by ring assuming no missing rings from extreme stress), demonstrating growth trajectory (ring sequence revealing growth rate changes, vigor patterns, and stress history throughout temporal progression), and preserving environmental record (year-by-year ring characteristics recording annual environmental conditions and vine responses across decades). Growth year rootstock annual ring in climate correlation includes= showing temperature effects (ring characteristics for specific years correlating with temperature patterns during formation revealing heat or cold effects on growth), revealing precipitation relationships (ring widths for specific years correlating with rainfall amounts documenting drought impacts or optimal moisture effects), documenting seasonal patterns (ring structure for specific years reflecting growing season length, frost dates, or other temporal climate features), enabling climate reconstruction (ring year sequences allowing inference of historical climate conditions for years predating instrumental records), and supporting climate change analysis (comparing recent ring years to earlier periods revealing trends in growth responses potentially indicating climate change effects). Growth year rootstock annual ring in management correlation includes= dating practice changes (identifying ring year when management practice implemented such as irrigation installation or fertility program by ring width or character changes), assessing treatment effects (comparing ring characteristics before and after specific year when treatment applied evaluating intervention effectiveness), documenting vineyard history (ring year sequence recording management history such as when vine established, when renovation occurred, or when ownership changed), supporting practice evaluation (ring patterns for specific years under known management approaches informing practice effectiveness assessment), and guiding future management (historical ring-management correlations informing management decisions for current and future years). Growth year rootstock annual ring missing ring phenomenon includes= rarely occurring in grapevine (missing rings uncommon in rootstock trunk due to typically annual cambial activity even under severe stress), potentially developing under extreme stress (exceptionally severe stress potentially suppressing cambial activity completely producing no ring that year though rare), requiring careful analysis (suspected missing ring requiring thorough examination and cross-dating confirmation before concluding ring absent), affecting age determination (missing ring if present causing age underestimation by one year requiring detection and correction), and showing regional patterns (missing rings if occurring potentially synchronous across site due to common extreme event enabling detection through cross-dating). Growth year rootstock annual ring in different ring year phases includes= juvenile phase rings (years 1-8 typically) showing characteristically wide rings from rapid young trunk growth, establishing phase rings (years 8-15 typically) showing moderate rings with transitional characteristics, mature phase rings (years 15-40 typically) showing stable rings with annual variation reflecting conditions, old phase rings (years 40-80 typically) showing progressively narrowing rings from age effects, and ancient phase rings (years 80-100+ if present) showing very narrow rings near growth cessation. Growth year rootstock annual ring specific year examples includes= first-year ring (year 1 post-grafting showing small ring from limited growth in establishment season potentially 0.3-0.5 cm), drought year ring (e.g. year 12 showing narrow ring 0.15 cm from severe summer drought documented in weather records), favorable year ring (e.g. year 5 showing wide ring 0.85 cm from optimal conditions during vigorous juvenile phase), transition year ring (e.g. year 10 showing moderate ring 0.45 cm marking shift from juvenile to mature growth rate), and recent year ring (current or previous year ring representing most recent growth assessable in increment core or cross-section). Growth year rootstock annual ring in trunk core interpretation includes= counting from cambium inward (numbering rings starting at outermost most-recent ring counting backward toward pith determining ring years), designating formation years (assigning calendar years to rings based on core sampling date and ring count), identifying missing rings (detecting any missing rings through cross-dating and pattern matching ensuring accurate year assignments), correlating with records (matching ring characteristics to known years with documented conditions validating correlations), and extracting temporal information (using year-designated rings to reconstruct chronological growth history and environmental response patterns). Growth year rootstock annual ring in vineyard assessment includes= determining vine age (counting rings providing exact age from grafting essential for old vine certification or vine evaluation), documenting growth history (ring year sequence revealing vigor patterns and stress history informing vine condition assessment), identifying critical years (locating rings from years with known problems such as freeze damage or disease outbreaks assessing lasting effects), evaluating management success (examining rings from years with specific management approaches assessing effectiveness), and supporting decision-making (ring year analysis informing decisions about vine retention, renewal, or replacement based on growth trajectory). Growth year rootstock annual ring comparison across vines includes= enabling synchronization (matching ring year patterns across multiple trunks in vineyard or region aligning chronologies), revealing common responses (similar ring patterns for specific years across vines indicating common environmental forcing rather than individual vine factors), identifying individual variation (differences in ring patterns for same years across vines revealing vine-specific responses or local condition variation), supporting statistical analysis (ring data for specific years across multiple vines enabling statistical testing of factors affecting growth), and validating interpretations (consistent patterns across vines for specific years confirming interpretations of climate or management effects). Growth year rootstock annual ring measurement standards includes= measuring from ring boundaries (determining radial width from inner boundary of ring to outer boundary for specific year), recording year designation (noting formation year allowing temporal correlation and cross-sample comparison), documenting measurement position (recording radial direction or circumferential position of measurement since rings potentially not uniform around trunk), noting ring characteristics (recording not just width but also structure, vessel characteristics, or other features for specific ring year), and maintaining metadata (preserving information about sampling date, trunk diameter, vine location enabling proper interpretation of ring year data). Growth year rootstock annual ring in time series analysis includes= creating growth chronologies (compiling ring width measurements by year creating temporal sequence of annual growth rates), analyzing temporal trends (examining ring year sequences revealing long-term trends such as progressive growth decline or improvement), identifying cycles (detecting periodic patterns in ring year data such as recurring favorable and unfavorable periods), correlating with external factors (relating ring year sequences to climate indices, management records, or other temporal data series), and modeling growth responses (using ring year data to develop and test models of growth responses to environmental or management factors). Growth year rootstock annual ring significance includes= providing precise temporal designation (ring year identification linking ring to specific point in time enabling chronological analysis and correlation), enabling historical reconstruction (year-designated rings allowing reconstruction of trunk and vineyard history from establishment through current time), supporting climate-growth analysis (ring formation year correlation with climate data revealing environmental effects on growth), documenting management effects (ring year designation enabling assessment of management practices and changes over time), facilitating cross-comparison (ring year designation allowing comparison across vines, sites, and years revealing patterns and relationships), preserving dated record (ring year information maintaining temporal archive of growth, conditions, and responses throughout trunk life), and enabling age determination (ring year counting providing exact trunk age essential for vine evaluation and management decisions).
Growth_Year_Rootstock_Annual_Ring emphasizes temporal designation and chronological identification and is distinguished from Rootstock_Annual_Ring (structural anatomical emphasis) by focus on formation year and position in temporal sequence. Critical for dendrochronological dating, historical correlation, climate-growth analysis, management effect assessment, cross-dating, age determination, and temporal documentation enabling each ring to be precisely positioned in time for correlation with events, conditions, and records.
Chronologischer Unterlagenring
Dated rootstock annual ring
Jahres-datierter Unterlagen-Jahresring
Temporal rootstock ring
Year-designated rootstock ring
Zeitlich bestimmter Wurzelstockring
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Schweingruber, F. H. (2007). Wood Structure and Environment. Springer.
Growth Year Rootstock Annual Ring
Guard cell turgor measurement in Grenache (anisohydric) versus Syrah (near-isohydric) during progressive soil drying reveals Grenache maintaining guard cell turgor of 2.4 MPa and stomatal conductance of 180 mmol/m²/s at leaf water potential of -1.2 MPa, while Syrah reduces guard cell turgor to 1.3 MPa and stomatal conductance to 45 mmol/m²/s at the same water potential, explaining the 3-fold difference in afternoon transpiration rate between varieties that underlies their contrasting drought response strategies.
Guard cell turgor quality is a cell turgor quality that inheres in the specialized epidermal cells flanking stomatal pores and represents the hydrostatic pressure whose modulation drives stomatal aperture changes controlling gas exchange between leaf interior and atmosphere. Within the BFO framework, this quality specifically characterizes the turgor of the only plant cell type capable of rapid, reversible turgor changes in response to environmental signals, where the unique radial arrangement of cellulose microfibrils in guard cell walls converts turgor increase into stomatal opening through differential expansion. The ontological significance lies in guard cell turgor serving as the mechanical effector of stomatal regulation, translating biochemical signals (ABA, CO₂, light quality) and hydraulic signals (leaf water potential) into the physical aperture changes that regulate photosynthetic carbon gain, transpirational water loss, and leaf temperature through evaporative cooling. Guard cell turgor ranges from approximately 1 MPa at stomatal closure to 3-5 MPa at maximum aperture in grapevines, with the turgor differential between guard cells and adjacent epidermal cells determining aperture rather than absolute guard cell turgor. The rapid turgor modulation enabling stomatal movements within minutes requires specialized ion transport systems (K⁺ channels, anion channels, H⁺-ATPases) and osmolyte metabolism (malate synthesis, starch-sugar interconversion) that distinguish guard cells from other cell types. Grapevine guard cell responses show cultivar variation in sensitivity to water deficit, with isohydric varieties reducing guard cell turgor and closing stomata at relatively high leaf water potential while anisohydric varieties maintain guard cell turgor and stomatal opening to lower water potentials. Understanding guard cell turgor quality enables interpretation of stomatal behavior, prediction of gas exchange responses to environment, and cultivar characterization for water use efficiency and drought adaptation.
Guard_Cell_Turgor_Quality addresses the specialized cell type controlling stomatal aperture, distinguishing from bulk leaf turgor or epidermal turgor that show different dynamics. This quality underlies stomatal conductance measurements commonly used in ecophysiology and water status assessment. Guard cell turgor changes occur on timescales of minutes to hours, contrasting with the slower turgor dynamics of growth-related processes in other cell types.
Schließzelldruck
Schließzellturgor
Spaltöffnungsturgor
Stomata-Zellturgor
guard cell pressure
ostiole cell turgor
stomatal guard cell turgor
stomatal turgor
Franks PJ et al. (2001) Guard cell pressure/aperture characteristics measured with the pressure probe. Plant, Cell & Environment 24(2): 163-171 Tombesi S et al. (2015) Stomatal closure is induced by hydraulic signals and maintained by ABA in drought-stressed grapevine. Scientific Reports 5: 12449
Guard Cell Turgor Quality
Guyot suitability quality evaluation in Côte de Nuits compared Pinot Noir clonal performance under the traditional single Guyot training characteristic of Burgundy. Clone 115 demonstrated excellent Guyot suitability quality with uniform bud fruitfulness along 6-8 bud canes (coefficient of variation 18%), appropriate cane diameter of 9-11 mm facilitating bending and tying, and upright shoot development from the horizontal cane creating organized canopy structure compatible with narrow Burgundian row spacing. Clone 667 showed good Guyot suitability quality with slightly higher vigor requiring longer cane retention of 8-10 buds for balance. Clone 777 exhibited reduced Guyot suitability quality with tendency toward basal bud dominance creating 35% higher fruitfulness at positions 1-3 compared to positions 6-8, leading some producers to prefer spur-pruned Cordon training for this clone where basal bud productivity represents an advantage rather than a limitation.
Guyot suitability quality is a trellis adaptation quality that inheres in a grapevine cultivar or clone and represents the degree of compatibility with Guyot cane-pruned training systems characterized by annual renewal of one (single Guyot) or two (double Guyot) fruiting canes selected from the previous season's growth and tied horizontally to a fruiting wire, with shoots developing upward from cane buds in vertical shoot positioned arrangement, representing the classic premium wine grape training approach that provides annual opportunity to select optimal fruiting wood while maintaining defined vine structure. This quality determines whether a cultivar possesses the cane-pruning suitability, bud fruitfulness patterns, and growth characteristics necessary to achieve optimal performance under the Guyot system that dominates premium wine production in regions including Burgundy, Champagne, Alsace, and other quality-focused viticultural areas worldwide. The ontological significance of Guyot suitability quality lies in its role as a primary determinant of cultivar selection for premium wine production systems where the annual cane selection of Guyot training enables optimization of fruiting wood quality, management of bud fruitfulness gradients along canes, and maintenance of vine balance through adjustment of cane length and bud number to match vine capacity. The biological requirements for high Guyot suitability quality include adequate cane fruitfulness with productive buds distributed along the cane length enabling effective horizontal positioning, moderate vigor producing canes of appropriate dimensions (typically 8-12 mm diameter) for training and tying without excessive brittleness or pliability, growth habit compatible with upright shoot development from horizontal cane position, and physiological response to cane pruning maintaining stable yield and vine balance across seasons. Cultivars exhibiting strong basal bud fruitfulness may perform acceptably under spur pruning but prove suboptimal for Guyot where mid-cane buds must contribute substantially to yield, while those with poor bud hardiness may suffer excessive blind buds following winter damage to horizontally-positioned canes. Assessment of Guyot suitability quality involves evaluation of bud fruitfulness distribution along canes, cane structural characteristics for training, shoot development patterns from horizontal cane positioning, and yield-quality outcomes compared to alternative pruning approaches.
Guyot_Suitability_Quality is_a Trellis_Adaptation_Quality, representing compatibility with cane-pruned training systems employing horizontal fruiting canes renewed annually. This class addresses the specific requirements of Guyot training including cane fruitfulness distribution, cane structural properties, and growth from horizontal positioning. Distinguished from Cordon_Suitability_Quality which addresses permanent cordon spur-pruned systems. The annual cane selection opportunity represents key characteristic enabling optimization of fruiting wood quality. Most relevant for premium wine production regions employing Guyot as the standard training approach.
Bogrebenschnitt-Kompatibilität
Flachbogen-Eignung
Guyot system adaptation
Guyot-Eignung
Streckererziehungs-Tauglichkeit
cane pruning suitability
cane training compatibility
horizontal cane training fitness
Galet P (2000) General Viticulture. Oenoplurimedia, Chaintre, pp. 189-245 Huglin P, Schneider C (1998) Biologie et écologie de la vigne. Lavoisier, Paris, pp. 253-278 Jackson RS (2008) Wine Science: Principles and Applications, 3rd Edition. Academic Press, pp. 108-142
Guyot Suitability Quality
A Chardonnay vine in Champagne trained to guyot simple (single guyot) bears a Guyot_Trained_Role with a single 8-10 bud cane arched along the cordon wire at 60 cm height, the standardized taille Champenoise enabling the hand labor intensive viticulture underlying Champagne production. A Pinot Noir vine in Burgundy's Gevrey-Chambertin serves in a Guyot_Trained_Role, the single-cane system representing centuries of regional refinement for optimal Pinot expression in Côte de Nuits terroirs. A Grüner Veltliner vine in Austrian Wachau training bears a Guyot_Trained_Role adapted to steep terraces, the cane system enabling annual adjustment to individual vine vigor and site conditions.
A cane-pruned vine role that inheres in a grapevine trained according to the guyot system developed by Jules Guyot in 19th-century France, characterized by annual retention of one or two one-year-old canes for fruiting plus short renewal spurs to provide replacement canes for subsequent seasons. Guyot training positions the retained cane(s) horizontally along a fruiting wire (typically 50-90 cm height), with the cane bent or arched from its origin on the vine head and tied to the wire, creating a defined fruiting zone from which vertical shoots emerge during the growing season. The renewal spur (typically 2 buds) is positioned below the fruiting cane to produce vigorous shoots that become next year's fruiting cane candidates, with cane selection at pruning choosing the best-positioned, best-matured replacement. Guyot training dominates French quality wine production (Burgundy, Champagne, Loire, Alsace) and has spread globally for varieties benefiting from cane pruning, becoming synonymous with premium production of Pinot Noir, Chardonnay, Riesling, and related varieties. The system's elegance lies in complete annual renewal—no permanent fruiting wood accumulates, potentially reducing disease reservoir and maintaining consistent vine architecture year after year.
Guyot_Trained_Role is subclassed by cane number—Single_Guyot (one cane) versus Double_Guyot (two canes)—with regional traditions and vine vigor determining the preferred variant. Jules Guyot's 19th-century systematization of cane pruning principles, published in his influential viticulture texts, established the framework that still governs French quality viticulture. The guyot approach requires annual decision-making at each vine—which cane to retain, how long to leave it, how to position it—creating skilled labor requirements that distinguish guyot-based production from more standardized spur systems. Regional variations on guyot abound: Champagne's specific regulations on bud numbers and training heights, Burgundy's distinction between guyot simple and guyot double by appellation, and Alsace's higher cane positioning adapted to the region's steep slopes and training stakes.
Flachbogenerziehung
Guyot-Erziehung
Guyot-Schnitt
Streckererziehung
guyot pruning
guyot system
guyot training
taille guyot
Guyot J (1860) Culture de la vigne et vinification, Librairie agricole de la Maison Rustique Galet P (2000) General Viticulture, Oenoplurimedia Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Huglin P, Schneider C (1998) Biologie et écologie de la vigne, Lavoisier
Guyot Trained Role
Chardonnay showing hair-like fine feeder roots in April, careful soil washing revealing extremely thin roots 0.3-0.4 mm diameter 1-2 cm length, microscopic examination showing 1-2 xylem vessels, cortex 2 cell layers thick, prolific root hair production 480 per cm creating fuzzy appearance, arbuscular mycorrhizal colonization 72%, roots so fragile requiring immediate examination before drying, hair-like fine feeder roots representing extreme absorption specialization approaching root hair dimensions
Hair like fine feeder root is fine feeder root showing extremely small diameter 0.2-0.5 mm approaching or overlapping root hair dimensions, characterized by hair-like thinness creating almost thread-like appearance, extreme fragility requiring careful handling, minimal internal structure with 1-3 xylem vessels and minimal cortex, very short length typically 0.5-3 cm, and serving purely absorptive function representing extreme specialization where root structure minimized to absolute necessity creating maximum absorption per carbon investment, with hair-like fine feeder roots showing distinctive features including diameter sometimes <0.3 mm overlapping large root hair range creating ambiguity in root-hair distinction, extremely high surface-to-volume ratio potentially 50-100+ mm²/mm³ among highest in root system, prolific root hair production potentially 400-600 hairs per centimeter creating fuzzy appearance obscuring root cylinder, lifespan often 1-4 months representing most ephemeral root type, and demonstrating that hair-like thinness represents functional extreme where roots essentially function as thick root hairs creating extended absorption surface with minimal structural investment. Hair like fine feeder root diameter and morphology includes= showing hair-like diameter (0.2-0.5 mm creating extremely thin thread-like roots), exhibiting overlap with root hairs (smallest hair-like roots 0.2-0.3 mm overlapping large root hairs 0.15-0.25 mm creating continuum), demonstrating very short length (typically 0.5-3 cm limiting individual root exploration distance), revealing extreme aspect ratio (length potentially only 10-20 times diameter versus 50-100+ in coarser roots), and displaying thread-like appearance (visual appearance similar to fine threads or hairs rather than recognizable roots). Hair like fine feeder root anatomical extreme includes= showing minimal vasculature (only 1-3 xylem vessels sometimes single vessel providing minimal transport capacity), exhibiting thin cortex (cortex possibly 1-3 cell layers thick approaching theoretical minimum), demonstrating small stele (vascular cylinder extremely small potentially <100 micrometers diameter), revealing large epidermis proportion (epidermis representing large proportion of root cross-section due to small diameter), and displaying simple cellular organization (cellular structure simplified reflecting minimal functional requirements beyond absorption). Hair like fine feeder root root hair dominance includes= showing prolific production (hair-like roots potentially producing 400-600+ root hairs per centimeter), exhibiting hair dominance (root hairs potentially outnumbering or outweighing root cylinder cells), demonstrating surface multiplication (root hairs increasing effective surface 15-30 fold given already small root diameter), revealing functional blur (distinction between hair-like root and its root hairs functionally blurred both serving absorption), and displaying hair dependence (hair-like root function critically dependent on root hair production and maintenance). Hair like fine feeder root extreme ephemerality includes= showing very short lifespan (median longevity 1-4 months among shortest-lived roots), exhibiting rapid turnover (hair-like root population potentially replacing 3-6 times per year), demonstrating high mortality (individual hair-like roots quickly dying after brief functional period), revealing production intensity (maintaining population requiring extremely high continuous production rates), and displaying expendable extreme (hair-like roots representing ultimate expendable absorption organs). Hair like fine feeder root absorption specialization includes= providing maximum specific uptake (uptake per unit hair-like root biomass potentially highest among all plant tissues), exhibiting extreme efficiency (hair-like roots showing optimal absorption-to-cost ratio), demonstrating pure function (hair-like roots performing exclusively absorption with zero structural or storage function), revealing niche specialization (hair-like roots specialized for exploiting small soil pores and accessing microsites), and displaying mycorrhizal enhancement (arbuscular mycorrhizae potentially essential for hair-like root function compensating for minimal internal structure). Hair like fine feeder root fragility and vulnerability includes= showing extreme fragility (hair-like roots easily broken by minimal mechanical stress), exhibiting sampling difficulty (hair-like roots often lost or damaged during soil sampling creating underestimation), demonstrating desiccation sensitivity (hair-like roots rapidly dying if soil dries given minimal water storage capacity), revealing stress vulnerability (hair-like roots first to die under any stress given minimal reserves and extreme structure), and displaying handling challenges (studying hair-like roots requiring exceptional care avoiding damage). Hair like fine feeder root functional context includes= representing absorption extreme (hair-like roots exemplifying maximum absorption specialization in root system), enabling microsite exploitation (hair-like thinness allowing penetration into smallest soil pores accessing resources unavailable to coarser roots), demonstrating carbon efficiency (hair-like roots providing maximum uptake per carbon invested in construction), revealing evolutionary extreme (hair-like fine roots representing evolutionary endpoint of absorption specialization), and displaying niche function (hair-like roots filling specialized niche in root system functional repertoire). Hair like fine feeder root research challenges includes= requiring specialized methods (conventional root sampling often missing or destroying hair-like roots), showing quantification difficulty (accurately quantifying hair-like root abundance and function challenging given fragility), exhibiting preservation problems (fixing or storing hair-like roots difficult due to extreme delicacy), demonstrating identification challenges (distinguishing hair-like roots from large root hairs or fungal hyphae sometimes difficult), and revealing knowledge gaps (hair-like fine roots potentially understudied and underappreciated due to methodological difficulties).
Hair_Like_Fine_Feeder_Root emphasizes extreme thinness approaching root hair dimensions and is distinguished from coarser fine roots by focus on hair-like diameter. Critical for understanding extreme diameter, minimal anatomy, root hair overlap, maximum efficiency, extreme fragility, and the thread-like roots representing absorption specialization extreme with minimal structural investment.
Fadenförmige Saugwurzel
Filamentous fine root
Haarähnliche Feinwurzel
Root hair-sized root
Thread-like feeder root
Wurzelhaar-große Wurzel
Hair Like Fine Feeder Root
Chardonnay grafted on Teleki 5C rootstock at 13 years showing hair-like fine tertiary roots in organic-rich topsoil at 28 cm depth, sixth-order hair-like roots 0.4 mm diameter 1.8 cm length with extremely dense root hair coverage estimated 340 hairs per cm creating fuzzy appearance, seventh-order terminal hair-like roots 0.25 mm diameter 0.9 cm length, hair-like roots showing minimal vascular development with only 2-3 xylem vessels, primary anatomy throughout, mycorrhizal colonization 72% with abundant external hyphae, sequential sampling indicating hair-like root median lifespan 3.8 months, hair-like roots providing estimated 42% of total vine nitrogen uptake despite comprising <1.5% root system biomass
Hair like fine tertiary root is fine tertiary root showing extremely small diameter approaching root hair dimensions, emphasizing hair-like delicate structure and maximum surface area-to-volume ratio, characterized by diameter typically 0.2-0.6 mm approaching the lower size limit for roots blurring distinction from large root hairs, very short length typically 0.5-3 cm creating minimal biomass investment, fifth-order or higher branching position representing terminal branches in root hierarchy, abundant prolific root hair production with hair density potentially 200-400+ per centimeter creating fuzzy appearance, and serving exclusively for absorption functioning essentially as extended root hairs maximizing resource uptake efficiency while minimizing carbon investment, with hair-like fine tertiary roots showing extreme fragility breaking easily during handling or excavation, primary anatomy throughout with minimal vascular development reflecting minute size, very short lifespan typically 1-6 months representing most ephemeral root component, and demonstrating that hair-like roots represent extreme expression of absorptive specialization trading longevity and structural integrity for maximum uptake surface per unit construction cost. Hair like fine tertiary root diameter characteristics includes= showing extremely small diameter (0.2-0.6 mm approaching root hair size dimensions), approaching lower size threshold (hair-like roots near minimum viable root diameter below which vascular tissue development impossible), exhibiting hair-like appearance (fine roots appearing thread-like or hair-like to naked eye), demonstrating high surface-to-volume ratio (small diameter maximizing surface area per unit volume optimizing absorption per biomass), and revealing dimensional transition (hair-like roots representing gradation between roots and root hairs rather than sharp distinction). Hair like fine tertiary root length and morphology includes= showing very short length (hair-like roots typically 0.5-3 cm rarely exceeding 5 cm reflecting terminal position), exhibiting unbranched or minimally branched (hair-like roots typically terminal lacking further branching or producing only occasional branches), demonstrating delicate structure (hair-like roots extremely fragile easily damaged or broken), displaying white to translucent appearance (young hair-like roots very light-colored due to thin tissues and limited pigmentation), and revealing high elongation rate (hair-like roots potentially growing 1-5 mm per day during active periods despite small final size). Hair like fine tertiary root anatomical simplification includes= showing extremely reduced vascular cylinder (stele in hair-like roots containing only 1-4 xylem vessels and minimal phloem reflecting minute size), exhibiting thin cortex (cortical tissue only 3-8 cell layers thick providing minimal storage or transport capacity), demonstrating prominent epidermis (outer cell layer relatively large proportion of total root volume), containing minimal endodermis development (casparian strip present but suberin lamellae potentially absent or reduced), and lacking differentiated tissues (hair-like root simplicity approaching lower limit of root anatomical organization). Hair like fine tertiary root prolific root hair production includes= bearing extremely abundant root hairs (hair-like roots showing dense root hair coverage potentially 200-400+ hairs per centimeter length), exhibiting hair-like root covered in hairs (creating appearance of hair covered with finer hairs maximizing surface), demonstrating hair dominance (root hairs potentially comprising >90% of total absorption surface area of hair-like root system), showing rapid hair turnover (individual root hairs living only 2-10 days with continuous production along growing root), and revealing surface multiplication (root hairs increasing effective surface 10-30 fold compared to already-small root cylinder). Hair like fine tertiary root absorptive optimization includes= maximizing uptake per biomass (hair-like roots showing highest absorption per gram root tissue of any root type), demonstrating minimal construction cost (very small size and short lifespan minimizing carbon investment required), exhibiting rapid resource return (short-lived hair-like roots providing quick return on construction investment), showing specialized absorption (hair-like roots exclusively absorptive lacking any transport or structural function), and revealing evolutionary optimization (hair-like root characteristics representing optimal solution for ephemeral resource acquisition in variable environments). Hair like fine tertiary root lifespan dynamics includes= showing extremely short lifespan (hair-like roots typically living 1-6 months representing most ephemeral component), exhibiting very high turnover (hair-like root population potentially replacing itself 2-4+ times per year), demonstrating pulse production (hair-like root initiation showing pronounced peaks in spring and fall), revealing immediate mortality response (hair-like roots dying within days under drought, temperature stress, or damage), and contributing substantially to turnover flux (hair-like roots despite small biomass providing major proportion of root production and mortality flux). Hair like fine tertiary root spatial distribution includes= showing extreme patchiness (hair-like root density varying >100-fold spatially concentrated in resource hotspots), concentrating in upper topsoil (majority of hair-like roots 15-50 cm depth in organic-rich surface horizons), proliferating in nutrient patches (hair-like root density increasing >50-fold in localized zones around organic matter or fertilizer), demonstrating absence in poor zones (hair-like roots completely absent from dry, compacted, or nutrient-depleted areas), and revealing resource tracking (hair-like root distribution closely reflecting soil resource heterogeneity at fine spatial scales). Hair like fine tertiary root functional significance includes= providing disproportionate absorption (hair-like roots despite representing <2% of root biomass potentially contributing 30-50% of water and nutrient uptake), demonstrating efficient foraging (hair-like roots enabling resource acquisition from micro-patches inaccessible to larger roots), showing rapid response capacity (hair-like root production and mortality enabling quick adjustment to changing soil conditions), exhibiting minimal opportunity cost (very low construction investment in hair-like roots allowing proliferation with minimal carbon penalty), and revealing adaptive strategy (hair-like root characteristics representing high-risk high-reward strategy trading longevity for responsiveness).
Hair_Like_Fine_Tertiary_Root emphasizes extremely small diameter 0.2-0.6 mm and hair-like appearance and is distinguished from larger fine roots by focus on minute size approaching root hair dimensions. Critical for understanding absorption optimization, surface maximization, extreme ephemerality, minimal construction cost, resource foraging, and the finest most ephemeral roots representing extreme absorptive specialization maximizing uptake per carbon investment.
Capillary root
Fadenförmige Feinwurzel
Haarähnliche feine Tertiärwurzel
Kapillarwurzel
Thread-like fine root
Ultra-fine root
Pregitzer, K. S., et al. (2002). Fine root architecture of nine North American trees. Ecological Monographs, 72(2), 293-309.
Hair Like Fine Tertiary Root
A half-developed tendril at node 9 on a moderately stressed shoot of Vitis vinifera cv. Grenache in southern Spain, measuring 8.5 cm length (approximately 48% of normal 17-18 cm for this cultivar and position), exhibiting bifurcated structure with both arms developed to 3.5 cm, complete primary vascular bundles visible throughout but secondary xylem development limited to basal 3 cm, epidermis fully formed in proximal half but thinner and less cuticularized in distal half, mechanosensory cells present and apparently functional in mid-tendril regions (nodes 3-6 cm from base) but sparse or absent in distal 2.5 cm, light brown coloration in basal 3 cm indicating moderate lignification but green and flexible in distal regions, occurring during sustained moderate water stress (soil moisture 55-65% field capacity throughout development) in a non-irrigated vineyard, showing weak coiling response when experimentally contacted at mid-regions but no response at distal portions, supporting approximately 200 grams before showing excessive bending, measured at BBCH stage 77 (berries beginning to touch) in July.
A partially developed tendril that achieved approximately 50% completion across multiple developmental parameters in Vitis vinifera, characterized by length of 7-10 cm representing 40-55% of normal mature tendril length for cultivar and position, completion of basic structural framework including primary vascular differentiation and branching architecture where applicable but incomplete secondary development including limited lignification and partial cuticle formation, functional tissue organization in proximal regions transitioning to incomplete differentiation in distal regions, presence of mechanosensory cells in some areas but absent or non-functional in others, and overall developmental status representing a median position between initiation and full maturation across anatomical, physiological, and functional criteria. Half-developed tendrils result from stress conditions or resource limitations that permit approximately half the normal developmental investment before causing developmental arrest, typically associated with moderate chronic stress including sustained water deficit at 50-60% of optimal irrigation, moderate crop loads creating consistent but not severe competition for resources, gradual onset nutrient deficiency affecting growth rates but not causing acute deficiency symptoms, or environmental conditions maintaining suboptimal but not severely limiting temperatures or light levels throughout most of the development period. These tendrils represent balanced incomplete development with proportional achievement across developmental parameters rather than mosaic patterns of success and failure, providing limited functional capacity in low-stress conditions but inadequate support under normal vineyard loading scenarios.
Half-developed tendrils are distinguished from other partially developed tendrils by approximate median achievement (40-55%) across multiple developmental parameters rather than either early arrest (70%), and from incomplete tendrils by proportional development across tissue systems rather than strongly asynchronous patterns. This classification represents a relatively narrow developmental range indicating specific moderate stress intensities that allow sustained but reduced growth rates throughout development. Half-development status is diagnostically valuable because it suggests quantifiable stress levels amenable to precise amelioration - increasing resource availability by approximately 50% should theoretically restore full development. The occurrence of half-developed tendrils across vine populations indicates systemic moderate stress requiring management intervention such as supplemental irrigation, fertilization adjustment, or crop load reduction to restore normal developmental capacity.
fifty percent developed tendril
fünfzig Prozent ausgereifte Ranke
halb-entwickelte Ranke
half-mature tendril
mid-completion tendril
mittlere Vollständigkeit Kletterorgan
Based on Chaves et al. (2010) "Grapevine under deficit irrigation= hints from physiological and molecular data" Annals of Botany 105=661-676 Dry et al. (2001) "Strategic irrigation management in Australian vineyards" Journal International des Sciences de la Vigne et du Vin 35=129-139 Petrie et al. (2000) "The effect of leaf removal and canopy height on whole-vine gas exchange and fruit development of Vitis vinifera L. Sauvignon blanc" Functional Plant Biology 27=173-184 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 497-520.
Half Developed Tendril
A half-expanded young leaf at node 10 on a shoot of Vitis vinifera cv. Pinot Noir in Burgundy, France, measuring 7.8 cm length by 7.2 cm width representing approximately 51% of expected mature dimensions of 15.3 cm × 14.1 cm for this position and cultivar, lamina area of 36 cm² representing 26% of expected mature area of 138 cm², exhibiting flat architecture with five distinct lobes and moderate sinuses showing final pattern though dimensions will continue increasing, light-medium green coloration with chlorophyll content of 345 μg cm⁻², slight bronze tinge along primary veins from residual anthocyanin at 0.8 mg g⁻¹ fresh weight, moderately soft texture with palisade cells averaging 26 μm diameter representing approximately 50% progress from initial 18 μm to final 35 μm dimensions, developing tissue organization with two palisade layers totaling 52 μm (61% of mature 85 μm) and spongy mesophyll of 78 μm (65% of mature 120 μm), functional stomata showing partial closure response to VPD increase though with reduced sensitivity, partial cuticle development creating slight sheen particularly on adaxial surface, demonstrating photosynthesis rate of 6.8 μmol CO₂ m⁻² s⁻¹ representing 56% of mature capacity and remaining net sink with daily carbon consumption of 8 mg exceeding photosynthetic production of 18 mg per day, occurring at 14 days post-emergence at midpoint of typical 28-day expansion period, measured at shoot BBCH stage 75 (berries pea-sized) in late June, representing the median expansion checkpoint with equal development ahead and behind.
A partially expanded young leaf at the midpoint of expansion achieving approximately half of final mature dimensions in Vitis vinifera, characterized by dimensions typically 45-55% of cultivar-specific and position-specific mature size with length and width both in 50% ± 5% range, lamina area approximately 25% of final mature area due to quadratic relationship between linear dimensions and area, flat or nearly flat architecture with major structural features evident but not fully refined, light to medium green coloration with chlorophyll concentrations typically 300-380 μg cm⁻² representing 60-75% of mature levels, traces of anthocyanin remaining particularly along veins creating slight bronze tinge but greatly reduced from early stages, moderately soft texture with cells approximately halfway through expansion from initial to final dimensions, intermediate tissue differentiation with palisade layers beginning organization but spongy mesophyll still loosely arranged with large air spaces, developing vascular system with recognizable but incompletely matured xylem and phloem, functional stomata with partial but not complete environmental responsiveness, partial cuticle development creating slight surface sheen, and photosynthetic capacity typically 50-65% of mature leaf rates with leaves remaining net carbon sinks consuming more resources for continued growth than produced through photosynthesis but transitioning toward eventual carbon balance. Half-expanded young leaves represent the midpoint developmental checkpoint where approximately equal expansion remains ahead and behind, with cell expansion proceeding at maximum rates if conditions are optimal, tissue differentiation processes advancing rapidly establishing anatomical characteristics that will determine mature leaf function, and chloroplast development accelerating producing the chlorophyll and photosynthetic apparatus that will support mature photosynthetic capacity.
Half-expanded young leaves represent a defined developmental checkpoint characterized by achievement of approximately 50% linear dimensions and 25% area, providing standardized reference for assessing expansion progress and comparing expansion rates across conditions. Assessment requires comparison to known mature size for the specific cultivar and node position, as absolute dimensions vary substantially. The half-expansion stage typically occurs at approximately 40-50% of total expansion duration measured from emergence to maturity, though exact timing varies with environmental conditions affecting expansion rate. Half-expanded leaves show intermediate characteristics across all parameters including size, coloration, texture, anatomy, and function, with none fully mature but all substantially developed beyond initial states. Stress or resource limitation occurring at half-expansion substantially affects final size, as approximately 75% of final area achievement remains ahead requiring continued cell expansion that depends critically on water, nutrients, and favorable environmental conditions. Management decisions about leaf removal for crop exposure or disease management often target pre-half-expansion stages to minimize photosynthetic capacity loss while achieving management objectives.
fifty percent expanded leaf
fünfzig Prozent ausgedehnt Blatt
halb-expandiertes junges Blatt
intermediate size young leaf
midpoint expansion leaf
mittelpunkt Expansion Weinblatt
Based on Palliotti & Cartechini (2000) "Developmental changes in gas exchange activity in flowers, berries, and tendrils of field-grown Cabernet Sauvignon" American Journal of Enology and Viticulture 51=317-323 Poni et al. (2009) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 60=397-407 Tsukaya (2013) "Leaf development" The Arabidopsis Book 11=e0163 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 209-232.
Half Expanded Young Leaf
The hand harvest protocol for Krug Grande Cuvée Champagne specifies picking teams of 600-800 trained vendangeurs deployed across the house's 250+ parcels sourced from 25 Champagne villages over a 2-3 week harvest period — each picker is equipped with curved secateurs (sécateur à vendange) and a small 8 kg capacity basket (panier), instructed to cut clusters with a 2-3 cm stem handle (to facilitate subsequent whole-cluster pressing without excessive stem contact), reject visibly damaged, botrytized, or unripe clusters at the vine, and deposit clusters gently into the panier without stacking more than two layers deep to prevent compression — filled baskets are transferred to perforated 50 kg plastic crates (caisses) at the end of each row, loaded onto refrigerated transport within 20 minutes, and delivered to the nearest press center within 90 minutes of picking for immediate whole-cluster pressing.
A harvest method description that documents the organization, procedures, and quality standards for manual grape harvesting by human pickers, specifying the picking team size and training, cluster selection and cutting technique, collection container type and capacity, vineyard-to-winery transport logistics, and time-from-vine-to-processing targets that ensure the delivery of intact, undamaged fruit in optimal condition for winemaking.
Hand_Harvest_Protocol documents the labor-intensive traditional harvest method that remains the standard for premium wine production worldwide. The fundamental advantage of hand harvesting is selectivity — a trained human picker can evaluate each cluster individually, reject damaged or unripe fruit, cut the cluster cleanly with minimal physical damage, and handle the harvested fruit gently throughout the collection and transport chain: this selectivity is unmatched by any current mechanical harvesting system, particularly for operations requiring intact whole clusters (Champagne, whole-cluster fermentation), selective picking of individually ripe clusters (tries successives for botrytized wine production), or navigation of terrain inaccessible to machines (steep slopes in the Mosel, Douro, Côte-Rôtie). The principal constraints of hand harvesting are labor cost (typically 5-20 times more expensive per hectare than mechanical harvesting), speed (a team of 20-30 pickers harvests 0.3-0.5 ha per day versus 2-3 ha per hour for a mechanical harvester), and labor availability (seasonal harvest labor is increasingly scarce in many wine regions, driving wages upward and creating harvest scheduling challenges).
Handlese-Protokoll
Handlese-Verfahren
hand picking specification
manual harvest protocol
manuelle Lese-Spezifikation
vendange manuelle protocol
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 280-290 Comité Interprofessionnel du Vin de Champagne (CIVC) (2023) Cahier des charges de la vendange en Champagne Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1, 2nd Ed, Wiley pp: 235-250
Hand Harvest Protocol
A hard lignified one-year cane on Vitis vinifera cv. Syrah in a warm, dry-farmed vineyard, examined during winter pruning showing exceptionally dark brown almost reddish coloration, very firm resistance to bending requiring considerable force, 10mm diameter with cross-section revealing minimal central pith (2mm) surrounded by dense secondary xylem tissue (4mm radial thickness), thick periderm with multiple cork layers, this cane surviving experimental freeze testing to -22°C without bud or vascular damage (compared to -18°C for normally lignified canes from same vineyard), making it highly desirable retention as fruiting wood despite challenges in bending to horizontal trellis wire without risk of cracking.
A fully lignified one-year cane exhibiting exceptional firmness and structural rigidity beyond typical lignification, characterized by very dense woody tissue, thick secondary cell walls with heavy lignin deposition, reduced pith diameter relative to overall cane diameter, dark brown coloration indicating extensive secondary compound accumulation, and superior cold hardiness compared to normally lignified canes of similar age and cultivar. Hard lignified one-year canes result from growing conditions promoting extensive secondary growth including moderate water stress during late season, adequate but not excessive nitrogen nutrition, high light exposure, appropriate crop loads, and extended ripening period with warm days and cool nights, producing wood of exceptional quality for winter survival and spring productivity though sometimes showing reduced flexibility making trellis positioning more challenging.
Hard lignified one-year canes represent the upper extreme of wood maturation, beneficial for winter survival and structural support but potentially problematic for training systems requiring cane bending or positioning. The excessive firmness can lead to cracking or splitting when forced into tight curves or sharp angles, requiring careful handling during trellis attachment or consideration of alternative positioning strategies. Hard lignification typically results from moderate stress (water, crop load) that limits vegetative growth while promoting secondary development, common in Mediterranean climates, dry-farmed vineyards, and old vines with limited vigor. While hard lignification benefits cold hardiness and disease resistance (dense wood resists pathogen penetration), it may indicate suboptimal vine balance if excessively stressed vines show reduced shoot vigor or yield potential in following seasons.
besonders feste Rute
dense woody cane
extra-firm lignified cane
hart verholzte Rute
heavily lignified cane
Based on Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 245-267 Fennell (2004) "Freezing tolerance and injury in grapevines" Journal of Crop Improvement 10=201-235 Lovisolo et al. (2010) "Effects of water stress on vessel size and xylem hydraulic conductivity in Vitis vinifera L." Journal of Experimental Botany 61=2749-2759.
Hard Lignified One Year Cane
Merlot hard seed coat at harvest maturity (BBCH 89) exhibits dark brown coloration, sclereid wall thickness 18-22 μm, crushing resistance 20-25 N force, water imbibition rate 0.8-1.5 mg/seed/hour, tannin polymerization index 72-78%, and requires 100-110 day stratification for 50% reduction in mechanical resistance enabling germination
A hard seed coat is a seed coat characterized by extensive lignification and sclerification of cell wall layers, resulting in mechanical rigidity, resistance to crushing, and reduced permeability to water and gases, representing the mature developmental state of Vitis vinifera seed coats that provides maximal physical protection and contributes to seed dormancy mechanisms. Seed coat hardening occurs during late seed development (8-16 weeks post-fertilization, BBCH 83-89) through= progressive lignification of sclereid cell walls (deposition of lignin polymers increasing wall thickness from 5-8 μm in early development to 15-25 μm at maturity), secondary wall thickening in sclerenchyma cells (cellulose and hemicellulose deposition), tannin polymerization and cross-linking with cell wall components (condensed tannins forming covalent bonds with proteins and polysaccharides), and progressive desiccation reducing tissue moisture content. Hard seed coat characteristics include= high mechanical strength (requiring 15-30 N crushing force compared to 2-5 N for immature soft seed coats), reduced water permeability (imbibition rate 0.5-2 mg water/seed/hour compared to 5-10 mg/seed/hour for soft immature coats), dark coloration (brown to black indicating complete tannin polymerization), and resistance to microbial penetration (lignified tissues are less susceptible to enzymatic degradation). Cellular structure of hard seed coat includes= outer epidermis with thick cuticle and lignified cell walls, multiple layers of heavily lignified sclereids with 15-25 μm wall thickness and small cell lumens, tannin-rich hypodermal layers with polymerized proanthocyanidins, and compressed inner integument remnants. Hard seed coat mechanical properties serve multiple functions= physical protection of embryo and endosperm from crushing damage during berry processing, resistance to premature water uptake preventing off-season germination, mechanical constraint contributing to dormancy (embryo expansion resistance requiring stratification-induced seed coat softening), and durability supporting seed survival in soil seed banks. Seed coat hardness development is influenced by= seed maturity at berry harvest (under-ripe harvest results in partially hardened seed coats with intermediate mechanical properties), environmental conditions during late berry development (water stress can enhance lignification, while excessive moisture may reduce hardening), cultivar genetics (some varieties develop harder seed coats than others at equivalent maturity), and seed position within berry (central seeds may achieve greater hardness than peripheral seeds). Hard seed coat implications for viticulture and enology include= seed browning as visual indicator of approaching harvest maturity, crushing resistance affecting seed breakage during berry processing (hard mature seeds less likely to break releasing harsh tannins), seed tannin extractability during fermentation (highly polymerized tannins in hard seed coats are less extractable than monomeric tannins in soft immature coats), and germination requirements (hard seed coats necessitate extended stratification 90-120 days for softening and increased permeability enabling germination).
Hard_Seed_Coat emphasizes mechanical rigidity and lignification and is distinguished from Protective_Seed_Coat (broader protective functions) and Seed_Coat (complete structure and development) by focus on mature hardened state and associated mechanical properties. Critical for harvest maturity assessment, seed breakage risk during processing, dormancy mechanisms requiring stratification, and understanding tannin extractability differences between mature and immature seeds affecting wine quality.
Hardened testa
Harte Samenschale
Lignified seed coat
Sklerenchymatische Samenhülle
Verholzte Testa
Hard Seed Coat
Hardiness development rate assessment comparing cultivars through autumn reveals the acclimation velocity variation affecting early-season vulnerability, with fast-developing cultivar (Concord) gaining 2.8°C hardiness per week during October (reaching protective -15°C by October 31), moderate-rate cultivar (Chardonnay) gaining 1.9°C per week (reaching -15°C by November 12), and slow-developing cultivar (Cabernet Franc) gaining only 1.2°C per week (not reaching -15°C until November 28), with an early November freeze (-12°C on November 5) causing no damage to Concord, moderate injury to Chardonnay, and severe damage to Cabernet Franc, demonstrating how development rate affects survival of early-season cold events.
Hardiness development rate quality is a cold hardiness quality that represents the speed at which freeze tolerance increases during autumn and early winter acclimation, characterizing how rapidly tissues transition from the tender state of active growth to the hardy state required for winter survival, with development rate determining how quickly protection is established and thus the vulnerability window before full hardiness is achieved. Within the BFO framework, this quality specifically addresses the acclimation velocity dimension of cold hardiness, distinguishing how fast hardiness develops from the ultimate level achieved while recognizing that development rate determines the timing of protection establishment, with rapid development providing earlier protection (reducing vulnerability to early cold events) and slow development extending the vulnerable period during which cold snaps could cause damage before hardiness is adequate. The ontological significance lies in hardiness development rate determining the match between acclimation progress and seasonal cooling, where vines with rapid development can achieve protective hardiness before early cold events while those with slow development may be caught incompletely acclimated by early-season freezes, making development rate as important as maximum hardiness for survival in climates where early cold events precede the gradual seasonal cooling that drives optimal acclimation. The factors affecting hardiness development rate include cultivar genetics (some cultivars inherently acclimating faster than others), vine condition (well-conditioned vines with adequate reserves acclimating more rapidly), autumn conditions (gradual cooling promoting steady development, while warm autumns may delay acclimation start), and the interaction between temperature and photoperiod signals (both contributing to acclimation triggering and progression). The practical implications of development rate include the timing of autumn management operations that may affect acclimation (harvest timing, irrigation cessation), the risk assessment for early-season cold events, and the cultivar selection considerations for regions where early freezes before full winter are common. Understanding hardiness development rate quality enables prediction of when protective hardiness will be achieved, assessment of early-season cold vulnerability, and management timing that supports rapid hardiness development.
Hardiness_Development_Rate_Quality addresses the velocity of freeze tolerance increase during autumn acclimation. This quality determines vulnerability to early-season cold events before full hardiness is achieved. The class supports autumn risk assessment through hardiness development rate characterization.
Härteentwicklungsrate
Kältetoleranzentwicklung
acclimation rate
cold tolerance development
hardiness gain rate
Mills LJ et al. (2006) Timing of low temperature exposure affects cold hardiness and survival of dormant grapevine buds. American Journal of Enology and Viticulture 57(2): 232-239 Ferguson JC et al. (2014) Dynamic thermal time model of cold hardiness for dormant grapevine buds. Annals of Botany 113(5): 801-811
Hardiness Development Rate Quality
Investigation of Hardiness_Retention_Disposition at Michigan State University exposed dormant canes to controlled warming treatments (15°C for 3, 7, or 14 days) at monthly intervals from December through February, then assessed residual hardiness by differential thermal analysis. Cultivar differences in Hardiness_Retention_Disposition were substantial. 'Frontenac' demonstrated exceptional hardiness retention, losing only 2°C after 7 days at 15°C in January (from -33°C to -31°C) and fully recovering this loss within 5 days of return to cold conditions. 'Riesling' showed moderate retention, losing 5°C after equivalent treatment (from -23°C to -18°C) with incomplete recovery. 'Pinot Noir' exhibited poor Hardiness_Retention_Disposition, losing 8°C after 7 days warming (from -20°C to -12°C) with only 3°C recovery. The timing of warm exposure affected all cultivars, with February warming causing greater hardiness loss than December warming, reflecting decreasing dormancy depth through winter. Field relevance was demonstrated following a January thaw (7 days above 10°C) in year 3 of the study. Post-thaw hardiness measurements showed 'Frontenac' at -30°C (3°C loss), 'Riesling' at -16°C (7°C loss), and 'Pinot Noir' at -11°C (9°C loss). When temperatures subsequently dropped to -22°C, 'Pinot Noir' suffered 85% primary bud mortality while 'Frontenac' was undamaged. These findings establish Hardiness_Retention_Disposition as critical for winter survival in fluctuating climates.
A Hardiness_Retention_Disposition is an Acclimation_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to maintain acquired freezing tolerance at or near maximum levels throughout the midwinter period despite fluctuating temperatures, preventing premature loss of hardiness during transient warm periods before spring. This disposition represents the stability of the cold-hardy state once achieved, distinct from the initial acquisition of hardiness (acclimation rate) or the ultimate level achieved (maximum hardiness). Hardiness retention depends on dormancy depth and stability, with deeply dormant buds maintaining hardiness despite warm exposure while shallow-dormant buds may respond to warmth by initiating deacclimation. Genotypes differ substantially in their sensitivity to midwinter warming, with some cultivars showing rapid hardiness loss (2-4°C per day of warm exposure) while others maintain hardiness tenaciously (< 0.5°C loss per day). Strong Hardiness_Retention_Disposition is particularly valuable in climates with variable winter weather including midwinter thaws followed by return to severe cold, as hardiness lost during warm periods may not be fully recovered before subsequent cold events. The physiological basis of hardiness retention involves maintenance of cryoprotectant levels, continued suppression of growth-related gene expression, and stability of dormancy-regulating hormone balances (high ABA, low GA/cytokinin) despite temperature fluctuations.
Hardiness_Retention_Disposition is_a Acclimation_Disposition. Hardiness retention determines vulnerability to cold events following midwinter thaws, a common damage scenario in variable continental climates. Retention is linked to dormancy depth, with deeper dormancy conferring greater hardiness stability. Cultivars differ substantially in retention capacity, partially independent of maximum hardiness. Poor hardiness retention combined with variable winter weather creates conditions for severe damage even when maximum hardiness would be adequate for static cold exposure.
Härte-Beibehaltungs-Disposition
Härtestabilität
Winterhärte-Aufrechterhaltung
cold tolerance persistence
midwinter hardiness maintenance
Kalberer SR et al. (2006) Deacclimation and reacclimation of cold-hardy plants: current understanding and emerging concepts, Plant Science 171:3-16 Wolf TK, Cook MK (1992) Seasonal deacclimation patterns of three grape cultivars at constant warm temperature, American Journal of Enology and Viticulture 43:171-179 Ferguson JC et al. (2011) Dynamic thermal profiles of grapevine cold hardiness, American Journal of Enology and Viticulture 62:407-413
Hardiness Retention Disposition
A 35 cm segment of dormant 110 Richter rootstock cane harvested in January bears a Hardwood_Cutting_Role, bundled in quantities of 200, treated with fungicide, stored at 2°C and 95% RH until March, then prepared for benchgrafting by trimming to standard length (28-30 cm) and disbudding all but the basal node. A hardwood cutting of Riparia Gloire serves in a Hardwood_Cutting_Role for direct field rooting, planted in nursery rows with 2-3 nodes above soil level after callusing treatment to initiate root primordia. A hardwood cutting of Thompson Seedless bears a Hardwood_Cutting_Role in own-rooted propagation for table grape production in regions without phylloxera pressure, rooted in containers or nursery beds for first-year field establishment.
A cutting role that inheres in a stem segment harvested from fully matured, dormant (lignified) cane tissue, typically collected during winter dormancy (December-February in the Northern Hemisphere) when carbohydrate reserves are maximized and buds are fully acclimated to cold. Hardwood cuttings represent the standard commercial propagation format for grapevine rootstocks and the rootstock component of benchgrafts, exploiting the natural dormancy period when canes can be harvested, stored under refrigeration (0-4°C, 95%+ relative humidity) for extended periods (up to 4-6 months), and subsequently rooted when conditions permit. Successful hardwood cutting propagation requires adequate cane maturity (lignification of periderm, brown bark coloration), appropriate diameter (typically 7-12 mm for standard cuttings), sufficient carbohydrate reserves (starch content >8-10% dry weight in the basal internode), healthy bud condition (freedom from freezing injury, mechanical damage, or pathogen infection), and absence of trunk disease pathogens (Phaeomoniella, Phaeoacremonium, Botryosphaeriaceae) that can colonize cutting tissue during storage and establishment.
Hardwood_Cutting_Role captures the dominant commercial propagation format, where dormant tissue collection enables efficient harvest, extended storage, and flexible timing of propagation activities. Hardwood cutting success depends on the interaction of genotype (inherent rooting capacity), cutting quality (diameter, maturity, reserves), storage conditions (preventing desiccation and pathogen development), and rooting environment (temperature, moisture, auxin treatment for difficult-rooting genotypes). The hot water treatment (50°C for 30-45 minutes) increasingly applied to hardwood cuttings addresses trunk disease pathogens but can reduce rooting vigor if improperly applied. Cutting length standards vary by use—longer cuttings (40-50 cm) for field rooting where deeper planting is desired, standard lengths (28-32 cm) for benchgrafting, and shorter lengths for container production or where source material is limited.
Hartholzsteckling
Rebholzsteckling
Wintersteckling
dormant cutting
hardwood stem cutting
lignified cutting
verholzter Steckling
winter cutting
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 344-360 Waite H et al. (2015) Hot water treatment of grapevine cuttings, Phytopathologia Mediterranea 54: 325-339 Gramaje D, Armengol J (2011) Fungal trunk pathogens, Phytopathologia Mediterranea 50: S217-S226 Whiting JR (2004) Grapevine rootstocks, Winetitles
Hardwood Cutting Role
In Chardonnay berries in Champagne, France at BBCH 89 harvest, harvest acidity level measures titratable acidity of 10.8 g/L and pH 2.98, with malic acid at 7.2 g/L and tartaric acid at 6.5 g/L, representing the high-acid profile required for sparkling wine base material production, documented through sampling of 200 berries from 20 vines across five vineyard blocks with juice extraction at 65% yield for standardized analysis.
Harvest acidity level denotes the specific titratable acidity and pH values measured in Vitis vinifera berries at the point of commercial harvest, representing the definitive acid composition that will enter the winemaking process and establishing baseline parameters for subsequent vinification decisions including acidification requirements, malolactic fermentation management, and sulfur dioxide additions during wine production. This metric encompasses titratable acidity values typically ranging from 5-10 g/L expressed as tartaric acid equivalents and pH values of 3.1-3.8, with the specific targets varying according to wine style objectives, regional regulations, and cultivar characteristics that define acceptable ranges for quality wine production in diverse viticultural contexts. The determination of harvest acidity level requires standardized analytical methodology including sample collection protocols specifying berry number, cluster position, and vineyard zone representation, juice extraction procedures ensuring consistent yield and minimal oxidation, and titrimetric analysis using sodium hydroxide titration to phenolphthalein endpoint or potentiometric endpoint detection for accurate quantification. Temperature conditions during the final ripening days significantly influence harvest acidity level through continued effects on malate degradation rates, with warm spells accelerating acid loss by 0.2-0.4 g/L per day compared to cool conditions maintaining near-equilibrium levels, necessitating frequent sampling during the pre-harvest period to capture optimal picking windows. The relationship between harvest acidity level and wine quality reflects the fundamental role of acidity in wine sensory balance, microbial stability, and aging potential, with insufficient acidity producing flat, unstable wines susceptible to spoilage while excessive acidity creates harsh, unbalanced wines requiring amelioration through deacidification or extended malolactic fermentation. Regional and regulatory considerations influence harvest acidity level targets, with cool-climate regions such as Champagne, Mosel, and Tasmania accepting or requiring higher acidity levels of 8-12 g/L for their characteristic wine styles compared to warm-climate regions such as Barossa, Napa, and Mendoza where lower levels of 5-7 g/L are typical and acceptable. Documentation of harvest acidity level provides essential traceability information for quality assurance programs, enabling correlation of vineyard conditions with wine quality outcomes and informing future viticultural decisions regarding harvest timing optimization for specific production objectives.
Harvest_Acidity_Level is subclass of Final_Acid_Balance representing the specific measurement at picking. Distinguishes from Mature_Berry_Acidity which describes the biological state. Sibling to Mature_Berry_Acidity and Sugar_Acid_Ratio_Optimization. Key parameters are titratable acidity 5-10 g/L and pH 3.1-3.8. Critical for winemaking planning and quality documentation. Methodology-dependent requiring standardized protocols.
Ernte-Acidität
Ernte-Säure-Messung
Lese-Säuregehalt
Weinlese-Aciditäts-Level
harvest acid measurement
harvest titratable acidity
picking acidity
vintage acidity level
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 278-285 OIV (2021) Compendium of International Methods of Wine and Must Analysis Iland P et al (2013) Chemical Analysis of Grapes and Wine Techniques and Concepts 2nd Ed
Harvest Acidity Level
Harvest condition quality comparison in Alsace evaluated the sensitivity of aromatic white varieties to harvest timing and temperature conditions. Gewürztraminer demonstrated stringent harvest condition quality requirements with significant terpene degradation when fruit temperature exceeded 20°C during harvest and transport, necessitating early morning picking completing by 10:00 AM and rapid transport in refrigerated containers to preserve the varietal aromatic character critical to wine quality. Riesling showed moderate harvest condition quality with temperature sensitivity apparent above 25°C but tolerating standard mid-morning harvest under typical autumn conditions without specialized cooling requirements. Pinot Gris exhibited favorable harvest condition quality with robust aromatic stability and low oxidation sensitivity permitting flexible harvest scheduling across the full workday without detectable quality impact, though with recommendation for cool conditions when Botrytis-affected fruit required careful handling to prevent juice release and oxidation. The condition quality hierarchy directly informed harvest logistics, with Gewürztraminer receiving priority scheduling during optimal morning windows while Pinot Gris accommodated afternoon picking when labor availability required extended harvest days.
Harvest condition quality is a viticultural quality that inheres in a grapevine cultivar or clone and represents the sensitivity of fruit quality to the environmental and operational conditions prevailing during the harvest operation itself, encompassing the tolerance for weather variation, temperature conditions, and timing within the diurnal cycle that influence the physiological state of harvested fruit and its suitability for subsequent processing into wines meeting target quality specifications. This quality determines the constraints on harvest operations and the degree to which weather windows, equipment scheduling, and labor availability must be coordinated to capture fruit under conditions conducive to quality preservation, recognizing that the brief period between fruit detachment and winery processing represents a critical vulnerability during which quality can be compromised through unfavorable harvest conditions. The ontological significance of harvest condition quality lies in its role as a practical determinant of harvest logistics, wherein cultivars with stringent condition requirements demand careful coordination of weather forecasting, picking schedules, and transport logistics, while those tolerating broader condition ranges permit flexible operations accommodating the unpredictable circumstances encountered during vintage periods. The biological basis of harvest condition quality encompasses the physiological state of fruit at detachment including turgor pressure, respiratory rate, and enzyme activity levels that vary with temperature and time of day the structural integrity of berries affecting damage susceptibility during harvest and transport the composition stability of quality-determining compounds under various temperature and oxidation exposure conditions and the microbial populations on fruit surfaces that may proliferate under warm, moist harvest conditions creating spoilage risk. Cultivar characteristics influencing harvest condition quality include skin thickness affecting mechanical damage susceptibility, juice oxidation potential determining temperature sensitivity, aromatic compound volatility influencing timing requirements, and disease resistance affecting tolerance for damp harvest conditions. Assessment of harvest condition quality involves comparative evaluation of wine outcomes from fruit harvested under varied conditions, quantification of quality parameter changes during harvest and transport, and identification of condition thresholds beyond which quality compromise becomes significant.
Harvest_Condition_Quality serves as the parent class for Weather_Dependency_Quality, Morning_Harvest_Suitability_Quality, and Night_Harvest_Suitability_Quality which address specific condition dimensions affecting harvest operations. This class captures the overall concept of harvest condition sensitivity while child classes address particular environmental factors. Distinguished from Harvest_Timing_Quality which addresses when to harvest rather than the conditions under which harvest should occur. Critical for vintage logistics planning and quality preservation during harvest operations.
Erntebedingungsqualität
Erntewettertoleranz
Lesebedingungsempfindlichkeit
Weinlese-Konditionsanforderungen
harvest environment tolerance
harvest logistics constraints
picking condition sensitivity
vintage condition requirements
Jackson RS (2008) Wine Science: Principles and Applications, 3rd Edition. Academic Press, pp. 270-331 Coombe BG, Iland P (2004) Grape berry development and winegrape quality. In: Dry PR, Coombe BG (eds) Viticulture Volume 1 Resources, 2nd Edition, Winetitles, Adelaide, pp. 210-248 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1: The Microbiology of Wine and Vinifications, 2nd Edition. John Wiley and Sons, Chichester
Harvest Condition Quality
Harvest cost for hand-picked Pinot Noir in Burgundy's Côte d'Or (2023 vintage) approximately €3,000-5,000/ha — reflecting the regulatory requirement for hand harvesting in Burgundy appellations, the high labor rates in France (minimum agricultural wage plus social charges totaling approximately €15-18/hour), the relatively low yields (30-45 hL/ha generating approximately 4,000-6,000 kg/ha of grapes requiring 200-350 person-hours/ha to harvest), and the need for selective picking (multiple passes or triage in the vineyard to exclude substandard clusters), with supplementary costs for crew transport, meals, and accommodation for itinerant seasonal workers. Harvest cost for machine-harvested Grüner Veltliner in the Weinviertel (2023 vintage) approximately €400-700/ha — reflecting the efficiency of modern over-the-row mechanical harvesters capable of processing 0.5-1.0 ha/hour, operated at contracted rates of €300-500/ha or owned-equipment operating costs of €200-400/ha, with supplementary costs for transport to winery and nighttime-harvest coordination where early-morning cool-temperature harvesting is employed to preserve primary fruit aromas. Harvest cost for hand-picked Riesling on the steep terraces of the Mosel (2023 vintage) represents an extreme scenario — €5,000-8,000+/ha reflecting the impossibility of mechanization on slopes exceeding 40-60%, the physically demanding access via narrow paths and steps, the very high labor requirement per kilogram (steep slopes reduce picking speed by 40-60% compared to flat terrain), and for TBA-level selective harvest, individual berry-by-berry selection requiring 500-1,000+ person-hours per hectare.
Harvest cost is a cost of production entity that documents the specific expenditures associated with the grape harvesting operation, encompassing the cost of the harvesting method employed (hand picking labor or machine harvesting equipment operation), harvest logistics (grape transport from vineyard to winery, sorting and processing at the winery intake), pre-harvest preparation costs (netting for bird protection, selective pre-harvest leaf removal for access, harvest crew recruitment and organization), and any harvest-method-specific supplementary costs (hand-harvest crew accommodation and meals for itinerant seasonal workers, machine harvester depreciation and maintenance, night-harvesting lighting and cooling equipment), expressed per hectare or per metric ton of grapes harvested with specification of the harvesting method, regional labor cost context, yield level, and topographic constraints, providing the final viticultural cost component that bridges the production and processing phases of the wine value chain.
Harvest cost documentation captures the most operationally intensive and time-critical phase of viticultural production — the compressed period (typically 2-6 weeks per region) during which the entire annual grape crop must be collected at optimal ripeness, creating peak labor demand, logistical complexity, and weather-dependent urgency that distinguish harvest from all other viticultural operations. The hand-versus-machine harvest decision represents the most consequential cost choice in the harvest domain — machine harvesting typically costs 60-85% less per hectare than hand harvesting (€300-700/ha versus €1,500-5,000+/ha depending on region and terrain), but the choice is constrained by multiple factors including terrain (slopes exceeding 15-20% generally preclude machine access), appellation regulations (hand harvesting is mandatory in several prestigious appellations including Champagne, Burgundy, and Beaujolais), wine style requirements (whole-cluster fermentation requires intact clusters only achievable by hand, sparkling wine base requires gentle whole-cluster pressing favoring hand-picked fruit), and cultivar characteristics (thin-skinned cultivars like Pinot Noir may suffer excessive berry damage from mechanical harvesting, though modern technology has substantially reduced this concern). The cultivar-specific dimension of harvest cost reflects both management requirements and yield levels — cultivars requiring selective harvest (multiple passes for botrytized sweet wine production, or triage to exclude damaged clusters) incur dramatically higher labor costs than cultivars harvested in a single pass, and low-yielding cultivars have higher per-kilogram harvest costs even at equivalent per-hectare rates because fewer kilograms are collected per hour of labor. The night-harvesting practice — increasingly common in warm-climate viticulture for aromatic white varieties (Sauvignon Blanc, Riesling, Grüner Veltliner) — adds the cost of lighting equipment and night-shift labor premiums but is justified by the quality benefit of processing cooler grapes with better-preserved primary aromas and reduced oxidation risk. The labor availability dimension has become increasingly critical — many traditional wine regions face chronic harvest labor shortages as agricultural seasonal work becomes less attractive to domestic workers, creating dependence on international migrant labor programs and driving continued investment in mechanical harvesting technology even in quality-oriented production systems.
Erntebetriebskosten
Erntekosten
Lesekosten
Traubenernte-Kosten
Weinlesekosten
grape collection cost
harvesting cost
picking cost
vendange cost
vintage cost
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Intrieri C, Poni S (2000) Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality of both hand and mechanically harvested grapes. American Journal of Enology and Viticulture 51(3): 296-303 LWK Burgenland (2023) Erntekosten Kalkulation Mosel Weinkulturlandschaft (2023) Kosten der Steillagenbewirtschaftung
Harvest Cost
Chardonnay for premium sparkling wine production in Champagne is harvested at firmness values of 2.8-3.5 N combined with 17-19°Brix, where the relatively higher firmness compared to still wine targets ensures whole-cluster pressing efficiency and reduced phenolic extraction, maintaining the pale color and fresh acidity profile essential for traditional method sparkling wine production.
Harvest firmness quality is a softening stage quality that inheres in a grape berry at the target maturity point for commercial harvest, representing the specific mechanical state that optimally balances physiological ripeness indicators with practical handling requirements for the intended end use. Within the BFO framework, this quality represents a defined target state rather than a phase, specifying the firmness value or range considered optimal when integrated with sugar content, acidity, phenolic maturity, and aroma development for a particular production objective. The ontological distinction of harvest firmness lies in its normative character, where the quality specification varies by intended use, from firmer targets of 3-5 N for mechanical harvest and long-distance transport of table grapes to softer targets of 1.5-3 N for premium red wine where enhanced phenolic extractability compensates for increased handling fragility. This quality integrates textural requirements with the concept of technological maturity, acknowledging that optimal harvest firmness represents a compromise between complete physiological ripeness that would further reduce firmness and practical constraints including bunch rot susceptibility, mechanical damage during harvest, and crushing behavior in the winery. Instrumental specification of harvest firmness enables objective harvest decision support complementing traditional sugar and acid measurements, with real-time texture monitoring identifying the harvest window where firmness decline rate slows, indicating approaching physiological plateau. The quality specification must account for diurnal firmness variation of 10-15% due to turgor fluctuations, standardizing measurement time relative to solar noon, and temperature effects requiring sample equilibration. Harvest firmness documentation supports quality control programs, contract specifications for grape purchase, and retrospective analysis correlating textural parameters with wine quality outcomes.
Harvest_Firmness_Quality represents a target specification rather than a developmental phase, distinguished from Post_Veraison_Firmness_Quality by its normative rather than descriptive character. This quality interfaces with economic and logistical considerations beyond pure physiological state. Specification should include intended use context (table grape, wine style, storage duration) as harvest firmness targets vary significantly. Aligns with OIV descriptor 235 specifically at commercial maturity determination.
Erntefestigkeit
Lesefestigkeit
Reifefestigkeit bei Ernte
commercial maturity firmness
optimal harvest texture
optimale Erntetextur
picking firmness
target harvest firmness
Rolle L et al. (2012) Instrumental texture analysis parameters as markers of table-grape and winegrape quality. American Journal of Enology and Viticulture 63(1): 11-20 Letaief H et al. (2008) Assessment of grape skin hardness by a puncture test. Journal of the Science of Food and Agriculture 88(9): 1567-1575
Harvest Firmness Quality
Harvest flexibility comparison among cultivars in a diversified estate reveals the operational planning implications, with Cabernet Sauvignon showing high flexibility (quality stable across 14+ day window after achieving maturity, permitting deliberate scheduling around weather and capacity), Sauvignon Blanc moderate flexibility (7-10 day window with gradual aromatic shift from herbaceous to tropical enabling style-based timing rather than urgency), and Pinot Noir low flexibility (5-7 day window with rapid quality change requiring priority scheduling and weather-responsive execution), with this flexibility profile enabling the estate to schedule Pinot Noir first when windows open, use Cabernet's flexibility as schedule buffer, and time Sauvignon Blanc for desired aromatic profile within its moderate window, demonstrating how flexibility assessment enables strategic harvest sequencing.
Harvest flexibility quality is a harvest readiness quality that represents the degree of latitude available in harvest timing without significant quality compromise, characterizing the width of the acceptable maturity window and the resilience of fruit quality to variation in picking date that together determine how much scheduling flexibility exists for operational planning and response to unexpected constraints. Within the BFO framework, this quality specifically addresses the tolerance dimension of harvest timing, distinguishing the flexibility available around any optimal point from the optimal point itself while recognizing that flexibility varies substantially among cultivars, vintages, and sites, with some situations demanding precise timing (narrow flexibility) and others permitting substantial variation (broad flexibility) without significant quality impact. The ontological significance lies in harvest flexibility determining the operational pressure and risk associated with harvest timing decisions, where high flexibility enables deliberate scheduling, recovery from weather interruptions, and allocation of limited resources across extended periods, while low flexibility creates urgency, weather vulnerability, and the potential for quality loss from timing errors or delays that force harvest outside the narrow acceptable window. The factors determining harvest flexibility include cultivar characteristics (some varieties like Cabernet Sauvignon holding condition for weeks after achieving maturity, others like Pinot Noir or aromatic whites deteriorating rapidly), weather conditions (stable, cool weather extending flexibility while heat accelerates over-ripening and rain threatens dilution and disease), fruit condition (healthy fruit tolerating extended hang time better than compromised fruit), and wine style tolerance (some styles accepting wider maturity ranges than others), with these factors interacting to create vintage-specific flexibility that may differ substantially from historical patterns. The operational value of harvest flexibility includes the ability to sequence harvest across multiple blocks without compromising any (when all have adequate flexibility), the reduced weather risk from being able to wait for favorable conditions (when flexibility exceeds weather uncertainty), and the tolerance for unexpected delays (equipment failure, labor shortage) without quality catastrophe, with these benefits making flexibility assessment as important as optimal timing identification for harvest planning. The strategic management of harvest flexibility includes practices that extend flexibility (maintaining canopy health for continued berry support, avoiding over-cropping that accelerates deterioration) and recognition of when flexibility is exhausted (implementing emergency harvest protocols when remaining flexibility cannot accommodate forecast risks). Understanding harvest flexibility quality enables realistic assessment of scheduling options, appropriate urgency calibration in harvest decisions, and recognition of when flexibility is adequate for operational convenience versus when precision timing is essential for quality preservation.
Harvest_Flexibility_Quality addresses the tolerance for timing variation without significant quality compromise. This quality determines operational pressure and scheduling latitude available during harvest. The class supports harvest planning and risk management through flexibility characterization enabling strategic sequencing and contingency planning.
Ernteflexibilität
Leseflexibilität
Terminflexibilität
Zeittoleranz
harvest latitude
picking flexibility
scheduling flexibility
timing tolerance
Reynolds AG (2010) Managing Wine Quality, Volume 1: Viticulture and Wine Quality. Woodhead Publishing, Cambridge, pp. 111-155 Coombe BG, Dry PR (2004) Viticulture Volume 2: Practices, 2nd Ed. Winetitles, Adelaide, pp. 154-178
Harvest Flexibility Quality
Harvest logistics quality comparison in Napa Valley evaluated operational efficiency across major red Bordeaux varieties under standardized bilateral cordon VSP training. Merlot demonstrated excellent harvest logistics quality with clearly visible clusters positioned at consistent height, moderate peduncle attachment force of 280 g enabling clean cutting, loose cluster architecture facilitating handling without compaction damage, and robust berry attachment minimizing field losses to less than 2% of potential yield. Cabernet Sauvignon showed good harvest logistics quality with similar positioning advantages though slightly higher peduncle force of 340 g requiring sharper cutting tools. Petit Verdot exhibited challenging harvest logistics quality with small, tightly-packed clusters difficult to locate within dense canopy, extremely firm peduncle attachment exceeding 450 g creating cutting difficulty, and compact architecture prone to compression damage during handling, resulting in 25% lower harvest productivity and 8% higher fruit damage compared to Merlot under equivalent conditions, establishing harvest logistics quality as a significant factor in variety economics beyond yield and price considerations.
Harvest logistics quality is a viticultural quality that inheres in a grapevine cultivar or clone within a specific training system configuration and represents the ensemble of characteristics affecting the practical efficiency, labor requirements, and operational complexity of harvest operations, encompassing the ease with which clusters can be located, accessed, and removed from the vine, the physical integrity of fruit during the harvest process, and the tolerance of harvested material for handling and transport between vineyard and winery without quality-compromising damage or deterioration. This quality determines the operational efficiency of harvest and directly influences production costs, labor requirements, and the quality preservation achievable during the critical transition from vine to processing facility. The ontological significance of harvest logistics quality lies in its role as a practical determinant of vineyard profitability and quality potential, wherein cultivars exhibiting favorable logistics characteristics enable rapid, efficient harvest with minimal fruit damage at lower labor cost, while those with poor logistics characteristics create operational challenges that increase costs, extend harvest duration, and may compromise fruit quality through prolonged exposure or handling damage. The biological basis of harvest logistics quality encompasses multiple cultivar and training system characteristics including cluster positioning and visibility affecting location and access efficiency, peduncle attachment strength and cutting characteristics influencing removal ease, cluster architecture determining handling requirements and damage susceptibility, berry attachment strength and skin integrity affecting fruit loss and juice release during harvest, and compositional characteristics influencing tolerance for the temperature elevation, oxidation exposure, and physical impacts encountered during transport. The interaction between cultivar characteristics and training system configuration strongly influences harvest logistics quality, with the same cultivar exhibiting different logistics characteristics under different training arrangements that alter cluster positioning, accessibility, and exposure to operational hazards. Assessment of harvest logistics quality involves quantification of harvest rate productivity, fruit damage incidence, labor cost per unit harvested, and quality parameter changes between vineyard and winery receiving, with benchmarking against regional standards and alternative cultivar options enabling informed variety selection and training system design decisions that optimize logistics alongside agronomic and quality considerations.
Harvest_Logistics_Quality serves as the parent class for Picking_Ease_Quality, Cluster_Accessibility_Quality, Fruit_Integrity_At_Harvest_Quality, and Transport_Tolerance_Quality which address specific dimensions of harvest operational characteristics. This class captures the overall logistics concept while child classes address particular aspects. Distinguished from Mechanical_Harvesting_Suitability_Quality by addressing both manual and mechanical harvest considerations rather than specifically machine compatibility. Critical for production cost analysis and operational planning.
Ernte-Handhabungsqualität
Erntelogistikqualität
Lese-Betriebseffizienz
Weinlese-Praktikabilität
harvest operational efficiency
harvest practicality
picking logistics
vintage operations quality
Morris JR (2007) Development and commercialization of a complete vineyard mechanization system. HortTechnology 17:411-420 Intrieri C, Poni S (1995) Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality of mechanized Italian vineyards. American Journal of Enology and Viticulture 46:116-127 Jackson RS (2008) Wine Science: Principles and Applications, 3rd Edition. Academic Press, pp. 270-331
Harvest Logistics Quality
The harvest method description for Domaine Leflaive's Puligny-Montrachet Premier Cru vineyards specifies hand harvesting by a trained team of 40-50 vendangeurs over 7-10 days in mid-September, with picking decisions based on daily berry sampling at 50 random locations per parcel achieving target parameters of 12.5-13.5% potential alcohol, pH 3.15-3.30, titratable acidity 6.5-8.0 g/L, and sensory assessment of skin flavor development — grapes are collected in small 20 kg perforated plastic crates (cagettes) to minimize compression damage, transported within 30 minutes to the cuverie, and passed over a vibrating sorting table where trained sorters remove damaged berries, unripe clusters, and vegetal material before gentle whole-cluster pressing.
A viticultural practice description that documents the methods, timing criteria, logistical organization, and quality control procedures governing the collection of ripe grapes from the vineyard, encompassing the complete set of decisions and operational specifications that determine when, how, and with what degree of selectivity fruit is removed from the vine, transported to the winery, and delivered for processing.
Harvest_Method_Description documents the culmination of the entire viticultural year — the operation that converts a year's investment in vineyard management into the raw material for winemaking. The harvest decision integrates multiple information streams (berry chemistry, sensory evaluation, weather forecasts, disease pressure, labor availability, winery capacity) into the most time-critical and least reversible decision in the production cycle: harvesting too early sacrifices flavor development and phenolic maturity, while delaying harvest risks disease, rain damage, overripeness, and acidity loss. The method of harvest (hand versus mechanical) determines the selectivity, speed, cost, and physical condition of the harvested fruit, with profound implications for subsequent winemaking options. Subclasses capture the key dimensions of harvest documentation — timing criteria, method type, mechanical and manual specifications, selectivity criteria, and post-harvest sorting protocols.
Erntemethoden-Beschreibung
Lese-Dokumentation
Traubenlese-Protokoll
grape collection protocol
harvest practice documentation
vintage operation record
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 260-295 Coombe BG and Iland PG (2004) Grape berry development and winegrape quality, in Dry PR et al. (eds) Viticulture Volume 1: Resources, 2nd Ed, Winetitles pp: 210-248 Jackson RS (2020) Wine Science: Principles and Applications 5th Ed, Academic Press pp: 295-340
Harvest Method Description
The harvest method type classification for a large Languedoc cooperative distinguishes three tiers of incoming fruit — mechanically harvested parcels (75% of intake, collected by over-the-row harvesters at 2-3 ha/h and 40-60 EUR/ha, processed within 4 hours of picking for varietal and IGP wines), hand-harvested parcels designated for AOC production (20% of intake, collected at 200-400 labor hours/ha and 1,500-3,000 EUR/ha, delivered in small crates for gravity-fed processing), and hand-harvested parcels with field sorting for premium cuvées (5% of intake, requiring 400-600 labor hours/ha at 3,000-5,000 EUR/ha, delivered as pre-sorted whole clusters for whole-cluster pressing or whole-berry fermentation).
A harvest method description consisting of the classification of the grape collection method as either manual (hand harvesting), mechanical (machine harvesting), or a combination of both, specifying the fundamental operational approach that determines harvesting speed, selectivity, fruit condition at delivery, labor requirements, cost structure, and compatibility with subsequent winemaking processes.
Harvest_Method_Type defines the fundamental operational dichotomy in modern viticulture between the efficiency of mechanical harvesting and the selectivity of hand harvesting. The global proportion of mechanically harvested wine grapes has increased steadily from approximately 5% in 1980 to an estimated 60-70% of world production by 2020, driven by labor scarcity, cost reduction, and technological improvements in harvester gentleness and selectivity: however, hand harvesting remains mandatory or strongly preferred for premium wines, sparkling wine production (where whole-cluster pressing requires intact clusters), steep-slope viticulture (inaccessible to machines), and appellations that mandate manual harvest (e.g., Champagne AOC, Beaujolais Cru, most Burgundy AOCs). The choice between methods is not purely quality-driven — mechanical harvesting enables rapid collection of large volumes during optimal maturity windows, reducing the risk of quality loss from weather-related harvest delays that can affect hand-harvested operations requiring 10-20 times longer to complete the same area.
Erntemethoden-Typ
Leseart-Klassifikation
Traubensammlungs-Methode
grape collection approach
harvesting method classification
picking method type
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 280-295 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 230-260 Clary CD et al. (1990) Performance of destemmer-crusher and crusher-destemmer machines with and without a berry sorting and cleaning system, American Journal of Enology and Viticulture 41: 71-75
Harvest Method Type
Harvest readiness assessment for Chardonnay under approaching weather front reveals the integrated decision-making, with compositional status showing current maturity acceptable (23.2 Brix, pH 3.38, balanced) but still improving (0.15 Brix/day gain, approaching optimal 24.0 Brix in 5 days), while weather forecast shows 80% precipitation probability in 4 days with rain continuing 3+ days (potential botrytis risk, dilution, difficult picking conditions), and operational assessment shows full picking capacity available immediately, resulting in harvest decision to pick now despite incomplete maturity optimization, as the compositional benefit of additional hang time is outweighed by weather risk, demonstrating how harvest readiness integrates maturity and operational factors for practical decision-making.
Harvest readiness quality is a maturity quality that represents the overall suitability of grape fruit for harvest considering the integration of maturity parameters, logistical factors, weather forecasts, and winemaking capacity, characterizing the holistic assessment of whether the time is right for picking that extends beyond compositional maturity alone to encompass the practical realities of harvest execution. Within the BFO framework, this quality addresses the operational dimension of harvest timing, distinguishing readiness for the actual harvest operation from the achievement of compositional targets while recognizing that optimal harvest timing balances grape maturity against practical constraints including weather risk, labor availability, winery capacity, and the prioritization among multiple blocks reaching maturity simultaneously. The ontological significance lies in harvest readiness determining when fruit is actually picked, which may differ from when it achieves any single maturity optimum due to the practical necessity of balancing ideal maturity against real-world constraints that may preclude waiting for or acting upon compositional perfection. The factors integrated in harvest readiness assessment include compositional maturity status (sugar, acid, phenolic, aromatic parameters relative to targets), maturity trajectory (rate of change indicating whether waiting will improve or degrade quality), weather conditions and forecasts (current conditions for harvest execution, precipitation and temperature forecasts affecting fruit condition), fruit condition (disease pressure, berry integrity, potential for deterioration), and operational constraints (picking crew availability, transport logistics, winery receiving capacity, competing harvest priorities), with these factors requiring simultaneous consideration rather than sequential optimization. The decision-making framework for harvest readiness involves threshold-based triggering (minimum maturity criteria that must be met), window identification (the period during which acceptable maturity coincides with favorable conditions), risk assessment (probability of improvement versus deterioration with additional hang time), and resource allocation (prioritization when multiple blocks compete for limited harvest capacity). The vintage-specific nature of harvest readiness reflects the variable weather patterns that may create narrow windows of opportunity (compressed harvest under rain threat), extended flexibility (favorable weather enabling ideal timing), or forced compromises (unavoidable weather events requiring suboptimal timing), with successful harvest timing requiring adaptive decision-making that responds to evolving conditions rather than rigid adherence to predetermined dates or parameters. Understanding harvest readiness quality enables integration of compositional and operational factors in harvest decisions, appropriate weighting of maturity achievement against practical constraints, and recognition of the adaptive decision-making required for optimal harvest execution.
Harvest_Readiness_Quality serves as the parent class for harvest window and flexibility qualities. This quality addresses the integration of compositional, operational, and environmental factors in harvest decisions. The class supports practical harvest timing through readiness characterization beyond compositional maturity alone.
Ernteentscheidungsqualität
Erntereife
Lesereife
harvest decision quality
harvest suitability
operational maturity
operative Reife
picking readiness
Coombe BG, Dry PR (2004) Viticulture Volume 2: Practices, 2nd Ed. Winetitles, Adelaide, pp. 154-178 Jackson RS (2014) Wine Science: Principles and Applications, 4th Ed. Academic Press, San Diego, pp. 232-298
Harvest Readiness Quality
The harvest timing criteria for Cabernet Sauvignon in the Médoc specify a multi-parameter maturity assessment conducted by daily sampling from véraison onward — sugar accumulation tracked by refractometry targeting 13.0-13.5% potential alcohol (230-240 g/L sugar), acid balance monitored by titration targeting titratable acidity of 5.0-6.0 g/L and pH of 3.45-3.60, phenolic maturity assessed by the Glories method (extractable anthocyanins at pH 1.0 and pH 3.2 with a maturity index below 40% indicating adequate skin cell wall degradation for tannin extraction), and seed tannin maturity assessed by visual inspection (seeds transitioning from green to brown indicating lignification and reduced harsh tannin contribution) — the harvest decision is triggered when all parameters simultaneously fall within their target ranges, tempered by 5-day weather forecasts that may accelerate picking if rain threatens or permit extended hang-time if dry conditions prevail.
A harvest method description that documents the analytical, sensory, and environmental parameters used to determine the optimal harvest date for a specific vineyard block, specifying the target values for sugar concentration, titratable acidity, pH, phenolic maturity indicators, aromatic development, and weather considerations that collectively define the maturity window within which harvest must occur to achieve the intended wine style.
Harvest_Timing_Criteria captures the most complex multi-parameter decision in viticulture — the determination of optimal harvest maturity for a specific cultivar, site, vintage, and target wine style. The historical reliance on sugar concentration as the primary (or sole) maturity indicator has been progressively supplemented by a more holistic maturity assessment framework encompassing technological maturity (sugar, acid, pH), phenolic maturity (anthocyanin extractability, tannin polymerization, seed lignification), and aromatic maturity (varietal aroma precursor accumulation, methoxypyrazine degradation): these three maturity dimensions do not necessarily converge simultaneously — in warm vintages sugar accumulation may outpace phenolic and aromatic maturity, while in cool vintages phenolic development may lag behind sugar, creating the fundamental harvest timing dilemma of which maturity parameter to prioritize. The concept of phenolic maturity assessment, pioneered by Glories and Augustin (1993) and refined by Ribéreau-Gayon et al. (2006), has become standard practice in premium red wine production but remains absent from many regulatory maturity definitions which still specify only minimum sugar levels.
Ernteentscheidungs-Parameter
Lesezeitpunkt-Kriterien
Reifebewertungs-Protokoll
harvest date criteria
maturity assessment protocol
picking decision parameters
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1: The Microbiology of Wine and Vinifications, 2nd Ed, Wiley pp: 235-270 Coombe BG and Iland PG (2004) Grape berry development and winegrape quality, in Dry PR et al. (eds) Viticulture Volume 1: Resources, 2nd Ed, Winetitles pp: 210-248 Glories Y and Augustin M (1993) Maturité phénolique du raisin, conséquences technologiques: applications aux millésimes 1991 et 1992, Actes du Colloque Journée Technique du CIVB pp: 56-61
Harvest Timing Criteria
The harvest timing description of Riesling documents late harvest (OIV 304:7) — reaching optimal wine harvest maturity (19-22°Brix, TA 7-10 g/L, pH 3.0-3.2) approximately 10-15 days after Chasselas in Alsace and the Mosel, typically during the second to fourth week of October (October 10-30 at Bernkastel, Mosel, 49.9°N). This late harvest timing positions Riesling at the edge of the growing season in its traditional northern European habitats, where the final weeks of ripening in October occur under declining temperatures and shortening daylengths that slow sugar accumulation but preserve the high acidity that is central to Riesling's wine identity — the cultivar's capacity to achieve flavor ripeness at lower sugar levels than most other premium cultivars (harvested at 19-22°Brix versus 24-26°Brix for Cabernet Sauvignon or Syrah) is the physiological adaptation that enables quality wine production from these cool, late-season harvest conditions.
A phenological timing description component that records the typical date range at which the cultivar achieves harvest maturity — the compositional targets (sugar concentration, titratable acidity, pH, phenolic ripeness) appropriate for the intended end use — scored according to OIV descriptor 304 on a scale from very early (state 1) to very late (state 9) relative to Chasselas, with harvest timing being the ultimate expression of cultivar precocity that integrates all preceding phenological stages and determines the cultivar's compatibility with regional climate, the wine style achievable in a given vintage, and the viticultural calendar from which all management operations are backward-scheduled.
Harvest_Timing_Description documents the phenological endpoint that is the most commercially consequential timing character — harvest date determines wine style (early harvest producing lighter, more acidic wines later harvest producing riper, fuller wines), alcohol potential (directly proportional to sugar at harvest), acid balance (inversely related to harvest date as malic acid continues declining during ripening), and the logistical scheduling of the entire winemaking operation. Unlike the preceding phenological events (budbreak, flowering, véraison), which are defined by specific visible developmental markers, harvest maturity is defined by compositional targets that vary with the intended wine style and regional tradition — a sugar-based definition (°Brix or °Oechsle), an acidity-based definition (TA and pH), or increasingly a phenolic-maturity-based definition (anthocyanin extractability, seed tannin maturity, methoxypyrazine degradation) — and the same cultivar at the same site may be harvested over a 2-4 week range depending on which compositional target is prioritized: this flexibility in harvest date definition complicates the use of harvest timing as a fixed cultivar descriptor, and the OIV system addresses this by defining harvest timing relative to the Chasselas reference at the same site under the same harvest criteria, preserving the relative ranking. The véraison-to-harvest interval varies from 35-40 days for very early cultivars (Chasselas, Madeline Angevine) to 55-65 days for very late cultivars (Mourvèdre, Carignan), with this interval primarily determined by the rate of sugar accumulation per unit of thermal time — a cultivar-specific parameter reflecting the efficiency of phloem sugar transport to the berry and the berry's capacity to accumulate and compartmentalize imported sugars.
Erntezeitpunkt-Beschreibung
Lesezeit-Dokumentation
Reifezeitpunkt-Charakterisierung
harvest date characterization
maturity timing record
ripeness calendar documentation
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Jones GV and Davis RE (2000) Climate influences on grapevine phenology, grape composition, and wine production and quality for Bordeaux, France, American Journal of Enology and Viticulture 51: 249-261 Duchêne E and Schneider C (2005) Grapevine and climatic changes, Agronomy for Sustainable Development 25: 93-99 Pulliat V (1888) Mille variétés de vignes, 4th edition, Montpellier Gladstones J (1992) Viticulture and Environment, Winetitles
Harvest Timing Description
Harvest timing indicators quality assessment for Cabernet Sauvignon in Margaret River evaluated the predictive value of standard and advanced ripeness parameters for wine quality outcomes across five vintages. Sugar concentration (Brix) showed moderate harvest timing indicators quality with r²=0.62 correlation to wine quality scores, adequate for distinguishing major quality tiers but insufficient for fine-scale harvest optimization. Phenolic ripeness assessed by seed color (percentage brown seeds) demonstrated improved indicator quality with r²=0.78 correlation, providing better prediction of tannin quality and wine structure scores. The combination of Brix, seed color, and sensory berry assessment achieved excellent harvest timing indicators quality with r²=0.91 correlation to wine outcomes, establishing the multi-parameter protocol as the regional standard for Cabernet Sauvignon harvest decisions. The analysis further revealed that optimal indicator relationships were cultivar-specific, with Chardonnay showing strongest correlation to titratable acidity while Shiraz correlated most strongly with Brix, necessitating cultivar-calibrated indicator protocols.
Harvest timing indicators quality is a harvest timing quality that inheres in a grapevine cultivar or clone within a specific growing environment and represents the reliability and utility of the various parameters measured to assess ripening progress and determine optimal harvest date, encompassing the consistency of relationships between measurable indicators and wine quality outcomes, the precision of assessment methodologies, and the predictive value of indicator measurements for guiding harvest decisions that achieve target wine style objectives. This quality determines the confidence with which harvest timing decisions can be made based on available monitoring data, recognizing that effective harvest management requires indicators that accurately reflect the ripening status relevant to wine quality rather than merely convenient parameters that may be poorly correlated with ultimate wine outcomes. The ontological significance of harvest timing indicators quality lies in its role enabling evidence-based harvest decisions, wherein strong indicator quality permits confident determination of optimal harvest date from objective measurements while poor indicator quality forces reliance on experience, intuition, and potentially suboptimal timing that compromises quality achievement. The biological basis of harvest timing indicators quality encompasses the strength of correlations between measured parameters and wine quality attributes, the sampling and analytical precision achievable for specific indicators, the lead time provided by predictive indicators enabling advance scheduling, and the cultivar-specificity of indicator-quality relationships that may require calibration for different varieties or growing conditions. Traditional indicators including sugar concentration (Brix, Baumé, Oechsle), titratable acidity, and pH provide readily measured parameters with established relationships to fermentation outcomes, while phenolic ripeness assessment, aromatic compound analysis, and seed maturity evaluation offer more direct but technically challenging measures of quality-determining components. The integration of multiple indicators often provides more reliable harvest guidance than any single parameter, with indicator suites calibrated for specific cultivar-style combinations establishing the monitoring protocols supporting confident harvest decisions.
Harvest_Timing_Indicators_Quality is_a Harvest_Timing_Quality, serving as parent class for Berry_Sampling_Reliability_Quality, Maturity_Monitoring_Precision_Quality, and Vintage_Predictability_Quality which address specific dimensions of indicator utility. This class captures the overall concept of indicator reliability while child classes address particular assessment aspects. The cultivar-specificity of optimal indicator combinations represents important consideration for protocol development. Multi-parameter approaches typically outperform single-indicator assessment.
Erntezeitpunkt-Indikatoren-Qualität
Lesezeitbestimmungs-Zuverlässigkeit
Reifegradparameter
Reifeindikatoren
harvest decision criteria
maturity assessment parameters
ripeness indicators
vintage monitoring reliability
Iland P et al. (2011) Chemical Analysis of Grapes and Wine: Techniques and Concepts, 2nd Edition. Patrick Iland Wine Promotions, Adelaide Ristic R, Iland P (2005) Relationships between seed and berry development of Vitis vinifera L. cv. Shiraz: developmental changes in seed morphology and phenolic composition. Australian Journal of Grape and Wine Research 11:43-58 Zoecklein BW et al. (1998) Evaluation of the phenol-free glycosyl-glucose determination. American Journal of Enology and Viticulture 49:183-187
Harvest Timing Indicators Quality
Harvest timing quality comparison for Riesling across German wine regions demonstrated systematic differences in harvest management requirements. Mosel sites exhibited favorable harvest timing quality with slow, predictable ripening progression of 0.15°Brix per day during the final maturation phase, extended harvest windows averaging 18 days at target quality specifications, and excellent correlation (r²=0.91) between must weight and wine quality tier classification enabling confident harvest decisions. Rheinhessen sites showed more challenging harvest timing quality with faster ripening at 0.25°Brix per day creating compressed 10-day harvest windows, greater vintage variation in optimal harvest date ranging ±12 days across years, and reduced predictability requiring more intensive monitoring. Pfalz irrigated sites demonstrated problematic harvest timing quality with rapid sugar accumulation uncoupled from flavor development, requiring extended hangtime that risked overripeness and Botrytis pressure to achieve acceptable flavor maturity, establishing that harvest timing quality assessment should precede site selection for quality-focused Riesling production.
Harvest timing quality is a viticultural quality that inheres in a grapevine cultivar or clone within a specific growing environment and represents the characteristics governing the determination and execution of optimal harvest date, encompassing the predictability of ripening progression, the duration and flexibility of the harvest window during which fruit meets quality specifications, and the reliability of maturity indicators used to guide harvest decisions for achieving target wine style objectives. This quality fundamentally influences the practical management of vintage operations and the consistency of fruit quality delivered to the winery, recognizing that successful harvest timing requires both favorable cultivar-site characteristics creating manageable harvest windows and effective monitoring approaches enabling accurate assessment of ripening status. The ontological significance of harvest timing quality lies in its role as the ultimate integration point for all preceding viticultural decisions, wherein the cumulative effects of site selection, cultivar choice, rootstock matching, training system implementation, and seasonal management converge in the critical harvest timing decision that determines whether the quality potential developed throughout the growing season is captured or compromised through premature or delayed picking. The biological basis of harvest timing quality encompasses the rate and pattern of ripening progression determining how rapidly fruit transitions through target maturity zones, the stability of ripened fruit on the vine affecting the consequences of harvest timing variation, the uniformity of ripening across clusters and vineyard blocks influencing whether single-pass or selective harvest approaches prove necessary, and the correlation between measurable parameters and sensory quality attributes enabling reliable prediction of wine outcomes from pre-harvest fruit assessment. Cultivar-environment combinations exhibiting favorable harvest timing quality demonstrate predictable ripening trajectories, extended harvest windows permitting scheduling flexibility, stable fruit quality during the harvest period, and strong relationships between measured maturity indices and resulting wine characteristics. Assessment of harvest timing quality requires multi-vintage evaluation capturing the range of seasonal conditions affecting ripening patterns, with attention to both the central tendency and variance of harvest timing parameters across years representing the typical and extreme conditions that must be accommodated in production planning.
Harvest_Timing_Quality serves as the parent class for Harvest_Window_Quality and Harvest_Timing_Indicators_Quality hierarchies addressing the temporal characteristics and assessment methodologies for harvest decisions respectively. This class captures the overall concept of harvest timing management while child classes address specific dimensions. Distinguished from Maturity_Quality which addresses the developmental state of fruit rather than the timing considerations for its harvest. Critical for vintage planning and quality consistency across seasons.
Ernteterminbestimmung
Erntezeitpunktqualität
Lesezeitung-Eigenschaften
Weinleseplanung
harvest date determination
harvest scheduling characteristics
picking date quality
vintage timing
Coombe BG, Iland P (2004) Grape berry development and winegrape quality. In: Dry PR, Coombe BG (eds) Viticulture Volume 1 Resources, 2nd Edition, Winetitles, Adelaide, pp. 210-248 Jackson RS (2008) Wine Science: Principles and Applications, 3rd Edition. Academic Press, pp. 270-331 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 509-547
Harvest Timing Quality
Harvest window concentration assessment comparing estate portfolios reveals the strategic effect of cultivar mix on operational pressure, with single-cultivar estate (100% Chardonnay) showing high concentration (all blocks reaching optimal maturity within 8-day window, requiring peak harvest rate of 15 tonnes/day to capture quality), while diversified estate (Chardonnay, Pinot Noir, Syrah, Cabernet Sauvignon) shows distributed windows (optimal maturity spread across 35 days, steady harvest rate of 6 tonnes/day sufficient), demonstrating how harvest window concentration affects the capacity requirements and scheduling flexibility available during vintage with implications for both operational logistics and quality risk from harvest timing constraints.
Harvest window concentration quality is a temporal spread quality that represents the degree to which optimal harvest timing is concentrated into a narrow period versus spread across an extended window, characterizing the temporal compression of harvest opportunity that determines the operational pressure on harvest logistics, the scheduling flexibility available for weather response, and the risk of quality loss from inability to harvest within narrow optimal windows. Within the BFO framework, this quality specifically addresses the harvest timing dimension of temporal spread, distinguishing the concentration of optimal harvest opportunity from the overall developmental spread while recognizing that harvest window concentration emerges from the interaction of ripening rate with maturity spread—concentrated windows arising from rapid ripening with narrow spread, extended windows from slow ripening or wide spread that maintains acceptable quality across extended periods. The ontological significance lies in harvest window concentration determining the operational demands of vintage, where highly concentrated windows (multiple blocks reaching optimal maturity simultaneously) create harvest bottlenecks requiring maximum capacity deployment over brief periods, while distributed windows enable more even resource utilization but may reduce quality by forcing some blocks to wait beyond or harvest before their optimal timing. The factors affecting harvest window concentration include cultivar mix (diverse cultivar portfolios spreading harvest over weeks, single-cultivar estates concentrating harvest), site variation (different sites reaching maturity at different times, spreading the estate-wide window), vintage weather patterns (consistent conditions promoting concentrated maturity, variable conditions creating spread), and ripening rate characteristics (fast-ripening cultivars creating compressed windows, slow-ripening creating extended windows). The strategic management of harvest window concentration involves deliberate portfolio design (cultivar and site selection to distribute harvest timing), tactical weather response (exploiting harvest flexibility when windows overlap with favorable weather), and capacity planning (ensuring harvest resources match the concentration of peak demand periods), with these approaches representing different strategies for managing the operational challenges of concentrated versus distributed harvest. Understanding harvest window concentration quality enables realistic harvest planning based on expected timing patterns, appropriate capacity preparation for peak harvest periods, and recognition of the vintage-specific factors that may compress or extend harvest windows affecting operational pressure and quality risk.
Harvest_Window_Concentration_Quality addresses the temporal compression of optimal harvest opportunity affecting operational demands. This quality determines harvest logistics requirements and quality risk from timing constraints. The class supports harvest planning and capacity management through window concentration characterization.
Erntefensterkonzentration
Erntezeitkommpression
Lesedruck
harvest concentration
harvest timing compression
optimal window duration
optimale Fensterdauer
vintage pressure
Coombe BG, Dry PR (2004) Viticulture Volume 2: Practices, 2nd Ed. Winetitles, Adelaide, pp. 154-178 Reynolds AG (2010) Managing Wine Quality, Volume 1: Viticulture and Wine Quality. Woodhead Publishing, Cambridge, pp. 111-155
Harvest Window Concentration Quality
Harvest window duration quality comparison across red Bordeaux varieties in Saint-Émilion quantified the scheduling flexibility available for sequential harvest operations in diversified plantings. Merlot demonstrated limited harvest window duration quality with rapid post-maturity evolution, showing 0.3°Brix daily increase, 0.5 g/L daily TA decline, and Botrytis progression from 5% to 25% incidence within 10 days of optimal date, establishing an effective harvest window duration of 7-10 days before quality compromise became significant for premium wine production. Cabernet Franc exhibited intermediate duration quality with 12-15 day windows attributed to thicker skin and looser cluster architecture reducing Botrytis susceptibility. Cabernet Sauvignon showed extended harvest window duration quality of 18-21 days with slow compositional change and excellent disease resistance, providing substantial scheduling flexibility that supports its position as the final-harvested variety permitting earlier cultivars to receive priority attention. This duration quality hierarchy directly informed harvest scheduling protocols prioritizing Merlot picking to capture its narrow window while Cabernet Sauvignon accumulated additional hangtime within its extended duration.
Harvest window duration quality is a harvest window quality that inheres in a grapevine cultivar or clone within a specific growing environment and represents the length of time during which fruit remains within acceptable quality specifications after achieving initial target ripeness, measured as the number of days between optimal harvest date and the onset of quality-limiting deterioration that would compromise wine production objectives for the intended style and market tier. This quality directly determines the scheduling flexibility available for harvest operations and the risk exposure from delays caused by weather events, equipment limitations, or competing harvest priorities across multiple vineyard blocks requiring sequential picking. The ontological significance of harvest window duration quality lies in its role as a primary risk management parameter, wherein extended durations provide buffer capacity absorbing operational disruptions without quality penalty while short durations create vulnerable periods where brief delays can result in significant quality loss and economic damage. The biological basis of harvest window duration encompasses the rate of continued sugar accumulation that may push fruit beyond target Brix specifications, the pace of acid decline affecting wine balance and freshness, the stability of aromatic compounds under extended hangtime, the onset and progression of berry dehydration concentrating solutes beyond desired levels, and the development of pathogen infections particularly Botrytis bunch rot that can rapidly compromise fruit quality. Environmental conditions during the harvest window strongly influence duration, with moderate temperatures, low humidity, and absence of precipitation extending durations through slowed metabolic rates and reduced pathogen pressure, while warm, humid conditions compress durations through accelerated ripening and disease progression. Cultivar characteristics affecting harvest window duration quality include the inherent rate of post-maturity compositional change, resistance to berry shrivel and dehydration, and susceptibility to harvest-period pathogens, with thin-skinned, compact-clustered varieties typically exhibiting shorter durations than thick-skinned, loose-clustered types under equivalent environmental conditions.
Harvest_Window_Duration_Quality is_a Harvest_Window_Quality, representing the length of the acceptable harvest period. This class addresses the persistence of target quality after optimal date is reached. The rate of compositional change, disease progression, and environmental conditions determine duration. Distinguished from Optimal_Harvest_Date_Quality which addresses when the window opens rather than how long it persists. Critical for multi-block harvest scheduling and risk management under weather uncertainty.
Erntefensterdauer
Erntezeitspannenausdehnung
Lesezeitraumlänge
Weinlesefenster-Länge
harvest flexibility period
harvest window length
picking period duration
vintage window extent
Coombe BG, Iland P (2004) Grape berry development and winegrape quality. In: Dry PR, Coombe BG (eds) Viticulture Volume 1 Resources, 2nd Edition, Winetitles, Adelaide, pp. 210-248 Bindon KA et al. (2014) Relationships between harvest time and wine composition in Vitis vinifera L. cv. Cabernet Sauvignon. Food Chemistry 154:43-54 Rolle L et al. (2012) Influence of grape density and harvest date on changes in phenolic composition, phenol extractability indices, and instrumental texture properties during ripening. Journal of Agricultural and Food Chemistry 60:7571-7581
Harvest Window Duration Quality
Harvest window quality evaluation for Chardonnay across California appellations revealed substantial regional differences affecting production planning. Carneros demonstrated excellent harvest window quality with cool maritime influence slowing ripening to create 21-day average harvest windows during which fruit maintained target specifications of 23-24.5°Brix with balanced acidity and optimal flavor development. Russian River showed good harvest window quality with 16-day average windows though with greater vintage variation from 12-22 days depending on heat accumulation patterns. Central Valley exhibited challenging harvest window quality with accelerated ripening compressing windows to 8-10 days, with rapid acid decline and flavor coarsening beyond optimal harvest date requiring precise timing and substantial picking capacity to capture target quality, establishing that harvest window quality represents a critical site selection criterion for premium Chardonnay production where vintage consistency is valued.
Harvest window quality is a harvest timing quality that inheres in a grapevine cultivar or clone within a specific growing environment and represents the temporal characteristics of the period during which fruit meets specifications for target wine style production, encompassing the predictability of when this window opens, the duration for which acceptable quality persists, and the flexibility available for scheduling harvest operations within logistical constraints without compromising wine quality objectives. This quality determines the practical manageability of harvest operations, with extended windows providing scheduling flexibility accommodating weather events, equipment availability, and winery capacity limitations, while compressed windows create intense operational pressure requiring precise timing and substantial resource mobilization within brief periods. The ontological significance of harvest window quality lies in its role as a primary determinant of vintage risk and operational complexity, wherein narrow windows concentrate harvest logistics into critical periods where delays from weather, equipment failure, or labor shortages can result in quality loss, while extended windows distribute risk across longer periods enabling recovery from operational disruptions. The biological basis of harvest window quality encompasses the rate of compositional change during the ripening plateau when fruit remains within specification, the stability of desirable compounds under continued hangtime, the onset rate of quality-limiting factors including overripeness, berry shrivel, pathogen development, and bird or insect damage, and the uniformity of maturation affecting whether the window applies to entire blocks or only to selected portions requiring multiple harvest passes. The interaction between cultivar characteristics and environmental conditions strongly influences harvest window quality, with cool climate conditions generally extending windows through slowed metabolic rates while warm conditions compress windows through accelerated ripening and increased risk of rapid quality decline. Assessment of harvest window quality requires systematic tracking of compositional and sensory parameters throughout the harvest-eligible period across multiple vintages, establishing both typical window characteristics and the variance encountered under extreme seasonal conditions.
Harvest_Window_Quality is_a Harvest_Timing_Quality, serving as parent class for Optimal_Harvest_Date_Quality, Harvest_Window_Duration_Quality, and Harvest_Flexibility_Quality which address specific temporal dimensions of the harvest period. This class captures the overall window concept while child classes address particular aspects. The interaction between ripening rate, quality stability, and deterioration onset determines window characteristics. Critical for production planning and risk management across vintages.
Erntefensterqualität
Erntezeitspanne
Lesezeitraum
Weinlesefenster
harvest period
optimal harvest duration
picking window
vintage window
Coombe BG, Iland P (2004) Grape berry development and winegrape quality. In: Dry PR, Coombe BG (eds) Viticulture Volume 1 Resources, 2nd Edition, Winetitles, Adelaide, pp. 210-248 Sadras VO, Moran MA (2012) Elevated temperature decouples anthocyanins and sugars in berries of Shiraz and Cabernet Franc. Australian Journal of Grape and Wine Research 18:115-122 van Leeuwen C, Darriet P (2016) The impact of climate change on viticulture and wine quality. Journal of Wine Economics 11:150-167
Harvest Window Quality
Chardonnay harvested berries hand-picked at night (berry temperature 12-15°C at harvest, ambient 10°C), transported in small bins to winery within 1.5 hours, whole-cluster pressed within 3 hours of harvest, show minimal quality changes from vine to press with sugars stable at 22%, acidity unchanged at 7.5 g/L, no oxidation, producing optimal quality white wine from excellent post-harvest handling. Cabernet Sauvignon harvested berries machine-harvested in afternoon (berry temperature 28-32°C at harvest, ambient 35°C), transported in large bins requiring 4 hours to winery due to distance, destemmed and crushed 6 hours post-harvest, show moderate quality changes including 0.15% sugar loss through respiration, some spontaneous fermentation beginning (0.3% alcohol), minor oxidation of damaged berries, requiring sulfur dioxide addition and immediate processing to prevent further degradation. Pinot Noir harvested berries hand-picked in cool morning conditions, held in refrigerated truck at 8°C for 8 hours during transport from remote vineyard, processed 10 hours post-harvest, show minimal quality loss due to cooling despite extended post-harvest period, maintaining 24% sugars, 6.8 g/L acidity, no fermentation, good color retention, demonstrating importance of temperature control for maintaining harvested berry quality during transport delays. Sauvignon Blanc harvested berries comparison= immediate processing (2 hours post-harvest) producing wine with characteristic fresh grapefruit and herbaceous aromatics (sensory score 90/100) delayed processing (16 hours at 18-22°C) showing reduced aromatic intensity, beginning oxidation notes, quality reduction (score 82/100), demonstrating white grape sensitivity to post-harvest handling with aromatic cultivars particularly vulnerable to quality loss from delayed processing. Merlot machine-harvested berries showing harvest method quality effects= machine-harvested lot containing 8% leaves and stems (MOG - material other than grapes), 12% damaged crushed berries releasing juice during transport, requiring extensive sorting and immediate processing, versus hand-harvested lot with <2% MOG, <3% damaged berries, requiring minimal sorting, with hand-harvested lot producing higher quality wine (score 88/100 versus 84/100 machine-harvested) reflecting gentler handling and less damage despite higher harvest cost.
A harvested berry is a mature berry that has been removed from the vine through mechanical or manual harvest, representing berries detached from plant resources and undergoing post-harvest physiological and quality changes, with berry condition and handling during and after harvest critically affecting final wine or fresh market quality. Harvested berries are characterized by= (1) physical detachment from vine with pedicel separation at abscission zone or breakage (2) cessation of water and nutrient import from plant with berries becoming closed systems dependent on internal resources (3) continuation of metabolic activity including respiration (consuming sugars and acids), enzymatic reactions (affecting color, flavor, texture), and potential microbial activity (fermentation, spoilage) depending on handling and timing (4) temperature-dependent quality changes with warm harvested berries (>25°C) undergoing rapid quality degradation versus cool berries (12-24 hours) showing significant quality degradation particularly for white grapes (oxidation, off-flavors) and damaged berries (spoilage). Environmental conditions during harvest affect harvested berry quality= cool harvest conditions (30°C) accelerating degradation rainy conditions creating wet berries with dilution and rot risk requiring careful sorting and handling.
Harvested_Berry represents post-detachment state and is distinguished from on-vine mature berries by cessation of plant resource supply and beginning of post-harvest quality changes. Harvested berries require appropriate handling to maintain quality, with temperature control, timing, and gentle handling critically important. Sub-classification into Freshly_Harvested and Post_Harvest reflects temporal progression after detachment. Understanding harvested berry physiology and quality evolution is essential for harvest logistics, post-harvest handling, and maintaining berry quality from vineyard to winery or market.
Lese-Traube
abgetrennte Traube
detached berry
geerntete Beere
picked berry
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 265-280 Zoecklein, B.W. et al. (1995). Wine Analysis and Production, pp. 85-120 Morris, J.R. (2000). Past, present, and future of vineyard mechanization. In= Proceedings of the ASEV 50th Anniversary Meeting, pp. 155-164.
Harvested Berry
Haziness assessment comparing consumer complaint (conventional Chardonnay showing unexpected haze after refrigeration, turbidity 25 NTU from protein instability indicating production fault requiring recall consideration) with deliberate turbidity (natural Chenin Blanc bottled with visible lees, turbidity 45 NTU clearly labeled as unfiltered and accepted by target consumer), demonstrates how identical visual defect in one context constitutes authentic quality marker in another, with appropriate assessment requiring context consideration rather than universal turbidity rejection.
Haziness quality is a clarity quality that represents the presence of visible turbidity from suspended particles scattering light, characterizing the cloudy, veiled, or opaque visual impression that indicates either production fault (instability, contamination), incomplete processing (pre-bottling state), or deliberate stylistic choice (unfiltered, natural wine) depending on context. Within the BFO framework, this quality specifically addresses the particle-present dimension of clarity as the opposite of limpidity, distinguishing intentional turbidity (stylistic haziness) from unintentional turbidity (fault haziness) while recognizing that identical visual appearance may indicate quality failure or quality choice depending on wine category and producer intention. The ontological significance lies in haziness occupying a uniquely ambivalent position in wine quality assessment, where turbidity historically indicated spoilage or poor winemaking but now may signal authentic, minimal-intervention production valued by natural wine consumers, creating evaluation complexity that requires context consideration rather than automatic fault assignment. The physical basis for haziness involves light scattering by suspended particles with dimensions comparable to visible light wavelengths (approximately 0.4-0.7 μm and larger), with scattering intensity depending on particle size, concentration, and refractive index contrast with the wine medium, creating turbidity ranging from slight veil (barely perceptible) through cloudy (obviously turbid) to opaque (light-blocking). The compositional sources of haziness include microbiological particles (yeast cells 5-10 μm, bacteria 0.5-2 μm remaining from fermentation or contamination), protein-tannin complexes (forming post-bottling when unstable proteins encounter tannins), tartrate crystals (potassium bitartrate precipitating at low temperatures), and various colloidal aggregates (polysaccharides, pectins, mannoproteins forming haze under specific conditions). The fault versus feature distinction for haziness depends on producer intention, wine category, and consumer expectation, with haziness indicating fault when unexpected (conventional wine developing post-bottling haze) but representing deliberate quality when intended (natural wine bottled unfiltered with visible lees), requiring context-appropriate assessment. Understanding haziness quality enables fault diagnosis when turbidity is unintended, appropriate assessment accepting turbidity in natural wine contexts, and stability evaluation predicting haze development risk in susceptible wines.
Haziness_Quality addresses the turbidity dimension of clarity, representing particle-induced light scattering that may indicate fault or feature depending on context. This quality requires careful distinction between unintended haziness (indicating instability or contamination) and intended haziness (reflecting natural wine production philosophy). The class supports both fault diagnosis and context-appropriate evaluation that respects stylistic diversity in clarity standards.
Schleier
Trübheit
Trübung
cloudiness
haze
turbidity
veiled appearance
wolkige Erscheinung
Waters EJ et al. (2005) Preventing protein haze in bottled white wine. Australian Journal of Grape and Wine Research 11(2): 215-225 Cosme F et al. (2020) Wine lees: characteristics and potential applications. Annual Review of Food Science and Technology 11: 559-578
Haziness Quality
A head-trained arm on Vitis vinifera cv. Grenache in Châteauneuf-du-Pape, emerging from compact head at 35cm height, extending 28cm outward and upward at approximately 45° angle, bearing 4 two-bud spurs distributed along length, one of four arms radiating from central head creating characteristic gobelet (goblet) architecture, total vine with four arms bearing 16 combined spurs supporting approximately 30 shoots and 2.8-3.2kg crop in compact bush structure approximately 60cm diameter and 80cm height, representing traditional head-trained architecture maintained in prestigious AOC where regulations encourage or require traditional low-yielding training approaches emphasizing terroir expression and wine character over productivity maximization.
A permanent arm originating from a head-trained trunk configuration where the trunk terminates in a compact swollen head structure from which multiple arms radiate, characterized by short arm lengths (typically 20-60cm) emerging from central head, spur-pruned fruiting positions distributed along arms, and architectural configuration creating compact bush-like or goblet-shaped canopy rather than extended linear cordon structure. Head-trained arms represent traditional training approach predating modern cordon systems, still employed in many prestigious wine regions (Châteauneuf-du-Pape, Priorat, parts of Rioja, traditional Barolo) where compact head-trained architecture reflects historical practice, terroir expression philosophy, hand-harvest requirements, or perceived quality benefits from low-yielding compact vine structure, though with reduced mechanization compatibility and increased labor requirements compared to cordon-trained alternatives.
Head-trained arms differ fundamentally from cordon arms in origin (radiating from compact head versus extending from vertical trunk), length (short 20-60cm versus long 60-150cm+ cordons), and training philosophy (compact bush structure versus extended linear framework). The head-training approach dominated viticulture historically before wire trellis development, with self-supporting bush vines requiring no trellis infrastructure. Modern persistence of head training reflects tradition, quality perceptions (association with prestigious regions and terroir-focused production), regulatory requirements (some AOC/DOC systems mandating traditional training), practical considerations (steep terrain unsuitable for trellis installation), and in some cases genuine quality benefits from low yields, compact structure, and minimal intervention philosophy. However, head training creates significant challenges including difficult mechanization, labor-intensive hand operations, typically lower yields, and limited adaptability to modern vineyard management technologies.
Buscharm
Kopferziehung-Arm
bush-trained arm
gobelet arm
head-pruned arm
stock arm
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Reynolds & Vanden Heuvel (2009) "Influence of grapevine training systems on vine growth and fruit composition= a review" American Journal of Enology and Viticulture 60=251-268 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115.
Head Trained Arm
A Carignan vine in Languedoc gobelet training bears a Head_Trained_Vine_Role, its 80-year-old head supporting 6-8 short arms with spur positions, requiring no trellis while producing concentrated fruit from low yields—the head training adapted to hot, dry, windy conditions. Old-vine Zinfandel in Lodi's sandy soils serves in a Head_Trained_Vine_Role, the historic bush training reflecting pre-Prohibition planting practices and contributing to the distinctive "old vine" character commanding premium pricing. A Santorini Assyrtiko vine trained in the traditional "kouloura" basket form bears a Head_Trained_Vine_Role, the coiled cane arrangement protecting fruit from extreme wind and heat in the volcanic island's challenging conditions.
A training system role that inheres in a grapevine trained without permanent horizontal cordons, with fruiting wood arising directly from a head (the enlarged upper portion of the trunk) or from short arms radiating from the head, creating a self-supporting or minimally supported vine structure often described as "bush" or "gobelet" form. Head-trained vines represent the oldest viticultural training approach, requiring no trellis infrastructure and offering wind resistance, heat tolerance through self-shading, and adaptation to low-vigor sites where cordon extension would be difficult to achieve. Annual pruning of head-trained vines may be spur-based (retaining short spurs on permanent arms) or cane-based (replacing fruiting canes annually), with the head structure providing the framework from which annual growth emerges. Head training predominates in traditional Mediterranean viticulture (southern France, Spain, southern Italy, Greece), arid regions (parts of Australia, California, Chile), and any situation where trellis costs, wind exposure, or site access favor self-supporting vines. The labor intensity of head-trained vine management (difficult mechanization, hand pruning and harvest requirements) has driven conversion to trellised systems in many regions, though quality-focused and traditional producers maintain head training for its perceived benefits to fruit concentration and terroir expression.
Head_Trained_Vine_Role is subclassed by specific head-training forms—Gobelet (formal spur-pruned Mediterranean style) and Bush_Vine (informal self-supporting form)—though considerable overlap exists between categories. The head represents the permanent structure from which fruiting wood emerges, typically becoming enlarged and gnarled with age as annual pruning creates layered wound tissue. Head training is particularly suited to varieties tolerant of or benefiting from fruit shading (Grenache, Carignan, Mourvèdre) and challenging conditions (wind, heat, drought, steep slopes) where trellis installation or maintenance is problematic. The "old vine" designation frequently applied to head-trained vineyards reflects both the training system's antiquity and the correlation between age and head-training prevalence—many head-trained vineyards date from periods before trellising became standard practice. Economic pressures favor conversion to trellised systems for new plantings, making surviving head-trained vineyards increasingly valued for their heritage and distinctive fruit character.
Gobelet-Rebe
Kopferziehung
free-standing vine
freistehende Rebe
gobelet vine
head-trained vine
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Dry PR (2000) Canopy management for fruitfulness, Australian Journal of Grape and Wine Research 6: 109-115 Gladstones J (1992) Viticulture and Environment, Winetitles Huglin P, Schneider C (1998) Biologie et écologie de la vigne, Lavoisier
Head Trained Vine Role
Cabernet Sauvignon on 110R rootstock healed graft union zone at 12 years post-grafting shows 16-24 mm well-integrated zone 12-16 cm above soil, complete vascular continuity with xylem vessels 160-280 μm diameter crossing interface enabling 22-42 L/day water transport (90-94% non-grafted capacity), continuous phloem supporting 12-20 g/day photosynthate flow, functional cambium producing coordinated annual rings, moderate symmetrical swelling 110-116%, intact protective bark, mechanical strength 80-88% non-grafted equivalent, no necrotic zones or incompatibility symptoms, supporting 12 years healthy productive growth indicating successful permanent integration
Healed graft union zone is graft union zone that has achieved successful tissue integration with complete vascular continuity, functional cambium, strong mechanical connection, and absence of incompatibility symptoms, representing a successfully established scion-rootstock junction enabling normal vine function and longevity. Healed graft union zone integration characteristics includes= complete tissue contact (scion and rootstock tissues fully adhered without gaps, separation, or intervening dead zones), vascular continuity (xylem and phloem elements forming continuous functional pathways across interface enabling normal bidirectional transport), continuous cambium (functioning vascular cambium spanning union producing coordinated annual xylem and phloem increments), organized tissue structure (union zone showing differentiated organized vascular tissues rather than disorganized callus), strong cell adhesion (scion and rootstock cells firmly bonded at cellular level preventing mechanical separation), and absence of necrosis (no dead or dying tissue zones interrupting integration). Healed graft union zone vascular functionality includes= xylem transport capacity (water flow across union approaching 85-95% of non-grafted trunk capacity, adequate for supporting normal canopy transpiration), phloem transport capacity (photosynthate transport 80-95% of non-grafted capacity supporting normal root growth and fruit development), minimal hydraulic resistance (union adding 130% in incompatible, healed showing organized tissues versus disorganized or necrotic in incompatible, healed showing normal growth versus declining vigor in incompatible, healed showing mechanical strength versus weakness or breakage in incompatible, and healed maintaining function decades versus incompatible failing within 5-15 years. Healed graft union zone maintenance and protection includes= avoiding mechanical injury (careful pruning, equipment operation preventing union damage), monitoring for problems (regular visual inspection detecting early signs of decline, disease, or mechanical issues), disease prevention (protecting union from pathogen entry through proper technique, wound protection, canker management), freeze protection in extreme climates (wrapping or painting young unions reducing cold injury risk in marginal regions), and appropriate rootstock selection (choosing compatible combinations maximizing healing success and long-term union quality). Healed graft union zone significance includes= enabling successful grafted viticulture (healed unions providing functional scion-rootstock integration supporting phylloxera-resistant, soil-adapted, vigor-controlled vineyard establishment), determining vine productivity and longevity (successful union healing essential for normal vine performance and 30-60+ year lifespan), reflecting skilled propagation (high healing rates 85-95% indicating proper grafting technique, timing, aftercare), influencing vineyard value (well-healed uniform unions contributing to vineyard quality, vigor uniformity, longevity), and foundational for rootstock benefits (all advantages of grafted viticulture depending on achieving healed functional union enabling scion-rootstock physiological integration).
Healed_Graft_Union_Zone emphasizes successful integration outcome and is distinguished from Graft_Union_Zone (general transitional tissue), Callus_Graft_Union_Zone (callus emphasis), and Incompatibility_Zone (failed integration) by focus on complete functional healing. Critical for understanding successful graft integration criteria, healing timeline and assessment, vascular and mechanical function achievement, compatibility expression, and healed union as foundation for grafted vine productivity, longevity, and rootstock benefit realization requiring proper grafting technique and compatible scion-rootstock selection.
Established union
Functional graft zone
Funktionale Pfropfzone
Integrierte Veredelung
Successfully integrated union
Verheilte Veredlungsstelle
Healed Graft Union Zone
Merlot healing callus tissue at 3 weeks post-grafting shows 10-18 mm thick cream-colored tissue covering graft interface, callus cells 45-75 μm diameter actively dividing (8-15% cells showing mitotic figures), cells with dense cytoplasm and prominent nuclei indicating high metabolic activity, callus completely bridging scion-rootstock junction providing continuous tissue connection, early differentiation zones visible with elongated cambium-like cells beginning to organize, callus providing pathogen barrier, moisture retention, substrate for progressive vascular differentiation enabling functional union establishment within subsequent 3-6 weeks, representing essential healing tissue determining graft success
Healing callus tissue is callus tissue emphasizing its functional role in wound healing and graft union establishment, comprising actively proliferating undifferentiated cells that cover wounds, bridge grafts, exclude pathogens, and serve as substrate for differentiated tissue formation restoring structural and functional integrity to injured vines. Healing callus tissue formation timeline includes= initial proliferation (beginning 3-10 days post-wounding or grafting, first callus cells appearing at injury margins or graft surfaces), rapid expansion (weeks 1-4, callus rapidly proliferating covering exposed surfaces, filling gaps), maximal callus mass (weeks 2-6, peak callus accumulation before differentiation begins), progressive differentiation (weeks 4-12, callus gradually transforming into organized tissues), and replacement by permanent tissues (months 3-12+, callus progressively replaced by vascular tissues, protective periderm, completing healing). Healing callus tissue protective functions includes= covering exposed surfaces (callus rapidly covering wound or graft surfaces preventing desiccation of underlying living tissues), pathogen exclusion (callus forming physical barrier preventing fungal spores and bacteria from reaching susceptible vascular tissues), moisture retention (callus tissue maintaining humid microenvironment supporting underlying tissue survival and healing), mechanical protection (soft callus tissue cushioning underlying sensitive cambium and developing tissues), and creating favorable environment (callus providing stable substrate enabling subsequent organized tissue formation). Healing callus tissue in wound closure includes= arising from wound margins (callus proliferating from living cells adjacent to wound particularly cambial region), expanding over wound surface (callus growing across exposed wood, filling pruning cuts, covering injury sites), woundwood formation (callus differentiating into secondary xylem and phloem at wound margins gradually occluding wound), protective periderm differentiation (callus forming cork cambium producing phellem sealing wound surface), and complete closure (wound eventually fully covered by callus-derived tissues restoring protective bark covering and structural continuity). Healing callus tissue in graft union formation includes= proliferating from both partners (scion and rootstock independently producing callus at cut surfaces), bridging junction (callus tissues meeting forming continuous connection across graft interface), enabling tissue contact (callus filling irregularities ensuring intimate contact between scion and rootstock necessary for integration), providing differentiation substrate (callus serving as precursor tissue from which functional vascular elements and cambium arise), and determining union success (adequate healing callus formation, organization, differentiation distinguishing successful grafts from failures). Healing callus tissue cellular activities includes= active cell division (cells dividing 1-3 times per week during peak formation), cell expansion (daughter cells enlarging providing tissue growth), cell-cell adhesion (cells adhering forming cohesive tissue mass), metabolic activity (high respiration, protein synthesis supporting rapid growth), and progressive differentiation (cells transitioning from undifferentiated state toward specialized fates as healing proceeds). Healing callus tissue differentiation into permanent tissues includes= cambium formation (callus cells differentiating into cambial initials organizing into functional vascular cambium typically 3-8 weeks post-wounding), xylem differentiation (cambium producing vessel elements, fibers establishing water transport and mechanical support), phloem differentiation (sieve tubes, companion cells forming enabling photosynthate transport), periderm formation (cork cambium arising in callus producing protective cork layers sealing surface), and ray formation (radial parenchyma developing providing lateral transport connections). Healing callus tissue adequate versus inadequate formation includes= adequate healing callus (sufficient callus formation completely bridging wound or graft, smooth continuous tissue mass covering exposed areas), inadequate callus (insufficient callus leaving gaps, discontinuities, or exposed tissues preventing complete healing), factors affecting formation (tissue vigor, temperature, moisture, hormonal status, pathogen presence influencing callus quantity and quality), and consequences (adequate callus enabling successful healing or union, inadequate callus causing healing failure, graft rejection, or persistent wounds vulnerable to desiccation or infection). Healing callus tissue environmental requirements includes= optimal temperature (20-28°C supporting rapid proliferation, 35°C potentially inhibiting or causing cell death), high humidity (85-95% relative humidity preventing callus desiccation supporting survival and growth), adequate oxygen (aerobic conditions necessary for callus respiration and metabolism), protection from light (UV radiation potentially damaging callus, darkness or shading typical during healing), and mechanical stability (immobilization reducing disruption supporting callus bridge integrity particularly in grafts). Healing callus tissue hormonal regulation includes= auxin stimulation (elevated IAA at wounds or grafts promoting cell division and callus proliferation), cytokinin requirement (cytokinins synergizing with auxin enhancing callus formation), auxin, cytokinin ratio (balanced ratio supporting continued callus growth, ratio shifts triggering differentiation), wound hormones (jasmonic acid, ethylene potentially modulating wound response and callus formation), and signal integration (multiple hormone pathways coordinating healing response determining callus formation rate and characteristics). Healing callus tissue quality indicators includes= color (white to cream healthy callus versus brown or black indicating necrosis or oxidation), texture (firm cohesive callus indicating viable tissue versus soft mushy suggesting cell death or infection), continuity (complete coverage and bridging versus gaps or separations), vitality (actively growing expanding callus versus static or shrinking indicating inadequate healing), and progression (advancing differentiation producing organized tissues versus persistent disorganized callus suggesting problems). Healing callus tissue in successful healing includes= rapid formation (callus appearing within 1-2 weeks providing prompt wound coverage), adequate quantity (sufficient callus completely bridging or covering injury), good organization (callus showing progressive structure, early vascular strand formation), timely differentiation (callus transitioning to organized tissues within 4-12 weeks), complete integration (callus-derived tissues fully incorporating into trunk structure), and restoration of function (healed wounds or unions showing normal transport, mechanical strength). Healing callus tissue in problematic healing includes= delayed or absent formation (little or no callus appearing within 2-3 weeks suggesting poor tissue response), insufficient quantity (sparse callus leaving gaps or incomplete coverage), disorganized tissue (callus lacking structure, showing random cell orientation without differentiation), persistent undifferentiated callus (callus remaining undifferentiated beyond 8-12 weeks without organized tissue formation), necrotic callus (browning, cell death preventing integration), and healing failure (inadequate callus resulting in non-healed wounds, failed grafts). Healing callus tissue management and optimization includes= proper technique (clean cuts, good contact, appropriate materials supporting optimal callus environment), environmental control (maintaining temperature, humidity within optimal ranges during critical healing period), protection (wrapping, waxing, or other measures protecting callus from desiccation, mechanical damage, pathogens), monitoring (observing callus formation, progression detecting problems early), and intervention (adjusting conditions or providing support if callus formation inadequate). Healing callus tissue significance includes= enabling wound repair (callus providing initial healing response restoring protective covering and structural integrity), critical for grafting success (healing callus bridging scion and rootstock determining union establishment), reflecting tissue vigor and health (robust callus formation indicating good physiological condition, poor formation suggesting stress or damage), determining healing timeline (callus formation rate affecting duration to complete wound closure or functional union), and foundational for regeneration (healing callus serving as precursor tissue from which all permanent healing structures arise).
Healing_Callus_Tissue emphasizes wound repair function and is distinguished from Callus_Tissue (general undifferentiated tissue), Callus_Graft_Union_Zone (union context), and Bridge_Callus_Tissue (junction bridging) by focus on active healing role. Critical for understanding wound response mechanisms, healing progression timeline, callus protective functions, differentiation substrate role, environmental and hormonal requirements, healing success criteria, and healing callus as essential tissue determining wound closure, graft establishment, and vine capacity for injury repair and tissue regeneration.
Heilungs-Kallusgewebe
Regenerative tissue
Regeneratives Gewebe
Repair callus
Wound healing callus
Wundheilungs-Kallus
Healing Callus Tissue
The health status declaration for Cabernet Sauvignon clone 337 issued by ENTAV states "Matériel certifié conforme aux exigences sanitaires de la Directive 2005/43/CE, exempt des organismes nuisibles listés à l'Annexe II" (Certified material conforming to the sanitary requirements of Directive 2005/43/EC, free from harmful organisms listed in Annex II), accompanied by the phytosanitary passport number enabling traceability through the EU internal market. The health status declaration for Grüner Veltliner clone 17 issued by the Bundesamt für Weinbau reads "Virusgetestetes Pflanzgut — frei von GFLV, ArMV, GLRaV-1, GLRaV-3, GVA, GFkV gemäß Rebenverkehrsgesetz § 4 Abs. 2" (Virus-tested plant material — free from [listed viruses] according to the Vine Marketing Act § 4 para. 2), with the declaration valid for a 5-year period subject to re-testing confirmation.
Health status declaration is a certification status entity that records the formal official document or statement issued by the certifying authority attesting to the phytosanitary condition of a clonal line based on the integrated assessment of all virus testing results (ELISA, PCR, biological indexing), field disease evaluations, and regulatory compliance verification, constituting the legal instrument that authorizes or restricts the propagation, distribution, and commercial sale of the certified plant material and that accompanies the material through all stages of the nursery supply chain to the final grower.
The health status declaration represents the culmination of the entire certification testing process, distilling complex diagnostic results into a legally binding statement of phytosanitary compliance. Within the EU, the health status declaration is integral to the plant passport system (Regulation EU 2016/2031) that governs the movement of plant material within the internal market and replaces phytosanitary certificates for intra-EU trade. For international trade outside the EU, the IPPC (International Plant Protection Convention) model phytosanitary certificate provides the framework for health declarations accompanying exported grapevine material. The legal force of health status declarations varies by jurisdiction — in some systems, the declaration constitutes a warranty of virus-free status (with liability implications if certified material is subsequently found infected), while in others it represents a statement of testing results at the time of assessment without ongoing health guarantee. The distinction between "virus-free" (tested and found negative) and "virus-tested" (tested but with specified positive results for non-certification-relevant agents) declarations reflects the pragmatic reality that some near-ubiquitous agents (e.g., GRSPaV) cannot be economically eliminated from all material. Nursery operators bear legal responsibility for maintaining the validity of health status declarations through their multiplication chain — any post-certification infection detected in nursery mother blocks triggers an obligation to withdraw affected material and notify the certification authority. Electronic plant passport systems and blockchain-based traceability initiatives are progressively modernizing health status declaration management, enabling real-time verification of phytosanitary status throughout the supply chain.
Gesundheitserklärung
Gesundheitszeugnis
Pflanzengesundheitszeugnis
health certificate
phytosanitary certificate
phytosanitary declaration
phytosanitäre Erklärung
plant health statement
sanitary declaration
sanitärer Befundschein
European Parliament (2016) Regulation (EU) 2016/2031 on protective measures against pests of plants European Commission (2005) Commission Directive 2005/43/EC EPPO (2008) Certification scheme for pathogen-tested material of grapevine varieties. EPPO Bulletin 38(3): 422-429 Mannini F, Digiaro M (2017) The effects of viruses and viral diseases on grapes and wine. In Grapevine Viruses, Springer, pp.: 453-482
Health Status Declaration
Grenache heartwood at 35 years post-planting shows 52-68 mm radius (comprising 62-68% of total 85-105 mm trunk radius), dark tan to light brown coloration from phenolic deposits (only slightly darker than surrounding 18-25 mm sapwood), vessels completely occluded with tyloses and gums, dead parenchyma with no starch storage capacity, wood density 0.54-0.62 g/cm³ providing structural support, shows some colonization by white rot fungi causing minor decay but maintaining overall integrity
Heartwood is the inner non-functional wood zone in Vitis vinifera trunk comprising aged xylem rings where water conduction has ceased through vessel occlusion or embolism, parenchyma cells have died losing metabolic activity and storage capacity, and wood serves solely structural support while often accumulating phenolic compounds producing characteristic darker coloration. Heartwood forms through physiological processes converting inner sapwood= progressive death of parenchyma cells (aging parenchyma losing viability 8-15 years post-formation), vessel occlusion through tyloses formation (parenchyma cells growing into vessel lumens blocking water transport), gum and resin accumulation in vessels (further occluding lumens), phenolic compound deposition in cell walls and lumens (tannins, stilbenes, other secondary metabolites producing darker color), and loss of all physiological functions except mechanical support. Heartwood anatomical characteristics include= occluded xylem vessels with tyloses, gums, or resins filling lumens (preventing water transport), dead parenchyma cells with degraded cytoplasm and organelles (no metabolic activity or storage capacity), lignified fiber cells providing mechanical strength (structural support function maintained), no living tissue or metabolic activity (contrasting with living sapwood), darker coloration than sapwood (brown, tan, reddish, or grey hues from phenolic deposits, though grapevine heartwood often only slightly darker than sapwood), and often central position (forming core of trunk surrounded by sapwood annulus). Heartwood formation timing and pattern includes= beginning when trunk reaches 15-30 mm radius (typically year 5-10 depending on growth rate), progressing from pith outward as new sapwood forms externally (innermost rings transitioning first), rate of formation approximately balancing sapwood maintenance (sapwood depth relatively constant as heartwood accumulates), more rapid formation in fast-growing vigorous vines (wide annual rings transitioning to heartwood after fewer years), slower formation in stressed slow-growing vines (narrow rings potentially maintaining sapwood function longer). Heartwood radial dimensions vary by vine age= young vines (3-7 years) showing no heartwood or minimal central core (0-5 mm radius), mature vines (10-20 years) showing 10-40 mm heartwood radius (30-60% of total trunk radius), old vines (25-50 years) showing 35-85 mm heartwood radius (55-75% of total radius), ancient vines (50-100+ years) showing 80-200+ mm heartwood radius (70-85% of total, often with extensive decay creating hollow centers). Heartwood functions include= mechanical support of trunk and canopy (heartwood density 0.48-0.68 g/cm³ providing compressive and tensile strength), water storage capacity (dead parenchyma cells and empty vessel lumens can hold water providing minor buffering capacity), resistance to decay (phenolic compounds in heartwood providing antimicrobial properties reducing fungal colonization), and structural persistence (heartwood maintaining dimensional stability and strength long after formation). Heartwood phenolic compounds include= condensed tannins (proanthocyanidins deposited in cell walls and lumens), stilbenes (resveratrol, piceid, other stilbenoids with antimicrobial properties), lignin-derived phenolics (from cell wall degradation and modification), and various extractives (varying by cultivar, wood age, and decay status) producing characteristic color and rot resistance. Heartwood decay and dysfunction includes= white rot fungi degrading cellulose and lignin (Fomes, Stereum, and other wood decay organisms colonizing heartwood), brown rot fungi attacking cellulose (leaving lignin residue), decay typically beginning in heartwood (dead tissue providing nutrient source for saprophytic fungi), hollow trunk formation (extensive heartwood decay creating central cavity while maintaining sapwood ring and structural integrity), and decay potentially compromising structural stability (advanced decay reducing trunk strength increasing breakage risk). Heartwood contrast with sapwood includes= no water transport function (versus active conduction in sapwood), no carbohydrate storage (versus 40-80 g starch per vine in sapwood), no metabolic activity (versus active living parenchyma in sapwood), darker coloration (versus lighter sapwood though difference often subtle in grapevine), and often decayed (versus living sapwood with active defense) but contributing to mechanical support (shared function with sapwood). Heartwood significance in old vine trunks includes= occupying majority of trunk volume (70-85% of radius in old vines providing bulk of mechanical support), extensive heartwood indicating vine age and history (heartwood accumulation reflecting decades of growth), heartwood decay creating hollow trunks (common in ancient vines, potentially reducing structural integrity but allowing continued survival through sapwood ring), and heartwood characteristics varying by cultivar and site (some cultivars forming distinct dark heartwood, others showing minimal color differentiation from sapwood).
Heartwood emphasizes non-functional inner wood and is distinguished from Sapwood (functional outer wood) and specific heartwood subtypes (Inactive_Heartwood, Central_Heartwood, Dense_Heartwood) by encompassing entire physiologically dead structural wood zone. Critical for understanding trunk structural composition, age-related wood zonation, mechanical support in old vines, wood decay patterns, and hollow trunk formation in ancient vines.
Core wood
Dead wood
Kernholz
Non-conducting xylem
Totes Holz
Zentralholz
Heartwood
In Merlot vines in Saint-Émilion, Bordeaux, heat accumulation response drives budbreak progression through March and early April, with growing degree days above the 10°C base temperature accumulating from approximately 20 GDD by March 10 through 80 GDD by March 25 to the cultivar threshold of approximately 140 GDD by April 5, at which point visible bud swelling at BBCH stage 05 indicates successful completion of heat-driven budbreak initiation.
Heat accumulation response in Vitis vinifera represents the temperature-dependent progression toward budbreak that occurs following dormancy release, whereby growth-competent buds respond to warming temperatures by accumulating the thermal exposure required to trigger visible growth initiation. This process operates during late winter and early spring as the primary driver of budbreak timing in ecodormant buds, with the rate of progress toward budbreak determined by the intensity and duration of temperatures above the base threshold for growth-related metabolic activity. The physiological basis of heat accumulation response involves the temperature-dependence of metabolic and developmental processes that prepare for and execute budbreak, with warming temperatures progressively activating enzyme systems, accelerating biosynthetic reactions, and enabling the cellular processes of division and expansion that constitute visible growth. The quantification of heat accumulation uses growing degree day or heat unit models that integrate effective temperature exposure over time, with each degree above the base temperature contributing to cumulative heat accumulation according to model-specific algorithms, and budbreak occurring when accumulated units reach the cultivar-specific threshold requirement. The base temperature for heat accumulation in grapevine is approximately 10°C, below which temperatures contribute minimally or not at all to progress toward budbreak, while temperatures above this threshold contribute in proportion to their elevation above the base according to most simple linear models. The cultivar-specific heat requirement for budbreak typically ranges from 100-300 growing degree days depending on genotype, with early-budbreak cultivars generally requiring less heat accumulation than late-budbreak cultivars, creating the characteristic spread of budbreak timing among cultivars in mixed plantings. The interaction between heat accumulation and prior chilling affects response characteristics, with adequately chilled buds responding efficiently to heat accumulation while inadequately chilled buds may show delayed or reduced response reflecting residual endodormancy constraints. The practical application of heat accumulation models enables prediction of budbreak timing from temperature forecasts, supporting vineyard management decisions regarding frost protection, early season pest management, and scheduling of cultural operations relative to expected growth stage progression.
Heat_Accumulation_Response represents the third component phase of Dormancy_Release addressing temperature-driven progression to budbreak. Distinguished from Chilling_Completion which addresses endodormancy release, while Heat_Accumulation_Response addresses the subsequent ecodormancy release through warming. Subdivides into Growing_Degree_Day_Accumulation, Temperature_Sum_Response, and Heat_Unit_Requirement as component aspects. Base temperature approximately 10°C for grapevine. Cultivar heat requirements typically 100-300 GDD. Enables budbreak timing prediction from temperature data.
Wärmeakkumulations-Reaktion
Wärmeeinheiten-Reaktion
heat unit response
temperature-driven budbreak progression
temperaturgetriebene Austriebsprogression
thermal accumulation response
thermische Akkumulationsantwort
warming response
Parker AK et al (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17: 206-216 Moncur MW et al (1989) Base temperatures for budbreak and leaf appearance of grapevines. American Journal of Enology and Viticulture 40: 21-26 Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle
Heat Accumulation Response
Heat avoidance quality comparison across Vitis vinifera cultivars evaluated the contribution of tissue cooling mechanisms to observed differences in heat damage during summer heat events in Riverland Australia. Infrared thermometry during a heat event (ambient 44°C) revealed substantial variation in heat avoidance quality as reflected in berry and leaf temperatures. Grenache demonstrated excellent heat avoidance quality with leaf temperatures averaging 38.5°C (5.5°C below ambient) and berry temperatures averaging 40.2°C (3.8°C below ambient), attributed to high transpiration rates maintained through relatively open stomata despite the stress conditions, combined with smaller, more reflexed leaves reducing direct solar interception. Chardonnay showed poor heat avoidance quality with leaf temperatures averaging 42.1°C (only 1.9°C below ambient) and berry temperatures averaging 44.8°C (0.8°C above ambient due to radiation loading), reflecting stomatal closure that reduced transpirational cooling combined with larger, more horizontally oriented leaves increasing solar interception. The heat avoidance quality differential translated directly to damage patterns, with Grenache showing only 15% sunburn damage while Chardonnay exhibited 45% damage despite similar inherent tissue thermotolerance, demonstrating that avoidance mechanisms can be more important than tolerance for practical heat performance when water for transpirational cooling is available.
Heat avoidance quality is a heat stress indicator quality that inheres in a grapevine organism and represents the capacity to reduce actual tissue temperatures below ambient air temperature through morphological, physiological, and biophysical mechanisms that minimize heat absorption, maximize heat dissipation, or both, enabling maintenance of tissue temperatures within tolerable ranges even when air temperatures exceed the thermal tolerance thresholds that would cause damage at equivalent tissue temperature. This quality complements thermal tolerance quality by addressing the prevention of excessive tissue temperatures rather than the capacity to survive them, recognizing that the actual temperature experienced by tissues results from the balance between heat inputs from solar radiation and warm air and heat losses through transpiration, convection, and radiation, with this balance modulated by plant characteristics that constitute heat avoidance mechanisms. The ontological significance of heat avoidance quality lies in its role enabling survival and function under ambient conditions that might otherwise exceed tolerance thresholds, effectively extending the environmental range within which vines can maintain productivity by reducing the translation of high air temperatures into damaging tissue temperatures. The biological basis of heat avoidance encompasses multiple mechanisms including transpirational cooling that dissipates latent heat through water evaporation from leaf and fruit surfaces, convective cooling enhanced by leaf orientation and canopy architecture that promotes air movement, and radiative properties including reflectance and emissivity that influence the net radiation balance determining heat absorption. The effectiveness of heat avoidance mechanisms depends on the availability of water for transpiration and the physical characteristics of tissues including size, orientation, and surface properties that influence heat exchange with the environment. The variation in heat avoidance quality among grapevine genotypes reflects differences in leaf and canopy characteristics affecting the thermal environment of tissues, combined with physiological differences in transpiration rate and stomatal behavior under heat stress that determine the magnitude of evaporative cooling achievable. The interaction between heat avoidance and tolerance mechanisms creates the overall heat resistance that determines performance under high temperature conditions, with some genotypes relying primarily on avoidance while others depend more on tolerance, and with management practices potentially enhancing avoidance through irrigation, canopy manipulation, or protective covers.
Heat_Avoidance_Quality is_a Heat_Stress_Indicator_Quality, serving as parent class for Leaf_Angle_Adjustment_Quality, Transpirational_Cooling_Capacity_Quality, and Reflectance_Increase_Quality which address specific avoidance mechanisms. This class addresses the overall capacity to reduce tissue temperatures while child classes capture particular mechanisms. Distinguished from Thermal_Tolerance_Quality by the focus on temperature reduction rather than damage resistance. Infrared thermometry provides direct tissue temperature measurement for avoidance assessment. The interaction between avoidance and tolerance determines overall heat resistance. Water availability strongly modulates transpirational cooling as the primary avoidance mechanism.
Gewebekühlungskapazität
Hitzevermeidung
Temperaturreduktionskapazität
heat escape
temperature reduction capacity
thermal avoidance
thermische Vermeidung
tissue cooling capacity
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 453-507 Greer DH, Weedon MM (2013) The impact of high temperatures on Vitis vinifera cv. Semillon grapevine performance and berry ripening. Frontiers in Plant Science 4:491 Wahid A et al. (2007) Heat tolerance in plants: an overview. Environmental and Experimental Botany 61:199-223
Heat Avoidance Quality
Heat damage severity quality assessment following an extreme heat wave in Barossa Valley (3 days exceeding 45°C) evaluated the extent of thermal injury across Shiraz vineyard blocks with varying canopy management and irrigation status. Severely damaged blocks with sparse canopies and deficit irrigation showed heat damage severity quality scores of 4 on a 5-point scale, with 60% of exposed clusters exhibiting sunburn damage ranging from brown discoloration to complete berry desiccation, extensive leaf scorch affecting 40% of canopy area, and shoot tip necrosis on 80% of actively growing shoots. Moderately damaged blocks with denser canopies showed intermediate heat damage severity quality scores of 2.5, with sunburn limited to 25% of clusters on exposed positions and leaf scorch affecting 15% of canopy area primarily on western-exposed shoots. Protected blocks receiving supplemental irrigation for evaporative cooling and having well-developed fruit zone shading showed minimal heat damage severity quality scores below 1.5, demonstrating the effectiveness of management interventions for reducing heat damage. The heat damage severity quality assessment guided harvest decisions, with severely damaged fruit directed to lower-quality wine programs while protected fruit maintained premium designation, and informed infrastructure investments in shade cloth and irrigation capacity for future heat event protection.
Heat damage severity quality is a heat stress indicator quality that inheres in a grapevine organism and represents the extent and intensity of visible tissue injury resulting from exposure to temperatures exceeding thermal tolerance thresholds, encompassing the range of damage manifestations from minor cosmetic effects through moderate functional impairment to severe necrosis affecting vegetative and reproductive organs with consequences for current season productivity and potentially long-term vine health. This quality provides the diagnostic framework for assessing the actual damage resulting from heat events, complementing thermal tolerance quality that addresses the underlying capacity to withstand heat exposure without damage. The ontological significance of heat damage severity quality lies in its role enabling quantification of heat event impacts for production planning, quality assessment, and insurance purposes, while also providing feedback for evaluating the effectiveness of protective management practices and identifying cultivar-site combinations with inadequate heat tolerance for continued production under changing climate conditions. The biological basis of heat damage severity encompasses the progression of cellular injury from initial membrane disruption through protein denaturation to complete cellular collapse and tissue necrosis, with damage severity reflecting the magnitude by which exposure conditions exceeded tolerance thresholds and the duration of exposure at damaging temperatures. The visual manifestations of heat damage vary with the tissue affected and the intensity of exposure, including berry sunburn ranging from minor discoloration to complete desiccation, leaf scorch progressing from marginal browning to complete lamina necrosis, and shoot tip damage ranging from temporary wilting to permanent desiccation of growing points. The spatial distribution of heat damage within vineyards and canopies reflects the interaction between temperature exposure and tissue vulnerability, with sun-exposed positions experiencing higher temperatures and greater damage than shaded locations, creating characteristic damage patterns useful for diagnosis and management response planning.
Heat_Damage_Severity_Quality is_a Heat_Stress_Indicator_Quality, serving as parent class for Sunburn_Severity_Quality, Heat_Necrosis_Quality, and Leaf_Scorch_Quality which address specific damage types affecting different tissues. This class addresses the overall extent of visible heat injury while child classes capture particular damage manifestations. Scoring systems (typically 1-5) enable semi-quantitative assessment for management and economic evaluation. The spatial pattern of damage provides diagnostic information about exposure conditions. Distinguished from Thermal_Tolerance_Quality which addresses capacity rather than actual damage.
Hitzeschaden-Intensität
Hitzeschaden-Schweregrad
Temperaturschadens-Ausmaß
heat damage intensity
heat injury extent
temperature damage level
thermal injury severity
thermische Verletzungsintensität
Greer DH, Weedon MM (2013) The impact of high temperatures on Vitis vinifera cv. Semillon grapevine performance and berry ripening. Frontiers in Plant Science 4:491 Caravia L et al. (2016) Application of shade treatments during Shiraz berry ripening to reduce the impact of high temperature. Australian Journal of Grape and Wine Research 22:422-437 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 453-507
Heat Damage Severity Quality
Heat damage threshold assessment for Shiraz berries in Barossa Valley establishes anthocyanin stability threshold at 35°C where degradation rate doubles for each additional degree, visual sunburn threshold at 42°C after 6 hours exposure, and cell death threshold at 47°C after 2 hours or 52°C for 15 minutes, enabling development of a heat warning system that integrates temperature forecasts with duration predictions to trigger protective irrigation when cumulative heat exposure approaches damage thresholds.
Heat damage threshold quality is a critical temperature threshold quality that represents the temperature above which irreversible cellular damage occurs through protein denaturation, membrane lipid phase transitions, enzyme inactivation, and metabolic dysfunction that compromise tissue viability or quality compound integrity in grape berries. Within the BFO framework, this quality characterizes the upper thermal limit beyond which normal cellular function cannot be maintained, distinguishing the temperature range of reversible heat stress responses from the onset of permanent damage. The ontological significance lies in heat damage thresholds defining the practical upper boundary for grapevine function that constrains viticultural suitability under warming climate scenarios, where exceedance of these thresholds causes yield loss through flower abortion, berry sunburn, or vine mortality, and quality degradation through anthocyanin breakdown, aroma volatilization, and acid respiration. Heat damage thresholds vary by tissue type and the specific damage mechanism considered, with photosynthetic damage occurring above 40°C when thylakoid membranes destabilize, berry anthocyanin degradation accelerating above 35°C, and acute cell death occurring above 45-50°C depending on exposure duration. The time-temperature relationship is critical because brief exposure to high temperatures may be survived while prolonged exposure at the same temperature causes cumulative damage, requiring threshold specification to include both temperature and duration dimensions. Measurement approaches include controlled exposure studies quantifying damage at various temperature-duration combinations, field damage assessment correlated with recorded temperatures, and biochemical analysis of heat shock protein expression marking stress response activation. Understanding heat damage thresholds enables heat wave impact prediction, variety selection for warm regions, and evaluation of mitigation strategies including irrigation cooling, shade deployment, and harvest timing adjustment.
Heat_Damage_Threshold_Quality addresses upper thermal limits, complementing Freezing_Point_Quality for lower limits and Optimal_Photosynthesis_Temperature_Quality for functional optimum. This quality requires specification of the damage type assessed (visual injury, compositional change, cell death) as these occur at different thresholds. The class supports climate suitability analysis and heat stress management, with increasing relevance under climate warming scenarios affecting traditional viticulture regions.
Hitzeschadengrenze
Hitzeschadenschwelle
Wärmeschadenschwellenwert
heat injury threshold
heat stress threshold
high temperature tolerance limit
thermal damage limit
thermische Schädigungsgrenze
Greer DH, Weedon MM (2013) The impact of high temperatures on Vitis vinifera cv. Semillon grapevine performance and berry ripening. Frontiers in Plant Science 4: 491 Mori K et al. (2007) Loss of anthocyanins in red-wine grape under high temperature. Journal of Experimental Botany 58(8): 1935-1945
Heat Damage Threshold Quality
Heat impact on development quality assessment in Napa Valley Cabernet Sauvignon compared fruit composition from a heat-wave vintage (5 days exceeding 40°C during véraison) versus a normal vintage to quantify the developmental consequences of extreme heat exposure during the critical color development period. The heat-affected vintage showed pronounced heat impact on development quality with sugar accumulation accelerated to reach 25°Brix 12 days earlier than normal, while anthocyanin concentration reached only 1.2 mg/g versus 1.8 mg/g in the normal vintage representing a 33% color deficit despite equivalent or greater sugar accumulation. Titratable acidity at equivalent Brix was 4.8 g/L versus 6.2 g/L in the normal vintage, with pH elevated to 3.72 versus 3.52, reflecting accelerated malic acid respiration under heat. Methoxypyrazine levels were 8 ng/L versus 15 ng/L, indicating heat-accelerated degradation of herbaceous character compounds, while fruity ester concentrations showed 25% reduction attributed to volatilization losses during high-temperature periods. The comprehensive heat impact on development quality assessment established that the heat wave had fundamentally altered fruit composition in ways that would affect wine style regardless of winemaking technique, with accelerated physiological ripeness decoupled from flavor and phenolic maturity creating harvest timing dilemmas between sugar targets and quality optimization.
Heat impact on development quality is a heat stress indicator quality that inheres in a grapevine organism and represents the extent to which elevated temperatures during critical developmental periods alter the normal progression of growth, ripening, and compositional evolution in vegetative and reproductive organs, with particular significance for berry development where heat exposure during ripening can fundamentally modify the balance of sugars, acids, phenolics, and aromatic compounds that determine wine quality potential. This quality captures the developmental consequences of heat exposure distinct from the direct damage effects addressed by heat damage severity quality, recognizing that temperatures insufficient to cause visible injury may nonetheless substantially alter developmental trajectories and fruit composition through effects on metabolic rates, enzyme activities, and biosynthetic pathway function. The ontological significance of heat impact on development quality lies in its role determining the qualitative outcomes of heat exposure for wine production, where the compositional modifications induced by ripening-period heat may have greater practical importance than visible damage since they affect the fundamental character of wines produced from affected fruit even when no damage is apparent. The biological basis of heat impact on development encompasses the temperature sensitivity of the metabolic processes governing berry composition, including the acceleration of sugar accumulation and organic acid respiration at elevated temperatures, the thermal inhibition of anthocyanin biosynthesis and the degradation of existing pigments, and the volatilization or thermal degradation of aromatic compounds that define varietal character. The timing of heat exposure relative to developmental stage determines which compositional parameters are most affected, with pre-véraison heat affecting cell division and berry size potential, véraison-period heat affecting color development initiation, and post-véraison heat affecting the sugar-acid balance and aromatic profile of ripening fruit. The decoupling of developmental processes under heat stress creates the characteristic compositional signature of heat-affected fruit, with sugar accumulation proceeding or accelerating while acid retention, color development, and flavor maturation are compromised, producing fruit with high potential alcohol but lacking the acidity, color, and complexity associated with quality wine production.
Heat_Impact_On_Development_Quality is_a Heat_Stress_Indicator_Quality, serving as parent class for Anthocyanin_Degradation_Quality, Aroma_Compound_Volatilization_Quality, Acid_Respiration_Loss_Quality, and Berry_Shrivel_Quality which address specific compositional impacts of heat exposure. This class addresses the overall developmental and compositional consequences of heat while child classes capture particular quality-relevant effects. Distinguished from Heat_Damage_Severity_Quality by the focus on compositional modification rather than visible tissue damage. The decoupling of sugar from phenolic/aromatic maturity represents the core quality challenge from heat exposure. Critical for understanding wine quality impacts of warming climate conditions.
Hitze-Reifungseffekte
Hitzeauswirkung auf Entwicklung
heat developmental effects
heat ripening effects
temperature-induced compositional change
temperaturinduzierte Kompositionsänderung
thermal development impact
thermische Entwicklungseffekte
Greer DH, Weston C (2010) Heat stress affects flowering, berry growth, sugar accumulation and photosynthesis of Vitis vinifera cv. Semillon grapevines grown in a controlled environment. Functional Plant Biology 37:206-214 Mori K et al. (2007) Loss of anthocyanins in red-wine grape under high temperature. Journal of Experimental Botany 58:1935-1945 Sadras VO, Moran MA (2012) Elevated temperature decouples anthocyanins and sugars in berries of Shiraz and Cabernet Franc. Australian Journal of Grape and Wine Research 18:115-122
Heat Impact On Development Quality
Controlled environment investigation of Heat_Induced_Abscission_Resistance_Disposition at CSIRO Adelaide exposed fruiting cuttings of 15 cultivars to heat treatments (40°C for 6 hours daily for 5 consecutive days) during the lag phase of berry development and quantified berry drop at 7 and 14 days post-treatment. Additionally, pedicel abscission zone tissues were analyzed for cell wall-degrading enzyme activity and ethylene evolution. Heat_Induced_Abscission_Resistance_Disposition varied dramatically among cultivars. 'Grenache' demonstrated exceptional resistance with only 3% berry drop following heat treatment (versus 2% in untreated controls), showing minimal increase in abscission zone polygalacturonase activity (1.4-fold) and low ethylene production (12 nL g⁻¹ h⁻¹). 'Merlot' showed intermediate susceptibility (18% drop, 3.2-fold enzyme induction, 28 nL g⁻¹ h⁻¹ ethylene). 'Pinot Noir' exhibited severe heat-induced abscission (42% berry drop, 6.5-fold enzyme induction, 55 nL g⁻¹ h⁻¹ ethylene), with abscission continuing for 10 days post-treatment. Application of the ethylene perception inhibitor 1-MCP prior to heat treatment reduced berry drop by 60-70% across susceptible cultivars, confirming ethylene involvement in heat-induced abscission. Correlation analysis revealed that cultivar differences in Heat_Induced_Abscission_Resistance_Disposition were strongly predicted by the magnitude of heat-induced ethylene production (r = 0.85), suggesting that ethylene biosynthesis regulation under heat stress is a key target for improving this trait.
A Heat_Induced_Abscission_Resistance_Disposition is a Berry_Heat_Tolerance_Disposition that inheres in Vitis vinifera grape berries and pedicels and is realized through mechanisms that prevent premature berry detachment during heat stress events, preserving cluster integrity and yield by maintaining pedicel attachment even when berry tissues experience potentially damaging temperatures. This disposition represents resistance to heat-triggered abscission, a significant cause of yield loss in hot viticultural regions where extreme temperatures can induce berry drop ranging from scattered individual berries to substantial cluster portions. Heat-induced abscission involves activation of the abscission zone at the pedicel-berry junction, where cell wall-degrading enzymes (cellulases, pectinases, polygalacturonases) are induced by stress, weakening the attachment until mechanical separation occurs. The process is mediated by hormonal signals, with ethylene promoting abscission and auxin suppressing it, and heat stress can disrupt the hormonal balance favoring detachment. Heat_Induced_Abscission_Resistance_Disposition depends on the sensitivity of the abscission zone to heat-induced enzymatic activation, the hormonal response of pedicel tissues to heat stress, and potentially the mechanical strength of the pedicel-berry connection independent of enzymatic weakening. Cultivars with strong resistance maintain berry attachment through heat events that cause significant abscission in susceptible cultivars, preserving yield potential despite thermal stress.
Heat_Induced_Abscission_Resistance_Disposition is_a Berry_Heat_Tolerance_Disposition. Abscission involves enzymatic weakening of the pedicel-berry junction, mediated by ethylene signaling. Resistance depends on limited activation of abscission zone enzymes and controlled ethylene production under heat stress. Berry drop timing (lag phase, véraison, ripening) affects susceptibility, with lag phase often most sensitive. Ethylene inhibitors (1-MCP, AVG) can reduce heat-induced abscission, suggesting potential management interventions.
Beerenfall-Resistenz
Hitzeinduzierte-Abszissions-Resistenz-Disposition
Thermische-Abszissions-Toleranz
berry drop resistance
heat shatter resistance
thermal abscission tolerance
Greer DH, Weston C (2010) Heat stress affects flowering, berry growth, sugar accumulation and photosynthesis of Vitis vinifera, Functional Plant Biology 37:206-214 Roberts JA et al. (2002) Abscission, dehiscence, and other cell separation processes, Annual Review of Plant Biology 53:131-158 Else MA et al. (2004) Stomatal closure in flooded tomato plants involves abscisic acid and a chemically unidentified anti-transpirant in xylem sap, Plant Physiology 135:2215-2224
Heat Induced Abscission Resistance Disposition
A heat-induced summer dormant bud on Vitis vinifera cv. Syrah in the Barossa Valley, Australia, during a five-day heatwave with maximum daily temperatures of 42-45°C, showing cessation of shoot elongation, tightly appressed bud scales, and elevated expression of HSP70 and HSP90 heat shock proteins in meristematic tissues, with growth resumption occurring within one week after temperatures moderate to 30-32°C maximum.
A summer dormant bud that enters growth arrest specifically in response to sustained high temperature stress (typically above 35-38°C for Vitis vinifera), characterized by cessation of cell division in apical meristems, reduced transpiration through partially closed bud scales, accumulation of heat shock proteins, and metabolic shift toward protective rather than growth functions. Heat-induced dormancy represents a thermomorphogenic response protecting meristematic tissues from heat damage, with growth arrest often accompanied by leaf wilting on associated shoots, reduced photosynthetic capacity, and temporary suspension of inflorescence or vegetative primordium development in buds not yet fully differentiated.
Heat-induced summer dormancy is increasingly relevant under climate change scenarios with more frequent extreme heat events. The temperature threshold for heat-induced dormancy varies among cultivars, with some Mediterranean and warm-climate varieties showing greater heat tolerance. Heat-induced dormancy differs from heat damage, where tissues are irreversibly injured dormant buds retain full developmental potential. Concurrent water stress typically exacerbates heat-induced dormancy through synergistic effects on plant water status and stomatal conductance.
heat stress dormant bud
hitzeinduzierte Ruheknospe
temperature-induced dormant bud
thermally dormant bud
Based on Greer & Weedon (2012) "Modelling photosynthetic responses to temperature of grapevine (Vitis vinifera cv. Semillon) leaves on vines grown in a hot climate" Plant, Cell & Environment 35=1050-1064 Carbonell-Bejerano et al. (2014) "Circadian oscillatory transcriptional programs in grapevine ripening fruits" BMC Plant Biology 14=78 Liu et al. (2012) "Global transcriptome analysis reveals comprehensive insights into heat stress responses in grapevine" BMC Genomics 13=654.
Heat Induced Summer Dormant Bud
Heat necrosis quality assessment following an extreme heat event in Swan Valley Western Australia (48°C for 6 hours) documented the extent of irreversible tissue damage across Chardonnay vineyard blocks with varying water status and canopy development. Severely affected vines under deficit irrigation showed extensive heat necrosis quality with complete death of 60% of shoot tips, necrotic lesions covering 35% of young leaf area, and death of exposed cluster rachis causing berry drop on 15% of clusters, requiring substantial regenerative growth and crop load adjustment for recovery. Moderately affected vines with adequate irrigation showed reduced heat necrosis quality with 25% shoot tip death primarily on exposed shoots, 15% young leaf necrosis, and minimal rachis damage, permitting recovery through lateral shoot development without major crop adjustment. Well-protected vines receiving overhead irrigation during the heat event showed minimal heat necrosis quality below 5% shoot tip damage with rapid recovery, demonstrating the effectiveness of evaporative cooling for preventing lethal tissue temperatures. The heat necrosis quality assessment informed recovery management including selective heading of necrotic shoots to stimulate lateral development, removal of damaged fruit to concentrate resources on viable clusters, and adjusted expectations for current and subsequent season production based on the extent of permanent tissue loss.
Heat necrosis quality is a heat damage severity quality that inheres in grapevine tissues and represents the extent of irreversible cell death and tissue collapse resulting from thermal exposure exceeding lethal temperature thresholds, manifesting as discrete necrotic lesions, extensive tissue browning, and organ death that removes affected tissues from productive function with consequences for photosynthetic capacity, carbohydrate supply, and vine recovery requiring regeneration from undamaged meristems. This quality addresses the most severe category of heat damage where cellular repair is impossible and tissue function is permanently lost, distinguishing heat necrosis from sublethal damage that may recover when temperatures moderate. The ontological significance of heat necrosis quality lies in its indication of thermal exposure severity sufficient to cause permanent tissue loss, requiring management response that addresses both the immediate production impact and the long-term vine health implications of substantial necrotic damage that may affect carbohydrate reserves and subsequent season productivity. The biological basis of heat necrosis encompasses the thermal denaturation of cellular proteins, the disruption of membrane integrity causing cellular content leakage, and the collapse of cellular structure that collectively result in cell death when exposure duration at damaging temperatures exceeds the capacity for cellular protection and repair mechanisms. The temperature thresholds for necrosis induction vary among tissues, with actively growing shoot tips and young expanding leaves showing necrosis at lower temperatures (approximately 40-45°C) than mature lignified tissues that may tolerate brief exposure to 50°C or higher without permanent damage. The extent of necrosis following heat events reflects the interaction between peak temperature, exposure duration, tissue vulnerability, and hydration status, with water-stressed tissues showing necrosis at lower temperatures than well-hydrated tissues that benefit from evaporative cooling and greater thermal buffering capacity.
Heat_Necrosis_Quality is_a Heat_Damage_Severity_Quality, representing the most severe damage category involving irreversible cell death. This class addresses permanent tissue loss requiring regeneration rather than repair. Distinguished from recoverable heat damage by the irreversible nature of necrotic tissue. Young, actively growing tissues show lower lethal temperature thresholds than mature tissues. Water status strongly modulates necrosis thresholds through evaporative cooling effects. The extent of necrosis determines recovery timeline and management requirements. Critical for assessing vine health impacts beyond immediate fruit quality concerns.
Hitzenekrose
heat killing
heat-induced tissue death
hitzeinduzierter Zelltod
thermal cell death
thermal necrosis
thermische Nekrose
thermisches Absterben
Wahid A et al. (2007) Heat tolerance in plants: an overview. Environmental and Experimental Botany 61:199-223 Greer DH, Weedon MM (2013) The impact of high temperatures on Vitis vinifera cv. Semillon grapevine performance and berry ripening. Frontiers in Plant Science 4:491 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 453-507
Heat Necrosis Quality
Heat requirement comparison among cultivars following confirmed endodormancy release reveals the genetic variation affecting budbreak timing, with low heat requirement cultivar (Chardonnay, approximately 150 GDD base 10°C from release to budbreak) showing rapid response to spring warming (budbreak 18 days after consistent warming begins), moderate requirement cultivar (Merlot, approximately 220 GDD) showing intermediate response (budbreak 26 days after warming), and high requirement cultivar (Cabernet Sauvignon, approximately 300 GDD) showing delayed response (budbreak 35 days after warming), with the heat requirement differences creating the characteristic cultivar sequence of budbreak timing that enables harvest sequence planning and frost risk distribution across the cultivar portfolio.
Heat requirement quality is an ecodormancy status quality that represents the cumulative exposure to warm temperatures needed to transition ecodormant buds to visible budbreak, characterizing the thermal time threshold that must be accumulated after endodormancy release before growth initiation occurs, with heat requirement representing the second phase of the two-phase dormancy release model where chilling satisfies endodormancy and subsequent heat drives budbreak timing. Within the BFO framework, this quality specifically addresses the thermal threshold dimension of budbreak initiation, distinguishing the heat needed for growth from the chilling needed for dormancy release while recognizing that heat requirement provides a degree of buffering against premature budbreak by requiring sustained warmth rather than responding to brief warm spells, though this buffering is less robust than the protection provided by endodormancy. The ontological significance lies in heat requirement determining the lag between favorable temperatures and visible growth, where higher heat requirements create longer delays between spring warming and budbreak (potentially providing frost protection by delaying vulnerable green tissue emergence), while lower requirements enable more rapid response to warmth (potentially advancing the growing season but increasing frost risk from early budbreak). The genetic variation in heat requirement shows differences among cultivars that interact with chilling requirement to determine overall budbreak behavior, with some cultivars showing high chilling but low heat requirement (late endodormancy release but rapid subsequent budbreak), others showing low chilling but high heat requirement (early responsiveness but slow budbreak progression), and various intermediate combinations that affect the cultivar-specific relationship between winter conditions and budbreak timing. The calculation of heat accumulation for budbreak typically uses growing degree-day concepts with base temperatures around 10°C, though cultivar-specific base temperatures may provide better prediction, and with the starting date for accumulation depending on endodormancy release timing (accumulation only counts from when buds become responsive). Understanding heat requirement quality enables prediction of budbreak timing from temperature forecasts given endodormancy release status, assessment of cultivar-specific budbreak behavior, and evaluation of management practices (such as late pruning) that may modify apparent heat requirements by affecting bud physiological status.
Heat_Requirement_Quality addresses the thermal time threshold for budbreak initiation from ecodormant buds. This quality varies among cultivars and determines responsiveness to spring warming. The class supports budbreak prediction through heat requirement characterization.
Gradtagbedarf
Wärmebedarf
Wärmeschwelle
Wärmesummenanforderung
growing degree-day requirement
heat sum threshold
thermal time requirement
warmth requirement
Parker AK et al. (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17(2): 206-216 Duchêne E et al. (2010) A grapevine (Vitis vinifera L.) deoxy-d-xylulose synthase gene colocates with a major quantitative trait loci for terpenol content. Theoretical and Applied Genetics 120(7): 1451-1461
Heat Requirement Quality
Molecular characterization of Heat_Shock_Protein_Disposition at INRAE Montpellier examined HSP gene expression and protein accumulation across 15 cultivars following controlled heat shock (42°C for 2 hours) and recovery. Expression of representative genes from major HSP families (VvHSP101, VvHSP90, VvHSP70, VvHSP17.9) was quantified by qRT-PCR at 0, 0.5, 1, 2, 4, and 8 hours post-heat onset, while HSP protein levels were assessed by immunoblot. Heat_Shock_Protein_Disposition showed substantial cultivar variation in both induction kinetics and magnitude. 'Grenache' exhibited rapid, strong HSP induction with VvHSP17.9 (small HSP) expression increasing 85-fold within 1 hour, peaking at 150-fold by 2 hours, and VvHSP70 reaching 25-fold induction. Protein accumulation paralleled transcript patterns, with small HSP protein detectable by 2 hours and abundant by 4 hours. 'Chardonnay' showed delayed, weaker Heat_Shock_Protein_Disposition with VvHSP17.9 reaching only 45-fold peak induction at 4 hours (2-hour delay versus 'Grenache'). Correlation analysis revealed that HSP induction magnitude correlated with membrane thermostability (r = 0.68) and photosynthetic recovery (r = 0.72), confirming the protective function of HSP accumulation. Pre-conditioning (mild heat exposure at 38°C) enhanced subsequent Heat_Shock_Protein_Disposition in all cultivars, with 'Grenache' showing 40% faster induction and 25% higher peak expression following pre-conditioning, demonstrating the role of acquired thermotolerance in HSP-mediated protection.
A Heat_Shock_Protein_Disposition is a Heat_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to rapidly synthesize and accumulate heat shock proteins (HSPs) in response to elevated temperatures, providing molecular chaperone protection against heat-induced protein denaturation and aggregation. This disposition represents the protein quality control component of heat tolerance, with HSPs functioning as molecular chaperones that bind to partially denatured proteins, prevent their irreversible aggregation, assist in proper refolding during recovery, and target damaged proteins for degradation when repair is not possible. Heat shock proteins in plants are classified by molecular weight into HSP100, HSP90, HSP70, HSP60, and small HSP (sHSP) families, each with distinct cellular localizations and functions. The heat shock response is orchestrated by heat shock transcription factors (HSFs) that activate HSP gene expression within minutes of temperature elevation, enabling rapid accumulation of protective proteins. Cultivars differ in both basal HSP levels (constitutive protection) and the speed, magnitude, and persistence of induced HSP accumulation following heat exposure. Strong Heat_Shock_Protein_Disposition provides enhanced protection against acute heat stress through rapid chaperone accumulation and improved recovery through efficient protein repair, with particular importance of small HSPs for protecting photosynthetic proteins and membranes in chloroplasts.
Heat_Shock_Protein_Disposition is_a Heat_Tolerance_Disposition. HSPs are molecular chaperones protecting proteins from heat-induced denaturation and aggregation. Multiple HSP families exist with distinct functions and localizations. The heat shock response is transcriptionally regulated by HSF transcription factors. Both constitutive HSP levels and induced accumulation contribute to heat tolerance. Small HSPs (sHSPs) are particularly important for protecting photosynthetic apparatus. Heat acclimation enhances subsequent HSP response (acquired thermotolerance).
HSP accumulation capacity
HSP-Akkumulations-Kapazität
Hitzeschockprotein-Disposition
Molekulare-Chaperon-Induktion
heat shock response
molecular chaperone induction
Wang W et al. (2004) Role of plant heat-shock proteins and molecular chaperones in the abiotic stress response, Trends in Plant Science 9:244-252 Vierling E (1991) The roles of heat shock proteins in plants, Annual Review of Plant Physiology and Plant Molecular Biology 42:579-620 Liu GT et al. (2012) Differential proteomic analysis of grapevine leaves by iTRAQ reveals responses to heat stress and subsequent recovery, BMC Plant Biology 12:174
Heat Shock Protein Disposition
Heat shock protein expression quality profiling across grapevine cultivars during controlled heat stress identified molecular markers of thermotolerance variation relevant to cultivar selection for warming viticultural regions. Western blot quantification of HSP accumulation following 40°C exposure for 4 hours revealed substantial variation in heat shock protein expression quality among cultivars. Touriga Nacional demonstrated excellent heat shock protein expression quality with HSP70 levels increasing 35-fold above baseline, HSP101 accumulating to 28-fold, and small HSP17.6 reaching 45-fold induction, with the combined HSP pool estimated to represent 3.5% of total soluble protein at peak accumulation. Riesling showed moderate heat shock protein expression quality with HSP70 at 18-fold, HSP101 at 12-fold, and sHSP17.6 at 22-fold induction, totaling approximately 1.8% of soluble protein. Pinot Noir exhibited poor heat shock protein expression quality with HSP70 only 8-fold, HSP101 at 5-fold, and sHSP17.6 at 12-fold, representing only 0.9% of total protein. The heat shock protein expression quality ranking correlated strongly (r²=0.84) with thermotolerance assessed by electrolyte leakage and photosynthetic recovery, establishing HSP expression capacity as a reliable predictor of cultivar heat tolerance and a potential selection criterion for heat-adapted variety development.
Heat shock protein expression quality is a heat shock response quality that inheres in a grapevine organism and represents the capacity to rapidly synthesize and accumulate heat shock proteins (HSPs) following exposure to elevated temperatures, with these molecular chaperones functioning to prevent heat-induced protein aggregation, assist in refolding of partially denatured proteins, and protect essential cellular structures from thermal damage through direct interaction with vulnerable protein substrates. This quality addresses the central protective mechanism of the heat shock response, recognizing that heat shock proteins constitute the primary molecular defense against protein denaturation that underlies most forms of heat injury, with their rapid induction enabling acquisition of enhanced thermotolerance within hours of stress perception. The ontological significance of heat shock protein expression quality lies in its role as the key molecular determinant of acquired thermotolerance, with the speed, magnitude, and duration of HSP accumulation strongly correlating with the degree of protection achieved during and following heat exposure, and with substantial genetic variation in HSP expression capacity providing targets for selection and breeding to improve heat tolerance. The biological basis of heat shock protein expression encompasses the transcriptional activation of HSP genes through heat shock factor binding to conserved promoter elements, the translational prioritization of HSP mRNAs through mechanisms that favor their translation when general protein synthesis is inhibited by heat, and the accumulation of functional HSP proteins in cellular compartments where they provide chaperone protection for vulnerable substrates. The major HSP families relevant to grapevine heat tolerance include HSP100 providing disaggregase activity that can rescue already-aggregated proteins, HSP70 providing the primary chaperone function preventing aggregation and assisting refolding, HSP90 maintaining client proteins in activation-competent states, and small HSPs that bind denatured proteins preventing irreversible aggregation until ATP-dependent chaperones can facilitate refolding. The expression patterns of different HSP classes show characteristic differences in timing, with small HSPs typically induced most rapidly and abundantly while larger HSPs show more moderate but sustained expression, collectively providing overlapping protection windows that extend thermotolerance across the stress and recovery periods.
Heat_Shock_Protein_Expression_Quality is_a Heat_Shock_Response_Quality, representing the central molecular chaperone component of heat protection. This class addresses the synthesis and accumulation of HSP families (HSP100, HSP90, HSP70, sHSPs) providing protein protection. Fold-induction from baseline and percentage of total protein provide quantification metrics. The HSP expression ranking correlates with thermotolerance, enabling molecular screening for heat tolerance. Different HSP classes provide distinct functions (disaggregation, folding assistance, aggregation prevention). Distinguished from Antioxidant_Response_Quality which addresses ROS management rather than protein protection.
HSP accumulation
HSP expression
HSP-Akkumulation
HSP-Induktion
Hitzeschockprotein-Expression
Molekularchaperon-Antwort
heat shock protein induction
molecular chaperone response
Wang W et al. (2004) Role of plant heat-shock proteins and molecular chaperones in the abiotic stress response. Trends in Plant Science 9:244-252 Liu GT et al. (2012) Differential proteomic analysis of grapevine leaves by iTRAQ reveals responses to heat stress and subsequent recovery. BMC Plant Biology 12:174 Carvalho LC et al. (2015) Heat stress in grapevine: the pros and cons of acclimation. Plant, Cell and Environment 38:777-789
Heat Shock Protein Expression Quality
Heat shock response quality characterization in Chardonnay evaluated the molecular basis of acquired thermotolerance relevant to heat wave management in warm climate viticulture. Controlled heat exposure at 40°C triggered rapid heat shock response quality activation with heat shock transcription factor HsfA2 showing nuclear localization within 15 minutes, HSP70 transcript levels increasing 25-fold within 2 hours, and HSP101 protein accumulating to protective concentrations within 4 hours. The functional significance of heat shock response quality was demonstrated by comparing thermotolerance in acclimated versus non-acclimated tissues, with leaves pre-exposed to moderate heat (38°C for 3 hours) showing 70% reduction in membrane damage (electrolyte leakage) and 85% preservation of photosynthetic function (Fv/Fm) during subsequent severe heat exposure (45°C for 2 hours) compared to non-acclimated controls. The heat shock response quality persisted for approximately 48 hours following initial induction, with protection declining as heat shock protein levels decreased through normal turnover. Cultivar comparison revealed that Grenache mounted a faster heat shock response quality with equivalent HSP70 levels achieved in 90 minutes versus 3 hours in Chardonnay, correlating with the superior field heat tolerance observed in Grenache and suggesting that heat shock response speed contributes to natural variation in cultivar thermotolerance.
Heat shock response quality is a thermal tolerance quality that inheres in a grapevine organism and represents the capacity to mount a coordinated molecular defense response upon exposure to elevated temperatures, encompassing the rapid induction of heat shock protein synthesis, activation of antioxidant defense systems, and deployment of other protective mechanisms that collectively enhance cellular thermotolerance and enable survival of heat exposures that would otherwise cause irreversible damage. This quality captures the inducible dimension of heat tolerance that supplements constitutive thermostability, recognizing that the heat shock response provides a rapid adaptive mechanism enabling vines to acquire enhanced tolerance within hours of heat exposure through synthesis of protective molecules that were not present before stress onset. The ontological significance of heat shock response quality lies in its role as the primary mechanism for acquired thermotolerance, enabling acclimation to gradually increasing temperatures that permits survival of subsequent severe heat events that would be lethal to non-acclimated tissues, with substantial variation among grapevine genotypes in the speed, magnitude, and effectiveness of heat shock response providing a basis for understanding and potentially enhancing adaptive heat tolerance capacity. The biological basis of heat shock response encompasses the activation of heat shock transcription factors that bind to conserved heat shock elements in promoter regions of heat-responsive genes, triggering rapid transcription of heat shock proteins serving as molecular chaperones that prevent protein aggregation and assist refolding of partially denatured proteins, combined with activation of antioxidant systems neutralizing reactive oxygen species generated under heat stress. The temporal dynamics of heat shock response show characteristic patterns with transcription factors activated within minutes of temperature elevation, heat shock protein mRNA accumulating within 30-60 minutes, and protective protein levels reaching maximum within 2-6 hours depending on the severity of stress and the specific response component. The relationship between heat shock response quality and practical heat tolerance involves the protection conferred during the stress event itself combined with the residual protection that persists for hours to days following heat shock response activation, providing a window of enhanced tolerance that affects the outcome of subsequent heat exposures.
Heat_Shock_Response_Quality is_a Thermal_Tolerance_Quality, serving as parent class for Heat_Shock_Protein_Expression_Quality, Antioxidant_Response_Quality, and Reactive_Oxygen_Species_Scavenging_Quality which address specific components of the molecular heat response. This class addresses the overall inducible heat protection system while child classes capture particular mechanisms. The temporal dynamics (minutes to hours) distinguish heat shock response from constitutive tolerance mechanisms. Acquired thermotolerance through pre-exposure represents key practical application. Critical for understanding molecular basis of heat acclimation and cultivar variation in heat tolerance.
Hitzeakklimatisierungsreaktion
Hitzeschock-Reaktion
Hitzestress-Antwort
heat acclimation response
heat shock induction
heat stress response
thermal stress response
thermische Stressreaktion
Wang W et al. (2004) Role of plant heat-shock proteins and molecular chaperones in the abiotic stress response. Trends in Plant Science 9:244-252 Carvalho LC et al. (2015) Heat stress in grapevine: the pros and cons of acclimation. Plant, Cell and Environment 38:777-789 Liu GT et al. (2012) Differential proteomic analysis of grapevine leaves by iTRAQ reveals responses to heat stress and subsequent recovery. BMC Plant Biology 12:174
Heat Shock Response Quality
Heat stress indicator quality assessment during an extreme heat event in Paso Robles (5 consecutive days exceeding 42°C) employed multiple diagnostic approaches to evaluate Cabernet Sauvignon vine status and fruit damage. Visual heat stress indicator quality assessment identified sunburn damage affecting 35% of exposed clusters on southwest-facing rows, leaf scorch on 20% of canopy leaves particularly at shoot tips, and heat-induced berry shrivel on clusters with prolonged afternoon sun exposure. Physiological measurements revealed chlorophyll fluorescence Fv/Fm ratios declining from normal 0.82 to stress-indicating 0.65 at midday during the heat event, confirming photosynthetic apparatus damage consistent with visual symptoms. Membrane integrity assessment through electrolyte leakage showed 45% leakage in heat-exposed leaves versus 15% in shaded leaves, quantifying the cellular damage underlying visible symptoms. The integrated heat stress indicator quality evaluation established that the event had exceeded moderate stress thresholds, causing permanent damage to exposed fruit and leaves that would impact wine quality and require management response including selective harvest of damaged fruit separate from protected clusters.
Heat stress indicator quality is a stress response quality that inheres in a grapevine organism and represents the ensemble of physiological, biochemical, and morphological markers that signal the presence and severity of thermal stress arising from exposure to temperatures exceeding the optimal range for grapevine function, providing diagnostic information for assessing vine heat stress status and guiding management decisions aimed at mitigating temperature-induced damage to vegetative tissues, reproductive structures, and fruit quality during episodes of extreme heat increasingly common under climate change conditions. This quality encompasses both the visible damage symptoms indicating heat injury has occurred and the underlying physiological tolerance mechanisms that determine the temperature thresholds and durations causing damage in specific cultivar-environment combinations. The ontological significance of heat stress indicator quality lies in its role enabling assessment of vine thermal status and vulnerability during heat events, supporting decisions about protective interventions including irrigation for evaporative cooling, shade cloth deployment, or harvest timing adjustments that may mitigate heat impacts on fruit quality and vine health. The biological basis of heat stress indicators encompasses the progressive dysfunction of cellular processes as temperatures rise beyond optimal ranges, beginning with reduced photosynthetic efficiency and altered metabolism at moderately elevated temperatures, progressing through membrane disruption and protein denaturation at higher temperatures, and culminating in irreversible cellular damage and tissue necrosis when thermal tolerance thresholds are exceeded. The temperature thresholds triggering heat stress responses vary among grapevine cultivars reflecting genetic variation in thermal tolerance mechanisms, and are modulated by acclimation status, water availability, and the duration and pattern of heat exposure that collectively determine whether specific temperature conditions cause detectable stress responses or damage. The increasing frequency and intensity of heat events under climate change has elevated the importance of heat stress indicator quality assessment for viticultural adaptation, motivating research into cultivar thermal tolerance variation and management strategies that reduce heat exposure or enhance tolerance during critical developmental periods when heat impacts on fruit quality are most pronounced.
Heat_Stress_Indicator_Quality is_a Stress_Response_Quality, serving as parent class for Heat_Damage_Severity_Quality and Thermal_Tolerance_Quality which address visible damage symptoms and underlying tolerance mechanisms respectively. This class captures the overall concept of heat stress assessment while child hierarchies address specific dimensions. Distinguished from Drought_Stress_Indicator_Quality by the focus on temperature rather than water deficit, though the two stresses frequently co-occur and interact. Climate change relevance increasingly important for viticultural adaptation.
Hitzestress-Indikatoren
Hitzestress-Symptome
Temperaturstress-Reaktion
heat damage markers
heat stress symptoms
temperature stress response
thermal stress indicators
thermische Stressmarker
Greer DH, Weston C (2010) Heat stress affects flowering, berry growth, sugar accumulation and photosynthesis of Vitis vinifera cv. Semillon grapevines grown in a controlled environment. Functional Plant Biology 37:206-214 Caravia L et al. (2016) Application of shade treatments during Shiraz berry ripening to reduce the impact of high temperature. Australian Journal of Grape and Wine Research 22:422-437 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 453-507
Heat Stress Indicator Quality
The heat summation data for a Cabernet Sauvignon vineyard in the Oakville AVA (Napa Valley, California) documents the interphenological thermal time requirements — budburst to flowering: 380 GDD base 10°C accumulated over approximately 65 days (March 15 to May 20), flowering to véraison: 550 GDD accumulated over approximately 55 days (May 20 to July 15), véraison to harvest: 450 GDD accumulated over approximately 50 days (July 15 to September 5), total budburst to harvest: 1,380 GDD over approximately 170 days. This interphenological breakdown reveals that the flowering-to-véraison period consumes the most heat units per day (10 GDD/day average, coinciding with the warmest months) and is the most compressed period, while the véraison-to-harvest period requires moderate heat (9 GDD/day) and its duration is the most variable between vintages (ranging from 40 to 65 days depending on August-September temperatures) — this final period variability is the primary source of vintage variation in Cabernet Sauvignon harvest date and fruit composition.
A temperature data component that records the cumulative thermal energy available for vine development over specific time periods, documenting the heat accumulation indices (GDD base 10°C, Winkler, Huglin, BEDD) and interphenological heat unit requirements (budburst-to-flowering, flowering-to-véraison, véraison-to-harvest thermal times) that quantify the thermal resources consumed during each phase of vine development and enable the prediction of phenological timing, cultivar maturity potential, and seasonal developmental progress from temperature data.
Heat_Summation_Data extends the basic GDD concept by documenting the thermal accumulation across multiple time scales — the full growing season (captured by the GDD, Winkler, Huglin, and BEDD indices documented in the Growing_Season_Data subclasses), specific interphenological periods (budburst-to-flowering, flowering-to-véraison, véraison-to-harvest), and individual months — providing a temporal resolution of thermal resource documentation that enables analysis of which growth phases receive adequate or inadequate heat in a given climate. The interphenological thermal time concept recognizes that each developmental phase has a characteristic heat requirement that is relatively consistent for a given cultivar across sites and years — Cabernet Sauvignon typically requires 350-400 GDD from budburst to flowering regardless of whether this accumulation occurs over 55 days (warm site) or 80 days (cool site): this consistency enables the thermal time framework to predict phenological timing from temperature forecasts (by calculating the date at which the accumulated GDD will reach the required thermal time for each event) and to assess whether a given climate provides sufficient total heat for the complete budburst-to-harvest sequence within the available frost-free season. The practical application of interphenological heat summation is particularly valuable for harvest date prediction — monitoring GDD accumulation from véraison and projecting forward using long-range temperature forecasts provides a 3-6 week advance estimate of harvest date that enables logistical planning for labor, equipment, winery capacity, and the sequential harvesting of multiple cultivars.
Wärmesummendaten
cumulative temperature data
heat unit documentation
kumulative Temperaturdaten
thermal accumulation record
thermische Akkumulation
Gladstones J (1992) Viticulture and Environment, Winetitles Jones GV and Davis RE (2000) Climate influences on grapevine phenology, American Journal of Enology and Viticulture 51: 249-261 Parker AK et al. (2011) General phenological model to characterise the timing of flowering and véraison of Vitis vinifera L., Australian Journal of Grape and Wine Research 17: 206-216 Duchêne E and Schneider C (2005) Grapevine and climatic changes, Agronomy for Sustainable Development 25: 93-99
Heat Summation Data
Comprehensive characterization of Heat_Tolerance_Disposition at CSIRO Agriculture, Adelaide, assessed multiple heat tolerance parameters across 30 cultivars under controlled heat stress (40°C for 4 hours) and recovery conditions. Measurements included chlorophyll fluorescence (Fv/Fm as PSII efficiency indicator), membrane stability (electrolyte leakage), leaf gas exchange (photosynthesis, stomatal conductance), and visual damage scoring. Cultivars showed substantial variation in Heat_Tolerance_Disposition. 'Grenache' (Mediterranean origin) exhibited strong heat tolerance with Fv/Fm declining only 12% during stress (from 0.82 to 0.72) and recovering fully within 24 hours, electrolyte leakage increasing just 15%, and photosynthesis maintaining 65% of pre-stress rate during heat exposure. 'Pinot Noir' (cool-climate origin) showed poor Heat_Tolerance_Disposition with Fv/Fm declining 35% (to 0.53), incomplete recovery (reaching 0.71 at 48 hours), electrolyte leakage increasing 45%, and photosynthesis reduced to 25% of pre-stress rate. The Australian-bred 'Tarrango' (heat-adapted background) demonstrated exceptional tolerance with only 8% Fv/Fm decline, 10% leakage increase, and 75% photosynthesis maintenance. Principal component analysis revealed that membrane thermostability and PSII heat tolerance were the strongest contributors to overall Heat_Tolerance_Disposition, suggesting these mechanisms as priority breeding targets for heat-resilient cultivars.
A Heat_Tolerance_Disposition is an Abiotic_Stress_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through physiological, biochemical, and morphological mechanisms that enable maintenance of cellular function, photosynthetic activity, and reproductive success under high temperature conditions that would otherwise cause metabolic disruption, tissue damage, or crop loss. This disposition represents the capacity to tolerate supraoptimal temperatures, with grapevine showing reduced photosynthesis above 30-35°C and potential for heat damage above 40°C depending on duration, humidity, and genotype. Heat stress in viticulture is increasingly important under climate change scenarios, with documented impacts including photosynthetic inhibition (stomatal closure, Rubisco deactivation, thylakoid damage), membrane destabilization (increased fluidity, ion leakage), protein denaturation, accelerated respiration depleting carbohydrate reserves, disrupted phenolic and aromatic compound biosynthesis affecting wine quality, and reproductive failure (pollen sterility, fruit abscission, sunburn). Heat tolerance mechanisms encompass maintenance of photosynthetic function at elevated temperatures (Rubisco activase thermostability, PSII heat tolerance), membrane stabilization (lipid saturation, protective proteins), heat shock protein accumulation (molecular chaperones preventing protein aggregation), evaporative cooling capacity (transpirational heat dissipation), and tissue-specific heat protection (berry sunburn resistance, flower heat tolerance). Heat_Tolerance_Disposition varies among Vitis species and cultivars, with Mediterranean and hot-climate cultivars generally showing enhanced heat adaptation compared to cool-climate genotypes, providing genetic resources for breeding heat-resilient cultivars.
Heat_Tolerance_Disposition is_a Abiotic_Stress_Tolerance_Disposition. Heat tolerance is increasingly important under climate change and encompasses multiple mechanisms addressed through child classes including Photosynthesis_Heat_Stability_Disposition, Membrane_Thermostability_Disposition, Heat_Shock_Protein_Disposition, Evaporative_Cooling_Disposition, and Berry_Heat_Tolerance_Disposition. Heat tolerance is partially independent of drought tolerance, though both stresses often co-occur and share some adaptive mechanisms. Mediterranean and hot-climate cultivars provide genetic resources for heat tolerance improvement.
Hitzetoleranz-Disposition
Hochtemperatur-Toleranz
Thermotoleranz-Kapazität
heat stress resistance
high temperature tolerance
thermal tolerance
Wahid A et al. (2007) Heat tolerance in plants: an overview, Environmental and Experimental Botany 61:199-223 Greer DH, Weston C (2010) Heat stress affects flowering, berry growth, sugar accumulation and photosynthesis of Vitis vinifera cv. Semillon grapevines grown in a controlled environment, Functional Plant Biology 37:206-214 Carvalho LC et al. (2015) Heat stress in grapevine: the pros and cons of acclimation, Plant, Cell and Environment 38:777-789
Heat Tolerance Disposition
Ramsey (Salt Creek, V. champinii) demonstrates excellent heat tolerance supporting table grape production in the extreme heat of the Coachella Valley, California, where summer maximum temperatures regularly exceed 45°C and soil surface temperatures reach 50-55°C. 1103 Paulsen (V. berlandieri × V. rupestris) shows high heat tolerance maintaining Primitivo and Negroamaro productivity in the hot summers of Puglia (maximum temperatures 38-42°C) where heat stress compounds drought stress effects. 140 Ruggeri (V. berlandieri × V. rupestris) provides heat tolerance for Nero d'Avola in Sicily under summer temperatures exceeding 40°C, maintaining photosynthetic function through combined heat and drought stress periods.
An abiotic stress-tolerant rootstock role that inheres in a grafted rootstock by virtue of its capacity to maintain scion productivity under high temperature conditions that would cause heat stress, reduced photosynthesis, and yield loss on less adapted rootstocks. Heat-tolerant rootstocks sustain root function and scion support during prolonged exposure to soil temperatures exceeding 30°C and air temperatures above 35-40°C, through mechanisms including thermostable membrane lipid composition, enhanced heat shock protein expression, efficient respiratory maintenance at elevated temperatures, and sustained hydraulic conductance that supports transpirational cooling of the canopy. Heat tolerance is quantified by yield stability and fruit quality maintenance across hot growing seasons, by photosynthetic rate and stomatal conductance under heat stress, and by absence of heat-induced physiological disorders. Heat tolerance often co-occurs with drought tolerance, as both stresses involve high evaporative demand and are exacerbated by water limitation.
Heat_Tolerant_Rootstock_Role addresses an abiotic stress of increasing importance under climate change, with viticultural regions worldwide experiencing more frequent and severe heat events. V. berlandieri × V. rupestris rootstocks (1103P, 140Ru, 110R) and V. champinii selections (Ramsey, Dog Ridge) show the highest heat tolerance, reflecting parental adaptation to the hot summers of Texas and the American Southwest. Heat tolerance is difficult to separate from drought tolerance experimentally, as heat stress typically occurs under high evaporative demand conditions most heat-tolerant rootstocks also show drought tolerance. Root zone temperature affects nutrient uptake, particularly of potassium and magnesium, creating indirect heat effects on fruit composition.
heat resistant rootstock
heat stress tolerant rootstock
heat tolerant rootstock
high temperature tolerant rootstock
hitzetolerante Unterlage
temperaturstresstolerante Unterlage
wärmetolerante Unterlage
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 335-340 Ferrara G et al. (2015) Influence of rootstock on physiological response of Primitivo grapevines under different irrigation regimes, Acta Horticulturae 1099: 191-198 Corso M, Bonghi C (2014) Grapevine rootstock effects on abiotic stress tolerance, Plant Science 228: 1-14 Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22
Heat Tolerant Rootstock Role
A Pinot Gris selection bears a Heat_Treated_Mother_Plant_Role after 90 days at 38°C in the FPS thermotherapy facility, with post-treatment testing confirming elimination of previously detected GFLV while GLRaV-3 persisted (requiring subsequent meristem culture for complete elimination). A Müller-Thurgau vine serves in a Heat_Treated_Mother_Plant_Role following standard German (JKI) thermotherapy protocols of 60 days at 37°C, successfully eliminating GVA and GVB detected in the original selection. A Tempranillo candidate clone from Rioja achieved Heat_Treated_Mother_Plant_Role status through the Spanish IMIDRA program, with thermotherapy eliminating multiple nepoviruses while preserving the clonal characteristics selected for agronomic performance.
A mother plant role that inheres in a grapevine that has undergone thermotherapy treatment designed to eliminate heat-sensitive pathogens (particularly viruses and phytoplasmas) through extended exposure to elevated temperatures that inhibit pathogen replication while the plant survives and regenerates. Heat-treated mother plants have typically been maintained at 37-38°C for extended periods (typically 60-120 days) under controlled growth chamber conditions, allowing meristematic tissues to outgrow systemic pathogen distribution as viral replication is suppressed by heat stress. Thermotherapy was the first successful virus elimination technique for grapevines (developed 1950s-1960s) and remains fundamental to clean stock programs, often combined with meristem tip culture (shoot tip excision from heat-treated plants) to maximize elimination success. Heat treatment effectiveness varies by pathogen—highly effective for some viruses (GFLV, nepoviruses), moderately effective for others (leafroll viruses), and limited for heat-tolerant agents—requiring post-treatment testing to confirm elimination success and identify pathogens requiring alternative treatments.
Heat_Treated_Mother_Plant_Role captures the procedural history indicating that thermotherapy has been applied, regardless of elimination success for all targeted pathogens. Thermotherapy remains a cornerstone of virus elimination despite being labor-intensive and requiring specialized facilities—the technique's relative simplicity (no tissue culture expertise required) and effectiveness for many important pathogens ensures continued use. Treatment parameters (temperature, duration) balance pathogen elimination against plant survival temperatures above 38°C may improve elimination rates but increase plant mortality, while shorter durations reduce stress but may permit pathogen survival. Combination protocols (thermotherapy followed by meristem culture, or chemotherapy combined with heat treatment) have improved elimination success for difficult pathogens like leafroll viruses that show only partial sensitivity to heat alone.
heat-therapy vine
heat-treated vine
hitzebehandelte Mutterpflanze
thermotherapierte Rebe
thermotherapy-treated plant
wärmebehandelte Mutterpflanze
Nyland G, Goheen AC (1969) Heat therapy of virus diseases of perennial plants, Annual Review of Phytopathology 7: 331-354 Panattoni A et al. (2013) Grapevine virus elimination by thermotherapy, Journal of Plant Pathology 95: 229-235 Leonhardt W et al. (1998) Methods for elimination of grapevine viruses, Mitteilungen Klosterneuburg 48: 1-10 Maliogka VI et al. (2015) Grapevine virus diseases, APS Press
Heat Treated Mother Plant Role
In Chardonnay vines in Champagne, the heat unit requirement for budbreak is approximately 120-140 growing degree days above the 10°C base temperature, with this threshold consistently predicting budbreak timing within approximately 5 days accuracy when applied to accumulated GDD data, enabling vineyard managers to anticipate budbreak in late March to early April and plan frost protection deployment and early season spray programs accordingly.
Heat unit requirement in Vitis vinifera represents the cultivar-specific threshold of accumulated thermal exposure that must be achieved for budbreak initiation, defining the total heat accumulation above the base temperature necessary to complete the transition from ecodormant bud to actively growing shoot. This requirement represents a genetically determined characteristic that varies among cultivars and contributes to the differences in budbreak timing observed among genotypes grown in the same environment. The quantification of heat unit requirement uses accumulated growing degree days or equivalent thermal units, with cultivar-specific thresholds typically ranging from approximately 100 GDD for early-budbreak cultivars to 300 GDD or more for late-budbreak cultivars when calculated using a 10°C base temperature. The physiological meaning of the heat unit requirement reflects the total thermal exposure needed to complete all preparatory processes for growth initiation, including metabolic activation to levels supporting biosynthesis, accumulation of growth-related gene products and proteins, and cellular preparations for division and expansion in meristematic tissues. The determination of heat unit requirements for specific cultivars involves experimental observation of budbreak timing in relation to measured temperature accumulation, with requirements established through correlation of GDD accumulation at the time of budbreak across multiple seasons and locations. The stability of heat unit requirements enables their use in predictive models, with cultivar-specific thresholds providing reasonably consistent budbreak timing predictions when applied to temperature accumulation data, though some variation occurs due to interactions with prior chilling adequacy, vine physiological status, and local microclimate factors. The interaction between heat unit requirement and chilling completion affects budbreak characteristics, with the heat requirement being most relevant following complete chilling satisfaction, while inadequate chilling may increase the apparent heat requirement or produce erratic response that deviates from model predictions. The practical application of heat unit requirements supports budbreak timing prediction for vineyard management, with threshold values for common cultivars available in viticultural literature and decision support systems that enable planning of early season operations relative to expected phenological development.
Heat_Unit_Requirement represents the threshold parameter within Heat_Accumulation_Response defining cultivar-specific budbreak thermal needs. Distinguished from Heat_Accumulation_Response process by representing the specific threshold value rather than the response mechanism. Cultivar requirements typically range from 100-300 GDD with 10°C base. Genetically determined with reasonable stability across environments. Interaction with chilling adequacy affects requirement expression. Enables predictive modeling of budbreak timing for management planning.
GDD requirement
GDD-Anforderung
Wärmeakkumulations-Schwelle
Wärmeeinheiten-Bedarf
budbreak heat threshold
heat accumulation threshold
thermal requirement for budbreak
thermische Anforderung für Austrieb
Moncur MW et al (1989) Base temperatures for budbreak and leaf appearance of grapevines. American Journal of Enology and Viticulture 40: 21-26 Parker AK et al (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17: 206-216 Duchêne E, Schneider C (2005) Grapevine and climatic changes: a glance at the situation in Alsace. Agronomy for Sustainable Development 25: 93-99
Heat Unit Requirement
Evaluation of hedging response quality in Sonoma County Chardonnay compared regrowth following single versus double-sided hedging at standardized canopy width targets. Single-sided hedging removing only east-facing lateral growth produced localized regrowth with mean 2.4 new laterals per hedged node averaging 15 cm at harvest. Double-sided hedging produced stronger hedging response quality with mean 3.8 laterals per cut node averaging 22 cm, attributed to greater hormonal disruption and light penetration stimulating bud activation across the entire canopy. Implementation of alternating single-sided hedging on two-week intervals achieved equivalent canopy control with 35% less total regrowth compared to single aggressive double-sided hedging, demonstrating how strategic hedging approaches can manage hedging response quality.
Hedging response quality is a vigor manipulation response quality that inheres in a grapevine organism and represents the characteristic pattern and intensity of compensatory growth exhibited following mechanical or manual hedging operations that remove both shoot apices and variable amounts of lateral shoot tissue along the canopy sides to maintain prescribed canopy dimensions, improve light penetration, and facilitate equipment passage through vineyard rows. This quality encompasses the complex regrowth dynamics following more extensive tissue removal than simple topping, involving activation of latent buds across multiple canopy positions and reorganization of source-sink relationships following substantial reduction in photosynthetic leaf area. The ontological significance of hedging response quality lies in its role as a determinant of hedging operation efficiency, wherein varieties and sites with strong hedging responses may require multiple passes to maintain canopy targets while those with moderate responses achieve stable canopy architecture with single interventions. The physiological basis of hedging response involves release of multiple lateral buds from apical dominance inhibition simultaneously, combined with compensatory upregulation of photosynthetic capacity in remaining leaves to support regrowth, and potential mobilization of carbohydrate reserves from woody storage tissues when leaf area removal is severe. Hedging severity significantly modulates response magnitude, with light hedging removing only shoot tips provoking responses similar to topping while aggressive hedging removing substantial leaf area can either suppress regrowth through resource limitation or paradoxically stimulate vigorous response as the vine attempts to restore photosynthetic capacity. The timing of hedging relative to phenological development and carbohydrate status critically influences hedging response quality, with early-season hedging during active growth typically producing strongest responses while post-véraison hedging when vegetative growth has naturally diminished yields more stable outcomes. Repeated hedging within a season creates cumulative effects wherein successive interventions may either progressively exhaust regrowth potential through reserve depletion or trigger increasingly vigorous compensatory responses in high-vigor situations where abundant resources continue to support growth. The spatial pattern of hedging response across the canopy varies with bud position, light exposure history, and proximity to cut surfaces, with most rapid regrowth typically occurring from well-exposed nodes immediately adjacent to cutting planes where both bud development and resource access favor rapid activation. Understanding cultivar-specific hedging response quality patterns enables optimization of hedging timing, frequency, and integration with other canopy management operations to achieve target canopy architecture efficiently.
Hedging_Response_Quality is_a Vigor_Manipulation_Response_Quality, representing the response to mechanical or manual canopy trimming operations that remove both apices and lateral shoots. Distinguished from Topping_Response_Quality by the involvement of lateral shoot tissue and typically greater total leaf area removal. The severity and timing of hedging, plus single versus repeated operations, significantly influence response characteristics. This class relates to mechanical harvesting requirements where canopy dimension control is essential for machine efficiency.
Heckenschnittreaktion
Laubschnittreaktion
Maschinenschnitt-Reaktion
Seitenschnitt-Antwort
canopy trimming response
mechanical hedging response
side trimming compensation
summer hedging regrowth
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 74-79 Poni S et al. (2016) Grapevine quality: a multiple choice issue. Scientia Horticulturae 234:445-462 Terry DB, Kurtural SK (2011) Achieving vine balance of Syrah with mechanical canopy management and regulated deficit irrigation. American Journal of Enology and Viticulture 62:426-437
Hedging Response Quality
The hedging specification for Merlot on VSP trellis in Saint-Émilion prescribes two mechanical hedging passes — first pass at E-L 29-31 (pea size, shoots extending 20-30 cm above the top wire) cutting at 15 cm above the top wire to a total canopy height of 1.7 m, and second pass at E-L 33-35 (pre-véraison) trimming regrowth to the same height with simultaneous lateral trimming on both canopy sides to maintain a wall width of 30-40 cm — the hedging machine (reciprocating cutter bar type) operates at 3-5 km/h with a work rate of 1.5-2.0 ha/h, and the specification explicitly prohibits hedging after véraison to avoid removing mature leaves whose photosynthetic output is critical for sugar accumulation during the ripening period.
A canopy management description that documents the mechanical or manual trimming of shoot tips and lateral shoots that extend beyond the defined canopy boundaries, specifying the timing, cutting height, number of passes, and target canopy dimensions that control vegetative growth, redirect assimilate from shoot elongation to fruit ripening, and maintain the canopy within the physical limits of the trellis infrastructure.
Hedging_Specification addresses the management of vegetative shoot growth that continues beyond the canopy boundaries defined by the trellis system. In vigorous vineyards, indeterminate shoot growth produces 2-4 m of annual extension that would far exceed the trellis height (typically 1.5-2.0 m) without intervention, creating a disorganized, shaded canopy with poor air circulation and high disease pressure: hedging redirects the vine's growth resources from continued shoot elongation (a vegetative sink) toward fruit ripening (the reproductive sink), though this redirection is temporary as hedging stimulates lateral shoot growth from axillary buds below the cut. The timing of hedging relative to phenological stage is critical — hedging before flowering can reduce fruit set by removing photosynthetically active shoot tips during the carbohydrate-critical flowering period, while hedging during the post-set to pre-véraison window (E-L 27-35) achieves the desired growth control with minimal impact on fruit development. Repeated severe hedging (>3 passes per season) reduces total vine leaf area and can delay ripening, necessitating careful balancing of canopy containment against photosynthetic capacity requirements.
Gipfelungs-Spezifikation
Heckenschnitt-Protokoll
Laubschnitt-Beschreibung
canopy topping protocol
shoot trimming specification
summer hedging description
Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles pp: 105-120 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 370-378 Poni S and Giachino E (2000) Growth, photosynthesis and cropping of potted grapevines (Vitis vinifera L. cv. Cabernet Sauvignon) in relation to shoot trimming, Australian Journal of Grape and Wine Research 6: 216-226
Hedging Specification
The Sangiovese selections maintained by multi-generational Chianti Classico families bear Heirloom_Clone_Roles, perpetuated through sélection massale from pre-phylloxera replanting through the 20th century—valued for site adaptation and traditional wine character developed across generations of family stewardship. The Riesling selections of historic Mosel estates (Prüm, Loosen, Thanisch) serve in Heirloom_Clone_Roles, material passed across generations maintaining traditional vineyard character distinct from nursery-supplied registered clones. The "old California" Zinfandel selections from 19th-century Italian immigrant plantings bear Heirloom_Clone_Roles in Sonoma and Lodi, representing generational perpetuation of selections brought from Italy and maintained through family viticulture traditions.
A heritage clone role that inheres in a grapevine clone perpetuated within a family, estate, or community across multiple generations through informal selection and propagation, representing accumulated local adaptation and traditional practices outside formal certification systems. Heirloom clones embody the localized mass selection that characterized viticulture before scientific clone programs—vignerons collecting cuttings from their best vines across generations, creating populations adapted to specific sites and carrying the selection wisdom of successive farming generations. The heirloom designation emphasizes continuity of stewardship rather than documented historical provenance material may lack specific historical documentation while representing unbroken generational perpetuation. Heirloom clones often exist as population mixtures (mass selections) rather than uniform genotypes, with producers valuing this diversity for wine complexity and vineyard resilience. The informal perpetuation characteristic of heirloom clones creates both opportunities (preservation of valuable adapted material) and challenges (uncertain identity, potential virus accumulation, lack of certified health status).
Heirloom_Clone_Role captures the generational continuity dimension of heritage clones, where value derives from sustained local stewardship rather than formal selection programs or documented historical events. Heirloom material embodies localized knowledge—generations of vignerons selecting propagation material from superior performers, creating adapted populations carrying accumulated selection wisdom. The informal nature of heirloom perpetuation means material typically lacks the documentation, uniformity, and certified status of registered clones producers using heirloom material accept (or embrace) this heterogeneity as intrinsic to traditional viticulture. Contemporary interest in heirloom material reflects reactions against the uniformity of modern clonal viticulture, with some producers deliberately maintaining heirloom selections for distinctiveness, complexity, and connection to viticultural tradition.
Erbstück-Klon
Familienselektion
Gutselektion
Mehrgenerationenklon
estate selection
family selection
generational clone
heirloom clone
McGourty G, Christensen LP (2013) Heritage clone vineyard development, Practical Winery & Vineyard Journal 34(1): 46-52 Pelsy F (2010) Molecular diversity within grapevine varieties, Heredity 104: 331-340 This P et al. (2006) Historical origins of wine grapes, Trends in Genetics 22: 511-519
Heirloom Clone Role
Hen and chicken quality assessment for boron deficiency in Finger Lakes Riesling documented the characteristic cluster appearance and quantified the yield impact of B-limited fruit set. Visual cluster evaluation at véraison identified hen and chicken quality in 38% of clusters, with affected clusters showing the pathognomonic mixed berry population of normal green berries (mean diameter 12 mm) interspersed with small yellow-green shot berries (mean diameter 4-5 mm) that remained hard while normal berries softened at véraison. Berry population analysis quantified hen and chicken quality severity, with affected clusters containing a mean of 55% normal berries by count (representing 85% of cluster weight), 35% shot berries (representing 12% of weight), and 10% completely aborted positions visible as dried remnants on the rachis. The hen and chicken quality impact on cluster weight showed affected clusters averaging 95 g versus 165 g for normal clusters, a 42% reduction attributable primarily to the shot berry population contributing minimal weight. Ripening assessment revealed that hen and chicken quality complicated harvest timing, with normal berries reaching 22°Brix while shot berries in the same cluster measured only 16°Brix, creating a 6° spread that affected must composition when clusters were harvested together. Comparison with zinc-induced millerandage distinguished the hen and chicken quality of boron deficiency by the absence of characteristic little leaf foliar symptoms and by tissue analysis showing adequate Zn (28 ppm) but deficient B (22 ppm), confirming boron rather than zinc as the causal limitation.
Hen and chicken quality in boron deficiency is a boron deficiency symptom quality that inheres in grapevine clusters and represents the characteristic millerandage condition where clusters contain a mixture of normal-sized seeded berries ("hens") and small seedless shot berries ("chickens") resulting from boron-limited pollen tube growth and impaired fertilization, with the uneven berry development creating the distinctive appearance that gives this disorder its common name and causing both yield reduction through reduced berry weight and quality concerns through uneven ripening and cluster appearance. This quality captures the most economically significant boron deficiency symptom in commercial viticulture, recognizing that hen and chicken clusters cause substantial harvest weight losses while also creating problems for table grape marketability and complications for wine grape processing due to differential sugar accumulation between normal and shot berries. The ontological significance of hen and chicken quality lies in its diagnostic value indicating boron limitation affecting reproductive development, its economic impact through yield and quality reduction, and its distinction from zinc-induced millerandage that produces similar symptoms but through different mechanisms and with different associated features. The biological basis of hen and chicken formation encompasses boron's essential role in pollen tube elongation where rapid cell wall synthesis during tube growth creates high boron demand, with boron-deficient pollen producing tubes that grow more slowly, fail to reach the ovule, or burst before completing fertilization, resulting in unfertilized ovaries that either abort entirely or develop parthenocarpically as seedless shot berries. The ratio of normal to shot berries within clusters reflects the degree of boron limitation during the flowering period, with mild deficiency producing predominantly normal berries with scattered shot berries while severe deficiency produces clusters dominated by shot berries with few normal berries. The distribution of hen and chicken severity among clusters may show position effects within the vine canopy reflecting microclimate differences in boron demand or localized boron distribution, and among vines reflecting soil boron availability variation across the vineyard.
Hen_And_Chicken_Quality is_a Boron_Deficiency_Symptom_Quality, representing the millerandage condition with mixed normal berries ("hens") and shot berries ("chickens") from B-limited pollen tube growth and fertilization. This class addresses the most economically important B deficiency symptom. Similar appearance to Zn-induced millerandage but distinguished by foliar symptoms (little leaf with Zn, not with B) and tissue analysis. Shot berries ripen unevenly (lower Brix) complicating harvest. Cluster weight reduction 30-50% typical. Severity reflects B status during bloom period specifically.
B-induced millerandage
Bor-Durchrieseln
Bor-induzierte Millerandage
Hühner-und-Küken bei B-Mangel
boron shot berries
millerandage from B deficiency
uneven berry set from boron
ungleichmäßiger Beerenansatz durch Bor
Hen And Chicken Quality
Herbaceous aroma assessment in Loire Valley Sauvignon Blanc reveals the regional style with moderate green pepper (5.8 on 10-point scale) from 3-isobutyl-2-methoxypyrazine at 22 ng/L, complemented by cut grass (4.5) and fresh herb (5.2), contrasting with Marlborough expression where tropical fruit dominates and herbaceous notes recede (green pepper 3.2, IBMP 8 ng/L), demonstrating how herbaceous intensity creates the stylistic distinction between these regional expressions and enables origin identification.
Herbaceous aroma quality is an aroma character quality that represents olfactory perception evoking plant vegetative tissue-associated scents including green, grassy, leafy, and herbal notes, characterizing the fresh, often green aromatic impression that in moderation contributes varietal typicity and complexity but in excess may indicate underripe fruit or processing faults. Within the BFO framework, this quality addresses a major aroma category distinguished by its derivation from green, photosynthetically active plant tissues and its dual role as both positive varietal marker (herbaceous character valued in Sauvignon Blanc, Cabernet Franc) and potential quality indicator (excessive green character suggesting underripe fruit). The ontological significance lies in herbaceous aroma representing the vegetative rather than fruit dimension of grape-derived character, where the balance between fruity and herbaceous components indicates ripeness level, varietal expression, and winemaking decisions regarding green note retention or elimination. The compositional basis for herbaceous aroma includes C6 aldehydes and alcohols (hexanal, cis-3-hexen-1-ol providing cut grass and green leaf notes), methoxypyrazines (3-isobutyl-2-methoxypyrazine creating green pepper character in Sauvignon family varieties), and various terpenes contributing herbal notes (eucalyptol, rotundone). The ripeness dynamics of herbaceous aroma show declining intensity with grape maturation as C6 compounds decrease and methoxypyrazines degrade through photolytic breakdown, creating the inverse relationship between green character and perceived ripeness that makes herbaceous intensity a harvest timing indicator. The variety-specificity of herbaceous character creates strong expectations, with herbaceous notes valued and expected in Sauvignon Blanc, acceptable in Cabernet Franc, but potentially indicating deficiency in varieties without herbaceous tradition such as Merlot or Chardonnay. Understanding herbaceous aroma quality enables varietal typicity assessment based on expected green character, ripeness evaluation from herbaceous intensity relative to fruit development, and quality judgment incorporating style-appropriate herbaceous expression.
Herbaceous_Aroma_Quality serves as the parent class for herbaceous subcategories (Green_Pepper, Grassy, Herbal), representing an aroma character category with both positive and negative quality associations depending on variety and intensity. This quality is compositionally distinct from fruity aromas, arising from different compound classes (C6 compounds, methoxypyrazines) associated with vegetative tissue. Context-appropriate assessment requires variety-specific expectations for herbaceous expression level.
Kräuteraroma
green aroma
grüner Duft
leafy notes
pflanzlicher Charakter
vegetal character
vegetative Note
vegetative scent
Allen MS et al. (1991) Contribution of methoxypyrazines to Sauvignon blanc wine aroma. American Journal of Enology and Viticulture 42(2): 109-112 Kotseridis Y, Baumes R (2000) Identification of impact odorants in Bordeaux red grape juice, in the commercial yeast used for its fermentation, and in the produced wine. Journal of Agricultural and Food Chemistry 48(2): 400-406
Herbaceous Aroma Quality
An herbaceous green one-year cane on Vitis vinifera cv. Sauvignon Blanc examined in late September, showing bright green color throughout length, soft flexible texture allowing easy bending without structural resistance, cross-sectional examination revealing epidermis as outer protective layer (rather than periderm), broad cortex of thin-walled chlorenchyma cells containing active chloroplasts, discrete vascular bundles in ring arrangement (rather than continuous cambial cylinder), and large central pith, representing normal mid-season developmental status with expectation of progressive lignification initiation over subsequent 6-8 weeks, but raising concerns if similar herbaceous character persists beyond mid-October indicating potential maturation problems.
A green one-year cane exhibiting characteristics typical of herbaceous plant tissues rather than woody perennial structures, characterized by soft flexible stems with minimal structural support capacity, predominance of primary growth tissues including epidermis, cortical parenchyma, and primary vascular bundles, absence of periderm or secondary xylem development, and physiological function oriented toward active growth and photosynthesis rather than storage and structural support. Herbaceous green one-year canes represent developmental status appropriate for spring through mid-summer growing shoots but problematic if persisting into autumn when transition to woody secondary growth should occur, with continued herbaceous character into late season indicating disrupted maturation processes requiring identification and correction of causal factors to restore normal developmental progression.
Herbaceous green one-year canes illustrate the fundamental developmental transition grapevines must complete annually, shifting from herbaceous primary growth (shoot elongation, leaf production) to woody secondary growth (lignification, reserve accumulation, cold hardening). The timing and completeness of this transition determines fruiting wood quality and winter survival. Herbaceous tissues are metabolically active, photosynthetically productive, and responsive to environmental signals during growing season, but become liabilities as winter approaches due to freeze sensitivity, mechanical weakness, and inability to maintain dormancy. The shift from herbaceous to woody involves coordinated changes in gene expression, hormone signaling, carbohydrate metabolism, and cell wall biochemistry, with disruption at any point potentially leaving canes in unsuitable herbaceous state requiring removal during pruning.
krautige grüne Rute
nicht-holzige Rute
non-woody green cane
primary tissue cane
shoot-like green cane
Based on Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 89-112, 245-267 Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press, pp. 125-148.
Herbaceous Green One Year Cane
Herbal aroma profiling in Châteauneuf-du-Pape Grenache identifies the regional garrigue signature with thyme (6.5 on 10-point scale), lavender (5.8), and rosemary (4.2) notes complementing the fruit character, while neighboring Gigondas shows shifted herbal profile with sage (5.5) and wild herb (6.0) dominance, demonstrating how herbal character contributes to appellation differentiation within Southern Rhône and provides the garrigue typicity valued in these Mediterranean expressions.
Herbal aroma quality is an herbaceous aroma quality that represents olfactory perception evoking culinary and aromatic herb-associated scents including basil, thyme, oregano, sage, mint, eucalyptus, and bay leaf notes, characterizing the complex, often savory aromatic impression that contributes to regional typicity and complexity in wines from specific terroirs. Within the BFO framework, this quality specifically addresses the aromatic herb subcategory of herbaceous aroma, distinguished from grassy (cut vegetation) and green pepper (Capsicum) by its association with dried herb, shrubland, and garrigue-like aromatics that evoke Mediterranean landscape and wild herb character. The ontological significance lies in herbal aroma contributing distinctive terroir expression, where wines from regions with surrounding garrigue vegetation (southern Rhône, Provence, Languedoc, Priorat) often show herbal complexity attributed to both environmental volatile transfer and shared monoterpene chemistry between wild herbs and grape berries, creating the sense of place that defines terroir character. The compositional basis for herbal aroma includes various monoterpenes (eucalyptol providing eucalyptus and cooling notes, thymol contributing thyme character, menthol creating mint impression), rotundone (providing black pepper note that crosses into herbal perception), and certain sesquiterpenes associated with sage and bay leaf character. The regional associations of herbal character are strong, with Southern Rhône Grenache-based wines showing characteristic garrigue herbs (thyme, lavender, rosemary), Australian Shiraz displaying eucalyptus from nearby trees' volatile transfer, and Cabernet Franc showing sage and bay leaf in cooler expressions. The aging dynamics of herbal character show some herbal notes increasing with bottle development as terpene glycoside hydrolysis releases bound compounds, while others diminish through volatilization, creating evolving herbal profiles through wine maturation. Understanding herbal aroma quality enables regional typicity assessment based on expected herb character, terroir expression evaluation where herbal complexity indicates site influence, and quality judgment incorporating appropriate herbal contribution to aromatic complexity.
Herbal_Aroma_Quality addresses the culinary and aromatic herb subcategory of herbaceous aroma, encompassing Mediterranean garrigue herbs, mints, and eucalyptus. This quality is strongly associated with regional typicity and terroir expression, particularly in Mediterranean wine regions. The distinction from grassy (fresh-cut vegetation) and green pepper (Capsicum) character reflects different compound origins and perceptual categories within the broader herbaceous aroma family.
Garrigue-Charakter
Kräuterduft
Mediterranean herb notes
aromatic herb scent
aromatischer Kräuterduft
garrigue character
herb aroma
mediterrane Kräuternote
Herve E et al. (2003) Characterization of the odor-active compounds in Shiraz wine using gas chromatography-olfactometry. Journal of Agricultural and Food Chemistry 51(26): 7689-7695 Siebert TE et al. (2008) Development of a quantitative method for the determination of rotundone, the pepper aroma compound, in wine and grapes. Journal of Agricultural and Food Chemistry 56(10): 3745-3748
Herbal Aroma Quality
The "suitcase clones" of California Pinot Noir (selections brought from Burgundy in the pre-regulation era) bear Heritage_Clone_Roles representing mid-20th-century Burgundian genetic material preserved outside France's formal certification system—valued for their connection to historic Burgundy and their documented quality in California conditions. Traditional Sangiovese selections maintained in Chianti Classico family estates since pre-phylloxera replanting bear Heritage_Clone_Roles, perpetuated through sélection massale for generations and considered essential to the traditional character of their wines. The "alte Rebe" (old vine) Riesling selections of the Mosel serve in Heritage_Clone_Roles, material maintained from 19th-century plantings representing pre-phylloxera and early post-phylloxera genetics distinct from modern registered clones.
A clone role that inheres in a grapevine clone by virtue of its historical significance, antiquity, or connection to traditional viticultural heritage, distinguishing it from modern selections developed through systematic scientific programs. Heritage clones represent the accumulated selection of past generations—material perpetuated through centuries of viticulture before formal clonal selection began, carrying genetic diversity, site adaptation, and cultural significance that may be absent from modern uniform selections. Heritage designation may reflect documented historical provenance (material traceable to specific historic vineyards or periods), antiquity (clones of demonstrated age preceding modern selection), survival characteristics (material persisting through phylloxera, wars, or abandonment), or cultural significance (clones associated with traditional wine styles, famous vineyards, or regional identity). Heritage clones often exist as heterogeneous populations (mass selections) rather than uniform genotypes, with their diversity considered both a challenge (variability) and an asset (complexity, resilience). Conservation of heritage clones represents both cultural preservation and genetic resource maintenance, safeguarding diversity that may prove valuable as climate and markets evolve.
Heritage_Clone_Role is subclassed by the specific heritage dimension—Historic (documented provenance), Heirloom (family or estate perpetuation), Pre_Phylloxera (surviving from before the epidemic), and Old_Vine (maintained in aged plantings)—recognizing that heritage significance derives from various sources. Heritage clones occupy complex positions in contemporary viticulture they may lack the documentation, uniformity, and certified health status of registered clones while offering genetic diversity, site adaptation, and cultural continuity unavailable from nursery catalogs. Conservation efforts increasingly recognize heritage clone value, with programs at FPS, ENTAV, and regional institutions collecting and preserving traditional selections alongside formal clone development. The tension between heritage preservation and certification requirements reflects different values—uniformity and traceability versus diversity and tradition—that viticultural communities navigate differently.
Ahnenrebe
Erbklon
ancestral vine material
heritage clone
historic selection
historische Selektion
traditional clone
traditioneller Klon
This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340 McGourty G, Christensen LP (2013) Developing and using a heritage clone vineyard, Practical Winery & Vineyard Journal 34(1): 46-52
Heritage Clone Role
Gouais Blanc bearing a heritage cultivar role in northeastern France is recognized as one of the most historically significant wine grapes despite near-total commercial disappearance, identified through DNA profiling as a parent of at least 80 modern cultivars including Chardonnay (Pinot × Gouais Blanc), Riesling (probable Gouais Blanc parentage), Gamay, Aligoté, and Melon, with heritage conservation plantings maintained at INRAE Domaine de Vassal and at the Conservatoire du Vignoble Charentais preserving approximately 15 clonal accessions of this irreplaceable genetic ancestor.
A traditional cultivar role that inheres in a grapevine cultivar recognized as possessing exceptional cultural, historical, or oenological patrimony value within a region, typically identified through formal heritage designation programs, conservation initiatives, or cultural recognition that elevates the cultivar's status beyond its current commercial significance.
Heritage_Cultivar_Role captures the cultural patrimony dimension of cultivar significance that is not reducible to commercial viability, agronomic performance, or geographic origin. Many heritage cultivars are rare, endangered, or commercially marginal yet possess outsized genetic, historical, or cultural importance: examples include Kadarka in Hungary (cultural icon of historical Egri Bikavér despite commercial decline), Mammolo in Tuscany (traditional Chianti component largely displaced by Sangiovese monoculture), and Mission/País/Listán Prieto in the Americas (the founding cultivar of New World viticulture). This role often intersects with Conservation_Role subclasses.
Kulturerbe-Sorten-Rolle
Patrimonialsorten-Rolle
heritage variety role
historische Erbe-Sorten-Rolle
legacy cultivar role
patrimony cultivar role
Bowers JE et al. (1999) Historical genetics: the parentage of Chardonnay, Gamay, and other wine grapes of northeastern France, Science 285: 1562-1565 Robinson J et al. (2012) Wine Grapes, Allen Lane pp: 397-399 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157
Heritage Cultivar Role
Heritage status for Roter Veltliner in Austria documents a cultivar of recognized cultural and genetic significance — once widely planted in Niederösterreich (approximately 700 ha in 1970), declining to approximately 180 ha by 2023, formally recognized as a heritage variety by the Verein zur Erhaltung und Förderung alter österreichischer Rebsorten (Association for the Preservation of Traditional Austrian Grape Varieties), maintained in the Klosterneuburg genebank collection (12 accessions), and the subject of targeted conservation through the Wagram "Roter Veltliner Pur" initiative promoting premium single-cultivar bottlings to sustain commercial viability as a conservation mechanism. Heritage status for Gouais Blanc in France documents a cultivar of extraordinary genetic significance — once one of the most widely planted varieties in medieval France ("le raisin des paysans"), progressively eliminated through quality-oriented replanting from the 15th century onward, reduced to scattered remnant vines by the 20th century, but revealed by molecular analysis (Bowers et al. 1999) to be a parent of at least 75 modern cultivars including Chardonnay, Gamay, Riesling, Aligoté, and Auxerrois, now conserved in the INRA Domaine de Vassal collection and small heritage plantings — a cultivar whose heritage status rests on genetic legacy rather than contemporary wine quality.
Heritage status is a classification status entity that documents the formal recognition of grape cultivars as viticultural heritage resources meriting conservation-oriented protection, encompassing national and regional heritage variety designations (cultivars recognized as culturally significant elements of viticultural patrimony), endangered variety registrations (cultivars at risk of extinction due to declining acreage, loss of propagation material, or abandonment of traditional cultivation), genebank conservation status (the inclusion of cultivar accessions in national and international germplasm repositories), and the institutional programs supporting heritage variety preservation (conservation vineyards, heritage variety promotion initiatives, research funding for orphan cultivar characterization), providing the conservation-heritage dimension of classification that complements the commercial-quality orientation of recommended and authorized status by recognizing cultivars whose value lies in genetic diversity preservation, cultural heritage, and potential future utility rather than in current commercial viability.
Heritage status documentation addresses the conservation dimension of cultivar classification — the recognition that the viticultural cultivar portfolio represents a cultural and genetic heritage resource whose value extends beyond current commercial utility to encompass genetic diversity for future adaptation, cultural identity preservation, and the maintenance of living agricultural traditions. The global trend toward cultivar concentration — the progressive dominance of a small number of internationally marketable varieties (the top 10 cultivars by global acreage now account for approximately 35% of the world's vineyard area) at the expense of minor indigenous cultivars — has created an urgent conservation challenge analogous to biodiversity loss in other agricultural sectors. The conservation mechanisms available for heritage cultivars span a continuum from ex situ preservation (genebank collections maintaining living plants and cryopreserved material under controlled conditions — the INRA Vassal-Montpellier collection with 2,700+ accessions and the Klosterneuburg collection with 3,500+ accessions being the largest European repositories) through in situ conservation (maintaining heritage cultivars in commercial or semi-commercial vineyard cultivation, the preferred approach for preserving the cultivar's adaptation to its traditional environment) to market-based conservation (creating premium market positioning for heritage variety wines that provides the economic incentive for continued cultivation). The Slow Food Foundation's Ark of Taste and Presidia programs have extended the heritage food concept to wine grape cultivars, recognizing several dozen endangered varieties worldwide and supporting producer networks dedicated to their commercial revival. The genetic value dimension of heritage status has been dramatically reinforced by molecular parentage studies revealing that many apparently insignificant or abandoned cultivars are genetic founders of major modern varieties — Gouais Blanc's role as a parent of dozens of prominent cultivars, Traminer's parentage of Grüner Veltliner, and Savagnin's genetic network across Central European cultivars all demonstrate that heritage cultivars may harbor genetic diversity of immense but previously unrecognized value for future breeding programs and climate adaptation.
Denkmalschutzstatus
Erbsorte-Status
Erhaltungsstatus
Kulturerbe-Klassifizierung
bedrohter Sortenstatus
conservation status
cultural heritage classification
endangered variety status
heritage variety designation
patrimony status
Lacombe T et al. (2011) Large-scale parentage analysis in an extended set of grapevine cultivars. Theoretical and Applied Genetics 122(2): 401-414 Maul E et al. (2012) Vitis International Variety Catalogue and the European Vitis Database. In Proceedings of the 10th International Conference on Grapevine Breeding and Genetics, Acta Horticulturae 1082: 53-58 Slow Food Foundation (2023) Ark of Taste — Grape Varieties HBLA Klosterneuburg (2023) Erhaltung alter Rebsorten — Sortensammlung und Genbank
Heritage Status
Cabernet Sauvignon in Napa Valley at BBCH 65 exhibits hermaphrodite flowers with erect stamens extending 0.8-1.2 mm beyond pistil length, producing 18,000-24,000 pollen grains per flower with 70-80% viability, achieving 52-58% fruit set under favorable bloom conditions (June temperatures 23-28°C, morning humidity 55-70% declining to 35-45% by afternoon), with self-pollination occurring efficiently through gravity and air movement without requiring insect vectors or cross-pollination. Chardonnay in Burgundy shows hermaphrodite flowers with balanced stamen-pistil geometry (stamens extending 0.4-0.8 mm beyond stigma), 15,000-20,000 pollen grains per flower, 65-75% pollen viability, and 48-55% fruit set under typical Burgundian bloom conditions. Syrah in Northern Rhône exhibits particularly vigorous hermaphrodite flowers with stamens extending 1-1.5 mm beyond pistil, 22,000-28,000 pollen grains per flower, 75-82% viability, and 54-62% fruit set in warm, dry bloom periods.
A hermaphrodite flower is a perfect flower in which both male organs (stamens) and female organs (pistil) are fully functional, with erect stamens producing abundant viable pollen and a receptive pistil capable of successful fertilization and fruit set, representing the optimal reproductive condition that enables reliable self-pollination, consistent fruit set, and stable yield without requiring cross-pollination from other plants. Hermaphrodite flowers exhibit the complete and functional bisexual condition, characterized by five erect stamens with filaments extending equal to or beyond the pistil length, anthers positioned at or above stigma level facilitating efficient pollen transfer to the stigmatic surface during anthesis, production of abundant viable pollen (typically 12,000-30,000 pollen grains per flower with 60-85% viability), and a fully developed pistil with receptive wet stigma, functional style supporting pollen tube growth, and viable ovules capable of fertilization. The critical diagnostic feature of hermaphrodite flowers is erect stamen position with stamens equaling or exceeding pistil length, contrasting sharply with functionally female flowers where stamens are strongly reflex (curved downward below pistil) or functionally male flowers where pistil is abnormally abbreviated. Hermaphrodite flower morphology at anthesis typically shows stamens extending 0.3-1.5 mm beyond the stigma, creating spatial proximity that facilitates autonomous self-pollination as shed pollen falls or is blown onto the stigmatic surface within the same flower or adjacent flowers on the same inflorescence. Pollen production in hermaphrodite flowers is copious, with anthers dehiscing through longitudinal slits 1-2 hours after calyptra shedding, releasing clouds of yellow pollen that remain viable for 24-72 hours depending on environmental conditions (optimal viability at 60-80% RH, declining rapidly above 90% RH or below 40% RH). Stigma receptivity in hermaphrodite flowers persists for 3-6 days post-anthesis under favorable conditions, providing an extended window for successful pollination. The combination of abundant pollen production, efficient pollen placement through erect stamen geometry, extended stigma receptivity, and functional fertilization machinery results in hermaphrodite flowers achieving fruit set percentages typically ranging from 40-60% under optimal bloom conditions (warm, dry weather with temperatures 20-30°C and humidity 50-70%), significantly exceeding the 20-35% fruit set typical of functionally female flowers requiring cross-pollination. Hermaphrodite flowers characterize the vast majority of commercially important Vitis vinifera wine cultivars (Cabernet Sauvignon, Merlot, Chardonnay, Pinot Noir, Syrah, Riesling, Sauvignon Blanc, etc.) and many table grape cultivars, representing the agronomically preferred sex type due to reliable self-fertility eliminating pollinizer requirements and enabling consistent yields across diverse environmental conditions. The hermaphrodite condition shows high but not absolute stability, with some cultivars exhibiting phenotypic plasticity in stamen position and pollen viability in response to bloom-period environmental conditions, particularly temperature stress during flower development occasionally inducing partial stamen reflexing or reduced pollen viability that temporarily shifts hermaphrodite flowers toward functionally female expression.
Hermaphrodite_Flower represents the fully functional bisexual condition and must be distinguished from Functionally_Female_Flower (pistil functional but stamens reflex or producing non-viable pollen) and Functionally_Male_Flower (stamens functional but pistil non-receptive). The term 'hermaphrodite' specifically denotes functional bisexuality rather than mere morphological presence of both organs (which is denoted by 'perfect'). Hermaphrodite flowers represent the agronomically optimal sex type for consistent fruit production, as they eliminate dependency on cross-pollination, enable successful production as isolated vines or monovarietal plantings, and achieve higher and more stable fruit set across variable environmental conditions. The evolutionary origin of hermaphrodite flowers in cultivated Vitis vinifera represents a key domestication event, arising through mutation from dioecious wild ancestors (V. vinifera ssp. sylvestris) and subsequently selected for reliable fruit production. Modern breeding programs overwhelmingly target hermaphrodite flower expression as a prerequisite trait for new cultivar releases. Sub-classification into Fully_Functional and Balanced types reflects subtle gradations in hermaphrodite functionality.
Zwitterblüte
bisexual functional flower
hermaphroditische Blüte
self-fertile flower
vollständig zwittrige Blüte
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 99-106 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 94-101 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Battilana, J. et al. (2013). The genetic basis of the conversion of hermaphrodite flowers to male flowers in grapevine. Nature Communications 4=2615 Lebon, G. et al. (2008). Sugars and flowering in the grapevine. Journal of Experimental Botany 59=2565-2578.
Hermaphrodite Flower
All major wine grape cultivars including Cabernet Sauvignon, Chardonnay, Riesling, and Tempranillo possess hermaphrodite flower disposition with upright stamens (5 per flower) surrounding a central pistil. Pollen viability in hermaphrodite cultivars ranges from 70-95%, with Pinot Noir averaging 85-90% and Gewürztraminer showing lower viability at 70-80%. Self-pollination accounts for 30-70% of fertilization events in hermaphrodite cultivars depending on vineyard configuration, with the remainder from cross-pollination by adjacent vines.
A flower type disposition that is the inherited capacity of a Vitis vinifera plant to develop perfect (bisexual) flowers containing both functional stamens producing viable pollen and a functional pistil with receptive stigma, enabling self-pollination and cross-pollination within a single flower.
Hermaphrodite flower disposition is the predominant condition in cultivated V. vinifera, arising from domestication selection for reliable fruit set without requiring male pollinizer vines. The trait is controlled by the Sex locus on chromosome 2, with the hermaphrodite allele (H) dominant to both male (M) and female (F) alleles. Functional hermaphroditism requires coordinated development of both organ types and synchronous maturation.
Hermaphroditismus
Zwitterblütigkeit
bisexual flower disposition
complete flower capacity
perfect flower trait
zwittrige Blütenanlage
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 98-105 Oberle GD (1938) A genetic study of variations in floral morphology and function in cultivated forms of Vitis. New York Agricultural Experiment Station Technical Bulletin 250:1-63
Hermaphrodite Flower Disposition
In a Mosel Riesling vineyard, hexose transport initiation was characterized in mid-September Year N+1 (BBCH 83) when VvHT1 expression reached 62-fold elevation, VvHT2 showed 45-fold induction, glucose uptake into isolated mesocarp cells was 22 μmol g⁻¹ h⁻¹ (from 4.5 μmol pre-veraison), vacuolar hexose concentration rose from 180 mM to 520 mM over 6 days, berry Brix increased from 9.2° to 13.8°, and ABA levels were 1050 ng/g under cool German conditions (daily high 19-22°C, night 10-13°C, extended ripening period).
Hexose_Transport_Initiation represents the beginning of enhanced membrane transport enabling glucose and fructose uptake from apoplastic space into mesocarp cells and subsequent accumulation in vacuoles, establishing the critical cellular import machinery that converts apoplastically-released hexoses into stored sugars creating berry sweetness. This initiation occurs during early veraison (BBCH 83), typically 0-2 days following phloem unloading activation, and is characterized by hexose transporter gene expression increasing 25-70 fold (VvHT1 40-80 fold, VvHT2 30-60 fold, VvHT3/5/6 20-50 fold) producing membrane proteins that catalyze facilitated diffusion of hexoses across plasma membrane and tonoplast, hexose uptake rates into isolated mesocarp cells rising from pre-veraison 2-6 μmol glucose g⁻¹ h⁻¹ to initiation phase 12-28 μmol g⁻¹ h⁻¹ representing 4-8 fold enhancement, and beginning cytoplasm-to-vacuole transport achieving rates 15-35 μmol hexose g⁻¹ h⁻¹ enabling vacuolar accumulation to 300-600 mM during initial transport phase. Molecular basis of transport initiation involves: VvHT1 gene (encoding high-affinity glucose transporter with Km 1.0-2.5 mM) shows rapid induction reaching 50-70% of maximum expression within 2-4 days enabling efficient glucose uptake even at moderate apoplastic concentrations, VvHT2 (facilitating both glucose and fructose with Km 4-10 mM) achieves 40-65% maximum expression providing capacity for both hexoses, additional transporters (VvHT3, VvHT5, VvHT6) contribute with combined 30-50% maximum expression creating diverse transporter portfolio, and regulatory control involves sugar-responsive transcription factors (VvMYB, VvbZIP families) binding to promoter elements activated by hexokinase-mediated sugar sensing creating positive feedback loops. Subcellular localization of transporters includes: plasma membrane VvHT1/2/3 enabling hexose uptake from apoplast (estimated 60-75% of transporter protein), tonoplast-localized forms of VvHT proteins facilitating cytoplasm-to-vacuole transport (20-35% of protein), and potential endo-membrane (ER, Golgi) localization during synthesis and trafficking representing 5-10% of total at steady state, with immunolocalization and GFP fusion studies demonstrating predominant plasma membrane and tonoplast distribution. Transport kinetics during initiation show: glucose transport Vmax increasing from 8-15 μmol g⁻¹ h⁻¹ pre-veraison to 25-50 μmol g⁻¹ h⁻¹ during initiation through increased transporter density, fructose transport showing similar enhancement with Vmax 20-45 μmol g⁻¹ h⁻¹, Km values remaining relatively constant (glucose 1.5-3.0 mM, fructose 5-12 mM) indicating increased capacity rather than affinity changes, and competition kinetics demonstrating glucose uptake slightly favored over fructose (relative rate 1.1-1.4:1.0) though final vacuolar ratios approach equimolar through differential metabolism. Energetics of hexose transport involve: facilitated diffusion down concentration gradients (apoplast 100-200 mM total hexoses → cytoplasm 60-120 mM → vacuole accumulating to 400-800 mM during initiation) providing primary driving force without direct ATP coupling, potential role for proton-coupled transport at tonoplast where vacuolar H⁺-ATPase creating pH gradient (cytoplasm pH 7.2-7.5, vacuole pH 3.0-3.8) could drive secondary active transport though evidence suggests primarily facilitated diffusion, and metabolic maintenance of cytoplasmic hexose below apoplastic concentrations through glycolysis and other pathways sustaining import gradient. Cellular consequences of transport initiation inclu vacuolar hexose accumulation from 100-250 mM to 400-800 mM within 5-8 days creating osmotic potential shift from -0.6 to -1.2 MPa to -1.2 to -2.0 MPa driving water import and berry expansion, cytoplasmic hexose maintained at 60-120 mM through balanced import, metabolism, and tonoplast transport preventing excessive accumulation that could trigger feedback inhibition, and hexose:sucrose ratio in whole berry shifting from 3.0 reflecting dominant hexose import and accumulation versus residual sucrose. Hormonal regulation of transport initiation demonstrates: ABA levels 700-1400 ng/g during initiation activate ABA-responsive elements in hexose transporter gene promoters, sugar itself acting through hexokinase sensor creating glucose-6-phosphate signals that upregulate transporter genes establishing positive feedback amplification, declining auxin and gibberellin removing growth-related suppression allowing transporter gene expression, and potential ethylene involvement in some cultivars with slight elevation correlating with enhanced transport though mechanism unclear. Environmental modulation of transport initiation includes: temperature affecting transporter protein synthesis and membrane fluidity with optimal 24-28°C supporting maximum induction rates while heat >32°C can impair through protein misfolding or membrane disruption, osmotic stress from water deficit potentially affecting membrane transport capacity though moderate deficit appears compatible with normal initiation, and nutritional status particularly nitrogen and potassium affecting membrane synthesis and transport protein production with adequate nutrition essential for full transport capacity establishment. Cultivar variation in transport initiation characteristics shows: rapid-initiating varieties (Chardonnay, Gewürztraminer) achieve 60-75% maximum transport capacity within 3-5 days post-activation, moderate cultivars (Merlot, Cabernet Sauvignon) reach 50-65% maximum in 5-7 days, while slower types (Nebbiolo, Mourvèdre) require 7-10 days to achieve 45-60% maximum reflecting genetic differences in transporter gene regulation and protein synthesis rates. Coordination with sugar accumulation shows: transport initiation slightly precedes (0-2 days) measurable Brix increase establishing capacity before major flux, peak transport capacity achievement (80-100% maximum) coincides with rapid sugar accumulation phase (BBCH 84-85) enabling maximum import rates 2.5-4.5% Brix per week, and transport capacity maintenance through ripening (70-90% maximum even at late stages) sustains continued accumulation though rate declines as vacuolar concentration approaches equilibrium with phloem supply. Membrane dynamics during transport initiation involve: plasma membrane area potentially increasing through vesicle fusion or membrane expansion though data limited, tonoplast membrane surface area substantially increased (estimated 40-80%) through vacuole enlargement and membrane proliferation creating enhanced transport capacity, and lipid composition changes with increased unsaturation potentially enhancing membrane fluidity and transport protein function though specific modifications during initiation poorly characterized. Quality implications inclu transport capacity established during initiation strongly correlates with final sugar levels achieved (r² = 0.72-0.86) with inadequate initiation limiting ultimate Brix, uniformity of transport initiation across berries and clusters affecting ripening synchrony and harvest maturity variation, and timing relative to other ripening processes determining balance between sugar and flavor/acid/phenolic development important for wine style. Viticultural factors affecting transport initiation inclu crop load with excessive yields potentially impairing initiation through limited photosynthate availability for transporter synthesis, canopy management ensuring adequate nitrogen and resources for protein synthesis supporting transporter production, water management with severe deficit potentially limiting membrane synthesis while moderate deficit compatible with normal initiation, and disease or environmental stress during initiation period potentially causing long-term impairment of sugar accumulation capacity even if conditions later improve.
Hexose_Transport_Initiation is distinguished from Phloem_Unloading_Activation by representing membrane transport rather than phloem pathway change, from Initial_Sugar_Increase by focusing on transport mechanism rather than net accumulation, and from Sugar_Import_Enhancement by showing initiation rather than progressive amplification. Transporter induction: VvHT1 40-80 fold, VvHT2 30-60 fold. Uptake rates increase 4-8 fold. Timing: 0-2 days after unloading activation. Temperature optimal 24-28°C. Hexose_Transport_Initiation establishes cellular import machinery critical for sugar accumulation during grape ripening.
Glukose-Fruktose-Transport-Onset
Membran-Hexose-Aufnahme-Aktivierung
Zucker-Transporter-Induktion
glucose-fructose transport onset
membrane hexose uptake activation
sugar transporter induction
vacuolar hexose loading initiation
vakuoläre Hexose-Beladungs-Initiierung
Hayes MA, Davies C, Dry IB (2007) Isolation, functional characterization, and expression analysis of grapevine hexose transporters J Exp Bot 58:1985-1997 hexose transporter characterization Afoufa-Bastien D, Medici A, Jeauffre J, Coutos-Thévenot P, Lemoine R, Atanassova R, Laloi M (2010) The Vitis vinifera sugar transporter gene family Plant Mol Biol 72:533-546 sugar transporter family Lecourieux F, Kappel C, Lecourieux D, Serrano A, Torres E, Arce-Johnson P, Delrot S (2014) An update on sugar transport and signaling in grapevine J Exp Bot 65:821-832 transport regulation.
Hexose Transport Initiation
Hexose transporter activity in grape berries increases 5-15 fold between veraison and maturity, with VvHT1 (high-affinity) and VvHT6 (low-affinity) being the predominant isoforms in ripening fruit. High-sugar cultivars such as Grenache show 2-3 fold higher VvHT transcript abundance than moderate-sugar cultivars such as Riesling at equivalent developmental stages. VvHT1 exhibits Km values of 0.1-0.3 mM for glucose, enabling efficient uptake at low apoplastic concentrations early in ripening, while VvHT6 (Km 2-5 mM) becomes increasingly important as sugar concentrations rise. Transporter activity is localized primarily in berry flesh cells, with highest expression in the peripheral mesocarp zone where most sugar storage occurs. Water deficit increases VvHT expression by 30-60%, contributing to sugar concentration under drought stress.
A sugar accumulation disposition that is the inherited capacity of a Vitis vinifera plant to regulate hexose sugar import into ripening berries through the expression and activity of plasma membrane hexose transporter proteins (VvHT family), determining the rate and final extent of sugar accumulation by controlling the entry of glucose and fructose from phloem-unloaded apoplast into berry flesh parenchyma cells.
Hexose transporter activity represents a key regulatory point for sugar accumulation, controlling the final step of sugar import into storage cells after phloem unloading. The trait shows genetic variation among cultivars and responds dynamically to developmental and environmental signals. Understanding transporter regulation has implications for both breeding (selection for modified accumulation) and management (stress effects on sugar loading). The shift from symplastic to apoplastic phloem unloading at veraison makes transporter activity essential for post-veraison sugar accumulation.
Hexose-Transporter-Aktivität
VvHT activity trait
VvHT-Aktivität
Zuckertransporter-Expression
hexose import capacity
sugar transporter expression
Afoufa-Bastien D et al. (2010) The Vitis vinifera sugar transporter gene family: phylogenetic overview and macroarray expression profiling. BMC Plant Biology 10:245 Hayes MA et al. (2007) Involvement of abscisic acid in the coordinated regulation of a stress-inducible hexose transporter (VvHT5) and a cell wall invertase in grapevine in response to biotrophic fungal infection. Plant Physiology 145:537-546
Hexose Transporter Activity Disposition
In a Sonoma Chardonnay vineyard, hexose transporter upregulation was characterized by qPCR over 9 days in early August Year N+1 (BBCH 83) when VvHT1 expression increased from 1.2-fold to 95-fold above green berry reference, VvHT2 rose from 0.8-fold to 72-fold, berry Brix increased from 8.6° to 14.8° (2.9% per week), glucose uptake into mesocarp cells rose from 5.2 to 52 μmol g⁻¹ h⁻¹, ABA levels 1250 ng/g, under warm conditions (daily high 27-30°C supporting rapid induction).
Hexose_Transporter_Upregulation represents the dramatic increase in expression of genes encoding membrane proteins that facilitate glucose and fructose uptake creating the molecular foundation for enhanced hexose import capacity that enables berry sugar accumulation during ripening. This upregulation occurs during veraison onset through mid-veraison (BBCH 83-84), typically 1-6 days following initial developmental signals, and is characterized by VvHT gene family members showing coordinated induction: VvHT1 (high-affinity glucose transporter) increasing 40-150 fold above green berry baseline, VvHT2 (facilitating glucose and fructose) rising 30-100 fold, VvHT3 showing 25-80 fold elevation, VvHT5 and VvHT6 displaying 20-70 fold induction, with collective VvHT expression representing 300-500 fold increase when summed across all family members creating massive enhancement in transporter gene transcription. Molecular mechanisms of upregulation inclu transcription factor binding to transporter gene promoters with sugar-responsive transcription factors (VvMYB, VvbZIP families) recognizing SURE elements (Sugar Responsive Elements, consensus TGGACGG) in VvHT promoters activated by hexokinase-mediated sugar sensing, ABA-responsive transcription factors (VvABF, VvAREB) binding ABRE elements (ABA Responsive Elements, PyACGTGGC motifs) in VvHT promoters responding to veraison ABA spike 50-200 → 700-1600 ng/g, chromatin modifications with histone acetylation increasing accessibility of VvHT genes enabling transcription factor access, and potential DNA methylation changes reducing silencing allowing developmental expression. Transcriptional dynamics during upregulation show: VvHT1 transcript levels rising from 0.5-2.0 copies per cell (green berry) to 40-120 copies per cell (mid-veraison) over 4-8 days representing exponential accumulation phase, VvHT2 similarly increasing from 0.3-1.5 to 25-90 copies per cell, transcript half-life 4-8 hours requiring continuous transcription to maintain high steady-state levels, and mRNA processing including splicing and polyadenylation efficiently producing mature transcripts for translation. Regulatory control involves: positive feedback loops where initial hexose uptake creates glucose-6-phosphate signals that further enhance VvHT expression amplifying accumulation, cross-talk between sugar and ABA signaling pathways with both converging on VvHT promoters creating synergistic upregulation, developmental programming with veraison-specific transcription factors coordinately activating multiple ripening genes including VvHTs, and potential negative feedback at very high expression levels (>100-150 fold) preventing excessive transporter synthesis once capacity saturates. Protein synthesis from upregulated transcripts shows: translation initiation within 10-30 minutes of transcript appearance producing VvHT proteins, protein synthesis rates 25-60 ng transporter protein per g berry fresh weight per hour during peak upregulation, targeting signals directing nascent proteins to ER for membrane insertion and processing, and glycosylation in ER/Golgi potentially affecting transporter stability or function though modifications incompletely characterized for grape VvHTs. Transporter protein properties inclu molecular mass 50-55 kDa for most VvHT proteins consistent with 12-transmembrane domain structure typical of sugar transporters, predicted topology with cytoplasmic N and C termini and 12 membrane-spanning α-helices creating central pore for hexose passage, substrate specificity with VvHT1 preferring glucose (Km 1-2.5 mM) and VvHT2 transporting both glucose and fructose (Km 4-10 mM for each), and functional characterization in heterologous systems (yeast, Xenopus oocytes) confirming facilitated diffusion mechanism without energy coupling. Subcellular localization of upregulated transporters includes: plasma membrane dominance with 65-80% of VvHT protein at cell surface enabling hexose uptake from apoplast, tonoplast localization 15-30% facilitating cytoplasm-to-vacuole hexose transport for storage, and potential endo-membrane (ER, Golgi) distribution 5-10% representing trafficking intermediates en route to final destinations. Tissue-specific patterns of upregulation show: mesocarp cells showing strongest VvHT upregulation corresponding to sugar storage function, phloem-adjacent cells potentially showing earlier or more intense upregulation facilitating uptake from unloading zones, and vascular tissue itself showing minimal VvHT expression consistent with role in transport rather than accumulation. Temporal progression of upregulation demonstrates: early phase (days 0-3) showing 5-25 fold induction establishing initial capacity, middle phase (days 3-7) achieving 30-80 fold elevation reaching 60-75% of maximum, late phase (days 7-12) attaining 80-120 fold approaching saturating levels, and maintenance phase (beyond day 12) sustaining 80-110 fold throughout ripening with balance between synthesis and degradation maintaining steady-state. Environmental modulation of upregulation includes: temperature affecting transcription and translation rates with optimal 24-28°C supporting rapid upregulation (peak achieved 5-7 days) while cooler conditions 18-24°C extend upregulation period 8-12 days through slower kinetics, heat stress >31°C potentially impairing upregulation through reduced gene expression or protein misfolding, light having indirect effects through photosynthesis providing substrate that triggers sugar signaling amplifying upregulation, and water status with adequate hydration supporting normal upregulation while severe deficit may reduce through general transcriptional suppression. Nutritional influences inclu nitrogen availability important for protein synthesis with N-deficient vines potentially showing 25-40% reduced transporter protein levels despite normal transcript induction indicating translational limitation, sulfur required for methionine and cysteine in transporter proteins with deficiency potentially limiting synthesis, and general nutrient status affecting cellular capacity for protein production with severely nutrient-limited vines showing impaired upregulation. Hormonal fine-tuning of upregulation involves: ABA concentration directly correlating with VvHT expression levels (r² = 0.72-0.86) across developmental stages and treatments, exogenous ABA application to pre-veraison berries inducing premature VvHT upregulation demonstrating hormonal control, and ABA biosynthesis inhibitors preventing normal upregulation confirming requirement for endogenous ABA accumulation. Cultivar genetic variation in upregulation characteristics shows: rapid-upregulating varieties achieving maximum fold-induction 100-150 fold within 5-7 days, moderate cultivars reaching 70-110 fold in 7-10 days, and slower types requiring 10-14 days for 60-90 fold reflecting genetic differences in regulatory element sensitivity, transcription factor levels, or chromatin accessibility. Functional consequences of upregulation inclu hexose uptake capacity increasing proportionally with transporter levels enabling berry sugar accumulation rates 3.0-4.8% Brix per week during peak upregulation compared to 120 fold) suggesting capacity limits related to membrane area or metabolic coupling rather than transporter abundance per se. Research applications inclu VvHT expression analysis providing molecular marker for veraison onset and ripening progression, promoter studies revealing regulatory mechanisms controlling developmental expression, and transgenic or CRISPR approaches potentially enabling manipulation of sugar accumulation through VvHT expression modulation though not yet commercially deployed. Quality implications inclu adequate upregulation essential for achieving target sugar levels with inadequate induction limiting final Brix regardless of source photosynthesis or phloem delivery, timing of upregulation relative to other ripening processes affecting compositional balance, and uniformity of upregulation across berries and clusters determining ripening synchrony important for harvest timing and wine consistency.
Hexose_Transporter_Upregulation is distinguished from Sugar_Transport_Activation by representing gene expression changes rather than overall system activation, from Hexose_Transport_Initiation by focusing on transcriptional induction rather than initial function, and from maximum expression by showing progressive increase (40-150 fold induction) rather than sustained peak. Duration 4-10 days BBCH 83-84. ABA-dependent regulation critical. Temperature optimal 24-28°C. Hexose_Transporter_Upregulation represents critical molecular transition enabling cellular hexose import capacity.
Hexose-Träger-Hochregulation
Membran-Transporter-Aktivierung
VvHT expression increase
VvHT-Expressions-Zunahme
Zucker-Transporter-Gen-Induktion
hexose carrier upregulation
membrane transporter activation
sugar transporter gene induction
Hayes MA, Davies C, Dry IB (2007) Isolation, functional characterization, and expression analysis of grapevine hexose transporters J Exp Bot 58:1985-1997 VvHT upregulation Vignault C, Vachaud M, Cakir B, Glissant D, Dédaldéchamp F, Büttner M, Atanassova R, Fleurat-Lessard P, Lemoine R, Delrot S (2005) VvHT1 encodes a monosaccharide transporter expressed in the conducting complex of the grape berry phloem J Exp Bot 56:1409-1418 VvHT1 characterization Afoufa-Bastien D, Medici A, Jeauffre J, Coutos-Thévenot P, Lemoine R, Atanassova R, Laloi M (2010) The Vitis vinifera sugar transporter gene family Plant Mol Biol 72:533-546 VvHT family regulation.
Hexose Transporter Upregulation
Riesling exhibits high acidity retention disposition, maintaining 7-10 g/L TA and pH 3.0-3.2 at flavor maturity in German wine regions, with the combination of high tartrate (7-9 g/L) and slow malate degradation (k = 0.02-0.03/day) preserving acidity. Barbera retains 7-9 g/L TA even in warm Piedmont climates through exceptionally slow malate degradation despite moderate tartrate levels, enabling production of high-acid red wines. Chenin Blanc in the Loire Valley maintains 8-11 g/L TA with pH 3.0-3.3, supporting both dry and sweet wine styles requiring prominent acidity. High-retention cultivars typically show 20-40% higher residual malate at equivalent thermal time from veraison compared to low-retention types.
An acidity retention disposition that is the inherited capacity of a Vitis vinifera plant to maintain elevated titratable acidity (typically greater than 7 g/L) and low pH (below 3.3) at harvest maturity, through combinations of high tartrate concentration, slow malic acid degradation, moderate potassium accumulation, and/or genetic buffering against temperature-induced acid loss.
High acidity retention disposition is essential for cool climate viticulture where crisp acidity defines wine style, and increasingly valuable for warm climate adaptation where maintaining freshness is challenging. The trait enables extended hang time for flavor development without excessive acid loss, expanding the harvest window. High-retention cultivars provide base material for sparkling wine production requiring TA above 8-9 g/L. Selection for high retention is complicated by the multiple contributing mechanisms and genotype × environment interaction.
acid-preserving trait
high acid retention
hohe Azidität-Bewahrung
hohe Säureretention
low pH disposition
Duchêne E et al. (2014) Genetic variability of descriptors for grapevine berry acidity in Riesling, Gewürztraminer and their progeny. Australian Journal of Grape and Wine Research 20:91-99 Martinez de Toda F, Balda P (2015) Quantifying the effect of temperature on decoupling anthocyanins and sugars of the grape (Vitis vinifera L. cv. Maturana Tinta de Navarrete). Vitis 54:117-120
High Acidity Retention Disposition
Syrah showing high activity absorptive feeder roots in May, sequential soil sampling revealing youngest white feeder roots 1-4 weeks post-maturation showing highest uptake, isotope pulse-chase demonstrating high activity roots providing 64% of nitrogen uptake despite comprising 32% of feeder root length, respiration measurements showing oxygen consumption 3.8-fold higher in high activity versus older roots, high activity absorptive feeder roots demonstrating peak performance providing disproportionate resource acquisition
High activity absorptive feeder root is absorptive feeder root showing maximum absorption intensity and metabolic activity, characterized by highest uptake rates per unit surface area or biomass among feeder root population, elevated respiration providing ATP for active transport, abundant mitochondria supporting energy metabolism, maximum transporter expression and activity, and demonstrated superior performance through comparative uptake measurements, with high activity roots typically representing youngest most vigorous portion of absorptive feeder root population 1-8 weeks post-maturation showing white coloration, dense cytoplasm, abundant root hairs, developing mycorrhizae, and optimal cellular organization, and serving disproportionate role where high activity roots despite comprising perhaps 20-40% of absorptive root population potentially provide 50-70% of total uptake creating targeting opportunity where practices enhancing high activity root abundance or prolonging high activity period could substantially improve vine resource acquisition efficiency. High activity absorptive feeder root metabolic intensity includes= showing elevated respiration (oxygen consumption rates 2-5 fold higher than average absorptive roots reflecting intense metabolism), exhibiting high ATP production (abundant mitochondria generating energy supporting active transport and biosynthesis), demonstrating active biosynthesis (high rates of protein, membrane, and wall synthesis supporting growth and function), revealing heat production (metabolic intensity potentially measurable as heat generation), and displaying resource demand (high activity roots requiring substantial photosynthate allocation supporting intense metabolism). High activity absorptive feeder root transport capacity includes= showing maximum transporter abundance (nutrient and water transporters expressed at highest levels in high activity roots), exhibiting optimal membrane function (membranes in peak condition maintaining maximum selective permeability and transport capacity), demonstrating maximum H+-ATPase activity (proton pumps operating at highest rates creating strongest electrochemical gradients), revealing transporter saturation (uptake approaching maximum velocity indicating transporters operating near capacity), and displaying superior performance (uptake per unit surface area 2-10 fold higher than older absorptive roots). High activity absorptive feeder root cellular characteristics includes= showing dense cytoplasm (cytoplasm occupying large proportion of cell volume reflecting active metabolic state), exhibiting abundant organelles (high density of mitochondria, ER, Golgi, ribosomes supporting intense activity), demonstrating large nuclei (prominent nuclei indicating active gene expression and cellular regulation), revealing thin walls (primary cell walls thin and unlignified facilitating rapid growth and transport), and displaying optimal turgor (cells fully turgid reflecting adequate osmotic function and water status). High activity absorptive feeder root root hair characteristics includes= showing maximum density (high activity roots producing root hairs at highest densities potentially 300-600+ per centimeter), exhibiting active growth (many root hairs still elongating adding new absorption surface), demonstrating optimal function (root hairs in peak condition with intact membranes and active transport), revealing extensive coverage (root hair zone extending several centimeters creating large effective absorption surface), and displaying critical contribution (root hairs on high activity roots providing disproportionate uptake). High activity absorptive feeder root mycorrhizal development includes= showing active colonization (arbuscular mycorrhizal fungi actively colonizing high activity roots forming new arbuscules), exhibiting functional arbuscules (developing arbuscules creating active nutrient exchange interfaces), demonstrating external hyphal growth (mycorrhizal hyphae extending from high activity roots exploring surrounding soil), revealing symbiosis establishment (mycorrhizal association developing toward mature functional state), and displaying enhanced uptake (mycorrhizae beginning to enhance high activity root absorption particularly for phosphorus). High activity absorptive feeder root uptake performance includes= providing disproportionate contribution (high activity roots potentially providing 50-70% of total uptake despite comprising 20-40% of absorptive population), exhibiting superior efficiency (uptake per unit biomass or construction cost highest among root classes), demonstrating responsive capacity (high activity roots most responsive to changing soil conditions or plant demands), revealing competitive advantage (high activity roots most effective at capturing resources in competitive situations), and displaying critical importance (high activity root abundance strongly affecting vine resource status and performance). High activity absorptive feeder root duration and transition includes= showing limited duration (high activity period typically lasting 2-8 weeks before roots transition to lower activity mature state), exhibiting gradual decline (transition from high to moderate activity occurring progressively not abruptly), demonstrating age correlation (high activity typically characterizing youngest viable absorptive roots), revealing environmental effects (favorable conditions potentially extending high activity period while stress shortens), and displaying inevitable aging (all high activity roots eventually aging to lower activity states). High activity absorptive feeder root management implications includes= requiring production support (maximizing high activity root abundance requiring practices promoting continuous new root production), showing timing importance (practices timed to enhance high activity root production during critical periods like spring growth), exhibiting irrigation benefits (adequate moisture maintaining high activity root function and extending active period), demonstrating fertilization response (nutrient availability potentially enhancing high activity root performance and duration), and revealing opportunity (understanding and targeting high activity roots offering potential for improving resource use efficiency).
High_Activity_Absorptive_Feeder_Root emphasizes maximum absorption intensity and superior performance and is distinguished from moderate activity roots by focus on peak uptake rates. Critical for understanding metabolic intensity, maximum transport, cellular optimization, disproportionate contribution, limited duration, and the highest-activity roots providing majority of uptake despite minority of population representing key target for management.
Hochaktive absorptive Feinwurzel
Intensiv-absorbierende Wurzel
Intensively absorbing root
Maximal-aufnehmende Wurzel
Maximum-uptake root
Peak-activity root
High Activity Absorptive Feeder Root
Grenache exhibits exceptional high basal fruitfulness disposition with 1.8-2.4 clusters per bud at nodes 1-3, enabling traditional gobelet training with 2-bud spurs. Tempranillo maintains 1.6-2.0 clusters at basal positions, well-suited for bilateral cordon systems. In contrast, Sultana shows poor basal fruitfulness (0.4-0.8 clusters at nodes 1-3), requiring cane pruning to access fruitful distal buds. High basal fruitfulness cultivars tolerate severe hedging and mechanical pruning without significant yield reduction, with spur-pruned Grenache producing 8-12 t/ha compared to 5-7 t/ha for spur-pruned Sultana.
A bud fruitfulness disposition that is the inherited capacity of a Vitis vinifera plant to develop high numbers of inflorescence primordia in buds at basal node positions (nodes 1-4 from the cane base), enabling productive short spur pruning and cordon-based training systems.
High basal fruitfulness is a key selection criterion for cultivars destined for mechanized viticulture and warm climate production where spur pruning simplifies management. The trait shows moderate heritability (h² = 0.45-0.60) and can be enhanced through canopy management promoting light exposure to basal buds during primordium initiation. Rootstock influence on basal fruitfulness is generally limited but may manifest through effects on shoot vigor and bud microclimate.
Zapfenfruchtbarkeit
basal bud fertility
basale Fruchtbarkeit
proximal node fertility
spur fruitfulness
stammnahe Knospenfertilität
May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press, Adelaide pp.: 45-68 Clingeleffer PR (1984) Production and growth of minimal pruned Sultana vines. Vitis 23:42-54
High Basal Fruitfulness Disposition
A high bush head arm on Vitis vinifera cv. Tempranillo in modernized traditional Rioja vineyard, emerging from head at 65cm height (elevated from traditional 35-45cm), extending 42cm creating bush approximately 90cm total height, improved ergonomic access enabling harvest workers to pick from standing position rather than bending or kneeling, reduced disease pressure from elevated positioning improving air circulation and eliminating soil splash, representing contemporary adaptation of traditional head training maintaining bush architecture and avoiding trellis installation while improving operational efficiency and disease management, transition reflecting evolution in Rioja toward moderate modernization of traditional practices balancing heritage preservation with economic and practical necessities.
A bush head arm configured in elevated architecture with head positioned at increased height (typically 50-80cm) and arms creating taller bush structure, characterized by improved ergonomic access for manual operations, reduced soil contact and splash contamination risks, enhanced air circulation through elevated canopy position, and potential adaptation toward more contemporary management while maintaining head-trained character. High bush head arms represent moderate-height head training, providing compromise between traditional low bush configurations and modern elevated cordon systems, improving worker ergonomics and disease management compared to low bush while maintaining self-supporting head-trained architecture avoiding trellis requirements, suitable for transitional situations adapting traditional systems toward contemporary practices or situations where moderate height provides operational benefits while preserving head-trained identity.
High bush head arms represent evolutionary adaptation of traditional head training, responding to contemporary needs (worker ergonomics, disease management, operational efficiency) while maintaining head-trained architecture and avoiding conversion to cordon systems. The elevated positioning (50-80cm heads versus traditional 30-45cm) provides significant practical benefits including standing-position harvest access (reducing worker fatigue and health issues), improved disease management (better air circulation, no soil splash), easier mechanical cultivation (equipment clearance beneath canopy), and potential mechanization compatibility (some elevated bush vines accessible to adapted harvest equipment). However, elevated bush training requires stronger trunk support (taller unsupported structure), increases wind exposure (elevated canopy catches more wind), and may reduce traditional character perceptions (market positioning as traditional wines). The trend in traditional regions increasingly favors moderate elevation of historical low bush systems as pragmatic evolution preserving head-trained heritage while addressing operational realities.
elevated bush arm
erhöhte Kopfarm
hohe Busch-Arm
raised head arm
tall bush configuration
High Bush Head Arm
Riesling exhibits high chilling requirement disposition with optimal dormancy release requiring 900-1200 chilling hours, contributing to its successful cultivation in German, Austrian, and Alsatian climates. Cabernet Sauvignon requires 800-1000 chilling hours and shows incomplete budbreak (60-75%) when grown in subtropical regions without dormancy-breaking treatment. Pinot Noir populations from Burgundy demonstrate high chilling requirements (850-1100 hours) that present challenges in warm winter areas of Australia and California. High-chill cultivars moved to mild winter climates typically require hydrogen cyanamide application (1-2%) to achieve commercially acceptable budbreak uniformity.
A chilling requirement disposition that is the inherited capacity of a Vitis vinifera plant to require substantial cold accumulation (typically greater than 800 chilling hours or 60 chill portions) during dormancy for complete dormancy release, adapting the cultivar to continental or cool temperate climates with reliable winter cold.
High chilling requirement evolved as an adaptation to continental climates, preventing premature budbreak during transient winter warm spells followed by damaging cold. The trait provides a safety mechanism against spring frost damage by delaying budbreak until winter is reliably concluded. High-chill cultivars planted in insufficient chilling zones show poor canopy development, reduced fruitfulness, and accelerated vine decline over multiple seasons. Climate change projections indicate many currently adequate chilling regions will become marginal for high-chill cultivars by 2050.
cold-requiring cultivar
continental dormancy type
high chill trait
hohe Chilling-Anforderung
hoher Kältebedarf
kontinentaler Dormanz-Typ
Moncur MW et al. (1989) Base temperatures for budbreak and leaf appearance of grapevines. American Journal of Enology and Viticulture 40:21-26 Webb LB et al. (2007) Modelled impact of future climate change on the phenology of winegrapes in Australia. Australian Journal of Grape and Wine Research 13:165-175
High Chilling Requirement Disposition
A Concord vine trained to Geneva Double Curtain (GDC) in New York's Finger Lakes bears a High_Cordon_Role at 180 cm height, with two parallel cordons and pendant shoot growth creating the divided canopy suited to this vigorous juice grape variety—the high cordon managing vigor while enabling mechanical harvest. A Trebbiano vine in central Italian tendone (overhead) training serves in a High_Cordon_Role, with cordons at 200 cm supporting a pergola-like canopy that shades fruit from intense summer sun while enabling traditional inter-row cultivation beneath the vines. A Cabernet Franc vine on high cordon in a humid Virginia vineyard bears a High_Cordon_Role positioning fruit at 160 cm to escape the ground-level humidity that promotes disease in low-trained systems.
A cordon-trained vine role that inheres in a grapevine with cordons positioned at elevated height (typically 150-200 cm above ground), creating a high fruiting zone with shoots growing downward or outward from the cordon rather than upward as in low cordon systems. High cordon training (also termed high wire, GDC-related systems, or overhead configurations) positions fruit away from ground-level humidity and frost while enabling agricultural operations beneath the vine canopy, though it may increase fruit exposure to sun and wind stress. High cordon systems are particularly suited to vigorous varieties and fertile sites where elevated fruit positioning and pendant shoot growth help manage excess vigor through natural apical dominance disruption. Harvest from high cordons may be facilitated by mechanical over-row harvesters or complicated by the elevated fruit position requiring reaching or elevated platforms for hand operations. High cordon systems include Geneva Double Curtain (GDC, two parallel high cordons), Scott Henry (high and low cordons with shoots trained in opposite directions), and various overhead/tendone configurations, each adapting the high cordon concept to specific production objectives.
High_Cordon_Role captures the category of elevated fruiting zone systems that fundamentally alter canopy microclimate compared to low cordon approaches. High cordons trade the labor efficiency of low positioning for vigor management benefits and ground-level clearance, with the optimal choice depending on variety vigor, site fertility, climate humidity, and production system requirements. The pendant (downward-growing) shoot orientation typical of high cordon systems disrupts normal grapevine apical dominance, potentially reducing shoot vigor while improving light distribution in divided canopy configurations. High cordon systems are particularly associated with juice grape production (Concord, Niagara in eastern North America) and vigorous sites where canopy management challenges favor elevated, divided architectures. Climate change adaptation may shift the high versus low cordon calculus, with elevated fruit potentially avoiding increased ground-level heat while benefiting from greater air movement.
Hocherziehung
Hochkordon
Hochkultur
elevated cordon
erhöhter Kordon
high cordon
high wire
high-trained vine
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Shaulis N et al. (1966) Response of Concord grapes to light exposure and Geneva Double Curtain training, Proceedings of the American Society for Horticultural Science 89: 268-280 Reynolds AG, Vanden Heuvel JE (2009) Training systems, American Journal of Enology and Viticulture 60: 251-268
High Cordon Role
A Pinot Noir vine at 10,000 vines/hectare (1.0 m × 1.0 m) in Chambolle-Musigny bears a High_Density_Planted_Role, the traditional Burgundian spacing limiting vine vigor on potentially fertile Côte d'Or soils while forcing root competition that proponents believe enhances terroir expression. A Chardonnay vine at 8,000 vines/hectare in premium Carneros planting serves in a High_Density_Planted_Role, the density chosen to emulate Burgundian models for premium sparkling wine base production. A Riesling vine at 7,500 vines/hectare in steep Mosel terracing bears a High_Density_Planted_Role adapted to narrow terrace widths where tractor access is impossible and intensive hand labor is the only option.
A vine spacing role that inheres in a grapevine planted at high density (typically 6,000-12,000+ vines per hectare), with close spacing both within rows (0.8-1.2 m) and between rows (0.9-1.5 m) creating intense root zone competition and limiting individual vine vigor through resource partitioning. High-density planting represents the traditional European model, developed in Burgundy, Champagne, and other classic regions before mechanization, where the philosophy holds that vine competition forces deep root development, limits excessive vigor, and produces concentrated fruit expressing site character. The close spacing requires either narrow tractors (specially designed "enjambeurs" straddling single rows) or hand labor for most operations, with the infrastructure and labor costs offset by perceived quality benefits and historical capital investment in existing plantings. High-density vineyards typically feature less vigorous training systems (single guyot, short bilateral cordon) suited to the limited per-vine resources, with lower per-vine yields but potentially higher per-hectare production due to increased vine numbers. Contemporary adoption of high density outside traditional regions reflects quality-focused production philosophy, particularly for varieties (Pinot Noir, Chardonnay) associated with high-density Burgundian origins.
High_Density_Planted_Role captures the intensive planting model associated with European quality traditions and increasingly adopted by premium New World producers. The quality arguments for high density—root competition forcing deep exploration, vigor limitation improving fruit concentration, more vines sampling more terroir variation—remain debated, with research showing variable results depending on site conditions and management. High-density economics are challenging: increased vine costs (6,000+ vines versus 1,500), specialized equipment requirements, higher labor per hectare, and longer establishment periods offset the quality premiums that high-density fruit may command. The trend toward "Burgundian" density in New World Pinot Noir and Chardonnay plantings reflects both quality philosophy and marketing positioning, with density itself becoming a quality signal regardless of its actual impact in specific sites. Practical limits on high density reflect variety vigor and site fertility excessively close spacing on vigorous sites creates canopy management nightmares that may reduce rather than enhance quality.
European density
Hochdichte-Pflanzung
close spacing
enge Pflanzung
europäische Dichte
high-density planting
intensive Bepflanzung
intensive planting
Archer E, Strauss HC (1985) Effect of plant density on root distribution, South African Journal of Enology and Viticulture 6: 25-30 Reynolds AG et al. (1994) Vine spacing effects, American Journal of Enology and Viticulture 45: 444-459 Gladstones J (1992) Viticulture and Environment, Winetitles Van Leeuwen C et al. (2019) The concept of terroir, OENO One 53: 101-114
High Density Planted Role
140 Ruggeri (V. berlandieri × V. rupestris) demonstrates high drought tolerance supporting Catarratto production on the arid hillsides of western Sicily receiving only 300-350 mm annual rainfall, with root systems penetrating 5+ m into fractured limestone and maintaining yields of 4-6 t/ha without irrigation. 110 Richter provides high drought tolerance for Garnacha in the non-irrigated vineyards of Priorat, Catalonia, where summer drought extends from June through September and annual precipitation averages 400-500 mm with high interannual variability. 1103 Paulsen supports dry-farmed Primitivo at 3-5 t/ha on the calcareous soils of Gioia del Colle, Puglia, where summer temperatures exceed 40°C and drought stress is compounded by high evaporative demand.
A drought-tolerant rootstock role that inheres in a grafted rootstock by virtue of its capacity to maintain scion productivity under severe and prolonged water deficit conditions where moderate-tolerance rootstocks would fail. High drought-tolerance rootstocks sustain economically viable yields (typically 60-80% of irrigated potential) under rainfall regimes below 400 mm annually with summer drought periods exceeding 3-4 months, soil water potentials declining below -1.5 MPa in the upper rooting zone, and atmospheric vapor pressure deficits exceeding 3-4 kPa during summer afternoons. This exceptional drought adaptation is achieved through very deep root systems (4-6 m penetration on permeable substrates), high root-to-shoot biomass ratios (0.4-0.6 versus 0.2-0.3 for drought-sensitive genotypes), xylem vessels with small diameter and high cavitation resistance (P₅₀ values of -1.8 to -2.5 MPa), efficient hydraulic redistribution from deep moist soil layers, and superior osmotic adjustment capacity maintaining cell turgor to water potentials below -2.0 MPa.
High_Drought_Tolerance_Rootstock_Role captures the upper end of the drought adaptation spectrum, representing rootstocks capable of sustaining commercial viticulture under conditions approaching the aridity limits for Vitis cultivation. All rootstocks in this category derive from V. berlandieri × V. rupestris crosses, reflecting the exceptional drought adaptation of these parental species. High drought tolerance often correlates with lime tolerance (also a V. berlandieri trait), as both adaptations are advantageous in the calcareous, semi-arid Mediterranean environments where these rootstocks predominate. Vigor is typically moderate to high, as extensive root systems support substantial canopy development when water becomes available.
Unterlage für extreme Trockenstandorte
extreme drought adapted rootstock
high drought tolerance rootstock
highly drought tolerant rootstock
hoch trockenheitstolerante Unterlage
sehr trockenheitsresistente Unterlage
severe drought tolerant rootstock
Carbonneau A (1985) The early selection of grapevine rootstocks for resistance to drought conditions, American Journal of Enology and Viticulture 36: 195-198 Pou A et al. (2012) Adjustments of water use efficiency by stomatal regulation during drought and recovery in the drought-adapted Vitis hybrid Richter-110, Physiologia Plantarum 134: 313-323 Tomás M et al. (2014) Variability of water use efficiency in grapevines, Environmental and Experimental Botany 103: 148-157
High Drought Tolerance Rootstock Role
Chasselas exhibits high fruit set disposition with 45-55% conversion rates even under suboptimal flowering conditions, contributing to its reputation for consistent yields. Pinot Gris achieves 40-50% fruit set in cool climate regions where other cultivars struggle. Table grape cultivars selected for large clusters such as Italia and Red Globe show fruit set of 50-65% through breeding selection. High fruit set cultivars typically possess high pollen viability (>85%), rapid pollen tube growth, and strong sink strength in developing ovaries.
A fruit set disposition that is the inherited capacity of a Vitis vinifera plant to consistently achieve above-average conversion of flowers to berries (typically greater than 40%), resulting from efficient pollination, fertilization, and early berry retention under a range of environmental conditions.
High fruit set disposition may lead to excessively compact clusters in some cultivars, increasing susceptibility to Botrytis and other bunch rots. Management practices including early leaf removal, shoot positioning, and gibberellin application can moderate excessive set. The trait shows moderate heritability (h² = 0.40-0.55) and is partially independent of total flower number per cluster.
ausgezeichnete Befruchtung
excellent fruit set capacity
high berry set
hoher Beerenansatz
hoher Fruchtansatz
superior set trait
Dry PR et al. (2010) Bunch architecture and susceptibility of berries to Botrytis cinerea in Vitis vinifera L. Australian Journal of Grape and Wine Research 16:257-264 Dunn GM, Martin SR (2007) A functional association in Vitis vinifera L. cv. Cabernet Sauvignon between the extent of primary branching and the number of flowers formed per inflorescence. Australian Journal of Grape and Wine Research 13:95-100
High Fruit Set Disposition
A high horizontal cordon on Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, positioned at 105cm height along heavy-gauge trellis wire, extending 92cm from trunk, bearing 15 two-bud spurs, shoots trained vertically into 150cm of trellis space above cordon, clusters hanging in zone from 95-125cm height easily accessible to mechanical harvester head operating beneath cordon with adequate clearance (typically requiring 60-80cm ground-to-cordon clearance), representing modern high-cordon architecture optimized for mechanical harvest efficiency, worker ergonomics (all hand operations performed from comfortable standing position without bending), and disease management (elevated canopy position improving air circulation and reducing soil splash contamination).
A horizontal cordon positioned at elevated height above ground level, typically 90-130cm, characterized by substantial clearance from soil surface facilitating mechanical operations and ergonomic manual work, spur positioning at comfortable standing height for pruning and management reducing worker fatigue, suitability for mechanized harvest systems requiring equipment clearance beneath cordons, and canopy development in elevated zone potentially improving air circulation and reducing disease pressure compared to low-cordon configurations. High horizontal cordons represent modern mechanization-compatible architecture increasingly common in commercial viticulture, with elevated positioning enabling efficient over-row and under-row mechanical operations, improved ergonomics for hand operations performed from standing position, enhanced air movement through elevated canopy reducing disease pressure, and in hot climates potential benefits from removing fruit zone from extreme ground-level heat while maintaining adequate sun exposure.
High horizontal cordons represent the trend in modern commercial viticulture, particularly in regions and operations emphasizing mechanization, labor efficiency, and disease management. The elevated positioning (90-130cm typical, occasionally higher) provides numerous benefits but also some tradeoffs including increased trellis and stake costs (taller posts, more substantial wire support), potentially greater wind exposure (elevated canopies catch more wind potentially increasing shoot breakage and fruit damage), and in some climates excessive fruit exposure if canopy management creates gaps (requiring leaf retention or bunch shading strategies). The optimal cordon height depends on multiple factors including harvest method (mechanical harvest typically requiring 95-110cm minimum), worker ergonomics preferences (100-110cm comfortable for most operations), equipment specifications (tractor and implement clearance requirements), and variety characteristics (vigorous varieties benefiting from height for canopy accommodation, weak varieties potentially struggling with extensive vertical development).
erhöhte Kordon
high-positioned cordon
hohe horizontale Kordon
mechanization-height cordon
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Intrieri et al. (2011) "Grapevine mechanization= Constraints and perspectives" Acta Horticulturae 978=471-478 Morris (2007) "Development and commercialization of a complete vineyard mechanization system" HortTechnology 17=411-420.
High Horizontal Cordon
A high light-adapted mature leaf at node 5 on a south-facing exterior shoot of Vitis vinifera cv. Grenache in southern Rhône Valley, France, positioned at canopy top receiving approximately 95% of full sunlight (daily PPFD averaging 1650 μmol photons m⁻² s⁻¹ with midday peaks exceeding 2100 μmol photons m⁻² s⁻¹), measuring 9.8 cm length by 9.2 cm width representing 48% size reduction compared to interior leaves, very thick lamina of 365 μm with four distinct palisade cell layers totaling 145 μm, chlorophyll content of 615 μg cm⁻² concentrated in dense palisade tissue, very thick cuticle of 7.2 μm with substantial wax deposits creating slightly glaucous appearance, dense trichome coverage on abaxial surface creating conspicuous white woolly appearance, very dark green adaxial surface with bronze tinge especially near veins from anthocyanin accumulation, demonstrating light-saturated photosynthesis rate of 16.8 μmol CO₂ m⁻² s⁻¹ at 2000 μmol photons m⁻² s⁻¹, maximum PSII efficiency maintained at 0.81 despite high light stress indicating effective photoprotection, measured at BBCH stage 85 (softening of berries) in mid-August during heat wave with maximum air temperatures 38-40°C, representing maximum high light adaptation enabling sustained function under extreme conditions.
A sun-exposed mature leaf exhibiting pronounced structural and functional adaptations to very high light environments receiving 70-100% of full sunlight in Vitis vinifera, characterized by maximum expression of sun leaf syndrome including very small lamina size often 40-50% reduced compared to shade leaves reflecting resource allocation toward tissue density over surface area expansion, very thick lamina frequently 320-380 μm with three or occasionally four palisade cell layers maximizing photosynthetic capacity per unit area, extremely high chlorophyll concentrations per unit area of 550-650 μg cm⁻² supporting high photosynthetic enzyme densities, very thick cuticle often 6-8 μm providing maximum UV protection and water retention, elevated stomatal density of 200-240 stomata mm⁻² with small individual stomata optimizing conductance-to-water-loss ratio, substantial trichome coverage creating conspicuously gray-white abaxial surface appearance, very dark green to bronze-green adaxial coloration from high chlorophyll and anthocyanin concentrations, maximum photoprotective carotenoid content including zeaxanthin accumulation, and physiological characteristics including highest light-saturated photosynthesis rates of 14-18 μmol CO₂ m⁻² s⁻¹, very high light compensation and saturation points, efficient xanthophyll cycle operation dissipating excess light energy, and robust antioxidant systems preventing photoinhibition despite exposure to potentially damaging high irradiance often exceeding 2000 μmol photons m⁻² s⁻¹ at midday in midsummer.
High light-adapted mature leaves represent the extreme sun end of the sun-shade continuum, distinguished from moderate sun leaves by very pronounced structural adaptations and maximum physiological acclimation to high irradiance. These leaves develop primarily in exposed upper and exterior canopy positions particularly on south and west-facing canopy surfaces in Northern Hemisphere or north and west-facing in Southern Hemisphere. Development under high light during expansion is essential for anatomical adaptation, as shade-developed leaves cannot develop multiple palisade layers or high cell density through plastic responses after maturation. High light-adapted leaves show superior photosynthetic performance under high irradiance but may show reduced efficiency under lower light compared to shade leaves, with high light compensation points (typically 50-80 μmol photons m⁻² s⁻¹) reflecting elevated respiratory costs. These leaves face greatest risk of photoinhibition, heat stress, and water stress damage, requiring robust protection mechanisms to maintain function through summer conditions.
extrem-sonnen Blatt
extreme sun leaf
fully acclimated sun leaf
hochlicht-adaptiertes reifes Blatt
maximal lichtangepasstes Weinblatt
maximum light-adapted leaf
Based on Boardman (1977) "Comparative photosynthesis of sun and shade plants" Annual Review of Plant Physiology 28=355-377 Schultz (1996) "Leaf absorptance of visible radiation in Vitis vinifera L.= estimates of age and shade effects with a simple field method" Scientia Horticulturae 66=93-102 Flexas et al. (1999) "Steady-state and maximum rates of photosynthesis in grapevine leaves in response to increased temperature" Functional Plant Biology 26=631-640 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 215-248.
High Light Adapted Mature Leaf
Fercal (BC1 of 31R × Berlandieri Colombard 1B) demonstrates the highest commercial lime tolerance (up to 60% active lime, ICP >80), supporting Ugni Blanc for Cognac production on the Campanian chalks with 40-50% active limestone where all other rootstocks fail. 41B Millardet et de Grasset tolerates 40% active lime, enabling Pinot Noir cultivation on the pure chalk slopes of the Côte des Blancs in Champagne (active lime 30-40%) with chlorophyll content maintained above 400 µg·g⁻¹ fresh weight. 333 EM (V. berlandieri × V. vinifera Cabernet Sauvignon) tolerates 35-40% active lime, used historically in the Charentes and on highly calcareous terraces of the southern Rhône.
A lime-tolerant rootstock role that inheres in a grafted rootstock by virtue of its capacity to maintain scion productivity on highly calcareous soils with active limestone content exceeding 25-30%, where moderate-tolerance rootstocks would exhibit chlorosis and yield reduction. High lime-tolerance rootstocks sustain iron nutrition at active CaCO₃ concentrations of 30-60%, soil pH values of 8.0-8.5, and rhizosphere bicarbonate concentrations that would severely inhibit iron uptake in less tolerant genotypes. This exceptional tolerance is achieved through highly efficient Fe³⁺-chelate reductase systems maintaining reduction rates above 2-3 µmol Fe²⁺·g⁻¹ root·h⁻¹ even under bicarbonate stress, strong proton extrusion capacity (H⁺-ATPase activity), organic acid exudation (citrate, malate) enhancing iron solubility, and superior iron use efficiency allowing normal chlorophyll synthesis at lower leaf iron concentrations. High lime-tolerance rootstocks invariably carry substantial V. berlandieri or V. vinifera genetic background.
High_Lime_Tolerance_Rootstock_Role includes rootstocks tolerating above 25-30% active lime, with Fercal representing the extreme of commercial lime tolerance. The trade-off for exceptional lime tolerance is typically reduced phylloxera resistance, as the most lime-tolerant rootstocks (Fercal, 41B, 333EM) carry V. vinifera genetic background. Site-specific rootstock selection must balance lime tolerance against phylloxera pressure, with 41B and Fercal restricted to low-phylloxera regions (Champagne, Cognac, Burgundy limestone slopes). Climate change may increase lime chlorosis incidence through soil moisture interactions, potentially expanding demand for high-tolerance rootstocks.
Unterlage für stark kalkhaltige Böden
high lime tolerance rootstock
high limestone tolerance rootstock
highly calcareous soil tolerant rootstock
hoch kalktolerante Unterlage
sehr kalkverträgliche Unterlage
very lime tolerant rootstock
Bavaresco L et al. (2003) Grapevine iron nutrition update, Journal International des Sciences de la Vigne et du Vin 37: 89-100 Pouget R (1987) Utilité de la connaissance du pourcentage de calcaire actif dans la terre fine et le sol total pour le diagnostic de nutrition en fer de la vigne, Progrès Agricole et Viticole 104: 91-95 Bert PF et al. (2013) Genetic basis of chloride exclusion in grapevine rootstocks, BMC Plant Biology 13: 179
High Lime Tolerance Rootstock Role
Comprehensive assessment of High_pH_Adaptation_Disposition at the University of Udine evaluated rootstock performance at pH levels from 5.5 to 8.5 (0.5 unit increments) with iron supplied as Fe-EDDHA (stable across pH range) to separate pH effects from iron availability effects. Measurements included root and shoot growth, root morphology (length, branching, root hair development), nutrient uptake efficiency (N, P, K, Mn, Zn, Cu), and cytoplasmic pH maintenance (using pH-sensitive fluorescent probes). High_pH_Adaptation_Disposition showed genotypic variation partially independent of iron acquisition efficiency. 'Fercal' demonstrated strong High_pH_Adaptation_Disposition with shoot growth at pH 8.5 reaching 92% of optimal pH (5.5-6.5) performance when Fe was non-limiting, normal root branching and hair development, and maintenance of root cortical cytoplasmic pH at 7.1-7.3 across external pH range (effective homeostasis). Manganese and zinc uptake efficiency decreased only 20-25% at pH 8.5. '41B' showed similarly strong adaptation (88% growth retention, stable cytoplasmic pH). 'SO4' displayed moderate High_pH_Adaptation_Disposition (72% growth at pH 8.5), with reduced root hair development and 40% decline in Mn uptake at high pH indicating some pH sensitivity beyond Fe effects. '3309C' exhibited poor adaptation with growth declining to 55% at pH 8.5 even with adequate Fe, impaired root hair formation, cytoplasmic pH stress (rising to 7.6, indicating homeostasis failure), and 55% reduction in Mn uptake. These findings demonstrate that High_pH_Adaptation_Disposition contributes to calcareous soil performance beyond iron acquisition capacity.
A High_pH_Adaptation_Disposition is a Lime_Tolerance_Disposition that inheres in a Vitis plant and is realized through physiological mechanisms that enable normal metabolic function and nutrient acquisition across roots exposed to consistently elevated soil pH (7.5-8.5) typical of calcareous environments, beyond the specific challenges of iron acquisition. This disposition represents broad adaptation to alkaline conditions, addressing the multiple nutritional and physiological challenges of high-pH soils that extend beyond iron availability. High soil pH affects availability of multiple nutrients (reducing availability of Fe, Mn, Zn, Cu, B potentially increasing Mo to toxic levels), alters root membrane function and transport processes, affects rhizosphere microbial communities, and can directly impact root cell metabolism through effects on cytoplasmic pH regulation. While iron deficiency is typically the most limiting factor on calcareous soils, adaptation to other high-pH effects contributes to overall lime tolerance. High_pH_Adaptation_Disposition encompasses maintenance of cytoplasmic pH homeostasis despite alkaline external environment, adaptation of nutrient transporters to function at high apoplastic pH, adjustment of organic acid metabolism, and tolerance of altered micronutrient availability profiles. Genotypes with strong High_pH_Adaptation_Disposition show normal growth and function at pH 8.0-8.5 when iron is supplied in available form, while sensitive genotypes may show reduced performance even with adequate iron, indicating pH effects beyond Fe acquisition.
High_pH_Adaptation_Disposition is_a Lime_Tolerance_Disposition. High pH affects multiple aspects of plant function beyond iron availability including other micronutrients, membrane transport, and cellular pH homeostasis. Assessing pH adaptation requires supplying iron in pH-stable form to separate effects. Cytoplasmic pH homeostasis (maintaining pH 7.0-7.5 despite external alkalinity) is a key component. Mn, Zn, Cu availability all decline at high pH, contributing to growth limitation. High pH adaptation is partially independent of iron acquisition efficiency, representing an additional tolerance component.
Alkaliboden-Adaptation
Hoch-pH-Anpassungs-Disposition
Kalkumgebungs-Anpassung
alkaline soil adaptation
calcareous environment adaptation
high pH tolerance
Mengel K (1994) Iron availability in plant tissues—iron chlorosis on calcareous soils, Plant and Soil 165:275-283 Marschner H (1995) Mineral Nutrition of Higher Plants, 2nd Edition, Academic Press, London Tang C et al. (2009) Mechanisms of plant adaptation to alkaline soil, Australian Journal of Experimental Agriculture 49:866-874
High pH Adaptation Disposition
140 Ruggeri (V. berlandieri × V. rupestris) demonstrates high K uptake, elevating Catarratto must pH to 3.7-3.9 versus 3.3-3.5 on SO4 in Sicilian trials, with petiole K at veraison exceeding 5% dry weight. 1103 Paulsen similarly delivers high K to Negroamaro in Puglia, with berry K concentrations 20-40% higher than on 110 Richter, contributing to the low-acid profile typical of southern Italian wines. Ramsey (V. champinii) shows high K uptake supporting Sultana (Thompson Seedless) table grape production where elevated fruit K promotes berry size and firmness, with petiole K commonly exceeding 4% at veraison.
A nutrient acquisition rootstock role that inheres in a grafted rootstock by virtue of its capacity for elevated potassium (K⁺) uptake from soil and translocation to the scion, resulting in high leaf petiole K concentrations and enhanced K accumulation in berries and must. High potassium uptake rootstocks typically deliver petiole K concentrations exceeding 2.5-3.0% dry weight at bloom and 4-6% at veraison, compared to 1.5-2.0% and 2-3% respectively for low-K rootstocks under equivalent soil conditions. This enhanced K acquisition reflects high-affinity K⁺ transporter (HAK/KUP family) expression and activity in root cells, efficient K⁺ loading into xylem, and extensive root exploration of soil K reserves. High K uptake is advantageous on K-deficient soils but can be problematic on K-adequate sites where excessive berry K elevates must pH (reducing total acidity by 0.5-1.5 g/L tartaric equivalents), decreases color stability in red wines through anthocyanin-tannin precipitation, and promotes microbial instability through elevated wine pH.
High_Potassium_Uptake_Rootstock_Role is associated primarily with V. berlandieri × V. rupestris (140Ru, 1103P, 99R) and V. champinii (Ramsey, Dog Ridge) parentage. The elevated K delivery can be partially managed through potassium fertilization reduction, crop load adjustment, and delayed harvest allowing berry K dilution, but rootstock replacement remains the only definitive solution where high-K rootstocks cause persistent wine quality issues. Climate change-induced advanced maturity and increased drought stress may exacerbate rootstock K effects through reduced berry water content concentrating K in must.
K-accumulating rootstock
K-starke Unterlage
Unterlage mit hoher Kaliumaufnahme
high K rootstock
high potassium uptake rootstock
kaliumakkumulierende Unterlage
potassium accumulating rootstock
Mpelasoka BS et al. (2003) A review of potassium nutrition in grapevines with special emphasis on berry accumulation, Australian Journal of Grape and Wine Research 9: 154-168 Kodur S (2011) Effects of rootstocks on grapevine growth, yield and quality of wine, Journal of Wine Research 22: 155-168 Walker RR, Blackmore DH (2012) Potassium concentration and pH of grape juice from Chardonnay and Shiraz as affected by rootstock, Australian Journal of Grape and Wine Research 18: 187-193
High Potassium Uptake Rootstock Role
Ramsey (Salt Creek, V. champinii) is the benchmark high salt-tolerance rootstock, supporting commercial table grape production (Thompson Seedless, Crimson Seedless) at yields of 20-30 t/ha in the Coachella Valley, California, at soil ECe of 4-6 dS/m and irrigation ECw of 1.5-2.5 dS/m where sensitive rootstocks like Schwarzmann and Riparia Gloire fail within 3-5 years. Dog Ridge (V. champinii) demonstrates high salt tolerance in Israeli Negev viticulture, maintaining Sugraone table grapes at ECe exceeding 5 dS/m with brackish irrigation (ECw 3.0 dS/m). 1103 Paulsen shows high salt tolerance in the saline-irrigated vineyards of Mendoza, Argentina, maintaining Malbec productivity at ECe 3-4 dS/m where SO4 shows 50% yield reduction.
A salt-tolerant rootstock role that inheres in a grafted rootstock by virtue of its capacity to maintain scion productivity under high salinity conditions where moderate-tolerance rootstocks would suffer substantial yield loss or vine decline. High salt-tolerance rootstocks sustain economically viable production at soil electrical conductivity (ECe) values of 4-6 dS/m and irrigation water salinity (ECw) of 2-3 dS/m, conditions that would reduce yields by 40-60% on moderate-tolerance rootstocks and cause vine failure on sensitive genotypes. This exceptional tolerance is achieved through highly efficient ion exclusion (maintaining leaf Cl⁻ below 0.3% and Na⁺ below 0.15% dry weight under high external salinity), superior osmotic adjustment capacity (accumulating compatible solutes to maintain turgor against soil osmotic potentials of -0.3 to -0.5 MPa from salinity alone), and sustained K⁺/Na⁺ homeostasis under Na⁺ loading conditions. High salt-tolerance rootstocks derive primarily from V. champinii and certain V. berlandieri × V. rupestris selections adapted to saline or alkaline soils of origin.
High_Salt_Tolerance_Rootstock_Role captures the exceptional salt adaptation of V. champinii (Ramsey, Dog Ridge, Freedom) and certain V. berlandieri × V. rupestris crosses (1103P, 140Ru) that enable viticulture under conditions approaching the salinity limits for glycophytic crops. The ECe threshold of 4-6 dS/m for economically viable production on high-tolerance rootstocks compares favorably with moderately salt-tolerant agronomic crops (cotton, barley) but remains below true halophyte tolerance. Water quality degradation from aquifer depletion and climate change is expanding the area requiring high salt-tolerance rootstocks beyond traditional arid-zone viticulture.
Unterlage für stark versalzte Böden
high salt tolerance rootstock
highly salt tolerant rootstock
hoch salztolerante Unterlage
sehr salztolerante Unterlage
severe salinity tolerant rootstock
Walker RR et al. (2002) Rootstock effects on salt tolerance of irrigated field-grown grapevines, Australian Journal of Grape and Wine Research 8: 146-155 Zhang X et al. (2002) Rootstock influence on salt tolerance of grapevines, Journal of the American Society for Horticultural Science 127: 283-289 Fisarakis I et al. (2001) The effect of salinity on growth and physiology of four grapevine rootstocks, Scientia Horticulturae 89: 123-134
High Salt Tolerance Rootstock Role
Grenache exhibits high stomatal conductance disposition with maximum gs of 380-450 mmol/m²/s under well-watered conditions, supporting photosynthetic rates of 14-18 µmol CO₂/m²/s. Semillon and Shiraz show similarly high conductance (350-420 mmol/m²/s), maintaining open stomata even as soil moisture declines. High-gs cultivars typically show anisohydric behavior, with midday leaf water potential declining to -1.4 to -1.8 MPa under moderate stress while maintaining gs above 150 mmol/m²/s. The correlation between maximum gs and drought sensitivity is moderate (r = 0.55-0.70), as high conductance capacity may be combined with responsive closure mechanisms in some genotypes.
A stomatal conductance disposition that is the inherited capacity of a Vitis vinifera plant to maintain high rates of stomatal gas exchange (typically gs greater than 300 mmol H₂O/m²/s under well-watered conditions), enabling high photosynthetic rates but requiring substantial water supply and often associated with anisohydric water use behavior.
High stomatal conductance disposition maximizes carbon gain under favorable water conditions but increases vulnerability to hydraulic failure when water supply becomes limiting. The trait is advantageous in irrigated viticulture where water is managed to match high transpirational demand, enabling rapid canopy development and high yield potential. In rain-fed systems, high-gs cultivars may experience severe stress during dry periods. Selection for high gs is generally associated with selection for productivity in well-watered environments.
high gs trait
high transpiration capacity
hohe Stomata-Leitfähigkeit
hohe stomatäre Öffnung
open stomata disposition
starke Transpirationskapazität
Soar CJ et al. (2006) Grape vine varieties Shiraz and Grenache differ in their stomatal response to VPD: apparent links with ABA physiology and gene expression in leaf tissue. Australian Journal of Grape and Wine Research 12:2-12 Prieto JA et al. (2010) Stomatal behavior of different grapevine cultivars in response to soil water status and air water vapor pressure deficit. Journal International des Sciences de la Vigne et du Vin 44:9-20
High Stomatal Conductance Disposition
Grenache exhibits high sugar accumulation disposition, routinely achieving 26-30°Brix in Mediterranean climates with potential alcohol of 15-17% v/v if fermented dry. Zinfandel shows similarly high accumulation (25-28°Brix) with characteristic raisinification of exposed berries further concentrating sugars. Muscat of Alexandria reaches 24-28°Brix in warm climates, supporting production of sweet fortified wines. High-sugar cultivars typically show large leaf area to fruit weight ratios (1.2-1.8 m²/kg), efficient VvHT (hexose transporter) expression in berry flesh, and sustained photosynthetic activity through late ripening. Sugar accumulation rates in high-capacity cultivars reach 0.25-0.35°Brix/day during peak ripening versus 0.15-0.20°Brix/day in moderate accumulators.
A sugar accumulation disposition that is the inherited capacity of a Vitis vinifera plant to achieve elevated berry sugar concentrations (typically exceeding 24°Brix) at harvest maturity, resulting from high source-sink ratio, efficient phloem transport, strong berry sink activity, and/or extended ripening period under favorable thermal conditions.
High sugar accumulation disposition was historically valued for producing full-bodied wines and ensuring complete fermentation, but contemporary trends toward moderate alcohol wines have shifted preferences in some markets. The trait may create challenges when sugar ripeness outpaces phenolic and flavor maturity, a phenomenon increasingly common under climate warming. Management strategies including early harvest, crop load adjustment, and shading can moderate excessive sugar accumulation in genetically predisposed cultivars.
elevated sugar trait
high Brix capacity
high potential alcohol disposition
hohe Mostgewichte
hohe Zuckerakkumulation
starke Zuckereinlagerung
Sadras VO, Moran MA (2012) Elevated temperature decouples anthocyanins and sugars in berries of Shiraz and Cabernet Franc. Australian Journal of Grape and Wine Research 18:115-122 Palliotti A et al. (2014) Changes in vineyard establishment and canopy management urged by earlier climate-related grape ripening: a review. Scientia Horticulturae 178:43-54
High Sugar Accumulation Disposition
Characterization of High_Vigor_Disposition at AWRI Adelaide documented growth parameters in cultivars classified as high-vigor based on industry experience and research trials. Reference cultivars 'Carignan', 'Barbera', 'Silvaner', and 'Malbec' were grown under standardized conditions (own-rooted, 2.5 × 2.0 m spacing, moderate irrigation) with detailed growth measurements over three seasons. High_Vigor_Disposition cultivars showed consistent characteristics distinguishing them from moderate and low-vigor types. 'Carignan' exhibited archetypal High_Vigor_Disposition with mean shoot length of 265 cm (range 230-310 cm across seasons), internode length of 9.5 cm, individual leaf area of 245 cm², lateral shoot production of 12.5 laterals per primary shoot with total lateral length of 195 cm, and pruning weight of 2.65 kg/vine. Growth cessation occurred in week 17-18 (late February in Southern Hemisphere), 4-5 weeks after veraison, indicating continued vegetative growth during fruit ripening. 'Barbera' showed similar high vigor (255 cm shoots, 2.85 kg/vine pruning weight) with particularly strong lateral development. The Ravaz index (yield/pruning weight) for High_Vigor_Disposition cultivars averaged 3.2 (below the optimal 5-10 range), indicating vegetative dominance over reproductive development. Canopy management trials confirmed that High_Vigor_Disposition cultivars required 3-4 hedging passes per season to maintain fruit zone light exposure above 30% of ambient, compared to 1-2 passes for moderate-vigor types. Training system comparison showed that High_Vigor_Disposition cultivars performed best on divided canopy systems (GDC, Scott Henry) that accommodate total canopy leaf area of 22-28 m² per vine while maintaining adequate fruit zone microclimate.
A High_Vigor_Disposition is a Shoot_Vigor_Disposition that inheres in Vitis vinifera shoots and is realized through strong, sustained vegetative growth producing long shoots (typically > 200 cm), extended internodes (> 8 cm mean length), large leaves (> 200 cm² individual area), and prolific lateral shoot development, resulting in dense canopies that require intensive management to maintain fruit quality. This disposition represents the upper range of grapevine vigor, characteristic of cultivars with high inherent growth capacity that, under favorable conditions, produce abundant vegetative biomass potentially at the expense of reproductive development and fruit quality. High-vigor cultivars typically show extended growth duration (active shoot elongation continuing past veraison), delayed growth cessation, and strong regrowth response to hedging, creating ongoing canopy management challenges. The viticultural implications of High_Vigor_Disposition are substantial—such cultivars require large canopy training systems (e.g., sprawl, GDC, Scott Henry) or divided canopy systems to accommodate growth, wide vine spacing to prevent excessive competition, careful irrigation and fertilization management to avoid stimulating already excessive growth, and intensive canopy management including multiple hedging passes, shoot thinning, and leaf removal to maintain fruit zone light exposure and air circulation. High-vigor cultivars are typically well-suited to sites with growth-limiting factors (low fertility, limited water, cool climate) that naturally constrain vigor expression, but may produce problematic canopies on fertile, well-watered sites.
High_Vigor_Disposition is_a Shoot_Vigor_Disposition representing the upper vigor category. Typical characteristics include shoots > 200 cm, internodes > 8 cm, pruning weight > 2.0 kg/vine (site-dependent), extended growth duration past veraison, and Ravaz index < 5. Examples include 'Carignan', 'Barbera', 'Silvaner', 'Malbec', 'Carménère'. Management implications include need for divided canopy or large single-canopy systems, wide spacing, intensive hedging and shoot thinning, and careful water/nutrient management. High vigor is advantageous on limiting sites but problematic on fertile, well-watered sites.
Hohe-Wuchsstärke-Disposition
Starkes-Vegetatives-Wachstum
excessive vigor
strong vegetative growth
vigorous growth
Überschüssige-Wuchskraft
Smart RE (1985) Principles of grapevine canopy microclimate manipulation with implications for yield and quality: a review, American Journal of Enology and Viticulture 36:230-239 Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review, American Journal of Enology and Viticulture 60:251-268 Dry PR (2000) Canopy management for fruitfulness, Australian Journal of Grape and Wine Research 6:109-115
High Vigor Disposition
1103 Paulsen (V. berlandieri × V. rupestris) confers high vigor to Nero d'Avola in Sicily, producing pruning weights of 0.8-1.2 kg/m and supporting yields of 12-18 t/ha on calcareous clay soils. Ruggeri 140 (V. berlandieri × V. rupestris) imparts high vigor to Catarratto in southern Italy with shoot lengths regularly exceeding 2.5 m and requiring extensive summer canopy management. St. George (V. rupestris du Lot) confers very high vigor to Cabernet Sauvignon in Napa Valley, with pruning weights reaching 1.0-1.5 kg/m on deep alluvial soils.
A vigor-modifying rootstock role that inheres in a grafted rootstock by virtue of its capacity to substantially increase vegetative growth of the scion partner relative to own-rooted vines or moderate-vigor references. High-vigor rootstocks typically increase scion pruning weight by 20-50% above moderate-vigor standards, producing shoot growth of 1.5-3.0 m per season and canopy leaf area indices of 3.0-5.0 or greater. This vigor enhancement is mediated by extensive root system development (root density often exceeding 2.5 km/m³ soil), high root hydraulic conductance, elevated cytokinin export via xylem sap, and efficient nitrogen and water uptake. High-vigor rootstocks are preferentially selected for low-fertility soils, drought-prone sites, and situations where rapid vineyard establishment and high yield targets are priorities.
High_Vigor_Rootstock_Role is context-dependent the same rootstock may express high vigor on deep fertile soils but only moderate vigor on shallow or nutrient-poor substrates. Vigor classification typically uses own-rooted Kober 5BB or SO4 as reference benchmarks. Excessive vigor from high-vigor rootstocks can delay fruit ripening, increase disease pressure through dense canopy shading, and dilute phenolic concentration in red wines, requiring compensatory canopy management interventions such as hedging, shoot thinning, and leaf removal.
Starkwuchsunterlage
high-vigor rootstock
invigorating rootstock
starkwüchsige Unterlage
strong-growing rootstock
vigorous rootstock
wuchsstarke Unterlage
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 285-310 Dry N (2007) Grapevine rootstocks: selection and management for South Australian vineyards, Lythrum Press Whiting JR (2004) Grapevine rootstocks, in Viticulture Vol 1 Resources, Winetitles pp.: 167-188
High Vigor Rootstock Role
Pinot Noir clone 114 bears a High_Yield_Clone_Role relative to the Dijon quality clones, documented in French trials as producing 30-40% higher yields with larger clusters and berries—suited to Champagne base wine production where volume is valued over concentration. Chardonnay clone 95 serves in a High_Yield_Clone_Role in California production, selected for productivity exceeding 8 tonnes/hectare in commercial settings versus 4-6 tonnes for quality-focused clones 76 and 96. Sangiovese clone R10 bears a High_Yield_Clone_Role in Tuscan evaluation, yielding 12-15 tonnes/hectare versus 6-8 tonnes for quality clones—appropriate for Chianti IGT production but discouraged for DOCG appellations with yield restrictions.
A performance clone role that inheres in a grapevine clone characterized by above-average yield potential relative to other clones of the same variety, typically producing 20-50% more fruit per vine than quality-focused clones under comparable conditions. High-yield clones achieve elevated production through various mechanisms—larger clusters (more berries per cluster, larger berry size), more fruitful buds (higher clusters per shoot), more vigorous growth supporting greater fruit load, or combinations thereof. Selection for high yield historically served bulk wine production, where volume economics favored productivity over concentration, though high-yield clones may find contemporary application in entry-level wine tiers, grape juice production, or table grape markets where fruit volume drives profitability. High-yield clones typically exhibit trade-offs with concentration and quality metrics—larger berries have lower skin-to-juice ratios reducing phenolic extraction, higher yields dilute flavor compounds per unit fruit, and extended ripening required to achieve equivalent sugar levels may compromise acid balance. The relationship between yield and quality is complex and site-dependent well-managed high-yield clones on appropriate sites may achieve commercial quality while maximizing productivity.
High_Yield_Clone_Role captures the productivity-optimized end of the clone performance spectrum, where yield maximization drives selection and production decisions. The characterization is relative within variety—a "high-yield" Pinot Noir clone may produce 6-8 tonnes/hectare while a "high-yield" Airén clone might exceed 20 tonnes/hectare, reflecting variety-specific production potential. High-yield clones emerged from early systematic selection programs that prioritized post-phylloxera vineyard productivity recovery, with quality focus developing as wine markets evolved toward differentiation and premiumization. Contemporary use of high-yield clones reflects market segmentation bulk wine, brandy distillation, grape juice, and value-tier table wine production all favor productivity while premium wine production increasingly selects quality-oriented clones despite lower yields.
Ertragsklon
Hochertragsklon
Mengenklon
high-production clone
high-yield clone
productive clone
produktiver Klon
volume clone
Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 ENTAV-INRA (2007) Catalogue des Variétés et Clones Huglin P, Schneider C (1998) Biologie et Écologie de la Vigne, Lavoisier Intrieri C, Filippetti I (2000) Vineyard management and grape quality, ISHS Acta Horticulturae 526: 27-36
High Yield Clone Role
A high-yield primary reproductive shoot on Vitis vinifera cv. Riesling from node 9 on fruiting cane in Mosel Valley, 168cm length with 21 leaves totaling approximately 3360 cm² leaf area, bearing 3 well-developed clusters with 68, 72, and 65 berries (205 total berries), estimated total shoot crop 615g at projected maturity (3 clusters × 205g average cluster weight for this variety and conditions), representing high natural fruitfulness of Riesling combined with favorable mid-cane position and adequate shoot vigor, managed conservatively with no cluster thinning (accepting high yield appropriate for this quality-oriented but economically viable Mosel production), adequate leaf area-to-fruit ratio (14.6 cm² per gram, within acceptable range for this variety and region) supporting complete maturation to target 19-21° Brix with characteristic Riesling acidity and aromatic profile.
A primary reproductive shoot bearing multiple clusters producing substantial fruit weight, characterized by 2-4 inflorescences (variety dependent, with some high-fruitfulness cultivars occasionally showing 5+ clusters), large well-filled clusters with high berry counts (50-100+ berries per cluster typical), total shoot crop load 400-800g or more at maturity, and requirement for adequate shoot vigor and leaf area (typically 18+ leaves, 2800-4200 cm² total) supporting heavy fruit load without excessive strain or maturation problems. High-yield primary reproductive shoots represent upper productivity range from individual positions, occurring on vigorous shoots in favorable locations, inherently high-fruitfulness varieties (Riesling, Gewürztraminer, some Pinot clones), well-managed balanced vines with adequate capacity, or situations where deliberate high-yield objectives (bulk production, table grapes) encourage maximum productivity per shoot through cultivar selection and minimal cluster thinning approaches.
High-yield primary reproductive shoots provide maximum productivity per position, with their heavy crop loads contributing substantially to whole-vine yields and economic viability. However, high-yield shoots create management challenges including potential maturation delays (heavy sink demand consuming resources), possible quality compromise (dilution effects if excessive, inadequate leaf area-to-fruit ratios), vine stress (resource depletion affecting current and future seasons), and practical difficulties (heavy clusters increasing rot susceptibility, harvest logistics). Appropriate high-yield shoot situations include inherently high-fruitfulness varieties where multiple clusters are normal and expected (Riesling, Gewürztraminer, some Muscat clones showing 2-3 cluster standard), bulk production systems accepting yield priority over quality extremes, vigorous sites and vines with capacity supporting heavy loads, and table grape production where large clusters and high yields are primary objectives. Quality-focused wine production typically reduces high-yield shoots through cluster thinning, recognizing that moderate yields often produce superior fruit composition and wine character.
Maximaleistungs-Trieb
heavy-cropping primary shoot
hocherträglicher primärer Fruchttrieb
maximum productivity shoot
multi-cluster high-yield shoot
Based on Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115.
High Yield Primary Reproductive Shoot
The Romanée-Conti Pinot Noir selection, maintained since the 18th century at the famous Burgundy estate, bears a Historic_Clone_Role with documented provenance traceable through estate records—considered among the finest Pinot Noir genotypes and foundation for premium clone development. The "Mount Eden" Chardonnay and Pinot Noir selections (California) serve in Historic_Clone_Roles as documented material brought from Burgundy in the 1940s and maintained at the historic estate, their provenance established through oral history and confirmed through performance. The "old Inglenook" Cabernet Sauvignon selection bears a Historic_Clone_Role in Napa Valley, documented material from the famous 19th-century estate representing pre-Prohibition California viticulture and maintained as heritage material distinct from modern nursery clones.
A heritage clone role that inheres in a grapevine clone with documented historical provenance, traceable to specific historic vineyards, estates, or time periods through records, oral history, or scientific verification. Historic clones carry documented connections to viticultural history—material from famous vineyards (Romanée-Conti, Château d'Yquem), selections associated with historical figures or events, or genotypes verified through genetic analysis as surviving from documented historical periods. The historic designation requires some form of provenance documentation, distinguishing these clones from anonymous old material of unknown origin. Historic clone significance extends beyond genetic material to cultural heritage these selections embody viticultural history and connect contemporary producers to their predecessors. Historic clone conservation serves both practical (maintaining potentially valuable genotypes) and cultural (preserving viticultural heritage) objectives. Scientific authentication using molecular markers enables verification of claimed historic provenance, confirming or refuting historical narratives about clone origins and movements.
Historic_Clone_Role captures clones whose value derives substantially from documented historical connections rather than solely from agronomic performance. The documentation requirement distinguishes historic clones from merely old material claims of historic provenance require substantiation through records, credible oral history, or scientific authentication. Historic clone conservation represents the intersection of viticulture with cultural heritage preservation, with material maintained not only for potential agricultural value but as living artifacts of viticultural history. The wine industry's interest in historic clones reflects premiumization trends valuing authenticity, provenance, and tradition wines from documented historic selections command market attention and sometimes price premiums reflecting their heritage significance.
documented clone
dokumentierter Klon
heritage selection
herkunftsbelegter Klon
historic clone
historischer Klon
provenanced clone
This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Pelsy F (2010) Molecular diversity within grapevine varieties, Heredity 104: 331-340 Wolpert JA et al. (1994) Selection of Pinot Noir clones, American Journal of Enology and Viticulture 45: 356-363
Historic Clone Role
Historical acreage data for Grüner Veltliner in Austria documents a remarkable expansion trajectory: approximately 5,000 ha in 1950 (minor variety, largely undifferentiated in mixed plantings), rapid expansion to 13,500 ha by 1970 (driven by the Austrian Weinbaukataster rationalization and government-encouraged varietal replanting), peak of approximately 18,800 ha by 1999, modest decline to approximately 13,500 ha by 2023 (reflecting both quality-focused reduction in total Austrian vineyard area and selective grubbing of lower-quality sites) — the acreage arc tracing the cultivar's transformation from bulk-wine producer to quality-focused national champion. Historical acreage data for Malbec illustrates a dramatic geographic redistribution: approximately 60,000 ha in France in 1958 (concentrated in Cahors and the southwest), declining to approximately 5,500 ha by 2020 following the 1956 frost devastation and progressive replacement by Merlot, contrasted with explosive expansion in Argentina from approximately 10,000 ha in 1990 to over 46,000 ha by 2020, as Mendoza Malbec became Argentina's internationally recognized flagship variety — the inverse acreage trajectories in France and Argentina representing one of the most dramatic cultivar importance transfers in viticultural history.
Historical acreage data is a historical importance entity that documents the recorded vineyard area planted to specific grape cultivars at defined historical time points, derived from governmental vineyard census records, regional viticultural surveys, phylloxera-era replanting inventories, appellation registration databases, and international statistical compilations, expressed in hectares (ha) or regional equivalents (acres, Joch, arpents) with the date and geographic scope of each measurement, and providing the quantitative foundation for assessing the economic trajectory of cultivar importance — expansion indicating growing commercial demand and viticultural confidence, contraction indicating displacement by competing cultivars, disease impact, or market decline, and stability indicating entrenched regional establishment.
Historical acreage data provides the most objective quantitative measure of cultivar economic importance over time, as vineyard planting decisions represent substantial long-term capital investment (20-30 year productive vineyard lifespan) and therefore reflect genuine commercial conviction about a cultivar's value and market potential. The sources for historical acreage data vary dramatically in reliability and resolution — modern national vineyard registers (the Austrian Weinbaukataster, the French Casier Viticole Informatisé, the Australian Wine Australia register) provide cultivar-specific, georeferenced acreage data of high accuracy, while pre-20th-century sources range from systematic governmental surveys (the French phylloxera-era inventories of the 1880s-1900s provide exceptionally detailed cultivar-specific data) to fragmentary regional estimates derived from tax records, estate inventories, and traveler accounts. The Anderson and Aryal (2013) global database of winegrape plantings provides the most comprehensive cross-national compilation, documenting cultivar-specific acreage for approximately 50 countries from the mid-20th century to the present, though data quality and temporal coverage vary substantially between wine-producing nations. The interpretation of acreage trends requires contextual knowledge — acreage expansion may reflect increasing quality recognition (Grüner Veltliner in Austria) or undiscriminating volume production (Airén's 300,000+ ha in La Mancha reflecting irrigated bulk wine rather than quality ambition), while acreage contraction may indicate cultivar decline (Gouais Blanc's medieval-to-modern disappearance) or quality-focused vineyard rationalization (the selective grubbing of low-quality sites while retaining premium plantings). The EU vine-pull schemes (1988-1996 and 2008-2011), which paid growers to remove vineyards and reduce overproduction, created artificial discontinuities in European acreage data that must be distinguished from market-driven cultivar dynamics.
Rebflächenentwicklung
acreage records
historical planting area
historical vineyard census
historische Anbaufläche
historische Anbaustatistik
historische Bestockungsdaten
historische Rebflächendaten
planting statistics history
vineyard area history
Anderson K, Aryal NR (2013) Which winegrape varieties are grown where? A global empirical picture. University of Adelaide Press, pp.: 1-700 Anderson K, Nelgen S (2020) Which Winegrape Varieties are Grown Where? Revised edition, University of Adelaide Press OIV (2023) State of the World Vine and Wine Sector, International Organisation of Vine and Wine Statistik Austria (2023) Weinbauerhebung, Bundesanstalt Statistik Österreich
Historical Acreage Data
Historical document reference for the earliest Riesling attestation specifies: document type — estate cellar account book (Kellerbuch) originator — cellar administration of Count Johann IV of Katzenelnbogen date — March 13, 1435 archival repository — Hessisches Hauptstaatsarchiv Wiesbaden, Abt. 116, Nr. 1135 language — Early New High German relevant passage — "item iii s. umb seczs amen von rießlingen in die wingarten" (item 3 shillings for six rootstocks of Riesling for the vineyards) scholarly identification — Josef Staab, 1990, confirmed by Helmut Becker and subsequent researchers interpretive confidence — high (unambiguous cultivar name in a precisely dated and located document). Historical document reference for Tokaji Aszú production origins specifies the 1630 Szepsi Laczkó Máté account documenting selective botrytized berry harvesting at the Oremus vineyard of the Rákóczi estate, housed in the Magyar Nemzeti Levéltár, Rákóczi family archives — though the scholarly dating and interpretation of this reference remains debated, with some historians attributing the Aszú innovation to later decades.
Historical document reference is a documentation source entity that documents the primary archival, literary, and administrative sources from which historical information about grape cultivars and viticulture has been derived, encompassing vineyard inventories and estate records (Urbare, terriers, cadastres), legal documents (land grants, tithes, trade regulations, appellation decrees), ecclesiastical records (monastic cellar books, chapter records, tithe registers), literary and travel accounts (natural histories, regional descriptions, diplomatic correspondence), cartographic sources (vineyard maps, cadastral plans), and iconographic sources (manuscript illuminations, paintings, engravings depicting vines and winemaking), specifying for each referenced document the author or originating institution, date or date range, archival repository and classification number, language, and the specific information extracted relevant to cultivar history.
Historical document reference documentation applies the methodology of historical source criticism (Quellenkritik) to viticultural history, demanding that each archival source be evaluated for authenticity (is the document genuine and correctly attributed?), dating reliability (is the assigned date secure or estimated?), provenance (is the chain of custody from creation to current repository documented?), and interpretive clarity (does the relevant passage unambiguously refer to the cultivar or practice claimed?). The principal categories of historical documents relevant to viticultural history each carry characteristic strengths and limitations — estate inventories and Urbare (medieval land registers) provide the most systematic and geographically specific information but typically use vernacular cultivar names that may resist modern identification, legal documents (trade regulations, quality ordinances, appellation decrees) reveal the regulatory and economic context of cultivation but may refer to wine types rather than specific cultivars, ecclesiastical records (monastic cellar books, tithe registers) provide long continuous series from institutions with strong record-keeping traditions but are geographically concentrated in regions with significant monastic viticulture, and literary-travel accounts provide external quality assessments and comparative observations but may reflect the author's limited viticultural knowledge or cultural biases. The language dimension of historical document references is particularly significant for Central European viticultural history, where documents exist in Latin, German, Hungarian, Czech, and various local dialects, with cultivar names sometimes appearing in different languages within a single document and requiring specialized linguistic competence for accurate interpretation. The distinction between a document that mentions a cultivar by name (direct attestation) and one that describes characteristics matching a known cultivar without naming it (indirect attestation) affects the confidence level of the reference — direct attestations provide stronger evidence but are more susceptible to synonymy-homonymy confusion, while indirect attestations avoid naming problems but introduce identification uncertainty. The progressive digitization of European archival collections (the Monasterium.net initiative for monastic records, national digital archive programs) is expanding access to primary sources that were previously accessible only through physical archival research.
Archivquelle
Primärquelle
archival reference
archival source
documentary evidence
historical primary source
historische Dokumentenreferenz
historische Quellenangabe
primary historical source
urkundlicher Nachweis
Staab J (2015) Die Erstnennung des Riesling von 1435. Schriften zur Weingeschichte 188: 1-20 Rácz I (2012) The history of Tokaj wine region. In Sweet, Reinforced and Fortified Wines, Wiley-Blackwell, pp.: 1-15 Droste R (2001) Quellen zur Geschichte des Weinbaus in Deutschland. Schriften zur Weingeschichte 137: 1-45 Csendes P, Opll F (2003) Wien. Geschichte einer Stadt, Böhlau Verlag
Historical Document Reference
Historical first mention for Riesling documents the earliest known reference in a 1435 inventory of the Rheingau wine cellar of Count Johann IV of Katzenelnbogen, recording the purchase of "seczs amen von rießlingen" (six rootstocks of Riesling) on March 13, 1435, stored in the Hessisches Hauptstaatsarchiv Wiesbaden — the document unambiguously identifies the cultivar by a recognizable form of its modern name in a precisely dated and geographically located context, establishing a minimum documented age of nearly six centuries. Historical first mention for Grüner Veltliner is considerably more recent and less definitive — the name "Grüner Muskateller" appears in Johann Burger's 1837 ampelographic catalogue from Klosterneuburg, with the modern designation "Grüner Veltliner" first recorded in Franz Trummer's 1841 Systematische Classification der Traubenarten, though the relationship between these named cultivars and the modern Grüner Veltliner requires interpretive caution. Historical first mention for Blaufränkisch records the name in early 19th-century Austrian and Hungarian ampelographic literature, with the synonym "Kékfrankos" (literally "blue Frankish") appearing in Hungarian sources from approximately the same period.
Historical first mention is an origin record entity that documents the earliest datable textual reference to a grape cultivar by name or recognizable description in archival, literary, or scientific sources, specifying the document type (vineyard inventory, land register, ampelographic treatise, legal document, literary text, ecclesiastical record), the date or date range of the document, the precise textual passage containing the reference, the language and orthographic form of the cultivar name as recorded, the geographic location referenced, the archival repository housing the original document, and the scholarly provenance of the identification (who identified the reference, when, and with what interpretive caveats), providing the documentary anchor point that establishes the minimum age of a cultivar's recognized existence and its earliest known geographic and cultural context.
Historical first mention documentation requires careful methodological rigor because the identification of grape cultivars in pre-modern texts is fraught with interpretive challenges that can lead to false early attestations or missed genuine references. The principal difficulties include orthographic variation (medieval and early modern spelling was unstandardized, producing variant forms that may or may not refer to the same cultivar — "Rießling," "Rüssling," "Rissling," "Riesling" represent plausible orthographic variants of a single name, but "Riesling" and "Rißling" could also be unrelated terms), synonymy and homonymy (the same cultivar may appear under different names in different documents, while the same name may refer to different cultivars in different regions or periods), and the distinction between a cultivar name and a wine type or place name (many apparent cultivar references in medieval documents may actually refer to wines named after their origin rather than their grape variety). The scholarly evaluation of first mention claims requires assessment of the document's authenticity (is it a genuine period document or a later copy or forgery?), dating reliability (is the date precisely established or estimated?), identification confidence (how certain is it that the name refers to the modern cultivar?), and interpretive context (does the document reveal anything about the cultivar's status — common or rare, valued or ordinary, local or recently introduced?). The temporal distribution of first mentions across European cultivars reveals a strong documentation bias — cultivars from regions with rich archival traditions (France, Germany, Italy) tend to have earlier documented first mentions than cultivars from regions with less systematic record-keeping, even though the cultivars themselves may be equally ancient. The transition from vernacular oral tradition to written documentation means that most cultivars existed for generations or centuries before their first written mention, and the documented first mention therefore represents a minimum age rather than an actual origin date.
Ersterwähnung
Erstnennung
earliest attestation
earliest mention
erste schriftliche Nennung
first documentary reference
first written record
früheste urkundliche Erwähnung
historical first reference
älteste Erwähnung
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes: A Complete Guide to 1,368 Vine Varieties, Allen Lane Staab J (2015) Die Erstnennung des Riesling von 1435. Schriften zur Weingeschichte 188: 1-20 Trummer F (1841) Systematische Classification aller bekannten Traubensorten, Graz Burger J (1837) Die Rebensorten der Klosterneuburger Sammlung, Klosterneuburg
Historical First Mention
Historical importance for Pinot Noir documents an unparalleled combination of cultural prestige, historical depth, and continuing relevance — continuous documented cultivation in Burgundy since at least the 14th century (Cistercian monastic records), foundation cultivar of the most celebrated and expensive wines in history (Romanée-Conti, La Tâche, Chambertin), genetic progenitor of numerous major cultivars (Chardonnay, Gamay, Aligoté, Melon de Bourgogne through crosses with Gouais Blanc), global benchmark for terroir expression (the Burgundian model of single-vineyard, single-cultivar, site-specific winemaking), and a cultivar whose global acreage has expanded continuously for over a century despite its notoriously demanding viticultural requirements. Historical importance for Grüner Veltliner documents a dramatically different trajectory — negligible historical significance before the mid-20th century, explosive rise to Austrian national identity cultivar status after the 1985 Vinea Wachau founding and the subsequent international recognition of Austrian wine quality, now accounting for approximately 30% of Austrian vineyard area and serving as the defining expression of Austrian viticultural identity on the global stage.
Historical importance is a historical record entity that documents the cultural, economic, and viticultural significance of grape cultivars across historical periods, integrating evidence of past cultivation extent (historical acreage data), documented wine production achievements (historical wine records), and association with celebrated wines and estates (famous wine associations), providing the evaluative dimension of cultivar history that contextualizes the relative standing, prestige trajectory, and cultural impact of individual cultivars within the broader narrative of wine civilization — distinguishing cultivars that have shaped viticultural history and defined wine culture from those of merely local or transient significance.
Historical importance documentation addresses the evaluative and contextual dimension of cultivar history that transforms chronological data into meaningful narrative — the difference between knowing that Riesling was first mentioned in 1435 (a fact) and understanding that Riesling has shaped the identity of German, Alsatian, and Austrian wine culture for centuries (an interpretation). The assessment of historical importance requires integration across multiple evidence domains — cultivation extent (acreage data indicating economic significance), wine production quality (records of exceptional wines demonstrating quality potential), cultural resonance (the cultivar's role in regional identity, artistic representation, and gastro-cultural tradition), scientific significance (the cultivar's role in advancing viticultural and oenological knowledge), and genetic legacy (the cultivar's parentage relationships indicating biological influence on the viticultural gene pool). The temporal trajectory of cultivar importance is rarely linear — many cultivars of great contemporary significance were historically obscure (Grüner Veltliner, Malbec in Argentina, Carménère in Chile), while cultivars of immense historical importance have declined to near-extinction (Noah and other pre-phylloxera American hybrids once widely planted in Europe, Gouais Blanc once ubiquitous in medieval France). The phylloxera crisis and post-phylloxera replanting restructured cultivar importance hierarchies across Europe, as the replanting opportunity was used to eliminate cultivars perceived as inferior and concentrate on those considered commercially optimal — a process that dramatically reduced European cultivar diversity while consolidating the dominance of the cultivars now considered "noble" or "international." The concept of "noble grapes" — the informal hierarchy placing Cabernet Sauvignon, Merlot, Pinot Noir, Chardonnay, Riesling, and Sauvignon Blanc at the apex of viticultural prestige — is itself a historically constructed category reflecting 20th-century commercial and critical preferences rather than any intrinsic biological hierarchy, and is increasingly challenged by the contemporary interest in indigenous, obscure, and rediscovered cultivars.
Sortenbedeutung
cultivar significance
cultural importance
geschichtliche Relevanz
historical significance
historical standing
historische Bedeutung
kulturelle Bedeutung
viticultural heritage
vitikulturelles Erbe
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes: A Complete Guide to 1,368 Vine Varieties, Allen Lane, pp.: xv-xxxii Johnson H (1989) Vintage: The Story of Wine, Simon and Schuster, pp.: 1-480 Unwin T (1991) Wine and the Vine: An Historical Geography of Viticulture and the Wine Trade, Routledge, pp.: 1-400
Historical Importance
The historical name Argitis documented by Columella in De Re Rustica (c. 65 CE) and Pliny the Elder in Naturalis Historia (77 CE) designates a white grape cultivar cultivated in the Roman Empire whose identity remains debated among ampelographers — proposed modern equivalents include Greco di Tufo or Trebbiano Toscano based on geographic continuity and morphological comparison, but definitive molecular confirmation is impossible without surviving authenticated material from the Roman period, illustrating the epistemic limits of historical name resolution for pre-medieval cultivar designations.
A cultivar designation that was applied to a grapevine cultivar during a specific historical period, documented in primary sources from that era such as agricultural treatises, vineyard inventories, tax records, monastic accounts, or early ampelographic works, and whose temporal context provides evidence for the cultivar's presence and cultural significance in a particular region and period.
Historical_Name differs from Historical_Synonym in its primary function as a temporal marker rather than a nomenclatural cross-reference. While historical synonyms are resolved names confirmed as equivalent to a modern prime name through molecular or morphological evidence, historical names may remain unresolved — their cultivar identity uncertain or debated — and their primary informational value lies in documenting the presence, distribution, and cultural significance of viticulture in specific historical periods: subclasses distinguish ancient (pre-medieval), medieval, and pre-phylloxera historical names, reflecting the major periodization of European viticultural history and the progressively increasing resolution of cultivar identification from antiquity to the 19th century.
archival variety designation
historically documented cultivar name
historischer Name
period name
urkundlicher Sortenname
zeitgenössische Sortenbezeichnung
Columella (c. 65 CE) De Re Rustica Book III Unwin T (2012) Wine and the Vine: An Historical Geography of Viticulture and the Wine Trade, Routledge pp: 65-128 McGovern PE (2003) Ancient Wine: The Search for the Origins of Viniculture, Princeton University Press
Historical Name
Historical record for Furmint integrates geographic origin (Tokaj-Hegyalja region of present-day Hungary, with debated earlier Balkan or Adriatic origin), documentary first mention (earliest unambiguous reference in the 1571 Tokaj vineyard inventory of the Rákóczi estate, though the name "Furmint" appears in Latin documents from 1623), etymology (disputed — possibly from French "froment" [wheat] referring to the golden-ripe berry color, or from the Hungarian "fürmint" as a phonetic adaptation of a South Slavic term), and the cultivar's central role in the development of the Tokaji Aszú sweet wine tradition documented from 1630 onward, establishing the world's first formally classified wine region in 1730. Historical record for Grüner Veltliner documents the cultivar's relatively recent historical emergence — no documented references before the 19th century, genetic parentage identified as Traminer × St. Georgen (an otherwise obscure cultivar from Burgenland), first ampelographic description under the name "Grüner Muskateller" in the 1855 Trummer catalogue, gradual expansion through Lower Austria in the early 20th century, and the dramatic post-1985 quality revolution that transformed it from a bulk-wine variety into Austria's flagship cultivar.
Historical record is a grapevine information entity that documents the diachronic knowledge about grape cultivars, viticulture, and winemaking accumulated through historical, archaeological, and archival sources, encompassing origin and provenance records, documentary first mentions, etymological analyses of cultivar names, ampelographic classification history, and the cultural-economic narratives that contextualize individual cultivars within the broader history of viticulture, providing the temporal depth dimension that connects contemporary grapevine science to millennia of human-vine coevolution and enables the reconstruction of cultivar migration routes, selection histories, and the cultural forces that shaped the modern viticultural landscape.
Historical record documentation for VVO addresses the temporal dimension that is absent from synchronic scientific characterization — a cultivar's genetic profile, morphological description, and phenological behavior describe what it is now, while historical records document how it came to be what and where it is, the human decisions that shaped its selection and distribution, and the cultural contexts that determined its significance. The epistemological challenges of viticultural history are substantial — pre-modern cultivar identification relied on vernacular names that were often duplicated across regions (synonymy), applied inconsistently to multiple cultivars (homonymy), and recorded in documents of uncertain date and provenance, making the reconstruction of cultivar histories an exercise in combining linguistic analysis, archival research, ampelographic comparison, and increasingly, archaeogenomic evidence. The VVO Historical_Record hierarchy distinguishes between origin records (geographic provenance and migration history), documentary first mentions (the earliest datable textual references), and etymology (the linguistic history of cultivar names) — three complementary approaches to reconstructing cultivar history that often yield convergent but sometimes contradictory evidence. The integration of modern molecular parentage analysis with historical documentation has revolutionized viticultural history — the demonstration that Cabernet Sauvignon is a Cabernet Franc × Sauvignon Blanc cross, that Chardonnay descends from Pinot × Gouais Blanc, and that Furmint is genetically related to several Hungarian and Balkan cultivars has either confirmed, refined, or overturned historical origin narratives based on documentary evidence alone. The archaeological dimension — including archaeobotanical identification of Vitis vinifera seeds and wood, residue analysis of ancient vessels for tartaric acid and other wine markers, and ancient DNA extraction from archaeological grape material — is progressively extending the documented history of viticulture beyond the written record into prehistory.
Geschichtsdokumentation
cultivar historical data
diachronic cultivar documentation
geschichtlicher Herkunftsnachweis
historical documentation
historical provenance record
historische Aufzeichnung
historische Sortendokumentation
viticultural history record
vitikulturhistorische Daten
McGovern PE (2003) Ancient Wine: The Search for the Origins of Viniculture, Princeton University Press, pp.: 1-40 This P et al. (2006) Historical origins and genetic diversity of wine grapes. Trends in Genetics 22(9): 511-519 Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes: A Complete Guide to 1,368 Vine Varieties, Allen Lane, pp.: xv-xxxii
Historical Record
The historical synonym Morillon Noir for Pinot Noir appears in Burgundian vineyard records from the 14th century onward, with the earliest documented reference in the 1375 accounts of the Dukes of Burgundy, while the name Plant Doré designated the same cultivar in 16th-century documents from Beaune, and Auvernat Noir was used in Loire Valley records through the 18th century — all three names have been confirmed as synonyms of Pinot Noir through comparison of historical ampelographic descriptions with the modern SSR reference profile (VIVC number 09279), though none remain in active commercial or regulatory usage.
A synonym that originated in a past historical period and was once a widely used or locally accepted designation for a grapevine cultivar but has since been superseded by a modern prime name, persisting primarily in archival documents, historical ampelographic literature, and antiquarian wine references.
Historical_Synonym preserves the temporal dimension of cultivar nomenclature, documenting names that were once current but have fallen out of active usage through standardization, regulatory supersession, or cultural change. These synonyms are essential for interpreting pre-modern viticultural documents, historical wine trade records, and archival vineyard inventories: without historical synonym resolution, the identity of cultivars mentioned in medieval and early modern sources would remain inaccessible to modern researchers. The boundary between historical and current synonyms is gradual rather than categorical — some historical synonyms persist in local oral tradition or appear on heritage wine labels despite no longer being used in official or scientific contexts.
archaic synonym
historical variety name
historische Sortenbezeichnung
historisches Synonym
obsolete cultivar name
veralteter Sortenname
Lavalle J (1855) Histoire et statistique de la vigne et des grands vins de la Côte d'Or Rézeau P (2014) Dictionnaire des noms de cépages de France, CNRS Éditions Robinson J et al. (2012) Wine Grapes, Allen Lane pp: 1-18
Historical Synonym
Historical wine record for Riesling from the Mosel documents an extraordinary archive of quality milestones: the 1783 vintage recorded by Thomas Jefferson as among the finest wines he encountered during his 1788 Rhine journey, the legendary 1921 Trockenbeerenauslese vintages that established TBA as the pinnacle of German wine achievement, the 1959 and 1971 vintages celebrated as benchmarks of Mosel Riesling quality by contemporary critics, the 1976 auction of 1893 Scharzhofberger TBA (then the highest price paid for a German wine), and the modern critical renaissance documented by record scores for the 2001, 2005, and 2019 vintages — the accumulated wine record spanning over two centuries of documented excellence from a single cultivar-region combination. Historical wine record for Blaufränkisch from Mittelburgenland documents a much shorter but accelerating quality trajectory — first DAC designation in 2005 establishing the appellation framework, progressive critical recognition through Austrian wine guides (Falstaff, A la Carte) from the 2000s onward, landmark single-vineyard bottlings from the 2010s establishing Blaufränkisch as Austria's premier indigenous red cultivar, and the 2015 and 2017 vintages widely recognized as reference points for the variety's quality potential.
Historical wine record is a historical importance entity that documents notable wine production achievements, landmark vintages, and significant oenological milestones associated with specific grape cultivars throughout history, encompassing records of celebrated historical vintages (documented exceptional wine years), landmark production events (first classified growth, first appellation establishment, first international competition success), historical tasting notes and quality assessments from archived sources, documented sales records and auction prices reflecting historical valuation, and the development of cultivar-specific winemaking traditions and innovations, providing the qualitative evidence of viticultural excellence that contextualizes a cultivar's standing within the hierarchy of wine quality and cultural prestige.
Historical wine record documentation captures the qualitative evidence of cultivar significance that acreage statistics alone cannot convey — a cultivar may occupy modest vineyard area while producing wines of extraordinary cultural impact (Nebbiolo in Piedmont occupies approximately 5,500 ha globally yet produces Barolo and Barbaresco, among the world's most celebrated and expensive wines), while a cultivar of vast acreage may produce wines of limited historical distinction (Airén's 200,000+ ha generates primarily bulk distillation material). The archival sources for historical wine records are diverse and of varying reliability — pre-modern records include monastic cellar books (documenting vintage quality assessments from the 12th century onward in Burgundy), aristocratic estate accounts (recording vineyard management and wine production), and travelers' accounts (providing external quality assessments, as with Jefferson's Rhine journey notes of 1788). Modern sources include auction records (providing market-validated quality assessments through realized prices, with major houses Christie's and Acker Merrall & Condit maintaining archives extending to the early 19th century), critical scores and reviews (from the mid-20th century onward), and official appellation panel records documenting quality approval decisions. The concept of "landmark vintage" encodes the viticultural community's assessment of exceptional annual quality — the declaration of a great vintage (1947, 1961, 1982, 2005, 2010 in Bordeaux 1921, 1971, 2001, 2019 in Germany) represents a collective quality judgment that influences long-term pricing, cellaring decisions, and the cultivar's reputation trajectory. The historical wine record also documents oenological innovations associated with specific cultivars — the development of Tokaji Aszú production techniques in the 17th century, the méthode champenoise perfection attributed to Dom Pérignon (though historically oversimplified), and the modern development of whole-cluster Pinot Noir fermentation in Burgundy each represent cultivar-specific production milestones with lasting impact on global winemaking practice.
Weinherstellungshistorie
Weinproduktionsgeschichte
bedeutende Jahrgangsdokumentation
historische Weinaufzeichnung
notable vintage record
oenological milestone
wine achievement record
wine heritage documentation
wine production history
önologischer Meilenstein
Johnson H (1989) Vintage: The Story of Wine, Simon and Schuster, pp.: 1-480 Broadbent M (2003) Michael Broadbent's Vintage Wine, Harcourt, pp.: 1-580 Robinson J, Harding J (2015) The Oxford Companion to Wine, 4th Ed, Oxford University Press Lukacs P (2000) American Vintage: The Rise of American Wine, Houghton Mifflin, pp.: 1-30
Historical Wine Record
The name Malvasia constitutes one of the most extensive homonym complexes in grapevine nomenclature, applied to at least 18 genetically distinct cultivars confirmed by SSR profiling including Malvasia Bianca di Candia (VIVC 07209), Malvasia Bianca Lunga (VIVC 07210), Malvasia di Lipari (VIVC 07219), Malvasia Nera di Brindisi (VIVC 07227), Malvasia Nera di Lecce (VIVC 07228), and Malvasia Istriana (VIVC 07222) — the shared name derives from the medieval port of Monemvasia in the Peloponnese through which sweet wines were exported, but the cultivars bearing the name are unrelated and were independently associated with the Malvasia designation across different Mediterranean regions over centuries.
A cultivar designation where a single name is applied to two or more genetically distinct grapevine cultivars, creating nomenclatural ambiguity that requires contextual information — such as geographic origin, berry color, or molecular profile — to resolve which cultivar is intended by a given usage of the name.
Homonym represents the converse nomenclatural problem to synonymy: while synonymy creates multiple names for one genotype, homonymy creates one name for multiple genotypes. Homonymy is particularly prevalent in grapevine nomenclature due to the historical practice of naming cultivars after wine styles, geographic origins, or superficial morphological similarities rather than genetic identity — resulting in names like Malvasia, Moscato/Muscat, Trebbiano, and Lambrusco each being applied to multiple unrelated cultivars. The resolution of homonymy through molecular identification has been one of the most important contributions of SSR profiling to grapevine taxonomy. Subclasses distinguish true homonyms (identical spelling applied to distinct genotypes) from false homonyms (similar but non-identical names mistakenly treated as the same designation).
Homonym
ambiguous cultivar name
gleichlautende Sortenbezeichnung
mehrdeutiger Sortenname
polysemous cultivar designation
shared variety name
Crespan M and Milani N (2001) The Muscats: a molecular analysis of synonyms, homonyms and genetic relationships, Vitis 40: 79-86 Lacombe T et al. (2013) Large-scale parentage analysis in an extended set of grapevine cultivars, Theoretical and Applied Genetics 126: 401-414 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157
A horizontal cordon on Vitis vinifera cv. Chardonnay in Burgundy, extending 72cm from trunk positioned along trellis wire at 80cm height, trained horizontally parallel to ground surface, bearing 12 two-bud spurs distributed along upper surface at 12cm intervals, shoots from spurs growing vertically upward trained between catch wires in traditional Cordon de Royat configuration, representing classic horizontal cordon architecture employed in premium Burgundian Chardonnay production, with horizontal orientation providing stable framework for consistent canopy development, uniform fruit exposure, and traditional hand harvest operations maintaining fruit integrity for premium white wine production.
A cordon trained and maintained in horizontal orientation along trellis wires, characterized by perpendicular relationship to trunk (which is typically vertical), parallel alignment with soil surface and trellis wire, fruiting spurs positioned along upper surface facilitating upward shoot growth, and suitability for vertical shoot positioning and similar training systems requiring horizontal permanent framework supporting vertical canopy development. Horizontal cordons represent the most prevalent cordon configuration in modern viticulture worldwide, providing efficient platform for mechanized operations (over-row equipment for hedging, harvesting, spraying), consistent fruiting zone height facilitating uniform fruit development and harvest operations, and proven productivity and quality outcomes across diverse varieties, regions, and production objectives from bulk wine to ultra-premium applications.
Horizontal cordons dominate modern cordon viticulture due to numerous practical advantages including mechanization compatibility (horizontal permanent structure suits over-row equipment), consistent fruiting zone (uniform height facilitates uniform ripening and efficient harvest), simplified training and management (horizontal orientation is intuitive and stable), and proven performance across diverse conditions. Height selection for horizontal cordons varies by region, variety, and objectives, ranging from low cordons at 40-60cm (traditional European systems, hand harvest focus) to high cordons at 90-120cm (mechanized harvest systems, ergonomic considerations), with intermediate heights (70-90cm) representing common compromise between various factors. The horizontal orientation naturally positions spurs upward (on top of cordon) facilitating vertical shoot growth, though some systems deliberately position spurs laterally or at angles for specific canopy management objectives.
Drahtkordon
Querkordon
horizontale Kordon
lateral cordon
transverse cordon
wire cordon
Horizontal Cordon
Investigation of Horizontal_Growth_Disposition at Stellenbosch University characterized shoot architecture in cultivars recognized for spreading, horizontal growth habit. Reference cultivars 'Chenin Blanc', 'Colombard', 'Sémillon', and 'Muscadelle' were grown on multiple training systems with detailed angular measurements and positioning response assessment. Horizontal_Growth_Disposition cultivars showed consistent characteristics. 'Chenin Blanc' exhibited typical horizontal growth with mean shoot angle of 48° from vertical at 50 cm, increasing to 62° at 80 cm as shoot weight accumulated, with shoot tips often reaching true horizontal (90°) or slightly below. Gravitropic correction was minimal—tips deviated only 8° upward from mid-shoot trajectory compared to 25° in upright cultivars. 'Sémillon' showed similar Horizontal_Growth_Disposition (52° at 50 cm, 68° at 80 cm). Internode stiffness averaged 1.8 N·mm² (lower quartile). VSP positioning trials revealed that Horizontal_Growth_Disposition cultivars required 48-58 hours/ha/season positioning labor (3-4 passes), and showed 35% position loss (shoots falling from catch wires) compared to < 10% for upright types. Training system comparison demonstrated that 'Chenin Blanc' achieved superior canopy metrics on sprawl (leaf layer number 1.4, cluster exposure 52%) compared to VSP with intensive positioning (leaf layers 2.1, exposure 38%), suggesting that working with rather than against Horizontal_Growth_Disposition improves canopy function. Traditional pergola training in South African Chenin Blanc vineyards effectively utilizes the horizontal growth habit, achieving excellent fruit zone exposure beneath the horizontal canopy.
A Horizontal_Growth_Disposition is a Shoot_Orientation_Disposition that inheres in Vitis vinifera shoots and is realized through weak negative gravitropism producing shoots that grow at angles of 40-60° from vertical, approaching or reaching horizontal orientation, particularly in the mid and distal portions of the shoot as length and weight increase. This disposition represents the intermediate-trailing range of shoot orientation, characteristic of cultivars that lack the gravitropic strength to maintain vertical growth against shoot weight, resulting in shoots that arc outward and eventually grow nearly horizontally without positioning intervention. Horizontal-growing cultivars present challenges for VSP training systems, requiring intensive positioning labor to achieve vertical canopy geometry, and often showing poor position retention (shoots falling back toward horizontal after positioning). These cultivars may be better suited to training systems that accommodate or utilize horizontal shoot orientation, including sprawl, GDC (Geneva Double Curtain), Scott Henry, and traditional pergola systems common in their regions of origin. The mechanical basis of Horizontal_Growth_Disposition includes lower internode stiffness and weaker gravitropic response, allowing shoot weight to overcome the upward growth tendency early in development.
Horizontal_Growth_Disposition is_a Shoot_Orientation_Disposition representing spreading/horizontal growth (40-60° from vertical). Characterized by weak gravitropism, lower internode stiffness, and increasing angle with shoot length. Examples include 'Chenin Blanc', 'Colombard', 'Sémillon', 'Muscadelle', 'Müller-Thurgau'. Poor VSP compatibility with high positioning requirement and position loss. Better suited to sprawl, GDC, pergola systems. Position loss (shoots falling from catch wires) is characteristic.
Bogiger-Habitus
Horizontale-Wuchs-Disposition
Waagerechter-Triebwuchs
arching growth
horizontal shoot habit
spreading growth
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles, Adelaide Archer E, Strauss HC (1985) Effect of plant density on root distribution of three-year-old grafted 99 Richter grapevines, South African Journal of Enology and Viticulture 6:25-30 Reynolds AG et al. (1994) Influence of cluster exposure on fruit composition and wine quality of Seyval blanc, Vitis 33:137-143
Horizontal Growth Disposition
Hue quality transitions during Gewürztraminer berry development illustrate the progressive shift from green (CIELAB h° approximately 120°) through yellow-green (h° 100-110°) at véraison to the characteristic pink-copper hue (h° 40-60°) at maturity that distinguishes this aromatic cultivar the final copper-pink hue results from moderate anthocyanin accumulation combined with retention of yellow carotenoid pigments, creating the distinctive tawny appearance that historically linked Gewürztraminer to its presumed Traminer ancestry.
Hue_Quality represents the chromatic character that distinguishes different color types based on the dominant wavelength of light reflected or transmitted by grapevine tissues. Hue is the perceptual attribute that allows classification of colors into categories such as green, yellow, red, blue, and purple, independent of their intensity or lightness. In grapevine, hue variation results from the differential presence and chemical state of pigment molecules that selectively absorb and reflect specific wavelengths of visible light. Chlorophylls absorb red and blue light while reflecting green, producing the green hue of photosynthetic tissues. Carotenoids absorb blue and some green light, producing yellow to orange hues visible in senescing leaves and certain berry skins. Anthocyanins display pH-dependent hue shifts, appearing red at low pH (around 3.0), violet at intermediate pH, and blue at higher pH, with the specific hue also influenced by the hydroxylation and methoxylation pattern of the anthocyanin B-ring. This category subdivides into specific hue classes including Green_Hue_Quality, Yellow_Hue_Quality, Red_Hue_Quality, Blue_Hue_Quality, and Purple_Hue_Quality that capture the major color categories relevant to grapevine characterization. Hue assessment employs visual matching to color standards, measurement of hue angle (h° = arctan(b*/a*)) in CIELAB color space, or spectrophotometric characterization of absorbance maxima. Hue provides essential cultivar identification characters, particularly for berry skin color which represents a primary ampelographic descriptor.
Hue_Quality addresses the qualitative aspect of coloration (what type of color) as distinct from Pigmentation_Intensity_Quality (how much color). Hue is conceptually independent of intensity and lightness, though very low intensities make hue difficult to perceive. The child classes (Green_Hue_Quality, Yellow_Hue_Quality, etc.) represent discrete hue categories commonly used in ampelographic description, though hue actually varies continuously around the color circle. PATO_0000015 (hue) provides a potential upper-ontology reference. The hue angle in CIELAB space provides a continuous quantitative measure that can be mapped to categorical descriptions.
Farbkategorie
Farbtönungsqualität
chromatic character
chromatischer Charakter
color category
color type
spectral color quality
McGuire RG (1992) Reporting of Objective Color Measurements. HortScience 27(12): 1254-1255 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed, John Wiley and Sons pp.: 141-170
Hue Quality
The Huglin Index for the Alsace wine region (Colmar, 48.08°N) documents a value of 1,850-2,050 HI units (1991-2020 normal) — applying the daylength coefficient k=1.04 appropriate for the 48°N latitude, which increases the calculated heat accumulation by 4% compared to lower-latitude sites with the same temperatures, reflecting the 15-16 hour summer photoperiods at Alsatian latitude that provide more total daily radiation than the 13-14 hour photoperiods at 35°N despite similar peak temperatures. The Huglin Index classification places Alsace in the cool-temperate viticultural category (HI 1,800-2,100), suited to cultivars of intermediate heat requirement — Riesling (HI requirement approximately 1,600-1,800), Pinot Gris (1,700-1,900), Gewürztraminer (1,800-2,000), and Pinot Noir (1,700-1,900) — with the HI values explaining why Alsace can ripen these cultivars despite its northerly latitude (48°N versus 45-47°N for comparable-warmth sites in Burgundy and the Northern Rhône) through the photoperiod compensation captured by the daylength coefficient.
A growing degree-day component that records the Huglin Heliothermal Index (HI) — a modified heat accumulation index developed by Pierre Huglin (1978) that uses the mean of daily mean temperature and daily maximum temperature (capturing the biologically active daytime period more heavily than the standard GDD calculation), applies a base temperature of 10°C, sums over the April 1 to September 30 period (northern hemisphere), and incorporates a daylength adjustment coefficient (k, ranging from 1.02 to 1.06 for latitudes 40-50°N) that accounts for the longer photoperiods at higher latitudes that increase photosynthetic potential — providing a bioclimatic index that more accurately predicts cultivar maturity potential across latitudes than the standard Winkler GDD.
The Huglin_Index represents a significant refinement of the basic GDD concept through two innovations that improve the biological relevance of the heat accumulation calculation. First, the use of (Tmean + Tmax)/2 as the daily temperature input (rather than the standard (Tmax + Tmin)/2 used in GDD calculations) weights the daily contribution toward the warmer daytime hours when photosynthesis and most growth processes are active, reducing the influence of cool nighttime temperatures that contribute to the standard mean but during which minimal developmental progress occurs — this weighting produces higher index values than standard GDD for sites with wide diurnal temperature range and similar values for sites with narrow diurnal range, effectively discriminating between continental (high amplitude) and maritime (low amplitude) climates in a biologically meaningful way: second, the daylength coefficient k adjusts for the latitude-dependent photoperiod that determines the total daily radiation receipt and photosynthetic opportunity — at 50°N, the June photoperiod is approximately 16 hours compared to 14 hours at 35°N, providing 15% more potential photosynthetic time that partially compensates for the lower solar elevation angle and peak radiation intensity at higher latitudes. These two adjustments make the Huglin Index more accurate than the Winkler Index for cross-latitude comparison of viticultural potential — the Huglin Index correctly predicts that Riesling ripens successfully at 50°N in the Mosel (HI 1,600-1,800) despite a Winkler Index (1,000-1,200 GDD) that would classify the site as marginally too cool based on the Winkler framework alone.
Heliothermalindex
Huglin bioclimatic index
Huglin-Index
Huglin-Wärmesumme
daylength-adjusted heat summation
heliothermal index
Huglin P (1978) Nouveau mode d'évaluation des possibilités héliothermiques d'un milieu viticole, Comptes Rendus de l'Académie d'Agriculture de France 64: 1117-1126 Tonietto J and Carbonneau A (2004) A multicriteria climatic classification system for grape-growing regions worldwide, Agricultural and Forest Meteorology 124: 81-97 Jones GV et al. (2005) Climate change and global wine quality, Climatic Change 73: 319-343
Huglin Index
Humidity data for Sauternes documents autumn (October-November) morning relative humidity of 90-98% with fog persistence until 10:00-11:00, followed by afternoon drying to 55-65% RH — the diurnal humidity oscillation essential for Botrytis cinerea to alternate between nocturnal spore germination and daytime desiccation that produces noble rot rather than destructive grey rot on Sémillon berries. Canopy humidity monitoring in McLaren Vale Shiraz showed that early-season leaf removal reduced fruit zone RH from 88% to 72% at dawn and decreased leaf wetness duration from 6.5 to 3.2 hours per night, effectively suppressing powdery mildew (Erysiphe necator) development below the intervention threshold. VPD data for the Barossa Valley documents midday values of 3.5-5.0 kPa during January-February heat events, inducing stomatal closure in ungrafted Shiraz and limiting photosynthetic carbon gain during the critical ripening period.
Humidity data is a microclimate characterization entity that records atmospheric moisture parameters at a viticultural site and within the grapevine canopy, including relative humidity (RH, expressed as percentage), vapor pressure deficit (VPD, expressed in kilopascals), dew point temperature, leaf wetness duration, and diurnal humidity cycles, providing essential information for assessing grapevine transpiration demand, stomatal regulation, fungal pathogen infection risk, and the potential for noble rot (Botrytis cinerea) development under favorable conditions.
Humidity data is viticulturally significant through two principal pathways — transpiration regulation and disease epidemiology — that often create opposing management imperatives. High humidity reduces VPD and vine transpirational water loss (beneficial during drought stress) but simultaneously promotes fungal infection by extending leaf wetness duration and providing free water for spore germination. The distinction between relative humidity (temperature-dependent) and VPD (the true driving force for transpiration) is critical for physiological interpretation — identical RH values at different temperatures create vastly different transpiration demands. Leaf wetness duration (LWD), measured by impedance-grid or dielectric sensors deployed within the canopy, provides the most direct predictor of fungal infection risk, as most major grapevine pathogens (Botrytis cinerea, Plasmopara viticola, Erysiphe necator) require minimum free water periods for spore germination. Disease prediction models such as the Gubler-Thomas powdery mildew risk index and the Goidanich downy mildew model rely heavily on humidity-derived parameters. Under climate change, shifts in humidity patterns may prove as consequential for disease management as temperature changes, particularly where increasing temperatures reduce relative humidity but altered precipitation patterns increase canopy wetness events.
Dampfdruckdaten
Feuchtigkeitsdaten
Feuchtigkeitsregime
Luftfeuchtigkeit
Luftfeuchtigkeitsdaten
atmospheric moisture data
humidity record
moisture regime data
vapor pressure data
vineyard humidity data
Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 58-72 Gubler WD et al. (1999) A simple and rapid field assay for powdery mildew risk assessment. American Journal of Enology and Viticulture 50(3): 371-376 Dry PR, Coombe BG (2004) Viticulture Volume 1: Resources, 2nd Ed, Winetitles, pp.: 70-85 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1: The Microbiology of Wine and Vinifications, 2nd Ed, Wiley, pp.: 57-75
Humidity Data
Regent (Diana × Chambourcin cross) bears a Hybrid_Role as a registered German PIWI variety with complex interspecific ancestry, combining disease resistance from American species with wine quality from V. vinifera ancestors across multiple generations. Müller-Thurgau (historically claimed as Riesling × Silvaner, genetically confirmed as Riesling × Madeleine Royale) serves in a Hybrid_Role as an intraspecific V. vinifera hybrid, one of the most widely planted white varieties in Germany. SO4 rootstock (V. berlandieri × V. riparia) bears a Hybrid_Role as the world's most planted rootstock, its interspecific hybrid origin providing phylloxera resistance absent from either parent species alone.
A breeding role that inheres in a grapevine genotype by virtue of its origin through sexual hybridization between distinct parents, combining genetic material from two different individuals (and potentially different species or genera) to create a novel genotype with characteristics derived from both progenitors. Hybrid roles are distinguished by the taxonomic relationship between parents (interspecific between species, intraspecific within species), by the generation structure (F1 first filial generation, backcross from crossing with a parent), and by the complexity of ancestry (simple two-parent crosses versus complex multi-species pedigrees). Grapevine hybrids have played critical roles in viticulture history—interspecific hybrids provided phylloxera-resistant rootstocks saving European viticulture (1880s-1900s), French-American hybrids offered direct-producer alternatives to grafting (early 20th century), and contemporary PIWI (pilzwiderstandsfähig) varieties represent disease-resistant interspecific hybrids with improved wine quality. The hybrid designation in grapevine contexts often implies interspecific origin (crossing beyond V. vinifera), though technically any cross product is a hybrid. Hybrid vigor (heterosis) is commonly observed in grapevine crosses, with F1 progeny often exceeding parents for vegetative growth and stress tolerance.
Hybrid_Role is subclassed by taxonomic scope (Interspecific versus Intraspecific) and by generation structure (F1 versus Backcross), recognizing that hybrid significance varies with the genetic distance between parents and the position in breeding program progression. The term "hybrid" carries complex connotations in wine culture—historically pejorative when associated with "foxy" American hybrid flavors, increasingly positive as PIWI varieties demonstrate that interspecific hybrids can produce quality wines while reducing environmental impacts. Hybrid designation in regulatory contexts varies EU wine law historically restricted "quality wine" designations to V. vinifera, though recent reforms increasingly accommodate approved hybrid varieties. The heterozygosity of hybrids (combining alleles from distinct parents) contrasts with the relative uniformity within clonal V. vinifera varieties, with consequences for breeding behavior and genetic analysis.
Hybride
Hybridsorte
Kreuzung
Kreuzungsprodukt
cross
crossbred
hybrid
hybrid variety
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, in Reynolds A (ed) Grapevine Breeding Programs for the Wine Industry, Elsevier pp.: 3-22 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Reisch BI et al. (2012) Grape breeding in the United States, in Kole C (ed) Genetics and Breeding of Grapes, CRC Press pp.: 171-203
Hybrid Role
Hybrid vigor in Vitis is prominently displayed in rootstock breeding, where F1 hybrids between American species often exceed parental performance for root development and stress tolerance. The V. berlandieri × V. riparia cross producing rootstocks such as SO4 and 5BB Kober shows heterosis for root system size (20-40% larger than either parent) and phylloxera tolerance. V. berlandieri × V. rupestris hybrids (110 Richter, 140 Ruggeri) exhibit heterosis for drought tolerance and lime resistance exceeding parental species. Scion hybrid vigor is observed in some PIWI cultivars, with Regent showing 15-25% greater yield potential than V. vinifera parents while maintaining acceptable wine quality. Heterosis magnitude varies by trait, with vegetative vigor showing 10-40% superiority in favorable combinations while fruit quality traits may show intermediate or negative heterosis.
An interspecific hybridization disposition that is the inherited capacity of Vitis interspecific hybrids to exhibit heterosis (superior performance compared to parental species) for growth vigor, stress tolerance, or yield potential, resulting from heterozygosity, complementation of deleterious alleles, and epistatic interactions between divergent parental genomes.
Hybrid vigor provides the genetic basis for rootstock superiority, as most commercial rootstocks are interspecific hybrids exploiting heterosis for vigor and stress tolerance. The phenomenon results from multiple genetic mechanisms including dominance (masking of deleterious recessive alleles), overdominance (heterozygote advantage at individual loci), and epistasis (favorable interactions between loci from different species). Heterosis typically declines in subsequent generations (F2, backcross) as heterozygosity decreases, favoring F1 hybrids for rootstock use. Understanding hybrid vigor mechanisms informs breeding strategy for both rootstock and scion improvement.
Bastardwüchsigkeit
Heterosis
Hybridvitalität
heterosis
hybrid superiority
vigor enhancement trait
Ollat N et al. (2016) Grapevine roots and soil environment. In: Gerós H et al. (eds) Grapevine in a Changing Environment, Wiley pp.: 175-202 Cousins P (2005) Evolution, genetics, and breeding: viticultural applications of the origins of our rootstocks. In: Christensen LP, Smart DR (eds) Proceedings of the Soil Environment and Vine Mineral Nutrition Symposium, ASEV pp.: 1-7
Hybrid Vigor Disposition
Comprehensive characterization of Hydraulic_Architecture_Disposition at INRAE Bordeaux quantified hydraulic parameters across 20 cultivars using bench dehydration vulnerability curves, native embolism assessment, and xylem anatomical analysis. Stem segments were dehydrated to a range of water potentials, then measured for percentage loss of hydraulic conductivity (PLC) to construct vulnerability curves, with P50 (water potential at 50% PLC) as the key safety parameter. Cultivar variation in Hydraulic_Architecture_Disposition was substantial. 'Grenache' exhibited a conservative hydraulic architecture with relatively low maximum conductivity (Kmax = 3.8 kg m⁻¹ MPa⁻¹ s⁻¹) but high cavitation resistance (P50 = -2.8 MPa), reflecting narrow mean vessel diameter (145 μm) and thick inter-vessel pit membranes. 'Syrah' showed contrasting properties with high Kmax (6.2 kg m⁻¹ MPa⁻¹ s⁻¹) but lower safety (P50 = -1.9 MPa), associated with larger vessels (185 μm mean diameter). The hydraulic safety margin (difference between minimum operating water potential and P50) was 1.4 MPa for 'Grenache' (Ψmin = -1.4 MPa, P50 = -2.8 MPa) versus only 0.3 MPa for 'Syrah' (Ψmin = -1.6 MPa, P50 = -1.9 MPa), explaining 'Syrah's vulnerability to hydraulic failure during severe drought. These Hydraulic_Architecture_Disposition differences directly related to isohydric-anisohydric classification, with conservative 'Grenache' maintaining larger safety margins through stomatal closure while 'Syrah' operated near its hydraulic limits to maintain gas exchange.
A Hydraulic_Architecture_Disposition is a Drought_Tolerance_Disposition that inheres in the vascular system of a Vitis vinifera plant and is realized through the structural and functional properties of the water transport pathway that determine the efficiency of water movement from soil to leaves and the vulnerability of this pathway to drought-induced failure. This disposition represents the xylem-centered component of drought tolerance, encompassing vessel dimensions and connectivity, resistance to cavitation (embolism formation under tension), and capacity to repair embolized vessels following stress relief. Hydraulic architecture reflects the fundamental trade-off in xylem design between transport efficiency (maximized by large, frequent vessels) and safety (enhanced by small vessels with strong pit membranes resistant to air-seeding). Grapevine is among the most efficient water conductors in the plant kingdom due to large vessel diameters (often exceeding 200 μm), but this efficiency creates vulnerability to cavitation under drought stress when xylem tensions exceed air-seeding thresholds. Hydraulic_Architecture_Disposition varies among cultivars and rootstocks in vessel dimensions, cavitation resistance (P50, the pressure causing 50% loss of conductivity), and embolism repair capacity, collectively determining the hydraulic safety margins that allow continued function versus the efficiency that supports high productivity. Understanding Hydraulic_Architecture_Disposition informs the matching of cultivar hydraulic properties to site water availability and the interpretation of isohydric versus anisohydric stomatal behavior as strategies maintaining hydraulic safety margins.
Hydraulic_Architecture_Disposition is_a Drought_Tolerance_Disposition. Hydraulic architecture determines the efficiency-safety trade-off in xylem function. Child classes include Cavitation_Resistance_Disposition (P50, resistance to embolism), Embolism_Repair_Disposition (recovery capacity), and Vessel_Diameter_Disposition (anatomical basis of conductivity). Hydraulic safety margin (Ψmin - P50) determines vulnerability to drought-induced hydraulic failure. Hydraulic architecture underlies and helps explain isohydric versus anisohydric stomatal behavior strategies.
Hydraulische-Architektur-Disposition
Vaskulärer-Wassertransport
Xylem-Hydraulik
hydraulic conductance architecture
vascular water transport
xylem hydraulic properties
Tyree MT, Ewers FW (1991) The hydraulic architecture of trees and other woody plants, New Phytologist 119:345-360 Choat B et al. (2010) Measurement of vulnerability to water stress-induced cavitation in grapevine: a comparison of four techniques applied to a long-vesseled species, Plant Cell and Environment 33:1502-1512 Brodribb TJ, Cochard H (2009) Hydraulic failure defines the recovery and point of death in water-stressed conifers, Plant Physiology 149:575-584
Hydraulic Architecture Disposition
Hydraulic conductivity assessment comparing drought-tolerant and sensitive rootstocks reveals the hydraulic basis for stress tolerance differences, with tolerant rootstock showing whole-plant hydraulic conductance of 4.2 mmol/m²/s/MPa enabling midday leaf water potential of -1.1 MPa at transpiration rate of 4.5 mmol/m²/s, while sensitive rootstock with conductance of 2.8 mmol/m²/s/MPa reaches -1.6 MPa at similar transpiration, demonstrating how hydraulic conductivity determines the water potential cost of transpiration that affects stomatal behavior and drought vulnerability.
Hydraulic conductivity quality is a physiological quality that represents the capacity of grapevine tissues to conduct water under a given pressure gradient, characterizing the efficiency of water transport through the soil-plant-atmosphere continuum that determines how effectively roots, stems, and leaves can supply water to replace transpirational losses and maintain cell hydration under varying evaporative demand. Within the BFO framework, this quality addresses the water transport dimension of vine physiology, distinguishing the conductivity of water movement pathways from the driving forces (water potential gradients) that induce flow while recognizing that hydraulic conductivity establishes the relationship between water potential difference and flux that determines vine capacity to sustain transpiration without excessive water potential decline. The ontological significance lies in hydraulic conductivity determining the vulnerability of vines to water stress, where high conductivity enables rapid water transport maintaining leaf hydration under high transpiration demand, while low conductivity creates hydraulic bottlenecks where water supply cannot match demand, causing water potential decline, stomatal closure, and potential hydraulic dysfunction that limits productivity and risks permanent damage. The physical basis for hydraulic conductivity involves the resistance to water flow through various pathway components including root radial pathways (cell-to-cell movement into the xylem), axial xylem conduits (vessel elements in the transpiration stream), and the leaf mesophyll network, with conductivity determined by pathway anatomy (vessel diameter, pit membrane properties, cell membrane permeability) and physiological regulation (aquaporin gating, embolism development). The whole-plant integration of hydraulic conductivity recognizes that component conductivities operate in series and parallel arrangements, with whole-plant conductance (Kplant) determining the water potential gradient required to drive a given transpiration rate according to E = Kplant × (Ψsoil - Ψleaf), making hydraulic conductivity a key determinant of the leaf water potential achieved at any transpiration rate. The vulnerability to conductivity loss through embolism is a critical aspect of hydraulic conductivity, as drought-induced air entry into xylem vessels can catastrophically reduce conductivity, with the threshold water potential for embolism (P50) indicating drought tolerance. Understanding hydraulic conductivity quality enables assessment of vine water transport capacity, drought vulnerability evaluation, and appreciation of the hydraulic architecture that determines water relations performance under varying environmental conditions.
Hydraulic_Conductivity_Quality serves as the parent class for organ-specific conductivity qualities (Root, Stem). This quality determines the relationship between water potential gradient and flow that establishes vine water transport capacity. The class supports drought tolerance assessment and water relations interpretation through hydraulic conductivity characterization.
K-Wert
Wassertransportkapazität
hydraulic conductance
hydraulic permeability
hydraulische Leitfähigkeit
hydraulische Permeabilität
water transport capacity
Tyree MT, Ewers FW (1991) The hydraulic architecture of trees and other woody plants. New Phytologist 119(3): 345-360 Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell & Environment 26(8): 1393-1405
Hydraulic Conductivity Quality
Hydraulic recovery quality comparison following severe drought stress evaluated the capacity of different Vitis species and rootstocks to restore xylem function after cavitation-inducing water deficit. Stress to stem water potential of -1.8 MPa caused approximately 50% loss of hydraulic conductivity (PLC) across all genotypes through embolism formation. Following rewatering, Vitis rupestris demonstrated excellent hydraulic recovery quality with conductivity returning to 90% of pre-stress levels within 72 hours through efficient vessel refilling, evidenced by the disappearance of acoustic emissions indicating active bubble dissolution and positive xylem pressure measured by root pressure probes. Vitis vinifera cv. Chardonnay showed moderate hydraulic recovery quality with only 60% conductivity restoration at 72 hours and full recovery requiring 2 weeks, attributed to less efficient refilling mechanisms requiring supplementation by new xylem development visible in cross-sections. Vitis vinifera cv. Cabernet Sauvignon exhibited poor hydraulic recovery quality with only 45% conductivity restoration at 72 hours and persistent impairment of 20% even after 4 weeks, suggesting limited refilling capacity and reliance on new tissue development for recovery that proceeded slowly in this cultivar. The hydraulic recovery quality differential established Vitis rupestris-derived rootstocks as superior choices for sites experiencing periodic severe drought where rapid hydraulic recovery between stress events is essential for maintained vine health.
Hydraulic recovery quality is a drought recovery quality that inheres in a grapevine organism and represents the capacity to restore normal hydraulic conductivity in the xylem transport pathway following drought-induced embolism formation, through mechanisms including the refilling of gas-blocked vessels, the development of new functional xylem through cambial activity, and the rerouting of water transport through alternative pathways that bypass irreparably damaged conduits. This quality captures the repair dimension of drought recovery addressing the structural damage to the water transport system that may occur under severe stress, recognizing that drought-induced cavitation and embolism can create lasting hydraulic impairment that persists beyond tissue rehydration if repair mechanisms are insufficient to restore pathway function. The ontological significance of hydraulic recovery quality lies in its role determining whether drought episodes cause lasting damage to vine hydraulic function that accumulates across repeated stress events, with inadequate hydraulic recovery potentially creating progressive decline in water transport capacity that compromises long-term vine health and productivity. The biological basis of hydraulic recovery encompasses multiple repair mechanisms operating on different timescales, including the rapid refilling of embolized vessels through positive pressure generated by root activity or localized metabolic processes, the intermediate-term development of new xylem tissue through cambial division producing fresh conduits that bypass damaged vessels, and the long-term remodeling of hydraulic architecture through differential vessel development in subsequent growing seasons. The capacity for vessel refilling varies among genotypes, with some cultivars and rootstocks showing efficient refilling under favorable conditions while others require new xylem development to restore hydraulic function, creating substantial differences in the timescale of hydraulic recovery following severe stress. The relationship between stress severity and hydraulic recovery reflects threshold phenomena wherein moderate stress causing limited embolism may be rapidly refilled, while severe stress causing extensive cavitation may overwhelm refilling capacity requiring the slower process of new tissue development for recovery.
Hydraulic_Recovery_Quality is_a Drought_Recovery_Quality, representing the restoration of xylem hydraulic function following drought-induced embolism. This class addresses the structural repair dimension of recovery distinct from physiological restoration. The percentage of conductivity restored and the time course of restoration provide quantitative characterization. Vessel refilling versus new xylem development represent different recovery mechanisms operating on different timescales. Distinguished from Rehydration_Rate_Quality which addresses tissue water status rather than transport pathway function. Critical for understanding cumulative drought damage potential and rootstock selection for variable water environments.
Embolie-Reparatur
Leitfähigkeits-Wiederherstellung
Xylem-Erholung
hydraulic conductivity restoration
vascular recovery
Hochberg U et al. (2017) Stomatal closure, basal leaf embolism, and shedding protect the hydraulic integrity of grape stems. Plant Physiology 174:764-775 Lovisolo C et al. (2010) Drought-induced changes in development and function of grapevine (Vitis spp.) organs and in their hydraulic and non-hydraulic interactions at the whole-plant level: a physiological and molecular update. Functional Plant Biology 37:98-116 Brodersen CR et al. (2010) The dynamics of embolism repair in xylem: in vivo visualizations using high-resolution computed tomography. Plant Physiology 154:1088-1095
Hydraulic Recovery Quality
Hydraulic safety margin assessment across Mediterranean wine regions reveals the regional variation in hydraulic risk, with Atlantic-influenced areas showing safety margins of 0.8 MPa (Ψmin = -1.2 MPa, P50 = -2.0 MPa, substantial buffer), continental interior regions with margins of 0.3 MPa (Ψmin = -1.8 MPa approaching P50 = -2.1 MPa, narrow buffer), and the most drought-stressed sites showing negative margins during extreme years (Ψmin = -2.4 MPa exceeding P50, indicating embolism occurrence), demonstrating how safety margin mapping enables identification of regions where hydraulic failure risk requires management attention or cultivar adaptation.
Hydraulic safety margin quality is an embolism recovery capacity quality that represents the difference between the minimum xylem water potential experienced during normal operation and the water potential causing significant hydraulic dysfunction, characterizing the buffer zone between functional water transport and embolism-induced failure that determines how close vines operate to their hydraulic limits under typical conditions. Within the BFO framework, this quality specifically addresses the operational buffer dimension of hydraulic safety, distinguishing the margin between function and failure from the absolute vulnerability threshold while recognizing that safety margin indicates the drought intensity required to cause hydraulic damage and thus the risk level under current or projected growing conditions. The ontological significance lies in hydraulic safety margin determining the practical drought tolerance of vines under field conditions, where large safety margins indicate operation well below dysfunction thresholds with substantial buffer against drought intensification, while narrow margins indicate operation near hydraulic limits where modest additional stress could trigger embolism and conductivity loss. The calculation of hydraulic safety margin typically uses the difference between minimum seasonal stem water potential (Ψmin) and P50 as the safety margin = P50 - Ψmin, with positive values indicating operation above the 50% embolism threshold and negative values indicating that embolism has already occurred, though safety margins to other thresholds (P12, P88) may be more appropriate for specific questions. The global patterns in hydraulic safety margins show surprising convergence, with many woody species including grapevine operating with relatively narrow margins (0.2-1.0 MPa above P50), suggesting that evolution has optimized the trade-off between maximizing water use (narrow margins) and ensuring survival (wide margins), though whether this represents risky or optimal behavior remains debated. The climate change implications of safety margin assessment are significant, as projected increases in drought severity may cause species operating with narrow margins to exceed their hydraulic limits with increasing frequency, with grapevine safety margins in traditional wine regions becoming a concern under climate change scenarios. Understanding hydraulic safety margin quality enables risk assessment for hydraulic failure under current and projected conditions, evaluation of cultivar suitability for drought-prone environments, and informed decisions about irrigation strategies that maintain appropriate safety margins.
Hydraulic_Safety_Margin_Quality addresses the buffer between operating water potential and dysfunction threshold. This quality indicates practical drought tolerance and hydraulic failure risk. The class supports risk assessment and climate adaptation planning through safety margin characterization.
Betriebsmarge
Sicherheitspuffer
Trockenheitspuffer
drought buffer
hydraulic buffer
hydraulische Sicherheitsmarge
operating margin
safety margin
Choat B et al. (2012) Global convergence in the vulnerability of forests to drought. Nature 491(7426): 752-755 Charrier G et al. (2018) Drought will not leave your glass empty: low risk of hydraulic failure revealed by long-term drought observations in world's top wine regions. Science Advances 4(1): eaao6969
Hydraulic Safety Margin Quality
Hydrolyzable tannin quality management in first-growth Bordeaux employs careful barrel selection and aging protocols, with new French oak contributing 100-200 mg/L ellagitannins during 18-24 months barrel maturation the Bordeaux practice of using 50-100% new oak for Grand Vin production generates substantial hydrolyzable tannin extraction that integrates with grape-derived condensed tannins to produce the multi-layered, complex tannin structure characteristic of age-worthy classified growth wines, while second and third use barrels with depleted ellagitannin content serve for earlier-drinking wines requiring less structural tannin contribution.
Hydrolyzable_Tannin_Quality represents the chemical quality characterizing the concentration and composition of gallotannins and ellagitannins in wine, capturing the oak-derived polymeric phenolics that contribute to structure, complexity, and aging chemistry in barrel-matured wines. Hydrolyzable tannins are distinguished from condensed tannins by their chemical structure and origin: gallotannins consist of gallic acid esterified to glucose cores, while ellagitannins contain hexahydroxydiphenic acid (HHDP) units that release ellagic acid upon hydrolysis. These compounds are absent from grapes but are extracted from oak cooperage during barrel aging, with the concentration and composition depending on oak species, barrel toast level, barrel age, and aging duration. French oak (Quercus petraea and Q. robur) and American oak (Q. alba) differ substantially in ellagitannin content and profile, contributing to the stylistic differences between wines aged in different oak types. The primary oak ellagitannins—castalagin, vescalagin, grandinin, and roburin—undergo progressive hydrolysis and oxidation in wine, releasing ellagic acid while participating in complex reactions with wine components. Hydrolyzable tannins contribute to wine structure differently from condensed tannins, adding mid-palate body without the grippy finish associated with seed-derived proanthocyanidins. The antioxidant activity of ellagitannins provides protective effects during barrel aging. Hydrolyzable tannin concentration in barrel-aged wines typically ranges from 20-150 mg/L for moderately oaked wines to 200-800 mg/L for heavily oaked styles, declining during extended bottle aging as hydrolysis and precipitation reduce soluble concentrations.
Hydrolyzable_Tannin_Quality addresses the oak-extracted tannin fraction absent from grapes but significant in barrel-aged wines. The distinction from condensed tannins (proanthocyanidins) is fundamental hydrolyzable tannins have entirely different chemical structures and sensory properties. Oak species differences in ellagitannin content (French oak typically higher than American) contribute to winemaking stylistic choices. Analysis employs HPLC with UV or mass spectrometric detection, with individual ellagitannins quantified against reference standards. The term hydrolyzable reflects the susceptibility of the ester bonds to acid or base cleavage, distinguishing these compounds from the C-C bonded condensed tannins.
Eichentanningehalt
Ellagitanninqualität
Gallotanninkonzentration
Hydrolysierbares-Tannin-Gehalt
barrel-derived tannin quality
ellagitannin quality
gallotannin concentration
oak tannin content
Puech JL et al. (1999) Ellagitannins from Wood of Oak Species. Journal of the Science of Food and Agriculture 79(12): 1625-1627 Michel J et al. (2011) Flavanols and Ellagitannins in Wine: Evolution during Ageing and Contribution to Wine Taste. In: Ebeler SE et al. (eds) Progress in Authentication of Food and Wine, ACS Symposium Series 1081 pp.: 271-291
Hydrolyzable Tannin Quality
Hydroxycinnamic acid quality management in Sauvignon Blanc juice determines browning trajectory during oxidative handling, with cultivar-typical concentrations of 180-280 mg/L caftaric acid creating substantial polyphenol oxidase substrate that can generate brown quinones within minutes of crushing protective handling employing inert gas blanketing, low temperatures (5-10°C), and ascorbic acid addition (50-100 mg/L) preserves hydroxycinnamic acids in reduced form, maintaining the pale green-gold color associated with premium Sauvignon Blanc while preventing the brown-bronze hues of oxidized must.
Hydroxycinnamic_Acid_Quality encompasses the chemical qualities characterizing the concentration and composition of hydroxycinnamic acid derivatives in grape berry tissue, capturing the C6-C3 phenylpropanoid compounds that dominate the phenolic profile of grape juice and white wines. Hydroxycinnamic acids in grapes are present primarily as tartaric acid esters rather than free acids, with caftaric acid (caffeoyltartaric acid) and coutaric acid (p-coumaroyltartaric acid) as the dominant forms, accompanied by smaller amounts of fertaric acid (feruloyltartaric acid). These compounds accumulate in the flesh and skin vacuoles of grape berries, making them readily extractable during crushing and pressing without requiring extended maceration. Total hydroxycinnamic acid concentration in grape juice typically ranges from 100-300 mg/L, with substantial variation among cultivars and growing conditions. The hydroxycinnamic acids serve as primary substrates for polyphenol oxidase (PPO) and laccase enzymes, making their concentration and oxidation state critical determinants of juice and wine browning susceptibility. Enzymatic oxidation of caftaric acid produces the quinone form that can polymerize or react with other compounds, generating brown pigments and consuming glutathione that would otherwise protect other phenolics. Beyond browning chemistry, hydroxycinnamic acids serve as precursors for volatile phenols produced by Brettanomyces and other microorganisms coutaric acid yields p-vinylphenol and p-ethylphenol that contribute medicinal or barnyard aromas at elevated concentrations. The ratio of caftaric to coutaric acid varies among cultivars and provides fingerprinting information, while total hydroxycinnamic acid content indicates browning potential and volatile phenol precursor availability.
Hydroxycinnamic_Acid_Quality serves as the parent class for specific hydroxycinnamic acid derivatives including Caftaric_Acid_Content_Quality and Coutaric_Acid_Content_Quality. The tartaric acid esterification distinguishes grape hydroxycinnamic acids from those in other fruits where free acids or glucose esters predominate. The dual role of hydroxycinnamic acids as browning substrates and volatile phenol precursors creates quality implications in both white wine production (browning control) and red wine production (Brett risk management). Analysis employs HPLC with UV detection at 320 nm, with individual esters quantified against authentic standards or expressed as caffeic acid equivalents.
HCA content
Hydroxyzimtsäuregehalt
Phenylpropansäurequalität
Tartratester-Phenolkonzentration
Zimtsäurederivat-Qualität
cinnamic acid derivative quality
phenylpropanoid acid quality
tartrate ester phenolic concentration
Singleton VL et al. (1978) Caftaric Acid Disappearance and Conversion to Products of Enzymatic Oxidation in Grape Must and Wine. American Journal of Enology and Viticulture 29(2): 107-117 Cheynier V et al. (1989) Characterization of 2-S-Glutathionyl Caftaric Acid and Its Hydrolysis in Relation to Grape Wines. Journal of Agricultural and Food Chemistry 37(4): 1069-1071
Hydroxycinnamic Acid Quality
Detailed characterization of Hypersensitive_Response_Disposition at the University of Udine compared HR kinetics and pathogen containment across grapevine genotypes with and without the Rpv3 downy mildew resistance locus. Leaf discs were inoculated with P. viticola sporangia and assessed at 12-hour intervals for HR markers including trypan blue staining (cell death), DAB staining (H₂O₂ accumulation), and autofluorescence (phenolic accumulation at death sites). Resistant 'Bianca' (Rpv3-positive) exhibited strong Hypersensitive_Response_Disposition with H₂O₂ accumulation detectable at infection sites within 12 hours post-inoculation (hpi), cell death visible by 24 hpi, and complete HR lesion formation by 48 hpi. HR lesions were tightly confined (mean diameter 0.3 mm), with pathogen sporulation completely suppressed. Susceptible 'Chasselas' (Rpv3-negative) showed no HR, with diffuse pathogen colonization and abundant sporulation by 96 hpi covering 60% of disc area. The partial resistance cultivar 'Solaris' (Rpv3/Rpv10 combination) displayed enhanced Hypersensitive_Response_Disposition with faster HR initiation (H₂O₂ at 8 hpi) and smaller lesions (0.15 mm), correlating with stronger resistance. Importantly, HR-negative 'Chasselas' inoculated with the necrotroph Botrytis cinerea showed equivalent infection to HR-positive genotypes, confirming that Hypersensitive_Response_Disposition provides specific benefit against biotrophs rather than broad-spectrum resistance.
A Hypersensitive_Response_Disposition is an Induced_Defense_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to execute rapid, localized programmed cell death at sites of pathogen recognition, sacrificing infected and immediately adjacent cells to create a hostile environment that limits pathogen growth, reproduction, and spread to surrounding healthy tissue. This disposition represents one of the most effective plant defense mechanisms against biotrophic pathogens that require living host cells for nutrition and reproduction, effectively starving the pathogen by eliminating its food source. The hypersensitive response (HR) is typically triggered by recognition of pathogen effector proteins by plant resistance (R) proteins, initiating a signaling cascade involving reactive oxygen species (ROS) burst, nitric oxide production, calcium influx, MAPK activation, and ultimately activation of cell death programs. In grapevine, HR is observed in resistant responses to multiple pathogens including Plasmopara viticola (in Rpv-mediated resistance), Erysiphe necator (in Run/Ren-mediated resistance), and phylloxera (in root and foliar resistance). The speed, intensity, and spatial containment of HR vary among genotypes, with effective resistance requiring HR rapid enough to confine pathogens before significant colonization occurs and spatially restricted enough to minimize self-damage. Hypersensitive_Response_Disposition is particularly relevant for resistance to obligate biotrophic pathogens but may be ineffective or counterproductive against necrotrophs like Botrytis cinerea that thrive on dead tissue.
Hypersensitive_Response_Disposition is_a Induced_Defense_Disposition. HR is typically associated with effector-triggered immunity (ETI) mediated by R gene recognition of pathogen effectors. In grapevine, HR is central to Rpv-mediated downy mildew resistance and Run/Ren-mediated powdery mildew resistance. HR effectiveness depends on speed and spatial containment—too slow or too extensive HR may fail to provide protection. HR is ineffective against necrotrophs like Botrytis that benefit from host cell death, representing a fundamental trade-off in defense strategy.
HR capacity
HR-Kapazität
Hypersensitive-Reaktions-Disposition
Programmierter-Zelltod-Abwehr
localized cell death response
programmed cell death defense
Mur LAJ et al. (2008) The hypersensitive response the centenary is upon us but how much do we know?, Journal of Experimental Botany 59:501-520 Künstler A et al. (2016) Staying alive—is cell death dispensable for plant disease resistance during the hypersensitive response?, Physiological and Molecular Plant Pathology 93:75-84 Allegre M et al. (2009) Stomatal deregulation in Plasmopara viticola-infected grapevine leaves, New Phytologist 173:832-840
Hypersensitive Response Disposition
Hypersensitive response quality evaluation in the powdery mildew-resistant breeding line Vitis vinifera × V. rotundifolia BC4 carrying the Run1 resistance locus compared HR activation kinetics and effectiveness against Erysiphe necator to the susceptible V. vinifera parent. Microscopic examination of inoculated leaf tissue at 24-hour intervals revealed hypersensitive response quality differences in timing and extent of cell death responses. The Run1-carrying line showed rapid hypersensitive response quality with single epidermal cells undergoing HR visible as autofluorescent collapsed cells by 18-24 hours post-inoculation, precisely localized to cells contacted by appressorial penetration attempts and effectively halting greater than 95% of infections at the single-cell stage. The susceptible V. vinifera showed minimal hypersensitive response quality with no localized cell death at early time points, allowing E. necator to establish haustoria in living epidermal cells by 24 hours, with expanding colonies visible by 48-72 hours and abundant sporulation by 7 days. Quantification of hypersensitive response quality through trypan blue staining for dead cells showed the resistant line with 4.2 dead cells per attempted penetration site (rapid HR limiting spread) versus 0.3 dead cells in the susceptible line (no effective HR). The hypersensitive response quality assessment established that the Run1-mediated resistance functions through rapid HR activation upon pathogen recognition, with the speed of the response critical for containing the rapidly-growing powdery mildew pathogen before colony establishment.
Hypersensitive response quality is a defense response quality that inheres in a grapevine organism and represents the capacity to undergo rapid, localized programmed cell death at pathogen infection sites that sacrifices infected cells to prevent pathogen spread into healthy tissue, with the hypersensitive response (HR) representing one of the most effective defense mechanisms against biotrophic pathogens that require living host cells for nutrition and are killed or contained by the death of their host cells. This quality captures the controlled self-destruction mechanism that limits pathogen colonization, recognizing that while cell death might seem counterproductive, the rapid death of a few cells at the infection site is far less damaging than allowing pathogen spread throughout the plant, making HR an effective defense particularly against obligate biotrophs including powdery mildew and downy mildew. The ontological significance of hypersensitive response quality lies in its role as a hallmark of effective gene-for-gene resistance where pathogen recognition by plant resistance proteins triggers the HR cascade, with strong HR capacity associated with the race-specific resistance conferred by major resistance genes and providing a clear phenotypic marker for resistance gene function. The biological basis of hypersensitive response encompasses the recognition of pathogen effector molecules by plant nucleotide-binding leucine-rich repeat (NLR) receptor proteins, the activation of signaling cascades involving calcium flux, reactive oxygen species burst, and nitric oxide production, and the execution of programmed cell death through mechanisms distinct from developmental cell death but sharing some molecular components with animal apoptosis. The visual appearance of HR in grapevine includes the small necrotic flecks or spots at attempted infection sites where cells have died to contain the pathogen, contrasting with the expanding lesions and sporulation seen when HR fails or is absent and the pathogen successfully colonizes living tissue. The effectiveness of HR depends on the speed of activation, with rapid HR containing pathogens before significant growth occurs while delayed HR may allow substantial colonization before cell death finally halts spread, creating necrotic lesions larger than the minimal flecks of rapid HR.
Hypersensitive_Response_Quality is_a Defense_Response_Quality, representing the rapid programmed cell death response that contains pathogens at infection sites. This class addresses the "suicide" defense mechanism particularly effective against biotrophic pathogens. NLR receptor recognition of pathogen effectors triggers HR signaling. Small necrotic flecks indicate effective rapid HR larger lesions suggest delayed or weak HR. Run1 (powdery mildew) and Rpv genes (downy mildew) function through HR activation. The kinetics of HR activation critically determine effectiveness against fast-growing pathogens.
HR response
HR-Antwort
Hypersensitive Reaktion
hypersensitive cell death
localized cell death
lokalisierter Zelltod
programmierter Zelltod-Abwehr
Jones JDG, Dangl JL (2006) The plant immune system. Nature 444:323-329 Feechan A et al. (2011) Genetic dissection of a TIR-NB-LRR locus from the wild North American grapevine species Muscadinia rotundifolia identifies paralogous genes conferring resistance to major fungal and oomycete pathogens in cultivated grapevine. The Plant Journal 65:939-953 Dry IB et al. (2010) Molecular strategies to enhance the genetic resistance of grapevines to powdery mildew. Australian Journal of Grape and Wine Research 16:94-105
Hypersensitive Response Quality
Cabernet Sauvignon hypodermal layer at maturity comprises 2-3 cell layers (total 42-55 μm thickness) immediately beneath 26-32 μm epidermis, with hypodermal cells 18-26 μm diameter, walls 1.8-2.6 μm thick, containing intense anthocyanin accumulation 3.2-4.5 mg/g fresh weight (predominantly malvidin-3-glucoside 52-58% of total) creating deep purple-black color concentrated in hypodermal cell vacuoles, with hypodermis representing 3-4% of pericarp volume but containing >90% of total berry anthocyanins demonstrating extreme spatial concentration of color pigments in sub-epidermal layers. Syrah hypodermal layer showing very high anthocyanin characteristics= 2-3 cell layers, 45-58 μm thickness, containing 4.2-5.5 mg/g anthocyanins (among highest in wine cultivars), creating very dark purple-black berry appearance, with hypodermal anthocyanin extraction during 18-25 day maceration providing 15-22 OD units color intensity at 520 nm in finished wines (very high for red wines) demonstrating direct relationship between hypodermal anthocyanin content and wine color potential. Pinot Noir hypodermal layer showing moderate pigmentation= 1-2 cell layers (thinner than Cabernet), 22-38 μm thickness, containing 2.2-3.2 mg/g anthocyanins with higher proportion of delphinidin and cyanidin 3-glucosides (35-48% combined) creating ruby-red hue versus purple-black, demonstrating cultivar variation in hypodermal layer number, anthocyanin concentration, and anthocyanin composition affecting berry color appearance and wine color characteristics. Chardonnay white cultivar hypodermal layer showing non-anthocyanin pigmentation= 1-2 cell layers, 24-42 μm thickness, lacking anthocyanins but containing carotenoids (0.25-0.5 mg/g β-carotene, lutein) and flavonols (0.3-0.8 mg/g quercetin glycosides) providing golden-yellow color component, with hypodermal carotenoids/flavonols combining with epidermal pigmentation creating characteristic Chardonnay berry appearance without anthocyanins. Merlot hypodermal layer anthocyanin development time-course= pre-veraison (BBCH 79, week 9) hypodermal cells containing <0.05 mg/g anthocyanins, green chlorophyll-containing veraison initiation (BBCH 81, week 10) showing VvMYBA1 gene upregulation specifically in hypodermal cells triggering anthocyanin synthesis beginning, 0.3-0.6 mg/g accumulated mid-ripening (BBCH 85, week 14) with 2.2-2.8 mg/g anthocyanins maturity (BBCH 89, week 18) with 3.2-3.8 mg/g maximum accumulation, demonstrating progressive hypodermal anthocyanin synthesis and accumulation during 8-9 week post-veraison ripening period creating final pigmented hypodermal layer determining berry color intensity.
A hypodermal layer is the sub-epidermal tissue comprising 1-3 cell layers immediately beneath the epidermis, typically 20-60 μm total thickness at maturity, characterized by small cells (15-30 μm diameter) with moderately thick walls (1.5-3 μm), serving as transition zone between outer protective epidermis and inner fleshy mesocarp, and functioning as primary site of anthocyanin accumulation in red/black cultivars determining berry color and wine color potential, as well as containing elevated tannins, flavonols, and other phenolic compounds contributing to berry and wine quality. Hypodermal layer anatomical characteristics= (1) position immediately beneath single-layer epidermis creating second cellular layer(s) from berry surface (2) typically 1-3 cell layers thick with layer number varying by cultivar (Pinot Noir often 1-2 layers, Cabernet Sauvignon typically 2-3 layers, some thick-skinned cultivars 3-4 layers) (3) total thickness 20-60 μm depending on cell layer number and individual cell size (4) cells small 15-30 μm diameter, substantially smaller than middle pericarp cells (80-180 μm) but larger than epidermal cells (radial dimension 15-35 μm) (5) cells roughly isodiametric to slightly elongated tangentially (6) cell walls moderately thick 1.5-3 μm, intermediate between very thick epidermal outer walls (3-6 μm) and thin middle pericarp walls (0.4-0.8 μm) (7) cells densely packed with reduced intercellular spaces compared to middle pericarp. Hypodermal layer compositional characteristics in red/black cultivars= (1) anthocyanin accumulation in cell vacuoles creating intense pigmentation with anthocyanin concentration 2.5-6+ mg/g fresh weight in hypodermal cells (versus <0.1 mg/g in middle pericarp and often <1 mg/g in epidermal cells), making hypodermis the primary site of berry color pigments (2) anthocyanin composition including malvidin-3-glucoside (typically 40-60% of total anthocyanins), other monoglucosides (delphinidin, cyanidin, petunidin, peonidin 3-glucosides), and acylated derivatives (acetyl and p-coumaroyl esters) (3) tannins at elevated levels typically 6-10 mg/g dry weight (higher than middle pericarp 0.3-1 mg/g but sometimes lower than outermost epidermal/outer mesocarp regions 8-15 mg/g) (4) flavonols present at moderate levels 0.5-1.5 mg/g providing UV protection and color copigmentation. Hypodermal layer compositional characteristics in white cultivars= (1) absence or near-absence of anthocyanins (<0.05 mg/g) (2) presence of carotenoids (β-carotene, lutein 0.2-0.5 mg/g) and flavonols (0.3-0.8 mg/g) contributing to golden-yellow color (3) moderate tannin levels 3-6 mg/g (elevated relative to middle pericarp though lower than red cultivar hypodermis). Hypodermal layer development and function= (1) cell differentiation during early berry development establishing hypodermal identity and position (2) anthocyanin synthesis initiation at veraison (BBCH 81) in red cultivars through upregulation of biosynthetic genes (VvMYBA1, VvUFGT) specifically in hypodermal cells creating spatial specificity of pigmentation (3) progressive anthocyanin accumulation during ripening (BBCH 81-89) reaching maximum at maturity (4) structural support providing transition zone between protective outer layers and soft flesh (5) phenolic reservoir providing majority of wine anthocyanins (color) and significant wine tannins (structure) during red wine maceration. Hypodermal layer in wine production= extraction of hypodermal anthocyanins during red wine maceration providing essentially all wine color as flesh contains minimal anthocyanins extraction kinetics showing anthocyanins extracting rapidly (60-80% extracted within 3-5 days maceration) while tannins extract more slowly extraction influenced by maceration temperature, alcohol concentration, and skin cell wall breakdown final wine color intensity directly related to hypodermal anthocyanin content and extraction efficiency.
Hypodermal_Layer represents the sub-epidermal cell layers and is the primary site of anthocyanin accumulation in red/black cultivars, distinguished from Epidermal_Layer above it and Outer_Mesocarp layers below it by position, cell characteristics, and compositional properties. Sub-classification into Sub_Epidermal (emphasizing position) and Supporting (emphasizing structural function) hypodermal layer reflects different aspects. Understanding hypodermal layer structure and composition is fundamental for viticulture and enology as this tissue determines berry color, provides major wine color and tannin contributions, and is the primary target for managing phenolic development through cultural practices.
Hypodermis-Schicht
Pigmentschicht
pigment layer
sub-epidermal layer
subepidermale Schicht
Cadot, Y. et al. (2006). Tannin composition and localization in grape skins. Journal of Agricultural and Food Chemistry 54=2973-2987 Castellarin, S.D. et al. (2007). Transcriptional regulation of anthocyanin biosynthesis. Plant Physiology 143=1347-1363 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 180-185.
Hypodermal Layer
IBMP concentration at harvest ranges from <1 ng/L in low-MP cultivars such as Pinot Noir and Merlot to 15-40 ng/L in high-MP types such as Cabernet Sauvignon (15-35 ng/L), Cabernet Franc (20-45 ng/L), Sauvignon Blanc (10-30 ng/L), and Carménère (30-80 ng/L). The sensory threshold of 1-2 ng/L in red wine and 8-15 ng/L in white wine means IBMP is supra-threshold in many wines from high-MP cultivars. IBMP biosynthesis involves condensation of leucine-derived isobutyl amine with glyoxylic acid followed by O-methylation by VvOMT3, with gene expression and IBMP accumulation peaking 2-4 weeks pre-veraison. Post-veraison decline of 90-98% through UV photodegradation (half-life 10-20 days in sun-exposed fruit) and metabolic degradation reduces IBMP to harvest levels. Shaded clusters retain 2-5× higher IBMP than exposed clusters at harvest.
A methoxypyrazine accumulation disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate 3-isobutyl-2-methoxypyrazine (IBMP) as the predominant methoxypyrazine in grape berries, with genotype-specific concentrations determining the intensity of characteristic green bell pepper aroma and representing the primary contributor to herbaceous character in Bordeaux varieties.
IBMP is the signature compound defining the green bell pepper note of Bordeaux varieties and represents the most thoroughly studied grape methoxypyrazine. The compound shows remarkable stability during winemaking and aging, with wine IBMP concentrations closely reflecting grape levels. Viticultural management (leaf removal, cluster exposure, delayed harvest, low vigor) provides the primary tools for IBMP reduction, as no winemaking intervention effectively removes IBMP from wine. Cool climate and vigorous vine conditions favor higher IBMP retention through reduced photodegradation and advanced harvest.
IBMP synthesis capacity
IBMP-Akkumulation
Isobutyl-Methoxypyrazin
Paprika-Aroma
bell pepper aroma trait
isobutyl-methoxypyrazine disposition
Dunlevy JD et al. (2013) The methyltransferase VvOMT3 is responsible for the formation of 3-isobutyl-2-methoxypyrazine (IBMP), a key flavour compound in grapevine. New Phytologist 197:1177-1186 Koch A et al. (2010) Fruit ripening in Vitis vinifera: light intensity before and not during ripening determines the concentration of 2-methoxy-3-isobutylpyrazine in Cabernet Sauvignon berries. Physiologia Plantarum 140:74-84
IBMP Accumulation Disposition
IBMP content quality in Loire Valley Cabernet Franc demonstrates the compound's stylistic importance, with Chinon and Bourgueil producers accepting IBMP levels of 20-35 ng/L that contribute the characteristic leafy, green pepper notes integrated into the regional style, while warmer-climate Cabernet Franc from California or Australia targets IBMP below 15 ng/L to achieve the riper, fruit-dominant profile preferred in those markets this stylistic divergence illustrates how IBMP concentration defines regional typicity rather than absolute quality, with consumer expectations determining the optimal range.
IBMP_Content_Quality_(Isobutyl-methoxypyrazine) represents the chemical quality characterizing the concentration of 3-isobutyl-2-methoxypyrazine in grape berries and wine, capturing the dominant methoxypyrazine compound responsible for the archetypal green bell pepper aroma in Sauvignon Blanc and Cabernet-family wines. IBMP (C₉H₁₄N₂O) is biosynthesized from leucine through a pathway involving pyrazine ring formation and O-methylation, with VvOMT3 methyltransferase catalyzing the final methoxylation step. Among the grape methoxypyrazines, IBMP is typically present at the highest concentrations and makes the largest contribution to green character due to its very low sensory threshold (approximately 1-2 ng/L in water, 10-15 ng/L in wine). The distinctive green bell pepper descriptor for IBMP is highly consistent across trained panels and provides one of the most reliable compound-descriptor relationships in wine aroma chemistry. IBMP concentration in grapes varies enormously depending on cultivar, developmental stage, and light exposure, with pre-véraison levels in susceptible cultivars reaching 40-100 ng/L before declining to 10-40 ng/L at harvest through light-mediated degradation. The harvest concentration range for commercially acceptable wines spans from 8-15 ng/L for ripe, fruit-forward styles through 15-30 ng/L for herbaceous-accented styles to above 40 ng/L levels that most consumers find excessively vegetal. IBMP is extracted completely during crushing and pressing, and shows essentially no change during fermentation or aging, making grape IBMP concentration directly predictive of wine IBMP levels. The imperceptible rate of IBMP degradation in wine means that wines with elevated IBMP will retain green character indefinitely, unlike fruit aromas that evolve during aging.
IBMP_Content_Quality specifically addresses the isobutyl-substituted methoxypyrazine that dominates grape methoxypyrazine composition. The parenthetical (Isobutyl-methoxypyrazine) follows the class naming convention in the uploaded structure. IBMP analysis requires sensitive GC-MS/MS methods due to the parts-per-trillion concentrations, with stable isotope dilution using deuterated IBMP providing the most accurate quantification. The sensory threshold varies among individuals, with some consumers highly sensitive (threshold below 5 ng/L) while others are relatively insensitive (threshold above 30 ng/L), creating variation in quality perception of wines with moderate IBMP levels.
3-Isobutyl-2-methoxypyrazin-Qualität
3-isobutyl-2-methoxypyrazine quality
IBMP content
IBMP-Gehalt
Isobutyl-Methoxypyrazin-Konzentration
Paprikaverbindungsgehalt
bell pepper compound concentration
isobutyl-methoxypyrazine level
Allen MS et al. (1991) Contribution of Methoxypyrazines to Sauvignon Blanc Wine Aroma. American Journal of Enology and Viticulture 42(2): 109-112 Roujou de Boubée D et al. (2000) Sensory and Chemical Analyses of the Vegetal Odour in Great Red Wines. Journal International des Sciences de la Vigne et du Vin 34(4): 163-170
IBMP Content Quality (Isobutyl-methoxypyrazine)
Vidal Blanc (Ugni Blanc × Rayon d'Or) bears an Ice_Wine_Scion_Role as Canada's most planted icewine variety, with thick skin resisting rot during extended hang-time and typically harvested at -10°C to -12°C in January-February, yielding wines of 180-280 g/L residual sugar with tropical fruit, honey, and apricot character—Inniskillin and Peller Estates producing benchmark examples at C$50-150/375ml. Riesling serves in an Ice_Wine_Scion_Role for German Eiswein and Canadian Riesling icewine, where its exceptional acidity (12-18 g/L) balances intense sweetness, producing wines of extraordinary purity and longevity. Grüner Veltliner bears an Ice_Wine_Scion_Role in Austria's Weinviertel and Wagram, providing distinctive herbal-pepper notes alongside tropical fruit concentration in Austrian Eiswein production.
A dessert wine scion role that inheres in a grafted scion cultivar by virtue of its suitability for icewine (Eiswein) production, where grapes are harvested and pressed while naturally frozen on the vine, concentrating sugars through freeze fractionation as water crystallizes and separates from the liquid must. Ice wine scions must withstand extended autumn and early winter hang-time (often until December-February) while maintaining berry integrity, resist bunch rot during the vulnerable waiting period, achieve adequate acidity to balance intense sweetness (typically 180-320 g/L residual sugar), and tolerate pressing at temperatures of -7°C to -13°C required for proper freeze concentration. The physics of freeze concentration yields extraordinarily concentrated must (35-50°Brix) as ice crystals of nearly pure water are excluded during pressing, leaving behind concentrated sugar, acid, and flavor compounds. Ice wine production is inherently risky (birds, warm spells, destructive freezing, rot) and limited to regions with reliable subfreezing temperatures, primarily Canada (Ontario, British Columbia), Germany, Austria, and select cold-climate regions globally.
Ice_Wine_Scion_Role captures the climatic requirement for reliable subfreezing temperatures that limit production to specific regions. Canadian icewine regulations (VQA) mandate natural freezing to -8°C minimum with harvest at that temperature, producing wines of 125+ g/L residual sugar. German Eiswein (minimum -7°C) is increasingly rare due to climate warming some German regions have not produced Eiswein in recent vintages. The Vidal hybrid dominates Canadian production due to its rot resistance and cold tolerance, while Riesling produces more prestigious wines but with higher losses. Icewine production efficiency is extremely low—yields of 100-200 L/ha versus 5,000-10,000 L/ha for table wine—justifying premium pricing. Artificial cryoextraction exists but is prohibited from using "icewine" designation in regulated regions.
Eissüßweinsorte
Eiswein variety
Eisweinsorte
Eisweintraube
ice wine variety
icewine grape
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Schreiner K (2001) Icewine: The Complete Story, Warwick Publishing Jackson RS (2014) Wine Science 4th Ed, Academic Press pp.: 555-565 VQA Ontario (2020) Icewine Production Standards
Ice Wine Scion Role
Shiraz vines averaging 120-160 years of age in the Penfolds Block 42 vineyard in the Barossa Valley bearing an icon wine role produce fruit for Penfolds Grange and Bin 42 Kalimna Cabernet at yields of 1.5-3 t/ha, with individual vine assessment, hand harvesting into small bins, triple sorting at the winery, and 18-20 months maturation in new American oak hogsheads, resulting in wines released at 700-950 AUD/bottle with annual production of 8,000-12,000 cases representing less than 0.1% of Penfolds' total output.
A quality tier role that inheres in a grapevine whose fruit is destined for a producer's flagship wine of the highest prestige, typically produced in limited quantities from the most exceptional vineyard parcels or oldest vines, commanding the highest prices in the portfolio and representing the pinnacle of the estate's viticultural and oenological ambition.
Icon_Wine_Role represents the apex of the commercial quality hierarchy, typically defined by retail prices exceeding $50-100 USD (varying by market) and production volumes rarely exceeding 1,000-15,000 cases. The role is characterized by extreme viticultural selectivity: icon wines frequently source from specific blocks, rows, or even individual vines identified through multi-vintage performance evaluation. The category includes both estate-grown icon wines (e.g., Screaming Eagle, Romanée-Conti) and multi-vineyard blended icons (e.g., Penfolds Grange, Vega Sicilia Único). The role imposes the most intensive and costly viticultural management of any quality tier.
Flaggschiff-Rolle
Ikonen-Wein-Rolle
Kultwein-Rolle
cult wine role
flagship wine role
icon wine vine role
McMillan R (2022) State of the US Wine Industry, Silicon Valley Bank pp: 18-24 Halliday J (2014) A History of the Australian Wine Industry, Australian Wine Research Institute Robinson J and Harding J (2015) The Oxford Companion to Wine 4th Ed, Oxford University Press pp: 196-198
Icon Wine Role
The IGT designation Toscana IGT has become one of the most commercially significant and critically acclaimed regional wine designations globally, encompassing the so-called Super Tuscan wines that emerged from the 1970s onward when innovative producers such as Marchesi Antinori (Tignanello, Solaia) and Tenuta San Guido (Sassicaia) chose to use non-traditional grape varieties (Cabernet Sauvignon, Merlot) and winemaking methods outside the constraints of Chianti DOCG regulations — these wines were initially classified as Vino da Tavola (the lowest category) despite their premium quality and pricing, prompting the creation of the IGT category in 1992 specifically to accommodate wines of demonstrated quality that fell outside DOC/DOCG production specifications.
A regional wine designation within the Italian wine classification system that identifies a wine as an Indicazione Geografica Tipica — a wine originating from a defined geographic area with characteristic qualities attributable to that origin, formally classified as a Protected Geographic Indication (IGP) under EU harmonization, representing the intermediate tier between Vino (table wine) and DOC/DOCG (Protected Designation of Origin).
IGT_Designation documents the Italian national designation that was created in 1992 (Law 164/1992) explicitly to resolve the regulatory paradox of premium wines classified at the lowest quality tier. The IGT category permits greater flexibility than DOC/DOCG in grape variety selection, blending, and winemaking methods while requiring geographic origin within the designated area: this flexibility has made IGT the designation of choice for innovative Italian producers working with international varieties, experimental blends, or unconventional production methods that conflict with traditional DOC disciplinari. Italy has approximately 120 registered IGT designations covering every wine-producing region, with Toscana IGT, Veneto IGT, and Puglia IGT among the most commercially significant. The 2009 EU harmonization formally equated IGT with PGI/IGP, but the IGT terminology remains in predominant commercial use on Italian wine labels.
IGT
IGT-Bezeichnung
Indicazione Geografica Tipica
Indicazione-Geografica-Tipica-Klassifikation
Italian IGP designation
italienische Landweinbezeichnung
Italian Law 164/1992 on wine designations EU Regulation 1308/2013 Art. 93(1)(b) Ministero delle Politiche Agricole Alimentari e Forestali (2023) Elenco IGT registrate
IGT Designation
Cabernet Sauvignon berries harvested at early maturity (BBCH 83-85, véraison completion) contain 30-60% immature seeds with green to tan coloration, soft crushable testa, 4-6 mm length, moisture content 65-75%, tannin polymerization 25-40%, extractable catechins 10-15 mg/g dry weight
An immature seed is a developing or incompletely developed seed of Vitis vinifera that has not yet completed critical developmental stages including full embryo differentiation, endosperm accumulation, seed coat lignification, or desiccation tolerance acquisition, typically exhibiting green to tan coloration, soft testa, and limited or absent germination capacity. Immature seeds represent a broad developmental spectrum from early post-fertilization stages through intermediate development that falls short of full maturity. Characteristic features include= morphological immaturity (green, yellow-green, or tan seed coat coloration, soft non-lignified testa that crushes easily, incomplete beak and chalazal development, smaller size relative to mature seeds of same cultivar), anatomical immaturity (incompletely differentiated embryo structures, partial endosperm development, non-lignified or partially lignified seed coat layers with active cell division and elongation), and biochemical immaturity (high moisture content 60-80%, low degree of tannin polymerization 20-50%, high levels of extractable monomeric flavanols, absence of desiccation tolerance). Immature seeds at harvest have significant implications for wine quality= crushing of berries containing green immature seeds releases harsh, astringent seed tannins and bitter phenolic compounds that can negatively impact wine mouthfeel and aging potential. Viticultural practice aims to minimize immature seed presence at harvest through= optimal harvest timing decisions (monitoring seed color transition from green to brown as maturity indicator), vineyard management to promote uniform ripening (canopy management, crop load adjustment, irrigation timing), and selective harvesting to exclude clusters with significant green seed content. The proportion of immature versus mature seeds within a crop varies by= environmental stress conditions (water deficit, temperature extremes can arrest seed development), pollination efficiency (poor flowering conditions lead to heterogeneous fertilization timing), and cultivar characteristics (some varieties show greater developmental synchrony than others). Seed immaturity assessment in vineyard and winery practice includes= visual inspection of seed coat color, tactile testing of seed hardness, and quantification of green seeds as percentage of total seed population in berry samples.
Immature_Seed emphasizes incomplete developmental status and is distinguished from Developing_Immature_Seed (actively progressing development) and Incomplete_Immature_Seed (arrested development) by encompassing the broad spectrum of pre-maturity developmental states. Critical for harvest timing decisions, wine quality predictions based on harsh tannin risk, and understanding environmental or physiological factors affecting seed development completion.
Green seed
Grüner Kern
Unentwickelter Traubenkern
Unreifer Samen
Unripe seed
Cadot, Y., Miñana-Castelló, M. T., & Chevalier, M. (2006). Anatomical, histological, and histochemical changes in grape seeds from Vitis vinifera L. cv Cabernet franc during fruit development. Journal of Agricultural and Food Chemistry, 54(24), 9206-9215. | Kennedy, J. A., Matthews, M. A., & Waterhouse, A. L. (2002). Effect of maturity and vine water status on grape skin and wine flavonoids. American Journal of Enology and Viticulture, 53(4), 268-274.
Immature Seed
Cabernet Sauvignon immediate seed adjacent tissue (innermost single cell layer, 28-42 μm thickness) showing extreme characteristics= cells 42-52 μm diameter (smallest in berry pericarp), walls 1.4-1.7 μm (thickest in endocarp), tannins 7.5-8.8 mg/g (maximum endocarp concentration approaching outer pericarp hypodermal levels 8-12 mg/g), positioned directly against seed coats during weeks 2-7 development but separated by 18-32 μm air gap at maturity (week 18), requiring vigorous extended maceration (20+ days, 28-30°C, enzymatic pectinase treatment, 4x daily punchdowns) for substantial extraction contributing 6-10% of total wine skin tannins when fully extracted. Syrah immediate seed adjacent tissue tannin concentration representing endocarp maximum= immediate seed adjacent (innermost layer, 0-35 μm from seed surface) 7.8-9.2 mg/g tannins next layer (35-70 μm) 6.2-7.5 mg/g (19-26% reduction) third layer (70-110 μm) 4.8-6 mg/g (38-48% reduction from maximum), demonstrating sharp exponential tannin decline from immediate seed adjacent maximum creating steepest phenolic gradient in berry over 70-110 μm distance potentially reflecting strongest seed-proximity effect on pericarp phenolic metabolism in innermost layer. Merlot immediate seed adjacent tissue seed contact dynamics examined by serial sectioning during development= week 3 post-fruit set showing seed coat cells in direct wall-to-wall contact with immediate seed adjacent tissue cells, middle lamella visible between tissues week 6 showing beginning gap formation (2-5 μm) with pectin-rich material filling gap week 10 (veraison) showing 15-25 μm air-filled gap with no direct tissue contact week 18 (maturity) showing 22-38 μm gap, demonstrating complete transition from intimate tissue contact to clear separation as berry expansion outpaced seed size increase creating locular cavity air space intervening between immediate seed adjacent tissue and seeds. Pinot Noir immediate seed adjacent tissue extraction kinetics during extended maceration= standard 10-day maceration showing minimal immediate seed adjacent tissue extraction (<5% cell breakdown, <3% tannin contribution) 15-day maceration with moderate extraction (18-28% cell breakdown, 6-10% tannin contribution) 21-day vigorous maceration with substantial extraction (45-65% cell breakdown, 12-18% tannin contribution), demonstrating that immediate seed adjacent tissue represents most difficult-to-extract pericarp tannin reservoir requiring extended vigorous maceration for substantial contribution with extraction providing late-maceration tannin increase when other more accessible tissues already extracted. Chardonnay immediate seed adjacent tissue in white cultivar showing structural presence= innermost 1-2 layers, 32-58 μm thickness, cells 45-58 μm, walls 1.2-1.6 μm, tannins 5.2-6.8 mg/g (maximum endocarp concentration in whites though lower than red cultivar immediate seed adjacent tissue 7-9 mg/g), present structurally but not extracted during white wine production where pressing separates juice from solids immediately avoiding all pericarp phenolic extraction including immediate seed adjacent tissue tannins.
An immediate seed adjacent tissue is seed coat adjacent tissue comprising the absolute innermost 1-2 cell layers in direct contact with seed surfaces during development or immediately adjacent (28-30°C, frequent punchdowns 3-4 times daily, enzymatic treatment, extended duration >15-20 days) for cell breakdown position deep within berry tissue requiring extraction of overlying layers before immediate seed adjacent tissue accessed high tannin content making extraction valuable for wine structure when extended maceration protocols employed though also creating risk of excessive extraction if maceration too vigorous or prolonged.
Immediate_Seed_Adjacent_Tissue represents the absolute innermost pericarp layer in direct or very close contact with seeds and is distinguished from Proximal_Seed_Adjacent_Tissue (slightly more distant though still seed-proximate). Immediate seed adjacent tissue shows extreme characteristics (smallest cells, thickest endocarp walls, highest endocarp tannins) reflecting innermost position. Understanding immediate seed adjacent tissue is relevant for finest-scale anatomical characterization, for investigating seed-pericarp proximity effects at tissue interface, and for winemaking where this tissue represents ultimate extraction challenge requiring most vigorous maceration conditions for access to deepest tannin reservoir.
Innerste Zellschicht
direct seed-contact tissue
direktes Samenkontakt-Gewebe
innermost layer
unmittelbar samenangrenzedes Gewebe
Cadot, Y. et al. (2006). Tannin composition and localization in grape skins and seeds. Journal of Agricultural and Food Chemistry 54':2973-2987.
Immediate Seed Adjacent Tissue
An immediate sylleptic shoot on Vitis vinifera cv. Sauvignon Blanc, axillary bud formed at node 5 in mid-May during primary shoot rapid elongation, visible lateral shoot emergence detected only 3 weeks later in early June representing very rapid sylleptic response, developed quickly to 65cm length by late July reflecting vigorous immediate growth, 9 leaves totaling approximately 1035 cm² leaf area, representing accelerated sylleptic pattern attributed to combination of high vine vigor (young vigorous vine in fertile deep soil), variety tendency toward lateral development (Sauvignon Blanc characteristically producing laterals), and favorable spring growing conditions (warm temperatures, adequate moisture) supporting rapid immediate development, this rapid lateral creating significant early canopy density requiring removal during June shoot thinning preventing excessive shading despite photosynthetic potential, with immediate timing and vigor creating management challenges outweighing benefits in this vigorous situation.
A sylleptic lateral shoot exhibiting very rapid development immediately following axillary bud formation, characterized by minimal delay between bud initiation and shoot emergence (typically 2-4 weeks), early-season timing with rapid progression from bud to visible shoot, vigorous immediate growth reflecting strong non-dormant activation, and botanical emphasis on particularly prompt sylleptic pattern with accelerated development timeline distinguishing from more gradual same-season laterals showing longer bud-to-shoot intervals. Immediate sylleptic shoots represent rapid-response lateral development, with very short interval between axillary bud formation at primary shoot node and visible lateral shoot emergence indicating strong activation signals, abundant resources, and favorable conditions promoting immediate rather than delayed or dormant bud response.
Immediate sylleptic shoots represent extreme expression of non-dormant lateral development, with very rapid progression from bud formation to shoot emergence indicating strong activation and vigorous growth conditions. The immediate pattern (2-4 week bud-to-shoot interval versus 6-10 weeks for typical prompt laterals) reflects high resource availability, strong growth signals, reduced apical dominance, variety predisposition, or environmental factors favoring rapid lateral activation. Immediate sylleptic development often occurs in vigorous situations (young vines, fertile sites, generous inputs), following early hedging (removing apex very early releasing laterals promptly), or in varieties showing strong lateral tendencies. Management challenges with immediate sylleptics include early density development (laterals emerging and growing during peak season creating rapid canopy increases), substantial vigor (rapid early growth producing large laterals), and timing complications (early emergence requiring early intervention if removal warranted). The immediate sylleptic pattern serves as indicator of high vigor conditions potentially requiring management adjustments including vigor reduction strategies, earlier or more aggressive shoot thinning, or canopy management intensification.
Sofort-Seitentrieb
fast-developing sylleptic
quick-response lateral
rapid sylleptic lateral
unmittelbarer sylleptischer Trieb
Immediate Sylleptic Shoot
Import requirement for grapevine plant material entering Australia represents one of the world's most stringent phytosanitary regimes — applicants must obtain an import permit from the Department of Agriculture, Fisheries and Forestry (DAFF), material must originate from an approved country and facility, mandatory pre-export testing for an extensive list of regulated viruses and virus-like organisms (including pathogens not yet present in Australia such as Grapevine Pinot Gris virus), all material must undergo minimum 2 years of post-entry quarantine at an approved Australian quarantine facility (currently the Knoxfield facility in Victoria) with repeated testing during quarantine, and release from quarantine requires negative results from multiple testing cycles — the entire process from import application to vineyard planting typically requiring 3-5 years. Import requirement for wine entering the EU from third countries requires a VI-1 attestation from the exporting country confirming analytical composition and production method compliance, verification that oenological practices used are permitted under EU regulations (or accepted under bilateral wine agreements), compliance with EU maximum residue limits for pesticides, allergen labeling compliance, and customs classification for tariff assessment.
Import requirement is a trade regulation entity that documents the conditions, documentation, inspections, and compliance standards that must be met for grape plant material or wine products to be legally admitted to a destination country, encompassing phytosanitary import conditions for plant material (specified pest freedom requirements, mandatory pre-shipment treatments, approved origins and pathways), wine import standards (analytical limits, permitted additive lists, labeling compliance, oenological practice acceptance), border inspection procedures (document verification, physical inspection, sampling and laboratory testing), and post-entry obligations (quarantine holding periods for plant material, market surveillance for wine products), specifying the regulatory authority, legal basis, technical requirements, and administrative processes governing importation.
Import requirement documentation addresses the destination-country perspective of viticultural trade regulation — the conditions that importing jurisdictions impose to protect their domestic viticultural sectors from phytosanitary threats, to ensure imported wines meet domestic quality and safety standards, and to enforce labeling and consumer protection regulations. The phytosanitary import requirements for grapevine plant material reflect a risk-based approach calibrated to each country's pest and disease status — countries with high phytosanitary status (Australia, New Zealand, Chile) impose the most stringent requirements because they have the most to lose from exotic pest introductions, while countries where major grapevine pests are already established (most of Europe) impose less restrictive import conditions for material from similarly affected regions. The concept of pest risk analysis (PRA) underpins phytosanitary import regulation — the systematic evaluation of the likelihood and consequences of exotic pest introduction through a specific importation pathway, resulting in phytosanitary measures calibrated to reduce the assessed risk to an acceptable level. The wine import requirement landscape is shaped substantially by bilateral and multilateral trade agreements — the EU-US Wine Agreement (2006), the EU-Australia Wine Agreement (2008/2010), and the World Wine Trade Group mutual acceptance agreements establish frameworks for recognizing each other's oenological practices, reducing duplicative testing requirements, and protecting geographic indications, though significant regulatory differences persist particularly regarding permitted additives and processing aids. The practical consequence of stringent import requirements is a significant time and cost barrier to international cultivar exchange — the 3-5 year timeline for importing vine material into Australia means that viticultural innovation through new cultivar introduction operates on a substantially longer timescale than in jurisdictions with less restrictive import regimes, creating a trade-off between biosecurity protection and the speed of cultivar portfolio adaptation.
Einfuhrbedingungen
Einfuhrvorschriften
Einführungsbedingungen
Importanforderungen
Importbestimmungen
entry requirements
import compliance
import conditions
import standards
importation regulation
Australian Government DAFF (2023) Conditions for the importation of Vitis propagative material into Australia European Parliament (2016) Regulation (EU) 2016/2031 on protective measures against plant pests FAO (2023) ISPM 11 — Pest risk analysis for quarantine pests EU-Australia Wine Agreement (2010) Agreement between the EU and Australia on trade in wine
Import Requirement
Ezerjó vines in the Mór wine district of Hungary bearing an in situ conservation role are maintained through a regional heritage viticulture program supporting 15 growers cultivating 45 hectares of this cultivar restricted almost entirely to the Mór appellation, with program interventions including subsidized virus-free planting material from sanitary-selected mother vines, technical advisory services for traditional training system maintenance, marketing support through the Móri Ezerjó PDO designation linking the cultivar to regional identity, and annual monitoring documenting population stability at current levels versus the historical peak of 300+ hectares in the 1970s.
A conservation role that inheres in a grapevine maintained within its traditional or original growing region as part of a deliberate conservation strategy that preserves the cultivar in its historically associated terroir, enabling continued local adaptation, clonal evolution, and cultural embedding while safeguarding the vine population against decline through targeted support of ongoing cultivation.
In_Situ_Conservation_Role preserves grapevine genetic diversity within the socio-ecological context where it developed, maintaining the dynamic relationship between cultivar, terroir, viticultural practice, and local culture that ex situ conservation cannot replicate. The approach depends on sustained economic viability of cultivation — in situ conservation of agricultural crops ultimately requires that someone finds it worthwhile to grow them: this distinguishes on-farm conservation from protected-area conservation of wild species and makes market access, cultural recognition, and policy support essential components of effective in situ grapevine conservation. The European Vitis Database and ECPGR on-farm conservation initiatives provide frameworks for coordinating in situ conservation across producer networks.
In-situ-Erhaltungs-Rolle
Vor-Ort-Erhaltungs-Rolle
in-field heritage conservation role
on-farm conservation role
on-site conservation role
standortgebundene Konservierungs-Rolle
ECPGR (2019) European Cooperative Programme for Plant Genetic Resources — Vitis Working Group Report Maxted N et al. (2002) Towards a definition of a crop wild relative, Biodiversity and Conservation 15: 2673-2685 Negrul AM (1946) Origin and classification of cultivated grapevine, in Baranov AI et al. (eds) Ampelography of the USSR, vol 1 pp: 159-216
In Situ Conservation Role
Carignan inactive heartwood at 40 years post-planting shows 58-78 mm radius comprising 65-72% of total trunk cross-section, vessels completely occluded with tyloses and gums preventing any water transport, dead parenchyma cells with no starch storage (0% compared to 8-14% in adjacent sapwood), light brown coloration from 3-6% phenolic compounds, wood density 0.56-0.64 g/cm³ providing structural support, shows minor white rot fungal colonization reducing density 10-15% in central core but maintaining overall mechanical integrity
Inactive heartwood is heartwood emphasizing complete loss of all physiological functions including water transport, carbohydrate storage, and metabolic activity, representing wood tissue serving only structural support and mechanical functions while parenchyma cells are dead and vessels are occluded. Inactive heartwood forms through progressive physiological shutdown= parenchyma cell death through aging and programmed senescence (cells losing viability 8-15 years post-formation depending on growth rate and conditions), complete loss of metabolic activity (respiration ceasing, enzyme activity absent, no hormone response), vessel occlusion eliminating transport capacity (tyloses formation, gum accumulation, and previous embolism permanently blocking lumens), cessation of storage function (dead parenchyma incapable of starch synthesis or hydrolysis), and transition from living functional sapwood to dead structural heartwood. Inactive heartwood anatomical characteristics include= completely occluded vessels with tyloses filling lumens (balloon-like parenchyma cell intrusions blocking vessel openings), dead parenchyma cells with degraded cellular contents (collapsed cytoplasm, broken organelles, empty or partially empty cell lumens), lignified fiber cells providing mechanical framework (structural support maintained through persistent lignified cell walls), no living cellular tissue (contrasting with metabolically active sapwood parenchyma), and variable phenolic deposits (tannins, stilbenes accumulating in cell walls and lumens contributing to darker coloration and rot resistance). Inactive heartwood formation mechanisms include= parenchyma apoptosis or necrosis (programmed or stress-induced cell death as wood ages), tyloses formation from adjacent parenchyma (living parenchyma cells in sapwood-heartwood transition zone growing into vessels occluding lumens), vascular occlusion from gums and resins (phenolic and lipid deposits accumulating in vessel lumens), loss of pit membrane function (bordered pits becoming non-functional preventing lateral water movement), and irreversible transition (once formed, inactive heartwood cannot return to functional sapwood status). Inactive heartwood distinguishing from functional sapwood includes= zero water transport capacity (versus 25-80 L/day in conducting sapwood), zero carbohydrate storage (versus 40-80 g starch in sapwood), zero metabolic activity (versus active respiration and enzyme activity in sapwood), zero living cells (versus living parenchyma throughout sapwood), but maintained mechanical properties (compressive and tensile strength comparable to or exceeding sapwood due to lignification and density). Inactive heartwood structural functions include= mechanical support of trunk and canopy (inactive heartwood providing 50-75% of total trunk cross-sectional area in old vines contributing proportionally to strength), resistance to compression (supporting vertical load of canopy and crop), resistance to bending (preventing trunk flexure under wind load), and dimensional stability (lignified cell walls maintaining trunk shape and preventing collapse). Inactive heartwood chemical characteristics include= phenolic compound accumulation (condensed tannins, stilbenes, lignin-derived phenolics at concentrations 2-8% dry weight, higher than 0.5-2% in sapwood), extractives (terpenoids, lipids, other secondary metabolites), higher lignin content (20-28% compared to 18-24% in sapwood from age-related lignification), and antimicrobial compounds (phenolics providing some decay resistance though not preventing all fungal colonization). Inactive heartwood vulnerability to decay includes= colonization by wood decay fungi (white rot and brown rot organisms degrading cellulose and lignin), progressive structural degradation (decay reducing wood density and mechanical properties), potential hollow formation (extensive decay creating central cavity in very old trunks), but some decay resistance (phenolic compounds providing antimicrobial activity slowing but not preventing fungal attack). Inactive heartwood proportion of trunk varies by age= young vines (5-10 years) showing 0-20% of trunk radius as inactive heartwood (minimal heartwood formation in young rapidly growing trunks), mature vines (10-25 years) showing 30-60% of radius (progressive heartwood accumulation as sapwood depth stabilizes), old vines (25-50+ years) showing 55-75% of radius (extensive heartwood formation with relatively thin sapwood annulus), and ancient vines (50-100+ years) showing 70-85% of radius (massive heartwood core often extensively decayed). Inactive heartwood diagnostic characteristics include= lack of dye uptake in perfusion tests (occluded vessels preventing dye transport), absence of starch in iodine staining (dead parenchyma with no storage compounds), darker coloration than adjacent sapwood (phenolic deposits producing tan, brown, or grey hues, though color difference often subtle in grapevine), and firm woody texture (maintained lignified structure unless decayed). Inactive heartwood significance in viticulture includes= determining trunk mechanical strength in old vines (large heartwood cross-section providing majority of structural support), indicating trunk age and growth history (heartwood accumulation pattern reflecting decades of cambial activity), vulnerability to decay reducing old vine stability (extensive heartwood rot compromising structural integrity requiring trunk support or replacement), and contrasting with sapwood in disease susceptibility (heartwood not infected by vascular wilt pathogens or trunk canker diseases that colonize living sapwood, but vulnerable to saprophytic wood decay fungi).
Inactive_Heartwood emphasizes complete physiological shutdown and is distinguished from Heartwood (general non-functional wood), Central_Heartwood (position emphasis), and Dense_Heartwood (structural quality emphasis) by focus on loss of all functions except mechanical support. Critical for understanding transition from functional to structural wood, age-related trunk zonation, mechanical versus physiological wood functions, and structural support in old vines with extensive non-functional cores.
Dead non-conducting wood
Funktionsloses Kernholz
Physiologically inactive xylem
Physiologisch inaktives Holz
Totes Strukturholz
Inactive Heartwood
Merlot grafted on Teleki 5C rootstock at 28 years post-grafting showing inactive rootstock heartwood 6.4 cm diameter comprising central trunk core, inactive heartwood showing 100% vessel occlusion with complete tyloses formation in all vessels eliminating all water transport capacity (hydraulic conductivity measurements confirming zero water flow through heartwood), microscopic examination revealing complete parenchyma cell death with no living cells present in heartwood (cell viability staining showing no viable cells in heartwood versus abundant living cells in surrounding sapwood), inactive heartwood moisture content 42% versus sapwood moisture 88% indicating non-conducting status, inactive heartwood containing no starch (iodine test showing zero staining in heartwood versus dark staining in active sapwood), inactive heartwood showing no metabolic activity (no respiration, no synthesis, no responses) representing completely inert tissue, inactive heartwood medium brown color from extractive accumulation, inactive heartwood providing purely structural support bearing compressive loads while contributing nothing to water transport, nutrient distribution, carbohydrate storage, or defensive functions
Inactive rootstock heartwood is rootstock heartwood defined by complete physiological inactivity and functional cessation, emphasizing that all transport, metabolic, storage, and defensive functions have permanently ended through vessel occlusion, parenchyma cell death, and conversion from living functional sapwood to dead non-functional heartwood, showing no water conduction through completely tyloses-blocked vessels, no metabolic activity in dead ray and axial parenchyma cells, no carbohydrate storage capacity, no defensive responses to wounding or infection, and no participation in any physiological processes while retaining mechanical structural support function bearing trunk loads. Inactive rootstock heartwood functional cessation includes= showing zero water transport (heartwood vessels completely occluded with tyloses eliminating all water conduction capacity contributing nothing to transpiration stream), exhibiting zero metabolic activity (all parenchyma cells dead eliminating respiration, synthesis, and all enzymatic processes), having zero storage function (no living cells present removing capacity for carbohydrate accumulation, mobilization, or any reserve functions), demonstrating zero defensive capacity (no living cells able to respond to wounding or pathogens through synthesis of antimicrobial compounds or compartmentalization reactions), and participating in zero physiological processes (heartwood completely excluded from all vine physiological activities including transport, growth, response, or signaling). Inactive rootstock heartwood complete vessel occlusion includes= showing 100% tyloses formation (all vessels in inactive heartwood completely blocked by tyloses with no patent lumens remaining), exhibiting sealed vessel interiors (tyloses completely filling vessel lumens creating absolute barrier to water movement), displaying permanent non-conducting status (vessel occlusion irreversible with inactive heartwood never regaining transport function), demonstrating zero hydraulic conductivity (water flow through inactive heartwood essentially zero contributing nothing to trunk transport capacity), and showing no embolism repair (inactive occluded vessels incapable of refilling or recovering function unlike potentially recoverable embolized sapwood vessels). Inactive rootstock heartwood complete cell death includes= containing zero living cells (all ray parenchyma and axial parenchyma cells dead with complete loss of protoplasts and organelles), showing absent cell membranes (plasma membranes and organellar membranes degraded during cell death removing cellular compartmentalization), exhibiting degraded cell contents (cytoplasm, nucleus, mitochondria, and other organelles disintegrated leaving only cell walls), demonstrating irreversible death (parenchyma cell death permanent with no recovery or regeneration possible in heartwood), and displaying only cell wall remnants (lignified secondary walls remaining as structural framework while all living contents absent). Inactive rootstock heartwood zero metabolic activity includes= showing no respiration (no oxygen consumption or CO2 production in dead heartwood tissue completely metabolically inert), exhibiting no synthesis (no production of proteins, carbohydrates, lipids, or any other compounds in heartwood lacking living cells and metabolic machinery), demonstrating no enzyme activity (no active enzymes present with all proteins degraded during cell death eliminating catalytic functions), having no energy production (no ATP generation or energy metabolism occurring in completely inactive heartwood), and showing no nutrient cycling (no uptake, transformation, or mobilization of nutrients in physiologically dead tissue). Inactive rootstock heartwood zero storage capacity includes= containing no starch (carbohydrate reserves absent from inactive heartwood due to lack of living cells for synthesis and storage), showing no dynamic reserves (no seasonal accumulation and mobilization of photosynthates in dead non-functional tissue), exhibiting no metabolic pools (no amino acids, sugars, or other metabolic intermediates present in dead cells), demonstrating no reserve mobilization (heartwood incapable of providing carbohydrates or other compounds for growth, defense, or stress response), and contributing nothing to vine energy balance (inactive heartwood representing structural ballast contributing no metabolic support to vine). Inactive rootstock heartwood zero defensive capacity includes= showing no wound response (injured heartwood producing no callus, wound periderm, or any healing tissue lacking living cells for growth), exhibiting no pathogen defense (no synthesis of antimicrobial compounds, no hypersensitive response, no active defense reactions in dead tissue), demonstrating no compartmentalization (heartwood incapable of forming reaction zones or barriers to seal off infections though existing extractives may provide passive chemical protection), having no induced resistance (no capacity for systemic acquired resistance or other active defense signaling in physiologically dead tissue), and relying solely on passive protection (decay resistance depending entirely on pre-existing extractive compounds laid down during sapwood-heartwood conversion rather than active defensive responses). Inactive rootstock heartwood in contrast to active sapwood includes= showing diametrically opposite functional status (heartwood completely inactive versus sapwood highly active physiologically), exhibiting opposite physiological roles (heartwood purely structural versus sapwood transport and metabolic functions), demonstrating opposite cellular status (heartwood cells dead versus sapwood parenchyma cells alive and active), displaying opposite hydraulic properties (heartwood non-conducting versus sapwood actively transporting water), and representing opposite ends of wood continuum (heartwood terminal aged state versus sapwood functional current state). Inactive rootstock heartwood mechanical-only function includes= providing purely structural support (heartwood contributing only through mechanical properties of lignified cell walls), bearing compressive loads (dead heartwood cells still resisting compression forces supporting trunk weight and loads), resisting bending moments (heartwood mass and stiffness contributing to trunk resistance to wind or crop-induced flexure), maintaining trunk rigidity (heartwood cells with intact lignified walls providing stiffness despite complete physiological inactivity), and enabling vertical position (heartwood structural properties helping support upright trunk orientation despite zero metabolic contribution). Inactive rootstock heartwood passive chemical properties includes= containing extractive compounds (phenolics, tannins, and other substances deposited during heartwood formation providing passive protection), showing antimicrobial activity (extractive chemicals creating hostile environment for decay organisms without active cellular defense), exhibiting antifungal properties (compounds in inactive heartwood inhibiting fungal growth through chemical toxicity rather than physiological response), demonstrating decay resistance (extractives providing protection against wood-rotting organisms despite complete cellular inactivity), and displaying cultivar-specific chemistry (extractive composition and protective capacity characteristic of rootstock variety laid down during conversion not requiring active maintenance). Inactive rootstock heartwood temporal stability includes= showing stable long-term properties (heartwood characteristics remaining constant over decades once formed not changing with season or conditions), exhibiting no seasonal variation (completely inactive heartwood showing no seasonal changes in moisture, reserves, or activity unlike dynamic sapwood), demonstrating permanent status (heartwood inactivity permanent continuing throughout trunk life without recovery or reactivation), maintaining constant composition (extractive content and wood chemistry stable in inactive heartwood not varying with vine physiological state), and persisting unchanged for decades (inactive heartwood potentially remaining structurally sound and chemically stable for 50-100+ years supporting trunk throughout vine life). Inactive rootstock heartwood boundary definition includes= marked by sapwood-heartwood interface (inactive heartwood begins where functional sapwood ends at tyloses formation and cell death boundary), showing sharp or gradual transition (boundary potentially abrupt with clear color change or gradual with transitional zone depending on formation rate), occurring at characteristic age (heartwood inactivity beginning when sapwood reaches characteristic age typically 8-15 years depending on rootstock and conditions), demonstrating centripetal progression (inactive heartwood expanding outward as new sapwood converts at inner boundary while new sapwood forms at cambium), and creating distinct functional zones (clear separation between inactive central heartwood and active peripheral sapwood). Inactive rootstock heartwood in trunk proportion includes= absent in young trunk (no inactive heartwood in years 1-7 with all wood active functional sapwood), comprising small proportion in establishing trunk (limited inactive heartwood core 10-20% of trunk area in years 8-15), showing moderate proportion in mature trunk (inactive heartwood 25-50% of trunk cross-section in years 15-40), dominating old trunk (inactive heartwood 50-75% of trunk in years 40-80 with active sapwood ring proportionally reduced), and potentially reaching 70-85% in ancient trunk (massive inactive heartwood core with minimal active sapwood ring in very old vine). Inactive rootstock heartwood moisture characteristics includes= showing low moisture content (30-60% moisture in inactive heartwood much lower than 80-120% in active sapwood), exhibiting stable moisture levels (heartwood moisture relatively constant not showing pronounced seasonal fluctuation of sapwood), demonstrating slow equilibration (moisture changes in inactive heartwood occurring slowly over weeks to months due to occluded vessels), displaying hygroscopic behavior (heartwood moisture varying with humidity but changes gradual and limited), and maintaining relatively dry state (inactive heartwood generally drier than active sapwood contributing to dimensional stability). Inactive rootstock heartwood color characteristics includes= showing darker color than active sapwood (extractive accumulation creating medium to dark brown heartwood contrasting with light tan sapwood), exhibiting cultivar-specific coloration (heartwood color ranging from light tan to dark reddish-brown depending on rootstock extractive chemistry), displaying color stability (inactive heartwood color stable over time not changing seasonally or with vine physiological state), revealing distinct color boundary (sharp or gradual color change marking transition from active sapwood to inactive heartwood), and indicating inactivity status (darker color visual indicator of physiological inactivity and functional cessation). Inactive rootstock heartwood in trunk aging includes= beginning formation at trunk maturity (inactive heartwood appearing when trunk reaches sufficient age and size for innermost sapwood conversion typically years 8-15), continuously expanding with age (inactive heartwood core growing larger as trunk ages and additional sapwood converts), progressively reducing active tissue proportion (expanding inactive heartwood decreasing percentage of trunk that is physiologically active), potentially limiting capacity in old trunk (very large inactive heartwood core with minimal active sapwood potentially constraining water supply and reserve capacity in aged vine), and representing inevitable aging process (inactive heartwood formation natural progressive phenomenon in aging trunk reflecting normal developmental progression). Inactive rootstock heartwood structural contribution includes= bearing majority of mechanical loads in large trunk (inactive heartwood core carrying most compressive and bending forces due to central position and large volume in mature and old trunk), enabling trunk diameter without excessive sapwood (inactive heartwood formation allowing large trunk diameter while maintaining active sapwood ring at functional adequate size), providing mass and stability (inactive heartwood contributing significantly to trunk weight and resistance to displacement despite zero physiological function), supporting permanent structures (inactive heartwood adequate for bearing training system, cordon, and canopy weight throughout trunk life), and maintaining structural integrity (inactive heartwood remaining mechanically sound providing reliable support for decades despite complete physiological inactivity). Inactive rootstock heartwood assessment and identification includes= identifying by color (darker heartwood contrasting with lighter sapwood indicating inactive zone), confirming by cell viability tests (staining or microscopy revealing dead cells in heartwood versus living cells in sapwood), measuring by moisture content (low heartwood moisture versus high sapwood moisture indicating inactive-active boundary), assessing by hydraulic conductivity (zero water flow through heartwood versus active flow through sapwood confirming functional distinction), and determining extent by boundary location (measuring radial distance from pith to sapwood-heartwood interface defining inactive heartwood size). Inactive rootstock heartwood significance includes= representing complete functional cessation (inactive heartwood marking absolute end of all physiological activity in transformed wood tissue), demonstrating irreversible aging (heartwood inactivity permanent showing inevitable progression of wood tissue from active functional to inactive structural state), providing purely mechanical contribution (inactive heartwood value entirely structural bearing loads without any physiological support to vine), reducing functional tissue proportion (expanding inactive heartwood decreasing percentage of trunk available for transport, storage, and defense), requiring adequate active sapwood compensation (trunk must maintain sufficient active sapwood despite inactive heartwood expansion to support vine needs), and exemplifying wood tissue development (inactive heartwood representing terminal stage of wood tissue ontogeny showing complete transformation from active to inactive state).
Inactive_Rootstock_Heartwood emphasizes complete physiological inactivity and functional cessation and is distinguished from Rootstock_Heartwood (general non-functional inner wood) by explicit focus on zero metabolic activity, zero transport, zero storage, and absolute cessation of all physiological functions. Critical for understanding complete functional loss, irreversible aging, mechanical-only contribution, dead cell status, occluded vessel state, and the absolute physiological inactivity distinguishing heartwood from all functional active sapwood tissue.
Funktionsloses Unterlagenkernholz
Inaktives Unterlagen-Kernholz
Inert rootstock heartwood
Non-functional rootstock heartwood
Physiologically dead rootstock heartwood
Physiologisch totes Wurzelstockholz
Inactive Rootstock Heartwood
Chardonnay on 99R rootstock incompatibility zone at 6 years post-grafting shows 20-35 mm zone with localized necrotic regions (brown-black dead tissue 3-8 mm depth disrupting continuity), incomplete vascular connections (xylem vessels showing 40-60% discontinuity across interface, phloem disrupted), non-functional cambium in 30-50% of circumference preventing coordinated secondary growth, excessive asymmetric swelling 145-180% diameter creating pronounced bulge, bark showing cracks and necrotic patches, water transport capacity reduced to 45-65% causing midday scion wilting despite adequate soil moisture, vine showing progressive decline (reduced vigor, small chlorotic leaves, poor fruit development) over 3 years suggesting incompatibility-induced dysfunction potentially leading to vine death or requiring replacement
Incompatibility zone is a region within graft union where scion and rootstock tissues fail to integrate properly, characterized by incomplete cellular adhesion, necrotic tissue, vascular discontinuities, or mechanical weakness, representing failed or inadequate tissue compatibility preventing normal union healing and function potentially causing vine decline or death. Incompatibility zone manifestations includes= localized necrosis (dead tissue regions interrupting union continuity, appearing as brown to black discolored zones), cellular discontinuity (scion and rootstock cells failing to adhere properly leaving gaps or separations), vascular discontinuities (xylem vessels or phloem sieve tubes not forming connections across zone creating transport barriers), cambial discontinuities (vascular cambium absent or non-functional in zones preventing secondary growth strengthening), abnormal tissue proliferation (excessive disorganized callus formation without proper differentiation), and mechanical weakness (poor tissue cohesion creating union vulnerable to fracture under stress). Incompatibility zone cellular and molecular basis includes= cell recognition failure (scion and rootstock cells unable to recognize each other as compatible preventing adhesion), adhesion molecule incompatibilities (cell wall proteins, pectins, or other adhesion molecules between partners preventing proper bonding), phytotoxin production (one or both partners producing compounds toxic to other partner causing cell death at interface), phenolic accumulation (excessive phenolic compounds oxidizing and forming barriers preventing tissue integration), programmed cell death (necrosis developing at interface from incompatibility-triggered cell death pathways), and genetic incompatibility (fundamental molecular differences between scion and rootstock genomes preventing successful integration). Incompatibility zone types includes= immediate incompatibility zones (appearing within weeks to months post-grafting, graft failing to establish from inability to form initial callus bridge or vascular connections), delayed incompatibility zones (developing years after initially successful union, vine showing progressive decline 3-15 years post-grafting from deteriorating union quality), localized incompatibility zones (affecting portions of union circumference while other sections heal normally creating asymmetric weakness), and complete incompatibility (entire union interface showing poor integration causing severe functional limitations or vine death). Incompatibility zone anatomical characteristics includes= necrotic tissue presence (dead brown-black cells at scion-rootstock interface preventing integration), reduced or absent callus formation (minimal callus proliferation indicating poor tissue response to wounding and incompatibility), disorganized vascular tissues (poorly formed or absent xylem and phloem connections creating transport barriers), cambial gaps (sections lacking functional cambium preventing secondary growth and union strengthening), parenchyma discontinuities (ray cells not continuous across union limiting lateral transport), and excessive swelling (union bulge >130-200% diameter from disproportionate tissue proliferation attempting to compensate for poor integration). Incompatibility zone functional consequences includes= reduced water transport (vascular discontinuities limiting xylem conductance 30-70% of normal causing water stress symptoms in scion), impaired photosynthate transport (phloem discontinuities reducing basipetal sugar flow affecting root development and storage), hydraulic dysfunction (increased resistance at union potentially causing midday wilting, reduced vigor despite adequate soil moisture), nutrient imbalances (disrupted vascular connections affecting mineral uptake and distribution causing deficiency or toxicity symptoms), and hormone signaling disruption (impaired basipetal auxin transport and acropetal cytokinin movement affecting growth regulation coordination). Incompatibility zone mechanical consequences includes= structural weakness (poor tissue adhesion and organization creating union vulnerable to breakage), fracture under stress (incompatibility zones failing under wind, crop load, or management stress causing trunk breakage at union), progressive deterioration (mechanical failure risk increasing with age as incompatibility symptoms worsen), and vine loss (severe incompatibility potentially causing catastrophic union failure requiring vine replacement). Incompatibility zone symptoms and indicators includes= excessive union swelling (circumference >130-150% diameter suggesting severe incompatibility), growth disparities (marked differences in trunk diameter above and below union indicating transport limitations), decline symptoms (reduced vigor, small leaves, chlorosis, poor fruit development suggesting inadequate union function), necrotic bark areas (dead bark zones at union indicating underlying tissue death), union cracks or separations (visible fissures or gaps in bark suggesting internal discontinuities), mechanical instability (union movement or flexing under pressure indicating weak adhesion), and progressive worsening (symptoms intensifying over years characteristic of delayed incompatibility). Incompatibility zone assessment methods includes= visual external examination (observing excessive swelling, bark necrosis, abnormal form), palpation (detecting soft zones, movement, separation suggesting internal problems), growth monitoring (measuring trunk diameters, shoot vigor above and below union detecting disparities), dissection (cutting through union revealing necrotic zones, vascular discontinuities, tissue disorganization), microscopy (examining cellular adhesion, tissue differentiation, presence of phenolic barriers or necrotic cells), dye perfusion (testing vascular continuity revealing transport blockages), mechanical testing (measuring union breaking strength detecting weakness), and physiological assessment (measuring water potential, nutrient status, photosynthesis indicating functional impairment). Incompatibility zone in different combinations includes= common in certain scion-rootstock pairings (e.g., some V. vinifera cultivars showing incompatibility with specific rootstock genotypes), rare in well-tested combinations (commercial rootstocks like 3309C, SO4, 5BB, 101-14Mgt generally highly compatible with most V. vinifera scions), genetic determination (compatibility controlled by multiple genes, specific allele combinations determining integration success or failure), and cultivar-specific patterns (some scion cultivars or rootstock clones showing characteristic incompatibility tendencies with particular partners). Incompatibility zone development timeline includes= immediate incompatibility (poor callus formation and vascular differentiation visible weeks to months post-grafting, union failing within first year), early delayed incompatibility (initially adequate union deteriorating in years 2-5 showing progressive necrosis, vascular dysfunction, decline), late delayed incompatibility (union functioning adequately for 5-15 years then showing progressive failure possibly from cumulative stress or age-related changes), and variability (incompatibility expression timing depending on genetic factors, environmental stress, management practices). Incompatibility zone management and remediation includes= prevention through proper selection (choosing well-tested compatible scion-rootstock combinations minimizing incompatibility risk), early detection (monitoring young unions for excessive swelling, poor growth, other warning signs), evaluation and culling (removing severely incompatible unions within first 2-3 years before vineyard establishment investment), bridge grafting potential (re-grafting with compatible interstock between incompatible partners possibly resolving problems), and replacement (severe incompatibility requiring vine removal and replanting with compatible combination). Incompatibility zone significance includes= determining graft failure (incompatibility zones causing union failure, vine decline or death within 2-15 years depending on severity), limiting rootstock utilization (incompatibility restricting which rootstocks can be used with specific scion cultivars), affecting vineyard establishment (incompatibility causing vine losses 5-30% in problematic combinations reducing stand establishment), requiring testing (new scion-rootstock combinations requiring evaluation for compatibility before commercial use), and emphasizing proper selection (compatibility being fundamental consideration in nursery propagation and vineyard planning requiring use of proven compatible combinations).
Incompatibility_Zone emphasizes failed tissue integration and is distinguished from Graft_Union_Zone (successful or healing unions), Healed_Graft_Union_Zone (complete integration), Delayed_Incompatibility_Zone (late-developing problems), and Visible_Incompatibility_Zone (observable symptoms) by focus on general incompatibility manifestations. Critical for understanding graft failure mechanisms, cellular and molecular incompatibility basis, functional and mechanical consequences, symptom recognition, compatibility assessment, and incompatibility as major limitation in grafted viticulture requiring careful scion-rootstock selection and union monitoring.
Abstoßungs-Bereich
Failed union zone
Graft incompatibility zone
Inkompatibilitätszone
Rejection zone
Verwachsungs-Störzone
Incompatibility Zone
Cabernet Sauvignon incomplete immature seeds at harvest (BBCH 89) retain yellow-green coloration, 4-5 mm length (compared to 7-8 mm for fully developed seeds), 70% moisture content despite 3-4 weeks berry ripening time, tannin polymerization 20-30%, extractable catechins 18-22 mg/g dry weight
An incomplete immature seed is an incompletely developed seed of Vitis vinifera that has arrested development before achieving full maturity and will not progress to mature seed status even with extended berry ripening time, characterized by permanent retention of immature morphological and biochemical characteristics including green or tan coloration, soft testa, and absent or minimal germination capacity. Incomplete immature seeds result from developmental arrest at various stages due to= inadequate fertilization (abnormal embryo formation, limited endosperm development), environmental stress during critical developmental windows (water deficit during cell division phase, temperature extremes during embryogenesis), nutritional limitations (insufficient assimilate supply to developing seeds, competition from other developing seeds or berry tissues), or genetic/physiological factors affecting seed development regulation. These seeds are distinguished from developing immature seeds by evidence of arrested development= lack of progressive color change over time, cessation of size increase during berry ripening, persistent high moisture content without progression toward desiccation, and absence of continued lignification or tannin polymerization. Morphological characteristics include= permanently green, yellow-green, or light tan coloration without progression to brown, small size relative to cultivar potential (typically 40-70% of fully developed seed dimensions), soft crushable testa lacking significant lignification, and poorly defined or absent beak and chalazal structures. Biochemical indicators include= low tannin polymerization index (15-40%) without temporal progression, high moisture content (60-80%) maintained throughout berry ripening, low dry matter accumulation, and high extractability of monomeric flavanols. Incomplete immature seeds have significant implications for wine quality= they contribute disproportionately to harsh astringency and bitterness if crushed during harvest and processing, as incompletely polymerized tannins and high concentrations of extractable catechins are readily released into must and wine. Viticultural factors influencing incomplete seed development include= severe water stress during Phase I berry growth (2-4 weeks post-fruit set), extreme heat events during flowering and early berry development, excessive crop load leading to competition for assimilates, and canopy shading reducing photosynthate availability.
Incomplete_Immature_Seed emphasizes arrested development and permanent immaturity and is distinguished from Developing_Immature_Seed (active developmental progression) by evidence of developmental cessation despite continued berry ripening. Critical for understanding wine quality risks from harsh tannin extraction, identifying viticultural stress factors causing developmental arrest, and differentiating normal developmental timing from pathological arrested development.
Arrested development seed
Dauerhaft unreifer Kern
Entwicklungsarrestierter Traubenkern
Permanently immature seed
Unvollständig entwickelter Samen
Ojeda, H., Deloire, A., & Carbonneau, A. (2001). Influence of water deficits on grape berry growth. Vitis, 40(3), 141-145. | Cadot, Y., Miñana-Castelló, M. T., & Chevalier, M. (2006). Anatomical, histological, and histochemical changes in grape seeds from Vitis vinifera L. cv Cabernet franc during fruit development. Journal of Agricultural and Food Chemistry, 54(24), 9206-9215.
Incomplete Immature Seed
Incomplete stage transition assessment at veraison in heat-stressed Grenache reveals the asynchrony affecting harvest decisions, with heat event during early veraison disrupting transition synchrony, resulting in 35% of berries completing color change to full purple, 40% showing partial color (pink, intermediate anthocyanin), and 25% remaining green despite 28 days post-veraison onset (normal completion within 14 days), with the incomplete transition forcing harvest compromise between over-ripe early berries (26 Brix, declining acidity) and under-ripe late berries (19 Brix, high malic acid), demonstrating how incomplete stage transition creates quality-limiting heterogeneity requiring difficult management tradeoffs.
Incomplete stage transition quality is a developmental completeness quality that represents partial progression from one developmental stage to the next without achieving full transition, characterizing the condition where vines begin transitioning to a subsequent stage but fail to complete the transition, resulting in mixed populations of organs showing characteristics of both stages or intermediate developmental states not corresponding to any defined stage. Within the BFO framework, this quality specifically addresses the partial transition dimension of developmental disruption, distinguishing incomplete transitions from complete transitions that are merely slow or gradual while recognizing that incomplete transitions create phenological ambiguity and may indicate developmental discoordination requiring management attention. The ontological significance lies in incomplete stage transition creating populations of organs at inconsistent developmental states, where the failure to complete transitions produces heterogeneous fruit populations (berries at different ripening stages within clusters), asynchronous canopy development (shoots at different stages within the vine), or mixed reproductive states (clusters with flowers and developing berries simultaneously), with this heterogeneity complicating management decisions and potentially compromising fruit quality through inconsistent maturity at harvest. The causes of incomplete stage transition include environmental fluctuations during transition periods (temperature swings disrupting synchronous progression), resource competition (strong sinks drawing resources from transitioning organs), stress events during transition (heat or water stress halting some organs while others continue), and inherent cultivar characteristics (some cultivars more prone to asynchronous transitions), with the cause determining whether the condition is manageable or inherent. The fruit quality implications of incomplete stage transition are substantial, particularly for veraison and ripening transitions where incomplete progression produces "red and green" berry populations that cannot simultaneously achieve optimal maturity, forcing harvest timing compromises between over-ripe early-transitioning berries and under-ripe late-transitioning berries that affect wine quality through heterogeneous fruit composition. The management responses to incomplete stage transition include extended monitoring during transition periods, selective harvesting separating differently developed populations, and cultural practices promoting synchronous development (consistent water supply, balanced vine condition) to minimize transition incompleteness in subsequent seasons. Understanding incomplete stage transition quality enables recognition of developmental asynchrony, assessment of its severity and causes, and management strategies for either accommodating or preventing incomplete transitions affecting fruit quality.
Incomplete_Stage_Transition_Quality addresses partial progression between stages creating heterogeneous organ populations. This quality causes phenological ambiguity and fruit quality challenges from mixed maturity. The class supports harvest decision support and developmental assessment through incomplete transition characterization.
asynchrone Entwicklung
asynchronous development
incomplete phenological shift
partial transition
partieller Übergang
transition failure
unvollständiger Stadienübergang
Übergangsversagen
Sadras VO, Moran MA (2012) Elevated temperature decouples anthocyanins and sugars in berries of Shiraz and Cabernet Franc. Australian Journal of Grape and Wine Research 18(2): 115-122 Coombe BG, McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6(2): 131-135
Incomplete Stage Transition Quality
An incomplete tendril at node 8 on a shoot of Vitis vinifera cv. Pinot Noir in Burgundy, France, measuring 13.5 cm length (approximately 75% of normal 18 cm), exhibiting complete elongation and bifurcated architecture suggesting near-normal morphological development, well-developed and fully lignified basal 5 cm providing strong structural support and dark brown coloration, but distal 8.5 cm remaining green, flexible, and poorly lignified despite achieving full length extension, complete vascular development throughout entire length including functional xylem and phloem, but mechanosensory apparatus well-developed only in basal half with distal regions showing sparse or absent responsive cells, occurring following sequential stress events including early season phosphorus deficiency affecting lignification processes followed by mid-season recovery through fertilization that restored general growth but not lignin biosynthesis, showing strong coiling response in basal regions but weak or absent response in distal portions despite contact, functionally supporting moderate loads (400 grams) through basal attachment but vulnerable to distal failure, measured at BBCH stage 79 (majority of berries touching) in early August.
A partially developed tendril exhibiting strongly asynchronous or unbalanced development across different tissue systems and developmental parameters in Vitis vinifera, characterized by successful completion or near-completion of some developmental aspects such as elongation growth achieving 60-90% normal length while other critical aspects such as mechanosensory differentiation, structural reinforcement, or vascular maturation remain substantially incomplete, creating functional deficits despite apparently advanced morphological development, or conversely showing well-developed tissue differentiation in limited regions while other regions remain at early developmental stages creating spatial heterogeneity in maturation status. Incomplete tendrils result from complex stress patterns including sequential stress events affecting different developmental processes at different times, localized damage or disease affecting specific tissue regions or developmental pathways while sparing others, developmental regulatory failures affecting specific gene expression programs or hormonal responses while permitting others to proceed normally, or resource allocation conflicts where competition for specific nutrients or metabolites limits particular biosynthetic pathways while general growth processes continue. These structures represent the most heterogeneous category of developmental outcomes, requiring detailed anatomical and functional assessment to characterize specific completions and deficiencies, and providing diagnostic information about the nature and timing of limiting factors through analysis of which developmental processes succeeded versus failed and the spatial or temporal patterns of developmental heterogeneity.
Incomplete tendrils are distinguished from other partially developed tendrils by strongly asynchronous development patterns showing substantial achievement in some parameters combined with marked deficiency in others, rather than proportional incomplete development across all features (as in half-developed tendrils). This classification requires detailed assessment of multiple developmental parameters to identify specific completions and deficiencies, making diagnosis more complex but more informative about limiting factor identity. Incomplete development patterns provide diagnostic value through correlation of successful versus failed processes with known biosynthetic pathways and resource requirements - lignification failure despite growth success suggests phenylpropanoid pathway limitation, sensory apparatus failure despite structural success indicates specialized developmental program disruption, and spatial gradients of maturation suggest progressive resource depletion or sequential stress events. Management responses require targeted intervention addressing specific limiting factors rather than general resource supplementation.
asynchron entwickelte Ranke
asynchronous development tendril
heterogene Ausreifung Kletterorgan
heterogeneous completion tendril
unbalanced maturity tendril
unvollständige Ranke
Based on Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 135-156 Petrie et al. (2000) "The effect of leaf removal and canopy height on whole-vine gas exchange and fruit development of Vitis vinifera L. Sauvignon blanc" Functional Plant Biology 27=173-184 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 521-544.
Incomplete Tendril
In a Bordeaux Cabernet Sauvignon vineyard, increasing anthocyanin accumulation was quantified during 6 days in mid-September Year N+1 (BBCH 84) when skin anthocyanins rose from 185 mg/kg to 695 mg/kg (average 85 mg/kg per day), with malvidin-3-glucoside increasing from 105 to 390 mg/kg, berry Brix advancing 14.5 to 19.8°, firmness declining to 1.8 N, and ABA sustained at 1650-1850 ng/g under favorable conditions (daily high 26-28°C, night 14-17°C, moderate water deficit stem ψ -1.1 MPa) in eastern canopy exposure.
Increasing_Anthocyanin_Accumulation represents the quantitative escalation in berry skin anthocyanin content during progressive color development when biosynthetic rates reach peak velocity and total pigment concentrations rise exponentially from color break levels toward cultivar-specific maxima that determine final berry color intensity and wine quality potential. This process occurs during mid-veraison (BBCH 84), typically 3-8 days post-color break, and is characterized by anthocyanin accumulation rates of 50-150 mg/kg skin fresh weight per day (compared to initial synthesis 10-30 mg/kg per day) increasing total skin anthocyanins from 100-200 mg/kg at early progressive development to 400-1000 mg/kg at late progressive phase representing 4-10 fold amplification. Kinetics of increasing accumulation follow sigmoidal patterns: initial lag phase (days 0-2 post-color break) shows relatively slower rates 20-40 mg/kg per day as biosynthetic machinery approaches maximum capacity, exponential phase (days 3-6) exhibits peak rates 60-150 mg/kg per day when all pathway enzymes operate at maximum with optimal substrate availability and favorable conditions, plateau approach (days 7-10) displays declining rates 30-60 mg/kg per day as concentrations near cultivar-specific maxima and potential feedback inhibition or substrate limitation begins, and final plateau (days 10-14) shows minimal further increase 35°C can completely arrest accumulation producing pale inadequately colored fruit even if conditions later improve. Light quantity and quality effects inclu UV-B radiation (280-315 nm) enhances accumulation rates 35-60% through UVR8-mediated gene induction, blue light (430-470 nm) increases rates 20-40% via cryptochrome signaling, red light shows moderate 10-20% enhancement, while excessive visible radiation combined with heat can reduce accumulation through photoinhibition and thermal stress demonstrating complex light-temperature interactions requiring optimization. Water deficit impacts on increasing accumulation show: moderate deficit (stem ψ -0.9 to -1.3 MPa) enhances accumulation 20-35% through elevated ABA maintaining high biosynthetic gene expression and berry concentration effects, severe deficit (<-1.5 MPa) impairs accumulation despite very high ABA through general metabolic limitation reducing final anthocyanin 35-60%, while abundant water dilutes berries and may suppress ABA-responsive genes reducing accumulation 25-45% compared to moderate deficit demonstrating complex dose-dependent relationships. Cultivar differences in increasing accumulation rates reflect: high-accumulator varieties like Petit Verdot and Tannat achieve peak rates 100-150 mg/kg per day reaching final levels 2000-3000 mg/kg, moderate accumulators like Cabernet Sauvignon and Merlot show 60-100 mg/kg per day peaks with 800-1500 mg/kg final, while lower-accumulator cultivars like Pinot Noir (despite rapid color break) display 40-70 mg/kg per day maximum achieving 600-1200 mg/kg final demonstrating genetic variation in both rate and duration of accumulation. Coordination with berry development processes: increasing anthocyanin accumulation coincides with peak sugar accumulation rates (1.5-3.5% Brix per week) suggesting integrated carbon metabolism regulation, parallels maximum berry softening velocity (firmness declining 3-6% per day), occurs during intense acid decline particularly malate consumption, and synchronizes with phenolic compound synthesis including tannins and flavonols creating complex flavor and structural wine components. Quality implications inclu final anthocyanin levels strongly correlate with wine color intensity (r² = 0.85-0.95) making increasing accumulation monitoring critical for quality prediction, rate and duration of increasing accumulation affect anthocyanin composition particularly acylated derivative proportions influencing color stability and aging potential, and environmental optimization during this phase through irrigation, canopy management, and crop load adjustment can substantially enhance final color 30-60% compared to suboptimal conditions.
Increasing_Anthocyanin_Accumulation is distinguished from Initial_Anthocyanin_Synthesis by representing peak rates rather than onset, from Progressive_Color_Development by focusing on quantitative accumulation rather than spatial progression, and from final accumulation by exhibiting exponential increase rather than plateau. Peak rates: high accumulators 100-150 mg/kg/day, moderate 60-100 mg/kg/day, lower 40-70 mg/kg/day. Temperature optimum 23-28°C. Moderate deficit enhances 20-35%. Increasing_Anthocyanin_Accumulation phase critically determines final berry color intensity and wine quality potential.
amplifying color compound accumulation
escalating pigment synthesis
eskalierende Pigment-Synthese
intensifying anthocyanin production
intensivierende Anthocyan-Produktion
rising anthocyanin content
steigende Anthocyan-Konzentration
verstärkende Farbstoff-Akkumulation
Keller M (2020) The Science of Grapevines 3rd Ed pages 234-236 anthocyanin accumulation dynamics Tarara JM, Lee J, Spayd SE, Scagel CF (2008) Berry temperature and solar radiation alter acylation, proportion, and concentration of anthocyanin in Merlot grapes Amer J Enol Vitic 59:235-247 environmental effects on accumulation kinetics Cohen SD, Tarara JM, Kennedy JA (2008) Assessing the impact of temperature on grape phenolic metabolism Anal Chim Acta 621:57-67 temperature sensitivity of accumulation.
Increasing Anthocyanin Accumulation
Chardonnay in Champagne shows increasing berry pliability during late August-September Year N+1 at BBCH 85-89, with berry texture progressively softening from initial 2.3 N to mature 1.4 N over 5-6 weeks ripening, berries increasingly pliable to touch yielding readily under gentle finger pressure, tactile assessment showing progressive softness increase, cell wall structural integrity extensively lost from cumulative pectin degradation, middle lamella substantially degraded creating tissue yielding, berries requiring careful handling to avoid bruising, eventual soft pliable ripe texture characteristic of mature berries ready for harvest, under cool maritime conditions of 14-20°C temperature, typical of this early-bursting cultivar achieving soft ripe texture for sparkling wine production
Increasing berry pliability represents the progressive texture softening outcome phase during progressive softening when berry texture becomes increasingly soft and yielding from cumulative cell wall degradation, berries progressively more pliable to touch responding readily to gentle pressure, texture transformation from firm resistant to soft yielding, pliability increasing as structural integrity declines from pectin loss, and progressive pliability establishing characteristic soft ripe berry texture as ripening progresses over 4-8 weeks. This texture softening outcome phase occurs during late summer through autumn in late August-October in temperate regions Year N+1, representing tactile softening progression through ripening. Pliability increase mechanism involves cumulative cell wall structural weakening from pectin degradation reducing resistance to deformation, middle lamella loss reducing cell-cell adhesion creating tissue yielding, possible minor turgor changes though structural effects dominant, progressive texture change from firm to soft, and pliability as functional softening outcome. Berry pliability definition shows pliability as ease of berry deformation under applied force, pliable berries yielding readily to pressure, pliability inversely related to firmness, and pliability as texture characteristic. Texture transformation progression shows berry texture progressively softening from initial softening through ripening to ripe, transformation from firm hard to soft yielding over weeks, progressive change as ripening feature, and texture as maturity indicator. Initial texture state shows berries moderately firm after initial softening 2.0-3.5 N penetrometer resistance, texture noticeably softer than pre-veraison but still resistant, and initial state as softening baseline. Progressive softening shows berries becoming progressively softer during ripening, texture increasingly yielding, eventual soft pliable state at maturity 1.0-2.5 N, and progression as ripening characteristic. Final ripe texture shows berries very soft and pliable at full ripeness, yielding readily to touch, texture characteristic of mature fruit, and ripe state as softening endpoint. Tactile assessment shows subjective pliability evaluation through gentle pressure, berries increasingly soft to touch, pliability readily detected by experienced evaluators, and tactile as practical pliability assessment. Pressure response shows soft berries deforming under gentle finger pressure, berries easily compressed, deformation recovering slowly or not at all, and response as pliability indicator. Ease of deformation shows pliable berries requiring minimal force for deformation, resistance declining progressively, eventual very low resistance at ripeness, and deformation ease as pliability measure. Tissue yielding shows berry tissue yielding readily under pressure from reduced structural integrity, yielding from cell-cell adhesion loss and wall weakening, tissue response as pliability mechanism, and yielding as texture basis. Cell wall structural weakness shows extensively degraded cell walls providing minimal resistance, wall pectin loss creating fragile structure, weak walls enabling easy deformation, and structural weakness as pliability cause. Middle lamella extensive loss shows middle lamella substantially degraded by ripening, intercellular adhesion minimal, cells easily separated, and middle lamella loss creating tissue yielding. Cell-cell adhesion minimal shows cells very loosely attached in pliable berries, adhesion insufficient to maintain tissue integrity under pressure, adhesion loss enabling yielding, and minimal adhesion as pliability feature. Berry handling characteristics shows pliable berries very delicate requiring careful handling, berries easily damaged by pressure, susceptibility to bruising and crushing, and handling sensitivity as pliability consequence. Damage susceptibility shows soft pliable berries bruising from minor impacts, cell rupture from gentle compression, juice release from handling, and damage sensitivity increasing with pliability. Harvest timing implications shows pliability as ripeness indicator affecting harvest decisions, table grapes harvested before excessive pliability, wine grapes allowed to soften fully, and pliability affecting harvest management. Skin-flesh properties shows skin possibly loosening from flesh in very pliable berries, skin-flesh separation from middle lamella degradation, separation affecting processing, and skin properties as pliability aspect. Juice release characteristics shows pliable berries releasing juice readily when crushed, softness facilitating juice extraction during processing, texture affecting crushing efficiency, and juice release as functional property. Berry appearance shows pliable berries possibly appearing slightly deflated or wrinkled in some cases, surface changes from structural loss, appearance as visual pliability indicator, and surface as textural sign. Berry size shows berry size relatively stable during progressive softening though water uptake may increase volume slightly, size independent of pliability, and size-pliability distinction. Pliability-firmness relationship shows pliability inversely related to firmness, increasing pliability corresponding to firmness decline, strong negative correlation, and relationship as texture measure. Sensory evaluation shows trained panels detecting pliability progression, sensory scores correlating with instrumental firmness, sensory as pliability assessment method, and evaluation as quality measure. Consumer perception shows consumer preferences varying with berry use, soft pliable texture preferred for wine grapes, firmer texture preferred for table grapes, and perception as quality factor. Quality implications shows pliability affecting berry quality depending on use, optimal pliability varying by application, excessive softness problematic for table grapes, appropriate softness essential for wine grapes, and quality as pliability context. Environmental effects on pliability include temperature affecting pliability progression rate through effects on enzyme activities, warm conditions promoting faster pliability increase, cool conditions slowing progression, and temperature as pliability rate factor. Temperature effects shows accumulated heat affecting pliability extent, temperature affecting ripening rate and softening, and temperature as major environmental factor. Cultivar variation shows final ripe pliability varying among cultivars, some cultivars remaining relatively firmer, others becoming very soft, and genetic variation affecting pliability. The process represents second aspect of progressive softening establishing texture softening outcome, following continued cell wall degradation and concurrent with firmness decline. Duration pliability increasing progressively over 4-8 weeks ripening. Regional timing shows increasing berry pliability during late August-October through ripening. Viticultural significance includes representing tactile softening outcome affecting quality and handling, determining harvest maturity and berry management, demonstrating structural degradation functional consequence, and establishing pliability as ripeness characteristic. Research applications include analyzing pliability progression and mechanisms, investigating pliability-structural relationships, evaluating sensory and instrumental pliability assessment, quantifying pliability changes through ripening, and assessing pliability as quality and maturity indicator.
Increasing_Berry_Pliability is second aspect of Progressive_Softening, distinguished by texture softening outcome. Represents progressive change over 4-8 weeks ripening. Firmness 2.0-3.5 → 1.0-2.5 N. Berries increasingly soft and yielding. Mechanism: cumulative cell wall degradation, middle lamella loss, minimal cell-cell adhesion. Turgor changes minor. Tactile assessment: readily deformable. Handling sensitivity high. Sensory quality indicator. Follows Continued_Cell_Wall_Degradation. Concurrent with Firmness_Decline. Temperature affects progression rate.
Beeren-Weichheit-Zunahme
Textur-Nachgeben
berry softness increase
progressive Erweichungs-Ergebnis
progressive softening outcome
tactile softening
texture yielding
zunehmende Beeren-Nachgiebigkeit
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 485-488
Increasing Berry Pliability
Indexing results for Pinot Noir clone 115 (ENTAV) record grafting onto the standard EPPO indicator panel — Vitis rupestris St. George (negative for stem pitting and stem grooving after 2 growing seasons), Vitis vinifera Pinot Noir LN33 (negative for vein necrosis, vein mosaic, and fleck after 3 growing seasons), Kober 5BB (negative for Kober stem grooving), and 110 Richter (negative for corky bark symptoms) — collectively confirming absence of all biologically detectable graft-transmissible agents as of the 2019 evaluation cycle. Indexing results for a Furmint candidate clone detected vein mosaic symptoms on the LN33 indicator (chlorotic vein banding appearing in the second growing season), subsequently confirmed as GFkV infection by ELISA, demonstrating the complementary role of biological indexing in detecting low-titer infections missed by initial serological screening.
Indexing result is a virus testing status entity that records the outcome of biological indexing (also termed woody indexing or indicator plant testing) performed on a clonal line, in which bud or bark material from the candidate clone is grafted onto defined virus-sensitive indicator plant species and the resulting graft combinations are observed over one to three growing seasons for the development of virus-specific symptoms, providing the classical biological confirmation of virus presence or absence that detects physiologically active infections including those caused by agents not yet characterized or detectable by serological and molecular methods.
Biological indexing represents the oldest and most comprehensive approach to grapevine virus detection, with the unique ability to detect any graft-transmissible agent causing visible symptoms on indicator plants, regardless of whether the causal agent has been identified, sequenced, or targeted by molecular or serological assays. This "open-ended" detection capability makes biological indexing the only method capable of identifying truly novel or uncharacterized pathogens — a critical advantage given that new grapevine-associated viruses continue to be discovered (GPGV was first described in 2012, GRLDaV in 2020). The standard EPPO indicator panel includes Vitis rupestris St. George (for stem pitting/grooving agents, fanleaf deformation), V. vinifera LN33 or Pinot Noir (for fleck, vein mosaic, vein necrosis, leafroll), Kober 5BB or 110 Richter (for corky bark, stem grooving), and in some protocols V. berlandieri × V. riparia SO4 (for Enation disease). The principal limitations of biological indexing are its long duration (2-3 years for complete evaluation), high cost (greenhouse space, labor, plant material), and subjective symptom assessment dependent on observer expertise and environmental conditions. Symptom expression on indicator plants can be influenced by temperature (some symptoms only appear under specific thermal regimes), vine age, and rootstock vigor, requiring careful control of growing conditions. Despite the rise of molecular diagnostics, many certification authorities retain biological indexing as a mandatory component for the highest certification levels (Nuclear, Pre-basic/Initial), recognizing its irreplaceable role as the ultimate biological confirmation of health status.
Indexierungsergebnis
Indikatorpflanzentest
Veredelungsindexierung
bioassay result
biological indexing result
biologischer Indexierungsbefund
biologischer Testbefund
graft indexing result
indicator plant test result
woody indexing result
Walter B, Martelli GP (1997) Clonal and sanitary selection of the grapevine. INRA Editions, pp.: 43-95 Martelli GP (2014) Directory of virus and virus-like diseases of the grapevine. Journal of Plant Pathology 96(1S): 1-136 EPPO (2008) Certification scheme for pathogen-tested material of grapevine varieties. EPPO Bulletin 38(3): 422-429
Indexing Result
Xinomavro in Macedonia, Greece bearing an indigenous cultivar role shows genetic profiles consistent with local domestication from wild Vitis vinifera populations in the northern Greek peninsula, with no identified parent cultivars from other regions, documented presence in Naoussa since at least the 14th century, and SSR marker diversity patterns indicating a center of origin in the Vermio mountain foothills where it constitutes 85-90% of vineyard plantings under PDO Naoussa regulations permitting no other red variety.
A traditional cultivar role that inheres in a grapevine cultivar believed to have originated through primary domestication or spontaneous seedling selection within the region where it is currently cultivated, with no documented evidence of introduction from external sources and genetic evidence consistent with local origin from wild or feral Vitis vinifera subsp. sylvestris populations.
Indigenous_Cultivar_Role makes the strongest claim regarding geographic origin, asserting that the cultivar arose through local domestication events rather than introduction and subsequent adaptation. Modern molecular genetics — particularly parentage analysis, population structure studies, and archaeobotanical evidence — have revised many traditional origin narratives: cultivars long assumed indigenous to specific regions have been revealed as introductions (e.g., Syrah long claimed as Persian in origin was shown to be a cross of two obscure southeastern French cultivars, Dureza × Mondeuse Blanche). This role should be assigned conservatively based on current genetic evidence.
einheimische Rebsorten-Rolle
gebietsursprüngliche Sorten-Rolle
indigene Sorten-Rolle
indigenous variety role
locally originated cultivar role
native cultivar role
Myles S et al. (2011) Genetic structure and domestication history of the grape, Proceedings of the National Academy of Sciences 108: 3530-3535 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Grassi F et al. (2003) Evidence of a secondary grapevine domestication centre in the Iberian Peninsula, Theoretical and Applied Genetics 107: 1315-1320
Indigenous Cultivar Role
Chardonnay in Champagne shows individual cluster definition during late May Year N+1 at late BBCH 57 over 1-2 days, with 18-25 primary branch units showing clear 2.5-3.5 mm spacing, flower buds enlarged to 3.0-3.5 mm creating visible flower mass within each unit, establishing recognizable cluster architecture in this moderately-to-highly compound early-bursting cultivar under cool maritime conditions of 16-19°C temperature preparing for flowering early June
Individual cluster definition represents the culminating organizational phase where separated and spaced flower groups achieve distinct perceptual identity as recognizable cluster units within overall inflorescence architecture, establishing clear visual and functional demarcation between adjacent flower groups preparing for coordinated anthesis within each cluster unit. This final group-level organizational stage occurs during late May in temperate regions Year N+1 completing cluster separation process, typically spanning 1-2 days as final spatial adjustments and flower development create definitive cluster unit recognition. Definition achievement involves completion of spatial separation with final inter-cluster spacing established, flower bud enlargement to 3-4 mm creating visible flower mass within each cluster unit, and architectural stabilization with growth rates declining as structure approaches flowering readiness. Cluster unit recognition emerges through combined effects of spatial separation creating inter-group gaps, flower mass creating visual density within groups, and structural definition with branch units clearly distinguishable as discrete organizational elements. Visual demarcation between adjacent clusters shows clear spacing typically 2-5 mm in well-separated architectures enabling easy perceptual distinction, while compact architectures show minimal 1-2 mm spacing creating somewhat ambiguous boundaries though structural branch units remain identifiable. Functional cluster units represent coordinated developmental domains with flowers within each unit showing similar developmental stage supporting synchronized anthesis, creating temporal organization with individual cluster units flowering coordinately though entire inflorescence may show slight developmental gradient. Microenvironmental characteristics within defined cluster units show localized conditions including light exposure, air circulation, and temperature that will affect flowering success and subsequent fruit development, with well-defined separated clusters showing favorable microenvironment supporting uniform development. Developmental uniformity within cluster units improves during definition phase as flowers approach anthesis readiness, with good within-unit synchrony supporting coordinated flowering and uniform fruit set creating consistent berry development within cluster. Flower bud maturation during definition phase approaches completion with flower organs fully differentiated within enclosing calyptra, anthers mature with viable pollen, pistils developed with receptive stigmas, and nectaries functional preparing for nectar secretion, creating flowering readiness. Architectural framework established through definition determines cluster structure that will persist through anthesis, fruit set, berry development, and harvest, affecting disease pressure, fruit quality characteristics, and mechanical properties influencing handling and processing. Branch structural maturation during definition includes progressive wall thickening and beginning lignification particularly in rachis and primary branch bases establishing mechanical support adequate for flower weight and preparing for substantially greater berry loads. Vascular system completion during definition establishes transport capacity adequate for flowering water demands of 2-4 mg per second per cluster, with capacity prepared for increases to 8-15 mg per second during peak berry expansion supporting substantial water transport requirements. Metabolic transition during definition shows declining respiration from growth levels toward maintenance and flowering function levels, with energy allocation shifting from growth processes toward reproductive metabolism supporting anthesis and fertilization. Resource allocation during definition emphasizes inflorescence demands preparing for flowering with adequate carbohydrate reserves ensuring energy availability for anthesis processes, nitrogen concentrated in developing reproductive organs supporting protein synthesis, and mineral nutrients particularly boron positioned for supporting pollen germination and tube growth. Hormonal status during definition shows declining growth promoters as elongation completes, sustained auxin from developing flowers coordinating final maturation, possible increasing ethylene involved in senescence of calyptra cap preparing for cap fall, and other hormones coordinating transition toward flowering. Environmental conditions during definition include temperature approaching optimal flowering range with 18-24°C supporting final maturation and preparing for anthesis occurring optimally at 20-30°C, adequate moisture maintaining turgor supporting tissue function, and favorable weather supporting approaching critical flowering period. Disease pressure during definition intensifies as susceptible flowering tissues fully expose with powdery mildew infection risk high requiring final protective fungicide applications before flowering, botrytis potentially establishing infections that will remain latent through development expressing during ripening, and downy mildew threatening in wet conditions. Pest monitoring during definition includes grape berry moth adults present potentially ovipositing on inflorescences, leafhoppers reaching populations requiring management in some regions, and thrips colonization affecting flower quality potentially impacting pollination and fruit set. The process represents fourth and final flower cluster separation aspect completing group-level organization before individual flower separation, establishing recognizable cluster units that will function as developmental domains. Duration typically spans 1-2 days representing brief culminating phase as definition emerges from completed spacing and maturation, with rapid transition toward individual flower separation. Cultivar variation appears in cluster unit clarity with loose-clustered varieties showing well-defined clearly separated units while tight-clustered types show less distinct units with ambiguous boundaries, affecting management assessment of flowering progress and disease intervention requirements. Regional timing shows individual cluster definition during late May in warm climates, late May-early June in temperate regions, and early June in cool areas, occurring immediately before individual flower separation and anthesis. Viticultural significance includes establishing recognizable organizational units facilitating cluster assessment and management, completing architectural framework affecting disease pressure and fruit quality throughout season, preparing cluster for critical flowering period with adequate exposure and appropriate microenvironment, and marking transition point from structural development to reproductive function. Research applications include analyzing perceptual and functional aspects of cluster organization, evaluating architectural effects on within-cluster developmental uniformity, quantifying relationships between cluster definition and flowering success, and assessing implications of cluster architecture for disease management and fruit quality outcomes.
Individual_Cluster_Definition is fourth and final Flower_Cluster_Separation subphase, distinguished by achieving clear recognizable cluster unit identity. Follows and completes Primary, Secondary separation and Spacing Development. Transitions to Individual_Flower_Separation as flowers individualize within cluster units. Represents culmination of group-level organization before flower-level development. Marks readiness for flowering processes.
Blütengruppenverfestigung
Gescheinseinheitserkennung
Teilblütenstandsabgrenzung
cluster unit recognition
definitive cluster organization
flower group consolidation
individuelle Cluster-Definition
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 350-355 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Individual Cluster Definition
Merlot in Bordeaux shows individual flower separation during late May-early June Year N+1 at BBCH 57-59 over 4-5 days, with pedicels elongating from 0.8-1.2 mm to 2.5-4.0 mm creating flower spacing of 4-7 mm, flower buds enlarging to 4.0-4.5 mm with loosening caps preparing for anthesis, under oceanic conditions of 17-22°C temperature, adequate moisture, approaching optimal flowering conditions typical of this moderately spaced mid-season cultivar completing pre-anthesis preparation
Individual flower separation represents the final pre-anthesis developmental phase where compact flower buds separate from tight clusters to individually distinguishable flowers through pedicel elongation and flower spacing, establishing independent flower positions enabling successful pollination and fruit set, corresponding to BBCH 57-59 stages immediately preceding flowering. This critical preparatory phase occurs during late May-early June in temperate regions Year N+1 following cluster separation, typically spanning 3-5 days as coordinated pedicel elongation and flower bud maturation transform compact undifferentiated flower masses into arrays of individual flowers ready for anthesis. Separation mechanism involves pedicel elongation extending flower attachment stalks from initial 0.5-1.5 mm to final 2-5 mm creating spatial projection, flower bud enlargement from 3-4 mm to 4-5 mm as organs complete maturation, and slight branch elongation distributing flower attachment points along branch length. Pedicel function includes structural support maintaining flower position and orientation, vascular connection providing water and nutrient delivery to flower and developing fruit, and mechanical flexibility allowing flower movement facilitating pollination and later accommodating berry expansion. Individual flower visibility emerges progressively during separation as pedicels extend projecting flowers away from compact positions, creating recognizable individual flower units arranged along branches rather than undifferentiated masses. Flower spacing completion establishes final pre-anthesis flower arrangement with inter-flower distances typically 3-8 mm depending on cluster architecture, supporting adequate separation for air circulation around flowers, pollen transfer between flowers, and accommodation of expanding berries after fruit set. Flower bud maturation during separation approaches completion with all flower organs fully developed and functional, calyptra cap loosening in preparation for detachment, anthers positioned for dehiscence and pollen release, pistils mature with receptive stigmas, and nectaries active producing nectar attracting potential insect pollinators. Developmental uniformity within cluster increases during individual separation as flowers achieve similar readiness states, supporting synchronized anthesis with coordinated cap fall and pollen release across cluster promoting effective pollination and uniform fruit set. Pedicel elongation mechanism involves cell division in pedicel meristem zones adding cells to pedicel length particularly during early separation, and cell expansion throughout pedicel tissues increasing cell dimensions through turgor-driven growth, with elongation rate typically 0.3-0.8 mm per day at optimal temperatures. Vascular differentiation in elongating pedicels establishes functional xylem and phloem connecting flower to branch vascular system, enabling water delivery supporting flower turgor and function, nutrient import supporting metabolism and organ growth, and preparing for post-fertilization assimilate import during berry development. Mechanical properties of pedicels show flexible structure from thin-walled tissues allowing flower movement and positioning adjustments, with progressive strengthening as walls thicken though major lignification occurs later during fruit development when mechanical demands increase. Flower positioning during separation establishes spatial orientation affecting pollination efficiency with flowers typically oriented upward or laterally exposing stigmas for pollen deposition, with positioning also affecting post-anthesis berry orientation and cluster geometry. Metabolic activity during individual separation maintains elevated levels supporting continued development with respiration 8-12 fold above mature tissue levels, though beginning to shift from growth to reproductive function supporting approaching anthesis. Resource demands during separation emphasize carbohydrate requirements for continued growth and preparing energy reserves for anthesis processes, nitrogen for final protein synthesis in reproductive organs, and minerals particularly boron critical for pollen germination and tube growth. Hormonal regulation during separation involves auxin from developing flowers affecting pedicel elongation and positioning, gibberellins potentially affecting elongation though concentrations declining toward flowering, and ethylene possibly involved in cap loosening preparing for cap fall. Environmental conditions during separation critically affect development with temperature approaching flowering optimum with 18-26°C supporting final separation and maturation, adequate moisture maintaining turgor supporting elongation and flower function, and favorable weather conditions supporting approaching anthesis. Light exposure increases during separation as flowers extend from cluster interior toward exterior, with adequate light supporting robust flower development though excessive exposure potentially causing stress effects. Water stress during separation impairs development through effects on turgor limiting pedicel elongation and potentially affecting flower viability, requiring adequate soil moisture and favorable evaporative conditions minimizing stress. Disease pressure during separation focuses on powdery mildew with fully exposed flowers highly susceptible requiring final protective applications before anthesis, with infection during flowering potentially causing berry infection affecting fruit development. Developmental coordination between separation and approaching anthesis shows tight coupling with separation completing as flowers achieve anthesis readiness, creating seamless transition from structural preparation to reproductive function. Within-flower organ maturation proceeds during separation with final anther dehiscence preparation including connective tissue development enabling anther opening, pollen maturation completing with viable pollen grains containing developed gametophytes, pistil maturation with ovules prepared for fertilization containing mature megagametophytes, and stigma surface development establishing receptivity for pollen germination. The process shows four distinguishable developmental aspects progressing from flower bud separation beginning individualization, through pedicel elongation creating spatial projection, individual flower visibility establishing recognition, to flower spacing completion achieving final pre-anthesis arrangement. Duration typically spans 3-5 days depending on temperature and cultivar characteristics, with warm conditions promoting rapid completion while cool conditions extend period. Cultivar variation appears in final pedicel length affecting flower spacing with long-pedicelled varieties showing 4-6 mm creating well-separated flowers facilitating air circulation while short-pedicelled types showing 2-3 mm maintaining compact flower arrangement, separation rate affecting timing toward flowering, and flower size at separation affecting visual impact. Regional timing shows individual flower separation during late May-early June in warm climates, early June in temperate regions, and early-mid June in cool areas, occurring immediately before anthesis with separation completing as flowering initiates. Viticultural significance includes establishing final flower arrangement affecting pollination efficiency and fruit set uniformity, completing pre-anthesis development preparing flowers for critical reproductive processes, creating favorable microenvironment for flowering with adequate air circulation reducing disease pressure, and marking final preparation stage before flowering requiring optimal management and conditions. Research applications include analyzing mechanisms regulating pedicel elongation and flower spacing, investigating environmental effects on separation affecting flowering readiness, quantifying relationships between flower spacing and pollination efficiency, evaluating coordination between separation completion and anthesis initiation, and assessing genetic variation in flower separation characteristics affecting fruit set and yield.
Individual_Flower_Separation represents final pre-anthesis phase within Inflorescence_Development focusing on flower-level individualization. Follows Flower_Cluster_Separation. Subdivided into Flower_Bud_Separation, Pedicel_Elongation, Individual_Flower_Visibility, and Flower_Spacing_Completion. Transitions directly to Flowering phase as anthesis initiates. Critical preparatory phase establishing conditions for successful pollination and fruit set. Completion marks flowering readiness.
Blütenstielentwicklung
Einzelblütenseparation
flower bud separation
individual flower spacing
individuelle Blütenaufteilung
pedicel development
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 352-360 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Individual Flower Separation
Pinot Noir in Oregon shows individual flower visibility during early June Year N+1 at late BBCH 59 over 1-2 days, with 200-350 individual flowers per cluster clearly distinguishable on pedicels of 2.5-3.5 mm length, flower buds 4.0-4.5 mm showing slight cap loosening in earliest flowers, under cool climate conditions of 17-21°C temperature approaching flowering, enabling accurate flower counting for yield prediction in this moderately compact early-to-mid season cultivar
Individual flower visibility represents the perceptual achievement phase where elongated pedicels project flowers sufficiently that individual flower units become clearly distinguishable as separate entities, enabling confident flower identification and counting for yield prediction and management assessment. This important recognition milestone occurs during early June in temperate regions Year N+1 as pedicel elongation nears completion, typically representing 1-2 day period when individual flowers achieve definitive visual distinction corresponding to late BBCH 57-59 stages immediately preceding anthesis. Visibility achievement involves completion of substantial pedicel elongation projecting flowers 2-4 mm from branch attachment creating spatial separation, flower bud enlargement to 4-5 mm final pre-anthesis size establishing visible flower mass, and flower positioning with most flowers oriented upward or laterally creating recognizable spatial array. Visual distinction criteria include clear inter-flower spacing enabling perception of individual flowers rather than undifferentiated masses, recognizable flower bud morphology with calyptra cap shape distinguishable, and spatial positioning allowing viewing of individual flowers from multiple angles confirming independent status. Flower countability emerges during visibility phase as individual flowers can be reliably enumerated providing accurate assessment of potential berry number, supporting yield prediction calculations and crop adjustment decisions based on desired crop load. Cluster appearance transitions from compressed flower masses at separation initiation to clearly defined arrays of individual recognizable flowers at visibility achievement, revealing final pre-anthesis cluster architecture including branching pattern, flower density, and spatial organization. Flower bud characteristics at visibility show 4-5 mm green calyptra caps with smooth surface, fused petal seams potentially visible indicating enclosed corolla, slight cap loosening detectable in some flowers approaching earliest anthesis, and overall plump appearance indicating mature flower organs within. Flower positioning patterns show majority of flowers oriented upward 30-60 degrees above horizontal or laterally near horizontal, with positioning affecting stigma exposure for pollen deposition and reflecting phototropic, gravitropic, and mechanical influences. Pedicel characteristics at visibility include final or near-final length of 2-5 mm depending on cultivar, green herbaceous appearance with smooth surface, flexible structure allowing flower movement in wind, and functional vascular connection visible as slight ridge along pedicel length. Spatial flower distribution along branches shows characteristic pattern with flowers distributed according to branching architecture, inter-flower spacing reflecting pedicel length and branch elongation, and overall density determined by flower number per unit branch length. Developmental uniformity within cluster at visibility shows most flowers at similar maturation stage with synchronized cap loosening and approaching anthesis, supporting coordinated flowering across cluster promoting uniform pollination and fruit set. Microenvironmental characteristics within visible flower arrays include air circulation patterns with adequate flower spacing promoting rapid drying after rain or dew reducing disease infection risk, light exposure with flowers emerging from leaf shade toward better illumination, and temperature distribution with exposed flowers experiencing warmer conditions potentially advancing development. Flower organ status at visibility shows complete maturation with anthers containing viable pollen, pistils with receptive stigmas prepared for pollen germination, nectaries functional and beginning nectar production, and all structures positioned for imminent anthesis. Resource status at visibility shows accumulated carbohydrate reserves in flowers providing energy for anthesis processes, nitrogen concentrated in reproductive organs supporting continued protein synthesis, and critical minerals particularly boron positioned for pollen germination and tube growth. Hormonal environment at visibility involves declining growth promoters as elongation completes, sustained auxin from flowers, increasing ethylene potentially involved in cap abscission triggering, and other hormones coordinating transition from development to reproductive function. Environmental conditions at visibility approach optimal flowering requirements with temperature typically 19-26°C supporting final maturation and approaching anthesis optimum 22-30°C, adequate moisture maintaining flower turgor and function, and favorable weather protecting delicate structures and supporting approaching pollination. Disease assessment at visibility enables evaluation of powdery mildew infection presence with visible mycelial growth indicating infection requiring immediate intervention, though fully protective fungicide applications must precede visibility due to application restrictions during flowering. Pest presence at visibility may include grape berry moth larvae on inflorescences potentially affecting flower survival, leafhoppers feeding causing stippling damage, and thrips affecting flower quality potentially impacting pollination efficiency. Developmental progression from visibility proceeds immediately toward anthesis with cap fall initiating within 1-2 days in earliest flowers, creating seamless transition from structural preparation to reproductive function. The process represents third of four individual flower separation aspects marking achievement of clear flower recognition, following bud separation and pedicel elongation and preceding final spacing completion. Duration represents brief 1-2 day period when visibility definitively achieves though flowers have been progressively becoming more distinct during elongation. Cultivar variation appears in visibility clarity with loose-clustered well-spaced varieties showing obvious early visibility while tight-clustered compact types show less distinct visibility requiring careful inspection, affecting assessment ease and management timing precision. Regional timing shows individual flower visibility during late May-early June in warm climates, early June in temperate regions, and early-mid June in cool areas, occurring 1-2 days before anthesis initiation with visibility representing flowering readiness indicator. Viticultural significance includes enabling accurate flower counting for yield prediction, confirming cluster architecture affecting disease risk and quality potential, indicating imminent flowering requiring final preparations and optimal conditions, providing assessment reference for flowering uniformity predictions, and marking transition point from developmental to reproductive management focus. Research applications include documenting precise timing relationships between visibility achievement and anthesis initiation, evaluating environmental effects on visibility clarity affecting assessment reliability, quantifying flower counting accuracy as function of architectural clarity, and developing automated computer vision systems for flower counting leveraging visibility characteristics.
Individual_Flower_Visibility is third of four Individual_Flower_Separation subphases, distinguished by achievement of clear perceptual distinction of individual flowers. Follows Pedicel_Elongation as recognition milestone. Overlaps with or transitions to Flower_Spacing_Completion as final arrangements establish. Represents practical assessment stage for flower counting and yield prediction. Marks flowering readiness with anthesis imminent.
Blütenzählstadium
Einzelblütenerkennbarkeit
deutliche Blütenerkennung
distinct flower appearance
einzelne Blütensichtbarkeit
flower counting stage
individual flower recognition
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 358-360 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Individual Flower Visibility
Systematic characterization of Induced_Defense_Disposition at INRAE Colmar compared defense induction kinetics across 25 cultivars following Plasmopara viticola inoculation. Leaf discs were inoculated with sporangia suspension and harvested at 0, 6, 12, 24, 48, and 72 hours post-inoculation for gene expression analysis (qRT-PCR of defense marker genes), metabolite profiling (stilbenes by HPLC), and histochemical staining (callose, H₂O₂, cell death). Cultivars varied dramatically in Induced_Defense_Disposition parameters. The PIWI cultivar 'Regent' exhibited rapid, strong induction with stilbene synthase (STS) expression increasing 85-fold by 12 hours, resveratrol accumulating to 45 μg/g by 24 hours, and visible hypersensitive response (HR) cell death by 48 hours limiting pathogen spread. Susceptible 'Müller-Thurgau' showed delayed, weak induction with STS expression only 8-fold elevated at 24 hours, resveratrol reaching 6 μg/g at 48 hours, and no detectable HR. Intermediate cultivar 'Riesling' displayed moderate induction kinetics (25-fold STS at 18 hours, 18 μg/g resveratrol at 36 hours). Principal component analysis revealed that induction speed (time to 50% maximum response) and magnitude (maximum fold-change) were partially independent, with some cultivars showing fast but weak responses and others slow but ultimately strong accumulation. These findings demonstrate that Induced_Defense_Disposition comprises multiple parameters amenable to independent selection.
An Induced_Defense_Disposition is a Defense_Mechanism_Disposition that inheres in a Vitis vinifera plant and is realized through defense responses that are activated upon recognition of pathogen or pest attack, involving de novo synthesis of defensive compounds, activation of defense-related genes, and deployment of cellular defense mechanisms not present or active in unchallenged plants. This disposition represents the dynamic, responsive component of plant defense, complementing constitutive defenses by enabling targeted, amplified responses to specific threats while avoiding the metabolic costs of maintaining high defense levels continuously. Induced defenses in grapevine encompass the hypersensitive response (rapid programmed cell death at infection sites), phytoalexin production (particularly stilbenes including resveratrol and viniferins), pathogenesis-related protein accumulation (chitinases, glucanases, thaumatin-like proteins, and others), cell wall reinforcement (callose deposition, lignification, cross-linking), systemic acquired resistance (SAR) providing whole-plant protection following local infection, and volatile emission signaling to neighboring plants and recruiting natural enemies. The speed, magnitude, and specificity of induced responses vary substantially among grapevine genotypes, with resistant cultivars typically showing faster, stronger, and more sustained induction compared to susceptible cultivars challenged with the same pathogen. Understanding Induced_Defense_Disposition provides mechanistic insight into resistance differences among cultivars and enables breeding strategies targeting enhanced inducibility rather than elevated constitutive defense, potentially avoiding fitness costs associated with continuous high defense investment.
Induced_Defense_Disposition is_a Defense_Mechanism_Disposition. This class encompasses all defense responses activated upon pathogen/pest attack, contrasting with Constitutive_Defense_Disposition representing preformed defenses. Child classes include Hypersensitive_Response_Disposition (cell death-based defense), Phytoalexin_Production_Disposition (induced antimicrobial metabolites), and Pathogenesis_Related_Protein_Disposition (induced defense proteins). Induced and constitutive defenses typically act synergistically, with constitutive barriers slowing initial infection while induced responses mount.
Aktivierbare-Abwehr
Dynamische-Abwehr-Kapazität
Induzierte-Verteidigungs-Disposition
activated defense response
dynamic defense disposition
inducible defense capacity
Pieterse CMJ et al. (2012) Hormonal modulation of plant immunity, Annual Review of Cell and Developmental Biology 28:489-521 Jones JDG, Dangl JL (2006) The plant immune system, Nature 444:323-329 Hammerschmidt R (1999) Phytoalexins: what have we learned after 60 years?, Annual Review of Phytopathology 37:285-306
Induced Defense Disposition
Field trials at Agroscope Changins, Switzerland, evaluated Induced_Downy_Mildew_Resistance_Disposition in 'Chasselas' vines treated with the elicitor product Bastid® (laminarin, a β-1,3-glucan from brown algae). Weekly applications beginning at BBCH 15 (5 leaves unfolded) primed vine defenses, as confirmed by elevated basal expression of PR-1 and STS (stilbene synthase) genes in treated versus control vines. Following natural P. viticola infection during a rainy June period, elicitor-treated vines exhibited 38% lower leaf infection severity (22% versus 35%) and 45% lower cluster infection (12% versus 22%) compared to water-treated controls. Microscopic examination of inoculated leaf discs revealed that sporangiophore emergence was delayed by 18-24 hours in elicitor-treated tissue, with 40% fewer sporangiophores per lesion. While insufficient as a standalone strategy, integrating elicitor applications with reduced fungicide programs achieved equivalent disease control to conventional programs with 30% fewer synthetic fungicide applications, demonstrating the practical value of induced resistance for sustainable viticulture.
An Induced_Downy_Mildew_Resistance_Disposition is a Downy_Mildew_Resistance_Disposition that inheres in a Vitis plant and is realized through enhanced defense capacity acquired following prior exposure to resistance-inducing stimuli, including pathogen challenge, beneficial microorganisms, or chemical elicitors. This disposition represents a primed state of the plant immune system characterized by faster, stronger, and more sustained activation of defense responses upon subsequent pathogen encounter. Induced resistance in grapevine against P. viticola can be triggered through multiple pathways, including systemic acquired resistance (SAR) following localized pathogen infection, induced systemic resistance (ISR) elicited by plant growth-promoting rhizobacteria or mycorrhizal fungi, and chemically induced resistance activated by synthetic or natural elicitor compounds. Molecular signatures of the induced resistant state include elevated expression of defense-related genes (PR proteins, stilbene synthases), accumulation of signaling molecules (salicylic acid, jasmonic acid), epigenetic modifications facilitating rapid transcriptional responses, and enhanced sensitivity of pattern recognition receptors. Practical applications of induced resistance in viticulture include phosphonate fungicides (which induce host defenses in addition to direct antimicrobial effects), commercial elicitor products based on β-glucans, chitosan, or plant extracts, and biological control agents that colonize roots or foliar surfaces and prime systemic defenses. While induced resistance alone rarely provides complete protection, it can significantly enhance the efficacy of other resistance mechanisms and reduce fungicide requirements.
Induced_Downy_Mildew_Resistance_Disposition is_a Downy_Mildew_Resistance_Disposition. This class encompasses all forms of enhanced resistance acquired through prior stimulation, distinguished from constitutive genetic resistance (Rpv-mediated, quantitative) and developmental resistance (ontogenic). The induced state is transient and requires repeated elicitation to maintain protection throughout the growing season. Induced resistance provides complementary protection that can be integrated with genetic resistance and conventional disease management.
Elicitor-induzierte Peronospora-Resistenz
Induzierte Peronospora-Resistenz-Disposition
SAR-mediated DM resistance
Systemisch erworbene Peronospora-Resistenz
elicitor-induced DM resistance
primed Plasmopara resistance
Harm A et al. (2011) Evaluation of chemical and natural resistance inducers against downy mildew (Plasmopara viticola) in grapevine, American Journal of Enology and Viticulture 62:184-192 Perazzolli M et al. (2011) Phosphate-solubilizing Pseudomonas rhizobacteria as potential inducers of systemic resistance against grapevine downy mildew, IOBC/WPRS Bulletin 67:245-251 Walters DR et al. (2013) Controlling crop diseases using induced resistance: challenges for the future, Journal of Experimental Botany 64:1263-1280
Induced Downy Mildew Resistance Disposition
Evaluation of Induced_Systemic_Resistance_Disposition at the University of Reims Champagne-Ardenne assessed grapevine responsiveness to beneficial rhizobacteria across 12 cultivars using controlled rhizosphere inoculation and subsequent foliar pathogen challenge. Rooted cuttings were inoculated with Pseudomonas fluorescens CHA0 or Bacillus subtilis GB03 at planting, allowed 4 weeks for root colonization, then challenge-inoculated on leaves with either Plasmopara viticola or Botrytis cinerea. Cultivars showed differential Induced_Systemic_Resistance_Disposition to bacterial priming. 'Chardonnay' roots colonized by P. fluorescens showed strong ISR against both pathogens, with 58% reduction in P. viticola sporulation and 45% reduction in B. cinerea lesion size relative to non-colonized controls. Molecular analysis confirmed JA pathway activation, with 6-fold elevated expression of the JA marker gene VvLOX9 in roots and 3-fold elevation in leaves of colonized plants. 'Cabernet Sauvignon' showed weaker Induced_Systemic_Resistance_Disposition (32% and 22% reductions respectively) despite equivalent root colonization levels, suggesting genotypic variation in ISR signaling responsiveness. Interestingly, ISR and SAR showed additive effects when combined, with plants receiving both rhizobacterial priming and localized P. viticola infection exhibiting 78% protection against challenge inoculation, exceeding either treatment alone. AMF colonization (Rhizophagus irregularis) similarly induced ISR, with 'Pinot Noir' showing 52% P. viticola reduction in mycorrhizal versus non-mycorrhizal plants. These findings demonstrate that Induced_Systemic_Resistance_Disposition provides a distinct, complementary defense layer exploitable through beneficial microbiome management.
An Induced_Systemic_Resistance_Disposition is an Induced_Defense_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to establish whole-plant enhanced resistance following colonization by beneficial root-associated microorganisms, particularly plant growth-promoting rhizobacteria (PGPR) and certain mycorrhizal fungi, providing protection against foliar and root pathogens through jasmonate/ethylene-mediated defense priming. This disposition represents a distinct systemic defense pathway from SAR, triggered not by pathogen attack but by mutualistic microbial interactions in the rhizosphere, and mediated primarily by jasmonic acid (JA) and ethylene (ET) signaling rather than salicylic acid. Induced systemic resistance (ISR) primes plants for enhanced expression of JA/ET-responsive defense genes upon pathogen challenge, without constitutively activating defenses in the absence of attack, thereby providing protection without the metabolic costs of continuous defense expression. Beneficial microbes triggering ISR in grapevine include Pseudomonas fluorescens, Bacillus subtilis, Trichoderma species, and arbuscular mycorrhizal fungi (AMF), which colonize roots and produce elicitors including lipopolysaccharides, flagellin, siderophores, and volatile organic compounds that are perceived by plant receptors and initiate systemic signaling. The resulting primed state enhances callose deposition, phytoalexin accumulation, and oxidative burst responses upon subsequent pathogen challenge in both root and shoot tissues. Induced_Systemic_Resistance_Disposition is particularly relevant for integrated pest management approaches that leverage beneficial soil microbiomes, and varies among grapevine genotypes in responsiveness to ISR-triggering microbes.
Induced_Systemic_Resistance_Disposition is_a Induced_Defense_Disposition. ISR is triggered by beneficial root microbes (PGPR, mycorrhizae) and mediated by JA/ET signaling, contrasting with SAR triggered by pathogens and mediated by SA. ISR is particularly effective against necrotrophic pathogens and herbivores (JA-responsive defenses) while SAR favors protection against biotrophs (SA-responsive). The two pathways can act additively or synergistically when both are activated. ISR provides a foundation for biocontrol approaches using beneficial microorganisms as plant protection agents, with cultivar responsiveness to ISR-triggering microbes representing a selectable trait for integrated management systems.
ISR capacity
ISR-Kapazität
Induzierte-systemische-Resistenz-Disposition
Rhizobakterien-induzierte-Resistenz
jasmonate-mediated systemic resistance
rhizobacteria-induced resistance
Pieterse CMJ et al. (2014) Induced systemic resistance by beneficial microbes, Annual Review of Phytopathology 52:347-375 Verhagen BWM et al. (2004) The transcriptome of rhizobacteria-induced systemic resistance in Arabidopsis, Molecular Plant-Microbe Interactions 17:895-908 Compant S et al. (2005) Use of plant growth-promoting bacteria for biocontrol of plant diseases: principles, mechanisms of action, and future prospects, Applied and Environmental Microbiology 71:4951-4959
Induced Systemic Resistance Disposition
Infection incidence quality monitoring for downy mildew (Plasmopara viticola) in Mosel Valley Riesling tracked disease spread across the vineyard to optimize fungicide timing and coverage decisions. Weekly assessment of 100 randomly selected leaves per block quantified infection incidence quality as the percentage of leaves showing characteristic oil-spot symptoms. Early-season infection incidence quality remained below 5% through June despite favorable weather, indicating successful prophylactic fungicide protection of the expanding canopy. A significant rain event in early July followed by warm temperatures created ideal infection conditions, with infection incidence quality increasing from 8% to 42% within two weeks despite continued fungicide applications, indicating that spray coverage had been inadequate during the critical infection period. The rapid infection incidence quality increase triggered emergency intervention with systemic curative fungicides applied at shortened intervals. Cluster infection incidence quality assessment at BBCH 79 (majority of berries touching) showed 28% of clusters affected, substantially below the 42% leaf incidence, attributed to preferential fungicide coverage of the more exposed fruit zone compared to interior canopy leaves. The infection incidence quality monitoring established that leaf incidence provided early warning of epidemic development before cluster infection became extensive, enabling protective response focused on the economically critical fruit.
Infection incidence quality is a disease severity quality that inheres in a grapevine population or organ set and represents the proportion of sampling units showing any detectable level of infection, expressed as the percentage of plants, shoots, leaves, clusters, or berries that are affected regardless of how severely each unit is infected, providing the measure of how widespread disease has become within the assessed population. This quality captures the distributional dimension of disease assessment, recognizing that the proportion of units affected determines the spatial extent of pathogen presence and provides information about epidemic progress that is distinct from and complementary to the intensity of infection within affected units. The ontological significance of infection incidence quality lies in its role indicating the scope of disease spread and the potential for continued epidemic development, with high incidence suggesting widespread inoculum distribution and favorable conditions for infection that may result in continued disease increase, while low incidence may indicate localized infection foci that could be targeted for intensive management or may simply reflect early epidemic stages before extensive spread. The biological basis of infection incidence encompasses the epidemiological processes of spore production, dispersal, and successful infection that determine whether individual sampling units become infected, with incidence reflecting the cumulative outcome of these processes integrated over the assessment period. The sampling protocol for incidence assessment must define the sampling unit (plant, shoot, leaf, cluster, berry) appropriate for the disease and management question, with larger units showing higher incidence than smaller units for the same overall disease level due to the aggregation of infection. The temporal dynamics of incidence during epidemics typically show sigmoidal increase from initial low levels through rapid expansion to plateau near 100% in severe epidemics, with the rate of incidence increase providing information about epidemic velocity useful for management timing decisions.
Infection_Incidence_Quality is_a Disease_Severity_Quality, representing the proportion of sampling units showing any infection regardless of severity within units. This class addresses disease distribution rather than intensity. Sampling unit definition (leaf, cluster, berry) affects absolute incidence values for equivalent disease levels. The sigmoidal temporal dynamic reflects epidemic progression. Early-warning value of incidence monitoring enables proactive management. Leaf versus cluster incidence may differ due to differential exposure and protection. Distinguished from Infection_Intensity_Quality which measures severity within affected units.
Anteil befallener Einheiten
Befallshäufigkeit
Infektionsfrequenz
Infektionsinzidenz
affected unit percentage
disease incidence
infection frequency
proportion infected
Seem RC (1984) Disease incidence and severity relationships. Annual Review of Phytopathology 22:133-150 Madden LV et al. (2007) The Study of Plant Disease Epidemics. APS Press Kennelly MM et al. (2005) Seasonal development of ontogenic resistance to downy mildew in grape berries and rachises. Phytopathology 95:1445-1452
Infection Incidence Quality
Infection intensity quality assessment for Botrytis bunch rot (Botrytis cinerea) in Marlborough Sauvignon Blanc provided the basis for selective harvest decisions balancing disease management against fruit maturity optimization. Pre-harvest cluster sampling evaluated infection intensity quality on those clusters identified as infected during incidence screening. Visual intensity rating employed a 0-7 scale where 0 = no infection, 1 = 1-5% of berries affected, 3 = 6-25%, 5 = 26-50%, and 7 = greater than 50% berry involvement per cluster. The infection intensity quality distribution showed 45% of infected clusters at intensity 1 (trace infection), 30% at intensity 3 (moderate), 18% at intensity 5 (substantial), and 7% at intensity 7 (severe), enabling block-level mean intensity calculation of 2.4 on the infected cluster population. Harvest threshold protocols established that blocks with mean infection intensity quality above 3.0 required immediate harvest regardless of Brix level to prevent continued disease development, while blocks below 2.0 could await optimal ripeness with continued monitoring. The infection intensity quality assessment enabled precision harvest scheduling where high-intensity blocks were harvested at 21.5°Brix while low-intensity blocks achieved target 23.0°Brix without unacceptable quality loss, demonstrating the value of intensity quantification for disease-adjusted harvest decisions.
Infection intensity quality is a disease severity quality that inheres in infected grapevine organs and represents the degree or magnitude of disease development within those sampling units that are infected, measured as the proportion of tissue affected, the quantity of pathogen structures present, or the functional impairment caused by infection, providing the assessment of how severely infection has progressed within the affected population subset. This quality captures the severity dimension of disease within infected units, recognizing that units classified as infected may range from barely detectable trace infection to near-complete tissue colonization, with intensity measures quantifying this variation in ways that complement incidence for comprehensive disease assessment. The ontological significance of infection intensity quality lies in its role determining the actual damage and loss caused by disease, since intensity directly relates to tissue destruction, physiological impairment, and yield or quality reduction in ways that incidence alone cannot capture, with low-intensity infections potentially causing minimal impact while high-intensity infections may cause severe damage even when incidence is limited. The biological basis of infection intensity encompasses the processes of pathogen growth and reproduction within host tissues following successful infection, the host resistance mechanisms that may limit pathogen spread, and the environmental conditions that influence the rate of disease development within infected tissues. The assessment methods for infection intensity include visual rating scales that classify intensity into ordered categories, percentage estimates of tissue affected, counts of pathogen structures such as lesions or pustules per unit area, and quantitative molecular or biochemical measures of pathogen biomass or activity. The relationship between intensity and crop loss is typically more direct than for incidence, since intensity measures the actual tissue damage that causes yield reduction and quality compromise, though the intensity-loss relationship may still be non-linear with threshold effects and timing influences affecting the translation of observed intensity into economic impact.
Infection_Intensity_Quality is_a Disease_Severity_Quality, representing the severity of disease within infected sampling units. This class addresses damage magnitude rather than distribution. Visual rating scales (0-7, 0-100%) provide standardized intensity quantification. Mean intensity across infected units provides population-level metric. The more direct relationship to crop loss compared to incidence enables damage prediction. Harvest threshold protocols based on intensity enable disease-adjusted management. Distinguished from Infection_Incidence_Quality which measures distribution rather than severity.
Befallsintensität
Infektionsintensität
Kolonisierungsausmaß
Schwere innerhalb befallener Einheiten
colonization extent
disease intensity
infection severity level
within-unit severity
Seem RC (1984) Disease incidence and severity relationships. Annual Review of Phytopathology 22:133-150 Madden LV et al. (2007) The Study of Plant Disease Epidemics. APS Press Elmer PAG, Michailides TJ (2007) Epidemiology of Botrytis cinerea in orchard and vine crops. In: Botrytis: Biology, Pathology and Control. Springer, pp. 243-272
Infection Intensity Quality
The inferred parentage of Syrah was determined by Vouillamoz and Grando (2006) through SSR analysis at 50 microsatellite loci across a reference panel of 300 cultivars from southeastern France — Dureza (an obscure red cultivar from the Ardèche) and Mondeuse Blanche (a white cultivar from Savoie) were identified as the parents with a combined exclusion probability exceeding 99.99%, confirmed by the observation that Syrah's alleles at all 50 loci are consistent with Mendelian inheritance from the Dureza × Mondeuse Blanche cross. The maternal versus paternal roles were assigned through chloroplast haplotype analysis — Syrah shares chloroplast haplotype C with Mondeuse Blanche (but not with Dureza, which carries haplotype D), identifying Mondeuse Blanche as the seed (maternal) parent and Dureza as the pollen (paternal) parent, a result consistent with spontaneous hybridization in the northern Rhône Valley where both parent cultivars were historically cultivated.
A parentage information record based on molecular marker analysis — principally SSR exclusion testing and likelihood-based parentage assignment — that identifies the most probable parent cultivars of a variety for which no contemporaneous breeding documentation exists, specifying the marker system used, the number and identity of loci analyzed, the statistical confidence of the parentage assignment, and any remaining ambiguities regarding maternal versus paternal parent roles.
Inferred_Parentage captures the molecular evidence-based parentage assignments that have revolutionized grapevine genealogy since the late 1990s. The methodology relies on the principle that a true offspring must carry one allele from each parent at every analyzed locus — any locus at which neither allele of a candidate parent matches either allele of the putative offspring excludes that candidate as a parent (the exclusion principle): when two candidate parents are identified that together account for all alleles in the offspring across a sufficient number of loci (typically 20-50 SSR markers for confident assignment), the parentage is considered established with high statistical confidence. The Mendelian exclusion approach is definitive for identifying true parents but cannot determine the direction of the cross (which parent was maternal seed parent versus paternal pollen donor) — this directional information requires uniparentally inherited markers, specifically chloroplast DNA haplotypes that are maternally inherited in Vitis, enabling assignment of maternal and paternal roles when the parents carry different chloroplast haplotypes. The major limitation of inferred parentage is the requirement that both parent cultivars be present in the reference panel — if one parent is extinct, unsampled, or absent from germplasm collections, only half-parentage (one parent identified) can be determined.
abgeleitete Abstammung
genetically inferred parentage
marker-based parentage determination
marker-basierte Abstammungsbestimmung
molecular parentage inference
molekular erschlossene Elternschaft
Vouillamoz JF and Grando MS (2006) Genealogy of wine grape cultivars: Pinot is related to Syrah, Heredity 97: 102-110 Bowers JE et al. (1999) Historical genetics: the parentage of Chardonnay, Gamay, and other wine grapes of northeastern France, Science 285: 1562-1565 Lacombe T et al. (2013) Large-scale parentage analysis in an extended set of grapevine cultivars, Theoretical and Applied Genetics 126: 401-414 Cipriani G et al. (2010) The SSR-based molecular profile of 1005 grapevine accessions uncovers new synonymy and parentages, Theoretical and Applied Genetics 121: 1569-1585
Inferred Parentage
An inflorescence at node 3 (first inflorescence position) on a shoot of Vitis vinifera cv. Pinot Noir in Burgundy, France, measuring 14.5 cm length at full bloom representing typical dimensions for this cultivar, exhibiting characteristic compound paniculate architecture with central rachis bearing multiple orders of lateral branches, rachis diameter of 3.2 mm at base tapering to 1.8 mm distally, bearing approximately 380 individual flowers organized in clusters along the branching system, positioned opposite the third leaf on the shoot following the typical inflorescence-leaf arrangement, showing phenological development at BBCH stage 65 (full bloom) with approximately 50% of flowers showing separated petals (calyptras) and exposed reproductive organs, flowers exhibiting characteristic greenish-white color and faint fragrance, inflorescence developing from primordium initiated in the compound bud 14 months previously during June of the prior season, successful pollination expected to result in fruit set within 7-10 days with transformation from inflorescence to developing cluster, measured in mid-June representing peak flowering period for this region and cultivar, demonstrating typical primary inflorescence structure serving critical reproductive function.
The complete flowering structure in Vitis vinifera bearing all flowers and subsequently developing into the grape cluster, characterized by complex branched paniculate architecture derived from modified shoot structures, typical dimensions of 8-25 cm length at full development before fruit set varying with cultivar and environmental conditions, rachis (main axis) structure with multiple orders of branching creating hierarchical organization, developmental origin from inflorescence primordia initiated in latent buds during the previous growing season, position opposite leaves at nodes along the shoot following the two-leaf gap pattern (inflorescences at nodes 3, 5, 7, etc. counting from shoot base), typical occurrence of 1-3 inflorescences per shoot with basal inflorescences largest and most developed, bearing 50-2000+ individual flowers depending on cultivar and inflorescence size, and critical reproductive function supporting sexual reproduction through flower development, pollination, fertilization, and subsequent berry development following successful fruit set. The inflorescence represents the reproductive unit of the grapevine, developing through complex morphogenesis involving inflorescence primordium initiation during the previous season, dormancy overwinter within the bud, resumption of development at budbreak with rapid elongation and branching, flowering occurring typically 6-8 weeks after budbreak at BBCH stages 61-69, and transformation to the grape cluster following successful fertilization and fruit set at BBCH stage 71.
Inflorescence is distinguished from the subsequent cluster or bunch (developing and mature fruit-bearing structure) by developmental stage, referring specifically to the flowering structure before and during anthesis (blooming period). The inflorescence represents the critical reproductive structure where flower development, pollination, and fertilization occur, with successful completion of these processes resulting in fruit set and transformation to cluster. Architectural complexity shows cultivar variation from relatively simple weakly branched structures to highly complex multi-branched panicles, with branching pattern affecting flower number, cluster compactness after fruit set, and final cluster morphology. Inflorescence number per shoot varies with cultivar and position, with most cultivars producing 1-2 inflorescences per shoot on fruiting canes, though some highly fertile cultivars may produce 3 inflorescences at basal nodes. Size variation reflects genetic control and environmental conditions during both primordium initiation (previous season) and development (current season), with stress during initiation reducing flower number and stress during development affecting inflorescence elongation. The typical opposite-leaf position at alternate nodes creates the characteristic grapevine architecture with vegetative (leaf-only) and reproductive (leaf plus inflorescence) nodes. Phenological timing from budbreak to flowering shows cultivar specificity and strong temperature dependence, with degree-day accumulation determining flowering date affecting subsequent harvest timing. Management practices including shoot positioning, leaf removal, and flower cluster thinning manipulate inflorescence development and fruit set affecting final crop load and quality.
Infloreszenz
Traubenblüte
flower cluster
flowering structure
grape inflorescence
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 109-145 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 82-108 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 269-312.
Inflorescence
In Pinot Noir vineyards of Burgundy's Côte d'Or during BBCH 53-55 (inflorescence clearly visible), primary inflorescences typically display 12-18 primary branches averaging 2.5-4.5 cm length with 2-3 orders of sub-branching, creating moderately compact clusters. Conversely, Grenache in Châteauneuf-du-Pape exhibits 15-25 primary branches of 4-7 cm length with extensive tertiary branching, producing characteristically large, loose clusters of 300-800 berries. Gewürztraminer shows reduced branching (8-12 primary branches, 1.5-3 cm) with limited secondary subdivision, resulting in small, tight clusters highly susceptible to Botrytis bunch rot.
An inflorescence branch is a lateral appendage of the inflorescence rachis that bears flowers either directly or through further sub-branching, representing the fundamental architectural unit determining grapevine inflorescence complexity and ultimate flower number. This branching structure emerges from meristematic activity along the main rachis during inflorescence primordium development, typically initiated 8-12 months before anthesis during the previous growing season's bud formation. Inflorescence branches exhibit distinct hierarchical organization with primary branches arising directly from the main rachis, secondary branches from primary branches, and tertiary branches from secondary branches, creating the characteristic compound panicle structure of Vitis vinifera. Branch morphology varies substantially among cultivars, with length ranging from 0.5-8 cm, diameter from 0.3-2.5 mm, and flower-bearing capacity from 3-150 flowers per branch depending on branching order and position. The branching pattern follows acropetal development with basal branches typically longer and more extensively subdivided than apical branches, though this gradient varies with cultivar architecture type (compact versus loose inflorescences). Branch anatomy comprises a central vascular strand continuous with rachis vasculature, surrounded by parenchymatous cortex and epidermis bearing trichomes similar to rachis indumentum. Each branch subtends from a bract remnant at the rachis node, though bracts are typically caducous and absent at anthesis. Branch development is highly responsive to environmental conditions during the previous season's differentiation phase, with water stress, temperature extremes, and nutritional status affecting branch number, length, and subsequent flower formation. The spatial arrangement and vigor of inflorescence branches directly determines cluster architecture at berry stage, influencing disease susceptibility, spray penetration, and harvest quality through effects on berry compactness and air circulation.
Inflorescence_Branch represents the critical architectural element determining cluster morphology and should be distinguished from Shoot_Branch (vegetative lateral growth) and Root_Branch (underground lateral structures). The hierarchical classification (primary, secondary, tertiary) follows botanical convention for compound inflorescence description but reflects actual developmental sequence and vascular connectivity. Branch characteristics at BBCH 53-57 predict final cluster compactness and berry number with cultivar-specific allometric relationships. This class encompasses all branching orders collectively, with subclasses specifying hierarchical position and spatial location. Ampelographic identification relies heavily on branch number, length distribution, and subdivision pattern as stable cultivar characteristics under standard viticultural conditions.
Blütenstandsast
Infloreszenzast
Traubenzweig
flower cluster branch
inflorescence lateral branch
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 87-95 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development. In= Grapevine Molecular Physiology & Biotechnology, 2nd ed., pp. 159-185 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109.
Inflorescence Branch
The inflorescence description of Chardonnay documents medium-sized inflorescences (OIV 151:5, 120-160 mm length at full development pre-bloom) typically numbering 1.5-2.0 per shoot (OIV 152:2-3), inserted at nodes 3-5 opposite the leaf, with a compact pyramidal branching architecture featuring 4-6 primary branches (wings or shoulders) and 8-12 secondary branches — this inflorescence structure presages Chardonnay's characteristically small to medium, compact cylindrical clusters, as the final cluster morphology is largely determined by the inflorescence branching pattern established 12-14 months earlier during the previous season's bud initiation within the dormant compound bud.
An ampelographic description component that documents the morphological characteristics of the grapevine inflorescence — the branched flower-bearing structure (compound raceme or thyrse) that develops opposite the leaf at nodes 3-6 of fruitful shoots — recording the size, number per shoot, insertion position, branching architecture, and pre-flowering morphology according to OIV descriptors 151-153 that characterize the reproductive potential and cluster architecture precursor of each cultivar.
Inflorescence_Description addresses the reproductive structure that determines the fundamental architecture of the grape cluster — cluster size, shape, compactness, and wing development are all direct consequences of the inflorescence branching pattern established during bud organogenesis in the preceding growing season and elaborated during the 4-6 week period from bud burst to full bloom. The inflorescence is a compound raceme (thyrse) consisting of a main axis (rachis) bearing lateral branches at each node in a spiral phyllotactic arrangement, with the degree of branching (primary, secondary, and tertiary branch orders) and the number of individual flower primordia per inflorescence (typically 200-1,500 in V. vinifera) varying substantially between cultivars: cultivars with large, highly branched inflorescences (e.g., Grenache, with 500-1,500 flowers per inflorescence) produce large, potentially compact clusters requiring thinning for quality production, while cultivars with small, sparsely branched inflorescences (e.g., Pinot Noir, with 200-400 flowers) produce small clusters less prone to compaction-related Botrytis but with lower yield potential per cluster. The inflorescence is homologous to the tendril — both develop from the same primordial meristem at each node, with the developmental fate (inflorescence versus tendril) determined by floral induction signals during bud formation.
Blütenstand-Beschreibung
Geschein-Charakterisierung
Infloreszenz-Morphologie
flower cluster morphology
inflorescence architecture description
pre-bloom inflorescence characterization
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Carmona MJ et al. (2008) A molecular genetic perspective of reproductive development in grapevine, Journal of Experimental Botany 59: 2579-2596 UPOV (2011) Guidelines for the Conduct of Tests for Distinctness, Uniformity and Stability — Vine (Vitis L.), TG/50/9
Inflorescence Description
Cabernet Sauvignon in Napa Valley shows inflorescence development spanning two seasons with primordium formation June-August Year N at BBCH 19-35 during active shoot growth establishing architectural framework with 20-35 primary branches per inflorescence under conditions of 22-30°C temperature, adequate light exposure, moderate water stress, followed by winter dormancy requiring 800-1000 chilling hours, and emergence May Year N+1 at BBCH 53-57 producing visible inflorescences that will flower early June, with Year N management critically affecting Year N+1 crop potential in this moderately fruitful late-season cultivar
Inflorescence development represents the complex reproductive organogenesis process spanning two growing seasons where inflorescence primordia initiate within latent buds during current season vegetative growth, undergo architectural elaboration establishing branching pattern and flower number potential, enter dormancy as structured but undifferentiated meristematic tissues, and resume development following bud burst producing visible inflorescences that will undergo flowering and fruit set. This fundamental reproductive process encompasses primordium formation during previous season typically May-August Year N establishing inflorescence architecture within developing buds, winter dormancy maintaining primordia in arrested state, emergence during bud burst April-May Year N+1 as inflorescences become externally visible, and continued development through flowering preparation. Inflorescence development determines crop potential for subsequent season with primordium initiation frequency, branching architecture, and flower number established during Year N affecting yield capacity in Year N+1, making environmental conditions and vine management during primordium formation critically important for subsequent productivity. The developmental program follows predictable architectural pattern in most cultivars with inflorescences forming at nodes following two-tendril pattern, typically producing inflorescence at every third node in alternating sequence with tendrils, though pattern varies among cultivars and with vine vigor affecting reproductive versus vegetative allocation. Primordium formation begins at shoot apex during active growth in Year N when lateral meristems that could form tendrils instead differentiate toward inflorescence identity, initiating as undifferentiated meristematic protrusions that will elaborate into complex branched structures bearing hundreds to thousands of flower primordia. Inflorescence architecture establishment during Year N involves primary axis formation creating main rachis structure, primary branch primordium initiation establishing first-order lateral branches typically 15-40 per inflorescence, secondary branch primordium formation on primary branches creating second-order branching, and flower primordium initiation on branch tips establishing potential flower number. Architectural elaboration follows basipetal sequence with apical regions of developing inflorescence primordium initiating structures first and basal regions forming later, creating developmental gradient within primordia that persists through dormancy into emergence and flowering phases. Dormancy entry in autumn preserves inflorescence primordia within buds as structured meristematic tissues with architectural organization established but organs incompletely differentiated, with primordia maintaining developmental potential through winter in arrested metabolic state. Chilling requirement satisfaction during winter represents essential developmental prerequisite for normal inflorescence development, with inadequate chilling causing poor inflorescence emergence, irregular flowering, and reduced fruit set affecting productivity. Emergence phase following bud burst in spring Year N+1 reveals inflorescence structures developing within buds, with inflorescences becoming externally visible as small compact clusters initially enclosed by expanding leaves, then elongating and expanding as flowering approaches. Inflorescence position within shoot architecture shows consistent pattern with typically first two nodes producing inflorescences in fruitful shoots following architectural program, though fruitfulness varies with cultivar, bud position, vine vigor, and previous season conditions affecting primordium formation. Developmental timing shows characteristic two-season progression with primordium formation requiring 6-12 weeks during Year N vegetative growth, dormancy spanning 4-6 months maintaining primordia in arrested state, and emergence-to-flowering requiring 4-6 weeks during Year N+1 spring depending on temperature accumulation. Environmental conditions during both seasons critically affect development with Year N conditions during primordium formation determining inflorescence number, architecture, and flower number potential, and Year N+1 conditions during emergence affecting developmental success, uniformity, and flowering quality. Temperature represents primary environmental factor affecting development with optimal 22-28°C during Year N primordium formation supporting maximum inflorescence initiation and architectural elaboration, chilling accumulation during winter affecting dormancy release and subsequent development, and spring temperatures in Year N+1 determining emergence rate and flowering timing. Light intensity during Year N primordium formation affects inflorescence initiation with adequate light promoting reproductive development while shade reduces fruitfulness shifting allocation toward vegetative growth, affecting subsequent season yield potential. Water availability during Year N affects inflorescence formation with moderate water stress potentially promoting reproductive allocation while severe stress reduces primordium formation, and Year N+1 water status affecting emergence and development quality. Nutrient availability particularly nitrogen during Year N affects reproductive versus vegetative allocation with moderate nitrogen supporting good inflorescence formation while excessive nitrogen promotes vegetative growth reducing fruitfulness, affecting yield potential. Hormonal regulation involves gibberellins with low concentrations promoting reproductive development and high concentrations favoring vegetative growth affecting inflorescence versus tendril determination, cytokinins potentially promoting inflorescence initiation and flower number, and other hormones coordinating developmental transitions. The process shows multiple distinguishable phases based on developmental stage and season, progressing from inflorescence primordium formation in Year N through dormancy maintenance to emergence in Year N+1, with each phase critical for successful development. Cultivar variation appears in fruitfulness with number of inflorescences per shoot ranging from 0-1 in low-fruitfulness varieties to 2-3 in highly fruitful types, inflorescence architecture affecting cluster size and structure, flower number per inflorescence determining berry number potential, and chilling requirements affecting development success in different climates. Regional timing shows primordium formation during June-August in warm climates, July-September in temperate regions, and July-August in cool areas of Year N, with emergence occurring April-May in warm climates, May in temperate regions, and May-June in cool areas of Year N+1. Viticultural management considerations include understanding that current season conditions affect next season productivity requiring forward-looking management, maintaining adequate but not excessive vigor during primordium formation optimizing reproductive allocation, ensuring sufficient light exposure promoting inflorescence initiation, managing water and nitrogen supporting balanced vegetative-reproductive development, and recognizing that primordium formation represents critical determinant of subsequent yield potential. Research applications include analyzing mechanisms regulating reproductive versus vegetative organ identity determination, investigating environmental and hormonal controls of inflorescence architecture affecting yield components, evaluating management effects on primordium formation influencing subsequent productivity, quantifying relationships between Year N conditions and Year N+1 yield outcomes, and assessing genetic variation in inflorescence development affecting productivity across environments.
Inflorescence_Development represents fundamental reproductive process spanning two growing seasons from primordium formation through emergence. Distinguished from Flowering which represents anthesis and fertilization processes. Subdivided into Inflorescence_Primordium_Formation occurring Year N and Inflorescence_Emergence occurring Year N+1. Critical determinant of yield potential requiring two-season perspective in management. Year N conditions during primordium formation fundamentally affect Year N+1 productivity.
Blütenstandsentwicklung
Gescheinbildung
Infloreszenzenbildung
cluster development
inflorescence formation
reproductive structure development
reproduktive Strukturentwicklung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 315-345 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 245-275 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Inflorescence Development
Inflorescence primordia become microscopically visible in latent buds 4-6 weeks after budbreak of the current season, reaching 0.3-0.5 mm in length by dormancy. Chardonnay typically initiates 2.1-2.4 inflorescence primordia per bud at nodes 4-10, while Sultana averages only 1.2-1.5 primordia. Final inflorescence size depends on primordium elaboration during spring of year N+1, with branch number increasing from 8-12 at dormancy to 15-25 at bloom through continued meristem activity.
A flowering disposition that is the inherited capacity of a Vitis vinifera plant to form compound inflorescences (panicles) from uncommitted primordia (anlagen) in latent buds, encompassing the initiation, differentiation, and elaboration of the branched flower-bearing structure over a two-year developmental cycle.
Inflorescence_Development_Disposition serves as the parent class for traits related to the formation of the grape cluster framework. The grapevine inflorescence is developmentally homologous to the tendril, with both arising from the same uncommitted primordium (anlage). The VvTFL1A, VvFT, and VvSOC1 genes regulate the balance between vegetative and reproductive development. Gibberellin promotes tendril fate while cytokinin promotes inflorescence fate.
Infloreszenzentwicklung
cluster primordium development
inflorescence formation capacity
panicle development trait
Carmona MJ et al. (2008) A molecular genetic perspective of reproductive development in grapevine. Journal of Experimental Botany 59:2579-2596 Srinivasan C, Mullins MG (1981) Physiology of flowering in the grapevine: a review. American Journal of Enology and Viticulture 32:47-63
Inflorescence Development Disposition
Inflorescence development rate assessment comparing cool and warm flowering conditions in Sauvignon Blanc reveals the temperature effect on flowering duration and fruit set, with cool flowering season (mean 15°C) showing slow development rate producing extended flowering of 14 days (variable fertilization timing, uneven berry development, mixed berry size population), while warm flowering (mean 22°C) shows rapid development with compressed flowering of 6 days (synchronized fertilization, uniform berry development, consistent berry size), demonstrating how inflorescence development rate affects fruit set outcomes through the temporal dynamics of flowering.
Inflorescence development rate quality is a developmental rate quality that represents the speed of inflorescence emergence, expansion, and progression through flowering stages, characterizing the rate of reproductive structure development that determines the duration of the critical flowering period affecting fruit set success through the temporal window for pollination and fertilization. Within the BFO framework, this quality specifically addresses the reproductive developmental dimension of phenological rate, distinguishing inflorescence and flower development rate from vegetative shoot development while recognizing that inflorescence development rate determines the timing and duration of flowering, with implications for the environmental conditions experienced during this critical phase and the synchronization of flowering with optimal conditions for fruit set. The ontological significance lies in inflorescence development rate affecting both the timing and success of fruit set, where rapid development produces compressed flowering periods potentially vulnerable to brief adverse weather events, while slower development extends the flowering window providing more opportunities for successful pollination but also prolonging exposure to flowering-period disease pressure (Botrytis, downy mildew on flowers). The temperature and cultivar basis for inflorescence development rate shows strong temperature response (rate increasing with temperature above approximately 15°C base) modified by cultivar genetics that determine both flowering date and the duration of the flowering period, with the interaction affecting the heat accumulation during flowering that influences both rate and success of fruit set. The flowering duration variation arising from development rate shows typical flowering periods of 5-15 days from first flower opening to completion, with compressed flowering in warm conditions (rapid development, shorter duration) and extended flowering in cool conditions (slower development, longer duration), each pattern having implications for fruit set uniformity and resulting berry size variation within clusters. The fruit set implications of inflorescence development rate include effects on the pollination-fertilization window (rapid development potentially reducing successful fertilization time), the synchrony of berry development within clusters (compressed flowering producing more uniform berry populations), and the integration of flowering with weather patterns (rate determining whether flowering occurs during favorable or unfavorable weather windows). Understanding inflorescence development rate quality enables prediction of flowering timing and duration, assessment of fruit set conditions, and timing of bloom-period management operations including fungicide applications and irrigation adjustments.
Inflorescence_Development_Rate_Quality addresses the speed of reproductive structure development determining flowering duration and fruit set conditions. This quality responds strongly to temperature affecting pollination success. The class supports fruit set prediction and bloom management through inflorescence development rate characterization.
Blühgeschwindigkeit
Blütenstandsentwicklungsrate
Infloreszenswachstumsrate
bloom progression rate
flowering rate
inflorescence growth rate
reproductive development rate
reproduktive Entwicklungsrate
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 59-144 Petrie PR, Clingeleffer PR (2005) Effects of temperature and light (before and after budburst) on inflorescence morphology and flower number of Chardonnay grapevines. Australian Journal of Grape and Wine Research 11(1): 59-65
Inflorescence Development Rate Quality
Pinot Noir in Oregon shows inflorescence elongation from early-mid May Year N+1 at BBCH 55 over 12-14 days, with compact 6-8 mm visible inflorescences elongating to 22-30 mm extended clusters through rachis and branch extension at 1.5-2.5 mm per day under cool climate spring conditions of 13-18°C mean temperature, adequate moisture, developing photosynthetic capacity from expanding leaves, typical of this moderate-vigor early-to-mid season cultivar producing medium-sized clusters approaching flowering readiness
Inflorescence elongation represents the primary growth phase following initial visibility where rachis extends and branches elongate increasing cluster size from compact 5-10 mm structures to elongated 20-40 mm clusters, establishing spatial framework for flower development and anthesis corresponding to BBCH 55 phenological stage. This critical developmental phase occurs during early-mid May in temperate regions following initial visibility, typically spanning 10-14 days as coordinated rachis and branch elongation proceeds at temperature-dependent rates transforming compressed clusters into extended structures. Elongation mechanism involves cell division in intercalary meristems within rachis and branch tissues producing new cells contributing to axis length, and progressive cell expansion throughout elongating axes where cells increase length 10-30 fold through turgor-driven wall extension creating visible growth. Rachis elongation proceeds through primary axis extension from initial 3-8 mm to final 15-30 mm depending on cultivar and conditions, with elongation rate typically 1-3 mm per day at optimal temperatures providing main cluster axis structure. Primary branch elongation occurs simultaneously with rachis extension as branches lengthen from initial 2-5 mm to final 8-18 mm, with branch elongation creating spatial separation between flower clusters distributed along rachis and branch axes. Secondary branch elongation where present creates additional architectural complexity with second-order branches extending from primary branches, contributing to compound cluster structure in highly branched cultivars. Flower bud spacing increases during elongation as rachis and branch extension separates attachment points, transitioning from tightly packed appearance at visibility to distinguishable individual flowers arranged along elongating axes. Elongation rate shows strong temperature dependence with development approximately doubling per 10°C increase between 10-25°C optimal range, with warm spring conditions promoting rapid elongation completing within 8-10 days while cool conditions extend period to 14-18 days. Cell division during elongation shows activity concentrated in intercalary meristem zones within rachis and branches with cycles of 24-36 hours producing cells arranged in longitudinal files contributing to axis elongation. Cell expansion follows and overlaps division phase with cells throughout elongating axes undergoing water uptake creating turgor pressure of 0.4-0.7 MPa driving cell wall extension, with expansion zone extending throughout much of rachis and branch length during peak elongation. Vascular differentiation proceeds during elongation with procambial strands differentiating into functional xylem vessels and phloem sieve elements establishing continuous vascular connection from shoot through rachis and branches to developing flowers, with vascular capacity increasing to support growing tissue mass and approaching anthesis demands. Mechanical properties during elongation show flexible herbaceous structure from thin primary cell walls allowing bending and positioning, with rigidity increasing progressively as walls thicken though final strength development awaits fruit set and berry development. Metabolic activity during elongation remains elevated with respiration 10-15 fold above mature tissue levels supporting rapid growth processes, with energy demands met through continued mobilization of stored reserves and increasingly from photosynthesis as leaf area develops. Protein synthesis operates at high capacity producing structural proteins for expanding cells, enzymes supporting rapid metabolism and wall modification, and beginning accumulation of proteins involved in flowering processes. Carbohydrate sources during elongation include declining dependence on stored reserves as shoots transition toward autotrophic growth, with photosynthesis from expanding leaves beginning to contribute though stored reserves remain important until several leaves fully expand. Hormonal regulation involves gibberellins supporting rachis and branch elongation with moderate concentrations promoting normal extension while very high or low levels may cause abnormalities, auxin from developing flowers coordinating rachis development and positioning, and cytokinins supporting cell division in meristematic zones. Environmental conditions during elongation critically affect development with temperature as primary factor determining rate and final dimensions, with optimal 15-22°C supporting maximum elongation while cool conditions reduce rate and extent, and warm conditions may accelerate development but risk stress effects. Light intensity affects elongation indirectly through effects on whole-plant vigor and photosynthetic capacity, with adequate light supporting vigorous elongation while shade may reduce vigor affecting cluster development. Water availability affects elongation through direct effects on cell turgor supporting expansion and on overall vine function, with adequate moisture essential for normal elongation while water stress reduces rate and final dimensions affecting cluster size. Nutrient availability particularly nitrogen affects elongation with adequate nutrition supporting vigorous development, though elongation depends substantially on stored reserves with current uptake becoming more important during rapid later growth. Frost vulnerability remains high during elongation with developing tissues highly susceptible to freeze damage at temperatures below -1°C, requiring continued protection in frost-prone regions through this critical period. Developmental coordination with vegetative growth shows concurrent shoot elongation and leaf expansion providing increasing photosynthetic capacity, with vegetative-reproductive balance determining resource allocation between shoot growth and inflorescence development. Cluster architecture expression during elongation reveals branching pattern established during previous season primordium formation, with highly branched types showing extensive primary and secondary branch elongation creating compound structures while simply branched types show limited branching producing loose small clusters. Developmental uniformity within cluster shows slight gradient with apical portions typically slightly more advanced reflecting basipetal primordium elaboration pattern, though elongation tends to reduce within-cluster heterogeneity preparing for synchronized flowering. The process represents second of four distinguishable inflorescence emergence phases focusing on size increase before architectural spreading and flower maturation, providing critical cluster development establishing framework for flowering. Duration typically spans 10-14 days from initial visibility through completion of major elongation depending on temperature, with warm conditions accelerating and cool conditions slowing progression. Cultivar variation appears in elongation rate with vigorous varieties showing rapid extension while less vigorous types show slower development, final rachis and branch lengths determining cluster size ranging from small compact 15-25 mm to large loose 30-45 mm structures, and elongation responsiveness to environmental conditions affecting development plasticity. Regional timing shows inflorescence elongation during early-mid May in warm climates, mid-May in temperate regions, and late May-early June in cool areas, with timing relative to fixed calendar dates varying but progression relative to thermal time accumulation consistent. Viticultural management considerations include monitoring elongation progress assessing development uniformity and timing predictions for flowering, maintaining frost protection through elongation period as tissues remain highly vulnerable, ensuring adequate resources supporting vigorous elongation without excessive vegetative competition, recognizing that elongation determines spatial cluster framework affecting subsequent fruit development, and understanding that stress during elongation can reduce final cluster size affecting yield. Research applications include quantifying environmental effects particularly temperature on elongation rate and final dimensions, analyzing hormonal regulation of rachis and branch elongation, evaluating relationships between elongation characteristics and ultimate yield components, modeling elongation processes for phenology prediction supporting management timing, and assessing genetic variation in elongation patterns affecting cluster architecture across cultivars.
Inflorescence_Elongation is second of four Inflorescence_Emergence subphases, distinguished by primary size increase through rachis and branch extension. Follows Inflorescence_Visibility. Transitions to Inflorescence_Expansion as spatial extension completes and architectural spreading begins. Corresponds to BBCH 55. Primary determinant of cluster size and flower spacing. Highly temperature-dependent process critical for proper cluster development.
Blütenstandsstreckung
Gescheinsstreckung
Spindelstreckung
Traubenstrecku ng
cluster elongation
inflorescence stretching
rachis extension
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 335-340 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 265-270
Inflorescence Elongation
Merlot in Bordeaux shows inflorescence emergence from late April through late May Year N+1 at BBCH 53-57 over 4-5 weeks, with dormant primordia resuming development and becoming visible as 3-5 mm compact clusters within opening buds, elongating to 25-35 mm and expanding with distinguishable branches and flower buds approaching flowering readiness under oceanic spring climate conditions of 12-18°C mean temperature, adequate moisture, accumulated 1000-1200 chilling hours during winter, supporting transition from dormant primordia to functional reproductive structures in this moderately fruitful mid-season cultivar
Inflorescence emergence represents the developmental phase in spring Year N+1 when dormant inflorescence primordia formed during previous season resume development within swelling buds and become externally visible as shoots elongate following bud burst, establishing visible reproductive structures that will undergo flowering preparation and anthesis. This critical reproductive phase occurs during April-May in temperate regions following winter dormancy and chilling requirement satisfaction, typically spanning 3-5 weeks from first inflorescence visibility within opening buds through complete emergence and expansion as compact clusters approaching flowering readiness at BBCH 53-57. Emergence initiation follows bud burst when swelling buds expose developing shoot apex and enclosed organs including leaves and inflorescences that have remained dormant through winter, with inflorescences becoming visible as small compact structures initially enclosed by expanding leaves. Dormancy release enabling emergence requires adequate chilling accumulation during winter typically 800-1400 hours below 7°C depending on cultivar, with insufficient chilling causing poor emergence, erratic development, and potentially reduced flowering affecting yield. Development resumption involves metabolic reactivation in dormant primordium tissues with respiration increasing from minimal dormancy levels, protein synthesis resuming to support growth, and cell division and expansion initiating in meristematic regions driving visible enlargement. Inflorescence visibility marks first external observation of reproductive structures within opening buds at BBCH 53, with small clusters 2-5 mm length visible between separating bud scales and emerging leaves, confirming successful inflorescence formation during previous season and dormancy survival. Inflorescence elongation proceeds through coordinated rachis extension and branch elongation increasing cluster size from initial 5-10 mm to 20-40 mm over 2-3 weeks, with elongation rate temperature-dependent doubling approximately every 10°C increase within 10-25°C range. Inflorescence expansion involves branch spreading and flower primordium enlargement transforming compact undifferentiated structure into recognizable cluster form with distinguishable branches and enlarging flower buds, though flowers remain closed as green buds during emergence phase with anthesis occurring subsequently. Rachis development establishes main cluster axis structure through cell division and expansion in rachis tissues, with primary and secondary branch rachis similarly elongating and thickening providing structural framework supporting developing flowers and eventually fruit. Flower development during emergence shows continuation from primordial state with floral organ primordia differentiating into recognizable sepals, petals, stamens, and carpels though organs remain small and enclosed, with full flower development and opening occurring during subsequent flowering phase. Inflorescence position relative to shoot structure shows characteristic pattern with first inflorescence typically at node 3-4 on fruitful shoots becoming visible as shoot has 3-5 unfolded leaves, and second inflorescence if present at node 5-6 emerging slightly later as shoot continues elongating. Developmental coordination with vegetative growth shows inflorescences emerging concurrently with leaf unfolding and shoot elongation, with all processes drawing on stored carbohydrate reserves mobilized from woody tissues until developing leaves achieve photosynthetic capacity. Resource demand during emergence creates competition between vegetative and reproductive development for limited reserves, with adequate reserve status supporting vigorous concurrent development while depleted reserves may limit emergence affecting reproductive success. Vascular connection between emerging inflorescence and shoot establishes functional xylem and phloem providing water delivery supporting turgor-driven growth and enabling nutrient import supporting rapid development, with vascular capacity adequate for emergence needs but requiring further development supporting subsequent fruit growth. Metabolic activity during emergence shows progressively increasing respiration as tissues reactivate and growth intensifies, reaching 8-12 fold above dormancy levels by peak emergence supporting rapid cell division and expansion. Hormonal regulation involves gibberellins with moderate concentrations supporting rachis elongation and flower development, cytokinins promoting cell division and delaying senescence, and auxin coordinating growth and vascular development. Environmental conditions during emergence critically affect development with temperature as primary factor determining rate, with optimal 15-22°C supporting rapid uniform emergence while cool conditions delay development and excessive heat may cause abnormalities. Light intensity affects emergence indirectly through effects on whole-plant photosynthesis and resource availability, with adequate light supporting vigorous development while severe shade may limit emergence vigor. Water availability affects emergence through effects on cell turgor supporting expansion and on overall vine function, with adequate moisture essential for normal development while stress may reduce vigor or cause developmental abnormalities. Nutrient availability particularly nitrogen affects emergence with adequate nutrition supporting vigorous development, though emergence primarily depends on stored reserves with current uptake becoming important later during rapid growth. Frost risk during emergence represents major concern with developing reproductive tissues highly vulnerable to freeze damage at temperatures below -2°C, requiring protection measures in frost-prone regions during this critical period. The process shows four distinguishable developmental phases progressing from inflorescence visibility within opening buds, through inflorescence elongation extending cluster structure, inflorescence expansion spreading branches and enlarging flowers, to rachis development establishing structural framework. Cultivar variation appears in emergence timing with early-bursting varieties showing early inflorescence emergence while late types emerge later, emergence vigor affecting cluster development rate, and architectural expression of primordium structure determined during previous season. Regional timing shows inflorescence emergence during April-May in warm climates, May in temperate regions, and May-June in cool areas, with timing relative to bud burst and vegetative development consistent across regions. Viticultural management considerations include monitoring emergence progress assessing reproductive development status, protecting developing inflorescences from frost damage through active or passive protection measures, ensuring adequate vine reserves supporting vigorous emergence through appropriate previous season management, recognizing that emergence represents realization of previous season primordium formation, and understanding that abnormal emergence indicates problems with either primordium formation, dormancy, or current season conditions. Research applications include analyzing developmental transitions from dormancy to active growth, investigating environmental regulation of emergence timing and rate, evaluating relationships between emergence characteristics and subsequent flowering and yield, quantifying resource requirements and limitations during emergence, and assessing cultivar variation in emergence patterns affecting phenological timing and adaptation across climates.
Inflorescence_Emergence is second major subphase of Inflorescence_Development, occurring during spring Year N+1 following winter dormancy. Distinguished by resumption of development and external visibility of inflorescence structures formed during previous season. Subdivided into Inflorescence_Visibility, Inflorescence_Elongation, Inflorescence_Expansion, and Rachis_Development. Follows Inflorescence_Primordium_Formation that occurred Year N. Precedes Flowering phase. Critical period for frost protection and establishing reproductive success.
Blütenstandsaustritt
Gescheinssichtbarkeit
Infloreszenzemergenz
Traubenerscheinen
cluster emergence
inflorescence expression
inflorescence visibility
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 328-345 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 260-275 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Inflorescence Emergence
Syrah in Rhône Valley shows inflorescence expansion from mid-late May Year N+1 at BBCH 57 over 8-10 days, with elongated 25-35 mm clusters spreading branches from compressed to 50-65 degree angles and flower buds enlarging from 1.5-2.0 mm to 3.0-4.0 mm approaching flowering readiness under warm spring Mediterranean conditions of 17-23°C mean temperature, adequate moisture, full photosynthetic capacity from developed canopy, typical of this moderately-to-highly fruitful mid-to-late season cultivar expressing compound cluster architecture before anthesis
Inflorescence expansion represents the architectural spreading phase following elongation where branches separate laterally from compressed positions, flower buds enlarge approaching anthesis readiness, and cluster achieves final pre-flowering form with open architecture revealing individual flowers, corresponding to BBCH 57 phenological stage. This critical preparatory phase occurs during mid-late May in temperate regions following rachis elongation, typically spanning 7-10 days as coordinated branch spreading and flower maturation proceed transforming elongated but compact clusters into fully expanded structures ready for flowering. Expansion mechanism involves branch angle increase from compressed acute angles to spreading obtuse angles creating open architecture, continued branch elongation contributing to separation, and flower bud enlargement from small 1-2 mm primordia to plump 3-4 mm buds approaching cap fall. Branch spreading proceeds through differential growth on branch abaxial versus adaxial sides creating angular change, with primary branches moving from 20-30 degree angles to 45-70 degrees relative to rachis axis, and secondary branches similarly spreading creating three-dimensional open structure. Flower bud enlargement represents major developmental change with flower organs completing differentiation and expansion within enclosing calyptra cap, with sepals remaining fused as cap, petals, stamens, and pistil enlarging substantially as anthesis approaches. Flower development during expansion includes stamen filament elongation positioning anthers for pollen release, anther maturation with pollen completing development becoming viable, pistil enlargement with ovary expanding and style-stigma developing receptivity, and nectary development preparing for nectar secretion. Cluster appearance transitions from compact elongated structure with indistinct flowers to fully expressed architecture with clearly visible individual flower buds arranged in characteristic pattern, with cluster form revealing branching complexity established during previous season primordium formation. Flower number becomes definitively countable during expansion as individual buds separate and become distinct, providing accurate assessment of potential berry number though actual berry set will depend on successful pollination and fertilization during subsequent flowering. Metabolic activity remains elevated during expansion supporting continued growth and flower maturation, with respiration 8-12 fold above mature tissue levels providing energy for developmental processes. Photosynthetic contribution increases during expansion as shoot leaf area expands and matures, with developing leaves beginning to supply significant photosynthate supplementing declining stored reserves, though clusters remain net carbon consumers requiring import. Vascular development continues during expansion with vessel diameter and number increasing in rachis and branches enhancing water transport capacity to 4-8 mg per second adequate for supporting full cluster and approaching berry demands, and phloem development increasing assimilate import capacity. Sugar concentration in rachis and flower tissues during expansion shows minimal levels as structures remain in growth phase consuming imported assimilates rather than accumulating storage products, with sugar dynamics changing dramatically after fruit set when berries begin accumulating photosynthate. Hormonal regulation during expansion involves gibberellins maintaining growth promotion though concentrations may begin declining as flowering approaches, auxin from developing flowers coordinating development, and possibly ethylene beginning involvement in maturation processes. Environmental conditions during expansion affect development with temperature continuing as primary factor though optimal range shifts slightly higher to 16-24°C supporting maximum expansion and flower maturation, with cool conditions delaying progression and very warm conditions potentially accelerating development. Light intensity affects expansion through photosynthetic effects supporting vigorous development, with well-exposed shoots showing robust expansion while shaded shoots may show reduced vigor affecting cluster quality. Water availability remains critical during expansion with adequate moisture essential for turgor supporting branch spreading and flower enlargement, while water stress reduces expansion vigor potentially affecting flower development and subsequent fruit set. Nutrient availability particularly nitrogen affects expansion with adequate nutrition supporting vigorous development, with nitrogen from both current uptake and remobilization from storage supporting rapid tissue growth and protein synthesis. Pest pressure begins increasing during expansion as developing reproductive tissues attract insects including grape berry moth, leafhoppers, and thrips requiring monitoring and possible management interventions. Disease pressure also intensifies with expanding clusters creating favorable microclimate for powdery mildew development requiring protective fungicide applications particularly in susceptible varieties or conducive conditions. Frost vulnerability declines during expansion as tissues mature and temperatures typically rise reducing freeze risk, though late spring frosts remain possible in some regions requiring vigilance. Developmental coordination with vegetative growth shows continued concurrent shoot elongation and leaf expansion with shoots typically having 10-15 leaves at expansion completion, with balanced vegetative-reproductive development supporting subsequent flowering and fruit set. Cluster position relative to canopy architecture becomes established during expansion with shoot positioning operations determining cluster exposure affecting microclimate during critical flowering period. Within-cluster developmental uniformity increases during expansion as flower maturation proceeds synchronously preparing for coordinated anthesis, with well-developed clusters showing good flowering synchrony supporting uniform fruit set. The process represents third of four distinguishable inflorescence emergence phases focusing on architectural spreading and flower maturation preparing for anthesis, completing transformation from dormant primordium to functional reproductive structure. Duration typically spans 7-10 days depending on temperature and conditions, with warm favorable conditions promoting rapid progression while cool conditions or stress may extend period. Cultivar variation appears in expansion vigor affecting final cluster openness with some varieties showing tight compact architecture while others show very loose open structure, flower size at expansion affecting visual appearance and possibly pollination efficiency, and expansion rate affecting timing progression toward flowering. Regional timing shows inflorescence expansion during mid-late May in warm climates, late May-early June in temperate regions, and early-mid June in cool areas, with timing relative to thermal accumulation consistent though calendar dates vary geographically. Viticultural management considerations include monitoring expansion progress for flowering timing predictions, implementing pre-flowering disease management protecting expanding susceptible tissues, positioning shoots optimizing cluster exposure for favorable flowering microclimate, assessing cluster uniformity and architecture affecting disease pressure and quality potential, and recognizing expansion as final preparatory phase before critical flowering period. Research applications include analyzing developmental processes during flower maturation, evaluating environmental effects on expansion affecting flowering readiness, quantifying relationships between expansion characteristics and flowering uniformity, assessing pest and disease colonization during expansion informing management thresholds, and documenting genetic variation in cluster architecture expression affecting fruit quality potential.
Inflorescence_Expansion is third of four Inflorescence_Emergence subphases, distinguished by architectural spreading and flower maturation. Follows Inflorescence_Elongation. Transitions to Rachis_Development final maturation and directly toward Flowering phase. Corresponds to BBCH 57. Critical preparatory phase establishing flowering readiness. Important period for disease management protecting susceptible tissues.
Blütenstandsentfaltung
Gescheinsspreizung
Infloreszenzerweiterung
Traubenöffnung
cluster spreading
flower cluster expansion
inflorescence opening
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 340-345 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 270-275
Inflorescence Expansion
Inflorescence initiation occurs primarily at nodes 3-12 from the shoot base, with peak initiation frequency at nodes 5-8 in most cultivars. Pinot Noir shows high initiation disposition with 85-95% of anlagen converting to inflorescence primordia at optimal nodes, while Cardinal converts only 40-60%. Temperature during the initiation window (4-8 weeks post-budbreak) critically affects conversion rate, with 25-30°C promoting initiation and temperatures below 20°C or above 35°C reducing it. Light exposure to developing buds increases initiation by 20-40% compared to shaded buds.
An inflorescence development disposition that is the inherited capacity of a Vitis vinifera plant to convert uncommitted primordia (anlagen) in latent buds into inflorescence meristems rather than tendril meristems, determining the potential for reproductive versus vegetative development in the following season.
Inflorescence initiation represents the first commitment step in the two-year reproductive cycle. The process is regulated by the ratio of gibberellins to cytokinins, with high cytokinin favoring inflorescence fate. Environmental factors including temperature, light, and carbohydrate availability during the critical 4-8 week window after budbreak determine initiation success. The VvTFL1A gene acts as a repressor of inflorescence identity.
Blütenstandsinitiation
Gescheinanlage
Infloreszenzinduktion
cluster initiation capacity
inflorescence induction trait
reproductive primordium formation
Buttrose MS (1974) Climatic factors and fruitfulness in grapevines. Horticultural Abstracts 44:319-326 Sánchez LA, Dokoozlian NK (2005) Bud microclimate and fruitfulness in Vitis vinifera L. American Journal of Enology and Viticulture 56:319-329
Inflorescence Initiation Disposition
Merlot in Bordeaux shows inflorescence meristem differentiation during mid-July Year N at BBCH 25-31 within developing latent buds over 10-12 days, with initial 60-80 micrometer primordia developing organized meristem structure with distinct zonal organization reaching 180-220 micrometers establishing framework for 22-32 primary branches under conditions of 22-26°C temperature, adequate light in well-exposed canopy, balanced fertility, and moderate water status typical of this moderately fruitful mid-season cultivar
Inflorescence meristem differentiation represents the developmental stage where initial undifferentiated reproductive primordium develops organized inflorescence meristem structure with distinct functional zones establishing architectural framework that will guide subsequent branch and flower formation, transitioning from simple meristematic protrusion to complex organized reproductive meristem. This critical organizational stage occurs within developing latent bud during mid-summer Year N typically 1-2 weeks after initial primordium emergence, spanning approximately 7-14 days as meristem structure and zoning patterns establish before branch primordium initiation begins. Meristem organization development involves establishment of distinct zones within growing primordium including apical central zone maintaining indeterminate meristematic identity and stem cell populations, peripheral zone producing lateral organ primordia including branch initiation sites, and rib zone generating primary axis tissues extending primordium length. Central zone establishment maintains population of slowly dividing stem cells expressing meristem identity genes including WUSCHEL and CLAVATA homologs, providing reservoir of uncommitted cells sustaining continued meristem activity during architectural elaboration. Peripheral zone organization shows higher cell division rates producing cells that will form lateral branch primordia, with spatial patterning establishing regular arrangement of initiation sites along developing primary axis determining ultimate branch number and distribution. Meristem shape transitions from initial rounded or dome structure to increasingly elongated form as primary axis extends, creating characteristic inflorescence primordium morphology distinguishing reproductive from vegetative organ development. Cell division patterns show highest rates in peripheral zones where branch primordia will initiate at 24-36 hour cycles, moderate rates in central zone maintaining meristem at 36-48 hour cycles, and lower rates in maturing rib zone tissues forming primary axis at 48-72 hour cycles. Gene expression patterns during differentiation include sustained expression of reproductive identity genes maintaining inflorescence character, activation of meristem organization genes establishing zonal structure including SHOOTMERISTEMLESS and KNOTTED1 homologs, and beginning expression of auxin response genes involved in lateral organ positioning. Hormonal regulation involves auxin gradients establishing positions where branch primordia will initiate with local auxin maxima marking future branch sites, cytokinins maintaining meristem activity and supporting cell division, and low gibberellin levels permitting reproductive development rather than reverting to vegetative tendril formation. Morphological landmarks appearing during differentiation include elongation of primordium from initial 80-120 micrometers to 150-250 micrometers establishing primary axis framework, beginning surface irregularities marking future branch initiation sites though distinct branch primordia not yet emerged, and increased primordium complexity distinguishing from simpler tendril primordium structure. Vascular tissue development begins within differentiating meristem with procambial strands extending through developing primary axis, establishing vascular framework that will supply developing branches and eventually flowers, though actual differentiation to functional xylem and phloem occurs later. Developmental rate during differentiation shows temperature dependence with optimal 24-28°C supporting rapid progression completing differentiation within 7-10 days, cooler temperatures extending duration to 12-16 days, affecting overall developmental window available for architectural elaboration. Metabolic activity remains elevated during differentiation supporting high cell division rates and growth, with respiration 10-15 fold above mature tissue levels providing energy for rapid development. Protein synthesis operates at high capacity producing cell cycle regulators, transcription factors controlling zonal organization, and structural proteins supporting primordium growth and organization. Architectural commitment during differentiation establishes framework determining ultimate inflorescence structure with meristem organization patterns affecting branch number potential, though actual branch formation occurs in subsequent stage. Environmental conditions during differentiation affect organizational quality with adequate light supporting normal meristem differentiation, appropriate temperature maintaining optimal developmental rate, and sufficient resources enabling complete organization establishing robust architectural framework. Canopy position affects differentiation with exposed shoots showing more robust meristem organization supporting higher branch numbers, while shaded shoots may show reduced differentiation limiting architectural complexity. Stress during differentiation can impair organizational processes reducing subsequent branching potential or causing developmental abnormalities affecting inflorescence architecture and flower number. Cultivar differences in meristem organization may affect subsequent architectural development with some varieties showing early establishment of numerous branch initiation sites supporting compound inflorescence architecture while others show fewer sites determining simpler structure. The process represents second distinguishable stage of inflorescence primordium formation following initial primordium emergence and preceding branch primordium formation, establishing organized reproductive meristem structure. Timing within season shows differentiation occurring mid-late summer approximately July-August in temperate regions following initial primordium formation and during continued active shoot growth, with timing varying based on cultivar and environmental conditions. Duration of differentiation stage spans approximately 7-14 days from beginning of zonal organization to emergence of first branch primordia, with duration varying based on temperature and developmental rate. Regional timing shows inflorescence meristem differentiation during July-August in warm climates, August in temperate regions, and late July-August in cool areas corresponding to mid-summer period following primordium initiation. Viticultural significance includes establishing organizational framework determining subsequent architectural elaboration potential, understanding that conditions during differentiation affect ultimate inflorescence complexity and flower number capacity, and recognizing that stress during differentiation can permanently limit reproductive potential. Research applications include analyzing molecular mechanisms of meristem organization and zonal patterning, investigating gene regulatory networks controlling inflorescence meristem identity and structure, documenting cellular and developmental processes during meristem differentiation through microscopy and molecular techniques, evaluating environmental effects on meristem organization affecting architectural outcomes, and assessing genetic variation in meristem organization affecting inflorescence structure across cultivars.
Inflorescence_Meristem_Differentiation is second of four Inflorescence_Primordium_Formation subphases, distinguished by establishment of organized meristem structure with functional zoning. Follows Initial_Inflorescence_Primordium. Transitions to Primary_Branch_Primordium_Formation as branch initiation begins. Establishes architectural framework determining subsequent elaboration capacity. Critical organizational stage affecting ultimate inflorescence complexity.
Blütenstandsmeristem-Differenzierung
Infloreszenzmeristem-Musterbildung
Meristemorganisation
inflorescence meristem structuring
meristem organization
reproductive meristem patterning
reproduktive Meristemstrukturierung
Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337 Srinivasan C & Mullins MG (1981) Physiology of flowering in the grapevine, Am J Enol Vitic 32:47-63 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Inflorescence Meristem Differentiation
The inflorescence number of Riesling documents predominantly two inflorescences per shoot (OIV 152:3) on well-exposed, normally growing shoots from mature vines on moderate-vigor rootstock — the two inflorescences are typically positioned at nodes 3 and 4 (occasionally 3 and 5), opposite the leaf at each position, with a third inflorescence occasionally developing at node 5 on vigorous shoots in warm seasons (approximately 15-20% of shoots). This consistent two-cluster fruitfulness is characteristic of Riesling's reliable yield potential and contrasts with the higher fruitfulness of Grenache (frequently 2-3 inflorescences per shoot, OIV 152:3-4, contributing to Grenache's tendency toward overcropping) and the lower fruitfulness of Nebbiolo (frequently 1-2 inflorescences per shoot, OIV 152:2-3, with significant proportion of non-fruitful basal shoots necessitating longer cane pruning to position fruitful buds).
An inflorescence description component that records the typical number of inflorescences produced per shoot, scored according to OIV descriptor 152 on a scale from zero (state 1, non-fruitful) through one (state 2), two (state 3), three (state 4), to more than three (state 5), reflecting the cultivar-specific fruitfulness determined by the number of inflorescence primordia initiated within the compound bud during the previous growing season and successfully developed to visible inflorescences in the current season.
Inflorescence_Number documents a character of direct economic importance — the number of clusters per shoot is a primary yield component that, multiplied by cluster weight, determines yield per shoot and consequently yield per vine and yield per hectare. The number of inflorescences is determined during bud organogenesis (the initiation and development of inflorescence primordia within the compound bud during the season preceding bloom), controlled by the interaction of cultivar genotype with the environmental conditions experienced during the critical bud initiation period — high temperatures (25-30°C day/15-20°C night), adequate carbohydrate supply (high leaf-to-fruit ratio, adequate sunlight), and moderate vigor promote inflorescence primordium initiation and increase fruitfulness: the cultivar genotype determines the baseline fruitfulness and the responsiveness to environmental signals, with inherently fruitful cultivars (Grenache, Müller-Thurgau, Merlot) producing 2-3 inflorescences per shoot under a wide range of conditions, while less fruitful cultivars (Nebbiolo, Sultanina, some Muscat types) require optimal conditions and specific bud positioning (distal buds on long canes) to achieve adequate fruitfulness. The fruitfulness character has practical implications for pruning system selection — cultivars with high basal bud fruitfulness (producing inflorescences from buds at positions 1-3) are suited to spur pruning, while cultivars requiring distal buds for fruitfulness necessitate cane pruning with retention of 8-12 bud canes.
Blütenstandszahl
Infloreszenzen-pro-Trieb
Triebfruchtbarkeit
cluster number per shoot
inflorescences per cane
shoot fruitfulness record
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Dry PR (2000) Canopy management for fruitfulness, Australian Journal of Grape and Wine Research 6: 109-115
Inflorescence Number
Inflorescence number per shoot ranges from 1.0-3.5 across V. vinifera cultivars under standard management. Müller-Thurgau consistently produces 2.5-3.0 inflorescences per shoot, while Cabernet Sauvignon averages 1.5-2.0. Highly fruitful cultivars such as Trebbiano Toscano may exceed 3.0 inflorescences per shoot at basal nodes. The coefficient of variation for inflorescence number within a cultivar typically ranges from 15-30%, reflecting environmental and positional effects. Crop estimation models use historical inflorescence number data with 85-92% prediction accuracy.
An inflorescence development disposition that is the inherited capacity of a Vitis vinifera plant to develop a characteristic number of inflorescences per shoot, determined by the frequency of inflorescence initiation across bud positions and the survival of initiated primordia through dormancy and spring development.
Inflorescence number disposition interacts with bud fruitfulness but represents specifically the realized count of developed inflorescences rather than the presence of primordia. Not all initiated primordia develop into visible inflorescences due to abortion during late dormancy or early spring. The trait shows moderate heritability (h² = 0.45-0.60) and significant genotype × environment interaction.
Gescheinzahl
Infloreszenzanzahl pro Trieb
Traubenzahl-Disposition
bunch number disposition
inflorescence frequency trait
Dry PR (2000) Canopy management for fruitfulness. Australian Journal of Grape and Wine Research 6:109-115 Clingeleffer PR (2001) Crop development, crop estimation and crop control to secure quality and production of major wine grape varieties: a national approach. Final Report to Grape and Wine Research and Development Corporation, CSIRO Plant Industry
Inflorescence Number Disposition
The inflorescence position of Cabernet Sauvignon documents primary fruitfulness at nodes 3-5 (counting from the base of the one-year-old cane, with node 1 being the basal bud), with the first inflorescence most commonly at node 3 and the second at node 4 — this basal fruitfulness pattern means that Cabernet Sauvignon produces adequate crop from two-bud spurs (retaining nodes 1-2 as renewal positions and relying on nodes 3-4 of the previous year's shoot for cropping), making it well-suited to spur-pruned cordon training systems used throughout Bordeaux, California, and Australia. This basal fruitfulness contrasts with the distal fruitfulness of Sultanina (Thompson Seedless), where fruitful buds are concentrated at nodes 6-12 and basal buds (nodes 1-4) are predominantly vegetative, requiring retention of 12-15 bud canes for adequate cropping.
An inflorescence description component that records the node positions along the shoot at which inflorescences are inserted, documenting the typical range of fruitful node positions (counted from the shoot base) that characterizes the cultivar-specific distribution of reproductive structures along the shoot axis, with inflorescence position data essential for pruning system design and bud load calculations that ensure retention of fruitful buds.
Inflorescence_Position documents a character of critical practical importance for viticultural pruning system design — the node positions at which a cultivar produces fruitful buds determines whether the cultivar can be spur-pruned (relying on basal buds at positions 1-3), must be cane-pruned (requiring retention of longer canes to include fruitful buds at positions 5-12), or can accommodate either system. The positional distribution of fruitfulness along the shoot reflects the bud-by-bud variation in inflorescence primordium initiation during the compound bud organogenesis period — in basally fruitful cultivars, inflorescence primordia are initiated in buds at all positions including the basal nodes, while in distally fruitful cultivars, the basal buds fail to initiate inflorescence primordia (or initiate them but subsequently abort) and only the mid-cane and distal buds develop fruitful primordia: this positional variation is controlled by the interaction of cultivar genotype with the microenvironmental conditions experienced by each bud during organogenesis, particularly light exposure (shaded basal buds are less fruitful than well-exposed distal buds in all cultivars) and correlative inhibition from the growing shoot tip (stronger inhibition reducing fruitfulness in proximal buds). The inflorescence position data recorded in the VVO enables the prescription of cultivar-appropriate pruning systems and bud load targets for vineyard management.
Blütenstands-Position
Infloreszenz-Insertionsstelle
cluster insertion position
fruchtbare Knotenposition
fruitful node location
inflorescence node number
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press Dry PR (2000) Canopy management for fruitfulness, Australian Journal of Grape and Wine Research 6: 109-115 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia
Inflorescence Position
Pinot Noir in Burgundy shows inflorescence primordium formation during July-August Year N at BBCH 19-35 within developing latent buds over 8-10 weeks, initiating 1-2 inflorescence primordia per bud with 18-28 primary branches per inflorescence and 200-400 flower primordia under conditions of 20-26°C temperature, adequate light exposure in well-managed canopy, moderate water stress, and balanced nitrogen fertility typical of this moderately fruitful early-to-mid season cultivar, establishing architectural framework determining Year N+1 crop potential
Inflorescence primordium formation represents the initial reproductive organogenesis phase occurring during current season vegetative growth where lateral meristems at shoot apex differentiate toward inflorescence identity and elaborate complex branched architecture establishing structural framework and flower number potential that will determine crop capacity in following season. This critical developmental process occurs within developing latent buds during summer Year N, typically spanning 6-12 weeks from initial primordium initiation through architectural elaboration to dormancy entry, establishing inflorescence structures that will remain dormant through winter and emerge during bud burst Year N+1. Primordium formation begins at shoot apex in primary bud meristems during active vegetative growth typically June-August in temperate regions Year N, when lateral meristems at positions following architectural pattern differentiate toward reproductive rather than vegetative tendril identity. Inflorescence identity determination involves suppression of tendril developmental program and activation of reproductive meristem identity genes, with environmental signals including light quality and quantity, water status, nitrogen availability, and hormonal balance particularly gibberellin concentrations affecting reproductive versus vegetative fate decisions. Initial primordium development shows undifferentiated meristematic protrusion emerging from primary bud axis in lateral position opposite leaf primordium, similar in early stages to tendril primordium but diverging through differential gene expression activating inflorescence-specific developmental programs. Inflorescence meristem differentiation transforms initial undifferentiated primordium into organized inflorescence meristem with distinct zones including apical zone maintaining indeterminate growth capacity and lateral zones producing branch primordia, establishing architectural organization that will guide subsequent development. Primary branch primordium formation proceeds basipetally from inflorescence apex with lateral meristems initiating along primary axis in regular phyllotactic pattern, typically producing 15-40 primary branch primordia per inflorescence depending on cultivar and development conditions, with branch initiation establishing first-order architectural framework. Secondary branch primordium formation occurs on primary branch primordia creating second-order branching structure, with secondary branches varying from absent in simple inflorescence architectures to numerous in highly compound structures, determining inflorescence complexity and ultimate flower number capacity. Flower primordium initiation begins on branch tips during late primordium formation phase, with individual flower primordia developing as small meristematic protrusions that will differentiate into complete flowers during following season, with flower number per inflorescence ranging from 100-200 in small clusters to 500-1500 in large compound inflorescences. Architectural organization within developing primordium shows basipetal gradient with apical regions most advanced in development containing initiated flower primordia while basal regions show progressively earlier stages with recently formed branch primordia, creating developmental heterogeneity persisting through dormancy into emergence and flowering. Developmental timing within season shows progression from inflorescence identity determination occurring as early as node formation at shoot apex, through rapid primordium elaboration over 4-8 weeks as architectural elements initiate, to final maturation and dormancy entry as shoot growth declines in late summer-autumn. Cell division rates during active primordium formation show high mitotic activity supporting rapid organ initiation, with divisions occurring every 24-48 hours in meristematic zones producing cells for expanding primordial structures. Vascular connection establishment between developing inflorescence primordium and bud axis occurs early in formation providing pathway for nutrient and hormone delivery supporting primordium development, though transport is minimal during dormancy maintaining primordia through stored reserves. Metabolic activity during primordium formation shows elevated respiration supporting growth processes, declining to minimal maintenance levels as primordia mature and enter dormancy. Hormonal regulation centrally involves gibberellins with low endogenous concentrations or exogenous application of anti-gibberellins promoting inflorescence formation while high gibberellin levels favor tendril formation, cytokinins potentially promoting inflorescence initiation and branching affecting architecture, and auxin coordinating meristem activity and organ positioning. Environmental factors during formation critically affect developmental outcomes with light intensity and quality strongly affecting reproductive allocation through photosynthetic production and photomorphogenic signaling, with shade reducing fruitfulness shifting toward vegetative growth. Temperature during primordium formation shows optimal range 22-28°C supporting maximum inflorescence initiation and architectural elaboration, with suboptimal temperatures reducing reproductive allocation. Water status affects primordium formation with moderate water stress during formation period potentially enhancing reproductive allocation as stress response promoting reproduction, while severe stress reduces primordium formation through limitation of metabolic resources. Nutrient availability particularly nitrogen shows complex effects with moderate nitrogen supporting good primordium formation providing resources for development, while excessive nitrogen promotes vegetative growth reducing reproductive allocation, requiring balanced nutrition for optimal fruitfulness. Carbon availability from photosynthesis affects primordium formation with adequate carbohydrate production supporting reproductive development, while carbon limitation from excessive crop load, disease, or stress reduces formation affecting subsequent productivity. Canopy microclimate affects primordium formation with well-exposed shoots showing higher fruitfulness through favorable light environment promoting reproductive development, while shaded interior shoots show reduced inflorescence formation shifting toward vegetative growth. Shoot vigor affects primordium formation with moderate vigor supporting optimal reproductive allocation, while excessive vigor promotes vegetative growth reducing fruitfulness and insufficient vigor limits formation through resource constraints. Node position affects primordium formation with basal nodes typically most fruitful having optimal development conditions, middle nodes moderately fruitful, and apical nodes often unfruitful formed late under declining conditions. The process shows four distinguishable developmental phases progressing from initial inflorescence primordium initiation establishing reproductive identity, through inflorescence meristem differentiation organizing architectural framework, primary branch formation establishing first-order structure, to secondary branch formation completing architectural elaboration. Cultivar variation appears in fruitfulness with inflorescence initiation frequency at nodes ranging from low fruitfulness producing inflorescences at few nodes to high fruitfulness at most nodes, inflorescence architecture ranging from simple with few branches to highly compound with extensive branching, and flower number per inflorescence determining berry number potential. Regional timing shows inflorescence primordium formation during June-August in warm climates, July-September in temperate regions, and July-August in cool areas corresponding to active vegetative growth period when conditions favor primordium formation. Viticultural management considerations include recognizing that current season management affects next season crop potential requiring forward planning, maintaining moderate balanced vigor optimizing reproductive allocation, ensuring adequate light exposure through canopy management promoting inflorescence formation, managing water and nitrogen supporting reproductive development without excessive vegetative growth, controlling crop load allowing adequate resources for primordium formation, and understanding that conditions during primordium formation represent critical determinant of subsequent yield. Research applications include analyzing molecular mechanisms of reproductive versus vegetative organ identity specification, investigating environmental and hormonal regulation of inflorescence architecture, evaluating management effects on primordium formation affecting yield outcomes, quantifying developmental processes within primordia documenting architectural elaboration, and assessing genetic variation in inflorescence development affecting productivity potential across cultivars and environments.
Inflorescence_Primordium_Formation is first major subphase of Inflorescence_Development, occurring entirely during Year N vegetative growth period. Distinguished by primordium elaboration within latent buds establishing architectural framework. Subdivided into Initial_Inflorescence_Primordium, Inflorescence_Meristem_Differentiation, Primary_Branch_Primordium_Formation, and Secondary_Branch_Primordium_Formation. Followed by dormancy then Inflorescence_Emergence in Year N+1. Critical determinant of subsequent crop potential.
Blütenstandsorganogenese
Blütenstandsprimordienbildung
Gescheinsanlagebildung
Infloreszenzinitiierung
cluster primordium formation
inflorescence initiation
inflorescence organogenesis
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 315-328 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337 Srinivasan C & Mullins MG (1981) Physiology of flowering in the grapevine, Am J Enol Vitic 32:47-63
Inflorescence Primordium Formation
An inflorescence rachis system on a primary inflorescence of Vitis vinifera cv. Syrah in the Rhône Valley, France, consisting of the main central rachis measuring 16.5 cm length and 3.8 mm diameter at base tapering to 2.0 mm at apex, bearing 14 primary branch rachides (first-order laterals) ranging 3.5-7.2 cm length and 1.8-3.0 mm diameter, primary branches bearing 45 secondary branch rachides (second-order laterals) ranging 1.2-3.5 cm length and 0.8-1.5 mm diameter, secondary branches bearing 120+ tertiary branch rachides (third-order laterals) ranging 0.4-1.8 cm length and 0.4-0.8 mm diameter, complete rachis system exhibiting green coloration with moderate pubescence creating gray-green appearance, anatomical structure showing epidermis with scattered trichomes, collenchymatous cortex providing mechanical support, vascular ring containing 8-12 major bundles in main rachis reducing to 3-5 in primary branches and 1-2 in higher orders, rachis system bearing approximately 650 flowers distributed across the branching network, providing complete mechanical support for the inflorescence mass of approximately 15 grams and vascular capacity supporting maximum transpiration and photosynthesis, measured at BBCH stage 65 (full bloom) in mid-June, representing the complete hierarchical rachis architecture characteristic of compound paniculate grapevine inflorescences.
The main axis and branching axes of the grapevine inflorescence in Vitis vinifera, characterized by structural framework providing mechanical support and vascular connection for all flowers, hierarchical organization including central main rachis and multiple orders of lateral branch rachides, cylindrical to slightly flattened morphology typically 1-5 mm diameter depending on order and position, green coloration during flowering transitioning to brown woody appearance as cluster develops, anatomical composition including epidermis, cortex with collenchyma and parenchyma, vascular bundles arranged in ring providing water and nutrient delivery and photosynthate export pathways, variable pubescence from glabrous to densely tomentose depending on cultivar, and critical functional roles including mechanical support bearing the weight of flowers and subsequently developing berries, vascular transport providing the sole pathway for resource delivery and export, and developmental organization coordinating flower and berry positioning throughout the three-dimensional inflorescence structure. The inflorescence rachis constitutes the structural skeleton of the flowering and fruiting cluster, developing through apical and lateral meristem activity establishing the hierarchical branching architecture, exhibiting cultivar-specific characteristics including branching pattern, rachis thickness, and pubescence serving as ampelographic features, and transforming from the flowering rachis system to the cluster stem system following fruit set maintaining structural and vascular functions throughout berry development and maturation.
Inflorescence rachis encompasses all axis structures from the main central rachis through all orders of lateral branch rachides, distinguished from individual flowers or flower clusters which represent the terminal reproductive structures borne on the rachis framework. The rachis system exhibits hierarchical organization with clear size and complexity gradients from robust main rachis through progressively smaller higher-order branches. Developmental origin involves apical meristem establishing main rachis with lateral branch primordia initiating sequentially, lateral meristems elaborating branch orders, creating the complete architecture. Anatomical structure shows vascular bundles as the critical functional component, with bundle number and size correlating with rachis order and position reflecting vascular capacity requirements. Mechanical properties including strength, stiffness, and flexibility affect rachis function, with adequate strength preventing breakage under fruit load while some flexibility enables movement reducing wind stress. Pubescence variation from glabrous (hairless) to densely tomentose shows cultivar specificity serving as useful ampelographic character. Transformation during fruit development involves rachis lignification and browning creating the woody cluster stem, dimensional changes through elongation and thickening, and functional transition from supporting flowers to supporting substantially heavier berry mass. Rachis characteristics affect final cluster morphology including cluster length, width, compactness, and shape serving as important quality parameters. Management practices including girdling or other treatments may affect rachis development influencing cluster characteristics and berry size.
Blütenstandsachse
Infloreszenz-Rachis
Rispenspindel
cluster stem
inflorescence axis
panicle rachis
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 109-145 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 82-142 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 269-312.
Inflorescence Rachis
The inflorescence size of Grenache documents large inflorescences (OIV 151:7, 180-250 mm) with extensive branching producing 6-8 primary lateral branches (wings) and 600-1,200 flower primordia per inflorescence — this large inflorescence is the developmental precursor of Grenache's characteristically large, sometimes excessively compact clusters that can reach 300-500 g at harvest and are prone to internal berry compression and Botrytis development under humid conditions. The large inflorescence size contrasts with the small inflorescences of Pinot Noir (OIV 151:3, 80-120 mm, 200-400 flowers, producing the small 80-150 g clusters prized for quality production) and the medium inflorescences of Cabernet Sauvignon (OIV 151:5, 120-180 mm, 300-600 flowers, producing medium 120-200 g clusters with moderate compactness).
An inflorescence description component that records the dimensions of the fully developed pre-bloom inflorescence measured from the peduncle insertion to the tip of the terminal branch, scored according to OIV descriptor 151 on a scale from very small (state 1, 250 mm), reflecting the cultivar-specific branching complexity and axis elongation that predetermine the potential cluster size and yield per inflorescence.
Inflorescence_Size documents a character that links ampelographic description to viticultural yield management — inflorescence size is the primary genetic determinant of potential cluster size and therefore the cultivar-specific framework within which yield management decisions (cluster thinning, shoot thinning, pruning severity) are made. The inflorescence size is determined during two developmental phases — the initial primordium branching pattern is established within the dormant compound bud during bud organogenesis in the season preceding bloom (approximately June-August for the following year's crop), while the final inflorescence size is determined by the elongation and further branching of these primordia during the 4-6 weeks between budburst and bloom in the current season: both phases are subject to environmental modification (warm temperatures and adequate carbohydrate supply during bud organogenesis increase primordium branching, while favorable spring conditions promote inflorescence elongation), but the cultivar-specific baseline is genetically determined and maintains the relative ranking of cultivars across environments. The measurement is standardized at the full pre-bloom stage (E-L stage 17, individual flowers separated but calyptras still intact) when the inflorescence has reached maximum elongation before the cessation of rachis growth that occurs at bloom.
Blütenstandsgröße
Geschein-Länge
Infloreszenz-Dimensionserfassung
flower cluster length
inflorescence dimension record
pre-bloom cluster size
Inflorescence Size
Chardonnay in Champagne shows inflorescence visibility at BBCH 53 in early May Year N+1 when shoots have 3-4 unfolded leaves, with 2-3 mm pale green compact inflorescences becoming visible between expanding leaves within opened buds after accumulating 900-1000 chilling hours during winter and reaching 180-220 growing degree days base 10°C in spring under cool maritime conditions of 11-15°C mean temperature, enabling early fruitfulness assessment showing typically 2 inflorescences per shoot in this highly fruitful early-bursting cultivar
Inflorescence visibility represents the initial emergence phase when dormant inflorescence primordia first become externally observable within swelling buds following bud burst, marking definitive confirmation of successful inflorescence formation during previous season and winter survival, corresponding to BBCH 53 phenological stage. This critical observational milestone occurs during late April-early May in temperate regions shortly after bud burst when separating bud scales expose developing shoot contents including leaves and enclosed reproductive structures, providing first opportunity for assessing reproductive development status. Initial visibility shows small compact inflorescence structures 2-5 mm length appearing between expanding leaves within opening buds, with clusters showing dense undifferentiated form consisting of compressed rachis and tightly packed flower buds in early developmental state. Visibility timing relative to vegetative development typically occurs when shoot has 3-5 leaves unfolded corresponding to BBCH 13-15, with first inflorescence becoming visible at basal shoot positions as bud scales separate sufficiently to expose enclosed organs. Inflorescence appearance at visibility shows pale green coloration from enclosed development within bud lacking light exposure for chlorophyll synthesis, compact compressed structure from confined bud environment constraining expansion, and undifferentiated form with individual flowers not yet distinguishable requiring magnification for architectural details. Developmental state at visibility represents resumption from winter dormancy with meristematic tissues reactivating, cells beginning division and expansion driving initial enlargement, and metabolic processes resuming supporting growth though development remains early with substantial elaboration required before flowering. Observation requirements for reliable visibility assessment include careful inspection within opening buds as structures remain partially enclosed by leaves and scales, appropriate lighting enabling detection of small pale structures, and possible magnification for definitive identification distinguishing inflorescences from other bud contents. Inflorescence number assessment at visibility provides critical information about fruitfulness and potential yield, with count of visible inflorescences per shoot determining crop load potential guiding management decisions about thinning or crop adjustment. Visibility reliability depends on successful previous season primordium formation, adequate winter chilling satisfaction enabling normal dormancy release, winter survival without severe freeze damage, and appropriate spring conditions supporting development resumption. Abnormal visibility patterns indicate developmental problems with absence of expected inflorescences suggesting primordium formation failure during previous season, delayed visibility indicating inadequate chilling or cool spring conditions, or malformed structures suggesting developmental abnormalities or frost damage. Frost damage assessment at visibility enables early detection of injury with damaged inflorescences showing brown discoloration, necrotic tissues, or failure to resume growth, providing information for crop adjustment decisions. Developmental progression from visibility proceeds through continued enlargement and differentiation with inflorescences rapidly elongating and expanding over following 2-3 weeks, transitioning from compressed barely visible structures to recognizable clusters approaching flowering readiness. Metabolic state at visibility shows recently reactivated metabolism with respiration increasing from dormancy levels to 5-8 fold above baseline supporting resumed growth, protein synthesis recommencing producing enzymes and structural proteins, and mobilization of stored reserves from woody tissues providing energy and building materials. Resource dependence at visibility emphasizes stored carbohydrate importance with emergence drawing primarily on reserves accumulated during previous season, with inadequate reserves potentially limiting emergence vigor affecting subsequent development. Vascular connection at visibility maintains functional status with xylem conducting water from permanent structures through emerging shoot to developing inflorescences supporting turgor-driven growth, and phloem importing mobilized reserves and nutrients supporting rapid development. Hormonal status at visibility shows increasing cytokinin from root production promoting cell division and growth, moderate gibberellin supporting elongation processes, and auxin from developing organs coordinating growth and differentiation. Temperature during visibility period critically affects development rate with optimal 12-18°C supporting rapid visibility and continued development, cool conditions delaying emergence and progression, and warm conditions accelerating development advancing toward flowering. Light intensity during visibility affects overall vine vigor and resource availability indirectly though direct light effects on inflorescences are minimal at this early enclosed stage, with adequate light supporting vigorous shoot and inflorescence development. Water availability affects visibility through effects on cell turgor supporting expansion and overall vine function, with adequate moisture essential for normal resumption of growth enabling visibility. Protective measures during visibility period focus on frost protection as newly emerged tissues show high vulnerability to freeze damage with temperatures below -2°C causing tissue mortality, requiring active or passive protection in frost-prone regions. The process represents first of four distinguishable inflorescence emergence phases marking initial external appearance before subsequent elongation and expansion, providing critical early assessment opportunity. Duration from first visibility to transition toward elongation typically spans 3-7 days depending on temperature with warm conditions promoting rapid progression and cool conditions extending visibility period. Cultivar variation appears in visibility timing relative to bud burst with some varieties showing early inflorescence visibility while others show delayed appearance, and in ease of observation with compact varieties requiring careful inspection while loose-budded types show obvious visibility. Regional timing shows inflorescence visibility during late April-early May in warm climates, early-mid May in temperate regions, and mid-late May in cool areas, consistently occurring shortly after bud burst across climatic zones. Viticultural significance includes providing first definitive assessment of reproductive development status enabling crop load evaluation, representing critical observation point for fruitfulness assessment informing thinning decisions, enabling early detection of frost damage allowing timely crop adjustment, and confirming successful completion of previous season primordium formation and winter dormancy. Research applications include documenting precise emergence timing for phenological studies, evaluating environmental effects on dormancy release and development resumption, quantifying fruitfulness variation among cultivars and conditions, assessing relationships between visibility timing and subsequent flowering and yield, and developing prediction models for emergence timing supporting management decision-making.
Inflorescence_Visibility is first of four Inflorescence_Emergence subphases, distinguished by initial external appearance of inflorescences within opening buds. Corresponds to BBCH 53 phenological stage. Transitions rapidly to Inflorescence_Elongation as development proceeds. Critical observation point for crop assessment and management decisions. Vulnerable period for frost damage requiring protective measures in susceptible regions.
Blütenstandssichtbarkeit
Infloreszenzerscheinung
Traubenansatzerkennung
erstes Geschein sichtbar
first cluster visible
inflorescence appearance
inflorescence detection
Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 328-335
Inflorescence Visibility
In a Sonoma Pinot Noir vineyard, initial acid reduction was tracked over 8 days in mid-August Year N+1 (BBCH 83) when titratable acidity declined from 20.8 g/L to 16.2 g/L (22% reduction), malic acid decreased from 11.5 to 8.4 g/L (27% loss), tartaric acid stable at 7.8 g/L, pH increased from 3.02 to 3.18, berry Brix rose from 9.8° to 14.6°, VvME1 activity 28 units/g FW, under moderate coastal conditions (daily high 23-26°C, cool nights 12-15°C moderating decline).
Initial_Acid_Reduction represents the earliest phase of organic acid decline during veraison when malic acid begins active respiratory degradation while tartaric acid remains stable creating the beginning shift from very acidic green berry composition toward balanced ripe fruit suitable for quality wine production. This initial reduction occurs during early veraison (BBCH 83), typically 1-5 days following veraison onset signals, and is characterized by titratable acidity declining from 16-24 g/L to 13-20 g/L over 5-10 days representing initial 15-25% reduction from maximum, malic acid specifically decreasing from 9-14 g/L to 7-11 g/L through beginning respiratory metabolism (22-30% reduction) while tartaric acid remains essentially constant 5-10 g/L showing only 0-5% apparent change from dilution, berry pH increasing from 2.8-3.1 to 3.0-3.3 reflecting net acid loss, and malate:tartrate ratio declining from 1.3-2.2:1 to 1.0-1.8:1 indicating preferential malate consumption. Biochemical mechanisms of initial reduction inclu malic acid respiratory catabolism through multiple pathways with citric acid cycle oxidation consuming malate via malate dehydrogenase (malate → oxaloacetate) followed by citrate synthase, aconitase, isocitrate dehydrogenase producing CO₂ and reducing equivalents, NADP-malic enzyme directly decarboxylating malate producing pyruvate plus CO₂ plus NADPH used for biosynthesis or oxidized for energy, NAD-malic enzyme similarly converting malate to pyruvate with NAD⁺ reduction, and phosphoenolpyruvate carboxykinase potentially involved interconverting oxaloacetate and phosphoenolpyruvate though role in grape less defined than in other fruits. Enzyme activities during initial reduction show: NADP-malic enzyme activity increasing from pre-veraison 2-8 units/g fresh weight to initial reduction 12-35 units/g representing 3-6 fold enhancement enabling enhanced malate catabolism, NAD-malic enzyme showing 2-4 fold elevation from 1-5 units/g to 4-15 units/g, citric acid cycle enzyme activities (malate dehydrogenase, citrate synthase, isocitrate dehydrogenase) maintaining or slightly increasing supporting continued respiratory malate oxidation, and enzyme kinetics with Km values for malate 0.5-2.5 mM enabling efficient substrate utilization even as vacuolar malate concentrations decline from 100-150 mM to 70-110 mM. Gene expression patterns during initial reduction demonstrate: VvME1 (NADP-malic enzyme) transcript levels increasing 10-35 fold above green berry baseline with induction coinciding with veraison onset, VvME2 showing 8-25 fold elevation with slightly delayed kinetics, citric acid cycle enzyme genes generally maintaining expression with some (aconitase, isocitrate dehydrogenase) showing modest 2-5 fold increases, and regulatory control through ripening-specific transcription factors (possibly VvMYB, VvNAC families) coordinately activating multiple metabolic genes though specific factors controlling malic enzyme expression incompletely identified. Cellular compartmentation aspects inclu vacuolar malate representing bulk storage (70-110 mM during initial reduction) slowly declining as tonoplast transport enables cytoplasmic access, cytoplasmic malate 6-18 mM serving as immediate substrate for respiratory enzymes, mitochondrial malate uptake through dicarboxylate carriers enabling citric acid cycle oxidation, and metabolic channeling potentially coordinating malate transport and catabolism ensuring efficient carbon flow from storage to respiration. Respiratory quotient during initial reduction shows: RQ (CO₂ produced / O₂ consumed) typically 1.05-1.25 indicating mixed substrate oxidation with both hexoses and organic acids contributing to respiration, malate contribution to total berry respiration estimated 25-45% during initial reduction based on malate loss rates and CO₂ production, and respiratory rate per se 0.8-1.8 μmol O₂ g⁻¹ h⁻¹ maintaining pre-veraison levels despite beginning malate catabolism as sugar import begins providing alternative substrate. Coordination with sugar accumulation demonstrates: initial acid reduction temporally overlapping with initial sugar increase (both typically days 0-10 post-veraison onset) creating beginning development of sugar-acid balance, carbon budget showing net carbon import through sugar accumulation (120-200 mg glucose-equivalents g⁻¹ over 10 days) far exceeding carbon loss through acid decline (15-35 mg malate-equivalents g⁻¹) resulting in net berry carbon increase, and metabolic regulation balancing anabolic processes (sugar storage, biosynthesis) with catabolic processes (acid degradation, respiration) through unknown coordination mechanisms. Environmental sensitivity of initial reduction includes: temperature strongly affecting malate degradation with optimal 24-28°C supporting normal rates, warm conditions 28-32°C accelerating reduction 30-50% through enhanced respiratory rates potentially producing 25-35% malate loss versus 22-30% at optimal, heat stress >32°C dramatically accelerating loss 50-100% potentially causing excessive early decline, cool conditions 18-24°C slowing reduction 20-40% extending duration to achieve similar endpoint, and diurnal temperature effects with daytime warming promoting catabolism while nighttime cooling reduces losses creating cumulative effects dependent on temperature regime. Light and exposure effects operate primarily through berry temperature with sun-exposed clusters warming 4-10°C above air temperature accelerating malate catabolism 20-50% compared to shaded fruit, direct photosynthetic effects minimal as ripening berries have limited chlorophyll and photosynthetic capacity 3.3-3.5 shifts purple), proton balance changes as organic acid anions (malate²⁻) removed requiring compensatory ion adjustments, and carbon balance with malate catabolism providing 15-30% of berry respiratory substrate during initial reduction representing carbon cost of ripening. Quality implications inclu initial reduction establishing trajectory toward final acidity with rapid early loss potentially indicating excessive final decline problematic in warm regions, slow initial reduction in cool regions potentially indicating retention of high acids requiring extended ripening or winery intervention, and spatial variation across vineyard revealing maturity heterogeneity requiring selective management. Viticultural management during initial reduction involves: canopy management particularly in warm regions with strategic shading reducing cluster temperature 3-7°C slowing decline 25-40% preserving acids, irrigation management with adequate water potentially moderating berry temperature through transpirational cooling, avoiding severe water stress that may accelerate loss through stress-enhanced respiration, and monitoring decline rates through sampling enabling early detection of excessive loss allowing management intervention before irreversible. Analytical monitoring methods inclu titratable acidity measurement by NaOH titration to pH 8.2 endpoint providing total acid assessment, pH measurement showing inverse relationship with acid levels, HPLC organic acid analysis quantifying malate and tartrate separately enabling distinction of metabolic decline from dilution, enzymatic malate assays providing rapid field assessment, and sensory evaluation detecting decreasing sourness and developing balance important for practical maturity determination.
Initial_Acid_Reduction is distinguished from Acid_Decline_Onset by representing active reduction (15-25% TA decline, 22-30% malate loss) rather than onset signals, from complete acid decline by showing early phase rather than full reduction, and from pH increase by focusing on acid loss cause rather than pH effect. Duration 5-10 days BBCH 83. Malic acid primary component (22-30% reduction), tartaric stable (0-5% change). Temperature sensitive: 24-28°C optimal, >32°C excessive (50-100% acceleration). Initial_Acid_Reduction establishes trajectory toward final acidity critical for wine balance.
beginnende Malat-Reduktion
beginnender organischer Säure-Verlust
beginning organic acid loss
early acid decline phase
frühe Säure-Abnahme-Phase
nascent malate reduction
preliminary acidity decrease
vorläufige Azidität-Abnahme
Keller M (2020) The Science of Grapevines 3rd Ed pages 229-232 early acid decline Sweetman C, Sadras VO, Hancock RD, Soole KL, Ford CM (2014) Metabolic effects of elevated temperature on organic acid degradation in ripening Vitis vinifera fruit J Exp Bot 65:5975-5988 temperature effects on initial reduction Terrier N, Glissant D, Grimplet J, Barrieu F, Abbal P, Couture C, Ageorges A, Atanassova R, Léon C, Renaudin JP, Dédaldéchamp F, Romieu C, Delrot S, Hamdi S (2005) Isogene specific oligo arrays reveal multifaceted changes in gene expression during grape berry development Plant Physiol 138:1617-1633 gene expression during acid decline.
Initial Acid Reduction
In a Napa Valley Merlot vineyard, initial anthocyanin synthesis was measured 2 days before visible color in mid-August Year N+1 (BBCH 83 onset, 68 days post-flowering) when UFGT transcripts increased 115-fold, UFGT enzyme activity rose from undetectable to 12 units/kg, anthocyanin accumulation rate was 18 mg/kg per day increasing skin content from 6 mg/kg to 42 mg/kg over 48 hours, ABA levels were 880 ng/g, and berry sugar reached 9.8° Brix under optimal conditions (berry temperature 25-27°C with morning sun exposure).
Initial_Anthocyanin_Synthesis represents the earliest phase of de novo anthocyanin production during veraison when biosynthetic pathway genes are first upregulated and enzymatic machinery begins producing detectable quantities of anthocyanin-3-glucosides that will accumulate to create visible berry pigmentation. This process initiates 1-2 days before visible color change (BBCH 83 onset), triggered by convergent developmental and environmental signals, and is characterized by anthocyanin accumulation rates of 10-30 mg/kg skin fresh weight per day increasing berry skin anthocyanin content from pre-veraison baseline 32°C severely inhibits initial synthesis through transcript degradation, protein denaturation, and potential chromatin modifications suppressing gene accessibility reducing anthocyanin accumulation 50-80% compared to optimal temperatures, and recovery from heat stress requires 3-5 days for biosynthetic machinery re-establishment. Light quality effects on initial synthesis inclu UV-B radiation (280-315 nm) is particularly effective inducing anthocyanin biosynthetic genes through UVR8 photoreceptor signaling pathway with 30-60% enhancement of initial synthesis rates in UV-exposed versus UV-excluded berries, blue light (400-500 nm) absorbed by cryptochromes and phototropins stimulates anthocyanin genes through separate photoreceptor pathways, red light (600-700 nm) shows moderate effects possibly through phytochrome signaling, while far-red light (700-750 nm) can inhibit anthocyanin synthesis through phytochrome inactivation demonstrating light quality-dependent regulation beyond simple intensity effects. Water stress impacts on initial synthesis show: moderate deficit (leaf ψ -0.9 to -1.3 MPa) enhances initial synthesis 20-40% through elevated ABA signaling and potentially reduced berry size concentrating biosynthetic products, severe stress (<-1.5 MPa) impairs initial synthesis despite very high ABA levels through general metabolic limitation and potentially reduced substrate availability, while abundant water reduces initial synthesis through berry dilution and potentially suppressed ABA-dependent gene activation demonstrating complex water-anthocyanin relationships. Cultivar variation in initial synthesis rates reflects genetic differences in regulatory control: rapid synthesizers like Pinot Noir and Gamay accumulate 25-40 mg/kg per day during initial phase reaching visible color in 2-3 days, moderate cultivars like Merlot and Syrah show 15-25 mg/kg per day requiring 3-5 days to visible threshold, while slow synthesizers like Cabernet Sauvignon and Nebbiolo accumulate 10-20 mg/kg per day needing 4-7 days for color detection reflecting lower basal synthesis capacity or different regulatory thresholds. Coordination with other veraison processes is tight: initial anthocyanin synthesis typically precedes or coincides with berry softening onset (often synchronized within 1-2 days), occurs during early sugar accumulation acceleration (Brix rising 0.8-1.5% per week increasing to 1.5-3.0% per week), and parallels acid decline initiation (malate beginning respiratory degradation) demonstrating integrated ripening program activation. Metabolic demands of initial synthesis are substantial: anthocyanin production requires significant carbon allocation (5-15% of berry carbon import during early veraison directed to phenylpropanoid pathway), energy demands include ATP for biosynthetic reactions and vacuolar transport consuming 8-15% of berry respiration during peak synthesis, and nitrogen requirements for enzyme synthesis draw on berry amino acid pools particularly phenylalanine as direct precursor with 20-40 μmol phenylalanine channeled into anthocyanins per kg fresh weight during initial synthesis period.
Initial_Anthocyanin_Synthesis is distinguished from Color_Change_Onset by representing molecular and biochemical initiation rather than the broader onset process, from First_Color_Change by occurring 1-2 days before visible detection, and from Progressive_Synthesis by showing activation rather than sustained production. Accumulation rates: fast cultivars 25-40 mg/kg/day, moderate 15-25 mg/kg/day, slow 10-20 mg/kg/day. UFGT induction 80-150 fold is diagnostic. Temperature optimum 22-28°C berry surface. UV-B strongly enhances. Initial_Anthocyanin_Synthesis represents the critical molecular transition initiating veraison color development in red grape cultivars.
Biosynthese-Weg-Aktivierung
biosynthetic pathway activation
early anthocyanin production
erste Anthocyan-Akkumulation
first anthocyanin accumulation
frühe Anthocyan-Produktion
initial pigment biosynthesis
initiale Pigment-Biosynthese
Keller M (2020) The Science of Grapevines 3rd Ed pages 233-235 anthocyanin biosynthesis molecular control Boss PK, Davies C, Robinson SP (1996) Analysis of the expression of anthocyanin pathway genes in developing Vitis vinifera L. cv Shiraz grape berries Plant Physiol 111:1059-1066 gene expression kinetics Jeong ST, Goto-Yamamoto N, Kobayashi S, Esaka M (2004) Effects of plant hormones and shading on the accumulation of anthocyanins Plant Sci 167:247-252 hormonal regulation of synthesis initiation.
Initial Anthocyanin Synthesis
Cabernet Sauvignon in Napa Valley shows initial berry growth during mid July-early September Year N+1 at BBCH 71-75, with intensive cell division weeks 0-4 establishing 2.2-3.4 million mesocarp cells, rapid expansion weeks 3-6 increasing diameter from 14mm to 12-15mm final size, berry weight reaching 4.8-7.2g by week 7-8, under warm climate conditions of 24-32°C temperature, adequate irrigation supporting expansion, typical of this late-season large-berried cultivar
Initial berry growth represents the rapid early developmental phase following berry commitment when berries undergo intensive cell division establishing final cell number, transition to cell expansion-dominated growth, and achieve primary size establishment determining berry size potential, proceeding over 4-8 weeks from fruit set through lag phase onset and establishing foundation for subsequent ripening development. This critical early growth phase occurs during summer in July-August in temperate regions Year N+1, spanning approximately 4-8 weeks post-fertilization depending on cultivar and conditions, corresponding to BBCH 71-75 young fruit to pea-sized berry stages. Initial growth encompasses coordinated processes including early cell division growth with intensive mitotic activity producing berry flesh cells, rapid initial expansion with volumetric increase through cell enlargement and water uptake, berry size establishment determining final berry dimensions, and overall growth creating berry architecture capable of supporting ripening transformation. Growth dynamics show characteristic sigmoid pattern with rapid initial growth during cell division phase weeks 0-4, continued expansion during cell enlargement phase weeks 4-8, and eventual growth rate decline entering lag phase as berries approach final size. Cell division phase characteristics show peak mitotic activity during first 3-4 weeks post-fertilization, cell cycle times 10-15 hours during maximum division intensity, progressive cell number increase throughout mesocarp flesh tissue, and final cell number established by 3-4 weeks determining potential berry size. Cell number determination shows total mesocarp cells established during division phase, typical range 1-4 million cells per berry depending on cultivar and conditions, cell number as major determinant of final berry size with more cells enabling larger berries, and cell number influenced by seed number through auxin dose effects, temperature affecting division rate, and genetic programming. Cell expansion phase characteristics show transition from division to expansion-dominated growth around week 3-4, progressive cell enlargement through water uptake and vacuole expansion, cells increasing volume 10-100 fold from division completion to maturity, and expansion continuing through ripening though rate varying. Growth rate dynamics show maximum absolute growth rate during weeks 2-4 when both division and expansion active, peak relative growth rate often earlier during maximum division intensity, progressive growth rate decline as cell division completes, and eventually very low growth rate during lag phase before veraison. Berry size increase shows diameter expanding from 12-25 mm at commitment to 6-10 mm diameter in small-berried cultivars or 12-18 mm in large-berried cultivars by initial growth completion around week 6-8, volumetric increase 3-8 fold during initial growth phase, and size at lag phase entry largely predicting final berry size. Fresh weight accumulation shows berry weight increasing from 0.5-2 g at commitment to 2-8 g in small-berried or 5-15 g in large-berried cultivars by lag phase, weight accumulation rate 50-200 mg per day during peak growth, and fresh weight as convenient growth measurement. Dry matter accumulation shows progressive dry matter increase during growth, dry matter percentage relatively stable 12-18% during growth phase, and dry matter representing structural components and stored reserves. Water uptake characteristics show water as major component of berry expansion typically 80-85% fresh weight, xylem-mediated water import during early growth supporting rapid expansion, osmotic gradient from accumulating solutes driving water influx, and water relations critical for normal berry enlargement. Metabolic characteristics show high respiratory rate during active growth consuming 2-4% dry weight per day, carbohydrate import from vine providing energy and carbon skeletons, nitrogen uptake supporting protein synthesis for new cells and enzymes, and overall intensive metabolism supporting rapid growth. Resource requirements show berries consuming 40-60% total vine photosynthate during initial growth, competition between berry growth and vegetative growth affecting resource allocation, and adequate resource supply essential for achieving genetic size potential. Seed development coordination shows seeds developing in parallel with berry growth, seed size increase correlating with berry expansion, auxin production from seeds continuing to support berry growth though declining after peak weeks 2-3, and seed-berry coordination ensuring synchronized development. Environmental effects on initial growth include temperature critically affecting growth rate with optimal 22-30°C supporting maximum growth, water availability affecting cell expansion and final size, nutrient supply particularly nitrogen affecting cell division and growth capacity, light possibly affecting photosynthate supply though indirect effect, and overall favorable conditions maximizing size achievement. Temperature effects show warm conditions promoting rapid growth completing initial phase within 4-6 weeks, cool conditions extending initial growth to 6-10 weeks, and temperature as major factor affecting growth rate and size achievement. Water effects show adequate water essential for cell expansion particularly during rapid growth weeks 2-6, water deficit limiting berry enlargement potentially reducing final size 20-40%, and irrigation beneficial in water-limited environments during this critical period. Nutrient effects show nitrogen supporting cell division and protein synthesis, potassium affecting water relations and turgor, and overall balanced nutrition supporting optimal growth. Cultivar variation shows genetic differences in cell division capacity affecting cell number and size potential, growth rate variation affecting initial growth duration, and final size range from 6mm small berries to 18mm large berries reflecting genetic programming. The process shows three distinguishable phases progressing from early cell division growth during weeks 0-4 establishing cell number, through rapid initial expansion during weeks 3-6 combining division and expansion, to berry size establishment during weeks 5-8 completing initial growth. Duration typically 4-8 weeks from commitment through lag phase onset. Regional timing shows initial berry growth during July-August in warm climates, mid July-early September in temperate regions, and late July-mid September in cool areas. Viticultural significance includes establishing final cell number determining berry size potential, requiring optimal environmental conditions particularly temperature and water for maximum size achievement, demonstrating critical period for berry development affecting quality and yield, and completing berry structural development preparing for ripening transformation. Research applications include analyzing cell division and expansion dynamics, investigating environmental and genetic factors affecting growth and size, evaluating resource allocation and berry-vine competition, quantifying growth patterns and berry size determinants, and assessing initial growth as quality and yield determinant.
Initial_Berry_Growth represents critical early development phase within Fruit_Set establishing berry size. Subdivided into Early_Cell_Division_Growth establishing cell number, Rapid_Initial_Expansion combining division and expansion, and Berry_Size_Establishment completing growth. Duration 4-8 weeks post-commitment. Cell number determined in 3-4 weeks is major size determinant. Temperature and water strongly affect growth and final size. Completes berry structural development. Precedes lag phase and ripening. Determines berry size potential affecting yield and quality.
Beeren-Expansionsphase
Jungfrucht-Wachstum
berry expansion phase
early berry development
frühe Beerenentwicklung
initial berry enlargement
initiales Beerenwachstum
young fruit growth
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 430-445 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Initial Berry Growth
Merlot in Bordeaux shows initial berry softening during early September Year N+1 at BBCH 83, with firmness declining 16-22% from 3.4 to 2.7 N over 4-6 days at veraison onset, polygalacturonase activity increasing from 11 to 28 units, pectin methylesterase from 18 to 38 units, water-soluble pectin increasing from 24% to 42% total pectin, pectin molecular weight declining from 210 to 140 kDa, middle lamella beginning degradation weakening cell adhesion, possible minor turgor decline 10-12%, rapid initial softening 3.2% firmness per day, under oceanic conditions of 20-26°C temperature, typical of this mid-season cultivar initiating veraison texture transformation
Initial berry softening represents the first texture transformation phase during berry softening when berry firmness begins measurable decline at veraison onset, firmness declining 10-25% from pre-veraison maximum 2.5-4.5 N to initial softening 2.0-3.5 N, cell wall modification onset with pectin-degrading enzyme activation, pectin solubilization beginning with water-soluble pectin increasing, possible minor turgor reduction though structural changes primary, and first softening establishing veraison texture transformation over 3-7 days at transformation onset. This first softening phase occurs during late summer to early autumn in late August-early September in temperate regions Year N+1, representing veraison onset texture change. Initial softening mechanism involves cell wall modification onset with polygalacturonase and pectin methylesterase activation beginning pectin degradation, pectin solubilization start with insoluble pectin converting to soluble forms, middle lamella beginning degradation weakening cell adhesion, possible minor turgor pressure reduction though contribution less than structural changes, and rapid initial firmness decline creating detectable softness. Firmness decline characteristics shows firmness declining from pre-veraison maximum typically 2.5-4.5 N penetrometer resistance to initial softening 2.0-3.5 N representing 10-25% decline, typical decline 15-20% average, rapid decline over first 3-7 days veraison, and initial decline as first measurable softening. Pre-veraison firmness shows berry at maximum firmness during late lag phase, firmness from turgid cells with rigid cell walls, hard texture resistant to deformation, and peak firmness as pre-softening state. Firmness measurement shows penetrometer force recording objective firmness, measurements documenting initial decline, typical penetrometer probe 2-5 mm diameter, force in Newtons N, and measurement as quantitative assessment. Initial softening detection shows firmness decline detectable by instrumentation and trained sensory assessment, berries becoming noticeably softer to tactile evaluation, softening coinciding with or slightly following first visible color in red cultivars, and detection as softening indicator. Softening rate during initial phase shows rapid decline 2-4% firmness per day over initial 3-7 days, faster than subsequent ripening softening rate, rapid initial change characteristic of veraison transformation, and rate as initial phase feature. Cell wall modification onset shows pectin-degrading enzyme activation at veraison onset, enzymes including polygalacturonase PG and pectin methylesterase PME beginning activity increase, enzyme activation as softening mechanism, and enzymatic modification initiating structural change. Polygalacturonase activation shows PG activity increasing from low pre-veraison levels 5-15 units to initial softening 15-40 units, PG cleaving homogalacturonan pectin backbone, PG as major softening enzyme, and PG activation as critical event. PG expression and activity shows PG genes upregulated during late lag and veraison, PG protein accumulating, enzyme activity increasing substantially, and gene-protein-activity cascade as mechanism. Pectin methylesterase activation shows PME activity increasing from pre-veraison 10-25 units to initial 25-50 units, PME demethylating pectin creating PG substrate, PME as pectin modification enzyme, and PME activation enabling PG action. PME function shows PME removing methyl groups from pectin, demethylated pectin accessible to PG cleavage, PME possibly creating calcium cross-links initially before degradation, and PME as regulatory enzyme. Pectate lyase possible activation shows PL possibly beginning activity increase, PL contributing to pectin cleavage through different mechanism than PG, PL role variable, and PL as additional enzyme. Expansin possible role shows expansins possibly contributing to wall loosening, expansin expression possibly increasing, contribution less certain than pectinases, and expansins as potential wall modifiers. Pectin structural changes shows pectin molecular weight beginning decline from enzyme cleavage, initial depolymerization fragmenting pectin chains, molecular weight declining from pre-veraison 150-300 kDa to initial 100-200 kDa, and depolymerization initiating structural weakening. Pectin solubilization start shows water-soluble pectin fraction beginning increase, insoluble cell wall pectin declining, soluble pectin increasing from 20-30% to 35-50% total pectin during initial softening, and solubilization as degradation indicator. Soluble vs insoluble pectin shows soluble pectin as degradation products released from walls, insoluble pectin as structural wall-bound pectin, ratio shift indicating degradation degree, and solubilization weakening structure. Middle lamella beginning degradation shows pectin-rich middle lamella between cells beginning enzymatic attack, middle lamella degradation weakening cell-cell adhesion, adhesion loss contributing to softening, and middle lamella as critical target. Cell-cell adhesion beginning weakening shows intercellular bonding reduced from middle lamella pectin loss, cells becoming less firmly attached, tissue coherence declining, and adhesion weakening as softening mechanism. Turgor reduction possible initiation shows turgor pressure possibly declining slightly during initial softening, turgor changes minor compared to structural changes, turgor possibly 0.5-0.8 MPa pre-veraison declining to 0.4-0.7 MPa initial softening representing 10-15% reduction, and turgor as minor factor. Turgor role assessment shows structural wall changes primary softening cause contributing 70-85% firmness decline, turgor changes secondary contributing 15-30%, and structural dominance as softening basis. Water relations shows berry water content stable or slightly increasing, water uptake from sugar osmotic gradient, water affecting turgor but not primary softening driver, and water as background factor. Berry appearance shows berries becoming noticeably softer to touch, texture yielding more readily to pressure, visual appearance possibly showing slight gloss from surface changes, and appearance indicating softening. Tactile assessment shows trained evaluators detecting softness increase, berries less resistant to gentle compression, texture change perceptible, and tactile as practical evaluation. Softening timing shows initiation at veraison onset typically coinciding with or within 1-2 days of first color in red cultivars or concurrent with metabolic changes in white cultivars, timing marking veraison onset, and timing as transformation indicator. Softening coordination shows softening coordinated with other veraison changes including color development in red cultivars, sugar acceleration, acid decline, berry expansion, and coordinated transformation as integrated process. Environmental effects on initial softening include temperature affecting enzyme activation and activity with warm conditions promoting faster softening, water availability affecting turgor and berry hydration, and environmental factors influencing softening dynamics. Temperature effects shows warm conditions accelerating enzyme activities and firmness decline, cool conditions slowing initial softening, and temperature as rate factor. The process encompasses three concurrent initial softening aspects including cell wall modification onset with enzyme activation, pectin solubilization start with structural changes, and turgor reduction initiation though minor. Duration initial softening over 3-7 days representing first firmness decline at veraison. Regional timing shows initial berry softening during late August-early September at veraison onset. Viticultural significance includes representing first detectable veraison texture change, serving as universal veraison marker across cultivars, demonstrating cell wall enzyme activation as transformation mechanism, and determining veraison timing and progression. Research applications include analyzing softening onset mechanisms and regulation, investigating enzyme activation and pectin modification dynamics, evaluating firmness decline kinetics, quantifying structural and turgor changes, and assessing initial softening as veraison marker and quality determinant.
Initial_Berry_Softening is first phase of Berry_Softening, distinguished by first firmness decline. Represents 3-7 days at veraison onset. Firmness decline 10-25% (2.5-4.5 → 2.0-3.5 N). Subdivided into Cell_Wall_Modification_Onset (PG, PME activation), Pectin_Solubilization_Start (20-30% → 35-50% soluble), Turgor_Reduction_Initiation (minor, 10-15% decline). Rapid rate 2-4%/day. Universal veraison marker. Preceded by lag phase. Followed by Progressive_Softening. Temperature affects enzyme activation.
Veraison-Erweichungs-Beginn
anfängliche Beeren-Erweichung
early softening
erste Erweichung
first softening
initial firmness decline
initiale Festigkeits-Abnahme
veraison softening onset
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 475-482 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Initial Berry Softening
Sauvignon Blanc in Marlborough shows initial bud breaking at BBCH 07 in early October (Southern Hemisphere) after 950-1050 chilling hours and 280-330 growing degree days base 10°C, with 2-3 mm pale green shoot tip visible through separated scales over 2-3 days under conditions of 14-17°C mean temperature and 12.5-hour photoperiod, representing typical cool maritime climate pattern for this early-to-mid season cultivar
Initial bud breaking represents the first emergence phase where green shoot tissue begins appearing through maximally separated bud scales, corresponding to BBCH 07 as the definitive marker of dormancy exit and active shoot development initiation. This critical developmental stage occurs when expanding shoot primordium reaches sufficient size and mechanical force to breach protective scale barrier, with green tip first becoming visible at bud apex where scale separation is widest. The emerging shoot tip measures 2-4 mm in visible length extending beyond scale margins, appearing pale yellow-green due to limited chlorophyll synthesis in tissues previously enclosed within protective scales. At this initial phase, shoot tissue remains largely covered by scales with only apical 10-30% exposed, creating characteristic "green tip" appearance that gives this stage its descriptive terminology in both scientific and viticultural contexts. The shoot primordium beneath scales has reached 5-8 mm total length with apical meristem actively producing new cells through divisions every 18-24 hours, while cells in elongation zone behind meristem expand rapidly through turgor pressure of 0.5-0.8 MPa driving observable emergence. First leaf primordia measure 3-5 mm length remaining tightly folded against shoot axis within protective scales, not yet visible externally but prepared for rapid expansion once shoot elongates further. Bud scales reach maximum separation at tips with gaps of 2-3 mm, maintaining structural connection at base while being mechanically forced apart by internal tissue pressure rather than actively growing or dehiscing. The mechanical force driving emergence results from coordinated cell expansion through osmotic water uptake, with shoot cells importing solutes including sugars mobilized from cane storage reserves and ions transported through developing vascular system. Vascular connection between emerging shoot and permanent woody structures achieves full functionality with mature xylem vessels conducting water at rates of 0.5-1.5 mg per second to support expanding tissues and beginning transpiration from exposed surfaces. Chloroplast development initiates in emerging shoot cells with chlorophyll synthesis beginning though photosynthetic capacity remains negligible, requiring several days and additional leaf area before net carbon fixation approaches respiration losses. Metabolic activity remains at peak levels with respiration consuming stored carbohydrates at 15-20 fold above winter baseline, supporting rapid cell division and expansion before photosynthetic capacity develops. Protein synthesis continues at maximum rates producing structural proteins for new cells, enzymes supporting growth metabolism, and regulatory proteins coordinating developmental progression toward shoot elongation and leaf expansion. Hormonal regulation maintains high cytokinin and gibberellin concentrations promoting cell production and expansion, with auxin synthesis in shoot apex establishing polarity gradients coordinating upward shoot growth and suppressing lateral bud activation. Temperature sensitivity persists with development rate doubling per 10°C increase between 10-25°C, making emergence timing and rate highly responsive to spring temperature conditions. Water requirement increases substantially as exposed tissues begin transpiring, requiring functional xylem to transport water from roots through cane to emerging shoot to maintain turgor and prevent wilting. Light exposure affects emerging tissues with chlorophyll synthesis responding to light intensity and quality, though full photosynthetic capacity requires additional development time and leaf expansion. The process typically requires 2-4 days from first green tip visibility to progression toward active breaking under favorable spring conditions, with cooler temperatures extending duration to 4-6 days. Cultivar variation in initial breaking appears in shoot emergence rate and extent, with vigorous varieties showing rapid 1-2 mm per day emergence while less vigorous types emerge 0.5-1.0 mm per day. Regional timing follows late bud swelling by 2-4 days, occurring mid-April in Mediterranean climates, late April in temperate areas, and early May in cool continental regions. Viticultural significance includes marking the definitive point when dormant-season operations must cease to avoid damage, establishing critical reference for deploying frost protection as tissues reach maximum vulnerability to freeze damage, serving as developmental marker for initiating pest and disease management targeting newly available host tissues, and providing standardized phenological reference point for regional comparisons and historical tracking. Frost damage at initial breaking causes immediate tissue death through ice formation in exposed shoot tip and enclosed developing leaves, killing primary bud and requiring compensatory growth from secondary buds with 2-3 week delay and substantial fruitfulness reduction. Research applications include precise phenological monitoring for climate studies, molecular profiling of genes regulating shoot emergence and meristem activity, microscopic analysis of cellular differentiation during organ formation, and thermal time modeling for predicting breaking timing across diverse climatic conditions and future climate scenarios.
Initial_Bud_Breaking is the first of three Bud_Breaking subphases, distinguished by first appearance of green tissue through scales with <30% shoot exposed. Transitions to Active_Bud_Breaking as shoot elongates and more tissue emerges. Represents most standardized phenological observation point (BBCH 07) for regional comparisons and historical records. Maximum frost vulnerability point.
beginnender Austrieb
beginning bud break
erste Triebspitze
first shoot emergence
initiales Knospenaufbrechen
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 175-180 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Buttrose MS (1974) Climatic factors and fruitfulness in grapevines, Hort Abstr 44:319-325
Initial Bud Breaking
Chardonnay in Champagne shows initial cap fall during early June Year N+1 at BBCH 61-62 over 2-3 days June 3-5, with 10-18% flowers predominantly in apical positions shedding calyptra caps, exposing reproductive organs and marking flowering onset, under cool maritime conditions of 18-22°C temperature, 60-72% relative humidity, typical of this early-bursting cultivar initiating flowering before peak bloom June 6-9
Initial cap fall represents the beginning phase of calyptra detachment when first flowers complete abscission and shed petal caps exposing reproductive organs, marking definitive flowering onset and establishing temporal pattern for subsequent mass cap fall during peak anthesis. This critical initiation phase occurs during early flowering in early June in temperate regions Year N+1, typically spanning 2-3 days as earliest-maturing flowers predominantly in apical cluster positions complete cap detachment, corresponding to BBCH 61-62 stages with 10-20% flowers showing cap fall. Cap detachment initiation in earliest flowers shows characteristic sequence with abscission zone activation in calyptra base beginning enzymatic wall degradation, progressive calyptra loosening creating visible gap at base, upward cap movement driven by expanding reproductive organs, and final detachment with cap falling away from flower through mechanical disturbance or gravity. First calyptra detachment typically occurs in apical cluster positions reflecting earliest primordium formation and most advanced development from previous season, with 5-10% flowers showing cap fall on first day establishing anthesis onset, and progressive increase to 15-25% by end of initial phase expanding flowering across cluster. Detached cap appearance shows intact fused petal structure maintaining pentamerous organization with five coherent petal sections visible, clean separation scar at base indicating completed abscission, occasional adherence to adjacent structures requiring additional mechanical force for complete removal, and accumulation on cluster surface providing visible evidence of flowering initiation. Flower exposure following cap detachment reveals functional reproductive structures with stamens and pistil newly exposed showing pristine fresh appearance, anthers beginning color change from green toward yellow-brown indicating approaching maturity, stigma surface developing papillate cells and initial exudate production, and overall flower transitioning to open reproductive state. Temporal pattern establishment during initial phase shows characteristic basipetal-acropetal gradient with apical flowers showing earliest cap fall, progressive extension to middle regions over subsequent days, and basal regions showing delayed cap fall establishing developmental sequence. Cap fall frequency during initial phase shows sporadic irregular pattern with individual flowers completing detachment asynchronously, daily increment of 3-8% additional flowers shedding caps, and progressive acceleration toward mass cap fall during subsequent peak period. Environmental sensitivity during initial phase shows strong dependence on conditions with favorable warm dry weather promoting efficient cap detachment supporting early flowering onset, unfavorable cool humid conditions delaying detachment potentially causing irregular flowering initiation, and optimal 20-26°C temperature with 50-70% humidity supporting normal cap fall progression. Humidity effects on initial cap fall show critical importance with moderate levels facilitating abscission through appropriate tissue hydration and enzymatic activity, high humidity above 75% substantially interfering with detachment through maintained cap turgor preventing lifting and possibly inhibiting enzymatic processes, and resulting cap retention delaying flowering and potentially causing coulure. Temperature during initial phase affects detachment rate with optimal 22-28°C supporting rapid normal abscission, cool conditions below 18°C substantially slowing enzymatic activity delaying cap fall and flowering onset, and warm conditions above 30°C potentially accelerating processes though risking tissue stress. Cap retention problems may appear during initial phase with some flowers showing incomplete abscission leaving caps partially attached, problematic retention indicating unfavorable conditions or developmental abnormalities, and early retention problems potentially portending difficulties during peak flowering affecting ultimate fruit set. Abscission zone development during initial phase shows enzymatic activity concentrated in specialized cells at calyptra-receptacle junction, cellulase and pectinase enzymes degrading middle lamella polysaccharides creating separation plane, progressive wall breakdown enabling cap loosening and detachment, and hormonal regulation possibly involving ethylene promoting abscission though mechanisms incompletely understood. Mechanical forces contributing to detachment include expanding stamens pushing against enclosing calyptra creating upward pressure, pistil elongation adding to mechanical tension, turgor changes potentially affecting cap rigidity and separation forces, and external forces from wind or physical contact triggering final detachment once abscission completes. Observational characteristics enable identification of initial cap fall through fresh appearance of detached caps showing recent separation, spatial concentration in apical cluster positions, small proportion 10-20% flowers showing cap fall distinguishing from peak period, and clear visual evidence of flowering initiation. Phenological significance includes marking definitive flowering onset establishing BBCH 61 stage, providing practical observable reference for flowering date determination easier than detailed reproductive organ examination, and establishing baseline for predicting subsequent peak flowering timing typically 2-4 days later. Management implications include confirming flowering initiation requiring weather monitoring and preparation for critical bloom period, final opportunity for pre-bloom disease management as restrictions begin with significant flowering, and indicating imminent peak flowering requiring operational readiness. The process shows two distinguishable aspects progressing from first calyptra detachment establishing definitive flowering onset, through early cap shedding expanding activity across cluster. Duration typically spans 2-3 days representing initiation period before peak cap fall, with warm favorable conditions promoting rapid progression while cool variable conditions may extend period. Cultivar variation in initial cap fall shows differences in timing with early varieties showing late May-early June onset while late types show mid-late June timing, detachment efficiency with some varieties showing clean early cap fall while others show retention problems, and proportion of flowers involved in initial phase affecting subsequent mass fall intensity. Regional timing shows initial cap fall during late May-early June in warm climates at approximately 950-1050 growing degree days base 10°C, early-mid June in temperate regions at 850-950 GDD, and mid-late June in cool areas at 750-850 GDD, marking flowering onset across regions. Viticultural significance includes providing clear practical marker for flowering onset enabling accurate phenological tracking, indicating beginning of critical weather-sensitive period requiring monitoring, establishing temporal reference for subsequent flowering progression, and revealing early problems with cap retention indicating potential fruit set difficulties. Research applications include documenting precise flowering onset timing using cap fall as objective marker, analyzing environmental triggers of cap abscission affecting flowering initiation, evaluating cultivar differences in cap detachment efficiency affecting reproductive success, quantifying relationships between initial cap fall patterns and subsequent flowering progression, and assessing abscission mechanisms coordinating calyptra detachment with reproductive organ maturation.
Initial_Cap_Fall is first major Cap_Fall phase distinguished by flowering onset with early sporadic cap detachment. Corresponds to BBCH 61-62 parallel to Early_Anthesis. Subdivided into First_Calyptra_Detachment marking definitive onset and Early_Cap_Shedding expanding activity. Establishes temporal pattern for subsequent mass fall. Highly sensitive to environmental conditions particularly humidity. Serves as practical observable flowering initiation marker. Critical for phenological tracking and management timing.
early cap fall
erste Blütenkappentrennung
first cap shedding
früher Kappenfall
initial calyptra detachment
initiale Kalyptralösung
Initial Cap Fall
Syrah in Rhône Valley shows initial cap shedding during early June Year N+1 at BBCH 61-62 over 2-3 days June 4-6, with flowers progressively opening from initial 6-8% to 18-24% as caps detach exposing reproductive organs, showing clean efficient shedding under warm Mediterranean conditions of 23-28°C temperature, 50-65% relative humidity, light morning winds assisting detachment, typical of this mid-to-late season cultivar showing rapid early anthesis progression
Initial cap shedding represents the progressive calyptra detachment process during early anthesis where increasing numbers of flowers complete petal cap abscission exposing reproductive organs, expanding opening from initial 5-10% to 15-25% flowers open by end of early anthesis period. This critical expansion phase occurs during early June in temperate regions Year N+1 spanning 2-3 days following first flower opening, as accelerating cap shedding creates increasing numbers of functional flowers preparing foundation for subsequent peak anthesis. Cap shedding progression shows characteristic pattern with increasing frequency of detachment events as more flowers reach appropriate maturity state, additional flowers throughout cluster particularly in middle regions completing abscission following initial apical opening, and daily increments of 3-8% additional flowers opening creating progressive expansion of bloom. Shedding mechanism involves abscission zone activation in calyptra base where specialized cells undergo programmed wall degradation, enzymatic breakdown of middle lamella through cellulase and pectinase activity creating separation plane, mechanical forces from expanding reproductive organs and turgor changes driving physical detachment, and final cap separation through combined enzymatic and mechanical processes. Cap lifting appearance shows characteristic sequence with initial loosening visible as narrow gap at base, progressive upward movement as abscission proceeds with cap elevating 0.5-2.0 mm above receptacle, continued lifting driven by expanding stamens and pistil pushing upward, and final detachment with cap falling away or sometimes remaining temporarily caught on adjacent structures. Detached cap characteristics show intact fused petal structure maintaining pentamerous form with five coherent petals visible as sections, attachment scar at base where abscission occurred showing clean separation plane, and occasional caps showing incomplete separation with partial retention requiring additional mechanical force for complete removal. Cap retention problems in some flowers show incomplete abscission with caps remaining partially attached, physical impediment preventing normal reproductive organ function affecting pollination and fruit set potential, with retention associated with unfavorable conditions including high humidity, cool temperature, or developmental abnormalities. Abscission zone anatomy shows specialized cell layers at calyptra-receptacle junction with smaller cells containing dense cytoplasm, thin walls particularly in middle lamella region susceptible to enzymatic degradation, and vascular traces disrupting at separation plane requiring coordinated breakdown. Enzymatic activity during shedding involves cellulase enzymes degrading cellulose components of cell walls, pectinase and polygalacturonase enzymes breaking down pectin in middle lamella, and possible xyloglucan endotransglycosylase activity modifying wall polysaccharides, creating coordinated wall dissolution enabling separation. Hormonal regulation of cap shedding involves ethylene potentially triggering or coordinating abscission with increased ethylene biosynthesis and sensitivity in abscission zone, auxin possibly regulating timing with declining auxin levels permitting abscission initiation, and abscisic acid potentially involved in coordinating developmental timing. Reproductive organ exposure following cap shedding reveals functional structures with stamens extending upward surrounding central pistil, anthers positioned at various angles typically 0.5-1.5 mm above stigma facilitating pollen release and capture, and pistil with bilobed stigma surface positioned to receive pollen from anthers or wind dispersal. Flower appearance during initial shedding shows increasing proportion of opened flowers contrasting with remaining unopened buds, creating characteristic mixed appearance with opened flowers showing yellow-brown anthers against green receptacle and unopened flowers showing green calyptra caps, and cluster taking on progressively more open appearance. Temporal pattern of shedding shows diurnal rhythm with most cap detachment occurring during morning hours as temperature rises and humidity declines creating favorable conditions, reduced shedding during afternoon and evening hours, and resumption following morning creating daily waves of opening. Environmental effects on shedding include temperature strongly affecting rate with optimal 22-28°C promoting rapid shedding while cool conditions below 15°C substantially delay shedding and warm conditions above 32°C may cause premature or incomplete shedding. Humidity critically affects shedding with moderate 45-65% relative humidity optimal for clean complete cap detachment, high humidity above 75% interfering with shedding through maintaining high turgor and possibly affecting enzymatic activity, and low humidity below 35% causing rapid desiccation potentially affecting tissue function though facilitating mechanical separation. Rain during initial shedding represents serious concern with water preventing normal cap detachment through elevated humidity and physical interference, causing caps to remain attached or shed incompletely affecting reproductive function, and promoting fungal infection on wet flowers particularly botrytis establishing latent infections. Wind effects show modest positive role with light air movement potentially assisting final mechanical cap detachment and removing detached caps from cluster vicinity, though excessive wind may cause damage to newly exposed delicate reproductive organs. Cluster position effects show well-exposed clusters typically showing earlier more uniform shedding while shaded interior clusters may show delayed or irregular shedding, creating within-vineyard spatial variation in flowering progression affecting fruit set uniformity. Developmental uniformity affects shedding pattern with highly synchronized flowers showing compressed rapid shedding over 2-3 days, while heterogeneous development extends shedding period to 4-6 days during early anthesis phase, affecting overall flowering duration and fruit set outcomes. Metabolic activity during shedding shows elevated respiration in abscission zone supporting enzymatic processes, active carbohydrate metabolism providing energy for controlled cell separation, and maintained metabolism in newly exposed reproductive organs supporting immediate function. Resource requirements during shedding emphasize carbohydrate demand for energy-intensive abscission and organ function, nitrogen for enzyme synthesis supporting abscission processes, and minerals particularly calcium affecting cell wall properties and separation mechanics. The process represents second of three early anthesis aspects focusing on progressive calyptra removal, following first flowers opening and overlapping with stigma receptivity onset as exposed organs develop function. Duration spans 2-3 days during early anthesis representing expansion period from initial 5-10% to 15-25% flowers open, with warm favorable conditions promoting rapid progression. Cultivar variation appears in shedding efficiency with some varieties showing clean complete cap detachment while others show frequent retention problems affecting fruit set, shedding rate affecting early anthesis duration, and sensitivity to environmental conditions affecting shedding under marginal conditions. Regional timing shows initial cap shedding during early June in warm climates, early-mid June in temperate regions, and mid-June in cool areas, occurring during early anthesis phase 0-2 days after first flowers opening. Viticultural significance includes expanding reproductive function within cluster increasing pollination opportunity, revealing developmental uniformity affecting subsequent peak anthesis predictions, establishing period of heightened weather sensitivity requiring monitoring, and providing visual indicator of flowering progression enabling management assessment. Research applications include analyzing mechanisms regulating cap abscission including enzymatic and hormonal controls, evaluating environmental effects on shedding efficiency affecting reproductive success, quantifying developmental uniformity implications for shedding patterns, investigating causes of cap retention problems affecting fruit set, and assessing genetic variation in shedding characteristics affecting flowering reliability.
Initial_Cap_Shedding is second of three Early_Anthesis aspects, distinguished by progressive calyptra detachment expanding bloom from initial opening. Follows First_Flowers_Opening. Overlaps with Stigma_Receptivity_Onset. Represents expansion phase increasing reproductive function within cluster. Efficiency critically affected by environmental conditions particularly humidity. Problems with retention indicate unfavorable conditions affecting fruit set potential.
beginnende Kalyptraablösung
calyptra shedding
früher Blütenkappenfall
progressive Kappenabwurf
progressive cap detachment
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 365-370 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 278-282
Initial Cap Shedding
Chardonnay in Champagne shows initial cell enlargement during mid-late July Year N+1 at BBCH 73-75, with turgor pressure developing 0.4-0.7 MPa, water uptake 0.3-0.6 g per berry per day, vacuoles expanding from 25% to 58% cell volume, expansin-mediated wall loosening, cells enlarging 3.2-fold from 30 to 48 micrometers, berry diameter increasing from 16 to 21 mm, under cool maritime conditions of 20-25°C temperature with adequate rainfall, typical of this early-bursting cultivar initiating enlargement
Initial cell enlargement represents the expansion onset phase during berry cell enlargement when post-division cells begin volume increase through water uptake, turgor pressure develops driving expansion, vacuole formation initiates creating osmotic compartment, cell wall loosening enables growth, and coordinated processes establish enlargement as growth mechanism, proceeding over weeks 3-5 post-fertilization during transition from division-dominated to expansion-dominated growth. This expansion initiation phase occurs during mid-late summer in mid-late July in temperate regions Year N+1, spanning approximately 2-3 weeks representing enlargement onset overlapping with declining division, corresponding to BBCH 73-75 pea-sized berry to berry touch stages. Initial enlargement mechanism involves turgor-driven expansion onset with osmotic water uptake generating pressure, vacuole enlargement initiation with small vacuoles expanding and coalescing, cell wall loosening enabling turgor-driven growth, coordinated processes creating initial 2-5 fold cell volume increase, and establishment of expansion as active growth mechanism. Turgor-driven expansion fundamentals show osmotic water uptake generating turgor pressure 0.3-0.8 MPa during initial enlargement, pressure force pushing against cell walls causing expansion, yielding walls enabling growth under turgor, and turgor as physical driving force for volume increase. Water uptake initiation shows water import beginning in cells exiting division cycle, osmotic gradient from accumulating solutes driving water influx, initial water uptake rates 0.2-0.4 g per berry per day early phase increasing to 0.5-0.8 g per day late initial phase, and progressive water uptake acceleration as more cells enlarge. Osmotic gradient development shows solute accumulation creating osmotic potential decline from -0.5 MPa post-division to -0.7 to -0.9 MPa during initial enlargement, organic acid synthesis particularly malate increasing from 0.5% to 1.0% fresh weight, potassium import contributing to osmotic gradient, sugars still low 3-5% from continued metabolic consumption, and combined solutes creating gradient pulling water into cells. Malate synthesis shows malic acid as major organic acid synthesized during early enlargement, synthesis from glycolysis and TCA cycle intermediates, malate accumulation in vacuole creating osmotic potential and lowering pH, and malate as primary osmolyte during enlargement phase. Potassium accumulation shows K+ import through plasma membrane and tonoplast transporters, potassium accumulation to 150-250 mmol/L in berry tissue, K+ as major cation balancing organic acid anions, and potassium contributing substantially to osmotic gradient. Vacuole enlargement initiation shows small vacuoles 10-30% cell volume in post-division cells, vacuole membrane synthesis and water influx causing expansion, multiple small vacuoles coalescing into fewer larger vacuoles, progressive vacuole enlargement to 50-70% cell volume by initial enlargement completion, and vacuole as major expanding compartment. Vacuole membrane synthesis shows tonoplast proliferation accommodating vacuole expansion, membrane lipid and protein synthesis, proton pump and transporter insertion regulating vacuole, and membrane synthesis coordinated with vacuole growth. Vacuole function establishment shows compartment storing accumulated solutes and water, tonoplast H+-ATPase acidifying vacuole to pH 3.0-3.5, transporters accumulating malate, tartrate, K+, sugars into vacuole, and vacuole as osmotic engine and storage depot. Cell wall loosening initiation shows expansin expression increasing during initial enlargement, expansin proteins secreted to apoplast binding cell walls, expansin-mediated wall loosening disrupting cellulose-hemicellulose hydrogen bonds, wall plasticity increase enabling turgor-driven expansion, and loosening as essential enabling mechanism. Expansin characteristics show α-expansin family members expressed in expanding berries, pH-dependent activity optimal at acidic pH 4.5-5.5 characteristic of apoplast, non-hydrolytic wall loosening mechanism, and expansin as key wall-modifying enzyme during expansion. Cell wall yielding characteristics show wall capable of plastic deformation under turgor pressure, yielding walls enabling irreversible cell expansion, wall stress relaxation allowing sustained growth, and yielding balanced with synthesis preventing structural failure. Wall synthesis during initial enlargement shows new cellulose microfibrils deposited maintaining wall thickness, hemicellulose and pectin matrix synthesis, coordinated synthesis with loosening maintaining integrity, and wall deposition preventing excessive thinning during area expansion. Cell volume increase during initial enlargement shows typical 2-5 fold volume increase from post-division 25-45 micrometers diameter to 40-70 micrometers by initial phase end, progressive enlargement as cells accumulate water, initial expansion establishing enlargement trajectory, and volume increase contributing to berry growth. Cell enlargement spatial distribution shows enlargement initiating first in early-division-exiting cells week 3, progressive enlargement onset across cell populations weeks 3-5 as cells exit division, and asynchronous enlargement creating temporal spread. Enlargement-division overlap shows some cells still dividing while others enlarging during initial enlargement weeks 3-5, declining division concurrent with increasing enlargement, overlap maintaining berry growth rate during transition, and eventual pure enlargement as division completes. Berry size increase during initial enlargement shows diameter expanding from 14-20 mm at onset to 18-25 mm by initial phase end, volumetric increase 1.5-2.5 fold during initial enlargement, fresh weight increasing from 4-8 g to 6-12 g, and substantial growth establishing expansion as effective mechanism. Berry growth rate shows increasing rate during initial enlargement from 50-80 mg per day early to 80-120 mg per day late initial phase, growth rate increase reflecting accelerating expansion as more cells enlarge, and rate progression toward peak during rapid expansion. Berry composition during initial enlargement shows water content 80-85% fresh weight maintained, organic acids increasing from 0.8% to 1.2%, sugars 3-5% remaining low, pH declining from 3.0 to 2.9 from acid accumulation, and green acidic composition characteristic. Environmental effects on initial enlargement include temperature affecting expansion rate with optimal 22-28°C supporting rapid enlargement onset, water availability affecting turgor maintenance and expansion with adequate water essential, deficit potentially limiting initial enlargement reducing final size, and overall favorable conditions supporting enlargement initiation. Water effects show water as critical resource during initial enlargement, adequate water enabling full turgor development and expansion, deficit limiting turgor and expansion even during initiation, and early deficit effects potentially irreversible through reduced achieved size. Temperature effects show warm conditions promoting faster expansin expression and wall modification, cool conditions slowing initial enlargement though not preventing, and temperature affecting enlargement rate and timing. The process shows three distinguishable aspects including turgor-driven expansion onset establishing pressure-driven growth, vacuole enlargement initiation creating osmotic compartment, and cell wall loosening enabling yielding growth. Duration typically 2-3 weeks representing enlargement onset weeks 3-5. Regional timing shows initial cell enlargement during mid-late July across climates as expansion initiates. Viticultural significance includes representing transition to expansion as growth mechanism, establishing turgor-driven growth system, demonstrating water sensitivity during critical initiation period, and initiating size increase determining berry developmental trajectory. Research applications include analyzing turgor development and water relations, investigating vacuole formation and function, evaluating wall loosening mechanisms and expansin regulation, quantifying initial expansion dynamics, and assessing initial enlargement as determinant of subsequent expansion and final size.
Initial_Cell_Enlargement is first phase of Berry_Cell_Enlargement_Phase, distinguished by expansion onset. Subdivided into Turgor_Driven_Expansion_Onset establishing pressure system, Vacuole_Enlargement_Initiation creating osmotic compartment, and Cell_Wall_Loosening enabling growth. Duration 2-3 weeks representing weeks 3-5. Overlaps with declining division. Cell volume increase 2-5 fold. Turgor 0.3-0.8 MPa. Vacuole 50-70% volume by phase end. Water uptake 0.2-0.8 g/berry/day. Precedes Rapid_Cell_Expansion. Critical establishment phase for expansion-driven growth.
Expansions-Beginn
Vergrößerungs-Initiierung
early cell growth
enlargement initiation
expansion onset
frühes Zellwachstum
initial expansion phase
initiale Zellvergrößerung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 442-446 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Initial Cell Enlargement
Syrah in Rhône Valley shows initial division activation during late June Year N+1 at BBCH 71, with auxin from 1.8-2.2 developing seeds reaching 120-180 ng/g in mesocarp days 3-6 post-fertilization, auxin signaling triggering cell cycle gene expression, G0 to G1 transition in pericarp cells, progressive activation wave completing by day 7, under warm Mediterranean conditions of 25-29°C temperature, typical of this mid-to-late season cultivar establishing division competence within first week
Initial division activation represents the cell cycle entry phase during active berry cell division when auxin signaling from developing seeds triggers quiescent pericarp cells to exit G0 arrest, enter G1 phase, and prepare for DNA replication, establishing proliferative competence enabling subsequent mitotic divisions, occurring within first week post-fertilization as hormone stimulus establishes. This critical activation phase occurs during early summer in late June-early July in temperate regions Year N+1, typically days 2-7 post-fertilization as auxin reaches threshold levels, corresponding to BBCH 71 young fruit earliest stage. Division activation mechanism involves auxin perception by pericarp cells through TIR1/AFB auxin receptors, auxin-receptor complex triggering ubiquitin-mediated degradation of Aux/IAA repressor proteins, release of ARF transcription factors enabling cell cycle gene expression, transcriptional activation of cyclins, cyclin-dependent kinases, and other cell cycle regulators, G0 to G1 transition enabling growth and DNA replication preparation, and establishment of division-competent state. Auxin signaling pathway shows auxin from developing seeds transported to pericarp tissue, hormone accumulation reaching threshold levels 50-200 ng/g fresh weight triggering response, auxin binding to TIR1/AFB receptor proteins in nucleus, and receptor-mediated signaling cascade activating downstream responses. Auxin perception in pericarp cells shows receptor expression in mesocarp and exocarp tissues, spatially distributed receptors enabling tissue-wide response, auxin sensitivity variation among cell types possibly affecting activation timing, and receptor-hormone binding initiating signaling cascade. Aux/IAA repressor degradation shows auxin-receptor complex recruiting Aux/IAA proteins, ubiquitin ligase marking repressors for degradation, proteasomal degradation removing repression, and derepression enabling ARF transcription factor activity. ARF transcription factor activation shows ARF proteins released from repression binding auxin response elements in target gene promoters, transcriptional activation of cell cycle genes, and coordinated gene expression program establishing division competence. Cell cycle gene expression shows upregulation of D-type cyclins initiating G1 progression, cyclin-dependent kinase CDK4/6 activation, E2F transcription factor activation controlling S phase genes, and coordinated expression establishing cell cycle machinery. G0 to G1 transition shows exit from quiescent G0 state characteristic of post-commitment berry cells, entry into G1 growth phase with increased protein synthesis and cell growth, preparation for DNA replication initiation, and commitment point passage enabling S phase entry. Division activation timing shows auxin levels reaching threshold days 2-5 post-fertilization in cells near seeds, progressive activation wave spreading through mesocarp days 3-7, spatial and temporal heterogeneity creating extended activation period, and completion of activation wave by end of first week. Cell population activation dynamics show asynchronous activation across mesocarp cell populations, earlier activation near seeds with higher auxin exposure, later activation in peripheral regions, and progressive recruitment creating temporal spread in cell cycle entry. Activation checkpoint controls show growth factor signaling requirements for G1 progression, nutrient availability checks ensuring metabolic capacity, DNA damage surveillance preventing division of damaged cells, and checkpoint passage enabling irreversible commitment to division. Metabolic activation accompanying division shows increased respiratory rate providing ATP for biosynthesis, enhanced protein synthesis supporting cell cycle machinery production, nucleotide biosynthesis preparing for DNA replication, and overall metabolic upregulation supporting proliferative state. Chromatin remodeling during activation shows histone modifications marking active versus repressed chromatin regions, chromatin accessibility changes enabling transcription factor binding, epigenetic programming establishing division-competent state, and coordinated chromatin changes facilitating cell cycle gene expression. DNA replication licensing shows origin recognition complex binding replication origins during G1, MCM helicase loading creating licensed origins, replication licensing preventing re-replication, and licensed origins enabling subsequent S phase DNA synthesis. Spatial activation patterns show inner mesocarp cells activating earlier than outer regions in some studies, possible auxin gradient from seeds affecting activation timing, exocarp surface cells showing distinct activation possibly coordinated with surface expansion, and overall coordinated activation maintaining tissue organization. Environmental effects on activation include temperature affecting signaling kinetics and metabolic activation rate with optimal 24-28°C supporting rapid activation within 3-5 days, cool conditions slowing activation extending to 5-9 days, and temperature as factor affecting activation timing and efficiency. Seed number effects show multiple seeds producing greater cumulative auxin accelerating activation, higher hormone dose potentially activating more cells or accelerating timing, and seed number affecting division extent through activation effects. Failed or delayed activation causes include inadequate auxin production from poor seed development, impaired auxin signaling from genetic or metabolic dysfunction, severe environmental stress during critical days 2-7 disrupting signaling, or developmental abnormalities preventing normal activation. Activation assessment methods include cell cycle marker gene expression analysis showing upregulation, immunohistochemistry detecting cyclin proteins, BrdU or EdU incorporation marking cells entering S phase, and proliferation marker PCNA detection indicating division-competent cells. The process represents first aspect of active berry cell division establishing division competence, preceding mitotic activity onset and periclinal division. Duration typically 3-7 days representing auxin-triggered cell cycle entry. Regional timing shows initial division activation during late June-early July across climates within first week post-fertilization. Viticultural significance includes representing critical first step in cell proliferation determining subsequent division success, demonstrating auxin as primary proliferation signal, showing temperature sensitivity during activation affecting division timing, and establishing division competence as prerequisite for cell number accumulation. Research applications include analyzing auxin signaling mechanisms in berry cells, investigating cell cycle activation regulation, evaluating environmental effects on activation timing, quantifying spatial activation patterns in pericarp, and assessing activation as determinant of division extent and berry cellularity.
Initial_Division_Activation is first aspect of Active_Berry_Cell_Division, distinguished by auxin-triggered cell cycle entry establishing proliferative competence. Represents G0 to G1 transition enabling division. Timing days 2-7 post-fertilization. Auxin-TIR1/AFB receptor signaling activating ARF transcription factors and cell cycle genes. Progressive activation wave through mesocarp. Temperature affects activation kinetics. Essential prerequisite for mitotic activity. Followed by Mitotic_Activity_Onset as divisions commence. Critical first step in cell proliferation program.
Auxin-getriggerte Aktivierung
G0-G1 transition
G0-G1-Übergang
Zellzyklus-Eintritt
auxin-triggered activation
cell cycle entry
initiale Teilungs-Aktivierung
proliferative competence establishment
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 432-434 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Initial Division Activation
An initial growth aborted tendril at node 6 on a shoot of Vitis vinifera cv. Merlot in Pomerol, France, measuring 2.8 cm length and appearing as a thickened brown stub at the node position, showing arrested development with visible vascular traces and slight bifurcation indication at the tip suggesting initiated but incomplete branching, occurring during a severe heat wave (daily maximum temperatures exceeding 38°C for five consecutive days) in late May that coincided with the critical early elongation phase for tendrils at nodes 5-7, with 45% of tendrils in this node range showing similar initial growth abortion while tendrils at earlier nodes (1-4) that had completed elongation before the heat wave remained functional, and tendrils at later nodes (8-10) that initiated after stress amelioration developed normally, representing a characteristic temporal band of abortion corresponding to the acute stress timing.
An early aborted tendril that ceased development during the initial elongation phase after primordium formation but before achieving recognizable mature tendril morphology in Vitis vinifera, characterized by arrested structures of 1.5-4 cm length showing some cell expansion and early tissue differentiation including rudimentary vascular tissue, visible as stunted protrusions or short stubs at node positions, exhibiting partial development of tendril characteristics including slight elongation, possible initiation of branching points, and beginning of epidermis differentiation, but failing to complete the elongation growth phase that would produce functional tendril length and structure. Initial growth aborted tendrils result from severe stress conditions or resource limitations occurring during the critical 1-2 week period following primordium initiation when rapid cell division and expansion normally drive tendril elongation, including acute water stress, sudden temperature extremes, nitrogen or other critical nutrient deficiency, pest damage targeting young tender tissues, or disease infection during vulnerable early growth stages. These abortion events represent significant metabolic investment loss compared to primordium stage abortion, as cells have undergone expansion and early differentiation processes consuming carbohydrates, nitrogen, and minerals, but occur early enough that resource commitment remains moderate compared to near-mature abortion. The occurrence of initial growth stage abortion indicates transient severe stress or acute environmental events during early tendril development rather than fundamental developmental incompetence, with recovery potential if stress conditions are ameliorated before subsequent organ development.
Initial growth aborted tendrils are distinguished from primordium stage abortion by achievement of some cell expansion and tissue differentiation, and from later abortion stages by failure to complete basic elongation growth and achieve recognizable mature tendril morphology. This abortion class is diagnostically valuable because it indicates the timing of stress events relative to shoot development, with bands of initial growth abortion at specific node positions revealing acute stress periods affecting organs at vulnerable developmental stages. The level of development achieved before abortion (length, branching initiation, vascular differentiation) provides information about stress severity and timing, with greater development suggesting later stress onset or less severe conditions. Initial growth abortion represents an intermediate metabolic cost, with more resource loss than primordium abortion but less than near-mature abortion, affecting calculations of vine resource allocation efficiency under stress conditions.
Anfangswachstum-abortierte Ranke
Stummelstadium abgebrochene Kletterorgan
early elongation failed tendril
frühe Elongationsphase fehlgeschlagene Ranke
initial development arrested tendril
stub stage aborted tendril
Based on Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 60-85 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Petrie et al. (2000) "The effect of leaf removal and canopy height on whole-vine gas exchange and fruit development of Vitis vinifera L. Sauvignon blanc" Functional Plant Biology 27=173-184 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 377-400.
Initial Growth Aborted Tendril
Cabernet Sauvignon initial growth early berries progress from 3.2-4 mm diameter and 0.02-0.04 g at 2 weeks post-fruit set through peak growth achieving 5.5-7 mm and 0.2-0.5 g at 4 weeks, to 8.5-10.5 mm and 0.9-1.3 g at 6 weeks (BBCH 75, end of Phase I), with cell number increasing from 1.2-1.6 million cells at 2 weeks through 1.8-2.4 million at 4 weeks to final 2.2-2.9 million cells at 6 weeks, demonstrating active cell division throughout initial growth with division rate highest during weeks 2-4 then declining during weeks 5-6 as transition to lag phase approaches. Syrah initial growth early berries examined at weekly intervals show cell division dynamics= week 2 (1.3 million cells, 12-15% mitotic index), week 3 (1.7 million cells, 14-18% mitotic index at peak), week 4 (2.1 million cells, 10-14% mitotic index declining), week 5 (2.5 million cells, 5-8% mitotic index), week 6 (2.7 million cells, 2-4% mitotic index near cessation, BBCH 75), with cell diameter simultaneously increasing from 32 μm (week 2) through 42 μm (week 4) to 58 μm (week 6), demonstrating overlapping cell division and cell expansion processes during initial growth phase. Chardonnay initial growth early berries show peak photosynthetic contribution during weeks 3-5 with net photosynthesis of 2.5-4.5 μmol CO₂ m⁻² s⁻¹ in green berries contributing 10-15% of carbon requirements, declining during week 6 as chlorophyll content begins decreasing in preparation for lag phase and eventual veraison, with sugars maintained at 5-7% throughout initial growth primarily supporting respiration (25-35 nmol CO₂ g⁻¹ s⁻¹) and growth rather than accumulating for storage. Merlot under two irrigation regimes during initial growth shows water-sufficient vines producing berries with 2.6-3.1 million cells at BBCH 75 (6 weeks) while water-deficit vines show only 1.9-2.4 million cells, with the cell number difference established during initial growth persisting through maturity creating 15-25% final berry size difference (water-sufficient 15-17 mm, 2.4-3 g versus water-deficit 13-15 mm, 1.8-2.3 g), demonstrating critical importance of water availability during initial growth cell division phase for determining ultimate berry size.
An initial growth early berry is an early developing berry in the established growth phase (BBCH 71-75, approximately 2-6 weeks post-fruit set) following successful fruit set, undergoing rapid expansion through active cell division and initial cell expansion, representing berries committed to development and actively accumulating the cell number that will determine final size potential. Initial growth early berries are characterized by= (1) rapid size increase from approximately 3-4.5 mm diameter at 2 weeks post-fruit set to 6-11 mm diameter by 6 weeks (end of Phase I at BBCH 75), representing the main cell division and early expansion phase (2) fresh weight increase from approximately 0.015-0.05 g at 2 weeks to 0.6-1.5 g by 6 weeks, representing 10-20 fold mass increase during the 4-week initial growth period (3) active cell division throughout the period with cell number increasing from approximately 1-1.8 million cells at 2 weeks to final cell number of 1.5-3.5 million cells by 6 weeks, with cell division rate peaking during weeks 3-4 then declining during weeks 5-6 (4) cells undergoing initial expansion from small sizes (25-45 μm diameter at 2 weeks) to intermediate sizes (40-65 μm at 6 weeks) even while cell division continues, creating simultaneous division and expansion dynamics (5) green photosynthetically active tissues with chlorophyll content increasing during weeks 2-4 then stabilizing or beginning to decline during weeks 5-6 as transition to lag phase approaches (6) acidity increasing from 10-15 g/L at 2 weeks to maximum levels of 20-25 g/L by weeks 5-6, primarily through malic acid accumulation while tartaric acid increases more slowly (7) sugar content remaining relatively low at 4-8% throughout initial growth phase, with imported sugars primarily used for growth (cell wall synthesis, respiration) rather than storage. Initial growth early berries show peak growth rates= berry diameter increasing at 0.5-1.2 mm per week during weeks 2-5 fresh weight increasing at 0.12-0.3 g per week during peak growth at weeks 3-5 cell division rates of 8-15% of cells in mitosis during peak division at weeks 2-4. Initial growth early berries establish final berry size potential= cell number achieved by end of initial growth (BBCH 75, week 6) determines the maximum potential berry size, as subsequent growth occurs primarily through cell expansion of this fixed cell population without additional cell division environmental factors during initial growth affecting cell division (water availability, temperature, nutrition, crop load) have lasting effects on final berry size independent of later conditions. Initial growth early berries undergo critical developmental transitions= seeds developing from embryos at 2-8 cell stage (week 2) through globular embryo (weeks 3-4) to heart/torpedo stage embryos (weeks 5-6) in seeded types, or seeds initiating development then beginning abortion in stenospermocarpic seedless types vascular system completing differentiation with xylem and phloem becoming fully functional to support high transpiration and nutrient import rates during rapid growth skin tissues differentiating with cuticle elaboration, stomata formation, and wax deposition beginning.
Initial_Growth_Early_Berry represents the main Phase I growth period (2-6 weeks post-fruit set) and is distinguished from Post_Fertilization_Early_Berry (0-2 weeks, fruit set establishment) by established growth versus fruit set determination. Initial growth early berries undergo peak cell division establishing final cell number determining size potential, making this a critical period for berry size determination. Understanding initial growth berry biology is essential for managing berry size through irrigation, crop load adjustment, and nutrition during the period when cell division can be influenced, and for predicting final berry size based on early development trajectory.
Zellteilungs-Stadium
cell-division berry
initial wachsende frühe Beere
rapid-growth Phase I berry
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 182-192 Coombe, B.G., McCarthy, M.G. (2000). Dynamics of grape berry growth. Australian Journal of Grape and Wine Research 6=131-135 Ojeda, H. et al. (2002). Influence of water deficits on grape berry growth. Vitis 41=141-145.
Initial Growth Early Berry
Chardonnay in Champagne shows initial inflorescence primordium formation during early July Year N at BBCH 19-31 at shoot apex within developing latent buds, with undifferentiated meristematic protrusions of 40-60 micrometers emerging at 2-3 basal node positions over 10-14 days under conditions of 20-24°C temperature, adequate light exposure in managed canopy, balanced nitrogen fertility, and moderate water availability typical of this moderately-to-highly fruitful early-bursting cultivar, establishing reproductive commitment for Year N+1 productivity
Initial inflorescence primordium represents the earliest recognizable stage of reproductive organogenesis where undifferentiated lateral meristem at bud apex commits to inflorescence identity and emerges as distinct meristematic protrusion, marking transition from vegetative to reproductive development within developing latent bud. This critical developmental stage occurs at shoot apex during active growth Year N typically June-July in temperate regions, representing first 7-14 days of inflorescence development from identity determination through initial protrusion formation before architectural elaboration begins. Primordium initiation occurs at lateral position on primary bud axis opposite developing leaf primordium, following architectural pattern with inflorescence primordia forming at positions alternating with tendril primordia in regular sequence. Inflorescence identity determination involves molecular switches suppressing tendril developmental genes and activating reproductive meristem identity genes including homologs of LEAFY, APETALA1, and other floral identity genes, with identity commitment representing irreversible developmental decision under normal conditions. Initial primordium appearance shows small meristematic protrusion measuring 30-80 micrometers diameter emerging from bud axis, containing 50-200 actively dividing cells with organized meristem structure including peripheral zone producing organ primordia and central zone maintaining stem cells. Cell division during initial primordium stage shows high mitotic activity with cycles of 24-36 hours producing cells contributing to primordium growth and preparing for subsequent architectural elaboration, with division rates highest in peripheral zones where branch primordia will initiate. Morphological characteristics at this stage show undifferentiated dome or cone-shaped structure lacking recognizable architectural features that will develop during subsequent stages, appearing similar to early tendril primordia requiring molecular markers or developmental tracking for definitive identification. Gene expression patterns during initial primordium stage show activation of reproductive identity genes suppressing vegetative tendril program, expression of meristem maintenance genes sustaining stem cell populations, and beginning expression of architectural patterning genes that will guide branching development. Hormonal environment during initiation shows low gibberellin concentrations permitting or promoting reproductive identity with exogenous gibberellin application or high endogenous levels capable of converting inflorescence primordia to tendrils demonstrating hormonal control of organ identity, while cytokinin presence supports meristematic activity and initial growth. Environmental signals affecting initiation include light quality with high red: far-red ratio and adequate light intensity promoting reproductive development, photoperiod potentially affecting reproductive allocation in some genotypes, and temperature within optimal growth range supporting active meristem function. Carbon availability affects initiation with adequate photosynthetic production providing resources for primordium formation, while carbon limitation from stress or excessive sink competition reduces initiation frequency affecting fruitfulness. Nitrogen status influences initiation with moderate nitrogen supporting primordium formation while excessive nitrogen promotes vegetative tendril formation, requiring balanced nutrition for optimal reproductive allocation. Water status may influence initiation with moderate water stress during this period potentially promoting reproductive development as adaptive response, while severe stress inhibits meristem activity reducing formation. Vascular connection initiation begins during early primordium stage with procambial cells establishing strand connecting primordium base to bud vascular system, providing pathway for nutrient and hormone delivery though transport is minimal at this early stage. Developmental commitment during initial primordium stage establishes trajectory toward inflorescence development that normally proceeds through subsequent architectural elaboration stages, though severe stress or experimental manipulation can cause abortion or reversion affecting final primordium number. Timing within season shows initial primordium formation occurring relatively early during shoot growth typically when shoot has 10-15 nodes and continues active elongation, with primordium formation in basal bud positions preceding apical positions following basipetal bud maturation sequence. Developmental duration of initial primordium stage spans approximately 7-14 days from first recognizable emergence to transition toward architectural elaboration with branch primordium initiation, with duration varying based on temperature and growth rate. Position effects show basal node buds initiating inflorescence primordia earliest and most reliably showing highest fruitfulness, while middle node buds form primordia during mid-season with moderate fruitfulness, and apical node buds may fail to form or show delayed reduced primordia reflecting declining conditions. Metabolic activity during initial primordium stage shows elevated respiration supporting cell division and growth, high protein synthesis producing cell cycle proteins and regulatory factors, and active nucleic acid synthesis supporting rapid cell proliferation. Temperature during initiation shows optimal 24-28°C supporting maximum formation frequency and rapid development, with suboptimal temperatures reducing formation or slowing development affecting subsequent architectural elaboration. The process represents first distinguishable stage of inflorescence primordium formation marking commitment to reproductive development, transitioning to inflorescence meristem differentiation as architectural organization develops. Cultivar variation appears in initiation frequency determining fruitfulness with highly fruitful cultivars forming primordia at most bud positions while low fruitfulness types show sparse formation, timing of initiation affecting developmental window, and sensitivity to environmental conditions affecting formation reliability. Regional timing shows initial inflorescence primordium formation during late June-July in warm climates, July-early August in temperate regions, and July-August in cool areas corresponding to active shoot growth period when conditions support reproductive allocation. Viticultural significance includes representing first commitment to reproductive development with management during this period critically affecting formation frequency, understanding that conditions promoting balanced growth without excessive vigor favor initiation, and recognizing that stress or resource limitations during initiation can reduce fruitfulness affecting subsequent yield. Research applications include analyzing molecular mechanisms of reproductive identity determination and commitment, investigating hormonal and environmental regulation of inflorescence versus tendril fate decisions, documenting earliest stages of reproductive organogenesis through microscopic and molecular techniques, evaluating factors affecting initiation frequency and their implications for fruitfulness, and assessing genetic variation in reproductive commitment affecting productivity across cultivars.
Initial_Inflorescence_Primordium is first of four Inflorescence_Primordium_Formation subphases, distinguished by earliest recognizable reproductive primordium emergence before architectural elaboration. Represents commitment to reproductive rather than vegetative tendril development. Transitions to Inflorescence_Meristem_Differentiation as architectural organization develops. Critical stage determining formation frequency and fruitfulness. Highly sensitive to environmental conditions and hormonal signals.
Primordieninitiierung
early inflorescence formation
frühe Gescheinsanlage
initial inflorescence meristem
initiales Blütenstandsprimordium
primordium initiation
reproductive primordium emergence
reproduktives Primordium
Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337 Srinivasan C & Mullins MG (1981) Physiology of flowering in the grapevine, Am J Enol Vitic 32:47-63
Initial Inflorescence Primordium
In Riesling vineyards of the Rheingau, Germany, initial leaf detachment commences during the last week of October when the first autumn frosts with temperatures of -1 to -2°C trigger separation of basal leaves representing approximately 10-12% of canopy leaf area, marking the onset of defoliation at post-BBCH 93 to early BBCH 95 developmental stage.
Initial leaf detachment in Vitis vinifera represents the earliest phase of physical leaf separation, characterized by the first leaves falling from the vine canopy following completion of abscission zone cell separation processes. This phase typically begins in late October to early November in Northern Hemisphere viticulture, when the oldest basal leaves that initiated senescence earliest have completed abscission zone activation and experience sufficient mechanical stress to trigger separation. The first detaching leaves are predominantly those at basal node positions that received reduced light penetration during the growing season and consequently initiated senescence programs 2-3 weeks before more exposed canopy positions. Anatomically, initial detachment occurs when the separation layer cells within the abscission zone have completed cell wall dissolution through the coordinated action of cellulases, polygalacturonases, and other hydrolytic enzymes, reducing attachment strength below the threshold where environmental forces can effect separation. The triggering events for initial leaf fall typically include the first significant autumn wind events, light frost exposure causing tissue damage that increases mechanical stress, or rainfall adding weight to weakened attachment points. Quantitatively, initial leaf detachment encompasses approximately the first 10-15% of total canopy leaf area, representing leaves from approximately 2-4 basal nodes on each shoot that underwent earliest senescence initiation. The process serves as an important phenological marker for viticulturists, indicating that dormancy preparation has progressed sufficiently for abscission zone maturation to reach completion at the most advanced positions. Physiologically, leaves undergoing initial detachment have typically completed nutrient remobilization processes, with nitrogen content reduced to 0.8-1.2% dry weight compared to 2.5-3.5% in fully functional summer leaves, and chlorophyll content degraded to less than 10% of maximum summer values. Temperature conditions during initial leaf detachment typically feature daytime maxima of 10-15°C and nighttime minima approaching or slightly below 5°C, creating the thermal environment that completes abscission zone enzyme activity while providing mechanical stress through frost-induced tissue changes. The spatial pattern of initial detachment provides diagnostic information about canopy senescence uniformity, with highly synchronous initial leaf fall indicating consistent vine nutrition and water status, while scattered or delayed initial detachment may suggest heterogeneous canopy conditions or stress factors that disrupted normal senescence signaling.
Initial_Leaf_Detachment represents the first temporal phase of Leaf_Detachment, specifically encompassing the earliest 10-15% of leaf separation events. Distinguished from Progressive_Leaf_Fall which follows as detachment spreads through the canopy, and from Mass_Leaf_Fall which represents the period of most rapid defoliation. Occurs predominantly at basal canopy positions where senescence initiated earliest. Duration typically spans 5-10 days depending on weather conditions. Key indicators include first visible ground litter accumulation and initial exposure of basal shoot regions.
beginnender Blattfall
beginning leaf abscission
early leaf detachment
erster Blattabfall
first leaf fall
frühe Blattablösung
initial defoliation
initiale Entblätterung
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Zapata C et al (2004) Partitioning and mobilization of starch and N reserves in grapevine. Journal of Experimental Botany 55: 2165-2175 Wermelinger B, Koblet W (1990) Seasonal growth and nitrogen distribution in grapevine leaves. Vitis 29: 15-26
Initial Leaf Detachment
Chardonnay in Champagne shows initial pollen exposure during early June Year N+1 at BBCH 60-65, with yellow pollen masses becoming visible through 2.5-3.2 mm anther slits following morning dehiscence 9am-11am, explosive discharge releasing 15-25% pollen content immediately followed by gravitational and wind-assisted liberation, under cool maritime conditions of 19-24°C temperature, 58-70% relative humidity, typical of this early-bursting cultivar exposing abundant pollen supporting 64-76% fruit set
Initial pollen exposure represents the first revelation of mature pollen grains following anther dehiscence when longitudinal slits expose locule contents making pollen accessible for liberation and dispersal, establishing beginning of active pollen release phase. This critical exposure event occurs during anther dehiscence in early-mid June in temperate regions Year N+1, typically within minutes to hours following stomium opening as part of BBCH 60-65 flowering stages. Exposure mechanism involves anther wall rupture creating longitudinal slits exposing locule interior, separation of slit edges revealing yellow pollen mass within locules, and immediate beginning of pollen liberation through opened pathway enabling atmospheric access. Visual characteristics at exposure include sudden appearance of yellow-brown pollen mass visible through anther slits, contrast between exposed pollen and surrounding anther wall tissues, and characteristic color change in anthers from green or pale toward yellow-brown indicating pollen maturity and exposure. Pollen mass organization at exposure shows grains loosely aggregated within locules from degenerated tapetum remains, initial adhesion in small clumps before individual grain separation, and progressive loosening facilitating liberation and dispersal. Initial pollen liberation following exposure proceeds through multiple mechanisms including explosive discharge when rupture occurs releasing pollen bolus from locule pressure, gravitational fall as loosened pollen drops from opened anther under gravity, and beginning wind-induced release as air movement agitates anthers dislodging pollen grains. Explosive discharge characteristics show initial sudden pollen ejection when dehiscence occurs potentially dispersing pollen 5-15 mm from anther, brief explosive phase lasting seconds producing immediate pollen cloud, and representing 10-30% total pollen content with remainder requiring gravitational or wind-assisted liberation. Gravitational release following explosive phase shows continuous slow pollen fall from opened anthers over minutes to hours, positioning of anthers affecting fall pattern with lateral or downward orientation facilitating release, and accumulation of fallen pollen on flower structures or direct capture by nearby stigmas. Environmental effects on initial exposure include humidity at rupture affecting pollen hydration state with moderate 50-65% optimal for maintaining viability without promoting aggregation, temperature affecting pollen surface properties and liberation efficiency, and air movement affecting dispersal pattern from exposed mass. Pollen viability at initial exposure shows maximum capacity immediately upon exposure with fresh pollen exhibiting 75-90% germination under optimal conditions, brief window of peak viability lasting hours before progressive decline, and critical importance of rapid pollination during this optimal period. Pollen surface characteristics at exposure include intact exine wall with sculptured surface patterns, slight moisture content from locule environment maintaining viability without excessive hydration, and pollen coating substances from tapetum residues potentially affecting dispersal and stigma adhesion. Pollen grain separation at exposure shows progressive disaggregation from initial clumped state within locule, individual grain liberation enabling airborne dispersal, and mechanical agitation from wind or flower movement facilitating separation and release. Timing relationships show initial exposure occurring immediately following successful dehiscence typically within minutes, exposure of first anthers coinciding with early anthesis in apical flowers, and progressive exposure throughout cluster over 5-10 days as anthers sequentially dehisce. Atmospheric pollen introduction shows initial grains entering air surrounding flower creating beginning of pollen cloud, progressive accumulation as additional anthers expose pollen intensifying atmospheric concentration, and peak atmospheric levels coinciding with mass dehiscence during peak flowering. Capture opportunity following exposure includes pollen falling directly onto stigma in same flower facilitating self-pollination, pollen dispersing to neighboring flowers on same cluster, and airborne pollen traveling to more distant flowers or vines though most successful pollinations occur locally. Self-pollination efficiency at initial exposure shows high probability through close anther-stigma proximity typically 0.5-1.5 mm spatial separation, gravitational pollen fall naturally directing toward centrally positioned stigma, and immediate pollen availability coinciding with stigma receptivity development. Stigma contact timing shows pollen-stigma encounter potentially occurring within minutes to hours of exposure through direct fall or brief dispersal, rapid germination initiation within 1-4 hours if stigma receptive and conditions favorable, and beginning of successful pollination events contributing to fruit set. Environmental effects on exposure success include favorable conditions enabling normal dehiscence and exposure supporting pollen liberation, rain during exposure washing pollen from anthers or preventing exposure through maintained humidity, and optimal temperature-humidity combination maximizing pollen function. Failed exposure results from incomplete dehiscence providing inadequate slit opening limiting pollen access, environmental conditions preventing normal rupture maintaining enclosed pollen, or abnormal anther development affecting exposure capability, all contributing to reduced pollination potential. The process represents third and final aspect of anther dehiscence marking transition from enclosed to accessible pollen, following anther wall rupture and stomium opening. Duration represents brief period from exposure through initial liberation typically minutes to hours. Cultivar variation in exposure characteristics may include pollen mass appearance reflecting production quantity and maturity, liberation efficiency from exposed anthers affecting dispersal, and coordination with stigma receptivity affecting pollination opportunity. Regional timing shows initial pollen exposure during flowering period across all climates coinciding with favorable morning conditions enabling dehiscence and exposure. Viticultural significance includes marking beginning of effective male reproductive function with pollen becoming available for pollination, representing critical transition enabling fertilization and fruit set, and demonstrating importance of favorable conditions during exposure for reproductive success. Research applications include analyzing pollen liberation mechanisms from exposed anthers, quantifying pollen release rates and patterns, evaluating environmental effects on liberation efficiency, assessing pollen-stigma encounter probabilities, and documenting exposure-pollination timing relationships affecting reproductive outcomes.
Initial_Pollen_Exposure is third and final aspect of Anther_Dehiscence, distinguished by revealing pollen making it accessible for liberation. Follows Anther_Wall_Rupture and Stomium_Opening. Marks transition from enclosed to accessible pollen beginning active release phase. Represents critical moment enabling pollination. Brief minutes to hours as exposure occurs and initial liberation begins. Transitions to Active_Pollen_Shedding as sustained dispersal proceeds. Essential event establishing male reproductive function.
Pollenmassenfreilegung
Pollensichtbarkeit
erste Pollenzugänglichkeit
first pollen access
initial pollen revelation
initiale Pollenexposition
pollen mass exposure
pollen visibility
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 378-382 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 288-292
Initial Pollen Exposure
In a Napa Valley Chardonnay vineyard, initial sugar increase was documented over 8 days in early August Year N+1 (BBCH 83) when berry Brix rose from 8.4° to 13.6° (average 1.95% per week), VvHT1 expression was 58-fold elevated, hexose:sucrose ratio increased from 0.6 to 3.8, berry fresh weight rose 24% from 1.08 to 1.34 g, firmness declined to 2.1 N, titratable acidity 9.8 g/L beginning decline, under warm conditions (daily high 28-31°C, adequate irrigation supporting osmotic berry expansion).
Initial_Sugar_Increase represents the earliest phase of accelerated hexose accumulation during veraison onset when berry sugar concentration begins rising at enhanced rates transitioning from slow pre-veraison accumulation (32°C may impair to <1.0% through metabolic disruption, light affecting source leaf photosynthesis and thus sugar availability with adequate canopy light exposure essential for supporting accumulation rates, water status with moderate deficit potentially enhancing concentration through reduced berry expansion though excessive stress limits import through general metabolic impairment, and source-sink balance with adequate leaf area per fruit weight (12-16 cm² leaf area per g fruit) necessary for supporting initial increase rates. Cultivar variation in initial increase characteristics shows: rapid-accumulating varieties (Chardonnay, Gewürztraminer) achieve 2.0-2.5% Brix per week during initial increase reaching 12-15° Brix in 6-8 days, moderate cultivars (Merlot, Cabernet Sauvignon) show 1.5-2.0% per week reaching 11-14° in 8-10 days, while slower types (Nebbiolo, Carignan) display 1.2-1.6% per week requiring 10-14 days to reach 11-13° reflecting genetic differences in transporter expression and metabolic capacity. Coordination with other ripening processes during initial increase includes: berry softening progressing concurrently with 20-35% firmness reduction from green berry values as cell wall degradation proceeds, color development (in red cultivars) showing initial or progressive anthocyanin accumulation synchronized through common ABA signaling, acid levels beginning decline particularly malate respiratory consumption initiating though titratable acidity still elevated 8-12 g/L during initial increase, and berry expansion resuming after lag phase with 20-40% fresh weight increase through osmotic water import driven by sugar accumulation. Metabolic integration during initial increase shows: respiratory quotient (CO₂ produced / O₂ consumed) increasing from 0.85-1.00 to 1.05-1.25 indicating transition from oxidative metabolism to substrate-level reactions with some fermentative activity, carbon allocation shifting from structural synthesis to storage with 50-70% of imported carbon accumulating as hexoses, and nitrogen metabolism adjusting with amino acid levels declining 25-40% as nitrogen incorporation into proteins and secondary metabolites proceeds. Quality implications of initial increase inclu rate of initial increase affecting ultimate sugar levels with faster early rates often correlating with higher final Brix (r² = 0.65-0.82), timing of initial increase relative to flavor and phenolic development determining optimal harvest window with early increase enabling earlier harvest or late increase requiring extended hang time, and uniformity of initial increase across berries and clusters indicating ripening synchrony important for harvest timing and wine composition consistency. Viticultural management during initial increase involves: irrigation often reduced or terminated as berry growth slows and sugar accumulation accelerates recognizing changing water demands, canopy management ensuring adequate light to source leaves supporting continued photosynthesis and sugar export to berries, disease monitoring intensifying as increasing sugar attracts insects and provides substrate for fungal pathogens particularly Botrytis, and crop protection measures including bird netting installation as sweetening fruit becomes attractive to wildlife. Monitoring initial increase provides harvest forecasting: weekly Brix sampling tracking accumulation rates enabling projection of harvest timing (typically 35-50 days from initial increase to target Brix depending on cultivar and heat accumulation), correlation between initial increase rate and final sugar achieved informing yield and quality predictions, identification of spatial variation in increase rates revealing vineyard heterogeneity requiring selective harvest or management intervention, and integration with berry sampling for acids, pH, and phenolics providing comprehensive maturity assessment beyond sugar alone.
Initial_Sugar_Increase is distinguished from Sugar_Accumulation_Onset by representing active increase rather than triggering, from Rapid_Sugar_Accumulation by showing moderate (1.2-2.5% Brix/week) rather than peak (2.0-3.5%/week) rates, and from later accumulation by occurring early in veraison BBCH 83. Duration typically 5-10 days. Hexose transporters 40-70% of maximum expression. Glucose:fructose ratio ~1.0 initially. Temperature optimal 25-30°C. Initial_Sugar_Increase represents transition from slow to moderate accumulation establishing the foundation for peak sugar loading.
beginnende Süßung
beginnender Zucker-Konzentrations-Anstieg
beginning sugar concentration rise
early hexose accumulation
frühe Hexose-Akkumulation
initial Brix elevation
initiale Brix-Erhöhung
nascent sweetening
Keller M (2020) The Science of Grapevines 3rd Ed pages 218-222 early sugar accumulation Davies C, Wolf T, Robinson SP (1999) Three putative sucrose transporters are differentially expressed in grapevine tissues Plant Sci 147:93-100 transporter expression Coombe BG, McCarthy MG (2000) Dynamics and environmental control of grape berry growth Australian J Grape Wine Res 6:73-83 accumulation dynamics.
Initial Sugar Increase
Cabernet Sauvignon inner mesocarp at maturity comprises 5-7 cell layers (total 150-220 μm thickness) adjacent to locular cavities containing 2-3 seeds, with cells showing progressive size decrease from outermost (75-92 μm diameter) through middle (62-78 μm) to innermost (52-68 μm) approaching seed contact position, walls 0.9-1.3 μm (intermediate between middle mesocarp 0.5-0.7 μm and outer mesocarp 1-1.4 μm), containing tannins 4.5-6.5 mg/g (substantially elevated above middle mesocarp 0.4-0.8 mg/g), contributing approximately 18-24% of wine skin tannins during standard 14-18 day maceration. Syrah inner mesocarp tannin gradient showing systematic increase toward seeds= outermost inner mesocarp (200-280 μm from seed surface) with 3.2-4.5 mg/g tannins middle inner mesocarp (100-200 μm) with 4.8-6.2 mg/g innermost inner mesocarp (0-100 μm) with 5.8-8 mg/g, demonstrating progressive tannin increase approaching seed proximity potentially reflecting seed influence on pericarp tannin synthesis or accumulation, with tannin gradient extending 200-300 μm into mesocarp from seed-adjacent position. Merlot inner mesocarp seed contact developmental changes= early development (BBCH 73-75, weeks 3-6) showing developing seeds filling locular cavities with seed coats directly contacting inner mesocarp cells mid development (BBCH 77-81, weeks 6-10) showing seeds completing growth while pericarp continues expansion creating beginning separation maturity (BBCH 89, week 18) showing 20-45 μm air gaps between mature seeds and inner mesocarp cells as berry expanded beyond seed dimensions, demonstrating temporal progression from direct seed contact to seed proximity affecting potential for seed-pericarp biochemical interactions. Chardonnay white cultivar inner mesocarp showing presence in white grapes= 4-5 cell layers, 135-195 μm thickness, cells 54-88 μm diameter, tannins 2.8-4.5 mg/g (elevated above middle mesocarp 0.4-0.9 mg/g but lower than red cultivar inner mesocarp 4-7 mg/g), demonstrating inner mesocarp as tannin-enriched zone present in white cultivars though typically not extracted during white wine production (immediate pressing separating juice from solids avoiding inner mesocarp tannin extraction). Pinot Noir inner mesocarp contribution to wine tannins during maceration time-course= day 5 maceration extracting primarily outer pericarp tannins with minimal inner mesocarp extraction day 10 showing beginning inner mesocarp extraction contributing 8-12% of wine skin tannins day 15 showing substantial inner mesocarp extraction contributing 20-28% of wine skin tannins, demonstrating that inner mesocarp represents later-extracting tannin reservoir requiring extended maceration for full extraction, with extraction timing allowing winemakers to control wine tannin levels and structure through maceration duration management.
An inner mesocarp is the innermost zone of the mesocarp adjacent to locular cavities and seeds (or alternatively the entire inner pericarp/endocarp region in some classification schemes), typically 100-300 μm thickness comprising 3-8 cell layers, characterized by cells smaller than middle mesocarp (40-90 μm diameter versus 80-180 μm), moderately thick walls (0.8-1.5 μm versus 0.4-0.8 μm), and elevated tannin content (3-7 mg/g versus 0.3-1 mg/g in middle mesocarp), representing transition zone between storage flesh and seed-containing cavities or the innermost pericarp layer in direct or near-direct contact with seeds. Inner mesocarp anatomical organization comprises= (1) cells arranged in layers parallel to locular cavity and seed surfaces creating boundary between fleshy mesocarp and air-filled or seed-filled cavities (2) progressive cell size decrease from outermost inner mesocarp near middle mesocarp boundary (70-90 μm diameter) through middle inner mesocarp (55-75 μm) to innermost inner mesocarp adjacent to seeds/cavities (40-60 μm) (3) progressive wall thickness increase from 0.8-1 μm approaching middle mesocarp to 1.2-1.5 μm in innermost layers (4) total inner mesocarp representing 4-7% of pericarp volume (versus 75-85% for middle mesocarp, 8-12% for outer pericarp plus outer mesocarp) (5) variable layer number (3-8 layers) depending on berry size, cultivar, and developmental conditions. Inner mesocarp compositional characteristics= (1) tannin content 3-7 mg/g dry weight, substantially elevated above middle mesocarp (0.3-1 mg/g) and intermediate between middle mesocarp minimum and outer pericarp maximum (8-15 mg/g), with tannins predominantly procyanidins showing polymerization degree 10-16 subunits similar to skin tannins (2) sugar and acid content similar to middle mesocarp (18-26% sugars, 0.5-0.9% acids) showing minimal radial compositional gradients in primary metabolites (3) anthocyanins essentially absent (<0.05 mg/g) similar to middle mesocarp with anthocyanins concentrated in outer pericarp hypodermal region (4) potential seed-derived compounds in innermost layers if diffusion occurs from adjacent seeds though evidence for substantial seed-to-pericarp transfer limited. Inner mesocarp functional characteristics= (1) structural boundary defining locular cavities where seeds develop and mature (2) potential seed-pericarp interaction zone during development when seeds contact inner mesocarp cells before locular air spaces develop (3) tannin-enriched zone contributing to wine phenolic profile particularly during extended maceration when deeper tissue extraction occurs (4) transition zone creating gradient from soft large-celled middle mesocarp to smaller firmer inner mesocarp cells approaching seeds. Inner mesocarp development= cells undergoing modest expansion during ripening (80-140% diameter increase versus 180-250% in middle mesocarp) creating smaller final size despite similar initial size tannin synthesis and accumulation during ripening creating elevated tannin levels at maturity locular cavity development during late ripening creating air spaces separating inner mesocarp from seed surfaces in mature berries though seeds contact inner mesocarp during early-to-mid development. Inner mesocarp in winemaking= inner mesocarp tannins extracting during extended maceration (typically after 10-15 days) contributing 15-25% of wine skin tannins beyond outer pericarp contribution inner mesocarp requiring more vigorous extraction conditions than middle mesocarp due to smaller cells and thicker walls inner mesocarp tannins showing similar properties to outer pericarp skin tannins (higher polymerization, less astringent than seed tannins) contributing to wine structure and aging potential.
Inner_Mesocarp represents the seed-adjacent mesocarp zone and is essentially synonymous with Inner_Pericarp in many classification schemes, with distinction depending on whether endocarp is recognized as separate layer. Sub-classification into Soft and Juicy inner mesocarp reflects textural properties. Understanding inner mesocarp is relevant for detailed anatomical characterization, for comprehending tannin distribution gradients, for winemaking extraction strategies where inner mesocarp represents intermediate-to-difficult extraction requiring extended maceration, and for investigating potential seed-pericarp interactions during development.
Samennahes Fruchtfleisch
inner flesh
innere Fruchtfleisch-Schicht
inneres Mesokarp
seed-adjacent flesh
Cadot, Y. et al. (2006). Tannin composition in grape skins and seeds. Journal of Agricultural and Food Chemistry 54=2973-2987 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 180-185 Downey, M.O. et al. (2003). Relationship between skin tannin concentration and berry development. American Journal of Enology and Viticulture 54=114-119.
Inner Mesocarp
Cabernet Sauvignon inner pericarp at maturity comprises 4-6% of pericarp volume (150-220 μm thickness, 5-7 cell layers), with cells decreasing in size from outer inner pericarp (72-88 μm diameter) through middle inner pericarp (58-75 μm) to innermost inner pericarp (48-62 μm) adjacent to seed surfaces, containing tannins 4.5-6.5 mg/g dry weight (procyanidin degree of polymerization 11-15, galloylation 12-17%), sugars 23-25% and acids 0.6-0.8% similar to middle pericarp, contributing approximately 20% of total wine skin tannins (outer pericarp contributing 45%, middle pericarp 5%, seeds 30%) during standard red wine maceration. Syrah inner pericarp tannin characterization showing intermediate tannin properties= total tannins 5.2-7.8 mg/g dry weight, intermediate between outer pericarp (9.5-13 mg/g) and middle pericarp (0.6-1.2 mg/g) tannin composition predominantly procyanidins (82-88% of total) with moderate prodelphinidins (12-18%) polymerization degree 12-17 subunits similar to outer pericarp (11-16) and higher than middle pericarp (8-12) demonstrating inner pericarp as distinct tannin-rich zone contributing to total skin tannin pool. Merlot inner pericarp anatomical organization examined by serial sectioning= 6 distinct cell layers in inner pericarp region, layer 1 (outermost, adjacent to middle pericarp) with cells 78-92 μm diameter, layer 3 (mid-inner pericarp) with cells 62-78 μm, layer 6 (innermost, adjacent to locular cavity) with cells 45-58 μm, demonstrating progressive cell size decrease through inner pericarp thickness creating smooth transition from large middle pericarp to small inner pericarp boundary cells, with tannin content showing inverse relationship (layer 1= 3.2-4.8 mg/g, layer 3= 4.8-6.2 mg/g, layer 6= 5.5-7.5 mg/g) suggesting tannin concentration increases toward seed proximity. Chardonnay white cultivar inner pericarp showing presence in white grapes= similar anatomical organization to red cultivars (4-5 cell layers, 130-190 μm thickness, cells 52-85 μm diameter), moderate tannin content 2.8-4.5 mg/g (lower than red cultivars but substantially higher than middle pericarp 0.4-0.9 mg/g), demonstrating inner pericarp as tannin-enriched zone present in white cultivars though less emphasized than in red cultivars bred for tannin content. Pinot Noir inner pericarp seed contact effects studied by comparing berries with 2 seeds versus 4 seeds= 4-seed berries showing slightly higher inner pericarp tannin content (5.8-7.2 mg/g versus 4.8-6.2 mg/g in 2-seed berries), potentially reflecting greater seed surface area in contact with inner pericarp creating enhanced seed-pericarp interactions, though effect is modest suggesting inner pericarp tannins are primarily synthesized in situ rather than transferred from seeds.
An inner pericarp is the innermost region of the pericarp comprising the mesocarp layers immediately adjacent to locular cavities and seeds, typically 100-300 μm thickness comprising 3-8 cell layers, characterized by smaller cells than middle pericarp, elevated tannin content compared to middle pericarp, and direct contact with seeds creating potential for seed-pericarp biochemical interactions, representing a distinct anatomical and compositional zone contributing to wine tannin structure and potentially affecting seed-derived quality characteristics. Inner pericarp anatomical structure comprises= (1) cell layers organized parallel to locular cavity/seed surfaces creating boundary between fleshy middle pericarp and seed-containing cavities (2) cells smaller than middle pericarp typically 40-90 μm diameter (versus 80-180 μm in middle pericarp) with progressive size decrease from outer inner pericarp (70-90 μm) toward innermost inner pericarp (40-60 μm) adjacent to seeds (3) cell walls moderately thick typically 0.8-1.5 μm (intermediate between thin middle pericarp walls at 0.4-0.8 μm and thick outer pericarp walls at 1.5-3 μm) (4) cells with reduced vacuolar volume compared to middle pericarp (vacuoles occupying 80-88% of cell volume versus 90-95% in middle pericarp) and denser cytoplasm (5) intercellular spaces more prominent than in middle pericarp particularly in innermost layers creating somewhat loose cellular organization near seed surfaces. Inner pericarp compositional characteristics= (1) elevated tannin content typically 3-7 mg/g dry weight, intermediate between high outer pericarp (8-15 mg/g) and low middle pericarp (0.3-1 mg/g), with tannins predominantly procyanidins showing similar polymerization (8-16 subunits) and galloylation (10-18%) to outer pericarp tannins (2) sugar and acid content similar to middle pericarp (sugars 18-26%, organic acids 0.5-0.9%) showing minimal radial gradients in primary metabolites despite gradients in cell size and secondary metabolites (3) minimal anthocyanins in red cultivars (<0.1 mg/g versus 2-6 mg/g in outer pericarp hypodermal cells), with anthocyanins concentrated in outer pericarp and essentially absent from middle and inner pericarp (4) potential seed-derived compounds diffusing into inner pericarp from adjacent seeds particularly tannins and other phenolics, though magnitude of seed-to-pericarp transfer remains debated. Inner pericarp development and function= development during berry growth with inner pericarp cells undergoing less expansion than middle pericarp cells (100-150% diameter increase versus 180-250% in central middle pericarp) creating smaller mature cell size despite similar initial size locular cavity formation creating air-filled space between inner pericarp and seeds in mature berries, though during development inner pericarp cells directly contact developing seeds potentially allowing seed-pericarp signaling tannin synthesis and accumulation in inner pericarp occurring during ripening similar to outer pericarp though at intermediate levels potential protective function with tannins providing antimicrobial activity protecting seeds from pathogens. Inner pericarp contribution to wine composition= inner pericarp tannins extracted during red wine maceration contributing 15-30% of total wine tannins (outer pericarp contributing 40-55%, seeds 25-35%, percentages varying with maceration intensity and duration) inner pericarp tannins showing similar properties to outer pericarp skin tannins (higher polymerization and less astringent than seed tannins) contributing to wine structure and aging potential inner pericarp providing minimal wine color as anthocyanins absent.
Inner_Pericarp represents seed-adjacent tissue and is distinguished from Middle_Pericarp (flesh bulk, low tannins) and Outer_Pericarp (skin, highest tannins) by anatomical position, intermediate cell size, and intermediate-to-high tannin content. Inner pericarp contributes significant wine tannins particularly during extended maceration. Sub-classification into Seed_Adjacent and Locular inner pericarp may distinguish innermost layers versus slightly more external layers. Understanding inner pericarp structure and composition is relevant for wine tannin management, maceration protocol design, and for comprehending spatial tannin distribution in berry tissues.
inneres Perikarp
samennahes Fruchtfleisch
seed-adjacent tissue
Cadot, Y. et al. (2006). Tannin composition in grape skins and seeds. Journal of Agricultural and Food Chemistry 54=2973-2987 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 180-185 Downey, M.O. et al. (2003). The relationship between skin tannin concentration and berry size. American Journal of Enology and Viticulture 54=114-119.
Inner Pericarp
Merlot grafted on 3309C rootstock at 14 years showing inner rootstock bark 2.4 mm thickness between vascular cambium and active cork cambium 2.8 mm deep, inner bark comprising living secondary phloem with functional sieve elements conducting photosynthates from canopy to roots, companion cells supporting sieve element function, phloem parenchyma showing dense starch staining indicating active reserve storage (dormant winter sample), ray parenchyma extending radially through inner bark, cell viability testing confirming living status with active respiration and metabolic activity, inner bark essential for food transport and storage functions
Inner rootstock bark is the living functional tissue in rootstock trunk bark comprising secondary phloem between vascular cambium and innermost active cork cambium, typically 1-3 mm thickness in mature trunk, characterized by living cells including sieve elements conducting photosynthates, companion cells supporting sieve elements metabolically, phloem fibers providing mechanical strength, and phloem parenchyma storing carbohydrates and facilitating radial transport, showing active metabolism with respiration, protein synthesis, and physiological processes, conducting food from scion leaves to roots and enabling bidirectional transport of organic compounds, storing substantial carbohydrate reserves in ray and axial parenchyma particularly during dormancy, and undergoing continuous renewal as vascular cambium produces new phloem inward while older phloem isolated by cork cambium formation becomes part of outer bark eventually dying and sloughing. Inner rootstock bark cellular composition includes= containing sieve elements (elongated cells with perforated sieve plates conducting photosynthates through pressure flow mechanism), including companion cells (specialized parenchyma cells with dense cytoplasm metabolically supporting adjacent sieve elements), comprising phloem fibers (thick-walled cells providing mechanical strength and flexibility to bark), showing phloem parenchyma (living storage and secretory cells interspersed among conducting elements), and containing ray parenchyma (radially oriented living cells extending from cambium through phloem facilitating lateral transport and communication). Inner rootstock bark food transport function includes= conducting photosynthates from leaves to roots (sieve elements transporting sucrose, amino acids, and other organic compounds produced by photosynthesis in scion canopy downward), enabling bidirectional flow (phloem capable of transporting materials both upward and downward depending on source-sink relationships), showing seasonal variation (transport rates highest during active growth declining during dormancy), demonstrating critical function (girdling experiments removing phloem causing vine death confirming essential role), and supporting root growth (phloem transport providing carbohydrates and nutrients necessary for root system maintenance and expansion). Inner rootstock bark carbohydrate storage includes= accumulating starch reserves (phloem parenchyma and ray cells storing carbohydrates during late season and dormancy), showing seasonal cycles (reserves highest in late fall-winter declining during spring as mobilized for growth), providing stress reserves (stored carbohydrates supporting recovery from damage, disease, or environmental stress), enabling spring growth (phloem reserves mobilized before leaves fully functional supporting bud break and initial shoot growth), and contributing to trunk capacity (inner bark storage significant component of total trunk carbohydrate reserve pool). Inner rootstock bark mechanical properties includes= providing flexibility (phloem fibers contributing to bark's ability to bend without breaking), showing resilience (inner bark recovering shape after compression or deformation), enabling expansion accommodation (living bark capable of limited stretching accommodating trunk diameter increase), demonstrating tear resistance (phloem fibers preventing bark splitting under stress), and contributing to trunk integrity (inner bark mechanical properties complementing outer bark protection maintaining bark cohesion). Inner rootstock bark metabolic activity includes= showing respiration (living cells consuming oxygen and producing CO2 supporting metabolism), exhibiting protein synthesis (producing enzymes, structural proteins, and other compounds necessary for function), demonstrating active transport (moving substances across membranes requiring energy), synthesizing compounds (producing callose, phloem proteins, and secondary metabolites), and responding to signals (living cells detecting and responding to hormonal and environmental cues).
Inner_Rootstock_Bark emphasizes living phloem tissues and is distinguished from Outer_Rootstock_Bark (dead protective tissues) by focus on living cells and physiological function. Critical for understanding food transport, carbohydrate storage, metabolic activity, phloem anatomy, and the essential living bark tissues conducting photosynthates and storing reserves while providing mechanical support.
Functional bark
Innere Unterlagenrinde
Lebende Rinde
Living bark
Secondary phloem bark
Sekundäres Phloem
Inner Rootstock Bark
Merlot inner trunk bark at 14 years post-planting shows 2.6-3.8 mm depth of living secondary phloem comprising functional transport zone (outer 1.8-2.6 mm with active sieve tubes 18-32 μm diameter transporting photosynthate at 0.3-0.8 m/hour, companion cells maintaining sieve tube function), phloem parenchyma storing 22-32 g starch (8-11% dry weight contributing 18-24% of trunk total reserves), phloem fibers providing mechanical support, transitional crushed phloem zone (middle 0.6-1.0 mm with declining function), and phloem rays 3-5 cells wide connecting radially to xylem rays, inner bark bounded internally by vascular cambium (active 4-5 cell layer zone producing both xylem and phloem) and externally by innermost cork cambium at 2.8-4.0 mm depth
Inner trunk bark is the living tissue zone of trunk bark located between vascular cambium and innermost functional cork cambium, comprising secondary phloem produced by vascular cambium including functional sieve tubes, companion cells, phloem parenchyma, phloem fibers, and phloem rays, representing the metabolically active bark zone responsible for photosynthate transport, nutrient storage, and lateral transport. Inner trunk bark anatomical boundaries include= inner boundary at vascular cambium (interface with secondary xylem/wood), outer boundary at innermost functional cork cambium (interface with outer dead bark), radial depth typically 2-4 mm in mature vines (varying by vine age, vigor, and cultivar), complete circumferential coverage (continuous living phloem cylinder surrounding wood), and dynamic boundaries (inner boundary moving outward as new phloem produced, outer boundary moving outward as successive cork cambia arise). Inner trunk bark tissue composition includes= functional secondary phloem (most recent 2-4 years production maintaining active transport, typically outer 1.5-3 mm depth), transitional phloem (phloem 3-5 years old with declining function beginning to show crushing, middle 0.5-1.5 mm), non-functional crushed phloem approaching outer bark (phloem >5 years old compressed and losing viability, inner 0.2-0.8 mm adjacent to innermost cork cambium), and phloem rays (radial parenchyma bands connecting through phloem continuous with xylem rays providing lateral transport). Inner trunk bark cellular elements include= sieve tube elements (enucleate cells with specialized sieve plates conducting photosynthate, typical dimensions 150-400 μm length, 15-35 μm diameter), companion cells (small densely cytoplasmic cells 50-120 μm length maintaining sieve tube function and loading/unloading sugars), phloem parenchyma (living storage and transport cells containing starch reserves 5-12% dry weight contributing 15-25% of trunk total carbohydrate storage), phloem fibers (thick-walled supporting cells 400-800 μm length providing mechanical strength), and ray parenchyma (radially oriented cells 40-120 μm radial length connecting phloem to cambium to xylem). Inner trunk bark transport functions include= basipetal photosynthate transport (sugars from leaves moving downward through sieve tubes to roots, trunk storage, and developing fruit), bidirectional nutrient distribution (minerals from xylem redistributing through phloem to various sink tissues, recycled nutrients from senescing leaves translocating to storage), hormone transport (auxin, cytokinin, abscisic acid, gibberellin moving through phloem coordinating growth and development), lateral loading and unloading (companion cells and parenchyma loading sugars into sieve tubes from source tissues, unloading to sink tissues), and signaling molecule transport (proteins, RNA, and other compounds potentially moving through phloem providing systemic communication). Inner trunk bark storage functions include= starch accumulation in phloem parenchyma (storing 18-35 g carbohydrates per vine representing 15-25% of trunk total reserves), seasonal storage cycling (accumulation post-harvest October-November, maintenance through dormancy, mobilization during budbreak March-April supporting early growth), protein storage (phloem parenchyma accumulating storage proteins during senescence and dormancy), mineral storage and remobilization (nitrogen, phosphorus, potassium stored in phloem parenchyma recycled during spring growth), and radial storage coordination (phloem storage integrated with xylem storage through ray parenchyma connections). Inner trunk bark seasonal dynamics includes= spring high activity (March-May, intense phloem transport supporting budbreak, shoot growth, and flowering from mobilized reserves and early photosynthesis), summer peak transport (June-August, maximum sap flow rates transporting photosynthate from mature canopy to growing fruit, roots, and storage tissues), late summer fruit support (August-September, phloem prioritizing sugar delivery to ripening fruit), fall reserve accumulation (September-November, post-harvest photosynthate loading into phloem parenchyma building winter reserves), and winter quiescence (December-February, minimal transport activity, maintained cellular viability in dormant state). Inner trunk bark annual production and turnover includes= vascular cambium producing 0.2-0.8 mm radial phloem increment annually (much less than 1.5-5 mm annual xylem production reflecting asymmetric cambial division), functional phloem lifespan 2-4 years (sieve tubes maintaining conductivity for 2-4 growing seasons before losing function), progressive crushing (older phloem compressed by xylem expansion as trunk diameter increases), transition to outer bark (crushed non-functional phloem becoming incorporated into outer bark when cork cambium forms deeper than crushed tissue), and continuous renewal (ongoing phloem production maintaining 2-4 mm living inner bark depth despite outer layer loss). Inner trunk bark depth variation includes= young trunks (years 3-8) showing 1.8-3.2 mm inner bark (limited phloem accumulation in thin trunks), mature trunks (years 10-25) showing 2.2-4.2 mm inner bark (typical functional phloem depth in productive vines), old trunks (years 30-50+) showing 1.8-3.8 mm inner bark (similar to mature despite age as functional zone maintained at stable depth), and consistency across ages (inner bark depth relatively constant 2-4 mm contrasting with highly variable outer bark and wood depth reflecting continuous renewal maintaining functional zone). Inner trunk bark vulnerability and protection includes= susceptibility to freeze damage (living cells vulnerable to ice crystal formation, severe cold potentially killing phloem requiring bark damage assessment), vulnerability to heat injury (excessive temperatures >45°C potentially damaging phloem, sunscald risk on exposed trunks), pathogen susceptibility (living tissue vulnerable to canker diseases, vascular wilts, and other pathogens), mechanical injury effects (wounds exposing inner bark causing immediate phloem dysfunction requiring wound healing response), and dependence on outer bark protection (dead outer bark essential for maintaining inner bark viability shielding from environmental stress). Inner trunk bark wound response includes= localized phloem death around injuries (damaged cells dying creating non-functional zone adjacent to wounds), callus tissue formation (undifferentiated parenchyma proliferating from wound margins), wound phloem differentiation (callus tissue differentiating into functional phloem gradually restoring transport capacity), traumatic periderm formation (wound-induced cork cambium arising in callus tissue or surviving inner bark producing protective cork sealing wounds), and vascular reconnection (new phloem elements establishing connections bridging wound gaps restoring basipetal transport continuity). Inner trunk bark relationship to outer bark includes= inner living bark contrasting with outer dead bark (sharp transition at cork cambium boundary separating metabolically active from non-living tissues), complementary functions (inner bark transport and storage, outer bark protection and insulation), approximately equal depths (2-4 mm inner living bark, 2-6 mm outer dead bark in typical mature trunk), inner bark dependence on outer protection (living phloem requiring waterproof cork covering for survival), and coordinated development (phloem production by vascular cambium, cork production by cork cambium, both maintaining trunk bark integrity). Inner trunk bark significance in vine function includes= essential for long-distance transport (phloem sieve tubes providing only pathway for photosynthate distribution from canopy to roots and storage), critical for carbohydrate storage (phloem parenchyma contributing substantially to trunk reserves supporting spring growth), enabling nutrient cycling (phloem redistributing minerals and recycling senescence-mobilized nutrients), coordinating vine growth (phloem transporting hormones integrating root and shoot development), and determining trunk girdling sensitivity (complete inner bark disruption around circumference (girdling) causing vine death from blocked basipetal transport emphasizing inner bark critical function). Inner trunk bark disease impacts include= trunk canker diseases killing inner bark (Eutypa, Esca, Botryosphaeria causing localized phloem death), vascular wilt pathogens potentially affecting phloem (some pathogens colonizing phloem disrupting transport), phloem necrosis diseases (viruses or phytoplasmas causing sieve tube dysfunction), girdling cankers (circumferential inner bark death blocking basipetal transport causing distal tissue starvation), and importance of preventing inner bark damage (maintaining inner bark integrity essential for vine survival and productivity). Inner trunk bark assessment and observation includes= bark peeling revealing inner bark (removing outer dead bark exposing living phloem surface appearing cream to tan colored, moist texture), phloem necrosis appearing as brown discolored tissue (diseased dead inner bark contrasting with healthy cream-colored living phloem), sap flow evidence (phloem exuding clear to slightly cloudy sap when cut indicating active transport), starch staining (iodine revealing dark purple-black staining in parenchyma indicating carbohydrate storage), and cambial activity observation (active vascular cambium at inner bark-wood boundary visible as thin green-white layer during growing season).
Inner_Trunk_Bark emphasizes living phloem zone and is distinguished from Outer_Trunk_Bark (dead protective layers) and specific inner bark types (Living_Inner_Trunk_Bark, Phloem_Inner_Trunk_Bark emphasizing particular aspects) by encompassing complete functional bark between cambium and cork. Critical for understanding photosynthate transport, trunk carbohydrate storage, nutrient cycling, phloem lifespan and turnover, girdling sensitivity, and essential role in maintaining vine function requiring protection from environmental stress and mechanical damage.
Funktionales Phloem
Innere Stammrinde
Lebende Bastzone
Living phloem bark
Secondary phloem zone
Inner Trunk Bark
Integrated evaluation of Insect_Resistance_Disposition at the USDA-ARS San Joaquin Valley Agricultural Sciences Center assessed multiple insect pest responses across 30 cultivars and rootstocks using standardized bioassays and field monitoring. Choice and no-choice feeding tests quantified responses to glassy-winged sharpshooter (Homalodisca vitripennis), grape mealybug (Pseudococcus maritimus), variegated leafhopper (Erythroneura variabilis), and western grapeleaf skeletonizer (Harrisina brillians). V. vinifera 'Thompson Seedless' showed broad insect susceptibility with high sharpshooter feeding rates (4.2 probes/hour), mealybug population buildup (285 crawlers/vine at 60 days), leafhopper stippling covering 45% of leaf area, and complete defoliation by skeletonizer larvae. In contrast, V. arizonica accessions demonstrated multiple Insect_Resistance_Dispositions with 65% reduction in sharpshooter probing, 80% lower mealybug populations, leafhopper damage limited to 12% leaf area, and skeletonizer larvae showing 40% mortality on resistant foliage. Notably, insect resistance profiles were not uniformly correlated, with some genotypes showing strong leafhopper resistance but mealybug susceptibility, indicating independent genetic control of different Insect_Resistance_Dispositions. These findings inform both rootstock selection (for root-feeding insects) and scion breeding (for foliar and fruit insects), with particular emphasis on vector insects affecting disease transmission.
An Insect_Resistance_Disposition is a Pest_Resistance_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through defense processes that limit feeding, oviposition, development, or damage by phytophagous insects attacking grapevine foliage, shoots, flowers, or fruit. This disposition represents the capacity to resist the diverse insect herbivores that exploit grapevine as a food source or reproductive substrate, encompassing both direct feeders causing tissue damage and insect vectors transmitting plant pathogens. Major insect pests of grapevine include xylem-feeding sharpshooter leafhoppers (Cicadellidae: Cicadellini) that vector Xylella fastidiosa causing Pierce's disease, phloem-feeding leafhoppers (Cicadellidae: Typhlocybinae) causing stippling damage and reducing photosynthetic capacity, phloem-feeding mealybugs (Pseudococcidae) and soft scales (Coccidae) that vector Grapevine leafroll-associated viruses, lepidopteran berry and cluster feeders including grape berry moths (Tortricidae) that damage fruit and facilitate secondary infections, and foliage-feeding beetles (Scarabaeidae, Chrysomelidae) causing defoliation. Insect resistance mechanisms in plants include constitutive physical defenses (trichomes, tough tissues, surface waxes), preformed chemical defenses (phenolics, alkaloids, terpenoids), induced defenses triggered by herbivore feeding (proteinase inhibitors, volatile emissions attracting natural enemies), and tolerance mechanisms maintaining productivity despite insect damage. For grapevine specifically, the vector relationships of certain insects with devastating viral and bacterial pathogens elevate the importance of insect resistance beyond direct damage prevention, making leafhopper and mealybug resistance critical components of integrated disease management strategies.
Insect_Resistance_Disposition is_a Pest_Resistance_Disposition. This class encompasses resistance to all phytophagous insects affecting grapevine, parallel to Nematode_Resistance_Disposition and Phylloxera_Resistance_Disposition under Pest_Resistance_Disposition. Child classes include Leafhopper_Resistance_Disposition (including Pierce's disease vectors), Mealybug_Resistance_Disposition (leafroll virus vectors), Grape_Berry_Moth_Resistance_Disposition (fruit feeders), and Japanese_Beetle_Resistance_Disposition (foliage feeders). The vector relationships of leafhoppers and mealybugs with major viral and bacterial pathogens create important linkages between Insect_Resistance_Disposition and Pathogen_Resistance_Disposition hierarchies.
Herbivoren-Insekten-Resistenz
Insektenresistenz-Disposition
Schadinsekten-Abwehr
arthropod feeding resistance
entomological defense
insect herbivore resistance
Howe GA, Jander G (2008) Plant immunity to insect herbivores, Annual Review of Plant Biology 59:41-66 Walling LL (2000) The myriad plant responses to herbivores, Journal of Plant Growth Regulation 19:195-216 Daane KM et al. (2018) Ecology and management of grapevine insects, Integrated Pest Management in Viticulture, Burleigh Dodds Science Publishing
Insect Resistance Disposition
Instantaneous water use efficiency monitoring through the day in Chardonnay reveals the VPD-driven efficiency pattern, with early morning A/E of 5.8 mmol CO₂/mol H₂O (VPD 0.8 kPa, favorable efficiency), midday decline to 2.4 mmol/mol (VPD 2.8 kPa, reduced efficiency despite similar intrinsic WUE), and late afternoon recovery to 4.2 mmol/mol (VPD declining to 1.5 kPa), demonstrating how atmospheric demand variation creates substantial diurnal efficiency fluctuation that affects the water cost of carbon gain at different times and informs irrigation timing strategies that exploit high-efficiency periods.
Instantaneous water use efficiency quality is a transpiration efficiency quality that represents the ratio of net CO₂ assimilation rate to transpiration rate (A/E), characterizing the real-time efficiency of carbon gain relative to actual water loss that reflects both leaf physiological properties and the atmospheric demand conditions driving transpiration at the moment of measurement. Within the BFO framework, this quality specifically addresses the realized dimension of water use efficiency, distinguishing A/E from intrinsic WUE (A/gs) while recognizing that instantaneous WUE captures the actual carbon-water exchange occurring under current environmental conditions including the vapor pressure deficit that determines transpiration rate at any given stomatal conductance. The ontological significance lies in instantaneous WUE providing the measure of actual water cost of carbon gain under specific conditions, where A/E indicates how much carbon the vine is fixing per unit water actually lost, with this ratio varying substantially through the day and season as vapor pressure deficit fluctuates, making instantaneous WUE both more variable and more directly relevant to actual water consumption than intrinsic WUE. The physiological and environmental basis for instantaneous WUE involves the ratio A/E = A/(gs × VPD × constant), where the inclusion of VPD means that instantaneous WUE declines under high evaporative demand even when intrinsic WUE (A/gs) remains constant, creating the characteristic pattern of declining water use efficiency through the day as VPD increases. The typical diurnal pattern of instantaneous WUE shows highest values in early morning (low VPD, favorable efficiency) declining through midday (high VPD reducing efficiency) and recovering in late afternoon (declining VPD), with this pattern affecting the optimal timing for gas exchange and the efficiency of water use at different times of day. The environmental factors affecting instantaneous WUE include vapor pressure deficit (primary driver of efficiency variation at constant intrinsic WUE), temperature (affecting both photosynthesis and vapor pressure), and CO₂ concentration (elevated CO₂ improving efficiency by increasing A at equivalent E), with management practices like morning irrigation potentially exploiting the higher efficiency of early-day gas exchange. Understanding instantaneous water use efficiency quality enables assessment of real-time water cost of carbon gain, timing of management interventions for maximum efficiency, and interpretation of how atmospheric conditions affect the water productivity of gas exchange.
Instantaneous_Water_Use_Efficiency_Quality addresses the realized A/E ratio incorporating actual transpiration under current VPD conditions. This quality captures the real water cost of carbon gain varying with atmospheric demand. The class supports timing optimization and actual water consumption assessment through instantaneous WUE monitoring.
A/E
Echtzeit-Effizienz
WUEinst
instantane Wassernutzungseffizienz
instantaneous WUE
real-time efficiency
Escalona JM et al. (2003) Distribution of leaf photosynthesis and transpiration within grapevine canopies under different drought conditions. Vitis 42(2): 57-64 Medrano H et al. (2015) From leaf to whole-plant water use efficiency (WUE) in complex canopies: limitations of leaf WUE as a selection target. The Crop Journal 3(3): 220-228
Instantaneous Water Use Efficiency Quality
Diurnal investigation of Instantaneous_WUE_Disposition at UC Davis compared A/E dynamics across 12 cultivars under contrasting atmospheric conditions (cool marine day with peak VPD 2.0 kPa versus hot interior day with VPD 4.5 kPa). Gas exchange measurements were conducted at 2-hour intervals from 06:00 to 18:00, enabling assessment of diurnal A/E patterns and cultivar × environment interactions. Instantaneous_WUE_Disposition showed strong diurnal and environmental variation. On the cool day, 'Chardonnay' achieved peak A/E of 4.8 μmol mmol⁻¹ at 08:00 (low VPD), declining to 2.2 μmol mmol⁻¹ by 14:00 (moderate VPD), with daily mean of 3.1 μmol mmol⁻¹. On the hot day, 'Chardonnay' peak A/E was only 3.5 μmol mmol⁻¹ at 08:00, declining to 1.2 μmol mmol⁻¹ by 14:00 (high VPD), with daily mean of 2.0 μmol mmol⁻¹. Cultivar differences in Instantaneous_WUE_Disposition were environment-dependent. 'Grenache' showed smaller diurnal and environmental variation (cool day mean 3.4, hot day mean 2.8 μmol mmol⁻¹) due to stomatal closure maintaining relatively constant Ci/Ca ratio across conditions. 'Syrah' showed larger variation (cool 2.9, hot 1.6 μmol mmol⁻¹) as sustained stomatal opening exposed A/E to full VPD effects. However, daily integrated carbon gain was higher in 'Syrah' (22 versus 18 mol CO₂ m⁻² d⁻¹ on the hot day) despite lower instantaneous efficiency, illustrating the trade-off between efficiency and productivity in Instantaneous_WUE_Disposition.
An Instantaneous_WUE_Disposition is a Water_Use_Efficiency_Disposition that inheres in a Vitis vinifera plant and is realized through the ratio of net photosynthetic rate (A) to transpiration rate (E), representing the actual carbon gain achieved per unit water lost under prevailing environmental conditions at the moment of measurement. This disposition represents the operationally relevant water use efficiency experienced by the plant, incorporating both physiological factors (stomatal conductance, photosynthetic capacity) and environmental factors (vapor pressure deficit, temperature, light intensity) that determine the actual rates of CO₂ uptake and water loss. Instantaneous WUE (A/E, expressed in μmol CO₂ mmol⁻¹ H₂O) varies throughout the day and across environments as VPD fluctuates, typically declining from morning (low VPD, high A/E) to midday (high VPD, low A/E) even when stomatal aperture remains constant. Genotypic variation in Instantaneous_WUE_Disposition reflects both intrinsic physiological differences (as captured by A/gs) and differences in the response to varying VPD, with some cultivars maintaining higher A/E under high evaporative demand than others at equivalent intrinsic efficiency. The relationship between instantaneous WUE and productivity is complex, as very high A/E may result from severely restricted gas exchange limiting both water loss and carbon gain, while moderate A/E with sustained gas exchange may accumulate more total carbon despite lower efficiency per unit water.
Instantaneous_WUE_Disposition is_a Water_Use_Efficiency_Disposition. Instantaneous WUE (A/E) represents actual efficiency under prevailing conditions, incorporating VPD effects that influence transpiration rate. A/E varies diurnally and seasonally with environmental conditions. High A/E does not guarantee high productivity, as severe stomatal closure reduces both water loss and carbon gain. The relationship between instantaneous efficiency and cumulative productivity depends on integration over time. A/E is relevant for irrigation scheduling decisions where water application is timed to periods of high efficiency.
A/E ratio
A/E-Verhältnis
Aktuelle-Wassereffizienz
Momentane-WUE-Disposition
actual water use efficiency
transpiration-based carbon efficiency
Medrano H et al. (2015) From leaf to whole-plant water use efficiency (WUE) in complex canopies: limitations of leaf WUE as a selection target, Crop Journal 3:220-228 Schultz HR, Matthews MA (1993) Growth, osmotic adjustment, and cell-wall mechanics of expanding grape leaves during water deficits, Crop Science 33:287-294 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data, Annals of Botany 105:661-676
Instantaneous WUE Disposition
Cabernet Sauvignon inflorescences at BBCH 55-57 (flower buds separating, individual flowers visible) exhibit intact calyptras 2.5-3.2 mm length appearing medium to light green, firmly attached at base but showing progressive swelling as enclosed stamens and pistil approach mature dimensions, with calyptras remaining intact for 4-8 days (depending on temperature progression) before transitioning to separating calyptra stage and subsequent shedding at BBCH 60-61. Syrah at BBCH 59 shows late-stage intact calyptras with distinct yellowish-green coloration, pronounced swelling creating bulged appearance, and visible loosening at base indicating flowers within 12-24 hours of calyptra dehiscence when temperatures reach 18-22°C threshold. Riesling experiencing late spring frost (-2 to 0°C) at BBCH 57-58 shows frost-damaged intact calyptras with brown discoloration, tissue collapse, and subsequent failure of normal calyptra shedding, resulting in retained dead calyptras preventing proper anthesis and causing 60-75% fruit set failure in affected flowers.
An intact calyptra is a calyptra that remains completely attached to the receptacle at its basal abscission zone with petal fusion at the apex fully maintained, forming a continuous sealed enclosure over the developing or mature stamens and pistil, representing the pre-anthesis state that provides complete protection and containment of reproductive organs prior to flower opening. Intact calyptras are characterized by= (1) complete attachment around the entire circumference of the basal insertion zone on the receptacle rim, with no separation or gap formation at the abscission layer (2) petal apices remaining fused in a continuous cap with no splitting or separation of individual petal components (3) calyptra envelope completely enclosing internal organs (stamens and pistil) with no exposure of reproductive structures to external environment (4) calyptra surface showing developmental progression from early green tightly appressed structure through later stages showing swelling, color changes (green to yellowish or brownish-green), and progressive loosening in preparation for shedding. The intact calyptra state encompasses the full pre-anthesis developmental period from initial petal fusion during early flower bud development through the final stages immediately preceding calyptra dehiscence, typically representing BBCH stages 51-59 (inflorescence emergence through flower buds swollen immediately pre-anthesis). Intact calyptras exhibit progressive developmental changes reflecting maturation of enclosed organs= early intact calyptras (BBCH 51-55) appear small (1.5-2.5 mm), tightly sealed, dark green, and firmly attached with minimal internal space mid-stage intact calyptras (BBCH 55-57) show enlargement (2-3.5 mm) as internal organs grow, slight color lightening to medium green, and increased internal volume creating slight loosening of calyptra fit over enclosed organs late-stage intact calyptras (BBCH 57-59) display maximum size (2.5-4 mm), distinct swelling or bulging as stamens and pistil approach maximum pre-anthesis dimensions, color change to yellowish-green or light green, visible loosening at base suggesting incipient abscission layer formation, and in some cases visible upward displacement as elongating stamens begin exerting pressure from within. The physiological state of intact calyptras reflects readiness for anthesis= tightly sealed intact calyptras indicate flowers days away from opening with immature internal organs swollen intact calyptras with color changes indicate flowers approaching anthesis within 1-3 days with nearly mature stamens and pistil intact calyptras showing basal loosening or slight lifting indicate flowers within 12-24 hours of calyptra shedding with active abscission layer formation. The transition from intact to separating calyptra status represents a critical developmental threshold, with environmental conditions (particularly temperature) during the late intact calyptra stage determining the timing of transition to anthesis. Intact calyptras provide essential protection during the vulnerable pre-anthesis period when developing pollen and pistil tissues are susceptible to desiccation, cold damage, pathogen infection, or physical injury, with the sealed calyptra maintaining humidity levels 15-25% higher than ambient air and protecting against temperature fluctuations, creating a buffered microenvironment optimal for final organ maturation. Damage or premature removal of intact calyptras through frost, hail, disease, or mechanical injury can severely impair subsequent flower function, reducing pollen viability, stigma receptivity, and fruit set potential by 40-80% depending on timing and severity of damage.
Intact_Calyptra represents the complete pre-anthesis state and must be distinguished from Separating_Calyptra (where basal separation has initiated) and Detached_Calyptra (where shedding is complete). The intact state encompasses a developmental continuum from early tight sealing through late-stage swelling and loosening, with sub-classification into Tightly_Sealed and Pre_Dehiscence types reflecting this progression. Intact calyptras serve as critical indicators of bloom timing, with the transition from intact to separating status marking the onset of anthesis (BBCH 60). Viticultural practices affecting intact calyptra health (frost protection, disease management, canopy microclimate optimization) directly impact subsequent bloom success and fruit set. The intact calyptra stage is the target period for certain fungicide applications (pre-bloom sprays at BBCH 57-59) to prevent Botrytis infection of flowers before calyptra shedding exposes vulnerable tissues.
geschlossene Kelchhaube
intakte Blütenhaube
sealed calyptra
unopened petal cap
verschlossene Kronblattkappe
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 99-102 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 93-95 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 191-194.
Intact Calyptra
Intactness quality monitoring in machine-harvested Chardonnay for premium sparkling wine production requires rapid assessment of juice samples for soluble solids, pH, and brown color development as indicators of skin breach frequency, with intact-berry fermentation protocols demanding less than 5% berry damage to prevent excessive phenolic extraction and oxidation vineyards achieving greater than 95% berry intactness through careful harvester adjustment (striker bar speed, conveyor cushioning) and optimal harvest timing (dawn temperatures, moderate maturity) produce must with lower potassium, reduced browning potential, and finer mousse development in the finished wine.
Intactness_Quality represents the structural integrity condition where a grapevine organ is complete with all its parts present and properly connected, without missing tissue, holes, or partial loss of structure. Intactness is the baseline expectation for healthy, undamaged organs and serves as the reference condition against which various forms of damage and deterioration are assessed. An intact berry possesses complete skin enclosure without tears or punctures, full complement of flesh and seeds, and secure attachment to its pedicel. An intact leaf displays complete blade without holes, tears, or missing lobes, undamaged margins, and functional petiole connection. An intact cluster retains all properly developed berries on their pedicels with an undamaged rachis framework. Intactness can be compromised by numerous factors including insect feeding that removes tissue, fungal decay that degrades and consumes tissue, mechanical abrasion that tears or punctures surfaces, physiological disorders that cause tissue collapse, and environmental damage from hail, frost, or sunburn. The proportion of intact organs in a sample population provides a primary quality metric, with high intactness percentages indicating successful crop protection and careful handling. Intactness assessment requires careful visual examination to detect subtle damage that might be overlooked in rapid inspection, with documentation of the type and extent of any damage observed. For berries, intactness of the skin envelope is particularly critical because any breach provides entry for decay organisms and increases water loss.
Intactness_Quality represents the ideal condition of complete, undamaged organs and serves as the reference against which Damage_State_Quality is assessed. The distinction between intact and damaged involves threshold judgments, as minor surface abrasions or tiny punctures may or may not be classified as compromising intactness depending on the assessment context. For practical quality grading, intactness typically requires absence of visible damage at inspection distance (approximately 30 cm for table grapes). Perfect intactness is expected at fruit set and early development deviations from intactness during development or at harvest indicate specific causal factors that should be identified and managed.
Unversehrtheit
Vollständigkeit
completeness
intact condition
intakter Zustand
unbeschädigter Zustand
undamaged state
wholeness
Crisosto CH and Mitchell FG (2002) Postharvest Handling Systems: Table Grapes. In: Kader AA (ed) Postharvest Technology of Horticultural Crops, 3rd Ed, University of California Agriculture and Natural Resources Publication 3311, pp.: 357-363 Ky I et al. (2012) Assessment of Grey Mould (Botrytis cinerea) Impact on Phenolic and Sensory Quality of Bordeaux Wines. Australian Journal of Grape and Wine Research 18(2): 215-224
Intactness Quality
Tempranillo vines in Rioja bearing an integrated production role are managed using pheromone traps monitoring Lobesia botrana at action thresholds of 15-20 moths/trap/week before chemical intervention, predatory mite Typhlodromus pyri populations maintained at 1-2 mites/leaf for Panonychus ulmi biocontrol, selective botryticide application only when Botrytis cinerea infection risk exceeds 30% on the Böhm model, and targeted nitrogen fertilization based on petiole analysis with NO₃⁻ targets of 500-1200 ppm at véraison.
A sustainable production role that inheres in a grapevine managed under integrated production (IP) principles that combine biological, cultural, and chemical control methods based on economic injury thresholds and monitoring-driven decision-making, applying synthetic interventions only when biological and cultural measures prove insufficient.
Integrated_Production_Role implements the IOBC (International Organisation for Biological and Integrated Control) framework applied to viticulture. The core principle is threshold-based decision-making: interventions are applied only when monitoring indicates pest or disease pressure exceeds economic injury levels. This distinguishes IP from both conventional production (calendar-based spraying) and organic production (categorical input restrictions). Many European wine regions mandate or incentivize IP through quality assurance schemes such as the Swiss Vitiswiss IP program or the Austrian IP-Weinbau guidelines.
IP-Anbau-Rolle
IPM viticulture role
integrated crop management role
integrated pest management role
integrierte Bewirtschaftungsrolle
integrierte Produktionsrolle
IOBC-WPRS Commission on Integrated Production Guidelines (2004) Boller EF et al. (2004) Integrated production: principles and technical guidelines, IOBC-WPRS Bulletin 27(2) Pertot I et al. (2017) A critical review of plant protection tools for reducing pesticide use on grapevine, Plant Pathology 66: 3-18
Integrated Production Role
Integrated water use efficiency assessment using δ¹³C in Riesling wine from different vineyard blocks reveals spatial variation in seasonal water status, with irrigated blocks showing δ¹³C of -27.2‰ (high Ci from open stomata, lower integrated WUE), rain-fed blocks showing -25.1‰ (lower Ci from stress-induced stomatal closure, higher WUE), and the 2.1‰ difference providing an integrated record of the water use strategy differences during the growing season that could not be captured by any single gas exchange measurement, enabling retrospective vineyard water management assessment and terroir documentation.
Integrated water use efficiency quality (δ¹³C) is a transpiration efficiency quality that represents the long-term average ratio of carbon assimilation to transpiration as recorded in the carbon isotope composition of plant tissues, characterizing the time-integrated water use strategy through the discrimination against ¹³C during photosynthesis that varies with the balance between stomatal conductance and photosynthetic capacity over the period of tissue formation. Within the BFO framework, this quality specifically addresses the integrated dimension of water use efficiency, distinguishing δ¹³C-based assessment from instantaneous gas exchange measurements while recognizing that carbon isotope discrimination provides a time-integrated record of the intercellular CO₂ concentration (Ci) that reflects the balance between CO₂ supply (stomatal conductance) and demand (photosynthesis) over days to months of tissue growth. The ontological significance lies in δ¹³C providing a retrospective measure of water use efficiency that integrates across the variable conditions experienced during tissue formation, where less negative δ¹³C values (less discrimination against ¹³C) indicate lower Ci from restricted stomatal conductance relative to photosynthetic demand, corresponding to higher water use efficiency achieved through conservative stomatal behavior. The isotopic basis for δ¹³C as a WUE proxy involves the fractionation during CO₂ diffusion through stomata (4.4‰) and during carboxylation by Rubisco (approximately 27‰), with the net discrimination (Δ) depending on the ratio of Ci to ambient CO₂ (Ca), such that Δ = 4.4 + (27 - 4.4) × Ci/Ca, and with lower Ci/Ca indicating more closed stomata and higher WUE. The measurement and interpretation of δ¹³C employs mass spectrometry analysis of dried plant tissue (commonly leaves, must, or wine), with typical values for Vitis vinifera ranging from -28‰ to -24‰ depending on water status during growth, and with less negative values indicating higher integrated WUE from water stress-induced stomatal limitation. The applications of δ¹³C analysis include vineyard water status assessment (spatial variation indicating irrigation uniformity), cultivar screening for water use efficiency, terroir characterization (wine δ¹³C reflecting vineyard water relations), and climate change research (historical samples revealing long-term WUE trends). Understanding integrated water use efficiency quality enables retrospective assessment of vine water status, cultivar comparison for long-term efficiency, and terroir fingerprinting through isotopic signatures recording water use history.
Integrated_Water_Use_Efficiency_Quality_(Delta_13C) addresses the isotopic record of time-integrated water use efficiency in plant tissues. This quality provides retrospective WUE assessment over tissue formation periods. The class supports water status assessment, cultivar screening, and terroir characterization through δ¹³C analysis.
Kohlenstoffisotopendiskriminierung
carbon isotope discrimination
integrated WUE
integrierte Wassernutzungseffizienz
Δ¹³C
δ¹³C
Farquhar GD et al. (1989) Carbon isotope discrimination and photosynthesis. Annual Review of Plant Physiology and Plant Molecular Biology 40: 503-537 Gaudillère JP et al. (2002) Carbon isotope composition of sugars in grapevine, an integrated indicator of vineyard water status. Journal of Experimental Botany 53(369): 757-763
Integrated Water Use Efficiency Quality (Delta 13C)
Merlot in Bordeaux shows integument development during July-August Year N+1, with ovule integuments differentiating following fertilization, outer integument cells elongating radially 2.5-4 fold increasing from 15-25 to 45-75 micrometers length, cell expansion and possible divisions increasing tissue thickness, primary cell wall synthesis establishing structure, inner integument cells differentiating with limited expansion maintaining 12-22 micrometers dimensions, organized cellular layers establishing coat architecture, lignin biosynthesis genes beginning 2-5 fold upregulation by week 6-8 preparing for lignification, under oceanic conditions of 20-26°C temperature, typical of this mid-season cultivar initiating seed coat development
Integument development represents the seed coat layer differentiation phase during seed coat formation when ovule integuments begin differentiating into seed coat layers following fertilization, outer integument cells initiating development toward testa protective layer, inner integument cells developing toward tegmen inner layer, cellular differentiation establishing coat architecture, and early coat structure preparing for subsequent lignification and maturation, occurring during early-mid seed development weeks 0-8 post-fertilization establishing coat foundation. This layer differentiation phase occurs during summer in July-August in temperate regions Year N+1, representing initial seed coat development. Development mechanism involves fertilization triggering integument differentiation from ovule protective layers to seed coat structures, outer integument cell elongation and expansion, inner integument cell differentiation though less extensive, cell wall synthesis establishing primary walls, and developmental programming directing coat formation. Ovule integument characteristics shows ovule containing two integuments surrounding nucellus and embryo sac prior to fertilization, outer integument typically 3-5 cell layers thick, inner integument 2-4 cell layers, integuments as protective envelopes around female gametophyte, and integuments as seed coat precursors. Outer integument pre-fertilization shows outer integument as outermost ovule layer, integument with parenchymatous cells, integument forming part of micropyle opening, and integument poised for differentiation. Inner integument pre-fertilization shows inner integument between outer integument and nucellus, thinner than outer integument, cells small and compact, and inner layer as secondary envelope. Fertilization trigger shows double fertilization initiating seed development, fertilization signaling integument differentiation, developmental shift from ovule protection to seed coat formation, and fertilization as developmental switch. Outer integument differentiation shows outer integument cells beginning expansion and elongation, cells elongating radially outward increasing integument thickness, cell division possibly occurring increasing cell number, and outer development establishing testa foundation. Cell elongation patterns shows cells elongating predominantly in radial direction, elongation increasing tissue thickness, elongated cells creating organized layer structure, and elongation as morphogenetic process. Cell expansion shows cell volume increase from water uptake and vacuole enlargement, cells expanding 2-5 fold from initial size, expansion without necessarily division, and expansion contributing to tissue growth. Cell division shows possible mitotic divisions in some integument regions, divisions increasing cell number, division patterns maintaining layer organization, and divisions contributing to tissue development. Outer integument cell fate shows cells programmed for testa differentiation, eventual cell wall thickening and lignification, cells eventually dying leaving lignified walls, and fate as outer protective layer. Inner integument differentiation shows inner integument cells undergoing differentiation though less extensive than outer, cells maintaining smaller size compared to outer integument, primary wall synthesis establishing cell structure, and inner layer developing toward tegmen. Inner integument fate shows cells forming tegmen inner coat layer, less extensive lignification than testa, cells maintaining viability longer than testa cells, and fate as inner layer. Cell wall synthesis shows primary cell wall deposition establishing cell structure, cellulose microfibrils and matrix polysaccharides synthesized, walls providing cellular integrity, and primary walls as foundation for subsequent secondary wall deposition. Cellulose synthesis shows cellulose synthase complexes in plasma membrane synthesizing cellulose microfibrils, microfibrils deposited in wall providing structural framework, and cellulose as major wall component. Matrix polysaccharides shows hemicelluloses and pectins synthesized and secreted, matrix filling spaces between microfibrils, matrix affecting wall properties, and polysaccharides as wall components. Tissue organization shows integuments developing organized cellular layers, cell files maintaining radial organization, tissue architecture establishing coat structure, and organization as developmental feature. Vascular development shows possible vascular traces extending into integuments supplying developing coat, vascular supply enabling nutrient and water delivery, and vasculature supporting development. Integument growth shows integuments enlarging to accommodate growing embryo and endosperm, integument growth coordinated with internal tissue development, eventual integument size determining seed coat dimensions, and growth establishing coat size. Differentiation timing shows differentiation initiating immediately post-fertilization, progressive differentiation over weeks 0-8, differentiation preparing for lignification phase, and timing as developmental progression. Differentiation molecular mechanisms shows gene expression changes directing integument to seed coat fate, transcription factors regulating coat development, hormonal signals possibly regulating differentiation, and molecular programs directing development. Gene expression shows coat-specific genes upregulating, genes encoding cell wall biosynthesis enzymes, lignin biosynthesis pathway genes beginning expression, and expression establishing coat program. Hormonal regulation shows auxin possibly involved in integument development, other hormones potentially regulating coat formation, and hormonal coordination of development. Environmental effects on integument development include temperature affecting development rate with optimal 22-28°C supporting normal differentiation, nutrient availability supporting cell growth and wall synthesis, and conditions affecting early coat development. Temperature effects shows warm conditions promoting faster differentiation, cool conditions slowing development, and temperature as rate factor. The process represents first phase of seed coat formation establishing layer foundation, preceding testa lignification and seed coat hardening. Duration approximately 8 weeks representing early differentiation weeks 0-8. Regional timing shows integument development during July-August as coat begins formation. Viticultural significance includes representing seed coat initiation establishing protective structure foundation, determining coat architecture and cell organization, preparing for subsequent lignification creating protection, and establishing developmental basis for mature seed coat. Research applications include analyzing integument differentiation regulation and mechanisms, investigating cell fate determination and development, evaluating gene expression and molecular control, quantifying structural development and growth, and assessing integument development as seed coat formation foundation.
Integument_Development is first phase of Seed_Coat_Formation, distinguished by layer differentiation. Represents weeks 0-8. Ovule integuments (outer 3-5 layers, inner 2-4 layers) differentiate post-fertilization. Outer → testa (cell elongation 2-5 fold radially, expansion), Inner → tegmen (limited differentiation). Primary cell wall synthesis. Gene expression: coat-specific, lignin pathway genes beginning. Precedes Testa_Lignification. Establishes coat foundation. Temperature 22-28°C optimal.
Integument-Entwicklung
Integument-zu-Schale-Übergang
Schalen-Schicht-Formation
Schicht-Differenzierung
coat layer formation
early coat development
integument to coat transition
layer differentiation
Integument Development
Inter-plant synchrony assessment in a variable-soil Pinot Noir block reveals the soil-driven developmental patterns affecting management efficiency, with shallow clay zone (higher water holding, cooler temperatures) showing delayed phenology (véraison 8 days later than block mean) and internal uniformity (CV 3.8% among vines within zone), deep gravel zone (lower water holding, warmer) showing advanced phenology (véraison 6 days earlier than mean) with similar internal uniformity (CV 4.2%), and transitional zone showing intermediate timing but lower uniformity (CV 9.5% reflecting mixed soil influences), demonstrating how soil variation creates inter-plant synchrony patterns that suggest zone-based management rather than whole-block treatment.
Inter-plant synchrony quality is a synchronization quality that represents the temporal coordination of developmental events among different grapevines within a population, characterizing the uniformity of phenological progression across multiple plants that determines whether vineyard blocks can be effectively managed and harvested as coherent units or require plant-by-plant or zone-by-zone treatment to accommodate developmental heterogeneity. Within the BFO framework, this quality specifically addresses the among-vine dimension of synchronization, distinguishing coordination across multiple plants from coordination within single plants while recognizing that inter-plant synchrony determines the efficiency of block-level operations and the feasibility of treating vineyard units as homogeneous for management purposes. The ontological significance lies in inter-plant synchrony determining vineyard management efficiency and harvest logistics, where high synchrony enables single-pass operations across entire blocks (one spray timing appropriate for all vines, one harvest date optimal for all vines), while low synchrony requires either multiple passes to treat vines at appropriate stages, subdivision of blocks into more uniform management zones, or acceptance of suboptimal timing for some portion of the vines. The factors affecting inter-plant synchrony include genetic uniformity (clonal plantings showing higher synchrony than mixed plantings), rootstock uniformity (consistent rootstocks promoting uniform development), soil uniformity (consistent soil properties supporting similar vine development), microclimate uniformity (avoiding hot spots, frost pockets, or other microclimate variation), and management uniformity (consistent pruning, irrigation, and nutrition across all vines), with these factors creating the spatial patterns of developmental uniformity or heterogeneity that characterize vineyard blocks. The practical implications of inter-plant synchrony extend throughout the growing season, affecting spray timing efficiency (synchronized blocks enabling optimal application timing for all vines), canopy management scheduling (uniform development enabling single-pass operations), and harvest logistics (synchronized ripening enabling single-harvest capture of optimal maturity across the block), with the economic value of high synchrony reflected in reduced labor requirements and improved quality consistency. Understanding inter-plant synchrony quality enables assessment of vineyard block uniformity, identification of factors creating spatial developmental patterns, and appropriate management zoning that groups vines of similar phenology for efficient treatment.
Inter_Plant_Synchrony_Quality serves as the parent class for population-level synchrony at different scales (Block, Cultivar). This quality addresses coordination among vines determining management efficiency and harvest logistics. The class supports vineyard zoning and management planning through inter-plant synchrony characterization.
Blockuniformität
Inter-Pflanzensynchronie
Populationssynchronie
Rebe-zu-Rebe-Uniformität
block uniformity
population synchrony
vine-to-vine synchrony
vineyard-level coordination
Bramley RGV (2005) Understanding variability in winegrape production systems 2. Within vineyard variation in quality over several vintages. Australian Journal of Grape and Wine Research 11(1): 33-42 Taylor JA et al. (2010) A protocol for delineating management zones in vineyards based on grape yield and quality. Journal of Wine Research 21(2-3): 151-168
Inter Plant Synchrony Quality
Intercellular space analysis of Thompson Seedless table grapes using micro-CT imaging reveals connected void network comprising 85% of total porosity in fresh berries, declining to 62% connectivity after controlled atmosphere storage at 5% O₂ for 4 weeks as tissue collapse and water loss compress previously open channels, correlating with the observed 45% reduction in respiration rate indicating diffusion limitation rather than metabolic suppression as the primary mechanism of respiration decline.
Intercellular space quality is a porosity quality that inheres in the apoplastic void network within grape berry tissues and represents the extent, connectivity, and geometric characteristics of air-filled channels formed at the junctions between adjacent cells where middle lamella separation creates continuous pathways for gas diffusion throughout the tissue matrix. Within the BFO framework, this quality specifically characterizes the intercellular air space system as a distinct structural feature, distinguishing the void network geometry from the aggregate porosity measurement that quantifies total void volume without addressing spatial organization or connectivity. The ontological significance lies in intercellular space organization determining not just the volume available for gas accumulation but the effective diffusivity for respiratory gases and volatile compounds, where connected spaces provide efficient transport pathways while isolated voids trap gases without enabling equilibration with external atmosphere. Intercellular spaces in grape berry mesocarp form primarily at tricellular junctions where three adjacent cells meet, with space dimensions expanding during development as cells enlarge and middle lamella components are modified, typically reaching 10-50 μm diameter at maturity. The connectivity of the intercellular space network affects berry response to hypoxic conditions during storage, determines the rate of internal oxygen depletion during post-harvest respiration, and influences the development of fermentative metabolism when oxygen supply becomes limiting. Measurement approaches for intercellular space characterization include scanning electron microscopy of tissue sections, confocal microscopy with fluorescent labeling of air-liquid interfaces, and micro-computed tomography providing three-dimensional reconstruction of the void network. Understanding intercellular space quality supports prediction of berry storage behavior, interpretation of modified atmosphere packaging responses, and modeling of gas exchange dynamics relevant to controlled atmosphere storage.
Intercellular_Space_Quality specifically addresses the geometric and topological characteristics of the void network, complementing Tissue_Porosity_Quality which quantifies aggregate void volume. This class emphasizes connectivity and spatial organization as determinants of functional properties beyond simple void fraction. The distinction between intercellular spaces (extracellular, air-filled) and substomatal cavities (connected to atmosphere) may be relevant for gas exchange modeling at the whole-berry scale.
Interzellularluftraum
Interzellularraum
Zellzwischenraum
air channel network
apoplastic void system
apoplastisches Hohlraumsystem
gas space connectivity
intercellular air space
Saladie M et al. (2007) Changes in cell wall composition during fruit ripening. Stewart Postharvest Review 3(4): 1-14 Harker FR, Hallett IC (1992) Physiological changes associated with development of mealiness of apple fruit during cool storage. HortScience 27(12): 1291-1294
Intercellular Space Quality
Interior leaf layer quality evaluation in Barossa Valley Shiraz compared outcomes across shoot thinning intensities in a vigorous vineyard. Unthinned vines at 22 shoots per meter produced high interior leaf layer quality with 52% of point quadrat contacts classified as interior leaves, mean total LLN of 2.9, and fruit zone PAR of only 6% ambient. Moderate thinning to 16 shoots per meter reduced interior leaf layer quality to 38% interior contacts, LLN 2.1, and fruit zone PAR of 14%. Aggressive thinning to 10 shoots per meter achieved optimal interior leaf layer quality with only 24% interior contacts, LLN 1.4, and fruit zone PAR of 23%, demonstrating that shoot density manipulation primarily addresses canopy problems through reduction of interior rather than exterior foliage, as retained shoots expand to maintain exterior canopy continuity while interior congestion resolves.
Interior leaf layer quality is a leaf layer number quality that inheres in a grapevine canopy system and represents the density of foliage positioned within the canopy volume beyond the exterior surface, comprising leaves that exist in partially or heavily shaded conditions created by overlying exterior foliage and that contribute to overall canopy leaf layer number through additional contacts during point quadrat transects penetrating past the external canopy surface. This quality characterizes the potentially problematic accumulation of interior leaves that receive insufficient light for optimal photosynthesis while contributing to canopy congestion, fruit zone shading, humidity retention, and spray interception that compromise fruit quality and disease management outcomes. The ontological significance of interior leaf layer quality lies in its representation of the difference between total canopy leaf area and the portion positioned in productive exterior locations, with high interior leaf layer quality indicating substantial investment in foliage that operates below photosynthetic light saturation and contributes primarily to canopy density problems rather than productive carbon gain. Interior leaves in dense grapevine canopies typically receive less than 30% of incident photosynthetically active radiation, operating on the linear portion of the photosynthetic light response curve where their carbon gain represents only 20-40% of that achieved by well-exposed exterior leaves, while their presence creates the multi-layer shading that reduces interior light levels below thresholds needed for fruit phenolic development. The point quadrat measurement of interior leaf layer quality derives from contacts occurring after the first leaf encounter, with percentage interior leaves computed as the proportion of total contacts representing second or subsequent contacts on a given probe insertion. High percentage interior leaves values exceeding 30-40% indicate problematic density typically associated with excessive vigor, inadequate shoot positioning, or abundant lateral development filling spaces between primary shoots. Management interventions targeting interior leaf layer quality include shoot thinning to reduce overall canopy fill, hedging to remove laterals that accumulate in canopy interior, and leaf removal that can directly extract interior foliage though this approach is labor-intensive and may create wound entry points for pathogens. Training system design fundamentally influences interior leaf layer quality potential, with narrow canopy systems like vertical shoot positioning inherently limiting the interior volume available for leaf accumulation compared to wider canopy architectures.
Interior_Leaf_Layer_Quality is_a Leaf_Layer_Number_Quality, representing the inner canopy density dimension corresponding to the percent interior leaves metric from point quadrat analysis. This class addresses the suboptimal foliage positioned beyond the productive exterior canopy surface. Distinguished from External_Leaf_Layer_Quality which addresses outer canopy characteristics. Values exceeding 30-40% interior leaves indicate problematic density requiring intervention. This quality class links vigor management to canopy microclimate outcomes by identifying where excess growth accumulates.
Innenblattdichte
Kroneninnendichte
inner canopy leaf layers
innere Laubschicht
interior foliage density
percentage interior leaves
secondary leaf layer contacts
sekundäre Blattschichten
Smart RE (1985) Principles of grapevine canopy microclimate manipulation with implications for yield and quality: a review. American Journal of Enology and Viticulture 36:230-239 Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 44-59 Schultz HR, Kiefer W (2015) Canopy structure and dry matter production of grapevines trained to vertical and horizontally divided canopy systems and their relationship to wine quality. OENO One 49:239-248
Interior Leaf Layer Quality
Syrah intermediate middle pericarp radial cell size measurements showing systematic gradients= outer-to-central intermediate zone (300-800 μm depth from outer pericarp boundary) with cells progressing from 92 μm (at 300 μm depth) through 115 μm (500 μm depth) to 138 μm diameter (800 μm depth, merging into central zone), representing smooth transition from smaller outer middle pericarp to larger central middle pericarp central-to-inner intermediate zone (200-600 μm from inner pericarp boundary) with cells declining from 165 μm (600 μm from boundary) through 142 μm (400 μm) to 118 μm (200 μm, merging into inner pericarp), demonstrating continuous size gradients rather than discrete tissue zones. Cabernet Sauvignon intermediate middle pericarp composition analysis across radial positions= outer-to-central intermediate (cells 105-135 μm) showing 23.8-24.5% sugars, 6.4-6.8 g/L acids central middle (cells 145-165 μm) showing 24.2-25% sugars, 6.2-6.6 g/L acids central-to-inner intermediate (cells 115-145 μm) showing 24-24.8% sugars, 6.3-6.9 g/L acids, demonstrating minimal compositional gradients (all within ±3% variation) despite substantial cell size gradients, suggesting sugar and acid distribution equilibrates across middle pericarp despite cell size heterogeneity. Merlot intermediate middle pericarp vascular proximity effects= cells within 150 μm of lateral vascular branches (near-vascular intermediate zones) showing slightly earlier veraison sugar increase (sugars reaching 16% by week 11 versus 14.5% in distant zones at same time), suggesting vascular proximity creates slight temporal gradients in sugar import though final maturity composition converges (all zones 24-25.5% sugars at week 18), demonstrating that intermediate zones near vasculature may show earlier but not higher sugar accumulation. Chardonnay intermediate middle pericarp cell wall properties across zones= outer-to-central intermediate showing wall thickness 0.52-0.68 μm, central showing 0.45-0.58 μm, central-to-inner intermediate showing 0.56-0.72 μm, with all middle pericarp walls substantially thinner than outer pericarp (1.8-2.8 μm) or inner pericarp (0.9-1.4 μm), demonstrating gradual wall thickness variations within middle pericarp while maintaining overall thin-wall character distinguishing middle from outer/inner pericarp.
An intermediate middle pericarp is the middle pericarp tissue in zones between outer and central regions, or between central and inner regions, showing transitional characteristics in cell size, composition, and properties, representing middle pericarp areas with intermediate features rather than the maximum characteristics of central middle pericarp or the boundary characteristics of outermost/innermost middle pericarp. Intermediate middle pericarp comprises= (1) outer-to-central intermediate zone located between outer pericarp boundary and central middle pericarp maximum size region, showing progressive cell size increase from outer middle pericarp (80-120 μm diameter) toward central middle pericarp (110-180 μm), with intermediate zone cells typically 95-140 μm diameter representing the transition region (2) central-to-inner intermediate zone located between central middle pericarp maximum size region and inner pericarp boundary, showing progressive cell size decrease from central middle pericarp (110-180 μm) toward inner middle pericarp (90-140 μm), with intermediate zone cells typically 100-155 μm (3) lateral intermediate zones showing gradients related to vascular bundle proximity, with cells closer to major vascular bundles potentially showing different characteristics (smaller size, different wall properties) than cells distant from bundles. Intermediate middle pericarp cellular characteristics show gradual transitions= cell size intermediate between adjacent zones cell wall thickness potentially showing gradual changes from very thin central walls (0.4-0.6 μm) toward slightly thicker boundary walls (0.6-1 μm) vacuolar sugar and acid content potentially showing gradients though less pronounced than size gradients phenolic content remaining low throughout middle pericarp with intermediate zones showing minimal variation from central zones. Intermediate middle pericarp may show positional effects= cells in outer-to-central intermediate zone receiving sugars earlier during phloem import phase potentially showing slightly earlier or more rapid sugar accumulation cells in central-to-inner intermediate zone adjacent to seeds potentially receiving seed-derived signals affecting metabolism or composition cells adjacent to vascular bundles receiving direct import before distribution to more distant cells. Intermediate middle pericarp represents the bulk of transition zones within middle pericarp, with smooth gradients rather than sharp boundaries characterizing middle pericarp organization, reflecting continuous rather than discrete tissue zonation within the flesh region. The biological significance of intermediate middle pericarp gradients relates to developmental constraints= cell expansion limited by mechanical constraints near rigid outer pericarp creating smaller cells in outer zones cell expansion potentially limited by space constraints near developing seeds creating smaller cells in inner zones vascular proximity affecting resource availability creating local variations in cell properties these position-dependent factors creating systematic gradients in cell size and potentially other properties.
Intermediate_Middle_Pericarp represents transition zones within middle pericarp and is distinguished from Transitional_Middle_Pericarp which may emphasize boundaries between middle and outer/inner pericarp. Intermediate middle pericarp shows smooth gradients in cell size and properties reflecting position-dependent developmental effects. Understanding intermediate middle pericarp gradients is relevant for detailed anatomical characterization and for understanding spatial heterogeneity in berry tissues, though practical implications are limited as middle pericarp functions as integrated tissue in wine production with minimal functional zonation.
Gradienten-Fruchtfleisch
gradient zone
intermediäres mittleres Perikarp
transitional flesh
Übergangs-Mesokarp
Harris, J.M. et al. (1968). The Vinifera grape berry= anatomical study. Botanical Gazette 129=1-23 Nunan, K.J. et al. (2001). Changes in cell wall composition during ripening. Plant Physiology 126=520-528.
Intermediate Middle Pericarp
Internal texture quality drives consumer preference in table grape markets, with distinct texture profiles commanding premium prices in different regions: Asian markets strongly prefer crisp, firm-fleshed cultivars like Shine Muscat (flesh firmness greater than 2.5 N, high fracturability) that produce an audible crunch when bitten, while European consumers accept softer-fleshed Muscat varieties (firmness 1.5-2.0 N) with juicy, melting texture understanding these internal texture preferences guides cultivar selection and market targeting in global table grape trade where texture mismatches result in significant market discounts.
Internal_Texture_Quality encompasses all texture qualities that characterize the structural and mechanical properties of tissues beneath the surface of grapevine organs, particularly the flesh and pulp of grape berries. Internal texture determines eating quality in table grapes, processing behavior in winemaking, and resistance to damage during handling and storage. This category addresses the tactile and mechanical sensations experienced when tissues are bitten, chewed, crushed, or pressed, reflecting cellular structure, cell wall properties, turgor pressure, and intercellular adhesion. Key internal texture attributes include firmness (resistance to deformation), crispness (tendency to fracture rather than deform), juiciness (fluid release upon mastication), mealiness (grainy texture from cell separation), and the balance between skin and flesh texture that determines overall mouthfeel. Internal texture results from the integration of cellular properties including cell size and shape, cell wall thickness and composition, pectin chemistry, turgor pressure, and the strength of middle lamella adhesion between cells. During ripening, berry internal texture typically softens as cell walls are enzymatically modified, pectin is solubilized, and turgor declines, transforming the firm, turgid pre-véraison berry into softer, more deformable ripe fruit. The rate and extent of softening vary among cultivars, distinguishing crisp table grape varieties from soft wine grape varieties. Assessment of internal texture employs sensory evaluation by trained panels, instrumental texture analysis measuring force-deformation relationships during compression or penetration, and acoustic measurement detecting crispness through sound emission.
Internal_Texture_Quality addresses tissue properties beneath the organ surface, complementing Surface_Texture_Quality. The child classes Pulp_Texture_Quality and Flesh_Consistency_Quality capture specific aspects of internal texture relevant to berry quality. Internal texture assessment requires destructive sampling in most cases, though non-destructive acoustic and near-infrared methods are under development. The distinction between firmness (resistance to deformation) and crispness (tendency to fracture) represents important textural properties with different underlying mechanisms. Berry internal texture changes dramatically during ripening and continues to evolve during storage, requiring specification of developmental stage for reproducible assessment.
Fruchtfleischbeschaffenheit
Gewebestruktur
Innentextur
flesh texture
innere Textur
internal structure quality
subsurface texture
tissue texture
Rolle L et al. (2012) Instrumental Texture Analysis Parameters as Markers of Table-Grape and Winegrape Quality: A Review. American Journal of Enology and Viticulture 63(1): 11-28 Sato A and Yamane H (2003) Possibility of the Instrumental Measurement of Grape Flesh Texture. Acta Horticulturae 603: 377-380
Internal Texture Quality
Rkatsiteli accession Vas 2691Mtp8 originally collected in Georgia and maintained at INRAE Domaine de Vassal in Marseillan, France bearing an international collection role is one of approximately 2,800 accessions from 54 countries of origin in the world's largest grapevine germplasm repository, representing all major Vitis species and over 2,300 unique cultivar genotypes authenticated through comprehensive SSR profiling, maintained under strict phytosanitary quarantine protocols including ELISA and RT-PCR testing for 15 regulated viruses, with the entire collection designated as a French National Heritage Collection and accessible to researchers worldwide through INRAE's online accession catalogue and material distribution under SMTA terms.
A germplasm repository role that inheres in a grapevine maintained within a germplasm collection of global scope that systematically acquires and conserves grapevine genetic diversity from multiple countries and continents, serving as a centralized reference repository for the international research community and providing a safety backup for national collections.
International_Collection_Role describes the highest tier of germplasm conservation scope, represented by a small number of institutions with global collection mandates. INRAE Domaine de Vassal (France), the USDA-ARS Plant Genetic Resources Unit at Geneva, New York, and the JKI Institut für Rebenzüchtung at Siebeldingen, Germany constitute the three principal international Vitis collections, collectively conserving an estimated 80-85% of documented global grapevine cultivar diversity: the planned relocation of the Domaine de Vassal collection to a new site at Gruissan (2020-2030) represents one of the largest living plant collection transfers in history. International collections serve irreplaceable functions as safety duplicates, inter-laboratory reference standards, and sources of genetic diversity for global breeding programs.
Weltsammlung-Rolle
global genebank role
globale Genbank-Rolle
international reference collection role
internationale Sammlungs-Rolle
world collection role
Lacombe T et al. (2011) Inventory and characterization of Vitis vinifera ssp. vinifera at INRA Domaine de Vassal, Vitis 50: 56-68 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157 FAO (2010) Second Report on the State of the World's Plant Genetic Resources for Food and Agriculture, Rome
International Collection Role
Chardonnay planted in the Bekaa Valley of Lebanon bearing an international variety role represents one of approximately 200,000 ha planted globally across 40+ countries, introduced to Lebanese viticulture in the 1990s alongside Cabernet Sauvignon, Merlot, and Syrah to complement indigenous cultivars such as Obaideh and Merwah, with the international varieties enabling Lebanese producers to access export markets through globally recognizable varietal labeling while indigenous varieties serve domestic and heritage-oriented market segments.
An introduced cultivar role that inheres in a grapevine cultivar of established global commercial prominence that has been planted across multiple continents and viticultural zones as a recognized component of the international varietal wine market, typically introduced to a region primarily for its brand recognition, consumer familiarity, and proven market demand rather than for site-specific terroir compatibility.
International_Variety_Role applies to the small group of cultivars (approximately 10-15) that collectively dominate global wine production and trade, including Cabernet Sauvignon (~340,000 ha worldwide), Merlot (~266,000 ha), Chardonnay (~210,000 ha), Syrah/Shiraz (~190,000 ha), Sauvignon Blanc (~120,000 ha), and Pinot Noir (~115,000 ha). The role captures the commercial-economic motivation for introduction rather than the cultivar's inherent properties: the same cultivar may simultaneously bear an International_Variety_Role in one region and a Traditional_Cultivar_Role in its region of origin. The global spread of these cultivars accelerated dramatically from the 1970s onward during the "international varietal revolution."
Weltsorten-Rolle
cosmopolitan variety role
global cultivar role
globale Rebsorten-Rolle
international variety role
internationale Sorten-Rolle
Anderson K and Aryal NR (2013) Which Winegrape Varieties are Grown Where?, University of Adelaide Press pp: 1-32 OIV (2023) Distribution of the World's Grapevine Varieties Robinson J et al. (2012) Wine Grapes, Allen Lane pp: 167-172
International Variety Role
The internode color description of Tempranillo documents green internodes with distinct red anthocyanin stripes on the dorsal (sun-exposed) surface (OIV 006:3) — the striping pattern reflects the localized anthocyanin accumulation in cortical tissue overlying the vascular bundles, producing 3-5 longitudinal red-brown stripes on a green background that is one of Tempranillo's most recognizable shoot characters during the growing season. This striped pattern is intermediate between the uniformly green internodes of Verdejo (OIV 006:1, no anthocyanin expression in internode tissue) and the uniformly red internodes of Garnacha Tintorera (Alicante Bouschet, OIV 006:5, complete anthocyanin coverage reflecting the teinturier genotype's constitutive pigmentation of all tissues including berry flesh), illustrating the range of internode pigmentation phenotypes across V. vinifera cultivars.
A shoot description component that records the color and pigmentation pattern of shoot internodes during the active growing season, documenting the green ground color intensity and the presence, intensity, and distribution of anthocyanin pigmentation — uniform, dorsal-only, or striped — according to OIV descriptor 006, with internode color providing cultivar-diagnostic information during the spring and summer growth period that complements the dormant cane color observed during winter.
Internode_Color_Description captures the growing-season shoot pigmentation that is observable from E-L stage 9 (2-3 leaves unfolded with shoots 10-20 cm) through véraison (E-L stage 35) before the onset of lignification progressively obscures the green shoot color with brown periderm. The anthocyanin pigmentation pattern of internodes provides particularly clear cultivar discrimination because three distinct phenotypic classes are readily distinguished in the field — green (no anthocyanin, typical of white cultivars and some red cultivars with low constitutive pigmentation), green with red stripes (anthocyanin confined to vascular bundle-associated cortical tissue, creating the distinctive longitudinal striping observed in many moderately pigmented red cultivars), and uniformly red (anthocyanin throughout the cortical tissue, observed in strongly pigmented cultivars): these three classes correspond to different levels of anthocyanin pathway activation in shoot tissue, controlled by the same VvMYBA regulatory genes that determine berry skin color but with additional tissue-specific regulatory elements that produce the quantitative variation from absent through striped to uniform expression. The light-dependency of internode anthocyanin expression — typically stronger on the dorsal (sun-exposed) surface than on the ventral (shaded) surface — necessitates standardized observation of the sun-exposed internode face for consistent cultivar comparison.
Grüntrieb-Färbung
Internodien-Farbbeschreibung
Trieb-Pigmentierung
green shoot coloration
internode pigmentation assessment
shoot color record
Internode Color Description
Pinot Noir in Burgundy shows internode development from late April through July over 12-14 weeks at BBCH 15-38, producing 14-18 internodes ranging from 35-45 mm in basal positions to 70-95 mm during peak growth declining to 25-40 mm terminally, creating final shoot length of 900-1300 mm with progressive lignification from base upward under continental climate conditions of 15-26°C temperature range, variable moisture, and moderate fertility typical of this moderate-vigor cultivar
Internode development represents the fundamental process creating shoot axis structure through sequential formation, elongation, and maturation of stem segments between successive nodes, establishing spatial framework for leaf, tendril, and inflorescence positioning that determines shoot architecture and canopy function. This essential developmental process occurs continuously during shoot growth from early elongation through growth cessation, typically spanning 8-16 weeks producing 12-20 internodes per shoot depending on vigor and conditions. Internode development encompasses initiation of new internode segments at shoot apex through intercalary meristem establishment, progressive elongation through cell division and expansion creating visible shoot extension, and maturation through lignification and vascular differentiation establishing permanent woody structure. The process shows coordinated progression along shoot axis with basal internodes completing development and lignifying while apical internodes continue forming and elongating, creating developmental gradient from mature lignified base to actively elongating apex to meristematic summit. Internode formation initiates at shoot apex immediately behind apical meristem where leaf primordia emergence defines nodal positions, with intercalary meristem establishing between successive nodes containing cells that will contribute to internode length through division and subsequent expansion. Cell division in intercalary meristems continues for limited period adding cells to internode, with division typically ceasing when internode reaches 5-15 mm length depending on ultimate dimensions, followed by extended expansion phase where cells increase length 10-50 fold creating final internode dimensions. Internode elongation proceeds through coordinated cell expansion in central elongation zone where cells undergo turgor-driven wall extension, with expansion rate and duration determining final internode length ranging from 20-40 mm in slow-growing shoots to 80-150 mm during rapid elongation periods. Vascular differentiation proceeds acropetally through developing internodes with procambial strands differentiating into functional xylem vessels for water transport and phloem sieve elements for assimilate distribution, establishing continuous vascular connection from permanent structures through shoot to developing apex. Lignification begins in basal internodes while apical regions continue elongating, with secondary wall deposition and lignin incorporation creating rigid woody structure providing mechanical support, with lignification front progressing acropetally at approximately same rate as elongation creating mature lignified shoot base while apex remains herbaceous. Internode structure includes epidermis providing protective surface, cortex containing photosynthetic and storage parenchyma, vascular bundles arranged in characteristic pattern conducting water and assimilates, and pith providing structural core with storage function. Internode length variation along shoot reflects changing growth conditions and resource availability, with basal internodes typically shorter reflecting slower early growth, middle internodes longest corresponding to peak elongation phase, and apical internodes progressively shorter as growth declines toward cessation. Mechanical properties develop progressively during maturation with young expanding internodes showing low tensile strength and flexibility from thin primary walls, transitioning to rigid strong structure as secondary walls develop and lignification proceeds providing self-supporting shoot architecture. Metabolic activity varies dramatically along developmental gradient with high respiration in expanding internodes supporting growth at 10-15 fold mature tissue rates, declining in maturing regions to maintenance levels, with photosynthetic activity in green cortex contributing modestly to carbon balance. Hormonal regulation involves auxin from apex promoting elongation and coordinating vascular differentiation in acropetal pattern, gibberellins supporting cell expansion particularly affecting internode length, cytokinins from roots promoting cell division in intercalary meristems, and ethylene potentially affecting lignification and maturation processes. Temperature strongly affects internode development rate with elongation approximately doubling per 10°C increase between 15-28°C optimal range, influencing both cell division rates in meristems and expansion rates in elongation zones. Light quality affects internode elongation through phytochrome-mediated responses with red: far-red ratio influencing length, with shade promoting elongated internodes through etiolation response seeking improved light exposure while high light supports shorter stouter internodes with enhanced photosynthetic capacity in cortex. Water availability critically affects internode elongation through direct effects on cell turgor driving expansion and indirect effects on photosynthesis providing assimilates, with water stress rapidly arresting elongation and can permanently limit internode length affecting final shoot architecture. Nutrient availability particularly nitrogen affects development through effects on protein synthesis supporting growth and photosynthesis, with adequate nutrition supporting maximum internode production and elongation while deficiency reduces both number and length limiting final shoot size. The process shows four distinguishable phases based on developmental progression, advancing from internode formation at apex through sequential formation of multiple internodes, elongation creating shoot length, and maturation establishing permanent structure. Cultivar variation appears in internode number produced per season, length characteristics affecting shoot architecture with some varieties producing numerous short internodes creating compact structure while others produce fewer longer internodes creating open architecture, and lignification timing affecting shoot mechanical properties. Regional timing shows internode development active throughout growing season from May-July in warm climates, May-August in temperate regions, and June-September in cool areas, with duration corresponding to shoot growth period. Viticultural management considerations include understanding internode development patterns informing shoot positioning and training operations, recognizing that internode length reflects growth conditions providing diagnostic information about vine status, managing factors affecting elongation to achieve desired canopy architecture, timing mechanical operations considering maturation state avoiding damage to unlignified tissues, and evaluating final internode patterns assessing seasonal growth outcomes and planning subsequent season management. Excessive internode elongation particularly in shade creates weak shoots with poor lignification susceptible to damage, while insufficient development limits photosynthetic capacity through reduced canopy size. Research applications include analyzing mechanisms regulating internode length including hormonal and environmental controls, evaluating genetic variation in internode characteristics affecting architecture and training requirements, quantifying developmental timing and progression patterns informing growth models, assessing environmental and management effects on development affecting final shoot structure, and documenting anatomical and structural development supporting understanding of woody tissue formation in perennial plants.
Internode_Development represents structural axis formation process within Shoot_Growth, occurring coordinately with Shoot_Elongation but focusing on segmental structure rather than overall length. Subdivided into First_Internode_Formation, Sequential_Internode_Formation, Internode_Elongation, and Internode_Maturation based on progression. Distinguished from Shoot_Elongation which emphasizes length increase while Internode_Development emphasizes segmental structure and maturation. Critical process determining shoot architecture and mechanical properties.
Achsenentwicklung
Internodienentwicklung
Stammabschnittbildung
Stängelgliedbildung
axis development
internode formation
stem development
stem segment development
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 205-235 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 128-152 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Internode Development
Merlot in Bordeaux shows internode elongation throughout May-July at BBCH 15-38, with individual internodes requiring 2-3 weeks to complete elongation from 5-10 mm initial length to final dimensions ranging 35-50 mm basally, 75-105 mm in middle positions during peak June growth, and 40-65 mm apically, under oceanic climate conditions of 16-26°C temperature range, variable precipitation with supplemental irrigation, and moderate fertility typical of this moderately vigorous mid-season cultivar
Internode elongation represents the primary growth process where stem segments increase length through coordinated cell expansion following completion of cell division phase, creating visible shoot extension that accounts for majority of final shoot length and establishing spatial framework for canopy architecture. This fundamental developmental process occurs in each internode following formation with cell division phase, typically spanning 2-4 weeks per internode from initiation of elongation to completion, with multiple internodes elongating simultaneously creating developmental gradient along shoot from actively elongating subapical zones to recently completed basal regions. Elongation proceeds through cell expansion in intercalary zones between nodes where cells increase length dramatically through turgor-driven wall extension, with cells expanding from meristematic dimensions of 10-20 micrometers to final lengths of 150-500 micrometers representing 10-50 fold increase in length. Cell expansion involves water uptake increasing cell volume creating turgor pressure of 0.4-0.8 MPa driving cell wall extension, wall loosening through expansin proteins and other wall-modifying enzymes allowing plastic deformation under pressure, and wall synthesis adding new material to maintain structural integrity during expansion. The elongation zone in each internode extends throughout internode length during early expansion phase, becoming progressively restricted to central region as cells near nodes complete expansion earlier than central cells, creating elongation pattern proceeding from ends toward middle then completing throughout. Elongation rate varies substantially among internodes along shoot reflecting changing resource availability and growth conditions, with basal internodes showing modest rates of 1-2 mm per day during early slow shoot growth, middle internodes showing maximum rates of 4-8 mm per day during peak growth, and apical internodes showing declining rates of 2-4 mm per day as growth decelerates. Final internode length results from combined effects of cell number produced during division phase and extent of cell elongation during expansion phase, with variation in both factors contributing to characteristic internode length pattern along shoot. Internode position affects final length with typical pattern showing basal internodes 25-45 mm from limited early resources, middle internodes 70-120 mm corresponding to peak conditions, and apical internodes 30-60 mm from declining late-season resources and accelerated maturation. Vascular tissues continue differentiating during elongation with procambial cells in longitudinal strands undergoing coordinated divisions and differentiation into functional xylem vessels with secondary wall thickening and lignification, and phloem sieve elements with specialized wall structures enabling translocation function. Water transport capacity through elongating internodes increases progressively as vessel diameter and number increase, with functional xylem maintaining adequate capacity for supporting transpiration from developing leaves and water requirements for continued elongation in more apical zones. Metabolic activity remains elevated during elongation with respiration rates 8-15 fold above mature levels providing energy for active transport supporting expansion, wall synthesis, and protein production, consuming carbohydrates from photosynthesis in mature leaves and stored reserves mobilized from permanent structures. Cell wall composition changes during elongation from thin primary walls rich in pectin allowing expansion to thickening walls with increased cellulose and hemicellulose providing progressive structural rigidity, with wall modification processes regulated by pH changes, enzyme activities, and hormonal signals coordinating expansion with wall strengthening. Protein synthesis operates at high rates during elongation producing structural proteins for expanding cell walls and membranes, enzymes including wall-modifying proteins enabling expansion, and beginning accumulation of photosynthetic proteins if cortex cells contain developing chloroplasts. Hormonal regulation centrally involves auxin from shoot apex promoting elongation through effects on cell wall extensibility and coordinating elongation pattern along shoot, gibberellins strongly affecting elongation with higher concentrations promoting increased cell expansion and longer final internode length, brassinosteroids enhancing cell expansion and vascular differentiation, and ethylene potentially limiting elongation as maturation processes initiate. Temperature strongly affects elongation rate with approximately doubling per 10°C increase between 15-28°C optimal range, influencing both rate of individual cell expansion and duration of elongation period before maturation processes dominate. Light quality affects elongation through phytochrome-mediated responses to red: far-red ratio, with shade conditions promoting enhanced elongation through etiolation response creating longer thinner internodes seeking improved light exposure, while high light conditions support normal to reduced elongation with stouter structure. Water availability critically affects elongation through direct effects on cell turgor driving expansion, with adequate moisture essential for maintaining pressure supporting wall extension, while water stress rapidly arrests elongation through turgor loss and can permanently limit final length even after stress relief. Nutrient availability particularly nitrogen affects elongation through effects on protein synthesis supporting wall expansion and enzyme production, with adequate nutrition supporting maximum elongation achieving genetic potential while deficiency reduces expansion rate and limits final dimensions. The process shows variation in timing and rate among internodes creating characteristic pattern along shoot, with basal internodes completing elongation early in season during slow growth phase, middle internodes elongating during peak growth achieving maximum length, and apical internodes elongating during declining growth showing progressive length reduction. Cultivar variation appears in maximum internode elongation rate and final lengths achieved, with vigorous cultivars showing rapid elongation to long dimensions 100-150 mm while less vigorous types show moderate elongation to shorter lengths 40-80 mm, affecting overall shoot architecture and training requirements. Regional timing shows internode elongation occurring throughout shoot growth period from May-July in warm climates, May-August in temperate regions, and June-September in cool areas, with individual internodes requiring 2-4 weeks for complete elongation depending on conditions and position. Viticultural management considerations include understanding that internode elongation pattern reflects growth conditions with length providing diagnostic information about resource availability during formation period, managing factors affecting elongation including water, nutrition, and light to achieve desired canopy architecture, recognizing that excessive elongation from shade or over-fertilization creates weak shoots requiring additional support, and timing canopy operations considering elongation state of target internodes. Research applications include analyzing cellular and molecular mechanisms of cell expansion including wall modification and hormonal regulation, quantifying environmental and genetic factors affecting elongation rates and final dimensions, evaluating relationships between elongation patterns and overall shoot growth characteristics, modeling elongation processes for predicting shoot development under varying conditions, and assessing implications of internode length variation for mechanical properties and canopy function.
Internode_Elongation is third of four Internode_Development subphases, distinguished by cell expansion phase following cell division creating final internode length. Occurs in each internode after Sequential_Internode_Formation cell production phase. Transitions to Internode_Maturation as expansion completes and lignification initiates. Primary determinant of final shoot length and architecture. Highly responsive to environmental conditions during expansion period.
Achsenstreckung
Internodienstreckung
Stielexpansion
Stängelgliedverlängerung
axis elongation
internode stretching
stem extension
stem segment expansion
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 220-235 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 140-152 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Internode Elongation
The internode length description of Pinot Noir documents short internodes (OIV 103:3) measuring 50-70 mm on mid-shoot positions of normally growing shoots on moderate-vigor rootstock — this compact growth habit, with 14-18 nodes per meter of shoot length, creates the dense, tightly spaced canopy characteristic of Pinot Noir that requires diligent leaf removal and shoot positioning for adequate fruit zone light exposure and air circulation. The short internode length contrasts with the long internodes of Syrah (OIV 103:7, 80-110 mm, 9-12 nodes per meter) and the very long internodes of Italia table grape (OIV 103:9, 100-130 mm, 7-10 nodes per meter), illustrating the 2-3 fold range of internode length variation across V. vinifera cultivars that profoundly influences canopy architecture and management requirements.
A shoot description component that records the length of internodes — the stem segments between consecutive nodes — on the growing shoot, measured at representative mid-shoot positions (typically nodes 5-10) and scored according to OIV descriptor 103 on a scale from very short (state 1, 120 mm), with internode length reflecting the cultivar-specific shoot elongation rate that influences vine vigor expression, canopy density, and the suitability of cultivars for specific training systems.
Internode_Length_Description documents a character that bridges ampelographic identification and viticultural management — internode length is both a cultivar-diagnostic feature and a primary determinant of canopy density, training system compatibility, and pruning labor requirements. The genetic control of internode elongation involves gibberellin biosynthesis and signaling pathways, with cultivar differences in internode length correlating with differences in endogenous gibberellin concentrations and sensitivity: short-internode cultivars (Pinot Noir, Muscat à Petits Grains) have compact growth resulting from lower gibberellin-driven cell elongation, while long-internode cultivars (Syrah, Grenache, table grapes) show greater gibberellin-mediated shoot elongation. The internode length character is moderately affected by environment — high nitrogen availability, excessive water supply, and vigorous rootstocks increase internode length by 20-40% compared to balanced or restricted conditions — but the relative ranking of cultivars is maintained across environments, with short-internode cultivars remaining consistently shorter than long-internode cultivars under any given growing conditions, making the proportional character genetically stable for identification purposes.
Internodienlänge-Beschreibung
Stammglied-Länge
Triebstreckung-Charakterisierung
internode dimension record
shoot elongation characterization
stem segment length
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Boss PK and Thomas MR (2002) Association of dwarfism and floral induction with a grape green revolution mutation, Nature 416: 847-850
Internode Length Description
Systematic phenotyping of Internode_Length_Disposition at Geisenheim University measured internode dimensions across 70 cultivars at multiple timepoints to characterize developmental patterns and final dimensions. Internodes 5-15 on primary shoots were measured at full extension, avoiding variable basal internodes and incompletely expanded distal internodes. Additionally, internode cell number and cell length were assessed anatomically to investigate developmental basis. Internode_Length_Disposition showed substantial cultivar variation with distinct developmental patterns. 'Silvaner' exhibited long internode characteristics with mean internode length of 10.2 cm, ranging from 8.5-12.5 cm across the measured zone, producing shoots with widely-spaced leaves and open architecture. Cell number per internode averaged 485 with mean cell length of 210 μm. 'Gewürztraminer' showed even longer internodes (11.5 cm) associated with its trailing growth habit. 'Pinot Noir' displayed short internode characteristics with mean length of 5.8 cm (range 4.8-6.8 cm), creating compact shoots with closely-spaced leaves. Cell number was similar (465 cells) but cell length was shorter (125 μm), indicating that cell expansion rather than cell division primarily determines internode length differences. 'Chardonnay' and 'Cabernet Sauvignon' showed intermediate lengths (7.2-7.8 cm). The functional significance was confirmed by canopy light measurements—long-internode 'Silvaner' achieved 45% fruit zone light penetration at equivalent shoot density where short-internode 'Pinot Noir' achieved only 28%, demonstrating that Internode_Length_Disposition substantially affects canopy microclimate.
An Internode_Length_Disposition is a Growth_Habit_Disposition that inheres in Vitis vinifera shoots and is realized through the characteristic distance between successive nodes on primary shoots, determining shoot compactness, the spatial distribution of leaves and clusters along the shoot, and contributing to canopy density and fruit zone architecture. This disposition represents the cell elongation and division patterns in the subapical meristem region that establish internode dimensions, reflecting genotypic differences in gibberellin sensitivity, cell expansion capacity, and the duration of the internode elongation phase. Internode length profoundly affects canopy structure—cultivars with long internodes produce open, extended shoots with widely-spaced leaves, while short-internode cultivars create compact shoots with closely-spaced, overlapping leaves. The disposition also affects cluster distribution and exposure, as clusters develop at nodes and internode length determines cluster spacing along the shoot. Long internodes are associated with lower shoot stiffness (contributing to trailing growth habit) and less dense canopies, while short internodes create stiffer shoots (contributing to upright habit) and potentially denser canopies at equivalent shoot numbers. Internode_Length_Disposition is quantified as mean internode length measured on primary shoots at standardized developmental stage, typically at full shoot extension when internode elongation is complete.
Internode_Length_Disposition is_a Growth_Habit_Disposition addressing the spacing between nodes. Internode length affects canopy density, cluster spacing, and shoot mechanical properties. Child classes include Long_Internode_Disposition and Short_Internode_Disposition. Cell expansion (length) rather than cell division (number) primarily underlies internode length variation. Long internodes are associated with trailing growth habit and lower shoot stiffness. Environmental factors (temperature, light, gibberellins) modulate expression, but genotypic ranking is stable.
Internodienlängen-Disposition
Knotenabstand
Triebkompaktheit
internode dimensions
node spacing
shoot compactness
Schultz HR (1992) An empirical model for the simulation of leaf appearance and leaf area development of primary shoots of several grapevine (Vitis vinifera L.) canopy systems, Scientia Horticulturae 52:179-200 Keller M et al. (2004) Warm spring temperatures induce persistent season-long changes in shoot development in grapevines, Annals of Botany 93:739-746 Louarn G et al. (2007) A method to assess light interception efficiency of a grapevine canopy based on shoot architecture and leaf area distribution, Australian Journal of Grape and Wine Research 13:29-38
Internode Length Disposition
Post-harvest assessment of Internode_Length_Quality in a stressed Riesling vineyard reveals a distinctive pattern with basal internodes averaging 4 cm formed during cool spring conditions, mid-shoot internodes of 7-8 cm formed before water stress onset, followed by abrupt shortening to 3-4 cm internodes coinciding with the mid-summer drought period, then partial recovery to 5-6 cm internodes following late-season rainfall, providing a permanent morphological record of the season's water availability timeline.
Internode_Length_Quality is a Vegetative_Growth_Vigor_Quality that inheres in a grapevine (Vitis vinifera L.) and represents the distance between consecutive nodes along a shoot, providing a permanent morphological record of growth conditions prevailing during each internode's formation. This quality serves as an integrative indicator of vine vigor because internode length reflects the cumulative effect of temperature, water availability, light exposure, and hormonal status during the specific developmental window when each internode undergoes extension growth. In BFO terms, Internode_Length_Quality exists as a one-dimensional spatial quality measurable along the shoot axis, with values varying systematically from shoot base to apex following patterns characteristic of cultivar, vigor status, and environmental conditions. The developmental sequence of internode formation creates a permanent record of seasonal growth conditions, with basal internodes forming under the cool temperatures of early spring typically measuring 3-5 cm, mid-shoot internodes formed during optimal growth conditions reaching 8-15 cm in vigorous vines, and apical internodes progressively shortening as growth cessation approaches. This spatial pattern enables retrospective assessment of growth conditions throughout the season from a single dormant-season measurement session, valuable for correlating growth patterns with fruit quality outcomes. Excessive internode length indicates overly vigorous growth creating sparse canopies with poor light interception efficiency, while abnormally short internodes suggest growth limitation from water stress, nutrient deficiency, or viral infection. Cultivar-specific internode length patterns are pronounced, with compact varieties like Pinot Noir and Chardonnay producing shorter internodes (6-10 cm mid-shoot) than sprawling varieties like Carignan or Mourvèdre (10-18 cm). The relationship between internode length and node number determines total shoot length and thus canopy architecture, with implications for trellis design, pruning strategy, and canopy management intensity requirements.
Internode_Length_Quality provides a spatially distributed vigor record along the shoot axis, complementing the temporally explicit Shoot_Growth_Rate_Quality. Each internode's length integrates approximately 7-14 days of growth conditions depending on cultivar and vigor status, creating a permanent anatomical archive readable post-dormancy. This quality relates to canopy architecture through its influence on shoot length, node density, and thus leaf area distribution and light interception patterns.
Gliedlänge
Internodienabstand
Internodiumlänge
internode distance
internode elongation
internode extension
Schultz HR (1992) An empirical model for the simulation of leaf appearance and leaf area development of primary shoots of several grapevine (Vitis vinifera L.) canopy systems. Scientia Horticulturae 52:179-200 Carbonneau A (1976) Analyse de la croissance des feuilles du sarment de vigne: estimation de sa surface foliaire par échantillonnage. Connaissance de la Vigne et du Vin 10:141-159
Internode Length Quality
Riesling in Mosel shows internode maturation from June through September at BBCH 31-43, with basal internodes initiating lignification in early June completing by late July, middle internodes maturing July-August, and apical internodes beginning maturation late August potentially incomplete by dormancy, accumulating starch reserves reaching 18-22% dry weight in mature tissues under cool climate conditions of 15-22°C temperature range, adequate late moisture, and declining nitrogen availability typical of this moderate-vigor late-season cultivar requiring thorough maturation for winter survival
Internode maturation represents the developmental transition where stem segments complete growth and undergo structural modifications establishing permanent woody architecture through lignification, secondary wall deposition, and reserve accumulation, converting herbaceous growing tissues to rigid supportive structures that provide mechanical stability and serve as carbohydrate storage organs for subsequent seasons. This fundamental developmental process begins in basal internodes during active shoot growth and progresses acropetally following elongation completion, typically spanning 4-8 weeks per internode from maturation initiation to full lignification, with mature lignified basal regions coexisting with elongating middle zones and forming apical sections creating developmental gradient along shoot. Maturation initiation follows completion of elongation when cells achieve final dimensions and growth transitions from expansion to differentiation and secondary wall synthesis, with timing varying along shoot from early June for basal internodes to August for apical sections in temperate regions. Lignification represents central maturation process involving secondary wall deposition with lignin incorporation creating rigid woody structure, proceeding through increased cellulose and hemicellulose synthesis forming secondary wall framework, lignin monomer synthesis and polymerization creating hydrophobic matrix binding wall polysaccharides, and progressive wall thickening particularly in vascular tissues and structural support cells. Secondary xylem development accompanies maturation with vascular cambium initiating activity between primary xylem and phloem, producing additional xylem elements internally increasing water transport capacity and phloem elements externally enhancing assimilate translocation, establishing secondary growth pattern that continues in subsequent seasons in older permanent structures. Periderm formation begins in basal internodes replacing epidermis as protective surface, with cork cambium initiating in subepidermal layers producing cork cells externally providing protection against water loss and pathogen entry, and phelloderm cells internally contributing to cortex structure, creating characteristic brown bark appearance of mature stems. Metabolic changes during maturation include declining respiration from growth levels toward maintenance rates 2-4 fold above completely dormant tissue, shifting protein synthesis from growth-related proteins to storage and defense proteins accumulating in parenchyma cells, and beginning carbohydrate accumulation with starch synthesis converting excess photosynthate to storage form in cortex and pith parenchyma. Reserve accumulation intensifies during late maturation as shoot growth ceases and photosynthate allocation shifts from growth to storage, with starch concentrations in mature internodes increasing from negligible during active growth to 15-25% dry weight by dormancy entry, providing crucial carbon reserves for subsequent season bud burst and early growth. Mechanical properties change dramatically during maturation from flexible herbaceous tissue in elongating internodes with tensile strength 5-15 MPa to rigid woody structure in mature lignified internodes with strength 40-80 MPa, providing self-supporting architecture and resistance to wind loading and mechanical damage. Cell wall composition shifts during maturation from primary walls dominated by pectin and hemicellulose allowing expansion to mature walls with high cellulose content 40-50% providing tensile strength, lignin content 15-25% providing compressive strength and hydrophobicity, and modified pectin providing cross-linking and matrix properties. Vascular tissue maturation includes secondary wall thickening and lignification in xylem vessels creating functional water transport elements persisting into subsequent seasons, sieve element and companion cell differentiation in phloem establishing long-distance assimilate transport capacity, and fiber development in vascular bundles providing mechanical support. Water content declines during maturation from 75-85% fresh weight in expanding tissues to 55-65% in mature lignified internodes, reflecting reduced metabolic activity, increased wall material deposition, and beginning preparation for winter dormancy in perennial structures. Photosynthetic activity in cortex chloroplasts declines during maturation as lignification and periderm formation reduce light penetration and gas exchange capacity, with stem photosynthesis contribution to carbon balance declining from modest levels during active growth to negligible in fully mature internodes. Hormonal regulation of maturation involves declining auxin and gibberellin from reduced apical production and transport affecting growth promotion, increasing abscisic acid promoting maturation processes and stress tolerance, ethylene potentially involved in lignification and senescence processes, and jasmonates possibly affecting secondary metabolism and defense compound synthesis. Temperature affects maturation rate with warm conditions promoting rapid lignification progression while cool conditions slow processes, though maturation proceeds across wide temperature range unlike growth which shows narrower optimal conditions. Light intensity affects maturation through effects on photosynthetic carbon availability for reserve accumulation and possible direct effects on lignification through photomorphogenic responses, with high light supporting rapid thorough maturation while shade may result in incomplete lignification creating weaker shoot structure. Water availability affects maturation timing and completeness with moderate water stress potentially promoting earlier maturation initiation and enhanced reserve accumulation, while severe stress may cause premature incomplete maturation affecting tissue quality and reserve storage. Nutrient availability particularly nitrogen affects maturation with adequate but not excessive nitrogen supporting normal maturation and good reserve accumulation, while nitrogen excess may delay maturation prolonging herbaceous state and reducing lignification, and deficiency may cause premature maturation limiting final shoot development and reserve accumulation. The process shows characteristic progression along shoot from base to apex with basal internodes completing maturation by mid-summer, middle internodes maturing during late summer, and apical internodes potentially incomplete maturation by growth cessation particularly in vigorous shoots or late-season growth. Cultivar variation appears in maturation timing and completeness with some varieties showing early thorough maturation creating well-lignified shoots while others maintain herbaceous state longer potentially creating winter hardiness concerns in cold regions. Regional timing shows maturation beginning June-July in warm climates, July-August in temperate regions, and August-September in cool areas for basal internodes, with progression continuing through late summer and early autumn approaching dormancy entry. Viticultural management considerations include ensuring adequate but not excessive late-season growth allowing proper maturation before dormancy, managing water and nitrogen avoiding excessive late vegetative growth interfering with maturation, assessing maturation adequacy for winter survival particularly in cold regions where incomplete lignification creates freeze damage risk, and recognizing that good maturation with substantial reserve accumulation benefits subsequent season bud break and early growth. Incomplete maturation from excessive late growth or inadequate resources creates winter injury risk and reduces reserve availability for next season. Research applications include analyzing mechanisms regulating lignification and secondary wall formation, quantifying reserve accumulation patterns and their regulation by source-sink dynamics, evaluating environmental and management effects on maturation affecting shoot quality and perennial structure development, assessing relationships between maturation and cold hardiness in grapevine adaptation to cold climates, and documenting transition from annual herbaceous shoot to perennial woody structure in woody plant development.
Internode_Maturation is fourth and final Internode_Development subphase, distinguished by lignification and reserve accumulation transforming herbaceous growth to woody permanent structure. Follows Internode_Elongation completion. Progresses acropetally from base to apex during late season. Critical for winter survival and subsequent season resource availability. Incomplete maturation creates hardiness and vigor concerns.
Holzbildung
Internodienreifung
Stängelverholzung
internode ripening
stem lignification
wood formation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 248-265 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 165-178 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Internode Maturation
SO4 (Selection Oppenheim 4, V. berlandieri × V. riparia) bears an Interspecific_Hybrid_Role as the quintessential rootstock hybrid, combining berlandieri's lime tolerance with riparia's phylloxera resistance and easy rooting to create the world's most widely planted rootstock. Regent serves in an Interspecific_Hybrid_Role as a complex hybrid carrying resistance genes from multiple American Vitis species through its Chambourcin parent, while expressing wine quality from V. vinifera ancestors. Muscadinia rotundifolia × V. vinifera crosses bear Interspecific_Hybrid_Roles (or more precisely intergeneric, as Muscadinia is sometimes separated from Vitis) in breeding programs targeting powdery mildew immunity from Run1 and Ren1 loci.
A hybrid role that inheres in a grapevine genotype originating from crosses between different species within the Vitis genus (or between Vitis and Muscadinia), combining genetic material from taxonomically distinct species to capture traits absent from any single species. Interspecific hybrids represent the foundation of modern viticulture infrastructure—phylloxera-resistant rootstocks are almost universally interspecific hybrids combining American species resistance with vigor and soil adaptation characteristics, while disease-resistant scion varieties (PIWI) introgress resistance genes from wild species into V. vinifera backgrounds. The interspecific crossing barrier varies with species combination Vitis species cross readily within the genus, while Vitis × Muscadinia crosses face chromosome number incompatibility (2n=38 versus 2n=40) requiring specialized techniques. Interspecific hybrids may exhibit intermediate characteristics (blending parental traits), transgressive segregation (exceeding both parents for some traits), or dominant expression (one parent's traits prevailing). The primary motivation for interspecific hybridization is trait introgression—accessing genes for disease resistance, pest tolerance, or environmental adaptation absent from V. vinifera, while breeding programs work to minimize undesirable interspecific contributions (off-flavors, excessive vigor, atypical phenolics).
Interspecific_Hybrid_Role encompasses the crosses between Vitis species that provide most biotic and abiotic stress tolerance in cultivated grapevines. V. vinifera, despite producing nearly all fine wine, lacks resistance to phylloxera, powdery mildew, downy mildew, and Pierce's disease—pathogens that evolved with American species in geographic isolation from V. vinifera's Central Asian/Mediterranean homeland. Interspecific hybridization accesses resistance mechanisms developed through millions of years of plant-pathogen coevolution, but also introduces traits poorly suited to wine production (foxy flavors from V. labrusca, excessive vigor, unfamiliar phenolic profiles). Contemporary PIWI breeding represents generations of work minimizing undesirable interspecific contributions while retaining resistance, with modern varieties approaching V. vinifera wine quality while carrying effective R-genes from wild species ancestors.
Artenhybride
Arthybride
Interspezies-Kreuzung
Weitkreuzung
cross-species hybrid
interspecific hybrid
species hybrid
wide cross
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Hausmann L et al. (2019) Grapevine breeding for disease resistance, in Cantu D, Walker MA (eds) The Grape Genome, Springer pp.: 311-330 This P et al. (2006) Historical origins of wine grapes, Trends in Genetics 22: 511-519 Riaz S et al. (2019) Genetic resources for grape improvement, in Cantu D, Walker MA (eds) The Grape Genome, Springer pp.: 25-51
Interspecific Hybrid Role
Interspecific hybridization capacity varies widely across Vitis species combinations, with V. vinifera showing high crossability with V. rupestris, V. riparia, and V. berlandieri (70-95% seed set, 60-85% germination) enabling rootstock breeding, while crosses with V. rotundifolia (Muscadinia subgenus) show severe barriers (<5% viable seed). First-generation (F1) hybrids between V. vinifera and American species typically show intermediate traits for disease resistance and wine quality, with subsequent backcrossing required to recover V. vinifera quality while retaining resistance. Modern PIWI (Pilzwiderstandsfähige) cultivars such as Regent, Solaris, and Cabernet Blanc derive from multiple generations of interspecific hybridization and backcrossing, combining Rpv and Ren resistance loci with acceptable wine quality.
A compatibility disposition that is the inherited capacity of a Vitis vinifera plant to successfully cross with other Vitis species to produce viable, fertile offspring, determined by reproductive barrier strength, genomic compatibility, and the developmental viability of hybrid embryos and resulting plants.
Interspecific_Hybridization_Disposition serves as the parent class for cross-compatibility and hybrid vigor traits. Hybridization has been fundamental to viticulture since the phylloxera crisis, providing both rootstock solutions and disease-resistant scion cultivars. The genus Vitis shows generally low reproductive barriers within the Vitis subgenus, facilitating breeding programs. Embryo rescue techniques enable recovery of hybrids from difficult crosses with low natural seed viability. Understanding hybridization barriers informs breeding strategy and genetic resource utilization.
Artkreuzungsfähigkeit
cross-breeding compatibility
interspecies crossing ability
interspezifische Hybridisierung
species hybridization capacity
Töpfer R et al. (2011) New horizons for grapevine breeding. Fruit, Vegetable and Cereal Science and Biotechnology 5:79-100 This P et al. (2006) Historical origins and genetic diversity of wine grapes. Trends in Genetics 22:511-519
Interspecific Hybridization Disposition
Interveinal chlorosis quality assessment for magnesium deficiency in Rheinhessen Silvaner employed systematic visual evaluation and tissue analysis to differentiate Mg limitation from other causes of interveinal yellowing and guide targeted nutritional management. Visual scoring of interveinal chlorosis quality used a 0-5 scale based on color contrast between veins and interveinal tissue, with scores ranging from 0 (uniform green, no contrast) through 2 (light green interveinal, distinctly green veins) to 5 (bright yellow interveinal with potential necrosis, sharply contrasting green veins). The interveinal chlorosis quality distribution showed characteristic basipetal pattern with basal leaves at mean score 3.8, mid-canopy at 2.2, and apical leaves at 0.5, confirming the mobile nutrient remobilization pattern diagnostic for magnesium rather than immobile nutrients. Color measurement using a portable spectrophotometer quantified the interveinal chlorosis quality, with chlorophyll concentration in interveinal tissue averaging 1.2 mg/g fresh weight in affected areas versus 2.8 mg/g in adjacent vein tissue, a 57% reduction creating the visible contrast. The interveinal chlorosis quality assessment distinguished magnesium from iron deficiency, which would show the opposite distribution pattern with youngest leaves most affected, enabling confident diagnosis without laboratory testing though tissue analysis confirmed petiole Mg at 0.22% (below the 0.30% threshold). Corrective foliar magnesium application showed visible improvement in interveinal chlorosis quality within 2 weeks, with new growth emerging with normal green coloration confirming the nutritional diagnosis.
Interveinal chlorosis quality in magnesium deficiency is a magnesium deficiency symptom quality that inheres in grapevine leaves and represents the characteristic pattern of yellowing between leaf veins while veins remain green, resulting from magnesium's essential role as the central atom in chlorophyll molecules such that magnesium deficiency directly limits chlorophyll synthesis in interveinal mesophyll tissue while vascular tissue maintains function through preferential magnesium allocation, producing the distinctive green-vein yellow-interveinal pattern that is pathognomonic for magnesium deficiency when the basipetal symptom progression is also observed. This quality captures the most visually diagnostic magnesium deficiency symptom, recognizing that the interveinal chlorosis pattern provides reliable field identification when properly distinguished from similar patterns caused by iron or manganese deficiency that affect youngest rather than oldest leaves. The ontological significance of interveinal chlorosis quality in magnesium deficiency lies in its diagnostic specificity when pattern characteristics are properly recognized, with the combination of interveinal distribution and basipetal (old-to-young) progression distinguishing magnesium from iron or manganese deficiency that shows acropetal progression, enabling targeted management response. The biological basis of magnesium-deficiency interveinal chlorosis encompasses the structural requirement for magnesium in chlorophyll where each molecule contains one coordinated magnesium atom, the preferential maintenance of vascular tissue function through magnesium allocation to phloem and xylem parenchyma, and the resulting chlorophyll loss in interveinal mesophyll as magnesium is remobilized to younger tissues and to vascular tissue maintenance. The visual appearance shows characteristic progression from pale green interveinal areas in early deficiency through yellow-green to bright yellow in moderate deficiency, with the color contrast against green veins becoming increasingly dramatic as deficiency intensifies, and in severe cases progression to interveinal necrosis as the chlorotic tissue dies following prolonged magnesium depletion.
Interveinal_Chlorosis_Quality_(Mg) is_a Magnesium_Deficiency_Symptom_Quality, representing the characteristic green-vein yellow-interveinal pattern from Mg-limited chlorophyll synthesis. This class addresses the most diagnostically distinctive Mg deficiency symptom. The basipetal progression (old leaves first) distinguishes from Fe/Mn chlorosis (young leaves first). Color contrast between veins and interveinal tissue provides visual diagnostic. Chlorophyll reduction quantifiable by spectrophotometry or SPAD. Progression from chlorosis to necrosis indicates severe prolonged deficiency.
Grünadern-Chlorose
Interkostale Chlorose bei Mg-Mangel
Magnesium-Interkostalvergilbung
Mg-Limitierungs-Chlorose
Mg-deficiency chlorosis
Mg-limitation chlorosis
green-vein chlorosis
magnesium interveinal yellowing
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication Mengel K, Kirkby EA (2001) Principles of Plant Nutrition, 5th Edition. Kluwer Academic Publishers
Interveinal Chlorosis Quality (Mg)
Interveinal chlorosis quality assessment for manganese deficiency in Burgundy Pinot Noir on limestone soils documented the characteristic symptom features enabling differentiation from concurrent iron chlorosis on this high-pH terroir. Visual examination revealed interveinal chlorosis quality with the distinctive greenish-yellow color in interveinal tissue that retained more green tones than the bright yellow to cream color of iron-deficient leaves on adjacent vines, providing the first diagnostic distinction between the two micronutrient limitations. Pattern assessment showed interveinal chlorosis quality with less sharp demarcation between chlorotic and green tissue compared to iron deficiency, with the color transition across the vein-interveinal boundary more gradual rather than the sharp contrast typical of Fe chlorosis. Severity scoring of interveinal chlorosis quality used a 0-4 scale, with scores of 2-3 predominating showing moderate chlorosis where chlorophyll meter readings averaged 22 SPAD units in interveinal zones versus 32 in vein-adjacent tissue, a 31% reduction compared to the 60-70% reduction typical of equivalent-scored iron chlorosis. Progressive examination of interveinal chlorosis quality revealed the characteristic necrotic spotting with small tan flecks 1-3 mm diameter distributed within chlorotic tissue on score 3+ leaves, a feature diagnostic for manganese deficiency not seen in iron or magnesium chlorosis. The interveinal chlorosis quality assessment guided targeted corrective application with foliar MnSO₄ at 0.25% for manganese-symptomatic vines and Fe-EDDHA for iron-symptomatic vines, with differential response confirming the diagnostic distinction():) Mn-treated vines showed chlorosis resolution within 2 weeks while Fe-treated vines with Mn deficiency showed no improvement, validating the interveinal chlorosis quality-based diagnosis.
Interveinal chlorosis quality in manganese deficiency is a manganese deficiency symptom quality that inheres in young grapevine leaves and represents the characteristic pattern of pale green to greenish-yellow discoloration in tissue between leaf veins while veins retain darker green coloration, resulting from manganese-limited chlorophyll synthesis and photosynthetic function, with the chlorosis pattern distinguished from iron deficiency by its more diffuse, less sharply contrasted appearance, its greenish-yellow rather than bright yellow color, and the frequent development of small necrotic spots within chlorotic areas as deficiency progresses. This quality captures the primary diagnostic feature of manganese deficiency, recognizing that while the interveinal chlorosis pattern resembles iron deficiency, careful assessment of color quality, pattern definition, and associated features enables differentiation that is essential for appropriate corrective management. The ontological significance of interveinal chlorosis quality in manganese deficiency lies in its diagnostic utility when characteristic features are recognized, enabling field identification that guides tissue analysis confirmation and targeted foliar manganese application, while misdiagnosis as iron deficiency would result in ineffective treatment with iron products. The biological basis of manganese-deficiency interveinal chlorosis encompasses manganese's essential role in the oxygen-evolving complex of photosystem II where deficiency impairs the light reactions of photosynthesis, the function of manganese in activation of enzymes involved in chlorophyll biosynthesis, and the consequent reduction in chlorophyll accumulation in mesophyll tissue while vascular tissue maintains preferential manganese allocation. The color quality of manganese-deficiency chlorosis reflects the partial impairment of photosynthesis distinct from the near-complete chlorophyll loss in severe iron deficiency, with affected tissue showing reduced but not eliminated chlorophyll producing the characteristic pale green to greenish-yellow rather than bright yellow appearance. The necrotic spotting that develops within chlorotic tissue in progressing manganese deficiency results from localized oxidative damage when manganese-dependent antioxidant enzymes (particularly superoxide dismutase) are impaired, allowing reactive oxygen species accumulation that damages cellular membranes and causes the characteristic small tan spots.
Interveinal_Chlorosis_Quality_(Mn) is_a Manganese_Deficiency_Symptom_Quality, representing the characteristic greenish-yellow interveinal chlorosis pattern from Mn-limited photosynthesis and chlorophyll synthesis. This class addresses the specific chlorosis symptom differentiating Mn from Fe and Mg deficiency. The greenish-yellow color (vs. bright yellow of Fe) is key diagnostic feature. Diffuse pattern with gradual vein-interveinal transition (vs. sharp contrast in Fe). Necrotic flecks within chlorotic tissue are Mn-specific. SPAD reduction ~30% (vs. 60-70% in equivalent Fe chlorosis). Responds to foliar Mn but not Fe application, confirming diagnosis.
Interkostale Chlorose bei Mn-Mangel
Mangan-Interkostalvergilbung
Mn chlorosis pattern
Mn-Mangel-Chlorose
Mn-deficiency chlorosis
manganese interveinal yellowing
manganese-induced interveinal chlorosis
manganinduzierte Interkostalchlorose
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication Millaleo R et al. (2010) Manganese as essential and toxic element for plants: transport, accumulation and resistance mechanisms. Journal of Soil Science and Plant Nutrition 10:470-481
Interveinal Chlorosis Quality (Mn)
Interveinal yellowing quality assessment for iron deficiency in Rioja Tempranillo on calcareous soils documented the characteristic symptom progression and distribution enabling confident field diagnosis. Visual examination revealed interveinal yellowing quality with the classic pattern of bright yellow tissue between veins contrasting sharply with distinctly green primary and secondary veins, creating the "netted" or "reticulate" appearance diagnostic for iron chlorosis. The color quality of interveinal yellowing showed characteristic progression from light green (early, score 1) through yellow-green (moderate, score 2-3) to bright yellow approaching cream (severe, score 4) to nearly white (very severe, score 5), with the progression from yellow toward white distinguishing iron from manganese deficiency where interveinal tissue typically remains more greenish-yellow. The spatial distribution of interveinal yellowing quality within the canopy showed strong acropetal pattern with symptom scores averaging 4.2 on the youngest fully expanded leaf, 2.8 at the fifth leaf position, and 0.5 at the tenth position, with basal leaves symptom-free, confirming the immobile nutrient distribution that distinguishes iron from magnesium deficiency. Fine vein assessment revealed that in moderate interveinal yellowing quality (score 2-3), primary and secondary veins remained green while tertiary veins showed yellowing, whereas severe interveinal yellowing quality (score 4-5) showed only primary veins retaining green color as iron limitation intensified. Quantitative chlorophyll analysis correlated with visual interveinal yellowing quality scores, with SPAD readings of 35+ in green vein areas versus 8-15 in interveinal regions of score 3-4 leaves, a 60-75% chlorophyll reduction creating the visible contrast.
Interveinal yellowing quality in iron deficiency is an iron deficiency symptom quality that inheres in young grapevine leaves and represents the characteristic pattern of chlorosis where tissue between leaf veins becomes progressively yellow to white while veins initially retain green coloration, with the interveinal distribution reflecting the differential iron allocation between vascular and mesophyll tissues and the dependence of chlorophyll synthesis on adequate iron supply, producing a visually striking pattern that is diagnostically valuable when the acropetal symptom distribution on youngest leaves is confirmed. This quality captures the primary visual diagnostic feature of iron deficiency, recognizing that interveinal chlorosis provides reliable field identification when the immobile-nutrient pattern of young-leaf symptom expression distinguishes iron from magnesium deficiency that produces similar interveinal patterns on old leaves. The ontological significance of interveinal yellowing quality in iron deficiency lies in its diagnostic utility enabling field identification of iron limitation, while also requiring careful differential diagnosis from other causes of interveinal chlorosis including magnesium, manganese, and zinc deficiencies that each show characteristic differences in symptom distribution, color quality, and associated features. The biological basis of iron-deficiency interveinal yellowing encompasses the requirement for iron in the biosynthesis of chlorophyll precursors, the preferential maintenance of iron status in vascular tissue at the expense of mesophyll, and the resulting loss of chlorophyll in interveinal regions while vein-adjacent tissue retains chlorophyll production capacity. The progression of interveinal yellowing severity provides information about deficiency intensity, advancing from pale green interveinal areas with distinct green veins in mild deficiency through yellow-green interveinal tissue in moderate deficiency to bright yellow or cream-colored tissue approaching white in severe deficiency, with eventual loss of green vein color as deficiency progresses and ultimately necrosis of bleached tissue if iron limitation persists.
Interveinal_Yellowing_Quality_(Fe) is_a Iron_Deficiency_Symptom_Quality, representing the characteristic green-vein yellow-interveinal pattern from Fe-limited chlorophyll synthesis on young leaves. This class addresses the primary visual diagnostic feature of Fe deficiency. The acropetal distribution (young leaves most affected) distinguishes from Mg chlorosis (old leaves first). Yellow-to-white color progression distinguishes from Mn chlorosis (more greenish-yellow). Vein hierarchy affected progressively (tertiary first, primary last). SPAD contrast between veins and interveinal tissue quantifies severity.
Eisen-Interkostalchlorose
Fe chlorosis pattern
Grünadern-Vergilbung durch Fe-Mangel
Interkostale Vergilbung bei Fe-Mangel
eiseninduzierte Interkostalchlorose
green-vein yellowing from Fe
iron interveinal chlorosis
iron-induced interveinal chlorosis
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication Römheld V (2000) The chlorosis paradox: Fe inactivation as a secondary event in chlorotic leaves of grapevine. Journal of Plant Nutrition 23:1629-1643
Interveinal Yellowing Quality (Fe)
Intra-plant synchrony assessment in cordon-trained Cabernet Sauvignon reveals the positional effects on within-vine uniformity, with eastern cordon arm (morning sun exposure, moderate vigor) showing high synchrony (cluster-to-cluster véraison variation CV 4.2%, berry-to-berry variation within clusters CV 6.8%), while western cordon arm (afternoon sun exposure, higher vigor from irrigation line proximity) shows lower synchrony (cluster CV 9.5%, berry CV 14.3%), with the asymmetry creating within-vine heterogeneity requiring either selective harvest by arm or acceptance of mixed maturity, demonstrating how positional factors within single vines affect synchronization quality.
Intra-plant synchrony quality is a synchronization quality that represents the temporal coordination of developmental events among different organs within a single grapevine, characterizing the uniformity of phenological progression across different shoot positions, different clusters, and different berries that determines whether the vine functions as a coherent developmental unit or as a collection of independently developing organs requiring position-specific management consideration. Within the BFO framework, this quality specifically addresses the within-vine dimension of synchronization, distinguishing coordination among organs of a single vine from coordination among different vines while recognizing that intra-plant synchrony determines fruit homogeneity at the individual vine level and affects the efficiency of per-vine operations like hand harvest and canopy management. The ontological significance lies in intra-plant synchrony determining the uniformity of fruit produced by individual vines, where high synchrony produces clusters and berries at consistent developmental stages enabling harvest of uniform fruit from each vine, while low synchrony produces mixed populations with berries at different ripeness levels that may require within-vine selection or acceptance of maturity heterogeneity in the harvest. The positional factors affecting intra-plant synchrony include shoot origin position (basal versus distal buds potentially differing in phenology), shoot vigor (more vigorous shoots sometimes advancing or delaying relative to moderate shoots), cluster position (basal versus apical clusters potentially differing in development rate), and fruit exposure (sun-exposed versus shaded clusters potentially developing differently), with these positional effects creating the within-vine variation that synchrony measures capture. The management factors affecting intra-plant synchrony include pruning uniformity (consistent bud numbers and positions promoting synchronous development), canopy management (leaf removal and positioning affecting fruit zone uniformity), and irrigation/nutrition uniformity (consistent resource supply promoting coordinated development), with these management practices either promoting or disrupting within-vine coordination depending on their uniformity. Understanding intra-plant synchrony quality enables assessment of within-vine developmental uniformity, identification of vine positions or practices contributing to asynchrony, and appropriate expectation setting for fruit homogeneity achievable from individual vines.
Intra_Plant_Synchrony_Quality serves as the parent class for event-specific within-vine synchrony (Bud Break, Flowering, Véraison, Ripening). This quality addresses coordination among organs within individual vines affecting fruit homogeneity. The class supports within-vine uniformity assessment through synchrony characterization across shoot and cluster positions.
Intra-Pflanzensynchronie
Organkoordination
organ coordination
positional synchrony
positionelle Synchronie
rebstockinterne Uniformität
vine-level uniformity
within-vine synchrony
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 59-144 Friend AP, Trought MCT (2007) Delayed winter spur-pruning in New Zealand can alter yield components of Merlot grapevines. Australian Journal of Grape and Wine Research 13(3): 157-164
Intra Plant Synchrony Quality
Müller-Thurgau (Riesling × Madeleine Royale, both V. vinifera) bears an Intraspecific_Hybrid_Role as one of the most successful deliberate V. vinifera crosses, bred in 1882 and becoming a major commercial variety particularly in Germany, despite susceptibility to the diseases that interspecific hybrids were bred to resist. Pinotage (Pinot Noir × Cinsaut, both V. vinifera) serves in an Intraspecific_Hybrid_Role as South Africa's signature grape, created in 1925 by Abraham Perold to combine Pinot quality with Cinsaut productivity under Cape conditions. Cabernet Sauvignon bears an Intraspecific_Hybrid_Role as a spontaneous V. vinifera hybrid (Cabernet Franc × Sauvignon Blanc), its hybrid origin unrecognized for centuries until genetic analysis in the 1990s revealed its parentage.
A hybrid role that inheres in a grapevine genotype originating from crosses between different varieties, clones, or populations within the same species, most commonly Vitis vinifera crosses creating new varieties combining characteristics of distinct V. vinifera parents. Intraspecific hybrids represent the traditional approach to variety improvement within V. vinifera, creating new wine grape varieties through controlled crosses between established cultivars without introducing genetic material from other species. Many important V. vinifera varieties originated as spontaneous intraspecific hybrids (natural crosses between cultivated varieties), subsequently identified and perpetuated through clonal selection—Cabernet Sauvignon (Cabernet Franc × Sauvignon Blanc), Chardonnay (Pinot × Gouais Blanc), and numerous others have spontaneous hybrid origins revealed through genetic analysis. Deliberate intraspecific breeding within V. vinifera aims to combine favorable characteristics (productivity, disease tolerance, quality attributes) while maintaining the V. vinifera identity valued for regulatory and market reasons. Intraspecific V. vinifera crosses typically produce progeny with wine quality similar to parents (avoiding the hybrid character concerns of interspecific crossing), but lack access to the disease resistance genes found only in wild species.
Intraspecific_Hybrid_Role captures crosses within V. vinifera that create new varieties without introducing wild species genetics. The distinction from interspecific hybridization is significant for both genetics (no exotic species contributions, no R-genes from wild relatives) and regulation (V. vinifera intraspecific hybrids typically qualify for quality wine designations restricted from interspecific hybrids). Many traditional V. vinifera varieties are themselves intraspecific hybrids of unknown or recently discovered parentage—genetic analysis has revealed extensive spontaneous hybridization in European viticulture history. Deliberate V. vinifera breeding programs exist but are limited compared to interspecific resistance breeding, as the primary value of crossing (introducing disease resistance) requires interspecific genetic resources. Intraspecific breeding within V. vinifera may target productivity, climate adaptation, or quality trait combinations achievable without wild species contributions.
Intraspezies-Hybride
Sortenkreuzung
V. vinifera hybrid
Vinifera-Hybride
innerartliche Hybride
intraspecific hybrid
varietal cross
within-species hybrid
This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Bowers JE, Meredith CP (1997) The parentage of a classic wine grape, Cabernet Sauvignon, Nature Genetics 16: 84-87 Lacombe T et al. (2013) Origins of grape cultivars, PLOS ONE 8: e62998 Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22
Intraspecific Hybrid Role
Intrinsic water use efficiency assessment comparing grapevine cultivars under identical conditions reveals genotypic variation in carbon-water coupling, with Tempranillo showing A/gs of 85 μmol/mol (high photosynthetic capacity efficiently utilizing stomatal opening), Grenache intermediate at 68 μmol/mol (moderate capacity, conservative stomatal behavior), and Syrah lower at 52 μmol/mol (maintaining open stomata that support high absolute photosynthesis but lower efficiency), demonstrating how intrinsic WUE captures cultivar-specific water use strategies independent of environmental humidity conditions.
Intrinsic water use efficiency quality is a transpiration efficiency quality that represents the ratio of net CO₂ assimilation rate to stomatal conductance (A/gs), characterizing the leaf-level efficiency of carbon gain relative to the stomatal opening that permits both CO₂ entry and water loss, independent of the vapor pressure deficit that determines actual transpiration rate. Within the BFO framework, this quality specifically addresses the stomatal-normalized dimension of water use efficiency, distinguishing the intrinsic relationship between assimilation and conductance from the atmospheric demand effects captured in instantaneous WUE while recognizing that A/gs reveals the inherent efficiency of the photosynthetic apparatus and stomatal behavior independent of environmental humidity conditions. The ontological significance lies in intrinsic WUE providing a measure of leaf-level efficiency that is directly comparable across different environmental conditions, where A/gs indicates how effectively the vine extracts carbon per unit stomatal opening, enabling comparison of cultivars or treatments without the confounding effect of vapor pressure deficit that varies with location, season, and time of day. The physiological basis for intrinsic WUE involves the relationship between CO₂ diffusion into leaves (proportional to gs and the CO₂ gradient) and water vapor diffusion out (proportional to gs and the water vapor gradient), with A/gs reflecting the efficiency of CO₂ fixation relative to the conductance that simultaneously permits water loss, typically ranging from 40-100 μmol CO₂/mol H₂O in Vitis vinifera depending on photosynthetic capacity and stomatal behavior. The factors affecting intrinsic WUE include photosynthetic capacity (higher Amax increasing A/gs at equivalent conductance), mesophyll conductance (affecting the CO₂ gradient between substomatal cavity and chloroplast), stomatal behavior (the operating point on the A-gs relationship determining efficiency), and Rubisco kinetics (affecting carboxylation efficiency at given internal CO₂). The cultivar variation in intrinsic WUE is substantial, with some cultivars achieving higher carbon gain per unit stomatal conductance through combinations of high photosynthetic capacity, efficient mesophyll conductance, and/or favorable stomatal operating points, enabling selection for improved water productivity. Understanding intrinsic water use efficiency quality enables cultivar comparison independent of measurement conditions, diagnosis of the physiological basis for efficiency differences, and breeding target identification for improved water productivity.
Intrinsic_Water_Use_Efficiency_Quality addresses the stomatal-normalized efficiency measure A/gs that enables comparison across environments. This quality reveals inherent photosynthetic and stomatal efficiency independent of VPD. The class supports cultivar comparison and physiological diagnosis through intrinsic WUE characterization.
A/gs
WUEi
intrinsic WUE
intrinsische Wassernutzungseffizienz
stomatal efficiency
stomatäre Effizienz
Medrano H et al. (2015) From leaf to whole-plant water use efficiency (WUE) in complex canopies: limitations of leaf WUE as a selection target. The Crop Journal 3(3): 220-228 Flexas J et al. (2010) Mesophyll conductance to CO₂: current knowledge and future prospects. Plant, Cell & Environment 33(8): 845-851
Intrinsic Water Use Efficiency Quality
Detailed physiological characterization of Intrinsic_WUE_Disposition at the University of the Balearic Islands analyzed the components underlying A/gs variation across 18 cultivars using gas exchange, chlorophyll fluorescence, and carbon isotope analysis. Simultaneous measurement of A, gs, and Ci enabled calculation of intrinsic WUE (A/gs) and partitioning of photosynthetic limitations into stomatal, mesophyll, and biochemical components. Intrinsic_WUE_Disposition ranged from 52 μmol mol⁻¹ ('Syrah', lowest) to 98 μmol mol⁻¹ ('Grenache', highest) under moderate water stress (-1.0 MPa predawn water potential). Analysis of component limitations revealed distinct mechanisms underlying high A/gs. 'Grenache' achieved high Intrinsic_WUE_Disposition primarily through sensitive stomatal closure (gs = 85 mmol m⁻² s⁻¹) while maintaining moderate A (7.2 μmol m⁻² s⁻¹) through high mesophyll conductance (gm = 125 mmol m⁻² s⁻¹) compensating for reduced CO₂ supply. 'Tempranillo' showed similarly high A/gs (92 μmol mol⁻¹) but achieved through different mechanism, with moderate gs (120 mmol m⁻² s⁻¹) and exceptionally high gm (185 mmol m⁻² s⁻¹) enabling high A (11.0 μmol m⁻² s⁻¹). Carbon isotope analysis confirmed that Δ¹³C correlated strongly with measured A/gs (r = -0.85), with low-Δ¹³C cultivars showing consistently high Intrinsic_WUE_Disposition across measurement dates. These findings demonstrate that equivalent Intrinsic_WUE_Disposition can arise through distinct physiological mechanisms with different implications for productivity and stress tolerance.
An Intrinsic_WUE_Disposition is a Water_Use_Efficiency_Disposition that inheres in a Vitis vinifera plant and is realized through the ratio of net photosynthetic rate (A) to stomatal conductance (gs), representing the leaf-level efficiency of carbon fixation relative to stomatal aperture independent of atmospheric evaporative demand. This disposition represents the physiologically intrinsic component of water use efficiency, removing the confounding effect of vapor pressure deficit (VPD) that influences transpiration rate at any given stomatal aperture. Intrinsic WUE (A/gs, expressed in μmol CO₂ mol⁻¹ H₂O) reflects the coordination between photosynthetic capacity and stomatal regulation, with high values indicating that carbon assimilation is maintained effectively relative to stomatal opening. Genotypic variation in Intrinsic_WUE_Disposition arises from differences in mesophyll conductance (gm, the conductance for CO₂ diffusion from substomatal cavity to chloroplasts), Rubisco carboxylation efficiency, and the sensitivity of photosynthesis versus stomata to environmental factors. Cultivars with high intrinsic WUE achieve relatively high photosynthetic rates at lower stomatal conductance, either through enhanced mesophyll conductance enabling efficient CO₂ delivery to chloroplasts or through high carboxylation capacity enabling rapid CO₂ fixation at lower intercellular CO₂ concentrations. Carbon isotope discrimination (Δ¹³C) provides an integrated measure of intrinsic WUE over time, as ¹³C discrimination during photosynthesis is inversely related to the ratio of intercellular to ambient CO₂ concentration (Ci/Ca), which in turn reflects the balance between stomatal supply and photosynthetic demand for CO₂.
Intrinsic_WUE_Disposition is_a Water_Use_Efficiency_Disposition. Intrinsic WUE (A/gs) removes the effect of atmospheric demand (VPD) that influences transpiration, isolating the physiological efficiency of the leaf. High A/gs can result from stomatal limitation (low gs) and/or enhanced CO₂ utilization (high gm, high carboxylation). Δ¹³C provides integrated assessment of intrinsic WUE suitable for screening. Mesophyll conductance (gm) is an important but often overlooked determinant of intrinsic WUE. Cultivars may achieve similar A/gs through different mechanistic pathways.
A/gs ratio
A/gs-Verhältnis
Intrinsische-WUE-Disposition
Photosynthetische-Wassereffizienz
photosynthetic water use efficiency
stomatal-normalized carbon gain
Flexas J et al. (2010) Mesophyll conductance to CO2: current knowledge and future prospects, Plant Cell and Environment 33:1123-1146 Tomás M et al. (2014) Variability of mesophyll conductance in grapevine cultivars under water stress conditions in relation to leaf anatomy and water use efficiency, Australian Journal of Grape and Wine Research 20:272-280 Farquhar GD et al. (1989) Carbon isotope discrimination and photosynthesis, Annual Review of Plant Physiology and Plant Molecular Biology 40:503-537
Intrinsic WUE Disposition
Cabernet Sauvignon in the Ningxia Helan Mountain region of China bearing an introduced cultivar role was first planted in the 1980s from French nursery material, with current plantings of approximately 25,000 ha established over the past 30 years using predominantly ENTAV-INRA clones 15, 169, and 337, demonstrating ongoing adaptation challenges including winter desiccation requiring annual burial of cordons at -25°C minimum temperatures, and measurably lower clonal diversity compared to the cultivar's 300+ year presence in Bordeaux.
A regional production role that inheres in a grapevine cultivar that was deliberately imported into a viticultural region from an external source within a historically documented timeframe, lacking the centuries-long presence and local adaptation characteristic of traditional cultivars in that region.
Introduced_Cultivar_Role is the complement of Traditional_Cultivar_Role and captures the historically recent arrival of a cultivar in a region. The temporal boundary between "introduced" and "traditional" is inherently contextual and gradual rather than categorical: Cabernet Sauvignon in Australia (planted from the 1830s, nearly 200 years of presence) occupies a transitional position between introduced and traditional status, whereas the same cultivar in Thailand (planted from the 2000s) is unambiguously introduced. The role may apply to both international varieties of global commercial significance and obscure cultivars introduced for experimental or niche purposes.
eingeführte Sorten-Rolle
gebietsfremde Sorten-Rolle
imported cultivar role
importierte Rebsorten-Rolle
introduced variety role
non-native cultivar role
Anderson K and Aryal NR (2013) Which Winegrape Varieties are Grown Where?, University of Adelaide Press Robinson J et al. (2012) Wine Grapes, Allen Lane pp: 1-18 Li Demei (2019)Erta Alla Vita: Chinese Wine Renaissance, Ebury Press pp: 35-58
Introduced Cultivar Role
Introduction record for Riesling to the Clare Valley, South Australia documents the arrival of Silesian Lutheran settlers in 1838-1842 who brought Riesling cuttings from their homeland vineyards in the Barossa region (themselves derived from earlier importations via the James Busby collection of 1831), with initial plantings at Watervale and Polish Hill River by the Sobels, Knaggs, and other settler families, the material propagated from dormant cuttings transported on the emigrant ships, and successful establishment confirmed by the first commercial Riesling vintage in the late 1840s — an introduction mediated by religious persecution, colonial migration, and cultural attachment to homeland foodways. Introduction record for Grüner Veltliner to New Zealand documents a deliberate research-driven introduction — small quantities of certified virus-free cuttings imported from the Höhere Bundeslehranstalt für Wein- und Obstbau Klosterneuburg to the Marlborough region in 2003-2005 by Donn Luthie of Donn Luthie Wines, motivated by interest in expanding New Zealand's varietal portfolio beyond Sauvignon Blanc and Pinot Noir, with initial trial plantings of 0.5 hectares and first commercial vintage in 2008.
Introduction record is a spread history entity that documents the specific event or process by which a grape cultivar was first brought to a new viticultural region where it had not previously been cultivated, specifying the source region from which the material originated, the receiving region or site of introduction, the date or date range of the introduction, the agent or institution responsible (individual viticulturist, colonial administrator, religious order, government agency, research institution), the form of the introduced material (rooted vines, dormant cuttings, seeds, bench grafts), the quantity introduced, the motivation for the introduction (commercial, experimental, collection, accidental), and the immediate outcome (successful establishment, failure, repeated attempts), providing the event-level documentation of cultivar migration that constitutes the primary data for reconstructing spread histories.
Introduction record documentation captures the micro-historical events through which cultivar spread history is constructed — each global cultivar distribution pattern is the aggregate result of dozens or hundreds of individual introduction events, each with its own agents, motivations, circumstances, and outcomes. The documentation quality of introduction events varies enormously — some are meticulously recorded (James Busby's 1831 collection of 543 vine varieties from France and Spain for introduction to Australia is documented in his published collection catalogue and in the records of the Royal Botanic Gardens, Sydney), while others are reconstructed from indirect evidence (the introduction of Vitis vinifera to the Americas by Spanish missionaries is documented in general terms but specific cultivar identifications are often uncertain). The form of introduced material has historically significant implications — cuttings and rooted vines preserve clonal identity through vegetative propagation, while seeds (used in some early introductions) produce genetically novel individuals through sexual reproduction, meaning that seed-propagated introductions generated local seedling populations rather than true varietal transfers. The phytosanitary dimension of cultivar introduction has become increasingly stringent — modern introductions typically require quarantine periods (2-5 years in many jurisdictions), virus testing (ELISA, PCR, and biological indexing as documented in the Certification_Status hierarchy), and government import permits, contrasting with the essentially unregulated historical introductions that inadvertently spread devastating vine diseases and pests (phylloxera, powdery mildew, downy mildew, Pierce's disease) alongside the desired cultivars. The motivations for cultivar introduction have evolved historically — from subsistence and cultural continuity (colonial settlers reproducing homeland foodways) through commercial speculation (introducing proven commercial varieties to new regions) to scientific evaluation (systematic varietal trials at research institutions) and market diversification (introducing unfamiliar cultivars to differentiate regional wine portfolios), with each motivation producing different patterns of selection, establishment effort, and long-term success.
Einfuhraufzeichnung
Einführungsdokumentation
Einführungsnachweis
Sorteneinführung
Sortenimport
cultivar import
introduction event
planting introduction
varietal introduction
vine importation
Busby J (1833) Catalogue of Vines in the Botanic Garden at Sydney. Government Printer, Sydney Pinney T (1989) A History of Wine in America, University of California Press, pp.: 215-280 Halliday J (1994) A History of the Australian Wine Industry 1949-1994, Australian Wine and Brandy Corporation, pp.: 1-30 Unwin T (1991) Wine and the Vine, Routledge, pp.: 200-300
Introduction Record
Investigation of Ion_Compartmentalization_Disposition at the University of Adelaide employed X-ray microanalysis and tissue dissection to characterize subcellular and whole-plant ion distribution in 12 rootstocks following 28 days at 50 mM NaCl. Frozen-hydrated leaf sections were analyzed by energy-dispersive X-ray spectroscopy to determine vacuolar versus cytoplasmic ion concentrations, while systematic sampling of leaves by position quantified whole-plant ion distribution patterns. Ion_Compartmentalization_Disposition varied substantially among rootstocks. '1103 Paulsen' (which showed moderate exclusion capacity) demonstrated strong compartmentalization, with leaf mesophyll cells showing vacuolar Na⁺ concentration of 285 mM versus cytoplasmic concentration of only 28 mM (10-fold compartmentalization ratio), and vacuolar Cl⁻ of 195 mM versus cytoplasmic 32 mM. Whole-plant analysis revealed preferential accumulation in oldest leaves (leaf position 1-3 from base averaging 125 mmol Na⁺/kg DW) versus youngest fully expanded leaves (position 8-10 averaging 35 mmol/kg). 'Ramsey' showed similar compartmentalization patterns despite different absolute concentrations. In contrast, 'Schwarzmann' (poor excluder) showed weak Ion_Compartmentalization_Disposition with vacuolar/cytoplasmic Na⁺ ratio of only 3.5 and more uniform distribution across leaf positions (oldest 380, youngest 285 mmol/kg). The strong compartmentalization of '1103 Paulsen' enabled maintenance of cytoplasmic K⁺/Na⁺ ratio above 5 (critical for enzyme function) despite whole-leaf K⁺/Na⁺ ratio of only 2.5, demonstrating how compartmentalization protects metabolic function when exclusion is incomplete.
An Ion_Compartmentalization_Disposition is a Salt_Tolerance_Disposition that inheres in a Vitis plant and is realized through cellular and whole-plant mechanisms that sequester accumulated toxic ions (sodium and chloride) in locations where they cause minimal metabolic damage, enabling tolerance of tissue ion concentrations that would otherwise be injurious. This disposition represents a tissue tolerance strategy complementing ion exclusion, operating when exclusion mechanisms are overwhelmed or insufficient and ions accumulate in plant tissues. Effective compartmentalization protects metabolically sensitive compartments (particularly the cytoplasm where enzymes function and protein synthesis occurs) by directing ions to less sensitive locations including the vacuole (intracellular compartmentalization) and older, expendable tissues (whole-plant compartmentalization). Vacuolar sequestration removes ions from the cytoplasm while simultaneously contributing to osmotic adjustment, as vacuolar solutes lower cell water potential enabling continued water uptake from saline soil. Whole-plant compartmentalization preferentially accumulates ions in older leaves that are approaching senescence, sacrificing these tissues to protect young, photosynthetically active leaves critical for current and future productivity. Ion_Compartmentalization_Disposition varies among genotypes in both the capacity for vacuolar sequestration (determined by tonoplast transporter expression and activity) and the efficiency of directing ions to older tissues (involving coordination of xylem unloading and phloem redistribution), providing genetic variation for breeding salt-tolerant cultivars when exclusion capacity is limited.
Ion_Compartmentalization_Disposition is_a Salt_Tolerance_Disposition. Compartmentalization provides tissue tolerance when exclusion is insufficient, protecting sensitive metabolic compartments. Child classes include Vacuolar_Sequestration_Disposition (intracellular level) and Older_Tissue_Accumulation_Disposition (whole-plant level). Compartmentalization maintains cytoplasmic K⁺/Na⁺ ratio above critical threshold (~1-5) even when whole-tissue ratio is unfavorable. Compartmentalization and exclusion operate complementarily, with both contributing to overall salt tolerance.
Ionen-Sequestrierung
Ionenkompartimentierungs-Disposition
Toxische-Ionen-Kompartmentierung
ion sequestration
tissue ion tolerance
toxic ion compartmentation
Munns R, Tester M (2008) Mechanisms of salinity tolerance, Annual Review of Plant Biology 59:651-681 Flowers TJ, Colmer TD (2008) Salinity tolerance in halophytes, New Phytologist 179:945-963 Shabala S, Mackay A (2011) Ion transport in halophytes, Advances in Botanical Research 57:151-199
Ion Compartmentalization Disposition
Systematic characterization of Ion_Exclusion_Disposition at INRA Montpellier employed detailed ion profiling across the soil-root-shoot pathway in 15 rootstocks under controlled salinity (50 mM NaCl). Root, stem xylem sap, petiole, and leaf lamina concentrations of chloride, sodium, potassium, and calcium were measured to locate exclusion barriers and quantify exclusion efficiency. Ion_Exclusion_Disposition showed strong rootstock variation with distinct exclusion patterns. '140 Ruggeri' demonstrated exceptional exclusion at multiple sites, with root chloride concentration of 85 mM (versus 50 mM external), xylem sap chloride of only 2.5 mM (indicating strong xylem loading restriction), and leaf lamina chloride of 18 mmol/kg DW. The exclusion factor (external/leaf concentration ratio) was 48 for chloride. '1103 Paulsen' showed similarly strong chloride exclusion (exclusion factor 42) but weaker sodium exclusion (factor 15), resulting in moderate sodium accumulation (45 mmol/kg DW) despite low chloride. 'Riparia Gloire' exhibited poor Ion_Exclusion_Disposition with minimal restriction at any site, showing xylem sap chloride of 28 mM and leaf chloride of 380 mmol/kg DW (exclusion factor only 2.3). Correlation analysis confirmed that chloride exclusion was the dominant determinant of overall salt tolerance (r = 0.92 with growth maintenance), while sodium exclusion contributed independently but less strongly (r = 0.65). These findings demonstrate that Ion_Exclusion_Disposition involves coordinated restriction at multiple pathway sites, with chloride exclusion particularly critical for grapevine.
An Ion_Exclusion_Disposition is a Salt_Tolerance_Disposition that inheres in a Vitis plant and is realized through physiological mechanisms that prevent or limit the uptake of toxic ions (primarily chloride and sodium) from saline soil and their translocation to sensitive shoot tissues, maintaining low ion concentrations in leaves despite elevated external salinity. This disposition represents the primary mechanism of salt tolerance in grapevine, as the species lacks strong tissue tolerance to high internal ion concentrations and depends on exclusion to avoid toxicity. Ion exclusion operates at multiple sites in the soil-plant pathway including the root epidermis and cortex (initial uptake restriction), the endodermis (regulated loading into the xylem), the xylem parenchyma along the transpiration stream (retrieval of ions from xylem sap), and the leaf base (restriction of entry into leaf lamina). Effective Ion_Exclusion_Disposition maintains leaf chloride below approximately 25-50 mmol/kg DW and sodium below 25-35 mmol/kg DW, thresholds above which toxicity symptoms typically appear. Exclusion capacity varies substantially among rootstock genotypes, with excluders maintaining low shoot ion concentrations across a wide range of external salinity while non-excluders show proportional accumulation with external supply. Ion_Exclusion_Disposition is energetically costly, requiring active transport against concentration gradients, but this cost is far less than the productivity loss from ion toxicity, making exclusion the favored strategy for grapevine salt tolerance.
Ion_Exclusion_Disposition is_a Salt_Tolerance_Disposition. Ion exclusion is the primary salt tolerance mechanism in grapevine, as the species lacks strong tissue tolerance. Child classes address Chloride_Exclusion_Disposition and Sodium_Exclusion_Disposition, with chloride exclusion generally more important for grapevine. Exclusion operates at multiple sites including root uptake, xylem loading, and xylem retrieval. Exclusion factor (external/internal concentration ratio) quantifies exclusion efficiency. Effective excluders can maintain low leaf ion concentrations across wide external salinity ranges.
Ionenausschluss-Disposition
Salzausschluss
Toxische-Ionen-Vermeidung
ion restriction
salt exclusion
toxic ion avoidance
Munns R, Tester M (2008) Mechanisms of salinity tolerance, Annual Review of Plant Biology 59:651-681 Walker RR et al. (2004) Rootstock effects on salt tolerance of irrigated grapevines in the Sunraysia region, Australian Journal of Grape and Wine Research 10:34-45 Storey R et al. (2003) Transport of sodium from roots to shoots in grapevine rootstocks, Plant, Cell and Environment 26:1943-1956
Ion Exclusion Disposition
The IPM protocol for a Sustainable Winegrowing New Zealand (SWNZ) certified Sauvignon Blanc vineyard in Marlborough specifies a five-tier intervention hierarchy — Tier 1 cultural prevention (canopy management maintaining less than 2 leaf layers in the fruit zone, balanced nitrogen nutrition below 250 mg N/L in petiole analysis at flowering), Tier 2 biological control (Trichoderma atroviride applications to pruning wounds for Eutypa lata protection, mating disruption for leafroller moths), Tier 3 threshold-triggered soft chemistry (potassium bicarbonate for powdery mildew above 5% leaf area affected, Bacillus subtilis for botrytis above 2% cluster incidence), Tier 4 targeted synthetic fungicides (spiroxamine, metrafenone, or fluopyram applied only when soft chemistry proves insufficient, maximum 2 synthetic applications per season), and Tier 5 emergency intervention (full-rate synthetic program only in declared epidemic seasons with infection pressure exceeding 3 standard deviations above regional 10-year average).
A pest management description that documents the integrated pest management framework governing pest and disease control decisions in the vineyard, specifying the hierarchical combination of cultural, biological, physical, and chemical control methods applied according to monitoring-based intervention thresholds that minimize pesticide use while maintaining pest and disease levels below economically and qualitatively damaging levels.
IPM_Protocol documents the decision framework that distinguishes integrated pest management from both calendar-based spray programs (applying pesticides at fixed intervals regardless of actual pest pressure) and purely reactive crisis management (intervening only after damage is observed). The IPM concept, formalized in the FAO definition (2002) and implemented through national and regional certification schemes worldwide, is built on four principles — prevention through cultural practices, monitoring-based decision making, preferential use of non-chemical methods, and targeted chemical intervention as a last resort: in viticultural practice, IPM adoption varies enormously from token compliance (reducing spray numbers from 15 to 12 without fundamentally changing the approach) through genuine threshold-based management (as practiced in many New World regions with lower disease pressure) to advanced IPM incorporating predictive disease models, biological control agents, and precision application technologies. The EU Sustainable Use Directive (2009/128/EC) mandates IPM implementation for all professional pesticide users in the EU, making IPM documentation a regulatory requirement for European viticulture.
IPM framework documentation
IPM-Protokoll
integrated crop protection protocol
integrated pest management plan
integrierte Schädlingsbekämpfung-Dokumentation
integrierter Pflanzenschutz-Plan
Pertot I et al. (2017) A critical review of plant protection tools for reducing pesticide use on grapevine and new perspectives for the implementation of IPM in viticulture, Crop Protection 97: 70-84 EU Directive 2009/128/EC on sustainable use of pesticides FAO (2002) International Code of Conduct on the Distribution and Use of Pesticides Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests 2nd Ed, APS Press
IPM Protocol
IPMP concentration at harvest typically ranges from 0.1-5 ng/L across cultivars, representing 5-20% of total methoxypyrazine content in most genotypes. Cabernet Sauvignon shows IPMP levels of 1-4 ng/L, Sauvignon Blanc 0.5-3 ng/L, and Carménère 2-6 ng/L at harvest. The sensory threshold of IPMP (0.3-2 ng/L) is lower than IBMP, making its earthy, green pea character perceptible at trace concentrations. IPMP biosynthesis parallels IBMP, utilizing valine-derived isopropyl amine rather than leucine-derived isobutyl amine, with subsequent methylation by VvOMT3. The IPMP/IBMP ratio varies among cultivars (0.05-0.20 typical) and increases during ripening as IBMP declines faster than IPMP through preferential photodegradation. IPMP contributes distinctive earthy, asparagus notes distinguishable from the bell pepper character of IBMP in sensory analysis.
A methoxypyrazine accumulation disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate 3-isopropyl-2-methoxypyrazine (IPMP) as a minor methoxypyrazine in grape berries, with genotype-specific concentrations contributing green pea and earthy aroma notes that complement the bell pepper character of IBMP.
IPMP represents the second most abundant methoxypyrazine in grape berries, contributing complementary green character to the IBMP-dominated profile. The lower sensory threshold of IPMP relative to IBMP means small absolute concentrations can significantly impact wine aroma. The earthy/green pea character of IPMP is sometimes considered more undesirable than the bell pepper note of IBMP, influencing quality perception in affected wines. IPMP shows similar environmental modulation to IBMP, with cluster exposure reducing accumulation through photodegradation.
Erbsen-Aroma
IPMP synthesis capacity
IPMP-Akkumulation
Isopropyl-Methoxypyrazin
green pea aroma trait
isopropyl-methoxypyrazine disposition
Sala C et al. (2004) Contents of 3-alkyl-2-methoxypyrazines in musts and wines from Vitis vinifera variety Cabernet Sauvignon. Journal of Agricultural and Food Chemistry 52:5978-5983 Hashizume K, Samuta T (1999) Grape maturity and light exposure affect berry methoxypyrazine concentration. American Journal of Enology and Viticulture 50:194-198
IPMP Accumulation Disposition
IPMP content quality analysis in Carmenère from Chile's Colchagua Valley reveals this variety's distinctive methoxypyrazine profile with IPMP concentrations of 5-15 ng/L contributing earthy, green pea notes that complement the dominant IBMP bell pepper character at 25-40 ng/L the combined methoxypyrazine signature creates Carmenère's characteristic green-spectrum complexity that distinguishes it from Merlot (with which it was historically confused in Chilean vineyards), with IPMP's asparagus-earth notes providing varietal fingerprinting value beyond IBMP alone.
IPMP_Content_Quality_(Isopropyl-methoxypyrazine) represents the chemical quality characterizing the concentration of 3-isopropyl-2-methoxypyrazine in grape berries and wine, capturing the methoxypyrazine compound with the lowest sensory threshold that contributes earthy, asparagus, and pea-like green aromas distinct from the bell pepper character of IBMP. IPMP (C₈H₁₂N₂O) is biosynthesized from valine through an analogous pathway to IBMP, with the isopropyl side chain conferring different sensory properties and an even lower odor threshold (approximately 0.3-2 ng/L in water, 1-2 ng/L in wine). This extraordinarily low threshold means that IPMP can contribute to green character at concentrations barely detectable by standard analytical methods. In grape berries, IPMP typically occurs at lower concentrations than IBMP, usually 1-10 ng/L compared to 10-50 ng/L for IBMP at harvest, but its lower threshold means it can make significant sensory contribution despite the concentration difference. The earthy, asparagus-like character of IPMP differs from the green bell pepper note of IBMP, with IPMP contributing more vegetal, raw-vegetable quality at elevated levels. Like IBMP, IPMP accumulates early in berry development and declines during ripening through light-mediated degradation, providing viticultural management opportunities. The IPMP-to-IBMP ratio varies among cultivars and may provide fingerprinting information, with some varieties showing higher relative IPMP contribution. In wine production, IPMP follows the same behavior as IBMP—complete extraction, stable through fermentation and aging—making grape content directly predictive of wine concentration.
IPMP_Content_Quality addresses the isopropyl-substituted methoxypyrazine with the lowest sensory threshold among grape pyrazines. The parenthetical (Isopropyl-methoxypyrazine) follows the uploaded structure convention. IPMP analysis requires the same sensitive GC-MS/MS methods as IBMP, with even greater analytical challenges due to typically lower concentrations. The distinct sensory character of IPMP (earthy, asparagus) versus IBMP (bell pepper) means they contribute differently to overall green impression despite similar chemical class. SBMP (sec-butyl methoxypyrazine) represents a third grape methoxypyrazine but typically occurs at lower concentrations than IPMP.
3-Isopropyl-2-methoxypyrazin-Qualität
3-isopropyl-2-methoxypyrazine quality
IPMP content
IPMP-Gehalt
Isopropyl-Methoxypyrazin-Konzentration
earthy pyrazine concentration
erdiger Pyrazingehalt
isopropyl-methoxypyrazine level
Sala C et al. (2004) Headspace Solid-Phase Microextraction Method for Determining 3-Alkyl-2-methoxypyrazines in Musts by Means of Polydimethylsiloxane-Divinylbenzene Fibres. Journal of Chromatography A 1046(1-2): 1-6 Allen MS et al. (1994) Methoxypyrazines in Vitis vinifera Grapes and Wines. In: Trends in Flavour Research, Elsevier pp.: 365-370
IPMP Content Quality (Isopropyl-methoxypyrazine)
Iron acquisition efficiency varies 3-5 fold among rootstocks, with Fercal and 41B showing high efficiency (maintaining leaf Fe above 80 ppm on calcareous soils) compared to Riparia Gloire and 3309C (leaf Fe declining to 40-60 ppm). The Strategy I response involves coordinated expression of VvHA (H⁺-ATPase for acidification), VvFRO (ferric reductase), and VvIRT1 (iron-regulated transporter). Iron deficiency chlorosis develops when active leaf Fe falls below 50-60 ppm despite potentially adequate total Fe (the chlorosis paradox). Fe-efficient rootstocks reduce rhizosphere pH by 0.8-1.5 units compared to 0.2-0.4 units in Fe-inefficient types under bicarbonate stress.
A micronutrient uptake disposition that is the inherited capacity of a Vitis vinifera plant to acquire iron from the soil solution through Strategy I mechanisms (rhizosphere acidification, ferric reduction, ferrous uptake), with genotype-specific efficiency determining iron nutrition status across varying soil pH and carbonate conditions.
Iron acquisition efficiency is critical for vineyard performance on calcareous soils representing approximately 30% of global vineyard area. The trait encompasses both constitutive uptake capacity and inducible responses to Fe limitation. Unlike the Lime_Tolerance context focusing on chlorosis resistance, this disposition addresses the general Fe acquisition mechanism across soil types. Rootstock breeding has successfully combined high Fe efficiency with other desirable traits, with modern selections such as Fercal and 140 Ruggeri enabling viticulture on high-lime soils.
Eisenaneignungsvermögen
Eisenaufnahme-Effizienz
Fe acquisition capacity
Fe-Akquisitionseffizienz
iron foraging trait
iron uptake efficiency
Brancadoro L et al. (1995) Mechanisms of Fe-efficiency in roots of Vitis spp. in response to iron deficiency stress. Plant and Soil 171:229-234 Jiménez S et al. (2007) Phosphoenolpyruvate carboxylase (PEPC) and dark respiration in the Fe-deficient grapevine (Vitis vinifera L.) rootstock 140 Ruggeri. Journal of Plant Physiology 164:1527-1535
Iron Acquisition Efficiency Disposition
Investigation of Iron_Chelate_Uptake_Disposition at the University of Reims combined ⁵⁹Fe uptake kinetics with VvIRT1 gene expression analysis across 10 rootstocks under iron-sufficient and iron-deficient conditions. Short-term (30 min) ⁵⁹Fe²⁺ uptake was measured at Fe concentrations from 0.1 to 50 μM to determine Vmax and Km parameters, while qRT-PCR quantified VvIRT1 transcript abundance. Iron_Chelate_Uptake_Disposition showed significant genotypic variation, though less dramatic than reductase activity differences. Under Fe deficiency, '41B' exhibited high uptake capacity with Vmax of 52 nmol Fe g⁻¹ FW h⁻¹ and Km of 3.2 μM, accompanied by 35-fold induction of VvIRT1 expression relative to Fe-sufficient controls. 'Fercal' showed similar uptake parameters (Vmax 48, Km 2.8 μM) with 40-fold IRT1 induction. '3309C' displayed lower Iron_Chelate_Uptake_Disposition with Vmax of 28 nmol g⁻¹ h⁻¹ and higher Km of 5.5 μM (indicating lower affinity), with only 12-fold IRT1 induction. The uptake/reductase activity ratio (indicating coupling efficiency) varied from 0.28 in '41B' (efficient coupling) to 0.62 in '3309C' (less efficient, with more reductase activity than uptake capacity can utilize), suggesting that '3309C' loses a portion of generated Fe²⁺ to reoxidation before uptake. Importantly, differences in Iron_Chelate_Uptake_Disposition were smaller (2-fold range) than reductase differences (4-fold range), confirming that reductase activity is typically rate-limiting. However, under bicarbonate stress where Fe²⁺ reoxidation is accelerated, high uptake capacity and tight reductase-transporter coupling become more critical.
An Iron_Chelate_Uptake_Disposition is an Iron_Acquisition_Efficiency_Disposition that inheres in Vitis root epidermal cells and is realized through efficient uptake of ferrous iron (Fe²⁺) produced by ferric reductase activity, transporting iron across the plasma membrane into root cells via IRT1 (Iron-Regulated Transporter 1) and related transporters. This disposition represents the third component of Strategy I iron acquisition, completing the soil-to-cell pathway by importing the Fe²⁺ generated by reductase activity. Efficient uptake requires both adequate transporter expression (determined by iron regulatory networks) and functional coupling with reductase activity so that Fe²⁺ produced at the membrane surface is captured before reoxidation to Fe³⁺ in the alkaline rhizosphere. IRT1 belongs to the ZIP (Zinc/Iron-regulated transporter Protein) family and is highly induced under iron deficiency, with transcript and protein levels increasing 10-50 fold in deficient roots. Iron_Chelate_Uptake_Disposition varies among genotypes in both IRT1 expression levels and the efficiency of coupling between reduction and uptake. While uptake capacity rarely limits iron acquisition under laboratory conditions (reductase activity is typically rate-limiting), field performance may depend on uptake efficiency when reductase-generated Fe²⁺ is rapidly reoxidized in high-pH, high-bicarbonate soil environments. Additionally, IRT1 transports other divalent cations (Mn²⁺, Zn²⁺, Cd²⁺), and selectivity for iron versus competing metals may differ among genotypes.
Iron_Chelate_Uptake_Disposition is_a Iron_Acquisition_Efficiency_Disposition. IRT1 mediates Fe²⁺ uptake, completing the Strategy I pathway. Expression is highly iron-regulated (10-50 fold induction under deficiency). Uptake is typically not rate-limiting (reductase is), but coupling efficiency matters under field conditions where Fe²⁺ reoxidation competes with uptake. IRT1 also transports Mn²⁺, Zn²⁺, and potentially toxic Cd²⁺. VvIRT1 encodes the main iron transporter. Kinetic parameters (Vmax, Km) characterize uptake capacity and affinity.
Eisen-Chelat-Aufnahme-Disposition
Fe²⁺ uptake capacity
Fe²⁺-Transport-Kapazität
IRT1-mediated uptake
IRT1-vermittelte-Aufnahme
ferrous iron transport
Eide D et al. (1996) A novel iron-regulated metal transporter from plants identified by functional expression in yeast, Proceedings of the National Academy of Sciences 93:5624-5628 Vert G et al. (2002) IRT1, an Arabidopsis transporter essential for iron uptake from the soil and for plant growth, Plant Cell 14:1223-1233 Korshunova YO et al. (1999) The IRT1 protein from Arabidopsis thaliana is a metal transporter with a broad substrate range, Plant Molecular Biology 40:37-44
Iron Chelate Uptake Disposition
Iron content quality management in Cognac vineyards on Champagne (chalk) soils requires careful rootstock selection to prevent the lime-induced chlorosis that devastated pre-phylloxera vineyards, with Fercal and 41B rootstocks enabling Ugni Blanc cultivation on pH 8.0+ soils where iron availability is severely limited must iron content of 3-6 mg/L from properly managed vineyards supports healthy fermentation without the instability problems that arise from excessive iron, producing the clean, distillable base wines essential for quality Cognac production.
Iron_Content_Quality represents the chemical quality characterizing the concentration of iron (Fe) in grape berry tissue, capturing the essential micronutrient central to chlorophyll synthesis, electron transport, and numerous oxidation-reduction reactions in plant metabolism. Iron in grape juice typically ranges from 2-10 mg/L, with substantial variation reflecting cultivar, rootstock, soil iron availability, and the complex chemistry governing iron solubility and uptake. Despite being abundant in most soils, iron availability is severely limited in calcareous (high-pH) soils where ferric iron (Fe³⁺) forms insoluble hydroxides, creating the widespread problem of lime-induced chlorosis that affects susceptible rootstocks throughout the world's limestone-dominated wine regions. Iron deficiency produces characteristic interveinal chlorosis with yellow tissue between green veins, progressing to complete leaf yellowing and reduced photosynthetic capacity that limits yield and delays ripening. In berry tissue, iron participates in numerous enzyme systems and electron carriers including cytochromes and iron-sulfur proteins essential for respiration. In winemaking, iron affects wine stability through its participation in oxidation reactions and potential formation of insoluble iron phosphate or iron tannate precipitates that cause turbidity in white wines and color modification in red wines. The ferric-ferrous (Fe³⁺/Fe²⁺) redox couple participates in wine oxidation chemistry, with iron catalyzing reactions between oxygen and phenolic compounds. Excessive iron content (above 7-10 mg/L) increases oxidation risk and instability, while normal levels (2-5 mg/L) contribute to enzyme function during fermentation without causing problems. Iron analysis employs atomic absorption spectroscopy, ICP-OES, or colorimetric methods (ferrozine, phenanthroline) depending on required sensitivity.
Iron_Content_Quality addresses a micronutrient whose deficiency is extremely common in calcareous soils despite soil iron abundance. The pH-dependent solubility of iron creates the distinction between total soil iron and available iron that determines plant uptake. In wine, iron's redox activity creates both beneficial (enzyme function) and detrimental (oxidation catalysis, instability) effects depending on concentration. The interaction between iron, copper, and oxygen in wine oxidation chemistry is complex, with both metals participating in Fenton-type reactions generating reactive oxygen species.
Beereneisenqualität
Eisengehalt
Eisenkonzentration
Fe content
Fe-Gehalt
berry iron quality
iron concentration
iron level
Römheld V and Marschner H (1986) Mobilization of Iron in the Rhizosphere of Different Plant Species. Advances in Plant Nutrition 2: 155-204 Danilewicz JC (2003) Review of Reaction Mechanisms of Oxygen and Proposed Intermediate Reduction Products in Wine: Central Role of Iron and Copper. American Journal of Enology and Viticulture 54(2): 73-85
Iron Content Quality
Iron deficiency symptom quality assessment in Burgundy Chardonnay on Kimmeridgian limestone soils documented the characteristic symptoms and severity gradient associated with lime-induced chlorosis in this classic calcareous terroir. Visual iron deficiency symptom quality evaluation revealed the diagnostic interveinal chlorosis pattern on young leaves, with bright yellow interveinal tissue contrasting sharply against green primary and secondary veins, symptoms concentrated on the apical 4-6 leaves while older leaves remained normally green. Symptom severity scoring of iron deficiency symptom quality on a 0-5 scale ranged from 2 (moderate interveinal yellowing with distinct green veins) through 4 (severe yellowing approaching white, veins losing green) to 5 in worst-affected areas (complete bleaching of young leaves, growing point necrosis). The spatial distribution of iron deficiency symptom quality correlated with active lime content variation, with symptoms absent where active lime was below 8% (pH 7.6), moderate at 10-15% active lime (pH 8.0-8.2), and severe above 18% active lime (pH 8.4+). Tissue analysis confirmed the iron deficiency symptom quality diagnosis with young leaf Fe at 38 ppm in symptomatic areas versus 85 ppm in asymptomatic areas (sufficiency threshold 60 ppm), though total soil Fe exceeded 15,000 ppm demonstrating that availability rather than quantity limited iron nutrition. Yield assessment showed iron deficiency symptom quality impact with 35% yield reduction in severely affected zones compared to unaffected areas, attributed to reduced photosynthetic capacity, poor fruit set, and smaller berry size, establishing the substantial economic significance of iron chlorosis in this high-value vineyard.
Iron deficiency symptom quality is a micronutrient deficiency symptom quality that inheres in a grapevine organism and represents the characteristic visual symptoms resulting from inadequate iron availability for chlorophyll synthesis and electron transport functions, including the diagnostic interveinal chlorosis on youngest leaves where leaf tissue between veins becomes yellow to white while veins initially remain green, progressing in severe cases to complete bleaching of young leaves and necrosis of growing points, with iron deficiency representing the most common and economically significant micronutrient limitation in viticulture particularly on calcareous soils where high pH and bicarbonate ions interfere with iron uptake and utilization. This quality captures the grapevine response to iron limitation, recognizing that iron deficiency (commonly termed "lime-induced chlorosis" when associated with calcareous soils) affects extensive vineyard areas in major wine regions including Champagne, Burgundy, Cognac, Rioja, and parts of California, causing yield reductions of 20-50% in severely affected vineyards. The ontological significance of iron deficiency symptom quality lies in its economic impact and the management challenge it presents, where soil iron content may be adequate but chemical unavailability prevents plant acquisition, requiring specialized management approaches including tolerant rootstock selection, soil acidification, and chelated iron application rather than simple iron fertilization. The biological basis of iron deficiency symptoms encompasses the essential role of iron in chlorophyll biosynthesis where iron is required for protochlorophyllide synthesis though not present in the final chlorophyll molecule, the function of iron in cytochromes and iron-sulfur proteins of the photosynthetic and respiratory electron transport chains, and the consequent impairment of both chlorophyll production and photosynthetic function when iron is limiting. The phloem immobility of iron means that deficiency symptoms appear first and most severely on youngest leaves that cannot acquire iron through redistribution from older tissues, creating the characteristic acropetal symptom progression that distinguishes iron from magnesium deficiency despite similar interveinal chlorosis patterns.
Iron_Deficiency_Symptom_Quality is_a Micronutrient_Deficiency_Symptom_Quality, serving as parent class for Lime_Induced_Chlorosis_Quality and Interveinal_Yellowing_Quality_(Fe) which address specific aspects of iron deficiency. This class addresses the overall iron deficiency syndrome while child classes capture particular symptom types and causation. Most common micronutrient deficiency in viticulture due to prevalence of calcareous soils. Phloem immobility means symptoms on youngest leaves first (opposite of Mg interveinal chlorosis). Tissue Fe threshold ~60 ppm soil Fe often adequate but chemically unavailable. Rootstock selection for lime tolerance is primary management strategy on calcareous sites.
Eisen-Limitation
Eisenchlorose
Eisenmangel-Symptome
Fe deficiency symptoms
Fe-Mangel-Zeichen
ferric deficiency indicators
iron chlorosis
iron limitation signs
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Mengel K (1994) Iron availability in plant tissues — iron chlorosis on calcareous soils. Plant and Soil 165:275-283 Bavaresco L et al. (2010) Grapevine iron nutrition: rootstock effects on scion response to lime-induced chlorosis. Acta Horticulturae 868:115-122
Iron Deficiency Symptom Quality
Fercal (BC1 of 31R × Berlandieri Colombard 1B) demonstrates exceptional iron efficiency with FCR activity maintained above 3 µmol Fe²⁺·g⁻¹ root·h⁻¹ even at bicarbonate concentrations of 20 mM, supporting chlorophyll-rich Ugni Blanc foliage (>500 µg·g⁻¹) on the highly calcareous soils of Cognac where other rootstocks show severe chlorosis. 41B Millardet et de Grasset shows high iron efficiency maintaining Chardonnay on Champagne chalk without chlorosis symptoms despite active lime exceeding 30%. 140 Ruggeri (V. berlandieri × V. rupestris) provides iron efficiency for Grillo on the calcareous clay soils of western Sicily, maintaining leaf Fe concentrations above 80 mg·kg⁻¹ dry weight where sensitive rootstocks decline below 40 mg·kg⁻¹.
A nutrient acquisition rootstock role that inheres in a grafted rootstock by virtue of its enhanced capacity for iron (Fe) acquisition, translocation, and delivery to the scion, maintaining adequate leaf iron status and preventing chlorosis under conditions of limited soil Fe availability or high soil pH that restricts Fe solubility. Iron-efficient rootstocks overcome Fe limitation through Strategy I iron acquisition mechanisms including elevated Fe³⁺-chelate reductase (FCR) activity in root plasma membranes, enhanced proton efflux capacity for rhizosphere acidification, increased Fe²⁺ transporter (IRT1) expression, and efficient Fe translocation via citrate and nicotianamine chelation in xylem sap. Iron efficiency is quantified by FCR activity per unit root surface area (efficient genotypes maintaining >2 µmol Fe²⁺·g⁻¹ root·h⁻¹ under bicarbonate stress), by leaf chlorophyll concentration under Fe-limiting conditions (efficient genotypes >400 µg·g⁻¹ fresh weight), and by absence of interveinal chlorosis symptoms on calcareous soils. Iron efficiency substantially overlaps with lime tolerance but involves distinct physiological components.
Iron_Efficient_Rootstock_Role captures the physiological capacity for Fe acquisition that underlies lime tolerance but may be expressed even on non-calcareous soils where other factors (waterlogging, compaction, nematode damage) limit Fe uptake. V. berlandieri parentage contributes the highest iron efficiency, as this species evolved on highly calcareous Edwards Plateau soils. Iron efficiency is a complex trait involving root Fe³⁺ reduction, rhizosphere acidification, and shoot Fe utilization rootstocks may show different rankings for efficiency under different forms of Fe stress. Foliar Fe applications can partially compensate for rootstock Fe inefficiency but do not address root zone causes of deficiency.
Eisenchlorose-resistente Unterlage
Fe efficient rootstock
Fe-effiziente Unterlage
chlorosis tolerant rootstock
eiseneffiziente Unterlage
iron chlorosis resistant rootstock
iron efficient rootstock
Bavaresco L et al. (2003) Grapevine iron nutrition update, Journal International des Sciences de la Vigne et du Vin 37: 89-100 Rombolà AD et al. (2002) Iron source affects iron reduction and re-greening of kiwifruit leaves, Journal of Plant Nutrition 25: 1771-1785 Tagliavini M, Rombolà AD (2001) Iron deficiency and chlorosis in orchard and vineyard ecosystems, European Journal of Agronomy 15: 71-92 Brancadoro L et al. (1995) Mechanisms of Fe efficiency in roots of Vitis spp., Plant and Soil 171: 229-234
Iron Efficient Rootstock Role
Kinetic characterization of Iron_Reductase_Activity_Disposition at the University of Milan measured ferric chelate reductase (FCR) activity across 18 rootstocks using the standard Fe(III)-EDTA/BPDS assay under varying conditions of iron status (Fe-sufficient, Fe-deficient) and bicarbonate (0, 5, 10 mM NaHCO₃). Root segments were incubated with Fe(III)-EDTA substrate and the Fe²⁺ product was trapped and quantified by bathophenanthroline disulfonate (BPDS) formation. Iron_Reductase_Activity_Disposition showed remarkable genotypic variation. '41B' exhibited exceptional reductase capacity with activity increasing from 28 nmol Fe²⁺ g⁻¹ FW h⁻¹ under Fe sufficiency to 185 nmol g⁻¹ h⁻¹ under deficiency (6.6-fold induction), with activity localized to the root hair zone and subapical regions as demonstrated by histochemical staining. 'Fercal' showed similar strong induction (7.2-fold, reaching 210 nmol g⁻¹ h⁻¹). '110 Richter' displayed moderate Iron_Reductase_Activity_Disposition (125 nmol g⁻¹ h⁻¹, 4-fold induction). '3309C' exhibited weak activity with only 45 nmol g⁻¹ h⁻¹ under deficiency (2.1-fold induction) and reached maximum at lower substrate concentration, suggesting lower enzyme abundance. Bicarbonate sensitivity differed among rootstocks: '41B' retained 65% of reductase activity at 10 mM HCO₃⁻ while '3309C' was inhibited to 30%. Gene expression analysis showed that VvFRO2 transcript levels under Fe deficiency were 8-fold higher in '41B' than '3309C', correlating with activity differences (r = 0.91 between expression and activity across genotypes). These findings establish Iron_Reductase_Activity_Disposition as a key discriminating trait for lime tolerance.
An Iron_Reductase_Activity_Disposition is an Iron_Acquisition_Efficiency_Disposition that inheres in Vitis root epidermal cells and is realized through high activity of plasma membrane-bound ferric chelate reductase enzymes (FRO family) that reduce solubilized Fe³⁺-chelates to Fe²⁺, the form required for uptake by IRT1 iron transporters. This disposition represents the second and often rate-limiting component of Strategy I iron acquisition, providing the essential reduction step linking Fe³⁺ solubilization (by acidification) to Fe²⁺ uptake (by IRT transporters). Ferric reductase activity is obligatory because IRT1 transporters accept only Fe²⁺, while iron in calcareous soils and most organic chelates exists as Fe³⁺. The ferric reductase operates at the root cell plasma membrane, accepting electrons from cytoplasmic NADPH and transferring them across the membrane to reduce extracellular Fe³⁺-chelate complexes, releasing Fe²⁺ for immediate uptake by co-localized IRT1 transporters. Iron_Reductase_Activity_Disposition varies substantially among rootstock genotypes in both constitutive activity levels and the magnitude of upregulation under iron deficiency, with lime-tolerant genotypes typically showing 4-8 fold induction of reductase activity under Fe deficiency compared to 1.5-3 fold in sensitive genotypes. The enzyme is sensitive to bicarbonate inhibition, contributing to the chlorosis-inducing effect of high soil HCO₃⁻, with tolerant genotypes showing less bicarbonate sensitivity.
Iron_Reductase_Activity_Disposition is_a Iron_Acquisition_Efficiency_Disposition. Ferric reductase provides the obligatory reduction of Fe³⁺ to Fe²⁺ for uptake. Activity is highly iron-regulated, with tolerant genotypes showing stronger induction. VvFRO genes encode the ferric reductase enzymes. Bicarbonate directly inhibits reductase activity, with tolerant genotypes showing less sensitivity. Reductase activity often correlates better with field lime tolerance than acidification alone. Activity is localized to root hair zone and subapical epidermis.
Eisenreduktase-Aktivitäts-Disposition
FCR activity
FRO enzyme capacity
FRO-Enzym-Kapazität
Ferri-Chelat-Reduktase
ferric chelate reductase
Brancadoro L et al. (1995) Mechanisms of Fe-efficiency in roots of Vitis spp. in response to iron deficiency stress, Plant and Soil 171:229-234 Robinson NJ et al. (1999) A ferric-chelate reductase for iron uptake from soils, Nature 397:694-697 Ksouri R et al. (2006) Genotypic variability within Tunisian grapevine varieties (Vitis vinifera L.) facing bicarbonate-induced iron deficiency, Plant Physiology and Biochemistry 44:315-322
Iron Reductase Activity Disposition
Iron uptake efficiency comparison among rootstocks on calcareous soil (45% active lime, pH 8.3) reveals the dramatic genotypic variation determining scion iron nutrition, with Fercal showing high Fe uptake efficiency (leaf Fe 85 mg/kg, no chlorosis, strong rhizosphere acidification and ferric reductase activity), 41B intermediate (leaf Fe 62 mg/kg, mild transient chlorosis), and SO4 showing poor efficiency (leaf Fe 38 mg/kg, severe interveinal chlorosis, 40% yield reduction), demonstrating how rootstock iron uptake efficiency determines viticulture viability on calcareous soils through Strategy I mechanism effectiveness.
Iron uptake efficiency quality is a micronutrient uptake efficiency quality that represents the capacity of grapevine roots to acquire iron from soil despite its extremely low solubility in aerobic conditions, characterizing the effectiveness of iron mobilization and uptake strategies that determine vine iron status affecting chlorophyll synthesis, electron transport, and numerous enzyme functions essential for photosynthesis and respiration. Within the BFO framework, this quality specifically addresses the iron component of micronutrient uptake, distinguishing iron acquisition from other trace elements while recognizing that iron presents unique challenges from its precipitation as insoluble hydroxides at neutral to alkaline pH, making iron deficiency (lime-induced chlorosis) the most common micronutrient disorder in viticulture on calcareous soils. The ontological significance lies in iron uptake efficiency determining access to this essential element required for chlorophyll biosynthesis, cytochrome function, and Fe-S cluster enzymes central to electron transport, where iron deficiency causes characteristic interveinal chlorosis as chlorophyll synthesis fails while veins remain green (iron immobility preventing redistribution), severely limiting photosynthetic capacity and vine productivity. The mechanistic basis for iron uptake in grapevine involves Strategy I mechanisms characteristic of dicots, including plasma membrane H⁺-ATPase acidification of the rhizosphere (increasing Fe³⁺ solubility), ferric chelate reductase (FRO) reducing Fe³⁺ to Fe²⁺, and IRT (Iron-Regulated Transporter) proteins importing Fe²⁺ into root cells, with each component potentially limiting iron acquisition depending on conditions. The soil pH effect on iron availability is dramatic, with soluble iron decreasing approximately 1000-fold for each unit increase in pH above 4, such that calcareous soils (pH 7.5-8.5) may have soluble iron concentrations of only 10⁻¹⁰ M despite high total iron, creating severe acquisition challenges that only efficient Strategy I mechanisms can address. The rootstock genotype critically affects iron uptake efficiency, with rootstocks containing Vitis berlandieri genetics often showing susceptibility to lime-induced chlorosis (LIC) on high-pH soils, while certain selections (41B, Fercal) demonstrate enhanced iron acquisition through more effective rhizosphere acidification and ferric reductase activity. Understanding iron uptake efficiency quality enables assessment of lime-induced chlorosis risk, rootstock selection for calcareous soils, and management strategies including iron chelate application for sites where root iron acquisition is limiting.
Iron_Uptake_Efficiency_Quality addresses the critical challenge of acquiring iron from calcareous soils where lime-induced chlorosis limits viticulture. This quality involves Strategy I mechanisms (acidification, reduction, transport) determining chlorosis susceptibility. The class supports rootstock selection for calcareous sites through Fe uptake efficiency characterization.
Chloroseresistenz
Eisenaufnahmeeffizienz
Fe uptake efficiency
Fe-Akquisition
Strategie-I-Effizienz
Strategy I efficiency
chlorosis resistance
iron acquisition
Brancadoro L et al. (1995) Mechanisms of Fe-efficiency in roots of Vitis spp. in response to iron deficiency stress. Plant and Soil 171(2): 229-234 Jiménez S et al. (2007) Influence of iron chlorosis on fruit quality in peach. International Journal of Food Science and Technology 42(1): 43-49
Iron Uptake Efficiency Quality
Shiraz vines in the Barossa Valley bearing an irrigated role receive supplemental water via subsurface drip lines delivering 0.5-2.5 ML/ha/season depending on vintage conditions, with irrigation scheduling guided by midday stem water potential measurements targeting -1.0 to -1.4 MPa during the pre-véraison period, enabling commercially viable production at 8-14 t/ha in a region receiving only 250-350 mm of growing season rainfall.
A production system role that inheres in a grapevine receiving supplemental water from an artificial delivery system at any point during the growing season, whether for full crop water requirement replacement, deficit management, or frost protection, distinguishing the vine from dry-farmed management.
Irrigated_Role encompasses all forms of supplemental water delivery regardless of volume, timing, or technology. The role is mutually exclusive with Dry_Farmed_Role. In many traditional European wine regions, irrigation was historically prohibited or severely restricted by appellation regulations: the EU progressively relaxed these restrictions from 2006 onward in response to climate change pressures. Subclasses distinguish the primary irrigation technology (drip, flood) and management strategy (deficit irrigation), reflecting fundamentally different approaches to vine water supply with distinct implications for fruit quality, yield, and resource efficiency.
Bewässerungsanbau-Rolle
bewässerte Produktionsrolle
irrigated vine role
irrigated viticulture role
künstlich bewässerte Rolle
supplementally watered production role
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 498-520 Williams LE and Matthews MA (1990) Grapevine, in Stewart BA and Nielsen DR (eds) Irrigation of Agricultural Crops, ASA Monograph 30 pp: 1019-1055
Irrigated Role
The irrigation protocol description for a premium Cabernet Sauvignon vineyard in the Napa Valley specifies a subsurface drip irrigation system with 2 L/h emitters at 0.9 m spacing on buried laterals at 0.3 m depth, regulated deficit irrigation (RDI) strategy applying 50% of reference crop evapotranspiration (ETc) from fruit set through véraison and 30% of ETc from véraison through harvest, soil moisture monitoring via neutron probe access tubes at 0.3, 0.6, and 0.9 m depths with irrigation triggered when soil water content at 0.6 m falls below 60% of field capacity, and predawn leaf water potential targets of -0.3 to -0.5 MPa during the pre-véraison deficit period — the protocol integrates weather station ET data, soil moisture measurements, and plant-based water status indicators into a decision framework that manages vine water stress to optimize berry size reduction, anthocyanin concentration, and tannin polymerization.
A viticultural practice description that documents the system, scheduling, strategy, and quantitative parameters governing the application of supplemental water to grapevines, encompassing the complete set of decisions and technical specifications that determine when, how much, and by what method water is delivered to the vineyard to manage vine water status in relation to developmental stage, target wine style, and water resource constraints.
Irrigation_Protocol_Description addresses one of the most actively debated and regionally variable domains in modern viticulture. The role of irrigation ranges from prohibited (many traditional European appellations including all French AOC until regulatory relaxation in 2006) through supplemental (used only to prevent vine damage during extreme drought) to essential (semi-arid and arid wine regions where viticulture is impossible without irrigation): this regulatory and philosophical diversity reflects fundamentally different perspectives on whether irrigation constitutes legitimate terroir management or artificial manipulation of wine character. The progressive relaxation of irrigation restrictions in European appellations — driven by climate change increasing drought frequency and severity — and the simultaneous development of precision irrigation technologies in New World regions have created a convergence toward managed deficit irrigation as the dominant paradigm in quality-oriented viticulture globally. Subclasses capture the key informational components of irrigation documentation.
Beregnungsmanagement-Dokumentation
Bewässerungspraxis-Beschreibung
Bewässerungsprotokoll-Beschreibung
irrigation management record
irrigation practice documentation
vineyard water management protocol
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 381-420 Williams LE and Ayars JE (2005) Grapevine water use and the crop coefficient are linear functions of the shaded area measured beneath the canopy, Agricultural and Forest Meteorology 132: 201-211 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data, Annals of Botany 105: 661-676
Irrigation Protocol Description
The irrigation scheduling protocol for Sauvignon Blanc in Marlborough integrates three data streams — daily reference evapotranspiration (ETo) from an on-site weather station multiplied by a crop coefficient (Kc) of 0.3-0.7 depending on canopy development stage to calculate crop water demand (ETc), continuous capacitance soil moisture probe readings at 0.3 m and 0.6 m depths compared against a refill threshold of 50% plant-available water, and fortnightly predawn leaf water potential measurements targeting -0.2 to -0.3 MPa during vegetative growth and -0.4 to -0.6 MPa during the post-véraison deficit period — irrigation is triggered when any two of the three indicators simultaneously signal water need, with the multi-parameter approach reducing the risk of over-irrigation that single-indicator scheduling can produce.
An irrigation protocol description that documents the decision rules, monitoring methods, and trigger criteria used to determine when irrigation events are initiated and how long they last, specifying the integration of atmospheric demand data (reference evapotranspiration), soil moisture status, plant water status indicators, and phenological stage into a scheduling framework that matches water supply to vine requirements throughout the growing season.
Irrigation_Scheduling_Protocol addresses the most complex real-time decision-making component of irrigation management — determining when to irrigate and how much water to apply at each event. Scheduling approaches range from simple calendar-based schedules (irrigate every 7 days regardless of conditions) through soil-based scheduling (irrigate when soil moisture falls below a defined threshold) and atmospheric-demand-based scheduling (replace a percentage of calculated ETc) to integrated plant-soil-atmosphere monitoring systems: the trend in precision viticulture is toward multi-parameter scheduling that combines the strengths of different monitoring approaches while compensating for the limitations of any single indicator. The phenological stage provides the overarching framework — scheduling parameters are typically adjusted at budburst, flowering, fruit set, véraison, and harvest to match the changing water requirement profile of the vine and the quality objectives associated with each developmental phase.
Beregnungszeitplan
Bewässerungsentscheidungs-Rahmenwerk
Bewässerungssteuerungs-Protokoll
irrigation decision framework
irrigation timing protocol
water scheduling method
Williams LE and Ayars JE (2005) Grapevine water use and the crop coefficient are linear functions of the shaded area measured beneath the canopy, Agricultural and Forest Meteorology 132: 201-211 Acevedo-Opazo C et al. (2010) Effects of grapevine (Vitis vinifera L.) water status on water consumption, vegetative growth and grape quality, Irrigation Science 28: 231-243 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 395-420
Irrigation Scheduling Protocol
The irrigation system type for a premium Shiraz vineyard in McLaren Vale is specified as surface drip irrigation with pressure-compensating emitters (Netafim DripNet PC 2.0 L/h) at 0.6 m spacing on polyethylene laterals positioned 0.3 m from the vine row on the northern (sun-exposed) side — drip irrigation achieves application efficiencies of 85-95% compared to 60-75% for overhead sprinkler and 40-60% for furrow irrigation, enables precise deficit irrigation management by controlling wetted soil volume (typically 30-40% of total root zone under drip versus 100% under overhead), and avoids wetting the canopy and fruit zone that would increase Botrytis risk in disease-prone environments.
An irrigation protocol description consisting of the name and technical classification of the water delivery infrastructure used to apply irrigation water to the vineyard, distinguishing among drip (surface and subsurface), micro-sprinkler, overhead sprinkler, furrow, and flood irrigation methods that differ fundamentally in water application efficiency, uniformity, root zone wetting pattern, and compatibility with precision water management strategies.
Irrigation_System_Type defines the physical infrastructure that constrains all subsequent irrigation management decisions. The global transition from flood and furrow irrigation to drip systems in premium viticulture represents one of the most significant technological shifts of the past 50 years — beginning in Israel in the 1960s (Netafim's invention of commercial drip irrigation in 1965) and progressively adopted across Australian, Californian, Chilean, South African, and increasingly European vineyards: drip irrigation's advantages include water savings of 30-50% compared to overhead systems, elimination of canopy wetting, precise spatial control of water application, compatibility with fertigation (nutrient delivery through irrigation water), and the ability to implement deficit irrigation strategies that require controlled rather than generous water supply. Subsurface drip irrigation (SDI), with emitter lines buried at 0.2-0.4 m depth, further reduces evaporative losses and eliminates surface wetting but requires higher installation investment and complicates system monitoring and maintenance.
Beregnungsanlagen-Art
Bewässerungsinfrastruktur-Klassifikation
Bewässerungssystem-Typ
irrigation delivery method
irrigation infrastructure classification
water application system type
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 381-395 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 168-195 Netafim (2023) Drip Irrigation for Vineyards technical documentation
Irrigation System Type
Mechanistic investigation of Isohydric_Behavior_Disposition at INRAE Bordeaux compared the archetypal isohydric cultivar 'Grenache' with anisohydric 'Syrah' under controlled drought progression, examining the physiological basis of water potential homeostasis. Potted vines were subjected to soil drying over 21 days with daily measurements of predawn water potential (Ψpd, indicator of soil water status), midday water potential (Ψmd), stomatal conductance, and xylem sap ABA. 'Grenache' exhibited classic Isohydric_Behavior_Disposition with Ψmd maintained between -1.2 and -1.4 MPa throughout drought despite Ψpd declining from -0.2 to -0.9 MPa. This homeostasis was achieved through progressive stomatal closure, with conductance declining from 280 to 35 mmol m⁻² s⁻¹ (87% reduction) as drought intensified. ABA analysis revealed that 'Grenache' accumulated xylem ABA earlier (detectable increase at Ψpd = -0.3 MPa) and more strongly (peak 2800 ng L⁻¹) than 'Syrah' (increase at -0.6 MPa, peak 1400 ng L⁻¹). Guard cell sensitivity assays confirmed that 'Grenache' stomata were 2.3-fold more responsive to ABA than 'Syrah' stomata. Hydraulic vulnerability curves showed that 'Grenache' maintained Ψmd above its P50 (water potential causing 50% loss of hydraulic conductivity = -1.8 MPa) throughout drought, whereas 'Syrah' Ψmd approached and briefly exceeded its P50 (-2.1 MPa). The Isohydric_Behavior_Disposition of 'Grenache' thus reflects combined early ABA accumulation and high stomatal ABA sensitivity, prioritizing hydraulic safety through strict Ψmd regulation.
An Isohydric_Behavior_Disposition is a Stomatal_Regulation_Disposition that inheres in a Vitis vinifera plant and is realized through tight stomatal control that maintains relatively constant midday leaf water potential regardless of soil water availability, prioritizing hydraulic safety and tissue hydration over carbon assimilation during drought. This disposition represents the water-conserving end of the stomatal regulation spectrum, with isohydric cultivars responding to early drought signals by closing stomata to prevent the decline in leaf water potential that would otherwise occur as soil water becomes limiting. The term "isohydric" (Greek: equal water) refers to the maintenance of stable leaf water status across varying soil moisture conditions, achieved through sensitive stomatal responses to hydraulic and chemical signals. Isohydric behavior involves high sensitivity to abscisic acid (ABA), rapid stomatal closure in response to declining xylem water potential, and possibly root-to-shoot chemical signaling that triggers stomatal closure before leaf water potential declines significantly. The adaptive value of Isohydric_Behavior_Disposition lies in protection against hydraulic failure (xylem cavitation and embolism) and maintenance of leaf tissue integrity during drought, enabling rapid recovery when water becomes available. However, this conservative strategy incurs costs in reduced photosynthesis and growth during drought periods, potentially limiting productivity in environments where moderate drought is common but not severe. Classic isohydric grapevine cultivars include 'Grenache' and 'Tempranillo'.
Isohydric_Behavior_Disposition is_a Stomatal_Regulation_Disposition. Isohydric behavior maintains stable leaf water potential through sensitive stomatal closure, prioritizing hydraulic safety over carbon gain. The mechanism involves early ABA accumulation and high guard cell ABA sensitivity. Isohydric cultivars are well-suited to severe drought environments where hydraulic failure is a significant risk. Trade-offs include reduced photosynthesis and growth under moderate drought where stomata close preemptively. 'Grenache' and 'Tempranillo' are classic isohydric cultivars.
Isohydrisches-Verhalten-Disposition
Konservative-Wassernutzung
Wasserpotential-Homöostase
conservative water use
isohydric stomatal behavior
water potential homeostasis
Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought, Plant Cell and Environment 26:1393-1405 Tardieu F, Simonneau T (1998) Variability among species of stomatal control under fluctuating soil water status and evaporative demand, Journal of Experimental Botany 49:419-432 Soar CJ et al. (2006) Grape vine varieties Shiraz and Grenache differ in their stomatal response to VPD: apparent links with ABA physiology and gene expression in leaf tissue, Australian Journal of Grape and Wine Research 12:2-12
Isohydric Behavior Disposition
Isohydric behavior quality characterization in Châteauneuf-du-Pape compared Grenache and Syrah responses to progressive soil drying under controlled conditions, revealing the contrasting strategies underlying their field performance differences. Grenache demonstrated pronounced isohydric behavior quality with midday leaf water potential remaining within the narrow range of -1.2 to -1.4 MPa across soil moisture conditions spanning field capacity to severe deficit, achieved through stomatal conductance reductions from 250 mmol m⁻² s⁻¹ under wet conditions to below 50 mmol m⁻² s⁻¹ under moderate stress, representing 80% closure to maintain constant water potential. The isohydric behavior quality of Grenache provided hydraulic safety benefits but limited photosynthesis to 35% of potential under moderate stress when Syrah at equivalent soil moisture maintained 65% of maximum photosynthesis through its contrasting anisohydric behavior. Field observations confirmed that Grenache's isohydric behavior quality resulted in earlier visible stress symptoms as stomatal closure limited cooling, but protected the cultivar from hydraulic damage during extreme stress events that caused xylem cavitation in the more risk-tolerant Syrah, demonstrating the survival advantages of the isohydric strategy under severe drought conditions even at the cost of reduced productivity under moderate stress.
Isohydric behavior quality is a stomatal response to drought quality that inheres in a grapevine cultivar or clone and represents the drought response strategy characterized by sensitive stomatal regulation that maintains relatively constant leaf water potential across a range of soil water availability conditions, closing stomata early and progressively as soil dries to limit transpirational water loss and prevent the decline in leaf water status that would occur if stomata remained open under increasing evaporative demand with diminishing supply. This quality captures one endpoint of the isohydric-anisohydric behavioral continuum, with strongly isohydric cultivars exhibiting tight stomatal control that sacrifices carbon assimilation to maintain hydraulic stability, representing a conservative drought strategy that prioritizes survival and hydraulic safety over productivity under water-limited conditions. The ontological significance of isohydric behavior quality lies in its characterization of a fundamental drought adaptation strategy with specific implications for vine performance, water use, and management requirements under deficit irrigation, where isohydric cultivars may maintain safer water status but at the cost of reduced photosynthesis that could limit fruit ripening capacity if stress is not carefully managed. The biological basis of isohydric behavior involves high sensitivity of stomatal guard cells to the signals indicating developing water deficit, including root-sourced ABA that increases as soil dries and leaf hydraulic signals that trigger closure before significant water potential decline occurs, combined with relatively conservative hydraulic architecture that amplifies these protective responses. The characteristic feature of isohydric behavior is the maintenance of relatively stable midday leaf water potential (within approximately 0.2 MPa range) across a wide range of soil moisture conditions, achieved through progressively greater stomatal restriction that reduces transpiration as soil water availability declines, creating the diagnostic pattern of constant water potential despite variable soil conditions that defines the isohydric strategy.
Isohydric_Behavior_Quality is_a Stomatal_Response_To_Drought_Quality, representing the conservative drought strategy maintaining stable leaf water potential through sensitive stomatal closure. This class addresses one endpoint of the isohydric-anisohydric behavioral spectrum. The diagnostic feature is constant midday water potential (±0.2 MPa) across variable soil moisture. Distinguished from Anisohydric_Behavior_Quality which represents the opposite strategy allowing water potential to vary with conditions. Grenache is the classic isohydric Vitis vinifera cultivar. The strategy provides hydraulic safety at the cost of reduced productivity under moderate stress but confers survival advantages under severe drought.
Wasserpotential-Regulation
hydraulic safety behavior
hydraulische Sicherheitsstrategie
isohydric strategy
isohydrisches Verhalten
konservative Wasserstrategie
water potential regulation
Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell and Environment 26:1393-1405 Tardieu F, Simonneau T (1998) Variability among species of stomatal control under fluctuating soil water status and evaporative demand: modelling isohydric and anisohydric behaviours. Journal of Experimental Botany 49:419-432 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data. Annals of Botany 105:661-676
Isohydric Behavior Quality
Sangiovese clone R24 bears an Italian_Clone_Role as a VCR Rauscedo selection, evaluated through Tuscan trials and registered in the Registro Nazionale for its suitability to premium Brunello di Montalcino and Chianti Classico production—demonstrating the Italian system's emphasis on regional quality objectives. Nebbiolo clone CVT 71 serves in an Italian_Clone_Role from the Conegliano research station, selected for optimal anthocyanin development and tannin structure in Barolo and Barbaresco production. Prosecco (Glera) clone VCR 101 bears an Italian_Clone_Role as a high-performing selection for the massive Prosecco industry, VCR selection supporting the billions of bottles produced annually from northeastern Italian vineyards.
A regional clone role that inheres in a grapevine clone registered within the Italian national clone certification system, coordinated through the Registro Nazionale delle Varietà di Vite (National Registry of Grapevine Varieties) and implemented by authorized regional selection programs including VCR (Vivai Cooperativi Rauscedo), CRA-VIT (formerly Istituto Sperimentale per la Viticoltura, Conegliano), and regional agricultural research stations. Italian clones are identified by variety name, institutional prefix, and numeric code (e.g., Sangiovese R24 for Rauscedo selection, Nebbiolo CVT 71 for Conegliano selection), with nomenclature varying by selecting institution. The Italian clone system reflects the country's extraordinary variety diversity (over 500 authorized varieties) and regional viticultural traditions, with selection programs targeting both major international varieties and regionally important cultivars. Italian clone evaluation emphasizes wine quality assessment, with extended winemaking trials central to registration decisions. VCR Rauscedo has become the world's largest grapevine nursery, distributing Italian-registered clones globally alongside major rootstock production that supplies international markets.
Italian_Clone_Role captures the distributed Italian clone system where multiple authorized institutions conduct selection and registration within a national coordination framework. The Italian approach reflects regional viticultural diversity different institutions have focused on different variety portfolios (VCR Rauscedo on international varieties and rootstocks, Conegliano on northeastern varieties, regional stations on local cultivars). Italian clone nomenclature is less standardized than the French numeric system, with various prefixes (R, VCR, CVT, I-xxx) indicating institutional origin. Italy's extraordinary variety diversity (second only to Georgia) means the clone system addresses many locally important varieties rarely encountered elsewhere, supporting regional wine identities that would be lost if only internationally traded varieties received selection attention.
Italian clone
Italian registered clone
Registro clone
Registro-Klon
VCR clone
italienisch registrierter Klon
italienischer Klon
Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 Ministero delle Politiche Agricole (2020) Registro Nazionale delle Varietà di Vite Schneider A et al. (2008) Clonal variation in grapevine, ISHS Acta Horticulturae 779: 397-404 Crespan M et al. (2006) Molecular characterization of Italian grapevine cultivars, Vitis 45: 69-73
Italian Clone Role
Systematic evaluation of Japanese_Beetle_Resistance_Disposition at Cornell University's New York State Agricultural Experiment Station compared feeding damage across 45 Vitis accessions under natural Japanese beetle pressure over five seasons in Geneva, NY. Defoliation was assessed weekly during the July-August beetle flight period using a 0-100% visual scale, with cumulative feeding expressed as beetle-days calculated from trap catches × defoliation integration. V. vinifera 'Concord' and V. labrusca hybrids showed high susceptibility with 65-80% defoliation at season peak and cumulative damage equivalent to 850-1,100 beetle-days. In contrast, V. vinifera 'Cayuga White' demonstrated strong Japanese_Beetle_Resistance_Disposition with only 15-25% peak defoliation and cumulative damage of 180-280 beetle-days despite equivalent beetle pressure. Laboratory choice tests confirmed field rankings: beetles offered paired leaf discs consistently consumed 3.8× more 'Concord' than 'Cayuga White' tissue over 24-hour trials. Phytochemical analysis revealed that 'Cayuga White' leaves contained 2.4-fold higher concentrations of quercetin glycosides, compounds with documented beetle-deterrent activity. Foliar application of quercetin extract to susceptible 'Concord' reduced beetle feeding by 45% in subsequent choice tests, providing proof-of-concept that the resistance mechanism involves feeding deterrent chemistry that could potentially be enhanced through breeding or management practices.
A Japanese_Beetle_Resistance_Disposition is an Insect_Resistance_Disposition that inheres in a Vitis plant foliage and is realized through defense processes that limit feeding, aggregation, or damage by the Japanese beetle (Popillia japonica), an invasive scarabaeid beetle that has become a significant defoliating pest of grapevine in eastern North America and poses ongoing invasion threats to other regions including Europe. This disposition represents resistance to an insect whose adult stage causes the economically important damage through voracious feeding on leaf tissue between veins, producing characteristic "skeletonized" leaves with only the vascular network remaining. Japanese beetles are gregarious feeders, with initial feeding damage releasing aggregation pheromones that attract additional beetles, potentially resulting in rapid, severe defoliation of preferred host plants including grapevine. Heavy defoliation reduces photosynthetic capacity, delays fruit ripening, increases sunburn on exposed clusters, and weakens vines entering dormancy. Resistance mechanisms may include leaf chemical composition deterring feeding initiation or continuation, leaf physical properties (thickness, toughness, trichome density) reducing palatability, volatile profiles less attractive to host-seeking beetles, and induced defenses reducing nutritional quality or producing deterrent compounds following initial damage. Substantial variation in Japanese beetle feeding preference among Vitis species and V. vinifera cultivars has been documented, with some cultivars consistently showing reduced damage providing opportunities for cultivar selection and breeding. Japanese_Beetle_Resistance_Disposition is particularly relevant for eastern North American viticulture where the pest is established and expanding, and for European regions preparing for potential invasion.
Japanese_Beetle_Resistance_Disposition is_a Insect_Resistance_Disposition. P. japonica is an invasive pest from Asia established in eastern North America since 1916, with recent spread to Europe (first detections in Italy 2014, Switzerland 2017). Adult beetles cause damage through foliage skeletonization larvae feed on grass roots and do not affect grapevine directly. Variation in susceptibility among Vitis species and cultivars provides opportunities for cultivar selection in affected regions. The beetle's invasive spread trajectory suggests increasing importance for European viticulture in coming decades, making resistance research a proactive priority.
Japanese beetle tolerance
Japankäfer-Fraßresistenz
Japankäfer-Resistenz-Disposition
Popillia resistance
Popillia-japonica-Abwehr
scarab beetle resistance
Hammons DL et al. (2009) Attraction of Japanese beetles to host plants and conspecific females, Journal of Economic Entomology 102:585-593 Loughrin JH et al. (1996) Volatile compounds induced by Japanese beetle feeding on susceptible plants, Journal of Chemical Ecology 22:639-658 Potter DA, Held DW (2002) Biology and management of the Japanese beetle, Annual Review of Entomology 47:175-205
Japanese Beetle Resistance Disposition
Concord bears a Juice_Production_Scion_Role as the dominant variety for American grape juice production (over 300,000 tonnes annually), with its intense "grapey" labrusca aroma (from methyl anthranilate at 2-5 mg/L), deep purple color (anthocyanins exceeding 1500 mg/kg), and cultural association with grape juice defining the category—Welch's grape juice being the iconic product since 1869. Niagara serves in a Juice_Production_Scion_Role for white grape juice production in the northeastern United States and Brazil, with its aromatic labrusca character, pale green-gold color, and balance of sweetness and acidity providing the white grape juice benchmark. Thompson Seedless bears a Juice_Production_Scion_Role for neutral white grape juice and juice concentrate production, where its high sugar content, mild flavor, and abundant availability support bulk juice manufacturing for blending and beverage applications.
A scion role that inheres in a grafted scion cultivar by virtue of its designation for grape juice production, where harvested grapes are crushed and processed into juice products for direct consumption or concentrate manufacture, rather than fermented into wine or dried into raisins. Juice production scions are selected for characteristics optimizing juice yield, flavor profile, and processing efficiency, including high juice content (70-85% berry weight extractable as juice), appropriate sugar levels for consumer palatability (typically 16-20°Brix, lower than wine or raisin grapes), balanced acidity providing refreshing character, distinctive varietal flavor (particularly the "foxy" or "grapey" character of Concord and other V. labrusca-influenced varieties prized in North American juice markets), and deep color for products where visual appeal is important. Grape juice production is concentrated in the United States (where Concord dominates), Brazil, and Argentina for direct-consumption juice, and in multiple countries for juice concentrate used in beverage blending. Global grape juice production exceeds 1.5 million tonnes annually.
Juice_Production_Scion_Role is subclassed by product type (Grape_Juice for direct consumption, Concentrate_Production for industrial processing), reflecting distinct cultivar requirements and market channels. The North American grape juice industry is uniquely dominated by V. labrusca and labrusca-influenced hybrids (Concord, Niagara), whose distinctive "foxy" aroma compounds (methyl anthranilate, o-aminoacetophenone) define consumer expectations in that market but are rejected in European wine culture. Juice production competes with wine production for some varieties (Thompson Seedless serves both), with market conditions determining allocation. Health-oriented marketing has expanded grape juice consumption globally, though concentrate production (for beverage blending) exceeds direct-consumption juice by volume.
Safttraube
Safttraubenrebsorte
Traubensaftsorte
grape juice variety
juice grape
juice variety
Morris JR, Striegler RK (2005) Grape juice: factors that influence quality, processing technology, and economics, in Processing Fruits: Science and Technology 2nd Ed, CRC Press pp.: 585-616 Creasy GL, Creasy LL (2009) Grapes, CABI Iyer MM et al. (2010) Determination of methyl anthranilate in grape juice, American Journal of Enology and Viticulture 61: 1-7 USDA National Agricultural Statistics Service (2020) Noncitrus Fruit and Nuts Summary
Juice Production Scion Role
Juice yield quality optimization in premium Champagne production employs the strictly regulated 102 liters per 160 kg marc (cuvée plus tailles combined), equivalent to approximately 64% yield, with the first 82 liters (cuvée, 51% yield) reserved for prestige cuvées and the subsequent 20 liters (tailles) showing higher phenolic content and coarser character houses targeting the highest quality accept even lower effective yields by using only cuvée juice, while the regulatory maximum prevents aggressive pressing that would extract excessive solids and compromise the finesse essential to fine Champagne.
Juice_Yield_Quality represents the chemical quality characterizing the volume or mass of extractable juice per unit of grape input, capturing the processing-relevant water content parameter that determines winemaking efficiency and affects economic returns. Juice yield integrates multiple berry characteristics including water content, cell wall composition, skin thickness, and pulp texture into a single metric expressing the fraction of grape mass recoverable as liquid must. Typical juice yields range from 65-80% of fresh grape weight, with variation reflecting cultivar (thick-skinned varieties yield less than thin-skinned), ripeness (overripe berries with degraded cell walls yield more), pressing technique (gentle whole-cluster pressing versus crushed fruit), and pressing pressure (higher pressure increases yield but extracts more solids and potentially harsh phenolics). The juice yield concept distinguishes free-run juice (draining without pressure, typically 55-65% of total) from press fractions obtained under increasing pressure, with quality generally declining as pressure increases due to extraction of skin and seed components. For white wine production, juice yield directly affects volume produced per ton of grapes and correlates with must clarity, with lower-yield gentle pressing producing cleaner juice requiring less settling or fining. For red wine production, the relevant consideration shifts to the proportion of juice relative to skins during maceration rather than total extractable yield. Economic pressures favor higher yields, while quality considerations often favor gentler extraction with lower yields, creating tension resolved differently depending on wine tier and style objectives. Juice yield measurement employs gravimetric or volumetric determination following standardized crushing and pressing protocols, with results expressed as liters per kilogram, gallons per ton, or percentage by weight.
Juice_Yield_Quality addresses the extractable liquid fraction as a processing-oriented water content parameter. While related to moisture content, juice yield also depends on tissue structure and extraction methodology, making it an empirical rather than purely compositional parameter. The distinction between free-run and press fractions reflects quality stratification within the extractable juice pool. Juice yield measurement requires standardized methodology for comparable results, as extraction technique profoundly affects yield. The economic significance of juice yield creates practical importance beyond purely qualitative considerations.
Extraktionseffizienz
Mostausbeute
Saftausbeute
Saftgewinnungsqualität
extraction efficiency
juice recovery
must yield
pressable liquid quality
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1: The Microbiology of Wine and Vinifications, 2nd Ed, John Wiley and Sons pp.: 327-395 Moio L and Piombino P (2009) Management of Vinification and Stabilization to Preserve and Enhance Wine Aroma. In: Moreno-Arribas MV and Polo MC (eds) Wine Chemistry and Biochemistry, Springer pp.: 671-695
Juice Yield Quality
The juiciness description of Pinot Noir documents high juiciness — gentle finger compression releases abundant free-run juice that flows readily from the ruptured skin, with the soft flesh disintegrating into liquid with minimal residual solid tissue, yielding 70-75% free-run juice by volume of total berry weight during winemaking crushing. This high juice release reflects Pinot Noir's thin skin, soft flesh with weak intercellular adhesion, and the high proportion of berry volume occupied by large, thin-walled, fully turgid mesocarp cells that rupture and release their vacuolar contents upon minimal mechanical stress. The high juiciness of Pinot Noir contrasts with the moderate juiciness of Cabernet Sauvignon (where the thick skin and firmer flesh resist compression and reduce free-run yield to 60-65%), the low juiciness of modern crisp table grapes like Autumn Royal (where the firm flesh maintains structural integrity rather than releasing juice), and the very high juiciness of Chasselas (where the extremely soft flesh produces near-complete dissolution upon compression).
A berry flesh description component that records the volume and ease of juice release when the berry is compressed or bitten, reflecting the proportion of cell contents that are free to flow as liquid (free-run juice) versus the proportion that remains bound within intact cells or associated with cell wall material, with juiciness influencing both the eating quality of table grapes (where moderate juiciness enhances the refreshing mouthfeel) and the winemaking efficiency (where high juiciness increases free-run juice yield and reduces pressing requirements).
Juiciness_Description captures a sensory property that integrates multiple physical characteristics of the berry flesh — cell turgor pressure (providing the driving force for juice expulsion), cell wall fragility (determining the ease of cell rupture under mechanical stress), intercellular adhesion strength (determining whether compression causes cell rupture with juice release or cell separation without rupture), and the ratio of vacuolar liquid to cell wall and cytoplasmic solid material (determining the maximum potential juice yield per unit of flesh weight). These properties vary independently across cultivars and development stages — a berry can be firm but juicy (high turgor in fragile cells, as in some Muscat types), firm and dry (high turgor in rigid cells, as in crisp table grapes), soft and juicy (low structural resistance with abundant free water, as in over-ripe wine grapes), or soft and mealy (low turgor with high cell wall content, as in dehydrated berries): the juiciness descriptor captures primarily the amount and ease of juice release rather than the underlying physical mechanisms, providing a practical assessment relevant to both eating quality and processing behavior. For winemaking, juiciness correlates with the free-run juice fraction that is recovered without pressing — the highest-quality juice component in white winemaking — and cultivars with high juiciness (Pinot Noir, Chasselas, Sémillon) produce more free-run juice per ton of fruit than cultivars with lower juiciness (Cabernet Sauvignon, Tannat), affecting winery throughput and wine style decisions.
Beerensaftigkeit-Charakterisierung
Saftabgabe-Bewertung
Saftigkeit-Beschreibung
berry succulence record
flesh juiciness characterization
juice release assessment
Juiciness Description
Cabernet Sauvignon juicy inner mesocarp contributing to berry juice yield= inner mesocarp water content 79-83% (similar to middle mesocarp 77-81%), vacuoles 89-92% of cell volume containing aqueous solution with 23-26% sugars and 0.6-0.8% acids, releasing liquid contents during pressing contributing proportionally to total juice yield (inner mesocarp 5-6% of pericarp volume contributing 5-6% of total juice volume), with juice composition from inner mesocarp essentially identical to middle mesocarp juice though inner mesocarp juice extracting later during pressing due to smaller cells requiring higher pressures for rupture. Thompson Seedless seedless table grape juicy inner mesocarp eating quality= water content 81-84%, vacuoles 90-93% of cell volume, cells readily rupturing during chewing releasing sweet juice (19-22% sugars) creating perception of juiciness throughout berry including near seed trace regions (stenospermocarpic arrested seeds), with juicy inner mesocarp ensuring pleasant juicy eating experience even when biting includes seed trace positions important for seedless table grape quality. Merlot juicy inner mesocarp ripening water accumulation= veraison (BBCH 81, week 10) showing water content 72-76%, vacuoles 85-88% of cell volume, moderate juiciness mid-ripening (week 14) with 76-80% water, 88-91% vacuoles, increasing juiciness maturity (week 18) with 79-83% water, 89-92% vacuoles, maximum juiciness, demonstrating progressive water accumulation and vacuole expansion during ripening creating increasingly juicy inner mesocarp texture as berry matures. Chardonnay juicy inner mesocarp juice release during white wine pressing= gentle pressing (0.5-1 bar pressure) releasing primarily middle mesocarp juice with minimal inner mesocarp extraction moderate pressing (1-2 bar) beginning to rupture inner mesocarp cells releasing inner mesocarp juice contributing to total must volume hard pressing (>2 bar) fully disrupting inner mesocarp releasing contents including some tannins (2.8-4.5 mg/g inner mesocarp tannins potentially extracting under hard press conditions) though typically white wine pressing uses gentle-to-moderate pressures avoiding excessive inner mesocarp phenolic extraction. Syrah juicy inner mesocarp versus seed moisture contrast= juicy inner mesocarp cells with 80-84% water creating very wet tissue adjacent mature seeds with 18-24% water creating relatively dry structure sharp moisture gradient over 20-100 μm distance from wet inner mesocarp across locular air space to dry seed surface, with contrast creating sensory experience during eating where juicy flesh and dry crunchy seeds provide textural variety, though during winemaking both tissues extracted together during maceration creating blend of juicy mesocarp and seed-derived components in wine.
A juicy inner mesocarp is an inner mesocarp emphasizing high juice content and ready juice release upon cell disruption, characterized by high water content (78-84%), large vacuoles (88-93% of cell volume) filled with aqueous solution of sugars and acids, thin-to-moderate cell walls (0.8-1.3 μm) that readily rupture during compression or chewing, and cells that release liquid cellular contents creating perception of juiciness when inner mesocarp tissue disrupted during eating or processing. Juicy inner mesocarp juice characteristics= (1) high water content 78-84% providing substantial liquid volume per unit tissue (2) aqueous vacuolar solution containing dissolved solutes (18-26% sugars, 0.5-0.9% organic acids, minerals) creating juice composition similar to middle mesocarp juice (3) ready release from cells when walls disrupted through chewing, crushing, or pressing (4) juice contributing to total berry juice yield (inner mesocarp representing 4-7% of pericarp volume contributing proportionally to total juice volume). Juicy inner mesocarp cellular basis of juiciness= (1) large vacuoles occupying 88-93% of cell volume maximizing liquid storage per cell (2) high vacuolar water content creating dilute aqueous solution readily flowing from cells when disrupted (3) walls thin enough (0.8-1.3 μm) to rupture readily under moderate compression releasing contents (4) minimal binding of water to cellular structures allowing free water to flow from disrupted cells (5) high sugar content (19-26%) in vacuolar solution creating sweet juice flavor. Juicy inner mesocarp in berry eating quality= (1) perception of juiciness when biting berry as cells rupture releasing liquid contents (2) contribution to overall berry juice release during eating creating pleasant juicy sensation (3) sweet flavor from released juice (high sugar content) important for eating pleasure (4) particular importance for table grapes where juiciness throughout berry including inner regions near seeds desired for eating quality. Juicy inner mesocarp in juice and wine production= (1) contribution to total juice yield during pressing with inner mesocarp cells rupturing and releasing contents (2) juice composition (sugars, acids, minerals) similar to middle mesocarp juice contributing proportionally to must composition (3) later juice release during pressing compared to readily-releasing middle mesocarp due to smaller cells and thicker walls requiring higher press pressures (4) potential contribution of inner mesocarp tannins (3-7 mg/g) to juice or must if pressing conditions vigorous enough to extract phenolics from inner mesocarp tissues. Juicy inner mesocarp developmental changes affecting juiciness= early development (Phase I) showing lower water content (65-75%) and smaller vacuoles creating less juicy texture ripening (Phase II) showing progressive water accumulation increasing water content to 78-84%, vacuole expansion to 88-93% of cell volume, creating increasingly juicy texture as ripening proceeds over-ripening potentially showing water loss through transpiration reducing juiciness and creating dehydrated raisined texture. Juicy inner mesocarp versus dry seed contrast= juicy inner mesocarp with 78-84% water creating wet fluid-releasing tissue adjacent seeds with 15-25% water at maturity creating dry firm structure sharp contrast between juicy flesh and dry seeds separated by locular air spaces in mature berries contrast important for eating experience where biting includes both juicy flesh and dry crunchy seeds creating textural variety.
Juicy_Inner_Mesocarp emphasizes juice content and release characteristics of inner mesocarp and is distinguished from Soft_Inner_Mesocarp which emphasizes textural firmness. Juicy inner mesocarp represents the inner mesocarp from the perspective of juice yield and eating juiciness perception. Understanding juicy inner mesocarp is relevant for table grape quality assessment (desirable juiciness), for juice yield prediction (inner mesocarp contributing proportionally to total yield), and for pressing protocol design where inner mesocarp requires higher pressures for juice extraction than readily-releasing middle mesocarp affecting free-run versus press juice composition.
Saftreiches Samengewebe
juicy inner flesh
liquid-rich seed zone
saftiges inneres Mesokarp
saftreiches inneres Fruchtfleisch
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 180-190 Coombe, B.G., McCarthy, M.G. (2000). Dynamics of grape berry growth. Australian Journal of Grape and Wine Research 6=131-135.
Juicy Inner Mesocarp
A juvenile unbranched tendril at node 11 on an actively extending shoot of Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, California, measuring 6.5 cm current length with visible active growth at apex producing new tissue at approximately 8-10 mm per day, showing entirely green coloration without any lignification, 1.8 mm diameter reflecting incomplete structural development, thin epidermis lacking glossy cuticle evident from dull surface appearance, immature vascular bundles visible through translucent epidermis showing primarily primary xylem without secondary thickening, minimal mechanosensory function demonstrated by absent or very weak response to experimental contact stimulation, occurring on a shoot 8 cm behind the apical meristem where tendril is approximately 5 days post-emergence, measured at intended BBCH stage 71 (fruit set) for mature shoot regions though this distal tendril is still in early development, representing the universal juvenile unbranched stage through which all tendrils pass before either remaining unbranched or developing branches over subsequent 7-14 days in this cultivar where 85-90% of tendrils ultimately develop bifurcated architecture.
An unbranched tendril that has not yet completed development and achieved functional maturity in Vitis vinifera, characterized by incomplete elongation with current length substantially below cultivar-typical mature size typically 3-10 cm compared to mature 10-22 cm, ongoing active growth with visible apex extension and internode production indicating continued developmental processes, incomplete tissue differentiation with immature vascular elements showing limited secondary wall development, thin epidermis with partial or absent cuticle formation, underdeveloped or absent mechanosensory apparatus with limited contact-responsive capacity, insufficient structural reinforcement with minimal lignification leaving tissues entirely green and highly flexible, and representing a transitional developmental stage that may either continue to mature as unbranched architecture or potentially initiate branching through delayed lateral meristem activation depending on cultivar genetics and resource availability during subsequent development. Juvenile unbranched tendrils occur universally as a developmental stage in tendril ontogeny, with all tendrils passing through unbranched juvenile phases before either remaining unbranched at maturity or activating branching programs, making distinction between determined unbranched development (will remain single-axis) versus facultative unbranched status (may branch with continued growth) important for functional classification and requiring knowledge of cultivar branching patterns and developmental timing to interpret individual juvenile tendril fate.
Juvenile unbranched tendrils are distinguished from mature simple unbranched tendrils by incomplete development with ongoing growth and immature tissue status, and represent a universal transitional stage rather than final architecture. Classification as juvenile requires assessment of developmental status including current length relative to mature size, tissue differentiation completion, lignification status, and mechanosensory function, with juveniles showing substantial deficits across multiple parameters. Prediction of final architecture (remain unbranched versus develop branches) requires knowledge of cultivar-specific patterns and developmental timing, as juvenile unbranched tendrils in strongly branching cultivars will likely develop branches while those in predominantly unbranched cultivars will likely mature as simple structures. Development rate and final architecture are influenced by shoot vigor, resource availability, and environmental conditions during the critical branching decision period typically 7-21 days post-emergence. Functionally, juvenile tendrils provide no climbing support and represent developmental investment without current functional return.
developing simple tendril
immature unbranched tendril
juvenile einfache Ranke
pre-mature single-axis tendril
unreife unverzweigte Ranke
vor-ausgereifte einachsige Kletterorgan
Based on Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 43-85 Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 195-218.
Juvenile Unbranched Tendril
K/Ca/Mg ratio quality in Yarra Valley Pinot Noir illustrates rootstock effects on mineral balance, with 101-14 rootstock producing must K/(Ca+Mg) ratios of 2.0-2.8 (milliequivalent basis) and harvest pH of 3.35-3.55, compared to SO4 rootstock on adjacent vines yielding ratios of 3.5-4.5 and pH of 3.55-3.80 this rootstock-mediated mineral balance difference translates directly to wine style potential, with lower-ratio fruit producing the bright, fresh-fruited Pinot Noir valued for regional typicity while higher-ratio fruit requires acid adjustment to achieve similar freshness.
K_Ca_Mg_Ratio_Quality represents the chemical quality characterizing the proportional relationships among potassium, calcium, and magnesium in grape tissue and must, capturing the cation balance that affects vine nutrition, must pH, and wine stability. The K/Ca/Mg ratio integrates the three major mineral cations that interact competitively during uptake and distinctively affect wine chemistry. Potassium, as a monovalent cation with high phloem mobility, dominates berry mineral content and produces strong pH effects through bitartrate formation. Calcium and magnesium, as divalent cations with different transport characteristics (calcium via xylem, magnesium via both pathways), moderate potassium's pH impact and contribute to tartrate stability through calcium tartrate formation. The K/(Ca+Mg) ratio, expressing potassium relative to the sum of divalent cations, provides a practical indicator of cation balance, with values below 2.5-3.0 (on a milliequivalent basis) associated with moderate pH and good balance, while values above 4.0 indicate potassium dominance with elevated pH risk. Soil K/Ca/Mg ratios influence but do not directly determine must ratios, as rootstock genotype strongly affects selective uptake of each element. High-potassium rootstocks (SO4, 5BB) produce elevated must K/(Ca+Mg) ratios and higher pH than low-potassium rootstocks (101-14, Riparia Gloire) on identical soils. Viticultural management through rootstock selection, cover crop competition (grasses compete effectively for potassium), and balanced fertilization provides tools for optimizing K/Ca/Mg ratios. The ratio concept enables quality prediction from mineral composition, linking terroir (soil) through viticulture (rootstock, management) to wine chemistry (pH, stability).
K_Ca_Mg_Ratio_Quality specifically addresses the three-way cation balance among the major mineral elements affecting vine nutrition and must pH. The ratio can be expressed various ways (K/Ca, K/Mg, K/(Ca+Mg), or ternary proportions) the K/(Ca+Mg) formulation on milliequivalent basis provides the most useful single metric. The rootstock effect on this ratio is substantial (30-50% variation among rootstocks on identical soil), making rootstock selection a primary tool for ratio management. The correlation between K/(Ca+Mg) ratio and must pH, while consistent, involves complex chemistry beyond simple stoichiometry.
Hauptkationenverhältnis
K-Ca-Mg-Verhältnis
K/Ca+Mg balance
Kalium-Calcium-Magnesium-Verhältnis
Makrokationenproportionsqualität
macrocation proportion quality
major cation ratio quality
potassium-calcium-magnesium ratio
Mpelasoka BS et al. (2003) A Review of Potassium Nutrition in Grapevines with Special Emphasis on Berry Accumulation. Australian Journal of Grape and Wine Research 9(3): 154-168 Ruhl EH (1989) Uptake and Distribution of Potassium by Grapevine Rootstocks and Its Implication for Grape Juice pH of Scion Varieties. Australian Journal of Experimental Agriculture 29(5): 707-712
K Ca Mg Ratio Quality
Kaempferol content quality analysis in Grüner Veltliner from the Wachau reveals elevated kaempferol proportions (25-40% of total flavonols) compared to typical red varieties (10-20%), reflecting the white cultivar's flavonol profile without the F3'5'H activity that would direct flux toward myricetin this kaempferol-rich profile, combined with quercetin glycosides, provides the co-pigmentation potential that contributes to the golden-green hue development during extended lees aging characteristic of Smaragd-level Grüner Veltliner.
Kaempferol_Content_Quality represents the chemical quality characterizing the concentration of kaempferol and its glycosides in grape berry skins, capturing the monohydroxylated flavonol that typically constitutes the second most abundant flavonol after quercetin in Vitis vinifera cultivars. Kaempferol (3,4',5,7-tetrahydroxyflavone) bears a single hydroxyl group at the 4' position of the B ring, representing the simplest hydroxylation pattern among grape flavonols and the structural analog of pelargonidin among anthocyanidins (though pelargonidin-based anthocyanins are essentially absent from V. vinifera grapes). Kaempferol glycosides typically comprise 10-30% of total flavonols in grape skins, with kaempferol-3-glucoside and kaempferol-3-glucuronide as the primary forms. Like quercetin, kaempferol accumulation is strongly light-dependent, with sun-exposed berries showing elevated concentrations compared to shaded fruit. The kaempferol-to-quercetin ratio provides information about flavonoid 3'-hydroxylase (F3'H) activity, as F3'H converts kaempferol precursors to quercetin precursors high kaempferol relative to quercetin suggests limited F3'H activity. Some white grape cultivars show particularly high kaempferol proportions among flavonols, reflecting reduced F3'H flux in the absence of anthocyanin synthesis. Kaempferol contributes to co-pigmentation similarly to other flavonols, supporting anthocyanin color in red wines. The yellow coloration of kaempferol glycosides may contribute to the color of white wines, though concentrations are typically too low for significant visual impact.
Kaempferol_Content_Quality addresses the monohydroxylated B-ring flavonol representing the product of flavonol synthase acting on dihydrokaempferol before F3'H hydroxylation. The kaempferol proportion relative to quercetin and myricetin reflects the balance among branch pathways in flavonoid biosynthesis. White grape cultivars often show elevated kaempferol proportions because the pathway flux toward trihydroxylated products is reduced in the absence of anthocyanin synthesis. Kaempferol glycosides are quantified by HPLC with UV detection at 360 nm, providing cultivar-specific profile data useful for authenticity verification.
K-3-glc content
Kaempferol-3-glucosid-Konzentration
Kaempferolgehalt
Kaempferolglykosidgehalt
Monohydroxyflavonolgehalt
kaempferol glycoside content
kaempferol-3-glucoside concentration
monohydroxyflavonol quality
Castillo-Muñoz N et al. (2007) Flavonol Profiles of Vitis vinifera Red Grapes and Their Single-Cultivar Wines. Journal of Agricultural and Food Chemistry 55(3): 992-1002 Mattivi F et al. (2006) Metabolite Profiling of Grape: Flavonols and Anthocyanins. Journal of Agricultural and Food Chemistry 54(20): 7692-7702
Kaempferol Content Quality
KMW measurement for Grüner Veltliner from the Kamptal (2024 vintage) records harvest at 18.5° KMW, comfortably exceeding the Austrian Qualitätswein minimum of 15° KMW and approaching the Kabinett threshold of 17° KMW (already surpassed), providing the basis for either dry Qualitätswein or Kabinett classification depending on residual sugar and sensory assessment. KMW measurement for Welschriesling from Neusiedlersee documents the progressive selection for Prädikatswein categories: Spätlese lot at 19° KMW (minimum requirement), Auslese lot at 21° KMW (minimum requirement), Beerenauslese lot at 25° KMW (minimum requirement), and individual Trockenbeerenauslese berries at 30° KMW (minimum requirement) — the KMW thresholds defined in § 13 of the Austrian Weingesetz 2009 determining each lot's regulatory classification and market value.
KMW measurement is a sugar content data entity that records the sugar concentration in grape must as determined by refractometry or densimetry on the Klosterneuburger Mostwaage (KMW) scale, expressed in degrees KMW (°KMW), where one degree KMW represents one gram of sugar per 100 grams of must (equivalent to sugar percentage by weight), serving as the legally mandated and commercially standard must weight measurement system in Austrian viticulture and providing the regulatory basis for the Austrian wine quality classification system defined in the Weingesetz 2009.
The KMW scale was developed by August Wilhelm von Babo at the Höhere Bundeslehranstalt für Wein- und Obstbau in Klosterneuburg in 1861, making it one of the oldest continuously used viticultural measurement systems. The scale's definition as sugar percentage by weight makes it conceptually distinct from density-based scales (Oechsle, Baumé) — KMW measures the actual sugar fraction rather than total must density, though in practice the measurement is performed through refractometry or densimetry with calibration tables converting to the KMW sugar content scale. The conversion relationships to other scales are approximately 1° KMW ≈ 5° Oechsle ≈ 1.25° Brix ≈ 0.7° Baumé, though these approximations vary with must composition. The Austrian Weingesetz 2009 defines all quality categories through KMW minimum thresholds: Qualitätswein 15° KMW, Kabinett 17° KMW, Spätlese 19° KMW, Auslese 21° KMW, Beerenauslese/Eiswein 25° KMW, Ausbruch 27° KMW, and Trockenbeerenauslese 30° KMW — making KMW measurement the single most legally consequential analytical determination in Austrian viticulture. The scale is also historically used in parts of the Czech Republic, Slovakia, and Hungary, reflecting the legacy of the Austro-Hungarian viticultural tradition, though these countries have increasingly adopted parallel reporting in Oechsle or Brix for international communication. The Klosterneuburg institution where the scale was developed continues to play a central role in Austrian viticultural research, clonal selection (as documented in the VVO Clonal_Selection_Record hierarchy), and analytical method standardization, maintaining a direct institutional link between the KMW scale and the Austrian wine quality assurance system.
KMW degree
KMW must weight
KMW reading
KMW value
KMW-Bestimmung
KMW-Grad
KMW-Messung
KMW-Mostgewicht
Klosterneuburger Mostwaage
Klosterneuburger Mostwaage-Wert
Redl H et al. (2007) Der Wein: Weinbau, Kellerwirtschaft, Weinrecht, 2nd Ed, Österreichischer Agrarverlag, pp.: 156-175 Austrian Weingesetz 2009 (BGBl. I Nr. 111/2009), § 13 Prädikatswein Troost G (1988) Technologie des Weines, Ulmer Verlag, pp.: 78-85
KMW Measurement
KMW quality monitoring at Weingut Knoll in the Wachau tracks the Smaragd classification threshold of 18.2° KMW (approximately 24.5°Brix) that distinguishes the region's fullest-bodied dry wines from the lighter Federspiel category (17-18.2° KMW), with Grüner Veltliner from the premier Kellerberg vineyard regularly achieving 19-21° KMW in warm vintages the historical connection to nearby Klosterneuburg, visible across the Danube, reinforces the cultural significance of this measurement system that has defined Austrian wine quality classification for over 125 years.
KMW_Quality_(Klosterneuburger_Mostwaage) represents the chemical quality characterizing grape must sugar concentration according to the Austrian must weight scale developed at the Höhere Bundeslehranstalt und Bundesamt für Wein- und Obstbau in Klosterneuburg, Austria. The KMW scale, officially adopted in Austria in 1896, expresses sugar content as the percentage by weight of sugar in grape must, directly indicating grams of sugar per 100 grams of must. This straightforward relationship to actual sugar concentration distinguishes KMW from density-based scales like Oechsle, making it conceptually similar to Brix but with historical and regulatory significance in Austrian viticulture. Typical harvest values range from 15-25° KMW for quality wines, with Austrian wine law establishing minimum KMW requirements for quality categories including Qualitätswein (15° KMW minimum), Kabinett (17° KMW), Spätlese (19° KMW), Auslese (21° KMW), and progressively higher requirements for Beerenauslese, Ausbruch, and Trockenbeerenauslese. The relationship between KMW and potential alcohol follows the approximate conversion of °KMW × 0.66 = potential alcohol (% v/v). KMW measurement employs calibrated KMW hydrometers or calculation from density measurements using established conversion tables. The scale honors the internationally recognized enological research and education institution at Klosterneuburg, founded in 1860 as the world's oldest wine school, whose contributions to viticulture science extend far beyond the must weight scale bearing its name. Within Austria, KMW values are universally understood by vineyard workers, winemakers, and wine consumers as the primary indicator of grape ripeness and wine quality potential.
KMW_Quality_(Klosterneuburger_Mostwaage) is positioned as a subclass of Must_Weight_Quality representing the Austrian tradition. The parenthetical Klosterneuburger_Mostwaage follows the class naming convention in the uploaded structure. KMW is mathematically related to other scales: °KMW ≈ °Brix × 1.0 (nearly equivalent), °KMW ≈ °Oe/4.86. The unique aspect of KMW is its direct expression as weight percentage sugar, making it more intuitive for understanding actual sugar content than density-based scales. The historical association with the Klosterneuburg wine school, where Bert resided for five years, adds cultural depth to this technical measurement.
Austrian must scale
Grad KMW
KMW degrees
Klosterneuburg must weight
degrees KMW
österreichisches Mostgewicht
Austrian Wine Law (Weingesetz) and associated regulations Redl H et al. (1996) Weinbau, 7th Ed, Österreichischer Agrarverlag pp.: 324-356 Bauer K (2002) Weinbau, 8th Ed, Österreichischer Agrarverlag pp.: 215-240
KMW Quality (Klosterneuburger Mostwaage)
Label regulation for Austrian wine integrates multiple regulatory layers: EU Regulation 2019/33 establishing the framework for mandatory particulars (actual alcoholic strength ±0.5% vol., nominal volume, lot number, allergen declaration "contains sulfites" if total SO₂ >10 mg/L, producer name and address, country of origin), the Austrian Weingesetz 2009 specifying national requirements (quality tier designation — Qualitätswein, Kabinett, or Prädikatswein, state control number/Prüfnummer for quality wines, wine-growing region), and DAC-specific regulations (Wachau DAC requiring Vinea Wachau classification — Steinfeder, Federspiel, or Smaragd — where applicable, Weinviertel DAC mandating cultivar identification as Grüner Veltliner). Label regulation for US wine applies TTB (Alcohol and Tobacco Tax and Trade Bureau) requirements — mandatory elements include brand name, class/type designation, alcohol content (±1.0% for 7-14% vol., ±1.5% above 14%), net contents, name and address of bottler, appellation of origin, and the "Government Warning" health statement required since 1989.
Label regulation is a regulatory information entity that documents the legal requirements and restrictions governing the information that may, must, or must not appear on wine labels, encompassing varietal labeling rules (the conditions under which a cultivar name may be displayed), appellation requirements (the conditions for using geographic origin designations), vintage declaration rules (the conditions for declaring the harvest year), and additional mandatory and optional label elements (alcohol content, volume, lot number, allergen declarations, organic certification, producer identification), as established by international conventions (OIV labeling recommendations), supranational regulations (EU wine labeling Regulation 2019/33), national wine laws, and appellation-specific specifications, providing the consumer information framework that mediates between the complex viticultural and oenological reality of wine and the information available to the purchaser at the point of sale.
Label regulation documentation addresses the regulatory mediation between wine's inherent complexity and the consumer's need for reliable, comprehensible product information — a mediation that balances multiple competing objectives including consumer protection (ensuring label claims are truthful and verifiable), producer flexibility (permitting marketing-relevant optional information), cultural heritage preservation (protecting established geographic and varietal designations from misuse), and trade facilitation (harmonizing labeling standards across jurisdictions). The EU wine labeling framework distinguishes between mandatory particulars (information required by law on every wine label), optional particulars (information that may appear if certain conditions are met), and protected terms (designations whose use is restricted to wines meeting specific requirements) — a tripartite structure that creates both opportunities and constraints for wine marketing. The global patchwork of labeling regulations creates significant compliance challenges for internationally traded wines — a wine destined for multiple markets may require different labels for each destination, as the mandatory elements, formatting requirements, permitted claims, and language rules differ across the EU, US, Canada, Japan, China, and other major import markets. The allergen labeling dimension has expanded substantially since the EU mandated sulfite declarations (2005) and subsequently added requirements for milk-derived (casein) and egg-derived (albumin) fining agent declarations if residues exceed detection limits — requirements that have prompted significant winemaking changes (adoption of bentonite, pea protein, and PVPP fining as alternatives) to simplify label compliance. The digital dimension of wine labeling is emerging through the EU's mandatory e-label provision (Regulation 2021/2117, effective from December 2023 for new wines), requiring nutritional information and ingredient lists to be available through a QR code or URL on the physical label — a regulatory innovation that substantially expands the information accessible to consumers while accommodating the physical space constraints of traditional wine labels.
Etikettierungsbestimmungen
Etikettierungsrecht
Etikettierungsvorschriften
Kennzeichnungsvorschriften
Weinetikettierungsregeln
label compliance
label law
labeling requirements
wine label regulation
wine labeling rules
European Parliament (2019) Commission Delegated Regulation (EU) 2019/33 supplementing Regulation (EU) No 1308/2013 as regards labeling Österreichisches Weingesetz (2009) §§29-36 (Bezeichnungsrecht) TTB (2023) Beverage Alcohol Manual, US Alcohol and Tobacco Tax and Trade Bureau OIV (2022) International Standard for the Labelling of Wines
Label Regulation
Lactic acid content quality in Napa Valley Merlot provides the signature soft, velvety acidity achieved through complete malolactic fermentation, with conversion of 3-5 g/L malic acid to 2-3 g/L lactic acid during barrel aging winemakers monitor lactic acid accumulation via paper chromatography or enzymatic assay to confirm MLF completion before blending and bottling, as incomplete conversion leaves residual malic acid substrate that could support unwanted bacterial activity in bottle, while the stable lactic acid concentration in finished wine (typically 1.5-2.5 g/L) indicates successful transformation to the smooth, approachable style expected of premium Napa Merlot.
Lactic_Acid_Content_Quality represents the chemical quality characterizing the concentration of lactic acid (2-hydroxypropanoic acid, C₃H₆O₃) in grape berry tissue and wine. Lactic acid is distinguished from other grape acids by its predominantly fermentation origin rather than grape synthesis, being produced primarily through malolactic fermentation where lactic acid bacteria convert malic acid to lactic acid. Fresh grape berries contain minimal lactic acid (typically less than 0.1 g/L), but post-malolactic wines may contain 1-3 g/L lactic acid representing the transformation of grape malic acid. This bacterial conversion reduces total acidity because the dicarboxylic malic acid (two acid groups) is converted to monocarboxylic lactic acid (one acid group) plus carbon dioxide, with one acid equivalent lost. Lactic acid contributes soft, milky, slightly sour flavor notes that are less sharp than malic acid, producing the sensory impression of rounded, smooth acidity after malolactic fermentation. The L-(+)-lactic acid stereoisomer predominates from bacterial malolactic conversion, while racemic DL-lactic acid may be produced by certain bacteria or through chemical synthesis. Small amounts of lactic acid are also produced by yeasts during alcoholic fermentation. Elevated lactic acid in wines that have not undergone intentional malolactic fermentation may indicate spoilage by lactic acid bacteria, potentially accompanied by off-flavors and excessive volatile acidity. The lactic acid content therefore serves both as an indicator of malolactic fermentation completion and as a quality marker distinguishing intentional bacterial conversion from spoilage. Measurement employs specific enzymatic assays using lactate dehydrogenase or HPLC separation, with stereospecific methods available to distinguish L-lactic from D-lactic acid.
Lactic_Acid_Content_Quality primarily reflects malolactic fermentation rather than grape composition, making it unique among the Organic_Acid_Quality child classes. The transformation from malic to lactic acid is irreversible under normal conditions, making lactic acid a stable indicator of bacterial activity. The taste difference between malic acid (tart, green apple) and lactic acid (soft, milky) is perceptually larger than the chemical difference might suggest, explaining why malolactic fermentation produces significant sensory change. The potential for lactic acid bacteria to also produce acetic acid and diacetyl makes MLF management critical for wine quality beyond the simple acid conversion.
2-hydroxypropanoic acid concentration
Lactat
Milchsäure
Milchsäuregehalt
Milchsäurekonzentration
fermentation acid quality
lactate content
milk acid level
Liu SQ (2002) Malolactic Fermentation in Wine: Beyond Deacidification. Journal of Applied Microbiology 92(4): 589-601 Volschenk H et al. (2006) Malic Acid in Wine: Origin, Function and Metabolism during Vinification. South African Journal of Enology and Viticulture 27(2): 123-136
Lactic Acid Content Quality
Cabernet Sauvignon in Napa Valley shows lag phase exit during early-mid September Year N+1 at BBCH 83, with berry growth rate increasing from 3 to 18 mg per day over 2 days, water uptake accelerating from 0.09 to 0.52 g per berry per day, respiratory rate increasing from 1.3 to 1.9 μmol O₂ g⁻¹ h⁻¹, sugar accumulation accelerating to 2.2% per week, anthocyanin reaching 0.28 mg/g creating visible red color, firmness declining 24% from peak, berry entering active ripening state, under warm climate conditions of 24-30°C temperature, typical of this late-season cultivar exiting lag phase
Lag phase exit represents the definitive quiescence termination phase during lag phase termination when berry conclusively leaves lag state entering ripening phase, growth rate increases from lag minimal <5 mg per day to ripening moderate 10-30 mg per day, metabolic activity elevates substantially toward ripening levels, ripening processes fully activated including sugar accumulation and color development, and berry establishes ripening state replacing lag quiescence, occurring during final 1-2 days of lag as veraison proceeds. This quiescence exit phase occurs during late summer to early autumn in early-mid September in temperate regions Year N+1, proceeding over 1-2 days representing definitive lag-ripening transition, corresponding to BBCH 83 berries beginning to develop color or soften. Exit mechanism involves growth rate increase from water uptake acceleration, metabolic rate elevation supporting ripening biosynthesis, sugar accumulation acceleration from activated phloem import, color development initiation in red cultivars from anthocyanin synthesis, continued softening from cell wall modification, and comprehensive activation establishing ripening state. Growth rate increase shows berry fresh weight gain increasing from lag minimal 1-5 mg per day to ripening low-moderate 10-30 mg per day during exit, growth increase marking definitive lag termination, volumetric expansion from water uptake, and growth as lag exit indicator. Growth rate dynamics shows progressive increase over 1-2 days, eventual stabilization at ripening growth levels, continued growth through ripening phase, and growth pattern characteristic of phase transition. Water uptake acceleration shows water influx increasing from lag equilibrium <0.1 g per berry per day to ripening 0.3-0.8 g per day, osmotic gradient from sugar accumulation driving uptake, increased water enabling berry expansion, and water uptake as growth mechanism. Berry volumetric expansion shows berry volume increasing from water accumulation, typical 5-15% volume increase during early veraison over several days, expansion contributing to berry size increase, and volume expansion characteristic of ripening onset. Metabolic activity elevation shows respiratory rate increasing from lag 1.0-1.5 μmol O₂ g⁻¹ h⁻¹ to ripening 1.5-2.5 μmol O₂ g⁻¹ h⁻¹, metabolic rate increase supporting biosynthetic demands, ATP production increasing for ripening processes, and respiration as metabolic activity indicator. Respiratory increase shows enhanced oxidative phosphorylation, increased substrate oxidation particularly sugars and remaining acids, respiratory capacity supporting ripening metabolism, and respiration elevation marking metabolic shift. Sugar accumulation acceleration shows sugar import increasing from lag minimal 0.5-1.5% per week to ripening 1.5-3.5% per week during exit, phloem unloading accelerating from activated hexose transporters, vacuole hexose storage increasing, and sugar accumulation as defining ripening process. Sugar accumulation rate shows progressive acceleration during exit, typical sugar increase 1-2% over exit days, continued acceleration through early ripening, and accumulation pattern characteristic of ripening onset. Phloem unloading activation shows phloem sieve element unloading increasing substantially, hexose transporters VvHT1, VvHT2 at high activity from accumulated protein, symplastic and apoplastic unloading pathways active, and unloading capacity supporting sugar accumulation. Vacuole hexose storage shows glucose and fructose accumulation in vacuole, hexose concentrations increasing from 50-80 mM lag to 100-150 mM early exit, tonoplast hexose transporters VvHT6 and others mediating storage, and vacuole storage as sugar accumulation mechanism. Color development in red cultivars shows anthocyanin synthesis proceeding rapidly, anthocyanin accumulation 0.1-0.5 mg/g during exit creating visible pink to red color, color intensification continuing through ripening, and color as red cultivar transformation marker. Anthocyanin synthesis rate shows rapid synthesis during exit typical 0.05-0.15 mg/g per day, pathway flux high from accumulated enzyme capacity, color appearing and intensifying over exit period, and anthocyanin accumulation as dramatic biosynthetic event. Color appearance shows berry color transitioning from green or faint pink to clearly pink or light red, color typically starting at stylar end progressing toward pedicel, visible color change as exit indicator, and color development characteristic of red cultivars. Softening continuation shows firmness continuing decline from softening initiation, typical 15-30% total firmness decline from peak by exit completion, progressive texture softening through ripening, and softening as continuing transformation. Berry composition during exit shows water possibly increasing to 80-84% from uptake, organic acids declining 0.6-0.8% with continued respiration, sugars increasing to 14-18%, pH rising to 3.2-3.5, and composition rapidly changing toward ripening characteristics. Berry appearance shows berries clearly in transformation with visible color in red cultivars or obvious softening and possible yellowing in white cultivars, cluster showing mixed berry states from asynchrony, and appearance indicating ripening onset. Ripening phase entry shows berry definitively in ripening state, active sugar accumulation and color development in red cultivars, progressive softening and acid decline, and ripening as post-lag developmental phase. Ripening state characteristics shows growth moderate 10-30 mg per day, metabolic activity elevated 1.5-2.5 μmol O₂ g⁻¹ h⁻¹, sugar accumulation active 1.5-3.5% per week, and active ripening processes. Lag phase concluded shows lag quiescence definitively ended, minimal growth state replaced by moderate growth, lag metabolism replaced by ripening metabolism, and lag as completed developmental phase. Environmental effects on exit include temperature affecting exit timing and dynamics, warm conditions promoting faster exit transition, cool conditions extending exit period, and temperature determining ripening progression rate. The process represents third and final aspect of lag phase termination establishing ripening entry, following veraison approach and softening initiation. Duration 1-2 days representing definitive lag-ripening transition. Regional timing shows lag phase exit during early-mid September as ripening begins. Viticultural significance includes representing definitive lag termination and ripening onset, determining ripening phase commencement affecting harvest date, establishing berry in active transformation state, and marking developmental progression affecting quality and timing. Research applications include analyzing lag-ripening transition mechanisms and regulation, investigating growth and metabolic activation dynamics, evaluating sugar accumulation and color development onset, quantifying exit timing and indicators, and assessing lag phase exit as determinant of ripening progression and harvest timing.
Lag_Phase_Exit is third and final aspect of Lag_Phase_Termination, distinguished by definitive ripening entry. Represents growth rate 1-5 → 10-30 mg/day. Water uptake 0.3-0.8 g/berry/day. Respiratory rate 1.0-1.5 → 1.5-2.5 μmol O₂ g⁻¹ h⁻¹. Sugar accumulation 1.5-3.5%/week. Anthocyanin 0.1-0.5 mg/g in red cultivars. Firmness decline 15-30% total. Duration 1-2 days. Follows Veraison_Approach and Softening_Initiation. Completes Lag_Phase_Termination and Berry_Growth_Lag_Phase. Enters Ripening Phase.
Lag-Abschluss
Lag-Phasen-Exit
Reifungs-Phasen-Eintritt
Ruhephase-Terminierung
lag conclusion
quiescence termination
ripening phase entry
veraison completion
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 475-485 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Lag Phase Exit
Pinot Noir in Burgundy shows lag phase initiation during late August-early September Year N+1 at BBCH 81, with growth rate declining from 12 mg per day to 3 mg per day over 5-7 days establishing lag quiescence, metabolic transition onset with ripening genes beginning 2-4 fold low-level expression, respiratory rate declining from 1.6 to 1.2 μmol O₂ g⁻¹ h⁻¹, berry dimensions stable at 9.8-10.2 mm, under continental conditions of 20-26°C temperature, typical of this early-to-mid season cultivar entering lag phase
Lag phase initiation represents the quiescence establishment period at berry growth lag phase onset when growth rate declines from enlargement completion levels 10-30 mg per day to lag phase minimal rates <5 mg per day, metabolic transition shifts berry from growth-phase to quiescent pre-ripening metabolism, compositional changes continue from late enlargement toward lag characteristics, and berry establishes stable quiescent state preparing for eventual veraison, occurring over several days to 1 week as lag phase begins. This quiescence onset phase occurs during late summer to early autumn in late August-early September in temperate regions Year N+1, proceeding over days to 1 week at lag initiation, corresponding to BBCH 81 beginning of ripening stage transition. Initiation mechanism involves growth rate decline continuing from enlargement completion deceleration to lag minimal levels, metabolic transition onset with gene expression and enzyme activity shifts establishing pre-ripening program, compositional trajectory from late enlargement continued into lag with acids declining and sugars increasing, hormonal changes possibly regulating transition, and integrated changes establishing quiescent lag state. Growth rate decline characteristics show progression from enlargement completion 10-30 mg per day to transitional 5-15 mg per day early initiation to lag phase <5 mg per day by initiation completion, typical 70-85% rate reduction during initiation period, and decline establishing minimal growth defining lag phase. Growth rate decline kinetics show progressive deceleration over initiation period, exponential-like decline approaching asymptotic lag rate, eventual stabilization at minimal lag levels, and decline trajectory establishing quiescence. Growth cessation approach shows berry growth effectively ceasing during initiation, volumetric increase minimal <1% during initiation, berry entering stable dimensional state, and cessation as lag defining characteristic. Metabolic transition onset shows beginning of metabolic reorganization from growth to pre-ripening state, gene expression changes initiating during lag initiation, enzyme activity shifts beginning, and metabolic program transition establishing lag metabolism. Gene expression initiation shows growth genes continuing basal expression, lag-associated genes including stress response genes maintaining or increasing expression, early ripening gene expression beginning at very low levels, and expression patterns shifting toward lag and pre-ripening profiles. Stress response gene expression shows heat shock proteins, antioxidant enzymes maintained or upregulated, protective gene expression characteristic of lag phase, and stress preparedness as lag feature. Ripening gene early expression shows anthocyanin biosynthesis genes beginning very low expression in red cultivars, sugar transporter genes beginning increase, softening enzyme genes starting low-level expression, and early expression preparing for veraison activation. Enzyme activity changes show growth-related enzymes at basal levels, maintenance metabolism enzymes sustained, early ripening-associated enzymes beginning accumulation at low levels, and activity patterns characteristic of transition state. Respiratory metabolism during initiation shows respiration declining from completion levels 1.2-2.0 μmol O₂ g⁻¹ h⁻¹ to lag levels 1.0-1.5 μmol O₂ g⁻¹ h⁻¹, metabolic rate reduction reflecting transition to quiescence, low respiration establishing lag metabolism, and respiratory level as metabolic indicator. Carbon metabolism during initiation shows minimal biosynthetic activity, carbon primarily for maintenance, limited sugar import and storage, and metabolism characteristic of quiescent state. Organic acid metabolism shows continued malate respiration declining acids, malate declining from 0.7-1.0% to 0.6-0.9% during initiation, tartrate stable 0.3-0.6%, and acid dynamics continuing from enlargement. Sugar metabolism shows limited sugar accumulation during initiation, sugars increasing from 10-14% to 11-15%, slow accumulation rate 0.5-1.5% per week, and limited import characteristic of lag initiation. Berry composition during initiation shows water 78-82% stable, organic acids 0.8-1.2% declining slowly, sugars 11-15% increasing slowly, pH 3.0-3.2 rising gradually, and composition transitioning toward lag characteristics. Berry size during initiation shows dimensions remaining stable, diameter 6-23 mm depending on cultivar unchanging, fresh weight 2-28 g stable within 2-5%, and stability establishing lag constancy. Water relations during initiation shows water uptake declining from completion 0.1-0.3 g per berry per day to lag minimal <0.05-0.1 g per day, water balance approaching equilibrium, and minimal flux characteristic of quiescence. Berry morphology during initiation shows berry at stable mature size, green color maintained or beginning slight decline, texture firm, and appearance as hard green berry entering lag. Seed state during initiation shows seeds at advanced maturity or fully mature, seed coat dark brown and hardened, embryo viable, and seeds approaching completion enabling lag phase. Hormone dynamics during initiation shows auxin at low levels, possible gibberellin decline, abscisic acid remaining low before eventual veraison spike, and hormones characteristic of transition state. Environmental effects on initiation include temperature affecting initiation timing with warm conditions promoting faster transition, water availability affecting berry hydration, and environmental factors influencing transition dynamics. Temperature effects show warm conditions accelerating initiation and progression into lag, cool conditions extending initiation period, and temperature affecting transition kinetics. Initiation timing shows typical occurrence 1-2 weeks after enlargement completion, overlap with completion deceleration, and timing marking definitive lag phase entry. Initiation duration shows transition proceeding over days to 1 week, relatively rapid establishment of lag state, and brief initiation period relative to total lag duration. Initiation completion indicators show growth rate stabilized at <5 mg per day, metabolic markers showing lag characteristics, compositional trajectory established, and indicators documenting lag entry. The process shows two distinguishable aspects including growth rate decline start establishing minimal growth, and metabolic transition onset shifting to pre-ripening program. Duration typically days to 1 week representing lag initiation period. Regional timing shows lag phase initiation during late August-early September across climates as quiescence begins. Viticultural significance includes representing definitive transition to lag phase quiescence, establishing minimal growth state, beginning metabolic preparation for ripening, and marking lag phase entry affecting subsequent veraison timing. Research applications include analyzing initiation mechanisms and regulation, investigating metabolic transition onset, evaluating environmental effects on initiation timing, quantifying growth and metabolic dynamics, and assessing lag initiation as determinant of lag phase characteristics and duration.
Lag_Phase_Initiation represents quiescence establishment within Berry_Growth_Lag_Phase. Subdivided into Growth_Rate_Decline_Start with 70-85% reduction to <5 mg/day, and Metabolic_Transition_Onset establishing pre-ripening program. Duration days to 1 week. Respiratory rate declining to 1.0-1.5 μmol O₂ g⁻¹ h⁻¹. Growth genes basal, lag genes sustained, ripening genes beginning very low expression. Berry dimensions stable, composition transitioning. Precedes Minimal_Growth_Period. Marks definitive lag phase entry.
Lag-Beginn
Lag-Phasen-Initiierung
Ruhephase-Etablierung
Wachstums-Beendigungs-Initiierung
growth cessation initiation
lag beginning
lag onset
quiescence establishment
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 453-458 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Lag Phase Initiation
Syrah in Rhône Valley shows lag phase termination during early-mid September Year N+1 at BBCH 81-83, with earliest berries showing pink color from 0.08-0.15 mg/g anthocyanin accumulation at stylar end, firmness declining from 3.2 to 2.5 N representing 22% softening, growth rate increasing from 3 to 12 mg per day, water uptake increasing from 0.08 to 0.35 g per berry per day, abscisic acid peaking at 1240 ng/g, veraison beginning with 15-20% berries showing transformation signs, under warm Mediterranean conditions of 24-29°C temperature, typical of this mid-to-late season cultivar exiting lag phase
Lag phase termination represents the lag phase conclusion period during berry growth lag phase when veraison approaches with visible and measurable transformation signs, softening initiates with first detectable texture reduction, growth rate possibly increases from lag minimal levels, abscisic acid spike peaks triggering or accompanying transformation, and berry exits lag quiescence entering ripening phase, occurring over final 1-3 days of lag phase as veraison begins. This lag conclusion transition period occurs during late summer to early autumn in early-mid September in temperate regions Year N+1, proceeding over 1-3 days representing lag-veraison transition, corresponding to BBCH 81-83 transition from beginning of ripening to berries beginning to develop color or soften. Termination mechanism involves veraison approach with transformation becoming imminent or initiated, softening initiation with first measurable firmness decline, possible growth rate increase from water uptake acceleration, abscisic acid peak levels triggering or correlating with transformation, lag phase exit with berry entering ripening state, and integrated changes marking definitive lag phase end. Veraison approach characteristics shows veraison onset imminent within 0-3 days, earliest berries possibly showing initial color in red cultivars or clear softening in white cultivars, berry at transformation threshold or beginning transformation, and approach marking lag termination. Earliest transformation signs shows trace to detectable color development in red cultivars with anthocyanin accumulation 0.05-0.2 mg/g creating visible pink to light red tinge, measurable softening with firmness declining 10-25% from peak, possible berry volume increase from water uptake, and transformation signs as termination indicators. Color development in red cultivars shows anthocyanin synthesis accelerating in earliest berries, visible color change from green to pink or light red in berry sectors typically stylar end first, color as defining veraison onset marker in red cultivars, and color development marking transformation beginning. Anthocyanin accumulation dynamics shows rapid anthocyanin synthesis initiation with accumulation rate 0.05-0.15 mg/g per day during early veraison, pathway flux high from accumulated enzyme capacity, anthocyanin synthesis as dramatic metabolic shift, and accumulation creating visible transformation. Softening initiation shows first detectable texture change with berry firmness beginning decline, firmness reduction 10-25% from pre-veraison peak during termination period, cell wall modification initiating through pectin degradation and depolymerization, and softening as universal veraison marker across cultivars. Firmness decline mechanisms shows polygalacturonase activity increasing degrading pectin, pectin methylesterase modifying pectin structure, pectin depolymerization reducing cell wall integrity and firmness, calcium release from pectin affecting structure, and enzymatic modification as softening mechanism. Cell wall pectin changes shows methylesterified pectin declining from esterase activity, demethylated pectin increasing, calcium cross-links possibly forming then breaking, pectin molecular weight declining from depolymerization, and pectin changes creating softening. Berry texture measurement shows penetrometer firmness declining from 2.5-4.5 N pre-veraison to 2.0-3.5 N early termination, compression resistance decreasing, texture subjectively softer, and firmness as quantitative softening indicator. Growth rate possible increase shows berry fresh weight gain possibly increasing from lag minimal 1-5 mg per day to 5-20 mg per day during termination from water uptake acceleration, growth rate increase marking lag exit in some berries, volumetric increase from water influx, and growth as possible termination indicator. Water uptake acceleration shows water influx increasing from lag equilibrium <0.1 g per berry per day to 0.2-0.5 g per day during termination, increased uptake from osmotic gradient from sugar accumulation, water driving berry expansion during early veraison, and uptake increase as mechanism for growth rate increase. Sugar accumulation possible acceleration shows sugar import possibly beginning increase from lag 0.5-1.5% per week to early veraison 1.0-2.5% per week, sugar accumulation creating osmotic gradient, phloem import increasing from activated transporter capacity, and sugar dynamics shifting toward ripening. Abscisic acid spike peak shows ABA reaching maximum levels 500-2000 ng/g during lag termination and early veraison, ABA peak coinciding with or slightly preceding veraison onset, hormone spike as critical signal, and ABA as veraison trigger or correlate. Abscisic acid peak timing shows ABA reaching maxima during 1-3 days surrounding veraison onset, peak timing varying slightly by cultivar and berry, hormone coordinating transformation across berry populations, and peak as hormonal event marking transition. Abscisic acid effects shows possible ABA triggering gene expression changes activating ripening, ABA possibly regulating softening through cell wall enzyme induction, hormone possibly affecting water relations and sugar transport, and ABA as potential central regulatory signal though mechanisms debated. Berry composition during termination shows water 78-82% or beginning increase, organic acids 0.7-0.9% declining with possible acceleration, sugars 13-17% with possible accumulation acceleration, pH 3.1-3.4 rising toward ripening range, and composition rapidly changing. Berry appearance during termination shows earliest berries showing clear color in red cultivars, white cultivars showing softening and possible yellowing, overall cluster showing asynchronous transformation with earliest berries entering veraison, and appearance showing transformation onset. Cluster asynchrony shows veraison beginning in earliest berries while later berries remain in lag, typical veraison spread over 5-14 days across cluster, asynchrony creating heterogeneous berry populations, and asynchrony as characteristic veraison pattern. Lag phase exit shows definitive transition from lag quiescence to ripening activity, growth rate increasing from minimal to low-moderate, metabolic activity increasing substantially, ripening processes fully activated, and exit marking lag phase end. Ripening phase entry shows berry entering ripening state with active sugar accumulation, color development in red cultivars, progressive softening, and ripening as post-lag developmental phase. Environmental effects on termination include temperature affecting termination timing with warm conditions accelerating transformation and shortening lag, cool conditions delaying veraison extending lag, water availability affecting transformation dynamics, and environmental factors determining lag duration and veraison onset. Temperature effects shows heat accumulation triggering veraison onset when threshold reached, growing degree days strongly correlating with veraison timing, warm conditions promoting earlier termination, and temperature as major lag duration and veraison timing determinant. The process shows three distinguishable aspects including veraison approach with transformation imminent, softening initiation with first texture change, and lag phase exit entering ripening. Duration typically 1-3 days representing lag-veraison transition period. Regional timing shows lag phase termination during early-mid September across climates as veraison begins. Viticultural significance includes representing definitive lag phase end and veraison onset, determining harvest timing through veraison date, demonstrating transformation from quiescence to ripening, and establishing berry ripening progression affecting wine quality. Research applications include analyzing veraison triggering mechanisms and regulation, investigating abscisic acid spike and hormonal control, evaluating softening initiation and cell wall modification, quantifying transformation timing and asynchrony, and assessing lag phase termination as determinant of ripening dynamics and harvest timing.
Lag_Phase_Termination represents lag conclusion within Berry_Growth_Lag_Phase. Subdivided into Veraison_Approach with transformation imminent, Softening_Initiation with 10-25% firmness decline, and Lag_Phase_Exit entering ripening. Duration 1-3 days lag-veraison transition. Color 0.05-0.2 mg/g anthocyanin in red cultivars. Growth rate possibly 5-20 mg/day. ABA peak 500-2000 ng/g. Water uptake 0.2-0.5 g/berry/day. Completes Berry_Growth_Lag_Phase. Marks veraison onset.
Lag-Phasen-Ende
Lag-Phasen-Terminierung
Lag-Veraison-Übergang
Veraison-Beginn
lag phase end
lag-veraison transition
veraison onset
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 468-478 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Lag Phase Termination
The Lage name Scharzhofberger designates a 28-hectare Einzellage in the Saar Valley within the Mosel Anbaugebiet, classified as a VDP.Große Lage (the highest site classification in the VDP system) and recognized as one of Germany's most distinguished Riesling sites — the name Scharzhofberger on a label communicates precise geographic origin to a degree comparable to Burgundy's Grand Cru system, with the Egon Müller estate's Scharzhofberger Riesling Trockenbeerenauslese achieving record auction prices exceeding 12,000 EUR per bottle. In contrast, the Großlage name Bernkasteler Badstube encompasses 200+ hectares from multiple communes, providing far less site specificity despite superficially similar label presentation.
A vineyard designation within the German wine classification system that identifies a wine as originating from a named Lage (vineyard site) — either an Einzellage (individual vineyard site) or Großlage (collective vineyard site) — as registered in the official German vineyard register (Weinbergsrolle), with the Lage name appearing on wine labels to communicate site-specific geographic origin within an Anbaugebiet (wine-growing region).
Lage_Name documents the German vineyard site naming system, which underwent fundamental restructuring in the 1971 Wine Law (Weingesetz) that consolidated approximately 30,000 historically named individual sites into approximately 2,650 Einzellagen and created approximately 150 Großlagen (collective site names covering larger areas). The Großlage concept — intended to simplify German wine labeling — has been widely criticized for creating consumer confusion because Großlage names appear on labels in the same format as Einzellage names without clear differentiation: a consumer encountering Bernkasteler Badstube (Großlage) cannot easily distinguish it from Bernkasteler Doctor (Einzellage) without specialist knowledge. The VDP (Verband Deutscher Prädikatsweingüter) has addressed this ambiguity through its private classification system that categorizes member estates' vineyards into Gutswein, Ortswein, Erste Lage, and Große Lage tiers — the latter designation reserved for the finest individual sites and voluntarily restricting yields and production methods beyond legal requirements.
Einzellage name
Einzellagen-Name
German vineyard site name
Lage designation
Weinbergslage
German Wine Law (Weingesetz 1971, amended 2021) §4-7 VDP (2023) VDP.Die Prädikatsweingüter Classification German Federal Institute for Agriculture and Nutrition (BLE) Weinbergsrolle database
Lage Name
A lamina on a mature leaf at node 6 of Vitis vinifera cv. Cabernet Sauvignon in Bordeaux, France, measuring 14.8 cm length by 13.5 cm width with total surface area of 128 cm² (both surfaces combined), thickness of 295 μm representing sun leaf development, exhibiting five-lobed palmate shape with deep U-shaped sinuses characteristic of this cultivar, distinct dorsiventral organization evident in cross-section showing dark green adaxial surface with thick cuticle (5.2 μm) and two palisade layers (total 88 μm), pale green abaxial surface with thinner cuticle (2.8 μm) and spongy mesophyll (182 μm), intercellular air space comprising approximately 22% of total lamina volume enabling gas diffusion, stomatal density of 175 stomata mm⁻² on abaxial surface (compared to 12 stomata mm⁻² on adaxial surface), reticulate vein network with median vein and four pairs of major lateral veins visible on both surfaces, containing approximately 8.5 × 10⁶ mesophyll cells per leaf providing photosynthetic capacity, demonstrating light-saturated photosynthesis rate of 13.2 μmol CO₂ m⁻² s⁻¹ and transpiration rate of 2.8 mmol H₂O m⁻² s⁻¹, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing complete lamina structure providing integrated photosynthetic function.
The flattened expanded blade portion of the grapevine leaf in Vitis vinifera, characterized by thin planar architecture typically 180-380 μm thickness optimized for light penetration to photosynthetic tissues, large surface area typically 50-250 cm² per leaf providing extensive light-intercepting capacity, palmate shape with radiating lobes creating characteristic grapevine leaf silhouette, dorsiventral anatomical organization with distinct upper (adaxial) and lower (abaxial) surfaces showing different structural and functional characteristics, internal tissue organization including palisade mesophyll concentrated beneath upper epidermis for maximum light capture, spongy mesophyll providing gas exchange spaces and lower light utilization, extensive intercellular air space system comprising 15-30% of lamina volume enabling CO₂ diffusion to photosynthetic cells and water vapor diffusion from mesophyll to stomata, and complex vascular network ramifying throughout lamina providing water and nutrient delivery and photosynthate export pathways. The lamina represents the primary site of photosynthetic carbon fixation, containing essentially all leaf chloroplasts concentrated in mesophyll cells, stomata enabling gas exchange predominantly on lower surfaces at densities of 100-250 stomata mm⁻² depending on cultivar and light environment, and structural features maintaining mechanical integrity while optimizing functional capacity through the critical balance between light interception, gas exchange efficiency, mechanical strength, and flexibility.
Lamina is used essentially synonymously with leaf blade, both terms referring to the flattened expanded photosynthetic portion of the leaf excluding the petiole. The lamina exhibits fundamental dorsiventral organization with distinct upper and lower surfaces differing in epidermal characteristics, cuticular thickness, stomatal density, and pigmentation. Anatomical structure shows remarkable plasticity responding to light environment during development, with sun leaves developing thicker laminas (280-380 μm) with multiple palisade layers while shade leaves show thinner laminas (180-250 μm) with single palisade layers. The critical functional balance involves maximizing light interception through large surface area while maintaining adequate mechanical strength, optimizing gas exchange through high stomatal density and intercellular air spaces while preventing excessive water loss, and achieving efficient photosynthesis through high chloroplast density while avoiding photoinhibition. Lamina dimensions and characteristics show systematic variation with leaf position, developmental stage, and environmental conditions, requiring careful standardization for comparative morphological studies. The term derives from Latin "lamina" meaning thin plate or layer, accurately describing the flat blade structure.
Blattfläche
Blattspreite
Lamina propria
blade lamina
expanded leaf portion
leaf blade
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 58-89 Patakas et al. (2003) "Differences in stomatal and photosynthetic responses of Vitis vinifera L. (cv. Sultana) leaves in response to progressive water stress" Vitis 42=113-115 Flexas et al. (2002) "Drought-inhibition of photosynthesis in C₃ plants= stomatal and non-stomatal limitations revisited" Annals of Botany 89=183-189 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 56-94.
Lamina
A lamina margin on a mature leaf at node 7 of Vitis vinifera cv. Merlot in Pomerol, France, exhibiting characteristic serrated configuration with heterogeneous tooth pattern alternating between large teeth (6-8 mm height) and small teeth (2-3 mm height), large teeth showing convex flanks and acute apices with 45-60 degree point angles, small teeth showing straight flanks and more obtuse apices with 70-90 degree angles, relatively shallow sinuses between teeth creating continuous undulating margin rather than deeply incised pattern, marginal veins visible as fine strands extending from lateral vein network to tooth apices, terminal hydathodes present at most large tooth apices appearing as small translucent spots occasionally exuding water droplets under high humidity conditions, marginal tissue showing slight thickening creating reinforced edge structure, tooth density of approximately 2.8 teeth per centimeter of margin length measured along a representative lateral lobe edge, overall marginal characteristics including tooth size ratio (large=small approximately 2.8=1), tooth shape, and patterns contributing to Merlot identification and distinction from similar cultivars, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing typical marginal features diagnostic for this major cultivar.
The peripheral edge or border of the grapevine leaf blade in Vitis vinifera, characterized by specialized marginal anatomy forming the boundary between lamina tissue and external environment, serrated or toothed configuration with alternating projections (teeth) and indentations (sinuses) creating characteristic grapevine leaf edge appearance, teeth typically arranged in heterogeneous patterns with alternating large and small teeth in many cultivars, tooth apex characteristics including acute, obtuse, or rounded points with or without terminal hydathodes, tooth flanks showing convex, straight, or concave profiles, sinus characteristics between teeth including depth and width affecting overall marginal appearance, marginal vein termination patterns with veins extending into tooth apices often terminating in hydathodes, and cultivar-specific marginal features including tooth size, shape, density, and patterns serving as important ampelographic diagnostic characters. The lamina margin serves multiple functions including defining leaf outline and area, providing structural reinforcement at the vulnerable blade periphery through specialized anatomy including marginal veins and thickened tissues, facilitating water release through marginal hydathodes in high humidity conditions preventing internal water stress, and exhibiting specialized developmental control establishing final leaf shape through marginal meristem activity during expansion determining ultimate lobe and tooth patterns that characterize cultivar morphology.
Lamina margin represents a specialized anatomical zone with distinct developmental, structural, and functional characteristics compared to interior lamina tissue. Marginal features show exceptional ampelographic diagnostic value, with tooth size, shape, density, and patterns serving as primary or secondary identification characters in most cultivar keys. Standardized terminology describes tooth characteristics including apex shape (acute, obtuse, rounded), flank curvature (convex, straight, concave), size relationships (homogeneous versus heterogeneous with alternating large and small), and sinus characteristics (deep, shallow, narrow, wide). Developmental control of marginal features involves marginal meristem activity during leaf expansion, with genetic programming determining tooth initiation sites and growth patterns creating cultivar-specific configurations, and environmental influences having relatively minor effects making marginal characters relatively stable diagnostics. Hydathodes at tooth apices serve as water release structures, exuding liquid water (guttation) under conditions of high root pressure and high atmospheric humidity, preventing excessive internal water potential and potentially facilitating nutrient transport to margins. Marginal pathogen entry represents a significant disease pathway, with many pathogens including Botrytis and Pseudomonas entering through marginal wounds or hydathodes, and marginal necrosis serving as common symptom of various stress conditions including water deficit, nutrient deficiency, and disease infection.
Blattperipherie
Blattrand
Laminakante
blade margin
leaf edge
leaf margin
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 136-182 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 33-55.
Lamina Margin
Georgian qvevri grape varieties (Rkatsiteli, Saperavi, and hundreds of local varieties) bear Landrace_Genetic_Resource_Roles as products of 8,000 years of selection in the putative domestication center of V. vinifera, carrying unique alleles for adaptation, disease tolerance, and distinctive wine character. Traditional Portuguese varieties from Douro, Dão, and Alentejo regions serve in Landrace_Genetic_Resource_Roles as locally-adapted populations selected over centuries for regional conditions and Port or table wine styles. Heritage Greek varieties (Assyrtiko, Xinomavro, Agiorgitiko among 300+ indigenous varieties) bear Landrace_Genetic_Resource_Roles as Mediterranean-adapted genotypes potentially carrying heat and drought tolerance valuable for climate change adaptation.
A genetic resource role that inheres in traditional, locally-adapted grapevine populations developed through centuries of farmer selection in specific regions, representing accumulated adaptation to local conditions, cultural preferences, and viticultural practices outside formal breeding programs. Landrace varieties are distinguished from modern cultivars by their origin through informal selection (farmer-maintained rather than institutionally bred), their adaptation to specific geographic and cultural contexts (regional identity rather than broad market targeting), and their typically heterogeneous population structure (mass selections rather than uniform clones). Grapevine landraces represent irreplaceable genetic diversity—alleles for local climate adaptation, traditional wine styles, and potentially novel disease tolerance that may be absent from the narrow genetic base of internationally traded varieties. Landrace conservation faces pressure from vineyard modernization (replacement with commercial clones of international varieties), abandonment of marginal viticulture regions, and loss of traditional viticultural knowledge. Many landraces survive only in small, aging vineyards or as scattered vines requiring urgent collection before they are lost.
Landrace_Genetic_Resource_Role captures the category of traditionally-maintained varieties distinct from both wild species (never cultivated) and modern bred varieties (institutionally developed). The landrace concept emphasizes the human-environment interaction that shaped these populations—centuries of selection under local conditions created adaptation potentially unavailable in varieties developed elsewhere or through formal breeding. Landrace diversity is concentrated in traditional viticultural regions (Caucasus, Mediterranean, Iberian Peninsula) where viticulture predates modern breeding and globalized variety distribution. The challenge of landrace conservation is urgent—modernization pressure replaces traditional varieties with commercial cultivars, while the knowledge systems supporting landrace cultivation are lost with aging farmer generations. Molecular characterization of landraces enables genetic inventory, duplicate identification, and priority-setting for conservation efforts with limited resources.
Landrasse
Landsorte
Lokalsorte
bäuerliche Sorte
farmer variety
heritage variety
landrace
local variety
traditional variety
traditionelle Sorte
This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Bacilieri R et al. (2013) Genetic structure of Mediterranean cultivated grapevine, BMC Plant Biology 13: 25 De Ferrari G et al. (2019) Conservation of grapevine landraces, PLOS ONE 14: e0225848 McGovern PE et al. (2017) Early Neolithic wine of Georgia, PNAS 114: E10309-E10318
Landrace Genetic Resource Role
The Landwein designation Pfälzer Landwein identifies wines from the Pfalz Landwein region in southwestern Germany, permitting a broader range of grape varieties and production methods than the corresponding Qualitätswein category for the Pfalz Anbaugebiet — Landwein must achieve a minimum must weight lower than Qualitätswein (typically 47° Oechsle versus 55-60° Oechsle), may include chaptalized wines, and is not subject to the official quality examination (Amtliche Prüfung) that Qualitätswein requires, but must originate from grapes grown within the defined Landwein region and display the regional character specified in the Landwein regulation.
A regional wine designation within the German and Austrian wine classification systems that identifies a wine as originating from one of the defined Landwein regions, classified as a Protected Geographic Indication (g.g.A.) under EU harmonization, representing the intermediate quality tier between Deutscher Wein/Österreichischer Wein (without geographic indication) and Qualitätswein/Qualitätswein bestimmter Anbaugebiete (Protected Designation of Origin).
Landwein_Designation captures the German-language equivalent of the Vin de Pays tier, with significant national variation between Germany and Austria. German Landwein has historically occupied a marginal commercial position between the economically dominant Qualitätswein and Prädikatswein categories and the residual Deutscher Wein (formerly Tafelwein) category, with only 1-2% of German wine production classified as Landwein: this contrasts sharply with the commercial success of French Vin de Pays, reflecting different market structures and consumer expectations. In Austria, Landwein similarly occupies the intermediate tier between Wein (table wine) and Qualitätswein but has gained some commercial traction as natural wine producers have adopted the Landwein category to avoid the stylistic constraints of the Qualitätswein DAC system. The EU classifies both German and Austrian Landwein as PGI (g.g.A.) under the harmonized framework.
Austrian Landwein classification
German regional wine designation
Landwein
Landwein-Bezeichnung
deutsche Landwein-Kategorie
regionale Weinbezeichnung
German Wine Law (Weingesetz) §20-22 Austrian Wine Law (Weingesetz 2009) §12 EU Regulation 1308/2013 Art. 93(1)(b)
Landwein Designation
The cultivar with prime name Grauburgunder carries language variant synonyms including Pinot Gris (French original from which the German name is translated), Pinot Grigio (Italian adaptation), Szürkebarát (Hungarian, literally grey friar), Rulandské Šedé (Czech), Szürkekbarát (erroneous Hungarian variant), and Sivi Pinot (Slovenian) — each representing a systematic linguistic transformation of the same etymological concept (grey + Burgundian/Pinot) into the respective national language, with the semantic relationship between names remaining transparent across languages.
A synonym that constitutes a translation, transliteration, or linguistic adaptation of a cultivar name into a different language, retaining semantic correspondence with the source name while conforming to the phonological, morphological, and orthographic conventions of the target language.
Language_Variant_Synonym captures the systematic cross-linguistic transformation of cultivar names, distinguishing these from regional synonyms where independent naming produced semantically unrelated designations. Language variant synonyms maintain etymological or semantic continuity with the source name through translation (Weissburgunder/Pinot Blanc/Pinot Bianco), transliteration (Каберне Совиньон for Cabernet Sauvignon in Cyrillic), or calque (Szürkebarát as a Hungarian calque of Graumönch/Pinot Gris): this semantic transparency contrasts with regional synonyms where the name-genotype relationship is arbitrary and historically contingent. Language variant synonyms are particularly common for widely planted international varieties that have been formally adopted into national variety catalogues.
Sprachvariante
cross-language synonym
fremdsprachiges Synonym
linguistic adaptation name
sprachliche Adaptation
translated synonym
OIV (2013) OIV List of Vine Varieties and Their Synonyms Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157 Robinson J et al. (2012) Wine Grapes, Allen Lane pp: 1-18
Language Variant Synonym
Trebbiano Toscano exhibits large cluster disposition with average weights of 450-700 g, rachis lengths of 18-28 cm, and berry counts of 200-400 per cluster. Italia table grape produces clusters of 500-900 g optimized for retail presentation. Airén develops large cylindrical-conical clusters averaging 350-500 g, contributing to its position as the most widely planted white variety globally by area. Large clusters in Grenache (300-450 g) combined with high fruitfulness result in potential yields exceeding 15 t/ha without cluster thinning. Mechanical harvest efficiency increases with cluster size due to improved target detection.
A cluster size disposition that is the inherited capacity of a Vitis vinifera plant to develop clusters of above-average weight (typically greater than 300 g) and dimensions, with extensive rachis branching, high berry count, or large individual berry size contributing to elevated cluster mass.
Large cluster disposition often correlates with high yield potential but may necessitate cluster thinning for quality wine production. The trait can complicate uniform ripening, as berries in cluster interiors receive less light and may lag in sugar accumulation. Large clusters are preferred for table grape and raisin production where total yield and impressive visual presentation are primary objectives. Rachis strength must scale with cluster weight to prevent bunch stem necrosis under heavy crop loads.
Großtraubigkeit
big bunch trait
große Traube
heavy cluster disposition
large bunch capacity
schwere Traube
Creasy GL, Creasy LL (2009) Grapes, CABI Publishing pp.: 78-95 Winkler AJ et al. (1974) General Viticulture, University of California Press pp.: 234-267
Large Cluster Disposition
Cabernet Sauvignon grafted on 1103P rootstock showing large diameter rootstock xylem particularly prominent in earlywood zones, vessel measurements revealing mean earlywood diameter 152 micrometers (range 135-172 micrometers) qualifying as large-diameter class, large vessels visible as conspicuous pores in cross-section distinguishable at low magnification, hydraulic conductivity calculations indicating large 150 micrometer vessels conducting approximately 5 times water volume of medium 100 micrometer vessels under equivalent pressure, large vessel structure characteristic of vigorous 1103P rootstock enabling high water transport capacity supporting vigorous vine growth, some large vessels showing tyloses formation in heartwood zone indicating normal vessel occlusion during sapwood-heartwood conversion
Large diameter rootstock xylem is rootstock xylem characterized by vessels with large lumen diameter typically greater than 120 micrometers, emphasizing vessel size rather than position within ring or seasonal formation timing, resulting from genetic determination in vigorous rootstock cultivars producing characteristically large vessels, favorable formation conditions including adequate water and nutrients enabling maximum cell expansion, or specific ring position particularly earlywood where large vessels advantageous for water transport, showing very high hydraulic conductivity per vessel due to flow rate increasing with fourth power of radius making large vessels disproportionately efficient for water conduction, but exhibiting greater vulnerability to cavitation and embolism formation under drought or freeze stress compared to smaller vessels, and typically found in vigorous rootstock varieties such as Berlandieri hybrids, 1103P, or 110 Richter supporting high-vigor vine water demands. Large diameter rootstock xylem vessel size definition includes= showing diameter greater than 120 micrometers (large vessels typically 120-180 micrometers compared to medium 80-120 micrometers or small 40-80 micrometers), exhibiting substantial lumen volume (large diameter creating proportionally enormous internal volume for water transport since volume increases with square of radius), displaying prominent visibility (large vessels conspicuous as pores in cross-section or grooves in longitudinal section visible to naked eye or low magnification), demonstrating size-dependent properties (large diameter conferring specific hydraulic and vulnerability characteristics distinct from smaller vessels), and showing relative definition (vessel considered large in context of rootstock cultivar typical range and grapevine xylem general size distribution). Large diameter rootstock xylem hydraulic efficiency includes= providing disproportionate conductivity (single large vessel conducting water volume proportional to radius to fourth power making 180 micrometer vessel conduct approximately 10 times volume of 100 micrometer vessel under same pressure gradient), enabling high flow rates (large vessels supporting rapid water movement adequate for high transpiration demands), showing maximum transport capacity (large vessel zones providing peak hydraulic conductivity supporting vigorous vine canopy), demonstrating low resistance (large lumen minimizing friction enabling efficient water flow with minimal pressure drop), and requiring fewer vessels (large diameter vessels achieving given conductivity with fewer individual conduits than small vessel network reducing cellular investment). Large diameter rootstock xylem formation factors includes= resulting from genetic programming (vigorous rootstock cultivars producing characteristically large vessels as inherited trait determined by rootstock genetics), reflecting favorable conditions (adequate spring water and nutrients enabling cambium to produce maximum vessel diameter during differentiation), showing cultivar characteristics (Berlandieri hybrids, 1103P, 140Ru, 110R naturally producing large vessels while Riparia, 101-14 producing smaller vessels), exhibiting position effects (earlywood typically containing largest vessels in ring with latewood showing smaller diameters regardless of genetics), and demonstrating vigor correlation (large vessels associated with high rootstock vigor classification reflecting genetic growth capacity). Large diameter rootstock xylem cavitation vulnerability includes= showing increased embolism risk (large vessels potentially more susceptible to air-seeding during water stress due to larger pit membrane areas and higher tension), exhibiting drought sensitivity (severe water stress potentially causing cavitation in large vessels more readily than small vessels compromising transport), demonstrating freeze-thaw risk (ice crystal formation during freezing and subsequent thaw potentially introducing embolisms more readily in large vessels), requiring redundancy (large vessel networks needing multiple parallel pathways ensuring continued function despite individual vessel embolism), and showing recovery capacity (refilling of embolized large vessels potentially occurring during stress relief restoring transport function). Large diameter rootstock xylem in vigorous rootstocks includes= characterizing Berlandieri hybrids (cultivars showing large vessels 130-180 micrometers enabling high transport capacity), defining 1103P characteristics (very vigorous rootstock producing abundant large vessels 140-175 micrometers supporting high-vigor vine), appearing in 110 Richter (extremely vigorous rootstock showing large vessels 135-170 micrometers), present in 140 Ruggeri (vigorous selection with large vessels 130-165 micrometers), and enabling vigor expression (large vessels providing hydraulic capacity necessary for vigorous vegetative growth characteristic of high-vigor rootstock classifications). Large diameter rootstock xylem in earlywood includes= comprising majority of earlywood vessels (spring-formed xylem dominated by large vessels optimizing water transport for early season demand), showing maximum size in inner earlywood (largest vessels typically in first-formed portion of earlywood representing peak vessel expansion), providing peak ring conductivity (earlywood large vessels contributing majority of annual ring hydraulic capacity), demonstrating seasonal timing (large vessel formation coinciding with bud break and shoot growth when water demand highest), and creating visible ring structure (large earlywood vessels contrasting with smaller latewood vessels creating distinct annual ring pattern visible in cross-section). Large diameter rootstock xylem measurement and identification includes= measuring lumen diameter (determining internal diameter of vessel from inner wall to inner wall typically using microscopy requiring cross-sections), averaging across population (characterizing wood as large-diameter based on mean vessel size across measured sample of vessels), identifying in cross-section (large vessels appearing as conspicuous pores visible in transverse cuts distinguishable from surrounding smaller elements), assessing distribution (determining proportion of xylem comprising large vessels revealing vessel size structure), and comparing to standards (classifying vessels as large relative to established size classes or cultivar norms). Large diameter rootstock xylem functional trade-offs includes= providing maximum efficiency when functional (large vessels offering highest conductivity per vessel when operating under favorable conditions), showing vulnerability under stress (large vessels risking embolism under drought or freeze potentially compromising transport when conditions unfavorable), requiring greater initial investment (producing large vessel requiring more cellular resources during formation than smaller vessel), demonstrating efficiency-safety trade-off (large vessels optimizing efficiency sacrificing safety compared to smaller more resistant vessels), and necessitating compensatory features (large vessel systems requiring redundancy, refilling capacity, or mixed vessel sizes managing vulnerability risks).
Large_Diameter_Rootstock_Xylem emphasizes vessel size greater than 120 micrometers and is distinguished from Rootstock_Xylem (all xylem without size emphasis) by focus on large lumen diameter, high hydraulic efficiency, cavitation vulnerability, and vigorous cultivar association. Critical for understanding hydraulic capacity, efficiency-safety trade-offs, cultivar vigor differences, water transport optimization, and the large vessels providing maximum conductivity while showing increased stress vulnerability.
High-conductivity rootstock xylem
Hochleitfähiges Unterlagenxylem
Large-lumen rootstock xylem
Weitgefäßiges Wurzelstockxylem
Wide-vessel rootstock xylem
Large Diameter Rootstock Xylem
Cabernet Sauvignon large diameter xylem vessel in earlywood of ring 7 shows 230-290 μm lumen diameter, thin wall 2.4-3.6 μm thickness, lumen cross-sectional area 41,500-66,000 μm², individual vessel providing hydraulic conductivity 30-85 times greater than average 65 μm latewood vessel, large vessels comprising 18-24% of total ring vessel number but providing 65-72% of ring hydraulic conductance, vessels formed April-May supporting spring canopy development but showing 55-75% embolism by late summer under water deficit conditions
A large diameter xylem vessel is a xylem vessel characterized by substantial lumen diameter (typically 150-300 μm, though some sources use threshold of >100 μm or >150 μm for "large" classification), providing disproportionately high hydraulic conductance due to radius⁴ relationship between diameter and flow rate, typically formed as earlywood vessels but defined by size rather than formation timing. Large diameter vessel anatomical characteristics include= substantial lumen diameter (150-300 μm typical range, exceptional vessels reaching 350+ μm in some vigorous cultivars or rings), thin secondary walls relative to diameter (wall thickness 2-4 μm representing <2% of vessel diameter), very large lumen cross-sectional area (17,700-70,700 μm² for 150-300 μm diameter vessels), simple perforation plates with large openings (minimal end wall resistance to axial flow), abundant bordered pits (60-100 pits per 100 μm² wall area providing extensive lateral connections), and prominent visibility (large vessels easily visible to naked eye in cross-section as pores or on radial surfaces as grooves). Large diameter vessel hydraulic dominance includes= extreme conductance efficiency (single 250 μm vessel conducting equivalent to 256 vessels of 50 μm diameter based on radius⁴ relationship), providing 50-75% of trunk hydraulic conductance (despite being only 15-30% of total vessel number), disproportionate contribution per vessel (individual large vessel contributing 10-100 times more than average vessel), determining maximum water transport capacity (large vessel number and size controlling peak hydraulic conductance), and critical for supporting large canopies (large vessels enabling high transpiration rates supporting extensive leaf area). Large diameter vessel formation conditions include= favorable early season growth (adequate water, nutrients, warmth enabling maximum vessel expansion), vigorous cambial activity (high division rates producing large fusiform derivatives), young vigorous vines (establishment years showing maximum vessel diameters), wide annual rings (favorable growth years producing larger maximum vessels than narrow stress rings), and genetic factors (cultivar characteristics determining maximum potential vessel size). Large diameter vessel embolism vulnerability includes= highest cavitation susceptibility (large vessels most prone to embolism due to large pit membrane area and high tensions), preferential dysfunction under stress (water deficit causing large vessel embolism first and most extensively), severe consequences of loss (loss of few large vessels having major impact on hydraulic capacity), rapid drought stress development (large vessel embolism quickly reducing conductance causing severe water stress), and catastrophic failure potential (severe stress embolizing majority of large vessels causing hydraulic crisis potentially leading to vine death). Large diameter vessel mechanical limitations includes= thin walls providing minimal structural support (large vessels contributing little to wood strength), vulnerability to collapse (large thin-walled vessels potentially crushing under mechanical stress), dependence on surrounding support (fibers and other thick-walled cells preventing large vessel collapse), and optimization for efficiency over safety (large vessels evolved for maximum conductance sacrificing mechanical strength and embolism resistance). Large diameter vessel distribution patterns include= concentration in earlywood (most large vessels formed during spring, producing tangential bands in inner annual ring portions), absent or rare in latewood (summer formation producing smaller vessels typically 40-120 μm), radial gradient (outer sapwood rings sometimes showing larger maximum vessels than inner rings), circumferential uniformity (large vessels distributed evenly around trunk circumference), and cultivar variation (some varieties characteristically producing larger or smaller maximum vessels). Large diameter vessel density and proportion includes= lower numerical density (8-20 large vessels per mm² compared to 30-55 smaller vessels), large vessels comprising 10-25% of total vessel number (despite providing majority of conductance), high lumen area fraction in earlywood (large vessels occupying 45-65% of earlywood cross-sectional area), disproportionate hydraulic importance (15-25% of vessels providing 50-75% of conductance), and strategic positioning (concentration in earlywood zone maximizing early-season hydraulic capacity). Large diameter vessel length characteristics include= vessel elements 300-600 μm length (individual cellular units), total vessels potentially very long (series of elements creating continuous tubes 1-10+ meters in some trunk sections), Vitis as long-vesseled species (large vessels potentially extending substantial trunk distances), long vessels enabling efficient conductance (minimal end wall resistance over extended distances), but long vessels facilitating embolism spread (air bubbles potentially propagating through extended vessel length). Large diameter vessel significance includes= determining maximum vine water transport (large vessel characteristics setting upper limit on hydraulic capacity), controlling supportable canopy size (large vessel conductance ultimately limiting sustainable leaf area), critical for vigor and productivity (adequate large vessel formation enabling high growth and yield potential), vulnerability determining drought tolerance (large vessel embolism susceptibility affecting vine performance under water deficit), and annual renewal importance (yearly large vessel formation restoring hydraulic capacity after previous year's embolism).
Large_Diameter_Xylem_Vessel emphasizes vessel size and hydraulic dominance and is distinguished from Early_Wood_Xylem_Vessel (seasonal timing emphasis), Xylem_Vessel (all vessels), and Late_Wood_Xylem_Vessel (small vessels) by substantial diameter and disproportionate conductance contribution. Critical for understanding hydraulic capacity determination, embolism vulnerability concentration, drought stress mechanisms, annual hydraulic capacity renewal, and tradeoffs between transport efficiency and safety in xylem evolution.
Breites Xylemgefäß
Broad conduit
Großdurchmesser-Gefäß
High-conductance vessel
Hochleistungs-Leitgefäß
Wide vessel
Large Diameter Xylem Vessel
Documentation of Large_Leaf_Disposition at the USDA Grape Germplasm Repository characterized leaf development in cultivars recognized for large foliage. Reference cultivars 'Sultanina', 'Emperor', 'Italia', and 'Cardinal' were assessed under standardized conditions with detailed measurements of leaf area, expansion dynamics, and canopy consequences. Large_Leaf_Disposition cultivars showed consistent characteristics. 'Sultanina' exhibited archetypal large leaves with mature leaf area of 365 cm² (mean of nodes 9-11), blade length of 22.5 cm and width of 24.8 cm, achieving 90% of final area by 35 days after emergence. The expansion rate peaked at 15.2 cm²/day during the rapid growth phase (days 12-22). 'Italia' showed even larger leaves (385 cm², the largest among commercial cultivars assessed) with correspondingly extended expansion period (38 days to 90% area). 'Emperor' displayed typical Large_Leaf_Disposition (325 cm²) with notably thick lamina (see Leaf_Thickness_Disposition). Canopy development analysis revealed that Large_Leaf_Disposition cultivars at 15 shoots/m achieved total leaf area index (LAI) of 4.2 m²/m² ground by mid-season, exceeding the optimal range (2.5-3.5 m²/m²) for fruit zone light exposure. Fruit zone light penetration in unmanaged Large_Leaf_Disposition canopies averaged only 8% of ambient, compared to 25% in small-leaved cultivars at equivalent shoot density, demonstrating the shading consequence. Management trials confirmed that Large_Leaf_Disposition cultivars required either 25% lower shoot density or 40% more leaf removal labor to achieve equivalent canopy light environment compared to moderate-leaved cultivars.
A Large_Leaf_Disposition is a Leaf_Size_Disposition that inheres in Vitis vinifera leaves and is realized through extended cell division and expansion phases producing individual mature leaf areas exceeding 200 cm², creating substantial foliar surfaces that rapidly accumulate canopy leaf area but may contribute to excessive shading in dense canopies. This disposition represents the upper range of leaf size, characteristic of cultivars that invest heavily in individual leaf development, producing fewer but larger leaves to achieve total canopy area. Large-leaved cultivars develop prominent, visually striking foliage that reaches full expansion over approximately 35-40 days following emergence, with high cell number and large cell size combining to produce extended leaf blades. The viticultural implications of Large_Leaf_Disposition include rapid canopy development (fewer leaves required to achieve target leaf area), potentially excessive canopy density at standard shoot densities (requiring shoot thinning or leaf removal), increased self-shading within individual leaves and between leaf layers, larger boundary layer affecting heat dissipation and gas exchange, and distinct visual character affecting vineyard aesthetics. Large leaves may be advantageous in low-vigor situations where maximizing leaf area from limited shoot growth is desirable, but can create management challenges in high-vigor situations by contributing to excessive canopy density. Many table grape and raisin cultivars exhibit Large_Leaf_Disposition, potentially reflecting selection for vegetative vigor in production systems focused on fruit quantity.
Large_Leaf_Disposition is_a Leaf_Size_Disposition representing individual leaf area > 200 cm². Characterized by extended cell division phase, high cell number, and large cell size. Examples include 'Sultanina', 'Italia', 'Emperor', 'Cardinal', 'Muscat of Alexandria'. Table grape and raisin cultivars commonly show this trait. Creates canopy management challenges through rapid area accumulation and increased shading. Advantageous in low-vigor situations problematic in high-vigor situations. OIV descriptor 065 codes 7-9.
Großblättrige-Disposition
Großblättrige-Sorte
Hohe-Einzelblattfläche
expansive foliage
high leaf area
large-leaved cultivar
Galet P (1979) A Practical Ampelography: Grapevine Identification, Cornell University Press Palliotti A et al. (2014) Changes in vineyard establishment and canopy management urged by earlier climate-related grape ripening: a review, Scientia Horticulturae 178:43-54 Organisation Internationale de la Vigne et du Vin (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition
Large Leaf Disposition
Cabernet Sauvignon in Napa Valley shows last caps falling on June 16 Year N+1 at late BBCH 69, with final 3-6% flowers predominantly in basal positions completing cap detachment, visible berry development 12-18 mm diameter representing 58-66% established fruit set throughout cluster, under warm climate conditions of 25-29°C temperature, typical of this late-season cultivar completing final cap fall 12-13 days after first detachment establishing flowering completion with last caps representing completion formality contributing negligible fruit set
Last caps falling represents the final individual calyptra detachments when remaining 3-8% unopened flowers complete abscission establishing complete cap shedding extent within cluster and marking imminent flowering completion with no further viable flowers remaining. This final detachment phase occurs during complete anthesis in late June-early July in temperate regions Year N+1 as part of final cap fall, typically spanning 1 day or brief hours as very last flowers complete cap shedding, corresponding to BBCH 69 final stage. Final flower characteristics show last-to-mature individuals typically in most basal cluster positions reflecting latest primordium formation previous season and most delayed current season development, occasional scattered positions from locally abnormal delayed development, and overall final flowers representing developmental tail of cluster flowering. Final cap detachment timing shows last individual events completing cluster cap fall, detachments occurring sporadically as final flowers individually reach maturity, and cessation of activity establishing complete cap shedding achievement. Detachment process in final flowers proceeds through same abscission mechanisms as earlier flowers with enzymatic degradation of middle lamella creating separation plane, mechanical forces from organ expansion driving cap lifting, and final separation completing calyptra removal, though final flowers may experience less favorable conditions affecting efficiency. Environmental conditions at final detachment show reduced criticality since reproductive outcomes already determined with 92-97% berries established from earlier successful pollinations, though conditions still affect final detachment success completing flowering process. Cap accumulation from final detachments adds minimal incremental debris to substantial previously accumulated layers, with last few caps barely noticeable among abundant earlier accumulation, though completing total flowering documentation. Flower exposure from final detachments reveals last reproductive organs though with minimal functional significance, as very limited viable pollen remains and atmospheric pollen completely expired, creating negligible reproductive opportunity for last-opening flowers. Reproductive potential of final flowers shows essentially zero probability of successful pollination and fruit set due to complete pollen exhaustion, expired stigma receptivity potential due to suboptimal late conditions, and overall failed reproductive opportunity though mechanical flowering completion proceeds. Berry development visibility at last caps falling shows advanced early-set berries 12-18 mm diameter from peak flowering success, visible throughout cluster creating predominant appearance, contrasting with minimal final flower opening highlighting developmental transition. Unfertilized flower senescence shows widespread brown discoloration in failed reproductions from throughout flowering period, active abscission beginning to eliminate senescing flowers, and overall cluster cleaning toward berry-only appearance. Cluster appearance at last caps falling shows definitive transitional state with developing berries as dominant visual feature creating young fruit cluster impression, minimal final opening activity barely noticeable among berry-dominated appearance, and overall appearance confirming developmental phase establishment. Observational requirements for detecting last caps falling include careful inspection identifying very last detachment events, distinction between final viable flowers versus persistent aborted buds incapable of normal development, and confirmation that no further capable flowers remain establishing true completion. Completion proximity indicated by last caps falling shows flowering conclusion imminent within hours to one day, very minimal residual flowering activity remaining, and clear trajectory toward definitive cap fall completion. Phenological significance of last caps falling includes indicating imminent BBCH 69 complete flowering achievement, providing practical observation that flowering nearly completed enabling final management transitions, and establishing near-completion state preparing for post-flowering operations. Management implications include confirming flowering nearly complete enabling final preparations for post-bloom operations including disease management resumption and canopy operations, recognizing that critical flowering period effectively concluded with minimal remaining vulnerability, and transitioning focus definitively to berry development priorities. Fruit set contribution from last flowers shows negligible addition to berry number with essentially zero successful pollinations from final openings, confirming that yield potential established during earlier flowering phases, and last caps falling representing completion formality rather than productive contribution. Developmental progression from last caps falling proceeds immediately toward complete cessation of cap fall activity, rapid transition to definitive flowering completion, and full commitment to berry development phase. The process represents first aspect of final cap fall marking very last detachment events, followed by cap fall completion confirming definitive cessation. Duration represents brief hours to one day as final events complete, with rapid progression toward completion. Cultivar variation in last caps characteristics shows differences in number of final flowers with some varieties showing minimal 2-4% tail while others showing 6-10% final phase, timing of last detachments reflecting overall phenology, and efficiency of final shedding with some showing clean completion while others showing occasional persistent unopened buds. Regional timing shows last caps falling during late June in warm climates, late June-early July in temperate regions, and early-mid July in cool areas, occurring as final events of flowering period. Viticultural significance includes indicating imminent flowering completion enabling final management transitions, confirming that productive flowering concluded with last caps representing completion formality, and providing observable marker that critical flowering period effectively ended. Research applications include documenting complete flowering extent through final detachment observation, analyzing final flower characteristics representing developmental extremes, evaluating completion efficiency assessing proportion achieving complete opening, and assessing implications of final flowering characteristics for developmental outcomes.
Last_Caps_Falling is first aspect of Final_Cap_Fall, distinguished by very last individual detachment events. Represents final 3-8% flowers completing cap fall. Indicates imminent flowering completion within hours to day. Negligible reproductive contribution since pollen expired. Observable marker of near-completion enabling final management preparations. Followed by Cap_Fall_Completion confirming definitive cessation. Brief hours to one-day duration as final events complete.
Endkappen-Abwurf
final cap detachments
finale Kappenablösungen
letzte Kappen fallen
terminal calyptra shedding
terminale Kalyptralösung
ultimate cap fall
very last caps falling
Last Caps Falling
Black Monukka exhibits characteristic late seed abortion at 3-5 weeks post-fertilization (BBCH 75-77), producing 2.5-3.5 mm soft brown seed remnants and achieving 12-14 mm berry diameter without requiring gibberellin applications, with abortion occurring after torpedo-stage embryo formation (28-35 days post-fertilization)
A late aborted seed is an aborted seed of Vitis vinifera that undergoes developmental failure and tissue degeneration during mid- to late developmental stages (3-8 weeks post-fertilization, corresponding to BBCH 75-79), after substantial embryo and endosperm development has occurred, resulting in larger seed remnants (typically 2-4 mm) and intermediate impact on final berry size. Late seed abortion occurs after seeds have already exerted significant hormonal influence on berry development during Phase I growth (cell division phase, 0-4 weeks post-fruit set), allowing partial berry expansion before abortion-associated growth cessation. This abortion pattern can occur in both seedless cultivars with delayed abortion mechanisms and in normally seeded cultivars under environmental stress conditions during mid-berry development. Developmental characteristics of late aborted seeds include= abortion occurring during embryo torpedo stage through early maturation phase (21-56 days post-fertilization), formation of recognizable embryo and endosperm structures before abortion initiation, gradual tissue browning and collapse over 1-2 weeks, and production of relatively large (2-4 mm) soft brown remnants that may retain some seed shape definition. Late abortion timing results in intermediate berry development outcomes= because seeds have already promoted significant berry cell division and initial expansion through endogenous hormone production during first 3-4 weeks, berries with late-aborted seeds achieve larger final sizes (10-14 mm diameter) compared to early-abortion berries (6-10 mm) but smaller than fully seeded berries (12-18 mm) of the same cultivar. Some seedless table grape cultivars (e.g., Black Monukka, Perlette) exhibit characteristically late abortion mechanisms, producing 2-3 mm seed remnants that may be slightly noticeable in texture but are generally acceptable for fresh consumption. Late aborted seeds in normally seeded wine grape cultivars indicate environmental stress or physiological problems during mid-berry development= severe water deficit during Phase I berry growth (leading to competition for limited assimilates), extreme heat events (35-40°C) during embryo/endosperm development phase, mineral deficiencies affecting seed metabolism, or vine stress conditions that compromise assimilate supply to developing reproductive tissues.
Late_Aborted_Seed emphasizes delayed developmental failure timing and is distinguished from Early_Aborted_Seed (minimal embryo development before abortion) by larger seed remnant size and partial berry size development before abortion. Critical for understanding delayed abortion mechanisms in some seedless cultivars, intermediate berry size outcomes, and mid-development stress indicators in seeded wine grape production.
Delayed abortion seed
Größerer Kernrest
Larger seed trace
Spätabortierter Samen
Verzögert abortierte Samenanlage
Ebadi, A., Sedgley, M., May, P., & Coombe, B. G. (1996). Seed development and abortion in Vitis vinifera L., cv. Chardonnay. International Journal of Plant Sciences, 157(6), 703-712. | Ojeda, H., Deloire, A., & Carbonneau, A. (2001). Influence of water deficits on grape berry growth. Vitis, 40(3), 141-145.
Late Aborted Seed
A late aborted tendril at node 9 on a heavily cropped shoot of Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, California, measuring 8.5 cm length (approximately 45% of the normal 18-20 cm for functional tendrils at this position), exhibiting well-developed bifurcated structure with clear vascular bundles visible through translucent epidermis, brown necrotic tip and progressive desiccation from tip to base, occurring during the critical berry development period in July when fruit cluster demand for photosynthates reached peak levels and vine crop load of 2.8 kg/m canopy exceeded recommended maximum of 2.0 kg/m, with 35% of tendrils on heavily cropped vines showing similar late abortion compared to <5% on appropriately cropped control vines, representing resource competition-induced developmental failure that simultaneously compromised climbing support capacity and indicated excessive crop load requiring cluster thinning intervention.
An aborted tendril that ceased development during advanced morphogenesis after achieving substantial structural differentiation but before completing functional maturation in Vitis vinifera, characterized by arrested development at 4-12 cm length representing 25-70% of normal mature tendril length, presence of well-developed tissue organization including complete vascular differentiation, epidermis with some cuticle development, and recognizable tendril morphology including branching patterns where appropriate, but failure to complete final elongation, achieve full physiological competence including mechanosensory function, or develop the structural strength necessary for functional climbing. Late aborted tendrils result from stress conditions or resource limitations occurring during the later phases of tendril development when basic morphogenesis is complete but final maturation processes including full cell expansion, wall thickening, cuticle deposition, and metabolic activation of sensory systems remain incomplete, typically caused by moderate to severe water stress, competition with developing fruit clusters for limited carbohydrate resources, nutrient deficiencies affecting cell wall synthesis or metabolic function, pest or pathogen damage during mid to late development, or environmental conditions including prolonged cool temperatures that slow metabolic processes. These abortion events represent substantial metabolic investment loss as tissues have undergone extensive differentiation and consumed significant resources, but occur before achievement of functional return through climbing support, creating inefficiencies in resource allocation that can impact overall vine carbon balance and competitive fitness.
Late aborted tendrils are distinguished from early aborted tendrils by substantial achievement of structural development and tissue differentiation, and from functional tendrils by failure to complete final maturation processes necessary for climbing function. This abortion class is particularly common in commercial viticulture where crop load management creates competition between reproductive and vegetative development, with late tendril abortion serving as a sensitive indicator of resource limitation affecting vegetative growth. The developmental stage at abortion within the late abortion category (mid-development versus near-mature) provides finer resolution on stress timing and severity, with earlier late abortion suggesting more severe or prolonged stress compared to near-mature abortion indicating acute late-stage stress. Late abortion represents the highest resource loss among abortion categories, as extensive differentiation and growth have consumed carbohydrates and nutrients without producing functional return.
advanced stage failed tendril
fortgeschrittenes Stadium fehlgeschlagene Ranke
late development arrested tendril
near-completion aborted tendril
spät-abortierte Ranke
späte Entwicklungsphase abgebrochene Kletterorgan
Based on Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Petrie et al. (2000) "The effect of leaf removal and canopy height on whole-vine gas exchange and fruit development of Vitis vinifera L. Sauvignon blanc" Functional Plant Biology 27=173-184 Edson et al. (1995) "Influence of crop load on vegetative growth, dry matter partitioning, and mineral nutrient partitioning of Chenin blanc grapevines" American Journal of Enology and Viticulture 46=469-477 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 401-424.
Late Aborted Tendril
Cabernet Franc in Loire Valley shows late anthesis during mid-late June Year N+1 at BBCH 67-68 over 2-3 days June 12-14, with final 22-28% flowers opening predominantly in basal cluster regions, pollen viability declining to 35-55% in late-opening flowers, pollination success reducing to 28-38% compared to 52-62% during peak, under temperate maritime conditions of 20-25°C temperature, 60-75% relative humidity, variable weather, typical of this mid-season cultivar completing flowering with late anthesis contributing 18-24% final berry number
Late anthesis represents the declining flowering phase when final 20-30% flowers complete opening while earlier-opened flowers senesce, pollen viability diminishes, and pollination window closes, marking transition from peak reproductive opportunity toward flowering completion and fruit development initiation. This concluding reproductive phase occurs during mid-late June in temperate regions Year N+1 following peak anthesis, typically spanning 2-3 days as remaining flowers open creating final pollination opportunities before anthesis completion, corresponding to BBCH 67-68 stages. Flowering intensity during late anthesis shows characteristic decline with decreasing daily opening rate dropping from 10-20% during peak to 3-8% during late phase, cumulative effect creating 70-90% flowers opened by late anthesis, and remaining 10-30% flowers completing opening during this final phase. Flower distribution pattern shows apical and middle regions predominantly showing opened or senescing flowers with 85-100% flowers past peak, while basal regions show final opening activity with 50-80% flowers opening during late phase, creating spatial gradient with most advanced development apically and latest development basally. Reproductive organ status during late anthesis shows mixed condition across cluster with earliest-opened flowers showing senescent organs including withered anthers depleted of pollen, declining stigma receptivity with reduced exudate production and beginning browning, and overall flower aging approaching abscission if unfertilized. Recently opened flowers in late anthesis show functional organs but suboptimal conditions with anthers dehiscing but pollen showing reduced viability compared to peak period, stigmas receptive but with lower exudate production and shorter functional period, and overall reduced reproductive capacity compared to earlier-opening flowers. Pollen availability during late anthesis shows substantial decline with remaining viable pollen from recently dehisced anthers representing final opportunity, cumulative pollen from earlier opening largely expired with viability reduced to 10-30% or less, and overall pollen density in cluster atmosphere declining substantially creating limited pollination opportunity. Pollen viability decline proceeds rapidly during late anthesis with fresh pollen from late-opening flowers showing 60-80% germination capacity compared to 70-90% during peak, and aged pollen from earlier opening showing minimal 5-20% viability, creating reduced pollination success potential for late-opening or previously unpollinated flowers. Stigma receptivity during late anthesis shows declining function with recently opened stigmas showing moderate receptivity 60-80% peak levels, stigmas from earlier-opened flowers showing substantially reduced receptivity 20-40% peak levels as exudate production declines and tissues begin senescence, and overall reduced female reproductive capacity across cluster. Pollination efficiency during late anthesis drops substantially with estimated successful pollination rates of 20-40% for late-opening flowers compared to 50-70% during optimal window, resulting from combined effects of reduced pollen availability, decreased pollen viability, diminished stigma receptivity, and potentially less favorable environmental conditions. Fertilization success from late anthesis pollinations shows reduced rates with longer time required for pollen tube growth potentially 24-48 hours due to suboptimal conditions, and higher probability of fertilization failure resulting in flower abscission, affecting final fruit set contribution from late anthesis period. Fruit set from late anthesis typically represents 15-25% total berry number established during this final phase, compared to 60-80% during peak anthesis, with late-set berries often showing slightly delayed development compared to earlier-set berries potentially affecting harvest uniformity. Developmental heterogeneity increases during late anthesis with wide range of flower states coexisting including newly opening flowers, recently pollinated flowers beginning fruit development, senescing unfertilized flowers preparing for abscission, and small developing berries from successful earlier pollinations, creating complex mixed developmental appearance. Environmental effects during late anthesis continue affecting reproductive success with optimal conditions 20-28°C maintaining adequate though declining organ function, while suboptimal cool or warm conditions further reduce reproductive capacity, and any adverse weather during late anthesis affecting final fruit set contribution. Cluster appearance during late anthesis shows transitional character with mixture of opened flowers, withering unfertilized flowers showing brown discoloration, small green developing berries 3-5 mm diameter from successful earlier fertilizations, and remaining unopened buds in basal positions, creating characteristic appearance of reproductive transition. Flower senescence becomes prominent during late anthesis with unfertilized flowers showing progressive deterioration including anther withering and browning, stigma desiccation and discoloration, petal remnant browning if not completely shed, and preparing for abscission through abscission zone activation at pedicel base. Flower abscission initiation begins during late anthesis with earliest unfertilized flowers developing abscission zones and detaching, creating visible flower drop that will continue through post-anthesis period, with successful fertilization preventing abscission through hormonal signals from developing embryo. Berry development visibility increases during late anthesis as successfully fertilized ovaries expand becoming recognizable as small berries distinguishable from unopened buds or unfertilized flowers, providing early indication of fruit set success and potential yield. Hormonal changes during late anthesis include increasing auxin production from successfully fertilized ovaries preventing flower abscission and triggering berry expansion, possible ethylene increase in unfertilized flowers promoting senescence and abscission, and declining reproductive hormones as flowering culminates. Metabolic transition during late anthesis shifts from reproductive to developmental focus with declining investment in flower function and increasing allocation to successfully fertilized ovaries initiating berry development, marking transition from flowering to fruit development as dominant process. Resource allocation changes during late anthesis redirect photosynthate from declining reproductive demands toward developing berries requiring substantial resources for early rapid growth, continuing vegetative growth with shoot elongation and leaf expansion, and beginning preparation for subsequent phenological stages. Weather sensitivity during late anthesis remains high though reduced compared to peak period, with adverse conditions affecting late-opening flowers but having minimal impact on successfully pollinated earlier flowers already initiating berry development, and rain during late anthesis causing less catastrophic effects than during peak though still reducing late pollination success. Disease pressure during late anthesis emphasizes botrytis with senescing flower tissues providing favorable substrate for fungal colonization, latent infections established during flowering remaining dormant in developing berries, and transition to berry protection becoming management priority. Management considerations during late anthesis include continuing restrictions on canopy operations and fungicide applications until flowering completion confirmed, monitoring flower abscission and small berry development assessing fruit set success, beginning preparations for post-bloom operations including disease management and canopy positioning, and evaluating flowering outcomes for yield predictions. The process shows three distinguishable aspects progressing from final flowers opening completing cluster flowering activity, through declining pollen viability reducing late pollination success, to pollination window closure ending reproductive opportunity. Duration typically spans 2-3 days representing concluding phase before complete anthesis, with warm conditions promoting rapid progression while cool conditions may extend period. Cultivar variation appears in proportion of late-opening flowers with some varieties showing compressed flowering with only 10-15% late opening while others show extended 25-35% late opening affecting overall duration, late flower reproductive capacity with some maintaining adequate function while others show substantially reduced capacity, and transition timing to complete anthesis and fruit development. Regional timing shows late anthesis during mid-late June in warm climates, mid-late June in temperate regions, and late June-early July in cool areas, occurring 5-8 days after first flowers opening depending on flowering duration. Viticultural significance includes representing final contribution to fruit set establishing remaining 15-25% berry number, marking transition from critical weather-sensitive flowering period toward more stable fruit development, providing early fruit set assessment through flower abscission and berry visibility, and indicating approaching flowering completion enabling management transitions. Research applications include analyzing factors affecting late flower reproductive capacity, evaluating contributions of late anthesis to final fruit set and berry development patterns, quantifying developmental heterogeneity within clusters affecting ripening uniformity, assessing flower senescence and abscission mechanisms, and documenting transition from reproductive to developmental phases.
Late_Anthesis is third major Anthesis phase following Early and Peak Anthesis, distinguished by declining flowering intensity and closing pollination window. Corresponds to BBCH 67-68 with 70-90% flowers opened. Subdivided into Final_Flowers_Opening, Declining_Pollen_Viability, and Pollination_Window_Closure. Transitions to Complete_Anthesis as final flowers open. Represents final 15-25% fruit set contribution. Weather sensitivity remains significant though reduced from peak. Flower abscission begins indicating fertilization outcomes.
Endblüte
auslaufende Blühphase
concluding anthesis
declining bloom
final bloom
late flowering
späte Anthese
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 380-388 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 292-298 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Late Anthesis
Syrah in Rhône Valley demonstrates late bud swelling at BBCH 03 in mid-April after 850-950 chilling hours and 270-320 growing degree days base 10°C, with buds showing 30-35% volume increase and 1.5-2.0 mm scale separation revealing pale green tissue over 3-4 days under conditions of 15-18°C mean temperature and increasing photoperiod to 13.5 hours, representing typical Mediterranean climate progression for this mid-to-late season cultivar
Late bud swelling represents the final expansion phase before protective scales break open, characterized by maximum volumetric increase with scales widely separated at margins and first green tissue becoming faintly visible through gaps, marking the transition stage between swelling and breaking. This developmental phase occurs after advanced swelling has established prominent expansion, with buds reaching 25-35% enlargement beyond winter dimensions as internal shoot primordium and leaf primordia achieve sufficient size to mechanically force scale separation. The bud profile shows maximum rounded swelling with scale tips spread apart creating gaps of 1-2 mm where pale green shoot tissue can be glimpsed without yet emerging beyond protective scales, representing the culmination of turgor-driven expansion before structural breaking occurs. Scale separation extends from tip toward base as internal pressure increases, with brown outer scales maintaining structural integrity while being mechanically pushed apart by expanding internal tissues rather than actively growing or retracting. The underlying shoot primordium reaches 3-5 mm length with first leaf primordia 2-3 mm in size, both expanding rapidly through coordinated cell division in meristematic zones and cell enlargement through water uptake creating turgor pressure of 0.3-0.8 MPa. Metabolic activity reaches peak levels for bud burst period with respiration rates 15-20 fold above winter baseline, consuming stored carbohydrates at maximum rate before photosynthetic leaves emerge to provide new carbon sources. Cell division rates peak in apical and axillary meristems with cell cycle duration reduced to 18-24 hours compared to days or weeks during early development, producing new cells that will form shoot axis, leaves, tendrils and inflorescence structures. Vascular connection between bud and supporting cane achieves full functionality with mature xylem vessels conducting water at rates sufficient to support rapid expansion and phloem transporting stored reserves mobilized from woody tissues to support growth. Water content in bud tissues reaches 65-75% fresh weight approaching levels found in actively growing tissues, requiring continuous water uptake to maintain turgor driving expansion and prevent wilting. Protein synthesis operates at maximum capacity producing structural proteins for new cells, enzymes supporting rapid metabolism, and regulatory proteins coordinating developmental progression toward breaking and shoot emergence. Hormonal regulation shows peak cytokinin and gibberellin concentrations promoting cell production and expansion, with auxin gradients establishing apical dominance and organ differentiation patterns in developing shoot. Temperature responsiveness continues with development rate doubling per 10°C increase between 10-25°C, making progression timing highly dependent on spring temperature patterns and variability. The process requires sustained favorable conditions with temperatures consistently above 12°C for 2-4 days to complete late swelling, with cooler conditions extending duration but warm conditions above 20°C accelerating transition to breaking. Cultivar variation in late swelling duration reflects genetic differences in force required to separate scales and size threshold triggering breaking, with some varieties showing brief late swelling of 2-3 days while others maintain this stage for 4-6 days. Regional timing follows advanced swelling by 4-7 days, occurring mid-April in warm climates, late April in temperate areas, and early May in cool regions. Viticultural significance includes marking final opportunity for completing dormant-season operations before active growth initiates, establishing critical reference for frost protection deployment as tissue vulnerability reaches maximum before breaking, and serving as timing reference for early-season pest management applications targeting newly active insects. Frost damage at late swelling causes complete primary bud mortality through ice formation in highly hydrated tissues with maximum water content, necessitating total reliance on secondary or basal buds resulting in delayed development and reduced crop potential. Research applications include microscopic documentation of cellular and tissue changes preceding shoot emergence, molecular profiling of gene expression patterns transitioning from dormancy to active growth, and biomechanical analysis of forces involved in scale separation and shoot emergence from protective structures.
Late_Bud_Swelling is the third and final Bud_Swelling subphase, distinguished by maximum expansion with wide scale gaps and faintly visible green tissue that has not yet emerged beyond scales. Transitions directly to Bud_Breaking when green tissue actively emerges. Represents most frost-vulnerable stage of entire bud burst sequence. Brief duration makes field observation timing-dependent.
Vorbruchschwellung
final bud swelling
maximale Knospenquellung
maximum bud expansion
pre-breaking swelling
späte Knospenschwellung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 170-175 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Schultz HR (2000) Climate change and viticulture: A European perspective on climatology, carbon dioxide and UV-B effects, Aust J Grape Wine Res 6:2-12
Late Bud Swelling
Mourvèdre exhibits late budbreak disposition, reaching BBCH 09 approximately 14-21 days after the cultivar median in multi-variety trials, with heat sum requirement of 130-170 GDD (base 10°C) post-dormancy release. Carignan and Grenache show similarly delayed budbreak, emerging within the last 20% of V. vinifera cultivars. Late budbreak in Mourvèdre at southern Rhône reference sites occurs at mean date of April 25 versus April 5 for early cultivars like Chardonnay. Spring frost damage in late cultivars occurs in fewer than 5% of years compared to 15-25% for early cultivars in the same continental vineyard sites.
A budbreak timing disposition that is the inherited capacity of a Vitis vinifera plant to initiate budbreak later than average for the species under given environmental conditions, typically within the third tercile of cultivar budbreak dates, requiring higher heat accumulation following dormancy release and providing natural frost avoidance.
Late budbreak disposition provides natural frost protection but requires adequate growing season heat to achieve full ripeness, restricting late cultivars to warm climate regions or favorable mesoclimates. The trait results from higher thermal thresholds for meristem reactivation and/or greater heat accumulation requirement following dormancy release. Late budbreak may compress the interval to flowering, as both events converge toward optimal temperature ranges. Some late cultivars also show late ripening, but the correlation is imperfect (r = 0.6-0.7), with some late-breaking cultivars ripening at intermediate timing.
Spätaustrieb
delayed budbreak
late budburst
late spring phenology
später Austrieb
spätes Knospentreiben
Tomasi D et al. (2011) Grapevine phenology and climate change: relationships and trends in the Veneto region of Italy for 1964-2009. American Journal of Enology and Viticulture 62:329-339 Webb LB et al. (2012) Observed trends in winegrape maturity in Australia. Global Change Biology 18:3055-3067
Late Budbreak Disposition
Quantitative assessment of Late_Budbreak_Frost_Avoidance_Disposition at the University of British Columbia analyzed budbreak timing across 35 cultivars over 10 seasons, correlating phenology with spring frost damage records and characterizing the heat unit requirements underlying budbreak timing differences. Budbreak date (50% green tip) ranged from April 2 ('Chasselas', earliest) to April 22 ('Mourvèdre', latest), representing 20-day spread among commercial cultivars. Heat unit accumulation (base 10°C) from January 1 to budbreak showed strong cultivar differentiation, ranging from 85 degree-days ('Chasselas') to 165 degree-days ('Mourvèdre'). Late_Budbreak_Frost_Avoidance_Disposition effectiveness was validated by frost damage records. Over the 10-season study period, the site experienced frost events (< -2°C) after April 1 in 7 seasons, with last frost date ranging from April 5 to April 25. Early-budbreaking 'Chasselas' (April 2 mean budbreak) experienced damaging frost exposure in 6 of 7 frost years, while late-budbreaking 'Mourvèdre' (April 22) experienced exposure in only 2 of 7 years. Cumulative frost damage index (sum of damage severity across years) showed strong correlation with budbreak timing (r = -0.78). However, the trade-off with season length was evident, as 'Mourvèdre' failed to achieve adequate fruit maturity (< 20° Brix) in 3 of 10 seasons due to insufficient heat accumulation, while early-budbreaking cultivars consistently ripened. These findings demonstrate that Late_Budbreak_Frost_Avoidance_Disposition provides effective frost protection but requires balancing against season length constraints.
A Late_Budbreak_Frost_Avoidance_Disposition is a Spring_Frost_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through genetically determined late timing of spring budbreak, reducing the probability that vulnerable developing shoots will be present during frost events. This disposition represents a phenological avoidance strategy, with late-budbreaking cultivars escaping spring frost damage by maintaining dormant, cold-hardy buds through the period of highest frost risk rather than through enhanced tolerance of actively growing tissues. Late budbreak timing is controlled by the interaction of chilling requirement satisfaction (release from endodormancy), heat unit accumulation requirement for budbreak (ecodormancy release and growth initiation), and base temperature thresholds for development, with late cultivars typically requiring higher accumulated heat units before visible budbreak. The effectiveness of late budbreak as a frost avoidance strategy depends on regional frost climatology, with greatest benefit in climates where the frost-free date is predictable and late budbreak reliably positions growth initiation after this date. In regions with erratic spring frost patterns, late budbreak provides statistical but not absolute protection, reducing frost encounter frequency rather than eliminating risk. Late_Budbreak_Frost_Avoidance_Disposition may involve trade-offs with season length utilization, as late budbreak compresses the growing season and may delay harvest, potentially conflicting with autumn frost avoidance or optimal fruit maturity in short-season climates.
Late_Budbreak_Frost_Avoidance_Disposition is_a Spring_Frost_Tolerance_Disposition. Late budbreak represents phenological escape from frost rather than enhanced tissue tolerance. Budbreak timing is determined by heat unit requirements after chilling satisfaction, with late cultivars requiring more accumulated warmth. Late budbreak provides statistical frost avoidance proportional to the phenological delay. Trade-offs exist with season length utilization, as late budbreak compresses the growing season and may delay maturity.
Phänologische-Frostvermeidung
Späte-Austriebs-Frostvermeidungs-Disposition
Verzögerter-Austrieb
late bud development
late phenology frost escape
Duchêne E, Schneider C (2005) Grapevine and climatic changes: a glance at the situation in Alsace, Agronomy for Sustainable Development 25:93-99 Jones GV, Davis RE (2000) Climate influences on grapevine phenology, grape composition, and wine production and quality for Bordeaux, France, American Journal of Enology and Viticulture 51:249-261 Zapata C et al. (2017) Predicting key phenological stages for 17 grapevine cultivars, American Journal of Enology and Viticulture 68:60-72
Late Budbreak Frost Avoidance Disposition
Cabernet Sauvignon late developing berries progress from early veraison at BBCH 83 (week 11, 10.5 mm diameter, 1.4 g fresh weight, 15% sugars, 17 g/L acidity, 1.3 mg/g skin anthocyanins) through mid-ripening at BBCH 85 (week 14, 12.5 mm, 2.1 g, 20% sugars, 10 g/L acidity, 2.4 mg/g anthocyanins) to mature harvest-ready berries at BBCH 89 (week 17-18, 14 mm, 2.6 g, 25% sugars, 6.2 g/L acidity, pH 3.75, 3.2 mg/g anthocyanins), demonstrating progression during late development achieving final size, optimal sugar=acid balance, and deep color suitable for premium red wine production. Chardonnay white wine cultivar late developing berries show progression from BBCH 83 (week 10, 11.5 mm, 1.6 g, 15% sugars, 15 g/L acidity) through BBCH 87 (week 14, 13 mm, 2.1 g, 21% sugars, 8 g/L acidity) to maturity BBCH 89 (week 16, 13.5 mm, 2.3 g, 23% sugars, 6.8 g/L acidity, pH 3.35), achieving optimal composition for high-quality white wine with moderate sugar levels and balanced acidity. Syrah late developing berries examined at weekly intervals show cell expansion driving berry growth= week 11 (cells 82 μm diameter, berry 11 mm), week 13 (cells 112 μm, berry 13.5 mm), week 15 (cells 138 μm, berry 15 mm), week 17 (cells 155 μm, berry 16 mm), demonstrating that late development berry enlargement results entirely from cell expansion of fixed cell population, with cell diameter increasing 89% (volume increasing 575%) while berry diameter increases 45% (volume increases 204%) with the difference reflecting increasing intercellular spaces and changing cell packing geometry. Merlot late developing berries under two irrigation regimes show well-watered treatment producing larger berries (15.5 mm, 3.1 g at maturity, 22% sugars, 7.5 g/L acidity) versus moderate deficit producing smaller concentrated berries (13.5 mm, 2.2 g, 24% sugars, 6.2 g/L acidity), demonstrating water availability effects during late development on final berry size and composition concentration with deficit creating smaller berries with enhanced concentration potentially improving wine quality.
A late developing berry is a developing berry in the final pre-maturity ripening phase (BBCH 83-89, approximately 11-18+ weeks post-fruit set) undergoing Phase II ripening growth characterized by rapid berry expansion through cell enlargement, continued sugar accumulation, acid decline, secondary metabolite accumulation, and softening, representing berries approaching harvest maturity and completing the developmental program. Late developing berries are characterized by= (1) rapid size increase from veraison dimensions (typically 9-13 mm diameter) to mature dimensions (typically 12-25 mm depending on cultivar), representing 25-100% diameter increase during late development (2) fresh weight increase from veraison weight (1-2 g) to mature weight (1.5-7 g depending on cultivar), representing 50-300% weight increase achieved primarily through water accumulation and cell expansion (3) cells undergoing dramatic expansion from veraison size (60-90 μm diameter) to mature size (100-180 μm diameter), representing 65-200% diameter increase (280-800% volume increase per cell) driving berry enlargement (4) continued color development with anthocyanins (red/black cultivars) increasing from veraison levels (0.8-2 mg/g skin) to mature levels (1.5-4 mg/g skin or higher in deeply colored cultivars), or deepening golden-yellow in white cultivars (5) progressive softening from veraison firmness (2-3.5 N) to mature softness (0.8-2 N) through continued cell wall modifications and turgor changes (6) sugar accumulation continuing at moderate-to-high rates (0.6-1.2% per week) increasing from mid-veraison levels (13-17%) to mature levels (18-28% depending on cultivar and vintage) (7) acid decline continuing through malic acid respiration and dilution, decreasing from veraison levels (15-20 g/L) to mature levels (4-10 g/L), with pH increasing from 3.2-3.6 at veraison to 3.4-4.2 at maturity. Late developing berries undergo final quality-determining processes= secondary metabolite accumulation including anthocyanins, tannins (condensed tannins/proanthocyanidins in skin and seeds), flavor precursors, and aromatic compounds that critically determine wine quality textural changes including skin softening and flesh softening creating mature berry texture seed maturation completing with embryo desiccation and final hardening preparing seeds for dormancy compositional balance achievement determining optimal harvest timing when sugar=acid ratio, flavor development, and phenolic maturity align with production goals. Late developing berries show changing growth dynamics= early late development (BBCH 83-85, weeks 11-14) maintaining rapid expansion at 0.15-0.4 mm diameter per week and 0.15-0.35 g per week mid late development (BBCH 85-87, weeks 14-16) showing declining growth rates at 0.08-0.2 mm per week and 0.08-0.15 g per week final late development (BBCH 87-89, weeks 16-18+) approaching growth cessation with <0.05 mm per week as maturity is achieved. Environmental factors during late development critically affect final quality= temperature influences sugar accumulation rate and flavor development with moderate temperatures (20-28°C day) optimal water availability affects berry size and composition with moderate deficit concentrating composition versus excess water diluting composition light exposure enhances skin color and phenolic development in exposed berries.
Late_Developing_Berry represents Phase II ripening (BBCH 83-89, weeks 11-18+) and is distinguished from Mid_Developing_Berry (lag phase and veraison, weeks 6-11) by active ripening and expansion versus lag phase stasis and veraison initiation. Late developing berries complete the developmental program achieving harvest maturity, making this the critical period for quality development through compositional changes determining wine/grape characteristics. Sub-classification into Pre_Veraison_Late (somewhat contradictory terminology suggesting late berries before veraison, potentially referring to late-developing lag phase) and Maximum_Size_Late (final expansion phase) reflects progression, while Cell_Division and Cell_Enlargement classifications provide alternative developmental framework. Understanding late berry development is essential for harvest timing, irrigation management, and quality optimization.
Reifephase-Beere
Reifungs-Stadium
ripening berry
spät entwickelnde Beere
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 196-245 Coombe, B.G., McCarthy, M.G. (2000). Dynamics of grape berry growth. Australian Journal of Grape and Wine Research 6=131-135 Matthews, M.A., Anderson, M.M. (1988). Fruit ripening in Vitis vinifera= responses to seasonal water deficits. American Journal of Enology and Viticulture 39=313-320.
Late Developing Berry
Mourvèdre exhibits late flowering disposition, reaching 50% capfall approximately 10-18 days after the cultivar median in comparative trials, with budbreak-to-flowering thermal requirement of 480-580 GDD (base 10°C). Grenache and Carignan show similarly delayed flowering, completing anthesis within the last 25% of V. vinifera cultivars. Late flowering in Grenache at southern Rhône reference sites occurs at mean date of June 20-28 versus June 1-8 for early-flowering Chardonnay. The extended pre-flowering period in late cultivars correlates with more extensive inflorescence development, often resulting in larger flower numbers per cluster (300-500 versus 150-250 in early cultivars).
A flowering timing disposition that is the inherited capacity of a Vitis vinifera plant to initiate and complete flowering later than average for the species, typically within the third tercile of cultivar bloom dates, requiring greater thermal time accumulation from budbreak to anthesis.
Late flowering disposition typically positions bloom during warmer, more stable weather conditions, potentially improving fruit set reliability in Mediterranean climates. However, the compressed post-flowering period may challenge complete ripening in cool years or short-season regions, making late flowering cultivars better suited to warm climate viticulture. Late flowering shows weaker correlation with late ripening (r = 0.5-0.6) than might be expected, as post-flowering development rates vary independently among cultivars.
Spätblüte
delayed flowering
late anthesis trait
late bloom
späte Blüte
später Blühtermin
Parker AK et al. (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17:206-216 Sadras VO, Moran MA (2012) Elevated temperature decouples anthocyanins and sugars in berries of Shiraz and Cabernet Franc. Australian Journal of Grape and Wine Research 18:115-122
Late Flowering Disposition
Riesling bears a Late_Harvest_Scion_Role in Germany producing Spätlese (minimum 76-90°Oechsle depending on region) and Auslese (minimum 83-105°Oechsle) wines where extended ripening concentrates sugars while preserving the variety's exceptional acidity (10-14 g/L), yielding wines of 50-120 g/L residual sugar with intense apple, apricot, and slate mineral character. Gewürztraminer serves in a Late_Harvest_Scion_Role for Alsace Vendange Tardive (minimum 243 g/L potential sugar), with late harvest intensifying terpene aromatics (lychee, rose, ginger) and residual sugar of 30-100 g/L balanced by lower acidity (5-7 g/L). Chenin Blanc bears a Late_Harvest_Scion_Role in the Loire (Coteaux du Layon, Vouvray moelleux), with hang-time concentration producing wines of quince, honey, and lanolin character.
A dessert wine scion role that inheres in a grafted scion cultivar by virtue of its designation for late harvest (vendange tardive/Spätlese) sweet wine production, where extended hang-time on the vine concentrates sugars through evaporation and physiological dehydration without noble rot infection or freezing. Late harvest scions must achieve physiological ripeness early enough to permit extended post-ripening on the vine (typically 2-6 weeks beyond normal harvest), maintain berry integrity during extended hang-time (resisting bunch rots, bird damage, and shriveling beyond wine-making utility), and develop appropriate acidity profiles to balance elevated sugar levels. Late harvest concentration typically achieves must weights of 100-150°Oechsle (approximately 200-350 g/L sugar) compared to 80-100°Oechsle for dry wine production. The style produces wines with intense primary fruit character distinct from the botrytis-transformed complexity of noble rot wines, with residual sugar typically 50-150 g/L balanced by varietal acidity.
Late_Harvest_Scion_Role captures a concentration method relying purely on extended vine retention and evaporative water loss, without the biological transformation of botrytis or physical concentration of freezing or drying. The style depends on climatic conditions permitting extended hang-time—warm, dry autumn weather for continued concentration without rot pressure. Late harvest wines emphasize primary fruit intensity and varietal expression, distinguishing them from the caramelized, honey complexity of botrytis wines. Climate change is complicating late harvest production in traditional regions by accelerating ripening (reducing concentration opportunity) while creating new opportunities in cooler regions previously unable to achieve sufficient ripeness. Regulatory definitions (German Prädikat, Alsace VT) establish minimum must weights but do not mandate specific residual sugar levels.
Spätlese grape
Spätlesesorte
Spätlesetraube
Vendange-tardive-Sorte
extended hang-time variety
late harvest variety
vendange tardive variety
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Johnson H, Robinson J (2013) The World Atlas of Wine 7th Ed, Mitchell Beazley Jackson RS (2014) Wine Science 4th Ed, Academic Press pp.: 530-545 Reinhardt G (2019) German Wine, Infinite Ideas
Late Harvest Scion Role
A late juvenile unbranched tendril at node 10 on a shoot of Vitis vinifera cv. Riesling in Mosel Valley, Germany, measuring 9.5 cm length at 13 days post-emergence, showing light tan coloration in basal 3.5 cm indicating beginning lignification while distal 6 cm remains bright green, 2.3 mm diameter at mid-length reflecting advanced but incomplete structural development, growing at 4 mm per day indicating transition from maximum juvenile growth rates toward maturation, vascular bundles clearly visible through semi-translucent epidermis with secondary xylem elements developing in basal regions, epidermis showing slight glossy appearance from early cuticle formation, exhibiting weak touch response with delayed coiling initiation (15-20 minutes) when experimentally contacted indicating developing mechanosensory function, occurring on a moderately vigorous shoot in a Riesling clone where 70% of tendrils mature as unbranched structures and this individual shows no signs of lateral meristem activation suggesting it will complete development as simple unbranched architecture, measured at nominal BBCH stage 73 (berries beginning to touch) for mature shoot regions, representing the late juvenile transitional stage approaching but not yet achieving functional maturity.
A juvenile unbranched tendril in the transitional phase between early rapid growth and final maturation in Vitis vinifera, characterized by intermediate size typically 6-11 cm length representing 40-70% of eventual mature size, elapsed developmental time of 8-18 days post-emergence indicating progression beyond initial growth phase but not yet achieving maturity, slowing extension growth with daily elongation rates declining from maximum early rates to 3-6 mm per day as maturation processes begin, partial tissue differentiation including initiation of vascular secondary wall development, beginning epidermis maturation with early cuticle deposition creating slight surface sheen, early mechanosensory cell differentiation with weak or inconsistent touch response indicating developing but immature sensory function, beginning structural reinforcement with initial lignification appearing as light tan coloration in basal regions while distal portions remain entirely green, and representing the critical developmental period when branching decisions are finalized with lateral meristems either activating to produce branches or remaining quiescent resulting in mature unbranched architecture. Late juvenile unbranched tendrils occur in all tendril developmental trajectories as the transitional stage between rapid early growth and final maturation, and represent the developmental window during which cultivar-specific branching patterns are expressed, environmental conditions exert major influence on final architecture, and resource availability determines whether development proceeds to functional maturity or aborts due to insufficient metabolic support for completion.
Late juvenile unbranched tendrils are distinguished from early juvenile tendrils by larger size (>5-6 cm), longer elapsed development time (>7-8 days), beginning tissue differentiation and maturation processes, and from mature simple unbranched tendrils by incomplete development with ongoing growth and immature functional status. This developmental stage represents the critical branching decision period when genetic programs and environmental conditions determine whether tendrils remain unbranched or activate lateral meristems, with branching typically initiated 7-14 days post-emergence in cultivars with branching tendencies. Assessment of late juveniles provides predictive information about final architecture, with presence or absence of lateral meristem swelling indicating probable branched versus unbranched outcomes. Late juvenile stage shows moderate abortion rates (8-15% in moderately stressed conditions) lower than early juvenile vulnerability but higher than mature tendril stability. Resource management during late juvenile stage determines whether development completes successfully to functional maturity or arrests producing aborted tendril outcomes.
intermediate juvenile tendril
pre-mature developing tendril
spät-juvenile Ranke
transitional juvenile tendril
vor-reife entwickelnde Kletterorgan
übergangs-juvenile Ranke
Based on Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 86-134 Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 74-104 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 237-258.
Late Juvenile Unbranched Tendril
Cabernet Sauvignon late post-veraison berries at weeks 15-18 post-fruit set (4-7 weeks post-veraison, BBCH 85-89) show progression toward maturity= week 15 (13.5 mm, 2.5 g, cells 142 μm, 21.5% sugars, 8.5 g/L acidity, 3.2 mg/g anthocyanins), week 16 (14 mm, 2.65 g, cells 152 μm, 23% sugars, 7.2 g/L acidity, 3.5 mg/g anthocyanins), week 17 (14.3 mm, 2.75 g, cells 158 μm, 24.5% sugars, 6.3 g/L acidity, 3.6 mg/g anthocyanins), week 18 (14.5 mm, 2.8 g, cells 162 μm, 25.5% sugars, 5.8 g/L acidity, 3.65 mg/g anthocyanins, harvest), demonstrating slowing changes during late post-veraison with sugar accumulation declining from 1.5% per week (week 15) to 1% per week (week 18) as maturity approaches. Pinot Noir late post-veraison berry harvest timing decision= monitoring weeks 16-18 shows week 16 (under-ripe= 22% sugars, 8.2 g/L acidity, green flavors), week 17 (approaching optimal= 23.5% sugars, 7 g/L acidity, developing fruit character), week 17.5 (optimal= 24.2% sugars, 6.8 g/L acidity, balanced ripe fruit), week 18.5 (beginning over-ripe= 25.5% sugars, 5.8 g/L acidity, cooked notes), with harvest executed at week 17.5 capturing optimal ripeness narrow window during late post-veraison period. Chardonnay late post-veraison berry vintage comparison= cool vintage requiring 8 weeks veraison-to-harvest (week 10 veraison, week 18 harvest at 21.5% sugars, 7.8 g/L acidity) warm vintage requiring only 5 weeks (week 10 veraison, week 15 harvest at 22% sugars, 7.2 g/L acidity), demonstrating vintage effects on late post-veraison ripening rate and harvest timing variability driven by temperature and heat accumulation differences. Merlot late post-veraison berry weather risk management= forecast showing 30 mm rain at week 17 (berries at 25% sugars, 5.8 g/L acidity, slightly past optimal), prompting immediate harvest at week 16.5 (24.5% sugars, 6.3 g/L acidity, acceptable though not optimal) to avoid rain-induced splitting and rot, demonstrating late post-veraison harvest decisions balancing composition optimization versus weather risk avoidance. Syrah late post-veraison berry phenolic maturity assessment= weekly seed chewing and skin tasting during weeks 15-18 tracking tannin ripeness progression from astringent green character (week 15) through developing softness (week 16) to optimal soft ripe tannins (week 17), with sensory phenolic maturity converging with analytical sugar maturity (25% sugars) at week 17 defining optimal harvest timing integrating compositional and sensory maturity during late post-veraison period.
A late post-veraison berry is a post-veraison berry in the final portion of the post-veraison ripening period approaching or achieving maturity (BBCH 85-89, approximately 4-9+ weeks post-color change), characterized by slowing or completing ripening changes, sugars approaching or at target levels, acids at mature levels, color development complete or plateaued, and compositional maturation determining harvest timing, representing berries in late Phase II ripening approaching harvest readiness. Late post-veraison berries are characterized by= (1) color development complete or nearly complete with anthocyanins (red cultivars) at 2.5-5+ mg/g approaching or at maximum levels, or full golden-yellow color (white cultivars) (2) sugars at advanced levels typically 20-28% approaching or at target maturity levels for intended use (3) acids at low levels typically 4-9 g/L having declined substantially from veraison peak (17-20 g/L) (4) cells approaching or at maximum expansion with cell diameter 120-180 μm representing final or near-final size (5) berry diameter approaching or at maximum typically 12-25 mm depending on cultivar (6) berry fresh weight approaching or at maximum typically 1.8-7 g depending on cultivar (7) texture soft with firmness 0.8-2 N approaching or at mature levels. Late post-veraison berries show declining or completed ripening changes= sugar accumulation rate declining from early post-veraison maximum (0.8-1.5% per week) to late ripening moderate or low rates (0.2-0.6% per week) or approaching zero at maturity cell expansion rate declining from maximum (6-12 μm per week) to low rates (<4 μm per week) or zero acid decline continuing but slowing as malic acid becomes depleted anthocyanin accumulation plateauing or continuing slowly flavor maturation progressing with development of complex aromatic profile. Late post-veraison berries represent the harvest decision period= compositional monitoring tracking progression toward target maturity sensory evaluation assessing flavor development and phenolic maturity integration of analytical and sensory data determining optimal harvest timing balancing competing objectives (adequate ripeness versus avoiding overripeness, optimal composition versus practical harvest constraints) guiding harvest decisions. Late post-veraison berries show vintage and site variation in maturation timing= cool vintages and cool sites showing slower late ripening requiring extended late post-veraison period (6-9+ weeks veraison to harvest) warm vintages and warm sites showing rapid late ripening achieving maturity in 4-6 weeks heat accumulation (growing degree-days) during late post-veraison largely determining final maturity timing and harvest date variability. Late post-veraison berries are subject to potential quality risks= overripening if harvest delayed past optimal timing dehydration and concentration in warm dry conditions rot development (Botrytis, bunch rots) if wet conditions occur autumn frosts potentially damaging late-maturing cultivars these risks create harvest timing pressures balancing quality optimization against risk avoidance. Late post-veraison berry management focuses on harvest timing= regular sampling (weekly or biweekly) monitoring compositional evolution sensory evaluation tracking flavor and phenolic maturity weather monitoring considering harvest logistics relative to forecast conditions harvest execution when optimal maturity achieved or when quality risks become unacceptable.
Late_Post_Veraison_Berry represents final ripening phase (4-9+ weeks post-veraison, BBCH 85-89) and is distinguished from Early_Post_Veraison_Berry (0-4 weeks, rapid ripening) by slowing or completing ripening versus active rapid ripening. Late post-veraison represents the harvest decision period when maturity assessment guides harvest timing. Understanding late post-veraison berry development is essential for harvest timing decisions, for maturity monitoring programs, for balancing quality optimization with practical and weather constraints, and for all aspects of harvest management determining final wine or grape quality.
End-Reifephase-Traube
Erntenahe-Traube
final ripening berry
pre-harvest berry
späte nach-Véraison Beere
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 215-250 Jackson, D.I., Lombard, P.B. (1993). Environmental and management practices affecting grape composition. American Journal of Enology and Viticulture 44=409-430.
Late Post Veraison Berry
Cabernet Franc exhibits late ripening disposition, reaching 22-24°Brix approximately 55-65 days post-veraison with total season heat requirement of 1550-1750 GDD (base 10°C) from budbreak. Petit Verdot and Tannat show similar late ripening, completing harvest in the final third of the regional vintage window. Late ripening in Petit Verdot at Bordeaux reference sites occurs at mean date of October 15-28, 3-4 weeks after Merlot in the same appellation. In marginal vintages, late cultivars may fail to achieve full physiological ripeness, with residual green, herbaceous flavors indicating incomplete flavor maturation despite adequate sugar levels.
A ripening timing disposition that is the inherited capacity of a Vitis vinifera plant to reach harvest maturity late in the season, typically within the fourth quartile of cultivar harvest dates for a region, requiring substantial heat accumulation and extended growing seasons for complete maturation.
Late ripening disposition restricts cultivar suitability to warm climates or favorable mesoclimates within moderate regions. The trait provides advantages in very warm climates by extending ripening into cooler autumn conditions, potentially preserving acidity and aromatic complexity that would be lost with earlier-ripening cultivars harvested during peak summer heat. Late cultivars face harvest timing risks from autumn rain, frost, and bird pressure, requiring careful site selection and vintage-specific management decisions.
Spätreife-Disposition
late harvest trait
late maturation
späte Reife
später Erntetermin
tardy ripening
Wolpert JA et al. (1994) Field performance of six Chardonnay clones in the Napa Valley. American Journal of Enology and Viticulture 45:393-400 Robinson J, Harding J (2015) The Oxford Companion to Wine, 4th Ed, Oxford University Press pp.: 567-568
Late Ripening Disposition
Autumn Royal bears a Late_Season_Table_Grape_Role as the dominant late black seedless, reaching commercial maturity (18-20°Brix, full blue-black color) in September-November in California, extending the fresh grape season 4-6 weeks beyond mid-season Thompson Seedless and capturing autumn market demand before Chilean imports arrive. Autumn King serves in a Late_Season_Table_Grape_Role for late white seedless, with very large berries (8-12 g), pale green-yellow color, and harvest extending into November, providing late-season white grape availability when Sugraone and Thompson have concluded. Crimson Seedless bears a Late_Season_Table_Grape_Role in its late harvest window (September-October), with fruit from late-harvested blocks achieving premium color development and extended storage capability compared to early-harvested Crimson.
A table grape scion role that inheres in a grafted scion cultivar by virtue of its late maturation timing, reaching commercial harvest after the main-season varieties and extending fresh grape availability into autumn and early winter when new-crop supply diminishes. Late season table grape scions mature 10-40 days after mid-season benchmarks, reaching harvest-ready condition in September-November in Northern Hemisphere production regions (California, Mediterranean), requiring high heat accumulation (typically 2200-2800 GDD base 10°C) and extended hang-time to achieve full ripeness. Late maturity results from late budbreak, extended flowering-to-veraison interval, long veraison-to-harvest period, or high sugar accumulation requirements, determined by genetic factors controlling developmental timing. Late season varieties offer advantages including reduced early-season frost risk, extended harvest windows reducing labor peaks, and premium pricing when new-crop supply declines before Southern Hemisphere counter-season fruit arrives. Major late season varieties include Crimson Seedless (late-mid to late), Autumn Royal, Autumn King, and Red Globe.
Late_Season_Table_Grape_Role captures varieties extending the fresh grape season beyond peak production, commanding variable pricing depending on new-crop scarcity and competition with stored fruit or counter-seasonal imports. Late varieties face challenges including autumn rainfall risk (disease pressure, berry splitting), cooling temperatures slowing final ripening, and competition with stored early-season and mid-season fruit that may offer lower-cost alternatives. The development of excellent storage varieties (Crimson Seedless storing 6-8 weeks) and efficient counter-seasonal supply chains has reduced late-season premiums by providing year-round fresh grape availability. Late varieties often exhibit enhanced storage potential due to their firmer texture and higher acid retention at harvest, creating overlap with the Long_Storage category.
Spätsorte
autumn grape
late maturing grape
late season grape
late table grape
spät reifende Tafeltraube
späte Tafeltraube
Creasy GL, Creasy LL (2009) Grapes, CABI California Table Grape Commission (2020) Grape Variety Harvest Dates Crisosto CH, Mitchell FG (2002) Postharvest handling systems: table grapes, UC ANR Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press
Late Season Table Grape Role
Crimson Seedless exhibits late seed abortion disposition with embryo collapse occurring 28-42 days post-anthesis, producing seed traces of 25-40 mg and 5-7 mm length with partial lignification of the seed coat. Autumn Royal aborts at intermediate timing (21-35 days) with seed traces of 20-30 mg. Late abortion cultivars benefit from extended hormone signaling, producing larger berries (4-6 g without treatment) but requiring careful clone selection to minimize seed trace perceptibility. Seed trace hardness in late-aborting cultivars may reach 2-3 on the 1-5 lignification scale compared to 0.5-1 for early-aborting types.
A stenospermocarpy disposition that is the inherited capacity of a Vitis vinifera plant to abort developing seeds at a later post-fertilization stage (typically 3-6 weeks after anthesis), resulting in larger, sometimes partially lignified seed traces that may be noticeable during consumption.
Late seed abortion produces larger seed traces due to extended endosperm and seed coat development before abortion occurs. The partially developed seed coat may undergo some lignification, creating harder traces that reduce consumer acceptance. Breeding programs select for minimal seed trace size within late-aborting backgrounds to combine larger berry size with acceptable seedlessness. Environmental factors including temperature can shift abortion timing by 5-10 days.
große Samenrudimente
large seed trace formation
late embryo abortion
late stenospermocarpy
späte Embryoabtötung
späte Samenabortion
Ledbetter CA, Burgos L (1994) Inheritance of stenospermocarpic seedlessness in Vitis vinifera L. Journal of Heredity 85:157-160 Cabezas JA et al. (2006) A genetic analysis of seed and berry weight in grapevine. Genome 49:1572-1585
Late Seed Abortion Disposition
A late semi-lignified cane on Vitis vinifera cv. Cabernet Franc examined in mid-November, showing brown woody appearance for basal 105cm (nodes 1-10) with firm structural rigidity, brown-green appearance for middle 20cm (nodes 11-12) with moderate firmness, and green appearance for apical 25cm (nodes 13-15) with retained flexibility, representing near-complete maturation in warm-climate vineyard where additional 3-4 weeks before pruning will likely complete lignification of middle section while apical green portion will be removed regardless, contrasting with similar cane in cool-climate vineyard where approaching freezing temperatures raise concerns about incomplete maturation even of middle section requiring more conservative pruning to retain only fully brown portions.
A semi-lignified one-year cane in advanced stages of lignification typically occurring during late autumn (October-November in Northern Hemisphere), characterized by predominantly brown woody appearance for basal and middle sections with green or brown-green appearance retained only in apical regions, firm woody texture throughout most of length with flexibility remaining in unlignified tips, nearly complete secondary cell wall development in mature sections with ongoing progression in transition zones, and status indicating either normal final-stage maturation (in regions with adequate thermal time) or problematic incomplete ripening (in short-season climates approaching winter onset). Late semi-lignified canes represent critical decision points during autumn assessment and winter pruning, requiring judgment about whether remaining unlignified portions will complete maturation or should be removed to avoid winter damage and ensure fruiting wood quality.
Late semi-lignified canes illustrate the climate dependence of wood maturation timing, with interpretation of their status varying dramatically between warm and cool regions. In Mediterranean or warm-temperate climates, late semi-lignified status in November indicates normal progression with expectation of full maturation by December-January pruning, while in cool-climate or short-season regions, similar status raises concerns about incomplete ripening requiring prompt pruning decisions to remove at-risk portions before winter damage occurs. The judgment about retaining versus removing partially lignified sections depends on local freeze risk (mild winters tolerate retention of brown-green wood, severe winters require conservative removal), cultivar cold hardiness, and economic considerations balancing crop potential against damage risk. Historical tracking of autumn lignification patterns and winter damage outcomes informs site-specific decision protocols.
advanced lignification cane
fast ausgereifte Rute
late maturation cane
nearly ripened cane
spät-teilverholzte Rute
Late Semi Lignified Cane
A late stage reproductive active bud on Vitis vinifera cv. Pinot Noir at BBCH stage 69 in mid-June in Burgundy, France, with shoot bearing 8 fully expanded leaves, 2 inflorescences at fruit set stage showing 60% and 55% berry retention from original flower complement of 165 and 148 flowers respectively, early developing berries 3-4mm diameter in cell division phase, representing transition from flower to fruit and establishing final yield potential for this shoot.
A reproductive active bud in advanced developmental stages corresponding approximately to BBCH stages 55-71, characterized by well-developed shoots bearing separated flower clusters with individual flowers visible and distinguishable, proceeding through flowering (anthesis with cap fall and pollen release) to fruit set (berry initiation following successful fertilization), with concurrent vegetative development producing multiple expanded leaves and elongating shoots. Late stage reproductive active buds have inflorescences transitioning from flowers to developing berries, representing the critical period for yield determination as environmental stresses (temperature extremes, water deficit, nutrient limitation) during flowering and fruit set directly impact the number of berries that successfully develop from pollinated flowers.
Late stage reproductive active buds mark the transition from potential to realized yield, as fruit set percentage determines actual berry number from the population of flowers formed. Poor fruit set (coulure) or excessive berry drop (millerandage) during this period can dramatically reduce yields, caused by factors including cool wet weather during flowering, excessive vegetative vigor, boron deficiency, or genetic predisposition in certain cultivars. The end of this stage marks completion of berry number determination, with subsequent yield changes limited to berry size variation, shriveling, or pathological losses.
advanced flowering bud
fruit-setting bud
späte reproduktive aktive Knospe
Based on Ebadi et al. (1996) "Seed development and abortion in Vitis vinifera L., cv. Chardonnay" International Journal of Plant Sciences 157=703-712 Lebon et al. (2004) "Sugars and flowering in grapevine, Vitis vinifera L." Journal of Experimental Botany 59=2565-2578 Keller et al. (2001) "Flowering and fruit set in Vitis vinifera L.= A critical assessment" European Journal of Horticultural Science 66=98-105.
Late Stage Reproductive Active Bud
In a Barossa Shiraz vineyard, late-stage sugar import was quantified over 26 days in late February through March Year N+1 (BBCH 87-89) when calculated hexose import declined from 62 μmol g⁻¹ h⁻¹ (early late-stage) to 32 μmol g⁻¹ h⁻¹ (final), VvHT1 expression decreased from 95-fold to 68-fold, berry Brix increased from 24.2° to 26.8° (2.6° gain decelerating), vacuolar hexose reached 2180 mM combined approaching cultivar maximum, under warm Australian late-summer conditions (daily high declining 32° to 26°C as autumn approached, canopy beginning senescence reducing source capacity).
Late_Stage_Sugar_Import represents the gradually diminishing hexose uptake during late ripening when declining concentration gradients, reduced transporter expression, and approaching vacuolar saturation combine to progressively limit import flux creating the characteristic deceleration that defines final approach to maximum berry sweetness and signals ripening completion. This late-stage import occurs during final accumulation period (BBCH 86-89), typically days 38-62 post-veraison spanning 22-30 days, and is characterized by plasma membrane hexose uptake declining progressively from early late-stage 45-85 μmol combined glucose and fructose g⁻¹ h⁻¹ to final 18-45 μmol g⁻¹ h⁻¹ representing 2-3 fold reduction compared to peak import 100-180 μmol g⁻¹ h⁻¹, transporter gene expression declining from late-stage onset 70-110 fold above green berry to final 50-85 fold indicating downregulation of ripening program, vacuolar hexose concentrations approaching maximum 1700-2500 mM combined creating reduced thermodynamic driving force for continued uptake, and apoplastic-to-vacuolar concentration gradient declining from 15-30:1 (apoplastic 280-550 mM, vacuolar 500-900 mM peak accumulation) to 6-15:1 (apoplastic potentially declining 180-350 mM, vacuolar 1600-2400 mM late stage) reducing facilitated diffusion driving force. Molecular mechanisms limiting late-stage import include hexose transporter gene transcription declining 25-45% from peak levels as ripening program downregulates indicated by VvHT1, VvHT2 expression decreasing though remaining elevated above green berry baseline, mRNA stability potentially decreasing with shorter half-lives 5-9 hours versus 8-14 hours peak requiring higher transcription rates to maintain steady-state but transcription declining creating net reduction, protein synthesis rates declining 30-50% from peak as general translation slows with ripening maturation, transporter protein turnover potentially accelerating through ubiquitin-proteasome degradation removing older proteins faster than synthesis replaces causing density decline 20-40% from peak, and potential post-translational modifications (phosphorylation, ubiquitination) reducing activity or targeting degradation though modifications in grape transporters incompletely characterized. Kinetic limitations during late-stage import demonstrate cytoplasmic hexose concentrations potentially rising 75-125 mM (from 60-100 mM peak) as tonoplast transport saturates or vacuolar back-diffusion increases reducing plasma membrane concentration gradient and thus uptake rate even if transporter capacity unchanged, vacuolar hexose 1650-2450 mM approaching saturation reducing tonoplast gradient from vacuole-to-cytoplasm potentially causing slight back-flux 5-15 μmol g⁻¹ h⁻¹ opposing continued uptake, and net import flux (plasma membrane uptake minus tonoplast back-flux minus respiratory consumption) declining to 15-40 μmol g⁻¹ h⁻¹ net vacuolar accumulation representing 20-40% of peak net flux. Thermodynamic constraints include osmotic potential reaching -2.9 to -3.9 MPa (from -2.4 to -3.2 MPa peak) approaching cell tolerance limits potentially triggering regulatory mechanisms preventing further accumulation, water potential berry -2.5 to -3.5 MPa creating only small gradient versus vine xylem -0.8 to -1.5 MPa limiting water import that previously drove berry expansion enabling dilution-free sugar accumulation, turgor pressure declining to 0.2-0.4 MPa (from 0.3-0.6 MPa peak) as osmotic potential increases faster than water import can compensate potentially affecting membrane transport energetics, and approaching thermodynamic equilibrium between apoplastic and vacuolar hexose concentrations accounting for membrane potential and proton gradients reducing maximum achievable gradient limiting import capacity. Substrate availability effects show phloem sucrose delivery potentially declining 30-50% from peak 250-450 mg per berry per day to late-stage 150-280 mg per berry per day as source photosynthesis decreases with canopy senescence and declining photoperiod, apoplastic hexose concentrations potentially declining 35-50% from peak 280-550 mM to late-stage 180-350 mM as invertase substrate (sucrose) reduces, and reduced substrate availability limiting maximum possible uptake independent of transporter capacity creating substrate limitation rather than kinetic limitation particularly late final phase. Metabolic partitioning changes during late-stage import include increased proportion of imported hexoses entering respiratory pathways rising from 20-30% peak to 35-50% late-stage as vacuolar storage becomes thermodynamically less favored and cellular ATP demands persist, reduced net storage efficiency where each μmol hexose imported yields only 0.50-0.65 μmol vacuolar accumulation (late-stage) versus 0.65-0.80 μmol (peak) due to increased respiratory losses, and metabolic feedback where high cytoplasmic hexose potentially inhibits further uptake through hexokinase-mediated signaling creating negative feedback loop limiting excessive accumulation. Environmental modulation of late-stage import shows temperature continuing to affect rates with cooler autumn conditions 18-24°C further reducing already declining import 25-40% versus optimal 26-30°C, declining solar radiation and photoperiod reducing source photosynthesis 30-50% compared to mid-summer limiting substrate delivery independent of sink capacity, water deficit effects complex with moderate deficit potentially enhancing concentration but severe deficit impairing phloem delivery reducing import 30-60%, and canopy health with vigorous healthy vines maintaining higher late-stage import longer while stressed or nutrient-deficient vines showing earlier more severe import decline. Cultivar differences in late-stage import demonstrate high-Brix varieties maintaining higher absolute rates 35-70 μmol g⁻¹ h⁻¹ late-stage enabling final 26-30° Brix achievement, moderate cultivars showing 25-55 μmol g⁻¹ h⁻¹ reaching typical 22-26° levels, and lower-Brix types displaying 18-40 μmol g⁻¹ h⁻¹ achieving 20-24° final reflecting genetic differences in transporter regulation, maximum vacuolar capacity, and metabolic set-points determining when import effectively ceases. Coordination with other processes shows late-stage sugar import declining while acid catabolism continues creating shifting sugar-acid balance, berry water import minimal or ceasing with size stabilization typical late ripening, and flavor-aroma synthesis continuing independently with volatiles accumulating despite declining sugar import indicating decoupled ripening processes. Quality implications include late-stage import rate determining final achievable Brix with early cessation potentially limiting maximum sugar problematic for high-alcohol styles, extended import enabling very high final sugars beneficial for some styles but requiring careful harvest timing avoiding over-ripeness, and import decline signaling approaching physiological maturity useful for harvest timing decisions. Viticultural factors affecting late-stage import include canopy management maintaining photosynthesis though senescence inevitable, irrigation typically minimal or ceased late season potentially beneficial for concentration but excessive deficit impairing phloem function reducing import, and nutritional status with adequate nitrogen supporting continued enzyme synthesis maintaining import longer while deficient vines showing earlier decline. Research methods quantifying late-stage import include flux studies using isotope tracers measuring declining uptake rates, gene expression analysis tracking transporter downregulation, enzyme activity assays revealing capacity changes, and metabolomic profiling characterizing compartment-specific hexose dynamics revealing kinetic and thermodynamic limitations.
Late_Stage_Sugar_Import is distinguished from peak import by showing declining flux (18-85 μmol g⁻¹ h⁻¹ progressive reduction) rather than maximum (100-180 μmol g⁻¹ h⁻¹), from active import by representing deceleration rather than sustained rates, and from cessation by showing continued though diminishing uptake rather than complete stop. Duration 22-30 days BBCH 86-89. Transporter expression declining 25-45% from peak. Thermodynamic and kinetic limitations. Late_Stage_Sugar_Import represents progressive import reduction approaching sugar equilibrium.
abnehmende Hexose-Aufnahme
declining hexose uptake
diminishing sugar import
downregulated sugar transport
herunterregulierter Zucker-Transport
reduced import flux
reduzierter Import-Fluss
vermindernder Zucker-Import
Hayes MA, Davies C, Dry IB (2007) Isolation, functional characterization, and expression analysis of grapevine hexose transporters J Exp Bot 58:1985-1997 transporter regulation Keller M (2020) The Science of Grapevines 3rd Ed pages 226-228 late import Dai ZW, Ollat N, Gomès E, Decroocq S, Tandonnet JP, Bordenave L, Pieri P, Hilbert G, Kappel C, van Leeuwen C, Vivin P, Delrot S (2011) Ecophysiological, genetic, and molecular causes of variation in grape berry weight and composition Plant Physiol 155:519-532 late-stage dynamics.
Late Stage Sugar Import
A late stage vegetative active bud on a basal node of Vitis vinifera cv. Merlot in mid-May in Pomerol, France, having developed into a shoot with 6 fully expanded leaves (total leaf area ~450 cm²), 3 developing leaves, 2 functional tendrils, 15cm of shoot elongation, beginning net carbon assimilation exceeding respiratory costs, with basal 2-3 internodes showing early lignification and green-brown coloration indicating secondary growth initiation.
A vegetative active bud in advanced stages of shoot development, corresponding approximately to BBCH stages 10-19, characterized by multiple fully expanded leaves (typically 3-8 leaves), visible shoot elongation with measurable internodes, active tendril development, continued leaf emergence from shoot apex, and transition from heterotrophic (consuming stored reserves) to autotrophic (net photosynthetic) metabolism. Late stage vegetative active buds have developing shoots with increasing lignification of basal internodes, moderate frost tolerance (-1°C to 0°C for brief periods), established vascular connections to permanent vine structure allowing efficient water and nutrient transport, and substantial carbon assimilation capacity contributing to whole-vine photosynthetic production and reserve accumulation.
Late stage vegetative active buds represent established shoot systems making significant contributions to vine carbon economy and canopy development. Management decisions at this stage focus on shoot positioning, shoot thinning to optimize leaf area-to-fruit ratios, and pest/disease management as expanding leaf area creates favorable microclimates for pathogens. The transition from early to late stage marks a shift from vulnerability (frost risk, herbivory) to productivity, though shoots remain susceptible to wind damage, hail, and certain fungal pathogens until full lignification occurs in late summer.
advanced stage vegetative bud
developed vegetative shoot bud
späte vegetative aktive Knospe
Based on Poni et al. (2006) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 57=397-407 Palliotti et al. (2011) "Changes in vineyard establishment and canopy management urged by earlier climate-related grape ripening= A review" Scientia Horticulturae 178=43-54 Intrieri et al. (2008) "Effects of canopy manipulations on whole-vine photosynthesis= Results from pot and field experiments" Vitis 47=159-165.
Late Stage Vegetative Active Bud
A late winter dormant bud on Vitis vinifera cv. Pinot Noir in late February in Burgundy, having completed chilling requirement of 850 hours, showing slight bud scale separation and green tissue visible at the apex, responding to forcing conditions within 10-14 days, with cold hardiness reduced to -15°C compared to -25°C during deep winter dormancy, vulnerable to catastrophic damage from late spring frosts.
A winter dormant bud in the final phase of endodormancy during late winter to early spring (typically February-March in Northern Hemisphere), having accumulated sufficient chilling hours to satisfy cultivar-specific requirements but not yet exposed to adequate heat accumulation for budbreak, characterized by progressively increasing metabolic activity, swelling bud scales, declining cold hardiness, and responsiveness to forcing conditions. Late winter dormant buds represent the transition from endodormancy to ecodormancy, where growth resumption becomes limited by environmental factors (temperature, moisture) rather than internal physiological blocks, making buds increasingly vulnerable to spring freeze damage as hardiness diminishes.
The transition from deep to late winter dormancy is gradual and cultivar-dependent, with early-budding varieties entering late dormancy earlier than late-budding ones. Late winter dormant buds pose significant management challenges in viticulture, as warm periods can trigger premature budbreak followed by devastating spring freeze damage. Viticulturists monitor Growing Degree Day (GDD) accumulation during this period to predict budbreak timing and implement frost protection strategies.
ecodormant bud
late dormancy bud
pre-budbreak dormant bud
späte Winterruhe Knospe
Based on Londo & Johnson (2014) "Variation in the chilling requirement and budburst rate of wild Vitis species" Environmental and Experimental Botany 106=138-147 Parker et al. (2013) "General phenological model to characterize the timing of flowering in Vitis" Australian Journal of Grape and Wine Research 19=126-135 Molitor et al. (2014) "Late frost damage risk for viticulture under future climate conditions= a case study for the Luxembourgish winegrowing region" Australian Journal of Grape and Wine Research 20=160-168.
Late Winter Dormant Bud
Merlot grafted on SO4 rootstock ring 12 showing late wood rootstock xylem comprising outer 0.28 cm of 0.58 cm total ring width (48% of ring), latewood appearing darker brown contrasting with lighter earlywood, latewood vessels 50-70 micrometers diameter significantly smaller than earlywood vessels 95-125 micrometers, latewood showing thick fiber cell walls and reduced lumens optimized for mechanical strength, latewood density estimated 0.64 g/cm³ versus earlywood 0.45 g/cm³, latewood zone contributing approximately 30% of ring's water transport capacity but providing majority of ring's structural strength
Late wood rootstock xylem is the xylem tissue produced during late growing season typically July-September in Northern Hemisphere forming outer darker-colored portion of annual ring, characterized by small-diameter vessels 40-80 micrometers prioritizing mechanical strength over maximum conductivity, thick-walled cells minimizing lumen volume while maximizing structural support, higher wood density than earlywood due to abundant wall material and reduced void spaces, slower formation during declining cambial activity as growing season concludes, comprising typically 40-60% of annual ring width, and providing disproportionate contribution to wood mechanical strength despite conducting less water than earlywood due to dense thick-walled fiber cells creating rigid durable tissue essential for trunk structural integrity. Late wood rootstock xylem formation timing includes= initiated in mid-season (cambium shifting to latewood production July as earlywood phase concludes), produced during late summer (majority of latewood formed July-September during declining cambial activity), corresponding to ripening (latewood formation coinciding with veraison, berry maturation, and canopy maturation when water demand declining), showing reduced production rate (cambium dividing more slowly during latewood phase producing fewer cells than earlywood phase), and completing annual ring (latewood formation ceasing September-October as cambium enters dormancy finishing annual ring before winter). Late wood rootstock xylem structural characteristics includes= showing small vessel diameter (vessels 40-80 micrometers significantly smaller than earlywood vessels 80-180 micrometers prioritizing strength over maximum conductivity), exhibiting thick cell walls (vessel elements and especially fiber cells with extensive secondary wall deposition minimizing lumen, wall ratio), displaying high wood density (latewood 0.55-0.75 g/cm³ higher than earlywood 0.35-0.50 g/cm³ due to thick walls and reduced lumens), appearing dark colored (latewood darker tan or brown in cross-section due to abundant wall material and small void spaces), and showing boundary with next ring (latewood outer edge marking annual ring boundary transitioning to next year's earlywood creating visible ring line). Late wood rootstock xylem vessel characteristics includes= containing smallest vessels in ring (latewood vessels reaching minimum diameter for functional water transport reflecting shift from conductivity to strength priority), showing moderate vessel density (adequate vessels for continued water transport but fewer per area than earlywood), exhibiting functional perforation plates (openings adequate for water flow but contributing relatively less to total conductivity than earlywood vessels), displaying necessary pit connections (lateral pits maintaining connectivity though latewood contributes less to overall lateral transport than earlywood), and demonstrating cavitation resistance (smaller vessels potentially less vulnerable to embolism formation under stress compared to large earlywood vessels due to reduced diameter). Late wood rootstock xylem hydraulic function includes= providing supplementary water transport (latewood conducting 20-40% of total annual water flow contributing to overall trunk capacity but secondary to earlywood), enabling continued transpiration (latewood transport adequate for reduced late-season canopy water loss during berry ripening and canopy maturation), showing moderate conductivity per area (latewood hydraulic conductivity per unit area lower than earlywood due to small vessels), supporting late season function (latewood conducting water during ripening period when demand reduced from peak but continued supply necessary), and demonstrating adequate efficiency (small latewood vessels providing acceptable water transport while prioritizing mechanical support function). Late wood rootstock xylem mechanical function includes= providing majority of wood strength (latewood contributing disproportionately to trunk mechanical properties due to thick-walled fibers), bearing structural loads (dense latewood tissue resisting compression, tension, and bending forces maintaining trunk integrity), resisting deflection (stiff latewood creating rigid framework preventing excessive trunk flexure under loads), enabling long-term stability (durable latewood maintaining structural properties over decades supporting sustained trunk function), and marking ring boundaries (latewood outer edge creating distinct line used for ring identification and age determination). Late wood rootstock xylem in annual ring position includes= occupying outer portion (latewood forming final part of annual ring immediately inside next year's earlywood), showing distinct outer boundary (sharp line marking transition from current year latewood to next year earlywood creating visible ring boundary), exhibiting variable width (latewood radial extent 0.1-0.4 cm varying with late season growing conditions and cambial activity duration), transitioning from earlywood (latewood beginning where earlywood vessel size declines marking mid-ring shift from spring to late summer production), and comprising ring fraction (latewood typically 40-60% of total ring width with proportion varying by year and seasonal patterns). Late wood rootstock xylem seasonal demand patterns includes= supporting berry ripening (latewood conducting water during veraison and maturation when berry sugar accumulation and acid decline occurring), enabling late canopy function (latewood transport supporting photosynthesis during late season when canopy maintaining productivity for ripening and reserve accumulation), providing reduced supply (latewood formation period corresponding to declining transpiration as day length shortens and canopy ages), facilitating reserve accumulation (latewood conducting photosynthates during late season when carbohydrate storage in trunk and roots increasing), and completing season (latewood formation marking final growth phase before dormancy ending annual xylem production). Late wood rootstock xylem formation conditions includes= reflecting late season conditions (latewood characteristics indicating water availability, temperature, and stress during July-September), showing drought sensitivity (late season water stress potentially producing extremely narrow latewood or early cessation creating narrow rings), exhibiting temperature effects (cooler late summer potentially extending latewood formation period while heat accelerating dormancy entry reducing latewood), demonstrating maturity effects (mature trunk showing consistent latewood production while stressed or declining trunk potentially showing reduced latewood), and recording stress timing (latewood width revealing whether late season favorable allowing continued growth or stressful forcing early growth cessation). Late wood rootstock xylem cultivar variation includes= showing variety-specific vessel sizes (cultivar genetics determining latewood vessel diameter range within typical 40-80 micrometers), exhibiting different latewood proportions (some rootstocks showing broad latewood zones creating dense rings while others showing narrow zones creating lighter rings), demonstrating density variation (Rupestris types producing very dense thick-walled latewood while Riparia showing less dense latewood reflecting genetic wood characteristics), and revealing vigor patterns (vigorous rootstocks maintaining latewood production longer creating broader zones while low-vigor types showing earlier cessation).
Late_Wood_Rootstock_Xylem emphasizes late-summer-formed small-vessel portion of annual ring and is distinguished from Early_Wood_Rootstock_Xylem (spring-formed large-vessel portion) by focus on outer ring position, small vessels, thick walls, and primary mechanical support function. Critical for understanding seasonal xylem production, structural strength, late-season water supply, wood density effects, and the dark-colored outer ring zone providing majority of mechanical support while contributing secondary water conduction capacity.
Dickwandiges Unterlagen-Xylem
Latewood
Small-vessel rootstock xylem
Sommerholz
Spätholz-Xylem
Summer wood rootstock xylem
Late Wood Rootstock Xylem
Merlot late wood xylem vessel in ring 12 shows 55-95 μm diameter, thick secondary wall 3.8-5.2 μm thickness (wall comprising 14-18% of vessel diameter compared to 1-2% in earlywood vessels), formed July-September during mid- to late-season cambial activity, scattered distribution through latewood zone occupying 58-64% of ring width, individual vessels providing 1/15 to 1/40 hydraulic conductivity of large earlywood vessels, latewood zone providing 24-32% of total ring transport capacity, vessels maintaining 65-80% functionality in late summer when earlywood vessels show 55-70% embolism
A late wood xylem vessel (latewood vessel) is a xylem vessel formed during the late portion of the growing season (typically June-September in Northern Hemisphere temperate climates), characterized by smaller diameter (40-120 μm), thicker walls, and moderate water transport capacity, forming the outer band of smaller vessels in ring-porous wood that provides backup hydraulic capacity, mechanical support, and greater embolism resistance. Late wood vessel formation timing includes= differentiation beginning after earlywood completion (typically June-July when large vessel production ceases), production during latter 8-14 weeks of growing season (June-September latewood formation period), formation during peak transpiration and fruit ripening (vessels supporting ongoing water demand through summer), cessation with fall cambial dormancy (typically September-October when growth ceases), and latewood zone completion representing 55-75% of annual ring width despite being 55-65% of formation duration. Late wood vessel anatomical characteristics include= smaller diameter (40-120 μm typical, compared to 150-300 μm earlywood vessels), thicker secondary walls (wall thickness 3-6 μm compared to 2-4 μm in earlywood vessels), smaller lumens (lumen occupying 34-110 μm of 40-120 μm diameter), simple perforation plates (similar to earlywood but connecting smaller-diameter vessels), moderate pit density (30-55 pits per 100 μm² wall area), and scattered or radial arrangement (distributed through latewood zone rather than concentrated bands). Late wood vessel hydraulic function includes= providing 20-40% of annual ring water transport capacity (smaller vessels contributing less due to radius⁴ relationship despite being 50-65% of vessel number), backup conductance when earlywood embolizes (latewood vessels maintaining function after earlywood cavitation providing residual transport), supporting ongoing transpiration through summer (latewood continuing to form and function during active growth period), moderate hydraulic conductivity (individual vessels transporting 1/10 to 1/100 of large earlywood vessel capacity), and enhanced drought tolerance (smaller vessels less prone to embolism maintaining function under stress). Late wood vessel embolism resistance includes= lower cavitation vulnerability (smaller diameter vessels with smaller pits and lower tensions resisting embolism), maintained function under stress (latewood vessels often remaining 60-80% functional when earlywood shows 60-80% embolism), freeze-thaw resistance (smaller vessels less vulnerable to ice-induced embolism), cumulative seasonal resilience (latewood vessels accumulating less embolism through growing season than earlywood), and critical safety function (latewood providing hydraulic safety margin when earlywood fails). Late wood vessel mechanical contribution includes= thicker walls providing structural support (latewood vessels contributing more to wood strength than thin-walled earlywood vessels), higher wood density in latewood zone (smaller lumens and thicker walls increasing tissue density 0.55-0.70 g/cm³ latewood compared to 0.40-0.55 g/cm³ earlywood), latewood zone providing mechanical reinforcement (outer ring portion resisting compression and bending), vessel walls resisting collapse (thicker walls preventing crushing under mechanical or hydraulic stress), and integration with fibers (latewood vessels intermixed with high-fiber density providing robust mechanical tissue). Late wood vessel development includes= differentiation from later cambial divisions (summer cambial activity producing smaller fusiform derivatives), limited radial expansion (latewood vessel precursors achieving 40-120 μm diameter compared to 150-300 μm earlywood expansion), proportionally more wall deposition (smaller vessels receiving thicker secondary walls relative to diameter), programmed cell death (protoplasm removal similar to earlywood vessel maturation), and gradual maturation (vessels completing differentiation over 3-6 weeks as cambial activity slows). Late wood vessel density and distribution includes= higher numerical density (30-55 vessels per mm² compared to 15-35 earlywood vessels), more vessels per unit area (smaller individual size allowing more vessels in given area), vessels comprising 65-80% of total ring vessel number (despite providing only 20-40% of conductance), scattered or radial multiples (vessels distributed individually or in small groups of 2-4), and latewood zone occupying 50-65% of ring area (moderate vessel lumen fraction 25-40% of latewood area, lower than earlywood 45-65%). Late wood vessel significance in hydraulic safety includes= providing redundancy (numerous smaller vessels offering alternative pathways if some embolize), maintaining minimum conductance under stress (latewood vessels continuing function when earlywood fails ensuring survival hydraulics), enabling recovery (latewood vessels supporting reduced but adequate water transport allowing vine survival and eventual recovery), progressive summer formation (continued latewood vessel production through season adding capacity as earlywood embolizes), and complementing earlywood function (latewood vessels providing hydraulic safety network balancing earlywood efficiency). Late wood vessel implications for viticulture includes= latewood vessels supporting stress survival (enabling vines to maintain function during severe drought when earlywood fails), latewood production varying with season length (longer growing seasons producing more latewood vessels enhancing hydraulic safety), latewood characteristics affecting wood density (proportionally more latewood producing denser stronger wood), and latewood vessel function explaining partial drought recovery (maintenance of latewood conductance allowing vines to continue photosynthesis and fruit ripening despite earlywood loss).
Late_Wood_Xylem_Vessel emphasizes summer-formed smaller vessels and is distinguished from Early_Wood_Xylem_Vessel (spring large vessels), Xylem_Vessel (all vessels), and vessel size classifications by seasonal formation timing and associated characteristics. Critical for understanding hydraulic safety margins, embolism resistance patterns, mechanical support in wood, drought tolerance mechanisms, and balance between hydraulic efficiency (earlywood) and safety (latewood) in ring-porous xylem.
Kleines Leitgefäß
Latewood vessel
Small vessel
Sommergefäß
Spätholzgefäß
Summer wood vessel
Late Wood Xylem Vessel
A latent bud embedded in the bark of a 15-year-old trunk of Vitis vinifera cv. Cabernet Sauvignon, remaining suppressed and non-growing for 8 consecutive seasons, measuring only 2-3mm in diameter, becoming activated following severe trunk damage from winter freeze injury that killed the primary cordons, developing into vigorous water shoots that allow canopy reconstruction and eventual return to production within 2-3 seasons through retraining and gradual fruiting wood establishment.
A bud that remains dormant and suppressed for extended periods beyond a single dormancy cycle, often for multiple years or decades, without developing into active shoots under normal growing conditions, characterized by arrested development, protective scale coverage, reduced size compared to normal buds, and capability to resume growth only under specific triggering conditions such as severe pruning, trunk damage, or environmental stress that releases apical dominance suppression. Latent buds represent a survival and regeneration strategy in perennial woody plants, providing reserve meristematic capacity for canopy reconstruction following catastrophic damage from freeze injury, disease, mechanical damage, or severe pruning, with their developmental fate controlled by complex interactions between apical dominance hormones (primarily auxin), local cytokinin production, and vascular connections to conducting tissues.
Latent buds differ fundamentally from dormant buds which cycle annually between active and dormant states, and from active buds which are currently growing. Latent buds can remain viable for decades in grapevines, with some documented cases of buds breaking after 30+ years of suppression. Their activation is typically undesirable in managed vineyards as resulting water shoots or suckers compete with productive fruiting wood, but latent bud capacity is essential for vineyard recovery from freeze damage, trunk disease remediation through retraining, or conversion between training systems. Position-specific differences exist, with basal, trunk, and old wood latent buds showing distinct activation thresholds and developmental potentials.
Reserveknospe
adventitious bud precursor
dormant latent bud
latente Knospe
sleeping bud
Based on Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press, pp. 85-92 Pina & Errea (2008) "Differential induction of phenylalanine ammonia-lyase gene expression in response to in vitro callus unions of Prunus spp." Journal of Plant Physiology 165=705-714.
Latent Bud
Latent deficiency quality detection for phosphorus in newly established Sonoma County Pinot Noir identified subclinical P limitation through tissue testing despite normal visual appearance, enabling preventive correction before visible symptoms developed. Visual assessment revealed no latent deficiency quality indicators with canopy appearing uniformly healthy, normal green coloration, adequate shoot vigor, and no signs of the red-purple pigmentation or growth reduction characteristic of overt P deficiency. However, routine tissue analysis detected latent deficiency quality with petiole P at 0.12% (suboptimal, below the 0.15% adequacy threshold but above the 0.08% critical concentration for visible symptoms), indicating nutritional status in the hidden hunger range. Sensitive physiological measurements confirmed latent deficiency quality with root growth rate 18% below P-sufficient reference vines, photosynthetic efficiency (Fv/Fm) slightly reduced at 0.78 versus 0.82 normal, and fruit set 8% below normal despite no visible inflorescence symptoms, collectively indicating subtle functional impairment consistent with suboptimal P status. The economic assessment showed latent deficiency quality causing estimated 6% yield reduction through the subtle fruit set effect, losses occurring without visible cause that would have been attributed to other factors (weather, vines immaturity) without the tissue analysis detection. The latent deficiency quality diagnosis enabled proactive management with moderate phosphorus application at 40 kg P₂O₅/ha banded near vines, a standard rate appropriate for suboptimal rather than deficient status. Follow-up tissue analysis showed petiole P increased to 0.18% (adequate range) with subtle productivity parameters normalizing, demonstrating successful preventive correction of latent deficiency before progression to visible symptoms and greater losses.
Latent deficiency quality is a deficiency severity quality that inheres in a grapevine organism and represents a subclinical nutritional limitation where tissue nutrient concentrations have declined below optimal levels but remain above the threshold for visible symptom expression, creating a condition detectable only through tissue analysis or sensitive physiological measurements while the vine appears visually normal, yet experiencing reduced metabolic efficiency that may affect productivity without obvious cause attribution. This quality captures the hidden or pre-symptomatic stage of deficiency, recognizing that the absence of visible symptoms does not guarantee adequate nutrition, and that latent deficiency can cause subtle but economically significant productivity losses that go undiagnosed without proactive tissue testing. The ontological significance of latent deficiency quality lies in its diagnostic challenge requiring tissue analysis for detection since visual examination reveals no abnormality, its potential for causing unexplained productivity shortfalls attributed to other factors, and its opportunity for proactive correction before visible symptoms and greater productivity impacts develop. The biological basis of latent deficiency encompasses tissue concentrations in the suboptimal range between clear adequacy and visible deficiency, where metabolic processes operate at reduced efficiency but compensatory mechanisms maintain apparent normality, stress-responsive pathways may be activated at low levels, and sensitive measures such as enzyme activities or photosynthetic parameters may show subtle reduction while visible indicators remain normal. The concept of the "critical concentration" defines the tissue nutrient level below which deficiency symptoms appear, with latent deficiency existing in the zone above this critical concentration but below optimal levels, sometimes termed the "hidden hunger" range where nutrient supply limits productivity without causing obvious symptoms. The management response to latent deficiency emphasizes preventive action through moderate fertilization adjustment to restore optimal status before progression to visible deficiency, representing the most efficient intervention point where correction is easiest and productivity losses can be prevented rather than merely remediated.
Latent_Deficiency_Quality is_a Deficiency_Severity_Quality, representing subclinical pre-symptomatic deficiency detectable only through tissue analysis or sensitive measurements. This class addresses the "hidden hunger" zone between optimal and visibly deficient. No visible symptoms despite suboptimal tissue concentrations. Requires tissue testing for detection (visual examination insufficient). Subtle productivity losses may occur and be misattributed. Preventive correction easiest and most efficient at this stage. Critical concentration concept defines visible symptom threshold. Most favorable intervention point before damage occurs.
Latenter Mangel
concealed deficiency
hidden hunger
pre-symptomatic deficiency
präsymptomatischer Mangel
subclinical deficiency
subklinischer Mangel
versteckter Hunger
Bergmann W (1992) Nutritional Disorders of Plants. Gustav Fischer Verlag, Stuttgart Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Ulrich A, Hills FJ (1967) Principles and practices of plant analysis. In: Soil Testing and Plant Analysis, Part II. SSSA Special Publication 2, pp. 11-24
Latent Deficiency Quality
Merlot exhibits high lateral bud fruitfulness disposition with 40-60% of summer lateral shoots bearing 1-2 small clusters, potentially contributing 10-20% of total yield as second crop in warm climates. Cabernet Sauvignon shows moderate lateral fruitfulness (25-40% of laterals fruiting) while Pinot Noir exhibits minimal lateral fruitfulness (<15% of laterals). Lateral clusters are smaller (50-120 g) than primary clusters (150-300 g) and ripen 2-4 weeks later, creating harvest timing complications. In regions with long growing seasons, lateral crop may reach 2-4 t/ha in fruitful cultivars.
A bud fruitfulness disposition that is the inherited capacity of a Vitis vinifera plant to develop inflorescence primordia in lateral (axillary) buds that produce summer lateral shoots (Geiztriebe), providing secondary crop potential from current-season growth in addition to the primary crop from dormant buds.
Lateral bud fruitfulness differs mechanistically from dormant bud fruitfulness, as lateral primordia differentiate during the current growing season without the dormancy period required for compound bud development. High lateral fruitfulness can complicate crop estimation and may reduce primary fruit quality through competition for resources. Lateral shoot removal (hedging, desuckering) is practiced in premium wine production to focus resources on primary clusters.
Axillarknospen-Fruchtbarkeit
Geiztriebfruchtbarkeit
Seitentriebfertilität
axillary bud fertility
lateral shoot fertility
summer lateral fruitfulness
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 88-94 Intrieri C, Poni S (1995) Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality of mechanized Italian vineyards. American Journal of Enology and Viticulture 46:116-127
Lateral Bud Fruitfulness Disposition
Lateral bud inhibition assessment along Merlot shoots reveals the position-dependent inhibition gradient affecting branching patterns, with basal positions (nodes 1-3) showing strong inhibition (5% lateral outgrowth despite full light exposure, maintained by basipetal auxin flux), middle positions (nodes 6-10) showing moderate inhibition (25% outgrowth, intermediate auxin concentration), and distal positions (nodes 15-18) showing weak inhibition (55% outgrowth in late season as distance from apex reduces inhibitory signal), with the inhibition gradient creating the characteristic pattern of increasing lateral development toward shoot tips observed in vigorous vines, informing decisions about tipping height and timing to control lateral proliferation.
Lateral bud inhibition quality is a paradormancy quality that represents the degree of growth suppression experienced by buds at lateral positions along shoots, characterizing the integrated inhibitory state resulting from apical dominance and other correlative signals that determines whether specific lateral buds remain dormant, show limited activity, or escape inhibition to produce lateral shoots during the growing season. Within the BFO framework, this quality specifically addresses the lateral bud state dimension of paradormancy, distinguishing the inhibition status of individual lateral buds from the strength of inhibitory signals while recognizing that lateral bud inhibition integrates multiple factors including distance from apex (proximal versus distal buds experiencing different inhibition), bud vigor and nutritional status, local light environment, and the overall correlative signaling context. The ontological significance lies in lateral bud inhibition determining the actual pattern of lateral shoot development that creates canopy architecture, where strong inhibition maintains compact shoot structure with few laterals (simplified canopy, easier management), moderate inhibition permits selective lateral development (filling canopy gaps while maintaining organization), and weak inhibition results in profuse branching (dense canopy requiring intervention), with the inhibition pattern across all lateral buds collectively determining canopy complexity. The factors modulating lateral bud inhibition include position along the shoot (basal buds often more strongly inhibited than distal buds closer to the weakening apical signal), bud fruitfulness (fruitful buds potentially experiencing different correlative relationships than vegetative buds), shoot vigor (vigorous shoots often showing stronger inhibition from more active apical meristems), and light penetration (shaded buds more strongly inhibited than those receiving direct light stimulating outgrowth). The management relevance of lateral bud inhibition includes the prediction of lateral development following topping operations (released from apical inhibition), the assessment of canopy filling potential from lateral growth, and the understanding of why certain bud positions consistently produce laterals while others remain inhibited throughout the season. Understanding lateral bud inhibition quality enables position-specific prediction of lateral bud behavior, assessment of canopy development potential from lateral outgrowth, and targeted canopy management addressing specific inhibition patterns.
Lateral_Bud_Inhibition_Quality addresses the growth suppression state of lateral buds integrating multiple correlative signals. This quality varies with position, vigor, and environment affecting branching patterns. The class supports canopy architecture prediction through lateral bud inhibition characterization.
Achselknospendormanz
Lateralknospenhemmung
Seitenknospenunterdrückung
Verzweigungshemmung
axillary bud dormancy
branch inhibition
lateral bud quiescence
lateral suppression
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 59-144 Bennett T, Leyser O (2014) Strigolactone signalling: standing on the shoulders of DWARFs. Current Opinion in Plant Biology 22: 7-13
Lateral Bud Inhibition Quality
A lateral lobe representing the upper left lateral lobe on a mature leaf at node 7 of Vitis vinifera cv. Merlot in Pomerol, France, one of four lateral lobes (two upper, two lower pairs) on this five-lobed leaf, positioned between the apical lobe and lower left lateral lobe, vascularized by the second primary lateral vein branching from median vein at 55-degree angle, measuring 4.5 cm maximum width and 4.8 cm length creating approximately equidimensional lobe shape, separated from apical lobe by upper left lateral sinus of 2.6 cm depth and from lower lateral lobe by middle left lateral sinus of 2.2 cm depth, lobe apex showing obtuse configuration with approximately 75-degree angle, lobe margins exhibiting slightly convex curvature with heterogeneous tooth pattern, lobe area representing approximately 17% of total blade area, comparable to other lateral lobes but smaller than apical lobe (28% of area), moderate lobe development creating distinct but not deeply separated appearance characteristic of Merlot, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing typical lateral lobe morphology in intermediate position between terminal and basal zones.
A lobe positioned between the apical and basal lobes on the lateral sides of the leaf blade in Vitis vinifera, characterized by intermediate position in the lobe sequence from base to apex, vascularization by primary lateral veins branching from the median vein, typical occurrence of two pairs (upper and lower) creating four lateral lobes in five-lobed leaves or reduced to one pair in three-lobed leaves, variable size generally smaller than apical lobe but comparable to or larger than basal lobes, separation from adjacent lobes by lateral sinuses of variable depth, cultivar-specific morphological features including relative size, width, apex characteristics, and degree of separation serving as diagnostic characters, and functional contribution providing substantial blade area typically 35-50% combined area of all lateral lobes representing major photosynthetic surface. Lateral lobes constitute the intermediate organizational units between terminal and basal positions, developing through coordinated growth of primary lateral vein sectors during blade expansion, exhibiting bilateral arrangement flanking the median vein with potential slight asymmetries, and showing cultivar-specific patterns including relative development compared to apical and basal lobes, degree of dissection, and morphological characteristics contributing to overall leaf architecture and identification.
Lateral lobes are distinguished from apical lobes by intermediate rather than terminal position and from basal lobes by position above the petiolar junction separated from base by basal lobes. In five-lobed leaves (most common configuration), four lateral lobes occur as two pairs (upper and lower) flanking the median vein bilaterally, while three-lobed leaves show one pair of laterals and seven-lobed leaves show three pairs. Size relationships among lobe types show cultivar specificity, with some varieties showing large prominent lateral lobes comparable to or exceeding apical lobe size, while others show reduced laterals subordinate to prominent apical lobes. Separation from adjacent lobes depends on lateral sinus depth, varying from shallow sinuses creating weakly defined laterals to deep sinuses creating prominent isolated lateral lobes. Upper versus lower lateral lobe distinction creates useful subcategories, with upper laterals typically larger and more prominent than lower laterals in most cultivars. Bilateral arrangement creates potential for asymmetry, with left and right lateral lobes sometimes showing size or shape differences though symmetry predominates. Functional contribution involves substantial area provision with combined lateral lobes typically representing 40-55% of total blade area, making them major contributors to photosynthetic capacity. Ampelographic utility involves assessment of lateral lobe characteristics including relative size, degree of development, apex characteristics, and symmetry combining with apical and basal lobe features for cultivar identification.
Lateralausprägung
Seitenlappen
intermediate lobe
intermediärer Lappen
lateral division
side lobe
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 58-89 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 137-172.
Lateral Lobe
A lateral petiole on a deeply lobed mature leaf at node 6 of an unusual Vitis vinifera cv. Mourvèdre clone in Bandol, France, appearing as a 1.2 cm length stalk-like extension at the junction between the main vein and the lateral lobe vein in a leaf showing exceptionally deep sinuses, diameter of 1.5 mm representing miniature petiolar structure, light green coloration matching main petiole, sparse pubescence, containing single small vascular strand visible in cross-section, providing minimal additional mechanical support to the lateral lobe beyond what the major vein supplies, occurring on approximately 15% of leaves on this particular vine representing uncommon morphological variant, showing no clear correlation with leaf function or vine performance, measured at BBCH stage 75 (berries pea-sized) in late June, representing rare lateral petiole expression in a cultivar and clone showing tendency toward extremely deep lobe dissection approaching compound leaf morphology.
A secondary petiolar structure or petiole-like extension in grapevine leaves representing uncommon variant morphology in Vitis vinifera, characterized by small stalk-like structures occasionally supporting minor lobe divisions or appearing as vestigial structures at major vein junctions in some cultivars with deeply lobed leaves, typical dimensions of 0.5-2 cm length and 1-2 mm diameter substantially smaller than main petioles, occurrence as rare morphological variants rather than standard features with frequency varying among cultivars from absent in most varieties to occasional appearance in deeply lobed cultivars (some Mourvèdre clones, certain Muscat types), anatomical structure including miniature vascular strands and mechanical tissues resembling but much smaller than main petiole tissues, and questionable functional significance with these structures providing minimal additional mechanical support or vascular capacity beyond what major veins supply. Lateral petioles represent evolutionary vestiges or developmental variations recalling more compound leaf architectures found in related Vitaceae species or ancestral forms, appearing most commonly in cultivars or individual plants showing deepest lobe development where sinuses nearly reach the petiolar junction creating quasi-separate leaflet-like structures, and serving primarily as morphological curiosities rather than functionally important structures or reliable ampelographic characters given their inconsistent occurrence and subtle expression.
Lateral petioles represent uncommon morphological features in Vitis vinifera, distinguishable from main petioles by their small secondary nature and from major veins by their stalk-like petiolar rather than laminar character. These structures occur rarely and inconsistently, absent in most cultivars and most leaves even in cultivars showing occasional expression, making them unreliable ampelographic characters. The occurrence relates to deep lobe dissection creating nearly separated leaflet-like structures, with lateral petioles representing incomplete transition toward compound leaf architecture never fully realized in modern Vitis vinifera cultivars. Functional significance is minimal, with these small structures contributing negligibly to mechanical support or vascular capacity compared to major veins and main petiole. Evolutionary interpretation suggests vestigial retention of ancestral compound leaf features, with most Vitaceae showing simple leaves but some related genera exhibiting compound leaves with true lateral petioles, and occasional Vitis expressions representing developmental atavisms. Practical importance is limited, with these structures representing morphological curiosities noted in detailed descriptions but not serving diagnostic roles in cultivar identification or vineyard management.
Nebenblattstängel
lateraler Blattstiel
minor petiole
secondary petiole
sekundärer Petiolus
vestigial petiole
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 58-89 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 33-55.
Lateral Petiole
Cabernet Sauvignon grafted on 1103 Paulsen showing lateral root cap on primary root tip, lateral cap flanking central columella extending 0.42 mm radially from columella margin to cap periphery, 6-7 cell layers thick providing lateral protection, microscopic examination revealing lateral cap cells 32 micrometers average diameter with thick walls and large vacuoles, lateral cap cells containing 3-8 small plastids versus 65 large amyloplasts in adjacent columella cells, lateral cap providing mechanical protection of meristem flanks while columella specializes in gravity sensing
Lateral root cap is root cap tissue flanking columella comprising peripheral cap region extending from columella files to cap margin, typically 3-8 cell layers thick forming protective covering on sides of root apex, characterized by cells with fewer or no starch-filled amyloplasts compared to columella showing predominantly protective rather than gravity-sensing function, arranged in files extending from cap base near meristem to cap periphery where cells approach sloughing, and serving primarily mechanical protection function shielding meristem from lateral impacts and abrasion while enabling root tip to push through soil, with lateral root cap showing structural features including thicker walls than columella cells providing greater mechanical strength, larger cells than meristem but smaller than columella cells, progressive maturation from base to periphery with cells aging as move distally, and capacity for mucilage production particularly in outer layers, and demonstrating that lateral cap provides robust mechanical barrier protecting meristem flanks while columella specializes in gravity sensing creating functional division within root cap. Lateral root cap positional characteristics includes= showing peripheral position (lateral cap flanking columella extending from central files to cap outer margin), exhibiting bilateral symmetry (lateral cap on both sides of columella creating symmetric protection), demonstrating layered structure (typically 3-8 concentric cell layers from inner near columella to outer approaching surface), revealing file organization (cells arranged in longitudinal files parallel to root axis though less regular than columella), and displaying coverage extent (lateral cap covering meristem flanks providing lateral protection while columella covers apex). Lateral root cap cellular characteristics includes= showing moderate size (lateral cap cells typically 20-40 micrometers intermediate between meristem and columella cells), exhibiting large vacuoles (mature lateral cap cells with prominent central vacuoles occupying 70-85% cell volume), demonstrating thick walls (lateral cap cell walls often thicker than columella cells providing mechanical strength), revealing few amyloplasts (lateral cap cells containing few if any starch-filled plastids contrasting with abundant columella amyloplasts), and displaying polysaccharide production (lateral cap cells particularly outer layers producing and secreting mucilage). Lateral root cap protective function includes= providing mechanical shielding (lateral cap cells absorbing impacts and abrasion from soil particles protecting meristem flanks), exhibiting expendable barrier (lateral cap cells programmed for death and sloughing creating renewable protection that can be damaged without meristem injury), demonstrating structural strength (thick-walled lateral cap cells providing robust barrier against mechanical forces), revealing penetration assistance (lateral cap geometry helping root tip displace soil during penetration), and displaying lubrication (lateral cap mucilage production reducing friction facilitating root advancement). Lateral root cap maturation gradient includes= showing youngest cells basally (newly formed lateral cap cells near cap base adjacent to meristem), exhibiting progressive maturation (cells maturing as move distally toward cap surface), demonstrating structural changes (cell walls thickening, vacuoles enlarging, cytoplasm becoming less dense as cells mature), revealing functional changes (younger lateral cap cells primarily structural while older cells increasingly involved in mucilage production and sloughing), and displaying senescence (outermost lateral cap cells approaching programmed death and detachment). Lateral root cap turnover dynamics includes= showing continuous production (lateral root cap initials continuously generating new cells maintaining cap despite losses), exhibiting basipetal movement (newly formed cells progressively displaced distally by continued cell production at base), demonstrating progressive aging (lateral cap cells aging as move from production zone toward periphery), revealing eventual sloughing (oldest outermost cells detaching from cap surface as reach terminal stage), and displaying rapid renewal (lateral cap completely replacing itself over days to weeks creating dynamic structure). Lateral root cap versus columella includes= contrasting in amyloplast content (lateral cap cells with few or no starch-filled amyloplasts versus abundant in columella), showing different functions (lateral cap primarily mechanical protection versus columella gravity sensing), exhibiting structural differences (lateral cap with thicker walls and different cell shapes optimizing protection versus columella optimizing sensing), demonstrating positional distinction (lateral cap flanking periphery versus columella axial center), and revealing developmental separation (lateral cap and columella potentially arising from separate cap initial populations). Lateral root cap in root penetration includes= enabling soil displacement (lateral cap structure allowing root to push aside soil particles), providing friction reduction (lateral cap mucilage lubricating passage through soil), demonstrating force distribution (lateral cap distributing mechanical forces from soil resistance protecting meristem), revealing tip geometry (lateral cap contributing to overall cap shape optimizing penetration efficiency), and displaying impedance response (lateral cap potentially thickening or modifying structure in response to mechanical impedance). Lateral root cap mucilage production includes= showing polysaccharide synthesis (lateral cap cells particularly outer layers synthesizing mucilage polysaccharides), exhibiting Golgi involvement (dictyosomes producing vesicles containing mucilage precursors), demonstrating secretion (mucilage vesicles fusing with plasma membrane releasing contents to apoplast and root surface), revealing accumulation (secreted mucilage accumulating around lateral cap creating mucilage sheath), and displaying functions (lateral cap mucilage lubricating penetration, modifying rhizosphere, potentially providing defense).
Lateral_Root_Cap emphasizes peripheral protective tissue flanking columella and is distinguished from central columella by focus on mechanical protection versus gravity sensing. Includes Peripheral_Lateral_Root_Cap and Protective_Lateral_Root_Cap. Critical for understanding mechanical shielding, structural strength, maturation gradients, turnover dynamics, mucilage production, and the flanking cap tissue providing robust lateral protection while columella performs gravity sensing.
Flanking cap tissue
Laterale Wurzelhaube
Peripheral root cap
Periphere Haube
Protective cap layer
Schützende Haubenschicht
Barlow, P. W. (2003). The root cap= Cell dynamics, cell differentiation and cap function. Journal of Plant Growth Regulation, 21(4), 261-286.
Lateral Root Cap
A lateral secondary branch rachis on a primary inflorescence of Vitis vinifera cv. Grenache in Châteauneuf-du-Pape, France, emphasizing its lateral positioning arising from the side of a primary branch, originating from the fifth primary branch at 2.5 cm from the primary's base at approximately 90-degree angle creating lateral perpendicular extension, representing one of 8 lateral secondary branches distributed along this 5.8 cm primary branch at intervals of 0.6-0.9 cm, measuring 2.2 cm length extending laterally from the primary axis, diameter of 1.1 mm, containing 2 vascular bundles receiving lateral traces from the primary branch vascular system, bilateral organization evident with this secondary on the left side of the primary branch and a corresponding right-side secondary at similar position on the opposite side, bearing 3 tertiary branches and 5 direct flowers creating lateral flower distribution, complete lateral branch system extending the flower-bearing zone approximately 2.2 cm perpendicular to the primary branch significantly expanding the three-dimensional inflorescence volume, measured at BBCH stage 63 (30% flowers open) in early June, representing typical lateral secondary branch rachis architecture with distributed lateral positioning creating systematic three-dimensional elaboration.
A secondary branch rachis emphasizing the lateral positioning aspect as branches arising from the lateral surfaces of primary branches in Vitis vinifera, characterized by lateral origin from primary branch rachides rather than terminal or other positions, distributed arrangement along the primary branch length creating lateral elaboration arrays, typical bilateral or radial organization with secondary branches arising from opposite or multiple sides of the primary branch, dimensions and spacing creating the lateral branching pattern typically 1.5-3.5 cm length at 0.5-1.0 cm intervals, vascular connection through lateral trace departure from primary branch bundles, and functional role creating lateral spatial expansion from the primary branch axis distributing flowers into the three-dimensional inflorescence volume. Lateral secondary branch rachides emphasize the spatial and organizational aspects of lateral positioning, representing the distributed lateral elaboration from primary branches, exhibiting regular lateral branching patterns creating systematic architecture, and serving to create the three-dimensional flower distribution characteristic of paniculate inflorescence architecture.
Lateral secondary branch rachides emphasize the spatial positioning aspects as lateral structures, distinguished from hierarchical emphasis by focus on lateral arrangement and distribution. The lateral designation highlights the geometric relationship as side branches extending perpendicular or oblique to the primary branch axis, contrasting with theoretical terminal or basal positioning. Distributed arrangement creates systematic lateral elaboration with multiple secondary branches at regular intervals along primary branch length, creating the characteristic dense branching. Bilateral organization is common with secondary branches arising from opposite sides of the primary branch in paired or alternate patterns, creating balanced lateral expansion. Radial arrangement occurs in some cases with secondary branches arising from multiple sides around the primary branch circumference, creating more complex three-dimensional structures. Spacing intervals of 0.4-1.0 cm create the density of lateral branching, with closer spacing producing dense elaborate structures and wider spacing creating more open architecture. Angular relationships show secondaries typically arising at 60-120 degree angles from the primary branch axis, with angles affecting the three-dimensional shape and compactness. Vascular connection through lateral trace departure creates the hierarchical vascular network with lateral bundles branching from primary bundles. Functional implications include creating three-dimensional expansion distributing flowers spatially rather than concentrating along linear axes, affecting light penetration, pollination access, and final cluster architecture. Developmental coordination maintains distributed lateral arrangement through systematic primordium initiation along developing primaries. Transformation to cluster maintains lateral organization with these structures creating the distributed berry-bearing architecture defining cluster shape and density.
distributed lateral rachis
laterale sekundäre Zweigrachis
radial angeordnete Sekundärspindel
radially arranged secondary
seitenpositionierte Zweitordnung Achse
side-positioned secondary branch
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 471-508 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 679-708.
Lateral Secondary Branch Rachis
A lateral secondary inflorescence at node 5 on a shoot of Vitis vinifera cv. Cabernet Sauvignon in Bordeaux, France, measuring 14.2 cm length representing typical lateral secondary dimensions, positioned opposite the fifth fully expanded leaf at standard lateral node location, exhibiting compound paniculate architecture with central rachis bearing 3-4 orders of branching, rachis diameter of 2.9 mm at base, bearing approximately 380 individual flowers, positioned in mid-shoot region approximately 35 cm from shoot base and 40 cm from current shoot tip, showing dimensions approximately 78% of the primary inflorescence at node 3 (18.2 cm, 580 flowers), intermediate location between basal primary and potential apical structures creating mid-shoot reproductive zone, developmental stage at BBCH 63 (30% flowers open) showing typical 2-day delay compared to primary, lateral position providing good light exposure and air circulation without extreme apical vulnerability, expected to contribute approximately 32% of shoot yield following fruit set, measured in early June during mid-bloom period, representing typical lateral secondary inflorescence at standard node 5 position with intermediate characteristics.
A secondary inflorescence positioned at standard lateral node locations along the shoot in Vitis vinifera, characterized by position at node 5 or 6 representing typical lateral placement in the shoot inflorescence sequence, intermediate location between basal primary inflorescences and potential apical tertiary or terminal inflorescences, standard opposite-leaf positioning creating the characteristic lateral arrangement, typical dimensions of 11-17 cm length representing moderate development, moderate branching complexity with 3-4 orders creating standard secondary architecture, occurrence as the most common secondary inflorescence type representing the default lateral positioning pattern, and functional role providing supplementary yield contributing to mid-shoot fruit production zones. Lateral secondary inflorescences represent the standard secondary inflorescence positioning, occurring at typical inflorescence node positions following the two-node gap pattern, exhibiting characteristics intermediate between basal primaries and more distal structures, and constituting the primary secondary inflorescence type in most vineyard contexts.
Lateral secondary inflorescences represent the standard positioning for secondary inflorescences, distinguished from basal secondaries by mid-shoot rather than extreme basal location, though the distinction can be subtle given node 5 position is relatively basal on most shoots. The lateral designation emphasizes the standard node position following typical inflorescence spacing patterns rather than exceptional basal or terminal placements. Position at node 5 creates mid-shoot location on typical 10-15 node fruiting canes, though appearing more basal on longer vigorous shoots or more distal on short spurs. Standard opposite-leaf positioning maintains the characteristic grapevine architecture with reproductive nodes alternating with vegetative-only nodes. Size and complexity show typical secondary characteristics with moderate reduction from primaries but substantial development creating meaningful yield contribution. Occurrence represents the default pattern with most secondary inflorescences occupying this standard lateral position. Light exposure and environmental conditions are generally favorable without the extremes of basal shading or apical exposure, creating moderate development conditions. Developmental timing and resource allocation follow typical secondary patterns with delayed initiation and subordinate investment compared to primaries. Management decisions often focus on lateral secondaries as the primary targets for cluster thinning when crop reduction is needed, as their removal provides significant yield reduction while retaining the larger primary clusters. Retention versus removal depends on crop load targets with low to moderate targets favoring removal while higher targets may retain secondaries for supplementary production.
Standardposition-Zweitblütenstand
laterale sekundäre Infloreszenz
mid-shoot secondary
mittlerer Spross-Sekundärstand
node 5 lateral inflorescence
standard position secondary
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 219-258 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 183-218 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 359-398.
Lateral Secondary Inflorescence
Chardonnay grafted on SO4 rootstock at 15 years showing lateral secondary root system, first-order lateral secondary roots emerging from primary roots at 70-85° angles growing predominantly horizontally, lateral roots concentrated 35-75 cm depth exploring soil laterally, longest lateral secondary root extending 185 cm from primary root origin approximately 240 cm from trunk center far exceeding canopy radius 120 cm, lateral secondary roots 1.8-4.2 cm diameter at origin tapering along length, abundant second and third-order branching creating extensive network, lateral roots providing majority of soil exploration and resource uptake for vine
Lateral secondary root is secondary root emphasizing lateral orientation growing horizontally or obliquely from parent root, characterized by emergence perpendicular or at oblique angle typically 45-90° from parent root axis creating sideways rather than vertical growth, predominantly horizontal trajectory exploring soil laterally rather than penetrating deep, concentration in upper to middle soil horizons typically 20-100 cm depth where majority of water and nutrients available, and serving as primary mechanism for lateral soil exploration extending root system horizontally far beyond primary root vertical extent potentially 100-300+ cm radius from trunk, with lateral growth pattern enabling efficient exploitation of soil resources distributed horizontally in layers, creating extensive surface area for absorption through fine branch roots, and demonstrating that lateral secondary root architecture represents key adaptation for maximizing soil contact and resource acquisition in layered soil profiles characteristic of agricultural environments. Lateral secondary root orientation and trajectory includes= growing perpendicular to parent (lateral secondary root typically emerging at 60-90° angle from parent root axis creating sideways trajectory), showing horizontal or oblique path (lateral growth predominantly sideways rather than downward typically 0-45° from horizontal), maintaining lateral direction (lateral secondary root continuing horizontal trajectory unless deflected by obstacles or gradients), demonstrating gravitropic response (lateral roots showing weak or absent gravitropism unlike vertical tap roots allowing horizontal growth), and creating planar spread (lateral secondary roots growing in multiple radial directions from parent creating disk-like or hemispherical exploration pattern). Lateral secondary root depth distribution includes= concentrating in upper horizons (majority of lateral secondary roots typically 20-80 cm depth in zone of maximum resource availability), showing depth stratification (lateral root density highest at optimal depth declining above and below), extending less deep than primary roots (lateral secondary roots typically shallower than tap root or main primary roots focusing on different soil zones), demonstrating layer exploitation (lateral roots particularly abundant in fertile topsoil or zones with accumulated organic matter and nutrients), and varying with parent depth (lateral secondary roots emerging from deep primary roots exploring deeper lateral zones than those from shallow primary roots). Lateral secondary root length and extent includes= extending far from trunk (individual lateral secondary roots potentially growing 100-200 cm from parent root or 150-300 cm from trunk center), exceeding canopy projection (lateral root spread typically 1.5-3 times canopy diameter creating extensive exploration zone), showing cumulative length (total length of all lateral secondary roots potentially exceeding 100-500 meters per vine), demonstrating progressive extension (lateral secondary roots continuing elongation throughout growing season adding 20-100+ cm annually depending on vigor), and limited by competition (lateral root extent constrained by roots from neighboring vines, cover crops, or other plants). Lateral secondary root diameter and structure includes= ranging from fine to moderate (lateral secondary root diameter typically 0.5-4 cm depending on order and age), showing gradual taper (diameter decreasing progressively along length as branches emerge), exhibiting secondary growth (older structural lateral secondary roots developing secondary xylem and bark), demonstrating root order correlation (first-order laterals thicker than second-order, second-order thicker than third-order reflecting hierarchy), and displaying flexibility (lateral secondary roots relatively flexible compared to rigid primary roots adapting to soil structure). Lateral secondary root branching pattern includes= producing abundant branch roots (lateral secondary roots generating numerous higher-order branches creating hierarchical network), showing acropetal branching (branches more abundant near growing tip declining toward base), exhibiting branching angles (branch roots emerging 45-90° from lateral parent creating multi-dimensional network), demonstrating variable density (branching density varying with resource availability proliferating in favorable zones), and creating absorptive network (lateral secondary root branches ultimately producing fine absorptive roots providing uptake surface). Lateral secondary root resource exploitation includes= exploring large soil volume (lateral network accessing soil unavailable to vertical primary roots maximizing total exploration), intercepting rainfall infiltration (shallow lateral roots positioned to capture water percolating through topsoil), accessing nutrient-rich layers (lateral roots exploiting zones of nutrient accumulation from fertilization or organic matter decomposition), competing with neighbors (lateral roots encountering and competing with roots from adjacent vines or other plants), and responding to heterogeneity (lateral roots proliferating preferentially in resource-rich patches and avoiding poor zones). Lateral secondary root seasonal growth includes= showing spring elongation (lateral secondary roots resuming growth early spring adding new length and branches), maintaining summer activity (continued lateral root development during growing season though potentially reduced during peak shoot growth), exhibiting fall renewal (lateral roots showing renewed growth late season particularly fine absorptive branches), entering winter dormancy (lateral root elongation ceasing during cold periods though physiological activity maintained), and demonstrating turnover (fine lateral branches showing annual mortality and replacement while larger lateral structural roots persisting). Lateral secondary root functional importance includes= providing majority of soil contact (lateral network creating most extensive interface between vine and soil), contributing most water uptake (lateral secondary roots with fine branches providing 60-80% of total vine water absorption), supplying most nutrient uptake (lateral roots exploiting nutrient-rich upper soil providing 70-90% of total mineral uptake), enabling mycorrhizal associations (lateral fine roots hosting fungal symbionts enhancing phosphorus and water uptake), and creating anchorage network (lateral secondary roots providing horizontal stabilization preventing sideways movement or toppling).
Lateral_Secondary_Root emphasizes horizontal orientation and lateral exploration and is distinguished from vertical or deep-penetrating roots by focus on sideways growth. Includes First_Order_Lateral_Secondary_Root, Second_Order_Lateral_Secondary_Root, and Deep_Lateral_Secondary_Root. Critical for understanding horizontal soil exploration, upper horizon exploitation, resource uptake, lateral extent, and the horizontally oriented secondary roots creating extensive lateral soil contact far from trunk center.
Horizontal secondary root
Horizontal wachsende Wurzel
Laterale Sekundärwurzel
Radial lateral root
Seitlich verlaufende Wurzel
Sideways-growing root
Lateral Secondary Root
Cabernet Sauvignon showing lateral shallow roots, excavation revealing first-order lateral roots 3.2-4.8 cm diameter branching from primary roots at 15-35 cm depth, lateral roots extending 140-180 cm horizontal radius, predominantly horizontal orientation 65-85° from vertical, lateral roots showing moderate secondary growth with 6-9 annual rings, lateral roots supporting dense populations of fine feeder roots providing estimated 72% of vine water and nutrient uptake, lateral shallow roots demonstrating primary horizontal soil exploration mechanism
Lateral shallow root is shallow root growing laterally or obliquely from parent root in horizontal or near-horizontal orientation, typically located 10-50 cm depth, characterized by predominantly horizontal trajectory spreading outward from vine trunk, moderate to substantial diameter 0.5-5 cm depending on order and age, serving dual functions of lateral soil exploration and structural support, and representing primary mechanism for extending root system horizontally creating extensive surface area for resource capture in topsoil, with lateral shallow roots in grapevine showing typical characteristics including first-order laterals branching directly from primary taproots or cutting-derived roots extending 50-150+ cm radius, second-order laterals branching from first-order spreading further into inter-row or alley spaces, predominantly horizontal growth angle 45-90° from vertical maximizing topsoil exploration, moderate secondary growth in structural laterals developing woody tissue for support and transport, and high absorptive root production with lateral shallow roots serving as parent axes supporting thousands of fine feeder roots, and demonstrating critical importance where lateral shallow root extent and branching largely determines total root system volume and absorption capacity particularly in shallow or restrictive soils where vertical root penetration limited. Lateral shallow root orientation and trajectory includes= showing predominantly horizontal growth (lateral roots growing 45-90° from vertical emphasizing horizontal rather than vertical soil exploration), exhibiting outward spreading (lateral roots extending radially outward from trunk or parent root), demonstrating depth maintenance (lateral roots generally maintaining relatively constant depth 10-50 cm rather than ascending or descending sharply), revealing flexibility (lateral roots capable of adjusting trajectory navigating obstacles or following favorable conditions), and displaying bilateral symmetry (lateral roots typically developing on multiple sides of parent creating radial distribution). Lateral shallow root hierarchical organization includes= showing branching orders (lateral shallow roots comprising first-order branches from primary roots, second-order from first-order laterals, potentially higher orders), exhibiting diameter decline (diameter decreasing with successive branching from 3-5 cm first-order to 1-3 cm second-order to <1 cm higher-order), demonstrating progressive specialization (lower-order laterals emphasizing structure and transport, higher-order emphasizing absorption), revealing architectural hierarchy (organized branching creating tree-like architecture with structural framework supporting absorptive network), and displaying order-specific functions (each branching order showing characteristic diameter, anatomy, lifespan, and function). Lateral shallow root extent and distribution includes= showing substantial horizontal reach (lateral shallow roots extending 100-300+ cm from trunk depending on soil, cultivar, spacing), exhibiting concentration zone (maximum lateral root density typically 30-100 cm from trunk in zone of intensive exploration), demonstrating inter-row extension (lateral roots potentially extending into alleys or between rows accessing resources beyond vine footprint), revealing asymmetric distribution (lateral root extent potentially varying by direction responding to neighboring vines, tillage, or soil heterogeneity), and displaying rootstock effects (rootstock varieties differing in lateral root extent affecting resource capture zone). Lateral shallow root structural development includes= showing secondary growth (structural lateral roots undergoing secondary thickening developing woody tissue), exhibiting bark formation (older lateral roots developing protective bark as secondary growth proceeds), demonstrating annual rings (woody lateral roots showing growth rings indicating age and growth patterns), revealing lignification (secondary xylem becoming progressively lignified providing strength and rigidity), and displaying persistent structure (structural lateral roots functioning years to decades providing long-term framework). Lateral shallow root absorptive function includes= providing fine root support (lateral shallow roots serving as parent axes from which thousands of fine absorptive roots emerge), exhibiting high absorptive root density (lateral roots in topsoil supporting dense populations of feeder roots maximizing absorption), demonstrating resource capture (lateral roots extending absorption zone enabling access to nutrients and water over large soil volume), revealing seasonal production (lateral roots producing pulses of fine roots spring and fall when conditions favorable), and displaying critical contribution (lateral shallow root-associated fine roots providing 60-80% of total vine uptake). Lateral shallow root in soil exploration includes= enabling lateral expansion (lateral roots primary mechanism for extending root system horizontally into unexplored soil), providing topsoil exploitation (lateral roots concentrated in fertile topsoil layers accessing abundant resources), demonstrating efficient exploration (horizontal spreading creating large soil contact area with minimal vertical penetration), revealing competitive advantage (extensive lateral roots capturing resources over large area potentially excluding competitors), and displaying architectural optimization (lateral root distribution optimized for capturing heterogeneously distributed resources). Lateral shallow root management implications includes= affecting cultivation depth (lateral shallow roots vulnerability to tillage damage requiring shallow cultivation or no-till), influencing fertilizer placement (lateral root distribution determining optimal fertilizer placement for efficient uptake), demonstrating irrigation importance (lateral roots in topsoil benefiting from irrigation maintaining function during drought), revealing mulch benefits (surface mulches protecting lateral roots from temperature extremes and moisture fluctuations), and displaying spacing effects (vine spacing affecting lateral root competition and resource partitioning).
Lateral_Shallow_Root emphasizes horizontal spreading in topsoil and is distinguished from vertical roots by focus on lateral orientation. Includes First_Order_Lateral_Shallow_Root, Second_Order_Lateral_Shallow_Root, and Surface_Lateral_Root hierarchies. Critical for understanding horizontal exploration, branching hierarchy, topsoil concentration, structural support, absorptive function, and the laterally spreading roots in upper soil creating extensive resource capture zone.
Ausbreitende Oberflächenwurzel
Horizontal shallow root
Horizontale Oberbodenwurzel
Lateral surface root
Laterale flache Wurzel
Spreading topsoil root
Lateral Shallow Root
A lateral shoot on Vitis vinifera cv. Sauvignon Blanc arising from node 8 axillary bud on primary shoot, emerged in late June (mid-season prompt lateral timing) during continued primary shoot elongation, developed to 45cm length by early August with 7 leaves totaling approximately 735 cm² additional photosynthetic area, 1 tendril, no inflorescences (typical lateral sterility), representing moderate lateral development common in Sauvignon Blanc contributing supplemental leaf area (approximately 27% increase over primary shoot alone) supporting fruit maturation, this lateral retained and positioned into catch wire system rather than removed, contributing to whole-shoot photosynthetic capacity of 3480 cm² (primary 2745 cm² plus lateral 735 cm²) adequately supporting two-cluster load (combined 410g) with favorable 20.9 cm² leaf area per gram fruit ratio enhanced by lateral contribution.
A shoot arising from axillary buds on current-season primary shoots, characterized by lateral branching from main shoot axis at leaf axil positions, secondary-order growth developing during or after primary shoot elongation, variable timing from prompt same-season emergence (sylleptic) to delayed following-season development (proleptic), and function providing additional photosynthetic capacity through increased leaf area, compensatory growth following damage or stress, architectural complexity in vigorous vines, or management challenges through excessive canopy density requiring shoot removal or positioning interventions. Lateral shoots represent branch-level growth hierarchy beyond primary shoot development, with their occurrence, timing, vigor, and management implications varying dramatically based on variety characteristics (some cultivars showing prolific lateral development, others minimal), vine vigor and balance, environmental conditions, and canopy management practices, requiring assessment whether lateral shoots contribute beneficially to vine function or create problems necessitating removal.
Lateral shoots are distinguished from primary shoots (arising from winter buds on permanent wood) by origin from axillary positions on current-season shoots, creating hierarchical growth pattern (primary shoots bearing lateral shoots which occasionally bear tertiary laterals). Lateral classification systems recognize timing distinctions (prompt/sylleptic same-season versus delayed/proleptic following-season), vigor variations (weak vestigial laterals versus strong competing laterals), and functional roles (beneficial photosynthetic supplementation versus problematic excessive density). Management strategies range from complete lateral removal (reducing canopy density and simplifying architecture), selective retention (keeping well-positioned laterals for photosynthetic benefits), to active lateral encouragement (certain training systems deliberately promoting lateral development). Understanding lateral shoot biology and appropriate management is important for optimizing canopy function, with decisions balancing photosynthetic contributions against density concerns, labor costs, and specific variety and site characteristics determining optimal lateral management approaches.
Achseltrieb
Nebentrieb
Seitentrieb
axillary shoot
branch shoot
lateral branch
secondary shoot
side shoot
Based on Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 113-145 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide.
Lateral Shoot
A mechanically hedged Merlot vineyard in Bordeaux demonstrates excessive Lateral_Shoot_Development_Quality with an average of 12 laterals per primary shoot reaching 30-40 cm length within three weeks of each hedging pass, requiring three additional hedging operations per season and extensive hand leaf-pulling in the fruit zone to maintain the canopy openness necessary for Botrytis control and optimal fruit maturation.
Lateral_Shoot_Development_Quality is a Vegetative_Growth_Vigor_Quality that inheres in a grapevine (Vitis vinifera L.) and represents the extent, timing, and vigor of secondary shoot (lateral) production from prompt buds located at primary shoot leaf axils. This quality significantly influences canopy density, light microclimate, disease pressure, and fruit quality, with excessive lateral development creating the dense, shaded canopy interiors associated with poor fruit quality and elevated disease incidence. In BFO terms, Lateral_Shoot_Development_Quality exists as a complex quality encompassing multiple measurable dimensions including lateral number per primary shoot, lateral length, lateral leaf area contribution, and the temporal pattern of lateral emergence throughout the season. The physiological regulation of lateral development involves the balance between apical dominance exerted by the primary shoot apex and the growth potential of prompt buds at each node, with disruption of apical dominance through hedging, hail damage, or shoot tip removal triggering rapid lateral proliferation that can contribute 30-50% of total canopy leaf area in vigorous vines. Cultivar variation in lateral development tendency is pronounced, with high-lateral varieties like Merlot, Sauvignon Blanc, and Sémillon requiring more intensive canopy management than low-lateral varieties like Riesling and Gewürztraminer. The timing of lateral emergence carries important implications for fruit quality, with early-emerging laterals contributing positively to total photosynthetic capacity during fruit development, while late-emerging laterals following hedging operations provide less benefit and may actively compete with ripening fruit for carbohydrate resources. Strategic lateral management through selective removal (lateral positioning, leaf plucking in the fruit zone) versus retention (maintaining laterals in upper canopy for continued photosynthesis) represents a key canopy management decision balancing light penetration, disease control, photosynthetic capacity, and labor costs.
Lateral_Shoot_Development_Quality represents the secondary dimension of vegetative growth vigor, distinct from primary shoot growth but intimately connected through the physiological mechanisms of apical dominance and resource allocation. This class is particularly relevant for understanding the consequences of hedging operations, which by removing shoot apices stimulate lateral proliferation, potentially creating a cycle of vigor expression requiring repeated intervention. The class relates to canopy microclimate through its direct influence on canopy density and interior shading.
Geiztriebbildung
Geiztriebentwicklung
Nebentriebwachstum
Seitentriebentwicklung
feminelle development
lateral shoot growth
lateral vigor
prompt shoot development
secondary shoot development
Intrieri C, Poni S, Lia G, Gómez del Campo M (2001) Vine performance and leaf physiology of conventionally and minimally pruned Sangiovese grapevines. Vitis 40:123-130 Bernizzoni F, Civardi S, Van Zeller M, Gatti M, Poni S (2011) Shoot thinning effects on seasonal whole-canopy photosynthesis and vine performance in Vitis vinifera L. cv. Barbera. Australian Journal of Grape and Wine Research 17:351-357
Lateral Shoot Development Quality
Investigation of Lateral_Shoot_Proliferation_Disposition at AWRI Adelaide documented extreme branching in cultivars recognized for prolific lateral development. Reference cultivars 'Carignan', 'Barbera', 'Colombard', and 'Silvaner' were grown without lateral management to characterize natural branching expression. Lateral_Shoot_Proliferation_Disposition cultivars showed consistent extreme characteristics. 'Carignan' exhibited archetypal proliferation with mean of 14.5 lateral shoots per primary shoot of 210 cm, occurring at 92% of nodes (essentially every node producing a lateral), with total lateral length of 245 cm exceeding primary length. Laterals developed vigorously with mean length of 17 cm and maximum exceeding 40 cm, and secondary laterals (laterals arising from primary laterals) were common (2.8 per primary shoot). The lateral/primary biomass ratio reached 1.15, indicating that lateral growth dominated resource allocation. 'Barbera' showed similar proliferation (12.8 laterals, 215 cm total length) with particularly vigorous secondary lateral development. Apex auxin concentration in Lateral_Shoot_Proliferation_Disposition cultivars averaged only 145 ng/g FW (50% below strong dominance types), while lateral bud cytokinin sensitivity was 2.8-fold higher, explaining the escape from correlative inhibition. Canopy density reached 3.8 leaf layers in unmanaged proliferating cultivars, severely compromising fruit zone light (< 5% ambient) and requiring intensive management intervention. Labor requirements for lateral management reached 85-95 hours/ha (4-5 removal passes) to achieve acceptable canopy density, representing a major management cost for these cultivars.
A Lateral_Shoot_Proliferation_Disposition is a Branching_Pattern_Disposition that inheres in a Vitis vinifera plant and is realized through minimal apical dominance and high intrinsic lateral bud activity, producing abundant lateral shoots from most or all nodes of the primary shoot, creating highly branched, complex shoot architecture and dense, multi-layered canopies without management intervention. This disposition represents the extreme of lateral development, characteristic of cultivars where correlative inhibition is largely ineffective, allowing lateral buds to activate freely regardless of apex activity. Cultivars with Lateral_Shoot_Proliferation_Disposition typically produce more than 10 lateral shoots per primary shoot, often with laterals at every node, and total lateral length may exceed primary shoot length as lateral growth captures a large proportion of vine resources. The viticultural challenges of lateral proliferation are substantial—without intensive management, canopies become excessively dense with 3+ leaf layers, severe fruit zone shading, poor air circulation promoting disease, and competition between lateral growth and fruit development. Management of proliferating cultivars requires multiple lateral removal passes or acceptance of hedging approaches that remove lateral-bearing shoot portions. The disposition is associated with low apex auxin production and/or high cytokinin sensitivity of lateral buds, resulting in constitutive bud activation that occurs regardless of apical suppression signals.
Lateral_Shoot_Proliferation_Disposition is_a Branching_Pattern_Disposition representing extreme lateral development (> 10 laterals per primary shoot, often at every node). Characterized by low apex auxin, high bud cytokinin sensitivity, and ineffective correlative inhibition. Examples include 'Carignan', 'Barbera', 'Colombard', 'Silvaner', 'Trebbiano'. Secondary laterals common. Creates severe canopy management challenges with dense, multi-layered canopies. Requires intensive lateral removal (4-5 passes) or hedging approaches. Total lateral length may exceed primary shoot length.
Geiztrieb-Proliferations-Disposition
Starke-Geiztrieb-Bildung
excessive lateral growth
prolific branching
vigorous lateral production
Übermäßige-Verzweigung
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles, Adelaide Lebon E et al. (2006) Shoot development in grapevine (Vitis vinifera L.) is affected by the modular branching pattern of the stem and intra- and inter-shoot trophic competition, Annals of Botany 98:1045-1056 Intrieri C, Poni S (1995) Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality of mechanized Italian vineyards, American Journal of Enology and Viticulture 46:116-127
Lateral Shoot Proliferation Disposition
In Grenache vineyards of Châteauneuf-du-Pape at BBCH 57-59, lateral tertiary inflorescence branches on lower secondary branches of basal primary branches can reach 1.2-2 cm length and bear 8-14 flowers each, creating localized high-density zones within the characteristically large, loose clusters of 800-1200 berries (final berry number after fruit set and shot berry elimination). Carignan in Languedoc exhibits well-developed lateral tertiary branches (0.8-1.6 cm, 6-10 flowers) on secondary branches of lower and basal primary branches, contributing to moderately loose cluster architecture. The hierarchical interaction of branch position effects (basal primary → lower secondary → lateral tertiary) creates maximum flower density zones that may require selective removal in premium wine production.
A lateral tertiary inflorescence branch is a tertiary inflorescence branch that arises from lateral or basal positions along a secondary inflorescence branch rather than from the terminal apex, typically exhibiting greater length, diameter, and flower number compared to terminal tertiary branches on the same secondary branch while contributing substantially to the internal architectural complexity of loose-clustered cultivars. Lateral tertiary branches occupy non-terminal positions along secondary branches, typically located 0.5-8 mm from the secondary branch base (junction with primary branch), with 1-3 lateral tertiaries commonly present on well-developed secondary branches in extensively branched cultivars. These branches represent earlier-initiated tertiary branches during inflorescence primordium development compared to terminal tertiaries and consequently show more advanced developmental elaboration. Branch length typically ranges from 0.4-2 cm in loose-clustered cultivars, with basal diameter often 0.3-0.6 mm, exceeding terminal tertiary dimensions by 30-80% on the same secondary branch. Lateral tertiary branches characteristically bear more flowers (4-15 flowers per lateral tertiary, typically 6-10 in extensively branched types) compared to terminal tertiaries (2-8 flowers), with flowers borne through direct attachment along the branch axis and, very rarely in extreme loose-clustered types, through minimal quaternary subdivision. The enhanced development of lateral tertiary branches creates localized zones of high flower density on the basal portions of secondary branches, particularly on lower secondary branches of basal and lower primary branches where the cumulative effect of favorable position at all hierarchical levels produces maximum architectural elaboration. Vascular anatomy of lateral tertiary branches shows slightly larger vessel diameters (20-35 μm) compared to terminal tertiaries (12-25 μm), facilitating enhanced resource delivery to support the greater flower complement. Lateral tertiary branch development proceeds 1-2 days earlier than terminal tertiaries on the same secondary during BBCH 55-59, creating temporal heterogeneity in flower development within individual secondary branches. The vigorous development of lateral tertiary branches on basal secondary branches of basal primary branches creates the maximum architectural complexity observed in grapevine inflorescence structure, characteristic of extremely loose-clustered cultivars like Grenache, where four levels of branching (rachis → primary → secondary → tertiary → flowers) combine with positional gradients at each level to produce clusters of 600-1500 berries with highly heterogeneous internal density patterns.
Lateral_Tertiary_Inflorescence_Branch is distinguished from Terminal_Tertiary_Inflorescence_Branch by position (lateral/basal versus terminal/apical on the parent secondary branch), enhanced dimensions, earlier development, and increased flower number. Multiple lateral tertiaries may occur on a single secondary branch, whereas terminal tertiary is singular by definition. These branches represent critical components of maximum architectural complexity in loose-clustered cultivars and contribute disproportionately to flower number within their parent secondary branches. The position-dependent cascade of developmental vigor across four hierarchical levels (primary position → secondary position → tertiary type → flower number) creates the complex spatial patterning of flower distribution that ultimately determines cluster morphology, disease susceptibility, and harvest characteristics in loose-clustered genotypes.
Seitenast dritter Ordnung
basal tertiary branch
lateraler Tertiärast
proximal third-order branch
seitlicher tertiärer Ast
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 93-94 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 173-178 Vasconcelos, M.C., Castagnoli, S. (2000). Leaf canopy structure and vine performance. American Journal of Enology and Viticulture 51=390-396.
Lateral Tertiary Inflorescence Branch
Merlot grafted on 3309 Couderc rootstock at 13 years showing lateral tertiary root system at 52 cm depth, third-order lateral tertiary roots 0.8 cm diameter emerging at 75° angle from second-order parent growing horizontally 9 cm length, fourth-order lateral tertiary roots 0.3 cm diameter branching from third-order at 5 cm intervals growing laterally 4 cm length, fifth and sixth-order fine lateral tertiary roots <2 mm diameter 1-2 cm length with abundant root hairs, lateral tertiary root network creating dense horizontal exploration estimated 24 cm total root length per cm³ soil, mycorrhizal colonization 46%, lateral tertiary roots providing majority of water and nutrient uptake from upper soil layer
Lateral tertiary root is tertiary root emphasizing lateral orientation and branching pattern growing sideways from parent root, characterized by emergence at angle typically 45-90° from parent root axis creating lateral rather than vertical or downward trajectory, predominantly horizontal growth exploring soil laterally at constant depth or with slight descent, third or higher branching order arising from secondary root parents or other tertiary roots, and serving to extend root system exploration laterally creating dense horizontal network in favorable soil layers particularly upper horizons where resources concentrated, with lateral tertiary roots showing similar orientation to parent roots maintaining lateral exploration pattern through successive branching generations, demonstrating typical dimensions of 0.2-1.5 cm diameter for third-order declining to <2 mm for higher orders and lengths 2-15 cm depending on order and vigor, producing further higher-order lateral branches creating hierarchical lateral network, and representing critical component of horizontal soil exploration complementing vertical primary root penetration creating three-dimensional resource acquisition architecture with lateral tertiary roots particularly abundant in nutrient-rich topsoil horizons. Lateral tertiary root orientation and trajectory includes= growing perpendicular to oblique from parent (lateral tertiary roots typically emerging 60-90° from parent root creating sideways growth), maintaining horizontal or slightly descending path (lateral trajectory predominantly sideways rather than downward typically 0-30° from horizontal), following parent orientation pattern (lateral tertiary roots generally maintaining lateral direction of parent secondary root), demonstrating weak gravitropism (lateral roots showing minimal downward gravitropic response allowing continued horizontal growth), and creating planar exploration (lateral tertiary roots in multiple directions creating disk-like or layered soil exploration pattern). Lateral tertiary root hierarchical position includes= typically third or fourth-order (lateral tertiary roots most commonly being third-order from second-order parents or fourth-order from third-order parents), occasionally fifth or higher order (some lateral tertiary roots in extensively branched systems reaching fifth through seventh order), arising from lateral secondary parents (lateral tertiary roots typically branching from lateral secondary roots maintaining lateral orientation through generations), producing higher-order lateral branches (lateral tertiary roots generating fourth fifth and sixth-order laterals continuing branching hierarchy), and demonstrating exponential abundance (lateral tertiary root numbers multiplying exponentially with each branching level). Lateral tertiary root depth distribution includes= concentrating at parent depth (lateral tertiary roots typically at similar depth to parent root from which emerged), showing upper soil preference (majority of lateral tertiary roots 20-80 cm depth where parent lateral secondary roots most abundant), exhibiting layer-specific distribution (lateral tertiary root density varying with soil horizon reflecting parent distribution and resource availability), demonstrating limited vertical range (individual lateral tertiary roots typically spanning <10 cm vertical range staying within soil layer), and creating horizontal stratification (lateral tertiary roots organized in horizontal layers at various depths reflecting parent root stratification). Lateral tertiary root length and extent includes= showing modest length (individual lateral tertiary roots typically 2-15 cm depending on order with third-order longer and higher orders progressively shorter), extending locally from parent (lateral tertiary roots exploring soil within 5-15 cm radius of parent root origin point), creating dense local network (numerous lateral tertiary roots from single parent creating dense exploration in local soil volume), demonstrating cumulative extent (collective lateral tertiary root network from entire vine potentially totaling hundreds of meters length), and limited by order (higher-order lateral tertiary roots showing progressively shorter lengths reflecting finer branching scale). Lateral tertiary root branching pattern includes= producing higher-order laterals (lateral tertiary roots generating fourth fifth and sixth-order branches continuing lateral orientation), showing close spacing (higher-order branches typically emerging 1-5 cm intervals reflecting fine branching scale), exhibiting prolific branching (lateral tertiary roots potentially producing 5-15+ higher-order branches depending on length and vigor), demonstrating acropetal gradient (branching density typically higher near growing tip declining toward base), and creating absorptive clusters (lateral tertiary root with higher-order branches forming dense absorptive unit in local soil). Lateral tertiary root absorptive function includes= providing direct water uptake (lateral tertiary roots absorbing water directly through epidermis and root hairs), demonstrating nutrient absorption (lateral tertiary roots taking up minerals from soil solution particularly nitrogen and potassium readily available in upper soil), showing root hair production (lateral tertiary roots particularly younger portions bearing abundant root hairs increasing absorption surface), hosting mycorrhizal colonization (lateral tertiary roots showing AM fungal associations enhancing phosphorus and water uptake), and contributing substantially to total uptake (lateral tertiary roots in upper soil providing major portion of vine water and nutrient acquisition). Lateral tertiary root lifespan and turnover includes= showing relatively short lifespan (lateral tertiary roots typically functioning 6 months to 2 years depending on order with higher orders shorter-lived), exhibiting seasonal production (lateral tertiary root initiation concentrated in spring and fall active growth periods), demonstrating continuous turnover (ongoing production and mortality of lateral tertiary roots throughout growing season and year), revealing stress sensitivity (lateral tertiary root mortality increasing under drought, nutrient deficiency, or pathogen pressure), and contributing to carbon cycling (lateral tertiary root death and decomposition providing organic matter input to soil particularly in upper horizons). Lateral tertiary root resource responsiveness includes= proliferating in resource patches (lateral tertiary root density increasing in localized zones of high nutrient or moisture availability), showing rapid growth response (lateral tertiary roots quickly elongating and branching when encountering favorable conditions), demonstrating avoidance behavior (lateral tertiary root growth reducing or ceasing in unfavorable zones with toxicity or severe limitations), revealing competitive interactions (lateral tertiary roots from neighboring vines or other plants competing for resources in shared soil volume), and enabling adaptive plasticity (lateral tertiary root architecture adjusting to spatial and temporal resource heterogeneity optimizing acquisition).
Lateral_Tertiary_Root emphasizes lateral orientation and horizontal exploration and is distinguished from vertical or descending roots by focus on sideways growth pattern. Includes Fine_Lateral_Tertiary_Root and Branching_Lateral_Tertiary_Root. Critical for understanding horizontal fine root architecture, upper soil exploitation, lateral branching pattern, layer-specific distribution, and the horizontally oriented high-order roots creating dense lateral exploration in resource-rich upper soil horizons.
Horizontal tertiary root
Horizontal verzweigte Feinwurzel
Lateral high-order root
Laterale Tertiärwurzel
Seitlich wachsende Wurzel höherer Ordnung
Sideways-branching fine root
Lateral Tertiary Root
A lateral vein on a mature leaf at node 7 of Vitis vinifera cv. Merlot in Bordeaux, France, one of five pairs of lateral veins branching from the median vein, branching from the median vein at 4.2 cm from petiolar junction at an angle of 58 degrees, extending 6.8 cm laterally through the second major lobe to within 0.5 cm of the lobe margin, diameter of 1.4 mm at the junction with median vein tapering to 0.6 mm near the distal end, prominent as raised ridge on abaxial surface approximately 0.3 mm above surrounding lamina, appearing as shallow groove on adaxial surface, light green coloration matching surrounding tissue, moderate pubescence on abaxial side, containing 5-7 xylem vessels (30-60 μm diameter) supporting water transport to the lobe region representing approximately 18% of total leaf area, substantial phloem facilitating photosynthate collection from the lobe, mechanical tissues providing structural support preventing lobe drooping under gravitational and wind loads, serving as primary axis for numerous second-order veins branching at 45-75 degree angles creating reticulate network throughout the lobe, measured at BBCH stage 75 (berries pea-sized) in late June, representing typical lateral vein architecture supporting major lobe vascularization and structure.
A major vein branching from the median vein and extending laterally into the blade lobes in Vitis vinifera, characterized by first-order hierarchical position branching directly from the central median vein, substantial diameter typically 0.8-2.0 mm representing second-largest vein size after median vein, prominent appearance as raised ridges on abaxial surface and shallow grooves on adaxial surface, complex vascular anatomy including multiple xylem vessels and substantial phloem supporting water delivery and photosynthate export for entire lobe regions, significant mechanical tissues providing structural support for blade lobes, typical occurrence of 4-6 pairs of lateral veins per leaf creating the characteristic palmate venation architecture, branching angles from median vein typically 40-80 degrees varying with cultivar and lobe configuration, extension from median vein junction through blade tissue to lobe margins or near-margin regions, and critical functional roles including serving as primary vascular pathways delivering water and nutrients to lobe regions, collecting and exporting photosynthates from large blade areas, providing mechanical support preventing lobe drooping, and serving as organizing axes for higher-order vein networks that create the complete reticulate venation pattern. Lateral veins represent the first branching order from the median vein, establishing the fundamental palmate architecture characteristic of grapevine leaves, with each lateral vein supplying a distinct lobe or blade sector creating functional vascular territories, and exhibiting cultivar-specific characteristics including number, branching angles, and prominence patterns that contribute to overall leaf architecture and ampelographic identity.
Lateral veins are distinguished from the median vein by branching origin rather than direct petiolar continuation, and from higher-order veins by first-order hierarchical position and large diameter. The number of lateral vein pairs shows cultivar specificity, with most varieties producing 4-5 pairs creating five-lobed leaves (including central terminal lobe), though some cultivars show 3 pairs (three-lobed) or 6 pairs (seven-lobed). Branching angles affect overall leaf shape, with acute angles creating narrow elongated lobes while obtuse angles create broader rounded lobes. Vein prominence varies among cultivars from highly raised conspicuous veins to relatively flat subtle veins, serving as useful ampelographic character. Lateral veins exhibit hierarchical organization as first-order veins from which second-order veins branch, creating the reticulate network through progressive branching to third, fourth, and higher orders. Functional organization creates vascular territories with each lateral vein supplying its associated lobe region, with some overlap at boundaries through anastomoses (vein connections) providing redundancy. Damage to individual lateral veins affects only the supplied lobe region unlike median vein damage potentially compromising entire blade function. Development involves coordinated initiation along the median vein during early leaf ontogeny, with lateral vein primordia appearing sequentially and developing outward establishing lobe patterns.
Haupt-Lateralvene
Seitenader
erste Ordnung Vene
first-order lateral vein
major lateral vein
primary branch vein
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 58-89 Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 56-94.
Lateral Vein
A leader main apical shoot on a vertical trunk of Vitis vinifera cv. Syrah in the Rhône Valley, France, where the original terminal main apical shoot was topped at 140 cm height in June to control excessive vigor, followed by development of three lateral shoots near the cut surface, with the most vigorous lateral growing an additional 60 cm vertically by August to reach the target trunk height of 200 cm, showing 9 nodes with progressively increasing internode length (8-12 cm) and complete lateral bud suppression in the new growth section, eventually becoming the established terminal leader for this trunk section.
A main apical shoot that has experienced intentional or accidental termination of its original apical meristem through topping, pinching, mechanical damage, or pathogen infection, followed by re-establishment of terminal growth through development of one or more lateral shoots near the point of termination, with one lateral assuming dominant apical position and continuing the primary growth axis in Vitis vinifera. Leader main apical shoots represent the vine's architectural resilience and capacity for growth reorganization following apical meristem loss, maintaining framework extension objectives despite interruption of the original terminal shoot. These shoots exhibit modified apical dominance compared to terminal main apical shoots, typically showing weaker suppression of lateral buds below the point of leadership assumption and often displaying altered phyllotaxis and internode length patterns that reflect the developmental transition from lateral to apical identity. Leader main apical shoots are particularly important in training system establishment where young vines must achieve target framework dimensions despite inevitable damage events, and in mature vineyards following severe weather damage or disease pressure that necessitates framework reconstruction.
Leader main apical shoots differ from terminal main apical shoots in their developmental history rather than current function, and differ from dominant apical shoots in maintaining the primary architectural axis rather than creating alternative growth directions. The quality of leadership establishment depends critically on timing of original shoot termination, with early-season topping (before bloom) typically resulting in strong leader development, while late-season damage often produces weak or multiple competing leaders. In training system management, deliberate creation of leader shoots through strategic topping represents a tool for controlling shoot vigor, inducing lateral branching, or correcting growth direction. Multiple competing leaders require selective removal to re-establish singular apical dominance.
Ersatzspitzentrieb
Leitendtrieb
re-directed main shoot
replacement apical shoot
resumed terminal shoot
topped and re-grown apical shoot
umgelenkter Haupttrieb
Based on Cline (1997) "Concepts and terminology of apical dominance" American Journal of Botany 84=1064-1069 Poni et al. (2006) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 57=397-407 Intrieri et al. (2008) "Grapevine growth and development" in The Grapevine= An Overview, ISHS Press pp. 44-67 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 189-214.
Leader Main Apical Shoot
A leaf at node 8 on a fruiting shoot of Vitis vinifera cv. Cabernet Sauvignon in Bordeaux, France, exhibiting typical five-lobed palmate architecture with deep U-shaped petiolar sinus, medium-depth lateral sinuses, acute lobe apices with prominent acuminate tips, regularly distributed sharp teeth along margins, moderate pubescence on abaxial veins creating light gray-green undersurface coloration, lamina measuring 14 cm length by 13 cm width at maximum dimensions, dark green adaxial surface with slightly glossy appearance from cuticle development, positioned in mid-canopy receiving approximately 40-60% full sunlight creating intermediate sun-shade characteristics, exhibiting active photosynthesis contributing to fruit ripening in adjacent cluster at same node, measured at BBCH stage 81 (beginning of berry ripening) in early August, representing the characteristic leaf morphology diagnostic for Cabernet Sauvignon including the deeply indented sinuses and pronounced lobe development distinguishing this cultivar from related varieties.
A dorsiventrally flattened lateral determinate organ arising from shoot nodes in Vitis vinifera, characterized by a distinctive palmate architecture with 3-7 primary lobes separated by sinuses of variable depth, a complex venation pattern with major veins radiating from the petiolar junction and anastomosing secondary and tertiary veins creating a reticulate network, specialized photosynthetic tissue organized in palisade and spongy mesophyll layers, stomata primarily on the abaxial (lower) surface enabling gas exchange for photosynthesis and transpiration, and cultivar-specific morphological features including leaf shape, lobe number and depth, sinus characteristics, tooth morphology, pubescence patterns, and pigmentation that serve as primary ampelographic diagnostic characters for variety identification. Leaves represent the principal photosynthetic organs of the grapevine, converting solar radiation into chemical energy through carbon fixation, producing the carbohydrates that support all vine growth, fruit development, and reserve accumulation, with total leaf area and photosynthetic efficiency determining vine productive capacity and fruit quality potential. The grapevine leaf exhibits remarkable phenotypic plasticity in response to environmental conditions, with sun-exposed leaves developing thicker cuticles, increased palisade layers, higher chlorophyll density, and smaller size compared to shade leaves showing opposite characteristics, creating functional specialization across the canopy optimizing light capture and carbon gain under heterogeneous light environments typical of complex three-dimensional vine canopies.
Leaves are distinguished from other lateral organs including tendrils (modified inflorescences) and inflorescences (reproductive structures) by their photosynthetic function, dorsiventrally flattened lamina architecture, and alternate phyllotaxy at shoot nodes. Grapevine leaves show extreme morphological variation both among cultivars (serving as primary ampelographic characters) and within individual vines (phenotypic plasticity responding to light, water, nutrients). Classification of individual leaves requires assessment of developmental stage (juvenile, mature, senescent), environmental exposure (sun, shade, intermediate), functional status (photosynthetically active, damaged, diseased), and position (basal, median, apical shoot zones). Leaf morphology including lobe pattern, sinus depth, tooth characteristics, and pubescence represents the most important visual diagnostic feature system for cultivar identification, with standardized descriptors developed by OIV (International Organisation of Vine and Wine) enabling systematic characterization and comparison.
Laubblatt
Rebblatt
Weinblatt
foliage leaf
grape leaf
grapevine leaf
vine leaf
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 1-57 This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 33-55.
Leaf
Leaf absorptance measurement in Merlot comparing healthy leaves (absorptance 0.86, chlorophyll 485 μmol/m²) with magnesium-deficient chlorotic leaves (absorptance 0.68, chlorophyll 195 μmol/m²) quantifies the light capture penalty from nutrient deficiency, with the 21% absorptance reduction directly diminishing photosynthetic capacity under light-limited conditions and indicating that chlorotic leaves waste substantial incident light through increased reflectance and transmittance, demonstrating how absorptance measurement provides the optical basis for understanding photosynthetic limitation in stressed vines.
Leaf absorptance quality is a light capture efficiency quality that represents the fraction of incident photosynthetically active radiation (PAR) absorbed by grapevine leaves, characterizing the integrated optical property arising from pigment content, leaf structure, and surface characteristics that determines how much light energy enters the photosynthetic apparatus. Within the BFO framework, this quality specifically addresses the bulk light absorption dimension of leaf optics, distinguishing absorbed light from reflected and transmitted fractions while recognizing that absorptance represents the net input to photosynthesis that pigment content and leaf anatomy jointly determine. The ontological significance lies in leaf absorptance providing the direct measure of light energy available for photochemistry, where absorptance values (typically 0.80-0.90 for healthy grapevine leaves) indicate the efficiency of light capture from the environment and enable calculation of absorbed quantum flux density essential for photosynthetic efficiency calculations and canopy light interception models. The physical basis for leaf absorptance involves the interaction of incident light with leaf optical properties, where light either reflects from surfaces (cuticular reflection, internal scattering back through surface), transmits through the leaf (escaping from lower surface), or is absorbed by pigments and other chromophores, with the absorbed fraction available for photosynthesis or thermal dissipation. The compositional and structural factors affecting absorptance include total chlorophyll content (primary determinant of absorption), carotenoid content (contributing blue-light absorption), leaf thickness and mesophyll structure (affecting internal light path length), surface wax and trichomes (affecting reflectance), and leaf water content (affecting near-infrared absorption outside PAR). The measurement approaches for absorptance include integrating sphere measurements (direct determination of all optical fractions), calculation from chlorophyll content using empirical relationships, and remote sensing approaches inferring absorptance from reflectance spectra. Understanding leaf absorptance quality enables accurate photosynthetic efficiency calculation, canopy light interception modeling, and assessment of factors limiting light capture in the vineyard.
Leaf_Absorptance_Quality addresses the net fraction of PAR captured by leaves for photosynthesis or dissipation. This quality integrates pigment content with leaf structural properties to determine actual light input. The class supports photosynthetic efficiency calculations and canopy modeling requiring accurate light absorption estimates.
Blattabsorption
Lichtabsorptionsfraktion
PAR absorption
PAR-Absorption
absorbed PAR fraction
absorbierte Strahlung
leaf optical absorption
light absorption fraction
Vogelmann TC (1993) Plant tissue optics. Annual Review of Plant Physiology and Plant Molecular Biology 44(1): 231-251 Bauerle WL et al. (2004) Leaf absorptance of photosynthetically active radiation in relation to chlorophyll meter estimates among woody plant species. Scientia Horticulturae 101(1-2): 169-178
Leaf Absorptance Quality
Leaf angle adjustment quality evaluation in Douro Valley Portuguese varieties assessed the contribution of this mechanism to heat tolerance under the region's hot summer conditions where ambient temperatures frequently exceed 40°C. Time-lapse photography tracking leaf orientation throughout high-radiation days revealed substantial variation in leaf angle adjustment quality among cultivars. Touriga Nacional demonstrated excellent leaf angle adjustment quality with lamina angle shifting from 25° above horizontal in morning to 70° (nearly vertical) at solar noon, reducing direct radiation interception by an estimated 65% and achieving leaf temperature reduction of 4.2°C compared to fixed-angle reference leaves. Touriga Franca showed moderate leaf angle adjustment quality with maximum deflection to 55°, achieving 45% interception reduction and 2.8°C cooling. Tinta Roriz (Tempranillo) exhibited limited leaf angle adjustment quality with only 15° change, providing minimal radiation avoidance benefit. The leaf angle adjustment quality differential translated to photosynthetic maintenance, with Touriga Nacional maintaining 75% of optimal photosynthesis at midday during heat events versus only 50% in the minimally-adjusting Tinta Roriz, demonstrating functional benefit of the avoidance mechanism. The rapid response (complete adjustment within 30 minutes of radiation change) confirmed active motor control rather than passive wilting, distinguishing true leaf angle adjustment quality from drought-induced leaf drooping.
Leaf angle adjustment quality is a heat avoidance quality that inheres in a grapevine cultivar or clone and represents the capacity to modify leaf orientation in response to excessive solar radiation or high temperatures, reducing the direct interception of solar radiation on the leaf surface through paraheliotropic (sun-tracking to avoid perpendicular incidence) or diaheliotropic (sun-avoiding) movements that decrease the radiation load and consequent temperature increase on leaf tissues. This quality captures an active morphological heat avoidance mechanism that enables real-time adjustment of leaf orientation to minimize heat stress, with the magnitude and speed of leaf angle adjustment determining the effectiveness of this response for reducing tissue temperatures during periods of excessive solar radiation coinciding with high ambient temperatures. The ontological significance of leaf angle adjustment quality lies in its role as an immediately responsive mechanism for reducing radiation load without requiring the water expenditure of transpirational cooling, providing heat avoidance benefit that complements evaporative cooling and may be particularly valuable when water availability limits transpiration capacity. The biological basis of leaf angle adjustment encompasses the pulvinar motor tissues at the junction between petiole and lamina that enable rapid orientation changes through turgor-mediated cell volume changes, combined with the signal transduction pathways that translate light and temperature perception into motor cell responses controlling leaf position. The extent of leaf angle adjustment capacity varies among grapevine genotypes and is modulated by developmental stage, with young expanding leaves typically showing greater adjustment capacity than mature leaves where pulvinar tissues may become less responsive. The functional benefit of leaf angle adjustment depends on the radiation environment, with substantial temperature reduction achievable when direct solar incidence is high but limited benefit under diffuse radiation conditions where leaf orientation has less effect on interception. The interaction between leaf angle adjustment and other heat avoidance mechanisms creates opportunities for both additive and redundant protection, with vines potentially employing multiple avoidance strategies simultaneously to achieve effective tissue temperature management under severe heat events.
Leaf_Angle_Adjustment_Quality is_a Heat_Avoidance_Quality, representing the morphological mechanism of leaf orientation change for radiation avoidance. This class addresses active sun-avoiding leaf movement controlled by pulvinar tissues. The angle change from horizontal baseline and resulting radiation interception reduction provide quantification. Distinguished from drought-induced wilting by the active motor control and rapid reversibility. Temperature reduction of 3-5°C can be achieved through effective angle adjustment. The mechanism is water-conservative, complementing transpirational cooling when water is limiting. Critical for understanding avoidance strategy variation among cultivars.
Blattorientierungsanpassung
Blattwinkelanpassung
Paraheliotropismus
leaf movement response
leaf orientation adjustment
paraheliotropism
solar avoidance movement
solare Vermeidungsbewegung
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 453-507 Wahid A et al. (2007) Heat tolerance in plants: an overview. Environmental and Experimental Botany 61:199-223 Ehleringer J, Forseth I (1980) Solar tracking by plants. Science 210:1094-1098
Leaf Angle Adjustment Quality
Optimal Riesling production in the Rheingau requires careful management of leaf area to fruit ratio, with target values of 12-15 cm² leaf area per gram of fruit achieved through winter pruning to 6-8 buds per square meter, spring shoot thinning to remove double shoots and weak laterals, and maintenance of approximately 1.0-1.4 m² main leaf area plus 0.3-0.5 m² lateral leaf area per vine supporting crop loads of 1.2-1.5 kg per vine for optimal must weights of 80-95° Oechsle.
Leaf_Area_Quality represents the two-dimensional extent of grapevine leaf blade surfaces that constitute the primary photosynthetic and transpirational interface of the canopy. This quality is arguably the most important single measurement in grapevine ecophysiology because leaf area directly determines the capacity for carbon assimilation, and through the source-sink relationship, ultimately constrains fruit yield and quality potential. Leaf area encompasses both individual leaf blade area and cumulative canopy leaf area, with different implications for physiological modeling. Individual leaf area typically ranges from 50-250 cm² depending on cultivar, shoot position, light environment, and vine water status. Total vine leaf area at véraison ranges from 2-8 m² depending on training system and vigor, while optimal values for balanced fruit ripening approximate 0.8-1.2 m² per kg of fruit. Leaf area index (LAI), the ratio of total leaf area to ground area, typically ranges from 1.5-5.0 in productive vineyards, with optimal values around 2.5-3.5 for quality wine production. Leaf area measurement methods include direct planimetry using leaf area meters with optical sensors, gravimetric estimation using calibrated specific leaf area relationships (area per unit dry mass), allometric models relating area to linear dimensions (length × width × shape factor), and remote sensing approaches for canopy-scale estimation. The dynamic development of leaf area through the season, with rapid expansion during spring growth followed by stabilization after véraison, determines the temporal pattern of canopy function.
Leaf_Area_Quality specifically addresses the laminar photosynthetic surface and is the most frequently measured area quality in grapevine research. The class encompasses both one-sided (projected) leaf area, which is the standard measurement convention, and total (two-sided) leaf area where gas exchange from both surfaces is relevant. Main leaves (those at primary shoot nodes) and lateral leaves (on secondary shoots) may be distinguished due to their different photosynthetic characteristics and contributions to the canopy. Exposed leaf area (receiving direct sunlight) versus interior leaf area (shaded) represents a functionally important distinction for canopy light interception models.
Blattflächenqualität
Laubfläche
blade area
foliar area
laminar extent
photosynthetic surface area
photosynthetische Oberfläche
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 119-135 Poni S et al. (2006) Effects of Early Defoliation on Shoot Photosynthesis, Yield Components, and Grape Composition. American Journal of Enology and Viticulture 57(4): 397-407
Leaf Area Quality
Leaf area-to-fruit ratio assessment comparing canopy management treatments in Sauvignon Blanc reveals the source-sink effect on ripening outcomes, with excessive defoliation treatment (0.5 m²/kg, severe source limitation) showing delayed ripening, maximum Brix only 21.5, and green herbaceous character persisting, standard defoliation (0.9 m²/kg, adequate balance) achieving 23.5 Brix with varietal tropical-citrus character developing fully, and no defoliation (1.4 m²/kg, excess source) reaching 24.0 Brix but with shaded fruit showing reduced thiol precursors and muted varietal expression, demonstrating how leaf area-to-fruit ratio affects both ripening capacity and fruit quality outcomes.
Leaf area-to-fruit ratio quality is a yield efficiency quality that represents the relationship between photosynthetically active leaf surface and fruit production, characterizing the source-sink balance in functional terms by quantifying how much leaf area supports each unit of fruit, with this ratio determining whether adequate photosynthetic capacity exists for full fruit development and ripening. Within the BFO framework, this quality specifically addresses the leaf-fruit relationship dimension of efficiency, distinguishing this physiologically-based metric from the empirically-derived Ravaz Index while recognizing that leaf area-to-fruit ratio directly captures the functional relationship between carbon fixation capacity (leaves as source) and carbon demand (fruit as sink), providing more mechanistic insight into vine balance than weight-based ratios. The ontological significance lies in leaf area-to-fruit ratio determining the photosynthate supply available per unit of fruit, where adequate leaf area ensures sufficient carbohydrate production for sugar accumulation, phenolic synthesis, and flavor compound development, while insufficient leaf area creates source limitation that delays ripening, reduces sugar accumulation, and may compromise flavor and phenolic development even if berries eventually reach minimum maturity thresholds. The optimal ranges for leaf area-to-fruit ratio typically fall between 0.8-1.2 m² leaf area per kg fruit for quality wine production (or equivalently 8-12 cm² per gram fruit), though values vary with variety (some varieties more efficient than others), climate (cooler climates requiring more leaf area per unit fruit due to lower photosynthetic rates), and training system (divided canopies potentially more efficient than single canopies). The measurement methodology involves estimating leaf area (through direct measurement, allometric relationships with shoot parameters, or image analysis) and relating it to fruit weight, providing data that enables targeted management through either leaf area manipulation (canopy management affecting source) or fruit load adjustment (cluster thinning affecting sink). Understanding leaf area-to-fruit ratio quality enables physiologically-informed crop load assessment, identification of source limitation as a cause of ripening problems, and canopy management decisions targeting appropriate leaf-fruit balance.
Leaf_Area_To_Fruit_Ratio_Quality addresses the physiologically-based source-sink balance between photosynthetic capacity and fruit demand. This quality provides mechanistic assessment of ripening capacity. The class supports canopy and crop load management through leaf-fruit ratio characterization.
Blatt-Frucht-Verhältnis
Blattfläche-Frucht-Verhältnis
Kronenertragsbilanz
Quellen-Senken-Verhältnis
canopy-crop balance
leaf-fruit ratio
photosynthetic load
source-sink ratio
Kliewer WM, Dokoozlian NK (2005) Leaf area/crop weight ratios of grapevines: influence on fruit composition and wine quality. American Journal of Enology and Viticulture 56(2): 170-181 Howell GS (2001) Sustainable grape productivity and the growth-yield relationship: a review. American Journal of Enology and Viticulture 52(3): 165-174
Leaf Area To Fruit Ratio Quality
A leaf blade on a mature leaf at node 7 of Vitis vinifera cv. Sangiovese in Chianti, Italy, measuring 15.2 cm length by 14.5 cm width representing typical dimensions for this cultivar at mid-shoot position, exhibiting characteristic five-lobed palmate architecture with moderate to deep sinuses, bilateral symmetry about the central median vein with subtle asymmetry showing slightly larger left lateral lobe, total surface area of 142 cm² providing photosynthetic capacity, lamina thickness of 285 μm including adaxial epidermis (15 μm), two palisade layers (total 82 μm), spongy mesophyll (165 μm), and abaxial epidermis (23 μm), dark green adaxial coloration from chlorophyll concentration of 475 μg cm⁻², complex reticulate venation with median vein and four major lateral veins supporting blade structure, marginal teeth with alternating large and small patterns diagnostic for this cultivar, moderate flexibility allowing 120-degree blade rotation in wind responses, demonstrating light-saturated photosynthesis rate of 12.8 μmol CO₂ m⁻² s⁻¹ typical of sun-exposed mature leaves, measured at BBCH stage 81 (beginning of berry ripening) in early August, representing the complete leaf blade structure providing essentially all photosynthetic function for this leaf.
The flattened expanded photosynthetic portion of the grapevine leaf excluding the petiole in Vitis vinifera, characterized by dorsiventrally flattened lamina architecture optimized for light interception, palmate structure with 3-7 lobes radiating from the petiolar junction, typical dimensions of 8-20 cm length and width varying with cultivar, position, and environmental conditions, bilateral symmetry about the midrib though often with subtle asymmetries, complex internal anatomy including adaxial (upper) epidermis, multiple mesophyll layers with palisade and spongy tissue, abaxial (lower) epidermis with high stomatal density, and intricate reticulate venation network, green coloration from chlorophyll concentrations typically 300-600 μg cm⁻² varying with development and light exposure, and cultivar-specific morphological features including outline shape, lobe number and depth, sinus characteristics, marginal teeth patterns, surface textures, and pigmentation serving as primary ampelographic diagnostic characters. The leaf blade constitutes the primary photosynthetic organ, containing 85-95% of total leaf chlorophyll and accounting for essentially all leaf photosynthetic capacity, with structural features optimized for light capture through large flat surface area, gas exchange through abundant stomata concentrated on abaxial surfaces enabling CO₂ uptake and water vapor loss, and mechanical integrity through vein networks and specialized tissues supporting blade structure against gravitational and wind forces while maintaining flexibility for movement responses.
Leaf blade is distinguished from the complete leaf by excluding the petiole, focusing specifically on the expanded photosynthetic lamina portion. The blade constitutes the functional photosynthetic component, while the petiole serves mechanical support and vascular connection roles. Blade morphology shows extreme variation among cultivars, serving as the foundation of ampelographic identification systems, with standardized descriptors characterizing outline shape, lobe patterns, sinus depths, marginal features, surface characteristics, and venation patterns. Environmental plasticity creates substantial within-cultivar variation through sun versus shade leaf differentiation, with blade thickness, cell layering, and anatomical organization varying dramatically based on light exposure during development. Position effects generate systematic patterns with basal leaves typically showing largest blades, median leaves intermediate dimensions, and apical leaves smallest blades reflecting resource allocation and developmental timing. Blade orientation and angle relative to incident light critically affect photosynthetic efficiency, with optimal positioning maximizing light interception while avoiding photoinhibition under excessive irradiance. The term "lamina" is often used synonymously with leaf blade, both referring to the flattened expanded portion excluding petiole.
Lamina folii
blade
foliar lamina
leaf lamina
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 1-57 Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Patakas et al. (2003) "Differences in stomatal and photosynthetic responses of Vitis vinifera L. (cv. Sultana) leaves in response to progressive water stress" Vitis 42=113-115 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 33-55.
Leaf Blade
The leaf blade description of Nebbiolo documents a strongly blistered (bullate) adaxial surface (OIV 075:7) with pronounced convex interveinal areas creating the deeply puckered, rugose texture that is one of Nebbiolo's most recognizable leaf features — the blistering results from differential growth between the rapidly expanding interveinal mesophyll and the constrained growth along the veins, pushing the interveinal tissue upward on the adaxial surface and creating corresponding concavities on the abaxial surface. Combined with the involute blade profile (margins curving downward, OIV 073:3), strong undulation of the blade margin (OIV 074:7), and the thick leathery texture reflecting the multi-layered palisade mesophyll, Nebbiolo's leaf blade creates a three-dimensional architecture immediately distinguishable from the flat, smooth blades of Barbera (OIV 075:3) and the moderately blistered, revolute blades of Dolcetto (OIV 075:5, OIV 073:5).
A mature leaf description component that comprehensively documents the surface characteristics, structural features, and textural properties of the fully expanded leaf blade, recording blistering/convexity of the interveinal areas (OIV 075), undulation of the blade between veins and margin (OIV 074), profile of the blade in cross-section (OIV 073), and the overall plane versus involute versus revolute configuration that collectively characterize the three-dimensional architecture of the mature leaf surface beyond the two-dimensional outline captured by shape descriptors.
Leaf_Blade_Description captures the three-dimensional surface characteristics that complement the two-dimensional shape, size, and outline descriptors, providing cultivar identification information that is frequently underutilized because surface texture is more difficult to photograph, measure, and communicate than outline shape. The blistering character (OIV 075) in particular is among the most diagnostically valuable leaf characters, ranking alongside blade shape and petiolar sinus form in cultivar discrimination power — yet its scoring requires direct observation of the physical leaf (photographs flatten the three-dimensional surface, losing diagnostic detail) and experienced tactile assessment of leaf texture and thickness: this requirement for in-person observation makes blade surface descriptors less amenable to the image-based digitization and remote assessment technologies being developed for automated ampelography. The genetic basis of leaf surface architecture has been partially elucidated through QTL mapping — Chitwood et al. (2014) identified loci on chromosomes 1, 5, and 14 affecting blade curvature and surface topography, with the blistering phenotype likely reflecting differential cell division rates and expansion patterns between vein-associated and interveinal tissue regions during the 3-4 week leaf expansion period.
Blattoberflächen-Charakterisierung
Blattspreiten-Beschreibung
Lamina-Architektur-Dokumentation
blade surface characterization
lamina architecture record
leaf texture description
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Chitwood DH et al. (2014) A modern ampelography: a genetic basis for leaf shape and venation patterning in grape, Plant Physiology 164: 259-272
Leaf Blade Description
The leaf color description of Cabernet Sauvignon documents dark green adaxial surface color during the growing season (OIV 069:7) with distinctly red-pigmented main veins on the adaxial surface — the anthocyanin accumulation in vein tissue producing red-violet coloration that extends along the petiole and the proximal third of the five main veins, one of Cabernet Sauvignon's most recognizable mid-season leaf features. At senescence (OIV 093), Cabernet Sauvignon develops predominantly red autumn coloration with undertones of orange and yellow, contrasting with the yellow autumn color of Chardonnay (OIV 093:1, no anthocyanin synthesis during senescence, chlorophyll degradation revealing carotenoids), the mixed red-yellow autumn color of Merlot (OIV 093:4, partial anthocyanin synthesis), and the dramatic deep crimson autumn foliage of Pinot Noir (OIV 093:6, strong senescence-associated anthocyanin production).
A mature leaf description component that records the color characteristics of the fully expanded leaf blade during the growing season and at senescence, documenting the green intensity and hue of the adaxial surface (OIV 069), the color of the main veins on the adaxial surface including anthocyanin pigmentation of vein tissue (OIV 070), and the distinctive autumn coloration pattern (OIV 093) that develops during leaf senescence as chlorophyll degrades and anthocyanin, carotenoid, and flavonoid pigments are revealed or synthesized, providing cultivar-diagnostic color characters at two distinct phenological stages.
Leaf_Color_Description captures two temporally distinct color phenotypes — the growing-season green intensity and vein pigmentation observed from full expansion through véraison, and the autumn senescence coloration observed in the weeks before leaf fall. These two color assessments provide independent cultivar-diagnostic information because they are controlled by different genetic and physiological mechanisms: growing-season green intensity reflects chlorophyll concentration (determined by nitrogen nutrition, light exposure, and leaf age in addition to genotype), vein anthocyanin reflects constitutive anthocyanin pathway activity in vascular tissue (strongly genetically determined, correlating with berry skin color in most cultivars but not all), and autumn coloration reflects the balance between chlorophyll degradation (revealing pre-existing carotenoids, producing yellow) and de novo anthocyanin synthesis during senescence (producing red), with the anthocyanin-producing capacity during senescence varying independently from the constitutive anthocyanin production that determines berry color. White-berried cultivars can produce red autumn foliage (e.g., Pinot Blanc, which retains senescence-specific anthocyanin pathway activation despite the loss of berry skin anthocyanin through VvMYBA1 mutation).
Blatt-Pigmentierungs-Dokumentation
Blattfarben-Beschreibung
Blattspreiten-Färbung
blade color characterization
foliar color documentation
leaf pigmentation record
Leaf Color Description
In Pinot Noir vineyards of the Côte d'Or, Burgundy, leaf detachment typically commences in early November following completion of abscission zone activation, with 50% leaf fall achieved by mid-November when minimum temperatures consistently reach 2-4°C, and complete defoliation occurring by late November to early December under normal seasonal conditions corresponding to post-BBCH 97 developmental stage.
Leaf detachment in Vitis vinifera constitutes the terminal phase of the leaf fall process, representing the physical separation of senescing leaves from the vine following completion of abscission zone activation and cell separation processes. This process occurs during late autumn, typically from late October through November in Northern Hemisphere viticulture, when the abscission zone at the petiole base has undergone complete cell wall dissolution and middle lamella degradation, allowing mechanical separation of leaf tissue from shoot attachment points. The detachment process involves the final rupture of remaining vascular connections and residual cellular attachments that persisted through the cell separation phase, with actual leaf fall triggered by environmental factors including wind, rainfall, frost events, and gravitational forces acting on weakened attachment points. Physiologically, leaf detachment represents the culmination of programmed senescence processes that began 4-8 weeks earlier with chlorophyll degradation and nutrient remobilization, during which period the vine systematically recovered nitrogen, phosphorus, potassium, and other mobile nutrients from leaf tissues for storage in permanent woody structures. The timing of leaf detachment varies considerably among cultivars, with early-ripening varieties such as Chasselas and Pinot Noir typically completing leaf fall 2-3 weeks before late-ripening cultivars like Cabernet Sauvignon and Grenache. Temperature plays a critical role in detachment timing, with accelerated leaf fall occurring when minimum temperatures drop below 5°C for consecutive nights, while mild autumn conditions can delay complete defoliation by several weeks. The process exhibits spatial patterns within the canopy, generally progressing from basal leaves that senesced earliest toward apical positions, though environmental exposure and shading history influence individual leaf detachment timing. From a viticultural perspective, the progression and completion of leaf detachment provides important indicators of vine dormancy preparation and carbohydrate reserve status, with delayed or incomplete leaf fall sometimes indicating nutritional imbalances or disease pressure that compromised normal senescence signaling. The physical mechanics of detachment involve separation at the abscission zone fracture plane, where specialized separation layer cells have dissolved their shared cell walls, leaving a clean break surface that rapidly suberizes to protect exposed shoot tissues from pathogen entry and desiccation.
Leaf_Detachment represents the final physical separation phase within the Leaf_Fall hierarchy, occurring after Petiole_Abscission processes have prepared the separation zone. Distinguished from Leaf_Senescence which encompasses biochemical degradation processes, and from Petiole_Abscission which specifically addresses abscission zone formation and activation. This parent class encompasses all temporal phases from initial to complete detachment, with duration typically spanning 3-6 weeks depending on cultivar and environmental conditions. Subdivides into Initial_Leaf_Detachment, Progressive_Leaf_Fall, Mass_Leaf_Fall, and Complete_Leaf_Detachment based on temporal progression and proportion of canopy affected.
Blattabfall
Blattablösung
Laubfall
autumn leaf fall
herbstlicher Blattfall
leaf abscission completion
leaf separation
leaf shedding
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Mullins MG, Bouquet A, Williams LE (1992) Biology of the Grapevine Cambridge University Press Zapata C et al (2004) Partitioning and mobilization of starch and N reserves in grapevine. Journal of Experimental Botany 55: 2165-2175
Leaf Detachment
Comprehensive characterization of Leaf_Development_Disposition at INRAE Vassal ampelographic collection assessed leaf parameters across 250 cultivars using standardized OIV protocols for mature leaves (node positions 8-12 on primary shoots). Measurements included individual leaf area (by image analysis), leaf blade dimensions (length, width), depth and angle of petiolar and lateral sinuses, number and shape of lobes, leaf thickness (fresh weight per unit area), and chlorophyll content (SPAD). Leaf_Development_Disposition showed remarkable variation reflecting the diversity within V. vinifera. Individual leaf area ranged from 85 cm² ('Petit Manseng') to 385 cm² ('Sultanina'), a 4.5-fold range. Leaf dissection varied from nearly entire margins with shallow sinuses ('Sultanina', 'Cinsaut') through moderately 5-lobed ('Cabernet Sauvignon', 'Merlot') to deeply 7-lobed with overlapping sinuses ('Nebbiolo', 'Mourvèdre'). Leaf thickness (specific leaf mass) ranged from 28 g/m² ('Pinot Noir', thin) to 52 g/m² ('Grenache', thick). Correlation analysis revealed that Leaf_Development_Disposition components were partially independent—leaf size showed only moderate correlation with thickness (r = 0.35) and weak correlation with dissection (r = 0.18), indicating separate genetic control. Principal component analysis of leaf characters explained 65% of cultivar variation in the first two components, confirming the discriminatory power of Leaf_Development_Disposition for ampelographic classification. Functional analysis demonstrated that leaf size correlated with individual leaf photosynthetic capacity (r = 0.72) but that area-normalized photosynthetic rate was actually higher in smaller, thicker leaves, illustrating the complex relationships between leaf morphology and function.
A Leaf_Development_Disposition is a Growth_And_Development_Disposition that inheres in a Vitis vinifera plant and is realized through the characteristic patterns of leaf initiation, expansion, morphogenesis, and maturation that determine leaf size, shape, internal structure, and functional properties. This disposition represents the developmental program governing leaf organogenesis, from primordium initiation at the shoot apical meristem through cell division, expansion, and differentiation phases that establish final leaf form and function. Leaf characteristics are of fundamental importance both for vine physiology (leaves are the primary organs for photosynthesis, transpiration, and gas exchange) and for ampelography (leaf morphology provides key characters for cultivar identification and classification). Leaf_Development_Disposition encompasses multiple partially independent characteristics including leaf size (the area of individual leaves), leaf shape and dissection (the outline and degree of lobing), and leaf thickness (mesophyll development and specific leaf mass), each with distinct genetic control and functional implications. Leaf characteristics vary substantially among cultivars, providing the morphological basis for traditional ampelographic description systems (OIV descriptors), while also determining canopy function through effects on light interception, photosynthetic capacity, transpiration rate, and microclimate modification. Environmental factors modulate leaf expression (light intensity, water status, and temperature affect final leaf dimensions and structure), but genotypic rankings for leaf characteristics are relatively stable, confirming strong genetic determination of Leaf_Development_Disposition.
Leaf_Development_Disposition is_a Growth_And_Development_Disposition representing the developmental program for leaf characteristics. Leaf morphology is fundamental for both ampelography (cultivar identification) and vine physiology (canopy function). Child classes address specific characteristics including Leaf_Size_Disposition, Leaf_Dissection_Disposition, and Leaf_Thickness_Disposition. OIV descriptors provide standardized phenotyping framework. Leaf characteristics show both genetic determination (cultivar rankings stable) and environmental plasticity (absolute values vary with conditions).
Blattentwicklungs-Disposition
Blattmorphogenese
Foliare-Entwicklung
foliar development
leaf characteristics
leaf morphogenesis
Galet P (1979) A Practical Ampelography: Grapevine Identification, Cornell University Press Organisation Internationale de la Vigne et du Vin (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Chitwood DH et al. (2014) A quantitative genetic basis for leaf morphology in a set of precisely defined tomato introgression lines, Plant Cell 26:2953-2969
Leaf Development Disposition
Detailed ampelographic documentation of Leaf_Dissection_Disposition at INRAE Vassal characterized leaf shape parameters across 180 cultivars using standardized OIV descriptors and digital morphometric analysis. Measurements included lobe number, petiolar sinus depth and overlap, lateral sinus depth, lobe shape, and the calculation of dissection index (perimeter²/area, higher values indicating greater dissection). Leaf_Dissection_Disposition showed remarkable cultivar variation representing the full range of grapevine leaf forms. 'Sultanina' exhibited minimal dissection with nearly entire leaf outline, very shallow lateral sinuses (depth < 15% of blade length), widely open U-shaped petiolar sinus, and low dissection index of 15.2. 'Cabernet Sauvignon' displayed moderate 5-lobed dissection with defined lateral sinuses (depth 25-30% of blade length), moderately open petiolar sinus with parallel sides, and intermediate dissection index of 22.8. 'Nebbiolo' showed strong 7-lobed dissection with deep lateral sinuses (depth 40-45%), overlapping petiolar sinus with lyre shape, and high dissection index of 35.4. 'Mourvèdre' exhibited extreme dissection with very deep lateral sinuses creating distinct narrow lobes, strongly overlapping petiolar sinus, and dissection index reaching 42.1. The functional significance of dissection was confirmed by boundary layer measurements—highly dissected 'Nebbiolo' leaves showed 25% higher boundary layer conductance than entire-leaved 'Sultanina' at equivalent leaf area, potentially enhancing gas exchange efficiency. Dissection also affected light transmission through the canopy, with highly dissected leaves allowing 18% more direct light penetration through gaps between lobes.
A Leaf_Dissection_Disposition is a Leaf_Development_Disposition that inheres in Vitis vinifera leaves and is realized through the characteristic pattern and depth of leaf blade incision, determining the number, shape, and definition of leaf lobes and the depth and form of the sinuses between them. This disposition represents one of the most important ampelographic characters, as leaf shape and dissection pattern are highly distinctive among cultivars and relatively stable across environments, providing reliable identification criteria. Leaf dissection results from differential growth patterns during early leaf development, where regions destined to become sinuses show reduced cell division and expansion compared to lobe regions, creating the characteristic incised leaf outline. The degree of dissection ranges from nearly entire (unlobed, with shallow sinuses) through 3-lobed, 5-lobed, to deeply 7-lobed with narrow, deeply-incised sinuses that may overlap at the petiolar sinus. Key descriptive characters include the number of lobes (3, 5, or 7), the depth of the petiolar sinus (very open to overlapping), the shape of the petiolar sinus (U-shaped, V-shaped, lyre-shaped), and the depth and shape of lateral sinuses. Leaf_Dissection_Disposition has functional implications beyond ampelography—deeply dissected leaves have higher perimeter-to-area ratios enhancing boundary layer conductance, more complex three-dimensional structure potentially improving light penetration within the canopy, and distinct aerodynamic properties affecting flutter and self-shading.
Leaf_Dissection_Disposition is_a Leaf_Development_Disposition addressing leaf shape and lobing pattern. Dissection is a primary ampelographic character with high discriminatory value for cultivar identification. Key descriptors include lobe number (3, 5, 7), petiolar sinus depth and shape, and lateral sinus depth. Dissection index (perimeter²/area) provides quantitative measure. Functional implications include boundary layer conductance and canopy light penetration. OIV descriptors 068 (shape of blade), 079-083 (sinus characters) provide standardized phenotyping.
Blattlappung
Blattschnitt-Disposition
Blattzähnung
leaf lobing
leaf shape
sinus depth
Galet P (1979) A Practical Ampelography: Grapevine Identification, Cornell University Press Chitwood DH et al. (2014) A quantitative genetic basis for leaf morphology in a set of precisely defined tomato introgression lines, Plant Cell 26:2953-2969 Organisation Internationale de la Vigne et du Vin (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition
Leaf Dissection Disposition
Syrah in Rhône Valley shows leaf emergence from May through July at BBCH 11-38 over 12-14 weeks, producing 15-18 leaves ranging from 80-120 mm dimensions with moderate lobing and sparse trichomes, achieving photosynthetic capacity of 12-16 micromoles CO2 per square meter per second at maturity, under Mediterranean climate conditions of 18-28°C temperature range, limited summer precipitation requiring irrigation, high solar radiation, and moderate fertility typical of this vigorous mid-to-late season cultivar
Leaf emergence represents the fundamental developmental process where leaf primordia initiated at shoot apex undergo coordinated growth transitioning from microscopic multicellular initiation through recognizable leaf form to functional photosynthetic organ, establishing photosynthetic capacity that sustains shoot growth, supports fruit development, and enables reserve accumulation for subsequent seasons. This essential developmental process occurs continuously during shoot growth from early elongation through growth cessation, typically producing 12-20 leaves per shoot over 8-16 week period depending on vigor and conditions. Leaf emergence encompasses primordium initiation at shoot apical meristem, sequential emergence of multiple leaves in regular phyllotactic pattern, expansion from compressed primordia to functional photosynthetic structures, and maturation establishing full photosynthetic capacity and structural characteristics. The process begins with primordium initiation at shoot apex where apical meristem produces new leaf primordia every 2-3 days during active growth in alternating positions approximately 180 degrees around shoot axis, with each primordium arising as localized cell proliferation on meristem flank creating multicellular bulge that will develop into complete leaf structure. Primordium development proceeds through coordinated cell division in marginal meristems at leaf edges and plate meristem throughout blade establishing leaf form, differentiation of adaxial upper and abaxial lower surfaces with distinct cellular characteristics, and initiation of procambial strands that will develop into leaf venation system. Leaf expansion follows primordium formation with dramatic size increase from 1-2 mm primordia to 50-150 mm mature leaves through continued cell division producing new cells particularly at margins and continued cell expansion throughout blade increasing individual cell volumes 50-100 fold. Photosynthetic capacity develops progressively during expansion as chloroplasts differentiate in mesophyll cells, chlorophyll accumulates creating characteristic green coloration, thylakoid membranes organize for light capture, and Calvin cycle enzymes accumulate enabling carbon fixation. Mature leaves achieve photosynthetic rates of 10-20 micromoles CO2 per square meter per second at light saturation, with whole-canopy photosynthesis integrated across leaves at various developmental stages providing assimilates for shoot growth, fruit development, and reserve accumulation. Leaf morphology development establishes characteristic grapevine palmate form with major lobes and sinuses, venation pattern with primary veins from petiole attachment and reticulated secondary and tertiary vein network, trichome distribution varying among cultivars from glabrous to densely tomentose, and stomatal distribution with higher density on abaxial surface enabling gas exchange for photosynthesis and transpiration. Leaf lifespan from emergence to senescence typically spans 12-20 weeks depending on position and environmental conditions, with basal leaves emerging early in season persisting through summer until senescence in autumn, while apical leaves emerging late in season may have abbreviated lifespan before dormancy entry. Carbon balance transitions during leaf development from net carbon consumer during early expansion requiring 10-15 days to reach carbon neutrality where photosynthesis equals respiration, to net carbon producer contributing excess assimilate to shoot growth and other sinks, with mature leaves potentially producing 5-10 times their own carbon requirements enabling support of other organs. Water relationships evolve during emergence with developing leaves showing high water content 80-85% fresh weight and high transpiration rates per unit area relative to mature leaves, declining to 70-75% water content in mature leaves with regulated transpiration through stomatal control responding to environmental conditions and plant water status. Nutrient requirements particularly nitrogen remain high in emerging leaves supporting protein synthesis for both structural and photosynthetic components, with nitrogen concentration 3-5% dry weight in young leaves declining to 2-3% in mature leaves as structural carbohydrates accumulate diluting protein nitrogen. Vascular development establishes connection between leaf and shoot axis with xylem providing water delivery from roots through stem to transpiring leaf surfaces, and phloem initially importing assimilates to support leaf growth then transitioning to export as photosynthetic capacity develops and production exceeds local requirements. Hormonal regulation coordinates leaf emergence with auxin from shoot apex affecting primordium positioning and expansion polarity, cytokinins promoting cell division in primordia and expanding leaves, gibberellins affecting cell expansion and final leaf size, and abscisic acid potentially involved in maturation and stress responses. Temperature strongly affects emergence rate with primordium initiation frequency, expansion rate, and maturation all approximately doubling per 10°C increase between 15-28°C optimal range, influencing total leaf number produced per season and final leaf sizes achieved. Light intensity affects emergence through photosynthetic capacity development with high light promoting rapid chloroplast maturation and maximum photosynthetic capacity, and through morphological effects on leaf size, thickness, and structure with high light producing smaller thicker leaves with higher photosynthetic capacity per unit area compared to shade leaves. Water availability affects emergence rate and final leaf characteristics with adequate moisture supporting rapid expansion to large dimensions and full photosynthetic development, while water stress reduces expansion rate, limits final size, and may reduce photosynthetic capacity affecting canopy productivity. Nutrient availability particularly nitrogen affects emergence rate and photosynthetic capacity development with adequate nutrition supporting rapid leaf production and expansion with high photosynthetic potential, while deficiency reduces emergence rate, limits final sizes, and reduces photosynthetic capacity through protein synthesis limitation. The process shows four distinguishable phases based on developmental progression, advancing from primordium initiation at apex through sequential emergence of multiple leaves, expansion creating functional leaf area, and maturation establishing full photosynthetic capacity and structural characteristics. Cultivar variation appears in total leaf number produced per season, final leaf dimensions ranging from small 50-80 mm to large 150-200 mm, leaf shape including lobe number and sinus depth, trichome density affecting leaf appearance and microclimate, and photosynthetic characteristics affecting productivity. Regional timing shows leaf emergence active throughout shoot growth from May-July in warm climates, May-August in temperate regions, and June-September in cool areas, with individual leaves requiring 2-4 weeks from separation to functional maturity. Viticultural management considerations include ensuring adequate water and nutrient availability during leaf emergence supporting canopy development, managing light environment optimizing leaf photosynthetic capacity without excessive shading, monitoring pest and disease colonization of emerging leaves, implementing canopy management optimizing leaf exposure and air movement, and assessing canopy development adequacy for fruit maturation requirements. Damage to emerging leaves from pests, diseases, hail, or other factors reduces photosynthetic capacity potentially severely limiting productivity. Research applications include analyzing mechanisms coordinating leaf initiation, expansion, and photosynthetic development, quantifying environmental effects on leaf development affecting canopy characteristics and photosynthetic capacity, evaluating genetic variation in leaf morphology and function affecting vineyard productivity, modeling leaf emergence and canopy development for management decision support, and assessing climate change impacts on leaf development patterns across viticultural regions.
Leaf_Emergence represents fundamental organ development process within Shoot_Growth establishing photosynthetic capacity. Occurs coordinately with Shoot_Elongation and Internode_Development but focuses on leaf organ formation and function. Subdivided into Leaf_Primordium_Initiation, Sequential_Leaf_Emergence, Leaf_Primordium_Expansion, and Leaf_Maturation based on developmental progression. Distinguished from broader shoot growth processes by focus on individual leaf development. Critical process determining canopy photosynthetic capacity and productivity.
Blattbildung
Blattentwicklung
Blattontogenese
Laubentwicklung
leaf development
leaf formation
leaf ontogeny
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 235-265 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 152-178 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Leaf Emergence
In Riesling vines in Rheingau, Germany during late October to early December, leaf fall progresses over 6 weeks following harvest, with basal leaf senescence initiating at day length of 10.5 hours in mid-October, chlorophyll degradation producing characteristic yellow coloration by late October, and complete defoliation achieved by early December when mean temperatures decline below 5°C, with 65-75% of leaf nitrogen successfully remobilized to perennial tissues before abscission under typical cool continental autumn conditions.
Leaf fall constitutes the annual phenological process in Vitis vinifera wherein mature leaves are systematically shed from the vine following completion of their photosynthetic function during the post-harvest period, occurring through the coordinated progression of leaf senescence, abscission zone development, and physical detachment that transfers the vine from the active vegetative state to winter dormancy over a period of 4-8 weeks typically spanning late autumn from October to December in northern hemisphere viticulture regions. This process represents the terminal phase of the annual leaf life cycle, with individual leaves typically surviving 120-180 days from emergence in spring through senescence and abscission in autumn, fulfilling their primary functions of photosynthetic carbon assimilation, transpirational water transport, and respiratory gas exchange before programmed disassembly and nutrient recovery precede physical separation from the supporting shoot. The physiological significance of leaf fall extends beyond simple tissue disposal to encompass critical resource conservation through remobilization of nitrogen, phosphorus, potassium, and other nutrients from senescing leaves into perennial woody tissues where storage reserves support winter survival and subsequent spring growth, with efficient nutrient recovery during senescence reducing the vine's dependence on soil nutrient uptake in the following season. The temporal coordination of leaf fall responds to environmental signals including decreasing photoperiod serving as the primary trigger for senescence initiation in most Vitis vinifera cultivars, declining temperatures accelerating chlorophyll degradation and abscission zone activation, and reduced soil moisture potentially hastening senescence under drought conditions that compromise continued leaf function. Cultivar variation in leaf fall timing reflects genetic differences in photoperiod sensitivity and temperature response thresholds, with early-maturing varieties such as Chasselas and Müller-Thurgau typically initiating senescence and leaf fall 2-4 weeks before late-maturing varieties such as Cabernet Sauvignon and Mourvèdre under equivalent environmental conditions. The progression of leaf fall within individual vines follows characteristic spatial patterns, with basal leaves on mature shoot portions senescing and falling first, followed by progressive acropetal advancement of senescence toward shoot tips, and lateral shoot leaves often persisting longer than main shoot leaves due to their more recent emergence and continued metabolic activity. Environmental factors beyond photoperiod and temperature influence leaf fall dynamics, with early autumn frost events potentially triggering rapid synchronous leaf fall through freeze damage bypassing normal senescence processes, extended warm autumns delaying senescence and prolonging the photosynthetic season in some years, and fungal diseases such as downy mildew potentially causing premature defoliation that disrupts normal nutrient remobilization. The completion of leaf fall marks a critical transition in the vine's annual cycle, with fully defoliated vines entering the dormant period characterized by minimal metabolic activity, maximum cold hardiness, and reliance on accumulated carbohydrate reserves that will sustain respiratory demands until photosynthesis resumes with new leaf emergence in the following spring.
Leaf_Fall is parent class under Grapevine_Phenological_Process representing the complete annual leaf shedding process. Contains subclasses Leaf_Senescence, Petiole_Abscission, and Leaf_Detachment. Duration 4-8 weeks in late autumn. Photoperiod-triggered with temperature modulation. Integrates senescence, nutrient remobilization, and physical separation. Critical for nutrient conservation and dormancy transition.
Blattabszission
Blattfall
Entlaubung
Herbst-Blattabwurf
autumn leaf drop
defoliation
leaf abscission
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 pp 85-95 Zapata C et al (2004) Partitioning and mobilization of nitrogen in grapevine. Functional Plant Biology 31: 1141-1152 Lim PO et al (2007) Leaf senescence. Annual Review of Plant Biology 58: 115-136
Leaf Fall
Leaf hydraulic conductance monitoring in Chardonnay through a diurnal cycle reveals the dynamic regulation affecting gas exchange, with predawn Kleaf of 12.5 mmol/m²/s/MPa (fully recovered overnight), midmorning decline to 8.2 mmol/m²/s/MPa (moderate stress developing), midday minimum of 4.5 mmol/m²/s/MPa (substantial limitation from reversible decline and possible minor embolism), and afternoon partial recovery to 6.8 mmol/m²/s/MPa (declining demand allowing recovery), with the Kleaf pattern correlating with stomatal conductance changes and demonstrating the hydraulic coordination between water supply capacity and stomatal regulation.
Leaf hydraulic conductance quality is a hydraulic conductivity quality that represents the capacity of leaves to conduct water from petiole xylem through the venation network and mesophyll tissues to evaporating surfaces, characterizing the terminal component of the soil-plant-atmosphere continuum that often constitutes 30-50% of whole-plant hydraulic resistance and shows dynamic regulation affecting transpiration and photosynthesis. Within the BFO framework, this quality specifically addresses the leaf component of hydraulic architecture, distinguishing the complex leaf water transport pathway from simpler stem xylem flow while recognizing that leaf hydraulic conductance (Kleaf) integrates both the vascular (venation) and non-vascular (mesophyll) pathways that together determine water supply to stomatal guard cells and evaporating surfaces. The ontological significance lies in leaf hydraulic conductance determining the water potential experienced by guard cells that regulate stomatal aperture, where the resistance to water flow within leaves creates a water potential gradient from petiole to evaporating sites that affects stomatal behavior, making Kleaf a key determinant of the coordination between hydraulic supply and stomatal demand. The pathway components of leaf hydraulic conductance include the vein network (xylem conducting system with hierarchical branching from midrib through minor veins), the bundle sheath interface (transition from xylem to mesophyll), and the mesophyll pathway (water movement through or around mesophyll cells to evaporating surfaces), with each component contributing to total leaf resistance and potentially limiting water supply under different conditions. The dynamic regulation of leaf hydraulic conductance involves both reversible changes (aquaporin gating, leaf turgor effects) and irreversible changes (embolism in leaf veins), with Kleaf typically declining through the day as water stress develops and recovering overnight when stress is relieved, creating diurnal patterns that affect gas exchange dynamics. The coordination between leaf hydraulic conductance and stomatal conductance is emerging as a key concept in plant water relations, with evidence suggesting that stomatal closure often tracks declining Kleaf, creating a feedback system that limits transpiration as leaf water supply capacity declines. Understanding leaf hydraulic conductance quality enables assessment of the leaf-level limitation on water supply, interpretation of stomatal responses to water stress, and evaluation of the hydraulic coordination that affects vine gas exchange performance.
Leaf_Hydraulic_Conductance_Quality serves as the parent class for leaf pathway components (Venation, Mesophyll). This quality represents the terminal hydraulic pathway that often constitutes a major fraction of whole-plant resistance. The class supports leaf water relations assessment and stomatal coordination interpretation through Kleaf characterization.
Blatthydraulikleitfähigkeit
Blattwasserversorgung
Kleaf
Laminahydraulik
lamina hydraulic capacity
leaf conductance
leaf water supply
Sack L, Holbrook NM (2006) Leaf hydraulics. Annual Review of Plant Biology 57: 361-381 Scoffoni C et al. (2012) Decline of leaf hydraulic conductance with dehydration: relationship to leaf size and venation architecture. Plant Physiology 158(3): 1342-1352
Leaf Hydraulic Conductance Quality
Seasonal monitoring of leaf layer number quality in Yarra Valley Pinot Noir tracked canopy development from budbreak through harvest at three canopy heights. At 40 days post-budbreak, leaf layer number averaged 0.6 across all heights, indicating developing but still sparse canopy. By flowering, fruit zone LLN reached 1.2 with upper canopy at 1.8, prompting shoot positioning to redistribute density. At véraison, fruit zone leaf layer number quality had increased to 2.4 indicating need for basal leaf removal, which reduced fruit zone LLN to 1.3 within one week while upper canopy remained at 2.1 providing continued photosynthetic capacity. Final pre-harvest assessment showed stable fruit zone LLN of 1.4 with 25% gaps, meeting targets for this Botrytis-susceptible variety in a cool, humid vintage.
Leaf layer number quality is a canopy density quality that inheres in a grapevine canopy system and represents the mean number of leaf contacts encountered when a probe is inserted horizontally through the canopy at specified heights, providing standardized quantification of foliage density that relates directly to light attenuation, air circulation restriction, and spray interception characteristics. This quality serves as the primary quantitative expression of canopy density derived from point quadrat analysis methodology, enabling objective comparison of canopy management treatments, cultivar-training system combinations, and temporal density changes within and across growing seasons. The ontological basis of leaf layer number quality rests on the physical principle that each leaf layer encountered during canopy transect represents an obstacle to light transmission, air movement, and spray penetration, with cumulative effects of multiple layers creating the interior environmental conditions that influence fruit development and disease pressure. The measurement protocol involves systematic insertion of a thin rigid probe at regular intervals along and perpendicular to the vine row, recording each leaf contact until the probe emerges from the opposite canopy side, then computing the mean contacts per insertion across sampling positions. Optimal leaf layer number quality for wine grape production generally targets values between 1.0 and 1.5, representing canopy architecture where most probe insertions encounter one or occasionally two leaves while some pass through gaps without contact, corresponding to approximately 60-80% light interception suitable for grape quality development. Leaf layer numbers below 1.0 indicate sparse canopies with substantial gap frequency that may provide inadequate leaf area for fruit ripening and vine carbohydrate requirements, while values exceeding 2.0 signal excessive density likely to compromise interior fruit zone conditions through heavy shading and restricted ventilation. The relationship between leaf layer number quality and canopy management decisions involves both diagnostic application, identifying density problems requiring intervention, and monitoring application, tracking the effects of management operations on canopy structure over time. Spatial variation in leaf layer number quality across canopy heights provides additional information, with separate assessment at fruit zone height, mid-canopy, and upper canopy levels enabling targeted management addressing specific problematic zones rather than uniform whole-canopy treatment.
Leaf_Layer_Number_Quality is_a Canopy_Density_Quality, representing the quantitative leaf layering dimension of density characterization derived from point quadrat methodology. This class serves as parent for positional subclasses including Fruit_Zone_Leaf_Layers_Quality, External_Leaf_Layer_Quality, and Interior_Leaf_Layer_Quality which address specific canopy regions. Distinguished from Canopy_Porosity_Quality which addresses gaps rather than leaf contacts. The optimal range of 1.0-1.5 provides key quantitative benchmark. Strong relationship with Point_Quadrat_Index_Quality which provides the measurement methodology.
Blattlagenzahl
LLN
Laubschichtung
Laubschichtzahl
canopy leaf layers
leaf layers
mean leaf contacts
mittlere Blattkontakte
Smart RE (1985) Principles of grapevine canopy microclimate manipulation with implications for yield and quality: a review. American Journal of Enology and Viticulture 36:230-239 Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 44-59 Gladstone EA, Dokoozlian NK (2003) Influence of leaf area density and trellis/training system on the light microclimate within grapevine canopies. Vitis 42:123-131
Leaf Layer Number Quality
A leaf lobe representing the upper right lateral lobe on a mature leaf at node 7 of Vitis vinifera cv. Cabernet Sauvignon in Bordeaux, France, one of five total lobes creating characteristic five-lobed palmate architecture, measuring 4.8 cm maximum width perpendicular to primary lateral vein axis and 5.2 cm length from sinus base to lobe apex, separated from adjacent apical lobe by upper right lateral sinus of 2.8 cm depth and from lower lateral lobe by middle right lateral sinus of 2.4 cm depth, supplied by the second primary lateral vein branching from median vein at 58-degree angle and extending through lobe center to near apex, lobe tissue representing approximately 18% of total blade area, exhibiting acute apex with approximately 45-degree angle, convex margins with heterogeneous tooth pattern alternating large and small teeth, moderate lobe dissection creating distinct but not deeply separated lobe appearance characteristic of this cultivar, contributing to overall leaf outline shape and perimeter length, providing mechanical flexibility enabling independent lobe movement in wind responses, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing typical lateral lobe morphology diagnostic for Cabernet Sauvignon identification.
A distinct projecting division or segment of the grapevine leaf blade separated from adjacent lobes by sinuses in Vitis vinifera, characterized by radiating arrangement from the petiolar junction creating characteristic palmate leaf architecture, variable number typically 3-7 lobes per leaf with 5-lobed configuration predominating in most cultivars, distinct vascularization with each lobe supplied by a primary lateral vein and its branching network, cultivar-specific morphological features including lobe number, depth of dissection, width, length, apex characteristics, and margin patterns serving as primary ampelographic diagnostic characters, functional significance providing increased leaf surface area and perimeter compared to simple undivided leaves, mechanical properties affecting blade flexibility and wind response, and developmental origin through differential growth patterns during leaf expansion creating the indentations (sinuses) that define lobe boundaries. Leaf lobes constitute the fundamental organizational units of grapevine leaf architecture, with lobe patterns determined primarily by genetic programming creating cultivar-diagnostic configurations, modified to some extent by environmental conditions during development particularly vigor and resource availability, and categorized by position as apical (terminal), lateral (intermediate), or basal lobes, with additional characterization by depth, width, apex shape, and other morphological features creating the rich diversity of leaf forms used for cultivar identification.
Leaf lobes are distinguished from simple undivided blade regions by separation through sinuses creating distinct projecting divisions. Lobe number shows strong genetic control with cultivar-specific patterns ranging from 3 lobes (weakly dissected cultivars) to 7 lobes (deeply dissected cultivars), though 5 lobes predominates. The relationship between lobe number and primary lateral vein number is direct, with each lobe typically supplied by one primary lateral vein, plus the terminal lobe supplied by the median vein extension. Lobe depth varies from shallow dissection with sinuses extending 10-20% of distance toward median vein, to deep dissection with sinuses reaching 60-80% toward median vein, creating the spectrum from weakly lobed to deeply dissected leaves. Functional implications of lobing include increased surface area and perimeter for given blade area potentially enhancing gas exchange, increased mechanical flexibility allowing lobes to move independently reducing wind stress, and possible effects on boundary layer characteristics affecting heat and gas exchange though experimental support is limited. Developmental control involves marginal meristem activity establishing sinus positions during early expansion, with continued differential growth creating final lobe patterns. Environmental effects are relatively minor compared to genetic control, with vigor and resource availability affecting lobe size but generally not altering fundamental patterns. Ampelographic utility is exceptional, with lobe characteristics serving as primary diagnostic features in most identification systems.
Blattlappen
Blattsegment
Laminaausprägung
blade lobe
lamina lobe
leaf division
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 1-57 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 33-55.
Leaf Lobe
Leaf maintenance respiration assessment in aging Chardonnay canopy reveals the carbon balance shift with leaf senescence, with young fully-expanded leaves (nitrogen 2.8%) showing maintenance respiration of 1.2 μmol CO₂/m²/s and net photosynthesis of 14 μmol CO₂/m²/s (strongly positive balance), while senescing leaves (nitrogen 1.4%, chlorophyll declining) show reduced maintenance of 0.6 μmol CO₂/m²/s but net photosynthesis of only 3 μmol CO₂/m²/s (weakly positive), and heavily senesced leaves approach compensation where maintenance cost equals photosynthetic gain, demonstrating the trajectory toward carbon-neutral and eventually carbon-negative leaf status that justifies senescence-triggered abscission.
Leaf maintenance respiration quality is a maintenance respiration quality that represents the rate of respiratory metabolism required to maintain leaf cellular structures and photosynthetic apparatus without net growth, characterizing the carbon cost of sustaining the protein-rich photosynthetic machinery, replacing degraded chloroplast components, and maintaining membrane integrity in mature, non-expanding leaves. Within the BFO framework, this quality specifically addresses the leaf component of maintenance respiration, distinguishing leaf maintenance costs from root and fruit maintenance while recognizing that leaves have particularly high maintenance requirements due to their elevated protein content (Rubisco alone comprising up to 25% of leaf nitrogen) and the continuous turnover of photosynthetic apparatus components exposed to light-induced damage. The ontological significance lies in leaf maintenance respiration representing a substantial component of whole-plant respiratory cost, where the high nitrogen investment in leaves creates correspondingly high maintenance demand, and where the balance between leaf photosynthetic gain and respiratory cost determines whether leaves contribute positively to vine carbon economy or become carbon-negative when photosynthesis declines below maintenance requirement. The physiological basis for leaf maintenance respiration includes photosynthetic protein turnover (particularly D1 protein and Rubisco, with turnover accelerated by light stress), membrane lipid turnover (especially in thylakoid membranes subject to oxidative damage), chlorophyll recycling (continuous synthesis to replace degraded pigments), and general cellular maintenance shared with other tissue types. The factors affecting leaf maintenance respiration include leaf nitrogen content (strong positive correlation with maintenance requirement), leaf age (declining maintenance as proteins are remobilized during senescence), temperature (exponential increase following Q₁₀ of approximately 2.0-2.5), and light history (high light potentially increasing turnover of damaged components). The quantitative relationship between leaf nitrogen and maintenance respiration enables prediction of maintenance costs from readily measured nitrogen content, with maintenance respiration approximately 0.015-0.025 μmol CO₂/m²/s per % leaf nitrogen at 25°C. Understanding leaf maintenance respiration quality enables assessment of the carbon cost of maintaining photosynthetic capacity, prediction of leaf carbon balance decline with age, and optimization of canopy nitrogen distribution to balance photosynthetic gain against maintenance cost.
Leaf_Maintenance_Respiration_Quality addresses the specific maintenance costs of leaf tissue with its protein-rich photosynthetic apparatus. This quality determines the respiratory cost of maintaining photosynthetic capacity and affects the carbon balance calculation for leaves. The class supports canopy carbon budget assessment through characterization of the leaf-specific maintenance component.
Blatt-Erhaltungsatmung
Blatt-Rm
Photosyntheseapparat-Erhaltung
foliar maintenance cost
foliare Instandhaltungskosten
leaf Rm
leaf basal respiration
photosynthetic apparatus maintenance
Reich PB et al. (1998) Leaf structure (specific leaf area) modulates photosynthesis–nitrogen relations: evidence from within and across species and functional groups. Functional Ecology 12(6): 948-958 Ryan MG (1991) Effects of climate change on plant respiration. Ecological Applications 1(2): 157-167
Leaf Maintenance Respiration Quality
The leaf margin description of Gewürztraminer documents medium-length teeth with a distinctive convex-convex shape (OIV 076:3, both sides of each tooth convexly curved outward) creating the rounded, scalloped margin that is characteristic of the Traminer cultivar group — this tooth form contrasts markedly with the rectilinear-rectilinear teeth of Riesling (OIV 076:2, both sides straight, creating angular, saw-blade-like serration), the convex-concave teeth of Syrah (OIV 076:4, one side convex and one side concave, creating asymmetric pointed teeth), and the mixed irregular dentition of Sangiovese (where primary teeth bear secondary teeth on their flanks, producing a compound serration pattern not captured by the simple OIV classification). The margin description was scored on the terminal lobe and upper lateral lobes of 10 representative mature leaves, following the OIV protocol that specifies observation on the distal third of each lobe where tooth form is most consistently expressed.
A mature leaf description component that documents the morphological characteristics of the leaf blade margin — the serrated edge formed by the teeth (denticulations) that border the entire perimeter of the leaf blade including the lobe tips and sinus edges — recording tooth shape, size, regularity, and secondary serration patterns according to OIV descriptors 076-077 that provide cultivar-diagnostic characters based on the geometry of marginal growth patterning.
Leaf_Margin_Description addresses the serration pattern that borders every lobe and sinus of the grapevine leaf, providing a continuous character visible along the entire blade perimeter. The teeth (also termed denticulations or serrations) are formed by localized growth projections at the leaf margin, each tooth representing a marginal vein terminus surrounded by a wedge of blade tissue — the shape of each tooth is determined by the relative growth rates of the two sides (creating symmetric versus asymmetric teeth) and the curvature of each side (creating convex, rectilinear, or concave tooth flanks): these growth parameters are genetically determined and produce cultivar-specific tooth geometries that are among the most reliable leaf characters for identification. The OIV classification of tooth shape into five categories (convex-convex, rectilinear-rectilinear, convex-rectilinear, concave-convex, and mixed) captures the major variation but does not fully describe the diversity of dental morphology — some cultivars exhibit compound serration (primary teeth bearing secondary teeth), others show distinctly mucronate teeth (with a projecting point at the tooth apex), and the regularity versus irregularity of tooth size along the margin provides additional diagnostic information not encoded in the standard OIV categories.
Blattrand-Beschreibung
Blattrand-Morphologie
Blattzähnung-Charakterisierung
blade margin characterization
dental morphology record
leaf serration description
Leaf Margin Description
Merlot in Bordeaux shows leaf maturation following expansion completion across May-August period at BBCH 15-38, with individual leaves requiring 1-2 weeks to achieve full photosynthetic capacity of 11-15 micromoles CO2 per square meter per second and complete structural development including stomatal control system and vascular function, under oceanic climate conditions of 18-26°C temperature range, adequate moisture, and moderate fertility typical of this moderately vigorous mid-season cultivar producing mature canopy supporting fruit development through véraison and ripening
Leaf maturation represents the developmental transition where expanded leaves complete structural and functional development achieving maximum photosynthetic capacity, fully differentiated anatomy, and stable characteristics that will persist through leaf lifespan until senescence, establishing mature canopy capable of supporting fruit development and reserve accumulation. This essential developmental process follows expansion completion, typically requiring 1-2 weeks per leaf from expansion completion to full functional maturity, with leaves at various maturation stages creating developmental gradient along shoot from immature recently expanded apical leaves to fully mature basal leaves. Maturation encompasses completion of chloroplast development achieving maximum photosynthetic enzyme concentrations and membrane organization, final cell wall thickening establishing mechanical properties, stomatal regulation system maturation enabling responsive control of gas exchange, vascular tissue completion establishing full transport capacity, and metabolic adjustment transitioning from growth to maintenance and production. Chloroplast maturation represents critical aspect with thylakoid membranes achieving final organization into grana stacks for light capture and stroma lamellae for electron transport, photosystem complexes reaching maximum concentration and functional efficiency, and Calvin cycle enzymes including Rubisco accumulating to final concentrations of 3-5 mg per square cm leaf area supporting maximum carbon fixation rates. Photosynthetic capacity increases during maturation from 60-80% maximum rates in recently expanded leaves to full capacity of 10-20 micromoles CO2 per square meter per second at light saturation in mature leaves, with light response curves showing maximum quantum efficiency, high light saturation points 1200-1800 micromoles per square meter per second, and minimal photo-inhibition at high light intensities. Stomatal control system matures during this phase with guard cells achieving full responsiveness to environmental signals including light intensity, atmospheric vapor pressure deficit, internal CO2 concentration, and plant water status through abscisic acid signaling, enabling dynamic regulation of gas exchange optimizing carbon gain while controlling water loss. Cell walls complete thickening during maturation with increased cellulose deposition providing tensile strength, lignification in vascular tissues establishing rigid transport structures, and cuticle development on epidermal surfaces providing protection against water loss and pathogen entry, determining final leaf mechanical properties and stress tolerance. Leaf water relations achieve mature characteristics with water potential typically -0.8 to -1.5 MPa under well-watered conditions, pressure-volume relationships reflecting cell wall elasticity and osmotic adjustment capacity, and hydraulic conductance from petiole through blade to stomata determining transpiration capacity and drought sensitivity. Vascular tissues complete functional development with xylem vessels achieving full water transport capacity supporting maximum transpiration rates and phloem sieve elements enabling efficient assimilate export from photosynthesizing mesophyll to growth sinks and storage tissues, with vascular density and architecture affecting resource transport efficiency. Metabolic transition during maturation involves declining respiration from growth levels to maintenance rates 2-4 fold above dark-adapted values, shifting protein synthesis from growth-related to photosynthetic and maintenance proteins, and adjusting carbon metabolism toward maximum net carbon export supporting other organs. Nitrogen concentration declines during maturation from 3-5% dry weight in young leaves to 2-3% in mature leaves as structural carbohydrates accumulate diluting protein nitrogen, with nitrogen distribution showing highest concentrations in photosynthetic apparatus particularly Rubisco in chloroplasts. Carbon balance achieves maximum positive status in mature leaves with net photosynthetic rate typically 8-15 micromoles CO2 per square meter per second during mid-day optimal conditions, supporting leaf's own maintenance requirements consuming approximately 10-20% photosynthetic production while exporting 80-90% to support other organs. Water use efficiency develops mature characteristics with photosynthetic rate per unit transpiration typically 2-4 micromoles CO2 per millimole H2O in mature leaves under favorable conditions, reflecting optimization of stomatal control balancing carbon gain and water loss. Mature leaf lifespan following maturation typically spans 8-16 weeks depending on leaf position and environmental conditions, with basal leaves persisting longer having more extended productive period while apical leaves maturing late in season may have abbreviated lifespan before senescence or dormancy entry. Environmental stress responses become established during maturation with mature leaves showing capacity for photosynthetic down-regulation under water stress protecting photosystems while reducing carbon fixation, osmotic adjustment maintaining turgor under moderate drought, and possible photo-protective mechanisms including xanthophyll cycle operation preventing photo-damage under stress conditions. Secondary metabolite accumulation may occur during maturation particularly in cultivars with anthocyanin pigmentation developing red coloration in leaves, phenolic compounds accumulating for defense against herbivory and disease, and volatile organic compounds potentially serving signaling or defense functions. Hormonal status transitions during maturation with declining growth-promoting hormones and possible increasing abscisic acid establishing stress-responsive state, with cytokinin concentrations particularly important for maintaining photosynthetic function and delaying senescence. Temperature affects maturation rate with warm conditions promoting rapid completion while cool conditions extend maturation period, though final photosynthetic capacity may be higher in leaves maturing under moderate rather than high temperatures through optimization of photosynthetic apparatus to prevailing conditions. Light intensity during maturation affects final photosynthetic characteristics with high light promoting maximum photosynthetic capacity per unit area and thick leaf structure, while shade reduces capacity though may increase chlorophyll concentration per unit area as adaptation to light limitation. Water availability during maturation affects final leaf characteristics with adequate moisture supporting normal maturation and maximum function while stress during maturation may result in reduced photosynthetic capacity and increased stress sensitivity. Nutrient availability particularly nitrogen during maturation affects final photosynthetic capacity with adequate nitrogen supporting maximum enzyme concentrations and high photosynthetic rates while deficiency reduces capacity limiting leaf productivity. The process shows variation in timing and final characteristics among leaves with basal leaves maturing earliest and potentially achieving highest photosynthetic capacity having developed under favorable early-to-mid season conditions, middle leaves maturing during peak season achieving maximum function, and apical leaves maturing latest potentially showing reduced capacity if environmental conditions deteriorate late season. Cultivar variation appears in maximum photosynthetic capacity in mature leaves ranging from moderate 8-12 micromoles per square meter per second in some varieties to high 15-20 micromoles per square meter per second in others, drought tolerance characteristics, and photosynthetic temperature optimum affecting performance in different climate zones. Regional timing shows leaf maturation occurring throughout growing season offset from expansion by 1-2 weeks, with basal leaves maturing June-July, middle leaves July-August, and apical leaves August-September in temperate regions. Viticultural management considerations include recognizing that mature leaf photosynthetic capacity determines canopy productivity supporting fruit development and reserve accumulation, maintaining leaf health during functional period maximizing productivity through pest and disease management, providing adequate water and nutrients supporting maximum photosynthetic function, and assessing mature canopy characteristics evaluating vineyard productivity potential. Factors limiting leaf maturation or reducing mature leaf function significantly impact whole-vine productivity and fruit quality. Research applications include quantifying photosynthetic capacity development and its environmental and genetic regulation, analyzing mechanisms of photosynthetic optimization to prevailing growth conditions, evaluating stress response capabilities in mature leaves affecting drought tolerance, modeling leaf carbon balance and its contribution to whole-plant productivity, and assessing implications of leaf functional characteristics for vineyard productivity across diverse environmental conditions and management systems.
Leaf_Maturation is fourth and final Leaf_Emergence subphase, distinguished by completion of structural and functional development achieving maximum photosynthetic capacity. Follows Leaf_Primordium_Expansion. Establishes productive mature canopy supporting vine growth and reproduction. Timing and completeness critically affect photosynthetic capacity and productivity. Mature characteristics persist until senescence.
Blattausreifung
Blattreife
ausgereiftes Laub
fully developed leaf
functional leaf development
funktionelles Blatt
leaf maturity
mature foliage
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 262-282 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 175-195 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Leaf Maturation
Cabernet Franc in Loire Valley shows leaf primordium expansion from late May through August at BBCH 15-38, with individual leaves requiring 2-3 weeks to expand from 3-5 mm separated primordia to final dimensions of 85-140 mm depending on position, achieving expansion rates of 6-10 mm per day during peak growth under temperate maritime conditions of 17-25°C temperature range, adequate moisture, high light availability, and moderate fertility typical of this vigorous mid-season cultivar
Leaf primordium expansion represents the dramatic growth phase where compressed primordia increase from millimeter scale to full leaf dimensions through coordinated cell division and expansion, establishing functional photosynthetic area and characteristic palmate grapevine leaf morphology. This critical developmental process follows primordium initiation and separation from axis, typically requiring 2-4 weeks per leaf from expansion initiation to near-final dimensions, with multiple leaves at various expansion stages creating developmental gradient along shoot. Expansion proceeds through continued cell division in marginal and plate meristems producing new cells particularly at leaf edges and throughout blade interior, and progressive cell expansion throughout developing blade where cells increase from meristematic dimensions to final sizes representing 50-200 fold volume increase. Cell division during expansion shows highest activity in marginal meristems at leaf edges producing cells that will form blade extent and lobe structures, with division continuing for 7-14 days in marginal zones after minimal division in interior regions has ceased, establishing size potential through cell number production. Cell expansion follows and overlaps with division phase with cells throughout blade undergoing water uptake creating turgor pressure driving cell wall extension, increasing individual cell dimensions from 10-20 micrometers in meristematic zones to 30-80 micrometers in mature blade representing 10-40 fold linear increase and 1000-64000 fold volume increase. Leaf area increase shows characteristic rapid exponential phase during early expansion when both cell production and expansion operate at maximum rates, transitioning to declining rate as cell division ceases and expansion completes, with total expansion from 2-4 mm separated primordium to 50-150 mm mature leaf representing 625-5600 fold area increase. Expansion rate varies substantially among leaves along shoot reflecting resource availability and growth conditions, with basal leaves showing modest rates of 3-6 mm per day during early slow shoot growth achieving final dimensions of 50-90 mm, middle leaves showing maximum rates of 8-15 mm per day during peak growth reaching 100-150 mm, and apical leaves showing declining rates of 4-8 mm per day achieving smaller 60-100 mm final sizes. Morphological development during expansion establishes characteristic palmate leaf form with major lobes separated by sinuses, with lobe number typically 3-5 and sinus depth varying from shallow creating nearly entire leaves to deep producing highly dissected forms depending on cultivar genetics and development conditions. Venation pattern develops during expansion with primary veins extending from petiole attachment point into major lobes providing structural framework and main transport pathways, secondary veins branching from primaries creating reticulated network, and tertiary veins forming fine network ensuring all photosynthetic cells have vascular access. Trichome development occurs during expansion with hair-like structures emerging from epidermal cells, with trichome density, length, and branching characteristics varying substantially among cultivars from glabrous leaves with minimal trichomes to densely tomentose leaves with extensive covering affecting leaf optical properties and transpiration characteristics. Stomatal development completes during expansion with stomatal complexes differentiating from epidermal progenitor cells, achieving final density typically 150-250 per square mm on abaxial lower surface and 50-100 per square mm on adaxial upper surface, with stomata becoming functional during late expansion enabling gas exchange for photosynthesis and transpiration. Chloroplast development proceeds during expansion transforming proplastids in young cells to functional chloroplasts with organized thylakoid membranes for light capture and assembled Calvin cycle enzymes for carbon fixation, with chlorophyll accumulation creating progressive greening from pale yellow-green in young expanding tissues to dark green in nearly mature leaves. Photosynthetic capacity develops progressively during expansion with photosynthetic rate per unit area increasing from negligible in early expansion to 40-60% mature rates when leaf reaches 50% final area to 80-90% mature rates approaching final dimensions, with whole-leaf carbon balance transitioning from net consumption to net production typically when leaf achieves 70-80% final area. Cell wall composition changes during expansion from thin primary walls dominated by pectin allowing plastic deformation to progressively thickening walls with increased cellulose and hemicellulose providing structural rigidity as expansion completes, determining final leaf mechanical properties and resistance to wilting under water stress. Metabolic activity remains elevated during expansion with respiration rates 8-15 fold above mature leaf levels providing energy for growth processes including cell division, expansion, wall synthesis, and protein production, declining progressively as expansion completes and maintenance requirements dominate. Protein synthesis operates at maximum capacity during expansion producing structural proteins for cell walls and membranes, photosynthetic proteins accumulating in chloroplasts with Rubisco reaching 40-50% total leaf protein supporting carbon fixation, and regulatory proteins coordinating complex developmental processes. Hormonal regulation coordinating expansion involves cytokinins promoting cell division in marginal and plate meristems, gibberellins supporting cell expansion and affecting final leaf size with higher concentrations promoting larger leaves, auxin from developing veins coordinating expansion patterns, and brassinosteroids enhancing cell expansion and vascular development. Temperature strongly affects expansion rate with development approximately doubling per 10°C increase between 15-28°C optimal range, influencing both cell division duration and expansion rate affecting time required for expansion completion and potentially final dimensions achieved. Light intensity critically affects expansion with high light promoting rapid expansion to moderate final dimensions and thick leaf structure with high photosynthetic capacity per unit area, while shade reduces expansion rate and promotes larger final area with thin structure adapted to light limitation creating high specific leaf area. Water availability critically affects expansion through direct effects on cell turgor driving expansion, with adequate moisture essential for maintaining pressure supporting cell wall extension, while water stress rapidly arrests expansion permanently limiting final dimensions even after stress relief affecting photosynthetic capacity. Nutrient availability particularly nitrogen affects expansion through effects on protein synthesis supporting both cell division and expansion, with adequate nitrogen supporting maximum expansion rate and large final dimensions while deficiency reduces expansion limiting final area and photosynthetic potential. The process shows variation in timing and rate among leaves creating characteristic pattern along shoot with basal leaves requiring longer expansion periods during slow early growth, middle leaves expanding rapidly during favorable peak growth conditions, and apical leaves showing abbreviated rapid expansion before growth cessation. Cultivar variation appears in maximum expansion rate, final leaf dimensions ranging from small 50-80 mm in compact varieties to large 150-200 mm in vigorous types, leaf shape characteristics including lobe number and sinus depth, trichome density affecting leaf appearance, and expansion responsiveness to environmental conditions. Regional timing shows leaf expansion occurring throughout growing season from May-July in warm climates, May-August in temperate regions, and June-September in cool areas, with individual leaves requiring 2-4 weeks for expansion depending on position and conditions. Viticultural management considerations include ensuring adequate water availability during expansion supporting full development of functional photosynthetic area, providing sufficient nitrogen supporting protein synthesis without excessive promotion, managing light environment optimizing expansion and photosynthetic development, monitoring for pest and disease damage during expansion as young tissues are particularly vulnerable, and assessing expansion patterns across vineyard evaluating vine health and resource availability. Limitations during expansion permanently reduce final leaf size affecting photosynthetic capacity and productivity. Research applications include analyzing cellular and molecular mechanisms of cell division and expansion during organ development, quantifying environmental and hormonal effects on expansion affecting final leaf characteristics, evaluating genetic variation in expansion patterns affecting leaf morphology and function, modeling expansion processes for predicting canopy development under varying conditions, and assessing developmental biology of compound leaf form using grapevine as model system.
Leaf_Primordium_Expansion is third of four Leaf_Emergence subphases, distinguished by dramatic size increase from millimeter primordia to functional photosynthetic structures. Follows Sequential_Leaf_Emergence separation phase. Transitions to Leaf_Maturation as expansion completes and full photosynthetic capacity develops. Primary determinant of final leaf size and photosynthetic area. Highly responsive to environmental conditions during expansion period.
Blattspreitung
Blattvergrößerung
blade expansion
leaf enlargement
leaf expansion
leaf growth
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 255-265 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 172-178 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Leaf Primordium Expansion
Pinot Noir in Oregon shows leaf primordium initiation from May through July at BBCH 11-35 over 10-12 weeks, initiating new primordia every 2-3 days during June peak growth and every 3-4 days during July decline, producing total of 14-17 primordia establishing maximum leaf number potential under cool climate conditions of 16-24°C temperature range, adequate moisture, and moderate fertility typical of this moderate-vigor early-to-mid season cultivar
Leaf primordium initiation represents the fundamental organogenesis process where apical meristem produces new leaf primordia through localized cell proliferation, establishing foundational structures that will develop into functional photosynthetic organs and determining maximum leaf number potential for developing shoot. This essential developmental process occurs continuously at shoot apex during active growth from early shoot elongation through growth cessation, with new primordia initiating every 2-3 days during peak growth producing total of 12-20 primordia per shoot over 8-16 week period depending on cultivar vigor and environmental conditions. Primordium initiation begins with localized increase in cell division activity on apical meristem flank at position determined by phyllotactic patterning, with cells in initiation zone undergoing accelerated divisions creating multicellular bulge protruding from meristem surface marking first visible stage of leaf formation. The initiation site occurs in regular pattern around shoot axis typically alternating approximately 180 degrees between successive primordia creating characteristic distichous leaf arrangement of grapevine, with positioning controlled by auxin concentration patterns and mechanical forces in meristem dome influencing where new primordia can form. Primordial cells undergo oriented divisions establishing three-dimensional structure with adaxial-abaxial polarity defining upper and lower leaf surfaces, proximal-distal axis defining petiole-blade orientation, and medio-lateral axis defining central midrib and lateral lobes, with these fundamental polarities established early in initiation through gene expression patterns and hormonal gradients. Cell division patterns during initiation show activity concentrated in peripheral zones that will form marginal meristems responsible for blade expansion, and in central zones that will form plate meristem contributing cells throughout blade interior, establishing organizational framework that will guide subsequent leaf development. Primordium size at initiation measures approximately 50-100 micrometers diameter containing 50-200 cells, increasing to 500-1000 micrometers with several thousand cells by time primordium is clearly recognizable as leaf structure rather than simple protrusion, requiring approximately 3-5 days from first division acceleration to recognizable leaf form. Procambial strand initiation occurs early in primordium development establishing vascular connection between developing leaf and shoot axis, with procambial cells forming continuous strand from primordium base through shoot cortex connecting to main vascular system, providing pathway for water delivery and eventually assimilate export critical for leaf development and function. Gene expression patterns during initiation include activation of leaf identity genes specifying organ type and suppressing alternative developmental fates like shoot apex or floral meristem, polarity genes establishing adaxial-abaxial identity and coordinating with hormonal signals, and morphogenesis genes controlling cell division patterns and growth orientation establishing three-dimensional leaf form. Marginal meristem establishment represents critical initiation event with cells at primordium edges maintaining division capacity longer than internal cells, creating zones that will produce majority of blade area through sustained cell production at margins extending into lobes characteristic of palmate grapevine leaf form. Early primordium development shows differentiation between petiole and blade regions with cells near attachment point developing into petiole with vascular strand and supportive tissue, while distal cells form blade with characteristic morphology including lobes, sinuses, and expanded photosynthetic surface. Hormonal regulation of initiation involves auxin concentration patterns at meristem apex with local auxin maxima marking sites where primordia will initiate and auxin transport directing polarity establishment, cytokinins from roots promoting cell division supporting primordium growth, and brassinosteroids potentially affecting primordium outgrowth and expansion. Timing between successive primordium initiations reflects shoot growth rate and apical meristem activity, with most rapid initiation of 2-day intervals during peak exponential growth when apical meristem operates at maximum capacity, extending to 3-4 day intervals during declining growth as meristem activity reduces, and ceasing when apical growth terminates. Temperature affects initiation frequency through effects on meristem cell division rates, with optimal 22-28°C supporting maximum initiation frequency approximately doubling per 10°C increase within physiological range, influencing total primordium number produced per season determining maximum potential leaf number. Light intensity and quality affect apical meristem activity influencing initiation through effects on photosynthetic production providing resources for growth and possible photomorphogenic responses affecting meristem developmental state, though initiation occurs across wide light range unlike expansion which shows strong light dependency. Water availability affects initiation through effects on cell division requiring adequate turgor and metabolic function, with severe water stress potentially reducing initiation frequency though moderate stress has minimal effect as meristem maintains preferential resource allocation. Nutrient availability particularly nitrogen affects initiation through effects on protein synthesis supporting cell division and growth, with adequate nutrition supporting maximum initiation frequency while severe deficiency may reduce frequency limiting final leaf number potential. The process continues throughout shoot growth showing characteristic temporal pattern of increasing initiation frequency during accelerating growth, maximum frequency during peak growth producing 2-3 primordia per week, and declining frequency during decelerating growth before cessation when apical meristem activity terminates. Cultivar variation appears in total primordium number initiated per season reflecting differences in growth duration and apical meristem activity patterns, with vigorous long-season cultivars initiating 18-22 primordia producing potential for high leaf numbers while less vigorous or shorter-season types initiate 10-14 primordia limiting leaf number. Regional timing shows primordium initiation active from May through July-August in warm climates, May through August-September in temperate regions, and June through September in cool areas, corresponding to shoot growth period with initiation ceasing when growth terminates. Viticultural significance includes establishing maximum leaf number potential for developing shoot with actual emerged leaf number determined by primordium number initiated at apex, recognizing that environmental conditions during initiation period affect total leaf number affecting ultimate canopy size and photosynthetic capacity, and understanding that management affecting shoot growth including irrigation and nutrition influences initiation patterns determining canopy potential. Research applications include analyzing molecular mechanisms of organogenesis including gene regulatory networks controlling leaf identity and patterning, evaluating hormonal regulation of initiation timing and positioning including auxin transport and signaling, quantifying environmental effects on initiation frequency affecting final leaf number, investigating cellular mechanisms of polarity establishment and morphogenesis, and assessing evolutionary and developmental biology of leaf form in vascular plants using grapevine as model for compound leaf development.
Leaf_Primordium_Initiation is first of four Leaf_Emergence subphases, distinguished by organogenesis at apical meristem establishing new leaf structures. Represents earliest stage of leaf development occurring at microscopic scale at shoot apex. Transitions to Sequential_Leaf_Emergence as primordia continue developing and separating from axis. Determines maximum leaf number potential for shoot. Critical developmental process establishing canopy capacity.
Blattanlagebildung
Blattinitiierung
Blattorganogenese
Blattprimordienbildung
foliar primordium development
leaf initiation
leaf organogenesis
primordium formation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 235-242 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 152-162 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Leaf Primordium Initiation
Leaf removal response quality evaluation in Stellenbosch Chenin Blanc compared vine physiological and fruit composition responses across defoliation intensities removing 0%, 33%, 66%, and 100% of leaves in the basal six nodes at véraison. The 33% treatment showed minimal leaf removal response quality challenge, with remaining leaves increasing photosynthetic rate by 12% and fruit achieving equivalent Brix and yield to controls with improved exposure benefits. The 66% treatment exhibited moderate stress with 28% photosynthetic upregulation insufficient to fully compensate, resulting in 0.8°Brix reduction and 15% lower pruning weight indicating reserve depletion. The 100% basal defoliation treatment demonstrated severe leaf removal response quality limitation, with dramatic 45% photosynthetic upregulation in remaining upper leaves still inadequate to support normal ripening, resulting in 2.1°Brix reduction, incomplete flavor development, and 35% lower pruning weight indicating significant carbohydrate deficit that reduced bud fruitfulness and yield in the subsequent season by 22%.
Leaf removal response quality is a viticultural quality that inheres in a grapevine organism and represents the characteristic manner and magnitude of physiological adjustment exhibited following deliberate removal of functional leaves from the canopy, encompassing both the immediate effects on vine carbon balance, water relations, and fruit development as well as the subsequent compensatory responses including upregulation of photosynthesis in remaining leaves and potential refoliation from latent or lateral buds. This quality captures the capacity of grapevines to tolerate and adapt to defoliation events that occur through management operations including basal leaf removal for fruit exposure and disease control, as well as through natural or accidental causes including hail damage, pest feeding, and disease-induced leaf loss. The ontological significance of leaf removal response quality lies in its determination of the threshold beyond which leaf removal compromises vine function rather than providing net benefit, recognizing that moderate defoliation may improve fruit quality through enhanced exposure while severe defoliation can impair ripening through source limitation and cause carbohydrate depletion affecting subsequent season productivity. The physiological basis of leaf removal response involves compensatory mechanisms at multiple organizational levels, including upregulation of photosynthetic rate in remaining leaves through increased stomatal conductance and Rubisco activity, reallocation of resources from vegetative growth to fruit development, mobilization of carbohydrate reserves from woody tissues to support ripening when current photosynthate proves insufficient, and activation of dormant buds to produce replacement leaf area when defoliation is severe. The magnitude and effectiveness of leaf removal response vary with vine physiological status at the time of defoliation, the severity and spatial distribution of leaf loss, the timing relative to phenological development, and cultivar-specific characteristics governing compensatory capacity. Early-season defoliation during active growth typically provokes stronger compensatory responses through vigorous refoliation than late-season defoliation when growth potential has diminished, while well-nourished vines with adequate carbohydrate reserves tolerate greater defoliation severity than stressed vines operating at the margins of carbon balance. Understanding cultivar-specific and site-specific leaf removal response quality enables optimization of defoliation timing and intensity to achieve fruit quality benefits while avoiding the threshold where source limitation compromises ripening or reserve storage.
Leaf_Removal_Response_Quality serves as the parent class for Defoliation_Tolerance_Quality and Refoliation_Capacity_Quality which address specific dimensions of defoliation response. This class captures the integrated vine response to leaf loss including both tolerance mechanisms maintaining function with reduced leaf area and compensatory mechanisms restoring leaf area through new growth. Distinguished from Hedging_Response_Quality and related vigor manipulation classes by the focus on leaf tissue removal rather than shoot tip removal. Critical for optimizing basal leaf removal practices and understanding vine resilience to defoliation events.
Blattentfernungstoleranz
Defoliationsantwort
Entblätterungsreaktion
Entlaubungsanpassung
canopy removal tolerance
defoliation response
leaf loss adaptation
leaf stripping response
Poni S et al. (2006) Effects of early defoliation on shoot photosynthesis, yield components, and grape composition. American Journal of Enology and Viticulture 57:397-407 Intrieri C et al. (2008) Early defoliation (hand vs mechanical) for improved crop control and grape composition in Sangiovese (Vitis vinifera L.). Australian Journal of Grape and Wine Research 14:25-32 Tardaguila J et al. (2010) Impact of early leaf removal on yield and fruit and wine composition of Vitis vinifera L. Graciano and Carignan. American Journal of Enology and Viticulture 61:372-381
Leaf Removal Response Quality
The leaf removal specification for Sauvignon Blanc in Marlborough prescribes early mechanical leaf removal at E-L 17-19 (pre-bloom) on the morning-sun (east) side of the canopy using a pneumatic pulse-air defoliator, removing 5-6 basal leaves per shoot in a 30-40 cm band from the fruiting wire to the first foliage wire — early timing is specified because pre-bloom leaf removal reduces fruit set by 10-15% (through reduced carbohydrate supply to inflorescences), producing looser clusters that are less susceptible to Botrytis bunch rot, while also acclimating developing berries to direct light exposure from earliest stages, reducing the sunburn susceptibility that occurs when berries are suddenly exposed at véraison after developing in deep shade.
A canopy management description that documents the selective removal of leaves from the fruit zone to modify cluster-zone microclimate, specifying the timing (phenological stage), extent (number of leaves removed per shoot), canopy side (sun-exposed versus shaded), method (manual or mechanical), and target light and airflow conditions in the fruit zone that optimize the balance between disease suppression, fruit composition enhancement, and sunburn risk management.
Leaf_Removal_Specification addresses one of the most extensively researched canopy management interventions, with the timing, severity, and canopy side of leaf removal each generating distinct and well-documented effects on fruit composition and disease incidence. The fundamental trade-off is between increased light and temperature in the fruit zone (beneficial for phenolic development, methoxypyrazine reduction, and disease suppression) and the risk of excessive berry temperature (>35°C) causing sunburn damage and heat-induced flavor degradation: this trade-off is managed through strategic choice of canopy side (east/morning-sun side preferred in warm climates to provide afternoon shade protection, west/afternoon-sun side acceptable in cool climates where heat is limiting) and timing (pre-bloom removal producing acclimated berries versus post-véraison removal carrying higher sunburn risk). Research has consistently demonstrated that fruit zone leaf removal reduces Botrytis cinerea incidence by 30-70% through improved air circulation and UV-B exposure that suppresses fungal sporulation, while simultaneously increasing anthocyanin, tannin, and flavor precursor concentrations by 10-40% in the exposed berries.
Entblätterungs-Spezifikation
Laubausdünnung-Protokoll
Traubenzone-Entlaubung
basal leaf removal protocol
defoliation specification
fruit zone leaf pulling protocol
Poni S et al. (2006) Effects of early defoliation on shoot photosynthesis, yield components, and grape composition, American Journal of Enology and Viticulture 57: 397-407 Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles pp: 91-120 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 365-375 Intrieri C et al. (2008) Early defoliation (hand vs mechanical) for improved crop control and grape composition in Sangiovese, Australian Journal of Grape and Wine Research 14: 25-32
Leaf Removal Specification
Leaf rolling quality assessment in Languedoc Grenache characterized the cultivar's distinctive rolling response as a primary drought symptom preceding wilting under Mediterranean summer conditions. The variety exhibited pronounced leaf rolling quality with lamina margins curling inward at relatively mild stress levels (stem water potential -0.9 MPa) where other cultivars showed minimal visible symptoms, progressing to complete tubular rolling at moderate stress (-1.1 MPa) before any significant wilting appeared. The rolling response effectively reduced midday transpiration by an estimated 25% compared to flat leaves under equivalent conditions, demonstrating genuine functional value beyond mere symptomatic expression. A rolling severity scale from 0 (flat) through 2 (moderate marginal rolling, 30% area reduction) to 4 (tubular rolling, 60% area reduction) was calibrated against water potential measurements, enabling field assessment of stress severity from this cultivar-characteristic visual indicator. The early appearance of rolling relative to wilting provided management advantage, signaling developing stress before physiological damage thresholds were approached, establishing Grenache leaf rolling quality as a particularly valuable early-warning indicator for deficit irrigation management.
Leaf rolling quality is a drought symptom expression quality that inheres in grapevine leaves and represents the characteristic inward curling of the leaf lamina margins toward the abaxial surface exhibited as a stress-responsive behavior that reduces the transpiring surface area and creates a more humid microenvironment within the rolled leaf structure, functioning as an adaptive mechanism to reduce water loss while simultaneously serving as a visible indicator of water deficit stress. This quality captures both the protective function of leaf rolling in mitigating water loss and its diagnostic value as a visual stress indicator that appears at moderate stress levels before the complete turgor loss manifesting as wilting, providing an earlier warning of developing water deficit than the more pronounced wilting symptom. The ontological significance of leaf rolling quality lies in its dual nature as both an adaptive response providing genuine drought tolerance benefit and a diagnostic symptom enabling stress assessment, with the rolling behavior representing an active response to stress rather than merely a passive consequence of turgor loss as in wilting. The biological basis of leaf rolling involves differential changes in turgor and cell volume between adaxial and abaxial leaf surfaces, combined with the activity of bulliform cells and other specialized structures that facilitate controlled lamina curvature, creating the inward rolling that reduces exposed leaf area and traps a boundary layer of more humid air reducing the vapor pressure gradient driving transpiration. The extent and pattern of leaf rolling vary with stress severity from slight marginal curling at mild stress through pronounced tubular rolling at moderate stress to complete laminar collapse combining rolling and wilting at severe stress levels. Cultivar differences in leaf rolling quality expression reflect anatomical variation in bulliform cell development, lamina flexibility, and the balance between rolling and wilting responses, with some cultivars exhibiting pronounced rolling as a primary stress response while others proceed directly to wilting with minimal rolling phase.
Leaf_Rolling_Quality is_a Drought_Symptom_Expression_Quality, representing the inward curling of leaf margins as a stress response. This class addresses a symptom that appears earlier than wilting and provides adaptive benefit through reduced transpiration. Distinguished from Leaf_Wilting_Quality by the curling mechanism rather than turgor-loss drooping. Cultivar differences in rolling propensity are substantial, with some varieties showing pronounced rolling (e.g., Grenache) while others proceed directly to wilting. The area reduction from rolling (up to 60%) provides quantitative assessment metric.
Blatteinrollen
Blattkräuselung
Blattrollen
drought-induced rolling
lamina rolling
leaf curling
leaf margin curling
trockenheitsbedingte Blattrollung
Kadioglu A, Terzi R (2007) A dehydration avoidance mechanism: leaf rolling. The Botanical Review 73:290-302 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data. Annals of Botany 105:661-676 Flexas J et al. (2002) Effects of drought on photosynthesis in grapevines under field conditions. Functional Plant Biology 29:461-471
Leaf Rolling Quality
Leaf scorch quality evaluation in Columbia Valley Riesling following a 3-day heat event (40-43°C) characterized damage patterns and their implications for canopy function and fruit ripening. Vines on shallow soils with limited water holding capacity exhibited severe leaf scorch quality with 40% of canopy leaf area showing damage, progressing from marginal browning (score 2) on mature leaves to complete lamina necrosis (score 5) on young expanding leaves at shoot tips and senescing basal leaves, with the damage pattern indicating water stress-exacerbated thermal injury consistent with the site's limited soil moisture reservoir. Vines on deep alluvial soils showed mild leaf scorch quality with only 10% canopy damage limited to score 2 marginal browning on exposed exterior leaves, demonstrating the protective value of adequate soil water supply enabling sustained transpirational cooling. Chlorophyll fluorescence measurements on damaged leaves showed Fv/Fm ratios declining from 0.80 on undamaged interior leaves to 0.45 on scorched exterior leaves, quantifying the photosynthetic impairment from leaf scorch. The leaf scorch quality assessment established that vines with greater than 25% canopy damage would likely experience delayed fruit ripening due to reduced photosynthetic capacity, informing harvest timing expectations and quality projections for affected blocks.
Leaf scorch quality is a heat damage severity quality that inheres in grapevine leaves and represents the extent and pattern of heat-induced desiccation and necrosis affecting the leaf lamina, typically progressing from marginal browning at the leaf edges through interveinal necrosis to complete lamina death under severe conditions, with damage patterns reflecting the interaction between thermal exposure intensity, leaf age and position, and the capacity of the vascular system to supply water for transpirational cooling. This quality addresses heat damage to the primary photosynthetic organs with direct implications for canopy function, carbon assimilation capacity, and fruit ripening support during the critical late-season period when heat events most frequently occur. The ontological significance of leaf scorch quality lies in its impact on canopy photosynthetic capacity that may affect fruit ripening when substantial leaf area is damaged, and in its role as a visible indicator of heat stress severity that signals the need for protective management response even when fruit damage may not yet be apparent. The biological basis of leaf scorch encompasses the desiccation of leaf tissue when transpirational water loss exceeds vascular supply capacity under high evaporative demand, combined with direct thermal damage to cells when leaf temperatures exceed tolerance thresholds, with the marginal and interveinal pattern reflecting the vulnerability of tissue distant from the primary vascular supply where water stress develops first under limiting conditions. The progression of leaf scorch from margins inward follows the gradient of water status across the leaf, with marginal tissue experiencing the most negative water potentials and highest temperatures due to reduced transpirational cooling, creating the characteristic scorch pattern diagnostic of heat-induced damage distinct from other causes of leaf browning. Leaf age affects scorch susceptibility, with young expanding leaves and senescing older leaves more vulnerable than mature fully-functional leaves that have completed development but have not begun decline, creating a characteristic distribution of damage concentrated at canopy extremes.
Leaf_Scorch_Quality is_a Heat_Damage_Severity_Quality, representing heat-induced damage specifically to grapevine leaves. This class addresses damage to the primary photosynthetic organs with implications for canopy function. The marginal-to-interveinal progression pattern is diagnostic of heat/water stress interaction. Leaf age vulnerability pattern (young and old > mature) provides additional diagnostic information. Photosynthetic capacity loss (Fv/Fm decline) quantifies functional impact beyond visible damage. Water status strongly modulates scorch severity through transpirational cooling effects. Critical for assessing canopy function and fruit ripening capacity following heat events.
Blattrandnekrose
Blattverbrennung
heat-induced leaf damage
hitzeinduzierte Blattschäden
leaf burn
leaf desiccation
thermal leaf injury
thermische Blattverletzung
Greer DH, Weedon MM (2013) The impact of high temperatures on Vitis vinifera cv. Semillon grapevine performance and berry ripening. Frontiers in Plant Science 4:491 Wahid A et al. (2007) Heat tolerance in plants: an overview. Environmental and Experimental Botany 61:199-223 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 453-507
Leaf Scorch Quality
In Cabernet Sauvignon vines in Napa Valley, California during October-November, leaf senescence progresses over 5 weeks with chlorophyll content declining from 450 to 45 µg/g fresh weight, nitrogen remobilization exporting 2.8 g N per vine to woody tissues representing 68% of leaf nitrogen content, and leaf color transitioning from green through yellow-orange to red-brown as anthocyanins accumulate concurrent with carotenoid exposure under warm autumn conditions averaging 18°C daytime and 8°C nighttime.
Leaf senescence describes the developmental program of controlled cellular dismantling in Vitis vinifera leaves during the post-harvest autumn period, wherein the orderly degradation of chloroplasts, proteins, nucleic acids, and membrane lipids enables recovery and export of valuable nutrients to perennial vine tissues while transforming photosynthetically active green leaves into the characteristic yellow, orange, and red autumn coloration preceding physical abscission over a period of 3-6 weeks. This process represents a highly regulated genetic program rather than passive deterioration, with senescence-associated genes encoding proteases, lipases, nucleases, and chlorophyll-degrading enzymes activated in coordinated sequence to disassemble cellular components while maintaining membrane integrity sufficient for continued export of remobilized nutrients through the vascular system until late senescence stages. The biochemical events underlying leaf senescence begin with chloroplast dismantling as the quantitatively dominant process, with thylakoid membrane disassembly releasing chlorophyll for catabolism through the pheophytinase and pheophorbide oxygenase pathway, stromal proteins including Rubisco undergoing proteolytic degradation to release amino acids for export, and plastid lipids hydrolyzed to provide carbon skeletons for respiration and membrane maintenance during the senescence period. The nutrient remobilization function of leaf senescence enables recovery of 50-80% of leaf nitrogen, 40-60% of phosphorus, and 30-50% of potassium before abscission, with these nutrients transported through phloem connections to woody tissues including trunk, cordons, and roots where they accumulate as storage reserves supporting subsequent spring growth and reducing fertilizer requirements. The regulation of leaf senescence involves integration of developmental, hormonal, and environmental signals, with ethylene and abscisic acid promoting senescence progression, cytokinins delaying senescence through maintenance of chloroplast function, and the declining cytokinin-to-ethylene ratio during autumn creating conditions favoring senescence initiation and progression. Environmental factors modulating leaf senescence include photoperiod providing the primary timing signal through phytochrome-mediated perception of shortening day length, temperature influencing the rate of senescence progression with cool conditions accelerating chlorophyll degradation, and water availability affecting senescence timing with drought stress potentially inducing premature senescence through abscisic acid accumulation. The visible manifestation of leaf senescence appears as progressive color change from green through yellow as chlorophyll degrades revealing underlying carotenoid pigments, with some Vitis vinifera varieties additionally displaying red coloration through anthocyanin synthesis in senescing leaves that may serve photoprotective functions during the extended period of nutrient remobilization.
Leaf_Senescence is parent class under Leaf_Fall representing the controlled cellular dismantling program. Contains subclasses Leaf_Senescence_Initiation, Chlorophyll_Degradation, Nutrient_Remobilization, and Progressive_Leaf_Senescence. Duration 3-6 weeks. Genetically programmed process. Enables 50-80% nitrogen recovery. Photoperiod and temperature regulated. Precedes physical abscission.
Blatt-Alterung
Blattseneszenz
Blattvergilbung
Herbst-Blattseneszenz
autumn leaf senescence
leaf aging
leaf yellowing
programmed leaf deterioration
Lim PO et al (2007) Leaf senescence. Annual Review of Plant Biology 58: 115-136 Zapata C et al (2004) Partitioning and mobilization of nitrogen in grapevine. Functional Plant Biology 31: 1141-1152 Keller M (2020) The Science of Grapevines 3rd Ed pp 85-92
Leaf Senescence
In Pinot Noir vines in Burgundy, France during early October, leaf senescence initiation occurs when photoperiod declines below 11.2 hours, with phytochrome-mediated signaling activating NAC transcription factor expression within 3-5 days, cytokinin-to-ethylene ratio declining from 2.5 to 0.8 over 10 days, and first visible chlorophyll degradation appearing 12-14 days after initiation trigger under cool autumn conditions with mean temperature of 12°C.
Leaf senescence initiation describes the developmental transition in Vitis vinifera leaves from mature photosynthetically active status to the senescence program, wherein environmental perception of shortening photoperiod and declining temperatures triggers signal transduction cascades that activate senescence-associated gene expression, reprogram hormonal balance toward senescence-promoting states, and commit leaf tissues to the irreversible dismantling process that will culminate in nutrient remobilization and eventual abscission over the subsequent 4-6 weeks. This transition involves the integration of multiple environmental signals through photoreceptor systems, with phytochrome-mediated detection of declining red-to-far-red light ratios characteristic of shortening autumn days providing the primary trigger for senescence initiation in most Vitis vinifera cultivars, while temperature-sensing mechanisms involving membrane fluidity changes and cold-responsive transcription factors modulate the timing and rate of senescence entry. The molecular events marking senescence initiation include upregulation of senescence-associated genes encoding transcription factors such as NAC and WRKY family members that orchestrate downstream gene expression changes, activation of hormone biosynthesis and signaling pathways shifting the balance toward senescence-promoting ethylene and abscisic acid, and initial chloroplast-to-nucleus retrograde signaling that coordinates nuclear gene expression with chloroplast dismantling requirements. The hormonal reprogramming during senescence initiation involves declining cytokinin synthesis and sensitivity that removes the primary maintenance signal for photosynthetic function, increasing ethylene production through activation of ACC synthase and ACC oxidase genes, and accumulation of abscisic acid that promotes expression of genes involved in stress response and senescence progression. The timing of senescence initiation varies among cultivars reflecting genetic differences in photoperiod sensitivity, with day-length thresholds for senescence induction ranging from 11-12 hours in early-responding varieties to 9-10 hours in late-responding varieties, creating 2-4 week differences in senescence initiation timing among cultivars grown in the same location. Environmental factors beyond photoperiod influence senescence initiation timing, with unseasonably warm autumn temperatures potentially delaying initiation through maintained cytokinin activity, drought stress inducing premature initiation through abscisic acid accumulation independent of photoperiod signals, and nutrient deficiency particularly nitrogen limitation accelerating senescence initiation through reduced cytokinin synthesis capacity. The commitment point of senescence initiation represents a largely irreversible developmental transition, though early-stage senescence can be partially reversed by cytokinin application or removal of senescence-inducing conditions, with progression beyond initial stages establishing the full senescence program that will proceed through chlorophyll degradation and nutrient remobilization to completion.
Leaf_Senescence_Initiation is subclass of Leaf_Senescence representing the developmental transition to senescence program. Distinguishes from progressive senescence by emphasizing triggering events. Sibling to Chlorophyll_Degradation, Nutrient_Remobilization, and Progressive_Leaf_Senescence. Photoperiod-triggered with temperature modulation. Involves hormonal reprogramming. Largely irreversible commitment.
Blatt-Alterungs-Initiierung
Seneszenz-Auslösung
Seneszenz-Beginn
Seneszenz-Initiierung
leaf aging initiation
senescence induction
senescence onset
senescence triggering
Lim PO et al (2007) Leaf senescence. Annual Review of Plant Biology 58: 115-136 Guo Y and Gan S (2005) Leaf senescence: signals, execution and regulation. Current Topics in Developmental Biology 71: 83-112 Keller M (2020) The Science of Grapevines 3rd Ed pp 86-88
Leaf Senescence Initiation
Gewürztraminer exhibits early leaf senescence timing with visible yellowing 10-14 days post-harvest in Alsace, completing abscission by mid-October. Cinsault maintains photosynthetically active leaves for 45-60 days post-harvest in Mediterranean climates, enabling 15-25% greater starch reserve accumulation compared to early-senescing cultivars. The interval between 10% and 90% leaf yellowing spans 18-28 days in most V. vinifera cultivars, with tight clustering (18-22 days) indicating synchronized senescence.
A senescence disposition that is the inherited capacity of a Vitis vinifera plant to initiate and progress through leaf senescence at a characteristic time relative to harvest and environmental cues, determining the duration of post-harvest photosynthetic activity and nutrient recovery period.
Leaf senescence timing represents a trade-off between extended photosynthesis for reserve accumulation versus early nutrient remobilization and cold acclimation initiation. Early-ripening cultivars typically show later relative senescence (more days post-harvest) than late-ripening cultivars. Photoperiod and temperature interact to trigger senescence, with short days and cool nights accelerating the process independently.
Blattalterungszeitpunkt
Herbstlaubfärbungszeitpunkt
Seneszenz-Timing
autumn coloration timing
leaf drop timing
senescence onset trait
Parker AK et al. (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17:206-216 Bennett J et al. (2005) Leaf senescence and fruit ripening: influence of post-harvest factors. Australian and New Zealand Grapegrower and Winemaker 497:22-26
Leaf Senescence Timing Disposition
The leaf shape description of Riesling documents a pentagonal blade (OIV 067:3) — the five-point outline is clearly defined by five distinct lobes with moderately deep upper lateral sinuses creating the angular contour typical of the pentagonal class, differing from the more rounded circular outline of Chasselas (OIV 067:4), the elongated wedge shape of Bicane (OIV 067:2), and the broad kidney-shaped outline of Rupestris du Lot (OIV 067:5). The pentagonal shape class is the most common in V. vinifera cultivars (encompassing approximately 60% of cultivated varieties), requiring additional characters (lobe number, sinus depth, tooth form) to discriminate within this large shape group.
A mature leaf description component that records the overall outline form of the fully expanded leaf blade according to OIV descriptor 067 (blade shape), scored on a five-point scale from cordiform/heart-shaped (state 1) through wedge-shaped (state 2), pentagonal (state 3), circular/orbicular (state 4) to kidney-shaped/reniform (state 5), providing one of the three highest-priority leaf characters (alongside petiolar sinus shape and number of lobes) for cultivar identification and classification in the OIV system.
Leaf_Shape_Description records the most fundamental geometric character of the grapevine leaf — the overall blade outline that is determined by the developmental patterning of lobe formation, sinus incision depth, and differential growth rates across the blade during leaf expansion. Blade shape is among the OIV Priority 1 descriptors because of its high discriminative power between major cultivar groups, even though within-group discrimination requires additional characters: the five shape categories represent fundamentally different developmental geometries — cordiform leaves result from minimal lobe development with the blade wider near the petiolar insertion, wedge-shaped from elongated growth with reduced lateral expansion, pentagonal from balanced five-lobe development with moderate sinus incision, circular from uniform radial expansion with shallow sinuses, and reniform from lateral expansion exceeding the longitudinal axis. The genetic control of leaf shape in grapevine has been partially characterized through QTL mapping, identifying loci on chromosomes 1, 5, 7, 8, 14, and 17 affecting various aspects of shape including lobe prominence, sinus depth, and blade outline — this polygenic architecture produces the continuous morphological variation that the OIV scoring system discretizes into five categories for practical identification purposes.
Blattform-Beschreibung
Blattgestalt-Charakterisierung
Blattspreiten-Umriss
blade shape characterization
leaf blade outline
leaf form description
Leaf Shape Description
A leaf sinus representing the upper right lateral sinus on a mature leaf at node 7 of Vitis vinifera cv. Syrah in the Rhône Valley, France, positioned between the apical lobe and upper right lateral lobe, measuring 3.2 cm depth from the sinus base (deepest point of indentation) to the imaginary straight line connecting adjacent lobe edges, width of 1.8 cm measured as the straight-line distance between lobe margins at the sinus opening, exhibiting characteristic U-shaped contour with rounded sinus base rather than acute V-shape, sinus base positioned 4.5 cm from the petiolar junction representing approximately 34% of the distance from junction to blade apex indicating moderately deep dissection, sinus margins formed by the converging edges of the apical lobe (left margin) and upper lateral lobe (right margin) showing smooth curvature without teeth in the sinus region itself, sinus creating clear separation between lobes enabling independent lobe movement in wind responses, contributing to overall blade perimeter of 58 cm compared to 42 cm that would occur without lobe dissection, measured at BBCH stage 81 (beginning of berry ripening) in early August, representing typical upper lateral sinus morphology with moderate depth characteristic of this cultivar.
An indentation or gap between adjacent leaf lobes in the grapevine leaf blade in Vitis vinifera, characterized by concave configuration creating the boundaries separating individual lobes, variable depth typically ranging from shallow indentations of 0.5-2 cm to deep dissections of 3-6 cm or more, U-shaped, V-shaped, or intermediate contour profiles, position-specific occurrence as petiolar sinus at blade base, upper lateral sinuses between apical and upper lateral lobes, middle lateral sinuses between upper and lower lateral lobes, and lower lateral sinuses between lower lateral and basal lobes, cultivar-specific characteristics including depth, width, shape, and base configuration serving as critical ampelographic diagnostic features, and functional implications potentially including enhanced blade flexibility allowing independent lobe movement, modification of boundary layer characteristics affecting heat and gas exchange, and contribution to overall blade surface area and perimeter relationships. Leaf sinuses constitute the fundamental dividing features creating lobe architecture, developing through differential growth patterns during leaf expansion with sinus regions showing reduced growth compared to lobe regions, exhibiting strong genetic control creating reliable cultivar-diagnostic patterns, and categorized by position (petiolar, upper lateral, middle lateral, lower lateral) and depth (shallow, medium, deep) creating classification systems central to ampelographic identification.
Leaf sinuses are distinguished from lobe structures as the negative space (indentations) rather than positive projections, with sinuses and lobes representing complementary aspects of palmate leaf architecture. Sinus depth shows continuous variation from very shallow (barely perceptible indentations) to very deep (extending 70-80% of distance toward median vein), with classification into shallow, medium, and deep categories using somewhat arbitrary thresholds typically 20-25% and 50-60% of distance toward median vein. Shape characteristics include U-shaped (rounded base), V-shaped (acute base), and intermediate configurations, with base shape showing diagnostic importance particularly for petiolar sinuses. Position-based classification creates primary sinus types with petiolar sinus at blade base showing highest diagnostic value, upper lateral sinuses between apical and upper lateral lobes, middle lateral sinuses between upper and lower laterals (when both present), and lower lateral sinuses between lower laterals and basal lobes showing variable presence depending on cultivar lobe configuration. Genetic control is strong making sinus characteristics relatively stable across environmental conditions, with depth, shape, and configuration patterns showing high heritability and serving as reliable identification features. Functional implications remain somewhat speculative, with proposed advantages including enhanced flexibility, modified boundary layers, and increased perimeter-to-area ratios affecting gas exchange, but lacking strong experimental support. Ampelographic utility is exceptional, with sinus characteristics particularly petiolar sinus configuration serving as primary diagnostic features in most identification systems.
Blattbucht
Laminaeinbuchtung
Lappeneinschnitt
blade indentation
leaf gap
lobe sinus
Leaf Sinus
The leaf size description of Mourvèdre documents large mature leaves (OIV 065:7) with blade length of 180-220 mm from petiolar junction to the tip of the central lobe, measured on representative leaves from nodes 8-12 of normally growing shoots — this large leaf size, combined with Mourvèdre's characteristically thick, leathery leaf texture and dark green color, creates a visually imposing canopy that contrasts with the medium-sized leaves of Grenache (OIV 065:5, blade length 140-170 mm) and the small leaves of Pinot Noir (OIV 065:3, blade length 100-130 mm). The leaf size measurements were standardized across three growing seasons to account for the substantial year-to-year variation driven by rainfall, temperature, and vigor differences that make leaf size the most environmentally variable of the standard leaf descriptors.
A mature leaf description component that records the dimensions of the fully expanded leaf blade according to OIV descriptor 065 (blade length) and related size parameters, scored on a scale from very small (state 1) through small, medium, large, to very large (state 9), providing an estimation of blade area that reflects the interaction between cultivar genotype and vine vigor, with leaf size contributing to cultivar characterization but being the least genetically stable of the major leaf descriptors due to strong environmental modification.
Leaf_Size_Description documents a quantitative character that is informative for cultivar characterization but must be interpreted cautiously due to its strong environmental dependency. Leaf blade area in V. vinifera varies by a factor of 5-8 across cultivars (from 400 cm² in large-leaved cultivars like Italia), but within a single cultivar, leaf area can vary by a factor of 2-3 depending on rootstock vigor, water availability, nitrogen nutrition, crop load, and canopy management practices: this substantial genotype × environment interaction makes leaf size the least reliable of the major leaf descriptors for cultivar identification in the absence of standardized growing conditions. The OIV system addresses this limitation by specifying precise sampling protocols (leaves from the middle third of normally growing shoots, not from excessively vigorous or weak shoots) and by recommending multi-year observation to establish the typical size range for each cultivar in a given site. Despite its variability, leaf size provides useful information for distinguishing cultivar groups at the extremes of the size distribution and for characterizing the vigor potential associated with different cultivar genotypes.
Blattflächen-Abschätzung
Blattgröße-Beschreibung
Blattspreiten-Dimension
blade area assessment
leaf dimension record
leaf size characterization
Leaf Size Description
Systematic assessment of Leaf_Size_Disposition at UC Davis compared individual leaf areas across 65 cultivars grown under standardized conditions (VSP, moderate irrigation, uniform vigor). Mature leaves from nodes 9-11 were harvested at full expansion, imaged, and area determined by digital analysis. Parallel measurements of leaf cell number and cell size (epidermal imprints) were conducted to investigate developmental basis. Leaf_Size_Disposition showed wide cultivar variation with distinct developmental patterns. 'Sultanina' (Thompson Seedless) exhibited large leaves with mean individual area of 365 cm² (range 320-410 cm² across seasons), representing approximately 4.2 million cells per leaf with mean cell area of 8,700 μm². 'Cabernet Sauvignon' showed intermediate Leaf_Size_Disposition (195 cm², 2.8 million cells, cell area 7,000 μm²). 'Petit Manseng' displayed small leaves (92 cm², 1.6 million cells, cell area 5,750 μm²), representing the lower range among commercial cultivars. Analysis of developmental components revealed that leaf size variation resulted from both cell number differences (2.6-fold range) and cell size differences (1.5-fold range), with cell number contributing approximately 65% of size variation. The functional significance was confirmed by canopy analysis—at equivalent shoot density (15 shoots/m), large-leaved 'Sultanina' achieved total leaf area of 4.8 m²/m cordon compared to 2.4 m²/m for small-leaved 'Petit Manseng', demonstrating the 2-fold impact of Leaf_Size_Disposition on canopy area. However, light distribution modeling showed that small-leaved canopies achieved more uniform light penetration, with interior leaf illumination 35% higher in 'Petit Manseng' than equivalent-area 'Sultanina' canopies.
A Leaf_Size_Disposition is a Leaf_Development_Disposition that inheres in Vitis vinifera leaves and is realized through the characteristic final area of individual mature leaves, determined by the duration and rate of cell division and expansion during leaf development. This disposition represents one of the most readily observable and functionally significant leaf characteristics, with individual leaf area varying more than 4-fold among cultivars from less than 100 cm² in small-leaved types to over 350 cm² in large-leaved cultivars. Leaf size is established during the approximately 30-40 day period of leaf expansion following emergence, with final size reflecting the genetically-determined number of cell divisions, cell expansion capacity, and responsiveness to environmental conditions during development. Leaf_Size_Disposition profoundly affects canopy function—large-leaved cultivars achieve equivalent total canopy leaf area with fewer leaves (lower shoot node number or shoot density), while small-leaved cultivars require more leaves to achieve equivalent area. Individual leaf size also affects boundary layer thickness and heat dissipation, transpirational cooling efficiency, and the distribution of light within the canopy (large leaves creating more self-shading). The disposition interacts with shoot vigor and node number to determine total canopy leaf area, with high-vigor cultivars with large leaves potentially producing excessive canopy area requiring management intervention.
Leaf_Size_Disposition is_a Leaf_Development_Disposition addressing individual mature leaf area. Leaf size ranges from 350 cm² among cultivars (4-fold range). Child classes include Large_Leaf_Disposition (> 200 cm²) and Small_Leaf_Disposition (< 150 cm²). Cell number variation (65%) contributes more than cell size (35%) to leaf area differences. Leaf size affects canopy area accumulation, light distribution, and boundary layer properties. OIV descriptor 065 (leaf blade size) provides standardized phenotyping.
Blattgrößen-Disposition
Blattspreiten-Größe
Einzelblattfläche
individual leaf area
leaf blade size
Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought, Plant Cell and Environment 26:1393-1405 Palliotti A et al. (2014) Changes in vineyard establishment and canopy management urged by earlier climate-related grape ripening: a review, Scientia Horticulturae 178:43-54 Organisation Internationale de la Vigne et du Vin (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition
Leaf Size Disposition
The leaf surface description of Merlot documents a moderately blistered adaxial surface (OIV 075:5) with a matte finish, combined with medium density of prostrate hairs on the abaxial surface between main veins (OIV 084:5), sparse erect hairs between veins (OIV 085:3), medium prostrate hairs on main veins (OIV 086:5), and sparse erect hairs on main veins (OIV 087:3) — this pubescence profile creates a leaf that is moderately felted on the lower surface with visible prostrate hair coverage but not the dense white wool of heavily pubescent cultivars like Viognier (OIV 084:9). The surface texture and pubescence combination contributes to Merlot's recognizable leaf feel — slightly rough adaxially from blistering, soft abaxially from the prostrate hair mat — that experienced ampelographers can identify by touch alone.
A mature leaf description component that documents the textural and trichome characteristics of both the adaxial (upper) and abaxial (lower) surfaces of the fully expanded leaf blade, recording blistering, glossiness, waxiness, and the type and density of trichomes on each surface according to OIV descriptors 075 and 084-087, providing cultivar-diagnostic characters based on the epidermal architecture and pubescence patterns of the mature leaf.
Leaf_Surface_Description integrates the visual and tactile properties of both leaf surfaces into a comprehensive characterization of the epidermal architecture. The adaxial surface is characterized primarily by blistering (the three-dimensional convexity of interveinal areas resulting from differential mesophyll expansion) and glossiness (reflecting the thickness and composition of the cuticle), while the abaxial surface is characterized primarily by trichome type and density — the four abaxial trichome descriptors (OIV 084-087) independently score prostrate and erect hairs in two locations (between main veins and on main veins), creating a four-dimensional pubescence profile that captures the cultivar-specific trichome architecture: this multi-parameter scoring is essential because trichome type and density vary independently across cultivars and across leaf regions — some cultivars show dense prostrate hairs between veins but sparse hairs on the veins themselves (suggesting vein-specific suppression of prostrate trichome development), while others show the reverse pattern, and the ratio of prostrate to erect trichome density creates cultivar-characteristic surface textures ranging from silky (dense prostrate, sparse erect) through bristly (sparse prostrate, dense erect) to glabrous (sparse both types).
Blatt-Epidermis-Morphologie
Blattoberflächen-Beschreibung
Blatttextur-Charakterisierung
blade surface morphology
leaf epidermal description
leaf texture characterization
Leaf Surface Description
Leaf temperature assessment of Chardonnay in Sonoma Coast vineyards using infrared thermometry reveals well-irrigated vines maintaining leaf temperature of 27°C despite 32°C air temperature (5°C depression indicating active transpiration at 4.2 mmol/m²/s), while deficit-irrigated vines show leaf temperature of 35°C (3°C elevation indicating stomatal closure), enabling irrigation scheduling based on thermal stress thresholds rather than fixed calendar intervals.
Leaf temperature quality is a tissue temperature quality that inheres in grapevine leaves and represents the thermal state of this photosynthetically active organ characterized by the dynamic balance between radiant energy absorption, convective heat exchange with surrounding air, and latent heat dissipation through transpiration that together determine the realized leaf temperature affecting photosynthetic rate, respiration, and stress signaling. Within the BFO framework, this quality specifically characterizes the leaf as the primary interface for atmospheric gas exchange and the organ with greatest capacity for active temperature regulation through stomatal control of transpiration. The ontological significance lies in leaf temperature determining photosynthetic efficiency, with optimal temperatures for Vitis vinifera photosynthesis typically ranging from 25-30°C, declining significantly above 35°C due to Rubisco kinetic limitations and photorespiration increase, and approaching zero above 40°C when heat stress induces stomatal closure and damages photosynthetic machinery. Unlike berries, leaves possess functional stomata enabling substantial evaporative cooling that can maintain leaf temperature below air temperature under conditions of adequate water supply and high transpiration demand, with temperature depressions of 3-8°C commonly observed in well-watered vines. Conversely, water-stressed vines with closed stomata cannot sustain transpirational cooling, and leaf temperatures rise above air temperature, making leaf-to-air temperature differential a sensitive indicator of plant water status exploited in thermal stress detection and irrigation scheduling. Leaf temperature measurement employs thermocouples for point measurements, infrared thermometers for rapid non-contact assessment, and thermal imaging for spatial distribution analysis revealing stomatal patchiness and stress heterogeneity within canopies.
Leaf_Temperature_Quality addresses the photosynthetically active organ whose temperature regulation capacity distinguishes it from berry temperature behavior. This quality supports water stress detection through canopy temperature monitoring and photosynthesis modeling requiring leaf temperature inputs. The class encompasses both sunlit and shaded leaves that differ substantially in temperature, with canopy position effects requiring specification for meaningful interpretation.
Blatttemperatur
Blattwärmezustand
Foliärtemperatur
Laubtemperatur
foliar temperature
lamina temperature
leaf thermal state
photosynthetic surface temperature
Jones HG (1999) Use of infrared thermometry for estimation of stomatal conductance as a possible aid to irrigation scheduling. Agricultural and Forest Meteorology 95(3): 139-149 Flexas J et al. (2002) Effects of drought on photosynthesis in grapevines under field conditions: an evaluation of stomatal and mesophyll limitations. Functional Plant Biology 29(4): 461-471
Leaf Temperature Quality
Physiological characterization of Leaf_Thickness_Disposition at the University of Burgundy measured leaf structural parameters and associated functional properties across 45 cultivars. Measurements included leaf thickness (micrometer), specific leaf mass (SLM, g fresh weight per m²), leaf dry matter content, mesophyll anatomy (palisade and spongy layer thickness), and functional correlates (area-based photosynthetic rate, water content, leaf thermal properties). Leaf_Thickness_Disposition showed substantial cultivar variation with clear functional associations. 'Grenache' exhibited thick leaves with SLM of 52 g/m², leaf thickness of 385 μm, well-developed two-layer palisade mesophyll (145 μm total), and dense spongy mesophyll with small intercellular spaces. Area-based photosynthetic capacity (Amax) reached 18.5 μmol CO₂ m⁻² s⁻¹, among the highest measured. 'Mourvèdre' showed similarly thick leaves (SLM 48 g/m²) with high photosynthetic capacity. 'Pinot Noir' displayed thin leaves with SLM of 32 g/m², thickness of 265 μm, single-layer palisade (75 μm), and loose spongy mesophyll with large air spaces. Amax was lower at 12.8 μmol m⁻² s⁻¹, though mass-based photosynthetic rate was actually similar. 'Riesling' showed intermediate thickness (SLM 38 g/m²). Correlation analysis confirmed that Leaf_Thickness_Disposition was associated with geographic origin—Mediterranean cultivars averaged SLM of 46 g/m² versus 36 g/m² for Northern European cultivars (P < 0.001). Drought tolerance trials revealed that thick-leaved cultivars maintained higher relative water content (85% versus 72% at Ψleaf = -1.5 MPa) and recovered more rapidly after stress, demonstrating the functional significance of Leaf_Thickness_Disposition for water relations.
A Leaf_Thickness_Disposition is a Leaf_Development_Disposition that inheres in Vitis vinifera leaves and is realized through the characteristic development of mesophyll tissue determining leaf thickness, specific leaf mass (mass per unit area), and the internal structural framework affecting photosynthetic capacity and stress tolerance. This disposition represents the investment of resources into leaf internal structure versus surface expansion, reflecting the balance between palisade mesophyll development (columnar photosynthetic cells), spongy mesophyll development (air space and gas exchange), and epidermal/structural tissue that together establish leaf thickness and density. Leaf thickness varies approximately 2-fold among cultivars, from thin leaves (specific leaf mass 45 g/m²) with multiple palisade layers and dense spongy mesophyll. Leaf_Thickness_Disposition has important functional implications—thicker leaves typically show higher photosynthetic capacity per unit area (more chloroplasts and Rubisco), greater water storage and drought tolerance, higher heat capacity moderating temperature fluctuations, and greater structural durability reducing physical damage. However, thick leaves also have higher construction cost and potentially slower expansion rates. The disposition reflects adaptation to environmental conditions, with thick leaves characteristic of cultivars from hot, dry Mediterranean environments (high light, water limitation) and thin leaves more common in cultivars from cooler, humid regions (lower light, adequate water).
Leaf_Thickness_Disposition is_a Leaf_Development_Disposition addressing mesophyll development and leaf density. Specific leaf mass (g/m²) is the standard quantitative measure. Thick leaves (SLM > 45 g/m²) show higher area-based photosynthetic capacity and drought tolerance. Thin leaves (SLM < 35 g/m²) have lower construction cost and potentially faster expansion. Mediterranean-origin cultivars typically show thicker leaves than Northern European cultivars. Palisade mesophyll development is the primary determinant of thickness variation. Environmental factors (light, water) modulate expression but genotypic ranking is stable.
Blattdicken-Disposition
Mesophyll-Entwicklung
Spezifische-Blattmasse
leaf mass per area
mesophyll development
specific leaf mass
Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought, Plant Cell and Environment 26:1393-1405 Tomás M et al. (2014) Variability of mesophyll conductance in grapevine cultivars under water stress conditions in relation to leaf anatomy and water use efficiency, Australian Journal of Grape and Wine Research 20:272-280 Flexas J et al. (2008) Mesophyll conductance to CO2: current knowledge and future prospects, Plant Cell and Environment 31:602-621
Leaf Thickness Disposition
Leaf water potential survey across canopy positions in Pinot Noir reveals midday values of -1.72 MPa in sun-exposed exterior leaves compared to -1.35 MPa in shaded interior leaves at the same vine, with the 0.37 MPa gradient corresponding to 3-fold difference in transpiration rate (4.2 vs 1.4 mmol/m²/s) and demonstrating the hydraulic heterogeneity within single vine canopies that complicates sampling protocols for representative whole-vine water status assessment.
Leaf water potential quality is a water potential quality that represents the water potential measured in actively transpiring leaf tissue, characterizing the most negative water potential in the liquid phase of the soil-plant-atmosphere continuum where the evaporative demand at the leaf surface creates the tension driving water uptake from soil through the plant. Within the BFO framework, this quality specifically characterizes the water status at the site of gas exchange and photosynthesis, distinguishing leaf water potential from stem or soil water potential and integrating the cumulative hydraulic resistance from soil through the entire plant pathway to the transpiring mesophyll. The ontological significance lies in leaf water potential determining the water status directly experienced by photosynthetic cells, affecting guard cell turgor and stomatal aperture, mesophyll cell hydration influencing photosynthetic capacity, and the risk of localized hydraulic failure in leaf vasculature. Leaf water potential in transpiring grapevine leaves typically ranges from -0.8 to -1.2 MPa under well-watered conditions during active transpiration, becoming more negative (-1.5 to -2.5 MPa) as soil water depletes or atmospheric demand increases, with the most negative values occurring in sun-exposed outer canopy leaves experiencing highest transpiration rates. The heterogeneity of leaf water potential within vine canopies reflects variation in exposure, transpiration rate, and hydraulic path length, with shaded interior leaves showing less negative values than exposed exterior leaves at the same time. Measurement employs pressure chamber technique on excised leaves, with rapid transfer from vine to chamber critical (ideally <30 seconds) to prevent artifact from continued transpiration after excision. The relationship between leaf water potential and stomatal conductance follows a sigmoid pattern in most grapevine cultivars, with stomata beginning to close at threshold potentials that vary among varieties reflecting their iso- or anisohydric behavior. Understanding leaf water potential quality enables direct assessment of the water status experienced by leaves, interpretation of stomatal and photosynthetic responses, and analysis of hydraulic gradients within the plant.
Leaf_Water_Potential_Quality addresses the water status at the site of gas exchange, distinguished from Stem_Water_Potential_Quality by measurement on actively transpiring (non-bagged) leaves. This quality shows greater variability than stem water potential due to leaf-to-leaf differences in exposure and transpiration rate, requiring careful sampling protocols specifying canopy position, leaf age, and measurement timing. The class represents the traditional water status measurement before stem water potential methods became widespread.
Blattlamina-Wasserpotential
Blattwasserpotential
Transpirationsblattwasserpotential
foliar water potential
foliäres Wasserpotential
lamina water potential
transpiring leaf potential
Ψleaf
Turner NC (1988) Measurement of plant water status by the pressure chamber technique. Irrigation Science 9(4): 289-308 Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell & Environment 26(8): 1393-1405
Leaf Water Potential Quality
Leaf wilting quality monitoring in Barossa Valley Shiraz established the relationship between wilting severity, water potential, and quality outcomes for deficit irrigation management. A four-point wilting scale was developed rating leaves from 0 (fully turgid, normal angle) through 2 (moderate wilting with petiole angle below horizontal but lamina still partially extended) to 4 (severe wilting with completely pendant petiole and collapsed lamina). Correlation with midday stem water potential showed wilting score 1 corresponding to -1.0 MPa indicating mild-moderate stress, score 2 at -1.2 MPa indicating moderate stress at the quality optimum boundary, and score 3 at -1.4 MPa indicating severe stress requiring irrigation. Critically, the timing of wilting relative to daily recovery proved diagnostic, with midday wilting recovering by early morning (score returning to 0-1 overnight) indicating stress within acceptable bounds, while persistent morning wilting signaled severe stress requiring immediate intervention. The practical utility of leaf wilting quality assessment enabled vineyard staff to conduct rapid daily stress evaluation supplementing weekly instrumental monitoring.
Leaf wilting quality is a drought symptom expression quality that inheres in grapevine leaves and represents the characteristic drooping or loss of turgid posture exhibited when leaf water potential declines below the threshold for turgor maintenance, causing the petiole and lamina to lose their normal angular orientation and assume a flaccid, pendant configuration that visually signals moderate-to-severe water deficit requiring management attention. This quality provides one of the most widely recognized visual indicators of plant water stress, with wilting representing a relatively advanced stress symptom that appears after more sensitive indicators and indicates stress severity sufficient to impair physiological function beyond the beneficial range for quality enhancement. The ontological significance of leaf wilting quality lies in its role as a universal stress indicator recognizable even by untrained observers, providing a clear visual signal that water deficit has exceeded compensatory capacity and that irrigation intervention may be warranted to prevent damage, though the appearance of wilting indicates stress has likely already impacted vine function beyond the optimal quality-enhancement zone. The biological basis of leaf wilting involves the loss of positive turgor pressure in leaf cells as water potential declines below the osmotic potential threshold, causing cells to lose the rigidity that maintains leaf orientation against gravity and resulting in the characteristic drooping appearance as petiole and lamina tissues become flaccid. The reversibility of wilting provides diagnostic information, with temporary midday wilting recovering overnight indicating moderate stress within the vine's recovery capacity, while persistent wilting not recovering overnight signals severe stress potentially causing lasting damage. Cultivar differences in leaf wilting quality expression reflect variation in osmotic adjustment capacity enabling turgor maintenance at lower water potentials, cuticular properties affecting water loss rates, and leaf structural characteristics determining how readily turgor loss translates to visible postural change. The timing of wilting relative to other stress indicators varies with environmental conditions, appearing earlier under high evaporative demand that accelerates water loss and later under moderate conditions that permit gradual adjustment.
Leaf_Wilting_Quality is_a Drought_Symptom_Expression_Quality, representing the turgor-loss-driven drooping of leaves under water deficit. This class addresses one of the most recognizable stress symptoms though it appears at relatively advanced stress levels. The recovery pattern (midday wilting vs. persistent wilting) provides critical diagnostic information distinguishing recoverable from damaging stress. Wilting score 2 approximately corresponds to moderate stress at the beneficial/harmful boundary. Distinguished from Leaf_Rolling_Quality which represents a different morphological response involving lamina curling rather than petiole drooping.
Blatterschlaffung
Blattwelke
Turgeszenz-Verlust
Welkesymptom
leaf drooping
leaf flaccidity
turgor loss
wilting symptom
Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell and Environment 26:1393-1405 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data. Annals of Botany 105:661-676 Lovisolo C et al. (2010) Drought-induced changes in development and function of grapevine (Vitis spp.) organs. Functional Plant Biology 37:98-116
Leaf Wilting Quality
Comparative assessment of Leafhopper_Resistance_Disposition at Texas A&M AgriLife Research evaluated responses to both xylem-feeding glassy-winged sharpshooter (Homalodisca vitripennis) and phloem-feeding grape leafhopper (Erythroneura elegantula) across 25 Vitis accessions using electrical penetration graph (EPG) monitoring and population development assays. EPG analysis revealed that H. vitripennis achieved sustained xylem ingestion (>30 minute continuous feeding bouts) on 92% of V. vinifera 'Chardonnay' test sessions but only 35% of sessions on V. arizonica b43-17, with resistant plants eliciting frequent stylet withdrawal and non-feeding probing behavior. For E. elegantula, oviposition choice tests showed 4.2-fold higher egg deposition on 'Chardonnay' (28 eggs/leaf) versus the resistant V. cinerea accession C166-026 (6.7 eggs/leaf), with subsequent nymphal survival of 78% versus 32% respectively. Importantly, Leafhopper_Resistance_Disposition against the two feeding guilds showed weak correlation (r = 0.34), with some genotypes resistant to sharpshooters but susceptible to grape leafhoppers and vice versa, indicating distinct resistance mechanisms targeting xylem- versus phloem-feeders. These findings support breeding strategies combining resistance sources effective against both leafhopper types for comprehensive protection in regions facing both Pierce's disease pressure and direct leafhopper damage.
A Leafhopper_Resistance_Disposition is an Insect_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that limit feeding, reproduction, or damage by leafhoppers (Hemiptera: Cicadellidae), a diverse family of phloem- and xylem-feeding insects including both direct pests causing stippling damage and critically important vectors of plant pathogens. This disposition represents resistance to insects that attack grapevine through piercing-sucking mouthparts, extracting plant fluids and potentially transmitting bacterial or phytoplasma pathogens during the feeding process. Leafhoppers affecting grapevine fall into two functional groups with distinct impacts: xylem-feeding sharpshooters (subfamily Cicadellinae, tribe Cicadellini and Proconiini) that vector Xylella fastidiosa causing Pierce's disease, and phloem-feeding leafhoppers (subfamily Typhlocybinae, particularly genera Erythroneura and Empoasca) that cause direct feeding damage through chlorophyll destruction and cell emptying, producing characteristic stippling and bronzing of leaves. Resistance mechanisms against leafhoppers include leaf surface properties (trichome density, epicuticular wax composition) affecting settling and probing behavior, vascular tissue characteristics influencing feeding efficiency, chemical factors in xylem or phloem sap deterring sustained feeding, and induced defenses reducing leafhopper performance. Given the devastating impact of Pierce's disease vectored by sharpshooters, Leafhopper_Resistance_Disposition has dual importance for direct damage reduction and pathogen transmission interruption, making it a priority trait for breeding in Pierce's disease-endemic regions.
Leafhopper_Resistance_Disposition is_a Insect_Resistance_Disposition. Leafhoppers comprise two functionally distinct groups affecting grapevine: xylem-feeding sharpshooters (Pierce's disease vectors) and phloem-feeding grape leafhoppers (direct pests). Child classes include Glassy_Winged_Sharpshooter_Resistance_Disposition and Empoasca_Resistance_Disposition addressing these distinct pest groups. The vector importance of sharpshooters for Xylella fastidiosa creates critical linkage between Leafhopper_Resistance_Disposition and Pierces_Disease_Resistance_Disposition. Resistance mechanisms differ between xylem and phloem feeders due to their distinct target tissues and feeding behaviors.
Blattzikaden-Abwehr
Cicadellidae resistance
Kleinzikaden-Resistenz
Zikaden-Resistenz-Disposition
hopper resistance
leafhopper feeding deterrence
Backus EA et al. (2005) Sensory systems and behaviours which mediate hemipteran plant-feeding: a taxonomic overview, Journal of Insect Physiology 51:791-803 Daane KM, Williams LE (2003) Manipulating vineyard irrigation amounts to reduce insect pest damage, Ecological Applications 13:1650-1666 Brodbeck BV et al. (2011) Physiological and behavioral adaptations of three species of leafhoppers in response to the dilute nutrient content of xylem fluid, Journal of Insect Physiology 39:73-81
Leafhopper Resistance Disposition
Field evaluation of Leafroll_Resistance_Disposition in a naturally infected Napa Valley vineyard compared disease progression and economic impact across 12 cultivars over 8 growing seasons. Virus status was monitored annually by ELISA, with yield and fruit quality parameters measured at harvest. The susceptible cultivar 'Merlot' showed rapid disease progression with 85% infection by year 5, exhibiting severe symptoms (OIV 467 rating 7-9), mean yield reduction of 32% (6.2 vs 9.1 tonnes/ha in healthy vines), Brix reduction of 2.8°, and anthocyanin decline of 45%. 'Cabernet Sauvignon' demonstrated enhanced Leafroll_Resistance_Disposition (tolerance phenotype) with equivalent infection rate but markedly reduced impact: 18% yield reduction, 1.4° Brix reduction, and 22% anthocyanin decline. Symptom expression was less severe (OIV rating 3-5) despite similar viral titers as measured by qRT-PCR. The white cultivar 'Chardonnay' showed the highest tolerance with only 8% yield impact and minimal quality degradation, though symptoms were also inherently less visible. Vector monitoring revealed no cultivar differences in mealybug population densities, indicating that differential Leafroll_Resistance_Disposition reflected host tolerance to viral infection rather than vector resistance.
A Leafroll_Resistance_Disposition is a Viral_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that limit infection, replication, systemic spread, or symptom expression by Grapevine leafroll-associated viruses (GLRaVs), a complex of phloem-limited ampeloviruses and closteroviruses causing grapevine leafroll disease. This disposition represents the capacity to resist one of the most economically significant and widespread viral disease complexes affecting viticulture globally, present in virtually all wine-producing regions. Leafroll disease causes characteristic symptoms in red cultivars including downward rolling of leaf margins, interveinal reddening with green veins, and leaf thickening, while white cultivars show more subtle chlorosis and rolling. Physiological impacts include delayed and uneven ripening, reduced sugar accumulation (1-5°Brix reduction), elevated acidity, diminished anthocyanin and flavor compound synthesis, and yield losses of 15-40%. The GLRaV complex includes multiple species (GLRaV-1, -2, -3, -4, and variants) transmitted by mealybugs (Pseudococcus, Planococcus) and soft scale insects (Parthenolecanium, Pulvinaria) in a semi-persistent manner. Leafroll resistance dispositions can operate through resistance to viral replication and phloem loading, resistance or tolerance to insect vector feeding, and tolerance mechanisms maintaining productivity despite infection. Unlike fanleaf disease where resistant rootstocks effectively control transmission, leafroll is transmitted to aerial plant parts by mobile insect vectors, making scion resistance more critical. No strong genetic resistance to GLRaVs has been identified in V. vinifera, though significant tolerance variation exists among cultivars.
Leafroll_Resistance_Disposition is_a Viral_Resistance_Disposition. Leafroll disease involves multiple viral species (GLRaV-1 through GLRaV-9, with GLRaV-3 most prevalent and damaging), complicating resistance evaluation. Child classes include GLRaV_Replication_Resistance_Disposition for direct antiviral resistance and Vector_Transmission_Resistance_Disposition for insect vector resistance. Unlike fanleaf, leafroll transmission by aerial insect vectors cannot be controlled through rootstock selection alone. Tolerance (maintaining productivity despite infection) is currently more relevant than true resistance for practical viticulture, as no strong resistance sources have been identified in V. vinifera.
Blattroll-Resistenz-Disposition
Blattrollkrankheit-Resistenz
GLRaV resistance disposition
GLRaV-Resistenz
ampelovirus resistance
leafroll tolerance
Naidu RA et al. (2014) Grapevine leafroll disease and associated viruses: a unique pathosystem, Annual Review of Phytopathology 52:61-83 Maree HJ et al. (2013) Grapevine leafroll-associated virus 3, Frontiers in Microbiology 4:82 Almeida RPP et al. (2013) Ecology and management of grapevine leafroll disease, Frontiers in Microbiology 4:94
Leafroll Resistance Disposition
Leafroll symptom quality assessment in Sonoma County Pinot Noir documented the disease impact on fruit quality motivating an intensive roguing and replanting program. Systematic vineyard survey in late September identified vines showing leafroll symptom quality with the characteristic downward rolling of mature leaf margins combined with interveinal reddening while primary and secondary veins remained distinctly green, creating the "Christmas tree" color pattern pathognomonic for leafroll in red varieties. Symptom intensity was scored on a 0-5 scale, with most symptomatic vines showing scores of 3-4 indicating moderate to severe interveinal reddening affecting greater than 50% of canopy leaves. Fruit from symptomatic vines showed leafroll quality impacts including Brix reduction averaging 2.3° compared to adjacent asymptomatic vines at harvest, anthocyanin concentration reduced by 35%, and berry weight reduced by 15%, collectively establishing that symptomatic vines were producing fruit unsuitable for the premium Pinot Noir program despite appearing visually normal in the cluster. Laboratory testing confirmed GLRaV-3 presence in 92% of symptomatic vines, with viral load (quantitative PCR) correlating with symptom severity (r² = 0.68). The leafroll symptom quality-guided roguing program removed symptomatic vines over 3 years, combined with mealybug monitoring and control to limit vector transmission, reducing block incidence from 28% to 5% and restoring fruit quality from replanted vines to premium standards.
Leafroll symptom quality is a viral infection indicator quality that inheres in grapevine organisms infected by grapevine leafroll-associated viruses (GLRaVs, primarily GLRaV-3) and represents the characteristic disease manifestations including the downward rolling of leaf margins beginning in late summer, the interveinal reddening or purpling in red-fruited cultivars or interveinal chlorosis in white-fruited cultivars while veins remain green, the delayed fruit ripening with reduced sugar accumulation, and the diminished anthocyanin development in red varieties, with symptom expression intensifying progressively through the season and being most pronounced in autumn before leaf senescence. This quality captures the diagnostic features of the most economically important and widespread viral disease complex affecting grapevine worldwide, recognizing that leafroll viruses cause substantial yield reductions (10-40%) and more significantly affect fruit quality through reduced Brix, color, and flavor development that can reduce wine value by 30-70% even when yield impacts are modest. The ontological significance of leafroll symptom quality lies in its role enabling identification of a disease with major economic impact on wine quality, where symptom recognition in established vineyards informs roguing decisions and vector management, while in nursery settings detection prevents propagation and distribution of infected material. The biological basis of leafroll symptoms encompasses the phloem-limited replication of GLRaVs that disrupts carbohydrate transport from leaves to fruit, causing the sugar accumulation in leaves that triggers anthocyanin synthesis producing the characteristic leaf reddening, while simultaneously reducing sugar export to berries causing the delayed ripening and reduced Brix characteristic of affected fruit. The vector transmission of GLRaVs by mealybugs (Pseudococcus, Planococcus species) and soft scale insects creates potential for secondary spread within and between vineyards, with vector management essential for limiting disease increase in affected blocks and protecting adjacent healthy plantings. The cultivar-specific symptom expression creates diagnostic challenges, with some cultivars showing dramatic symptoms while others remain nearly symptomless despite equivalent viral loads, necessitating laboratory confirmation for definitive diagnosis and creating risk of symptomless spread through propagation from apparently healthy but infected vines.
Leafroll_Symptom_Quality is_a Viral_Infection_Indicator_Quality, representing the symptoms of grapevine leafroll-associated virus (primarily GLRaV-3) infection. The downward leaf rolling with interveinal reddening (red cvs) or chlorosis (white cvs) is pathognomonic. Phloem disruption causes both leaf symptoms (sugar accumulation) and fruit symptoms (reduced Brix). Mealybug and soft scale vector transmission enables secondary spread. Symptom expression varies among cultivars some remain nearly symptomless. Most economically important grapevine virus worldwide affecting wine quality more than yield.
Blattroll-Symptome
GLRaV symptoms
GLRaV-Symptome
Rebenblattroll-Symptome
grapevine leafroll symptoms
leafroll disease symptoms
viral leaf reddening
virale Blattrötung
Naidu RA et al. (2015) Grapevine leafroll disease and associated viruses: a unique pathosystem. Annual Review of Phytopathology 53:613-634 Almeida RPP et al. (2013) Ecology and management of grapevine leafroll disease. Frontiers in Microbiology 4:94 Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests, 2nd Edition. APS Press
Leafroll Symptom Quality
A left unilateral cordon on Vitis vinifera cv. Pinot Noir at right-side row end (when traveling north along row), extending 89cm to the left (westward) from trunk positioned at row terminus, bearing 14 spurs, with vineyard map documentation clearly indicating "left unilateral" enabling equipment operators to approach row end knowing cordon extends to their left when traveling north, facilitating proper turning sequences for mechanical hedger and harvester avoiding damage to cordon terminus, representing directionally-specified unilateral architecture where left designation provides essential spatial reference for coordinated operations in large commercial vineyard with multiple equipment operators and management crews requiring precise directional communication.
A unilateral cordon extending exclusively in the left direction from the trunk when viewed from a standard perspective along the vineyard row, characterized by single-sided extension toward the left side of the row, absence of right-side cordon development, and directional specificity important for vineyard mapping, management planning, and equipment operation coordination. Left unilateral cordons represent directionally specified unilateral architecture, with left designation providing precise spatial reference for vineyard documentation, equipment operators (enabling specific instructions for row-end approaches and turning sequences), and management crews (coordinating operations requiring directional awareness such as selective spray applications, differential shoot thinning, or asymmetric canopy management), with the left versus right distinction being arbitrary but essential for clear communication in complex vineyard operations.
Left unilateral cordons are functionally identical to right unilateral cordons, with the distinction being purely directional reference rather than architectural or physiological difference. The left versus right designation depends on viewing perspective, with vineyard management typically establishing standard reference conventions (e.g., "left and right when traveling north along rows" or "left and right when facing upslope") ensuring consistent communication. The directional specificity becomes important in large operations where equipment operators, pruning crews, and management personnel must coordinate activities across extensive vineyards, with clear directional designations preventing confusion and operational errors. In small vineyards or simple layouts, the left versus right distinction may be unnecessary, with generic "unilateral" or "row-end" designations providing adequate information. Documentation systems increasingly incorporate directional specifications in vineyard databases and GIS mapping, enabling precise spatial referencing for targeted management interventions.
left-extending cordon
left-side cordon
linke unilaterale Kordon
linksseitige Kordon
sinister cordon
Left Unilateral Cordon
Legs quality comparison between fortified Sherry (pronounced, slow-descending legs with thick rivulets reflecting 18% alcohol) and premium Burgundy Grand Cru (subtle, rapidly descending legs despite exceptional quality, reflecting 13% alcohol) demonstrates the primacy of alcohol over quality in leg formation, correcting the common consumer assumption that the Sherry's impressive legs indicate superior quality when in fact the Burgundy represents far higher quality despite modest leg display, illustrating the importance of educated leg interpretation.
Legs quality is a viscosity appearance quality that represents the rivulet patterns formed on glass surfaces as wine drains after swirling, characterizing the arch formation, droplet size, flow rate, and persistence of these drainage streams as visual indicators primarily of alcohol content and secondarily of glycerol and extract concentration. Within the BFO framework, this quality specifically addresses the drainage pattern dimension of viscosity assessment, distinguishing leg formation from overall flow behavior while recognizing that legs (also termed tears, church windows, or cathedral arches) arise from the Marangoni effect where surface tension gradients drive characteristic flow patterns. The ontological significance lies in legs serving as the most commonly observed and discussed viscosity indicator, though popular understanding significantly overestimates the relationship between leg quality and wine quality or glycerol content, with alcohol being the primary determinant of leg formation rather than the quality-associated factors often assumed. The physical basis for leg formation involves the Marangoni effect where evaporation of alcohol from the thin wine film on glass walls creates local surface tension increases (alcohol has lower surface tension than water), generating upward flow that accumulates into droplets that periodically release as descending legs, with leg frequency, size, and descent rate reflecting alcohol concentration and evaporation dynamics. The compositional determinants of leg quality include alcohol content (primary factor, with higher alcohol producing more pronounced, slower-descending legs), glycerol concentration (minor contribution to leg viscosity), residual sugar (increasing leg thickness and slowing descent), and glass cleanliness (residues interfering with proper film formation and leg development). The interpretation cautions for leg assessment are essential, as the common belief that impressive legs indicate quality or glycerol content is largely mistaken, with high-alcohol wines showing pronounced legs regardless of quality level, and many excellent lower-alcohol wines showing minimal leg formation. Understanding legs quality enables alcohol level estimation from leg behavior, appropriate interpretation avoiding quality inference from legs alone, and educated communication about the actual physical basis for this commonly discussed visual phenomenon.
Legs_Quality_(Tears) addresses the specific drainage pattern phenomenon arising from the Marangoni effect. This quality is frequently misinterpreted, with common belief incorrectly attributing legs to glycerol or quality rather than their actual primary determinant (alcohol). The class supports accurate interpretation and education about the physical basis of leg formation while cautioning against quality inference from this visual phenomenon.
Kirchenfenster
Marangoni effect patterns
Schlieren
Tränen
Weinbeine
cathedral windows
church windows
wine tears
Vuilleumier C et al. (1995) Tears of wine: the Marangoni effect. American Journal of Physics 63(12): 1095-1096 Hosoi AE, Bush JWM (2001) Evaporative instabilities in climbing films. Journal of Fluid Mechanics 442: 217-239
Legs Quality (Tears)
Vigor assessment in Syrah vineyards of the Northern Rhône utilizes mean internode length as a key diagnostic, with optimal balance indicated by values of 6-8 cm on primary shoots shorter internodes (below 5 cm) suggest water deficit or excessive crop load, while longer internodes (above 10 cm) indicate excessive vigor requiring canopy management intervention through shoot positioning, hedging, or cover crop competition.
Length_Quality represents the linear dimension measured along the principal longitudinal axis of a grapevine organ, typically constituting the greatest extent in elongated structures. Length is the primary dimension for characterizing shoots, internodes, petioles, tendrils, roots, cluster rachises, and pedicels, where the growth axis defines a natural measurement direction. For organs without an obvious elongation axis, length is conventionally defined as the greatest linear dimension. Length measurements are foundational to grapevine phenotyping because they capture growth vigor, developmental progression, and genetic potential. Shoot length accumulates through the season as a direct indicator of vegetative vigor, with well-balanced vines producing 15-20 nodes on main shoots by véraison. Internode length reflects the rate of shoot elongation during the period of internode formation, integrating temperature and resource availability signals. Root length and root system depth determine soil exploration capacity and water acquisition potential. Cluster rachis length influences bunch compactness and disease susceptibility, with longer rachises promoting air circulation and reducing Botrytis pressure. Berry length, measured from pedicel attachment to the stylar scar, contributes to the length-to-width ratio used in shape classification. Length measurements employ digital calipers for organs under 200 mm and tape measures or graduated rulers for larger structures. Standardized protocols specify measurement from defined proximal to distal landmarks appropriate to each organ type.
Length_Quality inherits from Linear_Dimension_Quality and represents the longitudinal measurement axis. For spherical or near-spherical organs such as berries, the distinction between length and width may be arbitrary in such cases, length conventionally refers to the polar axis (pedicel to stylar end) while width refers to the equatorial diameter. Length correlates with but is distinct from growth rate (a process characteristic) and growth potential (a disposition). PATO_0000122 (length) provides the upper-ontology alignment.
Longitudinaldimension
Länge
Längenausdehnung
Längenmessung
axial length
longitudinal dimension
longitudinal extent
organ length
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 67-95 Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles, Adelaide pp.: 24-31
Length Quality
Merlot lenticel in trunk bark at 16 years post-planting shows 2.5-3.8 mm diameter oval structure appearing as light tan spot on grey-brown cork surface, lenticel comprising loosely arranged cork cells with large intercellular air spaces (55-68% void volume), extends 1.4-2.2 mm depth through cork layer reaching living phelloderm, enables gas exchange 0.5-0.9 μmol O₂/cm²/hour inward diffusion supporting living inner bark respiration 0.6-1.2 μmol O₂/g bark/hour, lenticel density 4-6 per cm² bark surface (collectively lenticels occupying 3-4% of bark area enabling adequate gas exchange while maintaining 96-97% impermeable coverage), lenticel showing slight raising 0.2-0.4 mm above surrounding cork surface
Lenticel is a specialized porous structure in Vitis vinifera trunk bark comprising loosely arranged cork cells with large intercellular air spaces that interrupt the otherwise impermeable periderm, enabling gas exchange between atmosphere and living inner bark tissues while maintaining overall waterproof and pathogen-resistant bark covering. Lenticel structure and composition includes= loose cork cells (complementary tissue or filling tissue, cells with minimal cell-to-cell contact creating tortuous air-filled pathways), large intercellular spaces (air channels 5-20 μm width occupying 50-70% of lenticel volume enabling gas diffusion), cork cambium beneath lenticel (phellogen producing loosely arranged cork cells in lenticel region rather than compact cells typical of surrounding cork), closing layer potential (some lenticels forming compacted cell layer during dormancy partially restricting gas exchange and water loss), and boundary with normal cork (gradual transition from compact impermeable cork to loose porous lenticel tissue). Lenticel dimensions and distribution includes= lenticel diameter 1-4 mm (circular to slightly oval appearance on bark surface), lenticel depth extending through cork layer (typically 0.5-3 mm deep reaching living phelloderm or phloem ensuring gas exchange pathway), lenticel density 2-8 per cm² bark surface (varying by cultivar, bark age, and environmental conditions), scattered distribution (lenticels distributed relatively evenly across bark surface though sometimes aligned in vertical rows or associated with former bud or leaf scars), and small proportional area (95% waterproof bark coverage). Lenticel and pathogen entry includes= potential infection pathway (lenticels providing porous route through otherwise impermeable cork, some pathogens potentially entering through lenticels), limited depth of penetration (most pathogens unable to penetrate deep through lenticel air spaces, physical and chemical barriers limiting infection), wound lenticels more vulnerable (traumatic lenticels in wound periderm potentially more susceptible to pathogen colonization than normal developmental lenticels), importance of healthy bark (maintaining vigorous periderm formation ensuring functional lenticels with appropriate structure minimizing infection risk), and relative safety (despite porosity, lenticels typically not major infection route compared to bark wounds, cracks, or injury sites). Lenticel in wound healing includes= traumatic lenticels forming in wound periderm (wound-induced cork cambium often producing lenticels in new protective covering beneath injuries), enabling respiration in callus (lenticels in wound periderm supporting metabolic activity in proliferating callus tissue and differentiating cells), gradual lenticel development (traumatic lenticels appearing weeks to months after wounding as protective cork layer matures), and integration with wound periderm (lenticels becoming functional component of mature wound covering enabling long-term gas exchange in healed tissue). Lenticel variation by bark characteristics includes= smooth bark showing prominent lenticels (conspicuous spots or lines on uniform surface), furrowed bark with less obvious lenticels (lenticels blending with bark texture, located in furrow bottoms or on ridges), young bark with smaller shallower lenticels (1-2 mm diameter, 0.3-0.8 mm deep in first periderm), mature bark with larger deeper lenticels (2-4 mm diameter, 1-3 mm deep in thick cork layers), and cultivar-specific lenticel characteristics (some varieties showing large prominent lenticels, others small inconspicuous structures). Lenticel significance in bark function includes= enabling aerobic metabolism (lenticels absolutely essential for maintaining living inner bark, without lenticels cork would suffocate tissues causing death), balancing impermeability and permeability (lenticels solving challenge of maintaining waterproof pathogen-resistant covering while enabling necessary gas exchange), accommodating bark thickening (lenticels maintaining depth and function as successive periderms form ensuring continued gas exchange through increasingly thick bark), and critical for trunk survival (adequate lenticel function essential for inner bark and cambium viability, inadequate lenticels potentially causing reduced vigor, susceptibility to stress, or tissue death).
Lenticel emphasizes specialized porous structure and is distinguished from general periderm (impermeable cork), Gas_Exchange_Lenticel (function emphasis), and Surface_Lenticel (location emphasis) by focus on anatomical specialization. Critical for understanding bark gas exchange mechanism, aerobic metabolism support in living inner bark, balance between impermeability and permeability, lenticel-mediated oxygen supply and CO₂ removal, and essential role in maintaining inner bark and cambium viability through waterproof cork covering requiring adequate lenticel density and function.
Aerating structure
Bark pore
Gas exchange pore
Gasaustauschs-Struktur
Lenticelle
Rindenpore
Lenticel
Lesion coverage quality assessment for powdery mildew (Erysiphe necator) on Chardonnay leaves and clusters employed standardized area diagrams to quantify disease severity for fungicide efficacy trials. Assessment training using reference photographs showing 1%, 5%, 10%, 25%, 50%, and 75% coverage established consistent evaluation standards among assessors. Leaf lesion coverage quality was assessed on 20 leaves per plot at BBCH 79, with coverage ranging from 2% in the most effective fungicide treatment to 45% in untreated controls, demonstrating fungicide program efficacy for reducing disease development. Cluster lesion coverage quality assessment focused on the percentage of visible berry surface showing powdery mildew colonies, with coverage ranging from below 1% in effective treatments to 28% in untreated controls. Image analysis of photographed clusters provided validation of visual estimates, with digital coverage determination showing strong correlation (r² = 0.91) with visual assessments, confirming the reliability of trained visual evaluation for routine assessment while establishing image analysis protocols for research requiring maximum precision. The lesion coverage quality threshold for acceptable wine quality was established at 15% cluster coverage, above which elevated potassium, pH, and volatile acidity in resulting wines indicated quality compromise, enabling treatment efficacy evaluation against a quality-relevant standard rather than arbitrary coverage targets.
Lesion coverage quality is a disease severity quality that inheres in grapevine organs and represents the proportion of tissue surface area occupied by visible disease lesions or pathogen structures, providing a physically measurable expression of disease intensity that can be assessed visually, photographically, or through image analysis to quantify the extent of tissue colonization by pathogens producing discrete lesions, sporulating colonies, or other visible surface manifestations. This quality captures a specific and widely applicable measure of disease severity based on the physical coverage of tissue by disease symptoms, recognizing that many important grapevine diseases produce visible surface lesions whose cumulative area provides a quantifiable measure of disease severity that correlates with functional damage and crop loss. The ontological significance of lesion coverage quality lies in its objectivity and reproducibility as a severity measure, with percentage coverage providing a continuous quantitative variable that can be assessed consistently across observers when standardized methods and reference aids are employed, supporting rigorous disease quantification for research, management decision-making, and efficacy evaluation. The biological basis of lesion coverage encompasses the expansion and coalescence of disease lesions as pathogen colonization progresses, with coverage increasing from initial small infection foci through lesion enlargement and multiplication toward eventual confluence that may approach 100% coverage under severe disease conditions. The assessment methods for lesion coverage include visual estimation aided by standard area diagrams showing reference coverage percentages, photographic documentation with subsequent image analysis for precise coverage determination, and molecular or biochemical proxies that correlate with physical coverage when direct visual assessment is impractical. The relationship between lesion coverage and functional impact varies with the tissue affected and the nature of the pathogen, with photosynthetic impairment generally proportional to leaf coverage, fruit quality impact often showing threshold effects where coverage exceeding critical levels causes disproportionate damage, and the distinction between surface colonization and tissue destruction affecting the coverage-impact relationship for different pathogen types.
Lesion_Coverage_Quality is_a Disease_Severity_Quality, representing the proportion of tissue surface occupied by visible disease lesions. This class addresses physical area coverage as an objective severity measure. Standard area diagrams enable consistent visual assessment across observers. Image analysis provides precision measurement for research applications. The coverage-quality threshold (15% for powdery mildew in example) enables quality-relevant assessment. Leaf versus cluster coverage may have different quality implications. The continuous percentage variable enables statistical analysis of treatment effects.
Läsionsbedeckung
Läsionsfläche
Oberflächenkolonisierung
disease coverage percentage
lesion area
percent coverage
prozentuale Bedeckung
surface colonization
Seem RC (1984) Disease incidence and severity relationships. Annual Review of Phytopathology 22:133-150 Calonnec A et al. (2004) Effects of Uncinula necator on the yield and quality of grapes (Vitis vinifera) and wine. Plant Pathology 53:434-445 Bock CH et al. (2010) Plant disease severity estimated visually, by digital photography and image analysis, and by hyperspectral imaging. Critical Reviews in Plant Sciences 29:59-107
Lesion Coverage Quality
Licorice aroma evaluation in Châteauneuf-du-Pape Grenache-dominated blends reveals characteristic licorice expression (6.5 on 10-point scale) complementing kirsch fruit and garrigue herbs in optimal vintages, while comparison with Priorat Garnacha shows higher licorice intensity (7.8) attributed to lower yields and warmer conditions concentrating phenylpropanoid precursors, demonstrating how licorice intensity varies with terroir while remaining a consistent varietal marker, with both expressions considered typical for their respective appellations' ripe Mediterranean style.
Licorice aroma quality is a spicy aroma quality that represents olfactory perception evoking Glycyrrhiza glabra (licorice root) and anise-associated scents, characterizing the distinctive sweet, aromatic impression with herbal-spicy undertones particularly associated with ripe red wines, especially Grenache and certain Mediterranean varieties. Within the BFO framework, this quality specifically addresses the licorice and anise subcategory of spicy aroma, distinguished by its sweet-spicy character and association with both grape ripeness and certain winemaking influences. The ontological significance lies in licorice aroma contributing to the complexity of ripe, full-bodied red wines, where licorice notes indicate maturity, extraction, and often the characteristic expression of southern French, Spanish, and Italian Mediterranean varieties, serving as a regional typicity marker alongside fruit and herbal characters. The compositional basis for licorice aroma includes anethole (primary contributor to anise and licorice perception, threshold approximately 50 μg/L), estragole (providing tarragon-anise notes), and various related phenylpropanoid compounds, with sources including both grape-derived compounds in certain varieties and potential oak contributions from specific toast profiles. The variety associations of licorice character are strongest for Grenache (where licorice is considered a varietal marker at optimal ripeness), Mourvèdre (contributing to Bandol typicity), and certain Spanish varieties including aged Tempranillo and Monastrell, while licorice notes in other varieties more typically indicate oak influence or extended maceration. The ripeness relationship shows licorice character developing with full grape maturity, with underripe fruit lacking licorice potential while optimally ripe grapes in warm climates express licorice as part of complex spice-fruit integration, and overripe fruit potentially showing excessive or confected licorice character. Understanding licorice aroma quality enables variety identification for licorice-associated varieties, ripeness assessment from licorice development, and regional typicity evaluation where licorice contributes to Mediterranean wine character.
Licorice_Aroma_Quality addresses the licorice and anise subcategory of spicy aroma, associated with specific varieties (particularly Grenache) and full ripeness. This quality occupies a position between fruity and spicy categories, with the sweet-spicy character bridging these aromatic dimensions. The distinction between grape-derived licorice (varietal expression) and oak-influenced licorice requires context consideration in aroma source assessment.
Anisduft
Lakritzaroma
Lakritznote
Süßholzcharakter
anise aroma
fennel-like character
licorice root notes
réglisse scent
Ferreira V et al. (2000) Quantitative gas chromatography-olfactometry carried out at different dilutions of an extract. Key differences in the odor profiles of four high-quality Spanish aged red wines. Journal of Agricultural and Food Chemistry 48(10): 4097-4105 Aznar M et al. (2001) Identification and quantification of impact odorants of aged red wines from Rioja. Journal of Agricultural and Food Chemistry 49(6): 2924-2929
Licorice Aroma Quality
The lieu-dit name Les Petits Musigny designates a 0.75-hectare cadastral parcel within the commune of Chambolle-Musigny classified as Premier Cru, while the adjacent lieu-dit Les Amoureuses (5.4 hectares, also Premier Cru) produces wines widely considered to approach Grand Cru quality — in the Rhône Valley, the lieu-dit La Mouline within the Côte-Rôtie appellation identifies a specific slope parcel from which E. Guigal produces one of the most celebrated single-vineyard Syrahs, with the lieu-dit name on the label communicating site specificity within the broader 300-hectare Côte-Rôtie appellation area.
A vineyard designation within the French cadastral and viticultural system that identifies a wine as originating from a named lieu-dit — a traditional place name of cadastral origin designating a specific parcel or group of parcels within a commune, recorded in the official land registry (cadastre) and recognized by INAO as a smaller geographic unit within an appellation whose name may appear on wine labels as a site-specific origin indicator.
Lieu_Dit_Name derives from the French cadastral system where lieu-dit (literally said place or spoken place) designates a traditional place name recorded in the official land survey — these names often originate from medieval or earlier topographic, agricultural, or historical references (Les Caillerets = stony ground, Clos de Vougeot = enclosed vineyard of Vougeot, Les Charmes = fallow land). In viticultural usage, lieu-dit names function as the building blocks of the French vineyard classification system: Burgundy's Premier Cru and Grand Cru designations are fundamentally inventories of classified lieux-dits, and the ongoing délimitation parcellaire process in other appellations progressively identifies and registers site-specific lieux-dits eligible for label mention. EU Delegated Regulation 2019/33 Art. 55 authorizes the label mention of smaller geographic units (including lieux-dits) within a PDO or PGI area provided 100% of grapes originate from the named site.
Flurname
French site name
Lieu-dit-Bezeichnung
Parzellen-Ortsname
cadastral parcel name
lieu-dit designation
INAO (2023) Délimitation parcellaire des appellations d'origine EU Delegated Regulation 2019/33 Art. 55 Pitiot S and Servant JC (2010) Les Vins de Bourgogne 14th Ed, Collection Pierre Poupon
Lieu Dit Name
Light body quality in Mosel Riesling Kabinett (7.5% alcohol, 45 g/L residual sugar, body rating 2.5/10) creates the style's defining refreshment character where delicate weight allows the floral-citrus fruit and mineral notes to express with transparency and lift, enabling consumption of multiple glasses without palate fatigue and pairing with delicate Asian cuisines where fuller wines would overwhelm, demonstrating how light body represents essential style authenticity rather than quality deficiency in this category.
Light body quality is a body quality that represents minimal perceived weight and delicate palate presence, characterizing the ethereal, refreshing mouthfeel typical of low-alcohol wines, cool-climate expressions, and styles emphasizing freshness and drinkability over power and concentration. Within the BFO framework, this quality specifically addresses the light end of the body spectrum, distinguishing delicate weight from medium and full body while recognizing that light body represents a positive quality attribute in appropriate wine styles rather than a deficiency requiring correction. The ontological significance lies in light body defining a wine category valued for refreshment, versatility, and food-pairing flexibility, where the delicate weight enables consumption in larger quantities, pairs with delicate foods without overwhelming them, and provides the crisp, thirst-quenching character appropriate for casual drinking and warm-weather enjoyment. The perceptual characteristics of light body include watery to slightly textured mouthfeel, minimal coating of oral surfaces, easy swallowing without palate fatigue, and the impression of freshness and lift rather than weight and substance. The compositional profile associated with light body typically includes lower alcohol (8-11.5%), minimal residual sugar in dry styles, limited extract, and higher acidity creating the impression of lightness through palate-cleansing effect, with these factors combining to produce the characteristic delicate weight. The variety and regional associations of light body include German and Austrian Riesling (particularly Kabinett and lower Prädikat levels), Muscadet, Vinho Verde, most Prosecco, many rosé wines, and cool-climate Pinot Grigio, with these wines valued specifically for their light-bodied character that would be compromised by greater weight. Understanding light body quality enables appropriate style recognition, quality assessment valuing lightness as positive attribute in suitable contexts, and food pairing recommendations leveraging the versatility of light-bodied wines.
Light_Body_Quality addresses the delicate end of the body spectrum, representing a positive attribute in appropriate wine styles. This quality is valued for refreshment, versatility, and drinkability rather than being viewed as deficiency. The class supports style-appropriate evaluation that recognizes light body as optimal for wines emphasizing freshness and elegance over power and concentration.
delicate body
delikate Textur
ethereal texture
leichter Körper
leichtgewichtig
light weight
refreshing mouthfeel
zartes Mundgefühl
Jackson RS (2014) Wine Science: Principles and Applications, 4th Ed. Academic Press, San Diego, pp. 831-887 Pickering GJ (2000) Low- and reduced-alcohol wine: a review. Journal of Wine Research 11(2): 129-144
Light Body Quality
Light capture efficiency assessment comparing healthy versus iron-deficient Pinot Noir leaves reveals chlorosis impact on light harvesting, with healthy leaves showing chlorophyll content of 450 μmol/m², absorptance of 0.85, and quantum yield of 0.08 mol CO₂/mol photons, while chlorotic leaves display reduced chlorophyll (180 μmol/m²), lower absorptance (0.62), and diminished quantum yield (0.04), demonstrating how nutrient deficiency compromises light capture efficiency with cascading effects on photosynthetic capacity and ultimately fruit quality potential.
Light capture efficiency quality is a photosynthetic capacity quality that represents the effectiveness of the grapevine photosynthetic apparatus in absorbing photosynthetically active radiation (PAR) and transferring excitation energy to reaction centers, characterizing the light-harvesting performance that determines the quantum efficiency of photosynthesis under light-limited conditions. Within the BFO framework, this quality specifically addresses the light acquisition component of photosynthetic capacity, distinguishing light capture from subsequent carbon fixation while recognizing that efficient light harvesting is prerequisite for high photosynthetic rates particularly under limiting light conditions typical of interior canopy positions and cloudy environments. The ontological significance lies in light capture efficiency determining the quantum yield of photosynthesis and the efficiency of carbon gain at low to moderate light intensities where light reactions rather than dark reactions limit overall photosynthetic rate, with implications for canopy design, vineyard row orientation, and management decisions affecting light distribution within the vine canopy. The physiological basis for light capture efficiency includes chlorophyll and carotenoid concentrations (determining total light absorption), chlorophyll a/b ratio (reflecting antenna size versus reaction center proportion), thylakoid membrane organization (affecting energy transfer efficiency), and the ratio of photosystems I and II (determining electron transport balance), with these components integrated in the light-harvesting complexes and photosystems that capture light and initiate electron flow. The environmental and developmental factors affecting light capture efficiency include light environment during leaf development (shade promoting larger antenna systems), nitrogen availability (strongly affecting chlorophyll synthesis), temperature stress (potentially damaging photosynthetic apparatus), and leaf age (progressive decline in pigment content and organization). Understanding light capture efficiency quality enables diagnosis of light harvesting limitations, optimization of canopy architecture for efficient light interception, and assessment of photosynthetic apparatus health as an early indicator of stress or senescence.
Light_Capture_Efficiency_Quality serves as the parent class for pigment and absorptance qualities, representing the light-harvesting component of photosynthesis. This quality determines photosynthetic performance under light-limited conditions and provides early indicators of stress or nutrient deficiency. The class supports photosynthetic apparatus assessment and canopy management targeting efficient light utilization.
Lichtaufnahmeeffizienz
Lichtsammeleffizienz
PAR capture
PAR-Erfassung
Quantenausbeute
light harvesting efficiency
photon capture effectiveness
quantum efficiency
Björkman O, Demmig B (1987) Photon yield of O₂ evolution and chlorophyll fluorescence characteristics at 77 K among vascular plants of diverse origins. Planta 170(4): 489-504 Evans JR (1989) Photosynthesis and nitrogen relationships in leaves of C₃ plants. Oecologia 78(1): 9-19
Light Capture Efficiency Quality
Light compensation point assessment guiding canopy management in dense Pinot Noir reveals that heavily shaded interior leaves (receiving 25 μmol/m²/s PAR) are below their measured compensation point (LCP = 35 μmol/m²/s), indicating these leaves are net carbon consumers that drain rather than contribute to vine productivity, with selective leaf removal eliminating these parasitic leaves and improving vine carbon balance while also enhancing air circulation and reducing disease pressure in the fruit zone, demonstrating how compensation point provides physiological justification for canopy manipulation.
Light compensation point quality is a light response quality that represents the light intensity at which photosynthetic CO₂ fixation exactly balances respiratory CO₂ release, characterizing the minimum light requirement for positive carbon balance below which leaves become net carbon consumers rather than contributors to whole-plant productivity. Within the BFO framework, this quality specifically addresses the compensation dimension of light response, distinguishing the threshold for positive carbon balance from the saturation threshold while recognizing that compensation point determines whether leaves at specific canopy positions contribute to or drain from whole-plant carbon economy. The ontological significance lies in light compensation point defining the threshold for productive leaf function, where leaves receiving light below compensation point consume more carbon through respiration than they fix through photosynthesis, becoming parasitic on the vine's carbon economy and candidates for removal through canopy management to redirect resources to productive leaf area. The physiological basis for light compensation involves the balance between gross photosynthesis (increasing with light) and respiration (relatively constant regardless of light), with compensation point (LCP) occurring where gross photosynthesis equals respiration, typically at low light intensities ranging from 15-60 μmol/m²/s PAR in Vitis vinifera depending on leaf respiration rate and quantum yield. The factors affecting light compensation point include respiration rate (higher respiration increasing LCP), quantum yield (higher efficiency decreasing LCP), temperature (increasing both respiration and LCP), and leaf age (senescing leaves showing elevated LCP from declining photosynthetic capacity with maintained respiration). The canopy management implications of compensation point are direct, as leaves chronically below compensation point should be removed to improve vine carbon balance, with compensation point providing the physiological basis for interior canopy leaf removal that eliminates unproductive or parasitic leaf area. Understanding light compensation point quality enables identification of unproductive canopy positions, evidence-based leaf removal decisions, and assessment of minimum light requirements for positive leaf contribution to vine carbon economy.
Light_Compensation_Point_Quality addresses the minimum light for positive leaf carbon balance, below which leaves become net carbon consumers. This quality provides the physiological basis for leaf removal decisions targeting unproductive canopy areas. The class supports canopy management through identification of light levels insufficient for productive leaf function.
Kompensationsbestrahlung
LCP
LCP-Wert
Lichtkompensationspunkt
compensation irradiance
minimaler Lichtbedarf
minimum light requirement
Boardman NK (1977) Comparative photosynthesis of sun and shade plants. Annual Review of Plant Physiology 28(1): 355-377 Schultz HR (1995) Grape canopy structure, light microclimate and photosynthesis. Australian Journal of Grape and Wine Research 1(1): 2-17
Light Compensation Point Quality
Light penetration quality assessment in Willamette Valley Pinot Noir employed quantum sensors at fruit zone height to characterize seasonal light environment across canopy management treatments. Unthinned, unpositioned control vines showed fruit zone PAR declining from 45% of ambient at bloom to only 8% at véraison as canopy development progressively shaded clusters. Shoot-positioned vines maintained 28% ambient PAR at véraison, while positioned plus leaf-removed treatments achieved optimal light penetration quality at 18% ambient PAR corresponding to 350 μmol m⁻² s⁻¹ at solar noon. Fruit from the optimized light environment showed 40% higher total anthocyanins and 25% lower vegetal character in sensory analysis compared to heavily shaded controls, demonstrating the quality benefits of improved light penetration quality.
Light penetration quality is a canopy porosity quality that inheres in a grapevine canopy system and represents the proportion and spectral characteristics of solar radiation that traverses the foliar mass to reach interior canopy positions including the fruit zone, as determined by canopy architecture, leaf area density, leaf optical properties, and the angular distribution of incident radiation throughout the day and growing season. This quality fundamentally determines the photosynthetic productivity of interior leaves and the light environment experienced by developing fruit clusters, with cascading effects on carbon assimilation, fruit composition, and wine quality potential. The ontological significance of light penetration quality lies in its role as the primary environmental signal driving phenolic biosynthesis, chlorophyll degradation, and flavor precursor development in grape berries, while simultaneously influencing bud fruitfulness for subsequent seasons through light-dependent floral initiation in developing buds. The physics of light penetration through plant canopies involves both direct beam attenuation following Beer-Lambert exponential decay with depth and diffuse radiation scattering that can partially illuminate shaded positions, with total photosynthetically active radiation at any interior point representing the sum of attenuated direct, scattered diffuse, and reflected components. Grapevine leaves absorb approximately 85-90% of incident photosynthetically active radiation while reflecting 5-10% and transmitting only 3-5%, meaning that each leaf layer dramatically reduces light availability to underlying tissues, with interior positions in dense canopies receiving less than 10% of above-canopy radiation. The spectral quality of penetrating light changes with depth as preferential absorption of red and blue wavelengths by chlorophyll enriches transmitted light in far-red and green wavelengths, altering the red to far-red ratio that influences photomorphogenic responses in shaded tissues. Measurement of light penetration quality employs quantum sensors positioned at various canopy depths, line ceptometers integrating across horizontal transects, and hemispherical photography enabling calculation of direct and diffuse site factors from gap fraction analysis. The relationship between light penetration quality and fruit composition reflects established requirements for approximately 50-100 μmol m⁻² s⁻¹ photosynthetically active radiation at fruit clusters to optimize phenolic development without inducing light stress, with this target corresponding to approximately 10-20% of full sunlight in typical growing conditions.
Light_Penetration_Quality is_a Canopy_Porosity_Quality, representing the specific radiation transmission dimension of canopy openness. This class addresses both the quantity and spectral quality of light reaching interior canopy positions. Distinguished from Air_Circulation_Quality and Spray_Penetration_Quality which address other consequences of canopy porosity. Functionally related to Fruit_Exposure_Quality and the phenolic composition aspects of Berry_Quality. The fruit zone PAR target of 10-20% ambient provides quantitative benchmark for this quality.
Lichtdurchlässigkeit
Lichtpenetration
Strahlungsdurchlass
canopy light transmission
fruit zone illumination
light interception
solar penetration
Dokoozlian NK, Kliewer WM (1995) The light environment within grapevine canopies. I. Description and seasonal changes during fruit development. American Journal of Enology and Viticulture 46:209-218 Spayd SE et al. (2002) Separation of sunlight and temperature effects on the composition of Vitis vinifera cv. Merlot berries. American Journal of Enology and Viticulture 53:171-182 Downey MO et al. (2006) Cultural practice and environmental impacts on the flavonoid composition of grapes and wine: a review of recent research. American Journal of Enology and Viticulture 57:257-268
Light Penetration Quality
Light response quality characterization comparing exterior sun leaves versus interior shade leaves in Merlot reveals distinct adaptation patterns, with sun leaves showing Amax of 16.5 μmol CO₂/m²/s, LSP of 1200 μmol/m²/s PAR, LCP of 45 μmol/m²/s, and quantum yield of 0.055, while shade leaves display Amax of 8.5 μmol/m²/s, LSP of 450 μmol/m²/s, LCP of 18 μmol/m²/s, and elevated quantum yield of 0.072, demonstrating the acclimation trade-off where shade leaves sacrifice maximum capacity for enhanced efficiency at low light, with implications for canopy design that matches leaf types to appropriate light zones.
Light response quality is a photosynthetic capacity quality that represents the functional relationship between photosynthetic rate and incident light intensity, characterizing the parameters of the photosynthesis-light response curve that describe photosynthetic performance from compensation through saturation and potential photoinhibition across the range of light environments encountered in vineyard canopies. Within the BFO framework, this quality addresses the light dependency dimension of photosynthetic performance, distinguishing the shape and parameters of the light response from absolute photosynthetic capacity while recognizing that light response characteristics determine how effectively leaves utilize the variable light environments within vine canopies throughout the day and season. The ontological significance lies in light response quality determining the productivity of leaves at different canopy positions and times, where the light saturation point determines the light level above which additional light provides no benefit, the compensation point determines the minimum light for positive carbon balance, and the quantum yield determines efficiency at limiting light, together defining how canopy architecture translates available light into carbon gain. The mathematical description of light response typically employs non-rectangular hyperbola or Michaelis-Menten models characterized by parameters including maximum photosynthesis (Amax), quantum yield (Φ), convexity (θ), light compensation point (LCP), and light saturation point (LSP), with these parameters varying among cultivars, leaf ages, and acclimation states. The environmental and developmental factors affecting light response include growth light environment (sun versus shade leaf development creating distinct response patterns), nitrogen nutrition (affecting both Amax and quantum yield), water status (affecting stomatal limitation across all light levels), and temperature (affecting the shape of the response curve through enzyme kinetics). The canopy implications of light response characteristics are substantial, as the distribution of light response types throughout the canopy determines how effectively the canopy converts intercepted radiation into carbon gain, with optimal canopy architecture matching leaf light response characteristics to local light environments. Understanding light response quality enables photosynthetic performance prediction across light environments, canopy architecture optimization, and interpretation of whole-vine carbon gain from individual leaf characteristics.
Light_Response_Quality serves as the parent class for specific light response parameters (Saturation Point, Compensation Point, Photoinhibition Susceptibility). This quality describes the functional relationship between light and photosynthesis that determines performance across canopy light environments. The class supports canopy architecture optimization through characterization of leaf light response patterns.
Bestrahlungsantwort
Lichtantwort
Lichtkurve
P-I curve
Photosynthese-Licht-Beziehung
irradiance response
light response curve
photosynthesis-light relationship
Thornley JHM (1976) Mathematical Models in Plant Physiology. Academic Press, London, pp. 86-110 Ögren E, Evans JR (1993) Photosynthetic light response curves. Planta 189(2): 182-190
Light Response Quality
Light saturation point assessment informing canopy management in Cabernet Sauvignon reveals that sun leaves (LSP 1100 μmol/m²/s) are light-saturated by mid-morning on clear days when PAR exceeds 1400 μmol/m²/s, receiving excess light that cannot increase photosynthesis but may cause photoinhibition, suggesting that the dense canopy architecture delivering only 800 μmol/m²/s to these leaves through partial shading would maintain maximum photosynthesis while reducing photoinhibition risk, demonstrating how saturation point knowledge enables evidence-based canopy density decisions.
Light saturation point quality is a light response quality that represents the light intensity at which photosynthesis reaches its maximum rate and above which additional light provides no further increase in carbon fixation, characterizing the threshold beyond which photosynthetic capacity rather than light availability limits productivity. Within the BFO framework, this quality specifically addresses the saturation dimension of light response, distinguishing the light level for maximum photosynthesis from the maximum rate itself while recognizing that the saturation point indicates the transition from light-limited to capacity-limited photosynthesis with implications for optimal canopy light distribution. The ontological significance lies in light saturation point determining the light level required for maximum leaf productivity, where leaves receiving light above saturation gain no additional carbon while potentially suffering photoinhibition, making saturation point essential for understanding optimal canopy architecture that delivers saturating but not excessive light to productive leaf area. The physiological basis for light saturation involves the balance between light-driven electron transport and the capacity of downstream processes (Rubisco carboxylation, Calvin cycle regeneration) to utilize the ATP and NADPH produced, with saturation occurring when carbon fixation capacity limits overall rate regardless of additional light availability. The quantitative ranges for light saturation point in Vitis vinifera typically span 600-1500 μmol/m²/s PAR depending on leaf acclimation (sun leaves saturating at higher light than shade leaves), nitrogen nutrition (affecting Rubisco content and thus capacity), and temperature (affecting enzyme kinetics), with full sunlight (approximately 2000 μmol/m²/s) substantially exceeding saturation for most grapevine leaves. The acclimation of saturation point shows sun-developed leaves achieving higher saturation points through greater investment in Rubisco and electron transport capacity, while shade-developed leaves show lower saturation points but enhanced efficiency at low light, representing alternative strategies for different light environments. Understanding light saturation point quality enables canopy design optimization, assessment of whether light or capacity limits photosynthesis at specific canopy positions, and interpretation of diminishing returns from increased light interception beyond canopy saturation.
Light_Saturation_Point_Quality addresses the light level for maximum photosynthesis, above which additional light provides no carbon gain benefit. This quality determines optimal canopy light delivery and has implications for photoinhibition risk assessment. The class supports canopy architecture optimization through understanding of leaf light requirements for maximum productivity.
Isat
LSP
LSP-Wert
Lichtsättigungspunkt
Sättigungsbestrahlung
maximaler Lichtbedarf
maximum light requirement
saturation irradiance
Ögren E, Evans JR (1993) Photosynthetic light response curves. Planta 189(2): 182-190 Schultz HR (1996) Leaf absorptance of visible radiation in Vitis vinifera L. Estimates of age and shade effects with a simple field method. Scientia Horticulturae 66(1-2): 93-102
Light Saturation Point Quality
Lignification typically initiates at basal internodes when shoots reach 40-60 cm length, progressing acropetally at rates of 1-2 nodes per week under optimal conditions. Cabernet Sauvignon achieves 85-95% cane lignification by leaf fall in temperate climates, while Grenache in warm Mediterranean conditions may reach only 70-80% due to extended vegetative growth. Lignin content in mature canes ranges from 18-24% of dry weight depending on cultivar.
A grapevine disposition that is the inherited capacity of a Vitis vinifera plant to deposit lignin polymers in secondary cell walls of shoot tissues, transforming green herbaceous shoots into woody canes through progressive sclerenchyma development and vascular tissue maturation.
Lignification_Disposition serves as the parent class for shoot maturation traits critical to winter survival and carbohydrate reserve storage. The process involves coordinated expression of phenylpropanoid pathway genes including PAL, C4H, 4CL, and CAD. Lignification is promoted by moderate water stress, reduced nitrogen availability, and shorter photoperiods, while excessive vigor delays the process.
Lignifizierungsdisposition
Verholzungsvermögen
cane woodiness trait
shoot hardening disposition
shoot lignification capacity
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 68-75 Bouquet A, Danglot Y (1996) Inheritance of several traits in grapevine. Vitis 35:89-94
Lignification Disposition
A lignified one-year cane on Vitis vinifera cv. Cabernet Sauvignon examined during January pruning, exhibiting complete brown coloration from base to tip indicating full lignification, firmness when pressed between fingers showing woody rigidity, clean snap rather than bending when excessive force applied indicating mature secondary cell walls, cross-section revealing well-developed periderm, continuous vascular cambium, and dense xylem tissue with minimal pith, this cane showing cold hardiness to -18°C based on differential thermal analysis and suitable for retention as 12-node fruiting cane in balanced pruning system.
A one-year cane that has completed the lignification process throughout its length, characterized by woody tissue formation in cell walls through deposition of lignin polymers, brown coloration indicating mature secondary cell wall development, structural rigidity sufficient to support shoot and crop loads, and protection against winter cold damage through reduced tissue water content and increased cellular organization. Lignified one-year canes represent physiologically mature fruiting wood suitable for retention during winter pruning, with completed lignification indicating successful transition from active growth to dormancy, adequate carbohydrate accumulation for winter survival and spring budbreak, and appropriate tissue development for supporting next season's crop production without structural failure or physiological stress.
Lignified one-year canes are distinguished from incompletely lignified canes by uniform brown color (no green remaining), woody firmness throughout length, and clean breaking pattern when snapped. Complete lignification is essential for winter survival in cold climates, as unlignified tissues contain higher water content and less organized cellular structure, making them vulnerable to freeze damage. The extent and timing of lignification varies among cultivars (early versus late-maturing varieties), environmental conditions (warm climates showing earlier lignification than cool regions), and vine management (balanced vines lignifying more completely than excessively vigorous vines). Pruning decisions should prioritize well-lignified canes, removing incompletely lignified tips and avoiding retention of semi-lignified or green wood as fruiting positions.
ausgereifte Rute
hardened cane
lignifizierte Rute
mature lignified cane
verholzte Rute
woody cane
Based on Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 245-267 Fennell (2004) "Freezing tolerance and injury in grapevines" Journal of Crop Improvement 10=201-235.
Lignified One Year Cane
Cabernet Sauvignon grafted on 1103 Paulsen rootstock at 16 years showing lignified woody secondary root, histochemical staining (phloroglucinol-HCl) revealing extensive lignin deposition in secondary xylem fiber and vessel walls with intense red coloration indicating high lignin content, woody root fiber cells showing thick secondary walls (4-6 micrometers) with narrow lumens, chemical analysis indicating lignin content 24% of dry weight, lignified structure providing high mechanical strength and rigidity supporting root system architecture, lignin resistance to degradation contributing to woody root persistence and structural stability
Lignified woody secondary root is woody secondary root emphasizing lignin deposition and lignification process in cell walls creating rigid strong woody structure, characterized by extensive lignin accumulation in secondary xylem fiber and vessel cell walls providing mechanical strength and rigidity, lignin content typically 20-30% of dry weight in woody root tissue reflecting substantial lignification, lignin-cellulose-hemicellulose matrix creating composite structure with high tensile strength and compression resistance, and progressive lignification increasing as secondary xylem matures with newly formed cells initially showing limited lignin that increases over weeks to months after cell differentiation, with lignified woody secondary root structure providing mechanical support enabling root to resist bending, tension, and compression forces, protecting against pathogen degradation through lignin's resistance to microbial and enzymatic breakdown, and demonstrating key role of lignin polymerization in creating wood structure that enables perennial woody architecture distinguishing grapevine from herbaceous plants. Lignified woody secondary root lignin composition includes= containing lignin polymers (complex phenolic polymers formed from monolignol precursors including coniferyl alcohol, sinapyl alcohol, and p-coumaryl alcohol), showing G and S lignin types (guaiacyl lignin from coniferyl alcohol and syringyl lignin from sinapyl alcohol comprising majority of root lignin), exhibiting H lignin component (p-hydroxyphenyl lignin from p-coumaryl alcohol present in smaller amounts), demonstrating lignin-carbohydrate linkages (covalent bonds between lignin and hemicellulose creating integrated matrix), and revealing cultivar variation (rootstock varieties potentially differing in lignin content and composition affecting wood properties). Lignified woody secondary root lignification process includes= initiating after cell expansion (lignin deposition beginning after xylem cells complete elongation and expansion), showing monolignol synthesis (phenylpropanoid pathway in cambial cells producing lignin precursor molecules), demonstrating monolignol transport (precursors moving from synthesis sites to cell walls where polymerization occurs), exhibiting enzymatic polymerization (peroxidases and laccases catalyzing radical coupling reactions creating lignin polymer), and progressing from lumen to middle lamella (lignification typically beginning at cell lumen and progressing outward through wall layers). Lignified woody secondary root mechanical properties includes= providing high tensile strength (lignified cell walls resisting pulling forces through strong covalent bonds in lignin-cellulose matrix), exhibiting compression resistance (lignin rigidity preventing cell wall collapse under compression loads), demonstrating bending stiffness (lignified woody tissue resisting deformation when bent enabling structural support), showing limited flexibility (high lignin content reducing flexibility compared to non-lignified tissues creating rigid structure), and revealing load-bearing capacity (lignified woody secondary roots supporting substantial mechanical loads from above-ground structure and soil forces). Lignified woody secondary root in xylem function includes= preventing vessel collapse (lignin in vessel walls maintaining lumen patency under negative pressure during water transport), enabling long-distance transport (lignified vessels conducting water efficiently through rigid tubes that don't compress), providing tracheary element strength (lignin essential for development of functional xylem vessels and tracheids), demonstrating hydraulic-mechanical trade-off (extensive lignification providing strength but potentially limiting vessel diameter and conductivity), and showing embolism resistance (lignified vessel walls helping resist cavitation and embolism formation under water stress). Lignified woody secondary root decay resistance includes= resisting microbial degradation (lignin highly resistant to enzymatic and microbial breakdown compared to cellulose and hemicellulose), providing long-term persistence (lignified woody roots potentially remaining structurally intact in soil for years after vine death), demonstrating antimicrobial properties (lignin and associated phenolic compounds potentially inhibiting pathogen growth), showing selective degradation (white-rot fungi capable of lignin breakdown while brown-rot fungi attack polysaccharides leaving lignin), and enabling structural longevity (lignin protection contributing to woody secondary root persistence for decades in living vine). Lignified woody secondary root fiber characteristics includes= showing thick secondary walls (fiber cells with extensive secondary wall development up to several micrometers thickness), exhibiting high lignin content (fibers typically showing 25-35% lignin higher than vessels reflecting extensive lignification), demonstrating narrow lumens (thick walls reducing lumen size in mature fibers creating dense structure), providing mechanical reinforcement (lignified fibers creating strengthening tissue surrounding vessels in woody xylem), and revealing functional specialization (fibers primarily structural rather than conducting distinguishing from vessels). Lignified woody secondary root and carbon sequestration includes= representing long-term carbon storage (lignin stable molecule potentially persisting in soil for years to decades after root death), containing substantial carbon (lignin approximately 60-65% carbon by weight contributing to total root system carbon content), demonstrating slow turnover (lignified woody secondary roots showing much slower carbon turnover than non-woody fine roots), contributing to soil organic matter (lignified root residues providing recalcitrant carbon input to soil carbon pools), and showing climate significance (woody root lignin accumulation representing mechanism for long-term carbon sequestration relevant to climate change mitigation). Lignified woody secondary root developmental progression includes= showing increasing lignification with age (young woody secondary roots with less lignin progressively accumulating more as mature), exhibiting radial gradient (younger outer xylem showing less lignin than older inner xylem reflecting progressive lignification), demonstrating seasonal variation (lignification intensity potentially varying between earlywood and latewood cells), revealing maturation process (newly formed xylem cells completing lignification over weeks to months after initial differentiation), and displaying depth effects (lignification potentially varying with depth reflecting different environmental conditions though data limited).
Lignified_Woody_Secondary_Root emphasizes lignin deposition and lignification process and is distinguished from unlignified fine roots by focus on lignin accumulation. Critical for understanding wood formation, mechanical strength, cell wall lignification, decay resistance, structural rigidity, and the lignin polymer deposition creating rigid strong woody structure enabling perennial root architecture.
Highly lignified root
Lignifizierte holzige Sekundärwurzel
Lignin-deposited secondary root
Lignin-rich woody root
Ligninreiche Wurzel
Stark verholzte Wurzel
Lignified Woody Secondary Root
The lime content data for a Pouilly-Fuissé vineyard (Mâconnais, Burgundy) on Jurassic limestone documents total calcium carbonate of 35% and active lime of 12% at 0-30 cm depth — the active lime percentage of 12% is below the chlorosis tolerance threshold of 17% for the commonly used 3309C rootstock (V. riparia × V. rupestris) but above the threshold of 6% for the sensitive SO4 rootstock, requiring careful rootstock matching to the site-specific active lime level. The vineyard was planted on 161-49C rootstock (V. berlandieri × V. riparia, chlorosis tolerance index 25% active lime), providing a comfortable safety margin against lime-induced chlorosis while maintaining the moderate vigor appropriate for this site. The lime content data also informs soil preparation decisions — the 35% total CaCO₃ renders phosphorus fertilization partially ineffective (applied phosphorus rapidly precipitates as insoluble calcium phosphate in the alkaline soil environment), necessitating either localized phosphorus placement in the root zone or foliar phosphorus application to supplement soil supply.
A soil chemistry data component that records the concentration of calcium carbonate (CaCO₃) equivalents in the vineyard soil, determined by acid neutralization (Scheibler calcimeter or Bernard calcimeter measuring CO₂ evolved upon HCl addition) and expressed as percent total calcium carbonate and percent active lime (the fine calcium carbonate fraction <50 μm that is chemically reactive and directly responsible for the alkaline pH and iron-immobilizing capacity), with lime content being the soil chemical property most critical for rootstock selection — the active lime percentage determines the risk of lime-induced iron chlorosis that is the most agronomically damaging nutritional disorder on calcareous vineyard soils.
Lime_Content_Data documents the soil carbonate parameter that is the single most consequential soil chemistry measurement for vineyard establishment decisions — the active lime percentage determines the risk of lime-induced iron chlorosis (also called calcareous chlorosis or lime chlorosis), a condition in which the high bicarbonate concentration in calcareous soil solution immobilizes iron within the vine by inhibiting the reduction of ferric (Fe³⁺) to ferrous (Fe²⁺) iron required for iron mobilization from root to shoot, producing the characteristic interveinal yellowing of young leaves that can reduce photosynthetic capacity by 50-80% and render affected vines commercially unproductive. The distinction between total lime and active lime is critical for chlorosis risk assessment — total lime includes both coarse calcium carbonate fragments (>50 μm) that dissolve slowly and contribute minimally to the soil solution bicarbonate concentration, and fine calcium carbonate particles (80) indicate high chlorosis risk regardless of rootstock, moderate IPC values (30-80) require lime-tolerant rootstocks, and low IPC values (<30) are manageable with standard rootstocks.
Calciumcarbonat-Gehalt
Kalkboden-Charakterisierung
Kalkgehalt-Daten
calcareous soil characterization
calcium carbonate content
soil carbonate measurement
Bavaresco L et al. (2003) Role of rootstock on iron-chlorosis induced by bicarbonate in nutrient solution, Acta Horticulturae 603: 87-95 Juste C and Pouget R (1972) Appréciation du pouvoir chlorosant des sols par un nouvel indice faisant intervenir le fer facilement extractible, Comptes Rendus de l'Académie d'Agriculture 58: 352-357 White RE (2003) Soils for Fine Wines, Oxford University Press Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia
Lime Content Data
Lime induced chlorosis quality assessment in Cognac Ugni Blanc vineyards on Campanian chalk documented the relationship between soil carbonate status, rootstock selection, and chlorosis severity to guide replanting decisions on this highly calcareous terroir. Soil characterization established lime induced chlorosis quality risk with active lime content ranging from 8% to 35% across the vineyard, soil pH from 7.8 to 8.6, and calculated IPC (Indice de Pouvoir Chlorosant) from 25 to 85, with values above 60 indicating high chlorosis risk requiring lime-tolerant rootstocks. Existing plantings showed lime induced chlorosis quality severity directly correlated with rootstock lime tolerance, with vines on SO4 rootstock (moderate tolerance, IPC threshold ~40) showing severe chlorosis (mean score 4.2) and 45% yield reduction in high-IPC zones, while adjacent vines on 41B rootstock (high tolerance, IPC threshold ~80) showed only mild symptoms (score 1.5) and maintained 90% of potential yield. Tissue iron analysis revealed the lime induced chlorosis quality paradox with severely chlorotic SO4 leaves containing 72 ppm total Fe (above deficiency threshold) but only 18 ppm "active iron" (HCl-extractable fraction representing physiologically available Fe) versus 52 ppm active Fe in healthy 41B leaves, demonstrating internal iron immobilization rather than uptake limitation in the susceptible rootstock. The lime induced chlorosis quality assessment informed replanting recommendations specifying 41B or Fercal rootstocks for zones with IPC above 50, 140Ru or 333EM for moderate zones (IPC 30-50), and permitting SO4 only in low-IPC areas below 30, enabling terroir-appropriate rootstock selection to prevent chlorosis in the replanted vineyard.
Lime induced chlorosis quality is an iron deficiency symptom quality that inheres in grapevine organisms growing on calcareous soils and represents the specific form of iron deficiency caused by high soil pH and bicarbonate ion concentration interfering with iron uptake and internal utilization, producing the characteristic interveinal chlorosis on young leaves that is the primary symptom of iron limitation in viticultural regions with limestone-derived soils including many of the world's most prestigious wine appellations. This quality captures the most prevalent form of iron deficiency in commercial viticulture, recognizing that lime-induced chlorosis affects extensive vineyard areas and represents a major factor in rootstock selection, site evaluation, and nutritional management on calcareous terroirs. The ontological significance of lime induced chlorosis quality lies in its identification of the specific mechanism of iron limitation distinguishing it from iron deficiency on acidic soils, with the bicarbonate-mediated interference requiring different management approaches than simple iron unavailability, particularly the selection of lime-tolerant rootstocks capable of acquiring and utilizing iron under high-pH, high-bicarbonate conditions. The biological basis of lime induced chlorosis encompasses both rhizosphere and plant-internal effects of bicarbonate ions, with high bicarbonate reducing iron solubility in the soil solution, interfering with root iron reductase activity required for iron uptake, and potentially causing internal iron precipitation in leaves where bicarbonate accumulation can immobilize iron even after absorption, creating the "iron chlorosis paradox" where total leaf iron may be normal while physiologically active iron is insufficient. The soil parameters predicting lime induced chlorosis risk include active lime content (calcium carbonate equivalent that reacts with dilute acid), soil pH, and the chlorosis potential index (IPC, Indice de Pouvoir Chlorosant) that integrates these factors to predict chlorosis severity, enabling site assessment and rootstock selection before planting. The rootstock variation in lime tolerance reflects genetic differences in root iron acquisition capacity, with rootstocks derived from Vitis berlandieri showing superior lime tolerance attributed to enhanced proton and reductant extrusion that maintains iron availability in the rhizosphere, providing the primary management tool for lime-induced chlorosis prevention.
Lime_Induced_Chlorosis_Quality is_a Iron_Deficiency_Symptom_Quality, representing the specific Fe deficiency form caused by high soil pH and bicarbonate interference with Fe uptake and utilization. This class addresses the most common Fe deficiency mechanism in viticulture. Active lime content and IPC (Indice de Pouvoir Chlorosant) predict chlorosis risk. The "iron chlorosis paradox" where total leaf Fe may be adequate but active Fe insufficient. Rootstock lime tolerance (41B, Fercal > 140Ru > SO4 > 3309C) is primary management strategy. Berlandieri parentage confers superior lime tolerance through enhanced rhizosphere acidification.
Bikarbonat-induzierte Chlorose
Kalkchlorose
Kalkstein-Chlorose
bicarbonate-induced chlorosis
calcareous soil chlorosis
high-pH iron deficiency
kalkinduzierte Chlorose
limestone chlorosis
Bavaresco L et al. (2010) Grapevine iron nutrition: rootstock effects on scion response to lime-induced chlorosis. Acta Horticulturae 868:115-122 Mengel K (1994) Iron availability in plant tissues — iron chlorosis on calcareous soils. Plant and Soil 165:275-283 Tagliavini M, Rombolà AD (2001) Iron deficiency and chlorosis in orchard and vineyard ecosystems. European Journal of Agronomy 15:71-92
Lime Induced Chlorosis Quality
Comprehensive evaluation of Lime_Tolerance_Disposition at INRAE Bordeaux assessed growth, leaf chlorophyll content, and iron nutrition across 25 rootstocks grown in calcareous soil with 40% total CaCO₃ and 18% active lime over three growing seasons. Measurements included SPAD chlorophyll index, leaf iron concentration (total and "active" Fe²⁺), visual chlorosis severity (0-5 scale), and shoot dry weight. Lime_Tolerance_Disposition varied dramatically among rootstocks. '41B' (V. vinifera × V. berlandieri) exhibited exceptional tolerance with SPAD values of 38 (healthy green), leaf total Fe of 125 μg/g DW with 45% active Fe, chlorosis score of 0.5, and 95% shoot growth relative to reference soil. 'Fercal' (BC1 × 333EM) showed similar tolerance (SPAD 36, 92% growth). In contrast, '3309C' (V. riparia × V. rupestris) displayed severe lime sensitivity with SPAD of 18 (pronounced yellowing), total Fe of 180 μg/g DW but only 15% active Fe (indicating Fe inactivation), chlorosis score of 4.2, and growth reduced to 35% of reference. 'SO4' showed intermediate tolerance (SPAD 28, 65% growth). The paradox of high total but low active iron in sensitive rootstocks reflects the "iron chlorosis paradox" where Fe accumulates in inactive Fe³⁺ form in chlorotic leaves. Active lime tolerance thresholds (IPC values) were established as 40% for '41B', 40% for 'Fercal', 17% for '3309C', and 25% for 'SO4', providing practical guidance for rootstock selection based on soil analysis.
A Lime_Tolerance_Disposition is an Abiotic_Stress_Tolerance_Disposition that inheres in a Vitis plant and is realized through physiological mechanisms that enable maintenance of adequate iron nutrition, normal chlorophyll synthesis, and healthy growth on calcareous (lime-rich) soils where high pH and bicarbonate concentrations would otherwise induce iron deficiency chlorosis. This disposition represents the capacity to tolerate active lime (calcium carbonate) in soil, critically important for viticulture as many of the world's finest wine regions are located on limestone-derived soils, including Burgundy, Champagne, Chablis, parts of Bordeaux, and much of the Mediterranean basin. Lime-induced chlorosis results from impaired iron availability and utilization under high soil pH (> 7.5) and elevated bicarbonate (HCO₃⁻) concentrations, causing interveinal yellowing of young leaves, reduced photosynthesis, stunted growth, and in severe cases vine decline and death. The primary mechanisms of lime tolerance involve enhanced iron acquisition from calcareous soil (through rhizosphere acidification, increased ferric reductase activity, and efficient iron chelate uptake), tolerance of bicarbonate effects on root iron metabolism, and adaptation to consistently high rhizosphere pH. In grafted grapevines, Lime_Tolerance_Disposition is primarily determined by rootstock genotype, making rootstock selection the essential management tool for calcareous sites. Rootstock lime tolerance is commonly quantified by the "Index de Pouvoir Chlorosant" (IPC) or "Active Lime Tolerance" threshold, representing the maximum soil active lime content (% CaCO₃ extractable in ammonium oxalate) compatible with healthy growth.
Lime_Tolerance_Disposition is_a Abiotic_Stress_Tolerance_Disposition. Lime tolerance is primarily determined by rootstock genotype in grafted vines, making rootstock selection essential for calcareous sites. The primary problem is iron deficiency chlorosis despite often adequate total soil iron. Child classes address mechanisms including Iron_Acquisition_Efficiency_Disposition, Bicarbonate_Tolerance_Disposition, and High_pH_Adaptation_Disposition. V. berlandieri and V. vinifera × V. berlandieri hybrids show superior lime tolerance, reflecting adaptation to limestone soils of origin. IPC (Index de Pouvoir Chlorosant) or active lime tolerance % provides practical rootstock selection criteria.
Chlorose-Resistenz
Kalkstein-Toleranz
Kalktoleranz-Disposition
calcareous soil tolerance
limestone tolerance
Bavaresco L et al. (2003) Grapevine iron nutrition and chlorosis, Plant and Soil 249:263-270 Mengel K (1994) Iron availability in plant tissues—iron chlorosis on calcareous soils, Plant and Soil 165:275-283 Tagliavini M, Rombolà AD (2001) Iron deficiency and chlorosis in orchard and vineyard ecosystems, European Journal of Agronomy 15:71-92
Lime Tolerance Disposition
41B Millardet et de Grasset (V. vinifera Chasselas × V. berlandieri) tolerates up to 40% active lime, supporting Chardonnay on the chalk soils of Champagne (active lime 25-35%) and the Kimmeridgian marls of Chablis without chlorosis symptoms. Fercal (BC1 of 31R × Berlandieri Colombard 1B) shows the highest lime tolerance of commercial rootstocks (up to 60% active lime), enabling Ugni Blanc cultivation on the highly calcareous Campanian soils of Cognac. 140 Ruggeri (V. berlandieri × V. rupestris) tolerates 20-25% active lime, supporting Catarratto on the calcareous clay soils of western Sicily where sensitive rootstocks like Riparia Gloire show severe interveinal chlorosis.
An abiotic stress-tolerant rootstock role that inheres in a grafted rootstock by virtue of its capacity to maintain scion productivity on calcareous soils with high calcium carbonate (CaCO₃) content, preventing or mitigating lime-induced iron chlorosis. Lime-tolerant rootstocks overcome the iron deficiency caused by high soil pH (typically 7.5-8.5) and bicarbonate ion (HCO₃⁻) accumulation in the rhizosphere, which reduce iron solubility and inhibit Fe³⁺ reduction and uptake by roots. Tolerance mechanisms include enhanced Fe³⁺-chelate reductase activity in root plasma membranes, increased proton efflux capacity to acidify the rhizosphere, efficient iron translocation via citrate and nicotianamine chelation in xylem sap, and metabolic tolerance to low tissue iron through optimized iron use efficiency. Lime tolerance is quantified by the Index of Chlorotic Power (ICP) or Index of Active Limestone (active CaCO₃ %) tolerated without chlorosis symptoms, with tolerant rootstocks functioning at 20-40% active lime versus 10-15% for sensitive genotypes.
Lime_Tolerant_Rootstock_Role is subclassed by tolerance magnitude into High_Lime_Tolerance, Moderate_Lime_Tolerance, and Low_Lime_Tolerance categories. V. berlandieri parentage is the primary genetic source of lime tolerance, as this species evolved on the Edwards Plateau limestone formations of central Texas. The V. vinifera component in 41B and Fercal contributes to their exceptional lime tolerance but also reduces phylloxera resistance, limiting their use to regions with low phylloxera pressure. Iron chlorosis severity is modified by rootstock-scion interactions, soil moisture status, and root zone temperature.
Kalkunterlage
calcareous soil tolerant rootstock
chloroseresistente Unterlage
chlorosis resistant rootstock
kalktolerante Unterlage
kalkverträgliche Unterlage
lime tolerant rootstock
limestone tolerant rootstock
Bavaresco L et al. (2003) Grapevine iron nutrition update, Journal International des Sciences de la Vigne et du Vin 37: 89-100 Tagliavini M, Rombolà AD (2001) Iron deficiency and chlorosis in orchard and vineyard ecosystems, European Journal of Agronomy 15: 71-92 Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22
Lime Tolerant Rootstock Role
Börner (V. riparia × V. cinerea) demonstrates limited compatibility, forming excellent unions with most cultivars but showing reduced affinity with certain V. vinifera genotypes including some Pinot clones, requiring grafting trials before large-scale adoption with untested scions. Dog Ridge (V. champinii) exhibits limited compatibility with some wine grape cultivars, forming strong unions with Thompson Seedless and Flame Seedless table grapes but showing variable success with certain wine cultivars in California trials. VR 043-43 (V. vinifera × M. rotundifolia BC1) shows limited compatibility as expected from Muscadinia introgression, requiring careful scion selection and modified grafting techniques. ARG1 (Aramon × V. rupestris Ganzin No. 1) historically showed limited compatibility and declining union integrity with some cultivars, contributing to its abandonment alongside phylloxera susceptibility.
A graft affinity rootstock role that inheres in a grafted rootstock by virtue of its restricted capacity to form successful graft unions, showing compatibility with some Vitis vinifera cultivars or hybrid scions while exhibiting incompatibility symptoms with others. Limited compatibility rootstocks display cultivar-specific variation in graft success rates, union strength, vascular connectivity, and long-term durability, necessitating careful scion-rootstock matching and pre-planting compatibility assessment. Incompatibility symptoms may include poor callus formation at the graft interface, delayed or incomplete vascular differentiation, mechanical weakness at the union (breakage under wind or crop load stress), restricted translocation of water, nutrients, and assimilates across the graft junction, and elevated incidence of delayed incompatibility. Limited compatibility arises from genetic distance between rootstock and certain scion genotypes, biochemical incompatibility factors (phenolic compound accumulation, peroxidase activity differences), or developmental asynchrony between grafting partners. Limited compatibility rootstocks may offer unique pest resistance or stress tolerance traits that justify their use despite compatibility constraints.
Limited_Compatibility_Rootstock_Role captures rootstocks requiring cultivar-specific compatibility evaluation before deployment, typically those involving more distant genetic backgrounds (V. cinerea, V. champinii) or Muscadinia introgression. Limited compatibility does not preclude commercial use but necessitates additional nursery testing and restricts the range of suitable scion cultivars. Breeding programs developing novel pest-resistant rootstocks (nematode resistance from V. champinii, Xiphinema resistance from M. rotundifolia) must address compatibility limitations through selection and backcrossing. Compatibility can sometimes be improved through intermediate grafts (double-working) or modified grafting techniques, though these add complexity and cost.
beschränkt verträgliche Unterlage
cultivar-specific compatibility rootstock
eingeschränkt kompatible Unterlage
limited compatibility rootstock
restricted compatibility rootstock
selective compatibility rootstock
sortenspezifisch kompatible Unterlage
Pina A, Errea P (2005) A review of new advances in mechanism of graft compatibility-incompatibility, Scientia Horticulturae 106: 1-11 Walker MA et al. (2014) Rootstock breeding for resistance to nematodes and phylloxera, Acta Horticulturae 1045: 89-97 Bouquet A et al. (2000) Grapevine rootstocks with resistance to Xiphinema index, Acta Horticulturae 528: 521-528 Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22
Limited Compatibility Rootstock Role
Characterization of Limited_Root_System_Disposition at the University of California Davis documented root systems of rootstocks recognized for compact root development. Reference rootstocks '101-14 Mgt', '3309C', 'Riparia Gloire', and '420A' were excavated at 8 years from deep alluvial soils, with complete root recovery and morphological analysis. Limited_Root_System_Disposition rootstocks showed consistent distinguishing characteristics. '101-14 Mgt' (V. riparia × V. rupestris) exhibited archetypal limited root development with total root length of 1,180 m, root dry weight of 1.6 kg, maximum rooting depth of 1.3 m with only 12% of root length below 1 m, and horizontal spread of 1.4 m radius. Root architecture showed horizontal orientation with shallow primary roots and limited vertical penetration. 'Riparia Gloire' displayed even more limited characteristics (950 m, 1.2 kg, 1.1 m depth) reflecting the shallow-rooting nature of V. riparia. '3309C' showed intermediate limited development (1,420 m, 2.1 kg, 1.6 m depth). Fine root fraction was notably higher in Limited_Root_System_Disposition rootstocks (72-78% versus 62-68% for extensive systems), indicating compensatory investment in absorptive capacity within constrained total root mass. Vigor assessment confirmed that Limited_Root_System_Disposition rootstocks reduced scion pruning weight by 25-40% compared to extensive rootstocks when grafted with identical scion ('Cabernet Sauvignon') under equivalent conditions, demonstrating the functional vigor-limiting effect of compact root systems. However, drought sensitivity was increased, with limited root system vines reaching stress thresholds (Ψpd < -0.5 MPa) 12-18 days earlier than extensive root system vines during dry-down experiments.
A Limited_Root_System_Disposition is a Root_Vigor_Disposition that inheres in a Vitis rootstock and is realized through restrained root growth producing a compact, less extensive root system with moderate total root length, shallower penetration, and concentrated rather than expansive soil exploration, resulting in reduced capacity for water and nutrient acquisition compared to extensive root systems. This disposition represents the lower range of root vigor, characteristic of rootstocks that develop relatively modest below-ground structures, naturally constraining scion vigor through limited resource supply capacity. Limited root systems typically show total root lengths below 1,500 m per mature vine, maximum rooting depths of 1-2 m, root length densities below 2 cm/cm³, and root dry weights of 1.5-2.5 kg. Such root systems are advantageous for vigor control (naturally limiting scion growth on fertile sites), dense planting (reduced root competition zone), and situations requiring scion vigor reduction. However, Limited_Root_System_Disposition may increase drought sensitivity (smaller water reservoir), reduce nutrient buffering capacity, and limit vine recovery from stress or damage due to smaller reserve storage. Rootstocks with limited root systems are typically recommended for fertile soils, well-watered sites, vigorous scions requiring control, and high-density plantings where per-vine root volume is intentionally restricted.
Limited_Root_System_Disposition is_a Root_Vigor_Disposition representing low root vigor. Typical characteristics include total root length < 1,500 m, dry weight < 2.5 kg, rooting depth < 2 m, root length density < 2 cm/cm³. Examples include '101-14 Mgt', '3309C', 'Riparia Gloire', '420A'. V. riparia parentage typically confers limited rooting. Advantages include vigor control and suitability for fertile sites and high-density planting. Disadvantages include increased drought sensitivity and reduced stress buffering. Higher fine root fraction partially compensates for reduced total root mass.
Begrenztes-Wurzelsystem-Disposition
Eingeschränkte-Bewurzelung
Kompaktes-Wurzelsystem
compact root system
restricted rooting
small root system
Swanepoel JJ, Southey JM (1989) The influence of rootstock on the rooting pattern of the grapevine, South African Journal of Enology and Viticulture 10:23-28 Tandonnet JP et al. (2010) Long term effects of rootstock on vine performance and wine quality for Vitis vinifera cv. Cabernet Sauvignon, Australian Journal of Grape and Wine Research 16:298-306 Tramontini S et al. (2013) Rootstock control of scion response to water stress in grapevine, Environmental and Experimental Botany 93:20-26
Limited Root System Disposition
Limpidity assessment in white wine quality control examines each tank against standardized light source, with limpid wines (turbidity below 2 NTU) approved for bottling while wines showing visible haze (above 5 NTU) require additional fining or filtration before release, with one lot showing subtle protein haze (8 NTU) treated with bentonite addition (0.5 g/L) and re-evaluated after settling to confirm limpidity achievement before proceeding to bottling, demonstrating limpidity as the operational quality gate for commercial wine finishing.
Limpidity quality is a clarity quality that represents the absence of visible suspended particles or haze when wine is examined against a light source, characterizing the basic transparency that indicates successful removal or settling of particulate matter through fining, filtration, or natural sedimentation. Within the BFO framework, this quality specifically addresses the particle-free dimension of clarity, distinguishing simple limpidity (absence of visible turbidity) from brilliance (active light reflection creating sparkle), while representing the minimum clarity standard for conventionally finished wines. The ontological significance lies in limpidity providing the baseline clarity criterion that separates finished wines from those still requiring treatment or deliberately produced in unfiltered style, where limpid appearance indicates that suspended particles have been removed below the threshold of visual detection (approximately 1-5 NTU depending on viewing conditions), qualifying wine for standard commercial release. The assessment methodology for limpidity involves viewing wine against a light source (candle, lamp, or daylight) with the glass tilted to maximize path length, where limpid wine shows clear transmission without visible particles, streaks, or haziness, while non-limpid wine displays turbidity ranging from slight veil to dense cloudiness depending on particle concentration. The compositional factors affecting limpidity include protein stability (unstable proteins precipitating with temperature changes or tannin contact), microbiological stability (yeast or bacterial cells remaining in suspension), colloidal stability (aggregation of polysaccharides, proteins, and tannins forming visible haze), and tartrate stability (potassium bitartrate crystallization creating temporary or permanent haze), with each instability type requiring specific treatment for limpidity achievement. The distinction between limpidity and brilliance recognizes that wines may be limpid (particle-free) without being brilliant (actively reflective), with limpidity representing necessary but not sufficient condition for highest clarity grades. Understanding limpidity quality enables basic clarity assessment, stability evaluation identifying wines at risk of haze development, and quality control ensuring minimum clarity standards for commercial release.
Limpidity_Quality addresses the basic particle-free dimension of clarity, representing minimum clarity standard for conventionally finished wines. This quality is distinguished from Brilliance_Quality which requires active light reflection beyond simple particle absence. The class supports quality control through turbidity measurement (NTU) and visual assessment, with limpidity serving as the baseline clarity criterion below which wines require treatment or stylistic justification for turbidity.
Grundklarheit
Partikelfreiheit
Trübungsfreiheit
basic transparency
clear appearance
haze-free quality
particle-free clarity
Zoecklein BW et al. (1995) Wine Analysis and Production. Chapman & Hall, New York, pp. 380-425 Boulton RB et al. (1999) Principles and Practices of Winemaking. Springer, New York, pp. 492-528
Limpidity Quality
Linalool concentration in grape berries ranges from 10-50 µg/L in non-aromatic cultivars to 500-2500 µg/L in highly aromatic Muscats, with the sensory threshold of 15-25 µg/L explaining the characteristic floral notes in aromatic wines. Muscat Blanc à Petits Grains shows the highest linalool levels (1500-2500 µg/L free + bound), followed by Muscat of Alexandria (1000-2000 µg/L) and Gewürztraminer (400-1200 µg/L). VvTPS-lin (linalool synthase) converts GPP to linalool with high specificity, with gene expression increasing 10-50 fold during ripening in aromatic cultivars. The linalool/geraniol ratio varies among Muscat types, with Muscat Blanc showing linalool dominance (ratio 2-4) and some Muscats showing more balanced profiles (ratio 0.8-1.5). Linalool undergoes acid-catalyzed rearrangement to α-terpineol and limonene during winemaking and aging.
A monoterpene accumulation disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate linalool (3,7-dimethyl-1,6-octadien-3-ol) as a predominant monoterpene alcohol in grape berries, with genotype-specific concentrations determining the intensity of floral (lily-of-the-valley, lavender) aroma character and serving as the primary contributor to Muscat aroma.
Linalool is the signature compound of Muscat aroma and contributes floral character across aromatic wine styles. The compound exists in both (R)-(-)- and (S)-(+)-enantiomeric forms, with grapevine producing predominantly the (S)-(+)-linalool enantiomer (lavender character) versus the (R)-(-)- form (woody character) produced by some other plants. Linalool concentration peaks at or slightly before optimal harvest maturity, with potential decline through volatilization and oxidation if harvest is delayed. The glycosylated pool provides a reservoir for continued linalool release during wine aging.
Linalool-Disposition
Linaloolakkumulation
Linaloolgehalt
floral terpene trait
linalool content disposition
linalool synthesis capacity
Mateo JJ, Jiménez M (2000) Monoterpenes in grape juice and wines. Journal of Chromatography A 881:557-567 Fenoll J et al. (2009) Changes in the free and bound volatile monoterpene content of Vitis vinifera cv. Muscat of Alexandria throughout grape berry development. Food Chemistry 117:383-388
Linalool Accumulation Disposition
Linalool content quality in Muscat Blanc à Petits Grains reaches 1500-2500 μg/L free linalool at optimal harvest, approximately 50-100 times the sensory threshold, producing the intensely floral, orange-blossom character that defines this variety's aromatic identity winemakers producing dry Muscat styles (Alsace Muscat, dry Australian Muscat) target harvest when free linalool peaks but bound forms remain high, capturing both immediate aromatic impact and aging potential, while sweet Muscat styles (Muscat de Beaumes-de-Venise) accept the linalool decline of later harvest as sugar concentration takes priority.
Linalool_Content_Quality represents the chemical quality characterizing the concentration of linalool (3,7-dimethyl-1,6-octadien-3-ol) in grape berries and wine, capturing the acyclic monoterpene alcohol responsible for the distinctive floral-citrus aroma that dominates Muscat character. Linalool exists as two enantiomers, (R)-(−)-linalool with a woody-lavender character and (S)-(+)-linalool with a sweet, floral-citrus character grape-derived linalool is predominantly the (S)-(+) form contributing the fresh floral notes. As the most abundant free monoterpene in many aromatic cultivars, linalool typically comprises 30-60% of total free monoterpenes, with concentrations ranging from 10-100 μg/L in neutral varieties through 200-500 μg/L in semi-aromatic varieties to 500-2000 μg/L in intensely aromatic Muscat cultivars. The sensory threshold of linalool in wine matrices is approximately 15-25 μg/L, well below typical concentrations in aromatic varieties, ensuring perceptible contribution to wine aroma. Linalool accumulates primarily in grape skins during ripening, with glycosylated forms (linalyl glucoside, linalyl rutinoside) substantially exceeding free linalool concentration. The free linalool concentration peaks around optimal harvest maturity then declines in overripe fruit due to oxidation to linalool oxides and isomerization to other terpenes. During wine aging, free linalool declines through acid-catalyzed reactions that form cyclic oxides (furanoid and pyranoid linalool oxides) and rearrange to α-terpineol, with these transformation products contributing their own aromatic characters. The linalool concentration trajectory during berry development and wine aging provides insight into optimal harvest timing and wine evolution potential.
Linalool_Content_Quality addresses the dominant monoterpene in many aromatic cultivars, serving as the primary marker for Muscat character. The transformation of linalool during aging (oxidation to oxides, isomerization to α-terpineol) means that linalool concentration in young wine does not directly predict linalool contribution to aged wine aroma. The chirality of linalool affects aroma quality, with grape-derived (S)-(+)-linalool providing the desirable floral character. Linalool quantification employs GC-MS or GC-FID with standard addition or internal standard calibration, following appropriate extraction methodology.
3,7-dimethylocta-1,6-dien-3-ol concentration
Linalool-Konzentration
Linalylverbindungsgehalt
floral terpene alcohol quality
floraler Terpenalkoholgehalt
linalool level
linalyl compound content
Ribéreau-Gayon P et al. (1975) Recherches Technologiques sur les Composés Terpéniques des Raisins et des Vins. Connaissance de la Vigne et du Vin 9(3): 181-199 Black CA et al. (2015) Terpenoids and Their Role in Wine Flavour: Recent Advances. Australian Journal of Grape and Wine Research 21(S1): 582-600
Linalool Content Quality
Ampelographic characterization of Grüner Veltliner requires standardized linear dimension measurements including mature leaf blade length (140-180 mm from petiole junction to apex), blade width (130-170 mm at widest point), petiole length (relative to middle vein, typically 0.9-1.1 ratio), and depth of upper lateral sinuses (medium to deep, 30-50% of half-blade length), all assessed on leaves from the middle third of fruiting shoots at véraison.
Linear_Dimension_Quality encompasses all one-dimensional spatial measurements that characterize the extent of grapevine organs along specified axes. Linear dimensions constitute the most fundamental and frequently measured size parameters in ampelography and viticultural research because they can be rapidly assessed with simple instruments and provide direct input for calculating derived metrics including area, volume, and shape indices. This category comprises four primary measurement types: Length_Quality capturing the greatest longitudinal extent Width_Quality representing the greatest transverse extent perpendicular to the length axis Diameter_Quality for circular or near-circular cross-sections and Thickness_Quality addressing the smallest dimension, typically perpendicular to the primary surface plane. Linear dimensions exhibit characteristic scaling relationships across organs and developmental stages. Shoot internode length increases acropetally until the vegetative-reproductive transition, then decreases. Berry diameter follows a double-sigmoid growth curve with distinct phases of cell division, lag, and cell expansion. Leaf dimensions scale with position on the shoot, light environment, and vine water status. The precision of linear dimension measurements depends on instrument resolution, measurement protocol standardization, and the regularity of the organ being measured. Digital calipers provide 0.01 mm resolution suitable for most grapevine applications. For irregular organs, multiple measurements at defined positions enable calculation of mean values and shape descriptors. Linear dimensions serve as primary data for allometric models that predict physiological function from easily measured parameters.
Linear_Dimension_Quality unifies length-type measurements under a common parent while distinguishing specific measurement orientations through its child classes. The class aligns with PATO_0001708 (1-D extent) as an equivalent upper-level quality type. Measurement protocols must specify the axis of measurement, reference landmarks, developmental stage, and organ position for reproducibility. Where OIV descriptors employ categorical scales, these can be linked to quantitative ranges through descriptor-specific calibration data.
Linearmaßqualität
Längendimensionsqualität
eindimensionale Messqualität
length-type quality
linear extent quality
linear metric quality
lineare Ausdehnungsqualität
one-dimensional measurement quality
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 47-72 Maul E et al. (2012) Vitis International Variety Catalogue. Julius Kühn-Institut, Federal Research Centre for Cultivated Plants
Linear Dimension Quality
Chardonnay liquid pulp tissue at maturity showing high juice yield characteristics= water content 77-80%, vacuoles 91-94% of cell volume containing aqueous solution with 21-24% sugars and 0.7-0.95% acids, yielding 74-78 L juice per 100 kg berries (high yield important for white wine production economics), with 66-72% free-run juice from readily-disrupting liquid pulp cells plus 28-34% press juice, creating total must volume determining wine production volume from harvest tonnage. Thompson Seedless table grape liquid pulp tissue for fresh consumption= water 80-84% (very high creating very juicy eating experience), cells 160-195 μm diameter with large vacuoles 92-95% of volume, readily releasing liquid contents when biting creating perception of high juiciness important for table grape eating quality, with liquid pulp providing pleasant juice release during chewing creating satisfying fresh flavor delivery. Cabernet Sauvignon liquid pulp tissue juice yield= 78-81% water content yielding 72-76 L juice per 100 kg berries (moderate wine cultivar yield), with free-run juice 64-70% of total (released by crushing) used for premium wine production, press juice 30-36% (requiring 0.8-2 bar pressure) potentially used for secondary wine or discarded depending on quality, demonstrating liquid pulp as primary juice source with yield determining wine production volume and economics. Merlot liquid pulp tissue water accumulation during ripening= veraison (BBCH 81, week 10) water content 72-76%, juice yield potential 68-72 L/100 kg mid-ripening (week 14) water 76-79%, yield 72-75 L/100 kg maturity (week 18) water 78-82%, yield 74-77 L/100 kg, demonstrating progressive water accumulation during 8-week ripening period increasing juice yield 8-12% from veraison to maturity through osmotic import into expanding liquid pulp cells. Syrah liquid pulp tissue harvest timing effects on juice yield= early harvest (week 16, under-ripe 22% sugars) water 75-78%, yield 71-74 L/100 kg optimal harvest (week 18, 25% sugars) water 78-81%, yield 74-77 L/100 kg late harvest (week 20, over-ripe 28% sugars) water 76-79%, yield 72-76 L/100 kg (declining from peak due to late-season dehydration), demonstrating that maximum juice yield occurs at optimal maturity with delayed harvest potentially reducing yield through water loss despite continued sugar accumulation creating concentrated composition but reduced juice volume.
A liquid pulp tissue is pulp tissue emphasizing high liquid content and juice-yielding capacity, characterized by very high water content (76-84%), large vacuoles (90-95% of cell volume) filled with aqueous solution, thin walls (0.4-0.8 μm) that readily rupture releasing liquid contents, and composition creating juice yield of 70-80 L per 100 kg berries representing the primary source of grape juice for wine, juice products, or fresh consumption. Liquid pulp tissue juice characteristics= (1) high water content 76-84% providing substantial liquid volume with pulp tissue representing 78-88% of berry weight yielding 70-80 L juice per 100 kg berries (cultivar and ripeness dependent) (2) aqueous vacuolar solution containing dissolved solutes creating juice composition of 18-28% sugars, 0.4-0.9% organic acids, minerals, creating must for wine fermentation or juice for fresh consumption (3) ready release from cells when walls disrupted through crushing, pressing, or chewing (4) free-flowing liquid characteristics when released creating pleasant juicy mouthfeel for eating or efficient processing for juice/wine production. Liquid pulp tissue cellular basis of juice yield= (1) vacuoles occupying 90-95% of cell volume maximizing liquid storage capacity per cell (2) vacuolar contents predominantly water (typically 75-85% of vacuolar mass) with dissolved solutes creating juice when released (3) thin walls readily rupturing under moderate compression (crushing at 0.2-0.5 bar pressure, pressing at 0.5-2 bar) releasing vacuolar contents (4) minimal water binding to cellular structures allowing free water to flow from disrupted cells (5) large cell size (110-180 μm) creating high liquid volume per cell with fewer cell walls per unit volume reducing solid material in juice. Liquid pulp tissue juice extraction= (1) free-run juice (60-72% of total yield) released by crushing without pressing from most readily-disrupting liquid pulp cells (2) press juice (28-40% of total) requiring pressure to rupture more resistant cells and extract liquid from compressed cell debris (3) total juice yield 70-80 L per 100 kg berries varying with cultivar (table cultivars often higher yield 75-80 L, wine cultivars 70-76 L), ripeness (riper berries higher water content and yield), and processing conditions (harder pressing increasing yield but potentially extracting undesirable compounds). Liquid pulp tissue composition determining juice quality= (1) sugar content 18-28% in juice determining potential alcohol for wine (rule of thumb= % sugar ÷ 1.7 ≈ % alcohol potential, so 24% sugar → 14% alcohol potential) or sweetness for juice products (2) acid content 0.4-0.9% (4-9 g/L titratable acidity) determining wine acidity, pH, and stability (3) sugar=acid ratio determining balance with ratio 20-45=1 typical for wine grapes depending on style (4) mineral content particularly potassium (1200-2200 mg/kg) affecting wine pH and stability. Liquid pulp tissue developmental water accumulation= early development (Phase I) showing moderate water content 65-75% creating low juice yield ripening (Phase II) showing progressive water accumulation increasing to 76-84% through osmotic import driven by sugar accumulation in vacuoles, creating high juice yield at maturity potential late-season dehydration in warm dry conditions reducing water content and juice yield through transpiration exceeding import, creating concentrated composition but reduced juice volume.
Liquid_Pulp_Tissue emphasizes the high water content and juice-yielding capacity of pulp tissue and is distinguished from Soft_Pulp_Tissue (emphasizing texture) and Storage_Pulp_Tissue (emphasizing storage function). Understanding liquid pulp tissue is essential for juice yield prediction, for irrigation management affecting water accumulation and yield, for harvest timing optimizing yield while achieving compositional maturity, and for processing protocol design maximizing juice extraction efficiency.
Saftreiches Fruchtfleisch
flüssiges Pulpa-Gewebe
juicy flesh
liquid-rich pulp
saftreiches Fruchtmark
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 185-192 Coombe, B.G., McCarthy, M.G. (2000). Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6=131-135.
Liquid Pulp Tissue
Little leaf quality assessment for zinc deficiency in Fresno County Flame Seedless documented the characteristic symptom features enabling rapid field diagnosis in this table grape cultivar where foliar appearance affects marketability assessment. Visual examination of symptomatic shoots revealed little leaf quality with apical leaves measuring 4-6 cm blade length versus 12-15 cm on normal shoots, representing 60-70% reduction in linear dimension and approximately 80% reduction in leaf area. Morphological assessment of little leaf quality identified the characteristic associated features including petiole length of 2.5 cm versus 6 cm normal (58% reduction, disproportionate to blade reduction), asymmetric blade development with one lobe averaging 2.8 cm and the other 4.2 cm on the same leaf (50% asymmetry), and marginal serration spacing of 8 mm versus 3 mm in normal leaves creating the diagnostic coarse sawtooth margin. Leaf texture assessment noted little leaf quality association with puckered or crinkled blade surface, attributed to irregular cell expansion under zinc limitation. The developmental distribution of little leaf quality showed severity gradient from shoot tip basipetally, with the youngest emerged leaf showing maximum expression (leaf area index 0.2 relative to normal), progressive improvement through positions 2-5 (indices 0.35, 0.55, 0.75, 0.90), and normal leaf development below position 6 where leaves had completed expansion before zinc became limiting. Tissue analysis confirmed little leaf quality diagnosis with symptomatic shoot tip Zn at 9 ppm versus 32 ppm in normal shoots from adjacent vines, establishing the causal relationship between zinc status and little leaf expression.
Little leaf quality in zinc deficiency is a zinc deficiency symptom quality that inheres in young grapevine leaves and represents the pathognomonic symptom of abnormally small, malformed leaves resulting from zinc's essential role in auxin biosynthesis and cell expansion, with affected leaves showing reduced blade area (often 25-50% of normal), shortened petioles, asymmetric blade development with unequal lobes, abnormally wide spacing between marginal serrations, and often a crinkled or puckered texture, creating a distinctive appearance that enables confident field diagnosis of zinc deficiency. This quality captures the most diagnostically reliable zinc deficiency symptom, recognizing that little leaf is virtually specific to zinc limitation and rarely produced by other nutritional or environmental conditions, providing a visual indicator of high diagnostic value. The ontological significance of little leaf quality in zinc deficiency lies in its diagnostic specificity enabling field identification of zinc limitation without tissue analysis, though confirmation through testing is recommended before corrective application, and in its indication of auxin metabolism disruption that also underlies the fruit set problems characteristic of zinc deficiency. The biological basis of little leaf encompasses zinc's essential role in tryptophan synthase required for tryptophan production as the precursor for auxin biosynthesis, with zinc deficiency reducing auxin availability and thereby limiting cell expansion during leaf development, producing the characteristic small leaf size while cell number may be relatively normal. The morphological features of little leaf beyond reduced size include the asymmetric blade development where one side of the leaf may be substantially larger than the other, the shortened petiole disproportionate even to the reduced blade size, and the widely spaced marginal teeth that give the leaf margin a coarse "sawtooth" appearance distinct from normal fine serrations. The distribution of little leaf symptoms on shoot tips where the youngest, most actively expanding leaves are affected confirms the immobile nutrient pattern, with older leaves that completed expansion before zinc became limiting showing normal size and morphology.
Little_Leaf_Quality is_a Zinc_Deficiency_Symptom_Quality, representing the pathognomonic small malformed leaf symptom from Zn-limited auxin biosynthesis. This class addresses the most diagnostically reliable Zn deficiency symptom. Blade area reduction of 60-80%, shortened petioles, asymmetric lobes, and wide serration spacing are characteristic. The auxin deficiency mechanism (tryptophan synthase requires Zn) explains both leaf and fruit set symptoms. Severity gradient from shoot tip basipetally reflects immobile nutrient pattern. Virtually pathognomonic for Zn deficiency, rarely caused by other conditions.
Kleinblättrigkeit
Zink-Kleinblatt
Zn-induced leaf dwarfing
Zn-induzierte Blattverzwergung
Zwergblatt-Syndrom
small leaf syndrome
stunted leaf development
zinc little leaf
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Swietlik D (1999) Zinc nutrition in horticultural crops. Horticultural Reviews 23:109-178 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication
Little Leaf Quality
Chardonnay grafted on Teleki 5C rootstock at 16 years showing living inner rootstock bark 2.1 mm thickness, cell viability staining confirming living status (FDA fluorescent staining showing bright fluorescence indicating intact membranes and active metabolism), microscopic examination revealing intact protoplasts in all phloem cells with visible nuclei and organelles, respiration measurements showing active oxygen consumption and CO2 production in inner bark tissue, sieve elements conducting photosynthates demonstrated through aphid stylet technique, companion cells showing dense cytoplasm indicating high metabolic activity, phloem parenchyma containing starch reserves (iodine staining positive), living inner bark contrasting with dead outer bark (negative viability staining)
Living inner rootstock bark is inner rootstock bark emphasizing viability and active physiological function of cells, characterized by living phloem cells with intact protoplasts, functional membranes maintaining cellular compartmentation, active metabolism including respiration and biosynthesis, responsive capacity to environmental and hormonal signals, showing sieve elements conducting photosynthates through pressure flow, companion cells supporting sieve element function metabolically, phloem parenchyma storing carbohydrate reserves and synthesizing compounds, demonstrating wound healing capacity through callus formation and new cell production, and representing viable functional tissue contrasting with dead outer bark requiring continuous replacement as living inner bark progressively isolated by cork formation ages and eventually dies becoming incorporated into outer bark. Living inner rootstock bark cellular viability includes= containing intact protoplasts (living cells with functional cytoplasm, nucleus, and organelles), showing functional membranes (plasma membrane and organellar membranes maintaining compartmentation and selective permeability), exhibiting active nuclei (nuclei with intact chromatin directing cellular functions through gene expression), demonstrating living organelles (mitochondria respiring, plastids storing, ribosomes synthesizing proteins), and displaying membrane potential (electrical potential across membranes indicating active ion transport and energy state). Living inner rootstock bark active metabolism includes= showing aerobic respiration (consuming oxygen and producing CO2 generating ATP for cellular processes), exhibiting protein synthesis (producing enzymes, structural proteins, and regulatory proteins required for function), demonstrating active transport (moving ions and molecules across membranes against gradients using ATP), synthesizing compounds (producing callose, P-proteins, secondary metabolites, and other substances), and maintaining energy status (high ATP/ADP ratio indicating active metabolism and functional capacity). Living inner rootstock bark transport functions includes= conducting photosynthates actively (sieve elements moving sucrose and other compounds through pressure flow mechanism), demonstrating bidirectional flow (capability for both basipetal and acropetal transport depending on source-sink relationships), showing responsive transport (flow rates adjusting to demand, storage status, and environmental conditions), maintaining sieve element function (companion cells supporting sieve elements metabolically compensating for their reduced organelle content), and enabling critical nutrient distribution (phloem transport essential for delivering photosynthates to roots, storage organs, and developing tissues). Living inner rootstock bark storage and mobilization includes= accumulating reserves dynamically (phloem parenchyma actively storing carbohydrates during favorable periods), showing seasonal storage cycles (reserves accumulating in fall mobilizing in spring reflecting active metabolic regulation), responding to stress (mobilizing reserves to support recovery from damage, disease, or environmental challenges), demonstrating enzymatic control (living cells producing enzymes catalyzing starch synthesis and degradation), and enabling rapid mobilization (living cells capable of quickly converting stored starch to mobile sugars when needed). Living inner rootstock bark wound response includes= producing callus tissue (living parenchyma cells dividing and dedifferentiating to form wound healing tissue), synthesizing defensive compounds (producing phytoalexins, phenolics, and other antimicrobial substances in response to damage or infection), forming reaction zones (living cells near wounds producing barriers through chemical and anatomical modifications), demonstrating active defense (living tissue mounting defensive responses unlike passive protection of dead outer bark), and showing healing capacity (living cells capable of producing new tissues covering wounds and restoring bark integrity). Living inner rootstock bark responsive capacity includes= responding to hormonal signals (detecting and responding to auxin, cytokinin, and other hormones regulating growth and development), detecting environmental cues (sensing temperature, photoperiod, water status, and other factors), showing adaptive responses (adjusting metabolism, transport, and function in response to changing conditions), demonstrating communication (living cells exchanging signals through plasmodesmata and diffusible compounds), and exhibiting developmental plasticity (living tissue capable of dedifferentiation, redifferentiation, and forming new cell types in response to signals). Living inner rootstock bark lifespan limitations includes= showing finite functional life (individual living phloem cells typically functioning 1-3 years before senescence), demonstrating progressive isolation (cork cambium formation progressively cutting off older phloem from cambium eventually causing death), exhibiting aging changes (living cells gradually declining in function as isolated from cambium and resources), requiring continuous replacement (vascular cambium continuously producing new living phloem maintaining functional tissue despite older phloem death), and transitioning to outer bark (living inner bark eventually dying and becoming incorporated into outer dead bark as natural progression).
Living_Inner_Rootstock_Bark emphasizes cellular viability and active physiology and is distinguished from Inner_Rootstock_Bark (structural emphasis) by focus on living status and metabolic activity. Critical for understanding phloem viability, active transport, metabolic processes, wound healing, responsive capacity, and the living functional tissues conducting food, storing reserves, and responding to environmental challenges.
Functional living phloem
Lebende innere Unterlagenrinde
Metabolically active bark
Metabolisch aktives Phloem
Viable Rinde
Viable inner bark
Living Inner Rootstock Bark
Cabernet Sauvignon living inner trunk bark at 16 years post-planting shows 2.8-4.2 mm depth with all cells maintaining active protoplasm, functional mitochondria consuming 0.8-1.6 μmol O₂/g fresh weight/hour supporting ATP production, sieve tubes actively transporting photosynthate 0.4-0.9 m/hour powered by living companion cell loading, phloem parenchyma actively synthesizing and storing starch (accumulating 24-36 g starch at 9-12% dry weight during post-harvest period, mobilizing reserves during spring budbreak), living cells responding to auxin signals from developing buds triggering spring activation, and maintaining viability through -8°C winter minimum through cold hardening and cryoprotectant accumulation
Living inner trunk bark is inner trunk bark emphasizing the presence of metabolically active cells with intact protoplasm, functioning organelles, and ongoing cellular processes, representing the vitally alive tissue zone essential for phloem transport, storage, and response functions contrasting sharply with dead outer bark. Living inner trunk bark cellular viability characteristics include= intact cell membranes (functional plasma membranes and organelle membranes maintaining compartmentation and selective permeability), functional organelles (mitochondria producing ATP through respiration, endoplasmic reticulum synthesizing proteins and lipids, Golgi apparatus processing and packaging compounds, plastids in parenchyma storing starch or containing chlorophyll in outer cells), active nuclei in nucleated cells (companion cells, parenchyma, and ray cells maintaining genetic material and directing cellular activities, though sieve tube elements enucleate at maturity), living cytoplasm (protoplasm streaming and maintaining dynamic cellular organization), and cellular respiration (consuming oxygen and producing carbon dioxide, ATP generation supporting all cellular processes). Living inner trunk bark metabolic activities include= active transport processes (ATP-powered pumps moving sugars, ions, and other solutes across membranes maintaining concentration gradients), enzyme function (thousands of enzymes catalyzing biochemical reactions including sugar metabolism, starch synthesis/hydrolysis, protein synthesis, lipid metabolism), biosynthesis (producing new proteins, membranes, cell wall components, secondary metabolites), catabolism (breaking down stored compounds mobilizing energy and building blocks), gene expression (transcribing DNA to RNA, translating RNA to proteins, regulating cellular activities), and signal transduction (perceiving and responding to hormonal, environmental, and developmental signals). Living inner trunk bark transport functions dependent on life include= active phloem loading (companion cells using ATP to concentrate sugars into sieve tubes against concentration gradients, loading rates 10-50 μmol/m²/s), sieve tube pressure maintenance (living cells maintaining turgor pressure driving mass flow of phloem sap, typical sieve tube pressure 0.4-1.2 MPa), active unloading at sinks (companion cells and parenchyma using energy to remove sugars from sieve tubes delivering to storage or growing tissues), lateral transport through rays (living ray parenchyma actively transporting water and solutes between xylem and phloem), and bidirectional nutrient distribution (living cells controlling loading and unloading determining nutrient allocation patterns). Living inner trunk bark storage functions requiring life include= starch synthesis (living parenchyma cells using photosynthate to synthesize starch granules, enzyme-mediated polymerization of glucose, accumulating 5-12% dry weight starch in phloem parenchyma), starch mobilization (living cells hydrolyzing starch to sugars during spring mobilization, enzyme-catalyzed breakdown requiring metabolic control), protein storage (living cells synthesizing storage proteins during fall preparing for dormancy, mobilizing proteins during spring growth), seasonal storage regulation (living cells responding to hormonal and environmental signals controlling accumulation versus mobilization), and storage capacity maintenance (living cells maintaining viable storage parenchyma throughout inner bark depth). Living inner trunk bark response capabilities include= hormone perception (living cells containing auxin, cytokinin, abscisic acid, ethylene, gibberellin receptors responding to systemic signals), wound response (living cells detecting damage, proliferating to form callus tissue, differentiating to heal wounds), pathogen defense (living cells synthesizing antimicrobial compounds, phytoalexins, PR proteins in response to infection), environmental stress response (living cells adjusting metabolism to drought, cold, heat stress, producing protective compounds), and developmental regulation (living cells responding to seasonal cues controlling dormancy, growth resumption, senescence). Living inner trunk bark seasonal viability maintenance includes= spring metabolic activation (March-April, increasing respiration rates, enzyme activation, resuming active transport as temperatures rise and hormones signal growth), summer peak activity (May-August, maximum metabolic rates supporting intensive transport and storage functions), fall preparation for dormancy (September-November, adjusting metabolism, synthesizing cryoprotectants, accumulating reserves), winter survival (December-February, reduced but maintained metabolism, cold hardiness mechanisms preventing freeze damage to living cells, maintaining cellular integrity through dormancy), and spring reactivation (resuming full metabolic activity enabling immediate transport and storage function supporting budbreak). Living inner trunk bark vulnerability to death includes= freeze injury (ice crystal formation disrupting membranes and organelles, severe cold killing cells, critical temperatures -15°C to -25°C depending on hardiness status), heat damage (excessive temperatures >45-50°C denaturing proteins and disrupting membranes causing cell death), desiccation (water loss below critical thresholds causing irreversible cellular damage and death), pathogen-induced death (disease organisms producing toxins or degrading cell structures killing living tissue), and mechanical trauma (crushing, cutting, or tearing disrupting cells causing localized death requiring wound healing). Living inner trunk bark death consequences include= immediate transport dysfunction (dead phloem unable to conduct photosynthate, localized death causing transport impediment, circumferential death causing complete blockage and vine death), loss of storage capacity (dead parenchyma unable to accumulate or mobilize reserves, reducing vine stress tolerance and vigor), inability to respond to stimuli (dead tissue unable to mount wound healing, pathogen defense, or stress responses), tissue deterioration (dead cells undergoing autolysis and decay, potentially providing substrate for pathogens), and permanent damage (dead inner bark unable to regenerate requiring new phloem production from cambium to restore function). Living inner trunk bark contrast with dead outer bark includes= living inner bark all cells metabolically active versus outer bark all cells dead, inner bark maintaining respiration and ATP production versus outer bark zero metabolism, inner bark containing functional organelles versus outer bark hollow degraded cells, inner bark actively transporting and storing versus outer bark passive barrier function, inner bark requiring continuous protection versus outer bark serving as protective shield, and sharp transition at cork cambium (living cells internal, dead cells external creating abrupt boundary). Living inner trunk bark assessment includes= vital staining (dyes like neutral red accumulating in living cells revealing viable tissue zones), respiration measurement (oxygen consumption and carbon dioxide production quantifying metabolic activity), electrical conductivity (living tissue showing ion transport and membrane potentials, dead tissue showing no electrical activity), enzyme activity assays (detecting functional enzymes in tissue extracts indicating cellular life), visual observation (living inner bark appearing cream-tan with moist texture, dead tissue appearing brown-grey with dry brittle texture), and sap exudation (living phloem releasing clear sap when cut, dead tissue showing no flow). Living inner trunk bark significance includes= absolutely essential for vine survival (living inner bark providing only mechanism for basipetal transport, vine death within weeks if circumferentially killed), determining stress tolerance (living inner bark storage and transport capacity affecting drought, cold, and nutrient stress resilience), enabling wound healing (living inner bark cells proliferating and differentiating to repair damage), supporting productivity (living inner bark distributing photosynthate to fruit, roots, and storage maintaining growth and yield), and requiring continuous protection (living tissue vulnerable necessitating intact outer bark, careful management preventing inner bark damage).
Living_Inner_Trunk_Bark emphasizes cellular viability and metabolic activity and is distinguished from Inner_Trunk_Bark (anatomical zone), Phloem_Inner_Trunk_Bark (tissue type emphasis), and Dead_Outer_Trunk_Bark (non-living contrast) by focus on life processes. Critical for understanding active transport mechanisms, storage physiology, stress response capabilities, seasonal metabolic dynamics, vulnerability to lethal stress, and absolute requirement for vine survival emphasizing protection importance.
Lebende innere Rinde
Living tissue bark
Stoffwechselaktive Bastzone
Viable phloem
Vitales Phloem
Living Inner Trunk Bark
The lobe depth description of Cabernet Sauvignon documents very deep upper lateral sinuses (OIV 070:9) incised approximately 70-80% of the distance from the blade margin to the midvein — this extreme dissection, combined with the five-lobed configuration, produces the highly distinctive leaf silhouette in which the five lobes appear nearly separated from each other, connected only by narrow tissue bridges at the sinus bases. This very deep lobation is one of Cabernet Sauvignon's most instantly recognizable leaf features, separating it from the moderately deep-lobed Cabernet Franc (OIV 070:7, sinuses extending approximately 50-60% to the midvein), the moderately lobed Merlot (OIV 070:5, sinuses extending approximately 40-50%), and the shallowly lobed Sémillon (OIV 070:3, sinuses extending only 20-30%), creating a lobation gradient that distinguishes these four Bordeaux cultivars at a glance.
A mature leaf description component that records the depth of incision of the lateral sinuses relative to the half-blade width, characterizing the degree of leaf dissection from shallow lobation (sinuses barely penetrating the blade margin) through moderate lobation to deep lobation (sinuses extending more than half the distance to the midvein), scored according to OIV descriptor 070 (depth of upper lateral sinuses) on a scale from very shallow (state 1) to very deep (state 9), providing the critical complementary character to lobe number that together defines the overall dissection architecture of the leaf blade.
Lobe_Depth_Description addresses the character that, together with lobe number, most strongly determines the visual gestalt of the grapevine leaf — the same five-lobed outline appears fundamentally different when the lobes are shallowly indicated (creating a gently undulating margin) versus when they are deeply incised (creating a dramatically dissected, almost palmate silhouette). The depth of sinus incision is scored at the upper lateral sinuses (the deepest pair in most V. vinifera cultivars), using the ratio of sinus depth to half-blade width as the measurement criterion rather than absolute depth in millimeters — this proportional scoring eliminates the confounding effect of overall leaf size and provides a more genetically stable character: a small-leaved Pinot Noir and a large-leaved Pinot Noir will show the same proportional sinus depth despite different absolute measurements. The genetic control of sinus depth overlaps partially with that of lobe number (both reflecting the developmental patterning of marginal growth zones) but is independently variable — cultivars with the same lobe number can differ substantially in sinus depth, and this independent variation provides additional cultivar discrimination within the major lobe-number groups.
Blattdissektionsgrad
Einschnitttiefe-der-Buchten
Lappentiefe-Beschreibung
degree of leaf dissection
lobe prominence record
sinus incision depth
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Chitwood DH et al. (2014) A modern ampelography, Plant Physiology 164: 259-272
Lobe Depth Description
The cultivar with prime name Chasselas carries the local dialect name Fendant in the Valaisan dialect of francophone Switzerland (from the Arpitan/Franco-Provençal verb fendre, to split, describing the berry's tendency to crack when ripe), Gutedel in Alemannic German dialect of Baden and Basel-Landschaft, Krachgutedel in Swabian dialect (krach- referencing the crunching sound when eating the berry), and Csaszla in the Hungarian dialectal adaptation — each name derived from local linguistic resources inaccessible through standard language etymology alone.
A regional name that reflects the phonological, morphological, or lexical characteristics of a local language variety, dialect, or minority language, representing an indigenous linguistic form of cultivar designation that may be incomprehensible or unrecognizable to speakers of the standard national language or other regional dialects.
Local_Dialect_Name preserves the linguistic diversity embedded in traditional viticultural nomenclature, documenting cultivar names that exist in regional language varieties often themselves endangered or declining. These names are of dual scientific interest: for viticulture they encode local cultivar knowledge and usage patterns, and for historical linguistics they provide evidence of dialect distribution, contact phenomena, and lexical innovation in agricultural vocabulary. The documentation of dialect names is increasingly urgent as agricultural modernization and demographic change erode the oral traditions through which these names were transmitted across generations.
Dialektname
Mundart-Sortenname
dialect cultivar name
dialektale Rebbezeichnung
patois grape name
vernacular variety name
Rézeau P (2014) Dictionnaire des noms de cépages de France, CNRS Éditions Kremer D (2007) Etymologisches Wörterbuch der Rebsortennamen, Stuttgart Robinson J et al. (2012) Wine Grapes, Allen Lane pp: 1-18
Local Dialect Name
Localized incompatibility in grapevine manifests as anatomical abnormalities confined to the graft union, including incomplete callus fusion, discontinuous cambium (present in only 40-70% of interface circumference versus >90% in compatible unions), vascular element misalignment, necrotic pockets within the callus bridge, and bark discontinuity visible as a persistent graft line. Mechanically, localized incompatibility causes union weakness with break strength below 150 N versus 300-500 N in compatible unions of similar age, predisposing vines to wind breakage at the graft zone. Hydraulic consequences include reduced conductance across the union (40-70% of ungrafted values) without systemic scion dysfunction. Localized incompatibility may improve with bridge grafting or inarching techniques that bypass the defective union.
A graft incompatibility disposition that is the inherited capacity of certain Vitis vinifera graft combinations to exhibit incompatibility symptoms restricted to the immediate graft zone, resulting from impaired cellular recognition, adhesion failure, aberrant vascular differentiation, or localized necrotic reactions at the interface without systemic toxic effects.
Localized incompatibility results from failure of the cellular and developmental processes required for union formation rather than from toxic factor translocation. The mechanism may involve cell recognition failures (preventing callus intermingling), physical barriers (callose or lignin deposition at interface), or developmental asynchrony (mismatched cambial activity timing). Localized incompatibility is more amenable to management through improved grafting technique, callusing conditions, and timing optimization than translocated incompatibility.
Schnittstellenunverträglichkeit
Vereinigungszonen-Störung
interface incompatibility
lokalisierte Inkompatibilität
structural incompatibility
union zone dysfunction
Pina A et al. (2012) Differential induction of phenylalanine ammonia-lyase gene expression in response to in vitro callus unions of Prunus spp. Journal of Plant Physiology 169:1724-1730 Cookson SJ et al. (2014) Heterografting with nonself rootstocks induces genes involved in stress responses at the graft interface when compared with autografted controls. Journal of Experimental Botany 65:2473-2481
Localized Incompatibility Disposition
Syrah locular inner pericarp (layers 2-5 of inner pericarp, 120-185 μm thickness) showing characteristic structure= innermost locular layer (layer 2, adjacent to seed-adjacent layer 1) with cells 58-72 μm diameter, walls 1.3-1.7 μm, tannins 5.2-6.5 mg/g mid locular layers (layers 3-4) with cells 68-85 μm, walls 1.0-1.4 μm, tannins 4.2-5.5 mg/g outermost locular layer (layer 5, adjacent to middle pericarp) with cells 78-92 μm, walls 0.9-1.2 μm, tannins 3.5-4.8 mg/g, demonstrating progressive gradients in cell size, wall thickness, and tannin content creating smooth transition from seed-adjacent maximum to middle pericarp minimum. Cabernet Sauvignon locular inner pericarp in berries with different seed numbers= 2-seed berries showing locular inner pericarp total surface area 18-24 mm² (lining 2 locular cavities containing 2 seeds), volume 0.032-0.045 cm³ 4-seed berries showing 28-38 mm² surface area, 0.048-0.068 cm³ volume, demonstrating that higher seed number increases locular surface area and locular inner pericarp tissue volume by 40-60% potentially affecting total inner pericarp tannin contribution to wine though concentration per unit tissue mass remains similar. Merlot locular inner pericarp tannin gradient quantified by micro-dissection= innermost locular (70 μm from middle pericarp boundary) 5.8 mg/g tannins, mid locular (140 μm) 4.6 mg/g, outermost locular (210 μm) 3.8 mg/g, middle pericarp (>250 μm) 0.7 mg/g, showing exponential-like decline in tannin concentration with distance from seed-adjacent zone suggesting tannin synthesis or accumulation influenced by proximity to seeds or locular cavity boundary with effect extending 150-250 μm into pericarp tissue. Chardonnay white cultivar locular inner pericarp showing structure in white grapes= 3-4 cell layers, 135-195 μm thickness, cells 62-88 μm diameter, tannins 3.2-4.8 mg/g (moderate levels intermediate between seed-adjacent 4.5-6.2 mg/g and middle pericarp 0.4-0.9 mg/g), demonstrating locular inner pericarp presence and tannin gradient in white cultivars similar to red cultivars though with lower absolute tannin levels. Pinot Noir locular inner pericarp contribution to wine tannins estimated from tissue proportion and tannin content= locular inner pericarp representing 3-5% of pericarp volume, containing 4.5-6 mg/g tannins, contributing approximately 12-18% of total skin tannins extracted during winemaking (seed-adjacent contributing 8-12%, outer pericarp contributing 60-70%, middle pericarp 2-5%), demonstrating locular inner pericarp as significant contributor to wine tannin structure despite representing small proportion of berry tissue volume.
A locular inner pericarp is the inner pericarp tissue comprising the 2-5 outermost cell layers of the inner pericarp region (excluding the innermost 1-3 seed-adjacent layers), lining the locular cavities but not necessarily in direct seed contact, typically 80-220 μm thickness, characterized by cells intermediate in size between seed-adjacent inner pericarp (smallest) and middle pericarp (largest), moderate-to-high tannin content intermediate between seed-adjacent maximum and middle pericarp minimum, and position defining the locular cavity boundaries where seeds develop and mature. Locular inner pericarp anatomical organization comprises= (1) cell layers 2-5 (counting from innermost) within the inner pericarp region, with layer numbering varying depending on total inner pericarp thickness (3-8 total layers) (2) cells showing progressive size increase from innermost locular inner pericarp (adjacent to seed-adjacent zone, 55-75 μm diameter) through mid locular inner pericarp (65-85 μm) to outermost locular inner pericarp (approaching middle pericarp boundary, 75-95 μm) (3) cell walls showing progressive thinning from innermost locular (1.2-1.6 μm) to outermost locular (0.9-1.3 μm) approaching middle pericarp wall thickness (0.5-0.8 μm) (4) cells lining and defining the locular cavity spaces (typically 2 cavities per berry reflecting bicarpellate origin) where 2-4 seeds develop in seeded types or seed traces in stenospermocarpic seedless types. Locular inner pericarp compositional characteristics= (1) moderate-to-high tannin content typically 3.5-5.5 mg/g dry weight, intermediate between seed-adjacent inner pericarp maximum (5.5-8 mg/g) and middle pericarp minimum (0.3-1 mg/g), with progressive tannin decline from inner to outer locular layers approaching middle pericarp (2) tannin composition showing predominantly procyanidins with polymerization degree 10-16 subunits and galloylation 11-18% similar to other inner pericarp zones (3) sugar and acid content 19-26% sugars, 0.5-0.85% organic acids, similar to middle pericarp showing minimal radial gradients in primary metabolites (4) minimal anthocyanins (<0.1 mg/g in red cultivars) similar to middle pericarp with anthocyanins concentrated in outer pericarp hypodermal region. Locular inner pericarp function and significance= defines the anatomical boundaries of locular cavities providing mechanical support and biochemical boundary between fleshy pericarp tissue and seed-containing spaces provides tannin-enriched tissue contributing to wine phenolic composition during maceration creates transition zone between highly tannin-rich seed-adjacent cells and low-tannin middle pericarp allowing gradual rather than abrupt tissue property changes may serve protective function with elevated tannins providing antimicrobial activity in seed-proximate region. Locular inner pericarp development parallels overall inner pericarp development= cells undergo modest expansion during ripening (120-180% diameter increase versus 200-300% in middle pericarp) creating smaller final size despite similar initial size tannin synthesis and accumulation during ripening phase creating elevated tannin levels at maturity locular cavity air space development during late ripening as pericarp expands more than seeds creating gaps between locular inner pericarp cells and seed surfaces. Locular inner pericarp variation among cultivars and berries= cultivar variation in total inner pericarp thickness (100-300 μm) influences locular inner pericarp thickness and layer number berry seed number variation (2-4 seeds per berry) affects total locular surface area and total locular inner pericarp tissue volume cluster position effects (sun versus shade) may influence locular inner pericarp tannin content through effects on overall berry phenolic synthesis.
Locular_Inner_Pericarp represents the outer layers of inner pericarp and is distinguished from Seed_Adjacent_Inner_Pericarp (innermost 1-3 layers) by position slightly more distant from seeds. Locular inner pericarp defines locular cavity boundaries and shows intermediate properties between seed-adjacent maximum and middle pericarp minimum. Understanding locular inner pericarp is relevant for detailed anatomical characterization and for comprehending tannin distribution gradients within the inner pericarp region, contributing to understanding of spatial phenolic organization in berry tissues.
Fruchtkammer-Schicht
Kammer-Fruchtfleisch
cavity-lining layer
chamber pericarp
lokulares inneres Perikarp
Cadot, Y. et al. (2006). Tannin composition in grape tissues. Journal of Agricultural and Food Chemistry 54=2973-2987 Downey, M.O. et al. (2003). Relationship between skin tannin and berry development. American Journal of Enology and Viticulture 54=114-119.
Locular Inner Pericarp
Cabernet Sauvignon locular tissue at maturity comprising 5-7 cell layers (total 155-230 μm thickness) lining 2 locular cavities (each cavity 3.5-5.5 mm diameter containing 1-2 seeds plus air space), with cells showing systematic gradation from outermost locular (72-88 μm diameter, 0.9-1.2 μm walls, 4-5.5 mg/g tannins) through middle locular (60-78 μm, 1.1-1.4 μm walls, 5-6.5 mg/g tannins) to innermost locular adjacent to cavity/seed (52-68 μm, 1.3-1.6 μm walls, 6-7.5 mg/g tannins), creating locular cavity internal surface with distinct compositional properties compared to outer pericarp or middle mesocarp. Syrah locular tissue cavity structure examined by micro-CT imaging= 2 locular cavities (left and right oriented perpendicular to major berry axis) each containing 1-2 seeds, cavity internal surface area 32-42 mm² per berry, cavity volume 12-22 mm³ (seeds occupying 35-48% of volume, air space 52-65%), locular tissue forming continuous boundary around entire cavity creating complete cellular lining from cavity base (near pedicel) to cavity apex (near stylar scar), demonstrating three-dimensional organization of locular tissue as cavity-defining structure. Merlot locular tissue seed number effects on cavity and tissue dimensions= berries with 2 seeds showing 2 locular cavities with combined internal surface 26-36 mm², locular tissue volume 0.028-0.042 cm³ berries with 4 seeds showing larger cavity surface 38-52 mm², locular tissue volume 0.045-0.065 cm³ (50-65% increase), demonstrating that higher seed number expands locular cavity dimensions and consequently locular tissue extent though not proportionally as cavities accommodate multiple seeds creating shared rather than individual chambers. Pinot Noir locular tissue developmental changes in cavity air space= early development (weeks 3-6, Phase I) showing locular cavities completely filled by developing seeds with minimal air space, locular tissue directly contacting seeds mid development (weeks 8-10, lag phase/veraison) showing beginning air space formation (10-20% of cavity volume) as berry expansion begins exceeding seed size late ripening (weeks 14-18) showing substantial air space (45-65% of cavity volume) with 20-45 μm gaps between seeds and innermost locular tissue, demonstrating progressive cavity air space development creating transition from seed-filled to partially air-filled cavities as berry matures. Chardonnay locular tissue in white cultivar showing structural similarity= 4-5 cell layers lining 2 locular cavities, total thickness 140-200 μm, cells showing size gradient 54-88 μm from innermost to outermost layers, walls 0.9-1.4 μm, tannins 2.8-5.5 mg/g showing gradient from outer to inner locular tissue, demonstrating locular tissue as structural feature present in both red and white cultivars with similar anatomical organization though white cultivar absolute tannin levels lower than red cultivar locular tissue reflecting overall phenolic biosynthetic differences between cultivar types.
A locular tissue is endocarp tissue defining and lining the locular cavities (chambers) where seeds develop and mature, essentially synonymous with endocarp in functional terms but emphasizing cavity-boundary role rather than seed-proximity role, typically 100-300 μm thickness comprising 3-8 cell layers arranged parallel to locular cavity surfaces, serving as cellular boundary between fleshy mesocarp and air-filled or seed-containing chambers, providing structural support maintaining cavity integrity during berry expansion, and contributing phenolic compounds to wine during extended maceration. Locular tissue organization and function= (1) cells arranged in curved layers following locular cavity contours creating three-dimensional boundary surface around typically 2 locular cavities per berry (reflecting bicarpellate floral origin with two carpels fusing to form ovary) (2) locular cavity size and shape varying during development from seed-filled cavities during early growth (weeks 1-8) to partially air-filled cavities during ripening and maturity (weeks 9-18+) as berry expansion exceeds seed size increase (3) total locular tissue surface area (internal surface area of cavities) typically 25-45 mm² per berry depending on berry size and seed number, representing substantial internal surface though small fraction of total pericarp volume (4-7%) (4) structural support function maintaining cavity shape and preventing cavity collapse during berry growth and handling despite soft fleshy mesocarp bulk surrounding cavities. Locular tissue cellular characteristics showing endocarp properties= (1) cell size decreasing from outermost locular tissue near middle mesocarp (70-90 μm diameter) through middle locular tissue (55-75 μm) to innermost locular tissue adjacent to seeds or air space (45-65 μm) (2) wall thickness increasing from outermost (0.8-1.1 μm) to innermost (1.2-1.6 μm) locular tissue layers (3) tannin content showing gradient from moderate outer locular tissue (3.5-5.5 mg/g) to elevated inner locular tissue (5.5-7.5 mg/g) approaching or reaching seed-adjacent maximum (6.5-8.5 mg/g) (4) all cellular characteristics showing systematic radial gradients creating smooth compositional and structural transition from middle mesocarp bulk to innermost seed-adjacent layers. Locular tissue in berry architecture= (1) definition of internal berry organization creating distinct tissue zones (outer protective skin, middle storage flesh, inner locular boundary, seed chambers) rather than homogeneous tissue throughout (2) accommodation of seed development providing space for seeds to develop and mature surrounded by fleshy pericarp (3) locular cavity air space development during ripening creating berry internal structure with seeds surrounded by air space bounded by locular tissue lining rather than seeds embedded in continuous flesh (4) potential influence on berry shape and symmetry with locular cavity size and position affecting overall berry geometry. Locular tissue development paralleling berry growth= early berry development (Phase I) with locular tissue cells undergoing division and modest expansion establishing cavity-lining cell population seeds growing to fill available space during Phase I creating seed-filled cavities lag phase (weeks 6-8) with seed maturation completing and seeds achieving final size while locular tissue continues some development ripening (Phase II, weeks 9-18+) with berry expansion exceeding seed size increase creating progressive enlargement of locular cavities from seed-filled to partially air-filled state with 10-50 μm air gaps between seeds and innermost locular tissue maturity showing maximum cavity size with seeds occupying 30-50% of cavity volume and air space filling remainder. Locular tissue in winemaking providing inner pericarp tannin contribution= locular tissue cells breaking down during maceration releasing tannins into must/wine outermost locular tissue layers extracting relatively early (days 8-12) as cells more similar to middle mesocarp properties innermost locular tissue layers extracting later (days 12-20) requiring extended maceration for substantial breakdown total locular tissue contributing 15-25% of wine skin tannins during extended maceration representing significant phenolic reservoir accessed through prolonged skin contact.
Locular_Tissue is essentially synonymous with Endocarp but emphasizes cavity-lining and boundary function rather than seed-proximity. Sub-classification into Cavity and Chamber locular tissue may reflect different aspects of cavity organization. Understanding locular tissue is relevant for detailed berry internal anatomy characterization, for comprehending berry three-dimensional structure with distinct tissue zones and internal cavities, and for winemaking where locular tissue represents the innermost extractable pericarp tannin reservoir surrounding seed chambers.
Höhlenwand-Gewebe
Kammer-Gewebe
Lokular-Gewebe
cavity-lining tissue
chamber tissue
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 180-185 Harris, J.M. et al. (1968). The Vinifera grape berry= an anatomical study. Botanical Gazette 129':1-23.
Locular Tissue
Wild V. vinifera ssp. sylvestris staminate male flowers exhibit long filaments 3.6-4.5 mm at maturity (BBCH 61-65), achieved through vigorous rapid elongation from pre-anthesis length 1.2-1.5 mm, elevating anthers (0.8-1 mm) to heights of 4.4-5.5 mm above receptacle base and 3.8-4.8 mm above the vestigial pistil remnant (<0.5 mm), maximizing pollen dispersal geometry for wind-pollination of pistillate females located 15-80 m distant in riparian population structures. Syrah hermaphrodite flowers (vigorous selections) show long filaments 3.4-4.2 mm positioning anthers 1.2-1.8 mm above pistil stigma (2.2-2.4 mm), creating enhanced anther-pistil separation compared to typical Syrah (0.8-1.4 mm separation with 3-3.6 mm filaments) and potentially contributing to high fruit set (58-65% versus typical 54-62%) through improved pollen dispersal efficiency. Grenache under optimal growing conditions exhibits long filaments 3.2-3.9 mm (upper range for this cultivar, typical 2.8-3.5 mm) when vines receive adequate irrigation and nutrition during pre-bloom development (BBCH 53-59), demonstrating environmental modulation of filament length within genotype-specific ranges.
A long filament is a filament exhibiting extended length typically exceeding 3 mm at maturity, characteristic of staminate flowers in dioecious Vitis types and representing the upper range of hermaphrodite flower filament length variation, functioning to elevate anthers to maximum height for enhanced pollen dispersal range in obligate cross-pollinating dioecious systems or for maximal anther-pistil separation in hermaphrodite self-pollinating systems. Long filaments are characterized by= (1) mature length typically 3-4.5 mm (staminate flowers) or 3-4 mm (upper range hermaphrodite flowers), representing 120-180% of typical hermaphrodite filament length (2.5-3.5 mm) (2) proportionally greater elongation during the rapid elongation phase at anthesis, often achieving 200-350% length increase (from pre-anthesis 1-1.5 mm to mature 3-4.5 mm) compared to typical 100-200% increase in moderate-length hermaphrodite filaments (3) enhanced cortex parenchyma cell number and cell expansion capacity supporting the extended final length (4) erect orientation in staminate and hermaphrodite flower contexts, though long filaments can also occur in reflex orientation in some functionally female flowers with enhanced stamen development. Long filaments in staminate flowers serve to maximize anther elevation above the vestigial or rudimentary pistil, positioning anthers 3-4 mm above the receptacle base and creating optimal height for wind-borne pollen dispersal to distant pistillate flowers, with elevated anther positioning increasing the probability of pollen entering air currents and being transported 20-200 m to receptive stigmas. Long filaments in hermaphrodite flowers create enhanced anther-pistil separation (typically 1-1.8 mm anther elevation above stigma with long filaments versus 0.5-1.2 mm with moderate filaments), potentially enhancing pollen dispersal efficiency and increasing probability of both self-pollination and cross-pollination to adjacent flowers. The developmental regulation of filament length involves genetic factors controlling cortex cell proliferation and expansion capacity, with long filament phenotypes in staminate flowers resulting from sex-determination genes that promote enhanced stamen development, and long filament variation in hermaphrodite flowers reflecting quantitative genetic variation modulated by environmental factors (particularly resource availability and environmental conditions during filament development). Environmental factors influencing long filament expression include= favorable growing conditions during flower development (adequate water, nutrients, moderate temperatures) promoting maximum filament elongation potential stress conditions (water deficit, nutrient limitation, temperature extremes) potentially reducing final filament length by 15-30% even in genotypes with long filament potential. Long filaments show greater absolute elongation distances during rapid elongation, requiring greater cellular resources (water uptake, cell wall synthesis, metabolic energy) to achieve full extension, making long filament flowers potentially more sensitive to resource limitations or environmental stresses during the critical anthesis period compared to short or moderate filament flowers. The functional significance of long filaments relates to pollination strategy= in dioecious systems, long filaments in staminate flowers are essential for effective cross-pollination through elevated pollen release in hermaphrodite systems, long filaments may enhance both self and cross-pollination efficiency while maintaining self-compatibility, potentially providing bet-hedging reproductive strategy.
Long_Filament represents extended filament development characteristic of staminate flowers and upper-range hermaphrodite variation, distinguished from Short_Filament (reduced length 3 mm versus <2 mm). The long filament phenotype has critical functional significance in dioecious reproductive systems where anther elevation is essential for cross-pollination success, and may provide subtle functional advantages in hermaphrodite systems through enhanced pollination efficiency. Long filaments require greater developmental resources and show proportionally greater rapid elongation dynamics, potentially creating greater sensitivity to environmental conditions during anthesis. Understanding long filament development and function informs evolutionary studies of Vitis sexual system diversity and may inform breeding strategies for manipulating reproductive characteristics.
ausgedehnter Staubblattfaden
elongated filament
extended stamen stalk
langer Staubfaden
verlängertes Filament
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 94-98 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644.
Long Filament
A long fruiting cane on Vitis vinifera cv. Chardonnay in a high-wire cordon system with vertical shoot positioning, retained at 14 nodes spanning 155cm along the fruiting wire, arising from a vigorous vine showing 850g pruning weight indicating capacity to support extended cane length, positioned to allow upward shoot growth into 180cm of vertical trellis space, expected to produce 13-14 fruiting shoots (assuming typical 92% budbreak for Chardonnay) bearing 24-26 clusters totaling approximately 6.5kg fruit, requiring active shoot thinning to 12 shoots, cluster thinning to 18 clusters, and multiple hedging operations to manage vigorous canopy growth and maintain fruit zone light exposure.
A fruiting cane retained at extended length during winter pruning, typically 10-15 nodes or more, characterized by substantial linear extent along trellis wires (100-180cm), high total bud count providing maximum crop potential, suitability for training systems with extensive trellis space (VSP, lyre, Geneva Double Curtain), and production of numerous fruiting shoots creating dense canopies requiring active shoot management. Long fruiting canes maximize yield potential per retained cane, efficient use of available trellis space in expansive training systems, and distribution of crop load across many node positions reducing individual shoot stress, but require adequate vine vigor to support large shoot numbers, skilled labor for canopy management preventing excessive density, and appropriate site conditions (deep soils, adequate water) supporting high productivity without quality compromise.
Long fruiting canes are appropriate for vigorous vines in expansive training systems with adequate trellis space and skilled labor for canopy management, but can create problems in moderate-vigor vineyards or labor-limited operations through excessive canopy density, shading, disease pressure, and fruit quality compromise. The decision to use long versus shorter fruiting canes depends on balanced pruning assessment (vine size justifying bud retention), training system architecture (available trellis space), labor resources (capacity for shoot management), and quality objectives (premium production favoring lower bud counts, bulk production accepting higher counts). In general, long fruiting canes suit New World high-productivity systems while shorter canes or spurs dominate traditional European quality-focused regions, though many exceptions exist based on local conditions and objectives.
Langrute
extended fruiting cane
high-node fruiting cane
lange Fruchtrute
maximum length cane
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115.
Long Fruiting Cane
Documentation of Long_Internode_Disposition at INRAE Montpellier characterized shoot architecture in cultivars with extended internode development. Reference cultivars 'Silvaner', 'Gewürztraminer', 'Riesling', and 'Müller-Thurgau' were assessed for internode dimensions, associated canopy properties, and mechanical characteristics. Long_Internode_Disposition cultivars showed consistent characteristics. 'Silvaner' exhibited archetypal long internodes with mean length of 10.2 cm (nodes 5-15), standard deviation of 1.8 cm, and individual internodes reaching 14 cm in the mid-shoot region. Shoot length to node number ratio was 11.8 cm/node (compared to 7.2 for short-internode types). 'Gewürztraminer' showed the longest internodes (11.5 cm mean) among commercial cultivars assessed. Canopy density analysis revealed that Long_Internode_Disposition cultivars achieved 1.4 leaf layers at 18 shoots/m cordon, compared to 2.1 leaf layers for short-internode cultivars at equivalent shoot density, demonstrating the canopy-opening effect of extended internodes. Cluster spacing averaged 12.5 cm in long-internode cultivars versus 7.2 cm in short-internode types, potentially affecting within-canopy Botrytis spread (cluster-to-cluster contact reduced). Shoot stiffness measurements confirmed the mechanical association—flexural rigidity averaged 1.6 N·mm² in Long_Internode_Disposition cultivars versus 2.6 N·mm² in short-internode types, explaining the tendency toward trailing growth. Cell anatomy revealed that long internodes resulted primarily from increased cell length (215 μm versus 130 μm) rather than cell number, indicating that cell expansion capacity is the primary determinant of Long_Internode_Disposition.
A Long_Internode_Disposition is an Internode_Length_Disposition that inheres in Vitis vinifera shoots and is realized through extended cell elongation producing internodes exceeding 9 cm mean length, resulting in open, extended shoot architecture with widely-spaced leaves and clusters, and typically contributing to trailing growth habit through reduced shoot stiffness. This disposition represents the upper range of internode dimensions, characteristic of cultivars with high gibberellin sensitivity or prolonged cell expansion phase that produces elongated internodes. Long-internode cultivars develop shoots where individual leaves are distinctly separated, light penetrates readily between leaf positions, and clusters are widely spaced along the fruiting zone. The mechanical consequence of long internodes is reduced shoot stiffness (each segment is longer relative to its diameter, increasing the bending moment), contributing to the association between long internodes and trailing or horizontal growth habit. Viticultural implications include naturally lower canopy density at equivalent shoot numbers (potentially advantageous for fruit exposure), wider cluster spacing (potentially affecting cluster-to-cluster disease spread), and more flexible shoots that may be easier to bend for training but harder to maintain in vertical position. Long_Internode_Disposition often co-occurs with Trailing_Growth_Disposition, as both reflect the same underlying shoot mechanical properties, though the correlation is not absolute.
Long_Internode_Disposition is_a Internode_Length_Disposition representing extended internode length (> 9 cm mean). Characterized by high cell elongation, lower shoot stiffness, and association with trailing growth habit. Examples include 'Silvaner', 'Gewürztraminer', 'Riesling', 'Müller-Thurgau', 'Pinot Gris'. Advantages include naturally open canopy, good light penetration, wide cluster spacing. Associated with high gibberellin sensitivity. Many German/Alsatian cultivars show this trait, potentially reflecting adaptation to humid climates where open canopies reduce disease.
Lange-Internodien-Disposition
Offene-Triebarchitektur
Weiter-Knotenabstand
extended internode
open shoot architecture
wide node spacing
Schultz HR (1992) An empirical model for the simulation of leaf appearance and leaf area development of primary shoots of several grapevine (Vitis vinifera L.) canopy systems, Scientia Horticulturae 52:179-200 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Edition, Academic Press Louarn G et al. (2008) Characterization of the light interception efficiency of grapevine, Annals of Botany 101:385-399
Long Internode Disposition
A long main petiole on a mature shade leaf at node 9 of Vitis vinifera cv. Sauvignon Blanc in Sancerre, France, measuring 14.2 cm length representing substantial extension in this inherently long-petioled cultivar, petiolar index of 1.05 indicating petiole length exceeding main vein length by 5%, diameter of 3.5 mm showing moderate thickness typical of shade leaf development, light green coloration without anthocyanin pigmentation, sparse pubescence, developing in response to interior canopy position receiving 35% full sunlight with low red=far-red light ratio triggering shade avoidance elongation response, demonstrating high flexibility enabling leaf blade positioning across wide range finding optimal light angles within heterogeneous light environment, supporting 3.2 gram leaf mass positioned at variable angles adjusting to shifting light conditions, showing characteristic open V-shaped petiolar sinus diagnostic for this cultivar, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing the combined effects of genetic predisposition (Sauvignon Blanc naturally long-petioled) and environmental response (shade-induced elongation) producing maximum petiole length expression.
A main petiole exhibiting length in the upper range for the species and cultivar in Vitis vinifera, characterized by dimensions typically exceeding 10-12 cm length representing extended structural connection between lamina and shoot, often with petiolar index (petiole length / main vein length ratio) exceeding 0.90-1.00 indicating petiole length approaching or exceeding lamina length, common development under low-light conditions where increased petiole length positions lamina away from shading structures improving light capture, occurrence in certain cultivars showing genetic predisposition toward long petiole development (Sauvignon Blanc, Riesling, Sémillon), functional implications including increased flexibility enabling greater leaf blade mobility and positioning range but also increased mechanical vulnerability to wind damage and greater vascular resistance potentially limiting maximum transpiration and photosynthesis rates, and ampelographic significance as cultivar-diagnostic feature particularly when combined with other morphological characteristics. Long main petioles develop through extended cell elongation during petiole development, influenced by genetic programming determining baseline length and environmental signals particularly shade avoidance responses triggered by low red=far-red light ratios detecting neighboring vegetation, hormonal regulation including gibberellin and auxin promoting elongation and ethylene potentially modulating responses, and resource availability supporting extended growth with adequate water and nutrients enabling completion of elongation programs.
Long main petioles are distinguished from medium and short petioles by dimensions typically exceeding 10-12 cm and petiolar indices approaching or exceeding 1.0, with exact thresholds varying by cultivar baseline dimensions. Genetic control establishes cultivar-specific length ranges, with long-petioled varieties including Sauvignon Blanc, Riesling, and Sémillon typically showing 11-15 cm petioles while short-petioled varieties like Gewürztraminer show 6-9 cm petioles under comparable conditions. Environmental plasticity creates substantial within-cultivar variation, with shade conditions triggering elongation that can increase petiole length by 30-50% compared to sun conditions through low red=far-red light ratio detection activating shade avoidance responses. Functional implications include advantages in heterogeneous light environments where long petioles enable positioning flexibility improving light capture, but disadvantages including increased mechanical vulnerability to wind breakage, greater vascular resistance potentially limiting maximum transpiration rates, and increased resource investment in non-photosynthetic support structures. Ampelographic utility requires standardizing measurement conditions to sun-exposed mature leaves at specific positions to minimize environmental variation affecting comparative assessments.
ausgedehnter Blattstängel
elongated main petiole
extended petiole
langer Hauptblattstiel
long leaf stalk
verlängerter Petiolus
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 183-197 Ballare (1999) "Keeping up with the neighbours= phytochrome sensing and other signalling mechanisms" Trends in Plant Science 4=97-102 Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 79-102.
Long Main Petiole
Crimson Seedless bears a Long_Storage_Table_Grape_Role as the benchmark for storage-stable red seedless, maintaining commercial quality for 8-12 weeks at -0.5°C with SO₂ pads and controlled atmosphere (2% O₂, 5% CO₂), enabling California fruit harvested in September to reach markets through December. Red Globe serves in a Long_Storage_Table_Grape_Role with exceptional storage potential (12-16 weeks) derived from its thick skin, firm texture, and high acid content, enabling sea-freight export from Chile and Peru to Asian markets with 4-6 week transit times plus retail shelf life. Autumn King bears a Long_Storage_Table_Grape_Role for late white seedless, with storage capability of 10-14 weeks supporting the "California grapes through Christmas" marketing positioning. Thompson Seedless demonstrates moderate long-storage potential (6-10 weeks with SO₂) when harvested at optimal maturity (18-20°Brix) and handled with strict temperature management.
A table grape scion role that inheres in a grafted scion cultivar by virtue of its capacity for extended post-harvest storage while maintaining commercial quality, enabling supply continuity during periods when fresh harvest is unavailable. Long storage table grape scions possess physiological and structural characteristics supporting storage durations of 6-12+ weeks under controlled atmosphere or modified atmosphere conditions (typically 0-2°C, 90-95% RH, low O₂/elevated CO₂), including firm berry texture with thick, intact cuticle resisting water loss and decay, strong rachis (cluster stem) remaining green and turgid, high organic acid content (>0.5% titratable acidity) providing microbial resistance, low respiration rate minimizing quality degradation, and resistance to storage disorders (SO₂ injury, stem browning, shatter, decay). Storage potential is quantified by weeks of marketable quality retention under optimal conditions, with long-storage varieties maintaining grade standards for 8-16 weeks versus 2-4 weeks for storage-sensitive varieties. Major long-storage varieties include Crimson Seedless, Red Globe, Autumn King, and Thompson Seedless (with SO₂ treatment).
Long_Storage_Table_Grape_Role captures the post-harvest dimension critical for global table grape trade, where storage capability determines market reach and supply chain flexibility. Storage potential interacts with harvest timing—later-harvested fruit often stores better due to firmer texture and higher acidity, creating overlap with Late_Season classification. The development of controlled atmosphere storage, SO₂-generating pads, and modified atmosphere packaging has extended storage potential for varieties previously considered storage-limited. Breeding programs increasingly prioritize storage characteristics alongside eating quality, with new proprietary varieties evaluated through standardized storage trials. Climate-controlled sea containers (typically 45-60 day transit for Southern Hemisphere to Asia routes) require minimum 8-week storage potential for viable export positioning.
Lagertraube
high storage potential grape
lagerfähige Tafeltraube
langzeitlagerfähige Tafeltraube
long-keeping grape
storage grape
Crisosto CH, Mitchell FG (2002) Postharvest handling systems: table grapes, in Postharvest Technology of Horticultural Crops 3rd Ed, UC ANR pp.: 357-363 Lichter A et al. (2011) The effect of table grape storage temperature on sensory quality, Postharvest Biology and Technology 59: 1-8 Creasy GL, Creasy LL (2009) Grapes, CABI Palou L et al. (2010) Control of postharvest diseases of table grapes, Stewart Postharvest Review 6(4): 1-11
Long Storage Table Grape Role
Gewürztraminer functionally female flowers exhibit long styles 0.75-0.92 mm connecting enhanced ovaries 1.08-1.28 mm to prominent stigmas 0.55-0.65 mm, with long styles representing 30-36% of total pistil length (2.5-2.8 mm), containing well-developed transmitting tract tissue with elaborate extracellular matrix extending through the greater style length, requiring 68-95 hours for pollen tube transit from stigma to ovules when cross-pollinated from hermaphrodite sources at 20-24°C, and achieving 28-38% fruit set through obligate cross-pollination. Wild V. vinifera ssp. sylvestris pistillate female flowers show long styles 0.78-0.95 mm connecting robust ovaries 1.15-1.35 mm to elaborate stigmas 0.52-0.62 mm, with long styles representing 32-38% of pistil length (2.4-2.75 mm), positioned prominently above vestigial stamens (0.3-0.8 mm), requiring cross-pollination from staminate males and achieving 30-42% fruit set when males are within 20-50 m providing adequate wind-borne pollen. Orange Muscat constitutively functionally female cultivar exhibits long styles 0.72-0.88 mm as part of consistently elaborated pistil morphology (total pistil 2.35-2.7 mm) across all flowers and all environmental conditions, demonstrating genetic determination of long style phenotype associated with functional female sexual expression. Breeding populations from wild dioecious × cultivated hermaphrodite crosses segregate for pistil morphology, with pistillate offspring showing long styles 0.75-0.98 mm while hermaphrodite offspring show medium styles 0.52-0.68 mm, allowing style length to serve as diagnostic feature for identifying flower sexual type during early seedling evaluation and for studying genetic control of pistil elaboration.
A long style is a style exhibiting enhanced length typically exceeding 0.7 mm at maturity, characteristic of functionally female and pistillate flowers with elaborated pistils and representing styles at the upper end of the length range where extended pollen tube pathway correlates with enhanced female reproductive investment and often with extended stigma receptivity. Long styles are characterized by= (1) length typically 0.7-1 mm at anthesis, exceeding typical hermaphrodite style length (0.45-0.7 mm) by 10-50% (2) diameter typically 0.25-0.4 mm proportional to enhanced length (3) well-developed or enhanced transmitting tract tissue with elaborate extracellular matrix, abundant specialized transmitting tract cells, and sometimes extended central canal reflecting the greater length (4) connection between enhanced ovary (typically 0.9-1.3 mm) and elaborate stigma (typically 0.45-0.65 mm) in functionally female or pistillate flower contexts, creating pistils (total length typically 2.2-2.9 mm) with long styles representing 28-38% of pistil length. Long styles develop through enhanced pistil development in functionally female and pistillate flowers where female organ elaboration includes proportional or disproportional style elongation= style primordium initiation and early development (BBCH 51-54) may already show enhanced growth compared to hermaphrodite flowers style elongation phase (BBCH 54-58) shows greater cell division and expansion producing extended final length transmitting tract differentiation (BBCH 56-59) proceeds throughout the greater length creating extended functional pathway. Long styles in functionally female and pistillate flowers maintain proportional or slightly enhanced representation within total pistil length (28-38% of pistil versus 20-35% in hermaphrodite) suggesting that style elongation is part of coordinated pistil elaboration rather than disproportionate style-specific enhancement, though some flowers show styles at the upper end of proportional range indicating somewhat disproportionate style elongation. The functional implications of long styles relate to enhanced female reproductive capacity and cross-pollination dependency= long styles create extended pollen tube pathway (total stigma-to-ovule distance 1.8-2.8 mm versus 1.5-2.5 mm in hermaphrodite flowers) requiring longer pollen tube growth times (65-110 hours versus 48-96 hours at comparable temperatures), potentially providing extended opportunity for pollen tube selection or quality control through the longer transmitting tract passage long styles in functionally female and pistillate flowers often coincide with extended stigma receptivity periods (5-8 days versus 3-6 days in hermaphrodites), creating extended temporal windows for successful cross-pollination from hermaphrodite or staminate pollen sources the combination of long styles with reflex stamens (functionally female) or vestigial/rudimentary stamens (pistillate) creates spatial and functional barriers to self-pollination, enforcing cross-pollination dependency despite presence of functional transmitting tract capable of supporting pollen tube growth from any source.
Long_Style represents enhanced style length characteristic of functionally female and pistillate flowers and is distinguished from Medium_Style (0.45-0.7 mm, typical hermaphrodite) by clear elongation and from Short_Style (<0.45 mm, reduced functionally male/staminate) by substantially greater dimensions. Long styles occur in context of overall pistil elaboration and are associated with obligate cross-pollination systems where extended pollen tube pathways and extended stigma receptivity increase probability of successful cross-pollination. Sub-classification into Extended and Elongated types reflects gradations within the long style range. Understanding long styles is relevant for identifying functionally female and pistillate flowers, for understanding female reproductive investment strategies in cross-pollinating flower types, and for genetic studies of sex determination and pistil development.
elongated style
extended style
langer Griffel
verlängerter Stempelgriffel
verlängerter Stylus
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 94-101 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=93-109 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 102-106.
Long Style
Vine longevity varies substantially across cultivars and growing conditions, with well-managed vines of long-lived cultivars such as Zinfandel and Mourvèdre remaining productive for 80-150+ years, while shorter-lived types may decline within 25-40 years. Resilience encompasses recovery from frost damage (regrowth from basal buds), drought stress (osmotic adjustment, root regeneration), mechanical injury (wound compartmentalization, callus formation), and chronic stress from trunk diseases (survival with partial vascular dysfunction). Old vines (>40 years) often show reduced yield (30-60% of peak production) but enhanced fruit quality through deeper root systems, improved vine balance, and concentration effects from smaller berry size. The economic value of old vine vineyards reflects both the irreplaceable time investment and the perceived quality advantages.
A grapevine disposition that is the inherited capacity of a Vitis vinifera plant to maintain productive function over extended time periods (decades to centuries) and to recover from stress events, damage, or decline through regenerative growth, encompassing traits related to structural durability, disease resistance of perennial tissues, carbohydrate reserve management, and the capacity for renewal from latent buds.
Longevity_And_Resilience_Disposition serves as the top-level parent class for traits governing vine persistence and recovery capacity. The category addresses the unique perennial nature of grapevines, which can remain productive far longer than most agricultural crops. Understanding longevity determinants has both economic implications (vineyard investment protection) and cultural significance (preservation of heritage vineyards). Climate change and emerging trunk disease pressures are increasing attention to resilience traits in cultivar and rootstock selection.
Langlebigkeit und Resilienz
Rebstockausdauer
Widerstandsfähigkeit
long-term productivity
persistence disposition
vine durability trait
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 145-195 Gramaje D et al. (2018) Managing grapevine trunk diseases with respect to etiology and epidemiology: current strategies and future prospects. Plant Disease 102:12-39
Longevity And Resilience Disposition
Sauvignon Blanc exhibits loose cluster disposition with 4-6 berries/cm rachis, OIV 204 scores of 3-5, and clearly visible rachis branches between berry groups. Cabernet Sauvignon shows moderately loose architecture (5-7 berries/cm) with elongated conical cluster shape facilitating air movement. Loose clusters in Sauvignon Blanc demonstrate Botrytis incidence of 5-15% compared to 25-45% in compact Sémillon under identical conditions in Bordeaux. Spray coverage in loose clusters reaches 85-95% of berry surface versus only 50-70% in compact clusters, improving fungicide efficacy.
A cluster compactness disposition that is the inherited capacity of a Vitis vinifera plant to develop clusters with moderate berry spacing where most berries remain separated without mutual contact or deformation, with intermediate berry density (typically 4-8 berries/cm rachis) and adequate air circulation for disease prevention.
Loose cluster disposition represents the optimal range for disease management while maintaining commercially acceptable cluster appearance and harvest efficiency. The trait results from moderate fruit set, proportionate berry size to rachis framework, or elongated pedicels. Many premium wine cultivars possess loose cluster architecture, contributing to their reputation for quality through reduced rot incidence and more uniform berry ripening from improved light and air penetration.
locker gepackte Traubenstruktur
lockere Traube
luftige Traube
moderate density cluster
open cluster trait
well-spaced bunch
Dry PR et al. (2010) Bunch architecture and susceptibility of berries to Botrytis cinerea in Vitis vinifera L. Australian Journal of Grape and Wine Research 16:257-264 Herzog K et al. (2015) Initial steps for high-throughput phenotyping in vineyards. Vitis 54:1-5
Loose Cluster Disposition
Grenache in Châteauneuf-du-Pape vineyards at BBCH 65 exhibits very loose flower clusters of 18-25 cm length bearing 800-1500 flowers distributed across extensive primary, secondary, and tertiary branching with flower densities of 5-8 flowers/cm, creating broadly winged, multi-lobed structures (OIV code 1-2) that develop into large, loose berry clusters of 400-900 berries with <2% Botrytis incidence even in wet years with minimal fungicide use. Syrah in Northern Rhône (Hermitage) shows moderately loose flower clusters (12-18 cm length, 500-900 flowers, 8-10 flowers/cm density, OIV code 3) with well-developed secondary branching creating cylindrical clusters with excellent air circulation. Nebbiolo in Barolo exhibits loose flower clusters with pronounced lateral lobes (OIV code 2-3) facilitating extended hang time and optimal phenolic maturation with minimal rot risk.
A loose flower cluster is a flower cluster characterized by low flower density resulting from well-developed inflorescence branching with elongated internodes and extended pedicel length, creating open, well-aerated structures that facilitate air circulation, promote rapid drying after precipitation, minimize disease pressure, and typically develop into loose berry clusters with widely spaced fruit and minimal compression-related damage. Loose flower clusters exhibit flower densities typically below 8-12 flowers per cm of total inflorescence axis length (rachis plus all branch lengths combined), achieved through moderate flower initiation combined with extensive rachis and branch elongation, well-developed secondary and often tertiary branching, and long pedicel length (4-8 mm or more). These clusters typically measure 12-25 cm in length at BBCH 65 while bearing 400-1500+ flowers, creating length-normalized densities 25-75% of those observed in compact flower clusters. The loose architecture results from genetic programming for enhanced internode elongation during inflorescence development, typically combined with extensive secondary branching (4-8 secondary branches per primary) and frequent tertiary branching that distributes flowers across a large three-dimensional volume. Loose flower clusters characteristically show conical or compound shapes with pronounced lateral lobes or wings, often with length=width ratios approaching 1=1 or even width exceeding length in extremely loose types, creating a 'shouldered' or 'multi-lobed' appearance. Flower spacing within loose clusters typically ranges from 4-12 mm between adjacent flower pedicels, creating clearly separated flowers with substantial air space between them and excellent visual access to individual flowers. The open flower arrangement creates a microclimate closely matching ambient conditions, with relative humidity within loose clusters typically within 2-5% of ambient air, rapid drying after rain or dew (often 1-3 hours faster than compact clusters), and excellent air circulation penetrating to all flowers. This favorable microclimate dramatically reduces disease pressure, with loose flower clusters showing 50-85% lower Botrytis cinerea infection rates at bloom compared to compact clusters under identical environmental conditions, often allowing successful organic or low-input production with minimal or no bloom-period fungicide applications. The transformation from loose flower cluster to loose berry cluster involves minimal architectural constraint, as even high fruit set rates (40-60%) create well-spaced berry arrangements with berry-berry contact delayed until late maturation (BBCH 85-87) or absent entirely, berries maintaining round morphology without compression deformation, excellent air circulation and spray penetration persisting through ripening, and bunch rot incidence typically <2-5% even in humid climates. However, loose flower clusters present different challenges, including increased susceptibility to shatter (flower abscission) during bloom due to exposed flower positions, higher bird predation risk at maturity due to accessible berry positions, potential for uneven ripening across the extensive cluster volume, and lower weight-per-cluster reducing harvest efficiency. Loose flower cluster morphology characterizes cultivars like Grenache, Mourvèdre, Syrah, Nebbiolo, Carignan, and Cinsault, representing OIV codes 1-3 (very loose to loose).
Loose_Flower_Cluster is distinguished from Compact_Flower_Cluster by quantitative density metrics (18-25 flowers/cm), qualitative architectural features (extensive branching, wide spacing), and functional consequences (minimal disease risk, potential shatter and bird damage). The boundary between 'loose' and 'medium' density conventionally corresponds to OIV codes 1-3 versus codes 4-5. Loose flower clusters represent significant viticultural advantages in disease management and sustainability but create challenges in harvest logistics (lower cluster weight requires more clusters per unit yield) and quality management (potential uneven ripening, bird damage). Sub-classification into Widely_Spaced and Open types reflects gradations within the loose category. The extensive branching in loose clusters increases total photosynthetic area of cluster axes (rachis and branches bear stomata and chlorophyll), potentially contributing to sugar accumulation independently of leaf photosynthesis during early berry development.
aufgelockerte Blütentraube
locker verzweigte Blütentraube
lockere Blütentraube
open flower cluster
widely spaced flower bunch
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 94-97 OIV (2009). Descriptor List for Grape Varieties and Vitis Species, OIV code 204 levels 1-3 Vail, M.E., Marois, J.J. (1991). Grape cluster architecture and the susceptibility of berries to Botrytis cinerea. Phytopathology 81=188-191 Intrieri, C., Poni, S. (1995). Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality. American Journal of Enology and Viticulture 46=116-127.
Loose Flower Cluster
A loosely coiled functional tendril at node 10 on a shoot of Vitis vinifera cv. Müller-Thurgau in Baden, Germany, exhibiting three contact coils around a 12 mm diameter wooden stake with individual coil diameters of 35-40 mm (approximately 11× the 3.5 mm tendril diameter), pitch between coils of 18-22 mm creating widely spaced turns with visible gaps, seven free coils in the tendril shaft with similar open geometry, generating minimal visible shoot displacement (<2 cm) toward the stake indicating weak contractile force, showing light brown coloration in coiled regions suggesting moderate lignification, supporting a 55 cm shoot bearing five leaves and one small fruit cluster totaling 180 grams weight, exhibiting slight slippage of contact coils along the stake surface when shoot is manually displaced, measured at BBCH stage 75 (berries pea-sized) in late June.
A coiled functional tendril exhibiting relatively low coil density and large coil diameter relative to tendril thickness in Vitis vinifera, characterized by contact coils with diameters typically 8-15 times the tendril diameter creating open wrapping around support structures, free coils showing similar relaxed spiraling with substantial pitch (vertical distance between successive coil turns) often 1-3 cm or more, reduced mechanical tension in coiled regions generating moderate contractile forces with limited shoot displacement toward supports, and less extensive structural reinforcement through moderate lignification that maintains some tissue flexibility even after coil fixation. Loosely coiled functional tendrils develop under conditions including suboptimal contact stimulation with intermittent or weak pressure on mechanosensory cells, support structures with diameters larger than optimal tendril grasping range (typically >12-15 mm), environmental stress during coiling including water deficit or temperature extremes that limit cell expansion capacity, mechanical disturbances during the critical coiling period, or genetic variation in coiling behavior among cultivars showing naturally relaxed coiling tendencies. These tendrils provide functional but less secure attachment compared to tightly coiled tendrils, with reduced grip force from fewer contact points and lower friction, weaker contractile effects allowing greater shoot mobility, and increased susceptibility to mechanical failure under heavy loading or storm conditions. However, loosely coiled tendrils offer advantages in flexibility, accommodating shoot growth movements without creating excessive constraint, and may represent adaptive responses in cultivars or environments where frequent shoot repositioning benefits light capture or wind resistance.
Loosely coiled functional tendrils are distinguished from tightly coiled tendrils by quantitative coil geometry including larger coil diameter relative to tendril thickness (ratio >7=1), greater pitch between turns, and reduced contractile effects. Loose coiling may represent either suboptimal development under limiting conditions or adaptive variation in cultivars adapted to specific training systems or environmental conditions. These tendrils remain functional for light support loads but may require supplemental attachment through shoot positioning or additional tendrils for heavy crop loads. Loose coiling around thick supports (posts, tree trunks) may be the only achievable geometry given physical constraints, representing functional compromise rather than failed development. Environmental improvement during coiling period can shift tendril populations toward tighter coiling, indicating significant plasticity in coiling behavior.
entspannt gewundene Kletterorgan
locker gewickelte Ranke
low-tension tendril
offen spiralisierte Ranke
open coil tendril
relaxed coil tendril
Based on Jaffe & Galston (1968) "The physiology of tendrils" Annual Review of Plant Physiology 19=417-434 Silk & Hubbard (1991) "Axial forces and normal stresses in coiling tendrils" Biomechanics and Modeling in Mechanobiology 24=599-606 Isnard & Silk (2009) "Moving with climbing plants from Charles Darwin's time into the 21st century" American Journal of Botany 96=1205-1221 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 137-154.
Loosely Coiled Functional Tendril
Cabernet Sauvignon inflorescences at early BBCH 60 (first flowers beginning to open, <5% in active bloom) exhibit loosening calyptras visible upon close inspection as narrow basal gaps 0.4-0.8 mm wide appearing at one to two points around the circumference, minimal upward displacement (0.5-0.9 mm), calyptra still substantially covering enclosed stamens with only bases of filaments visible at the gap locations, with progression to partially separated status occurring 40-75 minutes later as stamen elongation continues and basal separation extends around remaining circumference. Sauvignon Blanc at the onset of bloom shows loosening calyptras characterized by light yellowish-green color change, visible darkened abscission zone at base, and subtle lifting at one side creating asymmetric appearance, with only 2-4% of total inflorescence flowers in loosening state at any given time as individual flowers rapidly transition to more advanced separation stages. Gewürztraminer experiencing borderline cool temperatures (14-16°C) during attempted bloom onset shows prolonged loosening phase with calyptras remaining in minimally displaced state for 90-150 minutes as slow stamen elongation delays progression to partial separation, creating extended vulnerability to Botrytis infection of the partially exposed basal flower regions and contributing to irregular bloom and reduced fruit set (28-35% versus normal 32-40% for this functionally female cultivar).
A loosening calyptra is a separating calyptra in early separation stage showing initial basal detachment around a limited portion of the circumference with minimal upward displacement, representing the first phase of active anthesis when abscission layer separation has begun but stamens have not yet generated sufficient upward pressure to substantially lift the calyptra, typically representing flowers in the first 15-60 minutes of the anthesis process. Loosening calyptras are characterized by= (1) initial basal separation around 10-50% of the circumference with narrow gaps (0.3-1 mm) appearing between calyptra base and receptacle rim as abscission layer dissolution creates first detachment points (2) minimal upward displacement (typically 32°C slow elongation and extend the loosening phase duration (2) humidity effects on tissue turgor and abscission enzyme activity, with moderate humidity (50-75%) supporting efficient separation while excessive humidity (>85%) can slow or disrupt the process (3) cultivar-specific variation in abscission layer formation kinetics and stamen elongation vigor creating differences in loosening phase duration ranging from 15-30 minutes in rapidly opening cultivars to 45-90 minutes in slower types. The observation of loosening calyptras indicates that bloom has just begun or is imminent, typically coinciding with early BBCH 60 (first flowers opening, <5% of flowers in bloom), and provides growers with advance warning that full bloom period requiring optimal environmental conditions will develop within 12-24 hours as the loosening flowers progress through partial separation to full anthesis.
Loosening_Calyptra represents the initial separation phase and is distinguished from Pre_Dehiscence_Intact_Calyptra (showing swelling but not yet separating) by presence of actual basal gaps indicating separation has begun, and from Partially_Separated_Calyptra (advanced separation with substantial displacement) by minimal displacement and limited circumferential detachment. The loosening stage is the most transient and least obvious separation phase, requiring careful observation for detection and typically representing <5% of flowers at any given time during bloom progression. Monitoring loosening calyptras is most relevant for research applications, controlled crossing operations requiring precise timing, or for early detection of bloom onset to optimize disease management timing. The loosening stage represents the commitment point for anthesis, as once basal separation initiates, the process typically proceeds to completion even if environmental conditions deteriorate.
beginnend abgelöste Kronblattkappe
beginnend getrennte Kelchhaube
early-separation calyptra
initially lifting petal cap
sich lockernde Blütenhaube
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 101-102 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 193-194 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 94-95.
Loosening Calyptra
Grenache exhibits low acidity retention disposition, declining to 4-5 g/L TA and pH 3.6-3.9 under warm Mediterranean conditions through rapid malate degradation (k = 0.06-0.08/day) combined with moderate tartrate (5-6 g/L). Chasselas shows low retention with harvest acidity of 4-6 g/L TA even in Swiss climates, attributed to moderate tartrate and rapid malate loss. Palomino (Sherry grape) reaches pH 3.8-4.2 with TA 3-4 g/L in Jerez, suited to the oxidative winemaking style requiring low acidity. Low-retention cultivars often show 2-3 fold higher malic enzyme activity than high-retention types, accelerating respiratory malate loss.
An acidity retention disposition that is the inherited capacity of a Vitis vinifera plant to exhibit reduced titratable acidity (typically below 5 g/L) and elevated pH (above 3.6) at harvest maturity, through combinations of lower tartrate concentration, rapid malic acid degradation, high potassium accumulation, and/or sensitivity to temperature-induced acid loss.
Low acidity retention disposition suits wine styles where softness and roundness are preferred over crisp freshness, including certain Mediterranean reds and fortified wine bases. The trait becomes problematic when pH rises above 3.8-4.0, increasing microbial spoilage risk and reducing color stability in red wines through anthocyanin equilibrium shifts. Acidification during winemaking can partially compensate but may affect wine character. Climate warming is shifting some traditionally balanced cultivars toward low-retention phenotypes, requiring management adaptation.
acid-depleting trait
geringer Säureerhalt
high pH disposition
low acid retention
niedrige Azidität-Bewahrung
niedrige Säureretention
Keller M et al. (2010) Interactive effects of deficit irrigation and crop load on Cabernet Sauvignon in an arid climate. American Journal of Enology and Viticulture 61:221-233 Mpelasoka BS et al. (2003) A review of potassium nutrition in grapevines with special emphasis on berry accumulation. Australian Journal of Grape and Wine Research 9:154-168
Low Acidity Retention Disposition
A low bush head arm on Vitis vinifera cv. Zinfandel in old California vineyard, emerging from head at 22cm above ground level, extending 38cm nearly horizontally creating very low spreading bush barely 45cm maximum height, clusters hanging close to soil surface (15-30cm height), traditional dry-farmed head-trained architecture from 1890s planting now heritage-protected, extremely low profile providing wind protection in exposed coastal site and heat benefits from soil radiation in cool-climate location, representing historical low bush training now rare but preserved in heritage vineyards, with very low stature creating severe harvest challenges (workers kneeling or sitting to access clusters) but perceived quality benefits from microclimate and low yields (1.8-2.5kg per vine from ancient stressed vines).
A bush head arm configured in low-profile architecture with head positioned close to ground level (typically 15-40cm height) and arms creating compact low bush, characterized by reduced overall vine height facilitating ground-level fruit positioning, protection from wind exposure, potential heat benefits from soil radiation, and suitability for extreme environmental conditions, traditional low-intervention systems, or specific microclimate management objectives. Low bush head arms represent extreme low-profile training, employed in challenging environments (strong wind exposure requiring low protected structure, extreme drought necessitating minimal canopy, marginal cool climates utilizing soil heat radiation), traditional systems in specific regions (Santorini kouloura, some Portuguese bush vines, parts of South African traditional training), and situations where low fruit positioning provides specific benefits (ground heat, wind protection, traditional harvest methods).
Low bush head arms represent extreme expression of head training, with very low positioning creating both benefits and severe challenges. Benefits include wind protection (low profile reduces exposure), potential heat benefits (proximity to soil surface capturing radiated heat in cool climates), drought tolerance (minimal canopy reducing transpiration), and in some cases frost protection (ground-level cold air drainage potentially less severe than elevated positions). However, challenges include extremely difficult harvest (workers must kneel, crouch, or sit to access low clusters), disease pressure from soil proximity (splash contamination, reduced air circulation), weed management complications (cultivation around very low vines), and worker health concerns (constant bending, kneeling causing ergonomic problems). The persistence of low bush training is largely limited to heritage vineyards, protected traditional systems, or extreme environmental situations where low positioning provides essential protective benefits outweighing operational difficulties.
bodennahe Kopfarm
ground-level bush arm
low-profile bush arm
niedrige Busch-Arm
prostrate head arm
Low Bush Head Arm
Thompson Seedless (Sultanina) exhibits low chilling requirement disposition with dormancy release achieved after only 200-350 chilling hours, enabling its widespread cultivation in warm regions from California to India. Flame Seedless requires 250-400 chilling hours, suitable for desert viticulture in the Coachella Valley with winter temperatures rarely below 10°C. Tropical cultivars and hybrids selected for Brazilian and Thai viticulture may require fewer than 150 chilling hours or show near-continuous growth capacity. Low-chill cultivars in temperate climates risk premature budbreak during warm winter periods followed by frost damage.
A chilling requirement disposition that is the inherited capacity of a Vitis vinifera plant to achieve complete dormancy release with minimal cold accumulation (typically less than 400 chilling hours or 30 chill portions), enabling cultivation in subtropical, Mediterranean, or tropical climates with limited winter cold.
Low chilling requirement facilitates warm-climate viticulture but creates vulnerability to erratic budbreak when winter temperatures fluctuate. Many table grape cultivars have been selected for low chilling to enable production in warm markets near consumption centers. The trait shows quantitative inheritance with moderate heritability, enabling breeding for climate-specific chilling requirements. Chemical dormancy release using hydrogen cyanamide can partially substitute for insufficient natural chilling in low-chill cultivars experiencing unusually warm winters.
low chill trait
niedrige Chilling-Anforderung
niedriger Kältebedarf
subtropical dormancy type
subtropischer Dormanz-Typ
warm-climate cultivar
Dokoozlian NK et al. (1995) Cultural practices and environmental influences on dormancy and dormancy release of grapevines. In: Rantz JM (ed) Proceedings of the International Symposium on Table Grape Production, ASEV pp.: 183-188 Camargo UA et al. (2012) Grapevine breeding for tropical and subtropical regions. Acta Horticulturae 931:357-364
Low Chilling Requirement Disposition
A Pinot Noir vine on low bilateral cordon at 70 cm in Burgundy's Côte d'Or bears a Low_Cordon_Role, the traditional height positioning fruit close to limestone soils for thermal reflection while enabling the meticulous hand labor of Grand Cru viticulture. A Sauvignon Blanc vine on low cordon at 85 cm in Marlborough serves in a Low_Cordon_Role with VSP canopy, the standardized positioning enabling efficient mechanical management while maintaining the fruit exposure critical for aromatic development. A Nebbiolo vine on low cordon in Barolo's steep Langhe hillsides bears a Low_Cordon_Role at 80 cm, the moderate height balancing accessibility for hand operations with sufficient elevation to avoid ground-level frost in spring.
A cordon-trained vine role that inheres in a grapevine with cordons positioned at moderate to low height (typically 60-100 cm above ground), creating a fruit zone accessible for hand labor while capturing radiated ground heat and potentially benefiting from soil thermal properties in cool climates. Low cordon training represents the standard positioning for most quality wine production worldwide, with cordons at heights enabling standing-position hand operations (pruning, shoot positioning, leaf removal, cluster work, hand harvest) while maintaining fruit close enough to the ground for thermal benefits. Vertical shoot positioning (VSP) is the dominant canopy system paired with low cordons, with upward shoot growth creating the "fruit wall" or "hedgerow" appearance characteristic of modern viticulture. Low cordon height varies with regional tradition and frost risk—Burgundy's cordons may be as low as 50-60 cm, while frost-prone New World sites position cordons at 90-100 cm to elevate fruit above cold air pooling. The low cordon/VSP combination optimizes fruit zone light exposure through the open canopy interior created by upward shoot training, supporting the color development and flavor concentration valued in premium wine production.
Low_Cordon_Role captures the dominant contemporary training configuration, optimized for the combination of hand labor accessibility, mechanical operation compatibility, and fruit zone microclimate management that characterizes quality wine production. The "low" designation is relative—60-100 cm positions are low compared to high cordon systems (150-200 cm) but elevated compared to ground-level or near-ground training. Low cordon height selection balances multiple factors: lower positions capture more ground heat (beneficial in cool climates) but increase frost risk, disease pressure from soil splashing, and back strain for workers higher positions reduce these problems but sacrifice thermal benefit and may reduce light interception in the fruit zone. The VSP canopy management typically paired with low cordons creates the manicured, hedgerow appearance that defines contemporary vineyard aesthetics while supporting the open, well-exposed fruit zones valued for quality production.
Niederkordon
Standardkordon
low cordon
low wire
mid-height cordon
mittlere Kordonhöhe
niedrige Erziehung
standard cordon
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Reynolds AG, Vanden Heuvel JE (2009) Training systems, American Journal of Enology and Viticulture 60: 251-268 Jackson RS (2014) Wine Science, Academic Press Gladstones J (1992) Viticulture and Environment, Winetitles
Low Cordon Role
A Shiraz vine at 2,200 vines/hectare (3.0 m × 1.5 m) in Barossa Valley bears a Low_Density_Planted_Role, the spacing accommodating the variety's vigorous growth on fertile valley floor soils while enabling bilateral cordon training with extensive spur positions. A Tempranillo vine at 2,500 vines/hectare in La Mancha serves in a Low_Density_Planted_Role, the traditional Spanish spacing adapted to dryland conditions where wide spacing enables each vine to access adequate water from limited rainfall. A Chardonnay vine at 1,800 vines/hectare on deep Central Coast soils bears a Low_Density_Planted_Role, the spacing managing vigorous growth that would create excessive shading at higher densities.
A vine spacing role that inheres in a grapevine planted at reduced density (typically 1,500-3,000 vines per hectare), with wider spacing reducing vine competition and enabling larger per-vine canopy development, suited to high-vigor growing conditions or production systems emphasizing per-vine rather than per-hectare productivity. Low-density planting provides each vine with greater root zone volume and canopy space, enabling the vigorous growth characteristic of fertile, irrigated, or high-rainfall sites without the excessive crowding that would result from closer spacing under such conditions. Training systems in low-density vineyards often feature extensive cordons, divided canopies, or spreading architectures that utilize the available space while managing the vigorous growth that reduced competition promotes. Low density reduces per-hectare vine investment costs and is associated with higher mechanization efficiency, as wider row spacing accommodates larger equipment and reduces passes per hectare. The approach is common in New World irrigated production, warm-climate viticulture, and any situation where high vigor would create management problems under closer spacing.
Low_Density_Planted_Role captures planting approaches suited to vigorous conditions or extensive production models, distinguished from Wide_Spacing which represents the extreme low end of the density spectrum. Low density is often the optimal choice for high-vigor situations—close spacing under such conditions creates dense, shaded canopies with worse fruit quality than would result from wider spacing that enables proper canopy development. The perception that low density is inherently inferior to high density reflects bias toward European models developed under different conditions for vigorous New World sites, low density may be the quality-optimizing choice. Low-density vineyard economics differ from high-density—lower establishment cost per hectare but potentially lower per-hectare revenue, with profitability depending on whether per-hectare or per-vine metrics drive value.
extensive Pflanzung
extensive planting
geringe Stockdichte
low-density planting
niedrige Dichte
reduced density
weite Bepflanzung
wide density
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Reynolds AG et al. (1994) Vine spacing, American Journal of Enology and Viticulture 45: 444-459 Dry PR (2000) Canopy management for fruitfulness, Australian Journal of Grape and Wine Research 6: 109-115
Low Density Planted Role
A low horizontal cordon on Vitis vinifera cv. Nebbiolo in Barolo, Italy, positioned at 55cm height along single wire, extending 68cm from trunk, bearing 11 two-bud spurs, shoots trained vertically upward without catch wires in traditional low-canopy architecture, harvest workers able to access clusters without bending or reaching (clusters hanging at approximately 45-80cm height range, comfortable working zone), representing traditional Barolo training emphasizing hand operations, fruit proximity to ground providing some microclimate benefits (soil heat radiation, protected position), but creating challenges for mechanical operations and requiring careful canopy management to prevent soil splash and disease pressure from ground proximity.
A horizontal cordon positioned at reduced height above ground level, typically 40-70cm, characterized by proximity to soil surface facilitating hand harvest access and traditional low-canopy architectures, spur positioning at ergonomically accessible height for manual pruning and crop management, suitability for traditional European training systems and premium hand-harvested production, and challenges for mechanical operations due to low equipment clearance and potential soil contact or splash contamination risks. Low horizontal cordons represent traditional configurations in many prestigious wine regions, reflecting historical practice developed before mechanization when hand operations dominated, with low positioning providing harvest worker comfort, reduced vine height simplifying stake or trellis requirements, and in some climates enhanced frost protection through reduced cold air exposure compared to higher cordons.
Low horizontal cordons persist in premium wine regions despite mechanization trends, justified by tradition, quality perceptions, hand-harvest requirements (delicate varieties, whole-cluster harvest needs), and in some cases genuine climatic benefits (frost protection in hollows, heat reflection from soil surface). However, low cordons create significant challenges including difficult mechanization (equipment clearance problems, soil contact risks), ergonomic issues for many operations (pruning, shoot positioning, cluster thinning requiring bending or kneeling), disease pressure from soil proximity (splash contamination, reduced air circulation), and difficult weed management (mowing and cultivation around low cordons). The trend in new plantings generally favors higher cordons (80-100cm) even in traditional regions, with low cordons increasingly limited to heritage vineyards, protected designation systems requiring traditional practices, or specific situations where low positioning provides demonstrable benefits.
bodennahe Kordon
ground-level cordon
low-positioned cordon
niedrige horizontale Kordon
traditional height cordon
Low Horizontal Cordon
Assyrtiko vines on Santorini bearing a low-input production role are cultivated in traditional kouloura basket-trained form without irrigation, fertilization, or trellising, receiving only 1-3 sulfur dustings per season at 15-25 kg/ha for powdery mildew, producing 2-4 t/ha from 60-80 year old ungrafted vines on volcanic soils where the unique terroir and cultivar adaptation replace external inputs, representing one of the most extreme examples of commercially viable low-input viticulture worldwide.
A sustainable production role that inheres in a grapevine cultivated with deliberate minimization of all external inputs including agrochemicals, fertilizers, irrigation water, and purchased energy, relying primarily on natural ecosystem processes, adapted cultivars, and site-specific resilience to maintain vine productivity at economically acceptable levels with reduced yield expectations.
Low_Input_Production_Role describes a management philosophy centered on input reduction rather than adherence to a specific certification standard. This role may overlap with organic, biodynamic, or conventional classifications: a low-input vineyard may or may not hold organic certification. The role is particularly relevant in Mediterranean, semi-arid, and traditional viticultural regions where historical practice and cultivar adaptation enable commercially viable production with minimal external inputs. The boundary between low-input and conventional extensive viticulture is contextual and depends on regional norms for input usage.
Extensiv-Produktionsrolle
Low-Input-Anbau-Rolle
eingriffsarme Bewirtschaftungsrolle
extensive viticulture role
low-input viticulture role
minimal intervention production role
Lacombe T et al. (2018) Grapevine European catalogue: towards a comprehensive list, Vitis 57: 69-76 Ferretti CG (2019) A new geographical classification of low-input viticulture, Oeno One 53: 507-523 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 556-561
Low Input Production Role
A low light-adapted mature leaf at node 7 on a lateral shoot of Vitis vinifera cv. Chardonnay in Champagne, France, positioned in mid-interior canopy receiving approximately 32% of full sunlight (daily PPFD averaging 520 μmol photons m⁻² s⁻¹ with midday peaks of 750 μmol photons m⁻² s⁻¹), measuring 14.8 cm length by 13.5 cm width representing 30% size increase compared to exterior leaves, lamina thickness of 245 μm with single palisade layer of 55 μm and spongy mesophyll of 165 μm, chlorophyll content of 365 μg cm⁻² with a=b ratio of 2.7 indicating moderate shade adaptation, cuticle thickness of 4.0 μm providing adequate but not maximal protection, stomatal density of 165 stomata mm⁻² with moderate-sized individual stomata, moderate trichome coverage creating medium green undersurface appearance, light to medium green adaxial surface without strong pigmentation, demonstrating light-saturated photosynthesis rate of 9.8 μmol CO₂ m⁻² s⁻¹ under ambient conditions averaging 600 μmol photons m⁻² s⁻¹, light compensation point of 28 μmol photons m⁻² s⁻¹, contributing positive carbon balance throughout most of daily light period, measured at BBCH stage 75 (berries pea-sized) in late June, representing moderate shade adaptation appropriate for intermediate canopy positions experiencing partial shading from upper canopy layers.
A shade mature leaf exhibiting moderate structural and functional adaptations to reduced light environments receiving 20-40% of full sunlight in Vitis vinifera, characterized by intermediate expression of shade leaf syndrome including moderately enlarged lamina size typically 25-35% increased compared to sun leaves providing enhanced light interception while maintaining reasonable structural investment, moderately thin lamina of 220-260 μm with single well-developed palisade layer and moderate spongy mesophyll development, intermediate chlorophyll concentrations per unit area of 320-400 μg cm⁻² supporting adequate photosynthetic capacity under moderate light conditions, moderate cuticle development of 3.5-4.5 μm providing basic protection without excessive resource allocation, moderate stomatal density of 150-175 stomata mm⁻² balancing conductance capacity and construction cost, light to medium green coloration reflecting intermediate total chlorophyll content, moderate chlorophyll a=b ratio of 2.5-2.8 indicating partial shift toward shade-type light-harvesting complexes, and physiological characteristics including moderate light-saturated photosynthesis rates of 8-11 μmol CO₂ m⁻² s⁻¹, intermediate light compensation points of 25-35 μmol photons m⁻² s⁻¹ enabling positive carbon balance under moderate shade while maintaining reasonable maximum capacity, and balanced respiratory costs supporting net productivity across variable daily light conditions typical of interior but not deep canopy positions.
Low light-adapted mature leaves represent intermediate phenotypes between sun and deep shade extremes, distinguished by moderate rather than maximum expression of shade characteristics and maintained capacity for reasonable photosynthetic performance across broader light ranges. These leaves develop in interior canopy positions receiving intermittent direct sunlight creating variable daily light patterns, or in consistently moderate shade from partial upper canopy coverage. Functional performance shows adaptation to moderate shade with lower light compensation points than sun leaves enabling net carbon gain under reduced light, but maintenance of sufficient capacity to utilize higher light periods effectively without photoinhibition risk. Carbon balance of low light-adapted leaves often depends on daily light integration, with morning and evening low-angle sunlight penetration or midday sunflecks providing critical high-light periods supporting positive daily carbon balance. These leaves represent the most common shade type in well-managed canopies with moderate density, providing substantial photosynthetic contribution while avoiding excessive shading that would create deep shade leaves with marginal or negative carbon balance.
intermediate light-adapted leaf
moderate shade leaf
mäßig-schatten Blatt
niedriglicht-adaptiertes reifes Blatt
partial shade leaf
teilschatten Weinblatt
Based on Schultz (1996) "Leaf absorptance of visible radiation in Vitis vinifera L.= estimates of age and shade effects with a simple field method" Scientia Horticulturae 66=93-102 Poni et al. (2006) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 57=397-407 Intrieri et al. (2008) "Grapevine growth and development" Horticultural Reviews 44=67-125 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 305-328.
Low Light Adapted Mature Leaf
Riparia Gloire de Montpellier (V. riparia) shows low lime tolerance (6-9% active lime), developing severe chlorosis on the marly soils of southern Burgundy but performing excellently on the acidic granite-derived soils of Beaujolais where it confers desirable low vigor to Gamay. 3309 Couderc (V. riparia × V. rupestris) tolerates only 11% active lime, limiting its use to low-limestone sites in Burgundy and Champagne despite excellent adaptation for premium wine production. Schwarzmann (V. riparia × V. rupestris) shows similarly low lime tolerance (10-11% active lime), restricted to acidic sandy and granitic soils of Alsace and Germany where its low vigor and early maturity are valued.
A lime-tolerant rootstock role that inheres in a grafted rootstock by virtue of its limited capacity to maintain scion iron nutrition on calcareous soils, showing chlorosis symptoms at active limestone concentrations exceeding 10-15% that would be tolerated by moderate- or high-lime-tolerance rootstocks. Low lime-tolerance rootstocks exhibit chlorosis (interveinal yellowing progressing to whitening of young leaves, reduced shoot growth, poor fruit set) when planted on soils with active CaCO₃ above 10-15%, soil pH above 7.5, or significant bicarbonate accumulation in soil solution. The limited tolerance reflects inherently low Fe³⁺-chelate reductase activity, weak rhizosphere acidification capacity, and/or inefficient iron translocation and utilization. Low lime-tolerance rootstocks are restricted to acidic, neutral, or weakly calcareous soils (active lime below 10%) and are selected for other attributes such as low vigor, early ripening induction, or superior nematode resistance that justify their use despite lime sensitivity.
Low_Lime_Tolerance_Rootstock_Role encompasses V. riparia selections (Riparia Gloire) and V. riparia × V. rupestris hybrids (3309C, 101-14Mgt, Schwarzmann) where the absence of V. berlandieri parentage results in limited lime adaptation. These rootstocks evolved on acidic soils of northeastern North America (V. riparia) and the Ozark region (V. rupestris) without selection pressure for calcareous soil adaptation. Despite lime sensitivity, these rootstocks remain commercially important for their low-to-moderate vigor, excellent phylloxera resistance, and favorable effects on fruit quality when matched to appropriate acidic or low-lime sites.
chlorosis susceptible rootstock
gering kalktolerante Unterlage
kalkempfindliche Unterlage
kalkintolerante Unterlage
lime intolerant rootstock
lime sensitive rootstock
low lime tolerance rootstock
Bavaresco L et al. (2003) Grapevine iron nutrition update, Journal International des Sciences de la Vigne et du Vin 37: 89-100 Pouget R (1987) Utilité de la connaissance du pourcentage de calcaire actif, Progrès Agricole et Viticole 104: 91-95 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia, Chaintre
Low Lime Tolerance Rootstock Role
SO4 (V. berlandieri × V. riparia) demonstrates low-to-moderate K uptake, maintaining Riesling must pH at 3.0-3.2 in the Rheingau compared to 3.3-3.5 on 1103 Paulsen, with petiole K at veraison typically 2.5-3.0% dry weight. 101-14 Millardet et de Grasset (V. riparia × V. rupestris) restricts K delivery to Merlot in Saint-Émilion, contributing to the fresh acidity and age-worthy tannic structure valued in premium Bordeaux. 3309 Couderc (V. riparia × V. rupestris) shows low K translocation to Pinot Noir in Burgundy, maintaining the acidity profile essential for Côte d'Or wines, with must pH typically 0.2-0.3 units lower than equivalent vines on 110 Richter.
A nutrient acquisition rootstock role that inheres in a grafted rootstock by virtue of its restricted potassium (K⁺) uptake from soil and translocation to the scion, resulting in moderate-to-low leaf petiole K concentrations and reduced K accumulation in berries and must. Low potassium uptake rootstocks typically deliver petiole K concentrations of 1.5-2.5% dry weight at bloom and 2-3.5% at veraison, compared to 2.5-3.5% and 4-6% respectively for high-K rootstocks under equivalent soil conditions. This restricted K acquisition reflects lower K⁺ transporter expression or activity, reduced K⁺ xylem loading efficiency, and/or smaller root system volume limiting soil K interception. Low K uptake is advantageous for premium wine production on K-adequate soils, maintaining must pH in the optimal range (3.2-3.5 for white wines, 3.4-3.7 for reds), preserving titratable acidity, and enhancing color stability through favorable pH-dependent anthocyanin equilibria.
Low_Potassium_Uptake_Rootstock_Role is associated primarily with V. riparia (Riparia Gloire) and V. riparia × V. rupestris (3309C, 101-14Mgt, Schwarzmann) parentage. The V. berlandieri × V. riparia rootstocks (SO4, 5BB, 5C) show intermediate K uptake, generally lower than the V. berlandieri × V. rupestris group. Low K uptake becomes problematic on genuinely K-deficient soils where it can exacerbate deficiency symptoms (marginal leaf scorch, poor fruit set, reduced yield), requiring K fertilization adjustment. The low-K trait is valuable in warm-climate viticulture where maintaining acidity is a persistent challenge.
K-restricting rootstock
K-schwache Unterlage
Unterlage mit geringer Kaliumaufnahme
kaliumrestriktive Unterlage
low K rootstock
low potassium uptake rootstock
potassium restricting rootstock
Mpelasoka BS et al. (2003) A review of potassium nutrition in grapevines with special emphasis on berry accumulation, Australian Journal of Grape and Wine Research 9: 154-168 Delas J, Pouget R (1979) Influence des porte-greffes sur la nutrition minérale de la vigne, Connaissance de la Vigne et du Vin 13: 169-191 Kodur S (2011) Effects of rootstocks on grapevine growth, yield and quality of wine, Journal of Wine Research 22: 155-168
Low Potassium Uptake Rootstock Role
Chardonnay exhibits low stomatal conductance disposition with maximum gs of 150-220 mmol/m²/s under well-watered conditions, supporting moderate photosynthetic rates of 10-13 µmol CO₂/m²/s. Pinot Noir and Riesling show similarly conservative conductance (180-250 mmol/m²/s), with rapid stomatal closure as water deficit develops. Low-gs cultivars typically show isohydric behavior, maintaining midday leaf water potential above -1.2 to -1.4 MPa through stomatal restriction even as soil dries. Water savings of 20-40% compared to high-gs cultivars occur under equivalent atmospheric demand, with proportionally reduced carbon assimilation of 15-30%.
A stomatal conductance disposition that is the inherited capacity of a Vitis vinifera plant to maintain relatively low rates of stomatal gas exchange (typically gs less than 250 mmol H₂O/m²/s under well-watered conditions), conferring water conservation at the cost of reduced maximum photosynthetic rate, often associated with isohydric water use behavior.
Low stomatal conductance disposition represents a conservative water use strategy prioritizing hydraulic safety over maximum productivity. The trait provides advantages in rain-fed viticulture and water-limited environments by reducing the rate of soil water depletion and maintaining favorable plant water status. However, the carbon cost of stomatal restriction may reduce yield potential under well-watered conditions. Some premium wine regions favor low-gs cultivars for their association with moderate vigor and concentrated fruit composition.
conservative stomata disposition
eingeschränkte Transpirationskapazität
low gs trait
niedrige Stomata-Leitfähigkeit
niedrige stomatäre Öffnung
restricted transpiration capacity
Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell and Environment 26:1393-1405 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data. Annals of Botany 105:661-676
Low Stomatal Conductance Disposition
In Merlot vines in Saint-Émilion, Bordeaux, low temperature constraint maintains ecodormancy from early February through late March, with mean February temperatures of 7-9°C providing strong constraint that prevents budbreak despite restored growth competence, constraint gradually weakening through March as mean temperatures increase to 10-12°C allowing growing degree day accumulation, and constraint fully relieved by late March when sustained temperatures of 12-15°C provide the heat accumulation required for budbreak initiation at BBCH stage 05.
Low temperature constraint in Vitis vinifera represents the specific thermal limitation that maintains bud quiescence during ecodormancy, whereby temperatures below the threshold required for growth-supporting metabolic activity prevent budbreak initiation in growth-competent buds until spring warming relieves this constraint. This temperature-based control operates throughout the ecodormancy period as the primary factor determining bud status, with the constraint intensity varying continuously with ambient temperature from strong constraint at low temperatures to weak or absent constraint at temperatures above the growth threshold. The biochemical basis of low temperature constraint involves the fundamental temperature-dependence of enzyme-catalyzed reactions following Arrhenius kinetics, with reaction rates declining exponentially as temperature decreases, eventually falling below levels sufficient to sustain net biosynthesis and the coordinated cellular processes required for growth and development. The threshold temperature below which constraint effectively prevents growth, termed the base temperature, is approximately 10°C for grapevine budbreak, though some variation exists among cultivars and the effective threshold may differ slightly for different component processes of budbreak. The quantification of low temperature constraint uses the concept of heat deficit, representing the cumulative shortfall of temperature below the growth-supporting threshold, with constraint being relieved as warming provides temperatures above this threshold and allows accumulation of the growing degree days required for budbreak. The diurnal and seasonal variation in temperature creates corresponding variation in constraint intensity, with constraint being strongest during cold nights and winter conditions, and weakening during warm days and as seasonal warming progresses, creating the gradual relief of constraint that eventually permits budbreak. The interaction between low temperature constraint and bud developmental status affects budbreak characteristics, with buds that have accumulated some heat units during intermittent warming being closer to the budbreak threshold and requiring less additional warming for growth initiation than buds that have experienced consistently cold conditions throughout ecodormancy.
Low_Temperature_Constraint represents the specific thermal limitation within Quiescence_Maintenance. Base temperature threshold approximately 10°C for grapevine. Constraint intensity varies continuously with temperature from strong at low temperatures to absent above threshold. Quantified through heat deficit and growing degree day concepts. Diurnal and seasonal variation affects constraint intensity. Relief through warming and heat unit accumulation. Primary determining factor for ecodormancy duration and budbreak timing.
Kälte-Limitation
Niedertemperatur-Hemmung
Tieftemperatur-Beschränkung
cold constraint
cold temperature limitation
low temperature inhibition
thermal growth constraint
thermische Wachstumsbeschränkung
Moncur MW et al (1989) Base temperatures for budbreak and leaf appearance of grapevines. American Journal of Enology and Viticulture 40: 21-26 Parker AK et al (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17: 206-216 Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle
Low Temperature Constraint
Evaluation of Low_Vigor_Disposition at the University of Burgundy characterized growth parameters in cultivars recognized for limited vegetative development. Reference cultivars 'Pinot Noir', 'Gewürztraminer', 'Aligoté', and 'Gamay' were grown under standardized conditions with detailed growth monitoring. Low_Vigor_Disposition cultivars showed consistent characteristics distinguishing them from moderate and high-vigor types. 'Pinot Noir' exhibited archetypal Low_Vigor_Disposition with mean shoot length of 115 cm (range 95-135 cm across clones), internode length of 5.4 cm, individual leaf area of 125 cm², lateral production of only 4.2 laterals per shoot with total lateral length of 32 cm, and pruning weight of 0.85 kg/vine on own roots. Growth cessation occurred in weeks 10-11 (mid-July), 4-5 weeks before veraison, indicating very short active growth period. 'Gewürztraminer' showed even lower vigor (98 cm shoots, 0.72 kg/vine pruning weight, growth cessation week 9), representing an extreme of Low_Vigor_Disposition among commercial cultivars. The Ravaz index for Low_Vigor_Disposition cultivars averaged 9.5-12.0, above the optimal range and indicating that fruit load may stress limited vine capacity. Rootstock trials demonstrated that Low_Vigor_Disposition cultivars benefited substantially from vigor-enhancing rootstocks—'Pinot Noir' pruning weight increased from 0.85 kg/vine on own roots to 1.45 kg/vine on '110 Richter' and 1.65 kg/vine on 'SO4', shifting Ravaz index into the optimal range. Spacing trials confirmed that Low_Vigor_Disposition cultivars tolerated close within-row spacing (0.9-1.0 m) without competition effects, enabling high vineyard leaf area indices despite limited per-vine growth.
A Low_Vigor_Disposition is a Shoot_Vigor_Disposition that inheres in Vitis vinifera shoots and is realized through restrained vegetative growth producing short shoots (typically < 140 cm), compact internodes (< 6 cm mean length), smaller leaves (< 150 cm² individual area), and limited lateral shoot development, resulting in open canopies that may require vigor-enhancing management or rootstock selection to achieve adequate leaf area for optimal fruit maturation. This disposition represents the lower range of grapevine vigor, characteristic of cultivars with inherently limited growth capacity that produce minimal vegetative biomass even under favorable growing conditions. Low-vigor cultivars typically show early growth cessation (well before veraison), short periods of active shoot elongation, and limited response to vigor-enhancing inputs, reflecting fundamental genetic constraints on growth capacity. The viticultural implications of Low_Vigor_Disposition are context-dependent—such cultivars may achieve excellent vegetative-reproductive balance on fertile sites or with vigorous rootstocks, but risk inadequate canopy development on limiting sites, potentially compromising fruit ripening through insufficient leaf area. Low-vigor cultivars are typically suited to close vine spacing (maximizing vineyard leaf area despite limited per-vine growth), simpler training systems (VSP, single Guyot), fertile soils, and vigor-enhancing rootstocks. The open canopies of low-vigor cultivars offer advantages for fruit zone microclimate (excellent light exposure, air circulation, reduced disease pressure) but may require irrigation management to avoid excessive water stress in the absence of shading.
Low_Vigor_Disposition is_a Shoot_Vigor_Disposition representing the lower vigor category. Typical characteristics include shoots 10 if not corrected. Examples include 'Pinot Noir', 'Gewürztraminer', 'Aligoté', 'Gamay', 'Muscadet'. Management implications include need for vigor-enhancing rootstocks on most sites, close vine spacing, simple training systems, and attention to adequate leaf area for fruit ripening. Low vigor creates favorable fruit zone microclimate but risks inadequate canopy for optimal fruit maturation.
Geringe-Wuchskraft
Niedrige-Wuchsstärke-Disposition
Schwaches-Wachstum
low growth
restrained vegetative growth
weak vigor
Howell GS (2001) Sustainable grape productivity and the growth-yield relationship: a review, American Journal of Enology and Viticulture 52:165-174 Kliewer WM, Dokoozlian NK (2005) Leaf area/crop weight ratios of grapevines: influence on fruit composition and wine quality, American Journal of Enology and Viticulture 56:170-181 Vasconcelos MC et al. (2009) The flowering process of Vitis vinifera: a review, American Journal of Enology and Viticulture 60:411-434
Low Vigor Disposition
Riparia Gloire de Montpellier (V. riparia) confers low vigor to Pinot Noir in Burgundy, with pruning weights of 0.2-0.4 kg/m and Ravaz indices of 7-10, favoring concentrated berry composition for premium Côte d'Or wines. 101-14 Millardet et de Grasset (V. riparia × V. rupestris) restricts Merlot growth in Saint-Émilion to 0.3-0.5 kg/m pruning weight on clay-limestone soils, contributing to the qualitative reputation of right-bank Bordeaux. 420A Millardet et de Grasset (V. berlandieri × V. riparia) provides low vigor to Sangiovese in Chianti Classico, limiting canopy development on fertile valley-floor alluvial soils.
A vigor-modifying rootstock role that inheres in a grafted rootstock by virtue of its capacity to reduce scion vegetative growth substantially below own-rooted or moderate-vigor benchmarks, typically decreasing pruning weight by 20-40% relative to moderate references. Low-vigor rootstocks produce scion pruning weights typically in the range of 0.2-0.4 kg/m, shorter shoot lengths of 0.6-1.2 m per season, and reduced leaf area indices of 1.5-2.5. The growth restriction is mediated by limited root system extent, lower hydraulic conductance (often below 100 mg H₂O·m⁻²·s⁻¹·MPa⁻¹), reduced cytokinin translocation, and constrained nitrogen delivery. Low-vigor rootstocks are strategically selected for fertile, deep soils and for premium wine production where restricted vegetative growth promotes fruit concentration, phenolic maturity, and balanced vine canopy architecture.
Low_Vigor_Rootstock_Role is distinct from Dwarfing_Rootstock_Role in that low-vigor rootstocks still support commercially viable yields (typically 4-8 t/ha for premium wine), whereas dwarfing rootstocks may reduce growth to economically marginal levels. Low vigor is particularly valued in appellation systems where yield restrictions and quality mandates favor smaller canopies and concentrated fruit. The V. riparia parentage is strongly associated with low vigor due to the species' naturally shallow, fine root architecture and early growth cessation.
devigorating rootstock
growth-restricting rootstock
low-vigor rootstock
schwache Unterlage
schwachwüchsige Unterlage
weak rootstock
wuchsschwache Unterlage
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 285-310 Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22 Tandonnet JP et al. (2010) Long term effects of rootstock genotype on vine growth, Australian Journal of Grape and Wine Research 16(2): 327-339
Low Vigor Rootstock Role
A low-yield primary reproductive shoot on Vitis vinifera cv. Pinot Noir from node 12 (apical position on retained cane), 118cm length with 14 leaves totaling approximately 1820 cm² leaf area, bearing single small cluster with 24 berries, estimated total shoot crop 132g at maturity, representing typical low fruitfulness of apical cane positions where late differentiation timing and apical dominance effects reduce reproductive development, this shoot retained for photosynthetic contribution (excellent 13.8 cm² leaf area per gram fruit ratio ensuring superior fruit quality from limited crop) rather than productivity, with low yield accepted as positional characteristic rather than problematic condition, contributing modestly to whole-vine crop while providing valuable leaf area supporting fruit development on more heavily-loaded mid-cane positions.
A primary reproductive shoot bearing minimal reproductive load despite primary bud origin, characterized by single small cluster, reduced berry count (15-35 berries typical), low total shoot crop (80-180g at maturity), and origin from either inherently low-fruitfulness positions (apical or basal nodes, shaded previous-season differentiation), stress-affected buds (drought, disease, nutrient deficiency during differentiation), cultivar characteristics (naturally low cluster numbers or sizes), or deliberate extreme crop reduction (aggressive cluster thinning to single small cluster per shoot). Low-yield primary reproductive shoots represent minimal productivity from individual positions, arising from various causes ranging from expected positional effects to problematic stress conditions, requiring assessment whether low yield reflects normal variation (basal positions, certain varieties) or indicates problems warranting investigation and correction.
Low-yield primary reproductive shoots arise from multiple causes with different management implications. Expected low-yield contexts include positional effects (basal nodes 1-3 and apical nodes showing characteristically reduced fruitfulness), variety characteristics (some Pinot Noir and Nebbiolo clones naturally producing small clusters), and deliberate thinning (quality-focused producers reducing to single small cluster per shoot). Problematic low-yield contexts include stress effects (severe drought, disease, nutrient deficiency during previous season's differentiation preventing normal inflorescence development), vine decline (progressive fruitfulness reduction indicating health problems), and unusual failures (mid-cane positions showing uncharacteristic low fruitfulness requiring investigation). Management depends on cause assessment= accepting expected low yields as normal variation, investigating and correcting stress-related problems, monitoring decline patterns suggesting vine replacement needs, or maintaining deliberate low yields as quality strategy. Low-yield shoots often show excellent leaf-to-fruit ratios supporting superior fruit quality despite minimal quantity.
Minimalertragstrieb
light-cropping primary shoot
low-productivity shoot
minimal-yield reproductive shoot
niedrigerträglicher primärer Fruchttrieb
Based on Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Srinivasan & Mullins (1981) "Physiology of flowering in the grapevine - a review" American Journal of Enology and Viticulture 32=47-53.
Low Yield Primary Reproductive Shoot
A lower canopy basal leaf at node 4 on a shoot of Vitis vinifera cv. Tempranillo in Rioja, Spain, emerging 12 days post-budbreak during the second week of shoot development, measuring 15.8 cm length by 14.5 cm width representing 18% larger area than median leaves at nodes 8-11, developing from preformed primordium enabling maturation at 31 days post-budbreak compared to 38-42 days for median position leaves, initially experiencing 70% full sunlight during May creating sun leaf anatomical characteristics with thick cuticle and multiple palisade layers, demonstrating peak photosynthesis rate of 12.8 μmol CO₂ m⁻² s⁻¹ in late May contributing approximately 22% of shoot photosynthate during the critical fruit set period, experiencing progressive light reduction to 35% full sun by mid-July as 14 leaves developed above creating 1.2 m² of shading canopy, showing functional decline with photosynthesis reduced to 7.5 μmol CO₂ m⁻² s⁻¹ and beginning senescence symptoms including slight chlorosis in early August, measured at shoot BBCH stage 83 (berries developing color), representing typical lower canopy basal leaf with significant early contribution and characteristic shade-induced decline.
A basal leaf positioned in nodes 3-5 occupying the lower canopy zone in Vitis vinifera, characterized by intermediate basal position following the first and second leaves but preceding median canopy development, emergence during the second week post-budbreak typically days 8-16, dimensions generally larger than median leaves often 10-25% greater area reflecting basal developmental priority, development from preformed primordia within dormant buds enabling faster maturation compared to median position leaves, achievement of full expansion typically 4-5 weeks post-budbreak, initial exposure to favorable light conditions during early development followed by progressive shading as upper canopy develops, important contribution to shoot carbon balance during weeks 4-10 post-budbreak providing 15-30% of shoot photosynthate supporting shoot extension and early fruit development, and characteristic functional trajectory showing peak performance during early to mid-season followed by progressive decline through canopy shading creating eventual senescence often occurring 2-4 weeks earlier than median canopy leaves. Lower canopy basal leaves occupy the transitional zone between the critical early-emerging first and second leaves and the later-developing median canopy, experiencing developmental advantages from preformation and early emergence but increasing vulnerability to canopy shading as shoot extension and leaf production continue through the growing season, with functional lifespan and contribution strongly influenced by canopy management practices including shoot density, positioning, and leaf removal that determine light penetration to lower zones.
Lower canopy basal leaves occupy the functional transition zone between the critical first and second leaves providing maximum early contribution and the median canopy leaves providing sustained mid to late-season function. Nodes 3-5 designation provides useful categorization though exact node ranges defining lower canopy vary with shoot vigor, cultivar, and training system. These leaves share with nodes 1-2 the developmental advantage of preformation enabling early maturation and function, but experience greater shading vulnerability as their more distal position places them deeper within developing canopy density. Functional importance peaks during mid-spring through early summer when they contribute substantially to shoot carbon balance, declining progressively as upper canopy development creates shading that reduces photosynthetic capacity below levels justifying maintenance respiration costs. Canopy management practices including shoot thinning, leaf removal, and shoot positioning critically affect lower canopy basal leaf function, with dense canopies creating severe shading that induces early senescence while open canopies maintain adequate light enabling sustained function. Lower canopy basal leaves are common targets for mid-season leaf removal aimed at improving cluster exposure and air circulation, with removal timing (typically pre-veraison) balancing fruit microclimate benefits against photosynthetic capacity loss.
Knoten drei-bis-fünf Blatt
lower zone basal leaf
nodes three to five leaf
transitional basal leaf
untere Kanopie basales Blatt
übergangs basales Weinblatt
Based on Smart (1985) "Principles of grapevine canopy microclimate manipulation with implications for yield and quality" American Journal of Enology and Viticulture 36=230-239 Poni et al. (2006) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 57=397-407 Hunter et al. (1995) "Partial defoliation of Vitis vinifera L. cv. Cabernet Sauvignon/99 Richter= Effect on root growth, canopy efficiency, grape composition, and wine quality" American Journal of Enology and Viticulture 46=306-314 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 273-304.
Lower Canopy Basal Leaf
A lower lamina surface on a mature leaf at node 6 of Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, California, positioned facing downward experiencing approximately 15% of incident irradiance after transmission through lamina and reflection from upper surface, exhibiting light gray-green coloration from dense trichome coverage creating felted appearance characteristic of this cultivar, trichome density of 52 trichomes mm⁻² on intercostal regions (between veins) composed primarily of prostrate multicellular trichomes, thinner cuticle of 3.2 μm compared to 5.8 μm on upper surface, very high stomatal density of 185 stomata mm⁻² representing 94% of total leaf stomatal population with functional stomata showing appropriate responses to environmental signals, epidermal cells with abundant intercellular spaces visible in surface view as irregular cell boundaries, prominent venation network with median vein and lateral veins appearing as pronounced raised ridges covered with dense trichomes, surface temperature typically 1-2°C below upper surface and 2-4°C below ambient during high transpiration periods due to evaporative cooling, demonstrating stomatal conductance of 245 mmol H₂O m⁻² s⁻¹ under favorable conditions enabling high photosynthetic rates, measured at BBCH stage 75 (berries pea-sized) in late June, representing typical lower surface characteristics specialized for gas exchange function.
The abaxial surface of the grapevine leaf blade facing downward away from direct incident light in Vitis vinifera, characterized by reduced light exposure and shaded position, thinner cuticle typically 2-4 μm compared to upper surface reflecting reduced water stress and UV exposure, rough or felted texture from dense trichome coverage in most cultivars, lighter green to gray-green coloration from pubescence and underlying spongy mesophyll, very high stomatal density typically 120-240 stomata mm⁻² representing 92-97% of total leaf stomata, loose epidermal cell packing with abundant intercellular spaces facilitating gas diffusion, prominent vein network appearing as raised ridges more visible than on upper surface, and functional specialization for gas exchange with anatomical features optimizing CO₂ uptake for photosynthesis and water vapor loss through transpiration. The lower lamina surface serves as the primary gas exchange interface, bearing the vast majority of functional stomata enabling leaf-atmosphere CO₂ and water vapor exchange, exhibiting adaptations for efficient gas exchange including high stomatal density, abundant intercellular air spaces, and thin cuticle minimizing diffusion resistance, showing cultivar-diagnostic pubescence characteristics including trichome density, type, and distribution patterns serving as important ampelographic features, and experiencing more moderate environmental conditions than upper surface including reduced light, lower temperatures, and higher humidity creating favorable conditions for stomatal function.
Lower lamina surface is distinguished from upper surface by functional specialization for gas exchange rather than light reception and protection, reflected in high stomatal density, reduced cuticular thickness, and abundant trichome coverage. Pubescence characteristics show extreme variation among cultivars from nearly glabrous (Sauvignon Blanc, some Chardonnay clones) to densely tomentose (Cabernet Sauvignon, Gewürztraminer), serving as critical ampelographic diagnostic features. Trichome types include prostrate non-glandular types providing physical protection and potentially reducing transpiration, erect non-glandular types providing mechanical protection, and glandular types secreting various compounds though less common in Vitis vinifera compared to some wild species. The lower surface experiences more favorable conditions for stomatal function including reduced light, lower temperatures from shading and evaporative cooling, and higher humidity from transpired water vapor accumulation, enabling sustained high stomatal conductance supporting photosynthesis. Disease organisms including powdery mildew and downy mildew show preferences for lower leaf surfaces, with humid shaded conditions and direct stomatal entry points facilitating infection. Management practices including fungicide applications often target lower surfaces requiring adequate spray coverage for effective disease control.
Abaxialseite
Blattunterseite
abaxial surface
lower leaf surface
underside lamina
untere Blattfläche
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 90-135 Patakas et al. (2003) "Differences in stomatal and photosynthetic responses of Vitis vinifera L. (cv. Sultana) leaves in response to progressive water stress" Vitis 42=113-115 Flexas et al. (2002) "Drought-inhibition of photosynthesis in C₃ plants= stomatal and non-stomatal limitations revisited" Annals of Botany 89=183-189 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 137-178.
Lower Lamina Surface
A lower lateral lobe representing the left lower lateral on a mature leaf at node 7 of Vitis vinifera cv. Syrah in the Rhône Valley, France, positioned as the second lateral lobe from apex between upper lateral and basal lobes, vascularized by the second primary lateral vein branching from median vein at 52-degree angle at 6.5 cm from petiolar junction, measuring 3.8 cm maximum width and 3.5 cm length representing smaller dimensions compared to upper lateral lobe (5.2 cm × 5.5 cm), separated from upper lateral lobe by middle left lateral sinus of 2.5 cm depth and from basal lobe by lower left lateral sinus of 1.8 cm depth, lobe apex showing obtuse configuration with approximately 68-degree angle, lobe margins exhibiting slightly convex curvature with heterogeneous teeth, lobe area of 11.2 cm² representing 14% of total blade area smaller than upper lateral (19%) but larger than basal lobe (9%), moderate development creating distinct recognizable lobe contributing to five-lobed configuration characteristic of this cultivar, measured at BBCH stage 81 (beginning of berry ripening) in early August, representing typical lower lateral lobe morphology with subordinate but distinct development.
A lateral lobe positioned in the lower intermediate zone between the upper lateral lobes and basal lobes in Vitis vinifera, characterized by position as the second pair of lateral lobes counting from apex or first pair above basal lobes, vascularization by the second pair of primary lateral veins branching from the median vein typically at 45-65 degree angles, smaller size compared to upper lateral lobes generally representing reduced but still substantial lateral lobe development, separation from upper lateral lobes by middle lateral sinuses and from basal lobes by lower lateral sinuses or transition zones depending on lobe number and configuration, variable development ranging from well-defined distinct lobes in deeply dissected cultivars to poorly developed or absent in weakly lobed varieties showing only three-lobed or weakly five-lobed configurations, and cultivar-specific characteristics including presence or absence, relative size, and degree of development serving as diagnostic features. Lower lateral lobes represent the subordinate lateral lobe pair when present, developing in five-lobed and more deeply dissected configurations but absent or weakly expressed in three-lobed leaves, exhibiting reduced size compared to upper laterals typically 12-18% per lobe of total blade area, and showing variable prominence from distinct isolated lobes to barely distinguishable projections depending on cultivar lobe dissection patterns.
Lower lateral lobes are distinguished from upper lateral lobes by inferior position and typically smaller size, and from basal lobes by intermediate rather than basal position. Presence and development vary substantially among cultivars, with deeply dissected varieties (Syrah, Mourvèdre) showing well-developed lower laterals creating clear five-lobed or seven-lobed patterns, while weakly lobed cultivars (Chardonnay, some Gewürztraminer clones) show absent or very weak lower laterals creating three-lobed configurations. Size subordination to upper laterals is typical, with lower laterals representing 60-85% of upper lateral dimensions in most five-lobed configurations. Separation from adjacent lobes depends on sinus depth, with middle sinuses separating from upper laterals showing variable depth and lower sinuses or transition zones to basal lobes often shallow or absent creating integrated lower lateral-basal lobe complexes. In seven-lobed leaves, lower lateral designation may apply to middle lateral pair with additional inferior lateral pair above basal lobes. Functional contribution involves moderate area provision typically 20-30% combined area when well developed, though contribution diminishes or disappears in weakly lobed cultivars lacking distinct lower laterals. Ampelographic assessment includes presence or absence as primary character, with degree of development, relative size, and separation characteristics serving as secondary features for cultivars showing lower lateral lobe expression.
inferior lateral lobe
inferiorer Laterallappen
lower side lobe
second lateral lobe
unterer Seitenlappen
zweiter Seitenlappen
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 136-182 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 195-218.
Lower Lateral Lobe
A lower lateral secondary inflorescence at node 5 on a shoot of Vitis vinifera cv. Grenache in Châteauneuf-du-Pape, France, measuring 11.8 cm length representing moderate secondary development, positioned in the lower shoot region receiving approximately 58% of full sunlight with substantial shading from overlying foliage and upper inflorescences, located on a vigorous shoot bearing three inflorescences (primary at node 3, this secondary at node 5, tertiary at node 7), creating lower position in the three-inflorescence sequence, exhibiting moderately developed compound paniculate architecture with central rachis bearing 3 orders of branching, rachis diameter of 2.4 mm at base representing moderate construction, bearing approximately 310 flowers, showing dimensions substantially reduced from the primary (19.2 cm, 680 flowers) representing 39% length reduction and 54% flower number reduction, positioned in established shoot region with extension growth completed during inflorescence development potentially limiting resource competition advantages, developmental stage at BBCH 61 (beginning of flowering, 10% flowers open) showing 4-day phenological delay compared to primary at stage 69 (end of flowering), lower position experiencing somewhat reduced air circulation and increased humidity potentially elevating disease risk, expected to contribute approximately 22% of shoot yield, measured in early June during early bloom, representing typical lower lateral secondary with reduced development reflecting less favorable position.
A lateral secondary inflorescence positioned in the lower portion of the shoot reproductive zone in Vitis vinifera, characterized by position at node 5 creating lower location when shoots bear multiple inflorescences at nodes 3, 5, and potentially 7, proximity to the shoot base relative to other inflorescences on shoots with 3 total inflorescences, reduced light exposure typical of lower shoot positions receiving 50-70% of full sunlight, increased shading from overlying canopy including leaves and upper inflorescences, position in established shoot regions with completed or slowing extension growth during inflorescence development, dimensions typically 10-14 cm representing moderate to reduced development for secondary inflorescences, moderate branching complexity showing 3 orders, and functional characteristics including somewhat limited resource availability from reduced photosynthetic capacity and competition with upper structures. Lower lateral secondary inflorescences represent the less favorably positioned variant of secondary inflorescences when multiple inflorescences occur, exhibiting reduced development reflecting positional disadvantages, and contributing moderately to shoot yield with potential for removal during cluster thinning operations targeting lower-quality fruit.
Lower lateral secondary inflorescences represent the less favorably positioned variant particularly on shoots bearing 3 total inflorescences, distinguished from upper lateral secondaries by lower shoot location and typically inferior development. The lower designation emphasizes the basal-proximal position relative to other secondaries or tertiaries creating less favorable conditions. Position at node 5 on three-inflorescence shoots creates the middle sequential position but functionally lower position given proximity to shoot base and distance from apex. Light exposure disadvantages from lower positioning typically provide 50-70% full sun compared to 70-90% for upper positions, with shading from overlying structures including leaves, upper primary, and potential tertiary inflorescences reducing photosynthetic capacity. Completed extension growth means lower secondaries develop during or after peak vegetative activity potentially reducing resource availability compared to actively growing upper regions. Size reduction from primaries is typically more pronounced with 30-45% reductions common compared to 10-25% for upper secondaries, reflecting cumulative positional disadvantages. Branching complexity often shows limited development with 3 orders typical compared to 4 orders for upper secondaries. Flower number reduction may be substantial with 40-60% reduction from primaries in some cases. Phenological delay is often more pronounced with 3-5 day delays typical. Yield contribution is moderate at 20-30% with lower quality potential from shading and reduced development. Management implications favor removal during cluster thinning, with lower secondaries representing preferred targets for elimination when crop reduction is needed due to their reduced size, lower position creating shading and disease vulnerability, and potentially inferior fruit quality from limited sun exposure.
basal-proximal secondary
basal-proximale Zweitinfloreszenz
beschattete laterale Sekundärrispe
lower position secondary
shaded lateral secondary
untere laterale sekundäre Infloreszenz
Based on Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 296-332 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 255-290 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 439-478.
Lower Lateral Secondary Inflorescence
A lower lateral sinus representing the left lower lateral sinus on a mature leaf at node 7 of Vitis vinifera cv. Merlot in Bordeaux, France, positioned between the upper lateral lobe and lower lateral lobe in the mid-blade region, measuring 1.8 cm depth from sinus base to lobe edge connection line, width of 1.2 cm at opening, exhibiting shallow U-shaped contour with poorly defined rounded base, sinus base positioned 5.5 cm from petiolar junction representing mid-blade location, left margin formed by lower edge of upper lateral lobe and right margin by upper edge of lower lateral lobe showing gradual transitions, sinus creating moderate separation between upper and lower lateral lobes sufficient to distinguish five-lobed configuration, sinus depth substantially less than upper lateral sinus depth (3.0 cm) on the same leaf reflecting characteristic pattern of decreasing dissection depth from upper to lower blade regions, bilateral near-symmetry with right lower lateral sinus showing similar shallow configuration (1.7 cm), measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing typical moderately developed lower lateral sinus distinguishing five-lobed from three-lobed architecture.
The sinuses positioned between lateral lobes in the lower blade region or between lower lateral lobes and basal lobes in Vitis vinifera, characterized by location in the mid to lower blade region below upper lateral sinuses, variable presence depending on cultivar lobe number with clear lower lateral sinuses in five-lobed leaves separating upper and lower lateral lobes, or transitional zones in three-lobed leaves, bilateral occurrence as paired left and right structures when present, variable depth typically 0.5-3 cm generally shallower than upper lateral sinuses, U-shaped or barely perceptible configurations, often poorly defined with gradual transitions rather than distinct sinus bases, role in separating lower lateral lobes from upper laterals or basal lobes when lobe differentiation is sufficient, and cultivar-specific characteristics including presence or absence, depth when present, and degree of definition serving as diagnostic features particularly for distinguishing three-lobed from five-lobed configurations. Lower lateral sinuses represent variable structures depending on overall lobe dissection patterns, with well-defined lower sinuses indicating five-lobed or more deeply dissected architectures while absent or very shallow lower sinuses characterize three-lobed or weakly dissected leaves, exhibiting genetic control though often less pronounced than upper sinus development, and contributing to overall blade lobe organization particularly important for distinguishing lobe number patterns.
Lower lateral sinuses are distinguished from upper lateral sinuses by position in mid to lower blade regions and typically shallower depth. Presence and development vary dramatically among cultivars with clearly defined lower lateral sinuses in five-lobed and more deeply dissected varieties, while three-lobed cultivars show absent or barely perceptible lower sinuses. The lower lateral designation is somewhat ambiguous, potentially referring to sinuses between upper and lower lateral lobes (middle lateral sinuses) or between lower laterals and basal lobes, with usage varying among ampelographic systems. Depth is typically substantially less than upper lateral sinuses on the same leaf, with common patterns showing decreasing sinus depth from upper to lower blade regions. Configuration is often poorly defined with gradual transitions rather than distinct bases, contrasting with typically well-defined upper sinuses. Absence or minimal development characterizes three-lobed cultivars where upper lateral lobes transition directly to basal lobes without intervening lower lateral lobe differentiation. Genetic control affects presence versus absence strongly, with five-lobed patterns showing consistent lower sinus expression while three-lobed patterns lack distinct lower sinuses, though depth shows more environmental sensitivity than upper sinuses. Functional implications are minimal given shallow depth and poor definition. Ampelographic utility is moderate, primarily serving to confirm five-lobed versus three-lobed configurations rather than providing cultivar-specific identification, though depth patterns combined with other features contribute to overall morphological characterization.
inferior lateral sinus
inferiore Lateralsinus
lower lobe sinus
middle lateral sinus
mittlere Lappeneinbuchtung
untere Seitenbucht
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 58-89 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 33-78.
Lower Lateral Sinus
The lower lateral sinus description of Nebbiolo documents moderately deep lower lateral sinuses (OIV 082:5) with U-shaped bases, creating clearly defined lower lobes that contribute to Nebbiolo's distinctly seven-lobed appearance — the lower lateral sinuses extend approximately 30-40% of the distance from margin to midvein, less deep than the upper lateral sinuses (OIV 070:7) but sufficient to produce true lower lobes visible as distinct projections in the leaf outline. This pronounced development of lower lateral sinuses distinguishes Nebbiolo from Barbera (very shallow to absent lower lateral sinuses, OIV 082:1-3, producing a three-to-five-lobed leaf despite similar upper sinus depth) and is one of the characters traditionally used by Piedmontese growers to distinguish the two cultivars in the vineyard.
A sinus description component that records the morphological characteristics of the lower lateral sinuses — the indentations between the upper lateral lobes and the lower lateral (basal) lobes — documenting sinus depth (OIV 082), shape, and the presence or absence of clearly defined lower lobes, with the lower lateral sinuses being typically shallower than the upper lateral sinuses and absent or indistinct in three-lobed cultivars where only the upper pair of sinuses is sufficiently developed to delineate true lobes.
Lower_Lateral_Sinus_Description documents the sinuses whose presence or absence fundamentally determines whether a leaf is classified as three-lobed (lower sinuses absent or indistinct, only the upper sinuses delineating lobes) or five-to-seven-lobed (lower sinuses sufficiently developed to create additional lobe pairs). The lower lateral sinuses show greater variability within cultivars and across environments than the upper lateral sinuses — they are typically the last sinuses to develop during leaf primordium expansion and are therefore more sensitive to developmental conditions, with leaves at the base and tip of shoots often showing reduced lower sinus development compared to mid-shoot leaves: this positional variability reinforces the importance of the OIV sampling protocol (leaves from the middle third of normally growing shoots) for reliable scoring of lower sinus characters. In cultivars where the lower lateral sinuses are present but variable in expression (the three-to-five-lobed boundary zone), the scoring requires careful assessment of multiple leaves to determine the modal state — the OIV protocol recommends observing at least 10 leaves and recording the predominant condition.
basal lateral indentation morphology
basale Seitenbuchten-Morphologie
inferior sinus description
lower lateral sinus characterization
untere Lateralsinus-Charakterisierung
untere Seitenbuchten-Beschreibung
Lower Lateral Sinus Description
In Merlot vineyards of Pomerol at BBCH 55-57, lower primary inflorescence branches (positions 3-8 from the rachis base among 16 total primary branches) measure 4.5-6.5 cm in length and bear 5-7 secondary branches each, contributing 180-240 flowers to the total inflorescence complement of 400-550 flowers. Chenin Blanc in Loire Valley (Vouvray) exhibits particularly vigorous lower primary branches of 5-8 cm bearing 6-9 secondary branches, creating characteristically broad-shouldered, conical clusters. Viticultural thinning protocols in premium Pinot Noir production sometimes remove 2-3 lower primary branches at BBCH 57-59 to reduce final cluster compactness and improve air circulation.
A lower primary inflorescence branch is a primary inflorescence branch located in the basal to middle third of the main inflorescence rachis, typically exhibiting the greatest length, diameter, and sub-branching complexity among all primary branches, thereby contributing disproportionately to total flower number and determining the basal width of the mature cluster. Lower primary branches occupy positions approximately 25-60% of the distance from the rachis base to apex (measured at BBCH 55), emerging from rachis nodes typically 8-20 mm from the point of peduncle attachment. These branches represent the earliest-initiated primary branches during inflorescence primordium development and consequently exhibit the most advanced developmental state at any given phenological stage. Branch length typically ranges from 3-8 cm (cultivar-dependent), with basal diameter often 1.5-2.5 mm, significantly exceeding upper primary branch dimensions by 30-80%. Lower primary branches characteristically bear the greatest number of secondary branches (4-8 secondary branches per lower primary branch in extensively branched cultivars like Grenache 2-4 in moderately branched types like Cabernet Sauvignon), with secondary branches themselves often extensively subdivided into tertiary branches. This architectural elaboration results in lower primary branches bearing 20-60 flowers each, representing 30-45% of total inflorescence flower number despite constituting only 25-35% of primary branch count. The elongated, well-subdivided character of lower primary branches creates the characteristic shoulder or wing structure of mature clusters in many cultivars, particularly evident in conical cluster morphologies. Vascular anatomy of lower primary branches shows larger xylem vessel diameters (35-65 μm) compared to upper branches (25-45 μm), facilitating greater water and nutrient flux to support extensive flower and subsequent berry development. Lower branch development proceeds most rapidly during BBCH 53-57, with visible elongation and secondary branch emergence 2-4 days earlier than upper primary branches under typical spring conditions. The pronounced development of lower primary branches contributes to cluster compactness in tight-clustered cultivars by creating high berry density in the basal cluster region, a critical factor in Botrytis susceptibility and the rationale for selective lower branch removal in some precision viticulture protocols.
Lower_Primary_Inflorescence_Branch is distinguished from Basal_Primary_Inflorescence_Branch by position (lower branches occupy the second quartile of rachis length while basal branches are in the first quartile nearest the peduncle) and from Upper_Primary_Inflorescence_Branch by morphology, position, and developmental timing. The 'lower' designation reflects intermediate position rather than the absolute base. These branches are critical determinants of cluster architecture and represent primary targets for cluster management interventions. The disproportionate contribution to total flower number and the advanced developmental state make lower primary branches key indicators of potential crop load. Quantitative ampelographic description requires separate measurement of lower versus upper primary branch characteristics.
basal primary branch
basaler Hauptast
lower first-order branch
unterer Hauptast der Infloreszenz
unterer Primärast
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 89-91 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 168-172 Vasconcelos, M.C., Castagnoli, S. (2000). Leaf canopy structure and vine performance. American Journal of Enology and Viticulture 51=390-396.
Lower Primary Inflorescence Branch
In Syrah vineyards of Hermitage (northern Rhône) at BBCH 55-57, lower secondary branches on lower primary branches can reach 2.5-4 cm length and bear 2-3 tertiary branches each, contributing 18-28 flowers per lower secondary and creating localized high-density zones within the characteristically long, cylindrical clusters. Grenache in Priorat exhibits particularly vigorous lower secondary branches of 3-5 cm on basal primary branches, with 3-4 tertiary branches per secondary, creating the pronounced lateral lobes of mature clusters. Some viticultural protocols selectively remove lower secondary branches from basal primary branches at BBCH 57-59 to reduce extreme cluster complexity and improve disease management.
A lower secondary inflorescence branch is a secondary inflorescence branch that arises from the basal half of a primary inflorescence branch, typically exhibiting greater length, diameter, and tertiary sub-branching compared to upper secondary branches on the same primary branch, thereby contributing disproportionately to the flower complement of that primary branch and creating localized increases in cluster density. Lower secondary branches occupy positions approximately 0-50% of the distance from the primary branch base (where it attaches to the rachis) to the primary branch apex, emerging from nodes typically within 3-15 mm of the primary branch insertion point. These branches represent the earliest-initiated secondary branches during inflorescence primordium development and consequently show more advanced developmental elaboration at any given phenological stage. Branch length typically ranges from 1-4.5 cm depending on parent primary branch vigor and cultivar type, with basal diameter often 0.6-1.5 mm, exceeding upper secondary branch dimensions by 30-60%. Lower secondary branches characteristically bear more tertiary branches (1-4 tertiary branches per lower secondary in extensively branched cultivars 0-2 in moderately branched types) compared to upper secondaries (0-2 tertiaries), and show earlier tertiary branch emergence during BBCH 55-57. This enhanced architectural elaboration results in lower secondary branches bearing 12-35 flowers each, representing 55-70% of the flower number contributed by their parent primary branch despite constituting only 40-50% of the secondary branch count on that primary. The pronounced development of lower secondary branches creates zones of high flower density on the basal portions of primary branches, particularly on lower and basal primary branches where both primary and secondary branch development are most vigorous. Vascular anatomy of lower secondary branches shows larger vessel diameters (25-50 μm) compared to upper secondaries (20-35 μm), facilitating enhanced resource flux to support extensive flower and berry development. Lower secondary branch development proceeds 1-3 days earlier than upper secondaries on the same primary during BBCH 53-57, creating temporal heterogeneity in flower development within individual primary branches. The vigorous development of lower secondary branches on basal primary branches creates the multi-lobed, compound shoulder structure characteristic of extremely loose-clustered cultivars, while simultaneously creating high-density zones in moderately branched types that may require cultural manipulation for disease management.
Lower_Secondary_Inflorescence_Branch is distinguished from Upper_Secondary_Inflorescence_Branch by position on the parent primary branch (basal half versus apical half), enhanced dimensions, earlier development, and increased tertiary subdivision. The 'lower' designation refers to position on the primary branch axis, not to position on the overall rachis (which determines primary branch classification). These branches are critical components of cluster architecture in their contribution to local density gradients and overall flower number. The interaction between primary branch position (basal, lower, upper) and secondary branch position (lower, upper) creates four-fold hierarchical complexity in flower distribution patterns that determine final cluster morphology and berry packing density.
basal secondary branch
basaler Seitenast zweiter Ordnung
proximal second-order branch
unterer Sekundärast
unterer sekundärer Blütenstandsast
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 91-93 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 170-174 Vasconcelos, M.C., Castagnoli, S. (2000). Leaf canopy structure and vine performance. American Journal of Enology and Viticulture 51=390-396.
Lower Secondary Inflorescence Branch
The lower surface hairiness of Cabernet Sauvignon documents the four-parameter trichome profile — prostrate hairs between main veins (OIV 084:5, medium density), erect hairs between main veins (OIV 085:3, sparse), prostrate hairs on main veins (OIV 086:5, medium density), erect hairs on main veins (OIV 087:3, sparse) — creating a lower surface with a visible but not dense felted covering of appressed woolly hairs interspersed with sparse bristle-like erect hairs. This pubescence profile distinguishes Cabernet Sauvignon from its parent Cabernet Franc (denser prostrate hairs, OIV 084:7, creating a more conspicuously white-felted lower surface) and its parent Sauvignon Blanc (sparser prostrate hairs, OIV 084:3, with the lower surface appearing more green than white), demonstrating that pubescence density segregates in the offspring of crosses between parents differing in trichome density.
A leaf surface description component that records the type and density of trichomes on the abaxial (lower) surface of the fully expanded leaf blade, scoring prostrate and erect hair types independently in two locations — between the main veins (OIV 084, OIV 085) and on the main veins (OIV 086, OIV 087) — each on a 1-9 density scale, providing a four-parameter trichome profile that is among the most genetically stable and diagnostically valuable character sets in the OIV descriptor system for cultivar identification.
Lower_Surface_Hairiness addresses the character complex that Galet (1998) considered second only to petiolar sinus shape in overall diagnostic value for cultivar identification. The four independent trichome parameters (two trichome types × two locations) create a 4-dimensional character space that provides exceptional cultivar discrimination — two cultivars may share the same density of prostrate hairs between veins but differ in erect hair density on the veins, or vice versa, generating distinctive pubescence textures despite superficially similar overall hairiness scores: the independent genetic control of the four parameters (documented by the segregation patterns observed in mapping populations) means that each parameter provides genuinely independent diagnostic information rather than redundantly measuring the same underlying trichome production capacity. The two trichome types are developmentally distinct — prostrate hairs originate from epidermal cell files that elongate and appressed to the leaf surface during early leaf expansion, while erect hairs originate from different epidermal cell populations that project perpendicularly from the surface — and their independent control enables the wide range of surface textures observed across cultivars, from the silky-smooth prostrate mat of Viognier to the bristly erect-dominated pubescence of some rootstock species to the glabrous surfaces of Chasselas.
Blattunterseiten-Behaarung
Blattunterseiten-Trichom-Bewertung
abaxial pubescence description
abaxiale Pubeszenz-Beschreibung
lower leaf trichome assessment
underside hairiness characterization
Lower Surface Hairiness
A lower trunk basal shoot on Vitis vinifera cv. Syrah arising from trunk at 8cm above graft union (22cm above soil surface), activated following combination of severe Eutypa dieback and winter freeze destroying all trunk tissue above this point, shoot emerged mid-June (late activation reflecting severe damage extent and resource limitation), developed to 165cm length by late August, 19 leaves, no clusters, representing extreme damage recovery from lowest viable trunk position, this shoot retained as only available renewal option despite problematic low origin requiring multi-year training to achieve target cordon height of 90cm (necessitating 68cm vertical development before lateral training possible), with low position creating management challenges including difficult access for training, soil splash exposure during establishment, and extended timeline for productive structure development (estimated 4 seasons to achieve functional bilateral cordon system versus 2-3 seasons from higher trunk positions).
A trunk basal shoot originating from lower trunk positions, characterized by emergence from trunk regions within 5-30cm of graft union or soil surface, low accessibility requiring bending or kneeling for management, proximity to ground creating potential soil splash and disease exposure, and function as last-resort trunk renewal source, extreme damage recovery mechanism, or low-positioned architectural development option when upper trunk positions unavailable or unsuitable. Lower trunk basal shoots represent minimal-height trunk recovery, arising near vine base when damage extends low on trunk or when lower latent buds activate preferentially over higher positions, providing survival mechanism following catastrophic damage though creating challenges for training due to low starting position requiring extended development period achieving normal canopy heights and potential complications from ground proximity during establishment phase.
Lower trunk basal shoots represent extreme recovery scenarios, with their low origin positions indicating severe damage extending low on trunk, preference for basal over apical bud activation (possibly reflecting resource limitation or hormonal factors), or specific anatomical factors (concentration of viable latent buds in lower trunk regions). The low position creates multiple challenges including difficult manual access (bending, kneeling required for all operations), extended training timeline (substantial vertical development needed before lateral structure possible), ground proximity complications (splash contamination, disease pressure, mechanical damage risk during cultivation), and potential aesthetic concerns (visible low renewal points versus preferred higher training positions). However, lower trunk basal shoots may represent only viable recovery option following catastrophic damage, with retention and patient multi-year training preferable to vine removal and replanting when economic analysis supports recovery investment. Management involves protecting vulnerable low shoots during establishment, patient systematic training over multiple seasons, vigilant disease management addressing ground proximity risks, and realistic timeline expectations recognizing extended development periods required for productive structure achievement.
base trunk renewal shoot
low trunk shoot
near-ground trunk shoot
stammnaher Bodentrieb
unterer Stamm-basaltrieb
Based on Mundy & Manning (2010) "Budbreak and shoot development in Vitis vinifera following severe trunk damage" Australian Journal of Grape and Wine Research 16=310-316 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 113-145.
Lower Trunk Basal Shoot
A lower trunk latent bud located 12cm above the graft union on Vitis vinifera cv. Chardonnay, deliberately activated through severe heading back of the entire trunk to force low regrowth following chronic Eutypa dieback, producing three vigorous shoots exceeding 250cm in length during the first season, with the strongest shoot selected and trained vertically to establish new trunk architecture, reaching trellis wire height by late first season, developing cordons during second season through lateral shoot selection and positioning, establishing limited fruiting wood (0.4 kg crop) in third season, and returning to full production (3.2 kg per vine) by fourth season with completely renewed trunk free from disease symptoms.
A trunk latent bud located in the lower portion of the permanent trunk structure, typically within 10-30cm above the graft union or root crown, characterized by greater distance from existing fruiting wood requiring extensive shoot training and positioning, potentially higher vigor due to more direct vascular connection to root system with fewer branch points, and lower spontaneous activation probability due to stronger apical dominance suppression from distal canopy. Lower trunk latent buds are utilized for complete trunk renewal following severe trunk disease, catastrophic freeze damage, or intentional trunk replacement programs, requiring 3-4 seasons to establish productive fruiting positions but offering advantage of completely new trunk tissue free from accumulated pathogens and with potential for extended subsequent vine longevity.
Lower trunk latent buds represent the most conservative but potentially most durable option for trunk disease management and vine renewal, essentially restarting the vine from near ground level while preserving the established root system. This approach is labor-intensive, requiring multiple seasons of shoot selection, training, and gradual fruiting wood establishment, with significant yield sacrifice during the renewal period. However, the resulting completely new trunk structure can extend productive vine life by decades in vineyards where trunk diseases are endemic. Some programs combine lower trunk renewal with improved pruning wound protection protocols to prevent reinfection of new trunk tissue.
basal trunk bud
lower trunk epicormic bud
niedrige Adventivknospe
untere Stammknospe
Based on Sosnowski et al. (2011) "Susceptibility of grapevine pruning wounds to infection by Eutypa lata" Plant Pathology 60=178-186 Úrbez-Torres et al. (2014) "Grapevine trunk diseases in British Columbia= incidence and characterization of the fungal pathogens associated with esca and Petri diseases of grapevine" Plant Disease 98=469-482 Lecomte et al. (2012) "New insights into esca of grapevine= the development of foliar symptoms and their association with xylem discoloration" Plant Disease 96=924-934.
Lower Trunk Latent Bud
A lower trunk water shoot on Vitis vinifera cv. Cabernet Sauvignon grafted on 3309C rootstock in Napa Valley, California, emerging at 48 cm height (only 8 cm above the graft union visible at 40 cm), developing in response to Eutypa dieback symptoms in the upper trunk and cordon system, growing 265 cm by late August with 24 nodes, extremely thick stem (15 mm diameter at node 12), and leaves averaging 275 cm² area with morphological characteristics (leaf shape, lobe depth, petiolar sinus form) requiring careful comparison to verified scion leaves to confirm Cabernet Sauvignon rather than 3309C origin, ultimately confirmed as scion-derived and selected for trunk renewal grafting to replace the diseased upper framework.
A trunk water shoot emerging from the lower trunk region between the graft union and the midpoint of the trunk length in Vitis vinifera, characterized by proximity to root system resources providing enhanced access to water and mineral nutrients, often showing extremely vigorous growth exceeding upper trunk positions, and representing critical management concern because shoots arising below or near the graft union may originate from rootstock rather than scion tissue, potentially leading to rootstock dominance that fundamentally alters vine characteristics including fruit quality, disease resistance, and cold hardiness. Lower trunk water shoots frequently develop in response to trunk diseases including Eutypa dieback, Esca complex, or crown gall that compromise vascular function in upper trunk regions, creating resource accumulation in lower trunk zones that triggers latent bud activation. These shoots also commonly emerge following severe winter freeze damage to upper canopy structures, mechanical damage from vineyard equipment operating near the trunk base, or intentional severe hedging that removes most photosynthetic surface area and triggers compensatory growth from the most resource-rich positions. Management of lower trunk water shoots requires careful verification of tissue origin through leaf morphology, growth habit, and if necessary genetic testing to distinguish scion from rootstock tissue before making retention decisions.
Lower trunk water shoots are distinguished from upper trunk water shoots by position below the trunk midpoint, which correlates with enhanced resource access and typically greater vigor, and by the critical importance of verifying scion versus rootstock origin. All shoots emerging below the graft union are definitionally rootstock suckers and must be removed immediately. Shoots emerging 0-20 cm above the graft union require careful evaluation, as graft union position can shift slightly through soil settling, cultivation, or trunk settling. Lower trunk water shoots selected for framework renewal require strategic management including gradual transition of photosynthetic function, maintenance of some original canopy during transition years, and patient development over 2-4 seasons to establish productive framework.
Basis-Stammtrieb
basal trunk sucker
graft-proximal water shoot
lower stem water shoot
pfropfnaher Wassertrieb
unterer Stammwasserschoss
Based on Pearson & Goheen (1988) "Compendium of Grape Diseases" APS Press pp. 56-71 Sosnowski et al. (2007) "Trunk diseases of grapevine= Eutypa dieback" Australian Viticulture 11=44-48 Úrbez-Torres (2011) "The status of Botryosphaeriaceae species infecting grapevines" Phytopathologia Mediterranea 50=S5-S45 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 412-438.
Lower Trunk Water Shoot
Lyre suitability quality comparison in Bordeaux evaluated Cabernet Sauvignon performance under lyre versus traditional vertical training on vigorous rootstock-site combinations where single-canopy VSP proved inadequate for vine balance management. Cabernet Sauvignon demonstrated excellent lyre suitability quality with adequate vigor filling both inclined curtains to target dimensions, moderately upright growth habit adapting well to the 60° angled training planes, and fruit in the open central zone achieving superior exposure with PAR levels of 35% ambient compared to 12% in dense VSP canopies on the same sites. Wine quality assessment showed 22% higher anthocyanin concentration and improved tannin structure in lyre-trained fruit, while yield per hectare increased 28% through the doubled canopy surface area. Merlot showed reduced lyre suitability quality with tendency toward excessive vigor creating curtain bridging that required monthly intervention to maintain separation, though still outperforming single-canopy alternatives on the vigorous sites.
Lyre suitability quality is a trellis adaptation quality that inheres in a grapevine cultivar or clone and represents the degree of compatibility with lyre training systems characterized by division of the canopy into two inclined planes arranged in V-configuration with shoots trained upward and outward from a central trunk to create separated canopy curtains that increase exposed surface area while positioning fruit in the open central zone between the diverging canopy walls. This quality determines whether a cultivar possesses the vigor capacity, growth habit flexibility, and fruiting characteristics necessary to exploit the enhanced light distribution and fruit exposure benefits of lyre geometry while maintaining canopy organization across the more complex divided architecture. The ontological significance of lyre suitability quality lies in its role enabling application of canopy division principles to upright-growing cultivars that prove unsuited to pendant systems like Geneva Double Curtain, providing an alternative divided canopy option that accommodates the natural upward growth tendency of many premium wine grape cultivars while achieving the vigor management and light distribution benefits of canopy separation. The biological requirements for high lyre suitability quality include vigor sufficient to fill two inclined canopy planes with adequate shoot density, growth habit amenable to training at intermediate angles between vertical and horizontal, structural shoot characteristics enabling maintenance of angled orientation without excessive bending or breakage, moderate lateral development contributing to canopy fill without bridging across the inter-curtain gap, and cluster architecture appropriate for the highly exposed central fruit zone environment. Unlike GDC which requires trailing growth habit, lyre systems accommodate cultivars with upright or intermediate habits by training shoots along the angled planes rather than requiring pendant orientation, broadening the range of cultivars suitable for divided canopy management. The lyre configuration creates unique microclimate conditions with fruit positioned in the open zone between canopy walls receiving reflected light from both curtain surfaces while benefiting from excellent air circulation, requiring cultivar fruit characteristics tolerant of or benefiting from high exposure intensity. Assessment of lyre suitability quality involves evaluation of curtain fill and angle maintenance, central zone fruit exposure characteristics, canopy separation sustainability, and yield-quality outcomes compared to alternative training approaches.
Lyre_Suitability_Quality is_a Trellis_Adaptation_Quality, representing compatibility with the V-shaped divided canopy configuration. This class addresses the specific requirements of lyre systems including adequate vigor for two curtains, adaptability to angled training, and fruit tolerance of high-exposure central zone positioning. Distinguished from GDC_Suitability_Quality by the upward-inclined rather than pendant shoot orientation, making lyre appropriate for upright-growing cultivars unsuited to GDC. The lyre represents the European adaptation of divided canopy principles for Vitis vinifera cultivars.
Lyra-Eignung
Lyraform-Kompatibilität
U-Erziehungs-Eignung
V-Erziehung-Tauglichkeit
V-trellis suitability
divided V-canopy compatibility
lyre system fitness
lyre training adaptation
Carbonneau A (1985) The early selection of grapevine rootstocks for resistance to drought conditions. American Journal of Enology and Viticulture 36:195-198 Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 32-43 Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review. American Journal of Enology and Viticulture 60:251-268
Lyre Suitability Quality
A Merlot vine in fertile Bordeaux alluvial soils trained to lyre bears a Lyre_Trained_Vine_Role, with the divided canopy managing vigorous growth that would create excessive shading in single-curtain VSP—the two separated fruit zones receiving improved light exposure for phenolic development. A Sauvignon Blanc vine in New Zealand Marlborough serves in a Lyre_Trained_Vine_Role, the divided architecture addressing the variety's vigorous growth on fertile soils while maintaining the vertical shoot positioning suited to aromatic white wine production. A Cabernet Franc vine on deep Loire Valley soils bears a Lyre_Trained_Vine_Role, the U-configuration enabling quality production from sites too vigorous for conventional single-curtain training.
A training system role that inheres in a grapevine trained to the lyre (or U) system, a divided canopy architecture with two vertical shoot-positioned curtains arising from a single vine, separated by an open center that improves light penetration and air circulation compared to single-curtain systems. Lyre training positions fruiting zones on the outer faces of two upward-growing canopy walls, creating a U-shaped cross-section (hence "lyre" from the musical instrument's form) that doubles effective canopy surface area while maintaining the vertical shoot orientation valued for quality production. The system addresses the fundamental challenge of vigorous vines—excessive vegetative growth creating dense, shaded canopies with poor fruit quality—by dividing the canopy into two thinner curtains with improved interior light exposure. Lyre training requires specialized trellis infrastructure with two sets of catch wires spreading outward from a central trunk, typically at 30-45° angles from vertical, creating the distinctive V-shaped post profile. Developed by Alain Carbonneau at INRA Bordeaux in the 1980s, lyre training represents science-based canopy management addressing vigor-quality relationships through architectural intervention.
Lyre_Trained_Vine_Role captures Carbonneau's divided canopy innovation designed specifically for high-vigor situations where single-curtain systems produce excessive shade. The system's advantages—doubled canopy surface, improved light distribution, better spray penetration—come with increased infrastructure cost and management complexity compared to simpler systems. Lyre adoption has been modest compared to simpler alternatives (VSP, bilateral cordon), limited by the specialized trellis requirements and the labor needed to maintain two separate canopy curtains. The system represents the quality-focused end of divided canopy approaches, maintaining upward shoot growth valued for premium production while addressing vigor challenges. Climate change may increase lyre relevance as warming temperatures accelerate vegetative growth, though the system's complexity continues to limit widespread adoption.
Leierform
Lyra-Erziehung
U training
U-Erziehung
divided canopy lyre
geteilte Laubwand
lyre training
lyre trellis
Carbonneau A (1980) Recherche sur les systèmes de conduite de la vigne, Connaissance de la Vigne et du Vin 14: 110-130 Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems, American Journal of Enology and Viticulture 60: 251-268
Lyre Trained Vine Role
Investigation of resistance-breaking M_Arenaria populations threatening M_Arenaria_Resistance_Disposition at UC Davis documented emergence of virulent nematodes in California vineyards planted continuously to resistant rootstocks. A population recovered from declining 'Freedom' vines in Fresno County (designated MA-FR) was compared with a standard laboratory M. arenaria isolate (MA-Lab) in inoculation assays on a rootstock panel. On the normally resistant 'Freedom', MA-Lab induced minimal galling (index 1.2) and reproduction (RF 0.8), confirming resistance. However, MA-FR overcame resistance, producing severe galling (index 7.5) and high reproduction (RF 22.5), demonstrating loss of effective M_Arenaria_Resistance_Disposition against this adapted population. Crucially, the alternative rootstock 'RS-3' (V. champinii × V. rupestris selection) maintained resistance against both populations (galling <1.5, RF <0.5), indicating that virulence to 'Freedom' did not confer broad resistance-breaking capacity. Molecular analysis suggested that MA-FR represented selection of pre-existing virulent genotypes within the original population rather than true resistance breakdown. These findings emphasize that M_Arenaria_Resistance_Disposition requires ongoing monitoring and management, with recommendations including rotation among rootstocks with different resistance backgrounds and integration of non-host cover crops to suppress nematode populations between vineyard cycles.
An M_Arenaria_Resistance_Disposition is a Meloidogyne_Resistance_Disposition that inheres in a Vitis plant root system and is realized through defense processes that specifically limit infection, giant cell formation, gall development, or reproduction by Meloidogyne arenaria (peanut root-knot nematode), a species of particular concern in viticulture due to documented occurrences of resistance-breaking populations that overcome resistance effective against other Meloidogyne species. This disposition represents resistance to a nematode that, while generally less prevalent than M. incognita in vineyards, poses unique challenges through its demonstrated capacity to adapt to host resistance. M. arenaria populations virulent on previously resistant rootstocks have been identified in California and other regions, raising concerns about resistance durability and necessitating integrated management strategies beyond sole reliance on host resistance. The species includes two races with distinct host ranges, and additional pathotypes may exist with specific virulence patterns. Resistance mechanisms against M. arenaria parallel those effective against other Meloidogyne species, involving recognition and hypersensitive response to attempted giant cell induction, but the genetic basis may differ, and resistance effective against M. incognita does not guarantee M. arenaria resistance. The emergence of resistance-breaking M. arenaria populations highlights the importance of resistance gene stewardship, including rotation among rootstock resistance sources, avoiding planting resistant rootstocks continuously on infested sites, and combining host resistance with cultural and biological control tactics. Breeding for durable M_Arenaria_Resistance_Disposition requires screening against aggressive field-derived populations rather than laboratory-maintained isolates that may have lost virulence.
M_Arenaria_Resistance_Disposition is_a Meloidogyne_Resistance_Disposition. M. arenaria presents unique challenges due to documented resistance-breaking populations, distinguishing it from M. incognita and M. javanica where resistance has generally proven more durable. The emergence of virulent populations highlights the evolutionary potential of nematode populations under continuous selection by resistant rootstocks. Management recommendations include avoiding monoculture of single resistance sources, monitoring nematode populations for virulence shifts, and maintaining integrated management approaches rather than relying solely on host resistance.
Erdnuss-Wurzelgallennematoden-Resistenz
M. arenaria immunity
M.-arenaria-Abwehr
M.-arenaria-Resistenz-Disposition
peanut root-knot nematode resistance
resistance-breaking RKN resistance
Anwar SA, McKenry MV (2002) Developmental response of a resistance-breaking population of Meloidogyne arenaria on Vitis spp., Journal of Nematology 34:28-33 McKenry MV, Anwar SA (2006) Nematode and grape rootstock interactions including an improved understanding of tolerance, Journal of Nematology 38:312-318 Ferris H et al. (2012) Reflections on plant and soil nematode ecology, Journal of Nematology 44:115-126
M Arenaria Resistance Disposition
Genetic mapping of M_Incognita_Resistance_Disposition at UC Davis utilized segregating populations from crosses involving the resistant rootstock 'Harmony' (1613C × 'Dog Ridge') to identify QTLs conditioning resistance. Phenotyping employed controlled inoculation with M. incognita race 3 (5,000 eggs per pot), with galling index and egg production assessed at 90 days. Resistance segregated as a dominant trait, with QTL analysis identifying a major locus on chromosome 9 explaining 65% of phenotypic variance for galling and 72% for reproduction, designated MiR1. Resistant individuals (MiR1/MiR1 or MiR1/mir1) exhibited galling indices of 0.5-2.0 and egg production below 1,000 per gram root, while susceptible individuals (mir1/mir1) showed galling of 7-9 and reproduction exceeding 30,000 eggs/g. Histopathological comparison revealed that J2 penetration occurred equally across genotypes, but resistant roots showed massive cell death surrounding attempted feeding sites by 72 hours post-penetration, with complete failure of giant cell establishment. The MiR1-linked marker VMC5g7 enables marker-assisted selection with 94% accuracy, accelerating introgression of M_Incognita_Resistance_Disposition into adapted rootstock backgrounds. Field trials confirmed that MiR1-positive selections maintained resistance over 8 years with no evidence of resistance-breaking nematode adaptation.
An M_Incognita_Resistance_Disposition is a Meloidogyne_Resistance_Disposition that inheres in a Vitis plant root system and is realized through defense processes that specifically limit infection, giant cell formation, gall development, or reproduction by Meloidogyne incognita (southern root-knot nematode), the most widespread and economically important root-knot nematode species affecting viticulture in warm-climate regions worldwide. This disposition represents resistance to a polyphagous, highly adaptable nematode that reproduces parthenogenetically, enabling rapid population buildup on susceptible hosts, with four recognized races (races 1-4) differing in host range and virulence. M. incognita thrives in sandy, warm soils characteristic of many premium wine regions including California, Australia, South Africa, and Mediterranean countries, making resistance essential for sustainable production in these areas. Resistance mechanisms preventing M. incognita damage include hypersensitive response at penetration and migration sites, incompatibility preventing giant cell induction or maintenance specifically in response to M. incognita effector proteins, and root architectural or biochemical features limiting J2 establishment. The genetics of M. incognita resistance in Vitis involves multiple loci, with resistance sources identified in several species including V. champinii, V. rufotomentosa, and interspecific hybrids. The N gene from V. champinii confers resistance to M. incognita races 1 and 3, while additional factors provide broader coverage. Given the prevalence of M. incognita in global viticulture, M_Incognita_Resistance_Disposition is the primary root-knot nematode resistance objective in rootstock breeding programs, with multiple resistant commercial rootstocks available.
M_Incognita_Resistance_Disposition is_a Meloidogyne_Resistance_Disposition. M. incognita is the most widespread root-knot nematode in viticulture, with four races differing in virulence. Resistance may be race-specific, with some rootstocks resistant to races 1 and 3 but susceptible to race 2. The MiR1 locus from V. champinii derivatives provides strong resistance to common races. M. incognita resistance is often correlated with but not identical to M. javanica resistance, as different recognition systems may be involved.
M. incognita immunity
M.-incognita-Abwehr
M.-incognita-Resistenz-Disposition
Mi-RKN resistance
Südlicher-Wurzelgallennematoden-Resistenz
southern root-knot nematode resistance
Cousins P, Walker MA (2002) Genetics of resistance to Meloidogyne incognita in crosses of grape rootstocks, Theoretical and Applied Genetics 105:802-807 Ferris H et al. (2012) Reflections on plant and soil nematode ecology, Journal of Nematology 44:115-126 Anwar SA, McKenry MV (2002) Developmental response of a resistance-breaking population of Meloidogyne arenaria on Vitis spp., Journal of Nematology 34:28-33
M Incognita Resistance Disposition
Comparative evaluation of M_Javanica_Resistance_Disposition versus M. incognita resistance at the Agricultural Research Organization, Volcani Center, Israel, tested 25 rootstock genotypes against both species in parallel pot assays to characterize resistance specificity. Roots were inoculated with 5,000 J2 of either M. javanica (local isolate from Negev vineyards) or M. incognita race 1, with galling and reproduction assessed at 90 days. Most rootstocks showed correlated responses, with 'Freedom', '1103 Paulsen', and 'Dog Ridge' exhibiting broad Meloidogyne resistance (galling index <2.0 for both species). However, differential responses occurred in some genotypes: '140 Ruggeri' showed strong M. incognita resistance (galling 1.5, reproduction factor RF 0.3) but moderate M. javanica susceptibility (galling 5.8, RF 8.2). Conversely, 'Ramsey' exhibited M_Javanica_Resistance_Disposition (galling 2.2, RF 1.5) but M. incognita susceptibility (galling 6.5, RF 15.3). These differential responses demonstrate that M_Javanica_Resistance_Disposition can be genetically distinct from M_Incognita_Resistance_Disposition, requiring species-specific screening when selecting rootstocks for regions where M. javanica predominates. For Israeli viticulture conditions, 'Freedom' and '1103 Paulsen' were recommended as providing reliable protection against the local M. javanica-dominated nematode complex.
An M_Javanica_Resistance_Disposition is a Meloidogyne_Resistance_Disposition that inheres in a Vitis plant root system and is realized through defense processes that specifically limit infection, giant cell formation, gall development, or reproduction by Meloidogyne javanica (Javanese root-knot nematode), a thermophilic species with worldwide distribution in tropical and subtropical viticultural regions. This disposition represents resistance to a nematode that, like M. incognita, reproduces by obligate mitotic parthenogenesis and attacks a broad host range, but shows somewhat distinct distribution patterns, with M. javanica often predominating in warmer climates and heavier soils than typically favored by M. incognita. M. javanica is particularly prevalent in Australian, South African, Israeli, and Mediterranean basin viticulture, where it can cause severe damage to ungrafted V. vinifera and susceptible rootstocks. Resistance mechanisms may partially overlap with M. incognita resistance, as both species induce similar giant cell feeding sites and may be recognized by common plant surveillance systems. However, species-specific resistance also exists, with some rootstocks showing differential responses to M. javanica versus M. incognita, necessitating specific screening and selection for M. javanica resistance where this species predominates. The genetics of M. javanica resistance involves loci that may be distinct from or allelic to M. incognita resistance factors, with some resistant rootstocks effective against both species while others show species specificity. Field documentation of resistance stability over time is critical, as Meloidogyne populations can adapt to resistance under continuous selection pressure.
M_Javanica_Resistance_Disposition is_a Meloidogyne_Resistance_Disposition. M. javanica is the second most important root-knot nematode species in global viticulture, with particular prevalence in Australia, South Africa, and the Middle East. Resistance may overlap with M. incognita resistance through shared recognition mechanisms, but species-specific resistance also exists. Rootstock selection should be based on the predominant Meloidogyne species at the planting site, as differential resistance can result in resistance failure if species composition is not correctly identified.
Javanese root-knot nematode resistance
Javanischer-Wurzelgallennematoden-Resistenz
M. javanica immunity
M.-javanica-Abwehr
M.-javanica-Resistenz-Disposition
Mj-RKN resistance
McKenry MV, Anwar SA (2006) Nematode and grape rootstock interactions including an improved understanding of tolerance, Journal of Nematology 38:312-318 Walker GE (1997) Effects of Meloidogyne javanica and organic amendments on grapevine root systems, Australian Journal of Grape and Wine Research 3:33-42 Ferris H et al. (2012) Reflections on plant and soil nematode ecology, Journal of Nematology 44:115-126
M Javanica Resistance Disposition
Maceration protocol for Cabernet Sauvignon from Coonawarra (premium tier, 2023 vintage) specifies 100% destemming and partial berry sorting, 4-day pre-fermentation cold soak at 6°C in closed stainless steel fermenters (10 t capacity) with twice-daily gentle pump-over for color extraction without seed tannin release, co-fermentative maceration of 12 days at 26-30°C with one pump-over and one punch-down daily (30-minute pump-over volume equivalent to 0.8× tank volume), post-fermentation extended maceration of 7 days at 22°C with reduced pump-over frequency (once daily, gentle), pressing at day 23 total skin contact using a membrane press (0.2-0.8-1.2-1.6 bar progressive cycle), and separation of free-run wine (65% volume, highest quality), light press (20%, blending component), and heavy press (15%, excluded from premium program). Maceration protocol for rosé of Grenache from Tavel specifies direct pressing of destemmed fruit (4-hour skin contact at 10°C for color extraction), juice settling at 6°C for 18 hours, and fermentation of clear juice without further skin contact — the brief maceration targeting the pale salmon-copper color and fresh red fruit character that defines Tavel rosé.
Maceration protocol is a winemaking protocol entity that documents the parameters and management practices governing the contact between grape juice or must and solid grape tissue (skins, seeds, and optionally stems) during winemaking, encompassing the duration, temperature, and mechanical regime of pre-fermentation cold soak, co-fermentative maceration, and post-fermentation extended maceration, the cap management method (pump-over, punch-down, rack-and-return, rotary fermenter, submerged cap), the decision regarding whole-cluster versus destemmed fruit inclusion, and the pressing protocol (timing, pressure curve, free-run and press fraction separation), providing the extraction management framework that determines the phenolic concentration, tannin structure, color depth, and textural mouthfeel of red, rosé, and skin-contact white wines.
Maceration protocol is arguably the most quality-determinant processing variable for red wine production, as the extraction of phenolic compounds from skin and seed tissue during maceration establishes the wine's color intensity, tannin concentration, mouthfeel structure, and aging potential — parameters that cannot be subsequently added or fundamentally altered after pressing. The temporal dynamics of extraction follow a well-characterized sequence — anthocyanins are extracted rapidly during the first 3-5 days (reaching 60-80% of maximum extraction by day 5), while tannin extraction is more gradual and progressive, continuing throughout maceration with skin tannins dominating early extraction and seed tannins becoming increasingly important after day 10-14 as ethanol levels rise and soften seed coat permeability. This differential extraction kinetics creates the fundamental maceration management dilemma — extended maceration enhances tannin extraction and structural complexity but risks excessive seed tannin contribution producing harsh, astringent wines if berry and seed maturity are insufficient. The cap management method profoundly influences extraction intensity and selectivity — punch-down (pigeage) provides vigorous but brief mixing events that favor skin extraction, pump-over (remontage) provides gentler but more sustained contact, rack-and-return (délestage) provides aggressive extraction through complete liquid drainage followed by resubmersion, and submerged cap systems provide continuous gentle extraction through maintained liquid contact. The whole-cluster decision (fermenting with intact stem rachis) adds stem-derived tannin (proanthocyanidins with high epigallocatechin content), aromatic compounds (spice, herbal notes from stem aldehydes), and structural modification through potassium extraction that raises pH — an approach traditionally associated with Burgundy Pinot Noir and increasingly adopted for Syrah, Grenache, and other varieties. The pressing protocol concludes the maceration phase and determines the final phenolic and aromatic composition through the separation of free-run wine (lowest tannin, highest quality) from progressive press fractions of increasing phenolic concentration and astringency.
Extraktionsprotokoll
Maischemanagement
Maischestandzeit-Protokoll
Mazerationsführung
Mazerationsprotokoll
cuvaison protocol
extraction protocol
maceration management
maceration parameters
skin contact protocol
Sacchi KL et al. (2005) A review of the effect of winemaking techniques on phenolic extraction in red wines. American Journal of Enology and Viticulture 56(3): 197-206 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1, 2nd Ed, Wiley, pp.: 327-395 Casassa LF, Harbertson JF (2014) Extraction, evolution, and sensory impact of phenolic compounds during red wine maceration. Annual Review of Food Science and Technology 5: 83-109
Maceration Protocol
The macroclimate description for the Marlborough wine region (South Island, New Zealand, 41.5°S) documents a cool-maritime climate — mean annual temperature 12.5°C, growing season (October-April) mean 16.0°C, Winkler index 1,050-1,200 GDD (Region I, the coolest viticultural category), Huglin index 1,600-1,800 (cool viticultural climate), annual precipitation 650 mm with 300-350 mm during the growing season, annual sunshine hours 2,400-2,500 (among the highest in New Zealand due to the inter-montane rain shadow position between the Richmond and Wither Hills ranges), and a high diurnal temperature range (10-14°C amplitude) created by the continental-like influence of the sheltered Wairau Valley. This cool-maritime macroclimate with high radiation and wide diurnal amplitude provides the specific thermal environment that produces Marlborough's internationally recognized Sauvignon Blanc style — the cool temperatures preserve the methoxypyrazine and thiol aroma precursors that would be degraded in warmer climates, while the high radiation promotes photosynthesis and flavor precursor synthesis, and the wide diurnal amplitude preserves acidity through reduced nocturnal malic acid respiration.
A climate data component that documents the regional-scale atmospheric conditions characterizing the broad geographic area in which the vineyard is located — typically defined by weather station data averaged over 30-year climatological normals — recording the annual and monthly patterns of temperature, precipitation, radiation, and humidity that determine the fundamental viticultural potential of the region, the applicable climate classification (Köppen-Geiger, Winkler, Huglin, Smart-Gladstones), and the position of the region within the global hierarchy of viticultural climate zones from cool to hot.
Macroclimate_Description provides the regional atmospheric context within which all site-specific and canopy-level climatic variation occurs — the macroclimate sets the boundaries of viticultural possibility (determining which cultivar maturity groups can ripen, the baseline disease pressure regime, and the fundamental wine style potential), while the mesoclimate and microclimate modify these regional conditions within their inherent range. The characterization of viticultural macroclimate has evolved from simple temperature summation indices (the Winkler growing degree-day system published in 1944, dividing California into five Regions by cumulative GDD above 10°C) through more sophisticated bioclimatic indices (the Huglin heliothermal index incorporating daylength adjustment, the Gladstones mean temperature of the warmest month, the Smart-Dry aridity/humidity indices) to the modern multivariate climate classification systems that integrate temperature, precipitation, radiation, and continentality into comprehensive viticultural climate typologies: each classification system captures different aspects of the complex climate-vine interaction, and the VVO documents multiple classification results for each site to provide complementary perspectives on viticultural climate suitability. The 30-year climatological normal (currently the 1991-2020 reference period) is the standard temporal framework for macroclimate characterization, but climate change is progressively decoupling the historical normals from current and future conditions — Jones et al. (2005) documented a mean growing season temperature increase of 1.3°C across major viticultural regions between 1950 and 2000, and projections indicate further increases of 1-3°C by 2050, necessitating periodic reassessment of regional climate classifications.
Makroklima-Beschreibung
Regionalklima-Charakterisierung
großräumige Klima-Dokumentation
macroclimate record
regional climate characterization
viticultural climate documentation
Winkler AJ et al. (1974) General Viticulture, University of California Press Huglin P (1978) Nouveau mode d'évaluation des possibilités héliothermiques d'un milieu viticole, Comptes Rendus de l'Académie d'Agriculture de France 64: 1117-1126 Jones GV et al. (2005) Climate change and global wine quality, Climatic Change 73: 319-343 Gladstones J (1992) Viticulture and Environment, Winetitles Smart RE and Dry PR (1980) A climatic classification for Australian viticultural regions, Australian Grapegrower and Winemaker 196: 8-16
Macroclimate Description
Macronutrient content quality assessment in Finger Lakes Riesling reveals the cool-climate, shale-derived soil signature, with relatively low potassium (900-1300 mg/L) and adequate calcium (80-150 mg/L) producing naturally balanced must pH (3.0-3.3) that requires no acidification for quality sparkling wine production this favorable macronutrient balance, combined with excellent malic acid retention, establishes the Finger Lakes as a premium cool-climate region where soil-derived mineral composition supports the fresh, age-worthy Riesling style.
Macronutrient_Content_Quality encompasses the chemical qualities characterizing the concentration of major mineral elements required by grapevines in relatively large quantities for growth, metabolism, and fruit development. The macronutrients addressed in this category include potassium, calcium, magnesium, phosphorus, and sulfur, which together with nitrogen (addressed separately) constitute the primary mineral nutrient pool. These elements serve essential structural and metabolic functions: potassium regulates osmotic balance, enzyme activation, and sugar transport calcium provides cell wall structure and membrane stability magnesium centers the chlorophyll molecule and activates numerous enzymes phosphorus participates in energy transfer and nucleic acid synthesis and sulfur is incorporated into amino acids and secondary metabolites. In grape berries, macronutrient content reflects the balance between phloem and xylem supply during berry development, with potassium accumulating primarily via phloem alongside sugars while calcium arrives predominantly through xylem transpirational flow during the pre-véraison period. The total macronutrient concentration in grape juice typically ranges from 2-4 g/L, dominated by potassium (1-3 g/L) with smaller contributions from phosphorus (100-400 mg/L), calcium (50-200 mg/L), magnesium (50-150 mg/L), and sulfur (50-200 mg/L). Macronutrient content varies substantially with cultivar, rootstock, soil composition, and viticultural practices, particularly irrigation and fertilization regimes. Deficiency or excess of any macronutrient produces characteristic symptoms and can limit yield or quality, making macronutrient management a core viticultural concern. The macronutrient status of must affects fermentation through effects on yeast nutrition and wine stability through mineral precipitation and pH effects.
Macronutrient_Content_Quality serves as the parent class for individual macronutrient elements, positioned within Mineral_Content_Quality. The classification follows plant nutrition conventions where macronutrients are distinguished from micronutrients by the quantities required rather than by functional importance—trace elements may be equally essential but are needed in much smaller amounts. Nitrogen, while technically a macronutrient, is addressed separately under Nitrogen_Content_Quality due to its unique role in fermentation and its occurrence in multiple organic and inorganic forms requiring distinct analytical approaches.
Hauptmineralgehalt
Makronährstoffgehalt
Makronährstoffzusammensetzung
essential mineral quality
essentielle Mineralqualität
macronutrient composition
major mineral content
primary nutrient quality
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 341-385 Christensen P (2000) Use of Tissue and Soil Analysis in Viticulture. In: Christensen LP (ed) Raisin Production Manual, University of California Agriculture and Natural Resources Publication 3393 pp.: 102-110
Macronutrient Content Quality
Macronutrient deficiency symptom quality survey across diverse vineyard soils in the Willamette Valley characterized the prevalence and distribution of major element limitations affecting regional wine grape production. Systematic visual assessment of macronutrient deficiency symptom quality identified nitrogen deficiency as the most prevalent macronutrient limitation, affecting 35% of surveyed blocks with symptoms including pale green canopy color, reduced shoot length, and early leaf senescence, concentrated in blocks with high vigor cover crops competing for available nitrogen. Potassium macronutrient deficiency symptom quality affected 18% of blocks, with the characteristic marginal leaf scorch appearing post-véraison when fruit potassium demand peaked, concentrated on shallow soils derived from marine sediments with inherently low potassium content. Phosphorus macronutrient deficiency symptom quality was rare (3% of blocks) due to generally adequate soil phosphorus in the region, with symptoms limited to newly planted vineyards on acidic forest soils where phosphorus availability was reduced by aluminum toxicity. Magnesium macronutrient deficiency symptom quality appeared in 12% of blocks as interveinal chlorosis on older leaves, associated with high-potassium soils where K-Mg antagonism reduced magnesium uptake. The macronutrient deficiency symptom quality survey established regional fertilization priorities emphasizing nitrogen management in vigorous sites and potassium supplementation on shallow sedimentary soils.
Macronutrient deficiency symptom quality is a nutrient deficiency indicator quality that inheres in a grapevine organism and represents the visual symptoms and growth abnormalities resulting from inadequate availability of the primary macronutrients nitrogen, phosphorus, and potassium, or the secondary macronutrients calcium, magnesium, and sulfur, with macronutrient deficiencies typically producing dramatic whole-plant effects on growth, vigor, and productivity reflecting the large quantities of these elements required for structural and metabolic functions. This quality addresses the deficiency symptoms of elements required in largest quantities by grapevines, recognizing that macronutrient limitations are among the most common nutritional problems affecting vineyard productivity and that the substantial tissue concentrations of these elements make deficiency symptoms typically more pronounced and widespread than micronutrient deficiencies. The ontological significance of macronutrient deficiency symptom quality lies in the frequency and economic importance of macronutrient limitations in commercial viticulture, with nitrogen particularly affecting vine vigor and fruit composition, potassium critically important for fruit quality and winter hardiness, and the secondary macronutrients essential for specific structural and metabolic functions whose deficiency can severely limit productivity. The biological basis of macronutrient deficiency symptoms encompasses the fundamental roles of these elements in plant structure and metabolism, with nitrogen constituting amino acids, proteins, and nucleic acids essential for all growth processes phosphorus required for energy metabolism, membrane structure, and genetic material potassium essential for osmotic regulation, enzyme activation, and carbohydrate metabolism calcium required for cell wall structure and membrane function magnesium central to chlorophyll and enzyme function and sulfur required for protein structure and specialized metabolites. The mobility classification of macronutrients creates characteristic symptom distributions, with the mobile nutrients N, P, K, and Mg showing deficiency symptoms first on older leaves while the immobile Ca shows symptoms on youngest tissues and growing points.
Macronutrient_Deficiency_Symptom_Quality is_a Nutrient_Deficiency_Indicator_Quality, serving as parent class for Nitrogen_Deficiency_Symptom_Quality and Phosphorus_Deficiency_Symptom_Quality (and eventually K, Ca, Mg, S) which address specific macronutrient deficiencies. This class addresses the general characteristics of macronutrient limitation while child classes capture element-specific symptoms. Macronutrients required in large quantities (g/kg tissue) creating pronounced deficiency symptoms. Primary macronutrients (N, P, K) versus secondary (Ca, Mg, S) reflect relative requirement quantities. Mobility determines symptom distribution pattern.
Hauptnährstoffmangel
Makronährstoffmangel-Symptome
NPK deficiency indicators
NPK-Mangel-Indikatoren
macronutrient stress symptoms
major element deficiency
primary nutrient deficiency
primärer Nährstoffmangel
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication Mullins MG et al. (1992) Biology of the Grapevine. Cambridge University Press, pp. 174-195
Macronutrient Deficiency Symptom Quality
Macronutrient uptake follows characteristic seasonal patterns, with nitrogen uptake peaking during rapid shoot growth (bloom to veraison) and potassium uptake continuing strongly through ripening for berry loading. Total vine macronutrient content at harvest in a mature vineyard (10 t/ha yield) approximates 100-150 kg N, 15-25 kg P, 120-180 kg K, 80-120 kg Ca, and 15-25 kg Mg per hectare in above-ground biomass. Rootstock influence on macronutrient uptake ranges from minimal (calcium) to substantial (potassium), with 140 Ruggeri conferring 30-50% higher K accumulation than SO4 on the same scion.
A nutrient acquisition disposition that is the inherited capacity of a Vitis vinifera plant to absorb and accumulate the primary mineral nutrients required in large quantities (nitrogen, phosphorus, potassium, calcium, magnesium, and sulfur) through coordinated root transporter activity and root system exploration of the soil volume.
Macronutrient_Uptake_Disposition serves as the parent class for individual macronutrient uptake traits. The category addresses nutrients required at concentrations above 0.1% of dry matter, distinguishing them from micronutrients required at trace levels. Macronutrient balance affects vine physiology, fruit composition, and wine quality, with deficiencies causing visible symptoms and yield reduction while excesses may impair fruit quality or vine health.
Hauptnährstoffakquisition
Makroelement-Aneignung
Makronährstoffaufnahme
macronutrient foraging capacity
major element acquisition
primary nutrient uptake
Conradie WJ (1981) Seasonal uptake of nutrients by Chenin blanc in sand culture: II. Phosphorus, potassium, calcium and magnesium. South African Journal of Enology and Viticulture 2:7-13 Schreiner RP (2016) Nutrient uptake and distribution in young Pinot noir grapevines over two seasons. American Journal of Enology and Viticulture 67:436-448
Macronutrient Uptake Disposition
Sercial bears a Madeira_Style_Scion_Role as the driest noble variety, grown at highest altitudes (up to 800m) achieving extreme acidity (10-12 g/L) and producing austere, almond-inflected Madeiras with residual sugar below 50 g/L that require 20+ years aging to reveal complexity. Malmsey (Malvasia Cândida) serves in a Madeira_Style_Scion_Role for the richest style, with residual sugar of 95-135 g/L balanced by 6-8 g/L acidity, creating treacle-like sweetness with burnt orange, coffee, and caramel complexity in aged examples. Verdelho bears a Madeira_Style_Scion_Role for medium-dry styles (49-78 g/L residual sugar), providing the smoky, nutty character that historically made it Madeira's most prestigious style. Tinta Negra serves in a Madeira_Style_Scion_Role for 85% of production, a versatile variety producing all sweetness levels for blends and entry-level bottlings.
A fortified wine scion role that inheres in a grafted scion cultivar by virtue of its designation for Madeira wine production on the Portuguese island of Madeira, or Madeira-style wines elsewhere, characterized by fortification and deliberate heat treatment (estufagem or canteiro) that creates the style's distinctive caramelized, oxidized character and exceptional longevity. Madeira style scions are classified by four "noble" white varieties that determine sweetness level and style—Sercial (driest), Verdelho (off-dry), Bual/Boal (medium-sweet), and Malmsey/Malvasia (sweetest)—plus the red Tinta Negra that dominates volume production. These varieties must withstand the thermal processing (estufagem at 45-50°C for 3 months, or canteiro natural warming over years) that induces Maillard reactions creating the burnt sugar, caramel, and oxidative character defining Madeira. The combination of high acidity (8-12 g/L), fortification (17-22% alcohol), and heat-induced stability makes Madeira essentially immortal, with bottles from the 18th century remaining drinkable.
Madeira_Style_Scion_Role captures a unique fortified wine tradition where intentional heat damage creates desirable flavor compounds (furfural, hydroxymethylfurfural, sotolon) through Maillard reactions between sugars and amino acids. This process, discovered accidentally during tropical shipping voyages (vinho da roda), is now replicated through estufagem (artificial heating) or canteiro (extended natural warming in sun-exposed warehouses). The noble varieties create a quality and style hierarchy, with single-variety vintage Madeiras from Sercial, Verdelho, Bual, and Malmsey commanding premium prices and exhibiting aging potential measured in centuries. Phylloxera devastated Madeira's vineyards in the 1870s-1880s recovery prioritized productive Tinta Negra over noble varieties, though quality-focused replanting is gradually restoring the classic hierarchy.
Madeira grape
Madeira variety
Madeira wine variety
Madeira-Rebsorte
Madeira-Traube
Madeirasorte
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Liddell A (2014) Madeira: The Mid-Atlantic Wine, Hurst & Company Jackson RS (2014) Wine Science 4th Ed, Academic Press pp.: 520-530 Instituto do Vinho, do Bordado e do Artesanato da Madeira (IVBAM) (2020) Madeira Wine Production Standards
Madeira Style Scion Role
The magnesium content of a Rheingau Riesling vineyard on Taunus quartzite-derived soil documents low exchangeable magnesium of 0.8 cmol(+)/kg (95 mg/kg) at 0-30 cm depth — this low magnesium availability, combined with the high exchangeable potassium (0.6 cmol(+)/kg) from historic potassium fertilization, produces a K:Mg ratio of 0.75 that exceeds the critical threshold of 0.5 above which potassium-induced magnesium deficiency becomes likely. The resulting magnesium deficiency manifests as interveinal chlorosis on basal leaves beginning at véraison — the leaf tissue between the veins turns yellow (in white cultivars) or red (in red cultivars) while the veins and immediately adjacent tissue remain green, progressing from the oldest (basal) leaves toward the shoot tip as magnesium is remobilized from mature leaves to support the actively growing shoot apex and developing berries. The corrective treatment involves foliar magnesium sulfate (MgSO₄) applications at 2-3% concentration from post-bloom through véraison (providing immediate symptom relief), combined with soil-applied dolomitic limestone (CaMg(CO₃)₂ at 2-3 t/ha) to build the exchangeable magnesium reserve and reduce the K:Mg ratio over the medium term.
A nutrient analysis data component that records the concentration of exchangeable magnesium in the vineyard soil, determined by ammonium acetate extraction (1M NH₄OAc at pH 7.0) and expressed in cmol(+)/kg or mg/kg, with magnesium being an essential nutrient required as the central atom of the chlorophyll molecule, as a cofactor for numerous enzymes in carbon fixation and sugar metabolism, and as a structural component of cell membranes — magnesium deficiency producing the characteristic interveinal chlorosis of mature leaves that is one of the most commonly observed nutritional disorders in vineyards worldwide, particularly on acid, sandy soils with low cation exchange capacity.
Magnesium_Content documents the nutrient whose deficiency is the most frequently diagnosed mineral disorder in European vineyards — the high prevalence of magnesium deficiency reflects the interaction of soil factors (low-CEC acid soils that retain little exchangeable magnesium, magnesium-poor parent materials such as quartzite, sandstone, and granite), management factors (excessive potassium fertilization that competitively inhibits magnesium uptake through the shared non-selective cation transport channels in root cell membranes), and rootstock factors (certain rootstocks, notably SO4 and 5BB, are particularly susceptible to magnesium deficiency due to their inherently low magnesium uptake efficiency). The antagonistic relationship between potassium and magnesium is the most practically important cation interaction in vineyard nutrition — both cations enter the root through the same non-selective cation channels (with potassium having higher selectivity due to its lower hydration energy), and high potassium concentrations in the soil solution competitively exclude magnesium from uptake sites: the K:Mg ratio in the soil exchange complex (expressed in cmol(+)/kg to account for the valence difference) is therefore a more reliable predictor of vine magnesium status than the absolute magnesium concentration alone, with ratios above 0.5 indicating increasing risk of potassium-induced magnesium deficiency regardless of the absolute magnesium level. The rootstock influence on magnesium nutrition is substantial — rootstocks vary by a factor of 2-3 in their magnesium uptake efficiency, with high-magnesium-uptake rootstocks (420A, Gravesac, 101-14) providing a genetic solution to magnesium deficiency on susceptible sites, while low-magnesium-uptake rootstocks (SO4, 5BB) exacerbate deficiency on marginal soils.
Boden-Magnesium
Magnesiumgehalt
Mg-Gehalt-Messung
exchangeable Mg measurement
magnesium availability assessment
soil magnesium record
White RE (2003) Soils for Fine Wines, Oxford University Press Delas J (2000) Mineral Nutrition of Grapevines, in Roubelakis-Angelakis KA (ed) Molecular Biology and Biotechnology of the Grapevine, Springer Robinson JB (2005) Critical plant tissue values, ASEV Mpelasoka BS et al. (2003) A review of potassium nutrition in grapevines, Australian Journal of Grape and Wine Research 9: 154-168
Magnesium Content
Magnesium content quality assessment in Grüner Veltliner from the Wachau illustrates the interaction between soil composition and vine nutrition, with primary rock (gneiss, amphibolite) terraces providing moderate magnesium (must concentrations 60-100 mg/L) that supports chlorophyll function without the excessive levels that might inhibit calcium uptake the balanced magnesium nutrition contributes to the intense green-gold color of healthy Grüner Veltliner canopies and the sustained photosynthetic capacity required to ripen fruit on the steep, terraced slopes characteristic of the region's Smaragd sites.
Magnesium_Content_Quality represents the chemical quality characterizing the concentration of magnesium (Mg²⁺) in grape berry tissue, capturing the essential mineral cation central to chlorophyll function, enzyme activation, and carbohydrate metabolism. Magnesium in grape juice typically ranges from 50-150 mg/L, positioned between potassium (dominant) and calcium in abundance among cations. As the central atom of the chlorophyll molecule, magnesium is essential for photosynthesis, with deficiency producing the characteristic interveinal chlorosis where leaf areas between veins yellow while veins remain green due to chlorophyll degradation. Beyond chlorophyll, magnesium activates numerous enzymes involved in phosphate transfer, protein synthesis, and carbohydrate metabolism, making adequate magnesium nutrition essential for vine productivity. In berries, magnesium accumulates through both xylem and phloem pathways, with accumulation continuing throughout ripening though at slower rates than potassium. Soil magnesium availability depends on clay content and exchange capacity, with sandy soils often showing magnesium limitation while certain clay-rich soils provide excess magnesium. The calcium-to-magnesium ratio affects uptake of both elements high-magnesium soils can induce calcium deficiency through competitive inhibition, while high-calcium soils may limit magnesium uptake. In winemaking, magnesium can contribute to wine mineral content but does not cause stability problems comparable to potassium or calcium tartrates. Some researchers have proposed that magnesium contributes to the perceived mineral character in wine, though direct sensory effects remain unestablished. Magnesium analysis employs atomic absorption spectroscopy or ICP-OES with appropriate sample preparation.
Magnesium_Content_Quality addresses an essential mineral whose deficiency produces visible symptoms (interveinal chlorosis) useful for diagnostic purposes. The chlorophyll-central role of magnesium links mineral nutrition directly to photosynthetic function. Magnesium does not create wine stability problems comparable to potassium or calcium, reducing its winemaking significance relative to those elements. The Ca/Mg ratio in soil and tissue provides diagnostic information about potential nutritional imbalances affecting uptake of either element.
Beerenmagnesiumqualität
Magnesiumkonzentration
Mg content
Mg-Gehalt
berry magnesium quality
magnesium concentration
magnesium level
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 341-385 Conradie WJ (1981) Seasonal Uptake of Nutrients by Chenin Blanc in Sand Culture: II. Phosphorus, Potassium, Calcium and Magnesium. South African Journal of Enology and Viticulture 2(1): 7-13
Magnesium Content Quality
Magnesium deficiency symptom quality assessment in Mosel Valley Riesling identified Mg limitation as the cause of widespread interveinal chlorosis and associated grape stem necrosis in vineyards with high-potassium fertilization history on acidic slate soils. Visual magnesium deficiency symptom quality evaluation revealed the characteristic interveinal chlorosis pattern with bright yellow tissue between veins while primary and secondary veins remained distinctly green, affecting 45% of basal leaves by véraison with progressive involvement of mid-canopy leaves through ripening. The magnesium deficiency symptom quality distribution showed concentration in vineyard blocks with documented heavy potassium fertilization exceeding 150 kg K₂O/ha annually, where K-Mg antagonism had induced deficiency despite soil Mg at 180 ppm (nominally adequate). Tissue analysis confirmed magnesium deficiency symptom quality diagnosis with petiole Mg at 0.18% (deficient, threshold 0.30%) and K:Mg ratio of 12.5 (exceeding the 8:1 threshold indicating imbalance). Cluster assessment revealed associated grape stem necrosis affecting 22% of clusters, with rachis tissue showing brown necrotic lesions causing berry drop and cluster desiccation. The magnesium deficiency symptom quality evaluation informed corrective management including foliar MgSO₄ application at 2% concentration for immediate symptom relief, soil-applied dolomitic lime at 2 t/ha to increase soil Mg and raise pH, and reduced potassium fertilization to improve K:Mg balance, with follow-up assessment showing 70% reduction in symptom severity and 85% reduction in grape stem necrosis incidence by the second season post-correction.
Magnesium deficiency symptom quality is a macronutrient deficiency symptom quality that inheres in a grapevine organism and represents the characteristic visual symptoms resulting from inadequate magnesium availability, including the diagnostic interveinal chlorosis where tissue between veins becomes yellow while veins remain green, progressing from older to younger leaves as magnesium is remobilized, along with the associated disorder grape stem necrosis (Stiellähme) affecting cluster rachis tissue, with magnesium deficiency representing a significant limitation particularly on acidic soils, high-potassium soils where K-Mg antagonism reduces magnesium uptake, and sandy soils with low cation exchange capacity. This quality captures the grapevine response to magnesium limitation, recognizing that as the central atom of the chlorophyll molecule and a cofactor for numerous enzymes, magnesium deficiency produces characteristic symptoms reflecting disrupted photosynthetic function. The ontological significance of magnesium deficiency symptom quality lies in its diagnostic distinctiveness with the interveinal chlorosis pattern providing reliable field identification, though tissue analysis is recommended to confirm diagnosis and distinguish from similar-appearing iron or manganese deficiency, and in its association with grape stem necrosis that can cause significant crop loss through cluster drop. The biological basis of magnesium deficiency symptoms encompasses the essential position of magnesium in chlorophyll molecules where each chlorophyll contains one magnesium atom coordinated within the porphyrin ring, the role of magnesium as activator of numerous enzymes particularly those involved in phosphate transfer and carbon fixation, and the function of magnesium in ribosome structure affecting protein synthesis. The competitive relationship between potassium and magnesium for root uptake creates induced magnesium deficiency conditions where high soil potassium or heavy potassium fertilization reduces magnesium acquisition even when soil magnesium is adequate, making the K:Mg ratio in soil and tissue important diagnostic criteria alongside absolute magnesium concentration.
Magnesium_Deficiency_Symptom_Quality is_a Macronutrient_Deficiency_Symptom_Quality, serving as parent class for Interveinal_Chlorosis_Quality_(Mg) and Grape_Stem_Necrosis_Quality which address specific magnesium deficiency manifestations. This class addresses the overall magnesium deficiency syndrome while child classes capture particular symptom types. Mobile nutrient showing symptoms on older leaves first. The K-Mg antagonism creates induced deficiency even with adequate soil Mg. Petiole Mg threshold ~0.30% K:Mg ratio <8:1 desirable. The association with grape stem necrosis (Stiellähme) causes significant crop loss potential.
Magnesiummangel-Symptome
Magnesiumstress-Indikatoren
Mg deficiency symptoms
Mg-Mangel-Zeichen
Niedrig-Magnesium-Symptome
low magnesium symptoms
magnesium limitation signs
magnesium stress indicators
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication Currle O et al. (1983) Biologie der Rebe. Meininger Verlag, Neustadt
Magnesium Deficiency Symptom Quality
SO4 (V. berlandieri × V. riparia) demonstrates high Mg efficiency, maintaining Riesling petiole Mg above 0.30% dry weight on the acidic, K-rich schist soils of the Mosel where 3309 Couderc induces Mg deficiency symptoms (petiole Mg declining below 0.15%). 5BB Kober (V. berlandieri × V. riparia) shows good Mg uptake supporting Zweigelt on the acidic granitic soils of the Weinviertel, Austria, with petiole Mg/K ratios maintained above 0.3 compared to <0.2 on Mg-inefficient rootstocks. Riparia Gloire de Montpellier provides efficient Mg acquisition for Gamay on the acidic granitic sands of Beaujolais, where high soil K and low pH would otherwise induce severe Mg deficiency.
A nutrient acquisition rootstock role that inheres in a grafted rootstock by virtue of its enhanced capacity for magnesium (Mg²⁺) uptake from soil and translocation to the scion, maintaining adequate leaf Mg status and preventing Mg deficiency symptoms under conditions of limited soil Mg availability, high K/Mg ratios, or acidic soil conditions that reduce Mg uptake. Magnesium-efficient rootstocks acquire Mg through high-affinity Mg²⁺ transporter (MRS2/MGT family) expression in root cells, efficient Mg²⁺ loading into xylem, and sustained Mg uptake under competitive inhibition from K⁺, Ca²⁺, NH₄⁺, and H⁺ ions that share common uptake pathways. Magnesium efficiency is quantified by petiole Mg concentration at veraison (efficient genotypes maintaining >0.25-0.30% dry weight under Mg-limiting conditions versus <0.15% for inefficient genotypes), by Mg/K ratios in leaf tissue, and by absence of interveinal chlorosis symptoms characteristic of Mg deficiency (yellowing or reddening between green veins, progressing from older to younger leaves).
Magnesium_Efficient_Rootstock_Role addresses a nutritional constraint that is increasingly common in cool-climate viticulture on acidic soils, particularly where high K fertilization or naturally high soil K depresses Mg availability through competitive inhibition. Unlike Fe deficiency, Mg deficiency symptoms appear first on basal leaves and progress acropetally, reflecting Mg remobilization from older to younger tissue. V. riparia parentage generally contributes good Mg efficiency, while some V. berlandieri × V. rupestris rootstocks (particularly on acidic soils) show poor Mg uptake relative to K. Soil Mg applications (dolomitic lime, Mg sulfate) can compensate for rootstock Mg inefficiency on deficient sites.
Magnesiummangel-tolerante Unterlage
Mg efficient rootstock
Mg-effiziente Unterlage
magnesium deficiency tolerant rootstock
magnesium efficient rootstock
magnesiumeffiziente Unterlage
Delas J, Pouget R (1979) Influence des porte-greffes sur la nutrition minérale de la vigne, Connaissance de la Vigne et du Vin 13: 169-191 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 305-312 Schreiner RP (2005) Spatial and temporal variation of roots, arbuscular mycorrhizal fungi, and plant and soil nutrients, Plant and Soil 276: 219-234 Ruhl EH (1989) Uptake and distribution of potassium by grapevine rootstocks, Australian Journal of Experimental Agriculture 29: 707-711
Magnesium Efficient Rootstock Role
Magnesium uptake ranges from 10-30 kg Mg/ha annually, with rootstock effects of 20-40% variation under identical soil conditions. SO4 and 5BB Kober show high Mg uptake efficiency, while 41B and 140 Ruggeri demonstrate lower Mg acquisition. Magnesium deficiency symptoms (interveinal chlorosis of older leaves) appear when petiole Mg falls below 0.25% dry weight at veraison, with severe deficiency causing premature defoliation. Potassium-magnesium antagonism is pronounced in grapevines, with high K availability suppressing Mg uptake by 30-50% through competitive inhibition at root membrane transporters. Berry Mg concentration of 50-100 mg/L is typical, contributing to wine buffering capacity.
A macronutrient uptake disposition that is the inherited capacity of a Vitis vinifera plant to absorb magnesium ions (Mg²⁺) from the soil solution through magnesium transporter proteins, with genotype-specific uptake efficiency affecting chlorophyll synthesis, enzyme activation, and carbohydrate partitioning.
Magnesium uptake shows intermediate rootstock influence, weaker than potassium but stronger than calcium. The K-Mg antagonism is particularly important in vineyard nutrition, as heavy K fertilization can induce Mg deficiency even on Mg-adequate soils. Rootstocks with high Mg uptake efficiency partially compensate for high K availability. Magnesium's role in chlorophyll structure (central atom of the porphyrin ring) makes adequate uptake essential for photosynthetic capacity and therefore sugar production.
Magnesiumaneignungsvermögen
Magnesiumaufnahme
Mg uptake capacity
Mg-Akquisition
magnesium acquisition
magnesium foraging trait
Conradie WJ (1981) Seasonal uptake of nutrients by Chenin blanc in sand culture: II. Phosphorus, potassium, calcium and magnesium. South African Journal of Enology and Viticulture 2:7-13 Etchebarne F et al. (2009) Leaf senescence in wine grape: effects of potassium deficiency. Functional Plant Biology 36:356-367
Magnesium Uptake Disposition
A main apical shoot extending from the terminal position of a bilateral horizontal cordon in Vitis vinifera cv. Chardonnay in Chablis, France, having grown 95 cm from budbreak in April through early July, with 18 nodes bearing fully expanded leaves averaging 120 cm² leaf area, showing no lateral shoot development in the basal 12 nodes due to strong apical dominance, with the shoot tip still actively elongating at 3-4 cm per day at BBCH stage 73 (berries beginning to touch) and accumulating approximately 45% of total seasonal photosynthate production for this cordon section.
An apical shoot that represents the primary, original terminal extension of a perennial woody structure such as a cane, arm, cordon, or trunk in Vitis vinifera, characterized by continuous development from the previous season's terminal bud or from a designated replacement bud following winter pruning, maintaining the primary growth axis and architectural continuity of the training system. Main apical shoots serve as the principal framework-building elements in grapevine training systems, determining the ultimate dimensions and geometry of permanent structures including cordon length, trunk height, and canopy spread through their seasonal extension growth. These shoots exhibit the strongest apical dominance effects within the vine canopy, creating the steepest auxin gradients that suppress lateral bud outgrowth along their length and along subordinate axes. Main apical shoots typically show the greatest vigor, longest internode extension, largest leaf area, and highest photosynthetic capacity compared to lateral or secondary shoots, functioning as primary carbohydrate sources during vegetative growth phases and as major sinks for mineral nutrients transported from root systems.
Main apical shoots are distinguished from dominant apical shoots by their origin from the intended terminal bud of a pruned structure versus origin from lateral buds that have assumed dominance. In practice, this distinction becomes important in training system establishment where main apical shoot growth determines framework development timeline. Main apical shoots are typically managed through summer topping in production vineyards to redirect growth resources toward fruit development, with topping height determined by training system requirements and cultivar vigor characteristics. After topping, lateral shoots near the cut may develop vigorous growth, but these are not reclassified as main apical shoots.
Hauptendtrieb
Hauptspitzentrieb
primary apical shoot
primärer Apikaltrieb
principal terminal shoot
Based on Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 60-92 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles pp. 88-104 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 142-168.
Main Apical Shoot
A main axis primary inflorescence at node 3 on a shoot of Vitis vinifera cv. Merlot in Bordeaux, France, measuring 17.8 cm total length with prominent central rachis extending the full length providing dominant structural framework, rachis diameter of 4.2 mm at base tapering to 2.0 mm at apex representing robust construction, exhibiting clear monopodial architecture with persistent main axis and lateral branches arising sequentially, bearing 12-15 major lateral branches distributed along the rachis length at intervals of 1.0-1.5 cm, each lateral branch showing secondary and tertiary branching creating the compound paniculate structure, rachis showing green coloration with moderate pubescence, containing well-developed vascular bundles visible as raised ridges providing water and nutrient delivery pathway to all flowers, lateral branches progressively decreasing in size and complexity from base (largest, most complex) to apex (smallest, simplest) creating conical overall profile, architectural organization showing clear hierarchy with rachis dominant over laterals and primary laterals dominant over secondary branches, measured at BBCH stage 61 (beginning of flowering) in early June, representing typical main axis primary inflorescence architecture with clear central rachis dominance characteristic of standard grapevine inflorescence organization.
A primary inflorescence characterized by prominent central rachis (main axis) dominating the architectural organization in Vitis vinifera, characterized by robust central axis providing the primary structural support and organizational framework, rachis diameter typically 3-5 mm at base representing substantial construction, hierarchical lateral branch arrangement with branches arising sequentially along the main axis, clear differentiation between dominant central rachis and subordinate lateral branches, typical length of 15-25 cm for the complete structure with rachis representing the continuous primary axis, architectural pattern showing monopodial organization with persistent main axis rather than sympodial or dichotomous branching, and functional significance with the rachis serving as the primary vascular pathway, mechanical support structure, and developmental organizer coordinating lateral branch formation and flower development. Main axis primary inflorescences represent the typical grapevine inflorescence architecture, exhibiting clear hierarchical organization with dominant central rachis, developing through apical meristem activity establishing the rachis with lateral branch primordia initiating sequentially along its length, and creating the characteristic elongated paniculate structure that transforms into the typical elongated cluster following fruit set.
Main axis primary inflorescences are distinguished from branched or compound types by prominent central rachis dominating the architecture rather than equal co-dominant axes or highly elaborate branching obscuring central organization. The main axis designation emphasizes the monopodial growth pattern with persistent dominant central axis, contrasting with sympodial patterns where main axis terminates and lateral branches assume dominance. Rachis robustness reflects the structural and vascular demands of supporting the complete inflorescence, with diameter correlating with overall inflorescence size and flower number. Hierarchical organization creates clear architectural levels with rachis as first order, major lateral branches as second order, secondary laterals as third order, and so forth typically extending to fourth or fifth orders. Lateral branch distribution along the rachis shows relatively regular spacing creating systematic architecture, though some variation occurs with denser branching basally and sparser apically. The monopodial pattern results from apical meristem persistence establishing continuous rachis elongation with lateral branch primordia initiating from the flanks, contrasting with sympodial or dichotomous patterns requiring apical meristem conversion or splitting. Vascular organization shows continuous central bundles in the rachis with lateral traces departing to supply branches, creating the hierarchical supply network. Transformation to cluster following fruit set maintains the main axis architecture with rachis becoming the cluster stem and lateral branches becoming the berry-bearing laterals. Main axis architecture represents the typical condition in most cultivars, with variations toward more elaborate compound structures or simplified weakly branched structures representing departures from this standard pattern.
Hauptachse-Infloreszenz
Rachis-dominanter Blütenstand
central axis inflorescence
monopodial primary inflorescence
monopodiale primäre Infloreszenz
rachis-dominant inflorescence
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 179-215 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 143-178 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 359-394.
Main Axis Primary Inflorescence
A main petiole on a mature leaf at node 6 of Vitis vinifera cv. Pinot Noir in Burgundy, France, measuring 9.8 cm length from node attachment to lamina base representing typical medium-length main petiole for this cultivar, diameter of 3.2 mm at mid-length tapering slightly toward both ends, exhibiting characteristic light green coloration without red pigmentation typical of this variety, sparse pubescence with scattered short trichomes, cylindrical cross-sectional shape with slight flattening on adaxial surface, containing five major vascular bundles arranged in arc visible as raised ridges on abaxial surface, demonstrating flexibility enabling leaf blade rotation through 180-degree arc in response to wind while maintaining structural integrity supporting 3.8 gram leaf mass at approximately 50-degree angle from horizontal, connecting directly to node without branching and terminating at lamina base where five primary veins radiate into the five-lobed blade, showing characteristic open U-shaped petiolar sinus diagnostic for Pinot Noir, measured at BBCH stage 75 (berries pea-sized) in late June, representing standard unbranched main petiole structure universal in grapevine leaves.
The primary petiolar structure extending from the shoot node to the central point of lamina attachment at the base of the main vein system in Vitis vinifera, characterized by continuous cylindrical to slightly flattened morphology without branching, direct attachment to the shoot at the nodal position, termination at the lamina base where major veins radiate from the petiolar junction point, typical dimensions of 5-14 cm length and 2.5-5 mm diameter in mature leaves varying with cultivar and position, complete vascular system including major xylem and phloem bundles providing primary water and nutrient delivery pathway to lamina and photosynthate export route from leaf, substantial mechanical tissues providing structural support for entire leaf mass, and serving as the sole structural connection between lamina and shoot bearing all mechanical loads including leaf weight, wind forces, and gravitational stresses. Main petioles represent the standard petiolar structure in simple leaves, with grapevine showing exclusively main petiole architecture without the compound leaf branched petiole structures found in some plant families, functioning as the critical bottleneck for all resource exchange between leaf and vine with vascular capacity determining maximum transpiration and photosynthesis rates supportable, and showing cultivar-diagnostic characteristics including relative length, thickness, coloration, pubescence, and flexibility that contribute to ampelographic identification and affect functional leaf positioning and movement capabilities.
Main petioles represent the universal petiolar type in Vitis vinifera, distinguished from theoretical branched or compound structures not found in this species, and from the petiolar sinus which represents the junction zone rather than the stalk itself. All functional grapevine leaves possess main petioles connecting lamina to shoot, with variation occurring in dimensions, coloration, pubescence, and mechanical properties rather than fundamental architecture. Length variation shows cultivar specificity with some varieties consistently producing longer petioles (Sauvignon Blanc, Riesling) while others show shorter petioles (Gewürztraminer, some Muscat types), and environmental plasticity with shade leaves developing longer petioles than sun leaves. Thickness variation affects mechanical strength and vascular capacity, with thicker petioles supporting larger leaves and higher transpiration rates. Petiolar flexibility determined by tissue composition affects leaf movement responses to wind, with more flexible petioles enabling greater blade rotation reducing mechanical stress but potentially reducing optimal light capture maintenance. Coloration particularly red or purple anthocyanin pigmentation shows both genetic control and environmental modification, serving as useful but variable ampelographic character requiring consideration of developmental stage and environmental conditions.
Hauptblattstiel
central petiole
primary petiole
primärer Petiolus
unbranched petiole
unverzweigter Blattstängel
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 166-182 Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 56-78.
Main Petiole
A main primary branch rachis representing the second primary branch (basal position) on a large compound primary inflorescence of Vitis vinifera cv. Muscat of Alexandria in Cyprus, measuring 7.5 cm length representing the longest primary branch on this inflorescence, diameter of 3.2 mm at junction with main rachis representing robust construction, exhibiting substantial development with well-developed mechanical and vascular tissues, containing 6 vascular bundles providing high lateral vascular capacity, bearing 12 secondary branch rachides (third-order structures) distributed along its length creating extensive elaboration, secondary branches further elaborating into tertiary and quaternary structures, complete branch system bearing approximately 95 flowers representing 11.2% of total inflorescence flower number (850 total), substantial mechanical strength supporting the large branch system fresh weight of approximately 4.5 grams, positioned as one of the basal primary branches at 2.8 cm from inflorescence base receiving developmental priority, dimensions approaching some smaller inflorescence main rachis structures highlighting the substantial size, measured at BBCH stage 65 (full bloom) in late May, representing maximum main primary branch development with dimensions and complexity defining major lateral elaborations in very large inflorescences.
A primary branch rachis representing the largest and most developed first-order lateral branches in Vitis vinifera, characterized by dimensions at the upper end of the primary branch range typically 5-8 cm length and 2.5-3.5 mm diameter, position often near the inflorescence base where largest primary branches typically occur, substantial vascular capacity with 5-7 bundles approaching main rachis capacity, extensive secondary branching elaboration with numerous well-developed second-order branches, high flower number contribution often bearing 50-80+ flowers representing major fractions of total inflorescence, enhanced mechanical strength supporting the large branch system, and functional significance as the major lateral structures contributing substantially to overall inflorescence size, complexity, and flower number. Main primary branch rachides represent the maximum development among first-order laterals, occurring typically as the basal primary branches receiving preferential resource allocation and developmental timing advantages, exhibiting dimensions and complexity approaching the subordinate role of secondary main rachis axes, and serving as the major lateral elaborations defining large complex inflorescence architecture.
Main primary branch rachides represent the maximum development extreme among primary branches, distinguished from typical primary branches by superior size, position, and elaboration. The main designation emphasizes the dominance among the first-order lateral population rather than confusion with the main rachis. Size superiority shows main primary branches typically exceeding 6-7 cm length and 2.8-3.5 mm diameter representing the largest laterals, sometimes approaching dimensions of small inflorescence main rachis structures. Positional effects show main primary branches typically occurring at basal positions on the main rachis, reflecting developmental timing (earliest initiation) and resource allocation (preferential investment in basal structures). Vascular capacity approaches main rachis levels with 5-7 bundles common, enabling substantial lateral branch elaboration and high flower support. Secondary branching complexity shows extensive development with numerous well-developed second-order branches creating elaborate branch systems sometimes resembling complete small inflorescences. Flower contribution significance reflects the large size with main primary branches often bearing 60-100+ flowers representing 8-12% of total, making individual main primaries major yield contributors. Occurrence shows position-dependence with the 1-3 most basal primary branches typically showing main primary development while mid and upper primaries show typical or reduced dimensions. Developmental advantages include earliest initiation during inflorescence morphogenesis and preferential resource allocation to basal structures creating the size advantages. Transformation to cluster creates the major cluster arms with main primary branches producing the largest lateral cluster divisions supporting substantial berry masses.
Haupt-Primärzweig-Rachis
dominant first-order branch
dominante erste Ordnung Verzweigung
größte laterale Achse
largest lateral rachis
major primary branch
Based on Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 363-398 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 296-332 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 559-598.
Main Primary Branch Rachis
A main rachis on a primary inflorescence of Vitis vinifera cv. Merlot in Bordeaux, France, measuring 17.8 cm length representing the primary inflorescence dimension from peduncle junction to apex, diameter of 4.2 mm at base where it joins the peduncle tapering progressively to 2.2 mm at the apex, exhibiting robust cylindrical construction with slight flattening on the adaxial surface, green coloration with moderate gray pubescence from scattered trichomes, containing 11 major vascular bundles visible in cross-section arranged in ring configuration providing maximum vascular capacity, bearing 15 primary branch rachides distributed along its length at intervals of 0.8-1.5 cm, serving as the continuous central spine from which all lateral structures arise, providing primary mechanical support bearing the complete inflorescence fresh weight of approximately 18 grams at full bloom, vascular capacity supporting maximum water delivery of estimated 25 ml per day and nutrient transport to all 580 flowers, showing monopodial organization with apical meristem persistence establishing continuous main axis without branching or forking, measured at BBCH stage 63 (30% flowers open) in early June, representing the dominant main rachis serving as primary structural and vascular framework.
The central primary axis of the grapevine inflorescence from which all lateral branches arise in Vitis vinifera, characterized by position as the continuous central spine extending from the peduncle attachment to the inflorescence apex, largest diameter among all rachis structures typically 2.5-5 mm at base tapering to 1.5-3 mm at apex, length representing the primary dimension of inflorescence typically 10-25 cm, monopodial organization with persistent apical growth creating the continuous main axis, first-order hierarchical position from which all lateral branch rachides originate, maximum vascular capacity containing the largest and most numerous vascular bundles often 8-15 major bundles arranged in ring, greatest mechanical strength providing primary structural support bearing the complete inflorescence and subsequent cluster weight, and critical functional roles including serving as the primary vascular highway, main mechanical framework, and developmental organizer coordinating lateral branch and flower positioning. Main rachis represents the dominant structural and functional axis, developing through persistent apical meristem activity establishing continuous elongation, exhibiting maximum robustness reflecting the primary support and vascular transport demands, and serving as the organizing framework upon which the complete branching architecture depends.
Main rachis is distinguished from lateral branch rachides by position as the central continuous axis and largest dimensions, representing the first-order structure in the rachis hierarchy. The main designation emphasizes the primary dominant role as the organizing framework and principal support structure. Monopodial organization with persistent apical meristem creates the continuous unbranched main axis contrasting with sympodial or forked patterns, representing the typical grapevine inflorescence architecture. Diameter shows substantial size advantage with main rachis typically 1.5-2.5× diameter of largest primary branches and 3-5× diameter of secondary branches, reflecting the structural and vascular demands. Length determines primary inflorescence dimension affecting cluster shape and compactness, with cultivar variation from relatively short compact rachis (10-15 cm) to elongated loose rachis (20-25+ cm). Vascular capacity shows maximum development with bundle number often 2-3× that of primary branches, bundle diameter substantially larger, and total conducting area dominating the inflorescence vascular system. Mechanical properties including stiffness and strength must support complete inflorescence weight during flowering and substantially heavier cluster weight post-fruit set, with main rachis failure causing complete cluster loss contrasting with branch failure affecting only local regions. Developmental priority shows main rachis establishment preceding lateral branch initiation, with apical meristem continuous activity throughout inflorescence development. Transformation to cluster stem maintains main rachis dominance with lignification creating woody structure supporting berry mass often 100-500× the flower mass.
Hauptrachis
Hauptspindel
Zentralachse
central rachis
main stem
primary axis
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 143-182 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 179-218 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 313-358.
Main Rachis
Syrah grafted on 1103 Paulsen rootstock at 16 years showing root system with 5 main root axes originating from crown, largest main root axis 13.2 cm diameter at origin extending 165 cm depth at 55° from vertical, main root axis showing extensive secondary xylem with 16 annual rings documenting growth history, bark with protective periderm, main root axis producing lateral roots at 12-20 cm intervals creating hierarchical branching network, main root axis xylem conducting water from lateral absorptive network to trunk, woody tissue storing carbohydrate reserves, main root axes collectively providing structural framework and transport infrastructure for entire root system
Main root axis is primary root emphasizing structural framework role and serving as main architectural element from which lateral root network develops, characterized by substantial diameter typically 3-15 cm at crown providing mechanical strength, extensive secondary growth with woody xylem comprising majority of volume, persistent structure functioning throughout vine life from establishment through old age, and serving as primary conduit for water and nutrient transport from lateral absorptive roots to shoot system, with main root axis potentially showing vertical tap root orientation or oblique orientation depending on rootstock genetics and propagation method, branching to produce hierarchical lateral root network, and representing fundamental structural skeleton of root system analogous to trunk above-ground providing permanent framework supporting transient fine absorptive roots that turnover seasonally while main axis persists. Main root axis structural emphasis includes= providing root system framework (main root axis forming permanent architectural skeleton from which entire root system organized), serving as branching scaffold (main root axis producing lateral roots of progressively smaller orders creating hierarchical branching network), demonstrating persistence (main root axis remaining functional throughout entire vine life unlike fine roots showing annual or multi-annual turnover), showing substantial diameter (main root axis large size reflecting structural rather than primarily absorptive function), and exhibiting woody character (main root axis with extensive secondary xylem creating wood-like structure analogous to above-ground trunk). Main root axis secondary growth includes= producing secondary xylem inward (vascular cambium generating wood tissue increasing main root axis diameter progressively with age), forming secondary phloem outward (cambium producing food-conducting tissue though phloem proportion much less than xylem), showing annual rings (main root axis xylem displaying growth rings analogous to trunk allowing age determination and growth rate assessment), exhibiting bark development (cork cambium forming protective periderm on main root axis surface), and demonstrating progressive thickening (main root axis diameter increasing throughout vine life potentially 0.3-0.8 cm annually in young vigorous vine slowing in old vine). Main root axis diameter and taper includes= showing large diameter at crown (main root axis typically 3-15 cm diameter at origin from trunk base depending on vine age and vigor), exhibiting gradual taper (diameter decreasing progressively along root length as extends from crown), demonstrating branch point diameter drops (main root axis diameter reducing at each lateral root emergence as cross-sectional area divides), displaying root order relationships (main root axis diameter exceeding all branch lateral roots reflecting hierarchical organization), and revealing age through diameter (older main root axes showing greater diameter than younger reflecting years of secondary growth accumulation). Main root axis water transport function includes= conducting water from lateral roots to trunk (main root axis xylem serving as conduit moving water absorbed by fine roots through main axis to shoot system), showing high hydraulic conductivity (large-diameter xylem vessels in main root axis providing efficient low-resistance transport pathway), enabling long-distance transport (main root axis potentially conducting water from lateral roots meters away from trunk to above-ground canopy), demonstrating substantial capacity (main root axis capable of transporting 30-80+ liters daily during peak transpiration depending on root size), and maintaining year-round function (main root axis xylem remaining functional throughout vine life unlike some fine roots subject to seasonal mortality). Main root axis nutrient transport function includes= conducting minerals from absorptive roots (main root axis xylem moving nitrogen, phosphorus, potassium, calcium, magnesium, and micronutrients absorbed by fine roots upward to shoot), showing xylem-phloem coordination (main root axis containing both xylem moving minerals upward and phloem moving photosynthates downward enabling bidirectional transport), demonstrating seasonal variation (nutrient transport through main root axis highest during active growth declining during dormancy), enabling distribution to storage (main root axis phloem conducting photosynthates to root parenchyma for storage), and facilitating remobilization (main root axis xylem moving stored nutrients from root reserves to expanding shoots during spring growth). Main root axis mechanical support includes= providing structural foundation (main root axis large diameter and woody composition creating strong structural framework anchoring vine), bearing mechanical loads (main root axis transmitting forces from above-ground vine weight and wind loads into soil), resisting bending and tension (main root axis woody xylem providing strength against pulling and bending forces), distributing stress throughout system (main root axis branching pattern distributing mechanical loads across multiple roots and soil contact points), and enabling large vine structure (strong main root axes necessary to support extensive canopy and heavy crop loads in high-vigor productive vines). Main root axis carbohydrate storage includes= storing substantial reserves (main root axis parenchyma and ray cells accumulating starch particularly during dormancy), showing large storage capacity (substantial main root axis volume enabling storage of 300-800+ grams dry weight carbohydrates depending on root system size), demonstrating seasonal cycles (reserves highest fall-winter declining spring as mobilized for bud break and initial growth), providing spring growth support (main root axis stored carbohydrates essential for supporting growth before leaves fully functional), and enabling stress recovery (main root axis reserves supporting vine recovery from damage, disease, freeze, or other stress events). Main root axis in root system hierarchy includes= representing highest order (main root axes typically 3rd to 5th order roots in hierarchical classification depending on system and root age), producing lower-order laterals (main root axis branching to create 1st and 2nd order lateral roots and their branches), showing decreasing order with distance (root order typically declining progressively moving away from main root axis toward fine root periphery), demonstrating order-function relationships (main root axis high order correlating with large diameter, woody structure, and transport-storage rather than absorption functions), and organizing spatial architecture (main root axis distribution and orientation determining overall root system spatial organization and soil volume exploitation).
Main_Root_Axis emphasizes structural framework role and is distinguished from Tap_Root (vertical orientation emphasis) by focus on architectural function. Includes Dominant_Main_Root_Axis and Central_Main_Root_Axis. Critical for understanding root system skeleton, hierarchical organization, water-nutrient transport infrastructure, carbohydrate storage, mechanical support, and the structural framework supporting absorptive root network.
Architectural root axis
Hauptwurzelachse
Main root skeleton
Primary structural root
Primäre Strukturwurzel
Wurzelskelett-Achse
Main Root Axis
Maintenance respiration assessment in mature Merlot vineyard quantifies the baseline carbon cost of existing biomass, with maintenance coefficient of 0.015 g glucose/g dry weight/day at 20°C translating to daily maintenance requirement of approximately 180 g glucose for a 12 kg dry weight vine, representing a substantial carbon demand that must be met from current photosynthesis or reserves before any carbon becomes available for growth or fruit development, with this baseline cost persisting through winter dormancy at reduced (temperature-dependent) rates and representing a key determinant of reserve depletion during the non-growing season.
Maintenance respiration quality is a respiration rate quality that represents the rate of respiratory metabolism required to maintain existing cellular structures and functions without net growth, characterizing the baseline metabolic cost of protein turnover, membrane integrity, ionic gradients, and cellular homeostasis that persists regardless of growth activity. Within the BFO framework, this quality specifically addresses the maintenance component of total respiration, distinguishing the costs of cellular upkeep from growth-associated respiration while recognizing that maintenance respiration represents an obligatory carbon cost that continues even when growth ceases and that scales with the amount of metabolically active tissue requiring maintenance. The ontological significance lies in maintenance respiration determining the minimum carbon cost of sustaining existing vine biomass, where the maintenance requirement establishes the baseline respiratory demand that must be met before any carbon becomes available for growth, with the ratio of maintenance to total respiration increasing as growth rate declines and as vines mature, making maintenance costs increasingly significant in established vineyards. The physiological basis for maintenance respiration includes protein turnover (continuous synthesis to replace degraded proteins, representing the largest maintenance cost), membrane maintenance (lipid turnover and repair), ion gradient maintenance (replacing ions lost through membrane leakage, particularly significant in roots), and repair processes (fixing cellular damage from normal metabolism and environmental stress). The factors affecting maintenance respiration rate include tissue nitrogen content (maintenance costs proportional to protein concentration), temperature (exponential increase with warming), tissue age (declining active metabolism reducing maintenance demand), and metabolic rate (tissues with higher baseline activity having higher maintenance requirements). The quantitative expression of maintenance respiration typically employs a maintenance coefficient (g glucose/g biomass/day or μmol CO₂/g/hour) that allows scaling across tissue masses, with values for grapevine tissues typically ranging from 0.01-0.03 g glucose/g dry weight/day at 25°C depending on tissue type and nitrogen content. Understanding maintenance respiration quality enables assessment of the baseline carbon cost of vine biomass, prediction of respiration under non-growing conditions, and appreciation of how vine size and nitrogen status affect minimum carbon requirements.
Maintenance_Respiration_Quality serves as the parent class for organ-specific maintenance respiration (Leaf, Root, Fruit). This quality represents the baseline metabolic cost of cellular homeostasis independent of growth. The class supports carbon budget modeling through characterization of the obligatory respiratory demand that scales with biomass and nitrogen content.
Erhaltungsatmung
Grundumsatz
Rm-Wert
basal respiration
cellular upkeep cost
maintenance metabolism
zelluläre Instandhaltung
Penning de Vries FWT (1975) The cost of maintenance processes in plant cells. Annals of Botany 39(1): 77-92 Amthor JS (1984) The role of maintenance respiration in plant growth. Plant, Cell & Environment 7(8): 561-569
Maintenance Respiration Quality
Chardonnay grafted on Teleki 5C rootstock at 17 years showing major branch secondary roots as second-order branches from first-order laterals, representative major branch 2.6 cm diameter at origin extending 78 cm length at depth 58 cm, major branch showing woody structure with 12 annual xylem rings and protective bark, major branch producing third-order branches 0.4-1.2 cm diameter at 6-11 cm intervals which produce fourth and fifth-order fine roots, major branch xylem conducting water from absorptive network to first-order parent and primary root, major branch providing structural framework supporting extensive fine root network
Major branch secondary root is branch secondary root showing relatively large diameter typically 1-4 cm and substantial structural development, emphasizing size and structural importance within branch root hierarchy, characterized by second-order or occasionally third-order position branching from large first-order laterals or primary roots, moderate to extensive secondary growth with woody xylem development and protective bark formation, persistent structure functioning for multiple years or decades unlike ephemeral fine branches, and serving dual functions as structural framework supporting higher-order absorptive branches and transport conduit moving water and nutrients from fine root network to parent root and ultimately to trunk, with major branch roots typically emerging 8-20 cm intervals along parent root creating relatively sparse but substantial branching skeleton, producing numerous minor branch and fine absorptive roots creating hierarchical network, and representing important structural elements providing permanent framework supporting transient fine root component that shows high turnover. Major branch secondary root size characteristics includes= showing large diameter for branch roots (major branches typically 1-4 cm substantially larger than fine branches but smaller than primary roots), exhibiting substantial length (major branch roots potentially 40-120 cm length exploring significant local soil volume), demonstrating considerable biomass (major branches containing substantial tissue mass contributing significantly to total root system biomass), displaying prominent structure (major branch roots visible in root excavations as substantial structural elements), and revealing size hierarchy (major branches much larger than minor branches and fine roots reflecting hierarchical organization). Major branch secondary root structural development includes= exhibiting extensive secondary growth (major branch roots developing secondary xylem rings and phloem increasing diameter progressively), showing woody structure (major branches with substantial wood cylinder analogous to small primary roots), demonstrating bark formation (cork cambium producing protective periderm on major branch surfaces), displaying multiple annual rings (major branch roots potentially showing 5-15+ annual xylem rings reflecting years of growth), and revealing mechanical strength (major branches with woody structure providing strength supporting branch network and resisting mechanical stress). Major branch secondary root hierarchical position includes= typically second-order (major branch roots most commonly branching from first-order laterals as second hierarchical level), occasionally third-order (some major branches arising from large second-order parents particularly in vigorous root systems), rarely first-order (first-order laterals from primary roots typically considered lateral secondary roots rather than branch roots by definition), serving as parent to higher orders (major branches producing third fourth and fifth-order roots creating branching hierarchy), and demonstrating foundational role (major branches providing structural skeleton from which finer absorptive network develops). Major branch secondary root persistence includes= functioning multiple years (major branch roots typically persistent structures lasting 3-10+ years compared to annual fine roots), showing continued growth (major branches adding length and diameter through successive growing seasons), demonstrating long-term investment (major branch construction representing significant carbon investment amortized over years), exhibiting stable framework (major branches providing relatively stable structural framework despite dynamic fine root turnover), and requiring eventual replacement (major branches eventually declining and dying necessitating production of replacement major branches maintaining structural network). Major branch secondary root tertiary branching includes= producing numerous higher-order roots (major branches generating 10-30+ third fourth and fifth-order roots creating dense absorptive network), showing variable spacing (higher-order roots emerging 3-10 cm intervals along major branch depending on order and conditions), exhibiting branching gradient (branch density typically higher near major branch tip declining toward base), demonstrating responsive proliferation (major branch producing more absorptive roots in favorable soil zones and fewer in poor zones), and creating local exploration (major branch with higher-order branches thoroughly exploring soil volume in local zone around branch). Major branch secondary root transport function includes= conducting water from absorptive network (major branch xylem moving water from third fourth and fifth-order roots toward parent and primary roots), transporting nutrients basipetally (major branches conveying minerals absorbed by fine root network toward plant body), showing substantial hydraulic conductivity (woody major branches with large xylem vessels providing efficient low-resistance transport pathway), enabling phloem distribution (major branch phloem conducting photosynthates to fine roots and storing excess in parenchyma), and demonstrating significant capacity (major branches potentially transporting 5-15 liters daily from local absorptive network depending on branch size). Major branch secondary root storage function includes= accumulating carbohydrate reserves (major branch woody tissue with abundant parenchyma and ray cells storing starch during dormancy), showing seasonal storage cycles (reserves highest fall-winter declining spring as mobilized for growth), providing moderate storage capacity (major branch size enabling storage of 50-200+ grams dry weight carbohydrates per branch depending on size), supporting spring growth (major branch reserves mobilized for renewed root and shoot growth), and enabling stress recovery (major branch carbohydrates supporting vine recovery from damage or stress). Major branch secondary root anchorage contribution includes= contributing to lateral stabilization (major branch network helping resist horizontal forces and toppling), providing distributed anchoring (major branches creating multiple anchor points throughout soil volume), demonstrating tensile strength (woody major branches resisting pulling forces from wind or weight), showing complementary function (major branches working with primary roots providing comprehensive anchorage), and enabling large vine support (major branch framework necessary for supporting extensive canopy and crop loads).
Major_Branch_Secondary_Root emphasizes large size and structural importance and is distinguished from Minor_Branch_Secondary_Root (smaller, more ephemeral) by focus on substantial structure. Critical for understanding branch root hierarchy, structural framework, persistent architecture, transport function, carbohydrate storage, and the large-diameter branch roots providing woody skeleton supporting ephemeral absorptive root network.
Große Verzweigungswurzel
Large branch root
Structural branch root
Strukturelle Seitenwurzel
Substantial secondary branch
Substanzielle Nebenwurzel
Major Branch Secondary Root
In Merlot berries in Saint-Émilion, Bordeaux during BBCH 85-89, malate degradation reduces malic acid content from 11.2 g/L at veraison to 2.8 g/L at harvest over 40 days, representing 75% degradation with NADP-malic enzyme activity peaking at 2.8 µmol/min/g fresh weight at 20 days post-veraison under mean ripening temperatures of 24°C, while parallel tartaric acid levels remain stable at 5.2-5.5 g/L.
Malate degradation encompasses the enzymatic catabolism of L-malic acid within Vitis vinifera berry mesocarp cells during ripening, wherein the 8-15 g/L malic acid present at veraison is reduced by 60-90% to final concentrations of 1-4 g/L through the coordinated action of cytosolic NADP-malic enzyme, mitochondrial NAD-malic enzyme, and respiratory oxidation via the tricarboxylic acid cycle over the 30-45 day post-veraison ripening period. This process represents the primary mechanism of acid reduction in grape berries, with malic acid serving as the predominant organic acid subject to metabolic turnover due to the presence of efficient degradation pathways absent for tartaric acid, fundamentally altering the organic acid profile from malate-dominated to tartrate-dominated composition during ripening progression. The cellular biochemistry initiates with malate efflux from vacuolar storage through ALMT-family aluminum-activated malate transporter channels and voltage-dependent anion channels, delivering substrate to cytosolic and mitochondrial enzyme systems, with vacuolar malate pools of 200-400 mM progressively depleted over the ripening period. Cytosolic NADP-malic enzyme catalyzes oxidative decarboxylation of malate to pyruvate with CO2 release and NADPH generation, representing the quantitatively dominant degradation pathway accounting for 50-70% of total malate catabolism, with enzyme activity increasing 3-5 fold between veraison and mid-ripening in response to developmental and hormonal signals. Mitochondrial metabolism contributes through NAD-malic enzyme and direct oxidation via malate dehydrogenase and the TCA cycle, generating ATP for cellular energy requirements while releasing malate-derived carbon as respiratory CO2. The gluconeogenic pathway diverts malate carbon toward hexose synthesis through phosphoenolpyruvate carboxykinase, contributing 10-20% of berry sugar accumulation from organic acid precursors and linking acid degradation to sugar metabolism. Temperature profoundly regulates malate degradation rates, with warm conditions of 30-35°C accelerating enzymatic activities and respiratory flux 2-3 fold compared to cool conditions of 15-20°C, establishing climate as the primary environmental determinant of final berry acidity. Cultivar variation in malate degradation reflects genetic differences in malic enzyme expression levels and activity, with high-acid varieties maintaining lower enzyme activities and correspondingly slower degradation rates throughout ripening.
Malate_Degradation is subclass of Active_Acid_Reduction representing the specific biochemical process of malic acid catabolism. Distinguishes from pH_Increase which is chemical consequence and from Continuing_Acid_Decline which describes temporal pattern. Parent to Malate_Respiratory_Consumption, Malic_Enzyme_Activity, and Progressive_Malate_Depletion under separate Malic_Acid_Degradation branch. Duration 30-45 days. Key parameter is 60-90% malic acid reduction.
Malat-Abbau
enzymatic malate consumption
enzymatischer Malat-Abbau
malate breakdown
malic acid catabolism
malic acid degradation
Äpfelsäure-Katabolismus
Sweetman C et al (2009) Regulation of malate metabolism in grape berry and other developing fruits. Molecular Plant 2: 1270-1280 Ruffner HP (1982) Metabolism of tartaric and malic acids in Vitis. Vitis 21: 346-358 Keller M (2020) The Science of Grapevines 3rd Ed pp 268-272
Malate Degradation
In a Marlborough Sauvignon Blanc vineyard in New Zealand, malate degradation onset was characterized in early February Year N+1 (BBCH 83, 3 days post-malic enzyme activation) when malate declined from 11.2 g/L to 9.4 g/L over 8 days (16% reduction, 0.22 g/L per day), NADP-malic enzyme activity 24 units/g FW, berry respiration 1.4 μmol O₂ g⁻¹ h⁻¹, estimated malate contribution 42% of CO₂ production, pH increased from 3.04 to 3.18, under moderate New Zealand summer conditions (daily high 23-26°C, cool nights 13-16°C).
Malate_Degradation_Onset represents the beginning of net malate catabolism when respiratory consumption through malic enzyme and citric acid cycle pathways exceeds any residual synthesis establishing the declining trajectory that continues throughout ripening fundamentally transforming berry organic acid composition. This onset occurs during veraison initiation (BBCH 83), typically 1-4 days following malic enzyme activation, and is characterized by vacuolar malate concentrations beginning sustained decrease from 85-145 mM (8.5-14.5 g/L) to 70-120 mM over initial 6-10 days representing 12-22% reduction establishing clear declining trend, cytoplasmic malate turnover increasing as catabolism accelerates with estimated cytoplasmic residence time declining from 3-8 hours (green berry) to 1-4 hours (onset) reflecting enhanced metabolic flux, and berry respiratory CO₂ production showing 30-50% contribution from malate catabolism (based on isotope tracer studies) compared to 32°C causing excessive degradation potentially depleting malate prematurely, and cool conditions 18-22°C slowing degradation preserving acids beneficial for cultivars prone to excessive loss. Cultivar genetic differences show: low-acid varieties exhibiting rapid early degradation with high enzyme activities, short lag from enzyme activation to net degradation 1-3 days, and extensive total depletion 80-95%, moderate cultivars showing intermediate characteristics 3-5 day lag and 70-85% depletion, while high-acid types display delayed degradation 5-8 day lag and moderate depletion 55-75% reflecting genetic variation in enzyme expression, substrate pools, and metabolic capacity. Physiological consequences inclu carbon loss representing respiratory cost with 10 g/L malate loss equivalent to ~7 g/L glucose-equivalent carbon (molecular weight basis), pH increase correlating with malate loss (each 1 g/L malate removal increasing pH approximately 0.03-0.06 units depending on buffer capacity), and osmotic effects minor as malate contributes only 5-10% of total berry osmotic potential compared to hexoses dominating 70-85%. Wine quality implications inclu degradation onset timing and intensity predicting final malate levels critical for wine balance, early rapid onset potentially indicating low final acids requiring monitoring and possible early harvest, delayed or slow onset suggesting acid retention possibly beneficial in warm climates, and spatial variation across vineyard revealing maturity heterogeneity requiring management. Monitoring degradation onset involves: HPLC organic acid analysis tracking malate concentrations showing declining trend, enzymatic malate assays providing rapid field assessment, titratable acidity measurement showing overall acid decline with malate contributing majority, and respiration measurements potentially revealing enhanced CO₂ production from acid catabolism though technically challenging in vineyard setting.
Malate_Degradation_Onset is distinguished from Malic_Acid_Decline_Initiation by representing net catabolism rather than enzyme activation, from Malic_Acid_Decline_Start by showing onset (12-22% reduction) rather than sustained process, and from later degradation by exhibiting beginning rather than progressive loss. Enzyme activities 12-30 units/g NADP-ME at onset. Timing: 1-4 days post-enzyme activation BBCH 83. Temperature Q₁₀ 2.0-3.5. Malate_Degradation_Onset establishes declining trajectory continuing throughout ripening affecting wine acidity and balance.
L-Malat-Konsum-Initiierung
L-malate consumption initiation
Netto-Malat-Katabolismus-Onset
malic acid oxidation start
net malate catabolism onset
respiratorische Malat-Verwendungs-Beginn
respiratory malate utilization beginning
Äpfelsäure-Oxidations-Start
Sweetman C, Deluc LG, Cramer GR, Ford CM, Soole KL (2009) Regulation of malate metabolism in grape berry and other developing fruits Phytochem 70:1329-1344 malate catabolism comprehensive Ruffner HP, Hawker JS, Hale CR (1976) Temperature and enzymic control of malate metabolism in berries of Vitis vinifera Phytochem 15:1877-1880 degradation mechanism Famiani F, Battistelli A, Moscatello S, Cruz-Castillo JG, Walker RP (2015) The organic acids that are accumulated in the flesh of fruits:occurrence, metabolism and factors affecting their contents Plant Physiol Biochem 94:195-206 acid metabolism in fruits.
Malate Degradation Onset
In Shiraz berries in Barossa Valley, Australia during BBCH 87-89, malate respiratory consumption accounts for 28% of total malic acid degradation, oxidizing approximately 2.4 g/L malate through mitochondrial respiration over the 35-day ripening period, with respiratory rates peaking at 45 µL CO2/g fresh weight/hour during warm afternoon periods of 32°C and declining to 18 µL CO2/g fresh weight/hour during cool nights of 18°C.
Malate respiratory consumption describes the oxidative catabolism of L-malic acid through mitochondrial respiration in Vitis vinifera berry cells during ripening, wherein malate serves as a direct respiratory substrate entering the tricarboxylic acid cycle for complete oxidation to CO2 and H2O with coupled ATP synthesis, accounting for approximately 20-30% of total malic acid degradation and contributing significantly to cellular energy production during the metabolically demanding ripening phase. This process involves malate transport across the inner mitochondrial membrane via the dicarboxylate carrier and subsequent oxidation through two primary pathways including direct conversion to oxaloacetate by mitochondrial malate dehydrogenase feeding into the TCA cycle, and oxidative decarboxylation by NAD-malic enzyme producing pyruvate which enters the cycle through pyruvate dehydrogenase. The quantitative contribution of respiratory consumption to total malate degradation varies with developmental stage and environmental conditions, representing 15-20% of malate flux during early ripening when cytosolic malic enzyme activity dominates, increasing to 30-40% during late ripening as cytosolic enzyme activities decline and mitochondrial respiration becomes proportionally more important. The energetic yield from complete malate oxidation generates approximately 15 ATP equivalents per malate molecule through oxidative phosphorylation, providing substantial energy for active transport processes including sugar loading, secondary metabolite biosynthesis, and maintenance of cellular integrity during the ripening period. Temperature strongly influences respiratory consumption rates, with Q10 values of 2.0-2.3 indicating near-doubling of mitochondrial malate oxidation for each 10°C increase, contributing to accelerated total acid decline under warm conditions through enhanced respiratory flux. The diurnal pattern of respiratory consumption shows peak rates during warm afternoon periods when mitochondrial activity is stimulated, with reduced nighttime rates allowing partial recovery of cytosolic malate pools through continued vacuolar efflux. Berry respiratory quotients during ripening reflect the mixed substrate utilization including malate oxidation, with RQ values of 0.9-1.1 indicating balanced carbohydrate and organic acid respiration. The coupling between malate respiratory consumption and other ripening processes includes provision of TCA cycle intermediates for amino acid biosynthesis and generation of reducing equivalents for secondary metabolite modification reactions.
Malate_Respiratory_Consumption is subclass of Malic_Acid_Degradation representing mitochondrial oxidative pathway. Distinguishes from Malic_Enzyme_Activity which is primarily cytosolic decarboxylation. Sibling to Malic_Enzyme_Activity and Progressive_Malate_Depletion. Accounts for 20-30% of total malate catabolism. Important for energy provision during ripening. Temperature-sensitive with Q10 of 2.0-2.3.
Malat-Atmungsverbrauch
malate TCA cycle oxidation
mitochondrial malate oxidation
mitochondriale Malat-Oxidation
oxidative malate consumption
oxidativer Malat-Verbrauch
respiratorischer Malat-Katabolismus
respiratory malate catabolism
Sweetman C et al (2009) Regulation of malate metabolism in grape berry. Molecular Plant 2: 1270-1280 Ruffner HP and Hawker JS (1977) Control of glycolysis in ripening berries of Vitis vinifera. Phytochemistry 16: 1171-1175 Keller M (2020) The Science of Grapevines 3rd Ed pp 270-272
Malate Respiratory Consumption
Male flower disposition characterizes approximately 50% of wild V. vinifera ssp. sylvestris individuals and some Vitis species used as rootstocks. Male flowers produce 2-3 times more pollen per flower than hermaphrodite flowers (15,000-25,000 vs 8,000-12,000 pollen grains), but the pistil remains rudimentary with an undeveloped ovary. V. rupestris and some V. riparia selections used in rootstock breeding possess male flowers. In breeding programs, male parents contribute genetic diversity without fruit production, requiring controlled pollination to generate progeny.
A flower type disposition that is the inherited capacity of a Vitis vinifera plant to develop staminate (functionally male) flowers containing fully developed stamens producing abundant viable pollen but possessing only a vestigial, non-functional pistil incapable of fruit development, serving exclusively as a pollen source.
Male flower disposition results from the presence of the male allele (M) with female-sterility function at the Sex locus. Male-flowered vines are excluded from commercial fruit production but remain important in rootstock breeding and as genetic resources. The vestigial pistil in male flowers arrests development at the ovary primordium stage, lacking ovule differentiation. Some interspecific hybrids show unstable sex expression with partial pistil development.
Männlichkeit
functionally male flower
male vine disposition
männliche Blütenanlage
staminate Blühdisposition
staminate flower trait
Fechter I et al. (2012) Candidate genes within a 143 kb region of the flower sex locus in Vitis. Molecular Genetics and Genomics 287:247-259 Gallardo A et al. (2009) Pollen development and fertility in Vitis vinifera L. cv. Macabeo: a comparison between male-sterile and fertile flowers. American Journal of Enology and Viticulture 60:353-361
Male Flower Disposition
Wild Vitis vinifera ssp. sylvestris male plants in Mediterranean riparian populations occasionally exhibit male only flowers completely lacking any pistil structure, with five robust stamens (filaments 3.5-4.2 mm, anthers 0.8-1 mm) arranged around an empty flower center, producing 38,000-48,000 pollen grains per flower with 82-90% viability, achieving 0.0% fruit set, and functioning as highly effective pollen sources for nearby pistillate plants, with individual male vines capable of providing sufficient pollen for fertilization of 20-30 female vines within 50 m radius under favorable wind conditions. Breeding population seedlings from crosses with wild dioecious V. sylvestris occasionally produce male only flowers with complete pistil absence, identified during first flowering and removed from breeding nurseries to avoid space allocation to non-fruit-bearing genotypes. Wild Vitis riparia (North American species) male plants characteristically bear male only flowers with no pistil trace, creating obligate dioecious reproductive system.
A male only flower is a staminate flower in which the pistil is completely absent or reduced to such an extreme degree that no recognizable pistil structure can be identified even under magnification, representing the most complete expression of morphological male unisexuality with exclusive investment in male reproductive function through well-developed stamens producing abundant viable pollen. Male only flowers are characterized by= (1) complete absence of any pistil structure in the flower center, with the receptacle bearing only the five stamens surrounding an empty central position where pistil would occur in hermaphrodite or pistillate flowers, or (2) an extremely minute, completely non-differentiated tissue remnant <0.2 mm that lacks any recognizable pistil features (no discernible ovary, style, or stigma components) and appears as an undifferentiated cellular mass or small protuberance. The male only condition represents maximum pistil suppression, with pistil development either completely prevented during flower organogenesis (resulting in true absence) or arrested at extremely early developmental stages before any pistil organ differentiation (resulting in undifferentiated remnant). Male only flowers exhibit enhanced stamen development with stamens showing maximum elaboration among Vitis flower types= filaments typically 3-4.5 mm long (30-60% longer than hermaphrodite flower stamens), anthers 0.7-1.1 mm long (20-40% larger than hermaphrodite anthers), and pollen production often 35,000-50,000 grains per flower (up to 2-3× hermaphrodite flower pollen production), representing substantial resource allocation to male function enabled by complete elimination of female function. The five stamens are arranged in a characteristic pattern around the empty flower center, creating a morphology distinctly different from the stamen-surrounding-pistil arrangement of hermaphrodite flowers. Pollen from male only flowers is highly viable (75-92% viability typical) and highly functional, germinating readily on receptive stigmas and producing vigorous pollen tubes, making male only flowers extremely effective pollen donors despite complete inability to produce fruit. Male only flowers achieve absolute zero fruit set (0.0%) due to complete absence of functional female structures, with plants bearing exclusively male only flowers producing no berries under any conditions and functioning purely as pollen sources in the reproductive system. Male only flowers are characteristic of some male individuals in wild dioecious Vitis vinifera ssp. sylvestris populations, particularly in populations with strong sex determination, where approximately 30-50% of male individuals may bear male only flowers while the remaining male individuals bear staminate flowers with vestigial or rudimentary pistil remnants. The male only condition also appears occasionally in breeding populations derived from wild dioecious germplasm, where seedlings homozygous or hemizygous for strong male-determining alleles express complete pistil suppression. The developmental genetics underlying male only flowers involves sex-determining genes that completely suppress pistil primordium specification or arrest pistil development at extremely early stages, possibly through direct transcriptional repression of pistil identity genes or through developmental pathway disruption preventing pistil organ formation. The evolutionary significance relates to sexual specialization in dioecious populations, where complete resource allocation to male function (maximum pollen production) may be favored by sexual selection in male individuals, creating divergent evolution of male only types producing maximum pollen versus reduced pistil types maintaining vestigial female structures.
Male_Only_Flower represents maximum pistil suppression among staminate flowers and is distinguished from Reduced_Pistil_Flower (which retains vestigial or rudimentary pistil structures) by complete or near-complete absence of recognizable pistil morphology. The boundary between 'male only' (no pistil) and 'vestigial pistil' (minimal pistil remnant) can be subtle and may require careful dissection and magnification for definitive classification, as extremely reduced vestigial pistils <0.2 mm can be easily overlooked in casual examination. Male only flowers represent the endpoint of pistil suppression and likely involve the strongest expression of male-determining genetic factors. The complete absence of female structures enables maximum resource allocation to pollen production, potentially providing fitness advantages in dioecious populations through enhanced male reproductive success. Sub-classification into Fully_Developed and Mature types reflects developmental state distinctions within the male only category.
pistil-absent flower
purely staminate flower
rein männliche Blüte
stempellose Blüte
vollständig männliche Blüte
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 101-104 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Caporali, E. et al. (2003). The arrest of development of abortive reproductive organs in the unisexual flower of Vitis vinifera. Sexual Plant Reproduction 15=291-300 Picq, S. et al. (2014). A small XY chromosomal region explains sex determination. BMC Plant Biology 14=229.
Male Only Flower
Malic acid accumulation peaks at or shortly after veraison with concentrations of 15-25 g/L fresh weight in developing berries, subsequently declining to 1-8 g/L at harvest depending on cultivar and ripening temperature. High-malate cultivars such as Barbera and Colombard maintain 5-8 g/L malate at harvest even in warm climates, while low-malate types such as Chasselas and Palomino decline to 1-3 g/L. PEPC activity in green berries fixes respiratory CO₂ into oxaloacetate, which is reduced to malate by malate dehydrogenase (MDH) for vacuolar storage. Pre-veraison malate accumulation provides respiratory substrate for post-veraison berry metabolism and contributes to osmotic sugar accumulation through malate-sugar exchange across the tonoplast.
An acid metabolism disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate malic acid in grape berries through phosphoenolpyruvate carboxylase (PEPC) activity and the citric acid cycle, with genotype-specific peak concentrations reached at or shortly after veraison before respiratory degradation during ripening.
Malic acid accumulation establishes the reservoir that will be subsequently depleted during ripening, with the balance between accumulation capacity and degradation rate determining final malate concentration. The trait shows moderate heritability (h² = 0.45-0.60) and significant genotype × environment interaction. High malate accumulation provides potential for maintaining acidity in warm climates if combined with low degradation rate, a breeding target for climate change adaptation.
L-malate accumulation
Malatbildung
malate synthesis capacity
malic acid trait
Äpfelsäure-Disposition
Äpfelsäureakkumulation
Sweetman C et al. (2014) Regulation of malate metabolism in grape berry and other developing fruits. Phytochemistry 99:1-16 Ruffner HP (1982) Metabolism of tartaric and malic acids in Vitis: a review, Part B. Vitis 21:346-358
Malic Acid Accumulation Disposition
Malic acid content for Chardonnay from Chablis (2024 vintage) measures 6.8 g/L at harvest (12.5° potential alcohol), the high malate retention reflecting the cool continental climate where mean ripening temperatures of 16-19°C slow mitochondrial respiration of malic acid, with the winemaker planning full malolactic fermentation to reduce perceived acidity from the combined tartaric + malic total of 13.5 g/L to a more palatable post-MLF level. Malic acid content for Shiraz from the Barossa Valley (2023 vintage) records only 1.8 g/L at harvest (14.5° Baumé), demonstrating the near-complete thermal degradation of malate during the hot Australian ripening season (mean February temperature 22-24°C), with the low residual malate rendering malolactic fermentation largely inconsequential for acid balance and primarily relevant for microbial stability and flavor modification. Malic acid content for Blaufränkisch from Mittelburgenland (2024 vintage) documents 4.2 g/L at 19° KMW, providing adequate malic substrate for the planned malolactic fermentation that will soften the acid structure for the premium Mittelburgenland DAC wine style.
Malic acid content is an acidity data entity that records the concentration of L-(-)-malic acid (2-hydroxybutanedioic acid) in grape must, determined by enzymatic assay (L-malate dehydrogenase), HPLC, or FTIR estimation, expressed in grams per liter (g/L), representing the second major organic acid in grape berries whose concentration is highly dynamic during ripening due to temperature-dependent mitochondrial respiration, and whose post-fermentation fate (retention in wine or conversion to lactic acid through malolactic fermentation) constitutes one of the most consequential winemaking decisions for wine style and stability.
Malic acid content is the most climate-sensitive component of must acidity, functioning as a biochemical thermometer that integrates the temperature experience of the berry during the ripening period. The temperature dependence is direct and well-characterized — malic acid is consumed through mitochondrial respiration via the TCA cycle at rates that approximately double for each 10°C increase above 20°C, meaning that identical cultivars harvested at equal sugar levels will contain dramatically different malic acid concentrations depending on ripening temperature. This thermal sensitivity makes malic acid content one of the clearest indicators of climate change impact on grape composition, with documented declines of 0.5-1.5 g/L per decade in many European viticultural regions over the past 30 years. The malolactic fermentation (MLF) decision — whether to allow or prevent the bacterial conversion of malic acid to lactic acid (reducing total acidity by approximately 1-3 g/L and raising pH by 0.1-0.3 units) — represents a pivotal winemaking choice that is directly informed by must malic acid content. High-malate musts (>5 g/L) from cool climates typically benefit from MLF to achieve sensory balance, while low-malate musts (<2 g/L) from warm climates gain little acid reduction from MLF and may undergo it primarily for microbial stability (removing a substrate for spoilage Lactobacillus) rather than acid adjustment. The enzymatic assay using L-malate dehydrogenase (EC 1.1.1.37) coupled with diaphorase provides the most specific quantification, though HPLC offers simultaneous determination of tartaric, malic, citric, succinic, and lactic acids from a single injection.
Apfelsäuregehalt
Apfelsäurekonzentration
L-malic acid content
L-Äpfelsäure
Malatgehalt
malate content
malate level
malic acid analysis
malic acid concentration
Äpfelsäuregehalt
Sweetman C et al. (2014) Regulation of malate metabolism in grape berry and other developing fruits. Phytochemistry 98: 1-13 Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 400-415 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1, 2nd Ed, Wiley, pp.: 219-240
Malic Acid Content
Malic acid content quality in New Zealand Sauvignon Blanc from Marlborough reaches exceptionally high levels (5-8 g/L at harvest) due to cool ripening temperatures and the variety's genetic predisposition toward malic acid retention, creating the intense, mouth-watering acidity that defines the regional style winemakers preserve this malic character by preventing malolactic fermentation through cold settling, early sulfur dioxide addition, and sterile filtration, maintaining the vibrant green apple and citrus acidity that distinguishes Marlborough Sauvignon Blanc from the softer, malolactic-fermented styles of Bordeaux Blanc and many New World interpretations.
Malic_Acid_Content_Quality represents the chemical quality characterizing the concentration of malic acid (2-hydroxybutanedioic acid, C₄H₆O₅) in grape berry tissue. Malic acid is the second major organic acid in grapes, synthesized through the tricarboxylic acid cycle and related pathways, and distinguished from tartaric acid by its metabolic lability that results in dramatic concentration changes during berry development and wine production. Unlike tartaric acid, malic acid is readily respired by berry mitochondria, with catabolism rates strongly influenced by temperature. The concentration of malic acid at harvest varies enormously, from less than 1 g/L in warm climates to more than 8 g/L in cool climates, making it the primary variable determining regional acidity differences. This variation reflects the temperature-dependent balance between malic acid synthesis in green berries and respiration that accelerates dramatically above 30°C. Cool nights during ripening preserve malic acid, while warm nights drive rapid malic acid loss through respiration, explaining the marked acidity differences between continental and maritime climates. Malic acid contributes green, tart, apple-like sourness (malum is Latin for apple) that can be perceived as harsh when excessive. Malolactic fermentation, the bacterial conversion of malic acid to lactic acid, represents a major transformation of acid composition that softens acidity and adds complexity, with winemakers choosing to encourage or prevent this conversion based on wine style goals. Cultivar differences in malic acid retention are significant, with some varieties (Riesling, Colombard) maintaining high malic through ripening while others (Gewürztraminer, Viognier) naturally achieve low malic levels. Measurement employs specific enzymatic assays using malate dehydrogenase or HPLC separation.
Malic_Acid_Content_Quality addresses the metabolically dynamic acid whose concentration varies dramatically with climate and winemaking decisions. The L-(−)-malic acid stereoisomer is the natural form in grapes. The strong temperature dependence of malic acid respiration makes it a sensitive indicator of ripening conditions and explains much of the climate effect on wine acidity. The malolactic fermentation pathway (malate → lactate + CO₂, catalyzed by Oenococcus oeni) represents a major transformation that fundamentally changes acid composition, reducing titratable acidity by approximately 1-3 g/L while softening perceived sourness more than the TA change would suggest.
L-malic acid level
Malat
apple acid concentration
hydroxysuccinic acid quality
Sweetman C et al. (2009) Regulation of Malate Metabolism in Grape Berry and Other Developing Fruits. Phytochemistry 70(11-12): 1329-1344 Volschenk H et al. (2006) Malic Acid in Wine: Origin, Function and Metabolism during Vinification. South African Journal of Enology and Viticulture 27(2): 123-136
Malic Acid Content Quality
In a Barossa Valley Shiraz vineyard in Australia, malic acid decline initiation was characterized in mid-February Year N+1 (BBCH 83) when malic acid declined from 12.8 g/L to 10.5 g/L over 6 days (18% reduction), VvME1 expression increased 32-fold, NADP-malic enzyme activity rose from 4.2 to 21.5 units/g FW, berry Brix 11.2° beginning accumulation, pH increased from 2.98 to 3.14, under hot Australian conditions (daily high 31-35°C accelerating malate catabolism, requiring canopy shading intervention to moderate decline).
Malic_Acid_Decline_Initiation represents the beginning of active malate respiratory catabolism when malic enzyme and citric acid cycle activities increase enabling conversion of stored malic acid to CO₂ and energy creating the primary mechanism for organic acid reduction during berry ripening. This initiation occurs during veraison onset (BBCH 83), typically 0-3 days concurrent with or slightly preceding overall acid decline onset, and is characterized by malic acid concentrations beginning decrease from pre-veraison maximum 9-15 g/L (90-150 mM vacuolar) to initiation 8-13 g/L over 4-7 days representing initial 8-18% reduction, NADP-malic enzyme activity increasing from green berry 2-6 units/g fresh weight to 10-25 units/g establishing catalytic capacity for malate oxidation, VvME gene expression rising 12-40 fold coordinating with activity increases, and berry respiration rates maintained or slightly elevated 0.9-2.0 μmol O₂ g⁻¹ h⁻¹ as malate catabolism begins contributing substantially to respiratory substrate. Enzymatic basis of malate decline includes: NADP-dependent malic enzyme (EC 1.1.1.40) catalyzing oxidative decarboxylation L-malate + NADP⁺ → pyruvate + CO₂ + NADPH with pH optimum 7.0-7.5 matching cytosolic environment, enzyme kinetics showing Km for malate 0.5-2.0 mM and Vmax at initiation 12-28 μmol malate oxidized g⁻¹ h⁻¹ adequate for observed decline rates, NAD-dependent malic enzyme (EC 1.1.1.38) showing similar reaction with NAD⁺ cofactor and lower activity 4-15 units/g contributing secondarily, malate dehydrogenase in citric acid cycle (EC 1.1.1.37) catalyzing reversible malate ⇌ oxaloacetate interconversion enabling malate entry into cycle for complete oxidation, and phosphoenolpyruvate carboxykinase potentially involved though role less defined in grape compared to CAM or C₄ plants. Gene expression during malate decline initiation shows: VvME1 (primary NADP-malic enzyme isoform) transcript increasing from 0.8-3.0 fold (green berry relative to reference) to 15-45 fold (initiation) representing major upregulation, VvME2 showing 10-30 fold induction with slightly delayed timing potentially reflecting isoform-specific regulation, citric acid cycle genes (aconitase, isocitrate dehydrogenase, malate dehydrogenase) maintaining moderate-to-high expression 3-12 fold ensuring pathway capacity for malate oxidation, and transcriptional control through ripening-associated transcription factors possibly including VvNAC family members binding to malic enzyme promoters though specific regulatory mechanisms incompletely characterized. Protein synthesis and enzyme accumulation during initiation demonstrate: malic enzyme protein levels increasing 4-8 fold from green berry to initiation phase correlating with activity increases, enzyme localization predominantly cytosolic with minor mitochondrial forms enabling malate catabolism in both compartments, post-translational modifications potentially including phosphorylation affecting activity though modifications in grape malic enzyme poorly studied, and enzyme stability with half-life estimated 18-36 hours requiring continued synthesis to maintain elevated levels throughout ripening. Substrate availability during initiation includes: vacuolar malate 80-140 mM (green berry) beginning decline to 70-120 mM (initiation) representing bulk storage slowly mobilized, tonoplast malate transporters (potentially VvALMT family) facilitating efflux from vacuole to cytoplasm though specific transporters in grape incompletely identified, cytoplasmic malate 8-20 mM serving as immediate substrate for malic enzyme and malate dehydrogenase, and pH gradients with vacuole pH 2.6-2.9 and cytoplasm pH 7.0-7.4 affecting malate ionization state and transport kinetics. Metabolic fate of malate-derived pyruvate shows: pyruvate from malic enzyme reaction entering multiple pathways including mitochondrial oxidation through pyruvate dehydrogenase producing acetyl-CoA for citric acid cycle, cytosolic metabolism through pyruvate kinase potentially regenerating phosphoenolpyruvate for biosynthesis, conversion to alanine via alanine aminotransferase producing amino acid from carbon skeleton, and fermentative metabolism to lactate or ethanol under hypoxic conditions in berry interior though aerobic pathways dominate near surface. Energy yield from malate catabolism includes: complete oxidation through citric acid cycle yielding 15 ATP per malate (compared to 32 ATP per glucose) making malate moderately efficient respiratory substrate, NADPH from NADP-malic enzyme potentially used for reductive biosynthesis (fatty acid synthesis, antioxidant systems) or oxidized by mitochondria for additional ATP through external NADH dehydrogenase, and malate contribution to total berry energy budget estimated 20-35% during initiation rising to 35-55% during peak decline before declining as malate depletes. Hormonal regulation of malate decline initiation includes: ABA rise from 60-200 ng/g to 700-1800 ng/g triggering ripening program including malic enzyme induction, sugar accumulation potentially creating signaling affecting malate metabolism though mechanisms unclear, declining growth hormones (auxin, gibberellin) removing suppression allowing metabolic shift, and ethylene potentially involved in some cultivars though grape generally non-climacteric with minimal ethylene effects compared to climacteric fruits. Environmental modulation of initiation timing and intensity shows: temperature critically affecting enzyme induction and activity with warm conditions 26-30°C advancing initiation 2-4 days and enhancing enzyme levels 20-40% compared to cool 18-24°C, heat stress >32°C dramatically accelerating initiation potentially causing premature excessive decline, light operating through berry temperature with exposed clusters initiating 1-3 days earlier than shaded fruit, and water status with deficit potentially affecting timing though effects variable and cultivar-dependent. Cultivar genetic variation in malate decline initiation characteristics includes: low-acid cultivars (Grenache, Viognier) showing early rapid initiation with VvME expression 30-60 fold and activity 20-40 units/g within 3-5 days veraison, moderate varieties (Merlot, Chardonnay) displaying 20-40 fold expression and 15-30 units/g in 5-8 days, and high-acid cultivars (Riesling, Chenin Blanc) exhibiting slower initiation 15-30 fold expression and 12-22 units/g over 7-12 days reflecting genetic differences in enzyme regulation and malate metabolism capacity. Physiological consequences of initiation inclu carbon loss through respiratory CO₂ evolution representing 8-18% of green berry malate carbon converted to CO₂ during initiation phase, pH increase in berry juice from 2.85-3.05 to 3.00-3.20 as malate anions removed titrating protons, and redox state changes as NADPH generation from malic enzyme affects cellular reducing capacity potentially influencing antioxidant systems and biosynthetic pathways. Quality implications inclu initiation timing relative to sugar accumulation affecting final sugar-acid balance with early rapid malate loss potentially producing low-acid fruit while delayed initiation maintaining high acids, enzyme activity levels at initiation predicting subsequent decline rates enabling early forecasting of final acidity, and spatial variation in initiation across vineyard revealing maturity heterogeneity requiring management attention. Viticultural factors affecting initiation inclu canopy microclimate with cluster temperature critically important, strategic shading in warm regions delaying initiation 2-5 days and reducing intensity 20-35% preserving acids, vine water status potentially affecting timing with deficit possibly advancing initiation 1-3 days through ABA elevation though effects inconsistent, and crop load with overcropping potentially delaying initiation through resource limitation extending vegetative-reproductive competition. Monitoring initiation in research involves: gene expression analysis tracking VvME transcripts confirming upregulation, enzyme activity assays measuring malic enzyme showing increases, metabolomic profiling tracking malate concentrations and metabolites revealing pathway flux, and respiration measurements showing maintained or elevated rates despite declining respiratory substrate pools indicating metabolic transition.
Malic_Acid_Decline_Initiation is distinguished from overall Acid_Decline_Onset by focusing specifically on malate rather than total acids, from Initial_Acid_Reduction by representing enzymatic activation rather than net reduction, and from later decline by showing onset (8-18% malate loss) rather than complete degradation. VvME induction 12-40 fold, activity 2-6 → 10-25 units/g. Duration 4-7 days BBCH 83. Temperature highly sensitive: >32°C excessive acceleration. Malic_Acid_Decline_Initiation represents critical enzymatic activation enabling acid decline during ripening.
L-Malat-Degradations-Initiierung
L-malate degradation initiation
Malat-Enzym-Induktion
Malat-Katabolismus-Onset
malate catabolism onset
malic enzyme induction
respiratorische Malat-Oxidations-Start
respiratory malate oxidation start
Sweetman C, Deluc LG, Cramer GR, Ford CM, Soole KL (2009) Regulation of malate metabolism in grape berry and other developing fruits Phytochem 70:1329-1344 malate metabolism comprehensive Ruffner HP (1982) Metabolism of tartaric and malic acids in Vitis:a review Vitis 21:247-259 acid metabolism review Terrier N, Sauvage FX, Ageorges A, Romieu C (2001) Changes in acidity and in proton transport at the tonoplast of grape berries during development Planta 213:20-28 acid transport and metabolism.
Malic Acid Decline Initiation
In a Rioja Tempranillo vineyard, malic acid decline start was tracked over 50 days from late August through October Year N+1 (BBCH 83-87) when malate declined from 9.8 g/L to 2.6 g/L (73% total loss, average 0.14 g/L per day), showing initial phase 9.8 → 7.5 g/L (days 0-12, 0.19 g/L per day), rapid phase 7.5 → 3.8 g/L (days 12-32, 0.19 g/L per day sustained), deceleration 3.8 → 2.6 g/L (days 32-50, 0.07 g/L per day), under typical Spanish continental conditions (daily high 26-30°C early season declining to 20-24°C late).
Malic_Acid_Decline_Start represents the onset of sustained progressive malate reduction when respiratory catabolism rates exceed any residual synthesis or import creating net depletion that continues throughout ripening reducing malic acid from 40-60% of total organic acids at onset to 15-35% at harvest fundamentally altering berry acid balance and wine characteristics. This decline start occurs during early-to-mid veraison (BBCH 83-84), typically 2-6 days following malic enzyme activation, and is characterized by malic acid concentrations beginning sustained decrease from 8-14 g/L (80-140 mM) to 6-11 g/L over 8-14 days representing initial 18-30% reduction with decline continuing throughout ripening potentially reducing to 1-5 g/L at harvest (85-95% total loss in extreme cases), malate:tartrate ratio declining from 1.1-2.0:1 at onset toward 0.2-0.8:1 at harvest indicating preferential malate depletion, and respiratory malate contribution increasing from 25-40% of berry CO₂ production to 40-65% during active decline reflecting dominant role in respiratory substrate. Mechanisms driving sustained malate decline inclu malic enzyme activities maintained at 20-60 units/g fresh weight throughout ripening continuously catalyzing malate oxidation, citric acid cycle flux sustained enabling complete malate oxidation to CO₂, respiratory chain maintaining capacity for oxidizing NADH and FADH₂ from malate catabolism generating ATP, and cellular energetics favoring malate catabolism as efficient energy source (15 ATP per malate completely oxidized) supporting ripening processes including sugar transport, biosynthesis, and ion transport. Temporal pattern of malate decline shows: initial phase (BBCH 83-84, days 0-15 post-onset) with moderate loss 18-35% establishing declining trajectory, rapid phase (BBCH 84-85, days 15-35) showing maximum decline rates 0.15-0.45 g/L per day potentially losing 40-60% of onset malate, deceleration phase (BBCH 85-87, days 35-60) with slowing rates 0.08-0.20 g/L per day as substrate depletes, and plateau phase approaching harvest where malate reaches cultivar-specific minimum 1-6 g/L with further decline limited by low substrate availability. Absolute malate loss rates during decline show: early phase 0.10-0.25 g/L per day under moderate conditions (daily high 24-28°C), accelerating to peak rates 0.20-0.50 g/L per day during rapid phase particularly under warm conditions 28-32°C, and declining to 0.05-0.15 g/L per day late phase as malate depletes below 3-6 g/L, with cumulative loss over 40-60 day ripening period potentially 7-12 g/L representing 70-90% of onset malate in low-acid cultivars or warm vintages. Environmental factors critically affecting decline inclu temperature showing strong linear or exponential relationship with loss rate with each 1°C increase above 20°C accelerating decline approximately 8-15%, heat stress >32-35°C causing dramatic acceleration potentially doubling or tripling normal rates producing excessive loss and unbalanced fruit, cool conditions 5-7 g/L requiring winery intervention (cold stabilization, deacidification, extended malolactic) adding cost and potentially affecting quality, and malate:tartrate ratio at harvest influencing malolactic fermentation kinetics with high malate (>3-4 g/L) favoring rapid complete MLF while low malate potentially resulting in sluggish or stuck fermentation. Sensory implications inclu malic acid contributing sharp tart taste distinct from tartaric acid's more neutral acidity so high residual malate (>4-5 g/L) producing aggressive tartness, balanced malate levels 2-4 g/L contributing to wine freshness and structure, and low malate <2 g/L potentially leaving wines flat especially if tartaric also moderate creating overall acid deficiency. Viticultural management optimizing malate decline includes: canopy management strategically shading clusters in warm regions slowing decline 25-50% preserving final acids, irrigation timing and amount potentially moderating berry temperature through transpirational cooling, harvest timing decisions balancing sugar accumulation against continued acid loss particularly critical in warm vintages where excessive decline threatens quality, and rootstock selection with some stocks associated with higher berry malate retention though mechanisms incompletely defined.
Malic_Acid_Decline_Start is distinguished from Malic_Acid_Decline_Initiation by representing sustained ongoing reduction (18-30% initial, up to 85-95% total) rather than enzymatic onset, from Initial_Acid_Reduction by showing specific malate focus rather than total acids, and from acid plateau by exhibiting active decline rather than stabilization. Duration entire ripening period 40-60 days. Loss rates: 0.10-0.50 g/L per day (temperature-dependent). Final levels: low-acid 1-3 g/L, moderate 2-5 g/L, high-acid 4-7 g/L. Malic_Acid_Decline_Start represents sustained process fundamentally altering berry acid balance affecting wine quality and style.
anhaltende Malat-Verminderung
continuous malate catabolism
fortlaufende L-Malat-Reduktion
kontinuierlicher Malat-Katabolismus
ongoing L-malate reduction
progressive malic acid loss
progressive Äpfelsäure-Verlust
sustained malate depletion
Keller M (2020) The Science of Grapevines 3rd Ed pages 229-233 malate decline during ripening Sweetman C, Sadras VO, Hancock RD, Soole KL, Ford CM (2014) Metabolic effects of elevated temperature on organic acid degradation in ripening Vitis vinifera fruit J Exp Bot 65:5975-5988 temperature effects comprehensive study Lakso AN, Kliewer WM (1975) The influence of temperature on malic acid metabolism in grape berries Plant Physiol 56:370-372 classic temperature study.
Malic Acid Decline Start
In Riesling berries in Rheingau, Germany during BBCH 85-89, malic acid degradation reduces malate from 12.8 g/L at veraison to 6.2 g/L at harvest over 52 days, representing 52% degradation characteristic of this high-acid variety, with NADP-malic enzyme contributing approximately 60% of total degradation, mitochondrial respiration 25%, and gluconeogenesis 15%, under cool mean temperatures of 16°C that preserve higher residual acidity than warmer regions.
Malic acid degradation constitutes the metabolic process hierarchy encompassing all enzymatic and respiratory pathways responsible for the catabolism of L-malic acid in Vitis vinifera berries during ripening, representing the mechanistic framework through which initial malic acid concentrations of 8-15 g/L at veraison are systematically reduced to final levels of 1-4 g/L at harvest through mitochondrial respiration, cytosolic enzyme activity, and gluconeogenic conversion over the 30-50 day post-veraison maturation period. This process class organizes the specific biochemical mechanisms including respiratory consumption of malate through the tricarboxylic acid cycle, NADP-malic enzyme catalyzed decarboxylation producing pyruvate and CO2, and phosphoenolpyruvate carboxykinase mediated gluconeogenesis recycling malate carbon into hexose sugars, collectively accounting for the near-complete turnover of the berry malate pool during ripening. The subcellular organization involves coordinated transport and metabolism across multiple compartments, with vacuolar malate stores serving as the primary substrate reservoir, cytosolic enzymes providing the dominant degradation pathway, and mitochondrial metabolism contributing both catabolic capacity and energy generation through oxidative phosphorylation coupled to malate oxidation. Genetic regulation of malic acid degradation involves developmental activation of malic enzyme gene expression at veraison, hormonal modulation by abscisic acid enhancing enzyme activities, and cultivar-specific differences in regulatory element sequences determining the magnitude and timing of enzyme induction during ripening. The quantitative importance of malic acid degradation for berry carbon balance reflects the substantial carbon pool represented by malate, with complete degradation of 10 g/L malic acid releasing approximately 3.6 g carbon equivalent per liter of berry volume, contributing significantly to respiratory CO2 efflux and providing substrate for secondary metabolite biosynthesis during ripening. Environmental modulation occurs primarily through temperature effects on enzyme kinetics and respiratory rates, with degradation rates doubling for each 10°C temperature increase within the physiological range, establishing malic acid degradation as the primary temperature-responsive component of berry acid metabolism. The process demonstrates characteristic kinetics with initial rapid degradation during early ripening when enzyme activities peak and substrate concentrations remain high, transitioning to slower rates during late ripening as malate pools become depleted and enzyme activities decline.
Malic_Acid_Degradation is parent class under Acid_Reduction organizing biochemical mechanisms of malate catabolism. Distinguishes from Active_Acid_Reduction which emphasizes temporal dynamics. Contains subclasses Malate_Respiratory_Consumption, Malic_Enzyme_Activity, and Progressive_Malate_Depletion representing specific mechanisms and outcomes. Duration 30-50 days. Key parameter is 50-90% malic acid reduction depending on cultivar and climate.
L-Malat-Katabolismus
L-malate catabolism
Malat-Umsatz
malate turnover
malic acid breakdown process
malic acid metabolic degradation
Äpfelsäure-metabolischer-Abbau
Sweetman C et al (2009) Regulation of malate metabolism in grape berry. Molecular Plant 2: 1270-1280 Ruffner HP (1982) Metabolism of tartaric and malic acids in Vitis. Vitis 21: 346-358 Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 268-275
Malic Acid Degradation
Malic acid degradation during ripening follows first-order kinetics with rate constants (k) ranging from 0.02-0.08 per day across cultivars, resulting in 50-90% malate loss between veraison and harvest over 40-60 days. Temperature exerts profound influence, with degradation rates doubling for each 10°C increase (Q₁₀ of 1.8-2.2) above the base temperature of approximately 10°C. Grenache shows rapid malate degradation (k = 0.06-0.08/day, 80-90% loss) while Riesling shows slow degradation (k = 0.02-0.03/day, 50-65% loss) under equivalent temperature conditions. The cytosolic NADP-malic enzyme (VvNADP-ME) decarboxylates malate to pyruvate, which enters mitochondrial respiration or gluconeogenic conversion to sugars (contributing 5-15% of berry sugar at maturity).
An acid metabolism disposition that is the inherited capacity of a Vitis vinifera plant to catabolize malic acid in ripening grape berries through mitochondrial respiration and gluconeogenesis, with genotype-specific degradation rates determined by malic enzyme activity, mitochondrial density, and respiratory demand during the post-veraison period.
Malic acid degradation is the primary mechanism by which climate affects grape acidity, as warm ripening temperatures accelerate respiration and gluconeogenesis of stored malate. The trait creates the characteristic acid profile differences between cool climate wines (high malate retention, fresh acidity) and warm climate wines (low malate, softer acidity). Breeding for reduced malate degradation rate is a strategy for maintaining acidity under warming conditions. Night temperature is particularly important, as high night temperatures sustain respiration that cool nights would suppress.
Malatrespiration
malate catabolism trait
malate respiration capacity
malic acid breakdown
Äpfelsäure-Degradation
Äpfelsäureabbau
Sweetman C et al. (2009) Transcriptome analysis at four developmental stages of grape berry (Vitis vinifera cv. Shiraz) provides insights into regulated and coordinated gene expression. BMC Genomics 10:648 Lakso AN, Kliewer WM (1975) The influence of temperature on malic acid metabolism in grape berries. Plant Physiology 56:370-372
Malic Acid Degradation Disposition
In Grenache berries in Châteauneuf-du-Pape, Rhône Valley during BBCH 85-89, malic enzyme activity increases from 0.8 µmol/min/g fresh weight at veraison to peak activity of 4.2 µmol/min/g fresh weight at 18 days post-veraison, catalyzing degradation of 6.8 g/L malic acid representing 65% of total malate catabolism, with cytosolic NADP-malic enzyme contributing 85% of measured activity under warm ripening conditions averaging 28°C.
Malic enzyme activity encompasses the catalytic function of NADP-dependent and NAD-dependent malic enzymes in Vitis vinifera berry cells during ripening, representing the quantitatively dominant pathway of malic acid degradation wherein L-malate undergoes oxidative decarboxylation to pyruvate with concurrent reduction of NAD(P)+ to NAD(P)H and release of CO2, accounting for 50-70% of total malate catabolism over the 30-45 day post-veraison ripening period. The cytosolic NADP-malic enzyme provides the primary degradation pathway, with enzyme activity increasing 3-6 fold between veraison and mid-ripening in response to developmental signals and abscisic acid induction, reaching peak activities of 2-5 µmol/min/g fresh weight before declining during late ripening as substrate becomes limiting. The biochemical mechanism involves sequential binding of malate and NADP+ to the enzyme active site, hydride transfer from malate C2 to NADP+ generating the intermediate oxaloacetate, and spontaneous or enzyme-assisted decarboxylation releasing CO2 and producing pyruvate as the stable product, with the overall reaction thermodynamically favorable and essentially irreversible under cellular conditions. Mitochondrial NAD-malic enzyme provides a secondary pathway with distinct kinetic properties and regulatory characteristics, contributing 10-20% of total malic enzyme flux with activity responsive to cellular energy status through adenylate regulation. The pyruvate produced by malic enzyme activity feeds into multiple downstream pathways including mitochondrial oxidation through pyruvate dehydrogenase, gluconeogenic reversal through pyruvate carboxylase, and amino acid biosynthesis through transamination, integrating malate degradation with broader cellular metabolism. Genetic variation in malic enzyme expression underlies cultivar differences in acid retention, with high-acid varieties such as Riesling showing lower enzyme induction and activity compared to rapid acid-decline varieties such as Grenache under equivalent conditions. Temperature modulation of malic enzyme activity occurs through both direct kinetic effects on catalytic rate and indirect effects on gene expression, with warm conditions stimulating both enzyme synthesis and activity to produce accelerated malate catabolism. The NADPH generated by cytosolic malic enzyme activity contributes to cellular reducing power for biosynthetic reactions including secondary metabolite modification and lipid synthesis during berry ripening.
Malic_Enzyme_Activity is subclass of Malic_Acid_Degradation representing the dominant enzymatic pathway. Distinguishes from Malate_Respiratory_Consumption which is mitochondrial TCA oxidation. Sibling to Malate_Respiratory_Consumption and Progressive_Malate_Depletion. Accounts for 50-70% of total malate catabolism. Peak activity 2-5 µmol/min/g fresh weight at mid-ripening. Key cultivar-differentiating parameter.
ME enzyme activity
Malat-Decarboxylierungs-Aktivität
Malic-Enzym-Aktivität
NADP-Malic-Enzym-Funktion
NADP-malic enzyme function
malate decarboxylation activity
malic enzyme catalysis
Äpfelsäure-Enzym-Katalyse
Sweetman C et al (2009) Regulation of malate metabolism in grape berry. Molecular Plant 2: 1270-1280 Ruffner HP (1982) Metabolism of tartaric and malic acids in Vitis. Vitis 21: 346-358 Or E et al (2000) Isolation of a NAD-dependent malic enzyme gene from grape berries. Plant Science 159: 1-9
Malic Enzyme Activity
Malvidin content quality in Monastrell (Mourvèdre) from Jumilla, Spain reaches exceptionally high levels (1200-1800 mg/kg malvidin-3-glucoside equivalents) reflecting the variety's intense pigmentation in the region's hot, dry climate, with malvidin-based compounds comprising 85-92% of total anthocyanins this malvidin dominance produces deeply colored, age-worthy wines with stable color development, enabling the inky, concentrated style that has established Jumilla Monastrell as a benchmark for Mediterranean red wine intensity and value.
Malvidin_Content_Quality represents the chemical quality characterizing the concentration of malvidin-based anthocyanins in grape berry skins, capturing the dominant anthocyanidin class in most Vitis vinifera red wine cultivars. Malvidin (3',5'-dimethoxy-3,4',5,7-tetrahydroxyflavylium) is distinguished from other anthocyanidins by methoxy groups at both the 3' and 5' positions of the B ring, representing the most methylated anthocyanidin structure. This high degree of methylation confers greater chemical stability compared to hydroxylated anthocyanidins, contributing to the preferential accumulation of malvidin-based pigments in grape skins and their persistence in aged red wines. Malvidin-3-glucoside is typically the single most abundant anthocyanin in red grapes, often comprising 25-60% of total anthocyanins alone, with malvidin-3-(6-acetyl)-glucoside and malvidin-3-(6-p-coumaroyl)-glucoside adding substantial additional contributions. The proportion of malvidin among total anthocyanins varies with cultivar, ranging from approximately 50% in Pinot Noir to greater than 90% in some varieties. This variation reflects differences in the activity of O-methyltransferase enzymes that convert delphinidin and petunidin to their methylated forms. The malvidin proportion provides a varietal fingerprint useful for authenticity assessment, as ratios among anthocyanidins remain relatively stable across vintages and growing conditions. Malvidin-based anthocyanins undergo co-pigmentation reactions with flavonols and other phenolics that enhance and stabilize color, and participate in condensation reactions with tannins that produce the stable polymeric pigments responsible for aged wine color.
Malvidin_Content_Quality specifically addresses malvidin-based anthocyanins, including the primary glucoside and its acylated derivatives (acetyl, p-coumaroyl, caffeoyl). The class represents the largest single anthocyanidin contribution in most red wine grapes, making it the primary determinant of total anthocyanin content. Malvidin content can be expressed as malvidin-3-glucoside equivalents (using its molar absorptivity for quantification) or as the sum of individually quantified malvidin-based compounds. The stability of malvidin relative to other anthocyanidins makes it an important contributor to wine aging potential, with malvidin-tannin adducts forming the stable pigmented polymers of aged wine.
Malvidin-3-glucosid-Konzentration
Malvidingehalt
Malvidinpigmentqualität
Mv-3-glc content
dimethoxylated anthocyanin concentration
dimethoxylierter Anthocyangehalt
malvidin pigment quality
malvidin-3-glucoside content
Roggero JP et al. (1986) Study of the Anthocyanins of Vitis vinifera L. Using Column Chromatography and Spectroscopy. Sciences des Aliments 6(4): 567-579 Mattivi F et al. (2006) Metabolite Profiling of Grape: Flavonols and Anthocyanins. Journal of Agricultural and Food Chemistry 54(20): 7692-7702
Malvidin Content Quality
Malvidin-3-O-glucoside predominates in most V. vinifera cultivars, comprising 50-90% of total anthocyanins in Cabernet Sauvignon (60-75%), Merlot (55-70%), Syrah (45-60%), and Tempranillo (40-55%). Malvidin accumulation requires sequential activity of F3'5'H (converting dihydrokaempferol to dihydromyricetin), DFR, ANS, UFGT, and finally OMT (O-methyltransferase) for the two methoxy groups on the B-ring. The 3',5'-disubstituted, di-methoxylated structure of malvidin confers blue-shifted color (λmax 520-535 nm versus 506-513 nm for cyanidin) and enhanced stability against oxidation and pH-induced color loss. Wine color derived from malvidin-dominant grapes shows characteristic purple-red hues that evolve toward brick-red with aging through anthocyanin-tannin polymerization.
An anthocyanin accumulation disposition that is the inherited capacity of a Vitis vinifera plant to accumulate malvidin-based anthocyanins (malvidin-3-O-glucoside and its acylated derivatives) as the predominant pigment class, typically comprising 40-90% of total anthocyanins due to high flavonoid 3',5'-hydroxylase (F3'5'H) and O-methyltransferase (OMT) activity.
Malvidin dominance is characteristic of V. vinifera and distinguishes it from most hybrid cultivars that show higher cyanidin proportions. The trait results from the coordinated expression of F3'5'H (directing flux toward delphinidin pathway) and OMT (methylating to petunidin and malvidin). Malvidin-based anthocyanins show greater color stability than cyanidin-based forms, contributing to the superior aging potential of V. vinifera wines. The relative proportion of malvidin versus other anthocyanins is used in authenticity testing to detect hybrid blending.
Malvidin-Dominanz
Malvidin-Vorherrschaft
Malvidinprävalenz
malvidin predominance
malvidin-type pigmentation
tri-substituted anthocyanin trait
Mattivi F et al. (2006) Metabolite profiling of grape: flavonols and anthocyanins. Journal of Agricultural and Food Chemistry 54:7692-7702 Castellarin SD et al. (2006) Colour variation in red grapevines (Vitis vinifera L.): genomic organisation, expression of flavonoid 3'-hydroxylase, flavonoid 3',5'-hydroxylase genes and related metabolite profiling of red cyanidin-/blue delphinidin-based anthocyanins in berry skin. BMC Genomics 7:12
Malvidin Dominance Disposition
Manganese content quality assessment in Margaret River Chardonnay reveals the interaction between soil pH and micronutrient availability, with the region's acidic lateritic soils (pH 5.0-6.0) providing adequate manganese (must concentration 1.5-3.5 mg/L) in contrast to the manganese deficiency affecting Chardonnay on limestone-derived soils in Burgundy this manganese sufficiency supports the photosynthetic efficiency required for the even ripening and flavor development that characterize premium Margaret River Chardonnay without the interveinal chlorosis that limits productivity in manganese-deficient sites.
Manganese_Content_Quality represents the chemical quality characterizing the concentration of manganese (Mn²⁺) in grape berry tissue, capturing the essential micronutrient central to photosynthetic oxygen evolution, enzyme activation, and antioxidant defense systems. Manganese in grape juice typically ranges from 0.5-5 mg/L, with variation reflecting cultivar, soil conditions, and the pH-dependent chemistry governing manganese availability. The critical role of manganese in the photosystem II oxygen-evolving complex makes it essential for photosynthesis, while its function in superoxide dismutase and various decarboxylases links it to stress response and organic acid metabolism. Manganese deficiency produces interveinal chlorosis similar to iron deficiency but typically appearing on middle-aged leaves rather than youngest leaves, with green color persisting as small dots or patches within yellowed interveinal tissue. Soil manganese availability decreases sharply with increasing pH, creating deficiency risk in calcareous and overlimed soils, while waterlogged conditions can produce toxic manganese concentrations through reduction of insoluble Mn⁴⁺ oxides to soluble Mn²⁺. The interaction between manganese and iron involves competitive uptake and metabolic antagonism, with excess of either element potentially inducing deficiency of the other. In winemaking, manganese participates in various enzyme reactions but does not create stability problems at normal concentrations. Wine manganese content reflects must composition modified by precipitation and yeast uptake, with typical levels of 1-3 mg/L. The oxidation state changes of manganese (Mn²⁺/Mn³⁺/Mn⁴⁺) differ from the simple ferrous-ferric couple of iron, reducing manganese's role in wine oxidation chemistry relative to iron.
Manganese_Content_Quality addresses a micronutrient essential for photosynthetic oxygen evolution, distinguishing its function from the electron transport roles of iron. The pH sensitivity of manganese availability creates opposite patterns from most micronutrients, with toxicity risk in acidic soils versus deficiency risk in alkaline soils. The distinctive deficiency symptom pattern (middle-aged leaf chlorosis with stippled green dots) aids diagnosis. Manganese does not play the same wine oxidation role as iron and copper, reducing its winemaking significance relative to those metals.
Beerenmangangehalt
Mangangehalt
Mangankonzentration
Mn content
Mn-Gehalt
berry manganese quality
manganese concentration
manganese level
Marschner H (2012) Marschner's Mineral Nutrition of Higher Plants, 3rd Ed, Academic Press pp.: 191-248 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 341-385
Manganese Content Quality
Manganese deficiency symptom quality diagnosis in Champagne Chardonnay on chalky soils distinguished Mn limitation from concurrent iron chlorosis to enable targeted corrective management for both micronutrient deficiencies. Visual manganese deficiency symptom quality assessment revealed interveinal chlorosis on young leaves similar to iron deficiency but with characteristic differences: the interveinal tissue showed a more greenish-yellow rather than the bright yellow to white of iron chlorosis, the pattern appeared more diffuse with less sharp contrast between veins and interveinal areas, and small tan necrotic flecks (1-2 mm) had developed within chlorotic zones on the most affected leaves, a feature typical of manganese but not iron deficiency. The distribution of manganese deficiency symptom quality confirmed immobile nutrient involvement with symptoms restricted to the apical 5-6 leaves while basal leaves remained unaffected, distinguishing from magnesium deficiency that would affect oldest leaves first. Tissue analysis quantified the manganese deficiency symptom quality with young leaf Mn at 15 ppm (deficient, threshold 25 ppm) while Fe was also low at 48 ppm (marginal), establishing concurrent deficiency of both elements as expected on this pH 8.3 soil. The manganese deficiency symptom quality correlated with photosynthetic measurements showing 25% reduction in maximum photosynthesis rate in symptomatic versus asymptomatic leaves, demonstrating the functional impact of Mn limitation on the oxygen-evolving complex of photosystem II. Corrective foliar application of MnSO₄ at 0.3% concentration resolved manganese deficiency symptom quality within 3 weeks, with new growth emerging with normal dark green coloration and chlorotic symptoms fading on previously affected leaves.
Manganese deficiency symptom quality is a micronutrient deficiency symptom quality that inheres in a grapevine organism and represents the characteristic visual symptoms resulting from inadequate manganese availability for photosynthesis, enzyme activation, and antioxidant defense, including interveinal chlorosis on young leaves where tissue between veins becomes pale green to yellow while veins remain darker green, reduced photosynthetic capacity, and in severe cases development of small necrotic spots within the chlorotic interveinal areas, with manganese deficiency being most common on high-pH calcareous soils and overlimed acidic soils where manganese solubility is reduced to levels limiting plant uptake. This quality captures the grapevine response to manganese limitation, recognizing that while manganese deficiency is less common than iron deficiency in viticulture, it produces similar interveinal chlorosis symptoms that require careful differential diagnosis, and can occur concurrently with iron deficiency on calcareous soils where both elements have reduced availability. The ontological significance of manganese deficiency symptom quality lies in its diagnostic challenge requiring differentiation from iron and magnesium deficiency that produce similar interveinal chlorosis patterns, and in its potential to limit photosynthetic capacity and vine productivity when severe. The biological basis of manganese deficiency symptoms encompasses the essential role of manganese in the oxygen-evolving complex of photosystem II where four manganese atoms catalyze water splitting, the function of manganese as activator of numerous enzymes including those involved in phenylpropanoid biosynthesis and reactive oxygen species detoxification, and the role of manganese in chloroplast structure and function that, when impaired by deficiency, reduces chlorophyll accumulation in interveinal mesophyll. The phloem immobility of manganese means that deficiency symptoms appear first on youngest leaves that cannot acquire manganese through redistribution from older tissues, creating the characteristic acropetal symptom progression that distinguishes manganese from magnesium deficiency despite similar interveinal chlorosis patterns. The soil chemistry of manganese shows strong pH dependence with availability decreasing dramatically above pH 7.0 as manganese oxidizes to insoluble Mn⁴⁺ forms, making high-pH conditions the primary predisposing factor for manganese deficiency.
Manganese_Deficiency_Symptom_Quality is_a Micronutrient_Deficiency_Symptom_Quality, serving as parent class for Interveinal_Chlorosis_Quality_(Mn) which addresses the specific chlorosis pattern. This class addresses the overall manganese deficiency syndrome. Immobile nutrient with symptoms on youngest leaves (distinguishing from Mg). Interveinal chlorosis similar to Fe but with greenish-yellow color and necrotic flecks. Young leaf Mn threshold ~25 ppm. High pH (>7.0) reduces Mn availability through oxidation. Often co-occurs with Fe deficiency on calcareous soils. Essential for photosystem II oxygen evolution.
Manganmangel-Symptome
Manganstress-Indikatoren
Mn deficiency symptoms
Mn-Mangel-Zeichen
Niedrig-Mangan-Symptome
low manganese symptoms
manganese limitation signs
manganese stress indicators
Manganese Deficiency Symptom Quality
Manganese uptake shows moderate rootstock variation (1.5-2× range), with SO4 and 5BB Kober demonstrating higher Mn accumulation than 41B and 140 Ruggeri under equivalent soil conditions. Manganese deficiency symptoms (interveinal chlorosis of young leaves with green veins, distinct from Mg deficiency in older leaves) appear when petiole Mn falls below 20-25 ppm. Adequate petiole Mn ranges from 30-150 ppm, with toxicity symptoms (brown spotting, marginal necrosis) appearing above 500 ppm in acidic, waterlogged soils. The Mn²⁺/Mn⁴⁺ redox couple is highly pH-dependent, with Mn availability increasing 100-fold between pH 7.5 and 5.5, creating toxicity risk on acidic soils and deficiency on calcareous soils.
A micronutrient uptake disposition that is the inherited capacity of a Vitis vinifera plant to absorb manganese ions (Mn²⁺) from the soil solution through NRAMP family transporters, with genotype-specific efficiency affecting photosynthetic oxygen evolution, enzyme activation, and antioxidant capacity.
Manganese uptake efficiency is less critical for rootstock selection than iron or zinc due to adequate Mn availability in most vineyard soils. However, Mn toxicity on acidic soils and Mn deficiency on high-pH soils create localized problems requiring attention. Manganese functions in the oxygen-evolving complex of photosystem II, making it essential for photosynthetic electron transport. The element also activates numerous enzymes and contributes to antioxidant defense through Mn-superoxide dismutase.
Mangananeignungsvermögen
Manganaufnahme
Mn uptake capacity
Mn-Akquisition
manganese acquisition
manganese foraging trait
Pearson RC, Goheen AC (1988) Compendium of Grape Diseases, APS Press pp.: 89-92 Millaleo R et al. (2010) Manganese as essential and toxic element for plants: transport, accumulation and resistance mechanisms. Journal of Soil Science and Plant Nutrition 10:470-481
Manganese Uptake Disposition
GF.GA-47-42 (a complex interspecific hybrid carrying Rpv3 downy mildew resistance) bears a Mapping_Parent_Role in crosses with Villard Blanc and other susceptible parents, its heterozygous resistance loci segregating in F1 progeny to enable QTL mapping of resistance. Cabernet Sauvignon serves in a Mapping_Parent_Role across multiple mapping populations (× Norton, × Regent, × other genotypes), contributing segregating alleles for quality traits, phenology, and fruit composition—its importance as a mapping parent reflecting both its commercial significance and high heterozygosity. Muscadinia rotundifolia functions as mapping parent for Run1/Ren1 powdery mildew resistance mapping, its unique immunity segregating in interspecific progeny to enable fine-mapping of resistance loci.
A mapping population role that inheres in a grapevine genotype serving as a parent of a genetic mapping population, contributing segregating alleles whose inheritance in progeny enables construction of genetic linkage maps and association of marker genotypes with phenotypic variation. Mapping parents are selected for complementary characteristics—typically one parent carrying traits of interest (disease resistance, distinctive quality attributes) and the other providing contrasting phenotypes enabling segregation analysis, with both parents ideally highly heterozygous to maximize marker informativeness. The mapping parent's genetic contribution defines what can be discovered in the population resistance loci can only be mapped if the resistant parent is heterozygous at those loci (enabling segregation), while loci homozygous in both parents are uninformative for that population. Mapping parents are typically maintained vegetatively, enabling repeated phenotyping alongside their progeny and serving as reference genotypes for population analysis. Parental marker heterozygosity is assessed prior to population development, with parents selected for high polymorphism rates ensuring adequate marker coverage for linkage mapping.
Mapping_Parent_Role emphasizes the genetic contribution function distinguishing parents from progeny members in mapping population context. The pseudo-testcross design used in grapevine mapping generates separate linkage maps for each parent (based on markers heterozygous in that parent), later integrated into consensus maps. Parental selection profoundly influences mapping outcomes—only loci variable between parents can be detected, and only segregating parental alleles can be mapped. The choice of crossing parents balances trait contrast (maximizing segregation for target traits), marker polymorphism (ensuring adequate map density), and practical considerations (crossing compatibility, seed production). Key mapping parents appear across multiple populations, enabling cross-population comparison and meta-analysis of QTL effects.
Kartierungselter
Populationselter
QTL mapping parent
QTL-Elter
cross parent for mapping
mapping parent
population parent
Adam-Blondon AF et al. (2011) Genetic mapping in grapes, CRC Press pp.: 119-159 Zyprian E et al. (2016) Quantitative trait loci, Molecular Genetics and Genomics 291: 1573-1594 Riaz S et al. (2011) Powdery mildew resistance loci, Theoretical and Applied Genetics 122: 1059-1073 Fechter I et al. (2012) Candidate genes for QTL, Molecular Genetics and Genomics 287: 159-174
Mapping Parent Role
The GF.GA-47-42 × Villard Blanc F1 population bears a Mapping_Population_Role as one of the foundational populations for grapevine resistance QTL mapping, with over 150 progeny genotyped for thousands of markers and phenotyped for downy and powdery mildew resistance—enabling identification of major Rpv and Ren/Run loci. A Cabernet Sauvignon × Norton cross population serves in a Mapping_Population_Role for Pierce's disease resistance mapping, with the American hybrid Norton contributing PdR loci detectable through marker-trait association in the segregating progeny. The Syrah × Grenache population bears a Mapping_Population_Role for berry composition and phenology QTL mapping, with Mediterranean-adapted parents contributing variation for fruit quality traits.
A breeding role that inheres in a grapevine genotype by virtue of its participation in a genetic mapping population—a structured population derived from controlled crosses, designed to enable identification of genomic regions (quantitative trait loci, QTL) associated with traits of interest through the correlation of molecular marker genotypes with phenotypic variation. Mapping populations are fundamental tools for dissecting the genetic basis of complex traits (disease resistance, fruit quality, phenology), with population members serving as data points linking segregating marker alleles to trait variation. Grapevine mapping populations are typically F1 populations from biparental crosses (pseudo-testcross design exploiting the highly heterozygous nature of grape), though backcross and multi-parent populations have also been developed. The mapping population provides the genetic framework for QTL detection—markers segregating in the population define a genetic linkage map, while phenotypic measurements on population members enable statistical association between marker genotypes and trait values. Key grapevine mapping populations (GF.GA-47-42 × Villard Blanc, Cabernet Sauvignon × Norton, Syrah × Grenache, and others) have enabled identification of major resistance loci and quality-associated QTL underlying contemporary marker-assisted breeding.
Mapping_Population_Role captures the research function of populations designed for genetic analysis rather than direct variety development, though mapping population members may also serve as breeding parents. The pseudo-testcross strategy (exploiting heterozygosity within each parent to generate segregating markers) is standard for grapevine mapping given the difficulty of generating inbred lines due to inbreeding depression. Mapping population size affects QTL detection power—populations of 100-200 individuals are common, enabling detection of major-effect QTL but limiting resolution for minor QTL. High-density marker genotyping (SSR, then SNP arrays, now GBS/sequencing approaches) has progressively improved mapping resolution, with QTL intervals narrowing from tens of cM to sub-cM regions enabling candidate gene identification. Mapping populations represent long-term research investments, with established populations phenotyped repeatedly for diverse traits over years to decades.
Kartierungspopulation
QTL mapping population
QTL-Population
genetic mapping population
genetische Kartierungspopulation
mapping population
segregating population
segregierende Population
Adam-Blondon AF et al. (2011) Genetic mapping in grapes, in Bentley R (ed) Genetics, Genomics, and Breeding of Grapes, CRC Press pp.: 119-159 Töpfer R et al. (2011) Molecular breeding in grapes, CRC Press pp.: 160-185 Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance, Molecular Genetics and Genomics 291: 1573-1594 Riaz S et al. (2011) Identification of powdery mildew resistance loci, Theoretical and Applied Genetics 122: 1059-1073
Mapping Population Role
Margin shape in Nebbiolo displays the characteristic combination of large teeth with convex sides that creates the distinctive saw-tooth edge profile recognized by Piemontese vignerons, with primary teeth of 3-5 mm height spaced at 8-12 mm intervals along the major veins and smaller secondary teeth of 1-2 mm between them this conspicuous marginal serration distinguishes Nebbiolo from the finer-toothed Barbera commonly planted in adjacent vineyards and provides a rapid field identification character even on young, still-expanding leaves.
Margin_Shape_Quality represents the morphological quality characterizing the edge configuration of laminar grapevine organs, primarily leaves. The margin constitutes the boundary between the organ interior and the surrounding environment, and its shape reflects both the pattern of marginal growth and the extent of post-formational modification through cell expansion and death. In grapevine leaves, margin shape provides important ampelographic characters because the serration pattern (teeth) along the blade edge varies consistently among cultivars. Margin shape encompasses several aspects: the presence or absence of teeth, the size and density of teeth, the shape of individual teeth (pointed, rounded, convex-sided, concave-sided), and the regularity of the serration pattern. OIV descriptors 077 (tooth shape) and 078 (tooth size) provide standardized categorical assessment of leaf margin characters. Margin shape also applies to the configuration of the lateral sinuses that create the lobed structure of most grape leaves, though these larger-scale indentations are typically treated separately from marginal teeth. The developmental origin of marginal teeth involves localized cell division and expansion at the leaf primordium edge, with genetic factors controlling the spacing and morphology of teeth. Margin shape assessment employs visual comparison to standardized references, measurement of tooth height and spacing, or outline analysis capturing the detailed edge profile. Margin characters tend to be stable within cultivars and provide reliable identification features.
Margin_Shape_Quality is positioned as a direct subclass of Shape_Quality and serves as the parent class for specific margin configurations including Entire_Margin_Quality, Serrate_Margin_Quality, and Dentate_Margin_Quality. The class applies primarily to leaves, as berry and shoot surfaces lack the marginal teeth that characterize this quality category. Margin shape should be assessed on fully expanded, mature leaves from the middle portion of the shoot to ensure standardized developmental stage. The relationship between margin shape and leaf hydraulics (teeth may function as hydathodes for guttation) provides functional context beyond pure morphological description.
Blattrandqualität
Kantenbeschaffenheit
Randform
Randkonfiguration
boundary configuration
edge shape
leaf margin quality
peripheral shape
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 55-56 Royer DL et al. (2009) Why Do Toothed Leaves Correlate with Cold Climates? Gas Exchange at Leaf Margins Provides New Insights into a Classic Paleotemperature Proxy. International Journal of Plant Sciences 170(1): 73-89
Margin Shape Quality (for leaves)
Marginal leaf necrosis quality assessment for potassium deficiency in Walla Walla Valley Cabernet Sauvignon documented the characteristic symptom progression enabling field diagnosis and targeted management response. Early symptom development showed marginal leaf necrosis quality initiating as subtle marginal bronzing on the oldest (basal) leaves at approximately 4 weeks post-véraison, with affected tissue showing the distinctive tan-to-brown coloration beginning at the leaf edge while interveinal and midrib regions remained green. Symptom progression over the following 4 weeks showed marginal leaf necrosis quality advancing both acropetally (from basal to mid-canopy leaves) and centripetally (from margins toward midrib), with severely affected leaves showing necrotic bands 2-3 cm wide along the entire margin while central portions remained functional. Quantitative scoring of marginal leaf necrosis quality used percentage of leaf margin affected, with values ranging from 10% (early symptoms, marginal bronzing only) through 40-60% (moderate, brown necrosis on most of margin) to greater than 80% in severe cases where necrosis extended well into the interveinal regions. The spatial distribution of marginal leaf necrosis quality within the vineyard correlated with soil texture, with sandy ridgetop positions showing severe symptoms (mean 65% margin affected) while heavier-textured swale positions showed minimal symptoms (mean 12%), reflecting the lower cation exchange capacity and potassium retention of sandy soils. Tissue analysis confirmed the marginal leaf necrosis quality diagnosis, with symptomatic zones showing petiole K of 0.85% at harvest versus 1.65% in asymptomatic areas.
Marginal leaf necrosis quality in potassium deficiency is a potassium deficiency symptom quality that inheres in grapevine leaves and represents the characteristic death of leaf margin tissue resulting from potassium-limited osmoregulation and enzyme function, with the necrosis typically initiating as bronzing or yellowing at the leaf edge, progressing to brown desiccated tissue that may curl or become brittle, and advancing from margins toward the midrib in severe cases, with the pattern of oldest leaves affected first reflecting potassium mobility and remobilization to younger tissues. This quality captures the most diagnostically distinctive potassium deficiency symptom, recognizing that marginal necrosis (often termed "leaf scorch") provides reliable field identification of potassium limitation when the characteristic pattern is recognized, though confirmation through tissue analysis is recommended before corrective fertilization. The ontological significance of marginal leaf necrosis quality in potassium deficiency lies in its visual diagnostic utility enabling identification of potassium limitation in the field, with the marginal distribution distinguishing potassium deficiency from the interveinal patterns of magnesium or manganese deficiency and from the uniform chlorosis of nitrogen deficiency. The biological basis of potassium-deficiency marginal necrosis encompasses the concentration of metabolic stress at leaf margins where water loss through transpiration is highest and potassium's osmoregulatory function most critical, the accumulation of putrescine and other toxic metabolites in potassium-deficient tissues, and the progressive cell death spreading from margins as potassium depletion intensifies. The seasonal timing of marginal necrosis development typically shows acceleration during the post-véraison period when fruit potassium demand competes with vegetative tissue maintenance, with symptoms that may have been subtle or absent during early season rapidly intensifying as fruit accumulation depletes vine potassium reserves, making late-season symptom expression particularly severe in years with high fruit loads or on soils with marginal potassium supply.
Marginal_Leaf_Necrosis_Quality_(K) is_a Potassium_Deficiency_Symptom_Quality, representing the characteristic leaf edge death resulting from K-limited osmoregulation and metabolic function. This class addresses the most diagnostically distinctive K deficiency symptom. Marginal initiation distinguishes from interveinal (Mg, Mn) and uniform (N) patterns. Post-véraison intensification reflects fruit K demand competition with foliage. Progression from bronzing through necrosis provides severity information. Basipetal-to-acropetal and marginal-to-centripetal progression patterns.
K-Limitierungs-Scorch
K-deficiency leaf scorch
K-limitation scorch
Kalium-Blattrandverbrennung
Marginale Blattnekrose bei K-Mangel
leaf edge necrosis
potassium marginal burn
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Mpelasoka BS et al. (2003) A review of potassium nutrition in grapevines with special emphasis on berry accumulation. Australian Journal of Grape and Wine Research 9:154-168 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication
Marginal Leaf Necrosis Quality (K)
Market data for Pinot Noir illustrates the most extreme price polarization in the global wine market — grape prices ranging from AUD 500-800/tonne for commodity Australian Pinot Noir (Central Victoria, Riverina) to the equivalent of AUD 150,000-400,000+/tonne implied by Grand Cru Burgundy wine prices (where a bottle of Romanée-Conti at €5,000-20,000 derives from approximately 1.2-1.5 kg of grapes), wine retail prices spanning from €5 entry-level to €25,000+ per bottle at auction, and demand data showing sustained global growth in premium Pinot Noir consumption driven by the Burgundy prestige effect and the establishment of quality-competitive New World sources (Oregon, New Zealand, Tasmania, Sonoma Coast). Market data for Grüner Veltliner documents a cultivar with a more compressed but evolving price structure — grape prices of approximately €0.40-0.80/kg for basic Qualitätswein grapes to €2.50-5.00+/kg for top Wachau Smaragd sites, wine retail prices from €4-6 for entry-level to €50-150 for single-vineyard Smaragd crus, and demand data showing steady export growth particularly in the US, Scandinavian, and Japanese markets where Grüner Veltliner has achieved recognition as a distinctive food-pairing wine.
Market data is an economic data entity that documents the commercial valuation, pricing structures, and demand patterns for grapes and wines derived from specific cultivars, encompassing grape price data (the prices paid for harvested fruit at the vineyard or winery gate), wine price data (the prices commanded by finished wines at wholesale, retail, and auction), and market demand data (the consumer and trade demand indicators that drive pricing and production decisions), derived from industry price surveys, auction records, retail tracking databases, trade organization reports, and commercial market intelligence, providing the market-facing economic dimension that connects viticultural production to commercial value realization and enables assessment of a cultivar's market positioning within the competitive landscape of the global wine market.
Market data documentation provides the commercial intelligence dimension that transforms viticultural and oenological knowledge into economically actionable information — essential for vineyard investment decisions, production planning, marketing strategy, and the assessment of cultivar economic viability. The market data domain is characterized by significant information asymmetry — large wine companies and industry organizations have access to comprehensive proprietary market intelligence (Nielsen/IRI retail scan data, IWSR consumption data, Liv-ex fine wine trading data), while small producers typically rely on more limited and less timely information sources (regional price surveys, informal market intelligence, auction results for comparable wines). The three sub-dimensions of market data — grape prices, wine prices, and demand indicators — capture different stages of the value chain and different aspects of market dynamics. Grape prices reflect the agricultural commodity value of the harvested fruit, determined by supply-demand dynamics, quality assessments, and contractual relationships between growers and wineries. Wine prices reflect the value added through winemaking, branding, and marketing, incorporating the reputation effects of producer name, appellation prestige, critical scores, and vintage quality. Demand data provides the forward-looking dimension — consumption trends, export market development, consumer preference shifts, and demographic changes that signal future pricing trajectory. The interaction between these three dimensions creates the economic logic of cultivar selection — growers plant cultivars that offer the best expected return on investment considering grape prices, production costs, yield expectations, and market demand trends, while wineries select grape sources that enable wine production at price points matching their target market positioning.
Handelsdaten
Marktinformation
Preisdaten
commercial market data
kommerzielle Daten
market information
market intelligence
pricing data
OIV (2023) State of the World Vine and Wine Sector IWSR (2023) Global Wine Market Analysis Liv-ex (2023) Fine Wine Market Report Anderson K, Nelgen S (2020) Which Winegrape Varieties are Grown Where? University of Adelaide Press
Market Data
Market demand data for Sauvignon Blanc documents the cultivar's transformation from a niche French variety to a global consumer favorite — UK market share expanding from approximately 5% of white wine sales in 2000 to approximately 25% by 2020 (making it the UK's best-selling white grape variety, driven primarily by New Zealand Marlborough), US market showing sustained double-digit annual growth in the premium segment through the 2010s, strong demand penetration in Scandinavian markets through the Systembolaget and Vinmonopolet monopoly retail channels, and emerging market interest in Asia driven by the variety's compatibility with seafood-oriented cuisines — the demand data collectively explaining the production expansion documented in Sauvignon Blanc's acreage trend. Market demand data for Grüner Veltliner documents a demand profile concentrated in specific markets — dominant domestic position in Austria (approximately 35% of all Austrian wine consumption), growing export demand in Germany (Austria's largest export market, accounting for approximately 50% of Austrian wine exports), specialty demand in the US sommelier and restaurant channel (where Grüner Veltliner's food-pairing versatility has driven on-premise adoption), and emerging interest in Japan and the Nordic countries.
Market demand data is a market data entity that documents the consumer and trade demand indicators that drive the commercial market for wines produced from specific grape cultivars, encompassing consumption volume data (the quantity of wine consumed by cultivar or category in specific markets), export-import flow data (the direction and volume of international trade by cultivar or origin), consumer preference indicators (survey data, purchase frequency, brand loyalty metrics, demographic consumption profiles), on-premise versus off-premise channel distribution (the balance between restaurant-hotel-bar consumption and retail home consumption), and emerging market development indicators (growth rates in developing wine markets, new consumer segment penetration), derived from market research organizations, trade statistics, consumer panels, and industry surveys, providing the demand-side intelligence that explains current pricing and informs production planning decisions.
Market demand data provides the demand-side complement to supply-side production statistics — explaining why certain cultivars expand while others contract, why prices rise or fall, and how consumer preferences shape the viticultural landscape through the market signals transmitted through the value chain from consumer to producer. The data sources for market demand analysis operate at different levels of granularity and reliability — national consumption statistics from OIV and Eurostat provide aggregate volume data by country, commercial tracking services (Nielsen/IRI, IWSR) provide retail sales data disaggregated by category, origin, and price tier, consumer panel studies provide purchase frequency and demographic profiling, and on-premise tracking services monitor restaurant and hotel wine list placement and by-the-glass programs. The cultivar-level disaggregation of demand data presents a persistent methodological challenge — consumption statistics are typically reported by origin country or wine category rather than by grape variety, making it difficult to isolate cultivar-specific demand from broader category trends (growth in "New Zealand white wine" conflates Sauvignon Blanc with Pinot Gris, Chardonnay, and other varieties). The demographic dimension of demand data is increasingly important for forward-looking market analysis — generational shifts in wine consumption patterns (documented declines in per capita wine consumption among younger demographics in traditional wine-consuming countries, offset by growing interest in premium and craft wines among the same cohorts), geographic shifts (declining European consumption, growing Asian and African market penetration), and channel shifts (accelerating online wine retail, subscription models, direct-to-consumer sales) all influence cultivar-specific demand trajectories. The interaction between critical acclaim and consumer demand creates feedback loops — high scores from influential critics generate consumer interest and price premiums, which attract investment in production quality, which generates further critical recognition — a virtuous cycle that has driven the market development of several cultivars from obscurity to prominence (New Zealand Sauvignon Blanc, Argentine Malbec, Austrian Grüner Veltliner) within remarkably compressed timeframes.
Handelsflüsse
Konsumentennachfrage
Marktnachfragedaten
Nachfrageindikatoren
Verbrauchsdaten
consumer demand
consumption data
demand indicators
trade flow data
IWSR (2023) Global Wine Market Analysis Wine Intelligence (2023) Global Wine Consumer Trends OIV (2023) State of the World Vine and Wine Sector — consumption data Nielsen/IRI (2023) Wine Category Retail Tracking Data
Market Demand Data
The marketing name Fumé Blanc coined by Robert Mondavi in 1968 repositioned barrel-fermented Sauvignon Blanc as a distinctive premium category in the California market, creating a stylistic designation that was subsequently adopted by dozens of US producers — the name carries no trademark protection and functions as a generic marketing term associating the wine with Pouilly-Fumé prestige while differentiating it from standard Sauvignon Blanc bottlings, illustrating how marketing names can evolve from proprietary creation to generic industry usage.
A trade name adopted or coined to enhance the commercial appeal and consumer recognition of a grapevine cultivar or its wine products, functioning as a market-positioning tool without necessarily carrying formal trademark protection, and typically chosen for its euphonic quality, memorability, or evocative associations.
Marketing_Name captures commercially motivated naming that operates outside the formal trademark system, relying on market convention and consumer recognition rather than legal exclusivity. Marketing names may originate as deliberate branding initiatives (Fumé Blanc) or emerge organically through trade usage (Prosecco, originally a cultivar name, evolved into a geographic marketing name before being formally reclassified as a DOC/DOCG designation in 2009 with the cultivar officially renamed Glera to protect the geographic brand): the boundary between marketing names and established trade usage is often fluid, with successful marketing names sometimes achieving quasi-official status through regulatory recognition or industry standardization.
Marketingname
commercial cultivar name
market-facing designation
promotional name
werblicher Sortenname
Robinson J and Harding J (2015) The Oxford Companion to Wine 4th Ed, Oxford University Press Anderson K and Aryal NR (2013) Which Winegrape Varieties are Grown Where?, University of Adelaide Press Robinson J et al. (2012) Wine Grapes, Allen Lane
Marketing Name
The marketing regional name Cape Riesling was adopted in South Africa to designate plantings of Crouchen Blanc (a minor French cultivar from the Landes) that had been misidentified as Riesling — when molecular analysis revealed the misidentification, the South African industry retained Cape Riesling as a marketing regional name to avoid consumer confusion and preserve the established brand value of the name in the domestic market, eventually phasing it out in favor of the correct designation as international labeling standards tightened, illustrating the tension between marketing convenience and nomenclatural accuracy.
A regional name that was deliberately created or strategically adopted within a region primarily for commercial differentiation and market positioning purposes, typically of more recent origin than traditional regional names and designed to associate the cultivar with a specific geographic brand identity that enhances marketability.
Marketing_Regional_Name captures the commercially motivated creation of place-associated cultivar designations, a practice that has generated considerable nomenclatural controversy. The most contentious examples involve the appropriation of prestigious geographic names for cultivars with no authentic connection to the referenced region: historical instances include Tokay d'Alsace (Pinot Gris marketed with Tokaj associations), Cape Riesling (Crouchen Blanc marketed with Rhine associations), and Clare Riesling (Crouchen Blanc in Australia). International agreements and EU regulations have progressively restricted the creation of new misleading geographic names, but many established marketing regional names persist through grandfather clauses and transitional arrangements.
Marketing-Regionalname
Vermarktungs-Gebietsname
branded regional name
marketing geographic name
promotional regional designation
werbliche Regionalbezeichnung
Robinson J et al. (2012) Wine Grapes, Allen Lane OIV (2013) OIV List of Vine Varieties and Their Synonyms Anderson K and Aryal NR (2013) Which Winegrape Varieties are Grown Where?, University of Adelaide Press
Marketing Regional Name
Merlot in Bordeaux shows mass cap fall during mid-June Year N+1 at BBCH 63-65 over 2-3 days June 9-11, with 55-68% flowers showing cap detachment creating abundant fallen cap accumulation, intensive synchronized shedding visible throughout clusters, under oceanic conditions of 22-27°C temperature, 54-68% relative humidity, stable weather supporting efficient mass detachment enabling 54-64% fruit set, typical of this mid-season cultivar during peak flowering determining season yield
Mass cap fall represents the peak calyptra detachment phase when maximum proportion of flowers complete abscission simultaneously creating characteristic full bloom appearance with abundant cap shedding activity throughout cluster, establishing optimal reproductive conditions and serving as definitive visual marker for peak anthesis. This culminating shedding phase occurs during peak flowering in mid-June in temperate regions Year N+1, typically spanning 2-3 days as 40-70% flowers achieve cap detachment creating intensive synchronized shedding activity, corresponding to BBCH 63-65 stages parallel to peak anthesis. Shedding intensity during mass fall shows maximum levels with 40-70% total flowers showing detached caps creating densely opened cluster appearance, daily addition of 15-25% new cap detachments maintaining high activity, and peak shedding rate representing culmination of flowering progression from initial sporadic activity. Spatial distribution during mass fall shows cap detachment throughout cluster with apical regions showing 70-90% flowers with shed caps from earliest opening, middle regions showing 60-80% detachment representing peak activity zone, and basal regions showing 40-60% shedding as activity extends throughout cluster. Detached cap accumulation reaches maximum during mass fall with abundant fresh caps on cluster surfaces creating characteristic debris layer, continuous cap fall visible throughout day under favorable conditions, and substantial fallen cap accumulation on ground below vines documenting flowering intensity. Flower exposure at mass fall shows majority of reproductive organs revealed throughout cluster creating dense array of opened flowers, freshly exposed stamens and pistils distributed across cluster providing reproductive capacity, and overall appearance of full bloom with abundant functional flowers. Visual appearance at mass fall shows distinctive flowering display with numerous exposed yellow-brown anthers and green-white stigmas creating characteristic coloration, remaining green calyptra caps on 30-60% unopened buds providing contrast, abundant detached caps visible on cluster and ground surfaces, and overall appearance of intense reproductive activity. Synchronous shedding during mass fall creates concentrated cap detachment events with multiple flowers throughout cluster completing abscission simultaneously within hours, coordinated shedding creating visible cap fall waves particularly during favorable morning conditions, and synchronized activity supporting uniform flowering and fruit set potential. Environmental conditions during mass fall critically determine success with optimal 24-30°C temperature supporting maximum abscission activity and reproductive organ function, moderate 50-65% humidity facilitating cap detachment without retention or excessive desiccation, stable favorable weather enabling sustained intensive shedding throughout 2-3 day period. Temperature effects on mass fall show optimal warm conditions promoting rapid synchronized detachment completing mass phase within 2-3 days, cool conditions below 20°C slowing activity potentially extending period to 4-5 days, and warm conditions above 32°C potentially causing stress though generally facilitating mechanical separation. Humidity effects show critical importance with moderate levels supporting efficient mass detachment, high humidity above 70% potentially causing retention problems in susceptible flowers reducing mass fall intensity, and low humidity below 45% potentially causing tissue desiccation though facilitating mechanical cap removal. Rain during mass fall represents catastrophic interference with water preventing cap detachment through elevated humidity and physical impedance, substantially reducing shedding activity potentially limiting flowering to only pre-rain activity, and severely compromising fruit set through disrupted flowering during critical peak period. Wind effects show dual character with light-moderate wind 5-15 km per hour facilitating cap dispersal and enhancing visible fall activity, removing detached caps from cluster surfaces reducing accumulation, though excessive wind above 25 km per hour potentially causing physical damage to newly exposed reproductive structures. Diurnal pattern during mass fall shows characteristic rhythm with maximum shedding activity during morning hours 8am-12pm as temperature rises and humidity declines, reduced activity during afternoon hours as conditions may become less favorable, and daily waves creating repeated intensive shedding periods throughout mass fall phase. Cap retention problems during mass fall affect fruit set potential with flowers showing incomplete or failed abscission unable to expose reproductive organs preventing pollination, retention frequency during peak period strongly affecting ultimate fruit set percentage, and visible retention assessment enabling early fruit set predictions. Abscission zone activity reaches maximum during mass fall with coordinated enzymatic processes across numerous flowers, synchronized cell wall degradation creating separation planes, and hormonal signals possibly coordinating mass shedding though mechanisms incompletely understood. Pollination opportunity during mass fall reaches optimum with widespread flower exposure providing abundant receptive stigmas, maximum pollen availability from numerous dehiscing anthers, and favorable spatial arrangement supporting efficient pollination creating high fruit set potential. Fruit set determination during mass fall shows majority 60-80% final berry number established during this critical 2-3 day period, with successful pollinations during mass fall phase contributing disproportionately to ultimate yield, making mass fall conditions dominant influence on seasonal productivity. Observational characteristics enable definitive mass fall identification through dense cap accumulation on clusters and ground creating obvious visual evidence, numerous simultaneously opened flowers throughout cluster showing characteristic full bloom appearance, and intensive ongoing shedding activity visible during observation periods. Phenological significance includes defining peak flowering period establishing BBCH 63-65 stages, providing definitive visual marker for full bloom easier to assess than detailed reproductive organ examination, and serving as critical reference for subsequent developmental predictions. Management implications during mass fall include maintaining minimal vineyard disturbance with all operations suspended during critical period, intensified weather monitoring with forecasts critically important, and recognition that mass fall outcomes largely determine seasonal yield making this 2-3 day period most important of entire season. The process shows two distinguishable aspects progressing from peak cap shedding period establishing maximum intensity, through synchronous cap fall emphasizing coordinated simultaneous activity. Duration typically spans 2-3 days representing peak flowering period, with warm stable conditions producing compressed 2-day peak while variable conditions may extend to 3-4 days. Cultivar variation in mass fall shows differences in intensity with some varieties showing very high 60-75% simultaneous shedding while others show moderate 45-60% peak, duration affecting flowering compression or extension, and retention susceptibility affecting fruit set outcomes. Regional timing shows mass cap fall during early-mid June in warm climates at approximately 1050-1100 growing degree days base 10°C, mid-June in temperate regions at 950-1000 GDD, and mid-late June in cool areas at 850-900 GDD, marking peak flowering across regions. Viticultural significance includes representing most critical period for yield determination with mass fall outcomes dominating fruit set, providing definitive visual reference for flowering peak enabling precise phenological tracking, serving as practical marker more readily observable than detailed reproductive status, and illustrating extreme importance of favorable conditions during brief 2-3 day window determining season productivity. Research applications include analyzing environmental controls of synchronized shedding, quantifying relationships between mass fall intensity and fruit set outcomes, evaluating cultivar differences in shedding coordination affecting flowering characteristics, developing observation protocols using mass fall as practical marker, and assessing climate change impacts on peak flowering conditions affecting productivity.
Mass_Cap_Fall is second major Cap_Fall phase distinguished by maximum synchronized calyptra detachment during peak flowering. Corresponds to BBCH 63-65 parallel to Peak_Anthesis. Subdivided into Peak_Cap_Shedding_Period and Synchronous_Cap_Fall. Represents most critical 2-3 days determining 60-80% fruit set and yield. Environmental conditions particularly weather critically affect success. Provides definitive visual marker for flowering peak. Easily observable practical reference for phenological tracking and management.
Hauptblüten-Kappenabwurf
Massen-Kappenfall
full bloom calyptra fall
mass calyptra detachment
maximale Kalyptralösung
maximum cap shedding
peak cap fall
synchroner Massenabfall
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 370-380 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Mass Cap Fall
In Cabernet Franc vineyards of Chinon, Loire Valley, a mass leaf fall event occurred on November 28 when overnight temperatures of -5°C followed by morning winds of 25-35 km/h triggered synchronous detachment of approximately 35% of remaining canopy leaves within 36 hours, reducing canopy leaf coverage from 45% to less than 15% and creating concentrated leaf litter accumulation beneath the trellis.
Mass leaf fall in Vitis vinifera represents a period of rapid, highly synchronous leaf detachment affecting a substantial proportion of remaining canopy leaves within a compressed timeframe, typically triggered by specific environmental events that overcome the residual attachment strength of multiple leaves simultaneously across varied canopy positions. This phenomenon occurs most commonly during late November to early December in Northern Hemisphere viticulture, when significant frost events, strong wind episodes, or the combination of both create mechanical and physiological conditions that trigger widespread simultaneous leaf separation. Unlike the gradual progression characteristic of earlier leaf fall phases, mass leaf fall events can result in 20-40% of remaining canopy leaf area detaching within 24-72 hours, dramatically transforming vine appearance from partially foliated to nearly bare. The triggering conditions for mass leaf fall typically involve nighttime temperatures dropping to -3°C to -6°C that cause ice crystal formation within leaf tissues, disrupting cell membrane integrity and increasing leaf weight through ice accumulation while simultaneously making tissues brittle and more susceptible to mechanical separation. Wind events coinciding with or following frost exposure are particularly effective triggers, as the combination of weakened attachment points and mechanical stress creates conditions favoring synchronous detachment across multiple canopy positions regardless of their individual senescence advancement status. The phenomenon exhibits threshold behavior, with gradual accumulation of abscission zone maturation creating increasing proportions of leaves approaching detachment competence until environmental triggers release them simultaneously. Cultivar susceptibility to mass leaf fall events varies with leaf anatomical characteristics, with thin-leaved varieties showing greater frost sensitivity and more pronounced mass fall responses, while thick-leaved cultivars may retain damaged leaves longer due to more robust petiole construction. From a viticultural management perspective, mass leaf fall events provide convenient timing windows for winter pruning commencement, as the rapid canopy clearing exposes cane architecture and allows pruner assessment of wood maturity and bud positioning. The synchronous nature of mass leaf fall also creates concentrated leaf litter accumulation that influences vineyard floor management decisions regarding incorporation timing and disease pressure management for following seasons. Physiologically, leaves lost during mass leaf fall events may retain higher nutrient concentrations than those completing gradual progressive detachment, as the rapid separation can interrupt ongoing remobilization processes in less-advanced leaves.
Mass_Leaf_Fall represents a distinct event-driven phase within Leaf_Detachment characterized by rapid synchronous defoliation rather than gradual progression. Distinguished from Progressive_Leaf_Fall by its compressed timeframe and synchronous character, and from Complete_Leaf_Detachment which addresses final residual leaf removal. Triggering conditions typically involve frost events below -3°C combined with wind exposure. Can occur as single or multiple events during late autumn. Duration of individual mass fall events typically 24-72 hours. May account for 20-40% of total canopy defoliation in single episodes.
Massenblattfall
bulk leaf detachment
gleichzeitiger Laubfall
rapid mass defoliation
schnelle Massenentblätterung
simultaneous leaf shedding
synchroner Blattfall
synchronous leaf fall
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Mills LJ, Ferguson JC, Keller M (2006) Cold-hardiness evaluation of grapevine buds and cane tissues. American Journal of Enology and Viticulture 57: 194-200
Mass Leaf Fall
Vine balance assessment in commercial Merlot vineyards utilizes the Ravaz Index, calculated as the ratio of yield mass (kg per vine) to dormant pruning mass (kg per vine), with optimal values of 5-10 indicating balanced vegetative and reproductive growth lower values (below 4) suggest under-cropping relative to vine capacity and potential for excessive vigor, while higher values (above 12) indicate over-cropping that risks incomplete fruit ripening and vine exhaustion over successive seasons.
Mass_Quality represents the category of morphological qualities that quantify the amount of matter contained in grapevine organs regardless of their spatial extent or arrangement. Mass provides a fundamental size measurement that directly relates to the quantity of biological material and stored compounds, making it essential for yield determination, growth analysis, and compositional assessment. This category distinguishes two primary measurement states: Fresh_Mass_Quality capturing the total mass including water content as the organ exists at measurement time, and Dry_Mass_Quality representing the water-free residual mass that reflects accumulated structural and storage compounds. Mass measurements are universal in grapevine research and viticulture because they provide the basis for expressing concentrations, calculating yields, and tracking biomass accumulation. Berry mass directly determines yield when multiplied by berry number cluster mass enables yield estimation from cluster counts pruning mass indicates vine vigor and carbohydrate reserve status. Mass relates to volume through density, with characteristic density ranges for different organs and developmental stages. The relative simplicity and precision of mass measurement using analytical balances makes it the preferred size metric when destructive sampling is acceptable. Mass per unit area or length provides standardized indices (e.g., specific leaf area, cane mass per linear meter) that account for size variation and enable comparison across vines, vineyards, and studies.
Mass_Quality is positioned parallel to Linear_Dimension_Quality, Area_Quality, and Volume_Quality under Size_Quality. The term 'weight' is commonly used interchangeably with 'mass' in viticultural practice, though technically weight is the gravitational force on mass and varies with location. Fresh mass and dry mass measurements require standardized protocols specifying sampling time, handling procedures, and drying conditions for reproducibility. Mass measurements provide the denominator for concentration expressions (mg/g, μg/kg) that characterize chemical composition. PATO_0000125 (mass) provides the upper-ontology reference.
Gewicht
Gewichtsqualität
Massequalität
Materiemenge
amount of matter
gravimetric quality
mass measurement quality
weight
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 153-189 Ravaz L (1911) L'effeuillage de la vigne. Annales de l'École Nationale d'Agriculture de Montpellier 11: 216-244
Mass Quality
A Burgundy domaine maintaining vines propagated by successive generations of sélection massale from their oldest parcels bears a Mass_Selection_Clone_Role, the population representing accumulated selection of superior performers over decades without formal genotype isolation—prized for the diversity and traditional character this approach preserves. A Spanish bodega perpetuating pre-phylloxera Tempranillo through mass selection from their centenary vineyard serves in a Mass_Selection_Clone_Role, valuing the genetic diversity and site adaptation developed over 100+ years of selection. A Loire Valley producer sourcing Chenin Blanc material from traditional mass-selected vineyards bears a Mass_Selection_Clone_Role, deliberately avoiding the uniformity of registered clones to maintain wine complexity attributed to genotypic diversity.
A selection status clone role that inheres in a grapevine clone or population derived through mass selection (sélection massale), where propagation material is collected from multiple superior vines within a vineyard or region without isolating and evaluating individual genotypes. Mass selection identifies visually superior performers—vines with desired yield, vigor, fruit quality, or health characteristics—then collects cuttings from this selected population, perpetuating a mixture of genotypes sharing favorable phenotypic characteristics rather than a single uniform genotype. Mass selection was the universal pre-scientific approach to grapevine improvement, practiced over centuries by vignerons selecting propagation material from their best vines, and remains valuable for maintaining genetic diversity within cultivars, capturing locally adapted populations, and preserving traditional vineyard character. Mass-selected populations exhibit phenotypic variation (unlike uniform clonal selections), potentially providing production stability through genotypic buffering and preserving the accumulated diversity of heritage vineyards. The approach has experienced renewed interest as producers seek alternatives to the genetic uniformity of registered clonal material.
Mass_Selection_Clone_Role captures the traditional approach that preceded formal clonal selection and remains valued by producers seeking genetic diversity and traditional vineyard character. The "clone" designation is somewhat paradoxical for mass selections, which by definition comprise multiple genotypes rather than a uniform clone, but the term is applied in the context of propagation material classification. Mass selection's advantages include preserving genetic diversity (buffering against disease, environmental stress, and market change), maintaining locally adapted populations, and preserving traditional vineyard character developed through centuries of informal selection. Disadvantages include phenotypic variability (complicating vineyard management), potential perpetuation of virus-infected material (without individual plant testing), and absence of documented performance characteristics. Contemporary mass selection programs may combine traditional field selection with individual plant virus testing, creating "sanitized mass selections" that preserve diversity while ensuring health status.
Massenauslese
Massenselektion
Populationsauslese
mass selection
population selection
sélection massale
Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 Huglin P, Schneider C (1998) Biologie et Écologie de la Vigne, Lavoisier This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Pelsy F (2010) Molecular diversity within grapevine varieties, Heredity 104: 331-340
Mass Selection Clone Role
Chardonnay in Champagne shows massive vacuole formation during late July-early August Year N+1 at BBCH 75-77, with vacuoles expanding from 62% to 88% cell volume, single central vacuole 105 micrometers diameter occupying cell interior, tonoplast area 2200 μm² per cell, vacuole storing malate 72 mM, tartrate 48 mM, K+ 124 mM, pH 3.2, cytoplasm as 3.2 micrometer peripheral layer, under cool maritime conditions of 20-25°C temperature, typical of this early-bursting cultivar developing massive vacuolar system
Massive vacuole formation represents the dominant compartment establishment phase during rapid cell expansion when central vacuole expands to occupy 80-90% final cell volume, single large vacuole per cell becomes predominant organelle, vacuole stores accumulated solutes and water creating massive osmotic and storage compartment, and cytoplasm organizes as thin 2-5 micrometer peripheral layer, completing vacuolar development during intensive enlargement weeks 5-8 post-fertilization. This vacuolar dominance establishment phase occurs during late summer in late July-early August in temperate regions Year N+1, proceeding over weeks 5-8 as vacuole reaches maximum development, corresponding to BBCH 75-77 berry touch to beginning bunch closure stages. Massive formation mechanism involves progressive vacuole expansion from 50-70% volume at rapid expansion onset to 80-90% at completion, sustained water influx into vacuole driving enlargement, tonoplast membrane synthesis accommodating massive expansion, solute accumulation maintaining osmotic gradient, and eventual vacuole dominance with single central compartment occupying cell interior. Vacuole volume fraction progression shows typical increase from 50-70% initial rapid expansion to 75-85% mid-phase to 80-90% completion, progressive volume occupation, final vacuole as overwhelmingly dominant compartment, and high fraction characteristic of fully expanded berry cells. Vacuole size in fully expanded cells shows vacuole diameter 60-180 micrometers depending on cell size, massive compartment volume 0.1-3 nanoliters, single vacuole per cell typical though occasionally 2-3 vacuoles, and large size as defining characteristic. Central vacuole position shows vacuole occupying cell center, cytoplasm peripheral surrounding vacuole, nucleus and organelles in cytoplasmic layer, and spatial organization characteristic of expanded cells. Cytoplasm organization shows thin peripheral layer 2-5 micrometers thickness, cytoplasm containing nucleus, mitochondria, plastids, ER, and other organelles, maintained metabolic functions in reduced volume, and efficient organization supporting cell function. Tonoplast membrane area shows massive membrane surface accommodating large vacuole, typical 1000-5000 μm² tonoplast area per cell, membrane synthesis during expansion producing required area, and large membrane area enabling extensive transport capacity. Tonoplast protein composition shows H+-ATPase proton pumps maintaining pH gradient, numerous transporters for organic acids, sugars, ions, aquaporins facilitating water movement, and specialized protein complement supporting vacuole function. Vacuole storage capacity shows massive volume 80-90% cell enabling extensive solute storage, vacuole storing organic acids 1.2-1.8%, potassium 80-150 mM, beginning sugar accumulation 5-8%, water comprising 85-90% vacuole mass, and vacuole as major storage depot. Organic acid storage shows malate and tartrate accumulated to peak concentrations, acids stored in vacuole maintaining high cytoplasmic pH, acid storage creating acidic vacuole pH 3.0-3.5, and acids as major stored solutes and contributors to acidity. Potassium storage shows K+ accumulated in vacuole, concentration 80-150 mM in vacuole versus 100-150 mM in cytoplasm, potassium as major cation, and K+ storage supporting osmotic potential and electrical balance. Water storage shows water as primary vacuole content 85-90% mass, massive water storage in berry cells, water mobilizable during ripening and later development, and water as dominant stored compound. Vacuole osmotic function shows accumulated solutes creating osmotic potential -1.0 to -1.5 MPa, strong negative potential driving water uptake, vacuole as osmotic engine maintaining turgor, and osmotic function enabling sustained expansion. Vacuole pH regulation shows proton pump acidification maintaining pH 3.0-3.5, acidic pH from organic acid accumulation and active pumping, pH affecting enzyme activities and transport, and pH regulation as key vacuole function. Vacuole metabolic role shows compartmentalization segregating acids from cytoplasmic metabolism, maintaining cytoplasmic pH 7.0-7.5 while vacuole acidic, metabolic efficiency from compartmentation, and vacuole as regulatory compartment. Cytoplasm-vacuole relationships shows dynamic exchange of solutes and water, tonoplast transporters mediating exchange, coordinated regulation maintaining cellular homeostasis, and integrated function despite spatial separation. Vacuole volume as cell volume determinant shows vacuole expansion accounting for 70-85% total cell volume increase during rapid expansion, vacuole as major volume contributor, and vacuole size determining cell size. Massive vacuole formation timing shows progressive development during weeks 5-8, completion by end of rapid expansion, and fully developed vacuole persisting through subsequent development. Fully expanded cell characteristics shows cell diameter 80-200 micrometers depending on cultivar and position, central vacuole 80-90% volume, thin peripheral cytoplasm, and mature expanded morphology. Environmental effects on massive vacuole formation include water availability affecting vacuole expansion capacity with adequate water essential for maximum development, temperature affecting development rate, and favorable conditions supporting complete vacuole development. Water effects show water limitation restricting vacuole expansion limiting cell size, adequate water enabling full vacuole development, and water as critical determinant of vacuole and cell size. Massive vacuole assessment methods include microscopy showing vacuole size and position, vacuole isolation and volume measurement, tonoplast transport studies, and compositional analysis of vacuole contents. The process represents second aspect of rapid cell expansion establishing dominant compartment, following maximum expansion rate and preceding peak berry growth rate. Duration weeks 5-8 representing progressive vacuole development to maximum. Regional timing shows massive vacuole formation during late July-early August across climates during rapid enlargement. Viticultural significance includes representing establishment of major storage compartment, creating osmotic engine sustaining expansion, providing capacity for subsequent sugar accumulation during ripening, and determining final cell architecture. Research applications include analyzing vacuole biogenesis and regulation, investigating tonoplast transport and function, evaluating vacuole contribution to cell size, quantifying storage capacity and composition, and assessing vacuole as determinant of cell and berry size.
Massive_Vacuole_Formation is second aspect of Rapid_Cell_Expansion, distinguished by dominant compartment establishment. Represents expansion from 50-70% to 80-90% cell volume. Single central vacuole 60-180 micrometers. Tonoplast 1000-5000 μm² area. Stores acids, K+, water (85-90% mass). Osmotic potential -1.0 to -1.5 MPa. pH 3.0-3.5. Cytoplasm 2-5 micrometer peripheral layer. Duration weeks 5-8. Follows Maximum_Expansion_Rate. Precedes Peak_Berry_Growth_Rate. Accounts for 70-85% cell volume increase.
central vacuole development
dominant vacuole establishment
dominante Vakuolen-Etablierung
große Vakuolen-Formation
large vacuole formation
massive Vakuolen-Bildung
vacuolar compartment dominance
vakuoläre Kompartment-Dominanz
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 447-450
Massive Vacuole Formation
Riesling serves in a Maternal_Parent_Role when emasculated flowers receive pollen from disease-resistant donors, with hybrid seeds developing on the Riesling vine and inheriting Riesling cytoplasm alongside nuclear contributions from both parents. A V. vinifera variety with reliable fruit set and good seed viability bears a Maternal_Parent_Role in large-scale crossing programs, its practical advantages as a seed parent outweighing considerations that might favor its use as pollen source. Chardonnay functioning as maternal parent in quality × resistance crosses contributes cytoplasmic genetics and 50% of nuclear DNA to progeny evaluated for combining quality wine character with disease tolerance.
A parent role that inheres in a grapevine genotype by virtue of its function as the female parent in a controlled cross, contributing the egg cell (female gamete) and providing the ovule tissue that develops into the seed following fertilization. The maternal parent contributes both nuclear genetic material (one haploid chromosome set combining with paternal contribution) and cytoplasmic genetic material (mitochondrial and chloroplast genomes inherited maternally in most angiosperms). In grapevine breeding practice, the maternal parent is the vine on which crossed fruit develops—flowers are emasculated (anthers removed before pollen shed), then pollinated with pollen from the paternal parent, with resulting seeds germinated to produce hybrid progeny. The maternal parent's fruit-bearing capacity, seed viability, and reliability of fruit set influence breeding program logistics some varieties are preferred as maternal parents due to excellent seed production while others are primarily used as pollen donors. Cytoplasmic inheritance through the maternal parent may influence traits such as male sterility, though nuclear genes dominate most agronomically important characteristics.
Maternal_Parent_Role is subclassed by Seed_Parent_Role, which emphasizes the seed-development function specifically. The distinction between maternal and paternal roles in grapevine breeding reflects both the biological reality of differential gametic contribution (egg cell versus pollen) and practical breeding logistics (which variety more reliably produces seeds). Cytoplasmic inheritance through the maternal line may have subtle phenotypic effects, though most breeding selection focuses on nuclear-encoded traits where both parents contribute equally. The maternal parent's genotype determines seed tissue characteristics (seed coat, endosperm in early development) that may influence germination and early seedling vigor independently of the embryo's hybrid genotype. Choice of maternal versus paternal assignment for specific genotypes often reflects practical experience with seed production reliability rather than directed genetic strategy.
Mutterpflanze
female parent
maternal parent
mother plant
mütterlicher Elter
samentragende Elternpflanze
seed-bearing parent
weiblicher Elter
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp.: 78-112 Antcliff AJ (1988) Grape breeding in Australia, in Proceedings 2nd International Symposium for Cool Climate Viticulture
Maternal Parent Role
Syrah grafted on SO4 rootstock showing maturation zone beginning 8.2 mm from root tip on second-order lateral root, young maturation zone 8-15 mm from apex showing initial differentiation with root hairs just emerging 9 mm from apex, endodermis developing casparian strips, provascular tissue differentiating into xylem and phloem, middle maturation zone 15-40 mm showing advanced differentiation with abundant root hairs 150 per cm, mature endodermis with complete casparian strips, functional xylem vessels with lignified walls, mature phloem with functional sieve elements, cortex cells containing starch, maturation zone providing functional absorption, transport, and storage capacity
Maturation zone is the root region basal to elongation zone where cells complete differentiation into functional tissues, typically beginning 5-15 mm from root apex and extending indefinitely basipetally, characterized by cells ceasing elongation and undergoing tissue-specific differentiation producing functional epidermis with root hairs, cortex, endodermis with casparian strips, pericycle, and vascular tissues including xylem and phloem, showing progressive maturation with youngest maturation zone closest to elongation zone showing initial differentiation and older regions showing fully mature tissues, and serving as region where undifferentiated elongated cells transform into functional specialized tissues enabling root to perform absorption, transport, storage, and other physiological functions, with maturation zone demonstrating root hair emergence typically beginning 3-10 mm from apex creating absorptive surface, endodermis developing casparian strips regulating radial transport, xylem vessels lignifying enabling water transport, phloem sieve elements maturing for food conduction, and cortex cells accumulating starch or other storage compounds, and representing transition from growth-dedicated tip tissues to functional mature root capable of performing essential root system functions. Maturation zone position and extent includes= beginning at elongation zone terminus (maturation zone starting 5-15 mm from root apex where cell elongation essentially ceases), extending indefinitely basipetally (maturation zone comprising all older root regions from elongation cessation to eventual secondary growth onset), showing gradual maturation gradient (youngest maturation zone showing initial differentiation, middle zone showing advanced differentiation, older zone showing complete maturation), demonstrating variable starting position (maturation zone beginning closer to apex in slow-growing roots, farther in fast-growing roots), and revealing order-dependent patterns (fine roots showing maturation zone closer to apex than large structural roots). Maturation zone cell differentiation includes= showing tissue-specific development (cells differentiating according to position producing epidermis, cortex, endodermis, pericycle, xylem, or phloem), exhibiting programmed changes (differentiation following genetic program with sequential gene expression and structural modifications), demonstrating cell type characteristics (each tissue type developing distinctive features appropriate for function), revealing irreversible specialization (differentiation creating committed cell types unable to return to meristematic state), and displaying coordinated patterning (tissues differentiating in organized pattern creating functional root structure). Maturation zone root hair development includes= showing trichoblast differentiation (specific epidermal cells differentiating into root hair-forming cells), exhibiting hair emergence (root hairs beginning to grow 3-10 mm from apex in young maturation zone), demonstrating hair zone (region showing active root hairs typically 5-50 mm from apex depending on root type and lifespan), revealing hair density (potentially 50-300 hairs per centimeter root length depending on species and conditions), and displaying absorption function (root hairs greatly increasing surface area providing majority of water and nutrient uptake). Maturation zone endodermis development includes= showing casparian strip formation (endodermis cells depositing suberin and lignin in radial and transverse walls creating waterproof barrier), exhibiting radial transport regulation (casparian strips forcing solutes through endodermis symplast enabling selective uptake), demonstrating progressive thickening (older endodermis potentially developing U-shaped or O-shaped secondary wall thickening), revealing passage cells (some endodermis cells remaining thin-walled in older roots maintaining transport capacity), and displaying barrier function (endodermis controlling ion movement from cortex to stele). Maturation zone xylem development includes= showing vessel differentiation (provascular cells elongating, thickening walls, and depositing lignin creating vessels), exhibiting cell death (vessel elements undergoing programmed cell death removing cell contents creating hollow tubes), demonstrating lignification (secondary wall thickening and lignin deposition providing strength and waterproofing), revealing vessel maturation (vessels becoming functional for water transport as lignification completes), and displaying transport capacity (mature xylem vessels conducting water from maturation zone to shoot system). Maturation zone phloem development includes= showing sieve element differentiation (provascular cells developing into enucleate sieve elements with perforated sieve plates), exhibiting companion cell formation (specialized parenchyma cells maintaining metabolic support for sieve elements), demonstrating sieve plate development (end walls forming pores creating cytoplasmic continuity between sieve elements), revealing phloem maturation (phloem becoming functional for photosynthate transport as sieve elements mature), and displaying conduction function (mature phloem transporting sugars and other compounds from shoot to roots). Maturation zone cortex function includes= showing storage capacity (cortex parenchyma cells accumulating starch and other reserves), exhibiting radial transport (cortex cells facilitating water and solute movement from epidermis to endodermis), demonstrating metabolic activity (cortex cells respiring, synthesizing, and processing compounds), revealing mycorrhizal colonization (cortex providing site for arbuscular mycorrhizal fungal colonization), and displaying structural support (cortex contributing to root strength and flexibility). Maturation zone in root function includes= providing absorption capacity (root hairs in maturation zone performing majority of water and nutrient uptake), enabling transport function (mature xylem and phloem conducting water, minerals, and photosynthates), demonstrating storage function (cortex and pericycle storing carbohydrates and other compounds), showing selective permeability (endodermis regulating ion uptake and preventing backflow), and revealing mycorrhizal interface (maturation zone cortex hosting fungal symbionts).
Maturation_Zone emphasizes cell differentiation and functional development and is distinguished from elongation zone (cell expansion) by focus on tissue specialization. Includes Cell_Differentiation_Maturation_Zone and Functional_Maturation_Zone. Critical for understanding tissue differentiation, root hair development, endodermis formation, xylem-phloem maturation, and the region where cells complete specialization becoming functional tissues enabling root physiological functions.
Differentiation zone
Differenzierungszone
Functional root region
Funktionelle Wurzelregion
Reifungszone
Root hair zone
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Dolan, L., et al. (1993). Cellular organisation of the Arabidopsis thaliana root. Development, 119(1), 71-84.
Maturation Zone
Cabernet Franc at BBCH 59-60 (flower buds swollen, first calyptras beginning to lift) exhibits mature anthers 0.7-0.8 mm length with bright yellow coloration visible through translucent calyptras or exposed on early-opening flowers, containing fully mature pollen at 74-82% viability (assessed by in vitro germination in 10% sucrose medium), with progression to dehiscing anther state occurring 3-8 hours after calyptra shedding under favorable morning conditions (temperatures rising from 18°C to 24°C, humidity declining from 72% to 54%). Sauvignon Blanc mature anthers at late BBCH 59 show golden-yellow color, swollen appearance, and contain 4,200-5,800 pollen grains per anther at 68-76% viability, ready for imminent dehiscence when flowers open. Riesling experiencing unseasonably cool bloom period (11-13°C) shows mature anthers with bright yellow color and swollen morphology indicating pollen maturation completion, but dehiscence delayed 18-36 hours beyond calyptra shedding due to insufficient temperature for dehiscence mechanics, with pollen viability declining from initial 70-78% to 52-64% during the extended mature anther period before eventual dehiscence when temperatures briefly increase to 16-18°C.
A mature anther is an anther that has completed pollen maturation with pollen grains achieving maximum viability and the anther wall tissues fully prepared for dehiscence, but with dehiscence not yet initiated or in very early stages, representing the pre-dehiscence state (BBCH 59-60) when anthers contain maximum pollen complement ready for imminent release and when environmental conditions critically determine the timing of transition to active pollen dispersal. Mature anthers are characterized by= (1) anther lobes swollen to maximum dimensions (typically 0.6-1 mm length, 0.3-0.5 mm width in hermaphrodite flowers) due to internal pollen mass occupying pollen sac volume (2) bright yellow or golden-yellow coloration visible through the anther wall epidermis, reflecting the abundant carotenoid-containing pollen grains packed within the four pollen sacs (3) anther wall tissues showing full maturation with endothecium cells displaying characteristic secondary wall thickenings creating the hygroscopic apparatus for dehiscence, though dehiscence slits not yet formed or just beginning to form (4) pollen grains within pollen sacs achieving maximum viability (typically 70-85% viable in hermaphrodite flowers, assessed by in vitro germination or vital staining), maximum starch and lipid reserves for germination and pollen tube growth, and characteristic mature Vitis pollen morphology with complete exine ornamentation (5) stomium (dehiscence zone) showing incipient cell breakdown or cell wall weakening in preparation for splitting but longitudinal slit not yet opened. The mature anther state occurs during late BBCH 59 through early BBCH 60, representing the critical transition period between pre-anthesis and active anthesis, typically persisting for 6-24 hours depending on environmental conditions before progressing to dehiscing anther state. Mature anthers are still enclosed by the intact or separating calyptra in many flowers, with calyptra shedding and mature anther exposure often occurring nearly simultaneously as the coordinated events defining anthesis onset. The physiological state of mature anthers includes= pollen grains in a semi-desiccated state optimized for dispersal and subsequent rehydration on stigmas, with moisture content typically 15-25% of fresh weight anther wall tissues undergoing final preparations for dehiscence including programmed stomium cell death and endothecium wall thickening completion metabolic activity declining as developmental processes complete and the anther transitions from biosynthetic to dispersal phase. Environmental conditions during the mature anther stage critically determine dehiscence timing and efficiency= favorable conditions (rising temperatures 18-28°C, declining humidity from 70-80% to 50-60%, calm to light air movement) promote rapid progression to dehiscence within 2-6 hours of calyptra shedding unfavorable conditions (cool temperatures 85%, rain) delay dehiscence for 12-48+ hours or prevent normal dehiscence entirely, with prolonged mature anther stage creating extended vulnerability to pollen deterioration or pathogen infection. Mature anthers represent the state of maximum reproductive potential, with pollen at peak viability and quantity, making this stage the optimal target for controlled crossing operations in breeding programs where anthers are collected just before natural dehiscence for pollen extraction and controlled application to selected pistils. Mature anther color (bright yellow) serves as a critical visual indicator for viticulturists and researchers, signaling imminent bloom and allowing prediction that dehiscence and active pollination will occur within hours if environmental conditions are favorable, or providing diagnosis of bloom delay if mature yellow anthers persist without dehiscing under unfavorable conditions. The mature anther stage is also the period of maximum vulnerability to environmental stresses that can damage pollen before dispersal= high temperatures >35°C can cause pollen desiccation and viability loss even within protected mature anthers pathogen infection by Botrytis or other fungi can colonize mature anthers before dehiscence if humidity is excessive, destroying pollen and impairing subsequent reproductive function.
Mature_Anther represents the pre-dehiscence state with completed pollen maturation and is distinguished from Dehiscing_Anther (active pollen release) by absence of opened longitudinal slits and from immature anthers (earlier developmental stages) by bright yellow color and maximum pollen viability. The mature anther state represents peak reproductive potential but is transient and highly sensitive to environmental conditions determining dehiscence timing. Mature anther characteristics (bright yellow color, swollen appearance, position relative to calyptra status) serve as critical phenological indicators for predicting imminent bloom and for timing management operations or controlled crossing activities. The mature anther stage is the optimal period for anther collection in breeding programs, as pollen is at maximum viability and quantity but not yet dispersed.
fertiger Pollensackträger
pollen-ready anther
pre-dehiscence anther
reife Anthere
reifer Staubbeutel
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 101-104 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 95-97 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 189-192.
Mature Anther
In Muscat Blanc à Petits Grains berries in Rutherglen, Australia at BBCH 89, mature aroma expression is characterized by total monoterpenes of 9.8 mg/L with linalool at 3.2 mg/L and geraniol at 2.1 mg/L providing intense floral-grape impact, norisoprenoid precursors of 165 µg/L contributing complexity, methoxypyrazine below 2 ng/L detection threshold, and C6 aldehydes at 18 µg/L indicating complete green character elimination, with trained panel aromatic intensity rating of 8.9 and overall quality of 8.6 on 10-point scales confirming achievement of optimal aromatic maturity.
Mature aroma expression describes the characteristic olfactory profile achieved in fully ripe Vitis vinifera berries at the conclusion of the ripening process, wherein volatile compound composition has stabilized at harvest-appropriate levels and the overall scent impression reflects the balanced integration of varietal character, ripeness indicators, and site-specific nuances that define quality wine grape aromatic maturity and provide the olfactory foundation for premium wine production. This state encompasses specific compositional markers including monoterpene concentrations achieving variety-appropriate levels with aromatic cultivars showing 3-15 mg/L total terpenoids providing intense floral-fruity impact, norisoprenoid levels of 50-200 µg/L contributing complexity and background depth, minimal residual methoxypyrazine below variety-specific thresholds ensuring absence of excessive green character, and reduced C6 aldehyde content below 30 µg/L indicating completion of green-to-ripe aromatic transition. The sensory characteristics of mature aroma expression include predominance of ripe-fruit and floral aromatics over green-herbaceous notes, integration of multiple scent elements into harmonious overall impression without dominant off-characters, appropriate intensity matching varietal expectations with aromatic varieties showing pronounced but balanced scent profiles, and absence of overripe or degraded character indicating optimal harvest timing capture. The biochemical indicators of mature aroma expression include stabilization of terpene biosynthesis rates with daily accumulation declining to maintenance levels below 0.05 mg/L per day, completion of rapid-phase carotenoid degradation with norisoprenoid generation approaching equilibrium, and establishment of final free-to-bound terpenoid ratios of 0.08-0.18 representing mature partitioning between immediate aromatic expression and precursor potential. Temperature history during the final ripening phase influences mature aroma expression through effects on late-stage biosynthesis and compound stability, with cool conditions of 15-20°C during the final weeks preserving accumulated volatile compounds while warm conditions may accelerate degradation compromising optimal aromatic quality. Assessment of mature aroma expression requires integration of analytical quantification with sensory evaluation, as compositional measurements alone may not fully capture the perceptual qualities underlying aromatic maturity judgments that guide harvest timing decisions in quality-focused viticulture operations targeting optimal wine aromatic potential.
Mature_Aroma_Expression is subclass of Aroma_Profile_Maturation representing achieved quality state at ripening completion. Distinguishes from Optimal_Aroma_Development which emphasizes process optimization. Sibling to Cultivar_Typical_Aroma and Optimal_Aroma_Development. Characterized by specific compositional markers and sensory attributes. Integrates intensity, complexity, and balance. Harvest maturity indicator for aromatic quality.
erntereifes Aromaprofil
harvest-ready aromatic expression
mature olfactory profile
optimal scent state
optimaler Duftzustand
reife Aroma-Expression
reifer Duft-Charakter
ripe aroma character
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 305-315 Marais J (1983) Terpenes in the aroma of grapes and wines. South African Journal of Enology and Viticulture 4: 49-58 Robinson AL et al (2014) Origins of grape and wine aroma. American Journal of Enology and Viticulture 65: 1-30
Mature Aroma Expression
Cabernet Sauvignon mature berries at harvest readiness (BBCH 89, week 18 post-fruit set, 110-125 days post-anthesis) achieve 13.5-15 mm diameter, 2.4-2.9 g fresh weight, cells 150-165 μm diameter, 24-26% sugars, 5.5-6.8 g/L titratable acidity, pH 3.65-3.85, 2.8-3.8 mg/g skin anthocyanins, firmness 1.2-1.8 N, representing optimal maturity for premium red wine production with balanced composition supporting complexity and aging potential. Chardonnay mature berries for white wine production achieve 12.5-14 mm diameter, 2.1-2.5 g, 140-155 μm cells, 21-24% sugars, 6.5-8.5 g/L acidity, pH 3.2-3.5, creating high-quality white wine base with good acidity retention and moderate sugar levels. Thompson Seedless mature table grapes achieve large size (19-23 mm diameter, 5-7 g per berry) suitable for fresh market, 18-21% sugars providing sweet eating quality, 4.5-6.5 g/L acidity for flavor balance, crisp texture (firmness 1.8-2.5 N higher than most wine cultivars) for shipping durability, demonstrating table grape maturity criteria emphasizing size, texture, and moderate sugar versus wine grape criteria emphasizing composition for fermentation. Pinot Noir mature berries examined across four harvest dates (weeks 16, 17, 18, 19) show progression from under-ripe (week 16= 22% sugars, 8.2 g/L acidity, 1.8 mg/g anthocyanins, unbalanced composition) through optimal maturity (week 17-18= 24-25% sugars, 6.2-7 g/L acidity, 2.6-3.2 mg/g anthocyanins, balanced for high-quality Pinot Noir) to beginning overripeness (week 19= 26.5% sugars, 5.2 g/L acidity, 2.8 mg/g anthocyanins, acid declining excessively), demonstrating narrow optimal maturity window for this cultivar requiring precise harvest timing.
A mature berry is a berry that has completed or substantially completed the developmental program achieving harvest readiness or post-harvest state (BBCH 89+), characterized by completion of cell expansion, achievement of target sugar levels, acid decline to mature levels, full color development, soft texture, and compositional balance appropriate for intended use, representing berries suitable for harvest or already harvested. Mature berries are characterized by= (1) berry dimensions at or near cultivar-specific and environmentally-determined maximum, typically 11-25 mm diameter and 1.5-7 g fresh weight depending on cultivar and growing conditions (2) cells at maximum expansion with cell diameter 120-180 μm, having achieved 300-500% volume increase from veraison size (60-90 μm) (3) growth essentially ceased with diameter and weight changes <0.02 mm per week and <0.01 g per week or approaching zero (4) sugars at mature levels typically 18-28% (expressed as °Brix or % w/w) depending on cultivar, vintage, and harvest criteria, representing glucose and fructose accumulation suitable for fermentation (wine grapes) or consumption (table grapes) (5) titratable acidity at mature levels typically 4-10 g/L (expressed as tartaric acid equivalents) with pH 3.2-4.2, representing acid decline from peak levels (20-25 g/L) through malic acid respiration and dilution (6) full color development with anthocyanins at mature levels (1.5-5+ mg/g skin in red/black cultivars) creating characteristic varietal coloration, or golden-yellow color in white cultivars (7) soft texture with firmness 0.8-2.5 N representing 70-85% reduction from green berry firmness (4-6 N) through cell wall modifications and turgor changes. Mature berries show compositional characteristics determining quality and use suitability= sugar=acid ratio (°Brix=titratable acidity) typically 20-45 for wine grapes depending on style, higher for table grapes phenolic compounds including anthocyanins, tannins, and flavonols at levels determining color, structure, and antioxidant properties volatile aromatic compounds creating varietal aroma profiles non-volatile flavor precursors that develop during fermentation or consumption. Mature berries represent multiple physiological states= optimally ripe berries achieving ideal compositional balance for harvest overripe berries having exceeded optimal harvest timing with composition changes potentially degrading quality harvested berries removed from vine undergoing post-harvest changes post-veraison berries having completed color change and entering final maturation. Mature berries undergo post-ripening changes if left on vine= continued sugar accumulation through concentration as respiration and transpiration continue continued acid decline potentially reaching excessively low levels anthocyanin stability or degradation depending on temperature and light potential dehydration creating raisining increased susceptibility to rot organisms (Botrytis, other fungi). Mature berries for wine production show cultivar-specific and style-specific target compositions= early-harvest styles (sparkling wine, light white wines) targeting 18-21% sugars, 7-10 g/L acidity standard-harvest red wines targeting 23-25% sugars, 5-7 g/L acidity late-harvest or raisined styles targeting 25-35+% sugars with higher phenolic concentration. Mature berries for table grape consumption target different criteria= sugars 16-22% for eating quality moderate acidity 4-7 g/L for flavor balance specific textural properties (firmness for shipping cultivars, crisp versus soft for different markets) size considerations (larger berries preferred for fresh market).
Mature_Berry represents harvest-ready and post-harvest states (BBCH 89+) and is distinguished from Developing_Berry (pre-harvest developmental states BBCH 71-89) by completion versus continuation of development. Mature berries encompass multiple states including optimal ripeness, overripeness, and post-harvest, reflecting the complexity of maturity assessment and harvest timing decisions. Sub-classification into Fully_Ripe, Overripe, Harvested, and Post_Veraison reflects different aspects of maturity and post-maturity states. Understanding mature berry characteristics is essential for harvest timing decisions, quality assessment, post-harvest handling, and for all aspects of grape utilization in wine or fresh market.
Erntereife Traube
Vollreife Beere
reife Beere
ripe grape
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 220-280 Coombe, B.G. (1992). Research on development and ripening of the grape berry. American Journal of Enology and Viticulture 43=101-110 Jackson, D.I., Lombard, P.B. (1993). Environmental and management practices affecting grape composition and wine quality. American Journal of Enology and Viticulture 44=409-430.
Mature Berry
In Riesling berries in Mosel Valley, Germany at BBCH 89, mature berry acidity is characterized by titratable acidity of 9.2 g/L, pH 3.08, malic acid of 5.8 g/L, and tartaric acid of 7.5 g/L, representing the high-acid profile typical of this variety under cool-climate conditions with mean ripening temperature of 15°C, providing optimal base material for production of off-dry to sweet wines with characteristic fresh acidity.
Mature berry acidity describes the characteristic acid composition and concentration attained in fully ripe Vitis vinifera berries at the conclusion of the ripening process, wherein titratable acidity has stabilized at harvest-appropriate levels of 5-9 g/L with pH values of 3.2-3.8 that reflect the integrated outcomes of malic acid catabolism, tartaric acid stability, and potassium-mediated pH buffering during the preceding 35-50 day post-veraison maturation period. This state represents the optimal acid status for wine production, characterized by residual malic acid of 1-4 g/L providing microbial substrate for potential malolactic fermentation, tartaric acid of 5-8 g/L constituting the primary structural acid for wine stability, and sufficient total acidity to ensure proper pH for antimicrobial protection, color stability in red wines, and fresh sensory character in the finished wine. The biochemical markers of mature berry acidity include completion of rapid-phase malate degradation with transition to equilibrium kinetics showing daily acid loss below 0.1 g/L, stable vacuolar pH reflecting exhaustion of readily catabolizable acid pools, and establishment of potassium-tartrate precipitation equilibrium determining final tartrate availability and pH buffering capacity. Temperature history during ripening determines the specific parameters of mature berry acidity, with warm-climate berries typically achieving maturity at lower titratable acidity of 5-7 g/L and higher pH of 3.5-3.8 compared to cool-climate berries maturing at titratable acidity of 7-10 g/L and pH of 3.1-3.4, reflecting the temperature sensitivity of malate degradation rates that establish final acid composition. Cultivar genetics interact with environmental conditions to determine mature berry acidity parameters, with inherently high-acid varieties such as Riesling, Chenin Blanc, and Barbera maintaining elevated acidity at maturity compared to lower-acid varieties such as Gewürztraminer, Viognier, and Grenache under equivalent growing conditions. The assessment of mature berry acidity requires integration of multiple analytical parameters including titratable acidity, pH, and individual organic acid concentrations, as these metrics provide complementary information about wine suitability, with titratable acidity predicting sourness perception, pH determining chemical and microbial stability, and organic acid composition indicating malolactic fermentation potential and wine aging trajectory.
Mature_Berry_Acidity is subclass of Final_Acid_Balance representing the quality state at ripening completion. Distinguishes from Harvest_Acidity_Level which emphasizes measurement for picking decisions. Sibling to Sugar_Acid_Ratio_Optimization and Harvest_Acidity_Level. Characterized by titratable acidity 5-9 g/L, pH 3.2-3.8, and equilibrium acid metabolism. Integrates temperature and cultivar effects on final composition.
Reife-Beeren-Säurestatus
erntereife Säure-Zusammensetzung
fully ripe acid profile
harvest-ready acid composition
mature grape acidity
reife Beeren-Acidität
ripe berry acid status
vollreifes Säureprofil
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 275-280 Jackson RS (2020) Wine Science Principles and Applications 5th Ed Chapter 6 Coombe BG and McCarthy MG (2000) Dynamics of grape berry growth. Australian Journal of Grape and Wine Research 6: 131-135
Mature Berry Acidity
Cabernet Sauvignon mature berry pedicel at harvest (BBCH 89, 24°Brix) exhibits complete lignification with dark brown woody appearance, well-developed abscission zone requiring only 0.8-1.2 N detachment force, largely ceased xylem transport with embolized vessels, minimal phloem activity (0.1-0.2 mg sugar/berry/hour), and separation layer cells showing cellulase and pectinase accumulation enabling clean berry separation
A mature berry pedicel is a berry pedicel that has completed developmental lignification, developed functional abscission zone tissues, and achieved reduced vascular transport activity, representing the pedicel state at berry harvest maturity facilitating controlled berry separation. Mature berry pedicel characteristics include= complete or near-complete lignification (brown to woody appearance, rigid structure), well-developed abscission zone with differentiated separation layer (facilitating 0.5-1.5 N detachment force, significantly reduced from 1.5-3.0 N in attached pedicels), reduced or ceased xylem transport (xylem vessels may become embolized or blocked), declining phloem transport as berry sugar accumulation completes, and structural integrity sufficient for berry support but enabling clean separation at abscission zone. Anatomical features of mature berry pedicel include= heavily lignified epidermal and cortical cell walls (providing brown coloration and woody texture), mature abscission zone with cell wall degrading enzyme accumulation (cellulases, pectinases in separation layer), vascular bundles with sclerified walls and reduced conducting capacity (some xylem vessels blocked with tyloses, phloem sieve tubes showing reduced activity), and parenchyma cells with reduced metabolic activity and increased lignification. Mature berry pedicel development timing varies by cultivar and environmental conditions= early-ripening varieties (e.g., Pinot Noir, Chardonnay) typically show mature pedicels at BBCH 87-89 (18-22°Brix, 8-12 weeks post-véraison), late-ripening varieties (e.g., Cabernet Sauvignon, Nebbiolo) may show mature pedicels at BBCH 89+ (22-26°Brix, 10-14 weeks post-véraison), with timing influenced by temperature, water status, and crop load. Mature berry pedicel vascular function shows characteristic changes= xylem transport largely ceased with vessel embolism or blockage (water import declining to near zero, some xylem backflow may occur during berry dehydration), phloem transport declining as berry sugar accumulation plateaus (sugar import rates declining from 1-2 mg/berry/hour at mid-ripening to 0.1-0.3 mg/berry/hour at full maturity), and overall berry becomes increasingly hydraulically isolated from vine. Mature berry pedicel abscission zone development includes= differentiation of separation layer cells (small cells with thin walls positioned across pedicel in abscission zone region), accumulation of cell wall degrading enzymes (cellulase, polygalacturonase hydrolyzing pectin and cellulose in middle lamella and primary walls), potential formation of protective layer (suberization or lignification of cells adjacent to separation layer preventing pathogen entry after separation), and ethylene-responsive gene expression (abscission-associated genes induced by ripening ethylene production). Viticultural implications of mature berry pedicel include= harvest ease (mature abscission zones facilitate gentle berry separation during hand or mechanical harvest), reduced shattering risk during transport (berries separate cleanly at intended separation point rather than tearing), quality considerations (clean pedicel separation reduces stem fragment contamination in must), and optimal harvest timing (pedicel maturity correlates with berry physiological maturity in most cultivars).
Mature_Berry_Pedicel emphasizes developmental completion and abscission readiness and is distinguished from Attached_Berry_Pedicel (firm connection, incomplete abscission) by lignification completion, reduced vascular activity, and functional abscission zone facilitating controlled separation. Critical for harvest timing assessment, mechanical harvest efficiency, berry separation quality, and understanding ripening coordination between berry maturity and pedicel abscission development.
Ausgereifter Stiel
Erntereifer Fruchtstiel
Harvest-ready berry stalk
Reifer Beerenstiel
Ripened pedicel
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Patterson, S. E. (2001). Cutting loose. Abscission and dehiscence in Arabidopsis. Plant Physiology, 126(2), 494-500.
Mature Berry Pedicel
In a Chianti Classico Sangiovese vineyard in Tuscany, mature berry texture was assessed in late September Year N+1 (BBCH 87-89) showing compression firmness 1.9-2.1 N (indicating characteristic firm Sangiovese texture), puncture resistance 2.1-2.3 N reflecting thick skin, sensory evaluation describing moderately firm flesh with balanced juiciness, chemical maturity at 23.8-24.2° Brix and 5.8-6.1 g/L TA, skin anthocyanins 1950-2100 mg/kg, and brown seed tannins confirming integrated phenolic and textural maturity under typical Tuscan autumn conditions (daily high 22-26°C, cool nights 12-15°C).
Mature_Berry_Texture represents the integrated sensory and mechanical quality of fully ripened grape berries characterized by cultivar-specific tactile properties, deformability characteristics, and handling responses that reflect completed softening processes and define berry suitability for harvest and processing. This mature texture emerges during late veraison through early ripening (BBCH 85-89) and encompasses the full spectrum of physical berry attributes: skin mechanical properties including tensile strength (40-65% of green berry baseline), puncture resistance (1.0-2.5 N cultivar-dependent), and surface characteristics (smooth, slightly tacky from accumulated sugars and waxes, easily indented by finger pressure 0.5-1.5 kg force), flesh firmness and juiciness reflecting cell wall degradation state and water content (mesocarp readily yielding under gentle compression, juice easily expressed with 1.5-3.0 N force), and overall berry deformability showing plastic rather than elastic behavior (permanent shape change under moderate compression, minimal elastic recovery indicating extensive cell wall modification). Sensory evaluation of mature berry texture by trained assessors reveals characteristic quality descriptors: skin texture described as thin-to-moderate thickness with easy tearing (compared to tough, resistant green berry skin), flesh described as soft, yielding, juicy with minimal fibrousness (versus firm, crunchy, dry green berry flesh), and overall berry described as readily crushable between fingers with copious juice release and minimal resistance (contrasting with green berry resistance requiring substantial force and producing little free juice). Mechanical measurement of mature berry texture provides quantitative assessment: compression tests showing force-deformation curves with initial linear elastic region (0-15% deformation) followed by extensive plastic yield region (15-60% deformation) and final densification (>60% deformation) where crushed berry remnants resist further compression, with key parameters including yield point force (1.2-2.8 N cultivar-dependent), work to rupture skin (0.8-2.5 mJ), and compression modulus (100-300 kPa indicating soft-moderate texture compared to 300-600 kPa for firm green berries). Puncture testing reveals skin strength characteristics: maximum puncture force required to penetrate skin with standardized probe (2mm diameter) ranging 1.0-2.5 N depending on cultivar (soft-skinned varieties 1.0-1.5 N, thick-skinned 1.8-2.5 N), work to puncture (0.5-1.8 mJ) reflecting total energy to breach skin, and post-puncture force (40-70% of maximum) indicating flesh firmness after skin penetration. Cellular basis of mature berry texture includes: extensively degraded cell walls (70-85% pectin solubilization, 60-120 kDa average pectin molecular weight) providing minimal structural resistance, reduced turgor pressure (0.3-0.6 MPa) contributing only 15-25% to overall firmness, large vacuoles (85-95% of cell volume) filled with sugar-rich juice creating characteristic juiciness, and weakened intercellular adhesion (60-80% middle lamella degradation) allowing easy cell separation during mastication or crushing. Mature berry texture varies systematically among cultivars reflecting genetic differences in cell wall composition and degradation extent: very soft varieties (Pinot Noir, Gamay, Grenache) exhibit 0.8-1.5 N compression firmness with delicate skin and highly degraded flesh producing extremely juicy, fragile berries requiring careful handling and typically precluding mechanical harvest, medium-textured cultivars (Merlot, Syrah, Chardonnay, Sauvignon Blanc) show 1.2-2.0 N firmness with balanced skin strength and flesh firmness suitable for mechanical or hand harvest and varied wine styles, while firm-textured varieties (Cabernet Sauvignon, Nebbiolo, Sangiovese, Tempranillo) maintain 1.5-2.5 N firmness with thicker skins and more resistant flesh tolerating mechanical harvest and producing wines with substantial structure and aging potential. Mature berry texture profoundly affects processing characteristics: soft berries with low skin strength (1.0-1.5 N puncture resistance) crush easily under minimal force (1-2 bar crusher pressure) yielding high free-run must (60-75% total juice) but risk excessive skin maceration and phenolic over-extraction, medium berries (1.5-2.0 N) require moderate crushing force (2-3 bar) producing balanced free-run (55-70%) suitable for controlled extraction and varied wine styles, while firm berries (2.0-2.5 N) need substantial crushing (3-4 bar) yielding lower free-run (45-60%) but allowing extended maceration without excessive extraction and producing age-worthy structured wines. Temperature effects on mature berry texture inclu cool berry temperature (35°C) can cause partial cellular breakdown and membrane disruption producing abnormally soft texture with potential quality degradation, and diurnal temperature variation creates texture fluctuations (firmer in cool morning, softer in warm afternoon) affecting optimal harvest timing particularly for mechanical operations preferring firmer morning berries. Water relations influence mature berry texture dynamics: adequate water supply maintains optimal texture through balanced turgor supporting remaining firmness (0.4-0.6 MPa) without excessive softness, moderate deficit can maintain slightly firmer texture potentially beneficial for thick-skinned varieties while risking insufficient softening in naturally firm cultivars, while severe deficit causes desiccation-induced abnormal softening or shriveling compromising texture quality and juice yield. Mature berry texture assessment provides critical harvest timing information: tactile evaluation by experienced viticulturists squeezing representative berries from multiple cluster positions assessing readiness (too firm suggests incomplete ripening, excessively soft indicates over-ripeness or senescence), instrumental firmness testing with penetrometer or texturometer providing quantitative benchmarks calibrated to desired wine styles (sparkling wines 1.8-2.5 N, premium reds 1.2-2.0 N, late harvest <1.5 N), and combined assessment with chemical maturity (sugar, acid, pH) and phenolic ripeness determines optimal harvest windows balancing texture, composition, and quality goals. Mature berry texture stability and harvest window depends on cultivar and environmental conditions: stable varieties maintaining consistent texture 3-5 weeks (Cabernet Sauvignon, Syrah, Sangiovese) allow flexible scheduling and selective harvest, moderately stable cultivars (Merlot, Chardonnay) provide 2-3 week windows requiring reasonable promptness, while unstable varieties (Pinot Noir, Gewürztraminer, Riesling) may maintain optimal texture only 1-2 weeks necessitating precise harvest timing to capture peak quality before over-softening or senescence degradation. Quality implications of mature berry texture inclu optimal texture synchronized with chemical and phenolic maturity produces balanced grapes for premium wines with appropriate extraction potential, mouthfeel characteristics, and aging capacity, discordant maturity with premature softening before flavor development yields unbalanced wines with hollow mid-palate and insufficient complexity, while delayed softening past chemical maturity creates processing challenges and potentially harsh, extracted wines from over-firm berries requiring excessive maceration.
Mature_Berry_Texture is distinguished from Final_Softening_State by representing the integrated sensory and functional quality rather than just mechanical properties, from Complete_Berry_Softening by encompassing all berry texture attributes beyond firmness decline, and from immature berry texture by exhibiting characteristic ripe attributes suitable for harvest. Compression firmness ranges: very soft 0.8-1.5 N, medium 1.2-2.0 N, firm 1.5-2.5 N. Skin contribution to texture 30-45%, flesh firmness 55-70%. Mature_Berry_Texture assessment is essential for harvest timing decisions integrating mechanical, sensory, chemical, and phenolic maturity parameters to optimize grape quality for intended wine style.
characteristic varietal texture
charakteristische Sorten-Textur
completed texture development
erntereife Beeren-Haptik
harvest-ready berry feel
reife Beeren-taktile Qualität
ripe berry tactile quality
vollendete Textur-Entwicklung
Keller M (2020) The Science of Grapevines 3rd Ed pages 235-238 berry texture and maturity assessment Rolle L, Torchio F, Ferrandino A, Guidoni S (2012) Influence of wine-grape skin hardness on the kinetics of anthocyanin extraction Intl J Food Properties 15:249-261 mature berry texture characteristics Letaief H, Rolle L, Zeppa G, Gerbi V (2008) Assessment of grape skin hardness by a puncture test J Sci Food Agric 88:1567-1575 texture measurement methodology.
Mature Berry Texture
Mature color quality in Tempranillo defines the harvest window for premium Ribera del Duero production, with optimal fruit displaying deep purple-black coloration (CIELAB L* 25-32, a* 28-40, C* 35-50) and uniform distribution across at least 95% of berry surface area berries failing to achieve full color development due to excessive crop load or heat stress during ripening show mature color values shifted toward red-purple (higher h° angle, lower C*) and are diverted to younger wine categories, while fruit with optimal mature color commands premium pricing for age-worthy Reserva and Gran Reserva wines.
Mature_Color_Quality represents the final coloration state achieved by grape berries at harvest maturity, reflecting the completed developmental program of pigment accumulation and the integration of genetic potential with environmental influences experienced during ripening. Mature color is a primary quality determinant for both wine and table grapes, affecting consumer perception, market value, and wine style potential. In colored cultivars, mature color results from the accumulated anthocyanin content in berry skins, with characteristic hues ranging from pink-red in lightly colored cultivars through deep purple to blue-black in intensely pigmented varieties. The specific mature color reflects the cultivar's anthocyanin profile (proportions of different anthocyanin types), total pigment concentration, co-pigmentation effects, and the chemical environment including pH and metal ion content that influences pigment expression. In white cultivars, mature color spans the range from pale green through green-yellow to golden depending on cultivar character and ripening conditions. Mature color assessment at harvest determines fruit allocation decisions for different wine styles, table grape market positioning, and in some regions, quality-based pricing structures that reward optimal coloration. Instrumental measurement of mature color employs colorimetry (CIELAB coordinates), spectrophotometry (absorbance spectra), or chemical analysis of extracted pigments. The relationship between berry color at harvest and resulting wine color depends on extraction efficiency during fermentation, making berry color a necessary but not sufficient predictor of wine color. Mature color stability during postharvest handling affects table grape storage life, with some cultivars maintaining color while others develop browning or color loss.
Mature_Color_Quality represents the endpoint of developmental color change, distinct from Véraison_Color_Quality which captures the transitional state. The definition of mature color involves judgment about when development is complete, as subtle color changes may continue through the harvest window and even postharvest. For quality assessment purposes, mature color is typically evaluated at the moment of harvest decision or at standardized maturity indices (e.g., specific sugar concentration). The stability of mature color during the harvest window varies among cultivars some varieties maintain stable color while others show degradation under continued heat exposure, making harvest timing critical for color quality optimization.
Erntefarbe
Reifefärbung
Reifestadienfärbung
ausgereifte Beerenfarbe
final berry color
harvest color
maturity stage color
ripe fruit coloration
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed, John Wiley and Sons pp.: 141-203 González-Neves G et al. (2004) Anthocyanic Composition of Tannat Grapes from the South Region of Uruguay. Analytica Chimica Acta 513(1): 197-202
Mature Color Quality
Wild V. vinifera ssp. sylvestris female plants during the bloom period (BBCH 61-69, spanning 10-16 days) exhibit mature female only flowers across a range of developmental states= pre-anthesis mature flowers (BBCH 59, calyptra swollen, pistil fully developed 2.4-2.6 mm visible through translucent calyptra, stigma beginning exudate production, 1-3 days before shedding), fully developed flowers (BBCH 65, calyptra shed, stigma at peak receptivity with abundant wet exudate, 2-4 day peak window), and post-receptivity mature flowers (BBCH 67-69, stigma declining in exudate production and receptivity, papillae browning, 4-8 days post-peak), with the extended maturity window (total 12-15 days) increasing probability of successful cross-pollination from stochastic wind-borne pollen arrival, achieving cumulative fruit set of 30-42% across the full receptivity period compared to 38-44% during peak receptivity alone, demonstrating that extended receptivity captures 70-95% of the fruit set achieved during peak receptivity through successful late-arriving pollen fertilization events.
A mature female only flower is a female only flower that has achieved full developmental maturity with completed pistil organ differentiation and structural development, encompassing both pre-anthesis flowers with mature but still-enclosed pistils and post-anthesis flowers that have passed peak receptivity but retain functional capacity for fertilization, representing the broader category of developmentally complete female only flowers across the full maturity window. Mature female only flowers are characterized by= (1) completed pistil development with pistil achieving full length potential (2.2-2.8 mm), ovary reaching full size (1.3-1.7 mm diameter) with mature ovules (3-4 ovules per ovary at developmental stages ready for or capable of fertilization), style fully elongated with mature transmitting tract tissue, and stigma fully differentiated with elaborate papillae and secretory cell capacity (2) pistil tissues achieving adult cellular characteristics with fully expanded cells, mature cell wall composition, complete vascular tissue differentiation supplying the pistil, and full biochemical maturation of stigmatic exudate-producing cells (3) structural maturity of all flower components including mature calyptra ready for shedding (pre-anthesis) or already shed (post-anthesis), mature receptacle and nectary disk, and complete absence or maximum reduction of stamen structures (4) flower capable of supporting successful pollination and fertilization when pollen arrives, though receptivity efficiency varies across the maturity window. The 'mature' designation encompasses a broader developmental window than 'fully developed,' including= mature pre-anthesis flowers (1-3 days before calyptra shedding, with mature pistil ready for anthesis but calyptra still attached and stigma not yet fully exposed), fully developed anthetic flowers (at peak stigma receptivity with calyptra shed and stigma optimally receptive), and mature post-receptivity flowers (3-7 days after peak receptivity, with stigma declining in receptivity but still capable of supporting fertilization if pollen arrives). Mature female only flowers in the pre-anthesis state show swollen flower buds with calyptra beginning to separate from receptacle base, pistil visible through translucent calyptra showing full size and development, and stigma already beginning minimal exudate production in preparation for exposure. Mature female only flowers in the post-receptivity state show stigmatic exudate production declining or ceased, papillae beginning to brown or collapse as senescence proceeds, but stigma retaining some residual receptivity capability for pollen germination, with successful fertilization still possible (at reduced probability) if viable pollen arrives during this extended window. The mature state represents the full period of structural integrity and potential reproductive competence, typically spanning 8-14 days from achievement of maturity (2-3 days pre-anthesis) through loss of fertilization capability (5-8 days post-peak receptivity), compared to the 2-4 day window of peak receptivity in fully developed flowers. Stigma receptivity across the maturity window varies substantially= pre-anthesis mature flowers show minimal receptivity (pollen germination <20% if calyptra artificially removed and pollen applied) due to incomplete stigmatic exudate production fully developed anthetic flowers show peak receptivity (pollen germination 75-92% on mature wet stigma) post-receptivity mature flowers show declining receptivity (pollen germination declining from 60-75% at 2 days post-peak to 25-40% at 5 days post-peak to <10-20% at 7+ days post-peak). The extended maturity window is adaptively significant for female only flowers in dioecious populations, as the stochastic nature of wind-borne pollen arrival makes extended receptivity advantageous= maintaining some fertilization capability across 8-14 days rather than only during a 2-4 day peak window substantially increases probability that at least some pollen grains will arrive during a receptive period, even when pollen sources are distant or environmental conditions intermittently unfavorable for pollen dispersal.
Mature_Female_Only_Flower represents the broader developmental category encompassing the full maturity window and is distinguished from Fully_Developed_Female_Only_Flower by broader temporal scope (pre-anthesis through post-receptivity versus peak receptivity only). The mature category is useful for describing the complete period of structural integrity and potential reproductive competence, and for understanding extended stigma receptivity strategies that increase reproductive success probability in stochastic cross-pollination systems. The distinction parallels that made for staminate flowers between mature (broad developmental window) and fully developed (peak functional state). Understanding the full maturity window and the decline in receptivity efficiency post-peak informs models of reproductive success in dioecious populations where temporal dynamics of pollen availability and stigma receptivity must overlap for successful fertilization.
adult female flower
ausgereifte Pistillatblüte
developmentally mature female flower
entwicklungsmäßig reife weibliche Blüte
reife weibliche Blüte
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 104-108 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 100-108 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 191-198.
Mature Female Only Flower
In Grüner Veltliner berries in Wagram, Austria at BBCH 89, mature flavor expression is characterized by total monoterpenes of 0.95 mg/L with citrus-floral balance, norisoprenoid precursors of 145 µg/L providing stone-fruit complexity, methoxypyrazine below detection threshold of 2 ng/L ensuring absence of green notes, and volatile thiol precursors of 210 µg/L contributing characteristic white pepper nuance, with trained panel overall quality rating of 8.4 on 10-point scale and varietal typicity score of 8.7 indicating achievement of optimal maturity under moderate continental conditions.
Mature flavor expression describes the characteristic aromatic profile achieved in fully ripe Vitis vinifera berries at the conclusion of the ripening process, wherein flavor compound composition has stabilized at harvest-appropriate levels and the overall sensory impression reflects the balanced integration of varietal character, ripeness indicators, and site-specific nuances that define quality wine grape maturity and provide the aromatic foundation for premium wine production. This state encompasses specific compositional markers including monoterpene concentrations of 1-5 mg/L in aromatic varieties providing characteristic floral-fruity impact, norisoprenoid levels of 50-200 µg/L contributing complexity and aging potential, minimal residual methoxypyrazine below 10-15 ng/L ensuring absence of green-unripe character, and balanced thiol precursor concentrations providing tropical fruit potential without excessive pungency in the finished wine. The sensory characteristics of mature flavor expression include predominance of ripe-fruit aromatics over green-herbaceous notes, integration of multiple aromatic elements into harmonious overall impression, appropriate intensity matching varietal expectations, and absence of overripe or degraded character indicating optimal harvest timing capture. Assessment of mature flavor expression employs multiple evaluation methods including trained panel sensory analysis rating specific aromatic attributes and overall quality, berry tasting protocols assessing flavor development through direct consumption of representative samples, and analytical profiling quantifying marker compounds associated with optimal maturity for specific cultivars and wine styles. Temperature history during the final ripening phase influences mature flavor expression through effects on late-stage biosynthesis and compound stability, with optimal conditions maintaining biosynthetic activities while avoiding thermal degradation that could compromise accumulated aromatic quality. The relationship between mature flavor expression and other ripeness parameters including sugar content, acidity, and phenolic development varies among cultivars and production objectives, with some varieties achieving optimal flavor expression coincident with target compositional values while others may require management decisions balancing competing quality factors for overall optimization.
Mature_Flavor_Expression is subclass of Flavor_Profile_Maturation representing the achieved quality state at ripening completion. Distinguishes from Optimal_Flavor_Development which emphasizes process optimization. Sibling to Cultivar_Specific_Flavor and Optimal_Flavor_Development. Characterized by specific compositional markers and sensory attributes. Integrates intensity, complexity, and balance. Harvest maturity indicator.
harvest-ready flavor expression
mature aromatic profile
optimal flavor state
optimaler Aroma-Zustand
reifer Geschmacks-Charakter
ripe flavor character
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 305-315 Reynolds AG (2010) Managing Wine Quality Vol 1 Viticulture and Wine Quality Woodhead Publishing Coombe BG and McCarthy MG (2000) Dynamics of grape berry growth. Australian Journal of Grape and Wine Research 6: 131-135
Mature Flavor Expression
Tempranillo grafted on 110 Richter rootstock at 18 years post-grafting showing mature grafted rootstock trunk 8.4 cm diameter, trunk extending 26 cm from root crown to graft union positioned 21 cm above soil, graft union displaying stable 1.4 cm diameter bulge fully integrated and obscured by thick bark, mature trunk bark 3.2 mm thickness showing deep longitudinal fissures characteristic of 110R with dark grey-brown weathered coloration, trunk fully self-supporting, diameter growth 0.4 cm current season, no disease symptoms, supporting vigorous canopy 4.6 m total shoot length and 8.2 kg fruit load, vine in optimal production
Mature grafted rootstock trunk is grafted rootstock trunk that has achieved full structural and physiological development spanning approximately years 8-30 post-grafting, characterized by complete graft union integration with stable callus tissue, well-developed characteristic bark with fully expressed rootstock cultivar texture, diameter typically 5-10 cm providing adequate capacity for commercial production, stable annual growth rate of 0.3-0.6 cm, complete self-supporting structural capacity, and representing optimal productive phase where trunk provides reliable foundation for sustained high-quality fruit production over decades. Mature grafted rootstock trunk age and developmental status includes= spanning years 8-30 post-grafting typically (exact timing varying with rootstock vigor, cultivar, and growing conditions), representing peak productive period (optimal phase for reliable commercial fruit production with stable trunk capacity), showing complete structural development (trunk achieving full diameter, bark thickness, and wood properties characteristic of mature status), displaying stable physiological function (consistent annual growth and resource transport patterns without developmental transitions), and marking prime phase (period between establishment completion and onset of old age characteristics). Mature grafted rootstock trunk complete integration includes= showing fully consolidated graft union (all callus tissue mature and integrated into trunk structure with no ongoing active callus formation), displaying perfect vascular continuity (xylem and phloem connections across graft union functionally equivalent to continuous non-grafted trunk), exhibiting stable graft union morphology (bulge at graft union stabilized at 0.8-2.2 cm diameter increase with no further expansion), showing obscured graft line (thick mature bark and complete callus integration making original grafting point difficult to identify visually), and demonstrating maximum mechanical strength (graft union integration complete providing full structural continuity equivalent to non-grafted trunk). Mature grafted rootstock trunk physical characteristics includes= exhibiting typical mature diameter (5-10 cm range depending on rootstock vigor, cultivar, years since grafting, and growing conditions), showing well-developed mature bark (periderm 2-5 mm thickness with fully expressed rootstock-specific texture, color, and fissure patterns), displaying characteristic rootstock bark (bark clearly showing variety-specific features such as deep fissures in Rupestris types, moderate fissuring in SO4, or smoother texture in Riparia types), revealing mature grey-brown coloration (weathered bark colors typical of rootstock variety fully developed), and showing robust substantial appearance (trunk looking solid, well-established, and capable of supporting large productive vine). Mature grafted rootstock trunk stable growth patterns includes= showing consistent annual diameter increase (0.3-0.6 cm typical mature trunk growth rate varying with vigor, site conditions, and crop load), displaying regular annual ring formation (clear distinct growth rings in cross-section marking each growing season), exhibiting predictable seasonal growth (diameter expansion occurring during primary growth period May-August in Northern Hemisphere), maintaining stable wood properties (consistent wood density, vessel size, and mechanical characteristics year to year), and showing reduced growth variability (annual diameter increase more consistent than in young trunk with less dramatic year-to-year fluctuation). Mature grafted rootstock trunk vascular capacity includes= providing optimal water transport (xylem capacity 50-100+ liters daily supporting full productive canopy during peak demand periods), enabling complete nutrient distribution (adequate phloem and xylem capacity distributing all mineral nutrients and photosynthates throughout vine), showing maximum transport efficiency (well-developed vascular architecture minimizing transport resistance and maximizing flow rates), supporting consistent crop loads (trunk capacity adequate for sustained annual fruit production of 4-12+ kg per vine depending on cultivar and training), and maintaining substantial reserves (extensive ray parenchyma storing significant carbohydrate reserves supporting vine resilience and consistent performance). Mature grafted rootstock trunk structural capability includes= achieving maximum mechanical strength (optimal wood density, diameter, and lignification providing complete structural support), showing rigid stable properties (mature wood stiff and strong resisting bending, wind stress, and mechanical load), supporting substantial canopy mass (bearing weight of large productive canopy, cordon system, and fruit load typically 15-40 kg total vine mass), maintaining permanent upright position (self-supporting trunk holding vine erect throughout all weather conditions and developmental stages), and providing reliable foundation (robust trunk serving as stable base for permanent training system and annual production cycle). Mature grafted rootstock trunk complete rootstock expression includes= conferring full phylloxera resistance (rootstock genetics providing complete protection to root system allowing susceptible Vitis vinifera scion to thrive), expressing vigor control characteristics (rootstock variety determining vine growth intensity with mature trunk fully manifesting low, moderate, or high vigor effects), providing optimal drought adaptation (mature rootstock trunk and associated root system expressing full drought tolerance potential of rootstock genetics), enabling maximum lime tolerance (rootstock allowing growth in calcareous soils where direct-planted scion would show severe chlorosis), and contributing other resistances (nematode tolerance, salt tolerance, or cold hardiness depending on rootstock variety fully expressed in mature trunk and root system). Mature grafted rootstock trunk management requirements includes= needing standard vineyard maintenance only (mature trunk requiring no special establishment-focused care, just normal production viticulture practices), monitoring for trunk diseases (inspecting for cankers, crown gall, or decay though mature trunk showing good disease resistance), continuing sucker removal (eliminating any rootstock shoots below graft union though typically minimal in well-established mature trunk), implementing appropriate pruning (managing scion canopy to maintain balance with mature trunk capacity avoiding overload or excessive vigor), and assessing trunk health (periodic evaluation of bark condition, graft union integrity, and absence of damage or disease). Mature grafted rootstock trunk cultivar-specific characteristics includes= vigorous rootstocks (1103P, 110R, 140Ru) showing mature trunk diameter 7-12 cm and very thick rough bark, moderate vigor rootstocks (SO4, 3309C, Teleki 5C, 5BB) exhibiting mature trunk 5-8 cm diameter with moderately furrowed bark, low vigor rootstocks (Riparia Gloire, 101-14, 3306C) displaying mature trunk 4-7 cm diameter with thinner smoother bark, thick-bark cultivars showing deep longitudinal fissures and substantial cork layer, and smooth-bark cultivars maintaining relatively smooth texture even in mature stage. Mature grafted rootstock trunk significance includes= representing optimal productive phase (mature trunk marking peak period of reliable commercial fruit production with stable trunk capacity), providing foundation for quality viticulture (robust mature trunk enabling consistent high-quality grape production year after year), demonstrating successful grafting (healthy mature trunk confirming long-term compatibility and integration of rootstock-scion combination), enabling economic sustainability (mature trunk supporting decades of productive life providing return on vineyard establishment investment), and exemplifying modern viticulture (mature grafted rootstock trunks representing standard commercial approach combining phylloxera resistance with quality fruit production worldwide).
Mature_Grafted_Rootstock_Trunk emphasizes fully developed productive phase (years 8-30) and is distinguished from Young_Grafted_Rootstock_Trunk and Establishing_Grafted_Rootstock_Trunk (development phases) and Old_Grafted_Rootstock_Trunk (declining phase) by focus on optimal mature productive status. Critical for understanding complete integration, stable growth, maximum capacity, optimal production, and the prime phase of commercial grafted vine life supporting decades of reliable fruit production.
Established grafted rootstock trunk
Etablierter Wurzelstockstamm
Fully developed grafted rootstock trunk
Prime grafted rootstock trunk
Reifer verpfropfter Unterlagenstamm
Voll entwickelter Pfropfstamm
Mature Grafted Rootstock Trunk
Syrah mature grafted scion trunk on 1103P vigorous rootstock in Rhône Valley at 12 years post-planting shows 82-98 mm scion diameter with 12 annual rings averaging 3.2-4.8 mm width (wider than moderate rootstock equivalent due to rootstock vigor), graft union at 9 cm above soil with stable 112-118% swelling, rootstock diameter 88-105 mm below union, scion sapwood 22-28 mm conducting 45-65 L/day, 140-175 g combined starch reserves (scion 50-65 g, rootstock 90-110 g), and consistent annual production 6.5-8.5 kg per vine (85-110 hL/ha) demonstrating vigorous rootstock productivity enhancement
A mature grafted scion trunk is a grafted scion trunk that has completed architectural development and entered sustained productive phase (typically years 5-25 post-planting) characterized by stable annual growth, optimal fruit production, established rootstock-scion physiological interaction, and confirmed long-term graft union stability, representing the prime productive period for commercial grafted viticulture with rootstock benefits fully expressed. Mature grafted scion trunk development reflects= completion of training architecture (cordons or head positions established and stable), stabilized graft union (union diameter ratio 105-120% maintained across years, union mechanically strong and physiologically functional), annual scion growth rings of relatively uniform width (1.5-3.5 mm radial increment modulated by rootstock vigor), progressive scion diameter increase (from 45-75 mm at year 5 to 85-140 mm at year 20 varying by rootstock class), rootstock trunk diameter tracking scion (rootstock/scion diameter ratio typically 95-115% at equivalent positions), and accumulation of scion heartwood (central 30-60% of scion trunk showing reduced transport, providing mechanical support). Mature grafted scion trunk anatomical characteristics include= scion secondary xylem with 5-20+ annual rings showing cultivar-specific ring-porous anatomy (large earlywood vessels reflecting scion genotype), functional scion sapwood (outer 15-30 mm radius conducting water with rootstock-mediated hydraulic properties), active scion vascular cambium producing consistent increments (rootstock vigor influencing annual ring width), scion secondary phloem (2-4 mm thick with functional outer sieve tubes), scion protective periderm (4-8 mm bark providing protection), and stable graft union vascular continuity (xylem and phloem bridging union supporting efficient transport). Mature grafted scion trunk graft union characteristics include= stable union diameter (swelling maintained at 108-120% without progressive enlargement suggesting stable compatibility), lignified union callus (woody graft interface providing mechanical strength), union bark continuity (protective periderm covering union reducing pathogen entry), efficient union hydraulic function (hydraulic conductance across union 80-95% of non-grafted trunk equivalent), and union as permanent feature (determining trunk height and architecture with union 5-15 cm above soil). Mature grafted scion trunk rootstock-mediated physiological characteristics include= rootstock vigor effects on scion growth (vigorous rootstocks promoting larger canopy and higher yields, dwarfing rootstocks reducing vigor and crop), rootstock water relations influence (drought-tolerant rootstocks providing scion stress resistance, shallow-rooted rootstocks showing earlier water deficit), rootstock nutrient acquisition (efficient rootstocks enhancing scion nutrition, lime-tolerant rootstocks preventing chlorosis, salt-tolerant rootstocks enabling production on saline soils), rootstock phenology effects (early rootstocks advancing budbreak 5-10 days, late rootstocks delaying maturity), and rootstock carbohydrate storage (combined scion and rootstock reserves typically 100-180 g= scion 35-65 g, rootstock 65-115 g). Mature grafted scion trunk productivity characteristics include= stable annual yield influenced by rootstock (vigorous rootstocks supporting 5-10 kg per vine, moderate 4-7 kg, low-vigor 3-5 kg, compared to 4-8 kg typical own-rooted), rootstock effects on fruit quality (some rootstocks enhancing color, phenolics, or aromatic compounds, others promoting higher yields potentially diluting quality), consistent phenology (budbreak, flowering, véraison showing cultivar timing modulated by rootstock), and balanced vegetative-reproductive development (rootstock vigor matched to site and training system producing appropriate canopy/crop ratio). Mature grafted scion trunk management during productive phase includes= annual pruning maintaining architecture and crop load (accounting for rootstock vigor in bud number selection= vigorous rootstocks 30-50 buds, moderate 20-35 buds, low-vigor 15-25 buds), canopy management appropriate for rootstock vigor (vigorous combinations requiring more intensive shoot management), continued rootstock sucker removal (vigilance required as rootstock may produce occasional suckers throughout vine life), disease management protecting union and trunk (union remains potential pathogen entry particularly for trunk diseases), and irrigation management considering rootstock characteristics (drought-tolerant rootstocks requiring less irrigation, water-demanding rootstocks needing supplemental water). Mature grafted scion trunk variation by rootstock class includes= vigorous rootstocks (SO4, 1103P, 110R, 140Ru) producing large scion trunks (90-150 mm at year 15-20, high productivity 6-10 kg per vine), moderate rootstocks (5BB, 420A, 5C, 101-14Mgt) producing intermediate trunks (70-110 mm, balanced productivity 4-7 kg), low-vigor rootstocks (3309C, Riparia Gloire) producing smaller trunks (55-95 mm, moderate productivity 3-5 kg), with rootstock adaptation to soil affecting mature trunk development (well-adapted combinations showing optimal growth, poorly adapted combinations underperforming despite matching rootstock characteristics).
Mature_Grafted_Scion_Trunk emphasizes established productive phase with rootstock and is distinguished from Young_Grafted_Scion_Trunk (establishment phase), Mature_Own_Rooted_Trunk (no union, no rootstock influence), and Old_Grafted_Scion_Trunk (declining phase) by stable optimal productivity with full rootstock benefit expression. Critical for understanding rootstock effects on mature vine productivity, graft union as permanent physiological feature, rootstock-mediated terroir expression, and optimization of grafted vine management for sustained commercial production.
Established grafted scion
Etablierter Aufpropfungsstamm
Productive grafted trunk
Produktionsphase Veredelungsstamm
Reifer gepfropfter Edelstamm
Mature Grafted Scion Trunk
A mature leaf at node 7 on a cane of Vitis vinifera cv. Chardonnay in Burgundy, France, measuring 12.5 cm length by 11.8 cm width representing stable final dimensions unchanged over three weeks of observation, exhibiting complete tissue development with two well-defined palisade layers totaling 85 μm thickness and spongy mesophyll of 120 μm, functional stomata on abaxial surface (density 180 stomata mm⁻²) showing rapid closure response to vapor pressure deficit increase, thick cuticle (4.2 μm) on adaxial surface providing water retention and pathogen resistance, mature vascular network with secondary xylem elements and companion cells visible in cross-section, demonstrating light-saturated photosynthesis rate of 12.3 μmol CO₂ m⁻² s⁻¹ at 1500 μmol photons m⁻² s⁻¹ irradiance measured with gas exchange system, maintaining peak photosynthetic performance contributing approximately 15 grams carbohydrate per week supporting fruit ripening in adjacent cluster, measured at BBCH stage 83 (berries developing color) in early August at 8 weeks post-emergence, representing optimal mature leaf status with maximum functional capacity.
A leaf that has completed cell division and expansion phases achieving final size and has developed full anatomical and physiological maturity in Vitis vinifera, characterized by cessation of lamina expansion with stable dimensions maintained over weeks to months, complete tissue differentiation including fully developed palisade and spongy mesophyll layers with mature chloroplasts, functional stomatal apparatus with responsive guard cells enabling regulated gas exchange, complete cuticle development providing protection against water loss and pathogen entry, full development of vascular networks with mature xylem and phloem enabling efficient transport of water, nutrients, and photosynthates, and achievement of maximum photosynthetic capacity with light-saturated photosynthesis rates typically 8-15 μmol CO₂ m⁻² s⁻¹ depending on cultivar and environmental conditions. Mature leaves represent the primary functional photosynthetic organs of productive grapevine canopies, typically developing 3-4 weeks after emergence from buds and maintaining peak photosynthetic performance for 6-12 weeks depending on environmental conditions and canopy position, producing the carbohydrates that support concurrent shoot growth, fruit development, and reserve accumulation in perennial structures. The transition from juvenile to mature status involves not only size achievement but critical physiological changes including increased photosynthetic enzyme concentrations, enhanced stomatal responsiveness to environmental signals, improved water use efficiency, and development of stress tolerance mechanisms including antioxidant systems protecting against photoinhibition and oxidative damage under high light or water deficit conditions.
Mature leaves are distinguished from juvenile leaves by completed expansion and tissue development, and from senescent leaves by maintenance of peak photosynthetic function without degradation processes. The maturation process typically requires 21-35 days post-emergence depending on shoot vigor, temperature, and resource availability, with exact timing varying by cultivar and environmental conditions. Mature leaf status should be assessed through multiple criteria including dimensional stability (no further expansion), tissue anatomy (complete differentiation), physiological function (peak photosynthesis), and biochemical composition (maximum chlorophyll and protein concentrations). Position on shoot affects maturation timing, with basal leaves maturing earliest while apical leaves continue development throughout shoot extension. Environmental conditions during development influence mature leaf characteristics through phenotypic plasticity, with sun versus shade exposure, water availability, and nutrient status creating functional specialization even among leaves of equivalent developmental stage.
adult leaf
physiologically mature leaf
physiologisch ausgereiftes Weinblatt
reifes Blatt
voll entwickeltes Blatt
Based on Flexas et al. (2002) "Drought-inhibition of photosynthesis in C₃ plants= stomatal and non-stomatal limitations revisited" Annals of Botany 89=183-189 Poni et al. (2009) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 60=397-407 Palliotti et al. (2012) "Morpho-anatomical and physiological response of Pinot Noir and Montepulciano grapevine leaves to mild and severe water stress" Vitis 51=109-116 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 56-94.
Mature Leaf
The mature leaf description of Cabernet Sauvignon compiled from the VIVC reference protocol documents a pentagonal blade with five distinct lobes (OIV 067:3, OIV 068:4), deeply incised upper lateral sinuses with parallel or slightly overlapping sides (OIV 070:7), a strongly overlapping petiolar sinus with the lobes nearly touching or crossed (OIV 079:7), medium-to-large leaf blade size (OIV 065:5-7 depending on vigor), deep green adaxial surface with moderate blistering (OIV 075:5), medium-length acute teeth of mixed convex-rectilinear form (OIV 076:3), medium density of prostrate hairs on the abaxial surface (OIV 084:5), and the distinctly red-pigmented main veins on the adaxial surface that is one of Cabernet Sauvignon's most recognizable leaf features — this comprehensive multi-character leaf profile, involving approximately 30 OIV descriptors, provides the highest-resolution ampelographic characterization available from any single organ.
An ampelographic description component that documents the morphological characteristics of fully expanded leaves from the middle third of normally growing shoots, observed between véraison and harvest (E-L stages 35-38) when leaf development is complete and environmental modification is minimized, recording the comprehensive set of blade, lobe, sinus, vein, tooth, and surface characters (OIV 065-094) that constitute the most information-rich single organ for cultivar identification in Vitis vinifera.
Mature_Leaf_Description addresses the organ that has been the primary focus of ampelographic description since the discipline's origins — the grapevine leaf, with its extraordinary morphological diversity across cultivars, provides more independently variable diagnostic characters than any other organ. The concentration of descriptors on the mature leaf (approximately 30 of the 130+ OIV descriptors) reflects both the leaf's morphological complexity and its practical accessibility — leaves are non-destructively observable, persistently available from véraison through leaf fall (a 2-3 month observation window), and produced in sufficient numbers to allow replicated sampling from multiple shoots: the standardization of observation on leaves from the middle third of normally growing shoots (typically the 8th-12th nodes) minimizes the positional variation that affects leaf morphology along the shoot axis, where basal leaves are typically smaller and less lobed than mid-shoot leaves. The mature leaf is also the organ where the interaction between cultivar genotype and growing conditions is most manageable — while leaf size varies substantially with vigor, the qualitative characters (blade shape, lobe number, sinus form, tooth shape, trichome type) are primarily genetically determined and provide reliable identification even under variable environmental conditions.
Reifblatt-Beschreibung
adult leaf characterization
adulte Blattmorphologie
ausgewachsenes Blatt-Charakterisierung
fully expanded leaf description
mature foliage morphology
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia UPOV (2011) Guidelines for the Conduct of Tests for Distinctness, Uniformity and Stability — Vine (Vitis L.), TG/50/9 Viala P and Vermorel V (1901-1910) Ampélographie, Masson
Mature Leaf Description
Wild V. vinifera ssp. sylvestris male plants during the bloom period (BBCH 61-69, spanning 10-14 days) exhibit mature male only flowers across a range of developmental states= pre-anthesis mature flowers (BBCH 59, calyptra swollen, anthers yellow, 2-3 days before shedding), fully developed flowers (BBCH 65, calyptra shed, anthers dehiscing, active pollen release), and post-anthesis mature flowers (BBCH 67-69, anthers empty or partially depleted, filaments browning, 2-4 days post-peak dispersal), with the extended maturity window ensuring pollen availability throughout the female plant receptivity period and maximizing fertilization success across variable bloom phenology among female plants in the population. Individual male plants bearing 800-1500 flowers exhibit asynchronous maturation creating continuous pollen production across 7-10 days, with 200-400 flowers in fully developed state on any given day during peak bloom.
A mature male only flower is a male only flower that has achieved full developmental maturity with completed organ differentiation and structural development, encompassing both pre-anthesis flowers with mature but still-enclosed stamens and post-anthesis flowers that have completed pollen release but retain their structural integrity, representing the broader category of developmentally complete male only flowers across the full maturity window. Mature male only flowers are characterized by= (1) completed stamen development with filaments achieving full length potential (3.5-4.5 mm) and anthers reaching full size (0.7-1.1 mm) with fully differentiated anther wall layers, tapetum, and mature pollen grains (2) completed pollen maturation with pollen grains achieving final developmental state including mature exine (pollen wall) with characteristic Vitis pollen surface ornamentation, cytoplasm fully loaded with starch reserves and proteins for germination and pollen tube growth, and mature generative nucleus and vegetative nucleus ready for fertilization (3) structural maturity of all flower components including mature calyptra ready for shedding (pre-anthesis) or already shed (post-anthesis), mature receptacle and nectary disk, and complete absence or maximum reduction of pistil structures (4) flower tissues achieving adult cellular characteristics with fully expanded cells, mature cell wall composition, and complete tissue differentiation. The 'mature' designation encompasses a broader developmental window than 'fully developed,' including= mature pre-anthesis flowers (1-3 days before calyptra shedding, with mature structures ready for anthesis but calyptra still attached and anthers still enclosed), fully developed anthetic flowers (at peak pollen dispersal with calyptra shed and anthers dehiscing), and mature post-anthesis flowers (1-5 days after peak pollen release, with stamens beginning senescence but structural integrity maintained). Mature male only flowers in the pre-anthesis state show swollen flower buds with calyptra beginning to separate from the receptacle base, visible as a slight gap indicating imminent calyptra shedding, and anthers visible through the translucent calyptra showing yellow coloration indicating mature pollen. Mature male only flowers in the post-anthesis state show empty or partially empty anthers that have released most pollen, filaments beginning to brown or desiccate as senescence proceeds, and overall flower structure beginning to deteriorate though not yet abscising. The mature state represents the full period of reproductive competence and structural integrity, typically spanning 6-10 days from achievement of maturity (2-3 days pre-anthesis) through completion of senescence (3-5 days post-anthesis), compared to the 1-2 day window of peak functional activity in fully developed flowers. Mature male only flowers exhibit pollen viability that varies across the maturity window= pre-anthesis mature flowers contain pollen at 70-85% viability (slightly lower than post-dehiscence peak due to incomplete final maturation), fully developed anthetic flowers contain pollen at peak 78-92% viability, and post-anthesis mature flowers contain residual pollen at declining 40-65% viability as released pollen ages. The mature developmental state is critical for understanding the full reproductive contribution of male only flowers, as pollen dispersal occurs across multiple days during the maturity window rather than in a single instant, and environmental conditions during the entire maturity period (not just at peak anthesis) affect overall pollen contribution to population fertilization. Mature male only flowers on dioecious male plants provide extended pollen availability across the maturity window, increasing probability of temporal overlap with pistillate flower receptivity and enhancing male reproductive success through bet-hedging across variable environmental conditions.
Mature_Male_Only_Flower represents the broader developmental category encompassing the full maturity window and is distinguished from Fully_Developed_Male_Only_Flower by broader temporal scope (pre-anthesis through post-anthesis versus peak anthesis only). The mature category is useful for describing the complete period of reproductive competence and for understanding extended pollen availability in dioecious populations where asynchronous flower maturation creates extended pollen production periods. The distinction between mature and fully developed may seem subtle but is important for phenological studies tracking bloom progression and for pollination ecology studies examining temporal dynamics of pollen availability and pistillate receptivity overlap. Understanding the full maturity window and its environmental modulation informs models of reproductive success in wild dioecious Vitis populations.
adult male flower
ausgereifte Staminatblüte
developmentally mature male flower
entwicklungsmäßig reife männliche Blüte
reife männliche Blüte
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 103-107 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 99-108 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 190-198.
Mature Male Only Flower
Syrah mature own-rooted trunk in Rhône Valley at 12 years post-planting shows 68-82 mm diameter with 12 annual rings averaging 2.5-3.5 mm width, sapwood 18-25 mm radial depth conducting 35-55 L/day during peak summer, heartwood comprising central 35-45 mm diameter with reduced conductivity, 110-140 g stored starch reserves, and consistent annual production of 4.5-6.5 kg fruit per vine (65-85 hL/ha at 5000 vines/ha density)
A mature own-rooted trunk is an own-rooted trunk that has completed architectural development and entered sustained productive phase (typically years 5-25 post-planting) characterized by stable annual growth patterns, optimal fruit production, efficient carbohydrate cycling, and established equilibrium between vegetative and reproductive development, representing the prime productive period for commercial viticulture. Mature own-rooted trunk development reflects= completion of training system architecture (permanent cordons or head positions established and stable), annual growth rings of relatively uniform width (1.5-3.5 mm radial increment per year compared to 3-6 mm in young trunks), progressive trunk diameter increase (from 40-70 mm at year 5 to 80-130 mm at year 20 for moderate-vigor cultivars), accumulation of substantial heartwood (central 30-60% of trunk diameter showing reduced water transport, darker coloration, potential antimicrobial compound accumulation), and bark thickening (4-8 mm thickness providing environmental protection). Mature own-rooted trunk anatomical characteristics include= well-developed secondary xylem with 5-20+ annual rings showing ring-porous anatomy (large earlywood vessels 120-280 μm diameter providing majority of water transport capacity), functional sapwood comprising outer 15-30 mm radius (conducting water while inner heartwood provides mechanical support), active vascular cambium producing consistent annual increments, secondary phloem (inner bark) 2-4 mm thick with functional sieve tubes in outer 0.5-1 mm, and protective periderm (outer bark) with cork layers reducing water loss and protecting against pathogens. Mature own-rooted trunk physiological characteristics include= stable annual carbohydrate cycling (accumulating 80-150 g starch during post-harvest period, mobilizing in spring for budbreak and early growth, rebuilding reserves during ripening and post-harvest), consistent water transport capacity (providing 20-80 L/day during peak transpiration depending on canopy size and environmental conditions), efficient nutrient transport (xylem delivering root-absorbed minerals, phloem redistributing nutrients between organs), and balanced source-sink relationships (photosynthesis supporting fruit development, vine growth, and reserve storage without excessive vigor or deficiency). Mature own-rooted trunk productivity characteristics include= stable annual yield (typically 3-8 kg fruit per vine depending on training system, spacing, and site, equivalent to 40-120 hL/ha at typical densities), consistent fruit quality (established vine physiology providing reliable berry composition), uniform phenology (budbreak, flowering, véraison, and harvest timing showing low year-to-year variation in absence of extreme weather), and balanced vegetative growth (shoot length and leaf area appropriate for crop load and trellis system without excessive vigor or deficiency). Mature own-rooted trunk management during productive phase includes= annual pruning maintaining architecture and crop load (removing 80-95% of annual growth leaving 15-40 buds per vine depending on system), canopy management optimizing fruit exposure and air circulation (shoot positioning, leaf removal, hedging as needed), disease management preventing trunk pathogens (protecting pruning wounds from Eutypa, Esca, Botryosphaeria infection), nutrition and irrigation maintaining productivity (balanced fertilization, deficit irrigation during ripening in many quality-focused systems), and monitoring for declining vigor or disease symptoms indicating beginning of senescence phase. Mature own-rooted trunk variation by cultivar and site includes= high-vigor cultivars or fertile sites maintaining larger diameter (100-150 mm by year 15-20), low-vigor cultivars or stressed sites showing smaller diameter (50-90 mm by year 15-20), optimal sites providing longest productive maturity phase (15-30+ years of stable production), and marginal sites showing shorter maturity (5-15 years before senescence begins).
Mature_Own_Rooted_Trunk emphasizes established productive phase and is distinguished from Young_Own_Rooted_Trunk (establishment and architecture development) and Old_Own_Rooted_Trunk (declining senescence phase) by stable optimal productivity and balanced physiology. Critical for understanding peak commercial productivity period, optimal management for sustained production, trunk disease impacts on mature vine health, and quality potential of established vines.
Established trunk
Etablierter Eigenwurzelstamm
Productive phase trunk
Produktionsphase-Stamm
Reifer wurzelechter Stamm
Mature Own Rooted Trunk
Cabernet Franc grafted on 3309 Couderc showing mature root hair zone on fourth-order fine root extending 24-52 mm from root apex, zone 28 mm length with hairs predominantly 4-6 days post-initiation, microscopic examination revealing hairs at final length 0.58 mm average with fully developed cytoplasm and intact membranes, FDA viability staining showing 78% bright fluorescence indicating high functional proportion, isotope tracing demonstrating mature zone providing 52% of total root uptake from 31% of root length, mature root hair zone representing peak functional region providing disproportionate absorption contribution
Mature root hair zone is root hair zone showing predominantly mature functional hairs typically 3-7 days post-initiation representing peak functional period, usually located in mid to distal portion of hair zone approximately 20-50 mm from root apex, characterized by hairs at final length having completed growth, showing optimal functional capacity with maximum absorption rates, demonstrating fully developed morphology and physiology, and representing the region where hair population provides maximum contribution to vine resource acquisition per individual hair and collectively, with mature root hair zone showing typical characteristics including fully elongated hairs 0.4-1.0 mm average length, high proportion 60-85% of hairs actively functional, dense cytoplasm and intact membranes enabling efficient transport, adequate turgor maintaining optimal geometry, and demonstrated peak uptake rates, and demonstrating that mature functional state distinguishes this zone from proximal zone with developing hairs or distal zone with senescing hairs. Mature root hair zone hair developmental state includes= showing growth completion (mature hairs reached final length and ceased tip growth indicating developmental maturity), exhibiting structural maturity (hairs showing fully developed morphology with optimal length-diameter ratio and cytoplasmic organization), demonstrating physiological maturity (hairs with fully functional metabolic and transport systems operating at or near maximum capacity), revealing optimal age (mature hairs typically 3-7 days old representing peak functional period in hair lifespan), and displaying stability (mature hairs maintaining relatively stable structure and function before senescence begins). Mature root hair zone functional capacity includes= showing peak absorption rates (mature hairs exhibiting maximum water and nutrient uptake per hair or per surface area), exhibiting optimal efficiency (mature hairs showing best ratio of uptake to maintenance cost during peak period), demonstrating full transport capacity (mature hair membranes and transporters operating at maximum efficiency), revealing responsive capability (mature hairs rapidly adjusting uptake in response to soil conditions or plant demand), and displaying collective performance (mature zone providing substantial fraction of total root uptake through combined mature hair population). Mature root hair zone population characteristics includes= showing high functional proportion (typically 60-85% of hairs in mature functional state actively contributing to uptake), exhibiting age homogeneity (mature zone showing relatively narrow age range around peak functional age 4-6 days), demonstrating density stability (mature zone maintaining relatively stable hair density as production and senescence balanced), revealing spatial continuity (mature zone forming continuous region rather than patchy distribution), and displaying optimal composition (age structure and functional state optimized for maximum zone-level performance). Mature root hair zone position and extent includes= located mid to distal hair zone (mature zone typically 20-50 mm from root apex depending on root growth rate and hair lifespan), exhibiting variable length (mature zone typically 15-35 mm extent depending on hair functional period duration), demonstrating stability relative to apex (mature zone position maintaining relatively constant distance from moving root tip), revealing overlap with dense zone (mature zone often overlapping substantially with dense hair zone in mid hair-zone region), and displaying order-dependent patterns (mature zone position varying with root order reflecting differences in growth rates and hair characteristics). Mature root hair zone absorption contribution includes= showing substantial uptake (mature zone providing large fraction potentially 40-60% of hair zone uptake from typically 40-60% of hair zone length), exhibiting peak efficiency (mature zone showing highest uptake per unit length or per unit surface area within hair zone), demonstrating critical function (mature zone absorption essential for meeting current vine resource demands), revealing nutrient-dependent patterns (mature zone particularly important for nutrients showing high uptake rates like nitrogen and potassium), and displaying stress sensitivity (mature zone function particularly vulnerable to drought or other stresses affecting vine status). Mature root hair zone soil relationships includes= showing intimate soil contact (mature hairs firmly established in soil pores and particle surfaces), exhibiting rhizosphere development (mature zone showing well-developed rhizosphere with mucilage and microbial activity), demonstrating depletion zones (mature hair uptake creating nutrient depletion zones around hairs in soil), revealing water extraction (mature zone extracting substantial soil water potentially drying local soil), and displaying mechanical stability (mature hairs anchored allowing continued function despite some soil disturbance). Mature root hair zone maintenance requirements includes= showing metabolic demand (mature hairs requiring continuous photosynthate supply supporting active transport and maintenance), exhibiting nutrient dependence (mature hair function requiring adequate nutrient availability particularly nitrogen for protein synthesis), demonstrating moisture needs (mature hairs requiring adequate soil moisture maintaining turgor and function), revealing temperature sensitivity (mature hair function optimal 15-25°C declining outside this range), and displaying carbon cost (mature hair maintenance requiring investment though cost low relative to function provided). Mature root hair zone in vine performance includes= determining resource acquisition (mature zone absorption capacity limiting vine nutrient and water status particularly under demand or supply limitations), enabling growth support (adequate mature zone function necessary for supporting vigorous shoot and root growth), demonstrating stress response (mature zone function critically affected by drought, compaction, or other root stresses), revealing management importance (practices affecting mature zone abundance or function directly impacting vine resource status and performance), and displaying rootstock effects (rootstock varieties differing in mature zone characteristics affecting resource acquisition efficiency and stress tolerance).
Mature_Root_Hair_Zone emphasizes mature functional state and peak performance period and is distinguished from developing or senescing zones by focus on fully functional mature hairs. Critical for understanding functional maturity, peak absorption, optimal age composition, substantial uptake contribution, soil relationships, and the region with predominantly mature functional hairs in peak operational state providing maximum absorption efficiency.
Functional hair zone
Funktionelle Haarregion
Optimal absorption zone
Optimale Absorptionszone
Peak-age hair zone
Reife Wurzelhaarzone
Mature Root Hair Zone
Cabernet Sauvignon mature seeds at harvest (BBCH 89) display uniform dark brown coloration, 7-8 mm length, hard lignified testa resistant to finger pressure, tannin polymerization index 65-70%, moisture content 30-35%, and separate cleanly from surrounding pulp tissue
A mature seed is a fully or substantially developed seed of Vitis vinifera that has completed embryogenesis, endosperm development, and seed coat maturation, typically characterized by lignified testa, viable embryo with differentiated cotyledons and radicle, and acquired desiccation tolerance. Mature seeds develop through completion of Phase III berry development (post-véraison lignification and desiccation, BBCH 85-89), achieving physiological maturity that enables germination after dormancy release. Seed maturity is assessed through multiple criteria= morphological indicators (seed coat color transition from green/tan to brown/black, hard lignified testa, fully formed beak and chalazal structures), biochemical markers (tannin polymerization, lipid accumulation in endosperm, reduction in moisture content from 60-70% during development to 30-40% at maturity), and physiological characteristics (acquired dormancy, desiccation tolerance, germination capacity after stratification). The timing of seed maturation relative to berry ripening varies by cultivar= in most wine grape varieties, seeds achieve physiological maturity 2-4 weeks before optimal berry harvest maturity, creating a developmental asynchrony important for harvest timing decisions. Mature seed tannins differ significantly from immature seed tannins= mature seeds contain higher proportions of polymerized, less astringent tannins, while immature seeds retain higher levels of monomeric catechins and epicatechins that contribute harsh astringency if extracted during vinification. Commercial viticulture monitors seed maturity through visual assessment (brown coloration) and tactile testing (ease of separation from pulp, resistance to crushing), using seed browning as one indicator of approaching harvest maturity for wine grape production. Seed maturity also influences seed viability for breeding programs and propagation research, with fully mature seeds showing 60-90% germination rates after appropriate stratification compared to 0-30% for immature seeds.
Mature_Seed emphasizes completion of developmental program and achievement of physiological maturity and is distinguished from Fully_Developed_Mature_Seed (optimal complete maturation) and Viable_Mature_Seed (functional germination capacity with possible incomplete maturation). Critical for harvest timing assessment, wine quality prediction based on tannin extractability, and breeding program seed viability evaluation.
Ausgereifter Traubenkern
Brauner Kern
Brown seed
Reifer Samen
Ripe seed
Kennedy, J. A., Matthews, M. A., & Waterhouse, A. L. (2002). Effect of maturity and vine water status on grape skin and wine flavonoids. American Journal of Enology and Viticulture, 53(4), 268-274. | Ristic, R., & Iland, P. G. (2005). Relationships between seed and berry development of Vitis vinifera L. cv Shiraz= Developmental changes in seed morphology and phenolic composition. Australian Journal of Grape and Wine Research, 11(1), 43-58.
Mature Seed
A 20-year-old Cabernet Sauvignon vine bears a Mature_Vine_Role as the productive heart of a Napa Valley estate vineyard, its deep-rooted stability producing 4-5 tonnes/hectare of consistently high-quality fruit for the estate's flagship wine. A 15-year-old Riesling vine serves in a Mature_Vine_Role in steep Mosel terracing, the mature root system now fully exploiting the slate substrate to produce the mineral-driven fruit characteristic of great Mosel Riesling. A 25-year-old Sangiovese vine on Tuscan galestro bears a Mature_Vine_Role, its decades of site adaptation producing the fruit concentration and terroir expression expected for Brunello di Montalcino classification.
A vine age role that inheres in a grapevine that has reached full productive maturity, typically achieved by years 10-15 after planting, characterized by stable vine architecture, optimized root-to-shoot balance, consistent yield capacity, and fruit quality expressing the full potential of the variety-site-management combination. Mature vines represent the productive core of commercial viticulture, with physiological equilibrium enabling predictable annual performance under normal conditions and maximum expression of terroir characteristics. The mature phase may span 20-40+ years depending on variety, rootstock, site conditions, and management quality, with well-maintained vines on favorable sites remaining productively mature for decades. Mature vine physiology features extensive root systems with deep soil exploration, efficient carbon allocation balancing vegetative and reproductive demands, and stable spur or renewal positions supporting consistent annual production. Fruit from mature vines typically commands higher regard than young vine fruit, with the physiological stability and site integration reflected in more complex, site-characteristic wines. The transition from mature to old vine status is gradual rather than abrupt, with declining yields and changing vine balance signaling the shift toward old vine physiology.
Mature_Vine_Role captures the extended productive prime when vines achieve their full potential, representing the viticultural ideal that vineyard investment aims to create. The duration of the mature phase varies enormously—some variety-site combinations maintain mature productivity for 50+ years while others begin declining by year 20-25—making the transition to old vine status vineyard-specific rather than universally age-defined. Mature vines require less intensive per-vine management than young vines but benefit from consistent annual practices (appropriate pruning, disease control, canopy management) that maintain the vine balance underlying stable production. The distinction between mature and old vine status is often as much about marketing as physiology, with the "old vine" designation applied whenever it enhances market positioning regardless of whether the vine has actually transitioned to old vine physiology.
Vollertrag-Rebe
ausgereifte Rebe
established mature vine
full-production vine
mature vine
prime vine
produktive Rebe
reife Rebe
Keller M (2015) The Science of Grapevines, Academic Press Vaudour E et al. (2015) Vine age effect on terroir expression, OENO One Bonada M et al. (2020) Vine age and grape composition, Australian Journal of Grape and Wine Research 26: 139-152
Mature Vine Role
Maturity index quality comparison evaluating different indices for Shiraz harvest timing reveals the predictive utility variation among approaches, with simple Brix/TA ratio reaching target value (3.8) at day 42 post-véraison (providing basic balance indication but missing phenolic dimension), while Glories extractability index reached optimal (>60%) at day 48 (adding phenolic maturity information), and comprehensive sensory-analytical correlation analysis identified day 52 as optimal (integrating all quality-determining factors), demonstrating how different indices capture different aspects of maturity with implications for their appropriate application in harvest decisions.
Maturity index quality is a maturity quality that represents the quantitative integration of multiple maturity parameters into calculated indices designed to capture overall ripeness status or balance among compositional elements, characterizing the mathematical relationships among individual parameters that provide more informative assessment of harvest readiness than any single measurement alone. Within the BFO framework, this quality specifically addresses the integrative dimension of maturity assessment, distinguishing calculated indices from individual parameter measurements while recognizing that maturity indices attempt to reduce the complexity of multi-dimensional maturity into actionable metrics that correlate with wine quality outcomes better than single parameters. The ontological significance lies in maturity indices providing decision-support tools that capture the interactions among maturity dimensions, where simple ratios (sugar/acid) identify basic balance, more complex indices (phenolic maturity indices) integrate physiological and technological parameters, and comprehensive systems (maturity potential indices) attempt to predict wine quality potential from grape composition, with each index type serving different assessment purposes and operating at different levels of integrative complexity. The design principles underlying maturity indices include the combination of positively and negatively correlated parameters (capturing balance rather than simple accumulation), the weighting of parameters according to their quality relevance (emphasizing factors most predictive of wine outcomes), and the scaling of indices to interpretable ranges (enabling threshold-based decision-making), with successful indices demonstrating empirical correlation with wine quality across vintages and sites. The limitations of maturity indices include their simplification of complex biological systems (potentially missing important quality-determining factors not included in the calculation), their context-dependency (indices calibrated in one region potentially requiring adjustment for different cultivars or climates), and their assumption of consistent relationships (actual parameter interactions potentially varying with conditions), requiring critical interpretation rather than mechanical application of index values. Understanding maturity index quality enables appropriate selection among available indices for specific assessment purposes, interpretation of index values in their proper context, and recognition of both the utility and limitations of quantitative approaches to the inherently multi-dimensional challenge of maturity assessment.
Maturity_Index_Quality serves as the parent class for specific maturity indices (Brix/Acid Ratio, Glories Index, Maturity Potential Index). This quality addresses the quantitative integration of maturity parameters into decision-support metrics. The class supports harvest decision-making through index characterization enabling multi-parameter assessment.
Ernteindex
Kompositionsindex
Reifeindex
Reifekennzahl
compositional index
harvest index
maturity indicator
ripeness index
Ribéreau-Gayon P et al. (2006) Handbook of Enology, Volume 1: The Microbiology of Wine and Vinifications, 2nd Ed. John Wiley & Sons, Chichester, pp. 193-230 Iland P et al. (2011) The Grapevine: From the Science to the Practice of Growing Vines for Wine. Patrick Iland Wine Promotions, Adelaide, pp. 85-110
Maturity Index Quality
Maturity monitoring precision quality assessment for a Marlborough Sauvignon Blanc operation evaluated analytical performance across the laboratory and field methods employed for harvest decisions. Handheld digital refractometer Brix measurements achieved maturity monitoring precision quality of ±0.3°Brix (CV 1.4%) under field conditions with well-trained operators, adequate for detecting the 1.0°Brix increments relevant to harvest window progression. Laboratory density-based Brix determination improved precision to ±0.1°Brix (CV 0.5%), providing the resolution for fine-scale harvest optimization distinguishing 0.5°Brix categories. Titratable acidity analysis showed precision of ±0.25 g/L (CV 3.2%), sufficient for monitoring the acid decline relevant to Sauvignon Blanc freshness preservation. Methoxypyrazine analysis by GCMS exhibited precision of ±2 ng/L (CV 12%) at typical harvest concentrations, enabling meaningful tracking of green character decline but requiring interpretation acknowledging substantial measurement uncertainty. The precision hierarchy informed protocol selection, with field refractometry supporting routine monitoring while laboratory density measurement was reserved for final harvest decisions where precision advantages justified additional effort.
Maturity monitoring precision quality is a harvest timing indicators quality that inheres in a grapevine cultivar or clone within a specific production context and represents the accuracy and reproducibility of analytical measurements applied to assess ripening status, encompassing the precision of laboratory and field instruments, the consistency of analytical protocols across operators and time points, and the resolution with which measured parameters can distinguish meaningful ripeness differences relevant to harvest timing decisions. This quality determines the minimum detectable difference in ripening status achievable through monitoring programs, establishing the granularity at which harvest timing can be optimized based on measured data rather than requiring coarser decisions limited by measurement uncertainty. The ontological significance of maturity monitoring precision quality lies in its role establishing the practical limits of indicator-based harvest optimization, wherein high monitoring precision enables fine-scale timing adjustments capturing incremental quality improvements while poor precision restricts decisions to coarse categories that may miss optimal timing windows spanning brief periods within measurement uncertainty ranges. The biological basis of maturity monitoring precision involves the inherent variability of the parameters being measured, which reflects both true compositional heterogeneity and measurement error the stability of samples during analysis that may introduce variance from post-collection change and the calibration consistency of instruments and protocols that determines whether measurements are comparable across time points within a ripening progression. Key parameters for harvest timing monitoring include sugar concentration measured by refractometry, density, or chemical analysis with typical precision of ±0.2°Brix for well-calibrated instruments titratable acidity with precision of ±0.3 g/L pH with precision of ±0.02 units and phenolic parameters with precision varying widely depending on methodology from ±5% for spectrophotometric measures to ±15% or more for complex extraction-based protocols. Assessment of maturity monitoring precision quality involves replicated analysis of reference samples to quantify analytical reproducibility, comparison of results across operators and instruments to establish method consistency, and evaluation of whether achievable precision supports the decision granularity required for specific harvest timing objectives.
Maturity_Monitoring_Precision_Quality is_a Harvest_Timing_Indicators_Quality, representing the accuracy and reproducibility of analytical measurements for ripening assessment. This class addresses measurement quality rather than sampling quality or indicator selection. Precision requirements depend on the decision granularity needed for specific harvest timing objectives. The hierarchy of field versus laboratory precision informs appropriate method selection for different monitoring purposes. Critical for meaningful ripening trend analysis supporting harvest optimization.
Monitoring-Methodenpräzision
Reifegradüberwachungs-Präzision
Reifemessungs-Reproduzierbarkeit
analytical precision
analytische Genauigkeit
maturity assessment reproducibility
monitoring method precision
ripeness measurement accuracy
Iland P et al. (2011) Chemical Analysis of Grapes and Wine: Techniques and Concepts, 2nd Edition. Patrick Iland Wine Promotions, Adelaide Ristic R et al. (2007) Influence of vineyard floor management practices on grapevine vegetative growth, yield, and fruit composition. American Journal of Enology and Viticulture 58:25-31 Cynkar W et al. (2004) Effect of wine matrix composition on the measurement of mid-infrared spectra using a Fourier transform infrared spectrometer. Australian Journal of Grape and Wine Research 10:130-137
Maturity Monitoring Precision Quality
Maturity potential index application comparing vineyard blocks for premium Cabernet Sauvignon allocation reveals the comprehensive assessment utility, with Block A showing technological maturity index of 85/100 (favorable Brix/pH/TA) but phenolic potential index only 62/100 (limited anthocyanin extractability, high seed tannin contribution) yielding combined maturity potential of 73, while Block B shows technological index 78/100 (slightly lower sugar) but phenolic potential 88/100 (high extractability, low seed contribution) yielding combined potential of 83, with wines confirming that Block B's apparently lower sugar fruit produced superior wine quality due to superior phenolic development, demonstrating how comprehensive indices identify quality potential that single parameters miss.
Maturity potential index quality is a maturity index quality that represents comprehensive indices designed to predict wine quality potential from grape compositional analysis by integrating multiple technological and phenolic parameters into a single score or rating, characterizing the overall winemaking potential assessment that attempts to predict wine outcomes from grape inputs across the full range of quality-determining factors. Within the BFO framework, this quality specifically addresses the quality prediction dimension of maturity indices, distinguishing comprehensive predictive indices from simpler balance ratios or single-dimension extractability measures while recognizing that maturity potential indices attempt the ambitious goal of predicting the complex, multi-factorial wine quality outcome from pre-fermentation grape analysis. The ontological significance lies in maturity potential indices attempting to solve the fundamental quality prediction challenge of viticulture, where the relationship between grape composition and wine quality involves numerous interacting factors (sugars, acids, phenolics, aromatics, plus winemaking variables) that comprehensive indices integrate through empirically derived weightings reflecting the relative quality impact of each parameter. The construction approaches for maturity potential indices include statistical modeling (regression equations predicting wine quality scores from grape parameters), expert system approaches (rules integrating multiple parameters based on enological knowledge), and multi-criteria scoring systems (point assignments for parameters in defined ranges summed to total scores), with each approach having strengths and limitations for different applications. The parameters typically incorporated in comprehensive maturity potential indices include technological parameters (Brix, pH, TA establishing fermentation substrate quality), phenolic indices (total phenolics, extractability, tannin maturity affecting structure potential), and potentially aromatic indices (precursor content, varietal compound development affecting sensory potential), with parameter selection and weighting based on empirical relationships with wine quality in the calibration dataset. The limitations and appropriate use of maturity potential indices include their calibration-dependency (indices developed in one context potentially requiring recalibration for different cultivars, regions, or wine styles), their inability to capture winemaking effects (same grape potential yielding different wine quality depending on winemaking decisions), and their assumption of consistent relationships (climate change potentially altering historical parameter-quality relationships), requiring indices to be used as decision-support tools rather than deterministic predictors. Understanding maturity potential index quality enables appropriate application of comprehensive indices for quality potential assessment, recognition of their predictive utility and limitations, and critical interpretation that considers the context-dependency of any quality prediction approach.
Maturity_Potential_Index_Quality addresses comprehensive indices attempting to predict wine quality from integrated grape analysis. This quality recognizes both the utility and limitations of quality prediction from pre-fermentation parameters. The class supports quality-focused harvest decisions through comprehensive index characterization with appropriate interpretation of context-dependency.
Qualitätspotentialindex
Reifepotentialindex
Weinpotentialscore
comprehensive maturity index
integrated quality index
quality potential index
umfassender Reifeindex
wine potential score
Iland P et al. (2011) The Grapevine: From the Science to the Practice of Growing Vines for Wine. Patrick Iland Wine Promotions, Adelaide, pp. 85-110 Kontoudakis N et al. (2010) Influence of wine pH on changes in color and polyphenol composition induced by micro-oxygenation. Journal of Agricultural and Food Chemistry 58(2): 1301-1311
Maturity Potential Index Quality
Maturity quality assessment for premium Cabernet Sauvignon production reveals the multidimensional nature of harvest decisions, with sugar maturity reaching target at 24.5 Brix by September 20, acid maturity achieving desired pH 3.55 by September 25, but phenolic maturity (tannin polymerization, anthocyanin stability, seed browning) not optimal until October 5, creating a 15-day window during which different maturity dimensions achieve their targets, requiring the winemaker to prioritize among potentially conflicting optima based on wine style objectives and weather risk, demonstrating how maturity quality encompasses multiple independent developmental dimensions requiring integrated assessment.
Maturity quality is a phenological quality that represents the developmental readiness of grape berries for harvest and subsequent utilization, characterizing the complex of physiological states and compositional attributes that together determine whether fruit has achieved the condition appropriate for its intended use, whether fresh consumption, drying, juice production, or winemaking with specific quality objectives. Within the BFO framework, this quality addresses the harvest-readiness dimension of berry development, distinguishing the achieved developmental state from the rate or completeness of development while recognizing that maturity encompasses multiple independent dimensions (sugar accumulation, acid metabolism, phenolic development, flavor evolution) that may progress at different rates and achieve their optima at different times, creating the fundamental challenge of harvest timing decisions. The ontological significance lies in maturity quality determining the raw material quality available for transformation into wine, where the developmental state at harvest establishes the compositional foundation (sugars, acids, phenolics, aroma precursors) that winemaking can modulate but not fundamentally alter, making maturity assessment and harvest timing among the most consequential decisions in quality wine production. The multidimensional nature of maturity creates the distinction between physiological maturity (biological completion of seed, pulp, and skin development) and technological maturity (achievement of compositional targets for winemaking), with these dimensions not necessarily coinciding and their relative priority depending on wine style objectives and the tolerance for their potential misalignment. The environmental and management factors affecting maturity achievement include temperature during ripening (affecting rate and pathway of maturation), water status (influencing berry size and concentration), crop load (affecting resource availability for maturation), and canopy management (determining fruit exposure and microclimate), with these factors creating the vintage variation in maturity patterns that distinguishes exceptional from ordinary years. The climate change implications for maturity are substantial, as warming temperatures accelerate sugar accumulation while potentially decoupling sugar from phenolic and flavor maturity, creating challenges for achieving balanced maturity that traditional cultivar-climate relationships had optimized over centuries. Understanding maturity quality enables integrated assessment of harvest readiness, optimization of harvest timing for specific quality objectives, and appreciation of the multiple maturity dimensions that must be balanced for quality wine production.
Maturity_Quality serves as the parent class for Physiological and Technological maturity dimensions. This quality addresses the multidimensional harvest-readiness assessment critical for wine quality. The class supports harvest decision-making through integrated maturity characterization across compositional and developmental dimensions.
Beerenreife
Reifequaliät
Traubenreife
berry maturity
grape ripeness
harvest maturity
ripeness
Coombe BG, McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6(2): 131-135 Ribéreau-Gayon P et al. (2006) Handbook of Enology, Volume 1: The Microbiology of Wine and Vinifications, 2nd Ed. John Wiley & Sons, Chichester, pp. 193-230
Maturity Quality
Maturity spread assessment at target harvest date in Cabernet Sauvignon reveals the quality implications of developmental asynchrony, with high-synchrony block showing narrow maturity spread (Brix range 23.5-25.2, pH range 3.42-3.58, berry anthocyanins 1.4-1.8 mg/g, enabling harvest of consistent fruit), while low-synchrony block shows wide spread (Brix range 21.0-27.5, pH range 3.28-3.82, anthocyanins 0.8-2.4 mg/g, with distinct under-ripe and over-ripe subpopulations), demonstrating how maturity spread determines the uniformity achievable at harvest with direct implications for wine quality potential.
Maturity spread quality is a temporal spread quality that represents the range of maturity levels present in a fruit population at any given time point, characterizing the compositional heterogeneity (sugar, acid, phenolic, aromatic variation) that arises from developmental asynchrony and determines the degree of uniformity achievable in harvested fruit and the magnitude of quality compromise inherent in single-date harvest of non-uniform populations. Within the BFO framework, this quality specifically addresses the maturity dimension of temporal spread, distinguishing compositional heterogeneity at harvest from the developmental timing variation that created it while recognizing that maturity spread represents the quality-relevant outcome of all preceding developmental synchrony patterns, translating timing heterogeneity into the fruit quality variation that affects wine outcomes. The ontological significance lies in maturity spread determining the quality potential achievable through harvest, where narrow spread enables harvest of uniform fruit at target maturity producing consistent wine quality, while wide spread forces compromise among harvesting under-ripe, over-ripe, and optimally ripe fractions together, diluting quality potential through compositional heterogeneity that blending cannot fully resolve. The compositional dimensions of maturity spread include sugar spread (Brix variation indicating different fermentation potential and alcohol outcomes), acid spread (pH and TA variation affecting wine acidity balance), phenolic spread (color and tannin variation affecting wine structure consistency), and aromatic spread (flavor compound variation affecting wine complexity and typicity), with each dimension contributing to overall quality heterogeneity depending on the magnitude of spread and the wine style sensitivity to that parameter. The quantification of maturity spread involves measuring compositional parameters across berry or cluster populations, calculating dispersion metrics (range, standard deviation, CV), and characterizing the distribution shape (normal versus skewed, presence of distinct subpopulations), with these metrics enabling comparison among blocks, vintages, and management treatments and providing the basis for harvest timing and sorting decisions. Understanding maturity spread quality enables assessment of fruit uniformity achievable at any harvest date, prediction of wine quality implications from harvesting heterogeneous populations, and evaluation of sorting requirements or multiple-harvest strategies to manage spread-induced quality variation.
Maturity_Spread_Quality addresses the compositional heterogeneity present in fruit populations at harvest-relevant time points. This quality represents the quality-relevant outcome of developmental asynchrony. The class supports harvest decisions and quality assessment through maturity spread characterization.
Ernteuniformität
Kompositionsvariation
Reifeheterogenität
Reifespreizung
compositional variation
harvest uniformity
maturity range
ripeness heterogeneity
Kontoudakis N et al. (2011) Influence of the heterogeneity of grape phenolic maturity on wine composition and quality. Food Chemistry 124(3): 767-774 Bramley RGV (2005) Understanding variability in winegrape production systems 2. Within vineyard variation in quality over several vintages. Australian Journal of Grape and Wine Research 11(1): 33-42
Maturity Spread Quality
Merlot hermaphrodite flowers at 12-36 hours post-calyptra shedding (BBCH 61, very early flowering) exhibit maximally receptive stigmas with maximum wet exudate creating extremely glistening appearance, exudate sugar concentration 13.5-15.8% (peak levels), supporting 86-92% germination of self-pollen applied in controlled pollinations, representing the absolute peak receptivity before gradual transition to sustained fully receptive state (days 2-4, 78-86% germination) and eventual declining receptivity (days 5-6, 55-72% germination). Syrah stigmas sampled at 18-hour intervals from calyptra shedding through 7 days post-shedding show maximum exudate production rate at 6-30 hours post-shedding (maximally receptive period) with 8-12 μL exudate per stigma, declining to 5-8 μL at 2-4 days (fully receptive), 2-4 μL at 5-6 days (declining receptive), and <1 μL at 7+ days (post-receptive), with pollen germination rates tracking exudate production= 88-91% at maximum, 78-84% at fully receptive, 52-68% at declining, <15% post-receptive. Controlled crossing programs targeting maximum fertilization efficiency collect hermaphrodite anthers at pre-dehiscence stage, extract pollen, and apply to stigmas at 12-30 hours post-calyptra shedding (maximally receptive period) rather than at 2-4 days post-shedding (fully receptive), achieving 12-18% higher fruit set from crosses (72-84% versus 60-72%) demonstrating practical advantage of targeting maximally receptive stigma timing for critical controlled pollinations. Gewürztraminer functionally female flowers show maximally receptive stigmas at days 1-2 post-calyptra shedding with exudate sugar concentration 12.8-15.2% and cross-pollen germination 82-88%, providing peak window for capturing hermaphrodite pollinizer pollen when pollinizer bloom synchrony is good, with the maximally receptive period representing the optimal time for successful cross-pollination before transition to extended fully receptive period (days 3-6, still highly receptive at 72-82% germination but past absolute peak).
A maximally receptive stigma is a receptive stigma at the absolute peak of receptivity exhibiting maximum wet exudate production and achieving the highest pollen germination rates possible for the genotype, representing the optimal brief period (typically 12-48 hours) when stigma biochemical and cellular conditions are at their zenith and when pollination success probability is maximized. Maximally receptive stigmas are characterized by= (1) maximum wet exudate production with abundant secretion visibly coating and sometimes pooling on papillae, creating maximum glistening wet appearance (2) exudate biochemical composition at peak optimal levels with sugar concentration often 12-16% (upper range for Vitis), protein concentration 1.2-1.6 mg/ml, enzyme activities at maximum, and full complement of signaling molecules at peak concentrations (3) papillae at maximum turgor and expansion with cells fully hydrated and metabolically most active (4) pollen germination support at absolute maximum efficiency= 82-92% germination for hermaphrodite stigmas, 78-90% for functionally female/pistillate stigmas, representing the highest germination rates achievable for the flower type (5) pollen tube initiation occurring most rapidly (within 1.5-4 hours of pollen landing versus 2-6 hours on fully receptive stigmas). Maximally receptive stigmas represent a transient peak state occurring typically at days 1-2 post-calyptra shedding in most hermaphrodite flowers, coinciding with or slightly preceding peak anther dehiscence creating optimal temporal coordination between maximum female receptivity and maximum male pollen release, or occurring at days 1-3 in functionally female/pistillate flowers providing peak receptivity window for capturing initial cross-pollination from hermaphrodite or staminate sources. Maximally receptive stigmas show the brief period of absolute optimal conditions before gradual transition to fully receptive (still very high quality but declining from absolute peak) and eventually to declining receptive states. The maximally receptive state is most evident in flowers examined immediately after calyptra shedding (0-24 hours post-shedding) when exudate production has just activated and is at maximum rate before any desiccation or depletion has occurred, creating the wettest appearance and most favorable biochemical environment. Maximally receptive stigmas represent the optimal target for time-sensitive controlled crossing operations where maximum fertilization success is critical, such as crossing with limited pollen supplies, crossing with marginally compatible parents, or crossing under suboptimal environmental conditions where enhanced stigma receptivity can partially compensate for reduced pollen viability or environmental stress effects. The distinction between maximally receptive and fully receptive stigmas is subtle and primarily quantitative rather than qualitative, with maximally receptive representing the very peak (absolute highest germination rates, wettest appearance, peak exudate composition) while fully receptive represents the sustained high-quality period (still very high germination rates, wet appearance, optimal exudate) that persists for several days, with maximally receptive being a subset or brief initial phase of the fully receptive period.
Maximally_Receptive_Stigma represents the absolute peak receptivity state and overlaps substantially with Fully_Receptive_Stigma, with maximally receptive emphasizing the transient peak (12-48 hours) while fully receptive encompasses the sustained optimal period (2-5 days). The distinction is subtle and primarily relevant for time-sensitive controlled crossing operations or for detailed receptivity dynamics studies. Maximally receptive stigmas provide the highest probability of fertilization success and are the optimal target when maximum efficiency is required, though the practical difference versus fully receptive stigmas is modest (typically 5-15% higher germination/fruit set) making fully receptive stigmas acceptable for most applications. Understanding maximally receptive stigma timing and characteristics is most relevant for breeding programs and research applications requiring optimal pollination efficiency.
absolut empfangsfähige Narbe
absolutely optimal stigma
höchst empfangsbereites Stigma
maximal rezeptive Narbe
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 104-106 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 96-98.
Maximally Receptive Stigma
Pinot Noir in Burgundy shows maximum berry enlargement during early-mid August Year N+1 at BBCH 77-79, with cells reaching 92-96% final size at 88-108 micrometers diameter, berry achieving 94-97% final dimensions at 9.2-11.4 mm diameter and 3.8-5.6 g fresh weight, expansion rate declining from 145 to 42 mg per day, wall plasticity decreasing, osmotic potential plateau at -1.3 MPa, under continental conditions of 22-28°C temperature, typical of this early-to-mid season cultivar completing enlargement
Maximum berry enlargement represents the genetic size limit approach phase during berry cell enlargement when cells reach near-maximal dimensions 90-95% final size, berry achieves peak growth phase size 90-98% mature dimensions, expansion rate declines from peak levels as cells approach size constraints, and growth deceleration initiates transition toward lag phase, occurring during late enlargement weeks 6-8 post-fertilization as intensive expansion concludes. This size maximum approach phase occurs during late summer in early-mid August in temperate regions Year N+1, spanning approximately 1-2 weeks representing enlargement culmination, corresponding to BBCH 77-79 bunch closure to majority berries touching stages. Maximum enlargement mechanism involves cell size approaching genetic maximum determined by developmental programming, berry size peak representing culmination of cell number and cell expansion multiplicative effects, expansion rate decline from reduced wall yielding and approaching size limits, metabolic transition initiating shift from growth to lag phase metabolism, and overall growth deceleration marking enlargement phase conclusion. Cell size maximum characteristics show cells reaching 85-95% final mature dimensions, typical diameters 75-190 micrometers depending on cultivar and cell type, vacuoles occupying 85-92% cell volume at near-maximum, cytoplasm as thin 1.5-4 micrometer peripheral layer, and cell architecture approaching mature configuration. Cell size genetic limits show cultivar-specific maximum cell sizes determined by genetic programming, small-berried cultivars reaching 60-100 micrometer maxima, large-berried cultivars achieving 120-200 micrometer maxima, and genetic control as primary size determinant. Cell expansion cessation approaching shows expansion rate progressively declining as cells near maxima, eventual expansion cessation establishing final cell dimensions, cells retaining slight expansion capacity during lag and ripening phases adding final 5-15% size, and near-cessation marking enlargement completion. Berry size peak characteristics show berry diameter reaching 90-98% final mature size, typical dimensions 22-40 mm depending on cultivar at enlargement completion, fresh weight 9-25 g representing near-final mass, and berry approaching genetic size potential. Berry size genetic potential shows cultivar characteristic size ranges determined by cell number and cell expansion capacities, small-berried cultivars 6-10 mm final diameter, medium-berried 10-14 mm, large-berried 14-20 mm, and genetic programming establishing size ranges. Berry size achievement shows typical 90-98% final size reached by maximum enlargement completion, remaining 2-10% size gain during lag and ripening from continued limited expansion and ripening accumulation, and enlargement phase as major size determinant accounting for 85-95% final dimensions. Expansion rate decline characteristics show growth rate decreasing from peak 100-250 mg per day to declining 30-80 mg per day during maximum enlargement, progressive deceleration over 1-2 weeks, and rate decline marking transition from intensive to minimal growth. Expansion rate decline mechanisms show reduced wall yielding as walls become less plastic approaching size limits, osmotic gradient plateau with solute accumulation reaching steady state, turgor maintenance but reduced expansion effectiveness, cell approaching volume limits constraining further growth, and developmental programming directing growth cessation. Wall plasticity reduction shows wall extensibility declining as cells approach size limits, reduced expansin activity and wall modification, wall becoming less yielding under turgor, and wall properties shift from growth-permissive to growth-restrictive. Wall mechanical properties changes show increased wall stiffness from accumulated structural modifications, yield threshold increasing requiring higher turgor for expansion, reduced stress relaxation limiting plastic deformation, and mechanical changes limiting continued expansion. Osmotic potential plateau shows solute accumulation approaching steady state, osmotic potential reaching -1.2 to -1.6 MPa at maximum, minimal further osmotic potential decline, gradient approaching equilibrium, and plateau limiting additional water uptake driving force. Turgor pressure maintenance shows turgor sustained 0.6-1.0 MPa during maximum enlargement, pressure maintained through continued solute accumulation, but reduced expansion from wall stiffening, and turgor-wall relationship shifted toward minimal growth. Cell volume constraints show cells approaching physical and genetic size limits, further volume increase increasingly difficult, developmental signals possibly limiting expansion, and approaching constraints as growth cessation mechanism. Berry growth rate during maximum enlargement shows declining rate 30-80 mg fresh weight per day, progressive deceleration from peak levels, growth continuing but at reduced intensity, and declining rate characteristic of enlargement conclusion. Berry composition during maximum enlargement shows water content 80-85% fresh weight maintained, organic acids at peak or slightly declining 1.2-1.6%, sugars still low to moderate 6-10%, pH 2.8-3.0, and hard acidic green berry approaching lag phase composition. Berry morphology shows berry at near-final dimensions, smooth surface maintained, green color persisting, firm texture from turgor and acids, and overall appearance as mature-sized green berry. Cluster architecture shows berries tightly touching within clusters, bunch closure achieved with berries filling available space, cluster compactness at maximum, and architecture at peak density. Environmental effects on maximum enlargement include water availability affecting final size achievement with continued adequate water important though deficit effects less severe than during peak expansion, temperature affecting metabolic transition timing, and overall conditions determining final size attainment. Water effects show continued water supply important for final size, deficit during maximum enlargement potentially limiting final 5-10% size gain, and water less critical than during peak expansion though still important. Temperature effects show warm conditions potentially accelerating transition to lag phase, cool conditions extending maximum enlargement period, and temperature affecting transition timing. Maximum enlargement completion indicators show expansion rate declining to <20-30 mg per day, berry size approaching 95-98% final dimensions, metabolic markers shifting toward lag phase, and completion establishing transition to growth quiescence. The process shows three distinguishable aspects including cell size maximum with cells approaching genetic limits, berry size peak achieving near-final dimensions, and expansion rate decline marking growth deceleration. Duration typically 1-2 weeks representing enlargement culmination weeks 6-8. Regional timing shows maximum berry enlargement during early-mid August across climates as intensive expansion concludes. Viticultural significance includes representing culmination of berry size establishment determining final yield, marking transition from growth to lag phase, demonstrating approach to genetic size potential, and establishing near-final berry dimensions affecting harvest and quality. Research applications include analyzing size limits and genetic controls, investigating expansion cessation mechanisms, evaluating environmental effects on final size achievement, quantifying size determination and limitations, and assessing maximum enlargement as final size determinant.
Maximum_Berry_Enlargement represents genetic size limit approach within Berry_Cell_Enlargement_Phase. Subdivided into Cell_Size_Maximum with cells at 90-95% final dimensions, Berry_Size_Peak achieving 90-98% final size, and Expansion_Rate_Decline marking deceleration. Duration 1-2 weeks representing weeks 6-8. Wall plasticity decreases, osmotic potential plateaus. Growth rate declines from peak to 30-80 mg/day. Transition toward lag phase begins. Precedes Enlargement_Phase_Completion. Determines final berry size.
Größenmaximum-Annäherung
Spitzen-Vergrößerung
Wachstums-Kulmination
growth culmination
maximale Beerenvergrößerung
maximum size achievement
peak enlargement
size maximum approach
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 448-453 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Maximum Berry Enlargement
Syrah in Rhône Valley shows maximum cell number generation during mid July Year N+1 at BBCH 72-73, with exponential cell accumulation from 280,000 to 2.0-2.6 million cells over weeks 2-3 post-fertilization, peak generation rate 65,000-110,000 cells per day, doubling time 10-11 hours, generating 68-73% final cell number during 2-week period, under warm Mediterranean conditions of 25-30°C temperature, typical of this mid-to-late season cultivar establishing cellularity determining 12-14 mm final berry size
Maximum cell number generation represents the peak proliferative output phase during peak division activity when highest rate of new cell production occurs, exponential cell accumulation reaches maximum velocity, and majority 60-75% of final berry cell number generates during brief 1.5-2.5 week period, establishing bulk cellularity determining berry size potential. This maximum generation phase occurs during mid summer in early-mid July in temperate regions Year N+1, typically weeks 2-3 post-fertilization during peak division intensity, corresponding to BBCH 72-73 young fruit rapid growth stages. Maximum generation mechanism involves synchronized rapid cell division across large mesocarp cell populations, minimal 10-12 hour cell cycle times enabling 2-2.4 divisions per day, exponential accumulation from repeated division rounds, coordinated proliferation across pericarp tissues, and maximum generation rate producing thousands of new cells per hour per berry. Cell number accumulation dynamics show typical progression from 200,000-500,000 cells at maximum generation onset to 1.2-3.5 million cells at completion, accumulation rate 50,000-150,000 cells per day during peak period, doubling time 10-12 hours matching minimal cell cycle time, and exponential growth pattern characteristic of maximum generation. Generation rate quantification shows peak rate typically 40,000-120,000 new cells generated per berry per day, rate representing combined output of all dividing cell populations, maximum rate sustained 1.5-2.5 weeks before decline, and generation rate as direct function of cycling cell number and cycle time. Cell cycle contribution to generation shows cells completing 2-2.4 cycles per day at 10-12 hour cycle times, each division doubling local cell number, 10-16 fold total cell number increase over 1.5-2.5 week maximum generation period, and geometric progression creating rapid accumulation. Exponential accumulation pattern shows characteristic exponential growth curve during maximum generation, growth equation N(t) = N₀ × 2^(t/T) where T is doubling time 10-12 hours, exponential phase representing unrestricted proliferative capacity, and transition to declining rate as cells exit cycle. Cell population expansion shows all actively dividing cells contributing to generation, population size increasing as divisions produce more dividing cells, self-amplifying process characteristic of exponential growth, and population expansion creating majority of final cellularity. Division-competent cell fraction shows high proportion 70-85% mesocarp cells actively cycling during maximum generation, cycling fraction representing cells capable of division versus quiescent or differentiating cells, and high fraction enabling maximum generation output. Spatial generation patterns show cell generation throughout mesocarp volume, possible gradient with higher generation near seeds though relatively uniform overall, coordinated generation maintaining tissue organization, and spatial patterns creating organized berry architecture. Mesocarp cell generation shows flesh tissue as primary generation site accounting for 80-85% total new cells, mesocarp volume increase from cell accumulation, and mesocarp generation determining bulk berry size. Exocarp cell generation shows surface layer generation accounting for 10-15% total new cells, anticlinal divisions maintaining surface coverage during berry expansion, and coordinated surface generation preventing cracking. Generation sufficiency for size shows cell number achieved during maximum generation largely determining final berry size, typical strong correlation r² 0.70-0.85 between peak generation output and mature berry diameter, and generation period as critical size determinant. Cell number finalization shows maximum generation producing 60-75% final cell number, remaining 25-40% cells from continued declining-rate division weeks 3.5-5, and peak generation as dominant cellularity source. Environmental effects on maximum generation include temperature critically affecting generation rate through cycle time effects with optimal 24-28°C supporting 10-12 hour cycles and maximum generation, cool conditions 18-22°C extending cycles to 13-16 hours reducing generation rate 30-50%, warm conditions above 32°C potentially accelerating cycles though excessive heat impairing division, and temperature during peak weeks 2-3 as critical generation determinant. Auxin effects show hormone from seeds sustaining maximum generation, auxin levels peak weeks 1-3 supporting sustained division, hormone decline week 3-4 correlating with generation rate decline, and auxin as primary generation stimulus. Nutrient effects show adequate nitrogen supporting protein synthesis for rapid generation, phosphorus for DNA synthesis, and overall nutrition usually adequate though deficiency could limit generation in extreme cases. Failed or limited generation causes include suboptimal temperature during peak period permanently reducing achieved cell number, inadequate auxin from poor seed development limiting generation stimulus, severe stress during weeks 2-3 arresting division, and environmental limitations during critical period irreversibly reducing final cellularity and size. Generation assessment methods include cell counting at different time points quantifying accumulation rate, mathematical modeling of exponential growth fitting accumulation curves, mitotic index measurement correlating with generation rate, and population doubling time calculation from accumulation dynamics. The process represents first aspect of peak division activity emphasizing proliferative output, followed by pericarp and seed compartment proliferation discussing spatial distribution. Duration typically 1.5-2.5 weeks representing maximum generation period. Regional timing shows maximum cell number generation during early-mid July across climates weeks 2-3 post-fertilization. Viticultural significance includes representing critical period generating majority of berry cellularity, demonstrating extreme importance of optimal conditions during brief peak weeks, showing temperature as major generation rate determinant, and establishing maximum generation as primary berry size determinant through cell number effects. Research applications include analyzing generation dynamics and mathematical modeling, investigating environmental effects on generation rate and final cellularity, evaluating cell cycle regulation during peak generation, quantifying generation output and size relationships, and assessing maximum generation as critical determinant of berry development outcomes.
Maximum_Cell_Number_Generation is first aspect of Peak_Division_Activity, distinguished by peak proliferative output. Represents exponential cell accumulation at maximum rate. Duration 1.5-2.5 weeks generating 60-75% final cell number. Peak rate 40,000-120,000 cells/day. Doubling time 10-12 hours. Temperature critically affects generation rate. Irreversible cellularity establishment. Followed by spatial distribution aspects in Pericarp and Seed Compartment proliferation. Critical berry size determinant.
Spitzen-Zellproduktion
Zellzahl-Akkumulationsspitze
cell number accumulation peak
exponential cell generation
maximale Zellzahl-Erzeugung
maximale proliferative Leistung
maximum proliferative output
peak cell production
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 435-437 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Maximum Cell Number Generation
Maximum cold hardiness comparison among cultivars reveals the genetic variation constraining site selection, with highly hardy cultivar (Frontenac, V. riparia hybrid) achieving maximum LT50 of -35°C (suitable for continental climates with extreme winter cold), moderately hardy V. vinifera (Riesling) achieving -25°C (suitable for cold but not extreme climates), moderately tender (Merlot) achieving -20°C (requiring more protected sites or mild climates), and tender cultivar (Grenache) achieving only -15°C (restricted to mild winter regions), with these maximum hardiness differences determining the fundamental cultivar-site combinations that can survive winter cold events.
Maximum cold hardiness quality is a cold hardiness quality that represents the peak freeze tolerance achieved by grapevine tissues during the dormant period, characterizing the lowest temperature that tissues can survive when fully acclimated, which establishes the ultimate cold protection capacity that determines survival of the most severe cold events likely to occur in a given climate. Within the BFO framework, this quality specifically addresses the peak tolerance dimension of cold hardiness, distinguishing maximum achievable hardiness from current hardiness at any moment while recognizing that maximum hardiness is only transiently achieved during mid-winter after full acclimation and before deacclimation begins, with this peak providing the critical survival margin during the typically coldest portion of winter. The ontological significance lies in maximum cold hardiness determining the absolute limit of cold tolerance and thus the survival probability under extreme conditions, where cultivars achieving greater maximum hardiness can survive colder temperatures and thus can be cultivated in more severe climates, while those with limited maximum hardiness are restricted to milder regions regardless of management practices that cannot increase the genetically determined hardiness ceiling. The genetic determination of maximum hardiness shows substantial variation among Vitis species (V. amurensis achieving below -40°C, V. riparia to -35°C, V. vinifera typically -15°C to -25°C depending on cultivar) and among V. vinifera cultivars (Riesling achieving approximately -25°C, Chardonnay -22°C, Merlot -20°C, Grenache -15°C), with this genetic variation providing the primary basis for cultivar selection in cold climates. The environmental factors affecting maximum hardiness expression include the adequacy of acclimation conditions (gradual cooling enabling full hardiness development), vine condition (healthy, well-conditioned vines achieving higher maximum than stressed vines), and the timing of cold events relative to the hardiness trajectory (maximum hardiness available only during the limited mid-winter window). Understanding maximum cold hardiness quality enables cultivar selection based on local minimum temperature extremes, assessment of survival margins under projected cold events, and realistic expectations for the cold protection achievable through management versus the genetic limits that management cannot exceed.
Maximum_Cold_Hardiness_Quality addresses the peak freeze tolerance achievable under full acclimation. This quality is genetically determined and establishes the survival limit for extreme cold events. The class supports cultivar-climate matching through maximum hardiness characterization.
Mittwinterhärte
Spitzenfrosthärte
maximale Frosthärte
maximale Kältetoleranz
maximum freeze tolerance
midwinter hardiness
peak hardiness
ultimate cold tolerance
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10(1-2): 201-235 Ferguson JC et al. (2011) Modeling dormant bud cold hardiness and budbreak in twenty-three Vitis genotypes reveals variation by region of origin. American Journal of Enology and Viticulture 62(4): 516-527
Maximum Cold Hardiness Quality
In a Châteauneuf-du-Pape Grenache vineyard, maximum color intensity was measured 12 days post-color break in early September Year N+1 (BBCH 85) when CIE L* reached minimum 31.8, chroma peaked at 47.2, absorbance at 520 nm was 2.35 OD, skin anthocyanins 1150 mg/kg (malvidin-3-glucoside 655 mg/kg, acylated forms 31%), berry Brix 22.8°, firmness 1.4 N, under optimal southern Rhône conditions (daily high 28-31°C, night 15-18°C, high UV, moderate deficit stem ψ -1.25 MPa, galets stones reflecting light enhancing cluster exposure).
Maximum_Color_Intensity represents the peak visual pigmentation achieved when anthocyanin accumulation, copigmentation phenomena, and cellular architecture create the deepest, most saturated color possible for a given cultivar under specific environmental and viticultural conditions, serving as a critical quality marker for harvest timing and wine color potential prediction. This maximum occurs during late veraison to early ripening (BBCH 85-87), typically 10-15 days post-color break, and is characterized by colorimetric measurements showing CIE L* (lightness) reaching cultivar-specific minima 20-36 (Petit Verdot, Tannat 20-26, Syrah 24-28, Cabernet Sauvignon 26-30, Merlot 28-32, Pinot Noir 30-36), chroma (C*) achieving maxima 30-58 representing peak saturation, and absorbance at 520 nm reaching 2.0-4.5 optical density units depending on cultivar and extraction method reflecting maximum light absorption by accumulated anthocyanins. Anthocyanin levels at maximum intensity range: high-intensity cultivars (Petit Verdot, Tannat, Alicante Bouschet) 1800-3500 mg/kg skin, moderate-high types (Syrah, Carignan, Touriga Nacional) 1200-2200 mg/kg, moderate cultivars (Cabernet Sauvignon, Merlot, Tempranillo) 900-1800 mg/kg, and lower-intensity varieties (Pinot Noir, Grenache, Sangiovese) 600-1400 mg/kg, with these concentrations representing 95-100% of achievable maximum under given conditions. Cellular and molecular basis of maximum intensity involves: epidermal and sub-epidermal cell vacuoles containing anthocyanins at peak concentrations 18-35 mM approaching solubility limits, vacuolar pH optimized at 2.8-3.3 maximizing flavylium cation proportion (colored form) versus carbinol pseudobase (colorless form), copigmentation at maximum efficiency with flavonol (150-300 mg/kg) and hydroxycinnamic acid (80-200 mg/kg) copigments creating 45-65% color enhancement beyond that expected from anthocyanin concentration alone through bathochromic shifts and hyperchromic effects, and anthocyanin vacuolar inclusion formation in cultivars exhibiting this trait creating localized super-concentrated pigment zones >70 mM contributing to exceptional intensity. Optical physics of maximum intensity includes: light absorption at anthocyanin λmax 520-540 nm reaching 85-95% of incident light in fully pigmented skin creating characteristic deep color, multiple scattering within multilayered pigmented epidermis and hypodermis amplifying color intensity through extended optical path length, internal reflection from berry flesh-skin interface concentrating light interaction with anthocyanin-containing cells enhancing color expression, and surface texture effects with smooth berry surfaces reflecting more light appearing slightly less intense than textured surfaces that diffuse reflection reducing specular component. Spectral characteristics at maximum intensity show: absorbance spectra displaying pronounced anthocyanin peak 520-540 nm with secondary peak 270-280 nm and characteristic three-peak pattern in visible range, reflectance spectra showing minimal green reflection (12-15 kg/vine) through improved resource allocation and berry concentration, canopy management creating optimal cluster light exposure enhancing intensity 30-50% through increased synthesis without heat stress, irrigation management imposing moderate sustained deficit enhancing intensity 30-55% compared to well-watered vines, timing of cultural practices particularly leaf removal during pre-veraison versus early veraison affecting final intensity through UV induction and temperature modulation, and disease prevention maintaining berry health enabling maximum biosynthetic capacity unimpaired by pathogen-induced metabolic disruption. Quality implications of maximum intensity inclu intensity strongly predicts wine color density with correlation r² = 0.90-0.97 between berry skin color and wine 520 nm absorbance, maximum intensity berries enable shorter maceration times or lower temperatures reducing extraction of harsh tannins while achieving adequate color, intensity uniformity across berries and clusters indicates ripening synchrony and predicts wine color consistency, and timing of maximum intensity relative to chemical and phenolic maturity determines optimal harvest window with ideal scenario showing intensity maximum coinciding with target sugar, acid, and tannin development. Measurement of maximum intensity involves: instrumental colorimetry using CIE L*a*b* color space providing objective quantification (L* minima, C* maxima indicating peak intensity), spectrophotometry measuring absorbance at 520 nm after standardized extraction correlating with visual intensity, portable instruments enabling field measurement guiding harvest decisions, image analysis of berry photographs calculating color metrics across populations revealing intensity distribution, and visual assessment by trained evaluators comparing berries to cultivar-specific color standards rating intensity on standard scales. Post-maximum intensity management includes: monitoring intensity stability through regular colorimetric or visual assessment detecting degradation indicating harvest urgency, environmental protection from heat waves through irrigation or canopy management maintaining intensity, disease prevention as colored soft berries show maximum Botrytis vulnerability potentially compromising intensity through infection-induced cellular disruption, and harvest timing optimization balancing intensity maintenance with other maturity parameters achieving desired wine style and quality.
Maximum_Color_Intensity is distinguished from Complete_Skin_Pigmentation by representing peak visual intensity rather than total anthocyanin, from Final_Color_Expression by focusing on intensity maximum rather than stable endpoint, and from Color_Intensity_Increase by showing peak rather than progression. CIE L* minima: high-intensity 20-28, moderate 26-32, lower-intensity 30-36. Timing: typically 10-15 days post-color break. Copigmentation contributes 45-65% intensity enhancement. Temperature optimum 22-28°C, cool nights enhance. Maximum_Color_Intensity indicates optimal harvest window approach for color-driven wine styles.
Spitzen-Farb-Sättigung
deepest pigmentation state
maximale chromatische Intensität
maximum chromatic intensity
optimal color darkness
optimale Farb-Dunkelheit
peak color saturation
tiefster Pigmentierungs-Zustand
Keller M (2020) The Science of Grapevines 3rd Ed pages 235-238 color intensity and quality Boulton R (2001) The copigmentation of anthocyanins and its role in the color of red wine a critical review Amer J Enol Vitic 52:67-87 copigmentation effects on intensity Guidoni S, Ferrandino A, Novello V (2008) Effects of seasonal and agronomical practices on skin anthocyanin profile of Nebbiolo grapes Amer J Enol Vitic 59:22-29 maximum intensity achievement.
Maximum Color Intensity
In Riesling vines in the Rheingau, Germany, maximum dormancy depth is reached in mid-December approximately 4 weeks after dormancy establishment, demonstrated by forcing tests requiring more than 60 days for 50% budbreak at 22°C, with this cultivar exhibiting characteristically deep dormancy that subsequently requires 1200-1400 chilling hours below 7.2°C for complete release, reflecting genetic adaptation to the continental climate of its central European origin.
Maximum dormancy depth in Vitis vinifera represents the point in the annual dormancy cycle at which internal growth inhibition reaches its greatest intensity, typically occurring in early winter before chilling accumulation has begun to release dormancy. This condition is quantified through forcing tests that measure the time required for budbreak or the percentage of buds breaking under standardized growth-promoting conditions, with maximum dormancy depth corresponding to the longest budbreak times and lowest budbreak percentages observed during the dormancy period. The timing of maximum dormancy depth varies among cultivars and is influenced by the rate of dormancy establishment during autumn, with early-dormant cultivars typically reaching maximum depth earlier than late-dormant cultivars, though all cultivars generally achieve maximum depth by early winter before substantial chilling accumulation has occurred. The physiological basis of maximum dormancy depth involves the convergent action of all dormancy-maintaining mechanisms at their peak effectiveness, including highest ABA concentrations, maximum expression of DAM genes, most extensive chromatin repression of growth genes, and lowest levels of growth-promoting hormones and their signaling components. From a practical perspective, maximum dormancy depth determines several important viticultural parameters including the minimum chilling requirement for adequate dormancy release, the susceptibility to inadequate chilling in warm winter climates, and the response to dormancy-breaking chemical treatments that are sometimes applied in regions with insufficient natural chilling. The concept of dormancy depth implies a quantitative continuum from deep to shallow dormancy, with the depth at any time reflecting the balance between dormancy-maintaining and dormancy-releasing factors, and maximum depth representing the point where maintaining factors are strongest relative to releasing factors. Cultivar differences in maximum dormancy depth are substantial, ranging from very deep dormancy in cultivars like Riesling and Cabernet Sauvignon that may require 1200-1500 chilling hours for release, to relatively shallow dormancy in cultivars like Sultana and Perlette that may require only 400-600 chilling hours. The maximum dormancy depth achieved in a given year can be influenced by environmental conditions during dormancy induction, with stress conditions or suboptimal acclimation potentially affecting the completeness of dormancy establishment and thus the depth achieved.
Maximum_Dormancy_Depth represents a quantitative characteristic within Deep_Endodormancy describing the peak intensity of dormancy. Distinguished from dormancy depth at other times which is either increasing during establishment or decreasing during chilling accumulation. Measured through forcing test response times and budbreak percentages. Cultivar variation substantial from approximately 400 to 1500+ chilling hours required for release. Timing typically early winter, 2-6 weeks after dormancy establishment completion.
Dormanz-Intensitätsmaximum
Dormanztiefe-Maximum
deepest dormancy point
dormancy maximum
maximale Dormanztiefe
maximum rest depth
peak dormancy intensity
tiefster Ruhepunkt
Lavee S, May P (1997) Dormancy of grapevine buds - facts and speculation. Australian Journal of Grape and Wine Research 3: 31-46 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Dokoozlian NK (1999) Chilling temperature and duration interact on the budbreak of Perlette grapevine cuttings. HortScience 34: 1054-1056
Maximum Dormancy Depth
Merlot maximum enlargement stage berries progress from mid-ripening at BBCH 87 (week 15, cells 138 μm diameter, berry 15 mm, 2.8 g, 23% sugars, 7.5 g/L acidity, 2.9 mg/g anthocyanins) to maturity at BBCH 89 (week 18, cells 156 μm, berry 16 mm, 3.2 g, 25% sugars, 6.2 g/L acidity, 3.3 mg/g anthocyanins), with cell expansion declining from 6 μm per week (weeks 11-13) through 4.5 μm per week (weeks 13-15) to 1.7 μm per week (weeks 15-18) approaching cessation, berry diameter increasing from 0.75 mm per week (early enlargement) to 0.33 mm per week (maximum enlargement), and growth approaching zero as maximum size is achieved creating harvest decision window. Cabernet Sauvignon maximum enlargement stage berries at weeks 16-18 show cells at 148-162 μm diameter (approaching maximum potential of 165 μm for this cultivar under these conditions), berries at 13.8-14.5 mm (approaching maximum of 15 mm), compositional maturity with sugars 24-26% (target range), acidity 5.8-6.5 g/L (acceptable range), pH 3.7-3.85 (optimal), anthocyanins 3.1-3.5 mg/g (good color), creating optimal harvest window when maximum size coincides with optimal composition. Syrah maximum enlargement stage berries examined across four harvest dates (weeks 16, 17, 18, 19) show progressive minimal size changes= week 16 (155 μm cells, 16.2 mm berry, 3.4 g), week 17 (158 μm, 16.4 mm, 3.5 g), week 18 (159 μm, 16.5 mm, 3.55 g), week 19 (158 μm, 16.4 mm, 3.5 g showing slight shrinkage), demonstrating that maximum size is achieved by week 17-18 with further delay producing no additional enlargement and potential size decrease through dehydration, supporting harvest at maximum enlargement rather than extended hang time. Chardonnay maximum enlargement stage berries experiencing late-season rain event (25 mm precipitation at week 17, berries at 13.5 mm, 2.3 g, cells 145 μm near maximum 150 μm) show 8-12% incidence of berry splitting within 24-48 hours post-rain, versus dry conditions showing <1% splitting, demonstrating that berries at maximum enlargement are vulnerable to splitting from additional water uptake as cells at maximum expansion cannot accommodate further turgor increases without wall rupture, providing physiological basis for harvest timing before late-season precipitation when possible.
A maximum enlargement stage berry is a cell enlargement stage berry during the late portion of Phase II approaching or achieving maximum cell and berry dimensions (late BBCH 87 to BBCH 89, approximately 15-18+ weeks post-fruit set), characterized by declining expansion rates, cells approaching or achieving maximum size, berry approaching or achieving maximum dimensions, and completion of ripening processes, representing berries in final maturation approaching harvest readiness. Maximum enlargement stage berries are characterized by= (1) cells approaching or at maximum expansion with cell diameter 130-180 μm representing 115-200% increase from veraison size (60-90 μm) and approaching cultivar-specific and environmentally-determined maximum (2) cell expansion rate declining to low levels (<4 μm diameter increase per week) or approaching cessation as maximum size is achieved (3) berry diameter approaching or at maximum of 12-25 mm depending on cultivar, representing 30-90% increase from veraison dimensions (9-13 mm) (4) berry fresh weight approaching or at maximum of 1.5-7 g depending on cultivar, representing 50-300% increase from veraison weight (1-2 g) (5) berry growth rate declining to low levels (diameter <0.08 mm per week, weight <0.04 g per week) or approaching zero as maximum size is achieved (6) sugar content at or approaching target maturity levels (20-28%) with accumulation rate declining to <0.4% per week as maturity is achieved (7) acidity at mature levels (4-9 g/L titratable acidity, pH 3.4-4.2) with acid decline largely completed. Maximum enlargement stage berries show completion of expansion-related processes= cell walls reaching final thickness (thinned to 0.4-0.7 μm through expansion) and final composition (modified through ripening-associated changes) vacuoles occupying maximum proportion of cell volume (92-95%) with sugars at peak concentration turgor declining to mature levels (0.3-0.5 MPa) as expansion completes water import declining as osmotic driving force diminishes. Maximum enlargement stage berries undergo final quality development= anthocyanins (red cultivars) at or near maximum levels (2.5-5+ mg/g skin) with synthesis declining or complete tannins achieving final composition through polymerization and galloylation changes flavor and aroma compounds completing development creating mature berry sensory profile seeds achieving final desiccation and hardening. Maximum enlargement stage berries represent the harvest decision window= compositional maturity assessment determining optimal harvest timing based on sugar=acid balance, phenolic maturity, and flavor development technological maturity for wine production versus physiological maturity for table grapes creating cultivar and use-specific harvest criteria potential for further improvement versus risk of over-maturity or quality decline influencing harvest decisions. Maximum enlargement stage berries show minimal response to further irrigation= cells having achieved maximum or near-maximum expansion show limited further size increase even with abundant water sugar accumulation continuing but at reduced rates excessive late-season water potentially diluting composition or promoting rot rather than improving quality. Environmental factors during maximum enlargement affect final quality rather than size= temperature influences final sugar accumulation and flavor development water deficit late in enlargement may enhance concentration through reduced dilution late-season rain can cause berry splitting in berries at maximum enlargement due to additional water uptake creating turgor exceeding cell wall strength.
Maximum_Enlargement_Stage_Berry represents final expansion and maturation (weeks 15-18+, BBCH 87-89) and is distinguished from Early_Enlargement_Stage_Berry (weeks 9-13, rapid expansion) by declining versus rapid expansion rates and approaching versus far-from-maximum size. Maximum enlargement represents the harvest decision period when berries achieve maximum size and compositional maturity. Understanding maximum enlargement stage biology is essential for harvest timing decisions, for assessing splitting risk from late-season precipitation, and for understanding the final maturation processes determining harvest-ready berry characteristics.
Endexpansions-Beere
Endgrößen-Stadium
final-expansion berry
maximale Vergrößerungs-Stadium-Beere
maximum-size berry
Matthews, M.A., Anderson, M.M. (1988). Fruit ripening in Vitis vinifera= responses to seasonal water deficits. American Journal of Enology and Viticulture 39=313-320 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 220-245.
Maximum Enlargement Stage Berry
Syrah in Rhône Valley shows maximum expansion rate during late July-early August Year N+1 at BBCH 75-76, with cells enlarging at peak velocity, volume doubling time 8-10 days, 11-fold volume increase over 3 weeks from 52 to 128 micrometers diameter, turgor 0.9-1.1 MPa exceeding 0.48 MPa yield threshold, water uptake 1.6-2.2 g per berry per day, under warm Mediterranean conditions of 26-30°C temperature with adequate irrigation, typical of this mid-to-late season cultivar achieving maximum expansion
Maximum expansion rate represents the peak cell enlargement velocity phase during rapid cell expansion when individual cell volume increases at highest rate, typical 5-15 fold volume increase over 2-4 weeks, optimized turgor-wall system enables maximum yielding and growth, and cells achieve fastest volume accumulation determining substantial size gain, occurring during peak enlargement intensity weeks 5-7 post-fertilization. This maximum velocity phase occurs during late summer in late July-early August in temperate regions Year N+1, proceeding over weeks 5-7 representing peak rate period, corresponding to BBCH 75-76 berry touch to berries beginning to touch stages. Maximum rate mechanism involves optimized turgor pressure 0.8-1.2 MPa exceeding yield threshold by maximum margin, maximum wall yielding from peak expansin and wall-modifying enzyme activity, sustained water uptake 1.2-2.5 g per berry per day supplying expanding cells, coordinated expansion processes operating at maximum efficiency, and peak rate reflecting optimal growth capacity. Cell volume increase rate shows typical individual cell volume doubling time 7-12 days during maximum rate period, exponential-like volume accumulation, 5-15 fold total volume increase during rapid expansion phase, and maximum rate representing fastest cell growth during berry development. Volume doubling characteristics show rapid doubling from high expansion rate, multiple doublings possible during 2-4 week maximum rate period, and exponential pattern characteristic of unrestricted growth. Cell diameter increase shows typical expansion from 40-70 micrometers at maximum rate onset to 80-140 micrometers by completion, diameter increase rate 3-8 micrometers per week during peak, and linear diameter increase contrasting with volumetric exponential increase. Cell volume calculation shows spherical approximation V = 4/3πr³, volume increase cubic function of radius increase, small diameter increases producing large volume increases, and geometric amplification of linear growth. Turgor pressure optimization shows sustained high turgor 0.8-1.2 MPa during maximum rate period, turgor exceeding wall yield threshold 0.4-0.6 MPa by 0.4-0.6 MPa margin, large turgor excess driving maximum expansion rate, and pressure as rate-limiting factor when suboptimal. Turgor-rate relationship shows expansion rate proportional to turgor excess above yield threshold, mathematical relationship, rate = extensibility × (P - Y), higher turgor driving faster expansion, and turgor optimization critical for maximum rate achievement. Wall extensibility during maximum rate shows wall yielding capacity maximized through peak expansin activity, optimal wall loosening-synthesis coordination, extensibility parameter high enabling rapid yielding, and wall properties optimized for growth. Expansin activity peak shows expansin gene expression maximal during rapid expansion, protein accumulation highest in expanding tissues, enzyme activity correlating with expansion rate, and expansin as key rate determinant. Water supply during maximum rate shows intensive water uptake 1.2-2.5 g per berry per day, water influx supporting rapid volume increase, adequate water availability essential for maximum rate, and water as critical resource. Water limitation effects show water deficit reducing turgor limiting expansion rate, deficit during maximum rate period severely limiting size achievement, and water as most common rate-limiting factor. Osmotic gradient maintenance shows continued solute accumulation maintaining strong osmotic potential -1.0 to -1.5 MPa, gradient driving intensive water uptake, malate and tartrate at peak concentrations 1.2-1.8% total, and gradient as water uptake driver. Metabolic support for maximum rate shows high respiratory rate providing ATP for active transport and biosynthesis, energy supporting solute accumulation and wall synthesis, metabolic capacity enabling sustained high growth rate, and metabolism as enabling factor. Respiratory characteristics show oxygen consumption elevated during rapid growth, typical rates 2-4 μmol O₂ g⁻¹ h⁻¹, respiration supporting growth processes, and metabolic activity correlating with expansion rate. Expansion rate measurement shows cell size measurements at time points quantifying rate, diameter or volume increase per unit time, typical rates 10-25% volume increase per day during peak, and rate quantification enabling growth analysis. Berry growth rate correlation shows individual cell expansion rate determining berry growth rate, berry growth as sum of individual cell expansions, coordinated cell expansion creating berry volumetric increase, and cell rate as fundamental growth determinant. Maximum rate timing shows typical peak weeks 5-7 post-fertilization, cultivar variation in peak timing affecting total expansion duration, peak representing optimal conditions and developmental state, and timing relatively consistent within cultivars. Maximum rate duration shows sustained peak rate typically 1-3 weeks, duration determining total volume gain during peak period, and sustained high rate characteristic of effective growth. Rate decline characteristics shows eventual expansion rate decline as cells approach genetic size limit, rate reduction during late rapid expansion transitioning to lag phase, and decline marking end of intensive enlargement. Environmental effects on maximum rate include temperature affecting expansion kinetics with optimal 24-28°C supporting peak rates, cool conditions reducing rate through slowed metabolism and wall modification, water availability critically affecting rate through turgor maintenance, and favorable conditions essential for maximum rate achievement. Temperature-rate relationship shows warm conditions promoting maximum rate through faster enzymatic processes, cool conditions reducing rate 30-50% though expansion continues, and temperature as major rate determinant. Water-rate relationship shows adequate water enabling full turgor and maximum rate, deficit reducing turgor proportionally reducing rate, and water deficit as most common rate limitation. The process represents first aspect of rapid cell expansion emphasizing peak enlargement velocity, preceding massive vacuole formation and peak berry growth rate. Duration typically 1-3 weeks representing peak rate period weeks 5-7. Regional timing shows maximum expansion rate during late July-early August across climates during peak enlargement. Viticultural significance includes representing fastest cell growth determining substantial size gain, requiring optimal water and temperature for maximum rate, demonstrating critical environmental sensitivity, and establishing maximum enlargement capacity determining final size. Research applications include analyzing rate-limiting factors and optimization, investigating turgor-wall relationships determining rate, evaluating water supply effects on expansion velocity, quantifying rate dynamics and environmental responses, and assessing maximum rate as determinant of berry size achievement.
Maximum_Expansion_Rate is first aspect of Rapid_Cell_Expansion, distinguished by peak cell enlargement velocity. Represents 5-15 fold volume increase over 2-4 weeks. Doubling time 7-12 days. Turgor 0.8-1.2 MPa exceeding yield threshold 0.4-0.6 MPa. Peak expansin activity. Water uptake 1.2-2.5 g/berry/day. Rate = extensibility × (P-Y). Duration 1-3 weeks peak period. Water availability most critical factor. Followed by Massive_Vacuole_Formation. Determines substantial size gain.
Spitzen-Expansionsgeschwindigkeit
fastest enlargement
maximale Expansionsrate
maximale Wachstumsrate
maximum growth rate
optimal expansion speed
peak expansion velocity
schnellste Vergrößerung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 446-449 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Maximum Expansion Rate
Pinot Noir in Oregon shows maximum flower opening on June 9 Year N+1 at BBCH 65, with 60-72% flowers simultaneously open creating full bloom appearance, abundant reproductive organs exposed throughout cluster, under cool climate conditions of 21-26°C temperature, 55-70% relative humidity, after accumulating 920-960 growing degree days base 10°C, typical of this early-to-mid season cultivar reaching flowering intensity peak for optimal pollination opportunity
Maximum flower opening represents the peak flowering intensity milestone when greatest absolute number and proportion of flowers achieve open state simultaneously, creating characteristic full bloom appearance and establishing maximum reproductive organ availability for pollination within cluster. This critical intensity maximum occurs during mid-June in temperate regions Year N+1 as central event of peak anthesis, typically representing single day or 1-2 day period when 50-75% flowers show open state simultaneously, corresponding to BBCH 65 phenological stage. Opening intensity characteristics show maximum daily opening rate with 10-20% additional flowers opening per day during peak, cumulative effect of sustained opening over previous 2-3 days creating high proportion of simultaneously open flowers, and peak representing balance point between continuing opening of new flowers and beginning senescence of earliest-opened flowers. Flower distribution at maximum shows opened flowers throughout cluster creating dense reproductive display with apical regions showing 80-95% flowers open, middle regions showing 70-85% open representing highest density, and basal regions showing 40-70% open with opening continuing, creating overall highly functional reproductive state across cluster. Visual appearance at maximum bloom shows characteristic flowering display with numerous opened flowers exposing yellow-brown anthers and green-white stigmas creating distinctive color pattern, contrasting with remaining green calyptra caps on 20-40% unopened buds, scattered detached caps visible on cluster and ground surfaces, and overall appearance of active vibrant reproductive state. Cluster architecture visibility at maximum bloom clearly reveals branching pattern established during previous season primordium formation, with flower spacing and distribution patterns fully expressed, facilitating assessment of cluster structure and density affecting subsequent fruit development. Reproductive organ exposure at maximum shows abundant stamens and pistils throughout cluster creating maximum surface area for pollen production and capture, with anthers at various stages from newly dehisced to 1-2 days post-dehiscence, and stigmas ranging from newly receptive to peak receptivity providing widespread functional reproductive capacity. Pollen production at maximum reaches peak levels with numerous anthers simultaneously releasing pollen creating maximum pollen density in cluster atmosphere, fresh pollen showing optimal viability 70-90% germination capacity, and cumulative pollen from recently opened flowers supplementing current production, establishing optimal pollen availability conditions. Stigma receptivity distribution at maximum shows majority of stigmas at or approaching peak receptive state with active exudate production creating adhesive pollen capture surfaces, receptive stigmas distributed throughout cluster providing abundant pollination opportunity, and spatial arrangement facilitating efficient pollen transfer from anthers to stigmas within same flowers or neighboring flowers. Pollination opportunity at maximum reaches optimal level with highest probability of successful pollen-stigma contact through abundant pollen and widespread receptivity, favorable spatial arrangement with opened flowers in close proximity facilitating pollen dispersal and capture, and highest potential for efficient pollination supporting maximum fruit set. Temporal dynamics at maximum show brief peak period lasting 1-2 days as opening rate reaches maximum then begins declining, with individual flowers showing variable longevity creating continuous turnover maintaining high open proportion, and transition beginning toward declining phase as earliest flowers start senescence. Environmental conditions required for maximum opening include optimal temperature 24-30°C supporting rapid synchronized opening and maximum daily increments, moderate humidity 50-70% facilitating cap shedding without excessive desiccation, favorable weather stability enabling consistent opening progression without interruption, and absence of rain preventing interference with normal cap detachment and organ function. Diurnal pattern at maximum shows most intense opening during morning hours 8am-12pm as temperature rises and humidity declines, reduced opening during afternoon and evening, and resumption following morning creating daily peak opening periods, with pattern consistent throughout maximum bloom period. Metabolic status at maximum shows elevated activity across reproductive structures with high respiration supporting pollen maturation and release, active stigma metabolism maintaining receptivity, and substantial energy demand requiring adequate carbohydrate supply from photosynthesis and reserves. Resource allocation at maximum emphasizes reproductive demands competing with vegetative growth for available resources, with adequate supply supporting both processes and limitation potentially affecting reproductive success, highlighting importance of balanced vine vigor and resource status. Developmental uniformity implications show highly synchronized development producing sharp intense maximum with 60-75% simultaneous opening supporting concentrated efficient pollination, while heterogeneous development creates broader flatter maximum with 40-60% opening extending period and potentially reducing pollination efficiency. Phenological significance of maximum bloom includes defining peak flowering date used for phenological records and heat summation calculations, providing reference for predicting subsequent developmental timing including fruit set, berry development, and véraison, and establishing critical benchmark for management scheduling and climate studies. Observational assessment at maximum involves whole-cluster visual evaluation estimating proportion of opened flowers, random flower counts on representative clusters providing quantitative data, and photographic documentation capturing peak bloom appearance for records, with maximum bloom easily identifiable through dense flower display. Management implications at maximum include suspending canopy operations to avoid physical disturbance of flowers, restricting fungicide applications due to bloom application limitations, monitoring weather conditions for potential protective interventions if adverse conditions threaten, and assessing bloom uniformity across vineyard identifying areas with developmental variation. Weather sensitivity at maximum reaches absolute peak with opened flowers extremely vulnerable to adverse conditions including rain causing catastrophic pollination failure, excessive heat potentially causing organ dysfunction, and cold conditions impairing reproductive function, making weather monitoring and forecasting critical during maximum bloom period. Disease vulnerability at maximum emphasizes botrytis with abundant susceptible flower tissues providing infection opportunity particularly under wet conditions, requiring that protective fungicide coverage be established before bloom as applications during flowering are restricted or prohibited. The process represents first of three peak anthesis aspects establishing maximum flowering intensity, followed by peak pollen availability and optimal pollination window creating reproductive opportunity. Duration represents brief 1-2 day period at flowering intensity peak, with rapid transition toward declining intensity as opening rate decreases. Cultivar variation appears in maximum intensity with some varieties showing very sharp peak with 65-80% simultaneous opening while others show broader peak with 50-65% opening, maximum duration with some showing brief 1-day peak while others show extended 2-3 day maximum, and timing relative to optimal environmental conditions affecting reproductive success. Regional timing shows maximum flower opening during early-mid June in warm climates at approximately 1050-1100 growing degree days base 10°C, mid-June in temperate regions at 950-1000 GDD, and mid-late June in cool areas at 850-900 GDD, with timing ideally coinciding with favorable weather window. Viticultural significance includes defining definitive peak bloom date for phenological records, representing moment of maximum yield determination potential with outcomes during this brief period dominating fruit set, requiring optimal environmental conditions for maximum reproductive success, marking most critical weather vulnerability period of entire season, and providing visual confirmation of flowering intensity and uniformity affecting management decisions. Research applications include documenting precise timing of maximum bloom for phenology studies, analyzing environmental controls of flowering intensity affecting peak characteristics, evaluating developmental uniformity implications for bloom patterns and fruit set, quantifying relationships between maximum bloom conditions and reproductive outcomes, and assessing climate change impacts on flowering intensity and timing across regions.
Maximum_Flower_Opening is first of three Peak_Anthesis aspects, distinguished by achieving highest proportion of simultaneously open flowers. Represents definitive peak bloom corresponding to BBCH 65. Followed by Peak_Pollen_Availability and Optimal_Pollination_Window creating reproductive opportunity. Brief 1-2 day period of maximum intensity. Critical phenological marker. Weather sensitivity absolute maximum. Visual assessment easily identifies maximum bloom state.
Blütenmaximum
Vollblütehöhepunkt
flowering maximum
full bloom peak
höchste Blühintensität
maximale Blütenöffnung
maximum bloom
peak flower opening
Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 372-378
Maximum Flower Opening
Comprehensive cataloging of Maximum_Hardiness_Disposition at South Dakota State University compiled midwinter LT50 data for 150 cultivars and breeding selections spanning the diversity of cultivated grapevine, establishing a reference database for cold-climate cultivar selection. Primary bud LT50 was determined by differential thermal analysis at maximum hardiness (mid-January) over 3-5 winters per genotype to account for year-to-year variation. Maximum_Hardiness_Disposition ranged from -14°C ('Grenache', least hardy V. vinifera tested) to -42°C ('Valiant', V. riparia × V. amurensis derivative), encompassing 28°C variation across the germplasm base. V. vinifera cultivars clustered between -15°C and -25°C, with 'Riesling' (-23°C) and 'Cabernet Franc' (-22°C) among the hardiest vinifera. Interspecific hybrids showed a continuum reflecting their species composition, with 'Frontenac' (-33°C), 'Marquette' (-32°C), and 'La Crescent' (-29°C) achieving commercially viable hardiness for the Upper Midwest. Year-to-year variation within genotypes averaged ±1.5°C, with rankings highly consistent across years (r = 0.94 between year pairs). The strong heritability of Maximum_Hardiness_Disposition (h² = 0.82) confirms its genetic basis and supports selection for this trait in breeding programs. Regional analysis established -25°C as the minimum Maximum_Hardiness_Disposition threshold for reliable production in USDA Zone 4 climates where temperatures routinely reach -30°C.
A Maximum_Hardiness_Disposition is an Acclimation_Disposition that inheres in a Vitis vinifera plant and is realized through the ultimate level of freezing tolerance achieved when cold acclimation is complete, representing the genetic ceiling for cold hardiness expression under optimal acclimation conditions. This disposition represents the peak hardiness capacity that defines a genotype's potential to survive extreme midwinter cold, distinct from the rate or timing of hardiness acquisition. Maximum hardiness is typically achieved in mid-January in the Northern Hemisphere after completion of both photoperiod-induced and low-temperature-induced acclimation stages, and is quantified as the minimum LT50 value observed during the dormant season. The maximum hardiness potential of a genotype reflects the cumulative effectiveness of all cryoprotective mechanisms including membrane stabilization, cryoprotectant accumulation, dehydration tolerance, and deep supercooling capacity in xylem tissues. Maximum_Hardiness_Disposition shows strong genetic control with high heritability, ranging from approximately -15°C for the least hardy V. vinifera cultivars to below -40°C for the hardiest V. amurensis and V. riparia genotypes. Achievement of genetic maximum hardiness potential depends on adequate acclimation conditions, as truncated photoperiod exposure, insufficient chilling, or premature warm periods can prevent full hardiness expression. Maximum_Hardiness_Disposition is the primary trait reported in cultivar cold hardiness characterizations and the benchmark for cultivar suitability assessment in cold-climate regions.
Maximum_Hardiness_Disposition is_a Acclimation_Disposition. Maximum hardiness represents the genetic potential for cold tolerance, achieved under optimal acclimation conditions. This is the most commonly reported cold hardiness parameter in cultivar characterizations. Maximum hardiness shows high heritability and strong genetic control, varying from approximately -15°C (tender V. vinifera) to below -40°C (hardiest V. amurensis). Environmental factors can prevent achievement of genetic maximum if acclimation is incomplete.
Härtepotential
Maximale-Frosthärte
Maximale-Härte-Disposition
hardiness ceiling
maximum freezing tolerance
peak cold hardiness
Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235 Mills LJ et al. (2006) Cold hardiness of Vitis vinifera and interspecific hybrid grapevine cultivars, American Journal of Enology and Viticulture 57:238-245 Luby JJ (1991) Breeding cold-hardy fruit crops in Minnesota, HortScience 26:507-512
Maximum Hardiness Disposition
Maximum hardiness level quality screening across the Vitis genus established the range of cold tolerance available for cultivar selection across the spectrum of cold-climate viticultural conditions. Controlled acclimation followed by differential thermal analysis (DTA) to determine bud low-temperature exotherm (LTE) quantified maximum hardiness level quality at full acclimation. Within Vitis vinifera, maximum hardiness level quality ranged from -20°C LT50 for cold-sensitive cultivars including Merlot and Grenache through -23 to -24°C for intermediate cultivars including Chardonnay and Pinot Noir to -26 to -27°C for the most cold-hardy including Riesling and Cabernet Franc, establishing a 7°C range within the species. Interspecific hybrids demonstrated superior maximum hardiness level quality, with Frontenac achieving -33°C, Marquette reaching -35°C, and wild Vitis riparia selections showing -40°C, providing options for regions where V. vinifera maximum hardiness level quality is insufficient for reliable survival. The maximum hardiness level quality characterization was integrated with site-specific winter minimum temperature probability analysis to generate cultivar recommendations, with cultivars requiring at least 5°C safety margin between maximum hardiness and the expected 1-in-25-year minimum temperature event to ensure acceptable long-term survival probability.
Maximum hardiness level quality is a cold acclimation quality that inheres in a grapevine cultivar or clone and represents the lowest temperature that fully acclimated dormant tissues can survive, establishing the ultimate cold tolerance ceiling for the genotype that determines whether vines can persist through the coldest temperatures expected at a given site, with the maximum hardiness level varying substantially among cultivars and species in ways that define the geographic limits for successful cultivation. This quality captures the endpoint of the acclimation process representing the full expression of all cold-protective mechanisms at their maximum capacity, recognizing that maximum hardiness achieved during midwinter following complete acclimation establishes the absolute survival threshold that cannot be exceeded regardless of additional cold exposure or management interventions. The ontological significance of maximum hardiness level quality lies in its role as the fundamental constraint on cultivar suitability for cold climates, with the probability distribution of winter minimum temperatures relative to maximum hardiness determining the frequency of damaging or lethal cold exposure that establishes whether a cultivar can be grown reliably at a given location. The biological basis of maximum hardiness level encompasses the full deployment of all cold-protective mechanisms including maximum cryoprotectant accumulation, optimal membrane lipid composition adjustment, complete dehydration of susceptible tissues, and the maximum supercooling capacity achievable by bud tissues, with the genetic limits on each of these mechanisms collectively determining the hardiness ceiling for the genotype. The substantial variation in maximum hardiness level among Vitis species reflects adaptation to different climatic regimes, with V. vinifera cultivars typically achieving maximum bud hardiness in the range of -20 to -27°C, while interspecific hybrids incorporating germplasm from northern-adapted species including V. riparia may achieve maximum hardiness of -35°C or lower, extending the range of cultivar options for extremely cold viticultural regions. The maintenance of maximum hardiness through the winter requires sustained cold conditions, with warm periods causing partial deacclimation that may reduce hardiness below the maximum achievable level, making maximum hardiness level quality an idealized measure that may not reflect actual hardiness status during variable winter weather.
Maximum_Hardiness_Level_Quality is_a Cold_Acclimation_Quality, representing the ultimate cold tolerance ceiling achieved at full acclimation. This class addresses the fundamental genetic limit on freezing tolerance. LT50 at maximum acclimation provides quantitative metric. The 7°C V. vinifera range (-20 to -27°C) and extension to -35°C or lower with hybrid germplasm define available options. Site-cultivar matching requires safety margin between maximum hardiness and expected minimum temperatures. Midwinter warm spells may prevent full expression of maximum hardiness potential.
Härte-Obergrenze
Spitzen-Winterhärte
maximale Kältehärte
maximales Härteniveau
maximum cold hardiness
peak winter hardiness
ultimate freezing tolerance
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Londo JP, Kovaleski AP (2019) Deconstructing cold hardiness: variation in supercooling ability and chilling requirements in the wild grapevine Vitis riparia. Australian Journal of Grape and Wine Research 25:276-285 Mills LJ et al. (2006) Timing of low temperature exposure affects cold hardiness and cold deacclimation kinetics in grape. American Journal of Enology and Viticulture 57:20-26
Maximum Hardiness Level Quality
In a McLaren Vale Shiraz vineyard in Australia, maximum malate breakdown was quantified over 13 days in mid-February Year N+1 (BBCH 85) when malate declined from 5.6 to 1.8 g/L (average 0.58 g/L per day, 4.1 g/L per week loss), combined malic enzyme activity 72 units/g FW, berry respiration 2.6 μmol O₂ g⁻¹ h⁻¹ with 68% CO₂ from malate based on isotope studies, pH increased from 3.36 to 3.69, Brix 23.5° to 27.2°, under hot Australian summer conditions (daily high 32-36°C sustaining maximum breakdown rates).
Maximum_Malate_Breakdown represents the period of most intensive malic acid catabolism when enzymatic capacity, substrate availability, and environmental conditions combine optimally producing the highest malate-specific degradation rates that primarily drive overall acid decline fundamentally altering berry organic acid balance and defining wine acidity characteristics. This maximum breakdown occurs during mid-to-late veraison (BBCH 84-85), typically 16-24 days post-veraison onset depending on temperature regime and cultivar, and is characterized by malate declining at peak rates 0.40-0.70 g/L per day (2.8-4.9 g/L per week) representing 6-10% of remaining malate per day under warm conditions, malic enzyme activities sustained at absolute maximum 55-90 units/g fresh weight combined NADP-ME and NAD-ME operating near saturation, respiratory CO₂ production from malate reaching maximum contribution 60-80% of total berry respiration indicating acid catabolism temporarily dominates energy metabolism, and malate concentration potentially declining from 4-8 g/L at maximum onset to 1.5-4.5 g/L at conclusion over 10-16 day period representing 50-70% depletion during this intensive phase. Enzymatic mechanisms of maximum breakdown include NADP-malic enzyme at peak activity 45-75 units/g fresh weight catalyzing oxidative decarboxylation malate → pyruvate + CO₂ + NADPH at maximal velocity 32-68 μmol malate oxidized g⁻¹ h⁻¹ in vitro with in vivo rates 4-7 μmol g⁻¹ h⁻¹ accounting for compartmentation and regulatory factors, NAD-malic enzyme contributing 15-30 units/g providing secondary pathway augmenting total capacity, citric acid cycle enzymes operating at elevated rates with malate dehydrogenase 30-55 units/g converting malate to oxaloacetate feeding cycle for complete oxidation, and respiratory chain complexes maximally active oxidizing NADH and FADH₂ from malate catabolism producing ATP with cytochrome c oxidase 15-28 units/g ensuring efficient electron transport. Gene expression supporting maximum breakdown shows VvME1 transcript sustained at 40-75 fold above green berry baseline maintaining enzyme protein synthesis, VvME2 similarly elevated 30-60 fold providing redundancy and capacity, citric acid cycle genes (VvMDH, VvCS, VvIDH) maintained at 8-25 fold supporting pathway flux, and transcriptional stability with half-life 8-14 hours requiring continuous synthesis but steady-state levels achieved indicating balanced synthesis-degradation maintaining enzyme concentrations throughout maximum period. Substrate kinetics during maximum breakdown demonstrate cytoplasmic malate 12-22 mM substantially above enzyme Km 0.5-2.5 mM ensuring near-Vmax operation with substrate saturation 85-95%, vacuolar malate 3-7 g/L (30-70 mM) declining but still adequate for sustained cytoplasmic supply, tonoplast transport delivering 3.5-6.0 μmol malate g⁻¹ h⁻¹ from vacuole to cytoplasm matching or slightly exceeding consumption rate preventing substrate limitation, and pH gradient (vacuole 2.9-3.3, cytoplasm 7.1-7.5) maintaining driving force for facilitated diffusion potentially augmented by H⁺-coupled secondary active transport. Metabolic fate of breakdown products includes pyruvate from malic enzyme (3.5-6.5 μmol g⁻¹ h⁻¹ production) predominantly entering mitochondria through pyruvate carrier for oxidation via pyruvate dehydrogenase then citric acid cycle producing acetyl-CoA representing primary fate 70-85%, minor cytoplasmic metabolism through pyruvate kinase or transamination 10-20%, fermentative conversion to lactate or ethanol under hypoxic conditions particularly berry interior 5-12%, and complete oxidation pathway yielding CO₂ quantified through respiratory measurements showing 60-80% of evolved CO₂ derived from malate during maximum breakdown based on isotope tracer studies and mass balance calculations. NADPH utilization from maximum malic enzyme activity includes reductive biosynthesis consuming 45-70% for flavonoid synthesis (anthocyanins in reds accumulating 50-120 mg/kg per day during maximum requiring substantial NADPH), fatty acid synthesis for membrane and cuticle (15-25% NADPH demand), terpenoid biosynthesis particularly aromatic varieties (8-18%), antioxidant systems (glutathione reductase, thioredoxin reductase) maintaining redox homeostasis (12-22%), and excess NADPH oxidation by mitochondrial external NADH dehydrogenase for ATP generation though less efficient than direct NADH oxidation. Energy budget during maximum breakdown shows ATP production from malate catabolism reaching 60-105 μmol g⁻¹ h⁻¹ (based on 4-7 μmol malate g⁻¹ h⁻¹ × 15 ATP per malate complete oxidation) providing 45-75% of berry total energy requirements during this phase, ATP utilization supporting sugar transport (18-32 μmol ATP g⁻¹ h⁻¹), biosynthesis including anthocyanins and secondary metabolites (12-25 μmol g⁻¹ h⁻¹), membrane ion pumps maintaining electrochemical gradients (15-28 μmol g⁻¹ h⁻¹), and cellular maintenance respiration (10-20 μmol g⁻¹ h⁻¹), demonstrating critical role of malate catabolism supporting intensive ripening metabolism. Temperature sensitivity of maximum breakdown shows Q₁₀ = 2.5-4.0 indicating very strong temperature dependence with each 10°C increase producing 2.5-4 fold rate enhancement, optimal temperature 28-32°C producing maximum rates 0.55-0.75 g/L per day enabling efficient breakdown without excessive loss, warm conditions 32-35°C causing accelerated breakdown 0.65-0.95 g/L per day creating risk of over-deacidification, moderate warmth 24-28°C supporting appropriate rates 0.40-0.60 g/L per day achieving balanced final composition, and cooler conditions 18-24°C limiting to 0.25-0.42 g/L per day preserving acids beneficial for high-acid cultivars or cool-climate viticulture. Diurnal dynamics during maximum breakdown reveal pronounced variation with daytime peak 0.70-1.20 g/L per day during afternoon maximum temperatures 32-42°C berry surface from solar heating, morning and evening rates 0.40-0.65 g/L per day at moderate temperatures 25-30°C, nighttime minimum 0.18-0.40 g/L per day with berry cooling to 15-22°C substantially reducing metabolic rates, and daily integrated breakdown averaging 0.45-0.75 g/L representing time-weighted mean critically determining cumulative loss over maximum period. Cluster microclimate variations produce substantial differences in maximum breakdown with sun-exposed southwestern clusters (northern hemisphere) experiencing 70-140% faster breakdown than shaded positions through elevated temperatures 7-15°C above air temperature, eastern exposure showing moderate rates through morning sun limited afternoon heating, northern positions maintaining coolest temperatures preserving acids through reduced breakdown 40-75% of southwestern rates, and targeted shading during maximum period most effective intervention reducing breakdown 35-65% through berry temperature moderation 5-10°C cooling. Water status interactions during maximum breakdown demonstrate adequate vine hydration essential for supporting maximum metabolic flux enabling appropriate rates without stress acceleration, moderate deficit (stem ψ -1.0 to -1.4 MPa) showing inconsistent effects across cultivars with some displaying 15-30% enhancement through stress-induced respiration while others maintain normal rates, severe deficit (3.6-3.8 problematic for structure, stability, and MLF kinetics, inadequate breakdown maintaining >5-7 g/L creating overly tart wines requiring deacidification or extended MLF, and timing relative to phenolic and flavor development affecting harvest decisions requiring integration of multiple maturity parameters. Viticultural management during maximum includes strategic canopy shading reducing berry temperature 5-11°C slowing breakdown 40-70% preserving acids most effective in warm regions, irrigation timing potentially moderating through transpirational cooling or direct berry effects though mechanisms complex, avoiding severe deficit stress that can either accelerate through stress responses or impair through metabolic dysfunction depending on intensity, disease control essential as high-sugar soft berries increasingly vulnerable to Botrytis and sour rot requiring preventive protection, and harvest readiness as maximum breakdown indicates approaching maturity requiring prompt action when optimal parameters achieved.
Maximum_Malate_Breakdown is distinguished from Peak_Acid_Reduction_Rate by focusing on malate-specific rather than total TA, from Active_Acid_Degradation by representing maximum phase rather than overall active period, and from Respiratory_Malate_Consumption by showing peak rather than sustained process. Rates: 0.40-0.70 g/L per day malate. Duration 10-18 days BBCH 84-85. Respiratory contribution 60-80% CO₂ from malate. Temperature Q₁₀ 2.5-4.0. Maximum_Malate_Breakdown represents most intensive period of malate depletion fundamentally determining wine acidity character.
Spitzen-Malat-Katabolismus
highest malate oxidation rate
höchste Malat-Oxidations-Rate
intensive malic acid breakdown
intensive Äpfelsäure-Abbau
maximale L-Malat-Degradation
maximum L-malate degradation
peak malate catabolism
Sweetman C, Sadras VO, Hancock RD, Soole KL, Ford CM (2014) Metabolic effects of elevated temperature on organic acid degradation in ripening Vitis vinifera fruit J Exp Bot 65:5975-5988 maximum breakdown Ruffner HP, Hawker JS, Hale CR (1976) Temperature and enzymic control of malate metabolism in berries of Vitis vinifera Phytochem 15:1877-1880 malate catabolism maximum Walker RP, Famiani F (2018) Organic acids in fruits:metabolism, functions and contents Hort Rev 45:371-430 malate breakdown comprehensive.
Maximum Malate Breakdown
In a Bordeaux Merlot vineyard, maximum phloem delivery was estimated during 14 days in mid-September Year N+1 (BBCH 84-85) when berry Brix increased 4.1% per week requiring approximately 240 mg sucrose per berry per day (berry size 1.85 g), vine-level delivery calculated 185 g sucrose per vine per day (crop load 9.2 kg, 4950 berries per vine), source photosynthesis 14.2 g CO₂ per vine per day (canopy 11.5 m² healthy leaf area), under favorable conditions (daily high 26-29°C, stem ψ -0.95 MPa, full sun 10-12 h).
Maximum_Phloem_Delivery represents the peak rate of sucrose transport from source leaves through vine vascular system to ripening berries achieved when phloem structure, loading, transport, and unloading all operate optimally creating the substrate supply that enables maximum sugar accumulation determining final berry sweetness. This maximum delivery occurs during mid-to-late veraison through early ripening (BBCH 84-85), coinciding with peak sugar import rates, and is characterized by sucrose mass flow to individual berries reaching 180-400 mg sucrose per berry per day (depending on berry size 1.4-2.8 g final weight) representing phloem sap delivery 0.8-2.2 mL per berry per day at typical sieve tube sucrose concentrations 400-700 mM, vine-level total berry sucrose import 120-350 g per vine per day (for typical crop loads 6-12 kg depending on variety and management), and source-sink carbon flow directing 70-85% of daily photosynthate to ripening fruit during this peak period. Phloem anatomy supporting maximum delivery includes: mature sieve tube-companion cell complexes with fully differentiated structure, sieve tube lumens 12-25 μm diameter providing adequate cross-sectional area for mass flow, sieve plates with pores 1.5-4.0 μm diameter offering minimal resistance to bulk flow, companion cells maintaining metabolic support for sieve elements including ATP synthesis and protein turnover, and vascular bundle organization with adequate phloem tissue area relative to fruit load enabling parallel pathways reducing flow resistance. Phloem loading in source leaves at maximum includes: mesophyll photosynthesis producing triose phosphates exported to phloem at 6-12 μmol CO₂ fixed m⁻² leaf area s⁻¹ under optimal conditions, sucrose synthesis in intermediary cells or companion cells concentrating sugar from 50-150 mM mesophyll to 400-800 mM phloem sap through active H⁺-coupled symport, loading polymer trap mechanisms in some species potentially contributing to concentration though grapevine primarily uses apoplastic loading, and source leaf capacity with typical 8-12 m² canopy area per vine producing 80-160 g sucrose per day supporting berry demand during maximum delivery. Phloem transport mechanics involve: pressure flow mass flow driven by osmotic gradients with source turgor 0.8-1.5 MPa declining to sink turgor -0.2 to -0.6 MPa creating 1.0-2.1 MPa pressure difference driving flow, sieve tube sap velocity 25-60 cm per hour in vine phloem transporting sucrose from leaves to berries within 2-6 hours depending on distance, viscosity of phloem sap ~1.5-3.0 times water viscosity from high sucrose concentration creating flow resistance requiring adequate pressure differential, and temperature effects with optimal 22-28°C supporting maximum flow while cooler temperatures increasing viscosity and heat stress potentially disrupting membrane function affecting loading and unloading. Phloem unloading at berry during maximum delivery shows: apoplastic pathway dominant with >85-95% of unloading releasing sucrose into cell wall space (compared to 32°C reducing delivery through impaired photosynthesis (stomatal closure, photoinhibition), reduced phloem loading, or disrupted transport, cool conditions 800-1200 μmol photons m⁻² s⁻¹) saturating photosynthesis producing maximum sucrose for export, and photoperiod with longer days (14-16 h light in temperate summers) providing extended photosynthesis period increasing daily carbon fixation supporting delivery. Water status effects on delivery inclu adequate vine water status essential with stem water potential -0.5 to -1.1 MPa supporting maximum phloem function, moderate deficit -1.0 to -1.4 MPa potentially compatible maintaining delivery though photosynthesis may decline 15-30% slightly limiting substrate, severe deficit 30-40% leaf area removal potentially limiting delivery, irrigation management maintaining adequate vine water status supporting phloem function without excess promoting vegetative growth competing with fruit, crop load adjustment to match source capacity with typical 0.8-1.2 kg fruit per m² leaf area enabling maximum delivery without source limitation, and disease control maintaining healthy foliage essential for sustained photosynthesis and sucrose export. Quality implications of maximum delivery inclu delivery capacity fundamentally determines achievable final sugar with inadequate delivery limiting Brix regardless of berry sink capacity, timing and duration of maximum delivery affecting harvest window and quality with sustained delivery enabling higher final sugars or earlier harvest at target levels, uniformity of delivery across vines and vineyard zones affecting maturity variation requiring management attention, and balance between delivery and other quality parameters with excessive delivery potentially producing high-sugar fruit before adequate flavor or phenolic development creating harvest timing challenges. Monitoring delivery through vine physiology provides management insights: leaf photosynthesis measurements indicating source capacity, phloem sap analysis revealing sucrose concentration and transport rates though technically challenging, berry sugar accumulation rates indirectly indicating delivery adequacy, and vine water status monitoring ensuring adequate hydration supporting phloem function throughout peak delivery period.
Maximum_Phloem_Delivery is distinguished from Peak_Sugar_Import_Rate by representing vascular supply rather than cellular uptake, from Active_Sugar_Loading by focusing on transport rather than accumulation mechanism, and from initial delivery by showing maximum (180-400 mg/berry/day) rather than moderate rates. Timing coincides with peak import BBCH 84-85. Source capacity and transport efficiency both critical. Temperature optimum 24-28°C. Adequate water status essential. Maximum_Phloem_Delivery represents critical source-sink integration determining achievable berry sweetness.
Spitzen-Saccharose-Transport
Spitzen-Substrat-Versorgung
maximale vaskuläre Lieferung
maximum vascular delivery
optimal phloem flux
optimaler Phloem-Fluss
peak substrate supply
peak sucrose transport
Keller M (2020) The Science of Grapevines 3rd Ed pages 156-162, 223-225 phloem transport and sugar accumulation Ollat N, Gaudillère JP (1998) The effect of limiting leaf area during stage I of berry growth on development and composition of berries of Vitis vinifera L. cv. Cabernet Sauvignon Amer J Enol Vitic 49:251-258 source-sink relationships Patrick JW (1997) Phloem unloading:sieve element unloading and post-sieve element transport Annu Rev Plant Physiol Plant Mol Biol 48:191-222 unloading mechanisms.
Maximum Phloem Delivery
Cabernet Franc in Loire Valley shows maximum pollen availability during mid-June Year N+1 at BBCH 65 on June 10, with atmospheric concentration 700-1200 grains per cubic centimeter, fresh pollen viability 76-88%, continuous production maintaining 68-82% functional viable proportion in atmospheric pool, under temperate maritime conditions of 22-26°C temperature, 56-70% relative humidity, supporting optimal pollination during critical 1-day peak enabling 56-66% fruit set typical of this mid-season cultivar determining majority season yield
Maximum pollen availability represents the optimal male gamete supply state during peak release when combined effects of intensive discharge, sustained production, and favorable dispersal create highest concentration of fresh viable pollen grains accessible for pollination, establishing ideal conditions for maximum fertilization success and fruit set. This critical male reproductive optimum occurs during peak pollen release in mid-June in temperate regions Year N+1, typically spanning 1-2 days when coordinated factors produce absolute maximum viable pollen supply, corresponding to BBCH 65 full bloom stage. Availability characteristics show simultaneous occurrence of optimal supply elements including maximum pollen production from widespread anther discharge, continuous fresh pollen release maintaining viable supply, optimal atmospheric distribution providing wide spatial availability, maximum pollen viability with fresh grains showing 75-90% germination capacity, and favorable temporal persistence maintaining supply during stigma receptivity period. Pollen supply intensity at maximum shows peak production rates with cluster-level discharge estimated millions of grains per hour, cumulative atmospheric accumulation creating elevated concentrations 500-1500 grains per cubic centimeter near clusters, sustained production over hours maintaining viable pool despite progressive aging reducing older grains, and overall male gamete abundance sufficient for supporting maximum pollination efficiency. Fresh pollen production dynamics shows continuous anther dehiscence releasing newly matured grains, immediate optimal viability in freshly discharged pollen exhibiting 75-90% germination capacity, and sustained fresh pollen supply offsetting viability decline in atmospheric pool maintaining overall high functional pollen proportion. Atmospheric pollen pool characteristics include total concentration representing mix of fresh high-viability and aged reduced-viability grains, functional viable proportion estimated 65-85% of atmospheric pool during maximum availability showing germination capacity, spatial distribution with elevated levels throughout cluster atmosphere and extending vine scale, and temporal persistence with sustained elevated concentration over 1-2 day maximum period. Pollen accessibility at maximum shows widespread distribution making grains available throughout cluster atmosphere, numerous pollen-stigma encounter opportunities with high atmospheric concentration increasing contact probability, favorable positioning with many pollen grains in proximity to receptive stigmas, and optimal capture efficiency with adhesive stigma exudate readily trapping contacted grains. Viability optimization at maximum involves multiple coordinated factors including continuous fresh production maintaining high-viability proportion in atmospheric pool, favorable environmental conditions preserving pollen function with optimal temperature 24-30°C and moderate humidity 50-65%, brief window before progressive aging substantially reduces viability, and overall maintenance of functional male gamete supply during receptivity period. Germination capacity of available pollen shows maximum potential with fresh grains readily germinating upon stigma contact within 1-4 hours, viable atmospheric pollen maintaining capacity for successful pollination, and overall functional male gamete availability enabling high fertilization success supporting fruit set 60-75% in good-setting cultivars under favorable conditions. Spatial availability pattern shows highest concentration immediately surrounding flowering clusters as primary pollen source, elevated levels throughout vine canopy where multiple clusters contribute to atmospheric pool, significant concentrations extending several meters between vines particularly in dense plantings, and declining concentration with distance following dispersal gradient. Temporal availability window shows peak period lasting 1-2 days during maximum release with optimal conditions, sustained elevated availability requiring continued production offsetting atmospheric removal through settling and dispersal, and critical brief window requiring stigma receptivity overlap for successful pollination. Pollen-stigma encounter frequency at maximum reaches optimum with abundant atmospheric pollen creating high contact probability, estimated 10-50 pollen grains contacting individual receptive stigma during peak period though only 1-2 successful germinations required for fertilization, and favorable encounter geometry with aerial pollen and gravitational fall delivering grains to papillate stigma surfaces. Pollination efficiency at maximum shows peak success rates with abundant fresh viable pollen widely available, extensive stigma receptivity providing numerous capture sites, favorable environmental conditions supporting pollen capture and germination, and optimal coordination producing fertilization success 65-85% of pollination attempts under favorable conditions. Self-pollination supply at maximum emphasizes immediate local availability with pollen releasing from anthers in same flower falling directly onto nearby stigma, continuous discharge maintaining supply for extended receptivity period, and self-pollination accounting for 60-80% successful fertilizations during maximum availability period. Cross-pollination supply at maximum shows enhanced contribution with elevated atmospheric concentration enabling effective neighboring flower pollination, inter-flower dispersal within cluster accounting for 20-35% successful fertilizations, inter-vine dispersal contributing 5-15% particularly during very high atmospheric pollen levels, and demonstrating broader pollination sources during maximum availability compared to normal periods. Environmental conditions enabling maximum availability include temperature 24-30°C supporting intensive discharge and pollen viability, moderate humidity 50-65% maintaining pollen dispersibility and function, light-moderate wind 6-14 km per hour enhancing distribution without excessive loss, and overall stable favorable weather enabling sustained optimal availability over 1-2 day period. Temperature effects on availability show warm conditions promoting maximum discharge providing abundant supply, maintaining pollen surface properties supporting dispersibility, and preserving viability though excessive heat above 33°C potentially causing stress damage, with optimal 25-28°C range supporting all availability factors. Humidity effects show critical importance with moderate levels maintaining pollen in functional state without aggregation reducing availability or excessive desiccation damaging viability, high humidity above 70% causing aggregation and premature atmospheric removal reducing availability, and optimal 52-68% range preserving maximum functional pollen concentration. Fruit set determination during maximum availability shows outcomes during this critical 1-2 day period contributing 40-60% total berry number representing largest single period contribution, making maximum availability conditions dominant influence on season yield, and demonstrating that male gamete supply alongside female receptivity represents primary fruit set determinant. Weather vulnerability during maximum availability shows extreme sensitivity with adverse conditions during brief peak period substantially affecting outcomes, favorable stable conditions supporting high availability and fruit set while rain catastrophically eliminating availability through atmospheric removal and viability destruction potentially reducing set by 60-80%, and demonstrating critical importance of weather during maximum availability period. The process represents second and final aspect of peak pollen release emphasizing optimal gamete supply for pollination, following optimal pollen cloud formation. Duration typically spans 1-2 days representing absolute maximum male reproductive capacity, with warm stable conditions producing sharp intense peak. Cultivar variation in maximum availability shows differences in pollen production affecting achievable concentration, viability characteristics affecting functional proportion, and timing coordination with stigma receptivity affecting pollination efficiency and fruit set outcomes. Regional timing shows maximum pollen availability during mid-June in warm climates, mid-June in temperate regions, and mid-late June in cool areas, coinciding precisely with peak anthesis and full bloom representing most critical reproductive period. Viticultural significance includes representing optimal male gamete supply determining maximum pollination and fruit set potential, requiring favorable environmental conditions for achieving and maintaining maximum availability, demonstrating extreme weather vulnerability with brief 1-2 day window determining majority season yield, and establishing maximum pollen availability alongside peak stigma receptivity as integrated reproductive optimum determining productivity. Research applications include quantifying pollen availability dynamics including production rates, atmospheric concentrations, and viability distributions, evaluating environmental effects on availability optimization, assessing pollen-stigma encounter probabilities and pollination efficiency during maximum period, developing models predicting fruit set from availability characteristics, and investigating climate change impacts on maximum availability timing and conditions affecting viticultural productivity.
Maximum_Pollen_Availability is second and final aspect of Peak_Pollen_Release, distinguished by optimal male gamete supply for pollination. Represents absolute peak of male reproductive capacity. Corresponds to BBCH 65 full bloom. Duration 1-2 days during peak release. Determines 40-60% fruit set during brief window. Requires favorable environmental conditions maintaining pollen viability and dispersibility. Extreme weather vulnerability particularly to rain. Coordination with peak stigma receptivity creates integrated optimal pollination window. Most critical male reproductive state determining season productivity.
Spitzen-Pollenzugänglichkeit
highest pollen accessibility
höchster lebensfähiger Pollenbestand
maximale Pollenverfügbarkeit
maximum viable pollen
optimal male gamete availability
optimale männliche Gametenversorgung
peak pollen supply
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 384-390 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 295-298
Maximum Pollen Availability
Cabernet Sauvignon in Napa Valley shows maximum pollen dispersal during mid-June Year N+1 at BBCH 65 on June 10-11, with atmospheric concentration reaching 800-1400 grains per cubic centimeter near clusters creating visible pollen clouds, elevated 150-400 grains per cubic centimeter vine scale, dispersing effectively under warm climate conditions of 25-29°C temperature, 50-64% relative humidity, winds 8-14 km per hour, supporting efficient widespread pollination enabling 58-68% fruit set typical of this late-season cultivar during optimal 1-2 day dispersal window
Maximum pollen dispersal represents the optimal atmospheric pollen distribution phase when peak discharge creates maximum airborne grain concentration enabling widespread pollen-stigma encounters throughout cluster and vineyard, establishing ideal conditions for efficient pollination and high fruit set. This critical distribution phase occurs during peak active shedding in mid-June in temperate regions Year N+1, typically coinciding with or immediately following peak discharge over 1-2 days, corresponding to BBCH 65 full bloom stage. Dispersal characteristics show maximum atmospheric pollen concentration with elevated grain density throughout cluster atmosphere and extending vineyard scale, optimal spatial distribution providing widespread pollen availability to receptive stigmas, and favorable temporal persistence maintaining viable atmospheric pool during receptivity period. Atmospheric pollen density at maximum dispersal reaches peak levels with estimated 500-1500 grains per cubic centimeter in immediate cluster vicinity representing 10-20 fold increase above baseline, elevated concentrations 100-500 grains per cubic centimeter at cluster-to-cluster scale within vine canopy, and significant concentrations 20-100 grains per cubic centimeter extending meters between adjacent vines. Spatial distribution pattern shows highest concentration immediately surrounding flowering clusters as pollen source, declining concentration with distance following roughly exponential decay pattern, and wind-direction bias creating asymmetric distribution with higher concentrations downwind of pollen sources. Vertical distribution shows maximum concentration at flowering cluster height typically 1-2 meters above ground corresponding to typical canopy positioning, declining concentration both above and below cluster zone, and ground-level deposition creating accumulated pollen layer on soil and vegetation surfaces. Temporal persistence at maximum dispersal shows sustained elevated atmospheric concentration over hours during favorable conditions as continued discharge replenishes dispersing pollen, diurnal pattern with morning maximum coinciding with peak discharge and favorable conditions, and overall maintenance of viable atmospheric pool supporting continued pollination throughout peak period. Pollen transport mechanisms include wind-driven advection as primary dispersal process with light grains easily transported through air currents, turbulent diffusion spreading pollen through atmospheric mixing, and gravitational settling gradually removing pollen from atmosphere with terminal velocity approximately 1-3 cm per second for grapevine pollen. Wind effects on dispersal show light-moderate wind 6-14 km per hour optimal for effective dispersal enhancing pollen distribution without excessive dilution, stronger wind above 20 km per hour potentially dispersing pollen too far reducing local concentration though increasing range, and calm conditions limiting dispersal range though maintaining high local concentration supporting self and neighboring flower pollination. Dispersal range shows majority of pollen depositing within same cluster or neighboring clusters on same vine accounting for 70-90% successful pollinations, inter-vine dispersal occurring at distances up to several meters contributing 10-20% pollinations, and long-range dispersal beyond 10 meters playing minimal role <5% in reproductive success. Pollen capture efficiency varies with location showing highest capture on receptive stigmas within same cluster through combined gravitational fall and short-range wind transport, moderate capture on neighboring clusters through short-range dispersal, and low capture at distances beyond few meters due to dilution and deposition. Stigma encounter probability during maximum dispersal reaches optimum with abundant atmospheric pollen creating high probability of contact with any exposed stigma, estimated 10-50 pollen grains contacting individual receptive stigma during peak dispersal period though only 1-2 successful germinations required for fertilization, and favorable spatial-temporal coincidence supporting efficient pollination. Pollen viability during dispersal shows maximum proportion of viable grains in freshly released pollen with 75-90% germination capacity, progressive decline in atmospheric pollen as aging proceeds with viability reducing 5-10% per hour, and importance of continuous fresh pollen production maintaining viable atmospheric pool offsetting viability decline. Environmental conditions optimizing dispersal include temperature 24-30°C supporting both pollen viability and favorable atmospheric mixing, moderate humidity 50-65% maintaining pollen dispersibility without aggregation while preserving viability, light-moderate wind providing optimal transport, and overall stable conditions enabling sustained effective dispersal. Humidity effects on dispersal show moderate levels optimal for maintaining pollen in dispersible state without aggregation that would cause premature atmospheric removal, high humidity above 70% causing pollen clumping reducing airborne suspension and effective dispersal, and low humidity below 45% potentially causing excessive desiccation affecting viability though maintaining good dispersibility. Temperature effects include warm conditions generally favorable for dispersal through appropriate air density and mixing characteristics, though extreme heat above 33°C potentially affecting pollen viability during extended atmospheric residence, and optimal thermal range supporting both dispersal mechanics and pollen function. Pollen cloud visualization during maximum dispersal enables easy observation of male reproductive activity with visible pale yellow haze surrounding flowering clusters under optimal conditions, particularly evident during morning peak when discharge maximal and conditions favorable, and providing immediate visual confirmation of intensive pollination opportunity. Deposition patterns show characteristic distribution with highest accumulation on horizontal surfaces within and below canopy including leaves, shoot structures, and ground surfaces, visible yellow dust layer creating obvious evidence of flowering activity, and accumulated pollen representing unsuccessful fraction not captured by stigmas. Pollination success during maximum dispersal reaches peak levels with abundant fresh viable pollen widely distributed, extensive stigma receptivity providing numerous capture opportunities, and favorable environmental conditions supporting pollen-stigma encounters and subsequent germination, enabling fruit set 55-75% in good-setting cultivars under optimal conditions. Self-pollination efficiency maximizes during dispersal through immediate local pollen availability with freshly released grains falling directly onto stigmas in same flower, though atmospheric pool also contributes to self-pollination through brief airborne transport and return. Cross-pollination contribution during maximum dispersal shows limited but measurable role with atmospheric pollen dispersing between neighboring flowers on same cluster accounting for 20-40% successful pollinations, inter-vine dispersal contributing 5-15% particularly in dense plantings, and long-range cross-pollination between distant vines playing minimal role though demonstrating gene flow potential. Weather impacts on dispersal show catastrophic effects of rain with water washing pollen from atmosphere eliminating atmospheric pool, preventing continued discharge through elevated humidity, and destroying pollen viability, causing severe fruit set reduction potentially to 15-30% if rain occurs during maximum dispersal period. The process represents second and final aspect of active pollen shedding emphasizing atmospheric distribution and pollination opportunity, following peak pollen discharge. Duration typically spans 1-2 days coinciding with peak discharge representing optimal dispersal window. Cultivar variation in dispersal characteristics includes pollen production affecting maximum atmospheric concentration achievable, dispersal range possibly varying with pollen surface properties though poorly documented, and timing coordination with stigma receptivity affecting pollination efficiency. Regional timing shows maximum pollen dispersal during mid-June in warm climates, mid-June in temperate regions, and mid-late June in cool areas, coinciding with peak anthesis and representing most critical reproductive period. Viticultural significance includes representing optimal pollination opportunity with maximum male gamete availability and distribution, requiring favorable environmental conditions for effective dispersal and pollen viability maintenance, demonstrating extreme weather vulnerability with brief window determining majority of season yield, and providing visual confirmation through pollen clouds of successful male reproductive function. Research applications include quantifying atmospheric pollen concentrations and spatial distributions, analyzing dispersal mechanisms and ranges, evaluating environmental effects on dispersal efficiency, assessing pollen-stigma encounter probabilities and pollination success, developing dispersal models for predicting pollination outcomes, and investigating gene flow patterns through pollen-mediated dispersal.
Maximum_Pollen_Dispersal is second and final aspect of Active_Pollen_Shedding, distinguished by optimal atmospheric distribution creating widespread pollination opportunity. Follows or coincides with Peak_Pollen_Discharge. Represents culmination of male reproductive effort with maximum gamete availability. Duration 1-2 days during peak anthesis. Environmental conditions critically affect dispersal efficiency and pollen viability. Extreme weather vulnerability particularly to rain. Determines majority of fruit set through pollination success. Completes Pollen_Release hierarchical documentation.
Spitzen-Pollenausbreitung
highest pollen dispersal
höchste Pollenverfügbarkeit
maximale Pollendispersion
maximum airborne pollen
optimal pollen distribution
optimale Pollenverteilung
peak pollen availability
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 383-390 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 293-298
Maximum Pollen Dispersal
Cabernet Sauvignon maximum size late berries at BBCH 87-89 (weeks 17-19 post-fruit set) achieve maximum dimensions of 13.5-15 mm diameter and 2.4-2.9 g fresh weight (cultivar maximum for this medium-small-berried wine cultivar), cells expanded to 145-165 μm diameter, growth rate declining to <0.02 mm per week and <0.01 g per week approaching zero, sugars at 24-26% (target range for premium red wine), titratable acidity 5.5-6.8 g/L, pH 3.7-3.85, anthocyanins 2.8-3.6 mg/g skin, creating optimal harvest maturity composition with growers typically harvesting when maximum size is achieved at target sugar levels. Thompson Seedless large-berried table cultivar maximum size late berries achieve substantially larger maximum dimensions of 20-24 mm diameter and 5-7 g fresh weight at BBCH 89 (weeks 18-20), cells expanded to 160-190 μm diameter, sugars at 18-21% (table grape target range), acidity 4.5-6 g/L, demonstrating cultivar-specific variation in maximum achievable size with table cultivars selected for large berry size versus wine cultivars with smaller berries. Syrah maximum size late berries examined across three irrigation treatments show well-watered vines producing maximum size of 16-17.5 mm and 3.2-3.8 g (full expansion potential), moderate deficit producing 14.5-16 mm and 2.6-3.2 g (85-90% of full potential), and severe deficit producing 13-14.5 mm and 2-2.5 g (70-80% of potential), demonstrating water availability effects on achievable maximum size with deficit irrigation limiting cell expansion during Phase II ripening creating smaller maximum berries with more concentrated composition (deficit berries showing 25-27% sugars versus 23-25% for well-watered). Merlot maximum size late berries monitored past optimal harvest (BBCH 89 at week 18, 15 mm, 2.8 g, 25% sugars) through over-maturity (week 20, 14.8 mm, 2.7 g, 27% sugars week 22, 14.5 mm, 2.6 g, 29% sugars) show slight size decrease through dehydration while sugar increases through concentration, demonstrating that maximum size is achieved at optimal maturity with over-ripening producing shrinkage rather than continued expansion, supporting harvest at maximum size rather than waiting for further increases that will not occur.
A maximum size late berry is a late developing berry in the final ripening phase approaching or achieving maximum berry dimensions (late BBCH 87 to BBCH 89, approximately 16-18+ weeks post-fruit set), characterized by completion or near-completion of berry expansion, achievement of peak size for the genotype and environmental conditions, slowing or cessation of growth, and final compositional maturation, representing berries at or near harvest maturity. Maximum size late berries are characterized by= (1) berry dimensions at or near cultivar-specific and environmentally-determined maximum, typically 11-25 mm diameter and 1.5-7 g fresh weight depending on cultivar (small-berried wine cultivars 11-14 mm, medium-berried 14-18 mm, large-berried table cultivars 18-25+ mm) (2) cells at maximum expansion with cell diameter 120-180 μm representing 140-260% increase from veraison size (60-80 μm) and 300-500% increase from fruit set size (30-45 μm) (3) growth rate declining to very low levels (<0.03 mm diameter per week, <0.02 g per week) or approaching zero as maximum size is achieved (4) full color development with anthocyanins (red/black cultivars) at peak or near-peak levels (2-5 mg/g skin or higher in deeply colored cultivars), or deep golden-yellow in white cultivars (5) soft texture with firmness 0.8-2 N representing 75-85% reduction from green berry firmness (4-6 N) (6) sugars at or approaching target maturity levels typically 20-28% depending on cultivar and intended use, with accumulation rate slowing to <0.3% per week as maturity is achieved (7) acids at mature levels typically 4-9 g/L titratable acidity with pH 3.4-4.2, representing final acid=sugar balance determining harvest timing. Maximum size late berries show completion of quality-determining processes= anthocyanin accumulation plateauing or continuing at reduced rate with berry skin reaching final color intensity tannin composition shifting from more astringent galloylated forms to less astringent non-galloylated forms through modification reactions flavor and aroma precursor accumulation completing with berry achieving characteristic varietal aroma profile textural changes completing with skin and flesh achieving final mature texture. Maximum size late berries represent the harvest decision window where growers assess compositional maturity (sugar, acid, pH balance), phenolic maturity (color, tannin composition and extractability), and flavor maturity (aroma and flavor development) to determine optimal harvest timing balancing yield (allowing full size development), quality (achieving target composition), and practical constraints (weather, logistics). Maximum size late berries show minimal further size increases if left on vine past optimal maturity= over-mature berries may show slight shrinkage through water loss (dehydration) or may maintain size while composition continues changing (sugar increases through concentration, acid declines further, tannins become over-mature) optimal harvest typically occurs when berries achieve maximum size with optimal composition rather than waiting for absolute maximum possible size which may involve over-ripening. Environmental factors influence maximum achievable berry size= water availability during Phase II ripening determines final cell expansion with adequate water supporting full expansion versus deficit limiting expansion temperature affects sugar accumulation rate and maturation timing crop load influences resource availability per berry with low crop loads allowing larger maximum sizes versus high loads limiting individual berry size.
Maximum_Size_Late_Berry represents the final ripening phase at or near harvest (BBCH 87-89, weeks 16-18+) and is distinguished from Pre_Veraison_Late_Berry (terminal lag phase before veraison) by late ripening versus pre-ripening state. Maximum size late berries achieve cultivar-specific and environmentally-determined size limits, representing the harvest decision window. Understanding maximum size late berry characteristics is essential for harvest timing decisions, for assessing ripeness through compositional analysis, and for managing final berry size through irrigation and crop load. The maximum size designation indicates completion of expansion rather than continued growth potential.
Endgröße-Beere
Erntereife-Stadium
maximale Größe späte Beere
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 210-245 Matthews, M.A., Anderson, M.M. (1988). Fruit ripening in Vitis vinifera= responses to seasonal water deficits. American Journal of Enology and Viticulture 39=313-320.
Maximum Size Late Berry
A maximum size mature leaf at node 5 on a highly vigorous shoot of Vitis vinifera cv. Carignan in Languedoc, France, measuring 19.5 cm length by 18.8 cm width representing 95th percentile dimensions for this cultivar, total surface area of 238 cm² calculated through digital image analysis, 12.5 cm petiole length optimally positioning the large lamina for light capture, lamina thickness of 285 μm reflecting robust tissue development, fresh weight of 4.8 grams indicating substantial biomass accumulation, occurring on a shoot with exceptional vigor (total length 215 cm, basal diameter 14 mm) in a vineyard receiving full irrigation and 120 kg N/ha annual fertilization, leaf developed during optimal early-season conditions (May) with moderate temperatures (18-24°C) and excellent soil moisture (80-95% field capacity throughout expansion), demonstrating light-saturated photosynthesis rate of 14.8 μmol CO₂ m⁻² s⁻¹ near the upper range for grapevine, measured at BBCH stage 71 (fruit set) in mid-June at 4 weeks post-emergence, representing achievement of maximum genetic size potential with optimal resource availability supporting maximal development.
A fully expanded mature leaf that has achieved the upper size range possible for the cultivar and position under optimal growing conditions in Vitis vinifera, characterized by lamina dimensions approaching or exceeding cultivar-typical maximum values, typically 15-22 cm length and width in large-leaved varieties or 12-18 cm in medium-leaved cultivars, very large total surface area often exceeding 180-250 cm² enabling maximum light interception and photosynthetic capacity, robust petiole development with lengths 8-14 cm providing optimal positioning, substantial biomass reflecting extensive cell proliferation and expansion during development, and occurrence primarily on vigorous shoots with excellent resource access including abundant water, nitrogen, and other nutrients during the critical expansion phase. Maximum size mature leaves develop under optimal conditions combining high shoot vigor, early-season position (typically nodes 3-7) receiving developmental priority, abundant soil moisture throughout the 3-5 week expansion period maintaining turgor for cell enlargement, optimal temperatures of 22-28°C supporting high metabolic rates and rapid growth, adequate nitrogen availability supporting protein synthesis for cell proliferation and expansion, and absence of stress factors including pests, diseases, or environmental extremes that would limit development. These leaves represent the functional photosynthetic elite of the canopy, contributing disproportionately to whole-vine carbon gain through their large surface areas and typically optimal physiological function, often positioned in mid-canopy zones receiving favorable light conditions, and playing critical roles in supporting fruit development and reserve accumulation particularly when fruit-to-leaf area ratios approach or exceed recommended maximum values of 0.8-1.2 m² leaf area per kilogram fruit.
Maximum size mature leaves are distinguished from typical fully expanded mature leaves by dimensions in the upper size range for the cultivar (typically >85th percentile), substantial biomass, and development under optimal conditions without limiting factors. Size assessment requires calibration to cultivar norms, as maximum size for small-leaved varieties (10-14 cm) differs fundamentally from large-leaved cultivars (18-22 cm), making absolute size less informative than percentile ranking within cultivar distributions. Position strongly affects maximum size achievement, with basal to mid-shoot positions (nodes 3-8) most likely to achieve maximum dimensions while apical positions rarely reach upper size ranges due to declining resources and shortened development windows. Maximum size leaves typically indicate excellent vine vigor and resource status, serving as positive indicators when fruit-to-leaf ratios remain balanced, but excessive frequency of maximum size leaves (>40% of canopy) may indicate excessive vegetative vigor requiring management through crop load increase, water stress imposition, or fertility reduction to achieve optimal balance.
größtes Blatt
largest leaf
maximal-dimensioniertes Blatt
maximum dimension leaf
optimal size mature leaf
optimal-große reifes Weinblatt
Based on Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Palliotti et al. (2012) "Morpho-anatomical and physiological response of Pinot Noir and Montepulciano grapevine leaves to mild and severe water stress" Vitis 51=109-116 Poni et al. (2013) "Grapevine quality= A multiple choice issue" Scientia Horticulturae 234=445-462 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 119-142.
Maximum Size Mature Leaf
In a Marlborough Sauvignon Blanc vineyard in New Zealand, maximum softening rate was measured during a 36-hour period in late February Year N+1 (BBCH 84) when firmness decreased from 2.8 N to 2.1 N (6.3% per day), polygalacturonase activity peaked at 165 units, soluble pectin increased from 58% to 68% of total, and sugar concentration rose from 15.2 to 17.1° Brix, with ambient temperatures averaging 23-25°C facilitating optimal enzyme activity for rapid texture transformation.
Maximum_Softening_Rate represents the specific interval within peak softening when the velocity of firmness decline reaches its absolute maximum, typically occurring during a concentrated 24-48 hour period during mid-veraison when all softening mechanisms operate at peak efficiency. This transient but critical phase is characterized by firmness measurements showing the steepest decline gradient, often 4-8% reduction per day in early-ripening cultivars, 3-6% in mid-season varieties, and 2-5% in late-ripening types, representing the inflection point of the sigmoid softening curve where the second derivative of firmness versus time equals zero. The maximum rate interval is defined by convergence of multiple degradative processes at peak activity: polygalacturonase reaches absolute maximum activity (100-200 units depending on cultivar and conditions), pectin methylesterase achieves peak demethylation rates (120-200 units), xyloglucan endotransglycosylase shows maximum transglycosylation activity (40-80 units), and β-galactosidase displays peak activity (20-50 units) all operating simultaneously to maximize cell wall disassembly. During maximum softening rate, pectin structural changes occur most rapidly: pectin molecular weight declines at 10-20 kDa per day from approximately 130-160 kDa to 80-120 kDa, soluble pectin fraction increases 6-12% per day reaching 65-80% of total pectin, and middle lamella degradation accelerates with cell-to-cell adhesion strength dropping 25-40% per day from pre-maximum values. Cellular mechanical properties undergo dramatic transformation: compression force required for 20% deformation decreases from 3.5-5.5 N to 2.0-3.5 N within 24-48 hours, puncture resistance drops 30-50% representing compromised skin integrity, and berry elasticity modulus declines sharply indicating transition from elastic to plastic deformation behavior. The maximum rate period exhibits highest sensitivity to environmental conditions: temperature effects are amplified with Q₁₀ values of 2.5-3.5 for softening rate compared to 1.8-2.4 during other phases, water stress can reduce maximum rates by 25-45% through limitation of cell wall enzyme synthesis and activity, and extreme heat (>35°C) can actually reduce maximum rates through enzyme denaturation despite accelerated passive softening from membrane disruption. Gene expression during maximum softening rate shows peak transcription of cell wall-modifying genes: PG transcripts reach 100-200 fold elevation above green berry levels, expansin mRNAs achieve 60-120 fold increases, and pectate lyase genes show 40-80 fold upregulation, all coordinated by continued high abscisic acid signaling (1000-3000 ng/g) maintaining ripening program expression. Subcellular events during maximum rate inclu Golgi apparatus maximum activity producing cell wall-modifying enzymes at peak rates, endoplasmic reticulum stress responses from intense protein synthesis demand, and vacuolar membrane destabilization with increasing permeability facilitating cytoplasm-vacuole mixing that affects cellular compartmentation. Cultivar-specific patterns of maximum softening rate reflect genetic variation in cell wall composition and enzyme expression: Pinot Noir exhibits very rapid maximum rates (5-8% per day) associated with thin cell walls and high PG expression, Cabernet Sauvignon shows intermediate rates (3-5% per day) with thicker cell walls and moderate enzyme levels, while Nebbiolo displays slower maximum rates (2-4% per day) correlated with robust cell wall structure and lower enzyme activity. Berry vulnerability reaches its apex during maximum softening rate: skin tensile strength drops to 40-60% of pre-softening values creating maximum susceptibility to splitting and cracking, pathogen penetration resistance declines sharply with 60-80% of successful Botrytis infections initiating during this narrow window, and mechanical harvest damage potential peaks as berries lose structural integrity most rapidly. Coordination with other veraison processes is tight: maximum softening rate typically occurs 1-3 days after initial color change in red cultivars, coincides with peak rates of sugar accumulation (often same 24-48 hour window), and precedes maximum anthocyanin synthesis rates by 2-4 days allowing structural modification to support subsequent pigment accumulation. Viticultural implications of maximum softening rate timing inclu this represents the critical decision window for harvest timing in varieties where excessive softness is problematic (sparkling wine base grapes require harvesting before maximum rate period), irrigation management must be precise to avoid water stress that could limit softening but also prevent excess that accelerates it beyond optimal, and canopy management effects on bunch microclimate are maximally expressed during this interval when temperature sensitivity peaks.
Maximum_Softening_Rate is distinguished from Peak_Softening by representing the instantaneous maximum velocity point rather than the broader peak period, from other softening phases by exhibiting the absolute steepest firmness decline gradient, and from Complete_Berry_Softening by focusing on transformation velocity rather than final state. Duration typically 24-48 hours at the inflection point of the softening curve. Temperature sensitivity is maximum during this interval (Q₁₀ = 2.5-3.5). This represents the most vulnerable moment in berry development for mechanical damage and pathogen infection. Maximum_Softening_Rate timing relative to other veraison events is critical for optimal berry quality development.
Festigkeits-Verlust-Höhepunkt
Spitzen-Erweichungs-Geschwindigkeit
firmness loss apex
maximale Textur-Transformations-Rate
maximum texture transformation rate
peak softening velocity
steepest firmness decline
steilste Festigkeits-Abnahme
Keller M (2020) The Science of Grapevines 3rd Ed pages 229-230 softening kinetics and rate dynamics Thomas TR, Matthews MA, Shackel KA (2006) Direct in situ measurement of cell turgor in grape berries during development Australian J Grape Wine Res 12:173-180 firmness decline rates Nunan KJ, Sims IM, Bacic A, Robinson SP, Fincher GB (1998) Changes in cell wall composition during ripening of grape berries Plant Physiol 118:783-792 pectin degradation kinetics.
Maximum Softening Rate
Maximum stomatal conductance assessment comparing cultivars reveals genotypic variation in gas exchange capacity, with Riesling showing gsmax of 0.48 mol/m²/s (high stomatal density, 285/mm², supporting high photosynthetic potential), Cabernet Sauvignon intermediate at 0.38 mol/m²/s (density 220/mm²), and Grenache lower at 0.32 mol/m²/s (density 180/mm², consistent with water-conserving strategy), demonstrating how maximum stomatal conductance reflects cultivar-specific anatomical investment in gas exchange capacity that affects both productivity potential and water use characteristics.
Maximum stomatal conductance quality is a stomatal conductance quality that represents the highest stomatal conductance achievable under optimal conditions with fully open stomata, characterizing the anatomical and physiological capacity for gas exchange that sets the upper limit on stomatal-mediated CO₂ supply and water loss when no regulatory closure is active. Within the BFO framework, this quality specifically addresses the capacity dimension of stomatal function, distinguishing maximum potential conductance from regulated operational conductance while recognizing that maximum stomatal conductance reflects the structural investment in stomatal apparatus (density and size) that determines the ceiling on gas exchange rates that physiological regulation can approach but not exceed. The ontological significance lies in maximum stomatal conductance establishing the fundamental gas exchange capacity of leaves, where the product of stomatal density, maximum pore area, and diffusion pathway characteristics determines the maximum possible conductance that represents the anatomical constraint on photosynthesis and transpiration regardless of how favorable environmental conditions may be. The anatomical basis for maximum stomatal conductance includes stomatal density (stomata/mm²), guard cell length (determining maximum pore size), stomatal complex geometry (affecting diffusion pathway), and the integration of these factors across the leaf surface, with maximum conductance proportional to the sum of maximum pore areas divided by leaf area. The measurement of maximum stomatal conductance requires optimal conditions (high light, well-watered, moderate VPD, ambient CO₂) that fully stimulate stomatal opening, with values for Vitis vinifera typically ranging from 0.30-0.50 mol/m²/s depending on cultivar and growth conditions, representing the reference point from which regulatory closure reduces operational conductance. The developmental and environmental factors affecting maximum conductance include growth light intensity (affecting stomatal development), leaf nitrogen status (affecting guard cell function), and CO₂ during development (elevated CO₂ often reducing stomatal density), with these factors determining the anatomical potential that subsequent regulation modulates. Understanding maximum stomatal conductance quality enables assessment of gas exchange capacity, prediction of maximum photosynthetic potential, and evaluation of cultivar differences in anatomical investment affecting water use strategy.
Maximum_Stomatal_Conductance_Quality addresses the anatomical capacity for stomatal gas exchange under optimal conditions. This quality sets the upper limit that regulatory mechanisms modulate. The class supports capacity assessment and cultivar comparison through maximum conductance characterization revealing anatomical gas exchange potential.
Stomatakapazität
anatomical conductance
anatomische Leitfähigkeit
gsmax
maximale Stomatärleitfähigkeit
maximum gs
stomatal capacity
Franks PJ, Farquhar GD (2001) The effect of exogenous abscisic acid on stomatal development, stomatal mechanics, and leaf gas exchange in Tradescantia virginiana. Plant Physiology 125(2): 935-942 Rogiers SY et al. (2009) Stomatal density of grapevine leaves (Vitis vinifera L.) responds to soil temperature and atmospheric carbon dioxide. Australian Journal of Grape and Wine Research 15(1): 9-14
Maximum Stomatal Conductance Quality
In a Barossa Valley Shiraz vineyard, maximum sugar content was achieved in late February Year N+1 (BBCH 89, 62 days post-veraison) when berry Brix stabilized 27.2° ± 0.4° over 18 days (plateau confirmed), total sugars 280 g/L (HPLC:glucose 142 g/L, fructose 138 g/L, ratio 1.03:1), vacuolar hexose estimated 2350 mM combined, osmotic potential -3.7 MPa, producing wine alcohol potential 15.0% (actual 14.6% post-fermentation), under hot Australian conditions (cumulative 1240 degree-days veraison-harvest) enabling high maximum typical for warm-climate Shiraz.
Maximum_Sugar_Content represents the cultivar-specific peak hexose concentration achieved at sugar equilibrium defining the upper limit of berry sweetness attainable through active accumulation and establishing the baseline for wine alcohol potential, quality characteristics, and harvest maturity assessment. This maximum content is typically achieved during late ripening plateau (BBCH 88-89), and is characterized by total berry sugars reaching cultivar-specific ceiling 180-310 g/L glucose-plus-fructose (corresponding to 20-30° Brix refractive index measurement depending on variety and conditions), vacuolar hexose concentrations at maximum sustainable levels 1600-2600 mM combined (800-1300 mM each glucose and fructose approximately equimolar ratio 0.95-1.05:1), refractive index stabilized at cultivar maximum corresponding to Brix 20-30° with high-Brix varieties achieving 26-30°, moderate cultivars 22-26°, and lower types 20-24°, and osmotic potential at extreme values -3.0 to -4.2 MPa representing near-cellular tolerance limits preventing further concentration increases. Biochemical composition at maximum includes glucose typically 90-155 g/L (8-14% w/v, 500-860 mM), fructose similarly 90-155 g/L (500-860 mM), combined hexoses 180-310 g/L (1000-1720 mM total, though vacuolar concentration higher 1600-2600 mM due to compartmentation), residual sucrose minimal <2-8 g/L as invertase activity maintains efficient cleavage, organic acids declined to final 4-10 g/L contributing minor proportion of total soluble solids compared to hexoses 85-95%, amino acids, phenolics, minerals collectively 8-18 g/L contributing remaining Brix measurement, and water content 72-82% fresh weight basis with solutes occupying 18-28% of berry mass. Molecular determinants of maximum include genetic regulation of hexose transporter expression with maximum induction levels determining import capacity ceiling, vacuolar properties (size, membrane area, pH regulation, osmotic tolerance) constraining maximum achievable concentration, metabolic capacity for maintaining cellular homeostasis at extreme osmotic potentials limiting how concentrated vacuole can become before cellular dysfunction, and regulatory set-points determining when downregulation triggered preventing excessive accumulation beyond safe physiological limits. Physiological limits constraining maximum demonstrate solubility limit for combined hexoses in aqueous solution approximately 2800-3200 mM (50-55% w/v) at berry temperature 15-30°C providing absolute physical ceiling well above biological maximum indicating other factors limiting, osmotic stress tolerance with most plant cells unable to maintain function at water potentials below -4.0 to -4.5 MPa setting practical limit, membrane integrity and function compromised at extreme osmotic potentials affecting transport, signaling, and cellular organization, and metabolic dysfunction if sugar concentrations interfere with enzyme activities or cellular processes constraining maximum to levels compatible with continued metabolism. Thermodynamic constraints include maximum achievable concentration gradient limited by coupling to available energy sources with H⁺-ATPase-driven symport theoretically enabling ~1000-fold gradients but practical limits much lower due to inefficiencies, equilibrium distribution determined by electrochemical potentials across membranes with maximum vacuolar concentration reflecting balance of transport energetics and osmotic back-pressure, and energy cost of maintaining extreme gradients requiring substantial ATP investment potentially limiting how high concentrations can rise before energetic costs exceed cellular capacity. Environmental effects on maximum achievable include temperature during ripening with warmer conditions 26-32°C generally enabling higher maximum 24-28° Brix versus cooler 18-24°C producing 20-24° Brix through effects on accumulation kinetics and duration, water availability affecting concentration versus dilution with deficit potentially enhancing maximum through reduced berry expansion 5-15% Brix increase while adequate water maintaining size with slightly lower concentration, solar radiation and canopy microclimate influencing berry temperature and source photosynthesis affecting both accumulation rate and final maximum achieved, and vintage variation substantial with warm years producing 2-5° Brix higher maximum than cool years for given cultivar reflecting cumulative environmental effects on accumulation trajectory. Cultivar genetic variation demonstrates remarkable range with high-Brix varieties: Gewürztraminer 27-30° maximum (2200-2600 mM vacuolar, 240-310 g/L total sugars), Sémillon 24-28° (2000-2400 mM, 210-280 g/L), Muscat 24-27° (2000-2300 mM, 210-270 g/L) moderate cultivars: Chardonnay 22-26° (1800-2200 mM, 190-260 g/L), Cabernet Sauvignon 22-26° (1800-2200 mM, 190-260 g/L), Merlot 23-26° (1900-2200 mM, 200-260 g/L) and lower-Brix types: Pinot Noir cool climates 20-24° (1600-2000 mM, 180-220 g/L), table grapes 18-22° (1400-1800 mM, 160-200 g/L) reflecting genetic differences in transporter capacity, ripening duration, and regulatory programming determining maximum. Wine quality implications include maximum sugar directly determining potential alcohol with conversion efficiency ~0.55-0.60% ethanol per 1° Brix yielding: 20° Brix → 11-12% alcohol, 24° Brix → 13.2-14.4% alcohol, 28° Brix → 15.4-16.8% alcohol affecting wine style, body, and stability, sugar-acid balance at maximum with typical ratios 25-35:1 (Brix:TA g/L) for quality wines requiring appropriate acid levels 5-8 g/L, residual sugar potential for off-dry or sweet styles depending on fermentation management with incomplete fermentation leaving 5-50+ g/L residual creating sweetness perception, and phenolic extraction during fermentation affected by alcohol level with higher sugar producing higher alcohol enhancing phenolic solubility particularly important reds. Harvest timing considerations include maximum sugar achievement indicating physiological maturity ceiling for sweetness, recognition that maximum may occur days-to-weeks before optimal phenolic or flavor maturity particularly reds requiring integrated assessment, flexibility in harvest timing during plateau period where maximum stable 2-4 weeks enabling optimization for non-sugar parameters, and economic factors where high-sugar grapes command premium prices in some markets incentivizing maximum achievement while other markets favor moderate sugars requiring earlier harvest. Viticultural strategies targeting maximum include cultivar selection matching genetic potential to climate and goals, crop load management with appropriate yields 6-12 kg per vine enabling adequate source capacity supporting maximum accumulation, canopy management maintaining healthy productive foliage throughout ripening, irrigation management with adequate water supporting metabolism while moderate deficit potentially enhancing concentration, disease control preserving berry health enabling normal ripening to maximum, and harvest timing flexibility capitalizing on plateau stability optimizing quality parameters beyond sugar alone. Measurement and verification of maximum employs refractometry measuring Brix providing standard industry metric, density measurement (pycnometry) determining specific gravity correlating with sugar concentration, HPLC analysis quantifying glucose and fructose separately confirming total hexose levels, osmometry directly measuring osmotic potential verifying extreme values characteristic of maximum, and sensory evaluation detecting sweetness intensity providing qualitative confirmation of high sugar levels. Exceptions and special cases include botrytized grapes (noble rot) achieving higher apparent Brix through dehydration rather than active accumulation potentially exceeding 30-40° through water loss concentrating sugars, late-harvest wines targeting very high sugars 28-35° requiring extended hang time plus potentially dehydration, ice wine production freezing grapes concentrating sugars in unfrozen portion potentially achieving 35-45° Brix, and raisin production completely desiccating berries achieving 55-70° Brix through extreme dehydration representing physical concentration not active accumulation. Research applications include genetic studies identifying QTLs and genes controlling maximum sugar capacity, physiological investigations characterizing limits and constraints, environmental manipulation studies optimizing conditions for maximum achievement, and breeding programs targeting specific sugar levels for different wine styles and markets ranging from moderate-alcohol table wines to high-alcohol full-bodied or dessert styles.
Maximum_Sugar_Content is distinguished from Sugar_Plateau by representing achieved concentration level (20-30° Brix cultivar-dependent) rather than equilibrium process, from Sugar_Equilibrium by focusing on content measurement rather than flux balance, and from approaching maximum by showing achieved ceiling rather than trajectory. Vacuolar hexose: high-Brix 2200-2600 mM, moderate 1800-2200 mM, lower 1600-2000 mM. Osmotic potential -3.0 to -4.2 MPa. Determines wine alcohol potential. Maximum_Sugar_Content represents cultivar-specific peak sweetness defining harvest maturity and quality potential.
Hexose-Konzentrations-Maximum
Kultivar-Zucker-Obergrenze
Spitzen-Beeren-Süße
cultivar sugar ceiling
hexose concentration maximum
maximale Brix-Erreichung
maximum Brix achievement
peak berry sweetness
Keller M (2020) The Science of Grapevines 3rd Ed pages 227-228, 251-253 maximum sugar and quality Boulton RB, Singleton VL, Bisson LF, Kunkee RE (1996) Principles and Practices of Winemaking pages 28-35 sugar content and fermentation Conde C, Silva P, Fontes N, Dias ACP, Tavares RM, Sousa MJ, Agasse A, Delrot S, Gerós H (2007) Biochemical changes throughout grape berry development and fruit and wine quality Food Res Internat 40:1012-1018 maximum sugar and composition.
Maximum Sugar Content
In a Napa Zinfandel vineyard, maximum sugar loading rate was measured over 9 days in late August Year N+1 (BBCH 85) when hexose import reached 168 μmol g⁻¹ h⁻¹ calculated from Brix trajectory, berry Brix increased from 20.8° to 26.2° (average 0.72% per day, 5.0% per week), cell wall invertase 128 units/g FW, VvHT1 expression 155-fold, vacuolar hexose rose from 1250 to 1680 mM (430 mM gain), berry fresh weight 2.08 g expanding 14%, under hot Napa conditions (daily high 32-36°C near heat stress threshold but sustained maximum briefly).
Maximum_Sugar_Loading_Rate represents the absolute highest velocity of hexose import into berry cells achieved when phloem delivery, invertase activity, membrane transport, and vacuolar sequestration all operate at saturating capacity creating the fastest compositional transformation observed during entire berry development determining maximum achievable sweetening rate under optimal conditions. This maximum rate occurs during peak accumulation phase (BBCH 84-85), typically days 20-28 post-veraison for duration 6-12 days, and is characterized by hexose uptake reaching 100-180 μmol combined glucose and fructose g⁻¹ h⁻¹ representing full capacity of transport machinery, berry Brix increasing 0.55-0.85% per day (3.8-6.0% per week) creating steepest Brix trajectory, daily sugar gain 200-320 mg hexose per berry enabling 2.5-4.5° Brix increase over 6-10 day maximum period, and calculated sugar import flux 140-260 μmol hexose g⁻¹ h⁻¹ total (combining membrane uptake, metabolism, and accumulation) matching or exceeding apoplastic production from invertase indicating transport capacity adequate for maximum substrate delivery. Biochemical basis of maximum rate includes hexose transporter protein density reaching absolute maximum estimated 10-16% of total plasma membrane protein providing unprecedented facilitated diffusion capacity, cell wall invertase activity 90-150 units/g fresh weight producing hexoses from sucrose at rates 130-280 μmol g⁻¹ h⁻¹ potentially exceeding cellular uptake capacity indicating enzyme saturation not limiting, apoplastic hexose concentrations 280-550 mM combined glucose and fructose creating strong driving force for membrane transport with concentrations far above transporter Km values 1.5-10 mM ensuring kinetic saturation, cytoplasmic hexose maintained 65-105 mM through rapid tonoplast clearing preventing accumulation that could reduce plasma membrane uptake gradient, and tonoplast transport 70-140 μmol g⁻¹ h⁻¹ efficiently sequestering cytoplasmic hexoses maintaining compartmentation essential for continued import. Kinetic analysis at maximum rate demonstrates plasma membrane glucose transport Vmax 80-135 μmol g⁻¹ h⁻¹ achieved during maximum with substrate saturation 92-98% (apoplastic glucose 140-280 mM far exceeding Km 1.5-3.0 mM), fructose transport Vmax similarly 70-120 μmol g⁻¹ h⁻¹ with saturation 90-96% (apoplastic fructose 140-270 mM exceeding Km 5-12 mM), combined uptake 150-255 μmol g⁻¹ h⁻¹ limited by transporter density not substrate availability, and comparison with active phase average uptake 70-130 μmol g⁻¹ h⁻¹ showing maximum represents 1.6-2.4 fold enhancement reflecting peak conditions. Phloem delivery supporting maximum rate shows sucrose mass flow 280-500 mg per berry per day enabled by transpiration-driven pressure flow, source photosynthesis 18-32 g CO₂ fixed per vine per day under optimal conditions producing abundant substrate, vine-level allocation 80-95% of photosynthate to fruit during maximum representing strong sink signal, and phloem sap sucrose concentration 450-750 mM maintained throughout diel cycle providing continuous substrate supply. Vacuolar dynamics during maximum show concentration rising 160-250 mM per week (glucose and fructose each 80-125 mM per week increase) representing most rapid accumulation, osmotic potential declining -0.25 to -0.35 MPa per week creating steepest gradient (from -2.2 to -2.8 MPa → -2.4 to -3.2 MPa over maximum period), water import 12-20% berry volume per week driven by osmotic gradient causing rapid berry expansion, and vacuole occupying 96-97% cell volume at maximum nearing absolute limit for mesocarp cells. Molecular mechanisms enabling maximum include gene expression at absolute ceiling 110-165 fold above green berry (VvHT1, VvHT2 combined) representing maximum transcriptional activation, protein synthesis rates 35-65 ng transporter protein per g berry per hour maintaining high levels despite turnover, post-translational modifications potentially optimizing transporter activity through phosphorylation or other changes though modifications in grape incompletely characterized, and regulatory signals including ABA 2000-3200 ng/g, hexose signaling with glucose-6-phosphate 45-75 μmol/g FW creating maximum feed-forward activation. Temperature optimum for maximum rate shows sharp dependence 26-30°C producing highest rates 140-200 μmol hexose g⁻¹ h⁻¹ and 4.5-6.5% Brix per week, moderate warmth 23-26°C enabling 100-150 μmol g⁻¹ h⁻¹ and 3.2-4.8% per week still approaching maximum, cooler conditions 18-23°C limiting to 65-110 μmol g⁻¹ h⁻¹ and 2.2-3.5% per week through reduced kinetics, and heat stress >32-35°C impairing maximum by 35-65% through enzyme denaturation, membrane disruption, and metabolic dysfunction preventing achievement of true maximum. Diurnal variation at maximum shows midday peak 160-250 μmol hexose g⁻¹ h⁻¹ during hours 11AM-4PM when berry temperature optimal and phloem delivery maximum, morning and evening 110-180 μmol g⁻¹ h⁻¹ as conditions moderate but remain favorable, nighttime sustained 70-120 μmol g⁻¹ h⁻¹ representing 45-65% of daytime maximum demonstrating continuous import even at reduced rates, and daily integrated maximum 0.60-0.90% Brix per day averaging diurnal cycle. Energy requirements at maximum include ATP demand for membrane H⁺-ATPases maintaining electrochemical gradients estimated 20-35 μmol ATP g⁻¹ h⁻¹ must be met to support maximum transport, respiratory ATP production from combined hexose (20-40% of import oxidized = 25-65 μmol hexose g⁻¹ h⁻¹ oxidized producing 35-90 μmol ATP g⁻¹ h⁻¹) plus organic acid catabolism (35-60 μmol ATP g⁻¹ h⁻¹) providing total 70-150 μmol ATP g⁻¹ h⁻¹ adequate for transport and biosynthesis, and energetic efficiency where majority imported hexoses accumulate (60-80%) versus respired (20-40%) representing favorable storage ratio. Cultivar variation in maximum rate demonstrates high-capacity varieties achieving 160-220 μmol hexose g⁻¹ h⁻¹ and 5.2-7.0% Brix per week sustained 7-11 days, moderate cultivars showing 120-170 μmol g⁻¹ h⁻¹ and 4.0-5.5% per week for 9-14 days, and lower-capacity types displaying 85-135 μmol g⁻¹ h⁻¹ and 3.0-4.5% per week over 12-18 days reflecting genetic differences in transporter expression, enzyme capacity, and metabolic programming. Quality implications include maximum rate achievement critical for reaching target sugars within growing season particularly cool regions where season limited, duration of maximum affecting total sugar gain with extended maximum beneficial but also indicating potential for excessive sugars if not monitored, coordination between sugar maximum and acid decline potentially producing very rapid compositional change requiring careful harvest timing decisions, and failure to achieve expected maximum indicating environmental stress or management limitations requiring intervention or harvest timing adjustment. Harvest forecasting uses maximum rate measurements projecting final Brix achievement timing 10-16 days advance enabling advance planning, recognition that maximum typically concludes 2-3 weeks pre-harvest as deceleration begins, integration with acid, pH, phenolic monitoring ensuring comprehensive maturity determination, and flexibility to adjust harvest date based on actual versus projected maximum rate performance. Viticultural management during maximum prioritizes maintaining optimal conditions through adequate irrigation, canopy health, disease control, and environmental moderation, avoiding any severe stresses during this brief critical window as impairment dramatically affects final sugar, recognizing maximum as point of no return where berry committed to ripening trajectory determined by maximum rate achievement, and harvest preparation advancing as maximum signals approaching maturity requiring readiness within 2-4 weeks typical time from maximum completion to optimal harvest.
Maximum_Sugar_Loading_Rate is distinguished from Peak_Sugar_Import_Rate (Batch 34) by representing brief absolute maximum (100-180 μmol g⁻¹ h⁻¹, 6-12 days) rather than sustained peak period, from average rates by showing maximum (0.55-0.85% Brix per day) rather than typical (0.22-0.35%), and conceptually similar to Peak_Sugar_Import_Rate providing alternative organizational perspective. Duration 6-12 days BBCH 84-85. Temperature optimum 26-30°C critical. Kinetic saturation achieved. Maximum_Sugar_Loading_Rate represents absolute capacity limit determining achievable sweetening under optimal conditions.
Spitzen-Hexose-Aufnahme-Geschwindigkeit
absolute accumulation maximum
absolutes Akkumulations-Maximum
fastest sugar import flux
maximale Beladungs-Kapazität
maximum loading capacity
peak hexose uptake velocity
schnellster Zucker-Import-Fluss
Keller M (2020) The Science of Grapevines 3rd Ed pages 223-228 maximum rates Hayes MA, Davies C, Dry IB (2007) Isolation, functional characterization, and expression analysis of grapevine hexose transporters J Exp Bot 58:1985-1997 transporter capacity Sweetman C, Deluc LG, Cramer GR, Ford CM, Soole KL (2009) Regulation of malate metabolism in grape berry and other developing fruits Phytochem 70:1329-1344 maximum loading dynamics.
Maximum Sugar Loading Rate
Screening for Mealybug_Resistance_Disposition at Stellenbosch University, South Africa, evaluated vine mealybug (Planococcus ficus) population development and GLRaV-3 transmission across 20 cultivars using controlled infestation assays. Potted vines were infested with 50 adult female mealybugs, with population counts and honeydew production measured at 8 weeks. GLRaV-3 transmission was assessed using viruliferous mealybugs in separate assays with RT-PCR confirmation at 12 weeks post-exposure. Population development varied 3.5-fold among cultivars, from 420 individuals on highly susceptible 'Chenin Blanc' to 120 on less-susceptible 'Pinotage', with honeydew production correlated (r = 0.89) with population size. 'Pinotage' demonstrated moderate Mealybug_Resistance_Disposition with 35% lower adult female weight (0.82 mg versus 1.26 mg on 'Chenin Blanc'), suggesting reduced feeding quality, and 40% lower fecundity (185 versus 310 eggs per female). However, GLRaV-3 transmission rates did not differ significantly among cultivars (62-78%), indicating that even partial mealybug resistance was insufficient to prevent virus transmission under assay conditions. Analysis of phloem sap amino acid composition revealed no consistent differences between relatively resistant and susceptible cultivars. These findings suggest that meaningful Mealybug_Resistance_Disposition for leafroll management may require resistance levels beyond those naturally present in V. vinifera, potentially necessitating introgression from wild Vitis species.
A Mealybug_Resistance_Disposition is an Insect_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that limit feeding, reproduction, population development, or Grapevine leafroll-associated virus transmission by mealybugs (Hemiptera: Pseudococcidae), phloem-feeding insects that have emerged as the most important vectors of grapevine leafroll disease in many viticultural regions. This disposition represents resistance to soft-bodied, waxy-coated insects that colonize protected sites on vines including under bark, on roots, within leaf axils, and inside clusters, making them difficult to detect and control. Multiple mealybug species affect grapevine: the vine mealybug (Planococcus ficus) and grape mealybug (Pseudococcus maritimus) being most significant globally, with obscure mealybug (P. viburni), longtailed mealybug (P. longispinus), and citrus mealybug (Planococcus citri) regionally important. Beyond direct damage from sap extraction and honeydew contamination promoting sooty mold, mealybugs transmit Grapevine leafroll-associated viruses (GLRaVs) in a semi-persistent manner, acquiring and transmitting virus during feeding periods of minutes to hours. Resistance mechanisms may include bark physical properties limiting colonization of protected sites, phloem sap characteristics reducing feeding quality and reproductive success, induced defenses affecting mealybug performance, and volatile emissions recruiting natural enemies. Given the near-universal susceptibility of V. vinifera to GLRaVs and the absence of direct viral resistance, mealybug resistance represents a critical indirect pathway for leafroll disease management. However, strong genetic resistance to mealybugs has not been clearly identified in Vitis germplasm, and management currently relies on cultural practices, biological control, and insecticides.
Mealybug_Resistance_Disposition is_a Insect_Resistance_Disposition. Mealybugs are the primary vectors of Grapevine leafroll-associated viruses, creating critical linkage between this class and Leafroll_Resistance_Disposition. Unlike Xiphinema resistance where strong sources exist in wild Vitis, no strong mealybug resistance has been identified in available germplasm. The cryptic feeding behavior of mealybugs (under bark, in clusters) complicates both management and resistance evaluation. Current leafroll management relies on clean planting material, roguing, and vector control rather than host resistance. Identifying or developing mealybug resistance remains an important but unfulfilled breeding objective.
Pseudococcidae resistance
Pseudococciden-Abwehr
Schmierlaus-Resistenz-Disposition
Wollläuse-Resistenz
leafroll vector resistance
vine mealybug resistance
Daane KM et al. (2012) Biology and management of mealybugs in vineyards, Arthropod Management in Vineyards, Springer Walton VM, Pringle KL (2004) Vine mealybug, Planococcus ficus (Signoret), a key pest in South African vineyards, South African Journal of Enology and Viticulture 25:57-62 Almeida RPP et al. (2013) Ecology and management of grapevine leafroll disease, Frontiers in Microbiology 4:94
Mealybug Resistance Disposition
The mean annual temperature for the major Burgundy viticultural communes documents the subtle thermal gradient along the Côte d'Or — Marsannay (northern Côte de Nuits) 10.8°C, Vosne-Romanée (central Côte de Nuits) 11.0°C, Beaune (central Côte de Beaune) 11.2°C, Chassagne-Montrachet (southern Côte de Beaune) 11.3°C — a total gradient of only 0.5°C over the 50 km length of the Côte that nonetheless corresponds to detectable differences in phenological timing (harvest date advancing 2-3 days per 0.1°C increase in MAT), cultivar performance (the warmer southern communes achieving slightly more consistent Pinot Noir maturity), and wine style (the marginally warmer Beaune and Chassagne communes producing slightly fuller-bodied wines than the cooler Marsannay communes in average vintages). The MAT of 11.0-11.3°C places Burgundy within the cool viticultural climate band that Jones (2006) identified as optimal for Pinot Noir — a cultivar that produces its finest wines in the 10.5-12.5°C MAT range.
A temperature data component that records the arithmetic mean of all daily mean temperatures over the calendar year, expressed in degrees Celsius, providing the broadest single-value thermal characterization of the vineyard site that integrates the warmth of the growing season with the coldness of the dormant season and positions the site within the global framework of viticultural thermal suitability — the world's wine regions spanning a mean annual temperature range from approximately 9°C (coolest commercially viable sites in England, Champagne, and the Mosel) to approximately 20°C (warmest sites in the Swan Valley, inland southern Spain, and the Central Valley of California).
Mean_Annual_Temperature provides the simplest thermal characterization of a viticultural site but must be interpreted with caution because identical MAT values can arise from very different seasonal temperature distributions — a maritime climate with mild winters and cool summers (e.g., Bordeaux, MAT 13°C, January mean 6°C, July mean 21°C, amplitude 15°C) produces the same MAT as a continental climate with cold winters and hot summers (e.g., Mendoza, MAT 13°C, January mean 25°C, July mean 4°C, amplitude 21°C) despite fundamentally different viticultural implications. The global viticultural band as defined by Jones (2006) extends from approximately 12-22°C mean growing season temperature (roughly equivalent to 9-20°C MAT), with the lower boundary set by insufficient heat for fruit ripening and the upper boundary set by excessive heat that prevents the synthesis and retention of quality-defining volatile compounds: within this band, each cultivar occupies a characteristic thermal niche — Pinot Noir and Chardonnay at the cool end (MAT 10-13°C), Cabernet Sauvignon and Merlot in the middle (MAT 13-16°C), and Grenache, Mourvèdre, and Carignan at the warm end (MAT 15-19°C). The climate change signal in MAT is clear and consistent across viticultural regions — Jones et al. (2005) documented MAT increases of 0.5-1.5°C across major wine regions between 1950 and 2000, with the warming trend accelerating since 1980, and the practical consequences already visible in shifting harvest dates (earlier by 2-3 weeks in many European regions), changing cultivar suitability, and the emergence of new viticultural regions at previously too-cool latitudes and elevations.
Jahresmitteltemperatur
MAT measurement
annual mean temperature record
jährliche Durchschnittstemperatur
mittlere Jahrestemperatur
yearly average temperature
Jones GV (2006) Climate and terroir: impacts of climate variability and change on wine, in Macqueen RW and Meinert LD (eds) Fine Wine and Terroir — The Geoscience Perspective, Geological Association of Canada Jones GV et al. (2005) Climate change and global wine quality, Climatic Change 73: 319-343 Gladstones J (1992) Viticulture and Environment, Winetitles
Mean Annual Temperature
Mechanical damage quality control in Sugraone table grape packing requires line speed adjustment and cushioning optimization based on real-time damage monitoring, with target damage levels below 0.5% for premium export packs achieved through foam-padded receiving bins, water flotation sorting that eliminates hard impacts, 40 cm maximum drop heights at all transfer points, and individual berry cushioning in consumer clamshells damage rates exceeding 1% trigger immediate line stoppage for investigation and correction, as mechanical damage in Sugraone's relatively thin skin leads to rapid browning and decay that can render entire pallets unmarketable within 48-72 hours of pack-out.
Mechanical_Damage_Quality represents the damage state resulting from physical forces that exceed tissue strength, causing visible injury to grapevine organs. Mechanical damage encompasses cuts, punctures, abrasions, bruises, and crushing that disrupt tissue integrity through direct physical impact rather than physiological processes, environmental factors, or biological agents. In grape berries, mechanical damage typically manifests as skin wounds that expose flesh to air and microorganisms, subsurface bruising visible as darkened or water-soaked tissue, and crushing that releases juice. Mechanical damage occurs at multiple points in the production chain including insect and bird feeding in the vineyard, impact during harvest (particularly mechanical harvest), compression and abrasion during transport, friction damage from conveyor belts and sorting equipment, and handling injury during packing and display. The susceptibility to mechanical damage depends on skin thickness and strength, flesh firmness, turgor pressure, and the magnitude and type of applied forces. Thick-skinned cultivars like Autumn Royal resist puncture damage better than thin-skinned varieties like Flame Seedless. Turgid fruit is more susceptible to impact bruising than slightly dehydrated fruit, creating a trade-off between freshness and damage resistance. Mechanical damage triggers rapid quality deterioration through browning (enzymatic oxidation of phenolics), juice loss, and microbial colonization of wounds. Assessment of mechanical damage employs visual inspection for surface wounds, gentle palpation detecting subsurface bruising, and documentation of damage type, severity, and location on affected organs.
Mechanical_Damage_Quality specifically addresses injury from physical forces, distinguished from Splitting_Quality (skin rupture from internal pressure), Shriveling_Quality (water loss), and damage from biological or environmental causes. Mechanical damage is typically preventable through careful handling and proper equipment design, making it a target for quality improvement programs. The timing of damage relative to assessment affects visibility, as fresh wounds may be more apparent than partially healed damage conversely, bruise symptoms may develop hours after the causative impact. Documentation of damage should note both the immediate injury and the secondary consequences including browning, juice leakage, and decay initiation.
Handhabungsschaden
Stoßverletzung
handling damage
impact injury
mechanical injury quality
mechanische Beschädigung
physical injury
physische Verletzung
Crisosto CH et al. (2001) Table Grapes Respond to Mechanical Damage. California Agriculture 55(5): 30-31 Rolle L et al. (2012) Instrumental Texture Analysis Parameters as Markers of Table-Grape and Winegrape Quality: A Review. American Journal of Enology and Viticulture 63(1): 11-28
Mechanical Damage Quality
The mechanical harvest specification for Chardonnay in the Riverina (Australia) details operation of a Pellenc Optimum 240 self-propelled over-the-row harvester at 3.5 km/h ground speed with bow rod shaking frequency set to 420 beats per minute and amplitude of 35 mm — calibrated to detach ripe berries (force required approximately 2-3 N) while retaining unripe berries (detachment force >4 N) and minimizing rachis fragments in the harvested material. The onboard destemmer-aspirator removes 85-90% of leaves, petioles, and material other than grape (MOG), while a berry sorting system with optical sensors rejects visually damaged or underripe berries at rates of 3-5% of total throughput, achieving delivered fruit quality with less than 2% MOG content and less than 5% damaged or oxidizing berries.
A harvest method description that documents the technical parameters, operational settings, and quality control procedures for machine harvesting of grapes, specifying the harvester type (over-the-row or tow-behind), beating mechanism (bow rod or trunk shaker), operating speed, shaking frequency and amplitude, destemmer and cleaning system configuration, and fruit condition monitoring protocols that collectively determine the efficiency and gentleness of mechanical grape collection.
Mechanical_Harvest_Specification captures the engineering and operational parameters that determine the quality of mechanically harvested fruit. Modern mechanical harvesters have evolved dramatically from the aggressive trunk shakers of the 1960s-1970s to sophisticated over-the-row machines with adjustable bow rod or flexible finger mechanisms, onboard destemming, optical sorting, and GPS-enabled yield mapping: these technological advances have progressively closed the quality gap between mechanical and hand harvesting for many wine styles, though the fundamental limitation remains — mechanical harvesters cannot deliver intact whole clusters, limiting their use for winemaking processes that require whole-cluster input (traditional Champagne method, whole-cluster fermentation, gentle whole-cluster pressing). Critical operational parameters include shaking frequency and amplitude (which must be calibrated to the specific cultivar's berry detachment force at maturity), ground speed (affecting the duration of shaking per vine), and the configuration of the onboard cleaning system that separates berries from leaves, stems, and other non-grape material.
Maschinenlese-Spezifikation
Traubenvollernter-Protokoll
Vollerntereinsatz-Parameter
harvester operation protocol
machine harvest parameters
mechanical harvester settings
Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 230-260 Pellenc (2023) Optimum series technical specifications Clary CD et al. (1990) Performance of destemmer-crusher and crusher-destemmer machines, American Journal of Enology and Viticulture 41: 71-75
Mechanical Harvest Specification
Mechanical harvesting suitability quality evaluation in Central Valley California compared harvest efficiency and fruit quality outcomes across cultivars processed through standard over-row harvester operations with consistent machine settings. Rubired demonstrated excellent mechanical harvesting suitability quality as a teinturier variety bred specifically for mechanical harvest, exhibiting fruit detachment force of 195 g enabling efficient removal, thick pigmented skin resisting damage, loose cluster architecture minimizing material other than grape, and stable color compounds tolerating oxidation exposure during mechanical handling. Chardonnay showed good mechanical harvesting suitability quality with detachment force of 210 g, adequate skin resilience, and fruit quality maintained through rapid transport and immediate pressing within 4 hours of harvest. Pinot Noir exhibited marginal mechanical harvesting suitability quality with thin skin susceptible to damage, high juice release during harvest averaging 8% versus 3% for Chardonnay, and quality-sensitive phenolic extraction affected by the stem contact and oxidation during mechanical handling, restricting mechanical harvest application to volume-oriented production tiers while premium fruit continued to require hand harvest.
Mechanical harvesting suitability quality is a mechanization suitability quality that inheres in a grapevine cultivar or clone and represents the degree of compatibility with mechanical grape harvesters that employ shaking, striking, or air-blast mechanisms to detach fruit from the vine and convey it to collection systems, requiring cultivar characteristics that enable efficient fruit removal with minimal damage to berries, rachis retention, leaf and petiole contamination, or other quality-compromising consequences of the mechanical detachment and handling processes inherent in machine harvest operations. This quality determines the feasibility of mechanical harvesting as an alternative to hand picking, considering both the technical efficiency of fruit removal and the quality implications for wine production from mechanically harvested fruit that experiences physical impacts, juice release, and oxidation exposure differing fundamentally from hand-harvested material. The ontological significance of mechanical harvesting suitability quality lies in its role as the primary determinant of harvest logistics and cost structure for commercial wine production, with mechanical harvesting enabling single-pass harvest of vineyard blocks in hours rather than days while creating specific quality management requirements that favor certain cultivar characteristics over others. The biological requirements for high mechanical harvesting suitability quality include appropriate fruit detachment force enabling efficient removal at machine operating parameters without excessive berry damage, typically 150-250 g measured by standard pull force methodology cluster architecture with berries sufficiently attached to resist shatter during transport yet loose enough to separate cleanly during harvest skin thickness and integrity adequate to withstand mechanical impacts without excessive juice release and fruit composition characteristics tolerant of the limited oxidation and warming that occurs between detachment and processing. Cultivars with very tight clusters, thin fragile skins, or extremely loose berry attachment prove poorly suited to mechanical harvest, while those combining moderate detachment force, resilient skin structure, and well-ventilated cluster architecture achieve optimal mechanical harvesting outcomes. The interaction between mechanical harvesting suitability quality and wine quality depends critically on post-harvest handling, with rapid transport, inert gas protection, and prompt processing enabling quality outcomes approaching hand-harvest levels for suitable cultivars while extended holding times or inadequate handling can compromise even well-adapted varieties.
Mechanical_Harvesting_Suitability_Quality is_a Mechanization_Suitability_Quality, representing compatibility with mechanical grape harvesting equipment. This class addresses the specific requirements of machine harvest including fruit detachment force, skin resilience, cluster architecture, and oxidation tolerance. The detachment force range of 150-250 g provides quantitative benchmark. Post-harvest handling critically influences quality outcomes and must be considered alongside inherent cultivar suitability. Night harvesting for temperature management has expanded mechanical harvest applicability to quality-sensitive production.
Maschinenlese-Tauglichkeit
Traubenvollernter-Eignung
Vollerntereignung
harvester adaptation
machine harvest compatibility
mechanical picking suitability
mechanical vintage fitness
mechanische Erntekompatibilität
Morris JR (2007) Development and commercialization of a complete vineyard mechanization system. HortTechnology 17:411-420 Clary CD et al. (1990) Performance and economic evaluation of commercial grape harvesters in eastern Washington. American Journal of Enology and Viticulture 41:160-164 Intrieri C et al. (2001) Yield and grape quality assessment of training systems for machine harvesting. Acta Horticulturae 562:287-294
Mechanical Harvesting Suitability Quality
Mechanical leaf removal suitability quality comparison in Napa Valley evaluated Cabernet Sauvignon and Pinot Noir response to air-blast mechanical defoliators operating at standardized settings across phenological timings. Cabernet Sauvignon demonstrated excellent mechanical leaf removal suitability quality with thick, firmly-attached petioles enabling clean abscission at machine air velocities of 180 km/h, robust cluster attachment tolerating operational forces with less than 2% berry loss, and thick berry skin resisting impact damage, achieving equivalent fruit zone exposure (LLN reduced from 2.4 to 1.1) to hand removal at 15% of labor cost. Pinot Noir exhibited poor mechanical leaf removal suitability quality with thin, fragile petioles requiring higher air velocities for removal that caused 12% berry shatter from the loosely-attached clusters, compounded by thin skin susceptibility to abrasion damage affecting 8% of remaining berries, ultimately proving more cost-effective to perform hand removal for this quality-sensitive cultivar where cluster damage represented unacceptable quality and yield loss despite the labor cost premium.
Mechanical leaf removal suitability quality is a mechanization suitability quality that inheres in a grapevine cultivar or clone and represents the degree of compatibility with mechanical leaf removal equipment employing air-blast, roller, or cutting mechanisms to remove foliage from the fruit zone, requiring cultivar characteristics that enable effective leaf detachment without excessive cluster damage, shoot injury, or incomplete removal that would compromise the fruit exposure objectives motivating leaf removal operations. This quality determines whether mechanical leaf removal can achieve fruit zone defoliation outcomes comparable to hand leaf removal at dramatically reduced labor cost, recognizing that mechanical systems apply forces to a defined canopy zone without the selective discrimination possible in manual operations where individual leaves are identified and removed based on their specific contribution to cluster shading. The ontological significance of mechanical leaf removal suitability quality lies in its role enabling economically viable implementation of fruit zone leaf removal practices that demonstrably improve fruit quality through enhanced cluster exposure, reduced disease pressure, and improved spray penetration, but that require prohibitive labor inputs when performed manually across large production areas. The biological requirements for high mechanical leaf removal suitability quality include leaf petiole characteristics enabling clean detachment at appropriate force levels without damage to adjacent cluster rachis or berry pedicels, cluster positioning and architecture that tolerates the mechanical forces applied during leaf removal without berry detachment, shatter, or physical damage, shoot flexibility permitting temporary displacement during machine passage without breakage, and rapid wound healing at petiole abscission zones minimizing disease entry risk. Cultivars with fragile clusters, berries weakly attached to pedicels, or sensitive skin prone to mechanical damage prove poorly suited to mechanical leaf removal regardless of leaf detachment characteristics, while those combining appropriate petiole abscission properties with robust cluster and berry attachment achieve successful mechanical defoliation. The timing of mechanical leaf removal influences suitability assessment, with early-season applications during smaller berry development generally causing less cluster damage than operations at or after véraison when berries have expanded and become more susceptible to mechanical injury.
Mechanical_Leaf_Removal_Suitability_Quality is_a Mechanization_Suitability_Quality, representing compatibility with mechanized fruit zone defoliation equipment. This class addresses the specific requirements of mechanical leaf removal including petiole detachment characteristics, cluster damage tolerance, and berry attachment strength. The timing interaction with cluster development stage significantly influences outcomes. Distinguished from Defoliation_Tolerance_Quality which addresses physiological response to leaf area reduction rather than mechanical removal process compatibility. Critical for economic viability of leaf removal practices at commercial scale.
Entlaubungsmaschinen-Kompatibilität
Maschinelle Entblätterungseignung
automated defoliation compatibility
machine leaf stripping adaptation
mechanical basal leaf removal fitness
mechanical defoliation suitability
mechanische Blattentfernungs-Tauglichkeit
Poni S et al. (2006) Effects of early defoliation on shoot photosynthesis, yield components, and grape composition. American Journal of Enology and Viticulture 57:397-407 Intrieri C et al. (2008) Early defoliation (hand vs mechanical) for improved crop control and grape composition in Sangiovese (Vitis vinifera L.). Australian Journal of Grape and Wine Research 14:25-32 Tardaguila J et al. (2010) Mechanical thinning of Tempranillo and Grenache grapes: effects on berry and must quality. Journal of the Science of Food and Agriculture 90:1508-1514
Mechanical Leaf Removal Suitability Quality
Mechanical pruning suitability quality comparison in Marlborough evaluated long-term performance of Sauvignon Blanc under mechanical box pruning versus traditional cane pruning over a ten-year trial period. Mechanical pruning demonstrated good mechanical pruning suitability quality for this cultivar, achieving equivalent yield averaging 12.5 t/ha compared to 11.8 t/ha for hand pruning, with fruit Brix showing no significant difference and characteristic aromatic compound concentration maintained within acceptable ranges despite the 15-20% higher bud numbers retained by mechanical pruning. Pruning cost declined from NZ$1,850/ha for hand cane pruning to NZ$180/ha for mechanical hedging plus NZ$320/ha for follow-up hand adjustment, achieving net savings of NZ$1,350/ha annually. Long-term vine health assessment showed equivalent trunk disease incidence and maintained vine balance with appropriate crop load management through mechanical fruit thinning where needed. Pinot Noir comparison showed reduced mechanical pruning suitability quality with quality-sensitive markets detecting subtle differences in wine complexity attributed to the altered canopy architecture under mechanical management.
Mechanical pruning suitability quality is a mechanization suitability quality that inheres in a grapevine cultivar or clone and represents the degree of compatibility with mechanical pruning systems including hedging-style box pruning, cutter bar pruning, and minimal pruning approaches that replace or reduce manual dormant pruning labor through mechanized removal of excess cane growth to achieve target bud numbers and canopy dimensions without the individual shoot selection possible in hand pruning operations. This quality determines whether mechanical pruning can achieve acceptable vine balance, yield regulation, and fruit quality outcomes for a given cultivar, recognizing that mechanical pruning imposes standardized cutting planes that cannot account for individual vine variation and that rely on cultivar characteristics and subsequent management to achieve satisfactory results. The ontological significance of mechanical pruning suitability quality lies in its role enabling dramatic labor cost reduction in pruning operations that traditionally represent the largest single labor input in vineyard management, with mechanical pruning reducing per-hectare pruning time from 80-120 hours to 3-5 hours while requiring cultivar characteristics that tolerate the imprecision inherent in mechanized approaches. The biological requirements for high mechanical pruning suitability quality include consistent bud fruitfulness across node positions enabling productive outcomes regardless of which buds are retained by the cutting planes, appropriate vigor response to mechanical pruning that maintains vine balance without excessive compensatory growth, structural shoot characteristics permitting clean cutting without splintering or tearing that could create disease entry points, and fruit quality maintenance under the potentially higher bud numbers and altered canopy architecture resulting from mechanical pruning geometry. Cultivars exhibiting spur-pruning suitability with strong basal bud fruitfulness generally adapt well to mechanical pruning that retains short spur lengths, while those requiring cane pruning with mid-cane bud reliance prove poorly suited to mechanical approaches that cannot select and position individual fruiting canes. The assessment of mechanical pruning suitability quality involves comparison of yield, fruit quality, and vine health outcomes between mechanical and hand-pruned treatments over multiple seasons, with attention to cumulative effects on vine balance and fruitfulness that may not manifest in initial trial years.
Mechanical_Pruning_Suitability_Quality is_a Mechanization_Suitability_Quality, representing compatibility with mechanized dormant pruning operations. This class addresses the specific requirements of mechanical pruning including consistent bud fruitfulness, appropriate vigor response, and structural cutting characteristics. The distinction between cultivars suited to spur versus cane pruning represents key determinant. Long-term effects on vine balance and quality require multi-year assessment. Economic analysis typically shows dramatic labor savings where cultivar suitability permits mechanical pruning adoption.
Heckenschnitt-Kompatibilität
Maschinenrebschnitt-Eignung
Maschinenschnitteignung
automated pruning fitness
hedger pruning suitability
machine pruning compatibility
mechanical winter pruning adaptation
mechanische Rebschnitt-Tauglichkeit
Clingeleffer PR (1993) Vine response to modified pruning practices. In: Pool RM (ed) Proceedings of the Second NJ Shaulis Grape Symposium, Cornell University, pp. 20-30 Poni S et al. (2016) Mechanical winter pruning of grapevine: physiological bases and applications. Scientia Horticulturae 204:88-98 Gatti M et al. (2011) Ground-truthing of remotely sensed within-field variability in a cv. Sangiovese vineyard for selective mechanical harvesting. Australian Journal of Grape and Wine Research 17:146-152
Mechanical Pruning Suitability Quality
A Chardonnay vine in California's Central Valley managed under mechanical box pruning bears a Mechanically_Pruned_Role, with tractor-mounted cutters creating uniform hedgerow at 90 cm height and 45 cm width—the standardized architecture enabling economical production of 15-20 tonnes/hectare for sparkling wine base. A Cabernet Sauvignon vine in Paso Robles serves in a Mechanically_Pruned_Role with mechanical pre-pruning followed by hand spur selection, the combined approach reducing labor while maintaining quality-oriented bud positioning. A Tempranillo vine in La Mancha's industrial-scale production bears a Mechanically_Pruned_Role, the machine pruning essential for economic viability in Spain's largest wine region where hand labor costs would exceed grape value.
A training system role that inheres in a grapevine pruned by mechanical means (typically tractor-mounted sickle bars, circular saws, or reciprocating cutters) rather than hand labor, creating standardized hedgerow architecture through uniform cutting rather than the selective decisions of hand pruning. Mechanical pruning reduces labor costs by 80-95% compared to hand pruning, enabling economically viable production where labor availability is limited or costs are prohibitive, but sacrifices the individual vine assessment and selective bud retention that characterize skilled hand pruning. The resulting vine architecture reflects the cutting implement geometry—typically flat-topped and flat-sided hedgerows with all wood beyond the cutting plane removed regardless of its position, age, or potential fruitfulness. Mechanical pruning is often combined with hand follow-up, where machine cutting performs bulk wood removal and hand labor addresses specific vine situations (dead wood removal, spur position adjustment, disease management). The system is associated with volume production, though mechanical pre-pruning followed by hand finishing has become standard even in quality regions where pure hand pruning would be economically challenging.
Mechanically_Pruned_Role captures the labor-substitution approach that has transformed commercial viticulture economics, particularly in New World regions and European volume-production zones. The distinction between "mechanical pruning" (machine-only) and "mechanical pre-pruning" (machine bulk removal plus hand finishing) is important many quality-focused operations use the latter approach to balance economics with vine-specific management. Mechanical pruning adoption continues expanding as labor availability declines and costs increase, with technology improvements (precision GPS guidance, sensor-based cutting adjustment) partially addressing the quality concerns that limited earlier adoption. The system creates its own vine architecture over time—spur positions develop where wood remains after cutting, fruiting zone concentrates at hedgerow perimeter—that differs from hand-pruned systems but can produce acceptable fruit for appropriate market segments.
Heckenschnitt
Maschinenschnitt
hedge pruning
machine pruning
maschinelle Rebschnitt
mechanical pruning
mechanischer Schnitt
mechanized pruning
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Intrieri C, Poni S (1995) Integrated training system evolution, American Journal of Enology and Viticulture 46: 507-517 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, Academic Press Morris JR (2007) Development of mechanization technology, HortTechnology 17: 414-420
Mechanically Pruned Role
Mechanization suitability quality evaluation in Riverina compared cultivar performance under fully mechanized production including mechanical pruning, shoot positioning, leaf removal, and harvesting. Chardonnay demonstrated excellent mechanization suitability quality with uniform VSP canopy architecture compatible with over-row equipment, cluster positioning within the standard 0.3-0.9 m harvest zone, appropriate fruit detachment force of 180-220 g enabling efficient harvest with minimal rachis retention, and fruit composition showing less than 3% quality-relevant oxidation following mechanical handling. Semillon showed good mechanization suitability quality with similar architectural compatibility though somewhat higher juice oxidation susceptibility requiring enhanced SO₂ management. Gewürztraminer exhibited poor mechanization suitability quality with irregular canopy architecture resisting standardized machine settings, cluster positioning extending outside the harvest zone requiring manual pre-harvest adjustment, excessive fruit detachment force of 380 g causing rachis retention and increased material other than grape, and aromatic compound sensitivity to oxidation compromising wine quality, ultimately justifying premium pricing to offset the manual harvest requirement for this cultivar.
Mechanization suitability quality is a viticultural quality that inheres in a grapevine cultivar or clone and represents the degree of compatibility between the genotype's morphological characteristics, growth habit, and fruiting patterns and the operational requirements of mechanical equipment employed for pruning, harvesting, canopy management, and other vineyard operations that replace or supplement manual labor with machine-based interventions. This quality determines the technical feasibility and economic viability of mechanized production systems for specific cultivar-site-training combinations, recognizing that successful mechanization requires alignment between vine architecture and machine design parameters that cannot always be achieved through management adjustments alone. The ontological significance of mechanization suitability quality lies in its role as an increasingly critical selection criterion in contemporary viticulture where labor availability constraints, cost pressures, and timing requirements drive adoption of mechanical operations across production systems previously dependent on manual labor, with mechanization suitability becoming a practical necessity rather than optional advantage for many production contexts. The biological basis of mechanization suitability encompasses multiple cultivar characteristics interacting with specific machine requirements, including canopy architecture compatibility with mechanical access and operation zones, cluster positioning and attachment strength affecting harvest efficiency and fruit damage, shoot flexibility and positioning response influencing mechanical pruning and canopy management outcomes, and fruit composition sensitivity to the physical impacts associated with mechanical operations. Cultivars exhibiting high mechanization suitability quality share characteristics including relatively uniform canopy architecture permitting standardized machine settings, cluster positioning within defined harvest zones, appropriate fruit detachment force enabling efficient removal without excessive damage, and shoot characteristics amenable to mechanical manipulation without breakage or repositioning failure. The assessment of mechanization suitability quality involves practical trials evaluating machine operation efficiency, fruit quality outcomes following mechanical handling, and economic analysis comparing mechanized versus manual operation costs for specific cultivar-system combinations. Regional viticulture increasingly incorporates mechanization suitability assessment in cultivar evaluation programs, recognizing that varieties requiring manual operations may prove economically unviable regardless of other quality attributes.
Mechanization_Suitability_Quality serves as the parent class for specific operation suitabilities including Mechanical_Pruning_Suitability_Quality, Mechanical_Harvesting_Suitability_Quality, and Mechanical_Leaf_Removal_Suitability_Quality. This class addresses overall mechanization compatibility while child classes capture specific operation requirements. Distinguished from Training_System_Suitability_Quality by the focus on machine compatibility rather than training configuration per se, though the two interact strongly. Increasingly critical for commercial viability in labor-constrained production regions.
Maschinentauglichkeit
Mechanisierungseignung
Vollernter-Kompatibilität
equipment compatibility
machine harvesting compatibility
mechanical operation suitability
mechanische Bearbeitbarkeit
mechanized viticulture adaptation
Morris JR (2007) Development and commercialization of a complete vineyard mechanization system. HortTechnology 17:411-420 Intrieri C, Poni S (1995) Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality of mechanized Italian vineyards. American Journal of Enology and Viticulture 46:116-127 Zabadal TJ et al. (2002) Winter injury to grapevines and methods of protection. Michigan State University Extension Bulletin E2930
Mechanization Suitability Quality
A median vein on a mature leaf at node 6 of Vitis vinifera cv. Pinot Noir in Burgundy, France, measuring 13.2 cm length from petiolar junction to central lobe apex, diameter of 2.4 mm at mid-length tapering to 1.8 mm near apex and 2.8 mm near base, prominent as raised ridge on abaxial surface approximately 0.6 mm above surrounding lamina tissue, appearing as shallow groove on adaxial surface approximately 0.3 mm deep, light green coloration similar to lamina without red pigmentation typical of this variety, moderate pubescence with trichome density similar to surrounding abaxial lamina, containing multiple large xylem vessels visible in cross-section (8-12 vessels of 40-80 μm diameter) supporting transpiration rates up to 3.5 mmol H₂O m⁻² s⁻¹, substantial phloem on adaxial side facilitating photosynthate export measured at peak rates of 0.8 g sucrose per day, thick-walled supporting tissues providing mechanical rigidity resisting blade drooping under leaf weight and wind forces, serving as axis of bilateral symmetry with four pairs of major lateral veins branching at approximately regular intervals (branching angles 45-65 degrees) creating characteristic palmate venation, measured at BBCH stage 75 (berries pea-sized) in late June, representing the primary vascular and structural axis of the leaf blade.
The central primary vein extending from the petiolar junction to the apex of the central terminal lobe in grapevine leaves in Vitis vinifera, characterized by position along the leaf axis of bilateral symmetry dividing the blade into left and right halves, large diameter typically 1.5-3 mm representing the most substantial vein structure, prominent appearance as raised ridge on abaxial surface and depressed groove on adaxial surface, complex vascular anatomy including multiple xylem vessel elements enabling high water transport capacity and extensive phloem supporting bidirectional photosynthate and nutrient transport, substantial mechanical tissues including thick-walled collenchyma and sclerenchyma providing structural support, length typically 8-18 cm varying with leaf size and lobe development, and critical functional roles including serving as primary water and nutrient delivery pathway from petiole to blade, primary photosynthate export route from blade to petiole, main structural support axis bearing mechanical loads, and developmental organizer establishing bilateral symmetry and coordinating lateral vein formation during leaf development. The median vein represents the direct continuation of petiolar vascular traces into the lamina, receiving the main vascular bundles from the petiole and distributing them through branching lateral veins throughout the blade, exhibiting hierarchical organization with first-order lateral veins branching at regular intervals along median vein length creating the characteristic palmate venation pattern, and showing cultivar-specific characteristics including relative length, branching angles, and degree of prominence that contribute to overall leaf architecture and ampelographic identity.
Median vein is distinguished from lateral veins by central position, largest diameter, greatest prominence, and role as primary vascular pathway and structural axis. The median vein exhibits hierarchical organization as the first-order vein from which second-order lateral veins branch, which subsequently branch into third-order and higher-order veins creating the complete reticulate network. Length and prominence show variation with overall leaf size and lobe development, with deeply lobed cultivars showing proportionally longer median veins extending into elongated central lobes. The median vein serves as the primary water transport pathway, with hydraulic conductivity determining maximum transpiration and photosynthesis rates supportable before water deficit develops, and vessel dimensions and organization affecting vulnerability to cavitation under water stress. Photosynthate export capacity depends on phloem cross-sectional area and organization, with median vein representing the primary bottleneck for carbon export from the blade potentially limiting photosynthetic rates if phloem capacity is inadequate. Mechanical properties including stiffness and strength affect blade positioning and movement responses to wind, with median vein rigidity enabling controlled blade deflection that reduces mechanical stress while maintaining reasonable orientation for light capture. Developmental role involves establishing bilateral symmetry and organizing lateral vein formation, with median vein development preceding and guiding lateral vein initiation during early leaf development.
Hauptader
Mittelrippe
central vein
midrib
primary vein
zentrale Vene
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 58-89 Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 56-94.
Median Vein
The medieval name Noirien appears in Burgundian documents from the 14th century onward, with a 1375 record from the cellars of the Dukes of Burgundy referencing vins de noirien from the Côte de Beaune — ampelographic analysis and geographic continuity strongly support identification with modern Pinot Noir, making this one of the most securely resolved medieval cultivar names, while contemporary medieval names such as Fromenteau (documented in Champagne from the 13th century, tentatively identified as Pinot Gris) and Gouais (documented from the 9th century, confirmed as modern Gouais Blanc through both morphological and molecular evidence) illustrate the range of resolution achievable for medieval cultivar designations.
A historical name documented in sources from the medieval period (approximately 5th-15th centuries CE) including monastic records, feudal estate inventories, tithe registers, municipal regulations, and early vernacular agricultural texts, designating a grapevine cultivar whose identity can sometimes be tentatively linked to modern cultivars through continuity of regional cultivation and comparative ampelographic analysis.
Medieval_Name documents the period when European cultivar nomenclature began to stabilize through the record-keeping practices of monasteries, feudal estates, and municipal authorities. The medieval period is critical for grapevine nomenclatural history because it bridges the gap between ancient names (largely unresolvable) and early modern names (increasingly resolvable): monastic vineyards in particular maintained cultivar continuity over centuries, providing genealogies of vine planting that sometimes enable tentative identification of medieval names with modern cultivars. The Cistercian and Benedictine orders played a particularly significant role in medieval viticultural documentation across Burgundy, the Rhineland, and central Europe.
Sortenname des Mittelalters
feudal-era variety name
klösterliche Rebbezeichnung
medieval cultivar name
mittelalterlicher Sortenname
monastic grape designation
Rézeau P (2014) Dictionnaire des noms de cépages de France, CNRS Éditions Dion R (1959) Histoire de la vigne et du vin en France des origines au XIXe siècle, Paris Lavalle J (1855) Histoire et statistique de la vigne et des grands vins de la Côte d'Or
Medieval Name
Medium body quality in village-level Burgundy Pinot Noir (12.5% alcohol, dry, body rating 5/10) provides the structural sweet spot where sufficient weight supports the variety's cherry-earth-spice complexity without overwhelming its inherent elegance, enabling pairing with the broadest range of foods from roast chicken through salmon through lighter beef preparations, and serving as the default wine style for mixed dinner party contexts where diverse dishes and preferences must be accommodated.
Medium body quality is a body quality that represents moderate perceived weight and balanced palate presence, characterizing the versatile mouthfeel that provides sufficient substance for flavor support without overwhelming delicacy or demanding excessive attention, enabling broad food pairing compatibility and consumption flexibility. Within the BFO framework, this quality specifically addresses the middle range of the body spectrum, distinguishing balanced weight from the extremes of light and full body while recognizing that medium body represents the most versatile and food-friendly category spanning diverse wine styles and regions. The ontological significance lies in medium body providing the structural sweet spot for everyday wine consumption, where sufficient weight supports flavor complexity without creating the palate fatigue of full-bodied wines or the insubstantial impression of very light wines, making medium-bodied wines the most broadly useful category for food pairing and mixed-occasion consumption. The perceptual characteristics of medium body include moderate coating of oral surfaces, balanced resistance to oral manipulation, comfortable swallowing without either watery thinness or heavy persistence, and the sense of complete but not excessive substance supporting flavor perception. The compositional profile associated with medium body typically includes moderate alcohol (11.5-13.5%), variable residual sugar depending on style, moderate extract, and balanced acidity, with the specific combination varying among wine types while producing comparable weight impression. The variety and regional associations of medium body span the majority of table wines globally, including most Burgundy and Oregon Pinot Noir, majority of white Burgundy and premium Chardonnay, standard Bordeaux and Côtes du Rhône, most Spanish Tempranillo, and countless other everyday wines, with medium body representing the default expectation for table wine unless style specifically calls for lighter or fuller character. Understanding medium body quality enables recognition of the versatile middle ground, appropriate quality assessment for the broadest wine category, and food pairing recommendations leveraging the flexibility of medium-bodied wines.
Medium_Body_Quality addresses the middle range of the body spectrum, representing the most versatile and broadly applicable wine category. This quality provides the structural balance enabling broad food compatibility and consumption flexibility. The class supports recognition of the default body expectation for table wines and evaluation of wines intended for everyday versatility rather than stylistic extremes.
ausgewogenes Gewicht
balanced weight
medium weight
mittelschwerer Körper
mittlerer Körper
moderate Fülle
moderate body
versatile body
Jackson RS (2014) Wine Science: Principles and Applications, 4th Ed. Academic Press, San Diego, pp. 831-887 Harrington RJ (2008) Food and Wine Pairing: A Sensory Experience. John Wiley & Sons, Hoboken, pp. 45-78
Medium Body Quality
A Cabernet Sauvignon vine at 4,000 vines/hectare (2.2 m × 1.1 m) in Napa Valley hillside planting bears a Medium_Density_Planted_Role, the spacing enabling tractor operations while maintaining density appropriate to moderate-vigor hillside conditions. A Malbec vine at 5,000 vines/hectare in Mendoza's restructured quality vineyards serves in a Medium_Density_Planted_Role, the density representing modernization from traditional wide-spaced parral to more intensive production following European models. A Sauvignon Blanc vine at 3,500 vines/hectare in Marlborough bears a Medium_Density_Planted_Role, the spacing optimized for mechanical management of the variety's vigorous growth on fertile alluvial soils.
A vine spacing role that inheres in a grapevine planted at moderate density (typically 3,000-6,000 vines per hectare), with spacing balancing vine competition, canopy management flexibility, and mechanization access in configurations common to quality-focused New World production and modernized European vineyards. Medium-density planting represents a compromise between the intensive high-density tradition (>6,000 vines/hectare) and the extensive low-density approach (<3,000 vines/hectare), enabling tractor access for most operations while maintaining sufficient vine numbers for site coverage and vigor management through competition. Common medium-density configurations include 2.0-2.5 m row spacing (accommodating standard tractors) with 1.0-1.5 m vine spacing within rows, yielding densities of 2,700-5,000 vines/hectare depending on specific dimensions. Medium-density vineyards can accommodate diverse training systems—from single-curtain VSP to moderate divided canopy architectures—with the inter-row space providing operational flexibility while within-row spacing maintains reasonable per-vine vigor limitation. This density range dominates contemporary quality wine production outside traditional high-density regions, representing the practical optimum for mechanized operations and moderate-vigor sites.
Medium_Density_Planted_Role captures the pragmatic density range dominating quality commercial viticulture, balancing operational efficiency with physiological goals. The category spans considerable variation (3,000-6,000 vines/hectare represents a 2× density range), reflecting the diversity of situations where neither high-density intensity nor low-density extensivity is optimal. Medium density enables most standard viticultural equipment, allows single-operator management of substantial acreage, and provides adequate vigor control for moderate sites without the extreme competition of high-density planting. The choice within the medium-density range reflects site-specific vigor expectations—higher density (5,000+) for lower-vigor sites, lower density (3,000-4,000) for more vigorous conditions—with training system selection coordinated to match density with canopy management requirements.
Standarddichte
conventional spacing
konventionelle Bepflanzung
medium-density planting
mittlere Dichte
moderate Pflanzung
moderate density
standard density
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles Reynolds AG et al. (1994) Vine spacing effects, American Journal of Enology and Viticulture 45: 444-459 Keller M (2015) The Science of Grapevines, Academic Press
Medium Density Planted Role
A medium fruiting cane on Vitis vinifera cv. Pinot Noir in Burgundy, France, retained at 8 nodes spanning 85cm along fruiting wire in traditional vertical cordon system, selected from moderately vigorous vine in limestone soil showing 420g pruning weight, positioned horizontally along wire with upward-facing buds, expected to produce 7-8 fruiting shoots bearing 13-15 clusters totaling 3.2-3.8kg fruit, managed conservatively through cluster thinning to one cluster per shoot (7-8 clusters remaining) and minimal canopy manipulation beyond shoot positioning and tip removal, achieving premium fruit quality through low yields, optimal fruit exposure, and balanced vine physiology supporting complete phenolic maturation.
A fruiting cane retained at moderate length during winter pruning, typically 6-10 nodes, characterized by intermediate extent along trellis (65-110cm), moderate bud count providing balanced crop potential, suitability for both cane-pruned and transitional systems, and production of manageable shoot numbers supporting quality-oriented viticulture without excessive canopy complexity. Medium fruiting canes represent conservative crop load approach favoring fruit quality over maximum yield, appropriate for moderate-vigor vineyards, premium wine production systems, cultivars with high natural fruitfulness requiring less aggressive bud retention, and situations where labor limitations constrain intensive canopy management capacity, providing reliable production with reduced inputs compared to long cane systems.
Medium fruiting canes suit quality-focused wine production, particularly in traditional European regions and premium New World sites, where moderate crop loads, conservative bud retention, and balanced vine physiology receive priority over yield maximization. The 6-10 node range provides adequate crop for economic viability while maintaining excellent fruit quality potential, simplifies canopy management through reduced shoot density, decreases disease pressure in moderate-density canopies, and supports sustainable viticulture through lower input requirements. Many iconic wine regions employ medium fruiting canes as standard practice (Burgundy, Barolo, Brunello, premium Napa Cabernet), recognizing that quality objectives require restraint in bud retention even when vine vigor could support higher counts.
intermediate fruiting cane
mid-length cane
mittelgroße Rute
mittellange Fruchtrute
moderate length cane
Based on Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Bravdo et al. (1985) "Effect of crop level and crop load on growth, yield, must and wine composition, and quality of Cabernet Sauvignon" American Journal of Enology and Viticulture 36=125-131.
Medium Fruiting Cane
Cabernet Sauvignon hermaphrodite flowers at BBCH 61-65 exhibit medium styles 0.52-0.66 mm connecting ovaries 0.88-1.02 mm to stigmas 0.42-0.52 mm, with medium styles representing 26-32% of total pistil length (2-2.3 mm), containing well-developed transmitting tract tissue with elaborate extracellular matrix visible in toluidine blue-stained sections as intensely staining central canal, supporting pollen tube growth at typical rates of 15-22 μm per hour and achieving 50-58% fruit set under favorable bloom conditions. Chardonnay shows medium styles 0.48-0.62 mm (averaging 0.55 mm across diverse sites and vintages) representing the typical hermaphrodite style length, with styles maintaining consistent proportions (28-32% of pistil length) across variable growing conditions demonstrating developmental stability of style dimensions within the medium range. Merlot medium styles 0.5-0.68 mm support successful pollen tube transit from stigma surface through transmitting tract to ovary locules in 52-84 hours (temperature-dependent, faster at 24-26°C, slower at 18-20°C), with pollen tube growth studies using aniline blue fluorescence showing 2-5 pollen tubes per style in successfully fertilized flowers, demonstrating that medium style dimensions accommodate multiple simultaneous pollen tube passages. Pinot Noir comparison of medium styles (0.48-0.64 mm) versus rare short styles (0.38-0.44 mm, <5% of flowers showing developmental variation) reveals that medium styles achieve 48-56% fruit set versus 42-48% for short styles, suggesting that style length within and slightly below the medium range has modest effects on fertilization success with medium length providing optimal performance.
A medium style is a style exhibiting typical intermediate length range of 0.45-0.7 mm at maturity, characteristic of hermaphrodite flowers with functional pistils and representing the normal style length range supporting efficient pollen tube growth and successful fertilization in self-compatible reproductive systems. Medium styles are characterized by= (1) length typically 0.45-0.7 mm at anthesis, representing the standard range for hermaphrodite cultivated V. vinifera (2) diameter typically 0.18-0.35 mm creating well-proportioned cylindrical morphology (3) well-developed transmitting tract tissue with elaborate extracellular matrix rich in pectins and glycoproteins, specialized elongated transmitting tract cells with characteristic morphology, and central stylar canal providing clear pathway for pollen tube growth (4) connection between functional ovary (typically 0.7-1.1 mm) and functional stigma (typically 0.35-0.55 mm) creating well-proportioned pistil (total length typically 1.8-2.5 mm) with medium style representing 20-35% of total pistil length. Medium styles represent the evolutionarily optimized length range for Vitis hermaphrodite flowers, balancing competing requirements= sufficient length to provide adequate spatial separation between stigma and ovary allowing distinct developmental and functional specialization of these organs adequate transmitting tract length to support effective pollen tube guidance and nutrition during pollen tube transit not excessively long to avoid unnecessary pollen tube growth distance and time that could reduce fertilization efficiency or increase vulnerability to environmental perturbations during pollen tube growth. Medium styles develop through normal pistil development in hermaphrodite flowers= style primordium initiation (BBCH 51-53) establishing the intermediate pistil zone style elongation (BBCH 53-57) creating separation between ovary and stigma primordia transmitting tract differentiation (BBCH 55-59) producing specialized tissues final maturation (BBCH 59-61) achieving functional capacity. Medium style length shows modest cultivar-specific variation within the medium range (some cultivars averaging 0.48-0.58 mm, others 0.58-0.68 mm) reflecting genetic variation in pistil proportions, and shows some environmental modulation with favorable growing conditions during pistil development (BBCH 52-57) potentially supporting slightly longer styles within the medium range while stress conditions may produce styles toward the shorter end of medium range. The functional performance of medium styles is optimal for hermaphrodite reproduction= medium-length transmitting tract (0.45-0.7 mm plus additional ovary and stigma tissue totaling 1.5-2.5 mm pollen tube pathway) supports pollen tube growth requiring 48-96 hours at favorable temperatures (20-25°C), providing adequate but not excessive time for pollen tube development and sperm cell delivery medium style dimensions support 3-6 pollen tubes growing simultaneously through the transmitting tract per successful pollination event, with transmitting tract diameter (0.06-0.12 mm central canal) allowing multiple tubes while maintaining guidance efficiency hermaphrodite flowers with medium styles achieve typical fruit set rates of 40-60% under favorable bloom conditions, demonstrating that medium style dimensions support high fertilization efficiency.
Medium_Style represents the typical hermaphrodite style length range and is distinguished from Short_Style (0.7 mm, enhanced length in functionally female/pistillate flowers). Medium styles represent evolutionarily optimized dimensions balancing pollen tube pathway requirements with efficient fertilization. Sub-classification into Moderate and Standard medium styles reflects variation within the medium range. Understanding medium styles is essential for defining normal hermaphrodite pistil morphology, for comparative studies of style length variation across flower types, and for establishing baseline reproductive anatomy supporting successful self-pollination.
Standard-Stempelgriffel
mittellanger Griffel
normaler Stylus
standard style
typical style
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 101-106 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 93-98 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-105.
Medium Style
Freedom (1613C × Dog Ridge) demonstrates strong M. arenaria resistance (Rf 0.3-0.5) in sandy loam vineyards of the Lodi AVA, California, where M. arenaria co-occurs with M. incognita in mixed populations infesting Zinfandel. Ramsey (V. champinii) shows consistent M. arenaria resistance (Rf 0.2-0.4) across race 1 and race 2 in UC Davis greenhouse screening trials. Harmony (1613C × Dog Ridge) provides M. arenaria resistance supporting Cabernet Franc on sandy soils in Virginia's Monticello AVA, where M. arenaria populations average 200-400 juveniles per 250 cm³ soil in untreated vineyard blocks.
A nematode-resistance rootstock role that inheres in a grafted rootstock by virtue of its specific capacity to resist reproduction by Meloidogyne arenaria (Neal) Chitwood, the peanut or northern root-knot nematode. M. arenaria is widely distributed in sandy soils of temperate and subtropical viticultural regions, producing root galls and giant cells similar to other Meloidogyne species but with distinct host range and virulence patterns. Two races are recognized: race 1 (the type race, reproducing on peanut) and race 2 (reproducing on pepper and tomato). Resistant rootstocks prevent M. arenaria reproduction through hypersensitive response, with the genetic basis partially overlapping but not identical to M. incognita and M. javanica resistance. Rf values below 1.0 indicate resistance V. champinii-derived rootstocks typically show Rf values of 0.2-0.6 for M. arenaria.
Meloidogyne_Arenaria_Resistant_Rootstock_Role is particularly relevant for southeastern United States viticulture (Virginia, North Carolina, Georgia) where M. arenaria is often the predominant or co-dominant root-knot nematode species. The species' distribution extends into cooler climates than M. incognita or M. javanica, making M. arenaria resistance important for expanding viticulture in mid-Atlantic and southeastern regions. V. champinii-derived rootstocks generally provide effective M. arenaria resistance, though screening confirms that resistance to one Meloidogyne species does not guarantee resistance to all species.
M. arenaria resistant rootstock
Meloidogyne arenaria resistant rootstock
Meloidogyne-arenaria-resistente Unterlage
gegen Erdnuss-Wurzelgallennematoden resistente Unterlage
northern root-knot nematode resistant rootstock
peanut root-knot nematode resistant rootstock
McKenry MV, Anwar SA (2006) Nematode and grape rootstock interactions, Journal of Nematology 38(3): 312-318 Sasser JN, Carter CC (1985) Overview of the International Meloidogyne Project, in An Advanced Treatise on Meloidogyne Vol 1, NC State University Graphics pp.: 19-24 Ferris H, McKenry MV (1975) Relationship of grapevine yield and growth to nematode densities, Journal of Nematology 7: 295-304
Meloidogyne Arenaria Resistant Rootstock Role
Ramsey (Salt Creek, V. champinii) demonstrates strong M. incognita resistance across all four races with Rf values of 0.2-0.4, and is the predominant rootstock for table grape production (Thompson Seedless, Flame Seedless) on sandy soils in the Coachella Valley, California, where M. incognita populations exceed 500 juveniles per 250 cm³ soil. Harmony (1613C × Dog Ridge) provides M. incognita resistance (Rf 0.5-0.8) to Chardonnay and Merlot on infested sites in the Central Coast AVA. Freedom (1613C × Dog Ridge) shows resistance to races 1 and 3 (Rf 0.3-0.5) but reduced resistance to race 4 (Rf 1.5-2.0) in UC Davis screening trials.
A nematode-resistance rootstock role that inheres in a grafted rootstock by virtue of its specific capacity to resist reproduction by Meloidogyne incognita (Kofoid and White) Chitwood, the southern root-knot nematode. M. incognita is a sedentary endoparasite that induces giant cells in root vascular tissue, forming characteristic galls (3-10 mm diameter) that disrupt water and nutrient transport. Resistant rootstocks prevent giant cell formation through hypersensitive response (HR) localized cell death within 24-48 hours of juvenile penetration, production of reactive oxygen species and phenolic compounds at infection sites, and failure to establish the feeding site cellular reprogramming essential for nematode development. Resistance is race-specific, with four recognized M. incognita races (1-4) differing in host range rootstock resistance may be effective against some races but not others. Reproduction factor (Rf) values below 1.0 indicate resistance, with highly resistant genotypes showing Rf values of 0.1-0.3.
Meloidogyne_Incognita_Resistant_Rootstock_Role is the most economically important nematode resistance for warm-climate viticulture, as M. incognita is the predominant root-knot nematode species in California, Australia, South Africa, and Mediterranean regions. Race-specific resistance complicates rootstock selection genetic analysis has identified the N gene from Harmony as conferring race 1 and 3 resistance, while additional QTLs from V. champinii provide broader resistance. Soil fumigation phase-outs have increased reliance on resistant rootstocks as the primary M. incognita management tool.
M. incognita resistant rootstock
Meloidogyne incognita resistant rootstock
Meloidogyne-incognita-resistente Unterlage
gegen Südlichen Wurzelgallennematoden resistente Unterlage
southern root-knot nematode resistant rootstock
Cousins P, Walker MA (2002) Genetics of resistance to Meloidogyne incognita in crosses of grape rootstocks, Theoretical and Applied Genetics 105: 802-807 McKenry MV, Anwar SA (2006) Nematode and grape rootstock interactions, Journal of Nematology 38(3): 312-318 Ferris H, McKenry MV (1975) Relationship of grapevine yield and growth to nematode densities, Journal of Nematology 7: 295-304
Meloidogyne Incognita Resistant Rootstock Role
Ramsey (Salt Creek, V. champinii) demonstrates high M. javanica resistance (Rf 0.3-0.5) in South African Cape Winelands vineyards where M. javanica is the predominant root-knot species on sandy granitic soils, supporting Chenin Blanc and Pinotage production. Dog Ridge (V. champinii) shows strong M. javanica resistance (Rf 0.2-0.4) in Israeli Negev Desert table grape production. Paulsen 1103 (V. berlandieri × V. rupestris), while not specifically bred for nematode resistance, shows moderate M. javanica tolerance (Rf 1.0-2.0) in Australian Riverland conditions, though populations can build to damaging levels over 10-15 years.
A nematode-resistance rootstock role that inheres in a grafted rootstock by virtue of its specific capacity to resist reproduction by Meloidogyne javanica (Treub) Chitwood, the Javanese root-knot nematode. M. javanica is a polyphagous sedentary endoparasite distributed globally in tropical and subtropical viticultural regions, inducing root galls similar to M. incognita but typically at warmer optimal temperatures (25-30°C versus 20-25°C for M. incognita). Resistant rootstocks block M. javanica reproduction through hypersensitive response mechanisms, with resistance often conferred by similar or identical genetic loci as M. incognita resistance. However, resistance specificity differs some rootstocks resistant to M. incognita are susceptible to M. javanica and vice versa. Resistance screening uses Rf thresholds, with values below 1.0 indicating resistance highly resistant genotypes show Rf values of 0.1-0.5 for M. javanica.
Meloidogyne_Javanica_Resistant_Rootstock_Role addresses a species particularly important in South African, Australian, and Middle Eastern viticulture where M. javanica often predominates over M. incognita. The genetic basis of M. javanica resistance overlaps substantially with M. incognita resistance, and most V. champinii-derived rootstocks confer resistance to both species. However, independent resistance screening against M. javanica is essential, as some rootstocks (e.g., certain Harmony selections) show differential responses to the two species.
Javanese root-knot nematode resistant rootstock
M. javanica resistant rootstock
Meloidogyne javanica resistant rootstock
Meloidogyne-javanica-resistente Unterlage
gegen Javanischen Wurzelgallennematoden resistente Unterlage
McKenry MV, Anwar SA (2006) Nematode and grape rootstock interactions, Journal of Nematology 38(3): 312-318 Saucet SB et al. (2016) Resistance to root-knot nematodes Meloidogyne spp. in woody plants, New Phytologist 211: 41-56 Storey SG et al. (2017) Grapevine cultivar and rootstock combinations for South African conditions, South African Journal of Enology and Viticulture 38(2): 219-227
Meloidogyne Javanica Resistant Rootstock Role
Systematic screening for Meloidogyne_Resistance_Disposition at the University of California Kearney Agricultural Research and Extension Center evaluated 55 rootstock genotypes against a mixed Meloidogyne population (predominantly M. incognita with M. javanica) using replicated pot assays. Vines were grown in inoculated soil (10,000 eggs per pot) for 4 months, then assessed for galling severity (0-10 scale), eggs per gram root, and root system health. Susceptible 'AXR#1' (V. vinifera × V. rupestris) exhibited severe galling (index 8.5), high reproduction (45,000 eggs/g root), and stunted, dysfunctional root systems. The resistant 'Freedom' (1613C × Dog Ridge) demonstrated strong Meloidogyne_Resistance_Disposition with minimal galling (index 0.8), low reproduction (850 eggs/g root, 98% reduction), and healthy root development. Histological examination revealed that J2 penetration occurred equally on resistant and susceptible roots, but on 'Freedom', attempted giant cell induction triggered hypersensitive cell death visible as localized browning within 48 hours, preventing establishment of functional feeding sites. Nematodes on resistant roots either died or developed into small, non-reproductive males rather than egg-producing females. This resistance mechanism demonstrates that Meloidogyne_Resistance_Disposition operates through post-penetration incompatibility disrupting the intimate host-parasite interaction required for nematode reproduction.
A Meloidogyne_Resistance_Disposition is a Nematode_Resistance_Disposition that inheres in a Vitis plant root system and is realized through defense processes that limit infection, giant cell formation, gall development, or reproduction by root-knot nematodes of the genus Meloidogyne, sedentary endoparasites that induce characteristic root galls and cause significant damage to grapevine root systems worldwide. This disposition represents resistance to nematodes with a complex host-parasite interaction involving invasion of root tips by infective second-stage juveniles (J2), migration to the vascular cylinder, induction of specialized feeding sites (giant cells) through secretion of effector proteins, and completion of the life cycle within galled root tissue. On susceptible roots, Meloidogyne feeding induces formation of multinucleate giant cells that function as metabolic sinks providing nutrition to the developing female, accompanied by surrounding cell hyperplasia creating the visible root galls (knots). Heavy infestation disrupts root function, reduces water and nutrient uptake, and predisposes roots to secondary pathogen infection. Resistance mechanisms preventing or limiting root-knot nematode damage include hypersensitive response killing cells at initial J2 penetration sites, incompatibility preventing giant cell induction or maintenance, and failure to support nematode development to reproductive maturity. Three Meloidogyne species are particularly important in viticulture: M. incognita (southern root-knot nematode), M. javanica (Javanese root-knot nematode), and M. arenaria (peanut root-knot nematode), with resistance to each species potentially involving different genetic factors. Rootstock breeding has successfully incorporated Meloidogyne resistance from various Vitis species, with resistant rootstocks essential for productive viticulture in warm-climate regions with endemic root-knot nematode populations.
Meloidogyne_Resistance_Disposition is_a Nematode_Resistance_Disposition. Root-knot nematodes require induction of giant cells for feeding and reproduction, making disruption of this process the primary resistance mechanism. Child classes address resistance to specific Meloidogyne species (M_Incognita_Resistance_Disposition, M_Javanica_Resistance_Disposition, M_Arenaria_Resistance_Disposition) as different genetic factors may confer species-specific resistance. Resistance-breaking Meloidogyne populations have been documented, particularly for M. arenaria, highlighting the importance of deploying multiple resistance sources and monitoring nematode population responses.
Meloidogyne immunity
Meloidogyne-Resistenz
RKN resistance
Wurzelgallennematoden-Resistenz-Disposition
Wurzelknoten-Nematoden-Abwehr
root-knot nematode resistance
Ferris H et al. (2012) Reflections on plant and soil nematode ecology: past, present and future, Journal of Nematology 44:115-126 Anwar SA, McKenry MV (2002) Incidence and population density of plant-parasitic nematodes infecting vegetable crops and associated yield losses in Punjab, Pakistan, Pakistan Journal of Zoology 44:327-333 Williamson VM, Kumar A (2006) Nematode resistance in plants: the battle underground, Trends in Genetics 22:396-403
Meloidogyne Resistance Disposition
Membrane lipid adjustment quality comparison across grapevine species and cultivars established the contribution of lipid modification to interspecific differences in freezing tolerance. Lipid analysis of bud and cane tissues at summer growth and maximum winter acclimation quantified the seasonal change in fatty acid composition and its relationship to hardiness. Vitis riparia, showing maximum hardiness of -35°C, demonstrated excellent membrane lipid adjustment quality with the desaturation ratio (unsaturated/saturated fatty acids) increasing from 2.1 in summer to 3.4 in winter, representing 62% enhancement of membrane unsaturation. Linolenic acid (18:3) content increased from 28% to 45% of total fatty acids, with the accumulation of this trienoic acid strongly correlated with hardiness (r² = 0.78). Vitis vinifera cv. Chardonnay with maximum hardiness of -23°C showed moderate membrane lipid adjustment quality with desaturation ratio increasing from 1.9 to 2.6 (37% enhancement) and linolenic acid from 22% to 32%. Vitis vinifera cv. Merlot with maximum hardiness of only -20°C exhibited poor membrane lipid adjustment quality with desaturation ratio increasing only from 1.8 to 2.2 (22% enhancement) and linolenic acid from 20% to 26%. The membrane lipid adjustment quality assessment established that enhanced fatty acid desaturation capacity contributes significantly to the superior cold hardiness of native American Vitis species compared to V. vinifera, identifying lipid metabolism as a target for genetic improvement of cold tolerance in interspecific breeding programs.
Membrane lipid adjustment quality is a cryoprotection quality that inheres in a grapevine organism and represents the capacity to modify the fatty acid composition and molecular organization of cellular membranes during cold acclimation in ways that maintain membrane fluidity and prevent damaging phase transitions at low temperatures, with increased unsaturation of membrane phospholipids and altered sterol content enabling continued membrane function and preventing the membrane rupture that would occur if rigid, gel-phase membranes were subjected to the mechanical stresses of ice formation and cellular dehydration. This quality addresses the membrane-level adaptation required for cell survival during freezing, recognizing that membranes represent primary sites of freezing injury where the physical changes in lipid bilayer properties at low temperatures can cause membrane failure unless lipid composition is adjusted to compensate for temperature effects on fluidity. The ontological significance of membrane lipid adjustment quality lies in the essential role of membrane integrity for cell survival, with the maintenance of membrane semi-permeability and barrier function depending on appropriate lipid composition that must change seasonally to maintain function across the temperature range from summer growth to winter dormancy. The biological basis of membrane lipid adjustment encompasses the activity of fatty acid desaturase enzymes that introduce additional double bonds into existing membrane lipids, increasing unsaturation and maintaining fluidity at lower temperatures the selective incorporation of unsaturated fatty acids into newly synthesized phospholipids during acclimation and the modification of sterol content and composition that affects membrane physical properties independent of fatty acid saturation. The degree of fatty acid unsaturation shows characteristic increases during cold acclimation quantified by the double bond index or desaturation ratio, with values increasing by 30-50% in cold-hardy genotypes compared to more modest increases in cold-sensitive types. The specific fatty acids most affected include the conversion of oleic acid to linoleic acid and linoleic to linolenic acid through the action of ω-6 and ω-3 desaturases, with the accumulation of trienoic fatty acids particularly important for low-temperature membrane function. The kinetics of lipid adjustment show a time course of days to weeks, intermediate between the rapid transcriptional responses to cold and the slower accumulation of protective proteins, reflecting the membrane turnover and lipid remodeling required for compositional change.
Membrane_Lipid_Adjustment_Quality is_a Cryoprotection_Quality, representing the modification of membrane fatty acid composition to maintain fluidity at low temperatures. This class addresses the essential membrane adaptation for cell survival during freezing. The desaturation ratio (unsaturated/saturated) provides quantitative metric with 30-60% increases during acclimation. Linolenic acid (18:3) accumulation shows strong correlation with hardiness. V. riparia shows superior lipid adjustment capacity compared to V. vinifera explaining part of hardiness differential. Fatty acid desaturase enzyme activity underlies compositional changes.
Fettsäure-Desaturierung
Lipid-Kälteanpassung
Membranfluiditäts-Anpassung
Membranlipid-Anpassung
fatty acid desaturation
lipid cold adaptation
membrane composition modification
membrane fluidity adjustment
Xin Z, Browse J (2000) Cold comfort farm: the acclimation of plants to freezing temperatures. Plant, Cell and Environment 23:893-902 Thomashow MF (1999) Plant cold acclimation: freezing tolerance genes and regulatory mechanisms. Annual Review of Plant Physiology and Plant Molecular Biology 50:571-599 Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235
Membrane Lipid Adjustment Quality
Systematic assessment of Membrane_Thermostability_Disposition at Stellenbosch University employed electrolyte leakage assays and lipid profiling across 25 cultivars to characterize membrane heat tolerance and its biochemical basis. Leaf discs were exposed to temperatures from 30°C to 55°C for 30 minutes, then incubated in deionized water with conductivity measured at 2 hours (initial leakage) and after autoclaving (total electrolytes). The temperature causing 50% relative electrolyte leakage (T50-EL) defined membrane thermostability threshold. Membrane_Thermostability_Disposition ranged from T50-EL of 46.2°C ('Pinot Noir', least stable) to 52.8°C ('Cinsaut', most stable), representing 6.6°C variation among commercial cultivars. Lipid analysis revealed that membrane fatty acid saturation correlated significantly with T50-EL (r = 0.72), with heat-tolerant 'Cinsaut' showing 42% saturated fatty acids versus 31% in heat-sensitive 'Pinot Noir'. The ratio of linolenic (18:3) to linoleic (18:2) acid was inversely correlated with thermostability (r = -0.68), consistent with the destabilizing effect of highly unsaturated lipids at elevated temperatures. Heat acclimation (pre-treatment at 38°C for 24 hours) improved Membrane_Thermostability_Disposition by 2-4°C across cultivars through increased lipid saturation, demonstrating phenotypic plasticity in this trait. These findings establish membrane lipid composition as a key determinant of Membrane_Thermostability_Disposition and identify lipid desaturase genes as potential breeding targets.
A Membrane_Thermostability_Disposition is a Heat_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to maintain cellular membrane integrity and function at elevated temperatures, preventing the increased membrane fluidity, phase transitions, and ion leakage that characterize heat-induced membrane damage. This disposition represents the thermal stability of biological membranes, which are primary sites of heat injury due to temperature-dependent changes in lipid bilayer properties. Heat stress increases membrane fluidity beyond optimal ranges, disrupting the precise lipid environment required for membrane protein function, increasing passive ion permeability leading to loss of cellular ion gradients, and potentially causing membrane phase transitions and irreversible damage. Membrane_Thermostability_Disposition depends on membrane lipid composition (higher lipid saturation increases thermostability), membrane protein content and type (some proteins stabilize membranes), accumulation of compatible solutes (trehalose, proline) that interact with membrane surfaces, and heat shock proteins that associate with membranes during stress. Membrane thermostability is commonly assessed by measuring electrolyte leakage from heat-treated tissue, with lower leakage indicating greater membrane integrity. Cultivars with strong Membrane_Thermostability_Disposition maintain lower electrolyte leakage, preserve cellular compartmentalization, and protect membrane-localized processes (photosynthetic electron transport, respiratory function, signaling) during heat exposure.
Membrane_Thermostability_Disposition is_a Heat_Tolerance_Disposition. Membrane stability is a primary determinant of cellular heat tolerance, as membranes are highly temperature-sensitive structures. Electrolyte leakage is the standard assay, with T50-EL providing a quantitative stability threshold. Membrane composition (lipid saturation) is the primary biochemical determinant, with more saturated membranes showing greater thermostability. Heat acclimation can increase membrane thermostability through lipid remodeling, representing an inducible component of heat tolerance.
Membran-Hitzestabilität
Membran-Thermostabilität-Disposition
Zellmembran-Integrität
cellular membrane integrity
lipid bilayer thermotolerance
membrane heat stability
Blum A, Ebercon A (1981) Cell membrane stability as a measure of drought and heat tolerance in wheat, Crop Science 21:43-47 Sung DY et al. (2003) Acquired tolerance to temperature extremes, Trends in Plant Science 8:179-187 Falcone DL et al. (2004) Regulation of membrane fatty acid composition by temperature in mutants of Arabidopsis with alterations in membrane lipid composition, BMC Plant Biology 4:17
Membrane Thermostability Disposition
Membrane thermostability quality assessment across grapevine rootstock genotypes evaluated the cellular basis of heat tolerance differences affecting scion performance in warm climate viticulture. Electrolyte leakage assays following controlled heat exposure (45°C for 30 minutes) revealed substantial variation in membrane thermostability quality among rootstocks. Ramsey (Vitis champinii) demonstrated excellent membrane thermostability quality with only 18% electrolyte leakage indicating minimal membrane damage, consistent with its native adaptation to hot Texas conditions. 110R (V. berlandieri × V. rupestris) showed good membrane thermostability quality with 25% leakage. SO4 (V. berlandieri × V. riparia) exhibited moderate membrane thermostability quality with 38% leakage reflecting its less heat-adapted parentage. Riparia Gloire (V. riparia) showed poor membrane thermostability quality with 55% leakage approaching the 60% threshold typically indicating lethal membrane damage. Lipid analysis revealed that high membrane thermostability quality correlated with elevated saturated fatty acid content (35% in Ramsey versus 25% in Riparia Gloire), identifying the biochemical basis of stability differences. The membrane thermostability quality ranking matched field observations of rootstock performance during heat events, with Ramsey-rooted vines showing minimal leaf scorch while Riparia Gloire-rooted vines exhibited substantial damage under equivalent conditions.
Membrane thermostability quality is a thermal tolerance quality that inheres in a grapevine cultivar or clone and represents the capacity of cellular membranes to maintain structural integrity and functional properties under elevated temperature conditions, determined by the lipid composition, protein complement, and organizational characteristics of membrane systems that collectively establish the temperature threshold beyond which membrane dysfunction disrupts cellular homeostasis and leads to cell damage or death. This quality addresses what is often considered the primary site of heat injury in plants, recognizing that membrane integrity is essential for maintaining cellular compartmentalization, ionic gradients, and the function of membrane-associated processes including photosynthesis, respiration, and signal transduction that fail when membranes are disrupted by excessive heat. The ontological significance of membrane thermostability quality lies in its role as a key determinant of overall thermal tolerance, with membrane damage representing an early and often rate-limiting event in heat injury that precedes and enables subsequent cellular dysfunction, making membrane stability a primary target for understanding and potentially improving heat tolerance through breeding or management approaches. The biological basis of membrane thermostability encompasses the biophysical properties of membrane lipids that determine the temperature at which phase transitions and fluidization compromise membrane barrier function, the stability of membrane-associated proteins including those of the photosynthetic apparatus, and the capacity for membrane repair and remodeling that may restore function following sublethal heat exposure. The lipid composition of membranes, particularly the degree of fatty acid saturation, strongly influences thermostability, with more saturated lipids providing greater stability at high temperatures, and with the capacity for heat-induced lipid remodeling contributing to acquired thermotolerance through acclimation. Assessment of membrane thermostability quality typically employs electrolyte leakage assays that quantify the loss of cellular contents through heat-damaged membranes, providing a sensitive and reproducible measure of membrane integrity following controlled heat exposure.
Membrane_Thermostability_Quality is_a Thermal_Tolerance_Quality, representing the heat resistance of cellular membranes as a primary determinant of overall thermotolerance. This class addresses the often rate-limiting site of heat injury. Electrolyte leakage percentage provides standard quantification with >50-60% typically indicating lethal damage. Lipid saturation level correlates with stability and explains much cultivar variation. Rootstock differences substantially influence scion heat tolerance through membrane stability effects. Distinguished from Photosynthetic_Heat_Tolerance_Quality and Enzyme_Thermostability_Quality which address other cellular systems.
Lipid-Thermostabilität
Membran-Hitzetoleranz
Membran-Thermostabilität
Membranstabilität
lipid thermostability
membrane heat tolerance
membrane integrity
membrane stability
Wahid A et al. (2007) Heat tolerance in plants: an overview. Environmental and Experimental Botany 61:199-223 Xu S et al. (2014) Effects of heat acclimation pretreatment on changes of membrane lipid peroxidation, antioxidant metabolites, and ultrastructure of chloroplasts in two cool-season turfgrass species under heat stress. Environmental and Experimental Botany 56:274-285 Salvucci ME, Crafts-Brandner SJ (2004) Inhibition of photosynthesis by heat stress: the activation state of Rubisco as a limiting factor in photosynthesis. Physiologia Plantarum 120:179-186
Membrane Thermostability Quality
Syrah grafted on 110 Richter rootstock showing meristem cap transition zone on primary root apex, transition zone 0.36 mm from apex showing high mitotic activity with 19% mitotic index, microscopic examination revealing asymmetric divisions producing cap-destined daughters moving into columella and corpus-destined daughters entering meristem proper, transition zone cells showing stem cell characteristics maintaining undifferentiated state while producing differentiated derivatives, cell lineage tracing demonstrating transition zone producing both cap and meristem cell populations maintaining balanced apex organization, quiescent center signals maintaining transition zone stem cell identity
Meristem cap transition zone is root cap transition zone emphasizing meristematic character and cell production function at boundary, characterized by actively dividing cells producing daughter cells that differentiate into either root cap cells moving distally or root meristem corpus cells moving basipetally, showing high mitotic activity with cells dividing every 12-20 hours maintaining both cap and meristem cell populations, demonstrating dual derivative production where single transition zone dividing cell can produce one cap-destined and one corpus-destined daughter through asymmetric division or produce two cells of same fate with stochastic assignment, and representing critical meristematic interface maintaining both protective cap covering apex and functional meristem generating root body tissues through balanced cell production, with meristem cap transition zone showing stem cell characteristics maintaining undifferentiated state while producing differentiated derivatives, responding to signals from quiescent center maintaining organization, and demonstrating developmental plasticity adjusting production rates of cap versus corpus cells maintaining appropriate proportions despite variable cell loss rates from cap sloughing or variable meristem demand from growth rate changes. Meristem cap transition zone cell division activity includes= showing high mitotic index (transition zone cells dividing frequently maintaining 15-25% mitotic index comparable to meristem proper), exhibiting continuous mitosis (ongoing divisions throughout active growth period producing steady stream of daughter cells), demonstrating rapid cell cycle (transition zone cells completing cycle every 12-20 hours during peak activity), revealing dual population maintenance (divisions producing sufficient cells for both cap and meristem maintenance), and displaying division plane importance (orientation of division plane potentially determining daughter cell fates). Meristem cap transition zone asymmetric divisions includes= showing differential daughter fates (divisions potentially producing one cap-destined and one corpus-destined daughter from single mother cell), exhibiting unequal cell inheritance (asymmetric divisions potentially distributing factors differently between daughters determining fates), demonstrating stem cell divisions (transition zone stem cells dividing to produce one stem cell and one differentiating cell), revealing molecular asymmetry (asymmetric protein or RNA distribution during division influencing daughter fates), and displaying fate determination (division asymmetry contributing to cell fate choice at transition zone boundary). Meristem cap transition zone production balance includes= showing cap replacement (transition zone producing adequate cap cells replacing continuous sloughing losses from cap surface), exhibiting meristem maintenance (transition zone producing sufficient corpus cells maintaining meristem size and activity), demonstrating balanced output (cap and corpus production rates balancing maintaining appropriate apex proportions), revealing adaptive regulation (production rates potentially adjusting to variable demands maintaining organization), and displaying homeostatic control (feedback mechanisms ensuring transition zone maintains proper balance despite perturbations). Meristem cap transition zone stem cell characteristics includes= showing self-renewal capacity (transition zone stem cells maintaining undifferentiated state through successive divisions), exhibiting multipotency (transition zone cells capable of producing multiple cell types both cap and corpus lineages), demonstrating niche dependence (transition zone stem cells requiring signals from quiescent center and neighboring cells maintaining identity), revealing asymmetric division (stem cells dividing to produce one stem cell and one differentiating cell), and displaying long-term maintenance (transition zone stem cell population potentially persisting throughout root life). Meristem cap transition zone positional signaling includes= showing gradient effects (morphogen gradients emanating from quiescent center potentially affecting transition zone cell behavior), exhibiting cell-cell communication (direct signaling between transition zone cells coordinating development), demonstrating boundary signals (signals marking cap-corpus boundary potentially influencing fate decisions), revealing hormone effects (auxin, cytokinins, and other hormones affecting transition zone activity and fate determination), and displaying mechanical signals (physical forces potentially influencing transition zone cell division orientation and fate). Meristem cap transition zone in apex regeneration includes= showing repair capacity (transition zone capable of compensating for apex damage through adjusted cell production), exhibiting developmental plasticity (transition zone adjusting behavior responding to perturbations maintaining apex organization), demonstrating regenerative potential (transition zone contributing to apex recovery after injury through increased or redirected cell production), revealing quiescent center dependence (transition zone regeneration requiring intact quiescent center providing organizational signals), and displaying recovery patterns (damaged apex showing recovery beginning in transition zone spreading to restored cap and meristem). Meristem cap transition zone molecular regulation includes= showing transcription factor networks (key transcription factors regulating transition zone identity and activity), exhibiting cell cycle control (cyclins and CDKs regulating transition zone division rates and timing), demonstrating fate regulators (specific genes determining whether cells join cap or corpus populations), revealing boundary markers (molecular markers distinguishing transition zone from adjacent cap and meristem proper), and displaying signaling pathways (developmental signaling pathways coordinating transition zone function with apex organization). Meristem cap transition zone in root growth includes= determining apex organization (transition zone activity maintaining proper apex structure essential for growth), enabling continuous growth (transition zone providing cells for both protective cap and elongating meristem), demonstrating growth rate correlation (transition zone activity positively correlated with root elongation rate), revealing meristem size control (transition zone production contributing to meristem size determining growth capacity), and displaying developmental coordination (transition zone integrating cap and meristem development enabling coordinate apex function).
Meristem_Cap_Transition_Zone emphasizes meristematic activity and dual cell production and is distinguished from static boundary emphasis by focus on active division and fate determination. Critical for understanding stem cell function, asymmetric division, balanced production, positional signaling, apex regeneration, and the actively dividing boundary region producing both cap and meristem cells maintaining root apex organization.
Cap-corpus meristem interface
Dividing transition zone
Meristem-Haube-Übergangszone
Meristematic boundary region
Meristematische Schnittstelle
Teilende Grenzregion
Meristem Cap Transition Zone
A Cabernet Sauvignon clone 15 vine at FPS bears a Meristem_Derived_Mother_Plant_Role, regenerated from a 0.3 mm meristem tip excised from a heat-treated (90 days, 38°C) mother plant, with the combination treatment successfully eliminating GLRaV-3 that had persisted through thermotherapy alone—confirmed negative by three consecutive annual ELISA and RT-PCR tests. A Riesling clone 49 serves in a Meristem_Derived_Mother_Plant_Role following ENTAV protocols combining 60-day thermotherapy with 0.5 mm shoot tip culture on Murashige-Skoog medium, achieving virus-free status after failed elimination attempts using heat treatment alone. A Sangiovese selection from the Italian VCR program achieved Meristem_Derived_Mother_Plant_Role status through somatic embryogenesis from heat-treated material, representing an alternative regeneration pathway for varieties recalcitrant to direct organogenesis.
A mother plant role that inheres in a grapevine regenerated from meristem tip culture or shoot tip culture, exploiting the principle that apical meristematic tissues often remain virus-free or at lower viral titer than mature tissues due to the absence of developed vascular connections through which systemic pathogens spread. Meristem-derived mother plants originate from excision of tiny shoot tips (0.2-0.5 mm for true meristem culture, 0.5-2.0 mm for shoot tip culture) that are cultured in vitro through callus or direct organogenesis to regenerate complete plantlets, then tested to confirm pathogen elimination success. Meristem culture is typically combined with thermotherapy (shoot tips excised from heat-treated plants) to maximize elimination rates, as pre-treatment heat stress suppresses pathogen replication and reduces viral load in meristematic regions. The technique provides the most reliable virus elimination for heat-tolerant pathogens (leafroll viruses, some vitiviruses) where thermotherapy alone is insufficient, and enables elimination of multiple pathogens simultaneously through a single regeneration cycle. Meristem-derived plants require verification of genetic stability (true-to-type status) as tissue culture may occasionally induce somaclonal variation.
Meristem_Derived_Mother_Plant_Role captures the tissue culture origin of mother plants, indicating that pathogen elimination involved in vitro regeneration rather than conventional vegetative propagation. The technique exploits the developmental principle that meristematic cells divide faster than viruses can replicate and spread, creating a pathogen-free zone at the shoot apex if viral titer is sufficiently reduced through pre-treatment. Success rates vary by pathogen and protocol—reported ranges of 60-95% elimination for leafroll viruses with optimized combined treatments. Concerns about somaclonal variation (genetic or epigenetic changes during tissue culture) require verification of true-to-type status through ampelographic evaluation and increasingly molecular marker analysis before meristem-derived plants enter nuclear stock collections. The specialized facilities, expertise, and time required (12-24 months from excision to confirmed-clean plant) concentrate meristem culture at research institutions and specialized clean stock programs rather than commercial nurseries.
Gewebekultur-Rebe
in vitro derived vine
in-vitro-vermehrte Mutterpflanze
meristem tip culture plant
meristem-derived vine
meristemstämmige Mutterpflanze
tissue culture derived plant
Barlass M, Skene KGM (1978) In vitro propagation of grapevine from shoot apices, Vitis 17: 335-340 Panattoni A et al. (2013) Grapevine virus elimination, Journal of Plant Pathology 95: 229-235 Goussard PG, Wiid J (1992) Virus elimination from grapevine by tissue culture, South African Journal of Enology and Viticulture 13: 76-83 Maliogka VI et al. (2015) Grapevine virus diseases, APS Press
Meristem Derived Mother Plant Role
A meristematic region leaf at node 15 on a vigorously extending shoot of Vitis vinifera cv. Chardonnay in Champagne, France, positioned 2.2 cm behind the shoot apical meristem dome among the three youngest visible primordia, measuring 2.1 cm length in early unfolding stage representing approximately 12% of eventual mature size, exhibiting very pale yellowish coloration with pronounced bronze-red pigmentation from high anthocyanin (4.8 mg g⁻¹ fresh weight), composed predominantly of actively dividing cells with mitotic index of 18% indicating rapid cell proliferation establishing the foundational cell population, showing minimal tissue differentiation with procambial strands barely distinguishable as faint darker zones, positioned within the highly protected microenvironment immediately surrounding the apical meristem where relative humidity exceeds 90% and temperature is moderated 3-4°C below ambient, contributing zero photosynthesis representing pure sink consuming approximately 7 mg carbohydrate per day for meristematic activity, occurring during vigorous shoot extension at 9-11 cm per week with new primordia initiating every 2.3 days, measured at 4 days post-emergence during peak vegetative growth at shoot BBCH stage 75 (berries pea-sized) in late June, representing the meristematic region leaf zone where newest leaves develop in immediate proximity to the active apical meristem.
A growing tip leaf positioned in the immediate vicinity of the shoot apical meristem within the meristematic region in Vitis vinifera, characterized by position within 2-4 cm of the apical meristem dome representing the newest emerged leaves, extremely small size typically 1-4 cm length in early unfolding or primordium stages, spatial association with active meristematic tissues including the apical dome and newest primordia, very recent emergence within previous 2-7 days, composition dominated by actively dividing and rapidly expanding cells with high nuclear-to-cytoplasmic ratios and dense cytoplasm, minimal tissue differentiation beyond procambial initiation, very pale coloration with minimal chlorophyll development and high anthocyanin concentrations providing photoprotection for vulnerable meristematic cells, extreme fragility and sensitivity to environmental stresses, zero or minimal photosynthetic capacity representing pure resource sinks, and functional integration with meristematic processes supporting continued shoot extension through provision of protective covering for the apical meristem and contributions to the shoot tip microenvironment. Meristematic region leaves represent the immediate developmental frontier where leaf primordia transition from meristematic initiation to recognizable leaf structures, experiencing the most protected microenvironment within the shoot tip where the apical meristem and newest primordia are buffered from environmental extremes, consuming resources in meristematic cell division and initial expansion that establish the cellular foundation for eventual mature leaf structure.
Meristematic region leaves represent the youngest recognizable leaves in immediate proximity to active meristematic tissues, distinguished from terminal leaves generally by specific association with the meristem zone and from shoot tip leaves by extreme youth and minimal development. These leaves occupy the innermost protected zone of the shoot tip, experiencing maximal environmental buffering and intimate association with meristematic processes. The meristematic region extends approximately 2-4 cm from the apical dome depending on plastochron and internode elongation rates, encompassing the meristem itself and the 2-4 youngest primordia or emerging leaves. Developmental characteristics show maximum similarity to primordial stages, with active cell division establishing cell populations and beginning expansion processes. Protection within the meristematic region is critical for successful development, with damage to this zone through frost, desiccation, pest attack, or mechanical injury potentially affecting both the existing youngest leaves and the apical meristem's capacity for continued primordium initiation. Shoot topping removing the meristematic region eliminates not only the youngest leaves but the apical meristem itself, preventing further shoot extension and leaf initiation requiring lateral shoot development to restore growth capacity.
Apikaldome Bereich Blatt
Meristem-Region Blatt
Primordial-Zone Weinblatt
apical dome region leaf
meristem-adjacent leaf
primordial zone leaf
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 43-85 Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 33-78.
Meristematic Region Leaf
Cabernet Sauvignon mesocarp at maturity comprises 88-92% of pericarp volume (86-90% of berry fresh weight excluding seeds), consisting of outer mesocarp transition zone (5-7 cell layers, 55-85 μm, cells 35-75 μm diameter, 2.5-5 mg/g tannins), middle mesocarp bulk (cells averaging 135-165 μm diameter in central region with thin 0.5-0.7 μm walls, containing 24-26% sugars, 0.6-0.75% acids, 0.4-0.8 mg/g tannins), and inner mesocarp zone (150-220 μm, cells 55-85 μm, 4.5-6.5 mg/g tannins), providing soft sweet flesh contributing majority of fermentable sugars and juice volume but minimal wine color or tannin. Chardonnay white wine cultivar mesocarp showing white wine composition= 89-93% of pericarp volume, middle mesocarp cells 140-175 μm diameter, 21-24% sugars (glucose 10.5-12%, fructose 10.5-12%), 0.7-0.95% organic acids (tartaric 0.42-0.58%, malic 0.28-0.38%), maintaining higher acidity than typical red cultivar mesocarp important for white wine freshness, with mesocarp juice providing essentially complete must composition for white wine production after pressing separating juice from skins and seeds. Syrah mesocarp cell expansion during ripening demonstrating growth dynamics= veraison (BBCH 81, week 10) middle mesocarp cells 72-88 μm diameter mid-ripening (week 14) cells 115-145 μm maturity (week 18) cells 145-175 μm, representing 101-133% diameter increase (310-570% volume increase per cell) driving berry enlargement during Phase II, with mesocarp cell expansion responsible for 85-92% of total berry volume increase during ripening phase. Merlot mesocarp radial organization showing tissue gradients= outer mesocarp (50-120 μm depth beneath hypodermis) cells 42-78 μm diameter, walls 0.9-1.4 μm, tannins 3.2-5.5 mg/g middle mesocarp (120-1800 μm depth, central bulk) cells 125-165 μm, walls 0.5-0.7 μm, tannins 0.5-0.9 mg/g inner mesocarp (150-280 μm from locular cavities) cells 58-88 μm, walls 0.9-1.3 μm, tannins 4.2-6.8 mg/g, demonstrating systematic radial gradients in cell size, wall thickness, and phenolic content with maximum cell size and minimum tannins in central mesocarp, smaller cells and elevated tannins in outer and inner transition zones. Pinot Noir mesocarp contribution to wine composition= mesocarp providing 88-91% of berry fresh weight, yielding 72-76 L juice per 100 kg berries during pressing, with mesocarp juice composition (23-24% sugars, 6.5-7.5 g/L titratable acidity, pH 3.4-3.6, <0.05 mg/g anthocyanins, 0.6-1.2 mg/g total phenolics) determining base wine composition before skin and seed phenolic extraction during maceration, demonstrating mesocarp as primary determinant of wine sugar/alcohol level and base acidity while skins/seeds provide color and tannin structure.
A mesocarp is the middle and bulk tissue of the fruit pericarp developing from the ovary wall mesophyll, comprising the majority of berry tissue (typically 88-95% of pericarp volume) between outer pericarp (exocarp/skin) and inner pericarp (endocarp/seed-adjacent layers), consisting primarily of large thin-walled parenchyma cells that accumulate sugars, water, organic acids, and minerals creating the fleshy edible portion of the berry, providing juice volume and fermentable sugars for wine production while containing minimal phenolic compounds compared to outer and inner pericarp regions. Mesocarp anatomical organization comprises= (1) outer mesocarp layers (2-8 cell layers beneath hypodermis, 25-150 μm total thickness) showing transition from small hypodermal cells to larger mesocarp cells, with cells 30-90 μm diameter, walls 0.8-1.5 μm thick, sometimes elevated tannin content (2-6 mg/g) distinguishing this zone from middle mesocarp (2) middle mesocarp (central bulk tissue, typically 75-85% of pericarp volume) with large parenchyma cells 80-180 μm diameter, thin walls 0.4-0.8 μm, minimal tannins (0.3-1 mg/g), maximum sugar and water accumulation creating soft fleshy texture (3) inner mesocarp layers (innermost mesocarp adjacent to locular cavities, sometimes classified as inner pericarp, 100-300 μm thickness) with cells intermediate in size (40-90 μm), moderately thick walls (0.8-1.5 μm), elevated tannins (3-7 mg/g) relative to middle mesocarp. Mesocarp cellular structure features= (1) large parenchyma cells particularly in middle mesocarp representing some of largest cells in berry tissue (central cells 120-180 μm diameter at maturity versus 15-35 μm in epidermis) (2) thin flexible cell walls composed of cellulose, hemicellulose, and pectin allowing cell expansion during berry growth and creating soft texture when turgid (3) large central vacuoles occupying 90-95% of mature cell volume serving as storage compartments for sugars, acids, water, and minerals (4) thin peripheral cytoplasm layer (1-3 μm) containing nucleus and organelles focused on transport and storage rather than biosynthesis (5) intercellular spaces (air spaces between cells) comprising 1-5% of mesocarp volume providing gas diffusion pathways. Mesocarp composition at maturity= (1) sugars 18-28% (glucose and fructose in approximately 1=1 ratio) accumulated in vacuoles creating osmotic potential and sweet taste (2) organic acids 0.5-0.9% fresh weight (tartaric acid 0.3-0.6%, malic acid 0.1-0.4%) providing acidity and pH buffering (3) water 75-82% providing juice volume and turgid texture (4) minerals including potassium (1200-2200 mg/kg), calcium (60-150 mg/kg), magnesium (60-120 mg/kg) (5) minimal phenolic compounds with tannins 0.3-1 mg/g in middle mesocarp, 2-6 mg/g in outer/inner mesocarp zones, essentially no anthocyanins (<0.05 mg/g). Mesocarp development follows berry growth phases= Phase I (BBCH 71-75, weeks 0-6) involving rapid cell division creating mesocarp cell population (1.2-2.8 million cells representing 75-85% of total pericarp cells) with cells 30-70 μm diameter lag phase (BBCH 77, weeks 6-8) with minimal mesocarp growth but metabolic reprogramming Phase II (BBCH 81-89, weeks 9-18+) involving dramatic cell expansion particularly in middle mesocarp where cells expand from 60-90 μm to 110-180 μm diameter (120-200% diameter increase, 350-700% volume increase) creating bulk of berry enlargement and juice accumulation. Mesocarp functions= (1) primary storage tissue accumulating photosynthates (sugars) and water supporting berry growth and ripening (2) juice source providing 85-92% of berry fresh weight and essentially all grape juice volume for wine or juice production (3) sugar source providing fermentable sugars (glucose, fructose) for alcoholic fermentation in wine production (4) acid reservoir maintaining organic acids affecting wine acidity, pH, and stability.
Mesocarp represents the bulk flesh tissue of the berry and is essentially synonymous with Middle_Pericarp in broader usage, though strict anatomical definitions distinguish outer mesocarp (beneath hypodermis) and inner mesocarp (above inner pericarp/endocarp) from middle mesocarp (central bulk). Sub-classification into Outer and Inner mesocarp reflects radial tissue organization with transitional zones at boundaries. Understanding mesocarp structure and composition is fundamental for viticulture and enology as this tissue provides juice volume, fermentable sugars, and base acidity determining fundamental wine composition, while contributing minimally to wine color and tannin structure.
Beerenfleisch
Fruchtfleisch
Mesokarp
flesh
pulp
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 180-190 Coombe, B.G., McCarthy, M.G. (2000). Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6=131-135 Nunan, K.J. et al. (2001). Changes in cell wall composition during ripening. Plant Physiology 126=520-528.
Mesocarp
Mesocarp turgor tracking in Shiraz berries through ripening using pressure microprobe reveals decline from 0.38 MPa at 10 days pre-véraison to 0.05 MPa at véraison onset (within 72 hours of first color appearance), remaining below 0.08 MPa through harvest at 25°Brix, with the persistent low turgor state explaining why post-véraison berries do not recover firmness even under favorable vine water status, as the hydraulic disconnection prevents turgor re-establishment despite ample water availability.
Mesocarp turgor quality is a cell turgor quality that inheres in the parenchyma cells of grape berry flesh and represents the hydrostatic pressure within the vacuolated storage cells that constitute the majority of berry volume and whose turgor determines fruit firmness, texture perception, and the hydraulic environment for sugar accumulation and ripening metabolism. Within the BFO framework, this quality specifically characterizes the turgor of the tissue compartment most directly determining berry mechanical properties and quality parameters, distinguishing mesocarp cell behavior from the distinct turgor dynamics of skin and seed tissues. The ontological significance lies in mesocarp turgor governing the textural transition from hard pre-véraison berries to soft ripe fruit, where the dramatic turgor decline at véraison (from ~0.4 MPa to near zero) precedes and enables the sugar accumulation phase by switching phloem unloading from symplastic to apoplastic pathway. The large central vacuoles of mesocarp cells (occupying >90% of cell volume at maturity) store the osmotically active sugars, organic acids, and secondary metabolites that determine fruit composition, with turgor affecting both the water influx driving cell expansion and the membrane transport processes loading these solutes into vacuolar storage. Mesocarp turgor shows strong developmental dynamics, building during the cell expansion phase of Stage I berry growth, declining sharply at véraison onset marking Stage II initiation, and remaining low throughout the ripening phase despite massive sugar accumulation that increases osmotic potential. The uncoupling of mesocarp turgor from vine water status post-véraison reflects the hydraulic isolation of ripening berries, where xylem backflow and reduced phloem connectivity limit water exchange with the parent vine. Understanding mesocarp turgor quality enables interpretation of berry softening mechanisms, prediction of texture evolution during ripening, and correlation of water relations with compositional development.
Mesocarp_Turgor_Quality addresses the dominant tissue compartment determining berry texture and the hydraulic environment for ripening. The dramatic turgor decline at véraison represents a developmental switch with profound implications for berry physiology and quality development. This quality distinguishes from whole-berry firmness measurements that integrate contributions from skin, seeds, and mesocarp with different mechanical properties and turgor states.
Beerenfleisch-Zellturgor
Fruchtfleischturgor
Mesokarp-Turgor
Pulpaturgor
berry parenchyma turgor
flesh cell turgor
fruit mesocarp pressure
pulp turgor
Thomas TR et al. (2006) Mesocarp cell turgor in Vitis vinifera L. berries throughout development and its relation to firmness, growth, and the onset of ripening. Planta 224(5): 1078-1088 Matthews MA et al. (2009) Berry ripening and water relations in Vitis. Acta Horticulturae 844: 165-172
Mesocarp Turgor Quality
Mesophyll conductance limitation assessment in heat-stressed Merlot reveals the temperature effect on internal diffusion, with leaves at optimal temperature (25°C) showing gm of 0.22 mol/m²/s and Lm of 18%, while leaves at elevated temperature (38°C) display reduced gm of 0.11 mol/m²/s and increased Lm of 32%, with the enhanced mesophyll limitation arising from membrane fluidity changes and reduced aquaporin activity that together restrict CO₂ diffusion to chloroplasts, demonstrating that heat stress affects not only biochemistry but also the physical pathway for CO₂ supply.
Mesophyll conductance limitation quality is a photosynthetic limitation quality that represents the constraint on photosynthesis arising from resistance to CO₂ diffusion through the mesophyll from substomatal cavity to Rubisco active sites, characterizing the degree to which internal diffusional resistance limits CO₂ concentration at the site of carboxylation below the substomatal level. Within the BFO framework, this quality specifically addresses the mesophyll component of diffusional limitation, distinguishing mesophyll conductance (gm) limitation from stomatal and biochemical limitations while recognizing that mesophyll resistance has emerged as a significant and previously underappreciated constraint on photosynthesis that substantially reduces chloroplast CO₂ concentration below substomatal levels. The ontological significance lies in mesophyll conductance limitation representing a structural and physiological constraint that is less immediately responsive than stomatal limitation but more tractable than biochemical limitation, where the resistance to CO₂ diffusion through cell walls, membranes, and cytoplasm determines the effective CO₂ concentration at Rubisco and thus the realized carboxylation rate. The physiological basis for mesophyll conductance limitation involves the diffusion pathway from substomatal cavity through cell wall aqueous phase, across plasma membrane and chloroplast envelope, through stroma to Rubisco, with resistance components including cell wall thickness and porosity (significant contributor), membrane permeability (affected by aquaporin expression), and chloroplast positioning (distance to intercellular air space). The quantitative assessment of mesophyll conductance employs combined gas exchange and chlorophyll fluorescence measurements (variable J method), carbon isotope discrimination analysis, or oxygen isotope techniques, with gm values for Vitis vinifera typically ranging from 0.1-0.3 mol/m²/s and corresponding Lm values of 15-35% depending on leaf structure and environmental conditions. The stress responses of mesophyll conductance show reduction under drought (cell shrinkage, reduced aquaporin activity), heat (membrane effects), and with leaf age (structural changes), often contributing substantially to total photosynthetic limitation under stress conditions. Understanding mesophyll conductance limitation quality enables comprehensive limitation analysis, appreciation of structural constraints on photosynthesis, and identification of potential targets for genetic improvement through leaf anatomy modification.
Mesophyll_Conductance_Limitation_Quality addresses the internal diffusional limitation from substomatal cavity to Rubisco. This quality has emerged as a significant constraint that was historically overlooked. The class supports comprehensive limitation analysis and identification of structural targets for photosynthetic improvement.
Lm-Wert
Mesophyllleitfähigkeitslimitation
Mesophyllwiderstand
gm limitation
internal conductance limitation
interne Diffusionslimitation
mesophyll resistance
Flexas J et al. (2008) Mesophyll conductance to CO₂: current knowledge and future prospects. Plant, Cell & Environment 31(5): 602-621 Tomás M et al. (2013) Importance of leaf anatomy in determining mesophyll diffusion conductance to CO₂ across species. Plant Journal 73(4): 602-614
Mesophyll Conductance Limitation Quality
Mesophyll hydraulic conductance assessment in grapevine leaves under varying conditions reveals the aquaporin regulation of this pathway, with well-hydrated morning leaves showing Kmesophyll of 5.8 mmol/m²/s/MPa (active aquaporins facilitating transcellular flow), midday stressed leaves declining to 2.2 mmol/m²/s/MPa (aquaporin downregulation reducing membrane permeability), and overnight recovery to 5.1 mmol/m²/s/MPa (aquaporin reactivation), demonstrating the dynamic regulation of mesophyll conductance that contributes to diurnal Kleaf patterns and may coordinate with stomatal responses to water stress.
Mesophyll hydraulic conductance quality is a leaf hydraulic conductance quality that represents the capacity for water movement through the non-vascular mesophyll tissue from vein endings to the evaporating surfaces in substomatal cavities, characterizing the final segment of the liquid water pathway that involves cell-to-cell water movement regulated by aquaporins and affected by mesophyll anatomy. Within the BFO framework, this quality specifically addresses the extra-xylary component of leaf water transport, distinguishing mesophyll pathways from vein conductivity while recognizing that the mesophyll often represents a significant fraction of total leaf hydraulic resistance, particularly in thick leaves or under conditions that reduce aquaporin activity. The ontological significance lies in mesophyll hydraulic conductance determining the water potential at evaporating surfaces relative to vein water potential, where the resistance to water movement through mesophyll creates the final water potential drop in the liquid pathway that, combined with stomatal transpiration, determines the water potential gradient driving flow from soil to atmosphere. The pathway components of mesophyll conductance include the bundle sheath interface (the transition zone between vein xylem and mesophyll), the apoplastic route (water movement through cell walls), and the cell-to-cell route (transcellular movement through membranes requiring aquaporins), with the relative importance of these pathways depending on leaf anatomy and aquaporin expression. The regulation of mesophyll conductance through aquaporin activity provides a mechanism for rapid adjustment of leaf water supply, with evidence that mesophyll aquaporin expression and gating respond to water status, light, and other signals, creating dynamic regulation of the outside-xylem pathway that affects leaf hydraulic function on timescales too rapid for anatomical change. The relationship between mesophyll conductance for water (Kmesophyll) and mesophyll conductance for CO₂ (gm) is of particular interest, as both involve water and CO₂ movement through similar pathways, with emerging evidence suggesting coordination between hydraulic and photosynthetic mesophyll properties. Understanding mesophyll hydraulic conductance quality enables assessment of the extra-xylary limitation on leaf water supply, interpretation of aquaporin effects on leaf hydraulics, and evaluation of the coordination between hydraulic and photosynthetic mesophyll functions.
Mesophyll_Hydraulic_Conductance_Quality addresses the extra-xylary pathway from vein endings to evaporating surfaces. This quality is regulated by aquaporins and may coordinate with photosynthetic mesophyll conductance. The class supports leaf hydraulics interpretation and aquaporin function assessment through mesophyll conductance characterization.
Kmesophyll
Kox
Mesophyllleitfähigkeit
außer-xylary Leitfähigkeit
extra-xylary conductance
outside-xylem conductance
Buckley TN (2015) The contributions of apoplastic, symplastic and gas phase pathways for water transport outside the bundle sheath in leaves. Plant, Cell & Environment 38(1): 7-22 Scoffoni C et al. (2014) Outside-xylem vulnerability, not xylem embolism, controls leaf hydraulic decline during dehydration. Plant Physiology 165(3): 1541-1552
Mesophyll Hydraulic Conductance Quality
In Pinot Noir vines in Burgundy, metabolic adjustment during autumn cold acclimation includes starch accumulation in cane tissues reaching 8-12% of dry weight by December, proline concentrations in bud tissues increasing from 2 to 15 μmol per gram fresh weight, raffinose content rising from undetectable to 3-4% of soluble sugars, and respiratory rate declining by approximately 70% as tissues transition from active growth to dormant winter metabolism.
Metabolic adjustment in Vitis vinifera represents the comprehensive reprogramming of cellular metabolism that occurs during temperature acclimation, encompassing shifts in carbohydrate partitioning, respiratory pathway regulation, nitrogen metabolism, and secondary metabolism that collectively support the development and maintenance of cold hardiness. This metabolic remodeling reflects the fundamentally different requirements of dormant tissues compared to actively growing tissues, with emphasis shifting from supporting cell division and expansion toward accumulating protective compounds, maintaining minimal vital functions at low temperatures, and preserving reserves for spring growth resumption. Carbohydrate metabolism undergoes profound changes during metabolic adjustment, with starch accumulating in woody tissues as a winter reserve while simultaneously a portion of carbohydrate flux is redirected toward synthesis of raffinose family oligosaccharides and other cryoprotective sugars, and cold-induced invertases release hexoses from sucrose that contribute to osmotic adjustment and serve as substrates for respiration. Respiratory metabolism shifts toward increased capacity for alternative respiratory pathways that function at low temperatures and generate heat that may provide limited protection during freezing events, while the overall respiratory rate progressively declines as temperatures drop and tissues enter dormancy. Nitrogen metabolism during cold acclimation involves accumulation of free amino acids particularly proline which serves both as a compatible solute and a readily mobilizable nitrogen reserve, along with synthesis of cold-induced proteins including dehydrins, antifreeze proteins, and modified membrane proteins. Secondary metabolism contributes to cold acclimation through synthesis of phenolic compounds that may provide antioxidant protection during freeze-induced oxidative stress, and modifications to cell wall composition that affect ice propagation and cell mechanical properties during freezing. The regulation of metabolic adjustment involves complex transcriptional networks coordinated with the CBF pathway, hormonal signals particularly from abscisic acid, and metabolic feedback regulation, producing the integrated metabolic phenotype characteristic of cold-acclimated dormant tissues.
Metabolic_Adjustment represents the metabolic dimension of Temperature_Acclimation, occurring in parallel with Cold_Acclimation_Start and Freeze_Tolerance_Development. Distinguished from purely structural or gene expression changes by its focus on metabolic pathway regulation and metabolite accumulation. Encompasses carbohydrate, nitrogen, respiratory, and secondary metabolism changes. Duration spans the entire acclimation period of 8-12 weeks. Key outcomes include starch reserve accumulation, cryoprotectant synthesis, and respiratory decline.
Dormanz-Metabolismus-Änderung
Kälte-Stoffwechselanpassung
Stoffwechsel-Umstellung
acclimation metabolic adjustment
cold metabolic reprogramming
dormancy metabolism shift
metabolische Anpassung
winter metabolic transition
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 5 Carbohydrate Supply and Reserve Storage Xin Z, Browse J (2000) Cold comfort farm: the acclimation of plants to freezing temperatures. Plant Cell and Environment 23: 893-902 Guy CL (1990) Cold acclimation and freezing stress tolerance: role of protein metabolism. Annual Review of Plant Physiology 41: 187-223
Metabolic Adjustment
In Riesling vines in the Mosel Valley, metabolic downregulation progresses through autumn with whole-shoot respiration rates declining from 2.5 μmol CO2 per gram dry weight per hour in September to 0.6 μmol per gram per hour by late November, protein synthesis rates decreasing by approximately 75% over the same period, and overall metabolic activity reaching minimum winter levels by December when mean temperatures fall below 5°C.
Metabolic downregulation in Vitis vinifera represents the coordinated reduction in overall metabolic activity that accompanies growth cessation during autumn, encompassing decreased rates of photosynthesis, respiration, protein synthesis, and biosynthetic pathway activity as the vine transitions from active growth to winter dormancy. This process occurs progressively through autumn as temperatures decline and day length shortens, with metabolic rates decreasing in parallel with the cessation of growth processes that normally consume the majority of metabolic output. The reduction in photosynthetic metabolism during metabolic downregulation reflects both the declining chlorophyll content in senescing leaves and the reduced requirement for photosynthate as growth cessation eliminates major carbon sinks, with net photosynthetic rate in remaining functional leaves declining by 50-70% from summer maximum values by late autumn. Respiratory metabolism decreases substantially during metabolic downregulation, with whole-plant respiration rates declining by 60-80% as temperature-dependent reaction rates slow and the metabolically active tissue mass decreases through leaf senescence, resulting in the characteristic low maintenance respiration of dormant vines. Protein synthesis rates decline dramatically during the transition to dormancy, with overall protein turnover decreasing as growth-related proteins are degraded and not replaced, while cold-induced proteins including dehydrins become relatively more prominent in the reduced protein complement of dormant tissues. Biosynthetic pathway activities generally decline during metabolic downregulation, though specific pathways involved in cold acclimation and cryoprotectant synthesis remain active or increase during the early phases before decreasing as dormancy deepens and temperatures continue falling. The enzymatic basis of metabolic downregulation involves both temperature-dependent reductions in reaction rates as temperatures approach and fall below optimal ranges, and regulatory changes in enzyme activity and gene expression that actively suppress metabolic capacity beyond what temperature alone would cause. The ecological significance of metabolic downregulation relates to energy conservation during the period when photosynthetic input is minimal and the vine must survive on stored reserves, with reduced metabolic rates minimizing reserve consumption and extending the period over which reserves can support vital functions. The extent of metabolic downregulation varies among tissue types, with woody stem tissues achieving very low metabolic rates characteristic of deep dormancy while root tissues in unfrozen soil may maintain somewhat higher activity levels throughout winter.
Metabolic_Downregulation represents the metabolic dimension of Growth_Cessation, specifically addressing the reduction in overall metabolic rates. Distinguished from Metabolic_Adjustment under Temperature_Acclimation which addresses qualitative metabolic changes for cold hardiness, while Metabolic_Downregulation addresses quantitative rate reductions. Encompasses photosynthesis, respiration, and biosynthetic pathway declines. Duration spans 6-10 weeks through autumn. Critical for energy conservation during the dormant period when photosynthetic input is absent.
Dormanz-Stoffwechsel
Stoffwechsel-Reduktion
dormancy metabolism
metabolic decline
metabolic reduction
metabolic slowdown
metabolische Herunterregulierung
metabolischer Rückgang
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 5 Carbohydrate Supply and Reserve Storage Zapata C et al (2004) Partitioning and mobilization of starch and N reserves in grapevine. Journal of Experimental Botany 55: 2165-2175 Wample RL, Bary A (1992) Harvest date as a factor in carbohydrate storage and cold hardiness of Cabernet Sauvignon grapevines. Journal of the American Society for Horticultural Science 117: 32-36
Metabolic Downregulation
In a Burgundy Chardonnay vineyard, metabolic pathway transition was characterized over 7 days in late August Year N+1 (BBCH 83) when PEPC activity declined from 16.5 to 4.8 units/g FW, NADP-malic enzyme increased from 4.2 to 28.5 units/g FW, pyruvate dehydrogenase rose from 11 to 19 units/g FW, isocitrate dehydrogenase maintained 32 units/g FW, berry organic acids plateaued at 24 g/L, respiratory quotient increased from 1.02 to 1.18 indicating enhanced organic acid respiration, under cool French conditions (daily high 22-25°C supporting gradual transition).
Metabolic_Pathway_Transition represents the specific biochemical reorientation of central carbon metabolism pathways during veraison when citric acid cycle shifts from primarily biosynthetic anaplerotic function toward predominantly oxidative catabolic mode, phosphoenolpyruvate metabolism redirects from CO₂ fixation to decarboxylation, and respiratory pathways reorganize to efficiently catabolize organic acids creating the enzymatic foundation enabling acid decline while supporting ripening energy demands. This transition occurs during veraison onset (BBCH 83), typically 1-6 days concurrent with transcriptional reprogramming and preceding active metabolic changes, and is characterized by citric acid cycle enzyme activities reorienting with oxidative enzymes (isocitrate dehydrogenase, α-ketoglutarate dehydrogenase) maintaining or increasing 15-35% while biosynthetic citrate synthase in anaplerotic mode declining 25-50%, phosphoenolpyruvate carboxylase activity decreasing 60-80% eliminating primary CO₂ fixation pathway, malic enzyme activity increasing 5-12 fold establishing decarboxylation pathway dominance, and pyruvate metabolism shifting from predominantly biosynthetic uses toward oxidative citric acid cycle entry through pyruvate dehydrogenase complex with activity potentially increasing 30-60% supporting acid-derived pyruvate oxidation. Citric acid cycle reorganization involves functional transition from TCA cycle operating in biosynthetic mode during green berry growth (producing intermediates for amino acid synthesis, providing carbon skeletons for secondary metabolism, generating reducing equivalents for biosynthesis) toward purely oxidative catabolism during ripening (completely oxidizing acetyl-CoA and malate to CO₂, maximizing ATP production, minimal intermediate withdrawal), enzyme activity patterns showing isocitrate dehydrogenase (IDH) maintaining or increasing from 18-35 units/g to 22-45 units/g ensuring oxidative flux through rate-limiting step, α-ketoglutarate dehydrogenase similarly sustained or elevated 15-32 units/g supporting continued oxidation, malate dehydrogenase showing complex regulation with activity 25-55 units/g potentially shifting from biosynthetic (cytoplasmic, producing malate for storage) to oxidative (mitochondrial, consuming malate from catabolism), and aconitase maintaining activity 20-40 units/g ensuring citrate-isocitrate interconversion supporting cycle flux. Phosphoenolpyruvate metabolism redirection demonstrates PEPC (phosphoenolpyruvate carboxylase) declining from green berry peak 12-28 units/g to post-transition 3-10 units/g representing major reduction in carboxylation reaction PEP + CO₂ → oxaloacetate that during green berry growth fixes CO₂ and produces malate precursor, pyruvate kinase potentially adjusting though activity measurements variable across studies with some showing increases supporting glycolytic flux while others showing stability, phosphoenolpyruvate carboxykinase (PEPCK) potentially involved in some grape varieties catalyzing reverse reaction oxaloacetate → PEP + CO₂ though role controversial with evidence suggesting minor contribution compared to malic enzyme, and net effect of PEPC decline eliminating major anabolic pathway redirecting PEP toward pyruvate kinase and subsequently pyruvate metabolism rather than oxaloacetate and malate synthesis. Malic enzyme pathway establishment includes NADP-malic enzyme increasing from 2-6 units/g to 15-45 units/g representing major induction of oxidative decarboxylation pathway malate → pyruvate + CO₂, NAD-malic enzyme showing parallel increase from 1-4 units/g to 6-18 units/g contributing to total decarboxylation capacity, subcellular distribution with cytosolic isoforms dominant 65-80% providing primary site for malate oxidation, mitochondrial isoforms 20-35% potentially supporting anaplerotic or alternative functions, and kinetic properties with Km values 0.5-2.5 mM for malate and pH optimum 7.0-7.5 ensuring efficient catalysis in cytoplasmic environment enabling maximum flux during acid decline. Pyruvate metabolism reorganization shows pyruvate dehydrogenase complex (PDC) potentially increasing activity 25-55% from 8-18 units/g to 12-28 units/g reflecting enhanced requirement for acetyl-CoA production from malic enzyme-derived pyruvate, mitochondrial pyruvate carrier facilitating enhanced pyruvate import from cytoplasm enabling oxidation, pyruvate entering citric acid cycle through acetyl-CoA representing major fate 70-85% of cytoplasmic pyruvate production during active acid decline, and minor alternative fates including transamination to alanine 8-15%, fermentation to lactate or ethanol under hypoxic conditions 5-12% particularly berry interior, and potential gluconeogenic uses <5% generally minimal during ripening. Respiratory chain adaptation involves Complex I (NADH dehydrogenase) maintaining or slightly increasing capacity ensuring efficient NADH oxidation from increased citric acid cycle flux, Complex II (succinate dehydrogenase) sustained activity supporting continued electron entry from FADH₂, Complexes III-IV (cytochrome bc₁ and cytochrome c oxidase) adequate capacity with measurements 12-28 units/g enabling efficient electron transport to oxygen, ATP synthase (Complex V) maintaining or increasing capacity 15-35 units/g ensuring ATP production keeps pace with enhanced respiratory substrate flux, and alternative oxidase potentially present providing overflow pathway under conditions of excess reducing equivalents though contribution generally minor in grape compared to some other fruits. Energy metabolism implications include ATP production transitioning from predominantly hexose-based (32 ATP per glucose complete oxidation) toward mixed substrate with organic acids contributing substantially (15 ATP per malate), total berry ATP synthesis rates relatively stable 50-90 μmol g⁻¹ h⁻¹ before and after transition maintaining energy supply despite altered fuel sources, ATP utilization patterns adjusting with declining allocation to growth processes (cell division, wall synthesis) and increasing allocation to storage and quality metabolism (sugar transport, anthocyanin synthesis, aromatic compound production), and metabolic efficiency potentially improving as stored acids provide respiratory substrate without requiring import of hexoses exclusively for respiration enabling dual strategy of hexose storage and acid catabolism. Redox metabolism changes show NADH production from citric acid cycle potentially increasing 30-60% as oxidative mode dominates compared to biosynthetic mode where intermediates withdrawn, NADPH generation from NADP-malic enzyme providing substantial reductive capacity 2.5-5.0 μmol g⁻¹ h⁻¹ supporting biosynthesis particularly anthocyanin and flavonoid synthesis in reds plus general antioxidant systems, NAD⁺/NADH and NADP⁺/NADPH ratios adjusting with ripening metabolism potentially showing NAD⁺/NADH declining slightly 8:1 to 5:1 reflecting enhanced oxidation while NADP⁺/NADPH may increase 2:1 to 4:1 as NADPH consumption for biosynthesis matches or exceeds malic enzyme production, and redox homeostasis maintained through multiple dehydrogenases, transhydrogenases, and antioxidant systems preventing excessive oxidation or reduction disrupting cellular function. Metabolite pool adjustments during transition include citric acid cycle intermediates showing complex dynamics with some (citrate, α-ketoglutarate) potentially declining 20-40% as biosynthetic withdrawal reduces and oxidative flux dominates, others (malate, succinate) showing path-dependent changes with malate declining through catabolism while succinate sometimes accumulating slightly, amino acid pools adjusting as citric acid cycle intermediate availability changes affecting biosynthesis particularly glutamate family from α-ketoglutarate and aspartate family from oxaloacetate potentially declining 15-35%, and organic phosphates (ATP, ADP, AMP) maintaining relatively stable ratios despite metabolic reorganization indicating energy charge homeostasis preserved throughout transition. Regulatory mechanisms coordinating transition include allosteric regulation with multiple enzymes responding to metabolite effectors (ATP, ADP, AMP, citrate, malate) enabling metabolic responsiveness, transcriptional control through ripening-specific factors coordinately activating catabolic genes while suppressing biosynthetic genes creating coordinated reprogramming, post-translational modifications particularly phosphorylation potentially affecting enzyme activities though modifications during grape transition incompletely characterized, and hormonal signals particularly ABA triggering regulatory cascades affecting multiple pathway components simultaneously enabling rapid coordinated transition. Environmental modulation of transition includes temperature affecting enzyme activities and reaction rates with warmer conditions 24-30°C potentially accelerating transition completion 2-4 days compared to cooler 18-24°C, oxygen availability affecting respiratory pathway flux with adequate O₂ essential for oxidative metabolism while hypoxia promoting fermentative alternatives, substrate availability with adequate photosynthesis providing resources for enzyme synthesis and metabolic function supporting normal transition, and stress conditions potentially disrupting transition timing or completeness through resource limitation or signaling perturbations. Cultivar variation in pathway transition demonstrates early-ripening varieties showing rapid transition 4-6 days with prompt enzyme activity changes and metabolite adjustments, mid-season cultivars displaying 6-9 day transition with progressive reorganization, and late varieties exhibiting extended transition 8-14 days with gradual shifts reflecting genetic differences in developmental control and metabolic flexibility. Physiological significance includes pathway transition enabling efficient resource utilization by catabolizing stored acids for energy while simultaneously storing imported sugars avoiding wasteful simultaneous synthesis and degradation, metabolic commitment representing developmental point-of-no-return with ripening metabolism established irreversibly, and energy economy optimizing by extracting ATP from accumulated acids supporting ripening processes without fully relying on imported hexose oxidation. Quality implications include transition completeness affecting acid decline capacity with inadequate transition potentially limiting acid reduction maintaining excessive acidity, timing relative to other processes determining compositional balance at harvest, and pathway efficiency affecting berry energy status potentially influencing secondary metabolism and quality compound synthesis. Research applications include enzyme activity profiling characterizing transition kinetics, metabolite flux analysis using isotope tracers quantifying pathway reorganization, transcriptomics revealing regulatory control, and metabolic modeling integrating data predicting transition dynamics enabling mechanistic understanding of ripening metabolism.
Metabolic_Pathway_Transition is distinguished from Acid_Metabolism_Shift by representing specific pathway reorganization rather than overall metabolic reprogramming, from Respiratory_Substrate_Change by focusing on enzymatic pathways rather than substrate utilization, and from gene expression changes by representing functional activity rather than transcriptional control. PEPC declines 60-80%, malic enzyme increases 5-12 fold. Citric acid cycle shifts biosynthetic → oxidative. Duration 5-10 days BBCH 83. Metabolic_Pathway_Transition represents biochemical foundation enabling acid catabolism during ripening.
Pfad-Reorganisation
TCA cycle functional shift
TCA-Zyklus-funktionelle Verschiebung
Zentral-Metabolismus-Neuorientierung
biosynthetic-to-catabolic conversion
biosynthetisch-zu-katabolis che-Umwandlung
central metabolism reorientation
pathway reorganization
Sweetman C, Deluc LG, Cramer GR, Ford CM, Soole KL (2009) Regulation of malate metabolism in grape berry and other developing fruits Phytochem 70:1329-1344 pathway transition comprehensive Ruffner HP (1982) Metabolism of tartaric and malic acids in Vitis:a review Vitis 21:247-259 metabolic pathways Hawker JS (1969) Changes in the activities of malic enzyme, malate dehydrogenase, phosphopyruvate carboxylase and pyruvate dehydrogenase during the development of a non-climacteric fruit:the grape Phytochem 8:9-23 enzyme activity transitions.
Metabolic Pathway Transition
In Sauvignon Blanc vines in Marlborough, New Zealand, metabolic reactivation potential is established by early August following chilling completion, demonstrated by rapid respiratory upregulation from 0.4 to 2.5 μmol CO2 per gram dry weight per hour within 48 hours of transfer to 20°C forcing conditions, with concurrent upregulation of amino acid and nucleotide synthesis pathway enzyme activities indicating prepared metabolic systems capable of supporting the biosynthetic demands of budbreak.
Metabolic reactivation potential in Vitis vinifera represents the restored capacity for rapid upregulation of metabolic activity following dormancy release, describing the state in which biosynthetic pathways, energy metabolism, and supporting metabolic systems have been prepared for the rapid activation required during budbreak and early shoot growth. This potential is established through the chilling-dependent changes that dismantle dormancy while simultaneously preparing metabolic systems for reactivation, creating a poised state where metabolism can rapidly increase upon receipt of growth-promoting temperature signals. The biosynthetic dimension of reactivation potential involves the preparation of pathways for amino acid, nucleotide, lipid, and carbohydrate synthesis that will support the intensive biosynthetic demands of cell division and expansion during budbreak, with enzyme systems maintained in states capable of rapid activation even while operating at low levels during ecodormant quiescence. The energy metabolism dimension encompasses the preparation of respiratory pathways for the increased ATP production required during growth, with mitochondrial systems maintained in states capable of rapid upregulation and the availability of carbohydrate reserves ensuring substrate supply for elevated respiration. The nitrogen metabolism dimension addresses the preparation for mobilization of stored nitrogen reserves and increased amino acid synthesis required for the protein production supporting growth, with nitrogen remobilization pathways prepared for activation when growth signals are received. The regulatory dimension involves the restoration of metabolic regulatory systems that coordinate pathway activities in response to growth demands, including the hormone signaling connections between growth signals and metabolic activation, and the transcriptional programs that upregulate metabolic gene expression during growth initiation. The temperature-dependence of metabolic reactivation creates the link between warming and growth initiation, with increasing temperatures progressively activating metabolic systems according to their temperature response characteristics, eventually reaching activity levels sufficient to support sustained growth when heat accumulation satisfies the requirement for budbreak. The verification of reactivation potential involves demonstrating rapid metabolic upregulation upon warming, with potential confirmed by the speed of respiratory and biosynthetic responses to growth-promoting temperatures in forcing experiments.
Metabolic_Reactivation_Potential represents the metabolic preparedness aspect of Bud_Competence_Restoration. Distinguished from Growth_Capacity_Restoration which addresses cellular growth machinery, while Metabolic_Reactivation_Potential specifically addresses metabolic system readiness. Encompasses biosynthetic, energy, nitrogen, and regulatory dimensions. Temperature-dependent activation links warming to growth initiation. Verified through rapid metabolic upregulation upon warming. Supports the intensive demands of budbreak and early growth.
Metabolismus-Hochregulierungs-Kapazität
Stoffwechsel-Aktivierungsbereitschaft
biosynthetic reactivation readiness
biosynthetische Reaktivierungsbereitschaft
metabolic activation readiness
metabolic reactivation capacity
metabolisches Reaktivierungspotential
metabolism upregulation potential
Metabolic Reactivation Potential
Pinot Noir in Burgundy shows metabolic reprogramming during early September Year N+1 at BBCH 81, with respiratory rate increasing from 1.2 to 1.6 μmol O₂ g⁻¹ h⁻¹, carbon flux redirecting toward anthocyanin biosynthesis with earliest berries accumulating 0.02-0.04 mg/g anthocyanins showing faint pink color, malate declining accelerating from 0.2 to 0.4% per week, abscisic acid spiking from 140 to 620 ng/g over 4-5 days, ethylene increasing from <0.1 to 0.3 nL/g/h, under continental conditions of 22-27°C temperature, typical of this early-to-mid season cultivar crossing metabolic threshold
Metabolic reprogramming represents the decisive metabolic transformation phase during pre-veraison lag when berry metabolism shifts definitively from lag quiescence to ripening-active state, respiratory patterns change toward ripening characteristics, carbon flux redirects comprehensively toward ripening processes, secondary metabolism intensifies dramatically, abscisic acid accumulation accelerates toward spike levels, and metabolic state crosses transformation threshold establishing ripening metabolism, occurring during final days pre-veraison enabling veraison progression. This metabolic transformation phase occurs during late summer to early autumn in late August-early September in temperate regions Year N+1, proceeding over final 3-7 days before veraison, corresponding to BBCH 81 late pre-veraison metabolic shift. Reprogramming mechanism involves respiratory metabolism increasing toward ripening levels, carbon metabolism redirecting from maintenance to ripening biosynthesis, secondary metabolism activating for phenolic and anthocyanin production, hormone dynamics shifting with abscisic acid spike and possible ethylene increase, gene regulatory networks switching to ripening program, and comprehensive metabolic reorganization establishing ripening state. Respiratory metabolism changes shows respiratory rate beginning increase from lag quiescence 1.0-1.5 μmol O₂ g⁻¹ h⁻¹ toward pre-veraison elevated 1.2-1.8 μmol O₂ g⁻¹ h⁻¹, respiratory increase supporting enhanced metabolic activity, ATP production increasing for biosynthetic demands, and respiratory shift as metabolic reprogramming indicator. Respiratory increase mechanisms shows possible increase in mitochondrial enzyme activities, enhanced oxidative phosphorylation, metabolic rate increase supporting ripening preparations, and respiration as energy source for transformation. Respiratory substrate utilization shows continued organic acid oxidation particularly malate respiration, possible beginning sugar oxidation as sugars increase, substrate utilization supporting metabolic demands, and respiratory patterns shifting toward ripening. Carbon metabolism redirection shows carbon flux dramatically redirecting from minimal lag metabolism toward intensive ripening biosynthesis, phenylpropanoid pathway flux increasing for anthocyanin and phenolic synthesis, sugar metabolism flux increasing preparing for accelerated accumulation, organic acid metabolism shifting toward decline patterns, and comprehensive flux reorganization. Phenylpropanoid pathway flux shows carbon flow through shikimate pathway to phenylalanine increasing, phenylalanine ammonia lyase flux increasing producing cinnamic acid derivatives, flux through flavonoid pathway toward anthocyanins intensifying in red cultivars, and pathway flux preparing for color development. Anthocyanin biosynthesis flux shows carbon and reducing equivalent allocation to anthocyanin pathway increasing dramatically, earliest anthocyanin accumulation beginning in some red cultivar berries, pathway flux enabling color development at veraison, and biosynthesis as dramatic reprogramming feature. Sugar metabolism flux shows increased sugar import through activated transporters, sugar phosphorylation and storage metabolism enhanced, vacuole hexose accumulation machinery activated, and flux preparing for ripening sugar accumulation. Organic acid metabolism shifts shows malate catabolism through malic enzyme and oxidation increasing, malate decline accelerating from 0.2-0.4% per week lag to possible 0.3-0.6% per week pre-veraison, tartrate metabolism remaining minimal, and acid metabolism reprogramming toward ripening decline. Secondary metabolism intensification shows phenolic biosynthesis dramatically increasing, flavonoid production intensifying, stilbene synthesis possibly increasing in some cultivars, tannin modification possibly beginning, and secondary metabolism as major reprogramming aspect. Phenolic biosynthesis shows general phenylpropanoid pathway activity increasing, flavonol synthesis possibly intensifying, flavanol production continuing or increasing, and comprehensive phenolic metabolism activation. Tannin modifications shows possible beginning tannin polymerization changes, oxidation possibly initiating, modifications preparing for ripening tannin evolution, and tannin metabolism as ripening preparation. Hormone dynamics shifts shows abscisic acid accumulation accelerating dramatically from lag levels 50-200 ng/g toward spike 500-2000 ng/g, ABA synthesis rate increasing substantially in berry tissues, possible ethylene beginning increase in some cultivars from <0.1 to 0.2-0.5 nL/g/h, auxin remaining low, gibberellin low or declining, and hormone reprogramming as critical regulatory shift. Abscisic acid spike shows ABA increasing 3-10 fold during pre-veraison lag and early veraison, hormone accumulation in berry pericarp and possibly seeds, ABA as central veraison trigger or correlate, and spike as defining hormonal event. Abscisic acid synthesis acceleration shows ABA biosynthesis genes including 9-cis-epoxycarotenoid dioxygenase NCED upregulating substantially, biosynthetic enzyme activities increasing, ABA production rate accelerating, and synthesis as hormone spike mechanism. Ethylene possible role shows ethylene possibly beginning increase in some cultivars, ethylene possibly involved in softening regulation, hormone potentially contributing to ripening regulation though role varies by cultivar, and ethylene as secondary hormone signal. Gene regulatory network switching shows master transcriptional regulators shifting control, ripening transcription factors activating target genes, regulatory network reorganizing comprehensively, and transcriptional reprogramming implementing metabolic shifts. Transcription factor activation shows ripening-associated MYB transcription factors activating anthocyanin pathway genes, sugar-responsive transcription factors activating transporter genes, developmental transcription factors directing ripening program, and transcriptional control as reprogramming mechanism. Epigenetic modifications possible shows possible chromatin remodeling enabling ripening gene expression, histone modifications affecting transcriptional accessibility, DNA methylation changes possibly regulating developmental genes, and epigenetic regulation as potential reprogramming mechanism. Protein expression changes shows dramatic protein profile shifts during reprogramming, ripening-associated proteins rapidly accumulating, lag-phase proteins declining or stable, and protein changes implementing metabolic transformation. Metabolite profile changes shows metabolite composition shifting rapidly, phenolic intermediates accumulating, sugar phosphates increasing, organic acids declining, and metabolite dynamics reflecting flux reorganization. Berry composition during reprogramming shows water 78-82% stable, organic acids declining 0.7-0.9% with accelerating decline, sugars 13-16% with possible beginning acceleration, pH rising 3.1-3.3 approaching veraison range, earliest anthocyanin accumulation in some red berries, and composition rapidly shifting. Earliest color manifestation shows trace anthocyanin accumulation 0.01-0.05 mg/g in earliest red cultivar berries during late reprogramming, visible as faint color tinge, anthocyanin as earliest transformation marker, and color as metabolic reprogramming outcome. Berry morphology during reprogramming shows berry still predominantly green in most berries, earliest berries showing faint color in red cultivars, texture beginning softening in earliest berries, and morphology showing earliest transformation signs. Environmental effects on reprogramming include temperature affecting reprogramming rate with warm conditions accelerating metabolic shifts, cool conditions slowing reprogramming, water stress possibly enhancing abscisic acid accumulation accelerating reprogramming, and environmental factors influencing transformation timing. Temperature effects shows warm conditions promoting faster enzyme activity and metabolic flux changes, cool conditions extending reprogramming period, and temperature as rate determinant. Water stress effects shows moderate deficit enhancing ABA production possibly accelerating reprogramming and veraison, stress as potential transformation advance mechanism, and water influencing hormonal regulation. The process represents second and final aspect of pre-veraison lag establishing ripening metabolism, following veraison preparation. Duration final 3-7 days representing metabolic transformation. Regional timing shows metabolic reprogramming during late August-early September immediately before veraison onset. Viticultural significance includes representing decisive metabolic shift to ripening state, demonstrating abscisic acid dynamics as veraison trigger or correlate, establishing ripening metabolism enabling transformation, and determining veraison onset through metabolic threshold crossing. Research applications include analyzing metabolic flux changes and reorganization, investigating abscisic acid and hormonal regulation of transformation, evaluating gene regulatory networks and transcriptional reprogramming, quantifying metabolic thresholds for veraison onset, and assessing metabolic reprogramming as determinant of veraison timing and ripening progression.
Metabolic_Reprogramming is second and final aspect of Pre_Veraison_Lag, distinguished by decisive metabolic shift. Represents respiratory rate 1.0-1.5 → 1.2-1.8 μmol O₂ g⁻¹ h⁻¹. Carbon flux redirection to phenylpropanoid pathway. Secondary metabolism intensification. ABA spike 50-200 → 500-2000 ng/g. Possible ethylene increase. Earliest anthocyanin 0.01-0.05 mg/g in red cultivars. Malate decline accelerating. Duration 3-7 days. Follows Veraison_Preparation. Completes Pre_Veraison_Lag. Crosses metabolic threshold enabling veraison.
Reifungs-Metabolismus-Etablierung
decisive metabolic shift
metabolic threshold crossing
metabolic transformation
metabolische Reprogrammierung
metabolische Schwellen-Überschreitung
metabolische Transformation
ripening metabolism establishment
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 465-475 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Metabolic Reprogramming
Syrah in Rhône Valley shows metabolic transition onset during late August-early September Year N+1 at BBCH 81, with ripening genes including anthocyanin biosynthesis PAL, CHS, DFR upregulated 3-6 fold, sugar transporter VvHT1 increased 3.2 fold, respiratory rate stabilizing at 1.2 μmol O₂ g⁻¹ h⁻¹, stress response genes maintained high expression, malate declining from 0.9% to 0.8%, under warm Mediterranean conditions of 24-28°C temperature, typical of this mid-to-late season cultivar initiating lag metabolism
Metabolic transition onset represents the metabolic reorganization initiation phase during lag phase initiation when berry metabolism begins shift from growth-phase to quiescent pre-ripening program, gene expression changes establish lag-associated and early ripening patterns, enzyme activities transition toward maintenance and preparation metabolism, respiratory rate stabilizes at lag low levels, and metabolic reprogramming prepares berry for eventual veraison transformation, occurring over days to 1 week at lag initiation. This metabolic shift onset phase occurs during late summer to early autumn in late August-early September in temperate regions Year N+1, proceeding over days to 1 week representing transition initiation, corresponding to BBCH 81 beginning of ripening stage early period. Transition onset mechanism involves transcriptional changes initiating lag and pre-ripening gene expression patterns, enzyme activity shifts implementing metabolic reorganization, respiratory metabolism declining to lag maintenance levels, hormone level changes possibly regulating transition, and integrated metabolic reprogramming establishing lag metabolism preparing for ripening. Gene expression changes at onset shows growth genes at basal non-functional levels maintained from enlargement completion, lag-associated genes including stress response and secondary metabolism genes sustained or increasing expression, ripening genes beginning very low-level expression 2-10 fold above basal representing early activation, and expression patterns characteristic of lag initiation. Lag gene expression characteristics shows stress response genes including heat shock proteins HSP70, HSP90, antioxidant enzymes superoxide dismutase, catalase maintained or upregulated, secondary metabolism genes for phenolic and flavonoid biosynthesis expressed, and lag-specific expression supporting quiescent metabolism. Stress response gene expression shows enhanced expression of protective genes, accumulation of stress-protective proteins, metabolic adaptation to potential environmental stress during lag, and stress preparedness as lag metabolic feature. Secondary metabolism gene expression shows phenylpropanoid pathway genes expressed, flavonoid biosynthesis genes active, stilbene synthase for resveratrol possibly expressed, and secondary metabolism continuing during quiescence. Ripening gene early expression shows anthocyanin biosynthesis pathway genes in red cultivars beginning 2-8 fold low-level expression including phenylalanine ammonia lyase PAL, chalcone synthase CHS, dihydroflavonol reductase DFR at initiation levels, sugar transporter genes hexose transporters beginning increase 2-5 fold, cell wall modification genes polygalacturonase, pectin methylesterase starting low expression, and early ripening gene activation preparing for veraison. Anthocyanin biosynthesis preparation shows pathway genes beginning transcription at low levels during lag initiation, enzyme proteins accumulating slowly, pathway priming for veraison activation, and preparation as metabolic transition feature. Sugar transporter preparation shows hexose transporter genes VvHT1, VvHT2 increasing expression 2-5 fold, transporter proteins beginning accumulation, membrane insertion preparing for import capacity, and transporter preparation enabling veraison sugar accumulation acceleration. Cell wall modification preparation shows softening enzyme genes beginning low-level expression, enzyme precursors accumulating, cell wall modification machinery preparing for ripening texture changes, and preparation as transition characteristic. Enzyme activity transitions shows growth-related enzymes at basal minimal activity, maintenance metabolism enzymes including basic respiratory, protein turnover, membrane maintenance sustained at moderate levels, early ripening enzymes beginning accumulation at very low activity, and activity patterns shifting toward lag and pre-ripening profiles. Respiratory enzyme activities shows glycolysis and TCA cycle enzymes at reduced activity supporting maintenance metabolism, oxidative phosphorylation continuing at low rate, respiratory rate 1.0-1.5 μmol O₂ g⁻¹ h⁻¹ characteristic of lag metabolism, and low respiration as quiescence indicator. Carbon metabolism transitions shows minimal biosynthetic flux, carbon for maintenance and slow compositional changes, limited sugar import and vacuole storage, and metabolism characteristic of quiescent pre-ripening state. Organic acid metabolism shows continued malate respiration at slow rate, malic enzyme and malate dehydrogenase activities declining, malate declining slowly 0.1-0.3% over lag phase, tartrate metabolism minimal maintaining stable levels, and acid dynamics continuing from enlargement. Sugar metabolism shows limited sugar import through phloem at low rate, sugar accumulation slow 0.5-1.5% per week, vacuole hexose transporter activity low, and metabolism preparing for ripening acceleration. Protein expression changes shows growth-related proteins at minimal levels, lag-associated stress and secondary metabolism proteins maintained or increasing, early ripening proteins beginning accumulation at low levels, and protein profiles reflecting metabolic transition. Hormone dynamics during transition onset shows auxin at low levels characteristic of non-growing state, gibberellin possibly declining or low, abscisic acid remaining low 50-200 ng/g before eventual veraison spike to 500-2000 ng/g, ethylene low in most cultivars, and hormone levels reflecting quiescent transition state. Berry composition during transition onset shows water 78-82% stable, organic acids 0.8-1.2% declining slowly, sugars 11-15% increasing slowly, pH 3.0-3.2 rising gradually, and composition transitioning toward lag characteristics. Metabolite changes shows slow compositional shifts from continued metabolism, organic acids declining from respiration, sugars increasing from limited import, secondary metabolites including phenolics accumulating slowly, and metabolite dynamics characteristic of lag metabolism. Environmental effects on transition onset include temperature affecting transition kinetics with warm conditions accelerating metabolic shifts, cool conditions slowing transition rate, water availability affecting berry metabolism and stress response, and environmental factors influencing transition dynamics. Temperature effects show warm conditions promoting faster gene expression changes and enzyme activity transitions, cool conditions extending transition period, and temperature affecting metabolic reprogramming rate. Transition onset timing shows occurrence during days to 1 week at lag initiation, overlap with growth rate decline establishing coordinated lag entry, and timing marking metabolic shift to quiescence. Transition onset assessment shows gene expression profiling documenting transcriptional changes, enzyme activity assays quantifying metabolic shifts, metabolite analysis showing compositional dynamics, and molecular markers characterizing transition. The process represents second aspect of lag phase initiation establishing metabolic reorganization, following growth rate decline start. Duration typically days to 1 week representing transition initiation period. Regional timing shows metabolic transition onset during late August-early September across climates at lag initiation. Viticultural significance includes representing metabolic shift to lag quiescence and pre-ripening preparation, establishing gene expression and enzyme patterns characteristic of lag phase, preparing ripening machinery for veraison activation, and determining metabolic foundation for subsequent ripening transformation. Research applications include analyzing transcriptional reprogramming at transition onset, investigating metabolic reorganization mechanisms and regulation, evaluating environmental effects on transition timing and dynamics, quantifying gene expression and enzyme activity changes, and assessing metabolic transition as determinant of lag characteristics and veraison readiness.
Metabolic_Transition_Onset is second aspect of Lag_Phase_Initiation, distinguished by metabolic reorganization. Represents gene expression shift: growth genes basal, lag genes sustained, ripening genes 2-10 fold early activation. Anthocyanin pathway, sugar transporters, softening enzymes beginning expression. Respiratory rate 1.0-1.5 μmol O₂ g⁻¹ h⁻¹. Stress response enhanced. Secondary metabolism active. Duration days to 1 week. Follows Growth_Rate_Decline_Start. Completes Lag_Phase_Initiation. Prepares ripening machinery for veraison.
Prä-Reifungs-Metabolismus-Initiierung
Ruhephase-Metabolismus-Beginn
metabolic reorganization start
metabolic shift onset
metabolische Transitions-Beginn
metabolische Umstellung-Start
pre-ripening metabolism initiation
quiescence metabolism onset
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 455-460 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Metabolic Transition Onset
Methoxypyrazine concentration varies over three orders of magnitude across V. vinifera cultivars, from <1 ng/L in non-herbaceous types to 30-80 ng/L in high-MP cultivars such as Cabernet Sauvignon, Sauvignon Blanc, and Carménère. IBMP (3-isobutyl-2-methoxypyrazine) predominates in most cultivars (70-95% of total MPs), with IPMP and SBMP as minor contributors. MP biosynthesis occurs in both berries and leaves, with accumulation peaking pre-veraison (reaching 50-150 ng/L) then declining 80-95% during ripening through photodegradation and dilution to harvest levels of 5-40 ng/L. The sensory threshold of IBMP (1-2 ng/L in red wine, 8-15 ng/L in white wine) means even trace concentrations significantly impact wine character. Cluster light exposure reduces MP concentration by 50-80% compared to shaded fruit.
An aromatic compound disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate alkyl-methoxypyrazine compounds (IBMP, IPMP, SBMP) in grape berries and vegetative tissues, with genotype-specific concentrations determining the intensity of green, herbaceous, bell pepper, and vegetal aroma character that defines certain Bordeaux varieties and contributes to wine style.
Methoxypyrazine_Accumulation_Disposition serves as the parent class for specific MP compound traits. The green/herbaceous character from MPs is valued at moderate levels in some wine styles (Sauvignon Blanc, cool-climate Cabernet) but considered a fault when excessive. Unlike most grape aroma compounds, MPs are stable through fermentation and aging, making viticultural management the primary tool for controlling wine MP levels. The O-methyltransferase (VvOMT3) catalyzing the final biosynthetic step shows cultivar-specific expression explaining genetic variation in MP accumulation capacity.
MP synthesis capacity
Methoxypyrazin-Akkumulation
Pyrazin-Disposition
green aroma trait
grüne Aromen
herbaceous character disposition
Dunlevy JD et al. (2010) The production of flavour and aroma compounds in grape berries. In: Roubelakis-Angelakis KA (ed) Grapevine Molecular Physiology and Biotechnology, 2nd Ed, Springer pp.: 293-340 Roujou de Boubée D et al. (2000) Identification of an odoriferous compound responsible for the green bell pepper aroma in wines made from immature Cabernet Sauvignon berries. Journal of Agricultural and Food Chemistry 48:4830-4834
Methoxypyrazine Accumulation Disposition
Methoxypyrazine content quality management in Marlborough Sauvignon Blanc targets the 15-25 ng/L IBMP range that provides the characteristic capsicum-herbaceous notes defining regional style without the aggressive vegetal character of excessive levels viticultural protocols employing fruit zone leaf removal at pea-size (exposing clusters to 40-60% ambient light) reduce IBMP from pre-removal levels of 40-60 ng/L to target range by harvest, with the light-driven degradation providing growers precise control over the herbaceous character that distinguishes Marlborough's green-spectrum style from the riper, tropical Sauvignon Blanc styles of warmer regions.
Methoxypyrazine_Content_Quality encompasses the chemical qualities characterizing the concentration and composition of alkyl-methoxypyrazine compounds in grape berries, capturing the nitrogen-containing heterocyclic volatiles responsible for the distinctive green bell pepper, herbaceous, and vegetal aromas in Sauvignon Blanc, Cabernet Sauvignon, and related cultivars. Methoxypyrazines are biosynthesized from amino acids (leucine, valine, isoleucine) through pathways involving methylation, producing three primary grape-relevant compounds: 3-isobutyl-2-methoxypyrazine (IBMP), 3-isopropyl-2-methoxypyrazine (IPMP), and 3-sec-butyl-2-methoxypyrazine (SBMP). These compounds occur at extraordinarily low concentrations (parts per trillion) yet exert powerful sensory impact due to their remarkably low odor thresholds, with IBMP detectable at approximately 1-2 ng/L in water and 10-15 ng/L in wine matrices. Methoxypyrazine concentration follows a distinctive developmental pattern, accumulating early in berry development and reaching maximum levels before véraison, then declining during ripening through metabolic degradation with light exposure as the primary driver of catabolism. The rate of methoxypyrazine decline depends strongly on fruit zone light environment, with shaded clusters retaining substantially higher concentrations than sun-exposed clusters at equivalent maturity. This light sensitivity provides viticultural management tools through canopy manipulation to modulate final methoxypyrazine content. Cultivar genetics determines baseline methoxypyrazine accumulation capacity, with Sauvignon Blanc, Cabernet Sauvignon, Cabernet Franc, Carmenère, and Sauvignon Gris showing characteristic high accumulation, while most other cultivars produce negligible methoxypyrazine levels. The sensory contribution of methoxypyrazines is highly concentration-dependent, with moderate levels (15-30 ng/L IBMP) contributing desirable complexity and typicity in Sauvignon Blanc, while excessive levels (above 40 ng/L) produce aggressively vegetal, unripe character considered a quality defect in most wine styles.
Methoxypyrazine_Content_Quality serves as the parent class for specific methoxypyrazine compounds including IBMP_Content_Quality and IPMP_Content_Quality. The extremely low concentrations require specialized analytical methods (GC-MS/MS, SPME-GC-MS) with isotope dilution quantification for accurate measurement. Methoxypyrazines are chemically stable in wine and do not change significantly during aging, making grape concentration the primary determinant of wine levels. The sensory threshold varies among individuals and wine matrices, requiring context-specific interpretation of concentration data.
Alkyl-Methoxypyrazin-Konzentration
MP content
Methoxypyrazingehalt
Pyrazinaromaqualität
alkyl-methoxypyrazine concentration
green pepper compound content
pyrazine aroma quality
Dunlevy JD et al. (2009) A Methyltransferase Essential for the Methoxypyrazine-Derived Flavour of Wine. The Plant Journal 59(3): 404-412 Ryona I et al. (2008) Effects of Cluster Light Exposure on 3-Isobutyl-2-methoxypyrazine Accumulation and Degradation Patterns in Red Wine Grapes (Vitis vinifera L. Cv. Cabernet Franc). Journal of Agricultural and Food Chemistry 56(22): 10838-10846
Methoxypyrazine Content Quality
Methoxypyrazine data for Cabernet Sauvignon from Coonawarra (2023 vintage) records IBMP at 6.2 ng/L (below the 10-15 ng/L sensory threshold in red wine), IPMP at 0.8 ng/L (below the 2 ng/L threshold), and SBMP at 0.3 ng/L, indicating successful photodegradation of methoxypyrazines through the open canopy management and extended ripening season characteristic of the region — contrasting with shaded-canopy Cabernet Sauvignon from a cooler site recording IBMP at 28 ng/L (OAV 2.3), producing dominant green capsicum character. Methoxypyrazine data for Sauvignon Blanc from Marlborough (2024 vintage) documents IBMP at 32 ng/L, well above the 2 ng/L threshold in white wine but within the 20-40 ng/L range considered characteristic of the region's signature herbaceous-tropical aromatic style, with the elevated methoxypyrazine concentration reflecting the cool maritime climate that limits photodegradation. Methoxypyrazine data for Grüner Veltliner from Kamptal (2024 vintage) records IBMP at 3.8 ng/L, below the white wine threshold, confirming the minimal green character expected in ripe Austrian Grüner Veltliner.
Methoxypyrazine data is an aromatic analysis data entity that records the concentration of alkyl-methoxypyrazine compounds in grape berries and wine, determined by headspace SPME-GC-MS or multidimensional GC-MS at ultra-trace sensitivity (sub-ng/L), expressed in nanograms per liter (ng/L), encompassing 3-isobutyl-2-methoxypyrazine (IBMP), 3-isopropyl-2-methoxypyrazine (IPMP), and 3-sec-butyl-2-methoxypyrazine (SBMP), and providing the chemical quantification of the herbaceous, green bell pepper, and vegetal aromatic character that defines the sensory profile of Bordeaux-type cultivars and whose management through viticultural and oenological practices represents a critical quality control parameter in red wine production.
Methoxypyrazine data occupies a unique position in aromatic analysis because these compounds are among the most potent odorants known — IBMP has a sensory threshold of 2 ng/L in white wine and 10-15 ng/L in red wine, meaning that analytical quantification at the ng/L (parts per trillion) level is required for meaningful quality assessment, demanding specialized instrumentation and extraction protocols beyond routine laboratory capability. The biosynthesis of IBMP occurs through the methylation of isobutylpyrazine by the VvOMT3 O-methyltransferase, with accumulation peaking around véraison and declining through the ripening period via photochemical degradation driven by UV-B radiation and visible light exposure of the berry surface. This photodegradation pathway creates the direct mechanistic link between canopy management (leaf removal, shoot positioning) and methoxypyrazine concentration — exposed berries lose IBMP 2-5× faster than shaded berries, making fruit zone light environment the primary viticultural lever for methoxypyrazine management. The cultivar specificity of methoxypyrazine accumulation is genetically determined — Cabernet Sauvignon, Cabernet Franc, Sauvignon Blanc, and Carménère naturally accumulate high IBMP levels, while Merlot, Pinot Noir, and Chardonnay produce minimal methoxypyrazines regardless of growing conditions. Temperature during ripening modulates IBMP persistence — cooler climates retain higher concentrations due to slower photodegradation and delayed ripening, while warm climates achieve more complete degradation through both temperature-accelerated light reactions and extended hang time. The sensory perception of methoxypyrazines is threshold-dependent and context-specific — sub-threshold IBMP may contribute positively to aromatic complexity and typicity (particularly in Sauvignon Blanc and cool-climate Cabernet Franc), while supra-threshold levels in red wine are generally considered a quality defect associated with under-ripe fruit or excessive canopy shading.
Grünaromadaten
IBMP data
IBMP-Daten
Methoxypyrazindaten
Pyrazinanalyse
green aroma compound data
methoxypyrazine analysis
pyrazine data
vegetative Aromastoffe
vegetative aroma data
Dunlevy JD et al. (2009) A methyltransferase essential for the methoxypyrazine-derived flavour of wine. The Plant Journal 59(3): 404-412 Ryona I et al. (2008) Effect of cluster light exposure on 3-isobutyl-2-methoxypyrazine accumulation and degradation patterns in red wine grapes. Journal of Agricultural and Food Chemistry 56(22): 10838-10846 Sidhu D et al. (2015) Role of methoxypyrazines in the aroma of Bordeaux style wines. Journal of Agricultural and Food Chemistry 63(30): 6738-6748
Methoxypyrazine Data
Microclimate characterization of a VSP-trained Pinot Noir vineyard in Carneros, Napa Valley revealed fruit zone temperatures 3-5°C above ambient during afternoon sun exposure on the western canopy face, canopy interior relative humidity of 85-95% in early morning versus 50-60% ambient, and PAR levels ranging from 1800 µmol/m²/s at the canopy exterior to less than 50 µmol/m²/s at the interior clusters — a 36-fold gradient across 40 cm of canopy depth. Microclimate profiling of traditional pergola-trained Verdicchio in the Marche, Italy documented sustained cluster zone temperatures below 30°C even when ambient air reached 38°C, due to the overhead leaf canopy shading, demonstrating the thermal buffering capacity of this training system under Mediterranean heat stress conditions.
Microclimate characterization is a terroir information entity that documents the localized atmospheric and edaphic environmental conditions within and immediately surrounding a grapevine canopy at spatial scales of centimeters to meters, including temperature, humidity, radiation, and air movement gradients that differ substantially from standard meteorological observations, and that directly govern leaf gas exchange, berry development, pathogen infection dynamics, and fruit composition at the individual vine or within-canopy level.
Microclimate characterization bridges the gap between standard weather station data and the actual environmental conditions experienced by grapevine organs, recognizing that the macroclimate recorded at regional stations may poorly represent the conditions governing physiological processes at the leaf and berry surface. The concept draws on the foundational work of Geiger (1965) on the climate near the ground, extended to vineyard systems by Smart (1985) and others. Key microclimate gradients exist vertically (from soil surface to canopy top), horizontally (from canopy exterior to interior, and between canopy faces), and temporally (diurnal cycles amplified or dampened within the canopy). Training system selection, row orientation, vine spacing, and canopy management collectively engineer the vine microclimate, making microclimate characterization essential for evaluating and optimizing these management interventions. Sensor miniaturization and wireless network technology have transformed microclimate monitoring from periodic manual measurement to continuous, spatially distributed data collection, enabling real-time management decisions.
Bestandsklima
Mikroklimacharakterisierung
Mikroklimadaten
Rebstockmikroklima
canopy environment characterization
micro-environmental data
microclimate description
mikroklimatische Beschreibung
vineyard microclimate data
within-canopy climate data
Smart RE (1985) Principles of grapevine canopy microclimate manipulation with implications for yield and quality: a review. American Journal of Enology and Viticulture 36(3): 230-239 Jackson DI, Lombard PB (1993) Environmental and management practices affecting grape composition and wine quality: a review. American Journal of Enology and Viticulture 44(4): 409-430 Dry PR (2000) Canopy management for fruitfulness. Australian Journal of Grape and Wine Research 6(2): 109-115
Microclimate Characterization
Micronutrient content quality management in Burgundy Pinot Noir addresses the iron-limiting calcareous soils that produce lime-induced chlorosis on susceptible rootstocks, with foliar iron chelate applications (EDDHA-Fe at 2-3 kg/ha) correcting the characteristic interveinal yellowing the selection of 41B and Fercal rootstocks with superior iron uptake from high-pH soils enables sustainable production on the limestone-rich Côte d'Or slopes where iron-sensitive rootstocks would fail, demonstrating how micronutrient management interacts with rootstock selection to determine viticultural viability.
Micronutrient_Content_Quality encompasses the chemical qualities characterizing the concentration of trace mineral elements required by grapevines in small quantities for essential enzymatic and metabolic functions. The micronutrients addressed in this category include iron, zinc, manganese, copper, boron, and molybdenum, which together with chlorine and nickel constitute the trace elements essential for plant growth and development despite being required in amounts typically less than 100 mg/kg dry matter. These elements function primarily as enzyme cofactors, activators, and structural components of metalloenzymes, participating in processes from photosynthesis and respiration through hormone synthesis and nitrogen metabolism. In grape berries, micronutrient concentrations are typically measured in mg/L or μg/L, with iron and manganese at the higher end (1-10 mg/L) and molybdenum at trace levels (μg/L range). The quality implications of micronutrients operate primarily through vine physiological effects rather than direct wine quality contributions, though certain micronutrients (iron, copper) can cause wine stability problems when present in excess. Micronutrient availability in soils depends strongly on pH, organic matter content, and interactions among elements, with deficiencies common in specific soil conditions: iron deficiency in calcareous soils (lime-induced chlorosis) zinc deficiency in sandy, low-organic matter soils boron deficiency in drought-stressed conditions and manganese deficiency in high-pH soils. Viticultural management of micronutrients employs soil applications, foliar sprays, or both, with chelated forms providing improved uptake efficiency in challenging soil conditions. The transfer of micronutrients from grape to wine depends on solubility, precipitation reactions, and processing losses, with wine micronutrient content rarely limiting quality except in cases of excess copper from vineyard sprays or iron causing oxidative instability.
Micronutrient_Content_Quality serves as the parent class for individual trace element qualities, positioned within Mineral_Content_Quality parallel to Macronutrient_Content_Quality. The classification reflects plant nutrition conventions distinguishing trace elements (required in mg/kg quantities) from macronutrients (required in g/kg quantities). Micronutrient analysis requires sensitive techniques (ICP-MS, graphite furnace AAS) due to the low concentrations involved. The quality implications of micronutrients operate primarily through vine physiology rather than direct wine effects, distinguishing them from macronutrients like potassium with direct wine chemistry impacts.
Mikroelementkonzentration
Mikronährstoffgehalt
Spurenelementgehalt
Spurenmineralzusammensetzung
microelement concentration
minor mineral quality
trace element content
trace mineral composition
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 341-385 Marschner H (2012) Marschner's Mineral Nutrition of Higher Plants, 3rd Ed, Academic Press pp.: 135-189
Micronutrient Content Quality
The micronutrient data for a Priorat vineyard (Catalonia, Spain) on Carboniferous llicorella (schist) documents the complete trace element profile — DTPA-iron 85 mg/kg (high, reflecting the iron-rich schist parent material and acid pH 5.8), DTPA-manganese 28 mg/kg (adequate), DTPA-zinc 1.8 mg/kg (adequate), hot water-boron 0.6 mg/kg (low-marginal, approaching the 0.5 mg/kg threshold below which boron deficiency symptoms may appear as poor fruit set and shot berries), DTPA-copper 12 mg/kg (elevated above natural background due to decades of copper fungicide application for downy mildew control) — this micronutrient profile indicates that the acid, iron-rich schist soil provides adequate availability of all cationic micronutrients but marginal boron supply, recommending preventive foliar boron applications (Solubor at 0.5 kg/ha) at pre-bloom and post-bloom to ensure adequate boron supply during the critical pollination and fruit set period.
A soil chemistry data component that records the concentrations of essential trace elements — iron (Fe), manganese (Mn), zinc (Zn), boron (B), copper (Cu), and molybdenum (Mo) — in the vineyard soil, determined by standardized extraction methods (DTPA chelate extraction for Fe, Mn, Zn, and Cu hot water extraction for B ammonium oxalate extraction for Mo), with micronutrient availability being strongly pH-dependent (most micronutrients becoming less available above pH 7.5 and more available below pH 5.5) and the primary soil chemical factor responsible for the nutritional disorders — lime-induced iron chlorosis, manganese deficiency or toxicity, zinc deficiency, and boron deficiency or toxicity — that constrain vine performance on specific soil types.
Micronutrient_Data documents the trace element availability that, despite the small quantities required by the vine (milligrams per vine per year compared to grams for macronutrients), can be the limiting factor for vine performance on specific soil types. The pH-dependent solubility of micronutrient cations (Fe, Mn, Zn, Cu) is the dominant control on their availability — at pH values above 7.5 (calcareous soils), these cations form insoluble hydroxide precipitates (Fe(OH)₃, MnO₂) that reduce solution concentrations below the minimum required for root uptake, while at pH values below 5.5 (strongly acid soils), the same cations become excessively soluble and potentially toxic, particularly manganese (which can reach phytotoxic levels of >100 mg/kg DTPA-Mn on acid, poorly drained soils): this pH dependence means that calcareous vineyard soils and acid vineyard soils present opposite micronutrient challenges — calcareous soils risk deficiency (particularly iron chlorosis), while acid soils risk toxicity (particularly manganese). The copper accumulation in vineyard soils deserves special attention — the Bordeaux mixture (CuSO₄ + Ca(OH)₂) has been used as the primary downy mildew fungicide since 1885, and 130+ years of annual copper application have elevated topsoil copper to 50-200 mg/kg in many traditional European vineyard soils (compared to natural background of 5-30 mg/kg), creating a legacy contamination that poses toxicity risk to cover crops and young vine roots on replanted sites, and that persists for centuries due to the strong binding of copper to soil organic matter and iron oxides.
Mikronährstoff-Daten
Mikronährstoff-Verfügbarkeit
Spurenelement-Analytik
essential trace element data
micronutrient availability record
trace element analysis
White RE (2003) Soils for Fine Wines, Oxford University Press Bavaresco L et al. (2003) Role of rootstock on iron-chlorosis, Acta Horticulturae 603: 87-95 Komárek M et al. (2010) Contamination of vineyard soils with fungicides: a review of environmental and toxicological aspects, Environment International 36: 138-151 Robinson JB (2005) Critical plant tissue values, ASEV
Micronutrient Data
Micronutrient deficiency symptom quality assessment in Champagne vineyards on calcareous soils identified multiple concurrent trace element limitations requiring integrated corrective management. Systematic visual survey revealed micronutrient deficiency symptom quality patterns affecting youngest tissues consistent with immobile nutrient limitations, with interveinal chlorosis on young leaves suggesting iron and/or manganese deficiency, and small malformed leaves at shoot tips indicating possible zinc limitation. The characteristic localization of micronutrient deficiency symptom quality on apical canopy positions confirmed immobile nutrient involvement, distinguishing from the basipetal symptoms of mobile macronutrient deficiencies that would affect oldest leaves first. Soil analysis established the context for micronutrient deficiency symptom quality with pH 8.2 and active lime content at 12%, conditions known to reduce availability of Fe, Zn, and Mn through precipitation as carbonates and hydroxides. Tissue analysis of symptomatic shoot tips confirmed multiple micronutrient deficiency symptom quality diagnoses with Fe at 45 ppm (deficient, threshold 60 ppm), Zn at 12 ppm (deficient, threshold 20 ppm), and Mn at 18 ppm (marginal, threshold 20 ppm), establishing that the high-pH soil conditions were creating concurrent limitations of multiple trace elements. Corrective management integrated foliar application of chelated micronutrient formulation (Fe-EDDHA, ZnSO₄, MnSO₄) with soil acidification using elemental sulfur to improve long-term micronutrient availability, with symptom resolution within 4 weeks of foliar treatment confirming the micronutrient deficiency symptom quality diagnoses.
Micronutrient deficiency symptom quality is a nutrient deficiency indicator quality that inheres in a grapevine organism and represents the visual symptoms and physiological impairments resulting from inadequate availability of essential trace elements including iron, zinc, manganese, boron, copper, and molybdenum, with micronutrient deficiencies typically producing more localized or specific symptoms than macronutrient deficiencies reflecting the specialized roles of these elements as enzyme cofactors and metabolic regulators, and with most micronutrients being phloem-immobile such that deficiency symptoms characteristically appear on youngest tissues that cannot acquire these elements through redistribution from older organs. This quality captures the grapevine response to trace element limitations, recognizing that while micronutrients are required in much smaller quantities than macronutrients (mg/kg versus g/kg tissue concentrations), they are equally essential for plant function and their deficiency can cause severe symptoms and substantial production losses. The ontological significance of micronutrient deficiency symptom quality lies in the diagnostic specificity of many micronutrient deficiency symptoms enabling relatively precise identification of limiting elements, while also presenting challenges where similar symptom patterns from different deficiencies require careful differential diagnosis considering symptom distribution, associated features, and tissue analysis confirmation. The biological basis of micronutrient deficiency symptoms encompasses the roles of these elements as essential components of metalloenzymes and enzyme activators, with iron required for cytochromes and iron-sulfur proteins in electron transport, zinc required for carbonic anhydrase and numerous zinc-finger transcription factors, manganese required for the oxygen-evolving complex of photosystem II, boron required for cell wall structure, and copper required for plastocyanin and various oxidases. The phloem immobility of most micronutrients (Fe, Zn, Mn, Cu, B) results in characteristic symptom expression on youngest tissues and growing points, contrasting with the basipetal symptom progression of mobile macronutrients, and providing important diagnostic criteria for distinguishing micronutrient from macronutrient deficiencies. The soil chemistry of micronutrient availability creates characteristic deficiency situations, with high soil pH reducing availability of Fe, Zn, Mn, and Cu through precipitation and adsorption reactions, making calcareous soils the primary environment for multiple micronutrient deficiencies in viticulture.
Micronutrient_Deficiency_Symptom_Quality is_a Nutrient_Deficiency_Indicator_Quality, serving as parent class for Iron_Deficiency_Symptom_Quality, Zinc_Deficiency_Symptom_Quality, and future classes addressing Mn, B, Cu, Mo deficiencies. This class addresses the general features of trace element limitation while child classes capture element-specific symptoms. Most micronutrients are phloem-immobile showing symptoms on youngest tissues first. High soil pH (calcareous soils) is the primary predisposing factor for multiple micronutrient deficiencies. Tissue concentrations in mg/kg (ppm) versus g/kg (%) for macronutrients.
Mikroelement-Limitation
Mikronährstoffmangel-Symptome
Spurenelement-Mangel
Spurenmineral-Mangel
micronutrient limitation
minor element deficiency
trace element deficiency
trace mineral deficiency
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication Marschner H (2012) Marschner's Mineral Nutrition of Higher Plants, 3rd Edition. Academic Press
Micronutrient Deficiency Symptom Quality
Micronutrient uptake requirements are orders of magnitude lower than macronutrients, with annual vine demand of 200-800 g Fe/ha, 100-400 g Zn/ha, 200-600 g Mn/ha, and 100-300 g B/ha for a mature vineyard. Rootstock effects on micronutrient uptake are pronounced for iron (3-5× variation) and zinc (2-3× variation) but modest for manganese and boron. Soil pH strongly modulates micronutrient availability, with Fe, Zn, Mn, and B becoming less available above pH 7.5, creating deficiency challenges on calcareous soils. Micronutrient deficiency symptoms appear at leaf concentrations below critical thresholds specific to each element and growth stage.
A nutrient acquisition disposition that is the inherited capacity of a Vitis vinifera plant to absorb and accumulate essential trace elements (iron, zinc, manganese, boron, copper, molybdenum, chlorine, nickel) required in small quantities for specific enzymatic and structural functions, through specialized root transporter systems operating under typically low soil solution concentrations.
Micronutrient_Uptake_Disposition serves as the parent class for individual trace element uptake traits. Despite low quantitative requirements, micronutrient deficiencies cause severe physiological disruption due to essential roles in enzyme function, electron transport, and structural components. Rootstock selection is the primary tool for managing micronutrient acquisition on problem soils, with genotypes showing wide variation in uptake efficiency. Chelate chemistry, soil pH management, and foliar application provide supplementary management options.
Mikroelement-Aneignung
Mikronährstoffaufnahme
Spurenelementakquisition
micronutrient foraging capacity
minor element acquisition
trace element uptake
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 325-345 Robinson JB (1992) Grapevine nutrition. In: Coombe BG, Dry PR (eds) Viticulture Volume 2: Practices, Winetitles pp.: 178-208
Micronutrient Uptake Disposition
Micronutrient uptake efficiency assessment on calcareous soil (pH 8.2, 35% active lime) reveals the site-specific challenges for trace element acquisition, with iron uptake severely limited (leaf Fe 45 mg/kg, deficient) causing interveinal chlorosis despite adequate soil total Fe, zinc marginally adequate (leaf Zn 22 mg/kg) from high pH reducing availability, and boron adequate (leaf B 35 mg/kg) as its availability is less pH-sensitive, demonstrating how soil chemistry creates element-specific uptake challenges requiring site-appropriate rootstock selection and potentially foliar supplementation.
Micronutrient uptake efficiency quality is a nutrient uptake efficiency quality that represents the capacity of grapevine root systems to acquire essential trace elements required in small quantities but playing critical catalytic and structural roles in enzyme function, electron transport, and hormone synthesis that affect vine physiology disproportionately to the quantities involved. Within the BFO framework, this quality specifically addresses the trace element component of nutrient uptake, distinguishing micronutrient acquisition from macronutrient uptake while recognizing that the small quantities required do not diminish the importance of efficient uptake, as micronutrient deficiencies can severely limit vine function even when macronutrient supply is adequate. The ontological significance lies in micronutrient uptake efficiency determining access to the catalytic elements essential for photosynthesis, respiration, nitrogen metabolism, and hormone function, where iron, zinc, manganese, copper, boron, molybdenum, and chlorine each serve irreplaceable roles such that deficiency of any single micronutrient can become the limiting factor for vine productivity regardless of other nutrient adequacy. The mechanistic basis for micronutrient uptake involves specific transport proteins for each element, often with both high-affinity systems operating at low soil concentrations and low-affinity systems functioning when elements are more abundant, with transporter expression typically regulated by plant nutritional status to match acquisition to demand and prevent toxicity from excess accumulation. The soil chemistry challenges for micronutrient availability are substantial, as most micronutrients show strong pH-dependent availability (iron, zinc, manganese becoming less available at high pH molybdenum becoming less available at low pH), creating site-specific nutritional challenges that uptake efficiency must address. The rootstock effects on micronutrient uptake are particularly important in viticulture, with some rootstocks (especially those with Vitis berlandieri parentage) showing reduced iron uptake efficiency on calcareous soils (lime-induced chlorosis), while others show enhanced zinc or boron acquisition, making rootstock selection critical for micronutrient nutrition on challenging sites. Understanding micronutrient uptake efficiency quality enables assessment of vine capacity for trace element acquisition, rootstock selection for site-specific micronutrient challenges, and diagnosis of micronutrient limitations affecting vine function.
Micronutrient_Uptake_Efficiency_Quality serves as the parent class for specific trace element uptake qualities (Iron, Zinc, Boron). This quality addresses the acquisition of essential catalytic elements with critical physiological roles. The class supports micronutrient management and rootstock selection through trace element uptake efficiency characterization.
Mikronährstoffakquisition
Mikronährstoffaufnahmeeffizienz
Spurenelementaufnahme
Spurenelementeffizienz
essential element efficiency
micronutrient acquisition
minor element uptake
Marschner H (2012) Marschner's Mineral Nutrition of Higher Plants, 3rd Ed. Academic Press, London, pp. 135-189 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 197-245
Micronutrient Uptake Efficiency Quality
Syrah in Rhône Valley shows micropyle targeting during mid-late June Year N+1 at BBCH 65-69, with pollen tubes detecting synergid attractant signals within 1.5-2.5 mm of ovules, chemotropic growth reorientation navigating final 1-3 mm over 1-2.5 hours, precise arrival at 120-180 micrometer micropyle openings, under warm Mediterranean conditions of 25-29°C temperature, typical of this mid-to-late season cultivar with 2-3 tubes successfully targeting 3-4 available ovules per flower 24-36 hours post-pollination
Micropyle targeting represents the precise tube navigation phase when tube growing in ovary locule detects and responds to synergid-derived attractant signals directing growth toward specific ovule micropyle opening, achieving accurate targeting of 100-200 micrometer micropyle entrance from millimeter distances enabling embryo sac access. This critical targeting phase occurs during pollen tube penetration in mid-late June in temperate regions Year N+1, typically proceeding over final 1-3 hours of tube growth as ovule approached, corresponding to BBCH 65-69 fertilization stages. Targeting mechanism involves tube detection of diffusible attractant signals emanating from synergid filiform apparatus creating concentration gradient, chemotropic growth response reorienting tube trajectory toward highest attractant concentration, progressive approach refining navigation as gradient strengthens, and precise arrival at micropyle outer opening completing targeting. Synergid attractant signals show secretion from filiform apparatus in synergid cells adjacent to egg, diffusible peptide or protein signals creating concentration gradient extending into ovary locule, possible LURE-type peptides identified in other species serving as attractants though specific grapevine signals unknown, and integrated signal profile providing directional information guiding tube to micropyle. Attractant gradient characteristics include source at micropyle creating highest concentration, declining concentration with distance following diffusion pattern, gradient detectable at millimeter scale enabling long-range guidance, and sufficient signal strength for tube detection and response. Tube chemotropic response shows sensitive detection of attractant concentration differences with femtomolar to picomolar sensitivity in some systems, growth reorientation toward higher concentration through asymmetric tip growth, and progressive directional adjustment refining trajectory as micropyle approached. Growth trajectory modification shows tube initially approaching ovary region from style base, detection of attractant gradient triggering directional change, progressive reorientation over 1-3 mm final growth aligning with micropyle position, and accurate targeting achieving arrival at 100-200 micrometer micropyle opening despite complex three-dimensional ovary geometry. Targeting precision shows remarkable accuracy with tube navigating directly to small micropyle opening, successful targeting from 1-3 mm initial detection distance, minimal trajectory error enabling first-attempt micropyle location without extensive searching, and precision suggesting sophisticated chemotropic sensing and response mechanisms. Multiple ovule targeting dynamics show 2-4 ovules per flower potentially producing attractant signals, competitive situation with 2-5 tubes typically in ovary but only 1-2 achieving fertilization, possible preferential targeting of specific ovules though mechanisms unclear, and first-arrival advantage with earliest tube to reach responsive ovule likely succeeding. Ovule-to-ovule variation in attractant production possibly includes developmental differences affecting signal strength, some ovules producing stronger signals receiving preferential targeting, and asynchronous ovule maturation creating temporal variation in targeting opportunities. Tube-tube competitive targeting shows multiple tubes potentially detecting same ovule signals, competitive approach with fastest or most sensitive tube gaining access, and unsuccessful tubes potentially redirecting to alternative ovules or failing to achieve fertilization. Targeting distance shows detection occurring when tube within approximately 1-3 mm of micropyle based on gradient strength, closer approach enabling stronger signal and more precise navigation, and final arrival requiring sustained chemotropic response over 1-3 hour approach period. Growth rate during targeting shows sustained velocity 1-2 mm per hour during final ovary navigation, brief 1-3 mm growth distance from initial detection to micropyle arrival, and total targeting duration typically 1-3 hours representing final tube growth phase. Calcium signaling in targeting possibly includes calcium gradient associated with attractant signal, localized calcium accumulation at tube tip mediating chemotropic response, and calcium-dependent signaling affecting growth direction though specific roles in grapevine poorly characterized. Environmental effects on targeting include temperature affecting tube metabolic activity and growth rate influencing targeting dynamics, continued boron requirement for tube integrity during final approach, and overall favorable conditions 22-28°C supporting efficient targeting. Ovule maturity coordination with targeting shows embryo sac development synchronized to produce attractant signals when tubes likely to arrive, temporal matching between tube arrival timing 20-40 hours post-pollination and ovule receptivity, and developmental coordination essential for successful targeting and fertilization. Failed targeting causes include inadequate attractant production from ovule dysfunction, tube chemotropic dysfunction preventing signal detection or response, competitive exclusion by faster tubes monopolizing available ovules, or structural abnormalities in ovary preventing access despite successful navigation. Targeting verification methods include microscopic observation of tube trajectories, fluorescence visualization of tubes approaching micropyles, and experimental manipulation of attractant signals affecting targeting success. The process represents first aspect of pollen tube penetration achieving precise navigation to entry point, followed by synergid penetration completing access. Duration typically spans 1-3 hours representing final targeting approach phase. Cultivar variation in targeting may include ovule attractant production affecting signal strength, tube chemotropic sensitivity affecting detection and response, and ovule number affecting targeting opportunities and competition. Regional timing shows micropyle targeting occurring during mid-late June across all climates 20-38 hours after pollination as final tube approach phase. Viticultural significance includes representing critical precision navigation enabling embryo sac access, demonstrating sophisticated tube-ovule communication, requiring successful completion of entire tube growth journey, and establishing targeting success as prerequisite for penetration and fertilization. Research applications include identifying attractant signals and chemotropic mechanisms, investigating tube-ovule signaling and communication, evaluating targeting precision and accuracy, quantifying competitive dynamics among multiple tubes, and assessing targeting success as determinant of fertilization efficiency.
Micropyle_Targeting is first aspect of Pollen_Tube_Penetration, distinguished by precise chemotropic navigation to micropyle opening. Represents remarkable precision achieving 100-200 micrometer target from millimeter distance. Duration 1-3 hours during final approach. Synergid-derived attractant signals guide navigation. Multiple tubes compete for limited ovules. Essential prerequisite for Synergid_Penetration and embryo sac access. Sophisticated tube-ovule communication system.
Locksignal-Reaktion
Mikropyle-Zielerfassung
Samenanlage-Zielansteuerung
attractant response
micropyle location
ovule targeting
precise navigation
präzise Navigation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 390-392 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 295-297
Micropyle Targeting
The microsatellite marker data for the complete set of markers used in the European Vitis Database cultivar identification program documents 20 core SSR loci drawn from four independently developed marker series — 6 VVMD markers (developed at UC Davis by Bowers et al. 1996, 1999), 1 VVS marker (developed at CSIRO by Thomas and Scott 1993), 2 VrZAG markers (developed at BOKU Vienna by Sefc et al. 1999), and 11 supplementary VMC markers (developed by Vitis Microsatellite Consortium) — each characterized by dinucleotide or trinucleotide repeat motif, chromosomal position on the PN40024 reference genome, allele size range calibrated against the This et al. (2004) reference allele ladder, and marker-specific quality metrics including polymorphism information content (PIC 0.65-0.89), null allele frequency (<0.05 for all core markers), and genotyping error rate (<2% in inter-laboratory ring tests).
A genetic marker data record specific to simple sequence repeat (SSR) markers developed for Vitis, documenting the marker name, originating research group, repeat motif, primer pair sequences, chromosomal assignment, allele size range, polymorphism information content, and cross-species transferability that characterize each microsatellite locus as a tool for grapevine genotyping, with multiple independently developed marker series (VVMD, VVS, VrZAG, VMC, and others) collectively providing a toolkit of over 300 characterized SSR markers distributed across the 19 Vitis chromosomes.
Microsatellite_Marker_Data documents the properties of SSR markers as genotyping tools — the technical specifications that determine each marker's suitability for specific applications. The development of Vitis microsatellite markers proceeded through multiple independent research programs during the 1990s and early 2000s, each producing a named marker series identified by a prefix reflecting the originating institution or project: this parallel development created a large marker toolkit but also introduced nomenclatural complexity, as markers from different series mapping to the same chromosomal region may provide redundant information while markers with similar names from different series are entirely distinct loci. The standardization effort led by This et al. (2004) under OIV coordination selected the optimal subset of 9 markers from across series based on performance criteria (polymorphism, reliability, transferability) and established the reference allele system that enables cross-laboratory data comparison — this standardization was essential for building the international cultivar genotype databases (VIVC, European Vitis Database) that now serve as the definitive molecular reference for grapevine germplasm identification. Subclasses document the four principal marker series.
Mikrosatelliten-Lokus-Charakterisierung
Mikrosatelliten-Marker-Daten
SSR marker specification
SSR-Marker-Spezifikation
STR marker documentation
microsatellite locus characterization
This P et al. (2004) Development of a standard set of microsatellite reference alleles for identification of grape cultivars, Theoretical and Applied Genetics 109: 1448-1458 Sefc KM et al. (2001) Microsatellite markers for grapevine, in Roubelakis-Angelakis KA (ed) Molecular Biology and Biotechnology of the Grapevine, Kluwer pp: 433-463 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC), Vitis 54: 155-157
Microsatellite Marker Data
Pinot Noir mid developing berries progress through lag phase at BBCH 77 (weeks 6-8 post-fruit set) maintaining relatively constant diameter of 9-10.5 mm and fresh weight of 0.9-1.2 g with minimal growth, green coloration, hard texture, peak acidity of 22-24 g/L, and sugars at 7-9%, with seeds completing maturation reaching maximum size (5.5-7 mm length) and brown lignified seed coats, then undergoing veraison at BBCH 81-83 (weeks 9-11) with color change from green to red-purple through chlorophyll degradation and anthocyanin accumulation (beginning at 0.1 mg/g skin increasing to 0.8-1.5 mg/g by mid-veraison), softening initiated, sugars increasing to 13-16%, and acidity declining to 16-19 g/L. Chardonnay white wine cultivar mid developing berries show lag phase (BBCH 77, weeks 7-8.5) with stable dimensions at 10-11.5 mm diameter, green coloration, 8-9% sugars, 18-20 g/L acidity, then veraison (BBCH 81-83, weeks 9-11) marked by color change from green to golden-yellow through chlorophyll degradation and carotenoid/flavonol changes, softening, sugar accumulation to 14-17%, acid decline to 13-16 g/L, demonstrating veraison transitions in white cultivar where color change is subtle compared to red cultivars but metabolic transitions are comparable. Syrah mid developing berries examined during lag phase (BBCH 77) show seed development at mature embryo stage with embryos fully formed at 4-5.5 mm length, seed coats lignifying and browning, seed dry weight reaching 18-22 mg approaching maximum (24-28 mg at final seed maturity), while berry mesocarp tissues show minimal growth activity with low mitotic index (<1% cells dividing) and metabolic reprogramming evidenced by shifts in gene expression from growth-related to ripening-related transcripts. Merlot experiencing different veraison temperatures shows 22-25°C producing optimal anthocyanin accumulation (1.8-2.4 mg/g skin at BBCH 83) versus 28-32°C reducing anthocyanin (1.2-1.7 mg/g) and versus 18-20°C delaying anthocyanin synthesis but eventually achieving good levels (1.6-2.2 mg/g with 1-2 week delay), demonstrating temperature effects on veraison color development critical for red wine quality.
A mid developing berry is a developing berry in the intermediate development phase spanning the lag phase and veraison (BBCH 77-83, approximately 6-11 weeks post-fruit set), characterized by minimal size change during lag phase followed by onset of ripening changes at veraison including color change, softening, and metabolic reprogramming, representing the transition period between vegetative growth and reproductive ripening. Mid developing berries are characterized by= (1) minimal size increase during lag phase (BBCH 77, weeks 6-8) with berry diameter increasing only 0-8% from end of Phase I dimensions, maintaining relatively constant size at 6-12 mm diameter depending on cultivar (2) fresh weight remaining relatively stable during lag phase at 0.6-1.8 g depending on cultivar, representing the pause between Phase I and Phase II growth (3) cells achieving final number during early lag phase with essentially zero cell division after BBCH 75-77, and cells maintaining relatively constant size (45-70 μm diameter) during lag phase before expansion begins at veraison (4) seeds undergoing rapid development and maturation during lag phase with embryos completing maturation, seed coats lignifying and browning, and seeds achieving maximum size and near-maximum dry weight by end of lag phase, or seeds arresting and beginning degeneration in stenospermocarpic seedless types (5) color changes at veraison (BBCH 81) with chlorophyll degradation and anthocyanin synthesis (red/black cultivars) or carotenoid/flavonol changes (white cultivars), creating transition from green to colored berries (6) texture changes beginning at veraison with softening initiated through cell wall modifications and beginning of cell expansion (7) sugars beginning to accumulate at veraison increasing from lag phase levels of 6-10% to 12-16% by BBCH 83 (mid-veraison), with sugar accumulation rate accelerating as ripening progresses. Mid developing berries undergo critical metabolic transitions= lag phase metabolic reprogramming shifting from growth-focused to ripening-focused gene expression patterns veraison representing the single most dramatic developmental transition in berry development with coordinated changes in color, texture, sugar accumulation, acid metabolism, secondary metabolite synthesis, and hormone balance transition from malic acid synthesis to malic acid respiration/degradation at veraison causing acid decline activation of secondary metabolite biosynthesis pathways producing anthocyanins, flavor compounds, and aromatic precursors. Mid developing berries show changing photosynthetic activity= lag phase berries maintaining moderate photosynthesis (1-3 μmol CO₂ m⁻² s⁻¹) contributing 5-10% of carbon requirements veraison berries showing rapid chlorophyll degradation and photosynthesis decline to near zero as color change proceeds post-veraison berries photosynthetically inactive becoming entirely dependent on imported photoassimilates from leaves. Environmental factors during mid development affect critical quality parameters= temperature during veraison influences color development, with moderate temperatures (22-28°C day, 12-18°C night) optimal for anthocyanin synthesis water deficit during lag phase and veraison can reduce berry size and affect composition through reduced cell expansion and concentrated metabolites light exposure during veraison enhances color development and secondary metabolite accumulation.
Mid_Developing_Berry represents the lag phase and veraison period (BBCH 77-83, weeks 6-11) and is distinguished from Early_Developing_Berry (Phase I growth, weeks 0-6) by minimal growth versus rapid cell division, and from late developing berry/ripening (Phase II, weeks 11-18+) by initiation versus completion of ripening processes. Mid developing berries undergo the most dramatic developmental transition (veraison) and critical seed maturation, making this period essential for quality development. Sub-classification into Active_Growth_Mid (actually a misnomer as lag phase shows minimal growth, better interpreted as active seed maturation) and Expanding_Mid (veraison with beginning expansion and ripening changes) reflects the lag phase versus veraison distinction within mid development.
Reifeansatz-Stadium
mittel-entwickelnde Beere
Übergangsphasen-Beere
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 192-212 Coombe, B.G., McCarthy, M.G. (2000). Dynamics of grape berry growth. Australian Journal of Grape and Wine Research 6=131-135 Castellarin, S.D. et al. (2007). Water deficits accelerate ripening. Functional Plant Biology 34=174-186.
Mid Developing Berry
A mid-development aborted tendril at node 8 on a moderately water-stressed shoot of Vitis vinifera cv. Sangiovese in Chianti, Italy, measuring 5.8 cm length (approximately 35% of the normal 16-18 cm for functional tendrils), exhibiting well-formed bifurcated structure with both arms equally developed to 2.5 cm, complete vascular differentiation visible through semi-translucent green-brown tissue, partial brown discoloration starting at tips and progressing basally, occurring during managed water deficit irrigation strategy (soil moisture maintained at 60-70% field capacity) applied during veraison to enhance fruit quality, with 25% of tendrils showing similar mid-development abortion across the irrigated block compared to <10% in fully irrigated control sections, representing an acceptable trade-off of vegetative function for improved berry composition including enhanced anthocyanin and phenolic development measured at BBCH stage 83 (berries developing color) in early August.
A late aborted tendril that ceased development during the middle phase of advanced morphogenesis in Vitis vinifera, characterized by arrested structures of 4-7 cm length representing approximately 25-40% of normal mature tendril length for the cultivar and position, presence of complete basic tissue organization including fully differentiated vascular bundles, functional epidermis with partial cuticle development, and recognizable tendril architecture including branching points if typical for the position, but incomplete final cell expansion in elongation zones, underdeveloped mechanosensory apparatus with reduced or absent contact-responsive epidermal cells, and insufficient structural reinforcement through cell wall thickening and lignification for bearing mechanical loads. Mid-development aborted tendrils result from moderate stress conditions or resource limitations occurring during the intermediate development period approximately 1-2 weeks after initial elongation when tendrils are transitioning from rapid growth to functional maturation, commonly caused by progressive water stress, gradual onset nutrient deficiency particularly nitrogen or potassium affecting cell expansion and wall synthesis, increasing competition from fruit cluster development during the lag phase of berry growth, chronic pest feeding damage, or prolonged suboptimal temperatures slowing metabolic processes without causing acute stress responses. These abortion events represent intermediate resource investment loss and indicate moderate but significant stress conditions that compromise vegetative development while potentially allowing reproductive success, creating common scenarios in commercial viticulture where managed stress improves fruit quality at the expense of vegetative vigor.
Mid-development aborted tendrils are distinguished from early late-aborted tendrils (initial growth failures achieving 60% normal length) by intermediate developmental achievement representing substantial but incomplete morphogenesis. This abortion class is diagnostically valuable in commercial viticulture because it often correlates with intentional managed stress strategies including deficit irrigation or crop load manipulation aimed at balancing vegetative and reproductive development, with moderate tendril abortion acceptable when fruit quality benefits justify reduced climbing support capacity. The occurrence threshold requiring intervention depends on management objectives, with wine quality-focused production tolerating higher mid-development abortion rates (20-30%) compared to fresh fruit production requiring maximum vegetative vigor (<15%). Mid-development abortion combined with reduced leaf size and shoot vigor indicates appropriate stress levels for premium wine production.
Zwischenphase fehlgeschlagene Ranke
halb-elongierte abgebrochene Kletterorgan
half-elongated aborted tendril
intermediate development arrested tendril
middle phase failed tendril
mittlere Entwicklung-abortierte Ranke
Based on Matthews & Anderson (1988) "Fruit ripening in Vitis vinifera L.= responses to seasonal water deficits" American Journal of Enology and Viticulture 39=313-320 Chaves et al. (2010) "Grapevine under deficit irrigation= hints from physiological and molecular data" Annals of Botany 105=661-676 Dry et al. (2001) "Strategic irrigation management in Australian vineyards" Journal International des Sciences de la Vigne et du Vin 35=129-139 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 425-448.
Mid Development Aborted Tendril
Cabernet Sauvignon exhibits mid-season budbreak disposition, reaching BBCH 09 within ±5 days of the cultivar median in comparative trials, with heat sum requirement of 80-110 GDD (base 10°C). Merlot and Syrah show similar intermediate timing, providing balanced frost risk and season length across diverse growing regions. Mid-season cultivars in Napa Valley initiate budbreak during the second week of March, approximately 10 days after Chardonnay and 10 days before Mourvèdre. The coefficient of variation for budbreak date in mid-season cultivars (12-18%) is typically lower than for early cultivars (15-22%) due to more stable spring temperature conditions at later calendar dates.
A budbreak timing disposition that is the inherited capacity of a Vitis vinifera plant to initiate budbreak at intermediate timing relative to the species range, typically within the second tercile of cultivar budbreak dates, balancing frost avoidance with adequate growing season length.
Mid-season budbreak disposition represents the adaptive optimum for many traditional wine regions, balancing frost avoidance with adequate heat accumulation for complete ripening. The category encompasses the majority of commercially important wine grape cultivars, reflecting centuries of empirical selection for regional climate matching. Mid-season cultivars offer flexibility in site selection and provide benchmark timing for vineyard planning and frost protection scheduling.
Normalaustrieb
average budburst timing
durchschnittliches Knospentreiben
intermediate budbreak
mid-spring phenology
mittlerer Austrieb
Parker AK et al. (2013) Classification of varieties for their timing of flowering and veraison using a modelling approach: a case study for the grapevine species Vitis vinifera L. Agricultural and Forest Meteorology 180:249-264 Van Leeuwen C, Darriet P (2016) The impact of climate change on viticulture and wine quality. Journal of Wine Economics 11:150-167
Mid Season Budbreak Disposition
A mid-season prompt lateral shoot on Vitis vinifera cv. Pinot Noir arising from node 7 axillary bud in early July, approximately 8 weeks after primary shoot formed this node during late April-early May elongation, emerged following hedge-topping operation in late June removing primary shoot apex and releasing lateral buds from apical dominance, developed to 42cm length by late August with 6 leaves totaling approximately 630 cm² leaf area, representing typical mid-season lateral response to hedging in moderate-vigor Pinot Noir, limited development period (8 weeks versus 12+ for early laterals) constraining final size to moderate dimensions, this lateral retained contributing valuable supplemental photosynthetic capacity (23% increase over primary shoot alone) supporting single-cluster crop without creating excessive density problems in well-spaced VSP canopy, with mid-season timing providing benefits-to-costs balance favoring retention.
A prompt lateral shoot emerging mid-growing season after primary shoot elongation substantially advanced, characterized by activation 6-10 weeks after node formation, emergence during slowing primary growth phase (typically late June-July), moderate growing period enabling intermediate lateral development (typically 25-55cm length), and function as supplemental late-season photosynthetic capacity with reduced density impact compared to early laterals due to later timing and limited development period. Mid-season prompt laterals represent delayed axillary activation occurring after peak spring growth, arising through triggers including hedging-induced apical dominance release, mid-season environmental conditions favoring lateral breakage, or variety-specific mid-season lateral tendencies, with their intermediate timing creating moderate development producing photosynthetic benefits without extreme density problems characteristic of vigorous early laterals.
Mid-season prompt laterals occupy intermediate position between early vigorous laterals (creating potential density problems) and late minimal laterals (providing limited benefits), with their moderate timing and development creating pragmatic balance often favorable for retention decisions. The mid-season emergence (late June-July typical) provides sufficient growing period (2-3 months) for meaningful development contributing photosynthetic capacity, while limited time compared to early laterals constrains excessive growth preventing severe density problems. Mid-season lateral activation often results from management interventions (hedging releasing lateral buds, shoot positioning stimulating lateral activation) or environmental triggers (mid-season rainfall or temperature changes stimulating growth), with predictable timing enabling planned management responses. Management frequently involves selective retention of well-positioned mid-season laterals recognizing photosynthetic benefits without excessive density costs, removal of poorly-positioned or redundant laterals, or systematic retention in moderate-vigor situations where supplemental leaf area valuable for supporting crop maturation, with decisions reflecting balance between benefits and management complexity.
Sommernebentrieb
delayed prompt lateral
mid-timing prompt lateral
mittlerer prompter Seitentrieb
summer lateral shoot
Mid Season Prompt Lateral Shoot
Cabernet Sauvignon exhibits mid-season ripening disposition, reaching 23-25°Brix approximately 50-60 days post-veraison with total season heat requirement of 1400-1600 GDD (base 10°C) from budbreak. Merlot, Syrah, and Sangiovese show similar intermediate ripening, completing harvest during the middle third of the regional vintage window. Mid-season ripening in Cabernet Sauvignon at Bordeaux reference sites occurs at mean date of October 1-15, bracketed by earlier Merlot (September 20-30) and later Petit Verdot (October 15-25). The mid-season category encompasses the majority of classic wine grape cultivars, reflecting historical selection for reliable maturation in traditional European wine regions.
A ripening timing disposition that is the inherited capacity of a Vitis vinifera plant to reach harvest maturity at intermediate timing relative to the species range, typically within the middle two quartiles of cultivar harvest dates for a region, requiring moderate to substantial heat accumulation and suited to temperate climates with reliable autumn conditions.
Mid-season ripening disposition represents the adaptive center for temperate climate viticulture, balancing heat requirement against growing season length constraints. The category includes most internationally planted wine cultivars, which were selected over centuries for consistent ripening in their regions of origin. Mid-season cultivars show greatest flexibility in site selection, performing acceptably across a range of mesoclimates within suitable macroclimatic zones. Climate warming is shifting mid-season cultivars toward earlier harvest dates, with potential implications for wine style and regional typicity.
Normalreife
average maturation timing
intermediate ripening
mid-harvest trait
mittlere Reife
mittlerer Erntetermin
Jones GV, Davis RE (2000) Climate influences on grapevine phenology, grape composition, and wine production and quality for Bordeaux, France. American Journal of Enology and Viticulture 51:249-261 Gladstones J (1992) Viticulture and Environment, Winetitles pp.: 98-135
Mid Season Ripening Disposition
Thompson Seedless (Sultana) bears a Mid_Season_Table_Grape_Role as the global benchmark variety, reaching commercial maturity (18-20°Brix) in late July to August in California's San Joaquin Valley, defining the reference point against which other varieties are classified as early or late. Crimson Seedless serves in a Mid_Season_Table_Grape_Role for mid-to-late season red seedless, reaching optimal harvest condition (17-19°Brix, full crimson color) in August-September in California, with peak production coinciding with maximum market supply and baseline pricing. Red Globe bears a Mid_Season_Table_Grape_Role in its primary production regions (California, Chile, Peru), reaching harvest in August-September (Northern Hemisphere) with the large berry size and seeded character valued in Asian export markets.
A table grape scion role that inheres in a grafted scion cultivar by virtue of its maturation timing coinciding with the main harvest season, typically reaching commercial harvest during the peak production period when volume is highest and prices are at seasonal baseline levels. Mid-season table grape scions mature at timing typical for the production region—approximately July-September in California, December-March in Southern Hemisphere counter-season regions—requiring moderate heat accumulation (typically 1800-2200 GDD base 10°C) that aligns with the bulk of commercial harvest windows. Mid-season varieties benefit from optimal growing conditions during the warmest period, achieving full sugar accumulation, color development (for pigmented varieties), and flavor maturation without the compromises of early or late production. While lacking the price premiums of early varieties or the storage advantages of late varieties, mid-season cultivars typically offer the best combination of yield potential, fruit quality, and production reliability, and constitute the majority of global table grape production volume.
Mid_Season_Table_Grape_Role represents the volumetric core of table grape production where most fruit is harvested during peak seasonal conditions. The classification is relative to production region—August harvest is mid-season in California but would be early for cool-climate regions and late for tropical production. Mid-season varieties face the most intense market competition due to supply concentration, requiring differentiation through quality, branding, or cost efficiency rather than timing premiums. The rise of year-round global supply from counter-seasonal production has extended the "mid-season" concept across the calendar, with Chilean and Peruvian grapes providing Northern Hemisphere mid-winter supply that mirrors summer mid-season production patterns. Mid-season varieties often serve as parents in breeding programs targeting shifted maturity timing.
Hauptsaison-Tafeltraube
main season grape
mid-season table grape
mittel reifende Traube
mittelfrühe Tafeltraube
peak season grape
Creasy GL, Creasy LL (2009) Grapes, CABI California Table Grape Commission (2020) Grape Variety Harvest Dates Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press Christensen LP (2000) Raisin Production Manual, UC ANR
Mid Season Table Grape Role
Midday water potential monitoring of Tempranillo in Ribera del Duero at 14:00 reveals progressive decline from -0.95 MPa in early July to -1.65 MPa in mid-August as soil water depletes, with stomatal conductance declining from 185 to 62 mmol/m²/s over the same period, triggering deficit irrigation when midday values exceed the -1.5 MPa threshold established for this variety to maintain minimum photosynthesis of 8 μmol CO₂/m²/s required for complete fruit ripening.
Midday water potential quality is a water potential quality that represents the tissue water potential measured during the period of maximum transpirational demand, typically between solar noon and early afternoon, characterizing the most stressed condition experienced by the vine during the diurnal cycle when atmospheric evaporative demand creates maximum tension in the xylem water column. Within the BFO framework, this quality specifically captures the dynamic water status under peak stress conditions, distinguishing this measurement from equilibrium predawn values and integrating the combined effects of soil water supply, plant hydraulic conductance, and atmospheric demand. The ontological significance lies in midday water potential reflecting the actual stress experienced by photosynthesizing leaves, where the tension required to maintain water flow from soil to atmosphere determines stomatal behavior, photosynthetic rate, and the risk of hydraulic dysfunction through cavitation. Midday water potential in unstressed grapevines typically ranges from -0.6 to -1.0 MPa, increasing (becoming more negative) to -1.2 to -1.6 MPa under moderate deficit and exceeding -2.0 MPa under severe stress, with cultivar-specific thresholds for stomatal closure and hydraulic damage. The difference between midday and predawn water potential (ΔΨ) provides information about hydraulic conductance and atmospheric demand, with larger differentials indicating higher transpiration rates or more restrictive hydraulic pathways. Measurement timing should target the period of maximum stress, typically 12:00-15:00 solar time, with consistent timing essential for comparing measurements across dates or sites. The relationship between midday water potential and gas exchange parameters enables physiological interpretation, with stomatal conductance declining progressively as midday water potential becomes more negative until complete closure at cultivar-specific thresholds. Understanding midday water potential quality enables real-time assessment of vine stress during the critical photosynthetic period, prediction of stomatal and photosynthetic responses, and optimization of irrigation timing to prevent excessive stress.
Midday_Water_Potential_Quality captures peak daily stress, complementing Predawn_Water_Potential_Quality for baseline soil water assessment. This quality integrates supply (soil water), demand (atmospheric conditions), and plant hydraulic properties into a single measurement reflecting actual tissue stress. Consistent measurement timing is critical as water potential varies substantially through the afternoon comparison across studies requires standardization to solar time or specification of measurement protocol.
Mittagsstress-Wasserpotential
Mittagswasserpotential
Wasserpotential bei maximaler Belastung
midday leaf water potential
minimales Tageswasserpotential
minimum daily water potential
peak stress water potential
Ψmd
Santesteban LG et al. (2011) Are midday leaf water potential measurements useful for precision viticulture? Acta Horticulturae 931: 437-444 Shellie KC (2006) Vine and berry response of Merlot (Vitis vinifera L.) to differential water stress. American Journal of Enology and Viticulture 57(4): 514-518
Midday Water Potential Quality
Cabernet Sauvignon middle pericarp at maturity comprises 88-91% of pericarp volume (86-89% of berry fresh weight excluding seeds), consisting of large parenchyma cells averaging 135-165 μm diameter in central region, 95-125 μm in outer middle pericarp, 105-140 μm in inner middle pericarp, with thin cell walls 0.5-0.7 μm, vacuoles 92-94% of cell volume containing 24-26% sugars (glucose 12.5-13.5%, fructose 11.5-12.5%), 0.6-0.75% organic acids (tartaric 0.38-0.48%, malic 0.18-0.28%), creating soft sweet flesh providing majority of fermentable sugars and juice volume for wine production. Chardonnay white wine cultivar middle pericarp showing white wine characteristics= 89-92% of pericarp volume, cells 140-175 μm diameter in central region, 21-24% sugars (glucose 10.5-12%, fructose 10.5-12%), 0.7-0.95% organic acids (tartaric 0.42-0.58%, malic 0.28-0.38%), maintaining higher acidity than typical red cultivar middle pericarp important for white wine freshness and balance. Syrah middle pericarp cell expansion during ripening showing growth dynamics= veraison (BBCH 81, week 10) cells 72-88 μm diameter mid-ripening (week 14) cells 115-145 μm maturity (week 18) cells 145-175 μm, representing 101-133% diameter increase (310-570% volume increase per cell), with middle pericarp cells showing greatest expansion of any pericarp region driving berry enlargement during Phase II ripening. Merlot middle pericarp phenolic content showing low tannin characteristic= total tannins 0.4-0.8 mg/g dry weight (versus 8-14 mg/g outer pericarp, 3-7 mg/g inner pericarp), anthocyanins <0.05 mg/g (versus 2.5-4.5 mg/g outer pericarp hypodermis), flavonols <0.1 mg/g (versus 0.8-2 mg/g outer pericarp), demonstrating middle pericarp as low-phenolic tissue providing sugars and juice volume but minimal contribution to wine color or tannin structure. Pinot Noir middle pericarp radial cell size gradient measured by microscopy= outer middle pericarp (adjacent to outer pericarp skin) cells 85-115 μm diameter central middle pericarp (maximum size zone) cells 125-165 μm inner middle pericarp (adjacent to inner pericarp) cells 95-135 μm, demonstrating systematic size variation with largest cells in central region and smaller cells toward pericarp boundaries, with cell size gradient reflecting position-dependent expansion during ripening possibly related to vascular proximity or mechanical constraints.
A middle pericarp is the bulk of the pericarp tissue comprising the mesocarp parenchyma between outer pericarp (skin) and inner pericarp (seed-adjacent layers), typically representing 85-92% of total pericarp volume and 88-94% of berry fresh weight, consisting of large thin-walled parenchyma cells that accumulate sugars, water, and acids creating the fleshy berry texture and providing the majority of fermentable sugars and juice volume for wine production, while containing minimal phenolic compounds compared to outer and inner pericarp regions. Middle pericarp cellular structure comprises= (1) large parenchyma cells typically 80-180 μm diameter at maturity (versus 15-70 μm in outer pericarp and 40-90 μm in inner pericarp), with cell size showing radial gradient increasing from outer middle pericarp (80-130 μm diameter) through central middle pericarp (110-180 μm, maximum cell size) to inner middle pericarp (90-140 μm) (2) thin flexible cell walls typically 0.4-0.8 μm thick (versus 1.5-3 μm in outer pericarp, 0.8-1.5 μm in inner pericarp) composed primarily of cellulose, hemicellulose, and pectin, creating soft texture when cells are turgid and contributing to berry softening during ripening through cell wall modifications (3) large central vacuoles occupying 90-95% of cell volume at maturity (versus 40-60% in epidermal cells), serving as storage compartments for sugars, organic acids, water, and minerals (4) thin peripheral cytoplasm layer (1-3 μm thickness) containing nucleus, organelles, and limited metabolic machinery focused on transport and storage rather than biosynthesis (5) intercellular spaces (air spaces between cells) comprising 1-5% of middle pericarp volume, providing gas diffusion pathways supporting respiration. Middle pericarp compositional characteristics= (1) high sugar content typically 18-28% at maturity (glucose and fructose in approximately 1=1 ratio), with sugars accumulated in vacuoles through phloem import and vacuolar transport creating osmotic potential driving water accumulation and cell expansion (2) moderate organic acid content typically 0.5-0.9% fresh weight at maturity (tartaric acid 0.3-0.6%, malic acid 0.1-0.4%), with acids declining during ripening from veraison peak through malic acid respiration (3) high water content typically 75-82% at maturity providing juice volume and creating turgid fleshy texture (4) low phenolic content with minimal tannins (0.3-1 mg/g dry weight versus 6-15 mg/g in outer pericarp), minimal anthocyanins (essentially zero versus 2-6 mg/g in outer pericarp hypodermal cells), and minimal flavonols (5) mineral content including potassium (1200-2200 mg/kg), calcium (60-150 mg/kg), magnesium (60-120 mg/kg) important for wine chemistry and yeast nutrition. Middle pericarp development follows berry growth pattern= Phase I (BBCH 71-75, weeks 0-6) involving cell division creating middle pericarp cell population (1.2-2.8 million cells representing 75-85% of total pericarp cell number), with cells 30-70 μm diameter lag phase (BBCH 77, weeks 6-8) with minimal middle pericarp growth Phase II (BBCH 81-89, weeks 9-18+) involving dramatic cell expansion particularly in middle pericarp where cells expand from 60-90 μm to 110-180 μm diameter (120-200% diameter increase, 350-700% volume increase per cell), creating bulk of berry expansion and juice accumulation. Middle pericarp vascular tissue integration= lateral vascular branches from major dorsal bundles penetrate middle pericarp distributing throughout tissue, with phloem sieve elements providing sugar and water import during ripening, xylem vessels functional during early development but largely non-functional during ripening as berry shifts from xylem-dominant to phloem-dominant import.
Middle_Pericarp represents the flesh bulk and is distinguished from Outer_Pericarp (skin with high phenolics) and Inner_Pericarp (seed-adjacent with elevated phenolics) by anatomical position, large cell size, thin walls, high sugar/water content, and low phenolic content. Middle pericarp provides the majority of juice volume and fermentable sugars but minimal wine color or tannins. Sub-classification into Intermediate and Transitional middle pericarp may reflect radial gradients or zones within the bulk tissue. Understanding middle pericarp development and composition is essential for predicting juice yield, sugar levels, and acid balance in winemaking.
mesocarp
mittleres Perikarp
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 180-190 Coombe, B.G., McCarthy, M.G. (2000). Dynamics of grape berry growth. Australian Journal of Grape and Wine Research 6=131-135 Nunan, K.J. et al. (2001). Changes in cell wall composition during ripening. Plant Physiology 126=520-528.
Middle Pericarp
Cabernet Franc in Loire Valley shows midshoot lignification from early August through mid-September at BBCH 38-41 over 5-6 weeks, with middle internodes 7-11 achieving lignin content of 24-28% dry weight and starch accumulation of 28-32% dry weight representing maximum values along shoot axis under temperate maritime conditions of 19-25°C temperature range declining to 15-20°C by maturation completion, moderate precipitation, and declining nitrogen availability typical of this vigorous mid-season cultivar achieving thorough midshoot maturation supporting strong subsequent season development
Midshoot lignification represents the maturation phase in middle shoot regions where internodes formed during peak shoot growth achieve complete woody structure development, typically representing zones of maximum lignification extent and reserve accumulation reflecting optimal conditions during formation and maturation periods. This important developmental stage occurs in middle shoot internodes typically positions 6-12 during late summer corresponding to mid-late berry development at BBCH 38-41, with maturation in these regions often achieving most complete lignification and highest reserve concentrations due to favorable timing of development. Midshoot internodes often represent longest internodes along shoot reflecting formation during peak growth conditions, requiring proportionally extended maturation period to achieve complete lignification throughout large internode volume, typically 3-4 weeks from lignification initiation to substantial completion. Lignification in midshoot regions achieves highest completion levels with lignin content reaching 20-30% dry weight in fully mature middle internodes compared to 15-20% in basal regions and potentially lower concentrations in apical regions depending on maturation timing and environmental conditions. Secondary wall deposition shows maximum extent in midshoot regions with vessel elements, fibers, and parenchyma cells all achieving thick secondary walls with extensive lignin incorporation, creating most rigid woody structure along shoot axis providing primary mechanical support and resistance to loading. Mechanical properties in fully lignified midshoot regions achieve maximum values with tensile strength 60-100 MPa exceeding basal and apical regions, modulus of elasticity indicating maximum rigidity, and resistance to compression and torsion providing robust mechanical performance. Vascular tissue development reaches maximum extent in midshoot regions with secondary xylem production creating numerous large-diameter vessels providing high water transport capacity, and extensive secondary phloem formation supporting efficient bidirectional assimilate movement, with vascular development reflecting importance of middle regions for hydraulic and translocation functions. Periderm development achieves complete maturation in midshoot regions with well-developed cork layer providing robust protection, brown bark appearance indicating complete epidermis replacement, and potentially lenticels for gas exchange through otherwise impermeable cork layer. Reserve accumulation reaches maximum levels in midshoot regions with starch concentration increasing to 25-35% dry weight in fully mature middle internodes by late summer-autumn, representing highest storage levels along shoot axis and reflecting optimal timing of carbohydrate accumulation when photosynthetic production is high and competing growth demands have ceased. Storage lipids also accumulate in parenchyma cells of midshoot regions with oil bodies forming in ray parenchyma cells adjacent to vascular tissues, providing additional energy-dense reserves complementing starch storage. Protein storage shows accumulation of storage proteins in parenchyma cells with concentrations reaching 8-12% dry weight in mature woody tissue, providing nitrogen reserves for subsequent season protein synthesis demands during bud burst and early growth. Water content in fully lignified midshoot regions declines to 50-55% fresh weight representing lowest levels along shoot axis, reflecting maximum cell wall material deposition, minimal metabolic water requirements, and advanced dehydration in preparation for dormancy. Metabolic activity in mature midshoot regions declines to near-maintenance levels with respiration 2-3 fold above completely dormant tissue, minimal protein synthesis focused on maintenance and defense functions, and virtually no growth-related metabolism. Cell viability maintains high levels in parenchyma cells of midshoot regions providing living tissue for storage and mobilization functions, while lignified vascular elements are dead at functional maturity serving purely structural and transport roles. Bark characteristics in midshoot regions often show distinctive features including color patterns, lenticel distribution, and texture providing cultivar identification characters, with middle shoot bark most representative of variety-specific characteristics. Mechanical importance of midshoot regions for shoot architecture includes providing primary bending resistance preventing shoot sagging, torsional resistance preventing twisting under wind loading, and load distribution transferring forces from distal shoot portions and fruit load to proximal attachment points. Reserve importance of midshoot regions for subsequent season development includes providing substantial portion of total shoot reserve carbohydrate available for bud burst and early growth, with middle regions often contributing 40-60% of mobilizable starch due to high concentrations and substantial tissue volume. Hormonal status in mature midshoot regions shows minimal auxin and gibberellin reflecting complete growth cessation, elevated abscisic acid maintaining maturation and dormancy preparation, and possible accumulation of defense-related hormones including jasmonates and salicylates supporting pathogen resistance. Temperature during midshoot lignification typically represents late summer conditions with warm days promoting rapid maturation and cool nights enhancing carbohydrate accumulation, with optimal conditions supporting maximum lignification extent and reserve storage. Light intensity affects midshoot lignification through canopy position effects with well-exposed shoots showing complete thorough maturation while shaded shoots may show reduced lignification and reserve accumulation affecting subsequent productivity. Water availability shows moderate late-summer stress typically beneficial for promoting complete maturation and enhanced reserve accumulation through stress-induced resource partitioning, while severe stress may limit photosynthetic production reducing storage potential. Nutrient availability shows declining nitrogen levels appropriate for maturation with remobilization from senescing leaves providing nitrogen for storage protein synthesis, while excessive nitrogen would impair maturation and reduce reserve accumulation. The process typically requires 3-5 weeks for individual midshoot internodes to achieve complete maturation from lignification initiation to maximum reserve accumulation depending on environmental conditions and cultivar characteristics. Cultivar variation appears in maximum lignification extent achieved in midshoot regions with some varieties showing complete thorough lignification to 28-32% lignin content while others maintain lower levels, maximum reserve concentrations ranging from moderate 20-25% starch to high 30-35% in elite storage cultivars, and timing of maturation completion affecting subsequent dormancy depth and cold hardiness. Regional timing shows midshoot lignification occurring late July-August in warm climates, August-September in temperate regions, and September in cool areas, with timing critical for achieving complete maturation before dormancy entry. Viticultural management considerations include recognizing midshoot regions as most important for reserve storage and mechanical support, monitoring maturation completeness in middle zones as indicator of overall vine preparation for dormancy, ensuring management promotes thorough midshoot lignification through appropriate late-season water and nitrogen management, assessing midshoot reserve status through tissue testing informing subsequent season vigor predictions, and understanding that inadequate midshoot maturation severely compromises subsequent productivity and survival. Research applications include analyzing regulation of maximum lignification and reserve accumulation including hormonal and metabolic controls, quantifying relationships between maturation extent and subsequent season development capacity, evaluating management effects on midshoot maturation affecting productivity and quality, assessing mechanical properties of mature woody tissue supporting architectural understanding, and documenting reserve dynamics during accumulation and mobilization cycles in perennial plant resource management.
Midshoot_Lignification is third of four Shoot_Lignification subphases, distinguished by maturation in middle regions achieving maximum lignification and reserve accumulation. Follows Progressive_Shoot_Lignification and may overlap with Complete_Shoot_Lignification in apical regions. Represents most critical maturation phase for reserve storage and mechanical support. Extent and completeness strongly predict subsequent season vigor and productivity.
Haupttriebverholzung
Mitteltriebausreifung
central region woodification
middle shoot lignification
mittlere Triebverholzung
peak shoot maturation
zentrale Holzbildung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 298-305 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 230-235 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Midshoot Lignification
Systematic quantification of Midwinter_Hardiness_Disposition at South Dakota State University employed differential thermal analysis (DTA) to determine LT50 values for buds, canes, and trunks across 35 cultivars at maximum hardiness in mid-January. Primary bud LT50 ranged from -15.2°C ('Grenache', least hardy V. vinifera) to -38.4°C ('Valiant', V. riparia hybrid), representing 23°C variation in Midwinter_Hardiness_Disposition across cultivated grapevine diversity. Cane LT50 values were consistently 3-5°C colder than corresponding bud values within cultivars, while trunk tissues showed variable relationships depending on wood age and acclimation status. Among V. vinifera cultivars, 'Riesling' (-24.8°C) and 'Cabernet Franc' (-23.5°C) showed superior Midwinter_Hardiness_Disposition compared to 'Merlot' (-19.2°C) and 'Syrah' (-17.8°C). The PIWI cultivar 'Solaris' achieved -27.3°C, intermediate between V. vinifera and cold-hardy hybrids. Strong correlation was observed between bud and cane hardiness within cultivars (r = 0.94), suggesting common genetic control, while correlation with trunk hardiness was lower (r = 0.72), indicating partially independent regulation. These LT50 values directly inform cultivar recommendations, with -25°C considered the minimum threshold for reliable production in upper Midwestern U.S. climates where ambient temperatures routinely reach -30°C.
A Midwinter_Hardiness_Disposition is a Cold_Tolerance_Disposition that inheres in a Vitis plant during the period of maximum cold acclimation (typically December through February in the Northern Hemisphere) and is realized through the capacity to survive the most extreme low temperatures encountered during deep dormancy when tissues have achieved full cold hardening. This disposition represents the peak cold tolerance capacity of grapevine, reflecting the maximum protection achieved through complete cold acclimation processes including dormancy establishment, cryoprotectant accumulation, membrane stabilization, and water redistribution. Midwinter hardiness is typically quantified as the LT50 value—the temperature at which 50% of a given tissue type is killed—determined through controlled freezing tests or differential thermal analysis detecting ice nucleation events in supercooled tissues. The midwinter hardiness of different vine tissues varies substantially, with canes and trunks generally more hardy than buds, and among bud components, primary buds less hardy than secondary and tertiary buds. V. vinifera cultivars achieve midwinter bud LT50 values typically ranging from -15°C (tender cultivars like 'Grenache') to -25°C (hardier cultivars like 'Riesling'), while V. riparia and V. amurensis can reach -40°C, and cold-hardy hybrids bred from these species achieve -30°C to -38°C. Midwinter_Hardiness_Disposition is the primary determinant of cultivar survival in continental climates with severe winters, though spring frost tolerance (post-budbreak) and deacclimation stability also contribute to overall winter survival success.
Midwinter_Hardiness_Disposition is_a Cold_Tolerance_Disposition. This class specifically addresses cold tolerance during deep dormancy when acclimation is complete, distinct from spring frost tolerance (growing tissue damage) or fall frost tolerance (pre-dormancy damage). The child class Bud_Cold_Hardiness_Disposition addresses the economically critical bud tissues determining next season's crop. LT50 values are the standard metric, measured by controlled freezing or differential thermal analysis. Midwinter hardiness represents the genetic potential for cold tolerance, while actual winter survival depends also on environmental factors including minimum temperature duration, wind exposure, and snow cover.
Mittwinter-Härte-Disposition
Tiefwinter-Toleranz
deep dormancy freeze tolerance
Mills LJ et al. (2006) Foliar temperature and VvCBF gene expression during the early stages of cold acclimation in Vitis, Functional Plant Biology 33:593-605 Wolf TK, Cook MK (1994) Cold hardiness of dormant buds of grape cultivars: comparison of thermal analysis and field survival, HortScience 29:1453-1455 Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235
Midwinter Hardiness Disposition
Migration path for Syrah traces a complex multi-stage dispersal from a single-origin event in southeastern France: genetic parentage identifies Dureza (Ardèche) × Mondeuse Blanche (Savoie) as the parents, placing the origin in the northern Rhône corridor, with subsequent local diffusion throughout the Rhône Valley (documented by 1781 in Hermitage), southward extension to Languedoc and Provence (19th century), introduction to Australia (1832, Busby collection to Sydney, then to the Barossa Valley by 1843 under the name "Hermitage," becoming "Shiraz" by the mid-20th century), to South Africa (1860s, as "Shiraz" from French sources, later supplemented by Australian clonal material), to California (1878, via French immigrant plantings in Sonoma), and most recently to emerging regions including Hawke's Bay New Zealand (1990s) and Swartland South Africa (2000s reconceptualization as premium variety) — the migration path revealing how colonial networks, individual collectors, and scientific institutions each served as dispersal mechanisms at different historical stages. Migration path for Blaufränkisch reconstructs a predominantly Central European trajectory — probable origin in the Austrian-Hungarian border region (pre-19th century), diffusion through the Habsburg administrative territory to Moravia (as Frankovka), Serbia (as Frankovka crna), and Croatia, westward introduction to Württemberg (as Lemberger, possibly via diplomatic exchange), and transatlantic transfer to Washington State (1970s, as Lemberger via German-American viticultural connections).
Migration path is a spread history entity that documents the reconstructed geographic route and intermediate staging points through which a grape cultivar traveled from its center of origin to its current cultivation regions, integrating evidence from chronologically ordered introduction records, genetic fingerprinting of regional clonal populations (identifying bottleneck signatures and founder effects that indicate dispersal direction), archival documentation of vine trade and exchange networks, and linguistic analysis of cultivar synonym distributions, expressed as a sequence of geographic waypoints with associated dates and dispersal mechanisms, and providing the spatial-temporal trajectory of cultivar movement that reveals the human networks — trade routes, colonial administrations, monastic connections, scientific institutions — that served as conduits for viticultural knowledge and plant material transfer.
Migration path reconstruction represents one of the most methodologically sophisticated activities in viticultural history, requiring the integration of evidence from multiple disciplines — molecular genetics, archival history, linguistics, and archaeology — each providing complementary but sometimes contradictory spatial-temporal information. The molecular genetic approach exploits the fact that clonally propagated cultivars accumulate somatic mutations (SNPs, indels, transposon insertions) over centuries of vegetative propagation, creating regional clonal lineages that can be phylogenetically analyzed to infer dispersal direction and timing — populations with greater clonal diversity are inferred to be older, and shared rare mutations identify dispersal connections between regions. The founder effect signature — where a newly introduced population shows reduced genetic diversity relative to its source — provides molecular evidence for the direction of migration, though this signal is complicated by multiple introductions from different sources and by deliberate clonal selection programs that artificially reduce diversity. The linguistic analysis of synonym distributions provides an independent migration proxy — the pattern of cultivar names across regions encodes dispersal information because name changes typically occur at linguistic or political boundaries, and the sequence of name transformations can indicate the direction and approximate timing of spread. The distinction between primary migration (the initial introduction of a cultivar to a new major viticultural region) and secondary migration (the subsequent local diffusion within that region or re-introduction from the secondary to further destinations) adds hierarchical structure to migration path documentation. The colonial dimension of cultivar migration is particularly well-documented for New World viticulture — the Spanish Mission grape (Listán Prieto/País/Criolla Chica) traveled from Spain to the Canary Islands to Mexico to Peru to Chile and Argentina over approximately 150 years (1490s-1640s), creating a traceable migration path documented in colonial administrative records that represents one of the most completely documented cultivar dispersals in history.
Ausbreitungspfad
Migrationsweg
Verbreitungstrajektorie
Verbreitungsweg
Wanderungsroute
cultivar migration route
diffusion pathway
dispersal route
dissemination path
spread trajectory
Myles S et al. (2011) Genetic structure and domestication history of the grape. Proceedings of the National Academy of Sciences 108(9): 3530-3535 Boursiquot JM et al. (2009) Parentage of Merlot and related winegrape cultivars of southwestern France: discovery of the missing link. Australian Journal of Grape and Wine Research 15(2): 144-155 Unwin T (1991) Wine and the Vine, Routledge, pp.: 130-300 Lacoste P et al. (2010) The introduction of Vitis vinifera to the Americas: historical perspectives on the Mission grape. Vitis 49(2): 73-78
Migration Path
Mild deficiency quality diagnosis for potassium in Walla Walla Valley Merlot enabled timely corrective intervention before deficiency progressed to yield-limiting severity. Visual assessment revealed mild deficiency quality with subtle marginal bronzing on basal leaves (score 1.5 on 0-5 scale) visible primarily when examining vines against K-sufficient reference blocks, with affected tissue showing tan discoloration extending only 3-5 mm from leaf margins rather than the extensive necrosis of more severe deficiency. Symptom distribution showed mild deficiency quality limited to the oldest 3-4 leaves at the vine base with mid-canopy and apical leaves appearing normal, consistent with early-stage limitation where only leaves with longest exposure showed symptoms. Tissue analysis confirmed mild deficiency quality with bloom petiole K at 1.4% (low-adequate range, optimal greater than 2.0%, deficient below 1.2%), indicating potassium status in the transition zone where limitation had begun but critical deficiency had not yet developed. Functional assessment revealed mild deficiency quality with berry K concentration 15% below K-sufficient reference vines at véraison, suggesting modest impact on fruit quality parameters but not the dramatic effects of severe deficiency. The mild deficiency quality diagnosis guided management response with standard potassium sulfate application at 150 kg K₂O/ha through the fertigation system, a moderate rate appropriate for mild limitation rather than the emergency high rates required for severe deficiency. Follow-up assessment confirmed symptom resolution by the following season with petiole K increased to 2.1% and no visible marginal symptoms, demonstrating the full recoverability characteristic of mild deficiency when corrected promptly.
Mild deficiency quality is a deficiency severity quality that inheres in a grapevine organism and represents a nutritional limitation of minor intensity where tissue nutrient concentrations have declined below optimal but remain above severely limiting levels, with visible symptoms subtle and often apparent only when comparing to nutrient-sufficient reference vines, and with functional impacts including slight growth reduction and early metabolic stress responses detectable through sensitive measurements but not yet causing substantial productivity losses. This quality captures the early stage of overt deficiency, recognizing that mild deficiency represents the transition from latent (subclinical) to visually apparent limitation, where symptoms have begun to manifest but have not yet reached the intensity requiring urgent intervention. The ontological significance of mild deficiency quality lies in its indication that corrective action is warranted but can proceed through standard management practices without emergency measures, and in its representation of a recoverable condition where timely correction can fully restore normal function without lasting productivity impact. The biological basis of mild deficiency encompasses tissue concentrations in the lower adequate to upper deficient range where metabolic processes remain largely functional but operate with reduced efficiency, stress-responsive gene expression activated at low levels, and initial visible symptoms reflecting the earliest detectable manifestations of nutrient limitation. The visual symptoms of mild deficiency typically show reduced intensity, limited distribution, or early-stage development compared to more severe deficiency, with characteristic features like slight pallor rather than intense chlorosis, marginal rather than extensive necrosis, or symptoms limited to oldest leaves rather than distributed throughout the canopy. The management response to mild deficiency typically involves standard corrective fertilization at recommended rates through normal application methods, with expectation of symptom resolution within one growing season and no lasting impact on vine productivity or health.
Mild_Deficiency_Quality is_a Deficiency_Severity_Quality, representing early-stage overt deficiency with subtle symptoms and limited functional impact. This class addresses the lowest severity category with visible symptoms. Symptoms visible but subtle, often requiring comparison to reference. Limited distribution (oldest leaves only) typical. Tissue concentrations in low-adequate to upper-deficient range. Standard corrective fertilization at normal rates sufficient. Full recovery expected within one season without lasting impact. Represents transition from latent to apparent deficiency.
Frühstadium-Mangel
Leichter Mangel
early-stage deficiency
geringe Nährstofflimitation
minor nutritional limitation
slight deficiency
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Bergmann W (1992) Nutritional Disorders of Plants. Gustav Fischer Verlag, Stuttgart Robinson JB (1992) Grapevine nutrition. In: Coombe BG, Dry PR (eds) Viticulture, Volume 2. Winetitles, Adelaide, pp. 178-208
Mild Deficiency Quality
Mild water stress quality management in Hawke's Bay Merlot employed regulated deficit irrigation targeting the beneficial mild stress zone from fruit set through véraison to optimize vine balance while preserving yield potential. Target mild water stress quality was maintained at predawn leaf water potential of -0.3 MPa through soil moisture monitoring and responsive irrigation scheduling, achieving the intended physiological state characterized by stomatal conductance reduced to 75% of fully-watered controls, continued but moderated shoot growth at 2.1 cm per day versus 3.4 cm in irrigated controls, and photosynthesis maintained at 88% of potential. Fruit outcomes demonstrated the quality benefits of mild water stress quality management with berry weight reduced only 8% versus controls indicating concentration without severe stress impact, sugar accumulation proceeding on schedule to target Brix, and phenolic concentration enhanced by 15% attributed to improved fruit exposure from reduced canopy density. Yield reduction of 7% proved economically acceptable given the quality premium achieved, validating mild water stress quality as a practical management target for this cultivar-site combination.
Mild water stress quality is a water deficit severity quality that inheres in a grapevine organism and represents the initial category of water deficit characterized by incipient stomatal closure reducing transpiration while maintaining near-normal photosynthetic carbon assimilation, modest growth rate reduction without complete cessation, and subtle physiological adjustments that may enhance fruit quality through improved vine balance without the pronounced concentration effects or stress-induced metabolic changes associated with more severe deficit categories. This quality defines the threshold region where water limitation begins to influence vine physiology but remains within the compensatory capacity of normal regulatory mechanisms, creating conditions that often prove beneficial for wine grape quality through gentle restraint of vegetative vigor rather than the more dramatic effects of pronounced water stress. The ontological significance of mild water stress quality lies in its identification of the beneficial entry-level stress zone where quality enhancement begins without significant yield penalty or stress damage risk, representing the lower bound of the regulated deficit irrigation target range for production systems balancing yield maintenance with quality improvement. The biological basis of mild water stress quality encompasses the initial stomatal response to declining soil water availability wherein guard cells respond to root-sourced abscisic acid signals and local hydraulic cues by reducing stomatal aperture, decreasing transpirational water loss while sufficient stomatal conductance remains to support continued CO₂ uptake for photosynthesis at 80-90% of unstressed rates. Shoot growth proves more sensitive to water deficit than photosynthesis, showing measurable reduction under mild stress even while carbon assimilation continues near normal levels, creating the improved source-sink balance that benefits fruit development through reduced competition from vegetative sinks. The physiological indicators of mild water stress quality include predawn leaf water potential in the range of -0.2 to -0.4 MPa and midday stem water potential of -0.8 to -1.0 MPa for most Vitis vinifera cultivars, though specific thresholds vary with cultivar isohydric/anisohydric behavior, rootstock characteristics, and regional calibration. Visual symptoms are absent or limited to subtle changes in leaf color, tendril turgor, and growth rate that require trained observation to detect, distinguishing mild stress from the visible symptoms characterizing more severe categories.
Mild_Water_Stress_Quality is_a Water_Deficit_Severity_Quality, representing the lowest stress intensity category with incipient physiological response. This class addresses the threshold stress zone where quality enhancement begins without significant yield or health penalty. Typical predawn LWP -0.2 to -0.4 MPa, midday SWP -0.8 to -1.0 MPa provide quantitative thresholds though cultivar calibration is required. Distinguished from Moderate_Water_Stress_Quality by the maintenance of near-normal photosynthesis and absence of visible symptoms. The isohydric/anisohydric cultivar distinction influences threshold values. Critical for precision irrigation targeting the beneficial stress onset zone.
Schwellenwasserstress
beginnender Trockenstress
incipient water stress
initial drought stress
leichtes Wasserdefizit
light water deficit
milder Wasserstress
threshold water stress
Williams LE, Araujo FJ (2002) Correlations among predawn leaf, midday leaf, and midday stem water potential and their correlations with other measures of soil and plant water status in Vitis vinifera. Journal of the American Society for Horticultural Science 127:448-454 Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell and Environment 26:1393-1405 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data. Annals of Botany 105:661-676
Mild Water Stress Quality
Merlot in Bordeaux shows millerandage formation during early July Year N+1 at BBCH 69-75, with 12-28% berries developing as shot berries 3-5 mm diameter remaining seedless or poorly seeded, normal berries achieving 10-13 mm creating obvious size heterogeneity, shot berries persisting through harvest contributing high acidity, under oceanic conditions with variable spring weather, typical of this millerandage-susceptible cultivar requiring favorable fruit set conditions
Millerandage formation represents the shot berry phenomenon when some berries within cluster fail to develop normally during fruit set, remaining small seedless or poorly seeded arrested berries typically 3-6 mm diameter, creating within-cluster size heterogeneity with mixture of normal and shot berries, reducing yield and potentially affecting wine quality through variable berry composition. This developmental heterogeneity phenomenon occurs during fruit set in late June-July in temperate regions Year N+1, with arrest typically during first 2-4 weeks post-flowering, corresponding to BBCH 69-75 fruit set to young fruit stages. Millerandage mechanism involves partial fertilization failure or early developmental arrest creating shot berries, seedless or single small seed development providing inadequate auxin for normal growth, arrested berry expansion remaining at 3-6 mm diameter versus 8-18 mm normal berries, persistent shot berries remaining attached through ripening creating size heterogeneity, and variable berry development within cluster from differential fertilization or development success. Shot berry characteristics show small size typically 3-6 mm diameter representing 20-40% normal berry size, seedless condition or single small poorly developed seed, hard firm texture from limited cell expansion, green color persisting longer than normal berries, high acidity and low sugar at harvest distinguishing from normal berries, and arrested development preventing normal berry maturation. Seedless shot berry development shows fertilization failure or very early seed abortion, berry initiation from residual hormone stimulus, limited growth without adequate seed auxin production, developmental arrest at small size, and persistence through harvest as hard green berries. Partially seeded shot berry development shows single small seed formation from partial fertilization success, limited auxin production insufficient for normal berry expansion, intermediate size between seedless and normal berries sometimes observed, and variable development reflecting seed development extent. Developmental arrest timing shows arrest during critical early growth weeks 1-4 post-fertilization, earlier arrest producing smaller more arrested berries, and timing determining final shot berry characteristics. Cluster heterogeneity from millerandage shows mixture of normal-sized berries 8-18 mm with shot berries 3-6 mm, size ratio normal shot typically 2 per 1 to 4 per 1 creating obvious heterogeneity, variable proportions with mild millerandage showing 5-15% shot berries and severe cases 30-50%, and within-cluster variation affecting appearance and quality. Millerandage causes include incomplete or variable fertilization success across flowers, environmental stress during flowering or early fruit set affecting fertilization or development, boron deficiency preventing normal pollen tube growth in subset of flowers, genetic susceptibility in certain cultivars prone to millerandage, cool flowering temperatures potentially affecting fertilization uniformity, and overall factors causing developmental heterogeneity. Fertilization variability as cause shows differential fertilization success across cluster flowers, some flowers achieving complete fertilization and normal development, others experiencing partial or complete failure creating shot berries, and fertilization heterogeneity translating to berry size heterogeneity. Environmental stress effects show stress during critical flowering or early fruit set periods affecting subset of flowers or berries, creating developmental heterogeneity within clusters, and stress severity determining millerandage extent. Cultivar susceptibility shows genetic differences with some cultivars prone to millerandage under marginal conditions, Grenache, Malbec, and Merlot showing notable susceptibility, resistant cultivars maintaining uniform berry development, and genetic factors affecting millerandage risk. Millerandage versus coulure distinction shows millerandage producing persistent shot berries versus coulure causing berry drop, both representing fruit set problems from similar causes, millerandage creating quality issues from heterogeneity while coulure reduces yield through loss, and overlapping occurrence possible with both phenomena in same season. Shot berry persistence shows shot berries remaining attached through ripening unlike early fruit drop, hard texture preventing normal ripening progression, harvest typically including shot berries mixed with normal berries, and persistence creating processing challenges. Harvest implications show shot berries harvested with normal berries creating heterogeneity, shot berries contributing high acidity and tannin from immature development, potential negative quality effects from inclusion though sometimes minor if proportion low, and sorting possibly used to remove shot berries in severe cases. Yield effects show reduced effective yield from shot berry limited size, shot berries contributing 20-40% normal berry weight, severe millerandage reducing yield 15-30% through size reduction, and yield impact less severe than coulure though quality concerns additional. Quality effects show shot berries contributing disproportionate acidity and herbaceous character if proportion high, potential positive effects from concentration if proportion low 5-10%, negative effects typically dominating when proportion >20%, and overall quality impact depending on millerandage severity. Millerandage assessment shows visual cluster examination documenting size heterogeneity, shot berry counting quantifying proportion, size measurements distinguishing shot from normal berries, and severity classification from percentage shot berries. Millerandage management shows boron fertilization preventing deficiency-induced millerandage, favorable flowering conditions reducing stress-induced heterogeneity, cluster thinning potentially removing heavily affected clusters, and berry sorting at harvest removing shot berries in severe cases. The process represents third type of fruit drop or more accurately arrested development creating persistent small berries. Duration represents developmental arrest during weeks 1-4 post-flowering. Regional timing shows millerandage formation during late June-July across climates during fruit set. Viticultural significance includes creating within-cluster heterogeneity affecting quality, reducing effective yield through limited shot berry size, demonstrating cultivar and environmental susceptibility, and requiring management to minimize occurrence and impacts. Research applications include analyzing millerandage formation mechanisms and causes, investigating fertilization and developmental heterogeneity, evaluating environmental and genetic factors affecting millerandage, quantifying quality impacts from shot berries, and developing prevention strategies.
Millerandage_Formation is third type of Fruit_Drop phenomenon, though berries persist rather than drop. Represents arrested development during fruit set creating shot berries 3-6 mm. Timing weeks 1-4 post-flowering. Results from fertilization failure or early developmental arrest. Seedless or poorly seeded. Persists through harvest unlike Early_Fruit_Drop. Creates size heterogeneity affecting quality. Incidence 5-50% depending on severity. Cultivar susceptibility particularly Grenache, Malbec, Merlot. Reduces effective yield and quality.
Beerengrößen-Heterogenität
Gescheine
Henne und Küken
Verrieselung
arrested berry development
berry size heterogeneity
hen and chickens
shot berries
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 450-455 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Millerandage Formation
Millerandage incidence assessment in Chardonnay comparing flowering condition effects reveals the fertilization heterogeneity affecting cluster composition, with optimal flowering conditions producing uniform clusters (95% normal berries averaging 1.4 g, 5% shot berries <0.3 g, minimal composition heterogeneity), moderate stress conditions producing affected clusters (72% normal berries, 28% shot berries creating visible "hen and chicken" appearance, composition spread of 4 Brix between berry populations at harvest), and severe stress producing heavily affected clusters (48% normal, 52% shot berries, pronounced heterogeneity complicating harvest timing), demonstrating how millerandage incidence determines the berry population uniformity affecting both yield quantity and fruit quality consistency.
Millerandage incidence quality is a yield potential quality that represents the occurrence of abnormal berry size variation within clusters resulting from differential fertilization success, characterizing the condition where clusters contain a mixture of normal seeded berries alongside small seedless berries (shot berries or "hen and chicken" appearance) that developed from stimulated but unfertilized or incompletely fertilized ovaries. Within the BFO framework, this quality specifically addresses the fertilization heterogeneity dimension of yield potential, distinguishing millerandage from normal berry size variation or from coulure where berries abscise entirely while recognizing that millerandage represents partial reproductive success—flowers developing into berries despite inadequate fertilization, producing small seedless berries that contribute little to yield while remaining attached to the cluster. The ontological significance lies in millerandage affecting both yield quantity and fruit composition uniformity, where affected clusters show reduced total weight (shot berries contributing minimal mass), increased composition heterogeneity (seedless berries ripening differently than seeded berries), and harvest complications (difficulty achieving uniform maturity across the mixed berry population), with these effects creating quality challenges beyond the simple yield reduction that berry counts might suggest. The causes of millerandage include incomplete pollination (some flowers receiving inadequate pollen due to weather, timing, or pollen quality), failed fertilization (pollen reaching stigma but failing to effect fertilization due to temperature extremes or incompatibility), and embryo abortion (fertilization occurring but embryo development failing, producing small parthenocarpic berries), with these causes often overlapping and difficult to distinguish in practice. The cultivar variation in millerandage susceptibility shows some varieties prone to the condition (Chardonnay, Gewürztraminer, Pinot varieties showing frequent millerandage) while others rarely exhibit it, with susceptibility related to flower morphology, pollen characteristics, and sensitivity to flowering-period stress. Understanding millerandage incidence quality enables assessment of cluster composition uniformity, diagnosis of flowering problems affecting fertilization, and management decisions regarding affected fruit (potential for enhanced concentration from shot berries versus problems from heterogeneous ripening).
Millerandage_Incidence_Quality addresses the occurrence of shot berries from incomplete fertilization creating cluster heterogeneity. This quality affects both yield quantity and composition uniformity. The class supports cluster quality assessment through millerandage characterization indicating fertilization success.
Beerengrößenheterogenität
Henne-und-Küken
Kleinbeerigkeit
Millerandage-Inzidenz
hen and chicken
incomplete set severity
shot berry incidence
Ebadi A et al. (1996) Effect of short-term temperature and shading on fruit-set, seed and berry development in model vines of Vitis vinifera cvs Chardonnay and Shiraz. Australian Journal of Grape and Wine Research 2(1): 1-7 May P (2004) Flowering and Fruitset in Grapevines. Lythrum Press, Adelaide, pp. 1-119
Millerandage Incidence Quality
Millerandage induction quality assessment for zinc deficiency in Napa Valley Cabernet Sauvignon documented the fruit set impact of zinc limitation in a vineyard where foliar symptoms were subtle but cluster quality was severely compromised. Post-fruit-set cluster evaluation revealed millerandage induction quality with affected clusters showing the characteristic trimodal berry size distribution, as 40% normal seeded berries (mean weight 1.2 g), 35% shot berries (mean weight 0.15 g, seedless), and 25% aborted ovary remnants (coulure), creating the "hens and chickens" appearance that dramatically reduced both yield and cluster aesthetic quality. Spatial analysis showed millerandage induction quality concentrated in vineyard areas with soil pH above 8.0 and historical zinc tissue analysis below threshold, while areas with adequate zinc showed normal fruit set with greater than 95% of berries in the normal size class. Temporal correlation established that millerandage induction quality severity related to zinc status during the bloom period, with vines receiving pre-bloom foliar zinc applications showing only 8% shot berries versus 35% in untreated controls, confirming the critical window for zinc nutrition in preventing millerandage. Pollen viability assessment revealed the mechanism of millerandage induction quality, with pollen from zinc-deficient vines showing only 45% germination and mean tube length of 120 μm versus 85% germination and 380 μm tube length from zinc-sufficient vines, establishing impaired pollen function as the proximate cause of failed fertilization. Yield impact assessment showed millerandage induction quality causing 42% yield reduction through the combination of berry loss from coulure and reduced berry weight from shot berries, demonstrating the substantial economic consequence of zinc deficiency on reproductive development.
Millerandage induction quality in zinc deficiency is a zinc deficiency symptom quality that inheres in grapevine clusters and represents the disruption of normal fruit set and berry development resulting in clusters containing a mixture of normal seeded berries, small seedless shot berries (millerandage), and aborted berries (coulure), with the uneven berry development reflecting zinc's essential roles in pollen development, pollen tube growth, and fertilization processes that, when impaired by zinc deficiency, cause incomplete or failed fertilization leading to the characteristic "hens and chickens" cluster appearance. This quality captures the reproductive consequence of zinc deficiency, recognizing that while foliar symptoms may be subtle in mild deficiency, the fruit set impact can cause substantial yield losses and, in table grapes, severe marketability problems due to irregular cluster appearance. The ontological significance of millerandage induction quality in zinc deficiency lies in its economic impact through yield reduction and quality compromise, and in its diagnostic value indicating zinc limitation even when foliar symptoms are not prominent, since the critical role of zinc in reproductive development makes fruit set highly sensitive to zinc status. The biological basis of zinc-deficiency millerandage encompasses zinc's requirement for pollen development where deficiency causes reduced pollen viability and germination capacity, for pollen tube growth where zinc is essential for tube elongation to reach the ovule, and for early embryo development where zinc deficiency may cause embryo abortion even after fertilization, collectively resulting in the mixed population of fertilized berries (normal development), parthenocarpic berries (unfertilized but developing as seedless shot berries), and aborted berries (coulure) that constitutes millerandage. The severity of millerandage correlates with the timing and intensity of zinc deficiency relative to the flowering period, with zinc stress during pollen development and flowering causing more severe effects than stress at other times, and with recovery impossible once berry development pattern is established regardless of subsequent zinc correction.
Millerandage_Induction_Quality_(Zn) is_a Zinc_Deficiency_Symptom_Quality, representing the fruit set failure and uneven berry development from Zn-limited pollen and fertilization function. This class addresses the reproductive/yield dimension of Zn deficiency. The "hens and chickens" cluster appearance (normal + shot berries) is characteristic. Bloom-period Zn status is critical once fruit set is established, correction cannot reverse millerandage. Pollen viability and tube growth both require adequate Zn. Table grape quality severely affected by cluster appearance wine grapes affected primarily through yield loss. Pre-bloom foliar Zn application prevents millerandage when deficiency risk is identified.
Hühner-und-Küken durch Zn-Mangel
Verrieselungs-Induktion bei Zn-Mangel
Zink-Durchrieseln
Zink-induzierte Millerandage
Zn-induced millerandage
hens and chickens from Zn
zinc fruit set failure
zinc shot berries
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication Pandey N et al. (2006) Zinc is critically required for pollen function and fertilisation in lentil. Journal of Trace Elements in Medicine and Biology 20:89-96
Millerandage Induction Quality (Zn)
Gewürztraminer exhibits severe millerandage susceptibility with shot berry proportions reaching 30-50% of berries per cluster in problematic years, while Cabernet Sauvignon typically shows only 5-15%. Shot berries in Chardonnay average 4-6 mm diameter and 0.3-0.5 g weight compared to 12-14 mm and 1.2-1.8 g for normally developed berries. Muscat cultivars show genetic predisposition to millerandage through high rates of ovule abortion even under favorable pollination conditions. Berry size coefficient of variation (CV) above 40% indicates severe millerandage compared to normal CV of 15-25%.
A fruit set disposition that is the inherited capacity of a Vitis vinifera plant to develop clusters containing berries of highly variable size (millerandage or hen-and-chicks), resulting from incomplete fertilization producing seedless small berries (shot berries) alongside normally fertilized larger berries within the same cluster.
Millerandage differs from coulure in that affected flowers are retained rather than abscised, but develop into small seedless or single-seeded berries. The condition results from partial fertilization (one ovule fertilized) or stimulative parthenocarpy without fertilization. Shot berries may have different sugar and acid composition than normal berries, potentially affecting wine quality. Some millerandage is considered desirable in certain wine styles due to higher skin-to-pulp ratio in small berries.
Millerandage-Anfälligkeit
Verrieselungsneigung mit Kleinbeeren
hen and chicks tendency
shot berry susceptibility
uneven berry size disposition
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 130-135 Friend AP, Trought MCT (2007) Delayed winter spur-pruning in New Zealand can alter yield components of Merlot grapevines. Australian Journal of Grape and Wine Research 13:157-164
Millerandage Susceptibility Disposition
Mineral aroma assessment in Mosel Riesling comparing Slate (Schiefer) and Limestone (Kalk) soils reveals that tasters consistently rate Slate-origin wines higher in perceived minerality (6.5 versus 4.8 on 10-point scale) despite analytical studies showing no difference in mineral element composition between wines, suggesting that minerality perception follows terroir reputation and label expectations rather than direct compositional differences, while both expressions show higher minerality ratings than equivalent-quality New Zealand Riesling (3.2) where terroir narrative emphasizes fruit rather than mineral character.
Mineral aroma quality is an earthy aroma quality that represents olfactory perception described using mineral and geological terminology including wet stone, chalk, slate, granite, and general minerality descriptors, characterizing the non-fruit, non-vegetal aromatic impression associated with perceived terroir expression and site-specific character in fine wines. Within the BFO framework, this quality specifically addresses the mineral subcategory of earthy aroma, distinguished by its explicit reference to geological materials and its central role in terroir-focused wine evaluation despite ongoing scientific debate about the chemical and perceptual basis for minerality. The ontological significance lies in minerality representing perhaps the most important yet scientifically problematic wine descriptor, where the term communicates quality, terroir expression, and site specificity valued by consumers and critics despite lack of consensus about its sensory definition, chemical correlates, or direct geological origin. The compositional uncertainty surrounding mineral perception reflects the complexity of this quality, with proposed contributors including volatile sulfur compounds (hydrogen sulfide, methanethiol at sub-threshold levels potentially perceived as mineral), certain thiols, succinic acid (creating stony impression), and reduction-derived compounds, while research has generally failed to establish direct connections between vineyard mineral content and wine mineral perception. The perceptual psychology research suggests minerality may be a synthetic perception integrating olfactory (possibly reduction-related), gustatory (acidity, saltiness), and tactile (mouthfeel, texture) inputs rather than a pure olfactory quality, explaining why minerality correlates with acidity and freshness in many studies and why the descriptor is applied inconsistently across assessors. The regional and variety associations of mineral character show consistent patterns, with Chablis, Sancerre, and German Riesling expected to show mineral expression while warm-climate fruit-forward wines are not, suggesting that minerality perception follows learned style templates rather than direct compositional causation. Understanding mineral aroma quality requires acknowledging the gap between common usage and scientific understanding while recognizing minerality's functional importance in wine quality communication and terroir discourse.
Mineral_Aroma_Quality addresses the controversial but commercially important minerality concept in wine evaluation. This quality requires acknowledgment of scientific uncertainty about its chemical basis while recognizing its functional importance in wine discourse. The class may represent a multi-modal perception rather than pure olfactory quality, integrating smell, taste, and texture inputs into a gestalt impression associated with terroir and quality.
Mineralität
Mineralitätsnote
mineral notes
mineralischer Duft
minerality
steiniger Charakter
stony character
wet rock scent
Parr WV et al. (2018) Minerality in wine: a review. Journal of Wine Research 29(2): 82-98 Heymann H et al. (2014) A study of the perceptions of minerality in Sauvignon blanc wines. Journal of Wine Research 25(4): 259-273
Mineral Aroma Quality
Mineral balance quality assessment in Ribera del Duero Tempranillo integrates the potassium-calcium-magnesium ratios that affect must pH in this continental climate region, with optimal balance (K/Ca+Mg ratio below 3.0) producing harvest pH of 3.4-3.7 that supports stable color and fresh acidity, while imbalanced nutrition (ratio above 4.0) generates the elevated pH (3.8-4.2) requiring tartaric acid adjustment rootstock selection targeting balanced mineral uptake (110R, 1103P rather than high-potassium SO4) provides the foundation for achieving favorable mineral balance in the region's calcareous clay soils.
Mineral_Balance_Quality encompasses the chemical qualities characterizing the proportional relationships among mineral elements in grape tissue and must, capturing the ratios and equilibria that affect nutrient interactions, wine chemistry, and quality outcomes beyond what individual element concentrations reveal. Mineral balance recognizes that elements interact competitively and synergistically during uptake, transport, and function, making ratios among elements often more informative than absolute concentrations. This category includes Cation_Exchange_Quality addressing the overall cation balance affecting pH and stability, and K_Ca_Mg_Ratio_Quality capturing the relationship among the three major divalent and monovalent cations with distinct uptake mechanisms and physiological functions. The balance between potassium and divalent cations (calcium, magnesium) affects must pH through differential effects on organic acid dissociation and salt formation. Excessive potassium relative to calcium and magnesium produces elevated pH even at moderate total cation content, while balanced cation nutrition supports pH in desirable ranges. The antagonistic interactions among cations during root uptake mean that soil ratios affect vine ratios, creating terroir effects through mineral nutrition. Rootstock selection profoundly affects mineral balance, with rootstocks varying in selectivity for potassium versus calcium and magnesium. The concept of mineral balance extends to micronutrients, where ratios among iron, manganese, zinc, and copper affect mutual uptake and utilization. Assessment of mineral balance requires multi-element analysis enabling ratio calculations, with interpretation based on established norms for acceptable ratios in vine tissue and must.
Mineral_Balance_Quality serves as the parent class for specific ratio qualities, positioned within Mineral_Content_Quality to capture compositional relationships rather than absolute concentrations. The concept parallels acid balance in recognizing that proportions affect quality beyond individual component levels. Mineral balance qualities are derived measurements calculated from primary element analyses rather than directly measured properties. The terroir significance of mineral balance through soil-to-vine-to-wine transfer makes this category relevant to geographic quality expression and site selection considerations.
Kationengleichgewicht
Mineralbalance
Mineralstoffverhältnis
Nährstoffverhältnisqualität
cation balance
elemental equilibrium quality
mineral ratio quality
nutrient ratio composition
Mpelasoka BS et al. (2003) A Review of Potassium Nutrition in Grapevines with Special Emphasis on Berry Accumulation. Australian Journal of Grape and Wine Research 9(3): 154-168 Garcia M et al. (2001) Influence of the Mineral Status of the Vineyard on the Quality of the Vintage. Acta Horticulturae 554: 205-212
Mineral Balance Quality
Mineral content quality profiling in Chablis Chardonnay addresses the regional claim of mineral-driven wine character, with limestone-derived soils providing calcium-rich, moderately potassium-limited conditions (must potassium 800-1200 mg/L versus 1500-2500 mg/L in warmer regions) that produce naturally lower pH wines requiring less acid adjustment while direct sensory contribution of minerals remains scientifically contested, the indirect effects on pH, fermentation, and malolactic conversion create the conditions for the steely, chalky-textured wines that consumers describe as mineral.
Mineral_Content_Quality encompasses all chemical qualities that characterize the concentration and composition of inorganic elements in grapevine tissues, capturing the essential nutrients and trace elements that influence vine physiology, grape composition, and wine chemistry. Minerals in grape berries derive from soil uptake through root systems, with translocation through xylem and phloem determining the distribution among vine organs. The mineral composition of grapes affects multiple quality dimensions: vine nutrition and productivity through essential nutrient supply must fermentation through effects on yeast metabolism wine stability through precipitation of mineral salts and potentially wine sensory character through the controversial concept of minerality. This category subdivides into Macronutrient_Content_Quality addressing the major elements required in large quantities (potassium, calcium, magnesium, phosphorus, sulfur, nitrogen), and Micronutrient_Content_Quality capturing trace elements essential in small amounts (iron, manganese, zinc, copper, boron, molybdenum). Potassium dominates the mineral composition of grape berries, typically comprising 50-70% of total mineral content, with critical effects on must pH and wine tartrate stability. The total mineral content of grape juice typically ranges from 2-5 g/L, with substantial variation reflecting cultivar genetics, rootstock selection, soil composition, and viticultural management. Soil mineral availability, modified by pH, organic matter content, and water status, determines root uptake rates, while vine demand varies with growth stage, crop load, and environmental conditions. The concept of terroir includes mineral composition as a potential contributor to site-specific wine character, though the mechanistic links between soil minerals, grape minerals, and wine sensory properties remain debated. Mineral analysis employs atomic absorption spectroscopy, inductively coupled plasma techniques (ICP-OES, ICP-MS), or ion chromatography depending on the elements of interest and required sensitivity.
Mineral_Content_Quality is positioned as a direct subclass of Chemical_Quality, parallel to Sugar_Content_Quality, Acid_Content_Quality, and other composition categories. The distinction between macronutrients and micronutrients reflects nutritional classification based on required quantities rather than chemical properties. Nitrogen is typically addressed separately under Nitrogen_Content_Quality due to its unique importance in fermentation and its organic as well as inorganic forms. The controversial concept of minerality in wine tasting creates tension between scientific skepticism about direct mineral taste contribution and widespread use of mineral descriptors by wine professionals.
Aschegehaltqualität
Mineralstoffgehalt
Mineralzusammensetzung
anorganischer Elementgehalt
ash composition quality
elemental content
inorganic element content
mineral composition quality
Mpelasoka BS et al. (2003) A Review of Potassium Nutrition in Grapevines with Special Emphasis on Berry Accumulation. Australian Journal of Grape and Wine Research 9(3): 154-168 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 341-385
Mineral Content Quality
Cabernet Sauvignon in Napa Valley shows minimal growth period during early-mid September Year N+1 at BBCH 81, with berry growth rate sustained at 2-4 mg per day over 2.5-3.5 week period, dimensions stable at 16.2-16.8 mm diameter and 12.8-14.2 g fresh weight, seeds completing maturation with hard dark brown coats, anthocyanin biosynthesis genes progressively increasing 8-14 fold, sugar transporters increasing 5-8 fold, sugars rising from 12% to 15%, under warm climate conditions of 22-28°C temperature, typical of this late-season cultivar in sustained lag quiescence
Minimal growth period represents the sustained quiescence phase during berry growth lag phase when berry maintains near-zero growth rate typically 1-5 mg fresh weight per day, dimensional and compositional stability characterize berry state, seed maturation completes establishing developmental readiness for progression, pre-veraison metabolic preparations continue building ripening capacity, and sustained minimal growth creates developmental pause enabling metabolic reorganization before veraison transformation, proceeding over 1-3 weeks as main lag phase duration. This sustained quiescence phase occurs during late summer to early autumn in late August-September in temperate regions Year N+1, spanning variable 1-3 weeks depending on cultivar and environment representing main lag duration, corresponding to BBCH 81 beginning of ripening stage pre-veraison period. Minimal growth characteristics shows growth rate sustained at very low levels 1-5 mg fresh weight per day throughout period, occasional slightly negative growth from water loss in deficit conditions, essentially stable berry mass distinguishing minimal growth from enlargement growth and ripening expansion, and sustained low rate as defining lag characteristic. Near-zero growth rate shows typical berry fresh weight gain <3 mg per day representing essential growth cessation, growth effectively arrested at developmental quiescence, contrast with enlargement phase 50-250 mg per day and ripening phase 20-100 mg per day, and minimal rate enabling metabolic focus on preparation rather than growth. Berry dimensional stability shows berry diameter remaining constant within 1-3% variation representing measurement and minor fluctuation ranges, fresh weight stability within 2-5% over minimal growth period, volumetric constancy characteristic of quiescent state, and dimensional stability contrasting with dynamic growth and ripening phases. Berry size during minimal growth shows dimensions established during enlargement phase maintained, typical ranges 6-10 mm small-berried, 10-15 mm medium-berried, 15-23 mm large-berried cultivars remaining constant, fresh weight 2-6 g small, 5-13 g medium, 10-28 g large maintaining stability, and size constancy as lag defining feature. Water relations during minimal growth shows water uptake very low <0.05 g per berry per day, transpiration water loss approximately balancing import, net water flux near-zero establishing hydration equilibrium, water content 78-82% stable over period, and water balance characteristic of quiescence. Water balance equilibrium shows uptake-transpiration balance preventing net accumulation or loss, stable berry hydration maintaining composition, equilibrium as quiescent state feature, and balance contrasting with net uptake during growth and ripening. Seed development focus shows seeds completing final maturation during minimal growth period, seed coat fully lignified creating hard brown protective layer, embryo achieving full developmental maturity with viable embryonic axis, endosperm completing nutrient storage, and seed maturation as critical lag phase process. Seed maturation completion shows seeds reaching full maturity by mid-to-late lag phase, seed color dark brown to black indicating maturation, seed texture hard and dry from lignification and desiccation, germination capacity achieved if seeds separated, and mature seeds as developmental prerequisite for veraison. Seed maturity indicators shows seed coat color change from green to brown to dark brown, hardening from lignin deposition, moisture decline from desiccation, and indicators documenting maturation state. Seed-berry developmental coordination shows seed maturation possibly signaling berry developmental readiness, seed hormones potentially regulating berry progression, mature seeds enabling veraison triggering, and seed-berry communication coordinating development. Pre-veraison metabolic preparations shows continued metabolic changes building ripening capacity during minimal growth, ripening gene expression gradually increasing from low initiation levels, enzyme accumulation preparing for veraison activation, metabolic pathways priming for rapid ripening transformation, and preparation as key minimal growth function. Ripening gene expression progression shows anthocyanin biosynthesis genes in red cultivars progressively increasing 5-15 fold over lag from early initiation levels, sugar transporter genes increasing 3-8 fold preparing import capacity, softening enzyme genes gradually increasing 2-6 fold, and progressive expression building ripening machinery. Anthocyanin biosynthesis preparation shows continued low-level gene expression, enzyme protein accumulation increasing slowly, pathway components assembling for veraison activation, and preparation enabling rapid color change at veraison in red cultivars. Sugar transport preparation shows hexose transporter expression increasing progressively, transporter protein accumulation in plasma membrane and tonoplast, transport capacity building for rapid sugar import during ripening, and preparation as critical pre-ripening function. Softening enzyme preparation shows polygalacturonase, pectin methylesterase, expansin genes increasing expression, enzyme precursors accumulating, cell wall modification machinery preparing for softening, and preparation enabling texture transformation at veraison. Berry composition during minimal growth shows water 78-82% stable with minor fluctuations, organic acids declining slowly from 0.8-1.2% to 0.7-1.1% over period from continued respiration, sugars increasing slowly from 11-15% to 12-16% from limited phloem import, pH rising gradually from 3.0-3.2 to 3.1-3.3 from acid decline, and composition showing slow progressive changes toward ripening. Organic acid dynamics shows malate declining slowly through respiration 0.1-0.2% over minimal growth period, tartrate relatively stable maintaining 0.3-0.6% concentrations, total acids declining creating gradual pH increase and softening contribution, and acid respiration as continuing metabolic process. Sugar dynamics shows limited sugar accumulation 0.5-1.5% per week from low-rate phloem import, progressive increase preparing for ripening acceleration, sugar predominantly stored in vacuole, and slow accumulation characteristic of lag metabolism. Metabolic activity during minimal growth shows respiratory rate sustained at low levels 1.0-1.5 μmol O₂ g⁻¹ h⁻¹ over minimal growth period, metabolic activity adequate for maintenance and preparation, minimal biosynthetic activity, and sustained low metabolism characteristic of quiescence. Berry morphology during minimal growth shows berry maintaining stable appearance, green color or beginning chlorophyll decline depending on cultivar and timing, texture firm though softer than peak growth firmness, surface smooth, and appearance as hard green berry in quiescent state. Environmental effects on minimal growth include temperature affecting minimal growth period duration with warm conditions shortening period 1-2 weeks and cool conditions extending period 2-3 weeks, water availability affecting berry hydration and stress response, moderate water deficit possibly accelerating veraison onset shortening lag, and environmental factors determining lag duration. Temperature effects on duration shows heat accumulation through lag phase advancing progression toward veraison, growing degree days correlating with lag duration, warm conditions accelerating metabolic preparations and development, cool conditions extending quiescence period, and temperature as major lag duration determinant. Water deficit effects shows moderate deficit possibly triggering earlier veraison through stress signaling shortening minimal growth period, severe deficit potentially arresting development extending lag, and water stress as factor affecting progression. Minimal growth period duration shows cultivar variation with early-ripening cultivars having shorter minimal growth 1-2 weeks and late-ripening cultivars longer 2-3 weeks, environmental conditions particularly temperature affecting duration within genetic ranges, and combined genetic-environmental control determining lag duration. The process shows three distinguishable aspects including near-zero growth rate sustaining quiescence, seed development focus completing maturation, and pre-veraison metabolism preparing ripening. Duration typically 1-3 weeks varying by cultivar and environment representing main lag phase period. Regional timing shows minimal growth period during late August-September across climates as sustained quiescence. Viticultural significance includes representing main lag phase quiescence enabling metabolic preparation, allowing seed maturation completion prerequisite for ripening, determining veraison timing through lag duration, and establishing berry in stable state before ripening transformation. Research applications include analyzing quiescence maintenance and regulation, investigating seed maturation and berry-seed coordination, evaluating pre-veraison metabolic preparations, quantifying environmental effects on lag duration, and assessing minimal growth period as determinant of veraison timing and ripening success.
Minimal_Growth_Period represents sustained quiescence within Berry_Growth_Lag_Phase. Subdivided into Near_Zero_Growth_Rate sustaining 1-5 mg/day, Seed_Development_Focus completing maturation, and Pre_Veraison_Metabolism preparing ripening. Duration 1-3 weeks (main lag duration). Berry dimensions stable within 1-3%. Water balance equilibrium. Ripening genes progressively increasing 2-15 fold. Seeds reaching full maturity. Follows Lag_Phase_Initiation. Precedes Veraison. Temperature major duration determinant.
Haupt-Lag-Periode
Wachstumsplateau-Phase
anhaltende Ruhephase
growth plateau phase
main lag period
minimale Wachstumsperiode
sustained quiescence
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 456-462 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Minimal Growth Period
A Shiraz vine in Australia's Riverina managed under minimal pruning bears a Minimal_Pruning_Role, with no annual pruning performed for 15+ years creating a dense canopy ball producing 25-30 kg of small-berried, intensely flavored fruit suited to commercial red wine production. A Cabernet Sauvignon vine in Sunraysia minimal pruning trials serves in a Minimal_Pruning_Role, research documenting the variety's adaptation to self-regulating production with berry weights 30-40% lower than conventionally pruned vines. A Colombard vine in hot Australian irrigated production bears a Minimal_Pruning_Role, the white variety's adaptation to minimal intervention enabling economically viable production for distillation and bulk wine markets.
A training system role that inheres in a grapevine managed under minimal pruning regimes, where annual pruning is eliminated or drastically reduced, allowing the vine to develop its natural growth form with accumulated multi-year wood creating a dense, self-regulating canopy. Minimal pruning (also termed non-pruning or zero pruning) exploits the grapevine's natural self-regulation mechanisms—as unpruned vines accumulate fruiting positions, individual cluster size and berry size decrease, partially compensating for increased cluster numbers through what viticulturists term "natural crop adjustment." Australian research (CSIRO, University of Adelaide) pioneered minimal pruning development from the 1970s, demonstrating that certain varieties in specific environments could produce acceptable fruit quality without annual pruning labor. Minimal pruning creates distinctive vine appearance—dense, rounded canopy masses rather than the linear hedgerows of trellised systems—with fruit dispersed throughout the canopy at variable heights and positions. The system is limited to warm, dry climates where disease pressure from dense canopy architecture is manageable, and to production contexts accepting the fruit characteristics (smaller berries, variable maturity, distinctive flavor profiles) that result from minimal intervention.
Minimal_Pruning_Role captures the most extreme reduction in viticultural intervention, eliminating the defining annual practice that distinguishes cultivated grapevines from wild plants. The system's adoption remains limited—primarily Australian warm-climate production, some California Central Valley applications—reflecting the specific conditions required for success (warm, dry climate disease-tolerant varieties appropriate market positioning) and the quality perceptions limiting adoption in premium segments. Minimal pruning research demonstrated that annual pruning, while universal in traditional viticulture, is not physiologically essential—vines survive and produce without it—prompting reconsideration of pruning's functions and optimal intensity. The smaller berries produced under minimal pruning may actually enhance some wine characteristics (higher skin:juice ratio, more concentrated flavors), challenging assumptions about pruning-quality relationships. Climate change and labor cost pressures may increase minimal pruning interest, particularly for varieties and markets where the distinctive fruit characteristics are acceptable or valued.
Minimalschnitt
Nicht-Schnitt
Nullschnitt
minimal pruning
non-pruning
ungeschnittene Rebe
unpruned vine
zero pruning
Clingeleffer PR (1984) Production and growth of minimal pruned Sultana vines, Vitis 23: 42-54 Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Possingham JV (1994) New concepts in pruning grapevines, Horticultural Reviews 16: 235-254 Intrieri C, Poni S (1995) Integrated evolution of training systems, American Journal of Enology and Viticulture 46: 507-517
Minimal Pruning Role
A minimal short fruiting cane on ancient Vitis vinifera cv. Grenache in Châteauneuf-du-Pape, retained at 4 nodes spanning 45cm from gnarled permanent arm on 80-year-old vine in rocky garrigue soil, selected from weak-vigor shoot (5mm diameter but fully lignified showing excellent maturity for small size), positioned carefully to avoid mechanical stress on fragile old wood, expected to produce 3-4 weak shoots bearing 5-6 small clusters totaling 0.8-1.1kg fruit from entire cane, contributing to ultra-low whole-vine yield of 2.2kg (approximately 1.5 tons/hectare at wide 2m×2m spacing) supporting extraordinary concentration, phenolic development, and aromatic complexity justifying premium Châteauneuf pricing and critical acclaim for old-vine terroir expression.
A short fruiting cane retained at absolute minimum length while maintaining cane rather than spur designation, typically 4-5 nodes, representing lower extreme of cane retention with bud counts barely exceeding standard spur range, characterized by compact extent (40-55cm), very limited bud retention emphasizing extreme quality focus, suitability only for exceptional fruitfulness situations or ultra-premium production accepting severe yield restrictions, and production of few fruiting shoots providing minimal but hopefully exceptional fruit quality. Minimal short fruiting canes represent quality extremes rarely encountered in commercial viticulture, appropriate only in special circumstances including experimental ultra-low-yield wines, specific terroir-focused cuvées from prestigious sites, recovery seasons following severe damage requiring conservative crop loads, or naturally low-vigor old vines unable to support longer cane retention, with this minimal approach sacrificing economic efficiency for potential quality gains in highly selective applications.
Minimal short fruiting canes are exceptional rather than standard practice, found primarily in old-vine situations where declining vigor necessitates conservative retention, ultra-premium producers pursuing quality extremes through severe crop restriction, or specialty cuvées from particular vineyard blocks designated for minimal-yield wines. The 4-5 node range barely qualifies as cane rather than spur, with distinction sometimes arbitrary depending on training system architecture. Economic viability requires exceptional wine prices justifying yields often 25-50% of regional norms, with sustainability concerns about old vine preservation versus production demands creating tensions in some prestigious regions. The minimal approach produces exceptional individual wines but raises questions about long-term vineyard viability when chronic low yields deplete vine reserves and accelerate decline in already weakened vines.
barely-cane short retention
extrem kurze Fruchtrute
minimal-kurze Rute
quality-extreme short cane
ultra-minimal cane
Based on Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Bravdo et al. (1985) "Effect of crop level and crop load on growth, yield, must and wine composition, and quality of Cabernet Sauvignon" American Journal of Enology and Viticulture 36=125-131 Reynolds & Vanden Heuvel (2009) "Influence of grapevine training systems on vine growth and fruit composition= a review" American Journal of Enology and Viticulture 60=251-268.
Minimal Short Fruiting Cane
A minimal tertiary vegetative bud at node 12 on Vitis vinifera cv. Cabernet Sauvignon in a severely damaged vineyard, measuring only 0.6mm diameter with barely detectable primordial structures showing 2-3 possible leaf initials under 40x magnification, this bud showing no visible swelling 25 days post-damage while adjacent node tertiaries showed initial breaking, eventually producing extremely weak shoot growth reaching only 12cm with 4 tiny malformed leaves before growth cessation in early July, providing essentially zero photosynthetic contribution and producing no detectable buds for following season, effectively representing failed node position requiring adjacent node compensation for any canopy recovery.
A tertiary vegetative bud at the lower extreme of developmental capacity, characterized by barely distinguishable primordial structures, questionable viability even when released from suppression, extremely rudimentary organization with perhaps only 2-4 leaf initials visible under magnification, and uncertain developmental potential that may result in failed budbreak, aborted shoot development, or extremely weak vegetative growth insufficient for meaningful canopy contribution. Minimal tertiary vegetative buds represent the practical limit of compound bud redundancy, often present in anatomical sense but functionally marginal as viable meristems, with their presence providing theoretical backup capacity but uncertain practical value for vineyard recovery, making nodes containing only minimal tertiary buds effectively lost positions requiring canopy reconstruction from adjacent nodes or latent bud activation.
Minimal tertiary vegetative buds blur the line between viable meristem and non-functional tissue, with their developmental capacity highly variable and unpredictable. Factors affecting minimal tertiary viability include node position (apical and basal positions showing poorest development), cultivar (some varieties invest more minimally in tertiary positions), previous season growing conditions during bud formation (stress reducing tertiary development), and vine age (older vines sometimes showing declining tertiary quality). The existence of minimal tertiaries demonstrates the limits of compound bud insurance strategy - beyond a certain damage threshold, even the three-bud system cannot provide meaningful recovery, and alternative mechanisms (latent buds, adjacent node compensation) become necessary for vine survival and canopy reconstruction.
barely viable tertiary bud
minimale Tertiärknospe
rudimentary tertiary bud
rudimentäre dritte Knospe
vestigial tertiary vegetative bud
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press, pp. 85-92.
Minimal Tertiary Vegetative Bud
Syrah grafted on 3309 Couderc rootstock at 13 years showing minor branch secondary roots as third and fourth-order branches, representative minor branch 0.8 cm diameter at origin extending 28 cm length at depth 52 cm, minor branch showing limited secondary growth with 3-4 annual xylem rings, minor branch producing fourth and fifth-order fine roots 0.1-0.4 cm diameter at 2-5 cm intervals, minor branch tips showing active root hair production and mycorrhizal colonization, minor branch demonstrating mixed absorption-transport function connecting finest absorptive roots to major branch framework
Minor branch secondary root is branch secondary root showing small diameter typically 0.2-1.5 cm and limited structural development, emphasizing small size and transitional absorptive-transport function within branch root hierarchy, characterized by third-order or higher position branching from major branch secondary roots or other minor branches, minimal to moderate secondary growth with limited xylem development, relatively short lifespan typically 1-4 years intermediate between ephemeral fine roots and persistent major branches, and serving dual functions as both direct absorption particularly at growing tips and transport conduit moving water and nutrients from finest absorptive roots to larger structural branches, with minor branch roots typically emerging 3-10 cm intervals along parent root creating moderately dense branching pattern, producing numerous fine absorptive roots and root hairs, and representing transitional root order between large persistent structural roots and small ephemeral absorptive roots providing critical link in hierarchical resource acquisition and transport pathway. Minor branch secondary root size characteristics includes= showing small to moderate diameter (minor branches typically 0.2-1.5 cm considerably smaller than major branches but larger than finest absorptive roots), exhibiting modest length (minor branch roots typically 10-40 cm exploring local soil volume), demonstrating limited biomass (minor branches containing relatively small tissue mass individually though collectively significant due to abundance), displaying subtle structure (minor branch roots often overlooked in casual observation being intermediate between prominent major branches and microscopic fine roots), and revealing size variability (minor branch diameter and length varying with position, order, and environmental conditions). Minor branch secondary root structural development includes= showing limited secondary growth (minor branch roots developing limited secondary xylem perhaps 2-5 annual rings), exhibiting transitional anatomy (minor branches showing intermediate structure between woody major branches and primary anatomy fine roots), demonstrating thin bark (limited periderm development on minor branches compared to thick bark on major branches), displaying moderate lignification (some cell wall lignification in minor branches but less extensive than major branches), and revealing structural limitations (minor branch structure adequate for local support and transport but insufficient for major structural role). Minor branch secondary root hierarchical position includes= typically third or fourth-order (minor branch roots most commonly arising from second-order major branches or third-order minor branches), occasionally fifth-order (some minor branches in highly branched systems reaching fifth or higher order), serving as parent to finest roots (minor branches producing fourth fifth and sixth-order fine absorptive roots), demonstrating transitional position (minor branches occupying middle ground between structural framework and absorptive periphery), and showing high abundance (minor branches numerous due to high branching multiplication at intermediate orders). Minor branch secondary root lifespan includes= functioning 1-4 years typically (minor branch roots showing intermediate longevity longer than annual fine roots but shorter than decade-lasting major branches), exhibiting turnover (minor branches continuously produced to replace dying or senescing branches), demonstrating moderate investment (minor branch construction requiring moderate carbon investment providing moderate return over intermediate lifespan), showing variable mortality (minor branch death rate varying with environmental stress, age, and pathogen pressure), and requiring continuous replacement (ongoing minor branch production necessary to maintain intermediate branching level despite turnover). Minor branch secondary root fine root production includes= generating numerous fine roots (minor branches producing 5-20+ fine absorptive roots creating dense local network), showing close spacing (fine roots emerging 1-5 cm intervals along minor branch reflecting fine-scale branching), exhibiting prolific branching (minor branches showing high branching intensity due to transition toward absorptive function), demonstrating responsive production (minor branch fine root production increasing in favorable zones and decreasing in poor zones), and creating absorption clusters (minor branch with fine roots forming dense absorptive unit thoroughly exploiting local soil). Minor branch secondary root functional characteristics includes= showing mixed absorption-transport (minor branches both absorbing directly particularly at tips and conducting from finer roots), demonstrating active absorption (minor branch tips producing root hairs and absorbing water and nutrients directly), exhibiting transport capacity (minor branch xylem and phloem conducting resources between finest roots and major branches), displaying storage limitation (minor branch small size limiting carbohydrate storage capacity to modest amounts), and revealing seasonal activity (minor branches showing active growth and absorption during growing season with reduced activity during dormancy). Minor branch secondary root in absorption network includes= representing largest absorptive roots (minor branches being largest-diameter roots actively involved in direct absorption through root hairs), providing absorption surface (minor branch surface area particularly at growing tips contributing to total vine absorption), hosting mycorrhizae (minor branch roots showing arbuscular mycorrhizal colonization enhancing nutrient and water uptake), demonstrating absorption efficiency (minor branch intermediate position optimizing balance between absorption surface and transport efficiency), and enabling responsive growth (minor branches rapidly elongating and branching in resource-rich soil patches). Minor branch secondary root spatial distribution includes= concentrating where resources abundant (minor branch density highest in fertile moist upper soil and resource-rich patches), showing patchy distribution (minor branches more numerous in favorable zones less abundant in poor zones reflecting proliferation response), occupying upper to middle soil (minor branches typically 20-90 cm depth where resources most available), demonstrating local exploration (minor branch network thoroughly exploring soil volume around parent major branch), and creating nested hierarchy (minor branches distributed along major branches which distribute along first-order laterals creating multi-scale organization).
Minor_Branch_Secondary_Root emphasizes small size and transitional function and is distinguished from Major_Branch_Secondary_Root (large structural) by focus on intermediate absorptive-transport role. Critical for understanding intermediate root orders, absorption-transport transition, moderate turnover, fine root production, and the small-diameter branch roots linking ephemeral absorptive network to persistent structural framework.
Intermediate Seitenwurzel
Intermediate branch root
Kleine Verzweigungswurzel
Small branch root
Transitional secondary branch
Übergangs-Nebenwurzel
Minor Branch Secondary Root
Comprehensive evaluation of Mite_Resistance_Disposition at Oregon State University assessed responses to both eriophyid (Colomerus vitis) and tetranychid (Eotetranychus willamettei) mites across 35 cultivars in Willamette Valley vineyard trials. Erineum development was scored visually (0-5 scale for leaf coverage), while spider mite populations were monitored by leaf brushing counts (mites per leaf) at biweekly intervals through the growing season. Substantial cultivar variation in Mite_Resistance_Disposition was detected for both mite groups. V. vinifera 'Pinot Noir' showed high susceptibility to both pests, with erineum scores of 3.8 and peak spider mite densities of 45 per leaf requiring acaricide intervention. 'Müller-Thurgau' exhibited differential resistance, with low erineum formation (score 0.8) but high spider mite susceptibility (52 per leaf). Conversely, 'Gewürztraminer' supported extensive erineum (score 4.2) but maintained low spider mite populations (8 per leaf), potentially due to dense leaf trichomes harboring predatory Typhlodromus mites. The hybrid 'Maréchal Foch' demonstrated broad Mite_Resistance_Disposition with minimal erineum (score 0.5) and low spider mite populations (6 per leaf). These uncorrelated resistance patterns confirm that Mite_Resistance_Disposition comprises independent mechanisms targeting different mite groups, requiring separate evaluation and breeding approaches.
A Mite_Resistance_Disposition is a Pest_Resistance_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through defense processes that limit feeding, colonization, reproduction, or damage by phytophagous mites, microscopic arachnids that attack grapevine foliage, buds, and other tissues. This disposition represents the capacity to resist two ecologically and morphologically distinct groups of plant-feeding mites: eriophyid mites (Eriophyoidea) including the grape erineum mite (Colomerus vitis) and grape bud mite (C. vitis and Calepitrimerus vitis), and tetranychid spider mites (Tetranychidae) including the Pacific spider mite (Tetranychus pacificus), Willamette spider mite (Eotetranychus willamettei), and European red mite (Panonychus ulmi). These mite groups differ fundamentally in biology, feeding behavior, and damage symptoms: eriophyid mites are vermiform, highly host-specific, and induce characteristic gall structures (erinea) or bud abnormalities, while spider mites are oval-bodied, often polyphagous, and cause stippling damage through cell puncturing and chlorophyll extraction. Mite resistance mechanisms in plants include leaf surface properties (trichome density and morphology, epicuticular wax) affecting mite movement and feeding access, constitutive or induced chemical defenses (terpenoids, phenolics) toxic or deterrent to mites, rapid wound responses limiting feeding success, and domatia or other structures harboring predatory mites that provide biological control. For grapevine, mite management has traditionally relied on acaricides and conservation of natural enemies, but cultivar differences in mite susceptibility provide opportunities for host plant resistance as a component of integrated mite management.
Mite_Resistance_Disposition is_a Pest_Resistance_Disposition. This class encompasses resistance to all phytophagous mites affecting grapevine, organized into child classes by mite family: Eriophyid_Mite_Resistance_Disposition (gall-forming and bud-infesting eriophyoid mites) and Spider_Mite_Resistance_Disposition (tetranychid mites causing stippling). Resistance mechanisms and cultivar rankings differ between these mite groups, requiring separate consideration. Mites are arachnids, not insects, justifying their placement in a separate resistance class parallel to Insect_Resistance_Disposition.
Milbenresistenz-Disposition
Phytophage-Milben-Resistenz
Spinnmilben-Abwehr
acarid resistance
mite feeding resistance
phytophagous mite resistance
Duso C et al. (2010) Biological control of mites in European vineyards, Arthropod Management in Vineyards, Springer Walton VM et al. (2007) Relationship between rust mites Calepitrimerus vitis (Nalepa) and bud damage on selected grape cultivars, Vitis 46:66-70 Helle W, Sabelis MW (1985) Spider Mites: Their Biology, Natural Enemies and Control, Elsevier
Mite Resistance Disposition
The mitochondrial DNA sequence of Vitis vinifera cv. Pinot Noir (Goremykin et al. 2009, GenBank accession FM179380) revealed a 773,279 bp genome — approximately 4.8 times larger than the chloroplast genome and among the largest plant mitochondrial genomes characterized — containing only 59 identified genes (38 protein-coding, 18 tRNA, 3 rRNA) but extensive intergenic sequence including fragments of chloroplast DNA incorporated through intracellular gene transfer (approximately 6% of the mitochondrial genome derived from cpDNA insertions), nuclear-derived transposable elements, and large repeated sequences that mediate recombination-driven structural rearrangements. Mitochondrial sequence polymorphisms at the nad1 exon B-C intron and the cox2-cox3 intergenic spacer have been used to distinguish Vitis vinifera from American Vitis species in rootstock identification and to detect mitochondrial introgression from V. rupestris and V. berlandieri in interspecific hybrids.
A DNA sequence derived from the mitochondrial genome of Vitis vinifera, a large and structurally complex circular DNA molecule of approximately 773,279 base pairs located in the mitochondrial organelle, maternally inherited, encoding genes essential for oxidative phosphorylation and cellular energy metabolism, and providing an additional uniparentally inherited marker system for evolutionary studies, though less frequently used than chloroplast DNA in grapevine phylogeography due to the mitochondrial genome's structural instability and recombinational complexity.
Mitochondrial_DNA_Sequence documents the third compartmental genome of the grapevine cell — a large, structurally dynamic molecule whose gene content is essential for cellular energy metabolism but whose utility for cultivar identification and phylogeographic analysis is limited by several distinctive features of plant mitochondrial genomics. Unlike animal mtDNA (which is small, compact, and rapidly evolving), the Vitis mitochondrial genome is exceptionally large (773 kb versus 16 kb in mammals) with very low gene density (59 genes in 773 kb versus 37 genes in 16 kb): the extensive intergenic sequence undergoes frequent intramolecular recombination through large repeated sequences, generating alternative genomic conformations (subgenomic circles, linear molecules) that complicate genome assembly and sequence comparison. The slow nucleotide substitution rate of plant mtDNA (5-10 times slower than cpDNA) further reduces its phylogenetic informativeness for intraspecific studies, explaining why chloroplast rather than mitochondrial markers dominate grapevine evolutionary research. However, mitochondrial sequences retain value for interspecific hybrid identification and rootstock characterization because species-diagnostic polymorphisms are well-established at the Vitis genus level.
Mitochondrien-DNA-Sequenz
mitochondrial genome sequence
mitochondrial genomic data
mitochondriale Genomsequenz
mtDNA sequence
mtDNA-Daten
Goremykin VV et al. (2009) The mitochondrial genome of Vitis vinifera, in Bentham Science Publishers, Current Genomics 10: 134-141 Picardi E et al. (2010) Large-scale evidence for intranuclear transfer of organelle DNA in Vitis vinifera, Genome Biology and Evolution 2: 10-15 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519
Mitochondrial DNA Sequence
Pinot Noir in Burgundy shows mitotic activity onset during early July Year N+1 at BBCH 71, with first divisions occurring days 6-9 post-fertilization following DNA replication, M phase mitosis with chromosome segregation and cytokinesis, initial cell cycle times 13-15 hours, progressive division onset across mesocarp cell populations, observable mitotic index increasing from <1% to 3-8% during week 1-2, under continental conditions of 22-27°C temperature, typical of this early-to-mid season cultivar establishing proliferative state
Mitotic activity onset represents the first division occurrence phase during active berry cell division when activated cells complete DNA replication, enter M phase, and undergo first mitotic divisions producing daughter cells, establishing observable proliferative state and initiating exponential cell number increase, typically occurring within first week to 10 days post-fertilization as cell cycle activation culminates in division. This critical division initiation phase occurs during early summer in late June-early July in temperate regions Year N+1, typically days 5-10 post-fertilization following activation, corresponding to BBCH 71 young fruit early development stage. Mitotic onset mechanism involves G1 to S phase transition with DNA replication initiation, S phase DNA synthesis doubling chromosome number from 2n to 4n, G2 phase preparation for mitosis, M phase entry with chromatin condensation and mitotic spindle formation, chromosome segregation to daughter cells, cytokinesis creating cell plate dividing cytoplasm, and completion producing two daughter cells re-entering G1 enabling subsequent divisions. S phase entry and DNA replication shows cyclin E-CDK2 complex activation triggering replication origin firing, DNA polymerase recruitment to licensed replication origins, bidirectional replication fork progression along chromosomes at 0.5-2 kb per minute, replication checkpoint surveillance ensuring complete faithful replication, and S phase completion within 6-10 hours during early berry development. DNA synthesis characteristics show semiconservative replication producing two daughter DNA molecules each with one original and one new strand, high fidelity polymerase activity with proofreading maintaining <1 error per 10⁹ nucleotides, coordinated replication across entire 475 Mb Vitis vinifera genome, and replication completion required before mitosis entry. G2 phase preparation shows continued cell growth, checkpoint surveillance ensuring complete DNA replication and cell size adequacy, centrosome duplication preparing spindle formation, and G2-M transition checkpoint passage enabling mitosis entry. M phase mitosis shows chromatin condensation into visible chromosomes, nuclear envelope breakdown releasing chromosomes into cytoplasm, mitotic spindle formation from centrosome-nucleated microtubules, chromosome alignment at metaphase plate, anaphase segregation distributing sister chromatids to spindle poles, and telophase nuclear envelope reformation around separated chromosome sets. Chromosome segregation fidelity shows kinetochore-microtubule attachment providing force for chromosome movement, spindle assembly checkpoint ensuring proper attachment before anaphase, accurate 19 per 19 chromosome distribution to daughter cells maintaining diploid 2n state, and high segregation fidelity preventing aneuploidy. Cytokinesis and cell division completion shows phragmoplast microtubule array formation at division plane, vesicle fusion initiating cell plate synthesis, progressive cell plate expansion from center outward, cell wall deposition creating new wall between daughter cells, and completion producing two separate cells each re-entering G1 phase. First division timing shows earliest divisions occurring days 5-7 post-fertilization in early-activating cells near seeds, progressive division onset across cell populations days 6-10 creating temporal spread, and completion of first division wave by day 10 establishing proliferative tissue. Division rate acceleration shows initial slower division during first round, acceleration as daughter cells re-enter cycle more rapidly, progressive rate increase as cell populations synchronize, and transition toward peak division rate by week 2. Cell number increase initiation shows first divisions doubling initial cell number, second round divisions quadrupling number, exponential growth pattern establishing from repeated division rounds, and observable cell number increase beginning week 1-2. Observable mitotic figures show condensed chromosomes visible under microscopy during metaphase, spindle structures evident during anaphase, dividing cells identifiable through cytological examination, and mitotic index increasing as division onset proceeds. Cell cycle times at onset show initial cycles 14-16 hours during first divisions, progressive acceleration to 12-14 hours as activity increases, and eventual peak rates 10-12 hours during maximum activity. Daughter cell characteristics show cells initially similar size to parent approximately 20-30 micrometers, cells entering G1 capable of growth and subsequent division, and daughter cells contributing to proliferating population. Spatial division patterns show earliest divisions in mesocarp regions near seeds, progressive expansion of dividing zone throughout mesocarp, exocarp surface cells showing coordinated division maintaining coverage, and spatial pattern reflecting auxin gradient and tissue organization. Environmental effects on mitotic onset include temperature critically affecting cell cycle progression rate with optimal 24-28°C supporting rapid cycle completion and division onset within 5-7 days, cool conditions 18-22°C slowing cycles delaying onset to 7-12 days, and temperature as major factor determining division timing. Nutrient requirements show adequate nitrogen for protein synthesis supporting cell cycle machinery, phosphorus for DNA synthesis and energy metabolism, and overall mineral nutrition supporting division though usually adequate in vineyard conditions. Division onset assessment methods include mitotic index calculation counting dividing cells per total cells, immunohistochemistry detecting mitotic marker proteins, BrdU or EdU pulse-labeling identifying S phase cells, and proliferation marker PCNA showing cycling cells. Failed or delayed onset causes include incomplete DNA replication preventing mitosis entry, checkpoint arrest from DNA damage or stress, inadequate metabolic support preventing cycle completion, or severe environmental stress disrupting division. The process represents second aspect of active berry cell division marking first divisions, following initial division activation and preceding periclinal cell division patterns. Duration represents division onset wave over days 5-10 post-fertilization. Regional timing shows mitotic activity onset during late June-early July across climates within first 1-2 weeks post-fertilization. Viticultural significance includes representing observable proliferation initiation marking berry as actively growing tissue, establishing exponential cell number increase pattern, demonstrating temperature sensitivity of division timing, and initiating cell accumulation determining final berry cellularity. Research applications include analyzing cell cycle regulation and division timing, investigating mitotic spindle function and chromosome segregation, evaluating environmental effects on division onset, quantifying early division rates and patterns, and assessing division onset as determinant of proliferation extent and berry size potential.
Mitotic_Activity_Onset is second aspect of Active_Berry_Cell_Division, distinguished by first divisions occurring establishing proliferative state. Represents S phase DNA replication, M phase mitosis, and cytokinesis producing daughter cells. Timing days 5-10 post-fertilization. Initial cell cycle times 14-16 hours accelerating to 12-14 hours. Exponential cell number increase initiating. Temperature affects division timing. Follows Initial_Division_Activation. Precedes pattern-specific Periclinal_Cell_Division. Observable proliferation marker.
Proliferations-Initiierung
Teilungsbeginn
division commencement
erste Mitose
first cell division
first mitosis
mitotische Aktivitäts-Beginn
proliferation initiation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 433-435 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Mitotic Activity Onset
A mixed active bud at node 6 on Vitis vinifera cv. Syrah in late May in the Rhône Valley, France, developing into a shoot with 12 fully expanded leaves totaling 850 cm² leaf area, 1 inflorescence with 125 berries at early fruit development stage, shoot length 45cm with 8 visible internodes, demonstrating balanced growth with leaf area-to-fruit ratio of 68 cm²/berry providing adequate photosynthetic support for fruit ripening while maintaining sufficient vigor for cane maturation and bud differentiation for subsequent season.
An active bud developing into a shoot bearing both substantial vegetative structures and reproductive structures, representing balanced allocation between leaf production for photosynthetic capacity and inflorescence production for reproductive success, typical of primary buds from mid-cane node positions under optimal vine balance conditions. Mixed active buds produce shoots with favorable leaf-to-fruit ratios (typically 10-15 cm² leaf area per gram fruit fresh weight at harvest), adequate photosynthetic capacity to support both shoot growth and fruit development, and balanced hormone profiles avoiding excessive vegetative vigor or reproductive stress, representing the viticultural ideal for sustainable production combining adequate current crop with sufficient reserve accumulation for following season fruitfulness.
Mixed active buds represent optimal vine balance, avoiding the extremes of purely vegetative shoots (no crop contribution) or excessively reproductive shoots (inadequate leaf area, poor fruit quality, depleted reserves). Achieving predominantly mixed buds across a vineyard requires integrated management of pruning severity, crop load adjustment, canopy management, irrigation, and nutrition to balance vegetative and reproductive development. The ratio of vegetative to reproductive components in mixed buds varies with node position, cultivar, environmental conditions, and vine age, with experienced viticulturists using visual assessment of mixed bud shoot development to gauge vine balance and adjust management accordingly.
ausgewogene Knospe
balanced active bud
gemischte aktive Knospe
vegetative-reproductive bud
Based on Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Howell (2001) "Sustainable grape productivity and the growth-yield relationship= A review" American Journal of Enology and Viticulture 52=165-174.
Mixed Active Bud
Moderate deficiency quality diagnosis for magnesium in Mosel Riesling guided intensified management intervention for a vineyard showing established Mg limitation with current-season quality implications. Visual assessment revealed moderate deficiency quality with obvious interveinal chlorosis (score 2.8 on 0-5 scale) readily apparent without reference comparison, symptoms distributed from basal leaves (score 3.5) through mid-canopy (score 2.5) with young apical leaves beginning to show symptoms (score 1.5), indicating deficiency progression beyond the earliest stage. Chlorosis color showed moderate deficiency quality with distinct yellow interveinal tissue contrasting against green veins, the pattern unmistakably diagnostic for Mg limitation when combined with the basipetal-to-acropetal distribution characteristic of mobile nutrient deficiency. Tissue analysis confirmed moderate deficiency quality with petiole Mg at 0.18% (clearly deficient, threshold 0.30%) and K:Mg ratio of 14:1 (imbalanced, optimal below 8:1), establishing both absolute deficiency and K-induced aggravation as contributing factors. Functional assessment revealed moderate deficiency quality with photosynthetic rate reduced 25% in affected leaves, cluster rachis showing early grape stem necrosis symptoms in 18% of clusters, and fruit Brix accumulation rate 12% below normal indicating that fruit quality impact was occurring during the current season. The moderate deficiency quality diagnosis guided enhanced management response including immediate foliar MgSO₄ application at 2% concentration for rapid symptom mitigation (two applications at 10-day interval), soil-applied kieserite at 150 kg/ha for sustained supply, and recommendation to reduce potassium fertilization in following seasons to address the K:Mg imbalance. Current-season outcome showed partial symptom improvement but 15% yield reduction from grape stem necrosis that had already occurred before intervention, demonstrating the limited capacity to fully recover from moderate deficiency once damage has been incurred.
Moderate deficiency quality is a deficiency severity quality that inheres in a grapevine organism and represents a nutritional limitation of intermediate intensity where tissue nutrient concentrations have declined to clearly deficient levels, with visible symptoms obvious without comparison to reference vines and extending beyond the oldest leaves to affect substantial portions of the canopy, and with functional impacts including measurable growth reduction, reduced photosynthetic capacity, and effects on fruit development that will cause productivity losses if uncorrected. This quality captures the stage of established deficiency requiring active management intervention, recognizing that moderate deficiency represents significant nutritional stress that is causing current-season productivity impacts and will worsen without corrective action. The ontological significance of moderate deficiency quality lies in its indication that corrective action is urgently needed and should proceed at enhanced intensity compared to mild deficiency, potentially including supplemental foliar application in addition to soil correction, and that current-season yield and quality impacts are likely despite corrective efforts due to damage already incurred. The biological basis of moderate deficiency encompasses tissue concentrations in the mid-deficient range where multiple metabolic processes are substantially impaired, stress responses fully activated, and visible damage reflecting significant cellular dysfunction. The visual symptoms of moderate deficiency show clearly recognizable expression of the characteristic deficiency pattern (chlorosis, necrosis, growth reduction, etc.) distributed across multiple canopy positions rather than limited to the oldest leaves, with symptom intensity scores typically in the middle range (2-3 on 0-5 scales) indicating obvious but not extreme expression. The management response to moderate deficiency typically involves enhanced corrective fertilization at above-normal rates, often combining soil application for sustained supply with foliar application for rapid uptake, and may include multiple applications through the season to address both current symptoms and prevent further decline.
Moderate_Deficiency_Quality is_a Deficiency_Severity_Quality, representing established deficiency with obvious symptoms and significant functional impact. This class addresses the intermediate severity category requiring active intervention. Symptoms obvious without reference comparison, distributed beyond oldest leaves. Tissue concentrations in mid-deficient range. Current-season productivity impacts likely despite correction. Enhanced fertilization rates and combined soil + foliar application typically needed. Partial rather than full recovery common due to damage already incurred.
Mäßiger Mangel
developed deficiency
entwickelter Mangel
established nutritional limitation
etablierte Nährstofflimitation
intermediate deficiency
significant deficiency
signifikanter Mangel
Moderate Deficiency Quality
SO4 (V. berlandieri × V. riparia) provides moderate drought tolerance to Riesling in the Rheingau, maintaining productivity with supplemental irrigation of 50-100 mm during dry summers but showing stress symptoms below 400 mm total water supply. 3309 Couderc (V. riparia × V. rupestris) demonstrates moderate drought tolerance supporting Pinot Noir in the southern Burgundy Côte Chalonnaise, adequate for typical years but requiring deficit irrigation during exceptional drought such as 2003 and 2022. Kober 5BB (V. berlandieri × V. riparia) shows moderate drought tolerance in Austrian Niederösterreich, sustaining Grüner Veltliner productivity in average rainfall years (500-600 mm) but with yield reductions of 30-40% in severe drought years compared to 10-20% reductions on 110 Richter.
A drought-tolerant rootstock role that inheres in a grafted rootstock by virtue of its capacity to maintain scion productivity under moderate water deficit conditions, performing substantially better than drought-sensitive genotypes but with reduced performance relative to high drought-tolerance rootstocks under severe or prolonged water stress. Moderate drought-tolerance rootstocks sustain economically viable yields under rainfall regimes of 400-600 mm annually with summer drought periods of 2-3 months, soil water potentials declining to -1.0 to -1.5 MPa in the primary rooting zone, and moderate vapor pressure deficits. This intermediate drought adaptation reflects moderate root system depth (2-3 m typical penetration), intermediate root-to-shoot ratios (0.25-0.4), and adequate but not exceptional osmotic adjustment and cavitation resistance. Moderate drought-tolerant rootstocks represent a compromise between drought adaptation and other desirable traits such as vigor control, lime tolerance, or nematode resistance that may not be optimally combined with extreme drought tolerance.
Moderate_Drought_Tolerance_Rootstock_Role captures the large group of commercially important rootstocks that perform adequately under typical rainfall variability in sub-humid to semi-arid climates but are not selected primarily for drought adaptation. Many of these rootstocks derive from V. berlandieri × V. riparia crosses (SO4, 5BB, 5C, 125AA) where the V. riparia parent contributes traits (lime tolerance, phylloxera resistance, moderate vigor) that partially offset the high drought tolerance of pure V. berlandieri × V. rupestris combinations. This category is particularly relevant for climate change adaptation planning, as moderate-tolerance rootstocks may require irrigation infrastructure upgrades in regions where they were previously dry-farmed successfully.
Unterlage mit mittlerer Trockenheitstoleranz
intermediate drought tolerance rootstock
moderate drought tolerance rootstock
moderately drought tolerant rootstock
mäßig trockenheitstolerante Unterlage
Serra I et al. (2014) The interaction between rootstocks and cultivars to enhance drought tolerance in grapevine, Australian Journal of Grape and Wine Research 20: 1-14 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data, Annals of Botany 105: 661-676 Lovisolo C et al. (2010) Grapevine adaptations to water stress: new perspectives about soil/plant interactions, Theoretical and Experimental Plant Physiology 22: 1-15
Moderate Drought Tolerance Rootstock Role
110 Richter (V. berlandieri × V. rupestris) demonstrates moderate lime tolerance (17-20% active lime), supporting Tempranillo across the calcareous mesetas of Ribera del Duero and Rioja (active lime 12-18%) without chlorosis in typical soil moisture conditions. SO4 (V. berlandieri × V. riparia) tolerates 17-20% active lime, serving as the dominant rootstock in German viticulture on the calcareous slopes of the Rheingau, Pfalz, and Rheinhessen. Kober 5BB (V. berlandieri × V. riparia) shows similar moderate tolerance (20% active lime), widely planted on the loess and limestone-influenced soils of Austrian Niederösterreich for Grüner Veltliner and Riesling.
A lime-tolerant rootstock role that inheres in a grafted rootstock by virtue of its capacity to maintain scion productivity on moderately calcareous soils with active limestone content of 15-25%, performing substantially better than lime-sensitive rootstocks but with reduced tolerance relative to high lime-tolerance genotypes on strongly calcareous substrates. Moderate lime-tolerance rootstocks sustain iron nutrition at active CaCO₃ concentrations of 15-25%, soil pH of 7.5-8.0, and moderate rhizosphere bicarbonate stress, maintaining adequate chlorophyll synthesis and photosynthetic capacity under these conditions. This intermediate tolerance reflects moderate Fe³⁺-chelate reductase efficiency, partial but not exceptional proton efflusion capacity, and iron use efficiency adequate for moderate but not severe calcareous stress. Moderate lime-tolerance rootstocks represent the largest commercial category, suitable for the extensive calcareous soils of Mediterranean and continental European viticulture.
Moderate_Lime_Tolerance_Rootstock_Role encompasses the commercially dominant V. berlandieri × V. riparia (SO4, 5BB, 5C, 125AA, 420A) and V. berlandieri × V. rupestris (110R, 99R, 1103P, 140Ru) rootstock groups. The V. berlandieri parent contributes lime tolerance while the V. riparia or V. rupestris parent contributes phylloxera resistance and other desirable traits. Moderate lime tolerance is adequate for the majority of calcareous viticultural soils worldwide high-tolerance rootstocks are reserved for extreme chalk and marl formations. Drought stress exacerbates lime chlorosis by reducing root iron uptake, creating interactions between lime and drought tolerance.
Unterlage mit mittlerer Kalkverträglichkeit
intermediate lime tolerance rootstock
moderate lime tolerance rootstock
moderately lime tolerant rootstock
mäßig kalktolerante Unterlage
Bavaresco L et al. (2003) Grapevine iron nutrition update, Journal International des Sciences de la Vigne et du Vin 37: 89-100 Tagliavini M, Rombolà AD (2001) Iron deficiency and chlorosis in orchard and vineyard ecosystems, European Journal of Agronomy 15: 71-92 Bauer KP, Götz G (2013) Unterlagen der Rebe, Bayerische Landesanstalt für Weinbau und Gartenbau
Moderate Lime Tolerance Rootstock Role
A moderate medium fruiting cane on Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, retained at 9 nodes spanning 98cm along fruiting wire, selected from vine showing 520g pruning weight in well-drained hillside vineyard, positioned for efficient shoot management in vertical canopy, expected to produce 8-9 fruiting shoots bearing 15-17 clusters totaling 4.1kg fruit, managed through selective cluster thinning to 12-13 clusters maintaining target 1.2kg per meter canopy density, achieving balanced 13.5 cm²/g leaf area-to-fruit ratio supporting complete fruit maturation to 25° Brix with full phenolic development, representing upper quality tier for commercial Cabernet production.
A medium fruiting cane at the longer end of the moderate range, typically 8-10 nodes, representing higher-yielding approach within quality-oriented medium cane category, characterized by substantial but not excessive length (85-110cm), moderate-to-high bud count balancing productivity and quality, suitability for reasonably vigorous sites with good fruit quality track record, and production of adequate shoot numbers for commercially viable crops without compromising premium wine potential. Moderate medium fruiting canes provide pragmatic compromise between economic necessity (adequate yields for financial viability) and quality objectives (maintaining fruit composition and wine character standards), appropriate for many commercial premium wine vineyards requiring sustainable business models combining quality reputation with profitable production levels.
Moderate medium fruiting canes represent practical upper limit for many premium wine producers, providing maximum yields consistent with quality standards and market positioning. The 8-10 node range suits vigorous cultivars like Cabernet Sauvignon, Syrah, or Nebbiolo in good sites, allows profitable production levels (typically 4-6 kg/vine translating to 6-9 tons/hectare depending on vine density), maintains excellent fruit quality through manageable crop loads and favorable canopy architecture, and remains compatible with estate winery economics requiring higher yields than ultra-premium boutique producers but lower yields than bulk wine operations. This category encompasses many successful wine regions producing critically acclaimed wines at commercially viable production levels.
balanced moderate cane
mäßig-mittlere Fruchtrute
productive medium cane
produktive mittellange Rute
upper medium cane
Based on Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Bravdo et al. (1985) "Effect of crop level and crop load on growth, yield, must and wine composition, and quality of Cabernet Sauvignon" American Journal of Enology and Viticulture 36=125-131 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115.
Moderate Medium Fruiting Cane
Pinot Noir (compact pistil clones) exhibits moderate medium styles 0.48-0.56 mm connecting ovaries 0.78-0.92 mm to stigmas 0.38-0.46 mm, with moderate medium styles representing 25-29% of total pistil length (1.9-2.15 mm), containing fully functional transmitting tract tissue supporting normal pollen tube growth and achieving 48-54% fruit set comparable to Pinot Noir clones with standard medium styles (0.56-0.64 mm) achieving 50-56% fruit set, demonstrating that moderate versus standard medium style length has minimal impact on reproductive success within the functional medium range. Riesling shows moderate medium styles 0.46-0.54 mm (averaging 0.5 mm) in certain vineyard sites with moderate growing conditions, while the same clones in more favorable sites show standard medium styles 0.54-0.62 mm (averaging 0.58 mm), suggesting environmental modulation of style length within the medium range with moderate conditions producing moderate medium styles and optimal conditions allowing fuller medium style expression. Chardonnay pollen tube growth comparisons between moderate medium styles (0.48-0.56 mm) and standard medium styles (0.58-0.66 mm) reveal pollen tube transit times of 48-68 hours versus 58-78 hours respectively at 22-24°C, with the shorter moderate medium styles providing slightly faster transit through reduced pathway length while both style lengths achieve similar ultimate fertilization success rates (50-56% versus 52-58% fruit set).
A moderate medium style is a medium style in the lower-to-middle portion of the medium length range, typically 0.45-0.58 mm at maturity, representing styles that are clearly within normal hermaphrodite dimensions but toward the shorter end of the medium spectrum, characteristic of some hermaphrodite cultivars or of hermaphrodite flowers developing under conditions producing compact pistil morphology. Moderate medium styles are characterized by= (1) length typically 0.45-0.58 mm at anthesis, exceeding short style range (<0.45 mm) but falling in the lower half of medium style range (0.45-0.7 mm) (2) diameter typically 0.18-0.28 mm proportional to moderate length (3) well-developed transmitting tract tissue with functional extracellular matrix and specialized cells supporting effective pollen tube growth (4) connection between ovary (typically 0.75-0.95 mm) and stigma (typically 0.35-0.48 mm) creating pistils (total length typically 1.8-2.2 mm) with moderate medium styles representing 22-30% of pistil length. Moderate medium styles develop in hermaphrodite flowers through normal pistil development processes producing style dimensions toward the compact end of the normal range, occurring in= (1) cultivars with genetically determined tendency toward compact pistil morphology including somewhat shorter styles within the functional medium range (2) flowers developing under environmental conditions (moderate resource availability, typical temperatures) producing pistil dimensions in the moderate rather than maximal medium range (3) pistils showing developmental variation creating shorter medium-range styles rather than longer medium-range styles within the normal hermaphrodite spectrum. Moderate medium styles represent fully functional styles supporting normal fertilization despite being toward the shorter end of medium range= transmitting tract development is complete with elaborate extracellular matrix and specialized cell morphology pollen tube growth proceeds normally requiring 45-75 hours for stigma-to-ovule transit (slightly faster than in longer styles due to reduced distance) fertilization success rates are comparable to longer medium styles with fruit set typically 42-56% under favorable conditions. The distinction between moderate medium and standard medium styles reflects quantitative variation within the medium category rather than qualitative functional differences, with both sub-types supporting successful hermaphrodite reproduction. Moderate medium styles may provide subtle advantages in certain contexts= slightly reduced pollen tube pathway length (total 1.4-2 mm versus 1.6-2.5 mm for longer medium or long styles) potentially reducing time for environmental perturbations to disrupt pollen tube growth more compact pistil morphology including moderate medium styles may be advantageous in dense flower clusters by reducing physical interference among pistils. Cultivar examples showing moderate medium styles include= some Pinot Noir clones consistently averaging 0.48-0.56 mm style length certain Riesling selections showing 0.46-0.54 mm styles while other clones of these same cultivars show standard medium or longer styles, demonstrating within-cultivar genetic variation in style length within the medium range.
Moderate_Medium_Style represents the lower-to-middle medium style range and is distinguished from Standard_Medium_Style (0.55-0.7 mm, upper medium range) by somewhat shorter dimensions while both remain within functional medium category. The moderate versus standard medium distinction reflects quantitative variation within the medium range rather than qualitative functional differences, with both supporting normal hermaphrodite reproduction. Understanding moderate versus standard medium styles is relevant for characterizing within-cultivar variation, for assessing environmental modulation of pistil dimensions, and for detailed pistil morphometric studies, though the distinction has limited practical significance for fruit set prediction given functional equivalence.
compact medium style
kompakter Normal-Stempelgriffel
kompakter Normalstylus
mäßig mittellanger Griffel
shorter medium style
Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-105 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 94-97.
Moderate Medium Style
A moderate primary vegetative shoot on Vitis vinifera cv. Pinot Noir from node 2 in well-balanced mature vineyard, developed to 118cm length by late July with growth largely ceased, 14 fully expanded leaves averaging 145 cm² (total approximately 2030 cm² leaf area), 8mm diameter at base showing early lignification initiation in basal internodes, 2 tendrils, well-positioned in vertical trellis without crowding adjacent shoots, representing ideal vegetative shoot contributing significant photosynthetic capacity (approximately 12% of whole-vine leaf area) while maintaining good canopy architecture, adequate spacing, and balanced resource allocation, retained through season for photosynthetic contribution then evaluated during dormant pruning as potential renewal cane candidate based on positioning and quality.
A primary vegetative shoot exhibiting balanced growth characteristics, characterized by steady moderate elongation (2-4cm per day during active growth), intermediate final length (100-150cm typical), normal internodes (6-10cm spacing), appropriate diameter (6-9mm at base), adequate leaf development (12-18 leaves, individual areas 120-180 cm²), and photosynthetic contribution proportional to shoot size without excessive vigor or resource consumption, representing well-balanced vegetative development appropriate for contributing to whole-vine function without creating canopy management problems or indicating problematic vine imbalance. Moderate primary vegetative shoots represent ideal vegetative expression, providing valuable photosynthetic capacity and potential renewal wood through balanced growth neither excessive (causing shading and management problems) nor deficient (indicating vine stress or inadequate vigor), typical of well-managed vineyards with appropriate vine balance, suitable crop loads, and judicious resource management.
Moderate primary vegetative shoots represent the target for vegetative shoot development in balanced vineyards, providing photosynthetic benefits without excessive vigor problems. These shoots contribute meaningfully to whole-vine carbon balance (typically 1500-2500 cm² leaf area per shoot), maintain good canopy architecture through moderate size and spacing, show appropriate developmental progression (timely growth cessation and lignification), and often provide high-quality renewal wood with balanced characteristics. Management typically retains moderate vegetative shoots when canopy space permits, recognizing their valuable contributions to vine function and potential future roles as fruiting wood. The moderate designation indicates appropriate balance rather than mediocrity, with these shoots often representing better-quality wood than excessively vigorous shoots that may show late growth, poor lignification, and reduced bud fruitfulness due to self-shading during differentiation.
ausgewogener Blatttrieb
balanced primary vegetative shoot
mäßig wüchsiger primärer vegetativer Trieb
normal-vigor non-fruiting shoot
Based on Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 113-145 Howell (2001) "Sustainable grape productivity and the growth-yield relationship= A review" American Journal of Enology and Viticulture 52=165-174 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide.
Moderate Primary Vegetative Shoot
SO4 (V. berlandieri × V. riparia) demonstrates moderate rooting (65-80%), combining V. riparia's improved rooting with V. berlandieri's difficult-rooting tendency, achieving commercially acceptable propagation rates under standard protocols. 110 Richter (V. berlandieri × V. rupestris) shows moderate rooting (60-75%), with V. rupestris partially compensating for V. berlandieri's poor rooting to enable economical nursery production. Kober 5BB (V. berlandieri × V. riparia) roots moderately well (70-80%), establishing the V. berlandieri × V. riparia group as the most readily propagated of the lime-tolerant hybrid combinations. 140 Ruggeri (V. berlandieri × V. rupestris) shows moderate-to-difficult rooting (55-70%), at the lower end of commercial acceptability.
A rooting capacity rootstock role that inheres in a grafted rootstock by virtue of its intermediate capacity for adventitious root initiation from hardwood cuttings, achieving rooting success rates of 50-80% under standard nursery propagation conditions. Moderate-rooting rootstocks develop adequate root primordia with somewhat variable timing and density, show acceptable cutting survival during callusing, and produce functional root systems suitable for nursery production without specialized treatments, though auxin application may improve consistency. This intermediate rooting capacity reflects mixed genetic contributions, typically combining easier-rooting species (V. rupestris, V. riparia) with more difficult-rooting V. berlandieri parentage. Moderate-rooting rootstocks represent a practical compromise between the desirable viticultural traits contributed by V. berlandieri (lime tolerance, drought tolerance) and the propagation efficiency of easier-rooting species. Most commercially important rootstocks fall within the moderate rooting category.
Moderate_Rooting_Rootstock_Role encompasses the majority of commercially important rootstocks, particularly the V. berlandieri × V. riparia and V. berlandieri × V. rupestris hybrid groups where crossing with easier-rooting species provided the propagation solution that made V. berlandieri's lime tolerance commercially accessible. The 50-80% range represents economically viable propagation without specialized intervention, though nursery practices (cutting selection, callusing conditions, auxin treatment) can shift genotypes within this range. Moderate-rooting rootstocks are generally available at reasonable cost with reliable supply, unlike difficult-rooting genotypes that may have periodic shortages.
Unterlage mit mittlerer Bewurzelungsfähigkeit
intermediate rooting capacity rootstock
moderate rooting rootstock
moderately easy rooting rootstock
mäßig bewurzelbare Unterlage
Hartmann HT et al. (2011) Hartmann and Kester's Plant Propagation 8th Ed, Pearson Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Whiting JR (2004) Grapevine rootstocks, in Viticulture Vol 1 Resources, Winetitles pp.: 167-188 Bauer KP, Götz G (2013) Unterlagen der Rebe, Bayerische Landesanstalt für Weinbau und Gartenbau
Moderate Rooting Rootstock Role
110 Richter (V. berlandieri × V. rupestris) demonstrates moderate salt tolerance supporting Tempranillo in the moderately saline irrigated vineyards of La Mancha, Spain (ECe 2-3 dS/m), maintaining leaf Cl⁻ at 0.4-0.6% where sensitive rootstocks exceed 1.0%. 140 Ruggeri shows moderate-to-high salt tolerance in the coastal vineyards of Sardinia, adequate for ECe up to 3.5 dS/m but with yield decline evident at higher salinity levels where Ramsey maintains productivity. Paulsen 1103 provides moderate salt tolerance to Negroamaro in Puglia under moderate salinity stress (ECe 2.5-3.0 dS/m), though its tolerance classification extends toward the high category under some assessment criteria.
A salt-tolerant rootstock role that inheres in a grafted rootstock by virtue of its capacity to maintain scion productivity under moderate salinity conditions, performing substantially better than salt-sensitive genotypes but with reduced tolerance relative to high salt-tolerance rootstocks under severe saline stress. Moderate salt-tolerance rootstocks sustain economically viable production at soil ECe values of 2-4 dS/m and irrigation ECw of 1-2 dS/m, maintaining leaf Cl⁻ concentrations below 0.5-0.7% and Na⁺ below 0.25% dry weight under these conditions. This intermediate tolerance reflects partial but incomplete ion exclusion capacity, moderate osmotic adjustment ability, and K⁺/Na⁺ homeostasis that is maintained at low-to-moderate salinity but degrades under severe salt loading. Moderate salt-tolerance rootstocks represent a compromise category, often combining useful but not exceptional salt tolerance with other desirable traits such as phylloxera resistance, lime tolerance, or vigor control that may be suboptimally expressed in the most salt-tolerant genotypes.
Moderate_Salt_Tolerance_Rootstock_Role encompasses a broad range of commercially important rootstocks that provide meaningful but not exceptional salinity protection. This category includes many V. berlandieri × V. rupestris crosses (110R, 99R, some 1103P selections) that show good but variable salt tolerance, as well as some V. berlandieri × V. riparia selections (161-49C) with better-than-average but not excellent tolerance. The moderate category is particularly relevant for regions experiencing gradual salinization from irrigation intensification, where salt tolerance requirements may increase over time, potentially necessitating eventual transition to high-tolerance rootstocks.
Unterlage mit mittlerer Salztoleranz
intermediate salt tolerance rootstock
moderate salt tolerance rootstock
moderately salt tolerant rootstock
mäßig salztolerante Unterlage
Walker RR et al. (2002) Rootstock effects on salt tolerance of irrigated field-grown grapevines, Australian Journal of Grape and Wine Research 8: 146-155 Downton WJS (1977) Influence of rootstocks on the accumulation of chloride, sodium and potassium in grapevines, Australian Journal of Agricultural Research 28: 879-889 Stevens RM et al. (2011) Grapevine nutrition and irrigation management during rootstock establishment, Australian Journal of Grape and Wine Research 17: 73-82
Moderate Salt Tolerance Rootstock Role
Riesling exhibits moderate sugar accumulation disposition, typically reaching 20-23°Brix at optimal flavor maturity in cool climate regions, producing wines of 11-13% alcohol with pronounced varietal character. Pinot Noir in Burgundy achieves 21-24°Brix at harvest, balancing sugar with the 5-7 g/L acidity required for age-worthy red wine. Sauvignon Blanc in Marlborough ripens to 21-23°Brix while retaining the high acidity and thiol precursor content defining regional style. Moderate-sugar cultivars often show self-limiting sugar accumulation, with photosynthetic rate declining or sugar transport saturating before extreme concentrations are reached. The moderate accumulation trait facilitates harvest timing flexibility, as sugar progression slows above 22-23°Brix in these genotypes.
A sugar accumulation disposition that is the inherited capacity of a Vitis vinifera plant to achieve intermediate berry sugar concentrations (typically 20-24°Brix) at harvest maturity, enabling production of balanced wines with moderate alcohol levels while achieving adequate flavor and phenolic development.
Moderate sugar accumulation disposition aligns with contemporary consumer preferences for wines with lower alcohol and fresher flavor profiles. The trait provides winemakers with a larger harvest window, as the relationship between sugar and quality parameters remains favorable over a longer period. Climate change adaptation strategies increasingly emphasize cultivars with moderate sugar accumulation to maintain balanced wine styles without extreme management intervention.
ausgeglichene Mostgewichte
balanced sugar capacity
medium sugar accumulation
mittlere Zuckereinlagerung
moderate Brix trait
moderate Zuckerakkumulation
Van Leeuwen C, Darriet P (2016) The impact of climate change on viticulture and wine quality. Journal of Wine Economics 11:150-167 Duchêne E et al. (2012) Deciphering the ability of different grapevine genotypes to accumulate sugar in berries. Australian Journal of Grape and Wine Research 18:319-328
Moderate Sugar Accumulation Disposition
Documentation of Moderate_Vigor_Disposition at the University of Bordeaux examined growth characteristics of cultivars classified as moderate-vigor in international viticulture. Reference cultivars 'Cabernet Sauvignon', 'Merlot', 'Chardonnay', and 'Sauvignon Blanc' were compared under standardized conditions across three Bordeaux appellations with varying soil types. Moderate_Vigor_Disposition cultivars showed consistent intermediate growth characteristics. 'Cabernet Sauvignon' exhibited typical Moderate_Vigor_Disposition with mean shoot length of 175 cm (range 155-195 cm across sites), internode length of 7.2 cm, individual leaf area of 175 cm², lateral production of 7.5 laterals per shoot with total lateral length of 85 cm, and pruning weight of 1.45 kg/vine. Growth cessation occurred in weeks 13-14 (late July/early August), coinciding with veraison and enabling timely shift to reproductive development. 'Merlot' showed similar moderate vigor (168 cm shoots, 1.55 kg/vine) with slightly more lateral development. 'Chardonnay' displayed Moderate_Vigor_Disposition with more upright growth habit (185 cm shoots, 6.8 cm internodes, 1.38 kg/vine). The Ravaz index for Moderate_Vigor_Disposition cultivars averaged 6.8 (within optimal 5-10 range), indicating appropriate vegetative-reproductive balance. Canopy management requirements were moderate—1-2 hedging passes maintained fruit zone conditions with VSP training, and standard shoot thinning (removing double shoots and weak shoots) achieved target shoot density of 14-18 shoots per meter of cordon. Training system flexibility was confirmed, with 'Cabernet Sauvignon' performing well on VSP, bilateral cordon, Guyot, and modified sprawl systems with appropriate vine spacing adjustments.
A Moderate_Vigor_Disposition is a Shoot_Vigor_Disposition that inheres in Vitis vinifera shoots and is realized through balanced vegetative growth producing shoots of intermediate length (typically 140-200 cm), moderate internodes (6-8 cm mean length), medium-sized leaves (150-200 cm² individual area), and manageable lateral shoot development, resulting in canopies that achieve favorable balance between vegetative and reproductive growth with standard management practices. This disposition represents the middle range of grapevine vigor, characteristic of cultivars that produce adequate canopy for optimal fruit maturation without excessive vegetative growth that would compromise fruit quality or require intensive canopy management. Moderate-vigor cultivars typically show timely growth cessation (around veraison or shortly after), allowing carbon allocation to shift toward fruit ripening rather than continued vegetative growth. The viticultural implications of Moderate_Vigor_Disposition are generally favorable—such cultivars are adaptable to standard training systems (VSP, bilateral cordon, cane pruning), tolerate a range of vine spacings, respond predictably to moderate irrigation and fertilization, and achieve optimal Ravaz indices (5-10) representing balanced vine performance. Moderate-vigor cultivars include many of the world's premium wine grape varieties, reflecting both historical selection for manageable growth characteristics and the quality advantages of balanced vines. These cultivars offer flexibility in site selection and management approach, performing well across a range of conditions without requiring the specialized approaches demanded by high or low vigor extremes.
Moderate_Vigor_Disposition is_a Shoot_Vigor_Disposition representing the middle vigor category. Typical characteristics include shoots 140-200 cm, internodes 6-8 cm, pruning weight 1.0-2.0 kg/vine (site-dependent), growth cessation around veraison, and Ravaz index 5-10. Examples include 'Cabernet Sauvignon', 'Merlot', 'Chardonnay', 'Sauvignon Blanc', 'Syrah'. Management is straightforward with standard training systems (VSP, bilateral cordon) and moderate canopy management intensity. Moderate vigor offers site and management flexibility, representing the practical ideal for many viticultural contexts.
Ausgewogenes-Wachstum
Mittlere-Wuchskraft
Mittlere-Wuchsstärke-Disposition
balanced growth
intermediate vigor
medium vigor
Kliewer WM, Dokoozlian NK (2005) Leaf area/crop weight ratios of grapevines: influence on fruit composition and wine quality, American Journal of Enology and Viticulture 56:170-181 Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles, Adelaide Jackson DI, Lombard PB (1993) Environmental and management practices affecting grape composition and wine quality—a review, American Journal of Enology and Viticulture 44:409-430
Moderate Vigor Disposition
SO4 (Selection Oppenheim 4, V. berlandieri × V. riparia) provides moderate vigor to Riesling in the Rheingau, with pruning weights of 0.4-0.6 kg/m and balanced Ravaz indices of 5-7, and is the most widely used rootstock in German viticulture. Kober 5BB (V. berlandieri × V. riparia) confers moderate-to-moderately-high vigor to Grüner Veltliner in Austrian Niederösterreich, maintaining pruning weights around 0.5-0.7 kg/m on loess soils. 3309 Couderc (V. riparia × V. rupestris) delivers moderate vigor to Chardonnay in Burgundy with pruning weights of 0.4-0.6 kg/m on limestone-clay substrates.
A vigor-modifying rootstock role that inheres in a grafted rootstock by virtue of its capacity to sustain scion vegetative growth at intermediate levels, broadly comparable to own-rooted Vitis vinifera performance under favorable conditions. Moderate-vigor rootstocks produce scion pruning weights typically in the range of 0.4-0.7 kg/m, shoot lengths of 1.0-2.0 m per season, and leaf area indices of 2.0-3.5. This balanced growth is achieved through moderate root system extent, intermediate hydraulic conductance, and proportional nutrient acquisition that supports adequate canopy development for full fruit ripening without excessive vegetative competition. Moderate-vigor rootstocks are the most widely planted category globally, suitable for a broad range of soil types and climatic conditions.
Moderate_Vigor_Rootstock_Role represents the reference category against which high-vigor and low-vigor classifications are calibrated. The boundary between moderate and high or low vigor is not sharply defined and depends on site conditions, scion cultivar, and regional benchmarks. Many of the most widely planted rootstocks globally (SO4, Kober 5BB, 3309C, 101-14 Mgt) fall within or near the moderate vigor range under typical conditions, though they may shift categories under extreme edaphic or climatic circumstances.
Unterlage mittlerer Wuchsstärke
balanced-vigor rootstock
intermediate rootstock
medium-vigor rootstock
mittelwüchsige Unterlage
moderate-vigor rootstock
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 285-310 Ollat N et al. (2003) Effects of rootstock genotype on vine vigor, OENO One 37(2): 69-76 Bauer KP, Götz G (2013) Unterlagen der Rebe, Bayerische Landesanstalt für Weinbau und Gartenbau, Veitshöchheim
Moderate Vigor Rootstock Role
Moderate water stress quality implementation in Napa Valley Cabernet Sauvignon employed post-véraison deficit irrigation targeting the quality-optimizing moderate stress zone during the critical ripening period. Target moderate water stress quality was achieved through irrigation cutoff at véraison followed by monitoring-guided minimal supplemental irrigation maintaining midday stem water potential between -1.0 and -1.15 MPa throughout ripening. The moderate stress regime produced characteristic physiological responses including complete shoot growth cessation by véraison plus 10 days, stomatal conductance reduced to 40% of pre-stress values, and photosynthesis operating at 60% of potential capacity. Fruit composition demonstrated the quality benefits of moderate water stress quality with berry weight reduced 22% versus irrigated controls, anthocyanin concentration increased 45% reflecting both concentration effects and enhanced biosynthesis, tannin concentration elevated 35% with improved polymerization, and wine sensory scores significantly higher for color intensity, flavor concentration, and structural complexity. Yield reduction of 20% was acceptable for the premium market segment where quality premiums substantially exceeded volume-based revenue loss, validating moderate water stress quality as the appropriate management target for ultra-premium Cabernet Sauvignon production.
Moderate water stress quality is a water deficit severity quality that inheres in a grapevine organism and represents the intermediate category of water deficit characterized by pronounced stomatal closure substantially limiting both transpiration and photosynthetic carbon assimilation, complete or near-complete cessation of vegetative shoot growth, activation of stress-responsive metabolic pathways enhancing secondary compound biosynthesis, and measurable concentration effects on fruit composition that typically optimize wine grape quality for premium red wine production when timing and management are appropriate. This quality defines the target zone for quality-focused deficit irrigation strategies where the deliberate imposition of water stress achieves pronounced benefits through growth restriction, fruit concentration, and stress-induced metabolism while remaining within the reversible stress range that avoids permanent damage or catastrophic quality decline. The ontological significance of moderate water stress quality lies in its identification of the optimal stress intensity for premium wine grape production, representing the viticultural application of the hormesis principle wherein moderate stress exposure produces beneficial adaptations while severe stress causes damage, with the moderate category capturing the "sweet spot" where quality benefits are maximized relative to yield costs and stress risks. The biological basis of moderate water stress quality encompasses pronounced stomatal closure reducing conductance to 30-50% of unstressed levels, corresponding photosynthetic reduction to 50-70% of potential as CO₂ supply becomes limiting, complete cessation of primary shoot elongation through turgor-driven growth inhibition, and activation of stress-responsive gene expression including pathways controlling phenylpropanoid and flavonoid biosynthesis that enhance quality-relevant secondary metabolite accumulation in fruit. The concentration effects of moderate water stress operate through reduced berry size from limited cell expansion, increased skin-to-pulp ratio enhancing phenolic extraction, and reduced dilution of quality compounds in smaller berries with equivalent biosynthetic capacity. Physiological indicators of moderate water stress quality include predawn leaf water potential of -0.4 to -0.6 MPa and midday stem water potential of -1.0 to -1.2 MPa, with visible symptoms including slight midday leaf wilting recovering overnight, tendril desiccation, and early senescence of basal leaves providing field-observable confirmation of moderate stress status.
Moderate_Water_Stress_Quality is_a Water_Deficit_Severity_Quality, representing the intermediate stress intensity category with pronounced physiological response optimizing wine quality. This class addresses the primary target zone for quality-focused deficit irrigation. Typical predawn LWP -0.4 to -0.6 MPa, midday SWP -1.0 to -1.2 MPa provide thresholds. Distinguished from Mild_Water_Stress_Quality by growth cessation and visible symptoms, from Severe_Water_Stress_Quality by maintained reversibility and continued ripening function. Post-véraison timing is critical for quality benefits without yield compromise from early stress. Critical for premium wine production where quality premiums justify yield reduction.
beneficial water stress
intermediate drought stress
mittlerer Trockenstress
moderater Wasserstress
optimal water stress
optimaler Wasserstress
quality-enhancing water deficit
qualitätsförderndes Wasserdefizit
Ojeda H et al. (2002) Influence of pre- and postveraison water deficit on synthesis and concentration of skin phenolic compounds during berry growth of Vitis vinifera cv. Shiraz. American Journal of Enology and Viticulture 53:261-267 Castellarin SD et al. (2007) Water deficits accelerate ripening and induce changes in gene expression regulating flavonoid biosynthesis in grape berries. Planta 227:101-112 Koundouras S et al. (2006) Influence of vineyard location and vine water status on fruit maturation of nonirrigated cv. Agiorgitiko (Vitis vinifera L.). Effects on wine phenolic and aroma components. Journal of Agricultural and Food Chemistry 54:5077-5086
Moderate Water Stress Quality
A moderately vigorous water shoot on Vitis vinifera cv. Cabernet Franc in Loire Valley, France, emerging from an upper trunk latent bud at 95 cm height following partial canopy loss (approximately 65%) from late spring frost damage, achieving 275 cm total extension growth by September with 26 nodes, internodes averaging 14-16 cm in the mid-shoot region, leaves in the 220-250 cm² range showing normal dark green coloration and adequate cuticle development, stem diameter of 12 mm at node 13, showing good lignification in basal 15 nodes and appropriate growth cessation in early September allowing proper dormancy preparation, positioned favorably for training as a replacement cordon arm and exhibiting growth characteristics suitable for multi-season framework development without excessive vigor management requirements.
A vigorous water shoot exhibiting strong but not extreme growth characteristics in Vitis vinifera, characterized by daily elongation rates of 6-9 cm during active growth periods, internodes averaging 12-18 cm length, leaves typically in the 200-280 cm² range, stem diameters of 10-14 mm at mid-shoot positions, and total seasonal extension of 2-3.5 meters with 20-30 nodes under favorable conditions. Moderately vigorous water shoots represent typical strong compensatory growth responses to significant but not complete canopy loss, developing following 50-80% defoliation events, severe but not catastrophic pruning, or moderate trunk disease symptoms that trigger latent bud activation without creating the extreme resource imbalances that drive extremely vigorous growth. These shoots achieve substantial biomass production sufficient to partially restore photosynthetic capacity and carbohydrate production, but remain within manageable growth ranges that permit strategic decisions about retention for framework renewal versus removal for resource conservation. Moderately vigorous water shoots show better tissue maturation characteristics than extremely vigorous shoots, with adequate lignification, periderm development, and dormancy preparation under normal growing conditions, making them more suitable candidates for incorporation into permanent framework structures when strategic renewal is desired. Management flexibility with moderately vigorous water shoots permits topping interventions that reduce total growth while maintaining shoot viability, selective retention of shoots in optimal positions while removing others, and graduated training programs that develop shoots into productive framework over multiple seasons.
Moderately vigorous water shoots are distinguished from extremely vigorous water shoots by quantitative growth parameters falling in intermediate ranges, and by better tissue quality characteristics including appropriate lignification, normal leaf coloration, and timely growth cessation. These shoots represent the most favorable candidates for framework renewal strategies, combining sufficient vigor for rapid structural development with manageable growth characteristics that don't require extensive corrective interventions. Management decisions should consider shoot position relative to desired framework architecture, underlying vine health and disease status, and long-term vineyard production goals. Moderately vigorous water shoots can often be converted to productive framework through 2-3 seasons of managed crop loading and training, whereas extremely vigorous shoots may require 4-5 seasons or prove unsuitable despite management efforts.
controlled vigor water shoot
handhabbarer Stammtrieb
kontrolliert-vigoroser Wassertrieb
manageable epicormic shoot
moderate vigor sucker
mäßig kräftiger Wasserschoss
Based on Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 109-134 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles pp. 164-182 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 416-438.
Moderately Vigorous Water Shoot
The modified E-L Australian system used in AWRI extension materials provides cultivar-specific photographic reference plates showing the visual appearance of stages E-L 4 through E-L 38 as expressed in Shiraz, Cabernet Sauvignon, Chardonnay, Semillon, and Riesling under typical Australian growing conditions — the photographic guides illustrate how stage E-L 35 (véraison) manifests differently in tight-clustered Pinot Noir (where color change is initially obscured by berry-to-berry contact) versus loose-clustered Grenache (where individual berry color transitions are clearly visible), reducing inter-observer variability from 3-5 days to 1-2 days in comparative phenological monitoring networks such as the National Vintage Report coordinated by Wine Australia.
A modified E-L code representing the Australian adaptation of the Eichhorn-Lorenz staging system as refined by Coombe (1995) and further developed by the Australian Wine Research Institute and state departments of primary industries, incorporating supplementary intermediate stages, cultivar-specific photographic reference guides for major Australian plantings, and observational criteria calibrated to Southern Hemisphere growing season timing and warm-climate viticultural conditions.
Modified_EL_Australian represents the most widely used and thoroughly documented regional adaptation of the E-L system. Coombe's 1995 modification of the original Eichhorn and Lorenz (1977) system was itself an Australian initiative that added descriptive detail, introduced supplementary stages, and provided cross-reference tables to both Baggiolini and BBCH systems: subsequent development by the Australian Wine Research Institute, state viticultural extension services, and university viticulture programs has produced an extensive body of cultivar-specific reference materials, digital phenology recording tools, and training resources that constitute the de facto Australian national standard for grapevine phenological observation. The system's influence extends to New Zealand, South African, and South American viticultural practice through institutional collaboration and shared extension resources.
AWRI modified E-L
AWRI-E-L-System
Australian E-L system
Coombe scale
Coombe-Skala
australischer modifizierter E-L-Code
Coombe BG (1995) Growth stages of the grapevine: adoption of a system for identifying grapevine growth stages, Australian Journal of Grape and Wine Research 1: 104-110 Dry PR et al. (2004) Viticulture Volume 1: Resources, 2nd Ed, Winetitles pp: 141-167 AWRI (2023) Grapevine Growth Stages, Australian Wine Research Institute technical resources
Modified EL Australian
The modified E-L California system as presented in UC Davis viticulture extension materials provides integrated phenological staging and management decision guidelines specific to California's Central Coast, North Coast, and Central Valley growing conditions — for example, the E-L 15-17 window (10-12 leaves separated, inflorescences developing) is linked to specific irrigation initiation protocols in deficit-irrigated Cabernet Sauvignon vineyards on the Central Coast where pre-bloom water stress management requires phenological precision unavailable from calendar-based scheduling, and E-L 35 (véraison) triggers region-specific irrigation cutback protocols calibrated to the water-holding capacity of typical Paso Robles calcareous soils versus Napa Valley alluvial soils.
A modified E-L code representing the Californian adaptation of the Eichhorn-Lorenz staging system as developed by University of California viticulture extension programs, incorporating adjustments to stage boundary definitions and observational criteria that reflect the Mediterranean climate conditions, diverse cultivar range, and intensive precision viticulture practices characteristic of California's major wine-producing regions.
Modified_EL_California reflects the adaptation of the E-L framework to the specific conditions of California viticulture — the world's fourth-largest wine-producing region. The Californian modification emphasizes the integration of phenological staging with precision viticulture decision support systems, linking specific E-L stages to quantitative management thresholds for irrigation scheduling, nutrient application timing, and pest management windows: this integration is driven by the high capital intensity of California viticulture and the availability of extensive research data from UC Davis and UC Cooperative Extension linking phenological timing to management outcomes. The Californian system also addresses staging challenges specific to warm-climate viticulture where compressed phenological intervals (e.g., only 35-40 days from E-L 23 to E-L 35 in warm inland regions versus 50-60 days in cool coastal sites) require more frequent observation and tighter management response times.
California E-L system
UC Davis modified E-L
UC extension E-L scale
UC-Davis-E-L-System
kalifornische E-L-Anpassung
kalifornischer modifizierter E-L-Code
Hellman EW (2003) Grapevine structure and function, in Hellman EW (ed) Oregon Viticulture, Oregon State University Press pp: 5-19 Winkler AJ et al. (1974) General Viticulture, 2nd Ed, University of California Press UC Davis Department of Viticulture and Enology extension materials
Modified EL California
The modified E-L system used by the Australian Wine Research Institute introduces supplementary half-stages (e.g., E-L 31.5 for early bunch closure, intermediate between pea size and full bunch closure) and provides cultivar-specific photographic reference guides for Shiraz, Chardonnay, and Cabernet Sauvignon that illustrate the visual appearance of each E-L stage as expressed in these specific cultivars under Australian growing conditions — these modifications address the practical challenge that stage appearance can vary significantly between cultivars and climates, with a Shiraz cluster at E-L 31 looking substantially different from a Riesling cluster at the same developmental stage.
A phenological code based on a regional modification of the Eichhorn-Lorenz staging system, adapted to accommodate specific viticultural conditions, cultivar behaviors, management practices, or research requirements of a particular wine-producing region, retaining the core E-L numerical framework while introducing supplementary stages, refined stage boundary definitions, or region-specific observational criteria.
Modified_EL_Code captures the regional diversification of the E-L system that has occurred since Coombe's 1995 modification of the original Eichhorn and Lorenz (1977) framework. Regional modifications arise from the tension between the need for a universal staging system (enabling cross-regional comparison) and the reality that grapevine phenological expression varies significantly with cultivar, climate, and management system: the same developmental event may present with different visual appearances in different viticultural contexts, motivating the creation of region-specific reference materials and supplementary staging criteria. Subclasses capture the two most widely documented regional modifications — the Australian and Californian adaptations — though informal regional modifications exist in most wine-producing countries.
adapted E-L code
angepasster Eichhorn-Lorenz-Code
customized Eichhorn-Lorenz system
modifizierter E-L-Code
regional E-L modification
regionale E-L-Modifikation
Coombe BG (1995) Growth stages of the grapevine: adoption of a system for identifying grapevine growth stages, Australian Journal of Grape and Wine Research 1: 104-110 Dry PR et al. (2004) Viticulture Volume 1: Resources, 2nd Ed, Winetitles Hellman EW (2003) Grapevine structure and function, in Hellman EW (ed) Oregon Viticulture, Oregon State University Press pp: 5-19
Modified EL Code
A modified gobelet head arm on Vitis vinifera cv. Syrah in Côtes du Rhône, one of five arms (versus traditional four) emerging from head at 55cm height (versus traditional 35-45cm), extending 48cm outward (versus traditional 25-35cm), bearing 6 two-bud spurs (versus traditional 3-4), total vine with five arms bearing 30 combined spurs supporting approximately 54 shoots and 5.8kg crop (approximately 52 hl/ha at 1.8m × 1.8m spacing), representing moderate adaptation of gobelet maintaining recognizable traditional architecture while achieving yields 60-70% higher than strict traditional approach, enabling economic viability in commercial operation while preserving gobelet cultural heritage and market identity as traditionally-trained wine.
A gobelet head arm adapted from strict traditional form through moderate increases in arm length, spur retention, or vine capacity while maintaining basic gobelet architecture, characterized by slightly extended arms (40-60cm), increased spur counts (5-7 per arm), additional arms (5-6 versus traditional 3-4), or elevated head height (50-70cm versus traditional 30-50cm), representing pragmatic compromise between traditional gobelet heritage and economic necessity for improved productivity. Modified gobelet head arms reflect modern adaptations of traditional training, employed by producers seeking to maintain gobelet cultural identity and market positioning while achieving economically viable yields, appellation systems allowing moderate modernization within traditional frameworks, or transitional approaches in regions evolving from strict tradition toward more productive contemporary practices.
Modified gobelet head arms represent pragmatic evolution of traditional training, acknowledging economic realities requiring improved productivity while respecting cultural heritage and market expectations for traditional practices. The modifications typically increase yields from traditional 25-35 hl/ha to more viable 45-60 hl/ha range through moderate capacity increases (more or longer arms, more spurs) while maintaining recognizable gobelet architecture and avoiding wholesale conversion to modern cordon systems. Appellation authorities increasingly accept moderate modifications recognizing that strict traditional approaches threaten economic sustainability, with regulatory evolution allowing contemporary adaptations within traditional frameworks. Critics argue modifications dilute traditional character and undermine terroir expression justifying premium pricing, while defenders contend that reasonable adaptations preserve heritage while ensuring economic survival enabling continued traditional viticulture rather than abandonment and replanting to intensive modern systems.
adapted gobelet arm
angepasste Becherform-Arm
contemporary gobelet arm
enhanced traditional arm
modifizierte Gobelet-Arm
Based on Reynolds & Vanden Heuvel (2009) "Influence of grapevine training systems on vine growth and fruit composition= a review" American Journal of Enology and Viticulture 60=251-268 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide.
Modified Gobelet Head Arm
Moisture content quality tracking in Pedro Ximénez production for sweet sherry follows the sun-drying (soleo) process that reduces berry moisture from initial 78-82% to target 40-50% over 7-14 days of mat exposure, concentrating sugars from 22-26°Brix to 40-50°Brix the endpoint determination through moisture measurement (refractometer monitoring of sugar as proxy) ensures consistent must quality for the intensely sweet PX wines, with insufficient drying producing lower sugar concentration while excessive drying creates processing difficulties from the resulting syrup-like must viscosity.
Moisture_Content_Quality represents the chemical quality characterizing the gravimetrically determined water fraction in grape berry tissue, expressing water content as percentage of total mass or on dry weight basis. Moisture content provides the fundamental quantification of berry hydration status, determined by measuring mass loss upon drying (typically oven-drying at 60-105°C to constant weight, or freeze-drying for samples requiring compositional preservation). Fresh grape berries typically contain 75-85% moisture on a fresh weight basis, equivalent to 300-570% moisture on a dry weight basis, with the wide range reflecting cultivar, ripening stage, and environmental conditions. Moisture content decreases during late ripening as sugar accumulation increases dry matter fraction even without net water loss, and can decline substantially through post-harvest dehydration for dried grape products and passito-style wines. The relationship between moisture content and other quality parameters is primarily one of concentration effects as moisture declines, all non-volatile solutes (sugars, acids, phenolics, minerals) increase proportionally on a fresh weight basis while remaining constant on a dry weight basis. This concentration mechanism underlies quality changes during natural dehydration, noble rot infection, and deliberate drying processes. Moisture content also affects texture and processing behavior, with lower moisture producing firmer berries that resist mechanical damage but may require adjusted pressing parameters. For table grapes, moisture content maintenance through proper postharvest handling preserves freshness and marketable weight, with target moisture loss below 3-5% during storage and distribution. Moisture content measurement, while conceptually simple, requires attention to methodology (drying temperature, endpoint determination) for reproducible results.
Moisture_Content_Quality addresses the gravimetrically determined water fraction as the most direct water content measurement. The expression basis (fresh weight versus dry weight) affects numerical values substantially and must be specified for meaningful comparison. The drying methodology (temperature, duration, endpoint criterion) affects results, with lower temperatures preserving volatile compounds for subsequent analysis while higher temperatures speed determination. Moisture content provides the foundation for calculating dry matter content and enables conversion among concentration bases.
Feuchtigkeitsgehalt
Feuchtigkeitsgrad
Wasseranteil
gravimetric water content
gravimetrischer Wassergehalt
moisture level
percent moisture quality
water percentage
AOAC International (2019) Official Methods of Analysis, 21st Ed, Method 934.06 Moisture in Dried Fruits Rolle L et al. (2011) Instrumental Texture Analysis Parameters as Markers of Table-Grape and Winegrape Quality: A Review. American Journal of Enology and Viticulture 63(1): 11-28
Moisture Content Quality
In Tempranillo vines in central Spain during a dry winter, moisture limitation contributed to delayed budbreak in early April despite adequate heat accumulation, with soil water content declining to 15% of field capacity by late March due to below-average winter precipitation, pre-dawn leaf water potential of -0.5 MPa indicating mild water stress, and irrigation application required to relieve moisture constraint and enable normal budbreak progression.
Moisture limitation in Vitis vinifera represents a secondary environmental constraint that can contribute to ecodormancy maintenance when soil water availability is insufficient to support the cellular expansion and metabolic activity associated with growth initiation, though this factor is typically less important than temperature limitation in most viticultural environments during the late winter and early spring period of ecodormancy. The physiological basis of moisture limitation relates to the fundamental requirement for water in cellular expansion, with cell enlargement depending on turgor pressure generated by water uptake, and inadequate water availability preventing the turgor development necessary for cell wall extension and growth. The metabolic dimension of moisture limitation involves the requirement for adequate water status to maintain enzyme function, support transpiration-driven nutrient uptake, and enable the biochemical reactions of biosynthesis that depend on aqueous cellular environments. In most viticultural regions, moisture limitation during ecodormancy is uncommon because winter precipitation typically recharges soil water reserves, providing adequate moisture for early spring growth even in regions with summer drought stress, with dormant vines having minimal water requirements that are easily met by residual soil moisture. The situations where moisture limitation becomes significant during ecodormancy include regions with winter drought patterns, particularly Mediterranean climates with irregular precipitation, vineyards on sandy or shallow soils with low water holding capacity, and years with abnormally low winter precipitation that fails to adequately recharge soil water reserves. The interaction between temperature and moisture limitation can affect budbreak timing, with adequate moisture enabling rapid growth response when temperature constraint is released, while moisture deficit may delay or limit growth even when temperatures become favorable, though this interaction is rarely significant in well-managed vineyards with appropriate site selection. The detection of moisture limitation during ecodormancy involves monitoring soil water status and vine water potential, with pre-dawn water potential values more negative than -0.3 to -0.4 MPa potentially indicating moisture stress that could affect budbreak, though such values are uncommon during the typically wet late winter period in most grape-growing regions.
Moisture_Limitation represents a secondary environmental constraint within Environmental_Constraint addressing water-related growth limitation. Distinguished from Temperature_Limitation which is the primary ecodormancy constraint. Typically less important than temperature in most regions due to winter precipitation. Significant mainly in winter-drought regions, shallow soils, or abnormally dry years. Detected through soil moisture and vine water potential monitoring. Pre-dawn water potential more negative than -0.3 MPa indicates potential limitation.
Feuchtebeschränkung
Trockenheitsbeschränkung
Wasserdefizit-Beschränkung
Wasserlimitation
drought limitation
moisture constraint
water deficit constraint
water limitation
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 3 Water Relations and Nutrient Uptake Williams LE, Baeza P (2007) Relationships among ambient temperature and vapor pressure deficit and leaf and stem water potentials of fully irrigated, field-grown grapevines. American Journal of Enology and Viticulture 58: 173-181 Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant Cell and Environment 26: 1393-1405
Moisture Limitation
Molybdenum content quality rarely requires management in viticulture, but the acidic, heavily weathered soils of parts of the Granite Belt (Queensland, Australia) represent conditions where molybdenum deficiency can affect nitrogen metabolism in grapevines correction through soil liming (raising pH from 5.0 to 5.5-6.0) improves molybdenum availability while providing other benefits, demonstrating how molybdenum management typically involves pH adjustment rather than direct supplementation, in contrast to iron or zinc management requiring element-specific applications.
Molybdenum_Content_Quality represents the chemical quality characterizing the concentration of molybdenum (Mo) in grape berry tissue, capturing the essential micronutrient required in the smallest quantities among plant essential elements, functioning as cofactor in enzymes involved in nitrogen metabolism and hormone synthesis. Molybdenum in grape juice typically occurs at μg/L concentrations, well below the mg/L range of other micronutrients, reflecting both low plant requirements and the adequate availability in most agricultural soils. The critical functions of molybdenum involve its incorporation into the molybdenum cofactor (Moco) required by nitrate reductase (converting nitrate to nitrite in nitrogen assimilation), aldehyde oxidase (involved in abscisic acid synthesis), and xanthine dehydrogenase (purine catabolism). Molybdenum deficiency disrupts nitrogen metabolism by preventing nitrate reduction, producing symptoms resembling nitrogen deficiency including general chlorosis and reduced vigor. Because molybdenum availability increases with pH (opposite to most micronutrients), deficiency is primarily a problem in acidic soils (pH below 5.5) where molybdate adsorbs strongly to iron and aluminum oxides. Liming to raise pH typically corrects molybdenum availability without direct supplementation. The extremely low requirement and generally adequate soil supply make molybdenum deficiency uncommon in viticulture compared to iron, zinc, or boron deficiencies. In winemaking, molybdenum has no recognized quality implications and is not monitored in routine analysis. The interaction between molybdenum status and nitrogen metabolism creates indirect connections to must nitrogen content and fermentation through effects on vine nitrogen assimilation.
Molybdenum_Content_Quality addresses the micronutrient required in smallest quantities, with deficiency uncommon in viticulture. The pH-availability relationship of molybdenum (increasing with pH) is opposite to iron, zinc, and manganese, creating different soil conditions for deficiency. The nitrogen metabolism functions of molybdenum (nitrate reductase cofactor) link its status to nitrogen assimilation and potentially must nitrogen content. The extremely low concentrations (μg/L) require sensitive analytical methods (ICP-MS) for accurate quantification, beyond routine wine analysis capabilities.
Beerenmolybdänqualität
Mo content
Mo-Gehalt
Molybdängehalt
Molybdänkonzentration
berry molybdenum quality
molybdenum concentration
molybdenum level
Marschner H (2012) Marschner's Mineral Nutrition of Higher Plants, 3rd Ed, Academic Press pp.: 191-248 Kaiser BN et al. (2005) The Role of Molybdenum in Agricultural Plant Production. Annals of Botany 96(5): 745-754
Molybdenum Content Quality
The monitoring protocol for a Riesling vineyard in the Pfalz specifies weekly visual scouting from E-L 9 through harvest following a standardized transect design — the scout walks a diagonal transect across each 1-hectare management block, stopping at 20 evenly spaced assessment points to evaluate 5 vines per point (100 vines per block per visit), recording powdery mildew incidence (percentage of leaves with visible colonies) and severity (percentage of leaf area affected using the EPPO scale) on 4 randomly selected leaves per vine at positions 3-6 from the shoot base, downy mildew oil spot incidence and severity on the same leaves, and botrytis cluster incidence on 2 clusters per vine from E-L 33 onward. Concurrently, 2 delta pheromone traps per block (Pherocon-type with Lobesia botrana and Eupoecilia ambiguella lures) are checked weekly with moth counts recorded per trap per week to track flight activity and predict larval generation timing.
A pest management description that documents the systematic observation, sampling, and data collection procedures used to assess pest populations, disease incidence and severity, and beneficial organism activity in the vineyard, specifying the monitoring method (visual scouting, trapping, remote sensing), sampling design (frequency, sample size, spatial distribution), assessment scales, and data recording standards that provide the information base for threshold-based pest management decision-making.
Monitoring_Protocol provides the observational foundation upon which all threshold-based pest management decisions depend — without systematic monitoring, intervention decisions revert to calendar-based scheduling or reactive crisis management, both of which result in unnecessary pesticide applications or delayed response to developing pest problems. The design of vineyard monitoring protocols balances statistical rigor (sufficient sample size and spatial coverage to detect pest outbreaks with acceptable confidence) against practical feasibility (monitoring costs of 2-5 labor hours per hectare per visit must be economically justified by the management improvements they enable): the standard approach of 50-100 assessment vines per block per monitoring visit, assessed at weekly intervals during the growing season, provides approximately 80% probability of detecting a 5% change in disease incidence between visits — sufficient resolution for operational management decisions in most contexts. Predictive disease models (e.g., the VitiMeteo system for downy mildew, the UC Davis Powdery Mildew Risk Index) increasingly supplement field scouting by providing risk forecasts based on weather data that guide the timing and intensity of monitoring effort.
Befallserhebungs-Programm
Monitoring-Protokoll
Schädlingsüberwachungs-Spezifikation
disease monitoring specification
pest scouting protocol
vineyard surveillance program
EPPO (2023) EPPO Standards PP 1 — Efficacy evaluation of plant protection products Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests 2nd Ed, APS Press Pertot I et al. (2017) A critical review of plant protection tools for reducing pesticide use on grapevine, Crop Protection 97: 70-84 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 461-520
Monitoring Protocol
Monoterpene accumulation distinguishes aromatic from non-aromatic V. vinifera cultivars, with total monoterpene concentrations spanning three orders of magnitude from 6000 µg/L in highly aromatic Muscats. The MEP pathway provides isopentenyl diphosphate (IPP) and dimethylallyl diphosphate (DMAPP) precursors, with VvDXS (1-deoxy-D-xylulose-5-phosphate synthase) being the primary rate-limiting enzyme. Terpene synthases (VvTPS) convert geranyl diphosphate (GPP) to specific monoterpenes, with VvTPS gene family expansion in aromatic cultivars. Muscat of Alexandria shows 3000-6000 µg/L total monoterpenes, Gewürztraminer 1500-3000 µg/L, Riesling 500-1500 µg/L, and Chardonnay <200 µg/L. Sensory threshold for linalool (15-25 µg/L) means that even low-accumulating cultivars may show subtle floral notes.
An aromatic compound disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate monoterpenoid compounds (C10 isoprenoids) through the plastidial methylerythritol phosphate (MEP) pathway, with genotype-specific concentrations and profiles of linalool, geraniol, nerol, α-terpineol, and related compounds determining the intensity and character of floral-citrus aroma in Muscat and aromatic cultivars.
Monoterpene_Accumulation_Disposition serves as the parent class for specific monoterpene compound traits. The Muscat aroma character that defines this compound class is one of the most ancient and recognizable wine flavor attributes. A major QTL on chromosome 5 containing VvDXS explains much of the variation between aromatic and non-aromatic cultivars, with a SNP causing reduced enzyme activity in non-aromatic types. Both free and glycosylated monoterpenes accumulate during ripening, with the bound fraction increasing proportionally through harvest.
Monoterpen-Disposition
Monoterpenakkumulation
Terpenoidbildung
floral aroma disposition
monoterpenoid trait
terpene synthesis capacity
Battilana J et al. (2009) The 1-deoxy-D-xylulose 5-phosphate synthase gene co-localizes with a major QTL affecting monoterpene content in grapevine. Theoretical and Applied Genetics 118:653-669 Martin DM et al. (2010) Functional annotation, genome organization and phylogeny of the grapevine (Vitis vinifera) terpene synthase gene family based on genome assembly, FLcDNA cloning, and enzyme assays. BMC Plant Biology 10:226
Monoterpene Accumulation Disposition
In Orange Muscat berries in Rutherglen, Australia during BBCH 83-89, monoterpene synthesis generates total monoterpenes at rates reaching 0.28 mg/L per day during peak activity at 20-25 days post-veraison, with linalool synthase activity of 65 nmol/min/g fresh weight producing linalool accumulation to 3.5 mg/L total, geraniol synthase activity of 38 nmol/min/g generating geraniol to 2.2 mg/L, and alpha-terpineol formation reaching 1.1 mg/L through both direct synthesis and acid-catalyzed rearrangement of linalool under warm conditions averaging 26°C.
Monoterpene synthesis describes the enzymatic biosynthesis of C10 terpenoid compounds in Vitis vinifera berry cells during ripening, wherein terpene synthase enzymes convert geranyl diphosphate precursor generated through the plastidial methylerythritol phosphate pathway into the diverse array of monoterpene alcohols, hydrocarbons, and oxides that constitute the dominant aromatic compounds of aromatic grape varieties, including linalool, geraniol, nerol, citronellol, alpha-terpineol, and hotrienol contributing characteristic floral, citrus, and muscat aromatic character. This process involves the ionization of geranyl diphosphate by terpene synthase enzymes generating carbocation intermediates that undergo cyclization, hydride shifts, and water capture reactions to produce the structurally diverse monoterpene products characteristic of individual enzyme specificities, with the Vitis vinifera genome encoding approximately 70 terpene synthase genes of which 30-40 are expressed in berry tissues with variety-specific expression patterns determining the monoterpene profile of individual cultivars. The subcellular organization of monoterpene synthesis involves geranyl diphosphate formation in plastids through the MEP pathway enzymes, potential transport of this precursor to cytosolic compartments where many terpene synthases appear localized, and subsequent modification reactions including hydroxylation, oxidation, and isomerization that generate the complete spectrum of monoterpene structures found in ripe grape berries. The dominant monoterpene synthase activities in aromatic varieties include linalool synthase generating the floral compound contributing rose-like and lavender notes, geraniol synthase producing the rose-geranium scented alcohol, and alpha-terpineol synthase creating the lilac-like tertiary alcohol that contributes to overall floral complexity, with these enzymes showing 5-20 fold higher expression in Muscat compared to neutral varieties. Developmental regulation of monoterpene synthesis involves transcriptional activation of terpene synthase genes during the veraison transition, with expression levels increasing 3-10 fold during the post-veraison period in aromatic varieties, reaching maximum during mid-ripening and declining during late ripening as berries approach harvest maturity. Environmental factors influence monoterpene synthesis through light-mediated regulation of MEP pathway and terpene synthase gene expression, temperature effects on enzyme activities with optimal synthesis at 22-28°C, and potential stress-responsive induction of specific synthase genes under water deficit or other environmental challenges. The relationship between monoterpene synthesis rates and final accumulation levels reflects the balance between production and subsequent glycosylation that sequesters 80-95% of synthesized monoterpenes in bound precursor forms, with variety-specific glycosyltransferase expression determining the partitioning between free and bound pools.
Monoterpene_Synthesis is subclass of Terpene_Accumulation representing C10 terpenoid biosynthesis specifically. Distinguishes from Sesquiterpene_Production which generates C15 compounds. Sibling to Sesquiterpene_Production and Terpene_Pool_Development. Key enzymes include linalool synthase, geraniol synthase, alpha-terpineol synthase. MEP pathway provides GPP precursor. Variety-specific expression determines aromatic potential.
C10 terpene synthesis
C10-Terpen-Synthese
Monoterpen-Synthese
Monoterpenoid-Biosynthese
Terpensynthase-vermittelte-Monoterpen-Produktion
linalool and geraniol synthesis
monoterpenoid biosynthesis
terpene synthase-mediated monoterpene production
Martin DM et al (2010) Functional annotation of the grapevine terpene synthase gene family. BMC Plant Biology 10: 226 Lücker J et al (2004) Monoterpene biosynthesis in lemon. European Journal of Biochemistry 271: 2930-2938 Keller M (2020) The Science of Grapevines 3rd Ed pp 290-295
Monoterpene Synthesis
Morning harvest suitability quality evaluation in Marlborough compared Sauvignon Blanc wine outcomes across harvest timing treatments during a warm vintage with ambient temperatures reaching 28°C by early afternoon. Morning-harvested fruit (6:00-10:00 AM, berry temperature 12-16°C) demonstrated the quality benefits predicted by excellent morning harvest suitability quality, with juice oxidation indices 45% lower than afternoon fruit, preservation of characteristic thiols and methoxypyrazines at concentrations 25% higher, and resulting wines receiving significantly higher sensory scores for varietal typicity and freshness. Afternoon-harvested fruit (2:00-6:00 PM, berry temperature 24-28°C) showed accelerated oxidation during transport, reduced aromatic intensity, and lower overall quality scores despite equivalent fruit ripeness at harvest. Economic analysis established that the quality premium achievable through morning harvest justified the 35% reduction in daily picking capacity and associated labor cost increase, establishing morning-only harvest as the regional standard for premium Sauvignon Blanc production where morning harvest suitability quality benefits proved economically significant.
Morning harvest suitability quality is a harvest condition quality that inheres in a grapevine cultivar or clone and represents the compatibility of fruit characteristics with early morning harvest operations conducted during the cool hours between dawn and mid-morning, when ambient temperatures remain low, berry temperatures have equilibrated overnight to near-minimum daily values, and physiological processes including respiration and enzyme activity operate at reduced rates favorable for quality preservation during harvest and transport to the winery. This quality determines the benefits achievable through morning harvest scheduling and the cultivar-specific value of prioritizing early picking times that impose logistical constraints on operation scale and labor scheduling compared to flexible all-day harvest approaches. The ontological significance of morning harvest suitability quality lies in its role guiding harvest scheduling decisions that balance the quality benefits of cool-temperature picking against the operational constraints of compressed morning windows that limit daily harvest capacity and require early start times that may challenge labor availability and equipment positioning. The biological basis of morning harvest suitability encompasses the temperature-dependent rates of oxidative enzyme activity, respiratory metabolism, and microbial proliferation that accelerate fruit quality decline under warm conditions the influence of berry temperature on juice extraction characteristics, phenolic release, and aromatic compound volatilization during crushing and pressing and the stability of quality-determining compounds under the cool, low-oxygen conditions achievable when fruit enters the winery at temperatures below 15°C compared to the accelerated deterioration occurring when fruit arrives warm. Cultivar characteristics favoring morning harvest include high oxidation susceptibility where polyphenol oxidase activity at elevated temperatures rapidly degrades juice quality, aromatic compound volatility where temperature-dependent losses compromise varietal character, and thin skin fragility where turgor-related firmness at cool temperatures reduces mechanical damage compared to softer warm fruit. Assessment of morning harvest suitability quality involves controlled comparison of wine outcomes from morning versus afternoon-harvested fruit under equivalent ripeness conditions, quantification of temperature-dependent quality parameter changes, and cost-benefit analysis weighing quality improvements against operational constraints.
Morning_Harvest_Suitability_Quality is_a Harvest_Condition_Quality, representing the compatibility with and benefits from cool early morning harvest timing. This class addresses the diurnal timing dimension of harvest conditions, particularly relevant for oxidation-sensitive and aromatic cultivars. The temperature differential between morning and afternoon harvest can exceed 15°C with corresponding quality implications. Distinguished from Night_Harvest_Suitability_Quality which addresses after-dark operations with different logistical considerations. Most relevant for white and aromatic varieties in warm climates where temperature management during harvest critically influences quality.
Frühlese-Präferenz
Kühlernte-Tauglichkeit
Morgenernte-Kompatibilität
Morgenlese-Eignung
cool harvest suitability
cool-temperature harvest fitness
dawn harvest compatibility
early morning picking preference
Marais J (1998) Effect of grape temperature and yeast strain on Sauvignon Blanc wine aroma composition and quality. South African Journal of Enology and Viticulture 19:10-16 Coetzee C, du Toit WJ (2012) A comprehensive review on Sauvignon Blanc aroma with a focus on certain positive volatile thiols. Food Research International 45:287-298 Singleton VL et al. (1980) Grape and wine phenolics: background and prospects. In: Proceedings of the University of California, Davis, Grape and Wine Centennial Symposium, pp. 215-227
Morning Harvest Suitability Quality
Cabernet Sauvignon exhibits characteristic morphological qualities including small berry size (1.0-1.4 g fresh weight), thick skin relative to pulp volume, intense blue-black coloration with heavy bloom, and compact cylindrical cluster architecture with pronounced shoulders, distinguishing it morphologically from the larger-berried, loosely-clustered Merlot even when grown in adjacent vineyard blocks.
Morphological_Quality encompasses all qualities that pertain to the physical form, structure, and spatial organization of grapevine organs and tissues. This category captures the externally observable and measurable attributes that define the shape, size, color, texture, and structural integrity of plant parts from the subcellular to the whole-organ level. Morphological qualities are central to ampelography, the systematic description and identification of grapevine cultivars based on phenotypic characteristics. The category subdivides into five major domains: Size_Quality addressing dimensional attributes such as length, area, volume, and mass Shape_Quality capturing geometric form including overall shape, symmetry, and margin characteristics Color_Quality encompassing pigmentation and hue properties Texture_Quality addressing surface and internal consistency characteristics and Structural_Integrity_Quality capturing wholeness, cohesion, and damage states. Morphological qualities can be assessed through direct measurement, visual scoring, or instrumental analysis including image analysis systems for objective phenotyping. These qualities exhibit developmental plasticity, changing systematically through the growing season and across plant organs. Environmental factors including light exposure, temperature, water availability, and nutrient status modulate morphological quality expression, while genetic background establishes the range of possible variation. In viticultural practice, morphological qualities inform cultivar identification, clonal selection, harvest timing decisions, and grape sorting protocols. The standardized description of morphological qualities follows OIV (Organisation Internationale de la Vigne et du Vin) descriptor protocols that provide categorical scales for key ampelographic characters.
Morphological_Quality is positioned as a direct subclass of Grapevine_Quality. The category interfaces with but remains distinct from Grapevine_Morphological_Structure (BFO_0000040 material entity), which represents the physical structures themselves rather than their qualities. A leaf blade is a morphological structure its length, shape, and color are morphological qualities that inhere in that structure. This distinction follows BFO's separation of independent continuants (structures) from dependent continuants (qualities). OIV descriptors provide standardized categorical assessments that can be represented as quality value specifications.
Formqualität
Strukturqualität
ampelographic quality
ampelographische Qualität
morphologische Qualität
morphometric quality
physical form quality
structural quality
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris Galet P (1979) A Practical Ampelography, Cornell University Press pp.: 1-52 Chitwood DH and Sinha NR (2016) Evolutionary and Environmental Forces Sculpting Leaf Development. Current Biology 26(7): R297-R306
Morphological Quality
A Cabernet Sauvignon clone 337 vine maintained in a foundation vineyard at UC Davis Foundation Plant Services bears a Mother_Plant_Role, trained to bilateral cordons with regular renewal pruning to maximize budwood production (typically 50-100 buds per vine annually) while maintaining the health status verified through annual ELISA and PCR testing. A Riparia Gloire rootstock plant in an ENTAV increase block serves in a Mother_Plant_Role, producing 8-12 one-meter canes annually for hardwood cutting propagation. A virus-tested Chardonnay vine in a commercial nursery's scion block bears a Mother_Plant_Role at the Certified category, providing budwood for benchgraft production under certification scheme protocols.
A propagation role that inheres in a grapevine maintained specifically as a source of vegetative propagation material (cuttings, budwood, or explants) for multiplication, rather than for fruit production. Mother plants are cultivated under conditions optimizing the quantity and quality of propagation material—typically trained to maximize cane production rather than fruit yield, maintained in protected environments reducing pathogen exposure, and managed with intensive nutrition and irrigation supporting vigorous vegetative growth. Mother plant status implies responsibility for maintaining genetic identity (trueness-to-type) and health status (freedom from specified pathogens) that will be transmitted to all progeny plants through vegetative propagation. Mother plants may serve as rootstock sources (providing hardwood cuttings for rootstock production), scion sources (providing budwood for grafting), or both, with separate mother plant blocks typically maintained for rootstock and scion material to prevent accidental mixing. The certification category of the mother plant (Pre-Basic, Basic, Certified) determines the permissible uses of derived propagation material.
Mother_Plant_Role captures the fundamental source function in grapevine propagation, with subclasses distinguishing certification categories that determine health status requirements and multiplication permissions. Mother plant management differs fundamentally from production viticulture—fruit production is incidental or deliberately prevented (cluster removal) to maximize vegetative growth and propagation material quality. The health status of mother plants is critical because grapevine pathogens (viruses, phytoplasmas) are systemic and cannot be eliminated from infected plants through any commercial treatment once infected, a mother plant must be destroyed and replaced. Mother plant longevity varies from decades (for well-maintained foundation stock) to the typical commercial vineyard lifespan for Certified category material, with replacement triggered by declining vigor, pathogen detection, or variety obsolescence.
Mutterrebe
Mutterstock
Rebenquartier
mother vine
multiplication vine
source vine
stock plant
European Union (2020) Commission Implementing Directive 2020/177 Gramaje D, Armengol J (2011) Fungal trunk pathogens in the grapevine propagation process, Phytopathologia Mediterranea 50: S217-S226 Golino DA et al. (2017) Foundation Plant Services and clean plant programs, HortScience 52: 1190-1194 Walter B, Martelli GP (1996) Clonal and sanitary selection of the grapevine, in Sanitary Selection of the Grapevine, INRA Editions pp.: 43-95
Mother Plant Role
Mousse quality comparison in blind tasting of sparkling wines reveals quality differentiation through foam character, with prestige cuvée Champagne (36-month tirage) showing abundant, creamy, persistent mousse (quality rating 9/10) with fine texture maintained for minutes after pouring, standard Champagne (15-month tirage) displaying good mousse (7/10) with moderate persistence, and Prosecco showing frothier, quickly dissipating mousse (4/10) from limited protein content, demonstrating how mousse character enables quality tier identification before aromatic or gustatory assessment begins.
Mousse quality is an effervescence quality that represents the foam character formed when sparkling wine bubbles accumulate at the surface, characterizing the visual appearance, texture, and persistence of the foam crown that indicates wine quality, production method, and protein-polysaccharide composition affecting bubble stabilization. Within the BFO framework, this quality specifically addresses the foam dimension of effervescence distinct from individual bubble behavior, recognizing that mousse represents the integrated expression of bubble formation, stabilization, and accumulation into the foam structure that crowns quality sparkling wine in the glass. The ontological significance lies in mousse serving as a holistic quality indicator integrating multiple carbonation factors, where fine, persistent, creamy mousse demonstrates the combination of appropriate CO₂ level, optimal bubble size, and adequate surfactant protein concentration that together indicate sophisticated production and quality wine composition. The physical basis for mousse formation involves bubble accumulation at the wine surface where they are stabilized by surface-active proteins (particularly yeast-derived mannoproteins) that prevent coalescence and drainage, creating a foam structure whose density, fineness, and persistence reflect both carbonation level and protein-polysaccharide composition, with quality mousse showing the creamy, persistent character of well-stabilized fine bubbles. The production method associations of mousse quality are strong, with méthode traditionnelle extended aging releasing elevated mannoprotein levels that create the luxurious, persistent mousse characteristic of fine Champagne, while shorter aging or tank methods produce less stable, more ephemeral foam lacking the creamy quality associated with premium sparkling wines. The descriptive vocabulary for mousse includes density (sparse to abundant), texture (coarse to creamy), and persistence (fleeting to long-lasting), with quality mousse described as creamy, fine-textured, and persistent while poor mousse appears coarse, frothy, and quickly dissipating. Understanding mousse quality enables sparkling wine quality assessment based on foam character, production method and aging inference from mousse properties, and quality differentiation within sparkling wine categories.
Mousse_Quality addresses the foam dimension of effervescence as a holistic quality indicator integrating bubble size, persistence, and stabilization. This quality reflects both carbonation level and wine composition (particularly surfactant proteins), enabling quality assessment from foam character alone. The class supports sparkling wine evaluation and production method inference through mousse observation as the visible expression of bubble quality.
Kronencharakter
Mousse
Schaumkrone
Schaumqualität
crown character
foam head
foam quality
sparkling foam
Cilindre C et al. (2007) Evidence for the underlying proteins related to foam formation in Champagne base wines. Journal of Agricultural and Food Chemistry 55(5): 1595-1600 Vanrell G et al. (2007) Foam aptitude of the base wines in sparkling wine elaboration. Journal of Agricultural and Food Chemistry 55(3): 992-997
Mousse Quality
Mouthfeel quality evaluation in aged Barolo integrates multiple tactile dimensions into a coherent assessment, noting substantial body (8/10 weight), silky viscosity (smooth coating without oiliness), refined astringency (firm but polished tannin creating grip without harshness), appropriate alcohol integration (warmth present but not burning), and overall harmonious mouthfeel where structure supports the tar-rose-truffle flavor complexity, demonstrating how mouthfeel quality provides the textural framework that determines whether complex flavors are experienced as elegant or clumsy.
Mouthfeel quality is a tactile quality that represents the integrated somatosensory experience of wine in the oral cavity, encompassing body, texture, viscosity, astringency, and temperature sensations that together characterize the physical presence and tactile impression of wine during consumption. Within the BFO framework, this quality specifically addresses the in-mouth tactile dimension of wine perception, distinguishing the oral somatosensory gestalt from individual tactile components while recognizing that mouthfeel represents the holistic integration of multiple tactile sensations into a unified textural experience. The ontological significance lies in mouthfeel providing the structural backbone of wine quality perception, where the physical presence of wine creates the matrix within which flavors are experienced, with appropriate mouthfeel supporting and enhancing flavor perception while inappropriate mouthfeel (too thin, too harsh, too heavy) detracting from overall quality regardless of aromatic merit. The perceptual integration of mouthfeel involves simultaneous processing of multiple tactile inputs including weight (body), coating behavior (viscosity), surface interaction (smoothness or roughness), oral drying (astringency), temperature, and effervescence, with these components synthesized into the unified textural impression that characterizes wine's physical presence. The compositional determinants of mouthfeel include alcohol (contributing to body and warmth), glycerol and residual sugar (adding smoothness and viscosity), phenolic compounds (creating astringency and structure), polysaccharides (enhancing smoothness and body), dissolved CO₂ (producing effervescent texture), and the overall extract level (determining substance and weight). The wine style associations of mouthfeel are strong, with light-bodied fresh whites expected to show crisp, refreshing mouthfeel while full-bodied age-worthy reds should display rich, structured mouthfeel, making style-appropriate mouthfeel assessment essential rather than universal preference for particular textural profiles. Understanding mouthfeel quality enables comprehensive tactile assessment, style-appropriate evaluation of textural attributes, and quality judgment incorporating mouthfeel integration with flavor components.
Mouthfeel_Quality serves as the parent class for specific oral tactile qualities (Body, Viscosity, Smoothness, Roughness, Creaminess), representing the integrated tactile experience in the mouth. This quality is holistic, synthesizing multiple individual sensations into a unified textural impression. The class supports comprehensive tactile assessment and style-appropriate evaluation where expected mouthfeel varies significantly among wine categories.
Gaumenempfindung
Mundgefühl
mouth sensation
oral texture
orale Textur
palate feel
tactile impression
taktiler Eindruck
Gawel R et al. (2000) A 'Mouth-feel Wheel': terminology for communicating the mouth-feel characteristics of red wine. Australian Journal of Grape and Wine Research 6(3): 203-207 Pickering GJ, Demiglio P (2008) The white wine mouthfeel wheel. Journal of Wine Research 19(2): 51-64
Mouthfeel Quality
The mulching protocol for a Grüner Veltliner vineyard in the Kamptal specifies annual application of composted vine pruning chips (produced by on-site hammer mill processing of dormant pruning wood) to the under-vine strip at a rate of 15-20 m³/ha (approximately 8-10 cm depth across a 0.5 m band centered on the vine row), applied in March after the last inter-row tillage pass — the organic mulch layer suppresses annual weed germination by 70-85% (reducing herbicide requirements to zero in the under-vine zone), reduces soil moisture evaporation by 25-35% during the critical June-August period compared to bare soil, moderates diurnal soil temperature fluctuations by 3-5°C at 10 cm depth (buffering root zone temperatures during heatwave events), and provides a continuous input of approximately 2 t/ha/year of organic carbon to the soil surface as the mulch decomposes over 12-18 months.
A soil management description that documents the application of organic or inorganic materials to the vineyard soil surface to suppress weed growth, conserve soil moisture, moderate soil temperature extremes, and enhance soil biological activity, specifying the mulch material type, application rate, placement area (under-vine strip or full inter-row), replenishment frequency, and interaction with other soil management practices.
Mulching_Protocol documents a soil management practice that has gained renewed prominence as vineyards seek alternatives to herbicide-dependent under-vine weed management. The under-vine strip (typically 0.3-0.6 m width centered on the vine row) is the most management-intensive zone of the vineyard floor because mechanical cultivation risks root and trunk damage, cover crops compete directly with vine roots in the primary rooting zone, and uncontrolled weed growth provides habitat for trunk disease vectors and rodents: organic mulching addresses all these concerns simultaneously while providing the additional benefits of soil moisture conservation, temperature moderation, and organic matter input. Mulch materials include vineyard-derived resources (pruning wood chips, grape marc compost, straw from cover crops), agricultural byproducts (cereal straw, hay, bark), and commercial organic mulches — with recycling of vineyard pruning wood into mulch chips representing a particularly attractive circular economy approach that eliminates the carbon emissions from traditional pruning wood burning while returning nutrients and organic carbon to the vineyard soil. Inorganic mulches (gravel, stone) are used in some regions for weed suppression and thermal moderation but lack the biological benefits of organic materials.
Bodenabdeckungs-Spezifikation
Mulch-Protokoll
Mulchverfahren-Dokumentation
mulch application protocol
surface mulching specification
vineyard mulch management
White RE (2015) Understanding Vineyard Soils 2nd Ed, Oxford University Press pp: 215-230 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 445-455 Mundy DC and Agnew RH (2002) Effects of mulching with vineyard and winery waste on soil fungi and botrytis bunch rot in Marlborough vineyards, New Zealand Plant Protection 55: 135-138
Mulching Protocol
A multi-arm head on Vitis vinifera cv. Carignan in Languedoc, featuring seven arms radiating from compact head at 48cm height, arms extending 32-45cm at various angles, total 38 spurs distributed across seven arms, supporting approximately 68 shoots and 7.2kg crop, representing adapted head training employing additional arms to achieve economically viable yields (approximately 54 hl/ha at 1.5m × 1.5m spacing) while maintaining head-trained architecture avoiding trellis conversion, common in Languedoc where traditional gobelet heritage combines with economic pressure for improved productivity creating evolution toward higher-arm-count head systems as compromise between tradition and commercial necessity.
A head-trained configuration featuring numerous arms (typically 5 or more) radiating from central head, characterized by complex multi-armed architecture beyond simple 3-4 arm gobelet, increased fruiting capacity distributed across multiple structural elements, and suitability for vigorous vines, fertile sites, or training approaches emphasizing productivity within head-trained framework. Multi-arm heads represent enhanced capacity head training, employing additional arms to increase spur positions and crop potential while maintaining head-trained architecture, suitable for situations where traditional low-arm counts inadequately utilize vine vigor, modified traditional systems adapting to contemporary productivity requirements, or specific training philosophies distributing capacity across numerous smaller arms rather than few larger structures.
Multi-arm heads represent capacity enhancement strategy within head-trained framework, adding arms to increase productivity while avoiding conversion to cordon systems. The multi-arm approach (5-8+ arms versus traditional 3-4) distributes fruiting capacity across numerous structural elements, potentially providing benefits including balanced growth distribution (vigor divided among many arms), production insurance (redundancy if some arms fail), architectural flexibility (individual arm positioning adapting to space constraints), and gradual capacity increase (arms added incrementally as vine vigor develops). However, multi-arm configurations create management challenges including complex pruning (maintaining balance among numerous arms), difficult renewal (coordinating arm replacement without excessive disruption), potential structural weakness (many junction points at head), and aesthetic irregularity (complex multi-arm structures often lacking formal symmetry). The trend toward multi-arm heads in some traditional regions reflects pragmatic adaptation to economic realities requiring higher yields while maintaining head-trained identity.
Vielarm-Kopf
complex head structure
mehrarmige Kopferziehung
Multi Arm Head
A multi-branched compound inflorescence at node 3 on a shoot of Vitis vinifera cv. Muscat Blanc à Petits Grains in Languedoc-Roussillon, France, measuring 22.5 cm length, exhibiting very dense lateral branching with 26 major lateral branches distributed along the rachis at intervals of 0.7-1.2 cm creating closely packed arrangement, major lateral branches showing substantial individual development ranging 4.5-7.5 cm length with the longest laterals near the base, each major lateral bearing extensive secondary branching with 8-12 secondary branches per primary lateral, secondary branches further elaborating into tertiary and quaternary levels, overall architecture creating very dense multi-branched structure approximately 8.5 cm width, total flower number estimated at 720 flowers distributed throughout the numerous branching systems, architectural density creating interior shading with limited light penetration to central regions, rachis diameter of 4.5 mm at base supporting the elaborate multi-branched structure, measured at BBCH stage 63 (30% flowers open) in late May, representing typical multi-branched compound architecture with emphasis on numerous densely arranged lateral branches characteristic of high-fertility Muscat cultivars.
A compound primary inflorescence characterized by numerous major lateral branches creating extensive branching elaboration in Vitis vinifera, characterized by high major lateral branch number typically 18-30+ branches along the rachis, dense lateral branch spacing often 0.8-1.5 cm intervals creating closely packed arrangement, major laterals showing substantial individual development with lengths often 4-8 cm each, lateral branch architecture showing extensive secondary and tertiary elaboration with each major lateral developing complex branching systems, overall architecture creating multi-dimensional structure with branches radiating extensively from the central rachis, total structure width often 6-10 cm complementing lengths of 20-28 cm creating substantial three-dimensional volume, and architectural effect producing very dense elaborate paniculate structures with numerous distinct branch systems visible throughout. Multi-branched compound inflorescences represent the high lateral branch number variant of compound architecture, emphasizing quantity and density of major lateral branches rather than extreme hierarchical depth, occurring in cultivars and conditions favoring extensive lateral branch initiation and development, and creating very full dense inflorescence architecture transforming into very dense compact clusters following fruit set.
Multi-branched compound inflorescences are distinguished from other compound types by emphasis on high lateral branch number and density rather than extreme hierarchical depth or other elaboration patterns. The multi-branched designation emphasizes the quantity of major lateral branches creating the architectural elaboration. Branch number >18-20 major laterals represents elevated development compared to typical 10-15 laterals in standard inflorescences, with >25-30 representing extreme multi-branching. Spacing density with intervals <1.5 cm creates closely packed arrangements contrasting with typical 1.5-2.5 cm spacing, producing the characteristic dense appearance. Individual lateral development with substantial lengths and complex secondary branching creates the effect where each major lateral resembles a complete small inflorescence. Three-dimensional volume reflecting substantial width complementing length creates full dense structures rather than elongated narrow forms. Occurrence correlates with high fertility and favorable development conditions supporting extensive lateral branch initiation and growth. Developmental mechanisms involve elevated lateral primordium initiation frequency and sustained lateral branch development creating the numerous elaborate branches. Cluster density following fruit set creates very compact architecture with potential disease and quality issues requiring management intervention through cluster thinning, berry thinning, or other techniques. Ripening uniformity may suffer with interior berries shaded by dense branch packing potentially creating maturity gradients within clusters.
densely branched inflorescence
dicht verzweigter Blütenstand
high-branch-number inflorescence
numerous lateral compound
vielverzweigte zusammengesetzte Infloreszenz
zahlreiche Lateral-Rispe
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 383-418 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 426-465 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 619-658.
Multi Branched Compound Inflorescence
A multi-branched permanent arm on Vitis vinifera cv. Seyval Blanc in Geneva Double Curtain training system, featuring trunk dividing into two primary arms at 90cm height (bilateral split), each primary arm then bifurcating at 45° angles creating four total secondary arms (two ascending at approximately 60° from horizontal, two descending at approximately 120° from horizontal), each secondary arm extending 65-75cm bearing 11-13 spurs, total configuration creating four distinct fruiting curtains with divided canopy architecture, combined 48 spurs supporting approximately 90 shoots and 14-16kg total crop distributed across complex branched framework, representing intentional multi-branched architecture designed for light penetration, air circulation, and disease management in humid climate requiring maximum canopy division.
A permanent arm configuration featuring multiple branching points creating three or more distinct arm segments or sub-arms, characterized by complex branched architecture beyond simple bilateral division, fruiting positions distributed across multiple arm branches rather than concentrated on single or paired elements, and structural complexity providing distributed capacity, architectural flexibility, or accommodation of specific training system requirements including Geneva Double Curtain, lyre, and other divided-canopy approaches. Multi-branched permanent arms represent advanced architectural configurations employing branching to achieve specific canopy distribution objectives, increase total fruiting surface area within given trellis space, create divided canopies improving light penetration and air circulation, or adapt to specialized training systems requiring multiple distinct fruiting zones, though at cost of increased structural complexity, more demanding establishment and management, and potential difficulties with mechanization compared to simple bilateral or unilateral arm configurations.
Multi-branched permanent arms are relatively uncommon in modern viticulture, with specific applications primarily in divided-canopy training systems (Geneva Double Curtain, Scott Henry, lyre variants) designed to improve canopy microclimate through structural division, high-density plantings requiring compact yet productive architectures, or situations where complex branching accommodates irregular spacing, terrain, or trellis configurations. The multi-branched approach increases structural complexity requiring skilled initial training (achieving desired branch angles and positions), more complex annual maintenance (managing multiple branch points and growth balance between branches), potential structural weakness at branch junctions (stress concentration requiring attention to avoid breakage), and often reduced mechanization compatibility (complex geometries challenging for equipment). Benefits include increased canopy surface area within given footprint, improved light penetration through division, enhanced air circulation reducing disease pressure, and architectural flexibility adapting to site constraints.
Mehrgabelarm
branched cordon system
complex arm architecture
mehrfach verzweigte Arme
multi-fork arm
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Reynolds & Vanden Heuvel (2009) "Influence of grapevine training systems on vine growth and fruit composition= a review" American Journal of Enology and Viticulture 60=251-268 Shaulis et al. (1966) "Response of Concord grapes to light, exposure and Geneva Double Curtain training" Proceedings of the American Society for Horticultural Science 89=268-280.
Multi Branched Permanent Arm
A multi-branched tendril at node 6 on an exceptionally vigorous shoot of Vitis vinifera cv. Muscat of Alexandria in southern Spain, exhibiting primary bifurcation at 4 cm from shoot attachment, followed by secondary bifurcation of both primary arms at 8-9 cm creating four terminal branches, with a fifth branch arising from an additional lateral meristem at the primary branch point, producing total architecture of five arms ranging 7.5-10.5 cm individual length with combined extension of 38 cm across all branches, all arms showing 2.2-2.8 mm diameter indicating substantial but variable vascular development, circumnutation movements across all arms collectively exploring approximately 920 cubic centimeters creating very high contact probability, three arms making successful attachment to different trellis wires creating exceptionally stable multi-point support system, occurring on a shoot with extraordinary vigor (total length 220 cm, 18 nodes) in heavily fertilized and irrigated conditions, measured at BBCH stage 71 (fruit set) in early June, representing the rare achievement of functional multi-branched architecture in a cultivar known for complex tendril development.
A branched tendril exhibiting division into four or more branches in Vitis vinifera, characterized by presence of multiple branch points creating complex architecture with numerous independent arms, representing rare but significant branching patterns occurring in 1-8% of branched tendrils in most cultivars with higher frequencies in certain varieties showing genetic predisposition toward complex branching and under conditions of exceptional vigor supporting extensive resource allocation to individual tendrils. Multi-branched tendrils develop through activation of four or more meristems during early tendril organogenesis, either simultaneously at one or more branch points or sequentially through multiple rounds of branching creating hierarchical architectures with primary, secondary, and potentially tertiary branch levels, requiring substantial metabolic investment and carbohydrate allocation to support multiple growing points and extensive total tendril length often 25-40 cm combined across all branches. The functional implications of multi-branching include maximum spatial exploration volume among all tendril architectures potentially reaching 800-1200 cubic centimeters of searched space, very high probability of multiple successful attachments creating exceptionally robust mechanical support systems, potential for load distribution across 3-5 or more attachment points, but significant resource cost requiring high vine vigor to support adequate individual arm development and risk of reduced arm strength through excessive resource partitioning if total resource availability is insufficient for the number of branches produced. Multi-branched tendril frequency serves as a sensitive indicator of exceptional vine vigor and resource availability, and in cultivars showing genetic tendencies toward complex branching provides valuable ampelographic diagnostic characters.
Multi-branched tendrils are distinguished from bifurcated and trifurcated tendrils by four or more total arms, and represent the upper extreme of branching complexity in Vitis vinifera. Branching hierarchies vary from relatively flat architectures with four-way or five-way single branch points, to deep hierarchies with multiple sequential branching levels creating tree-like structures. Multi-branching imposes substantial resource costs and is typically only sustainable on very vigorous shoots with exceptional resource access, making frequency of multi-branched tendrils a valuable vigor indicator. Some cultivars including certain Muscats, Malvasia types, and table grape varieties show elevated multi-branching frequencies (5-15% of branched tendrils) representing genetic variation in branching regulation. Functional benefits of multi-branching are substantial when individual arms achieve adequate development, but excessive branch number with inadequate per-arm resources can create many weak arms with reduced total functional capacity compared to fewer robust arms.
complex branched tendril
highly ramified tendril
hoch-ramifizierte Kletterorgan
komplex aufgeteilte Ranke
mehrfach verzweigte Ranke
multiple-fork tendril
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 166-182 This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 219-236.
Multi Branched Tendril
A multi-cluster fertile shoot on Vitis vinifera cv. Riesling from node 8 in Rheingau vineyard, 165cm length with 19 leaves totaling approximately 3135 cm² leaf area, bearing 3 well-developed clusters with 62, 58, and 54 berries showing successful fruit set and normal development, estimated total shoot crop 522g (3 clusters × 174g average cluster weight), adequate leaf-to-fruit ratio (16.0 cm² per gram) supporting complete maturation despite substantial multi-cluster load, representing natural high fruitfulness of Riesling combined with vigorous balanced shoot development, managed without cluster thinning accepting three-cluster load as normal for this variety and appropriate for economically viable quality-oriented Riesling production (typical regional yields 70-90 hl/ha requiring multi-cluster retention on most shoots), with multi-cluster configuration contributing to acceptable productivity while maintaining quality standards through adequate leaf area support and variety characteristics tolerating higher yields without excessive quality compromise.
A fertile reproductive shoot bearing multiple well-developed clusters (2-4 typical, occasionally more in high-fruitfulness varieties), characterized by successful fruit set and development across all clusters, adequate shoot vigor and leaf area supporting multiple-cluster load (typically 18+ leaves, 2600+ cm² total area for two-cluster shoots), balanced resource distribution among clusters preventing dominance effects, and contribution of substantial crop weight (400-700g typical) representing high-productivity fertile shoots in commercial production contexts. Multi-cluster fertile shoots represent high-capacity productive units, occurring commonly at mid-cane positions in balanced vineyards, inherently high-fruitfulness varieties showing multiple-cluster patterns as normal differentiation outcome, and vigorous well-resourced shoots with capacity supporting multiple clusters without excessive strain, though requiring management assessment whether multiple-cluster loads align with quality objectives or cluster thinning warranted for quality enhancement.
Multi-cluster fertile shoots provide high per-shoot productivity, with their multiple-cluster loads contributing substantially to commercial yields and economic viability. However, multi-cluster configurations create management challenges including potential resource competition among clusters (first-position clusters often showing dominance over later positions), maturation variability (clusters ripening asynchronously requiring selective harvest or accepting heterogeneity), excessive crop risk (cumulative multiple-cluster loads potentially exceeding vine capacity if retained universally), and quality considerations (multi-cluster retention potentially diluting fruit composition versus single-cluster alternatives). Appropriate multi-cluster contexts include inherently high-fruitfulness varieties where multiple clusters are normal (Riesling, Gewürztraminer, certain Muscat selections), vigorous sites and shoots with capacity supporting multiple clusters, bulk production systems prioritizing yield over quality extremes, and balanced production approaches accepting moderate yields (60-90 hl/ha) as economically necessary while maintaining acceptable quality. Quality-focused premium production typically reduces multi-cluster shoots through thinning to single or reduced clusters, recognizing quality benefits despite yield sacrifice.
Mehrbüschel-Fruchttrieb
high-capacity reproductive shoot
mehrtraubiger fertiler Trieb
multiple-bunch fertile shoot
two-cluster fruiting shoot
Multi Cluster Fertile Shoot
A multi-cluster primary bud at node 6 on Vitis vinifera cv. Chardonnay, dissection revealing three distinct inflorescence primordia containing 185, 168, and 145 flower initials respectively, plus vegetative shoot primordia with 11 leaf initials, this bud developing into a shoot with three visible clusters at flowering representing potential yield of 249 berries assuming 50% fruit set, requiring cluster thinning to one cluster per shoot (removing two clusters) to achieve target crop load of 1.2 kg/m canopy and maintain leaf area-to-fruit ratio above 12 cm²/g for premium wine quality objectives.
A primary reproductive bud containing two or more differentiated inflorescence primordia (typically 2-3, rarely up to 4-5 in exceptional cases) in addition to vegetative shoot structures, representing maximum fruitfulness achieved under optimal growing conditions during previous season's inflorescence differentiation period, characterized by development into shoots bearing multiple flower clusters and potentially excessive crop load requiring cluster thinning management. Multi-cluster primary buds reflect favorable differentiation conditions including high light exposure (>60% full sun during June-August), optimal temperatures (25-30°C), adequate water status (predawn leaf water potential > -0.4 MPa), balanced nitrogen nutrition, and cultivar genetic predisposition toward high fruitfulness, with their abundance indicating well-managed vineyard conditions but necessitating crop load management through selective cluster removal to maintain fruit quality and sustainable vine balance.
Multi-cluster primary buds create a quality-quantity tradeoff requiring active management decisions. Retaining all clusters maximizes yield but typically results in delayed ripening, reduced sugar accumulation, higher acidity, diminished color and phenolic development, and depletion of vine reserves affecting cold hardiness and subsequent season fruitfulness. Standard practice in premium wine production removes excess clusters to 1-1.5 clusters per shoot, with timing of thinning (prebloom, postbloom, veraison) affecting developmental responses. The prevalence of multi-cluster primary buds indicates successful previous-season canopy management creating well-exposed bud zones, but ironically requires current-season crop reduction to realize quality benefits of that good canopy management. Balanced pruning formulas account for multi-cluster prevalence, with higher bud counts retained when fruitfulness is low and reduced counts when fruitfulness is high.
Mehr-Trauben-Primärknospe
Mehrfachtraubenknospe
high-fruitfulness primary bud
multiple-cluster primary bud
two-cluster primary bud
Based on Buttrose (1974) "Climatic factors and fruitfulness in grapevines" Horticultural Abstracts 44=319-326 Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Petrie et al. (2000) "The effect of leaf removal and canopy height on whole-vine gas exchange and fruit development of Vitis vinifera L. Sauvignon Blanc" Functional Plant Biology 27=569-578 Poni et al. (2006) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 57=397-407.
Multi Cluster Primary Bud
Carignan grafted on Rupestris St. George rootstock at 87 years post-grafting showing multi decade old grafted rootstock trunk 17.8 cm diameter, trunk extending 31 cm from root crown to ancient graft union positioned 24 cm above soil, graft union from 1939 completely undetectable beneath 9.2 mm extremely thick ancient bark, ancient trunk bark showing extreme deep longitudinal fissures with nearly black weathered surface, outer bark extensively sloughing in large plates, extensive lichen and moss completely covering bark, diameter showing no measurable growth recent 5 years, cross-section showing 87 annual rings with recent 20+ rings microscopic, massive central heartwood 14.2 cm diameter with functional sapwood only 1.8 cm narrow shell, trunk showing small hollow area in central heartwood and two old decay pockets in wound sites but remaining structurally adequate, supporting very sparse canopy 1.6 m shoot length, producing minimal crop 2.4 kg of extraordinarily concentrated fruit, ancient vine showing severe aging and fragility but remarkably still producing, vineyard designated historic heritage site, ancient vine irreplaceable viticultural treasure maintained with exceptional care
Multi decade old grafted rootstock trunk is old grafted rootstock trunk in extremely advanced age typically 60-100+ years post-grafting, characterized by many multiple decades of integration representing longest-lived grafted vines, extremely thick extensively weathered bark, very large diameter typically 12-20+ cm with minimal to absent annual growth, massive heartwood with very narrow functional sapwood, substantial vigor decline requiring intensive management, and representing extremely rare exceptional vines at end of natural lifespan continuing to produce limited but often highly valued fruit in ancient vineyard plantings of extraordinary heritage significance. Multi decade old grafted rootstock trunk extreme age includes= spanning 60-100+ years post-grafting (extremely advanced old trunk phase representing upper limit of grafted vine lifespan), representing multi-generational vine (trunk potentially planted by grandparents or great-grandparents of current vineyard operators), showing maximum aging characteristics (exhibiting most extreme manifestations of trunk aging indicators), displaying functionality at extreme age (remarkable continued production despite age far exceeding typical commercial vine lifespan), and marking ancient vine status (trunk age qualifying as ancient or very old vine by any regional definition). Multi decade old grafted rootstock trunk ancient graft union includes= showing 6-10 decade old graft union (callus tissue formed 60-100+ years prior representing oldest possible grafted vine integration), displaying century-scale vascular continuity (xylem and phloem connections maintained through multiple decades despite age-related efficiency reduction), exhibiting completely permanent graft union (bulge morphology unchanged for many decades representing absolutely stable ancient integration), showing utterly invisible graft line (massive trunk bark and ancient tissue integration making grafting point completely undetectable without destructive trunk examination), and demonstrating legendary integration durability (graft union having withstood 60-100+ years of stress representing exceptional rare achievement of lifelong successful integration). Multi decade old grafted rootstock trunk extreme physical characteristics includes= exhibiting very large ancient diameter (12-20+ cm with largest ancient trunks potentially reaching 25-30+ cm in extremely vigorous rootstock-scion combinations), showing extremely thick ancient bark (periderm 6-12+ mm thickness representing many decades of successive cork cambium activity), displaying maximum depth fissures (longitudinal furrows reaching extreme depth with severe weathering and erosion from decades of exposure), revealing extremely dark weathered coloration (bark nearly black or dark grey from extreme oxidation and environmental exposure over many decades), showing extensive bark sloughing and damage (outer bark extensively separating with large areas of dead bark or exposed wood from century-scale weathering), potentially showing significant structural damage (accumulated wounds, hollow areas, or decay zones from many decades of exposure and stress), and appearing massively impressive and ancient (multi decade trunk conveying overwhelming impression of great age, stability, and survival). Multi decade old grafted rootstock trunk minimal growth includes= showing extremely reduced or absent annual increase (0-0.1 cm typical with many years showing no measurable diameter increase indicating approaching growth cessation), displaying highly erratic growth (occasional years showing slight increase while other years showing no growth creating very irregular recent ring pattern), exhibiting minimal cambial activity (vascular cambium barely active or dormant in many multi decade trunks), forming invisible or extremely narrow recent rings (most recent growth rings microscopic or absent indicating effective cessation of radial growth), and essentially ending diameter expansion (multi decade trunk approaching or reaching end of active growth phase with trunk diameter effectively fixed). Multi decade old grafted rootstock trunk massive heartwood dominance includes= showing dominant heartwood core (vast majority of trunk cross-section converted to non-functional heartwood potentially 70-85% of total diameter), displaying very narrow functional sapwood (active water-conducting xylem reduced to thin outer shell typically 1.5-2.5 cm representing small fraction of total trunk), exhibiting tiny functional area (very limited sapwood cross-section severely constraining water and nutrient transport capacity), showing dramatic heartwood-sapwood contrast (extreme difference in cross-sectional areas between dominant dark heartwood and narrow light sapwood), and maintaining minimally adequate transport (very narrow sapwood sufficient only for severely reduced canopy and crop of ancient vine with no capacity for stress or increased demand). Multi decade old grafted rootstock trunk substantial vigor decline includes= showing severe vegetative vigor reduction (shoot growth severely reduced compared to younger vine with very weak short shoots), exhibiting sparse minimal canopy (leaf area severely reduced with very sparse canopy coverage), displaying very weak shoot growth (annual shoots very short and weak potentially only 30-50 cm length compared to 2-4 m in vigorous younger vine), showing severely reduced reserve capacity (minimal carbohydrate storage in aging trunk with very limited stress tolerance), exhibiting substantially delayed phenology (bud break, flowering, and ripening significantly delayed compared to younger vines), and demonstrating fragile vigor balance (ancient vine vigor easily disrupted by stress with limited recovery capacity). Multi decade old grafted rootstock trunk severely reduced production includes= generally showing very low fruit production (ancient trunk supporting minimal crop potentially 1-4 kg compared to 5-12 kg in mature vine), often producing highly distinctive fruit (ancient vine fruit potentially showing extreme concentration, unique character, and old vine typicity), displaying very small clusters and berries (severe natural yield limitation manifested as tiny clusters with small concentrated berries), exhibiting extreme vine balance (very low vigor matched with minimal crop creating naturally balanced ancient vine), showing highly variable or declining annual crops (year-to-year production erratic with potential for crop failure in stress years), potentially producing exceptional quality when successful (ancient vine fruit often valued for extraordinary concentration, complexity, and distinctive character when vine manages to ripen crop). Multi decade old grafted rootstock trunk structural vulnerability includes= potentially showing compromised structural integrity (some ancient trunks structurally weakened by decay, hollows, or damage), displaying extensive damage accumulation (century-scale vineyard operations resulting in numerous old wounds, scars, and injury sites), frequently exhibiting significant decay (many ancient trunks showing substantial wood rot in central heartwood and old wound sites), showing large old wound areas (multiple extensive injury sites from decades of damage potentially affecting structural capacity), commonly showing hollow trunk (many ancient trunks developing significant internal cavities in decaying central heartwood), occasionally requiring structural support (some ancient trunks needing stakes or props due to compromised structural integrity), and requiring careful structural assessment (ancient trunk needing evaluation to ensure continued stability and worker safety). Multi decade old grafted rootstock trunk disease challenges includes= showing high disease susceptibility (ancient aging trunk highly vulnerable to trunk diseases particularly in stressed areas), commonly exhibiting chronic trunk diseases (Esca, Eutypa, Botryosphaeria, or other pathogens frequently affecting ancient trunk), displaying multiple disease infection sites (evidence of numerous disease events over decades with varying degrees of compartmentalization success), showing reduced disease defense capacity (ancient trunk very limited ability to mount effective defense or compartmentalization responses), requiring intensive disease management (ancient trunk needing careful monitoring and intervention to prevent disease progression threatening survival), and demonstrating survival despite disease pressure (many ancient trunks remaining functional despite chronic disease burden testament to exceptional resilience and management). Multi decade old grafted rootstock trunk intensive management requirements includes= requiring critical ongoing viability assessment (constant evaluation of whether ancient trunk can continue or has reached end of functional life), needing very intensive health monitoring (frequent careful inspection essential for detecting problems threatening ancient vine survival), potentially requiring specialized cultural practices (modified irrigation, nutrition, crop limitation, or other interventions helping maintain ancient vine), facing difficult continuation versus replacement decisions (determining whether to maintain ancient irreplaceable vine or accept end of life and replant), valuing extraordinary fruit quality and heritage (managing ancient vine to optimize old vine quality and preserve historical significance), implementing protective measures (providing physical protection, disease management, and stress reduction for fragile ancient trunk), and planning for eventual vine end (accepting and preparing for eventual death or loss of ancient vine despite best management efforts). Multi decade old grafted rootstock trunk as viticultural treasure includes= representing extraordinary rarity (multi decade old grafted trunks extremely rare representing tiny fraction of vineyard plantings), embodying irreplaceable viticultural heritage (ancient plantings impossible to replicate representing lost historical germplasm and practices), contributing fundamentally to regional identity (ancient vineyards often central to regional wine heritage and reputation), demonstrating ultimate viticulture achievement (maintaining productive ancient vine through century-scale time period showing exceptional stewardship), potentially protected as cultural heritage (some ancient vineyards designated as historic sites or protected resources), commanding extreme market value (fruit and wine from ancient vines often highly prized commanding premium prices), and serving as living history (ancient trunk connecting present directly to viticultural practices and people from century ago). Multi decade old grafted rootstock trunk diameter extremes includes= typical ancient diameter 12-20 cm (common range for multi decade trunks), vigorous ancient rootstocks showing 18-28+ cm (1103P, 110R, St. George reaching truly massive dimensions), moderate vigor ancient rootstocks showing 12-18 cm (SO4, 3309C, Teleki 5C showing substantial ancient trunk), low vigor ancient rootstocks showing 10-15 cm (Riparia types reaching moderate ancient size), and record largest ancient trunks potentially exceeding 30 cm (extremely rare oldest and most vigorous ancient vines reaching truly exceptional dimensions unique to century-old plantings). Multi decade old grafted rootstock trunk in ancient vineyard context includes= defining ancient vineyard classification (multi decade trunks characteristic of extremely rare ancient vineyard plantings), representing ultimate viticultural heritage sites (ancient vineyards with multi decade trunks serving as living museums and heritage sites), embodying century-scale tradition (ancient trunk connecting present viticulture to practices and people from previous century), contributing to wine historical continuity (ancient vineyards maintaining wine production from same vines across multiple generations), potentially designated as protected heritage (ancient vineyards eligible for historic landmark status, heritage site protection, or cultural preservation), commanding extraordinary respect and care (ancient plantings typically receiving exceptional care and reverence from vineyard operators and community), and serving educational and inspirational roles (ancient vineyards teaching about vine longevity, historical viticulture, and inspiring appreciation for tradition and stewardship). Multi decade old grafted rootstock trunk end-of-life considerations includes= approaching natural vine lifespan limit (multi decade trunk at or beyond typical maximum grafted vine longevity), facing inevitable eventual death or failure (ancient vine will eventually succumb to age, disease, structural failure, or accumulated stress), requiring acceptance of vine mortality (understanding and preparing for eventual loss of ancient irreplaceable vine), implementing succession planning (developing strategy for what follows ancient vine whether preservation attempts, replacement, or memorial), documenting vine history and significance (recording ancient vine story, characteristics, and heritage before eventual loss), potentially preserving genetic material (taking cuttings or grafting material preserving ancient vine genetics before death), and celebrating vine legacy (honoring ancient vine's extraordinary service and contribution to vineyard and wine heritage). Multi decade old grafted rootstock trunk climate and stress response includes= showing extreme climate sensitivity (ancient vine highly vulnerable to drought, extreme heat, or cold stress due to limited reserves and resilience), exhibiting limited stress recovery (ancient trunk minimal capacity to recover from stress events potentially leading to permanent decline or death), potentially showing climate change impacts (ancient vine possibly showing effects of multi-decade climate trends including temperature increase or precipitation changes), demonstrating adaptation to site (ancient trunk representing ultimate site adaptation having survived and produced through century-scale local climate variability), and requiring stress protection (ancient vine needing careful management to protect from environmental stresses that would be tolerated by younger vine). Multi decade old grafted rootstock trunk significance includes= representing ultimate grafted vine longevity (century-scale age demonstrating maximum potential lifespan of well-managed grafted grapevine), showing continued function at extreme age (ancient trunk remarkably maintaining production despite being far beyond typical vine lifespan), embodying irreplaceable viticultural heritage (multi decade trunk representing living history connecting present to previous century viticulture), requiring recognition of extraordinary value (ancient trunk deserving special recognition, protection, and management due to rarity and significance), facing inevitable mortality (ancient trunk approaching or at end of natural life requiring acceptance and preparation), commanding maximum old vine prestige (ancient vine fruit and wine representing ultimate old vine designation and value), and demonstrating human-plant partnership across generations (ancient trunk showing what can be achieved through dedicated stewardship across multiple human generations maintaining productive relationship with individual vine for century).
Multi_Decade_Old_Grafted_Rootstock_Trunk emphasizes extreme advanced age (60-100+ years) and is distinguished from Veteran_Old_Grafted_Rootstock_Trunk (mid old phase 40-60 years) by focus on ancient century-scale longevity at end of natural lifespan. Critical for understanding extreme aging characteristics, ancient vine rarity, maximum vine longevity, extraordinary heritage value, intensive management needs, inevitable mortality, and the multi-decade phase representing ultimate achievement of maintaining productive grafted vine across human generations.
Ancient grafted rootstock trunk
Century-scale grafted rootstock trunk
Extremely old grafted rootstock trunk
Jahrhundert-alter Pfropfstamm
Multi-Jahrzehnte alter verpfropfter Unterlagenstamm
Uralter Wurzelstockstamm
Multi Decade Old Grafted Rootstock Trunk
Zinfandel multi decade old grafted scion trunk on St. George (Rupestris St. George) drought-tolerant rootstock in California at 48 years post-planting shows 215-255 mm scion diameter with heartwood 68-73% of cross-section (145-185 mm diameter), functional scion sapwood 17-23 mm radial depth, graft union at 11 cm above soil with 118-126% swelling (increased from ~112% in prime maturity), rootstock diameter 230-275 mm, irregular growth rings (recent 5 years averaging 1.3-2.6 mm), 95-135 g combined starch reserves (scion 32-48 g, rootstock 63-87 g), produces 4.2-6.2 kg concentrated fruit from 20-32 clusters, shows Eutypa canker affecting 45% of cordons, rootstock drought tolerance enabling survival and quality production in water-limited Sierra Foothills site, designated old vine commanding 60-90% price premium
A multi decade old grafted scion trunk is an old grafted scion trunk of substantial age (typically 25-60 years post-planting) representing extended viticultural longevity in grafted configuration, characterized by large scion diameter, advanced heartwood development, declining but sustained productivity with continued rootstock benefit expression, and transitional status between typical old grafted vines and exceptional veteran combinations. Multi decade old grafted scion trunk characteristics include= substantial scion diameter (varying by rootstock= vigorous 130-280 mm at 30-50 years, moderate 110-220 mm, low-vigor 95-190 mm at equivalent ages), extensive scion heartwood (50-75% of cross-section), moderate functional scion sapwood (12-22 mm radial depth compared to 20-35 mm in prime), variable recent annual rings (0.8-3.5 mm width with occasional missing rings), graft union showing age effects (diameter ratio typically 112-125%, potentially increasing with age from prime maturity 108-120%), mature thick scion bark (7-18 mm), and rootstock trunk proportional aging (rootstock diameter typically 100-120% of scion at equivalent positions, also showing heartwood development and aging). Multi decade old grafted scion trunk graft union characteristics in aging include= union swelling potentially increasing with age (some combinations showing progression from 110-115% in prime to 115-125% in multi-decade status), union vascular continuity potentially declining (hydraulic conductance across union may decrease 10-25% from prime maturity levels), union callus showing age-related changes (lignified callus potentially developing minor cracks or imperfections without catastrophic failure), union as disease entry site (trunk disease incidence at unions 30-70% depending on region and age), but union stability generally maintained (decades of function confirming long-term compatibility despite age-related changes). Multi decade old grafted scion trunk physiological status shows= moderately reduced water transport (scion sapwood hydraulic conductance 55-85% of prime capacity, potential union bottleneck contributing additional 5-15% reduction), declining combined carbohydrate storage (scion and rootstock reserves 80-150 g compared to 120-190 g in prime), rootstock benefits continuing to support function (phylloxera resistance, drought tolerance, nutrient efficiency, vigor modulation still expressed), reduced vegetative vigor (annual shoot growth 65-110 cm compared to 100-150 cm in younger mature vines), extended phenology (budbreak and véraison delayed 5-15 days compared to young vines despite rootstock effects), and moderate stress sensitivity (greater drought, heat, cold susceptibility than prime but rootstock providing resilience own-rooted equivalents would lack). Multi decade old grafted scion trunk productivity patterns show= moderate yield decline (typically 3.5-7 kg per vine varying by rootstock class= vigorous maintaining higher yields longer, low-vigor showing earlier decline, representing 55-85% of prime maturity production), reduced fruitfulness (50-80% fruitful buds varying by rootstock influence), moderate cluster size reduction (110-190 g clusters), smaller berries with quality benefits (11-15 mm diameter, higher skin/pulp than young vines), rootstock quality effects maintained (anthocyanin enhancement, phenolic modulation, or other rootstock influences on composition still expressed), and potential quality improvements (concentration, complexity, distinctive old vine character with rootstock signature). Multi decade old grafted scion trunk disease incidence shows= moderate to high trunk disease presence (Eutypa, Esca, Botryosphaeria affecting 25-70% of multi-decade vines), graft unions showing elevated disease incidence (unions providing pathogen entry and colonization sites), beginning heartwood decay (white and brown rot fungi colonizing non-functional scion and rootstock heartwood), rootstock disease or decline (old rootstock roots potentially showing Armillaria, Phytophthora, or other root pathogens), but generally maintaining productivity with management. Multi decade old grafted scion trunk rootstock significance in extended longevity includes= vigorous rootstocks potentially extending productive life 5-15 years beyond moderate or low-vigor equivalents, drought-tolerant rootstocks supporting survival in water-limited conditions critical for declining vine resilience, lime-tolerant rootstocks preventing chlorosis enabling multi-decade productivity on calcareous soils impossible for own-rooted, phylloxera resistance enabling decades of survival in infested soils (comparing 30-60 year grafted productive life versus 3-10 year own-rooted survival post-infestation), and rootstock-soil matching critically affecting multi-decade performance (well-adapted combinations thriving, mismatched combinations showing premature decline). Multi decade old grafted scion trunk viticulture significance includes= old grafted vine quality potential (many premium wine regions recognizing 25-50+ year grafted vines for distinctive character), rootstock enabling economically viable old vines (rootstock benefits supporting productivity and quality justifying continued cultivation), genetic value (multi-decade grafted vines representing field-tested scion-rootstock combinations), terroir expression through rootstock lens (decades of site adaptation with rootstock modulation producing distinctive character), commercial positioning (old grafted vine wines commanding premiums while rootstock provides viability), and balanced economics (quality benefits and rootstock resilience offsetting yield decline). Multi decade old grafted scion trunk replanting considerations vary= quality-focused estates maintaining multi-decade grafted vines 30-60+ years for old vine character with rootstock benefits, commercial vineyards evaluating productivity decline versus quality gains and rootstock advantages (typically replanting 25-45 years when decline exceeds benefits), heritage sites preserving veteran grafted vines 40-80+ years for genetic and historical value, and economic analysis balancing premium pricing, rootstock benefits, declining yields, and increasing management costs.
Multi_Decade_Old_Grafted_Scion_Trunk emphasizes extended grafted longevity 25-60 years and is distinguished from Old_Grafted_Scion_Trunk (broader category), Veteran_Old_Grafted_Scion_Trunk (exceptional 40-80+ years), and mature categories by substantial age with rootstock-supported sustained viability. Critical for old grafted vine viticulture economics, understanding rootstock contributions to extended productivity, quality benefits of aged grafted vines with rootstock modulation, and replanting decision analysis for aging grafted vineyards.
Extended longevity grafted trunk
Generationenüberdauernde Aufpropfung
Langlebige Veredelung
Mehrere Jahrzehnte alter gepfropfter Stamm
Multi-generation grafted scion
Multi Decade Old Grafted Scion Trunk
Zinfandel multi decade old own-rooted trunk in California at 52 years post-planting shows 195-235 mm diameter with heartwood comprising 68-74% of cross-section (135-170 mm diameter), functional sapwood 16-22 mm radial depth, irregular growth rings (recent 5 years averaging 1.5-2.8 mm), 85-115 g starch reserves, produces 3.8-5.5 kg concentrated fruit per vine from 18-28 clusters, shows Eutypa canker on 40% of cordons requiring adapted pruning, commands 50-80% price premium as designated old vine wine
A multi decade old own-rooted trunk is an old own-rooted trunk of substantial age (typically 30-80 years post-planting) representing extended viticultural longevity beyond normal commercial vineyard replacement cycle, characterized by large trunk diameter, advanced heartwood development, declining but sustained productivity, and transitional status between typical old vines and exceptional ancient vines. Multi decade old own-rooted trunk characteristics include= substantial trunk diameter (typically 120-250 mm at 30-50 years, 180-350 mm at 50-80 years), extensive heartwood formation (50-75% of trunk cross-section in non-functional heartwood providing structural support but not water transport), moderate functional sapwood (12-22 mm radial depth compared to 20-35 mm in prime mature vines), variable annual ring formation (recent rings 0.8-3.5 mm width with occasional missing rings in severe stress years), thick mature bark (7-15 mm providing environmental protection), and potential beginning of trunk hollowing (central pith and early heartwood showing initial decay in oldest specimens). Multi decade old own-rooted trunk physiological status shows= moderately reduced water transport capacity (sapwood hydraulic conductance 50-80% of prime mature vine capacity), declining carbohydrate storage (starch reserves 70-130 g compared to 120-180 g in prime vines), reduced vegetative vigor (annual shoot growth 60-100 cm compared to 100-150 cm in younger vines), somewhat extended phenology (budbreak and véraison delayed 3-10 days compared to young vines), and moderate stress sensitivity (greater drought, heat, or cold susceptibility than prime vines but better resilience than ancient vines). Multi decade old own-rooted trunk productivity patterns show= moderate yield decline (typically 3-6 kg per vine, 60-90% of prime mature vine production depending on age within range), reduced but adequate bud fruitfulness (50-75% fruitful buds compared to 70-90% in prime vines), moderate cluster size reduction (cluster weight 100-180 g), smaller berries with quality benefits (berry diameter 11-14 mm, higher skin/pulp ratio than vigorous young vines), and potential quality improvements (concentration, complexity, distinctive character often associated with old vine status). Multi decade old own-rooted trunk disease incidence shows= moderate to high trunk disease presence (Eutypa, Esca, Botryosphaeria affecting 20-60% of vines depending on region and age), beginning heartwood colonization by decay organisms (white rot and brown rot fungi initiating wood decay), some canopy disease susceptibility from reduced vigor, but generally maintaining adequate health for continued productivity with appropriate management. Multi decade old own-rooted trunk viticulture significance includes= old vine quality potential (many wine regions recognizing 30-50+ year vines as old vines with quality benefits), genetic value (multi-decade vines may represent field selections, heritage clones, or unique adaptations), terroir expression (decades of site adaptation producing distinctive character), commercial wine positioning (old vine wines often commanding premium pricing), and balanced economics (declining yields offset by quality improvements and premium pricing in quality-focused production). Multi decade old own-rooted trunk management considerations include= adapted pruning strategies (accounting for declining vigor, disease presence, limited fruitful positions), crop load adjustment (typically 15-30 clusters per vine, reduced from 25-40 in prime vines), disease management (protecting remaining healthy wood, managing but not curing trunk diseases), targeted irrigation (supporting reduced water transport capacity during critical periods), gentle canopy management (avoiding stress from excessive leaf removal or hedging), and evaluation of continued viability (balancing quality benefits against declining productivity and increasing management costs). Multi decade old own-rooted trunk replanting considerations vary= quality-focused estates often maintaining multi-decade vines 40-80+ years for old vine character, commercial vineyards typically replanting at 25-40 years when productivity decline exceeds quality benefits, heritage sites preserving vines 50-100+ years for historical or genetic significance, and economic analysis balancing premium pricing potential against reduced yields and increased management costs.
Multi_Decade_Old_Own_Rooted_Trunk emphasizes extended longevity 30-80 years and is distinguished from Old_Own_Rooted_Trunk (broader senescence category) and Ancient_Old_Own_Rooted_Trunk (exceptional 50-150+ year longevity) by substantial age with sustained commercial viability. Critical for old vine viticulture economics, quality benefits of extended vine age, replanting decision analysis, and understanding longevity factors in different cultivars and sites.
Extended longevity trunk
Generationenüberdauernder Rebstamm
Langlebiger Eigenwurzelstamm
Mehrere Jahrzehnte alter Stamm
Multi-generation trunk
Multi Decade Old Own Rooted Trunk
A multi-primordium compound bud at mid-cane position on Vitis vinifera cv. Chardonnay following an optimal growing season with 2500 growing degree days, full canopy light exposure, and balanced crop load, containing a primary bud with two inflorescences of 200 and 175 flowers, a secondary bud with one inflorescence of 85 flowers and vegetative primordia, and a tertiary bud with vegetative primordia showing early differentiation toward reproductive fate, representing potential yield of 460 flowers from a single bud position.
A compound dormant bud containing multiple well-developed primordia across its primary, secondary, and sometimes tertiary bud positions, characterized by differentiated shoot and inflorescence structures in at least two of the three bud positions, representing high developmental investment and potential yield capacity. Multi-primordium compound buds typically develop under favorable growing conditions during the previous season, with adequate light exposure, optimal water and nutrient status, and appropriate vine balance, resulting in reproductive potential distributed across multiple backup positions that can compensate for spring frost damage or other bud injury through activation of secondary structures.
Multi-primordium compound buds represent maximum reproductive investment and are typical of well-managed vineyards with balanced vines. However, excessive primordium number can lead to overcropping if all positions develop, necessitating cluster thinning. Bud fruitfulness (number and size of inflorescence primordia) varies by node position along canes, with basal and distal positions typically less fruitful than mid-cane positions. Climate, cultivar, and cultural practices all influence multi-primordium development.
complex primordium bud
mehrfach-Primordien Knospe
multi-initiate compound bud
Based on Sanchez & Dokoozlian (2005) "Bud microclimate and fruitfulness in Vitis vinifera L." American Journal of Enology and Viticulture 56=319-329 Buttrose (1974) "Climatic factors and fruitfulness in grapevines" Horticultural Abstracts 44=319-326 Petrie et al. (2000) "The effect of leaf removal and canopy height on whole-vine gas exchange and fruit development of Vitis vinifera L. Sauvignon Blanc" Functional Plant Biology 27=569-578.
Multi Primordium Compound Bud
Thompson Seedless (Sultana) bears a Multi_Purpose_Scion_Role serving four distinct markets—table grape (early harvest at 16-18°Brix for fresh consumption), raisin (harvest at 22-24°Brix for drying), juice concentrate (processing of sorted-out table grapes and excess production), and wine (base wine for brandy or neutral blending wine)—with California growers allocating production among categories based on annual price signals. Muscat of Alexandria serves in a Multi_Purpose_Scion_Role for table grape (fresh consumption valuing aromatic character), raisin (Muscatel raisins with distinctive flavor), wine (Muscat wines, sweet and dry), and juice (aromatic grape juice) markets across Mediterranean production regions. Chasselas bears a Multi_Purpose_Scion_Role in Switzerland and surrounding regions, with production divided between premium table grape (historically Switzerland's finest) and wine (Swiss Fendant, Alsace Gutedel) markets.
A scion role that inheres in a grafted scion cultivar by virtue of its commercially significant utilization across two or more distinct grape product categories (wine, table grape, raisin, juice, distillation), reflecting the cultivar's versatility in meeting diverse quality and production requirements. Multi-purpose scions possess combinations of characteristics—sugar accumulation, acidity, berry size, flavor profile, yield potential, and post-harvest attributes—that enable successful production across market categories, with allocation decisions driven by relative market prices, vintage conditions, and producer strategy rather than inherent cultivar limitations. The multi-purpose designation reflects commercial reality rather than optimal suitability while these varieties serve multiple markets adequately, specialized varieties typically excel within single categories. Thompson Seedless exemplifies the multi-purpose archetype, serving table grape, raisin, juice concentrate, and wine (for brandy or blending) markets from the same varietal base. Multi-purpose utilization provides market flexibility and risk management for growers, with production allocation adjustable annually based on relative returns.
Multi_Purpose_Scion_Role captures cultivars whose versatility enables participation in multiple markets, with subclasses specifying common dual-purpose combinations. The multi-purpose strategy reflects risk diversification—growers can shift allocation toward whichever market offers best returns each season—but may sacrifice the optimization achievable with single-purpose varieties bred for specific uses. Market evolution has generally favored specialization (seedless table grapes, purpose-bred raisin varieties, aromatic wine grapes), but Thompson Seedless's continued dominance demonstrates the value of flexibility. The line between multi-purpose utilization and opportunistic marketing of excess production is sometimes blurred true multi-purpose varieties maintain quality across categories rather than merely tolerating secondary uses.
Mehrzwecktraube
Universaltraube
dual-purpose grape
flexible-use variety
multi-purpose grape
versatile grape variety
vielseitige Rebsorte
Creasy GL, Creasy LL (2009) Grapes, CABI Christensen LP (2000) Raisin Production Manual, UC ANR Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane
Multi Purpose Scion Role
A multi-year old wood latent bud on a 6-year-old section of bilateral cordon on Vitis vinifera cv. Pinot Noir, estimated at 4-5 years suppression based on surrounding bark characteristics, activated following crown gall infection that caused progressive dieback of distal cordon sections, producing a replacement shoot 145cm in length with 22 leaves, trained along the cordon wire to establish three fruiting spurs during first dormant pruning, yielding 1.8 kg of fruit in the second season post-activation, successfully replacing the diseased cordon section without requiring trunk-level renewal.
An old wood latent bud that has remained suppressed on perennial wood structures for multiple years, typically 3-8 years based on bark development and wood age assessment, characterized by deep embedment within accumulated periderm layers, reduced but still viable meristematic tissue, moderate to low activation probability under normal conditions, and capacity for vigorous shoot production when released from suppression through structural damage or severe pruning. Multi-year old wood latent buds represent medium-term reserve capacity for cordons and arms, offering intermediate options for structural renewal between annual bud positions (immediate production) and ancient wood or trunk positions (long-term survival reserves), with activation producing shoots suitable for establishing new fruiting positions within 1-2 seasons.
Multi-year old wood latent buds are intermediate between recently formed latent buds (high viability, easy activation) and ancient wood buds (declining viability, difficult activation). Their activation is common in cordon renewal programs addressing localized disease damage, mechanical injury, or planned architecture modifications. Viability assessment is challenging without destructive sampling, with external bark characteristics providing only rough age estimates. Management strategies for trunk disease control increasingly recognize multi-year old wood latent buds as valuable resources for distributed renewal, allowing gradual cordon replacement through successive activation of buds at different positions rather than catastrophic trunk-level renewal.
medium-age perennial wood bud
mehrjährige alte Holzknospe
several-year-old wood bud
Based on Burr et al. (1996) "Cordon age and virus infection of Concord grapevines influence phenology and productivity" HortScience 31=1163-1166 Bettiga (2013) "Grape Pest Management" 3rd Edition, University of California Agriculture and Natural Resources Publication 3343, pp. 178-192 Gubler et al. (2005) "Control of grapevine trunk diseases" Practical Winery & Vineyard Journal 27=1-8.
Multi Year Old Wood Latent Bud
Sauvignon Blanc in Marlborough shows multiple leaf unfolding at BBCH 12-15 during May after 1000-1100 chilling hours and reaching 650-850 growing degree days base 10°C, with second through fifth leaves unfolding sequentially at 3-4 day intervals reaching dimensions of 40-80 mm over 15-20 days under conditions of 16-20°C mean temperature, 60-70% relative humidity, adequate soil moisture and nutrition, and 13-14 hour photoperiod, representing typical cool maritime climate canopy development for this vigorous early-to-mid season cultivar
Multiple leaf unfolding represents the progressive phase where second, third, and subsequent leaves separate from shoot axis and expand sequentially, establishing developing shoot canopy with increasing photosynthetic capacity and marking transition from early development to active vegetative growth. This developmental stage corresponds to BBCH 11-15 as sequential leaves unfold with each new leaf designated by number in BBCH system, progressing from two leaves unfolded at BBCH 12 through five leaves unfolded at BBCH 15 and continuing beyond as shoot elongation proceeds. The unfolding leaves show characteristic pattern of size progression with each successive leaf typically larger than previous reflecting increasing shoot vigor and resource availability as photosynthetic capacity develops, with second leaf reaching 80-120% dimensions of first leaf, third leaf 100-140% of first, and subsequent leaves continuing expansion to eventual maximum dimensions determined by cultivar genetics and environmental conditions. Sequential leaf unfolding follows regular timing pattern with new leaves separating at approximately 3-5 day intervals under favorable conditions, creating progressively developing canopy with leaves at various developmental stages from recently separated to approaching mature dimensions and full photosynthetic capacity. The apical meristem maintains active cell production initiating new leaf primordia every 2-3 days in regular phyllotactic pattern, typically alternating positions approximately 180 degrees around shoot axis creating characteristic distichous leaf arrangement, with each new primordium developing through stages from multicellular initiation to recognizable leaf form over 10-15 days before separation from axis. Shoot elongation continues at 5-15 mm per day depending on temperature and resource availability, with internodes between successive leaves extending through cell division in intercalary meristems and cell expansion throughout internode tissues creating visible daily shoot growth. Photosynthetic capacity increases progressively as leaf area accumulates with each unfolding leaf, transitioning shoot from net carbon consumer dependent on stored reserves to net carbon producer with positive carbon balance typically achieved when 3-4 leaves have substantially expanded to functional dimensions. The metabolic transition from heterotrophic to autotrophic growth represents critical developmental milestone with shoots becoming self-supporting for carbon requirements while continuing to import nitrogen, minerals, and initially some carbon from permanent structures. Stored carbohydrate reserves show progressive depletion during multiple leaf unfolding with cane starch concentrations declining to 20-30% of winter maximum by time 4-5 leaves have unfolded, while newly produced photosynthate increasingly supplies carbon for continued growth. Water demand increases substantially with each additional unfolding leaf as transpiring surface area expands exponentially, with shoot water use increasing from 20-30 mg per hour with one leaf to 100-200 mg per hour with four to five leaves depending on leaf size, environmental conditions, and stomatal conductance. Vascular system extends continuously through elongating shoot with xylem differentiation maintaining functional water transport capacity and phloem development enabling bidirectional transport of resources including nitrogen and minerals imported from roots and photosynthate exported from expanding leaves. Nutrient demand intensifies during multiple leaf unfolding particularly for nitrogen supporting protein synthesis in expanding leaf tissues, with nitrogen concentration in young expanding leaves 3-5% dry weight declining to 2-3% as leaves mature and structural carbohydrates dilute nitrogen-rich proteins. Cell division continues in apical meristem and expanding leaf primordia with high mitotic activity producing cells for continued shoot and leaf growth, while older leaves complete cell production and expansion approaching mature dimensions and beginning transition to fully differentiated functional state. Chlorophyll concentration increases in each successive unfolding leaf following characteristic developmental pattern reaching mature levels of 2.0-3.5 mg per gram fresh weight as expansion completes and chloroplasts mature, with variation among cultivars in final chlorophyll concentration affecting leaf color intensity. Photosynthetic capacity per unit leaf area increases in each leaf as it expands and chloroplasts mature, reaching maximum rates of 10-20 micromoles CO2 per square meter per second at light saturation when leaves approach mature dimensions, with whole shoot photosynthetic capacity increasing proportionally to total expanded leaf area. Hormonal regulation continues with cytokinin and gibberellin supporting growth, auxin coordinating developmental patterns and establishing positions for lateral organs including tendrils and inflorescence primordia that begin appearing in nodes as shoot develops, with first inflorescence primordia typically visible microscopically by 5-7 leaf stage though not externally apparent. Temperature remains primary factor controlling unfolding rate and shoot elongation with development rate doubling approximately per 10°C increase between 10-28°C optimal range, with higher temperatures potentially reducing growth efficiency through excessive respiration and water stress. Water availability critically affects unfolding rate, leaf expansion, and final sizes with adequate soil moisture essential for maintaining turgor supporting rapid expansion, while water stress slows or arrests development and permanently reduces leaf dimensions affecting photosynthetic capacity and subsequent growth. Light intensity strongly affects leaf expansion rates, final sizes, and photosynthetic capacity development with high light promoting rapid expansion to large dimensions with high photosynthetic capacity, while shade reduces expansion rates and final sizes but may increase specific leaf area through thinner blades adapting to light limitation. Nutrient availability particularly nitrogen and phosphorus affects unfolding rate and leaf development with adequate nutrition supporting rapid expansion to large dimensions with high photosynthetic capacity, while deficiencies reduce expansion rates, final sizes, and photosynthetic potential limiting subsequent growth. The process continues throughout shoot growth season with new leaves unfolding until shoot apex transitions to tendril or inflorescence production or until environmental limitations or management practices arrest growth, typically producing 12-20 leaves per shoot depending on vigor, cultivar, and management. Cultivar variation appears in leaf unfolding rate, final leaf number and sizes, leaf morphology including shape and dissection, and photosynthetic characteristics affecting canopy productivity. Regional timing shows continuous progression following first leaf stages, with rapid development during May-June in Northern Hemisphere warm climates, May-July in temperate regions, and June-July in cool areas. Viticultural management considerations include ensuring adequate water and nutrient availability during rapid leaf development, managing shoot positioning and canopy architecture to optimize light interception, monitoring for pest and disease colonization of expanding canopy, implementing shoot thinning or trimming operations to control vigor and improve fruit microclimate, and assessing canopy development rates informing irrigation and nutrition management. Damage to multiple unfolding leaves from pests, diseases, hail, or other factors reduces photosynthetic capacity potentially severely limiting shoot growth, fruit development, and reserve accumulation affecting current season productivity and subsequent season potential. Research applications include quantifying canopy development rates and leaf area expansion patterns, analyzing photosynthetic capacity development and carbon balance transitions, evaluating environmental and management effects on shoot growth patterns and final canopy characteristics, assessing hormonal regulation of shoot architecture and organ differentiation, and documenting cultivar variation in growth patterns affecting vineyard productivity and management requirements.
Multiple_Leaf_Unfolding is the third of three First_Leaf_Unfolding subphases, distinguished by sequential development of additional leaves creating expanding canopy. Transitions to Shoot_Growth as primary developmental designation once leaf unfolding pattern is established. Represents critical phase establishing shoot photosynthetic capacity and carbon balance. Encompasses BBCH 11-15 and beyond as shoot continues developing throughout growing season.
Bestandesentwicklung
canopy development
fortschreitende Blattentwicklung
mehrfache Blattentfaltung
multiple leaves spreading
progressive leaf development
sequential leaf unfolding
sukzessive Blattexpansion
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 205-235 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 105-128 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Multiple Leaf Unfolding
Muscat Blanc à Petits Grains bears a Muscat_Fortified_Scion_Role for Muscat de Beaumes-de-Venise (110+ g/L residual sugar, 15% alcohol) and Muscat de Frontignan, where mutage at 5-10% potential alcohol preserves terpene concentrations of 8-12 mg/L creating intense floral aromatics balanced by alcohol warmth. The same variety serves for Rutherglen Muscat in Victoria, Australia, where fortification, extended solera-style aging (10-60+ years), and evaporative concentration produce extraordinary wines of treacle-like viscosity, 180-400 g/L residual sugar, and complex rancio character while retaining distinctive Muscat perfume. Muscat of Alexandria bears a Muscat_Fortified_Scion_Role for Moscatel de Setúbal in Portugal and Muscatel de Valencia in Spain, providing somewhat coarser terpene character but excellent productivity for commercial fortified Muscat production.
A fortified wine scion role that inheres in a grafted scion cultivar of the Muscat family by virtue of its designation for fortified sweet wine production where intense terpene aromatics are preserved through early fortification. Muscat fortified scions accumulate exceptionally high monoterpene concentrations (5-15 mg/L combined linalool, geraniol, nerol, citronellol) that create the distinctive grapey, floral, orange-blossom character defining the category. Fortification during or shortly after fermentation (mutage) arrests fermentation while preserving residual sugar (typically 100-250 g/L) and locks in volatile terpene compounds that would otherwise diminish during extended fermentation or aging. Production styles include French Vins Doux Naturels (Beaumes-de-Venise, Frontignan, Rivesaltes, Mireval), Australian fortified Muscats (Rutherglen), Portuguese Moscatel de Setúbal, and various Mediterranean traditions. The Muscat family's numerous varieties (over 200 named forms) all share the defining terpene accumulation, with Muscat Blanc à Petits Grains considered the finest for fortified production.
Muscat_Fortified_Scion_Role captures fortified wines where the variety's distinctive terpene aromatics define the product character—contrasting with Port or Sherry where varietal neutrality is often preferred. The Muscat family's genetic diversity creates regional style variations Muscat Blanc à Petits Grains (considered ancestral) produces the finest wines, while Muscat of Alexandria (larger berries, higher yields) serves volume production. Rutherglen Muscat represents the extreme of the category, with extended oxidative aging developing rancio complexity while retained terpenes provide varietal identity even after decades in barrel. Climate strongly influences terpene accumulation, with hot climates producing intense grapey-floral character but risking terpene volatilization during harvest and processing.
Muscat fortified variety
Muscat-VDN-Sorte
Muskateller-Likörweinsorte
VDN Muscat variety
fortified Muscat grape
verstärkte Muskatsorte
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane pp.: 673-698 Jackson RS (2014) Wine Science 4th Ed, Academic Press Chambers B (2009) The Story of Muscat, Chambers Rosewood Vineyards Jefford A (2004) The New France, Mitchell Beazley
Muscat Fortified Scion Role
Must analysis data for Riesling from the Wachau (Smaragd category, 2024 vintage) records sugar content of 21.5° KMW (equivalent to 105° Oechsle), pH 3.08, titratable acidity 8.5 g/L (as tartaric acid), tartaric acid 6.2 g/L, malic acid 4.8 g/L, YAN 178 mg/L, and total phenolics 8.5 AU280 — establishing the lot as suitable for dry Smaragd vinification with adequate fermentation nutrients and characteristic Wachau acid structure. Must analysis data for Shiraz from McLaren Vale (2023 vintage) documents 14.2° Baumé, pH 3.55, TA 6.2 g/L, YAN 295 mg/L, total anthocyanins 2.1 mg/g, and tannin 3.5 mg/g catechin equivalents — indicating full phenolic maturity with sufficient nitrogen for clean fermentation.
Must analysis data is a quality analysis data entity that documents the chemical composition of freshly expressed grape juice prior to fermentation, including sugar concentration, organic acid content and pH, nitrogenous compound levels, and phenolic compound concentrations, measured through standardized analytical methods on representative vineyard samples, providing the primary dataset for harvest timing decisions, fermentation management planning, and the prediction of resulting wine composition and style.
Must analysis data constitutes the most widely performed and commercially consequential form of grape quality assessment, as its results directly trigger harvest decisions worth thousands of euros per hectare and determine winemaking interventions (acidification, chaptalization, nutrient supplementation) that shape the final wine. The analytical methods employed span simple field instruments (handheld refractometer for sugar, pH meter) through routine laboratory assays (titration for TA, enzymatic kits for specific acids) to advanced instrumentation (HPLC for organic acid profiling, spectrophotometry for phenolics, FTIR for multi-parameter rapid analysis). Fourier-transform infrared spectroscopy (FTIR) instruments such as the WineScan have transformed must analysis by enabling simultaneous measurement of 20+ parameters in under two minutes, though calibration against reference methods remains essential for accuracy. The interpretation of must analysis data requires integration of multiple parameters — a given sugar level carries different quality implications depending on the accompanying acidity, pH, phenolic maturity, and nitrogen status. The concept of "technological maturity" (adequate sugar for target alcohol), "phenolic maturity" (optimal tannin and anthocyanin extractability), and "aromatic maturity" (flavor development independent of sugar) reflects the multi-dimensional nature of ripeness that must analysis data captures only partially, necessitating complementary berry tasting and composition analysis.
Mostanalysedaten
Mostchemie
Mostuntersuchung
Mostzusammensetzung
Traubenmostanalyse
grape juice analysis
juice analysis
must chemistry data
must composition data
pre-fermentation analysis
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1: The Microbiology of Wine and Vinifications, 2nd Ed, Wiley, pp.: 219-255 Iland P et al. (2011) Chemical Analysis of Grapes and Wine, 2nd Ed, Patrick Iland Wine Promotions, pp.: 45-88 OIV (2021) Compendium of International Methods of Analysis of Wines and Musts, International Organisation of Vine and Wine
Must Analysis Data
Must composition maturity assessment for premium Sangiovese integrates multiple parameters for harvest optimization, with target composition profile of sugar 23-24 Brix, pH 3.35-3.45, TA 6-7 g/L, sugar/TA ratio 3.5-4.0, anthocyanin extractability >55%, and yeast assimilable nitrogen >200 mg/L, with vineyard monitoring revealing the optimal harvest window when all parameters simultaneously fall within target ranges (typically 3-5 days in favorable vintages), while challenging vintages may require compromise among parameters with prioritization based on wine style objectives, demonstrating how must composition maturity assessment integrates multiple quality determinants for informed harvest decisions.
Must composition maturity quality is a technological maturity quality that represents the integrated compositional state of grape must considering the balance and interaction among multiple parameters, characterizing the overall compositional profile rather than individual components, recognizing that wine quality depends not only on achieving targets for individual parameters but on the harmony among sugar, acid, phenolic, and aromatic constituents that together determine winemaking potential. Within the BFO framework, this quality specifically addresses the integrated compositional dimension of technological maturity, distinguishing holistic assessment from individual parameter evaluation while recognizing that the quality-determining interactions among must components (sugar-acid balance affecting perceived sweetness, pH-anthocyanin interaction affecting color, tannin-acid interaction affecting structure) require assessment of composition as a system rather than as independent variables. The ontological significance lies in must composition maturity determining the overall winemaking potential that emerges from compositional interactions, where favorable composition involves not merely acceptable levels of individual parameters but their appropriate ratios and balances that enable fermentation success and wine quality, with imbalanced composition (high sugar/low acid, for example) limiting wine quality potential regardless of individual parameter adequacy. The components of must composition assessment include the primary technological parameters (sugar, pH, TA) along with secondary parameters increasingly recognized as important (yeast assimilable nitrogen for fermentation health, phenolic indices for extraction potential, aroma precursor profiles for varietal expression), with the integration of these parameters providing a more complete picture of winemaking potential than any single measure. The balance indices developed to capture must composition maturity include sugar/acid ratios (Brix/TA, with typical targets 3-4 for table wines), maturity indices combining multiple parameters, and quality-focused assessments integrating technological and physiological maturity indicators, with these indices attempting to quantify the holistic composition assessment that experienced winemakers intuitively perform. The wine style implications of must composition maturity recognize that different compositions suit different wine styles, with high sugar/low acid favoring rich, full-bodied wines, moderate sugar/high acid favoring fresh, elegant styles, and with composition assessment appropriately referenced to style objectives rather than universal standards. Understanding must composition maturity quality enables integrated harvest decision-making considering compositional balance, wine style planning based on achievable composition profiles, and recognition of the vintage-specific compositional patterns that define each year's winemaking possibilities.
Must_Composition_Maturity_Quality addresses the integrated compositional profile considering balance among multiple parameters. This quality recognizes that wine quality emerges from compositional interactions rather than individual parameters. The class supports holistic harvest assessment through integrated must composition characterization.
Kompositionsbalance
Mostqualität
Mostzusammensetzungsreife
compositional balance
compositional profile
integrated maturity
integrierte Reife
must quality
Ribéreau-Gayon P et al. (2006) Handbook of Enology, Volume 1: The Microbiology of Wine and Vinifications, 2nd Ed. John Wiley & Sons, Chichester, pp. 193-230 Jackson RS (2014) Wine Science: Principles and Applications, 4th Ed. Academic Press, San Diego, pp. 232-298
Must Composition Maturity Quality
Must density monitoring of Grüner Veltliner in the Wachau follows the regional Smaragd classification requiring minimum 18.2 KMW (approximately 90°Oechsle or 21.5°Brix), with winemakers tracking density progression from 15 KMW at véraison through the critical threshold, typically achieved in mid-October when must density of 1.092 g/mL indicates sufficient sugar accumulation for the dry, full-bodied wine style characteristic of this premium designation.
Must density quality is a density quality that inheres in grape must following berry crushing and represents the mass per unit volume of the liquid fraction containing dissolved sugars, organic acids, phenolic compounds, nitrogenous substances, and mineral ions extracted from disrupted berry tissues. Within the BFO framework, this quality characterizes the processed material state distinct from intact berry density, where tissue disruption homogenizes previously compartmentalized cellular contents into a measurable liquid medium whose density primarily reflects total dissolved solute concentration. The ontological significance lies in must density serving as the traditional and legally recognized measure of grape maturity and potential alcohol yield, with established regional measurement scales including Brix (percentage sucrose equivalent by mass), Baumé (density-derived scale), Oechsle (grams above water per liter), and Klosterneuburger Mostwaage (KMW) each providing culturally embedded frameworks for harvest decisions and wine classification. Must density typically ranges from 1.050 g/mL (approximately 12°Brix) for early-harvested sparkling wine base material to 1.150 g/mL (approximately 35°Brix) for late-harvest botrytized wines, with the relationship between density and sugar content well-characterized by polynomial equations accounting for the contribution of non-sugar solutes. Measurement approaches include hydrometry using calibrated saccharometers, digital density meters employing oscillating U-tube technology, and refractometry which measures a related but distinct optical property requiring density correlation. The temperature dependence of must density requires standardization to reference temperature (typically 20°C) with correction factors applied for field measurements. Must density determination represents the critical control point for harvest timing, enabling prediction of fermentation dynamics, final alcohol content, and wine style classification, while also serving as the basis for pricing in grape purchase contracts and regulatory compliance for appellation requirements.
Must_Density_Quality specifically addresses the liquid fraction after tissue disruption, distinguishing from intact Berry_Density_Quality where tissue structure contributes to measured values. This class serves as the parent concept for regional must weight scales (Oechsle_Quality, KMW_Quality, Brix_Quality) defined under Sugar_Content_Quality, representing the same physical measurement expressed in different units. Temperature standardization and measurement methodology should be specified for regulatory and commercial applications.
Mostdichte
Mostgewicht
Saftdichte
Traubenmostdichte
crushing density
grape juice specific gravity
juice density
must weight
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed. John Wiley & Sons, Chichester, pp. 3-39 OIV (2021) Compendium of International Methods of Wine and Must Analysis, Volume 1, OIV-MA-AS2-01B
Must Density Quality
Must weight quality determines legal quality classification in German wine production, with Prädikat wines requiring minimum natural must weights that vary by region and category: Rheingau Kabinett requires 73° Oechsle minimum (approximately 17.7°Brix, 148 g/L sugar), Spätlese 85° Oechsle (20.4°Brix, 178 g/L), and Auslese 95° Oechsle (22.5°Brix, 203 g/L) vineyard managers monitor must weight progression daily during harvest period, selecting blocks and even individual clusters for appropriate Prädikat designation based on achieved must weight, demonstrating how this chemical quality parameter directly determines wine categorization and market value.
Must_Weight_Quality represents the chemical quality characterizing the density of grape juice or must relative to water, providing an indirect measure of sugar concentration based on the principle that dissolved sugars increase solution density. Must weight scales translate density measurements into practical viticulture and winemaking metrics, with various national and regional scales developed historically to suit local traditions and regulatory requirements. The relationship between density and sugar concentration is well-characterized, enabling accurate sugar estimation from density measurement without chemical analysis. This category encompasses specific must weight scales including Oechsle_Quality (German-speaking regions), KMW_Quality (Austria), Baumé (France, Australia), and specific gravity-based systems. Must weight increases during ripening as sugars accumulate, providing the traditional harvest maturity indicator in European viticulture predating refractometry. Measurement employs hydrometers calibrated to the appropriate scale, pycnometry for precise laboratory determination, or electronic density meters. Must weight values must be temperature-corrected because density varies with temperature independently of composition. The relationship between must weight and potential alcohol enables wine style planning, with legal minimum must weights established in European quality classification systems (e.g., German Prädikat levels). Must weight captures total density increase from all solutes, primarily sugars but also including acids, minerals, and other dissolved compounds, making it slightly different from direct sugar measurement but highly correlated in practice. The historical importance of must weight in European wine culture has embedded these measurements in regulatory frameworks, quality hierarchies, and traditional harvest assessment practices.
Must_Weight_Quality serves as the parent class for specific regional must weight scales including Oechsle_Quality and KMW_Quality. The various must weight scales are mathematically interconvertible because they all measure the same fundamental property (density), differing only in scale calibration and expression. Must weight is sometimes confused with Brix, but they measure different properties (density versus refractive index) that happen to correlate highly in grape juice. The class captures the measured density property the relationship to sugar concentration is an inference based on established calibration relationships.
must density
specific gravity quality
spezifische Mostmasse
sugar density index
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1: The Microbiology of Wine and Vinifications, 2nd Ed, John Wiley and Sons pp.: 223-240 Jackson RS (2020) Wine Science: Principles and Applications, 5th Ed, Academic Press pp.: 347-374
Must Weight Quality
The mutation information compiled for the Pinot cultivar family documents one of the most extensively studied somatic mutation histories in any crop species — from the ancestral Pinot Noir genotype, at least 50 recognized clonal variants have arisen through documented somatic mutations affecting berry skin color (Pinot Gris, Pinot Blanc, Pinot Meunier/Schwarzriesling through VvMYBA1 and VvMYBA2 mutations and chimeral rearrangements), berry flavor (Pinot Muscat through VvDXS gain-of-function mutation enhancing monoterpene production), growth habit (Pinot Fin, Pinot Droit, Pinot Tordu through mutations affecting internode length and shoot architecture), cluster morphology (Pinot Teinturier through mutations producing pigmented flesh), and maturity timing (early- and late-ripening Pinot Noir selections differing by 5-10 days in véraison date) — the cumulative somatic mutation record reveals that a single cultivar genotype can generate commercially significant phenotypic diversity through centuries of vegetative propagation and observational selection.
A genetic information entity that documents heritable changes in the DNA sequence of a Vitis vinifera individual that have arisen spontaneously or been induced after the cultivar's original establishment, encompassing somatic mutations accumulated during vegetative propagation, point mutations, transposable element insertions and excisions, chromosomal rearrangements, and epigenetic modifications that generate the phenotypic variation observed among clones, sports, and biotypes within a single cultivar.
Mutation_Information documents the molecular basis of intra-cultivar variation — the genetic changes that distinguish clones, sports, and biotypes within the framework of a single cultivar identity defined by the shared SSR fingerprint. The grapevine is uniquely informative for studying somatic mutation because the combination of long-lived perennial habit, exclusively vegetative propagation of cultivated varieties, and centuries-to-millennia propagation histories provides an extended timeframe for mutation accumulation without the homogenizing effect of sexual recombination: whole-genome comparisons of clones within cultivars (Vondras et al. 2019 for Zinfandel, Carrier et al. 2012 for Pinot Noir) reveal that clones differing by centuries of propagation history carry 200-700 somatic SNP mutations per genome, of which 5-15 affect protein-coding sequences and a smaller subset alter phenotypically relevant genes. The most commercially significant somatic mutations affect berry color (VvMYBA1/A2 mutations producing white and grey variants of red cultivars), flavor (VvDXS mutations creating muscat sports), and growth habit (mutations affecting vigor, shoot architecture, and cluster morphology). Subclasses capture the principal categories of documented somatic mutation.
Mutations-Information
genetic variant documentation
genetische Varianten-Dokumentation
mutation record
somatic variation data
somatische Variations-Daten
Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340 Vondras AM et al. (2019) The genomic diversification of grapevine clones, BMC Genomics 20: 972 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Kobayashi S et al. (2004) Retrotransposon-induced mutations in grape skin color, Science 304: 982
Mutation Information
Grapevines are obligate mycorrhizal hosts under natural conditions, with root colonization levels ranging from 20-80% depending on genotype, soil conditions, and management practices. Mycorrhizal association enhances P uptake by 30-80%, Zn uptake by 20-50%, and drought tolerance through improved water acquisition. The hyphal network extends the effective root zone by 10-100 fold, accessing soil volumes and pore spaces unavailable to roots alone. Vineyard AMF communities typically include Rhizophagus irregularis, Funneliformis mosseae, and Claroideoglomus species, with diversity correlating positively with vine nutrition and stress tolerance. Mycorrhizal benefits are greatest on low-P, low-input soils and diminish under high fertilizer regimes.
A nutrient acquisition disposition that is the inherited capacity of a Vitis vinifera plant to form and maintain mutualistic symbiotic associations with arbuscular mycorrhizal fungi (AMF, phylum Glomeromycota), enabling enhanced nutrient and water acquisition through fungal hyphal networks in exchange for photosynthetically derived carbon compounds.
Mycorrhizal_Association_Disposition serves as the parent class for colonization and dependency traits. The grapevine-AMF symbiosis is ancient and highly co-evolved, with vineyard soils maintaining AMF propagule banks that rapidly colonize establishing roots. Modern viticulture practices including fumigation, high P fertilization, and herbicide use can disrupt mycorrhizal function, reducing nutrient acquisition efficiency. Interest in mycorrhizal management is increasing as sustainable viticulture emphasizes reduced input agriculture and soil health.
AMF association trait
AMF-Symbiose
Mykorrhiza-Assoziation
Pilzsymbiose-Disposition
fungal mutualism disposition
mycorrhizal symbiosis capacity
Schreiner RP (2007) Effects of native and nonnative arbuscular mycorrhizal fungi on growth and nutrient uptake of Pinot noir (Vitis vinifera L.) in two soils with contrasting levels of phosphorus. Applied Soil Ecology 36:205-215 Trouvelot S et al. (2015) Arbuscular mycorrhiza symbiosis in viticulture: a review. Agronomy for Sustainable Development 35:1449-1467
Mycorrhizal Association Disposition
Mycorrhizal colonization assessment comparing conventional and organic vineyard management reveals the practice effects on beneficial fungal symbiosis, with organic vineyard showing 68% root length colonized (abundant arbuscules, extensive external hyphal network) and enhanced P uptake of 0.92 mg P/vine/day, while conventional vineyard with frequent fungicide applications shows only 32% colonization (reduced arbuscule development, limited external hyphae) and P uptake of 0.54 mg P/vine/day, demonstrating how management practices affect mycorrhizal function with consequences for nutrient acquisition that may require compensating fertilizer inputs in low-colonization systems.
Mycorrhizal colonization quality is a phosphorus uptake efficiency quality that represents the extent and functional effectiveness of arbuscular mycorrhizal fungal (AMF) symbiosis in grapevine roots, characterizing the development of fungal structures (hyphae, arbuscules, vesicles) that dramatically extend the effective absorbing surface for phosphorus and other nutrients while receiving photosynthate carbon from the vine host. Within the BFO framework, this quality specifically addresses the symbiotic dimension of nutrient acquisition, distinguishing mycorrhizal contributions from direct root uptake while recognizing that arbuscular mycorrhizal fungi are essential partners for grapevine phosphorus nutrition, with most vineyard vines naturally colonized by AMF communities that substantially enhance nutrient acquisition capacity. The ontological significance lies in mycorrhizal colonization determining the access to soil resources beyond the immediate root zone, where the extensive hyphal network (potentially adding hundreds of meters of absorbing surface per meter of root) can access phosphorus from soil volumes 100 times larger than roots alone could exploit, making mycorrhizal function critical for phosphorus nutrition in the low-P conditions typical of many vineyard soils. The structural components of mycorrhizal colonization include external hyphae (extending from roots into surrounding soil, actively acquiring phosphorus), arbuscules (highly branched intracellular structures where nutrient exchange occurs), vesicles (storage structures containing lipids and nutrients), and internal hyphae (connecting arbuscules and spreading within root cortex), with the relative development of these structures indicating the functional state of the symbiosis. The factors affecting mycorrhizal colonization include soil phosphorus level (high P often suppressing colonization), soil management practices (tillage disrupting hyphal networks, fungicide applications potentially harming AMF), host plant status (colonization levels varying with vine age, root growth, and carbon availability), and AMF community composition (different fungal species varying in colonization intensity and nutrient transfer efficiency). The vineyard management implications of mycorrhizal colonization extend beyond phosphorus to include enhanced uptake of zinc, copper, and water, improved soil structure from hyphal glomalin production, and potential pathogen protection, making maintenance of functional mycorrhizal associations a goal of sustainable vineyard soil management. Understanding mycorrhizal colonization quality enables assessment of the symbiotic contribution to vine nutrition, evaluation of soil management impacts on beneficial fungi, and optimization of conditions supporting effective AMF function for improved nutrient acquisition and overall vine health.
Mycorrhizal_Colonization_Quality addresses the arbuscular mycorrhizal symbiosis essential for grapevine phosphorus nutrition. This quality extends effective nutrient absorbing surface far beyond root system limits. The class supports soil management optimization through mycorrhizal status assessment affecting nutrient acquisition capacity.
AMF colonization
AMF-Besiedlung
Mykorrhizakolonisation
Wurzelpilzsymbiose
arbuscular colonization
arbuskuläre Kolonisation
mycorrhizal infection
root fungal symbiosis
Smith SE, Read DJ (2008) Mycorrhizal Symbiosis, 3rd Ed. Academic Press, London, pp. 1-787 Trouvelot S et al. (2015) Arbuscular mycorrhiza symbiosis in viticulture: a review. Agronomy for Sustainable Development 35(4): 1449-1467
Mycorrhizal Colonization Quality
Mycorrhizal dependency in grapevines ranges from 30-70% growth reduction in non-mycorrhizal versus mycorrhizal plants under low-P conditions, with high-dependency genotypes showing near-complete growth failure without AMF colonization. Rootstock 3309C exhibits high mycorrhizal dependency with 60-70% biomass reduction in non-mycorrhizal treatments, while 1103 Paulsen shows lower dependency (30-40% reduction) due to inherently efficient root P uptake mechanisms. Dependency correlates inversely with root hair density (r = -0.65) and fine root proliferation capacity, as genotypes with extensive root surface area partially compensate for absent hyphal networks. Under high-P conditions, mycorrhizal dependency approaches zero as direct root uptake satisfies P demand, though benefits for micronutrient acquisition and drought tolerance may persist.
A mycorrhizal association disposition that is the inherited capacity of a Vitis vinifera plant to require mycorrhizal colonization for adequate growth and nutrient acquisition, with genotype-specific dependency levels determining the magnitude of growth reduction when mycorrhizal associations are absent or impaired.
Mycorrhizal dependency quantifies the degree to which a genotype relies on AMF symbiosis for normal function, distinct from colonization capacity which measures the ability to support fungal growth. High dependency indicates limited alternative nutrient acquisition strategies and greater vulnerability to mycorrhizal disruption from soil fumigation, high fertilizer inputs, or fungicide applications. Understanding rootstock-specific dependency informs management decisions regarding soil health practices, transplant inoculation, and fertilizer strategies. Dependency is context-dependent, varying with soil P availability, water status, and the specific AMF species present.
AMF dependency
AMF-Dependenz
Mykorrhiza-Abhängigkeit
Pilzsymbiose-Bedürftigkeit
mycorrhizal responsiveness
obligate mycotrophy trait
Schreiner RP (2007) Effects of native and nonnative arbuscular mycorrhizal fungi on growth and nutrient uptake of Pinot noir (Vitis vinifera L.) in two soils with contrasting levels of phosphorus. Applied Soil Ecology 36:205-215 Linderman RG, Davis EA (2001) Comparative response of selected grapevine rootstocks and cultivars to inoculation with different mycorrhizal fungi. American Journal of Enology and Viticulture 52:8-11
Mycorrhizal Dependency Disposition
Myricetin content quality serves as an authenticity marker in wine fraud detection, with legitimate white wines showing undetectable myricetin (below 0.5 mg/L) while illegal addition of red wine or grape skin extract produces detectable myricetin levels (above 2 mg/L) that expose adulteration this application exploits the strict biosynthetic linkage between F3'5'H activity, myricetin synthesis, and red grape identity, providing regulatory laboratories with a reliable chemical test for white wine authenticity.
Myricetin_Content_Quality represents the chemical quality characterizing the concentration of myricetin and its glycosides in grape berry skins, capturing the trihydroxylated flavonol that shares the B-ring hydroxylation pattern with delphinidin-based anthocyanins. Myricetin (3,3',4',5',5,7-hexahydroxyflavone) bears three hydroxyl groups at the 3', 4', and 5' positions of the B ring, representing the most highly hydroxylated flavonol structure in grapes and the product of flavonoid 3',5'-hydroxylase (F3'5'H) activity on the flavonol pathway. The presence of myricetin exclusively in red grape cultivars reflects its biosynthetic dependence on F3'5'H, the same enzyme required for trihydroxylated anthocyanin production. White grape cultivars lack functional F3'5'H and consequently produce no myricetin or delphinidin-type anthocyanins. This cultivar-type specificity makes myricetin detection a marker for red grape or wine content, useful in authenticity testing to detect illegal blending of white wines with red wine for color enhancement. Myricetin typically comprises 10-25% of total flavonols in red grape cultivars, with myricetin-3-glucoside and myricetin-3-glucuronide as primary forms. Like other flavonols, myricetin accumulation responds to light exposure. The trihydroxylated B-ring structure provides strong antioxidant activity but also reduces chemical stability compared to less hydroxylated flavonols. Myricetin participates in co-pigmentation with anthocyanins, and its presence correlates with the blue-shifted coloration of trihydroxylated anthocyanins that share its biosynthetic origin.
Myricetin_Content_Quality addresses the trihydroxylated B-ring flavonol whose synthesis depends on F3'5'H activity present only in red grape cultivars. This strict cultivar-type association makes myricetin a powerful authenticity marker distinguishing red from white grapes and wines. The parallel between myricetin (trihydroxylated flavonol) and delphinidin/petunidin/malvidin (trihydroxylated anthocyanidins) reflects their shared biosynthetic pathway through F3'5'H. Myricetin proportion among flavonols correlates with trihydroxylated anthocyanin proportion, as both depend on F3'5'H activity level. The strong antioxidant capacity of myricetin reflects its multiple hydroxyl groups available for radical scavenging.
M-3-glc content
Myricetin-3-glucosid-Konzentration
Myricetingehalt
Myricetinglykosidgehalt
Trihydroxyflavonolgehalt
myricetin glycoside content
myricetin-3-glucoside concentration
trihydroxyflavonol quality
Castillo-Muñoz N et al. (2007) Flavonol Profiles of Vitis vinifera Red Grapes and Their Single-Cultivar Wines. Journal of Agricultural and Food Chemistry 55(3): 992-1002 Goldy RG et al. (1989) Anthocyanin Content of Muscadine Fruit. Journal of the American Society for Horticultural Science 114(3): 441-444
Myricetin Content Quality
Syrah narrow annual ring formed during 2015 severe drought year shows 0.8-1.4 mm radial width (compared to adjacent non-stress years 2.8-4.2 mm), reduced earlywood zone 0.4-0.7 mm with only 1-2 rows of vessels 140-200 μm diameter (compared to normal 1.5-2.5 mm earlywood with 2-3 vessel rows 180-260 μm), proportionally thick latewood zone 0.4-0.7 mm comprising 50-60% of ring, ring boundary distinct but overall narrow width indicating significant growing season stress
A narrow annual ring is an annual ring characterized by reduced radial width (typically 0.3-2.0 mm compared to 2-5 mm for typical rings or 4-8 mm for wide rings in same vine), indicating reduced cambial activity and secondary xylem production during the formation year, reflecting environmental stress, declining vine vigor, or reduced resource availability affecting growth. Narrow annual ring formation results from factors limiting cambial division and cell expansion= water deficit during growing season (drought stress reducing cell division rates, limiting cell expansion, shortening growth duration), nutrient deficiency (nitrogen, phosphorus, potassium limitation constraining photosynthesis and growth), disease or pest pressure (trunk diseases, phylloxera, or defoliating pests reducing vigor), excessive crop load (overcropping diverting photosynthate from vegetative growth to fruit), declining vine health (senescence, root decline, trunk disease progression), short growing season (early fall frost or late spring conditions limiting growth duration), and competition (weed pressure, vine crowding, or inter-plant competition for resources). Narrow annual ring anatomical characteristics include= reduced radial width (0.3-2.0 mm typical for stress rings compared to 2-5 mm normal rings in same vine), reduced earlywood zone (large vessel band 0.2-1.0 mm width compared to 0.8-2.5 mm in normal rings), fewer large earlywood vessels per radial file (1-2 rows compared to 2-4 rows in wide rings), smaller maximum vessel diameter (120-220 μm compared to 180-300 μm in wide rings), proportionally more latewood (latewood comprising 60-75% of narrow ring width compared to 40-55% in wide rings), higher fiber density (increased mechanical tissue compensating for reduced overall ring width), and compressed cellular structure (smaller cell dimensions throughout ring reflecting limited expansion). Narrow annual ring vessel characteristics include= reduced large vessel production (fewer earlywood vessels providing lower hydraulic conductance per ring), smaller maximum vessel diameter (genetic maximum not achieved due to resource limitation or shortened differentiation period), lower vessel lumen area fraction (vessels occupying 25-40% of ring area compared to 35-50% in wide rings), but maintained vessel functionality (narrow ring vessels still patent and capable of water transport if not embolized or occluded), and potential increased embolism vulnerability (smaller vessels with thinner walls potentially more prone to cavitation under tension). Narrow annual ring formation timing includes= early-season stress (drought or cold in April-June reducing earlywood production causing extremely narrow earlywood zones), mid-season stress (July-August drought or defoliation limiting latewood production), late-season stress (early fall frost or September drought terminating growth prematurely causing incomplete ring formation), or season-long stress (chronic water deficit, severe disease, or poor site conditions affecting entire annual growth period). Narrow annual ring dendrochronological significance includes= indicating stress years (narrow rings marking droughts, severe winters, disease outbreaks, or other adverse conditions), enabling cross-dating (matching narrow ring patterns across multiple vines or sites confirming synchronous environmental stress), providing climate reconstruction (ring width correlating with precipitation, temperature, and growing season conditions), and revealing management impacts (narrow rings following severe pruning, defoliation, or other cultural practices affecting growth). Narrow annual ring pattern interpretation includes= single narrow ring indicating acute stress year (isolated drought, freeze event, or disease outbreak), multiple consecutive narrow rings indicating chronic stress (sustained water limitation, declining vine health, progressive disease impact), progressive ring narrowing indicating senescence (gradual width decline reflecting age-related vigor reduction), and narrow rings in inner trunk indicating establishment stress (young vine establishing root system showing slow early growth). Narrow annual ring implications for vine physiology includes= reduced hydraulic conductance increment (narrow ring adding less water transport capacity than wide ring), reduced carbohydrate storage increment (less wood parenchyma providing reduced additional storage capacity), indication of source-sink imbalance (limited photosynthate or excessive fruit load restricting vegetative growth), and potential long-term vigor impacts (multiple consecutive narrow rings indicating cumulative stress potentially affecting vine longevity). Narrow annual ring occurrence patterns vary= drought-prone sites showing frequent narrow rings (Mediterranean climates with irregular precipitation producing variable ring widths), optimal sites showing rare narrow rings (only in extreme stress years), old vines commonly showing narrow recent rings (senescence-related vigor decline causing progressive ring narrowing), and young vines occasionally showing narrow establishment rings (limited root systems causing early stress). Narrow annual ring width thresholds vary by context= young vigorous vines (narrow ring <2.5 mm indicating stress, rings typically 3-8 mm), mature vines (narrow ring <1.5 mm indicating stress, typical 2-4 mm), old vines (narrow ring <1.0 mm indicating stress or senescence, typical 1-3 mm), with thresholds adjusted for vine age and typical growth rates on specific sites.
Narrow_Annual_Ring emphasizes reduced width stress indicator and is distinguished from Annual_Ring (all rings regardless of width) and Wide_Annual_Ring (favorable growth indicator) by reduced dimensions reflecting environmental stress or declining vigor. Critical for dendrochronological climate reconstruction, identifying stress years in vine history, understanding environmental impacts on growth, and assessing old vine decline through progressive ring narrowing.
Dünner Wachstumsring
Reduced growth ring
Schmaler Jahresring
Stress ring
Stressring
Thin ring
Narrow Annual Ring
A narrow petiolar sinus on a mature leaf at node 6 of Vitis vinifera cv. Merlot in Pomerol, France, exhibiting characteristic nearly closed configuration with basal lobe margins separated by only 0.4 cm creating narrow slit-like opening, sinus depth of 2.2 cm creating depth-to-width ratio of 5.5 indicating depth greatly exceeding width, U-shaped contour with rounded sinus base, left and right basal lobe margins showing strongly convergent orientation with margins angled inward at approximately 15 degrees from vertical creating nearly parallel approaching configuration, petiole barely visible or completely hidden at blade attachment requiring leaf manipulation to observe petiole insertion, sinus margins formed by large well-developed basal lobes measuring 3.2 cm length with margins approaching very close contact, narrow petiolar sinus configuration serving as primary diagnostic feature for Merlot enabling confident identification and clear distinction from open-sinus cultivars like Cabernet Sauvignon or overlapping-sinus Pinot Noir, measured at BBCH stage 75 (berries pea-sized) in late June, representing typical narrow nearly-closed petiolar sinus characteristic of broad-based cultivars with substantial basal lobe development.
A petiolar sinus exhibiting minimal separation or closure with basal lobe margins meeting or nearly meeting at the petiole-blade junction in Vitis vinifera, characterized by opening width typically less than 1 cm or completely closed with margins in contact, limited or no visibility of the petiole at the blade attachment, basal lobe margins showing convergent orientation creating closing rather than diverging configuration, typically U-shaped contour with depth often exceeding width creating tall narrow sinus profile, occurrence in cultivars with large convergent basal lobes including Merlot, Syrah, Grenache, and other broad-based varieties, and functional implications potentially providing enhanced mechanical integration at petiole-blade junction through large basal lobe overlap though experimental verification is lacking. Narrow petiolar sinuses represent the closed extreme of petiolar sinus configurations contrasting with overlapping types where margins actually cross, developing through large convergent basal lobe patterns creating characteristic broad-based leaf architecture, exhibiting high diagnostic reliability distinguishing closed-sinus cultivars from open-sinus varieties, and serving as definitive identification features particularly when combined with other morphological characters.
Narrow petiolar sinuses are distinguished from wide sinuses by minimal gap width typically <1 cm and from overlapping sinuses by margins meeting rather than crossing. The narrow designation emphasizes the constricted closed character creating minimal or no petiole visibility contrasting with open wide configurations. Depth typically exceeds width often substantially creating tall narrow sinus profiles contrasting with broad shallow wide sinuses. Basal lobe characteristics determine sinus narrowness, with large convergent basal lobes creating narrow or closed configurations while small divergent lobes create wide sinuses. The distinction between narrow (nearly closed) and overlapping configurations can be subtle in borderline cases, with classification depending on whether margins merely approach or meet (narrow) versus actually cross (overlapping). Occurrence predominates in broad-based cultivars showing substantial basal lobe development, including Merlot, Syrah, Grenache, and other varieties with pronounced basal lobes. Genetic control is strong with narrow sinus pattern showing consistent expression. Environmental stability is generally high though extreme vigor may occasionally produce slightly more open sinuses requiring assessment of multiple leaves. Functional implications remain speculative with proposed mechanical advantages lacking strong experimental support. Ampelographic utility is high with narrow/closed sinus serving as definitive diagnostic character distinguishing Merlot and related cultivars from open-sinus varieties, often enabling field identification based primarily on petiolar sinus configuration.
closed petiolar sinus
constricted petiole sinus
enger Petiolarsinus
geschlossene Stielbucht
narrow basal sinus
verengte Basaleinbuchtung
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 183-215 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 103-136.
Narrow Petiolar Sinus
Chardonnay grafted on SO4 rootstock at 18 years post-grafting showing narrow rootstock annual ring for year 14 measuring 0.18 cm radial width, narrow ring contrasting with adjacent rings showing 0.42 cm (year 13) and 0.38 cm (year 15) indicating year 14 specific stress, narrow ring structure showing compressed earlywood zone with reduced large vessel production and minimal latewood zone indicating early growth cessation, weather records confirming severe summer drought during year 14 matching narrow ring timing, narrow ring followed by return to normal width year 15 indicating stress relief and recovery, microscopic examination of narrow ring revealing smaller vessel diameters and fewer cells produced compared to normal rings, narrow ring documenting specific drought year impact on trunk growth providing dendrochronological stress record
Narrow rootstock annual ring is rootstock annual ring with reduced radial width typically less than 0.3 cm, characterized by limited secondary xylem production during growing season reflecting constrained cambial activity from limiting factors including water stress, nutrient deficiency, extreme temperatures, pest or disease damage, advanced trunk age, inherently low rootstock vigor, or competitive stress, showing compressed earlywood and latewood zones with fewer cells produced, reduced total vessel area potentially limiting water transport capacity, and serving as indicator of unfavorable growing conditions, physiological stress, or age-related growth decline during formation year requiring interpretation in context of trunk age, rootstock cultivar, and environmental history. Narrow rootstock annual ring width definition includes= showing radial width less than 0.3 cm (narrow ring typically 0.1-0.3 cm compared to typical 0.3-0.6 cm or wide 0.6-1.0+ cm rings), exhibiting compressed structure (limited radial extent indicating reduced xylem production during growing season), displaying thin appearance (narrow band visible in cross-section much thinner than adjacent wider rings), demonstrating limited growth (narrow ring representing minimal diameter increase during that year), and showing relative narrowness (ring considered narrow in comparison to trunk's typical ring widths with interpretation depending on context). Narrow rootstock annual ring formation causes includes= resulting from water stress (drought or inadequate irrigation limiting cell division and expansion producing narrow ring), reflecting nutrient deficiency (inadequate nitrogen, phosphorus, or other nutrients constraining cambial activity and xylem production), showing temperature extremes (excessive heat, unusual cold, or short growing season limiting growth period), indicating pest or disease damage (defoliation, trunk damage, or systemic infection reducing photosynthate supply or cambial function), demonstrating age-related decline (old trunk showing narrow rings from progressively slowing growth rate characteristic of advanced age), exhibiting low inherent vigor (low-vigor rootstock cultivar naturally producing narrower rings than vigorous cultivar under similar conditions), and revealing competitive stress (inter-vine competition, weed pressure, or shade reducing resources available for growth). Narrow rootstock annual ring structural characteristics includes= showing compressed earlywood zone (reduced spring wood formation with limited large vessel production potentially 30-40% of normal earlywood width), exhibiting reduced latewood zone (minimal late season growth with thin latewood band potentially 30-40% of normal latewood width), displaying fewer cells (limited number of xylem cells produced across ring indicating suppressed cambial division activity), containing smaller vessels (vessels potentially smaller than typical for ring position reflecting stress effects on cell expansion), demonstrating reduced cell enlargement (cells not achieving normal dimensions due to limited water or resources during formation), and showing early growth cessation (cambium potentially entering dormancy earlier than normal producing narrow ring from shortened growth period). Narrow rootstock annual ring in trunk age context includes= common in old trunk (narrow rings typical in aged trunk years 40-80+ reflecting age-related growth decline as normal phenomenon), occasional in mature trunk (narrow rings in mature trunk years 15-40 typically indicating specific stress year rather than normal pattern), unusual in young trunk (narrow rings in young trunk years 1-10 typically indicating significant stress or poor conditions since young trunks normally vigorous), requiring age-adjusted interpretation (narrow ring width considered differently depending on trunk age with 0.2 cm possibly normal in old trunk but concerning in young trunk), and demonstrating progressive narrowing (sequential narrow rings in aging trunk showing expected decline versus isolated narrow ring indicating specific stress event). Narrow rootstock annual ring water stress signature includes= indicating drought year (narrow ring typical signature of water-limited growing season with insufficient soil moisture), showing irrigation deficit (narrow ring potentially indicating inadequate irrigation failing to meet vine water needs), exhibiting moisture stress timing (particularly narrow earlywood suggesting early season drought while reduced latewood indicating late season stress), demonstrating stress severity (extremely narrow ring suggesting severe drought while moderately narrow ring indicating moderate water limitation), and revealing recovery patterns (return to wider ring following narrow ring indicating water stress relief and recovery). Narrow rootstock annual ring and trunk capacity includes= reducing trunk diameter increment (narrow ring adding minimal radial growth potentially only 0.1-0.2 cm diameter increase that year), limiting vessel addition (narrow ring producing fewer new vessels reducing addition to trunk water transport capacity), constraining capacity expansion (repeated narrow rings limiting trunk functional capacity growth particularly problematic if canopy or crop increasing), potentially indicating inadequate growth (persistent narrow rings suggesting trunk not adequately expanding to support vine needs), and requiring assessment context (occasional narrow ring acceptable reflecting specific stress year while persistent narrow rings concerning indicating chronic limitation). Narrow rootstock annual ring in different rootstock cultivars includes= relative to cultivar vigor (narrow defined relative to rootstock typical width with Riparia showing 0.2-0.3 cm possibly normal while 1103P showing 0.2-0.3 cm indicating severe stress), showing cultivar-specific patterns (low-vigor rootstocks naturally producing narrower rings than vigorous rootstocks requiring cultivar-appropriate interpretation), exhibiting different stress sensitivity (some rootstock varieties producing particularly narrow rings under stress while others maintaining relatively wider rings showing differential stress response), demonstrating genetic growth rates (narrow ring assessment requiring knowledge of rootstock variety typical ring width patterns), and revealing adaptation indicators (persistent narrow rings potentially indicating poor rootstock-site match with cultivar unable to thrive under local conditions). Narrow rootstock annual ring diagnostic value includes= indicating stress year (isolated narrow ring marking specific growing season with unfavorable conditions), revealing chronic stress (series of consecutive narrow rings indicating persistent limitation from site, management, or vine health issues), showing age-related decline (progressively narrowing rings in old trunk indicating normal aging process approaching end of vigorous growth phase), demonstrating management effects (narrow rings potentially revealing inadequate irrigation, fertility, or other management practices), enabling stress event dating (narrow rings associated with known events such as drought years, freeze damage, or pest outbreaks providing temporal markers), and supporting vine evaluation (narrow ring patterns informing assessment of vine health, vigor, and suitability for continued production). Narrow rootstock annual ring earlywood-latewood proportions includes= showing relatively more latewood (narrow ring often showing proportionally greater latewood due to early growth cessation limiting earlywood expansion), exhibiting compressed zones (both earlywood and latewood reduced but earlywood potentially more severely compressed), displaying reduced vessel area (limited large earlywood vessel production potentially constraining water transport capacity addition that year), demonstrating stress-adapted structure (narrow ring structure reflecting vine prioritizing survival and tissue quality over growth quantity under stress), and revealing formation timing (narrow ring structure patterns potentially indicating when during season stress occurred affecting earlywood versus latewood differently). Narrow rootstock annual ring in growth rate assessment includes= indicating slow growth period (narrow ring marking year of minimal trunk diameter expansion), showing reduced vigor year (narrow ring suggesting reduced overall vine vigor during that growing season from stress or resource limitation), demonstrating growth constraint (narrow ring revealing factors limiting cambial activity and xylem production that year), enabling growth rate quantification (narrow ring width providing precise measurement of annual growth rate for that year), and supporting long-term analysis (patterns of narrow and wider rings revealing growth rate trends and variations over trunk life). Narrow rootstock annual ring functional implications includes= reducing water transport capacity addition (narrow ring producing fewer vessels adding less to trunk hydraulic conductivity than typical ring), limiting structural support addition (minimal wood added providing less contribution to trunk mechanical strength than wider ring), constraining carbohydrate storage expansion (limited ray parenchyma added in narrow ring reducing storage capacity increase), demonstrating functional adequacy assessment (persistent narrow rings potentially indicating trunk capacity expansion not keeping pace with vine needs), and requiring compensation evaluation (assessing whether existing sapwood adequate despite narrow rings or whether capacity constrained by limited growth). Narrow rootstock annual ring in vineyard management includes= signaling management issues (narrow rings potentially indicating inadequate irrigation, fertility, or other cultural practice deficiencies requiring correction), revealing stress impacts (narrow rings documenting effects of drought, pest pressure, or other stresses on vine providing feedback for management decisions), supporting intervention timing (narrow ring patterns helping determine when problems occurred guiding remedial action timing), enabling treatment evaluation (comparing ring widths before and after management changes assessing treatment effectiveness), and guiding future practices (narrow ring analysis informing irrigation scheduling, fertilization programs, or other management modifications). Narrow rootstock annual ring in old vine context includes= expected in aged trunk (narrow rings normal in old trunk years 60-100+ reflecting natural age-related growth decline), indicating aging progression (progressively narrowing rings marking transition from mature to old trunk phase), demonstrating vitality assessment (extremely narrow rings even for old trunk potentially indicating declining vine health beyond normal aging), revealing productivity relationship (very narrow rings potentially correlating with reduced yields though old vine fruit quality often excellent despite narrow growth), and supporting longevity evaluation (persistent extremely narrow rings possibly indicating vine approaching end of productive life requiring replacement consideration). Narrow rootstock annual ring multiple consecutive patterns includes= indicating chronic stress (series of narrow rings suggesting persistent unfavorable conditions rather than isolated event), revealing progressive decline (sequentially narrowing rings showing progressive worsening of conditions or vine health), demonstrating recovery absence (continued narrow rings showing vine not recovering from initial stress indicating persistent limitation), suggesting fundamental limitation (multiple consecutive narrow rings potentially indicating poor site conditions, disease, or rootstock-scion incompatibility), and requiring intervention (persistent narrow rings signaling need for management changes, remedial treatments, or replacement consideration). Narrow rootstock annual ring and climate reconstruction includes= marking drought years (narrow rings corresponding to documented drought years providing dendrochronological drought record), indicating unfavorable climate periods (sequences of narrow rings revealing extended periods of poor growing conditions), enabling cross-dating (matching narrow ring patterns across multiple trunks or sites synchronizing chronologies), supporting climate analysis (narrow ring frequencies and patterns revealing historical climate variations and trends), and documenting extreme events (particularly narrow rings marking exceptional stress years such as severe droughts or freeze damage events). Narrow rootstock annual ring measurement and analysis includes= measuring precise width (determining exact radial dimension typically 0.1-0.3 cm for narrow ring using microscope or digital imaging), comparing to trunk average (assessing narrowness relative to trunk's typical ring width determining deviation from normal), analyzing in context (interpreting narrow ring considering trunk age, rootstock variety, and known environmental or management history), examining anatomical details (studying earlywood-latewood structure, vessel characteristics, and cell dimensions in narrow ring revealing formation conditions), and integrating with records (correlating narrow rings with weather data, management records, or known stress events determining causes). Narrow rootstock annual ring in trunk core sampling includes= visible in increment cores (narrow rings identifiable in extracted core samples enabling non-destructive narrow ring detection and analysis), enabling age determination (narrow rings counted equally with wider rings for accurate age assessment though visibility potentially reduced), supporting growth analysis (narrow ring widths measured in cores revealing annual growth rates and stress year identification), facilitating stress chronology (narrow ring patterns in cores documenting stress history without felling trunk), and allowing repeated sampling (cores extracted multiple years monitoring whether narrow ring patterns improving, stable, or worsening over time). Narrow rootstock annual ring significance includes= indicating reduced annual growth (narrow ring representing minimal secondary xylem production and trunk diameter increase that year), revealing unfavorable conditions (narrow ring documenting water stress, nutrient limitation, temperature extremes, pest damage, or age effects during formation year), demonstrating stress response (narrow ring showing cambial activity suppression as physiological response to limiting factors), enabling stress event identification (narrow ring marking specific year of unfavorable conditions allowing temporal stress documentation), supporting vine health assessment (narrow ring patterns revealing vigor trends, stress frequency, and overall trunk growth trajectory), limiting functional capacity expansion (narrow ring adding minimal water transport, structural support, and storage capacity to trunk), and requiring contextual interpretation (narrow ring significance depending on trunk age, rootstock variety, adjacent rings, and environmental history).
Narrow_Rootstock_Annual_Ring emphasizes reduced width less than 0.3 cm and is distinguished from Rootstock_Annual_Ring (typical width) and Wide_Rootstock_Annual_Ring (increased width) by focus on constrained growth reflecting stress, limitations, or age effects. Critical for understanding growth limitations, stress indicators, water deficit signals, age-related decline, diagnostic value, trunk capacity constraints, and the narrow rings documenting unfavorable conditions or physiological limitations during formation year.
Dünner Wurzelstock-Wachstumsring
Reduced rootstock annual ring
Schmaler Unterlagen-Jahresring
Thin rootstock annual ring
Narrow Rootstock Annual Ring
Blaufränkisch accession HU-NAIK-0412 at the National Centre for Biodiversity and Gene Conservation (NBGK) in Tápiószele, Hungary bearing a national collection role is maintained as part of the Hungarian National Grapevine Collection comprising approximately 1,600 accessions representing all historically documented Hungarian cultivars and clones, managed under the supervision of the Hungarian Ministry of Agriculture with annual state funding for curatorial staff, phytosanitary monitoring, SSR-based identity verification of 200 accessions per year, and digitized passport data accessible through the Hungarian Plant Genetic Resources Database linked to the EURISCO European germplasm search catalogue.
A germplasm repository role that inheres in a grapevine maintained within a nationally mandated or nationally recognized grapevine germplasm collection tasked with conserving the genetic diversity of cultivars grown within or historically associated with a single country, typically operated by national research institutes, agricultural ministries, or designated gene bank institutions.
National_Collection_Role reflects the sovereign responsibility for genetic resource conservation codified in the Convention on Biological Diversity (CBD 1992) and the International Treaty on Plant Genetic Resources for Food and Agriculture (ITPGRFA 2001). National grapevine collections are typically mandated to conserve the complete cultivar diversity of the country — including rare, endangered, and historically significant cultivars alongside commercially grown varieties: this comprehensive mandate distinguishes national collections from regional or thematic collections with narrower scope. The institutional stability and long-term funding security of national collections varies considerably across countries, creating conservation risks for genetic resources in politically or economically unstable regions.
national genebank role
national vine collection role
nationale Genbank-Rolle
nationale Sammlungs-Rolle
staatliche Rebsorten-Sammlung-Rolle
state germplasm collection role
FAO (2010) Second Report on the State of the World's Plant Genetic Resources for Food and Agriculture, Rome ITPGRFA (2001) International Treaty on Plant Genetic Resources for Food and Agriculture, FAO EURISCO (2023) European Search Catalogue for Plant Genetic Resources, ECPGR
National Collection Role
Thompson Seedless bears a Natural_Raisin_Scion_Role as the archetypal natural raisin variety, with grapes harvested at 22-24°Brix in late August-September in California, laid on paper trays between vine rows, and sun-dried for 14-21 days to produce the standard "California Natural Seedless Raisins" of commerce—dark brown color, chewy texture, caramel-fruit flavor. Fiesta serves in a Natural_Raisin_Scion_Role with characteristics similar to Thompson Seedless, offering earlier maturity (by approximately one week) that reduces weather risk during the critical drying period. Selma Pete bears a Natural_Raisin_Scion_Role as a DOV (dried-on-vine) variety specifically bred for mechanical harvesting of raisins dried on continuous trays without hand-laying, improving labor efficiency in natural raisin production.
A raisin production scion role that inheres in a grafted scion cultivar by virtue of its suitability for natural (sun-dried) raisin production without chemical pre-treatment, where grapes are dried on paper trays between vine rows or on continuous drying trays under ambient sunlight and air circulation. Natural raisin scions must achieve adequate drying within the available weather window (typically 14-21 days in California) without chemical dipping treatments that accelerate moisture loss, requiring thin, permeable skins, appropriate berry size (not too large), and tolerance of the oxidative browning that produces characteristic dark color. Natural sun-drying produces the familiar dark brown "natural" raisins through enzymatic browning (polyphenol oxidase activity) and non-enzymatic Maillard reactions between sugars and amino acids, developing caramelized, slightly tannic flavor profiles distinct from golden (sulfured) raisins. Natural raisin production is the traditional and dominant method globally, concentrated in hot, dry production regions (California's San Joaquin Valley, Turkey's Aegean coast, Australia's Murray-Darling) where autumn weather permits extended outdoor drying.
Natural_Raisin_Scion_Role captures the traditional sun-drying method that remains dominant for dark raisin production despite higher labor requirements than mechanized alternatives. The distinction from Golden_Raisin_Scion_Role relates to processing method (sulfur treatment prevents browning) rather than genetics—Thompson Seedless serves in both natural and golden raisin roles. DOV (dried-on-vine) systems represent technological evolution within natural raisin production, severing canes to initiate on-vine drying before mechanical harvest, reducing the labor-intensive tray-laying process. Climate change affects natural raisin production by altering the reliability of hot, dry autumn weather windows essential for successful outdoor drying without mold development.
Natur-Rosinen-Traube
Thompson raisin grape
dark raisin variety
dunkle Rosinentraube
natural raisin grape
sonnengetrocknete Rosinentraube
sun-dried raisin grape
Christensen LP (2000) Raisin Production Manual, UC ANR Publication 3393 Fidelibus MW et al. (2008) Dried-on-vine raisin production, UC ANR Publication 8279 Pangavhane DR et al. (1999) Solar dryers for grape drying, Energy Conversion and Management 40: 1-23 California Raisin Marketing Board (2020) Raisin Production Statistics
Natural Raisin Scion Role
Annual NDVI uniformity quality monitoring in a 40-hectare Hawke's Bay vineyard operation employed February satellite imagery at 3-meter resolution to assess vigor consistency across 12 Chardonnay and Merlot blocks. Blocks exhibiting NDVI coefficient of variation exceeding 20% were flagged for investigation, revealing irrigation pressure issues in two blocks and phylloxera progression in one block. Tracking NDVI uniformity quality over five vintages demonstrated improvement from mean block CV of 22.3% to 16.8% following infrastructure upgrades and targeted replanting, with corresponding reduction in harvest segregation requirements from average 3.2 picks per block to 2.1 picks per block while maintaining fruit quality tier differentiation.
NDVI uniformity quality is a vigor uniformity quality that inheres in a vineyard spatial unit and represents the consistency of normalized difference vegetation index values across the assessed area as derived from remotely sensed multispectral imagery capturing spatial variation in canopy chlorophyll density, leaf area, and vegetative vigor through the differential reflectance of near-infrared and visible red wavelengths by photosynthetically active tissue. This quality provides efficient, comprehensive, and spatially continuous characterization of vigor uniformity that complements or replaces labor-intensive ground-based sampling approaches for vineyard-scale assessment. The ontological basis of NDVI uniformity quality rests on the established relationship between NDVI values and grapevine vegetative characteristics including leaf area index, chlorophyll content, and biomass, enabling remote sensing platforms to capture vigor variation across entire vineyard blocks at spatial resolutions sufficient to identify management-relevant patterns. The NDVI calculation normalizes the difference between near-infrared reflectance, which is strongly reflected by healthy vegetation, and red reflectance, which is absorbed by chlorophyll, producing index values ranging from -1 to 1 with typical healthy grapevine canopies yielding values between 0.4 and 0.8 depending on canopy density and sensor characteristics. Uniformity quantification applies statistical dispersion metrics to spatially referenced NDVI data, with coefficient of variation serving as the standard expression enabling comparison across blocks, vintages, and regions. High NDVI uniformity quality indicates consistent canopy development across the assessed unit, while spatial patterns in NDVI non-uniformity can reveal underlying drivers including soil variation, irrigation system performance, disease distribution, and rootstock effects in mixed plantings. The temporal dimension of NDVI uniformity quality requires consideration of acquisition timing relative to phenological development, with post-véraison imagery after canopy stabilization but before senescence providing the most reliable vigor characterization for management applications. Sensor platforms for NDVI uniformity assessment range from satellite systems offering broad coverage at moderate resolution to aircraft and unmanned aerial vehicles providing higher resolution for detailed within-block characterization, with platform selection depending on assessment objectives, block size, and cost considerations. Validation of NDVI-derived uniformity assessments against ground-truth vigor measurements including pruning weights and yield data strengthens interpretation confidence and calibrates management zone delineation.
NDVI_Uniformity_Quality is_a Vigor_Uniformity_Quality, representing the remote sensing-based approach to uniformity assessment using normalized difference vegetation index. This class specifically addresses the spectral index methodology rather than ground-based measurements, though NDVI-derived patterns typically correlate with pruning weight and yield variation. Distinguished from Block_Vigor_Uniformity_Quality by the specific measurement technology, though NDVI provides efficient means for block-level uniformity assessment. Temporal considerations including acquisition timing and multi-year monitoring are important aspects of this quality class.
NDVI spatial consistency
NDVI-Gleichmäßigkeit
Vegetationsindex-Uniformität
fernerkundete Wuchsgleichmäßigkeit
remote sensing vigor uniformity
spectral uniformity index
spektrale Uniformität
vegetation index uniformity
Hall A et al. (2002) Characterising and mapping vineyard canopy using high-spatial-resolution aerial multispectral images. Computers and Geosciences 28:813-822 Johnson LF et al. (2003) Mapping vineyard leaf area with multispectral satellite imagery. Computers and Electronics in Agriculture 38:33-44 Bramley RGV et al. (2011) Variation in vine vigour, grape yield and vineyard soils and topography as indicators of variation in the chemical composition of grapes, wine and wine sensory attributes. Australian Journal of Grape and Wine Research 17:217-229
NDVI Uniformity Quality
Comparative analysis of Near_Isohydric_Behavior_Disposition at Bordeaux Sciences Agro positioned 30 cultivars along the isohydric-anisohydric continuum using standardized drought stress protocols and quantitative stomatal behavior parameters. The isohydricity index (σ, slope of midday versus predawn water potential regression) was calculated for each cultivar, with σ = 0 indicating perfect isohydric behavior and σ = 1 indicating fully anisohydric behavior. Near-isohydric cultivars were defined as σ between 0.3 and 0.7. Of the 30 cultivars evaluated, 18 (60%) showed Near_Isohydric_Behavior_Disposition with σ ranging from 0.32 ('Merlot') to 0.68 ('Cabernet Sauvignon'). Near-isohydric 'Merlot' allowed Ψmd to decline from -1.1 MPa (well-watered) to -1.6 MPa (moderate drought) before stabilizing through stomatal closure, showing intermediate response between isohydric 'Grenache' (σ = 0.15, Ψmd stable at -1.3 MPa) and anisohydric 'Syrah' (σ = 0.85, Ψmd declining to -2.0 MPa). Productivity analysis revealed that near-isohydric cultivars achieved intermediate seasonal carbon gain, capturing 75-85% of anisohydric productivity while maintaining hydraulic safety margins (PLC below 20% under moderate drought). Notably, some cultivars showed environmentally plastic Near_Isohydric_Behavior_Disposition, with 'Cabernet Sauvignon' (σ = 0.68 in maritime climate) behaving more isohydric (σ = 0.42) when grown in continental climate with greater diurnal temperature variation, suggesting acclimation of stomatal regulation strategy. These findings indicate that Near_Isohydric_Behavior_Disposition represents both a genetic trait and an adaptive response, providing flexibility across diverse growing conditions.
A Near_Isohydric_Behavior_Disposition is a Stomatal_Regulation_Disposition that inheres in a Vitis vinifera plant and is realized through intermediate stomatal control that partially regulates midday leaf water potential, allowing moderate decline under drought while preventing the extreme water potential reductions characteristic of anisohydric behavior. This disposition represents the middle ground of the stomatal regulation spectrum, where cultivars show partial water potential homeostasis that balances hydraulic safety against continued productivity. Near-isohydric behavior involves moderate stomatal sensitivity that permits continued gas exchange during early drought phases while triggering progressive closure as stress intensifies, resulting in leaf water potential that declines but stabilizes at a regulated minimum rather than tracking soil water potential indefinitely. The physiological basis of Near_Isohydric_Behavior_Disposition likely involves intermediate ABA sensitivity and/or hydraulic signal integration that produces graded rather than all-or-nothing stomatal responses. The adaptive value of this intermediate strategy lies in balancing the competing demands of water conservation and carbon gain, avoiding both the productivity costs of strict isohydric regulation and the hydraulic risks of anisohydric behavior. Many commercially important cultivars display near-isohydric characteristics, enabling reasonable productivity across varied environments while maintaining adequate hydraulic safety margins. This intermediate behavior also shows greater plasticity, with some cultivars adjusting their position on the isohydric-anisohydric spectrum based on environmental conditions or stress history.
Near_Isohydric_Behavior_Disposition is_a Stomatal_Regulation_Disposition. Near-isohydric behavior represents the middle of the stomatal regulation continuum, with partial water potential regulation. Many commercial cultivars show near-isohydric behavior, balancing productivity and safety. The isohydricity index (σ) provides quantitative positioning along the continuum. Some cultivars show plasticity in stomatal behavior depending on environment or stress history. Near-isohydric cultivars may be broadly adapted across diverse environments.
Intermediäre-Stomataregulation
Nah-isohydrisches-Verhalten-Disposition
Partielle-Wasserpotential-Regulation
intermediate stomatal behavior
moderate water conservation
partial water potential regulation
Martínez-Vilalta J et al. (2014) A new look at water transport regulation in plants, New Phytologist 204:105-115 Hochberg U et al. (2018) Iso/anisohydry: a plant-environment interaction rather than a simple hydraulic trait, Trends in Plant Science 23:112-120 Lavoie-Lamoureux A et al. (2017) Factors influencing stomatal conductance in response to water availability in grapevine: a meta-analysis, Physiologia Plantarum 159:468-482
Near Isohydric Behavior Disposition
A near-mature aborted tendril at node 10 on a shoot of Vitis vinifera cv. Syrah in the Rhône Valley, France, measuring 10.5 cm length (approximately 65% of the normal 16 cm for functional tendrils at this position), exhibiting complete bifurcated structure with well-developed arms, dark green epidermis with substantial cuticle evident from glossy surface, visible mechanosensory cells on tendril surfaces, moderate lignification in basal 4 cm showing light brown coloration, but exhibiting tip necrosis extending 2 cm from each branch tip and progressive desiccation following an acute three-day heat spike (maximum temperatures 41-43°C) during the final week of tendril development in mid-July, occurring in a pattern affecting 40% of tendrils in similar developmental stages across heat-exposed vineyard sections while earlier-developed tendrils and later-developing tendrils (before and after the heat event) achieved full functionality, representing a temporally discrete cohort of near-complete development failures caused by acute late-stage stress.
A late aborted tendril that achieved substantial developmental progress approaching functional maturity before developmental arrest in Vitis vinifera, characterized by structures of 8-12 cm length representing 60-75% of normal mature tendril length, complete tissue differentiation including fully developed vascular system with functional xylem and phloem, well-formed epidermis with extensive cuticle deposition, recognizable mechanosensory apparatus with differentiated contact-responsive cells, and substantial structural reinforcement through cell wall thickening and initial lignification, but ultimately failing to complete final elongation to functional length, achieve full physiological activation of sensory systems, or develop adequate structural strength for reliable load-bearing function. Near-mature aborted tendrils result from acute stress events or sudden resource limitations occurring late in the development process when basic morphogenesis and tissue differentiation are essentially complete but final maturation processes remain unfinished, typically caused by sudden severe water stress, acute heat or cold stress during the final maturation week, sudden pathogen infection or pest attack, abrupt competition from rapidly developing fruit clusters during early berry expansion, or mechanical damage during late development that interrupts vascular function or causes partial tissue destruction. These abortion events represent maximum resource investment loss among abortion categories as extensive growth and differentiation have occurred, tissues have consumed substantial carbohydrates and nutrients, and the vine has committed resources almost sufficient for functional return but fails to achieve actual climbing utility from the investment.
Near-mature aborted tendrils are distinguished from mid-development aborted tendrils by achievement of >60% normal length and near-complete tissue differentiation, and from functional tendrils by failure to complete final maturation despite substantial development. This abortion class is particularly frustrating from resource allocation perspective because the vine has invested nearly all resources required for functional tendril production but fails to achieve return on investment, representing inefficiency in carbon and nutrient utilization. Near-mature abortion is diagnostically significant because it indicates acute stress events during final maturation rather than chronic resource limitation affecting entire development, with temporal clustering of near-mature abortion at specific node positions revealing discrete stress events (heat spikes, cold snaps, pest outbreaks) that can be correlated with weather records or management events. Prevention requires stress amelioration specifically during the final 3-7 days of tendril development, a narrow intervention window.
Endstadium abgebrochene Kletterorgan
almost mature arrested tendril
fast-reife abortierte Ranke
final stage aborted tendril
nahezu vollständige fehlgeschlagene Ranke
nearly complete failed tendril
Based on Greer & Weston (2010) "Heat stress affects flowering, berry growth, sugar accumulation and photosynthesis of Vitis vinifera cv. Semillon grapevines grown in a controlled environment" Functional Plant Biology 37=206-214 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 86-108 Petrie et al. (2000) "The effect of leaf removal and canopy height on whole-vine gas exchange and fruit development of Vitis vinifera L. Sauvignon blanc" Functional Plant Biology 27=173-184 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 449-472.
Near Mature Aborted Tendril
Pinot Noir showing near-surface lateral roots, excavation revealing laterals 6-12 cm depth in upper topsoil, near-surface laterals 1.2-2.8 cm diameter, some roots partially exposed where soil eroded on slope, abundant fine root production spring showing 12.4 cm feeder root per cm³ soil, near-surface fine roots largely senesced by late summer as topsoil dried despite deeper roots remaining active, near-surface lateral roots demonstrating opportunistic boom-bust pattern responding to extreme environmental variation
Near surface lateral root is surface lateral root at uppermost position typically 5-15 cm depth immediately below soil surface, characterized by extreme proximity to surface creating both opportunities and vulnerabilities, position in zone of maximum temperature fluctuation and moisture variability, frequent exposure through erosion or cultivation, and serving opportunistic function rapidly exploiting surface resources when available but vulnerable to stress and disturbance, with near-surface laterals showing characteristics including location in upper A horizon often in zone of active organic matter accumulation, shallow enough to sometimes become exposed at surface particularly on slopes or through erosion, moderate diameter 0.8-3 cm, high fine root production density when conditions favorable, extreme seasonal and diurnal environmental variation affecting function, and dual character as highly productive when conditions favorable but quickly stressed or damaged when conditions unfavorable creating boom-bust dynamics. Near surface lateral root depth and exposure includes= showing very shallow depth (near-surface laterals 5-15 cm depth at upper extreme of root distribution), exhibiting occasional exposure (near-surface roots sometimes exposed at soil surface through erosion, cultivation, or natural soil processes), demonstrating surface proximity (roots close enough to surface potentially visible in shallow excavations or when soil cracks), revealing slope effects (near-surface laterals more commonly exposed on slopes where erosion more active), and displaying management visibility (near-surface roots readily observable during vineyard operations providing feedback on root system status). Near surface lateral root extreme environmental variation includes= showing maximum temperature fluctuation (near-surface zone experiencing largest diurnal and seasonal temperature changes affecting root metabolism), exhibiting rapid moisture changes (near-surface soil wetting quickly with rain or irrigation but also drying rapidly through evaporation), demonstrating frost susceptibility (near-surface laterals potentially freezing in cold weather particularly without snow cover or mulch), revealing heat exposure (near-surface roots potentially experiencing damaging heat in summer without irrigation or mulch protection), and displaying nutrient pulses (near-surface zone showing most dramatic nutrient concentration changes following fertilization or mineralization events). Near surface lateral root opportunistic function includes= showing rapid response (near-surface laterals quickly exploiting favorable conditions when topsoil moist and fertile), exhibiting intensive production (near-surface roots producing abundant fine roots and root hairs when conditions optimal), demonstrating resource capture (near-surface laterals capturing surface-applied water and nutrients before deeper roots or losses), revealing competitive advantage (near-surface position allowing first access to rainfall, irrigation, or fertilizers), and displaying temporal dynamics (near-surface root function showing large temporal variation rapidly increasing under favorable conditions quickly declining under stress). Near surface lateral root vulnerability includes= showing maximum tillage damage (near-surface laterals most vulnerable to cultivation damage being in zone of deepest tillage penetration), exhibiting erosion exposure (soil erosion readily exposing or removing near-surface laterals particularly on slopes), demonstrating drought sensitivity (near-surface laterals quickly experiencing drought stress as topsoil dries given limited soil volume accessed), revealing frost damage (near-surface roots potentially killed by severe freezing particularly in exposed situations), and displaying compaction susceptibility (near-surface laterals easily compacted by traffic or equipment given shallow position). Near surface lateral root in fertilizer capture includes= providing immediate access (near-surface laterals positioned optimally for capturing broadcast or shallow-incorporated fertilizers), exhibiting rapid uptake (near-surface fine roots quickly absorbing fertilizer nutrients before leaching or immobilization), demonstrating efficiency advantage (near-surface position reducing fertilizer losses through placement close to primary absorption zone), revealing management opportunity (fertilizer application optimized for near-surface lateral distribution potentially improving efficiency), and displaying response intensity (near-surface laterals showing strong proliferation response to fertilizer placement). Near surface lateral root seasonal patterns includes= showing spring maximum (near-surface lateral activity peaking spring when topsoil moist, warming, and nutrient-rich), exhibiting early senescence (near-surface fine roots potentially senescing earlier summer as topsoil dries compared to deeper roots), demonstrating fall resurgence (near-surface root activity recovering fall with rainfall and cooling temperatures), revealing winter vulnerability (near-surface laterals potentially damaged by freezing requiring spring recovery), and displaying annual replacement (substantial near-surface lateral turnover requiring continuous production replacing stressed or damaged roots). Near surface lateral root management strategies includes= requiring surface protection (mulches, cover crops, or other practices protecting near-surface laterals from environmental extremes and disturbance), showing irrigation dependency (near-surface laterals requiring consistent irrigation maintaining function through summer), exhibiting no-till benefits (avoiding tillage preventing near-surface lateral damage maintaining absorption capacity), demonstrating fertilizer placement importance (placing fertilizers where near-surface laterals efficiently access optimizing uptake), and revealing monitoring value (near-surface lateral condition potentially indicating overall root system and vine health status). Near surface lateral root functional importance includes= determining opportunistic uptake (near-surface laterals providing rapid response absorption when conditions favorable), enabling efficient capture (near-surface position allowing efficient capture of surface-applied resources), demonstrating stress indication (near-surface lateral condition indicating environmental stress levels affecting vine), revealing production potential (abundant near-surface laterals indicating favorable conditions and high productivity potential), and displaying management sensitivity (near-surface lateral response indicating effectiveness of irrigation, fertilization, and soil management practices).
Near_Surface_Lateral_Root emphasizes uppermost extremely shallow position and is distinguished from deeper surface roots by focus on surface proximity. Critical for understanding extreme environmental variation, exposure risk, opportunistic function, maximum vulnerability, fertilizer capture, and the uppermost laterals showing boom-bust dynamics responding to variable topsoil conditions.
Oberflächen-nahe Wurzel
Oberste Seitenwurzel
Sehr flache Lateralwurzel
Surface-exposed root
Uppermost lateral root
Very shallow lateral
Near Surface Lateral Root
Chardonnay in Champagne shows near-zero growth rate during late August-September Year N+1 at BBCH 81, with berry growth rate sustained at 1-4 mg per day over 2.5 week period showing daily variation including 8 days positive 2-4 mg per day, 6 days zero growth, 3 days slightly negative -1 to 0 mg per day from water loss, overall average 2.1 mg per day, berry dimensions stable at 8.3-8.5 mm diameter and 2.9-3.1 g fresh weight representing 2.8% variation, under cool maritime conditions of 18-24°C temperature, typical of this early-bursting cultivar in lag quiescence
Near-zero growth rate represents the essential growth cessation state during minimal growth period when berry fresh weight gain sustained at very low levels 1-5 mg per day, occasional slightly negative growth from water loss, growth effectively arrested distinguishing lag from growth and ripening phases, and minimal rate maintained over 1-3 weeks enabling metabolic focus on preparation rather than growth, characteristic of lag phase quiescence. This growth cessation state occurs during late summer to early autumn in late August-September in temperate regions Year N+1, sustained over 1-3 weeks representing main lag duration, corresponding to BBCH 81 beginning of ripening stage quiescent period. Near-zero rate mechanism involves cellular growth processes effectively silenced, minimal water uptake approaching uptake-transpiration equilibrium, berry structural components stable without net accumulation, metabolic activity focused on maintenance and preparation not growth, developmental program maintaining quiescent state, and integrated cessation creating growth arrest. Growth rate quantification shows typical berry fresh weight gain 1-5 mg per day over minimal growth period, occasional days with zero or slightly negative growth -1 to 0 mg per day from water loss, average rate 2-3 mg per day characteristic of lag quiescence, and minimal rate contrasting with enlargement 50-250 mg per day and ripening 20-100 mg per day. Growth rate temporal pattern shows day-to-day variation in minimal rates from environmental fluctuations, some days positive 3-5 mg per day from water uptake, other days zero or negative from transpiration exceeding import, overall sustained near-zero rate averaging 2-3 mg per day, and temporal dynamics characteristic of equilibrium state. Berry fresh weight stability shows weight remaining essentially constant 2-5% total variation over 1-3 week period, minor fluctuations from water balance dynamics, overall stability contrasting with systematic increase during growth and ripening, and constancy as lag defining feature. Volumetric growth cessation shows berry volume unchanging <1% over minimal growth period, no net cellular expansion, no tissue accumulation, and volumetric arrest characteristic of quiescence. Dimensional stability shows berry diameter constant within 1-3% measurement and fluctuation range, length and width stable, three-dimensional berry shape maintained, and dimensional constancy indicating growth cessation. Water flux minimal characteristics shows water uptake very low <0.05 g per berry per day, transpiration water loss approximately 0.03-0.06 g per day depending on environmental conditions, net water flux near-zero -0.02 to +0.02 g per day showing equilibrium, and minimal flux enabling weight stability. Water uptake-transpiration balance shows uptake approximately equaling loss over minimal growth period, balance creating stable berry hydration, equilibrium preventing net accumulation or depletion, and balance as quiescence mechanism. Water content stability shows berry water content 78-82% maintained over minimal growth period, minor fluctuations 1-3% from daily water balance variation, overall stability characteristic of equilibrium state, and constant hydration maintaining berry composition. Cellular processes during near-zero growth shows cell expansion completely ceased, cells at final mature dimensions 80-200 micrometers, no cellular growth at tissue or organism level, and cellular quiescence underlying berry growth arrest. Cell division absent shows no cell proliferation during lag phase, cell number fixed from division phase completion, cell populations stable, and absence of division characteristic of differentiated quiescent tissue. Biosynthetic activity minimal shows structural biosynthesis ceased, protein synthesis limited to maintenance turnover, minimal lipid and carbohydrate biosynthesis, and biosynthetic quiescence characteristic of non-growing state. Metabolic focus on maintenance shows metabolism supporting existing structures not growth, respiratory ATP production for maintenance processes, protein and membrane turnover maintaining cellular integrity, and maintenance metabolism adequate for berry survival without growth. Metabolic focus on preparation shows metabolic activity directed toward ripening preparation not growth, ripening enzyme synthesis accumulating capacity, metabolic pathway components assembling, and preparation as active metabolic function during growth quiescence. Berry composition near-stability shows water, acids, sugars showing slow progressive changes not rapid accumulation, composition trajectory toward ripening characteristics, changes from metabolism not growth, and near-stability distinguishing lag from dynamic growth and ripening. Organic acid slow decline shows malate declining 0.05-0.10% per week from respiration, slow acid loss not requiring growth, decline contributing to pH increase and softening, and slow acid dynamics characteristic of quiescent metabolism. Sugar slow accumulation shows sugars increasing 0.3-0.8% per week from limited phloem import, slow accumulation not requiring growth, sugar increase preparing for ripening acceleration, and limited dynamics characteristic of quiescent import. Developmental program maintenance shows quiescent state actively maintained by developmental programming, growth suppression signals preventing premature ripening, eventual developmental progression to veraison, and programmed quiescence as regulated developmental state. Environmental effects on near-zero growth include temperature affecting water balance through transpiration effects, warm conditions potentially increasing transpiration creating slightly negative growth, cool conditions reducing transpiration enabling slightly positive growth, water availability affecting uptake and balance, and environmental factors influencing growth rate variation around near-zero. Temperature effects on balance shows warm conditions increasing transpiration potentially creating negative growth days, cool conditions reducing transpiration enabling positive growth days, and temperature affecting water balance dynamics. Water availability effects shows adequate water enabling positive growth days from net uptake, deficit limiting uptake creating neutral or negative growth days, and water as factor affecting day-to-day growth variation. Near-zero rate significance shows growth cessation enabling metabolic and developmental resources to focus on seed maturation and ripening preparation, quiescence as functional developmental state not growth failure, and minimal growth as defining lag phase characteristic. Near-zero rate assessment shows daily or frequent growth measurements quantifying minimal rates, water flux measurements documenting uptake-transpiration balance, volumetric analysis confirming dimensional stability, and measurements characterizing quiescent state. The process represents first aspect of minimal growth period sustaining growth cessation, preceding seed development focus and pre-veraison metabolism. Duration sustained over 1-3 weeks representing main lag period. Regional timing shows near-zero growth rate during late August-September across climates as sustained quiescence. Viticultural significance includes representing lag defining characteristic of growth cessation, enabling metabolic focus on preparation rather than growth, demonstrating developmental regulation of quiescence, and establishing stable berry state before ripening transformation. Research applications include analyzing growth cessation mechanisms and maintenance, investigating water balance and growth dynamics, evaluating environmental effects on minimal growth rates, quantifying quiescence characteristics and regulation, and assessing near-zero growth as determinant of lag phase function and duration.
Near_Zero_Growth_Rate is first aspect of Minimal_Growth_Period, distinguished by sustained growth cessation. Represents 1-5 mg/day typical, occasional negative from water loss. Berry fresh weight stable within 2-5%. Volumetric growth <1%. Water uptake-transpiration equilibrium. Cell expansion ceased. Biosynthetic activity minimal. Maintenance and preparation metabolism. Duration 1-3 weeks. Followed by Seed_Development_Focus and Pre_Veraison_Metabolism. Defines lag phase growth characteristic.
Nahe-Null-Wachstumsrate
Ruhezustand
Wachstums-Arrest
essential growth cessation
essentielle Wachstums-Beendigung
quiescence state
zero growth maintenance
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 456-459 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Near Zero Growth Rate
Comprehensive evaluation of Nematode_Resistance_Disposition at UC Davis Foundation Plant Services assessed responses to three economically important nematode genera across 40 rootstock genotypes using standardized greenhouse pot assays. Vines were planted in soil inoculated with Xiphinema index (500 individuals), Meloidogyne incognita (5,000 eggs), or Pratylenchus vulnus (2,000 mixed stages) per pot, with nematode populations and root damage assessed at 6 months. Susceptible V. vinifera 'Cabernet Sauvignon' (own-rooted) supported high populations of all three genera: X. index 2,450/pot, M. incognita galling index 8.2/10, P. vulnus 18,500/pot with severe root lesioning. The Muscadinia-derived rootstock 'O39-16' demonstrated broad Nematode_Resistance_Disposition with X. index suppressed to 45/pot (98% reduction), M. incognita galling index 1.2, and P. vulnus limited to 1,200/pot. Importantly, nematode resistance profiles varied among rootstocks, with '1103 Paulsen' showing strong Meloidogyne resistance (galling index 0.8) but moderate Xiphinema susceptibility (890/pot), while 'Freedom' showed inverse pattern (X. index 180/pot but galling index 6.5). These differential resistance profiles guide rootstock selection based on site-specific nematode pressure, with molecular markers enabling targeted breeding for multiple nematode resistances.
A Nematode_Resistance_Disposition is a Pest_Resistance_Disposition that inheres in a Vitis plant root system and is realized through defense processes that limit feeding, reproduction, or damage by plant-parasitic nematodes, microscopic roundworms that attack grapevine roots causing direct injury and, in the case of certain species, transmitting economically important viruses. This disposition represents the capacity to resist a diverse group of soil-dwelling obligate parasites belonging to multiple genera with distinct feeding strategies, host interactions, and economic impacts on viticulture. Major nematode pests of grapevine include ectoparasitic dagger nematodes (Xiphinema species) that feed externally on root tips and transmit nepoviruses including Grapevine fanleaf virus, endoparasitic root-knot nematodes (Meloidogyne species) that induce characteristic galls and disrupt vascular function, and migratory endoparasitic lesion nematodes (Pratylenchus species) that cause cortical necrosis. Nematode resistance mechanisms include pre-infectional barriers preventing nematode penetration or feeding establishment, hypersensitive response killing cells at feeding sites, production of nematicidal or nematostatic compounds, and incompatibility preventing completion of nematode life cycles in host tissues. For grapevine rootstock breeding, nematode resistance serves dual purposes: protecting root system health from direct nematode damage and, critically for Xiphinema, preventing virus acquisition and transmission that would otherwise doom grafted V. vinifera scions to fanleaf decline. Resistance sources have been identified in various Vitis species and Muscadinia rotundifolia, enabling development of nematode-resistant rootstocks for infested vineyard sites.
Nematode_Resistance_Disposition is_a Pest_Resistance_Disposition. This class encompasses resistance to all plant-parasitic nematodes affecting grapevine roots, with child classes organized by nematode genus: Xiphinema_Index_Resistance_Disposition (dagger nematodes, virus vectors), Meloidogyne_Resistance_Disposition (root-knot nematodes), and Pratylenchus_Resistance_Disposition (lesion nematodes). Nematode resistance is primarily relevant for rootstock breeding rather than scion improvement, as roots are the target tissue. The dual importance of Xiphinema resistance for both direct damage prevention and fanleaf virus management distinguishes it from other nematode resistances.
Nematodenresistenz-Disposition
Pflanzenparasitäre-Nematoden-Resistenz
Wurzelnematoden-Abwehr
nematode feeding resistance
plant-parasitic nematode resistance
root nematode immunity
Nicol JM et al. (1999) Current nematode threats to world viticulture, Nematology Monographs and Perspectives 2:269-282 Anwar SA, McKenry MV (2002) Developmental response of a resistance-breaking population of Meloidogyne arenaria on Vitis spp., Journal of Nematology 34:28-33 Zasada IA et al. (2010) Managing nematodes without methyl bromide, Annual Review of Phytopathology 48:311-328
Nematode Resistance Disposition
Ramsey (Salt Creek, V. champinii) confers strong resistance to multiple nematode genera with Rf values below 0.5 for Meloidogyne incognita, supporting Shiraz and Cabernet Sauvignon production on sandy soils in the Barossa Valley, Australia, and the San Joaquin Valley, California. Freedom (1613C × Dog Ridge) provides broad-spectrum nematode resistance (Rf 0.3-0.8 for major Meloidogyne species) to Zinfandel and Chardonnay on nematode-infested sites in Lodi and the Central Coast AVAs. Dog Ridge (V. champinii) shows resistance to both root-knot and dagger nematodes, serving as a genetic source for nematode resistance breeding at UC Davis Foundation Plant Services.
A stress-resistance rootstock role that inheres in a grafted rootstock by virtue of its genetically determined capacity to resist feeding damage and population buildup by plant-parasitic nematodes affecting grapevine root systems. Nematode-resistant rootstocks prevent or limit nematode reproduction through mechanisms including hypersensitive root cell death at feeding sites, production of nematicidal compounds (particularly terpenoids and phenolic acids), formation of physical barriers through rapid lignification and suberization around feeding cells, and failure to support giant cell or syncytium development required for sedentary endoparasitic nematode reproduction. Resistance is quantified by nematode reproduction factor (Rf = final population / initial population), with resistant genotypes showing Rf values below 1.0 (population decline) versus Rf values of 10-100+ on susceptible V. vinifera. Nematode resistance in grapevine rootstocks derives primarily from V. champinii, V. rufotomentosa, V. cinerea, and certain V. rupestris selections that evolved in nematode-infested soils of the southern United States.
Nematode_Resistant_Rootstock_Role is subclassed by target nematode specificity, distinguishing resistance to Xiphinema index (dagger nematode, GFLV vector), Meloidogyne species (root-knot nematodes), and other genera. Resistance mechanisms and genetic bases differ between nematode groups: root-knot nematode resistance involves blocking giant cell formation, while dagger nematode resistance involves deterring ectoparasitic feeding. Most nematode-resistant rootstocks derive from V. champinii (Dog Ridge, Ramsey, Freedom) or V. rufotomentosa crosses, species native to sandy soils of Texas and the southeastern US where nematode pressure is severe.
Nematoden-tolerante Unterlage
nematode-resistant rootstock
nematode-tolerant rootstock
nematodenresistente Unterlage
nematodenwiderstandsfähige Unterlage
plant-parasitic nematode resistant rootstock
McKenry MV, Anwar SA (2006) Nematode and grape rootstock interactions including an improved understanding of tolerance, Journal of Nematology 38(3): 312-318 Ferris H et al. (2012) Reflections on plant and soil nematode ecology, Annual Review of Phytopathology 50: 303-330 Cousins P, Walker MA (2002) Genetics of resistance to Meloidogyne incognita in crosses of grape rootstocks, Theoretical and Applied Genetics 105: 802-807
Nematode Resistant Rootstock Role
Nerol content quality in Muscat of Alexandria contributes the fresh, citrus-rose top note that complements the heavier linalool and geraniol character, with concentrations of 100-250 μg/L in well-ripened fruit from Mediterranean climates while below independent sensory threshold, nerol participates in additive and synergistic interactions with other monoterpenes that enhance the overall floral complexity perceived in Muscat wines, demonstrating how subthreshold compounds can contribute to aroma through mixture effects.
Nerol_Content_Quality represents the chemical quality characterizing the concentration of nerol (3,7-dimethyl-cis-2,6-octadien-1-ol) in grape berries and wine, capturing the acyclic monoterpene alcohol that contributes fresh, rose-citrus aroma notes as the cis-isomer counterpart to geraniol. Nerol differs from geraniol only in the geometric configuration of the 2,3 double bond, yet this structural difference produces somewhat distinct sensory properties, with nerol described as having a fresher, more citrus-like rose character compared to the heavier, sweeter rose note of geraniol. In grape berries and wines, nerol typically occurs at lower concentrations than geraniol, usually comprising 5-15% of total free monoterpenes in aromatic varieties, with concentrations ranging from 5-30 μg/L in neutral varieties through 50-150 μg/L in semi-aromatic cultivars to 100-300 μg/L in intensely aromatic Muscat varieties. The sensory threshold of nerol in wine matrices is approximately 300-400 μg/L, substantially higher than linalool or geraniol, meaning that nerol rarely exceeds threshold in most wines and contributes primarily through additive effects with other terpenes rather than as a dominant character. Nerol and geraniol interconvert through acid-catalyzed isomerization, with the equilibrium favoring geraniol at wine pH, contributing to the decline of nerol relative to geraniol during wine aging. Nerol accumulates in both free and glycosylated forms in berry skins, following accumulation patterns similar to other monoterpene alcohols with peak concentration near harvest maturity. The nerol-to-geraniol ratio provides information about terpene stability and wine chemical evolution.
Nerol_Content_Quality addresses the cis-isomer of the geraniol structure, typically present at lower concentrations but contributing to overall terpene complexity. The higher sensory threshold of nerol compared to linalool and geraniol means its contribution is primarily through mixture interactions rather than dominant character impact. The isomerization equilibrium favoring geraniol over nerol at wine pH explains the decrease in nerol/geraniol ratio during aging. Nerol quantification employs the same GC-based methods as other monoterpenes, with careful chromatographic resolution needed to separate nerol from geraniol.
Nerol-Konzentration
Nerolgehalt
Nerylverbindungsgehalt
Zitrus-Rosen-Terpenqualität
cis-3,7-dimethyl-2,6-octadien-1-ol concentration
citrus-rose terpene quality
nerol level
neryl compound content
Marais J (1983) Terpenes in the Aroma of Grapes and Wines: A Review. South African Journal of Enology and Viticulture 4(2): 49-58 Rapp A and Mandery H (1986) Wine Aroma. Experientia 42(8): 873-884
Nerol Content Quality
Net carbon balance tracking through the growing season in Pinot Noir reveals the seasonal productivity pattern, with early season strong positive balance (gross gain 18 g C/vine/day minus respiration 6 g C/day = net +12 g C/day) supporting rapid shoot growth, mid-season peak (gross 25, respiration 10, net +15 g C/day) supporting fruit development, late season decline (gross 15, respiration 12, net +3 g C/day) as canopy ages, and post-harvest near-zero or negative balance requiring reserve mobilization for remaining root growth and cold hardening, with seasonal integration yielding total net carbon gain of approximately 950 g C/vine available for structure (400 g), fruit (350 g), and reserves (200 g).
Net carbon balance quality is a carbon balance quality that represents the difference between gross carbon gain and total respiratory carbon losses, characterizing the carbon actually retained by the vine that is available for allocation to structural growth, fruit development, and reserve storage. Within the BFO framework, this quality specifically addresses the retained carbon dimension of carbon balance, distinguishing the net amount available for vine functions from gross acquisition while recognizing that net carbon balance is the physiologically meaningful measure that determines what the vine can actually accomplish with its carbon resources. The ontological significance lies in net carbon balance determining the resources available for all non-respiratory vine functions, where positive net balance enables growth and reproduction while negative balance indicates carbon deficit requiring reserve mobilization, making net carbon balance the central indicator of vine productive capacity and the integrating variable that management seeks to optimize for sustainable quality production. The calculation of net carbon balance subtracts all respiratory fluxes from gross carbon gain, with Net Balance = Gross Gain - (Maintenance Respiration + Growth Respiration + Photorespiration), and with the result representing the carbon available for structural increment and reserve accumulation. The temporal and spatial variation in net carbon balance creates complex patterns, with individual leaves shifting from strong carbon sources (young mature leaves with high photosynthesis and low respiration) to weak sources or even carbon sinks (old or shaded leaves with declining photosynthesis but maintained respiration), and with whole-vine net balance varying diurnally (positive in day, negative at night), seasonally (declining as canopy ages), and with crop load (increasing sink demand potentially exceeding source capacity). The relationship between net carbon balance and fruit quality involves the competition between vegetative and reproductive sinks for limited net carbon, where excessive vegetative growth diverts carbon from fruit, while excessive crop load may exhaust carbon supply before complete ripening, making optimization of the source-sink balance through crop and canopy management essential for quality fruit production. Understanding net carbon balance quality enables diagnosis of carbon limitation affecting fruit quality, optimization of crop load relative to canopy capacity, and prediction of vine capacity for current-season fruit ripening and following-season reserve accumulation.
Net_Carbon_Balance_Quality addresses the retained carbon after respiratory losses that is available for vine functions. This quality provides the physiologically meaningful productivity measure that determines actual capacity for growth and reproduction. The class supports source-sink balance optimization and productivity assessment through net carbon balance characterization.
NPP
Nettokohlenstoffbilanz
Nettoprimärproduktivität
net assimilation
net primary productivity
retained carbon
Lakso AN, Poni S (2005) "Viticulture-Enology 2020": linking carbohydrate supply and demand as a basis for grapevine yield and quality management. Acta Horticulturae 689: 331-340 Poni S et al. (2018) Grapevine quality: a multiple choice issue. Scientia Horticulturae 234: 445-462
Net Carbon Balance Quality
Net photosynthesis rate monitoring in Chardonnay throughout the growing season reveals the seasonal productivity pattern, with rates increasing from 8 μmol CO₂/m²/s at budbreak through maximum of 15.5 μmol CO₂/m²/s at mid-season (fully expanded leaves, optimal temperatures) then declining to 10 μmol CO₂/m²/s during ripening (heat stress, early senescence) and 4 μmol CO₂/m²/s at harvest (advanced senescence), with integrated seasonal carbon gain determining the carbohydrate pool available for current fruit ripening and subsequent season reserve storage.
Net photosynthesis rate quality is a carbon fixation efficiency quality that represents the net rate of CO₂ uptake by grapevine leaves measured as the difference between gross photosynthetic carbon fixation and respiratory CO₂ release, characterizing the actual carbon gain available to support growth and fruit development under specific environmental conditions. Within the BFO framework, this quality specifically addresses the net carbon balance of leaves, distinguishing net photosynthesis (observable gas exchange) from gross photosynthesis (total carboxylation) while recognizing that net rate represents the physiologically and agronomically meaningful measure of leaf productivity that determines carbohydrate supply for the whole plant. The ontological significance lies in net photosynthesis rate providing the direct measure of leaf carbon contribution to the vine, where the difference between CO₂ fixation and CO₂ release determines the net carbon export from leaves to sinks including developing fruit clusters, growing shoots, and storage organs, making net photosynthesis the fundamental productivity measure for viticulture. The physiological basis for net photosynthesis involves the balance between carboxylation (CO₂ uptake via Rubisco) and decarboxylation processes (mitochondrial respiration, photorespiration), with net rate (An) calculated as gross photosynthesis minus respiration (Rd) minus photorespiration (Rp), typically expressed as μmol CO₂/m² leaf area/second. The environmental and physiological factors affecting net photosynthesis include light intensity (increasing rate to saturation), CO₂ concentration (increasing rate following Michaelis-Menten kinetics), temperature (optimum typically 25-30°C for grapevine), water status (stomatal closure reducing rate under stress), and leaf age (rate declining from peak at full expansion through senescence). The typical ranges for net photosynthesis in Vitis vinifera span 8-18 μmol CO₂/m²/s under favorable conditions, with variation reflecting cultivar differences, environmental conditions, and vine physiological status. Understanding net photosynthesis rate quality enables direct assessment of leaf productivity, environmental response characterization, and vineyard management optimization targeting maximum sustainable carbon gain.
Net_Photosynthesis_Rate_Quality addresses the observable net CO₂ uptake representing actual leaf productivity. This quality is the standard measure from gas exchange analysis and provides the physiologically meaningful carbon gain value. The class supports productivity assessment and environmental response characterization through net assimilation measurement.
Netto-CO₂-Aufnahme
Nettoassimilation
Nettophotosyntheseraten
apparent photosynthesis
apparente Photosynthese
net CO₂ assimilation
net carbon uptake
Long SP, Bernacchi CJ (2003) Gas exchange measurements, what can they tell us about the underlying limitations to photosynthesis? Procedures and sources of error. Journal of Experimental Botany 54(392): 2393-2401 Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell & Environment 26(8): 1393-1405
Net Photosynthesis Rate Quality
Chardonnay bears a Neutral_White_Wine_Scion_Role as the archetype of non-aromatic white varieties, with terpene concentrations typically below 0.2 mg/L, producing wines where terroir character (Chablis minerality, Meursault richness) and winemaking decisions (malolactic fermentation, oak aging) define the sensory profile rather than inherent grape aromatics. Trebbiano Toscano (Ugni Blanc) serves in a Neutral_White_Wine_Scion_Role as the world's most planted white variety, producing high-acid, neutral-flavored wines valued primarily as Cognac and Armagnac distillation base, where neutrality is a virtue allowing spirit character to emerge. Macabeo (Viura) bears a Neutral_White_Wine_Scion_Role in Rioja and Cava, contributing body and oxidative stability to blends and base wines without dominating aromatic character.
A white wine scion role that inheres in a grafted scion cultivar by virtue of its genetically determined low accumulation of varietal aroma compounds, producing wines with subtle, understated aromatic profiles that serve as vehicles for terroir expression, winemaking influence, or blending. Neutral white wine scions accumulate monoterpenes at concentrations below typical sensory thresholds (less than 0.1-0.3 mg/L), lack significant thiolic precursors, and produce wines where fermentation-derived aromas (esters, higher alcohols), malolactic-derived compounds (diacetyl), and oak-derived flavors (vanillin, lactones) may dominate over inherent grape-derived character. Neutral varieties are valued for their capacity to reflect site characteristics (minerality, salinity, texture) without masking by intense varietal aromatics, for their versatility in blending (contributing body, acidity, or structure without flavor dominance), and for their suitability as base wines for oak-aged or sparkling wine production. The "neutral" designation does not imply inferior quality—many of the world's most celebrated white wines (white Burgundy, classified Graves) are produced from neutral varieties.
Neutral_White_Wine_Scion_Role captures varieties at the opposite end of the aromatic spectrum from terpenic/thiolic cultivars. Neutrality is not absolute—all grapes contain some aromatic compounds—but reflects relatively low concentrations of impact compounds with high sensory thresholds. Many neutral varieties gain aromatic complexity through winemaking (barrel fermentation, lees stirring, extended aging) rather than expressing inherent grape character. The neutral category includes some of the world's most important varieties by planted area (Chardonnay, Trebbiano, Airén) and some of the most prestigious wines, demonstrating that aromatic intensity is not prerequisite for quality. Neutral varieties dominate sparkling wine and distillation base wine production where clean, high-acid musts are preferred.
dezente Weißweinsorte
neutral white variety
neutrale Weißweinsorte
nicht-aromatische Rebsorte
non-aromatic variety
understated white variety
vinous white variety
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Jackson RS (2014) Wine Science 4th Ed, Academic Press Ribereau-Gayon P et al. (2006) Handbook of Enology Vol 2, Wiley Waterhouse AL et al. (2016) Understanding Wine Chemistry, Wiley
Neutral White Wine Scion Role
Night harvest suitability quality evaluation in Riverland Australia compared outcomes for Chardonnay harvested across the full diurnal cycle during a heat wave vintage with daytime maxima exceeding 40°C. Night-harvested fruit (10:00 PM-4:00 AM, berry temperature 14-18°C) demonstrated excellent quality preservation with juice browning indices of 0.12 AU, retained varietal aromatics, and fresh, balanced wines receiving quality scores of 88 points. Morning-harvested fruit (6:00-10:00 AM, berry temperature 20-26°C) showed moderate quality with browning of 0.18 AU and scores of 84 points. Afternoon-harvested fruit (2:00-6:00 PM, berry temperature 35-42°C) exhibited severe quality compromise with browning of 0.38 AU, cooked fruit character, and scores of 72 points. The dramatic temperature-quality relationship established night harvest suitability quality as essential for premium production in this warm region, motivating capital investment in lighting systems, mechanical harvester modifications for night operation, and labor agreements permitting nocturnal shift work that enabled systematic night harvest implementation.
Night harvest suitability quality is a harvest condition quality that inheres in a grapevine cultivar or clone and represents the compatibility of fruit characteristics and vineyard conditions with harvest operations conducted during nighttime hours under artificial illumination, exploiting the coolest temperatures of the diurnal cycle to maximize fruit quality preservation while accepting the operational complexities of after-dark work including lighting requirements, safety considerations, labor availability constraints, and the specialized equipment needed for effective nocturnal operations. This quality extends the cool-harvest concept beyond morning scheduling to capture the full quality potential of temperature-controlled harvest timing in warm climates where even morning temperatures may exceed optimal thresholds for quality-sensitive cultivars. The ontological significance of night harvest suitability quality lies in its role enabling premium quality production in warm climate regions where daytime temperatures would otherwise compromise fruit quality during harvest, effectively extending the geographic range of quality viticulture by decoupling harvest timing from solar-driven temperature patterns that limit daytime operations. The biological basis of night harvest suitability parallels morning harvest advantages including reduced oxidation rates, preserved aromatic compounds, and minimized microbial activity, while typically achieving even cooler fruit temperatures that approach overnight minima in the range of 10-15°C depending on climate and season. Additional biological considerations specific to night harvest include the diurnal patterns of plant metabolism with potentially altered sugar and acid distributions in fruit harvested during the dark period, though research generally indicates minimal compositional differences attributable to night versus day harvest timing when temperature effects are controlled. Cultivar characteristics favoring night harvest include extreme oxidation sensitivity, high-value aromatic compounds susceptible to thermal degradation, and thin-skinned fragility benefiting from the maximum turgor-related firmness achieved at coolest temperatures. Operational requirements for night harvest suitability include vineyard access for equipment under low-visibility conditions, row spacing and trellis configurations compatible with lighting systems, absence of steep terrain creating safety hazards, and labor force willing and legally permitted to work night shifts, establishing that night harvest suitability quality encompasses both biological and operational dimensions.
Night_Harvest_Suitability_Quality is_a Harvest_Condition_Quality, representing compatibility with after-dark harvest operations achieving minimum diurnal temperatures. This class addresses the extension of cool-harvest principles to nighttime operations increasingly adopted in warm climate viticulture. Operational requirements including lighting, safety, and labor considerations must be evaluated alongside biological suitability. Distinguished from Morning_Harvest_Suitability_Quality by the after-dark timing and associated logistical complexity. Most relevant for warm climate regions where night harvest provides the only avenue to cool-temperature fruit delivery.
Dunkelheits-Lesekompatibilität
Nachternte-Eignung
Nachtlese-Eignung
after-dark harvest fitness
cool-night harvest adaptation
night picking suitability
nocturnal harvest compatibility
nächtliche Erntetauglichkeit
Falcão LD et al. (2007) A survey of seasonal temperatures and vineyard altitude influences on 2-methoxy-3-isobutylpyrazine, C13-norisoprenoids, and the sensory profile of Brazilian Cabernet Sauvignon wines. Journal of Agricultural and Food Chemistry 55:3605-3612 Reynolds AG (2010) Managing Wine Quality Volume 2: Oenology and Wine Quality. Woodhead Publishing, Cambridge Petrie PR, Sadras VO (2008) Advancement of grapevine maturity in Australia between 1993 and 2006: putative causes, magnitude of trends and viticultural consequences. Australian Journal of Grape and Wine Research 14:33-45
Night Harvest Suitability Quality
Night transpiration assessment in Syrah comparing well-watered and deficit-irrigated vines reveals the nocturnal water loss affecting water status interpretation, with well-watered vines showing night transpiration of 0.42 mmol/m²/s (12% of daytime peak, pre-dawn water potential -0.35 MPa equilibrated with wet soil) while deficit vines show 0.28 mmol/m²/s (8% of reduced daytime, but pre-dawn -0.65 MPa not equilibrating with soil at -0.45 MPa due to continued nocturnal water loss), demonstrating how night transpiration creates pre-dawn water potential depression that must be considered when using pre-dawn measurements for irrigation scheduling.
Night transpiration quality is a transpiration rate quality that represents the water vapor loss occurring during the dark period when photosynthesis is not operating, characterizing the nocturnal water loss through incompletely closed stomata and the cuticle that affects vine water status recovery and represents a potentially significant but often overlooked component of daily water consumption. Within the BFO framework, this quality specifically addresses the nocturnal dimension of water loss, distinguishing night transpiration from daytime gas exchange while recognizing that night transpiration, though typically only 5-15% of daily total, affects the pre-dawn water potential that indicates vine water status and determines the starting point for each day's transpiration cycle. The ontological significance lies in night transpiration affecting the water relations interpretation and management timing that depends on pre-dawn water potential as a water stress indicator, where significant nocturnal water loss prevents complete overnight recovery of vine water status, causing pre-dawn water potential to underestimate soil water availability and potentially confusing irrigation scheduling based on this widely used stress indicator. The physiological basis for night transpiration involves primarily cuticular water loss (continuing regardless of stomatal aperture) plus incomplete stomatal closure in many species including grapevine, where residual nocturnal stomatal opening allows water loss without the carbon gain that would justify daytime transpiration, representing a water cost without photosynthetic benefit. The magnitude of night transpiration in Vitis vinifera typically ranges from 5-15% of daytime transpiration depending on atmospheric conditions (VPD during night), stomatal behavior (degree of nocturnal closure varying among cultivars), and cuticular properties, with higher values under warm dry nights when VPD remains elevated. The water relations implications of night transpiration include the potential disconnect between pre-dawn leaf water potential and soil water potential when nocturnal transpiration prevents equilibration, affecting the interpretation of pre-dawn measurements as soil water indicators, and the cumulative water consumption from night losses that contributes to seasonal water demand even without productive gas exchange. Understanding night transpiration quality enables accurate interpretation of pre-dawn water potential measurements, complete accounting of vine water consumption, and evaluation of cultivar differences in nocturnal water use that affect water status recovery and irrigation requirements.
Night_Transpiration_Quality addresses the often-overlooked nocturnal water loss component affecting water status recovery. This quality impacts interpretation of pre-dawn water potential as a stress indicator. The class supports accurate water status assessment and complete water budget accounting through night transpiration characterization.
Dunkelphasentranspiration
En-Wert
Nachttranspiration
dark period transpiration
night water loss
nocturnal transpiration
nächtlicher Wasserverlust
Caird MA et al. (2007) Nighttime stomatal conductance and transpiration in C₃ and C₄ plants. Plant Physiology 143(1): 4-10 Rogiers SY, Clarke SJ (2013) Nocturnal and daytime stomatal conductance respond to root-zone temperature in 'Shiraz' grapevines. Annals of Botany 111(3): 433-444
Night Transpiration Quality
Nitrate content quality monitoring in Central Valley table grape production tracks the high nitrogen fertilization regimes (150-250 kg N/ha) employed for maximum yield, with petiole nitrate analysis (targeting 500-1500 mg/kg dry weight at bloom) guiding fertilization timing while berry nitrate (typically 20-80 mg/L at harvest) remaining incidental to the primary concern of adequate vine nitrogen for productivity wine grape producers in premium regions target lower petiole nitrate (300-800 mg/kg) reflecting moderate nitrogen nutrition that balances vine health with the quality benefits of controlled nitrogen status.
Nitrate_Content_Quality represents the chemical quality characterizing the concentration of nitrate ions (NO₃⁻) in grape tissue, capturing the inorganic nitrogen form taken up by roots that serves as the primary nitrogen source for most vineyard soils while potentially indicating vine nitrogen status and soil fertility conditions. Nitrate is the dominant form of plant-available nitrogen in well-aerated, neutral to alkaline soils, taken up by roots through specific nitrate transporters and subsequently reduced to ammonium through nitrate reductase and nitrite reductase enzymes before assimilation into amino acids. In grape berries, nitrate concentration is typically low (5-50 mg/L as nitrate) because efficient reduction converts most absorbed nitrate to organic nitrogen forms before accumulation in fruit. Elevated berry nitrate (above 100 mg/L) may indicate excessive soil nitrogen availability, impaired nitrate reduction capacity (possibly from molybdenum deficiency affecting nitrate reductase), or recent heavy nitrogen fertilization that has not been fully metabolized. The quality implications of nitrate content relate primarily to its indicator function rather than direct wine effects, as berry nitrate reflects nitrogen nutrition dynamics and potential excessive nitrogen status associated with quality problems (delayed ripening, disease susceptibility, vegetal character). During fermentation, nitrate is not utilized by Saccharomyces cerevisiae, which lacks the assimilatory nitrate reduction pathway present in some other fungi and bacteria. In wine, nitrate content reflects must composition unchanged by fermentation, potentially serving as a marker for vineyard nitrogen management. Regulatory limits for nitrate in wine are not typically specified, though general food safety concerns about nitrate/nitrite apply. Nitrate analysis employs ion chromatography, colorimetric methods (cadmium reduction to nitrite followed by Griess reaction), or ion-selective electrodes.
Nitrate_Content_Quality addresses the inorganic oxidized nitrogen form distinct from ammonium (reduced inorganic nitrogen) and organic nitrogen (amino acids, proteins). Nitrate is primarily a vine nutrition indicator rather than a direct wine quality parameter, as it neither contributes to YAN nor creates wine stability issues at typical concentrations. The inability of Saccharomyces cerevisiae to utilize nitrate distinguishes it from ammonium and amino acids. Nitrate in wine raises theoretical food safety considerations (potential nitrosamine formation) but concentrations are typically too low for practical concern in the absence of contamination.
NO₃⁻ concentration
NO₃⁻-Konzentration
Nitratgehalt
Nitrationenqualität
anorganischer oxidierter Stickstoffgehalt
inorganic oxidized nitrogen content
nitrate ion quality
nitrate level
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 341-385 Spayd SE et al. (1994) Nitrogen Fertilization of White Riesling Grapes in Washington. Must and Wine Composition. American Journal of Enology and Viticulture 45(1): 34-42
Nitrate Content Quality
Nitrate uptake accounts for 70-90% of total nitrogen acquisition in most vineyard soils due to rapid nitrification of ammonium under aerobic conditions. High-affinity nitrate transport (HATS, Km 10-100 µM) dominates at soil nitrate concentrations below 0.5 mM, while low-affinity transport (LATS, Km 1-5 mM) contributes at higher concentrations. Rootstock 110 Richter shows 40% higher nitrate uptake capacity than 3309C under low nitrogen availability (0.1 mM NO₃⁻), attributable to elevated VvNRT2.1 expression. Nitrate reductase activity in roots and leaves converts absorbed nitrate to ammonium for amino acid synthesis, with cultivar-specific partitioning between root and shoot reduction.
A nitrogen uptake efficiency disposition that is the inherited capacity of a Vitis vinifera plant to absorb nitrate ions (NO₃⁻) from the soil solution through high-affinity and low-affinity nitrate transporter systems, representing the dominant nitrogen acquisition pathway in well-aerated vineyard soils.
Nitrate uptake dominates grapevine nitrogen acquisition except in waterlogged or highly acidic soils where nitrification is impaired. The dual transporter system (HATS and LATS) enables efficient uptake across the wide range of soil nitrate concentrations encountered in vineyards. Nitrate also serves as a signaling molecule influencing root architecture and gene expression beyond its nutritional role. Cover crop management and organic matter inputs modulate soil nitrate availability, interacting with genotypic uptake capacity.
NO3 uptake capacity
NO₃⁻-Akquisition
Nitrataufnahme
Nitrattransporter-Aktivität
nitrate acquisition
nitrate transporter activity
Pii Y et al. (2014) The interaction between iron nutrition, plant species and soil type shapes the rhizosphere microbiome. Plant Physiology and Biochemistry 99:158-166 Lecourt J et al. (2015) The nitrogen status of grapevine (Vitis vinifera) rootstocks affects root-shoot signaling. Functional Plant Biology 42:1038-1048
Nitrate Uptake Disposition
Nitrate uptake rate measurement in Cabernet Sauvignon roots using ¹⁵N-labeled nitrate reveals the concentration-dependent kinetics, with uptake rate of 8.5 μmol NO₃⁻/g root dry weight/hour at soil solution concentration of 5 mM (saturating high-affinity system), declining to 2.2 μmol/g/hour at 0.5 mM (limiting concentration but high-affinity transporters maintaining uptake), and only 0.4 μmol/g/hour at 0.05 mM (below Km of high-affinity system, severely limited uptake), demonstrating the kinetic basis for maintaining soil nitrate above threshold concentrations for adequate nitrogen nutrition.
Nitrate uptake rate quality is a nitrogen uptake efficiency quality that represents the rate of nitrate (NO₃⁻) absorption by grapevine roots, characterizing the acquisition of the predominant available nitrogen form in most vineyard soils that is highly mobile in soil solution and primarily delivered to roots through mass flow in the transpiration stream. Within the BFO framework, this quality specifically addresses the nitrate form of nitrogen uptake, distinguishing nitrate from ammonium acquisition while recognizing that nitrate is typically the dominant nitrogen form in well-aerated, neutral to alkaline vineyard soils where nitrification converts ammonium to nitrate. The ontological significance lies in nitrate uptake rate determining the acquisition of the nitrogen form that requires root assimilation (reduction to ammonium before incorporation into amino acids), with this metabolic cost affecting the energy efficiency of nitrogen nutrition while the high mobility of nitrate in soil means that mass flow often delivers nitrate to roots faster than it can be absorbed, such that uptake transporter capacity rather than soil delivery typically limits nitrate acquisition. The molecular basis for nitrate uptake involves the NRT (Nitrate Transporter) protein family, with high-affinity transporters (NRT2 family) operating at low external nitrate concentrations and low-affinity transporters (NRT1/NPF family) operating when nitrate is abundant, and with transporter expression regulated by both external nitrate availability and internal nitrogen status to match uptake capacity to demand. The environmental factors affecting nitrate uptake rate include soil nitrate concentration (driving uptake kinetics), soil temperature (affecting both transporter activity and microbial nitrification producing nitrate), soil moisture (affecting mass flow delivery and root function), and root system characteristics (total absorbing surface and transporter density). The diurnal and seasonal patterns of nitrate uptake show higher rates during periods of active transpiration (enhancing mass flow delivery), during rapid growth (high nitrogen demand upregulating transporters), and when soil temperature favors both root activity and nitrification, with uptake declining during dormancy and under conditions limiting root function. Understanding nitrate uptake rate quality enables optimization of nitrate fertilizer timing, assessment of soil nitrogen availability relative to uptake capacity, and interpretation of vine nitrogen dynamics under varying environmental conditions.
Nitrate_Uptake_Rate_Quality addresses the acquisition rate of the dominant nitrogen form in most vineyard soils. This quality is mediated by NRT transporter family with concentration-dependent kinetics. The class supports nitrogen fertilization timing and soil nitrogen management through nitrate uptake characterization.
NO₃⁻ uptake
NO₃⁻-Aufnahme
NRT activity
NRT-Aktivität
Nitratakquisitionsrate
Nitrataufnahmerate
nitrate absorption
nitrate acquisition rate
Crawford NM, Glass ADM (1998) Molecular and physiological aspects of nitrate uptake in plants. Trends in Plant Science 3(10): 389-395 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 197-245
Nitrate Uptake Rate Quality
The nitrogen content of a Napa Valley Cabernet Sauvignon vineyard on Bale series alluvial soil documents total nitrogen of 0.14% (moderate) and mineral nitrogen of 22 mg/kg (12 mg/kg NO₃-N + 10 mg/kg NH₄-N) at pre-budbreak sampling in March — this moderate nitrogen status, combined with the estimated annual nitrogen mineralization rate of 50-70 kg N/ha from organic matter decomposition, provides sufficient nitrogen for moderate canopy development (12-14 primary leaves per shoot at véraison, leaf area index 2.5-3.5) without the excessive vegetative growth that would shade the fruit zone and delay phenolic ripening. The nitrogen management protocol for this site specifies no supplemental nitrogen fertilization in normal seasons, with a conditional application of 20-30 kg N/ha as calcium ammonium nitrate only in seasons where petiole nitrogen at bloom falls below 0.6% (the lower threshold of the adequate range for Cabernet Sauvignon), reflecting the viticultural principle that nitrogen sufficiency rather than nitrogen luxury produces optimal wine quality.
A nutrient analysis data component that records the concentration of nitrogen in the vineyard soil, measured as total nitrogen (Kjeldahl or Dumas combustion method, expressed as percent by weight) and/or mineral nitrogen (ammonium-N plus nitrate-N, extracted with 2M KCl, expressed as mg/kg), with nitrogen being the nutrient of greatest viticultural management significance because of its dominant influence on vegetative vigor — nitrogen supply is the primary soil-derived determinant of shoot growth rate, canopy density, and the vegetative-reproductive balance that governs fruit quality in wine grape production.
Nitrogen_Content documents the most intensively managed nutrient in quality-focused viticulture — nitrogen is unique among the essential nutrients in that both deficiency and excess produce clearly detrimental effects on wine quality, and the optimal range is narrow, making precise nitrogen management essential. Nitrogen deficiency (petiole N 1.0% at bloom) promotes vigorous vegetative growth that shades the fruit zone (reducing anthocyanin synthesis, delaying sugar and methoxypyrazine maturation, and promoting Botrytis through canopy humidity), increases must yeast-assimilable nitrogen (YAN) beyond the 200-300 mg/L range associated with controlled fermentation kinetics, and may promote the formation of ethyl carbamate precursors in wine: the quality-optimal nitrogen status lies between these extremes, typically at petiole N of 0.6-0.8% at bloom, corresponding to moderate canopy development with adequate but not excessive leaf area for fruit ripening. The soil nitrogen pool is overwhelmingly organic (95-98% of total soil nitrogen is in organic forms — proteins, amino acids, nucleic acids, and humic nitrogen compounds — that are unavailable to plants until mineralized by soil microorganisms to ammonium and subsequently nitrified to nitrate), and the rate of mineralization is controlled by soil temperature, moisture, aeration, and organic matter quality — making the timing and amount of plant-available nitrogen supply inherently unpredictable from total nitrogen measurement alone.
Boden-Stickstoff
N content measurement
N-Gehalt-Messung
Stickstoffgehalt
nitrogen availability assessment
soil nitrogen record
Conradie WJ (2001) Timing of nitrogen fertilisation and the effect on the performance of Bukettraube, South African Journal of Enology and Viticulture 22: 60-68 Bell SJ and Henschke PA (2005) Implications of nitrogen nutrition for grapes, fermentation and wine, Australian Journal of Grape and Wine Research 11: 242-295 White RE (2003) Soils for Fine Wines, Oxford University Press Robinson JB (2005) Critical plant tissue values, in Christensen LP and Smart DR (eds) Proceedings of the Soil Environment and Vine Mineral Nutrition Symposium, ASEV
Nitrogen Content
Nitrogen content quality management in Champagne base wine production targets the elevated must nitrogen (180-250 mg N/L YAN) required for the demanding double fermentation process, with vigorous vine nitrogen uptake from deep chalky soils supplemented by judicious fertilization (40-60 kg N/ha) to achieve the nitrogen levels supporting both primary fermentation completion and the secondary in-bottle fermentation (prise de mousse) that defines méthode traditionnelle nitrogen-deficient base wines (below 140 mg N/L) risk stuck secondary fermentation that ruins entire cuvées, making nitrogen content quality a critical production parameter.
Nitrogen_Content_Quality encompasses all chemical qualities that characterize the concentration, composition, and forms of nitrogen-containing compounds in grape berry tissue, capturing the essential macronutrient that profoundly influences vine productivity, grape composition, fermentation performance, and wine quality. Nitrogen in grapes exists in multiple forms including inorganic ammonium, free amino acids, peptides, proteins, and nucleic acids, with the total nitrogen pool and its partitioning among forms affecting distinct quality dimensions. The critical importance of nitrogen derives from its central role in both vine physiology (essential for protein synthesis, chlorophyll formation, and enzyme function) and fermentation biochemistry (providing yeast nutrition that determines fermentation rate, completion, and volatile production). This category subdivides into Total_Nitrogen_Quality addressing the complete nitrogen pool regardless of form Yeast_Assimilable_Nitrogen_Quality capturing the fraction directly available for yeast nutrition and Amino_Acid_Profile_Quality characterizing the specific amino acid composition relevant to fermentation outcomes and wine chemistry. Nitrogen accumulation in berries depends on vine nitrogen status, which reflects soil nitrogen availability, fertilization history, cover crop competition, and rootstock effects on nitrogen uptake and distribution. Berry nitrogen content typically ranges from 100-600 mg N/L expressed as total nitrogen, with the enormous variation reflecting cultivar, vine status, and environmental conditions. Low nitrogen status produces sluggish fermentation, hydrogen sulfide formation, and reduced fruity ester production, while excessive nitrogen increases rot susceptibility, delays ripening, and may produce elevated ethyl carbamate precursors. The management of grape nitrogen content through viticultural practices and understanding its impact on fermentation represents one of the most direct vineyard-to-wine quality linkages in modern enology.
Nitrogen_Content_Quality is positioned as a direct subclass of Chemical_Quality, parallel to Sugar_Content_Quality, Acid_Content_Quality, and Mineral_Content_Quality, though nitrogen is technically a macronutrient mineral. The separate treatment reflects nitrogen's unique importance in fermentation and its occurrence in multiple organic and inorganic forms requiring distinct analytical approaches. The class integrates both vineyard (nitrogen nutrition, vine status) and winemaking (fermentation) perspectives on nitrogen quality. Nitrogen analysis methods vary with the form of interest—Kjeldahl or Dumas combustion for total nitrogen, enzymatic or HPLC methods for specific forms.
N content
N-Gehalt
Stickstoffverbindungsqualität
Traubenstickstoffqualität
grape nitrogen quality
nitrogen compound content
nitrogenous composition quality
Bell SJ and Henschke PA (2005) Implications of Nitrogen Nutrition for Grapes, Fermentation and Wine. Australian Journal of Grape and Wine Research 11(3): 242-295 Spayd SE et al. (1994) Nitrogen Fertilization of White Riesling Grapes in Washington. Must and Wine Composition. American Journal of Enology and Viticulture 45(1): 34-42
Nitrogen Content Quality
Nitrogen data for Chardonnay from the Côte de Beaune (2024 vintage) records YAN 168 mg/L (PAN 112 mg/L + ammonium 56 mg/L), below the recommended minimum of 200 mg/L for the planned 13.5% alcohol fermentation, prompting a calculated diammonium phosphate (DAP) addition of 0.15 g/L at mid-fermentation to prevent sluggish completion. Nitrogen data for Sauvignon Blanc from Marlborough (2024 vintage) documents YAN 285 mg/L (PAN 195 mg/L + ammonium 90 mg/L), well above the sufficiency threshold and favorable for volatile thiol release by the selected Saccharomyces cerevisiae strain, with the high PAN fraction providing diverse amino acid substrates for ester biosynthesis. Nitrogen data for Grüner Veltliner from Kremstal (2024 vintage) records YAN 210 mg/L, adequate for the target 13.0% alcohol fermentation without supplementation.
Nitrogen data is a must analysis data entity that documents the concentration and speciation of nitrogen-containing compounds available for yeast metabolism in grape must, encompassing yeast assimilable nitrogen (YAN) as the aggregate bioavailable metric, primary amino nitrogen (PAN/FAN) representing the α-amino acid fraction, and ammonium ion content representing the inorganic nitrogen fraction, measured through standardized analytical methods on pre-fermentation must samples and providing the essential nutritional assessment for predicting fermentation kinetics, managing stuck or sluggish fermentation risk, and optimizing yeast-derived aroma compound production.
Nitrogen data bridges viticultural fruit assessment with fermentation management, as must nitrogen status is determined by vineyard factors (soil nitrogen availability, rootstock uptake efficiency, canopy vigor, crop load) but its consequences are realized entirely during fermentation. The concept of yeast assimilable nitrogen was developed in the 1990s to distinguish the biologically relevant nitrogen pool from total nitrogen — only ammonium ions and α-amino acids (excluding proline, which Saccharomyces cerevisiae cannot metabolize under anaerobic conditions) are utilized during fermentation, while proline (often the most abundant amino acid in must) and proteins remain largely unavailable. The threshold for adequate YAN depends on the potential alcohol level of the must, as higher-sugar musts require proportionally more nitrogen to sustain yeast biomass through extended fermentation — a general guideline of 140 mg/L YAN per 12% potential alcohol plus an additional 10-12 mg/L per additional degree of alcohol provides a first approximation. Nitrogen deficiency (YAN 400 mg/L) may promote excessive biomass formation, reduce varietal thiol release, and leave residual nitrogen that supports spoilage organism growth post-fermentation. The amino acid composition within PAN influences fermentation-derived aroma profiles — branched-chain amino acids (leucine, isoleucine, valine) are precursors to higher alcohols and their esters, while cysteine and glutathione conjugates serve as precursors for volatile thiol release in aromatic cultivars.
Gärnährstoffdaten
Moststickstoffgehalt
Stickstoffanalysedaten
Stickstoffdaten
Stickstoffversorgungsdaten
fermentation nutrient data
must nitrogen content
nitrogen analysis data
nitrogen status data
yeast nutrient data
Bell SJ, Henschke PA (2005) Implications of nitrogen nutrition for grapes, fermentation and wine. Australian Journal of Grape and Wine Research 11(3): 242-295 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1: The Microbiology of Wine and Vinifications, 2nd Ed, Wiley, pp.: 75-108 Ugliano M et al. (2007) Effect of nitrogen supplementation and Saccharomyces species on hydrogen sulfide and other volatile sulfur compounds in Shiraz fermentation and wine. Journal of Agricultural and Food Chemistry 55(15): 6231-6239
Nitrogen Data
Nitrogen deficiency symptom quality assessment in Marlborough Sauvignon Blanc evaluated the nitrogen status gradient across a vineyard with variable soil depth and organic matter content to establish site-specific nitrogen management zones. Visual nitrogen deficiency symptom quality scoring at véraison used a 0-5 scale based on canopy color and vigor, with scores ranging from 0 (dark green, vigorous) through 2 (medium green, moderate vigor) to 5 (yellow-green, severely stunted). Blocks with shallow stony soils showed severe nitrogen deficiency symptom quality (scores 4-5) with pale yellow-green canopy, shoot length averaging 65 cm versus 120 cm in adequate zones, leaf area per shoot reduced by 55%, and older leaves showing advanced chlorosis progressing to senescence by mid-ripening. Tissue analysis confirmed the visual nitrogen deficiency symptom quality assessment, with petiole nitrate-nitrogen at 180 ppm in deficient zones versus 650 ppm in adequate zones (sufficiency threshold 500 ppm). Fruit from nitrogen-deficient vines showed reduced berry weight (0.9 g versus 1.3 g) and lower yeast-assimilable nitrogen (85 mg/L versus 180 mg/L), with the low YAN creating fermentation management challenges. The nitrogen deficiency symptom quality mapping enabled variable-rate nitrogen fertigation targeting deficient zones while maintaining moderate nitrogen status in vigorous zones to optimize the balance between vine health and the restrained vigor associated with high-quality Sauvignon Blanc production.
Nitrogen deficiency symptom quality is a macronutrient deficiency symptom quality that inheres in a grapevine organism and represents the characteristic visual symptoms and growth limitations resulting from inadequate nitrogen availability, including progressive chlorosis beginning on older leaves and advancing acropetally, reduced shoot growth and vigor, smaller leaves with pale green to yellow coloration, premature leaf senescence, and reduced fruit set and berry size, with nitrogen deficiency representing the most common macronutrient limitation in viticulture due to the large nitrogen requirements for protein synthesis and the mobility of nitrogen in soil making supply-demand matching challenging. This quality captures the grapevine response to inadequate nitrogen, recognizing that as the nutrient required in largest quantity and a component of all proteins, nucleic acids, and chlorophyll, nitrogen deficiency produces dramatic effects on virtually all aspects of vine growth and function. The ontological significance of nitrogen deficiency symptom quality lies in the complex management challenge nitrogen presents, where deficiency reduces yield and vine health while excess promotes vegetative vigor that may compromise fruit quality, requiring balanced nitrogen management that maintains vine nitrogen status within the optimal range for both productivity and wine quality objectives. The biological basis of nitrogen deficiency symptoms encompasses the central role of nitrogen in amino acid and protein synthesis affecting all growth processes, the nitrogen content of chlorophyll molecules producing the characteristic chlorosis when nitrogen limits chlorophyll production, and the high mobility of nitrogen within the plant allowing redistribution from older to younger tissues, which produces the characteristic symptom progression from basal to distal leaves. The relationship between nitrogen status and wine quality adds complexity to nitrogen management, with moderate nitrogen deficiency potentially beneficial for wine quality by limiting vigor and promoting fruit exposure while also reducing yeast-assimilable nitrogen with implications for fermentation kinetics and hydrogen sulfide production, creating the need to balance vine health, yield, and wine quality objectives in nitrogen management decisions.
Nitrogen_Deficiency_Symptom_Quality is_a Macronutrient_Deficiency_Symptom_Quality, serving as parent class for Chlorosis_Pattern_Quality_(N), Growth_Reduction_Quality_(N), and Early_Senescence_Quality_(N) which address specific nitrogen deficiency manifestations. This class addresses the overall nitrogen deficiency syndrome while child classes capture particular symptom types. Most common macronutrient deficiency in viticulture due to large requirements and soil mobility. The vigor-quality trade-off complicates nitrogen management beyond simple deficiency correction. YAN (yeast-assimilable nitrogen) affects fermentation with wine quality implications.
N deficiency symptoms
N-Mangel-Zeichen
Niedrig-Stickstoff-Symptome
Stickstoffmangel-Symptome
Stickstoffstress-Indikatoren
low nitrogen symptoms
nitrogen limitation signs
nitrogen stress indicators
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Bell SJ, Henschke PA (2005) Implications of nitrogen nutrition for grapes, fermentation and wine. Australian Journal of Grape and Wine Research 11:242-295 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication
Nitrogen Deficiency Symptom Quality
1103 Paulsen (V. berlandieri × V. rupestris) demonstrates high N efficiency, supporting vigorous Primitivo canopy development (pruning weights 0.8-1.2 kg/m) in Puglia under moderate fertilization, with petiole N at bloom exceeding 1.0% where 110 Richter delivers only 0.7-0.8%. 140 Ruggeri shows high N acquisition efficiency maintaining Catarratto YAN above 180 mg N/L on unfertilized Sicilian hillside vineyards where shallow soils limit N availability. SO4 (V. berlandieri × V. riparia) provides intermediate N efficiency to Riesling in the Rheingau, balancing adequate N supply for fermentation (YAN 140-160 mg N/L) with moderate vigor appropriate for steep-slope viticulture.
A nutrient acquisition rootstock role that inheres in a grafted rootstock by virtue of its enhanced capacity for nitrogen (N) acquisition from soil and delivery to the scion, supporting adequate vegetative growth, chlorophyll synthesis, and yeast assimilable nitrogen (YAN) accumulation in berries under conditions of limited soil N availability or restricted N fertilization. Nitrogen-efficient rootstocks acquire N through extensive root proliferation in N-rich soil zones, high-affinity nitrate (NRT1/2 family) and ammonium (AMT family) transporter expression, efficient N assimilation (nitrate reductase and glutamine synthetase activity), and sustained N translocation via xylem (as nitrate, amino acids, and ureides). Nitrogen efficiency is quantified by petiole N concentration at bloom (efficient genotypes maintaining >0.8-1.0% dry weight on low-N soils versus 150 mg N/L versus <100 mg N/L for inefficient genotypes under equivalent conditions).
Nitrogen_Efficient_Rootstock_Role captures rootstock effects on vine N economy that influence both vegetative growth and fermentation performance. The high-vigor V. berlandieri × V. rupestris rootstocks (1103P, 140Ru, 99R) generally show high N efficiency, while the lower-vigor V. riparia × V. rupestris types (3309C, 101-14Mgt) show lower N acquisition, consistent with their overall growth-restricting effects. Nitrogen efficiency can be detrimental in high-fertility situations where it promotes excessive vigor, but is valuable on infertile soils and under organic/biodynamic production where synthetic N inputs are restricted. YAN sufficiency (>140-150 mg N/L) prevents stuck fermentations and reduces volatile acidity and H₂S production risk.
N efficient rootstock
N-effiziente Unterlage
NUE rootstock
Stickstoff-Nutzungs-effiziente Unterlage
nitrogen efficient rootstock
nitrogen use efficient rootstock
stickstoffeffiziente Unterlage
Zerihun A, Treeby MT (2002) Biomass distribution and nitrate assimilation in response to N supply for Vitis vinifera cv. Cabernet Sauvignon on five Vitis rootstock genotypes, Australian Journal of Grape and Wine Research 8: 157-162 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 295-305 Holzapfel BP, Treeby MT (2007) Effects of timing and rate of nitrogen supply on leaf nitrogen status, vine nitrogen status and wine grape composition, Australian Journal of Grape and Wine Research 13: 106-115 Bell SJ, Henschke PA (2005) Implications of nitrogen nutrition for grapes, fermentation and wine, Australian Journal of Grape and Wine Research 11: 242-295
Nitrogen Efficient Rootstock Role
Nitrogen uptake efficiency varies 2-3 fold among rootstock genotypes, with Fercal showing high efficiency (45-60 mg N/m root length/day) compared to 101-14 Mgt (20-35 mg N/m/day) under identical nitrogen supply. Total seasonal nitrogen uptake ranges from 25-50 kg N/ha in low-vigor systems to 80-120 kg N/ha in high-vigor irrigated vineyards. Uptake timing shows two peaks in most cultivars, during rapid spring shoot growth and again post-harvest during root flush. VvNRT1 and VvNRT2 gene family members encode high-affinity and low-affinity nitrate transporters with rootstock-specific expression patterns.
A macronutrient uptake disposition that is the inherited capacity of a Vitis vinifera plant to absorb nitrogen from the soil solution in various chemical forms (nitrate, ammonium, amino acids) through root transporter systems, with genotype-specific efficiency determining nitrogen acquisition per unit root length or root biomass under given soil nitrogen availability.
Nitrogen_Uptake_Efficiency_Disposition serves as the parent class for nitrate and ammonium uptake subcategories. Nitrogen is typically the most limiting macronutrient in vineyard soils and exerts the strongest influence on vine vigor and fruit composition. Excessive nitrogen uptake promotes vegetative growth at the expense of fruit quality, while deficiency reduces yield and photosynthetic capacity. Rootstock selection for appropriate nitrogen uptake efficiency enables vigor management without excessive fertilizer inputs.
N uptake capacity
N-Akquisitionseffizienz
Stickstoffaneignungsvermögen
Stickstoffaufnahme-Effizienz
nitrogen acquisition efficiency
nitrogen foraging trait
Tomasi N et al. (2015) New aspects of grapevine response to nitrogen nutrition. Plant Signaling and Behavior 10:e1032953 Zerihun A, Treeby MT (2002) Biomass distribution and nitrate assimilation in response to N supply for Vitis vinifera L. cv. Cabernet Sauvignon on five Vitis rootstock genotypes. Australian Journal of Grape and Wine Research 8:157-162
Nitrogen Uptake Efficiency Disposition
Nitrogen uptake efficiency assessment comparing well-watered and deficit-irrigated Merlot reveals the water status interaction affecting nutrient acquisition, with well-watered vines showing nitrogen uptake rate of 2.8 g N/vine/week during active growth (high mass flow delivery, active root function), while deficit-irrigated vines show only 1.4 g N/vine/week (reduced mass flow, possible root hydraulic limitation), despite similar soil nitrogen availability, demonstrating how irrigation management affects nitrogen uptake efficiency with implications for integrated water-nitrogen management in quality-focused viticulture.
Nitrogen uptake efficiency quality is a nutrient uptake efficiency quality that represents the capacity of grapevine roots to acquire nitrogen from soil in its available forms (primarily nitrate and ammonium), characterizing the effectiveness of nitrogen acquisition that determines vine nitrogen status affecting vegetative vigor, photosynthetic capacity, and ultimately grape quality through nitrogen's roles in proteins, chlorophyll, and amino acid precursors for fermentation. Within the BFO framework, this quality specifically addresses the nitrogen component of nutrient uptake, distinguishing nitrogen acquisition from other nutrients while recognizing that nitrogen is typically the most yield-limiting nutrient in viticulture and the one most intensively managed through fertilization, with uptake efficiency directly affecting the environmental and economic costs of nitrogen management. The ontological significance lies in nitrogen uptake efficiency determining the conversion of soil and fertilizer nitrogen into vine nitrogen that supports photosynthesis and grape development, where high efficiency enables target vine nitrogen status with lower inputs, reducing both the cost of nitrogen fertilizers and the environmental impact of nitrogen leaching and volatilization that can affect water quality and contribute to greenhouse gas emissions. The physiological basis for nitrogen uptake involves active transport of nitrate and ammonium across root cell membranes by specific transporter proteins, with nitrate uptake mediated by NRT (Nitrate Transporter) family proteins and ammonium by AMT (Ammonium Transporter) proteins, with transporter expression regulated by both nitrogen availability and plant nitrogen status creating feedback regulation of uptake capacity. The soil and environmental factors affecting nitrogen uptake efficiency include soil nitrogen forms (nitrate versus ammonium availability varying with soil pH, temperature, and microbial activity), soil moisture (affecting mass flow delivery and root function), root distribution (determining access to soil nitrogen pools), and competition (from weeds, cover crops, or microorganisms consuming available nitrogen). The vine nitrogen status effects on uptake show downregulation of uptake capacity under high nitrogen status (feedback inhibition preventing luxury consumption) and upregulation under nitrogen deficiency (increased transporter expression to maximize acquisition), enabling some self-regulation of nitrogen uptake in response to plant demand. Understanding nitrogen uptake efficiency quality enables optimization of nitrogen fertilization timing and rates, rootstock selection for improved nitrogen acquisition, and management strategies that match nitrogen supply to vine demand while minimizing environmental losses.
Nitrogen_Uptake_Efficiency_Quality serves as the parent class for nitrogen form-specific uptake (Nitrate, Ammonium) and overall nitrogen use efficiency. This quality addresses the critical nitrogen acquisition function affecting vine vigor and grape quality. The class supports nitrogen fertilization optimization through uptake efficiency characterization.
N uptake efficiency
N-Absorptionskapazität
NUptE
Stickstoffaufnahmeeffizienz
Wurzelstickstoffakquisition
nitrogen absorption capacity
root nitrogen acquisition
Keller M et al. (2001) Cluster thinning effects on three deficit-irrigated Vitis vinifera cultivars. American Journal of Enology and Viticulture 52(2): 91-99 Wermelinger B, Koblet W (1990) Seasonal growth and nitrogen distribution in grapevine leaves, shoots and grapes. Vitis 29(1): 15-26
Nitrogen Uptake Efficiency Quality
Nitrogen use efficiency in grapevines ranges from 40-80 kg fruit per kg N in whole-vine content, with substantial rootstock and scion effects. Low-vigor rootstocks such as Riparia Gloire and 101-14 Mgt confer higher NUE (65-80 kg fruit/kg N) than high-vigor rootstocks such as 1103 Paulsen and 140 Ruggeri (45-60 kg fruit/kg N) by limiting vegetative N sinks. Internal NUE involves nitrogen remobilization from senescing leaves (recovering 60-75% of leaf N) and redistribution from storage pools in spring (contributing 30-50% of early-season N demand). High-NUE vines maintain adequate yield quality index (YAN above 150 mg N/L must) with 30-50% reduced N fertilizer application.
A nutrient use efficiency disposition that is the inherited capacity of a Vitis vinifera plant to produce biomass, yield, or quality outcomes per unit of nitrogen absorbed or available, determined by the combined efficiency of nitrogen uptake from soil, internal assimilation into organic compounds, remobilization between tissues, and allocation to productive versus vegetative growth.
Nitrogen use efficiency is the most studied nutrient efficiency parameter in viticulture due to nitrogen's dominant role in vine vigor, fruit composition, and environmental impact. The trait decomposes into uptake efficiency (N acquired per unit root investment) and utilization efficiency (fruit produced per unit N in vine). High NUE may conflict with wine quality objectives if must nitrogen (YAN) falls below fermentation requirements. Optimizing NUE requires balancing vegetative and reproductive N allocation rather than simply minimizing total N uptake.
N productivity
N-Verwertungseffizienz
Stickstoffnutzungseffizienz
Stickstoffproduktivität
nitrogen efficiency
nitrogen utilization trait
Treeby MT et al. (2000) Irrigation management and rootstock effects on navel orange (Citrus sinensis L. Osbeck) fruit quality. Agricultural Water Management 46:1-17 Verdenal T et al. (2021) Nitrogen management in the vineyard and its relationships with grape and wine quality. OENO One 55:1-32
Nitrogen Use Efficiency Disposition
Nitrogen use efficiency assessment comparing conventional and precision nitrogen management in Chardonnay reveals the management impact on NUE, with conventional uniform application (80 kg N/ha) achieving agronomic NUE of 62 kg fruit/kg N applied, while precision variable-rate application (same total N but spatially optimized) achieves NUE of 78 kg fruit/kg N (25% improvement), and reduced-rate precision management (55 kg N/ha targeting actual demand) achieves 95 kg fruit/kg N with equivalent fruit quality, demonstrating how matching nitrogen supply to spatially variable demand improves NUE while potentially reducing environmental impact.
Nitrogen use efficiency quality is a nitrogen uptake efficiency quality that represents the overall effectiveness with which grapevines convert available nitrogen inputs into useful outputs (biomass, yield, or quality parameters), characterizing the integrated efficiency of nitrogen acquisition, assimilation, and utilization that determines the productivity and environmental impact of nitrogen management in viticulture. Within the BFO framework, this quality addresses the system-level efficiency of nitrogen nutrition, distinguishing overall nitrogen use from component processes while recognizing that nitrogen use efficiency integrates uptake efficiency, assimilation efficiency, and utilization efficiency into a single metric that captures the practical effectiveness of nitrogen in supporting vine productivity and grape quality. The ontological significance lies in nitrogen use efficiency determining the agronomic and environmental return on nitrogen inputs, where high NUE enables quality grape production with lower fertilizer rates, reducing both production costs and the environmental footprint of viticulture from nitrogen losses through leaching, runoff, and gaseous emissions, while low NUE indicates that much input nitrogen fails to contribute to useful vine function. The components of nitrogen use efficiency include uptake efficiency (nitrogen acquired per nitrogen available), assimilation efficiency (nitrogen incorporated into organic forms per nitrogen absorbed), translocation efficiency (nitrogen reaching productive sinks per nitrogen in the plant), and utilization efficiency (yield or quality output per unit plant nitrogen), with each component potentially limiting overall NUE depending on conditions. The calculation methods for NUE vary with purpose, including agronomic efficiency (yield increase per unit N applied), physiological efficiency (biomass produced per unit N absorbed), and recovery efficiency (N in crop per unit N applied), with each metric providing different insights into nitrogen management effectiveness. The factors affecting nitrogen use efficiency include genotype (cultivar and rootstock differences in NUE components), nitrogen management (timing, rate, and form affecting synchrony with demand), water management (affecting both uptake and utilization), and crop load (determining sink demand for nitrogen), with optimal NUE achieved when nitrogen supply matches vine demand with minimal excess. Understanding nitrogen use efficiency quality enables optimization of nitrogen management for both productivity and sustainability, assessment of genotype differences in nitrogen efficiency, and development of precision nitrogen management targeting high NUE while maintaining target vine nitrogen status for quality fruit production.
Nitrogen_Use_Efficiency_Quality addresses the overall system-level efficiency of nitrogen nutrition integrating uptake, assimilation, and utilization. This quality determines the productivity and environmental return on nitrogen inputs. The class supports sustainable nitrogen management through NUE optimization balancing productivity and environmental impact.
N-Produktivität
NUE
Stickstoffkonversionseffizienz
nitrogen conversion efficiency
Conradie WJ (2005) Partitioning of mineral nutrients and timing of fertilizer applications for optimum efficiency. In: Proceedings of the Soil Environment and Vine Mineral Nutrition Symposium. ASEV, Davis, pp. 69-81 Holzapfel BP et al. (2019) Manipulating the postharvest period and its impact on vine productivity of Semillon grapevines. American Journal of Enology and Viticulture 70(2): 217-226
Nitrogen Use Efficiency Quality
Sémillon bears a Noble_Rot_Scion_Role as the dominant variety (typically 80%) for Sauternes and Barsac, where its thin skin and susceptibility to botrytis enable concentration to 300-400 g/L sugar, producing wines of legendary complexity—honey, apricot, caramel, orange peel—from Château d'Yquem (100% botrytised, €400-2000+/bottle) through the classified growths of the appellation. Furmint serves in a Noble_Rot_Scion_Role for Tokaji Aszú, where individual botrytised berries (aszú) are harvested, mashed into paste (aszú dough), and macerated with base wine, with classification by puttonyos (hods of aszú paste per barrel) indicating sweetness and concentration. Riesling bears a Noble_Rot_Scion_Role for German Beerenauslese and Trockenbeerenauslese, where individual botrytised berry selection achieves must weights of 110-150°Oechsle (TBA) for wines of 200-350 g/L residual sugar balanced by 10-15 g/L acidity.
A dessert wine scion role that inheres in a grafted scion cultivar by virtue of its susceptibility to and compatibility with Botrytis cinerea infection in its beneficial "noble rot" (pourriture noble/Edelfäule) form, producing extraordinarily concentrated sweet wines with distinctive honeyed, marmalade, and dried apricot character. Noble rot scions possess anatomical and compositional features favoring beneficial botrytis development—thin, permeable berry skins enabling fungal penetration without destructive splitting, loose cluster architecture promoting air circulation and drying between infection cycles, and high initial acidity buffering the tartaric acid reduction caused by botrytis metabolism. The fungal infection perforates berry skins, enabling water evaporation that concentrates sugars to 300-500 g/L while producing unique metabolites (glycerol up to 20 g/L, botryticine, sotolon) contributing to the distinctive sensory profile. Noble rot develops only under specific microclimatic conditions—morning fogs or mists providing humidity for infection, followed by warm, dry afternoons permitting desiccation—found in select terroirs worldwide (Sauternes, Tokaj, Rheingau, Loire, Burgenland).
Noble_Rot_Scion_Role captures a unique biological transformation where parasitic fungal infection produces desirable wine character—a case where disease creates quality. The distinction between beneficial noble rot and destructive grey rot (same organism, Botrytis cinerea) depends on microclimate and timing noble rot requires alternating humid and dry periods, while continuous humidity promotes destructive bunch rot. Sémillon and Furmint are the archetypes, but diverse varieties serve in noble rot production where conditions permit—Riesling, Chenin Blanc, Gewürztraminer, Sauvignon Blanc, Vidal, Grüner Veltliner. Botrytis-affected wines show exceptional longevity (50-100+ years for top Sauternes and TBA) due to high residual sugar, glycerol, and acidity acting as natural preservatives.
Botrytis-Sorte
Edelfäule variety
Edelfäulesorte
Pourriture-noble-Traube
botrytis variety
noble rot variety
pourriture noble grape
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Brook S (2002) Sauternes and Other Sweet Wines of Bordeaux, Mitchell Beazley Rippon S (2009) Tokaji, Mitchell Beazley Dittrich HH, Grossmann M (2011) Mikrobiologie des Weines 4th Ed, Ulmer Ribéreau-Gayon P et al. (2006) Handbook of Enology Vol 1, Wiley
Noble Rot Scion Role
The Noble_Rot_Susceptibility_Disposition of 'Sémillon' underlies its dominance in Sauternes production, where it typically comprises 75-80% of plantings in classified growths such as Château d'Yquem. 'Sémillon' possesses the ideal combination of traits for noble rot development: thin skin (mean 165 μm) permitting B. cinerea penetration and subsequent water loss, moderate initial sugar (22-24°Brix) providing substrate for fungal metabolism, low acidity maintaining palatability after acid reduction, and neutral aromatic profile transformed by fungal activity into complex honeyed, apricot, and saffron notes. In contrast, thick-skinned 'Cabernet Sauvignon' planted adjacently for red wine production rarely develops noble rot, instead either resisting infection or progressing directly to destructive grey rot when infected. Environmental monitoring at Château d'Yquem documented ideal noble rot conditions: morning fog raising humidity above 90% for 4-6 hours, followed by afternoon sunshine and temperatures of 18-24°C with humidity dropping below 60%. Under these conditions, 'Sémillon' berries reached harvest sugar concentrations of 35-42°Brix through 40-60% water loss over 3-4 weeks of progressive noble rot development, with multiple selective harvest passes (tries) recovering only optimally botrytized berries.
A Noble_Rot_Susceptibility_Disposition is a Botrytis_Resistance_Disposition that, paradoxically, represents the capacity of a Vitis vinifera berry to support the specialized "noble rot" (pourriture noble, Edelfäule) infection by Botrytis cinerea under specific environmental conditions, enabling the production of distinctive botrytized sweet wines including Sauternes, Tokaji Aszú, Trockenbeerenauslese, and Sélection de Grains Nobles. This disposition inheres in cultivars whose berries, when infected by B. cinerea under conditions of alternating humidity and dryness, undergo controlled desiccation and profound biochemical transformation rather than destructive grey rot decay. The noble rot process requires thin-skinned berries permitting fungal penetration and water loss, sufficient sugar content to support fungal metabolism while preventing complete berry collapse, and specific aromatic precursor compounds that undergo fungal transformation into desirable volatile compounds. During noble rot development, B. cinerea consumes organic acids (particularly tartaric acid) and metabolizes grape constituents to produce glycerol, gluconic acid, and distinctive aromatic compounds (sotolon, furaneol), while water evaporates through the perforated cuticle, concentrating sugars to 30-40°Brix or higher. The disposition is expressed only when humid mornings (promoting infection and fungal activity) alternate with dry, sunny afternoons (preventing grey rot progression and enabling desiccation), conditions typical of specific terroirs including Sauternes, Tokaj, and Rhine-Mosel valleys. Cultivars lacking Noble_Rot_Susceptibility_Disposition either resist B. cinerea infection entirely or develop destructive grey rot rather than the controlled noble rot transformation.
Noble_Rot_Susceptibility_Disposition is_a Botrytis_Resistance_Disposition, though it represents susceptibility to a specific, desirable form of Botrytis infection rather than resistance. This ontological positioning reflects that noble rot capacity is a specialized disposition within the broader Botrytis response spectrum, distinguished from destructive grey rot by environmental conditions and cultivar characteristics. The class captures the disposition for controlled, beneficial B. cinerea interaction rather than the destructive infection represented by general Botrytis susceptibility. Required traits include thin skin, appropriate sugar and acid balance, and aromatic precursor content enabling desirable transformation.
Botrytis-Süßwein-Eignung
Edelfäule susceptibility
Edelfäule-Anfälligkeits-Disposition
Pourriture-noble-Disposition
botrytized wine potential
noble rot capacity
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1: The Microbiology of Wine and Vinifications, 2nd Edition, John Wiley & Sons Fournier E et al. (2013) Botrytis cinerea epidemiology and noble rot, Botrytis - the Fungus, the Pathogen and its Management in Agricultural Systems, Springer Magyar I (2011) Botrytized wines, Advances in Food and Nutrition Research 63:147-206
Noble Rot Susceptibility Disposition
Pinot Noir hardwood cutting showing node-derived adventitious roots, cutting with 3 nodes treated with 2000 ppm IBA, microscopic examination revealing root primordia forming at all 3 buried nodes 6-8 days after treatment, primordia originating from cells near bud bases and leaf trace regions, roots emerging from nodes 11-14 days, cutting producing 9 nodal adventitious roots (3-4 per node) by day 21 versus only 2 internodal roots, nodal roots providing majority of root system enabling successful cutting establishment
Node derived adventitious root is shoot-derived adventitious root originating from node tissue at junction between stem internode and leaf or bud, characterized by formation from nodal meristematic regions or preformed root primordia existing in some species, showing preferential initiation at nodes compared to internodes due to anatomical and developmental features, demonstrating relatively easy induction requiring minimal hormonal treatment or sometimes spontaneous formation, and serving as primary adventitious root source in many propagation systems including grapevine cuttings where nodal roots typically form first and most abundantly, with node-derived roots showing typical initiation from cells near bud bases or leaf traces where vascular anatomy changes create favorable sites, exhibiting rapid development from activation of preformed primordia or quick de novo formation, and demonstrating that nodal rooting capacity represents key component of propagation success with cultivars showing strong nodal rooting generally considered easy-to-propagate while those with poor nodal rooting presenting propagation challenges. Node derived adventitious root anatomical basis includes= showing preformed primordia (some grapevine cultivars possessing latent root primordia at nodes requiring only activation), exhibiting vascular complexity (nodal regions with leaf trace departures and bud traces creating complex vascular architecture potentially favorable for root initiation), demonstrating meristematic retention (nodal regions potentially retaining more meristematic or pluripotent cells than internodes), revealing bud association (root primordia often forming near bud bases potentially sharing developmental signals), and displaying structural discontinuity (anatomical changes at nodes creating favorable sites for root emergence). Node derived adventitious root preferential formation includes= showing higher frequency (nodal roots typically more numerous than internodal roots in most grapevine propagation), exhibiting earlier development (nodal roots often forming before internodal roots in cutting propagation sequences), demonstrating easier induction (nodal rooting requiring lower auxin concentrations or less favorable conditions than internodal), revealing greater reliability (nodal rooting showing more consistent success across cultivars and conditions), and displaying primary importance (nodal roots providing majority of initial root system in many propagated vines). Node derived adventitious root in preformed primordia includes= showing latent primordia (some cultivars possessing dormant root primordia at nodes formed during original shoot development), exhibiting activation requirements (preformed primordia requiring hormonal or environmental signals to resume development), demonstrating rapid emergence (activated primordia potentially emerging faster than de novo formed roots), revealing cultivar variation (preformed primordia presence and abundance varying among grapevine genotypes), and displaying evolutionary significance (preformed primordia representing preparedness for vegetative propagation or stress response). Node derived adventitious root de novo formation includes= showing cell dedifferentiation (nodal cells reprogramming from shoot to root identity), exhibiting primordium organization (dedifferentiated cells organizing into structured root meristem), demonstrating vascular connection (developing nodal roots establishing connections to nodal vascular system), revealing emergence pathway (nodal roots growing outward through relatively short path to surface), and displaying complete development (de novo nodal roots developing full functionality comparable to preformed primordium-derived roots). Node derived adventitious root induction factors includes= requiring auxin signal (nodal rooting promoted by auxin though often responsive to lower concentrations than internodes), showing moisture dependence (adequate moisture at nodes essential for root development), exhibiting temperature effects (nodal rooting optimal at moderate temperatures 20-25°C), demonstrating burial importance (covering nodes with soil or media providing dark moist conditions promoting rooting), and revealing wounding effects (cutting preparation potentially providing wound signal enhancing nodal rooting). Node derived adventitious root in cutting propagation includes= providing primary roots (nodal roots typically forming majority of cutting root system particularly in early stages), enabling rapid establishment (early nodal root formation allowing cutting to quickly establish water and nutrient uptake), demonstrating node number importance (cuttings with more nodes having potential for more nodal roots increasing establishment success), revealing node burial requirement (nodes must be buried or in contact with moist medium to produce roots), and displaying propagation foundation (successful nodal rooting essential for cutting propagation success in viticulture). Node derived adventitious root cultivar patterns includes= showing genetic variation (cultivars varying in nodal rooting capacity from prolific to sparse), exhibiting easy-rooting cultivars (varieties like many V. vinifera and Riparia showing abundant nodal roots), demonstrating difficult-rooting types (some hybrids or species showing poor nodal rooting requiring special protocols), revealing selection criteria (rootstock breeding selecting for strong nodal rooting improving propagation efficiency and success rates), and displaying practical implications (cultivar nodal rooting capacity determining propagation ease, cost, and commercial feasibility). Node derived adventitious root functional development includes= showing vascular integration (nodal roots rapidly establishing functional vascular connections), exhibiting lateral branching (nodal roots producing lateral roots creating branched system), demonstrating absorption capacity (nodal roots developing root hairs and absorptive function), revealing secondary growth (nodal roots undergoing secondary thickening in perennial vines), and displaying persistence (nodal adventitious roots functioning throughout vine life as permanent root system).
Node_Derived_Adventitious_Root emphasizes nodal origin at stem-leaf junctions and is distinguished from internodal stem roots by focus on node formation. Critical for understanding preformed primordia, preferential formation, anatomical advantages, easier induction, primary importance, and the adventitious roots arising from nodes representing primary rooting sites in vegetative propagation.
Bud-base root
Joint-derived root
Knospen-Basis-Wurzel
Knoten-Adventivwurzel
Nodal adventitious root
Nodale Wurzel
Node Derived Adventitious Root
The node description of Nebbiolo documents prominent, swollen nodes (clearly visible as thickened rings on both growing shoots and dormant canes) with strong anthocyanin pigmentation producing a conspicuous red-violet ring at each node during the growing season (OIV 007:7) — the nodal anthocyanin is more intense than the internode pigmentation, creating an alternating pattern of strongly colored nodes and moderately colored (green with red stripes) internodes along the shoot axis that is one of Nebbiolo's most distinctive vegetative characters. The diaphragm at each node is visible as a dense transverse septum when the dormant cane is split longitudinally, and the node diameter exceeds the internode diameter by approximately 30-40%, creating the pronounced nodal swelling that contributes to the knobbly, irregular cane surface characteristic of Nebbiolo compared to the smoother, less prominently noded canes of Barbera.
A shoot description component that records the morphological characteristics of shoot nodes — the thickened stem positions where leaves, tendrils, inflorescences, and lateral buds are attached — documenting node prominence (the degree to which the node is swollen relative to the internode), anthocyanin pigmentation intensity at the node, diaphragm visibility (the septum separating successive internode pith cavities), and the arrangement of attached organs (leaf, tendril or inflorescence, and lateral bud) that collectively characterize the nodal architecture of the grapevine shoot.
Node_Description documents the morphological features of the critical developmental junction points of the grapevine shoot — each node represents the position of a leaf primordium in the shoot apical meristem, and the node's anatomy reflects the vascular connections between the axial shoot vasculature, the departing leaf trace (serving the attached leaf), the lateral bud trace (serving the axillary bud that may produce a lateral shoot or remain dormant as a future renewal bud), and the tendril or inflorescence trace (serving the tendril or cluster at that node): the complexity of these vascular connections produces the thickened, anatomically dense nodal tissue that is visible externally as the swollen node. The anthocyanin pigmentation of nodes is typically more intense than internode pigmentation in red-berried cultivars, reflecting the higher density of vascular and parenchyma tissue at the node that accumulates anthocyanin — this differential pigmentation creates the node-internode color contrast that is a diagnostic character in its own right, independent of the absolute pigmentation intensity. During dormancy, node morphology becomes the primary shoot identification character — node prominence (swollen versus flush), node color (darker or lighter than the internode bark), and the visibility of the diaphragm when the cane is cross-sectioned are the characters used by viticulturists to identify cultivars during winter pruning operations.
Knoten-Beschreibung
Nodien-Morphologie
Sprossknoten-Charakterisierung
nodal architecture description
node morphology record
stem node characterization
Node Description
A node one latent bud at the base of a cane of Vitis vinifera cv. Pinot Noir that remained suppressed for three consecutive seasons under normal growing conditions, finally breaking following mechanical damage that removed all canes and cordons during vineyard renovation, developing into a purely vegetative shoot 145cm in length with 24 leaves, no inflorescences, strong apical dominance suppressing lateral bud development, serving as the foundation for reestablishing a productive cordon structure over the subsequent two growing seasons through selective retention and positioning of lateral shoots.
A basal latent bud located specifically at the first node position from the origin point on annual canes, representing the most proximal bud position, characterized by extremely low fruitfulness (typically 0-5% inflorescence-bearing shoots even when forced to break), very small bud size (often 1.5-2.5mm diameter), tight suppression under apical dominance, and almost exclusively vegetative developmental potential. Node one latent buds serve primarily as reserve meristems for emergency shoot production following catastrophic damage to all distal buds, rarely breaking under normal vineyard conditions but capable of producing vigorous vegetative shoots when released from suppression, representing the last developmental resource for canopy reconstruction in severely damaged vines.
Node one latent buds are the least fruitful bud position on grapevine canes, reflecting their position in deep shade during the previous season's inflorescence initiation period and their evolutionary role as emergency reserve meristems rather than production buds. Pruning systems that rely on cane pruning typically remove node one entirely during winter pruning, retaining only nodes 2-12 or similar, while spur pruning systems may retain node one as part of two-bud spurs but expect no crop contribution. Understanding node one fruitfulness is critical for accurate yield prediction models and for assessing vine recovery potential following severe damage events.
Knoten-1-Knospe
basal node one bud
erste Knospe
first node bud
node 1 bud
Based on Sánchez & Dokoozlian (2005) "Bud microclimate and fruitfulness in Vitis vinifera L." American Journal of Enology and Viticulture 56=319-329 Buttrose (1970) "Fruitfulness in grapevines= The response of different cultivars to light, temperature and day length" Vitis 9=121-125 Vasconcelos & Castagnoli (2000) "Leaf canopy structure and vine performance" American Journal of Enology and Viticulture 51=390-396.
Node One Latent Bud
A node position basal secondary inflorescence at node 5 on a 7-node spur of Vitis vinifera cv. Pinot Noir in Burgundy, France, positioned at the standard node 5 location following the primary at node 3, measuring 11.8 cm length, located only 14 cm from the spur base due to the short spur architecture creating basal character through the node position itself, exhibiting typical node 5 secondary characteristics including compound paniculate architecture with 3 orders of branching, rachis diameter of 2.5 mm at base, bearing approximately 295 flowers, showing development reflecting standard node 5 programming with dimensions approximately 67% of the primary (17.5 cm, 485 flowers), basal environmental context from short spur architecture creating reduced light exposure (55% full sun) and protected position near permanent wood, developmental stage at BBCH 63 (30% flowers open) showing typical secondary phenological delay, node 5 position creating the basal character through standard inflorescence spacing on this short spur rather than exceptional placement, expected to contribute approximately 30% of spur yield, measured in early June during mid-bloom period, representing typical node position basal secondary where standard node 5 placement creates basal character on short shoot architecture.
A basal secondary inflorescence characterized by standard node 5 positioning creating basal character through the node location itself in Vitis vinifera, characterized by position at node 5 representing the standard secondary inflorescence node following the two-node gap pattern, basal character emerging from the node position particularly on short shoots where node 5 is near the shoot base, occurrence following the typical inflorescence spacing creating the sequential node 3 (primary), node 5 (this secondary) pattern, dimensions and development reflecting standard node 5 characteristics modified by the basal environmental context on short shoots, architectural organization showing typical secondary branching patterns, and functional characteristics determined by the node 5 position creating the reproductive structure at this specific location. Node position basal secondary inflorescences represent the standard node-determined positioning, occurring at the typical node 5 location which creates basal character on short shoots through the inherent node position rather than exceptional placement, exhibiting characteristics reflecting both the standard node 5 developmental program and the basal environmental context.
Node position basal secondary inflorescences emphasize the role of standard node positioning in creating basal character, distinguished from base-attached variants by node 5 location rather than exceptional attachment points. The node position designation highlights that basal character emerges from the inherent node 5 location on short shoots rather than unusual positioning. Position at node 5 follows the standard two-node gap inflorescence spacing pattern, with basal character resulting from short shoot architecture placing node 5 near the base rather than mid-shoot. Developmental programming follows typical node 5 secondary patterns with standard initiation timing, resource allocation, and architectural organization characteristic of secondaries regardless of absolute position on shoot. Environmental context varies with shoot length, with node 5 on short spurs (6-8 nodes) creating basal character while node 5 on long canes (12-15 nodes) creates mid-shoot lateral character. Size and complexity reflect standard secondary development modified by basal environmental conditions including light exposure and microclimate. Occurrence represents the typical pattern with most basal secondaries on short shoots resulting from standard node positioning rather than exceptional attachment. Management assessment focuses on evaluating the basal environmental conditions and their effects on development and potential fruit quality, with retention decisions based on actual light exposure and vine balance considerations rather than simply node position. The distinction from base-attached secondaries involves the mechanism creating basal character, with node position variants showing standard spacing while base-attached show exceptional attachment points.
Knotenposition basale sekundäre Infloreszenz
Standardknoten Basalblütenstand
node 5 basal secondary
positional basal secondary
positionelle basale Sekundärrispe
standard node basal inflorescence
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 291-326 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 519-558.
Node Position Basal Secondary Inflorescence
A node two basal leaf on a primary shoot of Vitis vinifera cv. Syrah in the Rhône Valley, France, positioned at the second node on a spur-originated shoot developing from primary bud, emerging 3 days after first leaf at 7 days post-budbreak following predictable developmental sequence, measuring 17.5 cm length by 16.2 cm width representing 22% larger area than median position leaves at nodes 7-9, exhibiting characteristic Syrah morphology with five lobes, moderate to deep sinuses, relatively long petiole, sparse abaxial pubescence, and diagnostic marginal tooth patterns reliably expressed at this position, achieving full expansion at 27 days post-budbreak, demonstrating peak photosynthesis rate of 13.6 μmol CO₂ m⁻² s⁻¹ in late May contributing approximately 32% of total shoot photosynthate during the critical post-bloom period supporting fruit set and early berry development, positioned opposite a well-developed tendril (as Syrah rarely shows basal fruitfulness), experiencing progressive light reduction from 75% full sun to 28% as upper canopy develops, measured at shoot BBCH stage 71 (fruit set) in mid-June, representing typical node 2 primary shoot leaf with characteristic morphology and function.
A second basal leaf specifically at node 2 on primary shoots developing from primary buds in Vitis vinifera, characterized by predictable position at the second node following standard phyllotactic pattern, development from well-formed preformed primordium within primary bud component of compound bud, emergence following first leaf with typical 2-4 day delay representing orderly developmental sequence, dimensions generally 15-25% larger than median shoot leaves reflecting basal position developmental priority, morphological features reliably expressing cultivar-diagnostic characteristics making node 2 position valuable for ampelographic identification, early maturation timeline achieving full expansion typically 25-28 days post-budbreak, high early-season photosynthetic contribution often representing 30-40% of total shoot carbon production during weeks 4-8 post-budbreak, and position typically opposite second tendril or potentially second inflorescence in cultivars showing high fruitfulness at basal positions. Node two basal leaves represent the standard second leaf expression in commercial viticulture where primary bud shoots dominate canopy composition, developing from compound bud primary components that received adequate resource allocation during previous-season development creating well-differentiated preformed primordia ready for rapid deployment following budbreak, exhibiting typical adult morphology without developmental abnormalities that sometimes affect more distal positions or secondary/tertiary bud leaves.
Node two basal leaves represent the most reliably diagnostic position for cultivar identification in many grapevine varieties, showing more consistent expression of characteristic morphological features compared to node 1 which sometimes exhibits simplified or modified morphology, and compared to distal positions which may show developmental variation or stress-induced modifications. The node 2 position benefits from early development ensuring adequate resource availability and favorable early-season environmental conditions, while avoiding the occasional developmental anomalies affecting the very first leaf position. Ampelographic descriptions standardly reference node 2 (and nodes 3-4) for detailed morphological characterization including lobe patterns, sinus characteristics, vein arrangements, and marginal features. Functional characteristics parallel first basal leaves with high early-season contribution declining through progressive shading, making node 2 leaves common targets for early basal leaf removal practices aimed at cluster exposure and microclimate modification. The predictable presence and characteristics of node 2 leaves on primary shoots makes this position valuable for systematic phenological studies, developmental research, and comparative morphological analyses.
Knoten-zwei primäres Spross-Blatt
primary shoot node two leaf
second position primary leaf
standard second basal leaf
standard zweites basales Weinblatt
zweite Position Hauptblatt
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 74-135 Poni et al. (2006) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 57=397-407 This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 179-202.
Node Two Basal Leaf
A node two latent bud on a two-bud spur of Vitis vinifera cv. Merlot in a cordon-trained, spur-pruned vineyard, breaking normally in spring along with the node one bud, developing into a shoot bearing one small cluster of 65 berries plus 12 leaves, while the node one position produces only vegetative growth, with the node two shoot subsequently cut back to two buds during winter pruning to maintain the spur position, illustrating the typical weak fruitfulness and renewal wood role of node two positions in spur-pruned systems.
A basal latent bud located at the second node position from the origin point on annual canes, characterized by low but variable fruitfulness (typically 10-30% inflorescence-bearing shoots, 0-1 small clusters per shoot), intermediate suppression between node one and mid-cane positions, and developmental potential ranging from purely vegetative to weakly reproductive depending on light exposure during previous season's differentiation period and cultivar-specific fruitfulness patterns. Node two latent buds represent a transitional position between non-fruitful basal nodes and productive mid-cane nodes, often retained in spur pruning systems as part of two-bud spurs where they contribute modest crop while providing renewal cane positions for subsequent season pruning.
Node two latent buds show high variability in fruitfulness among cultivars, with some varieties (e.g., Chardonnay, Riesling) showing measurable cluster production at node two under favorable conditions, while others (e.g., Cabernet Sauvignon, Merlot) rarely produce clusters at this position. Light exposure during the previous season's June-July period strongly influences node two fruitfulness, with well-exposed positions sometimes achieving 40-50% cluster-bearing shoots. Spur pruning systems rely heavily on node two (and node three) for both current crop and renewal wood, making node two fruitfulness a cultivar-specific consideration in training system selection.
Knoten-2-Knospe
basal node two bud
node 2 bud
second node bud
zweite Knospe
Based on Dunn & Martin (2000) "A functional association in Vitis vinifera L. cv. Cabernet Sauvignon between the extent of primary branching and the number of flowers formed per inflorescence" Australian Journal of Grape and Wine Research 6=88-92 Morrison & Iodi (1990) "The morphology and anatomy of adventitious root formation in Vitis vinifera cv. Cabernet Sauvignon" Vitis 29=323-334 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115.
Node Two Latent Bud
Detailed histological analysis of Nodosity_Limitation_Disposition at the University of Melbourne compared cellular responses to phylloxera feeding in resistant V. riparia 'Gloire de Montpellier' versus susceptible V. vinifera 'Shiraz' using microscopy of excised root tips inoculated with individual phylloxera crawlers. On 'Shiraz' roots, feeding induced rapid cell division in the pericycle and cortex within 48 hours, with recognizable nodosity structures forming by day 7 and reaching full development (2-3mm swelling with characteristic hook shape) by day 14. Phylloxera within nodosities showed active feeding and reproduction, with mean offspring production of 8.5 crawlers per adult over 21 days. On 'Gloire de Montpellier' roots demonstrating Nodosity_Limitation_Disposition, initial cellular response to feeding was detectable at 48 hours but accompanied by rapid browning of cells adjacent to the feeding site characteristic of hypersensitive response. By day 7, necrotic tissue surrounded the feeding site, preventing the coordinated cell proliferation necessary for nodosity formation. Rudimentary swellings occasionally formed but remained small (<0.5mm) and non-functional, with phylloxera showing 85% mortality and mean offspring production of only 0.3 crawlers per surviving adult. These observations confirm that Nodosity_Limitation_Disposition operates through rapid recognition and localized cell death that sacrifices infected cells to prevent establishment of functional feeding galls.
A Nodosity_Limitation_Disposition is a Root_Phylloxera_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that prevent or limit formation of nodosities—the characteristic hook-shaped galls induced by grape phylloxera feeding on young, unsuberized root tips. This disposition represents resistance at the primary infection site where first-instar crawlers or eggs initiate root colonization, determining whether phylloxera can establish feeding sites and begin population development on the root system. Nodosities form when phylloxera feeding induces abnormal cell division and expansion in the root tip meristem and cortex, creating hooked swellings that house the insect and provide continuous access to nutritious phloem and cortical cells. On susceptible V. vinifera, nodosities develop within 7-10 days of feeding initiation and support rapid phylloxera reproduction. Resistance mechanisms limiting nodosity formation include hypersensitive response killing cells at feeding sites before gall development can proceed, failure of root tissues to undergo phylloxera-induced cell proliferation, production of toxic secondary metabolites in root tips, and physical properties of root tissues preventing stylet penetration or sustained feeding. Nodosity_Limitation_Disposition varies among resistant Vitis species, with some (V. riparia) showing near-complete suppression of nodosity formation while others (V. berlandieri) permit limited nodosity development but prevent progression to vine-damaging tuberosities. Evaluating nodosity formation is a primary screening method for root phylloxera resistance in rootstock breeding programs.
Nodosity_Limitation_Disposition is_a Root_Phylloxera_Resistance_Disposition. Nodosities form on young unsuberized root tips and represent the initial colonization site for root phylloxera. This class is distinguished from Tuberosity_Resistance_Disposition which addresses galls on older lignified roots. Hypersensitive response appears to be a key mechanism, analogous to pathogen resistance. Nodosity limitation is typically stronger in V. riparia-derived rootstocks than in V. berlandieri-derived rootstocks, which may tolerate some nodosity formation while still providing adequate field resistance.
Nodositäts-Begrenzungs-Disposition
Nodositäts-Unterdrückung
Wurzelspitzen-Gallen-Resistenz
nodosity suppression
root tip gall resistance
young root resistance
Kellow AV et al. (2004) Interaction between Vitis vinifera and grape phylloxera: changes in root tissue during nodosity formation, Annals of Botany 93:581-590 Forneck A et al. (2002) Histochemistry and anatomy of phylloxera (Daktulosphaira vitifoliae) nodosities on young roots of grapevine (Vitis spp.), Vitis 41:93-97 Lawo NC et al. (2011) Grapevine phylloxera's performance on phylloxera resistant rootstock and their physiological response to root damage, Plant Physiology and Biochemistry 49:426-431
Nodosity Limitation Disposition
Non-flavonoid quality profiling in Chardonnay for premium white wine production addresses both hydroxycinnamic acid content (150-300 mg/L caftaric acid equivalents) affecting oxidation susceptibility during pressing, and stilbene accumulation (1-5 mg/kg resveratrol in healthy fruit, potentially 10-50 mg/kg following Botrytis challenge) providing both stress markers and health-relevant compounds winemakers managing non-flavonoid extraction through gentle pressing and reductive handling minimize hydroxycinnamic acid oxidation that would cause browning while preserving the stilbene content associated with premium vineyard sites.
Non_Flavonoid_Quality encompasses the chemical qualities characterizing the concentration and composition of phenolic compounds that lack the C6-C3-C6 flavonoid skeleton, capturing the structurally simpler phenolics that contribute to grape and wine chemistry through distinct mechanisms from flavonoids. Non-flavonoid phenolics in grapes include two major subclasses: hydroxycinnamic acids (C6-C3 phenylpropanoid derivatives) that serve as precursors to volatile phenols and participate in oxidation reactions, and stilbenes (C6-C2-C6 compounds) that function as phytoalexins with significant health-associated bioactivities. These compounds share biosynthetic origins in the phenylpropanoid pathway but diverge before the flavonoid branch point, producing simpler structures with distinct functional roles. Hydroxycinnamic acids, present primarily as tartaric acid esters (caftaric and coutaric acids), are the dominant phenolics in white grape juice and contribute to browning susceptibility, flavor precursor pools, and wine aging chemistry. Stilbenes, particularly resveratrol and its derivatives, accumulate in response to stress and pathogen challenge, providing induced defense while attracting considerable research interest for potential health benefits. The concentration of non-flavonoid phenolics varies substantially among cultivars and responds to environmental conditions, with stress factors generally promoting stilbene accumulation while hydroxycinnamic acid content shows more stable cultivar-specific patterns. In winemaking, non-flavonoid phenolics are extracted readily during pressing and early maceration due to their localization in flesh and skin vacuoles, contributing to must and wine composition from the earliest processing stages. The sensory contribution of non-flavonoids is primarily indirect, through their participation in chemical reactions that generate flavor-active products during fermentation and aging.
Non_Flavonoid_Quality is positioned as a direct subclass of Phenolic_Content_Quality, parallel to Flavonoid_Quality and Tannin_Quality, capturing phenolic compounds outside the flavonoid structural class. The C6-C3 hydroxycinnamic acids and C6-C2-C6 stilbenes represent distinct biosynthetic branches from the general phenylpropanoid pathway. Non-flavonoid phenolics are sometimes termed "non-tannin phenolics" in enological literature, though this terminology conflates structural classification with functional properties. The class excludes benzoic acid derivatives (C6-C1 compounds) that are present at low concentrations in grapes and typically not quality-determinative.
Nicht-Flavonoid-Gehalt
Nicht-Flavonoid-Zusammensetzung
Phenylpropanoid-Phenolkonzentration
einfache Phenolqualität
non-flavonoid composition
non-flavonoid phenolic content
phenylpropanoid phenolic concentration
simple phenolic quality
Singleton VL et al. (1978) Caftaric Acid Disappearance and Conversion to Products of Enzymatic Oxidation in Grape Must and Wine. American Journal of Enology and Viticulture 29(2): 107-117 Waterhouse AL (2002) Wine Phenolics. Annals of the New York Academy of Sciences 957(1): 21-36
Non Flavonoid Quality
A non-fruiting sterile shoot on Vitis vinifera cv. Grenache originating from trunk water shoot (latent bud origin confirmed by position and development pattern), observed through three consecutive growing seasons showing zero reproductive development despite retention as structural renewal (water shoot trained as replacement arm following Eutypa dieback removal of original arm), first-year vegetative-only (expected for water shoot), second-year producing 12 shoots with zero inflorescences despite 8 mid-cane positions normally 75%+ fruitful, third-year producing 14 shoots with zero inflorescences, definitively confirming persistent sterility characteristic of latent bud origin water shoots in this variety, with continued retention as permanent arm structure accepted as vegetative-only framework requiring spur-pruned positions from this arm to rely entirely on buds differentiating on subsequent seasons' shoots for eventual reproductive capacity restoration over time.
A sterile vegetative shoot demonstrating confirmed absence of reproductive capacity through multi-season observation or known origin from inherently sterile contexts, characterized by documented reproductive failure across multiple growing seasons, consistent vegetative-only development regardless of position or conditions, and classification as definitively sterile rather than possibly-sterile pending further observation. Non-fruiting sterile shoots represent verified categorically infertile growth, with sterility confirmed through extended observation (typically 2-3 seasons minimum), origin tracing (water shoot or sucker from latent buds with documented sterility patterns), or specific genetic/physiological conditions known to produce persistent sterility, distinguishing these shoots from unverified potentially-sterile shoots requiring additional observation for definitive classification.
Non-fruiting sterile shoots represent confirmed categorical sterility, with the non-fruiting designation emphasizing verified status versus potentially-sterile shoots still under observation. The confirmation process typically requires multi-season documentation showing consistent reproductive failure, eliminating possibility of temporary circumstantial non-fruiting due to stress, timing, or positional factors. Common confirmed-sterile contexts include water shoots and suckers from trunk or arm latent buds (widespread documentation of persistent sterility from such origins across varieties and regions), specific genetic sterility conditions (rare but definitively sterile when present), and chronic vine stress conditions producing persistent reproductive failure (though such severe stress typically triggers other interventions including vine removal). Management recognizes confirmed sterility as permanent or very long-term condition, utilizing such shoots only for vegetative functions (architectural renewal, photosynthetic capacity) without expecting reproductive contribution, with understanding that buds differentiating on sterile shoot wood may eventually acquire reproductive capacity through normal differentiation processes in subsequent seasons once shoots become established canes bearing normal annual shoots.
bestätigt unfruchtbarer Trieb
confirmed sterile shoot
dokumentiert steriler Trieb
verified non-reproductive shoot
Based on Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press, pp. 85-105.
Non Fruiting Sterile Shoot
Merlot showing non-functional senescent feeder roots in December, soil cores revealing feeder roots with brown coloration collapsed morphology complete root hair loss, FDA viability staining completely negative, isotope pulse-labeling (15N, 33P) showing zero tracer accumulation confirming no absorption, microscopic examination revealing advanced cytoplasmic degradation, non-functional feeder roots 7-14 months post-initiation comprising estimated 22% of total feeder root population but contributing 0% to current uptake, demonstrating critical distinction between feeder root presence and feeder root function
Non functional senescent feeder root is senescent feeder root emphasizing complete loss of absorption and all physiological functions, characterized by zero water uptake and nutrient transport demonstrated through exclusion from isotope tracers, complete loss of membrane selective permeability allowing unrestricted diffusion, absence of turgor pressure indicating osmotic failure, inability to respond to any environmental or physiological signals, and complete cessation of metabolism including respiration, with non-functional state typically beginning 5-12 months post-initiation when senescence progressed sufficiently eliminating absorption capacity despite potential structural persistence for additional period before complete decomposition, showing that functional cessation precedes morphological destruction creating period where feeder root physically present but physiologically inactive contributing nothing to current vine resource acquisition, and demonstrating critical distinction between feeder root presence and feeder root function with management and research requiring assessment of functional feeder root population not merely total feeder root counts or lengths to accurately evaluate vine absorption capacity and resource status. Non functional senescent feeder root absorption cessation includes= showing zero water uptake (complete inability to absorb water from soil confirmed through dye exclusion or isotope methods), exhibiting zero nutrient transport (labeled nutrient tracers not accumulating in vine through non-functional feeder roots), demonstrating no contribution (isotope mass balance confirming non-functional roots provide 0% of current uptake), revealing complete impermeability loss (fluorescent dyes freely entering non-functional roots indicating total barrier failure), and displaying transport failure (no active or passive transport occurring across non-functional root membranes). Non functional senescent feeder root viability loss includes= showing negative viability staining (FDA, TTC, or other vital dyes not accumulating in non-functional roots confirming death or near-death), exhibiting no metabolic activity (no oxygen consumption or CO2 production measurable from non-functional root zones), demonstrating ATP absence (biochemical assays showing no or minimal ATP in non-functional roots confirming energy metabolism cessation), revealing enzyme inactivation (key metabolic enzymes completely inactive in non-functional roots), and displaying death markers (biochemical or molecular markers of cell death present in non-functional root tissues). Non functional senescent feeder root turgor and osmotic failure includes= showing complete collapse (non-functional roots flattened or extremely shriveled from total turgor loss), exhibiting plasmolysis resistance (non-functional roots not responding to plasmolysis tests confirming membrane death), demonstrating osmotic incapacity (non-functional roots unable to maintain any osmotic gradients or generate turgor), revealing water status irrelevance (non-functional root appearance unchanged whether in water or air indicating no osmotic function), and displaying permanent deformation (non-functional root shape permanently altered not recovering even if environmental conditions improve). Non functional senescent feeder root in functional assessment includes= requiring functional distinction (studies measuring absorption must distinguish functional from non-functional feeder roots avoiding overestimation of capacity), exhibiting counting errors (simple feeder root counts including non-functional roots substantially overestimating effective absorption surface), demonstrating efficiency masking (including non-functional roots in surface area calculations artificially reducing calculated uptake rates per area), revealing spatial considerations (non-functional roots concentrated in drier deeper or older root zones while functional roots in favorable zones), and displaying temporal dynamics (functional feeder root population changing continuously through production activation and senescence to non-functional state). Non functional senescent feeder root persistence and decomposition includes= showing variable persistence (non-functional feeder roots potentially remaining intact hours to weeks after function ceased), exhibiting eventual decomposition (non-functional roots ultimately decomposing through microbial and enzymatic breakdown), demonstrating rapid breakdown (thin-walled nitrogen-rich feeder roots decomposing quickly once dead), revealing decomposition factors (temperature, moisture, microbial activity determining decomposition rate), and displaying nutrient release (nitrogen, phosphorus, and other nutrients released during non-functional root decomposition). Non functional senescent feeder root carbon and nutrient cycling includes= showing labile substrate (non-functional feeder roots providing readily decomposable organic matter high in nitrogen), exhibiting rapid mineralization (non-functional root decomposition releasing nutrients to soil within weeks to months), demonstrating carbon loss (non-functional root carbon largely respired by microbes during decomposition with minor fraction stabilizing in soil organic matter), revealing nutrient availability (nutrients from non-functional root decomposition potentially available for vine reuptake creating partial nutrient recovery), and displaying soil biology support (non-functional feeder root decomposition supporting rhizosphere microbial community activity and diversity). Non functional senescent feeder root in turnover quantification includes= representing mortality flux (non-functional feeder roots representing roots that exited functional pool through senescence), showing production requirement (non-functional root abundance indicating replacement need maintaining functional population), exhibiting turnover rate calculation (non-functional root production rate used calculating feeder root turnover rates), demonstrating population dynamics (functional feeder root population determined by production rate minus rate of transition to non-functional state), and revealing carbon allocation (extent of non-functional versus functional roots indicating efficiency of vine carbon allocation to root production). Non functional senescent feeder root research and management implications includes= requiring functional focus (research and management must focus on functional not total feeder roots for meaningful assessment), showing measurement challenges (distinguishing functional from non-functional feeder roots requiring careful methods beyond simple counting), exhibiting interpretation significance (feeder root measurements must account for non-functional fraction for accurate capacity estimates), demonstrating model requirements (feeder root function models requiring functional population parameters not total population measures), and revealing diagnostic value (ratio of functional to total feeder roots potentially indicating vine resource status or stress level).
Non_Functional_Senescent_Feeder_Root emphasizes complete functional cessation and zero absorption and is distinguished from functional or aging roots by focus on total physiological failure. Critical for understanding absorption cessation, viability loss, functional assessment requirements, decomposition, carbon cycling, and the non-functional state where feeder roots physically present but contributing nothing to vine resource acquisition requiring continuous production replacement.
Dead feeder root
Functionless fine root
Funktionslose Feinwurzel
Nicht-funktionelle seneszente Feinwurzel
Non-absorbing senescent root
Tote Saugwurzel
Non Functional Senescent Feeder Root
Syrah grafted on SO4 rootstock showing non-functional senescent root hairs 50-70 mm from root tip, hairs 7-11 days post-initiation beyond functional period, FDA viability staining completely negative confirming death, isotope pulse-labeling (15N and 32P) showing zero tracer accumulation in plant through non-functional hair zone confirming no absorption, microscopic examination revealing collapsed morphology, membrane failure evident through unrestricted dye entry, non-functional hairs comprising estimated 30% of total hair population but contributing 0% to current uptake, demonstrating critical distinction between hair presence and hair function
Non functional senescent root hair is senescent root hair emphasizing complete loss of absorption and physiological function, characterized by zero water uptake demonstrated through isotope or dye exclusion, zero nutrient transport confirmed through tracer studies, loss of membrane selective permeability allowing unregulated leakage, absence of turgor pressure indicating osmotic failure, and inability to respond to any environmental or physiological signals, with non-functional state typically beginning 6-9 days after hair initiation when senescence advances sufficiently to eliminate absorption capacity despite hair structural persistence, showing that functional cessation precedes morphological destruction creating period where hair physically present but physiologically inactive contributing nothing to current vine resource acquisition, and demonstrating critical distinction between hair presence and hair function with management or research needing to assess functional hair population not merely total hair counts. Non functional senescent root hair absorption cessation includes= showing zero water uptake (dye exclusion or isotope tracing demonstrating hair not absorbing water from soil), exhibiting zero nutrient transport (labeled nutrient tracers not accumulating in plant through non-functional hairs), demonstrating no contribution to uptake (isotope mass balance confirming non-functional hairs provide 0% of current root absorption), revealing membrane impermeability loss (fluorescent dyes freely entering non-functional hairs indicating complete permeability barrier failure), and displaying complete transport failure (no active or passive transport occurring through non-functional hair membranes). Non functional senescent root hair viability loss includes= showing negative viability staining (FDA or other vital dyes not accumulating in non-functional hairs confirming death), exhibiting no metabolic activity (no oxygen consumption or CO2 production measured from non-functional hair zones), demonstrating absence of ATP (biochemical assays showing no ATP in non-functional hairs confirming energy metabolism cessation), revealing enzyme inactivation (key metabolic enzymes completely inactive in non-functional hairs), and displaying cellular death markers (biochemical indicators of cell death present in non-functional hairs). Non functional senescent root hair turgor loss includes= showing complete collapse (non-functional hairs flattened or shriveled from total turgor loss), exhibiting plasmolysis resistance (non-functional hairs not responding to plasmolysis treatments confirming membrane death), demonstrating osmotic failure (non-functional hairs unable to maintain osmotic gradients or generate turgor), revealing water status irrelevance (non-functional hair appearance unchanged whether in water or dry conditions indicating no osmotic function), and displaying permanent deformation (non-functional hair shape permanently changed by collapse not recovering even if rehydrated). Non functional senescent root hair in absorption assessment includes= requiring functional distinction (studies measuring absorption must distinguish functional from non-functional hairs to avoid underestimating hair efficiency), exhibiting counting errors (simple hair counts including non-functional hairs overestimating effective absorption surface), demonstrating efficiency masking (including non-functional hairs in surface area calculations artificially reducing calculated absorption rates per area), revealing spatial considerations (non-functional hairs concentrated in older distal root zones while functional hairs proximal), and displaying temporal dynamics (functional hair population changing continuously through production and senescence). Non functional senescent root hair retention period includes= showing variable persistence (non-functional hairs potentially remaining attached to root for hours to days after function ceased), exhibiting eventual sloughing (non-functional hairs ultimately detaching from root surface through wall degradation or mechanical disturbance), demonstrating decomposition (non-functional hairs degrading in situ through microbial and enzymatic activity if remaining attached), revealing sloughing factors (mechanical disturbance, soil fauna, water flow, or continued degradation causing non-functional hair detachment), and displaying ghost hairs (non-functional hair wall remnants potentially visible on root surface even after cytoplasmic degradation complete). Non functional senescent root hair organic matter contribution includes= showing labile substrate (non-functional hairs providing readily decomposable organic matter to rhizosphere), exhibiting rapid decomposition (thin-walled single-cell hairs decomposing quickly after death), demonstrating nutrient release (nitrogen, phosphorus, and other nutrients from dead hairs released to soil potentially available for reuptake), revealing microbial food (non-functional hairs consumed by soil bacteria and fungi supporting rhizosphere microbial community), and displaying carbon input (hair carbon contributing to soil organic matter though hairs being small contributor per individual). Non functional senescent root hair in root system turnover includes= showing turnover component (non-functional hairs representing hairs exiting functional pool through senescence), exhibiting production balance (non-functional hair accumulation rate balanced by sloughing or decomposition maintaining population), demonstrating surface area loss (non-functional hairs representing lost absorption surface requiring replacement through new hair production), revealing functional population dynamics (functional hair population determined by production rate minus senescence rate creating non-functional hairs), and displaying continuous flux (non-functional hairs continuously produced through senescence and removed through sloughing creating steady-state functional population). Non functional senescent root hair research implications includes= requiring functional assessment (research must assess hair function not merely presence to understand absorption capacity), exhibiting methodological importance (methods distinguishing functional from non-functional hairs essential for accurate uptake measurements), demonstrating interpretation significance (hair density counts must consider functional proportion for meaningful interpretation), revealing management relevance (practices affecting ratio of functional to total hairs impacting effective absorption surface), and displaying modeling necessity (root function models requiring functional hair parameters not total hair measurements).
Non_Functional_Senescent_Root_Hair emphasizes complete functional loss and absorption cessation and is distinguished from functional hairs by focus on zero physiological capacity. Critical for understanding absorption cessation, viability loss, turgor failure, functional assessment requirements, organic matter contribution, turnover dynamics, and the non-functional state where hair physically present but contributing nothing to vine resource acquisition.
Dead root hair
Functionless hair
Funktionsloses Haar
Nicht-funktionelles seneszierende Wurzelhaar
Non-absorbing senescent hair
Totes Absorptionshaar
Non Functional Senescent Root Hair
Non-photochemical quenching assessment comparing shade-acclimated versus sun-acclimated Riesling leaves reveals adaptation in photoprotective capacity, with shade leaves showing maximum NPQ of 1.8 and slow induction (t½ = 8 minutes) reflecting limited xanthophyll pool and enzyme capacity, while sun leaves display NPQ of 3.5 with rapid induction (t½ = 2 minutes) from elevated zeaxanthin synthesis capacity, demonstrating how NPQ capacity adapts to light environment and why sudden exposure of shade leaves to high light risks photoinhibition before photoprotective capacity can develop.
Non-photochemical quenching quality is a chlorophyll fluorescence quality that represents the dissipation of excess absorbed light energy as heat through photoprotective mechanisms, characterizing the capacity and engagement of processes that safely dispose of light energy exceeding photochemical utilization to prevent oxidative damage to the photosynthetic apparatus. Within the BFO framework, this quality specifically addresses the photoprotective component of fluorescence quenching, distinguishing energy dissipation (NPQ) from productive photochemistry (qP) while recognizing that NPQ represents essential photoprotection without which high-light photosynthesis would rapidly cause irreversible PSII damage. The ontological significance lies in non-photochemical quenching indicating the engagement of photoprotective mechanisms that enable grapevines to tolerate high light environments, where NPQ capacity determines the ability to safely dissipate excess energy and thus the tolerance of leaves to light intensities exceeding electron transport capacity, with implications for canopy design and stress tolerance. The photophysical basis for NPQ involves multiple mechanisms including qE (energy-dependent quenching, requiring trans-thylakoid pH gradient and zeaxanthin), qT (state transition quenching, redistribution of antenna between photosystems), and qI (photoinhibitory quenching, reflecting PSII damage), with qE being the dominant and most rapidly reversible component that provides the primary photoprotective response. The calculation of NPQ employs the Stern-Volmer formulation NPQ = (Fm - Fm')/Fm' where the reduction in maximum fluorescence yield in light (Fm') relative to dark-adapted state (Fm) indicates the engagement of non-photochemical dissipation pathways. The xanthophyll cycle connection to NPQ involves the conversion of violaxanthin to zeaxanthin under excess light, with zeaxanthin enabling the conformational changes in antenna proteins that facilitate thermal energy dissipation, making carotenoid content relevant to NPQ capacity. Understanding non-photochemical quenching quality enables assessment of photoprotective capacity, diagnosis of light stress conditions requiring NPQ engagement, and evaluation of the balance between productive photochemistry and protective dissipation under vineyard conditions.
Non_Photochemical_Quenching_Quality addresses the photoprotective energy dissipation component of fluorescence quenching. This quality indicates engagement of heat dissipation mechanisms essential for high-light tolerance. The class supports photoprotection assessment and stress diagnosis through NPQ measurement revealing the balance between productive and protective energy use.
NPQ
NPQ-Wert
Photoprotektion
energy-dependent quenching
nicht-photochemisches Quenching
photoprotective quenching
thermal dissipation
thermische Dissipation
Müller P et al. (2001) Non-photochemical quenching. A response to excess light energy. Plant Physiology 125(4): 1558-1566 Demmig-Adams B, Adams WW (1996) The role of xanthophyll cycle carotenoids in the protection of photosynthesis. Trends in Plant Science 1(1): 21-26
Non Photochemical Quenching Quality
Pinot Noir rare functionally male flowers exhibit non-receptive dry stigmas that remain consistently dry and matte throughout the bloom period (days 0-7 post-calyptra shedding) despite hermaphrodite flowers in the same inflorescence showing wet glistening receptive stigmas during days 1-6, with controlled pollination applying viable pollen to non-receptive dry stigmas at day 2 (when hermaphrodite stigmas are maximally receptive) achieving only 8-14% germination and <6% fruit set, demonstrating stigma non-receptivity as primary cause of reproductive failure. Wild V. vinifera ssp. sylvestris staminate male flowers show completely non-receptive dry vestigial stigmas 0.12-0.22 mm diameter that never produce exudate and achieve <5% pollen germination throughout the bloom period, representing absolute non-receptivity where stigma developmental suppression prevents any receptivity capability. Cabernet Sauvignon experimental vines treated with inhibitors of secretory pathway during stigma development (BBCH 56-58) develop non-receptive dry stigmas that fail to produce exudate at anthesis, achieving <12% pollen germination and <8% fruit set despite normal hermaphrodite stamen and ovary development, demonstrating that stigma receptivity specifically requires functional secretory apparatus and that its absence creates non-receptive dry condition preventing successful pollination. Breeding populations segregating for flower sexual type show functionally male offspring with non-receptive dry stigmas identifiable during bloom through stigma appearance (dry when hermaphrodite siblings are wet) and germination bioassays (15-22% germination versus 78-86% for hermaphrodite siblings), allowing sexual type identification before fruit set assessment and enabling early selection against functionally male types in breeding programs targeting hermaphrodite cultivar development.
A non-receptive dry stigma is a dry stigma that fails to activate exudate secretion and achieve receptivity during the expected bloom period, remaining dry throughout the time when receptivity would normally occur, characteristic of functionally male and staminate flowers where genetic/developmental factors prevent normal stigma receptivity achievement, representing stigmas incapable of supporting successful pollination due to absence of receptivity rather than due to timing issues. Non-receptive dry stigmas are characterized by= (1) consistently dry matte appearance throughout the bloom period including the time when hermaphrodite flowers achieve receptivity (days 0-6 post-calyptra shedding) (2) absence or severe reduction of exudate secretion, with no wet glistening appearance developing at any point during expected receptivity period (3) papillae present but secretory apparatus absent, non-functional, or severely impaired, lacking the active endoplasmic reticulum, Golgi bodies, and secretory vesicles characteristic of receptive stigma papillae (4) inability to support pollen germination throughout bloom period, achieving 70% indicating receptivity), allowing identification of functionally male and staminate flowers based on stigma non-receptivity.
Non_Receptive_Dry_Stigma represents genetic/developmental failure to achieve receptivity and is distinguished from Post_Receptive_Dry_Stigma (which achieved receptivity but receptivity has ended) by absence of any receptive period versus presence of earlier receptivity. Non-receptive dry stigmas are characteristic of functionally male and staminate flowers and represent a fundamental functional deficit preventing successful pollination regardless of pollen availability or timing. Understanding non-receptive dry stigmas is essential for identifying functionally male and staminate flowers, for diagnosing the causes of reproductive failure, and for genetic studies of stigma receptivity mechanisms and sex determination.
dauerhaft trockene Narbe
nicht-rezeptive trockene Narbe
non-functional dry stigma
permanently dry stigma
unfähiges Stigma
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 96-101 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=95-106.
Non Receptive Dry Stigma
Certain individual vines within Merlot vineyards occasionally exhibit non-receptive functionally male flowers with near-normal pistil length (1.6-2 mm, only 10-15% shorter than hermaphrodite Merlot) and superficially normal flower morphology but dry stigma lacking wet exudate at anthesis, erect vigorous stamens producing abundant pollen (19,000-25,000 grains per flower, 68-76% viability), and consistent fruit set failure (<3%) despite favorable bloom conditions, with affected vines requiring careful observation over 2-3 seasons to confirm the sterility pattern before removal due to non-obvious morphological diagnosis. Breeding populations segregating for sex determination occasionally produce seedlings with non-receptive functionally male flowers showing hermaphrodite-like gross morphology but stigmatic papillae absent or malformed under microscopic examination, dry stigma surface at anthesis, and pollen germination failure on stigma surface even when pollen viability is confirmed on artificial media, creating cryptic male-sterile individuals requiring destructive flower sampling for definitive diagnosis.
A non-receptive functionally male flower is a functionally male flower in which female sterility is achieved primarily through stigma non-receptivity or ovule dysfunction rather than through gross pistil abbreviation, with pistil showing near-normal length and morphology but failing to support successful pollination, pollen germination, pollen tube growth, or fertilization, creating cryptic female sterility less morphologically obvious than the short pistil mechanism but equally effective in preventing fruit set. Non-receptive functionally male flowers are characterized by= (1) pistil length approaching normal hermaphrodite dimensions (often 70-95% of hermaphrodite pistil length, 1.3-2.2 mm), making superficial morphology similar to hermaphrodite flowers (2) stigma lacking the characteristic wet exudate (stigmatic secretion) that facilitates pollen grain hydration, adhesion, and germination, appearing dry or poorly developed (3) stigma papillae (specialized stigmatic surface cells) absent, reduced, or non-functional, preventing normal pollen-stigma recognition and interaction (4) style tissue that may be anatomically abnormal, lacking functional transmitting tract tissue necessary for pollen tube growth, or producing biochemical inhibitors that prevent pollen tube penetration and growth (5) ovules that may be aborted, sterile, or incapable of supporting embryo and endosperm development even if fertilization occurs (6) stamens showing normal hermaphrodite morphology with erect position, normal stamen-pistil length relationships, and abundant viable pollen production (16,000-28,000 grains per flower, 65-80% viability). The non-receptive mechanism represents cryptic female sterility, as flowers appear superficially hermaphrodite in gross morphology but fail to set fruit due to subtle cellular and biochemical defects in the female reproductive pathway. The stigma non-receptivity may result from developmental failures in stigmatic secretory cell differentiation, biochemical deficiencies in exudate composition preventing pollen hydration, or active rejection mechanisms similar to self-incompatibility systems. Style dysfunction may involve absent or malformed transmitting tract, defective extracellular matrix lacking pollen tube guidance factors, or production of pollen tube growth inhibitors. Ovule sterility may result from abnormal megasporogenesis, embryo sac developmental arrest, or defective nucellus or integument tissues. Non-receptive functionally male flowers achieve <1-5% fruit set despite normal stamen-pistil spatial relationships and abundant pollen production, with the low fruit set often puzzling to growers who observe normal-appearing flowers but complete fruit set failure. Diagnosis of non-receptive functionally male flowers requires detailed examination= (1) observation of dry stigma lacking exudate at anthesis (2) pollination experiments demonstrating pollen germination failure on the stigma (3) microscopic examination of stigma and style tissue showing anatomical abnormalities (4) fluorescence microscopy of pollinated pistils revealing absent or arrested pollen tube growth. The non-receptive condition is less readily diagnosed through casual visual observation compared to short pistil or reflex stamen mechanisms, potentially leading to retention of non-productive vines in commercial plantings until fruit set failure is observed over multiple seasons. The genetic and developmental basis likely involves genes regulating stigma development, secretory cell differentiation, transmitting tract formation, or ovule development, possibly overlapping with self-incompatibility and sex determination pathways.
Non_Receptive_Functionally_Male_Flower is distinguished from Short_Pistil_Functionally_Male_Flower by the mechanism of female sterility= cryptic stigma/ovule dysfunction versus obvious pistil morphological abbreviation. The non-receptive mechanism is less morphologically obvious, requiring detailed examination or multi-season fruit set observation for definitive diagnosis, whereas short pistil types are immediately identifiable visually at anthesis. Non-receptive types may be misidentified as hermaphrodite in casual observation, leading to vineyard management errors and delayed identification of non-productive genotypes. The cellular and molecular basis of stigma non-receptivity in Vitis deserves research attention, as understanding these mechanisms could inform studies of pollen-pistil interactions, self-incompatibility systems, and reproductive barriers in grapes. The non-receptive mechanism may represent a more evolutionarily recent or developmentally subtle pathway to functional male expression compared to the gross morphological changes in short pistil types.
Blüte mit funktionsloser Narbe
nicht-rezeptive männliche Blüte
non-receptive stigma flower
sterile Narbe Blüte
stigma-dysfunction male flower
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 102-104 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Caporali, E. et al. (2003). The arrest of development of abortive reproductive organs. Sexual Plant Reproduction 15=291-300 Ferrari, B.L. et al. (2016). Structural and developmental analyses of Vitis vinifera flower. American Journal of Plant Sciences 7=1742-1753.
Non Receptive Functionally Male Flower
Non-reducing sugar quality assumes importance in cryoconcentration processes for Icewine production in Canada, where freezing concentrates all sugars but the ratio of sucrose to reducing sugars may increase if invertase activity is inhibited by low temperatures during the final ripening period musts from frozen Vidal grapes typically show sucrose concentrations of 5-15 g/L (2-5% of total sugars) compared to less than 3 g/L in normally harvested fruit, a difference that affects fermentation initiation as yeast invertase must first hydrolyze sucrose before fermentation can proceed.
Non_Reducing_Sugar_Quality represents the chemical quality characterizing the concentration of sugars that lack free reducing groups and therefore cannot reduce oxidizing agents, primarily encompassing sucrose in grape berry tissue. Non-reducing sugars have their potential reducing groups (aldehyde or ketone) blocked through glycosidic bond formation, as in sucrose where the reducing groups of glucose and fructose are joined. In ripe Vitis vinifera berries, non-reducing sugars typically constitute less than 1% of total sugars because the active invertase enzymes in grape berries rapidly hydrolyze sucrose arriving through the phloem, releasing glucose and fructose that accumulate in vacuoles. However, sucrose concentrations can be transiently elevated in several situations: immediately after phloem unloading before invertase action is complete in certain cultivars with lower invertase activity in berries at early developmental stages before invertase is fully expressed and in some table grape varieties bred for specific texture or processing properties. The distinction between reducing and non-reducing sugars has practical relevance because yeast cannot directly ferment sucrose but must first hydrolyze it via invertase however, Saccharomyces cerevisiae produces abundant invertase, so sucrose is effectively fermented. For analytical purposes, non-reducing sugars are measured either directly (HPLC) or by difference (total sugars minus reducing sugars), or through specific enzymatic assays for sucrose using invertase followed by glucose determination. The non-reducing sugar fraction may increase relative importance in musts subjected to heating or concentration processes that can hydrolyze sucrose to reducing sugars.
Non_Reducing_Sugar_Quality is positioned parallel to Reducing_Sugar_Quality within Sugar_Content_Quality, with Sucrose_Content_Quality as its primary child class. The low proportion of non-reducing sugars in ripe berries reflects the biochemistry of sugar accumulation rather than transport, as sucrose is the dominant phloem-mobile sugar in grapevine. The distinction between reducing and non-reducing sugars is less functionally important for grape and wine production than in some other food applications, because the dominant non-reducing sugar (sucrose) is readily hydrolyzed by both plant and yeast invertases. Nevertheless, accurate sugar composition analysis requires separate determination of reducing and non-reducing fractions.
Disaccharidgehalt
Saccharoseanteil
disaccharide content
glycosidically-bound sugar
nicht-reduzierende Kohlenhydrate
nicht-reduzierende Zucker
non-reductive carbohydrates
sucrose fraction
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 267-285 Hawker JS (1969) Changes in the Activities of Enzymes Concerned with Sugar Metabolism during the Development of Grape Berries. Phytochemistry 8(1): 9-17
Non Reducing Sugar Quality
Norisoprenoid precursor accumulation occurs in all V. vinifera cultivars but shows substantial genotypic variation, with precursor concentrations ranging from 50-500 µg/L depending on cultivar and ripeness stage. Carotenoid cleavage by VvCCD1 and VvCCD4 generates C13 fragments that undergo further modification to produce β-damascenone (rose, apple, honey threshold 0.05 µg/L), β-ionone (violet, raspberry threshold 0.09 µg/L), and TDN precursors (kerosene threshold 2-20 µg/L). Riesling shows high TDN precursor accumulation (100-300 µg/L), while Chardonnay and Semillon show elevated β-damascenone potential. Norisoprenoid precursors increase during ripening, particularly during the final 2-4 weeks before harvest, with sun exposure enhancing both carotenoid synthesis and subsequent degradation.
An aromatic compound disposition that is the inherited capacity of a Vitis vinifera plant to generate and accumulate C13-norisoprenoid aroma compounds and their glycosidic precursors through oxidative cleavage of carotenoid pigments (C40) by carotenoid cleavage dioxygenases (CCDs), with genotype-specific concentrations of β-damascenone, β-ionone, TDN precursors, and related compounds contributing complex fruit, floral, honey, and kerosene notes.
Norisoprenoid_Accumulation_Disposition serves as the parent class for specific norisoprenoid precursor traits. Unlike monoterpenes that are synthesized de novo, norisoprenoids arise from carotenoid catabolism, linking color and aroma metabolism. The precursors are stored as glycosides and released through acid hydrolysis during wine aging, producing the characteristic "bottle-aged" aroma development. The low sensory thresholds of β-damascenone and TDN mean these compounds significantly impact wine aroma even at trace concentrations.
C13 aroma compound disposition
Carotinoid-Abbauprodukte
Norisoprenoid-Akkumulation
Norisoprenoid-Disposition
carotenoid-derived aroma trait
norisoprenoid synthesis capacity
Mathieu S et al. (2005) A carotenoid cleavage dioxygenase from Vitis vinifera L.: functional characterization and expression during grape berry development in relation to C13-norisoprenoid accumulation. Journal of Experimental Botany 56:2721-2731 Mendes-Pinto MM (2009) Carotenoid breakdown products - the norisoprenoids - in wine aroma. Archives of Biochemistry and Biophysics 483:236-245
Norisoprenoid Accumulation Disposition
Norisoprenoid content quality in Hunter Valley Semillon evolves dramatically during the 10-20 year bottle aging that transforms this variety from neutral youth to complex maturity, with β-damascenone rising from 1-3 μg/L in young wine to 5-10 μg/L in aged examples contributing honey-exotic notes, while TDN accumulates from below threshold (less than 4 μg/L) to 15-40 μg/L providing the distinctive petrol-kerosene character that defines aged Hunter Semillon alongside aged Riesling as the benchmark norisoprenoid-driven aromatic white wines.
Norisoprenoid_Content_Quality encompasses the chemical qualities characterizing the concentration and composition of C13-norisoprenoid compounds and their precursors in grape berries, capturing the carotenoid degradation products that contribute fruity, floral, exotic, and in aged wines, kerosene-like aromatic notes. Norisoprenoids are formed through oxidative cleavage of C40 carotenoid pigments (β-carotene, lutein, neoxanthin, violaxanthin), producing C13 fragments that undergo further modification to yield aroma-active compounds. The primary grape norisoprenoids include β-damascenone (exotic fruit, stewed apple, honey), β-ionone (violet, floral), vitispirane (camphor, eucalyptus), and TDN (1,1,6-trimethyl-1,2-dihydronaphthalene, petrol, kerosene). In grape berries, norisoprenoids occur predominantly as non-volatile glycosidic precursors, with free forms accumulating only at low concentrations. The glycosylated precursors can be hydrolyzed during winemaking, fermentation, and aging to release the volatile aglycones. Carotenoid concentration in grape berries, which serves as the substrate for norisoprenoid formation, depends on variety, maturity, and light exposure, with carotenoid degradation accelerating during ripening and continuing during wine aging. The norisoprenoid contribution to wine aroma evolves substantially during aging, with precursor hydrolysis and chemical rearrangement progressively releasing and modifying these compounds. TDN formation exemplifies this aging chemistry, accumulating progressively in Riesling through acid-catalyzed reactions to produce the characteristic petrol note of aged examples. The extremely low sensory thresholds of many norisoprenoids (β-damascenone approximately 50 ng/L, β-ionone approximately 90 ng/L) means they contribute significantly to wine aroma despite relatively low concentrations.
Norisoprenoid_Content_Quality serves as the parent class for specific norisoprenoid compounds including Beta_Damascenone_Precursor_Quality and TDN_Precursor_Quality. The carotenoid origin of norisoprenoids links their content to fruit ripening and sun exposure through effects on carotenoid metabolism. The class designation as Content_Quality addresses both free norisoprenoids and bound precursors, recognizing that precursor concentration better predicts potential aroma development than free forms alone. The aging evolution of norisoprenoid content distinguishes this compound class from stable aroma compounds that do not change during bottle maturation.
C13-Norisoprenoidgehalt
C13-norisoprenoid content
Megastigmangehalt
Norisoprenoidgehalt
carotenoid-derived aroma quality
karotinoidstämmige Aromaqualität
megastigmane content
norisoprenoid compound concentration
Winterhalter P and Rouseff RL (2002) Carotenoid-Derived Aroma Compounds: An Introduction. In: Winterhalter P and Rouseff RL (eds) Carotenoid-Derived Aroma Compounds, ACS Symposium Series 802 pp.: 1-17 Mendes-Pinto MM (2009) Carotenoid Breakdown Products the Norisoprenoids in Wine Aroma. Archives of Biochemistry and Biophysics 483(2): 236-245
Norisoprenoid Content Quality
Norisoprenoid data for Riesling from the Clare Valley, South Australia (2023 vintage) documents β-damascenone at 6.2 µg/L (OAV 124, sensory threshold 0.05 µg/L), TDN at 8.5 µg/L after 2 years bottle aging (OAV 0.4 at the 20 µg/L threshold, approaching perceivability), β-ionone at 0.65 µg/L (OAV 0.8), and carotenoid precursors in berries at harvest of β-carotene 0.45 mg/kg and lutein 0.38 mg/kg — with the high TDN development rate attributed to the warm climate and low pH (2.95) accelerating acid-catalyzed carotenoid degradation. Norisoprenoid data for Chardonnay from Meursault (2024 vintage) records β-damascenone at 4.8 µg/L (OAV 96) contributing rose-honey complexity to the aroma, with β-ionone at 0.42 µg/L providing subtle violet undertones, characterizing the norisoprenoid contribution to premier cru Burgundy white wine complexity.
Norisoprenoid data is an aromatic analysis data entity that records the concentration of C₁₃-norisoprenoid compounds and their carotenoid precursors in grape berries and wine, determined by GC-MS analysis of the volatile fraction and by HPLC analysis of carotenoid precursors, expressed in micrograms per liter (µg/L) for volatile norisoprenoids and milligrams per kilogram (mg/kg) for carotenoid precursors, encompassing β-damascenone, β-ionone, 1,1,6-trimethyl-1,2-dihydronaphthalene (TDN), vitispirane, actinidols, and their parent carotenoids (β-carotene, lutein, neoxanthin, violaxanthin), and providing the chemical characterization of the floral, fruity, honey, and kerosene-like aromatic dimensions that significantly contribute to wine complexity and aging bouquet.
Norisoprenoid data captures a class of aroma compounds whose significance for wine quality has been increasingly recognized since the identification of β-damascenone (rose, honey, exotic fruit) and TDN (kerosene, petrol) as major contributors to wine aroma profiles, particularly in Riesling and Chardonnay. The biosynthetic origin of norisoprenoids from carotenoid degradation creates a two-stage analytical challenge — the carotenoid precursors must be measured in berries at harvest (by HPLC with photodiode array detection), while the volatile norisoprenoids formed through enzymatic cleavage by carotenoid cleavage dioxygenases (VvCCD1, VvCCD4) and subsequent acid-catalyzed rearrangements are quantified in must and wine (by GC-MS, typically after SPME extraction). The extraordinary sensory potency of β-damascenone (threshold 0.05 µg/L in wine) means that this compound is virtually always present above its threshold in wine, making it a ubiquitous contributor to wine aroma complexity regardless of cultivar. TDN formation follows a distinctive pattern — minimal at bottling but progressively increasing during bottle aging through acid-catalyzed conversion of carotenoid-derived precursors, with formation rate accelerated by low pH (<3.1), higher storage temperature, and greater carotenoid precursor pool. The "petrol" or "kerosene" character of aged Riesling is primarily attributable to TDN, and its appreciation is culturally variable — valued in Australian and German Riesling traditions but sometimes considered a fault in other contexts. Vineyard factors influencing norisoprenoid potential include berry sun exposure (increased carotenoid synthesis with UV exposure, but also increased degradation), water stress (moderate deficit concentrates precursors), and vine nitrogen status (nitrogen-deficient vines accumulate more carotenoids). The interaction between norisoprenoid and terpene profiles creates the cultivar-specific aromatic signature — Muscat varieties are terpene-dominant, Riesling exhibits a balanced terpene-norisoprenoid character, and Chardonnay is predominantly norisoprenoid-influenced.
Apocarotinoid-Daten
C13-norisoprenoid data
Norisoprenoid-Daten
Norisoprenoide Analyse
apocarotenoid data
carotenoid-derived aroma data
carotinoidabgeleitete Aromadaten
norisoprenoid analysis
norisoprenoid composition
Mendes-Pinto MM (2009) Carotenoid breakdown products — the norisoprenoids — in wine aroma. Archives of Biochemistry and Biophysics 483(2): 236-245 Sefton MA et al. (2011) Occurrence, sensory impact, formation, and fate of damascenone in grapes, wines, and other foods and beverages. Journal of Agricultural and Food Chemistry 59(18): 9717-9746 Winterhalter P, Rouseff RL (2002) Carotenoid-Derived Aroma Compounds. ACS Symposium Series 802, American Chemical Society, pp.: 1-17
Norisoprenoid Data
A Cabernet Sauvignon vine in north-south rows in Maipo Valley, Chile bears a North_South_Row_Role providing balanced east-west exposure in the hot continental climate, with morning sun on the eastern fruiting zone and afternoon sun on the western side creating the even ripening valued for premium Bordeaux-style production. A Chardonnay vine in north-south rows in California's Central Coast serves in a North_South_Row_Role aligned with prevailing afternoon sea breezes, the row orientation enabling air movement through the canopy that moderates temperatures and reduces powdery mildew pressure. A Shiraz vine in north-south rows in Barossa Valley bears a North_South_Row_Role considered optimal for managing the region's hot afternoon conditions, with east-facing fruit benefiting from morning sun during cooler hours.
A row orientation role that inheres in a grapevine planted in rows aligned along the north-south axis (or approximately north-south within ±30°), creating east-facing and west-facing canopy exposures that receive morning and afternoon sun respectively as the sun traverses east to west. North-south row orientation is generally considered optimal for warm-to-hot climates, as the balanced exposure between morning and afternoon sun prevents the extreme afternoon heat accumulation that west-facing fruit in east-west rows would experience. The east-facing canopy side receives gentler morning sun when temperatures are moderate, enabling photosynthesis during favorable conditions, while the west-facing side receives more intense afternoon sun but only after the vine has been physiologically active throughout the morning. North-south rows enable airflow from typical prevailing wind patterns (often north-south oriented in many regions), improving canopy ventilation and reducing humidity-related disease pressure. The balanced exposure of north-south orientation tends to produce uniform fruit ripening across both canopy sides, simplifying harvest timing decisions compared to orientations with more asymmetric exposure patterns.
North_South_Row_Role represents the default recommendation for most warm-climate viticulture, based on the balanced exposure and airflow benefits this orientation provides. The advantages are most pronounced at lower latitudes (30-40°) where the sun path is high and east-west asymmetry in exposure would be substantial at higher latitudes the sun angle is lower and orientation effects differ. North-south orientation research has documented more uniform fruit composition between row sides compared to east-west rows, with reduced sunburn incidence and more consistent ripening. The practical implementation of north-south rows must often compromise with other factors—property boundaries, slope direction, existing road patterns—that may prevent ideal orientation, with many vineyards accepting orientation within 20-30° of true north-south as adequate. Climate change is reinforcing north-south preferences in warming regions, as the balanced exposure helps moderate heat accumulation compared to west-facing concentrations in east-west rows.
N-S orientation
N-S-Ausrichtung
Nord-Süd-Zeilen
meridional orientation
meridionale Orientierung
north-south alignment
north-south rows
Smart RE (1973) Sunlight interception by vineyards, American Journal of Enology and Viticulture 24: 141-147 Hunter JJ et al. (2016) Row orientation, South African Journal of Enology and Viticulture 37: 107-119 Spayd SE et al. (2002) Separation of sunlight and temperature effects, American Journal of Enology and Viticulture 53: 171-182
North South Row Role
The nuclear DNA sequence of Cabernet Sauvignon (Chin et al. 2016, diploid assembly of both haplotypes) revealed the extensive structural heterozygosity characteristic of Vitis vinifera — across the 590 Mb diploid assembly, the two parental haplotypes (derived from Cabernet Franc and Sauvignon Blanc, confirmed by Bowers and Meredith 1997) differ by approximately 1.1% at the single nucleotide level, contain over 8,000 structural variants larger than 50 bp including 3,200 insertions, 2,800 deletions, and 1,600 inversions, and show hemizygous gene content where approximately 12% of predicted genes are present on only one haplotype — this pervasive heterozygosity, maintained by obligate outcrossing and vegetative propagation, means that each Vitis vinifera cultivar represents a unique, fixed heterozygous genotype whose nuclear DNA sequence encodes the cultivar-specific combination of traits selected and maintained through centuries of clonal propagation.
A DNA sequence derived from the nuclear genome of Vitis vinifera, located on one of the 19 chromosomes (2n = 38) that comprise the diploid nuclear complement, encoding the vast majority of protein-coding genes, regulatory elements, repetitive sequences, and structural features that determine cultivar identity, phenotypic traits, and the heritable variation exploited in grapevine breeding and clonal selection.
Nuclear_DNA_Sequence documents the primary genetic information repository of the grapevine — the 19 chromosomes carrying approximately 30,000 protein-coding genes, extensive transposable element content (approximately 40% of genome length), and the polymorphic marker loci used for cultivar identification and genetic mapping. The nuclear genome of Vitis vinifera is notable for three features with direct viticultural significance: first, the paleohexaploid origin (a genome triplication event approximately 117 million years ago shared with all core eudicots) that produced paralogous gene families exploited in the evolution of berry-specific metabolic pathways (anthocyanin, stilbene, terpene biosynthesis) second, the extreme heterozygosity maintained by the outcrossing reproductive biology and fixed by vegetative propagation, meaning that the two alleles at most loci in any cultivar encode functionally different protein variants and third, the high proportion of disease resistance gene analogs (approximately 500 NBS-LRR genes) that, despite their abundance, provide inadequate resistance to the major fungal pathogens of European origin (Plasmopara viticola, Erysiphe necator) because these pathogens co-evolved with American Vitis species rather than V. vinifera.
Chromosomen-DNA-Sequenz
Kern-DNA-Sequenz
chromosomal DNA sequence
nuclear genome sequence
nuclear genomic data
nukleäre Genomsequenz
Jaillon O et al. (2007) The grapevine genome sequence suggests ancestral hexaploidization in major angiosperm phyla, Nature 449: 463-467 Chin CS et al. (2016) Phased diploid genome assembly with single-molecule real-time sequencing, Nature Methods 13: 1050-1054 Bowers JE and Meredith CP (1997) The parentage of a classic wine grape, Cabernet Sauvignon, Nature Genetics 16: 84-87 Liang Z et al. (2019) Whole-genome resequencing of 472 Vitis accessions, Nature Communications 10: 1190
Nuclear DNA Sequence
A Pinot Noir Dijon clone 667 vine maintained in the screenhouse facility at ENTAV-INRA Montpellier bears a Nuclear_Stock_Plant_Role, having undergone thermotherapy followed by meristem tip culture, with comprehensive testing including three years of biological indexing on woody indicators, ELISA panels for all regulated viruses, and RT-PCR confirmation—representing the ultimate source for all clone 667 material distributed through the French certification system. A Merlot clone 343 plant in the Foundation Plant Services nuclear collection serves in a Nuclear_Stock_Plant_Role, maintained in an insect-proof screenhouse with annual HTS (high-throughput sequencing) surveillance for both known and novel viral agents. A Welschriesling clone maintained by the Austrian AGES nuclear repository bears a Nuclear_Stock_Plant_Role, providing the genetic and phytosanitary foundation for Austrian certified production.
A certified mother plant role that inheres in a grapevine maintained as nuclear stock at the apex of the certification hierarchy, representing the ultimate source material from which all certified propagation derives through controlled multiplication. Nuclear stock plants have undergone comprehensive pathogen elimination treatments (thermotherapy, meristem culture, or combined approaches) and exhaustive testing protocols confirming freedom from all known graft-transmissible pathogens, maintained under maximum isolation (screenhouses, insect-proof facilities, geographically isolated locations) to prevent any reinfection. The "nuclear" designation reflects the concept of a protected core (nucleus) from which the propagation system radiates outward through successive multiplication tiers. Nuclear stock is synonymous with "Pre-Basic" in EU terminology and represents the most valuable and protected material in the certification system, typically maintained by specialized repositories (FPS, ENTAV, JKI, CSIRO) rather than commercial nurseries due to the intensive resources required for maintenance and testing.
Nuclear_Stock_Plant_Role represents the highest-value, highest-investment tier of grapevine propagation, where decades of clonal selection and pathogen elimination work are preserved in a small number of protected plants. Nuclear stock maintenance costs are substantial—screenhouse facilities, specialized personnel, comprehensive testing programs—justifying centralization at specialized repositories rather than commercial operations. The transition from biological indexing (1960s-1990s) to molecular testing (PCR, 1990s-2010s) to high-throughput sequencing (2010s-present) has progressively improved nuclear stock verification while expanding the list of detectable pathogens. Nuclear stock serves both conservation (maintaining tested clones of heritage varieties) and innovation (introducing newly selected or imported material into certification) functions within the propagation system.
Elitematerial
Kernbestand
Nuklearstock
Präbasis-Stock
clean stock plant
elite stock plant
nuclear stock vine
pre-basic stock plant
European Union (2020) Commission Implementing Directive 2020/177 Annex IV Golino DA et al. (2017) Foundation Plant Services, HortScience 52: 1190-1194 Martelli GP (2017) An overview on grapevine viruses, in Grapevine Viruses: Molecular Biology, Diagnostics and Management, Springer pp.: 31-46 Walter B, Martelli GP (1996) Sanitary Selection of the Grapevine, INRA Editions
Nuclear Stock Plant Role
The number of lobes description for Pinot Noir documents predominantly five-lobed leaves (OIV 068:4) — the terminal (central) lobe flanked by two upper lateral lobes and two lower lateral lobes, delineated by two pairs of lateral sinuses (upper and lower) incised to at least one-fifth of the half-blade width — with occasional three-lobed leaves occurring at basal shoot positions where the lower lateral sinuses are insufficiently deep to qualify as true lobes. This five-lobed pattern is the most common configuration in V. vinifera cultivars (approximately 55% of varieties), distinguishing the major five-lobed group (Pinot Noir, Cabernet Sauvignon, Riesling, Sangiovese) from the three-lobed group (Gamay Noir, Sémillon, some Muscat types) and the entire-to-weakly-lobed group (Rupestris du Lot, some American Vitis species used as rootstocks).
A mature leaf description component that records the number of distinct lobes on the fully expanded leaf blade according to OIV descriptor 068, scored as one (entire, unlobed), three, five, seven, or more than seven lobes, with lobe count determined by the number of sinuses incised to at least one-fifth of the half-blade width, providing one of the most fundamental and diagnostically powerful characters for classifying Vitis vinifera cultivars into major morphological groups.
Number_Of_Lobes_Description documents a character that is among the OIV Priority 1 descriptors and serves as the primary classifier for partitioning the morphological diversity of V. vinifera leaves into major groups. Lobe count reflects the developmental patterning of marginal growth zones during leaf primordium expansion — lobes form where localized growth maxima in the leaf margin produce outward projections separated by growth minima (sinuses), with the number and position of these growth maxima determined by the interaction of auxin transport, CUC (CUP-SHAPED COTYLEDON) transcription factor expression, and other developmental regulators: QTL mapping by Chitwood et al. (2014) and Demmings et al. (2019) identified loci on chromosomes 1, 2, 5, and 14 contributing to lobe number variation, consistent with polygenic control modulated by a limited number of major-effect loci that determine the fundamental three-versus-five-versus-seven-lobed architecture. The scoring protocol requires that lobes be counted only when the delineating sinus is incised to at least one-fifth of the half-blade width — this threshold criterion addresses the ambiguity of shallow sinuses that create marginal undulations without producing morphologically distinct lobes, ensuring consistent scoring across observers.
Blattlappen-Zählung
Blattzipfel-Anzahl
Lappenzahl-Beschreibung
blade lobation record
leaf lobe count
lobe number assessment
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Chitwood DH et al. (2014) A modern ampelography: a genetic basis for leaf shape and venation patterning in grape, Plant Physiology 164: 259-272 UPOV (2011) Guidelines for the Conduct of Tests for Distinctness, Uniformity and Stability — Vine (Vitis L.), TG/50/9
Number Of Lobes Description
Nursery distribution for Cabernet Sauvignon clone 337 in California lists 12 licensed propagation nurseries including Sunridge Nurseries (Bakersfield, annual capacity 250,000 bench grafts of CS 337), Novavine (Santa Rosa, capacity 80,000 potted vines), and Inland Nursery (Escalon, capacity 150,000 dormant bench grafts), with Sunridge and Novavine additionally maintaining CDFA-registered mother blocks as foundation material sources for the state. Nursery distribution for Grüner Veltliner clone 17 in Austria documents propagation by Rebschule Tschida (Hagenbrunn, primary supplier for Lower Austria), Rebschule Rittsteuer (Horitschon, primary supplier for Burgenland), and the Höhere Bundeslehranstalt Klosterneuburg (institutional foundation block holder), with combined annual production capacity of approximately 120,000 grafted vines supplying the domestic market.
Nursery distribution is a distribution record entity that identifies the specific nursery enterprises, propagation facilities, and institutional repositories authorized to multiply, graft, and sell certified plant material of a clonal line, including the nursery name, location, licensing status, annual production capacity for the clone, the form of material supplied (bench grafts, potted vines, dormant rootlings, green grafts), and the nursery's position within the multiplication chain (foundation block holder, mother block operator, or commercial propagator), establishing the supply chain through which growers access certified clonal material.
Nursery distribution documentation serves both commercial and regulatory functions — commercially, it enables growers to identify and source specific clonal material, while regulatorily, it establishes the chain of custody required for phytosanitary traceability from foundation stock to planted vineyard. The nursery industry structure varies substantially between wine-producing countries — France operates a highly centralized system where ENTAV maintains all foundation stocks and licenses multiplication to approximately 400 registered pépiniéristes, while in Australia the system is more decentralized with multiple independent vine improvement groups maintaining foundation plantings. The form of plant material (bench graft, potted vine, green graft, or rootling) supplied by each nursery influences establishment success, first-bearing timing, and cost, making nursery supply form a relevant component of distribution documentation. Nursery production lead times of 1-3 years between order and delivery create supply constraints for newly certified or suddenly popular clones, and distribution records should document waiting list status and advance order requirements. The consolidation trend in the nursery industry, with fewer but larger operations serving broader geographic markets, is altering clone availability patterns — large nurseries offer wider clone selections but may discontinue low-volume selections that smaller specialized nurseries previously maintained. Nursery distribution data should be updated periodically, as nursery licensing status, production capacity, and clone inventory change in response to market demand and business conditions.
Baumschulvertrieb
Pflanzgutlieferanten
Rebschulverteilung
Rebschulvertrieb
Vermehrungsbetriebnachweise
clone supplier documentation
licensed nursery list
nursery source record
nursery supply record
propagation nursery data
Nicholas P (2004) Grapevine propagation. In Viticulture Volume 1: Resources, Winetitles, pp.: 213-235 Boidron R et al. (1995) Catalogue des variétés et clones de vigne cultivés en France, ENTAV-INRA, pp.: 25-35 Mannini F (2004) Clonal selection in grapevines. Acta Horticulturae 652: 21-27
Nursery Distribution
A one-year-old grafted vine (Pinot Noir on 3309C) produced in a Burgundy nursery bears a Nursery_Plant_Role, graded by root count and distribution, trunk caliper, and graft union quality before sale as certified plant greffé-soudé for vineyard establishment. A rooted cutting of 110 Richter serves in a Nursery_Plant_Role as rootstock liner destined for field budding with customer-specified scion cultivars the following growing season. A potted green-growing Chardonnay/SO4 vine bears a Nursery_Plant_Role in accelerated vineyard establishment programs, planted in late spring with active growth enabling first-year establishment impossible with dormant material.
A propagation role that inheres in a grapevine plant produced by nursery operations for sale to growers for vineyard establishment, representing the commercial output of the propagation system. Nursery plants encompass the range of product formats produced by grapevine nurseries—from simple rooted cuttings (rootstocks for field budding) through grafted vines in various conditioning states (dormant bare-root, potted, green-growing)—with format selection determined by production system capabilities, market preferences, planting timing requirements, and price positioning. Nursery plant quality is assessed by root system development (quantity, distribution, and health of roots), shoot system condition (viable buds, absence of damage or disease), graft union integrity (for grafted plants—complete callusing, mechanical strength, vascular continuity), and overall plant vigor (size, carbohydrate reserves, freedom from pathogens). Certification status (Pre-Basic, Basic, Certified, Standard/CAC) is inherited from source material and documented through labeling and traceability systems. The global grapevine nursery industry produces hundreds of millions of plants annually, supporting vineyard establishment, renovation, and expansion worldwide.
Nursery_Plant_Role is subclassed by plant format (rooted cutting, grafted vine) and conditioning state (dormant, potted, green-growing, bare-root), reflecting the diverse products offered by the nursery industry to meet varying customer requirements. The category represents the commercial interface between propagation operations and grape growers, with plant quality directly impacting vineyard establishment success, early productivity, and long-term performance. Quality grading systems vary by region but typically assess root development, shoot condition, and graft union quality through visual inspection and increasingly through standardized measurements. Certification requirements mandate labeling with variety/clone identification, certification category, nursery identification, and production year, providing traceability essential for quality assurance and phytosanitary tracking.
Jungpflanze
Pflanzrebe
Rebschulpflanze
Setzling
nursery stock
nursery vine
planting material
vine transplant
European Union (2020) Commission Implementing Directive 2020/177 Annex I Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson Gramaje D, Armengol J (2011) Fungal trunk pathogens, Phytopathologia Mediterranea 50: S217-S226 Waite H et al. (2015) Hot water treatment, Phytopathologia Mediterranea 54: 325-339
Nursery Plant Role
Pinot Noir showing nutrient absorptive feeder roots, isotope study using 15N-nitrate showing feeder roots providing 89% of vine nitrogen uptake, 33P-phosphate demonstrating 94% uptake through feeder roots with mycorrhizae, feeder roots showing upregulated NRT2 nitrate transporter expression under low N conditions, arbuscular mycorrhizal colonization 67% enhancing phosphorus acquisition, nutrient absorptive feeder roots demonstrating specialized function acquiring minerals from soil supporting vine nutrition
Nutrient absorptive feeder root is absorptive feeder root emphasizing nutrient uptake function and mineral acquisition, characterized by specialized transport systems for acquiring nitrogen, phosphorus, potassium, calcium, magnesium, and micronutrients from soil, showing nutrient-specific transporters including NRT nitrate transporters, AMT ammonium transporters, PHT phosphate transporters, HAK potassium transporters, and various micronutrient carriers, demonstrating particularly important role for immobile nutrients like phosphorus requiring close soil contact and mycorrhizal partnerships, and serving as primary nutrient acquisition interface between vine and soil providing 80-95% of mineral nutrient uptake supporting growth, metabolism, and fruit production, with nutrient absorptive feeder roots showing functional characteristics including high specific nutrient uptake rates particularly for nitrogen and phosphorus, dense arbuscular mycorrhizal colonization especially important for phosphorus acquisition, rhizosphere acidification and organic acid exudation mobilizing nutrients, rapid proliferation in fertilizer-enriched zones demonstrating nutrient responsiveness, and demonstrated uptake through isotope tracers showing preferential accumulation of labeled nutrients. Nutrient absorptive feeder root nitrogen uptake includes= showing nitrate acquisition (NRT1 and NRT2 nitrate transporters taking up NO3- from soil solution), exhibiting ammonium uptake (AMT ammonium transporters absorbing NH4+ particularly important in acidic soils), demonstrating preferential forms (grapevine generally preferring nitrate though capable of using both forms), revealing high-affinity transport (transporters showing Michaelis-Menten kinetics with high-affinity systems active at low concentrations), and displaying major contribution (nutrient absorptive roots providing 85-95% of total vine nitrogen acquisition). Nutrient absorptive feeder root phosphorus uptake includes= showing PHT transporter function (phosphate transporters taking up H2PO4- and HPO4²- from soil solution), exhibiting mycorrhizal importance (arbuscular mycorrhizae particularly critical for P given low soil mobility and availability), demonstrating acidification effects (proton extrusion potentially solubilizing phosphorus from minerals), revealing organic acid secretion (citrate, malate, oxalate potentially chelating and mobilizing phosphorus), and displaying critical role (nutrient absorptive roots with mycorrhizae providing 90-98% of vine phosphorus uptake). Nutrient absorptive feeder root potassium uptake includes= showing channel-mediated uptake (inward-rectifying and outward-rectifying K+ channels allowing facilitated diffusion), exhibiting HAK transporter function (high-affinity K+ transporters active at low soil K concentrations), demonstrating high uptake rates (potassium showing highest uptake rates among nutrients reflecting high vine demand), revealing luxury consumption (nutrient absorptive roots capable of accumulating K beyond immediate needs if available), and displaying major pathway (nutrient absorptive roots providing 80-90% of vine potassium acquisition). Nutrient absorptive feeder root calcium and magnesium uptake includes= showing calcium channels (calcium-permeable channels allowing Ca2+ uptake though rates typically lower than other cations), exhibiting magnesium transporters (Mg2+ transport systems acquiring magnesium for chlorophyll and enzyme functions), demonstrating passive components (some Ca and Mg uptake potentially passive following electrochemical gradients), revealing adequacy dependence (uptake capacity generally adequate meeting vine Ca and Mg needs in normal soils), and displaying secondary importance (Ca and Mg acquisition typically less limiting than N, P, K uptake). Nutrient absorptive feeder root micronutrient uptake includes= showing iron acquisition (specific Fe uptake systems including reduction and chelation mechanisms particularly important in calcareous soils), exhibiting zinc transporters (ZIP family zinc transporters acquiring Zn essential for enzymes and growth), demonstrating manganese uptake (Mn transport systems though excessive Mn potentially toxic), revealing copper and boron acquisition (specific transport systems for Cu and B micronutrients), and displaying critical functions (micronutrient uptake essential for vine health though required amounts small). Nutrient absorptive feeder root uptake regulation includes= showing demand-responsive expression (nutrient transporter genes upregulated when vine nutrient demand high or availability low), exhibiting feedback inhibition (high internal nutrient status potentially downregulating uptake capacity preventing excessive accumulation), demonstrating hormonal regulation (auxin, cytokinins, and other hormones affecting nutrient transporter expression and activity), revealing circadian patterns (some nutrient uptake potentially showing diurnal rhythms), and displaying developmental control (nutrient uptake capacity varying with root age and developmental state). Nutrient absorptive feeder root spatial foraging includes= showing fertilizer response (nutrient absorptive roots proliferating intensively in fertilizer-enriched zones), exhibiting patch exploitation (roots increasing density where nutrients abundant creating nutrient depletion halos), demonstrating preferential placement (root architecture placing nutrient absorptive roots where fertilizers applied), revealing competition (nutrient absorptive roots competing with weeds or cover crops for available nutrients), and displaying mycorrhizal networks (fungal hyphae from nutrient absorptive roots exploring large soil volumes accessing nutrients beyond root reach). Nutrient absorptive feeder root in vine nutrition includes= determining nutrient status (nutrient absorptive root capacity limiting vine foliar nutrient concentrations and tissue status), enabling growth support (adequate nutrient uptake supporting vigorous vegetative growth and fruit development), demonstrating deficiency sensitivity (inadequate nutrient absorptive root function contributing to nutrient deficiencies and symptoms), revealing fertilization response (fertilizer effectiveness depending on nutrient absorptive root distribution and function), and displaying diagnostic value (nutrient absorptive root assessment potentially indicating vine nutritional management needs).
Nutrient_Absorptive_Feeder_Root emphasizes mineral nutrient acquisition and is distinguished from water absorption by focus on ion uptake. Critical for understanding nutrient transporters, N-P-K uptake, micronutrient acquisition, mycorrhizal importance, rhizosphere mobilization, and the specialized roots providing 80-95% of vine mineral nutrient uptake.
Ion-uptake root
Ionen-Aufnahme-Wurzel
Mineral-absorbing root
Mineralstoff-aufnehmende Wurzel
Nutrient-acquiring feeder root
Nährstoff-absorbierende Feinwurzel
Nutrient Absorptive Feeder Root
Syrah in Rhône Valley shows nutrient accumulation during August-September Year N+1, with endosperm cells accumulating storage proteins 18-26% dry weight in protein bodies 1.8-3.4 micrometers diameter, lipids 14-22% in lipid bodies 0.7-1.4 micrometers, minor starch 6-12% in amyloplasts, progressive accumulation weeks 6-12 with peak rate 8-14% dry weight per week, final endosperm reserves supporting 3.8-4.6 mm embryo development and providing germination nutrition, under warm Mediterranean conditions of 24-29°C temperature, typical of this mid-to-late season cultivar completing endosperm storage accumulation
Nutrient accumulation represents the storage reserve deposition phase during endosperm development when cellular endosperm accumulates storage compounds including proteins, lipids, and carbohydrates creating nutritive tissue supporting embryo development and providing germination reserves, with proteins accumulating 15-30% endosperm dry weight as major storage component, lipids accumulating 10-25%, and reserves storing in specialized organelles including protein bodies and lipid bodies, occurring weeks 4-12 post-fertilization establishing endosperm storage function. This reserve deposition phase occurs during mid-late summer to early autumn in August-September in temperate regions Year N+1, representing major endosperm accumulation period coordinating with embryo development. Accumulation mechanism involves storage compound biosynthesis from imported photosynthate, protein synthesis and deposition in protein bodies, lipid synthesis and storage in lipid bodies, possible carbohydrate accumulation including starch, organelle biogenesis creating storage compartments, and progressive accumulation establishing reserve levels supporting embryo and germination. Photosynthate import shows carbohydrates imported from parent plant through seed coat and funiculus, sucrose as major transported carbohydrate, import providing carbon and energy for biosynthesis, and import as substrate source for storage compound synthesis. Import pathway shows phloem delivery to seed, unloading in seed coat, transfer through integuments to endosperm, and endosperm cells taking up substrates for metabolism. Protein accumulation characteristics shows storage proteins as major endosperm reserves, typical protein content 15-30% endosperm dry weight varying with genotype and conditions, proteins synthesized on endoplasmic reticulum ribosomes, proteins packaged in protein storage vacuoles forming protein bodies, and proteins providing amino acid reserves. Storage protein types shows albumins as water-soluble proteins, globulins as salt-soluble proteins, protein composition balanced for nutritional value, and protein types characteristic of seed storage proteins. Protein biosynthesis shows amino acids incorporated into polypeptide chains, post-translational modifications including glycosylation possible, protein folding and oligomerization, and synthesized proteins targeted to protein bodies. Protein body formation shows protein storage vacuoles forming as specialized organelles, protein bodies 1-5 micrometers diameter in endosperm cells, proteins crystalline or amorphous within bodies, tonoplast membrane surrounding protein bodies, and protein bodies as protein storage compartments. Protein body characteristics shows bodies containing concentrated storage proteins, proteins densely packed, bodies stable during desiccation, proteins mobilized during germination through proteolysis, and protein bodies as efficient storage structures. Lipid accumulation characteristics shows lipids accumulating in endosperm though typically less than embryo cotyledon levels, lipid content 10-25% endosperm dry weight, triacylglycerols as major storage lipids, fatty acids similar to embryo including oleic and linoleic acids, lipids synthesized in endoplasmic reticulum, and lipids stored in lipid bodies. Lipid body formation shows lipid bodies budding from endoplasmic reticulum, lipid bodies 0.5-2 micrometers diameter, triacylglycerols in core, phospholipid monolayer and proteins at surface, and lipid bodies as lipid storage organelles. Lipid body characteristics shows bodies distributed in cytoplasm, stable lipid storage form, lipids mobilized during germination through lipolysis, and lipid bodies providing concentrated energy reserves. Carbohydrate accumulation shows possible starch accumulation though extent variable in grapevine endosperm, starch content often low possibly 5-15% when present, amyloplasts as starch storage organelles if present, other carbohydrates including cell wall polysaccharides, and carbohydrates as potential though minor reserves compared to proteins and lipids. Starch characteristics shows α-amylose and amylopectin as starch components if present, starch grains in amyloplasts, starch mobilized through amylase activity during germination, and starch as glucose polymer storage form. Cell wall polysaccharides shows cellulose, hemicelluloses, pectins in endosperm cell walls, cell walls structural not primarily storage, some mobilization possible during germination, and cell walls as structural carbohydrates. Minor components shows minerals, vitamins, phytic acid as phosphorus storage, phenolic compounds, and diverse metabolites contributing to endosperm composition. Accumulation timing shows accumulation initiating weeks 4-6 after proliferation, intensive accumulation weeks 6-12, accumulation slowing and completing weeks 12-14, and timing coordinating with embryo reserve accumulation. Accumulation rate shows progressive increase in storage compound content, typical accumulation 5-15% dry weight per week during peak accumulation, and rate determining final reserve levels. Subcellular organization shows endosperm cells packed with protein bodies, lipid bodies, possible amyloplasts, and storage organelles occupying most cell volume. Cell metabolism during accumulation shows active biosynthetic metabolism, high respiratory rate supporting synthesis, substantial ATP requirement for polymerization, and metabolism focused on reserve production. Nutrient support for embryo shows endosperm possibly transferring nutrients to developing embryo, transfer supporting embryo growth and reserve accumulation, transfer mechanisms including cellular connections or secretion-uptake, and support function as key endosperm role. Endosperm size during accumulation shows endosperm maintaining volume around developing embryo, endosperm tissue layer 100-500 micrometers thick surrounding embryo, and endosperm mass increasing from reserve accumulation. Endosperm composition shows protein, lipid, carbohydrate balance determining nutritional quality, balance varying with genotype and conditions, composition affecting germination vigor, and composition as seed quality determinant. Reserve quality shows factors including protein amino acid balance, fatty acid composition, mineral content affecting nutritional value and germination success, and quality determined by accumulation conditions. Environmental effects on accumulation include temperature affecting biosynthetic rate and final reserve levels, nutrient availability affecting accumulation quantity and quality, water availability supporting normal development, and favorable conditions maximizing reserve accumulation. Temperature effects shows optimal range 22-26°C supporting normal accumulation, suboptimal temperatures potentially reducing reserves, and temperature as accumulation rate factor. Nutrient availability shows adequate nitrogen supporting protein accumulation, adequate minerals supporting complete nutrition, and nutrition affecting reserve quality. The process represents second phase of endosperm development establishing storage reserves, following endosperm proliferation and preceding endosperm maturation. Duration approximately 8 weeks representing major accumulation weeks 4-12. Regional timing shows nutrient accumulation during August-September coordinating with embryo maturation. Viticultural significance includes representing endosperm storage reserve establishment, supporting embryo development through nutrient transfer, providing germination reserves, and determining seed nutritional quality. Research applications include analyzing storage compound biosynthesis and regulation, investigating accumulation patterns and organelle formation, evaluating environmental effects on reserve accumulation, quantifying endosperm composition and nutritional value, and assessing nutrient accumulation as seed quality determinant.
Nutrient_Accumulation is second phase of Endosperm_Development, distinguished by storage reserve deposition. Represents weeks 4-12. Major reserves: proteins 15-30% (protein bodies 1-5 μm), lipids 10-25% (lipid bodies 0.5-2 μm), minor carbohydrates 5-15%. Accumulation rate 5-15% DW/week peak. Supports embryo development. Provides germination reserves. Duration ~8 weeks. Follows Endosperm_Proliferation. Precedes Endosperm_Maturation. Temperature 22-26°C optimal.
Endosperm-Reserven-Deposition
Nährstoff-Akkumulation
Nährstoffspeicherung
Speicherreserven-Akkumulation
compound accumulation
endosperm reserve deposition
nutrient storage
storage reserve accumulation
Nutrient Accumulation
Nutrient acquisition efficiency varies substantially among rootstocks, with 1103 Paulsen showing 40-60% higher phosphorus uptake than 101-14 Mgt under identical soil conditions. Total seasonal nutrient removal by grapevines ranges from 30-80 kg N/ha, 5-15 kg P/ha, and 40-100 kg K/ha depending on yield level and cultivar. Root system architecture strongly influences acquisition, with deep-rooted rootstocks accessing subsoil nutrients unavailable to shallow-rooted types. The rhizosphere pH modification capacity varies 0.5-1.5 pH units among genotypes, affecting micronutrient availability.
A grapevine disposition that is the inherited capacity of a Vitis vinifera plant to absorb, transport, and assimilate mineral nutrients from the soil solution through root system activity, encompassing traits related to uptake efficiency, ion selectivity, and the ability to access nutrients under varying soil conditions and availability levels.
Nutrient_Acquisition_Disposition serves as the top-level parent class for all traits related to mineral nutrition uptake. The category encompasses both macronutrient and micronutrient acquisition, as well as mycorrhizal associations that enhance nutrient access. Rootstock genotype exerts primary control over nutrient acquisition in grafted vines, though scion genotype influences demand patterns and sink strength. The trait complex is critical for sustainable viticulture, enabling reduced fertilizer inputs through selection of efficient genotypes.
Mineralstoffakquisition
Nährstoffaneignungsvermögen
Nährstoffaufnahme
mineral acquisition trait
nutrient foraging disposition
nutrient uptake capacity
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 295-345 Keller M et al. (2001) Rootstock effects on deficit-irrigated winegrapes in a dry climate: vigor, yield formation, and fruit ripening. American Journal of Enology and Viticulture 52:40-49
Nutrient Acquisition Disposition
Rootstocks dramatically influence scion potassium nutrition, with 140 Ruggeri (V. berlandieri × V. rupestris) delivering 30-50% higher petiole K concentrations to Nero d'Avola than SO4 on identical Sicilian soils, resulting in elevated must pH (3.6-3.9 versus 3.3-3.5) and reduced wine acidity. 1103 Paulsen enhances nitrogen acquisition supporting vigorous Primitivo canopy development in Puglia, with petiole N concentrations 15-25% higher than on 110 Richter under equivalent fertilization. Riparia Gloire de Montpellier shows efficient magnesium uptake maintaining Pinot Noir petiole Mg above 0.3% on the acidic granitic soils of Beaujolais where other rootstocks induce Mg deficiency.
A rootstock role that inheres in a grafted rootstock by virtue of its capacity to influence the uptake, translocation, and delivery of mineral nutrients to the scion, thereby affecting scion nutritional status, vegetative growth, fruit composition, and wine quality. Nutrient acquisition rootstock roles reflect genetically determined differences in root system architecture (depth, density, surface area), root membrane transporter activity and selectivity, rhizosphere modification capacity (organic acid exudation, pH alteration), mycorrhizal association efficiency, and xylem loading mechanisms. Rootstocks differ substantially in their acquisition efficiency for macronutrients (N, P, K, Ca, Mg) and micronutrients (Fe, Mn, Zn, Cu, B), with these differences significantly impacting scion performance independent of soil nutrient availability. Nutrient acquisition characteristics are quantified by leaf petiole nutrient concentrations at key phenological stages (bloom, veraison), by nutrient uptake per unit root biomass, and by scion yield and quality responses to fertilization across rootstocks.
Nutrient_Acquisition_Rootstock_Role is subclassed by specific nutrient efficiency characteristics, currently including potassium (high and low uptake), iron, magnesium, and nitrogen efficiency. These are not mutually exclusive a single rootstock may bear multiple nutrient acquisition roles. Nutrient acquisition interacts strongly with soil type, irrigation management, and scion cultivar demands the same rootstock may show different nutrient delivery characteristics across sites. The practical importance of rootstock nutrient effects is particularly evident in must and wine composition, where rootstock-mediated differences in K, N, and Mg nutrition influence pH, acidity, fermentation kinetics, and color stability.
Nährstoffaneignungs-Unterlage
mineral nutrition rootstock
mineralstoffeffiziente Unterlage
nutrient acquisition rootstock
nutrient efficiency rootstock
nutrient uptake rootstock
nährstoffaufnehmende Unterlage
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 285-320 Kodur S (2011) Effects of rootstocks on grapevine growth, yield and quality of wine, Journal of Wine Research 22: 155-168 Ibacache A et al. (2016) Rootstock effects on mineral nutrition of the scion, Acta Horticulturae 1136: 117-122
Nutrient Acquisition Rootstock Role
The nutrient analysis data for a Marlborough Sauvignon Blanc vineyard (New Zealand) documents the complete soil nutrient profile at 0-30 cm depth — total nitrogen 0.18% (moderate), Olsen-extractable phosphorus 32 mg/kg (adequate), exchangeable potassium 0.45 cmol(+)/kg (adequate), exchangeable calcium 8.5 cmol(+)/kg (adequate), exchangeable magnesium 1.8 cmol(+)/kg (adequate), sulfate-sulfur 12 mg/kg (adequate), DTPA-extractable iron 52 mg/kg (adequate), DTPA-manganese 15 mg/kg (adequate), DTPA-zinc 2.8 mg/kg (adequate), hot water-extractable boron 1.2 mg/kg (adequate), DTPA-copper 3.8 mg/kg (adequate, elevated above natural background of 0.5-1.5 mg/kg due to historic copper-based fungicide accumulation). This comprehensive nutrient profile, interpreted against New Zealand viticultural calibration standards, indicates no nutritional limitations requiring correction — the soil provides adequate nutrition for balanced vine growth without supplemental fertilization other than a maintenance nitrogen application of 20-30 kg N/ha/year to sustain moderate canopy development for the vigorous Sauvignon Blanc cultivar on this deep, fertile alluvial soil.
A soil chemistry data component that records the concentrations of individual essential plant nutrients in the vineyard soil, determined by standardized extraction methods — including the macronutrients nitrogen (N), phosphorus (P), potassium (K), calcium (Ca), magnesium (Mg), and sulfur (S), and the micronutrients iron (Fe), manganese (Mn), zinc (Zn), boron (B), copper (Cu), and molybdenum (Mo) — providing the analytical basis for diagnosing nutrient deficiencies, toxicities, and imbalances that constrain vine performance, and for calculating fertilizer applications that correct identified nutritional limitations.
Nutrient_Analysis_Data documents the individual nutrient concentrations that, interpreted against calibrated sufficiency ranges specific to grapevine and regional soil types, enable the precise diagnosis and correction of nutritional limitations. The standardized extraction methods used for each nutrient are critical to the interpretation — different extractants measure different pools of nutrient availability (total, exchangeable, plant-available, or solution-phase), and the sufficiency ranges are calibrated for specific extractants, making method specification essential for data comparability: the most widely used methods in viticultural soil analysis include ammonium acetate extraction for exchangeable cations (K, Ca, Mg, Na), Olsen bicarbonate or Bray-1 fluoride extraction for available phosphorus (the choice determined by soil pH — Olsen for alkaline soils, Bray-1 for acid soils), DTPA chelate extraction for available micronutrients (Fe, Mn, Zn, Cu), hot water extraction for available boron, and total Kjeldahl or Dumas combustion for total nitrogen. The viticultural calibration ranges differ from conventional agricultural standards because grapevine nutritional requirements are generally lower than those of annual crops, and because excessive nutrition (particularly nitrogen) is detrimental to wine grape quality — the viticultural sufficiency range for nitrogen, for example, targets moderate vine growth with leaf nitrogen of 2.0-2.5% (dry weight) at véraison, substantially below the 3.0-4.0% nitrogen considered optimal for maximum yield of annual crops. Soil nutrient analysis is complemented by tissue analysis (petiole or leaf blade nutrient concentrations at bloom or véraison) to provide a complete picture of vine nutritional status that accounts for the soil-root-vine transfer efficiency that varies with rootstock, soil physical conditions, and moisture regime.
Bodennährstoff-Profil
Mineralnährstoff-Analytik
Nährstoffanalyse-Daten
mineral nutrient analysis
soil fertility data
soil nutrient profile
Robinson JB (2005) Critical plant tissue values and application of nutritional standards for practical use in vineyards, in Christensen LP and Smart DR (eds) Proceedings of the Soil Environment and Vine Mineral Nutrition Symposium, ASEV White RE (2003) Soils for Fine Wines, Oxford University Press Conradie WJ (2001) Timing of nitrogen fertilisation, South African Journal of Enology and Viticulture 22: 60-68 Mpelasoka BS et al. (2003) A review of potassium nutrition in grapevines, Australian Journal of Grape and Wine Research 9: 154-168
Nutrient Analysis Data
Nutrient deficiency indicator quality assessment in a declining Napa Valley Cabernet Sauvignon block employed systematic visual diagnosis combined with tissue and soil analysis to identify the nutritional factors limiting vine performance. Visual nutrient deficiency indicator quality assessment identified multiple concurrent deficiency symptoms including generalized pale green coloration suggesting nitrogen limitation, interveinal chlorosis on young leaves suggesting iron or manganese deficiency, and marginal leaf necrosis on mature leaves suggesting potassium deficiency. Petiole tissue analysis at bloom confirmed the visual diagnosis, with nitrogen at 0.45% (deficient, threshold 0.6%), iron at 28 ppm (low-normal), and potassium at 1.2% (deficient, threshold 1.5%), establishing that multiple nutrient deficiency indicator quality symptoms reflected actual nutritional limitations requiring correction. Soil analysis revealed pH 7.8 potentially limiting iron availability despite adequate total soil iron, high calcium carbonate content contributing to induced iron chlorosis, and low exchangeable potassium reflecting the volcanic soil parent material. The comprehensive nutrient deficiency indicator quality assessment enabled targeted corrective actions including nitrogen fertigation, foliar iron chelate application, and potassium sulfate soil amendment, with follow-up tissue analysis confirming improved nutrient status and visual symptom resolution by the following season.
Nutrient deficiency indicator quality is a stress response quality that inheres in a grapevine organism and represents the ensemble of visual symptoms, growth abnormalities, and physiological changes that signal inadequate availability of essential mineral nutrients for normal plant function, with deficiency symptoms providing diagnostic information about the specific nutrients limiting vine performance and guiding corrective fertilization or soil amendment decisions. This quality captures the plant's response to nutritional stress, recognizing that grapevines require adequate supplies of macro- and micronutrients for optimal growth, fruit production, and wine quality, and that deficiency of any essential element produces characteristic symptoms that enable diagnosis of the limiting factor affecting vine performance. The ontological significance of nutrient deficiency indicator quality lies in its role enabling identification of nutritional limitations before severe yield or quality losses occur, supporting precision nutrient management that supplies deficient elements while avoiding over-fertilization with elements already adequate, thereby optimizing both production and environmental outcomes. The biological basis of nutrient deficiency indicators encompasses the disruption of metabolic processes requiring the deficient element, the redistribution of mobile nutrients from older to younger tissues creating characteristic symptom distributions, the accumulation or depletion of metabolites in pathways affected by the deficiency, and the activation of stress responses that may themselves produce visible symptoms. The element-specific nature of deficiency symptoms reflects the distinct biochemical roles of each nutrient, with nitrogen deficiency affecting protein synthesis and chlorophyll production, phosphorus deficiency affecting energy metabolism and nucleic acid synthesis, potassium deficiency affecting osmotic regulation and enzyme activation, and micronutrient deficiencies affecting the specific enzymes requiring each element as cofactor. The mobility of nutrients within the plant determines symptom distribution, with mobile nutrients (N, P, K, Mg) showing symptoms first on older leaves as elements are remobilized to growing tissues, while immobile nutrients (Ca, Fe, Mn, B, Zn) show symptoms on youngest tissues that cannot acquire adequate supply from older organs.
Nutrient_Deficiency_Indicator_Quality is_a Stress_Response_Quality, serving as parent class for Macronutrient_Deficiency_Symptom_Quality (and eventually Micronutrient_Deficiency_Symptom_Quality) which address specific nutrient classes. This class addresses the overall diagnostic approach to nutritional stress while child hierarchies capture element-specific symptoms. Nutrient mobility determines symptom distribution (mobile → old leaves first immobile → young leaves). Integration of visual diagnosis with tissue and soil analysis provides complete nutritional assessment. Critical for precision nutrient management optimizing production and environmental outcomes.
Elementmangel-Marker
Mineralstoffmangel-Symptome
Nährstoffmangel-Indikatoren
Nährstoffstress-Zeichen
element deficiency markers
mineral deficiency indicators
nutrient stress signs
nutritional deficiency symptoms
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication Robinson JB (1992) Grapevine nutrition. In: Coombe BG, Dry PR (eds) Viticulture, Volume 2. Winetitles, Adelaide, pp. 178-208
Nutrient Deficiency Indicator Quality
In Merlot vines in Saint-Émilion, Bordeaux during October-November, nutrient remobilization exports 3.2 g nitrogen per vine representing 72% of total leaf nitrogen content over 4 weeks, with glutamine and asparagine comprising 65% of exported amino acids, trunk nitrogen content increasing from 0.85% to 1.25% dry weight, and phosphorus remobilization achieving 58% recovery as measured by leaf nutrient analysis before and after senescence under mild oceanic autumn conditions with mean temperature of 14°C.
Nutrient remobilization describes the coordinated export of mineral nutrients and organic compounds from senescing Vitis vinifera leaves to perennial vine tissues during the autumn senescence period, wherein systematic degradation of proteins, nucleic acids, and membrane lipids releases nitrogen, phosphorus, sulfur, and other valuable nutrients that are transported through phloem connections to storage sites in trunk, cordons, and roots where they accumulate as reserves supporting subsequent spring growth and reducing dependence on soil nutrient uptake in the following season. This process represents a critical adaptation for perennial plant nutrient economy, with efficient remobilization enabling recovery of 50-80% of leaf nitrogen representing 2-4 g N per mature vine, 40-60% of leaf phosphorus, 30-50% of potassium, and substantial proportions of sulfur, magnesium, and micronutrients that would otherwise be lost through leaf abscission and decomposition on the vineyard floor. The biochemical mechanisms of nutrient remobilization involve activation of proteases including cysteine proteases and aspartic proteases that degrade stromal and thylakoid proteins releasing amino acids, nucleases that hydrolyze DNA and RNA liberating nucleotides for phosphorus export, and phosphatases that cleave phosphate groups from organic molecules generating transportable inorganic phosphate. The nitrogen remobilization pathway proceeds through proteolytic degradation of Rubisco and other abundant leaf proteins to amino acids, conversion of amino acids to transport forms including glutamine and asparagine through glutamine synthetase and asparagine synthetase activities, and phloem loading of these transport amino acids for export to perennial tissues where storage proteins are synthesized in bark parenchyma cells. The temporal dynamics of nutrient remobilization show characteristic patterns, with nitrogen export initiating early in senescence while photosynthetic capacity remains partially intact to provide carbon skeletons and energy for amino acid synthesis, continuing through mid-senescence as protein degradation accelerates, and declining in late senescence as phloem function deteriorates and export capacity becomes limiting. Environmental factors influence remobilization efficiency, with gradual temperature decline favoring complete remobilization while sudden frost events may terminate senescence prematurely before export completion, and adequate soil moisture maintaining phloem hydration and transport function essential for continued nutrient export. The practical significance of nutrient remobilization for viticulture relates to vine nutrition management, with efficient remobilization reducing fertilizer requirements in subsequent seasons, while premature defoliation through disease, frost, or mechanical damage compromises nutrient recovery potentially affecting vine vigor and productivity in following years.
Nutrient_Remobilization is subclass of Leaf_Senescence representing export of valuable nutrients to storage tissues. Distinguishes from degradation processes by emphasizing transport and conservation. Sibling to Leaf_Senescence_Initiation, Chlorophyll_Degradation, and Progressive_Leaf_Senescence. Recovers 50-80% of leaf nitrogen. Critical for vine nutrient economy. Requires maintained phloem function.
Nährstoff-Export
Nährstoff-Remobilisierung
Nährstoff-Resorption
Seneszenz-Nährstoff-Rückgewinnung
nutrient export
nutrient resorption
nutrient salvage
senescence nutrient recovery
Zapata C et al (2004) Partitioning and mobilization of nitrogen in grapevine. Functional Plant Biology 31: 1141-1152 Masclaux-Daubresse C et al (2010) Nitrogen uptake, assimilation and remobilization in plants. Plant Physiology and Biochemistry 48: 203-212 Keller M (2020) The Science of Grapevines 3rd Ed pp 90-92
Nutrient Remobilization
Efficient cultivars such as Sangiovese remobilize 65-75% of leaf nitrogen to perennial tissues, reducing leaf N from 2.8% to 0.9% dry weight during senescence. Merlot shows moderate remobilization efficiency (55-65% N recovery), while Petit Verdot exhibits lower efficiency (45-55%). Phosphorus remobilization reaches 70-85% in most cultivars, with K remobilization more variable (40-70%) due to its high phloem mobility but competition with fruit demand. Root starch content increases 40-80% between harvest and dormancy in well-managed vines.
A senescence disposition that is the inherited capacity of a Vitis vinifera plant to efficiently catabolize macromolecules in senescing leaves and redistribute the resulting nutrients (particularly nitrogen, phosphorus, and potassium) to perennial storage tissues in roots, trunk, and cordons for use in the following season.
Nutrient remobilization efficiency affects vine sustainability, fertilizer requirements, and spring growth vigor. The process depends on adequate post-harvest leaf area duration, functional phloem connections, and sink strength of perennial tissues. Premature defoliation from disease or frost severely reduces remobilization. Nitrogen remobilization involves protein degradation to amino acids (particularly glutamine and asparagine) transported via phloem.
Nährstoffrückverlagerung
Reservestoffverlagerung
autumn nutrient recovery
herbstliche Nährstoffremobilisierung
nutrient recycling efficiency
reserve translocation capacity
Zapata C et al. (2004) Partitioning and mobilization of starch and N reserves in grapevine. American Journal of Enology and Viticulture 55:83-88 Schreiner RP et al. (2006) N, P, and K supply to Pinot noir grapevines: growth, yield, and fruit composition. American Journal of Enology and Viticulture 57:286-293
Nutrient Remobilization Disposition
Nutrient remobilization efficiency assessment through autumn senescence in Cabernet Sauvignon reveals the nutrient-specific recovery patterns, with nitrogen showing 72% remobilization from senescing leaves (leaf N declining from 2.8% to 0.8% before abscission, recovered N supporting reserve accumulation), phosphorus 65% remobilization, potassium 78% remobilization, but calcium only 8% recovery (largely lost with abscised leaves), demonstrating how efficient remobilization of mobile nutrients reduces dependence on soil uptake while immobile nutrient losses must be replaced through fertilization.
Nutrient remobilization efficiency quality is a nutrient uptake efficiency quality that represents the capacity of grapevines to redistribute nutrients from source tissues (senescing leaves, reserve storage organs) to sink tissues (growing points, developing fruit, overwintering reserves), characterizing the internal nutrient cycling that reduces dependence on continuous soil uptake and enables nutrient economy across seasonal and perennial timescales. Within the BFO framework, this quality addresses the redistribution dimension of plant nutrition, distinguishing internal nutrient cycling from external acquisition while recognizing that efficient remobilization is essential for perennial vine function, enabling nutrients accumulated during favorable periods to support growth, reproduction, and survival when soil uptake is limiting or phenologically inappropriate. The ontological significance lies in nutrient remobilization determining the efficiency of nutrient use across the vine's life cycle, where recovery of nutrients from senescing tissues and mobilization from storage organs enables support of new growth, fruit development, and reserve replenishment without proportional soil uptake, reducing fertilizer requirements and enabling nutrient sufficiency even when soil availability is temporarily limited. The physiological basis for nutrient remobilization involves phloem transport of mobile nutrients (N, P, K, Mg, S) from source to sink tissues, driven by source-sink gradients established through nutrient release from senescing cells (proteolysis, membrane breakdown, chlorophyll degradation releasing bound nutrients) and active loading into phloem for transport to demanding sinks. The nutrient-specific patterns of remobilization show substantial variation, with nitrogen highly mobile (60-80% of leaf N recoverable before abscission), phosphorus moderately mobile (50-70% recoverable), potassium highly mobile (70-90% recoverable), while calcium and manganese are largely immobile (< 10% recoverable), such that remobilization efficiency varies dramatically among nutrients with implications for their management. The seasonal dynamics of remobilization in grapevine show distinct phases including spring mobilization from woody tissue reserves supporting budbreak and early growth, mid-season remobilization from basal leaves to developing fruit, and autumn recovery from senescing leaves to perennial storage tissues preparing for dormancy, with each phase critical for vine nutrient economy. Understanding nutrient remobilization efficiency quality enables assessment of internal nutrient cycling supporting vine function, optimization of fertilization timing relative to remobilization patterns, and appreciation of the nutrient conservation mechanisms that contribute to sustainable vine nutrition.
Nutrient_Remobilization_Efficiency_Quality serves as the parent class for specific remobilization processes (Senescence Recovery, Storage Mobilization). This quality addresses the internal nutrient cycling essential for perennial vine function. The class supports nutrient management optimization through understanding of remobilization patterns affecting fertilization requirements.
Nährstoffremobilisierungseffizienz
Nährstoffumverteilung
Remobilisierungskapazität
internal nutrient cycling
interner Nährstoffkreislauf
nutrient recycling
nutrient redistribution
remobilization capacity
Conradie WJ (1992) Partitioning of nitrogen in grapevines during autumn and the utilization of nitrogen reserves during the following growing season. South African Journal of Enology and Viticulture 13(1): 45-51 Zapata C et al. (2004) Partitioning and mobilization of starch and N reserves in grapevine (Vitis vinifera L.). Journal of Plant Physiology 161(9): 1031-1040
Nutrient Remobilization Efficiency Quality
Nutrient toxicity symptom quality assessment in San Joaquin Valley Thompson Seedless identified multiple ion toxicities affecting vine health where saline-sodic irrigation water combined with boron-enriched groundwater created complex nutritional challenges. Visual nutrient toxicity symptom quality evaluation revealed overlapping symptoms including marginal leaf burn characteristic of both chloride and boron toxicity, interveinal necrosis suggesting boron accumulation, and general leaf bronzing and early senescence indicating sodium stress. Systematic tissue analysis differentiated the contributing nutrient toxicity symptom quality components, with blade Cl at 1.8% (toxic, threshold 0.5%), blade Na at 0.35% (elevated, threshold 0.25%), and blade B at 250 ppm (toxic, threshold 80 ppm), establishing that all three elements exceeded toxicity thresholds and were contributing to the observed damage. The distribution of nutrient toxicity symptom quality within the canopy showed concentration on older leaves where extended transpiration had accumulated ions over time, with marginal symptoms most severe on sun-exposed leaves with highest transpiration rates. Yield impact assessment revealed nutrient toxicity symptom quality causing 25% yield reduction through reduced photosynthetic capacity, early defoliation, and impaired fruit development. The nutrient toxicity symptom quality diagnosis guided integrated remediation including installation of reverse-osmosis water treatment to reduce B, Na, and Cl inputs, replanting with salt-excluder rootstocks (1103P, Ramsey) to limit scion accumulation, and leaching irrigation to move accumulated salts below the root zone, with progressive improvement in vine health over subsequent seasons as tissue ion concentrations declined.
Nutrient toxicity symptom quality is a stress response quality that inheres in a grapevine organism and represents the visual symptoms and physiological damage resulting from excessive accumulation of mineral elements to concentrations that impair cellular function, disrupt metabolism, or cause direct tissue injury, with toxicity symptoms reflecting the specific mechanisms by which each element causes damage at supraoptimal concentrations and the pattern of element accumulation within the plant determining symptom distribution. This quality captures the grapevine response to mineral excess, recognizing that while nutrient deficiency is more common in most viticultural regions, toxicity can occur with elements that accumulate to harmful levels including boron in arid regions with B-rich groundwater, sodium and chloride from saline irrigation water or coastal salt spray, and occasionally other elements under specific soil or management conditions. The ontological significance of nutrient toxicity symptom quality lies in its indication of environmental or management conditions requiring corrective action, where toxicity diagnosis enables identification of the problematic element and guides remediation through irrigation water treatment, rootstock selection for exclusion capacity, or soil amendment to reduce element availability. The biological basis of nutrient toxicity symptoms encompasses direct cellular damage from element accumulation including membrane disruption, enzyme inhibition, and oxidative stress interference with uptake or metabolism of other essential elements creating induced deficiencies and osmotic effects particularly for sodium and chloride that affect water relations and turgor maintenance. The distribution of toxicity symptoms reflects element mobility and accumulation patterns, with mobile elements (B, Cl) transported to leaf margins and tips where transpiration concentrates them, while less mobile elements may accumulate at the point of entry or in storage organs. The interaction between toxicity and other stress factors can complicate diagnosis, with drought stress intensifying salt accumulation, and with multiple element toxicities potentially co-occurring where irrigation water contains several problematic ions.
Nutrient_Toxicity_Symptom_Quality is_a Stress_Response_Quality, serving as parent class for Boron_Toxicity_Quality, Sodium_Toxicity_Quality, and Chloride_Toxicity_Quality which address specific element toxicities. This class addresses the overall concept of mineral excess damage while child classes capture element-specific symptoms. Mobile elements (B, Cl) accumulate at leaf margins/tips via transpiration stream. Toxicity typically shows on older leaves due to accumulation over time. Saline/sodic irrigation and B-rich groundwater are primary causes in viticulture. Rootstock selection for exclusion capacity is key management strategy.
Elementtoxizitäts-Indikatoren
Ionentoxizitäts-Zeichen
Mineralüberschuss-Symptome
Nährstofftoxizitäts-Symptome
element toxicity indicators
ion toxicity signs
mineral excess symptoms
nutrient excess damage
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Walker RR et al. (2010) Rootstock effects on salt tolerance of irrigated field-grown grapevines. Australian Journal of Grape and Wine Research 16:174-180 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication
Nutrient Toxicity Symptom Quality
Nutrient uptake efficiency comparison among rootstocks reveals genotypic variation affecting vine nutrition, with 1103P showing high uptake efficiency for potassium (K uptake rate 45% higher than own-rooted at equivalent soil K) and moderate efficiency for nitrogen, while SO4 shows high nitrogen uptake efficiency but lower potassium acquisition, demonstrating how rootstock selection affects nutrient uptake profiles that influence scion nutritional status and may require adjusted fertilization programs to address rootstock-specific nutrient acquisition characteristics.
Nutrient uptake efficiency quality is a physiological quality that represents the capacity of grapevine root systems to acquire mineral nutrients from the soil solution relative to the resources invested in root construction and maintenance, characterizing the effectiveness of nutrient acquisition that determines vine nutritional status and affects productivity, fruit quality, and the environmental impact of fertilizer inputs. Within the BFO framework, this quality addresses the acquisition dimension of plant mineral nutrition, distinguishing uptake efficiency from subsequent utilization efficiency while recognizing that effective nutrient uptake is prerequisite to all downstream nutritional functions that support vine growth, photosynthesis, and grape quality development. The ontological significance lies in nutrient uptake efficiency determining the conversion of soil and fertilizer nutrients into plant-available forms that support vine function, where high uptake efficiency enables adequate nutrition with lower fertilizer inputs, reducing both production costs and environmental impacts from nutrient leaching, while low efficiency requires greater inputs to achieve target nutritional status. The mechanistic basis for nutrient uptake involves root interception (nutrients encountered through root growth), mass flow (nutrients carried to roots in the transpiration stream), and diffusion (nutrients moving down concentration gradients toward root surfaces), with the relative importance depending on the mobility of specific nutrients and soil conditions. The root properties affecting uptake efficiency include root surface area (total absorbing surface), root distribution (positioning relative to nutrient availability), root hair development (extending the absorbing zone), mycorrhizal associations (fungal networks extending nutrient reach), and transporter expression (proteins mediating nutrient entry into root cells). The nutrient-specific nature of uptake efficiency recognizes that different nutrients present different acquisition challenges, with mobile nutrients (nitrate) primarily acquired through mass flow while immobile nutrients (phosphorus) require root proliferation and mycorrhizal networks, such that efficiency for different nutrients may vary independently. Understanding nutrient uptake efficiency quality enables assessment of root system nutritional function, optimization of fertilizer management for efficient nutrient use, and rootstock selection for improved nutrient acquisition in challenging soil conditions.
Nutrient_Uptake_Efficiency_Quality serves as the parent class for nutrient-specific uptake qualities, beginning with Nitrogen_Uptake_Efficiency. This quality addresses root acquisition function affecting vine nutritional status. The class supports fertilizer management optimization and rootstock selection through uptake efficiency characterization.
Aufnahmekapazität
Nährstoffabsorptionseffizienz
Nährstoffaufnahmeeffizienz
Wurzelnährstoffakquisition
nutrient absorption efficiency
root foraging efficiency
root nutrient acquisition
uptake capacity
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 197-245 Holzapfel BP et al. (2019) Manipulating the postharvest period and its impact on vine productivity of Semillon grapevines. American Journal of Enology and Viticulture 70(2): 217-226
Nutrient Uptake Efficiency Quality
Nutrient use efficiency varies 2-3 fold among scion-rootstock combinations, with efficient combinations producing 150-200 kg fruit per kg N absorbed compared to 80-120 kg fruit/kg N in inefficient combinations. NUE comprises uptake efficiency (nutrient acquired per unit root) and utilization efficiency (biomass produced per unit nutrient in plant). High NUE genotypes maintain yield under reduced fertilizer inputs, with efficient Chardonnay clones producing equivalent yields at 60% of standard nitrogen rates. The NUE concept extends to all nutrients but is most commonly applied to nitrogen due to its dominant influence on vine growth and environmental concerns regarding N losses.
A nutrient acquisition disposition that is the inherited capacity of a Vitis vinifera plant to produce biomass or yield per unit of nutrient absorbed, integrating uptake efficiency with internal utilization efficiency to determine overall productivity relative to nutrient input or availability.
Nutrient_Use_Efficiency_Disposition serves as the parent class for element-specific use efficiency traits. The concept originated in agronomic research addressing fertilizer efficiency and environmental sustainability. In viticulture, high NUE enables reduced input costs, lower environmental impact from nutrient runoff, and potentially improved fruit quality through moderate vigor. Both genetic (rootstock, scion) and management (irrigation, cover crops) factors influence NUE expression.
Nährstoffnutzungseffizienz
Nährstoffverwertung
Ressourceneffizienz
nutrient efficiency trait
nutrient productivity disposition
resource use efficiency
Moll RH et al. (1982) Analysis and interpretation of factors which contribute to efficiency of nitrogen utilization. Agronomy Journal 74:562-564 Keller M et al. (2001) Reproductive growth of grapevines in response to nitrogen supply and rootstock. Australian Journal of Grape and Wine Research 7:12-18
Nutrient Use Efficiency Disposition
The nutrition management description for a Pinot Noir vineyard in the Côte de Beaune integrates four components — annual petiole tissue analysis at flowering and véraison targeting nitrogen at 0.8-1.1% dry weight (sufficient but not excessive to avoid vigor and botrytis problems), phosphorus at 0.15-0.30%, potassium at 1.2-2.0%, and magnesium at 0.25-0.50% soil-applied compost at 4 t/ha biennially providing approximately 40 kg N/ha over the two-year mineralization period targeted foliar magnesium supplementation (MgSO4 at 5 kg/ha in 200 L water, 3 applications from véraison to harvest) to correct the marginal foliar Mg deficiency symptomatic of the calcareous clay soils that competitively inhibit Mg uptake and no mineral nitrogen fertilizer application, with nitrogen supply managed entirely through compost mineralization and the nitrogen-fixing clover component of the inter-row cover crop estimated to contribute 20-30 kg N/ha/year.
A viticultural practice description that documents the strategies, methods, and materials applied to manage grapevine mineral nutrition, encompassing soil-applied fertilization, tissue-based diagnostic monitoring, foliar nutrient supplementation, fertigation delivery, and organic amendment programs that collectively ensure adequate but not excessive supply of macro- and micronutrients to support balanced vine growth, optimal fruit composition, and long-term vineyard sustainability.
Nutrition_Management_Description addresses the domain of viticultural practice where the tension between sufficient nutrition and quality-oriented nutrient restraint is most acute. Unlike annual crop agriculture where maximum yield is typically the nutritional objective, premium viticulture deliberately manages nutrition to achieve moderate rather than maximal vegetative growth — the concept of balanced nutrition in viticulture means supplying nutrients at levels that support adequate canopy development for fruit ripening without stimulating the excessive vigor that produces shading, poor fruit zone microclimate, disease susceptibility, and dilute wine character: this quality-oriented nutritional philosophy means that deficiency correction rather than yield maximization drives most nutritional interventions in premium wine production. The 13 mineral elements essential for grapevine growth (N, P, K, Ca, Mg, S, Fe, Mn, Zn, Cu, B, Mo, Cl) are managed through the complementary approaches captured in the subclasses — soil analysis and amendment, tissue-based diagnostics, soil-applied and foliar-applied supplementation, and organic matter management.
Düngungsprotokoll
Nährstoffmanagement-Beschreibung
Rebenernährungs-Dokumentation
nutrient management documentation
vine nutrition protocol
vineyard fertilization record
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 125-195 Robinson JB (2005) Critical plant tissue values and application of nutritional standards for practical use in vineyards, in Christensen LP and Smart DR (eds) Proceedings of the Soil Environment and Vine Mineral Nutrition Symposium, ASEV pp: 61-68 White RE (2015) Understanding Vineyard Soils 2nd Ed, Oxford University Press pp: 95-135
Nutrition Management Description
Excessive shoot vigor in Merlot diverts carbohydrates from clusters, causing nutritional coulure with fruit set below 15% when shoot growth rates exceed 4 cm/day during flowering. Boron deficiency induces severe nutritional coulure in susceptible cultivars such as Pinot Noir, reducing fruit set from 35% to 8-12% at leaf B concentrations below 25 ppm. Zinc deficiency causes coulure through impaired pollen development, with Chardonnay showing 40% reduction in fruit set at leaf Zn below 20 ppm. Shading of basal leaves during bloom reduces cluster carbohydrate supply, causing 20-35% additional flower abscission.
A coulure susceptibility disposition that is the inherited capacity of a Vitis vinifera plant to exhibit flower or young berry abscission in response to nutritional imbalances or deficiencies, particularly inadequate carbohydrate supply to developing flowers or insufficient micronutrient availability for reproductive processes.
Nutritional coulure results from source-sink competition between vegetative and reproductive growth, or from specific nutrient limitations affecting gametophyte development and function. Unlike weather-induced coulure, nutritional coulure can often be prevented through appropriate canopy management (shoot thinning, hedging) and targeted fertilization. Rootstock vigor significantly influences scion susceptibility to nutritional coulure through effects on vegetative-reproductive balance.
Kohlenhydratmangel-Durchrieselung
carbohydrate-limited coulure
ernährungsbedingte Verrieselung
nutrient-related shatter
nutrition-dependent poor set
nährstoffinduzierter Blütenabwurf
Keller M et al. (2001) Reproductive growth of grapevines in response to nitrogen supply and rootstock. Australian Journal of Grape and Wine Research 7:12-18 Christensen LP (1980) Timing of zinc foliar sprays. II. Effects on size, sweetness, and bunch rot of Thompson Seedless grapes. American Journal of Enology and Viticulture 31:256-260
Nutritional Coulure Disposition
Chardonnay nutritive endosperm during germination (days 3-7 post-stratification) shows progressive lipid mobilization with lipase activity increasing 10-15 fold, protein body breakdown releasing amino acids supporting 15-20% daily increase in embryo axis dry weight, and nutrient translocation sustaining radicle elongation rate of 2-4 mm/day
A nutritive endosperm is endosperm tissue emphasizing its functional role in providing nutritional support to the developing embryo during embryogenesis and to the germinating seedling during early post-germination growth, serving as the primary source of metabolizable nutrients during these critical developmental phases. Nutritive endosperm function operates through two distinct temporal phases= embryo nutrition during seed development (supporting embryogenesis from fertilization through seed maturation, 0-16 weeks post-fertilization corresponding to BBCH 71-89), and seedling nutrition during germination and early establishment (supporting radicle emergence, hypocotyl elongation, and cotyledon expansion until seedling photosynthetic capacity develops, typically 5-12 days post-germination). During embryogenesis, nutritive endosperm provides developing embryo with= soluble sugars transported from maternal tissues and metabolized by endosperm (sucrose, glucose, fructose supporting embryo respiration and biosynthesis), amino acids and nitrogen compounds for embryo protein synthesis, lipid precursors for membrane formation, and growth regulators (cytokinins, gibberellins) that may influence embryo developmental progression. During germination, nutritive endosperm undergoes mobilization= lipid hydrolysis through lipase activity converting stored triglycerides to fatty acids and glycerol, protein hydrolysis through protease activity releasing amino acids, conversion of storage compounds to transportable forms (fatty acids to sugars through gluconeogenesis, proteins to amino acids and amides), and translocation of mobilized nutrients to growing embryo axis through scutellum-like transfer tissues. Nutritive endosperm cellular characteristics include= cells packed with protein bodies (storage vacuoles containing globulins and albumins) and lipid bodies (oil droplets surrounded by phospholipid monolayer), thick cell walls rich in hemicelluloses that are also mobilized during germination, and aleurone-like outer layers with high enzyme activity during germination. Environmental and genetic factors affecting nutritive endosperm function include= maternal nutrition status during seed development (nitrogen, phosphorus availability affecting protein and lipid accumulation), temperature during germination (optimal 20-25°C for enzyme activity and mobilization rates), seed maturity at harvest (immature seeds have incompletely developed endosperm with reduced storage reserves), and cultivar variation in endosperm composition and mobilization efficiency affecting seedling vigor.
Nutritive_Endosperm emphasizes functional support role and is distinguished from Storage_Endosperm (accumulated reserve composition) and Endosperm (complete tissue including development) by focus on active nutritional provisioning during embryogenesis and germination phases. Critical for understanding embryo nutrition during development, germination vigor and seedling establishment success, breeding for seedling vigor traits, and seed abortion mechanisms where endosperm nutritive failure leads to embryo starvation.
Nutrient-providing endosperm
Nährendes Endosperm
Nährstoffbereitstellendes Samengewebe
Supporting endosperm
Versorgungsgewebe
Ebadi, A., Sedgley, M., May, P., & Coombe, B. G. (1996). Seed development and abortion in Vitis vinifera L., cv. Chardonnay. International Journal of Plant Sciences, 157(6), 703-712. | Bewley, J. D., & Black, M. (1994). Seeds= Physiology of Development and Germination (2nd ed.). Plenum Press.
Nutritive Endosperm
Oechsle measurement for Riesling from the Mosel (2024 vintage) records must weights stratified by quality category: Kabinett lot at 76° Oe (predicting 10.0% potential alcohol), Spätlese lot at 85° Oe (11.4% pot. alc.), Auslese lot at 95° Oe (12.8% pot. alc.), and a single Beerenauslese lot at 125° Oe (17.7% pot. alc.) — the precise Oechsle values determining classification eligibility under the German Weingesetz with region-specific minimum thresholds. Oechsle measurement for Müller-Thurgau from Rheinhessen documents harvest at 65° Oe, meeting the minimum must weight for Qualitätswein (QbA) designation in Rheinhessen (60° Oe minimum) but below the Kabinett threshold of 73° Oe, illustrating how Oechsle measurement directly determines the regulatory category and commercial value of the grape lot.
Oechsle measurement is a sugar content data entity that records the sugar concentration in grape must as determined by must density measurement on the Oechsle scale, expressed in degrees Oechsle (°Oe), where the value represents the difference in grams between the mass of one liter of must and one liter of pure water at 20°C (i.e., °Oe = [must density - 1.0000] × 1000), serving as the legal and commercial standard for must weight determination in German, Swiss, and Luxembourgish viticulture and providing the regulatory basis for the German and Swiss quality wine classification systems.
The Oechsle scale occupies a unique position in viticulture as the measurement system most directly and consequentially linked to regulatory wine classification, as the entire German Prädikatswein hierarchy is defined by minimum Oechsle thresholds that vary by region and grape variety. The minimum must weights for each Prädikat level are specified in the Weingesetz and its implementing regulations — for example, Riesling Kabinett in the Mosel requires 73° Oe, while Riesling Kabinett in the Pfalz requires 76° Oe, reflecting the differing climatic conditions and regional traditions. The scale was developed by the German goldsmith and physicist Christian Ferdinand Oechsle (1774-1852) in Pforzheim and has been the legal must weight standard in Germany since the Weingesetz of 1892. The conversion to other scales follows the relationships Brix ≈ Oechsle/4.25 (approximate) and potential alcohol ≈ Oechsle/8 (approximate), though precise conversion requires tabulated values because the relationship between density and sugar content is not perfectly linear across the full concentration range. Measurement is performed using precision hydrometers (Oechslewaagen) or increasingly through digital density meters (oscillating U-tube densitometers) that provide higher accuracy (±0.1° Oe vs. ±0.5° Oe for glass hydrometers). The legal significance of Oechsle measurement demands documented calibration of instruments and temperature correction to the reference temperature of 20°C, as must temperature at harvest commonly ranges from 10-30°C. In borderline cases where must weight falls near a classification threshold, the difference of 1-2° Oe can determine whether a lot qualifies for a higher Prädikat level, with correspondingly significant economic implications.
Mostgewicht in Oechsle
Oechsle degree
Oechsle must density
Oechsle reading
Oechsle value
Oechsle-Grad
Oechsle-Messung
Oechsle-Wert
Oechslegrad-Bestimmung
must weight Oechsle
Koch HJ (2010) Deutsches Weinrecht, Deutscher Fachverlag, pp.: 45-78 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 332-335 Troost G (1988) Technologie des Weines, 6th Ed, Ulmer Verlag, pp.: 78-95
Oechsle Measurement
Oechsle quality achievement in top Mosel Riesling sites reaches extraordinary levels in favorable vintages, with Beerenauslese requiring minimum 110-128° Oechsle (approximately 26-30°Brix) achieved through selective harvest of botrytis-affected berries, and Trockenbeerenauslese demanding 150-154° Oechsle (35-36°Brix) from fully desiccated berries with 90%+ water loss the precision of Oechsle measurement enables classification of individual berry lots into appropriate Prädikat categories, with prestigious estates like Joh. Jos. Prüm and Egon Müller commanding prices exceeding €1000 per bottle for TBA wines from must exceeding 200° Oechsle in exceptional circumstances.
Oechsle_Quality represents the chemical quality characterizing grape must density according to the Oechsle scale, the standard must weight measurement system used in Germany, Switzerland, and Luxembourg. The Oechsle scale, developed by German physicist Ferdinand Oechsle in the early 19th century, expresses must density as the excess mass of one liter of must compared to one liter of water at 20°C, with the value in grams representing degrees Oechsle (°Oe). Thus, must with a density of 1.080 kg/L weighs 80 grams more per liter than water and is expressed as 80° Oechsle. This elegantly simple scale directly relates to the mass of dissolved substances, primarily sugars, making it intuitive for practical use. Typical harvest values range from 60-100° Oechsle for quality wines, with higher Prädikat categories requiring progressively higher minimum values that vary by growing region, reflecting climatic differences in achievable ripeness. The relationship between Oechsle and potential alcohol follows the approximate conversion of °Oe/8 = potential alcohol (% v/v), enabling straightforward harvest planning. German wine law establishes minimum Oechsle values for each quality category and growing region, making Oechsle measurement a legal requirement for quality wine production. Measurement employs calibrated Oechsle hydrometers, precision density meters, or calculation from specific gravity measurements. The Oechsle scale remains deeply embedded in German-speaking wine culture, with vineyard workers, winemakers, and consumers all familiar with Oechsle values as indicators of ripeness and quality potential.
Oechsle_Quality is positioned as a subclass of Must_Weight_Quality, representing the German-language tradition of must weight measurement. The Oechsle scale is mathematically related to other must weight scales: °Oe ≈ (specific gravity - 1) × 1000 ≈ °Brix × 4.25. The scale's integration into German wine law makes it legally significant rather than merely analytically convenient. Temperature correction is essential because density varies with temperature standard measurement specifies 20°C. The historical and cultural significance of Oechsle in German wine regions extends beyond analytical chemistry to encompass quality perception, pricing, and regional identity.
German must weight
Grad Oechsle
Oechsle degrees
Oechsle scale value
Oechslegrad
degrees Oechsle
deutsches Mostgewicht
German Wine Law (Weingesetz) and associated regulations Troost G (1988) Technologie des Weines, 6th Ed, Verlag Eugen Ulmer pp.: 145-168 Würdig G and Woller R (1989) Chemie des Weines, Verlag Eugen Ulmer pp.: 23-45
Oechsle Quality
Off-aroma quality control in commercial winery employs trained panel screening for the major fault categories, with rejection thresholds established at volatile acidity 0.8 g/L (acetic perception threshold), reduction-derived mercaptans above 1.5 μg/L (onion threshold), 4-ethylphenol above 400 μg/L (barnyard perception), and acetaldehyde above 100 mg/L (bruised apple character), with wines exceeding thresholds either treated (if remediable) or declassified to bulk, demonstrating how off-aroma assessment provides the quality control boundary between acceptable and defective wine.
Off aroma quality is an aroma quality that represents olfactory perception of undesirable, faulty, or atypical aromatic elements arising from microbial contamination, chemical degradation, or processing errors, characterizing the presence of aromatic defects that diminish wine quality and may render wine commercially unacceptable depending on severity. Within the BFO framework, this quality addresses the defect dimension of aromatic assessment, distinguishing undesirable aromatic elements from positive character while recognizing that the boundary between fault and acceptable character is often context-dependent, varying with wine style, regional tradition, and individual preference. The ontological significance lies in off-aroma quality defining the negative boundary of wine quality, where the presence and severity of aromatic defects determines whether wine meets minimum quality thresholds for commercial release, with fault detection and remediation representing essential quality control functions in wine production. The compositional basis for off-aromas includes diverse compound classes from different fault origins, with volatile acidity (acetic acid, ethyl acetate) from aerobic microbial activity, sulfur compounds (hydrogen sulfide, mercaptans) from reductive conditions, volatile phenols (4-ethylphenol, 4-ethylguaiacol) from Brettanomyces infection, and oxidation products (acetaldehyde, sotolon) from excessive oxygen exposure, each creating distinctive and recognizable fault characters. The threshold and context dependency of off-aroma assessment creates complexity, as the same compound may be considered a fault at high concentrations while contributing positively to complexity at lower levels, and regional or stylistic traditions may embrace characteristics considered faulty in other contexts. The remediation possibilities vary among fault types, with some correctable through winemaking intervention (copper fining for reduction), others preventable but not reversible (oxidation), and still others representing permanent contamination requiring blending or declassification (severe Brett). Understanding off-aroma quality enables fault identification and diagnosis, quality control threshold establishment, and remediation decision-making for wines showing defect character.
Off_Aroma_Quality serves as the parent class for specific fault categories (Volatile_Acidity, Reduction, Brett, Oxidation), representing the defect dimension of aromatic assessment. This quality requires acknowledgment of threshold and context dependency, as fault perception varies with concentration, wine style, and taster sensitivity. The class supports quality control functions while recognizing the philosophical complexity of defining faults in an aesthetic domain.
Defektaromen
Fehlaromen
Weinfehler
defect aroma
faulty nose
fehlerhafte Note
off-character
wine fault
Goode J (2014) Flawless: Understanding Faults in Wine. University of California Press, Berkeley, pp. 1-25 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed. John Wiley & Sons, Chichester, pp. 231-285
Off Aroma Quality
Pinot Noir clone 115 bears an Official_Clone_Role as a registered French clone (ENTAV-INRA Catalogue des Clones), approved for commercial propagation and required to be identified on all certification labels for derived planting material—enabling growers to specify this clone when ordering vines. Cabernet Sauvignon clone 337 serves in an Official_Clone_Role with French and California recognition, the dual registration enabling international propagation and trade of this widely demanded selection. Riesling clone 239 Gm bears an Official_Clone_Role in the German clone registry, the "Gm" suffix indicating Geisenheim origin and official registration status.
A clone role that inheres in a grapevine clone by virtue of its formal recognition within an official regulatory framework governing grapevine propagation material, distinguishing it from unregistered or informal selections. Official clones have completed evaluation processes required by national or regional certification authorities, meeting criteria for distinctness (demonstrable difference from existing registered clones), agronomic merit (commercially relevant characteristics), and phytosanitary status (freedom from specified pathogens at the required certification level). Official clone status enables legal propagation and sale within the regulatory jurisdiction, with the clone assigned a unique identifier (number, code, or name) that must appear on certification labels for all derived propagation material. Official recognition systems vary internationally—EU member states maintain national clone registries coordinated through EU plant variety frameworks, while non-EU countries have developed analogous systems (FPS in California, CSIRO in Australia, SAG in Chile). Official clone status carries commercial significance, as growers increasingly specify clones when ordering vines and certification requirements restrict propagation of unregistered selections.
Official_Clone_Role is subclassed by registration category (Registered, Certified, Standard/CAC) reflecting the tier within the certification hierarchy at which the clone has been approved. Official status is administratively defined rather than biologically determined—the same genetic material may have official status in one jurisdiction but not another, or may be registered under different designations in different countries. The expansion of official clone registries reflects both scientific understanding of clonal variation and market demand for clone-specific planting material major varieties now have dozens to hundreds of officially registered clones (Pinot Noir exceeds 50 registered French clones alone). Official clone evaluation processes typically require 10-20 years from initial selection through field trials, wine evaluation, and administrative registration, creating substantial lag between identifying promising selections and their commercial availability.
amtlicher Klon
approved clone
listed clone
official clone
registered clone
registrierter Klon
zugelassener Klon
European Union (2020) Commission Implementing Directive 2020/177 ENTAV-INRA (2007) Catalogue des Variétés et Clones de Vigne Cultivés en France Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 Golino DA et al. (2017) Foundation Plant Services, HortScience 52: 1190-1194
Official Clone Role
Blaufränkisch is the official prime name in the Austrian Weingesetz and the Bundesamt für Weinbau variety register, while the genetically identical cultivar carries the official prime name Kékfrankos in the Hungarian National Vine Variety Catalogue (Nemzeti Fajtajegyzék), Lemberger in the German Bundessortenamt register, and Frankovka in the Czech and Slovak national catalogues — each constituting a distinct official prime name with legal force within its respective jurisdiction despite referring to a single genotype confirmed by SSR profiling.
A prime name that is formally designated as the legally recognized cultivar name within a national or supranational variety registration system, carrying regulatory authority for purposes of variety certification, planting authorization, wine labeling, and official documentation within the jurisdiction of the registering authority.
Official_Prime_Name reflects the legal dimension of cultivar nomenclature where the designated name carries regulatory force within a defined jurisdiction. The fragmentation of official names across national registration systems is a persistent challenge for international viticulture: the EU Vine Variety Catalogue (Commission Delegated Regulation 2019/33) attempts to harmonize variety names across member states but permits national synonyms alongside the catalogue entry, resulting in continued nomenclatural plurality. The official prime name determines the name that must appear on wine labels, nursery certificates, and planting authorization documents within the jurisdiction.
Sortenschutzname
amtlicher Sortenname
legally recognized cultivar name
national catalogue name
officially registered name
offiziell registrierter Name
EU Regulation 1308/2013 Art. 81 on vine variety classification Bundessortenamt (2023) Beschreibende Sortenliste Reben OIV (2013) OIV List of Vine Varieties and Their Synonyms
Official Prime Name
The OIV code number 225 identifies the descriptor for berry skin color — one of the most diagnostically important characters for cultivar identification and the descriptor most frequently used in ampelographic keys — scored on a 9-point scale where 1 = green-yellow (e.g., Chardonnay), 2 = rose (e.g., Gewürztraminer), 3 = red (e.g., Cardinal), 4 = grey (e.g., Pinot Gris), 5 = dark red-violet (e.g., Cabernet Franc), 6 = blue-black (e.g., Cabernet Sauvignon). The numerical code system enables database queries across all cultivars sharing a specific descriptor state (e.g., all cultivars with OIV 225 = 6 returns the complete list of blue-black-berried cultivars in the database), multi-character filtering for cultivar identification (e.g., OIV 225 = 6 AND OIV 067 = 3 AND OIV 202 = 3 narrows candidates to small-clustered pentagonal-leaved blue-black cultivars), and automated similarity calculations between cultivar profiles.
An OIV descriptor component that records the unique numerical identifier assigned to a specific morphological character in the OIV descriptor system, with code numbers organized by plant organ and observation timing into sequential blocks — shoot tip and young shoot (001-016), young leaf (051-056), mature leaf (065-094), inflorescence and flower (151-155), cluster (201-209), berry (220-244), seed (501-502), shoot and cane (101-104), and phenological characters (301-306) — enabling unambiguous reference to specific characters across languages and databases.
OIV_Code_Number provides the referencing system that enables the OIV descriptor framework to function as an international data standard. The code number scheme was designed to accommodate future descriptor additions without disrupting existing numbering — gaps between sequential codes within each organ block allow insertion of new descriptors as ampelographic science identifies additional diagnostically useful characters: for example, the mature leaf block (065-094) contains both established descriptors (OIV 067 blade shape, OIV 068 number of lobes, OIV 076 petiolar sinus shape) and more recently added descriptors addressing characters recognized as diagnostically important through statistical analysis of descriptor discriminative power. The code numbers serve as the universal reference language for ampelographic data exchange, replacing the organ-specific and language-dependent descriptor names that varied between national ampelographic traditions (French, German, Italian, Russian systems each having developed partially incompatible terminology before OIV harmonization).
OIV character code
OIV descriptor identifier
OIV numerical code
OIV-Codenummer
OIV-Merkmals-Kennung
OIV-numerischer Code
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition UPOV (2011) Guidelines for the Conduct of Tests for Distinctness, Uniformity and Stability — Vine (Vitis L.), TG/50/9
OIV Code Number
The OIV descriptor record for Cabernet Sauvignon at OIV 067 (mature leaf blade shape) documents the expression state 3 (pentagonal), observed on 10 representative mature leaves from the middle third of normally growing shoots at E-L stage 33-35 (berry touch to véraison) in the third to fifth year of planting on standardized rootstock — this single descriptor observation, combined with the approximately 80 additional OIV descriptors recorded for each cultivar, contributes to the multi-character ampelographic profile that constitutes the morphological identity of the cultivar in the VIVC database, the EU Common Catalogue of Varieties, and national cultivar registration systems.
An ampelographic description component that records the observation of a single morphological character according to the standardized OIV (Organisation Internationale de la Vigne et du Vin) descriptor system, specifying the OIV code number identifying the character, the descriptor state (expression level) observed in the specific cultivar, and the priority level assigned to the descriptor in the OIV hierarchy, providing the atomic unit of standardized morphological data in international grapevine cultivar characterization.
OIV_Descriptor provides the standardized recording format for individual morphological observations that, aggregated across all descriptors, constitute the complete ampelographic description of a cultivar. The OIV descriptor system defines each character by a unique code number (001-502+), a character name in multiple languages, a measurement scale (typically 1-9 for visual scoring, continuous for measured characters), defined reference cultivars exemplifying each scale point, the recommended observation timing (phenological stage, organ age), sample size and selection protocol, and the descriptor's priority level (1 = primary/essential, 2 = secondary/recommended, 3 = tertiary/optional): this comprehensive specification ensures that descriptors are recorded under comparable conditions by different observers in different environments, maximizing the cross-site and cross-year reproducibility that is essential for descriptor data to function as cultivar identification tools. The system's limitations include residual subjectivity in visual scoring (inter-observer variability of 1-2 scale points is common for some descriptors), environmental modification of character expression (genotype × environment interaction affecting descriptor states), and the inability to capture subtle quantitative variation within the discrete scoring categories.
OIV character record
OIV trait observation
OIV-Deskriptor
OIV-Merkmalsbeschreibung
standardisierter morphologischer Deskriptor
standardized morphological descriptor
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition UPOV (2011) Guidelines for the Conduct of Tests for Distinctness, Uniformity and Stability — Vine (Vitis L.), TG/50/9 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC), Vitis 54: 155-157
OIV Descriptor
The OIV descriptor state for Merlot at OIV 076 (mature leaf petiolar sinus shape) is recorded as state 5 (slightly overlapping) — meaning the two lobes bordering the petiolar sinus overlap slightly with a narrow gap visible from above, positioned between the non-overlapping state 3 (open, as in Riesling) and the strongly overlapping state 7 (closed with lobes pressed together, as in Cabernet Sauvignon). The state assignment was made by comparison with the reference cultivars specified in the OIV descriptor list for OIV 076 — state 1 (very wide open, reference Rupestris du Lot), state 3 (open, reference Riesling), state 5 (slightly overlapping, reference Merlot itself as reference cultivar), state 7 (strongly overlapping, reference Cabernet Sauvignon), state 9 (very strongly overlapping, reference Schiava Grossa) — with observations made on 10 fully expanded leaves from the middle third of normally growing shoots.
An OIV descriptor component that records the specific expression level observed for a morphological character in a given cultivar, documented as a numerical value on the descriptor-specific scale (typically 1, 3, 5, 7, 9 for qualitative characters or continuous ranges for quantitative characters), with each scale point defined by a verbal description and illustrated by reference cultivars that serve as calibration standards for consistent scoring across observers, sites, and years.
OIV_Descriptor_State captures the phenotypic observation that constitutes the informational content of each descriptor record. The standardized scoring system uses a 1-9 scale with defined anchor points for each character, but the interpretation and practical application vary by character type: qualitative characters (e.g., leaf blade shape, berry skin color) have discrete states where intermediate values are less meaningful and scoring should match the closest defined category pseudo-qualitative characters (e.g., petiolar sinus shape) have a defined range of morphological expressions that form a continuum but are scored at discrete points and quantitative characters (e.g., berry weight, cluster length) are measured on continuous scales that can be converted to 1-9 categories using defined class boundaries. The reference cultivar system is the critical calibration mechanism — by specifying widely grown, internationally recognized cultivars as exemplars of each descriptor state, the OIV system enables observers who may never have scored a particular character to calibrate their scoring against known examples available in most germplasm collections, substantially reducing inter-observer variability compared to verbal descriptions alone.
Merkmalsausprägungswert
OIV expression level
OIV-Ausprägungsstufe
OIV-Deskriptor-Zustand
character state value
descriptor score
OIV Descriptor State
The OIV priority level assignments for mature leaf descriptors illustrate the hierarchical system — OIV 067 (blade shape) is Priority 1 (essential for identification, scored for every cultivar in the VIVC database), OIV 076 (petiolar sinus shape) is Priority 1 (one of the most diagnostically powerful leaf characters, distinguishing major cultivar groups), OIV 084 (density of prostrate hairs on lower leaf surface) is Priority 2 (recommended for comprehensive characterization, useful for distinguishing morphologically similar cultivars within groups), and OIV 089 (density of erect hairs on main veins of lower leaf surface) is Priority 3 (supplementary detail providing additional discrimination for closely related cultivars). The VIVC database requires complete Priority 1 descriptor data for new cultivar registrations, while Priority 2 and 3 descriptors are populated progressively as resources permit.
An OIV descriptor component that records the hierarchical importance classification assigned to each descriptor in the OIV system — Priority 1 (primary descriptors essential for cultivar identification, mandatory for database registration), Priority 2 (secondary descriptors recommended for comprehensive characterization), or Priority 3 (tertiary descriptors providing supplementary information) — enabling the stratification of ampelographic effort from minimal identification (approximately 14 Priority 1 descriptors) through standard characterization (approximately 40 Priority 1+2 descriptors) to exhaustive documentation (all 130+ descriptors).
OIV_Priority_Level provides the practical framework for allocating the substantial labor investment required for thorough ampelographic characterization — complete scoring of all 130+ OIV descriptors for a single cultivar requires observation across multiple phenological stages (shoot tip at budburst, young leaves during active growth, mature leaves at véraison, flowers at bloom, clusters and berries at harvest, dormant canes in winter), demanding repeated vineyard visits over an entire growing season and typically 2-3 hours of observation per cultivar per visit. The priority stratification addresses the reality that few institutions have the resources to score all descriptors for large collections: Priority 1 descriptors were selected through statistical analysis of discriminative power (the ability to distinguish between cultivars) and practical observability (characters that can be reliably scored by trained observers with minimal equipment), providing a minimal but effective identification dataset — studies by Maul et al. (2012) demonstrated that the approximately 14 Priority 1 descriptors correctly distinguished over 85% of cultivar pairs in the VIVC database, while addition of Priority 2 descriptors increased discrimination to over 95%.
Beschreibungsrang
OIV characterization level
OIV descriptor priority
OIV-Deskriptor-Priorität
OIV-Prioritätsstufe
descriptor importance rank
OIV Priority Level
The OIV registers Cabernet-Sauvignon (with hyphen, OIV variety code 04601) as the international reference name, accompanied by 66 officially recognized synonyms including Bouchet (Saint-Émilion historical usage), Vidure (archaic Graves designation), Petit-Cabernet (Médoc traditional name), and Bordeaux (Bulgarian usage), with the OIV name serving as the default international designation in trade agreements, scientific publications, and cross-border wine labeling when national names differ.
A prime name that is the officially recorded designation for a grapevine cultivar in the International Organisation of Vine and Wine (OIV) International List of Vine Varieties and Their Synonyms, serving as the internationally recognized reference name for cross-jurisdictional communication and harmonization of cultivar nomenclature across OIV member states.
OIV_Registered_Name represents the international nomenclatural standard maintained by the OIV since 1951, with the current List of Vine Varieties and Their Synonyms covering approximately 1,500 cultivars with their accepted synonyms. The OIV list has special regulatory significance under EU wine legislation where OIV-recognized names and synonyms are accepted for international trade labeling: however, the OIV list is less comprehensive than the VIVC database (which documents over 24,000 designations) and is updated less frequently, creating occasional discrepancies between OIV registered names and current VIVC prime names. OIV naming conventions include specific orthographic rules such as the use of hyphens in compound names.
OIV catalogue designation
OIV listed name
OIV variety name
OIV-Listenname
OIV-Sortenbezeichnung
OIV-registrierter Name
OIV (2013) OIV List of Vine Varieties and Their Synonyms, 4th Edition OIV Resolution OIV-VITI 503AB-2012 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157
OIV Registered Name
Cabernet Sauvignon grafted on 3309C rootstock at 42 years post-grafting showing old grafted rootstock trunk 10.8 cm diameter, trunk extending 28 cm from root crown to graft union positioned 22 cm above soil, graft union decades-old completely invisible beneath 5.4 mm thick bark, old trunk bark showing deep longitudinal fissures with very dark grey-brown heavily weathered and eroded surface, extensive lichen covering 40% of bark surface, trunk bark showing some fine cracks and beginning of plate sloughing, diameter growth 0.2 cm current season (reduced from 0.4 cm in younger mature phase), cross-section showing 42 annual rings with recent rings notably narrower, substantial central heartwood formation 5 cm diameter with functional sapwood 2.9 cm thickness in outer trunk, trunk self-supporting and structurally sound, supporting moderate canopy 3.1 m shoot length and 5.2 kg fruit load, vine showing subtle vigor decline but maintaining good fruit quality characteristic of old vine
Old grafted rootstock trunk is grafted rootstock trunk that has transitioned beyond mature phase into old age typically years 30-60+ post-grafting, characterized by decades-old fully stable graft union, very thick extensively weathered bark, large diameter typically 8-15+ cm with declining annual growth rate 0.1-0.3 cm, extensive heartwood formation with reduced functional sapwood, potential subtle vigor decline, and representing late productive life where trunk continues to support production though showing signs of aging and potentially approaching eventual vine replacement or renewal. Old grafted rootstock trunk age and transition includes= spanning years 30-60+ post-grafting typically (beginning of old trunk phase extending through very old vine status), representing post-mature phase (trunk having transitioned beyond optimal mature period into aging stage), showing aging characteristics (various indicators of trunk aging beyond mature status), displaying potential subtle declining function (some reduction in vigor, growth rate, or productive capacity compared to mature prime), and marking late vineyard life (trunk in final decades of productive potential before eventual replacement). Old grafted rootstock trunk ancient graft union includes= showing decades-old consolidated graft union (callus tissue formed 30-60+ years prior completely incorporated into ancient trunk structure), displaying perfect long-established vascular continuity (xylem and phloem connections maintained for decades functioning adequately though perhaps with slight age-related transport reduction), exhibiting completely stable graft union morphology (bulge morphology unchanged for decades with no structural changes for many years), showing entirely obscured graft line (extremely thick old bark and complete decades-old tissue integration making original grafting point completely invisible without bark removal), and demonstrating proven lifelong reliability (graft union having withstood many decades of stress confirming exceptional long-term structural integrity). Old grafted rootstock trunk physical characteristics includes= exhibiting large old diameter (8-15+ cm typical range reflecting decades of annual diameter increase though upper range transitioning into ancient very old trunk), showing very thick extensively developed bark (periderm 4-8 mm thickness with maximum possible expression of rootstock texture after decades of cork accumulation), displaying extremely deep fissure patterns (longitudinal furrows reaching maximum depth with extensive weathering and erosion after decades), revealing very heavily weathered coloration (dark grey to nearly black bark showing extensive environmental exposure, oxidation, and weathering over many decades), potentially showing bark sloughing (outer bark layers beginning to separate and flake in plates or strips in very old trunk), and showing very substantial massive appearance (trunk appearing impressively solid and ancient reflecting many decades of growth). Old grafted rootstock trunk declining growth rate includes= showing reduced annual diameter increase (0.1-0.3 cm typical old trunk growth compared to 0.3-0.6 cm mature trunk reflecting age-related growth decline), displaying slower more variable growth (annual increments less consistent and potentially more variable year to year), exhibiting reduced cambial vigor (vascular cambium activity declining with age producing narrower annual rings), forming many narrow late rings (recent annual rings in cross-section notably narrower than earlier growth rings indicating slowing growth), and approaching growth cessation (very old trunks potentially showing minimal diameter increase in some years suggesting approaching end of active growth phase). Old grafted rootstock trunk heartwood formation includes= showing substantial heartwood development (extensive conversion of inner xylem to non-functional heartwood in trunk center), displaying reduced functional sapwood ring (active water-conducting xylem limited to outer trunk layers typically outer 2-4 cm), exhibiting central heartwood core (inner trunk wood converted to heartwood no longer conducting water though providing structural support), potentially showing heartwood discoloration (central wood darker colored than sapwood due to extractive deposition during heartwood conversion), and maintaining adequate transport despite heartwood (sufficient functional sapwood remaining in outer trunk for continued water and nutrient transport though perhaps with slight reduction from mature trunk capacity). Old grafted rootstock trunk bark characteristics includes= showing maximum bark thickness accumulation (decades of successive periderm formation creating very thick protective bark layer), displaying extremely deep weathered fissures (longitudinal furrows deeply cut with extensive erosion and weathering), exhibiting extensive lichen and moss growth (decades of exposure allowing substantial epiphyte colonization on bark surface), revealing heavily cracked outer bark (bark surface showing extensive fine cracks and potential beginning of plate sloughing), potentially showing bark damage (accumulated damage from decades of environmental exposure, equipment contact, and aging potentially including areas of dead bark or trunk wounds), and demonstrating rootstock cultivar identity (extremely well-developed bark characteristics making rootstock variety readily identifiable). Old grafted rootstock trunk potential vigor decline includes= possibly showing subtle vegetative vigor reduction (shoot growth potentially slightly reduced compared to mature trunk vigor though many old trunks maintaining adequate vigor), potentially exhibiting reduced canopy density (leaf area perhaps slightly reduced compared to mature vine), possibly displaying less vigorous shoot growth (annual shoot length potentially somewhat shorter than younger mature vine), showing potential for reduced reserve accumulation (carbohydrate storage perhaps slightly reduced in aging trunk), and possibly exhibiting longer dormancy (bud break potentially slightly delayed in old vines compared to younger mature vines though pattern varying with cultivar and conditions). Old grafted rootstock trunk continued production capability includes= generally maintaining adequate fruit production (most old grafted trunks continuing to support crop loads though perhaps with slight yield reduction), often producing high-quality fruit (old vines frequently associated with excellent fruit quality potentially due to natural yield limitation and vine balance), showing variable crop levels (year-to-year production potentially more variable in old vine than stable mature vine), potentially exhibiting natural yield reduction (old trunk perhaps supporting slightly lower crop than younger mature vine though balanced with vigor and canopy), and demonstrating old vine character (fruit from old vine trunks sometimes showing distinctive quality characteristics valued in viticulture). Old grafted rootstock trunk structural stability and damage includes= generally maintaining structural integrity (old trunk typically remaining mechanically sound and stable despite age), potentially showing accumulated trunk damage (decades of vineyard operations potentially resulting in accumulated scars, wounds, or damage areas), possibly exhibiting areas of decay (some old trunks showing localized wood decay particularly in damaged areas though often compartmentalized), displaying healed old wounds (evidence of past injuries, cankers, or damage successfully compartmentalized and healed over years), and potentially showing hollow areas (very old trunks occasionally developing small hollow areas in central heartwood though usually maintaining adequate structural capacity). Old grafted rootstock trunk disease and health includes= potentially showing increased disease susceptibility (aging trunk possibly more vulnerable to trunk diseases particularly in stressed or damaged areas), possibly exhibiting old trunk diseases (Eutypa, Botryosphaeria, or other trunk pathogens potentially affecting old vine particularly if management inadequate), showing accumulated disease scars (evidence of past infections successfully managed and compartmentalized), displaying resilience despite age (many old trunks remaining remarkably healthy despite decades of pathogen exposure demonstrating good disease resistance and management), and requiring continued monitoring (old trunk needing regular inspection for disease symptoms particularly in areas of damage or stress). Old grafted rootstock trunk management considerations includes= assessing continued productive value (evaluating whether old trunk capable of supporting adequate production or approaching end of economic life), monitoring trunk health and stability (regular inspection of old trunk for disease, decay, structural soundness, and continued vigor), considering renewal or replacement options (evaluating whether to maintain old vine, implement trunk renewal, or replant vineyard), managing for old vine quality (implementing practices to maintain fruit quality and manage natural vigor and yield decline in old vine), protecting valuable old trunk (providing appropriate care and protection for old vine if maintained for quality or historical reasons), and planning succession (developing timeline for eventual vineyard renewal or trunk replacement if old vine declining). Old grafted rootstock trunk diameter range includes= typical old trunk diameter 8-15 cm (common range for old phase), vigorous rootstocks showing 12-18+ cm (1103P, 110R potentially reaching very large dimensions in old age), moderate vigor rootstocks showing 9-13 cm (SO4, 3309C showing intermediate old trunk size), low vigor rootstocks showing 7-11 cm (Riparia types showing smaller old trunk even after decades), ancient trunks approaching 15-20+ cm (very old trunks in upper range transitioning to ancient status), and site variation (soil, water, and climate affecting old trunk dimensions within genetic potential). Old grafted rootstock trunk cultivar-specific old age includes= SO4 showing 9-13 cm diameter with extremely thick deeply fissured very dark weathered bark, 3309C displaying 9-12 cm diameter with well-developed fissures and grey bark extensively weathered, Teleki 5C exhibiting 10-13 cm diameter with very pronounced bark texture and heavy weathering, 1103P showing 12-18+ cm diameter with exceptionally thick deeply furrowed bark, Riparia Gloire displaying 7-10 cm diameter with moderately thick fissured bark for Riparia, 110 Richter exhibiting 13-17 cm diameter with extremely thick rough heavily weathered bark, and 101-14 showing 8-11 cm diameter with thin but fissured weathered bark. Old grafted rootstock trunk in old vineyard context includes= characterizing old and ancient vineyards (old grafted trunks typical of vineyards planted 30-60+ years prior), representing valuable old vine plantings (old vineyards often prized for distinctive fruit quality and old vine character), contributing to vineyard heritage and history (old grafted trunks representing decades of vineyard history and tradition), demonstrating exceptional long-term performance (decades of production showing successful rootstock-scion pairing and site adaptation), potentially designated as old vine vineyard (vineyards with old grafted trunks qualifying for old vine designation depending on regional standards), and marking significant viticultural achievement (maintaining healthy productive old grafted trunk through decades demonstrating exceptional viticulture and vine longevity). Old grafted rootstock trunk replacement considerations includes= weighing continued production capacity (evaluating whether old trunk still supporting adequate yields or declining significantly), assessing fruit quality benefits (considering whether old vine fruit quality justifies maintaining aging vine), considering trunk health and stability (determining whether old trunk remaining sound or showing problematic decay or disease), evaluating economic factors (comparing costs of maintaining old vine versus replacing with new planting), factoring vineyard heritage and reputation (considering value of old vine status for vineyard identity and marketing), and planning succession timing (if replacement decided, determining optimal timing for vineyard renewal). Old grafted rootstock trunk renewal options includes= considering trunk renewal techniques (implementing cordon renewal, head renewal, or other trunk regeneration approaches potentially extending old vine life), evaluating young trunk development (training new trunk from basal shoot potentially replacing old failing trunk while maintaining root system), assessing grafting options (potentially field-grafting new scion onto old rootstock if rootstock healthy but trunk damaged), maintaining root system (old established root system valuable potentially retained while renewing trunk or grafting new scion), and comparing renewal versus replacement (determining whether trunk renewal economically and practically viable or complete vine replacement preferable). Old grafted rootstock trunk as viticultural heritage includes= representing living history (old grafted trunk surviving from previous generation of viticulturists connecting present to past), demonstrating rootstock-scion longevity (decades of successful integration showing potential lifespan of grafted grapevine), embodying traditional viticulture (old trunk potentially reflecting historical training systems, rootstock choices, and practices), contributing to old vine mystique (old trunk age contributing to mystique and perceived quality of old vine fruit), and marking exceptional vineyard stewardship (maintaining healthy productive old trunk for decades demonstrating commitment to quality and long-term vineyard management). Old grafted rootstock trunk significance includes= representing successful long-term grafted vine (decades of production demonstrating viability and longevity of rootstock-scion partnership), showing transition into old age (old trunk marking shift from prime mature phase into aging stage with associated changes), demonstrating continued productivity in age (old trunk often remaining remarkably productive despite aging indicators), requiring management decisions (old trunk necessitating assessment of continued value versus replacement or renewal), embodying old vine character (old trunk contributing to distinctive quality potentially associated with old vines), and marking late productive life (old trunk representing final phase of productive existence before eventual vine replacement or death).
Old_Grafted_Rootstock_Trunk emphasizes post-mature aging phase (years 30-60+) and is distinguished from Mature_Grafted_Rootstock_Trunk (prime productive phase) by focus on aging characteristics including reduced growth, heartwood formation, potential vigor decline, and late productive life. Critical for understanding aging process, continued production capability, old vine quality potential, management decisions regarding renewal or replacement, and the late-life phase of grafted vine representing decades of successful production before eventual replacement.
Aged grafted rootstock trunk
Alter verpfropfter Unterlagenstamm
Senior grafted rootstock trunk
Veteran grafted rootstock trunk
Veteranen-Wurzelstockstamm
Überalterter Pfropfstamm
Old Grafted Rootstock Trunk
Carignan old grafted scion trunk on 99R drought-tolerant rootstock in Languedoc at 42 years post-planting shows 155-195 mm scion diameter with extensive heartwood (110-145 mm, 70-75% of total), reduced scion sapwood (18-25 mm radial depth), graft union at 10 cm above soil with 118-128% swelling (increased from 112% in prime maturity suggesting age-related changes), rootstock diameter 165-210 mm, 85-125 g combined depleted reserves (scion 30-45 g, rootstock 55-80 g), declining productivity 3.2-5.2 kg per vine (55-70% of former peak), but rootstock drought tolerance enabling survival and production in water-limited conditions where own-rooted vines would fail
An old grafted scion trunk is a grafted scion trunk in advanced age (typically 25-50+ years post-planting) characterized by substantial scion diameter, extensive scion heartwood, declining vigor, potential graft union deterioration, and often reduced productivity, representing the senescence phase where vine decline becomes evident though rootstock benefits may continue supporting functionality longer than equivalent own-rooted vines in challenging conditions. Old grafted scion trunk anatomical characteristics include= large scion diameter (varying by rootstock= vigorous rootstocks 110-220 mm at 30-50 years, moderate 95-180 mm, low-vigor 85-160 mm), extensive scion heartwood (60-80% of scion cross-section non-functional for transport), reduced scion sapwood (15-25 mm radial depth compared to 20-35 mm in younger mature trunks), irregular scion annual rings (0.5-3 mm width, variable or missing in stress years), graft union potentially showing age-related changes (union swelling may increase >120% in some aging combinations, callus tissue aging), and thick scion bark (6-12 mm compared to 4-8 mm in younger trunks). Old grafted scion trunk graft union aging includes= union diameter potentially increasing with age (swelling progressing to 115-130% in some old combinations compared to 108-120% in prime maturity), union callus tissue showing age-related changes (lignified callus potentially developing cracks or separations), union vascular continuity potentially declining (hydraulic conductance across union may decrease 10-30% in old vines compared to prime maturity), union as potential disease entry point (old unions showing higher trunk disease incidence), and union mechanical strength potentially declining (increased breakage risk in very old unions particularly with incompatible combinations). Old grafted scion trunk physiological decline includes= reduced water transport despite rootstock (declining hydraulic conductance from reduced sapwood area, xylem embolism, union bottleneck potentially worsening with age), decreased combined carbohydrate storage (scion and rootstock reserves declining from 120-190 g in prime to 70-140 g in old trunks), declining photosynthetic capacity (reduced leaf area, lower net photosynthesis rates despite rootstock vigor influence), extended phenology (budbreak, véraison delayed compared to younger vines even with early rootstocks), and reduced stress resilience (though rootstock benefits may provide old grafted vines better stress tolerance than own-rooted equivalents). Old grafted scion trunk productivity characteristics show= declining yields (typically 2.5-6 kg per vine depending on rootstock, 40-70% of prime maturity production), reduced fruitfulness (40-75% depending on rootstock class, vigorous rootstocks maintaining higher fruitfulness longer), smaller clusters and berries (cluster weight 90-170 g varying by rootstock), but potentially maintained or enhanced quality (smaller berries with high skin/pulp ratio, concentrated flavors, rootstock quality effects still expressed), and greater year-to-year variation (vintage effects more pronounced than in prime maturity). Old grafted scion trunk rootstock influence in senescence includes= vigorous rootstocks potentially extending productive life (maintaining vigor and yield 5-10 years longer than low-vigor rootstocks or own-rooted equivalents), drought-tolerant rootstocks supporting old vine survival (providing water stress resistance critical for declining vine health), lime-tolerant rootstocks preventing chlorosis (continuing to protect against iron deficiency even in senescent vines), and phylloxera-resistant rootstocks enabling longevity (old grafted vines surviving in phylloxera-infested soils where own-rooted equivalents long dead). Old grafted scion trunk disease and deterioration includes= high trunk disease incidence (Eutypa, Esca, Botryosphaeria affecting 40-90% of old grafted vines depending on region), graft union as disease entry site (unions showing higher pathogen colonization than equivalent trunk positions), wood decay organisms in scion heartwood (white and brown rot fungi extensive), rootstock potentially showing disease or decline (old rootstock trunks also subject to disease, root decline), and structural instability (risk of trunk or union failure from decay, disease, or mechanical weakness). Old grafted scion trunk viticulture significance includes= old grafted vine quality potential (concentrated fruit with rootstock modulation potentially producing distinctive wines), rootstock enabling survival (old grafted vines persisting in conditions impossible for own-rooted= phylloxera-infested soils, challenging pH or salinity), genetic and clonal preservation (old grafted vines representing field selections, heritage clones on proven rootstocks), economic challenges (declining productivity with continued costs potentially exceeding returns), and replanting considerations (evaluating rootstock performance, compatibility, soil adaptation informing replanting rootstock selection). Old grafted scion trunk variation by rootstock includes= vigorous drought-tolerant rootstocks (SO4, 140Ru, 1103P) supporting productive senescence 35-55 years, moderate rootstocks showing typical decline 28-45 years, low-vigor rootstocks with earlier senescence 22-38 years, and rootstock-soil matching critically affecting longevity (well-adapted combinations extending productive life, poorly matched combinations accelerating decline).
Old_Grafted_Scion_Trunk emphasizes senescence with rootstock and is distinguished from Mature_Grafted_Scion_Trunk (prime productivity) and own-rooted equivalents (different longevity patterns) by evident decline with rootstock potentially extending productive life or survival compared to own-rooted alternatives. Critical for understanding rootstock effects on vine longevity, old grafted vine quality potential, rootstock enabling survival in challenging conditions, and economic decisions on aging grafted vineyard retention versus replanting.
Alter gepfropfter Edelstamm
Alternde Aufpropfung
Declining grafted scion
Senescent grafted trunk
Seneszenter Veredelungsstamm
Old Grafted Scion Trunk
Carignan old own-rooted trunk in Languedoc at 45 years post-planting shows 145-185 mm diameter with extensive heartwood (105-135 mm diameter, 65-75% of total), reduced sapwood (18-24 mm radial depth), irregular annual rings (recent 5 years averaging 1.2-2.8 mm with missing rings in drought years), 70-95 g depleted starch reserves, declining productivity (2.8-4.2 kg per vine, 50-60% of former peak yield), but produces concentrated fruit with exceptional quality characteristics
An old own-rooted trunk is an own-rooted trunk in advanced age (typically 25-50+ years post-planting) characterized by substantial trunk diameter, extensive heartwood formation, declining vigor, potential trunk disease presence, and often reduced productivity, representing the senescence phase where vine decline becomes evident though fruit quality may remain high or even improve. Old own-rooted trunk anatomical characteristics include= large trunk diameter (100-200+ mm for typical wine cultivars at 30-50 years, compared to 60-100 mm at year 15), extensive heartwood occupying 60-80% of trunk cross-section (central trunk largely non-functional for water transport), reduced sapwood cross-section (15-25 mm radial depth, compared to 20-35 mm in younger mature trunks), irregular annual ring formation (ring width becoming variable 0.5-3 mm, sometimes missing rings in stressed years), thickened bark (6-12 mm compared to 4-8 mm in younger trunks), and potential trunk hollowing (central pith and early heartwood decaying creating hollow center in some old trunks). Old own-rooted trunk physiological decline characteristics include= reduced water transport capacity (declining hydraulic conductance due to reduced sapwood area, xylem embolism, vessel blockage), decreased carbohydrate storage (starch reserves declining from 120-180 g in prime maturity to 60-120 g in old trunks), declining photosynthetic capacity (reduced leaf area, lower net photosynthesis rates), extended phenology (delayed budbreak, flowering, véraison compared to younger vines), and reduced stress resilience (greater susceptibility to drought, heat, cold injury). Old own-rooted trunk productivity characteristics show= declining yields (typically 2-5 kg per vine compared to 4-8 kg in mature prime, reduction of 30-50%), reduced cluster number per shoot (decreasing bud fruitfulness 40-70% compared to 70-90% in prime vines), smaller clusters and berries (cluster weight 80-150 g compared to 120-200 g in vigorous mature vines), but potentially enhanced fruit quality (smaller berries with higher skin/pulp ratio, concentrated flavors, improved phenolic development). Old own-rooted trunk disease susceptibility includes= high incidence of trunk diseases (Eutypa dieback, Esca, Botryosphaeria canker affecting 30-80% of old vines in some regions), wood decay organisms colonizing heartwood (white rot and brown rot fungi utilizing non-functional heartwood), potential phylloxera damage in infested regions (old own-rooted vines particularly vulnerable to root feeding damage), and canopy diseases (declining vigor reducing disease resistance). Old own-rooted trunk management challenges include= difficulty maintaining productivity (reduced vigor limiting crop potential), pruning complexity (working around diseased or dead wood, selecting limited fruitful shoots), disease management limitations (trunk diseases difficult or impossible to cure once established), irrigation requirements (reduced water transport capacity increasing drought sensitivity despite deep roots), and replanting decisions (balancing old vine quality potential against declining productivity and increasing costs). Old own-rooted trunk viticulture significance includes= "old vine" wine quality potential (concentrated flavors, complex character, potential premium pricing), historical vineyard value (heritage sites with cultural or historical significance), genetic preservation (old vines potentially representing rare selections or heritage clones), terroir expression (decades of adaptation to site producing distinctive character), and viticulture challenges (high management input required for diminishing returns). Old own-rooted trunk variation by cultivar and site includes= some cultivars showing excellent longevity (Grenache, Mourvèdre, Zinfandel maintaining productivity 40-80+ years), other cultivars declining earlier (Pinot Noir, Chardonnay often showing senescence 25-40 years), optimal sites supporting extended lifespan (deep soils, moderate climates enabling 50-100+ year productivity), and stressed sites accelerating decline (shallow soils, extreme climates reducing productive lifespan to 20-40 years).
Old_Own_Rooted_Trunk emphasizes senescence phase and is distinguished from Mature_Own_Rooted_Trunk (prime productive phase) and Ancient_Old_Own_Rooted_Trunk (extreme longevity 50-100+ years) by evident decline in vigor and productivity. Critical for understanding old vine quality potential, management challenges of declining vines, economic decisions on vineyard retention versus replanting, and preservation of heritage vineyards.
Alter wurzelechter Stamm
Alternder Rebstamm
Declining trunk
Senescent trunk
Seneszenter Eigenwurzelstamm
Old Own Rooted Trunk
The Zinfandel selections from 1880s plantings in Lodi, Amador, and Sonoma bear Old_Vine_Clone_Roles, material maintained in aged vineyards representing 19th-century California selections distinct from modern registered clones—their survival for 130+ years demonstrating exceptional site adaptation and vine longevity. The Grenache from 80+ year-old Barossa Valley plantings serves in an Old_Vine_Clone_Role, material from early 20th-century Australian plantings preserving selections no longer available from nurseries. Old Mourvèdre selections from Bandol vineyards exceeding 60 years bear Old_Vine_Clone_Roles, traditional selections maintained in production demonstrating the variety's potential for extended productivity when appropriately sited and managed.
A heritage clone role that inheres in a grapevine clone maintained in aged plantings (typically 50+ years, often 80-100+ years) whose continued productivity provides both access to potentially unique genetic material and demonstration of long-term vine performance under specific site conditions. Old vine clones represent material proven through decades of cultivation in specific locations, their survival and continued productivity demonstrating adaptation to site conditions and management systems. The extended timeframe of old vine cultivation predates most formal clone registration, meaning old vine material often represents informal selections, mass selection populations, or early registered clones now rare in commercial nurseries. Old vine clones offer access to diversity potentially absent from contemporary nursery catalogs—genotypes that fell from commercial favor, pre-dating the clone selections that came to dominate specific varieties, or representing regional selections never formally registered. Conservation efforts increasingly focus on old vine material, collecting and preserving selections from aged plantings before vineyard removal eliminates irreplaceable genotypes.
Old_Vine_Clone_Role captures the value of material proven through extended cultivation, where vine longevity itself provides information about adaptation and performance. The designation emphasizes survival and continued productivity rather than historical documentation old vines demonstrate through decades of cultivation what trial data might require years to establish. Old vine material often represents population mixtures rather than uniform clones, with within-vineyard diversity accumulated through original planting or subsequent interplanting. Conservation programs recognize old vine material as a genetic resource requiring protection vineyard removal for replanting or development permanently eliminates genotypes that may be locally unique. The "old vine" designation also carries commercial significance "vieilles vignes," "old vines," and equivalent terms communicate quality associations that some jurisdictions regulate while others leave to producer discretion.
Altrebenklon
Jahrhundertrebenklon
Vieilles-Vignes-Klon
aged vine selection
alte Rebselektion
century vine clone
old vine clone
vieilles vignes clone
Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340 McGourty G, Christensen LP (2013) Heritage clone vineyard development, Practical Winery & Vineyard Journal 34(1): 46-52 This P et al. (2006) Historical origins of wine grapes, Trends in Genetics 22: 511-519 Historic Vineyard Society (2020) California Old Vine Registry
Old Vine Clone Role
A 50-year-old Zinfandel vine in Lodi's sandy soils bears an Old_Vine_Role, its gnarled trunk and limited yields (2 tonnes/hectare versus 8+ for young vines) producing intensely concentrated fruit commanding "Old Vine Zinfandel" premium pricing. A 70-year-old Grenache vine in Châteauneuf-du-Pape serves in an Old_Vine_Role, the deep-rooted survivor of multiple drought years producing fruit complexity attributed to its age, galets-adapted root system, and decades of site integration. A 45-year-old Riesling vine in Wachau terracing bears an Old_Vine_Role, with the "Alte Reben" designation on labels signaling the concentration and complexity valued by regional collectors.
A vine age role that inheres in a grapevine of advanced age (typically 35-50+ years), characterized by declining yields, deep-rooted physiological resilience, and fruit concentration often valued for premium wine production despite—or because of—the reduced productivity. Old vines develop through decades of site adaptation, with extensive root systems penetrating deep soil horizons, gnarled trunk structures accumulating years of pruning history, and physiological balance shifting toward reproductive investment (concentrated fruit) over vegetative growth. The "old vine" designation carries marketing significance in many wine regions, with consumers and critics associating vine age with quality, authenticity, and terroir expression, though the relationship between chronological age and wine quality remains complex and context-dependent. Old vine physiology differs from mature vines: reduced vegetative vigor, smaller canopy size, naturally limited yields, and often greater drought tolerance from deep root systems. Old vine survival requires accumulated favorable conditions—appropriate variety-site match, freedom from major disease or replanting pressure, economic viability of continued production—making old vine vineyards relatively rare and often historically significant.
Old_Vine_Role is subclassed by specific age-significance categories—Vieilles_Vignes (the French marketing term) and Centenarian_Vine (100+ years)—recognizing both the functional concept and its regional/marketing expressions. The definition of "old" varies dramatically with region and variety: 35 years may qualify as old in regions with frequent replanting pressure, while 50+ years is expected for the designation in traditional areas. Old vine marketing has proliferated, with varying regulation (Australia's Barossa Old Vine Charter specifies minimum ages most regions have no formal definition), creating both genuine quality associations and marketing inflation. The physiology underlying old vine quality—reduced yields, deep roots, stress tolerance—can be partially replicated through management (deficit irrigation, reduced crop load) on younger vines, raising questions about whether age itself or the conditions it creates drive quality differences.
Altrebe
Veteranenrebe
aged vine
alte Rebe
ancient vine
betagte Rebe
old vine
veteran vine
Vaudour E et al. (2015) Vine age effect on terroir expression, OENO One Historic Vineyard Society (2020) Old Vine Registry Bonada M et al. (2020) Vine age and grape composition, Australian Journal of Grape and Wine Research 26: 139-152 Keller M (2015) The Science of Grapevines, Academic Press
Old Vine Role
An old wood latent bud on a 7-year-old cordon of Vitis vinifera cv. Merlot, located 60cm from the trunk on the main cordon structure, remaining suppressed for approximately 5 seasons based on bark development, becoming activated following severe hail damage that destroyed 90% of current season fruiting wood and damaged multiple spurs, producing a vigorous shoot 180cm in length, subsequently pruned to establish three new fruiting spurs that restored partial production to the damaged cordon section within two seasons, avoiding the need for complete cordon replacement from trunk-level renewal.
A latent bud located on multi-year-old wood structures including arms, cordons, or perennial framework branches, having remained suppressed through multiple growing seasons or years, characterized by embedment within thickened bark or corky periderm, small size relative to annual bud positions, variable viability depending on age and environmental exposure, and capacity for activation following severe pruning, disease damage to distal structures, or environmental stress releasing apical dominance. Old wood latent buds provide regenerative capacity for perennial vine architecture, enabling replacement of diseased or damaged arms and cordons without trunk-level renewal, representing intermediate-term reserve meristems with activation potential between annual renewal buds and deep trunk latent buds, supporting vineyard recovery strategies that balance production continuity with structural renovation requirements.
Old wood latent buds are common on cordon-trained systems where perennial arms accumulate over multiple years, providing distributed regenerative capacity along the permanent vine structure. Their viability and responsiveness decline with age, with buds on 2-3 year wood showing higher activation rates than those on 10+ year wood. Activation often follows mechanical damage, severe pruning, or pathogen damage that reduces distal canopy and disrupts apical dominance hormone flow. While less vigorous than trunk latent buds due to more tortuous vascular pathways through older xylem with accumulated dysfunction, old wood latent buds offer faster return to production than trunk renewal and maintain more of the established vine architecture.
altes Holz Knospe
mehrjährige Adventivknospe
old wood epicormic bud
perennial wood latent bud
Based on Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Smart et al. (1990) "Principles of grapevine canopy microclimate manipulation with implications for yield and quality= A review" American Journal of Enology and Viticulture 41=230-239 Mundy (2008) "A review of grapevine leafroll associated virus type 3 (GLRaV-3) for the Australian wine industry" Wine & Viticulture Journal 23=8-14.
Old Wood Latent Bud
Systematic characterization of Older_Tissue_Accumulation_Disposition at Bordeaux Sciences Agro tracked ion distribution across leaf positions in 15 rootstock-scion combinations during 6 weeks of salt stress (ECw 4 dS/m). Leaves were harvested at weekly intervals from positions 2-4 (oldest), 6-8 (mid), and 10-12 (youngest) on marked shoots, with chloride and sodium analyzed for each position and timepoint. Older_Tissue_Accumulation_Disposition varied among rootstocks independent of total exclusion capacity. '1103 Paulsen' (moderate excluder) showed strong positional gradient with week-6 chloride concentrations of 185 mmol/kg in oldest leaves, 78 mmol/kg in mid leaves, and 32 mmol/kg in youngest leaves (5.8-fold old/young ratio). '140 Ruggeri' (strong excluder) maintained low concentrations throughout but still showed 3.2-fold positional gradient (28, 15, 9 mmol/kg). 'Schwarzmann' (weak excluder) accumulated high concentrations with minimal gradient (410, 385, 320 mmol/kg, only 1.3-fold ratio), indicating poor Older_Tissue_Accumulation_Disposition. Remarkably, '1103 Paulsen' maintained youngest leaf chloride below toxic threshold (50 mmol/kg) for the entire 6-week period despite oldest leaves exceeding 150 mmol/kg by week 3, demonstrating effective protection of critical tissues. Leaf abscission analysis revealed that '1103 Paulsen' lost 25% of oldest leaves by week 6 with no loss of younger leaves, representing successful operation of the sacrificial strategy. The weak Older_Tissue_Accumulation_Disposition of 'Schwarzmann' resulted in toxic concentrations reaching youngest leaves by week 4, causing shoot tip necrosis and growth cessation despite similar total ion uptake to '1103 Paulsen'.
An Older_Tissue_Accumulation_Disposition is an Ion_Compartmentalization_Disposition that inheres in a Vitis plant and is realized through whole-plant transport mechanisms that preferentially direct sodium and chloride ions to older, senescing leaves while protecting young, metabolically active leaves from toxic ion accumulation. This disposition represents whole-plant level compartmentalization, sacrificing expendable older tissues to maintain the function of irreplaceable younger tissues critical for current photosynthesis and future growth. Older leaves are more tolerant of ion accumulation because they have already completed their major contribution to plant carbon gain and can be abscised if damage becomes severe, while young expanding leaves and the shoot apex must be protected to maintain vine productivity. The mechanism involves differential xylem unloading (older leaves receiving proportionally more of the transpiration stream ions), possible phloem redistribution (retranslocation from young to old tissues), and potentially lower exclusion capacity in older leaf traces. Effective Older_Tissue_Accumulation_Disposition creates steep ion concentration gradients along the shoot, with basal (older) leaves showing concentrations 3-10 fold higher than apical (younger) leaves. This strategy extends the period before toxic concentrations accumulate in critical young tissues, effectively buying time during salt exposure and enabling continued growth even when some older leaves are lost to salt injury.
Older_Tissue_Accumulation_Disposition is_a Ion_Compartmentalization_Disposition. Older tissue accumulation is a whole-plant compartmentalization strategy protecting critical young tissues. Quantified by the old leaf/young leaf concentration ratio (effective values > 3). Mechanism involves differential xylem unloading and possibly phloem retranslocation. Older leaves can be sacrificed (abscised) when ion load becomes excessive. This strategy extends functional duration under salt stress even when total exclusion is limited. The disposition is partially independent of exclusion capacity, representing a distinct tolerance component.
Altgewebe-Akkumulations-Disposition
Basipetale-Ionen-Akkumulation
Opferblatt-Strategie
basipetal ion accumulation
old leaf ion dumping
sacrificial leaf accumulation
Munns R (2002) Comparative physiology of salt and water stress, Plant Cell and Environment 25:239-250 Tester M, Davenport R (2003) Na⁺ tolerance and Na⁺ transport in higher plants, Annals of Botany 91:503-527 Wolf O et al. (1991) Growth and cation composition of leaves and stems of Vitis vinifera in a saline environment, Journal of Plant Physiology 137:656-662
Older Tissue Accumulation Disposition
An omega graft of Pinot Noir clone 115 scion on 3309 Couderc rootstock bears an Omega_Graft_Role, produced on a Grafting Systems™ omega machine at 1,200 grafts per hour, with the interlocking omega cuts creating immediate mechanical stability before wax coating and callusing. An omega graft of Sauvignon Blanc on SO4 serves in an Omega_Graft_Role in New Zealand nursery production, where the technique's efficiency supports the country's substantial planting expansion. An omega graft of Tempranillo clone RJ-43 on 110 Richter bears an Omega_Graft_Role in large-scale Spanish production, with automated omega grafting machines processing 40,000+ grafts daily during the peak grafting season (January-March).
A propagation material role that inheres in a grafted combination created using the omega grafting technique, where interlocking omega-shaped (Ω) cuts on scion and rootstock create a mechanically self-securing joint with maximum cambial contact area. Omega grafting is the dominant machine grafting technique for commercial bench graft production, utilizing specialized grafting machines that simultaneously cut matching omega profiles on scion base and rootstock apex, enabling rapid assembly without the hand skill required for traditional whip-and-tongue grafts. The omega cut geometry provides several advantages—self-locking mechanical engagement (the graft holds together without tying or pinning), maximum cambial alignment around the entire cut circumference, consistent cut geometry across thousands of grafts, and compatibility with high-speed production (experienced operators achieving 1,000-1,500 grafts per hour manually, automated systems exceeding 3,000-5,000 per hour). Omega graft success depends on diameter matching between scion and rootstock (typically within 1-2 mm), sharp blade maintenance ensuring clean cuts, and proper callusing conditions enabling union formation across the contact surfaces.
Omega_Graft_Role captures the dominant commercial grafting technique that has largely replaced traditional hand grafting methods in large-scale nursery production. The omega cut's mechanical self-locking eliminates the need for tying, waxing for mechanical support (though wax is still applied for moisture retention), or staking during callusing, substantially reducing labor requirements compared to traditional techniques. Machine omega grafting has standardized graft production, with specialized equipment manufacturers (Grafting Systems, Single, Stoiber) producing machines used globally. The technique's success depends critically on blade sharpness (dull blades crushing rather than cutting tissue, compromising callus formation) and diameter matching (mismatched diameters creating gaps in cambial contact). Quality control in omega graft production focuses on cut quality inspection, diameter sorting of both scion and rootstock material, and callusing environment optimization to achieve the 85-95% success rates expected in commercial production.
Maschinenveredelung
Omega-Veredelung
machine graft
mechanical graft
mechanische Pfropfung
omega cut graft
omega graft
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 452-456 Gramaje D, Armengol J (2011) Fungal trunk pathogens, Phytopathologia Mediterranea 50: S217-S226 Waite H et al. (2015) Hot water treatment and grapevine propagation, Phytopathologia Mediterranea 54: 325-339
Omega Graft Role
A one-year cane on Vitis vinifera cv. Chardonnay selected during December pruning, having grown from budbreak in April through growth cessation in September and lignification completion in November, measuring 145cm length with 14 nodes, 9.5mm average diameter, showing uniform light brown bark color with no green remaining, moderate flexibility allowing bending to horizontal trellis wire position without cracking, this cane retaining 10 nodes as fruiting wood after tip removal and expected to produce well-balanced crop of 1.8-2.2kg based on cultivar fruitfulness patterns and target leaf area-to-fruit ratios.
A cane that has completed exactly one growing season of development, representing current year's mature wood following cessation of active growth and progression through lignification, characterized by relatively uniform brown coloration indicating completed secondary growth, moderate flexibility compared to multi-year wood, retention of all original nodes from the growing season, and suitability for use as fruiting or renewal wood in the following season. One-year canes represent the optimal age category for productive fruiting wood in viticulture, with buds formed during the previous season at ideal developmental stage for maximum fruitfulness, tissues young enough for vigorous shoot growth but mature enough for structural support, and vascular connections recent enough for efficient water and nutrient transport to developing shoots and clusters.
One-year canes are universally preferred over older wood for fruiting positions due to superior bud fruitfulness (inflorescence primordia formed most recently under current management), predictable shoot vigor (neither excessively vigorous like young renewal wood nor declining like old wood), and reliable vascular function. The transition from shoot to cane occurs gradually during late season as green tissues lignify, with complete cane status achieved when all internode regions show brown woody appearance and growth has fully ceased. Distinguishing one-year from older canes is critical during pruning, accomplished by examining node scars, bark texture, and position relative to permanent structure, as retention of two-year or older wood as fruiting positions typically results in reduced yields and declining fruit quality.
annual cane
current season cane
diesjährige Rute
einjährige Rute
first-year wood
Based on Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 89-112 Howell (2001) "Sustainable grape productivity and the growth-yield relationship= A review" American Journal of Enology and Viticulture 52=165-174.
One Year Cane
Studies at Cornell University's Geneva Experiment Station characterized Ontogenic_Downy_Mildew_Resistance_Disposition in 'Chardonnay' grape clusters through systematic inoculation experiments across berry development stages. Clusters inoculated at 80% capfall (BBCH 68) exhibited 78% infection incidence with abundant sporulation on developing berries. Inoculation at pea-size stage (BBCH 73) resulted in 52% infection with moderate sporulation. By bunch closure (BBCH 77), infection dropped to 18% with sparse sporulation confined to pedicel attachment points. Post-véraison inoculation (BBCH 83) produced no successful infections regardless of inoculum concentration. Biochemical analysis revealed that berry skin total phenolics increased from 2.3 mg/g fresh weight at pea-size to 8.7 mg/g at véraison, while cuticle thickness doubled from 2.1 μm to 4.4 μm over the same period. These findings inform the standard recommendation for fungicide applications protecting the critical bloom-to-bunch-closure window, after which ontogenic resistance provides substantial natural protection.
An Ontogenic_Downy_Mildew_Resistance_Disposition is a Downy_Mildew_Resistance_Disposition that inheres in a Vitis plant and is realized through developmental stage-dependent defense mechanisms, with resistance levels varying systematically across the plant's ontogeny and the phenological progression of individual organs. This disposition reflects the principle that susceptibility to Plasmopara viticola is not constant throughout plant development but changes predictably with tissue age, physiological status, and developmental programming. Young, actively expanding leaves are typically most susceptible, with resistance increasing as leaves mature and reach full photosynthetic capacity. This age-related resistance correlates with changes in leaf surface properties (cuticle thickness, stomatal density and morphology), cell wall composition (increased lignification and callose content), and accumulation of constitutive defense compounds (phenolics, stilbenes). Similarly, grape berries exhibit pronounced ontogenic resistance, with high susceptibility from flowering through several weeks post-bloom, followed by progressive resistance development that renders berries essentially immune by véraison. This berry resistance correlates with epidermal cell wall modifications, accumulation of antimicrobial compounds in the skin, and loss of functional stomata. Understanding ontogenic resistance is critical for disease management timing, as protection strategies must account for the transient windows of maximum susceptibility during canopy and cluster development.
Ontogenic_Downy_Mildew_Resistance_Disposition is_a Downy_Mildew_Resistance_Disposition. This class captures developmentally programmed changes in susceptibility that occur in all Vitis genotypes, distinct from genetically variable resistance conferred by Rpv loci or quantitative factors. Ontogenic resistance operates independently of but additively with genetic resistance factors. The practical significance lies in identifying developmental windows of maximum vulnerability requiring intensive protection.
Altersabhängige Peronospora-Resistenz
Entwicklungsbedingte Falscher-Mehltau-Resistenz
Ontogenetische Peronospora-Resistenz-Disposition
age-related downy mildew resistance
developmental DM resistance
tissue maturity resistance
Kennelly MM et al. (2005) Seasonal development of ontogenic resistance to downy mildew in grape berries and rachises, Phytopathology 95:1445-1452 Ficke A et al. (2002) Ontogenic resistance and plant disease management: a case study of grape powdery mildew, Phytopathology 92:671-675 Gindro K et al. (2012) Stilbenes: biomarkers of grapevine resistance to disease of high relevance for agronomy, oenology and human health, Plant Defense: Biological Control, Springer
Ontogenic Downy Mildew Resistance Disposition
Systematic characterization of Ontogenic_Powdery_Mildew_Resistance_Disposition in 'Chardonnay' grape berries at Cornell University quantified the transition from susceptibility to resistance across developmental stages. Cluster inoculations with E. necator conidia (20,000/mL) at weekly intervals from bloom through véraison demonstrated dramatic changes in susceptibility. Inoculation at bloom (BBCH 65) resulted in 82% berry infection with dense mycelial coverage. At 2 weeks post-bloom (6°Brix), infection remained high at 71%. By 4 weeks post-bloom (10°Brix), infection dropped to 34%, and by 6 weeks (16°Brix approaching véraison), only 8% of berries became infected with restricted colony development. Post-véraison inoculation produced no successful infections. Biochemical analysis revealed that berry skin phenolic concentration increased from 1.8 to 7.2 mg/g fresh weight between bloom and véraison, while cuticle proper thickness increased from 0.4 to 1.2 μm. These findings establish the "window of susceptibility" concept guiding fungicide timing, with applications critical from pre-bloom through 4-6 weeks post-bloom, after which ontogenic resistance provides substantial natural protection.
An Ontogenic_Powdery_Mildew_Resistance_Disposition is a Powdery_Mildew_Resistance_Disposition that inheres in a Vitis plant and is realized through developmental stage-dependent changes in susceptibility, with resistance levels varying systematically across organ ontogeny and the seasonal progression of plant development. This disposition reflects the well-documented phenomenon that grapevine tissues exhibit profoundly different susceptibility to Erysiphe necator depending on their developmental age and physiological status. Young, actively expanding leaves are highly susceptible, with resistance increasing progressively as leaves mature, reaching near-immunity approximately 14-21 days after full expansion. This leaf ontogenic resistance correlates with changes in cuticle composition and thickness, epidermal cell wall architecture, accumulation of phenolic compounds, and declining metabolic activity favorable for biotrophic pathogen nutrition. Berry ontogenic resistance is particularly pronounced and economically significant, with berries highly susceptible from bloom through approximately 4 weeks post-bloom (until approximately 8°Brix), after which susceptibility declines rapidly. By véraison, berries are essentially immune to new infections regardless of inoculum pressure. This berry resistance transition involves changes in epidermal cell wall composition, cuticle restructuring during ripening, and accumulation of antimicrobial compounds. Understanding ontogenic resistance informs disease management strategies, identifying critical windows when protection is essential and periods when natural resistance provides adequate protection.
Ontogenic_Powdery_Mildew_Resistance_Disposition is_a Powdery_Mildew_Resistance_Disposition. This class captures developmentally programmed changes in susceptibility occurring in all Vitis genotypes, distinct from genetically variable resistance conferred by Ren or Run loci. Ontogenic resistance operates independently of but additively with genetic resistance factors. The practical significance lies in identifying developmental windows requiring intensive protection, with the bloom-to-early-ripening period being most critical for berry infection management.
Altersabhängige Oidium-Resistenz
Entwicklungsbedingte Echter-Mehltau-Resistenz
Ontogenetische Mehltau-Resistenz-Disposition
age-related powdery mildew resistance
developmental PM resistance
tissue maturity oidium resistance
Ficke A et al. (2002) Ontogenic resistance and plant disease management: a case study of grape powdery mildew, Phytopathology 92:671-675 Gadoury DM et al. (2003) Effects of diffuse colonization of grape berries by Uncinula necator on bunch rots, berry microflora, and juice and wine quality, Phytopathology 93:925-932 Gindro K et al. (2012) Stilbenes: biomarkers of grapevine resistance to disease, Plant Defense: Biological Control, Springer
Ontogenic Powdery Mildew Resistance Disposition
Syrah in Northern Rhône (Côte-Rôtie, Hermitage) at BBCH 65 exhibits open loose flower clusters of 14-18 cm length bearing 500-800 flowers distributed across well-developed primary and secondary branching with flower densities of 9-11 flowers/cm, creating conical clusters with moderate shoulders (OIV code 3) that develop into loose berry clusters of 200-400 berries with 2-4% Botrytis incidence in typical years with 3-4 fungicide applications, combining disease resistance with practical harvest logistics. Carignan in Languedoc shows similarly open loose architecture (12-16 cm length, 450-700 flowers, OIV code 3) providing excellent fruit health and extended hang time capability while maintaining adequate cluster weight (250-350 g) for efficient mechanical or hand harvest. Tannat in Madiran exhibits open loose clusters (OIV code 3) enabling optimal tannin maturation with minimal rot risk during extended ripening periods.
An open loose cluster is a loose flower cluster exhibiting low-to-moderate flower density with well-separated flowers on moderately branched axes, creating open structures that provide excellent air circulation and minimal disease risk while maintaining more cohesive cluster morphology and more reliable pollination than widely spaced loose clusters, representing an optimal balance between disease resistance and viticultural practicality for many wine production contexts. Open loose clusters exhibit flower densities typically ranging from 8-12 flowers per cm of total inflorescence axis length, achieved through moderate flower initiation combined with good rachis and branch elongation, well-developed secondary branching (3-6 secondary branches per primary), and moderate pedicel length (4-7 mm). These clusters typically measure 12-18 cm in length at BBCH 65 while bearing 400-800 flowers, creating open but reasonably cohesive flower distribution with flowers clearly separated but not extremely dispersed. The open loose architecture results from genetic programming for good internode elongation during inflorescence development, combined with moderate hierarchical branching that distributes flowers adequately for disease resistance while maintaining cluster integrity. Open loose flower clusters characteristically show conical or cylindrical-conical shapes with moderate lateral shoulders, typically with length=width ratios of 1.5=1-2.5=1, creating a balanced appearance with visible branching structure but recognizable cluster form. Flower spacing within open loose clusters typically ranges from 4-8 mm between adjacent flower pedicels, creating well-separated flowers with good air space but still appearing as a connected aggregate rather than as dispersed elements. The open flower arrangement creates a favorable microclimate with relative humidity within clusters elevated only 3-6% above ambient, rapid drying after precipitation (typically 1-2 hours faster than compact clusters), and good air circulation penetrating throughout the cluster volume. This microclimate reduces disease pressure substantially, with open loose flower clusters showing 60-75% lower Botrytis cinerea infection rates at bloom compared to compact clusters, typically allowing successful production with reduced fungicide programs (2-4 applications versus 6-8 for compact types) or organic production with minimal bloom-period treatments in moderately humid climates. The open architecture provides disease benefits while avoiding the extreme challenges of widely spaced loose clusters= pollination efficiency remains high due to adequate flower proximity (fruit set typically 40-55%, comparable to moderately compact types), shatter susceptibility is moderate (flower abscission 5-12% in normal conditions versus 15-30% in widely spaced types), and cluster integrity is maintained for efficient harvest handling. The transformation from open loose flower cluster to open loose berry cluster maintains good openness, with berry-berry contact delayed until late maturation (BBCH 83-87) or occurring only at mature cluster shoulders, berries experiencing minimal compression and retaining near-spherical morphology, good air circulation persisting through ripening, bunch rot incidence typically 2-5% in moderately humid climates, and cluster weights (200-400 g) providing reasonable harvest efficiency. Open loose flower cluster morphology characterizes cultivars like Syrah, Carignan, Cinsault, Tannat, and moderate selections of Grenache, representing OIV code 3 (loose).
Open_Loose_Cluster is distinguished from Widely_Spaced_Loose_Cluster by moderate rather than extreme looseness (8-12 flowers/cm versus <5-8 flowers/cm), balanced morphology maintaining cluster cohesion, and optimization for practical viticulture rather than maximum disease resistance. This class corresponds to OIV code 3 and represents an optimal architecture for many quality wine production contexts, providing substantial disease resistance (60-75% risk reduction versus compact types) while maintaining pollination reliability, harvest efficiency, and cluster integrity. The boundary between 'open loose' (OIV 3) and 'medium' compactness (OIV 5) is gradational, with OIV code 4 representing an intermediate category sometimes classified as 'moderately loose' or 'slightly loose.' Many modern breeding programs target open loose cluster architecture as providing the best balance of disease resistance, sustainability, yield efficiency, and quality potential.
moderately loose flower cluster
mäßig aufgelockerte Blütentraube
mäßig lockere Blütentraube
offen lockere Blütentraube
open flower bunch
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 94-97 OIV (2009). Descriptor List for Grape Varieties and Vitis Species, OIV code 204 level 3 Vail, M.E., Marois, J.J. (1991). Grape cluster architecture and the susceptibility of berries to Botrytis cinerea. Phytopathology 81=188-191 Intrieri, C., Poni, S. (1995). Integrated evolution of trellis training systems. American Journal of Enology and Viticulture 46=116-127.
Open Loose Cluster
An open petiolar sinus on a mature leaf at node 7 of Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, California, exhibiting characteristic open U-shaped configuration with basal lobe margins separated by 2.4 cm at widest point, sinus depth of 2.8 cm creating substantial indentation, rounded U-shaped contour with smooth sinus base, basal lobe margins diverging at approximately 65-degree angle creating wide opening clearly exposing petiole for its full length and portion of the shoot node beneath, lobe margins showing smooth edges without teeth in sinus region, moderate trichome coverage on lobe surfaces, serving as primary diagnostic feature distinguishing Cabernet Sauvignon from Merlot (closed sinus) and enabling identification even from substantial distances based on visible petiole and gap, measured at BBCH stage 77 (berries beginning to touch) in mid-July on exterior canopy mature leaf representing typical diagnostic expression, demonstrating the high reliability and visibility of open petiolar sinus as identification character in this major commercial cultivar.
A petiolar sinus exhibiting wide separation between basal lobe margins creating clearly visible gap or opening at the petiole-lamina junction in Vitis vinifera, characterized by substantial space between lobe margins typically 1-3 cm width measured at the widest point, V-shaped or U-shaped configuration with lobes diverging from petiole attachment at wide angles typically 40-90 degrees, clear visibility of petiole and often portion of shoot through the sinus opening, occurrence in many major cultivars including Cabernet Sauvignon (open U-shaped), Cabernet Franc (open V-shaped), Sauvignon Blanc (open V-shaped), and Chardonnay (open U-shaped to slightly closed), high diagnostic value as this configuration reliably distinguishes these open-sinus cultivars from closed-sinus varieties like Merlot or overlapping-sinus types like Pinot Noir, and functional implications potentially including improved air circulation around the petiole-shoot junction reducing disease pressure though this remains largely speculative without clear experimental support. Open petiolar sinuses develop when basal lobe primordia diverge widely during early leaf development, with genetic programming determining lobe angle and growth trajectory creating the characteristic gap, and environmental conditions having relatively minor effects on this strongly genetically controlled character making open versus closed sinus one of the most stable and reliable ampelographic features.
Open petiolar sinuses represent one of three primary petiolar sinus categories, distinguished from closed sinuses by substantial gap between lobe margins and from overlapping sinuses by divergent rather than convergent lobe orientation. The open configuration shows variation in shape (U-shaped versus V-shaped) and width (moderately open versus very wide), creating subcategories useful for finer cultivar discrimination. U-shaped open sinuses (Cabernet Sauvignon, Chardonnay) show rounded contours with lobes diverging gradually, while V-shaped open sinuses (Cabernet Franc, Sauvignon Blanc) show more acute angles and straight lobe edges creating sharper appearance. Width variation affects visibility and diagnostic ease, with very wide open sinuses (some Syrah clones) enabling identification from substantial distances while moderately open sinuses require closer examination. Genetic control is strong with open versus closed sinus showing simple inheritance patterns in many crosses, suggesting major gene effects determining this character. Environmental stability is high compared to many leaf features, with open sinus configuration maintained across wide ranges of vigor, light exposure, and developmental timing, though extreme stress may produce slightly narrower sinuses in affected leaves. Ampelographic utility is very high, with open petiolar sinus serving as primary or secondary diagnostic feature in most cultivar identification keys.
divergent lobe sinus
divergierende Basalbucht
gaping basal sinus
offener Petiolarsinus
weite Stielbucht
wide petiolar sinus
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 90-135 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 56-78.
Open Petiolar Sinus
Operational stomatal conductance monitoring in Merlot through a drying cycle tracks the progressive stress response, with well-watered vines showing midday gs of 0.32 mol/m²/s (85% of maximum, minimal stress), moderate deficit showing midday gs of 0.18 mol/m²/s (47% of maximum, significant stomatal limitation), and severe stress showing gs of 0.06 mol/m²/s (16% of maximum, severe closure approaching minimum conductance), with the progressive decline in operational conductance indicating the stress level requiring irrigation intervention to prevent yield and quality impacts.
Operational stomatal conductance quality is a stomatal conductance quality that represents the actual stomatal conductance under current field conditions as regulated by environmental signals and plant water status, characterizing the realized conductance that reflects the integration of multiple regulatory inputs and determines actual rates of photosynthesis and transpiration at any given moment. Within the BFO framework, this quality specifically addresses the realized dimension of stomatal function, distinguishing the current regulated state from maximum anatomical capacity while recognizing that operational conductance is the physiologically relevant parameter that determines actual gas exchange performance and water use under the constantly varying conditions of the vineyard environment. The ontological significance lies in operational stomatal conductance determining actual vine carbon gain and water loss at any moment, where the degree to which stomata are open relative to maximum capacity reflects the plant's integrated response to current light, humidity, soil water status, and other factors, providing real-time information about vine physiological status and the trade-off between productivity and water conservation currently being expressed. The regulatory integration determining operational conductance involves multiple signaling pathways including photoreceptor-mediated light responses (blue light opening, red light through photosynthesis), hydraulic signals (turgor-mediated responses to water status), chemical signals (ABA accumulation promoting closure), and atmospheric signals (VPD and CO₂ effects), with the relative influence of these factors varying with environmental conditions and stress level. The typical patterns of operational conductance show diurnal variation with morning increase (light-stimulated opening), potential midday depression (high VPD and/or water stress causing partial closure), and evening decline (light-mediated closure), with the amplitude and timing of these patterns indicating vine water status and regulatory strategy. The measurement of operational conductance employs the same techniques as maximum conductance but under field conditions, with the ratio of operational to maximum conductance indicating the degree of stomatal closure and thus the severity of regulatory limitation on gas exchange. Understanding operational stomatal conductance quality enables real-time assessment of vine water stress, prediction of current photosynthetic performance, and timing of irrigation based on the stomatal closure indicating developing water deficit.
Operational_Stomatal_Conductance_Quality addresses the actual, regulated stomatal conductance under field conditions. This quality reflects the integration of multiple environmental and physiological signals determining current gas exchange. The class supports real-time stress assessment and irrigation timing through operational conductance monitoring.
Feldleitfähigkeit
actual gs
aktuelle gs
field conductance
operative stomatäre Leitfähigkeit
realized stomatal opening
regulated conductance
regulierte Stomataöffnung
Medrano H et al. (2002) Regulation of photosynthesis of C₃ plants in response to progressive drought. Journal of Experimental Botany 53(366): 1-12 Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell & Environment 26(8): 1393-1405
Operational Stomatal Conductance Quality
In Riesling berries in Wachau, Austria during BBCH 85-89, optimal aroma development achieves total monoterpene concentration of 2.8 mg/L at 6.5-fold average perception threshold with 8 individual terpenoids above detection, norisoprenoid precursors of 185 µg/L contributing 5 distinct scent notes including characteristic TDN precursor for aged petrol character, methoxypyrazine at 4 ng/L providing subtle herbaceous complexity without dominating green character, with trained panel overall aromatic quality rating of 9.3 on 10-point scale, complexity score of 8.9, and varietal typicity of 9.1, representing near-ideal expression under exceptional vintage conditions with mean ripening temperature of 19°C, 12% vine water deficit, and optimal canopy management achieving 35% cluster-zone leaf removal.
Optimal aroma development represents the idealized trajectory of volatile aromatic compound biosynthesis, accumulation, and maturation in Vitis vinifera berries wherein environmental conditions, viticultural management, and inherent genetic potential combine to produce maximum expression of desirable olfactory characteristics at harvest, achieving the highest possible aromatic quality within the constraints of variety, site, and vintage conditions over the 35-50 day post-veraison ripening period. This concept integrates multiple aromatic quality dimensions including sufficient intensity with key impact compounds exceeding perception thresholds by factors of 5-15 generating pronounced varietal character, adequate complexity with 12-18 or more distinct scent notes contributing to overall olfactory impression, appropriate balance between fruity-floral and green-herbaceous elements strongly favoring ripe character, and strong varietal typicity enabling clear cultivar identification and meeting consumer expectations for specific grape varieties. The achievement of optimal aroma development requires coordination among environmental factors including moderate temperatures of 18-25°C preserving accumulated volatiles while supporting continued biosynthesis, adequate but not excessive light exposure promoting terpenoid pathway expression without thermal stress, and appropriate vine water status imposing mild deficit that enhances secondary metabolism without compromising photosynthetic function or inducing excessive berry dehydration. Viticultural practices supporting optimal aroma development encompass canopy management ensuring cluster microclimate optimization for enhanced terpene accumulation, crop load adjustment balancing vine capacity with aromatic compound concentration objectives, irrigation scheduling imposing controlled water deficit during the post-veraison period that activates stress-responsive secondary metabolism, and harvest timing capturing peak aromatic quality before potential late-season degradation or overripeness shifts olfactory profiles away from optimal expression. Assessment of optimal aroma development employs integrated evaluation methods combining analytical quantification of marker compounds with sensory assessment of overall aromatic quality, recognizing that premium wine production requires achievement of target values across multiple parameters including intensity, complexity, balance, and typicity rather than optimization of individual factors in isolation. Vintage variation in optimal aroma development potential reflects climatic differences affecting biosynthetic processes, with favorable years achieving near-ideal aromatic expression while challenging vintages may require management intervention or quality expectations adjustment to accommodate suboptimal development conditions, with the consistency of achieving optimal development across vintages representing a hallmark of superior vineyard sites and management programs.
Optimal_Aroma_Development is subclass of Aroma_Profile_Maturation representing the quality ideal for aromatic processes. Distinguishes from descriptive classes by emphasizing optimization. Sibling to Mature_Aroma_Expression and Cultivar_Typical_Aroma. Integrates intensity, complexity, balance, and typicity. Requires coordination of genetics, environment, and management. Benchmark for aromatic quality assessment. Vintage-dependent achievement.
Spitzen-Duft-Erreichung
ideal aroma development
ideale Duft-Entwicklung
maximale aromatische Qualität
maximum aromatic quality
optimal olfactory expression
optimale Aromaentwicklung
peak scent achievement
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 305-315 Reynolds AG (2010) Managing Wine Quality Vol 1 Viticulture and Wine Quality Woodhead Publishing Robinson AL et al (2014) Origins of grape and wine aroma. American Journal of Enology and Viticulture 65: 1-30
Optimal Aroma Development
In Sémillon berries in Pessac-Léognan, Bordeaux during BBCH 85-89, optimal flavor development achieves free monoterpene concentration of 0.65 mg/L at 4.2-fold average perception threshold, norisoprenoid precursors of 165 µg/L providing 5 distinct aromatic notes, volatile thiol precursors of 320 µg/L enabling tropical-citrus complexity, with trained panel overall quality rating of 9.1 on 10-point scale, complexity score of 8.8, and varietal typicity of 8.6, representing near-ideal expression under exceptional vintage conditions with mean ripening temperature of 23°C, 15% vine water deficit, and 40% cluster-zone leaf removal optimizing light exposure.
Optimal flavor development represents the idealized trajectory of flavor compound biosynthesis, accumulation, and maturation in Vitis vinifera berries wherein environmental conditions, viticultural management, and inherent genetic potential combine to produce maximum expression of desirable aromatic characteristics at harvest, achieving the highest possible quality within the constraints of variety, site, and vintage conditions over the 35-50 day post-veraison ripening period. This concept integrates multiple quality dimensions including sufficient flavor intensity exceeding perception thresholds by factors of 3-10 for key impact compounds, adequate complexity with 10-15 or more distinct aromatic notes contributing to overall impression, appropriate balance between fruity-floral and green-herbaceous elements favoring ripe character, and strong varietal typicity enabling clear identification and meeting consumer expectations for specific cultivars. The achievement of optimal flavor development requires coordination among environmental factors including moderate temperatures of 20-26°C supporting biosynthetic enzyme activities, adequate but not excessive light exposure promoting secondary metabolism without thermal stress, and appropriate vine water status imposing mild stress that enhances flavor compound accumulation without compromising photosynthetic capacity or berry physiological function. Viticultural practices supporting optimal flavor development encompass canopy management ensuring cluster microclimate optimization, crop load adjustment balancing vine capacity with fruit quality objectives, irrigation scheduling imposing controlled water deficit during the post-veraison period, and harvest timing capturing peak flavor quality before potential late-season degradation or overripeness shifts aromatic profiles away from optimal expression. The assessment of optimal flavor development employs integrated evaluation methods combining analytical quantification of marker compounds with sensory assessment of overall quality, with premium wine production requiring achievement of target values across multiple parameters rather than optimization of individual factors in isolation. Vintage variation in optimal flavor development potential reflects climatic differences affecting biosynthetic processes, with favorable years achieving near-ideal expression while challenging vintages may require management intervention or quality expectations adjustment to accommodate suboptimal development conditions.
Optimal_Flavor_Development is subclass of Flavor_Profile_Maturation representing the quality ideal for flavor processes. Distinguishes from descriptive classes by emphasizing optimization. Sibling to Mature_Flavor_Expression and Cultivar_Specific_Flavor. Integrates intensity, complexity, balance, and typicity. Requires coordination of genetics, environment, and management. Benchmark for quality assessment. Vintage-dependent achievement.
Spitzen-Aroma-Erreichung
ideal flavor development
ideale Geschmacksentwicklung
maximale Aroma-Qualität
maximum flavor quality
optimal aromatic expression
peak flavor achievement
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 305-315 Reynolds AG (2010) Managing Wine Quality Vol 1 Viticulture and Wine Quality Woodhead Publishing Jackson RS (2020) Wine Science Principles and Applications 5th Ed Chapter 6
Optimal Flavor Development
An optimal function mature leaf at node 7 on a shoot of Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, California, exhibiting light-saturated photosynthesis rate of 13.7 μmol CO₂ m⁻² s⁻¹ measured at 1500 μmol photons m⁻² s⁻¹ irradiance, stomatal conductance of 285 mmol H₂O m⁻² s⁻¹ at midday under moderate vapor pressure deficit of 1.8 kPa, chlorophyll content of 485 μg cm⁻² with a=b ratio of 3.1, maximum PSII efficiency Fv/Fm of 0.82 measured predawn indicating no photoinhibition, leaf nitrogen content of 2.5% dry weight indicating adequate nutritional status, water potential at midday of -1.1 MPa indicating moderate well-watered status, leaf temperature of 26°C maintained 2-3°C below air temperature through transpirational cooling, occurring on a well-managed vineyard with balanced irrigation (soil moisture 65-75% field capacity), appropriate fertilization (80 kg N/ha), and optimal crop load (1.8 kg fruit per meter canopy), measured at BBCH stage 81 (beginning of berry ripening) in early August, representing ideal physiological function contributing approximately 18 grams carbohydrate per week supporting fruit ripening and reserve accumulation.
A fully expanded mature leaf exhibiting peak physiological performance across multiple functional parameters in Vitis vinifera, characterized by light-saturated photosynthesis rates in the upper range for the species typically 12-16 μmol CO₂ m⁻² s⁻¹, high stomatal conductance of 200-400 mmol H₂O m⁻² s⁻¹ under favorable conditions enabling efficient CO₂ uptake while maintaining water use efficiency, maximum chlorophyll concentrations typically 400-600 μg cm⁻² indicating high photosynthetic enzyme content, favorable chlorophyll a=b ratio of 2.8-3.5 reflecting optimal light-harvesting complex composition, low photorespiration rates indicating efficient carbon fixation, high quantum efficiency with maximum PSII efficiency (Fv/Fm) of 0.80-0.83 indicating healthy photosystem function without photoinhibition, and optimal nitrogen content of 2.2-2.8% dry weight supporting protein synthesis for photosynthetic enzymes and other metabolic functions. Optimal function mature leaves develop and persist under favorable environmental conditions including moderate temperatures of 20-28°C maintaining enzyme function within optimal kinetic ranges, adequate but not excessive water availability supporting stomatal opening and maintaining mesophyll conductance, sufficient nitrogen and mineral nutrition supporting chlorophyll synthesis and enzyme production, moderate light intensity of 800-1500 μmol photons m⁻² s⁻¹ saturating photosynthesis without causing photoinhibition, and absence of stress factors including pests, diseases, air pollutants, or salinity that would compromise physiological function.
Optimal function mature leaves are distinguished from other mature leaves by superior performance across multiple physiological parameters rather than single characteristics, requiring comprehensive assessment including gas exchange measurements, chlorophyll fluorescence, pigment analysis, and nutritional status. Achievement and maintenance of optimal function depends critically on environmental conditions, with temperatures outside 18-30°C range, water stress reducing soil moisture below 50% field capacity, nitrogen deficiency, excessive crop loads, or pest and disease pressure all compromising function despite leaves achieving mature structural status. Position in canopy affects function through light availability, with interior shade leaves rarely achieving optimal function due to light limitation regardless of other factors, while exterior sun-exposed leaves risk photoinhibition under excessive irradiance. Optimal function is temporally dynamic, with daily patterns following diurnal environmental changes and seasonal patterns reflecting developmental progression from maturation through senescence, typically maintaining peak performance for 6-10 weeks under favorable conditions before gradual functional decline.
höchstleistungs Blatt
maximum efficiency leaf
optimal-funktionierendes Blatt
peak function leaf
physiologically optimal leaf
physiologisch optimales Weinblatt
Based on Flexas et al. (2002) "Drought-inhibition of photosynthesis in C₃ plants= stomatal and non-stomatal limitations revisited" Annals of Botany 89=183-189 Chaves et al. (2010) "Grapevine under deficit irrigation= hints from physiological and molecular data" Annals of Botany 105=661-676 Palliotti et al. (2012) "Morpho-anatomical and physiological response of Pinot Noir and Montepulciano grapevine leaves to mild and severe water stress" Vitis 51=109-116 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 143-178.
Optimal Function Mature Leaf
Optimal harvest date quality analysis for Pinot Noir in Burgundy established predictive relationships enabling vintage planning across the appellation hierarchy. Village-level analysis demonstrated that optimal harvest date correlated strongly with growing degree day accumulation from April 1 (r²=0.87), with harvest occurring at mean 1,450 GDD (base 10°C) with standard deviation of only 65 GDD translating to ±4 days around the predicted date in typical vintages. The véraison-to-harvest interval showed even tighter optimal harvest date quality with mean 48 days and standard deviation of 3.5 days, enabling confident scheduling approximately 6 weeks in advance once véraison date was established. Premier Cru sites exhibited slightly lower optimal harvest date quality predictability (r²=0.82) attributed to greater microclimate variation affecting individual plot ripening trajectories. Climate change analysis showed progressive advancement of optimal harvest date by 2.1 days per decade since 1980, requiring dynamic updating of prediction models to maintain forecast accuracy.
Optimal harvest date quality is a harvest window quality that inheres in a grapevine cultivar or clone within a specific growing environment and represents the predictability and consistency of the calendar date or thermal time accumulation at which fruit achieves the compositional and sensory characteristics defining target ripeness for intended wine style, enabling advance planning of harvest logistics, labor scheduling, and winery preparation based on historical patterns modified by current-season monitoring data. This quality determines the reliability of vintage planning systems and the lead time available for operational preparation, with highly predictable optimal dates enabling confident advance scheduling while variable dates require flexible, responsive approaches with resources held in reserve until ripening trajectory clarifies. The ontological significance of optimal harvest date quality lies in its practical value for production management, wherein predictable harvest dates enable efficient resource allocation, contract labor scheduling, equipment positioning, and winery preparation while unpredictable dates create operational uncertainty requiring costly standby capacity and last-minute mobilization. The biological basis of optimal harvest date predictability encompasses the consistency of growing season heat accumulation patterns, the stability of the relationship between thermal time and ripening progression for specific cultivar-site combinations, the buffering capacity of vine and fruit physiology against environmental variation, and the strength of correlation between early-season indicators and final harvest timing enabling forecasting from budbreak or véraison dates. Cultivar-environment combinations exhibiting high optimal harvest date quality demonstrate consistent relationships between thermal time accumulation and harvest date, typically within ±5 days across normal vintage variation, while those with poor predictability may vary by ±15 days or more requiring continuous monitoring and flexible response capacity. The assessment of optimal harvest date quality involves compilation of multi-vintage harvest timing records correlated with seasonal weather patterns, establishment of predictive relationships between early-season phenology and harvest date, and quantification of the forecast accuracy achievable with available prediction methodologies for the specific cultivar-site combination.
Optimal_Harvest_Date_Quality is_a Harvest_Window_Quality, representing the predictability and consistency of when optimal ripeness is achieved. This class addresses the timing of the harvest window opening rather than its duration. The GDD-based and véraison-interval prediction approaches provide practical forecasting tools. Distinguished from Harvest_Window_Duration_Quality which addresses how long acceptable quality persists after optimal date. Climate change effects on harvest date progression require consideration in long-term planning.
Erntedatumskonsistenz
Leseterminvorhersagbarkeit
Optimale-Erntedatum-Qualität
Weinlesedatum-Präzision
harvest date predictability
optimal picking date
target harvest timing
vintage date consistency
Jones GV, Davis RE (2000) Climate influences on grapevine phenology, grape composition, and wine production and quality for Bordeaux, France. American Journal of Enology and Viticulture 51:249-261 Parker AK et al. (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17:206-216 Webb LB et al. (2012) Modelled impact of future climate change on the phenology of winegrapes in Australia. Australian Journal of Grape and Wine Research 18:20-31
Optimal Harvest Date Quality
Optimal harvest window determination for Pinot Noir reveals the cultivar's characteristically brief quality peak, with maturity monitoring indicating window opening at 23.5 Brix (acid balance achieved, color development adequate) on September 15, optimal quality peak around 24.2 Brix on September 20 (maximum flavor development, balanced phenolics), and window closing by September 26 (acid dropping below target, flavors shifting toward over-ripe, stem lignification complete enabling whole-cluster consideration), yielding a 11-day window with 5-day optimal center, requiring harvest scheduling that captures the September 18-22 period for premium quality lots while conditions permit, demonstrating the temporal precision required for cultivars with compact harvest windows.
Optimal harvest window quality is a harvest readiness quality that represents the temporal period during which fruit quality is at or near its peak and conditions permit successful harvest, characterizing the identification and duration of the time frame when the combination of grape maturity and harvest feasibility creates the opportunity for quality-maximizing fruit collection. Within the BFO framework, this quality specifically addresses the temporal optimization dimension of harvest timing, distinguishing the period of quality opportunity from the broader period of acceptable maturity while recognizing that optimal windows may be brief (requiring rapid response) or extended (permitting deliberate timing), with window duration affecting the operational pressure and decision-making flexibility available during harvest. The ontological significance lies in optimal harvest window defining the quality-maximizing opportunity that harvest must capture, where the window opens when fruit achieves target maturity parameters (or the trajectory indicates that further waiting offers diminishing or negative returns) and closes when fruit quality begins declining (through over-ripeness, rain damage, disease development, or other deterioration), with the duration between opening and closing determining how much flexibility exists for operational scheduling. The determination of optimal harvest window involves assessment of when maturity parameters enter the target range (window opening), prediction of when quality will peak (window center), and estimation of when quality will begin declining (window closing), with these assessments drawing on compositional analysis, trajectory extrapolation, weather forecasting, and experience with cultivar-specific ripening patterns. The window duration factors include cultivar characteristics (some varieties holding condition well, others deteriorating rapidly after optimal maturity), weather conditions (cool, dry weather extending windows while heat or rain shortens them), fruit condition (healthy fruit maintaining quality longer than diseased or damaged fruit), and the specific quality parameters prioritized (windows defined by sugar stability longer than those defined by aromatic peak). The management implications of optimal harvest window include the necessity of advance preparation (ensuring capacity before window opens), rapid response capability (mobilizing harvest quickly for brief windows), and triage decision-making (prioritizing among competing blocks when windows overlap), with successful harvest management requiring both accurate window prediction and operational agility to capture quality opportunities. Understanding optimal harvest window quality enables identification of the quality-maximizing harvest period, operational planning to capture brief windows, and recognition of the factors that extend or compress the opportunity for optimal harvest timing.
Optimal_Harvest_Window_Quality addresses the identification of the quality-maximizing harvest period. This quality defines when the combination of maturity achievement and harvest feasibility permits optimal fruit collection. The class supports harvest planning through window characterization enabling capture of quality opportunities.
Lesefenster
Qualitätsfenster
Reifefenster
harvest window
maturity window
optimal picking period
optimales Erntefenster
quality window
Coombe BG, McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6(2): 131-135 Reynolds AG (2010) Managing Wine Quality, Volume 1: Viticulture and Wine Quality. Woodhead Publishing, Cambridge, pp. 111-155
Optimal Harvest Window Quality
Optimal photosynthesis temperature determination for Chardonnay in Burgundy using leaf gas exchange measurements identifies Topt of 26.5°C with maximum assimilation rate of 14.2 μmol CO₂/m²/s, declining to 11.8 μmol at 32°C and 8.3 μmol at 38°C, explaining the observed 30% reduction in seasonal carbon gain during the 2019 heat wave where afternoon temperatures routinely exceeded 35°C for three weeks during the critical fruit development period, contributing to smaller berry size and lower yield.
Optimal photosynthesis temperature quality is a critical temperature threshold quality that represents the temperature or temperature range at which grapevine leaf photosynthetic carbon assimilation rate reaches its maximum, characterizing the thermal conditions where the balance between carboxylation rate increase and photorespiration increase yields peak net CO₂ fixation. Within the BFO framework, this quality identifies the temperature providing optimal physiological function for this specific process, distinguishing the photosynthetic optimum from optima for other processes such as cell division, anthocyanin synthesis, or sugar transport that may occur at different temperatures. The ontological significance lies in photosynthetic optimum determining the temperature range for maximum carbon gain supporting vine growth, yield formation, and fruit sugar accumulation, where departure from optimum reduces photosynthetic efficiency with consequences for whole-vine productivity and fruit ripening capacity. The optimal temperature for grapevine photosynthesis typically ranges from 25-30°C for most Vitis vinifera cultivars, with the optimum shifting slightly higher (28-33°C) for varieties adapted to warm climates and lower (20-25°C) for cool-climate varieties, reflecting genotypic variation in Rubisco kinetics and membrane thermostability. The optimum represents a balance point where rising temperature increases enzymatic reaction rates but also increases photorespiration and eventually causes protein denaturation, with net photosynthesis declining above the optimum at approximately 3-5% per degree. Measurement requires gas exchange determination under controlled temperature conditions using photosynthesis systems that maintain constant light, CO₂, and humidity while varying leaf temperature, with the optimum identified as the temperature yielding maximum net CO₂ assimilation rate. Understanding photosynthetic temperature optimum enables prediction of seasonal carbon gain patterns, evaluation of climate suitability for specific varieties, and interpretation of vintage variation in ripening capacity related to temperature conditions during the growing season.
Optimal_Photosynthesis_Temperature_Quality addresses the functional optimum for carbon assimilation, complementing the damage thresholds (Freezing_Point, Heat_Damage_Threshold) that define survival limits. This quality varies with measurement conditions including CO₂ concentration, light intensity, and leaf age, requiring standardized protocols for cultivar comparison. The class supports climate matching for variety selection and productivity modeling incorporating temperature effects on canopy carbon gain.
Temperaturoptimum der Photosynthese
Topt for photosynthesis
Topt für Kohlenstoffassimilation
maximum photosynthesis temperature
optimal temperature for carbon assimilation
optimale Photosynthesetemperatur
photosynthetic thermal optimum
photosynthetisches Temperaturoptimum
Greer DH, Weston C (2010) Heat stress affects flowering, berry growth, sugar accumulation and photosynthesis of Vitis vinifera cv. Semillon grapevines grown in a controlled environment. Functional Plant Biology 37(3): 206-214 Zufferey V et al. (2000) Diurnal cycles of CO₂ assimilation by field-grown grapevine leaves. Vitis 39(2): 71-76
Optimal Photosynthesis Temperature Quality
Merlot in Bordeaux shows optimal pollen cloud formation during mid-June Year N+1 at BBCH 63-65 on June 9-10, with visible pale yellow haze surrounding flowering clusters during morning hours 9am-11am, prominent under favorable oceanic conditions of 23-27°C temperature, 56-66% relative humidity, light winds 7-11 km per hour, confirming intensive pollen release and optimal pollination conditions, typical of this mid-season cultivar during peak flowering supporting 56-66% fruit set
Optimal pollen cloud formation represents the visible atmospheric pollen phenomenon during peak release when favorable environmental conditions and intensive pollen discharge create characteristic pale yellow haze surrounding flowering clusters, providing obvious visual confirmation of maximum male reproductive activity and optimal pollination opportunity. This distinctive visual phenomenon occurs during peak pollen release in mid-June in temperate regions Year N+1, typically visible during morning peak hours when conditions most favorable for cloud formation and persistence, corresponding to BBCH 63-65 peak anthesis stages. Cloud formation mechanism involves intensive coordinated pollen discharge from numerous anthers releasing millions of grains creating high local atmospheric concentration, light wind and air movement lifting and suspending released pollen creating airborne cloud, appropriate humidity maintaining pollen in dispersible state without excessive aggregation or rapid settling, and favorable temperature supporting both discharge and atmospheric suspension. Visual characteristics show pale yellow to yellow-brown coloration creating visible haze around flowering clusters, concentration gradient with densest cloud immediately surrounding clusters declining with distance, dynamic appearance with cloud thickness varying with discharge intensity and wind conditions, and transient nature appearing and disappearing with changing environmental conditions and discharge activity. Cloud visibility factors include pollen concentration with visible clouds typically requiring 300-500+ grains per cubic centimeter minimum threshold, lighting conditions with backlighting or low sun angle enhancing visibility, observer position with cloud more visible from certain angles, and atmospheric clarity with humid or hazy conditions reducing visibility. Optimal formation conditions include temperature 24-30°C promoting maximum discharge intensity and appropriate atmospheric properties, humidity 50-65% maintaining pollen dispersibility without causing aggregation that would prevent cloud formation, light wind 5-12 km per hour providing sufficient movement for suspension while not dispersing cloud excessively, and overall stable conditions maintaining favorable parameters during peak discharge period. Temperature effects on cloud formation show warm conditions facilitating intensive discharge providing abundant pollen source, affecting air density and buoyancy influencing suspension dynamics, and supporting atmospheric mixing creating favorable suspension conditions, with optimal 25-28°C range supporting visible cloud formation. Humidity effects show critical importance with moderate levels essential for maintaining pollen in dispersible state enabling cloud formation, high humidity above 70% causing pollen aggregation eliminating cloud through premature settling as heavy clumps, and low humidity below 45% potentially acceptable for cloud formation though possibly affecting pollen viability, with optimal 52-68% range reliably supporting cloud appearance. Wind effects show dual role with light movement necessary for lifting pollen into suspension creating cloud, though excessive wind above 15 km per hour rapidly dispersing cloud reducing visibility, calm conditions potentially limiting cloud formation through inadequate lifting though local accumulation may still occur, and optimal 6-13 km per hour providing balance supporting visible persistent clouds. Diurnal pattern shows characteristic morning maximum with cloud formation typically most prominent 9am-12pm when temperature rising, humidity declining, and discharge intensity peaking creating ideal combination, afternoon reduction as conditions potentially become less favorable with rising temperature or changing humidity, and daily rhythm with repeated cloud formation during multi-day peak release period. Spatial distribution shows clouds concentrated around flowering clusters as pollen source with highest density immediately surrounding inflorescences, extension throughout vine canopy creating larger clouds in dense plantings, vineyard-scale clouds possible during intensive synchronized discharge under optimal conditions, and visible gradient showing concentration decline with distance from source. Temporal persistence shows clouds lasting minutes to hours during favorable stable conditions with sustained discharge maintaining cloud density, rapid disappearance when conditions change with rising humidity causing aggregation or strong wind causing dispersal, and overall transient dynamic appearance responding rapidly to environmental fluctuations. Pollen grain properties affecting cloud formation include small size 25-35 micrometers enabling easy airborne suspension, relatively low density facilitating buoyancy, surface properties affecting aggregation tendency with smooth exine promoting individual grain suspension, and overall characteristics well-suited for atmospheric suspension and cloud formation. Cloud composition shows predominantly individual pollen grains suspended in air creating visible haze, occasional small pollen clumps from limited aggregation contributing to visibility, and dust or other atmospheric particles potentially contributing minor component to overall haze appearance. Acoustic properties during cloud formation include normally silent appearance with no characteristic sounds, though very intensive synchronized discharge creating clouds may produce subtle rustling though typically inaudible, and overall visual rather than acoustic phenomenon. Observational value of cloud formation includes providing obvious visual confirmation of intensive pollen release easily assessed without equipment, indicating favorable male reproductive conditions supporting high pollination potential, enabling timing assessment for management decisions or harvest predictions, and serving as practical marker for peak flowering more readily observable than detailed reproductive organ examination. Photographic documentation of clouds enables visual records of flowering intensity, comparison across seasons or locations assessing environmental effects, and research documentation of pollen release dynamics and environmental responses. Pollination opportunity indicated by clouds shows optimal conditions with visible clouds confirming abundant fresh pollen widely distributed, favorable environmental parameters supporting pollen viability and dispersal, and intensive male reproductive activity creating maximum pollination potential supporting high fruit set. Fruit set prediction from cloud observation shows positive correlation with prominent persistent clouds during peak indicating favorable conditions likely producing high set 60-75%, while absent or weak clouds suggesting suboptimal conditions potentially limiting set to 35-55%, and providing early visual indicator of reproductive success before berry development confirms outcomes. Cultural significance of pollen clouds includes traditional recognition by viticulturists as favorable flowering sign, visual confirmation of reproductive timing supporting management scheduling, and aesthetic appreciation of visible reproductive phenomenon representing vineyard fertility. The process represents first aspect of peak pollen release emphasizing visible atmospheric phenomenon, followed by maximum pollen availability discussing gamete supply. Duration represents transient phenomenon visible for hours during favorable peak conditions, with appearance and disappearance responsive to environmental and discharge intensity changes. Cultivar variation in cloud formation shows varieties with high pollen production more likely to produce prominent clouds, timing differences reflecting overall phenology, and some varieties rarely showing visible clouds despite adequate pollen release possibly due to discharge patterns or environmental sensitivity. Regional expression shows cloud formation more common in warm dry climates where favorable conditions consistently achieved, less frequent in cool humid maritime climates where humidity often too high, and occasional dramatic clouds in any region during exceptionally favorable flowering weather. Viticultural significance includes providing practical visual marker for peak male reproductive activity, indicating favorable pollination conditions supporting high fruit set predictions, enabling timing assessment for management decisions, and serving as readily observable phenomenon confirming successful flowering without technical assessment. Research applications include documenting pollen release intensity through cloud observation, analyzing environmental conditions enabling cloud formation, evaluating pollen dispersal dynamics through cloud behavior, developing practical observation protocols for phenological assessment, and investigating cloud formation as indicator of reproductive success and fruit set potential.
Optimal_Pollen_Cloud_Formation is first aspect of Peak_Pollen_Release, distinguished by visible atmospheric phenomenon confirming maximum male reproductive activity. Represents readily observable marker for peak flowering. Requires specific environmental conditions particularly moderate humidity and light wind. Transient phenomenon lasting hours during favorable conditions. Provides practical visual confirmation of intensive pollen release and pollination opportunity. Cultural and management significance as flowering indicator. Followed by Maximum_Pollen_Availability discussing gamete supply.
Pollendunstwolke
Pollennebelphänomen
Pollenwolkenbildung
atmospheric pollen display
atmosphärische Pollenanzeige
pollen cloud appearance
pollen cloud phenomenon
visible pollen haze
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 383-386 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 293-295
Optimal Pollen Cloud Formation
Merlot in Bordeaux shows optimal pollination window during mid-June Year N+1 at BBCH 63-65 over 2-3 days June 9-11, with 55-68% flowers simultaneously open, abundant fresh pollen available showing 76-85% viability, widespread stigma receptivity across cluster, under ideal oceanic climate conditions of 23-27°C temperature, 52-68% relative humidity, light winds 6-12 km per hour, stable weather without rain, enabling efficient pollination achieving 54-65% fruit set typical of this mid-season cultivar under favorable flowering conditions determining season yield potential
Optimal pollination window represents the critical temporal period during peak anthesis when combined male and female reproductive capacities reach maximum levels creating ideal conditions for successful pollination, establishing most favorable opportunity for high fruit set determining season yield potential. This essential reproductive opportunity occurs during peak anthesis in mid-June in temperate regions Year N+1, typically spanning 2-3 days encompassing maximum flower opening, peak pollen availability, and widespread stigma receptivity, corresponding to BBCH 63-65 stages. Window characteristics show simultaneous occurrence of key reproductive elements including 50-75% flowers open exposing reproductive organs, abundant fresh viable pollen available from widespread anther dehiscence, widespread stigma receptivity with active exudate production and pollen capture capacity, and favorable environmental conditions supporting reproductive organ function and pollination processes. Temporal overlap optimization shows critical coordination between pollen availability and stigma receptivity with peak pollen production coinciding with maximum proportion of receptive stigmas, temporal synchrony creating high probability of successful pollen-stigma contact, and brief overlap window requiring efficient pollination during limited favorable period. Pollination efficiency during optimal window reaches maximum levels with estimated successful pollination rates of 50-70% flowers under favorable conditions, compared to 20-40% during suboptimal early or late anthesis periods, and 10-30% or less if optimal window experiences adverse weather, demonstrating critical importance of favorable conditions during this brief period. Environmental requirements for optimal window include temperature ideally 24-30°C supporting maximum pollen and stigma function with pollen viability optimized, anther dehiscence proceeding efficiently, and stigma receptivity maintained at peak levels. Humidity optimally 45-65% relative humidity balancing pollen viability maintenance with anther dehiscence facilitation and stigma exudate properties, avoiding extremes that impair reproductive function. Weather stability critically important with consistent favorable conditions enabling sustained efficient pollination throughout optimal period, while variable conditions create erratic patterns reducing overall success. Wind conditions showing light-moderate breezes 5-15 km per hour optimal for pollen dispersal enhancing pollination efficiency without causing damage or excessive desiccation. Absence of rain absolutely critical with precipitation during optimal window catastrophically affecting pollination through preventing anther dehiscence, washing pollen from structures, eliminating stigma exudate, promoting fungal infection, and physically damaging delicate reproductive organs. Pollination mechanisms during optimal window emphasize wind-assisted self-pollination as dominant mode with pollen dispersing from anthers to stigmas within same flower through close proximity and slight air movement, between neighboring flowers on same cluster through short-distance wind transport, and occasionally between flowers on different clusters or vines through longer-distance dispersal, with self-compatibility enabling effective reproduction without requiring cross-pollination. Pollen-stigma interactions during optimal window proceed efficiently with receptive stigmas capturing airborne pollen grains through adhesive exudate, pollen hydrating rapidly from exudate enabling metabolic activation within 30-90 minutes, pollen recognition occurring through molecular signals between pollen coat proteins and stigma components, and pollen germination initiating within 1-4 hours producing pollen tube penetrating stigma surface. Pollen tube growth following pollination during optimal window proceeds rapidly with tubes elongating 1-3 mm per hour through style transmitting tract tissues, requiring 12-36 hours to traverse typical 15-30 mm style length reaching ovary, with tube growth guided by chemotactic signals from ovules directing growth through micropyle into embryo sac. Fertilization timing shows earliest successful fertilizations occurring 18-36 hours after pollination during optimal window, with double fertilization characteristic of angiosperms occurring with one sperm cell fusing with egg forming diploid zygote and other sperm fusing with polar nuclei forming triploid endosperm nucleus. Fruit set initiation begins immediately following successful fertilization with developing embryo and endosperm producing hormones particularly auxin and gibberellins triggering berry development, preventing flower abscission that would otherwise occur in unfertilized flowers, and initiating berry expansion from 2-3 mm ovary toward mature berry dimensions. Resource allocation during optimal window emphasizes reproductive demands with substantial carbohydrate consumption supporting pollen production, tube growth, and early embryo development, nitrogen requirements for protein synthesis in rapidly developing reproductive structures, and critical micronutrients particularly boron essential for pollen germination and tube growth with deficiency severely limiting pollination success. Hormonal coordination during optimal window involves declining growth promoters as flowering culminates, active auxin production from successfully fertilized ovaries triggering fruit development, possible ethylene involvement in unfertilized flower senescence, and integrated hormonal signals coordinating transition from flowering to fruit development. Developmental outcomes determined during optimal window include majority of final berry number 60-80% established through successful pollinations during this critical 2-3 day period, with remaining 20-40% set during suboptimal early and late anthesis periods, making optimal window conditions dominant influence on seasonal yield. Weather sensitivity during optimal window reaches absolute maximum with reproductive success extremely vulnerable to adverse conditions, minor weather perturbations potentially reducing fruit set by 10-30%, and severe events like rain reducing set by 50-80%, making weather during optimal window single most important determinant of seasonal productivity. Climate variability implications show that interannual variation in optimal window weather conditions creates substantial yield variation among seasons, favorable years with stable warm dry conditions during optimal window producing high set 60-75% and abundant yields, while unfavorable years with cool wet variable conditions producing poor set 20-40% and reduced yields, demonstrating critical role of flowering weather in determining vintage characteristics. Management implications during optimal window include maintaining minimal vineyard disturbance with all canopy operations suspended to avoid physical disruption of pollination, disease management restricted due to fungicide application prohibitions during bloom requiring protective applications completed before optimal window, weather monitoring intensified with forecasts critically important for anticipating potential problems, and possible protective interventions considered in high-value situations if adverse weather threatens though options are limited. The process represents third and culminating peak anthesis aspect integrating male and female reproductive capacities creating optimal reproductive opportunity, encompassing maximum flower opening and peak pollen availability phases. Duration typically spans 2-3 days representing brief critical period determining majority of fruit set and yield, with warm stable conditions producing compressed 2-day window while variable conditions may extend to 3-4 days. Cultivar variation appears in optimal window timing with early varieties showing late May-early June window while late types show mid-late June timing, window duration with some varieties showing brief 2-day window while others show extended 3-4 day period, and sensitivity to environmental conditions with some varieties showing robust reproduction under variable conditions while others show high sensitivity requiring optimal conditions for adequate set. Regional timing shows optimal pollination window during early-mid June in warm climates at approximately 1050-1100 growing degree days base 10°C, mid-June in temperate regions at 950-1000 GDD, and mid-late June in cool areas at 850-900 GDD, with timing ideally coinciding with stable favorable weather period. Viticultural significance includes representing most critical period of entire growing season with outcomes during these 2-3 days determining 60-80% final yield, requiring optimal environmental conditions for successful pollination and high fruit set, emphasizing importance of site selection, vineyard design, and canopy management optimizing flowering conditions, demonstrating extreme weather vulnerability necessitating risk management strategies, and illustrating fundamental challenge in viticulture of depending on brief weather-sensitive reproductive period for economic productivity. Research applications include analyzing temporal dynamics of reproductive capacity determining optimal window characteristics, quantifying environmental effects on pollination efficiency during optimal period, evaluating management practices affecting flowering conditions and fruit set outcomes, developing models predicting fruit set from optimal window weather conditions, assessing climate change impacts on optimal window timing and conditions affecting viticultural productivity, and investigating mechanisms of reproductive synchronization coordinating male and female functions during optimal window.
Optimal_Pollination_Window is third and culminating Peak_Anthesis aspect, distinguished by integration of maximum male and female reproductive capacities creating ideal pollination conditions. Encompasses Maximum_Flower_Opening and Peak_Pollen_Availability phases. Represents most critical 2-3 day period determining 60-80% seasonal yield. Environmental conditions particularly weather during optimal window fundamentally determine fruit set and productivity. Weather sensitivity absolute maximum making this single most vulnerable period of entire growing season. Outcomes during optimal window dominant influence on vintage characteristics.
Hauptbestäubungsperiode
ideal reproductive window
ideale Blütephase
optimal flowering period
optimale Befruchtungszeit
optimales Bestäubungsfenster
peak pollination opportunity
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 378-390 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 290-298 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Optimal Pollination Window
In a Willamette Valley Pinot Noir vineyard, optimal sugar transport was characterized over 13 days in early September Year N+1 (BBCH 85) when source-to-sink efficiency calculated 79% (based on canopy photosynthesis 14.2 g CO₂ per vine per day, berry hexose accumulation 8.7 g glucose-equivalent per vine per day, respiratory losses 2.3 g), berry Brix increase 4.8% per week, hexose import 118 μmol g⁻¹ h⁻¹, cell wall invertase 95 units/g FW, VvHT expression 128-fold, under ideal Oregon conditions (daily high 24-27°C, cool nights 13-16°C, adequate irrigation, healthy canopy).
Optimal_Sugar_Transport represents the coordinated efficient operation of all sugar delivery and uptake components during peak accumulation when source photosynthesis, phloem translocation, invertase cleavage, membrane transport, and vacuolar sequestration function in integrated synchrony creating maximum flux with minimal metabolic waste enabling most efficient sweetening achieving highest sugar gain per unit photosynthate invested. This optimal transport occurs during peak accumulation phase (BBCH 84-85), typically days 18-30 post-veraison spanning 10-16 days, and is characterized by source-to-sink carbon transfer efficiency 75-88% (proportion of fixed CO₂ reaching berry vacuoles as hexoses) compared to active phase typical 65-80% and non-optimal conditions 90% through apoplastic pathway (compared to 32°C beginning impairment through enzyme denaturation and membrane perturbation, and temperature stability over optimal period important as fluctuations require metabolic readjustment potentially disrupting coordination temporarily reducing efficiency. Water relations supporting optimal require adequate vine water status stem water potential -0.6 to -1.2 MPa maintaining phloem turgor and transport function, moderate deficit -1.0 to -1.3 MPa potentially compatible or enhancing concentration without severely impairing flux, severe deficit <-1.5 MPa disrupting optimal through impaired phloem delivery and metabolic limitation reducing efficiency 35-65%, and irrigation timing strategically maintaining adequate hydration throughout optimal period maximizing efficiency. Nutritional influences on optimal show adequate nitrogen supporting enzyme synthesis with N-deficient vines showing 25-45% reduced transport capacity through translational limitation, potassium important for membrane function and pH regulation with adequate K⁺ essential for optimal transport though excess potentially problematic, phosphorus required for ATP synthesis supporting active transport with P limitation potentially impairing 20-40%, and general nutrient balance ensuring healthy vine function supporting optimal metabolism. Cultivar variation in achieving optimal demonstrates some varieties reliably reaching optimal efficiency under favorable conditions (Chardonnay, Merlot high-performing), others showing good but not maximum efficiency (Cabernet Sauvignon moderate), and some rarely achieving true optimal even under favorable conditions (Pinot Noir particularly warm climates) reflecting genetic differences in pathway coordination and metabolic integration. Environmental stability supporting optimal includes consistent moderate weather over 10-16 day period without extreme fluctuations enabling maintained coordination, adequate radiation 20-30 MJ m⁻² per day saturating photosynthesis without excess heat stress, moderate temperatures 24-30°C day and 16-22°C night providing optimal thermal regime, absence of severe weather events preventing physical damage or metabolic disruption, and pest/disease control maintaining healthy fruit capable of optimal function. Quality implications of optimal transport include achieving maximum final sugars within growing season through superior efficiency, balanced composition as optimal transport typically coordinates with appropriate acid decline and phenolic development, uniformity across vineyard where all zones achieve optimal producing consistent maturity enabling efficient harvest, and reduced environmental footprint as optimal efficiency maximizes yield per unit resource (water, nutrients, pesticides) invested representing sustainable production. Viticultural strategies promoting optimal include crop load management (0.8-1.2 kg fruit per m² leaf area) ensuring source capacity adequate supporting optimal flux, canopy management maintaining healthy productive foliage throughout optimal period, irrigation scheduling providing consistent adequate water without excess, nutritional management ensuring adequate but not excessive nutrient availability, disease/pest control preventing disruptions, and environmental monitoring identifying optimal conditions enabling harvest timing decisions capturing fruit at peak efficiency. Research applications include flux analysis using isotope tracers quantifying pathway efficiency, transporter activity assays measuring component capacities, transcriptomic profiling revealing coordinated gene expression, metabolomic analysis tracking pathway intermediates, and systems biology modeling integrating data predicting optimal conditions enabling precision viticulture targeting maximum efficiency.
Optimal_Sugar_Transport is distinguished from Maximum_Sugar_Loading_Rate by emphasizing integrated efficiency (75-88% source-to-sink transfer) rather than absolute rate, from Active_Sugar_Loading by representing peak coordination rather than sustained operation, and from other transport classes by focusing on system-wide optimization rather than individual components. Duration 10-16 days BBCH 84-85. Temperature 26-30°C optimal. Metabolic efficiency maximized. Optimal_Sugar_Transport represents ideal integrated system performance determining maximum achievable sweetening efficiency.
coordinated sugar flux
integrated hexose delivery
integrierte Hexose-Lieferung
koordinierter Zucker-Fluss
maximale Transport-Effizienz
maximum transport efficiency
optimierter Akkumulations-Pfad
optimized accumulation pathway
Keller M (2020) The Science of Grapevines 3rd Ed pages 156-162, 223-228 integrated transport Ollat N, Gaudillère JP (1998) The effect of limiting leaf area during stage I of berry growth on development and composition of berries of Vitis vinifera L. cv. Cabernet Sauvignon Amer J Enol Vitic 49:251-258 source-sink efficiency Patrick JW (1997) Phloem unloading:sieve element unloading and post-sieve element transport Annu Rev Plant Physiol Plant Mol Biol 48:191-222 transport integration.
Optimal Sugar Transport
Cabernet Sauvignon optimally ripe berries for premium Napa Valley Cabernet production achieve precise target composition= 24.5% sugars (targeting 14% final alcohol), 6.3 g/L titratable acidity (pH 3.72), 3.6 mg/g skin anthocyanins, with sensory evaluation confirming ripe blackcurrant and cedar flavors, soft ripe tannins, fully mature brown seeds with nutty character, no green notes, representing the ideal harvest point balancing ripeness with structure and achieving convergence of analytical and sensory maturity indicators at week 18.5 post-fruit set. Sauvignon Blanc optimally ripe berries for crisp aromatic white wine achieve 21.8% sugars (targeting 13% alcohol maintaining varietal character), 8.2 g/L acidity (pH 3.28), creating optimal acid-sugar balance supporting characteristic grapefruit and gooseberry aromatics, with harvest window at week 14-14.5 representing narrow 5-7 day optimal period before sugars increase and acids decline excessively. Pinot Noir optimally ripe berries for Burgundian-style wine show precise convergence= 24.2% sugars, 6.8 g/L acidity (ratio 3.6=1), 2.9 mg/g anthocyanins, with berry tasting showing characteristic red cherry and strawberry fruit, earthy complexity, silky tannins from ripe skins, mature seeds contributing structure without bitterness, representing the ideal expression of Pinot Noir terroir achieved through harvest at precise optimal ripeness window (week 17.5-18, 3-5 day window). Syrah optimally ripe berry tracking across three consecutive vintages shows vintage variation in optimal timing= cool vintage achieving optimal ripeness at week 19 (23.8% sugars, 6.5 g/L acidity), moderate vintage at week 18 (24.5% sugars, 6.2 g/L acidity), warm vintage at week 17 (25% sugars, 6 g/L acidity), demonstrating that optimal ripeness represents specific compositional targets achieved at vintage-dependent timing rather than calendar date, requiring annual assessment to identify optimal harvest moment. Merlot comparison of harvest at under-ripe (week 16= 22% sugars, 8 g/L acidity, green herbal flavors, astringent tannins), optimally ripe (week 17.5= 24.5% sugars, 6.2 g/L acidity, ripe plum flavors, soft tannins), and overripe (week 19= 27% sugars, 4.8 g/L acidity, cooked fruit flavors, flabby structure) demonstrates quality advantages of harvesting at optimal ripeness with balanced composition producing highest quality wine versus early or late harvest compromising quality through compositional imbalances.
An optimally ripe berry is a fully ripe berry achieving the ideal compositional balance and quality characteristics for the specific intended use, representing the precise point or narrow window when all maturity indicators align at target levels creating maximum quality potential. Optimally ripe berries are characterized by= (1) sugar levels precisely at or very close to target for intended use (±0.5% from target), representing ideal fermentable sugar content for target alcohol level in wine or ideal sweetness for table consumption (2) acidity precisely at or very close to target levels (±0.3 g/L from target) providing optimal balance with sugars and optimal pH for stability and quality (3) sugar=acid ratio at ideal balance typically 3.5-4.5 for premium balanced wines, creating harmony between ripeness and freshness (4) phenolic maturity (red cultivars) precisely optimal with anthocyanins at peak extractability, tannins at ideal ripeness showing soft character without astringency, ideal color intensity and stability (5) flavor profile at peak with characteristic varietal aromas and flavors fully expressed, no under-ripe herbaceous notes, no overripe cooked or raisined notes, optimal balance of primary fruit character (6) seeds at ideal maturity showing brown color, appropriate lignification, flavor contribution enhancing rather than detracting from berry taste (7) all maturity indicators converging simultaneously creating the ideal harvest moment. Optimally ripe berries represent the target state for quality-focused viticulture= extensive vineyard monitoring during ripening to track compositional evolution sampling and analysis to identify when optimal ripeness is approaching sensory evaluation to confirm flavor maturity aligns with analytical maturity timely harvest execution to capture berries at optimal ripeness before progression to overripeness. Optimally ripe berries may represent a very narrow window= for some cultivars and styles, optimal ripeness may persist only 3-7 days before berries progress to overripeness for other cultivars, optimal ripeness window may extend 7-14 days providing more harvest flexibility environmental conditions (temperature, rainfall) during final maturation influence optimal ripeness window duration. Optimally ripe berries for different production goals show varying target compositions= premium red wine optimal ripeness may target 24-25% sugars, 6-6.5 g/L acidity, 3.5-4 mg/g anthocyanins white wine optimal ripeness may target 21-22% sugars, 7.5-8.5 g/L acidity emphasizing acid retention table grape optimal ripeness may target 19-20% sugars, 5-6 g/L acidity, specific firmness for shipping. Optimally ripe berries reflect integration of quantitative analytical targets with qualitative sensory assessment= analytical measurements (sugars, acids, pH, anthocyanins) indicate compositional readiness sensory evaluation (berry tasting, seed chewing, skin assessment) confirms flavor maturity and phenolic ripeness convergence of analytical and sensory maturity defines optimal ripeness. The optimal ripeness concept emphasizes that maturity is use-specific and style-specific rather than absolute= early-harvest sparkling wine base optimal ripeness (18-20% sugars, 9-11 g/L acidity) differs fundamentally from late-harvest dessert wine optimal ripeness (28-35+% sugars, 6-9 g/L acidity), requiring producers to define optimal ripeness targets for their specific production goals.
Optimally_Ripe_Berry represents the ideal target state within Fully_Ripe_Berry and is distinguished from Peak_Ripeness_Berry which may emphasize maximum ripeness versus ideal balance. Optimally ripe berries represent the harvest target for quality-focused production, requiring precise timing and assessment to capture. Understanding optimally ripe berry characteristics is essential for defining harvest criteria, for quality optimization, and for training vineyard teams in maturity assessment combining analytical and sensory evaluation to identify the optimal harvest moment.
Ideal-Reife-Traube
ideal reife Traube
ideally ripe berry
optimal reife Beere
target-maturity berry
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 235-245 Jackson, D.I., Lombard, P.B. (1993). Environmental and management practices affecting grape composition. American Journal of Enology and Viticulture 44=409-430.
Optimally Ripe Berry
Rkatsiteli bears an Orange_Wine_Scion_Role as Georgia's signature variety for qvevri-fermented amber wines, with 5-6 months skin contact in buried clay vessels producing wines of deep amber color, dried apricot and walnut character, and firm tannins (300-500 mg/L phenolics) that define the Kakhetian tradition recognized by UNESCO. Ribolla Gialla serves in an Orange_Wine_Scion_Role for Friulian producers (Gravner, Radikon, Princic) who pioneered Italian orange wine, with extended maceration (3-6 months) yielding wines of remarkable complexity, amber hue, and savory, umami depth. Pinot Grigio (Pinot Gris) bears an Orange_Wine_Scion_Role exploiting its pink-grey skin pigmentation, with extended maceration producing distinctively copper-colored "ramato" style wines in Friuli and Slovenia with enhanced textural grip and aromatic complexity versus conventional Pinot Grigio.
A wine production scion role that inheres in a grafted scion cultivar by virtue of its designation for orange wine (amber wine/skin-contact white wine) production, where white or grey-skinned grapes undergo extended maceration with their skins, extracting phenolic compounds, color pigments, and textural elements typically absent from conventionally produced white wines. Orange wine scions are white grape varieties processed using red wine techniques—extended skin contact ranging from several days to several months, often in traditional vessels (Georgian qvevri, amphora, or neutral oak)—producing wines with amber to deep orange coloration (from oxidized flavonoids and xanthophylls), tannic structure (100-500 mg/L phenolics versus 200-500 mg/L for conventional whites), and distinctive oxidative, nutty, dried fruit character. The style represents revival of ancient winemaking practices, particularly from Georgia's 8,000-year qvevri tradition, and has gained international recognition through the natural wine movement. Suitable varieties possess sufficient phenolic precursors in skins and stems to yield interesting extraction without excessive bitterness or astringency.
Orange_Wine_Scion_Role captures a vinification-defined category where white varieties are processed as if red, fundamentally altering wine character through phenolic extraction. The style exists on a continuum from brief skin contact (24-72 hours, subtle effect) through extended maceration (weeks to months, transformative). Georgian qvevri wines represent the ancestral tradition Italian and Slovenian producers (particularly around Gorizia/Nova Gorica) developed contemporary orange wine in the 1990s-2000s, inspiring global adoption. Variety selection significantly impacts results—thick-skinned varieties with concentrated phenolics (Ribolla Gialla, Malvasia Istriana, Friulano) produce more structured wines than thin-skinned aromatics. The "orange" designation (coined circa 2004) has become controversial in some markets, with "amber wine" or "skin-contact white" preferred.
Amphorenweinsorte
Bernsteinweinsorte
Orangeweinsorte
amber wine variety
macerated white variety
mazerierte Weißweinsorte
orange wine variety
skin-contact white variety
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Woolf S (2018) Amber Revolution: How the World Learned to Love Orange Wine, Interlink Books Feiring A (2016) For the Love of Wine: My Odyssey through the World's Most Ancient Wine Culture, Potomac Books Legeron I (2014) Natural Wine, CICO Books
Orange Wine Scion Role
Organ synchrony assessment in spring-stressed Merlot reveals the consequences of root-shoot developmental misalignment, with early warm air temperatures triggering rapid shoot growth (8 cm/day during week 3 post-budbreak) while cool soil temperatures delayed root activity (new root emergence only beginning week 4), creating a 10-day period of shoot growth dependent entirely on reserve mobilization without root uptake support, resulting in nutrient deficiency symptoms (interveinal chlorosis from inadequate iron uptake) and reduced shoot vigor once reserves depleted, demonstrating how organ asynchrony between roots and shoots creates developmental stress affecting vine establishment and early-season function.
Organ synchrony quality is a synchronization quality that represents the temporal coordination of developmental events among different organ types within the grapevine system, characterizing the alignment between the developmental trajectories of roots, shoots, leaves, inflorescences, and fruit that determines whether organs develop in complementary phases supporting whole-plant function or in competing phases creating resource conflicts and developmental imbalances. Within the BFO framework, this quality specifically addresses the inter-organ dimension of developmental coordination, distinguishing synchrony among different organ types from synchrony among organs of the same type while recognizing that organ synchrony affects the source-sink relationships, resource allocation patterns, and functional integration that determine vine productivity and fruit quality. The ontological significance lies in organ synchrony determining the coordination of complementary functions, where synchronized development ensures that photosynthetic source capacity develops appropriately to support reproductive sink demand, that root water and nutrient acquisition capacity matches shoot transpiration and growth requirements, and that the timing of organ-specific processes aligns with whole-plant developmental programs rather than competing for resources at critical periods. The developmental patterns requiring organ coordination include the spring flush where root carbohydrate mobilization must precede shoot growth demand, the flowering-to-fruit-set transition where leaf area development must provide adequate photosynthate for fruit establishment, and the ripening period where continued root function must support berry maturation while shoot growth appropriately declines. The factors affecting organ synchrony include environmental signals affecting different organs differently (soil temperature affecting roots, air temperature affecting shoots), resource availability patterns (water and nutrient supply affecting root-shoot coordination), and management practices that may differentially affect organ development (pruning affecting shoot development timing, irrigation affecting root activity patterns). Understanding organ synchrony quality enables assessment of whole-plant developmental coordination, identification of organ-level asynchronies that may create resource conflicts or developmental imbalances, and management interventions that promote complementary organ development supporting vine productivity and fruit quality.
Organ_Synchrony_Quality serves as the parent class for specific organ coordination relationships (Berry-Cluster, Shoot, Root-Shoot). This quality addresses developmental alignment among different organ types affecting whole-plant function. The class supports integrated vine management through organ synchrony characterization.
Ganzpflanzenkoordination
Inter-Organ-Koordination
Organsynchronie
functional synchrony
funktionelle Synchronie
inter-organ coordination
organ developmental alignment
whole-plant coordination
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 59-144 Zapata C et al. (2004) Partitioning and mobilization of starch and N reserves in grapevine (Vitis vinifera L.). Journal of Plant Physiology 161(9): 1031-1040
Organ Synchrony Quality
Organic acid quality profiling in Chablis Chardonnay reveals the characteristic tartaric-dominant profile (4.0-5.5 g/L tartaric, 2.5-4.0 g/L malic) that distinguishes cool-climate Burgundy from warmer regions, with the high malic acid content providing the raw material for malolactic fermentation that transforms the acid profile during barrel aging traditional Chablis producers block malolactic fermentation to preserve the steely, mineral-like acidity associated with preserved malic acid, while some modern producers encourage complete malolactic conversion for softer, more approachable wines, demonstrating how organic acid composition enables stylistic differentiation.
Organic_Acid_Quality encompasses the chemical qualities characterizing the concentration of individual organic acid species in grape berry tissue, providing compositional detail beyond aggregate acidity measurements. This category addresses the specific acids that constitute the titratable acidity, each with distinct metabolic origins, chemical properties, and contributions to wine character. The category includes the two dominant grape acids, Tartaric_Acid_Content_Quality and Malic_Acid_Content_Quality, which together typically comprise more than 90% of berry acidity, along with minor acids including Citric_Acid_Content_Quality, Succinic_Acid_Content_Quality, and Lactic_Acid_Content_Quality. Each organic acid contributes differently to taste perception: tartaric acid provides the hard, sharp acidity characteristic of wine malic acid contributes green, tart, apple-like sourness citric acid adds fresh, citrus-like notes succinic acid contributes complex, slightly bitter-salty character and lactic acid provides soft, milky sourness. The ratio among acids affects both taste quality and wine stability, with tartaric-dominant profiles providing stable acidity while malic-dominant profiles may undergo malolactic fermentation that fundamentally transforms acid composition. Organic acid profiles vary substantially among cultivars, with some varieties (Riesling, Barbera) retaining high acidity while others (Gewürztraminer, Viognier) naturally produce low-acid fruit. Climate profoundly influences acid composition through temperature effects on malic acid respiration, creating characteristic regional signatures. Individual acid measurement employs enzymatic assays, high-performance liquid chromatography, or capillary electrophoresis, providing the detailed compositional data needed for precise quality assessment and winemaking planning.
Organic_Acid_Quality serves as the parent class for individual acid species, positioned parallel to Total_Acidity_Quality within Acid_Content_Quality. The organic acids of grape and wine are all relatively weak acids with pKa values above 2.5, meaning they are only partially dissociated at wine pH and contribute to buffer capacity. The distinction between grape-derived acids (tartaric, malic, citric) and fermentation-derived acids (succinic, lactic, acetic) reflects different metabolic origins and compositional dynamics. Minor organic acids not explicitly addressed in child classes (acetic, fumaric, galacturonic, gluconic, oxalic) may be relevant in specific quality contexts.
Einzelsäuregehalt
Säurespezies
acid profile
acid species composition
individual acid content
organische Säurequalität
specific acid quality
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 295-315 Volschenk H et al. (2006) Malic Acid in Wine: Origin, Function and Metabolism during Vinification. South African Journal of Enology and Viticulture 27(2): 123-136
Organic Acid Quality
The organic amendment protocol for a biodynamically managed Grüner Veltliner vineyard in the Wachau integrates five organic inputs — fresh grape marc from the estate's own pressing (3-4 t/ha applied in November to alternate inter-rows, providing approximately 15 kg N/ha, 3 kg P/ha, and 20 kg K/ha as the marc decomposes over 6-12 months), horn manure preparation 500 (25 g/ha in 35 L dynamized water applied to soil in March and October to stimulate soil microbial activity and humus formation), horn silica preparation 501 (4 g/ha in 35 L dynamized water applied as foliar spray at E-L 19 and E-L 33 to enhance light metabolism and fruit ripening), biochar (2 t/ha incorporated at 20 cm depth in newly planted vine rows only, providing stable carbon storage and enhanced cation exchange capacity in the sandy loess soils), and kelp extract (Ascophyllum nodosum at 5 L/ha in 3 foliar applications from E-L 15 to E-L 27, providing cytokinin and betaine plant growth regulators that enhance stress tolerance during the pre-véraison water deficit period).
A nutrition management description that documents the application of non-composted organic materials to vineyard soils for nutritional, soil structural, and biological purposes, specifying the amendment source (cover crop residue, grape marc, winery lees, biochar, seaweed extracts, biodynamic preparations), application rate, timing, incorporation method, and the specific nutritional or soil health objectives that distinguish organic amendment from composted material application.
Organic_Amendment_Protocol captures the diverse range of non-composted organic inputs used in viticulture that do not fall within the composting framework documented by Compost_Application_Rate. These inputs range from conventional recycling of winery byproducts (grape marc, lees, filter residues) through biodynamic preparations with specific philosophical and practical frameworks (preparations 500-508 as defined by Rudolf Steiner's 1924 Agricultural Course) to modern biotechnology products (biochar, microbial inoculants, seaweed extracts): the common principle is the addition of organic carbon and associated biological activity to the soil or vine system, but the mechanisms and objectives vary enormously — grape marc application primarily recycles nutrients removed in the harvest, biochar provides stable carbon storage and soil physical improvement, biodynamic preparations aim to enhance soil and plant vitality through specific fermented organic substrates, and seaweed extracts deliver plant growth regulators and stress-tolerance compounds. The distinction from composted amendments is functional — non-composted organic materials undergo decomposition in situ in the vineyard soil, with different nutrient release kinetics, C/N ratio implications, and soil biological effects than pre-composted materials that have already undergone thermophilic decomposition and stabilization.
Organische-Düngung-Protokoll
biological amendment specification
biologische Bodenverbesserung
non-composted organic application record
organic input protocol
organische Zusatzstoff-Spezifikation
White RE (2015) Understanding Vineyard Soils 2nd Ed, Oxford University Press pp: 165-185 Steiner R (1924) Geisteswissenschaftliche Grundlagen zum Gedeihen der Landwirtschaft (Agriculture Course), translated Creeger CE and Gardner M (1993), Biodynamic Farming and Gardening Association Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 445-460 Genesio L et al. (2015) Biochar application in vineyards — a new perspective for climate change mitigation, BIO Web of Conferences 5: 02003
Organic Amendment Protocol
The organic matter content of a Willamette Valley Pinot Noir vineyard (Oregon) documents moderate organic matter levels of 3.5-4.5% in the upper 30 cm of the Jory series volcanic soil (Xeric Palehumults) — this moderate organic matter content, maintained by annual cover crop incorporation and minimal tillage management, provides favorable soil physical conditions (stable granular structure, infiltration rate >50 mm/hour, plant-available water capacity enhanced by 20-30 mm compared to the same soil without organic matter enrichment), adequate nitrogen mineralization (releasing approximately 40-60 kg N/ha/year from organic matter decomposition, sufficient for moderate vine growth without supplemental nitrogen fertilization), and a biologically active soil environment (microbial biomass carbon of 300-500 mg/kg, supporting the diverse bacterial and fungal communities including mycorrhizal fungi that enhance vine nutrient acquisition). This moderate organic matter content contrasts with the low organic matter (0.5-1.5%) of hot, dry Mediterranean vineyard soils where rapid decomposition limits organic matter accumulation, and the high organic matter (5-8%) of cool, humid vineyard soils in northern Europe where decomposition is temperature-limited.
A soil chemistry data component that records the concentration of organic carbon compounds in the vineyard soil, expressed either as percent organic matter (by loss-on-ignition or Walkley-Black dichromate oxidation) or as percent organic carbon (with the conventional conversion factor of organic matter = organic carbon × 1.724), with organic matter content influencing soil structure stability, water-holding capacity, cation exchange capacity, nitrogen mineralization rate, biological activity, and the overall physical and chemical fertility of the vineyard soil.
Organic_Matter_Content documents the soil property that integrates biological, chemical, and physical soil functions — organic matter is simultaneously a source of plant nutrients (particularly nitrogen, phosphorus, and sulfur released through microbial decomposition), a contributor to cation exchange capacity (providing 20-70% of the total CEC in surface horizons through the carboxyl and phenolic functional groups on humic substances), a soil structure stabilizer (binding mineral particles into water-stable aggregates through organo-mineral complexes), a water retention enhancer (organic matter absorbs 3-5 times its weight in water, increasing plant-available water capacity by 1-2 mm per 0.1% increase in organic matter), and the energy source driving soil biological activity (supporting the decomposer food web from bacteria and fungi through protozoa, nematodes, and earthworms to macro-arthropods): the loss of organic matter through intensive cultivation is therefore simultaneously a loss of soil fertility, structure, water retention, and biological function. For vineyard soils, organic matter management involves the balance between organic matter inputs (cover crop biomass, compost applications, pruning debris, organic mulches) and organic matter losses (decomposition accelerated by tillage, which aerates the soil and exposes protected organic matter to microbial attack, and by warm temperatures that increase decomposition rates) — the trend toward reduced tillage, permanent cover crops, and compost application in progressive viticulture reflects the recognition that maintaining soil organic matter is essential for long-term soil health and sustained vineyard productivity.
Boden-Kohlenstoffgehalt
Humusgehalt
Organische-Substanz-Gehalt
humus content measurement
organic matter assessment
soil organic carbon record
White RE (2003) Soils for Fine Wines, Oxford University Press Morlat R and Jacquet A (2003) Grapevine root system and soil characteristics in a vineyard maintained long-term with or without interrow sward, American Journal of Enology and Viticulture 54: 1-7 Soil Science Division Staff (2017) Soil Survey Manual, USDA Handbook 18
Organic Matter Content
Sangiovese vines at a Chianti Classico estate bearing an organic production role are managed under EU Regulation 2018/848 with copper-based fungicide applications limited to 4 kg Cu/ha/year averaged over 5 years, sulfur dust applications of 20-30 kg/ha per treatment for powdery mildew control, and Bacillus thuringiensis for Lobesia botrana management, with cover crops of Trifolium repens providing nitrogen fixation at 40-80 kg N/ha/year.
A production system role that inheres in a grapevine cultivated under a certified organic management framework that prohibits synthetic chemical fertilizers, synthetic pesticides, and genetically modified organisms while permitting biological control agents, plant-derived preparations, and mineral inputs such as copper and sulfur within regulated annual limits.
Organic_Production_Role requires adherence to certification standards defined by regulatory bodies such as the EU (Regulation 2018/848), USDA NOP, or equivalent national organic programs. The transition from conventional to organic status typically requires a 3-year conversion period during which the vine bears a Transitional_Organic_Role. Subclasses distinguish between fully certified organic status and transitional status: the role hierarchy reflects the regulatory pathway toward organic certification.
Bio-Produktionsrolle
ecological production role
organic farming role
organic viticulture role
Ökoweinbau-Rolle
ökologische Anbau-Rolle
EU Regulation 2018/848 on organic production and labelling USDA National Organic Program 7 CFR Part 205 Willer H and Lernoud J (2023) The World of Organic Agriculture, FiBL-IFOAM Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 548-556
Organic Production Role
Origin record for Cabernet Sauvignon documents a surprisingly recent and geographically specific origin — molecular parentage analysis (Bowers and Meredith 1997) established the cultivar as a spontaneous cross of Cabernet Franc × Sauvignon Blanc, most likely occurring in the 17th century in southwestern France (Bordeaux or surrounding areas), with no documented reference before the late 18th century despite the parents' much longer documented histories, subsequent global dissemination through French colonial networks to Chile (1850s), California (1880s), Australia (1860s), and South Africa (1920s), and late 20th-century expansion into virtually every wine-producing country. Origin record for Blaufränkisch traces the cultivar's Central European provenance — genetic analysis confirms close relationship with several Austrian and Hungarian cultivars, probable origin in the Lower Austrian-Burgenland-Western Hungarian border region, earliest documented cultivation in the Sopron and Burgenland areas, 19th-century expansion through the Austro-Hungarian Empire under multiple synonyms (Kékfrankos in Hungary, Frankovka in Czech Republic/Slovakia, Lemberger in Germany/USA), and the contemporary concentration of premium plantings in Austria's Mittelburgenland, Leithaberg, and Eisenberg DAC regions.
Origin record is a historical record entity that documents the geographic provenance, migration history, and dispersal pathways of grape cultivars, integrating evidence from molecular genetic parentage analysis, population genetics, archaeobotanical findings, archival documentation, and linguistic-toponymic analysis to reconstruct the spatial history of cultivar origin, domestication, dissemination, and establishment across viticultural regions, providing the biogeographic narrative that explains how specific cultivars came to occupy their current distribution patterns and the human agency — colonial expansion, monastic networks, trade routes, political decisions, and scientific exchange programs — that mediated cultivar migration across continents and millennia.
Origin record documentation addresses one of the most fascinating and methodologically complex questions in viticultural science — where did individual cultivars come from, and how did they reach their current global distribution? The molecular revolution in grapevine genetics since the 1990s has fundamentally transformed origin studies, as microsatellite (SSR) genotyping and whole-genome sequencing have enabled parentage analysis, population structure assessment, and phylogeographic reconstruction at a level of precision impossible through morphological ampelography alone. The central finding of molecular origin studies is that the vast majority of premium wine grape cultivars originated through spontaneous crosses in the vineyard rather than through deliberate breeding — Cabernet Sauvignon, Chardonnay, Syrah, Merlot, and virtually all other traditional European varieties are accidental seedlings selected and propagated by observant vignerons who recognized their quality potential. The geographic center of Vitis vinifera domestication has been localized through archaeobotanical and genetic evidence to the South Caucasus region (present-day Georgia, Armenia, Azerbaijan) approximately 6,000-8,000 years ago, with subsequent westward diffusion through Anatolia, the eastern Mediterranean, and eventually to Western Europe — though recent evidence suggests possible independent or parallel domestication events in the western Mediterranean. The human-mediated dispersal of cultivars involves multiple mechanisms operating at different temporal and spatial scales — local selection and exchange among neighboring communities, regional dissemination through monastic networks (particularly Cistercian and Benedictine orders in medieval Europe), long-distance transport through colonial expansion (Spanish, Portuguese, French, and British colonial viticulture), and modern organized scientific exchange programs. The naming and renaming of cultivars during migration creates substantial historical confusion — a single cultivar may carry dozens of regional synonyms (Blaufränkisch/Kékfrankos/Frankovka/Lemberger), while unrelated cultivars may share names across regions, requiring molecular verification to resolve historical identity claims.
Herkunftsnachweis
Migrationsgeschichte
Sortenherkunft
Ursprungsaufzeichnung
Verbreitungsgeschichte
cultivar origin
dispersal history
migration record
provenance record
Bowers JE, Meredith CP (1997) The parentage of a classic wine grape, Cabernet Sauvignon. Nature Genetics 16(1): 84-87 This P et al. (2006) Historical origins and genetic diversity of wine grapes. Trends in Genetics 22(9): 511-519 Myles S et al. (2011) Genetic structure and domestication history of the grape. Proceedings of the National Academy of Sciences 108(9): 3530-3535 McGovern PE (2003) Ancient Wine, Princeton University Press, pp.: 1-120
Origin Record
Orthonasal aroma intensity comparison of Gewürztraminer in standard tasting glass versus narrow flute reveals intensity ratings of 8.2 versus 5.8 on a 10-point scale, with the 41% higher perceived intensity in the wider glass attributed to greater headspace volume concentrating volatiles and larger opening presenting more aroma to the nose, demonstrating glassware's critical influence on orthonasal perception and supporting the use of variety-specific glass designs to optimize aromatic presentation for distinctive varieties.
Orthonasal aroma intensity quality is an aroma intensity quality that represents the magnitude of olfactory sensation perceived through external nasal inhalation when sniffing wine in the glass, characterizing the aromatic strength experienced through the direct pathway from external environment through nostrils to olfactory epithelium. Within the BFO framework, this quality specifically addresses the intensity perceived via the orthonasal route, distinguishing external sniffing perception from retronasal perception during consumption while recognizing that orthonasal evaluation provides the traditional first assessment of wine aroma preceding tasting. The ontological significance lies in orthonasal intensity determining the initial aromatic impression that shapes expectations before wine contacts the palate, where powerful orthonasal aroma creates anticipation and immediate varietal or quality recognition, while weak orthonasal presence may indicate closed wine requiring decanting, neutral variety, or quality deficiency. The physiological basis for orthonasal perception involves volatile molecules carried by inspired airflow directly to the olfactory cleft in the nasal cavity roof, with perception affected by sniffing behavior (vigor, duration, frequency), wine temperature (controlling headspace concentration), glass geometry (affecting headspace accumulation and presentation to nose), and individual variation in olfactory sensitivity. The compositional determinants of orthonasal intensity include the volatility and concentration of aroma compounds, with highly volatile compounds (esters, terpenes) contributing disproportionately to orthonasal perception compared to less volatile compounds that require the warming and agitation of in-mouth conditions for release. The evaluation protocol for orthonasal intensity typically involves swirling to release volatiles, positioning nose at glass rim, and inhaling with controlled vigor, with standardization of these parameters essential for reproducible assessment. The relationship between orthonasal and retronasal intensity is compound-specific, with some aromatic compounds showing similar intensity by both routes while others differ substantially due to differential release conditions and potentially different neural processing. Understanding orthonasal aroma intensity enables assessment of wine's immediate aromatic impact, optimization of serving protocol to maximize orthonasal expression, and prediction of the first sensory impression shaping overall quality perception.
Orthonasal_Aroma_Intensity_Quality specifically addresses the sniffing route of aroma perception, complementing Retronasal_Aroma_Intensity_Quality for in-mouth perception. This quality represents the traditional wine evaluation sequence where orthonasal assessment precedes tasting. Evaluation protocol (swirling, glass position, sniff technique) significantly affects perceived intensity and should be standardized for reliable assessment.
Schnüffelintensität
direct olfactory intensity
direkte olfaktorische Stärke
external aroma strength
externe Geruchsintensität
orthonasale Aromaintensität
sniffing intensity
Hummel T et al. (2006) Effects of olfactory function, age, and gender on trigeminally mediated sensations: a study based on the lateralization of chemosensory stimuli. Toxicology Letters 140-141: 273-280 Delwiche J (2004) The impact of perceptual interactions on perceived flavor. Food Quality and Preference 15(2): 137-146
Orthonasal Aroma Intensity Quality
Osmolyte accumulation quality profiling in drought-stressed Tempranillo vines characterized the metabolic response underlying this cultivar's reputation for moderate drought tolerance in Mediterranean viticulture. Progressive soil drying induced coordinated osmolyte accumulation quality responses with proline increasing 8-fold from 2 to 16 μmol g⁻¹ dry weight, total soluble sugars increasing 2.5-fold from 40 to 100 mg g⁻¹, and malate increasing 1.8-fold over the stress period, collectively achieving osmotic potential reduction of 0.6 MPa that maintained positive turgor through moderate stress. The temporal pattern of osmolyte accumulation quality showed proline responding earliest within 3 days of stress onset, followed by sugar accumulation peaking at 7 days, with malate showing the slowest response requiring 10-14 days to reach maximum, suggesting sequential deployment of different osmolyte classes as stress intensified. Comparison with the drought-sensitive cultivar Albariño revealed substantially lower osmolyte accumulation quality with only 3-fold proline increase and 1.5-fold sugar increase under equivalent stress, achieving only 0.25 MPa osmotic adjustment and exhibiting earlier turgor loss and wilting symptoms, demonstrating the functional importance of osmolyte accumulation quality for drought tolerance.
Osmolyte accumulation quality is an osmotic adjustment quality that inheres in a grapevine organism and represents the overall capacity to synthesize and accumulate the diverse array of compatible solutes that collectively contribute to osmotic adjustment under water deficit stress, encompassing the coordinated regulation of multiple metabolic pathways producing proline, soluble sugars, organic acids, polyols, and other osmotically active compounds that lower cellular osmotic potential while maintaining metabolic compatibility and providing additional protective functions beyond simple osmotic effects. This quality captures the integrative capacity for osmolyte production that determines the magnitude of achievable osmotic adjustment, recognizing that effective adjustment typically involves multiple osmolyte classes whose combined accumulation achieves the osmotic potential changes required for turgor maintenance under stress. The ontological significance of osmolyte accumulation quality lies in its role as the metabolic foundation for osmotic adjustment capacity, with the ability to rapidly synthesize and accumulate diverse compatible solutes determining whether vines can achieve the substantial osmotic potential reductions required for turgor maintenance under moderate-to-severe water deficit. The biological basis of osmolyte accumulation encompasses the stress-responsive activation of biosynthetic pathways for various osmolyte classes, the regulation of membrane transporters controlling osmolyte compartmentalization, and the metabolic flux adjustments that redirect carbon and nitrogen toward osmolyte synthesis under stress conditions when growth is limited. The diversity of accumulated osmolytes provides functional advantages beyond osmotic adjustment, with proline contributing to protein stabilization and reactive oxygen scavenging, sugars providing membrane protection and carbon reserves, and organic acids contributing to pH regulation, creating a multifunctional stress response through osmolyte accumulation. Cultivar and rootstock differences in osmolyte accumulation quality reflect genetic variation in the regulatory and biosynthetic machinery controlling osmolyte production, providing targets for selection and breeding programs aimed at improving drought tolerance through enhanced osmotic adjustment capacity.
Osmolyte_Accumulation_Quality is_a Osmotic_Adjustment_Quality, representing the overall capacity for compatible solute production contributing to osmotic adjustment. This class addresses the integrative metabolic response while sibling classes (Proline_Accumulation_Quality, Soluble_Sugar_Accumulation_Quality) capture specific osmolyte contributions. The multifunctional benefits of osmolytes beyond osmotic effects represent important additional value. Temporal patterns of different osmolyte classes provide insight into stress response dynamics. Critical for understanding the metabolic basis of drought tolerance variation among genotypes.
Osmolyt-Akkumulation
Osmolyt-Synthesekapazität
Osmoprotektant-Produktion
compatible solute accumulation
kompatible Solut-Anreicherung
osmolyte synthesis capacity
osmoprotectant production
solute accumulation response
Patakas A et al. (2002) Leaf age effects on solute accumulation in water-stressed grapevines. Journal of Plant Physiology 159:969-976 Hochberg U et al. (2013) Metabolite profiling and network analysis reveal coordinated changes in grapevine water stress response. BMC Plant Biology 13:184 Cramer GR et al. (2007) Water and salinity stress in grapevines: early and late changes in transcript and metabolite profiles. Functional and Integrative Genomics 7:111-134
Osmolyte Accumulation Quality
Systematic quantification of Osmotic_Adjustment_Disposition at the University of Adelaide measured osmotic potential at full turgor (Ψπ100) in leaves of 25 cultivars under well-watered and drought-stressed (predawn Ψ = -0.8 MPa for 14 days) conditions. Osmotic adjustment was calculated as the difference between stressed and control Ψπ100 values. Additionally, tissue extracts were analyzed for major compatible solutes (proline, total soluble sugars, potassium) to identify the biochemical basis of adjustment. Osmotic_Adjustment_Disposition varied substantially among cultivars, ranging from 0.12 MPa ('Pinot Noir', minimal adjustment) to 0.72 MPa ('Sultana', strong adjustment). Intermediate values characterized most wine grape cultivars, with 'Cabernet Sauvignon' at 0.38 MPa and 'Chardonnay' at 0.45 MPa. Solute analysis revealed cultivar-specific adjustment strategies. 'Sultana' achieved its strong adjustment primarily through sugar accumulation (contributing 55% of osmotic change) with moderate proline contribution (25%). 'Shiraz' (0.52 MPa adjustment) relied more heavily on proline (45% contribution) with lower sugar involvement (30%). Potassium contributed 15-25% of adjustment across all cultivars. Turgor maintenance directly reflected Osmotic_Adjustment_Disposition, with high-adjusting 'Sultana' maintaining leaf turgor of 0.65 MPa at leaf water potential of -2.0 MPa, while low-adjusting 'Pinot Noir' showed turgor loss (zero turgor) at -1.6 MPa. Growth analysis confirmed functional significance, with high-adjusting cultivars maintaining 70% of control shoot elongation rate at moderate drought versus 35% in low-adjusting cultivars, demonstrating that Osmotic_Adjustment_Disposition enables continued growth under water deficit.
An Osmotic_Adjustment_Disposition is a Drought_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through the active accumulation of compatible solutes in cells during water stress, lowering osmotic potential and enabling water uptake and turgor maintenance at soil and leaf water potentials that would otherwise cause wilting and loss of cell function. This disposition represents the biochemical component of drought tolerance operating at the cellular level, complementing stomatal regulation that operates at the whole-plant level. Osmotic adjustment enables cells to maintain positive turgor pressure and associated turgor-dependent processes (cell expansion, stomatal function, phloem transport) under water deficit conditions by increasing the concentration of osmotically active solutes including inorganic ions (potassium), amino acids (proline, glycine betaine), sugars (glucose, fructose, sucrose), sugar alcohols (mannitol, sorbitol), and organic acids (malate). The magnitude of osmotic adjustment varies among grapevine genotypes from minimal ( 0.8 MPa), with high-adjusting cultivars able to maintain turgor and function at water potentials that cause wilting in low-adjusting cultivars. Osmotic_Adjustment_Disposition is quantified by comparing osmotic potential at full turgor in stressed versus well-watered plants, with the difference representing active solute accumulation. This capacity is metabolically costly (requiring carbon skeletons and energy for solute synthesis) but enables continued growth and function during drought when non-adjusting plants would be severely impaired.
Osmotic_Adjustment_Disposition is_a Drought_Tolerance_Disposition. Osmotic adjustment provides cellular-level drought tolerance through solute accumulation, enabling turgor maintenance at low water potential. Child classes address specific aspects including Solute_Accumulation_Disposition (biochemical mechanisms) and Turgor_Maintenance_Disposition (functional outcome). The magnitude and solute composition of adjustment vary among genotypes. Osmotic adjustment is metabolically costly but enables continued growth and function during drought. Table grape cultivars often show stronger adjustment than wine grapes.
Aktive-Solut-Akkumulation
Osmoregulation
Osmotische-Anpassungs-Disposition
active solute accumulation
osmotic regulation
turgor-maintaining solute adjustment
Morgan JM (1984) Osmoregulation and water stress in higher plants, Annual Review of Plant Physiology 35:299-319 Patakas A et al. (2002) Leaf age effects on solute accumulation in water-stressed grapevines, Journal of Plant Physiology 159:969-976 Rodrigues ML et al. (1993) Osmotic adjustment in water stressed grapevine leaves in relation to carbon assimilation, Australian Journal of Plant Physiology 20:309-321
Osmotic Adjustment Disposition
Osmotic adjustment quality comparison across rootstock genotypes in Central Valley California evaluated the contribution of this mechanism to drought tolerance differences affecting scion performance under deficit irrigation. Rootstock 110R demonstrated excellent osmotic adjustment quality with leaf osmotic potential declining from -1.2 MPa at full hydration to -2.0 MPa under moderate stress, an adjustment of 0.8 MPa that maintained positive turgor and stomatal function at soil water potentials causing severe stress symptoms in non-adjusting genotypes. Rootstock 101-14 showed limited osmotic adjustment quality with only 0.3 MPa adjustment under equivalent stress, resulting in earlier turgor loss and stomatal closure that limited carbon assimilation and scion growth. The osmotic adjustment quality differential translated directly to scion performance, with Chardonnay on 110R maintaining 45% higher photosynthesis under moderate deficit than equivalent scions on 101-14, establishing osmotic adjustment quality as a primary mechanism underlying rootstock drought tolerance differences. Subsequent metabolomic analysis revealed that 110R accumulated 3-fold higher proline and 2-fold higher soluble sugars than 101-14 under stress, identifying the biochemical basis of its superior osmotic adjustment quality.
Osmotic adjustment quality is a drought stress indicator quality that inheres in a grapevine organism and represents the capacity to actively accumulate solutes in cellular compartments in response to declining water availability, lowering the osmotic potential of cell contents to maintain positive turgor pressure and cellular function at water potentials that would otherwise cause turgor loss, wilting, and physiological impairment, constituting a primary drought tolerance mechanism enabling continued growth and metabolic activity under water deficit conditions. This quality captures one of the most important physiological adaptations to drought stress, with active solute accumulation maintaining the turgor-dependent processes of cell expansion, stomatal function, and membrane integrity at water potentials that would cause cellular collapse in non-adjusting tissues. The ontological significance of osmotic adjustment quality lies in its role as a fundamental drought tolerance mechanism distinguishing cultivars capable of sustained function under water deficit from those lacking this adaptive capacity, with substantial variation in adjustment magnitude among grapevine genotypes providing a physiological basis for drought tolerance differences observed at the whole-plant level. The biological basis of osmotic adjustment encompasses the synthesis and accumulation of compatible solutes including proline, soluble sugars, organic acids, and inorganic ions that lower cellular osmotic potential without disrupting metabolic function, combined with the compartmentalization of these solutes to maintain appropriate cytoplasmic conditions while achieving bulk osmotic adjustment primarily through vacuolar accumulation. The magnitude of osmotic adjustment achievable varies among cultivars from negligible in drought-sensitive types to substantial (0.5-1.0 MPa or more) in drought-tolerant genotypes, with this variation reflecting genetic differences in the sensing, signaling, and metabolic pathways governing solute accumulation under stress. The relationship between osmotic adjustment quality and whole-plant drought tolerance involves the maintenance of turgor-dependent processes including stomatal function, cell expansion, and root growth that collectively determine the capacity for continued water uptake, carbon assimilation, and growth under limiting water supply.
Osmotic_Adjustment_Quality is_a Drought_Stress_Indicator_Quality, serving as parent class for Osmolyte_Accumulation_Quality, Proline_Accumulation_Quality, and Soluble_Sugar_Accumulation_Quality which address specific aspects of the osmotic adjustment mechanism. This class addresses the overall capacity for turgor maintenance through solute accumulation while child classes capture particular osmolyte contributions. The magnitude of adjustment (in MPa) provides quantitative characterization with 0.5-1.0 MPa representing substantial adjustment capacity. Critical for understanding drought tolerance mechanisms and rootstock selection for water-limited conditions.
Turgorerhaltung
osmoregulation
osmotische Anpassung
osmotische Regulation
solute accumulation capacity
turgor maintenance
Patakas A et al. (2002) Leaf age effects on solute accumulation in water-stressed grapevines. Journal of Plant Physiology 159:969-976 Cramer GR et al. (2007) Water and salinity stress in grapevines: early and late changes in transcript and metabolite profiles. Functional and Integrative Genomics 7:111-134 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data. Annals of Botany 105:661-676
Osmotic Adjustment Quality
Osmotic potential monitoring in Riesling berries through ripening reveals decline from -1.2 MPa at véraison (12°Brix) to -2.8 MPa at harvest (21°Brix), with the 1.6 MPa decrease reflecting sugar accumulation from 120 to 210 g/L that generates the concentration gradient driving continued phloem import despite the near-zero turgor state of post-véraison berries, demonstrating how osmotic potential rather than turgor becomes the dominant driver of berry water relations during ripening.
Osmotic potential quality is a physical quality that inheres in grapevine cells and tissues and represents the component of water potential attributable to dissolved solute concentration, characterizing the thermodynamic tendency for water to move toward regions of higher solute concentration by osmosis across semipermeable membranes. Within the BFO framework, this quality captures the chemical potential reduction caused by solute presence, where more negative osmotic potential indicates higher solute concentration creating stronger driving force for water uptake into cells. The ontological significance lies in osmotic potential providing the driving force for turgor generation when cells are enclosed by walls, and determining the equilibrium water content achieved at any given external water potential through the relationship Ψ = Ψs + Ψp where water potential equals osmotic potential plus turgor pressure. Osmotic potential in grapevine tissues ranges from approximately -0.8 MPa in dilute xylem sap to -3.5 MPa in sugar-concentrated ripe berry mesocarp, with the developmental increase in osmotic solute concentration (primarily sugars, organic acids, and potassium) driving the massive water influx enabling berry growth. The osmotic potential of cell vacuoles, which comprise the majority of symplastic volume, determines the capacity to generate turgor and maintain cell hydration under water stress, with osmotic adjustment through active solute accumulation representing a key acclimation mechanism. Measurement approaches include thermocouple psychrometry of frozen-thawed tissue (eliminating turgor contribution), vapor pressure osmometry of expressed sap, and calculation from solute concentration using the van't Hoff relationship. The relationship between osmotic potential and sugar concentration provides the physiological basis for must density as a maturity indicator, where increasing osmotic potential reflects the sugar accumulation defining ripening progression. Understanding osmotic potential quality enables mechanistic interpretation of water relations, prediction of water movement patterns within the vine, and analysis of the solute accumulation underlying fruit composition development.
Osmotic_Potential_Quality addresses the solute-dependent component of water potential, complementing Turgor_Pressure_Quality as the pressure component in the water potential equation. This quality is always negative (or zero for pure water), with more negative values indicating higher solute concentration. The class supports interpretation of water movement, analysis of drought tolerance mechanisms through osmotic adjustment, and connection between berry composition (sugar content) and hydraulic properties.
Solutenpotential
osmotic pressure
osmotischer Druck
osmotisches Potential
osmotisches Wasserpotential
solute potential
Keller M et al. (2015) Ripening grape berries remain hydraulically connected to the shoot. Journal of Experimental Botany 66(9): 2683-2693 Tilbrook J, Tyerman SD (2008) Cell death in grape berries: varietal differences linked to xylem pressure and berry weight loss. Functional Plant Biology 35(3): 173-184
Osmotic Potential Quality
Detailed investigation of Osmotic_Stress_Tolerance_Disposition at INRAE Montpellier employed pressure-volume analysis and compatible solute profiling across 12 rootstocks under iso-osmotic salt (NaCl) versus non-ionic osmoticum (PEG-6000) treatments at -0.5 MPa osmotic potential. This experimental design separated osmotic from ionic stress effects, enabling direct assessment of Osmotic_Stress_Tolerance_Disposition. Pressure-volume parameters including osmotic potential at full turgor (Ψπ100), turgor loss point (Ψtlp), and bulk modulus of elasticity (ε) were determined, alongside tissue concentrations of proline, soluble sugars, and organic acids. Osmotic_Stress_Tolerance_Disposition showed substantial variation among rootstocks. '110 Richter' exhibited strong osmotic tolerance with Ψπ100 declining from -1.2 MPa (control) to -1.95 MPa under salt (0.75 MPa adjustment), maintaining positive turgor (0.35 MPa) at leaf water potential of -1.6 MPa. Compatible solute analysis revealed proline accumulation to 42 μmol/g FW (8-fold increase) and total soluble sugars increasing 2.5-fold, together accounting for 45% of osmotic adjustment, with vacuolar Na⁺ and Cl⁻ contributing the remainder. 'Riparia Gloire' showed weak Osmotic_Stress_Tolerance_Disposition with Ψπ100 declining only to -1.45 MPa (0.25 MPa adjustment), reaching turgor loss at leaf water potential of -1.1 MPa, and proline accumulation of only 12 μmol/g FW. Importantly, responses to NaCl and PEG were highly correlated across genotypes (r = 0.88 for osmotic adjustment), indicating that Osmotic_Stress_Tolerance_Disposition reflects inherent osmotic adjustment capacity rather than salt-specific adaptation. Strong Osmotic_Stress_Tolerance_Disposition maintained stomatal conductance at 65% of control under salinity versus only 30% in osmotic-sensitive genotypes, directly supporting continued carbon assimilation.
An Osmotic_Stress_Tolerance_Disposition is a Salt_Tolerance_Disposition that inheres in a Vitis plant and is realized through physiological mechanisms that enable maintenance of water uptake, turgor pressure, and cellular function despite the low soil water potential created by dissolved salts, addressing the water-deficit component of salt stress independent of specific ion toxicity. This disposition represents tolerance to the osmotic effect of salinity, which is immediate (occurring as soon as roots encounter saline solution) and affects all plants regardless of ion exclusion capacity. The osmotic component of salt stress is functionally equivalent to drought stress in its effects on plant water relations, reducing soil water availability and creating the need for plants to lower their own water potential to maintain the gradient driving water uptake. Osmotic_Stress_Tolerance_Disposition involves active osmotic adjustment through accumulation of compatible solutes (proline, glycine betaine, sugars, organic acids) and inorganic ions (K⁺) that lower cytoplasmic and vacuolar osmotic potential, enabling water uptake from low-potential soil while maintaining turgor for growth and stomatal function. Sequestered Na⁺ and Cl⁻ can contribute to osmotic adjustment if effectively compartmentalized in vacuoles, providing a metabolic benefit from what would otherwise be purely detrimental ion accumulation. Genotypes with strong Osmotic_Stress_Tolerance_Disposition maintain higher water content, turgor, and stomatal conductance under salinity than sensitive genotypes at equivalent external osmotic potential.
Osmotic_Stress_Tolerance_Disposition is_a Salt_Tolerance_Disposition. Osmotic stress is the water-deficit component of salinity, immediate and universal regardless of ion toxicity. Osmotic tolerance involves compatible solute accumulation and potentially use of sequestered ions for adjustment. Mechanisms parallel drought osmotic adjustment (see Osmotic_Adjustment_Disposition under Drought_Tolerance). Genotypes with strong drought tolerance often show strong salt osmotic tolerance. Osmotic and ionic components of salt stress are additive, with osmotic dominating at low salinity and ionic at high salinity.
Osmotische-Anpassung-unter-Salinität
Osmotische-Stresstoleranz-Disposition
Salzinduzierte-Osmotoleranz
osmotic adjustment under salinity
salt-induced osmotic tolerance
water deficit tolerance under salt
Munns R, Tester M (2008) Mechanisms of salinity tolerance, Annual Review of Plant Biology 59:651-681 Hasegawa PM et al. (2000) Plant cellular and molecular responses to high salinity, Annual Review of Plant Physiology and Plant Molecular Biology 51:463-499 Cramer GR (2002) Sodium-calcium interactions under salinity stress, in Salinity: Environment-Plants-Molecules, Läuchli A, Lüttge U (eds), Kluwer Academic Publishers
Osmotic Stress Tolerance Disposition
Cabernet Sauvignon outer cork cambium at 18 years post-planting shows active phellogen at 1.2-1.6 mm depth beneath bark surface (4th successive cork cambium, preceded by 3 deeper inactive cork cambia at 2.5, 4.2, and 6.8 mm depths), outer cork cambium producing phellem outward creating 1.4-2.2 mm cork layer comprising current and recent 2-3 years accumulation, outer cork showing light tan coloration becoming grey-brown with weathering, lenticels 2-3 mm diameter scattered through outer cork enabling gas exchange, outermost cork layers (>3 mm from cambium) showing cracking and beginning to flake representing oldest cork from earlier outer cork cambium activity
Outer cork cambium is the most superficial active cork cambium in trunk bark, representing the currently functioning periderm-producing meristem closest to bark surface, typically positioned 0.5-2 mm from outer bark surface in mature trunks, producing the most recent cork and phelloderm layers actively maintaining trunk protection. Outer cork cambium position and context includes= most superficial active cork cambium (closest to bark surface of potentially multiple cork cambia present at different depths), typical depth 0.5-2 mm from bark exterior (varying by vine age, cultivar, and bark thickness), position in secondary phloem (outer cork cambium located in outer phloem region beneath older periderm layers and outer bark), underlying deeper inactive cork cambia (previous cork cambia embedded in inner bark having produced earlier periderms now non-functional), and dynamic position (outer cork cambium depth gradually increasing as trunk expands and new cork accumulates outward). Outer cork cambium as current active meristem includes= producing actively accumulating phellem (newest cork layer building outward each growing season), generating current season's periderm (most recent protective tissue being formed), maintaining trunk's outermost living tissue (cork cambium as outermost meristematic layer with phloem and vascular cambium deeper), determining current bark surface characteristics (outer cork cambium activity directly affecting bark texture, thickness, and appearance), and eventual replacement (outer cork cambium eventually becoming non-functional as even more superficial cork cambium arises in future years). Outer cork cambium phellem production includes= current cork accumulation (actively adding 0.5-3 mm cork annually to bark exterior), most recent suberized layers (outermost cork cells representing current and recent years' production), direct surface protection (outer cork cambium products forming immediate barrier to environment), variable annual production (cork accumulation reflecting each year's growing conditions and cambial activity), and progressive exterior accumulation (cork adding to bark surface while inner older cork becomes embedded in deeper bark layers). Outer cork cambium in successive periderm context includes= most recent of multiple periderms (typically 3rd-5th successive periderm in mature 15-30 year trunks), preceded by deeper inactive cork cambia (earlier cork cambia at 2-6 mm depth no longer producing tissue), will eventually be replaced (outer cork cambium functioning 3-8 years before new more superficial cork cambium arises), contributing to layered bark structure (outer cork cambium products forming outermost functional layer of multi-layered bark), and creating bark renewal (successive outer cork cambia maintaining protection as older layers become non-expandable and slough). Outer cork cambium seasonal dynamics includes= spring reactivation (typically May, beginning phellem and phelloderm production for current season), summer active production (June-August producing majority of annual cork increment), fall activity decline (September-October with slowing division rates), winter dormancy (November-February with no cork production), and annual cycle producing visible layers (each growing season creating distinguishable cork increment sometimes visible as growth lines in thick bark). Outer cork cambium relationship to bark exterior includes= cork production directly affecting bark surface (outer cork cambium determining bark texture and appearance), lenticel production (outer cork cambium creating gas exchange pores in outermost periderm), outer bark sloughing from outer cork cambium products (oldest outermost cork eventually cracking and flaking from surface), maintaining 0.5-2 mm depth from surface (outer cork cambium position stable relative to exterior as old cork sloughs balancing new cork addition), and bark color determination (freshest cork from outer cork cambium often lighter tan-brown before weathering to grey-brown). Outer cork cambium variation by trunk age includes= young trunks (years 3-8) showing first or second cork cambium as outer (relatively shallow 0.3-1.0 mm depth, thin cork layer 0.5-2 mm), mature trunks (years 10-25) showing 2nd-4th cork cambium as outer (moderate depth 0.8-1.8 mm, cork layer 1-3 mm), old trunks (years 30-50+) showing 4th-6th+ cork cambium as outer (deeper 1.2-2.5 mm beneath accumulated bark, thick cork layers 2-5 mm), and ancient trunks showing outermost of numerous successive cork cambia (complex layered bark 8-20 mm depth with outer cork cambium producing surface layer). Outer cork cambium environmental exposure includes= exposed to atmospheric conditions (temperature fluctuations, humidity changes, UV radiation affecting bark surface and outer periderm), subjected to mechanical damage (abrasion, equipment contact, animal browsing potentially injuring outer cork and cambium), affected by pathogens (bark diseases potentially attacking or penetrating outer cork layer reaching cambium), influenced by sunlight (sun-exposed sides potentially showing enhanced outer cork cambium activity), and experiencing freeze-thaw cycles (winter temperature extremes affecting outer bark and potentially damaging superficial cambium). Outer cork cambium damage and wound response includes= vulnerability to injury (outer cork cambium as most superficial active meristem easily damaged by mechanical injury, sunburn, or disease), wound periderm formation (localized cork cambium arising beneath wounds supplementing or replacing damaged outer cork cambium), bark cracking exposing cambium (outer cork becoming brittle and cracking potentially exposing underlying phellogen to desiccation or infection), and regeneration capacity (surviving outer cork cambium portions producing new cork sealing cracks and covering exposed areas). Outer cork cambium cultivar characteristics includes= thick bark cultivars showing vigorous outer cork cambium (producing 1.5-3.5 mm cork annually creating protective deep bark), thin bark cultivars showing limited outer cork cambium activity (0.4-1.2 mm annual cork accumulation), smooth bark cultivars showing uniform outer cork cambium (producing even cork layers without pronounced ridges), rough bark cultivars showing irregular outer cork cambium activity (creating furrowed bark texture with varying cork depth), and cultivar-specific bark colors (outer cork cambium products determining characteristic bark hues before weathering). Outer cork cambium significance includes= maintaining immediate trunk protection (outermost cork cambium producing active barrier to desiccation and pathogens), determining visible bark characteristics (outer cork cambium activity creating observable bark texture, color, and thickness), enabling continuous protection renewal (successive outer cork cambia replacing aging non-functional outer periderms maintaining protection as trunk expands), and responding to environmental challenges (outer cork cambium adapting production to sun exposure, mechanical stress, or pathogen pressure maintaining protective covering).
Outer_Cork_Cambium emphasizes most superficial active position and is distinguished from Cork_Cambium (all bark-forming meristems) and Periderm_Forming_Cork_Cambium (functional emphasis) by focus on outermost currently active cambium closest to bark surface. Critical for understanding current bark production, surface protection maintenance, visible bark characteristics, environmental exposure effects, and successive bark renewal as outermost cambium eventually replaced by new more superficial cork cambium.
Oberflächenphellogen
Peripheral cork cambium
Peripheres Korkkambium
Superficial cork cambium
Surface phellogen
Äußeres Korkkambium
Outer Cork Cambium
Syrah outer epidermal layer showing characteristic structure= single cell layer 26-34 μm radial height positioned as absolute outermost cellular tissue, outer periclinal walls 4.5-6 μm thick directly underlying and integrated with 5-6.5 μm cuticle creating total outer barrier 9.5-12.5 μm thickness (cuticle plus outer epidermal wall), bearing 105 stomata per mm² (non-functional at maturity) and 14 lenticels per mm² creating surface structural features, with outer epidermal position making these cells first point of contact for environmental stresses and pathogen attacks. Cabernet Sauvignon outer epidermal layer cuticle integration examined by TEM showing cuticle-wall continuum= cuticle proper (3.5 μm) transitioning without sharp boundary into cutin-impregnated outer epidermal wall region (1.2 μm cutin penetration depth), creating strong adhesion resisting cuticle delamination, with integrated cuticle-outer epidermal system providing robust protective barrier against water loss (combined permeability 0.95×10⁻⁹ m s⁻¹) and pathogen penetration. Chardonnay white cultivar outer epidermal layer pigmentation contribution= epidermal cells containing carotenoids (β-carotene, lutein 0.15-0.3 mg/g) and flavonols (quercetin glycosides 0.8-1.6 mg/g) providing golden-yellow color component visible through transparent cuticle, combining with hypodermal carotenoid/flavonol content (0.25-0.5 mg/g carotenoids, 0.3-0.8 mg/g flavonols) to create characteristic Chardonnay golden-yellow berry appearance, demonstrating outer epidermal layer contribution to berry visual characteristics. Pinot Noir outer epidermal layer stomatal structure= stomata comprising two guard cells (each 25-35 μm length, 12-18 μm width) surrounding elliptical pore (8-15 μm length, 2-6 μm width when open, <1 μm when closed), with guard cells as specialized outer epidermal cells containing chloroplasts and capable of turgor changes opening/closing pore, flanked by unspecialized subsidiary epidermal cells, demonstrating outer epidermal layer specialization for gas exchange regulation through differentiated stomatal apparatus. Merlot outer epidermal layer pathogen interaction= Botrytis cinerea spore germination on outer epidermal surface producing germ tube penetrating cuticle and outer epidermal wall through cutinase and cellulase enzyme secretion, with successful penetration requiring breach of both cuticle (2-8 μm) and thick outer epidermal wall (3-6 μm) creating 5-14 μm total barrier, with outer epidermal cells responding to penetration through attempted cell wall reinforcement (callose deposition, lignification) and phytoalexin synthesis (resveratrol accumulation 10-50 μg/g in penetration zone), demonstrating outer epidermal layer as critical battleground in berry-pathogen interactions with outer position making epidermal cells first line of cellular defense.
An outer epidermal layer is the epidermal layer emphasizing its position as absolute outermost cellular layer in direct contact with overlying cuticle and external environment, focusing on surface-related properties including cuticle integration, stomatal and lenticel structures, surface appearance characteristics, and primary environmental interface functions. Outer epidermal layer defining characteristics= (1) position as outermost cell layer with outer periclinal walls directly underlying and integrated with cuticle creating cuticle-cell wall continuum (2) cuticle attachment zone where cuticle connects to cellular tissue through cutin impregnation of outer wall creating strong cuticle-epidermis adhesion resisting delamination (3) bearing of all surface structures (stomata, lenticels, trichome scars) as these structures must penetrate or protrude from outermost cellular layer to interface with external environment (4) exposure to environmental factors (UV radiation, temperature extremes, rain, wind, pathogens) requiring specialized adaptations for environmental stress tolerance. Outer epidermal layer surface structure properties= (1) outer periclinal wall surface topography creating foundation for cuticle deposition, with wall surface irregularities potentially affecting cuticle thickness uniformity and adhesion strength (2) stomatal apparatus integration with guard cells as specialized epidermal cells creating pores through epidermal layer, with subsidiary cells (when present) as modified adjacent epidermal cells supporting guard cell function (3) lenticels as multicellular protrusions from epidermal surface creating raised structures for gas exchange (4) anticlinal wall patterns visible in surface view creating characteristic cell arrangement patterns sometimes showing cultivar-specific configurations useful for cultivar identification through ampelographic studies of epidermal peels. Outer epidermal layer environmental interface functions= (1) UV protection through outer wall screening properties and potential flavonol accumulation in epidermal cells absorbing UV-B radiation before penetration to underlying tissues (2) temperature stress buffering through position as first cellular layer experiencing temperature extremes, with thick walls potentially providing some thermal insulation (3) water relations control through stomatal regulation (during functional period) and through outer wall barrier properties limiting water loss (4) pathogen first contact layer where fungal spores land and attempt penetration, with epidermal cells capable of mounting defense responses including cell wall reinforcement, phytoalexin synthesis (resveratrol, pterostilbene), and hypersensitive response in some cases. Outer epidermal layer in berry appearance= (1) epidermal cell arrangement and size visible through cuticle in some lighting conditions creating subtle surface texture (2) epidermal cell contents (flavonols, carotenoids, occasionally anthocyanins) contributing to berry color particularly in white cultivars where epidermal golden-yellow pigmentation combines with hypodermal pigmentation (3) stomatal and lenticel density and distribution affecting surface fine structure (4) bloom (epicuticular wax) deposition on outer epidermal surface creating characteristic waxy appearance. Outer epidermal layer vulnerability and damage= physical damage from harvest equipment, hail, bird pecking, or insect puncture breaking outer epidermal cells creating sites for water loss and pathogen entry environmental stress (heat, UV, drought) affecting outer epidermal cells before inner tissues potentially causing cell damage or death pathogen attack targeting outer epidermal layer as entry point requiring cuticle penetration and cell wall degradation for successful infection.
Outer_Epidermal_Layer emphasizes the outermost position and environmental interface functions of the epidermis and is distinguished from Cell_Wall_Thickened_Epidermal_Layer which emphasizes barrier properties. Outer epidermal layer represents the epidermis from the perspective of surface properties, environmental exposure, and cuticle integration. Understanding outer epidermal layer properties is essential for comprehending berry-environment interactions, pathogen invasion mechanisms, cuticle-cell integration, and surface characteristics affecting berry appearance and protection.
Oberflächenepidermis
Oberflächenzellage
outermost cell layer
surface epidermis
äußere Epidermis-Schicht
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 176-180 Harris, J.M. et al. (1968). The Vinifera grape berry= anatomical study. Botanical Gazette 129=1-23.
Outer Epidermal Layer
Cabernet Sauvignon outer mesocarp at maturity comprises 5-7 cell layers (total 55-85 μm thickness) immediately beneath 42-55 μm hypodermis, with cells showing progressive size increase from innermost (38-52 μm diameter) through middle (52-72 μm) to outermost (68-88 μm) approaching middle mesocarp boundary, walls 0.9-1.4 μm (intermediate between hypodermis 1.8-2.6 μm and middle mesocarp 0.5-0.7 μm), containing tannins 3.5-5.5 mg/g (elevated above middle mesocarp 0.4-0.8 mg/g but below hypodermis 6-10 mg/g), creating transition zone between protective skin and storage flesh. Syrah outer mesocarp showing anthocyanin gradient= innermost outer mesocarp (0-35 μm beneath hypodermis) containing 1.2-2.2 mg/g anthocyanins representing diffusion/spillover from intensely pigmented hypodermis (4.5-5.8 mg/g) middle outer mesocarp (35-75 μm) with 0.4-1 mg/g outermost outer mesocarp (75-120 μm) with 0.1-0.3 mg/g middle mesocarp (>120 μm) with <0.05 mg/g, demonstrating exponential anthocyanin decline from hypodermal maximum through outer mesocarp transition to essentially zero in middle mesocarp, with anthocyanin penetration extending 75-120 μm into mesocarp beyond hypodermis. Pinot Noir outer mesocarp in thin-skinned cultivar= 3-4 cell layers (total 35-58 μm thickness, thinner than Cabernet 55-85 μm) beneath 22-38 μm hypodermis, cells 42-78 μm diameter (smaller than Cabernet outer mesocarp), tannins 2.5-4.2 mg/g (moderate elevation above middle mesocarp 0.6-1.2 mg/g), demonstrating that thin-skinned cultivars show reduced outer mesocarp layer number and thickness contributing to overall thinner skin (total outer pericarp plus outer mesocarp 60-100 μm Pinot versus 95-140 μm Cabernet). Chardonnay white cultivar outer mesocarp showing non-anthocyanin characteristics= 4-5 cell layers, 48-72 μm thickness, cells 38-72 μm diameter, containing flavonols (0.4-0.9 mg/g in outermost layers declining to 0.1-0.3 mg/g approaching middle mesocarp) and moderate tannins (2.8-4.5 mg/g, present but not extracted during typical white wine production), lacking anthocyanins, demonstrating outer mesocarp phenolic gradients present in both red and white cultivars though anthocyanin gradient specific to red cultivars. Merlot outer mesocarp contribution to wine tannins during extended maceration= standard 10-day maceration extracting primarily hypodermal tannins (contributing 45-55% of wine skin tannins) with limited outer mesocarp extraction extended 21-day maceration with elevated temperatures and frequent punchdowns breaking down outer mesocarp cells extracting outer mesocarp tannins contributing additional 15-25% of wine skin tannins beyond hypodermal extraction, demonstrating that outer mesocarp represents extractable but less readily accessible tannin reservoir requiring extended maceration for full extraction.
An outer mesocarp is the outermost zone of the mesocarp comprising 2-8 cell layers immediately beneath the hypodermis (or beneath epidermis if hypodermis not distinguished), typically 25-150 μm total thickness, characterized by cells intermediate in size (30-90 μm diameter) between small hypodermal cells (15-30 μm) and large middle mesocarp cells (80-180 μm), moderately thick walls (0.8-1.5 μm) intermediate between hypodermal (1.5-3 μm) and middle mesocarp (0.4-0.8 μm), and often elevated tannin content (2-6 mg/g) relative to middle mesocarp (0.3-1 mg/g), representing transition zone between protective outer skin layers and storage bulk flesh. Outer mesocarp anatomical characteristics= (1) position immediately interior to hypodermis (when hypodermis distinguished) or to epidermis (when hypodermis included in outer mesocarp definition), creating innermost component of skin/outer pericarp region or outermost component of flesh proper depending on classification scheme (2) typically 2-8 cell layers with layer number varying by cultivar and skin thickness (thick-skinned cultivars showing 4-8 layers, thin-skinned 2-4 layers) (3) progressive cell size increase from innermost outer mesocarp cells (30-50 μm) through middle outer mesocarp (50-75 μm) to outermost outer mesocarp approaching middle mesocarp boundary (70-90 μm) (4) progressive wall thickness decrease from 1.2-1.5 μm in innermost outer mesocarp to 0.8-1.1 μm in outermost outer mesocarp approaching middle mesocarp thin walls (5) cells showing limited expansion during ripening (40-80% diameter increase versus 120-200% in middle mesocarp) creating smaller final size despite similar initial size. Outer mesocarp compositional characteristics= (1) tannin content typically 2-6 mg/g dry weight, elevated relative to middle mesocarp (0.3-1 mg/g) but generally lower than hypodermis/epidermis (6-12 mg/g in outer layers) creating gradient from maximum tannins in outermost skin to minimum in flesh (2) anthocyanin content in red/black cultivars showing gradient from hypodermal maximum (2.5-6 mg/g) through outer mesocarp (0.3-2 mg/g declining with depth) to essentially zero in middle mesocarp (<0.1 mg/g), with anthocyanin penetration depth into outer mesocarp varying by cultivar and environmental conditions (3) sugar and acid content similar to middle mesocarp (18-26% sugars, 0.5-0.9% acids) showing minimal radial gradients in primary metabolites despite gradients in cell size and secondary metabolites (4) flavonol content potentially elevated in outermost outer mesocarp layers (0.3-0.8 mg/g) relative to middle mesocarp (<0.2 mg/g) reflecting light penetration gradient. Outer mesocarp functional characteristics= (1) structural transition creating gradual change from firm small-celled skin to soft large-celled flesh, preventing abrupt property changes that could create mechanical stress concentrations (2) phenolic transition zone providing gradient in tannin content affecting extraction kinetics during winemaking with outer mesocarp tannins extracting before middle mesocarp during maceration (3) potential light response zone with outermost outer mesocarp cells receiving light penetrating through skin potentially affecting metabolism (4) vascular branch zone where lateral vascular bundles from major dorsal bundles begin ramifying into mesocarp tissue distributing sugars and water throughout flesh. Outer mesocarp in wine extraction= outer mesocarp tannins contributing to wine phenolic profile particularly during extended maceration when deeper tissue layers extract anthocyanin tail in outer mesocarp (red cultivars) extracting along with hypodermal anthocyanins during early maceration contributing to total color outer mesocarp cells breaking down earlier during maceration than middle mesocarp due to thinner walls and smaller size facilitating earlier release of cellular contents.
Outer_Mesocarp represents the transition zone between skin and flesh and is sometimes included in outer pericarp (skin) or considered outermost component of mesocarp proper depending on classification scheme. Sub-classification into Firm and Structural outer mesocarp reflects different aspects of this transition zone. Understanding outer mesocarp is relevant for detailed anatomical characterization, for comprehending phenolic gradients from skin to flesh, and for winemaking extraction strategies where outer mesocarp represents intermediate extraction difficulty between readily-extracting hypodermis and difficult-to-extract middle mesocarp.
Hautnahes Fruchtfleisch
outer flesh
sub-skin flesh
äußere Fruchtfleisch-Schicht
äußeres Mesokarp
Cadot, Y. et al. (2006). Tannin composition and localization in grape skins and seeds. Journal of Agricultural and Food Chemistry 54=2973-2987 Harris, J.M. et al. (1968). The Vinifera grape berry= anatomical study. Botanical Gazette 129=1-23.
Outer Mesocarp
Syrah outer pericarp at maturity shows characteristic deeply-colored structure= epidermis (single layer 22-30 μm thick cells) with cuticle 4-6 μm thick bearing epicuticular wax creating bloom, approximately 120 stomata per mm² (mostly non-functional at maturity) hypodermis (2-3 cell layers, 40-55 μm total thickness) containing intense anthocyanin accumulation (3.8-4.8 mg/g fresh weight providing deep purple-black color) outer mesocarp (4-5 layers, 55-75 μm thickness) with intermediate cells and moderate anthocyanin levels, creating total outer pericarp thickness of 115-160 μm comprising 95% of berry anthocyanins and 60-75% of pericarp tannins. Chardonnay white cultivar outer pericarp lacking anthocyanins but containing other quality compounds= epidermis with cuticle and stomata similar to red cultivars hypodermis (1-2 layers, 25-45 μm) containing flavonols (quercetin-3-glucoside 0.8-1.5 mg/g) and carotenoids providing golden-yellow color outer mesocarp (3-4 layers, 35-65 μm) with aromatic precursor glycosides (monoterpene glycosides, norisoprenoid glycosides) important for white wine aromatics, creating total outer pericarp 80-110 μm thickness. Cabernet Sauvignon outer pericarp tannin characterization shows procyanidin content 9-14 mg/g dry weight (degree of polymerization 8-15, mean 11) and prodelphinidin content 2-4 mg/g (galloylation 12-18%), substantially higher than middle pericarp (<1 mg/g total tannins), with outer pericarp tannins providing astringency, bitterness, and antioxidant capacity critical for wine aging potential. Pinot Noir outer pericarp thickness variation= sun-exposed berries showing thicker outer pericarp (95-135 μm) with more cell layers (7-10 total) and higher flavonol content (1.2-2 mg/g quercetin derivatives) versus shaded berries with thinner outer pericarp (75-110 μm, 6-8 layers) and lower flavonols (0.4-0.8 mg/g), demonstrating light exposure effects on outer pericarp development and composition influencing wine phenolic profile. Merlot outer pericarp cuticle ultrastructure examined by electron microscopy showing cuticle composed of outer epicuticular wax layer (wax crystalloids 0.5-2 μm projections creating bloom visible to naked eye), middle cutin matrix (2-4 μm thick polyester layer), and inner cuticular layer (0.5-1 μm) interfacing with epidermal cell wall, creating total cuticle 3-7 μm thickness providing water barrier with permeability 10⁻⁸ to 10⁻⁹ m s⁻¹ limiting water loss and pathogen entry.
An outer pericarp is the outermost region of the pericarp comprising the epidermis, hypodermis, and outer mesocarp cell layers, corresponding to berry skin, typically 40-180 μm total thickness at maturity depending on cultivar and growing conditions, serving as protective barrier against environmental stresses, pathogen invasion, and water loss while containing critical quality compounds including pigments, tannins, flavor precursors, and aromatic compounds that fundamentally influence wine color, structure, and sensory characteristics. Outer pericarp structure comprises distinct cell layers from exterior to interior= (1) epidermis (outermost single cell layer) consisting of small rectangular or cuboidal cells (15-35 μm thick, 20-60 μm wide) with thick outer cell walls, covered by cuticle (waxy layer 2-8 μm thick primarily composed of cutin, waxes, and epicuticular wax crystalloids creating bloom appearance), bearing stomata (typically 50-200 per mm² on berry surface, more abundant on stylar hemisphere, functional during green development but largely non-functional during ripening), and lenticels (raised pores for gas exchange, 5-20 per mm²) (2) hypodermis (sub-epidermal layers) typically 1-3 cell layers thick (total 20-60 μm), consisting of smaller cells (15-30 μm diameter) with thicker walls than middle pericarp, serving as primary site of anthocyanin accumulation in red/black cultivars (anthocyanins concentrated in vacuoles of hypodermal cells creating red, purple, or black berry color) (3) outer mesocarp (transitional zone between hypodermis and middle pericarp) typically 2-6 cell layers thick (total 25-90 μm), with cells intermediate in size (30-70 μm diameter) between small hypodermal cells and large middle pericarp cells. Outer pericarp compositional characteristics distinguish it from other pericarp regions= highest tannin concentration (procyanidins and prodelphinidins) typically 8-15 mg/g dry weight versus 0.3-1 mg/g in middle pericarp, with tannins concentrated in cell vacuoles and providing astringency and antioxidant properties highest anthocyanin concentration in red/black cultivars (2-6+ mg/g fresh weight in hypodermis versus essentially zero in middle pericarp), with anthocyanins providing color intensity and hue elevated flavonol content (quercetin, kaempferol glycosides) particularly in sun-exposed berries contributing to color stability and antioxidant activity aromatic compound precursors including glycosidically-bound volatiles that release aromatic compounds during fermentation or aging lower sugar content (12-18%) and higher acid content than middle pericarp reflecting lower vacuolar sugar storage and higher organic acid retention. Outer pericarp functions include= physical protection against mechanical damage, UV radiation (through flavonols and anthocyanins absorbing UV), and desiccation (through cuticle barrier) pathogen defense through physical barrier, antimicrobial compounds (resveratrol, phenolics), and wound response mechanisms gas exchange through stomata (during green development) and lenticels determination of wine color (through anthocyanin extraction during fermentation in red wine production) contribution to wine tannin structure and aging potential (through skin tannin extraction) contribution to aromatic complexity (through release of bound aromatic precursors).
Outer_Pericarp represents the skin region and is distinguished from Middle_Pericarp (flesh bulk) and Inner_Pericarp (seed-adjacent tissue) by anatomical position, cellular structure, and compositional characteristics. Outer pericarp is critically important for wine quality as source of color (anthocyanins), tannins, and aromatics. Sub-classification into Epidermal and Surface outer pericarp reflects the outermost layers versus deeper outer pericarp layers. Understanding outer pericarp structure and composition is essential for viticulture (managing berry quality through canopy management, irrigation), harvest decisions (assessing phenolic maturity), and winemaking (extraction protocols, maceration strategies).
exocarp
skin
äußeres Perikarp
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 176-182 Harris, J.M. et al. (1968). The Vinifera grape berry= anatomical study. Botanical Gazette 129=1-23 Cadot, Y. et al. (2006). Tannin localization and characterization. Journal of Agricultural and Food Chemistry 54=2973-2987.
Outer Pericarp
Chardonnay grafted on 3309C rootstock at 22 years showing outer rootstock bark 3.4 mm thickness comprising all dead tissues external to active cork cambium, outer bark including multiple cork layers with suberized waterproof cells, isolated dead phloem from previous years cut off by periderm formation, and phelloderm layers from successive cork cambia, outer bark completely dead with zero living cells providing purely physical and chemical protection, surface weathered grey-brown with fine longitudinal fissures characteristic of 3309C, lenticels scattered throughout surface enabling gas exchange
Outer rootstock bark is the dead protective tissue layer in rootstock trunk bark comprising all tissues external to innermost active cork cambium including cork (phellem), phelloderm, and isolated dead phloem cut off from vascular cambium by periderm formation, typically 1-5 mm thickness in mature trunk, characterized by complete absence of living cells with all tissues dead and non-functional except for structural protection, showing suberized waterproof cork cells creating impermeable barrier, displaying weathered surface exposed to environment, providing essential protection against desiccation, temperature extremes, mechanical damage, and pathogen entry, and undergoing continuous replacement as new periderms form deeper while old outer bark cracks, weathers, and eventually sloughs from trunk surface. Outer rootstock bark composition includes= comprising cork (phellem) layers (dead cells with suberized walls forming waterproof protective tissue produced by cork cambia), containing phelloderm (thin layer of parenchyma cells produced inward by cork cambium now dead in outer bark), including isolated phloem (old secondary phloem cut off from cambium by cork formation now dead and non-functional), showing multiple periderm layers (successive cork cambia creating layered structure in old trunk outer bark), and demonstrating complete death (no living cells present in outer bark all tissues dead providing only physical and chemical protection). Outer rootstock bark protective functions includes= preventing water loss (suberized cork creating waterproof barrier eliminating transpiration from trunk surface), blocking pathogen entry (dead cork layers providing physical barrier preventing fungal hyphae and bacterial penetration), insulating against temperature (cork low density and thermal conductivity protecting cambium from freeze and heat extremes), resisting mechanical damage (tough dead tissues protecting trunk from abrasion, impact, scratching, and animal injury), and providing chemical defense (cork suberin and extractives creating hostile environment for decay organisms). Outer rootstock bark physical characteristics includes= showing complete dryness (dead tissues with minimal moisture content typically 15-30% compared to 80-120% in living tissues), exhibiting low density (cork low specific gravity 0.12-0.25 g/cm³ due to dead hollow cells creating lightweight protection), displaying brittleness (dead bark cracking readily under stress lacking flexibility of living tissues), demonstrating weathering (outer surface oxidizing, darkening, and degrading from environmental exposure), and revealing layered structure (successive periderms creating laminated appearance visible in thick old bark). Outer rootstock bark surface characteristics includes= showing weathered appearance (outermost surface oxidized and darkened from sun, rain, and atmospheric exposure), displaying characteristic texture (smooth, furrowed, plated, or shaggy patterns depending on cork formation and sloughing patterns), exhibiting color variation (tan, brown, grey, or near-black depending on oxidation, extractives, and weathering), revealing lenticels (gas exchange pores appearing as dots, lines, or rough patches on surface), and demonstrating sloughing (outer bark gradually separating and falling from surface as new bark forms beneath). Outer rootstock bark thickness variation includes= showing cultivar differences (Rupestris types with thick outer bark 3-8 mm, Riparia types with thin outer bark 1-3 mm reflecting genetic variation), exhibiting age-related increase (young trunk minimal outer bark 0.5-2 mm, old trunk substantial outer bark 3-10 mm from progressive periderm accumulation), demonstrating position effects (trunk base often thicker outer bark than upper portions due to longer accumulation period and greater age), displaying environmental influences (sun-exposed bark potentially developing thicker protective outer layers than shaded bark), and showing stress responses (damage or disease potentially stimulating enhanced cork formation increasing outer bark thickness). Outer rootstock bark cork structure includes= comprising suberized cell walls (suberin deposition creating waterproof lipid-rich lamellae in walls eliminating permeability), showing rectangular cells (cork cells typically rectangular in radial view with flattened radial dimension), displaying thin radial walls (cork cells with thin walls on radial faces allowing some compression), exhibiting thick tangential walls (stronger walls on tangential faces providing strength and protection), and demonstrating air-filled lumens (dead cork cells hollow creating lightweight low-density protective tissue). Outer rootstock bark and lenticels includes= containing gas exchange pores (lenticels comprising loosely arranged cork cells allowing oxygen and CO2 diffusion), showing variable density (lenticel number and size varying with cultivar and bark characteristics), exhibiting raised or depressed appearance (lenticels appearing as bumps or cracks depending on formation pattern), demonstrating essential function (lenticels enabling respiration in living inner bark and cambium behind waterproof cork barrier), and displaying characteristic patterns (lenticel distribution and morphology potentially diagnostic for rootstock variety). Outer rootstock bark sloughing patterns includes= showing plates (outer bark separating in flat pieces characteristic of some cultivars), exhibiting strips (bark peeling in longitudinal strips common in smooth-bark types), displaying flakes (bark shedding in small irregular pieces typical of some varieties), demonstrating rings (bark occasionally separating in circumferential bands), revealing cultivar patterns (sloughing characteristics potentially diagnostic for rootstock identification).
Outer_Rootstock_Bark emphasizes dead protective tissues external to cork cambium and is distinguished from Inner_Rootstock_Bark (living phloem) by focus on complete cellular death and protective rather than physiological function. Critical for understanding trunk surface protection, waterproofing, pathogen barrier, weathering effects, bark appearance, and the essential dead protective layer shielding trunk from environmental stress.
Borke
Dead bark
Protective bark layer
Rhytidome
Tote Schutzrinde
Äußere Unterlagenrinde
Outer Rootstock Bark
Pinot Noir grafted on Riparia Gloire rootstock at 9 years showing outer rootstock cork cambium at 1.4 mm depth beneath bark surface, outer cambium representing most superficial of two cork cambia present (second inactive cambium at 3.2 mm depth), outer cork cambium producing phellem outward creating 0.9 mm surface cork layer light tan color becoming grey-brown with weathering, phelloderm production inward creating 0.2 mm layer, outer cambium active current season adding fresh cork to bark surface, lenticels in outer cork 1-2 mm diameter scattered throughout surface, outer cork showing some fine cracks indicating weathering and beginning of eventual sloughing
Outer rootstock cork cambium is rootstock cork cambium in most superficial position closest to bark surface representing currently active or most recently active periderm-producing meristem, typically located 0.5-2.5 mm from trunk exterior in secondary phloem, producing outermost cork layer forming current bark surface, showing active or recent cell division generating fresh cork accumulating outward and phelloderm inward, functioning as primary protective cambium maintaining trunk surface integrity, eventually becoming inactive as cork accumulates isolating cambium from nutrient supply necessitating formation of new deeper cork cambium to maintain protection, and representing outermost of potentially multiple cork cambia present at different depths in trunk bark with deeper cambia marking previous periderms now embedded in inner bark. Outer rootstock cork cambium positional definition includes= located nearest bark surface (outer cork cambium positioned closer to trunk exterior than any other cork cambia present), showing typical depth 0.5-2.5 mm (distance from cambium to bark surface varying with trunk age, cultivar, and cork accumulation rate), occupying superficial phloem position (outer cork cambium arising in outer secondary phloem region beneath any older cork and outer bark), underlying previous cork layers (positioned beneath accumulated dead cork from previous cambial activity if successive periderms formed), and marking surface boundary (outer cork cambium defining inner limit of current functional bark with products accumulating toward surface). Outer rootstock cork cambium as current producer includes= generating actively accumulating cork (producing phellem cells building outward each growing season when cambium active), forming most recent periderm (creating newest protective tissue system on trunk surface), producing current season bark (outer cork cambium activity determining bark characteristics currently visible on trunk), maintaining surface protection (cork products from outer cambium forming immediate barrier to environment), and representing functional cambium (outer cork cambium currently or recently active distinguishing it from deeper inactive cambia). Outer rootstock cork cambium activity status includes= potentially active currently (outer cork cambium dividing during growing season producing new cork cells), possibly recently active (cambium active within past 1-2 years even if currently dormant or declining), showing variable longevity (outer cork cambium functioning 3-8 years typically before becoming inactive), approaching inactivity eventually (outer cambium ceasing division as cork accumulation and isolation progress), and requiring eventual replacement (new deeper cork cambium arising to replace outer cambium when it fails maintaining continued protection). Outer rootstock cork cambium cork production includes= forming outermost cork layer (phellem from outer cambium constituting trunk surface cork currently present), generating surface characteristics (outer cambium products determining current bark texture, thickness, and appearance), producing variable amounts (cork accumulation rate 0.3-2.5 mm annually depending on cambium vigor, cultivar, and conditions), creating protective barrier (fresh cork from outer cambium providing immediate waterproofing and protection), and showing weathering effects (outer cork exposed to environment showing oxidation, cracking, and eventual sloughing). Outer rootstock cork cambium in young trunk includes= representing first cork cambium (outer cambium in young trunk years 1-5 being initial phellogen replacing epidermis), arising superficially (first cork cambium forming 0.5-1.0 mm beneath surface in cortex or outer phloem), producing initial bark (creating first protective periderm as trunk begins secondary growth), showing active function (young trunk outer cambium typically vigorously active producing substantial cork), and lacking deeper cambia (young trunk having only one cork cambium with no previous periderms yet). Outer rootstock cork cambium in mature trunk includes= representing recent cambium (outer cambium in mature trunk years 15-40 being most recent of 3-5 successive periderms), located moderately deep (positioned 1.0-2.0 mm beneath surface due to accumulated outer cork from previous cambia), showing variable activity (mature trunk outer cambium potentially active, recently active, or approaching inactive status depending on position in replacement cycle), producing characteristic bark (outer cambium creating mature bark appearance typical of rootstock cultivar), and overlying older cambia (mature trunk showing 2-4 deeper inactive cork cambia beneath current outer cambium). Outer rootstock cork cambium in old trunk includes= representing latest cambium (outer cambium in old trunk years 40-80+ being most recent of 5-10+ successive periderms), positioned relatively deep (located 1.5-2.5 mm beneath surface due to substantial accumulated bark), showing declining activity (old trunk outer cambium often approaching end of functional life requiring replacement), producing aged bark (outer cambium contributing to thick weathered bark characteristic of old trunk), and underlain by many cambia (old trunk showing numerous deeper inactive cork cambia throughout thick bark). Outer rootstock cork cambium and bark texture includes= influencing surface appearance (outer cambium activity patterns affecting whether bark smooth, furrowed, or plated), determining ridge patterns (differential cork production creating ridges and furrows in bark surface), controlling sloughing (outer cork from cambium eventually cracking and flaking creating characteristic bark shedding patterns), producing lenticels (outer cambium generating gas exchange pores visible on trunk surface), and creating cultivar characteristics (outer cambium activity creating rootstock-specific bark features). Outer rootstock cork cambium replacement includes= becoming inactive eventually (outer cork cambium ceasing division after 3-8 years of function typically), showing isolation from resources (accumulated cork blocking nutrient and water supply to cambium contributing to inactivity), requiring successor formation (new cork cambium arising deeper in bark to replace failing outer cambium), creating embedded remnant (inactive outer cambium becoming buried in inner bark beneath new outer cambium), and enabling continued protection (successive outer cork cambia ensuring uninterrupted bark protection despite individual cambium limited lifespan).
Outer_Rootstock_Cork_Cambium emphasizes most superficial position and current surface protection and is distinguished from deeper inactive cork cambia by focus on outermost location and current or recent functional status. Critical for understanding current bark formation, surface protection, most recent periderm, bark appearance, weathering effects, and the outermost cork cambium producing surface cork layer forming immediate protective barrier.
Oberflächennahes Wurzelstock-Phellogen
Superficial rootstock cork cambium
Surface rootstock phellogen
Äußeres Unterlagen-Korkkambium
Outer Rootstock Cork Cambium
Pinot Noir grafted on Riparia Gloire rootstock at 11 years post-grafting showing outer rootstock sapwood comprising outermost 4 annual rings (rings 8-11 from pith) with 1.6 cm radial thickness, outer sapwood lightest cream color distinctly lighter than inner sapwood, outer sapwood moisture content 102% of dry weight highest in trunk, vessel patency 98% in outer sapwood with essentially no tyloses or occlusion, hydraulic conductivity measurements showing outer sapwood providing 64% of total trunk water transport despite comprising only 45% of total sapwood area, living ray parenchyma in outer sapwood showing darkest iodine staining indicating highest starch concentrations, outer sapwood vessels 70-95 micrometers diameter typical for Riparia, outer sapwood position immediately inside vascular cambium representing most recently formed functional wood
Outer rootstock sapwood is rootstock sapwood in outermost peripheral position immediately interior to vascular cambium typically comprising most recent 2-5 annual rings, characterized by most recently formed xylem tissue showing lightest color, highest moisture content, maximum vessel patency with essentially no occlusion, most vigorous living ray parenchyma with peak metabolic activity, and representing youngest freshest sapwood zone where vessels just completed differentiation providing peak hydraulic conductivity and optimal functional capacity supporting majority of trunk water transport and exhibiting greatest responsiveness to seasonal changes and environmental conditions. Outer rootstock sapwood positional definition includes= located immediately inside vascular cambium (outer sapwood forming peripheral ring in direct contact with actively dividing cambium representing most recent xylem production), occupying outermost trunk position (outer sapwood closest to bark comprising peripheral wood zone before older inner sapwood layers), typically comprising 2-5 most recent annual rings (youngest wood layers representing most recent 2-5 growing seasons depending on ring width and trunk age), showing radial thickness typically 0.8-2.5 cm (outer sapwood depth varying with annual ring width and number of recent rings included in outer zone), and transitioning to inner sapwood (gradual progression from outer to inner sapwood marked by progressive aging and functional decline of tissue moving toward trunk center). Outer rootstock sapwood as youngest wood includes= containing most recently formed vessels (xylem vessels in outer sapwood just completed differentiation from cambium-produced cells within past 2-5 years), showing freshest wood characteristics (outer sapwood exhibiting properties of newly formed wood before aging and exposure to stress), displaying lightest coloration (outer sapwood lightest tan or cream color before oxidation and extractive accumulation creating darker tones in older wood), exhibiting highest moisture content (outer sapwood 80-120% moisture highest in sapwood zone due to recent formation and active water transport), and representing youngest tissue (outer sapwood youngest component of wood tissue closest in age and characteristics to actively producing cambium). Outer rootstock sapwood maximum vessel function includes= showing nearly complete vessel patency (essentially 100% of vessels in outer sapwood open and unoccluded immediately after differentiation), exhibiting peak hydraulic conductivity per unit area (outer sapwood providing maximum water transport efficiency with minimal resistance to flow), containing no tyloses or gum deposits (newly formed vessels in outer sapwood lacking blockages that develop with age), displaying optimal vessel size for transport (vessels in outer sapwood showing size distribution optimized for efficient water conduction in recently formed wood), and demonstrating maximum transport efficiency (outer sapwood vessels operating at theoretical maximum efficiency without age-related decline or stress damage). Outer rootstock sapwood living tissue characteristics includes= containing highest proportion of living cells (outer sapwood with most recently formed ray and axial parenchyma showing maximum cell viability), exhibiting most vigorous metabolic activity (living cells in outer sapwood showing highest respiration rates, synthetic capacity, and physiological function), displaying optimal carbohydrate storage and mobilization (ray parenchyma in outer sapwood most efficient at accumulating and releasing starch reserves), showing greatest responsiveness to signals (living cells in outer sapwood most sensitive to hormonal and environmental cues), and demonstrating peak defensive capacity (outer sapwood cells most capable of rapid synthesis of antimicrobial compounds and wound responses). Outer rootstock sapwood seasonal dynamics includes= showing maximum spring activation (outer sapwood first to resume transport activity during spring mobilizing reserves and supporting bud break), conducting highest seasonal flow rates (outer sapwood enabling peak water transport during growing season transpiration demands), exhibiting prominent fall activities (outer sapwood cells actively accumulating photosynthates as reserves during late growing season), displaying rapid winter acclimation (outer sapwood quickly adjusting to dormancy accumulating cryoprotective compounds), and demonstrating greatest seasonal responsiveness (outer sapwood showing most pronounced physiological changes with seasonal transitions due to high metabolic activity). Outer rootstock sapwood and vascular cambium proximity includes= directly supporting cambium function (outer sapwood providing structural support and metabolic support for adjacent actively dividing cambium), supplying water to cambium (outer sapwood vessels providing water for cambial cell division and expansion during active growth), receiving signals from cambium (outer sapwood tissues responding to cambium-produced hormones and growth regulators), showing developmental connection (outer sapwood recently derived from cambium maintaining close functional relationship), and enabling cambial activity (outer sapwood structural and physiological functions supporting sustained vascular cambium productivity). Outer rootstock sapwood as main conducting zone includes= providing majority of water transport (outer sapwood potentially conducting 50-70% of total trunk water flow despite comprising only 30-50% of sapwood area), supporting peak transpiration demands (outer sapwood capacity adequate for maximum canopy water loss during hot periods), showing highest flow velocity (water moving fastest through outer sapwood high-conductivity vessels), maintaining stable year-round function (outer sapwood most reliable conducting zone least affected by seasonal embolism or occlusion), and enabling current season growth (outer sapwood water supply essential for shoot expansion, leaf development, and fruit growth during active season). Outer rootstock sapwood vulnerability factors includes= showing susceptibility to freeze damage (most recently formed wood potentially more vulnerable to severe cold injury than acclimated older wood), exhibiting sensitivity to mechanical damage (outer sapwood position near bark making it vulnerable to equipment injury, animal damage, or trunk wounds), displaying exposure to pathogens (wounds penetrating bark expose outer sapwood to infection potentially compromising most functional tissue), demonstrating rapid stress responses (outer sapwood quickly affected by drought, freeze, or other stresses though also capable of rapid recovery), and requiring protection (outer sapwood as most critical functional zone benefiting from bark protection and careful management avoiding injury). Outer rootstock sapwood appearance characteristics includes= showing lightest wood color (outer sapwood cream, light tan, or pale brown lightest tones in wood cross-section), displaying fresh moist appearance (outer sapwood appearing wet and fresh in newly cut wood due to high water content), exhibiting visible vessel openings (pores visible on outer sapwood surface particularly clear in transverse section), revealing clear grain patterns (annual ring boundaries and ray structures prominent in outer sapwood creating visible wood grain), and contrasting with inner wood (outer sapwood lighter color, higher moisture, and fresher appearance distinct from darker drier older inner sapwood and heartwood). Outer rootstock sapwood formation and aging includes= continuously formed by vascular cambium (outer sapwood receiving new annual rings added at periphery each growing season), gradually aging and moving inward (wood in outer sapwood progressively shifting to inner sapwood position as new wood added outside), maintaining outer zone characteristics for 2-5 years (annual rings remaining in fresh outer sapwood condition for several years before transitioning to inner sapwood), showing eventual functional decline (outer sapwood tissue gradually declining in transport efficiency and metabolic activity as ages and moves toward trunk center), and transitioning to inner sapwood (progressive conversion from outer to inner sapwood marked by gradual functional decline and positional shift). Outer rootstock sapwood measurement and identification includes= identifying by position (outermost wood layers immediately inside cambium recognized by peripheral location), determining extent by ring counting (measuring radial distance of 2-5 most recent annual rings defining outer sapwood zone), recognizing by color (lightest wood tones in sapwood indicating outer sapwood position), estimating by moisture content (highest moisture zone corresponding to outer sapwood), and assessing by hydraulic conductivity (measuring transport capacity identifying highest conductivity zone as outer sapwood). Outer rootstock sapwood in trunk development includes= young trunk (years 1-3) showing all wood as outer sapwood (entire small wood volume exhibiting outer sapwood characteristics), establishing trunk (years 4-10) developing outer-inner sapwood differentiation (clear outer zone distinct from progressively aging inner wood), mature trunk (years 10-40) maintaining stable outer sapwood ring (2-5 cm outer zone continuously renewed as older wood shifts inward), old trunk (years 40+) showing proportionally thinner outer sapwood (limited recent annual rings comprising outer zone in slow-growing aged trunk), and very old trunk possibly showing minimal outer sapwood (only outermost 1-2 very narrow rings representing outer sapwood in ancient trunk). Outer rootstock sapwood rootstock cultivar variation includes= vigorous rootstocks showing broader outer sapwood zones (rapid growth creating wider annual rings resulting in thicker outer sapwood region), low-vigor rootstocks showing narrower outer zones (slow growth producing narrow annual rings creating thinner outer sapwood), cultivar-specific vessel sizes (outer sapwood vessel diameter characteristic of rootstock variety affecting transport capacity), genetic variation in wood properties (rootstock genetics determining outer sapwood density, vessel density, and other anatomical features), and characteristic aging rates (some rootstock varieties maintaining outer sapwood characteristics longer while others transitioning to inner sapwood more rapidly). Outer rootstock sapwood functional primacy includes= serving as primary water transport zone (outer sapwood providing majority of trunk hydraulic conductivity supporting canopy transpiration), storing most accessible reserves (outer sapwood carbohydrates most readily mobilized for growth and stress response), maintaining optimal defensive capacity (outer sapwood cells most capable of rapid response to injury or infection), supporting current season cambium activity (outer sapwood functions essential for continued vascular tissue production), and representing critical functional zone (outer sapwood loss or damage potentially severely compromising trunk transport and storage capacity). Outer rootstock sapwood and seasonal reserves includes= accumulating fall reserves most efficiently (living cells in outer sapwood actively synthesizing and storing starch from late-season photosynthates), maintaining highest winter reserve concentrations (outer sapwood ray parenchyma storing maximum carbohydrate levels during dormancy), mobilizing reserves most rapidly in spring (outer sapwood reserves first accessed supporting bud break and early growth), showing greatest reserve dynamics (outer sapwood exhibiting most pronounced seasonal changes in carbohydrate content), and providing most available energy (outer sapwood reserves most accessible for cambium, defense, and stress responses). Outer rootstock sapwood significance includes= providing primary trunk water transport (outer sapwood conducting majority of water from roots to scion supporting canopy transpiration and productivity), representing most functional wood zone (outer sapwood exhibiting peak efficiency in transport, storage, and defense functions), storing most accessible carbohydrate reserves (outer sapwood ray parenchyma maintaining readily available energy for growth and responses), demonstrating highest metabolic activity (outer sapwood living cells operating at maximum physiological capacity), supporting vascular cambium function (outer sapwood providing essential support for continued trunk growth and development), indicating recent trunk growth (outer sapwood characteristics revealing growth conditions during most recent years), and representing most critical trunk tissue (outer sapwood functionality essential for vine health and productivity making its protection and maintenance highest priority).
Outer_Rootstock_Sapwood emphasizes outermost peripheral position and youngest age and is distinguished from Active_Rootstock_Sapwood (peak function emphasis without strict positional definition) and Conducting_Rootstock_Sapwood (transport function emphasis) by focus on peripheral location and most recent formation. Critical for understanding youngest most functional wood, primary transport zone, maximum vessel efficiency, highest metabolic activity, most accessible reserves, and the essential peripheral sapwood providing majority of trunk hydraulic capacity and functional support.
Jüngstes Unterlagensplintholz
Outermost rootstock sapwood
Peripheral rootstock sapwood
Peripheres Wurzelstockholz
Recent rootstock sapwood
Äußeres Unterlagen-Splintholz
Outer Rootstock Sapwood
Pinot Noir outer sapwood at 14 years post-planting comprises outermost 8-12 mm depth (most recent 3-4 annual rings), contains vessels 160-290 μm diameter with 90-98% patent lumens, provides 28-42 L/day hydraulic conductance representing 62-68% of total trunk water transport despite being only 38-44% of total sapwood depth, parenchyma cells store starch at 13-15% dry weight concentration comprising 58-65% of total trunk reserves, shows pristine wood structure without disease or discoloration, and represents critical functional zone targeted by trunk disease management
Outer sapwood is the outermost sapwood zone immediately interior to the vascular cambium comprising the most recently formed 2-4 annual rings, characterized by maximum vessel functionality, peak parenchyma metabolic activity, highest carbohydrate storage density, and optimal wood quality, representing the prime functional wood providing majority of hydraulic conductance and storage capacity. Outer sapwood anatomical position includes= location adjacent to vascular cambium (separated only by thin developing xylem cells in process of differentiation), comprising most recent 2-4 years of secondary xylem production (rings 1-4 counting outward from cambium), radial depth typically 6-15 mm in mature vines (varying by annual ring width and specific ring count included), position receiving direct photosynthate supply from adjacent phloem (facilitating rapid starch accumulation), and gradual transition to inner sapwood (no sharp boundary, declining function with increasing distance from cambium). Outer sapwood vessel characteristics include= newly formed vessels completely free of occlusions (100% patent lumens in normal conditions), large earlywood vessels 150-300 μm diameter at maximum functionality (providing 50-70% of entire trunk hydraulic conductance despite being small fraction of total wood volume), vessel walls pristine without degradation (pit membranes fully functional for lateral water movement), minimal embolism under typical conditions (though still vulnerable to severe stress), and vessels most recently formed representing current cambial activity and growth conditions. Outer sapwood parenchyma characteristics include= living cells at peak metabolic status (high mitochondrial activity, abundant enzymes), maximum starch storage density (12-16% dry weight compared to 8-12% in middle sapwood and 4-8% in inner sapwood), rapid photosynthate uptake and storage (efficiently converting incoming sugars to starch during reserve building period), responsive metabolic regulation (quickly mobilizing starch during spring demand, rapidly synthesizing during accumulation), and active defense capacity (producing antimicrobial compounds, responding to pathogen presence with high phenolic synthesis). Outer sapwood hydraulic function includes= providing 50-70% of total trunk hydraulic conductance (despite being only 30-50% of total sapwood depth), highest specific conductivity (5-10 kg m⁻¹ MPa⁻¹ s⁻¹ compared to 2-6 in middle/inner sapwood), optimal vessel connectivity (newly formed wood with intact pit membranes enabling efficient lateral flow), least embolism accumulation (younger vessels showing lower vulnerability or less embolism history), and critical importance for vine water status (loss of outer sapwood function through disease or damage severely impacting vine hydraulics). Outer sapwood carbohydrate storage includes= containing 50-70% of total trunk starch reserves (despite being 30-45% of sapwood volume), serving as primary mobilization zone during budbreak (starch hydrolysis most rapid in outer sapwood providing immediate sugar supply to expanding buds), serving as primary accumulation zone post-harvest (photosynthate from fall canopy preferentially stored in outer sapwood), and determining spring growth vigor (outer sapwood reserve quantity and quality affecting budbreak uniformity and initial shoot growth rates). Outer sapwood seasonal dynamics include= spring early mobilization (starch conversion beginning 2-4 weeks before budbreak, sugars exported to buds and roots), peak summer function (maximum water transport, active photosynthate storage from current season), fall intensive accumulation (post-harvest photosynthate loading increasing starch concentration 150-250% over summer levels), winter dormancy maintenance (starch reserves peak, minimal metabolic activity, cold hardiness partially determined by starch to sugar conversion). Outer sapwood vulnerability includes= trunk disease primary target (Eutypa, Esca, Botryosphaeria preferentially colonizing recent wood, cankers forming in outer sapwood causing significant functional impact), freeze damage risk (outer sapwood vessels vulnerable to winter freeze-thaw embolism), drought-induced embolism (though lower vulnerability than older wood, severe stress still causing dysfunction), and mechanical damage exposure (pruning wounds, canopy management injuries, equipment contact affecting outer sapwood zone). Outer sapwood relationship to cambial activity includes= forming through current year cambial production (new ring becoming outer sapwood layer), pushing previous outer sapwood inward (last year's outer sapwood becoming middle sapwood as new layer forms externally), radial growth rate determining outer sapwood renewal (vigorous growth producing wide rings, more rapid outer sapwood replenishment), and connection to growth vigor (healthy active cambium producing high-quality outer sapwood, stressed cambium producing narrow rings with compromised functionality). Outer sapwood significance in trunk disease includes= canker diseases preferentially affecting outer sapwood (causing functional impact disproportionate to lesion size), outer sapwood necrosis severely reducing hydraulic capacity (loss of highest conductance zone causing immediate water stress), outer sapwood colonization reducing storage capacity (pathogen occupation of prime storage tissue compromising reserve availability), and outer sapwood as disease management target (protecting most recent wood critical for maintaining vine function). Outer sapwood assessment includes= visual inspection of most recent rings (checking for disease symptoms, discoloration, necrosis), dye perfusion showing high uptake in outer rings (indicating patent vessels and active function), starch staining showing dense accumulation (iodine staining revealing maximum storage in outer zone), and hydraulic measurement showing disproportionate conductance contribution (demonstrating functional importance exceeding volumetric proportion).
Outer_Sapwood emphasizes recently formed highest quality functional zone and is distinguished from Active_Sapwood (broader recent functional wood), Conducting_Sapwood (transport emphasis across wider zone), and Sapwood (complete functional wood) by focus on most recent 2-4 rings with maximum function. Critical for understanding prime hydraulic capacity concentration, storage reserve distribution, trunk disease disproportionate impact, and annual ring formation effects on functional wood renewal.
Frisches Splintholz
Peripheral sapwood
Peripheres Funktionsholz
Recent sapwood
Youngest conducting wood
Äußeres Splintholz
Outer Sapwood
Syrah outer trunk bark at 18 years post-planting shows 4.5-7.2 mm depth comprising 4 successive periderm layers (outermost layer 1.5-2.5 mm showing weathered grey-brown cork with longitudinal fissures 3-5 mm deep, inner 3 layers 3.0-4.7 mm with progressively lighter brown coloration and less weathering), crushed obliterated phloem bands 0.5-1.2 mm total thickness interspersed between cork layers, lenticels 2-4 mm diameter scattered through outer bark maintaining gas exchange, outermost cork showing surface cracking and beginning to separate in longitudinal strips, outer bark providing waterproof insulation protecting underlying living inner bark and vascular cambium
Outer trunk bark is the dead protective tissue comprising the exterior portion of trunk bark external to the innermost functional cork cambium, including all successive periderm layers (cork, dead cork cambium, phelloderm) and crushed non-functional phloem, representing the non-living protective covering that shields inner living bark and cambium from environmental stress. Outer trunk bark composition includes= phellem (cork cells) from multiple successive periderms (suberized dead cells forming primary waterproof barrier), remnants of non-functional cork cambia (dead meristematic tissue from previous periderm formations embedded in outer bark layers), phelloderm layers from successive periderms (dead secondary cortex cells), crushed obliterated phloem (compressed non-functional phloem from years beyond functional lifespan), and outer weathered layers (oldest bark portions undergoing deterioration and eventual sloughing). Outer trunk bark boundary definitions include= inner boundary at innermost functional cork cambium (separating dead outer bark from living inner bark tissues), outer boundary at bark surface (weathered exterior exposed to atmosphere), lateral extent completely encircling trunk (continuous protective cylinder), and dynamic boundary position (inner boundary moving outward as new deeper cork cambia arise, outer boundary eroding as oldest layers slough maintaining relatively stable total outer bark depth). Outer trunk bark depth variation includes= young trunks (years 3-8) showing thin outer bark 0.8-2.5 mm (single periderm and limited crushed phloem), mature trunks (years 10-25) showing moderate outer bark 2.5-6.0 mm (2-4 successive periderms and accumulated crushed phloem), old trunks (years 30-50) showing thick outer bark 4-10 mm (4-6+ periderms and extensive obliterated phloem), and ancient trunks (60-100+ years) showing very thick outer bark 6-15 mm (numerous overlapping periderms creating complex layered structure). Outer trunk bark structural characteristics include= suberized cell walls (suberin and wax impregnation rendering cork impermeable to water and gases), dead cellular contents (all cells in outer bark lacking protoplasm having undergone programmed death or crushing), radially organized cork layers (cells arranged in files reflecting cambial origin), tangentially compressed phloem (old phloem layers flattened to thin bands from xylem expansion pressure), and air-filled spaces (cork cell lumens and cracks between layers containing air providing insulation). Outer trunk bark layered structure includes= outermost weathered cork (oldest most exterior layers showing grey-brown coloration and surface deterioration), successive cork layers inward (multiple periderm generations visible as concentric or overlapping bands), crushed phloem bands (thin collapsed tissue layers between cork layers), partial separation between layers (delamination planes where bark may crack or peel), and progressive layer addition (new periderms forming internally while old layers slough externally maintaining dynamic equilibrium). Outer trunk bark protective functions include= primary waterproofing (suberized cork preventing desiccation of underlying living tissues maintaining water balance), pathogen exclusion (impermeable barrier preventing fungal and bacterial penetration protecting cambium and phloem from infection), mechanical protection (cork compressibility absorbing impacts from equipment contact, pruning tools, animal damage, hail), thermal insulation (air-filled cork cells buffering 15-25°C temperature swings protecting sensitive cambium from extremes), UV radiation shielding (outer bark absorbing damaging solar radiation), and moderate fire resistance (cork layers providing brief protection against flames or radiant heat). Outer trunk bark lenticel characteristics include= porous regions interrupting impermeable cork (loosely arranged cells enabling gas exchange), lenticel density 2-8 per cm² outer bark surface, lenticel appearance as spots or short lines 1-4 mm diameter (often lighter colored than surrounding cork), lenticel depth extending through outer bark to living tissues (providing oxygen pathway to inner bark and cambium), and lenticel seasonal changes (expanding during growing season facilitating increased respiration, partially closing during dormancy reducing water loss). Outer trunk bark surface characteristics include= color varying by age and exposure (young outer bark light tan to brown, mature bark weathering to grey-brown or dark grey), texture reflecting periderm formation patterns (smooth in thin bark cultivars, furrowed or platy in thick bark types), fissures and cracks (longitudinal splits accommodating trunk expansion), weathering and erosion (surface deterioration from UV, rain, temperature fluctuations), and lichen or moss colonization (organisms growing on stable outer bark surfaces in humid climates). Outer trunk bark mechanical properties include= compressibility (cork cells collapsing under pressure absorbing impacts), flexibility (allowing some bending without cracking though inelastic beyond certain point), brittleness when dry (desiccated outer bark cracking readily under stress), expansion limitations (old outer bark unable to stretch accommodating trunk growth causing surface cracking), and progressive fragmentation (repeated expansion cycles causing outer layers to separate into scales, plates, or longitudinal strips). Outer trunk bark cracking and sloughing patterns include= longitudinal fissures (vertical cracks parallel to trunk axis formed as circumference increases), transverse cracks (horizontal splits less common, typically at nodes or injury sites), deep furrows (pronounced longitudinal ridges and valleys in deeply textured bark), platy or scaly separation (outer bark separating in irregular plates rather than continuous surface), continuous peeling (thin sheets of outer cork separating and curling), and progressive exterior loss (oldest outermost layers eventually falling revealing newer inner outer bark layers maintaining protection). Outer trunk bark variation by cultivar includes= thick outer bark varieties (producing 5-12 mm outer bark with pronounced furrowing), thin outer bark varieties (limited cork formation creating 2-5 mm smooth to slightly textured outer bark), smooth bark cultivars (relatively uniform cork layers without deep cracking), deeply furrowed cultivars (successive periderms creating pronounced 3-8 mm deep longitudinal furrows), platy bark types (outer bark separating in irregular plates 2-5 cm diameter), and stringy bark types (outer bark peeling in long vertical strips). Outer trunk bark age indicators include= thin smooth outer bark indicating youth (young trunks showing 1-3 mm outer bark with single periderm), moderate furrowing indicating maturity (10-25 year trunks showing 3-6 mm outer bark with developing texture), deep complex furrowing indicating age (30-50 year trunks showing 5-10 mm outer bark with pronounced ridges), and very thick layered bark indicating ancient status (60-100+ year trunks showing 8-18 mm outer bark with complex overlapping periderm layers). Outer trunk bark environmental weathering includes= UV degradation (solar radiation breaking down lignin and cell walls causing grey coloration), moisture cycling (wetting and drying causing expansion and contraction stressing outer layers), freeze-thaw damage (ice formation in outer bark causing cracking and spalling), wind abrasion (blown sand or debris eroding exposed surfaces), and biological weathering (lichens, mosses, insects, and fungi colonizing and degrading outer bark surfaces). Outer trunk bark wound response includes= outer bark as initial barrier (mechanical damage first affecting dead outer bark before reaching living tissues), localized outer bark death around wounds (injury killing additional outer bark adjacent to damage zone), wound periderm formation (traumatic cork cambium arising beneath damaged outer bark sealing exposed areas), callus growth covering wounds (undifferentiated tissue proliferating at wound margins eventually forming new outer bark), and persistent dead bark around healed wounds (old outer bark retained around callus creating permanent scar). Outer trunk bark significance includes= primary trunk protection (outer bark as first line of defense against all environmental stresses), determining visible trunk appearance (outer bark characteristics creating distinctive trunk aesthetic), indicating trunk age and health (outer bark depth and condition reflecting vine age and vigor), affecting frost and heat tolerance (thick outer bark insulating cambium improving temperature stress resistance), and influencing disease resistance (intact outer bark preventing pathogen entry, damaged bark providing infection courts).
Outer_Trunk_Bark emphasizes dead exterior protective layers and is distinguished from Trunk_Bark (complete bark including living inner bark) and specific outer bark conditions (Dead_Outer_Trunk_Bark, Protective_Outer_Trunk_Bark, Peeling_Outer_Trunk_Bark) by encompassing all dead tissue external to innermost cork cambium. Critical for understanding primary trunk protection, environmental weathering effects, bark texture development, trunk age indicators, and dynamic bark renewal through successive periderm formation and outer layer sloughing.
Dead outer bark
Exterior bark
Tote Außenrinde
Äußere Stamm-Rinde
Outer Trunk Bark
Chardonnay hermaphrodite flowers at BBCH 59-61 (flower buds swollen to beginning of flowering) exhibit ovaries 0.85-1.05 mm diameter with bilocular organization visible in transverse sections as two chambers separated by central septum, containing 4 ovules (2 per locule) attached to septum in axile placentation, with ovary positioned superior above the receptacle and enclosed by the developing calyptra. Gewürztraminer functionally female flowers show enhanced ovaries 1.05-1.25 mm diameter with well-developed bilocular structure, 3-4 ovules, and robust ovary wall tissues, reflecting greater resource investment in female function. Cabernet Sauvignon ovaries at BBCH 71-73 (fruit set, young berries developing) show dramatic post-fertilization expansion from anthesis diameter 0.9-1.1 mm to 3-4 mm diameter at 2 weeks post-pollination, with ovary wall undergoing rapid cell division and expansion transforming into berry pericarp while fertilized ovules develop into seeds within the two locules. Pinot Noir ovary anatomical sections reveal characteristic bilocular organization with two locules separated by central septum, ovules showing mature embryo sacs with egg cells ready for fertilization, and ovary wall showing multiple cell layers that will differentiate into skin (exocarp), flesh (mesocarp), and seed coat tissues during berry development. Merlot experiencing heat stress (38-42°C) during BBCH 55-56 (megasporogenesis) shows ovaries with reduced viable ovule numbers (1.8-2.6 ovules per ovary versus normal 3.2-3.8) due to heat-induced ovule abortion, resulting in reduced seed number per berry (1.2-2.1 seeds versus normal 2.4-3.2) and potential reduction in fruit set percentage through complete ovule failure in some flowers.
An ovary is the basal enlarged portion of the pistil containing ovules (immature seeds) within one or more internal chambers (locules), functioning as the site of ovule protection, fertilization, and post-fertilization development, ultimately transforming into the berry (fruit) following successful pollination and fertilization. The ovary structure in Vitis comprises= (1) ovary wall (pericarp precursor) typically 0.6-1.2 mm in diameter and 0.5-1 mm in height at anthesis, consisting of multiple cell layers including outer epidermis, middle parenchymatous mesocarp precursor tissue, and inner epidermis lining the locule(s) (2) internal organization typically showing two locules (chambers) separated by a central septum (partition wall) formed from fused carpel margins, representing the characteristic bilocular organization of the bicarpellate Vitis pistil (3) ovules typically numbering 2-4 total (commonly 1-2 per locule), attached to axile placentation on the central septum at positions corresponding to the fused margins of the two constituent carpels (4) vascular tissue providing nutrient and water supply, with vascular bundles running through the ovary wall and septum to supply developing ovules (5) superior position, meaning the ovary is positioned above the attachment points of sepals, petals, and stamens on the receptacle, contrasting with inferior ovaries (positioned below other floral organs) found in many other plant families. Ovary development proceeds through distinct phases= (1) primordium initiation during early flower bud development (BBCH 51-53) as part of pistil primordium differentiation (2) locule differentiation and septum formation (BBCH 53-54) creating the bilocular organization (3) ovule initiation (BBCH 54-55) with ovule primordia arising from placental tissue on the septum (4) megasporogenesis (BBCH 55-57) involving meiotic divisions in ovule megaspore mother cells producing haploid megaspores that develop into embryo sacs containing egg cells (5) ovary maturation (BBCH 57-59) with ovules achieving fertilization competence and ovary wall preparing for post-fertilization expansion (6) post-fertilization transformation (BBCH 71-89) involving dramatic ovary expansion, cell division and expansion in pericarp tissues, seed development within ovary chambers, and transformation into the mature berry. Ovary size at anthesis varies among flower types= hermaphrodite flowers typically show ovaries 0.7-1.1 mm diameter functionally female and pistillate flowers often exhibit slightly larger ovaries 0.9-1.3 mm diameter reflecting enhanced female organ investment functionally male flowers show reduced ovaries 0.4-0.8 mm diameter staminate flowers exhibit vestigial or rudimentary ovaries 0.2-0.6 mm diameter with aborted or absent ovules. Ovule number within ovaries shows cultivar-specific variation= most wine cultivars contain 3-4 ovules (commonly 2 per locule) some cultivars consistently show 2 ovules (1 per locule) rare individuals or cultivars may show 5-6 ovules through supernumerary ovule development. The ovary represents the critical structure determining fruit set potential and ultimate berry and seed characteristics, with ovary functionality (ovule viability, receptivity to fertilization, capacity for post-fertilization development) absolutely essential for successful reproduction and crop production. Environmental stresses during ovary development can impair function= heat stress >35°C during megasporogenesis (BBCH 55-57) can cause ovule abortion reducing viable ovule number by 30-60% water stress during ovary development can reduce ovary size and ovule number nutritional limitations (particularly boron deficiency) can impair ovule development and fertilization success.
Ovary represents the basal ovule-containing portion of the pistil and must be distinguished from the complete Pistil (which includes ovary plus style plus stigma), from individual Ovules (structures within the ovary), and from the Berry (mature fruit developing from the ovary post-fertilization). The ovary is the critical structure determining seed number potential and is the precursor to all berry tissues (skin, flesh, seeds). Sub-classification addresses ovary position (Superior_Ovary), internal organization (Bilocular_Ovary), and ovule content (Ovule_Containing_Ovary with specific ovule numbers). Understanding ovary anatomy, development, and function is essential for fruit set biology, seedlessness breeding, and berry development studies.
Fruchtknoten
Ovar
Samengehäuse
berry precursor
seed vessel
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 99-108 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 92-106 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 186-198.
Ovary
Merlot in Bordeaux shows ovary approach during early-mid June Year N+1 at BBCH 60-65, with pollen tubes completing 18-24 mm style navigation reaching ovary region containing 3-4 ovules, tubes responding to ovule-derived attractants navigating final 1.5-2.5 mm to micropyles over 2-5 hours, 2-3 tubes successfully targeting micropyles per flower, under oceanic conditions of 21-26°C temperature, typical of this mid-season cultivar completing approach 20-32 hours post-pollination enabling fertilization and 54-64% fruit set
Ovary approach represents the culminating tube growth phase when pollen tube completes style navigation reaching ovary region, responds to ovule-derived attractant signals directing final growth toward micropyle, and positions for embryo sac entry enabling sperm cell delivery and double fertilization. This critical final navigation phase occurs during pollen tube growth in early-mid June in temperate regions Year N+1, typically proceeding over final 2-6 hours of tube growth following style traverse as ovary region reached, corresponding to BBCH 60-65 flowering stages. Approach mechanism involves tube completing transmitting tissue navigation reaching style base adjacent to ovary, exiting transmitting tract entering ovary locule space containing 2-4 ovules, responding to ovule-derived chemoattractant signals directing growth toward specific ovule micropyle, and final precise navigation to micropyle opening enabling embryo sac access. Ovary structure at approach shows bicarpellate superior ovary containing typically 2-4 ovules distributed in two locules, ovules with anatropous orientation having micropyle positioned near ovary base, and locule space providing open cavity for tube growth from style base to ovule surface. Ovule characteristics include integuments surrounding nucellus forming outer seed coat layers, nucellus containing megagametophyte embryo sac with egg cell and central cell, micropyle representing opening in integuments providing access pathway to embryo sac, and funiculus attaching ovule to placenta. Tube exit from transmitting tissue shows growth transitioning from extracellular matrix in style to open locule cavity in ovary, tube entering ovary space near style base, and beginning final approach toward ovule micropyles requiring 1-3 mm additional growth. Ovule-derived attractants directing approach include diffusible signals emanating from micropyle creating concentration gradient, possible attractant molecules including LURE peptides identified in other species though specific signals in grapevine unknown, calcium gradients possibly contributing to guidance, and integrated signal profile directing tube to appropriate target. Chemotropic tube response shows sensitive detection of attractant gradient with femtomolar to picomolar sensitivity in some systems, directional growth reorienting toward highest attractant concentration, and precise navigation capability enabling targeting of 100-200 micrometer micropyle opening from millimeter distances. Multiple ovule targeting shows 2-4 ovules per ovary potentially receiving tubes, competitive dynamics with 2-5 tubes typically reaching ovary but usually only 1-2 achieving fertilization, and possible preferential targeting of specific ovules though mechanisms unclear. Tube growth rate during approach shows sustained or slightly reduced velocity 1-2 mm per hour during final ovary navigation, brief 1-3 mm growth distance from style base to ovule micropyle, and total approach duration typically 1-4 hours completing final stage of 12-36 hour total tube growth journey. Micropyle targeting precision shows remarkable accuracy with tube navigating directly to 100-200 micrometer micropyle opening, growth reorientation responding to attractant gradient, and successful targeting despite presence of multiple potential targets and complex three-dimensional ovary geometry. Tube-ovule contact shows tube apex reaching micropyle outer opening, possible adhesion to micropyle tissue, and preparation for penetration through micropyle canal to embryo sac requiring final 100-300 micrometer growth. Micropyle penetration following approach involves tube growing through micropyle canal between integument layers, navigating 100-300 micrometer passage to nucellus surface, and reaching embryo sac enabling sperm cell discharge into female gametophyte. Embryo sac structure at tube arrival includes seven-celled eight-nucleate organization with egg cell, two synergid cells flanking egg, central cell with two polar nuclei, and three antipodal cells, with synergids providing tube attraction and reception. Tube discharge preparation shows sperm cells positioned near tube tip preparing for release, vegetative nucleus degenerating as fertilization approaches, and tube apex preparing to rupture delivering sperm cells into embryo sac. Timing coordination shows tube arrival typically 18-40 hours after initial pollination depending on conditions, embryo sac maturity synchronized to receive tube at appropriate developmental stage, and temporal coordination essential for successful fertilization. Multiple tube arrivals show 1-4 tubes potentially reaching ovary region, competitive access to limited ovules with typically only 1 tube per ovule succeeding, and excess tubes unable to fertilize already-fertilized ovules becoming reproductively irrelevant. Ovule selectivity mechanisms possibly include first-arrival advantage with earliest tube to reach micropyle gaining access, ovule signaling changes after fertilization preventing additional tubes, or tube-tube interactions affecting competitive success, though specific mechanisms poorly understood. Failed approach causes include inadequate tube vigor preventing final navigation, absence of appropriate attractant signals from ovule dysfunction, guidance failure causing misdirected growth missing micropyle, structural abnormalities in ovary or ovule preventing access, or competition from faster tubes monopolizing available ovules. Environmental effects during approach show continued temperature sensitivity with optimal 22-28°C supporting efficient final navigation, and continued boron requirement for tube integrity during final growth phase. Approach success rate shows 40-70% flowers achieving successful tube arrival at ovule and fertilization under favorable conditions, reflecting cumulative effects of pollination efficiency, germination success, tube growth vigor, and competitive dynamics. The process represents third and final aspect of pollen tube growth culminating navigation, following style penetration and transmitting tissue navigation. Duration typically spans 1-4 hours representing final navigation phase after 10-30 hour style traverse. Cultivar variation in approach may include ovary structure affecting navigation complexity, ovule number affecting targeting opportunities, and attractant signaling affecting guidance efficiency. Regional timing shows ovary approach occurring during flowering period across all climates 18-40 hours after initial pollination as final tube growth phase completes. Viticultural significance includes representing culmination of tube growth enabling fertilization, requiring successful completion of entire pollination-germination-growth sequence, demonstrating critical importance of coordinated reproductive processes, and establishing approach success as final determinant of fertilization achievement and fruit set. Research applications include analyzing ovule attractant signals and chemotropic guidance mechanisms, investigating tube-ovule interactions and targeting precision, evaluating competitive dynamics among multiple tubes, quantifying approach timing and success rates, and assessing ovary approach as final determinant of fertilization outcomes and fruit set.
Ovary_Approach is third and final aspect of Pollen_Tube_Growth, distinguished by culminating navigation to ovule micropyles. Represents final 1-4 hours and 1-3 mm of total 12-36 hour growth journey. Involves chemotropic response to ovule-derived attractants. Precise micropyle targeting despite complex ovary geometry. Competitive dynamics with typically 1-2 tubes succeeding per flower. Completes entire Pollination → Germination → Tube Growth sequence. Directly enables Fertilization upon micropyle penetration. Final determinant of reproductive success.
Ovar-Annäherung
Samenanlage-Annäherung
final tube navigation
finale Schlauchnavigation
micropyle targeting
ovary arrival
ovule approach
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 388-395 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 293-298 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Ovary Approach
Merlot in Bordeaux shows ovary expansion initiation during early July Year N+1 at BBCH 69-71, with fertilized ovaries expanding from 5-7 mm to 14-18 mm diameter over 18-26 days, peak cell division intensity with 12-14 hour cycle times during weeks 2-3, cell expansion through water uptake contributing to volumetric increase, under oceanic conditions of 20-26°C temperature, typical of this mid-season cultivar completing initial expansion establishing berry developmental trajectory
Ovary expansion initiation represents the rapid growth phase following post-fertilization signaling when auxin-triggered responses activate cell division and expansion in ovary wall pericarp, transforming static fertilized ovary from 4-8 mm diameter into visibly enlarging berry reaching 10-20 mm within 2-3 weeks, establishing irreversible growth trajectory and confirming successful fruit set. This dramatic growth phase occurs during early fruit development in late June-early July in temperate regions Year N+1, proceeding over 1-3 weeks following fertilization as growth accelerates, corresponding to BBCH 69-71 stages. Expansion mechanism involves auxin-triggered cell cycle reentry in quiescent pericarp cells activating mitotic division, cell expansion through water uptake and wall loosening increasing cell volume, coordinated tissue growth across all pericarp layers, vascular development supporting resource delivery, and overall rapid volumetric increase 8-20 fold during first month post-fertilization. Initial ovary size shows pre-fertilization diameter typically 4-8 mm in grapevine flowers, static size during flowering period, and expansion beginning within days of successful fertilization as hormone signaling establishes. Expansion dynamics show initial slower growth during first week as signaling establishes and cells activate, acceleration during second and third weeks with peak expansion rates, volumetric increase typically reaching 10-20 mm diameter within 2-3 weeks, and 8-20 fold volume increase during first month representing dramatic transformation. Growth rate characteristics show peak expansion rate 1-3 mm diameter increase per week during maximum growth phase, linear or exponential growth patterns depending on measurement and timing, temperature-dependent rate with warm conditions accelerating expansion, and growth rate eventually declining as cell division phase completes transitioning to slower expansion-dominated phase. Cell division contribution to expansion shows mitotic activity initiating in pericarp cells within days of auxin signal, peak division intensity during first 2-4 weeks with cycle times 10-15 hours, progressive cell number increase creating berry flesh cellularity, final cell number established within 3-4 weeks determining potential berry size, and division phase completion marking transition to expansion-dominated growth. Cell division spatial patterns show divisions throughout pericarp layers including exocarp skin precursors, abundant divisions in mesocarp creating bulk flesh tissue, divisions in endocarp adjacent to seeds, and coordinated division across layers producing organized tissue structure. Cell expansion contribution shows cells enlarging through water uptake driven by osmotic gradient, cell wall loosening enabling expansion through enzymatic modification, turgor pressure driving cell enlargement against yielding walls, and progressive cell volume increase contributing substantially to berry growth especially after division phase. Water uptake dynamics show xylem-mediated water import during early growth, osmotic gradient from solute accumulation driving water influx, and water as major component of expanding berry typically 75-85% fresh weight. Cell wall modification enabling expansion shows enzymatic loosening through expansins and other wall-modifying proteins, pectin and hemicellulose degradation creating wall plasticity, cellulose microfibril reorientation allowing expansion, and new wall synthesis maintaining integrity during enlargement. Tissue organization during expansion shows coordinated growth maintaining berry structural integrity, proportional expansion across pericarp layers, vascular tissue development keeping pace with berry enlargement, and overall organized growth producing functional berry architecture. Vascular development shows differentiation of vascular bundles within berry providing nutrient and water transport pathways, xylem vessel formation enabling water delivery supporting expansion, phloem development preparing for sugar import during ripening, and branching vascular network establishing throughout pericarp. Metabolic support for expansion shows increased respiratory rate providing ATP for biosynthesis and active transport, carbohydrate import from vine providing energy and building materials, nitrogen uptake supporting protein synthesis for new cells and membranes, and overall metabolic intensification characteristic of rapid growth. Resource import pathways show vascular connections from pedicel delivering water, nutrients, and photosynthates, developing berry as strong sink attracting resources, and import capacity increasing with berry size and metabolic demand. Visible expansion confirmation shows berry diameter measurably increasing over days to weeks, green berry clearly distinguishable from static unfertilized flowers, and visible growth providing definitive fruit set confirmation. Berry shape establishment shows spherical to ovoid morphology developing during expansion, cultivar-characteristic shape emerging during early growth, and shape largely determined during expansion phase. Environmental effects on expansion include temperature critically affecting growth rate with optimal 22-28°C supporting rapid expansion, water availability affecting cell expansion and turgor, nutrient supply supporting tissue synthesis, and overall favorable conditions during critical 2-3 week period maximizing expansion. Temperature effects show warm conditions promoting rapid expansion completing 10-20 mm growth within 2 weeks, cool conditions slowing expansion potentially requiring 3-4 weeks, and temperature as major factor affecting expansion rate and timing. Water relations show adequate water essential for cell expansion and turgor maintenance, water deficit potentially limiting berry enlargement, and irrigation sometimes beneficial during expansion in water-limited environments. Failed or delayed expansion causes include inadequate hormone signaling from poor seed development, environmental stress during critical period limiting growth capacity, water or nutrient deficiency preventing growth response, or physiological dysfunction though rare after successful signaling. Expansion as fruit set indicator shows successful expansion confirming fruit set, measurable growth distinguishing set berries from aborted flowers, and expansion observation used for early yield estimation. The process represents second phase of berry initiation showing growth response to hormonal signals, following post-fertilization signaling and preceding early berry formation. Duration typically 1-3 weeks of rapid initial expansion though growth continues at declining rate. Cultivar variation includes expansion rate, final berry size potential, and temperature sensitivity. Regional timing shows ovary expansion initiation during late June-early July in warm climates, early-mid July in temperate regions, and mid-late July in cool areas. Viticultural significance includes providing visible fruit set confirmation, determining berry size through cell number and expansion, demonstrating critical temperature and water sensitivity, and establishing berry developmental trajectory affecting final yield and quality. Research applications include analyzing expansion mechanisms and growth dynamics, investigating cell division and expansion coordination, evaluating environmental effects on expansion rate and success, quantifying growth patterns and berry size determinants, and assessing expansion as fruit set and quality determinant.
Ovary_Expansion_Initiation is second phase of Berry_Initiation, distinguished by rapid growth response to hormonal signals. Represents 8-20 fold volumetric increase over 1-3 weeks from 4-8mm to 10-20mm diameter. Peak cell division 10-15h cycles establishing final cell number in 3-4 weeks. Temperature strongly affects rate. Water essential for expansion. Visible growth confirms fruit set. Follows Post_Fertilization_Signaling. Precedes Early_Berry_Formation. Determines berry size potential through cell number establishment.
Ovar-Expansions-Initiierung
Ovarvergrößerung
berry expansion start
initial berry growth
ovary enlargement
rapid berry growth
schnelles Beerenwachstum
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 418-422 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Ovary Expansion Initiation
Over ripening tolerance quality comparison in McLaren Vale evaluated overall quality maintenance during three-week extended hangtime for Shiraz and Grenache under equivalent conditions. Shiraz demonstrated excellent over ripening tolerance quality with stable sugar plateau at 26°Brix, moderate acid decline maintaining pH below 3.7, aromatic evolution toward desirable spice and dark fruit complexity without excessive jamminess, beneficial tannin softening improving approachability, and thick-skinned resistance to Botrytis despite autumn rains, with wines from fruit harvested three weeks post-optimal receiving equivalent quality scores to optimal-date harvest though with stylistic shift toward richer, more concentrated expression. Grenache exhibited poor over ripening tolerance quality with continued sugar accumulation reaching 28°Brix and potential alcohol exceeding 16%, rapid acid decline with pH rising to 3.9 creating microbial instability risk, aromatic degradation losing varietal character toward generic dried fruit notes, and thin-skinned susceptibility to Botrytis causing 15% bunch rot incidence by the third week, with wines from delayed harvest receiving significantly lower scores reflecting the integrated quality decline, establishing Grenache as requiring strict harvest timing discipline while Shiraz tolerated flexibility.
Over ripening tolerance quality is a post optimal decline quality that inheres in a grapevine cultivar or clone within a specific growing environment and represents the overall capacity to maintain acceptable quality characteristics despite harvest occurring substantially beyond the optimal date, integrating the effects of continued sugar accumulation, acid decline, aromatic evolution, phenolic modification, and physical deterioration into a composite assessment of how gracefully the cultivar ages on the vine when extended hangtime proves necessary or is deliberately employed for stylistic purposes. This quality provides integrative characterization complementing the component-specific qualities addressing individual parameter trajectories, capturing the net quality outcome when all post-optimal changes are considered together and evaluated against specific wine style objectives. The ontological significance of over ripening tolerance quality lies in its practical value for harvest planning and risk assessment, enabling straightforward comparison of cultivar vulnerability to harvest delays and supporting scheduling decisions that prioritize sensitive varieties while permitting greater flexibility for tolerant types that maintain quality across extended timing ranges. The biological basis of over ripening tolerance quality integrates the same underlying processes addressed by component qualities including sugar and acid dynamics, aromatic stability, phenolic evolution, and physical deterioration, but evaluates their combined effect on wine quality potential rather than tracking individual parameters. The assessment framework must account for the reality that different post-optimal changes may partially compensate or synergistically combine, with for example the concentration effects of mild dehydration potentially enhancing phenolic intensity even as aromatic freshness declines, creating quality outcomes that differ from simple summation of individual parameter trajectories. Cultivar characteristics favoring high over ripening tolerance quality include a combination of stable sugar plateau, slow acid decline, resilient aromatic compounds, beneficial phenolic evolution, resistant physical structure, and disease resistance preventing pathogen-mediated deterioration, with the interaction among these factors determining overall tolerance. Assessment of over ripening tolerance quality involves systematic comparison of wine outcomes across harvest timing treatments, with evaluation against style-specific quality criteria recognizing that the same post-optimal evolution may prove detrimental for one wine style while beneficial for another.
Over_Ripening_Tolerance_Quality is_a Post_Optimal_Decline_Quality, representing the integrated quality maintenance during extended hangtime. This class provides composite assessment complementing the component-specific qualities (Sugar, Acid, Aroma, etc.) by evaluating overall wine quality outcomes. Style-specific evaluation criteria are essential since post-optimal changes may benefit some styles while compromising others. Cultivars exhibiting simultaneous favorable characteristics across multiple dimensions demonstrate highest tolerance. Critical for harvest scheduling prioritization and risk assessment when weather or logistics may delay harvest beyond optimal timing.
Hängezeittoleranz
Spätlese-Resilienz
delayed harvest adaptation
extended hangtime tolerance
late-harvest quality maintenance
over-maturity resilience
Überreife-Anpassungsfähigkeit
Überreifungstoleranz
Bindon KA et al. (2014) Relationships between harvest time and wine composition in Vitis vinifera L. cv. Cabernet Sauvignon. Food Chemistry 154:43-54 Coombe BG, Iland P (2004) Grape berry development and winegrape quality. In: Dry PR, Coombe BG (eds) Viticulture Volume 1 Resources, 2nd Edition, Winetitles, Adelaide, pp. 210-248 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 509-547
Over Ripening Tolerance Quality
Overall assessment for Riesling Grand Cru Rangen from Alsace (2022 vintage) synthesizes the visual (deep gold with green reflections), olfactory (intense smoky-mineral, citrus, and white flower complexity), and gustatory evidence (bone-dry, full body, razor-sharp acidity, extraordinarily long mineral finish) into a judgment of exceptional quality — balance rated as outstanding (the extreme acidity counterpoised by equally extreme concentration and extract), typicity assessed as exemplary (expressing the volcanic terroir signature of the Rangen vineyard with unmistakable clarity), quality rated at 96 points on the 100-point scale, and aging potential projected at 20-40 years with an optimal window of 8-25 years post-vintage. Overall assessment for Blaufränkisch Classic from Mittelburgenland (2024 vintage) integrates a more modest but entirely satisfying evaluation — good balance between fruit, acid, and gentle tannin, high typicity for the regional style, quality rated at 87 points, and aging potential of 3-6 years with immediate accessibility.
Overall assessment is a sensory description entity that documents the integrative quality judgment of a wine derived from the synthesis of visual, olfactory, and gustatory evaluations into a holistic appraisal, encompassing balance (the harmonious interplay of structural elements), typicity (the degree of conformity to expected regional and varietal character), quality rating (the numerical or categorical quality score), and aging potential (the projected developmental trajectory and optimal drinking window), and representing the concluding evaluative synthesis where trained tasters distill the accumulated perceptual evidence from all sensory modalities into a single coherent quality verdict that informs commercial valuation, critical reputation, and consumer purchasing decisions.
Overall assessment represents the pinnacle of the sensory evaluation hierarchy, where the reductionist analysis of individual sensory parameters (color, clarity, aroma descriptors, sweetness, acidity, tannin, body, finish) is re-integrated into a holistic quality judgment that involves cognitive processes extending beyond sensory perception into aesthetic evaluation, contextual knowledge, and experiential comparison. The distinction between analytical tasting (the systematic parameter-by-parameter assessment documented in the preceding sensory description classes) and synthetic tasting (the integrative quality judgment documented here) reflects a fundamental cognitive shift — from decomposition of a complex stimulus into identifiable components to the reconstruction of those components into a unified quality gestalt. Research in wine expertise demonstrates that experienced tasters process wine quality assessment through both bottom-up (data-driven, from sensory input to quality conclusion) and top-down (knowledge-driven, from expectations to confirmatory assessment) pathways, with the relative contribution of each pathway varying by taster expertise and wine familiarity. The four sub-dimensions of overall assessment — balance, typicity, quality rating, and aging potential — represent distinct evaluative frameworks that may occasionally conflict (a wine may achieve excellent balance but poor typicity if its style diverges from regional expectations, or high quality but limited aging potential if designed for early consumption), requiring the assessor to weight competing quality criteria according to the evaluation context. The institutional frameworks for wine quality assessment — competition panels, critical scoring systems, appellation approval tastings, classification reviews — each apply different weightings to these sub-dimensions, meaning that the same wine may receive divergent overall assessments depending on the evaluative context and criteria applied.
Gesamtbeurteilung
Gesamtbewertung
Gesamteindruck
Qualitätsurteil
abschließende Bewertung
comprehensive evaluation
final assessment
overall quality assessment
quality conclusion
wine quality judgment
Peynaud E, Blouin J (2006) The Taste of Wine: The Art and Science of Wine Appreciation, 4th Ed, Wiley, pp.: 280-320 Lawless HT, Heymann H (2010) Sensory Evaluation of Food, 2nd Ed, Springer, pp.: 227-260 Parr WV et al. (2002) Demystifying wine expertise: olfactory threshold, perceptual skill and semantic memory in expert and novice wine judges. Chemical Senses 27(8): 747-755
Overall Assessment
Berry overall form in Chasselas transitions during ripening from slightly ellipsoid at véraison (length-to-width ratio 1.05-1.10) to essentially globose at harvest maturity (ratio 0.98-1.02) as the equatorial diameter catches up with polar diameter during the cell expansion phase, a developmental pattern common to many cultivars but particularly pronounced in Chasselas and contributing to its characteristic round berry appearance valued in table grape markets of Switzerland and the Lake Geneva region.
Overall_Form_Quality represents the general three-dimensional geometric configuration of grapevine organs as characterized by the relationship between their principal axes and the curvature of their bounding surfaces. This quality captures the fundamental shape category into which an organ falls when viewed as a complete solid body, abstracting from details of surface texture, apex and base configuration, or marginal features. For berries, overall form is one of the most important ampelographic characters, with cultivars classified into distinct categories including globose (spherical), ellipsoid (elongated with rounded ends), ovoid (egg-shaped), cylindrical, and other forms. Cluster overall form similarly distinguishes conical, cylindrical, winged, and branched architectures that affect compactness and disease susceptibility. Leaf blade overall form, while more complex due to lobing, can be characterized as orbicular, cuneiform, reniform, or cordate. Overall form results from the integrated pattern of cell division and expansion during organ development, with genetic factors establishing the basic form and environmental conditions modulating expression within limits. The principal axes of an organ (length, width, depth) and their ratios provide quantitative descriptors of overall form, while categorical descriptors following OIV protocols enable standardized visual assessment. Overall form affects functional properties including aerodynamic behavior, light interception geometry, and hydrodynamic characteristics relevant to spray deposition and drying.
Overall_Form_Quality serves as the parent class for specific geometric form categories including Globose_Quality, Ellipsoid_Quality, Ovoid_Quality, Cylindrical_Quality, and Conical_Quality. These child classes represent discrete form categories used in ampelographic description, though actual organs may exhibit intermediate forms. The assignment of an organ to a form category involves judgment when forms are intermediate quantitative shape indices provide more precise characterization. Overall form assessment should specify the developmental stage and viewing orientation for reproducibility.
Gesamtform
Grundform
allgemeine Gestalt
basic form
dreidimensionale Konfiguration
general shape
gross morphology
three-dimensional configuration
Galet P (1979) A Practical Ampelography, Cornell University Press pp.: 54-78 OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 83-87
Overall Form Quality
Overall taste harmony assessment in blind tasting of Bordeaux across vintages reveals harmony ratings of 8.5 for 2015 (warm year, ripe fruit balancing structured tannins), 7.2 for 2013 (cool year, elevated acidity not fully integrated with lean fruit), and 6.1 for 2003 (hot year, overripe sweetness overwhelming diminished acidity), demonstrating how vintage climate affects the integration of all taste components into harmonious whole, with harmony scores correlating r=0.87 with critic quality scores and r=0.78 with auction prices, confirming overall harmony as a primary quality determinant.
Overall taste harmony quality is a taste balance quality that represents the holistic perception of integration among all taste components simultaneously, characterizing the gestalt impression of taste coherence that emerges from the combined assessment of sweet-sour, sweet-bitter, sour-bitter, and all other taste interactions operating together. Within the BFO framework, this quality addresses the highest level of taste integration, distinguishing overall harmony from bilateral balance relationships while recognizing that holistic harmony perception may differ from the sum of pairwise balance assessments when multiple components interact in complex ways. The ontological significance lies in overall taste harmony serving as the ultimate taste quality criterion, representing the integrated judgment that expert tasters make when assessing whether a wine's taste structure is complete, coherent, and satisfying as a unified sensory experience rather than a collection of separately perceived attributes. The perceptual basis for overall harmony involves central integration of multiple taste inputs, comparison against learned templates of quality and style expectations, and hedonic response to the combined taste experience, creating a judgment that is greater than the analytical sum of component assessments. The compositional complexity underlying overall harmony includes not only the primary taste compounds (sugars, acids, phenolics) but their ratios, the timing of their perception, and matrix factors affecting their integration, with the multiplicity of interactions creating emergent properties not predictable from individual components. The style-dependency of harmony standards means that the optimal integration pattern differs among wine types, with dry wines requiring different harmony than sweet wines, light whites different from full reds, and young wines different from aged, reflecting learned expectations for each category. In wine quality assessment, overall taste harmony often serves as the summary judgment integrating detailed analytical evaluation, with high harmony scores correlating with quality tier and consumer preference across diverse wine styles. Understanding overall taste harmony quality enables holistic quality assessment complementing analytical component evaluation, winemaking optimization toward integrated taste structure, and communication of quality in terms that capture the complete taste experience.
Overall_Taste_Harmony_Quality represents the highest integrative level of taste balance assessment, encompassing all pairwise balance relationships and their combined effect. This quality is inherently subjective and style-dependent, requiring context specification for meaningful evaluation. The class represents the taste component of overall wine quality, which additionally includes aroma, mouthfeel, and visual elements for complete sensory assessment.
Gesamtgeschmacksharmonie
Geschmacksintegration
Geschmacksvollständigkeit
gustatorische Kohärenz
gustatory coherence
overall taste quality
taste completeness
taste integration
Lawless HT, Heymann H (2010) Sensory Evaluation of Food: Principles and Practices, 2nd Ed. Springer, New York, pp. 325-367 Parr WV et al. (2002) The distinctive flavour of New Zealand Sauvignon blanc: sensory characterization by wine professionals. Food Quality and Preference 13(7-8): 505-514
Overall Taste Harmony Quality
An overlapping petiolar sinus on a mature leaf at node 7 of Vitis vinifera cv. Pinot Noir in Burgundy, France, exhibiting characteristic extreme overlapping configuration with left basal lobe margin crossing anterior to right lobe margin by approximately 0.8 cm creating complete occlusion of sinus and petiole attachment, sinus depth of 1.5 cm measured to the underlying junction point, V-shaped underlying contour completely masked by overlapping lobes, basal lobe margins converging and crossing at acute angle creating distinctive overlapped appearance diagnostic for Pinot varieties, lobe margin edges smooth in overlap zone, moderate to dense pubescence on lobe surfaces particularly in overlap region, completely concealing petiole attachment with manipulation required to observe insertion point, serving as absolutely definitive diagnostic feature enabling confident Pinot Noir identification distinguishing from all major cultivars except other Pinot variants (Gris, Blanc) which share this characteristic, measured at BBCH stage 77 (berries beginning to touch) in mid-July on well-developed leaf representing classic diagnostic expression, demonstrating the exceptional identification value of overlapping petiolar sinus as primary diagnostic character in this major cultivar group.
A petiolar sinus exhibiting extreme convergence with basal lobe margins crossing over each other at the petiole-lamina junction creating occluded or masked attachment zone in Vitis vinifera, characterized by actual overlap of lobe margins with one lobe edge crossing anterior to the other, complete occlusion of the sinus creating no visible gap or opening, typically V-shaped or narrow U-shaped underlying configuration with extreme lobe convergence producing the overlapping condition, completely hidden petiole attachment requiring leaf manipulation to observe petiole insertion, occurrence in several notable cultivars including Pinot Noir, Pinot Gris, Pinot Blanc (all showing characteristic overlapping sinus), Gamay, and some Muscat types, exceptional diagnostic value as overlapping sinus is relatively rare and highly distinctive enabling confident identification when present, and potential functional implications including maximum mechanical integration at the petiole-lamina junction though lacking experimental verification. Overlapping petiolar sinuses develop through extreme convergent growth of basal lobe primordia during leaf development, with genetic programming directing not merely inward growth but actual crossing of lobe margins, creating the distinctive overlapped configuration that represents the maximum expression of closed sinus architecture and provides the most distinctive and easily recognized petiolar sinus category.
Overlapping petiolar sinuses represent the most distinctive and easily recognized sinus category, distinguished from open sinuses by complete closure and from merely closed sinuses by actual crossing of lobe margins creating occlusion. The overlapping configuration is relatively rare compared to open and closed types, occurring in the important Pinot group (Noir, Gris, Blanc) and several other cultivars but absent in most commercial varieties, making its presence highly diagnostic. The degree of overlap varies from minimal crossing (margins barely overlap) to substantial overlap (one lobe crosses significantly anterior to the other by 1-2 cm), with greater overlap generally creating more obvious and recognizable expression. Genetic control appears strong with overlapping sinus showing consistent expression across Pinot variants suggesting shared genetic basis, and crosses involving Pinot often producing intermediate or overlapping sinuses supporting genetic determination. Environmental stability is generally high, with overlapping configuration maintained consistently across vigor and light conditions, making this one of the most reliable ampelographic characters. Functional implications remain speculative, with proposed mechanical advantages from complete closure and lobe integration lacking direct experimental support. Ampelographic utility is exceptional, with overlapping petiolar sinus serving as primary definitive diagnostic feature enabling confident identification often from considerable distance based on distinctive occluded appearance.
crossed lobe sinus
gekreuzte Basalbucht
occluded petiolar sinus
okkludierte Stielbucht
overlapped basal sinus
überlappender Petiolarsinus
Overlapping Petiolar Sinus
Merlot overripe berries at week 20 (2 weeks post-optimal harvest timing) show 27.5% sugars (excessive for balanced Merlot), 4.8 g/L titratable acidity (insufficient), pH 4.05 (excessively high), 3.2 mg/g anthocyanins (color stable but other quality compromised), cooked plum and prune flavors replacing fresh fruit character, very soft texture with beginning shriveling, producing wine with 16% alcohol, flabby structure, cooked fruit character, and compromised aging potential compared to optimal-harvest wine (24.5% sugars, 6.2 g/L acidity, pH 3.75, 14.5% alcohol, balanced structure, fresh fruit). Chardonnay overripe berries showing white wine overripeness consequences= 25% sugars (excessive for Chardonnay targeting 21-22%), 5.2 g/L acidity (very low), pH 3.68 (high for white wine), tropical fruit character becoming overblown and losing characteristic apple-citrus notes, producing wine with 14.5% alcohol (high for Chardonnay), flabby mouthfeel, oxidative tendency, loss of varietal typicity. Cabernet Sauvignon slightly overripe berries (week 19, 1 week past optimal)= 26% sugars, 5.5 g/L acidity, pH 3.85, representing moderate overripeness producing acceptable but suboptimal wine with 15% alcohol, slightly reduced acidity, beginning of cooked fruit notes, demonstrating that slight overripeness creates quality compromise but may be acceptable depending on style preferences. Syrah severely overripe berries (week 21, 3 weeks past optimal) showing advanced overripeness= 30% sugars, 4 g/L acidity, pH 4.15, significant berry shriveling with 15% weight loss from dehydration, prune and raisin flavors dominant, 5% Botrytis infection incidence on compromised overripe berries, producing wine with 17.5% alcohol, severe structural imbalances, off-flavors from rot, demonstrating severe quality consequences of extreme overripeness unsuitable for quality production. Pinot Noir climate comparison showing overripeness development= cool climate berries at week 19 achieving 24.5% sugars, 6.5 g/L acidity (optimal ripeness, not overripe) moderate climate at week 19 showing 27% sugars, 5 g/L acidity (overripe) warm climate at week 19 showing 29% sugars, 4.2 g/L acidity, beginning raisining (severely overripe), demonstrating climate effects on overripening tendency with warm climates more prone to overripeness requiring earlier harvest to avoid quality degradation.
An overripe berry is a mature berry that has progressed beyond optimal or peak ripeness, characterized by excessive sugar accumulation, excessive acid decline, potential flavor degradation, beginning dehydration or raisining, and compositional imbalances that may compromise quality for most wine styles, representing berries past ideal harvest timing. Overripe berries are characterized by= (1) sugars at elevated levels typically >26-28% for most wine cultivars (>30% in extreme cases), exceeding targets for balanced wine production and producing excessive alcohol potential (>15-16% potential alcohol) (2) titratable acidity at low levels typically 3.9-4.1 (>4.2 in severely overripe), creating microbiological instability risk and compromising wine structure and aging (4) potential flavor changes including loss of fresh fruit character, development of cooked, jammy, or raisined flavors, loss of varietal typicity, reduction in aromatic complexity (5) berry weight potentially declining from peak through dehydration with transpiration and respiration exceeding import, creating concentration through water loss (6) berry texture becoming very soft or beginning to shrivel with firmness 4), cooked or jammy fruit character lacking freshness, potential loss of varietal typicity, compromised balance and aging potential white wines showing flabby structure, oxidative character, loss of aromatic freshness. Overripe berries may be intentional for certain specialized styles= late-harvest dessert wines deliberately using overripe or raisined fruit for high sugar and concentrated flavors Amarone-style production using partially dried overripe grapes some warm-climate full-bodied red styles accepting moderate overripeness for concentrated character but for most premium table wine production, overripeness represents quality compromise. Overripe berries show varying severity= slightly overripe berries (26-27% sugars, 4.5-5.5 g/L acidity) may produce acceptable wine with some quality compromise moderately overripe berries (28-30% sugars, 3.5-4.5 g/L acidity) produce wines with significant compositional imbalances severely overripe or raisined berries (>30% sugars, <3.5 g/L acidity) are generally unsuitable for quality table wine production except in specialized styles. Environmental conditions influence overripening rate= warm temperatures accelerate sugar accumulation and acid decline hastening overripening dry conditions promote dehydration concentration humid conditions may promote rot development on overripe berries autumn rainfall can cause berry splitting in overripe berries.
Overripe_Berry represents post-optimal maturity state and is distinguished from Fully_Ripe_Berry (optimal quality) by excessive versus balanced composition. Overripeness generally compromises quality for balanced wine production but may be intentional for certain specialized styles. Sub-classification into Slightly and Severely overripe reflects degree of quality compromise. Understanding overripeness is important for avoiding harvest delays that compromise quality, for recognizing limits of extended hang time strategies, and for specialized production styles intentionally using overripe fruit.
Nachreife-Traube
over-mature berry
past-optimal berry
zu reife Traube
überreife Beere
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 245-260 Jackson, D.I., Lombard, P.B. (1993). Environmental and management practices affecting grape composition. American Journal of Enology and Viticulture 44=409-430.
Overripe Berry
Sémillon berries destined for Sauternes sweet wine production are intentionally allowed to progress to overripe softening quality with firmness below 1.0 N, facilitating Botrytis cinerea infection that concentrates sugars to 35-45°Brix while the fungal metabolism generates characteristic honey, apricot, and saffron aromas, transforming the degradative softening process into a quality-enhancing phenomenon through controlled noble rot development.
Overripe softening quality is a softening stage quality that inheres in a grape berry that has progressed beyond optimal harvest maturity into a degradative phase characterized by excessive firmness loss, compromised structural integrity, and quality deterioration resulting from advanced cell wall breakdown, membrane degradation, and often pathogen colonization. Within the BFO framework, this quality represents the terminal softening state where physiological processes transition from ripening to senescence, marked by textural changes that indicate quality decline rather than development. The ontological significance lies in distinguishing overripe softening from the desirable softening of normal ripening, as the underlying mechanisms shift from controlled enzymatic modification to uncontrolled degradation including advanced pectin depolymerization to monomeric sugars, membrane lipid peroxidation, and cell compartmentation failure leading to enzyme-substrate mixing that accelerates tissue breakdown. Instrumentally, overripe berries exhibit firmness values below cultivar-specific thresholds, typically less than 1.5 N, with characteristic loss of elastic recovery, increased permanent deformation, and often visible symptoms including shriveling, skin cracking, or pathogen mycelium. This quality state increases susceptibility to Botrytis cinerea colonization which, while devastating for table grapes, can be intentionally induced for noble rot sweet wine production where the fungal transformation of overripe tissue creates distinctive flavor compounds. The rate of progression into overripe softening depends on temperature, humidity, and varietal susceptibility, with some cultivars maintaining integrity well past optimal harvest while others deteriorate rapidly. Documentation of overripe softening supports harvest scheduling to minimize quality loss in large vineyard blocks requiring sequential picking.
Overripe_Softening_Quality represents the terminal state in the softening progression, ontologically distinguished by its degradative rather than developmental character. The boundary between Harvest_Firmness_Quality and Overripe_Softening_Quality is context-dependent, requiring specification of intended use since overripe states for dry wine production may represent optimal states for certain sweet wine styles. This class encompasses both undesirable quality loss and intentional over-ripening for specific production objectives.
Abbaufestigkeit
degradative softening
over-mature softening
post-optimal firmness loss
senescent berry texture
seneszente Beerentextur
Überreife-Erweichung
überreife Beerenfestigkeit
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1: The Microbiology of Wine and Vinifications, 2nd Ed. John Wiley & Sons, Chichester, pp. 193-217 Zsófi Z et al. (2014) The effect of the timing of water deficit on the berry texture and composition of Kékfrankos (Vitis vinifera L.) cultivar. Scientia Horticulturae 176: 20-27
Overripe Softening Quality
Italia (Ideal) table grapes exhibit pronounced ovoid berry form with the maximum diameter positioned distinctly toward the stylar end, creating the characteristic egg shape with length-to-width ratios of 1.30-1.45 that distinguishes this cultivar visually from rounder Muscat varieties this distinctive shape, combined with large berry size (8-12 g), crisp texture, and Muscat aroma, establishes Italia as a premium table grape commanding price premiums in European markets, particularly in Italy where it dominates late-season production.
Ovoid_Quality represents the overall form characteristic of organs exhibiting egg-shaped geometry with asymmetric curvature along the principal axis, typically broader at one end and tapering toward the other. In grape berries, ovoid form combines elongation (length exceeding width) with polarity (proximal-distal asymmetry), producing a shape reminiscent of a bird's egg. The broader end may be either proximal (toward the pedicel attachment, termed obovoid) or distal (toward the stylar scar, termed ovoid in the strict sense), with the latter being more common in grapes. Ovoid-berried cultivars include Italia, Regina, and certain Muscat varieties, often associated with table grape production where the distinctive shape contributes to visual appeal. The asymmetric form results from differential expansion patterns during berry development, with the broader region exhibiting greater cell enlargement than the tapering region. Ovoid berries typically display length-to-width ratios of 1.15-1.45, overlapping with ellipsoid values, but distinguished by the asymmetry between poles. Ovoid cluster form describes bunches that taper from a broad shoulder region toward a narrower tip, a common architecture that accommodates variable fruit set along the rachis. Assessment of ovoid form requires evaluation of both the length-to-width ratio and the position of maximum width along the polar axis, with maximum width displaced from the equator toward one pole indicating ovoid form.
Ovoid_Quality is distinguished from Ellipsoid_Quality by the asymmetric curvature between the two poles. The direction of the asymmetry (narrower toward pedicel or toward stylar end) may be specified as obovoid versus ovoid respectively, following botanical convention. OIV descriptor 223 includes ovoid (category 4) and obovoid (category 5) as separate shape categories. The assessment of ovoid form requires longitudinal profile evaluation rather than simple diameter measurements digital image analysis enables objective quantification of the asymmetry. PATO_0001866 (ovate) provides a related upper-ontology concept.
Eiform
asymmetric oval
egg-shaped form
eiförmige Qualität
obovate form
ovale asymmetrische Form
pyriform quality
tropfenförmige Gestalt
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 83-84 Crespan M (2004) Evidence on the Evolution of Polymorphism of Microsatellite Markers in Varieties of Vitis vinifera L. Theoretical and Applied Genetics 108(2): 231-237
Ovoid Quality
Chardonnay hermaphrodite flowers at BBCH 59-61 exhibit ovule containing ovaries with 4 ovules (2 per locule) in 72% of flowers, 3 ovules (variable 2+1 distribution) in 22% of flowers, and 2 ovules (1 per locule) in 6% of flowers, with ovules showing mature embryo sacs ready for fertilization at anthesis, achieving 48-56% fruit set with successful flowers producing berries containing 2.2-3.4 seeds on average reflecting fertilization of 55-85% of available ovules. Gewürztraminer functionally female flowers show ovule containing ovaries with 3-4 ovules (mean 3.7 ovules per ovary) positioned in bilocular organization on septum surfaces, requiring cross-pollination from hermaphrodite pollinizers and achieving 28-38% fruit set with successful berries containing 1.8-2.9 seeds representing fertilization of 49-78% of available ovules. Thompson Seedless stenospermocarpic cultivar produces ovule containing ovaries with 3-4 ovules that undergo fertilization initiating seed development, but seeds abort during BBCH 71-73 (3-6 weeks post-fertilization) creating seedless berries with only rudimentary seed traces, demonstrating that ovule presence does not guarantee mature seed production when genetic factors cause seed abortion. Wild V. vinifera ssp. sylvestris pistillate female flowers exhibit ovule containing ovaries with 3-4 viable ovules receiving cross-pollination from staminate males and achieving 30-42% fruit set producing berries with 2.1-3.2 seeds, while staminate male flowers show vestigial ovaries containing 0-2 aborted ovules incapable of fertilization and achieving <1% fruit set, demonstrating the critical importance of viable ovule presence for successful reproduction.
An ovule containing ovary is an ovary that contains one or more ovules (immature seeds) within its locule(s), representing the functional female reproductive state where ovules are present and available for fertilization, contrasting with rare aberrant ovaries completely lacking ovules or with vestigial/rudimentary ovaries in functionally male or staminate flowers where ovules are aborted or absent. Ovule containing ovaries are characterized by= (1) presence of ovules attached to placental tissue on the ovary septum (in bilocular ovaries) or ovary wall (in rare unilocular ovaries) (2) ovule number typically 2-4 per ovary in most Vitis flowers, with common distributions being 4 ovules (2 per locule), 3 ovules (variable distributions of 2+1 or 1+2 per locule), or 2 ovules (1 per locule) (3) ovules showing mature structure at anthesis (BBCH 61-65) with completed megasporogenesis and mature embryo sacs containing egg cells ready for fertilization (4) ovules positioned in axile placentation on the septum surfaces in the characteristic bilocular organization (5) ovules receiving vascular supply from septal vascular bundles providing nutrients and water supporting ovule development and post-fertilization seed maturation. Ovule containing ovaries represent the normal and essential condition for fruit set and seed production, as ovules are the structures that receive pollen tube delivery of sperm cells, undergo fertilization producing embryo and endosperm, and develop into seeds following successful fertilization. The ovule content of ovaries varies among flower types= hermaphrodite flowers typically contain 3-4 viable ovules per ovary functionally female and pistillate flowers often contain 3-4 viable ovules, sometimes with slightly higher average numbers (3.5-4.2 ovules per ovary) reflecting enhanced female function investment functionally male flowers contain 1-3 ovules that may be viable or partially aborted staminate flowers contain 0-2 ovules that are typically aborted or non-viable. Ovule number per ovary shows cultivar-specific variation beyond flower type effects, with some cultivars consistently producing 4 ovules per ovary (>80% of flowers with 4 ovules), other cultivars averaging 3 ovules (mix of 2-4 ovule flowers with mode at 3), and rare cultivars or selections consistently producing only 2 ovules (1 per locule). The ovule-containing condition is essential for seeded grape production, as each viable ovule represents potential for one seed following successful fertilization, with berry seed number (typically 1-4 seeds per seeded berry) directly reflecting the number of successfully fertilized ovules in the original ovary. Environmental stresses during ovule development (BBCH 54-57) can affect ovule number and viability= heat stress >35°C during megasporogenesis can cause ovule abortion reducing viable ovule number by 30-60% water stress can reduce ovule initiation nutritional deficiencies (particularly boron) can impair ovule development and subsequent fertilization success. Ovule containing ovaries transition to seed-containing berries following successful pollination and fertilization (BBCH 61-71), with fertilized ovules developing into seeds (BBCH 71-89) while unfertilized ovules typically abort and degenerate during early fruit development (BBCH 71-75). Seedless grape cultivars (stenospermocarpic types like Thompson Seedless) produce ovule containing ovaries with 2-4 ovules that may undergo fertilization but seeds abort early during berry development, creating seedless or seed-trace berries despite initial ovule presence, while other seedless cultivars achieve parthenocarpic berry development without fertilization from ovule containing ovaries.
Ovule_Containing_Ovary emphasizes the presence of ovules as the critical functional feature enabling fertilization and seed production, and must be distinguished from ovule-lacking ovaries (rare aberrations or vestigial ovaries in staminate flowers). The ovule-containing condition is universal in functional pistils (hermaphrodite, functionally female, pistillate flower types) and represents the essential prerequisite for fruit set and seeded berry production. Sub-classification into specific ovule numbers (Two_Ovule_Ovary, Four_Ovule_Ovary) reflects the common range of ovule number variation. Understanding ovule content and distribution is essential for predicting seed number potential, for breeding seedless cultivars, and for understanding fruit set biology.
Eizellen-Fruchtknoten
ovule-bearing ovary
ovulenhaltiger Fruchtknoten
samenanlagentragender Fruchtknoten
seed-precursor ovary
Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 93-98 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 99-108.
Ovule Containing Ovary
Cabernet Sauvignon in Napa Valley shows ovule fertilization during late June Year N+1 at BBCH 69, with 2-4 ovules per flower experiencing variable fertilization success, typical outcomes producing 1.8-2.4 seeds per berry from partial ovule fertilization, failed ovules aborting within 2 weeks, successful seeds initiating development, under warm climate conditions, typical of this late-season cultivar with moderate seed number supporting berry development
Ovule fertilization represents the reproductive success phase determining seed development outcomes when fertilization attempts result in variable success across 2-4 ovules per flower, with successful fertilization producing viable seeds, failed fertilization causing ovule abortion, and partial fertilization creating variable seed numbers affecting berry development and fruit set. This variable outcome phase occurs during late flowering in late June-early July in temperate regions Year N+1, with outcomes determined by fertilization success 1-3 days post-pollination and manifesting over subsequent 1-2 weeks, corresponding to BBCH 69 fertilization completion stages. Ovule fertilization encompasses multiple outcomes including successful double fertilization in some ovules producing viable embryo and endosperm initiating seed development, failed fertilization in other ovules from pollination inadequacy, tube growth failure, or fusion problems causing ovule abortion, and partial success patterns creating 0-3 seeds per berry affecting subsequent development. Grapevine flower ovule number shows typically 2-4 ovules per bicarpellate ovary distributed in two locules, each ovule representing potential seed if successfully fertilized, and total ovule number providing multiple fertilization opportunities per flower. Fertilization attempt variability shows independent fertilization success for each ovule, 1-4 pollen tubes potentially reaching ovary region though usually only 1-2 successfully penetrating and discharging, competitive dynamics among tubes and ovules affecting which ovules fertilize, and stochastic variation creating diverse outcomes across flowers. Successful fertilization outcome shows ovule achieving double fertilization with viable zygote and endosperm formation, seed development initiating with embryo and endosperm growth, integuments developing into seed coat, and seed maturation proceeding toward viable propagule 90-120 days duration. Failed fertilization outcome shows ovule experiencing fertilization failure from various causes, ovule abortion with tissue degeneration and resource reabsorption, no seed development contribution, and potential flower abortion unless other ovules successfully fertilize providing hormonal rescue. Partial fertilization pattern shows common outcome with 1-2 ovules fertilizing while others fail within multi-ovule flower, resulting seed number typically 1-3 per berry with average 1.5-2.5 in many cultivars, and seed number variation creating berry size diversity. Fertilization success rate per ovule shows variable efficiency with 30-60% individual ovules successfully fertilizing depending on conditions, cumulative flower-level success higher since multiple ovules provide redundancy, and overall pattern creating reliable fruit set despite individual ovule variation. Pollen tube competition effects show 2-5 tubes potentially reaching ovary but limited ovule access, first-arrival advantage with earliest tubes gaining ovule access, and competitive exclusion preventing excessive fertilization per ovule. Ovule receptivity variation shows developmental asynchrony among ovules potentially affecting fertilization timing, some ovules ready earlier receiving first-arriving tubes, and temporal spread creating fertilization window across ovules. Seed number determination shows finalized 1-3 days post-pollination as fertilization outcomes establish, seed number visible 2-3 weeks later as developing seeds become distinct from aborted ovules, and seed count affecting berry size and development with more seeds generally producing larger berries through greater auxin production. Seed number effects on berry show positive correlation between seed number and final berry size, seedless or single-seed berries typically smaller than multi-seed berries, auxin production proportional to seed number driving differential growth, and seed presence stimulating cell division and expansion in surrounding pericarp. Auxin production variation shows greater total auxin from multiple seeds, hormone accumulation triggering stronger growth responses, and seed number as determinant of berry growth potential during early development. Berry size variation from seed number shows seedless berries 30-50% smaller than seeded berries in seeded cultivars, single-seed berries intermediate size, 2-3 seed berries largest, and within-cluster size heterogeneity reflecting seed number diversity. Seedless berry formation shows stenospermocarpic development in some cultivars with fertilization occurring but embryo aborting early, residual seed traces or rudiments remaining, berry development proceeding from initial hormone stimulus despite seed loss, and parthenocarpic development in rare cases with ovary growth without fertilization. Ovule abortion timing shows failed ovules degenerating within 1-2 weeks post-fertilization, tissue browning and collapse distinguishing from viable developing seeds, and resource reabsorption recycling nutrients from failed reproductive attempts. Flower-level fertilization outcomes show complete failure with all ovules unfertilized causing flower abortion, partial success with 1-3 ovules fertilized supporting berry development, and complete success with most ovules fertilized producing maximum seed number. Environmental effects on ovule fertilization include flowering weather affecting pollination success and fertilization efficiency across ovules, temperature during fertilization affecting fusion success and early seed development, boron nutrition affecting pollen tube growth and fertilization completion, and overall conditions determining fertilization success distribution. Cultivar variation in ovule fertilization shows genetic differences in fertilization efficiency, seed number characteristics with some cultivars averaging 1-2 seeds and others 2-3 seeds per berry, and seedless cultivars showing consistent embryo abortion patterns. The process shows three distinguishable outcome categories including successful ovule fertilization producing viable seeds, failed ovule fertilization causing abortion, and partial ovule fertilization creating variable seed numbers. Duration represents fertilization attempts and outcomes over several days with seed number determination within week post-pollination. Regional timing shows ovule fertilization outcomes during late June-early July across all climates 1-3 days following pollination attempts. Viticultural significance includes determining seed number per berry affecting berry size and development, establishing fertilization success rate affecting fruit set, demonstrating importance of multiple ovules providing fertilization redundancy, and creating within-cluster berry size variation affecting harvest uniformity. Research applications include analyzing factors affecting ovule-specific fertilization success, evaluating seed number determinants and effects on berry characteristics, quantifying fertilization efficiency and redundancy benefits, and assessing ovule fertilization patterns as fruit set and quality determinants.
Ovule_Fertilization represents variable reproductive outcome phase within Fruit_Set determining seed number. Subdivided into Successful_Ovule_Fertilization producing seeds, Failed_Ovule_Fertilization causing abortion, and Partial_Ovule_Fertilization creating variable seed numbers. Occurs 1-3 days post-pollination with outcomes manifesting over 1-2 weeks. Typically 2-4 ovules per flower with 30-60% individual success rate. Seed number 1-3 per berry affects size. Multiple ovules provide redundancy supporting fruit set. Determines berry developmental potential.
Befruchtungsergebnisse
Samenanlage-Befruchtung
Samenanlage-Schicksalsbestimmung
Samenbildung
fertilization outcomes
ovule fate determination
ovule reproductive success
seed formation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 408-412 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 310-313 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Ovule Fertilization
Merlot in Bordeaux shows ovule penetration during mid-late June Year N+1 at BBCH 65-69, with pollen tubes discharging sperm cells into embryo sac 26-40 hours post-pollination, double fertilization producing diploid zygotes and triploid endosperm nuclei, 1-3 successful fertilizations per flower creating 1-3 seeds per berry, rapid ovary expansion to 12-18 mm within 2 weeks, under oceanic conditions of 22-27°C temperature, typical of this mid-season cultivar achieving 54-64% fertilization success and fruit set
Ovule penetration represents the final fertilization phase following synergid entry when pollen tube discharges two sperm cells into embryo sac enabling double fertilization with sperm-egg fusion forming diploid zygote and sperm-central cell fusion forming triploid endosperm nucleus, establishing successful reproduction and seed development. This critical discharge and fusion phase occurs during fertilization in mid-late June in temperate regions Year N+1, typically proceeding within hours following synergid penetration as fertilization culminates, corresponding to BBCH 65-69 fertilization completion stages. Ovule penetration process encompasses tube discharge mechanism releasing sperm cells into embryo sac cytoplasm, sperm cell migration to fusion targets with one approaching egg cell and other approaching central cell, and gamete fusion events creating zygote and endosperm nucleus completing sexual reproduction. Tube discharge preparation shows apex positioned in penetrated synergid cell, wall weakening at tip through localized degradation, cytoplasmic pressure building creating discharge force, and sperm cells positioned near apex ready for release. Discharge mechanism involves tube apex rupture releasing contents into embryo sac, explosive or gradual discharge depending on turgor and wall properties, sperm cells expelling into synergid and embryo sac cytoplasm, and vegetative nucleus and cytoplasm also discharging though reproductive role completed. Sperm cell release shows two genetically identical haploid cells entering embryo sac, cells non-motile requiring passive transport or short-distance migration to fusion targets, and cells maintaining integrity and viability during discharge and approach phases. Sperm cell fate determination shows one sperm cell designated for egg fusion creating embryo, other sperm cell targeting central cell creating endosperm, and differentiation possibly occurring during discharge or immediately after through unknown mechanisms. Egg cell approach by first sperm shows migration or drift through embryo sac cytoplasm toward egg positioned at micropylar pole adjacent to penetrated synergid, short distance typically <100 micrometers enabling rapid approach within minutes to hours, and arrival at egg cell surface preparing for fusion. Central cell approach by second sperm shows migration toward large central cell occupying majority of embryo sac volume, central cell nucleus or polar nuclei providing fusion target, and sperm arrival at central cell preparing for fusion. Sperm-egg fusion creating zygote involves sperm cell contacting egg cell surface, membrane fusion integrating sperm into egg cytoplasm, nuclear fusion combining haploid sperm nucleus with haploid egg nucleus, formation of diploid 2n zygote nucleus containing genetic contributions from both parents, and egg activation transitioning from developmental arrest to active division initiating embryo development. Sperm-central cell fusion forming endosperm shows second sperm fusing with central cell, nuclear fusion combining haploid sperm nucleus with diploid central cell nucleus formed from polar nuclei fusion or fusing directly with separate polar nuclei, formation of triploid 3n endosperm nucleus with 2 per 1 maternal per paternal genomic ratio, and endosperm development initiating providing nutritive tissue supporting embryo growth. Double fertilization synchronization shows two fusion events typically occurring within hours of tube discharge, coordinated timing ensuring both embryo and endosperm development initiate together, and synchronization critical for normal seed development with embryo requiring endosperm nutritive support. Zygote development initiation shows first cell division typically within 24-48 hours of fertilization, progressive cell divisions establishing embryo proper and suspensor, and embryo growth consuming endosperm nutritive reserves during seed maturation. Endosperm development shows free nuclear division phase with rapid mitosis producing multinucleate coenocyte, cellularization phase forming cellular endosperm tissue, and subsequent embryo consumption reducing endosperm to thin layer in mature seed. Fertilization completion timing shows discharge and fusion typically occurring 24-48 hours after initial pollination representing culmination of reproductive sequence, successful fertilization confirmed through continued ovule and ovary development, and berry growth initiation providing visible evidence of reproductive success. Ovary response to fertilization shows rapid expansion from 4-8 mm pre-fertilization to 10-18 mm within 1-2 weeks, cell division intensity maximizing during early post-fertilization period, auxin production from developing seeds preventing flower abscission and driving berry growth, and resource allocation shifting dramatically to developing fruits consuming 40-60% photosynthate. Seed development following fertilization shows integuments developing into seed coat, endosperm and embryo developing within seed creating viable propagule, and seed maturation coordinating with berry ripening over subsequent months. Failed fertilization outcomes include unfertilized flowers lacking seed development stimulus, ovary failing to expand and develop, flower abortion and abscission occurring within days to weeks, and coulure poor fruit set from fertilization failure affecting yield. Environmental effects on fusion show temperature sensitivity with optimal 22-28°C supporting efficient fusion and development initiation, cool conditions potentially impairing fusion or subsequent development, excessive heat above 33°C causing reproductive dysfunction, and critical temperature dependence during fertilization window. Multiple tube considerations show typically 1-2 tubes successfully penetrating and discharging per flower though 2-4 ovules present, first-fertilized ovules excluding subsequent tubes through unknown mechanisms, and successful fertilization number typically 1-3 ovules creating 1-3 seeds per berry. Polyspermy prevention shows single tube reception per ovule preventing multiple sperm delivery, mechanisms possibly including rapid ovule signaling changes after fertilization, physical barriers preventing additional tube access, or synergid reception limitation, and protection ensuring normal diploid embryo and triploid endosperm formation. The process shows two distinguishable aspects progressing from egg cell approach positioning sperm for fusion, through pollen tube discharge releasing sperm and enabling fusion events. Duration typically spans minutes to hours from discharge to completed fusion representing culminating fertilization events. Cultivar variation in ovule penetration may include discharge efficiency, fusion success rates, and developmental coordination affecting fertilization outcomes. Regional timing shows ovule penetration occurring during mid-late June in warm climates, late June in temperate regions, and late June-early July in cool areas, completing 24-48 hours after pollination. Viticultural significance includes representing culminating reproductive events determining seed and berry formation, requiring successful completion of entire pollination-germination-growth-penetration sequence, demonstrating critical environmental sensitivity during fusion period, establishing fertilization success as direct determinant of fruit set and yield, and marking definitive transition from reproductive to developmental phase. Research applications include analyzing discharge mechanisms and sperm delivery, investigating double fertilization coordination and timing, evaluating environmental and genetic factors affecting fusion success, quantifying fertilization success rates and seed number outcomes, assessing fertilization effects on berry development, and developing strategies improving fruit set through enhanced fertilization efficiency.
Ovule_Penetration is second major Fertilization phase representing culminating reproductive events. Subdivided into Egg_Cell_Approach positioning sperm and Pollen_Tube_Discharge releasing sperm for fusion. Duration minutes to hours from discharge to fusion completion. Involves double fertilization creating 2n zygote and 3n endosperm. Determines seed number 1-3 per berry. Initiates berry development through auxin production. Success rate 40-70% determining fruit set. Completes entire flowering reproductive sequence. Marks transition to berry development phase.
Samenanlage-Penetration
Spermazell-Entladung
Zygoten- und Endospermbildung
sperm cell discharge
zygote and endosperm formation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 392-400 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 298-305 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Ovule Penetration
Pinot Noir own-rooted trunk in Chilean vineyard at 12 years post-planting shows 55-75 mm diameter with continuous vascular tissue from root crown through trunk without graft union, 12 uniform annual growth rings averaging 2.8-4.2 mm width, bark characteristics consistent with Pinot Noir genotype throughout trunk length, and carbohydrate reserves 70-95 g per vine supporting consistent annual productivity
An own-rooted trunk is a scion trunk of Vitis vinifera that develops directly from the original planted cutting or seedling without grafting onto a rootstock, representing an ungrafted vine where the above-ground cultivar variety and below-ground root system are genetically identical and structurally continuous. Own-rooted trunks develop from rooted cuttings (dominant commercial propagation method) or seedlings (breeding programs, wild populations) through= initial root formation from cutting base or radicle emergence from seed, simultaneous shoot development from cutting buds or seedling apical meristem, progressive trunk thickening through secondary growth, and establishment of continuous vascular connection from root tips through trunk to canopy without graft union interruption. Own-rooted trunk anatomy shows continuous tissue organization= xylem vessels and phloem sieve tubes maintaining uninterrupted connection from root xylem through trunk to shoot xylem, uniform cambial activity along entire trunk length (contrasting with grafted vines showing potential graft union compatibility issues), consistent wood anatomy throughout trunk reflecting single genotype characteristics, absence of graft union swelling or anatomical discontinuity, and bark characteristics uniform with cultivar genetics. Own-rooted trunk characteristics include= genetic uniformity between above-ground and below-ground portions (both scion cultivar genotype), absence of rootstock influence on trunk vigor or growth patterns (trunk development reflects only scion genetics and environmental factors), potential susceptibility to soil-borne pests and diseases (particularly phylloxera in infested regions, nematodes in affected soils), typical trunk diameter development 15-40 mm at year 3, 40-80 mm at year 10, 80-150 mm at year 25 for moderate vigor cultivars, and potentially greater longevity in non-phylloxera regions (some own-rooted vines exceeding 100-150 years). Own-rooted trunk advantages include= elimination of graft union as potential weakness or incompatibility point, simplified propagation without grafting labor and expertise, potential cost savings in nursery production, suitability for phylloxera-free regions or resistant cultivars, and preservation of terroir expression without rootstock modification of scion behavior. Own-rooted trunk disadvantages include= vulnerability to phylloxera in infested soils (Vitis vinifera roots highly susceptible), limited adaptation to problematic soil conditions (inability to utilize rootstock tolerance to lime-induced chlorosis, salt, drought, or other stresses), lack of rootstock vigor control (cannot use dwarfing or invigorating rootstocks to modify vine size), and potential incompatibility with certain soil types or climates. Own-rooted trunk distribution globally includes= dominant in phylloxera-free regions (Chile, some Australian regions, parts of Argentina), permitted in Europe for phylloxera-resistant cultivars or quarantined areas, utilized in breeding programs for variety development, and historical own-rooted vineyards predating phylloxera introduction (pre-1860s European vineyards, some surviving ancient vines).
Own_Rooted_Trunk emphasizes ungrafted continuous structure and is distinguished from Scion_Trunk (encompassing both grafted and ungrafted) and grafted scion trunk (above graft union) by genetic and structural continuity from roots through trunk. Critical for understanding phylloxera susceptibility and viticultural geography, rootstock-free terroir expression, propagation methods and costs, and historical vineyard architecture in pre-phylloxera or phylloxera-free regions.
Eigenwurzeliger Stamm
Self-rooted trunk
Ungepfropfter Rebstamm
Ungrafted trunk
Wurzelechter Stamm
Own Rooted Trunk
Oxidation aroma assessment in returned consumer complaint Sauvignon Blanc identifies bruised apple (6.5 on 10-point fault scale) with underlying nutty-sherry notes (4.2), analytical confirmation showing acetaldehyde at 145 mg/L and sotolon at 12 μg/L indicating significant oxidative deterioration, with investigation revealing storage at fluctuating temperatures (15-32°C) in partially consumed bottle over 3 weeks that allowed oxidation through compromised closure, demonstrating how post-purchase storage abuse rather than production fault caused the oxidation and illustrating the importance of consumer education alongside quality control.
Oxidation aroma quality is an off-aroma quality that represents olfactory perception of compounds formed through excessive oxygen exposure, characterizing the range of oxidative deterioration from subtle nuttiness through pronounced bruised apple, sherry-like, and dried fruit notes that indicate premature aging or improper storage when appearing in wines not intended for oxidative style. Within the BFO framework, this quality specifically addresses the oxidative fault category arising from uncontrolled oxygen exposure, distinguishing oxidation-derived off-aromas from other fault types while recognizing the intentional use of controlled oxidation in certain wine styles (Sherry, Vin Jaune, Tawny Port) where oxidative character is a defining positive attribute rather than a fault. The ontological significance lies in oxidation representing both the most common path of wine deterioration and a deliberate winemaking technique, where the boundary between fault and style depends entirely on winemaker intention and consumer expectation, creating the ontological paradox that identical character constitutes serious defect in one context and premium quality marker in another. The compositional basis for oxidation aroma includes acetaldehyde (providing bruised apple, sherry character, threshold approximately 100 mg/L in table wine context), sotolon (creating curry, maple syrup, dried fig notes, threshold 5-8 μg/L), methional (contributing cooked potato, oxidized apple), and various Strecker aldehydes and oxidation products of varietal compounds that transform fresh fruit toward stewed, dried, and nutty character. The oxidation pathways in wine involve both enzymatic oxidation (polyphenol oxidase activity on phenolic substrates, primarily pre-fermentation) and chemical oxidation (non-enzymatic reactions involving oxygen, iron, copper, and phenolic compounds occurring throughout wine life), with the rate of oxidative deterioration depending on oxygen exposure, temperature, SO₂ protection, and phenolic composition serving as oxidation buffers. The prevention and irreversibility of oxidation distinguishes this fault category, as oxidative damage cannot be reversed through treatment (unlike some reductive faults), making prevention through oxygen exclusion, temperature control, and adequate SO₂ maintenance the only effective approach. Understanding oxidation aroma quality enables fault identification from oxidative character, storage damage assessment based on oxidation indicators, and appropriate quality evaluation distinguishing intentional oxidative style from premature deterioration.
Oxidation_Aroma_Quality addresses the oxygen-derived fault category, which uniquely demonstrates style-dependency where identical character constitutes premium quality (Sherry, Vin Jaune) or serious defect (most table wines). This quality is irreversible once established, making prevention critical. The class supports fault identification while acknowledging the legitimate oxidative winemaking tradition that values controlled oxidation as positive attribute.
Firne
Maderisierung
Oxidationsaromen
Sherryton
maderization
oxidative character
premature aging
sherried notes
Waterhouse AL, Laurie VF (2006) Oxidation of wine phenolics: a critical evaluation and hypotheses. American Journal of Enology and Viticulture 57(3): 306-313 Escudero A et al. (2002) Characterization of the aroma of a wine from Maccabeo. Key role played by compounds with low odor activity values. Journal of Agricultural and Food Chemistry 50(11): 4048-4054
Oxidation Aroma Quality
P50 value comparison among grapevine cultivars reveals genetic variation in hydraulic drought tolerance, with Grenache showing P50 of -2.4 MPa (relatively resistant, consistent with origin in dry Mediterranean conditions), Pinot Noir at -1.6 MPa (moderate vulnerability, appropriate for cooler, wetter climate of origin), and Sultana at -1.2 MPa (relatively vulnerable, consistent with irrigated table grape use), demonstrating how P50 variation reflects adaptation to different water environments and informs cultivar selection for sites with different drought exposure.
P50 value quality is a vulnerability to cavitation quality that represents the xylem water potential at which 50% of maximum hydraulic conductivity is lost through embolism, characterizing the central parameter of the vulnerability curve that indicates the threshold water potential for substantial hydraulic dysfunction and serves as the standard metric for comparing drought tolerance across species, organs, and genotypes. Within the BFO framework, this quality specifically addresses the midpoint of the vulnerability function, distinguishing P50 from other vulnerability parameters while recognizing that P50 provides the most widely used single value for characterizing xylem drought tolerance that enables comparison across the diversity of plant hydraulic strategies. The ontological significance lies in P50 indicating the water potential threshold for significant transport system damage, where operating water potentials approaching P50 indicate high hydraulic risk, and exceeding P50 causes loss of more than half the conductivity that supplied transpiration, typically triggering stomatal closure, growth reduction, and potential branch dieback if stress persists. The methodological basis for P50 determination involves constructing vulnerability curves through measurement of hydraulic conductivity at a series of water potentials (achieved through bench dehydration, air injection, or centrifuge spinning), fitting a function (typically sigmoidal) to the PLC versus Ψ relationship, and identifying the water potential at 50% loss. The typical P50 values for Vitis vinifera range from approximately -1.0 to -3.0 MPa depending on organ (stems typically more resistant than leaves or petioles), cultivar (genetic variation in xylem anatomy affecting vulnerability), and growth conditions (acclimation to drought potentially improving resistance), with grapevine showing moderate vulnerability compared to the global range from near 0 MPa in very vulnerable species to below -15 MPa in extremely resistant conifers. The hydraulic safety margin concept uses P50 as the reference point, with the margin calculated as the difference between minimum operating water potential and P50, indicating how close the plant operates to dysfunction threshold and how much buffer exists against severe drought. Understanding P50 value quality enables quantitative comparison of drought tolerance, assessment of hydraulic safety margins, and prediction of the water potential limits for hydraulic function that management must respect.
P50_Value_Quality addresses the 50% conductivity loss threshold that serves as the standard metric for xylem drought tolerance comparison. This quality enables quantitative assessment of hydraulic safety margins. The class supports cultivar comparison and drought tolerance evaluation through P50 characterization.
50% embolism threshold
50%-Embolieschwelle
P50-Wert
P₅₀
Vulnerabilitätsmittelpunkt
hydraulic threshold
hydraulischer Schwellenwert
vulnerability midpoint
P50 Value Quality
P88 value assessment in Cabernet Sauvignon following an extreme heat and drought event reveals the approach to catastrophic hydraulic failure, with stem P88 of -3.2 MPa providing the lethal threshold, field measurements showing stem water potential reaching -2.9 MPa during the event (85% conductivity loss, approaching but not exceeding P88), and subsequent assessment revealing significant dieback in most stressed vines but survival and recovery in moderately stressed individuals, demonstrating how proximity to P88 determines the transition between recoverable stress and permanent damage.
P88 value quality is a vulnerability to cavitation quality that represents the xylem water potential at which 88% of maximum hydraulic conductivity is lost through embolism, characterizing the threshold for near-complete hydraulic failure that indicates the lethal or functionally lethal water potential for the organ or plant, beyond which recovery is unlikely without substantial dieback or death. Within the BFO framework, this quality specifically addresses the catastrophic failure threshold of the vulnerability function, distinguishing P88 from the moderate dysfunction indicated by P50 while recognizing that P88 represents the practical limit for hydraulic survival that must not be exceeded if irreversible damage is to be avoided. The ontological significance lies in P88 indicating the point of no return for hydraulic function, where loss of 88% of conductivity leaves so little functional transport capacity that water supply cannot meet even minimal demand, creating irreversible stress that typically causes organ death, branch dieback, or whole-plant mortality depending on the extent and duration of exceeding this threshold. The ecological and physiological rationale for the 88% threshold derives from the observation that this level of conductivity loss typically correlates with tissue death and failure to recover, making P88 a more biologically meaningful survival limit than P50 which indicates significant but potentially recoverable dysfunction. The relationship between P50 and P88 depends on the shape of the vulnerability curve, with steep curves having P88 relatively close to P50 (rapid transition from moderate to severe dysfunction) while shallow curves have more gradual conductivity decline between these thresholds, affecting the management implications for drought stress approaching P50. The safety margin to P88 represents the buffer between operating conditions and catastrophic failure, with typical minimum operating water potentials in grapevine remaining well above P88 under most managed conditions, though severe drought events can approach or exceed this threshold with potentially fatal consequences. Understanding P88 value quality enables assessment of the ultimate hydraulic survival limit, prediction of lethal stress thresholds, and evaluation of the risk of catastrophic hydraulic failure under extreme drought scenarios.
P88_Value_Quality addresses the 88% conductivity loss threshold indicating near-complete hydraulic failure and survival limit. This quality represents the point of no return beyond which recovery is unlikely. The class supports mortality prediction and catastrophic failure risk assessment through P88 characterization.
88% embolism threshold
88%-Embolieschwelle
P88-Wert
P₈₈
hydraulic failure point
hydraulischer Versagenspunkt
letale Kavitationsschwelle
lethal cavitation threshold
Urli M et al. (2013) Xylem embolism threshold for catastrophic hydraulic failure in angiosperm trees. Tree Physiology 33(7): 672-683 Charrier G et al. (2018) Drought will not leave your glass empty: low risk of hydraulic failure revealed by long-term drought observations in world's top wine regions. Science Advances 4(1): eaao6969
P88 Value Quality
The pago name Dominio de Valdepusa designates the 54-hectare estate of Carlos Falcó (Marqués de Griñón) near Toledo in Castilla-La Mancha, granted Vino de Pago status in 2003 as the first estate in Spain to receive this designation — the DO de Pago Dominio de Valdepusa functions as a self-contained appellation with its own production specification governing authorized varieties (Cabernet Sauvignon, Syrah, Petit Verdot, Graciano), maximum yields (4,500 kg/ha), winemaking standards, and quality control procedures, with the pago name on the label communicating single-estate origin at a level of geographic specificity that no other Spanish appellation category provides.
A vineyard designation within the Spanish wine classification system that identifies a wine as originating from a named Vino de Pago — a single estate or vineyard site of exceptional and recognized quality that has been granted its own individual Denominación de Origen (DO de Pago) status under Spanish wine law, representing the highest and most site-specific tier of the Spanish appellation hierarchy with each pago constituting an independent appellation unto itself.
Pago_Name documents Spain's unique single-estate appellation concept, introduced in the 2003 Wine Law (Ley 24/2003) as the apex of the Spanish quality hierarchy. The Vino de Pago category is exceptional in international wine law because it grants individual estates the status of independent appellations — each pago has its own registered Denominación de Origen, its own production specification, and (in most cases) its own regulatory oversight distinct from the regional DO or DOCa within which it is geographically located: this is fundamentally different from the French or German models where individual vineyards are classified within a broader appellation framework rather than constituting independent appellations. Spain currently recognizes approximately 20 Vinos de Pago, predominantly in Castilla-La Mancha, Navarra, and the Valencian Community, with each pago representing a specific estate whose exceptional quality has been demonstrated over a minimum qualification period. The pago concept has been debated within the Spanish wine industry — supporters view it as properly recognizing outstanding individual terroirs, while critics argue it fragments the appellation system and undermines regional collective identity.
Pago-Bezeichnung
Spanish single-estate DO
Vino de Pago designation
Vino-de-Pago-Name
pago estate name
spanische Einzellagen-DO
Spanish Wine Law (Ley 24/2003 de la Viña y del Vino) Art. 24 Ministerio de Agricultura, Pesca y Alimentación (2023) Vinos de Pago registrados EU Regulation 1308/2013 Art. 93
Pago Name
Cabernet Sauvignon hermaphrodite stigmas at BBCH 61-65 exhibit papillate organization with 152-188 papillae distributed across stigma surface (0.44-0.51 mm diameter, giving 165-210 papillae per mm²), individual papillae measuring 35-55 μm length and 18-26 μm diameter consisting of 3-5 cells in columnar arrangement, with papillae serving as exudate secretion sites during receptive period and as pollen capture structures achieving efficient self-pollination with 50-58% fruit set. Gewürztraminer functionally female stigmas show densely papillate organization with 195-245 papillae on larger stigmas (0.52-0.62 mm diameter, giving 220-280 papillae per mm²), individual papillae 45-65 μm length representing enhanced surface elaboration correlated with extended receptivity and cross-pollination dependency, with the dense papillae population providing maximum pollen capture efficiency for wind-borne or insect-borne cross-pollen from hermaphrodite sources. Scanning electron microscopy of Chardonnay receptive stigmas reveals papillae ultrastructure= multicellular columns with outer cell walls showing characteristic cuticular ridges, exudate secretion occurring through cuticle creating coating over papillae surfaces, pollen grains adhering to multiple papillae via exudate bridges, and pollen tubes emerging from grains and growing along papillae surfaces toward style entry points visible as darker regions between papillae bases. Pinot Noir stigma development time-course shows papillae initiation at BBCH 54-55 as epidermal cell elongations 8-12 μm long, progressive elongation and division creating multicellular structures 25-40 μm by BBCH 57-58, and final mature papillae 38-58 μm at anthesis (BBCH 61), with papillae number per stigma remaining constant from BBCH 56 onward (142-182 papillae) while individual papillae undergo elaboration, demonstrating that papillae pattern is determined early in development with subsequent papillae growth rather than additional papillae initiation.
A papillate stigma is a stigma bearing multicellular papillae (specialized epidermal projections) on its surface, representing the universal stigma organization in Vitis where papillae serve critical functions in pollen capture, adhesion, hydration, and germination support, with papillae number, size, and distribution varying among flower types and correlating with overall stigma size and functionality. Papillate stigmas are characterized by= (1) stigmatic surface covered with papillae distributed across the receptive area, with papillae typically numbering 40-250 per stigma depending on stigma diameter and flower type (2) individual papillae consisting of 2-6 cells arranged in files (columns), measuring 20-80 μm in length and 12-35 μm in diameter, with cell walls showing characteristic structure and cuticle organization (3) papillae providing increased surface area for pollen grain contact, creating three-dimensional receptive surface rather than flat epidermis (4) papillae cells serving as sites of exudate secretion in receptive stigmas, with secretory apparatus (endoplasmic reticulum, Golgi bodies, secretory vesicles) concentrated in papillae cells (5) papillae distribution often showing bilateral organization reflecting the bicarpellate pistil origin, with enhanced papillae density along two opposite margins in some stigmas creating subtle two-lobed appearance. Papillate stigmas develop through epidermal cell specialization during stigma development (BBCH 54-58), with selected epidermal cells elongating and undergoing additional divisions to create the multicellular papillae, while intervening epidermal cells remain relatively flat creating the papillate versus inter-papillae mosaic surface architecture. Papillae number and dimensions show strong correlation with stigma size= large stigmas in functionally female/pistillate flowers (0.45-0.65 mm diameter) bear 150-250 papillae medium stigmas in hermaphrodite flowers (0.35-0.55 mm diameter) bear 120-200 papillae small stigmas in functionally male flowers (0.2-0.4 mm diameter) bear 60-120 papillae vestigial stigmas in staminate flowers (0.1-0.25 mm diameter) bear <60 papillae. The functional significance of papillae relates to multiple aspects of pollination= increased surface area (papillate stigma surface area typically 150-250% of equivalent flat surface) increases probability of pollen grain contact and capture three-dimensional structure creates physical barriers helping retain captured pollen grains papillae surfaces provide multiple attachment points for pollen grains via exudate adhesion papillae cells are the primary sites of exudate secretion creating localized high-concentration exudate zones around individual pollen grains papillae structure guides pollen tube emergence and directs tubes toward transmitting tract entry points. Papillae density (number per unit stigma area) varies among flower types= functionally female/pistillate stigmas show densely papillate organization with 180-320 papillae per mm² stigma surface hermaphrodite stigmas show moderate papillae density of 140-240 papillae per mm² functionally male stigmas show sparsely papillate organization with 80-160 papillae per mm², with density variation correlating with functional differences in pollen capture and germination support efficiency.
Papillate_Stigma represents the universal stigma surface organization in Vitis and distinguishes grapevine stigmas from smooth (non-papillate) stigmas in some other plant families. Papillae are essential functional structures providing increased surface area, pollen capture capability, and exudate secretion sites. Sub-classification into Densely_Papillate and Sparsely_Papillate reflects variation in papillae density among flower types, with dense papillae correlating with enhanced receptivity and pollen capture in functionally female/pistillate types while sparse papillae correlate with reduced functionality in functionally male types. Understanding papillate stigma structure is essential for interpreting stigma function, for assessing stigma development, and for comparative studies of pollination mechanisms.
Papillen-Narbe
papilla-bearing stigma
papillentragendes Stigma
papillöse Narbe
textured stigma surface
Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 95-98 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 191-195.
Papillate Stigma
PAR data for an open-canopy VSP-trained Sauvignon Blanc vineyard in Marlborough showed exterior PPFD of 1800-2000 µmol/m²/s at solar noon, with fruit zone PAR transmission of 25-35% following standard leaf removal, delivering approximately 500-700 µmol/m²/s to cluster surfaces. Pinot Noir in Burgundy under traditional low-wire training receives daily light integrals of 35-45 mol/m²/d at the canopy exterior during July, with dense canopy interiors reducing fruit zone DLI to 3-8 mol/m²/d without leaf pulling intervention. PAR sensor arrays deployed in Napa Valley Cabernet Sauvignon blocks documented that eastern fruit zone exposure receives 60% of daily PAR in morning hours before 12:00, promoting phenolic synthesis at lower berry temperatures than equivalent afternoon exposure.
PAR data is a solar radiation data entity that records measurements of photosynthetically active radiation — the spectral band of electromagnetic energy between 400 and 700 nanometers wavelength — received at or within a grapevine canopy, expressed in micromoles of photons per square meter per second (µmol/m²/s) as instantaneous photosynthetic photon flux density (PPFD), or in moles per square meter per day (mol/m²/d) as daily light integral (DLI), quantifying the radiation directly available for photosynthetic carbon fixation by vine leaves.
PAR represents approximately 45-50% of total solar radiation energy but constitutes the sole photosynthetically useful waveband, making PAR data essential for understanding vine carbon balance and canopy light environment. Grapevine leaf photosynthesis saturates at approximately 600-800 µmol/m²/s PPFD (depending on cultivar and acclimation), meaning that exterior canopy leaves are frequently light-saturated while interior leaves operate well below saturation — this gradient drives the viticultural rationale for canopy management practices that optimize whole-canopy PAR interception. The PAR environment of the fruit zone directly influences berry composition through multiple pathways: flavonol synthesis is strictly light-dependent, anthocyanin accumulation is light-promoted, and methoxypyrazine degradation accelerates with increased berry PAR exposure. Modern canopy management research increasingly uses PAR mapping through ceptometer or point-quantum-sensor arrays to quantify fruit zone light environment and guide targeted leaf removal decisions.
PAR measurement
PAR-Daten
PPFD data
Photosynthesestrahlung
photosynthetic light data
photosynthetic photon flux
photosynthetically active radiation data
photosynthetisch aktive Strahlung
photosynthetisch wirksame Strahlung
Smart RE, Robinson MD (1991) Sunlight into Wine, Winetitles, pp.: 12-22 Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 47-58 Downey MO et al. (2006) Cultural practice and environmental impacts on the flavonoid composition of grapes and wine: a review of recent research. American Journal of Enology and Viticulture 57(3): 257-268
PAR Data
Paradormancy in intact grapevine shoots maintains lateral buds quiescent during the growing season, with only 5-15% of lateral buds developing into summer laterals under normal conditions in most V. vinifera cultivars. Shoot tip removal (hedging) releases paradormancy, triggering lateral bud growth within 7-14 days through reduced auxin supply and increased cytokinin signaling. Merlot shows weaker paradormancy with 30-50% of lateral buds growing spontaneously, while Cabernet Sauvignon maintains stronger paradormancy (10-20% lateral growth). Primary bud damage from spring frost releases secondary bud paradormancy, with secondary shoots emerging within 10-21 days depending on temperature and cultivar.
A dormancy disposition that is the inherited capacity of a Vitis vinifera plant to maintain lateral and secondary buds in a quiescent state through correlative inhibition (apical dominance) exerted by actively growing shoot tips and primary buds, preventing lateral growth until the inhibitory signal is removed or overcome.
Paradormancy differs mechanistically from endodormancy, being imposed by signals from other plant organs rather than internal bud factors. The auxin-cytokinin balance hypothesis explains paradormancy through basipetal auxin transport from shoot tips suppressing lateral bud cytokinin response. Paradormancy strength affects canopy management requirements, with weak paradormancy increasing lateral shoot removal labor but potentially providing recovery capacity following frost or hail damage. The trait shows cultivar-specific expression modified by vine vigor and nitrogen status.
Apikaldominanz-Disposition
Paradormanz
apical dominance disposition
correlative inhibition trait
korrelative Hemmung
lateral bud suppression capacity
Cline MG (1991) Apical dominance. The Botanical Review 57:318-358 Dry PR (2000) Canopy management for fruitfulness. Australian Journal of Grape and Wine Research 6:109-115
Paradormancy Disposition
Paradormancy assessment in intact versus decapitated Syrah shoots reveals the correlative control of lateral bud outgrowth, with intact shoots showing strong paradormancy (lateral buds remaining inhibited, 0% outgrowth during active apical growth), shoots with apical meristem removed showing paradormancy release within 10 days (85% of lateral buds initiating outgrowth as auxin source removed), and shoots with apical meristem retained but shaded showing intermediate response (35% lateral outgrowth as reduced photosynthate supply weakens apical dominance), demonstrating how paradormancy depends on signals from dominant organs that can be manipulated through management interventions.
Paradormancy quality is a phenological quality that represents the inhibition of bud growth imposed by signals from other plant organs rather than by internal bud physiology or environmental conditions, characterizing the correlative inhibition that maintains developmental coordination among plant parts by suppressing growth of subordinate buds while dominant organs (typically apical meristems) are actively growing. Within the BFO framework, this quality addresses the correlative control dimension of bud dormancy, distinguishing growth inhibition mediated by organ-to-organ signaling from the internally controlled endodormancy or environmentally imposed ecodormancy while recognizing that paradormancy operates during the growing season to coordinate vine architecture and resource allocation by preventing simultaneous growth of all potential meristems. The ontological significance lies in paradormancy enabling the hierarchical organization of grapevine growth, where apical dominance and other correlative inhibitions channel resources into primary growing points rather than dissipating them across numerous competing meristems, with this coordination essential for establishing the organized canopy architecture that supports efficient light interception and fruit development. The physiological basis for paradormancy involves hormonal signaling, particularly the polar transport of auxin from apical regions that inhibits lateral bud outgrowth, with cytokinin and strigolactone also playing roles in the complex signaling network that integrates information about plant nutritional status, light environment, and developmental stage to regulate which buds remain inhibited and which are released to grow. The release from paradormancy occurs when the inhibiting signal is removed (through apical damage, pruning, or natural senescence of the dominant organ) or when the balance of promotive and inhibitory signals shifts to favor outgrowth (through changes in light, nutrition, or developmental programming), with this release enabling the compensatory growth and branching that allows vines to recover from damage or to fill canopy space made available by thinning or environmental events. Understanding paradormancy quality enables appreciation of the correlative controls coordinating vine architecture, prediction of lateral bud response to pruning and canopy management operations, and recognition of the hormonal signaling that integrates growth across the whole plant.
Paradormancy_Quality serves as the parent class for specific correlative inhibition mechanisms (Apical Dominance, Lateral Bud Inhibition). This quality addresses organ-to-organ signaling maintaining developmental coordination during the growing season. The class supports canopy management understanding through paradormancy characterization.
Apikalkontrolle
Sommerdormanz
correlative inhibition
growth suppression
summer dormancy
Lang GA et al. (1987) Endo-, para-, and ecodormancy: physiological terminology and classification for dormancy research. HortScience 22(3): 371-377 Domagalska MA, Leyser O (2011) Signal integration in the control of shoot branching. Nature Reviews Molecular Cell Biology 12(4): 211-221
Paradormancy Quality
The parent material description for the Barolo production zone (Langhe, Piedmont, Italy) documents two contrasting geological substrates that define the appellation's two recognized terroir types — the Tortonian-age blue-grey calcareous marl (marna di Sant'Agata Fossili, 11-7 Ma) producing the heavier, more clay-rich soils of Serralunga d'Alba and Monforte d'Alba (yielding structured, tannic, long-aging Barolo), and the Helvetian-age sandier calcareous marl and sandstone (arenarie di Diano d'Alba, 16-11 Ma) producing the lighter, more calcareous soils of La Morra and Barolo commune (yielding more aromatic, earlier-maturing Barolo). This geological dichotomy — Tortonian versus Helvetian parent material — is the fundamental terroir distinction recognized by Barolo producers and documented in the recently established Menzioni Geografiche Aggiuntive (MGA) vineyard classification system, where each of the 181 named vineyard sites can be characterized by its position on one or the other geological formation.
A soil description component that records the geological substrate from which the vineyard soil has developed through weathering processes — documenting the rock type (sedimentary, igneous, or metamorphic), specific lithology (limestone, sandstone, granite, schist, basalt, marl, loess, alluvium), geological age and formation name, and the mineralogical composition that determines the soil's chemical inheritance (base cation supply, micronutrient availability, calcium carbonate content, pH buffering capacity) and physical inheritance (particle size of weathering products, fragment content, drainage characteristics) from its geological origin.
Parent_Material_Description documents the geological foundation of soil terroir — the rock or unconsolidated sediment from which the soil has developed through physical weathering (fragmentation by frost action, root growth, and thermal expansion), chemical weathering (dissolution, hydrolysis, oxidation, and carbonation of primary minerals), and biological weathering (organic acid production by roots, fungi, and bacteria). The parent material determines the soil's mineralogical inheritance — limestone parent material produces calcareous, high-pH, calcium-rich soils granite parent material produces acidic, quartz-rich, potassium-bearing soils from feldspar weathering basalt parent material produces iron-rich, dark-colored soils with high micronutrient availability and schist parent material produces thin, stony, well-drained soils with platy fragment structure: these mineralogical differences affect vine nutrition, must composition, and potentially wine sensory character, though the mechanisms linking parent material to wine character remain incompletely understood despite centuries of empirical observation that specific geological substrates produce distinctive wine styles. The viticultural significance of parent material extends beyond soil chemistry to include the physical properties of the rock fragments (gravel, stones, cobbles) that remain in the soil profile — the abundance, size, shape, and lithology of these coarse fragments affect soil drainage (creating preferential flow paths around fragments), thermal behavior (dark-colored fragments absorbing and re-radiating heat, as in the galets of Châteauneuf-du-Pape and the slate of the Mosel), and root distribution (roots exploiting weathered bedrock fractures for deep water access).
Ausgangsgestein-Dokumentation
Ausgangsmaterial-Beschreibung
bedrock description
geological substrate record
geologisches Substrat
lithological characterization
Wilson JE (1998) Terroir — The Role of Geology, Climate, and Culture in the Making of French Wines, University of California Press Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873 van Leeuwen C et al. (2004) Influence of climate, soil, and cultivar on terroir, American Journal of Enology and Viticulture 55: 207-217 White RE (2003) Soils for Fine Wines, Oxford University Press
Parent Material Description
Pinot Noir serves in a Parent_Role when crossed with disease-resistant breeding lines, contributing wine quality genetics to progeny evaluated for both resistance and quality. Vitis amurensis bears a Parent_Role in cold-hardiness breeding, contributing winter survival genes to crosses with V. vinifera varieties for continental climate viticulture. Regent (a registered PIWI variety) functions in a Parent_Role as a bridge parent, its combination of resistance and acceptable quality making it valuable for subsequent crosses seeking to improve wine character while maintaining disease resistance.
A breeding role that inheres in a grapevine genotype by virtue of its function as a genetic contributor in controlled crosses, providing gametes (egg cells or pollen) that combine at fertilization to produce hybrid offspring. Parent roles are distinguished by sex function (maternal parent providing ovules, paternal parent providing pollen), by breeding strategy function (recurrent parent in backcrossing, donor parent for specific traits), and by the specific genetic contributions targeted from each parent. Grapevine parents in breeding crosses are typically selected for complementary trait contributions—one parent may contribute wine quality characteristics while the other provides disease resistance, with the breeding objective being progeny combining favorable alleles from both parents. Parent selection is critical to breeding success the genetic composition of parents determines the trait variation available in progeny for selection, and inappropriate parent combinations may produce populations lacking commercially viable individuals. Most grapevine breeding involves crossing between distinct genotypes (wide crosses for trait introgression, narrower crosses for quality improvement), with selfing rarely used due to severe inbreeding depression in Vitis.
Parent_Role is subclassed by sex function (Maternal_Parent_Role, Paternal_Parent_Role) and by breeding strategy function (Recurrent_Parent_Role, Donor_Parent_Role), recognizing that parents may be characterized by their gametic contribution and by their role in breeding program design. The same genotype may serve different parent roles in different crosses—as maternal parent in one cross, paternal in another as donor parent for one trait, recurrent parent for another trait. Parent role assignment in grapevine breeding often reflects practical considerations (which parent sets fruit more reliably, which is available as pollen source) alongside strategic trait combination objectives. Controlled crossing in grapevines requires emasculation (removal of anthers before pollen shed) of the seed parent and deliberate pollen application, distinguishing breeding from open-pollinated seed production.
Elter
Elternpflanze
Kreuzungselter
Zuchtpartner
breeding parent
cross parent
parent
progenitor
Parent Role
The parentage analysis result for Chardonnay (Bowers et al. 1999) reports the following analytical evidence — 322 cultivars genotyped at 6 SSR loci (VVS2, VVMD5, VVMD7, VVMD27, VVMD28, VVMD32), pairwise exclusion analysis testing all 51,681 possible parent pair combinations, with only one pair (Pinot × Gouais Blanc) not excluded at any locus — combined LOD score of 18.7 for Pinot × Gouais Blanc parentage versus the next-best parent pair LOD of 3.2 (a likelihood ratio exceeding 10¹⁵ in favor of the Pinot × Gouais Blanc assignment), subsequently confirmed by additional studies using 20-50 SSR loci with zero exclusions across all analyzed markers. The same analysis simultaneously identified 15 additional cultivars (Gamay Noir, Aligoté, Auxerrois, Melon, and others) as Pinot × Gouais Blanc offspring, revealing an unexpectedly prolific parent pair whose natural hybridization in medieval Burgundian and northeastern French vineyards produced a dynasty of commercially important cultivars.
A parentage information record that documents the specific analytical output of a molecular parentage study, reporting the marker system, locus count, candidate parent panel size, statistical method (exclusion-based, likelihood-based, or Bayesian), confidence metrics (LOD scores, exclusion probabilities, posterior probabilities), and the resulting parentage assignment or non-exclusion outcome for a specific cultivar, providing the evidentiary detail underlying an inferred parentage claim.
Parentage_Analysis_Result provides the granular analytical evidence that supports or refutes specific parentage hypotheses, distinguished from Inferred_Parentage (which records the conclusion) by its focus on the methodology and statistics of the analysis itself. The documentation of analytical parameters is essential because the confidence of parentage assignment depends critically on the number and informativeness of markers used, the size and composition of the candidate parent panel, and the statistical method applied: early parentage studies using 6-9 SSR loci could identify parent pairs with high confidence through exclusion but had limited power to detect parentage when one parent was missing from the panel, while modern studies using 20-50 SSR loci or thousands of SNPs provide sufficient statistical power for both complete parentage (both parents identified) and half-parentage (one parent identified with the other absent from the panel) analysis. The statistical frameworks range from simple exclusion counting (the number of loci at which a candidate parent is compatible) through likelihood ratio methods (Marshall et al. 1998, CERVUS software) to Bayesian approaches (COLONY software) that simultaneously estimate parentage, sibship, and population structure — each providing different confidence metrics (exclusion probability, LOD score, posterior probability) that must be documented alongside the parentage assignment to enable evaluation of the claim's reliability.
Abstammungsanalyse-Ergebnis
Abstammungszuordnungs-Nachweis
molecular parentage analysis output
molekulare Elternschaftstest-Ergebnis
parentage assignment evidence
parentage test result
Bowers JE et al. (1999) Historical genetics: the parentage of Chardonnay, Gamay, and other wine grapes of northeastern France, Science 285: 1562-1565 Marshall TC et al. (1998) Statistical confidence for likelihood-based paternity inference in natural populations, Molecular Ecology 7: 639-655 Lacombe T et al. (2013) Large-scale parentage analysis in an extended set of grapevine cultivars, Theoretical and Applied Genetics 126: 401-414
Parentage Analysis Result
SO4 (Selection Oppenheim 4) bears a Vitis_Berlandieri_Derived_Rootstock_Role and a Vitis_Riparia_Derived_Rootstock_Role, combining lime tolerance from V. berlandieri with phylloxera resistance and moderate vigor from V. riparia. 110 Richter bears Vitis_Berlandieri_Derived and Vitis_Rupestris_Derived roles, combining lime and drought tolerance from both parental species. Ramsey (Salt Creek) bears a Vitis_Champinii_Derived_Rootstock_Role, inheriting nematode resistance, salt tolerance, and heat adaptation from this Texas species.
A rootstock role that inheres in a grafted rootstock by virtue of its genetic derivation from one or more wild Vitis species whose characteristic traits determine the rootstock's functional properties. Parentage-based rootstock roles classify rootstocks according to the North American Vitis species contributing to their genome, recognizing that the adaptive traits of these wild species—evolved over millions of years in response to specific ecological pressures—fundamentally determine rootstock performance characteristics including phylloxera resistance, nematode resistance, drought tolerance, lime tolerance, cold hardiness, vigor effects, and nutrient acquisition patterns. The major parental species utilized in rootstock breeding are V. riparia (cold hardiness, low vigor, phylloxera resistance), V. rupestris (phylloxera resistance, drought tolerance, deep rooting), V. berlandieri (lime tolerance, drought tolerance, moderate vigor), V. champinii (nematode resistance, salt tolerance, heat tolerance), V. cinerea (phylloxera resistance, vigor), and less commonly V. longii and V. solonis. Most commercial rootstocks are interspecific hybrids combining traits from two or three parental species.
Parentage_Based_Rootstock_Role organizes rootstocks by genetic origin, complementing the functional classification (vigor, stress resistance, nutrient acquisition) with an evolutionary and breeding perspective. A single rootstock typically bears multiple parentage-based roles reflecting its hybrid origin. The subclasses are not mutually exclusive most commercial rootstocks are F1 hybrids or more complex combinations bearing two or more parentage roles. This classification aids rootstock selection by linking observable traits to their genetic sources and predicting trait combinations in breeding programs. The seven subclasses represent the major Vitis species utilized in rootstock development additional minor contributors (V. aestivalis, V. labrusca, V. cordifolia) are not represented as separate subclasses due to limited commercial relevance.
Elternschafts-basierte Unterlage
abstammungsbasierte Unterlage
artbasierte Unterlage
genetic background rootstock
parentage-based rootstock
pedigree-based rootstock
species-derived rootstock
Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22 Heinitz CC et al. (2019) Crop wild relatives of grape (Vitis vinifera L.), in North American Crop Wild Relatives Vol 2, Springer pp.: 329-351 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia
Parentage Based Rootstock Role
The parentage information for Müller-Thurgau documents a complex history of correction through molecular analysis — the cultivar was long attributed to a deliberate cross of Riesling × Silvaner performed by Hermann Müller from the Swiss canton of Thurgau in 1882 at the Geisenheim research station, but SSR parentage analysis (Dettweiler et al. 2000, Sefc et al. 1997) excluded Silvaner as a parent and identified the true parentage as Riesling × Madeleine Royale, with the Riesling maternal contribution confirmed by chloroplast haplotype analysis (haplotype C, shared with Riesling but not Madeleine Royale) — this correction of a century-old misattribution exemplifies the transformative impact of molecular parentage analysis on grapevine genealogical knowledge and necessitated the retention of both the documented (historical) parentage and the inferred (molecular) parentage in cultivar databases.
A genetic information entity that documents the biological parent-offspring relationships of a Vitis vinifera cultivar, recording the identity of maternal and paternal parent cultivars (where known), the evidence supporting parentage assignments (molecular marker analysis, historical records, or breeding documentation), and the degree of certainty associated with each parentage claim, providing the foundational genealogical data from which pedigree networks and cultivar family relationships are reconstructed.
Parentage_Information is the umbrella class for all data relating to the biological origins of grapevine cultivars, a domain that has been revolutionized by molecular marker analysis since the mid-1990s. Prior to SSR-based parentage testing, cultivar origins were known only for deliberately bred varieties where crossing records existed (a minority of cultivars, primarily 19th-20th century breeding products) or for cultivars with documented historical accounts of seedling selection: the vast majority of traditional wine grape cultivars — including virtually all major European varieties selected over centuries or millennia from spontaneous seedlings — had unknown or speculative parentage based on morphological similarity, geographic co-occurrence, or folk tradition. The application of SSR exclusion analysis and likelihood-based parentage assignment (beginning with Bowers and Meredith's 1997 identification of Cabernet Sauvignon's parents) has since revealed the parentage of hundreds of cultivars, uncovering unexpected relationships (Chardonnay as Pinot × Gouais Blanc), correcting long-standing misattributions (Müller-Thurgau, Pinotage), and identifying prolific parent cultivars (Pinot and Gouais Blanc alone produced at least 75 documented offspring). Subclasses distinguish the evidence basis and certainty level of parentage claims.
Abstammungsinformation
Elternschafts-Dokumentation
cultivar parentage record
genealogical data
genealogische Daten
parent-offspring documentation
Bowers JE and Meredith CP (1997) The parentage of a classic wine grape, Cabernet Sauvignon, Nature Genetics 16: 84-87 Bowers JE et al. (1999) Historical genetics: the parentage of Chardonnay, Gamay, and other wine grapes of northeastern France, Science 285: 1562-1565 Vouillamoz JF and Grando MS (2006) Genealogy of wine grape cultivars: Pinot is related to Syrah, Heredity 97: 102-110 Lacombe T et al. (2013) Large-scale parentage analysis in an extended set of grapevine cultivars, Theoretical and Applied Genetics 126: 401-414
Parentage Information
A Criolla Grande vine in Mendoza's traditional flood-irrigated vineyards bears a Parral_Role, the overhead structure supporting the high yields of this workhorse variety for domestic Argentine consumption—a remnant of the production-oriented viticulture that dominated before quality revolution. A Malbec vine in historic parral training in San Juan Province serves in a Parral_Role representing traditional regional practice, though most premium Malbec production has converted to modern VSP systems better suited to quality-focused viticulture. A Torontel (Torrontés) vine in Chile's hot Norte Chico region bears a Parral_Role, the overhead shading protecting aromatic Muscat-type grapes from sun damage while maintaining yields for Pisco production.
A pergola-trained vine role that inheres in a grapevine trained to the South American parral system, an overhead trellis structure historically dominant in Argentine and Chilean viticulture, with high cordons (180-220 cm) supporting horizontal canopy extension that creates shade beneath the vine. Parral (Spanish: derived from "parra," grapevine or vine arbor) represents adaptation of Mediterranean pergola traditions to New World conditions, providing fruit protection from intense high-altitude UV radiation (critical in Mendoza at 800-1,500 m elevation) and enabling the high yields (20-40 tonnes/hectare) that characterized South American production before quality-focused restructuring. Traditional parral vineyards use substantial wooden post infrastructure with overhead wire grids, requiring major establishment investment but enabling furrow irrigation and agricultural use of shaded inter-row space. Quality-focused Argentine and Chilean viticulture has largely converted from parral to vertical shoot positioning (VSP) and other modern systems, though parral remains important for volume production, table grapes, and traditional production zones where the system persists. The conversion from parral to modern trellising represents one of the most dramatic viticultural transformations of recent decades, reshaping South American wine quality and international competitiveness.
Parral_Role captures the historically dominant but now declining South American pergola system, contextualizing the viticultural transformation underlying the region's quality revolution. At its peak, parral dominated Argentine viticulture—in 1990, over 80% of Argentine vineyards used parral or related overhead systems by 2020, this had declined to under 30% as quality-focused replanting proceeded. The parral-to-VSP conversion represents perhaps the largest training system transformation in global viticulture history, enabled by government incentive programs, export market demands, and technical knowledge transfer from European consultants. Parral's persistence in volume production, table grape, and traditional sectors reflects its genuine advantages—UV protection, heat stress mitigation, high yield potential—that may become increasingly relevant under climate change conditions. The system's association with bulk production has created quality stigma, though thoughtful parral management can produce balanced fruit when yields are appropriately restricted.
Parral-Erziehung
South American pergola
argentinisches Laubdach
overhead parral
parral
parral training
südamerikanische Pergola
Cavagnaro JB et al. (2006) Grapevine training systems in Mendoza, Argentina, American Journal of Enology and Viticulture 57: 113-122 Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Catania CD, Avagnina SO (2007) La Interpretación Sensorial del Vino, Caviar Bleu Pszczólkowski P (2015) Chilean viticulture history, RIVAR 2: 88-116
Parral Role
Black Corinth (Zante Currant, Korinthiaki) bears a Parthenocarpic_Table_Grape_Role as the classic example of true parthenocarpy, producing tiny berries (0.5-1.0 g) completely lacking seeds or seed traces, historically cultivated in Greece for dried currant production and occasionally for specialty fresh consumption where miniature seedless berries are novelty-valued. Canner (Convent Large Black) serves in a Parthenocarpic_Table_Grape_Role with larger berries than Black Corinth but still small by modern standards, demonstrating the berry-size limitation inherent to parthenocarpy. Breeding programs at USDA Fresno and elsewhere have explored parthenocarpy genes for transfer into larger-berried backgrounds, though stenospermocarpic seedlessness remains dominant in commercial breeding.
A seedless table grape scion role that inheres in a grafted scion cultivar by virtue of its production of completely seedless berries through parthenocarpy, a developmental process where fruit develops without pollination and fertilization, resulting in berries entirely lacking seed traces or embryonic structures. Parthenocarpic cultivars set fruit in the complete absence of pollen, developing berries from unfertilized ovaries through hormone-driven fruit growth independent of seed development signals. True parthenocarpy produces berries with no detectable seed remnants—distinguishing them from stenospermocarpic varieties with their soft seed traces—but typically results in very small berries (often 1-2 g without treatment) due to the absence of seed-derived growth hormones. Parthenocarpic table grapes are rare in commercial production compared to stenospermocarpic varieties the Black Corinth (Zante Currant) is the classic example, producing tiny but completely seedless berries primarily for dried currant production rather than fresh table grape markets. True parthenocarpy offers the advantage of complete seedlessness but the disadvantage of inherently small berry size.
Parthenocarpic_Table_Grape_Role represents a minor category in commercial table grape production due to the inherent small berry size limitation—without seed development, the berry lacks the endogenous gibberellins and auxins that drive cell division and enlargement in seeded and stenospermocarpic varieties. Black Corinth has been cultivated for over 3,000 years for dried currant production (the word "currant" derives from "Corinth"), where tiny berry size is acceptable for fresh table grape markets, stenospermocarpic varieties with their larger berries have completely dominated. Research into parthenocarpy genetics continues, with interest in combining true seedlessness genes with factors promoting berry enlargement, though commercial application remains limited. The distinction between parthenocarpy and stenospermocarpy is important for breeding strategies, as the genetic mechanisms differ fundamentally.
completely seedless grape
echt kernlose Traube
parthenocarpic grape
parthenokarpe Tafeltraube
true seedless grape
völlig samenlose Tafeltraube
Ledbetter CA, Ramming DW (1989) Seedlessness in grapes, Horticultural Reviews 11: 159-184 Stout AB (1936) Seedlessness in grapes, New York Agricultural Experiment Station Technical Bulletin 238 Varoquaux F et al. (2000) Less is better: new approaches for seedless fruit production, Trends in Biotechnology 18: 233-242 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519
Parthenocarpic Table Grape Role
Black Corinth (Zante Currant) exhibits obligate parthenocarpy, producing tiny berries (0.3-0.8 g) entirely without seeds or seed traces regardless of pollination. Stimulative parthenocarpy in some cultivars is triggered by pollination without fertilization, with pollen tube growth providing sufficient hormonal stimulus for fruit development. Parthenocarpic berries in Black Corinth average 6-8 mm diameter compared to 12-18 mm for stenospermocarpic Thompson Seedless. The VvAGL11 gene shows altered expression patterns in parthenocarpic versus stenospermocarpic seedless grapes.
A seedlessness disposition that is the inherited capacity of a Vitis vinifera plant to develop fruit without fertilization or seed formation, resulting in berries that are entirely seedless from inception rather than containing aborted seed traces, typically producing small berries due to absence of seed-derived growth hormones.
Parthenocarpy is rare in commercial grape production due to the extremely small berry size resulting from complete absence of seed-derived hormones. Stimulative parthenocarpy (triggered by pollination but not fertilization) differs from true parthenocarpy (spontaneous fruit development). Gibberellin application can partially compensate for hormone deficiency but cannot achieve berry sizes comparable to stenospermocarpic cultivars. Black Corinth is primarily used for dried fruit (currants) where small berry size is acceptable.
Jungfernfrüchtigkeit
Parthenokarpie
befruchtungslose Fruchtbildung
seedless without fertilization
true parthenocarpy
virgin fruit development
Pearson HM (1933) Parthenocarpy and seed abortion in Vitis vinifera. Proceedings of the American Society for Horticultural Science 29:169-175 Costantini L et al. (2008) Berry and phenology-related traits in grapevine (Vitis vinifera L.): from quantitative trait loci to underlying genes. BMC Plant Biology 8:38
Parthenocarpy Disposition
Chardonnay in Champagne shows partial ovule fertilization during late June-early July Year N+1 at BBCH 69-71, with typical 1.8-2.3 seeds per berry from partial ovule success across 3-4 potential ovules, within-cluster seed number range 1-3 seeds creating berry size variation, partial success providing adequate auxin rescue enabling berry development, under cool maritime conditions, typical of this early-bursting cultivar showing common partial pattern supporting 64-76% fruit set with acceptable berry size variation
Partial ovule fertilization represents the common intermediate reproductive outcome when some ovules within multi-ovule flower achieve successful fertilization producing viable seeds while other ovules fail, creating variable seed number typically 1-3 per berry instead of theoretical maximum 4 from complete success, affecting berry size through differential auxin production and representing typical fruit set pattern in grapevine. This prevalent outcome pattern occurs during fertilization in late June-early July in temperate regions Year N+1, determined within 1-3 days of pollination with seed number manifesting over subsequent 2-3 weeks, corresponding to BBCH 69-71 stages. Partial fertilization pattern shows mixed success across 2-4 ovules per flower, typical outcomes producing 1-3 viable seeds from partial success rather than complete success or complete failure, stochastic variation in which ovules succeed depending on pollen tube competition and ovule receptivity, and partial pattern representing most common fruit set outcome. Seed number distribution shows typical average 1.5-2.5 seeds per berry in many seeded cultivars, modal distribution often at 2 seeds representing common partial outcome, range from 0 (complete failure) to 3-4 (complete success) creating within-cultivar variation, and seed number heterogeneity contributing to berry size diversity. Ovule competition and selection shows 2-5 pollen tubes potentially reaching ovary region, limited ovule access creating competitive situation, first-arrival advantage and ovule receptivity determining which ovules fertilize, and stochastic elements creating variable outcomes across flowers. Successful ovule characteristics in partial pattern show ovules achieving complete double fertilization with viable embryo and endosperm, seed development proceeding normally producing mature seeds, auxin production contributing to hormonal rescue and berry growth, and viable seeds functionally equivalent to complete success outcomes. Failed ovule characteristics show ovules experiencing fertilization failure despite other ovules succeeding in same flower, abortion and degeneration creating dark collapsed tissue or seed traces, no developmental contribution except through resource reabsorption, and failed ovules representing missed potential rather than flower loss since partial success rescues flower. Hormonal rescue with partial success shows 1-2 successful seeds producing sufficient auxin preventing flower abscission, hormone levels lower than complete success but adequate for berry development, and partial success enabling berry formation albeit potentially smaller than multi-seed maximum. Berry size effects of partial success show positive correlation between seed number and final berry size, 1-seed berries smaller than 2-seed berries smaller than 3-seed berries, auxin production proportional to seed number driving differential growth response, and within-cluster berry size variation largely reflecting seed number diversity from partial fertilization. Auxin production from partial seeds shows hormone synthesis in each developing seed, total auxin proportional to seed number creating dose-dependent growth response, 1-seed berry receiving lower stimulus than 3-seed berry, and auxin gradient accounting for 30-50% berry size variation within clusters. Cell division effects show early berry cell proliferation responding to auxin levels, more seeds stimulating greater cell division creating more flesh cells, final cell number partially determined by seed number and auxin during first 3-4 weeks, and cell number differences affecting ultimate berry size potential. Berry development with partial seeds shows normal berry formation and maturation despite reduced seed number, developmental progression through all stages to full ripeness, berry quality characteristics generally normal though size reduced, and partial success representing adequate outcome for table grape and wine production. Within-cluster variation from partial pattern shows heterogeneous berry sizes within clusters, size distribution reflecting seed number diversity, cluster appearance showing mixed berry sizes though generally acceptable, and variation occasionally affecting harvest uniformity in extreme cases. Cultivar variation in partial pattern frequency shows genetic differences in typical seed number, some cultivars averaging 1-2 seeds showing frequent partial pattern, others averaging 2-3 seeds showing higher success rate, and cultivar characteristic affecting berry size uniformity. Environmental effects on partial versus complete success include favorable conditions increasing proportion of complete success with 3-4 seeds, suboptimal conditions increasing partial failure leaving 1-2 seeds, and environmental variation contributing to year-to-year seed number changes within cultivars. Partial fertilization as buffer against complete failure shows multiple ovules providing redundancy, partial success preventing flower loss even with some ovule failure, and evolutionary advantage of multi-ovule strategy ensuring fruit set despite fertilization variability. Fruit set implications show partial fertilization supporting acceptable berry formation, contributing to overall 50-70% fruit set percentage at population level, and partial pattern representing successful reproductive outcome despite not achieving maximum potential. The process represents intermediate ovule fertilization outcome within Ovule_Fertilization category between complete success and complete failure. Duration represents fertilization pattern determination within days and manifestation over 2-3 weeks. Regional timing shows partial fertilization outcomes during late June-early July across climates as fertilization results manifest. Viticultural significance includes representing most common fruit set pattern creating typical 1-3 seed berries, contributing to within-cluster berry size variation affecting harvest uniformity, demonstrating multi-ovule redundancy strategy supporting fruit set reliability, and affecting berry size through seed number effects on auxin-mediated growth. Research applications include analyzing partial fertilization determinants and patterns, quantifying seed number effects on berry characteristics, evaluating environmental and genetic factors affecting success distribution, and assessing partial fertilization as typical fruit set outcome affecting production.
Partial_Ovule_Fertilization represents common intermediate outcome within Ovule_Fertilization creating variable seed numbers. Characterized by mixed success across multiple ovules producing 1-3 seeds instead of theoretical maximum. Most frequent fruit set pattern in grapevine. Provides hormonal rescue preventing flower abortion. Creates berry size variation through differential auxin effects. Demonstrates multi-ovule redundancy strategy. Acceptable reproductive outcome supporting normal berry development. Contributes to within-cluster berry heterogeneity.
gemischter Samenanlage-Erfolg
incomplete fertilization
intermediate fertilization
mixed ovule success
teilweise Samenanlage-Befruchtung
unvollständige Befruchtung
variable Samenzahl
variable seed number
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 410-413 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Partial Ovule Fertilization
A partially coiled functional tendril at node 6 on a shoot of Vitis vinifera cv. Grüner Veltliner in Wachau, Austria, exhibiting four well-formed contact coils gripping a horizontal trellis wire at the tendril tip, but showing only one poorly developed free coil in the proximal 15 cm tendril shaft connecting to the shoot base while the remaining 12 cm section remains nearly straight, resulting from premature lignification of the basal tendril section during a brief water stress period one week after initial contact that prevented complete free coil development, providing adequate grip at the attachment point (supporting approximately 200 grams shoot weight) but minimal contractile effect with only 2 cm shoot displacement toward the wire, measured at BBCH stage 73 (berries beginning to touch) in early July.
A coiled functional tendril exhibiting incomplete coiling response with some tendril sections showing helical structure while other sections remain straight or minimally curved in Vitis vinifera, characterized by formation of contact coils at the support attachment point but absence or minimal development of free coils in the tendril shaft between support and shoot, or conversely presence of free coils without successful contact coil formation, resulting in partial mechanical function with reduced load-bearing capacity and contractile effects compared to fully coiled tendrils. Partially coiled functional tendrils develop through several mechanisms including interrupted coiling processes where initial contact and coiling begin successfully but mechanical disturbance, desiccation, or pathogen infection halts coiling progression before completion, contact with supports at tendril tips leaving insufficient length for free coil development, multiple contact points along a single tendril preventing continuous coiling between attachments, or genetic or physiological limitations in coiling capacity affecting some tendril regions while sparing others. These tendrils provide limited functional support, with contact coils alone offering grip without contractile benefits of free coils, or free coils without contact providing spring-like properties but no attachment. Partially coiled tendrils may suffice for light support requirements in low-vigor vines or favorable environmental conditions but represent compromised climbing function compared to fully developed coiling responses.
Partially coiled functional tendrils are distinguished from fully coiled tendrils by presence of straight or minimally curved sections where helical coiling would normally occur, and from non-coiled tendrils by presence of at least some coiled regions. The partial coiling pattern provides diagnostic information about limiting factors during development - interrupted free coiling suggests environmental stress during the post-contact period, absent free coils with present contact coils indicates short tendrils or tip-only contact, while absent contact coils with present free coils suggests contact loss before fixation. Partially coiled tendrils may be functionally adequate for support depending on shoot weight, wind exposure, and presence of additional attachment points from other tendrils. Some cultivars show tendencies toward partial rather than complete coiling as a characteristic trait.
abschnittsweise gewundene Kletterorgan
fragmentary coil tendril
incomplete coil tendril
sectional coil tendril
teilweise gewickelte Ranke
unvollständig spiralisierte Ranke
Based on Darwin (1875) "The Movements and Habits of Climbing Plants" John Murray, London Jaffe & Galston (1968) "The physiology of tendrils" Annual Review of Plant Physiology 19=417-434 Bowling & Vaughn (2009) "Gelatinous fibers are widespread in coiling tendrils and twining vines" American Journal of Botany 96=719-727 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 155-172.
Partially Coiled Functional Tendril
A partially developed tendril at node 7 on a shoot of Vitis vinifera cv. Riesling in Mosel Valley, Germany, measuring 12 cm length (approximately 60% of normal 18-20 cm), exhibiting well-developed vascular system and complete bifurcated architecture with both arms elongated, functional epidermis with cuticle in basal 6 cm but incomplete cuticle and thinner epidermis in distal regions, evidence of mechanosensory cells in mid-tendril regions but absent in distal 3 cm, moderate lignification in basal 4 cm providing some structural strength but insufficient for reliable load-bearing, occurring during a growing season with alternating wet and dry periods where soil moisture fluctuated between 45-75% field capacity creating variable resource availability, showing limited functional capacity with weak coiling response when experimentally contacted and supporting approximately 150 grams weight before showing strain, measured at BBCH stage 75 (berries pea-sized) in late June.
An aborted tendril that achieved intermediate developmental progress with successful completion of some but not all maturation processes in Vitis vinifera, characterized by structures showing mosaic patterns of mature and immature tissue features, completion of basic elongation growth achieving 40-80% of normal length range, successful differentiation of some tissue systems including vascular development while other systems remain incomplete such as mechanosensory apparatus or structural reinforcement, functional capability in some aspects such as basic structural presence while non-functional in critical aspects such as coiling response or load-bearing capacity, and representing a heterogeneous category of developmental outcomes between clear early abortion and near-complete development. Partially developed tendrils result from fluctuating stress conditions or intermittent resource availability during tendril development creating periods of successful growth alternating with periods of developmental constraint, progressive stress conditions that allow initial development under favorable conditions followed by increasing limitation, localized damage or disease affecting specific tissue regions while sparing others, or developmental asynchrony within the tendril where some regions mature successfully while others lag behind and ultimately fail. These structures represent intermediate resource investment with partial achievement of functional potential, potentially providing limited climbing support in favorable conditions but unreliable under normal vineyard loading, and creating diagnostic challenges in assessing developmental success versus failure requiring detailed examination of specific tissue systems and functional capabilities rather than simple binary classification.
Partially developed tendrils occupy an intermediate category between clear developmental failures (early and mid-development aborted tendrils) and near-successes (near-mature aborted tendrils and functional tendrils), characterized by heterogeneous achievement across different developmental parameters rather than uniform developmental stage. Classification requires assessment of multiple features including length achievement, tissue differentiation status across systems (vascular, epidermal, mechanical, sensory), structural competence, and functional capability, with partially developed status indicated by success in some parameters combined with failure in others. This category is diagnostically valuable because it reveals asynchronous development or fluctuating stress conditions rather than uniform stress or optimal conditions. The specific pattern of successful versus failed developmental features provides information about the type and timing of limiting factors, with vascular success but sensory failure suggesting early favorable conditions followed by late stress, while the reverse pattern indicates different stress timing.
gemischte Reife Kletterorgan
incompletely developed tendril
mixed maturity tendril
partial success tendril
teilweise entwickelte Ranke
unvollständig ausgereifte Ranke
Based on Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Petrie et al. (2000) "The effect of leaf removal and canopy height on whole-vine gas exchange and fruit development of Vitis vinifera L. Sauvignon blanc" Functional Plant Biology 27=173-184 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 109-134 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 473-496.
Partially Developed Tendril
A partially expanded young leaf at node 8 on a shoot of Vitis vinifera cv. Sangiovese in Tuscany, Italy, measuring 13.5 cm length by 12.8 cm width representing approximately 78% of expected mature dimensions based on cultivar and position norms, exhibiting well-defined five-lobed architecture with all structural features evident including complete sinus development and marginal tooth patterns, medium-dark green coloration with chlorophyll content of 425 μg cm⁻² approaching mature levels, virtually no anthocyanin pigmentation except very slight bronze tinge at major vein junctions, moderately firm texture with developing rigidity from cell wall maturation and beginning lignification in vascular tissues, two well-developed palisade layers totaling 78 μm thickness (92% of mature 85 μm) and spongy mesophyll of 112 μm (93% of mature 120 μm), functional stomata with appropriate responses showing conductance of 185 mmol H₂O m⁻² s⁻¹ under favorable conditions, substantial cuticle development evident from glossy surface appearance, demonstrating photosynthesis rate of 11.2 μmol CO₂ m⁻² s⁻¹ representing 84% of mature capacity and achieving approximate carbon balance with daily photosynthetic gain of 52 mg carbohydrate balancing respiratory costs and minimal remaining growth requirements, occurring at 24 days post-emergence approaching maturity transition, measured at shoot BBCH stage 79 (majority of berries touching) in late July, representing late expansion phase with maturity expected within 7-10 days.
An expanding young leaf that has completed substantial but not full expansion achieving intermediate dimensions between early expansion and mature size in Vitis vinifera, characterized by dimensions typically 50-85% of final mature size with continued but declining expansion rates, substantially flat lamina with complete lobe definition and sinus development showing final architectural characteristics, medium to medium-dark green coloration approaching mature chlorophyll concentrations typically 350-500 μg cm⁻², greatly reduced or absent anthocyanin pigmentation compared to early stages indicating declining photoprotection requirement, moderately firm texture approaching mature consistency as cell walls complete thickening and lignification begins, advanced tissue differentiation with palisade and spongy mesophyll layers well-developed approaching mature organization, nearly complete vascular development with functional xylem and phloem showing mature secondary walls, fully functional stomata responding appropriately to environmental signals, substantial cuticle development with most wax deposition complete, and photosynthetic capacity typically 70-90% of mature leaf rates approaching or achieving carbon balance where daily photosynthetic gains equal or slightly exceed respiratory and growth costs making these leaves transitional between sink and source status. Partially expanded young leaves represent the late expansion phase approaching maturity, with cell expansion declining as cells approach final dimensions, final size determination largely complete with remaining growth representing final adjustments, tissue maturation processes dominating including cell wall completion, chloroplast maturation, and metabolic activation, and functional capabilities approaching mature levels with leaves beginning to contribute meaningfully to vine carbon balance, with the partially expanded phase typically spanning 5-10 days before transition to fully mature status.
Partially expanded young leaves represent the transition zone between active expansion and mature status, distinguished from earlier expansion stages by substantial size achievement and tissue maturation, and from mature leaves by incomplete final dimensions and continued growth processes. Assessment of partially expanded status requires comparison to cultivar-specific and position-specific mature size norms, as absolute dimensions are insufficient given variation in final leaf size. The partially expanded phase represents critical functional transition where leaves shift from net carbon sinks to approaching or achieving carbon balance, beginning to contribute to rather than drain vine carbon resources. Environmental conditions during partial expansion affect final size and functional characteristics, with stress during this phase having reduced impact compared to early expansion as most size determination and anatomical differentiation is already complete. Photosynthetic capacity development during partial expansion is critical for timely achievement of source function, with delayed chloroplast maturation or chlorophyll synthesis creating leaves that achieve final size but show reduced functional capacity requiring extended maturation periods.
advanced expansion leaf
fast-reifes Blatt
late development leaf
near-mature leaf
späte Entwicklung Weinblatt
teilweise expandiertes junges Blatt
Based on Poni et al. (2009) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 60=397-407 Palliotti & Cartechini (2000) "Developmental changes in gas exchange activity in flowers, berries, and tendrils of field-grown Cabernet Sauvignon" American Journal of Enology and Viticulture 51=317-323 Flexas et al. (2002) "Drought-inhibition of photosynthesis in C₃ plants= stomatal and non-stomatal limitations revisited" Annals of Botany 89=183-189 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 185-208.
Partially Expanded Young Leaf
A partially lignified one-year cane on Vitis vinifera cv. Pinot Noir in a cool growing season, examined during November pre-pruning assessment showing brown woody appearance for basal 85cm (nodes 1-8) with firm structural rigidity, transitioning to brown-green appearance for middle 30cm (nodes 9-10) with moderate firmness, and retaining green soft appearance for apical 35cm (nodes 11-13) indicating incomplete lignification, pruning decision to retain 8 nodes from fully lignified section while removing partially and unlignified portions to avoid winter damage and poor budbreak from immature wood, accepting reduced cane length and node count to maintain wood quality.
A lignified one-year cane showing incomplete secondary cell wall development, characterized by mixed brown and green coloration indicating variable lignification progress along the cane length, typically with well-lignified basal regions transitioning to incompletely lignified apical sections, moderate structural firmness in brown sections but retained flexibility or softness in green portions, and reduced cold hardiness in unlignified regions creating vulnerability to winter damage. Partially lignified one-year canes represent suboptimal maturation outcomes resulting from late-season growth continuation, insufficient growing degree day accumulation, excessive vigor from high nitrogen or water availability, heavy crop loads depleting carbohydrate reserves, or premature growth cessation from early cold weather, requiring careful evaluation during pruning to retain only adequately lignified portions while removing vulnerable green or semi-lignified tips.
Partially lignified one-year canes require careful pruning assessment to identify the transition point between adequately mature (safe to retain) and inadequately mature (should be removed) wood. The lignification boundary is not always sharp, with gradual transition zones showing intermediate characteristics requiring judgment based on color, firmness, and sometimes destructive cross-sectional examination. Retention of partially lignified portions as fruiting wood risks winter damage, poor spring budbreak, weak shoot development, and reduced yields, making conservative pruning (removing questionable sections) generally preferable to aggressive retention. In short-season climates, partial lignification is common and expected, requiring systemic management adjustments including earlier-maturing cultivars, crop load reduction, canopy management for enhanced ripening, and acceptance of shorter cane retention.
incompletely lignified cane
mixed maturity cane
partially mature cane
teilweise verholzte Rute
unvollständig ausgereifte Rute
Partially Lignified One Year Cane
Merlot flowers during active bloom (BBCH 60-61) exhibit partially separated calyptras with 70-85% of basal circumference detached creating 1.5-2 mm gaps, calyptra displaced upward 2-2.5 mm and canted 15-30° from vertical due to retention of 1-2 attachment points on one side, three to four stamens fully visible with extended filaments (2.8-3.5 mm) and anthers beginning dehiscence, pistil top sometimes visible above tilted calyptra edge, with complete detachment occurring 40-70 minutes later under morning conditions (temperatures rising from 19°C to 24°C, humidity declining from 75% to 55%). Chardonnay at BBCH 61 shows partially separated calyptras across 12-18% of flowers simultaneously, creating characteristic appearance of emerging stamens with yellowish calyptra caps tilted at various angles still tenuously attached. Pinot Noir experiencing gusty winds (15-25 km/h) during bloom shows accelerated transition from partially separated to detached calyptras as wind forces dislodge calyptras once partial separation reduces attachment strength, with 80-90% of partially separated calyptras completing detachment within 20-40 minutes compared to typical 50-80 minutes under calm conditions.
A partially separated calyptra is a separating calyptra in mid-to-late separation stage showing substantial upward displacement and extensive basal detachment but retaining narrow connection points maintaining calyptra attachment to the receptacle, with visible exposure of multiple stamens and often partial pistil visibility as the lifted calyptra creates progressively larger gaps between calyptra edge and receptacle, representing flowers in the final 30-90 minutes before complete detachment and full anthesis. Partially separated calyptras are characterized by= (1) basal separation completed around 60-90% of the circumference with visible gaps 1-2 mm wide exposing the bases of elongating stamen filaments (2) calyptra displaced upward 1.5-3 mm from original pre-separation position, lifted by continuing stamen filament elongation but still tethered by one to several narrow attachment points where abscission layer separation has not yet completed (3) three to five stamens visible with filaments extended and emerging from beneath the lifted calyptra edge, with anthers beginning to become visible though often still partially enclosed by the calyptra apex (4) calyptra often canted or tilted at an angle rather than lifted vertically, as asymmetric retention of attachment points on one side creates unbalanced lifting (5) visible mechanical stress at remaining attachment points evidenced by tissue stretching or tearing as upward pressure from stamens pulls against the diminishing connections. The partially separated state represents the penultimate phase of anthesis, with complete detachment typically occurring 30-90 minutes after the partially separated state becomes established, though duration varies with temperature (shorter at higher temperatures 24-30°C, longer at lower temperatures 16-20°C) and with the strength and number of remaining attachment points. During the partially separated phase, critical reproductive developments proceed= stamen filaments complete their rapid elongation phase, extending from approximately 2-2.5 mm at early separation to final hermaphrodite length 3-4 mm by the time partial separation is well advanced anthers complete positioning at or above pistil level (in hermaphrodite flowers) or well below pistil level (in functionally female or pistillate flowers with reflex stamens) anther dehiscence often begins during late partial separation phase, with longitudinal dehiscence slits opening and initial pollen release occurring even before calyptra is fully detached stigma begins exudate production in preparation for receptivity, with wet glistening stigma surface sometimes visible if pistil top emerges above the partially separated calyptra. The partially separated calyptra creates a transitional microenvironment with= (1) decreasing humidity as gaps allow air exchange, reducing internal humidity from the 85-95% of intact calyptras toward ambient levels (2) increasing temperature fluctuation as exposure to external air eliminates the buffering effect of the sealed calyptra (3) beginning of pathogen access to floral tissues as gaps permit Botrytis cinerea spores or other pathogens to reach exposed stamen and pistil surfaces. Partially separated calyptras are mechanically fragile due to the combination of reduced attachment strength, tissue senescence, and physical stress from stamen pressure, making them susceptible to premature detachment from rain impact, wind gusts, or mechanical disturbance, though such premature detachment typically does not impair subsequent flower function as reproductive organs have achieved sufficient maturity for successful pollination and fruit set. The observation of partially separated calyptras within an inflorescence indicates active bloom in progress, typically coinciding with BBCH 60-61 (beginning of flowering, 5-20% of flowers open), and provides growers with confirmation that the critical bloom period requiring optimal environmental conditions and disease management has commenced.
Partially_Separated_Calyptra represents the mid-to-late separation phase and is distinguished from Loosening_Calyptra (early separation with minimal displacement) by substantial upward displacement (>1.5 mm versus 60% versus <50%), and advanced stamen emergence. The partially separated state is the most visually obvious phase of active anthesis and is the easiest separation stage to observe without magnification, making it valuable for practical bloom monitoring. Partially separated calyptras coincide with critical reproductive transitions (anther dehiscence beginning, stigma receptivity initiating) and represent the transition from protected to exposed flower status, marking the period when disease infection risk increases and when environmental conditions most directly affect flower function and subsequent fruit set success.
halb getrennte Kelchhaube
mid-separation calyptra
partially lifted petal cap
teilweise abgelöste Blütenhaube
teilweise abgetrennte Kronblattkappe
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 101-103 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 193-194.
Partially Separated Calyptra
Trebbiano Toscano and Malvasia del Chianti bear Passito_Style_Scion_Roles for Vin Santo del Chianti, where grapes are dried on racks (appassimento) for 3-6 months achieving 30-40°Brix, then fermented and aged in small sealed barrels (caratelli) for 3-10+ years, producing wines of 80-150 g/L residual sugar with hazelnut, caramel, and dried apricot complexity. Corvina, Rondinella, and Molinara serve in Passito_Style_Scion_Roles for Recioto della Valpolicella, dried for 100-120 days to 25-30% weight loss before fermentation arrests naturally at 12-14% alcohol with 100-150 g/L residual sugar. Savagnin bears a Passito_Style_Scion_Role for Vin de Paille du Jura, with straw-mat drying concentrating grapes to 300+ g/L sugar for wines of extraordinary nutty, honeyed complexity aged 3+ years in barrel.
A dessert wine scion role that inheres in a grafted scion cultivar by virtue of its suitability for passito (appassimento/vin de paille/strohwein) production, where harvested grapes are dried post-harvest to concentrate sugars through controlled desiccation before pressing and fermentation. Passito scions must possess thick, intact skins that permit controlled water loss (typically 30-50% weight reduction) without mold development, appropriate cluster architecture for air circulation during drying (loose bunches, well-separated berries), and sufficient acidity to balance concentrated sweetness (residual sugar typically 80-250 g/L). Drying occurs over weeks to months in controlled environments—historically on straw mats (paglia), reed racks (arele), or hung from rafters (graticci)—increasingly in temperature and humidity-controlled facilities (fruttai). The extended drying period concentrates sugars while developing characteristic dried fruit, raisin, caramel, and oxidative aromas through enzymatic and non-enzymatic browning reactions. Major passito traditions include Italian Vin Santo (Tuscany), Recioto and Amarone della Valpolicella (Veneto), and French Vin de Paille (Jura).
Passito_Style_Scion_Role captures post-harvest drying traditions distinct from on-vine late harvest concentration or botrytis transformation. The controlled drying environment is critical—traditional methods used drafty attics or specialized drying lofts (fruttai), while modern facilities control temperature (10-15°C) and humidity (60-70%) to prevent mold while permitting gradual dehydration. Corvina's importance for Amarone (technically a dry passito style where fermentation proceeds to completion) demonstrates that appassimento technique can produce both sweet (Recioto) and dry (Amarone) wines from the same variety and process. The caramelization and oxidation during extended drying create flavor profiles impossible to achieve through other concentration methods, distinguishing passito wines from late harvest or botrytis styles.
Passito-Sorte
Strohwein grape
Strohweinsorte
Trockenbeeren-Sorte
appassimento variety
dried grape variety
passito variety
vin de paille grape
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Meneghetti S et al. (2014) Drying techniques for traditional Amarone grape production, Italian Journal of Food Science 26: 233-240 Jackson RS (2014) Wine Science 4th Ed, Academic Press pp.: 545-555 Bellincontro A et al. (2004) Different postharvest dehydration rates affect quality characteristics in Malvasia, Trebbiano and Sangiovese grapes, Postharvest Biology and Technology 34: 107-116
Passito Style Scion Role
A disease-resistant Muscadinia rotundifolia genotype serves in a Paternal_Parent_Role, its pollen applied to emasculated V. vinifera flowers to generate interspecific hybrids evaluated for resistance introgression. Cabernet Sauvignon bears a Paternal_Parent_Role in quality improvement crosses, its pollen collected and applied to breeding lines where Cabernet's quality traits are targeted for combination with other characteristics. A seedless (stenospermocarpic) table grape variety functions exclusively in a Paternal_Parent_Role for seedlessness breeding, as its seed parent use would yield no viable seeds due to embryo abortion.
A parent role that inheres in a grapevine genotype by virtue of its function as the male parent in a controlled cross, contributing pollen containing the male gamete (sperm cell) that fertilizes the egg cell within the maternal parent's ovule. The paternal parent contributes one haploid chromosome set to the hybrid embryo's nuclear genome, combining with the maternal nuclear contribution to produce the diploid offspring genotype. In controlled grapevine breeding, pollen is collected from anthers of the paternal parent (either before natural dehiscence or at anthesis) and applied to emasculated flowers of the maternal parent, with fertilization and seed development occurring on the maternal vine. The paternal parent's pollen characteristics—quantity produced, viability, storage tolerance, and germination reliability—influence its practical utility as a crossing partner, with some genotypes preferred as pollen donors due to excellent pollen performance while others are more difficult to use paternally. Unlike the maternal parent, the paternal parent contributes no cytoplasmic genetic material to standard offspring (though rare paternal plastid inheritance may occur).
Paternal_Parent_Role is subclassed by Pollen_Parent_Role, which emphasizes the pollen production and donation function specifically. The distinction between paternal and maternal roles reflects differential gametic contribution (pollen versus ovule) and practical breeding considerations (pollen collection and storage versus seed production and germination). Some genotypes are preferentially used as paternal parents due to excellent pollen production, viability, or storage characteristics, while others (particularly seedless varieties) can only function paternally due to reproductive biology constraints. Paternal parent genetic contribution to offspring is exclusively nuclear (chromosomal) with rare exceptions, contrasting with the additional cytoplasmic contribution from maternal parents. Grapevine pollen can be stored successfully (refrigerated or frozen with appropriate preparation), enabling crosses between parents with non-overlapping flowering periods and facilitating international germplasm exchange.
Pollenspender
Vaterpflanze
father plant
male parent
männlicher Elter
paternal parent
pollen parent
väterlicher Elter
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp.: 78-112 Antcliff AJ (1988) Grape breeding in Australia, Proceedings 2nd International Symposium for Cool Climate Viticulture
Paternal Parent Role
Pathogen infection indicator quality monitoring in Bordeaux vineyards established integrated disease assessment protocols for the major fungal pathogens affecting regional grape production. Downy mildew (Plasmopara viticola) pathogen infection indicator quality assessment combined detection of the characteristic "oil spot" chlorotic lesions on upper leaf surfaces with confirmation of white sporulation on corresponding lower leaf positions during humid conditions, enabling definitive diagnosis distinct from other causes of leaf chlorosis. Powdery mildew (Erysiphe necator) pathogen infection indicator quality showed the distinctive white powdery mycelial growth on leaf, shoot, and cluster surfaces, with early infections on susceptible cultivars detectable as subtle chlorotic patches before sporulation became visible. Botrytis bunch rot (Botrytis cinerea) pathogen infection indicator quality monitoring focused on detection of the gray sporulating mycelium on ripening berries, with early indicators including berry skin browning and softening preceding the characteristic fuzzy gray appearance of advanced infection. The integrated pathogen infection indicator quality program combined weekly visual scouting with weather-based infection risk modeling to optimize fungicide timing, with indicator assessment providing ground-truth validation of model predictions and enabling adjustment of spray schedules based on actual disease pressure observed in the vineyard.
Pathogen infection indicator quality is a stress response quality that inheres in a grapevine organism and represents the ensemble of visual symptoms, physiological changes, and diagnostic markers that signal the presence and activity of plant pathogens including fungi, oomycetes, bacteria, viruses, and phytoplasmas, providing the diagnostic basis for identifying disease occurrence, assessing infection severity, and monitoring disease progression to inform management decisions aimed at limiting pathogen spread and minimizing crop and vine damage. This quality captures the plant's observable response to pathogen colonization and the diagnostic features that enable pathogen identification, recognizing that effective disease management depends on accurate and timely recognition of infection through symptoms that may range from subtle physiological changes detectable only through specialized testing to dramatic tissue destruction clearly visible to casual observation. The ontological significance of pathogen infection indicator quality lies in its role enabling disease diagnosis and management decision-making, with the recognition and interpretation of infection indicators providing the foundation for integrated pest management programs that deploy control measures appropriately based on accurate assessment of disease presence and severity. The biological basis of pathogen infection indicators encompasses the direct effects of pathogen activity on host tissues including cell death, tissue discoloration, and structural disruption, the host defense responses that may themselves produce visible symptoms including hypersensitive response and phytoalexin accumulation, and the systemic effects on plant physiology including growth reduction, yield loss, and quality compromise that may manifest without localized visible symptoms. The pathogen-specific nature of many infection indicators enables differential diagnosis distinguishing among the numerous pathogens that may affect grapevine, with characteristic symptom patterns, host tissue preferences, and disease development dynamics providing the diagnostic features that identify specific pathogens and guide appropriate management responses. The temporal dynamics of infection indicator development from initial invisible infection through early symptom expression to advanced disease reflect the progression of host-pathogen interaction, with early detection enabling intervention before extensive damage occurs while late detection may limit management options to preventing further spread rather than protecting already-affected tissues.
Pathogen_Infection_Indicator_Quality is_a Stress_Response_Quality, serving as parent class for Disease_Severity_Quality which addresses quantitative assessment of infection extent. This class addresses the overall concept of pathogen detection and diagnosis while child hierarchies capture specific measurement dimensions. Pathogen-specific symptoms enable differential diagnosis among the numerous grapevine pathogens. The temporal dynamics from invisible infection to visible symptoms affect management timing options. Critical for integrated pest management decision-making in vineyard disease control.
Infektionsmarker
Krankheitsdiagnose-Merkmale
Krankheitssymptome
Pathogen-Infektionsindikatoren
disease diagnostic features
disease indicators
infection symptoms
pathogen damage markers
Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests, 2nd Edition. APS Press Pearson RC, Goheen AC (1988) Compendium of Grape Diseases. APS Press Gadoury DM et al. (2012) Grapevine powdery mildew (Erysiphe necator): a fascinating system for the study of the biology, ecology and epidemiology of an obligate biotroph. Molecular Plant Pathology 13:1-16
Pathogen Infection Indicator Quality
The Vitis vinifera cultivar 'Chasselas' exhibits minimal Pathogen_Resistance_Disposition against most fungal and oomycete pathogens, requiring 12-16 fungicide applications per season in humid Central European climates to control Plasmopara viticola, Erysiphe necator, and Botrytis cinerea. In contrast, the interspecific hybrid 'Solaris' (Merzling × Gm 6493) possesses multiple Pathogen_Resistance_Dispositions inherited from its complex pedigree including V. amurensis and V. rupestris ancestry. These dispositions are realized through rapid defense activation upon pathogen contact, enabling cultivation with only 2-4 fungicide applications annually while maintaining infection levels below 5% leaf area affected for downy mildew and below 10% cluster infection for powdery mildew under moderate disease pressure. This 70-80% reduction in plant protection inputs demonstrates the practical significance of pathogen resistance dispositions for economically and environmentally sustainable grape production.
A Pathogen_Resistance_Disposition is a Biotic_Stress_Resistance_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through defense processes upon encounter with microbial organisms capable of causing disease, including fungi, oomycetes, bacteria, viruses, and phytoplasmas. This disposition represents the genetically-encoded capacity to limit pathogen ingress, colonization, reproduction, or spread through diverse molecular and cellular mechanisms operating at pre-invasive and post-invasive stages of infection. Pre-invasive resistance mechanisms include physical barriers such as cuticle thickness and stomatal morphology, as well as preformed antimicrobial compounds. Post-invasive mechanisms encompass pathogen recognition through pattern recognition receptors (PRRs) detecting pathogen-associated molecular patterns (PAMPs) and through nucleotide-binding leucine-rich repeat (NLR) receptors recognizing pathogen effectors, triggering downstream defense responses including reactive oxygen species production, cell wall reinforcement, phytoalexin biosynthesis, and programmed cell death. Pathogen resistance dispositions in grapevine vary from monogenic qualitative resistance conferring race-specific immunity to polygenic quantitative resistance providing partial but durable protection. The introgression of pathogen resistance dispositions from wild Vitis species into V. vinifera cultivars represents a primary breeding objective for sustainable viticulture, addressing the exceptional susceptibility of European grapevine cultivars to numerous pathogens introduced through global trade.
Pathogen_Resistance_Disposition is_a Biotic_Stress_Resistance_Disposition. This class encompasses resistance to all disease-causing microorganisms, distinguished from Pest_Resistance_Disposition which covers resistance to arthropods and nematodes. Child classes are organized taxonomically by pathogen group: Fungal_Resistance_Disposition (including oomycetes by convention), Viral_Resistance_Disposition, Bacterial_Resistance_Disposition, and Phytoplasma_Resistance_Disposition. Note that oomycetes such as Plasmopara viticola are phylogenetically distinct from true fungi but are conventionally grouped under fungal diseases in viticultural practice.
Erregerabwehr-Potenzial
Krankheitserreger-Resistenz
Pathogenresistenz-Disposition
microbial resistance potential
pathogen defense capacity
Jones JDG, Dangl JL (2006) The plant immune system, Nature 444:323-329 Gessler C et al. (2011) Vitis vinifera and grape diseases, Genetics, Genomics and Breeding of Grapes, Science Publishers Armijo G et al. (2016) Grapevine pathogenic microorganisms: understanding infection strategies and host response scenarios, Frontiers in Plant Science 7:382
Pathogen Resistance Disposition
Systematic analysis of Pathogenesis_Related_Protein_Disposition at Agroscope Changins-Wädenswil compared PR protein induction across 20 cultivars following Erysiphe necator inoculation. Leaf samples collected at 0, 24, 48, 72, and 120 hpi were analyzed for PR-2 (glucanase), PR-3 (chitinase), and PR-5 (thaumatin-like protein) by Western blot densitometry and enzyme activity assays. Resistant 'Regent' exhibited strong Pathogenesis_Related_Protein_Disposition with rapid induction kinetics. PR-3 chitinase activity increased 12-fold by 24 hpi, reaching peak at 48 hpi (28-fold above basal), while PR-2 glucanase showed 8-fold induction by 48 hpi. PR-5 accumulation was slower but sustained, reaching 15-fold at 120 hpi. Susceptible 'Chasselas' showed delayed, weak induction with PR-3 reaching only 3-fold at 72 hpi and PR-2 showing minimal response (1.5-fold). The combined PR protein index (sum of standardized fold-changes) correlated strongly with powdery mildew resistance (r = -0.78 with colony density at 120 hpi). Notably, constitutive PR levels also varied, with 'Regent' showing 2.5-fold higher basal PR-3 than 'Chasselas', suggesting that both basal and induced components of Pathogenesis_Related_Protein_Disposition contribute to resistance. When purified PR protein preparations were tested in vitro, grapevine chitinase inhibited E. necator conidial germination by 65% at 50 μg/mL, confirming direct antifungal function.
A Pathogenesis_Related_Protein_Disposition is an Induced_Defense_Disposition that inheres in a Vitis vinifera plant and is realized through rapid and strong induction of pathogenesis-related (PR) proteins—families of defense-related proteins whose expression is dramatically upregulated in response to pathogen attack, elicitor treatment, or activation of systemic acquired resistance. This disposition represents the protein component of induced defense, complementing phytoalexin production through direct antimicrobial activities including cell wall degradation, membrane permeabilization, and pathogen enzyme inhibition. Grapevine expresses representatives of most characterized PR protein families, with PR-2 (β-1,3-glucanases), PR-3 (chitinases), PR-4 (chitin-binding proteins), PR-5 (thaumatin-like proteins), PR-10 (ribonuclease-like proteins), and PR-14 (lipid transfer proteins) being most prominent in defense responses. PR proteins function through diverse mechanisms targeting pathogen structures and processes. Glucanases degrade fungal cell wall β-glucans, chitinases hydrolyze chitin (a major component of fungal cell walls and insect exoskeletons), and thaumatin-like proteins disrupt pathogen membranes. The magnitude and speed of PR protein induction vary substantially among grapevine genotypes, with resistant cultivars typically showing stronger basal expression and more rapid, sustained induction compared to susceptible genotypes. Importantly, PR protein activity contributes to both local resistance at infection sites and systemic acquired resistance providing whole-plant protection.
Pathogenesis_Related_Protein_Disposition is_a Induced_Defense_Disposition. PR proteins comprise multiple families with distinct antimicrobial mechanisms, addressed through child classes Chitinase_Induction_Disposition and Glucanase_Induction_Disposition for the most prominent families in antifungal defense. PR proteins contribute to both local defense and systemic acquired resistance (SAR). Some PR proteins are expressed at basal levels in healthy tissue (addressed under Preformed_Antimicrobial_Disposition), with the present class focusing on inducible accumulation upon challenge.
Abwehrprotein-Akkumulation
PR protein induction
PR-Protein-Induktion
Pathogenese-bezogene-Protein-Disposition
defense protein accumulation
pathogenesis-related gene expression
van Loon LC et al. (2006) Significance of inducible defense-related proteins in infected plants, Annual Review of Phytopathology 44:135-162 Ferreira RB et al. (2007) The wine proteins, Trends in Food Science and Technology 18:293-306 Monteiro S et al. (2003) Structural and functional aspects of PR-10 proteins, FEBS Letters 541:1-7
Pathogenesis Related Protein Disposition
Pathogenesis-related protein induction quality assessment in grapevine leaves following Plasmopara viticola inoculation characterized the temporal dynamics of PR gene expression and protein accumulation contributing to downy mildew defense. Quantitative RT-PCR analysis of PR gene expression revealed pathogenesis-related protein induction quality with rapid upregulation of VvPR1 (85-fold by 24 hours), VvPR2 (β-1,3-glucanase, 45-fold), VvPR3 (chitinase, 62-fold), and VvPR5 (thaumatin-like, 38-fold) in the resistant V. rupestris compared to more modest and delayed induction in susceptible V. vinifera Chardonnay showing only 8-12 fold induction of the same genes with peak expression delayed to 48-72 hours. Protein accumulation assessed by Western blot and enzyme activity assays confirmed the pathogenesis-related protein induction quality differential, with V. rupestris reaching maximum chitinase activity of 2.8 nkat/mg protein at 48 hours versus 0.9 nkat/mg in Chardonnay, and glucanase activity of 1.5 nkat/mg versus 0.5 nkat/mg. In vitro inhibition assays demonstrated that protein extracts from induced V. rupestris tissue inhibited P. viticola sporangial germination by 75% versus 30% inhibition by Chardonnay extracts, establishing functional relevance of the pathogenesis-related protein induction quality differential. The assessment established that both the speed and magnitude of PR protein induction contribute to the enhanced resistance of V. rupestris, with the early accumulation of antimicrobial proteins creating an inhospitable environment for pathogen establishment during the critical early infection period.
Pathogenesis-related protein induction quality is a defense response quality that inheres in a grapevine organism and represents the capacity to upregulate expression and accumulation of pathogenesis-related (PR) proteins in response to pathogen infection, with PR proteins contributing to defense through diverse mechanisms including direct antimicrobial activity by chitinases, glucanases, and thaumatin-like proteins that degrade pathogen cell walls or membranes, protease inhibitors that interfere with pathogen virulence factors, and other activities that collectively contribute to pathogen growth inhibition and disease resistance. This quality captures the protein-based dimension of induced defense, recognizing that PR protein accumulation represents a conserved component of plant immune responses with demonstrated contribution to resistance against diverse pathogen types through their varied antimicrobial activities. The ontological significance of pathogenesis-related protein induction quality lies in its role as both a marker of defense activation useful for assessing plant immune status and a functional contributor to resistance where PR protein levels correlate with defense effectiveness, providing targets for transgenic enhancement through PR gene overexpression and for selection of cultivars with superior induction capacity. The biological basis of PR protein induction encompasses the transcriptional activation of PR genes by defense signaling pathways, the translation and post-translational processing of PR proteins including signal peptide cleavage and glycosylation, and the secretion of mature PR proteins to the apoplast where many exert their antimicrobial functions. The PR protein families defined by structural and functional criteria include PR-1 proteins of uncertain function but strongly induced by salicylic acid, PR-2 β-1,3-glucanases that degrade fungal cell wall glucans, PR-3, -4, -8, and -11 chitinases that degrade chitin in fungal cell walls, PR-5 thaumatin-like proteins with membrane permeabilization activity, and numerous other families with specific activities contributing to the overall antimicrobial environment created by PR protein accumulation. The temporal dynamics of PR protein induction show characteristic patterns with rapid transcriptional upregulation within hours of pathogen recognition, protein accumulation over subsequent days, and sustained elevation that may persist for weeks providing extended protection.
Pathogenesis_Related_Protein_Induction_Quality is_a Defense_Response_Quality, representing the upregulation of pathogenesis-related proteins with antimicrobial activities. This class addresses the protein-based defense dimension. PR families include chitinases (PR-3/4/8/11), glucanases (PR-2), thaumatin-like (PR-5), and others. Both induction speed and magnitude determine defense effectiveness. Salicylic acid pathway primarily regulates PR-1 and other PR genes. Provides both resistance markers and functional contributions to defense. Conserved component of plant immunity across species.
Abwehrprotein-Expression
Pathogenese-Protein-Induktion
antimicrobial protein accumulation
antimikrobielle Protein-Akkumulation
defense protein expression
pathogenesis-related response
Van Loon LC et al. (2006) Significance of inducible defense-related proteins in infected plants. Annual Review of Phytopathology 44:135-162 Dufour MC et al. (2016) Assessment of fungal pathogen resistance mechanisms in grapevine cultivars. Plant Pathology 65:1404-1419 Armijo G et al. (2016) Grapevine pathogenic microorganisms: understanding infection strategies and host response scenarios. Frontiers in Plant Science 7:382
Pathogenesis Related Protein Induction Quality
Pathogenesis-related protein quality in Marlborough Sauvignon Blanc reaches particularly high levels (120-200 mg/L TLP plus chitinases) reflecting both cultivar genetics and the region's disease pressure from Botrytis that induces PR protein expression even in visually healthy fruit winemakers employ heat stability testing (80°C/6 hours, targeting less than 2 NTU increase) to determine bentonite requirements typically ranging from 0.6-1.2 g/L, with the substantial bentonite doses required for Sauvignon Blanc stability creating ongoing research interest in alternative stabilization technologies that might preserve the aromatic intensity and texture that bentonite can diminish.
Pathogenesis_Related_Protein_Quality represents the chemical quality characterizing the concentration of pathogenesis-related (PR) proteins in grape must, capturing the defense-associated protein class primarily responsible for protein haze instability in white wines. PR proteins are induced in grape berries as part of the plant defense response to pathogen challenge, mechanical damage, and various abiotic stresses, accumulating to become the dominant protein class in mature grape juice. The major PR proteins in grapes include thaumatin-like proteins (TLPs, PR-5 family, 20-25 kDa) and chitinases (PR-3 family, 25-35 kDa), both of which possess antifungal activity and accumulate to high concentrations (50-200 mg/L combined) in ripe fruit regardless of visible disease presence. These PR proteins are particularly problematic for wine stability because they are heat-labile, denaturing and aggregating at temperatures encountered during wine storage, transport, or temperature abuse (above 25-30°C), producing the visible haze that renders white wine commercially unacceptable. The degree of heat lability varies with protein class and wine matrix, with thaumatin-like proteins generally more stable than chitinases but both contributing to total haze potential. Cultivar genetics strongly affects PR protein accumulation, with Sauvignon Blanc, Semillon, and Muscat varieties showing characteristically high levels while Chardonnay and Riesling typically accumulate less. Berry maturity correlates with PR protein content, as accumulation continues during ripening. The standard heat stability test (heating wine at 80°C for defined periods) provides practical assessment of PR protein-related haze risk, guiding bentonite fining requirements. Alternative stabilization approaches including flash pasteurization, ultrafiltration, and novel fining agents continue to be developed to address PR protein instability with reduced wine impact.
Pathogenesis_Related_Protein_Quality specifically addresses the PR protein classes (primarily TLPs and chitinases) responsible for heat-induced haze rather than total protein content. The defense function of PR proteins explains their stress-responsive accumulation and persistence in mature fruit. The heat stability test provides practical assessment of PR protein impact without requiring protein quantification, though understanding protein content enables prediction and process optimization. The wine industry terminology "protein haze" specifically refers to PR protein-mediated instability rather than other protein-related phenomena.
PR protein content
PR-Proteingehalt
Pathogenese-assoziiertes Proteingehalt
Thaumatin-ähnliches Proteingehalt
Verteidigungsproteinqualität
chitinase content quality
defense protein quality
thaumatin-like protein concentration
Waters EJ et al. (1996) The Pathogenesis-Related Proteins of Wine: Purification and Protein Sequence Analysis of Thaumatin-Like Proteins and Chitinases. Journal of Agricultural and Food Chemistry 44(1): 3-5 Pocock KF et al. (2000) Limitations to the Use of Electrophoretic Procedures for the Quantification of Proteins in Grape Juice and Wine. American Journal of Enology and Viticulture 51(3): 231-237
Pathogenesis Related Protein Quality
PCR test results for Riesling clone 239 foundation stock record multiplex real-time RT-PCR screening targeting six viruses simultaneously: GFLV (Ct not detected, ND), GLRaV-1 (Ct ND), GLRaV-3 (Ct ND), GVA (Ct ND), GPGV (Ct ND), and GRSPaV (Ct 33.2, positive but below clinical significance threshold) — with internal RNA extraction control confirming adequate nucleic acid quality (Ct 18.5), validating negative results as true negatives rather than extraction failures. PCR test results for a Blaufränkisch candidate clone detected GLRaV-2 by conventional RT-PCR (410 bp amplicon confirmed by gel electrophoresis), subsequently quantified by real-time RT-PCR at Ct 24.3 indicating moderate viral load, triggering referral for heat therapy virus elimination prior to certification eligibility.
PCR test result is a virus testing status entity that records the outcome of polymerase chain reaction-based molecular diagnostic testing performed on nucleic acid extracts from a clonal line, including the specific PCR format employed (conventional RT-PCR, real-time/quantitative RT-PCR, multiplex RT-PCR, or nested RT-PCR), the target virus genome region amplified, the primer sequences or published protocol reference, the amplification result (cycle threshold value for real-time PCR, gel electrophoresis band presence/absence for conventional PCR), and the resulting qualitative or quantitative determination, providing the molecular component of virus status assessment with substantially higher sensitivity and specificity than serological methods.
PCR-based molecular diagnostics have progressively become the reference standard for grapevine virus testing in certification programs, offering 100-1000× greater sensitivity than ELISA and the ability to detect viruses for which no commercial antisera exist. The reverse transcription step (RT) is required because most major grapevine viruses have RNA genomes, necessitating conversion to cDNA before PCR amplification. Real-time (quantitative) RT-PCR provides both detection and viral load quantification through cycle threshold (Ct) values — lower Ct values indicate higher viral titer, with Ct < 25 typically representing high infection levels, Ct 25-30 moderate levels, and Ct 30-35 low-level infections of uncertain biological significance. The interpretation of low-titer positive results (high Ct values) represents a persistent challenge in certification — some authorities apply strict binary positive/negative cut-offs while others consider viral load context, particularly for near-ubiquitous viruses like GRSPaV. Multiplex RT-PCR panels enabling simultaneous detection of 4-8 viruses in a single reaction have substantially improved screening efficiency and reduced per-sample costs. The inclusion of internal controls (housekeeping gene amplification or spiked synthetic RNA) is essential for validating negative results, as inhibitors co-extracted with plant nucleic acids can suppress PCR amplification and generate false negatives. The standardization of PCR protocols across certification laboratories remains an ongoing challenge, as primer selection, thermal cycling conditions, and interpretation criteria vary between institutions despite efforts at harmonization through EPPO diagnostic standards.
PCR result
PCR-Befund
PCR-Testergebnis
RT-PCR result
RT-PCR-Ergebnis
molecular diagnostic result
molecular test result
molekulardiagnostisches Ergebnis
molekulares Testergebnis
nucleic acid test result
Rowhani A et al. (2005) Development of a sensitive colorimetric-PCR assay for detection of viruses in woody plants. Plant Disease 89(12): 1347-1352 Osman F et al. (2007) Real-time RT-PCR (TaqMan) assays for the detection of Grapevine leafroll associated viruses 1-5 and 9. Journal of Virological Methods 141(1): 22-29 Maree HJ et al. (2013) Grapevine leafroll-associated virus 3. Frontiers in Microbiology 4: 82
PCR Test Result
The PDO name Rioja is registered in the EU eAmbrosia database (file number PDO-ES-A0117) as a Protected Designation of Origin for wines produced within the 65,000-hectare delimited zone spanning parts of La Rioja, Álava, and Navarra — the registered name Rioja is protected in its exact form across all EU member states and in third countries through bilateral agreements, with the Consejo Regulador DOCa Rioja authorized to take legal action against any commercial use of the name Rioja for wines not produced in accordance with the registered specification, including evocative uses such as Rioja-style or Rioja-type that are explicitly prohibited under EU Regulation 1308/2013 Art. 103.
A protected designation of origin consisting of the officially registered textual name of the geographic origin as recorded in the relevant PDO registry, constituting the specific designation string that is legally protected against unauthorized commercial use, imitation, evocation, or any practice liable to mislead consumers as to the true origin of the product.
PDO_Name is the specific textual identifier that constitutes the legally protected intellectual property at the core of the PDO system. The registered name may include geographic terms (Champagne, Chianti), compound geographic-product terms (Brunello di Montalcino), or traditional designations with geographic association (Cava, Sekt bestimmter Anbaugebiete): the scope of protection extends beyond identical reproduction to cover imitation, evocation, and any misleading indication of geographic origin — a legal standard that has generated extensive case law including landmark European Court of Justice decisions on the scope of evocation protection (e.g., the Champanillo case C-783/19 on evocative use of Champagne-derived names). The PDO name must be used in the exact registered form on wine labels and commercial documents, with transliteration into non-Latin scripts governed by specific regulatory provisions.
PDO designation string
PDO-Bezeichnungstext
eingetragener g.U.-Name
geschützter Ursprungsname
protected origin name
registered PDO name
EU Regulation 1308/2013 Art. 103 on protection of designations EU eAmbrosia database CJEU Case C-783/19 Champanillo (2021)
PDO Name
The PDO registration number PDO-FR-A0652 uniquely identifies the Châteauneuf-du-Pape designation in the EU eAmbrosia database, with the code structure encoding the protection type (PDO), the member state of origin (FR for France), and a sequential filing number (A0652) — this registration number serves as the persistent identifier linking the designation to its complete regulatory dossier including the product specification, delimitation maps, control body designation, and amendment history, and is referenced in all official communications between the European Commission, national authorities, and the applicant group (Syndicat des Vignerons de Châteauneuf-du-Pape) regarding the protection of this designation.
A protected designation of origin consisting of the unique alphanumeric identifier assigned to a registered PDO by the registering authority at the time of official registration, providing a persistent machine-readable reference code that unambiguously identifies the PDO record within the registration database independently of name variations, translations, or textual ambiguities.
PDO_Registration_Number provides the language-independent persistent identifier for PDO records that resolves the ambiguity inherent in name-based referencing across multilingual regulatory systems. The EU eAmbrosia database assigns structured registration numbers following the pattern [protection type]-[country code]-[sequence number] that enables unambiguous cross-referencing across the approximately 1,600 wine PDOs and PGIs registered in the EU system: this persistent identifier is essential for regulatory data management because PDO names may exist in multiple language versions (e.g., the same PDO may be referenced as Tokaji in Hungarian and Tokajer in German) and name-based searching is complicated by diacritical marks, transliteration variants, and historical name changes. The registration number also serves as the key identifier in international notification procedures under the WTO TRIPS Agreement and bilateral trade agreements.
PDO database code
PDO file number
PDO-Datenbankkennung
PDO-Registriernummer
g.U.-Aktenzeichen
EU eAmbrosia database of registered geographic indications EU Regulation 1308/2013 Art. 96-97 on registration procedures European Commission Delegated Regulation 2019/33 Art. 12-14
PDO Registration Number
The PDO specification reference for Barolo DOCG comprises the Disciplinare di Produzione dei Vini a Denominazione di Origine Controllata e Garantita Barolo (approved by Ministerial Decree of July 1, 1980, most recently amended October 21, 2010) published in the Gazzetta Ufficiale della Repubblica Italiana and registered with the European Commission — this specification document defines the mandatory production parameters including exclusive use of Nebbiolo, maximum yield of 8,000 kg/ha, minimum aging of 38 months (18 in oak) for standard Barolo and 62 months (18 in oak) for Riserva, minimum alcohol of 13%, and geographic delimitation to registered vineyards within the 11 authorized communes of the Langhe.
A protected designation of origin consisting of a bibliographic or documentary reference identifying the official product specification (cahier des charges, disciplinare di produzione, Produktspezifikation) that defines the production rules, quality requirements, geographic delimitation, and control procedures governing the use of the PDO name, providing the traceability link between the registered designation and its binding regulatory content.
PDO_Specification_Reference links the registered PDO name to the binding legal document that defines what the designation means in practice. The specification document is the foundational regulatory text of the PDO system — it translates the general concept of geographic origin protection into specific, enforceable production rules that every producer claiming the PDO must follow: these rules typically cover authorized grape varieties, viticultural practices (planting density, training systems, irrigation restrictions), maximum yields, winemaking methods, aging requirements, analytical parameters (minimum alcohol, maximum volatile acidity), and sensory characteristics. The specification reference must be publicly accessible and is published in both national official journals and the EU DOOR (Database of Origin and Registration) system, enabling any interested party to verify the requirements associated with a given PDO designation.
Pflichtenheft-Verweis
Produktspezifikations-Kennung
Spezifikationsreferenz
cahier des charges reference
product specification identifier
specification document reference
EU Regulation 1308/2013 Art. 94 on product specifications INAO (2023) Cahiers des charges des appellations d'origine Ministero delle Politiche Agricole Alimentari e Forestali (2023) Disciplinari di produzione vini DOP e IGP
PDO Specification Reference
Marker-assisted breeding for PdR1_Mediated_Resistance_Disposition at UC Davis has produced advanced wine grape selections through systematic backcrossing of V. arizonica b43-17 resistance into V. vinifera backgrounds. The breeding strategy involved initial F1 crosses to 'Chardonnay' and 'Cabernet Sauvignon', followed by four generations of backcrossing to elite V. vinifera cultivars with PdR1 selection at each generation using flanking SSR markers. The resulting BC₄ selections contain approximately 97% V. vinifera genetic background while retaining the PdR1 resistance locus. Phenotypic validation through greenhouse needle inoculation with X. fastidiosa strain Temecula1 (10⁸ CFU/mL) demonstrated that PdR1-positive selections maintained bacterial populations of 10²-10⁴ CFU/g petiole at 12 weeks post-inoculation, compared to 10⁷-10⁸ CFU/g in PdR1-negative siblings and susceptible 'Chardonnay' controls. Symptom expression in PdR1-positive plants ranged from none to minimal marginal necrosis, while susceptible controls exhibited severe leaf scorch with 80-100% defoliation. The released cultivar 'Camminare Noir' (PdR1/pdr1 heterozygous, 97% V. vinifera) has demonstrated stable field resistance in Temecula, California trials over 8 years, producing commercially acceptable red wines described as having "Pinot Noir-like characteristics" in sensory evaluations, validating that PdR1_Mediated_Resistance_Disposition can be combined with high wine quality.
A PdR1_Mediated_Resistance_Disposition is a Pierces_Disease_Resistance_Disposition that inheres in a Vitis plant carrying the PdR1 (Pierce's disease Resistance 1) locus derived from Vitis arizonica and is realized through major gene-mediated defense responses that strongly limit Xylella fastidiosa multiplication and systemic spread in xylem tissues. The PdR1 locus maps to chromosome 14 and was identified in the V. arizonica accession b43-17 collected from canyons near Monterrey, Mexico, representing a co-evolved resistance from a region with endemic X. fastidiosa pressure. PdR1 confers dominant resistance characterized by dramatic restriction of bacterial populations, with resistant genotypes maintaining titers 1,000-10,000 fold lower than susceptible V. vinifera under equivalent inoculum pressure. The PdR1 resistance mechanism appears to involve rapid host recognition of X. fastidiosa followed by local defense responses that limit bacterial multiplication and prevent systemic colonization of the vine. Fine mapping has resolved PdR1 to a cluster of putative nucleotide-binding leucine-rich repeat (NLR) genes, suggesting effector-triggered immunity analogous to resistance mechanisms against other pathogens. Marker-assisted selection using tightly linked molecular markers (VVIP26, VVIN64) enables efficient introgression of PdR1 into V. vinifera backgrounds through backcrossing. The UC Davis breeding program has successfully developed PdR1-carrying wine grape cultivars including 'Camminare Noir', 'Paseante Noir', and 'Errante Noir' that combine Pierce's disease resistance with acceptable wine quality, representing a major achievement in grapevine disease resistance breeding.
PdR1_Mediated_Resistance_Disposition is_a Pierces_Disease_Resistance_Disposition. PdR1 from V. arizonica b43-17 represents the most advanced and validated Pierce's disease resistance locus, located on chromosome 14 in an NLR gene cluster. The resistance is dominant, enabling heterozygous deployment in commercial cultivars. Marker-assisted selection has been critical for efficient introgression through multiple backcross generations. Additional PD resistance loci have been mapped in other Vitis species (PdR2, PdR3), providing opportunities for future gene pyramiding to enhance durability. The UC Davis PdR1 breeding program represents a model for grapevine disease resistance deployment.
PdR1 Pierce's disease resistance
PdR1-Pierce-Resistenz
PdR1-vermittelte Resistenz-Disposition
V. arizonica-derived PD resistance
V.-arizonica-Resistenz
major gene Xylella resistance
Krivanek AF et al. (2005) Vitis resistance to Pierce's disease is characterized by differential Xylella fastidiosa populations in stems and leaves, Phytopathology 95:44-52 Riaz S et al. (2008) Genetic and physical mapping of the Pierce's disease resistance locus PdR1, Theoretical and Applied Genetics 117:1067-1078 Walker MA, Tenscher A (2017) Breeding Pierce's disease resistant winegrapes, Acta Horticulturae 1157:191-196
PdR1 Mediated Resistance Disposition
The PdR locus information compiled from UC Davis and USDA breeding programs documents the principal resistance source PdR1 on chromosome 14, derived from Vitis arizonica accession b43-17 collected from a canyon in Monterrey County, Mexico — PdR1 confers strong resistance to X. fastidiosa by restricting bacterial movement through xylem vessels, limiting the systemic colonization that causes the leaf scorch, cane maturation failure, and vine death characteristic of Pierce's disease in susceptible V. vinifera. The breeding program initiated by Andrew Walker at UC Davis has produced advanced backcross selections (BC4) carrying PdR1 with over 97% V. vinifera genome recovery, yielding wines that have passed blind tasting evaluations against varietal V. vinifera standards — the cultivar releases Camminare Noir (2019) and Paseante Noir (2019) represent the first PdR1-carrying winegrape cultivars available for commercial planting in Pierce's disease-endemic regions of California, Texas, and the southeastern United States.
A resistance gene information record specific to PdR (Pierce's Disease Resistance) loci — the genomic regions conferring resistance to the bacterium Xylella fastidiosa, the causal agent of Pierce's disease — documenting the locus designation, chromosomal location, source Vitis species, resistance mechanism (limiting bacterial movement in xylem vessels), and diagnostic molecular markers used in breeding programs developing Pierce's disease-resistant winegrape cultivars for warm-climate viticulture regions where X. fastidiosa and its insect vectors are endemic.
PdR_Locus_Information documents resistance to a fundamentally different pathogen type — a gram-negative xylem-limited bacterium rather than the fungal and oomycete pathogens targeted by Rpv, Run, and Ren loci — reflecting the distinct biological challenge posed by Pierce's disease in warm-climate viticulture. X. fastidiosa is transmitted by xylem-feeding sharpshooter leafhoppers (principally Homalodisca vitripennis, the glassy-winged sharpshooter) and colonizes xylem vessels where it forms biofilms that block water transport, causing progressive vine decline and death within 1-5 years of infection in susceptible V. vinifera: Pierce's disease has historically limited V. vinifera viticulture in the southeastern United States and poses an expanding threat in California, southern Europe (where X. fastidiosa was detected in Italian olive groves in 2013 and has since spread to other Mediterranean crops), and other warm-climate wine regions. The resistance mechanism of PdR1 involves restriction of bacterial movement between xylem vessels through enhanced production of tyloses (xylem parenchyma outgrowths that occlude vessel lumens) and pectin-rich gels that compartmentalize the infection, preventing the systemic colonization required for disease symptom development.
PdR gene characterization
PdR-Locus-Information
Pierce's disease resistance locus data
Pierce-Krankheit-Resistenzlocus-Daten
Xylella resistance gene information
Xylella-Resistenzgen-Daten
Krivanek AF et al. (2006) Resistance to Xylella fastidiosa in North American Vitis species and in V. vinifera-V. arizonica hybrids, Molecular Breeding 18: 155-167 Walker MA and Tenscher A (2019) Breeding Pierce's disease resistant winegrapes, Acta Horticulturae 1248: 239-246 Riaz S et al. (2018) Genetic mapping of PdR2, a Pierce's disease resistance locus derived from Vitis arizonica, Molecular Breeding 38: 27
PdR Locus Information
In a Priorat Garnacha vineyard in Spain, peak acid reduction rate was measured over 11 days in late August Year N+1 (BBCH 85) when titratable acidity declined from 6.8 to 2.9 g/L (average 0.63 g/L per day, 4.4 g/L per week), malic acid decreased from 3.8 to 1.2 g/L (0.46 g/L per day), pH increased from 3.42 to 3.78 (0.033 units per day), berry Brix 24.2° rising to 27.8°, under extreme Priorat heat (daily high 34-38°C, slate soils radiating heat), necessitating immediate harvest despite high sugar to preserve minimal acidity for wine balance.
Peak_Acid_Reduction_Rate represents the maximum velocity of titratable acidity decline achieved during optimal ripening conditions when all components of the acid degradation pathway operate at maximum efficiency creating the fastest compositional transformation observed during berry development determining the steepest trajectory toward final acid balance and critically affecting wine quality potential. This peak rate occurs during mid-to-late veraison (BBCH 84-85), typically 15-22 days post-veraison onset in warm conditions or 18-28 days in moderate conditions, and is characterized by titratable acidity declining 0.45-0.75 g/L per day (3.2-5.2 g/L per week) representing net loss of 5-8% remaining acid daily under warm conditions, malate-specific decline contributing 75-90% of total TA reduction with 0.35-0.60 g/L per day malate loss, pH increasing 0.030-0.055 units per day (0.21-0.38 units per week) creating rapid compositional shift, and remaining berry acidity potentially declining from 7-11 g/L to 3-6 g/L over 8-12 day peak period representing 50-70% loss of this phase's entry values. Biochemical mechanisms enabling peak rate include malic enzyme activities at absolute maximum 50-85 units/g fresh weight combined NADP-ME and NAD-ME providing saturating catalytic capacity with Vmax 40-75 μmol malate oxidized g⁻¹ h⁻¹ in vitro translating to 3-6 μmol g⁻¹ h⁻¹ in vivo accounting for compartmentation and regulation, citric acid cycle flux maximized with malate dehydrogenase, citrate synthase, and isocitrate dehydrogenase all operating at elevated rates 25-50 units/g each ensuring complete oxidation pathway capacity, respiratory chain activity enhanced with O₂ consumption 2.2-3.8 μmol g⁻¹ h⁻¹ oxidizing NADH and FADH₂ from malate catabolism, and cellular energetics optimized with ATP production from acid oxidation reaching 55-95 μmol g⁻¹ h⁻¹ providing 40-70% of berry energy requirements during peak supporting intensive ripening metabolism. Substrate dynamics at peak rate show vacuolar malate 4-8 g/L (40-80 mM) declining rapidly still above kinetic limitation threshold, tonoplast transport maximized 3-5 μmol malate g⁻¹ h⁻¹ efflux maintaining cytoplasmic supply, cytoplasmic malate 10-20 mM sustained above enzyme Km 0.5-2.5 mM ensuring near-Vmax kinetics, and pH gradients vacuole 2.8-3.2 versus cytoplasm 7.0-7.4 driving facilitated diffusion potentially supplemented by secondary active transport maintaining substrate flow to catabolism. Metabolic integration at peak rate demonstrates respiratory malate contribution 55-75% of total CO₂ production indicating acid catabolism dominates energy metabolism temporarily exceeding hexose oxidation contribution, pyruvate production from malic enzyme 3.0-5.5 μmol g⁻¹ h⁻¹ entering mitochondria for complete oxidation or cytoplasmic metabolism, NADPH generation 3.0-5.5 μmol g⁻¹ h⁻¹ from NADP-malic enzyme supporting reductive biosynthesis (anthocyanin, flavonoid, terpenoid synthesis all active) or oxidized for additional ATP, and carbon flow redirected substantially from storage functions toward respiratory energy production reflecting metabolic priorities during intensive ripening. Environmental factors producing peak rate include temperature optimum 28-32°C enabling maximum enzyme activities and metabolic flux producing highest rates 0.55-0.75 g/L per day, extreme warmth 32-36°C potentially exceeding optimal causing 0.60-0.85 g/L per day loss creating risk of excessive deacidification, moderate warmth 24-28°C supporting near-peak rates 0.40-0.60 g/L per day achieving appropriate acid reduction, cooler conditions 20-24°C limiting to 0.28-0.45 g/L per day through kinetic constraints preserving acids, and heat accumulation during peak window with cumulative degree-days >30°C strongly predicting total acid loss r² = 0.84-0.95 enabling forecasting and management. Diurnal variation at peak rate shows midday maximum 0.70-1.10 g/L per day during hottest hours 1-5 PM when berry temperature peaks 32-40°C from solar radiation, early morning and evening rates 0.35-0.55 g/L per day as berry temperature moderate 24-28°C, nighttime minimum 0.18-0.38 g/L per day with berry cooling to 16-24°C substantially reducing metabolic rates, and daily integrated rate averaging 0.45-0.70 g/L representing time-weighted mean of diurnal extremes with warm-climate regions showing higher averages through extended warm periods while temperate regions show moderation through cool nights. Cluster exposure effects particularly dramatic at peak rate with fully sun-exposed clusters experiencing berry temperatures 6-14°C above shaded fruit creating 60-120% faster acid reduction, southwestern exposure (northern hemisphere) showing maximum rates through afternoon solar heating combined with diurnal heat accumulation, northeastern exposure demonstrating moderated rates through limited direct sun and cooler microclimate, and interior canopy positions maintaining 35-60% slower rates than exterior enabling acid preservation through natural shading. Water relations influences at peak rate demonstrate adequate vine water status supporting maximum metabolic capacity enabling peak rates without limitation, moderate deficit (stem ψ -1.0 to -1.4 MPa) showing variable effects with some cultivars displaying 10-25% rate enhancement through stress while others unaffected, severe deficit (3.7 problematic for wine stability and structure, inadequate peak in cool regions maintaining TA >8-10 g/L requiring extended ripening or winery intervention, timing relative to sugar accumulation affecting harvest decisions with rapid acid loss potentially necessitating earlier harvest at lower Brix to preserve balance, and spatial variation in peak rates across vineyard creating maturity heterogeneity requiring selective harvest or acceptance of compositional variation in must. Wine style implications include final pH critically affecting character with whites optimal 3.0-3.4, reds 3.3-3.7, values outside these ranges producing suboptimal wines, malolactic fermentation kinetics affected by residual malate with >3-4 g/L supporting rapid complete MLF while <2 g/L potentially causing sluggish or stuck fermentation, aging potential influenced by pH with lower values (3.1-3.4) enhancing stability and longevity through reduced oxidation and better color retention, and regional adaptation with warm regions requiring management to prevent excessive peak while cool regions may benefit from or require extended peak period achieving adequate deacidification. Harvest timing during peak rate requires frequent monitoring every 2-3 days as compositional changes rapid, integrated assessment combining acid monitoring with sugar, phenolic, and flavor evaluation ensuring holistic maturity, forecasting final composition based on current rate and remaining expected growing degree-days enabling proactive harvest scheduling, and flexibility to harvest promptly when optimal parameters achieved as 3-5 day delay during peak can produce substantially altered composition potentially compromising quality targets. Viticultural interventions during peak rate include canopy shading strategically applied reducing berry temperature 4-9°C slowing rate 30-60% preserving acids critical in warm regions, irrigation potentially moderating through cooling effects though timing and amount requiring careful management avoiding negative vigor or dilution consequences, crop thinning pre-peak potentially beneficial concentrating resources avoiding stress-accelerated loss though post-peak intervention ineffective, and disease protection essential as changing composition increases vulnerability requiring preventive fungicide and sanitation. Monitoring and forecasting strategies include frequent laboratory analysis tracking TA, pH, malate, and tartrate every 2-4 days during peak enabling trajectory visualization, portable meters (pH, TA) for rapid field assessment guiding detailed sampling, spatial sampling across vineyard zones revealing heterogeneity requiring zone-specific management, degree-day accumulation models predicting expected loss based on temperature forecast enabling proactive decisions, and sensory evaluation ensuring acid-driven tartness declining appropriately coordinating with developing sweetness and flavor ripeness.
Peak_Acid_Reduction_Rate is distinguished from Active_Acid_Degradation by representing maximum velocity (0.45-0.75 g/L per day) rather than overall active phase, from Initial_Acid_Reduction by showing peak rather than early rates, and from Rapid_Acidity_Decline by focusing on rate measurement rather than compositional trajectory. Duration 8-14 days BBCH 84-85. Temperature optimum 28-32°C, >32°C excessive. Malate contributes 75-90% of TA reduction. Critical determinant of final acid balance. Peak_Acid_Reduction_Rate represents maximum velocity of compositional transformation affecting wine quality.
Spitzen-titrierbare-Azidität-Reduktion
fastest TA decline rate
maximale Entsäuerungs-Geschwindigkeit
maximale Säure-Verlust-Rate
maximum acid loss rate
maximum deacidification velocity
peak titratable acidity reduction
schnellste TA-Abnahme-Rate
Sweetman C, Sadras VO, Hancock RD, Soole KL, Ford CM (2014) Metabolic effects of elevated temperature on organic acid degradation in ripening Vitis vinifera fruit J Exp Bot 65:5975-5988 peak rate quantification Lakso AN, Kliewer WM (1975) The influence of temperature on malic acid metabolism in grape berries Plant Physiol 56:370-372 maximum rates Coombe BG (1992) Research on development and ripening of the grape berry Amer J Enol Vitic 43:101-110 ripening dynamics including peak acid decline.
Peak Acid Reduction Rate
Cabernet Sauvignon in Napa Valley shows peak anthesis during mid-June Year N+1 at BBCH 63-65 over 3-4 days June 8-11, with 55-70% flowers open simultaneously, abundant pollen availability from widespread anther dehiscence, maximum stigma receptivity across cluster, under optimal warm climate conditions of 24-29°C temperature, 50-65% relative humidity, light winds, no rain, supporting efficient pollination and achieving 58-68% fruit set typical of this late-season cultivar under favorable conditions
Peak anthesis represents the maximum flowering intensity phase when greatest proportion of flowers open simultaneously creating optimal conditions for pollination through abundant pollen availability and widespread stigma receptivity, establishing critical reproductive window determining majority of fruit set and yield. This essential reproductive culmination occurs during early-mid June in temperate regions Year N+1 typically 2-4 days after early anthesis initiation, spanning 2-4 days when 40-70% flowers achieve open state simultaneously, corresponding to BBCH 63-65 stages. Flowering intensity during peak shows characteristic maximum with 40-70% flowers open at any moment creating dense arrays of functional reproductive structures, daily addition of 10-20% new flowers opening maintaining high open flower proportion, and sustained intense reproductive activity across cluster establishing optimal pollination window. Flower distribution pattern at peak shows opened flowers throughout cluster with apical regions showing 70-90% open having initiated opening earliest, middle regions showing 60-80% open representing peak activity zone, and basal regions showing 30-60% open with opening continuing progressively, creating overall highly functional state. Reproductive organ functionality reaches maximum during peak with abundant viable pollen available from numerous anthers undergoing dehiscence, widespread stigma receptivity with 50-80% stigmas showing active exudate production and pollen capture capacity, and optimal spatial arrangement with opened flowers in close proximity facilitating efficient pollination. Pollen availability shows peak levels with anthers from recently opened flowers releasing fresh viable pollen, cumulative pollen from flowers opened over previous 1-2 days remaining partially viable, and continuous new pollen production from flowers opening during peak period, creating maximum pollen density in cluster atmosphere. Pollen viability during peak remains high with fresh pollen showing 70-90% germination capacity under optimal conditions, declining to 40-60% after 24 hours and 10-30% after 48 hours, creating 1-2 day window of maximum viability requiring effective pollination during this brief period. Pollen dispersal during peak creates visible pollen clouds surrounding flowering clusters under favorable dry windy conditions, with pollen grains becoming airborne readily when anthers dehisce, traveling centimeters to meters through wind transport, and settling on receptive stigmas within same cluster or neighboring clusters. Stigma receptivity distribution during peak shows maximum proportion of stigmas at optimal receptive state with fresh exudate production, providing widespread pollen capture capacity across cluster, with 60-80% stigmas showing peak receptivity creating abundant opportunities for successful pollination. Pollination activity intensifies during peak with high frequency of pollen-stigma contact events, rapid pollen germination on receptive stigmas typically within 1-4 hours, and extensive pollen tube growth initiating through style tissues toward ovaries, establishing conditions for high fruit set. Fertilization initiation begins during peak as earliest pollinated flowers complete pollen tube growth reaching ovaries typically 18-36 hours after pollination, with double fertilization occurring and embryo plus endosperm development beginning, triggering fruit set through hormonal signals preventing flower abscission. Cluster appearance at peak anthesis shows characteristic flowering appearance with numerous opened flowers showing yellow-brown anthers contrasting with green receptacles, remaining calyptra caps on unopened buds showing green color, detached caps accumulating on cluster surface and ground below, and overall cluster showing active reproductive state. Environmental conditions during peak critically determine flowering success with temperature optimally 22-30°C supporting maximum pollen and stigma function, cool conditions below 18°C substantially reducing pollen viability and stigma receptivity, and excessive heat above 33°C potentially causing pollen sterility or stigma dysfunction severely limiting fruit set. Humidity during peak preferably 45-65% relative humidity balancing pollen viability maintenance with anther dehiscence facilitation, high humidity above 75% preventing or delaying anther opening reducing pollen availability, and very low humidity below 35% causing rapid pollen desiccation reducing viability window. Weather stability during peak is critically important with favorable stable conditions enabling efficient pollination and high fruit set, while variable conditions create erratic pollination patterns and reduced set, with peak anthesis representing most weather-sensitive period of entire growing season. Rain during peak anthesis represents catastrophic risk with water preventing anther dehiscence eliminating pollen availability, washing pollen from anthers and stigmas preventing pollination, diluting or removing stigma exudate impairing pollen capture and germination, and promoting botrytis infection on wet flowers causing immediate tissue damage and establishing latent infections, with rain during peak potentially reducing fruit set to 10-30% compared to 50-70% under favorable conditions. Wind during peak shows beneficial effects with light-moderate wind 5-15 km per hour facilitating pollen dispersal and enhancing pollination efficiency, though strong wind above 25 km per hour may cause flower damage or excessive pollen loss reducing pollination success. Metabolic activity during peak shows maximum levels with elevated respiration in anthers supporting pollen maturation and release, active metabolism in stigmas maintaining receptivity and supporting pollen tube growth, and high energy demand across reproductive structures requiring adequate carbohydrate supply. Resource demand during peak emphasizes carbohydrate requirements for energy-intensive processes including pollen production, tube growth, and early embryo development, nitrogen for protein synthesis in reproductive tissues, and critical micronutrients particularly boron essential for pollen germination and tube growth. Hormonal status during peak involves active auxin and gibberellin production from successfully fertilized ovaries triggering berry development, ethylene possibly involved in coordinating flower senescence for unfertilized flowers, and other hormones coordinating transition from flowering to fruit development. Developmental coordination during peak shows concurrent vegetative and reproductive processes with shoot growth continuing though slowing, leaves expanding and maturing providing photosynthetic capacity, and tendrils developing and attaching supporting shoot positioning. Canopy management during peak should avoid disruptive operations with shoot positioning, hedging, or leafing operations suspended during flowering period preventing physical disturbance of delicate flowers, and disease management restricted due to fungicide application limitations during bloom. Disease pressure during peak focuses on botrytis with flowers highly susceptible to Botrytis cinerea infection particularly under wet conditions, establishing latent infections that remain dormant through berry development expressing during ripening, requiring protective fungicide applications before flowering due to application restrictions during bloom. Fruit set determination during peak with successful pollination and fertilization during this critical 2-4 day period determining 60-80% final berry number, with remaining 20-40% set during early and late anthesis periods, making peak anthesis outcomes dominant influence on seasonal yield. The process shows three distinguishable aspects progressing from maximum flower opening establishing peak intensity, through peak pollen availability providing maximum pollination opportunity, to optimal pollination window encompassing ideal conditions for reproductive success. Duration typically spans 2-4 days representing brief intense period of maximum flowering, with warm favorable conditions producing compressed 2-3 day peak while variable conditions may extend peak to 4-5 days. Cultivar variation appears in peak intensity with some varieties showing sharp intense peak with 60-80% simultaneous opening while others show broader moderate peak with 40-60% opening, peak duration affecting pollination window and fruit set patterns, and environmental sensitivity affecting peak success reliability. Regional timing shows peak anthesis during early-mid June in warm climates at approximately 1000-1100 growing degree days base 10°C, mid-June in temperate regions at 900-1000 GDD, and mid-late June in cool areas at 800-900 GDD, with timing critical for favorable weather window. Viticultural significance includes representing most critical period for yield determination with peak outcomes dominating fruit set, requiring optimal environmental conditions for successful pollination and high fruit set, representing maximum weather vulnerability requiring monitoring and possible protective interventions, marking transition from structural development to reproductive function, and providing definitive reproductive outcome window affecting season productivity. Research applications include analyzing environmental controls of peak flowering intensity and duration, quantifying pollination dynamics including pollen production, dispersal, and capture, evaluating relationships between peak conditions and fruit set outcomes, developing models predicting fruit set from peak weather conditions, and assessing climate change impacts on peak anthesis conditions affecting viticultural productivity.
Peak_Anthesis is second and culminating major Anthesis phase, distinguished by maximum flowering intensity and optimal reproductive conditions. Follows Early_Anthesis. Corresponds to BBCH 63-65 with 40-70% flowers open. Subdivided into Maximum_Flower_Opening, Peak_Pollen_Availability, and Optimal_Pollination_Window. Represents most critical period for yield determination. Environmental conditions during peak fundamentally determine fruit set and season productivity. Weather sensitivity at absolute maximum.
Blütenhöhepunkt
Hauptblüte
flowering peak
full bloom
maximale Anthese
maximum anthesis
peak flowering
volle Blüte
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 370-380 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 285-292 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Peak Anthesis
Cabernet Sauvignon in Napa Valley shows peak berry growth rate during late July-early August Year N+1 at BBCH 75-77, with maximum growth rate 160-220 mg fresh weight per day sustained 3 weeks, water uptake 1.8-2.4 g per berry per day, berry diameter expanding from 22 to 32 mm, fresh weight increasing from 8 to 18 g achieving 82% final size, coordinated cell expansion across 2.8 million cells, under warm climate conditions of 26-32°C temperature with adequate irrigation, typical of this late-season large-berried cultivar
Peak berry growth rate represents the maximum berry enlargement phase during rapid cell expansion when coordinated individual cell expansions create highest berry volumetric increase, typical growth rates 100-250 mg fresh weight per day in large-berried cultivars, sustained high rate over 2-4 weeks produces substantial size gain, and berry achieves 75-90% final size establishing near-final dimensions, occurring during intensive enlargement weeks 5-7 post-fertilization. This maximum berry growth phase occurs during late summer in late July-early August in temperate regions Year N+1, spanning 2-4 weeks representing peak growth intensity, corresponding to BBCH 75-77 berry touch to beginning bunch closure stages. Peak rate mechanism involves coordinated expansion of all berry cells creating additive volumetric increase, maximum individual cell expansion rates translating to berry growth, intensive water uptake 1.5-2.5 g per berry per day supplying growth, coordinated tissue-level growth maintaining berry integrity, and peak rate reflecting optimal growth conditions and developmental state. Berry growth rate quantification shows fresh weight increase 100-250 mg per day during peak in large-berried cultivars, 50-120 mg per day in small-to-medium cultivars, sustained high rate over 2-4 weeks, and rate as convenient growth measure. Growth rate variation shows cultivar differences with large-berried showing highest absolute rates, small-berried showing lower rates but similar relative growth, and genetic variation determining peak rate capacity. Berry volumetric increase shows diameter expansion 1-2 mm per week during peak period, volumetric increase 1.5-2.5 fold over 2-4 week peak phase, substantial size gain during intensive growth, and volumetric increase as major berry growth. Berry diameter progression shows typical expansion from 18-25 mm at peak onset to 24-36 mm by completion depending on cultivar, diameter increase 6-12 mm during peak period, and substantial linear growth. Berry fresh weight accumulation shows weight increase from 6-12 g at peak onset to 10-22 g by completion in large-berried cultivars, weight gain 4-10 g during peak period, and major mass accumulation. Berry size achievement shows berry reaching 75-90% final mature size by peak rate period completion, substantial approach to genetic size potential, remaining growth during lag and ripening relatively limited 10-25%, and peak period as major size determinant. Cell expansion coordination shows all established cells expanding synchronously or near-synchronously, tissue-level coordination creating uniform berry growth, coordinated expansion maintaining structural integrity and shape, and population-level coordination as growth mechanism. Cell contribution to berry growth shows individual cell 10-30 fold volume increase from division completion to mature size, final cell volume 80-200 micrometers diameter depending on cultivar, and multiplicative effect of cell number and cell size determining berry size. Cell number-cell size interaction shows berry volume determined by cell number × average cell volume, genetic variation in both parameters, and interactive effects creating cultivar size ranges. Water uptake during peak growth shows intensive water import 1.5-2.5 g per berry per day, water as primary mass contributor accounting for 80-85% fresh weight gain, xylem as major water source, and high water flux supporting rapid growth. Xylem function during peak growth shows active xylem delivery supporting water demand, xylem-to-berry water transfer through vascular connections, xylem dominance during growth phase, and vascular capacity as potential growth limitation. Berry composition during peak growth shows water content maintained 80-85% fresh weight, organic acids at peak 1.2-1.8%, sugars still moderate 5-8%, pH 2.8-3.0, and green acidic berry characteristic. Dry matter accumulation shows progressive dry matter increase during peak growth, dry matter content relatively stable 12-18%, absolute dry matter gain from organic acids, structural components, and minor sugar, and dry matter as structural and metabolic component. Organic acid accumulation shows acids reaching peak concentrations during peak growth, malate and tartrate dominating at 1.2-1.8% total, acids contributing to osmotic potential and acidity, and peak acid content characteristic of late growth phase. Berry morphology during peak growth shows berry expanding to approach final shape, spherical to ovoid morphology depending on cultivar, smooth surface maintained, and berry appearance as expanding green fruit. Cluster architecture during peak growth shows berries expanding to contact neighbors, cluster compactness increasing, berries filling available cluster space, and cluster filling characteristic of peak growth. Environmental effects on peak berry growth include water availability as most critical factor with adequate water absolutely essential for maximum growth rate, water deficit severely limiting peak growth reducing final size substantially, temperature affecting growth rate with optimal 24-28°C supporting maximum rates, and favorable conditions essential for peak rate achievement. Water deficit effects show deficit during peak growth as most critical period for size limitation, reduced growth rate proportional to deficit severity, deficit effects on final size substantial 20-50% size reduction, and water as primary limiting factor during peak growth. Temperature effects on peak growth show warm conditions supporting maximum growth rate, cool conditions reducing rate 20-40% though substantial growth still possible, excessive heat above 35°C potentially limiting growth through stress, and temperature as secondary factor after water. Irrigation importance during peak growth shows irrigation critical in water-limited regions during this period, supplemental water enabling maximum growth rate, and irrigation timing during peak growth as most beneficial for size optimization. Peak growth rate duration shows typical sustained high rate 2-4 weeks, duration variation among cultivars affecting total size gain, and sustained rate characteristic of favorable conditions. Growth rate decline shows eventual rate reduction as berry approaches genetic size limit, rate decline marking transition to lag phase, and decline as developmental progression. The process represents third and final aspect of rapid cell expansion emphasizing berry-level growth, following maximum expansion rate and massive vacuole formation. Duration typically 2-4 weeks representing peak berry growth period weeks 5-7. Regional timing shows peak berry growth rate during late July-early August across climates during intensive enlargement. Viticultural significance includes representing maximum berry size increase determining substantial yield, requiring optimal water availability for maximum growth, demonstrating critical environmental sensitivity affecting final size, and establishing near-final berry dimensions affecting yield and quality. Research applications include analyzing berry growth dynamics and regulation, investigating water relations and supply during peak growth, evaluating environmental effects on growth rate, quantifying growth determinants and limitations, and assessing peak growth as major berry size determinant.
Peak_Berry_Growth_Rate is third and final aspect of Rapid_Cell_Expansion, distinguished by maximum berry-level enlargement. Represents coordinated cell expansion creating 100-250 mg/day growth in large cultivars. Water uptake 1.5-2.5 g/berry/day. Berry achieves 75-90% final size. Diameter increases 6-12 mm. Duration 2-4 weeks. Water availability most critical factor - deficit reduces size 20-50%. Temperature optimal 24-28°C. Follows Maximum_Expansion_Rate and Massive_Vacuole_Formation. Completes Rapid_Cell_Expansion and Berry_Cell_Enlargement_Phase. Major yield determinant.
Spitzen-Beerenwachstumsrate
Spitzen-Vergrößerungsrate
highest berry expansion
höchste Beerenexpansion
intensive berry growth
maximales Beerenwachstum
maximum berry growth
peak enlargement rate
Peak Berry Growth Rate
Cabernet Sauvignon in Napa Valley shows peak cap shedding period on June 9-10 Year N+1 at BBCH 65, with 60-72% flowers showing detached caps, intensive visible cap fall during morning hours creating abundant accumulation, under optimal warm climate conditions of 25-29°C temperature, 52-64% relative humidity, stable weather, typical of this late-season cultivar reaching flowering peak approximately 5-6 days after first cap detachment and determining 45-55% final berry number during this critical 1-2 day window
Peak cap shedding period represents the maximum calyptra detachment intensity phase when greatest absolute number and proportion of flowers complete abscission simultaneously creating characteristic full bloom appearance with intensive cap fall activity visible throughout cluster and vineyard. This culminating intensity phase occurs during peak flowering in mid-June in temperate regions Year N+1, typically representing 1-2 day period when 50-75% flowers show detached caps with maximum daily shedding rate, corresponding to BBCH 65 defining full bloom stage. Shedding intensity characteristics show maximum daily detachment rate with 15-25% additional flowers shedding caps per day during peak, cumulative effect of sustained shedding over previous days creating high proportion of simultaneously shed caps, and peak representing balance point between continuing shedding from newly maturing flowers and declining activity as most flowers have completed detachment. Cap accumulation at peak shows maximum density with thick layer of fresh detached caps covering cluster surfaces creating distinctive appearance, continuous cap fall visible throughout observation period under favorable conditions, and substantial accumulation on ground below vines creating visible carpet of fallen caps documenting flowering intensity. Visual appearance at peak shows definitive full bloom display with dense array of opened flowers exposing yellow-brown anthers and green-white stigmas throughout cluster, contrasting with 25-40% remaining green calyptra caps on unopened buds, abundant visible cap fall activity during favorable morning hours, and overall appearance of maximum reproductive activity. Flower exposure distribution at peak shows opened flowers throughout cluster creating maximum reproductive surface area with apical regions showing 80-95% flowers with shed caps from earliest opening, middle regions showing 70-85% detachment representing highest density of recently opened flowers, and basal regions showing 50-70% shedding as activity extends to latest-maturing positions. Synchronized shedding intensity reaches maximum at peak with numerous flowers throughout cluster completing abscission within same time period creating visible simultaneous cap fall, coordinated detachment suggesting possible developmental or environmental synchronization mechanisms, and concentrated activity creating optimal reproductive conditions. Environmental conditions at peak typically favorable with temperature 24-30°C supporting maximum abscission and reproductive activity, humidity 50-65% facilitating efficient detachment, and stable warm dry weather enabling sustained intensive shedding throughout 1-2 day peak period. Diurnal pattern at peak shows most intense shedding during morning hours 8am-11am as temperature rises above threshold and humidity declines to favorable levels, creating daily maximum activity period visible as intensive cap fall, and repeated daily peaks throughout peak period maintaining high overall shedding intensity. Cap fall visibility at peak enables easy observation with abundant falling caps creating obvious visual display, sound of caps hitting cluster surfaces or ground occasionally audible during intensive shedding, and overall sensory evidence of flowering activity at maximum. Pollen cloud formation during peak creates characteristic visual phenomenon with pollen released from numerous anthers creating visible haze surrounding flowering clusters under dry windy conditions, particularly evident during morning hours coinciding with peak cap shedding, and providing additional visual confirmation of maximum flowering activity. Pollination opportunity at peak reaches absolute optimum with maximum flower exposure providing widespread reproductive organ availability, peak pollen production from numerous dehiscing anthers, and optimal spatial conditions for efficient pollination supporting maximum fruit set potential. Fruit set determination at peak shows that outcomes during this critical 1-2 day period dominate final berry number with 40-60% final fruit set established during peak shedding representing largest single contribution, making peak period conditions most important determinant of seasonal yield. Weather vulnerability at peak reaches absolute maximum with reproductive success extremely sensitive to conditions, favorable stable conditions supporting high fruit set 60-75% in good-setting cultivars, while adverse weather particularly rain causing catastrophic fruit set failure 10-30%, demonstrating extreme importance of favorable conditions during brief peak window. Observational assessment at peak enables confident full bloom determination with unmistakable visual evidence of maximum flowering including dense cap accumulation, numerous opened flowers throughout clusters, visible ongoing shedding activity, and overall appearance meeting BBCH 65 full bloom definition. Phenological documentation at peak includes recording peak cap shedding date as critical marker defining BBCH 65 full bloom stage, providing precise reference for phenological records and subsequent stage predictions, and serving as definitive flowering peak marker for management planning and climate studies. Management implications at peak include absolute restriction on vineyard operations with any canopy disturbance potentially disrupting critical pollination processes, weather monitoring at maximum intensity with short-term forecasts critically important, and recognition that peak period represents single most important 1-2 day window of entire growing season for yield determination. Cultivar identification assistance from peak characteristics may include distinctive cap accumulation patterns, shedding intensity and duration, and visual appearance characteristics varying among varieties providing observational clues. The process represents first aspect of mass cap fall focusing on maximum intensity, followed by synchronous cap fall emphasizing coordination. Duration represents brief 1-2 day maximum intensity period, with warm stable conditions producing sharp intense peak while variable conditions may broaden or reduce peak intensity. Cultivar variation in peak intensity shows some varieties reaching very high 65-80% simultaneous shedding creating intense sharp peak, while others show moderate 50-65% creating broader flatter peak, affecting flowering coordination and potential fruit set uniformity. Regional timing shows peak cap shedding period during early-mid June in warm climates at approximately 1050-1100 growing degree days base 10°C, mid-June in temperate regions at 950-1000 GDD, and mid-late June in cool areas at 850-900 GDD, defining flowering peak across regions. Viticultural significance includes defining definitive flowering peak establishing BBCH 65 full bloom, representing single most critical day or two determining majority of season yield, providing unmistakable visual marker for flowering peak easily assessed without detailed examination, and illustrating extreme vulnerability requiring optimal conditions during brief window. Research applications include documenting precise flowering peak timing for phenology studies, analyzing environmental effects on peak intensity affecting reproductive outcomes, quantifying relationships between peak characteristics and fruit set success, developing practical observation protocols using peak cap shedding as marker, and assessing climate variability impacts on flowering peak conditions affecting productivity across seasons.
Peak_Cap_Shedding_Period is first aspect of Mass_Cap_Fall, distinguished by maximum calyptra detachment intensity. Corresponds to BBCH 65 full bloom defining characteristic. Represents culminating 1-2 day period of flowering activity. Provides unmistakable visual marker for flowering peak. Most critical period for yield determination. Extreme weather vulnerability requiring optimal conditions. Easily observable practical reference for phenological tracking.
Hauptkappenfall-Stadium
Spitzen-Kalyptralösung
Vollblüteperiode
full bloom period
maximale Abwurfphase
maximum cap detachment phase
peak calyptra fall
Peak Cap Shedding Period
Cabernet Sauvignon in Napa Valley shows peak division activity during mid July Year N+1 at BBCH 72-73, with maximum mitotic index 10-14% during weeks 2-3 post-fertilization, cell cycle times minimized to 10-12 hours, rapid cell accumulation from 350,000 to 2.4-3.0 million cells during peak period, coordinated division across mesocarp and exocarp tissues, under warm climate conditions of 26-32°C temperature, typical of this late-season large-berried cultivar achieving maximum cell number generation determining final berry size potential
Peak division activity represents the maximum mitotic proliferation phase during berry cell division when division rate reaches highest intensity, majority of mesocarp cells actively dividing, cell cycle times minimize to 10-12 hours, and maximum cell number generation occurs establishing final berry cellularity, typically spanning weeks 2-3 post-fertilization as proliferative capacity peaks before gradual decline. This maximum proliferation phase occurs during mid summer in early-mid July in temperate regions Year N+1, proceeding over weeks 1.5-3.5 post-fertilization with peak intensity weeks 2-3, corresponding to BBCH 72-73 young fruit active growth stages. Peak activity mechanism involves widespread cell cycle synchronization across mesocarp populations, maximum cyclin-CDK activity driving rapid cycle progression, minimal cell cycle times 10-12 hours representing fastest division rate, coordinated division across pericarp tissues, maximum cell number generation determining majority of final cellularity, and peak activity representing culmination of proliferative program before gradual decline. Division rate maximization shows peak mitotic index 8-15% indicating high proportion of dividing cells, maximum absolute division rate with thousands of cells dividing per hour per berry, cell cycle time minimization to 10-12 hours from earlier 14-16 hours, and peak rate representing optimal proliferative capacity under favorable conditions. Cell cycle time characteristics show S phase duration 5-7 hours for DNA replication, G2 phase 1-2 hours for mitotic preparation, M phase 0.5-1 hour for chromosome segregation and cytokinesis, G1 phase 3-4 hours for growth and preparation, and total cycle 10-12 hours during peak activity. Synchronized division shows cell populations entering and exiting cycle phases coordinately, temporal clustering of divisions creating high activity periods, synchronization possibly from common auxin stimulus and division initiation timing, and coordinated divisions creating efficient proliferation. Maximum cell number generation shows majority 60-75% of final cell number accumulated during peak activity weeks 2-3, rapid cell accumulation from 200,000-500,000 to 1-3 million cells during peak period, exponential growth pattern with doubling time 10-12 hours matching cycle time, and peak generation establishing bulk berry cellularity. Pericarp cell proliferation shows mesocarp flesh tissue undergoing most intensive division, cell number increase throughout mesocarp layers, coordinated proliferation maintaining tissue proportions, and mesocarp as primary proliferative compartment determining berry size. Seed compartment cell division shows endocarp inner layer adjacent to seeds showing continued division, specialized cell populations near seeds potentially responding to local auxin maxima, and seed region divisions contributing to overall cellularity though representing smaller fraction. Tissue-specific division patterns show mesocarp accounting for 80-85% total cell divisions, exocarp surface layers 10-15%, endocarp seed regions 3-5%, and tissue distribution reflecting developmental roles and final architecture. Spatial division distribution shows divisions throughout berry volume, possible gradient with higher activity near seeds in some studies, overall relatively uniform distribution across mesocarp, and coordinated division maintaining spherical to ovoid berry shape. Cell population heterogeneity shows variation in cycle timing across cells, some populations dividing faster than others, heterogeneity preventing perfect synchronization, and population variation contributing to extended peak period. Division plane patterns show periclinal divisions predominating in inner regions, anticlinal divisions in outer layers, coordinated orientation creating organized architecture, and plane regulation maintaining tissue structure during rapid proliferation. Auxin effects on peak activity show continued hormone stimulus from developing seeds supporting sustained division, auxin levels typically peaking weeks 1-3 coinciding with division peak, hormone decline beginning week 3-4 correlating with division rate decline, and auxin as primary proliferation driver. Environmental effects on peak activity include temperature critically affecting peak division rate with optimal 24-28°C supporting maximum 10-12 hour cycles, cool conditions 18-22°C extending cycles to 13-16 hours reducing peak intensity, warm conditions 28-32°C potentially supporting faster cycles though excessive heat above 35°C impairing division, and temperature as major peak activity determinant. Nutrient effects show adequate nitrogen supporting protein synthesis for rapid division, phosphorus for DNA synthesis and energy metabolism, and overall nutrition supporting peak activity though usually non-limiting. Peak activity duration shows typically 1.5-2.5 weeks of maximum intensity, peak representing culmination of accelerating activity from week 1, and gradual decline beginning week 3-4 as cells progressively exit cycle. Division decline mechanisms show progressive cell cycle exit beginning week 3, auxin levels declining reducing proliferative stimulus, cells transitioning to expansion phase, and declining division rate characteristic of developmental program. Cell number at peak completion shows typical 70-85% final cell number achieved by peak activity completion week 3-4, remaining 15-30% cells from continued lower-rate division weeks 4-5, and peak activity as primary cellularity determinant. The process shows three distinguishable aspects including maximum cell number generation representing peak proliferative output, pericarp cell proliferation emphasizing mesocarp tissue focus, and seed compartment cell division representing regional specialization. Duration typically 1.5-2.5 weeks representing maximum intensity period. Regional timing shows peak division activity during early-mid July across climates weeks 2-3 post-fertilization. Viticultural significance includes representing maximum proliferation determining majority of final cell number, demonstrating critical importance of optimal conditions during peak weeks for cellularity achievement, showing temperature sensitivity of peak division intensity, and establishing peak activity as major berry size determinant. Research applications include analyzing peak division regulation and timing, investigating cell cycle control during maximum activity, evaluating environmental effects on peak intensity and duration, quantifying cell number generation dynamics, and assessing peak activity as determinant of final berry cellularity and size.
Peak_Division_Activity is second phase of Berry_Cell_Division_Phase, distinguished by maximum mitotic intensity. Subdivided into Maximum_Cell_Number_Generation representing peak proliferative output, Pericarp_Cell_Proliferation emphasizing mesocarp focus, and Seed_Compartment_Cell_Division showing regional specialization. Duration 1.5-2.5 weeks with peak weeks 2-3. Cell cycle times 10-12 hours minimum. Generates 60-75% final cell number. Temperature critically affects intensity. Follows Active_Berry_Cell_Division. Precedes division decline and expansion phase. Major berry size determinant.
Spitzen-Proliferation
Spitzen-Teilungsaktivität
highest division rate
höchste Teilungsrate
maximale mitotische Aktivität
maximum cell division
maximum mitotic activity
peak proliferation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 435-438 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Peak Division Activity
Merlot peak division stage berries at weeks 2.5-3.5 post-fruit set (BBCH 72-73) exhibit maximum mitotic index of 16-18% (3.8 mm diameter, 0.055 g, 1.55 million cells at week 2.5 5 mm, 0.11 g, 1.95 million cells at week 3.5), with cell number increasing at 0.5-0.6 million per week representing population doubling every 8-10 days during this absolute peak period, maximum growth rates of 1-1.2 mm diameter per week and 0.055-0.075 g per week, and maximum expression of cyclin B and CDK genes as measured by qRT-PCR showing 4-8 fold higher expression than at fruit set or lag phase. Chardonnay peak division stage berries at week 3 post-fruit set show spatial distribution of division with immunohistochemistry revealing highest mitotic index in inner mesocarp (18-22% cells positive for phospho-histone H3) versus middle mesocarp (14-17% positive) versus outer mesocarp (8-11% positive), with overall berry average of 16% mitotic index, demonstrating that peak division is highest in central berry regions and that tissue-specific division patterns exist even during peak overall division period. Syrah peak division stage berries experiencing controlled temperature treatments show optimal 25°C producing maximum mitotic index of 17-19% and cell increase rate of 0.58-0.68 million per week, versus 30°C showing reduced mitotic index 13-15% and rate 0.42-0.52 million per week, versus 35°C heat stress showing severely impaired division with mitotic index 7-9% and rate 0.22-0.32 million per week, demonstrating temperature sensitivity of peak division with heat stress particularly damaging during this critical high-activity period. Cabernet Franc comparison of peak division stage berries (week 3, 16.5% mitotic index, 0.52 million cells per week increase) versus declining division stage berries (week 5, 6% mitotic index, 0.18 million per week) versus lag phase berries (week 7, 1.5% mitotic index, <0.05 million per week) demonstrates the transient nature of peak division as a brief high-activity window within the broader cell division stage, with activity declining sharply after peak and approaching cessation by lag phase onset.
A peak division stage berry is a rapid division stage berry at the absolute maximum of cell division activity (BBCH 72-73, approximately 2-3.5 weeks post-fruit set), characterized by maximum mitotic index, maximum cell number increase rate, and maximum growth vigor, representing the brief period of highest cell proliferation within the cell division stage. Peak division stage berries are characterized by= (1) maximum mitotic index typically 14-18% of mesocarp cells in M phase, representing the highest proportion of dividing cells during berry development (2) maximum cell number increase rate typically 0.4-0.6 million cells per week, with cell population doubling approximately every 7-12 days during peak division (3) cell number typically 1.4-2 million cells at peak division stage (mid-range within overall 0.8-2.8 million progression during Phase I) (4) maximum cell cycle progression rate with individual cells completing division cycles in 4-7 days during peak activity (5) berry diameter typically 3.5-5.5 mm during peak division, representing mid-Phase I size (6) berry fresh weight typically 0.035-0.12 g during peak division (7) maximum growth rate with diameter increasing at 0.8-1.3 mm per week and fresh weight increasing at 0.04-0.1 g per week during peak division period. Peak division stage berries show maximum expression of cell cycle regulatory genes= cyclin B (G2/M transition regulator) at peak expression cyclin-dependent kinases (CDKs driving cell cycle progression) maximally active DNA replication machinery genes (DNA polymerases, MCM proteins, PCNA) at maximum expression supporting rapid S phase progression mitotic machinery genes (kinesins, cohesins, separins) maximally expressed supporting M phase. Peak division stage berries show maximum resource demand= carbon import from leaves at peak rates supporting biosynthesis for new cell production nitrogen demand high for protein synthesis in proliferating cells high water import supporting cell expansion concurrent with division maximum respiration rates (45-60 nmol CO₂ g⁻¹ s⁻¹) providing ATP for biosynthesis. Peak division stage berries represent the moment of maximum opportunity for cultural interventions affecting final berry size= irrigation applied during peak division has maximum effect on final cell number by supporting maximum division rates crop thinning during peak division allows redistributed resources to support enhanced division in retained berries deficit irrigation during peak division creates maximum cell number reduction and size limitation if that is desired (e.g., for wine quality). Peak division stage berries show maximum sensitivity to stress= heat stress (>35°C) during peak division can severely impair division reducing final cell number by 30-50% water deficit during peak division has maximum impact on final size nutritional deficiencies during peak division limit division most severely. The peak division stage is brief (typically 1-2 weeks duration) representing the apex of a rise-and-fall pattern of division activity, with division accelerating during weeks 1-2, peaking during weeks 2-3.5, then declining during weeks 4-6.
Peak_Division_Stage_Berry represents the absolute apex of cell division activity (weeks 2-3.5, BBCH 72-73) within the broader Rapid_Division_Stage (weeks 1-4) and Cell_Division_Stage (weeks 0-6). Peak division represents a brief critical window of maximum mitotic activity and maximum sensitivity to factors affecting cell proliferation. Understanding peak division stage timing and characteristics is important for timing interventions affecting berry size, for understanding cellular mechanisms of berry growth, and for assessing stress effects during the most vulnerable high-division period.
Absolut-Maximal-Zellteilungs-Stadium
absolut maximale Proliferations-Beere
absolute-peak-division berry
maximale Teilungs-Stadium-Beere
maximum-mitotic berry
Ojeda, H. et al. (2002). Influence of water deficits on grape berry growth. Vitis 41=141-145 Coombe, B.G. (1976). Development of fleshy fruits. Annual Review of Plant Physiology 27=507-528.
Peak Division Stage Berry
Syrah in Rhône Valley shows peak pollen availability during mid-June Year N+1 during peak anthesis, with widespread anther dehiscence releasing abundant pollen creating visible pollen clouds under favorable conditions, fresh pollen showing 75-88% germination capacity, pollen density maximizing in cluster atmosphere, under warm Mediterranean conditions of 25-29°C temperature, 48-62% relative humidity, light winds facilitating dispersal, supporting efficient pollination typical of this mid-to-late season cultivar
Peak pollen availability represents the maximum pollen production and presence period when greatest quantities of viable pollen become available for pollination through widespread anther dehiscence and sustained pollen release, establishing optimal male reproductive capacity enabling efficient pollination across cluster. This critical male function maximum occurs during peak anthesis in mid-June in temperate regions Year N+1, typically coinciding with or slightly following maximum flower opening, spanning 1-2 days when pollen density reaches maximum levels through cumulative production from recently opened flowers. Pollen production intensity shows maximum levels with numerous anthers simultaneously undergoing dehiscence releasing mature pollen grains, typical flower producing 20,000-60,000 pollen grains from five anthers representing substantial reproductive investment, and cluster-level production reaching millions of pollen grains creating dense pollen atmosphere in favorable conditions. Anther dehiscence pattern at peak shows widespread anther opening throughout cluster with apical regions showing 70-90% dehisced anthers from earliest-opened flowers, middle regions showing 60-80% active dehiscence representing peak activity zone, and basal regions showing 30-60% beginning dehiscence as opening continues, creating overall maximum dehiscence proportion. Pollen release mechanism involves anther wall dehydration creating mechanical tension opening longitudinal stomium slits, sudden rupture of anther locules liberating pollen grains from internal positions, and physical agitation from wind or flower movement dislodging pollen from anthers dispersing into atmosphere. Pollen characteristics at peak include optimal structural condition with intact exine wall and cytoplasmic contents, tricellular maturity state containing vegetative cell and two sperm cells ready for fertilization, and maximum viability with 70-90% pollen showing germination capacity under optimal laboratory conditions. Pollen viability timing shows peak immediately upon release from freshly dehisced anthers declining progressively over hours to days, with viability reducing to 50-70% after 6-12 hours exposure to ambient conditions, 30-50% after 24 hours, and 10-30% after 48 hours, creating critical 12-24 hour window of maximum functional viability. Environmental effects on viability include temperature with optimal 20-28°C maintaining viability while excessive heat above 35°C causes rapid viability loss through protein denaturation, humidity with moderate 40-70% maintaining pollen hydration and viability while very low humidity below 30% causes rapid desiccation reducing viability and very high humidity above 80% promotes fungal growth on pollen surfaces, and rain catastrophically destroying viability through excessive hydration and physical washing. Pollen dispersal at peak creates characteristic pollen clouds visible surrounding flowering clusters under favorable dry windy conditions, with light pollen grains 25-35 micrometers diameter easily becoming airborne through wind transport, traveling distances from centimeters within same flower to meters between adjacent vines, and settling on receptive stigma surfaces throughout cluster and neighboring clusters. Pollen density in cluster atmosphere reaches maximum during peak with estimated concentrations of 100-1000 pollen grains per cubic centimeter in immediate cluster vicinity under favorable conditions, declining with distance from source but remaining elevated 10-100 grains per cubic centimeter at vine scale, providing abundant pollen availability for pollination. Pollen-stigma contact frequency increases substantially during peak through high pollen density and widespread stigma receptivity, with individual receptive stigma potentially receiving 10-50 pollen grains during peak period though only 1-2 successful germinations required for fertilization, creating high-probability pollination opportunity. Pollen germination on receptive stigmas proceeds rapidly during peak with optimal conditions supporting germination initiation within 1-4 hours after contact, pollen grain hydrating from stigma exudate enabling metabolic activation, and pollen tube emergence through aperture penetrating stigma surface beginning growth toward ovary. Pollination efficiency during peak reaches maximum with abundant fresh viable pollen and widespread receptive stigmas creating optimal probability of successful pollen-stigma contact, favorable environmental conditions supporting pollen viability and germination, and high spatial density of reproductive structures facilitating efficient pollination within and between flowers. Self-pollination predominance shows characteristic pattern with majority of successful pollinations occurring within individual flowers or between flowers on same cluster through close proximity, wind-assisted transfer playing important role in distributing pollen among neighboring flowers, and limited cross-pollination between vines occurring but representing minor proportion of total successful pollinations. Atmospheric conditions affecting pollen dispersal include wind speed with light-moderate 5-15 km per hour optimal for pollen transport enhancing pollination efficiency, calm conditions below 3 km per hour reducing dispersal though self-pollination remains effective, and strong wind above 20 km per hour potentially dispersing pollen excessively reducing local availability or causing pollen desiccation. Temperature effects during peak show optimal 24-30°C supporting maximum pollen production, release, and viability, cool conditions below 18°C reducing anther dehiscence and pollen viability, and excessive heat above 33°C potentially causing pollen sterility through heat stress effects on developing pollen or reducing viability through rapid desiccation. Humidity during peak preferably 45-65% relative humidity balancing pollen viability maintenance with anther dehiscence facilitation, high humidity above 75% interfering with dehiscence preventing pollen release, and low humidity below 35% causing rapid pollen desiccation though facilitating dispersal. Metabolic activity in pollen and anthers shows maximum levels with elevated respiration in anthers supporting dehiscence processes, stored reserves in pollen grains providing energy for germination and initial tube growth, and metabolic activation upon hydration enabling rapid germination on receptive stigmas. Resource requirements for pollen production emphasize substantial carbohydrate investment with pollen containing high concentrations of storage compounds supporting germination and tube growth, protein content supporting cytoplasmic function and tube development, and specialized lipids in pollen coat affecting stigma interactions. Boron nutrition critically affects pollen function with adequate boron essential for pollen germination and tube growth, deficiency causing pollen sterility or poor germination severely reducing fruit set even with abundant pollen production, requiring sufficient boron availability during pollen development and germination phases. Anthesis synchronization implications show that peak pollen availability coinciding with peak stigma receptivity creates optimal pollination window, while temporal mismatch between pollen and stigma peaks reduces pollination efficiency affecting fruit set, highlighting importance of developmental synchrony for reproductive success. The process represents second of three peak anthesis aspects focusing on male reproductive capacity, developing concurrently with or shortly after maximum flower opening and overlapping with optimal pollination window. Duration represents brief 1-2 day period of maximum pollen presence, with rapid viability decline necessitating efficient pollination during this window. Cultivar variation appears in pollen production quantity with some varieties producing abundant pollen supporting high fruit set while others show modest production potentially limiting set, pollen viability characteristics with some showing robust viability 80-95% while others show reduced 60-75% viability, and pollen release timing relative to stigma receptivity affecting pollination efficiency. Regional timing shows peak pollen availability during mid-June in warm climates, mid-June in temperate regions, and mid-late June in cool areas, coinciding with peak anthesis and requiring favorable weather for optimal function. Viticultural significance includes representing critical determinant of pollination success and fruit set, requiring optimal environmental conditions particularly temperature and humidity for maximum pollen viability and function, emphasizing boron nutritional importance for pollen germination, representing period of maximum weather vulnerability with rain catastrophically affecting pollen availability, and establishing male reproductive capacity as potential fruit set limitation in unfavorable conditions. Research applications include analyzing pollen biology including production, viability dynamics, and germination characteristics, investigating environmental effects on pollen function affecting reproductive success, evaluating nutritional requirements particularly boron for normal pollen development and function, quantifying pollen dispersal patterns and pollination efficiency under varying conditions, and assessing genetic variation in pollen characteristics affecting reproductive success and fruit set.
Peak_Pollen_Availability is second of three Peak_Anthesis aspects, distinguished by maximum male reproductive capacity through abundant viable pollen. Develops concurrently with or shortly after Maximum_Flower_Opening. Overlaps with Optimal_Pollination_Window. Represents critical male contribution to pollination success. Brief 1-2 day period of maximum viability requiring rapid pollination. Environmental conditions particularly temperature and humidity critically affect viability. Boron nutrition essential for function.
Pollenmaximum
maximale Pollenproduktion
maximum pollen production
optimal pollen presence
optimale Pollenpräsenz
peak pollen release
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 375-382 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 287-292 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Peak Pollen Availability
Syrah in Rhône Valley shows peak pollen discharge on June 10-11 Year N+1 at BBCH 65, with 58-72% anthers simultaneously releasing pollen creating visible pollen clouds during morning hours 9am-11am, atmospheric concentration estimated 700-1200 grains per cubic centimeter, under warm Mediterranean conditions of 26-30°C temperature, 48-62% relative humidity, light winds 7-13 km per hour, supporting intensive pollination enabling 52-66% fruit set typical of this mid-to-late season cultivar during optimal 1-2 day peak discharge determining majority yield
Peak pollen discharge represents the maximum pollen liberation intensity when greatest proportion of anthers simultaneously release pollen creating characteristic intensive shedding activity with abundant pollen liberation throughout cluster, establishing optimal male gamete availability supporting maximum pollination opportunity. This culminating release intensity occurs during peak active shedding in mid-June in temperate regions Year N+1, typically spanning 1-2 days when widespread coordinated anther dehiscence creates maximum simultaneous pollen release, corresponding to BBCH 65 full bloom stage. Discharge intensity characteristics show maximum daily pollen release with 40-70% anthers actively discharging pollen simultaneously creating peak liberation activity, cumulative effect of sustained shedding from recently dehisced anthers maintaining high atmospheric concentration, and peak representing optimal coordination of male reproductive capacity. Anther distribution at peak shows dehisced anthers throughout cluster with apical regions showing 70-90% anthers discharged from earliest maturation, middle regions showing 60-80% active discharge representing highest density of recently opened anthers, and basal regions showing 40-60% beginning discharge creating overall maximum cluster-level pollen availability. Pollen liberation rate at peak shows maximum output with continuous release from numerous anthers creating sustained high-intensity shedding, estimated individual anther releasing 50-70% total pollen content during first hours following dehiscence, and cluster-level production reaching millions of grains per hour during peak activity. Atmospheric pollen concentration at peak reaches maximum levels with estimated 500-1500 grains per cubic centimeter in immediate cluster vicinity under optimal conditions representing 5-10 fold increase above background, visible pollen clouds forming around flowering clusters under favorable dry conditions, and elevated concentrations 50-200 grains per cubic centimeter extending vine scale supporting efficient pollination. Visual appearance at peak discharge shows characteristic pollen cloud phenomenon with pale yellow haze visible surrounding flowering clusters, particularly evident during morning hours 9am-11am when discharge typically maximal, and obvious pollen fall visible as fine dust accumulating on surfaces below clusters. Pollen cloud dynamics show formation during favorable conditions when temperature 24-30°C and humidity 50-65% enable efficient liberation and airborne suspension, maximum visibility during peak discharge when release intensity highest, and dissipation when wind disperses pollen or humidity rises causing aggregation and settling. Temporal pattern at peak shows characteristic morning maximum coinciding with optimal environmental conditions as temperature rises and humidity declines, typical peak activity 9am-12pm in temperate regions, and daily rhythm with repeated peak periods throughout 1-2 day maximum discharge phase. Environmental conditions at peak include temperature 24-30°C optimal for maximum discharge through efficient tissue dehydration and pollen liberation, moderate humidity 50-65% maintaining pollen viability and dispersibility without aggregation or prevented liberation, light-moderate wind 6-14 km per hour enhancing dispersal without excessive loss, and overall stable favorable weather enabling sustained intensive discharge. Temperature effects on peak discharge show warm conditions promoting maximum liberation through optimal tissue mechanics and pollen surface properties, cool conditions potentially reducing discharge intensity, and excessive heat above 32°C potentially affecting pollen viability though generally facilitating release. Humidity effects show critical importance with moderate levels optimal for maintaining high discharge rate and pollen viability, elevated humidity above 70% potentially reducing discharge through maintained anther hydration and causing pollen aggregation, and low humidity below 45% potentially causing excessive desiccation affecting viability though facilitating liberation. Wind effects show beneficial role with light-moderate wind enhancing dispersal creating better distribution and potentially agitating anthers dislodging additional pollen, calm conditions allowing pollen cloud formation though reducing long-range dispersal, and strong wind potentially dispersing pollen excessively reducing local availability. Pollen viability at peak discharge shows maximum proportion of viable grains with fresh pollen exhibiting 75-90% germination capacity, continuous production maintaining viable atmospheric pool, and brief window of optimal viability before progressive decline affects aged airborne pollen. Pollination opportunity at peak reaches absolute optimum with maximum fresh viable pollen availability creating abundant male gametes, widespread stigma receptivity providing numerous capture targets, and favorable spatial and temporal conditions supporting efficient pollen-stigma encounters, enabling fruit set 55-75% in good-setting cultivars under optimal conditions. Fruit set determination at peak shows outcomes during this critical 1-2 day period contributing 40-55% total berry number representing single largest contribution period, making peak discharge conditions dominant influence on season yield similar to overall peak anthesis window. Coordinated discharge at peak suggests possible synchronization mechanisms with developmental uniformity producing anthers reaching maturity simultaneously, common environmental responses triggering coordinated dehiscence as conditions become favorable, and possible physiological coordination though mechanisms unclear, creating concentrated male reproductive activity. Self-pollination efficiency at peak maximizes through immediate pollen availability in same flower with freshly released pollen falling directly onto nearby stigma, close anther-stigma proximity facilitating rapid encounter, and continued discharge maintaining pollen supply for extended receptivity period. Neighboring flower pollination at peak intensifies through abundant atmospheric pollen dispersing short distances within cluster, concentrated pollen availability creating high encounter probability, and coordinated timing supporting effective pollination between nearby synchronized flowers. Weather vulnerability at peak reaches maximum with reproductive success extremely sensitive to conditions during this brief 1-2 day window, favorable stable conditions supporting high fruit set while adverse weather particularly rain causing catastrophic failure potentially reducing set to 15-30%, demonstrating extreme importance of favorable conditions during peak discharge. The process represents first aspect of active pollen shedding focusing on maximum intensity, followed by maximum pollen dispersal emphasizing atmospheric distribution. Duration typically spans 1-2 days representing peak intensity period, with warm stable conditions producing sharp intense peak while variable conditions may broaden or reduce intensity. Cultivar variation in peak discharge shows some varieties reaching very high intensity with coordinated massive release while others show moderate peak, timing relative to stigma receptivity peak affecting pollination efficiency, and discharge efficiency affecting atmospheric concentration achieved. Regional timing shows peak pollen discharge during mid-June in warm climates at approximately 1050-1100 growing degree days base 10°C, mid-June in temperate regions at 950-1000 GDD, and mid-late June in cool areas at 850-900 GDD, coinciding with peak anthesis and full bloom. Viticultural significance includes representing most critical male reproductive moment determining majority of fruit set, requiring optimal environmental conditions for maximum discharge efficiency and viability, demonstrating extreme weather vulnerability with brief window determining season outcomes, and providing visual confirmation through pollen clouds of maximum reproductive activity. Research applications include analyzing peak discharge mechanisms and coordination, quantifying pollen production rates and atmospheric concentrations, evaluating environmental effects on discharge intensity, assessing pollination efficiency during peak period, and developing models predicting fruit set from peak discharge characteristics and conditions.
Peak_Pollen_Discharge is first aspect of Active_Pollen_Shedding, distinguished by maximum liberation intensity. Represents culminating 1-2 day period of male reproductive activity. Corresponds to BBCH 65 full bloom. Creates visible pollen clouds under favorable conditions. Determines 40-55% fruit set during brief window. Extreme weather vulnerability particularly to rain. Followed by Maximum_Pollen_Dispersal discussing atmospheric distribution. Most critical male reproductive period.
Pollenabgabemaximum
Spitzen-Pollenabgabe
highest pollen discharge
höchste Pollenliberation
maximale Pollenfreisetzung
maximum pollen release
peak pollen liberation
pollen release maximum
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 382-385 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 292-295
Peak Pollen Discharge
Pinot Noir in Burgundy shows peak pollen release during mid-June Year N+1 at BBCH 63-65 over 2-3 days June 8-10, with 58-72% anthers simultaneously discharging creating visible pollen clouds during morning hours, atmospheric concentration 600-1100 grains per cubic centimeter, fresh pollen viability 78-88%, under continental conditions of 22-27°C temperature, 54-68% relative humidity, light winds 7-12 km per hour, supporting optimal pollination enabling 60-70% fruit set typical of this early-to-mid season cultivar during critical 2-3 day peak determining majority yield
Peak pollen release represents the culminating male reproductive phase when anther dehiscence, pollen discharge, and atmospheric dispersal reach maximum coordinated intensity creating optimal pollen availability and distribution, establishing ideal conditions for maximum pollination success and fruit set determination. This critical male reproductive culmination occurs during peak flowering in mid-June in temperate regions Year N+1, typically spanning 2-3 days when widespread coordinated pollen liberation creates maximum male gamete availability, corresponding to BBCH 63-65 peak anthesis stages. Peak characteristics show simultaneous occurrence of optimal male reproductive elements including widespread anther dehiscence with 50-75% anthers actively releasing pollen, maximum pollen discharge intensity producing abundant fresh viable grains, optimal atmospheric pollen concentration creating 500-1500 grains per cubic centimeter near clusters, and favorable environmental conditions supporting pollen viability and dispersal. Temporal coordination at peak shows critical convergence of male reproductive capacities with peak discharge timing coinciding with maximum atmospheric dispersal capability, sustained intensive pollen production maintaining viable atmospheric pool, and optimal overlap with widespread stigma receptivity creating ideal pollination window. Pollen production intensity reaches maximum at peak with numerous anthers simultaneously discharging creating cumulative cluster-level production estimated millions of grains per hour, continuous fresh pollen release maintaining viable atmospheric supply, and overall male gamete availability sufficient for supporting high fruit set potential. Atmospheric pollen characteristics at peak include maximum concentration providing abundant male gametes throughout cluster atmosphere, optimal spatial distribution with elevated levels extending vine and vineyard scale, visible pollen cloud formation under favorable conditions creating obvious evidence of intensive release, and sustained temporal persistence maintaining availability during stigma receptivity period. Pollen cloud phenomenon at peak shows characteristic visual display with pale yellow haze surrounding flowering clusters, particularly evident during morning peak hours 9am-11am when conditions optimal, sound occasionally audible during very intensive synchronized discharge though typically silent, and overall sensory confirmation of maximum male reproductive activity. Pollen viability at peak shows maximum proportion of viable grains with continuous fresh pollen production maintaining 75-90% germination capacity in atmospheric pool, brief optimal window before progressive aging reduces viability, and critical importance of pollination during peak period capturing highest-quality male gametes. Environmental conditions at peak typically optimal with temperature 24-30°C supporting efficient dehiscence, pollen viability, and atmospheric transport, moderate humidity 50-65% maintaining pollen dispersibility and viability without aggregation, light-moderate wind 6-14 km per hour enhancing distribution, and stable favorable weather enabling sustained intensive release over 2-3 day period. Temperature optimization at peak shows warm conditions promoting maximum dehiscence through efficient anther dehydration, maintaining pollen surface properties enabling dispersal, and supporting atmospheric mixing enhancing distribution, with optimal 25-28°C range supporting all pollen release processes simultaneously. Humidity optimization shows moderate levels critical for balancing multiple requirements including sufficient anther dehydration enabling dehiscence, maintained pollen viability without desiccation damage, appropriate dispersibility preventing aggregation and premature atmospheric removal, and optimal 52-68% range supporting coordinated male reproductive processes. Wind optimization shows light-moderate speeds ideal for enhancing dispersal without excessive pollen loss, facilitating atmospheric mixing improving distribution, potentially agitating anthers dislodging additional pollen, and optimal 7-13 km per hour range supporting effective male gamete distribution. Pollination opportunity at peak reaches absolute optimum with maximum fresh viable pollen widely distributed creating abundant male gamete availability, extensive stigma receptivity providing numerous capture sites, favorable spatial arrangement supporting efficient encounters, and optimal temporal coordination creating ideal reproductive window. Fruit set determination at peak shows outcomes during this critical 2-3 day period dominating final berry number with successful pollinations during peak contributing 50-70% total fruit set, making peak pollen release conditions primary determinant of season yield, and demonstrating critical importance of favorable male reproductive function. Self-pollination efficiency at peak maximizes through abundant immediate pollen availability in same flower with continuous discharge maintaining supply, close anther-stigma proximity facilitating rapid encounters, and fresh viable pollen ensuring high germination success supporting effective self-pollination accounting for 60-80% successful reproductions. Cross-pollination contribution at peak shows enhanced role during maximum atmospheric dispersal with elevated pollen concentration enabling effective neighboring flower pollination accounting for 20-35% successful reproductions, inter-vine dispersal contributing 5-15% particularly in dense plantings with high atmospheric pollen levels, and demonstrating secondary but significant role of atmospheric dispersal in reproductive success. Weather dependency at peak shows extreme sensitivity with reproductive success critically dependent on favorable conditions during brief 2-3 day window, optimal stable weather supporting high fruit set 60-75% in good-setting cultivars, while adverse conditions particularly rain causing catastrophic failure potentially reducing set to 15-30%, demonstrating that peak pollen release represents single most weather-vulnerable period alongside stigma receptivity peak. Rain effects at peak show devastating interference with water preventing anther dehiscence eliminating continued pollen production, washing pollen from atmosphere destroying atmospheric pool, eliminating pollen viability through excessive hydration, and promoting fungal infection, with rain during peak potentially reducing fruit set by 60-80% compared to favorable conditions. Developmental coordination at peak shows remarkable synchronization with peak pollen release coinciding with peak anthesis and maximum cap fall, widespread flower opening and stigma receptivity aligning temporally with maximum pollen availability, and integrated reproductive system coordination creating optimal pollination window determining season productivity. Metabolic activity at peak shows maximum respiratory demand in discharging anthers supporting intensive pollen release, elevated energy consumption in pollen maintaining viability and germination capacity, and overall high metabolic investment in male reproductive function during brief critical period. Resource requirements at peak emphasize carbohydrate demand for supporting intensive metabolic activity in anthers and pollen, nitrogen for maintaining protein function in pollen and reproductive tissues, critical boron for supporting subsequent pollen germination and tube growth, and adequate mineral nutrition supporting overall reproductive function. Hormonal environment at peak involves possible regulatory signals coordinating dehiscence timing though mechanisms poorly understood in grapevine, gibberellins potentially affecting anther development and pollen maturation, and overall hormonal context supporting male reproductive function during flowering. The process shows two distinguishable aspects progressing from optimal pollen cloud formation creating visual and atmospheric evidence of peak activity, through maximum pollen availability establishing ideal male gamete supply for pollination. Duration typically spans 2-3 days representing peak male reproductive period paralleling peak anthesis, with warm stable conditions producing compressed 2-day peak while variable conditions may extend to 3-4 days. Cultivar variation in peak release shows differences in intensity with some varieties producing very high pollen concentrations while others show moderate peaks, peak duration affecting temporal window for optimal pollination, and coordination with stigma receptivity affecting reproductive efficiency. Regional timing shows peak pollen release during mid-June in warm climates at approximately 1050-1100 growing degree days base 10°C, mid-June in temperate regions at 950-1000 GDD, and mid-late June in cool areas at 850-900 GDD, coinciding precisely with peak anthesis period. Viticultural significance includes representing most critical male reproductive period determining 50-70% fruit set and season yield, requiring optimal environmental conditions for efficient release and viability maintenance, demonstrating extreme weather vulnerability with brief window determining productivity, providing visual confirmation through pollen clouds of successful male function, and establishing peak pollen release alongside peak stigma receptivity as dominant yield determinants. Research applications include analyzing coordination mechanisms producing synchronized peak release, quantifying pollen production rates and atmospheric dynamics during peak, evaluating environmental effects on peak intensity and duration, assessing pollination efficiency during optimal period, developing models predicting fruit set from peak pollen release characteristics, and investigating climate change impacts on peak timing and conditions affecting productivity.
Peak_Pollen_Release represents culminating male reproductive phase parallel to Peak_Anthesis within Flowering. Corresponds to BBCH 63-65 stages. Subdivided into Optimal_Pollen_Cloud_Formation and Maximum_Pollen_Availability. Represents most critical 2-3 days for male contribution to fruit set determining 50-70% berry number. Extreme weather vulnerability particularly to rain. Requires optimal coordination of dehiscence, discharge, dispersal, and viability. Temporal alignment with peak stigma receptivity creates optimal pollination window. Most important male reproductive period.
Spitzen-Pollenfreisetzung
culminating male reproduction
maximale Pollenproduktionsphase
maximum pollen production phase
männlicher Reproduktionshöhepunkt
optimal pollen release period
optimale Pollenabgabeperiode
pollen release peak
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 380-388 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 290-298
Peak Pollen Release
Syrah peak ripeness berries in warm vintage achieve maximum development at week 18= 27% sugars (near maximum for this cultivar), 5.2 g/L acidity (acids declined substantially), pH 3.88 (high), 4.5 mg/g skin anthocyanins (peak color), intense blackberry and black pepper flavors at maximum concentration, fully mature desiccated brown seeds, representing absolute peak of ripening progression suitable for powerful concentrated wine style but potentially over-ripe for balanced fresh style. Zinfandel peak ripeness berries showing characteristic extreme ripeness= 28-30% sugars (very high for this late-ripening cultivar), 4.5-5.5 g/L acidity, pH 3.9-4.1, 3.5-4.5 mg/g anthocyanins, intensely concentrated jammy fruit flavors, with some berries showing beginning raisining, representing peak ripeness appropriate for traditional Zinfandel style emphasizing power and ripeness but challenging for balanced moderate-alcohol styles. Cabernet Sauvignon comparison of optimal ripeness (week 18= 24.5% sugars, 6.3 g/L acidity, pH 3.72, 3.6 mg/g anthocyanins, balanced ripe fruit character) versus peak ripeness (week 20= 27% sugars, 5.3 g/L acidity, pH 3.92, 4.2 mg/g anthocyanins, very intense concentrated fruit) demonstrates trade-off between maximum ripeness development and compositional balance, with optimal ripeness harvest producing more balanced wines while peak ripeness harvest produces more powerful concentrated wines with higher alcohol and lower acid potentially compromising balance and aging potential. Merlot peak ripeness berries in warm site showing beginning of senescence transition= 26.5% sugars representing near-maximum accumulation with rates declining from 0.8% per week to 0.2% per week, anthocyanins at 3.8 mg/g showing plateau, berry weight declining slightly from maximum (2.9 g to 2.85 g) through transpiration, firmness declining to very soft 0.9 N, representing peak ripeness verging on overripeness with harvest urgency to capture peak development before quality degradation. Pinot Noir peak ripeness timing across three climate zones shows climate effects on peak= cool climate achieves peak at 23.5% sugars, 7 g/L acidity (balanced peak) moderate climate at 25% sugars, 6 g/L acidity (concentrated peak) warm climate at 26.5% sugars, 5 g/L acidity (extreme peak potentially excessive for quality Pinot Noir), demonstrating that peak ripeness is climate-dependent with warmer climates producing higher peak ripeness potentially exceeding quality optima.
A peak ripeness berry is a fully ripe berry at the absolute maximum of ripeness development, characterized by maximum sugar accumulation, maximum flavor intensity, maximum phenolic development, and maximum physiological maturity, representing berries at the apex of the ripening progression before senescence or overripening begins. Peak ripeness berries are characterized by= (1) sugars at maximum or near-maximum levels achievable under the given environmental conditions, typically 25-28% for wine grapes in favorable vintages, representing the peak of sugar accumulation before rates slow dramatically or cease (2) flavor compounds at maximum intensity and complexity with fully developed varietal character, maximum aromatic compound concentration, optimal balance of fruit, floral, spice, and other flavor notes (3) anthocyanins (red cultivars) at or very near maximum concentration typically 3-5 mg/g skin or higher in deeply colored cultivars, representing peak color development (4) tannins at maximum ripeness with optimal polymerization state, minimal astringency, ideal extractability for wine production (5) acids declined to lower levels typically 5-6.5 g/L reflecting advanced malic acid respiration, with pH often 3.6-4 or higher (6) seeds at maximum maturity showing full lignification, brown color throughout, maximum desiccation (7) berry physiological state showing maximum development with berries beginning transition from active ripening to senescence. Peak ripeness berries may represent optimal ripeness for certain production styles emphasizing maximum ripeness= late-harvest styles, powerful concentrated wines, raisined or partially dried styles but may represent excessive ripeness for styles emphasizing balance and freshness. Peak ripeness berries show maximum resource accumulation= sugars at peak representing maximum carbon import and accumulation in vacuoles anthocyanins at peak representing maximum phenylpropanoid pathway activity flavor compounds at peak representing maximum biosynthesis of volatile and non-volatile secondary metabolites water content potentially declining from peak berry size through transpiration and respiration exceeding import. Peak ripeness berries represent a transitional state= brief period at maximum ripeness development before progression to overripeness some quality parameters may be optimal (sugar, flavor intensity, color) while others may be suboptimal (acids potentially too low, pH potentially too high, balance potentially compromised) harvest decision requires assessing whether maximum ripeness aligns with quality goals or whether earlier harvest at optimal balance is preferred. Peak ripeness timing varies with cultivar, climate, and vintage= early-ripening cultivars like Pinot Noir may reach peak ripeness at 23-25% sugars late-ripening cultivars like Cabernet Sauvignon may reach peak at 26-28% sugars warm climates and warm vintages promote earlier achievement of peak ripeness cool climates and cool vintages may prevent achievement of full peak ripeness. Peak ripeness versus optimal ripeness distinction= peak ripeness emphasizes maximum development potentially at cost of balance optimal ripeness emphasizes ideal balance even if some parameters (sugar, color) are not at absolute maximum quality-focused viticulture often targets optimal ripeness rather than peak ripeness to achieve balance and elegance.
Peak_Ripeness_Berry represents maximum ripeness development and is distinguished from Optimally_Ripe_Berry which emphasizes ideal balance versus maximum development. Peak ripeness may or may not align with optimal quality depending on production goals and style preferences. Understanding peak ripeness versus optimal ripeness distinction is important for harvest decisions, particularly for producers deciding between balance-focused earlier harvest versus ripeness-focused later harvest, and for understanding cultivar and climate effects on achievable ripeness levels and optimal harvest timing.
Höchstreife-Traube
Spitzenreife-Traube
apex-maturity berry
maximale Reife-Beere
maximum-ripeness berry
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 240-250 Jackson, D.I., Lombard, P.B. (1993). Environmental and management practices affecting grape composition. American Journal of Enology and Viticulture 44=409-430.
Peak Ripeness Berry
Syrah in Barossa Valley shows peak shoot elongation from mid-December to early January at BBCH 31-35 over 3 weeks, with shoots extending from 1400 mm at flowering to 2100-2400 mm following fruit set through sustained elongation rate of 42-52 mm per day, under warm climate conditions of 24-30°C mean temperature, irrigation maintaining adequate soil moisture, high solar radiation, and moderate fertility typical of this vigorous mid-to-late season cultivar in Southern Hemisphere warm region
Peak shoot elongation represents the maximum linear growth phase where elongation rate reaches highest sustained level before environmental limitations or developmental programming initiate growth decline, characterized by consistent rapid daily extension creating major portion of final shoot length and establishing definitive canopy architecture. This developmental stage typically occurs during mid-June to early-July in temperate regions corresponding approximately to BBCH 31-35 encompassing flowering through early fruit set, with elongation rate maintaining maximum of 40-60 mm per day for vigorous cultivars and 30-45 mm per day for moderate vigor types over 2-3 week period creating shoot length increase of 600-1200 mm. Peak elongation represents linear growth phase of sigmoid growth curve where rate remains at maximum for extended period as photosynthetic capacity, favorable environmental conditions, and developmental state combine to optimize growth resource availability. The apical meristem continues maximum activity producing new leaf primordia and contributing cells to shoot axis at consistent high rate, maintaining organized structure with active cell division cycles of 18-22 hours supplying cells for sustained rapid elongation. Internode lengths during peak elongation achieve maximum dimensions typically 80-150 mm reflecting optimal resource availability and growth conditions, with individual internode length varying based on position and microenvironmental conditions but generally remaining consistent during peak phase. Cell expansion in subapical zones operates at maximum sustainable rate with abundant water uptake and active wall loosening, expansion zone extending 25-35 mm behind apex with cells reaching final lengths of 300-600 micrometers depending on internode position and vigor characteristics. Leaves continue emerging and expanding at maximum rate with new leaves appearing every 2 days and rapidly expanding to large final dimensions, with mature leaf photosynthetic capacity fully established contributing strongly positive net carbon balance. Photosynthetic capacity operates at seasonal maximum with full canopy area developed and mature leaves functioning at peak efficiency, producing substantial excess assimilate beyond shoot growth requirements available for fruit development, root growth, and reserve accumulation. Carbon allocation during peak elongation shifts progressively from predominantly shoot growth toward increasing fruit development as reproductive demand accelerates following fruit set, with shoots consuming 40-60% their photosynthetic production for growth while contributing substantial excess to developing clusters and beginning significant reserve accumulation in permanent structures. Vascular system maintains maximum functionality with xylem conducting water at 4-8 mg per second supporting high transpiration and growth water requirements, and phloem transporting large assimilate quantities to multiple competing sinks including shoot growth, developing fruit, and storage tissues. Water demand reaches seasonal maximum during peak elongation as full canopy transpires under typically warm conditions, with shoot water use of 500-900 mg per hour requiring continuous adequate root uptake and soil moisture availability maintaining turgor and photosynthetic function. Nutrient demand continues at high levels particularly nitrogen supporting continued protein synthesis though declining as proportion of total demand as fruit development creates competing nutrient sink, phosphorus supporting energy metabolism in rapidly growing tissues, and potassium critical for fruit development quality attributes. Metabolic activity maintains elevated levels with respiration supporting growth processes though declining as proportion of photosynthetic production as mature leaves operate at maximum efficiency, creating favorable carbon balance supporting multiple growth processes simultaneously. Protein synthesis continues at high rates supporting shoot growth requirements while increasingly producing proteins for fruit development and reserve accumulation in permanent structures. Hormonal regulation maintains auxin from shoot apex supporting continued elongation though effects begin declining as reproductive hormones from developing fruit increasingly influence assimilate partitioning, gibberellins continue supporting cell expansion, and cytokinins promote continued cell division though root production may decline as root growth becomes limited by above-ground demands. Temperature during peak elongation typically remains in optimal 22-28°C range with mid-summer conditions, though excessive heat above 32°C can begin limiting growth efficiency through increased respiration and water stress. Light intensity remains high during long summer days supporting maximum photosynthetic production, with full canopy creating shading patterns affecting lower canopy leaves and potentially limiting whole-canopy photosynthetic efficiency if excessive density develops. Water availability becomes critical during peak elongation as maximum transpiration and growth demands coincide with typically warm dry conditions in many regions, requiring irrigation in water-limited regions maintaining adequate soil moisture preventing stress that rapidly arrests growth. Nutrient availability must remain adequate though excessive nitrogen during peak elongation can promote excessive vegetative vigor competing with fruit development, requiring balanced nutrition supporting both vegetative and reproductive growth. The process typically spans 2-3 weeks under favorable conditions with consistent optimal temperatures, adequate moisture, and balanced nutrition, with duration varying based on cultivar vigor characteristics and environmental limitations. Cultivar variation appears in maximum sustained elongation rate during peak phase, duration of peak elongation period, and sensitivity to environmental limitations or reproductive demand affecting transition to declining growth. Regional timing shows peak elongation occurring mid-June to early July in warm climates, late June to mid-July in temperate regions, and July in cool areas, typically spanning flowering through early fruit set developmental period when both vegetative and reproductive demands are high. Viticultural management considerations include maintaining adequate but not excessive water availability supporting growth while promoting moderate water stress initiating growth decline and improved fruit development, managing nitrogen availability avoiding excessive late vegetative growth competing with fruit quality development, implementing shoot trimming operations at appropriate timing removing excessive length while maintaining adequate leaf area for fruit maturation, monitoring canopy density managing excessive growth creating unfavorable fruit microclimate, and assessing growth uniformity across vineyard blocks identifying management requirement variations. Excessive peak elongation extending too long creates overly vigorous canopies with dense shading reducing fruit quality, while premature limitation of peak elongation reduces photosynthetic capacity potentially limiting fruit development and reserve accumulation. Research applications include analyzing mechanisms maintaining sustained maximum growth rate, quantifying assimilate partitioning between vegetative and reproductive sinks during competition period, evaluating environmental and management effects on peak elongation duration and transition to declining phase affecting canopy architecture, modeling growth dynamics for decision support optimizing vegetative-reproductive balance, and assessing genetic and environmental variation in growth patterns affecting productivity and quality potential.
Peak_Shoot_Elongation is third of five Shoot_Elongation subphases, distinguished by maximum sustained linear growth rate occurring typically during flowering through early fruit set. Transitions to Declining_Shoot_Elongation as environmental limitations or developmental programming reduce growth rate. Represents critical phase where vegetative-reproductive competition is highest. Timing and duration strongly affect final canopy architecture and fruit development balance.
Höchststreckungsphase
Spitzenvegetation
linear growth phase
lineare Wachstumsphase
maximale Triebstreckung
maximum elongation
peak vegetative growth
sustained rapid growth
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 235-248 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 152-165 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Peak Shoot Elongation
In a Burgundian Pinot Noir vineyard near Beaune, peak softening was documented during the third week of August Year N+1 (BBCH 83-84) when firmness measurements declined from 2.4 N to 1.5 N over 4 days (average 5.6% per day), coinciding with polygalacturonase activity reaching 125 units, pectin molecular weight dropping from 145 kDa to 85 kDa, and berry fresh weight increasing from 1.15 g to 1.35 g as sugar content rose from 17.5 to 20.8° Brix under warm conditions (daily maximum 26-28°C).
Peak_Softening represents the developmental interval during veraison when berry firmness decline reaches maximum velocity, characterized by the most rapid structural transformation from firm to pliable berry texture. This critical phase occurs during mid-veraison (BBCH 83-84), typically 2-5 days duration, when the rate of firmness reduction peaks at 3-6% per day compared to 2-4% during initial softening and 0.5-1.5% during progressive softening. The process is defined by simultaneous maxima in multiple degradative pathways: polygalacturonase activity reaches 80-150 units (5-10 fold above pre-veraison baseline), pectin methylesterase activity peaks at 100-180 units, and xyloglucan endotransglycosylase activity shows 4-8 fold elevation. Berry firmness measurements decline most precipitously during this interval, dropping from approximately 2.0-3.0 N to 1.2-2.0 N within 3-5 days, representing the steepest portion of the sigmoid softening curve. Cell wall structural changes reach maximum intensity: middle lamella pectin solubilization accelerates to 5-10% per day increase in soluble fraction (total soluble pectin 60-75%), pectin molecular weight drops rapidly from 120-180 kDa to 70-120 kDa, and cell-to-cell adhesion strength decreases by 40-60% from pre-peak values. The peak softening interval coincides with other veraison transformations including accelerated sugar accumulation (1.5-3.5% Brix increase per week), rapid anthocyanin synthesis in red cultivars (50-150 mg/kg per day in skin), and continued berry expansion (15-35 mg fresh weight per day). Temperature strongly influences peak softening dynamics: optimal conditions (22-28°C) support maximum enzyme activities and rapid texture transformation, while cooler temperatures (15-20°C) extend the peak period and reduce maximum rates, and heat stress (>32°C) can accelerate softening beyond optimal enzyme kinetics through enhanced membrane permeability and cellular disruption. Cultivar variation in peak softening characteristics is substantial: early-ripening varieties like Pinot Noir and Chardonnay exhibit sharper, more concentrated peaks (3-4 days) with maximum rates 4-7% firmness loss per day, mid-season cultivars like Cabernet Sauvignon show moderate peaks (4-5 days) with 3-5% daily decline, while late-season varieties like Mourvèdre display more gradual peaks (5-7 days) with 2-4% maximum rates. The peak softening phase marks the transition from structural berry to highly manipulable fruit, with significant viticultural implications: mechanical harvest feasibility declines sharply as berry integrity diminishes, rot susceptibility increases due to compromised skin strength and enhanced pathogen penetration, and bunch compression intensifies as individual berries lose mechanical resistance and conform to cluster geometry. Molecular regulation involves peak expression of softening-related genes: PG genes show 80-150 fold upregulation relative to green berry baseline, expansin genes reach 40-80 fold elevation, and xyloglucan endotransglycosylase genes achieve 20-50 fold increases. Hormonal orchestration is dominated by continued high abscisic acid levels (800-2500 ng/g fresh weight) maintaining ripening gene expression, while ethylene production shows cultivar-dependent patterns with slight elevation (0.5-2.0 nL g⁻¹ h⁻¹) in some varieties correlating with enhanced softening enzyme activity. Physiological consequences of peak softening extend beyond texture: increased membrane permeability facilitates enhanced metabolite transport including accelerated sugar import and anthocyanin accumulation, while reduced cellular compartmentalization allows mixing of vacuolar and cytoplasmic contents affecting flavor precursor availability and enzymatic reactions. Environmental stresses during peak softening can substantially alter berry quality development: water deficit moderately reduces softening intensity potentially maintaining firmer texture but may compromise berry expansion and metabolite dilution, while excessive irrigation can accelerate softening beyond optimal rates leading to burst-prone berries and diluted flavor compounds. Viticultural management targeting peak softening focuses on optimizing the timing and intensity of this transformation through careful irrigation scheduling to balance cell turgor with structural integrity, canopy management to moderate bunch zone temperatures and prevent excessive softening acceleration, and crop load adjustment to ensure adequate photosynthate supply supporting both structural transformation and metabolite accumulation without resource limitation that could compromise either process.
Peak_Softening is distinguished from Initial_Berry_Softening by representing the maximum velocity phase rather than onset, from Progressive_Softening by exhibiting peak rather than sustained moderate rates, and from Complete_Berry_Softening by focusing on the transformation intensity rather than final state achievement. Duration typically 2-5 days during mid-veraison. Maximum firmness decline rates: early cultivars 4-7% per day, mid-season 3-5% per day, late varieties 2-4% per day. Temperature is the primary rate-controlling environmental factor. Peak_Softening represents the most rapid structural transformation in berry development and marks the point of maximum vulnerability to mechanical damage and pathogen invasion.
Spitzen-Textur-Transformation
maximale Erweichungs-Intensität
maximale Festigkeits-Abnahme-Geschwindigkeit
maximum firmness decline velocity
maximum softening intensity
peak texture transformation
steepest firmness loss
steilster Festigkeits-Verlust
Keller M (2020) The Science of Grapevines 3rd Ed pages 228-232 berry softening dynamics and cell wall degradation Coombe BG, McCarthy MG (2000) Dynamics and environmental control of grape berry growth Australian J Grape Wine Res 6:73-83 firmness decline kinetics Nunan KJ, Davies C, Robinson SP, Fincher GB (2001) Expression patterns of cell wall-modifying enzymes during grape berry development Planta 214:257-264 enzyme activity peaks.
Peak Softening
In a Barossa Valley Shiraz vineyard, peak sugar accumulation was identified over 14 days in mid-February Year N+1 (BBCH 84-85) when berry Brix increased from 18.2° to 25.6° (7.4° gain, average 5.3% per week peak rate), hexose import calculated 128 μmol g⁻¹ h⁻¹ based on Brix trajectory and berry volume, VvHT1 expression 142-fold, cell wall invertase 118 units/g FW, berry fresh weight 1.68 g → 1.95 g (16% expansion during peak), under optimal Australian conditions (daily high 28-31°C, adequate irrigation, healthy canopy supporting maximum photosynthesis).
Peak_Sugar_Accumulation represents the maximum rate phase of hexose import and storage during mid-to-late veraison when all components of the sugar transport and accumulation pathway operate at optimal capacity producing the fastest sweetening observed during ripening that accounts for disproportionate fraction of total sugar gain achieved in compressed timeframe defining critical window for achieving commercial maturity levels. This peak accumulation occurs during mid-to-late veraison (BBCH 84-85), typically days 18-32 post-veraison onset spanning 10-18 days depending on cultivar and environmental conditions, and is characterized by berry Brix increasing at maximum rates 3.5-5.5% per week (0.50-0.78% per day) compared to active phase average 2.5-4.0% per week and late phase 1.2-2.8% per week, hexose import reaching maximum velocity 90-170 μmol combined glucose and fructose g⁻¹ h⁻¹ representing full saturation of transport machinery, daily sugar gain 180-280 mg hexose per berry per day (depending on berry size 1.6-2.4 g) enabling 4-8° Brix increase over 12-16 day peak period, and cumulative contribution where peak phase though brief in duration accounts for 30-50% of total ripening sugar gain demonstrating concentrated intensive accumulation. Biochemical characteristics at peak include hexose transporter genes (VvHT1, VvHT2, VvHT3, VvHT5, VvHT6) expressed at absolute maximum 100-150 fold above green berry collectively providing capacity substantially exceeding average active phase levels, cell wall invertase activity 80-140 units/g fresh weight producing hexoses from phloem sucrose at maximum velocity matching or exceeding cellular uptake capacity, plasma membrane hexose uptake operating near Vmax 85-150 μmol g⁻¹ h⁻¹ with substrate concentrations well above Km ensuring kinetic saturation, tonoplast transport maximized 65-120 μmol g⁻¹ h⁻¹ efficiently clearing cytoplasmic hexoses maintaining concentration gradients enabling continued plasma membrane uptake, and metabolic integration where partial hexose respiration 25-35% of import provides ATP supporting transport while majority 65-75% accumulates representing optimal storage efficiency. Source-sink dynamics at peak demonstrate phloem delivery maximized 250-450 mg sucrose per berry per day supported by canopy photosynthesis operating at seasonal maximum 15-28 g CO₂ fixed per vine per day under favorable conditions, sink strength at absolute maximum with berry demand overwhelming competing sinks receiving 75-90% of daily photosynthate, source-sink connectivity optimized through vascular development and phloem maturation enabling efficient translocation, and environmental conditions typically favorable during peak with moderate temperatures 25-30°C, adequate water, and high light supporting both source and sink processes. Cellular processes supporting peak include vacuolar expansion approaching maximum 95-97% cell volume creating storage capacity 1.8-3.2 nL per mesocarp cell, tonoplast membrane area at maximum following proliferation during earlier phases providing adequate surface for transport, osmotic potential declining most rapidly -0.22 to -0.32 MPa per week (from -2.0 to -2.6 MPa → -2.4 to -3.2 MPa over peak period) creating strong water potential gradient, turgor pressure maintained 0.3-0.6 MPa through balance of osmotic potential and cell wall resistance, and berry expansion continuing 12-22% during peak through osmotic water import demonstrating coupling between sugar accumulation and growth. Environmental factors producing peak conditions include temperature optimum 26-30°C enabling maximum enzyme activities and metabolic flux with this range occurring typically mid-to-late summer (northern hemisphere August, southern hemisphere February), photoperiod still long 13-15 hours providing extended photosynthesis period supporting substrate production, solar radiation high 20-30 MJ m⁻² per day saturating photosynthesis, water availability adequate with stem water potential -0.7 to -1.2 MPa supporting phloem function without severe stress, and absence of extreme weather events enabling uninterrupted accumulation during critical window. Hormonal regulation at peak shows ABA sustained at elevated plateau 1800-2800 ng/g maintaining maximum transporter expression and coordinating ripening processes, sugar signaling through hexokinase pathway with glucose-6-phosphate 35-65 μmol/g fresh weight creating strong feed-forward amplification, declining growth hormones having reached minimum maintaining metabolic focus on storage rather than structural development, and potential ethylene elevation slight 0.3-0.8 nL g⁻¹ h⁻¹ in some cultivars possibly fine-tuning ripening though grape generally non-climacteric with minimal ethylene effects. Cultivar variation in peak characteristics demonstrates high-Brix varieties (Gewürztraminer, Sémillon) achieving exceptional peak rates 4.8-6.2% Brix per week sustained 10-14 days with high transporter expression and efficient metabolism, moderate cultivars (Chardonnay, Merlot) showing 3.8-5.0% per week for 12-16 days representing typical commercial varieties, and lower-accumulation types (Pinot Noir cool climates, table grapes) displaying 3.0-4.2% per week over 14-20 days reflecting genetic differences in maximum capacity and duration. Quality implications of peak accumulation include timing and intensity determining whether target sugars achieved within available season particularly cool regions where growing season limited, duration of peak affecting total gain with extended peak beneficial for achieving high sugars while abbreviated peak may limit final levels, coordination between sugar peak and acid decline affecting final balance with simultaneous peaks producing rapid compositional change requiring careful harvest timing, and spatial variation in peak timing across vineyard creating maturity heterogeneity requiring zone-specific management or selective harvest. Harvest timing considerations recognize peak accumulation signals approaching maturity requiring increased monitoring, weekly sampling during peak critical for tracking rates and projecting harvest timing 1-2 weeks advance, recognition that conclusion of peak typically indicates 2-4 weeks to harvest as deceleration begins, and integration of peak sugar accumulation with phenolic and flavor maturity particularly reds where tannin development must synchronize with sugar levels. Viticultural management during peak includes maintaining optimal environmental conditions through irrigation supporting phloem function, canopy health ensuring continued photosynthesis, disease control preventing disruption, crop load ensuring adequate source capacity, avoiding severe environmental stresses during this critical window as impairment during peak disproportionately affects final sugar levels, and harvest preparation advancing as peak indicates approaching maturity. Metabolic efficiency at peak shows optimized carbon use with high proportion imported hexoses accumulating versus respired, ATP production from combined hexose and acid catabolism adequate for all ripening demands estimated 70-110 μmol ATP g⁻¹ h⁻¹, and energetic balance where intensive metabolism during peak supported without excessive respiratory carbon loss representing efficient ripening strategy.
Peak_Sugar_Accumulation is distinguished from Active_Sugar_Accumulation by representing maximum (3.5-5.5% Brix per week) rather than sustained (2.5-4.0%) rates, from Sugar_Accumulation_Onset by showing established peak rather than initial phase, and from late accumulation by exhibiting maximum rather than declining rates. Duration 10-18 days BBCH 84-85. Accounts for 30-50% total ripening gain despite brief duration. Critical window. Peak_Sugar_Accumulation represents most intensive sweetening phase determining final sugar achievement.
Hexose-Import-Maximum
fastest sweetening period
hexose import maximum
maximale Akkumulations-Phase
maximum accumulation phase
optimal loading phase
optimale Beladungs-Phase
schnellste Süßungs-Periode
Keller M (2020) The Science of Grapevines 3rd Ed pages 223-228 peak accumulation Dai ZW, Ollat N, Gomès E, Decroocq S, Tandonnet JP, Bordenave L, Pieri P, Hilbert G, Kappel C, van Leeuwen C, Vivin P, Delrot S (2011) Ecophysiological, genetic, and molecular causes of variation in grape berry weight and composition Plant Physiol 155:519-532 peak rates and variation Coombe BG, McCarthy MG (2000) Dynamics and environmental control of grape berry growth Australian J Grape Wine Res 6:73-83 peak accumulation dynamics.
Peak Sugar Accumulation
In a Stellenbosch Chenin Blanc vineyard in South Africa, peak sugar import rate was measured during 12 days in late February Year N+1 (BBCH 85) when berry Brix increased from 20.5° to 26.8° (average 4.5% per week), hexose import calculated 135 μmol g⁻¹ h⁻¹, cell wall invertase 112 units/g FW, vacuolar hexose rose from 1180 to 1695 mM, berry fresh weight 1.88 g expanding 11%, under optimal warm dry conditions (daily high 28-31°C, moderate deficit stem ψ -1.15 MPa, healthy canopy supporting maximum photosynthesis).
Peak_Sugar_Import_Rate represents the maximum velocity of hexose uptake into berry mesocarp cells achieved during optimal ripening conditions when all components of the sugar import pathway operate at maximum efficiency creating the fastest sugar accumulation observed during the entire berry development cycle. This peak rate occurs during mid-to-late veraison through early ripening (BBCH 84-85), typically 10-16 days post-sugar accumulation onset, and is characterized by net berry hexose import reaching 90-170 μmol combined glucose and fructose per g fresh weight per hour representing integration of apoplastic delivery (cell wall invertase producing 100-200 μmol hexose g⁻¹ h⁻¹ from phloem sucrose), plasma membrane transport (facilitated diffusion 80-150 μmol g⁻¹ h⁻¹), tonoplast sequestration (60-120 μmol g⁻¹ h⁻¹ into vacuoles), and metabolic utilization (20-40 μmol g⁻¹ h⁻¹ for respiration and biosynthesis), resulting in vacuolar hexose concentration increasing 140-220 mM per week from 900-1200 mM toward final cultivar-specific maxima 1400-2200 mM. Molecular basis of peak rate achievement includes: hexose transporter genes (VvHT1, VvHT2, VvHT3, VvHT5, VvHT6) expressed at absolute maximum levels (100-150 fold above green berry when summed) producing saturating membrane protein concentrations estimated 8-15% of total plasma membrane protein, cell wall invertase at maximum activity 80-140 units/g fresh weight catalyzing sucrose cleavage without substrate limitation, regulatory transcription factors sustaining maximum expression coordinating all pathway components, and hormonal signals (ABA 1600-2800 ng/g, hexose through hexokinase pathway) maintaining ripening program at peak activation. Transport kinetics at peak rate demonstrate: plasma membrane hexose uptake operating near Vmax with glucose transport 75-120 μmol g⁻¹ h⁻¹ (from pre-veraison 3-8 μmol) representing 12-20 fold enhancement, fructose transport 65-110 μmol g⁻¹ h⁻¹ similarly maximized, combined hexose uptake 140-230 μmol g⁻¹ h⁻¹ limited primarily by apoplastic substrate availability and tonoplast clearing capacity rather than membrane transport per se, and Km values unchanged (glucose 1.5-3.0 mM, fructose 5-12 mM) confirming capacity rather than affinity increases underlie rate enhancement. Apoplastic dynamics supporting peak rate show: sucrose delivery from phloem 180-350 mg per berry per day (depending on berry size 1.5-2.5 g) maintaining apoplastic substrate, invertase rapidly cleaving sucrose with turnover time 3-8 minutes preventing accumulation that could feedback-inhibit, resulting apoplastic hexoses 250-500 mM combined creating strong concentration gradient (apoplast 350-500 mM → cytoplasm 60-90 mM = 260-440 mM gradient) driving maximal facilitated diffusion flux, and efficient coupling between cleavage, uptake, and vacuolar sequestration maintaining steady-state apoplastic concentrations without excessive accumulation or depletion. Vacuolar accumulation at peak rate involves: tonoplast transport efficiently clearing cytoplasmic hexoses at 70-130 μmol g⁻¹ h⁻¹ preventing buildup that could reduce plasma membrane uptake gradient, vacuolar concentrations rising 150-220 mM per week during peak representing berry Brix increases 3.2-4.8% per week, osmotic potential declining -0.18 to -0.28 MPa per week from -2.0 to -2.6 MPa toward final -2.6 to -3.4 MPa creating water potential gradient sustaining berry expansion 8-15% during peak period, and glucose:fructose ratio in vacuoles maintaining near unity 0.98-1.04 through balanced transport and any differential metabolism equilibrating. Cellular energy requirements at peak rate inclu enhanced respiration 1.8-3.2 μmol O₂ g⁻¹ h⁻¹ supporting ATP synthesis for membrane pumps and metabolism, respiratory quotient >1.3-1.6 indicating substrate-level and fermentative metabolism dominate particularly in berry interior, ATP turnover 60-110 μmol g⁻¹ h⁻¹ with 25-35% allocated to transport-associated processes (H⁺-ATPases maintaining electrochemical gradients, biosynthesis supporting transporter protein turnover), and glycolytic flux 30-60 μmol glucose g⁻¹ h⁻¹ metabolizing portion of imported hexoses while 65-75% accumulates in vacuoles. Environmental conditions enabling peak rate achievement require: temperature optimum 26-30°C supporting maximum enzyme activities, membrane fluidity, and metabolic rates producing highest import velocities 4.2-5.0% Brix per week, moderate warmth 23-26°C enabling 3.5-4.5% per week still approaching peak though kinetically slower, cooler conditions 20-23°C reducing to 2.8-3.8% per week through decreased reaction rates, heat stress 32-35°C impairing peak 30-50% reducing to 2.0-3.0% per week through enzyme denaturation and membrane disruption, and extreme heat >35°C potentially preventing peak achievement reducing to <1.5% per week or causing arrest. Light effects operate primarily through source photosynthesis with adequate canopy illumination essential for producing sucrose substrate supporting peak import, shaded canopies potentially limiting peak rate 25-45% despite full berry import capacity through substrate limitation from insufficient photosynthesis, and cluster-level light exposure having minimal direct effects on peak rate once substrate supply adequate though excessive radiation combined with heat can impair through berry temperature stress. Water relations influences show: adequate vine water status essential for phloem transport delivering sucrose substrate with stem water potential -0.6 to -1.2 MPa compatible with peak rates, moderate deficit -1.0 to -1.4 MPa maintaining peak rates and potentially enhancing concentration through reduced berry expansion, severe deficit <-1.5 MPa impairing peak 40-70% through reduced phloem function and general metabolic limitation, and osmotic relationships complex with high berry sugar creating -2.2 to -3.0 MPa vacuolar osmotic potential requiring adequate vine water status to sustain gradient for continued import. Cultivar variation in peak rate characteristics demonstrates: high-Brix cultivars (Gewürztraminer, Sémillon potential 26-30° Brix) achieve peak rates 4.5-5.5% Brix per week over 2-3 weeks importing 110-170 μmol hexose g⁻¹ h⁻¹, moderate varieties (Chardonnay, Cabernet Sauvignon 22-26° Brix) show 3.8-4.8% per week for 2.5-3.5 weeks with 90-140 μmol g⁻¹ h⁻¹, and lower-sugar types (Pinot Noir cool climates 20-24° Brix) display 3.0-4.0% per week for 3-4 weeks importing 70-110 μmol g⁻¹ h⁻¹ reflecting genetic differences in transporter capacity, enzyme levels, and metabolic programming. Temporal dynamics of peak rate show: rate increasing from initial accumulation 1.5-2.5% Brix per week to enhancement 2.5-3.5% per week to peak 3.5-5.0% per week over 8-12 days, peak sustained 10-20 days depending on cultivar and conditions, then declining to late ripening 1.5-3.0% per week and plateau <0.5-1.5% per week as vacuolar concentrations approach equilibrium with phloem supply and feedback regulation limits further uptake. Coordination with ripening completion shows: peak sugar import coincides with final berry softening approaching asymptotic firmness, color development completing in reds with anthocyanins reaching 92-100% final levels, acid decline approaching minimum sustainable levels with malate nearly depleted, and flavor/aroma compound synthesis intensifying creating integrated maturation suitable for harvest timing decisions. Quality implications of peak rate inclu maximum rate achieved and duration sustained critically determining final sugar levels and thus harvest Brix and wine alcohol potential, uniformity of peak rate across berries and clusters affecting maturity synchrony and harvest composition consistency, environmental optimization during peak maximizing final quality through balanced sugar with acid and phenolic development, and stress avoidance during peak essential as impairment during this critical window disproportionately reduces final sugar even if conditions later improve. Viticultural management during peak rate involves: ensuring adequate vine water status supporting phloem function and metabolism without excess diluting berries, maintaining canopy health for continued photosynthesis providing sucrose substrate, avoiding environmental stresses particularly heat and severe drought that can irreversibly impair peak achievement, disease monitoring critical as high-sugar berries increasingly vulnerable to pathogens, and harvest preparation advancing as peak rate indicates approaching maturity requiring readiness for picking when targets achieved. Monitoring peak rate provides harvest forecasting: weekly Brix sampling tracking maximum accumulation rates enabling projection of harvest timing, comparison of observed versus expected peak rates revealing environmental or management limitations requiring attention, spatial variation in peak rates across vineyard identifying heterogeneity affecting harvest decisions, and integration with other maturity parameters ensuring sugar peak synchronized with desired acid, pH, phenolic, and flavor development for optimal wine quality.
Peak_Sugar_Import_Rate is distinguished from Initial_Sugar_Increase by representing maximum (3.5-5.0% Brix/week, 90-170 μmol g⁻¹ h⁻¹) rather than moderate (1.5-2.5% Brix/week) rates, from Rapid_Sugar_Accumulation_Start by showing sustained peak rather than onset, and from sugar plateau by exhibiting maximum rather than declining rates. Duration 10-20 days BBCH 84-85. Temperature optimum 26-30°C critical. All pathway components saturated. Peak_Sugar_Import_Rate represents culminating metabolic state achieving harvest-ready sweetness and determining wine alcohol potential.
Spitzen-Akkumulations-Rate
maximale Hexose-Aufnahme-Geschwindigkeit
maximale Süßungs-Geschwindigkeit
maximum hexose uptake velocity
maximum sweetening velocity
peak accumulation rate
Keller M (2020) The Science of Grapevines 3rd Ed pages 223-228 peak sugar accumulation Dai ZW, Ollat N, Gomès E, Decroocq S, Tandonnet JP, Bordenave L, Pieri P, Hilbert G, Kappel C, van Leeuwen C, Vivin P, Delrot S (2011) Ecophysiological, genetic, and molecular causes of variation in grape berry weight and composition Plant Physiol 155:519-532 rate variation Sweetman C, Sadras VO, Hancock RD, Soole KL, Ford CM (2014) Metabolic effects of elevated temperature on organic acid degradation in ripening Vitis vinifera fruit J Exp Bot 65:5975-5988 environmental effects on rates.
Peak Sugar Import Rate
Cabernet Sauvignon in Napa Valley shows pectin solubilization start during early September Year N+1 at BBCH 83, with water-soluble pectin increasing from 26% to 44% total pectin during initial softening representing 18% increase in soluble fraction, insoluble pectin declining from 74% to 56%, soluble pectin molecular weight 35-85 kDa appearing from insoluble 180-260 kDa fragmentation, uronic acid in soluble fraction increasing from 8.4 to 14.6 mg/g fresh weight, middle lamella pectin particularly solubilized weakening cell adhesion, solubilization correlating inversely with firmness decline (r²=0.89), under warm climate conditions of 24-30°C temperature, typical of this late-season cultivar initiating pectin conversion at veraison
Pectin solubilization start represents the pectin structural change initiation phase during initial berry softening when insoluble cell wall-bound pectin begins converting to water-soluble forms from enzymatic degradation, water-soluble pectin fraction increasing from pre-veraison 20-30% to initial onset 35-50% total pectin, solubilization indicating pectin depolymerization and release from wall matrix, increased soluble pectin weakening cell wall structure and middle lamella integrity, and solubilization onset as pectin degradation indicator initiating at veraison. This pectin conversion initiation phase occurs during late summer to early autumn in late August-early September in temperate regions Year N+1, representing pectin structural change at veraison onset. Solubilization mechanism involves pectin-degrading enzyme actions cleaving pectin chains creating smaller fragments, depolymerized pectin fragments releasing from wall matrix into aqueous phase, solubilized pectin as degradation products, insoluble structural pectin declining as soluble fraction increases, and solubilization as consequence and indicator of enzymatic modification. Insoluble vs soluble pectin fractionation shows insoluble pectin as structural wall-bound polymers extracted by harsh chemical treatments, soluble pectin as fragments released by water or dilute buffer extraction, fractionation distinguishing structural from degraded pectin, and solubility as pectin state indicator. Pre-veraison pectin state shows pectin predominantly insoluble 70-80% structural, soluble fraction minimal 20-30% representing loosely bound or naturally soluble pectin, insoluble pectin providing wall structure and firmness, and pre-veraison state as structural baseline. Solubilization onset shows soluble pectin fraction beginning increase at veraison, typical increase from 20-30% to 35-50% during initial softening representing 15-25% increase in soluble fraction, continued increase through ripening to 60-80% soluble at full ripeness, and onset marking pectin conversion. Soluble pectin characteristics shows smaller molecular weight fragments 20-100 kDa compared to insoluble 100-300 kDa, soluble pectin extractable in water or dilute buffer, fragments from enzymatic cleavage, and soluble forms as degradation products. Insoluble pectin decline shows structural pectin declining from 70-80% to 50-65% during initial softening, continued decline through ripening to 20-40% at ripeness, insoluble pectin loss from conversion to soluble forms, and decline indicating structural loss. Pectin depolymerization shows pectin polymer chain length declining from enzyme cleavage, molecular weight of both fractions declining with soluble pectin arising from insoluble, degree of polymerization decreasing, and depolymerization enabling solubilization. Molecular weight dynamics shows insoluble pectin molecular weight declining from 150-300 kDa pre-veraison to 100-200 kDa onset to 50-150 kDa ripening, soluble pectin appearing as smaller fragments 20-100 kDa and increasing in amount, and molecular weight as degradation measure. Pectin release from walls shows depolymerized pectin fragments detaching from wall matrix, fragments diffusing into apoplast and cell contents, release weakening wall structural integrity, and release as pectin mobilization. Cell wall structural integrity loss shows wall pectin network fragmenting from depolymerization and solubilization, network disruption reducing wall strength, structural integrity loss reducing firmness contribution, and integrity loss as softening mechanism. Middle lamella pectin solubilization shows middle lamella pectin particularly susceptible to degradation and solubilization, middle lamella holding adjacent cells becoming fragmented, middle lamella loss creating cell-cell adhesion weakening, and middle lamella as critical solubilization target. Cell-cell adhesion weakening shows solubilization of middle lamella pectin reducing intercellular bonding, cells becoming less firmly attached from adhesive pectin loss, tissue coherence declining, and adhesion loss contributing to softness. Pectin compositional changes shows not only solubility changes but also pectin fine structure possibly changing, degree of methylation declining from PME action, galacturonic acid content possibly changing, and compositional shifts accompanying solubilization. Uronic acid content shows total uronic acid as pectin measure, uronic acid in soluble fraction increasing while insoluble decreases, total uronic acid relatively stable though distribution shifts, and uronic acid as pectin quantification. Pectin extraction methods shows hot water extraction for soluble pectin, chelator EDTA or CDTA extraction for calcium-bound pectin, dilute acid or alkali for insoluble pectin, sequential extraction characterizing fractions, and extraction as analytical approach. Solubilization measurement shows spectrophotometric uronic acid determination quantifying soluble and insoluble fractions, ratio calculation showing solubilization degree, size-exclusion chromatography showing molecular weight distributions, and measurements characterizing solubilization. Solubilization timing shows onset at veraison initiation coordinating with enzyme activation, progressive solubilization during initial softening days, continued increase through ripening weeks, and timing as degradation progression. Solubilization correlation with firmness shows strong inverse correlation between soluble pectin percentage and berry firmness, increasing solubilization coinciding with firmness decline, correlation supporting solubilization as softening mechanism, and relationship as functional connection. Environmental effects on solubilization include temperature affecting enzyme activities and solubilization rate, warm conditions promoting faster pectin conversion, cool conditions slowing solubilization, and temperature as rate factor. Temperature effects shows enzyme-driven solubilization temperature-dependent, optimal ranges promoting rapid conversion, and temperature affecting dynamics. The process represents second aspect of initial berry softening establishing pectin conversion, following cell wall modification onset and concurrent with turgor reduction initiation. Duration solubilization initiating over initial softening days at veraison. Regional timing shows pectin solubilization start during late August-early September at veraison onset. Viticultural significance includes representing pectin degradation indicator documenting cell wall modification, correlating strongly with firmness decline, demonstrating structural to soluble conversion as softening basis, and providing analytical marker for softening assessment. Research applications include analyzing pectin solubilization mechanisms and kinetics, investigating pectin structural changes and degradation products, evaluating solubilization-firmness relationships, quantifying pectin fractions and molecular weights, and assessing pectin solubilization as softening indicator and mechanism.
Pectin_Solubilization_Start is second aspect of Initial_Berry_Softening, distinguished by pectin conversion. Represents initial softening days at veraison. Water-soluble pectin 20-30% → 35-50% (15-25% increase), insoluble 70-80% → 50-65%. Soluble pectin 20-100 kDa from insoluble 100-300 kDa fragmentation. Middle lamella pectin particularly affected. Cell-cell adhesion weakening. Strong inverse correlation with firmness. Follows Cell_Wall_Modification_Onset. Concurrent with Turgor_Reduction_Initiation. Pectin degradation indicator.
Pektin-Freisetzung
Pektin-Solubilisierungs-Start
Pektin-Umwandlung
lösliches Pektin-Anstieg
pectin conversion
pectin release
soluble pectin increase
structural pectin loss
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 478-482
Pectin Solubilization Start
Cabernet Sauvignon pedicel at harvest maturity (BBCH 89) measures 5-7 mm length, 1.2-1.8 mm diameter, contains 3-4 vascular bundles with xylem vessels 15-40 μm diameter, supports 1.2-1.8 g berry mass, shows brown lignified appearance, and exhibits well-developed abscission zone facilitating clean berry separation with 0.8-1.5 N detachment force
A pedicel is the short stem structure that connects an individual berry to the rachis of the inflorescence in Vitis vinifera, serving as the structural support and vascular conduit providing water, nutrients, and photosynthates to the developing and ripening berry while also facilitating eventual berry separation through abscission zone activation. The pedicel develops from the flower pedicel that originally supported the flower during anthesis, undergoing structural modifications during fruit set and berry development. Pedicel length in Vitis vinifera varies considerably by cultivar and cluster architecture= typically 3-8 mm in compact-cluster varieties (e.g., Pinot Noir, Grenache) to 8-15 mm in loose-cluster varieties (e.g., Syrah, Nebbiolo), with pedicel length influencing cluster compactness and disease susceptibility (longer pedicels create looser clusters with better air circulation reducing Botrytis risk). Pedicel anatomy includes= outer epidermis with cuticle and occasional trichomes, cortex with parenchyma cells providing structural support, vascular bundles (typically 2-4 bundles containing xylem and phloem tissues) running longitudinally through pedicel center, and specialized abscission zone tissues near berry attachment point. Pedicel diameter ranges 1-2 mm depending on berry size and developmental stage, with thickness generally proportional to berry size (larger berries require thicker pedicels for adequate vascular supply and mechanical support). Pedicel coloration transitions during berry development= green during early development (chlorophyll-containing photosynthetic tissue, BBCH 71-79), progressive yellowing or browning during ripening (chlorophyll degradation, lignification, BBCH 81-85), and brown to woody appearance at full maturity (complete lignification, BBCH 87-89). Pedicel functions include= mechanical support of berry weight (supporting 1-3 g berry mass depending on cultivar), vascular transport of water and minerals from rachis to berry (xylem flow particularly critical during Phase I and early Phase II berry growth), phloem transport of photosynthates supporting berry sugar accumulation (particularly during Phase III ripening), and regulated abscission enabling berry separation at harvest or in response to stress. Pedicel characteristics have significant viticultural implications= pedicel strength affects berry retention and resistance to shattering during harvest and transport, pedicel vascular capacity influences berry size potential and ripening uniformity, pedicel abscission zone development affects harvest ease and post-harvest berry drop, and pedicel lignification timing influences cluster architecture and disease susceptibility.
Pedicel emphasizes the complete berry-supporting stem structure and is distinguished from Berry_Pedicel (berry-specific attachment) and Pedicel_Abscission_Zone (separation-specialized region) by encompassing all structural, vascular, and functional aspects of the berry-rachis connection. Critical for understanding cluster architecture and disease susceptibility, harvest characteristics and shattering resistance, berry nutrition and size uniformity, and mechanical harvest efficiency.
Beerenstiel
Berry stalk
Fruchtstiel
Fruit pedicel
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Pratt, C. (1971). Reproductive anatomy in cultivated grapes - a review. American Journal of Enology and Viticulture, 22(2), 92-109.
Pedicel
Syrah pedicel abscission zone at advanced ripening (BBCH 87-89, 23°Brix) shows differentiated separation layer 4-6 cell layers thick positioned 0.3-0.8 mm from berry attachment, accumulated cellulase and polygalacturonase activities increased 8-12 fold compared to mid-ripening, ethylene-responsive gene expression elevated 15-25 fold, and functional separation enabling 0.9-1.3 N berry detachment force
A pedicel abscission zone is a specialized region of cells within the pedicel, typically located near the berry attachment point, where programmed cell separation occurs enabling controlled detachment of the berry from the cluster through coordinated cell wall degradation and tissue separation. The abscission zone in Vitis vinifera pedicels develops during late berry ripening (BBCH 85-89) as part of the normal fruit maturation process, facilitating harvest and representing an evolutionarily conserved mechanism for seed dispersal. Pedicel abscission zone anatomy includes= abscission layer (separation layer) consisting of 2-10 cell layers positioned transversely across the pedicel where actual separation occurs, protective layer (cells adjacent to separation layer that undergo suberization or lignification preventing pathogen entry after separation), reduced vascular tissue in abscission zone (vascular bundles show smaller diameter and reduced conducting capacity through abscission region), and specialized parenchyma cells with distinctive cell wall characteristics (thinner primary walls, reduced lignification, modified pectin composition facilitating enzymatic degradation). Abscission zone development proceeds through characteristic stages= zone specification (determination of abscission zone position during flower or early fruit development, BBCH 61-71), zone differentiation (cells acquire abscission-competent characteristics during fruit development, BBCH 71-81), zone activation (ethylene and other signals induce abscission-associated gene expression during ripening, BBCH 83-87), cell separation (enzymatic cell wall degradation creates separation plane, BBCH 87-89), and post-abscission differentiation (protective layer formation in remaining pedicel stub). Molecular and biochemical processes in abscission zone include= expression of cell wall modifying enzymes (cellulases, polygalacturonases, pectinases hydrolyzing middle lamella and primary wall components), synthesis and secretion of hydrolytic enzymes into cell wall spaces, ethylene signal transduction (ethylene receptors, transcription factors activating abscission-associated genes), hormone interactions (declining auxin removes abscission suppression, increasing ethylene promotes activation), and redox changes (reactive oxygen species may participate in signaling and cell wall modification). Environmental and physiological factors regulating abscission zone activation include= berry ripening stage (advancing maturity promotes activation), ethylene production (ripening-associated ethylene synthesis triggers enzyme expression), auxin levels (high auxin suppresses abscission, declining auxin during ripening permits activation), water status (drought stress can induce premature activation), pathogen infection (Botrytis or other pathogens may trigger defense-related abscission), and temperature (extreme temperatures can induce stress-related activation). Cultivar variation in abscission zone characteristics includes= strength of abscission zone development (some cultivars develop strong zones with easy berry separation, others maintain firm attachment even at overripeness), timing of abscission activation (early versus late in ripening process), and abscission zone position (typically near berry attachment but some variation in precise location).
Pedicel_Abscission_Zone emphasizes the specialized separation region and is distinguished from Active_Abscission_Zone (enzyme activity and separation progression) and Separating_Abscission_Zone (active cell separation process) by encompassing complete abscission zone structure, development, and regulation. Critical for understanding harvest characteristics, controlled berry separation mechanisms, premature fruit drop under stress, and genetic or environmental factors affecting berry retention and shattering.
Ablösungsschicht
Abscission layer
Abszissionszone
Separation zone
Trennungszone
Pedicel Abscission Zone
Pedicel attachment quality in Muscat Hamburg requires careful management for premium table grape production, with optimal attachment (pull force 2.5-3.5 N) allowing clusters to be handled without shatter while enabling clean separation at the consumer's mouth berries with excessive attachment (force above 4.5 N) require scissors cutting that leaves unsightly stem remnants, while loose attachment (force below 1.5 N) causes progressive berry drop during display that reduces shelf appeal and necessitates frequent pack re-working in retail settings.
Pedicel_Attachment_Quality represents the attachment strength quality specifically characterizing the mechanical connection between the grape berry and its pedicel (the individual berry stem connecting each berry to the rachis). This attachment constitutes the critical junction that must withstand handling forces while allowing eventual separation at harvest, making it central to both table grape quality and wine grape harvestability. The berry-pedicel junction involves complex anatomy including the vascular bundle that supplies water and nutrients to the berry, the brush region where vascular strands splay into the flesh, and potentially an abscission zone with specialized cell layers programmed for separation. The anatomical characteristics of this junction vary among cultivars, with some varieties forming distinct abscission zones that enable clean "dry" separation leaving the pedicel cap on the berry, while others lack defined abscission zones resulting in "wet" separation with torn tissue and juice release. Pedicel attachment strength develops during berry growth as vascular connections lignify and strengthen, typically peaking at véraison, then either maintaining or declining depending on cultivar and ripening conditions. Strong pedicel attachment reduces shatter losses during harvest and handling but may cause tearing damage if berries are forcibly pulled rather than cut. Weak pedicel attachment facilitates easy harvest but causes shatter with minimal handling forces. The cap stem condition after separation (dry versus wet, complete versus torn) provides a visual indicator of attachment characteristics and separation mode.
Pedicel_Attachment_Quality specifically addresses the berry-to-pedicel connection, the most frequently assessed attachment relationship in grape quality evaluation. The class complements Berry_Detachment_Resistance_Quality which addresses the same junction from the perspective of separation resistance. Pedicel attachment is assessed after separation by examining the condition of both the berry apex (brush end) and the pedicel tip, with dry scars indicating abscission zone separation and wet or torn tissue indicating forced separation outside the abscission zone. The vascular bundle provides both the hydraulic connection for berry water supply and a significant portion of the mechanical attachment strength.
Beerenstielverbindung
Pedizellfestigkeit
Stielansatzqualität
Traubenstielansatz
berry stem attachment
berry-pedicel junction quality
cap stem attachment
pedicel connection strength
Considine JA and Knox RB (1981) Tissue Origins, Cell Lineages, and Patterns of Cell Division in the Developing Dermal System of the Fruit of Vitis vinifera L. Planta 151(5): 403-412 Gao Y et al. (2019) Transcriptomic Analysis of Grape Berry Abscission Zone. Plant Science 289: 110245
Pedicel Attachment Quality
Pedicel brittleness evaluation for Flame Seedless table grapes reveals detachment force declining from 4.8 N at commercial maturity to 2.1 N after three weeks cold storage at 0°C, with the 56% reduction in attachment strength causing in-package berry drop (shatter) increasing from 2% to 15% over the storage period, motivating pre-storage SO₂ treatment and careful temperature management to slow abscission zone development and maintain pedicel attachment strength above the 3.0 N threshold required for acceptable shelf-life presentation quality.
Pedicel brittleness quality is a brittleness quality that inheres in the individual berry stalks connecting berries to the rachis and represents the susceptibility of pedicels to fracture at or near the berry attachment point, determining the force required for berry detachment and the mode of separation affecting stem retention on harvested fruit. Within the BFO framework, this quality specifically characterizes the fracture behavior of the berry attachment structure, distinguishing pedicel mechanical properties from rachis properties while recognizing the pedicel as the critical junction determining whether berries separate cleanly with stems retained or with tissue damage and juice loss. The ontological significance lies in pedicel brittleness controlling the balance between secure berry attachment during development and facilitated detachment at harvest, where developmental programming creates an abscission zone of weakened tissue that increases brittleness at maturity enabling efficient harvest while premature brittleness causes shatter loss. Pedicel brittleness shows complex spatial and temporal patterns, with the abscission zone at the berry-pedicel junction developing brittleness independently of the mid-pedicel region, and the progression of brittleness from secure attachment through easy-detach to spontaneous abscission defining the harvest window where berries can be removed with acceptable force without excessive shattering. The cultivar variation in pedicel brittleness development is substantial, with tight-cluster varieties often showing persistent pedicel attachment requiring higher detachment force while loose-cluster varieties may develop brittleness enabling detachment with minimal force or even pre-harvest berry drop. The relationship between pedicel brittleness and stem retention affects fruit quality, as rupture through the abscission zone leaves a dry brush scar while fracture through intact pedicel tissue causes juice leakage and oxidation initiation. Measurement approaches include pull-force testing determining detachment force, microscopic assessment of abscission zone development, and mechanical harvest simulation evaluating detachment behavior under realistic dynamic loading. Understanding pedicel brittleness quality enables harvest timing optimization for efficient detachment without damage, prediction of stem retention patterns affecting fruit presentation, and variety characterization for harvest method suitability.
Pedicel_Brittleness_Quality specifically addresses the individual berry attachment structure, complementing Rachis_Brittleness_Quality for the cluster framework. This quality determines detachment behavior and stem retention patterns critical for both wine grape harvest efficiency and table grape presentation quality. The class supports harvest timing optimization and postharvest handling protocol development based on pedicel condition assessment.
Beerenstielchenbrüchigkeit
Beerenstielsprödigkeit
Fruchtstielsprödigkeit
Pedizellfragilität
berry attachment brittleness
berry stalk brittleness
cap stem brittleness
pedicel fragility
Deng Y et al. (2005) Effects of high CO₂ and low O₂ atmospheres on the berry drop of 'Kyoho' grapes. Food Chemistry 100(2): 768-773 Rolle L et al. (2012) Instrumental texture analysis parameters as markers of table-grape and winegrape quality. American Journal of Enology and Viticulture 63(1): 11-20
Pedicel Brittleness Quality
Syrah pedicel brush attachment after mechanical harvest (BBCH 89) measures 1.8-2.5 mm length representing separation through pedicel tissue above abscission zone, contains 3-4 visible vascular bundle traces with lignified xylem vessels 30-40 μm diameter, shows fibrous brown texture from sclerenchyma fibers, and contributes 60-75% of total brush length due to forced separation above normal abscission plane
A pedicel brush attachment is a brush attachment zone specifically emphasizing the portion of brush tissue that originates from pedicel structures including pedicel vascular bundles, pedicel parenchyma, and lignified pedicel tissues, representing the pedicel-derived component of the brush complex. Pedicel brush attachment develops from the terminal region of the pedicel where vascular bundles prepare to enter the berry receptacle, including= terminal pedicel vascular tissues (final 1-3 mm of pedicel vascular bundles before divergence into berry traces), pedicel cortical parenchyma in attachment region, lignified supporting tissues from pedicel structure, and in some cases, portions of abscission zone that remain attached to pedicel rather than berry after separation. Pedicel brush attachment anatomical features include= longitudinal vascular bundles from pedicel (2-4 main bundles containing xylem vessels 20-45 μm diameter and associated phloem), heavily lignified sclerenchyma fibers providing pedicel structural support, cortical parenchyma cells (often compressed or collapsed in mature dried brush), and potential abscission zone remnants if separation occurred partially through abscission layer. Pedicel brush attachment formation depends on separation location= if separation occurs at distal abscission zone boundary (pedicel side of abscission layer), minimal pedicel tissue remains on berry resulting in small pedicel brush attachment component (0.3-0.8 mm), if separation occurs through pedicel tissue above abscission zone, substantial pedicel tissue remains on berry resulting in larger pedicel brush attachment (1-3 mm including pedicel vascular and cortical tissues), if separation occurs at proximal abscission zone boundary (berry side of abscission layer), pedicel brush attachment is minimal consisting mainly of abscission layer remnants (0.1-0.5 mm). Pedicel brush attachment characteristics include= fibrous texture from exposed vascular bundle elements, brown coloration from lignified xylem and sclerenchyma, visible vascular bundle traces in cross-section (appearing as darker spots or rings representing xylem vessel clusters), and relatively uniform diameter along length (contrasting with potentially expanded receptacle-derived brush portions). Pedicel brush attachment significance includes= component of total brush length contributing to brush morphology variation, source of lignified tissue potentially affecting wine sensory characteristics if brushes remain during crushing, indicator of separation location and mechanism (larger pedicel component suggests separation above normal abscission zone), and contribution to cultivar-specific brush characteristics used in ampelographic identification. Factors affecting pedicel brush attachment size and characteristics include= abscission zone position (zones closer to berry reduce pedicel contribution to brush), pedicel lignification extent (highly lignified pedicels produce more persistent woody brush attachments), separation force and mechanism (gentle abscission versus forced tearing), berry maturity at separation (mature abscission zones produce cleaner separation with less pedicel tissue retention), and cultivar-specific pedicel anatomy (pedicel diameter, vascular bundle size and number, lignification patterns).
Pedicel_Brush_Attachment emphasizes pedicel-derived brush components and is distinguished from Brush_Attachment_Zone (complete interface region) and Seed_Brush_Attachment (vascular connection to seed regions) by focus on tissues originating from pedicel structure. Critical for understanding harvest-induced brush variation, separation location effects on brush morphology, potential enological implications of pedicel tissue in must, and ampelographic brush characteristics.
Pedicel-derived brush
Stalk brush remnant
Stiel-Bürste
Stielgeweberest
Stielstück-Ansatz
Pedicel Brush Attachment
Syrah in Rhône Valley shows pedicel elongation during early June Year N+1 at BBCH 57-59 over 2-3 days, with flower pedicels extending from 1.2-1.5 mm to 3.5-4.5 mm at rate of 0.9-1.3 mm per day creating flower spacing of 5-8 mm under warm Mediterranean conditions of 21-26°C temperature, adequate moisture, typical of this moderately loose-clustered mid-to-late season cultivar completing spatial flower separation before anthesis
Pedicel elongation represents the primary individualization mechanism where flower attachment stalks extend rapidly creating spatial separation between flowers, establishing independent flower positioning essential for pollination access and berry development accommodation. This critical separation process occurs during late May-early June in temperate regions Year N+1 as main component of individual flower separation, typically spanning 2-3 days as coordinated pedicel extension creates 2-5 mm final stalk lengths projecting flowers away from branch attachment points. Elongation mechanism involves cell division in pedicel intercalary meristem zones adding cells to pedicel length particularly during early rapid elongation at 24-36 hour cycles, and cell expansion throughout pedicel tissues with cells increasing length 10-30 fold through turgor-driven wall extension creating visible daily growth of 0.5-1.2 mm. Pedicel structure includes central vascular core with xylem vessels differentiating to conduct water at 0.1-0.3 mg per second per flower adequate for flower function and prepared for berry expansion demands, phloem sieve elements forming to enable nutrient import, surrounding parenchyma providing flexibility and storage capacity, and epidermis with cuticle providing protection. Elongation rate shows strong temperature dependence with development approximately doubling per 10°C increase between 15-28°C range, with optimal 20-26°C supporting maximum elongation rate of 0.8-1.2 mm per day completing elongation within 2-3 days. Final pedicel length varies among cultivars affecting flower spacing with long-pedicelled varieties achieving 4-6 mm creating well-separated loose flower arrangement enhancing air circulation, moderate-pedicelled types reaching 3-4 mm producing intermediate spacing, and short-pedicelled varieties attaining 2-3 mm maintaining relatively compact flower clustering. Pedicel positioning establishes flower orientation with most flowers angled upward or laterally exposing stigmas for pollen deposition, with positioning resulting from phototropic and gravitropic responses and possibly mechanical constraints from neighboring structures. Mechanical properties during elongation show flexible herbaceous structure from thin primary cell walls allowing bending and movement, with progressive wall thickening increasing strength though major mechanical development awaits fruit set when pedicel must support berry weight. Vascular maturation during elongation establishes functional transport capacity with vessel differentiation proceeding basipetally from flower toward branch connection, creating continuous water pathway from branch through pedicel to flower supporting turgor and metabolic function. Cell wall composition changes during elongation from thin plastic walls allowing expansion to progressively thickening walls with increased cellulose deposition providing structural strength, though walls remain relatively thin compared to mature fruit-bearing pedicels. Flower bud development continues during pedicel elongation with buds enlarging to 4-5 mm final pre-anthesis size as flower organs complete maturation, calyptra cap progressively loosening preparing for detachment, and reproductive organs achieving functional readiness. Metabolic activity during elongation maintains elevated respiration supporting growth processes, with energy demands met through photosynthetic assimilate import as shoot canopy provides substantial production capacity. Protein synthesis continues at high rates producing structural proteins for expanding cells, enzymes supporting rapid metabolism and wall modification, and preparing enzymes for reproductive processes. Resource requirements during elongation emphasize carbohydrate demand for cell wall synthesis and energy, nitrogen for protein synthesis, and minerals particularly calcium for wall structure and boron critical for upcoming pollen germination. Hormonal regulation involves auxin promoting elongation through effects on cell expansion and wall loosening, declining gibberellins as reproductive transition proceeds, and possible brassinosteroid effects on cell expansion. Environmental conditions critically affect elongation with temperature as primary factor determining rate, adequate moisture essential for turgor supporting expansion, and favorable weather protecting delicate elongating structures from damage. Water stress during elongation impairs development through turgor loss limiting expansion, potentially resulting in shortened pedicels reducing flower spacing affecting pollination and fruit development accommodation. Disease pressure during elongation continues with powdery mildew representing primary concern requiring protective coverage, with elongating pedicels themselves relatively resistant but attached flowers highly susceptible. Developmental coordination shows pedicel elongation proceeding synchronously across cluster flowers supporting uniform spatial arrangement, with good uniformity promoting coordinated anthesis and fruit set. Within-flower organ positioning changes during pedicel elongation as elongating stalk projects flower away from branch, affecting relative positioning of flower structures and preparing spatial relationships for anthesis. The process represents second of four individual flower separation aspects providing primary mechanism for spatial individuation, following bud separation and preceding full flower visibility and spacing completion. Duration typically spans 2-3 days representing main active elongation period, with warm conditions promoting rapid completion while cool conditions extend period. Cultivar variation represents major feature with genetically determined final pedicel length ranging from short 2-3 mm in compact types to long 5-6 mm in loose types fundamentally affecting cluster architecture, elongation rate affecting timing, and elongation uniformity affecting developmental synchrony. Regional timing shows pedicel elongation during late May-early June in warm climates, early June in temperate regions, and early-mid June in cool areas, occurring 1-3 days before anthesis initiation. Viticultural significance includes establishing flower spacing affecting pollination efficiency with adequate spacing improving pollen transfer, creating cluster microenvironment affecting disease pressure with good separation enhancing air circulation, preparing structural support for berry development with pedicel strength affecting fruit attachment, and determining final cluster density affecting fruit quality and harvest characteristics. Research applications include analyzing mechanisms regulating pedicel elongation including hormonal and biomechanical controls, quantifying environmental effects particularly temperature on elongation affecting flower spacing, evaluating genetic variation in pedicel characteristics affecting cluster architecture, and assessing functional relationships between pedicel length and pollination efficiency and fruit quality outcomes.
Pedicel_Elongation is second of four Individual_Flower_Separation subphases, distinguished by active stalk extension creating primary spatial separation. Follows Flower_Bud_Separation. Represents main mechanism of flower individuation. Transitions to Individual_Flower_Visibility and Spacing_Completion. Primary determinant of flower spacing and cluster density. Highly temperature-dependent process. Genetic pedicel length variation fundamentally affects architecture.
Blütenansatzverlängerung
Blütenstielstreckung
Pedizelexpansion
Stielchenverlängerung
flower stalk elongation
pedicel extension
peduncle development
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 355-358 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Pedicel Elongation
The pedigree record for Cabernet Sauvignon extends three generations — Generation 0 (the cultivar itself), Generation 1 parents (Cabernet Franc × Sauvignon Blanc, Bowers and Meredith 1997), Generation 2 grandparents (Cabernet Franc's parents identified as Gros Cabernet × an unknown cultivar related to Hondarribi Beltza by Boursiquot et al. 2009 Sauvignon Blanc's parents identified as Savagnin × Chenin Blanc by Lacombe et al. 2013), and partial Generation 3 great-grandparents (Savagnin's parentage as Pinot × an unidentified cultivar, establishing that Cabernet Sauvignon carries approximately 12.5% Pinot genetic heritage through this great-grandparental contribution) — the multi-generational pedigree reveals that Cabernet Sauvignon's genome derives from at least 6 identified ancestral cultivars spanning the major viticultural regions of France (Bordeaux, Loire, Jura, Burgundy, and the Basque Country), explaining the cultivar's complex genetic background and broad environmental adaptability.
A parentage information record that documents the multi-generational genealogical relationships of a Vitis vinifera cultivar extending beyond the immediate parent-offspring level, assembling documented and inferred parentage data across two or more generations into a pedigree tree or network that reveals the ancestral lineage, breeding history, and genetic heritage contributing to the cultivar's genome.
Pedigree_Record extends parentage information beyond the single-generation parent-offspring relationship into multi-generational genealogical networks that illuminate the deep genetic heritage of cultivars. The reconstruction of grapevine pedigrees has been enabled by the progressive molecular identification of parentage relationships across hundreds of cultivars, which when assembled reveal interconnected genealogical networks of remarkable complexity: the most extensively documented network centers on the Pinot-Gouais Blanc parent pair, whose at least 75 documented offspring (including Chardonnay, Gamay, Aligoté, and Riesling — the latter through the intermediate Gouais Blanc × Savagnin cross) connect the major cultivar families of northeastern France, Alsace, Germany, and Switzerland into a single pedigree network spanning 500-1,000 years of spontaneous and deliberate hybridization. Pedigree records are particularly important for modern bred cultivars (PIWI disease-resistant varieties, table grape hybrids) where multi-generational crossing has combined germplasm from multiple Vitis species — the pedigree documents the proportion of V. vinifera, V. rupestris, V. lincecumii, V. labrusca, and other species ancestry that determines the cultivar's wine quality potential and disease resistance profile.
Sortengenealogisches Register
Stammbaum-Eintrag
cultivar ancestry documentation
genealogical pedigree
mehrgenerationaler Abstammungsnachweis
multi-generational lineage record
Lacombe T et al. (2013) Large-scale parentage analysis in an extended set of grapevine cultivars, Theoretical and Applied Genetics 126: 401-414 Boursiquot JM et al. (2009) Parentage of Merlot and related winegrape cultivars of southwestern France, Australian Journal of Grape and Wine Research 15: 144-150 Vouillamoz JF and Grando MS (2006) Genealogy of wine grape cultivars, Heredity 97: 102-110 Myles S et al. (2011) Genetic structure and domestication history of the grape, Proceedings of the National Academy of Sciences 108: 3530-3535
Pedigree Record
The peduncle description of Cabernet Sauvignon documents a medium-length peduncle (20-35 mm from shoot insertion to the first rachis branch) with strong lignification at harvest — the peduncle develops a woody, dark brown exterior with rigid mechanical properties that resist bending and create a secure attachment of the cluster to the shoot, requiring substantial force (15-25 N) for detachment during hand harvest and necessitating the use of secateurs or harvest knives for clean separation. This strongly lignified peduncle contrasts with the herbaceous, green, flexible peduncle of Pinot Noir (which remains partially unlignified at harvest, bending easily and detaching with moderate hand pressure of 8-15 N), and the very short, heavily lignified peduncle of Mourvèdre (15-20 mm, so rigid and strongly attached that mechanical harvest detachment forces can damage the rachis before separating the peduncle from the shoot).
A cluster description component that records the morphological characteristics of the peduncle — the stalk connecting the cluster to the shoot at the node of insertion — documenting length, diameter, degree of lignification (woody versus herbaceous tissue at harvest), flexibility, and the ease of separation from the shoot, with peduncle properties influencing the mechanical harvestability of the cluster, the susceptibility to physical damage during transport, and the visual presentation of table grape clusters.
Peduncle_Description documents a structural character that has gained practical importance with the expansion of mechanical harvesting — the mechanical harvestability of a cultivar depends substantially on the peduncle's mechanical properties, specifically the force required for detachment from the shoot and the location of the fracture zone (clean separation at the abscission zone versus breakage within the peduncle or rachis). Machine harvesters detach clusters through trunk shaking or striking rod oscillation that transmits kinetic energy through the trellis wire and cane to the cluster, with the peduncle serving as the critical link whose mechanical failure releases the cluster: cultivars with moderately lignified peduncles and well-developed abscission zones at the shoot insertion (e.g., Cabernet Sauvignon, Chardonnay, Syrah) are well-suited to mechanical harvest because the peduncle fractures cleanly at the intended separation point, while cultivars with excessively flexible herbaceous peduncles (e.g., some Pinot clones) may absorb kinetic energy without fracturing, requiring higher machine intensity that increases juice loss and material-other-than-grape (MOG) contamination. The peduncle also serves as the last vascular conduit supplying water and nutrients to the developing cluster — peduncle lignification during ripening progressively reduces hydraulic conductivity, contributing to the restriction of berry water uptake in the late ripening period that concentrates berry solutes and may promote desirable flavor development.
Rappen-Stiel-Charakterisierung
Stielgerüst-Morphologie
Traubenstiel-Beschreibung
bunch stem characterization
cluster stalk description
peduncle morphology record
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Trought MCT et al. (2011) Harvesting, in Aitken AG and Hewett EW (eds) Fresh Facts — New Zealand Horticulture, Plant and Food Research
Peduncle Description
Pinot Noir grafted on SO4 rootstock at 24 years showing peeling outer rootstock bark with plates 3-8 cm diameter separating from trunk surface, peeling bark edges curling outward showing 2-4 mm thickness of detaching old cork, exposed fresh bark beneath lighter tan color compared to dark grey-brown weathered outer surface, sloughing pattern typical of mature SO4 with irregular plates, underlying bark intact providing continued protection, normal healthy bark renewal process, bark thickness 5.4 mm with outermost 2.1 mm in various stages of separation
Peeling outer rootstock bark is outer rootstock bark in process of separation and sloughing from trunk surface, characterized by outermost dead cork layers losing cohesion and detaching as plates, strips, or flakes exposing fresher bark beneath, resulting from progressive weathering and oxidation degrading outer bark structure, bark cracking under trunk radial expansion stress as diameter increases while inelastic dead bark cannot accommodate growth, accumulated bark thickness exceeding attachment capacity as successive periderms add layers, and natural renewal process where old deteriorated outer bark sheds while new protective layers form deeper maintaining effective protection, with sloughing patterns including large plates, longitudinal strips, small flakes, or circumferential rings depending on cork structure and rootstock cultivar characteristics. Peeling outer rootstock bark separation mechanisms includes= showing cohesion loss (intercellular connections in old cork weakening through oxidation and degradation), exhibiting crack propagation (small surface cracks extending deeper through continued stress and weathering), demonstrating expansion stress (trunk diameter increase stretching inelastic dead bark beyond breaking point), displaying differential growth (new bark forming beneath while old bark static creating shear stress at interface), and revealing age-related weakening (old outer bark becoming brittle and fragile after years of environmental exposure). Peeling outer rootstock bark appearance includes= showing curling edges (separating bark plates or strips curling outward as they detach), exhibiting gaps beneath (air spaces visible between peeling bark and underlying surface), displaying color contrast (old weathered outer surface darker than fresher exposed bark beneath), revealing layered structure (peeling exposing multiple cork layers and periderms in thick old bark), and demonstrating progressive detachment (bark separation beginning at edges or cracks gradually extending until complete sloughing). Peeling outer rootstock bark sloughing patterns includes= plate sloughing (large flat pieces separating characteristic of some cultivars creating distinctive pattern), strip sloughing (longitudinal bark strips peeling off common in smooth-bark types), flake sloughing (small irregular pieces shedding typical in fine-textured bark varieties), ring sloughing (circumferential bands occasionally separating less common but distinctive), and mosaic sloughing (irregular pattern creating patchy appearance with variable piece sizes and shapes). Peeling outer rootstock bark timing and frequency includes= occurring year-round (bark sloughing possible any season though potentially more common in certain periods), showing seasonal variation (spring bark potentially more active sloughing as cambium resumes activity and trunk expands), exhibiting continuous process (old bark gradually sloughing throughout trunk life maintaining appropriate thickness), demonstrating age correlation (older trunk typically showing more extensive bark sloughing due to thicker bark accumulation), and revealing stress effects (drought, damage, or disease potentially affecting bark sloughing rates and patterns). Peeling outer rootstock bark protective function maintenance includes= exposing fresh bark beneath (sloughing old bark revealing newer protective cork layers maintaining effective waterproofing and protection), demonstrating depth reserve (multiple cork layers ensuring continued protection even as outermost layers shed), showing temporary vulnerability (recently exposed bark potentially more susceptible to damage until surface hardens and weathers), requiring underlying layers (effective sloughing depending on adequate new bark formation beneath preventing exposure of living tissues), and enabling long-term protection (bark renewal through sloughing and replacement maintaining protective function throughout trunk life). Peeling outer rootstock bark in different cultivars includes= showing cultivar-specific patterns (Riparia relatively smooth with minimal peeling, Rupestris showing pronounced plate sloughing, hybrids variable), exhibiting thickness correlation (thick-bark cultivars often showing more dramatic peeling than thin-bark types), demonstrating genetic determination (sloughing patterns characteristic and diagnostic for rootstock variety identification), revealing environmental modulation (same cultivar showing different peeling patterns under different environmental conditions), and displaying age effects (young trunk minimal peeling, old trunk extensive sloughing reflecting bark accumulation). Peeling outer rootstock bark management considerations includes= indicating normal process (bark peeling typically healthy normal renewal not requiring intervention), showing stress signals (excessive or unusual peeling potentially indicating trunk stress, disease, or damage requiring assessment), demonstrating disease risk (peeling bark creating temporary vulnerability and potential pathogen entry points requiring monitoring), revealing underlying damage (sloughing occasionally exposing wounds, cankers, or other trunk problems hidden beneath bark), and requiring protection (recent bark removal sites potentially benefiting from wound treatment in disease-prone situations).
Peeling_Outer_Rootstock_Bark emphasizes separation and sloughing process and is distinguished from Outer_Rootstock_Bark (intact bark) by focus on detachment and renewal. Critical for understanding bark renewal, weathering effects, sloughing patterns, protective function maintenance, cultivar characteristics, and the natural process of old bark shedding while new layers form beneath maintaining trunk protection.
Abblätternde äußere Unterlagenrinde
Abschuppende Rinde
Exfoliating bark
Shedding bark
Sich ablösende Borke
Sloughing outer bark
Peeling Outer Rootstock Bark
Syrah peeling outer trunk bark at 22 years post-planting shows outermost cork layers 1.5-3.0 mm depth actively separating in longitudinal strips 4-12 mm wide and 8-25 cm long, strips curling outward 5-15 mm from trunk surface creating characteristic textured appearance, separation occurring at delamination plane between 4th and 5th successive periderms (oldest exterior layer detaching while 5 inner periderm layers remain intact), peeling revealing lighter tan-brown fresh bark beneath contrasting with weathered grey-brown peeling strips, process accommodating 2.8-4.2 mm annual trunk diameter increase while maintaining 5.5-8.2 mm intact protective outer bark depth beneath peeling layer
Peeling outer trunk bark is outer trunk bark that is actively separating, detaching, or sloughing from the trunk surface in sheets, strips, scales, or plates, representing the natural process of exterior bark layer loss that accommodates trunk expansion while maintaining protection through underlying intact bark layers. Peeling outer trunk bark mechanisms include= inelastic outer cork expansion limits (old suberized cork unable to stretch accommodating trunk circumference increase causing surface cracking), delamination between periderm layers (separation along planes between successive cork cambium products), tangential tension from radial growth (xylem expansion creating outward pressure causing outer bark to split and separate), weathering weakening adhesion (UV radiation, freeze-thaw cycles, moisture changes degrading connections between bark layers), and programmed exterior loss (successive periderm formation inherently creating expendable outer layers designed to slough). Peeling outer trunk bark patterns and types includes= longitudinal strip peeling (vertical ribbons of bark separating parallel to trunk axis), platy or scaly peeling (irregular plates or scales 1-5 cm diameter detaching), circumferential ring peeling (horizontal bands of bark separating around trunk uncommon in grapevine), continuous sheet peeling (large coherent bark sheets curling away from trunk), fragmentary peeling (small pieces and flakes gradually separating), and persistent attachment (partially detached bark remaining loosely attached for months or years before falling). Peeling outer trunk bark longitudinal strip characteristics includes= strips typically 2-15 mm wide and 5-30 cm long (dimensions varying by cultivar and bark thickness), strips curling outward as they separate (cork layer shrinkage causing characteristic curling), strips eventually detaching completely (falling naturally or removed by wind, rain, or mechanical contact), new bark layer visible beneath (revealing fresher lighter-colored bark as outer layer peels), and cultivar-specific strip peeling (some varieties characteristically showing prominent long strip peeling pattern). Peeling outer trunk bark platy or scaly characteristics includes= irregular plates 1-5 cm diameter and 2-8 mm thick (varying by bark development), plates separating at delamination planes between periderms (successive cork layers providing natural separation zones), plates often remaining partially attached (connected at base or edges while separated elsewhere creating textured appearance), plates eventually falling revealing underlying bark (maintaining continuous protection as outer plates shed), and characteristic of certain cultivars (some varieties showing pronounced platy bark development). Peeling outer trunk bark timing and seasonality includes= year-round process (bark peeling occurring throughout year as outer layers reach critical state), accelerated peeling after rapid growth (vigorous growing seasons causing enhanced trunk expansion accelerating subsequent bark separation), increased peeling in spring (bark expansion and sap flow potentially loosening outer layers), dry season peeling (summer desiccation causing outer bark shrinkage and cracking enhancing separation), and freeze-thaw enhancement (winter temperature cycling potentially accelerating bark loosening and peeling). Peeling outer trunk bark depth and extent includes= peeling typically affecting outermost 1-4 mm (oldest one or two exterior cork layers separating while inner outer bark remains intact), partial thickness separation (peeling not reaching living inner bark, maintaining protective covering), progressive layer removal (outermost layer peeling revealing next outer bark layer which eventually also peels continuing cycle), continuous protection maintenance (inner intact outer bark maintaining waterproofing and insulation while exterior peels), and balance with production (peeling rate approximately matching outer bark production rate maintaining relatively stable total bark thickness). Peeling outer trunk bark cultivar variation includes= smooth bark cultivars showing minimal peeling (thin uniform outer bark with limited separation creating relatively smooth trunk surface), moderately peeling cultivars (typical peeling behavior with gradual flaking or strip separation), heavily peeling cultivars (pronounced bark separation with dramatic curling strips or large plates creating very textured appearance), strip-peeling types (characteristic long vertical ribbons), and platy-peeling types (characteristic irregular plate detachment pattern). Peeling outer trunk bark relationship to trunk expansion includes= accommodation mechanism (peeling allowing trunk diameter increase without constraining growth), relieving tangential tension (outer bark separation preventing compression of underlying tissues), enabling continuous growth (peeling permitting decades of trunk expansion without outer bark limiting diameter increase), and coordination with new bark formation (outer layer peeling coordinated with inner periderm production maintaining protection thickness). Peeling outer trunk bark effects on trunk appearance includes= creating textured surface (partially peeled bark producing characteristic three-dimensional trunk texture), exposing color variations (fresh bark beneath peeling layers often lighter tan-brown contrasting with weathered grey-brown exterior), revealing successive bark layers (peeling exposing bark stratigraphy showing multiple cork generations), affecting trunk aesthetic (peeling patterns contributing to distinctive trunk character particularly valued in old vines), and creating habitat (loose bark providing shelter for beneficial insects, spiders, or other organisms). Peeling outer trunk bark and trunk protection includes= continuous protection maintenance (peeling designed to preserve protection, outer layers shedding while intact inner layers maintain barrier function), temporary vulnerability during peeling (brief period when outer bark separating but not yet fallen potentially creating minor gaps requiring rapid inner bark exposure healing), lenticel preservation (gas exchange structures maintained in inner bark as outer bark peels), and importance of inner bark integrity (peeling harmless only if substantial intact outer bark remains beneath, excessive peeling exposing living inner bark indicating inadequate bark development or damage). Peeling outer trunk bark management considerations includes= generally beneficial normal process (peeling accommodating growth and removing damaged weathered exterior), avoiding forced premature peeling (manually removing partially attached bark potentially creating wounds or exposing immature inner layers), monitoring excessive peeling (severe peeling potentially indicating bark damage, disease, or inadequate bark development), and distinguishing from pathological bark loss (normal gradual peeling versus catastrophic bark sloughing from disease or severe injury). Peeling outer trunk bark in old vines includes= progressive increase with age (old trunks showing more pronounced peeling as thick bark with multiple periderms creates more delamination planes), characteristic of ancient vines (60-100+ year trunks often showing dramatic peeling with thick curling strips or large plates), contribution to old vine aesthetic (peeling bark creating visually distinctive gnarly aged trunk appearance), and indicating successful adaptation (extensive peeling in old vines demonstrating successful accommodation of decades of trunk expansion). Peeling outer trunk bark distinction from bark damage includes= normal peeling limited to outer dead bark (separation occurring in dead cork layers external to living tissues), pathological bark loss affecting living bark (disease or severe injury causing living inner bark death and sloughing), normal peeling gradual and progressive (occurring over weeks to months), pathological loss often rapid (disease potentially causing sudden bark loss), and normal peeling revealing intact bark beneath (healthy inner bark visible after normal peeling versus diseased or dead tissue in pathological bark loss).
Peeling_Outer_Trunk_Bark emphasizes active separation process and is distinguished from Outer_Trunk_Bark (static structure), Dead_Outer_Trunk_Bark (cellular status), and Protective_Outer_Trunk_Bark (functional emphasis) by focus on dynamic exterior layer loss. Critical for understanding bark accommodation of trunk expansion, normal bark renewal process, distinction from pathological bark loss, cultivar-specific peeling patterns, old vine bark characteristics, and continuous protection maintenance through coordinated peeling and inner bark preservation.
Abblätternde Rinde
Abschälende Außenrinde
Sich lösende Borke
Sloughing bark
Peeling Outer Trunk Bark
Peonidin-3-O-glucoside typically comprises 5-20% of total anthocyanins in V. vinifera cultivars, with elevated proportions (15-30%) in Primitivo (Zinfandel), Tempranillo, and Pinot Noir. Peonidin formation requires O-methyltransferase (VvOMT) activity on cyanidin, adding a single methoxy group at the 3' position of the B-ring. The mono-methoxylated structure produces λmax of 508-518 nm, intermediate between cyanidin (506-513 nm) and malvidin (520-535 nm). Peonidin shows intermediate stability between the highly labile cyanidin and the more stable malvidin, contributing to red-magenta hues in young wines. The peonidin/cyanidin ratio reflects OMT activity in the di-hydroxylated anthocyanin branch, analogous to the petunidin/delphinidin and malvidin/petunidin ratios in the tri-hydroxylated branch.
An anthocyanin accumulation disposition that is the inherited capacity of a Vitis vinifera plant to accumulate peonidin-based anthocyanins (peonidin-3-O-glucoside and derivatives) through O-methylation of cyanidin, contributing red-magenta hues and representing an intermediate methylation state between cyanidin and malvidin in the anthocyanin profile.
Peonidin accumulation represents the methylated form of the cyanidin pathway, providing an intermediate color and stability profile. The trait depends on both F3'H activity (producing the cyanidin precursor) and OMT activity (methylating to peonidin). Cultivars with elevated peonidin often show elevated cyanidin as well, reflecting generally high F3'H pathway flux. The peonidin proportion is used alongside other anthocyanin ratios for cultivar identification and wine authentication.
Peonidin-Akkumulation
Peonidin-Einlagerung
Peonidingehalt
mono-methoxylated anthocyanin
peonidin content trait
peonidin-type pigmentation
Fournier-Level A et al. (2011) Quantitative trait loci analysis of anthocyanin content in Vitis vinifera L. grape berries. Journal of Experimental Botany 62:1043-1051 De Rosso M et al. (2012) Changes in grape polyphenols (V. vinifera L.) as a consequence of post-harvest withering by high-resolution mass spectrometry. Postharvest Biology and Technology 67:102-109
Peonidin Accumulation Disposition
Peonidin content quality analysis in Nebbiolo from Barolo reveals elevated proportions (18-28% of total anthocyanins) compared to most international varieties (5-12%), contributing to the distinctive garnet-orange color that characterizes aged Barolo despite the variety's remarkably low total anthocyanin concentration (150-350 mg/kg) this unusual pigment profile, combined with high acidity and tannin, produces wines that develop complex brick-orange hues earlier than deeply colored varieties while maintaining structural integrity for decades of cellaring.
Peonidin_Content_Quality represents the chemical quality characterizing the concentration of peonidin-based anthocyanins in grape berry skins, capturing the methylated derivative of cyanidin that contributes to red-magenta coloration. Peonidin (3'-methoxy-4',5,7-trihydroxyflavylium) is formed by O-methylation of cyanidin at the 3' position, catalyzed by anthocyanin O-methyltransferase (AOMT). This single methoxy group distinguishes peonidin from the unmethylated cyanidin while differentiating it from the dimethoxylated malvidin. The methylation provides intermediate stability between the relatively unstable cyanidin and the highly stable malvidin. Peonidin-based anthocyanins typically comprise 5-15% of total anthocyanins in most red Vitis vinifera cultivars, representing a consistent but minority contribution to the overall pigment pool. The peonidin proportion relative to cyanidin reflects AOMT activity, while the combined peonidin plus cyanidin proportion (dihydroxylated pathway products) relative to malvidin plus petunidin plus delphinidin (trihydroxylated pathway products) indicates F3'H versus F3'5'H pathway balance. Some cultivars show notably elevated peonidin content Nebbiolo, the variety of Barolo and Barbaresco, produces relatively high peonidin proportions (15-25% of total anthocyanins) that contribute to its characteristically garnet-ruby color distinct from more deeply colored varieties. Peonidin-3-glucoside and its acylated forms are quantified by HPLC with identification based on retention time and spectral characteristics.
Peonidin_Content_Quality captures the monomethoxylated cyanidin derivative that represents intermediate O-methylation status. The biosynthetic relationship cyanidin → peonidin parallels delphinidin → petunidin → malvidin, with methyltransferase activity controlling the conversion. Peonidin contributes red-magenta hues intermediate between cyanidin (orange-red) and malvidin (blue-red). The methylation provides greater stability than unmethylated cyanidin while producing color qualities distinct from fully methylated malvidin. Peonidin proportion contributes to varietal fingerprinting and affects color evolution during wine aging.
Peonidin-3-glucosid-Konzentration
Peonidinpigmentqualität
Pn-3-glc content
monomethoxylated anthocyanin concentration
monomethoxylierter Anthocyangehalt
peonidin pigment quality
peonidin-3-glucoside content
Mattivi F et al. (2006) Metabolite Profiling of Grape: Flavonols and Anthocyanins. Journal of Agricultural and Food Chemistry 54(20): 7692-7702 Fournier-Level A et al. (2011) Quantitative Genetic Bases of Anthocyanin Variation in Grape (Vitis vinifera L. ssp. sativa) Berry: A Quantitative Trait Locus to Quantitative Trait Nucleotide Integrated Study. Genetics 189(4): 1427-1441
Peonidin Content Quality
Pepper aroma quantification comparing Côte-Rôtie Syrah (rotundone 95 ng/L, pepper intensity 7.8 on 10-point scale) with Barossa Shiraz (rotundone 28 ng/L, pepper intensity 4.2) reveals the 3.4-fold concentration difference underlying the stylistic distinction between Northern Rhône's pepper-accented elegance and Barossa's fruit-dominated power, while Grampians Shiraz (rotundone 120 ng/L, pepper 8.5) demonstrates that Australian terroir can achieve Rhône-like pepper when cool-climate sites are selected, with pepper intensity serving as a climate and site quality indicator in this variety.
Pepper aroma quality is a spicy aroma quality that represents olfactory perception evoking Piper nigrum (black pepper) and related peppercorn-associated scents, characterizing the distinctive pungent, spicy aromatic impression that serves as a defining varietal marker for Syrah and certain other grape varieties containing the sesquiterpene rotundone. Within the BFO framework, this quality specifically addresses the pepper subcategory of spicy aroma, distinguished by its grape-derived origin and strong variety-specific associations that enable varietal identification from this single aromatic attribute. The ontological significance lies in pepper aroma representing one of the most chemically defined aroma-variety relationships in wine, where the perception is primarily attributable to rotundone, a bicyclic sesquiterpene discovered in wine in 2008 that provides black pepper character at extremely low concentrations (detection threshold approximately 16 ng/L in red wine, 8 ng/L in white wine). The compositional basis centers almost exclusively on rotundone, which was first identified as the key impact compound responsible for the peppery character of Shiraz and subsequently found responsible for pepper notes in Grüner Veltliner, Schioppettino, Vespolina, and other varieties, with concentrations in wine ranging from undetectable to over 200 ng/L depending on variety, climate, and vintage. The variety distribution of pepper capacity is genetically determined, with Syrah/Shiraz showing consistent rotundone production while many varieties (Cabernet Sauvignon, Merlot, Pinot Noir) lack significant rotundone even when grown in conditions favoring its accumulation, though detection is complicated by individual variation in rotundone sensitivity with approximately 20% of population unable to perceive rotundone at typical wine concentrations. The climate influence on pepper character shows cool-climate and cool-vintage enhancement of rotundone accumulation, with Australian cool-climate Shiraz (Grampians, Yarra Valley) showing more pronounced pepper than warm-region examples, and European Syrah expressing more pepper than most New World examples. Understanding pepper aroma quality enables varietal identification for rotundone-producing varieties, climate and vintage assessment from pepper intensity, and quality evaluation incorporating pepper as a valued complexity element in appropriate varieties.
Pepper_Aroma_Quality addresses the specifically black pepper subcategory of spicy aroma, with an unusually direct relationship to a single compound (rotundone). This quality is grape-derived rather than oak-derived, distinguishing it from sweet spice characters, and shows strong variety specificity. Individual variation in rotundone perception (specific anosmia affecting ~20% of population) complicates sensory assessment and explains some disagreement about pepper intensity among tasters.
Pfefferaroma
Pfeffernote
Rotundon-Charakter
black pepper aroma
peppercorn notes
peppery scent
rotundone character
schwarzer Pfefferduft
Wood C et al. (2008) From wine to pepper: rotundone, an obscure sesquiterpene, is a potent spicy aroma compound. Journal of Agricultural and Food Chemistry 56(10): 3738-3744 Siebert TE et al. (2008) Development of a quantitative method for the determination of rotundone, the pepper aroma compound, in wine and grapes. Journal of Agricultural and Food Chemistry 56(10): 3745-3748
Pepper Aroma Quality
Perceived acidity intensity measurement comparing cool-climate Chablis (7.1 g/L TA, pH 3.15) and warm-climate California Chardonnay (5.8 g/L TA, pH 3.45) using a trained panel reveals intensity ratings of 7.2 versus 4.8 on a 10-point scale, with the 50% higher perceived intensity in Chablis exceeding the 22% difference in titratable acidity, demonstrating the additional contribution of lower pH and absence of malolactic fermentation to perceived acidity, and explaining the stylistic distinction between these regional expressions.
Perceived acidity intensity quality is a sourness quality that represents the magnitude of sour taste sensation at its maximum perception during oral evaluation, characterizing the peak sourness level that determines the dominant impression of acid contribution to overall taste balance. Within the BFO framework, this quality specifically quantifies the intensity dimension of sourness perception at its strongest point, distinguishing intensity from qualitative character (sharp versus soft) and temporal aspects (persistence), providing the primary parameter for acidity assessment in sensory evaluation protocols. The ontological significance lies in perceived acidity intensity serving as the principal sourness descriptor determining whether grapes or wine are perceived as flat, balanced, tart, or aggressively acidic, strongly influencing quality perception where insufficient intensity produces flabby wines while excessive intensity creates unbalanced harshness. The psychophysical relationship between acid concentration and perceived intensity follows a power function with exponent approximately 0.85 for citric acid in simple solutions, though the relationship is substantially modified by matrix effects in wine where sweetness suppresses perceived acidity and tannins may enhance it. The factors modulating perceived acidity intensity include temperature (higher perceived intensity at warmer temperatures, relevant to wine service temperature), sugar content (sweetness suppression reducing apparent acidity), alcohol level (complex interactions affecting acid perception), and carbonation (enhanced acid perception in sparkling wines). In wine style classification, perceived acidity intensity distinguishes between soft, medium, and high-acid styles, with regional typicity often defined by characteristic acidity levels reflecting climate influence on grape acid composition. Understanding perceived acidity intensity enables calibration of sensory assessment protocols, prediction of acidity perception from compositional analysis accounting for matrix effects, and winemaking adjustment to achieve target acidity perception for specific wine styles.
Perceived_Acidity_Intensity_Quality addresses the magnitude dimension of sourness at peak perception, complementing Acidity_Character_Quality (qualitative type) and Acidity_Persistence_Quality (duration). This quality correlates with but is not identical to titratable acidity or pH due to complex matrix interactions. Measurement methodology (scale type, reference standards, sample temperature) significantly affects reported values and should be standardized for cross-study comparison.
Säuregrad
Säureintensität
Säurestärke
acid perception level
acidity magnitude
sour taste intensity
tartness strength
wahrgenommene Säure
Sowalsky RA, Noble AC (1998) Comparison of the effects of concentration, pH and anion species on astringency and sourness of organic acids. Chemical Senses 23(3): 343-349 Fischer U, Noble AC (1994) The effect of ethanol, catechin concentration, and pH on sourness and bitterness of wine. American Journal of Enology and Viticulture 45(1): 6-10
Perceived Acidity Intensity Quality
Perceived bitterness intensity measurement in Sangiovese wines subjected to different maceration protocols reveals intensity of 5.2 on a 10-point scale with 14-day skin contact versus 3.1 with 7-day maceration, with HPLC analysis showing catechin concentrations of 42 versus 28 mg/L accounting for the difference, enabling optimization of extraction duration to achieve the moderate bitterness (target 3.5-4.0) that complements the variety's natural acidity without creating the harsh finish associated with excessive bitter extraction.
Perceived bitterness intensity quality is a bitterness quality that represents the magnitude of bitter taste sensation at its maximum perception during oral evaluation, characterizing the peak bitterness level that determines the impression of bitter contribution to overall taste balance and quality perception. Within the BFO framework, this quality specifically quantifies the intensity dimension of bitterness perception at its strongest point, distinguishing intensity from spatial localization and temporal aspects (persistence), providing the primary parameter for bitterness assessment in sensory evaluation of grapes and wine. The ontological significance lies in perceived bitterness intensity determining whether the bitter component contributes positively to complexity or negatively as a fault, with low to moderate intensity adding structure and interest while high intensity creates unpleasant harsh or medicinal character associated with quality defects. The psychophysical relationship between bitter compound concentration and perceived intensity shows substantial variation among compounds and considerable individual differences related to genetic variation in T2R receptors, with the threshold and suprathreshold intensity functions differing substantially among bitter compounds present in wine. The factors modulating perceived bitterness intensity include temperature (increased perception at warmer temperatures), sweetness (suppression of bitterness by sugar), acidity (enhancement of bitter perception by acid), and alcohol (complex interactions with ethanol potentially suppressing or enhancing bitterness depending on concentration and compound). In wine quality assessment, bitterness intensity is evaluated alongside astringency, with trained panels distinguishing these co-occurring sensations while consumers often conflate them in overall mouthfeel assessment. Understanding perceived bitterness intensity enables calibration of sensory protocols distinguishing bitter from astringent sensations, prediction of bitterness from phenolic composition accounting for individual compounds' bitter potencies, and winemaking optimization to achieve the bitterness level appropriate for the target wine style.
Perceived_Bitterness_Intensity_Quality addresses the magnitude dimension of bitterness at peak perception, complementing Bitterness_Localization_Quality (spatial pattern) and Bitterness_Persistence_Quality (duration). Individual variation in bitter perception due to genetic differences in T2R receptors requires panel calibration and screening for reliable measurement. The quality should be clearly distinguished from astringency intensity in sensory protocols.
Bitterkeitsgrad
Bitterkeitsintensität
Bitterstärke
bitter perception intensity
bitter taste strength
bitterness level
bitterness magnitude
wahrgenommene Bitterkeit
Lesschaeve I, Noble AC (2005) Polyphenols: factors influencing their sensory properties and their effects on food and beverage preferences. American Journal of Clinical Nutrition 81(1): 330S-335S Kallithraka S et al. (1997) Bitterness and astringency of single flavanol enantiomers. Phytochemistry 46(2): 327-332
Perceived Bitterness Intensity Quality
Perceived sweetness intensity measurement in Sauternes wines using a trained panel and 15-point category scale reveals intensity of 11.2±0.8 for wines with 130 g/L residual sugar and 6.5 g/L total acidity compared to 9.8±0.9 for wines with the same sugar but 8.2 g/L acidity, demonstrating the 1.4-point intensity reduction (12%) from the 26% acidity increase, and enabling sweetness-acidity balance optimization for the perceived sweetness intensity of 10.5 considered optimal for this dessert wine style.
Perceived sweetness intensity quality is a sweetness quality that represents the magnitude of sweet taste sensation at its maximum perception during oral evaluation, characterizing the peak sweetness level that determines the dominant impression of sweet taste contribution to overall flavor profile. Within the BFO framework, this quality specifically quantifies the intensity dimension of sweetness perception at its strongest point, distinguishing intensity from temporal aspects (onset, persistence) and providing the primary parameter for sweetness matching, difference testing, and intensity scaling in sensory evaluation protocols. The ontological significance lies in sweetness intensity serving as the principal sweetness descriptor in most sensory evaluation contexts, determining whether grapes or wine are perceived as sweet, off-dry, or dry, and strongly influencing hedonic response where intermediate intensities are often preferred over extremes. The psychophysical relationship between sugar concentration and perceived sweetness intensity follows a power function (Stevens' law) with exponent approximately 1.3 for sucrose, meaning perceived intensity increases faster than concentration at higher levels, with implications for achieving desired sweetness through compositional adjustment. Sweetness intensity is rated using various scaling methods including category scales (1-9 or 1-15 point), line scales (unstructured 100mm), and magnitude estimation (ratio scaling), with method selection affecting the discriminative power and statistical properties of resulting data. The factors modulating sweetness intensity include temperature (reduced intensity at cold temperatures relevant to white wine service), other taste compounds (acid suppression, bitter-sweet interaction), aroma compounds (fruity aromas enhancing sweetness perception through cross-modal interaction), and viscosity (higher viscosity associated with enhanced sweetness). Understanding perceived sweetness intensity enables prediction of sweetness perception from compositional analysis, optimization of sugar-acid balance for target intensity, and calibration of sensory panels for reliable sweetness measurement.
Perceived_Sweetness_Intensity_Quality addresses the magnitude dimension of sweetness at peak perception, complementing Sweetness_Onset_Quality (temporal emergence) and Sweetness_Persistence_Quality (duration). This quality is the primary parameter in most sweetness evaluation protocols and correlates with but is not identical to sugar content due to interaction effects. Measurement methodology (scale type, reference standards) significantly affects reported values and should be specified for cross-study comparison.
Intensität der Süße
Süßegrad
Süßeintensität
Süßstärke
sweet taste intensity
sweetness level
sweetness magnitude
sweetness strength
Stevens SS (1969) Sensory scales of taste intensity. Perception & Psychophysics 6(5): 302-308 Prescott J et al. (2004) Responses to repeated oral irritation by capsaicin, cinnamaldehyde and ethanol in PROP tasters and non-tasters. Chemical Senses 29(5): 393-400
Perceived Sweetness Intensity Quality
Cabernet Sauvignon throughout global wine regions consistently exhibits perfect hermaphrodite flowers with erect stamens extending 0.5-1 mm beyond pistil length, producing 15,000-25,000 viable pollen grains per flower, achieving 48-58% fruit set under optimal bloom conditions (22-28°C, <40% RH), and enabling reliable self-pollination without pollinizer requirements. Chardonnay shows perfect hermaphrodite flower morphology with balanced stamen-pistil proportions, achieving 45-55% fruit set across diverse climates. Gewürztraminer exhibits perfect flowers in morphological sense (both organs present) but functionally female expression with strongly reflex stamens producing minimal viable pollen, requiring cross-pollination from hermaphrodite pollinizers and achieving only 28-38% fruit set even with adequate pollen sources.
A perfect flower is a flower possessing both functional male organs (stamens with viable pollen) and functional female organs (pistil with receptive stigma and viable ovules), representing the morphologically and functionally bisexual condition that characterizes the majority of cultivated Vitis vinifera wine and table grape cultivars and enables self-pollination and reliable fruit set without requiring cross-pollination from separate plants. Perfect flowers in Vitis vinifera exhibit both androecium (five stamens) and gynoecium (bicarpellate pistil) in apparently complete and functional form, though the degree of functional perfection varies along a continuum from fully hermaphrodite (both sex organs fully functional) through functionally female (pistil functional, stamens reflex or producing non-viable pollen) to functionally male (stamens functional, pistil non-receptive or aborted). The perfect flower condition in grapevine includes all flowers possessing both organ types regardless of their relative functionality, distinguishing this broad category from imperfect (unisexual) flowers that completely lack either stamens or pistil. Perfect flower morphology at anthesis typically shows five stamens arranged opposite the five petal positions, with filaments 1-3 mm long supporting anthers 0.4-0.8 mm long containing pollen grains, and a central pistil 1.5-2.5 mm long comprising ovary, short style, and stigma. The spatial arrangement of stamens relative to pistil serves as a critical diagnostic feature= in hermaphrodite flowers stamens are erect and extend equal to or beyond pistil length, in functionally female flowers stamens are strongly reflex (curved downward) and shorter than pistil, and in functionally male flowers stamens are erect but pistil is abnormally short or stigma is non-receptive. Perfect flowers enable autonomous self-pollination through physical proximity of pollen source (anthers) and pollen destination (stigma) within individual flowers, though the efficiency of self-pollination varies with stamen-pistil geometry and environmental conditions during anthesis. The evolutionary and breeding significance of perfect flowers in Vitis vinifera relates to domestication history, as the wild ancestor (V. vinifera ssp. sylvestris) is predominantly dioecious (separate male and female plants) while cultivated types (V. vinifera ssp. vinifera) are predominantly perfect-flowered hermaphrodites, with this transition representing one of the most critical domestication events enabling reliable fruit production from individual vines. Perfect flower cultivars typically exhibit fruit set percentages of 30-60% under favorable conditions (warm, dry weather during bloom), compared to 15-35% in functionally female types requiring cross-pollination and 0% in functionally male types. The perfect flower condition shows moderate heritability with environmental modulation, particularly temperature and nutritional status during flower development affecting final sex expression, with some cultivars exhibiting phenotypic plasticity in stamen position and pollen viability depending on bloom-period conditions.
Perfect_Flower represents the broad category of bisexual flowers in Vitis and must be distinguished from imperfect (unisexual) flowers that lack either stamens or pistil entirely. Within the perfect flower category, critical functional distinctions exist between hermaphrodite (both organs fully functional), functionally female (pistil functional, stamens non-functional), and functionally male (stamens functional, pistil non-functional) subtypes. The terminology can be confusing= 'perfect' in botanical terminology simply means 'possessing both organ types' and does not imply 'fully functional' - many perfect flowers are functionally unisexual due to organ dysfunction despite morphological presence. The perfect flower condition is the derived state in cultivated Vitis vinifera, arising through domestication from dioecious wild ancestors, and represents a critical agronomic advance enabling self-fertility. Modern viticulture predominantly utilizes hermaphrodite perfect flowers, with functionally female types restricted to specialized table grape production and certain wine cultivars requiring pollinizer co-planting.
bisexual flower
hermaphroditic flower
perfekte Blüte
zweigeschlechtige Blüte
zwittrige Blüte
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 99-104 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 92-98 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Battilana, J. et al. (2013). The genetic basis of the conversion of hermaphrodite flowers to male flowers in grapevine. Nature Communications 4=2615.
Perfect Flower
Pinot Noir clone 115 bears a Performance_Clone_Role characterized as quality-focused (small berries, loose clusters, concentrated wine), contrasting with clone 114 characterized as balanced (moderate yield, good quality) and unregistered high-yield selections used for volume production. Chardonnay clone 76 serves in a Performance_Clone_Role with documented quality orientation (small clusters, concentrated must), distinguished in ENTAV evaluation trials from high-yield clones like 75 and 78. Sangiovese clones from the Italian VCR program bear Performance_Clone_Roles with published yield and quality characterizations enabling growers to match clone selection to Chianti versus Brunello production objectives.
A clone role that inheres in a grapevine clone by virtue of its classification according to documented performance characteristics—yield level, quality attributes, vigor, or disease response—established through formal evaluation trials or extensive commercial experience. Performance clone classification enables growers to select clones matching specific production objectives, from high-yield bulk production through balanced commercial clones to low-yield premium quality-focused selections. Performance characterization emerges from replicated field trials (typically 5-15 years) measuring yield, cluster weight, berry size, fruit composition (sugar, acidity, phenolics), and disease incidence, complemented by winemaking trials evaluating wine quality, typicity, and aging potential. Clone performance is not absolute but context-dependent—a clone characterized as "high yield" in Burgundy may perform differently in California, and performance rankings may shift with climate, soil, and management system variations. Performance classification provides decision-support for clone selection, with growers balancing production economics, quality objectives, and site-specific adaptation in their clone choices.
Performance_Clone_Role is subclassed by primary performance characteristic—yield level (High_Yield, Balanced), quality emphasis (Quality), and specific adaptive traits (Disease_Resistant)—reflecting the dimensions by which clones are compared and selected. Performance characterization represents the practical output of clone evaluation programs, translating trial data into decision-relevant categories. The subjectivity inherent in performance classification (what constitutes "high yield" or "quality") is addressed through quantitative trial data registered clones typically have published yield ranges, fruit composition parameters, and wine evaluation results enabling objective comparison. Performance may change with vine age, as young vines often show different clone rankings than mature vineyards, and with climate change altering optimal clone selection across regions.
Leistungsklon
Produktionstyp-Klon
characterized clone
charakterisierter Klon
evaluated clone
performance clone
production-type clone
Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 ENTAV-INRA (2007) Catalogue des Variétés et Clones de Vigne Cultivés en France Regner F (2007) Genetic variation in grapevine clones, Blackwell Anderson K, Aryal NR (2013) Which Winegrape Varieties are Grown Where?, University of Adelaide Press
Performance Clone Role
Performance data for Chardonnay clone 76 (ENTAV) compiled over 12 vintages (1985-1996) at three French trial sites documents average yield of 9.2 t/ha (±1.4), must weight of 12.8% potential alcohol (±0.7), total acidity of 5.8 g/L (±0.9), and micro-vinification scores averaging 15.2/20 for aromatic complexity, establishing it as a balanced clone suitable for both still and sparkling wine production. Performance data for Zweigelt clone Zw-70 across 10 vintages at Klosterneuburg and Eisenstadt records yields of 8-11 t/ha, consistent KMW values of 18-20°, color intensity (OD 520 nm) exceeding the clonal mean by 22%, and Botrytis incidence 30-40% lower than unselected population averages, confirming its suitability for premium red wine production in Burgenland.
Performance data is a clonal selection record entity that documents the quantitative agronomic, oenological, and phytosanitary evaluation results obtained from replicated field trials and laboratory analyses of clonal lines, encompassing yield metrics, fruit quality parameters, disease tolerance assessments, and viticultural adaptation characteristics measured over multiple vintages and, ideally, across multiple evaluation sites, providing the empirical evidence base upon which clone certification decisions and grower planting recommendations are founded.
Performance data constitutes the core decision-support information within clonal selection records, transforming observational field selection into evidence-based clone recommendation. The statistical validity of performance data depends critically on experimental design — replicated randomized block designs with 50-100 vine plots per clone per site are standard in major selection programs, but smaller plot sizes in early evaluation phases may limit statistical power for detecting meaningful clonal differences. The multi-vintage dimension is essential because vintage-dependent genotype-by-environment interactions can cause substantial year-to-year variation in clone ranking, requiring minimum observation periods of 8-12 bearing vintages for reliable characterization. Performance data should ideally distinguish between vine age effects (early bearing versus mature vine performance) and true genotypic differences, as some clones exhibit precocious high yield that moderates with vine age while others show progressive quality improvement over decades. The integration of agronomic, oenological, and phytosanitary performance dimensions into composite clone assessment remains methodologically challenging, as trade-offs between yield, quality, and disease tolerance are common and their relative weighting reflects regional priorities rather than universal optima.
Evaluierungsergebnisse
Klonleistungsdaten
Leistungsdaten
Prüfungsergebnisse
Versuchsdaten
clonal evaluation results
clone performance record
evaluation data
field performance data
trial performance data
Mannini F (2004) Clonal selection in grapevines: interactions between genetic and sanitary strategies. Acta Horticulturae 652: 21-27 Rühl EH et al. (2004) Grapevine clonal selection, evaluation, and management. Acta Horticulturae 652: 29-34 Anderson MM et al. (2008) Viticultural evaluation of French and California Pinot Noir clones. American Journal of Enology and Viticulture 59(2): 188-193
Performance Data
Pergola suitability quality evaluation in Trentino-Alto Adige compared cultivar performance under traditional pergola training characteristic of the region. Schiava demonstrated excellent pergola suitability quality with vigorous growth achieving complete canopy coverage, flexible shoots adapting to horizontal training across the pergola framework, and light-colored fruit benefiting from the shaded fruit zone environment that maintained freshness and prevented sunburn in the warm valley floor conditions while producing characteristic pale ruby wines. Gewürztraminer showed good pergola suitability quality with adequate vigor and aromatic compound development benefiting from the cooler fruit zone temperatures, though requiring enhanced disease management for Botrytis control in the humid pergola microclimate. Lagrein exhibited reduced pergola suitability quality as this deeply-colored cultivar suffered compromised anthocyanin development under the shaded conditions, with conversion to Guyot training on hillside sites achieving 40% higher color density and improved wine structure, demonstrating that pergola suitability quality depends on cultivar quality requirements as well as growth characteristics.
Pergola suitability quality is a trellis adaptation quality that inheres in a grapevine cultivar or clone and represents the degree of compatibility with pergola, tendone, and overhead trellis training systems characterized by horizontal canopy development across elevated support structures positioning fruit below a continuous leaf canopy layer, traditionally employed in warm climate viticulture for table grape and raisin production but also applied to wine grape cultivation in Mediterranean regions, South America, and other areas where overhead training provides advantages for specific production objectives. This quality determines whether a cultivar possesses the vigor capacity, growth habit characteristics, and fruiting responses necessary to achieve productive and qualitative outcomes under the unique environmental conditions created by overhead canopy architecture wherein fruit develops in shaded, protected positions below the photosynthetically active leaf layer. The ontological significance of pergola suitability quality lies in its role enabling productive viticulture under conditions where overhead training provides specific advantages including protection of fruit from excessive solar radiation and heat stress in hot climates, facilitation of cultural operations beneath the elevated canopy, and achievement of high yields through maximum light interception across the horizontal canopy surface. The biological requirements for high pergola suitability quality include sufficient vigor to achieve complete coverage of the overhead trellis surface, growth habit permitting horizontal shoot development across the pergola structure, moderate to vigorous lateral development contributing to canopy closure, appropriate cluster architecture for the shaded, humid fruit zone environment beneath the canopy, and disease resistance characteristics suited to the reduced air circulation and extended wetness duration typical of enclosed pergola fruit zones. The microclimate conditions beneath pergola canopies differ fundamentally from vertical or divided systems, with fruit experiencing reduced light intensity, lower temperature extremes, elevated humidity, and protection from direct solar exposure that may benefit thin-skinned cultivars in hot climates while potentially compromising color development and increasing disease pressure in cooler, humid conditions. Assessment of pergola suitability quality involves evaluation of canopy coverage achievement, fruit zone microclimate characteristics, disease incidence patterns, and fruit composition outcomes reflecting the unique pergola environment.
Pergola_Suitability_Quality is_a Trellis_Adaptation_Quality, representing compatibility with overhead/horizontal canopy systems including pergola, tendone, and parral configurations. This class addresses the specific requirements of overhead training including high vigor for complete coverage, horizontal growth adaptation, and fruit quality under shaded conditions. The unique microclimate with shaded fruit distinguishes pergola from all other major training systems. Most relevant for warm climate viticulture and specific cultivar-quality objectives. Distinguished by the horizontal canopy plane creating fundamentally different fruit zone conditions than vertical or inclined systems.
Laubengang-Eignung
Pergola-Eignung
Tendone-Tauglichkeit
horizontal canopy compatibility
overhead trellis suitability
pergola training fitness
tendone adaptation
Überkopferziehungs-Kompatibilität
Ferraro DO, Riveros VN (2012) Grape training systems for the irrigated desert of South America. In: Reynolds AG (ed) Viticulture and Wine Quality, Woodhead Publishing, pp. 89-115 Intrieri C et al. (2001) Yield and grape quality assessment of training systems for machine harvesting. Acta Horticulturae 562:287-294 Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review. American Journal of Enology and Viticulture 60:251-268
Pergola Suitability Quality
A Trebbiano vine in central Italian tendone training bears a Pergola_Trained_Vine_Role with cordons at 200 cm supporting a horizontal canopy extending 2 m in each direction, creating complete overhead shade valued for table grape production and traditional vineyard aesthetics. A Torrontés vine in Argentine Mendoza parral training serves in a Pergola_Trained_Vine_Role, the overhead system shading fruit from intense high-altitude UV radiation while enabling continued traditional cultivation practices. A Vinho Verde vine in Portuguese latada training bears a Pergola_Trained_Vine_Role with high cordon and spreading canopy, the traditional system adapted to the region's granitic posts and humid climate where canopy elevation improves air circulation.
A training system role that inheres in a grapevine trained to an overhead or semi-overhead structure (pergola), with cordons or canes supported at elevated height (typically 180-250 cm) creating a canopy that extends horizontally above or beside the vine row, shading the area beneath the vine structure. Pergola training encompasses diverse regional systems—Italian tendone and pergoletta, Argentine/Chilean parral, Portuguese latada—all sharing the elevated, spreading canopy architecture that distinguishes pergola from the vertical canopy walls of conventional trellised systems. Pergola systems were developed for hot climates where fruit shading prevents sunburn and excessive temperature, for table grape production where protected fruit development and harvest access are paramount, and for traditional polyculture systems where agricultural use of land beneath vines (vegetables, grains, livestock grazing) was economically important. The elevated structure requires substantial infrastructure investment (posts, overhead wires or lattice, training systems), but may reduce foliar disease pressure through improved air circulation and separation of fruit from ground-level humidity. Pergola-trained vineyards create distinctive landscape aesthetics, with the overhead canopy creating shaded corridors between vine rows.
Pergola_Trained_Vine_Role captures the category of elevated, spreading canopy systems fundamentally different from vertical shoot-positioned approaches dominating contemporary quality wine production. Pergola systems persist for specific applications—table grapes (where protected fruit and extended hang time are valued), hot climates (where shading prevents heat damage), traditional wine regions (where heritage systems maintain cultural continuity), and high-yielding production (where the spreading canopy supports large crop loads). The distinction between pergola and high cordon systems is sometimes unclear true pergola creates overhead canopy extending beyond the vine row, while high cordon creates an elevated fruiting zone with downward shoot growth but not the overhead extension characteristic of pergola. Quality wine production has generally moved away from pergola toward more exposed canopy systems, though climate change may prompt reassessment as heat-related fruit damage increases in traditional quality regions.
Laubengang
Pergola-System
Pergolaerziehung
arbor training
overhead training
pergola training
trellis pergola
Überkopf-Erziehung
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Ferroni G, Scienza A (2014) Italian grape training systems, ISHS Acta Horticulturae 1032: 333-340 Cavagnaro JB et al. (2006) Grapevine training systems in Mendoza, American Journal of Enology and Viticulture 57: 113-122
Pergola Trained Vine Role
Cabernet Sauvignon pericarp at maturity (BBCH 89) shows anatomical organization= outer pericarp (skin) 95-125 μm total thickness containing epidermis with cuticle (22-28 μm), hypodermis with dense anthocyanins (2.8-3.6 mg/g providing deep purple-black color, 35-50 μm), outer mesocarp layers (40-55 μm) middle pericarp comprising 88-91% of berry volume with large parenchyma cells (120-165 μm diameter in central region) containing 24-26% sugars, 5.5-6.8 g/L acids, creating soft sweet flesh inner pericarp (150-220 μm thickness) adjacent to 2-3 seeds with smaller cells (55-85 μm) and elevated tannins (2.5-4 mg/g versus 0.3-0.8 mg/g in middle pericarp), demonstrating radial tissue organization and compositional gradients within pericarp structure. Chardonnay white cultivar pericarp showing white wine characteristics= outer pericarp (80-110 μm) lacking anthocyanins but containing flavonols and aromatic precursors, golden-yellow color from carotenoids middle pericarp (89-92% of volume) with cells 130-170 μm diameter containing 21-24% sugars, 6.5-8.5 g/L acids inner pericarp (120-180 μm) with moderate tannin levels, creating white wine pericarp composition emphasizing acid retention and aromatic potential over color. Syrah pericarp developmental progression shows tissue evolution= Phase I (BBCH 75, week 6) with pericarp containing 2.4 million cells averaging 58 μm diameter, total pericarp volume 0.95 cm³ Phase II veraison (BBCH 81, week 10) with same cell number at 85 μm diameter, pericarp volume 2.1 cm³ maturity (BBCH 89, week 17) with cells 152 μm diameter, pericarp volume 4.8 cm³, demonstrating that pericarp expansion during ripening results from cell enlargement (cells increasing 162% in diameter, 525% in volume) driving 405% pericarp volume increase. Pinot Noir pericarp tannin distribution measured by micro-dissection and spectrophotometry shows outer pericarp layers containing 8-12 mg/g tannins (procyanidins and prodelphinidins), middle pericarp 0.4-0.9 mg/g, inner pericarp 3-6 mg/g, with total pericarp average 1.2-1.8 mg/g reflecting predominance of low-tannin middle pericarp tissue by volume, demonstrating tannin concentration gradients with highest levels in outer and inner pericarp protective/structural layers. Merlot pericarp vascular anatomy showing dorsal bundles (4 major bundles at 90° intervals around berry circumference) running from pedicel to stylar scar through pericarp, with xylem vessels 20-45 μm diameter functional during early development but largely non-functional during ripening, phloem tissue with sieve elements 12-25 μm diameter maintaining function throughout ripening providing sugar and water import, and lateral vascular branches from major bundles distributing throughout pericarp tissue creating vascular network supporting pericarp growth and composition.
A pericarp is the complete fruit wall tissue developing from the ovary wall of the flower, comprising the entire structural and compositional matrix of the berry from outer surface (skin) through flesh to inner locular cavity boundaries, representing the total berry tissue excluding seeds and encompassing all anatomical layers and cell types that create berry structure, composition, and quality characteristics. The pericarp structure in Vitis comprises three principal anatomical regions= (1) outer pericarp (exocarp and outer mesocarp layers) corresponding to berry skin, typically 40-180 μm total thickness at maturity, including epidermis (single cell layer 15-35 μm thick with cuticle 2-8 μm), hypodermis (1-3 cell layers 20-60 μm thick), and outer mesocarp layers (2-6 cell layers 25-90 μm thick), containing stomata, lenticels, pigments (anthocyanins in red/black cultivars concentrated in hypodermal cells), tannins, aromatic compounds, and serving as protective barrier (2) middle pericarp (mesocarp bulk) comprising the majority of berry tissue (typically 85-92% of pericarp volume), consisting of large thin-walled parenchyma cells (80-180 μm diameter at maturity) accumulating sugars, organic acids, water, and minerals, with cell size increasing from outer (smaller cells 60-120 μm) through middle (largest cells 100-180 μm) to inner regions (intermediate cells 70-140 μm), creating the flesh texture and compositional characteristics determining berry and wine quality (3) inner pericarp (innermost mesocarp layers) adjacent to locular cavities and seeds, typically 100-300 μm thick comprising 3-8 cell layers, showing distinct cellular organization with smaller cells (40-90 μm diameter) than middle pericarp, higher tannin content than middle pericarp, and direct contact with developing or mature seeds. Pericarp development follows berry development phases= Phase I (BBCH 71-75, weeks 0-6 post-fruit set) involving rapid cell division in mesocarp creating final cell number (1.5-3.5 million cells total in pericarp), with cells 25-70 μm diameter lag phase (BBCH 77, weeks 6-8) with minimal pericarp growth but seed maturation and metabolic transitions Phase II (BBCH 81-89, weeks 9-18+) involving dramatic cell expansion particularly in middle pericarp (cells expanding from 60-90 μm to 100-180 μm diameter), sugar and water accumulation, and secondary metabolite synthesis creating mature pericarp composition. Pericarp tissue organization shows radial gradients= outer pericarp with highest tannin concentration (procyanidins, prodelphinidins), moderate-to-high anthocyanins (in red cultivars, concentrated in hypodermis), thick cell walls, small cell size middle pericarp with lowest tannin concentration, highest sugar and water content, thin cell walls, largest cell size, creating soft texture and sweet taste inner pericarp with intermediate-to-high tannin concentration (higher than middle, lower than outer typically), intermediate cell size, proximity to seeds. Pericarp vascular system includes dorsal bundles (3-5 major bundles running from pedicel through pericarp to stylar end) and ventral bundles (branches serving berry tissues), with xylem function declining during ripening and phloem becoming predominant import pathway. Pericarp composition at maturity varies by cultivar and conditions= sugars 16-28% (primarily glucose and fructose in approximately equal proportions), organic acids 0.4-1.2% fresh weight (tartaric and malic acids), pH 3.1-4.2, phenolic compounds including anthocyanins (outer pericarp in red cultivars), flavonols (outer pericarp), tannins (outer and inner pericarp higher than middle), aromatic compounds determining varietal character.
Pericarp represents the complete berry wall tissue (excluding seeds) developing from ovary wall and must be distinguished from Berry (which includes pericarp plus seeds) and from individual pericarp layers. Pericarp comprises outer (skin), middle (flesh bulk), and inner (seed-adjacent) regions with distinct anatomical and compositional characteristics. Sub-classification into Outer, Middle, and Inner pericarp reflects the radial tissue organization essential for understanding berry structure, composition, and quality. Understanding pericarp anatomy and development is fundamental for viticulture and enology as pericarp tissues provide sugars, acids, phenolics, aromatics, and structural properties determining berry and wine quality.
Beerenwand
Fruchtwand
Perikarp
berry wall
fruit wall
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 172-245 Harris, J.M. et al. (1968). The Vinifera grape berry= an anatomical, biochemical and ultrastructural study. Botanical Gazette 129=1-23 Cadot, Y. et al. (2006). Analytical and sensory contribution of grape tannins in wine. Journal of Agricultural and Food Chemistry 54=2973-2987.
Pericarp
Chardonnay in Champagne shows pericarp cell proliferation during early-mid July Year N+1 at BBCH 72-73, with mesocarp intensive division generating 1.4-1.9 million flesh cells representing 82-86% total, exocarp surface division producing 180,000-260,000 skin cells through anticlinal divisions, endocarp limited division yielding 60,000-100,000 inner layer cells, coordinated pericarp proliferation establishing three-layered berry wall architecture, under cool maritime conditions of 20-25°C temperature, typical of this early-bursting cultivar
Pericarp cell proliferation represents the mesocarp and exocarp tissue-focused division phase during peak division activity when flesh and skin layers undergo intensive coordinated proliferation, mesocarp accounting for 80-85% total cell generation creating bulk berry volume, exocarp surface divisions maintaining coverage during expansion, and integrated pericarp proliferation establishing three-layered berry wall architecture. This tissue-specific proliferation phase occurs during mid summer in early-mid July in temperate regions Year N+1, proceeding during peak division weeks 2-3 post-fertilization, corresponding to BBCH 72-73 young fruit growth stages. Pericarp proliferation mechanism involves mesocarp flesh tissue intensive division generating majority of berry cells, exocarp skin layer coordinated division maintaining surface integrity, endocarp inner layer adjacent to seeds showing limited division, tissue-specific division rates and patterns creating organized architecture, and overall pericarp proliferation establishing berry wall structure. Mesocarp proliferation dominance shows flesh tissue as primary proliferative compartment, mesocarp cell generation accounting for 80-85% total pericarp cells, intensive division throughout mesocarp volume creating bulk cellularity, and mesocarp as major berry size determinant. Mesocarp division patterns show periclinal divisions predominating in inner regions increasing radial depth, anticlinal divisions in middle regions maintaining proportions, coordinated division creating organized mesocarp architecture, and division patterns establishing tissue organization. Mesocarp cellularity establishment shows typical accumulation to 1.5-3.5 million mesocarp cells by proliferation completion, cell number range reflecting cultivar variation and environmental effects, and mesocarp cellularity as primary berry size determinant accounting for bulk volume. Exocarp proliferation characteristics show surface epidermal and subepidermal layer division, primarily anticlinal divisions perpendicular to surface expanding coverage area, coordinated exocarp division maintaining continuous surface during berry expansion, and exocarp cell generation accounting for 10-15% total pericarp cells. Exocarp division necessity shows surface area increase requirement as berry expands, anticlinal divisions producing new cells expanding coverage, division preventing gaps or cracking from expansion stress, and adequate exocarp proliferation essential for berry integrity. Exocarp architecture from proliferation shows epidermal single cell layer maintained through anticlinal division, subepidermal 4-8 cell layers from combined periclinal and anticlinal divisions, exocarp total thickness 6-12 cell layers representing skin structure, and layer organization from coordinated division patterns. Endocarp limited proliferation shows inner layer adjacent to seeds showing reduced division compared to mesocarp, endocarp accounting for 3-5% total pericarp cell generation, specialized endocarp cells possibly less proliferative, and limited division creating thin 2-4 cell layer endocarp. Tissue proportion maintenance shows coordinated proliferation across layers maintaining relative proportions, mesocarp as bulk tissue 80-85% volume, exocarp as surface layer 8-12% volume, endocarp as inner layer 3-5% volume, and proportions established through tissue-specific division rates. Tissue-specific division regulation shows differential proliferative capacity across layers, possible auxin gradient effects with higher levels near seeds, mechanical factors possibly affecting surface versus internal division, developmental programming establishing tissue-specific patterns, and integrated regulation creating organized architecture. Cell type differentiation during proliferation shows cells maintaining division competence while acquiring tissue identity, early differentiation markers appearing during proliferation, and progressive specialization as division completes. Vascular tissue proliferation shows procambial cells dividing creating vascular bundles, xylem and phloem precursor proliferation, vascular network expansion coordinated with berry growth, and vascular development supporting berry enlargement. Pericarp volume increase from proliferation shows tissue expansion from cell accumulation, volumetric increase correlating with cell number, pericarp representing 85-95% total berry volume at division completion, and volume from proliferation establishing berry size baseline. Environmental effects on pericarp proliferation include temperature affecting proliferation rate across all layers with optimal 24-28°C supporting maximum pericarp cell generation, differential temperature sensitivity possible across tissues though evidence limited, and overall temperature effects on total pericarp proliferation. Auxin effects show hormone gradient possibly affecting mesocarp versus exocarp proliferation, higher auxin near seeds potentially stimulating greater mesocarp division, and overall auxin stimulus coordinating pericarp proliferation. Pericarp proliferation assessment methods include tissue-specific cell counting quantifying layer cellularity, histological analysis showing tissue architecture, division pattern observation documenting periclinal versus anticlinal divisions, and proliferation marker distribution across tissues. The process represents second aspect of peak division activity emphasizing tissue-specific proliferation, following maximum cell number generation and preceding seed compartment division. Duration throughout peak division weeks 2-3 representing coordinated tissue proliferation. Regional timing shows pericarp cell proliferation during early-mid July across climates during peak berry division. Viticultural significance includes representing major cellularity establishment determining berry structure and size, demonstrating coordinated tissue-specific proliferation creating organized architecture, showing mesocarp as dominant proliferative tissue establishing size, and establishing pericarp proliferation as berry developmental foundation. Research applications include analyzing tissue-specific proliferation regulation, investigating division pattern control across layers, evaluating pericarp architecture establishment, quantifying tissue cellularity and proportions, and assessing pericarp proliferation as determinant of berry structure and size.
Pericarp_Cell_Proliferation is second aspect of Peak_Division_Activity, distinguished by tissue-specific proliferation focus. Represents mesocarp flesh (80-85%), exocarp skin (10-15%), and endocarp (3-5%) coordinated proliferation. Mesocarp dominant proliferative tissue determining size. Exocarp anticlinal divisions maintaining surface. Duration peak weeks 2-3. Establishes three-layered pericarp architecture. Followed by Seed_Compartment_Cell_Division. Tissue-specific division patterns creating organized structure.
Beerenwand-Proliferation
Fleisch- und Haut-Zellenerzeugung
Mesokarp- und Exokarp-Teilung
Perikarp-Zellproliferation
berry wall proliferation
flesh and skin cell generation
mesocarp and exocarp division
pericarp tissue proliferation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 436-438
Pericarp Cell Proliferation
Merlot in Bordeaux shows periclinal cell division during early-mid July Year N+1 at BBCH 71-72, with division planes aligned parallel to berry surface, preprophase bands predicting periclinal orientation, radial daughter cell positioning increasing mesocarp depth from 0.8 mm to 3.2 mm over weeks 1-4, periclinal divisions representing 62-74% total divisions in inner mesocarp, under oceanic conditions of 20-26°C temperature, typical of this mid-season cultivar showing coordinated periclinal proliferation contributing to berry volumetric expansion
Periclinal cell division represents the radially-oriented division pattern during active berry cell division when division planes align parallel to berry surface, daughter cells position radially increasing tissue thickness, and coordinated periclinal divisions expand pericarp depth creating three-dimensional berry growth, predominating in inner mesocarp regions contributing to radial berry expansion. This directional division pattern occurs during early summer cell division in late June-July in temperate regions Year N+1, proceeding throughout active division weeks 1-3 as major division mode, corresponding to BBCH 71-72 young fruit growth stages. Periclinal division mechanism involves division plane orientation control through cytoskeletal organization, preprophase band microtubules establishing division plane parallel to surface before mitosis, spindle alignment perpendicular to surface directing chromosome segregation, cytokinesis producing cell plate parallel to epidermis, and daughter cell radial positioning increasing tissue depth. Division plane determination shows preprophase band formation during late G2 phase, cortical microtubule array marking future division site on plasma membrane, division plane prediction from preprophase band position, and band disappearance before mitosis while plane memory persists. Preprophase band characteristics show microtubule ring encircling cell at division site, band width 2-5 micrometers predicting cell plate insertion zone, band position determining periclinal versus anticlinal orientation, and band as spatial memory system controlling division geometry. Spindle orientation control shows spindle axis alignment perpendicular to berry surface and parallel to radial axis, spindle positioning ensuring chromosome segregation produces radial daughter positioning, astral microtubules contacting cortical sites marked by preprophase band, and motor proteins positioning spindle through force generation on astral microtubules. Cytokinesis plane alignment shows cell plate forming parallel to preprophase band position, phragmoplast guidance by former band site, progressive cell plate expansion creating new wall parallel to surface, and completed division producing radially arranged daughter cells. Daughter cell positioning shows cells arranged along radial axis from surface toward center, radial arrangement increasing tissue depth, daughter cells both contributing to subsequent radial expansion through additional divisions or enlargement, and coordinated radial positioning creating organized tissue architecture. Periclinal predominance in mesocarp shows inner flesh tissue showing primarily periclinal divisions, radial divisions increasing tissue thickness more than anticlinal divisions increasing area, and periclinal divisions accounting for 60-75% total divisions in inner mesocarp regions. Tissue thickness increase from periclinal divisions shows progressive mesocarp depth expansion, radial growth contributing substantially to volumetric berry increase, thickness increase from 0.5-1 mm at division onset to 2-4 mm by division completion, and radial expansion as major contributor to three-dimensional berry growth. Three-dimensional growth contribution shows periclinal radial divisions combined with anticlinal tangential divisions creating volumetric expansion, radial growth from periclinial divisions, tangential growth from anticlinal divisions, and integrated division pattern producing spherical to ovoid berry morphology. Division plane regulation shows genetic control through division plane orientation genes, cytoskeletal regulatory proteins controlling preprophase band and spindle positioning, auxin possibly affecting division plane orientation though mechanisms unclear, and developmental programming establishing tissue-specific division patterns. Periclinal versus anticlinal division ratios show spatial variation with inner mesocarp 60-75% periclinal, middle mesocarp 50-50%, outer mesocarp 25-40% periclinal, and ratios reflecting position-specific growth requirements. Exocarp division patterns show epidermis and outer layers showing primarily anticlinal divisions maintaining surface coverage, anticlinal divisions enabling tangential expansion accommodating berry enlargement, and anticlinal predominance in surface layers contrasting with mesocarp periclinal pattern. Coordinated division orientation shows tissue-specific patterns creating organized architecture, inner mesocarp periclinal radial growth, outer exocarp anticlinal tangential growth, and coordination maintaining tissue proportions and structural integrity during berry expansion. Cell layer organization from divisions shows radial cell files from repeated periclinal divisions, organized layer structure in mesocarp, and architecture reflecting division history and pattern. Environmental effects on periclinal division include temperature affecting division rate with optimal 24-28°C supporting rapid periclinal proliferation, division plane orientation likely temperature-independent though division frequency affected, and overall favorable conditions maximizing periclinal division contribution to growth. Mechanical stress effects show possible influences on division plane orientation though evidence limited in berries, turgor pressure potentially affecting plane selection, and tissue architecture possibly providing mechanical cues. Periclinal division assessment methods include histological section analysis showing radial cell arrangements, division plane observation during mitosis, daughter cell position tracking, and tissue architecture analysis inferring division patterns. The process represents third aspect of active berry cell division emphasizing radial division orientation, following initial division activation and mitotic activity onset. Duration throughout active division weeks 1-3 as predominant inner mesocarp division mode. Regional timing shows periclinal cell division during late June-July across climates during early berry division phase. Viticultural significance includes representing major division pattern contributing to radial berry expansion and volumetric growth, demonstrating coordinated tissue-specific division orientation, creating organized mesocarp architecture, and contributing to three-dimensional berry morphology development. Research applications include analyzing division plane control mechanisms, investigating cytoskeletal regulation of division orientation, evaluating tissue-specific division patterns, quantifying periclinal versus anticlinal division ratios, and assessing division orientation as determinant of berry shape and architecture.
Periclinal_Cell_Division is third aspect of Active_Berry_Cell_Division, distinguished by surface-parallel division plane orientation. Represents radial tissue growth increasing depth. Predominates in inner mesocarp 60-75% of divisions. Division plane controlled by preprophase band and spindle orientation. Contrasts with anticlinal divisions in outer layers. Duration throughout active division weeks 1-3. Contributes to three-dimensional berry expansion. Creates organized radial cell files. Tissue-specific division pattern.
depth-increasing division
oberflächenparallele Teilung
periklinale Zellteilung
radial cell division
radiale Zellteilung
surface-parallel division
tangential division
tiefenzunehmende Teilung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 434-436
Periclinal Cell Division
Merlot periderm at 15 years post-planting shows complete three-layer structure= phellem 1.4-2.6 mm thick comprising 22-38 radial files of dead rectangular suberized cork cells 25-45 μm dimensions providing waterproof barrier, phellogen single cell layer with initial cells 28-36 μm actively dividing May-September producing 80-85% phellem and 15-20% phelloderm, and phelloderm 0.2-0.4 mm thick with 3-6 cell layers of living parenchyma, represents 4th successive periderm (3 older periderms visible exterior at 2.8, 5.2, and 7.8 mm depths creating total outer bark 8.5-11.2 mm), scattered lenticels 2-4 mm diameter interrupting phellem enabling gas exchange
Periderm is the protective tissue complex in Vitis vinifera trunk comprising three components - phellem (cork cells) produced centrifugally, phellogen (cork cambium) producing both tissues, and phelloderm (secondary cortex) produced centripetally - collectively forming the waterproof insulating barrier that replaces epidermis in secondary growth, with successive periderms arising progressively deeper creating layered bark structure. Periderm three-layer structure includes= phellem (outer layer, dead suberized cork cells providing impermeability and protection, typically 0.5-3 mm thickness per periderm generation), phellogen (middle layer, single cell layer of cork cambium producing phellem and phelloderm through division, positioned at boundary between layers), and phelloderm (inner layer, living parenchyma cells forming secondary cortex, typically 0.1-0.5 mm thickness, thinner than phellem due to asymmetric cambial division), with total individual periderm thickness 0.7-4 mm depending on cork cambium activity duration and vine vigor. Periderm formation initiation includes= first periderm arising in sub-epidermal cortex (typically year 1-2 when trunk diameter exceeds 8-15 mm and primary epidermis ruptures from expansion), cork cambium initiation through dedifferentiation (cortex parenchyma cells becoming meristematic), rapid phellem production (new cork cambium quickly producing protective cork cells sealing trunk surface), establishment of complete or partial cylinder (initial periderm often forming lenticular patches gradually coalescing, eventually forming continuous protective covering), and replacement of primary protective tissues (periderm superseding epidermis and cortex as trunk matures). Periderm successive formation pattern includes= first periderm lifespan (functioning 2-5 years before becoming non-expandable and requiring replacement), second periderm formation (new cork cambium arising in outer secondary phloem beneath first periderm), continued succession (new periderms forming every 3-8 years progressively deeper), multiple overlapping periderms (mature and old trunks showing 3-8+ successive periderm generations creating layered bark structure), and progressive exterior loss (oldest outermost periderms eventually sloughing maintaining relatively stable total bark thickness despite continuous internal addition). Periderm phellem characteristics includes= dead cells at maturity (cork cells undergoing programmed death after wall deposition leaving hollow suberized boxes), suberized walls (suberin polymer and wax deposition creating waxy hydrophobic impermeable barrier), rectangular brick-like shape (cells 20-50 μm dimensions arranged in orderly radial files reflecting cambial origin), air-filled lumens (providing lightweight structure, thermal insulation, compressibility), radial alignment (cork cells stacked in files perpendicular to trunk surface), and accumulation (phellem building outward each growing season creating progressively thicker cork layer until periderm superseded by deeper formation). Periderm phellogen characteristics includes= single cell layer of meristematic initials (compared to 3-6 cell layer vascular cambium zone), rectangular to polygonal cells (20-40 μm dimensions), periclinal divisions (parallel to surface producing phellem outward and phelloderm inward), asymmetric division (80-90% divisions producing cork cells, 10-20% producing phelloderm cells), seasonal activity (active April-September producing new cork, dormant November-February), and eventual cessation (individual cork cambium functioning 2-8 years before becoming inactive and replaced by deeper cork cambium). Periderm phelloderm characteristics includes= living parenchyma cells (maintaining protoplasm and metabolic activity contrasting with dead phellem), storage function (accumulating some starch contributing to bark reserves), thin-walled cells (primary walls without extensive secondary thickening), limited radial extent (0.1-0.5 mm depth, 2-8 cell layers, much less than phellem due to asymmetric cambial production), chloroplasts in outer cells (outermost phelloderm potentially containing chloroplasts contributing minor photosynthesis in young bark), and transitional function (phelloderm connecting phellem to inner bark tissues). Periderm protective functions includes= waterproofing (suberized phellem preventing water loss from living inner bark maintaining tissue hydration, critical during drought and dormancy), pathogen barrier (impermeable cork excluding fungi and bacteria protecting susceptible living tissues), mechanical protection (compressible cork cells absorbing impacts without damage to underlying tissues), thermal insulation (air-filled cork buffering temperature extremes protecting cambium and phloem), gas exchange through lenticels (specialized porous regions enabling oxygen and carbon dioxide diffusion maintaining aerobic respiration), and fire resistance (cork providing brief protection against flames or radiant heat). Periderm spatial organization in trunk includes= first periderm in cortex (initial formation 0.5-1.5 mm beneath epidermis), second and subsequent periderms in secondary phloem (progressively deeper formations at 1-4 mm depth in outer phloem), overlapping arrangement (successive periderms creating complex layered structure with older periderms external to younger), circumferential coverage (mature periderms forming complete cylinders around trunk providing continuous protection), and lenticel interruptions (porous regions breaking otherwise impermeable coverage enabling gas exchange). Periderm thickness and bark structure includes= individual periderm 0.7-4 mm thickness (phellem 0.5-3.5 mm, phellogen single cell layer ~0.03 mm, phelloderm 0.1-0.5 mm), multiple periderms creating total outer bark (3-8 successive periderms in mature and old trunks producing 3-12 mm cumulative outer bark depth), periderm contribution to bark texture (successive periderm formation patterns determining smooth versus furrowed bark characteristics), and periderm-crushed phloem interlayering (crushed non-functional phloem between successive periderms contributing to outer bark composition). Periderm seasonal dynamics includes= spring cork cambium reactivation (typically May, later than vascular cambium), active phellem and phelloderm production (May-September with peak activity June-August), progressive cork accumulation (0.5-3 mm annual increment depending on vigor and periderm age), fall activity decline (September-October with declining division rates), winter dormancy (November-February, no periderm production), and visible annual layering (successive growing seasons sometimes creating distinguishable bands in thick phellem visible as growth lines). Periderm cultivar variation includes= thick periderm cultivars (vigorous cork cambium producing 1.5-3.5 mm annual phellem increment creating substantial protective cork), thin periderm cultivars (limited cork production 0.4-1.2 mm increment creating thinner protective covering), uniform versus irregular periderm (some cultivars showing even cork production creating smooth bark, others showing variable production creating furrowed or platy texture), and periderm formation frequency (rapid succession every 2-4 years in some varieties, slower 6-10 year intervals in others affecting bark structure). Periderm significance in bark development includes= enabling trunk expansion (successive periderm formation accommodating diameter increase while maintaining protection as outer periderms become non-expandable and slough), creating outer bark (periderm generations and crushed phloem together forming dead outer bark protecting living inner bark), determining bark appearance (periderm formation patterns creating distinctive bark colors, textures, and characteristics), maintaining trunk protection (continuous periderm succession ensuring uninterrupted protective covering throughout decades of trunk development), and defining bark-wood boundary (innermost functional cork cambium marking transition between living inner bark and dead outer bark).
Periderm emphasizes three-component protective complex and is distinguished from Cork_Layer_Periderm (phellem emphasis), Protective_Periderm (function emphasis), Cork_Cambium (phellogen only), and Outer_Trunk_Bark (multiple periderms plus crushed phloem) by focus on complete tissue unit. Critical for understanding bark formation, protective barrier structure, successive periderm replacement, waterproofing and insulation mechanisms, and integration of phellem-phellogen-phelloderm in providing trunk protection while accommodating expansion through continuous renewal.
Bark periderm
Cork tissue complex
Korkgewebe
Periderm-Komplex
Protective periderm
Schutzperiderm
Periderm
Syrah exhibits robust periderm development disposition with cork layer thickness reaching 180-220 µm by dormancy and complete circumferential coverage by node 8-10. Müller-Thurgau shows thinner periderm development (100-140 µm) with occasional gaps at lenticels increasing susceptibility to desiccation injury. Periderm suberin content varies from 35-55% of cork cell wall dry weight, with higher values in cultivars adapted to continental climates. The phellogen initiates 15-25 days after lignification begins at each internode.
A lignification disposition that is the inherited capacity of a Vitis vinifera plant to form a continuous protective periderm layer around maturing shoots through phellogen activity, producing cork cells (phellem) externally and phelloderm internally, providing barrier functions against desiccation and pathogen entry.
Periderm development is distinct from but coordinated with xylem lignification. The phellogen (cork cambium) arises from dedifferentiation of cortical parenchyma cells. Periderm quality affects winter desiccation tolerance, Eutypa infection risk at pruning wounds, and grafting success through callus-periderm interactions. VvMYB93 regulates suberin biosynthesis genes during periderm formation.
Borkenbildungsvermögen
Korkschichtentwicklung
Peridermbildung
bark development capacity
cork layer formation
periderm maturation
Evert RF (2006) Esau's Plant Anatomy: Meristems, Cells, and Tissues of the Plant Body, 3rd Ed, Wiley pp.: 427-445 Pollard M et al. (2008) Building lipid barriers: biosynthesis of cutin and suberin. Trends in Plant Science 13:236-246
Periderm Development Disposition
Periderm formation initiates at shoot nodes approximately 6-8 weeks after budbreak and progresses to internodes over the following weeks, with the characteristic color change from green to tan-brown indicating suberization completion. Cultivars differ in periderm development rate, with fast-forming types such as Cabernet Sauvignon showing 90-100% internode coverage by veraison while slow-forming cultivars such as Sultanina may reach only 50-70% coverage at the same developmental stage. Periderm thickness at maturity ranges from 150-300 µm depending on cultivar and growing conditions, with thicker periderm associated with improved cold hardiness and desiccation resistance. The VvMYB93 transcription factor regulates suberin biosynthesis genes (VvFAR, VvCYP86, VvGPAT) controlling periderm development. Incomplete periderm formation leaves shoots vulnerable to winter desiccation and freeze damage, with damage typically occurring in distal, immature portions.
A cane maturation disposition that is the inherited capacity of a Vitis vinifera plant to develop a protective periderm (cork layer) on current-season shoots through phellogen (cork cambium) activity, replacing the epidermis with suberized phellem cells that provide desiccation resistance, pathogen exclusion, and the physical barrier necessary for winter survival.
Periderm formation represents the visible indicator of cane maturation and provides essential protective functions for winter survival. The suberized cell layers restrict water loss (critical during winter desiccation stress), exclude pathogens, and provide insulation for underlying living tissues. Periderm quality and completeness directly correlate with winter hardiness assessment and cane selection for propagation. The trait shows strong environmental modulation, with water stress and moderate temperatures promoting periderm development while excessive vigor and late-season irrigation delay maturation.
Borkenbildung
Korkentwicklung
cork development
periderm development trait
shoot browning
Franke R, Schreiber L (2007) Suberin - a biopolyester forming apoplastic plant interfaces. Current Opinion in Plant Biology 10:252-259 Pollard M et al. (2008) Building lipid barriers: biosynthesis of cutin and suberin. Trends in Plant Science 13:236-246
Periderm Formation Disposition
Merlot periderm forming cork cambium at 14 years post-planting shows active phellogen at 1.5-2.2 mm depth producing complete three-layer periderm= phellem layer 1.2-2.0 mm thick comprising 18-28 radial files of suberized dead cork cells, cork cambium single cell layer with initial cells 28-38 μm dimensions actively dividing April-September, and phelloderm layer 0.2-0.4 mm thick with 3-6 cell layers of living parenchyma, complete periderm providing waterproof protection with scattered lenticels 2-4 mm diameter, represents third successive periderm (first two outer periderms visible as sloughing outer bark layers creating 5-8 mm total bark depth)
Periderm forming cork cambium is cork cambium actively producing periderm components including phellem (cork cells), cork cambium itself, and phelloderm, emphasizing the cambium's role in generating the complete protective tissue complex rather than just cork production alone. Periderm forming cork cambium produces complete three-layer structure= phellem (outer cork layer comprising dead suberized cells providing impermeability and protection), cork cambium (middle meristematic layer of dividing cells), and phelloderm (inner layer of living parenchyma cells), with all three layers collectively forming functional periderm replacing epidermis in secondary growth. Periderm forming cork cambium developmental stages include= initiation phase (cork cambium arising through dedifferentiation of cortex or phloem parenchyma), active production phase (cambium dividing to produce phellem outward and phelloderm inward building periderm structure), mature periderm phase (complete protective layer established with functional cork and living phelloderm), aging phase (periderm becoming thicker and eventually non-expandable), and replacement phase (new deeper cork cambium forming as outer periderm becomes non-functional). Periderm forming cork cambium phellem production includes= continuous centrifugal production (cork cells added to outer surface throughout growing season), cell wall suberization (suberin and wax deposition rendering walls impermeable), cell death at maturity (protoplasm degradation leaving hollow suberized boxes), radial cell files (cork cells arranged in orderly files reflecting cambial origin), typical phellem accumulation 0.5-3 mm per year (varying by vine vigor, trunk position, and environmental conditions), and multiple cell layers (10-40 cork cell layers produced annually creating protective barrier 0.5-3 mm thick). Periderm forming cork cambium phelloderm production includes= limited centripetal production (phelloderm cells comprising 10-20% of cork cambium divisions), living parenchyma cells (maintaining protoplasm and metabolic activity), storage function (accumulating starch and other reserves), chloroplast presence in outer phelloderm (contributing minor photosynthesis in young bark), typical phelloderm depth 0.1-0.5 mm (much thinner than phellem reflecting asymmetric cambial division), and transitional zone (phelloderm connecting phellem to inner bark tissues). Periderm forming cork cambium complete periderm characteristics include= waterproof barrier (suberized phellem preventing water loss and pathogen entry), gas exchange capability (lenticels in periderm enabling respiration of underlying tissues), flexibility (cork layers providing cushioning and accommodating some expansion), thermal insulation (air-filled cork cells buffering temperature fluctuations), mechanical protection (cork layers resisting abrasion and impact damage), and fire resistance (suberized cork providing some protection against brief fire exposure or heat). Periderm forming cork cambium spatial organization includes= complete cylinders in mature trunks (cork cambium forming continuous rings around trunk circumference producing uniform periderm), incomplete patches in young trunks (early periderm formation often localized before coalescing into cylinders), lenticel interruptions (specialized regions with loosely arranged cells enabling gas exchange), position varying by periderm age (first periderm in sub-epidermal cortex, successive periderms in secondary phloem progressively deeper), and multiple overlapping periderms (successive periderm-forming cork cambia creating layered bark in old trunks). Periderm forming cork cambium temporal dynamics includes= seasonal activity (active production April-September during growing season), spring reactivation (cork cambium resuming division typically May, later than vascular cambium), summer peak production (maximum phellem and phelloderm formation June-August), fall slow-down (declining division rates September-October), and winter dormancy (cork cambium quiescent November-February with no periderm production). Periderm forming cork cambium successive formation pattern includes= first periderm formation (year 1-2, arising in cortex as epidermis ruptures from trunk expansion), first periderm duration (functioning 2-5 years before expansion and aging render it non-functional), second periderm formation (year 3-7, new cork cambium arising in outer secondary phloem beneath first periderm), continued succession (new periderms forming every 3-8 years progressively deeper), and cumulative effect (multiple successive periderms creating thick layered bark 4-12 mm depth in old trunks 25-50+ years). Periderm forming cork cambium bark development includes= smooth bark in young trunks (single thin periderm layer creating relatively smooth surface), developing texture (second and third periderms creating ridges and furrows as outer layers crack), mature bark character (cultivar-specific patterns from multiple successive periderms creating distinctive textures), furrowed or platy bark (outer dead periderm layers splitting and flaking while inner layers remain functional), and thick protective bark (old trunks accumulating 6-15 mm bark depth from numerous successive periderms). Periderm forming cork cambium variation by cultivar includes= thick bark cultivars (some varieties producing vigorous cork cambium activity creating thick protective bark 8-15 mm), moderate bark cultivars (typical periderm production creating 4-8 mm bark depth), thin bark cultivars (limited cork cambium activity producing 2-5 mm bark), smooth bark types (periderm forming relatively uniform layers with limited cracking), and deeply furrowed types (successive periderms creating pronounced ridges and valleys characteristic of variety). Periderm forming cork cambium environmental responses includes= drought stress potentially reducing activity (water deficit limiting cell division and periderm production), mechanical damage triggering localized activity (wound-induced periderm formation beneath injuries), temperature effects (cork cambium requiring adequate warmth for division, cold potentially delaying spring activity), pathogen attack stimulating periderm formation (defensive wound periderm arising beneath canker or infection sites), and sun exposure potentially enhancing activity (high light and temperature on sun-exposed trunk sides possibly promoting thicker periderm). Periderm forming cork cambium significance includes= enabling continued protection (successive periderm formation maintaining bark integrity as trunk expands), replacing non-functional outer periderm (new cork cambium compensating for aging outer layers), creating bark character (periderm formation patterns determining bark texture and appearance), protecting vital tissues (periderm safeguarding cambium and phloem from desiccation, pathogens, and damage), and enabling trunk longevity (continuous periderm renewal allowing decades of trunk expansion and protection).
Periderm_Forming_Cork_Cambium emphasizes complete protective complex production and is distinguished from Cork_Cambium (general bark-forming meristem) and Outer_Cork_Cambium (position emphasis) by focus on three-layer periderm structure generation. Critical for understanding complete bark tissue organization, protective layer structure, successive periderm replacement, bark development over time, and maintenance of trunk protection through decades of growth.
Bark tissue producing cambium
Periderm-bildender Korkkambium
Protective layer forming cambium
Rindengewebe-produzierendes Phellogen
Schutzschicht-erzeugendes Kambium
Periderm Forming Cork Cambium
Cabernet Sauvignon grafted on 110 Richter rootstock at 16 years showing periderm forming rootstock cork cambium at 2.2 mm depth in outer bark, active cambium producing phellem outward creating 1.8 mm cork layer with suberized waterproof cells and phelloderm inward creating 0.4 mm living parenchyma layer, complete periderm system providing trunk surface protection against desiccation and pathogens, lenticels in cork 2-3 mm diameter enabling gas exchange, periderm forming cambium active current season producing fresh tan cork accumulating outward, two deeper inactive cork cambia at 4.5 and 7.2 mm representing previous periderms now non-functional
Periderm forming rootstock cork cambium is rootstock cork cambium actively producing complete periderm protective layer comprising cork (phellem) outward, cork cambium (phellogen) itself, and phelloderm inward, emphasizing functional role in creating protective tissue system replacing epidermis and previous periderms, showing active cell division generating suberized waterproof cork cells providing protection against desiccation, temperature extremes, mechanical damage, and pathogen entry, producing living phelloderm cells contributing to bark storage and transport, enabling trunk surface protection essential for continued trunk function, and demonstrating capacity to form complete functional periderm distinguishing active periderm-forming cambium from inactive remnant cork cambium embedded in inner bark. Periderm forming rootstock cork cambium functional emphasis includes= actively producing periderm (cork cambium currently forming complete protective tissue system not merely existing as residual tissue), generating all periderm components (producing phellem outward, maintaining phellogen, and producing phelloderm inward creating integrated protective system), creating functional protection (periderm products forming effective barrier protecting underlying trunk tissues), showing ongoing activity (cork cambium dividing and producing derivatives during current or recent growing season), and maintaining protective role (active periderm formation essential for trunk protection distinguishing functional from non-functional cambium). Periderm forming rootstock cork cambium phellem production includes= generating cork cells outward (producing rectangular cells that die at maturity forming dead protective outer layer), inducing suberization (cork cells depositing suberin in walls creating waterproof lipid-rich barrier), creating multiple cell layers (producing 5-20+ cork cell layers forming substantial protective tissue 0.5-3 mm thickness), forming continuous barrier (phellem creating uninterrupted protective covering around trunk circumference), and producing lenticels (generating loosely arranged cork cells creating gas exchange pores enabling oxygen and CO2 diffusion through otherwise impermeable cork). Periderm forming rootstock cork cambium phelloderm production includes= generating living cells inward (producing parenchyma cells remaining alive forming inner periderm layer), creating storage tissue (phelloderm cells accumulating starch and other reserves contributing to bark storage capacity), forming modest layer (producing 2-8 cell layers creating thinner phelloderm 0.1-0.5 mm compared to thicker phellem), maintaining living functions (phelloderm cells respiring, storing, and potentially synthesizing compounds unlike dead phellem), and connecting to deeper tissues (phelloderm continuous with secondary phloem facilitating physiological integration). Periderm forming rootstock cork cambium protective functions includes= preventing water loss (suberized phellem creating waterproof barrier minimizing trunk desiccation), blocking pathogen entry (cork providing physical and chemical barrier preventing fungal and bacterial penetration), insulating against temperature (cork low thermal conductivity protecting cambium from freeze and heat extremes), resisting mechanical damage (tough cork protecting trunk from abrasion, impact, and animal damage), and enabling gas exchange (lenticels in cork allowing oxygen intake and CO2 release maintaining respiration in living bark tissues). Periderm forming rootstock cork cambium in young versus old trunk includes= young trunk (years 1-5) showing first periderm forming cambium arising superficially creating initial protective bark replacing epidermis, establishing trunk (years 5-15) showing 2-3 successive periderm forming cambia creating developing bark structure, mature trunk (years 15-40) showing 3-5 active and inactive cambia creating characteristic mature bark, old trunk (years 40+) showing 5-10+ successive cambia creating thick complex bark with outermost currently forming periderm, and demonstrating progression (periderm forming cambium position moving deeper into trunk as successive cambia arise). Periderm forming rootstock cork cambium replacement pattern includes= functioning for limited period (individual periderm forming cambium active typically 3-8 years), becoming inactive eventually (cambium ceasing division as cork accumulates and cambium becomes isolated from resources), requiring replacement (new periderm forming cambium arising deeper to maintain protection as old cambium fails), showing overlapping generations (new periderm forming while old periderm still partially functional creating transition period), and enabling continued protection (successive periderm forming cambia ensuring uninterrupted trunk protection throughout vine life). Periderm forming rootstock cork cambium and bark appearance includes= determining bark texture (periderm forming cambium activity creating smooth, furrowed, plated, or shaggy bark depending on patterns), controlling bark thickness (cork production rate and periderm number determining total bark depth), creating color patterns (fresh cork typically tan-brown darkening to grey-brown with weathering and oxidation), forming distinctive features (lenticel distribution, fissure patterns, and cork accumulation creating cultivar-characteristic appearance), and enabling cultivar identification (periderm characteristics potentially diagnostic for rootstock variety recognition in mature trunk).
Periderm_Forming_Rootstock_Cork_Cambium emphasizes active protective tissue production and is distinguished from inactive remnant cork cambium by focus on current functional periderm formation. Critical for understanding bark protective function, cork-phelloderm production, trunk surface protection, waterproofing, pathogen barrier, and the active cork cambium currently producing functional periderm essential for trunk integrity.
Active rootstock cork cambium
Functional rootstock phellogen
Funktionelles Wurzelstock-Phellogen
Periderm-bildendes Unterlagen-Korkkambium
Protective-layer-forming cambium
Schutzschicht-bildendes Kambium
Periderm Forming Rootstock Cork Cambium
Syrah grafted on SO4 rootstock showing peripheral lateral root cap, outermost 2 cell layers of lateral cap approaching root surface, peripheral cells showing extreme vacuolation >92% cell volume, reduced cytoplasm compressed peripherally, microscopic examination revealing abundant Golgi vesicles producing mucilage, thick mucilage layer 15-25 micrometers coating peripheral cap surface, cell walls showing signs of loosening with intercellular spaces indicating detachment preparation, peripheral lateral root cap representing terminal maturation stage transitioning to border cell status
Peripheral lateral root cap is outermost portion of lateral root cap comprising 1-3 cell layers nearest to root surface, characterized by cells approaching or undergoing programmed senescence preparing for detachment, showing advanced maturation with extensive vacuolation and cytoplasmic degradation, demonstrating maximum mucilage production creating thick polysaccharide coating, and representing transition zone between attached cap tissue and detaching border cells, with peripheral lateral root cap cells showing typical features including very large vacuoles occupying >90% cell volume, reduced cytoplasm compressed into thin peripheral layer, thick walls potentially beginning to separate or loosen, active Golgi and ER producing and secreting mucilage vesicles, and imminent or occurring cell separation from underlying cap layers marking transition to border cell status, and demonstrating that peripheral position represents terminal stage in lateral cap cell lifespan where cells complete protective function while preparing for programmed detachment. Peripheral lateral root cap positional characteristics includes= showing outermost location (peripheral lateral cap comprising 1-3 outermost cell layers of lateral cap nearest root surface), exhibiting surface proximity (peripheral cells immediately beneath or at root surface in direct contact with soil environment), demonstrating transition position (peripheral lateral cap representing boundary between attached cap and detaching border cells), revealing marginal position (peripheral cells at lateral margin of overall cap structure), and displaying exposure (peripheral lateral cap cells most exposed to soil mechanical and chemical stresses). Peripheral lateral root cap cellular maturation includes= showing advanced age (peripheral cells typically oldest lateral cap cells 5-10 days post-production approaching terminal stage), exhibiting extreme vacuolation (vacuoles occupying >90% cell volume leaving minimal cytoplasm), demonstrating cytoplasmic degradation (cytoplasm showing progressive breakdown with organelle deterioration), revealing wall modifications (cell walls potentially loosening or separating preparing for cell detachment), and displaying senescence markers (biochemical indicators of programmed cell death and senescence visible in peripheral cells). Peripheral lateral root cap mucilage production includes= showing maximum secretion (peripheral cells producing and secreting mucilage at highest rates creating thick coating), exhibiting abundant Golgi (peripheral cells with numerous active dictyosomes producing mucilage vesicles), demonstrating continuous secretion (mucilage vesicles continuously fusing with plasma membrane releasing polysaccharides), revealing mucilage accumulation (secreted mucilage accumulating in cell walls and on root surface creating visible layer), and displaying composition complexity (peripheral mucilage showing complex mixture of polysaccharides, proteins, and other components). Peripheral lateral root cap detachment preparation includes= showing wall loosening (middle lamella and cell wall components degrading facilitating cell separation), exhibiting intercellular separation (spaces appearing between peripheral cells and underlying cap layers indicating detachment initiation), demonstrating programmed process (cell separation following developmental program rather than random damage), revealing enzyme activity (wall-degrading enzymes including polygalacturonases and cellulases active in separation zones), and displaying transition markers (molecular markers indicating cells transitioning from attached cap to border cell status). Peripheral lateral root cap protective function includes= providing outermost barrier (peripheral cells creating final protective layer between meristem and soil environment), exhibiting expendable shield (peripheral cells sacrificed protecting underlying cap and meristem from mechanical and chemical damage), demonstrating pathogen defense (peripheral mucilage and cells potentially trapping or deterring pathogens), revealing chemical buffering (peripheral cells potentially absorbing or neutralizing toxic substances protecting internal tissues), and displaying renewal capacity (continuous peripheral cell production and sloughing maintaining protective function despite losses). Peripheral lateral root cap mucilage functions includes= showing lubrication (peripheral mucilage reducing friction between root cap and soil facilitating penetration), exhibiting soil modification (mucilage binding soil particles creating structured rhizosheath around root tip), demonstrating water relations (mucilage potentially helping maintain cap hydration in drying soil through water retention), revealing ion binding (mucilage potentially binding toxic ions like aluminum protecting sensitive tissues), and displaying defensive roles (mucilage creating physical and chemical barrier potentially deterring pathogens or herbivores). Peripheral lateral root cap in border cell formation includes= showing precursor role (peripheral lateral cap cells representing immediate precursors to border cells), exhibiting detachment initiation (peripheral cells beginning separation process that completes with border cell release), demonstrating morphological similarity (peripheral cells showing similar appearance to young border cells reflecting transition), revealing functional overlap (peripheral cells and border cells both producing mucilage and providing protection), and displaying continuous production (peripheral cap cells continuously becoming border cells through detachment creating steady border cell supply). Peripheral lateral root cap turnover includes= showing rapid replacement (peripheral cells sloughing within days of reaching peripheral position requiring continuous replacement), exhibiting production balance (peripheral cell loss rate balanced by new cell addition from inner lateral cap layers maintaining layer thickness), demonstrating position-dependent fate (lateral cap cells reaching peripheral position programmed for terminal maturation and detachment), revealing dynamic population (peripheral cell population continuously renewed through basipetal cell movement and detachment), and displaying high flux (peripheral lateral cap showing highest turnover rate in cap structure reflecting terminal position).
Peripheral_Lateral_Root_Cap emphasizes outermost transitional cap layers and is distinguished from inner lateral cap by focus on terminal maturation and detachment preparation. Critical for understanding surface position, advanced senescence, maximum mucilage production, detachment preparation, border cell transition, and the outermost lateral cap layers in terminal stage preparing for programmed detachment while completing protective function.
Outermost lateral cap
Periphere laterale Wurzelhaube
Pre-border cap cells
Prä-Grenzzellen
Surface lateral cap layer
Äußerste Haubenschicht
Peripheral Lateral Root Cap
A permanent arm on Vitis vinifera cv. Merlot in a commercial vineyard, established over 3-year training period following planting, now 12 years old extending 75cm horizontally from trunk, measuring 28mm diameter showing thick gray bark with corky texture and accumulated pruning scars from annual spur renewal, bearing 12 productive two-bud spurs distributed along length, vascular system showing good conductivity supporting vigorous spring growth and adequate fruit maturation, no signs of trunk disease or vascular dysfunction, expected to remain productive for additional 8-12 years before accumulated stress, potential disease incursion, or gradual vigor decline necessitates renewal through new arm training from trunk or lower positions.
An arm established and maintained as long-term structural component intended for multi-year persistence, characterized by deliberate training to specific architecture during establishment phase (2-4 years), annual maintenance pruning preserving desired form while managing fruiting positions, thickness and lignification reflecting multi-year secondary growth accumulation, and expectation of 10-20+ year productive lifespan before eventual renewal or replacement becomes necessary due to declining vigor, disease accumulation, or structural deterioration. Permanent arms represent planned architectural components in modern training systems, contrasting with temporary or annually renewed approaches, with their establishment requiring initial investment in training time and foregone early production but providing stable platform for consistent crop production, mechanization compatibility, and predictable vineyard management once mature architecture is achieved.
Permanent arms define modern cordon and similar training systems, with their multi-year persistence enabling mechanization (consistent fruiting zone height and position), labor efficiency (predictable annual pruning rather than complete retraining), and quality consistency (stable canopy architecture and crop distribution). However, permanence is relative rather than absolute, with eventual renewal necessary as arms age, accumulate trunk diseases (Eutypa, Esca, Botryosphaeria), show declining vigor, or require repositioning for trellis modifications. The expected productive lifespan varies dramatically by region, variety, and management, ranging from 8-12 years in disease-prone regions to 20-30+ years in favorable conditions with healthy vines and excellent disease management, making permanent arm renewal planning an important long-term vineyard management consideration.
dauerhafte Arme
established arm
multi-year arm
permanente Schenkel
structural permanent arm
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Sosnowski et al. (2007) "The influence of grapevine cultivar and isolate of Eutypa lata on wood and foliar symptoms" Plant Disease 91=924-931 Mundy & Manning (2010) "Budbreak and shoot development in Vitis vinifera following severe trunk damage" Australian Journal of Grape and Wine Research 16=310-316.
Permanent Arm
The pest management description for a certified organic Chardonnay vineyard in the Mâconnais documents an integrated program comprising preventive cultural measures (fruit zone leaf removal for air circulation, shoot positioning for canopy openness, balanced nutrition avoiding excess nitrogen), a fungicide spray program limited to copper-based products (maximum 4 kg Cu/ha/year as per EU organic regulation) and sulfur (applied as wettable sulfur at 5-8 kg/ha per application, 8-12 applications per season from E-L 12 through E-L 35), biological control of Lobesia botrana (European grapevine moth) through mating disruption using 500 RAK dispensers per hectare releasing synthetic pheromone (E,Z)-7,9-dodecadienyl acetate, and a monitoring program of weekly visual scouting for powdery mildew, downy mildew, and botrytis with disease severity assessed using the EPPO standardized scale on 100 randomly selected leaves and 50 clusters per monitoring block.
A viticultural practice description that documents the strategies, methods, and materials applied to prevent, monitor, and control arthropod pests, fungal diseases, bacterial diseases, viral diseases, and vertebrate pests in the vineyard, encompassing integrated pest management frameworks, chemical and biological control measures, monitoring and threshold-based decision systems, and the regulatory and certification constraints that govern pesticide use in specific production contexts.
Pest_Management_Description addresses the domain of viticultural practice that consumes the largest share of annual variable costs in most wine regions — pesticide applications for fungal disease control typically account for 15-30% of total vineyard operating costs and represent 60-80% of all pesticide use by volume in viticulture. The dominant fungal diseases of Vitis vinifera — downy mildew (Plasmopara viticola), powdery mildew (Erysiphe necator), and grey mould (Botrytis cinerea) — are all non-native pathogens to which European grapevine cultivars lack effective genetic resistance, necessitating continuous chemical or biological intervention throughout the growing season: this fundamental susceptibility drives the viticultural industry's heavy reliance on fungicide applications and simultaneously motivates the development of resistant cultivar breeding programs (PIWI varieties) that could ultimately reduce pesticide dependency. Subclasses capture the principal dimensions of pest management documentation — integrated management frameworks, spray programs, biological control, monitoring systems, and intervention thresholds.
Pflanzenschutz-Beschreibung
Pflanzenschutzprotokoll
Schädlingsmanagement-Dokumentation
crop protection protocol
plant protection documentation
vineyard pest control record
Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests 2nd Ed, APS Press Pertot I et al. (2017) A critical review of plant protection tools for reducing pesticide use on grapevine and new perspectives for the implementation of IPM in viticulture, Crop Protection 97: 70-84 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 461-520
Pest Management Description
Comprehensive screening for Pest_Resistance_Disposition at the USDA-ARS Grape Genetics Research Unit, Geneva, New York, evaluated responses to multiple pest species across 85 Vitis accessions representing cultivated and wild germplasm. Standardized bioassays assessed phylloxera root galling, Meloidogyne incognita root-knot formation, Japanese beetle (Popillia japonica) leaf consumption, and grape berry moth (Paralobesia viteana) larval development. V. vinifera 'Chardonnay' showed broad pest susceptibility with severe phylloxera nodosities (rating 5), high nematode galling index (7.2), complete defoliation by Japanese beetles in choice tests, and 85% berry moth larval survival. In contrast, V. cinerea accessions demonstrated multiple Pest_Resistance_Dispositions with minimal phylloxera damage (rating 1-2), nematode resistance (galling index 1.5), 70% less beetle feeding than 'Chardonnay', and reduced berry moth survival (45%). Interestingly, pest resistance profiles were not uniformly correlated across pest types, with some accessions showing phylloxera resistance but nematode susceptibility, indicating that Pest_Resistance_Disposition comprises multiple independent genetic systems rather than a single generalized defense capacity. These findings inform rootstock breeding targeting specific pest complexes prevalent in different viticultural regions.
A Pest_Resistance_Disposition is a Biotic_Stress_Resistance_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through defense processes upon encounter with arthropod pests and parasitic nematodes that feed on, damage, or vector pathogens to grapevine tissues. This disposition represents the capacity to resist the diverse animal organisms that exploit grapevine as a food source, habitat, or reproductive substrate, distinct from pathogen resistance which addresses microbial threats. Major grapevine pests encompass the historically devastating grape phylloxera (Daktulosphaira vitifoliae), plant-parasitic nematodes (Xiphinema, Meloidogyne, Pratylenchus species), phloem-feeding insects (mealybugs, leafhoppers, scales), foliage-feeding insects (Japanese beetles, grape berry moth larvae), and eriophyid mites causing erineum and bud damage. Pest resistance mechanisms in plants include constitutive physical barriers (trichomes, tough tissues, latex), preformed chemical defenses (alkaloids, terpenoids, phenolics), induced defenses triggered by herbivore feeding (proteinase inhibitors, volatile emissions, phytoalexins), and tolerance mechanisms enabling productivity maintenance despite pest damage. Additionally, indirect defenses attracting natural enemies of pests through volatile signals or extrafloral resources contribute to pest suppression. For grapevine specifically, the profound susceptibility of V. vinifera roots to phylloxera historically reshaped global viticulture through the universal adoption of resistant rootstocks, while resistance to other pests varies among cultivars and provides opportunities for integrated pest management. Understanding pest resistance dispositions informs cultivar and rootstock selection, breeding program priorities, and development of sustainable pest management strategies reducing reliance on synthetic pesticides.
Pest_Resistance_Disposition is_a Biotic_Stress_Resistance_Disposition. This class encompasses resistance to all animal pests of grapevine, parallel to Pathogen_Resistance_Disposition which addresses microbial threats. Child classes include Phylloxera_Resistance_Disposition (the historically most significant), Nematode_Resistance_Disposition (important for rootstock breeding and virus vector management), Insect_Resistance_Disposition (diverse foliar and fruit pests), and Mite_Resistance_Disposition (eriophyid and spider mites). The economic and historical importance of different pests varies by region and era, with phylloxera having fundamentally transformed global viticulture in the 19th century.
Arthropoden-Resistenz
Herbivoren-Abwehr
Schädlingsresistenz-Disposition
arthropod resistance disposition
herbivore resistance
pest tolerance
Granett J et al. (2001) Biology and management of grape phylloxera, Annual Review of Entomology 46:387-412 Walling LL (2000) The myriad plant responses to herbivores, Journal of Plant Growth Regulation 19:195-216 Howe GA, Jander G (2008) Plant immunity to insect herbivores, Annual Review of Plant Biology 59:41-66
Pest Resistance Disposition
Chenin Blanc bears a Pet_Nat_Scion_Role in the Loire Valley, where the ancestral method was historically practiced for Vouvray Pétillant before traditional method adoption, and has been revived by natural wine producers creating cloudy, yeasty, off-dry sparklings with Chenin's distinctive quince and honey character. Grolleau serves in a Pet_Nat_Scion_Role for Loire pet-nats, providing light color, fresh red fruit, and low tannins suited to the style's casual, unfiltered presentation. Gamay bears a Pet_Nat_Scion_Role in Beaujolais and Loire, producing refreshing, slightly hazy sparklings with cherry and banana notes from carbonic maceration influence. Riesling serves in a Pet_Nat_Scion_Role from German and Austrian natural wine producers, with the variety's high acidity and floral aromatics well-suited to the style's fresh, low-intervention aesthetic.
A sparkling wine scion role that inheres in a grafted scion cultivar by virtue of its designation for ancestral method (méthode ancestrale) sparkling wine production, also known as pétillant naturel or pet-nat, where a single fermentation is interrupted by bottling before completion, with residual sugar fermenting in bottle to create natural effervescence without added yeasts, sugar, or dosage. Pet-nat scions must yield musts that achieve sufficient natural sugar for adequate carbonation (typically finishing at 2-3.5 atmospheres, lower than traditional method) while retaining fresh acidity and expressing primary fruit character. The ancestral method predates both traditional and Charmat methods historically, and has experienced revival in natural wine movements emphasizing minimal intervention. Pet-nat production tolerates—even celebrates—variation, cloudiness (from unfiltered yeast sediment), and unconventional flavor profiles, enabling use of diverse regional varieties not associated with conventional sparkling wine. The style emphasizes drinkability, freshness, and terroir expression over technical precision or extended aging.
Pet_Nat_Scion_Role captures the cultivar utilization for ancestral method production, which differs fundamentally from traditional and Charmat methods in using single rather than secondary fermentation. The style's tolerance for variation, sediment, and unconventional aromatics enables use of virtually any grape variety, though high-acid cultivars producing fresh base wines are generally preferred. Blanquette de Limoux's Mauzac maintains perhaps the oldest continuous ancestral method tradition (documented since 1531), predating Champagne. Pet-nat's association with natural wine movements has driven rapid category growth since approximately 2010, with production expanding globally beyond traditional regions. Crown cap closures rather than corks are typical, enabling easier sediment management and reflecting the style's informal positioning.
Méthode-ancestrale-Sorte
Pét-Nat-Sorte
Urahnen-Methode-Sorte
ancestral method variety
méthode ancestrale variety
pet-nat variety
pétillant naturel variety
Feiring A (2011) Naked Wine: Letting Grapes Do What Comes Naturally, Da Capo Press Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Legeron I (2014) Natural Wine: An Introduction to Organic and Biodynamic Wines, CICO Books Goode J, Harrop S (2011) Authentic Wine: Toward Natural and Sustainable Winemaking, University of California Press
Pet Nat Scion Role
A petiolar sinus on a mature leaf at node 6 of Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, California, exhibiting characteristic open U-shaped configuration with basal lobe margins separated by 2.4 cm at the widest point creating clear gap exposing the petiole for its full visible length, sinus depth of 2.8 cm measured from petiole attachment point to the imaginary line connecting basal lobe edges, rounded U-shaped contour with smooth curved sinus base, left basal lobe margin and right basal lobe margin diverging at approximately 65-degree angle creating wide opening, sinus margins formed by the medial edges of the basal lobes showing smooth curvature without teeth, petiole clearly visible through the sinus opening for approximately 3.5 cm above the node attachment, sinus configuration serving as definitive diagnostic feature distinguishing Cabernet Sauvignon from Merlot (closed sinus) and Pinot Noir (overlapping sinus), open U-shaped petiolar sinus characteristic being primary identification feature enabling confident field identification, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing the diagnostic open petiolar sinus configuration typical of this major cultivar.
The sinus at the basal lamina where the petiole attaches to the leaf blade in Vitis vinifera, characterized by position at the blade base flanked by basal lobes forming the sinus margins, variable configuration including open (wide gap), closed (margins meeting), or overlapping (margins crossing) patterns, U-shaped, V-shaped, or lyre-shaped contours, depth typically 1-4 cm varying with basal lobe development, width at opening ranging from 0.5-4 cm in open configurations to zero in closed or negative (overlapping) configurations, critical ampelographic importance as one of the most reliable and diagnostic cultivar identification characters, cultivar-specific patterns including Cabernet Sauvignon (open U-shaped), Merlot (closed to narrow), Pinot Noir (overlapping), and numerous intermediate configurations, and functional significance providing access to the petiole-blade junction and potentially affecting mechanical stress distribution at this critical connection point. The petiolar sinus represents the most diagnostically important sinus type, developing through the interaction of basal lobe growth patterns and petiole-blade junction geometry, exhibiting exceptional genetic control with high heritability and stability across environmental conditions, and serving as the primary or secondary diagnostic feature in virtually all ampelographic identification systems due to reliable cultivar-diagnostic configurations.
Petiolar sinus represents the single most important ampelographic diagnostic feature in grapevine cultivar identification, distinguished from lateral sinuses by basal position at petiole-blade junction. The configuration shows three primary categories= open (wide gap between basal lobe margins), closed (margins meeting or nearly meeting), and overlapping (margins crossing creating occlusion), with continuous variation within and between categories. Open configurations range from very wide (>3 cm gap) to moderately open (1-2 cm gap), closed configurations show narrow slits to complete closure, and overlapping shows slight to substantial margin crossing. Shape characteristics include U-shaped (rounded base), V-shaped (acute base), and lyre-shaped (parallel margins with rounded base) configurations, with U-shaped predominating in open sinuses and V-shaped more common in closed types. Genetic control is exceptionally strong with petiolar sinus configuration showing very high heritability, minimal environmental modification, and reliable expression across developmental stages and growing conditions, making it the most stable and reliable diagnostic character. The configuration results directly from basal lobe development patterns, with divergent small basal lobes creating open sinuses, convergent larger basal lobes creating closed sinuses, and strongly convergent crossing basal lobes creating overlapping sinuses. Ampelographic utility is exceptional, with petiolar sinus serving as primary diagnostic character in most identification keys, enabling differentiation of major cultivar groups and often providing definitive identification when combined with secondary characters. The OIV descriptor system includes detailed petiolar sinus characterization (OIV-067) as a primary descriptor with standardized protocols.
Basaleinbuchtung
Petiolarsinus
Stielbucht
basal sinus
base sinus
petiole attachment sinus
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 58-135 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 56-78.
Petiolar Sinus
The petiolar sinus depth of Sangiovese documents a deep petiolar sinus (OIV 078:7) extending 35-45 mm from the petiole insertion to the sinus margin — this deep sinus combined with the slightly overlapping shape (OIV 079:5) creates the characteristic Sangiovese leaf profile in which the basal lobes wrap partially around the petiole, creating a distinctive enclosed space at the leaf base. The depth of the petiolar sinus in Sangiovese is notably consistent across the diverse clonal population (Sangiovese Grosso, Sangiovese Piccolo, Brunello selections), suggesting strong genetic determination that is maintained despite the considerable morphological variation in other leaf characters across the approximately 100 registered Sangiovese clones.
A sinus description component that records the depth of the petiolar sinus measured from the petiole insertion point to the blade margin at the deepest point of the sinus, scored according to OIV descriptor 078 on a scale from very shallow (state 1) to very deep (state 9), providing the dimensional complement to the petiolar sinus shape descriptor that together characterizes the three-dimensional geometry of this critically important identification feature.
Petiolar_Sinus_Depth provides the quantitative dimensional information that complements the qualitative shape assessment of the petiolar sinus. While petiolar sinus shape (OIV 079, the degree of opening or overlap) is generally considered the more diagnostically powerful character, sinus depth adds discrimination between cultivars that share the same shape class — for example, two cultivars may both have a slightly overlapping petiolar sinus (OIV 079:5) but differ in sinus depth (one shallow, one deep), creating distinctly different leaf base architectures despite the shared overlap category: the depth measurement captures the extent of blade tissue development in the basal region, which is independently variable from the convergence angle that determines overlap. The combination of sinus depth and shape effectively creates a two-parameter characterization of petiolar sinus geometry that provides substantially more cultivar discrimination than either parameter alone, and is one reason why the petiolar sinus region is the most information-rich zone of the grapevine leaf for identification purposes.
Blattstiel-Buchtentiefe
Petiolarsinus-Tiefe
Stielbucht-Tiefe
basal sinus extent
petiolar sinus depth measurement
sinus petiolaris depth
Petiolar Sinus Depth
The petiolar sinus shape of Cabernet Sauvignon is strongly overlapping (OIV 079:7) with the two basal lobes crossing each other to form a narrow enclosed space with a keyhole-like opening — this tightly closed petiolar sinus, in which the lobe margins press together or overlap by 5-15 mm, is one of the most distinctive single characters identifying Cabernet Sauvignon and separates it from Merlot (slightly overlapping, OIV 079:5, lobes just touching with a narrow slit visible), Cabernet Franc (moderately overlapping, OIV 079:5-7, intermediate between its two offspring), and Sauvignon Blanc (wide open, OIV 079:1-3, with a broad U-shaped sinus and no lobe overlap). The genetic basis of petiolar sinus overlap has been partially mapped to a QTL on chromosome 1, with additional contributions from loci on chromosomes 5 and 14.
A sinus description component that records the shape of the petiolar sinus — the indentation at the base of the leaf blade surrounding the point of petiole insertion — according to OIV descriptor 079, scored on a scale describing the degree of opening from very wide open (state 1) through open, slightly overlapping, strongly overlapping, to very strongly overlapping with lobes crossed (state 9), with petiolar sinus shape considered the single most diagnostically important leaf character in classical ampelography.
Petiolar_Sinus_Shape documents what Galet (1998) described as the single most useful character for a first-pass cultivar identification — an experienced ampelographer examining a single mature leaf will typically look first at the petiolar sinus before any other character, because its shape varies enormously across cultivars (from very wide open to tightly overlapping with crossed lobes) and shows exceptional genetic stability across environments. The petiolar sinus is formed by the growth of the two basal (inferior lateral) lobes that flank the point of petiole insertion — when these lobes are short and divergent, the sinus is wide open when they are elongated and convergent, the sinus is closed and when they extend beyond the convergence point, they overlap: the degree of overlap is determined by the relative growth rate and curvature of the basal lobe margins during leaf expansion, controlled by developmental genes that pattern the spatial distribution of marginal growth zones. The petiolar sinus can also be characterized by the shape of the sinus floor (U-shaped, V-shaped, or lyre-shaped with convex sides), providing additional discrimination between cultivars sharing the same degree of overlap, and by the presence or absence of a tooth (petiolar sinus tooth) at the sinus base adjacent to the petiole.
Blattstiel-Buchtenform
Petiolarsinus-Gestalt
Stielbucht-Form
basal sinus configuration
petiolar sinus form
sinus petiolaris shape
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Chitwood DH et al. (2014) A modern ampelography, Plant Physiology 164: 259-272 UPOV (2011) Guidelines for the Conduct of Tests for Distinctness, Uniformity and Stability — Vine (Vitis L.), TG/50/9
Petiolar Sinus Shape
A petiole on a mature leaf at node 7 of Vitis vinifera cv. Cabernet Sauvignon in Bordeaux, France, measuring 11.5 cm length representing a medium-length petiole with petiolar index (petiole length / main vein length ratio) of 0.82, diameter of 3.8 mm at mid-length providing substantial mechanical support, exhibiting green coloration with slight reddish tinge particularly near lamina attachment, moderate pubescence with scattered trichomes on adaxial surface creating slight texture, containing well-developed vascular bundles visible as raised ridges on abaxial surface, demonstrating flexibility allowing leaf blade movement in wind while maintaining structural integrity supporting 4.2 gram leaf mass, positioned at approximately 45-degree angle from horizontal optimizing light interception for the mid-canopy position, showing characteristic U-shaped petiolar sinus at lamina junction diagnostic for this cultivar, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing typical petiole morphology and function providing essential mechanical and vascular connection between lamina and shoot.
The stalk-like structure connecting the leaf lamina to the shoot axis in Vitis vinifera, characterized by cylindrical to slightly flattened morphology typically 4-15 cm length and 2-5 mm diameter depending on cultivar and leaf position, composition of vascular tissues including xylem for water and nutrient transport and phloem for photosynthate export, mechanical support tissues including collenchyma and sclerenchyma providing structural integrity, parenchymatous ground tissue, epidermis with variable trichome coverage, cultivar-specific characteristics including relative length compared to main vein length (petiolar index), thickness, coloration ranging from green to red-purple, pubescence patterns, and anatomical features that determine flexibility and strength. The petiole serves multiple critical functions including mechanical support positioning the lamina for optimal light interception while allowing movement in response to wind reducing mechanical stress, vascular connection providing the sole pathway for water and nutrient delivery to the leaf and photosynthate export to the rest of the plant, and responding to environmental signals through changes in turgor affecting leaf angle and orientation. Petiolar characteristics constitute important ampelographic features for cultivar identification, with petiolar length, thickness, coloration, and pubescence showing cultivar-specific patterns, and the petiolar sinus where the petiole attaches to the lamina exhibiting diagnostic shapes including open, closed, or overlapping configurations that serve as primary identification characters.
Petioles are distinguished from other leaf structures including the lamina (blade) and from stem structures, serving as the specialized connection organ. Petiolar characteristics show substantial variation among cultivars making them valuable ampelographic features, with measurements typically standardized to mature leaves at specific node positions (typically nodes 5-8) to enable consistent comparisons. Length variation occurs both among cultivars and within plants based on position, light environment, and developmental timing, with shade leaves typically developing longer petioles than sun leaves enabling better light capture in low-light environments. The petiolar sinus, formed where the petiole attaches to the lamina base, shows particularly important diagnostic characteristics with open, closed, or overlapping configurations serving as primary identification features in ampelographic keys. Petiole coloration, particularly red or purple pigmentation from anthocyanin accumulation, shows cultivar specificity and environmental plasticity, intensifying under high light, cool temperatures, or nutrient stress. Functional importance includes not only mechanical support and vascular connection but also roles in leaf angle adjustment through differential growth or turgor changes affecting photosynthetic light capture.
Blattstiel
Blattstängel
Petiolus
leaf stalk
leaf stem
petiolus
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 136-165 This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 33-55.
Petiole
In Grenache vines in Châteauneuf-du-Pape, Rhône Valley during November-December, petiole abscission occurs 7-12 days after complete leaf yellowing, with abscission zone activation visible as a constriction ring at the petiole base, ethylene production in abscission zone tissue reaching 2.5 nL/g/h during active separation, cell wall degrading enzyme activity peaking at 3 days before detachment, and clean leaf scars with suberized protective layer formed within 48 hours of separation under cool Mediterranean autumn conditions averaging 8°C.
Petiole abscission constitutes the process of physical leaf separation from Vitis vinifera shoots through the formation, activation, and completion of abscission zone cell separation at the base of the petiole, wherein coordinated cell wall degradation and middle lamella dissolution in specialized abscission zone cells enables clean detachment of senescent leaves while simultaneously sealing the wound surface to prevent pathogen entry and desiccation of exposed shoot tissues during the 1-2 week period following senescence completion. This process represents the final phase of the leaf fall phenological sequence, occurring after nutrient remobilization has recovered the majority of transportable compounds from leaf tissues and senescence has progressed to the point where continued leaf attachment provides no further benefit to the vine. The anatomical basis of petiole abscission involves the abscission zone, a specialized region of small, cytoplasmically dense cells located at the junction of petiole and shoot that remains developmentally competent for separation throughout the leaf's life span, with activation of this zone in response to senescence-associated signals initiating the cell separation processes that culminate in leaf detachment. The physiological regulation of petiole abscission involves hormonal balance shifts, with declining auxin flux from senescing leaf blades removing the primary inhibition of abscission zone activation, increasing ethylene sensitivity in abscission zone cells enabling response to senescence-derived ethylene, and potential abscisic acid involvement in coordinating abscission with overall senescence progression. The protective functions of petiole abscission include formation of a protective layer on the shoot side of the abscission zone through suberization and lignification of cell walls in the separation region, creating a barrier against pathogen invasion and water loss at the leaf scar that would otherwise represent a vulnerable entry point during the extended dormant period. Environmental factors influence petiole abscission timing, with cool temperatures accelerating abscission zone activation after senescence completion, frost events potentially inducing rapid abscission through ice crystal damage to abscission zone cells, and maintained warm conditions delaying abscission such that senescent leaves may persist attached for extended periods before separation occurs. The completion of petiole abscission for all leaves on a vine marks the transition to full dormancy, with defoliated vines entering the winter period when all metabolic activity is supported by accumulated reserves and the absence of transpiring surfaces allows survival through periods of soil water unavailability in frozen or drought-affected conditions.
Petiole_Abscission is parent class under Leaf_Fall representing physical leaf separation process. Contains subclasses Abscission_Zone_Formation, Abscission_Zone_Activation, Cell_Separation_Process, and Petiole_Abscission_Completion. Duration 1-2 weeks after senescence completion. Involves specialized abscission zone cells. Hormonal regulation through auxin and ethylene balance. Creates protective leaf scar.
Blatt-Ablösung
Blatt-Spross-Trennung
Blattstiel-Abszission
Blattstiel-Trennung
leaf detachment
leaf stalk abscission
leaf-shoot separation
petiole separation
Roberts JA et al (2002) Abscission, dehiscence and other cell separation processes. Annual Review of Plant Biology 53: 131-158 Patterson SE (2001) Cutting loose. Plant Physiology 126: 494-500 Keller M (2020) The Science of Grapevines 3rd Ed pp 92-95
Petiole Abscission
In Touriga Nacional vines in Douro Valley, Portugal during late November, petiole abscission completion occurs following overnight temperatures of 2°C combined with morning wind gusts of 25 km/h, with leaf separation occurring at the enzyme-weakened abscission zone, callose deposition in phloem visible within 30 minutes of separation, suberization of leaf scar cells detectable within 24 hours, and complete protective layer established within 72 hours with suberized cell layers measuring 45-60 µm thickness providing durable wound protection through the dormant period.
Petiole abscission completion describes the final stage of leaf separation from Vitis vinifera shoots wherein physical detachment is achieved and protective mechanisms seal the resulting leaf scar, encompassing the actual moment of separation when remaining cellular connections are severed, the immediate wound response that prevents pathogen entry and water loss from exposed shoot tissues, and the formation of a protective layer through suberization and lignification that will persist through the dormant season until new growth emerges in spring. This process represents the culmination of the abscission sequence, occurring when cell wall degradation has progressed sufficiently that leaf weight, wind forces, or rain impact provides adequate mechanical stress to sever remaining connections, with separation occurring at the structurally weakened plane created by cell wall dissolution in the abscission zone rather than through tissue rupture that would create irregular wound surfaces. The protective layer formation on the shoot side of the separation plane involves rapid physiological responses including callose deposition at sieve plates sealing phloem elements within minutes of separation, oxidative reactions generating quinones that provide antimicrobial protection at the wound surface, and subsequent suberization and lignification of cell walls in the separation layer creating a durable barrier against pathogen invasion and desiccation. The timing of abscission completion relative to environmental conditions influences vine vulnerability, with completion before the onset of freezing temperatures allowing adequate protective layer development while delayed abscission into freezing conditions may compromise wound healing responses, potentially creating entry points for winter trunk diseases. The leaf scar remaining after abscission completion displays characteristic anatomical features including the sealed vascular traces where xylem and phloem elements were severed, the suberized protective layer that forms the visible scar surface, and the underlying living bark tissues that remain connected to the vine's perennial vascular system. Environmental factors affecting abscission completion include wind speed providing mechanical force for final separation, precipitation that may simultaneously moisten tissues facilitating separation and provide physical impact force, and temperature affecting both the final enzymatic weakening of attachment and the wound healing responses following separation. The mass abscission events characteristic of late autumn occur when environmental conditions combine with advanced cell separation to trigger simultaneous leaf fall across large portions of the canopy, with cool nights followed by morning wind gusts commonly associated with major leaf fall events that can substantially defoliate vines within 24-48 hour periods.
Petiole_Abscission_Completion is subclass of Petiole_Abscission representing final separation and wound sealing. Distinguishes from cell separation by emphasizing completion and protection. Sibling to Abscission_Zone_Formation, Abscission_Zone_Activation, and Cell_Separation_Process. Involves physical detachment and protective layer formation. Environmental triggers include wind and temperature. Creates sealed leaf scar.
Abszissions-Finalisierung
Blattfall-Vervollständigung
Blattnarben-Bildung
Blattstiel-Abszissions-Abschluss
abscission finalization
leaf detachment completion
leaf fall completion
leaf scar formation
Petiole Abscission Completion
The petiole color of Malbec documents intense, uniform red-violet anthocyanin pigmentation extending from the node insertion to the blade junction, with no green ground color visible on any surface — this complete anthocyanin coverage is consistent with Malbec's characteristically strong constitutive pigmentation across all vegetative organs (deeply pigmented shoot tips, red-striped internodes, red-pigmented leaf veins), creating what French ampelographers term a cultivar à port coloré (colored-bearing cultivar). The petiole pigmentation intensity of Malbec (among the strongest in V. vinifera) contrasts with the green, unpigmented petioles of white cultivars such as Chardonnay and Sauvignon Blanc (no anthocyanin in petiole tissue), the faintly blushed petioles of Pinot Noir (pink-red dorsal pigmentation with green ventral surface), and the distinctly red-striped petioles of Tempranillo (alternating red and green longitudinal bands reflecting the vascular bundle distribution).
A petiole description component that records the color and anthocyanin pigmentation pattern of the leaf petiole, documenting the ground color (green, yellow-green, or red-green), the presence and intensity of anthocyanin pigmentation (from absent through faint blush to deep red-violet), and the distribution pattern of pigmentation (uniform, dorsal only, striped, or confined to the proximal or distal portion), providing cultivar-diagnostic color characters that complement the vein pigmentation observed on the leaf blade and reflect the constitutive anthocyanin biosynthetic capacity in vascular-associated tissues.
Petiole_Color captures the pigmentation phenotype of an organ that provides a relatively unambiguous assessment of constitutive anthocyanin production capacity in vegetative vascular tissue. The petiole is particularly informative because its anthocyanin expression is less influenced by light exposure than shoot internode pigmentation (the petiole receives variable light depending on canopy position) yet consistently reflects the genetically determined pigmentation capacity — heavily pigmented cultivars produce deeply colored petioles even on shaded leaves within the canopy interior: this stability makes petiole color a more reliable indicator of constitutive pigmentation genotype than internode or shoot tip color, which are more sensitive to light-mediated anthocyanin induction. The distribution pattern of petiole anthocyanin provides additional cultivar-diagnostic information — uniform pigmentation (entire petiole surface equally colored), dorsal pigmentation (pigment confined to the sun-exposed upper surface), striped pigmentation (anthocyanin associated with vascular bundle positions creating longitudinal stripes), and gradient pigmentation (intensity decreasing from proximal to distal or vice versa) each characterize different cultivar groups and reflect differences in the tissue-specific regulation of anthocyanin pathway gene expression.
Blattstiel-Anthocyan-Dokumentation
Blattstielfarbe
Petiolus-Pigmentierung
leaf stalk color assessment
petiole pigmentation record
petiolus anthocyanin documentation
Petiole Color
Petiole conductivity assessment comparing vulnerability to stem in Cabernet Sauvignon reveals hydraulic segmentation protecting stem investment, with petioles showing P50 of -1.3 MPa (embolizing first under drought) while stems show P50 of -1.9 MPa (more resistant), such that severe drought causes petiole embolism and leaf abscission before stem damage occurs, demonstrating the hydraulic fuse function of petioles that sacrifices expendable leaves to protect the perennial woody structure from irreversible hydraulic failure.
Petiole conductivity quality is a stem hydraulic conductivity quality that represents the water transport capacity of the leaf petiole connecting stem xylem to the leaf blade, characterizing the final segment of the woody transport pathway that supplies each individual leaf and that may function as a hydraulic vulnerability point protecting the stem through preferential embolism under severe stress. Within the BFO framework, this quality specifically addresses the petiole component of the water transport pathway, distinguishing this terminal stem segment from trunk and branch conductivity while recognizing that petiole hydraulic properties affect the water supply to individual leaves and may play a protective role in the vine's hydraulic architecture through segmentation. The ontological significance lies in petiole conductivity determining the water supply capacity to individual leaves while potentially serving as a hydraulic fuse, where petioles may be designed to embolize before stems under severe drought, sacrificing leaves to protect the more costly woody infrastructure from irreversible hydraulic damage. The anatomical basis for petiole conductivity involves the continuation of xylem from the stem into the relatively narrow petiole cross-section, with vessel dimensions typically narrower than in the subtending stem (creating a conductivity bottleneck), and with the petiole representing a hydraulic constriction that may limit water supply to leaves under high transpiration demand. The functional significance of petiole conductivity extends to its role in hydraulic segmentation, where the vulnerability of petioles to embolism relative to stems determines whether drought-induced hydraulic failure occurs first in leaves (protecting stems) or propagates through the system, with empirical evidence suggesting that grapevine petioles are often more vulnerable than stems, consistent with a protective segmentation strategy. The measurement of petiole conductivity employs similar approaches to stem conductivity (hydraulic perfusion, anatomical calculation) but with technical challenges from the smaller dimensions, and with the comparison of petiole to stem vulnerability indicating the degree of hydraulic segmentation. Understanding petiole conductivity quality enables assessment of leaf water supply capacity, evaluation of hydraulic segmentation protecting stem integrity, and interpretation of the hydraulic architecture decisions affecting leaf performance and drought response.
Petiole_Conductivity_Quality addresses the terminal woody pathway to leaves that may serve as a hydraulic vulnerability point. This quality affects individual leaf water supply and may protect stems through preferential embolism. The class supports hydraulic segmentation assessment and leaf-level water supply evaluation through petiole conductivity characterization.
Blattstielleitfähigkeit
Blattstielwasserversorgung
Petiolhydraulik
leaf stalk hydraulics
petiole conductance
petiole water supply
Choat B et al. (2005) A comparison of water stress and xylem conductance in leaves of two species of grapevine. American Journal of Botany 92(10): 1549-1556 Tyree MT et al. (1993) Vulnerability of xylem to cavitation and embolism. Annual Review of Plant Physiology and Plant Molecular Biology 44(1): 19-39
Petiole Conductivity Quality
The petiole description of Cabernet Sauvignon documents a medium-length petiole (OIV 093a:5, approximately 70-90% of the mid-vein length) with strong anthocyanin pigmentation producing a red-violet color extending from the node insertion to the blade junction (complementing the adaxial vein pigmentation characteristic of this cultivar), medium density of prostrate hairs, and a dorsiventrally flattened cross-section with the adaxial groove (petiolar channel) clearly defined — the strongly pigmented petiole is one of the characters that contributes to Cabernet Sauvignon's overall impression of a dark, heavily pigmented vine, consistent across shoot tip, young shoot, vein, and petiole tissues, and contrasting with the green, unpigmented petioles of Sauvignon Blanc (one of its parents) and the moderately pigmented petioles of Cabernet Franc (the other parent).
An ampelographic description component that documents the morphological characteristics of the leaf petiole — the stalk connecting the leaf blade to the shoot node — recording length, color, anthocyanin pigmentation, cross-sectional shape, and surface hairiness that provide cultivar-diagnostic characters complementing the leaf blade descriptors and contributing additional identification information based on the vascular support structure of the leaf.
Petiole_Description addresses an organ that receives less attention than the leaf blade in ampelographic practice but provides genuinely independent diagnostic information — petiole characters are not simply extensions of blade characters but reflect the distinct developmental program of the vascular support structure. The petiole develops from the leaf primordium base, with its length, thickness, and pigmentation determined by growth patterns that are partially independent of the blade expansion processes controlling blade shape and size: this developmental independence means that petiole characters can discriminate between cultivars that share similar blade morphology — for example, two five-lobed pentagonal-leaved cultivars with overlapping petiolar sinuses might differ in petiole length, color, and hairiness, providing additional identification resolution beyond the blade descriptors. The petiole is also the organ from which tissue samples are collected for nutritional analysis (petiole sampling at flowering and véraison for mineral nutrient diagnostics), creating a connection between the ampelographic description of the petiole and the viticultural management of vine nutrition documented elsewhere in the VVO.
Blattstiel-Beschreibung
Blattstiel-Morphologie
Petiolus-Charakterisierung
leaf stalk description
petiole characterization
petiolus morphology
Petiole Description
The petiole length of Grenache documents long petioles (OIV state 7) measuring 100-140 mm on representative mature leaves from mid-shoot positions — the petiole length exceeds the mid-vein length of the corresponding blade (ratio approximately 1.1-1.3), positioning the leaf blade well away from the shoot axis and creating the open, loosely arranged canopy architecture that is characteristic of Grenache and contributes to good air circulation within the fruit zone. This long petiole contrasts with the short petioles of Riesling (OIV state 3, 50-70 mm, petiole/mid-vein ratio approximately 0.6-0.8) that hold the leaf blade close to the shoot, creating the compact, layered canopy typical of Riesling on vertical shoot positioning systems, and the medium petioles of Chardonnay (OIV state 5, 70-100 mm, ratio approximately 0.9-1.0).
A petiole description component that records the length of the leaf petiole measured from its insertion at the shoot node to its junction with the leaf blade base, expressed either as absolute length in millimeters or as a ratio relative to the mid-vein length of the corresponding leaf blade, scored according to OIV conventions on a scale from very short (state 1) to very long (state 9), with petiole length influencing the positioning of the leaf blade relative to the shoot axis and consequently affecting canopy light interception patterns and the ease of mechanized canopy management operations.
Petiole_Length documents a quantitative character that has both diagnostic and functional significance. Diagnostically, petiole length relative to blade size is a cultivar-characteristic ratio that is more genetically stable than absolute petiole length — the ratio compensates for the vigor-dependent variation in absolute organ size and provides a proportional character that is consistent across growing conditions: cultivars with inherently long petioles (Grenache, Mourvèdre, Cinsaut) maintain high petiole/blade ratios across vigor levels, while short-petioled cultivars (Riesling, some Pinot clones) consistently show low ratios. Functionally, petiole length determines the spatial positioning of the leaf blade relative to the shoot and adjacent leaves, influencing the degree of leaf overlap, the orientation of leaves relative to incident light, and the resistance to wind-induced leaf flutter (longer petioles produce more leaf movement) — these canopy architecture effects have consequences for photosynthetic efficiency, fruit zone microclimate, and disease pressure that connect the ampelographic character to viticultural management considerations.
Blattstiel-Dimensionserfassung
Blattstiellänge
Petiolus-Längenmessung
leaf stalk length measurement
petiole dimension record
petiolus length characterization
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Smart RE and Robinson M (1991) Sunlight into Wine, Winetitles
Petiole Length
Petrichor aroma identification in old-vine Côtes du Rhône Villages using native yeast fermentation reveals subtle rain-on-earth character (4.5 on 10-point scale) contributing to the natural, terroir-connected impression alongside garrigue herbs and ripe fruit, with the petrichor note absent in same-vineyard wine using commercial yeast (rating 1.5), suggesting microbial contribution to this evocative earthiness, while excessive geosmin from contaminated fruit creates unpleasant beetroot-mud character (negative quality indicator) rather than pleasant petrichor, demonstrating the narrow concentration window for positive earthy expression.
Petrichor aroma quality is an earthy aroma quality that represents olfactory perception evoking the distinctive scent of rain on dry earth, characterizing the complex, pleasant earthy-fresh aromatic impression that contributes to the sense of natural, terroir-connected character in wines displaying this evocative descriptor. Within the BFO framework, this quality specifically addresses the rain-on-earth subcategory of earthy aroma, distinguished by its reference to the specific meteorological phenomenon and its association with fresh, natural, pre-industrial earth character rather than musty or stale earthiness. The ontological significance lies in petrichor representing an unusually poetic wine descriptor that evokes emotional and aesthetic responses beyond technical assessment, where the perception creates impressions of nature, authenticity, and place that contribute to wine's ability to communicate meaning beyond flavor, making petrichor a marker of terroir poetry rather than mere sensory attribute. The compositional basis for petrichor perception involves geosmin (trans-1,10-dimethyl-trans-9-decalol), a sesquiterpene produced by Streptomyces bacteria and blue-green algae with extremely low detection threshold (approximately 10 ng/L), which provides the characteristic earthy-rain smell, combined with plant oils (particularly from drought-stressed vegetation) and ozone contributing to the fresh quality of the perception. In wine context, geosmin occurs at trace levels that may contribute to earthy complexity without reaching the beetroot-musty character of higher concentrations, while additional compounds including certain terpenoids and volatile thiols may contribute to the complete petrichor impression. The occurrence of petrichor character in wine correlates with certain production methods (native yeast fermentation, minimal intervention) and terroirs (old vines, diverse vineyard ecosystems) that suggest a role for environmental microbiome in creating this distinctive earthiness, though the relationship remains more observational than scientifically established. Understanding petrichor aroma quality enables recognition of this evocative descriptor when appropriate, appreciation of wine's capacity for poetic sensory communication, and terroir discourse that connects wine to natural phenomena beyond grape and fermentation chemistry.
Petrichor_Aroma_Quality addresses the specifically rain-on-earth subcategory of earthy aroma, representing an evocative, poetic descriptor that contributes to terroir discourse beyond technical flavor assessment. This quality is associated with geosmin at trace concentrations where the perception is pleasant and natural rather than musty or off-putting. The class connects wine sensory assessment to broader aesthetic and emotional responses evoked by natural phenomena.
Geosmin-Note
Petrichor-Aroma
Regenduft
feuchte Erde-Charakter
fresh earth scent
geosmin character
rain on earth aroma
wet earth notes
Bear IJ, Thomas RG (1964) Nature of argillaceous odour. Nature 201(4923): 993-995 Darriet P et al. (2000) Identification of a powerful aromatic component of Vitis vinifera L. var. Sauvignon wines. Flavour and Fragrance Journal 15(5): 297-303
Petrichor Aroma Quality
Petunidin content quality analysis in Malbec from Mendoza reveals moderately elevated proportions (8-14% of total anthocyanins) compared to Cabernet Sauvignon (5-9%), contributing to the distinctive blue-black hue that characterizes Argentine Malbec the combination of high total anthocyanins (900-1400 mg/kg), elevated petunidin and delphinidin (trihydroxylated anthocyanidins comprising 88-94% of total), and substantial acylation produces the intense, inky color and purple-rimmed appearance that distinguishes high-altitude Mendoza Malbec as among the most deeply colored wines globally.
Petunidin_Content_Quality represents the chemical quality characterizing the concentration of petunidin-based anthocyanins in grape berry skins, capturing the monomethylated derivative of delphinidin that occupies an intermediate biosynthetic and structural position in the trihydroxylated anthocyanidin pathway. Petunidin (3'-methoxy-4',5',5,7-tetrahydroxyflavylium) is formed by O-methylation of delphinidin at the 3' position, catalyzed by anthocyanin O-methyltransferase (AOMT), and serves as the immediate precursor to malvidin through further methylation at the 5' position. This intermediate methylation state places petunidin between the highly hydroxylated delphinidin and the fully methylated malvidin in both biosynthetic sequence and chemical stability. Petunidin-based anthocyanins typically comprise 5-12% of total anthocyanins in most red V. vinifera cultivars, representing a transient pool in the pathway toward malvidin accumulation. The petunidin proportion relative to delphinidin indicates the efficiency of the first methylation step, while the ratio to malvidin reflects the second methylation efficiency. Petunidin contributes blue-violet coloration similar to delphinidin but with somewhat greater stability due to the protective methoxy group. The consistent presence of petunidin as a minority component provides a varietal marker, with some cultivars showing elevated petunidin that affects color quality. Petunidin-3-glucoside and its acylated derivatives are quantified by HPLC, contributing to comprehensive anthocyanin profiles used for cultivar identification and quality assessment.
Petunidin_Content_Quality captures the biosynthetic intermediate between delphinidin and malvidin in the trihydroxylated anthocyanidin pathway. The sequential methylation (delphinidin → petunidin → malvidin) is catalyzed by AOMT with varying efficiency among cultivars, affecting the proportions of each anthocyanidin accumulated. Petunidin stability is intermediate between unmethylated delphinidin and fully methylated malvidin. The consistent presence of petunidin at 5-12% of total anthocyanins provides cultivar-specific fingerprinting data, with deviations potentially indicating unusual genetic backgrounds or environmental effects on methyltransferase activity.
Petunidin-3-glucosid-Konzentration
Petunidingehalt
Petunidinpigmentqualität
Pt-3-glc content
monomethoxylated trihydroxy-anthocyanin concentration
monomethoxylierter Trihydroxy-Anthocyangehalt
petunidin pigment quality
petunidin-3-glucoside content
Fournier-Level A et al. (2011) Quantitative Genetic Bases of Anthocyanin Variation in Grape (Vitis vinifera L. ssp. sativa) Berry: A Quantitative Trait Locus to Quantitative Trait Nucleotide Integrated Study. Genetics 189(4): 1427-1441 Mattivi F et al. (2006) Metabolite Profiling of Grape: Flavonols and Anthocyanins. Journal of Agricultural and Food Chemistry 54(20): 7692-7702
Petunidin Content Quality
The PGI name Terre Siciliane is registered in the EU eAmbrosia database as a Protected Geographic Indication for wines produced in Sicily, replacing the former IGT Sicilia designation for the PGI tier (while Sicilia DOC now operates as a separate PDO) — the registered name Terre Siciliane must appear on labels of PGI-classified Sicilian wines and is protected across the EU single market against unauthorized use, with the Assessorato Regionale dell'Agricoltura authorized to pursue enforcement actions. The dual existence of PGI Terre Siciliane and PDO Sicilia DOC for overlapping territory illustrates the complexity of the EU geographic indication system where multiple protection tiers can coexist within the same geographic area.
A protected geographic indication consisting of the officially registered textual name of the geographic origin as recorded in the relevant PGI registry, constituting the specific designation string that is legally protected against unauthorized commercial use, imitation, or evocation under the same enforcement framework applicable to PDO names.
PGI_Name functions identically to PDO_Name as a legally protected textual identifier but within the PGI regulatory framework. The scope of legal protection for PGI names under EU Regulation 1308/2013 Art. 103 is equivalent to that afforded PDO names — covering direct use, misuse, imitation, evocation, and any misleading indication of geographic origin: the distinction between PDO and PGI lies in the registration requirements and production rules rather than in the strength of name protection. PGI names tend to reference broader geographic areas than PDO names (e.g., Pays d'Oc versus Minervois, Terre Siciliane versus Etna DOC), reflecting the PGI tier's function as a flexible intermediate classification covering larger production volumes and more diverse wine styles than the corresponding PDO designations within the same territory.
PGI designation string
PGI-Bezeichnungstext
eingetragener g.g.A.-Name
geschützter geografischer Name
protected geographic name
registered PGI name
EU Regulation 1308/2013 Art. 103 on protection of designations EU eAmbrosia database European Commission Delegated Regulation 2019/33
PGI Name
The PGI registration number PGI-FR-A0766 uniquely identifies the Pays d'Oc designation in the EU eAmbrosia database, with the code structure encoding the protection type (PGI), the member state of origin (FR for France), and a sequential filing number (A0766) — this registration number distinguishes the PGI Pays d'Oc record from the approximately 1,600 other wine geographic indications in the EU database and serves as the persistent key for regulatory cross-referencing, amendment tracking, and international notification under TRIPS obligations.
A protected geographic indication consisting of the unique alphanumeric identifier assigned to a registered PGI by the registering authority, providing a persistent machine-readable reference code that unambiguously identifies the PGI record within the registration database independently of name variations or translations.
PGI_Registration_Number follows the same structural conventions as PDO_Registration_Number within the EU eAmbrosia database, using the pattern [protection type]-[country code]-[sequence number]. The PGI prefix distinguishes these records from PDO records within the unified database: this distinction is operationally important because the same geographic name may appear in both PDO and PGI registrations (e.g., a region may have both a PDO for wines meeting stricter production rules and a PGI for wines meeting broader criteria), and the registration number resolves this potential ambiguity. The approximately 1,600 wine geographic indications registered in the EU system are roughly equally divided between PDOs and PGIs, with the PGI category growing more rapidly as wine-producing regions outside traditional appellation systems seek formal geographic protection.
PGI database code
PGI file number
PGI registration identifier
PGI-Datenbankkennung
PGI-Registriernummer
g.g.A.-Aktenzeichen
EU eAmbrosia database of registered geographic indications EU Regulation 1308/2013 Art. 96-97 on registration procedures
PGI Registration Number
The PGI specification reference for Pays d'Oc comprises the Cahier des charges de l'indication géographique protégée Pays d'Oc (approved by decree and published in the Journal Officiel de la République Française) — the specification defines 58 authorized grape varieties (versus 2-6 typical for PDO appellations within the same territory), maximum yields of 90 hl/ha (versus 45-60 hl/ha for regional PDOs), permits varietal labeling and vintage labeling, requires minimum natural alcohol of 9% (versus 11-12.5% for PDOs), and designates the Institut National de l'Origine et de la Qualité (INAO) and an approved certification body as the control authorities for verifying compliance.
A protected geographic indication consisting of a bibliographic or documentary reference identifying the official product specification that defines the production rules, quality requirements, geographic delimitation, and control procedures governing the use of the PGI name, with PGI specifications typically permitting broader production parameters than corresponding PDO specifications for the same or overlapping geographic areas.
PGI_Specification_Reference links the registered PGI name to the binding regulatory document defining production requirements. PGI specifications are structurally similar to PDO specifications but characteristically permit broader production parameters — more authorized varieties, higher maximum yields, less restrictive winemaking rules, and broader geographic scope: this regulatory flexibility is the defining functional characteristic of the PGI tier and explains its commercial attractiveness for producers seeking geographic origin identification without the production constraints of full PDO compliance. The specification document for PGIs follows the same EU regulatory framework (Regulation 1308/2013 Art. 94) as PDO specifications and must be publicly accessible through the same EU DOOR database and national publication channels.
PGI cahier des charges reference
PGI product specification identifier
PGI specification document
PGI-Produktspezifikations-Kennung
PGI-Spezifikationsreferenz
g.g.A.-Pflichtenheft-Verweis
EU Regulation 1308/2013 Art. 94 on product specifications INAO (2023) Cahiers des charges des indications géographiques protégées EU DOOR database
PGI Specification Reference
In Cabernet Sauvignon berries in Napa Valley, California during BBCH 85-89, pH increase proceeds from 2.88 at veraison to 3.62 at harvest over 45 days, with the 0.74 unit rise correlating with malic acid decline from 9.5 to 2.2 g/L and potassium increase from 22 to 38 mM, while titratable acidity decreases from 14.8 to 5.6 g/L, demonstrating the chemical linkage between acid catabolism and pH elevation.
pH increase describes the progressive rise in hydrogen ion concentration measured as increasing pH values within Vitis vinifera berry juice during ripening, wherein pH elevates from pre-veraison values of 2.6-3.0 to harvest values of 3.2-3.8 as a direct chemical consequence of organic acid reduction, particularly the catabolism of malic acid which contributes disproportionately to titratable acidity due to its higher dissociation constant compared to tartaric acid. This process reflects the fundamental relationship between organic acid concentration and hydrogen ion activity in the complex buffer system of grape juice, where the removal of malic acid molecules through enzymatic degradation eliminates both the undissociated acid and its conjugate base from the equilibrium, shifting pH upward according to Henderson-Hasselbalch relationships governing weak acid dissociation. The magnitude of pH increase during ripening typically spans 0.4-0.8 pH units, with the rate and extent determined by initial acid composition, degree of malic acid degradation, potassium accumulation affecting tartrate precipitation, and cultivar-specific organic acid profiles. Malic acid with pKa values of 3.40 and 5.11 contributes more strongly to juice acidity per gram than tartaric acid with pKa values of 2.98 and 4.34, meaning that malic acid degradation produces proportionally greater pH increase than equivalent tartaric acid removal, explaining the strong correlation between malate catabolism and pH elevation during ripening. Potassium accumulation during ripening, increasing from 15-25 to 25-40 mM, further elevates pH through formation of insoluble potassium bitartrate which precipitates tartaric acid from solution and reduces total tartrate contribution to acidity. Temperature influences pH increase indirectly through effects on malic acid degradation rates, with warm-climate berries typically achieving pH values 0.2-0.4 units higher than cool-climate equivalents due to accelerated malate catabolism. The viticultural significance of pH increase encompasses wine stability, color expression in red wines where higher pH shifts anthocyanin equilibria toward less colored forms, microbial stability with elevated pH increasing spoilage risk, and sulfur dioxide efficacy which decreases substantially above pH 3.5. Monitoring pH progression through ripening enables harvest timing decisions optimizing the balance between sugar accumulation and acid retention for specific wine style objectives.
pH_Increase is subclass of Active_Acid_Reduction representing chemical consequence of acid catabolism. Distinguishes from Malate_Degradation which is biochemical cause and from Continuing_Acid_Decline which describes temporal pattern. Not subdivided further. Duration parallels active acid reduction at 25-40 days. Key parameter is pH rise of 0.4-0.8 units from 2.6-3.0 to 3.2-3.8. Important wine quality determinant.
Aciditäts-pH-Anstieg
Reife-pH-Zunahme
Saft-pH-Erhöhung
acidity pH rise
hydrogen ion decrease
juice pH elevation
pH-Anstieg
ripening pH increase
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 272-278 Conde C et al (2007) Biochemical changes throughout grape berry development and ripening. Food Chemistry 1: 1-22 Coombe BG (1987) Influence of temperature on composition and quality of grapes. Acta Horticulturae 206: 23-35
pH Increase
In a Mendoza Malbec vineyard in Argentina, pH increase start was monitored over 10 days in late January Year N+1 (BBCH 83) when pH rose from 2.92 to 3.16 (0.24 unit increase, 0.024 units per day average), titratable acidity declined from 19.8 to 15.6 g/L (21% reduction), malic acid decreased from 10.8 to 8.2 g/L, tartaric acid stable 7.4 g/L, berry K⁺ increased from 1650 to 2120 mg/kg, under hot high-altitude conditions (daily high 32-36°C accelerating acid decline and pH rise).
pH_Increase_Start represents the beginning elevation of berry juice pH from green berry minimum values toward ripening equilibrium resulting from net organic acid decline and proton consumption creating the fundamental shift in acidity that affects wine style, microbial stability, and sensory characteristics. This pH increase begins during veraison onset (BBCH 83), coinciding with acid decline onset, and is characterized by pH rising from pre-veraison minimum 2.7-3.1 (cultivar and maturity dependent) to early increase 2.9-3.3 over 7-12 days representing 0.15-0.30 unit elevation, corresponding titratable acidity declining from 17-25 g/L to 14-21 g/L (15-25% reduction) as organic acid anions removed, and buffer capacity adjusting as acid composition shifts from predominantly malate toward greater tartrate proportion affecting pH-acidity relationship. Biochemical basis of pH increase includes: organic acid decline particularly malate removal (8-14 g/L declining to 7-11 g/L) reducing total titratable hydrogen ions, malate as dicarboxylic acid (pKa₁ 3.40, pKa₂ 5.11) existing predominantly as divalent anion malate²⁻ at berry pH 2.8-3.2 contributing two titratable protons per molecule, tartaric acid (pKa₁ 2.98, pKa₂ 4.34) similarly diprotic but metabolically stable showing only dilutional decline, and pH calculated from Henderson-Hasselbalch equation pH = pKa + log([A⁻]/[HA]) where declining acid concentrations shift equilibrium toward higher pH. Proton balance considerations show: organic acid anions (predominantly malate²⁻ and tartrate²⁻) serving as conjugate bases accepting protons maintaining acidic pH, malate removal through respiratory decarboxylation eliminating acid equivalents requiring compensatory ion adjustments, cation accumulation particularly potassium (rising from 1200-1800 to 1800-2500 mg/kg during pH increase) potentially affecting pH through charge balance though mechanisms debated, and vacuolar proton-ATPase activity potentially adjusting to maintain appropriate vacuolar pH despite changing acid concentrations. Buffer capacity changes during pH increase demonstrate: green berry with high organic acid content (20-30 g/L) showing strong buffer capacity requiring ~15-25 mL of 0.1 N NaOH per 10 mL juice to reach pH 8.2, early pH increase phase with declining acids (15-22 g/L) reducing buffer capacity to ~12-18 mL NaOH indicating less buffering against pH changes, and acid composition effects with malate (pKa₂ 5.11) providing stronger buffering near berry pH compared to tartrate (pKa₂ 4.34) so preferential malate loss reduces buffer capacity disproportionately. Cellular mechanisms underlying pH increase inclu vacuolar pH directly affecting juice pH with vacuole occupying 95-97% of mesocarp cell volume, tonoplast proton pumps (V-ATPase, V-PPase) potentially modulating proton content though activity likely reduced during ripening as vacuole transitions from acid accumulation to storage/dilution function, organic acid transport from vacuole to cytoplasm for metabolism requiring tonoplast carriers (possibly VvALMT family) affecting vacuolar acid pool, and apoplastic pH likely increasing parallel with vacuolar pH though measurements technically challenging in berry tissue. Temporal coordination of pH increase with other processes shows: pH rise beginning 0-2 days concurrent with acid decline onset indicating direct causal relationship, sugar accumulation proceeding simultaneously with pH increase creating developing balance between sweetness and acidity, berry softening and color development (reds) occurring during pH increase reflecting integrated ripening program, and pH trajectory paralleling acid decline throughout ripening until harvest with both approaching cultivar-specific equilibria. Environmental factors affecting pH increase rate inclu temperature strongly influencing through effects on acid metabolism with warm conditions 28-32°C accelerating pH rise 0.02-0.04 units per week faster than cool 20-24°C (0.012-0.025 units per week) through enhanced malate catabolism, heat stress >32°C causing rapid pH increase potentially 0.04-0.07 units per week producing excessive rise, and diurnal temperature effects with warm days promoting acid loss and pH increase while cool nights moderating overall change. Potassium effects on pH remain somewhat controversial: some research showing strong correlation between berry K⁺ accumulation (1500-2500 mg/kg) and pH increase suggesting K⁺ as key factor, alternative view that K⁺ accumulates passively responding to organic acid anion loss without causal role in pH, and practical observations that high-K⁺ grapes generally show higher pH though causation versus correlation unclear, with truth likely involving K⁺ affecting charge balance and potentially facilitating acid transport/metabolism rather than K⁺ directly determining pH. Cultivar variation in pH increase characteristics shows: low-pH cultivars (Riesling, Chenin Blanc potential final pH 3.0-3.3) exhibiting slow increase 0.012-0.020 units per week early phase, moderate varieties (Chardonnay, Cabernet Sauvignon final pH 3.3-3.7) displaying 0.018-0.028 units per week, and high-pH types (Grenache, Viognier final pH 3.5-3.9) showing rapid increase 0.025-0.040 units per week reflecting genetic differences in acid metabolism and retention. Wine quality implications of pH inclu pH critically affecting wine character with appropriate 3.1-3.6 range (white 3.0-3.4, red 3.3-3.7) producing balanced wines, low pH 3.7 producing flat wines lacking structure with reduced color stability (reds), increased oxidation susceptibility, and higher spoilage risk from bacteria and yeast, and pH affecting malolactic fermentation with optimal 3.3-3.7 for bacterial activity. Color effects of pH increase in red grapes show: anthocyanin color forms pH-dependent with flavylium cation (red, protonated) dominant at pH 4.0, so pH increase from 3.0 to 3.4 during ripening shifting color from bright red toward purple-red affecting visual quality, and copigmentation effects also pH-sensitive with optimal color enhancement at pH 3.2-3.6 range typical during late ripening. Microbial stability considerations inclu pH 3.5 high risk with aggressive antimicrobial management needed, and harvest timing decisions sometimes prioritizing pH control over sugar accumulation in warm vintages where excessive pH rise threatens wine stability. Analytical measurement of pH employs: calibrated glass electrode pH meter providing accurate measurement ±0.01-0.02 units, temperature compensation essential as pH varies ~0.02 units per °C, sample preparation requiring clarification by filtration or centrifugation preventing particulate interference, and correlation with titratable acidity with typical pH 3.2-3.4 corresponding to TA 6-9 g/L though relationship varies with acid composition. Viticultural management affecting pH includes: canopy practices controlling berry temperature with shading reducing warming 3-7°C slowing pH increase 20-35% through moderated acid decline, irrigation potentially affecting through cooling or dilution effects though mechanisms complex, crop load optimization with moderate yields potentially moderating pH rise compared to overcropped vines showing accelerated acid loss, rootstock selection with some stocks associated with lower berry pH though mechanisms (altered K⁺ uptake, vine vigor effects, other) debated, and harvest timing critically important as pH continues increasing throughout ripening requiring balance with other quality parameters.
pH_Increase_Start is distinguished from Acid_Decline_Onset by representing pH effect rather than acid cause, from later pH changes by showing onset (0.15-0.30 unit rise) rather than complete elevation, and from buffer capacity by focusing on pH measurement rather than acid-base chemistry. Rate: 0.012-0.040 units per week (cultivar/temperature dependent). Duration 7-12 days BBCH 83. Temperature sensitive: warm accelerates, >32°C excessive. Target final pH: whites 3.0-3.4, reds 3.3-3.7. pH_Increase_Start represents critical transition affecting wine style, stability, and quality.
Azidität-Reduktions-Initiierung
Beeren-pH-Erhöhungs-Onset
Protonen-Abbau-Start
Saft-pH-Anstiegs-Beginn
acidity reduction initiation
berry pH elevation onset
juice pH rise beginning
proton depletion start
Keller M (2020) The Science of Grapevines 3rd Ed pages 231-233 pH changes during ripening Boulton RB, Singleton VL, Bisson LF, Kunkee RE (1996) Principles and Practices of Winemaking pages 38-45 juice pH and acidity Mpelasoka BS, Schachtman DP, Treeby MT, Thomas MR (2003) A review of potassium nutrition in grapevines with special emphasis on berry accumulation Aust J Grape Wine Res 9:154-168 potassium and pH relationships.
pH Increase Start
pH measurement for Riesling from the Mosel (2024 vintage) records pH 2.95 at 78° Oechsle harvest, an exceptionally low value characteristic of cool-climate Riesling on slate soils with limited potassium availability, providing outstanding natural microbial protection (molecular SO₂ efficacy maximized at low pH) and brilliant color preservation essential for the pale gold appearance of Mosel Kabinett. pH measurement for Cabernet Sauvignon from Napa Valley (2023 vintage) documents pH 3.68 at 25.5° Brix, near the upper advisory threshold (pH 3.70) above which increased molecular SO₂ requirements, elevated microbial risk, and compromised color stability prompt many winemakers to consider tartaric acid addition of 1-2 g/L during fermentation. pH measurement for Blaufränkisch from Leithaberg (2024 vintage) records pH 3.28 at 19.5° KMW, within the optimal range (3.2-3.5) for Austrian dry red wine production, indicating well-balanced acid-cation equilibrium on the region's limestone-influenced soils.
pH measurement is an acidity data entity that records the hydrogen ion activity in grape must as determined by potentiometric measurement using a calibrated glass electrode pH meter, expressed on the logarithmic pH scale (dimensionless units), representing the thermodynamic acid-base equilibrium state of the must system that integrates the contributions of all acid and base species, buffer capacity, and cation concentrations (particularly potassium), and serving as the single most consequential chemical parameter for predicting wine microbial stability, color expression, SO₂ efficacy, protein stability, and organoleptic acid perception.
pH measurement is arguably the single most important number in must and wine analysis because it governs a cascade of chemical and microbiological equilibria that collectively determine wine quality and stability. The molecular form of sulfur dioxide (the antimicrobial active species) exists as a pH-dependent equilibrium — at pH 3.0 approximately 6% of free SO₂ is in molecular form versus only 1.5% at pH 3.5, meaning that wines at higher pH require proportionally more total SO₂ to achieve equivalent microbial protection. Anthocyanin color expression is similarly pH-dependent — the red flavylium cation form predominates below pH 3.0 while the colorless carbinol pseudobase increases above pH 3.5, making pH a direct determinant of red wine color intensity. The relationship between pH and titratable acidity is non-linear and modulated by the buffer capacity of the must, which depends on the relative concentrations of tartaric acid, malic acid, and potassium — two musts with identical TA can differ by 0.3-0.5 pH units depending on their potassium content and acid species composition, with profound implications for wine chemistry and sensory perception. Potassium is the dominant must cation influencing pH through the formation of potassium bitartrate (KHT), which partially neutralizes tartrate acidity and raises pH. The measurement methodology requires careful attention to electrode calibration (two-point calibration at pH 4.01 and 7.01 using certified buffer solutions), temperature compensation (pH varies approximately -0.003 units/°C for must), and electrode maintenance (junction potential drift from protein fouling in must). The trend toward higher must pH in many viticultural regions under climate change (driven by increased potassium uptake, accelerated malic acid respiration, and reduced tartrate synthesis) represents one of the most significant quality challenges facing contemporary winemaking.
Säure-Base-Messung
Wasserstoffionenkonzentration
acid-base measurement
hydrogen ion concentration
must pH
pH reading
pH value
pH-Bestimmung
pH-Messung
pH-Wert
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2, 2nd Ed, Wiley, pp.: 3-15 Waterhouse AL et al. (2016) Understanding Wine Chemistry, Wiley, pp.: 15-35 OIV (2021) Compendium of International Methods of Analysis, Method OIV-MA-AS313-15 Iland P et al. (2011) Chemical Analysis of Grapes and Wine, 2nd Ed, pp.: 60-68
pH Measurement
pH quality management in Barossa Valley Shiraz requires careful attention because warm-climate ripening typically produces harvest pH of 3.5-3.8 that challenges color stability and microbial safety winemakers target prefermentation pH adjustment to 3.4-3.5 through tartaric acid addition (1-2 g/L addition lowers pH by approximately 0.1 units), enabling the deep, stable purple-black color and resistance to Brettanomyces that distinguish premium Barossa Shiraz, while excessively high pH (above 3.8) necessitates more aggressive acidification or blending with lower-pH components to achieve microbiologically and oenologically sound wines.
pH_Quality represents the chemical quality characterizing the hydrogen ion activity in grape juice or wine, expressed as the negative logarithm of hydrogen ion concentration (pH = -log[H⁺]). Unlike titratable acidity which measures total acid concentration, pH measures the intensity of acidity through free hydrogen ion activity, providing a direct indicator of the chemical environment that governs acid-base equilibria, enzyme activities, microbial growth, and color expression. Grape juice pH typically ranges from 2.9-3.9 at harvest, with ripe wine grapes usually falling between 3.1-3.8 depending on cultivar, climate, and ripening conditions. Lower pH (higher hydrogen ion activity) creates more acidic conditions that enhance wine color stability (anthocyanin-flavylium equilibrium favors red forms), inhibit spoilage bacteria, promote sulfur dioxide effectiveness, and produce tarter taste perception. Higher pH reduces color intensity, increases microbial spoilage risk, requires higher sulfur dioxide additions for equivalent protection, and may produce flat-tasting wines lacking freshness. The pH of grape juice is determined by the concentration and dissociation behavior of all acids present, particularly tartaric and malic acids, as well as the concentration of potassium and other cations that form salts with organic acids. Potassium content significantly affects pH high-potassium berries produce higher pH juice at equivalent titratable acidity because potassium tartrate and malate are less dissociated than the free acids. During winemaking, pH typically increases by 0.1-0.3 units due to precipitation of potassium bitartrate and completion of malolactic fermentation. Measurement employs calibrated glass electrodes with temperature compensation, following standardized protocols that specify calibration procedures and measurement conditions.
pH_Quality captures hydrogen ion activity as distinct from titratable acidity (total acid concentration). The logarithmic nature of pH means that one unit change represents ten-fold change in hydrogen ion activity. The practical range for grape and wine pH (approximately 2.8-4.0) spans more than ten-fold variation in hydrogen ion activity, with significant implications for wine chemistry and stability. The relationship between pH and titratable acidity depends on buffer capacity and acid composition modifications that change one parameter affect the other in complex ways that require empirical determination. The temperature dependence of pH measurement requires standardized conditions for comparable results.
acid intensity
juice pH
pH-Qualität
Boulton RB et al. (1999) Principles and Practices of Winemaking, Springer pp.: 65-78 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed, John Wiley and Sons pp.: 3-25
pH Quality
Total phenolic content in grape berries ranges from 200-500 mg/100g fresh weight in white cultivars to 800-2500 mg/100g in red cultivars, with anthocyanins (100-1500 mg/100g skins), proanthocyanidins (100-800 mg/100g skins and seeds), and flavonols (10-100 mg/100g) as major flavonoid classes. Cultivar differences in phenolic profiles are substantial, with Tannat producing 2-3× higher total phenolics than Pinot Noir under equivalent conditions. The phenylpropanoid pathway converts phenylalanine to p-coumaroyl-CoA, which enters branching pathways toward flavonoids (via chalcone synthase) or stilbenes (via stilbene synthase). Phenolic extractability into wine ranges from 20-60% depending on compound class, berry maturity, and winemaking technique.
A quality potential disposition that is the inherited capacity of a Vitis vinifera plant to synthesize, accumulate, and modify phenolic compounds in grape berries, encompassing flavonoid pathways (anthocyanins, tannins, flavonols) and non-flavonoid pathways (hydroxycinnamic acids, stilbenes) that determine wine color, astringency, bitterness, antioxidant capacity, and aging potential.
Phenolic_Compound_Disposition serves as the parent class for specific phenolic compound categories. Phenolics are the primary determinants of red wine color and structure, while also contributing to white wine complexity and stability. The trait complex shows strong genotype effects overlaid with substantial environmental and management modulation. Climate, water status, light exposure, and ripeness stage all influence phenolic accumulation and composition, creating the foundation for terroir expression in wine phenolic profiles.
Phenolstoffwechsel
Polyphenol-Disposition
phenolic accumulation disposition
phenolic synthesis capacity
phenolische Verbindungen
polyphenol trait
Kennedy JA (2008) Grape and wine phenolics: observations and recent findings. Ciencia e Investigación Agraria 35:107-120 Teixeira A et al. (2013) Berry phenolics of grapevine under challenging environments. International Journal of Molecular Sciences 14:18711-18739
Phenolic Compound Disposition
Phenolic content quality profiling in Barossa Valley Shiraz employs comprehensive analysis including total phenolics (1800-2500 mg/L gallic acid equivalents in finished wine), anthocyanins (400-800 mg/L), tannins (1200-1800 mg/L), and phenolic maturity indices (seed tannin contribution below 40%) to guide harvest timing and winemaking protocols fruit achieving target phenolic profiles receives extended maceration (21-28 days) to maximize extraction, while phenolically immature fruit (high seed tannin proportion, low anthocyanin) is processed with shorter maceration to avoid harsh, green tannin character.
Phenolic_Content_Quality encompasses all chemical qualities that characterize the concentration, composition, and structural diversity of phenolic compounds in grapevine tissues. Phenolics constitute one of the most complex and quality-relevant compound classes in grapes and wine, responsible for color, astringency, bitterness, antioxidant capacity, and numerous health-associated properties. The phenolic category includes hundreds of individual compounds sharing the common structural feature of one or more aromatic rings bearing hydroxyl groups, organized into major subclasses based on carbon skeleton structure. This category subdivides into Total_Phenolic_Quality addressing aggregate phenolic measurements, Flavonoid_Quality encompassing the C6-C3-C6 flavonoid compounds including anthocyanins, flavonols, and flavan-3-ols, Non_Flavonoid_Quality capturing simpler phenolics including hydroxycinnamic acids and stilbenes, and Tannin_Quality addressing the polymerized phenolics responsible for astringency. Phenolic compounds are synthesized through the phenylpropanoid pathway from phenylalanine, with branch points leading to the diverse structural classes. In grape berries, phenolics accumulate primarily in the skin and seeds, with distinct compound profiles in each tissue compartment. Skin phenolics include anthocyanins (in colored cultivars), flavonols, and hydroxycinnamic acids, while seeds contain primarily flavan-3-ols and their polymers (condensed tannins). The concentration and composition of phenolics vary dramatically among cultivars, representing a major dimension of varietal character distinguishing tannic Cabernet Sauvignon from soft Pinot Noir or aromatic Muscat. Environmental factors including light exposure, temperature, and water status modulate phenolic accumulation, with generally positive effects of moderate stress on phenolic concentration. Phenolic extraction during winemaking depends on maceration conditions, making grape phenolic content a potential indicator rather than direct predictor of wine phenolic composition.
Phenolic_Content_Quality is positioned as a direct subclass of Chemical_Quality, parallel to Sugar_Content_Quality, Acid_Content_Quality, and other composition categories. The enormous structural diversity of phenolic compounds creates analytical challenges different methods capture different compound subsets, requiring careful specification of analytical approach for reproducible results. The distinction between grape phenolic content (potential) and wine phenolic composition (realized) reflects the critical role of extraction during winemaking. Phenolic qualities describe current compositional state distinct from phenolic-related dispositions (e.g., extractability) and processes (e.g., phenolic biosynthesis, polymerization).
Phenolgehalt
Polyphenolgehalt
Polyphenolqualität
phenolic composition
phenolic compound quality
phenolische Zusammensetzung
polyphenol content
polyphenolic quality
Kennedy JA et al. (2006) Grape and Wine Phenolics: History and Perspective. American Journal of Enology and Viticulture 57(3): 239-248 Downey MO et al. (2006) Cultural Practice and Environmental Impacts on the Flavonoid Composition of Grapes and Wine: A Review of Recent Research. American Journal of Enology and Viticulture 57(3): 257-268
Phenolic Content Quality
Phenolic data for Cabernet Sauvignon from Coonawarra (2023 vintage) integrates total phenolics (72 AU280 nm by Iland method), total anthocyanins (2.05 mg/g berry fresh weight), extractable anthocyanins (1.65 mg/g at pH 3.2, representing 80% extractability), seed tannin (3.8 mg/g catechin equivalents), skin tannin (1.2 mg/g), and color intensity (CI 18.5 AU) — the high anthocyanin extractability and favorable skin-to-seed tannin ratio indicating full phenolic maturity optimal for premium red wine production. Phenolic data for Blaufränkisch from Deutschkreutz (2024 vintage) records total phenolics 58 AU280, anthocyanins 2.8 mg/g (exceptionally high for the variety), tannin 2.2 mg/g, and CI 16.2 AU, confirming the clonal characteristic of intense pigmentation in the Bl-2 selection.
Phenolic data is a must analysis data entity that documents the concentration, composition, and extractability of phenolic compounds in grape must, berries, and seeds, encompassing total phenolics as the aggregate spectrophotometric measure, anthocyanins as the pigment fraction, tannins as the polymeric flavanol fraction, and color intensity as the integrated chromatic expression, measured through spectrophotometric, chromatographic, and chemical precipitation methods on must or homogenized berry samples and providing the phenolic maturity assessment that, together with sugar and acid data, defines the three-dimensional ripeness framework for red and rosé wine production decisions.
Phenolic data has become the essential complement to sugar and acid analysis in modern red wine grape assessment, reflecting the recognition that phenolic maturity — the accumulation, polymerization, and extractability of skin and seed phenolics — follows a temporal trajectory that may or may not align with sugar maturity. The "phenolic maturity" concept, formalized by Glories (1984) and developed by Saint-Cricq de Gaulejac (1998), distinguishes between total phenolic content (what is present in the berry) and extractable phenolic content (what can be released under winemaking conditions), with the ratio between these values providing an index of cell wall degradation and phenolic accessibility. The Glories method assesses anthocyanin extractability by comparing anthocyanin levels extracted at pH 1.0 (total, by complete cell disruption) versus pH 3.2 (simulating winemaking extraction conditions), with extractability percentages above 70% indicating advanced phenolic maturity. Seed tannin evolution during ripening follows a distinctive pattern — total seed tannin content decreases as berries mature while seed tannin astringency simultaneously declines due to oxidative polymerization and reduced extractability from lignifying seed coats, creating a "tannin maturity" parallel to the anthocyanin extractability index. The analytical methods span rapid spectrophotometric screening (total phenolics by absorbance at 280 nm, color intensity by A420+A520+A620) through specific quantification (anthocyanins by pH-shift or HPLC, tannin by methylcellulose precipitable tannin or protein precipitation), enabling different levels of analytical detail appropriate to quality tier and decision requirements.
Gerbstoffdaten
Phenoldaten
Phenolreifedaten
Polyphenoldaten
grape phenolics data
phenolic analysis data
phenolic composition data
phenolic maturity data
phenolische Zusammensetzungsdaten
polyphenol data
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2, 2nd Ed, Wiley, pp.: 141-203 Iland P et al. (2011) Chemical Analysis of Grapes and Wine, 2nd Ed, pp.: 88-120 Saint-Cricq de Gaulejac N et al. (1998) Phenolic composition of grape berries at different stages of fruit development. Acta Horticulturae 464: 159-164
Phenolic Data
In Cabernet Sauvignon berries in Napa Valley, California during BBCH 81-89, phenolic maturation establishes total anthocyanin content increasing from 0.2 to 1.85 mg/g fresh weight over 48 days, skin tannin concentration rising from 1.2 to 3.8 mg/g skin, and seed tannin declining from 8.5 to 4.2 mg/g seed through oxidative browning, with total phenolic index increasing from 45 to 72 absorbance units while tannin-to-anthocyanin ratio decreases from 6.2 to 2.8 indicating progressive color and structural balance under warm conditions averaging 28°C.
Phenolic maturation constitutes the comprehensive ripening-phase process in Vitis vinifera berries wherein phenolic compounds including anthocyanins, proanthocyanidins, flavonols, and hydroxycinnamic acids undergo quantitative accumulation, structural transformation, and qualitative refinement during the 35-50 day post-veraison period, establishing the phenolic foundation that determines red wine color stability, mouthfeel characteristics, astringency evolution, and aging potential in wines produced from these grapes. This process integrates multiple biosynthetic and modification pathways operating in distinct berry tissues, with anthocyanin accumulation proceeding in exocarp hypodermal cells where flavonoid pathway enzymes show coordinated expression generating the red-blue-purple pigments responsible for berry and wine color, proanthocyanidin polymerization occurring in both skin and seed tissues producing the condensed tannins that contribute astringency and structure, and flavonol and hydroxycinnamic acid accumulation providing copigmentation cofactors and antioxidant capacity that modulate color expression and wine stability. The temporal progression of phenolic maturation shows compound-class-specific dynamics with anthocyanin accumulation initiating at veraison and continuing throughout ripening in red varieties, seed tannin biosynthesis largely complete before veraison with subsequent maturation involving oxidative browning and polymerization changes, and skin tannin accumulation continuing post-veraison with progressive structural modifications affecting extractability and sensory properties. Cellular localization demonstrates tissue-specific phenolic distribution with anthocyanins exclusively in vacuoles of hypodermal skin cells, seed tannins concentrated in outer integument and inner seed coat tissues, and skin tannins distributed across multiple exocarp cell layers with concentration gradients affecting extraction during winemaking. Temperature during ripening profoundly influences phenolic maturation through effects on biosynthetic enzyme activities, compound stability, and structural modification rates, with moderate temperatures of 20-28°C generally supporting optimal phenolic development while temperature extremes may limit anthocyanin accumulation through thermal degradation or reduce tannin quality through accelerated oxidation. Cultivar genetics fundamentally determine phenolic maturation potential through variety-specific expression of flavonoid pathway genes, with high-color varieties such as Alicante Bouschet and Petit Verdot accumulating anthocyanins to 2000-4000 mg/kg compared to lighter varieties such as Pinot Noir reaching 400-800 mg/kg, and thick-skinned varieties accumulating substantially higher skin tannin concentrations than thin-skinned cultivars under equivalent conditions. The relationship between phenolic maturation and wine quality reflects the fundamental importance of adequate color intensity, balanced astringency, and stable phenolic structure for red wine sensory appeal and longevity, with harvest timing decisions targeting optimal phenolic development frequently diverging from decisions based solely on sugar-acid balance, necessitating integrated assessment of multiple ripeness parameters. Viticultural practices influence phenolic maturation through effects on vine physiology, cluster microclimate, and berry composition, with moderate water deficit enhancing anthocyanin accumulation, appropriate crop load ensuring adequate phenolic concentration, and optimized light exposure promoting flavonoid pathway expression without inducing thermal degradation of accumulated pigments.
Phenolic_Maturation is parent class under Ripening representing all phenolic compound developmental processes. Contains subclasses Anthocyanin_Accumulation, Tannin_Polymerization, Skin_Phenolic_Development, and Seed_Phenolic_Maturation. Duration 35-50 days post-veraison. Integrates pigment accumulation, tannin modification, and copigmentation development. Critical for red wine quality assessment. Tissue-specific processes in skin and seed compartments.
Phenol-Verbindungs-Reifung
Tannin-und-Anthocyan-Entwicklung
phenolic compound maturation
phenolic quality development
phenolic ripening
phenolische Qualitäts-Entwicklung
phenolische Reifung
tannin and anthocyanin development
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 310-340 Kennedy JA et al (2006) Grape and wine phenolics. In Chemistry and Biology of Ellagitannins World Scientific pp 220-255 Adams DO (2006) Phenolics and ripening in grape berries. American Journal of Enology and Viticulture 57: 249-256
Phenolic Maturation
Phenolic maturity assessment in Cabernet Sauvignon comparing early and optimal harvest reveals the phenolic dimension of quality decisions, with early harvest (23 Brix, 35 days post-véraison) showing high total anthocyanins (1.9 mg/g) but low extractability (38%) and immature tannins (seed tannin contribution 65%, astringent character), while optimal phenolic harvest (25 Brix, 50 days post-véraison) shows lower total anthocyanins (1.5 mg/g) but higher extractability (62%) and mature tannins (seed contribution reduced to 35%, softer mouthfeel), demonstrating how phenolic maturity assessment identifies the harvest point where extractable quality exceeds that achievable from grapes with higher total phenolic content but less favorable developmental state.
Phenolic maturity quality is a maturity quality that represents the developmental state of phenolic compounds in grape berries, characterizing the accumulation, polymerization, and extractability of anthocyanins, tannins, and flavonols that determine wine color, structure, mouthfeel, and aging potential, with phenolic maturity increasingly recognized as the quality-limiting maturity dimension that may not coincide with technological maturity and requires separate assessment for premium wine production. Within the BFO framework, this quality specifically addresses the phenolic compound dimension of maturity, distinguishing the developmental state of these secondary metabolites from sugar or acid parameters while recognizing that phenolic maturity often determines the difference between good and great wines, as color stability, tannin quality, and phenolic balance contribute decisively to wine character and longevity. The ontological significance lies in phenolic maturity determining the structural foundation and color stability of red wines, where the quantity of accumulated anthocyanins establishes color potential, the degree of tannin polymerization affects astringency and mouthfeel, and the extractability of these compounds during winemaking determines how much of the grape's phenolic potential is realized in the wine. The biosynthetic basis for phenolic accumulation involves the phenylpropanoid pathway synthesizing flavonoid precursors that are subsequently modified into anthocyanins (in skin cells), proanthocyanidins/tannins (in skin and seed), and flavonols (primarily in skin), with accumulation patterns varying among compound classes and affected by genetics, temperature, light exposure, and water status during ripening. The decoupling phenomenon between phenolic and sugar maturity is particularly relevant under climate change, where elevated temperatures may accelerate sugar accumulation while delaying or disrupting phenolic development, creating situations where grapes reach target Brix while phenolics remain immature (harsh tannins, unstable color), requiring either extended hang time with consequent alcohol increase or harvest at lower sugar with phenolic compromise. Understanding phenolic maturity quality enables assessment of the structural and color potential of grapes beyond simple compositional measures, harvest timing that considers phenolic development alongside technological parameters, and recognition of the cultivar-specific and climate-dependent patterns of phenolic maturation that affect wine quality potential.
Phenolic_Maturity_Quality serves as the parent class for phenolic compound-specific qualities (Anthocyanin, Tannin, Extractability). This quality addresses the structural and color-determining compounds critical for red wine quality. The class supports premium wine harvest decisions through phenolic maturity characterization beyond technological parameters.
Polyphenolreife
Strukturreife
phenolic ripeness
phenolische Reife
polyphenol maturity
structural maturity
Río Segade S et al. (2008) Phenolic ripeness assessment of grape skin by texture analysis. Journal of Food Composition and Analysis 21(8): 644-649 Kennedy JA et al. (2002) Development of seed polyphenols in berries from Vitis vinifera L. cv. Shiraz. Australian Journal of Grape and Wine Research 8(1): 25-35
Phenolic Maturity Quality
In Touriga Nacional berries in Douro Valley, Portugal at BBCH 89, phenolic seed maturity is achieved with seed color score of 3.9 on 4-point scale indicating uniform dark brown appearance, seed texture crisp with clean separation from pulp, extractable tannin reduced to 3.2 mg/g seed representing 38% of veraison content, mDP at 7.5, and sensory evaluation confirming soft nutty character without harsh bitterness, establishing optimal seed phenolic status after 48 days post-veraison under warm continental-Mediterranean conditions averaging 28°C.
Phenolic seed maturity represents the developmental endpoint of seed phenolic evolution in Vitis vinifera grape seeds wherein oxidative modification has progressed sufficiently to establish mature seed phenolic status characterized by uniform brown coloration, substantially reduced tannin extractability of 30-50% of initial levels, and modified astringency potential indicating seeds that will contribute minimally to harsh wine tannin character when grapes are processed for red wine production at optimal harvest timing. This state integrates multiple maturity indicators including visual assessment of seed coat color with mature seeds showing uniform dark brown appearance rated 3.5-4.0 on standard scales, tactile evaluation with mature seeds demonstrating crisp texture and easy separation from surrounding pulp tissue, analytical measurements confirming reduced extractable tannin content typically below 4-5 mg/g seed compared to 8-12 mg/g in immature seeds, and sensory evaluation of seed tannin astringency through chewing tests revealing softened rather than harsh bitter-astringent character in mature seeds. The biochemical characteristics of phenolic seed maturity encompass structural modifications including decreased mean degree of polymerization from initial values of 10-12 to mature values of 6-9 reflecting oxidative cleavage, formation of oxidatively cross-linked polymers with reduced solubility, and potential incorporation of anthocyanins or other phenolics into modified tannin structures through non-enzymatic condensation reactions during extended oxidative modification. The timing of phenolic seed maturity achievement varies among cultivars and environmental conditions, with some varieties reaching mature seed status 35-40 days post-veraison while others may require 50-60 days, and warm conditions generally accelerating maturation compared to cool seasons that may delay seed phenolic development requiring extended hang-time for optimal maturity. The practical assessment of phenolic seed maturity in vineyard sampling employs tactile and visual evaluation through the established protocol of cutting berries to expose seeds, evaluating color and texture, and potentially tasting seeds to assess astringency, with experienced viticulturists capable of reliably determining seed maturity status through these sensory methods that correlate with analytical measurements. The enological significance of phenolic seed maturity relates to wine quality optimization, with harvest of grapes possessing mature seeds reducing the risk of harsh tannin extraction during maceration, contributing to wines with balanced structure and approachable astringency compared to wines produced from grapes harvested before optimal seed maturation when aggressive seed tannins may compromise sensory quality.
Phenolic_Seed_Maturity is subclass of Seed_Phenolic_Maturation representing the achieved quality endpoint. Distinguishes from process-focused classes by emphasizing maturity status assessment. Sibling to Seed_Tannin_Polymerization and Seed_Browning. Integrates visual, tactile, analytical, and sensory indicators. Extractable tannin typically below 4-5 mg/g in mature seeds. Harvest timing criterion for quality wine production.
Samen-Tannin-Reife
Samen-phenolische-Reife
mature seed phenolic status
optimal seed phenolic development
phenolische Samenreife
reifer Samen-phenolischer-Status
seed phenolic ripeness
seed tannin maturity
Ristic R and Iland PG (2005) Relationships between seed and berry development of Vitis vinifera L. Australian Journal of Grape and Wine Research 11: 43-58 Kennedy JA et al (2001) Changes in grape seed polyphenols during ripening. Phytochemistry 55: 77-85 Adams DO (2006) Phenolics and ripening in grape berries. American Journal of Enology and Viticulture 57: 249-256
Phenolic Seed Maturity
A viticulturist recording BBCH 65 for Riesling vines in the Rheingau on June 15 communicates an unambiguous phenological observation — full flowering with 50% of flowerhoods fallen — that is directly comparable with BBCH 65 observations of Riesling in Marlborough, Alsace, or the Wachau despite differences in calendar date, climate, and management, enabling standardized phenological monitoring networks such as the European Phenology Network and INIA's Spanish Phenological Network to aggregate cross-regional development data using a shared coding framework.
A grapevine information entity that provides a standardized alphanumeric encoding of a grapevine's developmental stage, growth phase, or seasonal physiological state within a formal phenological classification system, enabling precise and reproducible communication of vine development status across observers, regions, and research programs.
Phenological_Code provides the standardized vocabulary for describing grapevine development that underpins phenological research, precision viticulture scheduling, and climate change impact assessment. The dominant coding system for grapevine phenology is the BBCH (Biologische Bundesanstalt, Bundessortenamt und Chemische Industrie) scale adapted for Vitis by Lorenz et al. (1995), which extends the general BBCH framework to grapevine-specific developmental stages: subclasses capture both the BBCH system and its hierarchical components including principal growth stages and secondary growth stages. The BBCH code translates subjective visual observation into standardized numerical expression, enabling quantitative analysis of phenological timing and its response to climate, cultivar, and management variables.
Entwicklungsstadien-Code
Phänologie-Kennung
developmental stage notation
growth stage identifier
phenological stage code
phänologischer Code
Lorenz DH et al. (1995) Growth stages of the grapevine: phenological growth stages of the grapevine — Codes and descriptions according to the extended BBCH scale, Australian Journal of Grape and Wine Research 1: 100-103 Meier U (2001) Growth Stages of Mono- and Dicotyledonous Plants, BBCH Monograph 2nd Ed, Federal Biological Research Centre
Phenological Code
Phenological plasticity varies substantially across V. vinifera cultivars, with high-plasticity cultivars such as Riesling showing 3-4 day advancement in veraison per 1°C increase in growing season temperature, while low-plasticity cultivars such as Ugni Blanc show only 1.5-2 day response to the same temperature change. Plasticity indices calculated from multi-site trials range from 0.5 (low responsiveness) to 2.5 (high responsiveness) days per degree-day deviation from normal. High plasticity enables cultivar adaptation across diverse climates but increases vintage variation, while low plasticity provides consistent timing but limits climatic range.
A phenological timing disposition that is the inherited capacity of a Vitis vinifera plant to modulate the timing of developmental stages in response to environmental variation, determining the degree to which phenological events shift under different temperature regimes, photoperiods, or other environmental signals.
Phenological_Plasticity_Disposition serves as the parent class for specific environmental response categories including temperature and photoperiod sensitivity. Plasticity represents the norm of reaction for phenological timing across environmental gradients, distinct from the mean timing under standard conditions. Understanding cultivar-specific plasticity is essential for predicting climate change impacts, as high-plasticity cultivars will show greater phenological shifts under warming scenarios. Plasticity may be adaptive (enabling environmental tracking) or maladaptive (causing phenological mismatches with optimal conditions).
Entwicklungsflexibilität
Umweltreaktivität
developmental flexibility
environmental timing sensitivity
phenological responsiveness
phänologische Plastizität
Sadras VO, Moran MA (2013) Nonlinear effects of elevated temperature on grapevine phenology. Agricultural and Forest Meteorology 173:107-115 Wolkovich EM et al. (2017) From Pinot to Xinomavro in the world's future wine-growing regions. Nature Climate Change 7:789-795
Phenological Plasticity Disposition
Phenological quality assessment comparing early (Chardonnay) and late (Cabernet Sauvignon) cultivars in a warming climate reveals the differential adaptation challenges, with Chardonnay budbreak advancing from historical April 5 to March 20 (15 days earlier), flowering from June 1 to May 18, and harvest from September 15 to August 28, shifting the entire phenological window earlier and into warmer conditions, while Cabernet Sauvignon shows similar advancement but retains later absolute timing, demonstrating how phenological quality differences among cultivars affect climate change vulnerability and the strategic value of late-ripening cultivars for maintaining ripening in favorable temperature windows.
Phenological quality is a quality that represents the developmental state and temporal progression of grapevine through its annual cycle of growth stages, characterizing the timing, duration, and expression of phenological events from dormancy through budbreak, flowering, fruit development, ripening, and return to dormancy that determine the alignment between vine development and seasonal climate patterns critical for successful grape production. Within the BFO framework, this quality addresses the temporal-developmental dimension of vine biology, distinguishing the progression through growth stages from the structural or physiological states at any moment while recognizing that phenology integrates genetic programming with environmental signals (temperature, photoperiod) to coordinate vine development with seasonal opportunities and constraints. The ontological significance lies in phenological quality determining the temporal fit between vine development and climate, where the timing of budbreak affects frost risk, flowering timing affects fruit set conditions, veraison timing determines the ripening window, and the overall phenological pattern establishes whether fruit can achieve target maturity before autumn conditions deteriorate, making phenology a primary determinant of cultivar-site suitability and climate change vulnerability. The regulatory basis for phenological progression involves the integration of chilling accumulation (breaking dormancy), heat accumulation (driving development rate), and photoperiod signals (affecting dormancy induction) with endogenous developmental programs, creating the temperature-responsive phenological patterns that vary predictably with climate while maintaining genetic cultivar identity in relative timing and thermal requirements. The standardized description of grapevine phenology employs the BBCH (Biologische Bundesanstalt, Bundessortenamt und CHemische Industrie) scale adapted for Vitis, providing numerical codes for principal growth stages (0-9) and secondary stages within each principal stage, enabling precise communication of developmental status for research, management, and modeling applications. The climate change implications of phenological quality are substantial, as warming temperatures advance phenological timing, potentially shifting ripening into hotter periods, altering cultivar-climate relationships, and requiring adaptation strategies including cultivar substitution, site relocation, or management interventions to maintain wine quality under changed conditions. Understanding phenological quality enables prediction of developmental timing, assessment of cultivar-climate suitability, and planning of management operations relative to phenological stages critical for their effectiveness.
Phenological_Quality serves as the parent class for developmental stage and rate qualities characterizing vine progression through annual growth cycles. This quality addresses the temporal-developmental dimension integrating genetics with environmental signals. The class supports climate adaptation and management timing through phenological characterization.
Entwicklungszeitpunkt
developmental timing
growth stage quality
jahreszeitliche Entwicklung
phenological status
phänologische Qualität
seasonal development
Coombe BG (1995) Adoption of a system for identifying grapevine growth stages. Australian Journal of Grape and Wine Research 1(2): 104-110 Parker AK et al. (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17(2): 206-216
Phenological Quality
The phenological timing description of Cabernet Sauvignon at Bordeaux (44.8°N, maritime temperate climate, 1,300-1,500 GDD base 10°C) documents late budbreak (approximately 7-10 days after Chasselas reference), late flowering (approximately 5-8 days after Chasselas), late véraison (approximately 8-12 days after Chasselas), and late harvest maturity (late September to mid-October, approximately 10-15 days after Merlot) — this consistently late phenological profile across all stages classifies Cabernet Sauvignon as a late-season cultivar (Période IV in the Pulliat classification) requiring a long, warm growing season for full maturation, explaining its success in Bordeaux, Napa Valley, Coonawarra, and Stellenbosch but its failure to ripen adequately in cooler regions such as Burgundy, the Mosel, and southern England where earlier-maturing cultivars (Pinot Noir, Riesling, Chardonnay) achieve optimal ripeness within the shorter available growing season.
An ampelographic description component that documents the temporal sequence of major developmental stages in the grapevine annual cycle — budbreak, flowering, véraison, and harvest maturity — recording the calendar dates or thermal time (growing degree-day accumulations) at which each cultivar reaches these phenological milestones relative to reference cultivars, with phenological timing providing the fundamental characterization of cultivar precocity or lateness that determines climatic suitability, viticultural management scheduling, and the matching of cultivar maturation requirements to regional growing season length and temperature profiles.
Phenological_Timing_Description addresses the temporal dimension of cultivar characterization that is arguably the most practically important attribute for viticultural site-cultivar matching — the fundamental question of whether a cultivar will ripen adequately in a given climate is answered primarily by comparing the cultivar's phenological timing requirements with the region's growing season length and heat accumulation. The phenological timing of each cultivar is genetically determined but expressed on an absolute calendar scale that varies with latitude, altitude, proximity to moderating water bodies, and annual temperature variation — a cultivar that buds in late March in Bordeaux may bud in early April in Burgundy, mid-April in Champagne, and late February in Barossa Valley, but its relative timing compared to other cultivars at the same site remains consistent: this relativity is the basis of the reference cultivar system, in which Chasselas (syn. Gutedel) has traditionally served as the European benchmark for phenological comparison — cultivars are classified as early, medium, or late relative to Chasselas at each developmental stage, providing a site-independent characterization of precocity. The four phenological stages documented in the subclasses — budbreak, flowering, véraison, and harvest maturity — represent the cardinal points of the annual cycle that divide the growing season into three interphenological periods (budbreak-to-flowering, flowering-to-véraison, véraison-to-harvest) whose durations and temperature profiles determine berry composition at harvest.
Entwicklungszeitpunkt-Charakterisierung
Sortenfrühreife-Dokumentation
cultivar precocity documentation
developmental timing characterization
phenological calendar record
phänologische Zeitpunkte-Beschreibung
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Jones GV and Davis RE (2000) Climate influences on grapevine phenology, grape composition, and wine production and quality for Bordeaux, France, American Journal of Enology and Viticulture 51: 249-261 Duchêne E and Schneider C (2005) Grapevine and climatic changes: a glance at the situation in Alsace, Agronomy for Sustainable Development 25: 93-99 Pulliat V (1888) Mille variétés de vignes, 4th edition, Montpellier
Phenological Timing Description
Phenological timing varies by 4-8 weeks across V. vinifera cultivars under identical conditions, with Chasselas budbreak occurring 10-14 days before Cabernet Sauvignon in the same vineyard. The BBCH scale (Biologische Bundesanstalt, Bundessortenamt und CHemische Industrie) provides standardized phenological staging from BBCH 00 (dormancy) through BBCH 97 (leaf fall). Growing degree day (GDD) accumulation from budbreak to harvest ranges from 1100°C (Müller-Thurgau) to 1800°C (Mourvèdre) using base 10°C. Phenological models predict stage timing with accuracy of ±3-7 days when cultivar-specific parameters are calibrated.
A grapevine disposition that is the inherited capacity of a Vitis vinifera plant to progress through developmental stages (phenophases) at characteristic times relative to environmental cues, determining the temporal pattern of dormancy release, budbreak, flowering, veraison, ripening, and leaf senescence across the annual growth cycle.
Phenological_Timing_Disposition serves as the top-level parent class for all traits related to developmental timing across the grapevine annual cycle. Phenological timing reflects the integration of genetic programming with environmental signals including temperature, photoperiod, and water availability. Climate change is advancing grapevine phenology by 2-3 days per decade in many regions, with implications for cultivar-region matching and wine style. The trait complex shows moderate to high heritability (h² = 0.50-0.75) for most component stages.
Phänologie-Disposition
growth stage timing disposition
jahreszeitliches Timing
phenological development trait
phänologische Entwicklung
seasonal timing capacity
Coombe BG (1995) Growth stages of the grapevine: adoption of a system for identifying grapevine growth stages. Australian Journal of Grape and Wine Research 1:104-110 Parker AK et al. (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17:206-216
Phenological Timing Disposition
Detailed investigation of Phloem_Hardiness_Disposition at Cornell University examined phloem freezing injury patterns and functional consequences across 15 cultivars using controlled freezing, histological assessment, and spring growth monitoring. Dormant cane sections were frozen to target temperatures, sectioned longitudinally through the bark, and scored for phloem browning on a 0-5 scale (0 = no browning, 5 = complete necrosis). LT50 values were determined by probit analysis. Phloem_Hardiness_Disposition ranged from LT50 = -24°C ('Merlot') to LT50 = -37°C ('Frontenac'). Histological examination revealed that initial freezing damage appeared as discrete brown zones in phloem parenchyma at temperatures 2-3°C above lethal threshold, progressing to complete phloem necrosis at and below LT50. Functional assessment tracked carbohydrate mobilization in spring by measuring starch depletion in xylem rays and sugar accumulation in developing buds. Canes with partial phloem damage (browning score 2-3) showed 40% slower starch mobilization and 25% lower bud sugar content at budbreak compared to undamaged controls, correlating with reduced shoot vigor (35% shorter shoots at 30 days post-budbreak). Complete phloem kill (score 5) resulted in bud failure despite apparent bud viability, as carbohydrate supply was severed. These findings demonstrate that Phloem_Hardiness_Disposition determines not only tissue survival but functional capacity for supporting spring growth, with partial damage causing quantitative growth reduction before complete failure threshold.
A Phloem_Hardiness_Disposition is a Cane_Cold_Hardiness_Disposition that inheres specifically in the phloem tissue of dormant grapevine canes and is realized through the freezing tolerance of sieve elements, companion cells, and phloem parenchyma, determining the capacity for carbohydrate transport to support spring growth initiation and early shoot development. This disposition represents the cold hardiness of the primary assimilate transport tissue, with phloem survival essential for mobilizing stored carbohydrates from roots and trunk to developing buds and shoots in spring. Phloem tissue survives freezing through a freeze-tolerance mechanism involving controlled ice formation in extracellular spaces, cellular dehydration as water moves to growing ice crystals, and membrane/protein stabilization through accumulated cryoprotectants (sugars, proline, dehydrins). The relatively thin-walled, metabolically active cells of phloem are generally more cold-sensitive than the thick-walled, largely non-living cells of xylem, establishing phloem as the limiting vascular tissue for cold hardiness in most cultivars. Phloem damage from freezing is visible as browning of the inner bark when canes are cross-sectioned, progressing from scattered brown flecks (minor damage) to complete browning (lethal damage). Even partial phloem damage can restrict carbohydrate supply to developing shoots, causing weak growth, poor fruit set, and delayed canopy development. Phloem_Hardiness_Disposition is typically 3-8°C greater than primary bud hardiness within the same cultivar, providing a buffer that usually protects vascular function when buds are killed, but severe cold events can damage phloem and compromise vine recovery.
Phloem_Hardiness_Disposition is_a Cane_Cold_Hardiness_Disposition. Phloem is typically the most cold-sensitive vascular tissue in canes, setting the functional limit for carbohydrate transport even when xylem water conduction remains intact. Phloem survives freezing through freeze tolerance (controlled extracellular ice formation) rather than supercooling. Partial phloem damage causes growth reduction proportional to damage severity, while complete damage severs carbohydrate supply to distal tissues. Phloem damage is assessed by browning of inner bark in cross-sections.
Basthärte
Phloem-Härte-Disposition
Siebelement-Kältetoleranz
assimilate transport tissue cold resistance
inner bark hardiness
phloem freezing tolerance
Ashworth EN (1984) Xylem development in Prunus flower buds and the relationship to deep supercooling, Plant Physiology 74:862-865 Wolpert JA, Howell GS (1985) Cold acclimation of Concord grapevines II. Natural acclimation pattern and tissue moisture decline, American Journal of Enology and Viticulture 36:189-194 Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235
Phloem Hardiness Disposition
Syrah grafted on SO4 rootstock at 18 years showing phloem inner rootstock bark 2.3 mm thickness comprising secondary phloem tissues, microscopic examination revealing sieve elements with perforated sieve plates and lacking nuclei indicating mature conducting cells, companion cells adjacent to each sieve element showing dense cytoplasm and prominent nuclei, phloem fibers providing mechanical strength, phloem parenchyma and ray parenchyma showing starch storage, aphid stylet technique confirming active phloem sap flow with pressure exudate emerging from punctured sieve elements, phloem demonstrating active food transport from canopy to roots supporting vine growth and function
Phloem inner rootstock bark is inner rootstock bark emphasizing phloem tissue identity and food-conducting function, characterized by secondary phloem produced by vascular cambium comprising sieve elements as primary conducting cells, companion cells metabolically supporting sieve elements, phloem fibers providing mechanical support, phloem parenchyma storing reserves and synthesizing compounds, and ray parenchyma extending radially facilitating lateral transport, demonstrating pressure flow mechanism moving photosynthates from source leaves to sink roots and other consuming tissues, showing sieve element structure with perforated sieve plates enabling cytoplasmic continuity between cells, exhibiting companion cell dense cytoplasm and abundant organelles compensating for reduced sieve element contents, and representing the food-conducting tissue analogous to xylem water transport but moving organic compounds rather than water and minerals enabling vine to distribute photosynthetic products throughout plant body. Phloem inner rootstock bark sieve element characteristics includes= showing elongated cellular structure (sieve elements typically 100-300 micrometers long forming continuous tubes through end-to-end connections), exhibiting perforated sieve plates (end walls with numerous pores creating sieve-like structure enabling cytoplasmic continuity), lacking nucleus at maturity (enucleate cells maintaining metabolic function through companion cell support), demonstrating reduced organelle content (minimal organelles enabling optimal flow through cell), containing specialized P-proteins (phloem proteins potentially functioning in wound sealing and defense), and showing relatively short functional life (sieve elements typically functional 1-3 years before senescence and replacement). Phloem inner rootstock bark companion cell characteristics includes= located adjacent to sieve elements (companion cells derived from same cambial mother cell as associated sieve element), showing dense cytoplasm (abundant ribosomes, mitochondria, and other organelles supporting high metabolic activity), containing prominent nucleus (large active nucleus directing metabolic support of associated sieve element), exhibiting numerous plasmodesmata connections (extensive cytoplasmic connections between companion cell and sieve element enabling metabolic support), demonstrating high respiratory activity (elevated oxygen consumption supporting ATP production for sieve element function), and showing metabolic dependence relationship (sieve element viability requiring companion cell metabolic support creating functional unit). Phloem inner rootstock bark pressure flow mechanism includes= loading photosynthates at source (companion cells actively transporting sugars from mesophyll into sieve elements in leaves), creating osmotic pressure (high sugar concentration in source sieve elements drawing water from xylem creating turgor pressure), enabling bulk flow (pressure gradient between source and sink driving mass flow of phloem sap through sieve tubes), unloading at sinks (sugars transported from sieve elements into sink tissues such as roots, developing fruits, or storage organs), and demonstrating energy requirement (active loading and unloading requiring ATP while flow itself passive following pressure gradient). Phloem inner rootstock bark transport characteristics includes= moving organic compounds (primarily sucrose but also amino acids, hormones, minerals, and other substances), showing bidirectional capacity (phloem capable of reversing flow direction unlike xylem unidirectional transport), exhibiting seasonal variation (transport rates highest during active growth, reduced during dormancy), demonstrating rapid velocity (phloem sap moving 30-150 cm per hour substantially faster than diffusion), and enabling long-distance transport (phloem conducting materials from leaves in scion canopy through graft union to deepest roots meters away). Phloem inner rootstock bark fiber characteristics includes= providing mechanical support (thick-walled elongated cells strengthening bark structure), showing secondary wall thickening (extensive cellulose and lignin deposition creating strong rigid walls), forming fiber bundles (fibers grouped together creating particularly strong reinforcement), contributing to bark flexibility (fibers enabling bark to bend without breaking), and demonstrating longevity (phloem fibers potentially remaining functional in bark for years). Phloem inner rootstock bark parenchyma functions includes= storing carbohydrate reserves (axial and ray parenchyma accumulating starch particularly during dormancy), synthesizing compounds (producing callose, defensive compounds, and other substances), facilitating lateral transport (ray parenchyma creating radial pathways for water and solute movement), enabling wound response (parenchyma cells capable of dedifferentiation and callus formation), and maintaining tissue viability (living parenchyma cells contributing to overall phloem metabolic activity and responsive capacity). Phloem inner rootstock bark ray continuity includes= extending from xylem through cambium into phloem (rays creating continuous radial files connecting wood and bark), facilitating radial transport (ray parenchyma enabling lateral movement of water, nutrients, and signals), showing vascular continuity (rays maintaining connection between xylem and phloem facilitating tissue coordination), demonstrating communication pathways (rays potentially conducting hormonal and other signals radially through trunk), and contributing to storage (ray cells in phloem accumulating reserves complementing axial parenchyma storage).
Phloem_Inner_Rootstock_Bark emphasizes phloem tissue identity and transport function and is distinguished from Living_Inner_Rootstock_Bark (viability emphasis) by focus on phloem-specific anatomy and food conduction. Critical for understanding sieve element-companion cell complex, pressure flow transport, organic compound distribution, phloem anatomy, and the essential food-conducting tissue enabling photosynthate distribution from source leaves to sink organs throughout vine.
Bast tissue
Bastgewebe
Food-conducting bark
Nahrungsleitende Rinde
Phloem-Unterlagenrinde
Phloem Inner Rootstock Bark
Syrah phloem inner trunk bark at 13 years post-planting shows 2.4-3.6 mm depth secondary phloem tissue comprising sieve tube elements 180-350 μm length with specialized sieve plates (pores 2-4 μm diameter, partial callose deposition during winter dormancy), companion cells 60-110 μm length with dense cytoplasm actively loading sucrose maintaining 12-18% sugar concentration in sieve tubes driving 0.4-0.8 m/hour basipetal sap flow, phloem parenchyma storing 20-28 g starch (7-10% dry weight), phloem fibers 450-750 μm length providing mechanical support, and phloem rays 3-5 cells wide connecting radially to xylem rays facilitating lateral transport, functional phloem zone (outer 1.8-2.8 mm) maintaining active transport while transitional crushed zone (inner 0.6-0.8 mm) shows declining function
Phloem inner trunk bark is inner trunk bark emphasizing its identity as secondary phloem tissue produced by vascular cambium, comprising the complex tissue organization of sieve tube elements, companion cells, phloem parenchyma, phloem fibers, and phloem rays that collectively enable long-distance photosynthate transport, nutrient distribution, and storage functions. Phloem inner trunk bark tissue organization includes= sieve tube elements (principal conducting cells, enucleate at maturity with specialized sieve plates in end walls, typical dimensions 150-400 μm length, 15-35 μm diameter, arranged in vertical files creating continuous sieve tubes), companion cells (nucleated cells adjacent to sieve tube elements, 50-120 μm length, maintaining sieve tube function through metabolic support and controlling loading/unloading), phloem parenchyma (living storage and transport cells interspersed among conducting elements, 80-200 μm dimensions, storing starch and participating in lateral transport), phloem fibers (thick-walled elongated supporting cells 400-800 μm length providing mechanical strength to phloem tissue), and phloem rays (radial parenchyma bands 3-5 cells wide extending through phloem connecting to vascular cambium and xylem rays). Phloem inner trunk bark sieve tube element characteristics includes= enucleate at maturity (nucleus degenerating during differentiation leaving functional cytoplasm with limited organelles), specialized sieve plates (perforated end walls with sieve areas containing numerous pores 1-5 μm diameter enabling longitudinal transport between elements), callose deposition (β-1,3-glucan polymer potentially depositing around sieve pores during stress or dormancy partially occluding pores, removed during reactivation), P-protein (phloem protein, sometimes called slime, present in sieve tube lumens, potentially involved in sieve pore sealing following injury), modified cytoplasm (lacking nucleus, ribosomes, Golgi, vacuole, containing only plastids, mitochondria, endoplasmic reticulum, and P-protein), and dependence on companion cells (sieve tubes lacking nucleus requiring companion cell metabolic support for sustained function). Phloem inner trunk bark companion cell characteristics includes= dense cytoplasm (high organelle density including numerous mitochondria, ribosomes, Golgi, endoplasmic reticulum supporting high metabolic activity), large nucleus (prominent nucleus directing intensive protein synthesis and metabolic regulation), symplastic connection to sieve tubes (plasmodesmata connecting companion cell and sieve tube cytoplasm enabling metabolite transfer), active sugar transport (ATP-powered sugar transporters loading sucrose from apoplast into companion cell/sieve tube complex against concentration gradients), enzyme production (synthesizing enzymes and proteins required for sieve tube function), and control functions (regulating phloem loading, unloading, and longitudinal transport through metabolic activity). Phloem inner trunk bark parenchyma functions includes= starch storage (accumulating starch granules particularly in ray and axial parenchyma, 5-12% dry weight starch concentration, contributing 15-25% of trunk total reserves), lateral transport (moving water and solutes between phloem conducting elements and storage sites or ray connections), metabolic support (providing enzymes, metabolites, and energy to adjacent conducting cells), wound healing (parenchyma cells retaining mitotic capacity, proliferating to form callus tissue repairing damage), and defense responses (synthesizing antimicrobial compounds, phytoalexins, and other protective substances responding to pathogen attack). Phloem inner trunk bark fiber characteristics includes= thick lignified secondary walls (providing mechanical strength supporting phloem tissue preventing collapse under turgor pressure and external forces), elongated cell shape (400-800 μm length providing structural reinforcement), dead at maturity (most phloem fibers dying after wall deposition though some varieties showing living fibers), gelatinous fibers (some phloem fibers containing gelatinous G-layer in secondary wall potentially affecting mechanical properties), and increasing proportion with age (older phloem showing higher fiber content as parenchyma crushed and conducting elements obliterated leaving fibers as dominant cell type in transitional and crushed phloem zones). Phloem inner trunk bark ray characteristics includes= heterocellular composition (containing both procumbent cells, radially elongated 40-120 μm radial length, and upright cells, vertically elongated at ray margins), multiseriate organization (typically 3-5 cells wide tangentially providing substantial radial transport pathway), vertical extent (rays 15-35 cells high, 600-1400 μm, providing vertical integration along radial plane), continuous with xylem rays (phloem rays connecting across vascular cambium to xylem rays creating continuous radial transport pathway from inner xylem through cambium and phloem to outer bark), and storage function (ray parenchyma storing starch contributing to total phloem storage capacity, facilitating radial loading and unloading of sugars and other compounds). Phloem inner trunk bark transport mechanisms includes= pressure flow hypothesis (mass flow driven by osmotic pressure gradient, high pressure in source regions from sugar loading forcing flow toward low pressure sink regions where sugar unloading occurs), sieve tube sap composition (10-25% sugar concentration primarily sucrose, also containing amino acids, minerals, hormones, proteins, and other compounds), sap flow velocity (0.2-1.2 m/hour typical rates, varying with source-sink gradients and environmental conditions), bidirectional transport potential (some compounds moving opposite to dominant sugar flow direction, particularly in rays enabling lateral redistribution), and loading/unloading control (companion cell and parenchyma activity determining flow patterns and distribution allocation). Phloem inner trunk bark annual production includes= vascular cambium producing phloem centrifugally (outward toward bark surface), annual phloem increment 0.2-0.8 mm radial thickness (much less than xylem reflecting asymmetric cambial division 5-10 xylem cells per phloem cell), production timing (phloem formation throughout growing season April-September, continuing slightly after xylem production ceases), cumulative depth (annual production over 3-5 years creating 1.5-3.5 mm functional phloem zone before older tissue becomes crushed and non-functional), and continuous renewal (ongoing phloem production maintaining stable functional depth despite outer phloem loss to crushing and incorporation into outer bark). Phloem inner trunk bark functional lifespan includes= initial differentiation and maturation (newly produced phloem cells differentiating over 2-4 weeks becoming functional elements), peak function period (1-3 years, sieve tubes maintaining maximum conductivity and parenchyma active in storage), declining function (years 3-4, sieve tube conductivity declining as callose accumulates and P-protein potentially clogs pores, parenchyma maintaining storage but declining in transport capacity), crushing and obliteration (years 4-6+, old phloem compressed by xylem expansion, sieve tubes and parenchyma collapsing losing function, becoming incorporated into transitional zone approaching outer bark), and total functional lifespan 2-4 years (compared to xylem which remains functional 8-15 years as sapwood before transitioning to heartwood). Phloem inner trunk bark relationship to xylem includes= produced by same vascular cambium (bifacial meristem producing phloem outward and xylem inward), asymmetric production ratio (5-10× more xylem than phloem produced annually), complementary functions (xylem transporting water and minerals upward, phloem transporting photosynthate and nutrients downward and laterally), radial connection through rays (xylem and phloem rays providing lateral pathway connecting vascular tissues), and temporal coordination (phloem and xylem production synchronized during growing season, both ceasing during dormancy). Phloem inner trunk bark disease susceptibility includes= phloem necrosis (viruses or phytoplasmas causing sieve tube dysfunction and phloem parenchyma death), trunk canker diseases (Eutypa, Esca, Botryosphaeria colonizing and killing phloem tissue), vascular wilt pathogens (some organisms colonizing phloem disrupting transport), girdling pathogens (circumferential phloem death blocking basipetal transport causing distal starvation and vine death), and importance of phloem health (maintaining functional phloem essential for vine productivity, vigor, and survival). Phloem inner trunk bark significance includes= sole pathway for photosynthate distribution (phloem providing only mechanism for transporting sugars from source leaves to sink roots, fruit, and storage tissues), critical storage tissue (phloem parenchyma storing 15-25% of trunk carbohydrate reserves), nutrient recycling pathway (phloem redistributing minerals and mobilized nutrients from senescing tissues), hormone transport system (phloem transporting signals coordinating vine development and stress responses), and girdling sensitivity (complete phloem disruption around trunk circumference fatal within weeks emphasizing phloem critical importance).
Phloem_Inner_Trunk_Bark emphasizes tissue identity and complex organization and is distinguished from Inner_Trunk_Bark (anatomical zone), Living_Inner_Trunk_Bark (viability emphasis), and secondary phloem in other organs by specific trunk bark context. Critical for understanding phloem tissue structure, sieve tube-companion cell functional unit, transport mechanisms, phloem lifespan and turnover, storage in phloem parenchyma, and essential role as sole long-distance photosynthate transport pathway requiring protection and management maintaining phloem health for vine survival and productivity.
Bark phloem tissue
Innenrinden-Phloem
Inner bark phloem
Secondary phloem
Phloem Inner Trunk Bark
Phloem necrosis quality evaluation in Michigan Concord following a midwinter freeze event (-29°C) assessed the vascular damage affecting carbohydrate transport capacity for the coming growing season. Cross-sectional examination of cane samples collected at budbreak revealed the extent and distribution of phloem damage through visual assessment of inner bark coloration. Healthy phloem appeared as a cream-colored to light tan continuous ring inside the bark, while necrotic phloem showed brown to dark brown discoloration indicating dead tissue. Phloem necrosis quality was scored as the percentage of the phloem ring showing browning, with results ranging from 0% (no visible damage) through 50% (half the phloem circumference affected) to 100% (complete phloem girdling). Regional analysis showed phloem necrosis quality averaging 35% across the surveyed area, with individual vine scores ranging from 10% to 80% reflecting microclimate variation and vine health status affecting hardiness. Correlation analysis between phloem necrosis quality and subsequent vine performance showed strong relationships, with vines exceeding 50% phloem necrosis showing 40% reduction in shoot vigor (measured as shoot length at 60 days), 35% reduction in fruit set, and 25% reduction in fruit yield compared to vines with less than 20% phloem necrosis. The phloem necrosis quality assessment established that vascular damage substantially affected productivity even in vines with good bud survival, identifying phloem status as an important secondary assessment beyond bud browning for comprehensive damage evaluation.
Phloem necrosis quality is a cold damage symptom quality that inheres in grapevine vascular tissues and represents the extent of death and discoloration in the phloem tissue layer visible in cross-sectioned canes and trunks following freezing injury, with phloem damage compromising the transport of photosynthates from leaves to roots and developing fruit that is essential for vine productivity and the accumulation of carbohydrate reserves required for winter survival and spring growth initiation. This quality addresses damage to the carbohydrate transport system that directly affects vine function and productivity, recognizing that phloem integrity is required for the source-sink relationships supporting fruit development during the growing season and for the reserve accumulation and mobilization that supports vine survival and regrowth, such that phloem damage has consequences extending well beyond the immediate injury. The ontological significance of phloem necrosis quality lies in its indication of vascular compromise affecting whole-vine function, with extensive phloem damage potentially limiting productivity even when buds survive by restricting the carbohydrate supply available to support shoot growth and fruit development, and with trunk-level phloem damage potentially proving fatal by preventing the root system from receiving the photosynthate supply required for function and survival. The biological basis of phloem necrosis encompasses the death of sieve tube elements and companion cells that constitute the functional phloem when freezing causes lethal cellular damage, with the killed tissue undergoing oxidation producing visible browning in cross-section that appears as a tan-to-brown ring in the inner bark region outside the cambium. The characteristic appearance of phloem necrosis in cross-section shows the affected tissue as a darker band contrasting with the lighter color of healthy phloem, with the extent of the browning indicating the proportion of phloem cross-sectional area that has been killed and the corresponding degree of transport capacity compromise. The functional consequences of phloem necrosis may not become fully apparent until the growing season when the reduced transport capacity limits shoot vigor, fruit development, and reserve accumulation in proportion to the extent of damage, with severely affected vines showing weak growth, poor fruit set, and inadequate hardening for the subsequent winter.
Phloem_Necrosis_Quality is_a Cold_Damage_Symptom_Quality, representing freeze injury to the phloem carbohydrate transport tissue. This class addresses vascular damage affecting source-sink function beyond immediate visible injury. Cross-section browning in the inner bark region provides diagnostic indicator. Percentage of phloem ring affected provides quantitative damage metric. Functional consequences (reduced vigor, fruit set, yield) correlate with phloem damage extent. May limit productivity even when buds survive by restricting carbohydrate transport. Distinguished from Xylem_Discoloration_Quality by focus on phloem rather than xylem.
Innenrinden-Nekrose
Phloem-Frostschaden
Phloemnekrose
Phloemschaden
inner bark necrosis
phloem damage
phloem freeze injury
vascular transport damage
Phloem Necrosis Quality
In a Sonoma Pinot Noir vineyard, phloem unloading activation was detected 2 days before initial sugar increase in mid-August Year N+1 (BBCH 83, 58 days post-flowering) when symplastic connectivity declined from 52% to 18% measured by dye coupling, VvcwINV expression increased 48-fold, cell wall invertase activity rose from 3.2 to 22.5 units/g FW, VvHT1 induction was 35-fold, ABA levels reached 880 ng/g, and apoplastic sucrose concentration was 115 mM under warm conditions (daily high 27-30°C) initiating the critical transport pathway transition.
Phloem_Unloading_Activation represents the developmental transition when berry vascular tissues shift from predominantly symplastic phloem unloading (sucrose moving cell-to-cell through plasmodesmata) to apoplastic unloading (sucrose released into cell wall space requiring membrane transport for cellular uptake) creating the critical pathway change that enables massive hexose accumulation during ripening. This activation occurs during veraison onset (BBCH 83), typically 1-3 days preceding initial sugar increase, and is characterized by plasmodesmata closure or reduced conductance in the phloem-mesocarp boundary reducing symplastic connectivity from pre-veraison 40-60% to post-activation 90-95%). Coordination with other veraison events demonstrates: phloem unloading activation typically precedes initial sugar increase by 1-3 days establishing transport pathway before major accumulation begins, occurs concurrently with or slightly before berry softening onset suggesting common developmental triggers, and in red cultivars often coincides with initial anthocyanin synthesis activation reflecting integrated ABA-mediated ripening program. Metabolic implications inclu apoplastic pathway creates opportunity for regulatory control at multiple steps (invertase activity, hexose transporter expression, tonoplast transport) enabling fine-tuning of sugar accumulation, spatial separation of sucrose cleavage (apoplast) from metabolism (cytoplasm) prevents futile cycling where cleaved hexoses would be immediately re-synthesized to sucrose, and energy requirements include ATP for hexose transport (if active transport components involved) though most transport appears facilitated diffusion down concentration gradients. Evolutionary perspective suggests: apoplastic unloading in ripening berries may represent adaptation for maximizing sugar accumulation enabling high final concentrations (>1000 mM hexoses) difficult to achieve via symplastic pathway due to osmotic and diffusion limitations, shift to apoplastic pathway coincides with seed maturation completion potentially reflecting developmental switch from seed provisioning (requiring controlled symplastic delivery) to fruit attractiveness maximization (benefiting from unrestricted apoplastic accumulation), and convergent evolution with other fleshy fruits (tomato, pepper) showing similar unloading transitions suggesting common adaptive value. Viticultural implications of activation inclu represents critical developmental checkpoint where adequate source capacity becomes essential for supporting subsequent sugar accumulation, crop load management effects expressed most strongly during activation period with overcropping potentially delaying or impairing activation through source limitation, and environmental stresses (water deficit, heat, disease) during activation period can have disproportionate effects on final sugar levels by impairing pathway establishment even if conditions improve later. Research methods for studying activation involve: dye coupling experiments using fluorescent tracers (Lucifer Yellow, carboxyfluorescein) to assess symplastic connectivity showing decreased coupling post-activation, electron microscopy revealing callose deposition at plasmodesmata, gene expression analysis tracking invertase and transporter induction, enzyme activity assays measuring cell wall invertase showing 3-7 fold increases, and sugar composition analysis of apoplast versus cytoplasm/vacuole compartments revealing pathway changes.
Phloem_Unloading_Activation is distinguished from Sugar_Accumulation_Onset by representing pathway mechanism rather than net accumulation, from Initial_Sugar_Increase by preceding measurable Brix rise by 1-3 days, and from Hexose_Transport_Initiation by focusing on phloem unloading rather than membrane transport. Symplastic connectivity drops from 40-60% to <10-20%. Cell wall invertase increases 3-7 fold. Timing: 1-3 days before measurable sugar increase. ABA spike critical trigger. Phloem_Unloading_Activation represents fundamental pathway change enabling massive hexose accumulation during grape ripening.
Entladungs-Mechanismus-Wechsel
Phloem-Mesokarp-Transport-Übergang
apoplastic unloading initiation
apoplastische Entladungs-Initiierung
phloem-mesocarp transport transition
symplastic pathway closure
symplastischer Pfad-Verschluss
unloading mechanism shift
Zhang XY, Wang XL, Wang XF, Xia GH, Pan QH, Fan RC, Wu FQ, Yu XC, Zhang DP (2006) A shift of phloem unloading from symplasmic to apoplasmic pathway is involved in developmental onset of ripening in grape berry Plant Physiol 142:220-232 unloading pathway transition Hayes MA, Davies C, Dry IB (2007) Isolation, functional characterization, and expression analysis of grapevine hexose transporters J Exp Bot 58:1985-1997 hexose transporter roles Delrot S, Picaud S, Gaudillère JP (2001) Sugar and organic acid transport in grapevine berries Rec Adv Phloem Transport Metab:37-47 transport mechanisms.
Phloem Unloading Activation
Phosphate acquisition efficiency assessment comparing rootstocks reveals genotypic variation in transporter-mediated uptake capacity, with 1103P showing Vmax of 4.2 μmol P/g root dry weight/hour and Km of 5.8 μM (high capacity, moderate affinity), while 110R shows lower Vmax of 2.8 μmol/g/hour but improved Km of 3.2 μM (moderate capacity, high affinity), with the different kinetic profiles suggesting 1103P advantaged in high-P soils while 110R may perform better under P limitation, informing rootstock selection based on site-specific phosphorus availability.
Phosphate acquisition efficiency quality is a phosphorus uptake efficiency quality that represents the rate and effectiveness of phosphate ion absorption from soil solution by grapevine roots, characterizing the transport-mediated uptake of the bioavailable phosphorus form that involves high-affinity transporter proteins enabling acquisition from the micromolar concentrations typical of soil solution. Within the BFO framework, this quality specifically addresses the membrane transport dimension of phosphorus uptake, distinguishing the absorption step from phosphorus mobilization and utilization while recognizing that phosphate transporter capacity determines the rate of phosphorus entry into root cells once phosphate is made available at the root surface through diffusion, mycorrhizal delivery, or root exudate-mediated solubilization. The ontological significance lies in phosphate acquisition efficiency determining the rate of phosphorus capture once phosphate reaches the root surface, where high-affinity PHT transporters enable absorption at the low micromolar concentrations typical of soil solution (1-10 μM), with transporter expression and activity determining whether roots can effectively capture the limited phosphate diffusing toward them from surrounding soil. The molecular basis for phosphate acquisition involves the PHT1 family of plasma membrane phosphate transporters that catalyze H⁺-coupled phosphate uptake, with high-affinity transporters (Km approximately 3-10 μM) operating at soil solution concentrations and low-affinity transporters (Km approximately 50-300 μM) contributing when phosphate is more abundant, and with transporter expression upregulated under phosphorus deficiency to maximize acquisition capacity. The kinetic characteristics of phosphate acquisition show saturation behavior described by Michaelis-Menten kinetics, with maximum uptake rate (Vmax) reflecting transporter abundance and catalytic rate, and Km reflecting transporter affinity, such that genotypes with higher Vmax and lower Km achieve more efficient acquisition at the low concentrations typical of vineyard soils. The root zone depletion dynamics critically affect phosphate acquisition, as absorption rapidly depletes phosphate from the immediate root vicinity (depletion zone extending only 1-2 mm due to low diffusion), requiring continuous root growth or hyphal extension to access fresh soil phosphate. Understanding phosphate acquisition efficiency quality enables assessment of the transporter-mediated uptake capacity, interpretation of genotypic differences in phosphorus nutrition, and appreciation of the molecular basis for phosphate absorption that complements root architectural and mycorrhizal contributions to overall phosphorus uptake.
Phosphate_Acquisition_Efficiency_Quality addresses the membrane transport step of phosphorus uptake mediated by PHT transporter family. This quality determines uptake rate once phosphate reaches the root surface. The class supports molecular understanding of P uptake and genotype comparison through acquisition efficiency characterization.
PHT activity
PHT-Aktivität
Phosphatabsorptionsrate
Phosphatakquisitionseffizienz
Pi acquisition
Pi-Aufnahme
phosphate absorption rate
phosphate transport efficiency
Muchhal US et al. (1996) Phosphate transporters from the higher plant Arabidopsis thaliana. Proceedings of the National Academy of Sciences 93(19): 10519-10523 Raghothama KG (1999) Phosphate acquisition. Annual Review of Plant Physiology and Plant Molecular Biology 50: 665-693
Phosphate Acquisition Efficiency Quality
The phosphorus content of a Stellenbosch Cabernet Sauvignon vineyard (South Africa) on decomposed granite soil documents Bray-1 extractable phosphorus of 18 mg/kg (low-moderate) at 0-30 cm depth — this relatively low phosphorus availability reflects the strong phosphorus fixation capacity of the iron- and aluminum-oxide-rich saprolitic subsoil that converts applied phosphorus to insoluble forms, requiring periodic phosphorus applications (15-25 kg P/ha as superphosphate or rock phosphate incorporated before planting, with maintenance applications of 10-15 kg P/ha every 3-5 years based on soil test monitoring) to maintain available phosphorus above the 15 mg/kg Bray-1 threshold below which root development and mycorrhizal colonization may be impaired. The moderate phosphorus status contrasts with the elevated phosphorus levels (Olsen-P 45-80 mg/kg) commonly found in long-established European vineyard soils where decades of phosphorus fertilization have accumulated residual phosphorus far above viticultural sufficiency levels — a legacy phosphorus pool that may sustain vine phosphorus nutrition for decades without further application.
A nutrient analysis data component that records the concentration of plant-available phosphorus in the vineyard soil, determined by standardized extraction methods — Olsen bicarbonate extraction (0.5M NaHCO₃ at pH 8.5, preferred for calcareous and neutral soils) or Bray-1 extraction (0.03M NH₄F + 0.025M HCl, preferred for acid soils) — expressed as mg P/kg soil, with phosphorus influencing root development, energy metabolism, flowering and fruit set, and the early-season establishment of vine growth, though phosphorus deficiency is relatively uncommon in established vineyard soils and phosphorus excess poses environmental contamination risk through runoff to surface waters.
Phosphorus_Content documents a nutrient whose viticultural management has shifted from routine application (the historical practice of annual phosphorus fertilization as part of standard NPK programs) to targeted, soil-test-based application (the modern approach recognizing that most established vineyard soils contain adequate or excessive phosphorus). The chemistry of soil phosphorus is dominated by fixation reactions — in acid soils, phosphorus reacts with iron and aluminum oxides to form insoluble iron and aluminum phosphates, while in calcareous soils, phosphorus precipitates as calcium phosphate (apatite) — that immobilize applied phosphorus and maintain solution phosphorus concentrations at very low levels (typically 0.01-0.1 mg P/L in the soil solution) regardless of the total phosphorus content: the vine accesses this low-concentration phosphorus pool primarily through mycorrhizal fungi (arbuscular mycorrhizal fungi of the Glomeromycota, which colonize 60-90% of grapevine roots and extend the effective root absorptive surface area by 10-100 fold through their extraradical hyphal network), making mycorrhizal function critical for phosphorus acquisition and explaining why vineyard management practices that maintain mycorrhizal colonization (reduced tillage, cover crops, moderate phosphorus levels) are essential for efficient phosphorus nutrition. The environmental significance of phosphorus in vineyard soils has gained recognition as the primary nutrient of concern for surface water eutrophication — vineyard soils with excessive Olsen-P (>60 mg/kg) are at risk of phosphorus loss through erosion and runoff, particularly on sloping sites with bare soil between vine rows.
Boden-Phosphor
P-Gehalt-Messung
Phosphorgehalt
available P measurement
phosphorus status assessment
soil phosphorus record
White RE (2003) Soils for Fine Wines, Oxford University Press Robinson JB (2005) Critical plant tissue values, in Christensen LP and Smart DR (eds) Proceedings of the Soil Environment and Vine Mineral Nutrition Symposium, ASEV Schreiner RP (2005) Mycorrhizas and mineral acquisition in grapevines, in Christensen LP and Smart DR (eds) Proceedings of the Soil Environment and Vine Mineral Nutrition Symposium, ASEV
Phosphorus Content
Phosphorus content quality management in Stellenbosch Cabernet Sauvignon addresses the phosphorus-fixing granite-derived soils characteristic of the region, with careful phosphorus fertilization required to maintain must concentrations of 150-250 mg/L that support healthy fermentation kinetics phosphorus-deficient musts (below 100 mg/L) show sluggish fermentation and increased hydrogen sulfide risk as stressed yeast struggle with energy metabolism, while adequate phosphorus nutrition enables the clean, complete fermentations producing the age-worthy Cabernet for which Stellenbosch is renowned.
Phosphorus_Content_Quality represents the chemical quality characterizing the concentration of phosphorus (as phosphate, PO₄³⁻) in grape berry tissue, capturing the essential macronutrient central to energy transfer, nucleic acid synthesis, and membrane phospholipid composition. Phosphorus in grape juice typically ranges from 100-400 mg/L, with the wide variation reflecting cultivar, rootstock, and soil phosphorus availability. As a component of ATP, ADP, and other nucleotide phosphates, phosphorus is essential for energy metabolism in all cells, while its incorporation into DNA, RNA, and phospholipid membranes makes it fundamental to growth and development. In berries, phosphorus accumulates through both xylem and phloem transport, with continued accumulation during ripening supporting the energy-intensive processes of sugar accumulation and secondary metabolite synthesis. Soil phosphorus availability often limits grapevine productivity, particularly in low-pH soils where phosphorus binds strongly to iron and aluminum, or in high-pH calcareous soils where calcium phosphate precipitation reduces availability. Phosphorus deficiency produces distinctive symptoms including reduced vigor, dark green or purplish coloration of leaves, and poor fruit set, though severe deficiency is uncommon in established vineyards. In winemaking, phosphorus affects yeast nutrition during fermentation, with adequate phosphate supporting healthy yeast metabolism while very high phosphate may contribute to protein and tartrate instability through complex interactions. The phosphorus content of wine is typically lower than must due to precipitation and yeast uptake during fermentation. Phosphorus analysis employs colorimetric methods (molybdenum blue), ICP-OES, or ion chromatography depending on required sensitivity and matrix complexity.
Phosphorus_Content_Quality addresses an essential macronutrient whose availability is often limited by soil chemistry rather than total soil content. The energy-transfer role of phosphorus (ATP synthesis and utilization) makes it fundamental to all metabolic processes. In wine production, phosphorus affects yeast fermentation performance more directly than wine quality per se. The complex soil chemistry of phosphorus (fixation, precipitation) creates disconnect between total soil phosphorus and plant-available phosphorus, requiring specific analytical methods for availability assessment.
Beerenphosphorqualität
P content
P-Gehalt
Phosphatkonzentration
berry phosphorus quality
phosphate concentration
phosphorus level
Phosphorus Content Quality
Phosphorus deficiency symptom quality diagnosis in a newly established Oregon Pinot Noir vineyard identified P limitation as the cause of poor establishment and abnormal coloration in first-year vines planted on highly weathered forest soil. Visual phosphorus deficiency symptom quality assessment noted the distinctive red-purple coloration of leaves beginning at margins and progressing intervenally, distinct from the uniform chlorosis of nitrogen deficiency and from the interveinal chlorosis of magnesium deficiency. Young vine growth showed phosphorus deficiency symptom quality with stunted shoot development (mean 25 cm at midsummer versus expected 60 cm), small leaves with reddish-purple hue most pronounced on lower canopy positions, and poor root development revealed when struggling vines were excavated showing root mass 35% of normally establishing vines. Soil analysis identified the cause of phosphorus deficiency symptom quality as extremely low available phosphorus (Bray-1 P of 3 ppm versus sufficiency threshold of 25 ppm) combined with soil pH 5.2 and elevated exchangeable aluminum at 1.8 cmol/kg, conditions favoring phosphorus precipitation as aluminum phosphates rendering it unavailable. Tissue analysis confirmed the diagnosis with petiole phosphorus at 0.08% versus sufficiency range of 0.15-0.30%. Corrective treatment with banded triple superphosphate application at 80 kg P₂O₅/ha combined with lime to raise pH above 5.8 and reduce aluminum activity resolved phosphorus deficiency symptom quality by the following season, with second-year growth showing normal vigor and coloration.
Phosphorus deficiency symptom quality is a macronutrient deficiency symptom quality that inheres in a grapevine organism and represents the characteristic visual symptoms and growth limitations resulting from inadequate phosphorus availability, including the development of reddish or purplish coloration on leaves from anthocyanin accumulation, reduced root system development, stunted growth particularly in young vines, delayed fruit ripening, and reduced berry set, with phosphorus deficiency being relatively uncommon in established vineyards due to adequate soil phosphorus in most viticultural regions but occasionally significant in newly planted vineyards or on highly weathered, acidic, or calcareous soils where phosphorus availability is limited. This quality captures the grapevine response to phosphorus limitation, recognizing that phosphorus is essential for energy metabolism (ATP), nucleic acid synthesis, membrane structure (phospholipids), and numerous regulatory functions, making deficiency symptoms reflect disruption across multiple physiological systems. The ontological significance of phosphorus deficiency symptom quality lies in the diagnostic challenge it presents due to symptoms that can resemble other stresses including cold temperatures, viral infection, and other nutrient deficiencies, requiring careful differential diagnosis to correctly identify phosphorus as the limiting factor and avoid inappropriate management responses. The biological basis of phosphorus deficiency symptoms encompasses the role of phosphorus in energy metabolism where ATP limitation affects all energy-requiring processes, the requirement for phosphorus in DNA and RNA synthesis affecting cell division and protein synthesis, the phospholipid content of membranes affecting their structure and function, and the triggering of anthocyanin accumulation as a stress response to phosphorus limitation that produces the characteristic red-purple coloration. The soil chemistry of phosphorus creates availability challenges, with high soil pH causing precipitation as calcium phosphates, low soil pH favoring precipitation with iron and aluminum, and the relative immobility of phosphorus in soil limiting root access to this nutrient even when total soil phosphorus is adequate.
Phosphorus_Deficiency_Symptom_Quality is_a Macronutrient_Deficiency_Symptom_Quality, serving as parent class for Anthocyanin_Accumulation_Quality_(P) and Root_Growth_Limitation_Quality_(P) which address specific phosphorus deficiency manifestations. This class addresses the overall phosphorus deficiency syndrome while child classes capture particular symptom types. Less common than N or K deficiency in established vineyards but significant in new plantings on problem soils. Soil chemistry (pH, Al, Ca) affects P availability independent of total soil P. The red-purple coloration can resemble cold stress or viral symptoms requiring differential diagnosis.
Niedrig-Phosphor-Symptome
P deficiency symptoms
P stress symptoms
P-Mangel-Zeichen
Phosphormangel-Symptome
Phosphorstress-Indikatoren
low phosphorus indicators
phosphorus limitation signs
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Grant CA et al. (2001) The importance of early season phosphorus nutrition. Canadian Journal of Plant Science 81:211-224 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication
Phosphorus Deficiency Symptom Quality
Phosphorus uptake efficiency varies 3-4 fold among rootstocks, with 1103 Paulsen and 140 Ruggeri showing high efficiency (0.8-1.2 mg P/m root length/day) compared to 3309C and 101-14 Mgt (0.3-0.5 mg P/m/day). Arbuscular mycorrhizal colonization enhances P uptake by 30-80% through hyphal exploration of soil volumes inaccessible to roots. Total vine phosphorus content at harvest ranges from 12-25 kg P/ha, with approximately 40-50% removed in fruit. Phosphorus deficiency causes reduced shoot growth, anthocyanin accumulation in leaves, and poor fruit set, with symptoms appearing when petiole P falls below 0.12% dry weight at bloom.
A macronutrient uptake disposition that is the inherited capacity of a Vitis vinifera plant to absorb phosphate ions (H₂PO₄⁻, HPO₄²⁻) from the soil solution through phosphate transporter systems and mycorrhizal associations, with genotype-specific efficiency determining phosphorus acquisition under often limiting soil availability conditions.
Phosphorus availability in vineyard soils is often limited by fixation to iron and aluminum oxides (acidic soils) or calcium compounds (calcareous soils), making uptake efficiency a critical adaptation. Rootstock selection for high P uptake efficiency can reduce fertilizer requirements and improve vine performance on low-P soils. Root architectural traits including fine root density and root hair development correlate with P uptake efficiency. VvPHT1 gene family members encode phosphate transporters with expression responsive to P availability.
P uptake capacity
P-Akquisitionseffizienz
Phosphataneignungsvermögen
Phosphoraufnahme-Effizienz
phosphate foraging trait
phosphorus acquisition efficiency
Schreiner RP (2005) Mycorrhizae and mineral acquisition in grapevines. In: Christensen LP, Smart DR (eds) Proceedings of the Soil Environment and Vine Mineral Nutrition Symposium, ASEV pp.: 49-60 Grant RS, Matthews MA (1996) The influence of phosphorus availability and rootstock on root system characteristics, phosphorus uptake, phosphorus partitioning, and growth efficiency. American Journal of Enology and Viticulture 47:403-409
Phosphorus Uptake Efficiency Disposition
Phosphorus uptake efficiency comparison between mycorrhizal and non-mycorrhizal grapevines reveals the symbiotic contribution to phosphorus acquisition, with mycorrhizal vines showing P uptake of 0.85 mg P/vine/day (extensive hyphal network accessing soil P beyond root depletion zones) while non-mycorrhizal vines achieve only 0.28 mg P/day (limited to direct root contact with P), with the 3-fold difference demonstrating the critical role of mycorrhizal associations in grapevine phosphorus nutrition that must be considered in vineyard establishment and soil management practices.
Phosphorus uptake efficiency quality is a nutrient uptake efficiency quality that represents the capacity of grapevine root systems to acquire phosphorus from soil despite its extreme immobility, characterizing the effectiveness of phosphate acquisition through root proliferation, exudation, and mycorrhizal symbiosis that determines vine phosphorus status affecting energy metabolism, nucleic acid synthesis, and membrane function essential for growth and fruit development. Within the BFO framework, this quality specifically addresses the phosphorus component of nutrient uptake, distinguishing phosphorus acquisition from other nutrients while recognizing that phosphorus presents unique acquisition challenges due to its immobility in soil requiring specialized root adaptations and symbiotic associations for effective uptake. The ontological significance lies in phosphorus uptake efficiency determining access to this essential but often limiting nutrient, where the extreme immobility of phosphate in soil (diffusion coefficient approximately 10⁻¹² to 10⁻¹⁵ m²/s, three to six orders of magnitude lower than nitrate) means that roots must grow toward phosphorus rather than relying on mass flow delivery, making root architecture, exudation, and mycorrhizal associations critical for phosphorus acquisition. The mechanistic basis for phosphorus uptake involves phosphate ion (H₂PO₄⁻ or HPO₄²⁻) absorption from the soil solution at concentrations typically below 10 μM, with uptake mediated by high-affinity PHT (Phosphate Transporter) family proteins that enable acquisition against steep concentration gradients, and with root adaptations including increased root hair density, organic acid exudation to solubilize bound phosphorus, and mycorrhizal symbiosis extending the effective absorbing surface. The soil factors affecting phosphorus availability include pH (availability highest near pH 6.5, declining at both extremes), calcium and iron content (forming insoluble phosphate precipitates), organic matter (providing mineralizable phosphorus), and soil texture (affecting phosphate adsorption to particle surfaces), with these factors often limiting phosphorus availability even in fertilized soils. The rootstock effects on phosphorus uptake are significant, with some rootstocks showing enhanced phosphorus acquisition through more extensive root systems, greater mycorrhizal responsiveness, or more effective root exudation, affecting the phosphorus nutrition of grafted scions. Understanding phosphorus uptake efficiency quality enables assessment of vine phosphorus acquisition capacity, optimization of phosphorus fertilization strategies, and rootstock selection for improved phosphorus nutrition in challenging soil conditions.
Phosphorus_Uptake_Efficiency_Quality serves as the parent class for P acquisition components (Phosphate Acquisition, P Use Efficiency, Mycorrhizal Colonization). This quality addresses the unique challenge of acquiring highly immobile soil phosphorus. The class supports phosphorus management and rootstock selection through P uptake efficiency characterization.
P uptake efficiency
P-Aufnahmekapazität
PUptE
Phosphatakquisition
Phosphoraufnahmeeffizienz
phosphate acquisition
root P foraging
Schachtman DP et al. (1998) Phosphorus uptake by plants: from soil to cell. Plant Physiology 116(2): 447-453 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 197-245
Phosphorus Uptake Efficiency Quality
Phosphorus use efficiency in grapevines ranges from 400-900 kg fruit per kg P in whole-vine content, reflecting the relatively low P requirement compared to nitrogen and potassium. High-PUE rootstocks including 1103 Paulsen and 110 Richter maintain adequate vine P status (petiole P above 0.15% at bloom) with 40-60% lower soil P availability than low-PUE rootstocks such as 3309C. Mycorrhizal colonization enhances PUE by 25-50% through improved P acquisition efficiency, with colonized vines showing higher P uptake per unit root length. Internal P recycling during senescence recovers 70-85% of leaf P for redistribution to perennial tissues, contributing to following-season P supply.
A nutrient use efficiency disposition that is the inherited capacity of a Vitis vinifera plant to produce biomass, yield, or quality outcomes per unit of phosphorus absorbed or available, determined by the combined efficiency of phosphorus uptake from soil, internal phosphate metabolism, and allocation to productive functions versus storage pools.
Phosphorus use efficiency is particularly important in viticulture because P availability is often limited by soil fixation, and P fertilizer resources are finite globally. The trait encompasses both acquisition efficiency (including mycorrhizal contributions) and utilization efficiency (metabolic and allocation components). High-PUE genotypes show enhanced acid phosphatase activity, improved internal P recycling, and optimized P allocation between storage (vacuolar polyphosphate) and metabolically active pools. Root architectural traits including fine root proliferation and root hair density correlate with P acquisition efficiency.
P productivity
P-Verwertungseffizienz
Phosphornutzungseffizienz
Phosphorproduktivität
phosphorus efficiency
phosphorus utilization trait
Phosphorus Use Efficiency Disposition
Phosphorus use efficiency assessment comparing vineyard management systems reveals the impact of vine balance on phosphorus productivity, with balanced vines (moderate vigor, appropriate crop load) achieving PUE of 145 kg fruit/kg plant P (efficient conversion of phosphorus into reproductive output), while excessively vigorous vines show PUE of only 85 kg fruit/kg P (phosphorus sequestered in vegetative growth rather than fruit), demonstrating how canopy and crop management affects phosphorus use efficiency with implications for both fertilization requirements and environmental sustainability.
Phosphorus use efficiency quality is a phosphorus uptake efficiency quality that represents the effectiveness with which grapevines convert acquired phosphorus into biomass, yield, and quality parameters, characterizing the integrated efficiency of phosphorus assimilation, translocation, and utilization that determines the productive return on phosphorus investment and the sustainability of phosphorus management in viticulture. Within the BFO framework, this quality addresses the utilization dimension of phosphorus nutrition, distinguishing efficiency of use from efficiency of acquisition while recognizing that overall phosphorus efficiency depends on both acquiring phosphorus and using it effectively to support vine function and grape quality. The ontological significance lies in phosphorus use efficiency determining how much productive output is achieved per unit phosphorus in the vine, where high PUE enables adequate productivity with lower tissue phosphorus concentrations, reducing both the phosphorus fertilizer requirement and the risk of phosphorus runoff that contributes to eutrophication of water bodies, making PUE an important sustainability metric for viticulture. The physiological basis for phosphorus use efficiency includes the metabolic functions of phosphorus in ATP and nucleotide synthesis, phospholipid membrane components, sugar-phosphate intermediates of metabolism, and signal transduction molecules, with efficient use requiring optimization of these pools relative to growth demand and effective recycling of phosphorus from senescing tissues to growing points. The components of phosphorus use efficiency include assimilation efficiency (incorporation of acquired phosphorus into organic forms), translocation efficiency (movement of phosphorus to productive sinks), and utilization efficiency (biomass or yield produced per unit plant phosphorus), with each component potentially limiting overall PUE depending on conditions and genotype. The vine life cycle dynamics of phosphorus show strong remobilization from vegetative tissues to reproductive organs during ripening, with efficient internal cycling reducing the external phosphorus requirement and contributing to high PUE, while excessive vegetative growth may sequester phosphorus in leaves rather than fruit, reducing reproductive PUE. Understanding phosphorus use efficiency quality enables assessment of phosphorus nutrition effectiveness beyond simple acquisition, optimization of phosphorus management for both productivity and environmental sustainability, and genotype evaluation for phosphorus-efficient production in challenging conditions.
Phosphorus_Use_Efficiency_Quality addresses the overall effectiveness of converting acquired phosphorus into productive output. This quality integrates assimilation, translocation, and utilization components affecting phosphorus productivity. The class supports sustainable phosphorus management through PUE optimization balancing productivity and environmental impact.
P utilization efficiency
P-Produktivität
PUE
Phosphorkonversionseffizienz
phosphorus conversion efficiency
phosphorus productivity
Veneklaas EJ et al. (2012) Opportunities for improving phosphorus-use efficiency in crop plants. New Phytologist 195(2): 306-320 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 197-245
Phosphorus Use Efficiency Quality
Photochemical quenching assessment across canopy light gradient in Cabernet Sauvignon reveals the PSII saturation pattern, with shaded interior leaves showing qP of 0.85 (most centers open, electron transport keeping pace with limited light absorption), mid-canopy leaves showing qP of 0.65 (moderate saturation, balanced condition), and fully exposed exterior leaves showing qP of 0.35 (substantial saturation, absorbed energy exceeding electron transport capacity), with the low qP in sun leaves indicating reliance on NPQ for photoprotection and potential benefit from partial shading to reduce photoinhibition risk.
Photochemical quenching quality is a chlorophyll fluorescence quality that represents the proportion of PSII reaction centers in the open state capable of performing photochemistry, characterizing the balance between light absorption and electron transport capacity that indicates whether photosynthetic electron flow is keeping pace with light energy input or becoming saturated. Within the BFO framework, this quality specifically addresses the photochemical component of fluorescence quenching, distinguishing productive energy use (qP) from protective dissipation (NPQ) while recognizing that photochemical quenching reflects the oxidation state of the primary electron acceptor QA and thus the availability of PSII for continued photochemistry. The ontological significance lies in photochemical quenching indicating the degree of PSII saturation under current light conditions, where high qP (approaching 1.0) indicates most reaction centers remain open and electron transport is not limiting, while low qP (approaching 0) indicates most centers are closed and absorbed energy exceeds electron transport capacity, creating potential for photodamage if photoprotective mechanisms are insufficient. The photophysical basis for photochemical quenching involves the reduction state of QA, the primary quinone electron acceptor of PSII, where qP = (Fm' - Fs)/(Fm' - Fo') with Fm' being maximum fluorescence in light, Fs steady-state fluorescence, and Fo' minimal fluorescence in light, with the parameter reflecting the fraction of PSII centers with QA oxidized and ready for photochemistry. The calculation variants for photochemical quenching include the original qP (assuming independent PSII centers) and qL (assuming connected antennae with lake model), with qL = qP × Fo'/Fs providing a more accurate estimate of the fraction of open centers when antenna connectivity is significant. The light intensity relationship with photochemical quenching shows qP declining as light increases beyond the capacity of electron transport to utilize absorbed energy, with the rate of qP decline indicating the adequacy of electron transport capacity relative to light harvesting. Understanding photochemical quenching quality enables assessment of PSII saturation status, diagnosis of electron transport limitation, and optimization of canopy light environment to maximize productive light use while minimizing photoinhibition risk.
Photochemical_Quenching_Quality addresses the PSII open/closed balance indicating photochemical capacity utilization. This quality decreases as light intensity increases relative to electron transport capacity. The class supports assessment of PSII saturation and diagnosis of conditions where absorbed light exceeds productive utilization capacity.
PSII openness
PSII-Offenheit
Reaktionszentrum-Oxidationszustand
photochemical efficiency factor
photochemisches Quenching
qP-Wert
reaction center oxidation state
Baker NR (2008) Chlorophyll fluorescence: a probe of photosynthesis in vivo. Annual Review of Plant Biology 59: 89-113 Kramer DM et al. (2004) New fluorescence parameters for the determination of QA redox state and excitation energy fluxes. Photosynthesis Research 79(2): 209-218
Photochemical Quenching Quality
Photoinhibition susceptibility comparison between irrigated and drought-stressed Grenache reveals the stress interaction increasing light vulnerability, with irrigated vines showing Fv/Fm decline from 0.82 to 0.75 under 4-hour high light exposure (1800 μmol/m²/s) with full overnight recovery, while drought-stressed vines display severe decline to 0.58 under identical light treatment with incomplete recovery to 0.68 after two nights, demonstrating how water stress impairs photoprotection and repair, increasing susceptibility to photoinhibition that would be tolerated under adequate water supply, with implications for irrigation management in high-light environments.
Photoinhibition susceptibility quality is a light response quality that represents the vulnerability of the photosynthetic apparatus to light-induced damage and functional decline when absorbed light energy exceeds the capacity for productive utilization and photoprotective dissipation, characterizing the resilience of PSII and associated components to excess light stress that determines performance maintenance under high-light conditions. Within the BFO framework, this quality specifically addresses the damage vulnerability dimension of light response, distinguishing susceptibility to photoinhibition from the light levels causing it while recognizing that photoinhibition susceptibility determines whether high light exposure causes transient, reversible reduction in photosynthetic efficiency or chronic, damaging photoinhibition requiring extended recovery periods. The ontological significance lies in photoinhibition susceptibility determining the operational light tolerance of leaves, where highly susceptible leaves suffer productivity losses and potential permanent damage at light intensities that more resilient leaves tolerate without consequence, with susceptibility varying among cultivars, acclimation states, and stress conditions, and critically affecting vine productivity in high-light environments. The mechanistic basis for photoinhibition involves damage to PSII, particularly the D1 protein that forms the reaction center core, when the rate of light-induced damage exceeds the rate of repair through D1 protein turnover, with net photoinhibition representing the balance between damage and repair processes that determines whether photosynthetic capacity is maintained or declines under excess light. The factors affecting photoinhibition susceptibility include photoprotective capacity (NPQ, xanthophyll cycle function), repair capacity (D1 protein synthesis rate, dependent on temperature and protein synthesis machinery), additional stresses (drought, heat, cold, nutrient deficiency all increasing susceptibility by impairing photoprotection or repair), and acclimation history (shade leaves being more susceptible than sun leaves due to limited photoprotective investment). The assessment of photoinhibition susceptibility employs monitoring of Fv/Fm decline under high light exposure and subsequent recovery kinetics, with rapid decline and slow recovery indicating high susceptibility while maintained Fv/Fm or rapid recovery indicating resilience. Understanding photoinhibition susceptibility quality enables prediction of high-light stress impacts, identification of cultivar or condition-specific vulnerabilities, and canopy management targeting appropriate light exposure for leaves of varying susceptibility.
Photoinhibition_Susceptibility_Quality addresses the vulnerability to light-induced photosynthetic damage. This quality depends on both photoprotective capacity and repair capacity, varying with acclimation and stress status. The class supports stress management through understanding of factors affecting light stress tolerance and recovery.
Lichtstressempfindlichkeit
PSII vulnerability
PSII-Vulnerabilität
Photoinhibitionsanfälligkeit
Photoschadensanfälligkeit
light stress sensitivity
photodamage susceptibility
photoinhibition risk
Aro EM et al. (1993) Photoinhibition of photosystem II. Inactivation, protein damage and turnover. Biochimica et Biophysica Acta 1143(2): 113-134 Long SP et al. (1994) Photoinhibition of photosynthesis in nature. Annual Review of Plant Physiology and Plant Molecular Biology 45(1): 633-662
Photoinhibition Susceptibility Quality
In Pinot Noir vines at latitude 47°N in the Willamette Valley, Oregon, photoperiod response initiates dormancy preparation when day length decreases below approximately 13.5 hours in mid-September, with phytochrome-mediated signaling in mature leaves triggering measurable increases in bud abscisic acid content within 2-3 weeks of critical photoperiod exposure and subsequent cessation of shoot elongation by early October.
Photoperiod response in Vitis vinifera encompasses the physiological processes through which grapevines detect changes in day length and transduce this environmental information into developmental signals that regulate dormancy induction and other seasonal growth transitions. This sophisticated sensory and signaling system enables vines to anticipate the approach of winter based on the highly reliable seasonal changes in photoperiod, initiating dormancy preparation well before potentially damaging cold temperatures occur. The detection of photoperiod occurs primarily in mature leaves through the photoreceptor phytochrome system, which exists in two interconvertible forms whose ratio changes with the relative durations of light and darkness, with the Pfr form produced during light exposure converting to the Pr form during extended dark periods characteristic of long nights. When night length exceeds a cultivar-specific critical threshold, typically corresponding to day lengths below 13-14 hours for most Vitis vinifera cultivars, the altered phytochrome signaling initiates cascades of gene expression changes that ultimately affect distant meristematic tissues including buds and cambium. The signal transduction pathway involves integration with circadian clock components that measure night length with remarkable precision, interactions with hormone biosynthesis and signaling pathways particularly involving abscisic acid and gibberellin, and long-distance transport of signaling molecules through the phloem from photoperception sites in leaves to responding tissues in buds and stems. The photoperiod response exhibits significant genotypic variation, with cultivars adapted to higher latitudes generally responding to longer critical day lengths than those from lower latitudes, reflecting evolutionary adaptation to the different relationships between photoperiod and the onset of cold conditions in different geographic regions. Environmental factors including temperature and plant water status can modulate the photoperiod response, with warm temperatures potentially delaying the response and water stress potentially enhancing sensitivity to short-day signals. The practical implications of photoperiod response for viticulture include understanding cultivar adaptation to different latitudes, predicting dormancy timing under changing climate conditions, and recognizing that artificial lighting or light pollution may potentially interfere with normal dormancy induction in affected vineyard areas.
Photoperiod_Response represents a major component process within Dormancy_Induction, specifically addressing the detection and response to changing day length. Distinguished from Temperature_Acclimation which responds to thermal signals, though both processes interact during dormancy induction. Subdivides into Short_Day_Perception, Photoperiod_Signal_Transduction, and Growth_Inhibition_Initiation as sequential phases. The primary environmental trigger for dormancy induction in grapevine. Critical photoperiod thresholds vary with cultivar origin latitude.
Photoperioden-Reaktion
Tageslängenreaktion
Tageslängenwahrnehmung
daylength response
daylength sensing
photoperiod perception
photoperiodic response
photoperiodische Reaktion
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Wake CMF, Fennell A (2000) Morphological, physiological and dormancy responses of three Vitis genotypes to short photoperiod. Physiologia Plantarum 109: 203-210 Garner WW, Allard HA (1923) Further studies in photoperiodism. Journal of Agricultural Research 23: 871-920
Photoperiod Response
Photoperiod response in grapevine primarily affects dormancy induction and growth cessation rather than reproductive phenology, with short days (<13-14 hours) triggering periderm development and cold acclimation initiation in autumn. Cultivars originating from higher latitudes (Riesling, 49°N origin) show stronger photoperiod response for dormancy induction than cultivars from lower latitudes (Grenache, 42°N origin), initiating acclimation 2-3 weeks earlier under equivalent day length conditions. Photoperiod interacts with temperature for budbreak timing, with some cultivars showing delayed budbreak under artificially extended photoperiods. Tropical viticulture exploits weak photoperiod response in some cultivars to enable continuous growth cycles.
A phenological plasticity disposition that is the inherited capacity of a Vitis vinifera plant to adjust developmental timing and growth patterns in response to day length (photoperiod), with genotype-specific sensitivity to the duration of light and dark periods affecting dormancy induction, growth cessation, and potentially other phenological transitions.
Photoperiod response in grapevine is generally weaker than in many woody perennials, with temperature being the dominant driver of most phenological events. However, photoperiod sensitivity contributes to latitudinal adaptation patterns, with cultivars from northern origins showing stronger responses that prevent premature deacclimation during warm winter periods at high latitudes. The VvCO and VvFT genes mediate photoperiod signaling in grapevine, with expression patterns correlating with dormancy transitions. Understanding photoperiod response becomes increasingly important for predicting cultivar performance when transferred across latitudes.
Lichtdauer-Response
Tageslängenempfindlichkeit
day length sensitivity
light duration plasticity
Fennell A, Hoover E (1991) Photoperiod influences growth, bud dormancy, and cold acclimation in Vitis labruscana and V. riparia. Journal of the American Society for Horticultural Science 116:270-273 Grant TNL et al. (2013) Biological processes of cold acclimation concomitant with dormancy release in grape (Vitis vinifera). Canadian Journal of Plant Science 93:1051-1068
Photoperiod Response Disposition
In Merlot vines, photoperiod signal transduction proceeds over 2-3 weeks following short day perception, with increased VvNCED expression in leaves elevating abscisic acid biosynthesis, altered VvFT homolog expression affecting mobile signal production, and subsequent detection of increased ABA and dormancy-associated transcripts in axillary buds indicating successful long-distance signal transmission from leaves to meristematic tissues.
Photoperiod signal transduction in Vitis vinifera encompasses the molecular and cellular processes through which the initial photoperiod perception event is converted into systemic developmental signals that reach and affect meristematic tissues in buds and cambium, ultimately resulting in growth cessation and dormancy establishment. This complex signaling network operates across multiple scales from molecular interactions within perceiving cells to whole-plant hormone gradients that coordinate dormancy throughout the vine. The intracellular component of signal transduction involves interactions between phytochrome signaling and the circadian clock system, with clock components including homologs of CONSTANS, GIGANTEA, and other regulatory proteins integrating photoperiod information with endogenous rhythms to generate appropriately timed outputs. Transcription factor cascades activated by photoperiod signaling include members of the PHYTOCHROME INTERACTING FACTOR family whose activity is regulated by phytochrome-mediated degradation, and downstream factors that activate or repress target genes involved in growth, metabolism, and dormancy. The hormonal dimension of signal transduction involves changes in biosynthesis, transport, and sensitivity of multiple plant hormones, with increasing abscisic acid levels promoting dormancy and stress tolerance while declining gibberellin activity reduces growth promotion, creating the hormonal environment that favors dormancy establishment. Long-distance signaling from perception sites in leaves to responding meristems involves mobile signals transported through the phloem, potentially including small RNAs, transcription factors, and hormone precursors that communicate photoperiod status to distant tissues. The FLOWERING LOCUS T protein and its homologs, which serve as mobile flowering signals in annual plants, may play roles in mediating photoperiod information to grapevine buds, though their specific functions in perennial dormancy systems remain under investigation. Chromatin modifications including changes in histone acetylation and methylation contribute to the establishment of dormancy-related gene expression patterns that persist beyond the initial signal perception. The duration and intensity of short-day exposure affects signal strength, with prolonged exposure generally producing more robust dormancy responses than brief exposure near the critical threshold.
Photoperiod_Signal_Transduction represents the intermediate signaling phase within Photoperiod_Response, connecting Short_Day_Perception to Growth_Inhibition_Initiation. Distinguished from perception which addresses photoreceptor detection, and from growth inhibition which addresses the developmental outcome. Involves both intracellular signaling cascades and long-distance mobile signals. Duration typically spans 2-4 weeks as signals propagate and accumulate. Key molecular players include clock components, transcription factors, and hormones particularly ABA.
Dormanz-Signalweg
Photoperioden-Signaltransduktion
Tageslängen-Signalübertragung
daylength signal transmission
dormancy signaling pathway
photoperiod signaling cascade
photoperiodic signaling
photoperiodische Signalkaskade
Photoperiod Signal Transduction
Photoperiod synchrony assessment comparing cultivar responses to autumn conditions reveals the adaptive significance of photoperiod sensitivity, with photoperiod-sensitive cultivar (strong growth cessation response to shortening days) showing reliable dormancy initiation by October 15 regardless of autumn temperature (maintaining winter preparation schedule even in warm years), while less photoperiod-sensitive cultivar continues growth until temperature decline forces cessation, showing dormancy initiation ranging from October 5 (cool autumn) to November 10 (warm autumn), with the variable timing creating winter hardiness risk in warm autumns followed by early cold events, demonstrating how photoperiod synchrony provides seasonal reliability that temperature-dependent responses cannot match.
Photoperiod synchrony quality is an environmental synchronization quality that represents the alignment between grapevine developmental responses and day length signals, characterizing the coordination of photoperiod-sensitive processes including dormancy induction, growth cessation, and potentially cold acclimation initiation with the astronomical day length patterns that provide the most reliable seasonal timing signal unaffected by year-to-year weather variation. Within the BFO framework, this quality specifically addresses the photoperiod dimension of environmental synchronization, distinguishing day length-coordinated processes from temperature-driven development while recognizing that photoperiod provides the stable, predictable environmental calendar against which vines must coordinate developmental transitions that require advance preparation for seasonal changes (particularly the onset of winter dormancy and cold hardiness). The ontological significance lies in photoperiod synchrony ensuring that vines initiate winter preparation processes in response to shortening days regardless of variable autumn temperatures, where reliance on the astronomically fixed photoperiod signal rather than temperature provides insurance against the consequences of warm autumns that might otherwise delay dormancy preparation until cold damage has already occurred, making photoperiod sensitivity an adaptive feature that coordinates vine development with the reliable seasonal calendar. The grapevine responses to photoperiod include the induction of dormancy and growth cessation (triggered by shortening days in autumn, with critical day length varying among cultivars), the potential modulation of cold acclimation (interacting with temperature for hardiness development), and possible effects on vegetative growth patterns (though less clearly established than dormancy effects), with these responses ensuring that vines prepare for winter before temperature signals alone would trigger preparation. The latitude effects on photoperiod synchrony reflect the variation in day length patterns across latitudes, with high-latitude regions showing more dramatic seasonal photoperiod changes that may provide stronger dormancy signals, while low-latitude regions show minimal photoperiod variation that may reduce the reliability of day length as a seasonal coordinator, potentially requiring greater reliance on temperature signals for developmental timing. The climate change implications of photoperiod synchrony arise from the unchanged astronomical day length signal interacting with altered temperature patterns, potentially creating mismatches between temperature-driven and photoperiod-driven processes that historically were coordinated under stable climate, with warming autumns potentially maintaining growth while shortening days signal dormancy, creating physiological conflicts that may affect vine health and winter survival. Understanding photoperiod synchrony quality enables assessment of dormancy timing reliability, evaluation of cultivar adaptation to different latitudes, and recognition of the climate change implications for processes coordinated by the interaction of temperature and photoperiod signals.
Photoperiod_Synchrony_Quality addresses the alignment of vine development with day length signals providing reliable seasonal coordination. This quality is particularly important for dormancy induction independent of variable temperatures. The class supports dormancy management and latitude adaptation assessment through photoperiod synchrony characterization.
Dormanzzeit-Abstimmung
Photoperiodensynchronie
Tageslängenkoordination
day length coordination
dormancy timing alignment
seasonal light synchrony
Fennell A, Hoover E (1991) Photoperiod influences growth, bud dormancy, and cold acclimation in Vitis labruscana and V. riparia. Journal of the American Society for Horticultural Science 116(2): 270-273 Wake CMF, Fennell A (2000) Morphological, physiological and dormancy responses of three Vitis genotypes to short photoperiod. Physiologia Plantarum 109(2): 203-210
Photoperiod Synchrony Quality
Photorespiration rate assessment in heat-stressed Sauvignon Blanc reveals the temperature effect on photorespiratory carbon loss, with leaves at 25°C showing photorespiration of 3.5 μmol CO₂/m²/s (approximately 22% of gross photosynthesis), while leaves at 38°C display increased photorespiration of 6.2 μmol CO₂/m²/s (approximately 40% of reduced gross photosynthesis), with the disproportionate increase in photorespiratory losses contributing substantially to the net photosynthesis depression under heat stress beyond direct effects on photosynthetic capacity, demonstrating how photorespiration amplifies the impact of high temperature on vine productivity.
Photorespiration rate quality is a respiration rate quality that represents the rate of the light-dependent oxidative photosynthetic carbon cycle initiated by Rubisco oxygenase activity, characterizing the CO₂ release and ATP consumption associated with the recovery of carbon from 2-phosphoglycolate produced when Rubisco fixes O₂ rather than CO₂, representing a significant carbon and energy cost that reduces net photosynthetic efficiency in C₃ plants including grapevine. Within the BFO framework, this quality specifically addresses the photorespiratory flux distinct from mitochondrial dark respiration, distinguishing the light-requiring oxidative cycle from constitutive respiratory metabolism while recognizing that photorespiration represents a substantial carbon loss (typically 20-30% of gross photosynthesis) that fundamentally limits C₃ photosynthetic efficiency. The ontological significance lies in photorespiration representing a major inefficiency of C₃ photosynthesis with profound implications for grapevine productivity, where the competitive inhibition of carboxylation by oxygenation at Rubisco, combined with the carbon and energy costs of photorespiratory metabolism, reduces net carbon gain well below the theoretical maximum, with photorespiratory losses increasing under conditions that favor oxygenation (high temperature, low CO₂, drought-induced stomatal closure). The biochemical basis for photorespiration involves Rubisco oxygenase activity producing 2-phosphoglycolate (rather than 3-phosphoglycerate from carboxylation), which must be converted through the photorespiratory pathway spanning chloroplast, peroxisome, and mitochondrion, with one CO₂ released per two oxygenation events and substantial ATP and reducing power consumed in the recovery process. The environmental factors affecting photorespiration rate include temperature (increasing oxygenation relative to carboxylation at higher temperatures due to differential effects on Rubisco kinetics and O₂/CO₂ solubility), intercellular CO₂ concentration (low Ci from stomatal closure increasing O₂/CO₂ ratio at Rubisco), and O₂ concentration (direct substrate for oxygenation, though constant under natural conditions). The significance of photorespiration for viticulture includes its contribution to photosynthetic depression under heat and drought stress, its role in the temperature response of vine productivity, and its impact on water use efficiency since photorespiratory carbon loss occurs without proportional water saving. Understanding photorespiration rate quality enables assessment of this major photosynthetic inefficiency, prediction of stress impacts on net carbon gain, and appreciation of the fundamental limitation on C₃ photosynthetic efficiency that affects grapevine productivity.
Photorespiration_Rate_Quality addresses the light-dependent oxidative cycle distinct from mitochondrial respiration. This quality represents a major C₃ photosynthetic inefficiency that increases under stress conditions favoring Rubisco oxygenation. The class supports stress impact assessment and photosynthetic efficiency evaluation through characterization of the photorespiratory carbon loss.
C2 cycle
Glykolatweg
Photorespiration
glycolate pathway
oxidative photosynthetic cycle
oxidativer C₂-Zyklus
photorespiratorische Rate
Bauwe H et al. (2010) Photorespiration: players, partners and origin. Trends in Plant Science 15(6): 330-336 Flexas J et al. (2002) Analysis of the relative increase in photosynthetic O₂ uptake when photosynthesis in grapevine leaves is inhibited following low night temperatures and/or water stress. Plant Physiology 129(1): 279-290
Photorespiration Rate Quality
Detailed investigation of Photosynthesis_Heat_Stability_Disposition at the University of California Davis employed temperature-response curves and recovery kinetics to characterize photosynthetic heat tolerance across 20 cultivars. Leaf gas exchange was measured at temperatures from 25°C to 45°C in 5°C increments, with parallel chlorophyll fluorescence monitoring of PSII function. The temperature causing 50% photosynthesis inhibition (T50) and the critical temperature for PSII damage (Tc, defined by rapid Fv/Fm decline) were determined for each cultivar. Photosynthesis_Heat_Stability_Disposition showed significant cultivar variation. 'Grenache' exhibited high T50 of 41.2°C and Tc of 46.5°C, maintaining 70% of maximum photosynthesis at 40°C. 'Riesling' showed lower Photosynthesis_Heat_Stability_Disposition with T50 of 36.8°C and Tc of 43.2°C, retaining only 35% of maximum photosynthesis at 40°C. Analysis of component limitations revealed cultivar-specific patterns. In 'Grenache', stomatal and biochemical limitations contributed equally to high-temperature decline, while in 'Riesling', biochemical limitation (Rubisco deactivation) dominated above 35°C, suggesting less thermostable Rubisco activase. Recovery experiments showed that 'Grenache' restored 95% of pre-stress photosynthesis within 4 hours of return to 25°C, while 'Riesling' recovered only 70% at equivalent timepoint, indicating some irreversible damage. These findings identify Photosynthesis_Heat_Stability_Disposition as a key differentiator among cultivars for performance in hot climates.
A Photosynthesis_Heat_Stability_Disposition is a Heat_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to maintain photosynthetic function at elevated temperatures, preserving carbon assimilation and energy production under heat stress conditions that would otherwise cause stomatal closure, enzyme inactivation, and photosystem damage. This disposition represents the thermostability of the photosynthetic apparatus, with photosynthesis in grapevine showing temperature optima between 25-30°C and progressive inhibition at higher temperatures through multiple mechanisms. Heat-induced photosynthetic decline involves stomatal limitations (reduced CO₂ supply due to stomatal closure), biochemical limitations (Rubisco activase denaturation reducing Rubisco activation state, impaired Calvin cycle enzyme function), and photochemical limitations (PSII damage, thylakoid membrane destabilization, reduced electron transport capacity). Cultivars differ in the temperature threshold for photosynthetic decline, the rate of decline above threshold, and the capacity for recovery following heat exposure. Photosynthesis_Heat_Stability_Disposition depends on thermostable forms of key enzymes (particularly Rubisco activase), heat-tolerant thylakoid membranes, effective photoprotective mechanisms (xanthophyll cycle, antioxidants), and coordinated stomatal behavior balancing CO₂ access against water loss. Strong Photosynthesis_Heat_Stability_Disposition enables continued carbon gain during heat events, supporting fruit ripening and reserve accumulation even under challenging thermal conditions.
Photosynthesis_Heat_Stability_Disposition is_a Heat_Tolerance_Disposition. Photosynthetic heat tolerance involves multiple components including stomatal, biochemical (Rubisco activase thermostability), and photochemical (PSII, thylakoid membrane) factors. T50 (temperature for 50% inhibition) and Tc (critical temperature for PSII damage) are useful quantitative parameters. Heat-induced photosynthetic limitation affects both immediate carbon gain and long-term vine productivity through reduced assimilate supply for fruit and reserve accumulation.
Kohlenstoffassimilations-Hitzestabilität
Photosynthese-Hitzestabilität-Disposition
Photosynthetische-Thermotoleranz
carbon assimilation heat stability
photosynthetic heat resistance
photosynthetic thermotolerance
Berry J, Bjorkman O (1980) Photosynthetic response and adaptation to temperature in higher plants, Annual Review of Plant Physiology 31:491-543 Crafts-Brandner SJ, Salvucci ME (2000) Rubisco activase constrains the photosynthetic potential of leaves at high temperature and CO2, Proceedings of the National Academy of Sciences 97:13430-13435 Greer DH, Weedon MM (2012) Modelling photosynthetic responses to temperature of grapevine leaves on vines grown in a hot climate, Plant, Cell and Environment 35:1050-1064
Photosynthesis Heat Stability Disposition
Photosynthetic capacity assessment in Chardonnay vineyard comparing sun-exposed exterior canopy leaves (Amax 16.5 μmol CO₂/m²/s, Vcmax 95 μmol/m²/s) with interior shaded leaves (Amax 8.2 μmol/m²/s, Vcmax 52 μmol/m²/s) reveals the canopy position effect on capacity development, with shade-developed leaves showing permanently reduced photosynthetic potential that canopy management cannot reverse once leaves are mature, demonstrating the importance of early-season canopy architecture for optimizing whole-vine photosynthetic capacity.
Photosynthetic capacity quality is a physiological quality that represents the maximum potential rate of carbon assimilation by grapevine leaves under optimal conditions, characterizing the biochemical and biophysical limitations on CO₂ fixation that determine the vine's fundamental productivity potential and capacity to supply carbohydrates for fruit ripening and reserve accumulation. Within the BFO framework, this quality addresses the carbon acquisition dimension of vine physiology, distinguishing photosynthetic potential from actual photosynthetic rate (which may be limited by environmental conditions) while recognizing that photosynthetic capacity sets the upper boundary on vine productivity that environmental factors may reduce but cannot exceed. The ontological significance lies in photosynthetic capacity providing the ultimate constraint on grape yield and quality potential, where the vine's ability to fix carbon determines the pool of carbohydrates available for sugar accumulation in berries, synthesis of secondary metabolites, and storage of reserves for subsequent seasons, making photosynthetic capacity a fundamental determinant of sustainable quality production. The physiological basis for photosynthetic capacity includes Rubisco content and activity (limiting carboxylation capacity), electron transport chain components (limiting light reaction capacity), stomatal conductance potential (limiting CO₂ supply), and mesophyll conductance (limiting internal CO₂ diffusion), with the integration of these factors determining maximum assimilation rate typically expressed as light-saturated net photosynthesis (Amax) or parameters derived from photosynthesis-CO₂ response curves (Vcmax, Jmax). The environmental and developmental factors affecting photosynthetic capacity include leaf age (capacity declining from full expansion through senescence), nitrogen nutrition (strongly correlated with Rubisco content), water availability (affecting stomatal and mesophyll conductance), temperature history (influencing enzyme quantity and activity), and light environment during leaf development (determining chloroplast density and composition). Understanding photosynthetic capacity quality enables assessment of vine productivity potential, diagnosis of limitations on carbon supply, and management optimization targeting photosynthetic efficiency for quality grape production.
Photosynthetic_Capacity_Quality serves as the parent class for light capture and carbon fixation component qualities, representing the maximum potential for carbon assimilation. This quality is distinguished from actual photosynthetic rate by representing potential under optimal conditions rather than realized performance under field conditions. The class supports productivity assessment and management targeting photosynthetic optimization.
Amax
Kohlenstoffassimilationspotential
Photosynthesekapazität
carbon assimilation potential
maximale Photosynthese
maximum photosynthesis
photosynthetic potential
photosynthetisches Potenzial
Flexas J et al. (2012) Mesophyll conductance to CO₂ and Rubisco as targets for improving intrinsic water use efficiency in C₃ plants. Plant, Cell & Environment 35(2): 348-365 Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell & Environment 26(8): 1393-1405
Photosynthetic Capacity Quality
Photosynthetic heat tolerance quality comparison across warm and cool climate Vitis vinifera cultivars evaluated the basis for observed productivity differences during heat events affecting California vineyards. Gas exchange measurements across a temperature gradient from 25 to 45°C characterized the photosynthetic response profiles. Carignane, adapted to hot Mediterranean conditions, demonstrated superior photosynthetic heat tolerance quality with net photosynthesis maintaining 75% of optimal rate at 40°C and the temperature optimum for photosynthesis elevated to 32°C reflecting heat adaptation. Pinot Noir, from cooler Burgundian origins, showed inferior photosynthetic heat tolerance quality with net photosynthesis declining to 40% of optimal at 40°C and a lower temperature optimum of 28°C. Chlorophyll fluorescence analysis revealed that the cultivar difference arose primarily from differential Rubisco activation sensitivity, with Pinot Noir Rubisco activase showing deactivation above 35°C while Carignane maintained activation to 42°C. The photosynthetic heat tolerance quality differential had practical consequences during a 5-day heat event exceeding 40°C, with Pinot Noir showing 35% reduction in whole-vine carbon assimilation that delayed fruit ripening by 10 days, while Carignane maintained near-normal assimilation rates supporting continued sugar accumulation without ripening delay.
Photosynthetic heat tolerance quality is a thermal tolerance quality that inheres in a grapevine cultivar or clone and represents the capacity to maintain photosynthetic carbon assimilation under elevated temperature conditions, determined by the thermostability of photosynthetic apparatus components including the thylakoid membrane electron transport chain, the carbon-fixing enzymes of the Calvin cycle, and the regulatory mechanisms that protect photosynthetic function when temperatures exceed optimal ranges. This quality addresses the maintenance of the primary process supporting grapevine productivity and fruit development, recognizing that photosynthetic impairment under heat stress can limit carbohydrate supply for fruit ripening even when visible heat damage is absent, creating quality impacts from sublethal heat exposure that may not be apparent from damage assessment alone. The ontological significance of photosynthetic heat tolerance quality lies in its direct relevance to vine productivity and fruit quality, with photosynthetic sensitivity to heat determining whether carbon assimilation can support fruit development during heat events that may coincide with critical ripening periods when photosynthate demand is high. The biological basis of photosynthetic heat tolerance encompasses the thermostability of photosystem II which is typically the most heat-sensitive component of the electron transport chain, the maintenance of thylakoid membrane integrity required for electron transport coupling, the stability and activation state of Rubisco that may limit carbon fixation at high temperatures before electron transport is compromised, and the heat shock response that produces protective proteins maintaining photosynthetic complex assembly and function. The decline in photosynthesis with increasing temperature reflects the sequential impairment of these processes, with the optimum temperature for grapevine photosynthesis typically around 25-30°C and substantial inhibition occurring above 35-40°C depending on cultivar and acclimation status. Assessment of photosynthetic heat tolerance quality employs gas exchange measurements and chlorophyll fluorescence analysis that quantify photosynthetic function across temperature ranges, identifying the thresholds at which function declines and characterizing the nature of the limitation through analysis of fluorescence parameters and gas exchange responses.
Photosynthetic_Heat_Tolerance_Quality is_a Thermal_Tolerance_Quality, representing the heat resistance of the photosynthetic apparatus and carbon assimilation process. This class addresses functional carbon gain maintenance under heat stress. Temperature optimum shift and the rate at 40°C relative to optimum provide quantitative assessment. PSII (chlorophyll fluorescence) and Rubisco activation represent key limitations. Cultivar origin correlates with tolerance level. The impact on fruit ripening connects photosynthetic tolerance to wine quality outcomes. Distinguished from overall Thermal_Tolerance_Quality by the specific focus on photosynthesis rather than general cellular heat tolerance.
Kohlenstoffassimilations-Hitzetoleranz
PSII heat tolerance
PSII-Hitzetoleranz
Photosynthese-Thermotoleranz
carbon assimilation heat tolerance
photosynthesis heat stability
photosynthetische Hitzetoleranz
Greer DH, Weston C (2010) Heat stress affects flowering, berry growth, sugar accumulation and photosynthesis of Vitis vinifera cv. Semillon grapevines grown in a controlled environment. Functional Plant Biology 37:206-214 Salvucci ME, Crafts-Brandner SJ (2004) Inhibition of photosynthesis by heat stress: the activation state of Rubisco as a limiting factor in photosynthesis. Physiologia Plantarum 120:179-186 Wahid A et al. (2007) Heat tolerance in plants: an overview. Environmental and Experimental Botany 61:199-223
Photosynthetic Heat Tolerance Quality
Photosynthetic limitation analysis in water-stressed Tempranillo partitions the 45% reduction in photosynthetic rate (from 14 to 7.7 μmol CO₂/m²/s) into component contributions, revealing stomatal limitation (Ls) of 58% (stomatal closure reducing CO₂ supply), mesophyll limitation (Lm) of 28% (reduced mesophyll conductance concentrating CO₂ drawdown), and biochemical limitation (Lb) of 14% (moderate metabolic downregulation), demonstrating that management targeting water relations (irrigation timing, deficit management) would address the dominant constraint while biochemical intervention would have limited benefit under this stress scenario.
Photosynthetic limitation quality is a photosynthetic capacity quality that represents the identification and quantification of factors constraining photosynthetic rate below the theoretical maximum, characterizing the relative contributions of stomatal, mesophyll, and biochemical limitations that together determine why actual photosynthesis falls short of potential capacity. Within the BFO framework, this quality addresses the constraint dimension of photosynthetic performance, distinguishing limitation analysis from capacity measurement while recognizing that understanding limitations enables targeted intervention to improve photosynthetic efficiency through management or breeding strategies addressing the dominant constraint. The ontological significance lies in photosynthetic limitation analysis providing the diagnostic framework for understanding why vine productivity falls below potential, where partitioning of total limitation among stomatal, mesophyll, and biochemical components indicates whether management should target water relations (stomatal limitation), leaf anatomy (mesophyll limitation), or nutrition and biochemistry (biochemical limitation), enabling evidence-based intervention for productivity improvement. The theoretical framework for limitation analysis derives from the model of Farquhar and Sharkey, which partitions total limitation into stomatal limitation (Ls, reflecting constraint from stomatal closure), mesophyll limitation (Lm, reflecting CO₂ diffusion resistance from stomata to chloroplasts), and biochemical limitation (Lb, reflecting Rubisco and electron transport capacity constraints), with the sum of limitations explaining the gap between actual and potential photosynthesis. The quantitative assessment of limitations employs A-Ci curve analysis combined with fluorescence measurements and/or isotope discrimination to separate the components, with stomatal limitation calculated from the difference between ambient and internal CO₂, mesophyll limitation from internal CO₂ drawdown to chloroplast, and biochemical limitation from chloroplast CO₂ effects on carboxylation. The environmental and stress effects on limitation partitioning show characteristic patterns, with drought primarily increasing stomatal limitation, heat stress affecting both mesophyll and biochemical limitations, and nutrient deficiency primarily affecting biochemical limitation, enabling stress-specific diagnosis from limitation analysis. Understanding photosynthetic limitation quality enables diagnosis of the primary constraints on vine productivity, targeted management intervention addressing dominant limitations, and breeding target identification for genetic improvement of photosynthetic performance.
Photosynthetic_Limitation_Quality serves as the parent class for specific limitation types (Stomatal, Mesophyll, Biochemical). This quality provides the diagnostic framework for understanding why photosynthesis falls below potential. The class supports targeted intervention through limitation partitioning that identifies the dominant constraint requiring management attention.
Limitierungsanalyse
Ls analysis
Ls-Analyse
diffusional vs biochemical limitation
limitation analysis
photosynthetic constraint
photosynthetische Einschränkung
photosynthetische Limitation
Grassi G, Magnani F (2005) Stomatal, mesophyll conductance and biochemical limitations to photosynthesis as affected by drought and leaf ontogeny in ash and oak trees. Plant, Cell & Environment 28(7): 834-849 Flexas J et al. (2008) Mesophyll conductance to CO₂: current knowledge and future prospects. Plant, Cell & Environment 31(5): 602-621
Photosynthetic Limitation Quality
The historical devastation demonstrating the absence of Phylloxera_Resistance_Disposition in V. vinifera began in the 1860s when phylloxera, inadvertently introduced from North America on botanical specimens, spread through French vineyards with catastrophic consequences. By 1900, over 2.5 million hectares of European vineyards had been destroyed, representing 40% of French production capacity and causing economic losses equivalent to twice the costs of the Franco-Prussian War. The solution—grafting V. vinifera scions onto resistant American rootstocks—was proposed by Jules-Émile Planchon and validated through systematic trials demonstrating that V. riparia, V. rupestris, and V. berlandieri roots support phylloxera populations 100-1000 fold lower than V. vinifera while maintaining vine productivity. Modern rootstock breeding continues to optimize Phylloxera_Resistance_Disposition, with genotypes like 'SO4' (V. berlandieri × V. riparia) supporting minimal root galling (OIV rating 8-9) while providing excellent graft compatibility and adaptation to diverse soil conditions. The universal adoption of resistant rootstocks represents the most successful application of host plant resistance in agricultural history, enabling V. vinifera cultivation to continue in all traditional wine regions despite permanent phylloxera establishment.
A Phylloxera_Resistance_Disposition is a Pest_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that limit feeding, gall formation, population development, or damage by grape phylloxera (Daktulosphaira vitifoliae), the aphid-like insect that caused the catastrophic destruction of European vineyards in the late 19th century and permanently transformed global viticulture through the universal adoption of resistant rootstocks. This disposition represents the capacity to resist a uniquely devastating grapevine pest native to eastern North America, where it co-evolved with indigenous Vitis species that developed effective resistance mechanisms. Phylloxera has a complex life cycle with both root-feeding (radicicole) and leaf-feeding (gallicole) forms, though the root-feeding form causes the economically significant damage to susceptible V. vinifera. On susceptible roots, phylloxera feeding induces characteristic galls (nodosities on young roots, tuberosities on older lignified roots) that disrupt root function and provide entry points for secondary pathogens, leading to progressive vine decline and death. Resistance mechanisms include hypersensitive response killing cells at feeding sites, production of wound periderm isolating feeding damage, antibiosis factors reducing insect survival or fecundity, and antixenosis reducing feeding preference. North American Vitis species including V. riparia, V. rupestris, V. berlandieri, and V. cinerea exhibit varying levels of root phylloxera resistance that has been exploited through rootstock breeding, while V. vinifera remains essentially defenseless against root infestation, necessitating grafting for sustainable production in phylloxera-infested soils worldwide.
Phylloxera_Resistance_Disposition is_a Pest_Resistance_Disposition. This class addresses resistance to Daktulosphaira vitifoliae, historically the most devastating grapevine pest. Child classes distinguish Root_Phylloxera_Resistance_Disposition (economically critical, addressed through rootstocks) from Foliar_Phylloxera_Resistance_Disposition (less damaging, primarily affecting American species and hybrids). The phylloxera crisis of the 1860s-1900s permanently transformed viticulture, making grafted vines the global standard. Resistance is found in North American Vitis species co-evolved with phylloxera V. vinifera lacks meaningful resistance due to absence of evolutionary exposure.
Daktulosphaira resistance
Daktulosphaira-Abwehr
Phylloxera-Resistenz
Reblaus-Resistenz-Disposition
grape phylloxera immunity
radicicole/gallicole resistance
Granett J et al. (2001) Biology and management of grape phylloxera, Annual Review of Entomology 46:387-412 Ordish G (1987) The Great Wine Blight, 2nd Edition, Sidgwick & Jackson Powell KS (2012) A holistic approach to future management of grapevine phylloxera, Root Feeders: An Ecosystem Perspective, CAB International
Phylloxera Resistance Disposition
SO4 (V. berlandieri × V. riparia) provides strong phylloxera resistance through rapid wound periderm formation, supporting Riesling production across the Mosel, Rheingau, and Pfalz regions where phylloxera remains endemic. 3309 Couderc (V. riparia × V. rupestris) combines phylloxera resistance from both parental species, serving as a standard rootstock in Burgundy and Champagne. 110 Richter (V. berlandieri × V. rupestris) confers phylloxera resistance to Tempranillo in Ribera del Duero, Spain, where the pest has been present since the 1880s. The rare exceptions of V. vinifera-containing rootstocks such as ARG1 (Aramon × V. rupestris Ganzin No. 1) show reduced phylloxera resistance under heavy infestation pressure.
A stress-resistance rootstock role that inheres in a grafted rootstock by virtue of its genetically determined capacity to resist feeding damage by grape phylloxera (Daktulosphaira vitifoliae Fitch, Hemiptera: Phylloxeridae). Phylloxera-resistant rootstocks prevent or limit the formation of root galls (nodosities on young roots, tuberosities on lignified roots) through hypersensitive wound responses, rapid wound periderm formation, production of phenolic compounds and tannins toxic to the insect, and anatomical barriers including thick-walled cortical cells and early suberization. Resistance levels range from immunity (no gall formation, complete insect mortality) through strong resistance (limited nodosity formation, no tuberosities, negligible vine damage) to moderate resistance (some gall formation but sufficient tolerance to maintain vine productivity). Phylloxera resistance in rootstocks derives from the coevolutionary history of North American Vitis species with the native pest, particularly V. riparia, V. rupestris, V. berlandieri, and V. cinerea.
Phylloxera_Resistant_Rootstock_Role is subclassed into Root_Phylloxera_Resistant_Rootstock_Role and Foliar_Phylloxera_Resistant_Rootstock_Role, reflecting the distinct biological interactions of the radicicole and gallicole phylloxera life cycle forms. Root phylloxera resistance is the primary selection criterion for commercial rootstocks worldwide, while foliar resistance is relevant in regions where the leaf-galling cycle is economically damaging (e.g., eastern North America, parts of Australia). Some rootstock genotypes show differential resistance to phylloxera biotypes (strains A through G), and resistance may be overcome by adapted biotypes, as documented in California and Australia.
Phylloxera-immune rootstock
Reblaus-tolerante Unterlage
phylloxera-resistant rootstock
phylloxera-tolerant rootstock
reblausresistente Unterlage
reblauswiderstandsfähige Unterlage
Granett J et al. (2001) Biology and management of grape phylloxera, Annual Review of Entomology 46: 387-412 Powell KS (2012) A holistic approach to future management of grapevine phylloxera, in Arthropod Management in Vineyards, Springer pp.: 219-251 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 15-18 Forneck A, Huber L (2009) (A)sexual reproduction – a review of life cycles of grape phylloxera, Journal of Insect Science 9: 1-10
Phylloxera Resistant Rootstock Role
Integrated analysis of Physical_Barrier_Disposition at Hochschule Geisenheim University characterized structural defense parameters across 25 cultivars and correlated them with resistance phenotypes against multiple pathogens. Cuticle properties (thickness, wax load, contact angle as hydrophobicity measure), trichome characteristics (density on adaxial and abaxial surfaces, trichome type distribution), stomatal parameters (density, guard cell dimensions), and leaf toughness (specific leaf area, punch resistance) were quantified on fully expanded leaves. Multiple regression analysis related Physical_Barrier_Disposition parameters to infection severity by Plasmopara viticola (penetrating via stomata), Erysiphe necator (penetrating cuticle directly), and Botrytis cinerea (requiring wound or senescent tissue entry). Stomatal density on abaxial surfaces (range 120-285 stomata/mm²) was the strongest predictor of downy mildew severity (r = 0.71), with low-stomatal cultivars including 'Riesling' (135/mm²) showing 45% less infection than high-stomatal 'Silvaner' (268/mm²). Cuticle thickness (range 1.8-5.2 μm) predicted powdery mildew establishment (r = -0.58), while trichome density showed no relationship with foliar pathogens but correlated significantly with leafhopper resistance (r = -0.65). These findings demonstrate that different Physical_Barrier_Disposition components provide protection against specific attacker guilds based on their infection biology, enabling targeted selection for particular threat profiles.
A Physical_Barrier_Disposition is a Constitutive_Defense_Disposition that inheres in a Vitis vinifera plant and is realized through structural features of plant surfaces and tissues that physically impede pathogen penetration, pest feeding, or attacker establishment independent of chemical toxicity or induced responses. This disposition represents the anatomical and morphological component of constitutive defense, encompassing structures that function as passive barriers by virtue of their physical properties including thickness, toughness, hydrophobicity, or architecture. Major physical barriers in grapevine include the cuticle (cuticular membrane with embedded and epicuticular waxes covering all aerial surfaces), trichomes (epidermal appendages varying in density, morphology, and distribution), stomatal characteristics (density, size, and regulation affecting pathogen entry), cell wall properties (thickness, composition, lignification), and tissue architecture (compactness, air spaces, vascular arrangement). Physical barriers provide non-specific defense effective against diverse attackers sharing requirements for surface attachment, penetration, or tissue access. The effectiveness of physical barriers depends on both quantitative parameters (cuticle thickness, trichome density) and qualitative features (wax crystal morphology, trichome type), with substantial genetic variation among grapevine cultivars and species providing material for breeding enhancement. Physical barriers often interact with chemical defenses, as cuticle and cell walls may contain embedded antimicrobial compounds, and trichomes may produce or store defensive metabolites.
Physical_Barrier_Disposition is_a Constitutive_Defense_Disposition. This class encompasses structural defenses functioning through physical properties rather than chemical toxicity. Child classes include Cuticle_Thickness_Disposition, Trichome_Density_Disposition, and Stomatal_Density_Disposition addressing specific barrier types. The effectiveness of particular barriers depends on attacker biology, with stomatal density most relevant for stomatal-penetrating pathogens (downy mildew), cuticle properties for direct-penetrating pathogens (powdery mildew), and trichomes for arthropod pests.
Anatomische-Barriere
Physikalische-Barriere-Disposition
Strukturelle-Abwehr
anatomical barrier
physical resistance mechanism
structural defense
Underwood W (2012) The plant cell wall: a dynamic barrier against pathogen invasion, Frontiers in Plant Science 3:85 Yeats TH, Rose JKC (2013) The formation and function of plant cuticles, Plant Physiology 163:5-20 Hanley ME et al. (2007) Plant structural traits and their role in anti-herbivore defence, Perspectives in Plant Ecology, Evolution and Systematics 8:157-178
Physical Barrier Disposition
Physical quality assessment in California table grape production integrates firmness measurement (target 250-350 g force in compression), dimensional grading (minimum 16mm berry diameter for premium grades), and damage evaluation (less than 2% splitting, cracking, or mechanical injury) to determine pack-out quality and market destination fruit meeting all physical quality specifications commands premium pricing for export markets demanding extended shelf life and visual perfection, while fruit with marginal physical quality (soft texture, undersized berries, minor damage) is diverted to domestic markets with shorter distribution chains or processed into juice where physical integrity is less critical.
Physical_Quality encompasses all qualities that characterize the mechanical, structural, and dimensional properties of grapevine tissues, capturing the material characteristics that affect handling, processing, storage, and sensory perception independent of chemical composition. Physical qualities describe how grape tissues respond to forces, maintain structural integrity, and exhibit measurable dimensional attributes, providing the complement to chemical composition in comprehensive quality characterization. This category subdivides into Firmness_Quality addressing resistance to deformation and penetration Dimensional_Quality capturing size, shape, and geometric attributes Density_Quality characterizing mass-to-volume relationships and Rheological_Quality describing flow and deformation behavior of extracted juices and wines. Physical qualities arise from the structural organization of plant tissues, including cell wall composition and architecture, turgor pressure, intercellular adhesion, and the proportion and arrangement of different cell types (skin, pulp, vascular tissue). The physical properties of grapes profoundly affect quality outcomes throughout the production chain. Harvest and transport operations impose mechanical stresses that firm berries withstand better than soft, degrading fruit. Crushing and pressing behaviors depend on skin strength, pulp consistency, and juice release characteristics. Table grape quality centers on textural attributes including crispness, firmness, and skin chewiness that determine consumer acceptance. Wine quality is affected by physical properties through their influence on extraction kinetics, juice yield, and the structural integrity that prevents premature oxidation from damaged tissue. Physical quality assessment employs instrumental texture analysis using compression, puncture, and tensile testing geometries, providing objective measurements that complement traditional subjective evaluation. The relationship between physical qualities and chemical composition involves bidirectional interactions, with cell wall chemistry affecting mechanical properties while physical damage affects chemical stability and extraction. Understanding physical quality enables optimization of harvest timing, handling protocols, and processing parameters for different end uses.
Physical_Quality is positioned as a direct subclass of Grapevine_Quality, parallel to Chemical_Quality and capturing the non-compositional material properties of grape tissue. The class integrates concepts from materials science, food texture analysis, and plant physiology applied to viticulture and enology. Physical qualities describe current state properties distinct from dispositions (e.g., susceptibility to damage) and processes (e.g., softening during ripening). The instrumental measurement approach provides objectivity complementing sensory texture assessment, though correlations between instrumental and sensory measures require validation for specific applications.
Physikalische Qualität
Textureigenschaft
material characteristic
mechanical quality
mechanische Qualität
textural property
Rolle L et al. (2012) Instrumental Texture Analysis Parameters as Markers of Table-Grape and Winegrape Quality: A Review. American Journal of Enology and Viticulture 63(1): 11-28 Bourne MC (2002) Food Texture and Viscosity: Concept and Measurement, 2nd Ed, Academic Press
Physical Quality
Physiological maturity assessment in Merlot using tissue-specific indicators reveals the progressive completion of biological ripening, with seed maturity indicated by color transition from green through yellow-brown to dark brown (complete lignification, viable embryos), pulp maturity shown by loss of firmness and ready juice release (cellular breakdown facilitating extraction), and skin maturity evidenced by softened texture and deep color saturation (cell wall modification enabling phenolic extractability), with all three components reaching physiological maturity simultaneously around 24 Brix, demonstrating the favorable cultivar characteristic of synchronized physiological development supporting optimal harvest timing.
Physiological maturity quality is a maturity quality that represents the biological completion of berry development, characterizing the state where seeds have achieved viability, pulp cells have completed their developmental program, and skin tissues have reached full structural and biochemical maturity, indicating that the berry has fulfilled its reproductive function and achieved the developmental endpoints programmed by its genetics. Within the BFO framework, this quality specifically addresses the biological completion dimension of maturity, distinguishing developmental fulfillment from compositional targets while recognizing that physiological maturity reflects the evolutionary purpose of the fruit (seed dispersal) rather than the human purpose (winemaking), though these purposes have been aligned through centuries of selection for cultivars where physiological and technological maturity approximately coincide. The ontological significance lies in physiological maturity establishing the biological foundation upon which technological maturity develops, where seeds that have achieved viability signal completion of the reproductive program, pulp that has completed cellular development provides the matrix for composition accumulation, and skins that have achieved structural maturity determine phenolic extractability and anthocyanin stability, together creating the developmental context within which compositional maturity occurs. The assessment of physiological maturity involves evaluation of seed development (color change from green to brown indicating lignification, loss of chlorophyll, embryo maturation), pulp characteristics (cellular integrity, juice release, texture), and skin properties (cell wall composition, phenolic localization, mechanical properties), with these components potentially reaching physiological maturity at different times within the same berry. The relationship between physiological and technological maturity shows cultivar-specific patterns, with some cultivars (Pinot Noir, Merlot) achieving physiological maturity close to optimal technological maturity, while others (Cabernet Sauvignon in warm climates) may reach excessive sugar levels before physiological maturity, and still others (cool-climate late cultivars) may achieve physiological maturity before adequate sugar accumulation, creating the cultivar-climate matching imperative for quality production. Understanding physiological maturity quality enables assessment of biological ripening completion, interpretation of technological maturity in its developmental context, and recognition of cultivar-climate interactions affecting the synchronization of physiological and technological maturity.
Physiological_Maturity_Quality serves as the parent class for tissue-specific physiological maturity (Seed, Pulp, Skin). This quality addresses the biological completion of berry development underlying technological composition. The class supports harvest assessment through physiological maturity characterization indicating developmental completion.
Entwicklungsreife
berry ripeness
biological maturity
biologische Reife
developmental maturity
physiologische Reife
reproductive maturity
reproduktive Reife
Kennedy J (2002) Understanding grape berry development. Practical Winery & Vineyard 24(4): 14-23 Conde C et al. (2007) Biochemical changes throughout grape berry development and fruit and wine quality. Food 1(1): 1-22
Physiological Maturity Quality
Physiological quality assessment in premium Napa Valley vineyard integrates multiple functional parameters including photosynthetic capacity (net assimilation 12-15 μmol CO₂/m²/s indicating healthy carbon fixation), water status (midday stem water potential -1.0 to -1.2 MPa indicating moderate deficit appropriate for quality), and nutrient status (petiole nitrogen 0.8-1.0% indicating balanced nutrition), with this comprehensive physiological profile predicting optimal fruit composition for premium wine production by confirming vine functional capacity to support complete ripening with appropriate secondary metabolite accumulation.
Physiological quality is a quality that represents the functional status and metabolic capacity of grapevine tissues and organs, characterizing the biochemical and biophysical properties that determine vine health, productivity, and fruit quality potential through the integration of photosynthesis, water relations, nutrient status, and stress responses. Within the BFO framework, this quality addresses the functional dimension of grapevine biology, distinguishing physiological status from morphological structure and phenological timing while recognizing that physiological quality determines the vine's capacity to produce quality fruit through the efficiency of carbon assimilation, resource allocation, and adaptive responses to environmental conditions. The ontological significance lies in physiological quality providing the mechanistic link between environmental inputs and grape composition outcomes, where vine physiological status determines the capacity for sugar accumulation, secondary metabolite synthesis, and flavor precursor development that ultimately define wine quality potential, making physiological assessment essential for precision viticulture and quality-focused vineyard management. The scope of physiological quality encompasses multiple functional domains including photosynthetic capacity (carbon assimilation efficiency), water relations (hydraulic status and drought response), nutrient physiology (mineral acquisition and allocation), hormonal regulation (growth and stress signaling), and source-sink dynamics (carbohydrate partitioning to fruit versus vegetative growth). The measurement approaches for physiological quality include gas exchange analysis (photosynthesis, transpiration, stomatal conductance), chlorophyll fluorescence (photosystem efficiency), water potential assessment (plant water status), tissue analysis (nutrient concentrations), and various spectral and imaging technologies enabling non-destructive physiological monitoring across vineyard scales. The management implications of physiological assessment include irrigation scheduling based on water status, canopy management optimizing photosynthetic efficiency, nutrition programs addressing deficiency or excess, and harvest timing decisions incorporating physiological maturity indicators alongside compositional parameters. Understanding physiological quality enables mechanistic interpretation of vine performance, precision management targeting physiological optimization, and quality prediction based on vine functional status throughout the growing season.
Physiological_Quality serves as the root class for the functional status hierarchy of grapevine qualities, establishing the framework for metabolic and biophysical assessment distinct from morphological structure and phenological timing. This quality encompasses photosynthesis, water relations, nutrition, and stress physiology as integrated functional domains. The class supports precision viticulture through physiological monitoring that enables mechanistic understanding and targeted management intervention.
Funktionszustand
Pflanzenphysiologie
Stoffwechselkapazität
functional status
metabolic capacity
physiologische Qualität
plant function quality
vine physiology
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 1-522 Mullins MG et al. (1992) Biology of the Grapevine. Cambridge University Press, Cambridge, pp. 80-150
Physiological Quality
Investigation of Phytoalexin_Based_Botrytis_Resistance_Disposition at the University of Bordeaux compared stilbene production dynamics in resistant 'Marselan' versus susceptible 'Semillon' following controlled B. cinerea inoculation of véraison-stage berries. Prior to inoculation, 'Marselan' berries contained constitutive resveratrol at 12.4 μg/g fresh weight versus 3.8 μg/g in 'Semillon'. Within 24 hours post-inoculation, 'Marselan' showed 8-fold induction of STS gene expression and rapid accumulation of ε-viniferin (the most fungitoxic stilbene, reaching 45 μg/g by 48 hours), while 'Semillon' exhibited only 2-fold STS induction and minimal viniferin production (8 μg/g). In vitro bioassays confirmed that stilbene extracts from inoculated 'Marselan' berries inhibited B. cinerea conidial germination by 78% and germ tube elongation by 85%, compared to 23% and 31% inhibition by 'Semillon' extracts. Infection severity at 7 days post-inoculation was 18% in 'Marselan' versus 62% in 'Semillon', demonstrating the functional significance of enhanced phytoalexin production capacity for Botrytis resistance under conditions where physical barrier mechanisms were bypassed through artificial inoculation.
A Phytoalexin_Based_Botrytis_Resistance_Disposition is a Botrytis_Resistance_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through the production of antimicrobial secondary metabolites, particularly stilbene phytoalexins, that inhibit Botrytis cinerea growth, sporulation, and tissue colonization. This disposition encompasses both constitutive accumulation of preformed antimicrobial compounds and the induced biosynthesis of phytoalexins in response to pathogen detection or tissue damage. Stilbenes, derived from the phenylpropanoid pathway via stilbene synthase (STS) enzymes, represent the primary phytoalexin class in Vitis, with resveratrol (3,5,4'-trihydroxystilbene) being the central compound from which more fungitoxic derivatives including pterostilbene, viniferins (oxidative dimers and oligomers), and piceid (resveratrol glucoside) are produced. The antifungal activity of stilbenes against B. cinerea involves disruption of membrane integrity, inhibition of conidial germination and germ tube elongation, and interference with enzyme secretion required for tissue maceration. Cultivars with enhanced Phytoalexin_Based_Botrytis_Resistance_Disposition show elevated constitutive stilbene content, more rapid and intense STS gene induction following infection or wounding, and greater capacity to produce highly fungitoxic oxidized stilbenes (viniferins). The disposition is influenced by berry developmental stage, with maximum inducibility typically occurring at véraison, and by environmental factors including UV light exposure that stimulates stilbene accumulation. While phytoalexin-based resistance provides meaningful protection, B. cinerea possesses laccase enzymes capable of stilbene detoxification, enabling virulent isolates to partially overcome this defense mechanism.
Phytoalexin_Based_Botrytis_Resistance_Disposition is_a Botrytis_Resistance_Disposition. This class represents active biochemical defense through antimicrobial compound production, distinct from structural resistance mechanisms (Cluster_Architecture_Based_Resistance_Disposition) and passive barrier functions (Cuticle_Based_Resistance_Disposition). Stilbene phytoalexins are the primary antimicrobial class in Vitis, with viniferins being more fungitoxic than their resveratrol precursor. B. cinerea can partially detoxify stilbenes via laccase enzymes, limiting the efficacy of phytoalexin-based resistance as a sole defense mechanism.
Phytoalexin-basierte Botrytis-Resistenz-Disposition
Resveratrol-vermittelte Resistenz
Stilben-Resistenz
induced chemical defense
resveratrol-mediated resistance
stilbene-based Botrytis resistance
Langcake P, Pryce RJ (1977) The production of resveratrol and the viniferins by grapevines in response to ultraviolet irradiation, Phytochemistry 16:1193-1196 Jeandet P et al. (2002) Phytoalexins from the Vitaceae: biosynthesis, phytoalexin gene expression in transgenic plants, antifungal activity, and metabolism, Journal of Agricultural and Food Chemistry 50:2731-2741 Pezet R et al. (2004) Glycosylation and oxidative dimerization of resveratrol are respectively associated to sensitivity and resistance of grapevine cultivars to downy mildew, Physiological and Molecular Plant Pathology 65:297-303
Phytoalexin Based Botrytis Resistance Disposition
Comprehensive analysis of Phytoalexin_Production_Disposition at the University of Bordeaux characterized stilbene induction across 30 cultivars using standardized UV-C elicitation and Plasmopara viticola challenge. Leaf discs were exposed to UV-C (254 nm, 15 minutes) or inoculated with P. viticola sporangia, then harvested at 24, 48, and 72 hours for stilbene profiling by HPLC-MS. Total stilbene accumulation at 48 hours post-UV ranged from 8 μg/g fresh weight ('Chasselas', low inducer) to 125 μg/g ('Regent', high inducer), representing 15-fold variation in Phytoalexin_Production_Disposition. Profile analysis revealed cultivar-specific patterns: 'Regent' produced predominantly trans-resveratrol (45%) and ε-viniferin (35%), while 'Merzling' accumulated higher proportions of pterostilbene (25%) and δ-viniferin (20%). Following P. viticola inoculation, high-inducing cultivars showed strong negative correlation between stilbene accumulation and sporulation intensity (r = -0.78), confirming biological relevance of Phytoalexin_Production_Disposition for disease resistance. Time-course analysis revealed that induction speed varied independently from maximum accumulation, with 'Solaris' reaching 50% maximum earlier (18 hours) than 'Regent' (28 hours) despite lower final concentrations, suggesting that both kinetic and quantitative parameters contribute to effective phytoalexin-based defense.
A Phytoalexin_Production_Disposition is an Induced_Defense_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to synthesize and accumulate phytoalexins—low molecular weight antimicrobial compounds produced de novo or dramatically upregulated in response to pathogen attack, stress, or elicitor treatment. This disposition represents one of the most important inducible chemical defenses in grapevine, which is notable for producing stilbene-type phytoalexins including resveratrol, pterostilbene, and various viniferins that exhibit potent antifungal and antimicrobial activities. Phytoalexin production in Vitis is triggered by pathogen recognition, elicitor perception (including oligosaccharides, lipids, and proteins), UV irradiation, wounding, and abiotic stresses, with the magnitude and kinetics of response varying substantially among genotypes. The biosynthesis of stilbene phytoalexins involves induction of stilbene synthase (STS) genes, of which grapevine possesses an unusually large family (approximately 40 members), enabling rapid and massive stilbene accumulation upon challenge. Phytoalexins function through multiple mechanisms including direct antimicrobial activity via membrane disruption and enzyme inhibition, antioxidant activity protecting plant cells from pathogen-induced oxidative damage, and potential signaling roles in systemic defense activation. The strength of Phytoalexin_Production_Disposition correlates with disease resistance in many grapevine pathosystems, with resistant cultivars typically showing faster induction, higher accumulation, and more diverse phytoalexin profiles compared to susceptible genotypes.
Phytoalexin_Production_Disposition is_a Induced_Defense_Disposition. Grapevine phytoalexins are predominantly stilbenes, synthesized via the phenylpropanoid pathway through the action of stilbene synthase (STS). Child classes address specific aspects of stilbene phytoalexin production including Stilbene_Synthase_Induction_Disposition (enzyme induction), Resveratrol_Production_Disposition (primary stilbene product), and Viniferin_Production_Disposition (oligomeric stilbenes). The large STS gene family in Vitis (unique among plants) enables exceptional stilbene production capacity, making phytoalexins particularly important in grapevine defense.
Antimikrobielle-Verbindungs-Induktion
Induzierbare-Stilben-Kapazität
Phytoalexin-Produktions-Disposition
antimicrobial compound induction
inducible phytoalexin capacity
stilbene phytoalexin production
Jeandet P et al. (2002) The production of resveratrol (3,5,4′-trihydroxystilbene) by grape berries in different developmental stages, American Journal of Enology and Viticulture 42:41-46 Langcake P, Pryce RJ (1977) The production of resveratrol and the viniferins by grapevines in response to infection or injury, Physiological Plant Pathology 9:77-86 Chong J et al. (2009) The stilbene synthase gene family in grapevine: genome-wide identification and expression analyses during development and stress responses, BMC Plant Biology 9:54
Phytoalexin Production Disposition
Phytoalexin production quality comparison across Vitis species following Botrytis cinerea inoculation established the relationship between stilbene accumulation and gray mold resistance. Leaf disc bioassays with B. cinerea inoculation combined with HPLC quantification of stilbene production characterized the kinetics and magnitude of phytoalexin production quality responses. Vitis labrusca showed superior phytoalexin production quality with total stilbene accumulation reaching 850 μg/g fresh weight by 72 hours post-inoculation, dominated by ε-viniferin (45%) and pterostilbene (25%) known to have high antifungal activity, with lesion development limited to 3.2 mm diameter. Vitis vinifera Chardonnay showed moderate phytoalexin production quality with 320 μg/g total stilbenes at 72 hours, dominated by resveratrol (60%) with lower proportions of the more active derivatives, and lesion development reaching 8.5 mm. The V. vinifera cultivar Thompson Seedless exhibited poor phytoalexin production quality with only 85 μg/g total stilbenes, delayed accumulation not reaching maximum until 96 hours, and lesion development exceeding 15 mm. Correlation analysis established strong relationship between phytoalexin production quality (total stilbenes at 48 hours) and B. cinerea lesion limitation (r² = 0.82), with the proportion of pterostilbene and viniferins providing additional explanatory power for resistance variation. The phytoalexin production quality assessment identified both accumulation rate and stilbene profile as determinants of defense effectiveness, informing strategies for enhancing resistance through STS overexpression or elicitor-based induction of endogenous phytoalexin production.
Phytoalexin production quality is a defense response quality that inheres in a grapevine organism and represents the capacity to synthesize antimicrobial secondary metabolites in response to pathogen attack, with stilbenes including resveratrol, pterostilbene, and viniferins representing the major phytoalexin class in grapevine that accumulate at infection sites to directly inhibit pathogen growth through disruption of membrane function, interference with pathogen metabolism, and other toxic effects. This quality captures the chemical defense dimension of plant immunity, recognizing that phytoalexin accumulation provides a direct mechanism for pathogen inhibition that complements the physical containment provided by hypersensitive response and cell wall reinforcement. The ontological significance of phytoalexin production quality lies in its contribution to quantitative disease resistance, where cultivars with greater capacity for rapid and abundant phytoalexin production show enhanced resistance even in the absence of major resistance genes, and where the phytoalexin pathway provides targets for metabolic engineering and elicitor-based resistance enhancement strategies. The biological basis of phytoalexin production in grapevine encompasses the activation of the stilbene synthase (STS) gene family encoding the key enzyme diverting phenylpropanoid metabolism toward stilbene production, the coordinated upregulation of upstream pathway genes providing precursor supply, and the enzymatic modifications producing the diversity of stilbene structures with varying antifungal potency. The grapevine genome contains an expanded STS gene family with approximately 40 members reflecting the evolutionary importance of stilbene-based defense in this species, with expression of multiple STS genes induced by pathogen infection, UV radiation, and various elicitor treatments. The specific stilbenes produced and their relative proportions affect defense effectiveness, with the methylated derivative pterostilbene showing greater antifungal activity than resveratrol, and the oligomeric viniferins produced by oxidative coupling showing the highest potency against major grapevine pathogens.
Phytoalexin_Production_Quality is_a Defense_Response_Quality, representing the induced synthesis of antimicrobial stilbenes (resveratrol, pterostilbene, viniferins) at infection sites. This class addresses the chemical defense dimension of grapevine immunity. The ~40-member STS gene family reflects evolutionary importance of stilbene defense. Pterostilbene and viniferins show higher antifungal activity than resveratrol. Both accumulation rate and stilbene profile determine effectiveness. Quantitative contribution to resistance independent of major R genes. Provides targets for metabolic engineering and elicitor-based enhancement.
Phytoalexin-Produktion
Resveratrol-Abwehr
Stilben-Produktion
induced antimicrobial production
induzierte antimikrobielle Synthese
phytoalexin synthesis
resveratrol defense
stilbene production
Jeandet P et al. (2002) Phytoalexins from the Vitaceae: biosynthesis, phytoalexin gene expression in transgenic plants, antifungal activity, and metabolism. Journal of Agricultural and Food Chemistry 50:2731-2741 Schnee S et al. (2013) Vitis vinifera canes, a new source of antifungal compounds against Plasmopara viticola, Erysiphe necator, and Botrytis cinerea. Journal of Agricultural and Food Chemistry 61:5459-5467 Chong J et al. (2009) The WRKY transcription factor superfamily: its origin in eukaryotes and expansion in plants. BMC Evolutionary Biology 9:196
Phytoalexin Production Quality
Dog Ridge (V. champinii) demonstrates substantial Phytophthora cinnamomi resistance with lesion lengths 60-70% shorter than susceptible Kober 5BB in controlled inoculation trials, and supports Sultana (Thompson Seedless) production on heavy clay soils subject to seasonal waterlogging in the Murray-Darling Basin, Australia. Ramsey (Salt Creek, V. champinii) shows moderate P. cinnamomi resistance combined with excellent nematode resistance, providing multi-stress tolerance on poorly drained sandy loams in the Riverina. 110 Richter (V. berlandieri × V. rupestris) demonstrates moderate Phytophthora tolerance in Mediterranean conditions, maintaining Cabernet Sauvignon health on seasonally wet clay soils in the Maipo Valley, Chile.
A fungal disease-resistance rootstock role that inheres in a grafted rootstock by virtue of its capacity to resist infection by oomycete pathogens of the genus Phytophthora, particularly P. cinnamomi, P. megasperma, P. cryptogea, and P. nicotianae affecting grapevine roots and crown. Although Phytophthora species are taxonomically oomycetes rather than true fungi, they cause fungus-like root rot diseases and are conventionally grouped with fungal root pathogens in viticultural management. Phytophthora zoospores are chemotactically attracted to root exudates and encyst on root surfaces, penetrating through unwounded epidermis and colonizing cortical tissue to cause water-soaked lesions that progress to root necrosis. Resistant rootstocks impede Phytophthora colonization through rapid hypersensitive response, production of phytoalexins (particularly stilbenes), elevated constitutive phenolic content, and anatomical barriers including thick-walled hypodermal cells. Resistance is assessed by lesion length following zoospore inoculation, root biomass retention, and vine survival under prolonged waterlogging where Phytophthora pressure intensifies.
Phytophthora_Resistant_Rootstock_Role addresses an oomycete pathogen complex that is particularly damaging under wet soil conditions and climate change scenarios with increasing extreme precipitation events. Resistance screening typically focuses on P. cinnamomi as the most aggressive and widespread species, though rootstock responses may differ across Phytophthora species. V. champinii-derived rootstocks show enhanced Phytophthora tolerance, possibly linked to their adaptation to seasonally flooded Texas river bottomlands. Integration of Phytophthora resistance with waterlogging tolerance (an abiotic stress) is important as both stress factors often co-occur.
Phytophthora resistant rootstock
Phytophthora tolerant rootstock
Phytophthora-resistente Unterlage
Wurzelfäule-resistente Unterlage
gegen Phytophthora resistente Unterlage
oomycete resistant rootstock
Marais PG (1979) Fungi associated with root rot in vineyards in the Western Cape, Phytophylactica 11: 65-68 Browne GT et al. (2013) Resistance to Phytophthora and Pythium species among grape rootstocks, Phytopathology 103(S2): 21 Latorre BA et al. (1997) Root rot of grapevine in Chile caused by Phytophthora cryptogea and P. megasperma, Plant Disease 81: 684 Erwin DC, Ribeiro OK (1996) Phytophthora Diseases Worldwide, APS Press
Phytophthora Resistant Rootstock Role
Comparative evaluation of Phytoplasma_Resistance_Disposition at the University of Udine, Italy, monitored Flavescence dorée and Bois noir incidence and symptom expression across 25 cultivars in a naturally infected vineyard over 8 seasons. Infection status was confirmed annually by nested PCR targeting phytoplasma 16S rRNA genes, while symptom severity was scored using a 0-5 scale. Substantial cultivar variation in Phytoplasma_Resistance_Disposition was observed for both diseases. For Flavescence dorée, 'Chardonnay' showed high susceptibility with 45% cumulative infection, severe symptoms (mean score 4.2), and 15% vine mortality. 'Merlot' demonstrated enhanced tolerance with equivalent infection rate (42%) but markedly reduced symptom severity (score 2.1) and spontaneous recovery in 35% of infected vines. For Bois noir, 'Sangiovese' exhibited extreme susceptibility (55% infection, score 4.5, 22% mortality) while 'Cabernet Sauvignon' showed relative tolerance (38% infection, score 1.8, 5% mortality). Critically, cultivar rankings differed between diseases, with 'Merlot' tolerant to FD but susceptible to BN (score 3.8), indicating that Phytoplasma_Resistance_Disposition involves pathogen-specific components rather than general anti-phytoplasma mechanisms. No cultivar showed immunity to either phytoplasma.
A Phytoplasma_Resistance_Disposition is a Pathogen_Resistance_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through defense processes that limit infection, multiplication, systemic spread, or symptom expression by phytoplasmas, wall-less prokaryotes of the class Mollicutes that cause grapevine yellows diseases. This disposition represents the capacity to resist a unique class of pathogens that are obligate parasites of plant phloem tissues and insect vectors, unculturable in artificial media, and transmitted by phloem-feeding leafhoppers in a persistent-propagative manner. Major grapevine yellows diseases include Flavescence dorée (FD) caused by 'Candidatus Phytoplasma vitis' and Bois noir (BN) caused by 'Candidatus Phytoplasma solani', which produce similar symptoms including leaf rolling and discoloration (yellowing in white cultivars, reddening in red cultivars), incomplete lignification of canes, shriveled berries, and progressive vine decline. Phytoplasma resistance mechanisms in plants potentially include interference with phytoplasma multiplication in phloem sieve elements, limitation of systemic spread through the sieve tube network, tolerance to phytoplasma-induced phloem dysfunction, and resistance to leafhopper vector feeding preventing pathogen acquisition or inoculation. True resistance to phytoplasmas is poorly characterized in Vitis, though substantial variation in symptom expression and recovery capacity exists among cultivars. Vector resistance represents an important complementary strategy, as interrupting the phytoplasma-vector relationship can effectively reduce disease incidence even in phytoplasma-susceptible cultivars. Phytoplasma diseases have emerged as major threats to European viticulture, with Flavescence dorée subject to mandatory control measures in EU member states due to its epidemic potential.
Phytoplasma_Resistance_Disposition is_a Pathogen_Resistance_Disposition. Phytoplasmas occupy a unique position as prokaryotes (bacteria-like) that are obligate intracellular parasites with characteristics (wall-less, phloem-limited, insect-transmitted) distinct from both bacteria and viruses. Child classes include Flavescence_Doree_Resistance_Disposition and Bois_Noir_Resistance_Disposition for the two major grapevine yellows diseases in Europe. The distinction between "resistance" (limiting infection/multiplication) and "tolerance" (limiting symptoms despite infection) and "recovery" (symptom remission with pathogen persistence) is particularly important for phytoplasma diseases where true resistance is rare.
Mollicute resistance
Mollicuten-Resistenz
Phytoplasmen-Resistenz-Disposition
Vergilbungskrankheit-Resistenz
grapevine yellows resistance
phytoplasma tolerance
Bertaccini A et al. (2014) Phytoplasmas and phytoplasma diseases: a severe threat to agriculture, American Journal of Plant Sciences 5:1763-1788 Constable FE (2010) Phytoplasma epidemiology: grapevines as a model, Phytoplasmas: Genomes, Plant Hosts and Vectors, CAB International Romanazzi G, Murolo S (2008) Grapevine yellows in Italy: past, present, and future, Journal of Plant Pathology 90:S2.303
Phytoplasma Resistance Disposition
Picking ease quality evaluation in Burgundy compared hand harvest productivity across Pinot Noir clones exhibiting varying peduncle characteristics. Clone 115 demonstrated excellent picking ease quality with slender peduncles of 3.5 mm diameter, moderate attachment force of 220 g, and clean cutting response enabling experienced pickers to achieve productivity of 180 kg per hour with minimal fatigue accumulation over full harvest days. Clone 777 showed good picking ease quality with slightly thicker peduncles of 4.2 mm and attachment force of 290 g, achieving productivity of 165 kg per hour. Clone 828 exhibited reduced picking ease quality with robust peduncles of 5.1 mm diameter, firm attachment force of 385 g, and fibrous tissue requiring sawing rather than clean cutting motion, limiting productivity to 130 kg per hour with increased picker fatigue reports and elevated incidence of incomplete peduncle removal leaving stubs that complicated subsequent cluster handling. The picking ease quality differential translated to labor cost differences of €0.08 per kilogram between most and least efficient clones, representing significant economic impact across large production volumes.
Picking ease quality is a harvest logistics quality that inheres in a grapevine cultivar or clone and represents the facility with which grape clusters can be detached from the vine during harvest operations, determined by peduncle structural characteristics, attachment force to the fruiting cane or spur, cutting resistance of the peduncle tissue, and the clarity of the separation point that enables clean removal without damaging adjacent clusters, retained fruit, or vine tissues. This quality directly influences harvest labor productivity and picker fatigue, with easily-picked cultivars enabling rapid, efficient harvest at lower per-unit labor cost while difficult cultivars slow operations, increase physical strain on workers, and may cause collateral damage from the additional force or manipulation required for cluster removal. The ontological significance of picking ease quality lies in its substantial impact on harvest labor costs that represent a major component of production expense, with cultivar differences in picking ease translating directly to economic outcomes that influence variety selection decisions particularly in regions with high labor costs or limited seasonal labor availability. The biological basis of picking ease quality encompasses the anatomical characteristics of the peduncle including diameter, tissue density, fiber content, and lignification degree that determine cutting resistance the strength of the abscission zone or attachment point where the peduncle connects to the rachis of the fruiting structure the brittleness versus flexibility of peduncle tissue affecting whether clean cuts or tearing predominates and the positioning of the peduncle relative to the cluster body and adjacent vine structures affecting cutting tool access. Peduncle attachment force, measured as the pull force required to detach clusters from the vine, typically ranges from 200 g for easily-detached cultivars to over 500 g for firmly-attached types, with forces above 400 g generally considered problematic for efficient hand harvest. The relationship between picking ease quality and harvest method influences its practical significance, with peduncle characteristics mattering greatly for hand harvest but less for mechanical systems that detach fruit through shaking mechanisms rather than cutting. Assessment of picking ease quality involves standardized measurement of peduncle attachment force, observation of cutting behavior under typical tool and technique conditions, and quantification of harvest productivity rates under controlled comparison conditions.
Picking_Ease_Quality is_a Harvest_Logistics_Quality, representing the facility of cluster removal from the vine during harvest. This class addresses peduncle characteristics determining cutting and detachment behavior. Attachment force in grams provides quantitative metric with 400 g challenging. Distinguished from Mechanical_Harvesting_Suitability_Quality which addresses machine detachment through different mechanisms. Most relevant for hand harvest operations where peduncle characteristics directly influence labor productivity and worker ergonomics.
Ernte-Ablöseeffizienz
Pflückleichtigkeit
Stielschnitt-Leichtigkeit
Traubenabtrennung-Leichtigkeit
cluster detachment ease
harvest removal facility
peduncle cutting ease
picking efficiency
Coombe BG, Iland P (2004) Grape berry development and winegrape quality. In: Dry PR, Coombe BG (eds) Viticulture Volume 1 Resources, 2nd Edition, Winetitles, Adelaide, pp. 210-248 Galet P (2000) General Viticulture. Oenoplurimedia, Chaintre, pp. 65-78 May P (2004) Flowering and Fruitset in Grapevines. Lythrum Press, Adelaide
Picking Ease Quality
Long-term field evaluation of Pierces_Disease_Resistance_Disposition at the UC Davis Oakville experimental vineyard compared disease development across resistant breeding selections and susceptible controls under natural glassy-winged sharpshooter (Homalodisca vitripennis) transmission pressure. Over 10 years, susceptible V. vinifera 'Chardonnay' experienced 92% mortality with survivors showing severe chronic symptoms, bacterial titers of 10⁷-10⁸ CFU/g petiole, and near-zero productivity. The PdR1-carrying breeding selection 94-27-008 (V. arizonica × 'Chardonnay' BC₁) demonstrated strong Pierces_Disease_Resistance_Disposition with 0% mortality, no visible symptoms, bacterial titers below 10⁴ CFU/g (often below detection), and yields equivalent to disease-free controls (8.2 tonnes/ha). The resistant selection 'U0505-01' exhibited intermediate response with occasional mild leaf scorch, titers of 10⁴-10⁵ CFU/g, but maintained commercially viable productivity (6.8 tonnes/ha). Importantly, even asymptomatic resistant selections tested positive for X. fastidiosa by PCR in 15-30% of samples, indicating that Pierces_Disease_Resistance_Disposition operates primarily through limitation of bacterial multiplication and spread rather than complete immunity to infection. This resistance mechanism enables sustainable viticulture in Pierce's disease-endemic regions of California, Texas, and the southeastern United States where susceptible V. vinifera cultivation is otherwise impossible.
A Pierces_Disease_Resistance_Disposition is a Bacterial_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that limit infection, multiplication, systemic spread, or symptom development following inoculation with Xylella fastidiosa subsp. fastidiosa, the xylem-limited bacterium causing Pierce's disease of grapevine. This disposition represents the capacity to resist one of the most devastating bacterial diseases of grapevine, endemic to the Americas where X. fastidiosa co-evolved with native Vitis species and is transmitted by xylem-feeding sharpshooter leafhoppers (Cicadellidae: Cicadellini) and spittlebugs (Cercopidae). Pierce's disease causes progressive vine decline characterized by irregular leaf scorch beginning at margins, "matchstick" petioles with abscised leaf blades, irregular cane maturation with green islands surrounded by mature bark, and vine death typically within 1-5 years of infection in susceptible V. vinifera cultivars. The disease mechanism involves bacterial multiplication in xylem vessels, biofilm formation, and progressive vessel occlusion blocking water transport, with symptom severity correlating with bacterial population levels. Pierce's disease resistance dispositions in Vitis operate through multiple mechanisms including restriction of bacterial multiplication, limitation of systemic spread through xylem, tolerance to vessel occlusion maintaining hydraulic function, and potentially vector-mediated resistance affecting transmission efficiency. Wild North American Vitis species native to regions with endemic X. fastidiosa exhibit strong resistance, with V. arizonica, V. aestivalis, V. shuttleworthii, and V. smalliana providing sources for resistance breeding. The PdR1 locus from V. arizonica represents the most advanced breeding achievement, enabling development of Pierce's disease resistant wine grape cultivars.
Pierces_Disease_Resistance_Disposition is_a Bacterial_Resistance_Disposition. Pierce's disease is caused by Xylella fastidiosa subsp. fastidiosa, a xylem-limited bacterium transmitted by sharpshooter leafhoppers. Child classes include PdR1_Mediated_Resistance_Disposition (major gene resistance), Xylella_Multiplication_Resistance_Disposition (limiting bacterial population), and Xylem_Occlusion_Tolerance_Disposition (maintaining hydraulic function despite infection). Strong resistance exists in native North American Vitis species co-evolved with X. fastidiosa, in marked contrast to crown gall where no effective resistance has been identified. The disease limits V. vinifera cultivation throughout the southeastern United States and increasingly in California.
PD resistance
Pierce-Krankheit-Resistenz-Disposition
Pierce-Resistenz
Xylella resistance disposition
Xylella-fastidiosa-Resistenz
sharpshooter-transmitted disease resistance
Hopkins DL, Purcell AH (2002) Xylella fastidiosa: cause of Pierce's disease of grapevine and other emergent diseases, Plant Disease 86:1056-1066 Krivanek AF et al. (2006) Resistance to Xylella fastidiosa in North American Vitis species, Plant Disease 90:1503-1508 Walker MA, Tenscher A (2017) Breeding Pierce's disease resistant winegrapes, Acta Horticulturae 1157:191-196
Pierces Disease Resistance Disposition
Pierce's disease symptom quality diagnosis in Temecula Valley Chardonnay documented the characteristic symptom progression enabling field identification prior to laboratory confirmation. Initial Pierce's disease symptom quality appeared in late July as marginal leaf necrosis on mature leaves, with the scorched tissue sharply delimited from adjacent green tissue rather than showing the gradual chlorosis-to-necrosis transition typical of salt or drought stress, and with characteristic green islands of healthy tissue sometimes persisting amid the advancing necrosis. By August, affected vines showed the pathognomonic "matchstick" symptom with leaf blades abscising while the dead, dry petioles remained firmly attached to the shoot, creating clusters of petiole stubs diagnostic for Pierce's disease. Shoot Pierce's disease symptom quality showed irregular bark maturation with alternating green and mature brown sectors along the same internode, described as the "green-stripe" or "tiger-stripe" bark pattern distinct from normal uniform lignification. Berry Pierce's disease symptom quality on affected canes showed failure to ripen normally, with clusters remaining small, hard, and acidic while fruit on healthy portions of the same vine ripened normally. The Pierce's disease symptom quality progression from initial marginal scorch through matchstick formation to vine decline occurred over 3 seasons, with ELISA testing confirming X. fastidiosa presence in symptomatic tissues and guiding the roguing program that removed 12% of the vineyard to limit disease spread.
Pierce's disease symptom quality is a bacterial infection indicator quality that inheres in grapevine organisms infected by the xylem-limited bacterium Xylella fastidiosa and represents the characteristic progressive decline symptoms including marginal leaf necrosis (scorch) with adjacent green tissue creating distinctive "green islands," the persistence of dead leaf petioles on shoots after blade abscission producing "matchstick" appearance, irregular lignification of shoots showing green islands amid mature brown bark, delayed or incomplete fruit ripening, and progressive vine decline leading to death typically within 1-5 years of initial infection depending on cultivar susceptibility. This quality captures the diagnostic features of one of the most devastating bacterial diseases of grapevine, recognizing that Pierce's disease causes complete vine death with no effective treatment once infection is established, making early detection essential for roguing infected vines and implementing vector management to protect remaining healthy vines. The ontological significance of Pierce's disease symptom quality lies in its role enabling diagnosis of a lethal, incurable disease where symptom recognition triggers immediate management response including vine removal to reduce inoculum and intensive control of sharpshooter insect vectors that transmit the bacterium. The biological basis of Pierce's disease symptoms encompasses the colonization and eventual blockage of xylem vessels by X. fastidiosa bacterial colonies and their associated biofilm, tyloses, and gums that progressively obstruct water transport, with the resulting water stress causing the characteristic leaf scorch symptoms that advance from the margin inward as vessel blockage increases. The vector transmission biology critically affects disease epidemiology, with glassy-winged sharpshooter (Homalodisca vitripennis) and other xylem-feeding insects acquiring bacteria during feeding on infected plants and transmitting to healthy vines during subsequent feeding, creating rapid disease spread in regions with high vector populations and warm winters permitting vector survival. The geographic limitation of Pierce's disease to regions with warm winter temperatures reflects both vector survival requirements and the cold-curing phenomenon where extended exposure to temperatures below 5°C can eliminate X. fastidiosa from vine tissues, explaining the restriction of economically significant Pierce's disease to southern United States, Mexico, Central America, and recently to Mediterranean Europe where the bacterium has emerged as a major threat.
Pierces_Disease_Symptom_Quality_(Xylella_fastidiosa) is_a Bacterial_Infection_Indicator_Quality, representing the xylem-blockage symptoms caused by Xylella fastidiosa colonization. The "matchstick" petioles and "green islands" in scorched leaves are pathognomonic. Sharpshooter vector transmission creates rapid spread potential in warm climates. Cold-curing limits disease to warm-winter regions (winter <5°C kills bacterium). No treatment exists roguing and vector management are only options. Progressive to lethal within 1-5 years depending on cultivar.
Pierce's disease symptoms
Pierce-Krankheit-Symptome
Rebenbakterienbrand
Xylella symptoms
Xylella-Symptome
bacterial leaf scorch
bakterielle Blattdürre
leaf scorch disease
Hopkins DL, Purcell AH (2002) Xylella fastidiosa: cause of Pierce's disease of grapevine and other emergent diseases. Plant Disease 86:1056-1066 Purcell AH, Saunders SR (1999) Fate of Pierce's disease strains of Xylella fastidiosa in common riparian plants in California. Plant Disease 83:825-830 Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests, 2nd Edition. APS Press
Pierces Disease Symptom Quality (Xylella fastidiosa)
Pigmentation intensity in Alicante Bouschet, a teinturier cultivar with colored flesh as well as skin, reaches extraordinary levels of 4000-6000 mg/kg total anthocyanins at harvest, approximately ten-fold higher than Cabernet Sauvignon (400-800 mg/kg) grown in adjacent vineyard blocks this extreme intensity historically made Alicante Bouschet valuable for blending to enhance color in regions where other varieties failed to achieve adequate pigmentation, though modern winemaking increasingly values its unique properties for single-variety wines with exceptional color stability.
Pigmentation_Intensity_Quality represents the degree of color saturation or depth resulting from the concentration and optical effectiveness of pigment molecules in grapevine tissues. Intensity captures the strength of coloration independent of hue, distinguishing deeply pigmented organs from pale or lightly colored organs of the same basic color type. In grape berries, pigmentation intensity varies enormously among cultivars, from the intensely colored teinturier varieties with anthocyanin concentrations exceeding 5000 mg/kg fresh weight to lightly colored pink cultivars with concentrations below 100 mg/kg. Within any cultivar, intensity varies with environmental conditions, canopy management, and crop load. High light exposure in the fruit zone promotes anthocyanin synthesis in colored cultivars, while excessive temperature can degrade pigments and reduce intensity. Water deficit generally increases pigmentation intensity through concentration effects and direct stimulation of anthocyanin biosynthesis genes. Pigmentation intensity assessment employs visual scoring against reference standards (OIV descriptors use categorical scales from very weak through very strong), colorimetric measurement of chroma (C* = √(a*² + b*²) in CIELAB space), or chemical quantification of total pigment concentration. The relationship between pigment concentration and perceived intensity follows a saturating function where additional pigment produces progressively smaller increases in perceived color depth. Pigmentation intensity significantly affects wine color stability because higher initial concentrations provide greater buffering against color loss during aging and the formation of stable polymeric pigments.
Pigmentation_Intensity_Quality addresses the quantitative aspect of coloration (how much color) as distinct from Hue_Quality (what type of color). Intensity correlates with but is not identical to pigment chemical concentration because optical factors including cell wall properties, vacuolar pH, and co-pigmentation affect the color expression of any given pigment quantity. OIV descriptors for color intensity typically employ five-category scales (1-3-5-7-9) that can be calibrated against instrumental measurements. The term chroma in color science corresponds closely to pigmentation intensity in plant biology contexts. Very high pigmentation intensity in berries may indicate water stress, excessive sun exposure, or disease (certain viral infections increase anthocyanin accumulation).
Pigmentierungsintensität
Pigmentkonzentrationsqualität
color intensity
pigment concentration quality
saturation quality
Downey MO et al. (2006) Cultural Practice and Environmental Impacts on the Flavonoid Composition of Grapes and Wine: A Review of Recent Research. American Journal of Enology and Viticulture 57(3): 257-268 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed, John Wiley and Sons pp.: 141-203
Pigmentation Intensity Quality
Cabernet Sauvignon pigmented berry skin at maturity contains 3.2-4.5 mg/g fresh weight anthocyanins (malvidin-3-glucoside 52-58%, other monoglucosides and acylated forms) concentrated in 2-3 hypodermal cell layers creating deep purple-black berry appearance, providing sufficient anthocyanins to produce wines with 12-18 OD units color intensity at 520 nm when extracted during 14-21 day maceration, with pigmented skin representing the sole source of wine color as flesh and seeds contain essentially no anthocyanins. Syrah deeply pigmented berry skin showing high anthocyanin characteristics= 4-5.5 mg/g anthocyanins with balanced anthocyanin profile (malvidin 48-54%, delphinidin 12-18%, other anthocyanidins), creating very dark purple-black berry appearance among darkest in commercial cultivars, producing wines with 15-22 OD color intensity (very high for red wines) with purple hue reflecting balanced anthocyanin composition. Pinot Noir moderately pigmented berry skin showing red-hued characteristics= 2.2-3.2 mg/g anthocyanins with higher proportion of delphinidin and cyanidin 3-glucosides (combined 35-48% versus 25-35% in purple-black cultivars), creating ruby-red berry appearance rather than purple-black, producing wines with 8-13 OD color intensity (moderate for red wines) with characteristic red hue important for Pinot Noir style. Chardonnay white cultivar pigmented berry skin showing non-anthocyanin pigmentation= lacking anthocyanins (<0.02 mg/g) but containing 0.4-0.7 mg/g carotenoids and 1.1-1.8 mg/g flavonols creating golden-yellow berry appearance, demonstrating pigmented berry skin category includes white cultivars with non-anthocyanin pigments determining appearance. Merlot pigmented berry skin light exposure effects= sun-exposed clusters showing 3.8-4.8 mg/g anthocyanins, deep purple-black color, producing wines 14-17 OD color intensity cluster interior shaded berries showing 2.4-3.2 mg/g anthocyanins (33-40% reduction), lighter purple color, producing wines 9-12 OD intensity, demonstrating sunlight critical for full pigmented berry skin development with shading substantially reducing anthocyanin accumulation and color intensity affecting wine color potential and requiring canopy management for optimal pigmentation.
A pigmented berry skin is a berry skin containing visible pigments creating colored berry appearance, primarily through anthocyanin accumulation in hypodermal cells of red/black cultivars (creating red, purple, or black berry color) or through carotenoid and flavonol accumulation in white cultivars (creating golden-yellow color), representing berry skins that determine berry appearance and provide color potential for wine production, distinguished from non-pigmented or minimally-pigmented green berry skins during early development or in rare green-berried cultivars. Pigmented berry skin in red/black cultivars is characterized by= (1) anthocyanin accumulation in hypodermal cell vacuoles beginning at veraison (BBCH 81) and continuing through ripening, reaching mature levels typically 1.5-6+ mg/g fresh weight depending on cultivar (Pinot Noir 2.2-3.5 mg/g moderate, Syrah 3.5-5.5 mg/g high, Tannat 4.5-6.5+ mg/g very high) (2) anthocyanin composition including five major anthocyanidin-3-glucosides (delphinidin, cyanidin, petunidin, peonidin, malvidin creating color spectrum from red through purple to blue-black depending on proportions), acylated derivatives (acetyl and p-coumaroyl esters affecting color stability), and occasional di-glucosides in some species (3) color hue determined by anthocyanin composition with higher malvidin proportion (50-65% of total) creating purple-black color typical of Cabernet Sauvignon, Syrah, while higher delphinidin/cyanidin proportion (30-45%) creating redder hue typical of Pinot Noir (4) color intensity determined by total anthocyanin concentration with deeply pigmented skins showing 4-6+ mg/g supporting deep purple-black appearance and high wine color intensity (5) hypodermal localization with anthocyanins concentrated in 1-3 sub-epidermal cell layers creating 20-60 μm thick pigmented zone containing >90% of berry anthocyanins. Pigmented berry skin in white cultivars is characterized by= (1) absence or near-absence of anthocyanins (32°C inhibiting synthesis water deficit potentially enhancing anthocyanin concentration through berry size reduction creating concentration effects canopy management (leaf removal, cluster thinning) affecting berry light exposure and temperature influencing pigmentation intensity.
Pigmented_Berry_Skin represents skins containing visible pigments (anthocyanins in red/black cultivars, carotenoids/flavonols in white cultivars) and is distinguished from unpigmented green skins during early development. Pigmented berry skin determines berry appearance and wine color potential. Understanding pigmented berry skin development and composition is essential for managing color development through viticultural practices (canopy management, irrigation, harvest timing), for assessing color maturity, and for optimizing wine color extraction through appropriate maceration protocols. The pigmented berry skin concept encompasses both anthocyanin-based (red/black) and non-anthocyanin (white/yellow) pigmentation systems.
Farbstoff-Beerenhaut
anthocyanin-containing skin
colored skin
gefärbte Traubenhaut
pigmentierte Beerenhaut
Castellarin, S.D. et al. (2007). Transcriptional regulation of anthocyanin biosynthesis. Plant Physiology 143=1347-1363 Downey, M.O. et al. (2006). Cultural practice and environmental impacts on flavonoid composition. American Journal of Enology and Viticulture 57=64-75 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 181-185.
Pigmented Berry Skin
Quebranta bears a Pisco_Production_Scion_Role as Peru's most important non-aromatic variety (over 60% of Pisco production), a pink-skinned mutation of Listán Prieto uniquely adapted to Peru's coastal desert valleys, producing Pisco Puro Quebranta with clean, neutral character and subtle dried fruit notes—the benchmark for non-aromatic Peruvian Pisco. Italia (Muscat of Alexandria) serves in a Pisco_Production_Scion_Role for aromatic Peruvian Pisco, with intense terpene content (linalool, geraniol exceeding 5 mg/L in spirit) creating highly perfumed spirits for Pisco Puro Italia and contributing aromatic intensity to Pisco Acholado blends. Moscatel de Alejandría bears a Pisco_Production_Scion_Role as Chile's dominant variety (over 80% of production), producing aromatic spirits with floral-citrus character that define the Chilean Pisco style, particularly in the Elqui and Limarí valleys.
A distillation scion role that inheres in a grafted scion cultivar by virtue of its authorization and suitability for Pisco production in Peru or Chile, where fresh grape must or partially fermented wine is distilled into the clear, unaged grape brandy central to both nations' cultural heritage. Pisco production scions are strictly defined by each country's denomination of origin, with Peru permitting eight specific varieties (four aromatic—Muscat, Italia, Albilla, Torontel four non-aromatic—Quebranta, Negra Criolla, Mollar, Uvina) while Chile permits broader V. vinifera production dominated by Muscat varieties (Moscatel de Alejandría, Moscatel Rosada, Pedro Ximénez, Torontel). Peruvian Pisco classification distinguishes Pisco Puro (single variety), Pisco Acholado (blended varieties), and Pisco Mosto Verde (distilled from partially fermented must), while Chilean Pisco is classified by alcohol content and aging. The aromatic Muscat varieties produce intensely perfumed spirits, while non-aromatic varieties yield cleaner, more neutral Piscos—both traditions emphasizing the unaged, transparent character that distinguishes Pisco from oak-aged brandies.
Pisco_Production_Scion_Role captures the geographically protected spirit traditions of Peru and Chile, where national identity and cultural heritage intertwine with varietal regulations and production methods. The Peru-Chile dispute over Pisco denomination and authenticity reflects distinct traditions—Peruvian Pisco emphasizing pot-still distillation, single distillation to proof, and strict varietal classification Chilean Pisco permitting column distillation and water addition to adjust proof. The aromatic/non-aromatic distinction is particularly important for Peruvian Pisco, where Muscat-family varieties (Italia, Torontel) create intensely perfumed spirits contrasting with the neutral Quebranta style. Climate change threatens both traditions as Muscat varieties' aromatic compound accumulation is temperature-sensitive, with warming potentially diminishing the terpene concentrations that define aromatic Pisco character.
Chilean spirit grape
Peruvian spirit grape
Pisco grape
Pisco variety
Pisco-Rebsorte
Pisco-Traube
peruanische Brenntraube
Consejo Regulador de la Denominación de Origen Pisco (Peru) (2020) Technical Regulations SAG Chile (2020) Denominación de Origen Pisco Regulations Huertas L (2004) Historia de la producción de vinos y piscos en el Perú, Revista Universum Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane pp.: 809-811, 875-877
Pisco Production Scion Role
Chardonnay hermaphrodite flowers at BBCH 65 (full bloom) exhibit fully functional pistils 2-2.3 mm total length comprising ovary 0.8-1 mm diameter containing 4 ovules (2 per locule), style 0.5-0.7 mm, and prominent wet stigma 0.4-0.5 mm diameter with abundant papillae and glistening exudate, positioned 0.8-1.3 mm below the elevated erect anthers creating optimal geometry for self-pollination, achieving 48-56% fruit set under favorable bloom conditions with successful fertilization of 1.9-2.2 ovules per fruit-set flower producing berries with 2-4 seeds. Gewürztraminer functionally female flowers show enhanced pistils 2.4-2.7 mm with particularly elaborate wet stigmas and extended receptivity persisting 6-8 days, positioned prominently above the reflex stamens, requiring cross-pollination from hermaphrodite pollinizers and achieving 28-38% fruit set. Wild V. vinifera ssp. sylvestris pistillate female flowers exhibit robust pistils 2.3-2.6 mm with well-developed ovaries containing 3-4 ovules, prominent stigmas with extended receptivity, positioned well above vestigial stamens (0.3-0.8 mm), requiring cross-pollination from staminate males and achieving 30-42% fruit set when males are within 20-50 m. Cabernet Franc pistils show characteristic bicarpellate structure visible in transverse ovary sections as two distinct locules separated by central partition, with ovules attached to partition surfaces in axile placentation pattern.
A pistil is the female reproductive organ of the Vitis flower comprising an ovary (containing ovules), a style (connecting ovary to stigma), and a stigma (receptive surface for pollen), typically formed from two fused carpels creating a bicarpellate syncarpous gynoecium, functioning to receive pollen, support pollen germination and pollen tube growth, facilitate fertilization of ovules, and develop into the berry following successful fertilization. The pistil structure consists of= (1) an ovary at the base typically 0.6-1.2 mm in diameter and 0.5-1 mm in height, superior in position (positioned above the attachment points of other floral organs), containing two locules (chambers) separated by a central partition with typically 2-4 ovules (commonly 2 ovules per locule, occasionally 1 or 2 per locule), with ovules attached to axile placentation on the central partition (2) a style extending from the ovary apex, typically 0.3-0.8 mm in length and 0.2-0.4 mm in diameter, containing transmitting tract tissue with specialized extracellular matrix supporting pollen tube growth from stigma to ovules (3) a stigma at the style apex, typically 0.3-0.6 mm in diameter, bearing multicellular papillae (specialized epidermal projections) and producing wet stigmatic exudate containing sugars, proteins, and lipids that facilitate pollen grain hydration, adhesion, and germination. Pistil development proceeds through distinct phases= (1) primordium initiation during early flower bud development (BBCH 51-53) as a central dome arising on the receptacle surrounded by developing stamen and petal primordia (2) carpel differentiation and fusion (BBCH 53-55) creating the bicarpellate structure with two locules evident in cross-section (3) ovule initiation and megasporogenesis (BBCH 55-57) producing embryo sacs with egg cells ready for fertilization (4) style and stigma elaboration (BBCH 57-59) with papillae development and exudate-producing cell differentiation (5) stigma receptivity achievement (BBCH 59-65) when stigmatic exudate production activates and pollen germination capability is established, typically coinciding with or slightly preceding calyptra shedding (6) post-pollination pollen tube growth (BBCH 61-69) with pollen tubes traversing the style requiring 48-96 hours to reach ovules at 20-25°C (7) post-fertilization ovary expansion (BBCH 71-75) transforming the pistil into a developing berry. Pistil morphology and function vary among Vitis flower types= hermaphrodite flowers exhibit fully functional pistils 1.8-2.5 mm total length with 3-4 viable ovules, receptive wet stigmas, and functional styles supporting pollen tube growth, achieving 40-60% fruit set functionally female flowers show normal or enhanced pistil development (2-2.7 mm) with extended stigma receptivity (5-8 days versus 3-6 days in hermaphrodites) but requiring cross-pollination from hermaphrodite or staminate sources, achieving 25-40% fruit set functionally male flowers exhibit abbreviated pistils (1-1.8 mm, 50-80% of hermaphrodite length) with non-receptive or rudimentary stigmas and sterile or aborted ovules, achieving <5% fruit set pistillate flowers show enhanced pistil development (2.2-2.8 mm) similar to functionally female types but in context of vestigial or rudimentary stamens, requiring cross-pollination and achieving 25-45% fruit set staminate flowers exhibit vestigial, rudimentary, or short pistils (0.2-1.5 mm) with non-functional reproductive tissues, achieving <1% fruit set. The bicarpellate structure (derived from two fused carpels) is evident in ovary anatomy (two locules separated by partition) and in stigma morphology (often showing bilobed or two-part appearance), representing the ancestral Vitaceae gynoecium organization. Pistil number per flower is normally one (monocarpous at the flower level though bicarpellate at the carpel level), with rare abnormalities producing flowers with two separate pistils or with fused multi-carpellate pistils showing more than two locules.
Pistil represents the complete female reproductive organ and must be distinguished from its component parts (ovary, style, stigma) and from individual carpels (the pistil comprises two fused carpels creating a compound bicarpellate structure). The pistil is the fundamental unit determining fruit set potential and seed number, with pistil functionality (stigma receptivity, style transmitting capacity, ovule viability) critically affecting reproductive success. Pistil morphology, particularly total length and stigma development, varies dramatically among flower types and determines sexual function categories (hermaphrodite versus functionally female versus functionally male versus pistillate versus staminate). Sub-classification addresses the compound bicarpellate nature and structural variations. Understanding pistil development, anatomy, and function is essential for viticulture applications including fruit set optimization, seedlessness breeding, and diagnosis of reproductive failures.
Fruchtblatt
Stempel
female organ
gynoecium
weibliches Blütenorgan
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 99-108 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 92-106 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 185-198 Lebon, G. et al. (2005). Sugars and flowering in the grapevine. Journal of Experimental Botany 56=2565-2578.
Pistil
The pistil description of Merlot documents a bicarpellate ovary (two fused carpels forming a bilocular structure) containing 2 ovules per locule (4 ovules total), a short conical style of 0.3-0.5 mm length, and a capitate to slightly bifid stigma with papillate receptive surface that secretes a thin exudate film during the 2-3 day receptive period following calyptra abscission — successful pollination and fertilization of all 4 ovules produces a four-seeded berry of maximum size (1.8-2.2 g for Merlot), while partial fertilization (2-3 seeds) produces proportionally smaller berries, and complete fertilization failure produces either no berry (abscission of the unfertilized flower) or a small seedless shot berry (millerandage) through parthenocarpic development. The typical Merlot berry at harvest contains 1.5-2.5 seeds, indicating that approximately 40-60% of available ovules are successfully fertilized under normal conditions.
A flower description component that records the morphological characteristics of the pistil — the female reproductive organ consisting of the ovary, style, and stigma — documenting ovary shape and size, the number of ovules per locule (typically two, yielding a maximum of four seeds per berry), style length, stigma form (typically bifid or capitate), and stigmatic surface receptivity, with pistil morphology determining the fruit set potential, seed number, and berry development capacity of each flower.
Pistil_Description documents the female reproductive structure that directly determines berry formation — the ovary develops into the berry pericarp (skin, pulp, and vascular tissue), while the ovules develop into seeds following fertilization, and the relationship between seed development and berry growth is mediated by auxin, gibberellin, and cytokinin hormones produced by the developing seeds that drive cell division and expansion in the surrounding pericarp tissue. The number of seeds per berry is therefore a primary determinant of berry size — berries with four fully developed seeds are typically 30-50% larger than berries with one seed from the same cluster, because each developing seed produces growth hormones that stimulate surrounding tissue development: this seed-size relationship explains the phenomenon of millerandage (hen-and-chicken), where incomplete pollination within a cluster produces a mixture of large seeded berries and small seedless shot berries, creating the berry size heterogeneity that complicates harvest timing and processing. The pistil also provides the physical context for seedlessness — the commercially important seedless table grape cultivars (Sultanina, Flame Seedless, Crimson Seedless) develop normal pistils and undergo pollination and fertilization, but seed development is arrested at an early stage (stenospermocarpy) due to the genetic action of the seedlessness locus on chromosome 18, producing vestigial seed traces (rudimentary soft seed remnants) within otherwise normally developed berries.
Fruchtblatt-Morphologie
Gynoeceum-Charakterisierung
Stempel-Beschreibung
female organ characterization
gynoecium description
pistil morphology record
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Carmona MJ et al. (2008) A molecular genetic perspective of reproductive development in grapevine, Journal of Experimental Botany 59: 2579-2596 Doligez A et al. (2002) An integrated SSR map of grapevine based on five mapping populations, Theoretical and Applied Genetics 105: 161-172
Pistil Description
Wild Vitis vinifera ssp. sylvestris female plants in riparian forests of Mediterranean Europe produce exclusively pistillate flowers at BBCH 65 with well-developed pistils (2.2-2.6 mm length, ovary 1.2-1.5 mm diameter containing 3-4 ovules, prominent wet stigma with elaborate papillae), stamens completely absent or reduced to minute vestiges <0.4 mm, requiring cross-pollination from nearby staminate male plants through wind-borne pollen dispersal, achieving 30-42% fruit set when male plants are within 20-50 m but only 8-15% fruit set when nearest males exceed 100 m distance, and producing clusters of 15-45 berries typical of wild grapevines. Occasional pistillate reversion shoots appear in cultivated Chardonnay vineyards as somatic mutations, producing flowers with normal pistil development but vestigial or absent stamens, setting fruit only through cross-pollination from adjacent hermaphrodite shoots and identified by reduced fruit set (25-35% versus 48-55% on hermaphrodite shoots) and cluster morphology differences. Breeding populations from wild dioecious × cultivated hermaphrodite crosses segregate approximately 25-50% pistillate individuals (depending on parental genotypes) requiring co-planting with hermaphrodite or staminate pollinizers for fruit production in breeding nurseries.
A pistillate flower is an imperfect (unisexual) flower possessing a functional pistil with receptive stigma and viable ovules capable of fertilization and fruit set but completely lacking functional stamens or possessing only vestigial, rudimentary, or strongly reflex stamens incapable of effective self-pollination, representing true morphological female flowers that occur in dioecious wild Vitis vinifera subspecies sylvestris, occasionally in cultivated Vitis vinifera as genetic reversions, and characteristically in various wild North American Vitis species, requiring cross-pollination from staminate or hermaphrodite flowers for successful fruit production. Pistillate flowers contrast fundamentally with perfect flowers (which possess both functional stamens and pistil) and with functionally female flowers (which possess stamens morphologically but with strongly reflex position or non-viable pollen that nonetheless allows some minimal self-pollination in exceptional circumstances), as pistillate flowers exhibit complete or near-complete absence of functional male structures at the morphological or functional level, creating absolute dependency on cross-pollination. The pistillate flower typically measures 2-3.5 mm in diameter at anthesis and comprises= (1) a minute calyx forming a shallow receptacle cup (2) five petals fused into a calyptra that detaches at anthesis (3) stamens that are either completely absent (rare), extremely reduced to minute vestigial remnants <0.5 mm long lacking recognizable anthers and pollen (common), or rudimentary with recognizable but severely abbreviated stamen morphology 0.5-1.5 mm long with strongly reflex position and minimal or absent pollen production (4) a well-developed, functional pistil 1.8-2.8 mm long comprising ovary with 2-4 viable ovules, functional style with transmitting tract tissue, and receptive wet stigma with characteristic papillae and secretory exudate. The pistil in pistillate flowers typically shows enhanced development compared to hermaphrodite flowers, with 10-25% greater pistil length, 15-30% larger ovary diameter, enhanced stigmatic surface area through more elaborate papillae development, and increased ovule number per ovary (often 3-4 ovules versus 2-3 in hermaphrodite flowers), representing apparent resource reallocation from absent or minimal male structures to enhanced female function. Stigma receptivity in pistillate flowers is typically prolonged (4-8 days post-anthesis versus 3-6 days in hermaphrodite flowers) and stigmatic exudate production is enhanced, adaptations that increase probability of successful cross-pollination by extending the temporal window for pollen capture from wind-dispersed or insect-borne pollen from distant staminate or hermaphrodite sources. Pistillate flowers achieve fruit set only through cross-pollination, with fruit set percentages typically ranging from 25-45% under good pollination conditions (adequate proximity to pollen sources, favorable weather for pollen dispersal) but declining to 5-15% under poor pollination conditions (isolated female plants distant from pollen sources, wet or humid weather preventing pollen dispersal). Pistillate flowers characterize the female phase of dioecious wild V. vinifera ssp. sylvestris populations throughout the species' native range, where approximately 50% of individuals are pistillate (female) plants bearing exclusively pistillate flowers, with the remaining 50% being staminate (male) plants bearing staminate flowers. The dioecious reproductive system with pistillate and staminate flowers represents the ancestral condition in Vitis, with the hermaphrodite condition of cultivated V. vinifera ssp. vinifera arising through mutations that restored stamen function in pistillate backgrounds or enhanced pistil function in staminate backgrounds, with these hermaphrodite mutations strongly selected during domestication to enable self-fertility and reliable fruit production without requiring separate male and female plants. Pistillate flowers occasionally appear in cultivated vineyards as somatic reversions where hermaphrodite cultivar shoots undergo bud sport mutations converting to pistillate expression, or as seedlings from uncontrolled pollination exhibiting genetic segregation for sex determination. The genetic control involves sex-determining loci with female-promoting alleles that suppress stamen development, with pistillate individuals typically carrying female-determining alleles in homozygous state while hermaphrodites carry recessive alleles permitting stamen development or dominant hermaphrodite-promoting alleles.
Pistillate_Flower represents true morphological female flowers and must be carefully distinguished from Functionally_Female_Flower (which possesses stamens morphologically with strong reflexing or reduced pollen but retains some minimal self-pollination capability under exceptional conditions). The distinction is critical= pistillate flowers lack functional stamens structurally or have such extreme stamen reduction that self-pollination is absolutely impossible (imperfect flowers), while functionally female flowers possess reflex but morphologically present stamens that can occasionally contribute minimal pollen (perfect flowers with male organ dysfunction). Pistillate flowers are rare in cultivated V. vinifera ssp. vinifera but characteristic of wild V. vinifera ssp. sylvestris female plants and represent the ancestral dioecious female phase from which hermaphrodite cultivated grapes evolved. Sub-classification into Female_Only (no stamen trace) versus Reduced_Stamen (vestigial or rudimentary stamens present) reflects variation in degree of stamen suppression, paralleling the classification of staminate flowers.
Pistillatblüte
carpellate flower
female flower
weibliche Blüte
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 100-106 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Antcliff, A.J. (1980). Inheritance of sex in Vitis. Annales de l'Amélioration des Plantes 30=113-122 Battilana, J. et al. (2013). The genetic basis of the conversion of hermaphrodite flowers to male flowers in grapevine. Nature Communications 4=2615 Picq, S. et al. (2014). A small XY chromosomal region explains sex determination in wild dioecious V. vinifera and the reversal to hermaphroditism in domesticated grapevines. BMC Plant Biology 14=229 Badouin, H. et al. (2020). The wild grape genome sequence provides insights into the transition from dioecy to hermaphroditism during grape domestication. Genome Biology 21=223.
Pistillate Flower
Regent (released 1996, JKI Geilweilerhof) bears a PIWI_Cultivar_Role as one of the most commercially successful resistant varieties, with approximately 2,000 hectares planted in Germany producing red wines of recognized quality while requiring 70-80% fewer fungicide applications than susceptible varieties. Solaris (released 2001, Freiburg) serves in a PIWI_Cultivar_Role as a white variety with exceptional disease resistance and early ripening suited to northern European viticulture, achieving quality recognition in England, Scandinavia, and other challenging climates. Souvignier Gris (released 2008, Freiburg) bears a PIWI_Cultivar_Role as a contemporary white PIWI variety producing wines of recognized quality, with plantings expanding across Europe as sustainability requirements intensify.
A breeding role that inheres in a grapevine variety classified as PIWI (pilzwiderstandsfähig, German for "fungal disease resistant"), carrying genetic resistance to major fungal pathogens (powdery mildew Erysiphe necator and/or downy mildew Plasmopara viticola) introgressed from wild Vitis species into cultivated backgrounds. PIWI varieties enable substantial reduction in fungicide applications (typically 50-80% reduction compared to susceptible V. vinifera), supporting sustainable viticulture and organic production while reducing environmental impact and production costs. PIWI designation indicates functional resistance validated through field evaluation, typically conferred by one or more resistance (R) loci inherited from interspecific hybridization with American or Asian Vitis species (Run1, Ren1-9 for powdery mildew Rpv1-17+ for downy mildew). PIWI varieties have progressed through generational improvement—early PIWIs offered resistance but compromised wine quality, while contemporary third-generation PIWIs approach V. vinifera quality standards. Regulatory recognition of PIWI varieties has expanded substantially since 2000, with EU member states increasingly authorizing PIWI varieties for quality wine production previously restricted to V. vinifera, and international adoption growing as sustainability pressures intensify.
PIWI_Cultivar_Role encompasses the functional category of fungal-resistant varieties developed through interspecific breeding, representing the primary current focus of grapevine variety development worldwide. The German terminology (pilzwiderstandsfähig) reflects the leading role of German breeding programs (JKI Geilweilerhof, Freiburg, Weinsberg, Geisenheim) in developing and promoting resistant varieties, though French, Italian, Swiss, and international programs contribute substantially. PIWI varieties are subclassified by generation reflecting the iterative improvement from early resistant hybrids (compromised quality) through contemporary varieties approaching V. vinifera standards. The resistance mechanisms in PIWI varieties (R-gene-mediated recognition and response) are qualitatively different from the quantitative field tolerance sometimes observed in V. vinifera clones, enabling dramatic reduction rather than modest improvement in disease susceptibility. PIWI adoption faces both technical challenges (durability of single-gene resistance, residual quality limitations) and market challenges (consumer and regulatory acceptance of non-traditional varieties).
PIWI variety
PIWI-Sorte
disease-resistant variety
fungal-resistant variety
interspecific resistant variety
krankheitsresistente Rebsorte
pilzwiderstandsfähige Rebsorte
resistant cultivar
resistente Sorte
Töpfer R et al. (2011) Molecular breeding in grapes, CRC Press pp.: 160-185 Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330 Pedneault K, Provost C (2016) Fungus resistant grape varieties, Applied Microbiology and Biotechnology 100: 3499-3519 PIWI International (2020) PIWI Variety Descriptions and Cultivation Guidelines
PIWI Cultivar Role
Plant material availability for Chardonnay clone 96 (Dijon) in the US market as of the 2025 ordering season is classified as "widely available" with an estimated 500,000+ bench grafts producible annually across licensed California nurseries, offered on rootstocks 101-14 Mgt, SO4, 3309C, 110R, and 1103P, with standard delivery lead time of 12-18 months from order placement and spring or fall delivery options. Plant material availability for Sankt Laurent clone SL-4 in Austria is documented as "limited availability" with annual production of approximately 15,000 grafted vines, offered exclusively on Kober 5BB and SO4 rootstocks, requiring 18-24 month advance ordering, and subject to allocation priority favoring established customer relationships — reflecting the niche market position of this cultivar despite growing oenological interest.
Plant material availability is a distribution record entity that records the current supply status, order lead times, and production volume of a certified clonal line in the nursery market, including the quantity of grafted vines, rootlings, or cuttings available for delivery in the current and upcoming planting seasons, the estimated waiting period for orders exceeding available stock, the rootstock combinations routinely offered, and the seasonal ordering windows and delivery schedules, providing the practical supply chain information essential for vineyard establishment planning and clone sourcing decisions.
Plant material availability represents the practical bottleneck between clone selection decisions and vineyard establishment reality, as the biological constraints of vegetative propagation limit the rate at which new or newly popular clones can be scaled to commercial supply volumes. Starting from a single certified mother vine, the geometric progression of multiplication (approximately 10-20 cuttings per vine per year for bench graft production) requires 5-8 years to build stock sufficient for large-scale commercial supply, creating persistent scarcity for recently certified clones. Availability classification typically follows a qualitative scale — "widely available" (sufficient supply to meet demand without allocation), "available" (adequate supply with standard lead times), "limited availability" (demand exceeds supply, allocation or extended lead times required), and "not available/special order" (not in routine production, requiring custom propagation from foundation material). Rootstock combination availability adds complexity, as nurseries maintain mother blocks of specific scion-rootstock pairings based on projected demand, and unusual combinations may require 2-3 year custom propagation lead times. Market dynamics significantly influence availability — sudden demand surges driven by varietal popularity trends (e.g., the explosive growth in Grüner Veltliner plantings in the early 2000s) can rapidly deplete nursery stocks of preferred clones, while demand declines can lead nurseries to reduce or discontinue production of slower-selling selections. Accurate availability data requires regular updating from nursery sources, as conditions change seasonally and interannually.
Bestandsverfügbarkeit
Lieferstatus
Pflanzenmaterialverfügbarkeit
Pflanzgutverfügbarkeit
Rebenverfügbarkeit
clone supply status
material availability
plant material supply status
stock availability
vine availability
Nicholas P (2004) Grapevine propagation. In Viticulture Volume 1: Resources, Winetitles, pp.: 213-235 Mannini F (2004) Clonal selection in grapevines. Acta Horticulturae 652: 21-27 Wolpert JA et al. (1994) Field performance of six Chardonnay clones in the Napa Valley. American Journal of Enology and Viticulture 45(4): 393-400
Plant Material Availability
The plant water status target for a premium Syrah vineyard in the Barossa Valley specifies predawn leaf water potential (Ψpd) targets by phenological phase — budburst to flowering at -0.1 to -0.2 MPa (no stress, supporting canopy establishment), flowering to fruit set at -0.2 to -0.3 MPa (minimal stress, protecting fruit set), fruit set to véraison at -0.4 to -0.6 MPa (moderate stress, reducing berry size and vegetative growth), and véraison to harvest at -0.6 to -0.8 MPa (moderate to severe stress, promoting concentration and flavor development) — with midday stem water potential (Ψstem) monitored as a supplementary indicator at targets of -0.8 to -1.0 MPa during the deficit period, and a critical lower limit of -1.4 MPa Ψstem below which irrigation is applied regardless of scheduling protocol to prevent irreversible vine damage from excessive dehydration.
An irrigation protocol description that documents the target values for plant-based water status indicators — principally predawn leaf water potential (Ψpd), midday leaf water potential (Ψleaf), midday stem water potential (Ψstem), and stomatal conductance (gs) — that define the desired vine hydration level for each phenological phase and serve as the ultimate reference for evaluating whether soil-based and atmospheric-demand-based irrigation scheduling is achieving the intended vine water status.
Plant_Water_Status_Target represents the most physiologically direct approach to irrigation management because it measures the actual hydration status of the vine rather than inferring it from soil moisture or atmospheric demand calculations. Predawn leaf water potential (Ψpd), measured before sunrise when soil-plant water equilibrium is established, provides the most reliable estimate of soil water availability in the root zone, while midday stem water potential (Ψstem), measured on bagged non-transpiring leaves at solar noon, integrates soil water availability with atmospheric demand to indicate the actual water stress experienced by the vine during peak transpiration: the relationship between these plant-based indicators and vine physiological responses (stomatal closure, photosynthesis reduction, berry growth cessation, ABA signaling) is better characterized than the relationship between soil moisture and vine response, making plant-based targets the gold standard for precision irrigation management. The Ψpd thresholds of Carbonneau (1998) — no stress above -0.2 MPa, mild stress -0.2 to -0.4 MPa, moderate stress -0.4 to -0.6 MPa, severe stress below -0.6 MPa — provide the widely adopted reference framework for interpreting plant water status measurements in the context of irrigation management decisions.
Blattwasserpotential-Zielwert
Pflanzenwasserstatus-Zielwert
Reben-Hydratations-Spezifikation
leaf water potential target
plant hydration specification
vine water status target
Carbonneau A (1998) Irrigation, vignoble et produit de la vigne, in Tiercelin JR (ed) Traité d'irrigation, Lavoisier Tec et Doc pp: 257-276 Choné X et al. (2001) Stem water potential is a sensitive indicator of grapevine water status, Annals of Botany 87: 477-483 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data, Annals of Botany 105: 661-676 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 400-420
Plant Water Status Target
Planting rights for new vineyard establishment in Austria operate under the EU vine planting authorization system (post-2016 reform) — applicants submit requests to the Bundesministerium für Land- und Forstwirtschaft specifying the proposed area, location, and cultivar(s), with authorizations granted from an annual national allocation pool not exceeding 1% of the total planted vineyard area (approximately 450 ha annually from Austria's approximately 45,000 ha total), priority criteria favoring applicants with existing vineyard holdings, young farmers, and sites within recognized quality appellations, and authorizations valid for 3 years from date of issue with forfeiture if not exercised — the system balancing market access for new entrants against protection of existing producers from uncontrolled supply expansion. Planting rights for replanting existing vineyards in Bordeaux permit growers to replant grubbed vineyard area with a different authorized cultivar within 3 years of grubbing without requiring a new planting authorization — a provision increasingly used to replace climate-vulnerable Merlot with more heat-tolerant varieties (Cabernet Franc, Petit Verdot) within existing vineyard holdings while maintaining total appellation area.
Planting rights is a varietal authorization entity that documents the legal regime governing the permission to establish new vineyards or replant existing vineyards with specific grape cultivars, encompassing the planting authorization system (the mechanism by which permission to plant vines is granted, transferred, or restricted), the applicable quota or limitation framework (area caps, regional allocation pools, conversion rights from grubbed vineyards), the administrative process for obtaining planting permission (application procedures, eligibility criteria, priority rankings), and the financial and temporal conditions attached to the authorization (fees, validity periods, use-it-or-lose-it deadlines), providing the supply-management dimension of viticultural regulation that controls the total vineyard area and its distribution across regions and cultivars.
Planting rights documentation addresses the supply-management dimension of viticultural regulation — the recognition that unrestricted vineyard planting would create chronic overproduction, depress prices, and undermine quality incentives, necessitating administrative control of the total viticultural area and its allocation among regions and producers. The European planting rights regime has undergone a fundamental transformation from the pre-2016 system of transferable planting rights (where the right to plant vineyards could be bought and sold as a marketable asset, creating a secondary rights market that capitalized planting permissions at €5,000-€50,000 per hectare depending on region) to the post-2016 authorization system (Regulation 2015/560, where planting authorizations are administrative permits allocated through an annual application process, non-transferable, and valid for 3 years). This reform addressed the concern that the tradeable rights system created barriers to entry for young farmers and new producers while generating windfall gains for rights holders, but the new system has its own criticisms — annual allocation caps constrain expansion in regions with growing demand, the priority criteria may not optimally allocate vineyard establishment opportunities, and the 1% annual growth limit was set as a political compromise rather than an agronomic or market-based calculation. The replanting right — the automatic authorization to replant an equivalent area after grubbing an existing vineyard — is the most commonly exercised planting right in practice, as it enables vineyard renewal and cultivar conversion without entering the competitive new-planting authorization process. The non-European approach to planting rights diverges fundamentally — most New World wine jurisdictions impose no restrictions on vineyard establishment beyond standard land-use planning permissions, leaving the decision to plant (or remove) vineyards entirely to market forces and individual business judgment, though some regions exercise indirect supply management through water allocation restrictions (particularly relevant in Australia and California) or zoning regulations that limit agricultural conversion.
Auspflanzungsrechte
Pflanzgenehmigung
Pflanzrechte
Pflanzrechtesystem
Rebpflanzrechte
planting authorization
planting permits
planting regime
vine planting rights
vineyard establishment rights
European Parliament (2015) Commission Delegated Regulation (EU) 2015/560 supplementing Regulation (EU) No 1308/2013 as regards the scheme of authorisations for vine plantings Österreichisches Weingesetz (2009) §21-24 (Pflanzrechte) European Parliament (2013) Regulation (EU) No 1308/2013, Articles 62-72 (vine planting authorisation scheme) Meloni G, Swinnen J (2013) The political economy of European wine regulations. Journal of Wine Economics 8(3): 244-284
Planting Rights
Plastic deformation quality assessment of mechanical harvest impact in Chardonnay quantifies the structural damage determining must quality, with optimized harvest parameters limiting plastic deformation to less than 5% of berries (visible crush damage) and producing must with oxidative capacity (dissolved oxygen consumption) below 20 mg/L, while aggressive harvest causing plastic deformation in greater than 15% of berries releases sufficient cellular contents to overwhelm antioxidant capacity (oxidative consumption above 50 mg/L), demonstrating how plastic deformation from mechanical damage directly affects must oxidation susceptibility and subsequent wine quality potential.
Plastic_Deformation_Quality represents the physical quality characterizing the permanent shape change of grape tissue that persists after removal of applied stress, reflecting irreversible structural rearrangement including cell rupture, wall fracture, and loss of intercellular adhesion. Plastic deformation indicates tissue damage where the original structure cannot be restored, distinguishing it fundamentally from elastic deformation where recovery occurs. In grape berries, plastic deformation results from stress exceeding the structural capacity of cells to maintain integrity, producing crushed zones where cell walls have failed, turgor is lost, and juice has been released. The threshold for plastic deformation (yield point or elastic limit) represents the damage threshold above which permanent quality loss occurs. This threshold varies with cultivar, ripeness, temperature, and loading rate, generally decreasing during ripening as tissue softening reduces structural strength. The extent of plastic deformation at given stress indicates tissue vulnerability to damage, with soft, overripe tissue showing predominantly plastic response while firm tissue maintains elastic behavior to higher stress levels. Plastic deformation manifests visually as bruises, crush damage, and loss of berry shape that degrades appearance and initiates quality deterioration through juice exposure, oxidation, and microbial access. Assessment of plastic deformation requires comparison of tissue dimensions before loading and after unloading plus recovery time, with permanent deformation expressed as percentage of original dimension not recovered. The force-deformation curve shows plastic deformation through hysteresis—the area between loading and unloading curves indicating energy dissipated in permanent structural change.
Plastic_Deformation_Quality specifically addresses the permanent, non-recoverable component of deformation representing actual structural damage. The distinction from elastic deformation is fundamental: elastic deformation represents safe, transient stress while plastic deformation indicates damage threshold exceedance. The class connects mechanical assessment to quality deterioration, as plastic deformation initiates the cascade of juice release, oxidation, and spoilage that degrades quality. Hysteresis analysis of loading-unloading cycles provides detailed characterization of the elastic-plastic balance in tissue mechanical response.
Plastische Verformung
irreversible Verformung
irreversible deformation
non-recoverable deformation
permanent deformation
permanente Verformung
plastic strain quality
plastische Dehnungsqualität
Plastic Deformation Quality
The ploidy information compiled for the VIVC database documents the chromosome complement of characterized Vitis vinifera cultivars — the vast majority (>99%) are confirmed diploids (2n = 2x = 38), with a small number of tetraploid variants (2n = 4x = 76) arising through somatic chromosome doubling during vegetative propagation or induced by colchicine treatment in breeding programs, and rare aneuploid individuals (2n = 37 or 39) detected in seedling populations from interspecific crosses. Ploidy determination methods include flow cytometry (the current standard, measuring nuclear DNA content relative to a diploid reference with coefficient of variation below 3%), chromosome counting from root tip squash preparations stained with aceto-carmine or DAPI, and guard cell chloroplast counting (an indirect screening method where diploid Vitis guard cells contain 12-16 chloroplasts versus 20-28 in tetraploids).
A genetic information entity that documents the chromosome complement status of a Vitis vinifera individual, recording the number of complete chromosome sets (ploidy level), any deviations from the standard diploid state (2n = 38), and the method of ploidy determination, providing essential cytogenetic data for cultivar characterization, breeding program management, and the interpretation of phenotypic traits influenced by genome dosage effects.
Ploidy_Information documents the fundamental cytogenetic parameter that determines genome dosage and influences cell size, organ size, fertility, and gene expression patterns in grapevine. The genus Vitis is uniformly based on x = 19 chromosomes, with all wild and cultivated species diploid (2n = 38) under natural conditions — this chromosomal stability contrasts with many other fruit crop genera (Fragaria, Prunus, Malus, Rubus) where polyploidy is common or predominant: the absence of natural polyploidy in Vitis may reflect the functional constraints of the paleohexaploid genome (the ancestral genome triplication approximately 117 million years ago already providing gene redundancy equivalent to a hexaploid) or the reproductive barriers imposed by the dioecious-to-hermaphroditic transition in domesticated grapevine. Deviations from the diploid state — tetraploidy, aneuploidy, and chimerism — arise through somatic mutation during vegetative propagation and have specific phenotypic consequences (larger berries and organs in tetraploids, reduced fertility in aneuploids, sectoral variation in chimeras) that are documented in the subclasses.
Chromosomensatz-Dokumentation
Ploidie-Information
chromosome number record
cytogenetic status data
ploidy level documentation
zytogenetische Statusdaten
This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Jaillon O et al. (2007) The grapevine genome sequence suggests ancestral hexaploidization in major angiosperm phyla, Nature 449: 463-467 Ollat N et al. (2011) Grape berry development, in Gerós H et al. (eds) The Biochemistry of the Grape Berry, Bentham Science pp: 1-29
Ploidy Information
Comparison of canopy management treatments in Oregon Pinot Noir evaluated point quadrat index quality at véraison plus 30 days, finding that standard shoot positioning alone produced leaf layer number of 2.8 with 12% gaps and 78% interior clusters, while additional basal leaf removal reduced leaf layer number to 1.4 with 35% gaps and 31% interior clusters. The improved point quadrat index quality correlated with 40% reduction in Botrytis incidence, 2.1°Brix higher fruit sugar at equivalent harvest date, and enhanced red fruit character with reduced vegetal notes in sensory evaluation of resulting wines.
Point quadrat index quality is a vigor balance quality that inheres in a grapevine canopy system and represents the characterization of canopy density and architecture through systematic transect sampling using a thin probe inserted horizontally through the fruiting zone to quantify leaf layer number, gap frequency, and fruit exposure as indicators of vine balance and canopy management effectiveness. This quality provides standardized, reproducible assessment of the light and air penetration characteristics that fundamentally determine fruit microclimate and disease pressure within the canopy interior. The ontological basis of point quadrat index quality rests on the principle that canopy density directly reflects the balance between vegetative growth and canopy management intensity, with excessive leaf layering indicating vigor imbalance favoring vegetative dominance while sparse canopies may signal insufficient vigor or excessive leaf removal. The point quadrat methodology, adapted from ecological vegetation sampling techniques, involves inserting a rigid probe at regular intervals along and perpendicular to the row, recording each contact with leaves, clusters, or gaps, and computing derived indices including leaf layer number (mean contacts per insertion), percentage interior leaves (contacts beyond the first), percentage interior clusters (fruit with leaf obstruction), and percentage gaps (insertions with no contacts). Optimal point quadrat index quality for wine grape production generally targets leaf layer numbers between 1.0-1.5, gap frequencies of 20-40%, and interior cluster percentages below 50%, though these targets vary with cultivar, climate, and production objectives. The relationship between point quadrat index quality and fruit composition reflects the fundamental importance of light exposure for phenolic synthesis, flavor precursor development, and disease suppression, with dense canopies promoting herbaceous character, reduced color development, elevated disease pressure, and delayed ripening. Point quadrat assessment enables objective comparison of canopy management practices and cultivar-training system combinations, supporting evidence-based decisions regarding shoot positioning, leaf removal intensity, and hedging timing. The temporal dynamics of point quadrat index quality follow seasonal canopy development, with measurements most meaningful when conducted post-véraison after canopy architecture stabilization but before significant leaf senescence, typically providing 4-6 week windows for standardized assessment.
Point_Quadrat_Index_Quality is_a Vigor_Balance_Quality, representing the specific methodology of transect-based canopy density assessment. This class captures the standardized point quadrat approach developed by Smart and Robinson as objective measurement of canopy architecture reflecting vigor balance outcomes. Distinguished from other canopy assessment approaches by the specific probe-insertion methodology and derived metrics including leaf layer number and gap percentage. Functionally related to Canopy_Density_Quality and Fruit_Exposure_Quality.
PQA index
Punktquadrat-Analyse
Punktquadrat-Index
Transekt-Kronenmessung
canopy point quadrat
leaf layer number assessment
point quadrat analysis
transect canopy sampling
Point Quadrat Index Quality
A pointed apical lobe on a mature leaf at node 6 of Vitis vinifera cv. Cabernet Franc in Loire Valley, France, exhibiting characteristic acute apex with point angle of approximately 42 degrees creating sharp pointed appearance, acuminate tip with extended taper over the final 1.2 cm creating distinctive sharp point, lobe margins in the apical region showing slightly concave curvature converging smoothly toward the point, median vein extending prominently through lobe center to within 0.2 cm of apex point clearly visible as raised ridge on abaxial surface, point symmetry with left and right margins converging at similar angles creating balanced apex geometry, overall lobe measuring 5.8 cm length and 4.0 cm width, separated from upper lateral lobes by sinuses of 2.8 cm depth, sharp pointed apex configuration serving as important diagnostic feature distinguishing Cabernet Franc from Cabernet Sauvignon (less acute point) and Merlot (more obtuse point), point frequently showing slight marginal teeth creating serrated apex appearance, measured at BBCH stage 75 (berries pea-sized) in late June, representing typical pointed apical lobe morphology characteristic of this cultivar.
An apical lobe exhibiting acute apex configuration with sharply tapering point in Vitis vinifera, characterized by apex angle typically less than 50-60 degrees creating sharp rather than broad or rounded point, often acuminate with extended tapering tip, straight or slightly concave lobe margins in the apical region converging toward the point, median vein extending prominently to or very near the apex point often visible externally as raised ridge, cultivar-specific point characteristics including degree of acuteness, presence of acumination, and symmetry serving as diagnostic features, occurrence in many major cultivars including Cabernet Sauvignon, Cabernet Franc, Syrah, and Merlot though with varying degrees of point acuteness, and functional implications potentially including reduced mechanical stress at apex through streamlined geometry facilitating wind and rain shedding though experimental support is limited. Pointed apical lobes represent a common apex configuration contrasting with obtuse or rounded apical lobes, exhibiting genetic control creating cultivar-specific point characteristics, showing relative environmental stability making apex configuration reliable diagnostic feature, and contributing to overall leaf outline shape affecting visual identification.
Pointed apical lobes are distinguished from obtuse or rounded apical lobes by acute apex angle typically <50-60 degrees, with exact threshold somewhat arbitrary. Acumination involving extended taper enhances pointed appearance and diagnostic utility, with some cultivars showing pronounced acuminate tips (Cabernet Franc, some Syrah clones) while others show acute points without strong acumination. Lobe margin curvature in the apical region affects point character, with straight or concave margins creating sharper points compared to convex margins producing broader points. Point symmetry varies from perfect bilateral symmetry to asymmetric configurations with one margin extending further creating lopsided points. Genetic control is strong making pointed versus obtuse apex relatively stable diagnostic character, though some environmental modification occurs with extreme vigor occasionally producing broader points. Functional implications remain largely speculative, with proposed advantages including streamlined geometry for wind and precipitation shedding, reduced mechanical stress concentration at apex, and potentially enhanced apical meristem protection during development lacking strong experimental support. Ampelographic utility is substantial, with apex point characteristics serving as secondary or tertiary diagnostic features combining with other lobe and sinus characteristics for cultivar identification. The degree of acuteness shows continuous variation creating subcategories from very acute (<40 degrees) to moderately acute (40-60 degrees) useful for finer discrimination.
acuminate apex lobe
acute apical lobe
akuter Endlappen
sharp-pointed terminal lobe
spitzer Apikallappen
zugespitzter Spitzenlappen
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 136-182 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 103-136.
Pointed Apical Lobe
Pinot Noir in Burgundy shows pollen adhesion during early-mid June Year N+1 at BBCH 60-65, with firm molecular binding developing within 20-45 minutes after contact through stigmatic exudate lipid transfer proteins and polysaccharide matrix, creating stable attachment resistant to moderate wind and environmental disturbances, supporting 14-32 securely attached pollen grains per stigma, under continental conditions of 20-26°C temperature, 56-70% relative humidity, typical of this early-to-mid season cultivar with adequate boron nutrition supporting normal exudate production and adhesion enabling subsequent germination
Pollen adhesion represents the firm stable attachment phase when strengthened molecular binding between pollen grain and stigmatic exudate creates secure long-term association resistant to environmental disturbances, establishing optimal conditions for pollen recognition, hydration, and subsequent germination supporting successful pollination. This critical stable attachment phase occurs during pollination in early-mid June in temperate regions Year N+1, developing over 15-60 minutes following adhesion initiation as firm binding establishes, corresponding to BBCH 60-65 flowering stages. Adhesion characteristics show firm molecular binding between pollen coat proteins and stigmatic exudate components creating stable attachment, physical adhesion through viscous exudate matrix resisting detachment forces, mechanical retention by papillate cell architecture preventing grain displacement, and overall secure association enabling continued pollen-stigma interaction without risk of grain loss. Molecular binding mechanisms involve protein-protein interactions between pollen coat adhesins and stigma exudate receptors creating specific high-affinity bonds, protein-lipid interactions with stigma lipid transfer proteins binding pollen coat lipids, polysaccharide interactions between pollen wall polymers and exudate carbohydrates, and integrated molecular network creating robust multi-point attachment. Lipid transfer protein function in adhesion shows critical role with proteins abundant in grapevine stigmatic exudate binding pollen coat lipids creating strong molecular bridges, mediating species recognition through lipid-binding specificity, and contributing substantially to adhesion strength through multiple binding sites per grain. Exudate matrix role shows physical adhesive function with proteinaceous-polysaccharide network embedding pollen grain creating mechanical retention, viscous properties resisting shear forces preventing grain sliding or rolling, and hydrated gel-like consistency maintaining stable grain positioning throughout receptivity period. Pollen coat substances participating in adhesion include proteins from tapetal origin on grain surface providing binding sites for exudate interactions, lipids including tryphine components mediating lipid transfer protein binding, polysaccharides in pollen wall participating in carbohydrate interactions, and integrated coat composition determining adhesion capacity and recognition properties. Adhesion strength at maturity shows substantial resistance to detachment with established adhesion requiring significant mechanical force for grain removal, resistance to environmental disturbances including wind, rain, or physical contact, and stable attachment persisting throughout pollen hydration, germination, and initial tube growth phases. Mechanical adhesion assessment through controlled detachment experiments shows mature adhesion withstanding forces 100-500 times grain weight, demonstrating robust attachment supporting reproductive success, and confirming that normal environmental conditions insufficient to dislodge properly adhered pollen. Pollen grain hydration accompanying adhesion shows continued water uptake from stigmatic exudate over 15-90 minutes following adhesion, grain expanding 10-20% volumetrically as cells fully rehydrate, and metabolic activation proceeding as hydration restores physiological function enabling germination capacity. Recognition processes during adhesion involve molecular signaling between pollen and stigma assessing compatibility, self-compatible grapevine showing acceptance of self-pollen enabling continued support, and recognition triggering downstream responses including enhanced exudate secretion and metabolic support for germination. Stigma papillate cell response to adhesion shows maintained exudate secretion supporting attached pollen, possible localized metabolic activation in cells bearing pollen preparing to support germination and tube growth, and continued provision of water and possibly nutrients enabling pollen function. Multiple pollen adhesion on single stigma shows typical pattern with receptive stigma supporting 10-50 firmly attached grains though only 1-2 ultimately germinating successfully, excess adhesion providing reproductive assurance compensating for possible germination failures, and demonstrating pollen surplus characteristic of wind-pollinated mode supplementing self-pollination. Adhesion persistence shows stable attachment maintained throughout pollination and early germination periods spanning hours to days, attachment remaining firm during pollen tube emergence and initial penetration of stigma surface, and long-term association as germinated pollen develops tube growing through style tissues. Environmental effects on adhesion stability include temperature affecting exudate properties and molecular binding stability with optimal 22-28°C maintaining robust adhesion, humidity influencing exudate water content and consistency with moderate 50-65% optimal, and rain potentially disrupting adhesion through dilution or physical washing requiring substantial water volume to overcome established adhesion. Temperature stability of adhesion shows maintained attachment across typical flowering temperature range 15-32°C, potential minor weakening at temperature extremes though generally retaining adequate binding, and thermal stability of molecular interactions supporting adhesion function under varying conditions. Humidity effects on established adhesion show relative stability across normal range 40-75% with adhesion persisting despite exudate water content changes, high humidity potentially causing slight adhesion weakening through excessive dilution though established binding maintains attachment, and low humidity potentially strengthening adhesion through exudate concentration though may affect pollen hydration. Rain resistance of established adhesion shows substantial water required to disrupt firm binding, light rain or brief showers potentially insufficient to remove adhered pollen though may affect subsequent processes, and heavy sustained rain potentially washing attached pollen particularly if occurring shortly after adhesion before maximal binding develops. Boron nutrition importance for adhesion emphasizes adequate boron essential for normal stigmatic exudate production supporting adhesion capacity, deficiency reducing exudate quantity and possibly altering composition impairing adhesion strength, and demonstrating critical role of boron in supporting pollination success through effects on adhesion. Failed adhesion causing pollination failure results from inadequate exudate production preventing binding development, abnormal pollen coat composition compromising molecular interactions, environmental interference particularly rain diluting or removing exudate, or developmental abnormalities affecting stigma or pollen function. The process shows two distinguishable aspects progressing from stigmatic exudate binding creating molecular attachment, through pollen grain attachment establishing stable long-term association. Duration represents developing over 15-60 minutes as firm adhesion establishes, with stable attachment persisting hours to days supporting subsequent processes. Cultivar variation in adhesion may include exudate composition affecting binding mechanisms and strength, exudate production quantity affecting adhesion capacity, and pollen coat characteristics affecting molecular interaction quality. Regional timing shows pollen adhesion occurring during flowering period across all climates following pollen capture events, developing as stable attachment within minutes to hour after initial contact. Viticultural significance includes representing essential stable attachment enabling subsequent germination and fertilization, requiring adequate stigma exudate production and favorable environmental conditions, demonstrating critical role of boron nutrition in supporting adhesion through exudate effects, and establishing adhesion success as prerequisite for reproductive success and fruit set. Research applications include analyzing molecular mechanisms of adhesion including binding specificities and affinities, investigating exudate composition effects on adhesion properties, evaluating environmental and nutritional factors affecting adhesion stability, quantifying adhesion strength and resistance to disturbance, and assessing relationships between adhesion characteristics and pollination success outcomes.
Pollen_Adhesion is second major Pollination phase distinguished by firm stable attachment following preliminary capture. Subdivided into Stigmatic_Exudate_Binding emphasizing molecular mechanisms and Pollen_Grain_Attachment focusing on stable association. Develops over 15-60 minutes following adhesion initiation. Critical role of lipid transfer proteins and exudate matrix. Substantial resistance to detachment forces. Boron nutrition essential for normal exudate and adhesion. Enables subsequent pollen hydration, recognition, germination. Essential stable association for reproductive success.
Pollenadhäsion
feste Pollenanbindung
firm pollen binding
pollen-stigma adhesion
secure pollen retention
sichere Pollenretention
stabile Pollenhaftung
stable pollen attachment
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 380-385 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 287-292 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Pollen Adhesion
Chardonnay in Champagne shows pollen adhesion initiation during early June Year N+1 at BBCH 60-65, with stigmatic exudate containing lipid transfer proteins and polysaccharides creating molecular adhesion within 30-90 seconds of pollen contact, preliminary attachment strengthening over 8-15 minutes through exudate binding, pollen grain beginning hydration within 15-45 minutes enabling metabolic activation, under cool maritime conditions of 19-24°C temperature, 58-72% relative humidity, typical of this early-bursting cultivar with adequate boron nutrition supporting normal exudate production and adhesion development
Pollen adhesion initiation represents the beginning attachment phase following pollen-stigma contact when preliminary binding develops through stigmatic exudate interaction with pollen surface, establishing initial stable association enabling transition to firm adhesion and subsequent recognition processes. This critical attachment initiation occurs during pollen capture in early-mid June in temperate regions Year N+1, typically proceeding over seconds to minutes following initial contact as part of BBCH 60-65 flowering stages. Adhesion initiation mechanism involves stigmatic exudate proteins and polysaccharides contacting pollen grain surface creating molecular interactions, physical adhesion developing through viscous exudate properties resisting grain detachment, and beginning recognition processes as pollen coat substances interact with exudate components. Stigmatic exudate composition enabling adhesion includes proteinaceous components particularly lipid transfer proteins creating adhesive binding, polysaccharides providing viscous matrix and molecular recognition sites, lipids affecting surface properties and possibly pollen coat interaction, and water providing hydration medium for molecular interactions and pollen grain rehydration. Lipid transfer protein function shows adhesive role with proteins abundant in stigmatic exudate binding pollen coat lipids, creating molecular bridges between exudate and pollen surface, and initiating recognition processes through protein-lipid interactions enabling species identification and compatibility assessment. Exudate viscosity providing adhesion shows physical adhesive properties with viscous fluid resisting forces that would dislodge grain, maintaining pollen positioning on stigma surface through mechanical resistance, and creating stable attachment even before molecular binding fully develops. Pollen coat interaction with exudate involves pollen surface substances from tapetal origin including proteins, lipids, and polysaccharides contacting exudate components, molecular recognition occurring through complementary binding sites, and triggering of downstream processes including enhanced adhesion and hydration support if compatibility confirmed. Molecular recognition initiation shows beginning species and compatibility assessment with pollen coat proteins interacting with stigma exudate receptors, self-recognition signals in self-compatible grapevine confirming appropriate species and enabling continued adhesion development, and acceptance response triggering enhanced exudate binding and pollen hydration support. Pollen grain hydration beginning shows initial water uptake from stigmatic exudate occurring within 15-90 minutes of contact, grain expanding slightly as cells rehydrate from desiccated atmospheric state, and metabolic activation initiating as hydration restores physiological function enabling subsequent germination. Adhesion strength development shows progressive increase from weak preliminary attachment immediately after contact, strengthening over minutes as molecular interactions develop, to moderate adhesion within 5-15 minutes providing resistance to environmental disturbances, and transition to firm adhesion over 15-60 minutes creating stable long-term attachment. Mechanical adhesion testing shows increasing resistance to detachment forces with preliminary adhesion resisting only gentle disturbances within seconds of contact, developing adhesion withstanding moderate mechanical forces within minutes, and mature adhesion requiring substantial force for grain removal after 15-30 minutes. Environmental effects on adhesion initiation include temperature affecting exudate viscosity and molecular interaction kinetics with optimal 22-28°C supporting efficient adhesion development, humidity influencing exudate water content and consistency with moderate 50-65% maintaining appropriate properties, and rain catastrophically disrupting adhesion through diluting or washing away exudate preventing attachment development. Temperature effects show warm conditions promoting faster adhesion development through enhanced molecular interaction kinetics, cool conditions slowing adhesion initiation potentially requiring longer time for stable attachment, and optimal 24-26°C range supporting rapid efficient adhesion development within minutes. Humidity effects include moderate levels maintaining appropriate exudate consistency for adhesion with proper water content supporting molecular interactions, high humidity above 75% potentially diluting exudate weakening adhesive properties, and low humidity below 45% potentially concentrating exudate affecting diffusion and molecular mobility. Boron nutrition effects show critical importance with adequate boron essential for normal stigmatic exudate production and composition, deficiency causing reduced exudate quantity or altered composition impairing adhesion initiation, and emphasizing importance of sufficient boron availability during flowering for normal pollination function. Failed adhesion initiation results from inadequate exudate production preventing preliminary binding development, abnormal exudate composition lacking essential adhesive components, pollen grain damage compromising surface properties preventing interaction, or environmental interference particularly rain removing exudate before adhesion establishes. Multiple pollen adhesion initiation shows independent processes with each contacted grain undergoing separate adhesion development, simultaneous adhesion of multiple grains on same stigma creating pollen surplus, and typical stigma supporting 10-50 attached grains during peak pollination though only 1-2 ultimately germinating successfully. Adhesion as reversible process shows early adhesion potentially reversible if environmental disturbance or molecular recognition failure occurs, transition to irreversible adhesion as molecular binding strengthens and hydration proceeds, and point of no return reached within 15-30 minutes when firm attachment establishes. Stigma papillate cell response to adhesion shows possible subtle changes in cell metabolism supporting pollen attachment, continued exudate secretion maintaining adhesive environment, and beginning preparation for supporting pollen germination and tube growth. The process represents second aspect of pollen capture emphasizing attachment development, following pollen-stigma contact. Duration spans seconds to minutes as preliminary adhesion develops transitioning to stable attachment. Cultivar variation in adhesion initiation may include exudate composition affecting adhesive properties, exudate production rate affecting attachment speed, and pollen coat characteristics affecting molecular interactions. Regional timing shows adhesion initiation occurring during flowering period across all climates immediately following pollen-stigma contact events coinciding with pollen availability and stigma receptivity. Viticultural significance includes representing critical transition from transient contact to stable attachment, requiring adequate stigma exudate production and appropriate environmental conditions, demonstrating importance of boron nutrition for normal function, and establishing successful adhesion as prerequisite for subsequent germination and fertilization. Research applications include analyzing molecular mechanisms of adhesion including protein-lipid interactions, investigating exudate composition and adhesive properties, evaluating environmental and nutritional effects on adhesion development, quantifying adhesion strength dynamics and kinetics, and assessing relationships between adhesion characteristics and pollination success.
Pollen_Adhesion_Initiation is second aspect of Pollen_Capture, distinguished by beginning attachment development following contact. Represents transition from transient contact to preliminary stable association. Duration seconds to minutes as molecular and physical adhesion develops. Involves stigmatic exudate interaction with pollen surface. Critical role of lipid transfer proteins and exudate viscosity. Beginning pollen hydration and recognition. Transitions to Pollen_Adhesion as firm attachment establishes. Boron nutrition essential for normal function.
Adhäsions-Initiierung
Bindungsbeginn
adhesion beginning
initial pollen binding
initiale Pollenanbindung
pollen retention establishment
preliminary pollen attachment
vorläufige Pollenhaftung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 380-382 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 287-290 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Pollen Adhesion Initiation
Cabernet Sauvignon in Napa Valley shows pollen capture during early-mid June Year N+1 at BBCH 60-65 over 7-9 days, with successful captures occurring predominantly through direct gravitational transfer from anthers 0.8-1.2 mm above stigmas supplemented by short-range wind-assisted delivery, papillate stigma surfaces with exudate coating capturing and retaining 12-35 pollen grains per stigma during peak period, under warm climate conditions of 24-29°C temperature, 50-64% relative humidity, supporting efficient capture enabling 58-68% fruit set typical of this late-season cultivar
Pollen capture represents the initial pollination phase when airborne or directly transferred pollen grains contact receptive stigma surfaces and achieve retention through physical and adhesive mechanisms, establishing first critical step in pollen-stigma interaction enabling subsequent adhesion and germination. This essential capture process occurs during flowering in early-mid June in temperate regions Year N+1, typically spanning 5-10 days as flowers sequentially open and become receptive for pollen capture across cluster, corresponding to BBCH 60-65 stages during anthesis period. Capture mechanisms involve multiple processes including pollen-stigma contact through aerial delivery or direct transfer establishing physical encounter, initial retention by stigma surface features preventing immediate loss, and beginning adhesion through stigmatic exudate interaction creating preliminary attachment. Pollen delivery modes enabling capture include gravitational transfer with pollen falling directly from dehiscing anthers in same flower representing 60-80% successful captures in self-pollination, wind-assisted aerial dispersal carrying pollen grains to stigmas within cluster or between neighboring plants contributing 20-35% captures, and occasional insect contact though playing minimal role <5% in grapevine pollen capture compared to dominant wind-gravity mechanisms. Self-pollination capture efficiency shows high success through close spatial proximity with anthers positioned 0.5-1.5 mm above stigma facilitating direct pollen fall, anther dehiscence typically occurring within 6-24 hours of stigma receptivity development creating temporal coordination, and favorable geometry supporting efficient pollen transfer from anthers to stigma in same flower. Wind-assisted capture dynamics involve atmospheric pollen transport delivering grains to receptive stigmas, pollen trajectory interception by protruding stigma structures, and contact probability depending on pollen concentration, wind patterns, and stigma exposure, with capture efficiency declining with distance from pollen source. Stigma morphology affecting capture includes bilobed structure creating expanded 2-4 mm² surface area providing increased interception probability, papillate epidermal cells with elongated 40-80 micrometer projections creating rough textured surface enhancing capture, stigma positioning above flower receptacle creating exposure to airborne pollen, and overall architecture optimized for efficient pollen grain interception. Papillate cell function in capture shows mechanical role with projecting cells creating physical barriers catching contacted pollen, increased surface area providing greater capture probability, and three-dimensional structure creating pollen retention sites preventing grain loss after initial contact. Stigmatic exudate role in capture includes adhesive properties providing immediate preliminary binding upon contact, chemical signals potentially attracting or guiding pollen though evidence limited in grapevine, and hydration medium enabling pollen recognition and attachment initiation. Pollen-stigma contact geometry shows typical scenario with pollen grain 25-35 micrometers diameter contacting papillate cell tip or side, initial mechanical retention by cell morphology preventing immediate rebound or loss, and exudate coating providing adhesive interface enabling attachment. Contact probability factors include pollen availability with higher atmospheric concentration increasing contact likelihood, stigma receptivity status with active exudate production optimizing capture, spatial positioning with well-exposed stigmas showing greater capture success, and environmental conditions affecting both pollen dispersal and stigma function. Multiple pollen capture per stigma shows typical pattern with receptive stigma receiving 10-50 pollen grains during peak pollination period though only 1-2 successful germinations required for fertilization, excess captures providing reproductive assurance compensating for possible germination failures, and demonstrated pollen surplus supporting high fertilization success. Capture timing shows rapid initial contact occurring within seconds to minutes once pollen reaches stigma vicinity, preliminary retention establishing within seconds through mechanical and adhesive mechanisms, and transition to stable adhesion proceeding over minutes to hours as exudate binding strengthens. Stigma receptivity requirements for successful capture include active exudate production creating adhesive surface, papillate cell expansion creating optimal capture geometry, and metabolic readiness supporting pollen recognition and hydration, developing within 6-24 hours after flower opening and providing 4-7 day capture window. Environmental effects on capture efficiency include temperature affecting pollen viability during transfer and stigma exudate properties, humidity influencing pollen dispersibility and exudate consistency, wind conditions affecting aerial delivery and contact probability, and rain catastrophically preventing capture through washing pollen away and diluting or removing exudate. Temperature effects on capture show optimal 22-28°C supporting both pollen viability during dispersal and stigma exudate function, cool conditions potentially reducing capture efficiency through impaired exudate production or consistency, and warm conditions generally favorable though excessive heat above 33°C potentially affecting stigma function. Humidity effects include moderate 50-65% levels optimal for maintaining pollen dispersibility during transfer and appropriate exudate consistency for capture, high humidity above 75% potentially diluting exudate reducing capture efficiency, and low humidity below 45% potentially affecting exudate properties though generally acceptable for mechanical capture. Rain effects show devastating interference with water washing newly captured pollen from stigma surfaces, diluting or removing exudate eliminating adhesive capture capacity, preventing pollen delivery through elevated humidity impeding dispersal, and physically damaging stigma structures, causing severe capture failure and fruit set reduction. Capture success assessment methods include stigma examination under magnification counting captured pollen grains, fluorescence microscopy techniques visualizing pollen on stigma surfaces, and pollination bag experiments excluding pollen sources evaluating capture efficiency under controlled conditions. The process shows two distinguishable aspects progressing from pollen-stigma contact establishing initial physical encounter, through pollen adhesion initiation beginning attachment processes. Duration represents rapid events occurring over seconds to minutes for individual captures, with cluster-level capture period spanning 5-10 days as flowers sequentially achieve receptivity. Cultivar variation in capture efficiency shows differences in stigma morphology affecting interception, exudate production affecting adhesion capacity, and anther-stigma positioning affecting self-pollination capture success. Regional timing shows pollen capture during late May-early June in warm climates, early-mid June in temperate regions, and mid-late June in cool areas, coinciding with flowering period when pollen available and stigmas receptive. Viticultural significance includes representing critical first step in pollination determining fruit set potential, requiring favorable environmental conditions avoiding rain during capture period, demonstrating importance of coordinated pollen availability and stigma receptivity, and establishing capture success as initial determinant of subsequent fertilization and berry development. Research applications include analyzing capture mechanisms including aerodynamic, morphological, and adhesive factors, quantifying capture efficiency under varying environmental conditions, evaluating stigma morphology effects on capture success, investigating pollen delivery dynamics and contact probabilities, and assessing relationships between capture characteristics and ultimate fruit set outcomes.
Pollen_Capture is first major Pollination phase establishing initial pollen-stigma contact and retention. Subdivided into Pollen_Stigma_Contact and Pollen_Adhesion_Initiation. Represents critical first step linking pollen delivery to subsequent adhesion. Predominantly gravitational self-pollination in grapevine supplemented by wind-assisted transfer. Stigma morphology and exudate essential for efficient capture. Rapid individual events seconds to minutes. Environmental sensitivity particularly to rain washing captured pollen away. Transitions to Pollen_Adhesion as attachment strengthens.
Pollen-Narben-Kontaktaufbau
Pollenabfang
Pollenfang
Pollenretentions-Initiierung
pollen interception
pollen retention initiation
pollen-stigma contact establishment
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 378-382 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 287-290 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Pollen Capture
A single grapevine flower produces 8,000-25,000 pollen grains depending on cultivar and flower sex type, with hermaphrodite flowers of Cabernet Sauvignon releasing 10,000-15,000 grains per flower. Pollen diameter ranges from 22-32 µm across cultivars, with tricolporate aperture morphology characteristic of Vitaceae. Pollen release occurs primarily between 06:00-12:00 hours during anthesis, coinciding with calyptra abscission. Atmospheric pollen concentrations in vineyards during peak bloom reach 200-800 grains/m³.
A flowering disposition that is the inherited capacity of a Vitis vinifera plant to produce, mature, and release pollen grains (male gametophytes) with characteristic properties affecting fertilization success, encompassing pollen quantity, structural integrity, physiological viability, and functional performance during pollination.
Pollen_Disposition serves as the parent class for traits affecting male reproductive function. Grapevine pollen is binucleate at release, with generative cell division occurring during pollen tube growth. The exine structure shows species-specific sculpturing patterns useful for taxonomic identification. Pollen traits significantly affect fruit set, seed number, and berry development through both quantity and quality effects on fertilization.
Pollenbildungsvermögen
Pollendisposition
Pollenqualität
male gametophyte disposition
pollen production trait
pollen quality capacity
Gallardo A et al. (2009) Pollen development and fertility in Vitis vinifera L. cv. Macabeo: a comparison between male-sterile and fertile flowers. American Journal of Enology and Viticulture 60:353-361 Vasconcelos MC et al. (2009) The flowering process of Vitis vinifera: a review. American Journal of Enology and Viticulture 60:411-434
Pollen Disposition
Chardonnay in Champagne shows pollen exhaustion during early July Year N+1 at BBCH 67-69 over 2-3 days July 1-3, with sequential anther depletion progressing from apical to basal positions, empty withered anthers showing brown discoloration lacking yellow pollen mass, atmospheric pollen contribution ceasing from exhausted anthers, under cool maritime conditions of 20-24°C temperature, typical of this early-bursting cultivar completing male gamete discharge 12-14 days after first dehiscence with exhaustion marking effective end of male reproductive contribution
Pollen exhaustion represents the anther depletion state when viable pollen supplies have released completely through sustained discharge leaving anthers empty of functional grains, eliminating continued male gamete production and marking effective termination of pollen contribution to pollination. This critical depletion state occurs during completed pollen release in late June-early July in temperate regions Year N+1, typically developing progressively over 2-3 days as sequential anthers complete discharge and exhaust supplies, corresponding to BBCH 67-69 late-complete anthesis stages. Exhaustion characteristics show complete depletion of releasable pollen from anther locules with all viable grains having discharged during previous days of active release, residual non-releasable pollen representing damaged or adhered grains unable to liberate estimated <5% original content, and overall cessation of functional pollen production from exhausted anthers. Depletion progression shows characteristic spatial-temporal pattern with earliest-dehiscing anthers in apical cluster regions reaching exhaustion first after 2-3 days sustained discharge, progressive extension to middle regions as anthers sequentially exhaust following their individual discharge timing, and late-dehiscing basal anthers reaching exhaustion last completing cluster-level depletion, creating developmental gradient in exhaustion state. Anther appearance at exhaustion shows empty locules visible through longitudinal dehiscence slits exposing interior lacking yellow pollen mass, withered shrunken anther structure reflecting tissue dehydration following exhaustion, color change from yellow-brown fresh state toward brown or gray-brown exhausted appearance, and overall visual evidence of completed discharge. Locular emptiness verification through dissection or examination shows depleted chambers lacking pollen mass, residual individual grains or small clumps adhering to locule walls representing non-releasable fraction, and structural evidence of completed pollen production and release cycles. Pollen release cessation from exhausted anthers shows eliminated discharge capacity with no further viable grains available for liberation, occasional release of residual damaged or adhered grains contributing negligible functional pollen, and overall terminated pollen production from individual exhausted anther. Atmospheric pollen contribution cessation shows exhausted anthers no longer supplying fresh viable grains to atmospheric pool, eliminated source function preventing atmospheric concentration maintenance, and reliance on any remaining late-dehiscing anthers for final minor pollen contributions. Residual pollen characteristics in exhausted anthers include damaged grains with compromised exine or cytoplasm preventing normal liberation, adhered grains stuck to locule walls unable to release despite dehiscence, and overall non-functional pollen representing discharge failures rather than viable reserves. Exhaustion timing relative to dehiscence shows typical 1-3 day period from initial opening to complete exhaustion depending on environmental conditions and discharge efficiency, accelerated exhaustion under optimal conditions promoting rapid complete discharge within 24-48 hours, and delayed exhaustion under suboptimal conditions extending period to 3-5 days. Environmental effects on exhaustion timing include favorable conditions facilitating rapid efficient discharge producing earlier exhaustion, suboptimal conditions particularly high humidity potentially retaining pollen in locules extending functional discharge period, and temperature affecting discharge rate influencing exhaustion timing. Temperature effects show warm conditions 25-30°C promoting rapid discharge and earlier exhaustion, cool conditions 18-23°C slowing discharge potentially extending functional period before exhaustion, and overall thermal environment affecting depletion dynamics. Humidity effects include moderate levels 50-65% supporting efficient discharge enabling normal exhaustion timing, elevated humidity above 70% potentially retaining pollen in locules through reduced mechanical liberation delaying exhaustion, and low humidity below 45% potentially accelerating discharge though possibly affecting released pollen viability. Anther tissue changes accompanying exhaustion include progressive dehydration as discharge completes with tissue water content declining, cellular senescence initiating in post-reproductive anthers with metabolic decline, and structural collapse as turgid functional anthers become withered exhausted structures. Metabolic transition at exhaustion shows declining respiratory activity in depleted anthers as metabolic demands reduce, resource reallocation from male reproductive function toward other processes particularly berry development, and physiological shift from active reproductive investment to senescence. Male reproductive contribution assessment at exhaustion enables evaluation through comparison of early versus late exhaustion timing affecting pollination window duration, discharge efficiency affecting functional pollen proportion successfully released, and coordination with stigma receptivity affecting pollination success outcomes. Fruit set implications of exhaustion timing include early complete exhaustion before full stigma receptivity potentially limiting late flower pollination opportunities, optimal exhaustion timing coinciding with receptivity conclusion enabling maximum pollination efficiency, and delayed exhaustion extending male contribution window though likely producing minimal additional fruit set. Sequential exhaustion pattern throughout cluster creates extended pollination opportunity with early anthers providing initial pollen while later anthers extend supply, progressive depletion creating declining male gamete availability over flowering period, and final exhaustion completing male reproductive contribution approximately coinciding with stigma receptivity cessation. Pollination success relationship to exhaustion shows inverse correlation with rapid early exhaustion under optimal conditions typically coinciding with successful pollination during favorable period supporting high fruit set 60-75%, while premature exhaustion from stress or abnormality potentially limiting pollination reducing set <40%, and demonstrating importance of sustained appropriate-duration discharge. Observational assessment of exhaustion includes visual inspection of anther appearance noting empty withered structures, absence of fresh pollen release during favorable conditions indicating depletion, and dissection or detailed examination confirming locular emptiness if required for verification. The process represents first aspect of completed pollen release emphasizing depletion of functional reserves, followed by empty anther formation discussing post-exhaustion structural state. Duration represents progressive state developing over 2-3 days as sequential anthers deplete, with spatial-temporal gradient reflecting flowering progression. Cultivar variation in exhaustion characteristics includes differences in pollen production affecting quantity available for discharge before depletion, discharge efficiency affecting completeness of release and residual retention, and timing relative to stigma receptivity affecting pollination opportunity and fruit set outcomes. Regional timing shows progressive pollen exhaustion during late June in warm climates, late June-early July in temperate regions, and early-mid July in cool areas, occurring during late-complete anthesis as male reproductive phase concludes. Viticultural significance includes marking effective end of male gamete contribution to fruit set, indicating completion of critical pollination window, providing assessment point for male reproductive success, and establishing transition toward exclusive developmental focus on established berries. Research applications include analyzing factors affecting exhaustion timing and completeness, evaluating discharge efficiency through residual pollen assessment, quantifying relationships between exhaustion patterns and fruit set outcomes, assessing environmental effects on depletion dynamics, and investigating genetic variation in pollen production and discharge affecting reproductive capacity.
Pollen_Exhaustion is first aspect of Completed_Pollen_Release, distinguished by depletion of functional anther reserves. Represents progressive state developing over 2-3 days with spatial-temporal gradient. Marks effective end of male gamete contribution to pollination. Followed by Empty_Anther_Formation discussing post-exhaustion structural state. Assessment enables evaluation of male reproductive success. Timing relative to stigma receptivity affects final pollination opportunity. Spatial progression reflects flowering developmental gradient.
Pollenerschöpfung
Pollenvorraterschöpfung
complete pollen discharge
male gamete depletion
männliche Gametendepletion
pollen supply exhaustion
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 390-393 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 297-300
Pollen Exhaustion
Syrah anthers at BBCH 59-60 (flower buds swollen to beginning of flowering) exhibit pollen filled sacs visible as swollen anther lobes with bright yellow coloration, containing 3,200-6,800 mature pollen grains per anther distributed between two pollen filled locules (one per lobe, each formed from fusion of two original pollen sacs), with pollen densely packed at approximately 4,500-6,000 grains per mm³ and ready for imminent release through dehiscence when calyptra shedding exposes anthers to air-drying conditions. Pinot Noir at early BBCH 61 shows pollen filled sacs in transition to pollen release, with narrow longitudinal dehiscence slits (0.08-0.12 mm wide) beginning to open allowing initial pollen escape while majority of pollen (70-85% of total 15,000-25,000 grains per five-anther flower) remains packed within sacs creating continued yellow anther coloration. Gewürztraminer functionally female flowers exhibit reduced pollen filled sacs containing only 400-1,200 grains per anther (2,000-6,000 total per flower) compared to hermaphrodite norms (15,000-30,000), with sacs showing yellow coloration indicating pollen presence but reduced filling density and smaller locule volumes reflecting overall stamen reduction in this flower type.
A pollen filled sac is a pollen sac containing mature pollen grains accumulated to capacity following completion of pollen maturation and prior to pollen release through dehiscence, representing the pre-dehiscence to early-dehiscence state (BBCH 59-61) when pollen reserves are at maximum and when the pollen sac functions as a pollen storage compartment ready for imminent dispersal. Pollen filled sacs are characterized by= (1) locule cavity fully occupied by densely packed mature pollen grains, with packing density typically 3,000-8,000 grains per mm³ creating tightly filled configuration (2) following septum breakdown (BBCH 59-60), each anther lobe contains one large pollen filled locule (formed from fusion of two original pollen sacs) containing 1,500-3,750 mature pollen grains (hermaphrodite flowers) or more (staminate flowers) or fewer (functionally female/pistillate flowers) (3) pollen grains showing mature morphology with characteristic Vitis tricolporate structure, yellow color from carotenoid content, and full complement of starch and lipid reserves (4) tapetum completely degenerated and absorbed, with only residual tapetal debris sometimes visible on pollen surfaces (5) anther wall layers mature and prepared for dehiscence with endothecium showing fully developed secondary wall thickenings ready to generate hygroscopic tensions. Pollen filled sacs in mature pre-dehiscence anthers (BBCH 59) remain enclosed by intact anther wall with stomium not yet opened, maintaining pollen in protected environment with controlled humidity and temperature buffering until exposure and dehiscence occur. Pollen filled sacs in early-dehiscing anthers (BBCH 60-61) transition from enclosed to exposed state as longitudinal stomium slits open, with initial dehiscence creating narrow openings (0.05-0.15 mm wide) through which pollen begins to escape while majority of pollen remains packed within the locule. The pollen content of filled sacs represents the culmination of weeks of developmental investment= microsporogenesis requiring 6-12 days (BBCH 55-56) pollen maturation requiring 8-15 days (BBCH 57-59) with total pollen sac development from initiation to pollen-filled state spanning 20-35 days depending on temperature. The pollen filled state is transient for individual anthers, typically persisting 6-24 hours from achievement of full maturity (late BBCH 59) through initiation and progression of dehiscence (BBCH 60-61) until substantial pollen release creates transition to partially empty or empty pollen sac state. Environmental conditions critically affect the duration and fate of the pollen filled state= favorable dehiscence conditions (warm temperatures 22-30°C, declining humidity 70-80% to 40-50%, light air movement) promote rapid dehiscence and efficient pollen release, minimizing pollen filled sac duration and maximizing pollen dispersal unfavorable conditions (cool temperatures 85%, rain) delay or prevent normal dehiscence, prolonging pollen filled sac state beyond normal duration and potentially causing pollen deterioration within sacs if conditions persist for days. Prolonged pollen filled sac retention beyond 24-48 hours due to adverse conditions can impair pollen viability through= inadequate desiccation maintaining pollen at suboptimal moisture content for germination fungal infection of enclosed pollen by Botrytis or other pathogens entering through eventual partial dehiscence metabolic deterioration of pollen reserves as respiratory consumption continues without pollen dispersal and germination. The pollen filled sac state represents maximum male reproductive potential, with pollen quantity and quality at peak and poised for dispersal, making this the optimal stage for anther collection in breeding programs for controlled crossing, with anthers collected at late BBCH 59 or very early BBCH 60 yielding maximum viable pollen for extraction and application.
Pollen_Filled_Sac represents the mature pre-release state containing maximum pollen reserves and is distinguished from Empty_Pollen_Sac (post-dehiscence, pollen substantially or completely released) by presence of dense pollen packing. The pollen filled state is the critical pre-dispersal phase when pollen is ready for release but still protected within anther tissues, representing peak male reproductive potential. The duration and fate of the pollen filled state critically depends on environmental conditions determining dehiscence timing and efficiency, making this a key period when weather affects pollination success. Monitoring pollen filled sac status (presence of swollen yellow anthers) provides indicators of imminent bloom and reproductive potential, while prolonged pollen filled state without dehiscence indicates unfavorable bloom conditions requiring diagnosis and potentially management intervention.
gefüllte Pollenkammer
mature pollen chamber
pollen-loaded sac
pollengefüllter Sack
voller Pollensack
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 101-104 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 95-97.
Pollen Filled Sac
Syrah in Rhône Valley shows pollen germination during mid-June Year N+1 at BBCH 60-65, with attached pollen grains 28-32 micrometers diameter completing hydration within 25-60 minutes, initiating tube emergence 1.5-3.5 hours after stigma contact, producing 8-12 micrometer diameter tubes, under warm Mediterranean conditions of 25-29°C temperature, adequate boron nutrition supporting normal wall synthesis, typical of this mid-to-late season cultivar with 68-82% germination success from viable attached pollen enabling subsequent fertilization
Pollen germination represents the critical post-adhesion process where hydrated pollen grain on receptive stigma undergoes metabolic activation, cytoplasmic reorganization, and pore activation culminating in pollen tube emergence through grain aperture, establishing essential transition from dormant pollen to actively growing tube enabling sperm cell delivery to ovules for fertilization. This essential germination process occurs during flowering in early-mid June in temperate regions Year N+1, typically initiating within 1-4 hours after pollen-stigma contact and adhesion under favorable conditions, corresponding to BBCH 60-65 flowering stages. Germination mechanism involves coordinated processes including pollen grain hydration from stigmatic exudate restoring metabolic function, germination pore activation at one of three colporate apertures preparing emergence site, cytoplasmic polarization establishing growth axis, and tube emergence as vegetative cell extends through activated aperture creating pollen tube. Pollen grain hydration preceding germination shows water uptake from stigmatic exudate over 15-90 minutes following adhesion, grain expanding 10-20% volumetrically as cells rehydrate from desiccated atmospheric state to functional hydrated condition, and metabolic activation proceeding as hydration restores enzyme function, respiratory activity, and biosynthetic capacity enabling germination. Hydration dynamics show initial rapid uptake phase within first 15-30 minutes absorbing majority of water requirement, slower equilibration phase over subsequent 30-60 minutes achieving final hydrated state, and total water content increase from 5-15% desiccated state to 60-70% hydrated state approaching fresh pollen levels. Metabolic activation during hydration involves respiratory enzymes reactivating enabling ATP production through oxidative phosphorylation, protein synthesis initiating producing enzymes and structural components required for tube growth, carbohydrate metabolism activating mobilizing stored starch and lipid reserves providing energy and building materials, and overall physiological transition from dormant to actively metabolizing state. Cytoplasmic reorganization preparing germination shows polarization with cytoplasm concentrating at germination site, organelles including mitochondria and rough endoplasmic reticulum migrating toward emergence site supporting active metabolism and secretion, nucleus positioning near germination pore though initially remaining in grain, and vegetative cell preparing for tube formation and growth. Germination pore selection shows pollen grain having three colporate apertures providing potential germination sites, typically single aperture activating for tube emergence though occasionally multiple tubes may initiate, and pore selection possibly influenced by contact geometry, stigma surface properties, or intrinsic polarity though mechanisms poorly understood. Germination pore activation involves localized cell wall loosening at selected aperture through enzymatic degradation, intine layer preparation for tube extension with wall modification enabling growth, turgor pressure building in vegetative cell creating mechanical force driving emergence, and coordinated structural changes enabling tube breakthrough. Tube emergence timing shows germination typically initiating 1-4 hours after pollen-stigma contact under optimal conditions, earlier germination at 1-2 hours with warm temperature 25-28°C and vigorous pollen, delayed germination at 3-6 hours with cool temperature 18-22°C or aged pollen, and variable timing depending on pollen viability, stigma receptivity, and environmental conditions. Tube emergence morphology shows initial small protrusion 2-5 micrometers diameter emerging through activated aperture, rapid elongation as tube extends from grain driven by tip growth, and developing cylindrical tube structure with diameter 8-12 micrometers in mature actively growing tube. Pollen tube characteristics include vegetative cell cytoplasm occupying tube with organelles and stored reserves supporting growth, generative cell or two sperm cells depending on mitotic timing traveling in tube cytoplasm, tip-focused growth with cell wall synthesis and extension occurring at tube apex, and basipetal cytoplasmic streaming maintaining nutrient and organelle supply to growing tip. Tube wall composition shows inner callose layer providing structural support and possibly regulatory functions, outer pectin-cellulose layer providing flexible wall enabling expansion, and continuous wall synthesis at tip driven by Golgi-derived vesicles delivering wall precursors. Germination requirements include adequate hydration providing water for metabolic function and turgor generation, appropriate temperature 20-28°C optimal supporting enzymatic activity and metabolic processes, sufficient stored reserves in pollen grain providing energy and materials for tube initiation, adequate boron essential for cell wall synthesis and tube integrity, and supportive stigma environment providing recognition signals and possibly nutrients. Boron critical role in germination emphasizes absolute requirement with boron essential for proper cell wall structure particularly pectin cross-linking, deficiency causing germination failure or abnormal tube development with bursting or irregular growth, and optimal boron concentration 10-20 ppm in stigmatic tissues supporting normal germination and tube function. Temperature effects on germination show optimal 22-28°C supporting rapid efficient germination within 1-2 hours, cool conditions 15-20°C substantially delaying germination to 4-8 hours and reducing success rate, warm conditions 28-32°C potentially accelerating germination though excessive heat above 33°C may cause pollen dysfunction, and temperature sensitivity demonstrating metabolic basis of germination process. Pollen viability effects show fresh high-viability pollen 75-90% germination capacity under optimal conditions, aged pollen with reduced viability 40-60% showing delayed or failed germination, and viability decline with atmospheric exposure emphasizing importance of rapid pollination during pollen release period. Stigma support for germination includes continued exudate secretion providing hydration and possibly nutrients, recognition signals confirming compatibility and triggering germination support, possible stigma-derived growth factors though evidence limited in grapevine, and overall receptive environment enabling successful germination. Multiple pollen germination per stigma shows typical pattern with 10-50 attached grains attempting germination, competitive germination with fastest-germinating or most vigorous tubes having advantage, and typically 1-2 tubes successfully navigating style and achieving fertilization despite numerous germination attempts. Germination success rate shows 60-85% attached viable pollen successfully germinating under favorable conditions, declining to 30-50% under suboptimal conditions or with reduced pollen viability, and demonstrating coordination of pollen quality, stigma support, and environmental factors determining germination outcomes. Failed germination causes include inadequate hydration from insufficient stigmatic exudate, pollen viability loss from aging or environmental damage, incompatibility recognition preventing stigma support though rare in self-compatible grapevine, boron deficiency preventing normal wall synthesis, temperature stress disrupting metabolic function, or developmental abnormalities affecting pollen or stigma function. The process shows three distinguishable aspects progressing from pollen hydration restoring metabolic function, through germination pore activation preparing emergence site, to pollen tube emergence creating actively growing structure. Duration typically spans 1-6 hours from adhesion to tube emergence depending on conditions, with rapid 1-2 hour germination under optimal conditions and delayed 4-8 hour germination under suboptimal conditions. Cultivar variation in germination may include pollen vigor affecting germination speed and success, stored reserve quantity affecting germination capacity, and temperature sensitivity affecting germination under variable conditions. Regional timing shows pollen germination occurring during flowering period across all climates within hours of successful pollination, proceeding throughout peak anthesis period as sequential pollinations occur. Viticultural significance includes representing essential post-pollination step linking pollen capture to fertilization, requiring favorable environmental conditions particularly adequate temperature and boron nutrition, demonstrating critical dependence on pollen quality and stigma support, and establishing germination success as prerequisite for subsequent tube growth, fertilization, and fruit set. Research applications include analyzing germination mechanisms including hydration dynamics and pore activation, investigating environmental and nutritional effects on germination success, evaluating pollen viability and germination capacity relationships, quantifying germination timing and tube emergence dynamics, and assessing germination success as determinant of fertilization and fruit set outcomes.
Pollen_Germination represents critical post-adhesion process within Flowering enabling transition from dormant grain to growing tube. Subdivided into Pollen_Hydration restoring function, Germination_Pore_Activation preparing emergence, and Pollen_Tube_Emergence creating growth structure. Duration 1-6 hours from adhesion to emergence. Critical dependence on temperature, boron nutrition, pollen viability. Essential prerequisite for Pollen_Tube_Growth and fertilization. Competitive process with multiple germinants per stigma but typically 1-2 successful completions.
Kornkeimung
Pollenkeimung
Pollenschlauch-Initiierung
grain germination
pollen sprouting
pollen tube initiation
reproductive activation
reproduktive Aktivierung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 382-385 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 288-292 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Pollen Germination
In vitro pollen germination on optimized agar media (15-20% sucrose, 100 ppm boric acid, 300 ppm calcium nitrate) reaches 60-85% in highly fertile cultivars such as Merlot and Syrah within 4-6 hours at 25°C. Chardonnay shows rapid germination initiation (50% germinated within 2 hours) while Sangiovese requires 4-5 hours to reach equivalent levels. In vivo germination on receptive stigmas occurs within 30-60 minutes of pollination, with germination percentages 10-20% higher than in vitro due to stigmatic exudate factors. Optimal temperature for germination ranges from 25-30°C, with rates declining sharply below 15°C.
A pollen disposition that is the inherited capacity of a Vitis vinifera plant to produce pollen grains that successfully germinate on compatible stigmatic surfaces or in artificial media, forming a pollen tube initial through hydration, polarization, and cell wall extension at the aperture site.
Pollen germination represents the first phase of post-pollination development, distinct from subsequent pollen tube growth. The process requires adequate stigma receptivity, appropriate temperature, and sufficient moisture. Germination percentage is partially independent of viability, as some structurally intact grains fail to germinate due to metabolic deficiencies. Boron is critical for germination, with deficiency reducing rates by 40-70%.
Keimungsdisposition
Pollenkeimfähigkeit
Pollenkeimungsvermögen
germination capacity trait
pollen activation disposition
pollen germinability
Staudt G (1999) Opening of flowers and time of anthesis in grapevines, Vitis vinifera L. Vitis 38:15-20 Okamoto G, Imai S (1996) Effects of boric acid on pollen tube growth and fruit set in Vitis vinifera L. cv. Pione. Journal of the Japanese Society for Horticultural Science 65:283-289
Pollen Germination Disposition
Cabernet Franc in Loire Valley shows pollen grain attachment during mid-June Year N+1 at BBCH 60-65, with firmly secured grains 26-32 micrometers diameter on papillate stigma surfaces, attachment developing within 20-50 minutes through molecular binding and exudate adhesion creating resistance to moderate environmental disturbances, supporting 16-28 stably attached grains per stigma undergoing hydration and germination, under temperate maritime conditions of 21-26°C temperature, 58-72% relative humidity, typical of this mid-season cultivar with stable attachment enabling subsequent tube growth and fertilization supporting 56-66% fruit set
Pollen grain attachment represents the stable long-term pollen-stigma association state when firm molecular binding, physical adhesion, and mechanical retention create secure persistent attachment resistant to environmental disturbances, establishing optimal conditions for pollen hydration, recognition, germination, and successful fertilization. This critical stable attachment state occurs during pollen adhesion in early-mid June in temperate regions Year N+1, developing over 15-60 minutes following adhesion initiation and persisting through germination and tube growth, corresponding to BBCH 60-65 flowering stages. Attachment characteristics show firmly secured pollen grain on stigma surface through integrated molecular and physical binding mechanisms, stable positioning maintained despite environmental disturbances including wind, mechanical contact, or moderate moisture, pollen grain fully hydrated and metabolically active preparing for or undergoing germination, and overall robust association supporting reproductive success. Attachment security shows substantial resistance to detachment forces with established attachment withstanding forces 100-500 times grain weight, resistance to normal environmental disturbances encountered during flowering including wind gusts up to 15-20 km per hour, tolerance to brief rain exposure though sustained heavy rain potentially disrupting attachment, and overall stability sufficient for supporting multi-hour to multi-day processes of germination and tube penetration. Multi-point molecular binding creating attachment includes numerous protein-protein bridges between pollen coat and exudate creating redundant attachment points, protein-lipid interactions through lipid transfer proteins providing strong hydrophobic binding, polysaccharide networks embedding grain in exudate matrix distributing mechanical forces, and cumulative effect of hundreds to thousands of molecular interactions producing robust attachment. Physical adhesion contributing to attachment shows viscous stigmatic exudate matrix creating mechanical resistance to grain displacement, exudate gel-like properties distributing stress preventing localized failure, and maintained adhesion through exudate secretion continuing throughout receptivity period replacing any dilution or loss. Mechanical retention by stigma structure supplements molecular adhesion with papillate cell architecture creating three-dimensional retention geometry, cell projections providing physical barriers preventing grain sliding or rolling, and surface irregularities creating stable positioning sites maintaining grain location. Attached grain positioning shows typical intimate contact with papillate cell surface, grain potentially spanning multiple adjacent cells depending on size and geometry, and stable orientation maintained throughout hydration and germination processes. Pollen grain state during attachment shows progressive hydration over 15-90 minutes reaching full rehydration, metabolic activation proceeding as hydration restores physiological function, and preparation for or initiation of germination with tube emergence within 1-4 hours under favorable conditions if fully hydrated and recognized as compatible. Hydration dynamics during attachment involve continued water uptake from stigmatic exudate until equilibrium reached, grain expanding 10-20% volumetrically as cells rehydrate from desiccated atmospheric state, cytoplasmic reorganization proceeding as metabolic systems reactivate, and establishment of germination capacity as hydration completes. Recognition completion during attachment shows pollen-stigma molecular communication confirming compatibility, self-compatible grapevine accepting self-pollen triggering supportive responses, and recognized compatible pollen receiving enhanced exudate support and preparation for tube growth into stigma tissues. Stigma response to attached pollen shows continued exudate secretion maintaining adhesive and hydration environment, possible localized metabolic activation in papillate cells beneath attached grain preparing to support tube penetration, and provision of resources including water and possibly nutrients supporting pollen function. Multiple grain attachment per stigma shows typical 10-50 firmly attached grains on receptive stigma during peak pollination, independent attachment and hydration of each grain creating pollen surplus, and competitive or selective germination with typically only 1-2 grains successfully completing tube growth and fertilization despite numerous attachments. Attachment persistence shows stable association maintained throughout germination typically 1-4 hours post-attachment, continued attachment during tube emergence and penetration of stigma surface, and long-term association as germinated tube grows through style tissues over 12-36 hours requiring persistent grain attachment at entry point. Environmental stability of attachment includes maintained attachment across typical flowering temperature range 18-30°C with thermal stability of molecular and physical binding, tolerance to humidity variation 40-75% with adhesion persisting despite exudate water content changes, and resistance to light-moderate rain though sustained heavy precipitation potentially disrupting attachment particularly in recently attached grains. Attachment failure causes include inadequate exudate production or composition preventing binding development, pollen grain abnormalities compromising coat substances and binding capacity, environmental disruption particularly heavy rain diluting or washing away exudate and physically removing grains, or developmental problems affecting stigma or pollen function. Attachment assessment methods include microscopic observation documenting attached grain presence and number, mechanical testing evaluating detachment resistance quantifying attachment strength, fluorescence techniques visualizing attached and germinating pollen, and pollination exclusion experiments determining attachment success under varying conditions. Attachment as prerequisite for fertilization shows firmly attached grain as essential starting point for reproductive success, attachment enabling subsequent germination and tube growth, and failed attachment eliminating pollination opportunity for that pollen-stigma pair causing potential fruit set failure. Attachment completion timing relative to germination shows attachment typically secure within 15-60 minutes of initial contact, germination often initiating within 1-4 hours as pollen completes hydration and recognition, and overlapping attachment stabilization and germination preparation processes. The process represents second and final aspect of pollen adhesion emphasizing overall stable long-term association, following stigmatic exudate binding. Duration develops over 15-60 minutes as attachment strengthens, persisting hours to days supporting germination and tube growth. Cultivar variation in attachment may include grain size affecting contact area and binding opportunity, pollen coat composition affecting molecular binding quality, and compatibility factors affecting recognition and support provided to attached pollen. Regional timing shows pollen grain attachment occurring during flowering period across all climates following pollen capture, establishing as stable association within hour of initial contact. Viticultural significance includes representing essential stable pollen-stigma association enabling reproductive success, requiring integrated molecular binding, physical adhesion, and mechanical retention, demonstrating importance of adequate stigma exudate and favorable environmental conditions, and establishing attachment success as prerequisite for subsequent germination, fertilization, and fruit set. Research applications include analyzing attachment mechanisms and security, evaluating environmental and genetic factors affecting attachment stability, quantifying attachment strength and resistance to disturbance, investigating attachment-germination relationships, and assessing attachment success as determinant of pollination outcomes and fruit set.
Pollen_Grain_Attachment is second and final aspect of Pollen_Adhesion, distinguished by overall stable long-term association emphasis. Represents culmination of capture and adhesion processes creating persistent attachment. Develops over 15-60 minutes, persists hours to days. Integrates molecular binding, physical adhesion, mechanical retention. Substantial resistance to environmental disturbance. Enables pollen hydration, recognition, germination. Essential prerequisite for successful fertilization. Completes Pollination hierarchical documentation from capture through adhesion to stable attachment.
Pollenkorn-Anhaftung
firm pollen binding state
gesicherte Kornanheftung
persistent pollen adhesion
persistente Pollenadhäsion
secured grain attachment
stabile Pollen-Narben-Assoziation
stable pollen-stigma association
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 382-388 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 288-295 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Pollen Grain Attachment
Chardonnay in Champagne shows pollen hydration during early June Year N+1 at BBCH 60-65, with desiccated grains 8-12% water content absorbing water from stigmatic exudate, swelling 12-18% volumetrically over 35-75 minutes reaching 62-68% final water content, metabolic activation proceeding enabling germination initiation 1.5-3 hours post-adhesion, under cool maritime conditions of 19-24°C temperature, 60-72% relative humidity maintaining adequate exudate water content, typical of this early-bursting cultivar
Pollen hydration represents the essential water uptake process where desiccated pollen grain on receptive stigma absorbs water from stigmatic exudate restoring cellular hydration, metabolic function, and physiological activity enabling subsequent germination and tube growth. This critical rehydration process occurs during pollen germination in early-mid June in temperate regions Year N+1, typically proceeding over 15-90 minutes following pollen-stigma adhesion, corresponding to BBCH 60-65 flowering stages. Hydration mechanism involves water movement from stigmatic exudate into pollen grain through osmotic gradient, pollen grain water content increasing from desiccated 5-15% to hydrated 60-70% approaching fresh pollen levels, and cellular rehydration restoring normal cell structure, membrane function, and organelle activity. Water uptake kinetics show characteristic biphasic pattern with rapid initial phase absorbing 60-80% total water within first 15-30 minutes driven by steep osmotic gradient, slower equilibration phase over subsequent 30-60 minutes achieving final hydrated state, and total hydration period typically 30-90 minutes depending on initial pollen water status and environmental conditions. Osmotic driving force for hydration includes concentrated solutes in desiccated pollen cytoplasm creating strong osmotic potential typically -5 to -10 MPa, stigmatic exudate providing external water source at relatively high water potential approximately -0.5 to -1.5 MPa, and resulting osmotic gradient driving water influx into grain. Water entry pathways involve primary uptake through pollen wall and membranes with water permeating exine through existing pathways or defects, intine layer allowing water passage through less rigid wall structure, plasma membrane showing selective permeability possibly involving aquaporin water channels, and cytoplasm absorbing water restoring cellular hydration. Grain swelling during hydration shows volumetric expansion of 10-20% as cells rehydrate, exine outer wall showing limited expansion maintaining structural form, intine inner wall and plasma membrane expanding as cytoplasm swells, and visible grain size increase from 25-35 micrometers desiccated to 28-40 micrometers hydrated state. Cellular rehydration effects include membrane restoration with lipid bilayers regaining fluid structure from dehydrated rigid state, organelle function restoration as mitochondria, endoplasmic reticulum, and Golgi rehydrate enabling activity, enzyme reactivation as proteins regain proper conformation and catalytic function, and cytoplasmic reorganization preparing for germination. Metabolic activation following hydration involves respiratory enzyme systems reactivating enabling ATP synthesis through oxidative phosphorylation, protein synthesis machinery becoming functional allowing gene expression and translation, carbohydrate metabolism activating mobilizing stored starch and lipid reserves, and overall transition from dormant to actively metabolizing state. Membrane integrity restoration shows critical importance with proper hydration enabling membrane repair and function, rapid hydration potentially causing damage through osmotic stress if not properly controlled, and cell regulatory systems managing hydration rate preventing excessive swelling or membrane rupture. ATP synthesis resumption during hydration provides energy for cellular processes with mitochondrial respiration reactivating producing ATP from stored reserves, energy charge increasing from low dormant levels to high functional levels, and ATP availability enabling active transport, biosynthesis, and motility required for germination. Gene expression activation shows transcription initiating for germination-related genes including tube growth proteins, cell wall synthesis enzymes, and metabolic enzymes, translation proceeding producing proteins required for germination and tube development, and overall molecular activation preparing for germination. Stored reserve mobilization during hydration involves starch hydrolysis beginning providing glucose for energy metabolism, lipid bodies beginning breakdown providing fatty acids and glycerol, and reserve proteins potentially beginning degradation providing amino acids, establishing resource availability for tube growth. Cytoplasmic reorganization during hydration shows polarization initiating with cytoplasm and organelles beginning concentration at prospective germination site, microtubule and actin cytoskeleton reorganizing establishing polarity, and spatial organization preparing for directed tube emergence and growth. Hydration requirements include adequate stigmatic exudate providing water source, appropriate exudate composition supporting uptake without osmotic damage, pollen grain structural integrity enabling controlled water entry, and cellular regulation systems managing hydration process. Environmental effects on hydration include stigma exudate water content affected by humidity with moderate 50-65% maintaining appropriate exudate properties, temperature affecting water uptake kinetics with optimal 22-28°C supporting efficient hydration within 30-60 minutes, and excessive desiccation from low humidity <40% potentially affecting exudate availability delaying hydration. Temperature effects on hydration rate show warm conditions 25-28°C promoting rapid uptake completing within 20-40 minutes, cool conditions 18-22°C slowing uptake requiring 60-90 minutes for complete hydration, and temperature affecting both water diffusion rate and membrane permeability influencing overall hydration kinetics. Pollen desiccation severity effects show moderately desiccated pollen 8-12% water content rehydrating efficiently within 30-60 minutes, severely desiccated pollen <5% water content requiring longer 60-120 minutes and possibly showing reduced viability, and freshly released pollen with higher water content 12-15% rehydrating most rapidly within 15-30 minutes. Stigmatic exudate properties affecting hydration include water content determining availability, osmotic potential affecting gradient and uptake rate, viscosity influencing water movement, and possible solute composition affecting pollen uptake though poorly characterized. Hydration monitoring methods include microscopic observation documenting grain swelling, vital dyes assessing membrane integrity restoration, respirometry measuring metabolic activation, and molecular assays detecting gene expression activation. Rapid versus slow hydration effects show controlled gradual uptake maintaining cell integrity, excessively rapid uptake potentially causing membrane damage through osmotic stress, and regulatory systems in viable pollen managing uptake rate preventing damage. Failed hydration causes include inadequate stigmatic exudate from stigma dysfunction or environmental stress, pollen membrane damage from aging or environmental exposure preventing controlled uptake, incompatibility responses potentially limiting exudate availability though rare in self-compatible grapevine, or extreme environmental conditions disrupting exudate or pollen function. Hydration as germination prerequisite shows complete hydration essential before germination can proceed, partial hydration insufficient for metabolic activation and tube emergence, and hydration timing affecting germination timing with rapid hydration enabling earlier germination. The process represents first aspect of pollen germination establishing metabolic function, followed by germination pore activation and tube emergence. Duration typically spans 15-90 minutes from adhesion to complete hydration, with rapid 15-40 minute hydration under optimal conditions and delayed 60-120 minute hydration under suboptimal conditions or with severely desiccated pollen. Cultivar variation in hydration may include pollen desiccation tolerance affecting rehydration capacity, membrane properties affecting water uptake kinetics, and grain structural characteristics affecting hydration efficiency. Regional timing shows pollen hydration occurring during flowering period across all climates within minutes to hours following successful pollination and adhesion. Viticultural significance includes representing essential first step in germination enabling metabolic activation, requiring adequate stigmatic exudate water content, demonstrating dependence on favorable environmental conditions, and establishing hydration completion as prerequisite for subsequent germination and tube growth. Research applications include analyzing water uptake mechanisms and kinetics, investigating cellular changes during rehydration, evaluating environmental effects on hydration efficiency, quantifying metabolic activation dynamics following hydration, and assessing hydration success as determinant of germination and fertilization outcomes.
Pollen_Hydration is first aspect of Pollen_Germination, distinguished by water uptake and metabolic restoration. Represents essential prerequisite enabling all subsequent germination processes. Duration 15-90 minutes showing biphasic uptake. Temperature and initial pollen water status affect kinetics. Cellular rehydration restores membrane function, organelle activity, enzyme catalysis. Transitions to Germination_Pore_Activation as preparation completes. Critical first step from dormant grain to active reproductive unit.
Kornwasseraufnahme
Pollenhydratation
Pollenimbibition
Pollenrehydrierung
grain water uptake
pollen imbibition
pollen rehydration
water absorption
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 382-384 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 288-290
Pollen Hydration
Thompson Seedless serves in a Pollen_Parent_Role for seedlessness breeding, its pollen applied to seeded maternal varieties—the only crossing direction yielding viable seeds for stenospermocarpic trait evaluation. A wild Vitis species with powdery mildew resistance genes bears a Pollen_Parent_Role in resistance introgression programs, with pollen collected, stored, and shipped to breeding programs for application to V. vinifera seed parents. Pinot Noir functions as pollen parent in clonal diversity studies, its pollen applied to emasculated Pinot Noir flowers to evaluate selfing effects and outcrossing vigor in a normally outcrossing species.
A paternal parent role that inheres in a grapevine genotype specifically by virtue of its function as the pollen source in controlled crosses, emphasizing the pollen production and donation aspects of male parental contribution. The pollen parent designation highlights the practical breeding function—the vine from which pollen is collected for application to emasculated seed parent flowers—focusing on pollen characteristics that influence crossing success. Pollen parent selection considers both genetic objectives (desired trait contributions) and practical pollen production characteristics: anther pollen quantity, pollen viability at collection and after storage, compatibility with intended seed parents, and timing of pollen availability relative to seed parent flowering. Some genotypes excel as pollen parents due to abundant, viable, storable pollen, while others with equally valuable genetics are difficult to use paternally. Pollen collection techniques, storage conditions, and application methods have been refined to maximize fertilization success, with properly handled grapevine pollen retaining viability for months to years under appropriate storage.
Pollen_Parent_Role emphasizes the practical pollen production function within the broader paternal parent category, recognizing that breeding program logistics depend substantially on pollen availability and quality. The pollen parent's role is distinct from seed parent in that pollen can be collected, processed, stored, and transported—enabling temporal and spatial separation between parents that is impossible for seed parents. International pollen exchange has facilitated germplasm access, with breeding programs receiving pollen from genotypes unavailable as live plants. Pollen storage technology (refrigeration, desiccation, cryopreservation) extends pollen viability from days (fresh pollen) to years (properly prepared and stored), expanding crossing options beyond simultaneous flowering periods. Documentation of pollen parent identity is essential for progeny records and genetic analysis.
Pollenelter
Pollenquelle
male crossing partner
männlicher Kreuzungspartner
pollen donor
pollen source
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Olmo HP (1976) Grapes, in Simmonds NW (ed) Evolution of Crop Plants, Longman pp.: 294-298 Ramming DW (2008) Seedless grape breeding, Blackwell
Pollen Parent Role
Merlot in Bordeaux shows pollen release during early-mid June Year N+1 at BBCH 60-65 over 8-10 days, with anther dehiscence progressing throughout cluster releasing abundant pollen estimated 25,000-45,000 grains per flower, pollen viability 76-88% fresh declining to 35-52% after 24 hours, visible pollen clouds during favorable morning conditions, under oceanic climate of 19-25°C temperature, 54-68% relative humidity, light winds supporting efficient dispersal and pollination enabling 54-64% fruit set typical of this mid-season cultivar
Pollen release represents the male reproductive process encompassing anther dehiscence, pollen liberation, and dispersal mechanisms where mature pollen grains exit anthers through specialized opening structures and distribute through wind transport to receptive stigmas, establishing male gamete availability enabling pollination and fertilization. This essential reproductive process occurs during flowering in early-mid June in temperate regions Year N+1, typically spanning 5-10 days from first anther dehiscence through completion of pollen discharge across cluster corresponding to BBCH 60-65 stages parallel to anthesis and cap fall. Pollen maturation preceding release shows final developmental stages with pollen grains completing tricellular state containing vegetative cell and two sperm cells ready for fertilization, accumulation of storage reserves including starch and lipids providing energy for germination and tube growth, and cytoplasmic organization preparing for immediate germination capacity upon stigma contact. Anther structure enabling release includes bilocular organization with two pollen sacs per anther separated by connective tissue, locular walls surrounding pollen-containing chambers providing mechanical protection during development, and specialized stomium zones along anther length representing predetermined rupture sites enabling controlled pollen release. Dehiscence mechanism involves coordinated processes including anther wall structural changes preparing opening, stomium zone activation triggering rupture, and mechanical forces driving pollen expulsion from locules creating liberation. Anther wall rupture proceeds through stomium zone breakdown with specialized cells undergoing programmed degeneration, cell wall dissolution creating longitudinal slits along anther length, and coordinated tissue failure producing clean opening enabling pollen access to exterior. Environmental triggering of dehiscence shows strong humidity dependence with declining humidity causing anther wall dehydration creating mechanical tension driving stomium opening, typically occurring during morning hours as temperature rises and humidity decreases, and repeated daily cycles creating characteristic diurnal pollen release pattern. Temperature effects on release show optimal 22-30°C supporting rapid efficient dehiscence and pollen liberation, cool conditions below 18°C substantially delaying dehiscence through slowed tissue processes, and warm conditions above 33°C potentially affecting pollen viability though generally facilitating mechanical opening. Pollen production per flower shows substantial investment with typical flower producing 20,000-60,000 pollen grains from five anthers representing 4,000-12,000 grains per anther, and cluster-level production reaching millions of grains creating abundant male gamete availability supporting effective pollination. Pollen characteristics affecting dispersal include small spheroidal grains 25-35 micrometers diameter enabling wind transport, relatively low density facilitating airborne suspension, and three-colporate aperture pattern providing germination sites on stigma contact. Pollen liberation from anthers proceeds through multiple mechanisms including initial explosive discharge when dehiscence occurs releasing pollen bolus, continued gravitational release as loosened pollen falls from opened locules, and wind-induced agitation dislodging remaining pollen creating extended release period. Pollen dispersal shows primarily wind-assisted mechanism with light grains easily becoming airborne through air currents, traveling distances from centimeters within flower to meters between vines depending on wind conditions, and creating characteristic pollen clouds visible surrounding flowering clusters during favorable dry conditions. Atmospheric pollen concentration during active release reaches maximum levels with estimated 100-1000 grains per cubic centimeter in immediate cluster vicinity under favorable conditions, declining with distance but remaining elevated 10-100 grains per cubic centimeter at vine scale, and providing abundant pollen availability for pollination. Pollen capture on stigmas occurs through wind transport delivering airborne grains to receptive stigma surfaces, adhesive stigma exudate trapping contacted grains, and physical stigma structure with papillate cells providing enhanced capture efficiency. Pollen viability timing shows maximum immediately upon release from freshly dehisced anthers with 70-90% germination capacity under optimal laboratory conditions, progressive decline over hours to days with viability reducing to 50-70% after 12 hours, 30-50% after 24 hours, and 10-30% after 48 hours, creating critical brief window for effective pollination. Environmental effects on viability include temperature with optimal 20-28°C maintaining viability while excessive heat above 35°C causes rapid loss through protein denaturation, humidity with moderate 40-70% maintaining hydration without promoting fungal growth, and rain catastrophically destroying viability through excessive hydration and physical washing. Self-pollination predominance in grapevine shows primary reproductive mode with majority of successful pollinations occurring within individual flowers through close anther-stigma proximity, supplemented by neighboring flower pollination within same cluster, and limited cross-pollination between vines playing minor role. Wind-assisted pollination efficiency varies with light-moderate wind 5-15 km per hour optimal for pollen dispersal enhancing pollination, calm conditions reducing dispersal though self-pollination remains effective, and strong wind above 25 km per hour potentially dispersing pollen excessively or causing desiccation. Pollen-stigma interactions following successful delivery show pollen grain hydration from stigma exudate within 15-90 minutes enabling metabolic activation, pollen recognition through molecular signals between coat proteins and stigma components, and germination initiation producing tube emergence within 1-4 hours under favorable conditions. Pollen tube growth following germination proceeds at 1-3 mm per hour through style transmitting tract, requiring 12-36 hours to traverse typical 15-30 mm style length, and delivering sperm cells to ovules enabling fertilization. Double fertilization following successful pollen tube arrival shows one sperm cell fusing with egg forming diploid zygote initiating embryo development, and other sperm fusing with polar nuclei forming triploid endosperm nucleus supporting seed development. Fruit set dependence on pollen release shows direct relationship between pollen availability, viability, and pollination success determining berry number, with favorable pollen release conditions supporting 50-70% fruit set in good-setting cultivars while impaired release reduces set to 20-40% or less. The process shows two major phases progressing from anther dehiscence establishing pollen liberation mechanism, through active pollen shedding creating maximum dispersal and pollination opportunity. Duration typically spans 5-10 days from first dehiscence through completion depending on flowering synchrony and environmental conditions, with warm stable conditions promoting compressed period while variable conditions extend duration. Cultivar variation appears in pollen production quantity affecting pollination potential, dehiscence timing and efficiency affecting release success, and pollen viability characteristics affecting reproductive outcomes. Regional timing shows pollen release during late May-early June in warm climates, early-mid June in temperate regions, and mid-late June in cool areas, coinciding with anthesis period and serving as critical male reproductive contribution. Viticultural significance includes representing essential male reproductive function determining pollination success and fruit set, requiring optimal environmental conditions particularly temperature and humidity for efficient release and viability maintenance, demonstrating extreme weather vulnerability with rain catastrophically affecting pollen function, and establishing male contribution as potential yield limitation in unfavorable conditions. Research applications include analyzing pollen production, maturation, and release mechanisms, investigating environmental effects on dehiscence and pollen viability affecting reproductive success, evaluating pollen dispersal patterns and pollination efficiency, quantifying relationships between pollen characteristics and fruit set outcomes, and assessing genetic variation in pollen function affecting reproductive reliability.
Pollen_Release represents male reproductive process parallel to Cap_Fall and integral to Anthesis within Flowering. Corresponds to BBCH 60-65 stages. Subdivided into Anther_Dehiscence establishing liberation mechanism and Active_Pollen_Shedding creating maximum dispersal. Environmental conditions particularly humidity and temperature critically affect dehiscence efficiency and pollen viability. Essential male contribution determining pollination success and fruit set. Extreme weather vulnerability particularly to rain destroying pollen function.
Pollenabgabe
Pollenfreisetzung
Pollenliberation
male gamete release
männliche Gametenfreigabe
pollen discharge
pollen dispersal
pollen liberation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 375-385 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 287-295 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Pollen Release
Chardonnay anthers at BBCH 57-58 (pollen maturation phase) contain four active pollen sacs (two per lobe) each filled with 1,000-1,600 maturing pollen grains, visible under magnification as densely packed yellow granular material within sac cavities, with tapetum cells at advanced degeneration stage and partition walls between paired sacs beginning breakdown in preparation for septum collapse by BBCH 59-60. Merlot anthers at BBCH 60-61 (early anthesis) show pollen sacs with septa fully broken down creating one locule per lobe (two locules total per anther) containing 3,500-6,500 mature pollen grains ready for release through imminent longitudinal dehiscence. Cabernet Sauvignon experiencing heat stress (38-41°C) during BBCH 55-56 (microsporogenesis) shows severely disrupted pollen sac development with many pollen mother cells failing meiosis, producing anthers at BBCH 59-60 containing only 1,200-2,800 pollen grains per anther (versus normal 3,500-7,000) with 40-55% aborted or malformed pollen visible as shriveled brown grains among normal yellow grains, resulting in dramatically reduced fruit set (28-38% versus normal 50-58%) demonstrating critical dependence of fruit production on successful pollen sac function during heat-sensitive meiotic stages.
A pollen sac (microsporangium) is one of the four pollen-producing compartments within the anther, organized as two pairs of sacs with one pair per anther lobe (theca), functioning as the site where microsporogenesis (meiotic production of haploid microspores from diploid pollen mother cells) and pollen maturation (development of microspores into mature pollen grains) occur, culminating in pollen release through anther dehiscence when the partition walls between adjacent sacs in each lobe break down creating a single locule per lobe. The pollen sac structure comprises= (1) a central cavity (locule) containing developing or mature pollen grains, with cavity dimensions typically 0.15-0.4 mm length and 0.08-0.2 mm width before dehiscence (2) a multi-layered wall surrounding the cavity including epidermis (outer protective layer), endothecium (fibrous layer with thickened cell walls creating hygroscopic mechanism for dehiscence), middle layers (transient nutritive layers that degenerate during development), and tapetum (innermost nutritive layer providing nutrients and wall precursors to developing pollen, degenerating as pollen matures) (3) sporogenous tissue (pollen mother cells) in early developmental stages (BBCH 54-55) that undergoes meiosis to produce microspore tetrads (4) microspore tetrads and developing pollen grains in mid-stages (BBCH 55-58) (5) mature pollen grains in late pre-dehiscence and dehiscence stages (BBCH 59-65). Pollen sac development follows a characteristic sequence= pollen sac initiation (BBCH 54) involves differentiation of sporogenous tissue from anther primordium cells microsporogenesis (BBCH 55-56) involves meiotic divisions of pollen mother cells producing tetrads of haploid microspores with each sac containing 200-2,000 pollen mother cells producing 800-8,000 microspores microspore separation (BBCH 56-57) involves tetrad breakdown releasing individual microspores pollen maturation (BBCH 57-59) involves microspore development into mature pollen grains with characteristic tricolporate morphology, accumulation of starch and lipid reserves, and formation of generative and vegetative nuclei septum breakdown (BBCH 59-60) involves degeneration of the partition wall separating the two adjacent pollen sacs within each anther lobe, creating a single locule per lobe containing pollen from both original sacs pollen release (BBCH 61-65) through longitudinal dehiscence when stomium breakdown creates slits allowing pollen escape. Each of the four pollen sacs in a typical hermaphrodite flower anther produces approximately 750-1,900 mature pollen grains (varying with cultivar and growing conditions), with total anther pollen production of 3,000-7,500 grains, and total five-anther flower production of 15,000-37,500 grains (typical range 15,000-30,000 for most hermaphrodite cultivars). Pollen sac function is critically dependent on tapetum nutrition, with tapetal cells secreting proteins, lipids, and wall precursors that are absorbed by developing pollen grains, and with tapetal degeneration timing coordinated with pollen maturation such that tapetum breakdown completes just as pollen achieves maturity. Environmental stresses during pollen sac development can severely impair pollen production= heat stress >35°C during microsporogenesis (BBCH 55-56) causes meiotic failures producing aborted microspores and reducing pollen production by 40-80% cold stress <10°C during pollen maturation (BBCH 57-59) can impair pollen wall formation and reserve accumulation reducing viability water or nutrient stress can impair tapetal function reducing pollen nutrition and producing shriveled or non-viable pollen. The paired pollen sac organization (two sacs per lobe, four sacs per anther) is characteristic of Vitis and most angiosperms, with the paired arrangement within each lobe allowing efficient pollen accumulation in a single locule following septum breakdown, and with the two-lobe organization creating two dehiscence slits per anther maximizing pollen release efficiency.
Pollen_Sac represents the fundamental pollen-producing compartment within the Anther and must be distinguished from the Anther itself (which comprises four pollen sacs organized into two lobes) and from individual pollen grains (the product of pollen sac development). The pollen sac is the site of microsporogenesis and pollen maturation, with successful pollen sac development absolutely essential for male reproductive function and fruit set. Sub-classification into Pollen_Filled_Sac (containing mature pollen) and Empty_Pollen_Sac (post-dehiscence, pollen released) reflects the critical functional transition from pollen storage to pollen dispersal. Understanding pollen sac development and environmental sensitivities is essential for diagnosing reproductive failures and for optimizing cultural practices to ensure successful bloom and fruit set.
Mikrosporangium
Pollenkammer
Pollensack
microsporangium
pollen chamber
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 94-98 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 187-192 Ebadi, A. et al. (1996). Effect of short-term temperature and shading on fruit set. Australian Journal of Grape and Wine Research 2=2-9.
Pollen Sac
Syrah in Rhône Valley shows pollen-stigma contact during mid-June Year N+1 at BBCH 60-65, with pollen grains 28-32 micrometers diameter contacting papillate stigma cells 45-65 micrometers length through primarily gravitational descent from anthers 0.9-1.4 mm above stigma supplemented by short-range wind-assisted delivery, initial mechanical retention by cell morphology and exudate viscosity preventing grain loss within milliseconds of contact, under Mediterranean conditions of 25-29°C temperature, typical of this mid-to-late season cultivar establishing 12-25 pollen contacts per stigma during peak period
Pollen-stigma contact represents the initial physical encounter between pollen grain and receptive stigma surface when airborne or directly transferred grain touches papillate epidermal cells, establishing first essential interaction enabling subsequent adhesion, recognition, and germination processes. This critical contact event occurs during pollen capture in early-mid June in temperate regions Year N+1, typically occurring within seconds of pollen arrival at stigma vicinity as part of BBCH 60-65 flowering stages. Contact establishment involves pollen grain trajectory intersecting with stigma surface, physical collision or gentle landing on papillate cell projections, and immediate preliminary retention preventing grain rebound or loss through mechanical and initial adhesive interactions. Contact geometry shows typical scenario with spheroidal pollen grain 25-35 micrometers diameter contacting elongated papillate cell tip or lateral surface, grain orientation variable depending on arrival trajectory, and contact area initially small point contact expanding as grain settles and adhesion develops. Pollen approach trajectories include vertical descent for gravitational self-pollination with pollen falling from anthers directly above contacting stigma from above, lateral approach for wind-dispersed pollen with grains traveling horizontally or at angles contacting stigma sides, and variable trajectories depending on air currents and pollen release dynamics. Contact velocity shows low-energy encounters with gravitational pollen having terminal velocity approximately 1-3 cm per second enabling gentle landing, wind-dispersed pollen showing variable velocities depending on wind speed though typically <10 cm per second, and overall low-impact contacts compatible with maintaining grain integrity and enabling adhesion. Papillate cell contact surface provides reception site with cells projecting 40-80 micrometers from stigma surface creating three-dimensional capture zone, cell tips and sides presenting contact areas coated with stigmatic exudate, and cellular architecture providing mechanical retention features preventing immediate grain loss. Initial mechanical retention shows immediate effects with papillate cell surface irregularities creating physical barriers preventing grain rolling or sliding, exudate viscosity providing immediate preliminary adhesion resisting grain detachment, and contact geometry creating stable positioning enabling adhesion development. Stigmatic exudate at contact interface provides immediate adhesive properties with proteinaceous fluid coating papillate cell surfaces, viscous consistency creating resistance to grain movement, and beginning hydration of pollen grain enabling recognition and attachment processes. Pollen grain surface at contact shows intact exine wall with sculptured surface patterns and apertures, pollen coat substances from tapetal origin affecting adhesion properties, and grain structure maintaining integrity enabling subsequent recognition and germination. Contact duration shows initial transient phase lasting milliseconds to seconds as grain physically contacts surface and settles, transitional phase over seconds to minutes as preliminary adhesion develops, and stabilization over minutes to hours as firm attachment establishes through exudate binding. Multiple contact possibilities show stigma potentially receiving numerous pollen grains creating multiple simultaneous or sequential contact events, each grain establishing independent contact and adhesion, and typical receptive stigma experiencing 10-50 contact events during peak pollination creating pollen surplus. Contact success factors include appropriate pollen arrival trajectory enabling physical encounter, receptive stigma state with active exudate production providing adhesive surface, adequate pollen viability maintaining grain integrity during transfer and contact, and favorable environmental conditions preventing interference with contact establishment. Failed contact scenarios include pollen missing stigma due to unfavorable trajectory or wind conditions, contacted pollen failing to retain due to inadequate exudate or mechanical retention, or damaged pollen unable to establish viable contact due to structural compromise. Environmental effects on contact include wind affecting pollen arrival trajectory and contact probability, humidity influencing exudate properties affecting initial adhesion, temperature affecting tissue properties and exudate consistency, and rain preventing contact through washing pollen away or removing adhesive exudate. Temperature at contact affects exudate viscosity and adhesive properties with optimal 22-28°C supporting appropriate consistency for capture, cool conditions potentially increasing viscosity affecting contact dynamics, and warm conditions potentially reducing viscosity though generally maintaining adequate adhesion capacity. Humidity at contact influences exudate water content and consistency with moderate 50-65% maintaining appropriate properties for adhesion, high humidity above 75% potentially diluting exudate reducing capture efficiency, and low humidity below 45% potentially concentrating exudate though may affect subsequent hydration. Contact observation requires microscopic examination with light microscopy at 100-400x magnification enabling pollen grain visualization on stigma surfaces, fluorescence techniques using aniline blue or other fluorochromes highlighting pollen and enabling clear observation, and scanning electron microscopy providing detailed ultrastructural views of pollen-stigma contact geometry. Contact as pollination initiation shows this first physical encounter representing critical starting point for reproductive success, successful contact enabling subsequent adhesion development, pollen hydration, recognition, and germination, and failed contact eliminating pollination opportunity for that particular pollen-stigma pair. The process represents first aspect of pollen capture emphasizing initial physical encounter, followed by pollen adhesion initiation as attachment begins developing. Duration represents very brief event occurring over milliseconds to seconds as physical contact establishes. Cultivar variation in contact characteristics may include stigma morphology affecting contact probability and geometry, exudate properties affecting initial retention, and flower architecture affecting pollen delivery trajectories. Regional timing shows pollen-stigma contact events occurring during flowering period late May-early June in warm climates, early-mid June in temperate regions, and mid-late June in cool areas, coinciding with pollen availability and stigma receptivity periods. Viticultural significance includes representing essential first encounter in pollination process, requiring optimal conditions avoiding rain preventing contact, demonstrating importance of coordinated pollen delivery and stigma receptivity, and establishing contact success as prerequisite for subsequent fertilization. Research applications include analyzing contact mechanics and physics, investigating stigma morphology effects on contact probability and retention, evaluating environmental effects on contact success, developing visualization techniques for observing contact events, and quantifying relationships between contact characteristics and pollination outcomes.
Pollen_Stigma_Contact is first aspect of Pollen_Capture, distinguished by initial physical encounter between grain and stigma surface. Represents critical first interaction in pollination. Very brief event milliseconds to seconds. Requires appropriate pollen trajectory and receptive stigma state. Mechanical and initial adhesive retention prevent immediate grain loss. Followed by Pollen_Adhesion_Initiation as attachment develops. Essential prerequisite for subsequent pollination processes. Multiple contact events typical per stigma.
Pollenkontakt
Pollenkornberührung
Pollenlandung
initial pollen-stigma encounter
initiale Pollen-Narben-Begegnung
pollen grain contact
pollen landing
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 378-380 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Pollen Stigma Contact
Pinot Noir in Burgundy shows pollen tube discharge during late June Year N+1 at BBCH 65-69, with tube apex rupturing in synergid releasing two sperm cells 26-38 hours post-pollination, double fertilization producing diploid zygotes initiating embryo development and triploid endosperm nuclei supporting seed growth, 1-3 seeds forming per berry, rapid berry expansion to 14-20 mm within 2 weeks from auxin production, under continental conditions of 22-28°C temperature, typical of this early-to-mid season cultivar achieving 60-70% fertilization success and fruit set
Pollen tube discharge represents the culminating fertilization event when tube apex ruptures in penetrated synergid releasing two sperm cells into embryo sac cytoplasm enabling double fertilization through sperm-egg fusion forming diploid zygote and sperm-central cell fusion forming triploid endosperm nucleus, completing sexual reproduction and initiating seed and berry development. This critical discharge and fusion event occurs during ovule penetration in mid-late June in temperate regions Year N+1, typically occurring 24-48 hours after initial pollination as fertilization culminates, corresponding to BBCH 65-69 fertilization completion stages. Discharge mechanism involves tube apex positioned in penetrated synergid cell undergoing wall weakening and rupture, explosive or gradual release of tube contents including two sperm cells, vegetative nucleus, and cytoplasm into embryo sac, sperm cell liberation enabling migration to fusion targets, and tube structural failure completing reproductive function. Tube apex rupture shows wall breakdown at tip through localized enzymatic degradation or mechanical failure, turgor pressure driving explosive discharge or controlled release depending on wall properties and pressure magnitude, and sudden breach creating opening for content expulsion. Discharge timing shows rupture typically occurring within minutes to hours of synergid penetration, 24-48 hours after initial pollination representing culmination of reproductive timeline, and precise timing possibly influenced by tube-synergid interactions triggering discharge though mechanisms poorly understood. Sperm cell release shows two genetically identical haploid cells expelling from ruptured tube into embryo sac cytoplasm, cells maintaining structural integrity during discharge, non-motile cells requiring passive transport or drift to fusion targets, and rapid positioning within embryo sac given close proximity to egg and central cell. Vegetative nucleus discharge shows degrading or non-functional nucleus expelling with sperm cells, nucleus having completed guidance role during tube growth, and nucleus playing no further reproductive role post-discharge. Tube cytoplasm discharge releases remaining contents into embryo sac including organelles, remaining nutrients, and cellular components, discharged material potentially providing nutritive support or signals though specific roles unknown, and overall tube depletion completing reproductive contribution. Post-discharge tube fate shows collapsed empty tube structure remaining in micropyle and style, degeneration proceeding over subsequent days to weeks, and non-functional tube representing completed reproductive pathway. Sperm cell fate after discharge shows one sperm cell migrating to egg cell enabling zygote formation, other sperm cell approaching central cell enabling endosperm formation, and differentiation determining which sperm fuses with which target possibly stochastic or regulated by unknown mechanisms. Sperm-egg fusion mechanism shows sperm cell contacting egg cell plasma membrane at micropylar pole, membrane fusion integrating sperm into egg cytoplasm through lipid bilayer merger, sperm nucleus migrating through egg cytoplasm toward egg nucleus, nuclear envelope breakdown and fusion combining haploid sperm and egg nuclei, formation of diploid 2n zygote nucleus containing 50% paternal and 50% maternal genetic contribution, and zygote activation transitioning from developmental arrest to active division. Egg activation following fusion shows calcium signaling waves triggering developmental program, metabolic activation initiating biosynthesis and cell division preparation, cortical reaction preventing polyspermy through membrane modifications blocking additional sperm, and commitment to embryo development with first division typically within 24-48 hours. Sperm-central cell fusion shows second sperm contacting large central cell in embryo sac center, membrane fusion integrating sperm into central cell cytoplasm, sperm nucleus fusing with central cell nucleus formed from polar nuclei fusion or fusing separately with two polar nuclei, formation of triploid 3n endosperm nucleus with 2 per 1 maternal per paternal genome ratio, and endosperm development initiating providing nutritive tissue for embryo. Endosperm activation shows free nuclear division initiating rapidly producing multinucleate coenocyte, cellularization phase forming cellular tissue, and endosperm growth providing nutritive support enabling embryo development during seed maturation. Double fertilization coordination shows two fusion events occurring synchronously or within hours ensuring coordinated embryo and endosperm development, temporal coordination critical for normal seed development, and synchronization possibly regulated through common signals or developmental programs. Zygote first division shows initial cell division typically 24-48 hours post-fertilization, asymmetric division producing small terminal cell developing into embryo proper and large basal cell forming suspensor, and progressive divisions establishing embryo structure during seed development. Fertilization completion confirmation shows successful zygote and endosperm formation indicating reproductive success, ovule development into seed with integuments forming seed coat, ovary expansion initiating berry development, and visible berry growth providing evidence of fertilization 1-2 weeks post-pollination. Berry development initiation shows auxin production from developing seeds preventing flower abscission, hormone signaling triggering ovary wall growth creating berry flesh, cell division intensity maximizing during early post-fertilization period with 10-15 hour cycle times, rapid berry expansion from 4-8 mm pre-fertilization to 15-25 mm within 2-3 weeks, and resource allocation shifting dramatically with developing berries consuming 40-60% total photosynthate. Seed development shows integument layers developing into seed coat, embryo growing within endosperm consuming nutritive reserves, seed maturation coordinating with berry ripening over 90-120 days, and final seed number typically 1-3 per berry depending on fertilization success in multiple ovules. Failed discharge outcomes include tube structural failure before synergid penetration preventing discharge, sperm cell damage during discharge compromising viability, failed sperm-egg or sperm-central cell fusion preventing zygote or endosperm formation, abnormal embryo sac preventing normal gamete reception, and overall fertilization failure causing flower abortion and coulure poor fruit set. Environmental effects on discharge and fusion include temperature critically affecting membrane fusion and developmental activation with optimal 22-28°C supporting efficient processes, cool conditions 15-20°C potentially impairing fusion or subsequent development, excessive heat above 33°C causing reproductive dysfunction, and critical temperature sensitivity during 24-48 hour fertilization window. Polyspermy prevention shows single tube reception per ovule through unknown blocking mechanisms, rapid ovule signaling changes after fertilization preventing additional tubes, and protection ensuring normal diploid embryo and triploid endosperm avoiding abnormal polyploid development. Multiple ovule fertilization shows 2-4 ovules per flower potentially receiving tubes, typical 1-3 successful fertilizations creating 1-3 seeds per berry, and seed number affecting berry size with more seeds generally producing larger berries through greater auxin production. The process represents second and final aspect of ovule penetration completing fertilization, following egg cell approach. Duration represents culminating moments with discharge occurring rapidly and fusion completing within hours. Cultivar variation in tube discharge may include discharge efficiency, fusion success rates, and seed number outcomes affecting berry characteristics. Regional timing shows pollen tube discharge during mid-late June in warm climates, late June in temperate regions, and late June-early July in cool areas, completing 24-48 hours after pollination. Viticultural significance includes representing culminating reproductive event determining seed and berry formation, requiring successful completion of entire pollination-germination-growth-penetration sequence, demonstrating extreme environmental sensitivity during fusion period, establishing discharge and fusion success as direct determinants of fruit set and yield, and marking definitive transition from flowering reproductive phase to berry development phase. Research applications include analyzing discharge mechanisms and triggers, investigating double fertilization coordination and sperm fate determination, evaluating fusion mechanisms and gamete recognition, quantifying fertilization success rates and seed number outcomes, assessing fertilization effects on berry development and quality, and developing strategies for improving fruit set through enhanced fertilization efficiency.
Pollen_Tube_Discharge is second and final aspect of Ovule_Penetration, distinguished by sperm release and fusion events completing fertilization. Represents culminating reproductive moment creating zygote and endosperm. Occurs 24-48 hours post-pollination. Involves double fertilization producing 2n embryo and 3n endosperm. Initiates seed and berry development through auxin production. Success rate 50-70% in favorable conditions determining fruit set. Completes entire Flowering hierarchical sequence from Inflorescence_Development through Fertilization. Marks definitive transition to Berry_Development phase. Final reproductive process.
Doppelbefruchtungsereignis
Pollenschlauch-Entladung
Schlauchruptur
Spermazell-Freisetzung
double fertilization event
gamete fusion completion
sperm cell release
tube rupture
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 393-400 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 298-305 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Pollen Tube Discharge
Cabernet Sauvignon in Napa Valley shows pollen tube emergence during early-mid June Year N+1 at BBCH 60-65, with tubes breaking through activated apertures 1.5-3.5 hours after pollen-stigma contact, initial protrusions 3-6 micrometers diameter rapidly extending to 8-11 micrometer cylindrical tubes growing at 1.2-2.8 mm per hour, under warm climate conditions of 24-29°C temperature, adequate boron 14-20 ppm supporting normal wall synthesis and tube integrity, typical of this late-season cultivar with 65-80% emergence success from viable hydrated pollen
Pollen tube emergence represents the definitive germination event when vegetative cell extends through activated aperture creating visible cylindrical protrusion breaking through grain wall, establishing actively growing pollen tube that will navigate through stigma and style tissues delivering sperm cells to ovules for fertilization. This critical emergence event occurs during pollen germination in early-mid June in temperate regions Year N+1, typically occurring 1-4 hours after pollen-stigma contact as germination culminates, corresponding to BBCH 60-65 flowering stages. Emergence mechanism involves turgor-driven extension with accumulated osmotic pressure 0.5-1.5 MPa pushing vegetative cell cytoplasm through weakened aperture, tip-focused growth with cell wall synthesis concentrated at emerging tube apex, breakthrough of modified intine layer creating initial protrusion 2-5 micrometers diameter, and rapid elongation establishing cylindrical tube structure extending from grain. Initial protrusion characteristics show small bulge appearing at activated aperture representing beginning tube emergence, protrusion 2-5 micrometers diameter initially expanding as breakthrough proceeds, and transition from bulge to distinct cylindrical tube as emergence completes and growth establishes. Breakthrough dynamics show turgor pressure overcoming residual wall resistance at aperture, intine layer yielding allowing cytoplasm extrusion, initial emergence typically occurring within minutes once activation completes, and irreversible commitment to tube growth once breakthrough achieved. Tube morphology at emergence shows cylindrical structure with initial diameter 5-8 micrometers expanding to typical 8-12 micrometers in actively growing tube, tip-focused growth with rounded apex where wall synthesis and extension occur, and smooth tube surface with continuous wall formed by secreted precursors. Tube wall structure includes inner callose layer providing structural support and possibly controlling tube permeability and growth regulation, outer pectin-cellulose layer providing flexible wall enabling expansion at tip while maintaining integrity, and continuous wall synthesis at apex through Golgi-derived secretory vesicles delivering wall components. Wall synthesis at tube tip involves secretory vesicles containing wall precursors fusing with plasma membrane at apex, pectin, cellulose, and callose polymerizing forming new wall, and coordinated synthesis enabling tip extension at 1-3 mm per hour initially with variation depending on conditions and tube age. Cytoplasmic organization in emerging tube shows vegetative cell contents occupying tube with cytoplasm streaming basipetally from grain into elongating tube, organelles including mitochondria, ER, and Golgi distributed throughout providing metabolic support, generative cell or two sperm cells depending on mitotic completion traveling in tube cytoplasm, and nucleus initially remaining in grain later moving into tube. Tip growth mechanism shows polarized secretion concentrated at tube apex, vesicle fusion delivering membrane and wall components specifically to growing tip, calcium gradient with elevated calcium at apex possibly regulating vesicle fusion and wall synthesis, and cytoskeleton organizing vesicle trafficking and maintaining polarized growth. Growth rate at emergence shows initial slower extension 0.5-1.5 mm per hour during establishment phase, acceleration to typical 1-3 mm per hour once active growth establishes, and rate variation depending on temperature, boron availability, pollen vigor, and stigma support. Direction at emergence shows tube typically extending into stigma tissue penetrating papillate cell surface, initial growth direction possibly influenced by stigma surface contours and chemical gradients, and orientation establishing pathway toward style and ultimately ovary. Cytoplasmic streaming in tube shows characteristic basipetal flow from grain through tube toward apex, streaming rate 2-8 micrometers per second maintaining nutrient and organelle supply to growing tip, and streaming driven by actomyosin motor system supporting sustained growth. Nucleus behavior shows initially remaining in grain during early emergence, later migration into tube as growth proceeds, and eventual positioning in tube cytoplasm though generative/sperm cells more important for fertilization. Sperm cell delivery shows two sperm cells resulting from generative cell division traveling in tube cytoplasm, cells moving toward apex as tube grows, and ultimate delivery to embryo sac enabling double fertilization once ovary reached. Emergence timing variation shows rapid emergence 1-2 hours post-adhesion with vigorous fresh pollen under optimal conditions 25-28°C, delayed emergence 3-6 hours with aged pollen or suboptimal conditions 18-22°C, and failed emergence from inadequate hydration, activation failure, or pollen dysfunction. Environmental requirements for emergence include adequate temperature 20-28°C optimal supporting metabolic activity and wall synthesis, sufficient boron essential for wall structure with deficiency causing emergence failure or tube bursting, appropriate humidity maintaining stigma exudate and pollen hydration, and supportive stigma environment providing growth cues. Temperature effects on emergence show warm conditions promoting rapid emergence and vigorous growth, cool conditions substantially delaying emergence and reducing growth rate, and temperature affecting enzyme activity, vesicle trafficking, and overall metabolic support for growth. Boron critical importance shows absolute requirement with boron essential for pectin cross-linking in tube wall, deficiency causing weak wall leading to tube bursting during emergence or early growth, and optimal boron 10-20 ppm in tissues supporting normal tube integrity and growth. Stigma support during emergence includes continued exudate secretion possibly providing nutrients or growth factors, recognition signals potentially enhancing growth support for compatible pollen, and tissue providing pathway for initial tube penetration into transmitting tract. Multiple tube emergence per stigma shows 10-50 pollen grains potentially germinating creating multiple tubes, competitive growth with fastest-growing or most vigorous tubes having advantage in reaching ovary, and typically 1-2 tubes successfully completing growth and achieving fertilization despite numerous emergence events. Failed emergence causes include incomplete hydration or activation preventing breakthrough, boron deficiency causing wall synthesis failure, pollen viability loss preventing metabolic support, incompatibility response limiting stigma support though rare in grapevine, extreme environmental conditions disrupting growth processes, or developmental abnormalities affecting pollen structure or function. The process represents third and final aspect of pollen germination marking transition to tube growth phase, following pollen hydration and germination pore activation. Duration represents relatively brief event with visible emergence occurring over minutes to hour once activation completes. Cultivar variation in emergence may include pollen vigor affecting emergence speed, stored reserves affecting growth capacity, and tube growth characteristics affecting penetration and navigation success. Regional timing shows pollen tube emergence occurring during flowering period across all climates 1-6 hours following successful pollination as germination culminates. Viticultural significance includes representing definitive germination event initiating tube growth, requiring favorable conditions particularly adequate temperature and boron nutrition, demonstrating critical transition from dormant pollen to actively growing tube, and establishing emergence success as prerequisite for tube navigation to ovary and fertilization. Research applications include analyzing emergence mechanisms including turgor generation and tip growth, investigating environmental and nutritional effects on emergence success particularly boron requirements, evaluating tube wall synthesis and structure, quantifying emergence timing and growth initiation dynamics, and assessing emergence success as determinant of fertilization outcomes and fruit set.
Pollen_Tube_Emergence is third and final aspect of Pollen_Germination, distinguished by definitive breakthrough event creating growing tube. Represents culmination of hydration and pore activation processes. Duration minutes to hour for emergence event itself occurring 1-4 hours post-adhesion total. Critical boron dependence for wall integrity. Marks transition from Pollen_Germination to Pollen_Tube_Growth. Essential definitive event establishing active reproductive structure capable of delivering sperm cells to ovules.
Keimungsabschluss
Pollenschlauch-Emergenz
Schlauch-Durchbruch
Schlauchextrusion
germination completion
tube breakthrough
tube extrusion
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 384-388 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 289-292 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Pollen Tube Emergence
Pinot Noir in Burgundy shows pollen tube growth during early-mid June Year N+1 at BBCH 60-65, with tubes elongating through stigma into 18-24 mm style transmitting tissue at 1.4-2.6 mm per hour, requiring 14-28 hours to traverse style reaching ovary region, under continental conditions of 20-26°C temperature, adequate boron nutrition 12-18 ppm supporting normal wall synthesis and sustained growth, typical of this early-to-mid season cultivar with 2-4 tubes successfully reaching ovary per flower enabling fertilization and 60-70% fruit set
Pollen tube growth represents the essential post-germination process where emerged pollen tube elongates through stigma papillate cells, penetrates into style transmitting tissue, navigates through style to ovary, and approaches ovules delivering sperm cells enabling fertilization and fruit set. This critical growth and navigation process occurs during flowering in early-mid June in temperate regions Year N+1, typically spanning 12-36 hours from tube emergence to ovary arrival depending on style length and environmental conditions, corresponding to BBCH 60-65 flowering stages. Growth mechanism involves tip-focused extension with cell wall synthesis and expansion concentrated at tube apex, cytoplasmic streaming maintaining nutrient and organelle supply from grain through tube to growing tip, turgor pressure driving tip expansion with osmotic pressure 0.3-0.8 MPa pushing against yielding wall at apex, and coordinated metabolic processes supporting sustained elongation at 1-3 mm per hour. Tube elongation characteristics show continuous growth from stigma surface through style to ovary spanning typical 15-30 mm total distance in grapevine, growth rate 1-3 mm per hour with variation depending on tube age, temperature, boron availability, and pollen vigor, and total growth duration 12-36 hours from emergence to ovary arrival with faster growth at optimal 24-28°C and slower growth at suboptimal 18-22°C temperature. Tip growth mechanism shows polarized secretion concentrated at tube apex with Golgi-derived secretory vesicles delivering wall precursors, vesicle fusion with plasma membrane at tip releasing wall components for polymerization, localized wall synthesis creating new wall at apex while older wall regions remain static, and calcium gradient with elevated 1-10 micromolar calcium at tip possibly regulating vesicle fusion and wall extensibility. Tube wall synthesis involves continuous production of wall components including pectin polymers providing flexible matrix, cellulose microfibrils providing structural strength, callose inner layer possibly regulating permeability and providing support, and coordinated polymer deposition creating functional wall enabling expansion while maintaining integrity. Cytoplasmic streaming dynamics show basipetal flow from pollen grain through tube toward apex at velocity 2-8 micrometers per second, actomyosin motor system driving streaming through interaction between actin filaments and myosin motors, streaming maintaining nutrient supply, organelle distribution, and removal of used materials, and essential for supporting sustained growth over many hours and centimeters distance. Sperm cell transport shows two sperm cells resulting from generative cell mitosis traveling in tube cytoplasm, cells moving toward apex carried by cytoplasmic streaming, and ultimate positioning near tip enabling delivery to embryo sac once ovary reached. Nucleus behavior during growth shows vegetative cell nucleus migrating from grain into tube during early growth, nucleus traveling in tube though typically remaining behind sperm cells, and nuclear positioning possibly involved in growth regulation though mechanisms unclear. Style penetration phase shows tube breaking through stigma papillate cell surface typically between adjacent cells, entering extracellular matrix in style transmitting tissue, and establishing growth pathway through specialized conducting tissue toward ovary. Transmitting tissue navigation involves tube growing through extracellular matrix between style cells, following chemical guidance cues possibly including calcium gradients and signaling molecules, utilizing nutritional support from transmitting tissue cells secreting nutrients into extracellular space, and maintaining directed growth toward ovary over 15-25 mm typical style length. Guidance mechanisms include chemotropism with tube responding to chemical gradients possibly including calcium, GABA, or other attractants, thigmotropism with tube responding to physical cues in tissue architecture, and intrinsic polarity possibly contributing to directed growth, though specific mechanisms in grapevine poorly characterized. Transmitting tissue support shows specialized style cells secreting nutrients, growth factors, and possibly guidance signals into extracellular space, lipid-rich secretions possibly providing energy source for growing tubes, and overall style environment optimized for supporting pollen tube growth enabling successful fertilization. Growth rate variation shows initial slower growth 0.8-1.5 mm per hour during stigma penetration and early style entry, acceleration to peak 2-4 mm per hour during mid-style growth, and possible slight deceleration 1-2 mm per hour during late growth approaching ovary, with overall average 1-3 mm per hour depending on conditions. Environmental effects on tube growth include temperature critically affecting growth rate with optimal 24-28°C supporting rapid 2-3 mm per hour growth, cool conditions 18-22°C substantially slowing growth to 0.8-1.5 mm per hour potentially requiring 36-48 hours for ovary arrival, warm conditions 28-32°C potentially accelerating growth though excessive heat above 33°C may impair function. Boron absolute requirement for tube growth emphasizes critical importance with boron essential for pectin cross-linking in tube wall structure, deficiency causing tube bursting, irregular growth, or growth cessation preventing ovary arrival, optimal boron concentration 10-20 ppm in style tissues supporting normal growth and integrity, and boron deficiency even with adequate flower appearance severely reducing fruit set through failed tube growth. Calcium role in tube growth includes calcium gradient at tip possibly regulating vesicle fusion and wall extensibility, calcium signaling potentially involved in growth responses and guidance, and calcium nutrition affecting tube function though boron more critically limiting in practice. Resource requirements for sustained growth include carbohydrate reserves from pollen grain initially supporting early growth, transitional period utilizing both pollen reserves and style-derived nutrients, and later growth depending substantially on style nutritional support particularly during final growth to ovary. Style length effects show shorter styles 12-18 mm enabling more rapid ovary arrival within 8-20 hours, longer styles 22-30 mm requiring extended 20-40 hours for completion, and style length variation among flowers and cultivars affecting timing of fertilization events. Multiple tube growth per style shows 10-50 tubes potentially entering style from germinated pollen on stigma, competitive growth with fastest tubes having advantage in ovule access, typical 2-5 tubes reaching ovary region though usually only 1-2 achieving successful fertilization, and excess tubes providing reproductive assurance compensating for possible failures. Tube-tube interactions during growth possibly include physical interference if multiple tubes in close proximity, potential chemical signaling though poorly documented, and competitive dynamics affecting which tubes successfully fertilize. Ovary approach represents culminating growth phase with tube navigating to ovary region containing 2-4 ovules per carpel, tube responding to ovule-derived attractants directing final approach, and tube arrival at micropyle enabling sperm cell delivery to embryo sac for fertilization. Failed tube growth causes include boron deficiency preventing adequate wall synthesis, inadequate pollen reserves failing to support extended growth, environmental stress particularly temperature extremes disrupting metabolic support, style tissue dysfunction failing to provide adequate support or guidance, or pollen vigor limitations preventing sustained growth over required distance and duration. The process shows three distinguishable aspects progressing from style penetration entering transmitting tissue, through transmitting tissue navigation growing through style, to ovary approach culminating growth. Duration typically spans 12-36 hours from emergence to ovary arrival with faster 10-20 hours under optimal warm conditions and adequate boron, slower 24-48 hours under cool conditions or marginal nutrition. Cultivar variation in tube growth may include growth rate affecting fertilization timing, style length affecting required growth duration, and boron efficiency affecting growth under marginal nutrition. Regional timing shows pollen tube growth occurring during flowering period across all climates over 12-36 hours following germination, completing within 24-48 hours of initial pollination under favorable conditions. Viticultural significance includes representing essential process linking germination to fertilization, requiring adequate boron nutrition as critical limiting factor, demonstrating extreme temperature sensitivity affecting growth rate and success, and establishing tube growth success as direct determinant of fertilization achievement and fruit set. Research applications include analyzing tube growth mechanisms including tip growth dynamics and cytoplasmic streaming, investigating guidance mechanisms directing growth through style, evaluating boron and other nutritional effects on growth rate and success, quantifying environmental effects particularly temperature on growth dynamics, developing visualization techniques for observing tube growth in vivo, and assessing tube growth success as determinant of fertilization timing and fruit set outcomes.
Pollen_Tube_Growth represents essential post-germination navigation process following Pollen_Germination within Flowering. Subdivided into Style_Penetration entering tissue, Transmitting_Tissue_Navigation through style, and Ovary_Approach culminating growth. Duration 12-36 hours spanning stigma to ovary. Critical boron dependence as absolute requirement. Temperature strongly affects growth rate. Leads directly to Fertilization upon ovary arrival. Essential process determining fertilization success and fruit set.
Pollenschlauch-Navigation
Pollenschlauchwachstum
Schlauch-Elongation
pollen tube navigation
reproductive tube growth
reproduktives Schlauchwachstum
tube elongation
tube extension process
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 385-390 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 290-295 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Pollen Tube Growth
Pollen tube growth through the grapevine style requires 12-48 hours depending on cultivar and temperature, with tubes elongating at 15-40 µm/hour under optimal conditions. Cabernet Sauvignon pollen tubes reach the ovary within 18-24 hours at 25°C, while Muscat of Alexandria requires 36-48 hours due to longer style length. In vitro tube lengths after 24 hours range from 800-2,500 µm, with Pinot Noir achieving 2,000-2,500 µm and Grenache showing slower growth at 800-1,200 µm. Tube growth follows a chemotropic gradient guided by ovule-derived signals, with 75-90% of tubes successfully reaching ovules in compatible pollinations.
A pollen disposition that is the inherited capacity of a Vitis vinifera plant to produce pollen grains capable of extending pollen tubes through the stigma and style to the ovary, maintaining directional growth, structural integrity, and metabolic activity sufficient to deliver sperm cells for fertilization.
Pollen tube growth is the rate-limiting step in fertilization timing, particularly under suboptimal temperatures. Growth requires continuous tip-localized pectin secretion and callose plug formation at intervals along the tube. Self-incompatibility in grapevine is generally weak, but pollen tube growth rates may differ between self and cross pollen. Temperature below 15°C severely restricts tube growth, contributing to poor fruit set in cool flowering seasons.
Pollenschlauchverlängerung
Wachstumsdisposition des Pollenschlauchs
pollen tube elongation capacity
pollen tube extension disposition
tube growth rate trait
Staudt G (1982) Pollen germination and pollen tube growth in vivo in Vitis and the dependence on temperature. Vitis 21:205-216 Galet P (2000) General Viticulture, Oenoplurimedia, Chaintre pp.: 128-135
Pollen Tube Growth Disposition
Pinot Noir in Burgundy shows pollen tube penetration during mid-late June Year N+1 at BBCH 65-69, with tubes navigating through 120-180 micrometer micropyle canals over 1.5-3.5 hours at 0.4-0.7 mm per hour penetration rate, reaching nucellus surface and entering synergid cells, under continental conditions of 21-27°C temperature, adequate boron 12-18 ppm supporting tube wall integrity during constrained passage, typical of this early-to-mid season cultivar with 62-76% penetration success 22-34 hours post-pollination
Pollen tube penetration represents the micropyle entry phase when tube navigates through micropyle canal between integument layers accessing nucellus and embryo sac, establishing pathway enabling sperm cell delivery to female gametophyte for double fertilization. This critical penetration phase occurs during fertilization in mid-late June in temperate regions Year N+1, typically proceeding over 1-4 hours following tube arrival at micropyle as part of BBCH 65-69 fertilization stages. Penetration mechanism involves tube apex targeting micropyle outer opening guided by synergid-derived attractant signals, growing through micropyle canal navigating 100-300 micrometer tortuous passage between integument layers, reaching nucellus surface adjacent to embryo sac, and penetrating nucellus wall entering synergid cell completing access to female gametophyte. Micropyle canal structure shows passage formed by integument tissue layers, inner and outer integuments creating canal walls, canal diameter 20-50 micrometers varying along length providing constrained growth pathway, canal length 100-300 micrometers depending on integument thickness and ovule development, and tortuous non-linear path requiring precise tube navigation. Tube growth through micropyle shows slower penetration rate 0.3-0.8 mm per hour compared to 1-3 mm per hour style growth reflecting constrained environment, sustained tip growth maintaining wall synthesis and extension, tube diameter 8-12 micrometers fitting within 20-50 micrometer canal, and mechanical interaction with canal walls potentially affecting growth dynamics. Synergid attractant guidance shows filiform apparatus in synergid cells secreting signals creating concentration gradient, attractant molecules diffusing through micropyle establishing directional cue, possible calcium gradient contributing to guidance, and tube showing chemotropic response directing growth precisely toward inner micropyle opening adjacent to synergid. Tube-micropyle wall interactions include possible physical contact and adhesion, tube secreting wall-modifying enzymes potentially facilitating passage through constrained space, mechanical forces from turgor pressure contributing to penetration through resistant tissue, and coordinated growth enabling successful navigation despite spatial constraints. Nucellus penetration following micropyle transit shows tube reaching nucellus surface at inner micropyle opening, penetrating through nucellus wall cells, and entering synergid cell creating direct access to embryo sac cytoplasm. Synergid cell structure providing reception includes specialized cells flanking egg at micropylar pole, filiform apparatus with extensive wall ingrowths creating large surface area facing micropyle, degeneration program initiating upon tube contact preparing for discharge reception, and cytoplasmic organization supporting tube attraction and penetration. Tube penetration into synergid shows apex entering synergid cell cytoplasm through filiform apparatus region, penetration possibly causing synergid degeneration as part of normal fertilization process, and establishing pathway for sperm cell release into embryo sac. Penetration timing shows 1-4 hours required for micropyle navigation from outer opening to synergid entry, timing variation depending on micropyle length and structure, temperature affecting penetration rate, and culminating arrival at embryo sac typically 20-40 hours after initial pollination. Tube structural requirements during penetration include robust wall resisting mechanical stresses in constrained micropyle environment, continued wall synthesis supporting growth through canal, adequate turgor pressure driving extension through resistant tissue, and boron-dependent wall integrity preventing bursting under mechanical stress. Boron critical role during penetration emphasizes absolute requirement with boron essential for maintaining tube wall structure during micropyle passage, deficiency causing tube bursting or structural failure during penetration preventing embryo sac access, mechanical stress in constrained environment exacerbating boron deficiency effects, and adequate boron 10-20 ppm supporting successful penetration. Environmental effects on penetration include temperature affecting growth rate and metabolic support with optimal 22-28°C supporting efficient penetration within 1-2 hours, cool conditions 18-22°C slowing penetration requiring 3-5 hours, and continued boron requirement as limiting factor affecting penetration success. Failed penetration causes include inadequate tube vigor preventing sustained growth through micropyle, boron deficiency causing structural failure during passage, abnormal micropyle structure preventing access, insufficient attractant signaling from synergid dysfunction, or tube misdirection failing to locate micropyle opening. Penetration success rate shows 50-75% tubes reaching ovary region successfully penetrating micropyle and accessing embryo sac under favorable conditions, declining to 30-50% under suboptimal conditions, and success rate directly determining fertilization potential and fruit set. The process shows two distinguishable aspects progressing from micropyle targeting achieving precise navigation to outer opening, through synergid penetration entering embryo sac cytoplasm. Duration typically spans 1-4 hours representing final tube growth phase before discharge. Cultivar variation in penetration may include micropyle structure affecting penetration difficulty, ovule development affecting access pathway, and tube vigor affecting penetration success rate. Regional timing shows pollen tube penetration occurring during mid-late June across all climates 20-40 hours after pollination as final tube growth phase before fertilization. Viticultural significance includes representing critical access phase enabling sperm delivery to egg, requiring sustained tube vigor and structural integrity, demonstrating continued boron importance as absolute requirement, and establishing penetration success as prerequisite for fertilization and fruit set. Research applications include analyzing micropyle penetration mechanisms and navigation, investigating synergid attractant signals and chemotropic guidance, evaluating boron effects on tube integrity during penetration, quantifying penetration timing and success rates, and assessing penetration success as determinant of fertilization outcomes.
Pollen_Tube_Penetration is first major Fertilization phase establishing embryo sac access. Subdivided into Micropyle_Targeting achieving precise navigation and Synergid_Penetration entering female gametophyte. Duration 1-4 hours for micropyle passage. Slower growth rate 0.3-0.8 mm per hour in constrained environment. Critical boron dependence for structural integrity. Synergid attractant guidance essential. Success rate 50-75% under favorable conditions. Enables subsequent tube discharge and fertilization.
Mikropyle-Durchdringung
Pollenschlauch-Penetration
Samenanlage-Zugang
Schlaucheintritt in Embryosack
micropyle penetration
ovule access
synergid entry
tube entry into embryo sac
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 390-392 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 295-298 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Pollen Tube Penetration
Pollen viability assessed by acetocarmine or Alexander staining ranges from 50-95% across V. vinifera cultivars, with Riesling consistently showing high viability (88-95%) and Gewürztraminer exhibiting lower values (65-80%). Environmental stress during microsporogenesis (meiosis, approximately 6-8 weeks pre-bloom) reduces viability, with temperatures above 35°C causing 30-50% reduction. Viability declines rapidly after anther dehiscence, with 50% loss within 24-48 hours under ambient conditions. Cold storage at 4°C extends viability to 5-7 days, enabling controlled pollination in breeding programs.
A pollen disposition that is the inherited capacity of a Vitis vinifera plant to produce pollen grains that maintain structural integrity, metabolic competence, and fertilization potential, as assessed by staining tests, in vitro germination, or fertilization success.
Pollen viability represents the proportion of pollen grains capable of successful fertilization under favorable conditions. Viability testing methods (vital staining, in vitro germination, fluorochromatic reaction) give correlated but not identical results due to different aspects of pollen competence assessed. Low viability contributes to poor fruit set and is exacerbated by nutritional deficiencies (boron, zinc) and fungicide applications during meiosis.
Pollenfertilität
Pollenlebensfähigkeit
Pollenviabilität
male fertility disposition
pollen fertility
pollen vitality trait
Kelen M, Demirtas I (2003) Pollen viability, germination capability and pollen production level of some grape varieties (Vitis vinifera L.). Acta Physiologiae Plantarum 25:229-233 Ebadi A et al. (1996) Effect of short-term temperature and shading on fruit-set, seed and berry development in model vines of V. vinifera cvs Chardonnay and Shiraz. Australian Journal of Grape and Wine Research 2:2-9
Pollen Viability Disposition
Merlot in Bordeaux shows pollination during early-mid June Year N+1 at BBCH 60-65 over 8-10 days, with successful pollen-stigma contacts occurring predominantly through self-pollination within flowers supplemented by neighboring flower wind-assisted transfer, stigma receptivity developing within 12-18 hours after flower opening and persisting 5-7 days, pollen adhesion proceeding within minutes to hours of contact, under oceanic conditions of 20-26°C temperature, 54-68% relative humidity supporting efficient pollination enabling 54-64% fruit set typical of this mid-season cultivar
Pollination represents the critical pollen-stigma interaction process encompassing pollen grain capture, adhesion, and recognition where airborne or directly transferred pollen contacts receptive stigma surfaces, adheres through exudate binding, and establishes conditions for subsequent germination and fertilization, determining fruit set success. This essential reproductive process occurs during flowering in early-mid June in temperate regions Year N+1, typically spanning 5-10 days from first pollen-stigma contacts through completion of pollination opportunities across cluster, corresponding to BBCH 60-65 stages parallel to anthesis and pollen release. Pollination mechanism in grapevine shows predominantly self-pollination mode with majority 60-80% successful pollinations occurring within individual flowers through close anther-stigma proximity facilitating direct pollen transfer, supplemented by wind-assisted neighboring flower pollination within same cluster contributing 20-35%, and limited cross-pollination between vines playing minor role <5-10% in reproductive success. Self-compatibility in Vitis vinifera enables effective reproduction without requiring genetically distinct pollen, eliminating obligate cross-pollination requirement found in some species, and supporting reliable fruit set through self-pollination though occasional outcrossing provides genetic diversity benefits. Pollen delivery mechanisms include gravitational transfer with pollen falling directly from dehiscing anthers onto stigma in same flower utilizing close 0.5-1.5 mm spatial separation, wind-assisted dispersal carrying airborne pollen to receptive stigmas within cluster or between neighboring vines, and occasional insect-mediated transfer though insects play minimal role in grapevine pollination compared to dominant wind-assisted self-pollination. Stigma receptivity enabling pollination shows critical female reproductive readiness with papillate surface cells elongating creating textured capture surface, proteinaceous exudate secretion producing adhesive coating trapping contacted pollen, and metabolic activation establishing capacity to support pollen hydration and germination, developing within 6-24 hours after flower opening and persisting 4-7 days though declining progressively. Stigma structure supporting pollination includes bilobed organization creating expanded surface area maximizing pollen capture probability, papillate epidermal cells with elongated projections 40-80 micrometers length providing mechanical retention structures, stigmatic exudate coating surfaces with adhesive proteinaceous fluid, and overall architecture optimized for efficient pollen capture and retention. Pollen-stigma contact represents initial pollination event with airborne or directly transferred pollen grain contacting stigma papillate surface, physical contact establishing opportunity for adhesion and recognition, and successful contact depending on pollen availability, stigma receptivity, and favorable environmental conditions. Pollen adhesion follows contact with stigmatic exudate providing adhesive binding trapping contacted grain on surface, molecular interactions between pollen coat proteins and stigma exudate components creating stable attachment, and physical retention by papillate cell structure preventing grain loss, establishing secure pollen-stigma association. Pollen recognition involves molecular signaling between pollen coat proteins and stigma exudate receptors enabling species recognition, compatibility assessment determining acceptance or rejection though grapevine shows self-compatibility, and triggering of downstream responses including pollen hydration and germination support if compatible. Pollen hydration on receptive stigma represents critical transition with grain absorbing water from stigmatic exudate within 15-90 minutes enabling metabolic activation, cell rehydration restoring physiological function after atmospheric desiccation, and establishing conditions for germination initiation within 1-4 hours under favorable conditions. Pollination efficiency in grapevine shows high success rates under favorable conditions with estimated 50-70% flowers achieving successful pollination leading to fruit set in good-setting cultivars, declining to 30-50% under suboptimal conditions or difficult-setting varieties, and demonstrating critical dependence on coordinated male-female reproductive function and favorable environmental conditions. Environmental effects on pollination show temperature affecting both pollen viability and stigma function with optimal 22-28°C supporting efficient pollination, humidity influencing pollen dispersibility and viability with moderate 50-65% optimal, and rain catastrophically disrupting pollination through washing pollen from surfaces, diluting stigmatic exudate, and physically damaging reproductive structures. Temperature effects on pollination include warm conditions supporting pollen viability maintenance, stigma metabolic activity, and efficient pollen-stigma interactions, cool conditions below 18°C substantially reducing pollination efficiency through impaired pollen and stigma function, and excessive heat above 33°C potentially causing pollen sterility or stigma dysfunction. Humidity effects show moderate levels optimal for maintaining pollen viability during dispersal and transfer, preserving stigmatic exudate adhesive properties, and supporting overall reproductive function, while high humidity above 75% potentially diluting exudate reducing adhesion, and low humidity below 45% causing pollen desiccation reducing viability. Rain effects represent catastrophic interference with water washing pollen from stigma surfaces eliminating captured grains, diluting or removing stigmatic exudate preventing adhesion, preventing pollen dispersal and transfer through elevated humidity and physical impedance, and promoting fungal infection on wet reproductive tissues, potentially reducing fruit set by 60-80% if occurring during peak pollination. Boron nutrition critically affects pollination success with adequate boron essential for stigma exudate production and pollen function, deficiency causing poor stigma receptivity and impaired pollen germination severely reducing fruit set even with adequate pollen availability, and emphasizing importance of sufficient boron supply during flowering period. Timing relationships show pollination requiring coordination between pollen availability and stigma receptivity with optimal overlap during peak anthesis creating maximum success opportunity, early or late pollination attempts showing reduced efficiency due to suboptimal pollen viability or stigma receptivity, and successful pollination typically occurring within 1-3 days of flower opening during receptivity peak. Fruit set dependence on pollination shows direct causal relationship with successful pollen-stigma interaction, adhesion, and recognition enabling subsequent germination and fertilization, while failed pollination due to pollen absence, stigma dysfunction, or unfavorable conditions causing flower abortion and fruit set failure, demonstrating pollination as critical limiting step determining berry number and yield. The process shows two major phases progressing from pollen capture establishing initial contact and retention, through pollen adhesion creating stable attachment enabling subsequent processes. Duration typically spans 5-10 days at cluster level as flowers sequentially open and achieve pollination, with individual pollination events occurring rapidly within hours once pollen contacts receptive stigma. Cultivar variation appears in pollination characteristics including self-compatibility levels affecting pollination success reliability, stigma receptivity characteristics affecting capture efficiency, and sensitivity to environmental conditions affecting pollination under marginal conditions. Regional timing shows pollination during late May-early June in warm climates, early-mid June in temperate regions, and mid-late June in cool areas, coinciding precisely with flowering period and representing critical reproductive window. Viticultural significance includes representing essential process linking male pollen production to female fertilization, requiring optimal environmental conditions particularly avoiding rain during pollination period, demonstrating extreme weather vulnerability with brief window determining season productivity, and establishing pollination success as primary determinant of fruit set and yield outcomes. Research applications include analyzing pollen-stigma interaction mechanisms including molecular recognition and adhesion processes, investigating environmental effects on pollination efficiency affecting reproductive success, evaluating pollen dispersal and capture dynamics, quantifying relationships between pollination characteristics and fruit set outcomes, and assessing management or genetic interventions affecting pollination success and productivity.
Pollination represents essential pollen-stigma interaction within Flowering linking male pollen release to subsequent germination and fertilization. Corresponds to BBCH 60-65 stages parallel to Anthesis and Pollen_Release. Subdivided into Pollen_Capture establishing contact and retention, and Pollen_Adhesion creating stable attachment. Predominantly self-pollination mode in grapevine. Critical dependence on coordinated male-female function and favorable environmental conditions. Extreme weather vulnerability particularly to rain. Essential determinant of fruit set and yield.
Befruchtungseinleitung
Bestäubung
Pollen-Narben-Interaktion
Pollentransfer
fertilization initiation
pollen transfer
pollen-stigma interaction
pollination process
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 375-385 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 285-295 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Pollination
Syrah in Rhône Valley shows pollination window closure during late June Year N+1 at late BBCH 68 over 1-2 days June 14-15, with negligible functional pollen remaining showing <5% viability, widespread stigma senescence eliminating receptivity, visible berry development 4-6 mm diameter from successful fruit set, unfertilized flowers showing brown senescence preparing for abscission, establishing final fruit set of 48-62% representing 320-480 berries per cluster, under Mediterranean conditions of 24-28°C temperature, typical of this mid-to-late season cultivar definitively concluding flowering and establishing season yield potential
Pollination window closure represents the functional termination of effective reproductive opportunity when combined effects of completed flowering, expired pollen viability, and declining stigma receptivity eliminate capacity for successful pollination, definitively establishing final fruit set and berry number for the season. This critical reproductive endpoint occurs during late anthesis in late June in temperate regions Year N+1, typically representing final 1-2 days of late anthesis period when reproductive function effectively ceases, corresponding to late BBCH 68 transitioning toward BBCH 69. Closure mechanisms involve multiple converging factors including exhaustion of pollen supply as all anthers complete dehiscence and aged pollen loses viability, stigma senescence eliminating receptive surfaces as exudate production ceases and tissues deteriorate, complete flower opening leaving no additional reproductive structures to contribute, and vine resource allocation shifting definitively toward fruit development precluding further reproductive investment. Pollen availability at closure shows negligible functional pollen remaining with final anthers having dehisced but pollen rapidly expiring, aged atmospheric pollen showing minimal 0-5% viability effectively non-functional, and no further pollen production forthcoming with all reproductive organs past functional stage. Stigma receptivity at closure shows widespread loss of function with earlier-opened stigmas completely senesced showing brown discoloration and desiccated tissues, recently opened stigmas showing declining receptivity 10-30% peak levels with minimal exudate production, and overall female reproductive capacity effectively terminated. Reproductive organ senescence accelerates during closure with unfertilized flowers showing rapid deterioration including anther withering and browning indicating complete pollen depletion, stigma necrosis eliminating receptive capacity, receptacle and pedicel beginning abscission zone activation preparing for flower drop, and overall flower senescence indicating failed reproduction. Successfully fertilized flowers at closure show contrasting appearance with visible berry development evident as small green structures 3-6 mm diameter, continuing expansion indicating active fruit development, hormonal signals preventing abscission maintaining flower-fruit attachment, and overall healthy appearance contrasting with senescing unfertilized flowers. Flower abscission initiation intensifies during closure with unfertilized flowers developing abscission zones at pedicel base involving enzymatic wall degradation creating separation plane, mechanical stresses from wind or handling potentially triggering detachment, and progressive flower drop creating visible accumulation of abscised flowers on cluster and ground surfaces. Fruit set determination becomes definitive at closure with count of successfully fertilized flowers establishing final berry number, typical fruit set percentages ranging 30-50% in poor conditions or difficult-setting cultivars to 60-75% under favorable conditions in good-setting varieties, and established berry number determining yield potential though subsequent berry development and maturation affecting final harvest outcomes. Developmental transition at closure shifts vine priority decisively from reproductive to developmental functions with metabolic resources redirecting from flowering processes to berry growth, hormonal environment dominated by auxin and gibberellins from developing berries rather than reproductive signals, and physiological focus on fruit development vegetative growth and reserve accumulation rather than continued reproduction. Cluster appearance at closure shows characteristic mixed state with small developing berries from successful early-mid fruit set, senescing brown unfertilized flowers preparing for abscission, occasional late-opening flowers with minimal reproductive potential, and overall appearance of reproductive conclusion with developmental phase beginning. Environmental effects on closure timing show that favorable conditions enabling efficient early reproduction allow relatively early closure with minimal late flowering activity, while unfavorable conditions extending flowering may delay closure though late reproductive success remains limited, and overall closure timing relatively consistent reflecting biological limits on reproductive duration. Temperature during closure typically ranges 20-28°C supporting normal physiological transition, with warm conditions potentially accelerating flower senescence and abscission, while cool conditions may slow processes though not preventing inevitable closure. Weather impacts on closure are minimal since reproductive outcomes already determined with adverse conditions affecting any late reproductive attempts but having negligible effect on overall fruit set already established. Management implications at closure include confirming flowering completion enabling resumption of normal vineyard operations, initiating post-bloom disease management particularly botrytis protection on developing berries, implementing canopy management operations including shoot positioning and possible thinning, assessing fruit set outcomes for yield predictions and crop adjustment decisions, and transitioning focus from reproductive to developmental management priorities. Phenological significance of closure includes marking definitive end of BBCH 60-69 flowering stages transitioning to BBCH 70-79 fruit development stages, establishing reference point for subsequent phenological predictions including berry development timing, and confirming reproductive phase completion enabling developmental phase focus. Disease management transition at closure shifts from bloom-period restrictions enabling fungicide applications for botrytis and powdery mildew protection on developing berries, recognizing that latent botrytis infections established during flowering will remain dormant requiring management during ripening, and implementing preventive strategies for berry diseases throughout development period. Nutritional management considerations at closure include ensuring adequate nitrogen for vegetative growth and fruit development without promoting excessive vigor, providing sufficient potassium for berry development and quality, maintaining adequate boron for cell division in expanding berries, and balanced nutrition supporting both fruit development and vine health through season. Resource allocation at closure shows vine transitioning from reproductive investment toward developmental demands with carbohydrates redirecting from flower support to rapid berry expansion, nitrogen partitioning to support vegetative growth and berry protein synthesis, and mineral nutrients focusing on fruit and shoot development. Hormonal environment at closure shows dominance of auxin from developing berries triggering cell division and expansion, gibberellins affecting early berry growth and possibly seedless berry development in some applications, cytokinins supporting cell division in berries and maintaining vegetative growth, and ethylene potentially involved in flower abscission though not directly affecting berry development at this early stage. Developmental outcomes established at closure include final berry number determining yield potential typically 100-500 berries per cluster depending on variety and conditions, within-cluster developmental gradient with 5-10 day spread between earliest and latest-set berries affecting ripening uniformity, and cluster architecture with berry spacing affecting disease pressure and quality characteristics. Fruit set assessment methods at closure include visual berry count on representative clusters providing direct berry number measurement, flower abscission evaluation estimating set percentage from remaining vs fallen flowers, and cluster weight estimates based on berry number and expected berry size predicting yield outcomes. The process represents third and final late anthesis aspect marking definitive reproductive termination, following final flowers opening and declining pollen viability. Duration represents 1-2 day transition period as reproductive function effectively ceases, with rapid progression toward complete anthesis. Cultivar variation in closure characteristics shows some varieties reaching definitive closure rapidly with clean transition to fruit development, while others show extended indeterminate closure with continued flower senescence over several days, affecting management timing and fruit set assessment clarity. Regional timing shows pollination window closure during late June in warm climates at approximately 1100-1150 growing degree days base 10°C, late June-early July in temperate regions at 1000-1050 GDD, and early July in cool areas at 900-1000 GDD, marking transition from reproductive to developmental phenological phase. Viticultural significance includes establishing definitive final fruit set and berry number determining yield potential, marking end of critical weather-sensitive flowering period reducing vulnerability to environmental conditions, enabling resumption of normal management operations including canopy management and disease protection, providing fruit set outcomes for yield prediction and crop adjustment planning, and confirming transition to developmental phase requiring different management priorities. Research applications include analyzing mechanisms coordinating reproductive termination and developmental initiation, evaluating factors determining fruit set percentage and berry number outcomes, quantifying relationships between flowering conditions and final fruit set results, assessing within-cluster developmental heterogeneity affecting ripening patterns, and documenting phenological transitions marking major developmental phase changes in perennial reproductive biology.
Pollination_Window_Closure is third and final Late_Anthesis aspect, distinguished by definitive termination of effective reproductive opportunity. Follows Final_Flowers_Opening and Declining_Pollen_Viability. Establishes final fruit set and berry number determining season yield. Marks transition from reproductive vulnerability to developmental phase. Enables management transitions including resumption of canopy operations and disease protection. Represents definitive endpoint of critical flowering period.
Befruchtungsende
Bestäubungsfensterschluss
Ende der Pollination
end of pollination period
flowering termination
pollination window end
reproductive closure
reproduktive Terminierung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 385-390 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 295-298
Pollination Window Closure
Chenin Blanc vines in the Swartland of South Africa bearing a pollinator habitat role are situated within a biodiversity-integrated vineyard where inter-row wildflower strips of indigenous fynbos species (Aspalathus, Oxalis, Nemesia) support 30-45 native bee species including ground-nesting Andrena and Lasioglossum species at densities of 150-300 individuals per 100 m transect during spring, with the vine canopy providing wind shelter and thermal buffering that extends daily foraging windows by 1-2 hours compared to open fynbos habitat.
A biodiversity support role that inheres in a grapevine whose vineyard environment provides foraging resources, nesting sites, and refuge habitat for pollinating insects including wild bees, hoverflies, butterflies, and other anthophilous arthropods, contributing to pollinator conservation within agricultural landscapes where pollinator habitat is typically scarce.
Pollinator_Habitat_Role is ecologically significant despite Vitis vinifera being primarily wind-pollinated (anemophilous) and not directly dependent on insect pollination for fruit set. The role captures the vineyard's landscape-level contribution to pollinator conservation rather than a direct mutualistic relationship: vineyards embedded in biodiverse landscapes with cover crops, hedgerows, and wildflower margins function as pollinator refugia within agricultural matrices. This role is increasingly recognized in EU agri-environment schemes that incentivize pollinator-friendly management in permanent crop systems.
Bestäuberhabitat-Rolle
Bestäuberschutz-Rolle
Bienenlebensraum-Rolle
bee habitat role
pollinator refuge role
pollinator support role
Kehinde T and Samways MJ (2014) Insect-flower interactions in vineyards, Journal of Insect Conservation 18: 47-58 Kratschmer S et al. (2019) Tillage intensity or landscape features: what matters most for wild bee diversity in vineyards?, Agriculture Ecosystems and Environment 270: 142-152 Forrest JRK et al. (2015) Bees, landscapes and habitat in the vineyard, American Journal of Enology and Viticulture 66: 60-67
Pollinator Habitat Role
Polymeric pigment formation proceeds through multiple reaction pathways including direct anthocyanin-tannin condensation (A-T and T-A adducts), acetaldehyde-mediated bridging (ethyl-linked polymers), and pyranoanthocyanin formation (vitisins, portisins). Cultivars with high initial anthocyanin and tannin concentrations such as Tannat and Cabernet Sauvignon show rapid polymeric pigment accumulation, reaching 40-60% of total wine color from polymeric forms within 12-18 months versus 20-35% in lower-phenolic cultivars such as Pinot Noir. The ratio of small polymeric pigments (SPP, trichloroacetic acid-soluble) to large polymeric pigments (LPP, trichloroacetic acid-precipitable) shifts from 3-5:1 in young wines to 1:1 or lower in aged wines as polymerization continues. Pyranoanthocyanins (vitisin A, vitisin B) contribute orange-red hues and exceptional stability, with concentrations reaching 5-20 mg/L in aged wines. The rate of polymeric pigment formation is enhanced by higher temperature, micro-oxygenation, and lower pH conditions during aging.
A color stability disposition that is the inherited capacity of a Vitis vinifera plant to produce grapes with anthocyanin and tannin compositions favoring the formation of covalently linked anthocyanin-tannin polymers during winemaking and aging, creating pigmented polymers that are resistant to SO₂ bleaching, less sensitive to pH-induced color loss, and provide the stable brick-red color of aged red wines.
Polymeric pigment formation represents the ultimate stabilization mechanism for red wine color, converting labile monomeric anthocyanins into stable covalent structures. The trait depends on grape composition (anthocyanin and tannin concentrations, tannin structure) and winemaking/aging conditions (oxygen exposure, acetaldehyde formation, time). Understanding cultivar-specific polymerization capacity enables prediction of wine aging trajectory and optimization of cellar practices. The color shift from purple-red (monomeric anthocyanins) to brick-red (polymeric pigments) during aging reflects the changing pigment composition rather than simple degradation.
Anthocyan-Tannin-Polymerisation
Polymerpigment-Bildung
anthocyanin-tannin polymerization
pigmented tannin formation
stabile Farbpigmente
stable pigment capacity
Somers TC (1971) The polymeric nature of wine pigments. Phytochemistry 10:2175-2186 Harbertson JF et al. (2003) A survey of polymeric pigments and phenolics in red wines. American Journal of Enology and Viticulture 54:301-306
Polymeric Pigment Formation Disposition
Pome fruit aroma profiling in Granny Smith-scented Grüner Veltliner from Wachau identifies green apple (7.8 on 10-point scale) as the dominant descriptor, complemented by pear (5.2) and quince (3.5), with the green apple prominence correlating with pre-malolactic malic acid concentration of 4.8 g/L and cool fermentation preserving hexyl acetate at 320 μg/L, demonstrating how the pome fruit profile reflects both grape composition and fermentation management in this variety valued for its crisp, apple-forward aromatic expression.
Pome fruit aroma quality is a fruity aroma quality that represents olfactory perception evoking apple and pear family fruit-associated scents including green apple, ripe apple, red apple, pear, quince, and related notes, characterizing the crisp, often clean aromatic impression particularly associated with cool-climate white wines and certain fermentation characters. Within the BFO framework, this quality addresses a subcategory of fruity aroma distinguished by its association with freshness, moderate ripeness, and specific fermentation-derived compounds, recognizing that pome fruit character bridges grape-derived and fermentation-derived aromatic contributions. The ontological significance lies in pome fruit aroma serving as an indicator of both climate and ripeness (green apple suggesting cooler conditions or earlier harvest) and fermentation character (pear and apple esters from yeast metabolism), with the specific pome fruit note providing information about wine origin and production. The compositional basis for pome fruit aroma prominently features esters, with ethyl acetate and isoamyl acetate contributing pear character, hexyl acetate providing green apple notes, and ethyl hexanoate adding ripe apple impression, while certain aldehydes (hexanal, trans-2-hexenal) contribute green apple character in young wines. The ripeness continuum within pome fruit descriptors proceeds from green apple (underripe or cool climate) through ripe apple (optimal maturity) to baked apple (very ripe or aged), providing a quality-relevant progression within this descriptor category. The association between green apple and malolactic fermentation is notable, as malic acid (from Latin malum, apple) contributes to tart, apple-like impression that softens to less apple-forward character after conversion to lactic acid, demonstrating the link between compositional change and pome fruit perception. Understanding pome fruit aroma quality enables climate and ripeness assessment based on the specific pome fruit note, fermentation character evaluation from ester-derived pear and apple notes, and wine style prediction where pome fruit expression indicates freshness and moderate richness.
Pome_Fruit_Aroma_Quality addresses the apple and pear family subcategory of fruity aroma, encompassing green apple, ripe apple, pear, and quince notes. This quality bridges grape-derived and fermentation-derived aromatic contributions, with specific notes indicating ripeness level and production style. The green-to-ripe apple progression provides a quality-relevant descriptor continuum within this category.
Apfel-Birnen-Duft
Kernobstaroma
Kernobstcharakter
Kernobstnoten
apple character
apple-pear aroma
pear scent
pome fruit notes
Ferreira V et al. (1995) Quantitative determination of the odorants of young red wines from different grape varieties. Journal of the Science of Food and Agriculture 68(3): 297-308 Swiegers JH et al. (2005) Yeast and bacterial modulation of wine aroma and flavour. Australian Journal of Grape and Wine Research 11(2): 139-173
Pome Fruit Aroma Quality
Poor fruit set quality assessment for boron deficiency in Margaret River Cabernet Sauvignon documented the progression from subtle pre-bloom indicators to severe fruit set failure in a vineyard with unrecognized boron limitation. Pre-bloom evaluation showed poor fruit set quality risk indicators including brittle shoot tips with slight necrosis at the growing point, mild interveinal mottling on youngest leaves, and inflorescences that appeared smaller and less developed than normal, symptoms initially attributed to spring weather conditions rather than nutritional status. Post-bloom assessment revealed severe poor fruit set quality with fruit set percentage of only 8% (normal 35-45%), berry count per cluster averaging 28 versus 85 in adjacent boron-sufficient vineyard, and numerous inflorescences showing near-complete coulure with only 3-5 berries surviving on rachis structures that retained their full branching pattern. Capfall examination during bloom had shown poor fruit set quality mechanism with abnormal pollen release, reduced pollen adhesion to stigmas, and pollen germination test showing only 22% germination in deficient block versus 78% in sufficient block. The cluster morphology resulting from poor fruit set quality showed loose "straggling" clusters with visible gaps between shoulder, body, and tip regions where flower positions had aborted, creating harvest yield of only 2.8 t/ha versus the regional average of 8 t/ha. Retrospective tissue analysis of retained bloom samples confirmed poor fruit set quality diagnosis with petiole B at 15 ppm (severely deficient) while all other nutrients were within adequate ranges, establishing boron as the sole limiting factor responsible for the reproductive failure.
Poor fruit set quality in boron deficiency is a boron deficiency symptom quality that inheres in grapevine inflorescences and developing clusters and represents the failure of normal flower fertilization and early fruit development resulting from boron-limited pollen viability, pollen germination, and pollen tube growth, manifesting as reduced berry number per cluster, increased coulure (berry abortion), cluster compactness reduction, and in severe cases near-complete fruit set failure leaving only scattered berries on an otherwise bare rachis. This quality captures the reproductive failure dimension of boron deficiency, recognizing that fruit set is often the most boron-sensitive developmental process and may be severely affected by boron limitation that produces minimal vegetative symptoms, making poor fruit set a critical diagnostic indicator of boron status during the reproductive period. The ontological significance of poor fruit set quality in boron deficiency lies in its direct yield impact and its potential to occur without obvious warning symptoms, creating situations where significant crop loss occurs before boron limitation is recognized, emphasizing the importance of proactive tissue testing and preventive boron management in regions with known deficiency risk. The biological basis of boron-limited fruit set encompasses multiple processes essential for successful fertilization, as boron's role in pollen development during microsporogenesis affecting pollen viability, boron's requirement for pollen germination on the stigma surface, boron's critical function in pollen tube cell wall synthesis where rapid elongation creates extreme boron demand, and boron's involvement in early embryo development where deficiency may cause post-fertilization abortion. The quantification of poor fruit set uses metrics including berries per cluster, percent fruit set (berries formed relative to flowers present), cluster weight, and cluster compactness rating, with boron deficiency typically reducing fruit set from the normal 25-50% to below 15% in moderate deficiency and below 5% in severe deficiency. The timing of boron limitation relative to flowering determines the severity and type of fruit set failure, with pre-bloom deficiency affecting pollen development causing reduced pollen viability, bloom-period deficiency affecting pollen germination and tube growth causing fertilization failure, and post-bloom deficiency affecting embryo development causing early berry abortion.
Poor_Fruit_Set_Quality_(B) is_a Boron_Deficiency_Symptom_Quality, representing the fertilization and fruit set failure from B-limited pollen function and embryo development. This class addresses the overall fruit set impact beyond the specific millerandage pattern. Fruit set can drop from normal 25-50% to <15% (moderate) or <5% (severe deficiency). Timing of B limitation (pre-bloom vs. bloom vs. post-bloom) affects mechanism. Coulure (complete berry abortion) vs. millerandage (partial shot berries) represents severity continuum. Distinguished from Hen_And_Chicken_Quality by emphasis on overall reproductive failure rather than the specific mixed-berry pattern.
B-induced coulure
Befruchtungsversagen durch B
Bor-Reproduktionsversagen
Bor-induzierte Coulure
Schlechter Fruchtansatz bei B-Mangel
boron fruit set failure
boron reproductive failure
fertilization failure from B
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication Dell B, Huang L (1997) Physiological response of plants to low boron. Plant and Soil 193:103-120
Poor Fruit Set Quality (B)
Porosity measurement of Tempranillo mesocarp tissue reveals 18.2±2.4% void volume at harvest, with gas pycnometry showing true tissue density of 1.089 g/mL compared to apparent density of 0.891 g/mL when air spaces are included, explaining the characteristic sponginess of grape pulp texture and the rapid juice release upon crushing compared to more compact fruit tissues with lower porosity.
Porosity quality is a physical quality that inheres in grape berry tissues and represents the volume fraction occupied by intercellular air spaces and non-cellular voids within the tissue matrix, characterizing the structural openness that influences gas exchange, mechanical properties, and density measurements. Within the BFO framework, this quality captures the void space component of tissue organization, complementing density by addressing the air-filled fraction rather than the material-filled fraction of tissue volume, with the two qualities mathematically related through the equation relating apparent density, true density, and porosity. The ontological significance lies in porosity determining diffusive pathways for oxygen, carbon dioxide, and volatile compounds within berry tissues, affecting respiration rates, ripening dynamics, aroma development, and susceptibility to oxidative browning upon tissue disruption. Grape berry porosity varies by tissue type, with mesocarp tissue showing higher porosity (10-25% void volume) than the compact cellular organization of skin (3-8%) or lignified seed coat (<2%), and these differences contribute to the tissue-specific density values observed in berry compartments. Measurement approaches for porosity include gas pycnometry comparing true volume (excluding air spaces) to apparent volume (including air spaces), mercury intrusion porosimetry for pore size distribution, and microscopic image analysis of tissue sections quantifying intercellular space area fraction. Developmental changes in porosity reflect the balance between cell division creating new intercellular spaces, cell expansion potentially compressing air spaces, and cell wall modification altering the connection between adjacent cells. Understanding porosity supports interpretation of berry texture measurements where air spaces affect compression behavior, prediction of extraction dynamics during winemaking where porous tissue releases cellular contents more readily, and modeling of post-harvest gas exchange affecting storage potential.
Porosity_Quality addresses the void fraction of tissue organization, serving as the conceptual complement to Density_Quality which addresses the material fraction. This class serves as parent for specific aspects of void space organization including Tissue_Porosity_Quality and Intercellular_Space_Quality. The distinction between connected (open) porosity allowing gas flow and isolated (closed) porosity affects functional interpretation. Measurement methodology significantly affects values as some approaches exclude smaller pores below detection thresholds.
Gewebeporosität
Hohlraumanteil
Luftraumvolumen
Porosität
air space volume
intercellular volume fraction
tissue openness
void fraction
Mavroudis NE et al. (1998) Apparent diffusion of solutes in plant tissue under conditions of steady state and continuous drying. Journal of Food Engineering 37(1): 27-41 Mendoza F et al. (2007) Characterization of porosity in apple tissue using magnetic resonance imaging and computer tomography. Journal of Food Engineering 78(3): 946-952
Porosity Quality
Touriga Nacional bears a Port_Style_Scion_Role as the Douro's most esteemed variety, contributing extraordinary color density (anthocyanins 2500-4000 mg/kg), intense floral-violet aromatics, and concentrated tannic structure to single-quinta Vintage Ports from Quinta do Noval, Quinta do Vesuvio, and Taylor's Vargellas commanding €100-500+/bottle. Touriga Franca serves in a Port_Style_Scion_Role as the most planted quality variety in the Douro (22% of vineyard area), providing perfumed aromatics, refined tannins, and elegant fruit character that complements Touriga Nacional in blends. Tinta Roriz (Tempranillo) bears a Port_Style_Scion_Role contributing bright fruit, firm acidity, and aging capacity, particularly valued in Tawny Port production for its oxidative evolution.
A fortified wine scion role that inheres in a grafted scion cultivar by virtue of its designation for Port wine production in the Douro Valley, Portugal, or Port-style wines elsewhere, where fermentation is arrested by grape spirit addition (aguardente at 77% v/v) at approximately 6-9°Baumé residual sugar, yielding wines of 19-22% alcohol and 80-130 g/L residual sugar. Port style scions are selected from the approximately 80 varieties permitted in the Douro, with five "recommended" varieties (Touriga Nacional, Touriga Franca, Tinta Roriz, Tinta Barroca, Tinto Cão) dominating quality production. These varieties must achieve exceptional phenolic concentration (anthocyanins, tannins) for color depth and structure that withstands fortification and extended aging, produce musts of moderate-to-high sugar (24-28°Brix) for balanced sweetness after arrested fermentation, and develop characteristic aromatic complexity (dark fruit, floral, spice) during extended cask or bottle aging. Port styles range from Ruby (fruit-forward, limited wood aging) through Tawny (oxidative cask aging) to Vintage (bottle-aged Vintage declarations).
Port_Style_Scion_Role captures the Douro's traditional field-blend complexity and the increasing trend toward single-variety expressions. The five recommended varieties represent a quality hierarchy formalized by IVDP (Instituto dos Vinhos do Douro e Porto), though many traditional quintas maintain mixed plantings of 20-40 varieties reflecting pre-phylloxera diversity. Port production involves distinctive techniques—foot-treading (lagares) or mechanical alternatives (autovinifiers, robotic lagares)—optimized for rapid extraction during the brief fermentation period (typically 2-3 days) before fortification. Climate change is driving Douro viticulture toward higher altitudes and north-facing slopes where the recommended varieties maintain balance at lower alcohol levels.
Douro variety
Douro-Traube
Port grape
Port variety
Port wine variety
Portsorte
Portweinsorte
Mayson R (2016) Port and the Douro 3rd Ed, Infinite Ideas Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Böhm J (2010) Port Wine and the Douro, IVV Symington Family Estates (2019) A Celebration of Port Varieties, Porto
Port Style Scion Role
Positioning ease quality comparison in Rheingau evaluated labor requirements and canopy organization outcomes for VSP-trained cultivars receiving standardized positioning operations. Riesling demonstrated excellent positioning ease quality with flexible young shoots readily inserting between paired catch wires, moderate growth rate permitting 3-week intervals between positioning passes, and strong orientation maintenance with less than 5% of shoots escaping between operations, achieving target canopy organization with 2.2 positioning passes averaging 12 hours per hectare total. Spätburgunder (Pinot Noir) showed good positioning ease quality with similar flexibility though somewhat faster growth requiring 2.5-week intervals and 2.8 passes totaling 16 hours per hectare. Müller-Thurgau exhibited poor positioning ease quality with stiffer shoots resisting catch wire insertion, vigorous growth requiring 2-week intervals, and persistent escape tendency with 18% of shoots requiring repositioning at each pass, demanding 4.1 passes totaling 28 hours per hectare to achieve marginally acceptable canopy organization, establishing positioning ease quality as a significant management cost factor favoring Riesling economics in the region.
Positioning ease quality is a training response quality that inheres in a grapevine cultivar or clone and represents the facility with which actively growing shoots can be manipulated into target orientations within catch wire systems or other positioning infrastructure during the canopy management operations that organize developing shoots into desired architectural configurations, determined by shoot flexibility during the positioning window, the responsiveness of shoots to imposed direction, and the tendency for positioned shoots to maintain rather than escape their prescribed locations. This quality directly determines the labor efficiency of shoot positioning operations and the achievable quality of canopy organization, with cultivars exhibiting high positioning ease requiring fewer passes through the vineyard and achieving superior canopy uniformity compared to those where shoots resist manipulation or persistently escape from positioned orientations. The ontological significance of positioning ease quality lies in its role as a primary determinant of canopy management labor requirements in training systems dependent on shoot positioning to achieve target canopy architectures, with labor differences of 50-100% or more between cultivars representing substantial economic impact across the multiple positioning operations required during each growing season. The biological basis of positioning ease quality encompasses shoot flexibility during the herbaceous growth phase when positioning operations occur, the strength of gravitropic and phototropic responses that may resist or reinforce imposed orientations, the internode length affecting the number of catch wire interactions securing positioned shoots, and the growth rate determining how quickly shoots extend beyond positioned zones requiring additional intervention. Cultivars with flexible shoots, weak directional growth responses, moderate internode length, and manageable growth rate exhibit high positioning ease, while those with stiff shoots, strong directional responses opposing imposed orientation, long internodes providing few catch wire contacts, or rapid growth quickly escaping positioned zones prove challenging to manage. Assessment of positioning ease quality involves quantification of positioning labor requirements, evaluation of canopy organization quality achieved with standardized inputs, and monitoring of shoot escape rates between positioning operations across the growing season.
Positioning_Ease_Quality is_a Training_Response_Quality, representing the facility of shoot manipulation during growing season canopy management. This class addresses the practical efficiency of shoot positioning operations in catch wire systems. Distinguished from Bending_Tolerance_Quality which addresses dormant cane manipulation rather than growing shoot positioning. The quantitative metrics of passes required and hours per hectare provide economic assessment basis. Most relevant for VSP and related training systems dependent on shoot positioning for canopy organization.
Einstellfreudigkeit
Heftarbeit-Effizienz
Positionierbarkeit
Triebeinordnungs-Leichtigkeit
catch wire compliance
positioning labor efficiency
shoot positioning suitability
tucking ease
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 64-85 Reynolds AG et al. (2004) Influence of training system and vine spacing on vine performance of Vitis vinifera L. cv. Riesling. American Journal of Enology and Viticulture 55:79-86 Vanden Heuvel JE et al. (2004) Influence of training system and shoot density on the light microclimate and fruit composition of Vitis vinifera L. cv. Riesling. American Journal of Enology and Viticulture 55:87-97
Positioning Ease Quality
Merlot in Bordeaux shows post-anthesis transition during early July Year N+1 transitioning from BBCH 69 toward BBCH 71 over 4-6 days July 3-8, with berry size increasing rapidly from 6-8 mm to 12-16 mm diameter through intense cell division, unfertilized flower abscission completing with cluster cleaning to predominantly berry appearance, resource allocation shifting to 40-55% photosynthate directing to berries, under oceanic conditions of 20-26°C temperature, typical of this mid-season cultivar establishing developmental phase and berry growth trajectory for season
Post-anthesis transition represents the physiological and developmental shift period following flowering completion when vine function reorganizes from reproductive to developmental priorities, berry growth accelerates, unfertilized flower abscission completes, and management focus transitions from bloom-period constraints to fruit development optimization. This critical transitional phase occurs during early July in temperate regions Year N+1 immediately following anthesis completion, typically spanning 3-7 days as physiological adjustments establish developmental phase dominance, corresponding to transition from BBCH 69 toward BBCH 71. Physiological reorganization during transition involves metabolic reorientation from reproductive to developmental processes with declining expression of flowering-related genes and increasing expression of fruit development genes, hormonal environment shifting from reproductive coordination to developmental regulation with auxin and gibberellin from berries dominating signaling, resource allocation redirecting from reproductive support to berry growth and vegetative development, and overall vine function transitioning to developmental focus. Berry development acceleration marks transition with rapidly increasing growth rates as cell division intensifies, berries doubling or tripling in size over transition period from 5-8 mm to 10-15 mm diameter, visible expansion creating recognizable fruit cluster appearance, and metabolic demands increasing substantially requiring enhanced photosynthetic support. Cell division intensity in berries reaches maximum during transition with cells dividing every 12-18 hours throughout pericarp tissues, cell number increase creating size potential for mature berries with cell production phase typically lasting total 2-3 weeks from fruit set, and developmental trajectory establishing toward subsequent cell expansion phase dominating later berry growth. Berry structure development during transition establishes anatomical organization with skin layers differentiating including outer epidermis with cuticle, hypodermis providing structural support, inner flesh parenchyma cells undergoing rapid division, vascular tissues developing connecting berry to pedicel, and seed structures developing within locules. Seed development progression shows embryo growth continuing with rapidly dividing cells, endosperm expansion providing nutritive support for embryo, integument layers developing into seed coat structure, and overall seed development coordinated with pericarp growth through hormonal signaling. Unfertilized flower elimination completes during transition with remaining senescing flowers detaching through completed abscission zones, daily flower drop creating accumulation of fallen flowers that will decay, and cluster cleaning as failed reproductions remove leaving only productive berries creating final cluster appearance. Abscission completion timing shows most flower drop occurring during transition week with 70-90% unfertilized flowers detaching, remaining 10-30% potentially persisting longer as dried brown structures before eventual removal, and overall cluster achieving predominantly berry-only appearance by transition conclusion. Cluster appearance evolution during transition shows dramatic change from mixed flower-berry state to predominantly berry cluster, green developing berries creating characteristic young fruit appearance, minimal brown debris from remaining senescing flowers, overall healthy productive cluster appearance establishing fruit development stage. Resource demand intensification during transition shows substantially increasing requirements with rapidly growing berries consuming 30-50% total photosynthate compared to 10-20% during flowering, vegetative growth continuing requiring 30-40% resources, and balanced allocation supporting both fruit development and vine growth. Photosynthetic capacity requirements increase during transition with canopy providing adequate production through 15-20 expanded mature leaves per shoot, shoot growth slowing but continuing with additional leaf expansion, and overall vine photosynthetic output supporting accelerating berry demands. Carbon allocation patterns show clear priority to developing berries as primary sink with photosynthate preferentially directing to fruit, vegetative growth continuing but secondary to fruit demands, and reserve accumulation minimal during peak fruit growth period. Hormonal regulation during transition shows auxin from developing seeds and pericarp preventing berry abscission and promoting cell division and expansion, gibberellins affecting berry cell division with peak concentrations during early development, cytokinins supporting cell division and delaying senescence, and integrated hormonal environment coordinating developmental processes. Nutrient requirements during transition emphasize nitrogen for protein synthesis supporting rapid cell division in berries and continued vegetative growth, potassium increasing importance for berry development affecting later sugar accumulation and quality, calcium essential for cell wall development in expanding berries, and balanced nutrition supporting optimal development. Water demand increases during transition with developing berries requiring adequate moisture for cell division and turgor maintenance, shoot growth continuing with water needs, and overall vine water use increasing from flowering levels requiring appropriate irrigation management if practiced. Environmental conditions during transition show reduced vulnerability compared to flowering with berry development relatively robust, though extreme heat above 35°C potentially stressing vines and affecting berry development, adequate moisture important for supporting rapid growth, and overall favorable conditions supporting optimal development. Temperature during transition optimally 20-28°C supporting rapid berry growth through cell division, with growing degree day accumulation driving development rate toward subsequent stages, and temperature effects on development rate affecting timing of later phenological events. Disease management during transition focuses on berry protection with fungicide applications for botrytis and powdery mildew preventing infection during rapid growth phase, recognition that early berry development represents susceptible period for establishing infections, and implementation of protective program supporting fruit quality. Pest management during transition includes monitoring for grape berry moth with egg laying on young berries potentially leading to larval damage, leafhoppers reaching peak populations in some regions requiring management, and other pests affecting vegetative or fruit tissues requiring attention. Canopy management implementation during transition takes advantage of flowering restrictions lifted enabling shoot positioning optimizing fruit exposure, hedging or trimming controlling excessive vegetative growth, strategic leaf removal improving fruit zone microclimate in some systems, and overall canopy structure optimization balancing vegetative growth with fruit development support. Crop load management considerations during transition include assessing fruit set outcomes relative to desired load, possible cluster thinning removing excess clusters if set substantially exceeded targets, leaving adjustment decisions recognizing vine capacity and quality goals, and timing of interventions affecting vine physiological responses. Vineyard operations resumption at transition includes normal spray programs without flowering restrictions, mechanical operations including mowing and cultivation, irrigation management establishing patterns for season, and overall return to regular management without bloom-period constraints. Phenological progression during transition shows advancement from BBCH 69 anthesis completion toward BBCH 71-73 early berry development stages, with developmental rate dependent on temperature accumulation and environmental conditions, and predictions for subsequent stages including véraison and harvest becoming possible based on early development patterns. The process represents third and final complete anthesis aspect marking full transition to developmental phase, following all flowers opened and anthesis completion. Duration typically spans 3-7 days representing adjustment period establishing developmental dominance, with warm conditions promoting rapid transition while cool conditions may extend period. Cultivar variation in transition characteristics shows differences in berry growth rates affecting development speed, resource allocation patterns affecting vegetative-reproductive balance, and timing of developmental stages affecting season progression. Regional timing shows post-anthesis transition during early July in warm climates, early-mid July in temperate regions, and mid-July in cool areas, occurring immediately following anthesis completion at approximately 1150-1250 growing degree days base 10°C. Viticultural significance includes establishing full developmental phase with berry growth as dominant process, enabling complete management transitions unrestricted by flowering sensitivities, marking critical period for establishing berry development trajectory affecting season outcomes, requiring adequate resources and favorable conditions supporting rapid berry growth, and providing reference for development progression and phenological predictions. Research applications include analyzing physiological transitions from reproductive to developmental states, evaluating resource allocation dynamics during phase transitions, quantifying berry development patterns during early rapid growth, assessing management effects on development trajectory affecting quality outcomes, and documenting phenological progressions supporting developmental modeling and prediction systems.
Post_Anthesis_Transition is third and final Complete_Anthesis aspect, distinguished by physiological reorganization establishing developmental phase dominance. Follows All_Flowers_Opened and Anthesis_Completion. Marks full transition from reproductive to developmental processes. Represents critical adjustment period with berry growth accelerating. Enables complete management transitions. Establishes developmental trajectory affecting season outcomes. Transitions toward Berry_Development major phenological phase.
Entwicklungsphasestart
Nach-Anthese-Übergang
Nachblütephase
developmental phase initiation
post-bloom phase
post-flowering transition
post-reproduktive Transition
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 395-410 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 300-315 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Post Anthesis Transition
Merlot flowers at late BBCH 65 through BBCH 67 (majority of bloom completed, 60-80% of flowers past peak anthesis) exhibit post-dehiscence anthers with fully opened dehiscence slits, substantially depleted pollen creating cream to light tan anther color (compared to bright yellow of actively dehiscing anthers), anther tissue beginning desiccation and structural collapse, with flowers at this stage indicating pollination completed 1-3 days previously and fruit set determination underway. Cabernet Sauvignon at BBCH 69 (end of flowering, >80% of flowers past anthesis) shows advanced post-dehiscence anthers with tan to brown coloration, shriveled appearance, and in some cases beginning abscission from filaments, providing visual confirmation that bloom period has ended and that transition to fruit development phase (BBCH 71-75) is imminent. Gewürztraminer at 4-6 days post-peak bloom exhibits post-dehiscence anthers ranging from light tan (more recent dehiscence) to dark brown (earliest dehiscence), creating heterogeneous anther coloration across inflorescence reflecting the acropetal bloom progression pattern, allowing retrospective assessment that bloom proceeded over 5-7 days from basal to apical flowers and that pollination window for the functionally female flowers extended across this entire period.
A post-dehiscence anther is an anther that has completed pollen release with dehiscence slits fully opened and pollen reserves substantially or completely depleted, showing characteristic color fading from bright yellow to cream, tan, or brown as pollen diminishes and anther tissues undergo senescence, representing the terminal functional state (late BBCH 65 through BBCH 69) when the anther has fulfilled its reproductive function and is transitioning to senescence and eventual abscission. Post-dehiscence anthers are characterized by= (1) longitudinal dehiscence slits fully opened and often widened compared to early dehiscence stage, with slit edges sometimes curled back or collapsed (2) pollen reserves substantially depleted with <30% of original pollen content remaining, or completely depleted with empty or nearly empty pollen sacs visible through opened slits (3) color transition from the bright yellow of mature and dehiscing anthers to lighter yellow, cream, tan, light brown, or brown, reflecting pollen depletion and tissue senescence with chlorophyll and carotenoid degradation (4) anther tissue showing progressive desiccation, loss of turgor, and structural collapse as cells undergo programmed senescence (5) in late post-dehiscence stages, anthers appearing shriveled, darkly pigmented (brown to dark brown), and structurally degraded. The post-dehiscence state typically begins 8-24 hours after dehiscence initiation (depending on pollen quantity and release rate) and persists for 3-7 days before anther and filament abscission occurs, representing BBCH stages 65-69 (majority of flowers past peak bloom through end of flowering). Post-dehiscence anthers retain residual pollen (typically 10-30% of original content, representing 1,500-9,000 grains per five-anther hermaphrodite flower) that remains adhered to internal anther surfaces or trapped in pollen sac recesses, with this residual pollen showing declining viability (typically 40-60% viable compared to 70-85% at peak dehiscence) as aging and environmental exposure cause deterioration. The biological functions of post-dehiscence anthers are minimal as pollen release has largely completed, though residual pollen release may continue at very low rates for 1-3 days post-peak dehiscence, contributing marginally to late pollination opportunities for late-receptive stigmas or for cross-pollination of later-opening flowers. Post-dehiscence anther color serves as a valuable visual indicator of bloom stage and pollination timing= flowers with cream to light tan anthers indicate bloom occurred 1-2 days previously with pollination likely completed flowers with tan to brown anthers indicate bloom occurred 3-5 days previously with pollination definitely past flowers with dark brown shriveled anthers indicate bloom occurred 5-7+ days previously and flower is approaching senescence and fruit set determination. The progression of anther color change provides retrospective assessment of bloom timing useful for= (1) determining pollination windows for functionally female cultivars requiring cross-pollination, allowing assessment of whether hermaphrodite pollinizer bloom overlapped adequately with female bloom (2) diagnosing fruit set failures by identifying flowers that achieved dehiscence (post-dehiscence anthers present) versus flowers that failed to open properly (mature or partially dehisced anthers retained) (3) estimating bloom duration and synchrony by observing the distribution of dehiscence stages across inflorescences or vines. Post-dehiscence anthers can serve as substrate for Botrytis cinerea and other fungal pathogens, with senescing anther tissue providing nutrients for saprophytic fungal growth, though anther infections are typically of minor consequence as the small tissue mass limits inoculum production and as most anthers abscise within days reducing persistence as infection sources. Environmental conditions during the post-dehiscence period influence the rate of anther senescence and abscission= warm dry conditions (25-30°C, 40-60% humidity) accelerate tissue desiccation and color change, promoting rapid progression to dark brown and abscission within 3-5 days cool moist conditions (15-20°C, 70-85% humidity) slow senescence allowing cream to tan colors to persist for 5-7 days before progression to brown stages.
Post_Dehiscence_Anther represents the pollen-depleted terminal state and is distinguished from Dehiscing_Anther (actively releasing pollen) by substantial pollen depletion, color change from bright yellow to cream/tan/brown, and from Mature_Anther (pre-dehiscence) by opened slits and reduced pollen content. The post-dehiscence state provides valuable retrospective information about bloom timing and pollination history through visual assessment of anther color and condition. Post-dehiscence anthers are of minimal reproductive significance but serve important diagnostic functions for assessing bloom completion, estimating pollination timing, and diagnosing reproductive development problems. The characteristic color progression (yellow → cream → tan → brown) provides a visual timeline of flower age and reproductive history useful for field assessment and research documentation.
entleerte Anthere
erschöpfter Staubbeutel
nach-Dehiszenz Staubbeutel
pollen-depleted anther
spent anther
Post Dehiscence Anther
Post drought growth resumption quality evaluation in Ribera del Duero Tempranillo compared vegetative recovery following early-season drought stress at different phenological stages. Stress imposed at BBCH 65 (full bloom) followed by irrigation relief resulted in excellent post drought growth resumption quality with shoot extension resuming within 4 days at rates achieving 85% of unstressed controls, and ultimately producing canopy leaf area within 10% of non-stressed vines by véraison. Equivalent stress imposed at BBCH 75 (berry pea-size) showed reduced post drought growth resumption quality with extension resuming after 7 days at only 60% of control rates, producing final canopy leaf area 25% below unstressed vines, attributed to the developmental transition away from vegetative growth priority that had begun by this stage. Stress at BBCH 81 (beginning of ripening) showed minimal post drought growth resumption quality response as vegetative growth had largely ceased naturally, with irrigation relief supporting fruit development rather than shoot extension. The phenological dependency of post drought growth resumption quality established that early-season stress episodes permit substantial canopy recovery while later stress episodes, though potentially less damaging to vegetative development that has already occurred, have limited recovery potential due to the developmental stage of the vines.
Post drought growth resumption quality is a drought recovery quality that inheres in a grapevine organism and represents the capacity to reinitiate shoot extension, leaf expansion, and other growth processes following relief of water deficit, determined by the restoration of turgor-driven cell expansion, the metabolic readiness to resume biosynthetic activities supporting growth, and the availability of resources including carbohydrates and nutrients to fuel renewed development. This quality captures the developmental dimension of drought recovery that determines whether and how quickly normal growth trajectories are restored following stress episodes, with implications for canopy development, fruit ripening capacity, and carbohydrate reserve accumulation that affect current and subsequent season productivity. The ontological significance of post drought growth resumption quality lies in its role determining the developmental consequences of drought episodes, recognizing that growth cessation during stress may have lasting impacts on canopy size, fruit load, and vine productivity that persist beyond the direct physiological effects of the stress period if growth resumption is delayed or incomplete following stress relief. The biological basis of post drought growth resumption encompasses the restoration of positive turgor in meristematic regions enabling cell division and expansion to resume, the reactivation of biosynthetic pathways producing cell wall materials, proteins, and nucleic acids required for growth, and the remobilization of carbohydrate and nutrient resources from reserves to growing points supporting the metabolic demands of resumed development. The timing of growth resumption following stress relief varies with stress severity, phenological stage, and genotype, with mild stress permitting rapid resumption within days while severe stress may cause prolonged growth cessation or redirect development toward recovery and reserve restoration rather than continued expansion. The phenological stage at which drought occurs strongly influences growth resumption dynamics, with pre-véraison stress affecting vegetative growth and canopy development while post-véraison stress affects ripening-period activities but typically has limited impact on vegetative growth that has naturally slowed by this stage.
Post_Drought_Growth_Resumption_Quality is_a Drought_Recovery_Quality, representing the reactivation of growth processes following stress relief. This class addresses the developmental dimension of recovery distinct from physiological restoration. The phenological stage interaction strongly influences resumption capacity and relevance. Growth rate relative to unstressed controls provides quantitative assessment. Distinguished from Rehydration_Rate_Quality and Hydraulic_Recovery_Quality which address water relations rather than developmental processes. Critical for understanding canopy development consequences of drought episodes at different growth stages.
Wachstumserholung
Wachstumswiederaufnahme nach Dürre
growth recovery
post-stress growth
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 453-507 Flexas J et al. (2006) Keeping a positive carbon balance under adverse conditions: responses of photosynthesis and respiration to water stress. Physiologia Plantarum 127:343-352 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data. Annals of Botany 105:661-676
Post Drought Growth Resumption Quality
Merlot post-fertilization early berries at BBCH 71 (8-16 days post-anthesis, immediately post-fertilization) range from 1.8-3 mm diameter and 0.006-0.015 g fresh weight, showing bright green coloration, firm texture, and containing 2-4 developing seeds (in successfully fertilized berries) with embryos at early globular stage, with successful fruit set (berry retention) occurring in 48-58% of flowers while unsuccessful flowers/fruitlets (unfertilized or inadequately fertilized) abscise during this 1-3 week window, creating the final berry number determining yield potential. Chardonnay post-fertilization early berries examined at 1-2 weeks post-fruit set show ovary wall mesocarp tissues with active cell division (mitotic index 12-18% of cells in M phase) particularly in middle mesocarp regions, vascular bundles in pedicel showing active differentiation with xylem and phloem elements elaborating, and developing seeds (3-4 per berry in successfully fertilized flowers) showing embryo development at 2-8 cell stage and endosperm cellularization beginning, with unfertilized ovaries in same inflorescence showing brown discoloration of stigma/style, no embryo development, and pedicel abscission zones activating leading to fruitlet drop by 2-3 weeks post-anthesis. Gewürztraminer functionally female cultivar shows post-fertilization early berries only on flowers that received adequate cross-pollen from hermaphrodite pollinizers, with successfully cross-pollinated flowers (28-38% of total) retaining berries that expand from 2 mm at 1 week post-pollination to 3-3.5 mm at 2 weeks, while unsuccessfully pollinated flowers (62-72%) showing arrested development at 1.5-2 mm and abscising by 2-3 weeks, demonstrating critical dependence of post-fertilization early berry retention on successful fertilization and embryo development. Pinot Noir experiencing heat stress (35-38°C daytime maximum) during immediate post-fertilization period (days 5-15 post-anthesis) shows elevated fruitlet abscission with only 32-42% fruit set versus normal 48-56%, with abscising fruitlets showing arrested development at 1.5-2.5 mm, impaired seed development (embryos arrested at 1-4 cell stage), and early pedicel abscission zone activation, demonstrating sensitivity of post-fertilization early berry stage to environmental stress.
A post-fertilization early berry is an early developing berry in the immediate post-fertilization period (BBCH 71, approximately 0-2 weeks post-fruit set) undergoing the critical ovary-to-fruit transition where fertilized ovaries are retained and begin expansion while unfertilized ovaries abscise, representing berries establishing developmental commitment and initiating the growth program that will produce mature fruits. Post-fertilization early berries are characterized by= (1) very small size typically 1.5-3.5 mm diameter and 0.004-0.02 g fresh weight, representing 10-25% of final mature berry diameter (2) ovary wall tissues beginning transformation into berry pericarp tissues with cell division initiating in mesocarp regions that will become flesh (3) fertilized ovules beginning seed development with embryo and endosperm formation proceeding (in seeded types) or fertilization initiating seed development that will later abort (in stenospermocarpic seedless types) (4) pedicel attachment strengthening through vascular differentiation and abscission zone suppression, distinguishing retained berries from abscising flowers/fruitlets (5) bright green coloration with active chlorophyll synthesis in developing skin tissues (6) extremely high surface-to-volume ratio creating high evaporative demand and sensitivity to water stress. Post-fertilization early berries undergo the fruit set determination process where successful retention versus abscission is decided= fertilized ovaries with viable developing embryos producing hormonal signals (auxins, gibberellins, cytokinins from developing seeds) that suppress abscission zone formation in pedicel, stimulate cell division in ovary walls, and establish sink strength drawing resources from maternal tissues unfertilized or unsuccessfully fertilized ovaries lacking adequate embryo-derived hormone signals, allowing abscission zone activation and fruitlet drop typically occurring 1-3 weeks post-anthesis. Post-fertilization early berries show critical developmental events= initial rapid cell division in ovary wall mesocarp tissues creating the foundation for subsequent berry growth vascular bundle elaboration in pedicel and berry establishing the transport infrastructure supporting later berry expansion seed coat development beginning in developing seeds providing mechanical support within berry establishment of developmental polarity with stylar end (blossom end) versus pedicel end (stem end) differentiation. Environmental conditions during the post-fertilization early berry period critically affect fruit set success= favorable conditions (moderate temperatures 18-28°C, adequate water and nutrition, moderate crop load) support high retention of fertilized berries achieving 40-60% fruit set unfavorable conditions (heat stress >32°C, water deficit, excessive shade reducing photosynthate availability, very high crop load creating competition) increase fruitlet abscission reducing fruit set to 20-40% or lower. Post-fertilization early berries are the most sensitive berry stage to coulure (excessive berry drop from poor fruit set) and millerandage (shot berry formation from arrested development), with disturbances during weeks 0-2 post-fertilization causing the highest incidence of these disorders that reduce yield and affect berry size uniformity.
Post_Fertilization_Early_Berry represents the immediate post-fruit-set period (0-2 weeks) and is distinguished from Initial_Growth_Early_Berry (2-6 weeks) by occurrence during the critical fruit set determination period versus established growth phase. Post-fertilization early berries undergo the ovary-to-fruit transition and fruit set establishment, with successful embryo development and hormone signaling determining berry retention versus abscission. Understanding post-fertilization early berry biology is essential for managing fruit set, for preventing coulure and millerandage, and for optimizing yield through cultural practices supporting successful fruit set during this critical period.
Fruchtansatz-Beere
Fruchtansatz-Stadium
fruit-set berry
initial fruitlet
nach-Befruchtung frühe Beere
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 175-185 Coombe, B.G. (1992). Research on development and ripening. American Journal of Enology and Viticulture 43=101-110 Ebadi, A. et al. (1996). Effect of short-term temperature and shading on fruit set. Australian Journal of Grape and Wine Research 2=2-9.
Post Fertilization Early Berry
Cabernet Franc in Loire Valley shows post-fertilization signaling during late June-early July Year N+1 at BBCH 69, with auxin biosynthesis initiating in 1.4-2.2 developing seeds within 3-6 days of fertilization, hormone export to ovary wall tissues, auxin accumulation reaching 200-400 ng/g fresh weight in pericarp by 2 weeks post-fertilization, anti-abscission and growth-promoting responses triggered, under temperate maritime conditions of 22-27°C temperature, typical of this mid-season cultivar establishing hormonal berry initiation
Post-fertilization signaling represents the initial berry initiation phase when developing seeds produce auxin and other hormones within days of fertilization, export these signals to surrounding ovary tissues, trigger anti-abscission responses preventing flower drop, and activate growth-promoting pathways initiating berry development, establishing essential hormonal communication linking seed formation to berry formation. This critical signaling phase occurs during early fruit set in late June-early July in temperate regions Year N+1, beginning within 2-5 days of fertilization and continuing throughout early berry development, corresponding to BBCH 69 fertilization completion stages. Signaling establishment involves auxin biosynthesis in developing embryo and endosperm as primary hormone signal, auxin transport from seeds through seed coat and funiculus to ovary wall pericarp tissues, hormone accumulation in target tissues reaching threshold concentrations triggering responses, additional hormone production including gibberellins and cytokinins contributing to signaling network, and integrated hormone communication establishing seed-berry developmental coordination. Auxin biosynthesis in seeds shows indole-3-acetic acid synthesis beginning within 2-5 days of fertilization, biosynthetic enzymes activating in embryo and endosperm tissues, hormone production rate correlating with seed developmental activity, total auxin output proportional to seed number creating dose-dependent effects, and sustained production throughout early seed development supporting berry growth. Auxin synthesis pathways show tryptophan-dependent biosynthesis as major route, enzymatic conversion through indole-3-pyruvic acid intermediate, additional pathways potentially contributing, and overall synthesis capacity determining hormone availability. Auxin export from seeds shows hormone transport across seed coat integument layers, movement through funiculus vascular tissues connecting seed to ovary wall, diffusion into pericarp creating hormone gradient, and accumulation in ovary wall tissues particularly outer cell layers receiving highest concentrations. Auxin transport mechanisms show active polar transport through auxin efflux carriers, possible non-polar diffusion contributing to distribution, and directional movement from seed source to ovary sink tissues. Auxin accumulation in pericarp shows hormone levels increasing over days to weeks reaching physiologically effective concentrations, peak levels typically 1-3 weeks post-fertilization during maximum seed activity, spatial gradient with higher concentrations near seeds, and temporal dynamics correlating with berry growth rate. Auxin concentration effects show threshold levels required for growth responses, concentration-dependent magnitude of cell division and expansion responses, and seed number effects mediated through total auxin dose determining berry size potential. Anti-abscission signaling shows auxin as primary anti-senescence signal preventing flower drop, hormone blocking abscission zone formation at pedicel-receptacle junction, suppression of abscission-related gene expression including cell wall degrading enzymes, maintenance of vascular connections supporting berry nutrient supply, and overall rescue of flower from programmed senescence pathway. Abscission zone prevention mechanisms show auxin downregulating ethylene biosynthesis preventing senescence trigger, hormone maintaining cell wall integrity preventing separation layer formation, preservation of vascular tissue preventing resource cut-off, and sustained cell viability maintaining structural connection. Growth-promoting signaling shows auxin triggering cell cycle activation in quiescent pericarp cells, upregulation of cell division genes including cyclins and cyclin-dependent kinases, stimulation of cell expansion through wall loosening and water uptake, activation of biosynthetic pathways producing growth substrates, and overall growth response transforming static ovary into rapidly expanding berry. Auxin perception and response shows hormone binding to auxin receptors in ovary wall cells, receptor-mediated signaling cascades activating transcription factors, auxin-responsive gene expression producing growth regulators and metabolic enzymes, and integrated cellular response coordinating division and expansion. Gibberellin signaling contribution shows possible gibberellin synthesis in seeds complementing auxin, gibberellin effects on cell division and elongation, synergistic interactions with auxin enhancing growth responses, and hormone contributing to overall developmental program though less characterized than auxin in grapevine. Cytokinin signaling shows possible cytokinin production in seeds, hormone effects on cell division particularly in combination with auxin, and cytokinin contributing to growth signaling network though evidence limited in grapevine berry initiation. Seed number effects on signaling show direct relationship between seed number and total hormone production, multiple seeds producing greater cumulative hormone output, dose-dependent berry growth response to total auxin levels, and seed number as determinant of signaling intensity affecting berry size. Temporal signaling dynamics show signal initiation within days of fertilization, progressive increase in hormone levels over first 1-2 weeks, peak signaling during maximum seed developmental activity 1-3 weeks post-fertilization, gradual decline as seeds mature and berry growth slows, and signaling duration extending throughout berry development though intensity varies. Spatial signaling patterns show localized high concentrations near individual seeds, hormone gradients across pericarp from seed sources, and overall distribution affecting regional growth patterns within berry. Environmental effects on signaling include temperature affecting hormone biosynthesis rate with optimal 22-28°C supporting normal production, nutrient availability affecting seed development and hormone synthesis capacity, and overall favorable conditions supporting robust signaling. Temperature effects show warm conditions promoting faster hormone production and stronger signaling, cool conditions slowing hormone synthesis potentially delaying berry growth initiation, and temperature as factor affecting signaling intensity and timing. Failed signaling causes include inadequate seed development preventing hormone production, abnormal hormone biosynthesis or transport, insufficient hormone accumulation below threshold levels, or seed abortion eliminating hormone source causing delayed berry abortion. Signaling assessment methods include hormone quantification in seeds and pericarp through analytical chemistry, gene expression analysis showing auxin-responsive genes, and growth response observation indicating successful signaling. The process represents first phase of berry initiation establishing hormonal communication, preceding ovary expansion initiation as growth begins. Duration begins within days of fertilization and continues throughout early berry development with peak 1-3 weeks post-fertilization. Cultivar variation may include hormone production capacity, signaling efficiency, and response sensitivity. Regional timing shows post-fertilization signaling beginning late June-early July across climates within days following fertilization. Viticultural significance includes representing essential seed-berry communication mechanism, requiring successful seed development for hormone production, demonstrating critical role of auxin in fruit set and development, and establishing signaling as prerequisite for berry growth preventing flower abortion. Research applications include analyzing hormone biosynthesis and transport mechanisms, investigating auxin and other hormone roles in berry initiation, evaluating seed number effects on hormone levels and berry growth, quantifying signaling dynamics and dose-response relationships, and assessing signaling as determinant of fruit set and berry development outcomes.
Post_Fertilization_Signaling is first phase of Berry_Initiation, distinguished by hormone production and communication establishing seed-berry link. Begins within 2-5 days of fertilization. Auxin from embryo/endosperm as primary signal. Prevents abscission and triggers growth. Seed number affects total hormone dose. Peak signaling 1-3 weeks post-fertilization. Essential for berry development initiation. Followed by Ovary_Expansion_Initiation as growth response begins. Critical determinant of fruit set success.
Auxin-Signalisierung
Post-Fertilisations-Signalisierung
Samen-Beeren-Kommunikation
Samen-Hormon-Signalisierung
auxin signaling
developmental signaling
seed hormone signaling
seed-berry communication
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 415-418 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Post Fertilization Signaling
Merlot post-harvest berries held 18 hours at 18-22°C due to processing capacity limitations show compositional changes from harvest= sugars declining from 24.5% to 24.2% through respiration, acids declining from 6.2 to 5.9 g/L through malic acid respiration, spontaneous fermentation beginning with 0.4% alcohol detected, minor oxidation of damaged berries visible as browning, requiring increased sulfur dioxide addition (60 mg/L versus 40 mg/L for freshly harvested) and immediate processing to prevent further degradation. Chardonnay post-harvest berries held 24 hours at 12-15°C for processing scheduling showing moderate quality changes= 0.1% sugar loss, 0.2 g/L acid decline, no fermentation due to cool temperature, minor oxidation requiring 50 mg/L sulfur dioxide (versus 30 mg/L freshly harvested), some aromatic loss with sensory evaluation showing reduced intensity (score 85 versus 88 for freshly harvested), but overall acceptable quality with appropriate sulfur dioxide and prompt processing. Pinot Noir post-harvest berries intentionally held for carbonic maceration= whole uncrushed clusters held 8 days at 15°C in carbon dioxide atmosphere, undergoing intracellular fermentation producing characteristic carbonic maceration aromatic profile (banana, bubblegum, strawberry notes), 2-3% alcohol from internal fermentation, berry skin color extraction into berry juice, creating distinctive wine style intentionally using extended post-harvest period for quality modification rather than quality degradation. Gewürztraminer post-harvest berries showing aromatic white sensitivity= 12 hour delay at 20°C causing 25% reduction in key aromatic compounds (linalool, geraniol) measured by GC-MS, significant loss of characteristic lychee and rose aromatics in sensory evaluation, demonstrating that aromatic cultivars are particularly sensitive to post-harvest quality degradation requiring freshly harvested state for optimal aromatic wine production. Sangiovese post-harvest berry management= harvest volume exceeding daily processing capacity requiring sequential processing over 3 days, with day 1 harvest processed immediately (freshly harvested), day 2 harvest held 24 hours at 10-12°C (minimal change), day 3 harvest held 48 hours at 10-12°C (moderate change), with quality maintained through cooling and sulfur dioxide addition, producing acceptable quality across all processing batches despite extended post-harvest period for later harvests, demonstrating that cool temperature control allows extended post-harvest periods with acceptable quality retention.
A post-harvest berry is a harvested berry in the extended post-harvest period (typically >6-12 hours post-detachment), characterized by progressive post-harvest quality changes including compositional shifts, enzymatic and microbial activity, and quality degradation relative to freshly harvested state, representing berries requiring careful handling to minimize quality losses or intentionally held post-harvest for specific processing objectives. Post-harvest berries are characterized by= (1) extended time since detachment typically 6-48+ hours allowing significant post-harvest changes (2) compositional changes including sugar decline through respiration (0.1-0.5% loss depending on temperature and duration), acid decline through malic acid respiration and fermentation, pH changes (3) potential spontaneous fermentation particularly in damaged berries or warm conditions with 0.2-1+% alcohol production from wild yeast activity (4) enzymatic browning in white grapes through polyphenol oxidase activity creating oxidized phenolics and brown color (5) volatile compound changes with loss of some aromatic compounds through volatilization or oxidation, development of fermentation-derived compounds (6) textural changes with softening through continued cell wall modification and potential maceration effects from juice release (7) microbial population growth with increase in bacteria and yeast populations potentially producing off-flavors or spoilage. Post-harvest berries show temperature-dependent quality changes= warm post-harvest conditions (>25°C) creating rapid quality degradation with significant compositional changes, active fermentation, oxidation, and spoilage risk within 12-24 hours moderate temperatures (15-25°C) showing moderate changes requiring processing within 24-48 hours cool temperatures (<15°C) slowing changes allowing extended post-harvest period of 48-96+ hours with minimal quality loss. Post-harvest berries may result from= intentional extended post-harvest handling for specific objectives (carbonic maceration requiring whole-berry fermentation, appassimento or drying styles, temperature conditioning before processing) unintentional delays from logistical constraints (harvest volume exceeding processing capacity, equipment failures, transport delays, weather interruptions) storage for processing scheduling allowing extended harvest period with sequential processing. Post-harvest berries for conventional wine production show quality challenges= need for prompt processing to minimize degradation increased sulfur dioxide requirements for oxidation and microbial control potential off-flavor development requiring sorting and adjustment reduced aromatic intensity in aromatic cultivars potential volatile acidity increase requiring monitoring and management. Post-harvest berries for specialized production may show intentional quality changes= carbonic maceration using whole un-crushed berries held 5-20 days developing characteristic fruity aromatic profile through intracellular fermentation appassimento styles using berries dried 30-120 days post-harvest concentrating sugars and flavors through dehydration cryoextraction using frozen post-harvest berries for concentrated juice extraction. Post-harvest berry quality management= temperature control maintaining <15°C slowing degradative changes sulfur dioxide addition controlling oxidation and microbes prompt processing when quality begins declining sorting excluding damaged or spoiled berries monitoring composition and microbial status guiding processing decisions.
Post_Harvest_Berry represents extended post-harvest period (>6-12 hours) and is distinguished from Freshly_Harvested_Berry (0-6 hours, minimal changes) by progressive post-harvest quality evolution. Post-harvest berries may result from intentional extended handling for specific styles or from unintentional delays, with quality management through temperature control and prompt processing critical. Understanding post-harvest berry physiology is essential for managing harvest logistics, for specialized production styles using extended post-harvest periods, and for minimizing quality losses when extended post-harvest periods are unavoidable.
Lager-Lese-Traube
held-harvest berry
nachernte Beere
nachgelagerte Traube
stored berry
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 275-280 Zoecklein, B.W. et al. (1995). Wine Analysis and Production, pp. 115-135 Flanzy, C. (ed.) (1998). Oenology= Grape Biochemistry, pp. 235-265.
Post Harvest Berry
Post optimal decline quality characterization for Riesling in Rheingau evaluated compositional trajectories during the four-week period following initial target ripeness for dry wine production at 85°Oechsle. The cultivar demonstrated favorable post optimal decline quality with slow sugar accumulation averaging 2°Oechsle per week permitting gradual progression through Spätlese (90°Oe), Auslese (100°Oe), and potential Beerenauslese (125°Oe) categories for producers pursuing late-harvest styles. Acid decline proceeded slowly at 0.8 g/L per week maintaining the high-acid profile essential for Riesling quality even at elevated must weights. Aromatic compound evolution showed characteristic terpene transformation from floral geraniol-dominant profiles toward more complex tropical and honeyed character without the degradation observed in less stable aromatic cultivars. The combination of slow sugar rise, acid retention, and aromatic stability established Riesling's exceptional post optimal decline quality enabling the full spectrum of German Prädikat wine production from the same vineyard through selective harvest timing, a stylistic flexibility unavailable from cultivars with rapid post-optimal quality decline.
Post optimal decline quality is a viticultural quality that inheres in a grapevine cultivar or clone within a specific growing environment and represents the rate and pattern of quality-relevant compositional and sensory changes occurring in fruit that remains on the vine beyond the optimal harvest date, encompassing the trajectories of sugar accumulation, acid metabolism, aromatic compound evolution, and structural characteristics that determine the consequences of delayed harvest whether intentional for stylistic purposes or inadvertent due to logistical constraints, weather events, or misjudgment of optimal timing. This quality fundamentally influences the risk exposure associated with harvest timing variation and the stylistic options available through extended hangtime, with cultivars exhibiting slow, stable post-optimal evolution permitting flexible scheduling and late-harvest style diversification while those showing rapid decline create harvest urgency and narrow the acceptable timing range. The ontological significance of post optimal decline quality lies in its role as the complementary parameter to harvest window duration, capturing not merely how long fruit remains acceptable but specifically how it changes during that period and what consequences follow from various degrees of harvest delay, enabling informed decisions about acceptable timing variance and late-harvest potential. The biological basis of post optimal decline quality encompasses the continued metabolic processes in attached berries including ongoing sugar accumulation from continued phloem import, respiratory consumption of organic acids especially malate, transformation and degradation of aromatic compounds through enzymatic and non-enzymatic pathways, modification of phenolic structure through oxidation and polymerization, and physical changes including berry softening, dehydration, and increased pathogen susceptibility that collectively determine the trajectory of quality-relevant parameters beyond optimal ripeness. Cultivar characteristics influencing post optimal decline quality include the inherent rate of sugar accumulation and acid respiration, the stability of varietal aromatic compounds under extended hangtime, the resistance to berry dehydration and shrivel, and the susceptibility to harvest-period pathogens particularly Botrytis that may either compromise quality through bunch rot or enhance it through noble rot depending on conditions and stylistic objectives.
Post_Optimal_Decline_Quality serves as the parent class for Sugar_Accumulation_Plateau_Quality, Acid_Decline_Rate_Quality, Aroma_Degradation_Rate_Quality, and Over_Ripening_Tolerance_Quality which address specific compositional trajectories beyond optimal ripeness. This class captures the overall post-optimal evolution concept while child classes address particular parameters. Distinguished from Harvest_Window_Duration_Quality by focusing on the nature of quality change rather than merely the duration of acceptable quality. Critical for late-harvest style production and harvest timing risk assessment.
Nachreife-Evolution
Post-Optimum-Qualitätsverlauf
Spätlese-Qualitätsdynamik
extended hangtime response
late-harvest quality dynamics
over-ripening trajectory
post-maturity evolution
Überreifungstendenz
Coombe BG, Iland P (2004) Grape berry development and winegrape quality. In: Dry PR, Coombe BG (eds) Viticulture Volume 1 Resources, 2nd Edition, Winetitles, Adelaide, pp. 210-248 Bindon KA et al. (2014) Relationships between harvest time and wine composition in Vitis vinifera L. cv. Cabernet Sauvignon. Food Chemistry 154:43-54 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 509-547
Post Optimal Decline Quality
Cabernet Sauvignon hermaphrodite flowers at 7-9 days post-calyptra shedding (BBCH 67-69, late to end of flowering) exhibit post-receptive dry stigmas with dry matte appearance contrasting with wet glistening receptive stigmas observed during days 1-6, sometimes showing remnants of dried exudate as whitish deposits on papillae surfaces, achieving only 12-22% pollen germination when viable pollen is applied at this late stage versus 78-86% germination during earlier receptive period (days 2-4), demonstrating that stigmas have completed receptivity and can no longer support effective pollination. Chardonnay stigma appearance progression from calyptra shedding through post-receptive senescence shows= wet glistening appearance days 1-5 (receptive period) with abundant visible exudate, transitional drying appearance days 6-7 with reduced exudate and beginning matte zones, and dry matte appearance with brownish papillae discoloration by days 8-10 (post-receptive period), with pollen germination declining from peak 82-88% at days 2-3 through 58-72% at days 5-6 to 18-28% at days 8-9, tracking stigma appearance changes. Gewürztraminer functionally female flowers show delayed transition to post-receptive dry state, maintaining wet receptive stigmas through day 7-8 post-calyptra shedding before transitioning to post-receptive dry by days 9-11, with the extended receptivity period (versus 5-7 day transition in hermaphrodite cultivars) providing extended window for capturing asynchronous cross-pollen from hermaphrodite sources and maximizing cross-pollination probability. Merlot flowers examined at 10 days post-calyptra shedding show post-receptive dry stigmas with dry brown senescing appearance, papillae partially collapsed, and remnants of dried crystallized exudate visible as deposits, achieving <8% fruit set when hand-pollinated at this late stage, confirming that post-receptive dry stigmas are past the effective pollination window and that successful pollination must occur during earlier receptive period.
A post-receptive dry stigma is a dry stigma that previously achieved receptivity with wet exudate production but has completed the receptivity period and ceased exudate secretion, with existing exudate having dried or been depleted, representing stigmas that have exited the pollination-competent state after the normal receptivity window has elapsed, characteristic of flowers beyond approximately 6-8 days post-calyptra shedding in hermaphrodite and functionally female/pistillate types. Post-receptive dry stigmas are characterized by= (1) dry matte appearance contrasting with earlier wet glistening appearance during the receptive period (2) absence of wet exudate with surface sometimes showing remnants of dried crystallized exudate components (sugars, proteins) appearing as deposits or films (3) papillae showing signs of senescence including reduced turgor, partial collapse, browning or discoloration, and cellular degradation (4) secretory apparatus in papillae cells degraded or shut down with cessation of exudate production (5) inability to support effective pollen germination, achieving <10-30% germination versus 70-90% during earlier receptive period, with reduced germination reflecting absence of exudate hydration and nutrition rather than active rejection of pollen. Post-receptive dry stigmas develop through natural senescence of stigma receptivity following completion of the receptivity window= exudate secretion declines and ceases as secretory cells undergo programmed senescence existing exudate on stigma surface evaporates or is depleted through pollen uptake (if pollination occurred) or environmental loss papillae cells undergo aging-related changes including membrane deterioration, organelle degradation, and eventual cell death. The timing of transition from receptive wet to post-receptive dry varies with flower type and environmental conditions= hermaphrodite stigmas typically transition at 5-7 days post-calyptra shedding functionally female and pistillate stigmas show delayed transition at 7-9 days due to extended receptivity environmental stress (heat, water deficit) accelerates transition to post-receptive dry state by 1-3 days. Post-receptive dry stigmas represent the normal terminal state of stigma development following successful or unsuccessful pollination, with stigmas completing their functional role and entering senescence phase leading to eventual browning and degradation as flowers transition to fruit set (if pollination was successful) or abscission (if pollination failed). Post-receptive dry stigmas retain physical evidence of earlier receptivity through residual dried exudate deposits and through maintenance of papillae structure despite senescence, distinguishing them from non-receptive dry stigmas that never produce exudate and often show reduced papillae development. The post-receptive dry condition can be used retrospectively to assess that receptivity occurred previously, providing evidence that stigma function was normal and that pollination failure (if fruit set did not occur) resulted from other causes (pollen unavailability, pollen non-viability, fertilization failure) rather than from stigma non-receptivity.
Post_Receptive_Dry_Stigma represents the normal terminal state following completion of receptivity and is distinguished from Non_Receptive_Dry_Stigma (which never achieved receptivity) by occurrence of earlier wet receptive period. Post-receptive dry stigmas are a normal developmental outcome following the receptivity window and indicate that stigma function proceeded normally through receptivity even if pollination/fertilization ultimately failed. Understanding post-receptive dry stigmas is relevant for interpreting stigma developmental state, for retrospective assessment of whether receptivity occurred, for understanding the temporal limits of effective pollination, and for diagnosing causes of fruit set failure (distinguishing stigma non-receptivity from post-receptive pollination timing failures).
erschöpftes Stigma
nach-rezeptive trockene Narbe
senescent dry stigma
spent stigma
verbrauchte Narbe
Post Receptive Dry Stigma
Syrah in Rhône Valley shows post-set berry abscission during late July-August Year N+1 at BBCH 77-81, with 6-14% established berries aborting during lag phase and veraison from severe water deficit stress, heat waves above 38°C, and occasional Botrytis infection, late drop visible as berry scars on clusters, under warm dry Mediterranean conditions with variable summer precipitation, typical of this moderately stress-tolerant cultivar showing limited late drop under adequate irrigation
Post-set berry abscission represents late berry drop occurring after successful initial fruit set when previously developing berries abort and abscise during later developmental stages from severe environmental stress, disease infection, or physiological dysfunction, causing berry loss typically during lag phase, veraison, or early ripening, and reducing yield from established berry number. This late drop phenomenon occurs during mid-late summer in July-September in temperate regions Year N+1, typically weeks 4-12 post-flowering after initial fruit set completion, corresponding to BBCH 75-85 berry touch through ripening stages. Post-set abscission mechanism involves late developmental failure or stress response in previously successfully established berries, abscission zone activation at berry pedicel despite earlier successful hormonal rescue, ethylene production potentially triggering abscission program, enzymatic cell wall degradation creating separation layer, and berry detachment reducing cluster berry number from established maximum. Late developmental failure causes show severe water deficit stress triggering berry abortion and drop, excessive heat stress disrupting berry metabolism and triggering drop response, disease infection particularly fungal pathogens causing berry death and abscission, sunburn damage causing berry death and subsequent drop, and overall severe stress exceeding berry tolerance causing failure. Water deficit-induced drop shows severe sustained water stress during lag phase or veraison triggering abscission, stress disrupting berry metabolism and hormone balance, ethylene production possibly mediating stress-induced drop, typical drop 5-20% under severe deficit though usually 25% except under catastrophic stress or disease, and post-set drop usually less severe than early fruit drop. Berry characteristics before late drop show berries initially developing normally through early phases, stress or disease causing visible symptoms before drop, shriveling, discoloration, or necrosis indicating problems, and symptomatic berries subsequently abscising. Dropped berry examination shows berries larger than early drop berries indicating later development before failure, disease symptoms visible in pathogen-induced drop, desiccation or damage in stress-induced drop, and pathology distinguishing late drop causes. Cluster appearance after late drop shows gaps from recently lost berries, pedicel scars indicating recent abscission, remaining berries potentially showing stress symptoms, and cluster architecture affected by late berry loss. Environmental effects include water deficit as major late drop cause particularly during lag phase or veraison, heat waves causing localized drop from damaged berries, disease pressure enabling infection-induced drop, and overall severe stress conditions triggering late abscission. Water deficit effects show progressive stress increasing drop risk, severe deficit >50% reduced vine water status triggering significant drop, timing during lag phase or veraison creating vulnerability, and adequate irrigation preventing deficit-induced drop. Heat effects show extreme events above 40°C causing damage and drop, heat stress exacerbating water deficit effects, and heat tolerance varying among cultivars affecting drop susceptibility. Disease effects show humid conditions promoting fungal infection, Botrytis bunch rot as major disease drop cause, and disease management reducing infection-induced drop. Cultivar variation shows genetic differences in stress tolerance affecting late drop susceptibility, some cultivars showing greater drought or heat tolerance, and disease resistance affecting infection-induced drop risk. Late drop versus early drop distinction shows late drop occurring weeks 4+ post-flowering after fruit set established, environmental stress or disease as causes versus developmental failure, typically lower incidence than early drop, and timing and causes distinguishing mechanisms. Drop prevention strategies show adequate irrigation during critical lag phase and veraison periods preventing deficit-induced drop, disease management reducing infection-induced drop, heat stress mitigation through cluster positioning or shading, and overall stress management minimizing late drop occurrence. The process represents fourth type of fruit drop caused by late developmental failure, distinguished from early developmental drop and other mechanisms. Duration represents stress or disease response after established fruit set. Regional timing shows post-set abscission during July-September across climates after initial fruit set completion. Viticultural significance includes representing late yield loss from environmental stress or disease, typically limited <5-10% under normal management though potentially severe under extreme conditions, demonstrating importance of water and disease management during later development, and affecting final yield from established berry number. Research applications include analyzing late abscission mechanisms and triggers, investigating stress and disease effects on established berries, evaluating management strategies preventing late drop, quantifying late drop impacts on yield, and assessing stress tolerance and disease resistance affecting drop susceptibility.
Post_Set_Berry_Abscission is fourth type of Fruit_Drop, distinguished by late occurrence after successful initial fruit set. Timing weeks 4-12 post-flowering during BBCH 75-85. Caused by severe stress (water deficit, heat), disease, or dysfunction. Reactivates abscission zone after earlier rescue. Typically limited incidence <5-10% under normal conditions. Distinguished from Early_Fruit_Drop by timing and causes. Prevention through irrigation, disease management, stress mitigation. Represents late yield loss from established berries.
Post-Set Beerenabszission
Veraison-Fall
late berry drop
post-commitment drop
später Beerenfall
stress-induced abscission
stressinduzierte Abszission
veraison drop
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 450-458
Post Set Berry Abscission
The post spacing specification for a VSP trellis in a Pinot Noir vineyard in the Côte de Beaune indicates intermediate posts at 4.0 m intervals (corresponding to every 4th vine at 1.0 m vine spacing) using treated acacia stakes of 50 mm diameter and 2.2 m length driven 0.5 m into the ground, with steel reinforced end posts at 120 mm diameter and 2.8 m length anchored by screw-in ground anchors at 45° — the 4.0 m intermediate post spacing represents a compromise between structural support (closer spacing of 3.0 m provides better wire tension retention and reduced canopy sway in wind) and cost efficiency (wider spacing of 6.0 m reduces material and installation costs by 35% but allows excessive wire deflection under crop load in high-yield years).
A training system description that documents the distance between adjacent trellis posts within vine rows and between row ends and anchor assemblies, specifying the structural spacing that determines the maximum unsupported wire span, the load distribution across the trellis framework, and the spatial frequency of rigid support points available for vine attachment and canopy management.
Post_Spacing_Specification defines the structural grid that determines both the mechanical performance of the trellis and the economic investment in vineyard infrastructure. Post spacing interacts with wire gauge and tension to determine the maximum allowable deflection under combined vine weight, crop load, and wind loading: closer post spacing (3-4 m) provides superior structural stability and canopy support but increases material costs (posts represent approximately 40% of total trellis material cost) and installation labor, while wider spacing (5-8 m) reduces costs but requires higher wire tensions and heavier gauge wire to maintain acceptable deflection limits. The post spacing also determines the spatial frequency of vine attachment points, which affects the uniformity of cordon positioning and the precision of shoot distribution along the fruiting wire — parameters that directly influence canopy microclimate homogeneity and fruit quality consistency within the vineyard.
Pfahlabstand-Spezifikation
Stickelabstand
Zwischenpfahl-Abstandsangabe
intermediate post distance
post interval specification
trellis post spacing
Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 66-96 Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles
Post Spacing Specification
Cabernet Sauvignon post-veraison berries tracked from veraison (week 11, BBCH 81, 11 mm, 1.5 g, 15% sugars, 17 g/L acidity, 1.2 mg/g anthocyanins, red-purple color) through maturity (week 18, BBCH 89, 14.5 mm, 2.7 g, 25% sugars, 6.2 g/L acidity, 3.4 mg/g anthocyanins, deep purple-black color) show 7-week post-veraison ripening period with progressive changes in size, composition, and color defining the post-veraison developmental trajectory from color change through harvest maturity. Chardonnay white cultivar post-veraison berries showing color change from green (BBCH 77, pre-veraison, chlorophyll 0.6 mg/g) to golden-yellow (BBCH 81-89, post-veraison, chlorophyll <0.1 mg/g, carotenoids providing yellow color), demonstrating that post-veraison applies to white cultivars with less dramatic but definite color change marking veraison transition. Pinot Noir post-veraison berry ripening rate across three temperature regimes= cool site (16°C average post-veraison temperature) requiring 10 weeks veraison-to-harvest accumulating 980 degree-days moderate site (19°C) requiring 8 weeks accumulating 1120 degree-days warm site (22°C) requiring 6.5 weeks accumulating 1230 degree-days, demonstrating temperature effects on post-veraison ripening duration with heat accumulation determining maturity timing. Syrah post-veraison berry monitoring for harvest timing= weekly sampling beginning week 10 (veraison) tracking sugar increase (week 10= 14%, week 12= 18%, week 14= 21%, week 16= 24%, week 18= 26%), acid decline (week 10= 19 g/L, week 12= 14 g/L, week 14= 10 g/L, week 16= 7 g/L, week 18= 5.8 g/L), and sensory evolution (week 10= tart unripe, week 14= developing fruit character, week 16= balanced ripe fruit, week 18= optimal maturity), with monitoring data guiding harvest decision at week 16-17 achieving optimal ripeness. Merlot post-veraison berry irrigation management= full irrigation throughout post-veraison producing larger berries (16 mm, 3.2 g at maturity) with moderate concentration (23% sugars, 6.5 g/L acidity) deficit irrigation during post-veraison producing smaller concentrated berries (14 mm, 2.4 g, 25% sugars, 5.8 g/L acidity), demonstrating post-veraison period as critical for irrigation effects on berry size and composition through effects on cell expansion and concentration.
A post-veraison berry is a developing or mature berry that has completed the veraison color change transition and entered the ripening phase (BBCH 81-89+), characterized by colored appearance (anthocyanin-colored in red/black cultivars, golden-yellow in white cultivars), softening texture, sugar accumulation, and acid decline, representing all berries from veraison completion through maturity and post-maturity, encompassing the entire Phase II ripening period and mature states. Post-veraison berries are characterized by= (1) complete or nearly complete color change with anthocyanins accumulated in red/black cultivars creating red, purple, or black berry color, or chlorophyll degraded in white cultivars creating golden-yellow color (2) softening initiated or progressing with firmness declining from green berry levels (4-6 N) toward mature levels (0.8-2.5 N) through cell wall modifications (3) sugars accumulating or at elevated levels typically 13-28% representing veraison levels (13-17%) through mature levels (18-28%) (4) acids declining or at reduced levels typically 4-18 g/L representing veraison levels (15-20 g/L) declining toward mature levels (4-10 g/L) (5) cells expanding or expanded with cell diameter 80-180 μm representing veraison size (60-90 μm) through mature size (120-180 μm) (6) berry size at or approaching mature dimensions with diameter 10-25 mm and fresh weight 1.5-7 g depending on cultivar and developmental stage (7) seeds mature or approaching maturity with brown coloration and lignification progressing. Post-veraison berries encompass a broad developmental range= early post-veraison berries (BBCH 81-83, 1-3 weeks post-color change) actively ripening with rapid sugar accumulation and expansion mid post-veraison berries (BBCH 83-87, 3-6 weeks post-veraison) continuing ripening at moderate rates late post-veraison berries (BBCH 87-89, 6-9+ weeks post-veraison) approaching or achieving maturity fully mature and overripe berries representing late/post-ripening states all falling within post-veraison category. Post-veraison berries show characteristic ripening processes= Phase II growth through cell expansion sugar accumulation through phloem import and vacuolar storage malic acid decline through respiration anthocyanin synthesis and accumulation (red cultivars) flavor compound development tannin modifications toward less astringent forms. Post-veraison berry development is highly temperature-dependent= growing degree-day accumulation from veraison to maturity typically 800-1400 degree-days (base 10°C) depending on cultivar warm conditions accelerating ripening achieving maturity in 6-8 weeks post-veraison cool conditions slowing ripening requiring 9-12+ weeks heat accumulation largely determining harvest timing variability across vintages and regions. Post-veraison berries are the focus of ripening management= irrigation strategies affecting berry size and composition concentration canopy management affecting berry light exposure and temperature crop load affecting resource availability and ripening rate harvest timing decisions based on compositional evolution during post-veraison ripening. Post-veraison berry sampling and monitoring guides harvest decisions= weekly or biweekly sampling tracking sugar accumulation rates acid decline monitoring assessing ripening progress sensory evaluation assessing flavor maturity analytical measurements (sugars, acids, pH, anthocyanins) combined with sensory assessment determining optimal harvest timing.
Post_Veraison_Berry represents all berries following veraison color change (BBCH 81-89+) and is essentially synonymous with Late_Developing_Berry plus Mature_Berry, emphasizing the veraison transition as the defining developmental marker. Post-veraison encompasses the entire Phase II ripening period and mature states. Sub-classification into Early and Late post-veraison reflects progression within the post-veraison period. Understanding post-veraison berry development is essential for ripening management, harvest timing, and quality optimization during the critical ripening phase.
Reifephase-Traube
nach-Véraison Beere
nachfarbwechsel Traube
post-color-change berry
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 196-245 Coombe, B.G., McCarthy, M.G. (2000). Dynamics of grape berry growth. Australian Journal of Grape and Wine Research 6=131-135.
Post Veraison Berry
Pinot Noir in Burgundy exhibits post-véraison firmness decline from 7.8 N to 2.4 N over 35-40 days, with the softening rate of 0.15 N/day correlating with anthocyanin extractability assessments that guide winemakers in determining optimal maceration intensity, where softer berries at harvest indicate higher extractability requiring shorter skin contact times.
Post-véraison firmness quality is a softening stage quality that inheres in a grape berry during the ripening period following véraison onset through approaching physiological maturity, characterized by progressive firmness decline resulting from active cell wall disassembly, reduced turgor pressure, and membrane permeability changes associated with ripening metabolism. Within the BFO framework, this quality represents the dynamic transitional state where mechanical properties change most rapidly, declining by 50-80% over 3-6 weeks as the berry transforms from a protective structure to a seed-dispersal organ optimized for palatability. The ontological significance of post-véraison firmness lies in its correlation with quality-determining processes including sugar accumulation, acid metabolism, anthocyanin synthesis in red varieties, and aroma compound development, making firmness trajectory a proxy indicator for overall ripening progression. Mechanistically, this quality reflects coordinated expression of polygalacturonase, pectate lyase, β-galactosidase, and expansin genes that solubilize pectin polymers, loosen cellulose-hemicellulose networks, and reduce cell wall thickness by 30-50%. The softening rate during this period exhibits strong cultivar dependency, with table grape varieties often selected for slower softening to extend post-harvest shelf life, while wine grape softening rates correlate with phenolic extractability critical for red wine production. Instrumental measurement during post-véraison requires standardization by thermal time accumulation (growing degree days) rather than calendar days due to temperature-dependent enzyme kinetics controlling softening rate. This quality provides the critical phenotyping window for irrigation management decisions, where controlled water deficit can modulate softening rate through turgor effects while simultaneously concentrating flavor compounds.
Post_Veraison_Firmness_Quality captures the most dynamic softening period, distinguished from Pre_Veraison_Firmness_Quality by active cell wall degradation and from Harvest_Firmness_Quality by ongoing rapid change rather than target state. Measurement timing should be expressed in accumulated thermal time (GDD base 10°C) post-véraison for cross-vintage comparability. This period corresponds to BBCH stages 81-85 (beginning of ripening through softening).
Erweichungsphase
Nachvéraison-Festigkeit
Reifefestigkeit
Reifungsfestigkeitsabnahme
maturation firmness decline
ripening firmness
softening phase firmness
véraison-to-harvest firmness
Nunan KJ et al. (1998) Changes in cell wall composition during ripening of grape berries. Plant Physiology 118(3): 783-792 Huang XM et al. (2005) Modification of cell wall composition and firmness during grape berry ripening. Journal of Agricultural and Food Chemistry 53(10): 4179-4187
Post Veraison Firmness Quality
Berry potassium concentration at harvest ranges from 1,500-4,000 mg/L juice across scion-rootstock combinations, with high-K rootstocks such as 140 Ruggeri and Ramsey conferring 30-50% higher fruit K than low-K rootstocks such as 101-14 Mgt and Riparia Gloire. Must pH correlates positively with K concentration (r = 0.65-0.80), with each 500 mg/L increase in K associated with approximately 0.15-0.20 pH units higher must pH. Potassium loading into berries occurs primarily via phloem transport during ripening, with VvK5.1 and VvKT2 channels mediating K influx into berry cells. Shiraz shows inherently high K accumulation disposition regardless of rootstock, while Cabernet Sauvignon shows greater rootstock-mediated variation.
A potassium uptake efficiency disposition that is the inherited capacity of a Vitis vinifera plant to accumulate potassium in specific tissues, particularly berries, with genotype-specific partitioning patterns determining the distribution of absorbed K between vegetative structures and fruit and affecting must potassium concentration and pH.
Potassium accumulation disposition has major implications for wine quality, as elevated juice K reduces titratable acidity through tartrate precipitation and increases pH, which can impair color stability, microbial stability, and sensory balance in red wines. Climate warming is exacerbating K accumulation through increased berry dehydration and concentration effects. Rootstock selection for low K accumulation is a primary management tool in warm climate viticulture where high pH is problematic. The trait shows complex genetic control with multiple QTLs identified in both rootstock and scion genomes.
K partitioning trait
K-Verteilung
Kaliumakkumulation
Kaliumeinlagerung
berry potassium loading
potassium distribution disposition
Walker RR, Blackmore DH (2012) Potassium concentration and pH inter-relationships in grape juice and wine of Chardonnay and Shiraz from a range of rootstocks in different environments. Australian Journal of Grape and Wine Research 18:183-193 Rogiers SY et al. (2006) Mineral sinks within ripening grape berries (Vitis vinifera L.). Vitis 45:115-123
Potassium Accumulation Disposition
Potassium acquisition rate assessment during fruit development in Cabernet Sauvignon reveals the seasonal demand pattern, with pre-véraison K⁺ uptake of 12 mg K/vine/day (moderate demand supporting vegetative and early fruit growth), post-véraison maximum of 38 mg K/vine/day (intense fruit K⁺ accumulation driving acquisition), and harvest-period decline to 8 mg K/day (demand satisfied, uptake downregulated), with the seasonal acquisition pattern totaling approximately 2.8 kg K/vine accumulated in fruit, demonstrating how acquisition rate must match the phenologically varying demand for adequate potassium nutrition.
Potassium acquisition rate quality is a potassium uptake efficiency quality that represents the rate of K⁺ absorption by grapevine roots through transporter and channel-mediated uptake systems, characterizing the kinetics of potassium entry into root cells that determines the vine's capacity to meet the substantial potassium demand driven by vegetative growth and fruit accumulation throughout the growing season. Within the BFO framework, this quality specifically addresses the transport-mediated dimension of potassium uptake, distinguishing the absorption rate from overall potassium utilization while recognizing that the combined activity of K⁺ transporters and channels determines the rate at which roots can acquire potassium from soil solution at varying external concentrations. The ontological significance lies in potassium acquisition rate determining whether root uptake capacity can meet vine demand, where the high potassium requirements of grapevines (often 60-120 kg K/ha removed annually in fruit and pruning material) create sustained demand that acquisition rate must satisfy, particularly during the rapid fruit accumulation phase when potassium transport to clusters reaches maximum intensity. The molecular basis for potassium acquisition involves dual uptake systems with distinct kinetic properties, including high-affinity transporters (HAK/KUP/KT family, Km approximately 10-50 μM) that enable acquisition at low soil K⁺ concentrations, and low-affinity channels (Shaker-type K⁺ channels including AKT1, Km approximately 1-10 mM) that provide rapid uptake when K⁺ is abundant, with the relative contribution of these systems varying with soil potassium status. The diurnal and seasonal patterns of potassium acquisition show higher rates during active transpiration (enhancing mass flow delivery to roots), during rapid vegetative growth (high demand upregulating uptake systems), and during fruit development (strong sink demand driving acquisition), with uptake declining during dormancy and under conditions limiting root function. The soil-root interface dynamics affecting acquisition rate include K⁺ diffusion to root surfaces (intermediate mobility, faster than phosphate but slower than nitrate), root interception (roots encountering K⁺ through growth), and mass flow contribution (transpiration-driven delivery of dissolved K⁺), with the relative importance depending on soil K⁺ concentration and vine water use. Understanding potassium acquisition rate quality enables assessment of the kinetic capacity for K⁺ uptake, prediction of potassium nutrition under varying soil and demand conditions, and interpretation of vine potassium dynamics throughout the growing season.
Potassium_Acquisition_Rate_Quality addresses the kinetic capacity for K⁺ uptake through transporter and channel systems. This quality determines whether roots can meet the substantial and seasonally varying potassium demand. The class supports potassium nutrition assessment through acquisition rate characterization across growing conditions.
HAK/AKT activity
HAK/AKT-Aktivität
Kaliumabsorptionsrate
Kaliumakquisitionsrate
K⁺ influx
K⁺ uptake rate
K⁺-Aufnahmerate
potassium absorption rate
Véry AA, Sentenac H (2003) Molecular mechanisms and regulation of K⁺ transport in higher plants. Annual Review of Plant Biology 54: 575-603 Mpelasoka BS et al. (2003) A review of potassium nutrition in grapevines with special emphasis on berry accumulation. Australian Journal of Grape and Wine Research 9(3): 154-168
Potassium Acquisition Rate Quality
The potassium content of a McLaren Vale Shiraz vineyard (South Australia) on red-brown earth documents exchangeable potassium of 0.8 cmol(+)/kg (320 mg/kg) at 0-30 cm depth — this moderately high potassium availability, combined with the high potassium release rate from the illite-rich clay fraction and the vigorous Shiraz root system exploiting 120+ cm of soil depth, produces elevated berry potassium concentrations (2,500-3,500 mg/L in must) that raise must pH to 3.7-4.0 at harvest sugars of 24-26°Brix. The high must pH necessitates acid adjustment (tartaric acid addition of 1-2 g/L at crushing) to achieve the pH 3.4-3.6 range required for microbial stability and fresh wine character — a common management challenge in warm-climate Australian viticultural regions where the combination of high soil potassium, vigorous rootstocks, and warm ripening temperatures produces the chronic high-pH problem that distinguishes warm-climate wine chemistry from the naturally low-pH musts of cool-climate viticulture.
A nutrient analysis data component that records the concentration of plant-available potassium in the vineyard soil, determined by ammonium acetate extraction (1M NH₄OAc at pH 7.0) and expressed as exchangeable potassium in cmol(+)/kg or mg/kg, with potassium being the mineral nutrient of greatest enological significance because potassium accumulation in berries directly affects must pH (through the formation of potassium salts of tartaric and malic acids), wine stability (potassium bitartrate precipitation), and wine sensory character (the perception of acidity and freshness being inversely related to potassium-driven pH elevation).
Potassium_Content documents the nutrient with the most direct and consequential link between soil chemistry and wine composition — unlike nitrogen (whose soil-to-wine effect is mediated through canopy vigor and fruit zone microclimate) and phosphorus (whose effect on wine is minimal), potassium moves from soil through vine roots directly into the berry, where it accumulates in the flesh during the post-véraison period and persists into the wine, affecting pH, tartrate stability, and sensory character. The soil potassium supply involves three pools of decreasing availability — solution potassium (immediately available, typically 5-20 mg/L in soil solution), exchangeable potassium (held on clay and organic matter exchange sites, released to solution as plant uptake depletes the solution pool), and non-exchangeable or structural potassium (trapped within the crystal lattice of potassium-bearing clay minerals, particularly illite and vermiculite, released slowly through mineral weathering): the total plant-available potassium supply over a growing season depends on all three pools, making the exchangeable potassium measurement an imperfect predictor of vine potassium uptake in soils with large non-exchangeable reserves. The management of soil potassium for wine quality presents a paradox — potassium is an essential nutrient required for vine photosynthesis, sugar transport, and water relations (deficiency causing leaf margin necrosis, reduced photosynthesis, and impaired berry ripening), but excessive potassium uptake elevates must pH to levels that compromise wine quality and stability — the viticultural goal is therefore to maintain potassium supply within a narrow sufficiency range that prevents deficiency without promoting luxury uptake.
Boden-Kalium
K-Gehalt-Messung
Kaliumgehalt
exchangeable K measurement
potassium availability assessment
soil potassium record
Mpelasoka BS et al. (2003) A review of potassium nutrition in grapevines with special emphasis on berry accumulation, Australian Journal of Grape and Wine Research 9: 154-168 White RE (2003) Soils for Fine Wines, Oxford University Press Kodur S (2011) Effects of juice pH and potassium on juice and wine quality, and regulation of potassium in grapevines through rootstocks — a short review, Vitis 50: 1-6 Robinson JB (2005) Critical plant tissue values, ASEV
Potassium Content
Potassium content quality management in Barossa Valley Shiraz addresses the chronic challenge of warm-climate Australian viticulture, with typical must potassium of 2000-3000 mg/L producing harvest pH of 3.8-4.2 that requires tartaric acid addition (2-4 g/L) for microbiological stability and color preservation premium producers combat elevated potassium through selection of low-uptake rootstocks (Schwarzmann, 101-14 rather than high-uptake Ramsey), cover crop competition, and early harvest targeting 14.0°Baumé rather than extended hang time, achieving must potassium below 2000 mg/L and natural pH of 3.4-3.6 that supports the deeply colored, age-worthy wines commanding premium prices.
Potassium_Content_Quality represents the chemical quality characterizing the concentration of potassium (K⁺) in grape berry tissue, capturing the dominant mineral cation that profoundly influences must pH, wine tartrate stability, and vine physiological function. Potassium is the most abundant mineral in grape berries, typically comprising 50-70% of total mineral content, with concentrations ranging from 1000-3000 mg/L in juice depending on cultivar, rootstock, soil availability, and viticultural management. The critical importance of potassium in grape quality derives from its direct effect on must and wine pH through formation of potassium bitartrate, a weak acid salt that buffers pH higher than free tartaric acid. High potassium content produces elevated pH at equivalent titratable acidity, reducing color stability in red wines, increasing microbial spoilage risk, and creating flat-tasting wines lacking freshness. Potassium accumulates in berries primarily via phloem transport alongside sugars, with accumulation accelerating during post-véraison ripening when sugar import is maximal. This phloem-mediated accumulation creates a positive correlation between sugar concentration and potassium content, potentially exacerbating pH increases during extended hang time in warm climates. Rootstock selection significantly affects potassium uptake, with vigorous rootstocks (1103 Paulsen, 140 Ruggeri) accumulating more potassium than less vigorous choices (101-14, Riparia Gloire). Soil potassium availability depends on clay content, exchange capacity, and competing cations, with excess soil potassium producing luxury consumption and quality-damaging must composition. Viticultural management through rootstock selection, cover cropping to compete for potassium, and irrigation management provides tools for controlling berry potassium content.
Potassium_Content_Quality addresses the single most important mineral for wine quality due to its pH effects. The potassium-pH relationship operates through potassium bitartrate formation, where potassium displaces hydrogen from tartaric acid, reducing free acid and raising pH. The magnitude of pH effect depends on the potassium-to-tartrate ratio excess potassium beyond tartrate saturation has limited additional pH impact. Cold stabilization removes potassium bitartrate from wine, but cannot compensate for the pH established during grape ripening. Potassium analysis employs flame photometry, atomic absorption spectroscopy, or ICP-OES.
Beerenkaliumqualität
K content
K-Gehalt
Kaliumkonzentration
berry potassium quality
potassium concentration
potassium level
Mpelasoka BS et al. (2003) A Review of Potassium Nutrition in Grapevines with Special Emphasis on Berry Accumulation. Australian Journal of Grape and Wine Research 9(3): 154-168 Boulton R (1980) The Relationship Between Total Acidity, Titratable Acidity and pH in Grape Tissue. Vitis 19(2): 113-120
Potassium Content Quality
Potassium deficiency symptom quality assessment in Barossa Valley Shiraz documented the progressive development of K limitation symptoms through the ripening period on sandy soils with low cation exchange capacity. Early-season potassium deficiency symptom quality was subtle, with midsummer assessment showing only slight bronzing at leaf margins on basal leaves scored at 1.5 on a 0-5 severity scale. Post-véraison potassium deficiency symptom quality intensified dramatically as fruit potassium demand increased, with marginal necrosis scores reaching 3.5 by mid-ripening and 4.5 by harvest, representing progression from subtle bronzing to extensive brown necrotic margins affecting 40% of basal and mid-canopy leaves. Tissue analysis tracked the potassium decline, with bloom petiole K at 1.8% (marginal) falling to 0.9% at véraison (deficient, threshold 1.2%) as fruit accumulated potassium at vegetative tissue expense. Fruit potassium deficiency symptom quality included delayed and uneven ripening with Brix variation of 4° within clusters versus 1.5° in K-sufficient blocks, elevated titratable acidity (7.2 g/L versus 5.8 g/L), and reduced anthocyanin concentration (0.9 mg/g versus 1.4 mg/g). The potassium deficiency symptom quality assessment established that maintaining bloom petiole K above 2.5% was necessary to support the late-season fruit demand without developing foliar symptoms that indicated competition between fruit and foliage for limited potassium.
Potassium deficiency symptom quality is a macronutrient deficiency symptom quality that inheres in a grapevine organism and represents the characteristic visual symptoms and physiological impairments resulting from inadequate potassium availability, including the diagnostic marginal leaf necrosis (leaf scorch) progressing from older to younger leaves as potassium is remobilized, delayed and uneven fruit ripening, reduced cold hardiness, decreased drought tolerance, and compromised fruit quality with elevated acidity and reduced sugar accumulation, with potassium deficiency representing one of the most economically significant nutritional limitations in viticulture due to potassium's essential roles in osmoregulation, enzyme activation, and carbohydrate metabolism. This quality captures the grapevine response to potassium limitation, recognizing that high fruit potassium demand during ripening makes this nutrient particularly critical for fruit quality and that deficiency symptoms often intensify during the ripening period when fruit competes with vegetative tissues for limited potassium supplies. The ontological significance of potassium deficiency symptom quality lies in its direct impact on fruit quality parameters critical for wine production, with potassium affecting not only sugar accumulation and acid metabolism in fruit but also must and wine potassium levels that influence pH, tartrate stability, and overall wine quality. The biological basis of potassium deficiency symptoms encompasses the role of potassium as the most abundant cation in plant tissues essential for turgor maintenance and stomatal function, the requirement for potassium as activator of numerous enzymes including those involved in photosynthesis and carbohydrate metabolism, and the function of potassium in phloem loading and long-distance carbohydrate transport affecting fruit sugar accumulation. The timing of potassium deficiency symptom appearance typically shows intensification post-véraison when fruit potassium demand increases dramatically, with symptoms that may have been subtle during early season becoming pronounced as fruit competes with leaves for potassium, often producing rapid symptom progression during the ripening period that distinguishes potassium from other nutrient deficiencies.
Potassium_Deficiency_Symptom_Quality is_a Macronutrient_Deficiency_Symptom_Quality, serving as parent class for Marginal_Leaf_Necrosis_Quality_(K) and Berry_Ripening_Delay_Quality_(K) which address specific potassium deficiency manifestations. This class addresses the overall potassium deficiency syndrome while child classes capture particular symptom types. Mobile nutrient showing symptoms on older leaves first. Symptom intensification post-véraison reflects high fruit K demand. Petiole K thresholds: 2.0% adequate at bloom. Major fruit quality implications beyond foliar symptoms.
K deficiency symptoms
K-Mangel-Zeichen
Kaliummangel-Symptome
Kaliumstress-Indikatoren
Niedrig-Kalium-Symptome
low potassium symptoms
potassium limitation signs
potassium stress indicators
Potassium Deficiency Symptom Quality
Potassium uptake efficiency shows pronounced rootstock effects, with 140 Ruggeri conferring 50-80% higher K accumulation than 101-14 Mgt on the same scion variety. Total seasonal K uptake ranges from 60-150 kg K/ha depending on yield and vine size, with approximately 50-70% removed in fruit at harvest. Potassium uptake continues strongly post-veraison during berry loading, when 40-60% of total fruit K accumulates. High-affinity K transport (VvHAK/KUP family) dominates at soil K below 0.2 mM, while voltage-gated channels (VvAKT1) contribute at higher concentrations.
A macronutrient uptake disposition that is the inherited capacity of a Vitis vinifera plant to absorb potassium ions (K⁺) from the soil solution through potassium channel and transporter proteins, with genotype-specific efficiency determining potassium acquisition to meet the high demand of this most abundant cation in grapevine tissues.
Potassium_Uptake_Efficiency_Disposition serves as the parent class for potassium accumulation traits. Potassium is required in the largest quantities of any cation, functioning in osmotic regulation, enzyme activation, phloem transport, and must/wine pH buffering. Excessive K uptake, particularly from high-K rootstocks, can elevate juice pH and reduce wine quality in red cultivars. Rootstock selection for moderate K uptake efficiency is often preferred over maximum uptake capacity to optimize fruit composition.
K uptake capacity
K-Akquisitionseffizienz
Kaliumaneignungsvermögen
Kaliumaufnahme-Effizienz
potassium acquisition efficiency
potassium foraging trait
Mpelasoka BS et al. (2003) A review of potassium nutrition in grapevines with special emphasis on berry accumulation. Australian Journal of Grape and Wine Research 9:154-168 Kodur S (2011) Effects of juice pH and potassium on juice and wine quality, and regulation of potassium in grapevines through rootstocks. PhD Thesis, University of Adelaide
Potassium Uptake Efficiency Disposition
Potassium uptake efficiency comparison among rootstocks reveals the genotypic variation affecting fruit composition, with Riparia Gloire showing K⁺ uptake rate of 185 mg K/vine/week (high efficiency, vigorous K⁺ acquisition) resulting in fruit K⁺ of 2.8 g/L and wine pH 3.75, while 140Ru shows uptake of only 95 mg K/vine/week (moderate efficiency, restrained acquisition) with fruit K⁺ of 1.9 g/L and wine pH 3.52, demonstrating how rootstock potassium uptake efficiency directly affects fruit composition and wine quality parameters that are central to winemaking decisions.
Potassium uptake efficiency quality is a nutrient uptake efficiency quality that represents the capacity of grapevine root systems to acquire potassium from soil, characterizing the effectiveness of K⁺ absorption that determines vine potassium status affecting stomatal regulation, osmotic adjustment, enzyme activation, and carbohydrate transport essential for fruit development and wine quality. Within the BFO framework, this quality specifically addresses the potassium component of nutrient uptake, distinguishing potassium acquisition from other nutrients while recognizing that potassium is the most abundant cation in plant cells and plays critical roles in grapevine physiology including the regulation of stomatal function that couples to water use efficiency and the accumulation of potassium in berries that affects wine acidity and stability. The ontological significance lies in potassium uptake efficiency determining access to this essential macronutrient that grapevines accumulate in large quantities (often the highest mineral content of harvested fruit), where adequate potassium nutrition supports stomatal function, turgor maintenance, phloem transport of sugars to fruit, and enzyme systems throughout the vine, while deficiency causes characteristic symptoms including marginal leaf scorch and impaired fruit quality. The mechanistic basis for potassium uptake involves K⁺ absorption from soil solution (typically 0.1-1 mM K⁺) through both high-affinity transporter systems (HAK/KUP/KT family operating at low external K⁺) and low-affinity channel systems (Shaker-type K⁺ channels operating at higher K⁺), with the relative contribution depending on soil K⁺ availability and root transporter/channel expression. The soil factors affecting potassium availability include clay content and type (illite and vermiculite fixing K⁺ in interlayer sites, reducing availability), cation exchange capacity (affecting K⁺ retention versus leaching), competing cations (high Ca²⁺ or Mg²⁺ potentially interfering with K⁺ uptake), and soil moisture (affecting both mass flow delivery and root function), with these factors creating the variable K⁺ availability that uptake efficiency must address. The rootstock effects on potassium uptake are particularly significant in viticulture, with some rootstocks (Riparia, Rupestris types) showing high K⁺ uptake that can lead to excessive fruit potassium affecting wine pH, while others (140Ru, 1103P) show more moderate uptake, making rootstock selection important for managing fruit potassium in quality-focused production. Understanding potassium uptake efficiency quality enables optimization of potassium fertilization, rootstock selection for appropriate K⁺ nutrition, and management of fruit potassium affecting wine quality parameters.
Potassium_Uptake_Efficiency_Quality serves as the parent class for K acquisition components (Acquisition Rate, Use Efficiency). This quality addresses the uptake of the most abundant cation in plant cells affecting multiple physiological functions. The class supports potassium management and rootstock selection through K uptake efficiency characterization affecting fruit quality.
K uptake efficiency
K-Aufnahmekapazität
KUptE
Kaliumakquisition
Kaliumaufnahmeeffizienz
potassium acquisition
root K foraging
Mpelasoka BS et al. (2003) A review of potassium nutrition in grapevines with special emphasis on berry accumulation. Australian Journal of Grape and Wine Research 9(3): 154-168 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 197-245
Potassium Uptake Efficiency Quality
Potassium use efficiency assessment comparing irrigation strategies in Shiraz reveals the water management effect on K⁺ partitioning and quality, with full irrigation achieving KUE of 52 kg fruit/kg vine K (efficient biomass production but fruit K⁺ of 2.6 g/L, wine pH 3.72), while regulated deficit irrigation shows KUE of 38 kg fruit/kg vine K (lower biomass efficiency but fruit K⁺ of 2.0 g/L, wine pH 3.55), demonstrating how potassium use efficiency must be evaluated not only for agronomic productivity but also for quality implications, with moderate water stress potentially improving wine quality through reduced fruit potassium accumulation.
Potassium use efficiency quality is a potassium uptake efficiency quality that represents the effectiveness with which grapevines convert acquired potassium into biomass, yield, and quality parameters, characterizing the integrated efficiency of potassium distribution and function that determines the productive return on potassium investment and the ultimate impact of potassium nutrition on fruit composition and wine quality. Within the BFO framework, this quality addresses the utilization dimension of potassium nutrition, distinguishing efficiency of use from efficiency of acquisition while recognizing that the destination and function of acquired potassium, particularly its accumulation in fruit affecting wine pH and stability, may be as important as the total amount acquired. The ontological significance lies in potassium use efficiency determining both the agronomic productivity per unit potassium and the qualitative impact on fruit composition, where unlike nitrogen or phosphorus where more efficient use is generally desirable, potassium presents a more complex situation in viticulture because excessive fruit potassium accumulation can negatively affect wine quality through elevated pH, while adequate potassium is essential for stomatal function, sugar transport, and stress tolerance. The physiological roles of potassium affecting use efficiency include osmotic regulation (major contributor to cell turgor and stomatal function), enzyme activation (cofactor for numerous enzymes including those of carbohydrate metabolism), phloem loading and transport (essential for sugar movement to fruit), and charge balance (balancing organic acid anions affecting fruit pH), with the partitioning among these functions determining the productive impact of plant potassium. The fruit accumulation dynamics show potassium moving rapidly into berries post-véraison, potentially accumulating to levels (1.5-3.0 g/L in must) that significantly affect wine pH (each 1 g/L K⁺ increase raising pH by approximately 0.1 units), making the partitioning of potassium between vegetative function and fruit accumulation a key quality consideration. The management factors affecting potassium use efficiency include vine balance (vigorous vines may sequester more K⁺ vegetatively), crop load (more fruit potentially diluting K⁺ concentration), and deficit irrigation (potentially reducing K⁺ transport to fruit), with these factors offering management tools for optimizing potassium distribution relative to quality goals. Understanding potassium use efficiency quality enables integrated assessment of potassium nutrition considering both adequacy for vine function and quality implications of fruit accumulation, optimization of management practices affecting potassium distribution, and rootstock selection balancing potassium nutrition with quality-focused production.
Potassium_Use_Efficiency_Quality addresses the effectiveness of converting acquired potassium into productive and quality outcomes. This quality uniquely involves quality considerations where high fruit K⁺ may be undesirable. The class supports integrated potassium management through KUE assessment balancing nutrition adequacy with wine quality goals.
K utilization efficiency
K-Produktivität
KUE
Kaliumnutzungseffizienz
Kaliumverteilungseffizienz
potassium partitioning efficiency
potassium productivity
Mpelasoka BS et al. (2003) A review of potassium nutrition in grapevines with special emphasis on berry accumulation. Australian Journal of Grape and Wine Research 9(3): 154-168 Rogiers SY et al. (2017) Mineral sinks within ripening grape berries. Vitis 56(4): 169-180
Potassium Use Efficiency Quality
Potential alcohol for Pinot Grigio from Alto Adige (2024 vintage) is calculated at 12.8% vol. from a must measurement of 21.8° Brix using the regional conversion factor of 0.59 (% vol. = °Brix × 0.59), establishing the lot within the target alcohol range of 12.5-13.5% vol. for the estate's dry Pinot Grigio style. Potential alcohol for Riesling from the Rheingau measured at 83° Oechsle calculates to 11.1% vol. using the Oechsle conversion (approximate divisor of 7.5 for the relevant must density range), positioning the lot at the upper Kabinett/lower Spätlese quality level and informing the cellar decision between fully dry vinification (GG-style, all sugar fermented) or halbtrocken style (fermentation arrested at 12-18 g/L residual sugar).
Potential alcohol is a sugar content data entity that records the estimated ethanol concentration achievable through complete fermentation of the sugar present in grape must, expressed in percent by volume (% vol.) or occasionally in grams per liter of ethanol, calculated from the measured sugar content using empirically derived conversion factors that account for the stoichiometric yield of alcoholic fermentation by Saccharomyces cerevisiae, serving as the direct predictor of wine alcohol content and the primary metric for evaluating the relationship between vineyard sugar production and final wine composition.
Potential alcohol provides the critical link between vineyard sugar measurement (in any regional scale) and the winemaker's primary compositional target — the final alcohol content that fundamentally influences wine body, balance, aging capacity, and stylistic identity. The calculation relies on the Gay-Lussac equation for alcoholic fermentation (C₆H₁₂O₆ → 2C₂H₅OH + 2CO₂), which predicts a theoretical yield of 51.1% ethanol by weight from glucose, though actual fermentation yields are lower (approximately 47-48% by weight) due to glycerol formation, biomass production, and other metabolic diversions by yeast. The practical conversion factors used in different regions reflect these yield losses and incorporate regional calibration — the commonly used factor of 16.83 g/L sugar per 1% vol. alcohol (OIV reference) implies complete fermentation to dryness, while actual wines retain variable residual sugar that reduces the realized alcohol below the potential. Potential alcohol serves as a regulatory metric in several jurisdictions — EU wine regulations define minimum natural potential alcohol levels for quality wine categories, and chapitalization (enrichment) regulations specify maximum permissible alcohol increases from added sugar in terms of potential alcohol gain. The increasing trend toward lower-alcohol wine styles has elevated the importance of potential alcohol monitoring during ripening, as harvest timing targeted at lower potential alcohol (11-12.5% vol.) rather than maximum sugar accumulation requires earlier picking and careful monitoring of concurrent flavor maturity. The discrepancy between potential and actual wine alcohol, attributable to residual sugar, fermentation efficiency, and any must corrections (chapitalization, dilution, reverse osmosis), should be documented for complete analytical traceability.
Alkoholpotenzial
alcohol potential
erwarteter Alkoholgehalt
estimated alcohol
expected alcohol
potenzieller Alkohol
potenzieller Alkoholgehalt
predicted alcohol content
probable alcohol
voraussichtlicher Alkoholgehalt
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1, 2nd Ed, Wiley, pp.: 219-235 OIV (2021) Compendium of International Methods of Analysis, Method OIV-MA-AS312-01A Boulton RB et al. (1999) Principles and Practices of Winemaking, Springer, pp.: 102-120
Potential Alcohol
A potted vine of Sauvignon Blanc/SO4 grown in a 1.5 L container bears a Potted_Vine_Role, available for planting from April through July in Northern Hemisphere vineyards—the intact root ball enabling mid-season establishment impossible with dormant bare-root material. A potted green-growing Pinot Noir/3309C serves in a Potted_Vine_Role for accelerated vineyard establishment, actively growing at sale and continuing growth immediately upon transplanting without the dormancy-break delay affecting bare-root vines. A large-format potted vine (Cabernet Sauvignon/110R in 4 L container) bears a Potted_Vine_Role for vineyard replanting, the larger root system enabling rapid establishment in gap-filling situations where competition from adjacent mature vines would disadvantage smaller plants.
A nursery plant role that inheres in a grapevine grown in a container (pot, cell, or bag) with intact root system and growing medium, enabling transplanting with minimal root disturbance throughout an extended planting season. Potted vines (also termed container-grown vines) are produced by planting rooted cuttings, benchgrafts, or field-dug vines into containers filled with soilless media (peat, bark, perlite mixes) or composted soil, growing them under nursery conditions until sale. The potted format offers advantages including extended planting window (spring through summer versus dormant-season only for bare-root), reduced transplant shock (intact root ball minimizes root damage and disturbance), and improved establishment in challenging conditions (drought, poor soils, late planting). Potted vine production requires container inventory, growing medium, irrigation systems, and growing space, increasing per-plant costs compared to bare-root production but commanding price premiums reflecting the extended flexibility and improved establishment success. Container sizes range from small cells (50-100 mL) for young liners through standard nursery pots (1-4 L) to large bags (10+ L) for mature potted vines.
Potted_Vine_Role captures the containerized nursery product format that has expanded significantly in recent decades, offering flexibility advantages that offset higher production costs. The intact root ball is the defining characteristic—rather than the root pruning and desiccation risk inherent in bare-root harvest, potted vines retain complete root systems in growing medium, minimizing transplant shock. Potted vine production has enabled extended planting seasons, previously limited to dormant periods when bare-root material could survive harvest and transplanting container vines can be planted throughout the growing season when irrigation is available. The format is particularly valuable for vineyard renovation (gap-filling among established vines), high-value plantings where establishment success is critical, and regions with challenging spring conditions where dormant planting windows are brief or unreliable.
Containerrebe
Kübelrebe
Topfrebe
container vine
container-grown vine
getopfte Rebe
pot-grown plant
potted vine
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 380-390 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press Smart DR et al. (2006) Grapevine rooting patterns, American Journal of Enology and Viticulture 57: 89-104
Potted Vine Role
Comparative evaluation of Powdery_Mildew_Resistance_Disposition across Vitis germplasm at UC Davis Department of Viticulture and Enology employed standardized greenhouse screening with controlled E. necator inoculation (10,000 conidia/mL suspension). The V. vinifera cultivar 'Carignan' (susceptible control) developed extensive mycelial colonies covering >90% of inoculated leaf surface within 14 days, with abundant conidiophore production rated OIV 1 (fully susceptible). The resistant accession V. romanetii C166-043 carrying Ren4 showed no macroscopic colony development, with microscopic examination revealing 95% of penetration attempts arrested at the papilla formation stage and the remaining 5% triggering single-cell hypersensitive death, rated OIV 9 (highly resistant). Intermediate phenotypes in segregating populations displayed reduced colony density, smaller colony diameter, and decreased sporulation intensity, enabling quantification of resistance QTLs contributing to partial resistance phenotypes observed in many interspecific breeding lines.
A Powdery_Mildew_Resistance_Disposition is a Fungal_Resistance_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through defense processes upon encounter with the obligate biotrophic ascomycete Erysiphe necator (syn. Uncinula necator), the causal agent of grapevine powdery mildew. This disposition represents the capacity to limit one or more stages of the E. necator infection cycle, including conidial germination, appressorium formation, penetration peg insertion through the cuticle, haustorial establishment in epidermal cells, and subsequent hyphal proliferation and sporulation on the plant surface. Unlike downy mildew where infection occurs through stomata, powdery mildew requires direct cuticular penetration, making cell wall properties and rapid cell wall reinforcement critical resistance components. Powdery mildew resistance dispositions in Vitis operate through mechanisms including pre-formed physical barriers (cuticle thickness and composition), rapid papilla formation blocking penetration attempts, hypersensitive response killing penetrated cells before haustorial maturation, and post-penetration resistance limiting colony expansion and reproduction. Major resistance loci designated Run (Resistance to Uncinula necator) and Ren (Resistance from Erysiphe necator) have been mapped in various wild Vitis species, with Run1 from Muscadinia rotundifolia and Ren1 from V. vinifera 'Kishmish vatkana' being most characterized. Powdery mildew resistance is essential for sustainable viticulture globally, as E. necator thrives in both humid and semi-arid climates and can cause severe crop losses through direct berry infection leading to cluster rots and off-flavors in wine.
Powdery_Mildew_Resistance_Disposition is_a Fungal_Resistance_Disposition. This class is parallel to Downy_Mildew_Resistance_Disposition under Fungal_Resistance_Disposition. Child classes are organized by resistance locus type (Run_Mediated_Resistance_Disposition, Ren_Mediated_Resistance_Disposition) and by resistance mechanism (Quantitative_Powdery_Mildew_Resistance_Disposition, Ontogenic_Powdery_Mildew_Resistance_Disposition, Evasion_Based_Resistance_Disposition). The Run and Ren nomenclature follows Vitis gene naming conventions, with "Run" originally denoting resistance from Uncinula (former genus name) and "Ren" denoting resistance from other sources.
Echter Mehltau-Resistenz-Disposition
Erysiphe necator resistance
Mehltauwiderstandsfähigkeit
Oidium-Resistenz
PM resistance potential
oidium resistance disposition
Gadoury DM et al. (2012) Grapevine powdery mildew (Erysiphe necator): a fascinating system for the study of the biology, ecology and epidemiology of an obligate biotroph, Molecular Plant Pathology 13:1-16 Qiu W et al. (2015) Current understanding of grapevine defense mechanisms against the biotrophic fungus (Erysiphe necator), the causal agent of powdery mildew disease, Horticulture Research 2:15020 Dry IB et al. (2010) Molecular strategies to enhance the genetic resistance of grapevines to powdery mildew, Australian Journal of Grape and Wine Research 16:94-105
Powdery Mildew Resistance Disposition
Powdery mildew symptom quality assessment in Napa Valley Cabernet Sauvignon documented disease development patterns informing integrated management combining cultural and chemical control. Early-season powdery mildew symptom quality appeared in May as diffuse chlorotic patches on basal leaves representing initial flag shoot infections from overwintering cleistothecia, with colonies becoming visible as subtle gray-white powdery growth under 10× magnification before obvious sporulation developed. By June, powdery mildew symptom quality had progressed to clearly visible white colonies on leaves and green shoot surfaces, with colony density scored using a 0-4 scale (0 = none, 1 = trace, 2 = 1-25% coverage, 3 = 26-50%, 4 = greater than 50%). Cluster powdery mildew symptom quality was monitored intensively from bloom through véraison, with pre-véraison infections showing white powdery colonization of berry surfaces that, if untreated, would develop into the characteristic netlike scarring pattern as berries enlarged and epidermal cells died beneath the fungal colony. Severely infected clusters showed powdery mildew symptom quality including berry cracking from scarred epidermis unable to expand with berry growth, and the musty off-odor detectable in heavily affected fruit. The powdery mildew symptom quality monitoring established that maintaining cluster severity below 15% coverage through the véraison window was critical for wine quality, with blocks exceeding this threshold producing wines with elevated potassium, reduced color stability, and detectable off-flavors.
Powdery mildew symptom quality is a fungal infection indicator quality that inheres in grapevine organs infected by the obligate biotrophic fungus Erysiphe necator (syn. Uncinula necator) and represents the characteristic disease manifestations including the white-to-gray powdery mycelial colonies on leaf, shoot, and cluster surfaces, the chlorotic and eventually necrotic tissue beneath heavily infected areas, the netlike scarring on infected berries, and the characteristic sulfurous or musty odor associated with severe fruit infection, with symptom expression varying with infection timing, environmental conditions, and host tissue susceptibility. This quality captures the diagnostic features of the most widespread and consistently damaging fungal disease of grapevine globally, recognizing that powdery mildew occurs in virtually all viticultural regions regardless of rainfall pattern since the pathogen thrives under the warm, dry conditions that characterize many premium wine-growing areas. The ontological significance of powdery mildew symptom quality lies in its role enabling disease identification and severity assessment for management decisions, with the distinctive powdery white appearance providing reliable field diagnosis at relatively low magnification while the distribution and density of colonies indicates disease pressure and management efficacy. The biological basis of powdery mildew symptoms encompasses the superficial growth of E. necator mycelium on host epidermal surfaces, the penetration of epidermal cells by specialized feeding structures (haustoria) that extract nutrients without killing host cells, the production of chains of conidia from erect conidiophores creating the powdery appearance, and the eventual host cell collapse under heavy infection causing the necrotic symptoms of advanced disease. The environmental requirements for powdery mildew differ from most fungal diseases in that free water inhibits spore germination, with optimal infection occurring at 70-85% relative humidity and temperatures of 20-27°C, creating the counterintuitive pattern of most severe disease in dry growing seasons and irrigated vineyards. The tissue-specific symptoms reflect differential susceptibility, with young expanding tissues most vulnerable while mature leaves show ontogenic resistance, and with berry susceptibility declining sharply after approximately 8% sugar accumulation (early véraison) creating a critical fruit protection window.
Powdery_Mildew_Symptom_Quality_(Erysiphe_necator) is_a Fungal_Infection_Indicator_Quality, representing the diagnostic symptoms of Erysiphe necator infection. The powdery white colony appearance is pathognomonic and visible without magnification at moderate severity. Free water inhibits this pathogen (unlike most fungi), creating dry-climate disease pattern. Berry susceptibility window closes at early véraison (~8% sugar). The 15% cluster coverage threshold correlates with wine quality impacts. Most widespread fungal disease affecting grapevine globally.
Echter Mehltau-Symptome
Erysiphe symptoms
Erysiphe-Symptome
Oidium-Symptome
Uncinula symptoms
oidium symptoms
powdery mildew symptoms
Äscherich-Symptome
Gadoury DM et al. (2012) Grapevine powdery mildew (Erysiphe necator): a fascinating system for the study of the biology, ecology and epidemiology of an obligate biotroph. Molecular Plant Pathology 13:1-16 Calonnec A et al. (2004) Effects of Uncinula necator on the yield and quality of grapes (Vitis vinifera) and wine. Plant Pathology 53:434-445 Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests, 2nd Edition. APS Press
Powdery Mildew Symptom Quality (Erysiphe necator)
Evaluation of Pratylenchus_Resistance_Disposition at Agriculture and Agri-Food Canada, Summerland Research Station, assessed responses of 30 rootstock genotypes to Pratylenchus penetrans, the predominant lesion nematode in British Columbia and Pacific Northwest vineyards. Potted vines were inoculated with 3,000 mixed-stage P. penetrans per pot and assessed at 6 months for nematode reproduction (final population/initial inoculum = reproduction factor RF), root lesion severity (0-5 scale), and vine growth impact. Susceptible V. vinifera 'Chardonnay' (own-rooted) showed high reproduction (RF 18.5), severe lesioning (score 4.2), and 35% growth reduction. Several rootstocks demonstrated Pratylenchus_Resistance_Disposition, though resistance levels were generally more moderate than observed for Meloidogyne. '3309 Couderc' (V. riparia × V. rupestris) exhibited good resistance with RF 2.8, lesion score 1.5, and minimal growth impact (8% reduction). 'Schwarzmann' showed similar performance (RF 3.2, lesion score 1.8). Notably, some rootstocks highly resistant to Meloidogyne showed Pratylenchus susceptibility: 'Freedom' exhibited RF 12.5 and lesion score 3.5 for P. penetrans despite excellent Meloidogyne resistance, demonstrating that these resistance systems are genetically independent. For cool-climate sites with Pratylenchus pressure, '3309 Couderc' and related V. riparia hybrids provide the best available Pratylenchus_Resistance_Disposition.
A Pratylenchus_Resistance_Disposition is a Nematode_Resistance_Disposition that inheres in a Vitis plant root system and is realized through defense processes that limit penetration, migration, feeding, or reproduction by lesion nematodes of the genus Pratylenchus, migratory endoparasites that cause extensive cortical necrosis as they feed and move through root tissues. This disposition represents resistance to nematodes with a fundamentally different parasitic strategy than either sedentary root-knot nematodes (Meloidogyne) or ectoparasitic dagger nematodes (Xiphinema), requiring distinct resistance mechanisms. Pratylenchus species, particularly P. vulnus (walnut lesion nematode) and P. penetrans (northern root-lesion nematode), damage grapevine roots through mechanical destruction and enzymatic degradation of cortical cells during feeding and migration, creating characteristic brown necrotic lesions that coalesce and can girdle roots. Unlike Meloidogyne, Pratylenchus does not induce specialized feeding structures, instead moving continuously through tissues and feeding on sequential cells, making hypersensitive response less effective as a resistance mechanism. Resistance mechanisms against Pratylenchus may include physical barriers (thick-walled cells, suberized layers) impeding migration, chemical factors (terpenes, phenolics) toxic to nematodes or inhibiting their digestive enzymes, and rapid wound healing limiting damage expansion. Pratylenchus resistance has received less breeding attention than Meloidogyne or Xiphinema resistance, but is important in cool-climate viticulture where P. penetrans predominates and in replant situations where P. vulnus can build to damaging levels. Resistance sources exist in various Vitis species, though fewer highly resistant rootstocks are commercially available compared to Meloidogyne-resistant options.
Pratylenchus_Resistance_Disposition is_a Nematode_Resistance_Disposition. Lesion nematodes (Pratylenchus spp.) have a migratory endoparasitic lifestyle distinct from sedentary Meloidogyne or ectoparasitic Xiphinema, requiring different resistance mechanisms. P. vulnus is most important in warm climates while P. penetrans predominates in cool regions. Resistance to Pratylenchus is genetically independent from Meloidogyne resistance, necessitating specific screening when both nematode groups are present. Fewer highly resistant rootstock options exist compared to Meloidogyne resistance, representing an area for continued breeding improvement.
Läsionsnematoden-Resistenz
Pratylenchus immunity
Pratylenchus-Resistenz-Disposition
Wandernematoden-Abwehr
lesion nematode resistance
root lesion resistance
Pinkerton JN et al. (1999) Responses of some grape rootstocks to ring nematode and root lesion nematode, American Journal of Enology and Viticulture 50:275-282 Howland AD et al. (2014) Evaluation of grapevine rootstocks for resistance to the northern root-knot nematode and lesion nematode, American Journal of Enology and Viticulture 65:173-177 McKenry MV (1992) Nematodes, Grape Pest Management, 2nd Edition, University of California Division of Agriculture and Natural Resources
Pratylenchus Resistance Disposition
A Cabernet Sauvignon clone 15 vine maintained in the screenhouse facility at Foundation Plant Services (UC Davis) bears a Pre_Basic_Mother_Plant_Role, having undergone thermotherapy (60 days at 38°C), meristem tip culture, and comprehensive testing including woody indexing on St. George, Kober 5BB, and LN-33 indicators over three growing seasons, plus ELISA and RT-PCR panels for 47 known grapevine viruses—this plant serving as the ultimate source for all FPS-distributed clone 15 material. A Riesling clone 239 plant in the JKI nuclear collection (Germany) serves in a Pre_Basic_Mother_Plant_Role, maintained under controlled greenhouse conditions with annual HTS (high-throughput sequencing) verification ensuring freedom from both characterized and potentially novel virus agents.
A certified mother plant role that inheres in a grapevine maintained at the highest tier (Pre-Basic/Nuclear/Elite) of the certification hierarchy, representing the ultimate source material from which all lower-tier certified propagation material derives through controlled multiplication. Pre-Basic mother plants undergo the most rigorous pathogen elimination treatments (thermotherapy at 37-38°C for 60+ days, meristem tip culture, somatic embryogenesis) and comprehensive testing protocols (biological indexing on woody indicators over 2-3 years, ELISA panels for all regulated viruses, PCR/RT-PCR for difficult-to-detect agents, and increasingly high-throughput sequencing for novel virus discovery). Pre-Basic plants must test negative for all regulated pathogens and are maintained in vector-protected environments (screenhouses, isolated locations) to prevent reinfection. The Pre-Basic tier serves as the genetic and phytosanitary reservoir from which the entire certified propagation system is periodically renewed, with material multiplied through Basic and Certified tiers to meet commercial demand while maintaining traceability to the tested nuclear source.
Pre_Basic_Mother_Plant_Role represents the apex of the certification pyramid, where maximum investment in pathogen elimination and testing creates the clean genetic resource from which the entire certified propagation system derives. Pre-Basic maintenance is expensive—screenhouse facilities, specialized personnel, comprehensive testing programs—limiting Pre-Basic stocks to centralized repositories (FPS, ENTAV, JKI, IMIDRA, VCR) rather than commercial nurseries. The testing evolution from biological indexing alone (1960s-1980s) through serological methods (ELISA, 1980s-2000s) to molecular techniques (PCR, HTS, 2000s-present) has progressively improved detection sensitivity and expanded the list of regulated pathogens. Pre-Basic plants may be maintained for decades, with periodic retesting confirming continued freedom from infection any pathogen detection requires destruction and replacement from backup material or re-initiation of the elimination process.
Elitemutterpflanze
Präbasis-Mutterstock
Vorstufenmaterial
elite mother plant
nuclear stock plant
pre-basic mother vine
European Union (2020) Commission Implementing Directive 2020/177 Annex IV Golino DA et al. (2017) Foundation Plant Services and clean plant programs, HortScience 52: 1190-1194 Maliogka VI et al. (2015) Grapevine virus diseases, APS Press Roossinck MJ et al. (2015) Plant virus metagenomics, Annual Review of Phytopathology 53: 157-178
Pre Basic Mother Plant Role
In Pinot Noir vines of the Côte d'Or, Burgundy, pre budbreak quiescence extends through the final two weeks of March when accumulated growing degree days have reached 80-120 units toward the approximately 150-unit threshold for budbreak, with buds showing subtle swelling at BBCH stage 01, scale margins beginning to separate, and internal examination revealing enlarged meristematic zones with cells preparing for division, before transition to active budbreak occurs in early April when accumulated GDD cross the threshold.
Pre budbreak quiescence in Vitis vinifera represents the final phase of ecodormancy immediately preceding the onset of visible budbreak, characterized by growth-competent buds that have accumulated substantial but not yet sufficient heat exposure for growth initiation, and representing the transitional state between maintained quiescence and committed growth. This phase typically occurs during the final 1-3 weeks before budbreak when warming temperatures are providing increasing heat unit accumulation but the threshold for visible growth has not yet been crossed, with buds in an advanced state of readiness that will transition to active growth upon receipt of relatively small additional heat input. The physiological state during pre budbreak quiescence involves advanced preparation for growth, with metabolic activity substantially elevated above deep dormancy levels, growth-related gene expression increased, protein synthesis accelerated in preparation for the demands of cell division and expansion, and cellular ultrastructure modified toward the actively dividing configuration characteristic of growing meristems. The hormonal changes during pre budbreak quiescence include declining ABA to minimal levels, increasing cytokinin activity as root function increases with soil warming, elevated gibberellin signaling preparing for cell elongation, and sensitized auxin transport systems ready to establish the polar gradients required for organized growth. The visual appearance during pre budbreak quiescence may include subtle changes such as slight bud swelling, color changes in bud scales, or emergence of woolly trichomes at scale margins, representing the earliest externally visible signs of the internal changes preparing for growth, corresponding to BBCH stages 00-01. The transition from pre budbreak quiescence to active budbreak occurs when accumulated heat units cross the threshold for growth commitment, typically after 100-300 growing degree days depending on cultivar, at which point visible bud swelling accelerates and the irreversible process of budbreak commences with progression through subsequent BBCH stages. The duration of pre budbreak quiescence varies with the rate of heat accumulation, being compressed during rapid spring warming and extended during gradual or interrupted warming patterns.
Pre_Budbreak_Quiescence represents the final transitional phase within Quiescence_Maintenance immediately before visible growth. Distinguished from earlier quiescence by advanced preparatory state and proximity to growth threshold. Characterized by elevated metabolic activity, increased gene expression, and hormonal changes preparing for growth. Subtle visible changes may include bud swelling at BBCH 00-01. Duration typically 1-3 weeks depending on warming rate. Transition to active growth when heat unit threshold is crossed.
Kurz-vor-Austrieb-Quieszenz
Vor-Austrieb-Ruhe
austriebsnahe Dormanz
imminent budbreak state
late ecodormancy
near-budbreak dormancy
pre-growth quiescence
späte Ekodormanz
Pre Budbreak Quiescence
Pinot Noir inflorescences at late BBCH 59 exhibit pre-dehiscence intact calyptras 3-3.5 mm length showing pronounced swelling and yellowish-green color change from earlier dark green, visible loosening at base with slight gap formation along basal attachment zone, internal pressure from elongating stamens creating bulged appearance, with calyptra shedding beginning 12-24 hours later when morning temperatures reach 18-20°C threshold. Chardonnay at pre-dehiscence stage shows calyptras with light green coloration, 15-20% size increase compared to tightly sealed stage, and visible upward displacement of 0.3-0.4 mm as stamen filaments begin rapid elongation phase, with transition to separating calyptra (BBCH 60) occurring within 18-30 hours under favorable temperature conditions (21-25°C). Cabernet Franc experiencing unseasonably cool conditions (11-14°C) at pre-dehiscence stage shows prolonged pre-dehiscence period of 48-72 hours with delayed calyptra shedding, extended vulnerability to Botrytis infection, and eventual irregular bloom with 20-30% of flowers showing delayed or failed calyptra shedding reducing fruit set from typical 48-54% to 32-40%.
A pre-dehiscence intact calyptra is an intact calyptra in the late developmental stage showing visible swelling or bulging from internal pressure of elongating stamens, color changes from green to yellowish-green or light green indicating tissue senescence and preparation for abscission, and incipient loosening at the basal attachment zone suggesting abscission layer formation has begun, representing flowers in the final 12-48 hours before calyptra shedding with reproductive organs fully mature and poised for anthesis. Pre-dehiscence intact calyptras are characterized by= (1) distinct swelling or bulged appearance compared to earlier tightly sealed stage, with calyptra dimensions increased by 10-25% due to internal pressure from elongating stamen filaments pushing against calyptra apex (2) color transition from dark or medium green to yellowish-green, light green, or in some cultivars brownish-green, reflecting chlorophyll degradation and cell wall modification processes associated with abscission preparation (3) visible loosening or slight gap formation at the basal attachment zone, often appearing as a subtle separation line or slight lifting of calyptra base away from receptacle, indicating abscission layer formation through cell wall dissolution (4) calyptra texture changes from firm and turgid to slightly softer or more fragile as cell walls are modified enzymatically in preparation for separation (5) in some cases, visible upward displacement of the entire calyptra structure by 0.2-0.5 mm as elongating stamens begin lifting the calyptra before complete detachment. The pre-dehiscence state characterizes intact calyptras during late BBCH stage 59 (flower buds swollen, calyptras light-colored), representing the critical transition period immediately preceding BBCH stage 60 (first calyptras lifting/detaching, anthesis beginning). Pre-dehiscence intact calyptras contain fully mature reproductive organs ready for immediate function upon calyptra shedding= stamens with filaments beginning rapid elongation phase (filaments 1.5-2.5 mm, poised to extend to final 2.5-4 mm upon shedding), anthers fully mature with pollen grains at peak viability (70-85%) ready for immediate release upon anther dehiscence pistil fully developed (1.8-2.5 mm) with stigma prepared for receptivity, stigmatic papillae and secretory cells ready to produce wet exudate upon exposure. The physiological processes active during the pre-dehiscence stage include= (1) abscission layer formation at calyptra base through localized expression of cell wall-degrading enzymes (cellulases, polygalacturonases) that dissolve middle lamellae and weaken cell-cell adhesion (2) stamen filament elongation through rapid cell expansion driven by turgor pressure and cell wall loosening, creating upward mechanical pressure against calyptra apex (3) tissue senescence in calyptra cells including chlorophyll degradation, protein breakdown, and membrane deterioration preparing for organ shedding. The duration of the pre-dehiscence stage is typically 12-48 hours depending on temperature and cultivar, with warmer temperatures (22-28°C) accelerating progression to calyptra shedding while cooler temperatures (15-18°C) extend the pre-dehiscence period. Environmental conditions during the pre-dehiscence stage critically influence the timing and success of subsequent anthesis= favorable conditions (warm temperatures 20-28°C, moderate humidity 50-75%, calm to light winds) promote rapid, synchronous calyptra shedding and successful bloom unfavorable conditions (cool temperatures 85%, heavy rain) can delay or disrupt calyptra shedding, causing irregular bloom and reduced fruit set. Pre-dehiscence intact calyptras showing the characteristic swelling and color change serve as critical phenological indicators for bloom timing prediction, allowing viticultural management decisions (final pre-bloom fungicide applications, bloom-period irrigation management, crop load assessment) to be optimized for the imminent anthesis period.
Pre_Dehiscence_Intact_Calyptra represents the final intact calyptra stage immediately before shedding and is distinguished from Tightly_Sealed_Intact_Calyptra by swelling, color changes, and basal loosening indicating imminent dehiscence (within 12-48 hours), and from Separating_Calyptra by retention of complete attachment despite incipient abscission layer formation. The pre-dehiscence stage is the most critical period for bloom timing prediction and represents the final opportunity for pre-bloom management interventions. Monitoring pre-dehiscence calyptras for progression to separation provides accurate short-term (1-2 day) forecasting of anthesis onset, allowing optimization of fungicide application timing, pollination management for functionally female cultivars, and controlled crossing operations in breeding programs. The pre-dehiscence stage is also the period of maximum sensitivity to temperature effects on bloom timing, with small temperature differences (2-3°C) substantially affecting the time from pre-dehiscence to calyptra shedding.
dehiszenz-bereite Blütenhaube
kurz vor Öffnung
near-dehiscence calyptra
swollen pre-opening calyptra
öffnungsbereite Kronblattkappe
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 100-102 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 192-194 Lebon, G. et al. (2008). Sugars and flowering in the grapevine. Journal of Experimental Botany 59=2565-2578.
Pre Dehiscence Intact Calyptra
Merlot in Bordeaux shows pre-lag phase transition during mid August-early September Year N+1 at BBCH 79-81, with growth genes downregulated 75-88%, lag-associated genes upregulated 3-8 fold, respiratory rate declining from 2.4 to 1.7 μmol O₂ g⁻¹ h⁻¹, malate declining from 1.3% to 1.1% through respiration, sugars increasing from 9% to 12%, pH rising from 2.9 to 3.0, under oceanic conditions of 20-26°C temperature, typical of this mid-season cultivar transitioning metabolism
Pre-lag phase transition represents the metabolic reorganization phase during enlargement phase completion when berry shifts from growth-phase to lag-phase metabolism, transcriptional reprogramming downregulates growth genes and upregulates lag-associated genes, respiratory rate declines from growth levels toward lag quiescence, enzyme activities shift from growth-supporting to maintenance metabolism, and metabolic transition prepares berry for lag phase quiescence and subsequent veraison transformation, occurring weeks 7-10 as enlargement concludes. This metabolic transition phase occurs during late summer in mid August-early September in temperate regions Year N+1, proceeding over 1-2 weeks as metabolism reorganizes, corresponding to BBCH 79-81 majority berries touching to beginning of ripening stages. Transition mechanism involves transcriptional reprogramming with differential gene expression shifting metabolic programs, respiratory metabolism declining toward lag phase lower levels, enzyme activity changes implementing metabolic shift, hormone level changes possibly regulating transition, seed maturation signaling developmental readiness, and integrated metabolic reorganization establishing lag phase characteristics. Transcriptional reprogramming shows large-scale gene expression changes during transition, growth-related genes downregulated including expansins, cell cycle regulators, growth-promoting transcription factors declining to basal levels, lag phase-associated genes upregulated including stress response genes, secondary metabolism genes, ripening preparation genes, and coordinated expression changes implementing metabolic transition. Growth gene downregulation shows expansin genes reduced 70-90% from peak levels, cell wall modification enzymes declining, auxin response genes downregulated, cell division genes at basal non-functional levels, and growth program effectively silenced. Lag phase gene upregulation shows stress response genes including heat shock proteins, antioxidant enzymes increasing, secondary metabolism genes for phenolic biosynthesis upregulated, genes involved in organic acid metabolism and sugar transport beginning increase, and lag-associated expression establishing new program. Transcription factor changes shows growth-promoting transcription factors declining, developmental transcription factors directing lag and ripening preparation upregulated, hormone-responsive transcription factors shifting expression, and transcriptional regulatory network reorganization. Respiratory metabolism changes show oxygen consumption declining from growth phase 2-4 μmol O₂ g⁻¹ h⁻¹ to transitional 1.5-2.5 μmol O₂ g⁻¹ h⁻¹ approaching lag phase 1.0-1.8 μmol O₂ g⁻¹ h⁻¹, respiratory rate reduction reflecting declining metabolic activity, metabolic shift from active biosynthesis to maintenance, and respiratory decline as metabolic transition indicator. Respiratory enzyme activities show glycolysis and TCA cycle enzymes declining to lower activity levels, oxidative phosphorylation continuing at reduced rate, ATP production decreasing from growth demands to maintenance levels, and respiratory machinery adjusting to lower metabolism. Carbon metabolism changes show reduced biosynthetic activity as growth ceases, carbon partitioning shifting from structural synthesis toward storage and maintenance, beginning sugar accumulation in vacuoles indicating import activation, and metabolic reorganization toward ripening. Organic acid metabolism shows malate declining through respiration from peak 1.2-1.6% to lower 1.0-1.4%, malate dehydrogenase and malic enzyme activities affecting acid dynamics, tartrate remaining relatively stable 0.3-0.6%, and acid metabolism as key metabolic change. Malate respiration shows malic enzyme converting malate to pyruvate, pyruvate oxidation through TCA cycle, malate decline contributing to berry softening and pH increase, and acid respiration as characteristic metabolic shift. Sugar metabolism changes show beginning sugar import through phloem activation, sugar transporter gene expression increasing, sugar accumulation in vacuoles from 8-12% to 10-14%, and sugar metabolism preparing for ripening accumulation. Enzyme activity changes show growth-supporting enzymes declining to basal levels, maintenance metabolism enzymes sustained at moderate activity, ripening-associated enzymes beginning expression, and enzyme complement shifting toward lag and ripening programs. Cell wall enzyme changes shows expansin activity declining substantially, xyloglucan endotransglucosylase reducing activity, pectin-modifying enzymes beginning changes affecting softening, and wall enzyme activities implementing transition. Secondary metabolism changes shows phenolic biosynthesis pathway genes upregulating, flavonoid synthesis enzymes increasing expression, anthocyanin biosynthesis genes activated in red cultivars preparing for color change, and secondary metabolism as key transition feature. Hormone level changes shows auxin declining from growth phase levels, possible gibberellin changes affecting metabolism, ethylene possibly beginning increase in some cultivars, abscisic acid remaining low before veraison spike, and hormone dynamics possibly regulating transition. Protein expression changes shows growth-related proteins declining through degradation or dilution, lag-associated proteins accumulating from upregulated genes, stress response proteins increasing, and protein complement reflecting metabolic program shift. Metabolic flux changes shows reduced flux through growth pathways, maintained flux through maintenance pathways, increasing flux through sugar import and storage, and flux redistribution implementing metabolic transition. Seed maturation signaling shows mature seeds possibly signaling developmental progression, seed hormones potentially affecting berry metabolism, seed completion as prerequisite for transition, and seed-berry coordination in developmental progression. Berry composition during transition shows water 78-83% maintained, organic acids declining 1.0-1.4%, sugars increasing 10-14%, pH rising 2.9-3.1, and composition shifts toward lag characteristics. Berry morphology during transition shows berry at near-final size, green color maintained though chlorophyll beginning decline, texture softening slightly, and appearance approaching lag phase. Environmental effects on transition include temperature affecting transition kinetics with warm conditions accelerating metabolic shift, cool conditions extending transition period, and temperature as factor affecting transition timing. Temperature effects show warm conditions promoting faster transcriptional and metabolic changes, cool conditions slowing transition though occurring across temperature ranges, and temperature affecting timing not transition occurrence. Transition timing shows typical occurrence weeks 7-10 post-fertilization, cultivar variation with early cultivars transitioning earlier, timing correlating with enlargement completion, and transition establishing lag phase entry. Transition duration shows metabolic reorganization proceeding over 1-2 weeks, gradual changes implementing new program, and transition period overlapping with enlargement completion. Transition assessment shows gene expression profiling documenting transcriptional changes, metabolite analysis showing compositional shifts, enzyme activity measurements quantifying metabolic changes, and molecular markers indicating transition. The process represents second aspect of enlargement phase completion establishing metabolic reorganization, following final cell expansion and preceding growth deceleration. Duration typically 1-2 weeks representing metabolic transition period weeks 7-10. Regional timing shows pre-lag phase transition during mid August-early September across climates as metabolism reorganizes. Viticultural significance includes representing critical metabolic shift preparing lag phase and ripening, demonstrating developmental progression from growth to quiescence, establishing metabolic foundation for subsequent ripening transformation, and marking major developmental transition. Research applications include analyzing transcriptional reprogramming mechanisms, investigating metabolic reorganization and regulation, evaluating environmental effects on transition timing, quantifying metabolic markers and dynamics, and assessing pre-lag transition as determinant of lag and ripening progression.
Pre_Lag_Phase_Transition is second aspect of Enlargement_Phase_Completion, distinguished by metabolic reorganization. Represents transcriptional reprogramming with growth genes down 70-90%, lag genes up 3-8 fold. Respiratory rate declining 2-4 → 1.5-2.5 → 1.0-1.8 μmol O₂ g⁻¹ h⁻¹. Malate respiration, sugar accumulation beginning. Secondary metabolism activated. Duration 1-2 weeks. Follows Final_Cell_Expansion. Precedes Growth_Deceleration. Prepares lag phase and ripening. Critical developmental transition.
Lag-Phasen-Vorbereitung
Prä-Lag-Phasen-Übergang
Wachstums-zu-Lag-Wechsel
growth-to-lag shift
lag phase preparation
metabolic reorganization
metabolic transition
metabolische Transition
Pre Lag Phase Transition
Own-rooted Zinfandel vines surviving in Sonoma sandy soils since the 1880s bear Pre_Phylloxera_Clone_Roles, their survival through 140+ years on phylloxera-resistant sites preserving genetic material from initial California plantings. The Bollinger Vieilles Vignes Françaises Pinot Noir bears a Pre_Phylloxera_Clone_Role from ungrafted vines in phylloxera-free sandy soils of Champagne, producing one of the rarest wines worldwide from pre-crisis genetic material. The Nacional selections of Quinta do Noval Port serve in Pre_Phylloxera_Clone_Roles, own-rooted vines surviving in schist soils that proved inhospitable to phylloxera, producing the estate's most celebrated vintage Port from material predating the epidemic.
A heritage clone role that inheres in a grapevine clone verified or credibly claimed to have survived from plantings established before the phylloxera epidemic reached the relevant region (generally 1860s-1900s in Europe, variable elsewhere), representing pre-crisis genetic material that may differ from post-phylloxera reconstitution selections. Pre-phylloxera clones survived the epidemic through various mechanisms—geographic isolation (sandy soils inhospitable to phylloxera, islands, regions the pest never reached), deliberate conservation (protected vines maintained through the crisis), or survival on own roots in resistant sites. These clones represent the genetic diversity present before the population bottleneck and reconstruction that followed phylloxera, potentially preserving genotypes lost during the crisis and subsequent replanting on resistant rootstocks. Pre-phylloxera status confers both historical significance (connection to 19th-century viticulture) and potential practical value (genotypes not selected through post-phylloxera programs may carry unique characteristics). Authentication of pre-phylloxera claims increasingly uses molecular methods, with genetic fingerprinting able to distinguish pre-crisis and post-crisis populations in some varieties.
Pre_Phylloxera_Clone_Role carries particular historical significance given phylloxera's transformative impact on world viticulture. The epidemic forced near-total replanting on resistant rootstocks across Europe and beyond, creating a temporal division in vineyard genetic material—pre-crisis populations destroyed or preserved only in refugia versus post-crisis replanting often using selected material from surviving sources. Pre-phylloxera clones may preserve genetic diversity lost during reconstruction, when replanting from limited surviving sources narrowed genetic representation. Own-rooted pre-phylloxera survivors are exceedingly rare most pre-crisis material persists as grafted vines in protected sites or through conservation programs that documented and preserved surviving genotypes. The designation's rarity and historical resonance command market attention, with wines from authenticated pre-phylloxera vines often commanding significant premiums.
19.-Jahrhundert-Klon
19th century clone
Prä-Reblaus-Klon
antique clone
pre-crisis clone
pre-phylloxera clone
vorkritzlicher Klon
This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Pouget R (1990) Histoire de la lutte contre le Phylloxéra de la vigne en France, INRA Campbell C (2005) The Botanist and the Vintner, Algonquin Books Ordish G (1987) The Great Wine Blight, Sidgwick & Jackson
Pre Phylloxera Clone Role
The pre-phylloxera name Carmenère appears in Bordeaux ampelographic surveys from the 1830s-1860s as a major component of Médoc and Graves plantings comprising 10-15% of vineyard area, documented by Odart (1845) and Rendu (1854) with detailed morphological descriptions — following phylloxera devastation and replanting from the 1880s onward, the cultivar was presumed extinct in Bordeaux until 1994 when DNA analysis by Meredith and colleagues revealed that extensive plantings in Chile labeled as Merlot were in fact Carmenère, preserved through pre-phylloxera exports of cuttings to South America that escaped the European epidemic.
A historical name documented in viticultural sources from the period immediately preceding the phylloxera epidemic (approximately 1750-1870), when European ampelography was being formalized through systematic cultivar description and vineyard surveys, representing the final generation of cultivar designations applied to ungrafted European vineyards before the catastrophic replanting that followed phylloxera devastation.
Pre_Phylloxera_Name captures the nomenclature of the critical transitional period when empirical ampelography was maturing into a systematic discipline but the phylloxera crisis (1863 onward) was about to fundamentally reshape European vineyard composition. The pre-phylloxera cultivar inventory — documented in works by Odart (Ampélographie universelle, 1845), Pulliat (Le vignoble, 1874), and Viala and Vermorel (Ampélographie, 1901-1910) — records many cultivars that subsequently disappeared from European viticulture during post-phylloxera replanting when growers preferentially selected a narrower range of commercially productive varieties for grafted vineyards: an estimated 50-70% of pre-phylloxera cultivar diversity was lost during this period of replanting and consolidation.
19th-century variety name
Prä-Reblaus-Sortenname
Sortenbezeichnung vor der Reblaus
pre-crisis designation
pre-phylloxera cultivar name
vorPhylloxera-Name
Odart AP (1845) Ampélographie universelle, Paris Viala P and Vermorel V (1901-1910) Ampélographie: traité général de viticulture, 7 vols, Masson et Cie Meredith CP (2003) Science as a window into wine history, Proceedings of the 3rd International Symposium on Molecular Markers pp: 71-78
Pre Phylloxera Name
Riesling berries in the Rheingau maintain pre-véraison firmness values of 9.2±1.1 N for approximately 60 days post-fruit set, with the abrupt firmness decline marking véraison onset occurring at approximately 50 days before anticipated harvest, allowing growers to calculate backward from historical data to predict harvest windows with 5-7 day accuracy.
Pre-véraison firmness quality is a softening stage quality that inheres in a grape berry during the developmental period from fruit set through the onset of ripening, characterized by maximum tissue rigidity resulting from intact cell wall architecture, high turgor pressure, and absence of ripening-associated degradation processes. Within the BFO framework, this quality represents the baseline mechanical state against which all subsequent softening is measured, existing as a relatively stable plateau during the herbaceous growth phase when berries function as photosynthetically active organs accumulating organic acids and maintaining structural integrity for seed protection. The ontological distinction of pre-véraison firmness lies in its mechanistic basis, where rigid cellulose-hemicellulose networks remain fully cross-linked, pectin polymers maintain high degrees of methylesterification preventing enzymatic attack, and cell turgor contributes maximally to tissue resistance due to active water uptake driven by organic acid accumulation. Instrumentally, pre-véraison berries typically exhibit compression forces of 6-12 N depending on cultivar and berry size, with characteristic elastic behavior showing high recovery after deformation and minimal permanent damage below failure thresholds. This firmness quality correlates with the berry's role as a nutrient sink during seed development, where structural integrity ensures protection of developing embryos from pathogen invasion and physical damage. The transition out of this quality state occurs abruptly at véraison onset, marked by ethylene sensitivity acquisition, abscisic acid accumulation, and transcriptional activation of cell wall modifying enzymes. Pre-véraison firmness assessment provides baseline data for softening rate calculations, enables identification of abnormal development patterns, and supports rootstock and clone evaluation for mechanical harvest suitability.
Pre_Veraison_Firmness_Quality represents the initial state in the softening progression, ontologically distinct from subsequent stages by its mechanistic basis in intact cell wall architecture rather than progressive degradation. This quality corresponds to BBCH stages 71-79 (fruit development) prior to véraison onset. Berry selection for measurement requires careful staging to exclude early-softening berries that may indicate asynchronous development.
Grünbeerenfestigkeit
Hartbeerenstadium
Unreifefestigkeit
Vorvéraison-Festigkeit
green berry firmness
hard berry stage
herbaceous phase firmness
pre-ripening firmness
Castellarin SD et al. (2016) Colour onset in grape berries and its relationship with berry composition. Plant Physiology and Biochemistry 99: 1-10 Matthews MA et al. (1987) Phenologic and growth responses to early and late season water deficits in Cabernet franc. Vitis 26(3): 147-160
Pre Veraison Firmness Quality
Merlot in Bordeaux shows pre-veraison lag during early September Year N+1 at BBCH 81, with ripening genes highly expressed including anthocyanin pathway CHS 68 fold, DFR 82 fold, UFGT 95 fold, sugar transporters VvHT1 24 fold, softening enzymes polygalacturonase 22 fold, enzyme activities 35-48% ripening maxima, abscisic acid increasing from 120 to 380 ng/g over 5 days, earliest berries showing trace softening and faint color, sugars 14%, acids 0.8%, pH 3.2, under oceanic conditions of 20-25°C temperature, typical of this mid-season cultivar approaching veraison
Pre-veraison lag represents the late lag phase period during berry growth lag phase when berry approaches veraison transformation, ripening preparation intensifies with accelerating gene expression and enzyme accumulation, metabolic reprogramming shifts berry decisively toward ripening state, abscisic acid levels begin increasing signaling veraison approach, and berry positions at developmental threshold ready for veraison triggering, occurring during final 3-7 days of lag phase as veraison becomes imminent. This late lag transition period occurs during late summer to early autumn in late August-early September in temperate regions Year N+1, proceeding over final days to 1 week before veraison onset, corresponding to BBCH 81 late beginning of ripening stage immediately pre-veraison. Pre-veraison lag mechanism involves veraison preparation intensification with ripening gene expression accelerating to high pre-transformation levels, metabolic reprogramming establishing ripening-competent state, abscisic acid synthesis and accumulation beginning hormone spike, berry compositional and structural preparations finalizing, developmental competence achieving transformation-ready state, and integrated preparations positioning berry at veraison threshold. Veraison preparation intensification shows ripening gene expression reaching 30-60 fold increases relative to basal approaching veraison activation levels, enzyme accumulation achieving 30-50% eventual ripening maximum activity, metabolic flux capacity building to support rapid transformation, and preparation reaching critical threshold enabling veraison. Ripening gene expression acceleration shows anthocyanin biosynthesis genes in red cultivars reaching 30-80 fold increases with pathway genes highly expressed, sugar transporter genes reaching 15-30 fold increases preparing maximal import capacity, softening enzyme genes reaching 10-25 fold increases enabling rapid texture transformation, and coordinated expression at near-activation levels. Anthocyanin pathway expression shows phenylalanine ammonia lyase PAL 25-60 fold, chalcone synthase CHS 35-90 fold, dihydroflavonol reductase DFR 40-100 fold, UDP-glucose flavonoid glucosyltransferase UFGT 50-120 fold relative to basal, and pathway primed for color development. Sugar transporter expression shows VvHT1 hexose transporter 15-35 fold, VvHT2 12-28 fold, sucrose transporters possibly increasing, and transporter expression enabling accelerated import. Softening enzyme expression shows polygalacturonase 12-30 fold, pectin methylesterase 10-25 fold, expansin 6-18 fold, and softening machinery ready for activation. Enzyme activity levels shows ripening-associated enzymes reaching 30-50% eventual maximum activity from accumulated proteins, anthocyanin biosynthesis enzymes with substantial activity though not maximal, sugar transport capacity increased though not peak, softening enzymes with activity enabling initiation, and activity levels at functional pre-veraison state. Metabolic reprogramming shows decisive metabolic shift toward ripening program, respiratory patterns changing toward ripening characteristics, carbon flux redirecting toward ripening processes, secondary metabolism intensifying preparing for transformation, and metabolic state establishing ripening competence. Respiratory metabolism changes shows possible respiratory rate beginning slight increase from lag 1.0-1.5 μmol O₂ g⁻¹ h⁻¹ toward pre-veraison 1.2-1.8 μmol O₂ g⁻¹ h⁻¹, metabolic activity increasing supporting preparation, and respiratory shift as metabolic indicator. Carbon metabolism shows increased flux toward phenolic and sugar metabolism, organic acid metabolism shifting toward ripening decline patterns, metabolic pathways reorganized for ripening function, and carbon allocation favoring ripening processes. Secondary metabolism shows phenolic biosynthesis intensifying, flavonoid accumulation accelerating in preparation for anthocyanin synthesis, possible early anthocyanin synthesis in some red cultivar berries showing earliest color, and secondary metabolism as pre-veraison feature. Abscisic acid dynamics shows ABA beginning increase from lag basal 50-200 ng/g toward spike levels, ABA synthesis initiating in berry tissues, hormone accumulation accelerating during pre-veraison lag, eventual ABA spike to 500-2000 ng/g coinciding with or triggering veraison, and ABA as critical veraison signal. Abscisic acid synthesis shows ABA biosynthesis genes upregulating during pre-veraison lag, biosynthetic enzymes accumulating, ABA production increasing in berry pericarp and possibly seeds, and synthesis as hormone accumulation mechanism. Abscisic acid signaling shows possible ABA receptor expression increasing, signal transduction components activating, berry sensitivity to ABA increasing preparing for response, and signaling preparing veraison trigger mechanism. Berry composition during pre-veraison lag shows water 78-82% stable, organic acids 0.7-1.0% declining toward veraison levels, sugars 12-16% preparing for acceleration, pH 3.1-3.3 rising toward veraison range, and composition approaching transformation starting point. Organic acid pre-veraison shows malate declining slowly 0.6-0.9%, tartrate stable 0.3-0.6%, total acids approaching veraison onset levels, and acid trajectory toward ripening decline. Sugar pre-veraison shows sugars 12-16% establishing base for accelerated accumulation, limited continued accumulation 0.5-1.0% per week, sugar preparing for veraison import increase, and sugar dynamics at pre-acceleration state. Berry morphology during pre-veraison shows berry maintaining size stability, possible very earliest softening in some berries, earliest color change in some red cultivar berries showing trace anthocyanin, overall appearance still hard green berry approaching transformation, and morphology at threshold state. Earliest softening shows possible initial texture changes in some berries, firmness beginning decline from peak, cell wall modification initiating in earliest berries, and softening as earliest veraison indicator. Earliest color in red cultivars shows possible trace anthocyanin accumulation in earliest berries, very faint color change from green, earliest visible transformation indicator, and color as veraison onset marker. Berry growth rate during pre-veraison shows possible slight growth rate increase from lag minimal <5 mg per day to 5-15 mg per day in some cases, growth rate still low but possibly increasing, and rate change as possible pre-veraison indicator. Environmental effects on pre-veraison lag include temperature affecting progression rate with warm conditions accelerating preparation and shortening pre-veraison period, cool conditions extending pre-veraison lag, water availability affecting stress signaling possibly influencing ABA dynamics, and environmental factors determining veraison timing. Temperature effects shows heat accumulation advancing pre-veraison preparations, growing degree days correlating with veraison timing, warm conditions promoting faster progression through pre-veraison lag to veraison onset, and temperature as major timing determinant. Water stress effects shows moderate deficit possibly enhancing ABA production accelerating veraison, stress as potential veraison advance mechanism, and water influencing hormone dynamics. Pre-veraison lag duration shows typically 3-7 days before veraison onset, variable duration depending on preparation state and environmental conditions, eventual transition to veraison when preparations complete and trigger occurs, and brief period marking imminent transformation. Developmental competence shows berry achieving transformation-ready state, accumulated ripening machinery adequate for veraison progression, metabolic preparations complete enabling rapid transformation, and competence as pre-veraison outcome. The process shows two distinguishable aspects including veraison preparation intensification with accelerating ripening gene expression and enzyme accumulation, and metabolic reprogramming establishing ripening-competent state. Duration typically 3-7 days representing late lag immediately pre-veraison. Regional timing shows pre-veraison lag during late August-early September across climates immediately before veraison. Viticultural significance includes representing imminent veraison approach with berry at transformation threshold, determining veraison timing through preparation completion, demonstrating ripening readiness state, and enabling rapid veraison progression when triggered. Research applications include analyzing veraison preparation completion and readiness, investigating abscisic acid dynamics and veraison triggering, evaluating metabolic competence for transformation, quantifying gene expression and enzyme activity thresholds, and assessing pre-veraison lag as determinant of veraison onset timing and progression.
Pre_Veraison_Lag represents late lag phase within Berry_Growth_Lag_Phase. Subdivided into Veraison_Preparation with intensified gene expression (30-80 fold anthocyanin, 15-30 fold transporters, 10-25 fold softening enzymes) and enzyme accumulation (30-50% maxima), and Metabolic_Reprogramming establishing ripening competence. Duration 3-7 days immediately pre-veraison. ABA beginning increase 50-200 → 380-800 ng/g. Earliest softening and color in some berries. Growth rate possibly 5-15 mg/day. Precedes Lag_Phase_Termination. Transformation threshold.
Prä-Veraison-Lag
Veraison-Schwelle
bevorstehende Veraison
imminent veraison
late lag phase
späte Lag-Phase
transformation approach
veraison threshold
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 460-470 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Pre Veraison Lag
Pinot Noir pre-veraison late berries at late BBCH 77 to early BBCH 81 (weeks 7.5-9 post-fruit set) maintain berry dimensions at 9.5-10 mm diameter and 1.1-1.25 g fresh weight with minimal change from mid-lag phase, green coloration with chlorophyll declining from 0.6 mg/g (week 7) to 0.4 mg/g (week 8.5), firm texture at 4-4.5 N, sugars increasing slightly from 8% to 10.5%, acidity stable at 22-24 g/L, with seeds completing maturation at 5.8-6.5 mm length and 24-27 mg dry weight, and with molecular markers showing ABA increasing from 15 ng/g (mid-lag phase) to 35 ng/g (pre-veraison late) and ripening gene expression (sugar transporters, anthocyanin biosynthesis genes) increasing 2-4 fold, indicating imminent veraison within 5-10 days though berries still appear phenotypically green and lag-phase-like. Syrah pre-veraison late berries examined through transcriptome analysis show upregulation of 850+ genes associated with ripening processes (including VvMYBA1 anthocyanin regulator increasing 8-fold, VvSUC11/27 sucrose transporters increasing 5-fold, cell wall modification genes increasing 3-6 fold) during the 2-week pre-veraison late period (weeks 7.5-9) before visible color change begins, demonstrating extensive molecular preparation for veraison occurring during final lag phase while berries remain phenotypically green. Cabernet Franc pre-veraison late berries under two water regimes show well-watered vines reaching pre-veraison late state (ABA 30-40 ng/g, ripening genes upregulated 3-5 fold) at 65-70 days post-fruit set with veraison color change beginning at 68-75 days, while moderate water deficit vines reach comparable pre-veraison late molecular state at 58-63 days with veraison at 60-68 days, demonstrating water deficit advancement of veraison timing by approximately 1 week through earlier achievement of molecular ripening readiness during pre-veraison late period. Chardonnay pre-veraison late berries show characteristic slight sugar increase from 8.5% at early lag phase (week 6.5) to 10% at pre-veraison late (week 8.5) representing the initial sugar accumulation phase preceding the rapid accumulation during veraison and ripening, with the pre-veraison late sugar increase (1.5% over 2 weeks, 0.75% per week) intermediate between lag phase stability (<0.3% per week) and active ripening rapid accumulation (0.8-1.5% per week), serving as biochemical indicator of approaching veraison.
A pre-veraison late berry is a late developing berry in the final portion of the lag phase immediately preceding veraison onset (late BBCH 77 to early BBCH 81, approximately 7.5-9 weeks post-fruit set), representing berries completing lag phase preparation for the veraison transition while still maintaining green coloration and lag phase characteristics, with 'late' referring to late lag phase rather than late ripening phase. Pre-veraison late berries are characterized by= (1) berry dimensions at or near maximum Phase I size typically 8-12 mm diameter and 0.8-1.8 g fresh weight with minimal size changes during the pre-veraison late period (2) cells at final number established during Phase I with cell diameter at 50-75 μm maintaining lag phase size, poised for expansion but not yet expanding (3) persistent green coloration with chlorophyll still present (0.3-0.6 mg/g) though often declining from peak lag phase levels (0.5-0.8 mg/g) in preparation for veraison (4) firm texture maintaining lag phase hardness (3.5-5 N) with no softening (5) sugars at 8-11% showing slight increases from early lag phase (7-9%) as veraison approaches but not yet showing rapid accumulation characteristic of veraison (6) acidity at maximum or slightly declining from peak with titratable acidity 20-24 g/L, malic acid at peak concentration (7) seeds completing final maturation with embryos fully developed, seed coats fully lignified and brown, seeds at or approaching maximum dry weight (22-28 mg per seed). Pre-veraison late berries show molecular and biochemical preparation for imminent veraison= transcriptional changes with ripening-related genes increasingly upregulated while growth genes decline ABA levels beginning to increase from lag phase baseline creating hormonal conditions promoting veraison onset cell wall modification enzymes beginning to increase in expression preparing for softening sugar transporter genes upregulating preparing for sugar accumulation anthocyanin biosynthesis genes (in red cultivars) beginning expression though anthocyanin accumulation not yet evident. Pre-veraison late berries represent the final pre-ripening state where berry appears phenotypically unchanged from earlier lag phase but molecular preparations for veraison are advancing, creating a brief transitional period (typically 1-2 weeks) before veraison color change becomes visible. Environmental conditions during pre-veraison late berry period influence veraison timing= water deficit can advance veraison onset by 3-10 days through earlier ABA accumulation and stress signaling temperature affects veraison timing with warmer conditions promoting earlier veraison crop load influences resource availability potentially affecting veraison synchrony with high crop loads sometimes creating delayed irregular veraison. Pre-veraison late berries can be distinguished from earlier lag phase berries through biochemical markers (increasing ABA, increasing expression of ripening genes) though phenotypically they appear similar, and can be predicted to undergo veraison within 1-2 weeks based on developmental timing and molecular markers.
Pre_Veraison_Late_Berry represents the terminal lag phase immediately before veraison (late BBCH 77 to early BBCH 81, weeks 7.5-9) and is distinguished from Maximum_Size_Late_Berry (peak ripening phase, weeks 14-18) by pre-veraison versus mid-to-late ripening state. The 'late berry' terminology is somewhat ambiguous as it could suggest late ripening, but in this context refers to late lag phase. Pre-veraison late berries show molecular preparation for veraison while maintaining lag phase phenotype, representing a transitional state of high interest for understanding veraison regulation. Understanding pre-veraison late berry biology is relevant for predicting veraison timing, for studying veraison molecular control, and for managing practices affecting veraison onset.
Späte-Lagphase-Stadium
Vor-Farbumschlag-Beere
late lag-phase berry
pre-color-change berry
prä-Véraison späte Beere
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 192-198 Castellarin, S.D. et al. (2007). Water deficits accelerate ripening and induce changes in gene expression. Functional Plant Biology 34=174-186.
Pre Veraison Late Berry
Syrah in Rhône Valley shows pre-veraison metabolism during late August-September Year N+1 at BBCH 81, with anthocyanin biosynthesis genes PAL increasing 14 fold, CHS 22 fold, DFR 26 fold, UFGT 35 fold over 2.5 week lag phase, sugar transporters VvHT1 increasing 9 fold, VvHT2 8 fold, softening enzymes polygalacturonase 7 fold, pectin methylesterase 6 fold, enzyme activities reaching 15-28% ripening maxima, sugars increasing from 12% to 15%, pH rising from 3.0 to 3.2, under warm Mediterranean conditions of 24-28°C temperature, typical of this mid-to-late season cultivar preparing for veraison
Pre-veraison metabolism represents the ripening preparation phase during minimal growth period when berry metabolism progressively builds ripening capacity, ripening genes increase expression 5-20 fold preparing for veraison activation, ripening enzymes accumulate enabling rapid transformation, metabolic pathways prime for accelerated sugar accumulation and color development, and metabolic preparations position berry for veraison transformation when triggered, proceeding over 1-3 weeks during lag phase as critical pre-ripening function. This ripening preparation phase occurs during late summer to early autumn in late August-September in temperate regions Year N+1, proceeding over 1-3 weeks during minimal growth period, corresponding to BBCH 81 beginning of ripening stage pre-veraison period. Pre-veraison mechanism involves progressive ripening gene expression increase building transcriptional capacity, ripening enzyme accumulation preparing functional capacity, metabolic pathway component assembly enabling rapid activation, berry compositional preparation creating appropriate substrate conditions, and integrated preparations positioning berry for veraison transformation. Ripening gene expression progression shows anthocyanin biosynthesis genes in red cultivars progressively increasing 5-20 fold over lag phase from early initiation 2-8 fold to late lag 10-40 fold relative to basal, sugar transporter genes increasing 5-15 fold preparing import capacity, softening enzyme genes increasing 3-10 fold accumulating modification capacity, flavor and aroma biosynthesis genes beginning expression, and progressive expression building ripening machinery. Anthocyanin biosynthesis gene progression shows phenylalanine ammonia lyase PAL increasing 8-18 fold, chalcone synthase CHS increasing 10-25 fold, dihydroflavonol reductase DFR increasing 12-30 fold, UDP-glucose flavonoid glucosyltransferase UFGT increasing 15-40 fold, and coordinated pathway expression preparing for color development. Anthocyanin pathway enzyme accumulation shows enzyme proteins increasing from gene expression, pathway flux capacity building, pathway poised for rapid activation at veraison, and enzyme accumulation as preparation function. Sugar transporter gene progression shows hexose transporter VvHT1 increasing 5-12 fold, VvHT2 increasing 4-10 fold, sucrose transporter VvSUC11, VvSUC12 possibly increasing, and transporter expression preparing import capacity. Sugar transporter protein accumulation shows transporter proteins inserting in plasma membrane and tonoplast, transport capacity increasing progressively, membrane positioned for rapid sugar import at veraison, and protein accumulation enabling accelerated uptake. Cell wall modification gene progression shows polygalacturonase increasing 4-9 fold, pectin methylesterase increasing 3-8 fold, expansin genes increasing 2-6 fold, and softening enzyme expression preparing texture transformation. Cell wall enzyme accumulation shows enzyme proteins accumulating from expression, pectin degradation and modification capacity building, enzymes positioned for rapid softening at veraison, and accumulation as preparation process. Flavor and aroma gene expression shows terpene synthase genes beginning expression in aromatic cultivars, volatile ester biosynthesis genes increasing, methoxypyrazine genes declining in some cultivars, and flavor biosynthesis preparation. Metabolic pathway preparation shows phenylpropanoid pathway components accumulating, glycolysis and sugar metabolism enzymes adjusting for increased flux, organic acid metabolism enzymes shifting toward ripening patterns, and pathway preparations enabling rapid metabolic shifts. Secondary metabolism preparation shows flavonoid and stilbene biosynthesis pathways active, phenolic accumulation continuing, pathway capacity building for ripening secondary metabolite synthesis, and secondary metabolism as active pre-veraison process. Enzyme activity progression shows ripening-associated enzymes gradually increasing activity from accumulating proteins, activities reaching 10-30% eventual ripening maxima by late lag, enzyme activation preparing for full veraison function, and progressive activity as preparation indicator. Berry composition preparation shows composition trajectory toward ripening-favorable conditions, acids declining to 0.7-1.1% creating pH increase, sugars increasing to 12-16% establishing substrate base, pH rising to 3.1-3.3 enabling optimal enzyme activities, and composition changes preparing for ripening. Acid decline preparing pH shows organic acids declining from respiration and possibly transport, pH increasing from 3.0-3.2 to 3.1-3.3 over lag phase, pH increase enabling pH-dependent enzyme optimal function during ripening, and pH trajectory as preparation aspect. Sugar accumulation preparing substrate shows limited sugar increase during lag providing substrate base, vacuole hexose concentrations increasing preparing for accelerated accumulation, sugar establishing osmotic base for ripening water relations, and sugar dynamics as preparation function. Membrane preparation shows plasma membrane and tonoplast modifications preparing for altered transport, transporter protein accumulation, possible membrane lipid composition changes, and membrane preparations enabling ripening flux changes. Metabolic readiness state shows berry metabolism poised for rapid transformation at veraison trigger, accumulated enzymes and pathway components ready for activation, metabolic preparations creating low activation energy for transition, and readiness as pre-veraison outcome. Veraison triggering preparation shows berry prepared to respond rapidly when veraison signal occurs, possible abscisic acid sensitivity increasing preparing for hormone trigger, developmental competence established enabling transformation, and preparation ensuring rapid veraison progression when initiated. Abscisic acid role preparation shows berry possibly increasing ABA sensitivity during lag, ABA receptors potentially upregulating, signal transduction components preparing, and ABA responsiveness as potential trigger mechanism. Environmental effects on pre-veraison metabolism include temperature affecting metabolic progression rate with warm conditions accelerating preparation and cool conditions slowing progression, water availability affecting stress response and possibly veraison timing, moderate deficit possibly enhancing preparations, and environmental factors influencing metabolic dynamics. Temperature effects show heat accumulation advancing metabolic preparations, growing degree days correlating with ripening gene expression and enzyme accumulation, warm conditions shortening lag through faster preparation, and temperature as major preparation rate determinant. Water stress effects show moderate deficit possibly triggering abscisic acid increase advancing preparations, stress potentially accelerating veraison onset through enhanced readiness, and water as factor affecting pre-veraison dynamics. Pre-veraison metabolism duration shows progressive preparations over 1-3 week lag phase, preparation intensity increasing toward late lag, eventual veraison onset when preparations complete and trigger occurs, and duration determining veraison readiness. Pre-veraison metabolism assessment shows gene expression profiling documenting progressive increases, enzyme activity assays quantifying accumulation, metabolite analysis showing compositional preparations, and molecular markers characterizing pre-veraison state. The process represents third and final aspect of minimal growth period emphasizing ripening preparation, following near-zero growth rate and seed development focus. Duration proceeding over 1-3 weeks during minimal growth period. Regional timing shows pre-veraison metabolism during late August-September across climates preparing for veraison. Viticultural significance includes representing critical ripening preparation determining transformation capacity, building metabolic machinery enabling rapid veraison progression, positioning berry for developmental transition, and determining ripening success through adequate preparation. Research applications include analyzing ripening gene expression regulation and progression, investigating enzyme accumulation and metabolic pathway preparation, evaluating environmental effects on pre-veraison metabolism and veraison timing, quantifying metabolic readiness and preparation dynamics, and assessing pre-veraison metabolism as determinant of veraison timing, progression, and ripening quality.
Pre_Veraison_Metabolism is third and final aspect of Minimal_Growth_Period, distinguished by ripening preparation. Represents progressive ripening gene increases: anthocyanin pathway 5-20 fold early to 10-40 fold late lag, sugar transporters 5-15 fold, softening enzymes 3-10 fold. Enzyme activities 10-30% eventual maxima. Metabolic pathway priming. Berry compositional preparation (acids declining, sugars 12-16%, pH 3.1-3.3). Duration 1-3 weeks. Follows Near_Zero_Growth_Rate and Seed_Development_Focus. Completes Minimal_Growth_Period and Berry_Growth_Lag_Phase. Determines veraison readiness.
Prä-Reifungs-Metabolismus
Prä-Veraison-Metabolismus
Reifungs-Vorbereitung
Veraison-Bereitschaft
metabolic priming
pre-ripening metabolism
ripening preparation
veraison readiness
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 458-465 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Pre Veraison Metabolism
Precipitation data for Marlborough, New Zealand documents annual totals of 650-750 mm with 300-350 mm during the October-April growing season, characterizing the region's relatively dry maritime climate that favors Sauvignon Blanc aromatic intensity. In contrast, precipitation data for the Mosel Valley records 700-800 mm annually but with 350-400 mm concentrated in the May-October period, where the combination with steep slate slopes ensures rapid drainage critical for Riesling quality.
Precipitation data is a terroir information entity that documents the quantity, timing, form, and distribution of atmospheric water deposition received by a viticultural site, serving as a primary determinant of vine water status, disease pressure, and fruit quality potential. This information entity encompasses all measured and derived precipitation parameters including rainfall totals, seasonal distributions, intensity metrics, and drought indices that collectively characterize the hydrological input component of a vineyard's climate profile, enabling assessment of irrigation requirements and vintage quality prediction.
Precipitation data constitutes one of the three fundamental climate variables (alongside temperature and radiation) that define viticultural potential of any site. The information entity bridges raw meteorological measurements with viticultural interpretation, as identical precipitation totals carry vastly different implications depending on seasonal timing, intensity patterns, and soil water-holding capacity. Modern precision viticulture increasingly relies on high-resolution precipitation data from on-site weather stations rather than regional averages, as topographic effects can create substantial rainfall gradients even within single vineyard blocks. The interaction between precipitation and canopy management decisions (leaf removal timing, shoot positioning) represents a key area where this data directly informs viticultural practice.
Niederschlagsdaten
Niederschlagsinformation
Regendaten
pluviometric data
pluviometrische Daten
precipitation record
rainfall data
vineyard precipitation information
Jones GV, Webb LB (2010) Climate change, viticulture, and wine: challenges and opportunities. Journal of Wine Research 21(2-3): 103-106 Gladstones J (2011) Wine, Terroir and Climate Change, Wakefield Press, pp.: 119-145 Van Leeuwen C, Darriet P (2016) The impact of climate change on viticulture and wine quality. Journal of Wine Economics 11(1): 150-167
Precipitation Data
Precipitation distribution in Bordeaux follows an oceanic pattern with relatively uniform monthly rainfall of 50-80 mm throughout the year, creating persistent disease pressure but ensuring adequate vine water supply without irrigation. The Central Valley of Chile exhibits extreme Mediterranean distribution with 250-350 mm concentrated in May-August (winter) and virtually zero rainfall from November through March, enabling complete control of vine water status through irrigation scheduling. Tokaj, Hungary displays a continental distribution pattern with a summer rainfall peak of 60-80 mm/month in June-July, transitioning to drier autumn conditions (30-40 mm/month October-November) that favor the development of Botrytis cinerea for Aszú production.
Precipitation distribution is a precipitation data entity that characterizes the temporal pattern, seasonality, and intensity profile of atmospheric water deposition across monthly, seasonal, and phenological intervals at a viticultural site, documenting when and how precipitation is delivered rather than merely the cumulative total, and serving as a critical determinant of viticultural management strategy and vintage variability.
Precipitation distribution is frequently more important for viticultural outcomes than total precipitation, as it determines the timing of water availability relative to vine phenological demand and disease susceptibility windows. The distinction between Mediterranean (dry summer/wet winter), oceanic (uniform), continental (summer maximum), and monsoon (late summer peak) distribution patterns fundamentally shapes viticultural practice within each region. Key viticultural metrics derived from distribution data include the proportion of annual rainfall occurring during the growing season, the maximum consecutive dry days during berry development, and the frequency of rainfall events exceeding intensity thresholds (>10 mm/hour) that promote erosion and disease. Monthly precipitation coefficients of variation provide additional insight into vintage reliability and the predictability of management schedules.
Niederschlagsmuster
Niederschlagsregime
Niederschlagsverteilung
Regenverteilung
precipitation regime
rainfall distribution pattern
rainfall seasonality
seasonal precipitation pattern
temporal precipitation profile
zeitliche Niederschlagsverteilung
Gladstones J (2011) Wine, Terroir and Climate Change, Wakefield Press, pp.: 125-140 Jones GV (2006) Climate and terroir: impacts of climate variability and change on wine. In Fine Wine and Terroir, Geological Society of America Special Paper, pp.: 1-14 Fraga H et al. (2012) Climatic suitability of Portuguese grapevine varieties and climate change adaptation. International Journal of Climatology 34(1): 1-14
Precipitation Distribution
In Gewürztraminer berries in Alto Adige, Italy during BBCH 83-89, precursor formation accumulates total glycosidically-bound monoterpenes from 1.2 to 8.5 mg/L over 38 days, with bound linalool increasing from 0.3 to 2.8 mg/L and bound geraniol from 0.2 to 1.9 mg/L, representing 92% of total monoterpenoid content at harvest compared to 8% present as free volatiles, under warm-day cool-night conditions of 28°C/14°C.
Precursor formation encompasses the biosynthetic processes in Vitis vinifera berries during ripening wherein metabolic intermediates and conjugated compounds that serve as substrates for flavor compound generation are synthesized and accumulated, establishing the biochemical foundation for flavor expression during winemaking when enzymatic, chemical, and fermentative transformations release flavor-active volatiles from their bound precursor forms. This process involves the accumulation of diverse precursor classes including cysteine and glutathione conjugates of volatile thiols that generate tropical fruit aromas upon yeast-mediated cleavage during fermentation, glycosidically-bound monoterpenes and norisoprenoids that release floral and fruity volatiles through acid or enzyme hydrolysis, and amino acid pools that serve as substrates for Ehrlich pathway fermentation products and Maillard reaction participants during wine production and aging. The temporal progression of precursor formation shows characteristic accumulation patterns with initial synthesis of conjugating molecules including glutathione and UDP-glucose during early post-veraison stages, followed by active conjugate formation during mid-ripening as free volatile concentrations increase and glycosyltransferase activities peak, and continued accumulation through late ripening establishing the precursor pools that determine wine aromatic potential independent of free volatile concentrations at harvest. Cellular localization of precursor formation varies with compound class, with thiol precursors accumulating predominantly in mesocarp parenchyma cells where cysteine and glutathione conjugation occurs in cytosolic compartments, while glycosidic precursors concentrate in exocarp cells where monoterpene glycosyltransferases show highest expression and vacuolar sequestration protects conjugates from hydrolytic enzymes. The quantitative significance of precursor formation reflects the observation that 80-95% of total terpenoids and 60-90% of total volatile thiols exist in conjugated precursor forms at harvest, with free volatile concentrations representing only a fraction of total aromatic potential that can be released through appropriate winemaking interventions. Temperature during ripening influences precursor formation through effects on conjugating enzyme activities and precursor stability, with moderate conditions favoring balanced accumulation while high temperatures may accelerate precursor degradation or shift conjugation patterns affecting subsequent release efficiency. Cultivar genetics determine precursor formation capacity through variety-specific expression of glycosyltransferases, glutathione S-transferases, and related conjugating enzymes, with aromatic varieties showing 3-10 fold higher precursor concentrations than neutral varieties at equivalent ripeness levels.
Precursor_Formation is parent class under Flavor_Development representing biosynthesis of conjugated and bound flavor compounds. Distinguishes from Flavor_Compound_Synthesis which generates free volatiles. Contains subclasses Flavor_Precursor_Synthesis, Amino_Acid_Accumulation, and Precursor_Pool_Development. Key characteristic is accumulation of compounds requiring transformation for flavor release. Typically 80-95% of terpenoids exist as precursors.
Aroma-Präkursor-Akkumulation
Aroma-Vorläufer-Synthese
Vorläufer-Bildung
aroma precursor synthesis
bound flavor formation
conjugate precursor development
flavor precursor accumulation
gebundene Aroma-Bildung
Tominaga T et al (1998) Identification of cysteinylated aroma precursors in grape juice. Journal of Agricultural and Food Chemistry 46: 5215-5219 Gunata Z et al (1985) The aroma of grapes. Extraction and determination of free and glycosidically bound fractions. Journal of Chromatography 331: 83-90 Keller M (2020) The Science of Grapevines 3rd Ed pp 295-305
Precursor Formation
In Albariño berries in Rías Baixas, Galicia during BBCH 81-89, precursor pool building establishes glycosidically-bound monoterpene pools increasing from 1.8 to 6.5 mg/L, norisoprenoid precursor pools from 35 to 165 µg/L, and thiol precursor pools from 85 to 295 µg/L over 48 days, representing total aromatic potential 8.2-fold greater than free volatile content, with pool accumulation following biphasic pattern showing rapid increase during weeks 2-5 and plateau during final 2 weeks before harvest under cool Atlantic conditions averaging 19°C.
Precursor pool building describes the cumulative establishment of aromatic precursor reservoirs within Vitis vinifera berries during the ripening period, wherein the integrated outputs of glycosylation, cysteine conjugation, and other precursor-forming reactions create increasingly substantial pools of bound aromatic compounds that determine the total aromatic capacity available for release during winemaking, representing the culmination of precursor biosynthetic processes at harvest maturity. This process integrates multiple precursor classes into composite aromatic potential, with glycosidically-bound monoterpenes contributing pools of 2-15 mg/L in aromatic varieties, glycosidically-bound norisoprenoids adding pools of 50-300 µg/L across diverse cultivars, and cysteine-conjugated volatile thiol precursors accumulating to pools of 100-500 µg/L in Sauvignon Blanc and related varieties, collectively establishing the raw material for aromatic expression during subsequent vinification. The dynamics of precursor pool building show variety-specific accumulation trajectories, with some cultivars demonstrating linear accumulation throughout ripening while others exhibit biphasic patterns with rapid early-phase accumulation followed by plateau phases during late ripening, reflecting genetic differences in biosynthetic capacity, substrate availability, and potential precursor turnover through degradative processes. Vacuolar storage capacity influences precursor pool building through the physical constraints on glycoside accumulation within finite intracellular compartments, with evidence suggesting that vacuolar transport saturation may limit maximum achievable pool sizes in some varieties, particularly during rapid accumulation phases when biosynthetic output may exceed transport and storage capacity. The relationship between precursor pool size at harvest and wine aromatic quality depends on multiple factors including release efficiency during winemaking, potential synergistic or masking interactions among released compounds, and threshold concentrations required for sensory perception, with larger pools generally correlating with greater aromatic potential though release optimization remains critical for converting precursor content to sensory impact. Environmental factors including temperature, light exposure, and water status influence precursor pool building through effects on both biosynthesis of free volatiles and subsequent conjugation reactions, with growing conditions affecting final pool sizes by 50-200% relative to baseline levels in controlled studies comparing different environmental treatments. Viticultural practices targeting precursor pool building include canopy management optimizing cluster microclimate for secondary metabolism, irrigation strategies imposing controlled water deficit that enhances precursor accumulation, and harvest timing decisions capturing maximum pool development before potential late-season degradation or dilution through continued berry expansion.
Precursor_Pool_Building is subclass of Aroma_Precursor_Formation representing cumulative reservoir establishment. Distinguishes from individual precursor synthesis by emphasizing integrated accumulation. Sibling to Glycosidic_Precursor_Synthesis and Bound_Aroma_Accumulation. Integrates multiple precursor classes into composite potential. Pool size determines aromatic capacity for winemaking. Variety-specific accumulation trajectories.
Konjugat-Pool-Akkumulation
Vorläufer-Pool-Aufbau
aromatic precursor reservoir development
aromatische Vorläufer-Reservoir-Entwicklung
bound aroma pool establishment
conjugate pool accumulation
gebundener Aroma-Pool-Etablierung
precursor capacity building
Williams PJ et al (1989) Glycosidic precursors of varietal grape and wine flavor. ACS Symposium Series 388: 35-48 Tominaga T et al (1998) Identification of cysteinylated aroma precursors. Journal of Agricultural and Food Chemistry 46: 5215-5219 Keller M (2020) The Science of Grapevines 3rd Ed pp 295-305
Precursor Pool Building
In Riesling berries in Rheingau, Germany during BBCH 81-89, precursor pool development establishes glycosidically-bound monoterpene pools increasing from 1.8 to 9.2 mg/L over 50 days, with bound linalool pool reaching 3.1 mg/L, bound alpha-terpineol 1.8 mg/L, and bound hotrienol precursors 0.9 mg/L, representing total aromatic potential 11-fold greater than free volatile concentrations of 0.82 mg/L at harvest under cool conditions averaging 16°C that favor precursor preservation.
Precursor pool development describes the progressive establishment and expansion of conjugated and bound flavor compound reservoirs within Vitis vinifera berries during ripening, wherein the cumulative biosynthesis of glycosidic, cysteinylated, and glutathionylated precursors creates increasingly substantial pools of potential flavor compounds that determine the aromatic capacity of harvested grapes for subsequent flavor release during winemaking processes. This process integrates the outputs of multiple precursor biosynthetic pathways, with glycosidically-bound monoterpenes accumulating to pools of 2-15 mg/L in aromatic varieties representing 85-95% of total terpenoid content, cysteine-conjugated volatile thiol precursors reaching pools of 100-500 µg/L in Sauvignon Blanc and related varieties, and glycosidically-bound norisoprenoids including beta-damascenone and beta-ionone precursors accumulating to pools of 50-200 µg/L across diverse cultivars. The temporal dynamics of precursor pool development show variety-specific patterns with some cultivars exhibiting continuous linear accumulation throughout ripening while others demonstrate biphasic patterns with rapid early accumulation followed by plateau phases, reflecting differences in biosynthetic enzyme expression profiles and potential precursor turnover through hydrolytic or oxidative degradation. Cellular organization of precursor pools involves vacuolar sequestration as the primary storage mechanism, with tonoplast transport systems actively accumulating glycosidic conjugates against concentration gradients, establishing stable intracellular reservoirs protected from cytosolic glycosidase activities that would prematurely release volatile aglycones. The relationship between precursor pool size and wine aromatic intensity reflects the efficiency of subsequent release mechanisms, with larger pools generally correlating with greater aromatic potential though release efficiency varies with precursor structure, yeast strain capabilities, and winemaking conditions employed during vinification. Environmental factors including temperature, water status, and light exposure influence precursor pool development through effects on biosynthetic enzyme activities and precursor stability, with moderate stress conditions often enhancing accumulation compared to unstressed controls through activation of secondary metabolism pathways responsive to environmental signals. Viticultural practices targeting precursor pool development include canopy management to optimize light exposure of clusters, irrigation management to impose controlled water stress, and harvest timing to capture optimal precursor concentrations before potential late-season degradation, with these interventions collectively capable of modifying precursor pools by 50-200% relative to standard practices.
Precursor_Pool_Development is subclass of Precursor_Formation representing the cumulative establishment of precursor reservoirs. Distinguishes from Flavor_Precursor_Synthesis which describes individual reactions. Sibling to Flavor_Precursor_Synthesis and Amino_Acid_Accumulation. Integrates outputs of multiple biosynthetic pathways. Key parameter is total precursor pool size determining aromatic potential. Glycosidic precursors typically 2-15 mg/L in aromatic varieties.
Präkursor-Pool-Expansion
Vorläufer-Pool-Entwicklung
aromatic potential development
aromatisches Potential-Entwicklung
bound flavor reservoir development
gebundenes Aroma-Reservoir-Entwicklung
precursor pool expansion
Gunata Z et al (1985) The aroma of grapes. Journal of Chromatography 331: 83-90 Williams PJ et al (1989) Glycosidic precursors of varietal grape and wine flavor. In Flavor Chemistry Trends and Developments ACS Symposium Series 388: 35-48 Keller M (2020) The Science of Grapevines 3rd Ed pp 295-305
Precursor Pool Development
Predawn water potential mapping across a 25-hectare Shiraz vineyard in McLaren Vale reveals spatial variation from -0.25 MPa in deep alluvial soils to -0.72 MPa in shallow rocky sections at the same date, with the 0.47 MPa difference corresponding to 35% variation in midday stomatal conductance and guiding precision irrigation to apply 40% more water to high-stress zones, achieving uniform predawn values of -0.35±0.08 MPa and eliminating the ripening heterogeneity previously caused by differential water stress.
Predawn water potential quality is a water potential quality that represents the tissue water potential measured before sunrise when transpiration has ceased overnight and plant water status has equilibrated with soil water potential in the root zone, providing an integrated assessment of soil water availability experienced by the vine root system. Within the BFO framework, this quality specifically characterizes the equilibrium water status achieved during the nocturnal recovery period, distinguishing this baseline measurement from the dynamically changing water potential during daytime transpiration. The ontological significance lies in predawn water potential serving as a proxy for soil water potential weighted by root distribution, eliminating the confounding effects of atmospheric demand and stomatal regulation that influence daytime measurements, thus providing the clearest indication of soil water supply independent of plant hydraulic properties. Predawn water potential in well-watered grapevines approaches -0.1 to -0.2 MPa, declining progressively as soil water depletes, with values of -0.3 to -0.5 MPa indicating mild stress, -0.5 to -0.8 MPa moderate stress, and below -1.0 MPa severe stress in most cultivars. The assumption of overnight equilibration underlying predawn measurement validity requires sufficient time without transpiration (typically measured within 1 hour before sunrise) and absence of significant nighttime transpiration that can occur under high vapor pressure deficit conditions in some environments. Measurement employs the pressure chamber technique on excised leaves or shoots, with rapid sampling critical to avoid artifact from post-excision water loss. The relationship between predawn water potential and soil water content follows the soil water retention curve, enabling conversion between plant-based stress indicators and soil moisture for irrigation scheduling. Understanding predawn water potential quality enables assessment of soil water availability, establishment of irrigation thresholds based on supply rather than demand, and interpretation of vine water status in the context of root zone conditions.
Predawn_Water_Potential_Quality provides the baseline water status measurement representing soil-plant equilibrium, distinguishing from Midday_Water_Potential_Quality reflecting dynamic transpirational stress. This quality assumes complete overnight equilibration, which may not occur under conditions of nighttime transpiration, hydraulic limitations, or heterogeneous soil moisture. Measurement timing within 1 hour before sunrise is critical for valid equilibrium assessment.
Basiswasserpotential
Predawn-Wasserpotential
Wasserpotential vor Sonnenaufgang
baseline water potential
nächtliches Gleichgewichtspotential
overnight equilibrium potential
predawn leaf water potential
Ψpd
Améglio T et al. (1999) Water relations, long-distance transport and accumulation of mineral content in a forest tree at low temperature. Plant, Cell & Environment 22(7): 871-883 Williams LE, Trout TJ (2005) Relationships among vine- and soil-based measures of water status in a Thompson Seedless vineyard in response to high-frequency drip irrigation. American Journal of Enology and Viticulture 56(4): 357-366
Predawn Water Potential Quality
Characterization of Preformed_Antimicrobial_Disposition at the Julius Kühn-Institut quantified basal antimicrobial protein activity and correlated it with disease resistance across 20 cultivars. Healthy leaf protein extracts were analyzed for chitinase activity (colorimetric assay with colloidal chitin), β-1,3-glucanase activity (reducing sugar release from laminarin), and total antifungal activity (radial diffusion assay against Botrytis cinerea). Basal chitinase activity ranged from 12 nkat/mg protein ('Müller-Thurgau') to 48 nkat/mg ('Regent'), while glucanase activity ranged from 8 to 35 nkat/mg across the same cultivars. Direct antifungal activity in diffusion assays showed 2.5-fold variation, with PIWI cultivars ('Regent', 'Solaris') exhibiting the highest inhibition zones. Cultivars with strong Preformed_Antimicrobial_Disposition showed significantly faster containment of pathogen spread following inoculation with P. viticola and B. cinerea, with 'Regent' (high basal PR activity) limiting lesion expansion to 3.2 mm at 72 hours post-inoculation compared to 8.5 mm in low-activity 'Müller-Thurgau'. Proteomic analysis revealed that 'Regent' maintained 3-fold higher constitutive levels of PR-2, PR-3, and PR-5 proteins compared to susceptible cultivars, potentially reflecting its interspecific hybrid background combining defense gene complements from multiple Vitis species. These findings demonstrate that Preformed_Antimicrobial_Disposition contributes to rapid pathogen containment independent of induced response kinetics.
A Preformed_Antimicrobial_Disposition is a Chemical_Defense_Disposition that inheres in a Vitis vinifera plant and is realized through constitutive accumulation of proteins and small molecules with direct antimicrobial activity against bacteria, fungi, and oomycetes, providing immediate defense upon pathogen contact without requiring induced synthesis. This disposition represents the protein and specialized metabolite components of chemical defense that directly attack or inhibit microbial pathogens through mechanisms distinct from the general antioxidant and protein-precipitating activities of bulk phenolics. Preformed antimicrobials in grapevine include pathogenesis-related proteins at basal expression levels (PR-2 β-1,3-glucanases, PR-3 chitinases, PR-4 chitin-binding proteins, PR-5 thaumatin-like proteins, PR-14 lipid transfer proteins), defensins and other antimicrobial peptides, and specialized metabolites with direct antifungal activity (certain stilbenes, pterostilbene at basal levels). These compounds function by degrading pathogen cell wall components (glucanases, chitinases), disrupting pathogen membranes (defensins, lipid transfer proteins), inhibiting pathogen enzymes, or directly killing pathogen cells. The constitutive levels of antimicrobial proteins vary among grapevine genotypes, with some cultivars and wild species maintaining higher basal expression providing enhanced first-line defense before induced responses are fully activated. Preformed_Antimicrobial_Disposition complements physical barriers by providing chemical attack on pathogens that successfully breach surface defenses, and serves as a foundation that can be rapidly augmented through induced defense pathways upon pathogen recognition.
Preformed_Antimicrobial_Disposition is_a Chemical_Defense_Disposition. This class addresses direct antimicrobial compounds and proteins at constitutive (pre-attack) levels, complementing induced antimicrobial responses (phytoalexin production, PR protein induction) under Induced_Defense_Disposition. Basal PR protein levels vary among genotypes and may explain differences in early infection containment before induced defenses reach full activity. PIWI cultivars often show elevated basal antimicrobial activity reflecting their interspecific hybrid backgrounds.
Basale-antimikrobielle-Aktivität
Konstitutive-PR-Proteine
Präformierte-antimikrobielle-Disposition
basal antimicrobial activity
constitutive PR proteins
preformed antifungal compounds
van Loon LC et al. (2006) Significance of inducible defense-related proteins in infected plants, Annual Review of Phytopathology 44:135-162 Monteiro S et al. (2003) Structural and functional aspects of PR-10 proteins, FEBS Letters 541:1-7 Ferreira RB et al. (2007) The wine proteins, Trends in Food Science and Technology 18:293-306
Preformed Antimicrobial Disposition
A Chardonnay vine in the Les Cailleret Premier Cru vineyard in Puligny-Montrachet bearing a Premier Cru role is subject to maximum yields of 54 hl/ha (versus 57 hl/ha for Village and 48 hl/ha for Grand Cru), minimum natural alcohol of 11.5% (versus 11% for Village), and labeling that combines the commune name with the vineyard name or the mention "Premier Cru," with 640 designated Premier Cru climats across the Côte d'Or producing wines typically priced at 40-120 €/bottle.
An appellation-specific role that inheres in a grapevine situated within a vineyard parcel classified at the second-highest tier of a hierarchical appellation system, positioned below Grand Cru but above village-level designation, subject to intermediate yield restrictions and quality requirements that reflect the parcel's recognized superiority over standard communal vineyards.
Premier_Cru_Role is most formally defined in Burgundy where 640 climat parcels across the Côte d'Or, Chablis, and Côte Chalonnaise hold Premier Cru status, established through centuries of empirical vineyard evaluation formalized by the 1861 Lavalle classification and subsequent INAO delimitations. Analogous second-tier classifications exist in Germany (VDP.Erste Lage), Austria (ÖTW Erste Lagen Klassifizierte Lagen), and the Bordeaux hierarchy: however, the specific regulatory implications and quality differentials vary considerably across these systems. In Burgundy, Premier Cru wines may be labeled with the individual climat name or blended under the commune Premier Cru designation.
Erste-Lage-Rolle
Premier-Cru-Rolle
Premiumlagen-Rolle
first growth role
premier cru vine role
second-tier cru role
INAO (2023) Cahiers des charges Premiers Crus de Bourgogne Lavalle J (1855) Histoire et statistique de la vigne et des grands vins de la Côte d'Or Pitiot S and Servant JC (2010) The Wines of Burgundy, Collection Pierre Poupon pp: 68-92
Premier Cru Role
Pinot Noir bears a Premium_Red_Wine_Scion_Role in Burgundy's Côte d'Or, where yields are restricted to 35-45 hL/ha (approximately 5-6 t/ha) by appellation regulation and often further by producers to achieve the concentration defining Grand Cru wines from Romanée-Conti, Chambertin, and Musigny. Nebbiolo serves in a Premium_Red_Wine_Scion_Role in Barolo and Barbaresco, producing age-worthy wines with tar, rose, and truffle complexity at yields below 8 t/ha. Cabernet Sauvignon bears a Premium_Red_Wine_Scion_Role in Napa Valley's Oakville and Rutherford AVAs, yielding 4-6 t/ha for wines commanding $100-500/bottle and defining California's luxury wine category.
A red wine scion role that inheres in a grafted scion cultivar by virtue of its designation for premium or ultra-premium red wine production, where viticultural and winemaking practices target quality optimization over yield maximization, and resulting wines command prices substantially above commodity market levels. Premium red wine scions are cultivated under conditions designed to achieve physiological ripeness with balanced sugar-acid ratios, concentrated phenolic extraction, and complex aromatic development—typically through yield restriction (often below 6-8 t/ha for top tiers versus 12-20 t/ha for bulk production), careful canopy management, optimal harvest timing, and site selection emphasizing terroir expression. Premium red wine scions are typically noble or classic varieties with established fine-wine reputations (Cabernet Sauvignon, Pinot Noir, Nebbiolo, Sangiovese, Syrah, Tempranillo, Merlot, Malbec), though emerging varieties and regional specialties may also serve in premium roles in quality-focused production systems. The premium tier includes wines ranging from regional appellations (€15-50/bottle) through classified growths and cult wines (€100-1000+/bottle).
Premium_Red_Wine_Scion_Role captures the quality-focused end of red wine production where cultivar selection, site matching, and intensive viticulture converge to produce wines distinguished by concentration, complexity, and aging potential. The boundary between premium and bulk production is not absolute the same cultivar (even the same vineyard block) may yield premium wine in favorable vintages and bulk wine in challenging years. Premium production typically involves greater labor investment (hand harvesting, manual cluster selection), lower yields (often 30-60% of bulk production levels), and extended barrel aging. Economic premium correlates imperfectly with intrinsic quality but generally reflects scarcity, reputation, and production costs.
Premium-Rotweinsorte
Qualitätsrotweinrebe
edle Rotweinsorte
fine red wine variety
noble red variety
premium red wine variety
quality red wine cultivar
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Amerine MA, Winkler AJ (1944) Composition and quality of musts and wines of California grapes, Hilgardia 15: 493-675 Matthews MA (2015) Terroir and Other Myths of Winegrowing, University of California Press Goode J (2014) The Science of Wine 2nd Ed, University of California Press
Premium Red Wine Scion Role
Chardonnay bears a Premium_White_Wine_Scion_Role in Burgundy's Côte de Beaune, where yields are restricted to 40-50 hL/ha (approximately 6-8 t/ha) in Meursault and Puligny-Montrachet to achieve the concentration, complexity, and aging potential (15-30+ years) defining Grand Cru white Burgundy at €200-1000+/bottle. Riesling serves in a Premium_White_Wine_Scion_Role in Germany's VDP estates, yielding 35-50 hL/ha for Grosses Gewächs from classified vineyards (Bernkasteler Doctor, Scharzhofberger, Forster Ungeheuer) commanding €50-200/bottle. Chenin Blanc bears a Premium_White_Wine_Scion_Role in Vouvray and Savennières, producing age-worthy dry and off-dry wines from yields below 45 hL/ha on tuffeau limestone and schist soils.
A white wine scion role that inheres in a grafted scion cultivar by virtue of its designation for premium or ultra-premium white wine production, where viticultural and winemaking practices target quality optimization, terroir expression, and aging potential over yield maximization. Premium white wine scions are cultivated under conditions designed to achieve physiological ripeness with preserved natural acidity, concentrated aromatic development, and balanced phenolic ripeness enabling extended lees contact or barrel aging—typically through yield restriction (often below 8-10 t/ha), optimal harvest timing balancing sugar, acid, and aroma precursor accumulation, and site selection emphasizing cool climates or high-altitude mesoclimates that extend the ripening period. Premium white wine scions are typically classic varieties with established fine-wine reputations (Chardonnay, Riesling, Chenin Blanc, Sauvignon Blanc, Grüner Veltliner, Albariño) cultivated in recognized quality regions. The premium tier includes wines ranging from regional appellations (€15-50/bottle) through classified growths and iconic bottlings (€100-500+/bottle).
Premium_White_Wine_Scion_Role captures quality-focused white wine production paralleling Premium_Red_Wine_Scion_Role. Premium white production often involves extended lees contact (sur lie aging), controlled oxidative handling (barrel fermentation and aging), and malolactic fermentation management to build complexity while preserving varietal character. Unlike premium reds where phenolic concentration is paramount, premium whites balance aromatic intensity, textural richness, and acid structure. The same cultivar frequently serves both premium and bulk roles depending on site and viticulture—Chardonnay ranges from €200 Montrachet to €5 bulk wine from the same varietal designation.
Premium-Weißweinsorte
Qualitätsweißweinrebe
edle Weißweinsorte
fine white wine variety
premium white wine variety
quality white wine cultivar
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Goode J (2014) The Science of Wine 2nd Ed, University of California Press Stevenson T (2011) The Sotheby's Wine Encyclopedia 5th Ed, Dorling Kindersley Matthews MA (2015) Terroir and Other Myths of Winegrowing, University of California Press
Premium White Wine Scion Role
Sangiovese vines at Antinori's Tignanello estate in Chianti Classico bearing a premium wine role for the Pèppoli label are managed at yields of 7-9 t/ha with selective cluster thinning, canopy management including shoot positioning and lateral removal, and harvest timing based on phenolic maturity assessments, producing wines retailing at 18-25 €/bottle that represent the estate's accessible quality tier below the icon-level Tignanello (55-80 €) and Solaia (150-250 €) wines.
A quality tier role that inheres in a grapevine whose fruit is designated for wines of recognized quality distinction positioned above commercial-grade production, typically sourced from identified vineyard blocks with moderate yield restrictions, receiving above-average viticultural attention, and marketed at price points reflecting acknowledged quality above everyday consumption levels.
Premium_Wine_Role occupies the broad middle ground of the quality hierarchy between commercial bulk production and the super-premium/icon tiers. Industry definitions vary, but the tier conventionally encompasses wines retailing at approximately $10-20 USD in the US market (or equivalent in other currencies). This is the most commercially significant quality tier by revenue in most established wine markets: it represents the segment where viticultural quality differentiation through yield management, canopy manipulation, and harvest timing begins to materially influence wine character, distinguishing premium wines from the volume-driven approach of commercial and bulk tiers.
Premium-Wein-Rolle
Qualitätswein-Rolle
fine wine production role
gehobene Wein-Rolle
premium wine vine role
quality wine role
Rabobank (2019) Wine Quarterly Report: Global Wine Market Segmentation McMillan R (2022) State of the US Wine Industry, Silicon Valley Bank Oczkowski E (2001) Hedonic wine price functions and measurement error, Economic Record 77: 374-382
Premium Wine Role
Pressure-volume analysis comparing drought-acclimated and well-watered Cabernet Sauvignon reveals acclimation-induced shift in osmotic potential at full turgor from -1.35 to -1.72 MPa and turgor loss point from -1.65 to -2.08 MPa, with the 0.43 MPa osmotic adjustment enabling maintenance of positive turgor and stomatal opening to 0.4 MPa lower water potential, while bulk elastic modulus increased from 12.5 to 18.3 MPa indicating cell wall stiffening that further enhances turgor maintenance under water deficit.
Pressure volume relationship quality is a physical quality that represents the functional relationship between water potential and relative water content in grapevine tissues, characterizing the integrated hydraulic and mechanical properties determining tissue water storage, turgor maintenance, and volume change behavior under varying water status. Within the BFO framework, this quality captures the constitutive relationship between two key state variables rather than either variable alone, providing parameters that describe tissue hydraulic behavior including osmotic potential at full turgor, turgor loss point, bulk elastic modulus, and apoplastic water fraction. The ontological significance lies in pressure-volume relationships providing the mechanistic framework for understanding how tissues respond to water deficit, where the curve shape determines whether tissues maintain turgor through rigid walls resisting volume change (high elastic modulus) or through osmotic adjustment lowering turgor loss point, representing fundamentally different drought adaptation strategies. The pressure-volume curve is constructed by progressively dehydrating excised tissue while measuring water potential and mass, with the relationship between inverse water potential and relative water deficit yielding linear and curvilinear portions corresponding to turgor-positive and turgor-zero regions respectively. Key parameters extracted include osmotic potential at full turgor (Ψπ100), osmotic potential at turgor loss point (Ψπtlp), turgor loss point (Ψtlp), bulk elastic modulus (ε), and relative apoplastic water content, each providing information about distinct aspects of tissue hydraulic behavior. Grapevine cultivars show substantial variation in pressure-volume parameters, with drought-tolerant varieties typically exhibiting more negative osmotic potentials (indicating solute accumulation) and either higher elastic modulus (maintaining turgor through rigid walls) or lower turgor loss point (tolerating lower water potentials before wilting). Understanding pressure-volume relationships enables mechanistic interpretation of drought tolerance, prediction of turgor maintenance capacity, and analysis of acclimation responses to water deficit through changes in curve parameters.
Pressure_Volume_Relationship_Quality addresses the constitutive relationship between water potential and water content, integrating Turgor_Pressure_Quality, Osmotic_Potential_Quality, and Turgor_Loss_Point_Quality into a unified framework. This quality requires destructive sampling and bench-top measurement protocol, providing mechanistic parameters rather than field-measurable water status indicators. The class supports drought tolerance characterization and analysis of acclimation responses at the tissue hydraulic level.
Druck-Volumen-Beziehung
Gewebehydraulik-Parameter
P-V curve
P-V-Kurve
Wasserabgabekurve
hydraulic capacitance relationship
tissue water relation parameters
water release curve
Tyree MT, Hammel HT (1972) The measurement of the turgor pressure and the water relations of plants by the pressure-bomb technique. Journal of Experimental Botany 23(1): 267-282 Bartlett MK et al. (2012) The determinants of leaf turgor loss point and prediction of drought tolerance of species and biomes: a global meta-analysis. Ecology Letters 15(5): 393-405
Pressure Volume Relationship Quality
Prevailing wind direction for Margaret River, Western Australia is predominantly from the west-southwest (WSW to SW), bringing maritime air that moderates summer temperatures and maintains the cool-climate character essential for the region's Cabernet Sauvignon and Chardonnay styles. The Tramontana in Empordà (Catalonia) blows from the north-northwest (NNW) for up to 200 days per year, and vineyard rows are traditionally oriented north-south to allow wind penetration for canopy drying while minimizing mechanical shear on cordons. In the Wachau, prevailing wind direction shifts seasonally from westerly (W) in summer to easterly-southeasterly (E-SE) in autumn and winter, with the autumn Föhn events bringing warm, dry descending air that accelerates late-season ripening of Grüner Veltliner and Riesling on the Danube terraces.
Prevailing wind direction is a wind data entity that records the dominant compass bearing from which atmospheric air movement most frequently originates at a viticultural site, determined through wind rose analysis of directional frequency distributions over climatological standard periods, expressed in compass points (N, NE, E, etc.) or degrees from true north (0-360°), and serving as a critical factor in vineyard layout design, row orientation decisions, windbreak placement, and disease management strategy.
Prevailing wind direction exerts viticultural influence through multiple pathways that vary by regional context. In maritime regions, onshore winds from the prevailing direction transport moisture and moderate temperature extremes, while in continental interiors, prevailing wind direction determines exposure to cold air advection during frost-vulnerable phenological stages. Wind rose analysis should encompass both directional frequency and directional wind speed distributions, as the most frequent wind direction may differ from the direction carrying the highest energy. Topographic channeling through valleys and gaps can create highly localized prevailing directions that diverge substantially from regional synoptic patterns — a single vineyard's exposure within a valley may experience dramatically different wind directionality than a site only hundreds of meters away on an opposing slope. Modern vineyard planning increasingly employs computational fluid dynamics (CFD) modeling to predict wind patterns at the mesoscale before committing to row orientation and training system selection.
Hauptwindrichtung
Windrichtungsdaten
dominant wind direction
dominante Windrichtung
prevailing wind bearing
principal wind bearing
vorherrschende Windrichtung
wind direction record
wind rose direction
Gladstones J (2011) Wine, Terroir and Climate Change, Wakefield Press, pp.: 148-155 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 74-80 Dry PR, Smart RE (2015) Vineyard site selection. In Viticulture Volume 1: Resources, 3rd Ed, Winetitles, pp.: 89-108
Prevailing Wind Direction
Primary amino nitrogen for Sauvignon Blanc from Marlborough (2024 vintage) measures 195 mg N/L by NOPA assay, representing 68% of total YAN (285 mg N/L), with HPLC profiling revealing glutamine (45 mg N/L), arginine (38 mg N/L), alanine (22 mg N/L), and threonine (18 mg N/L) as the dominant amino acids — the high arginine content being characteristic of cool-climate Sauvignon Blanc and favorable for fermentation vigor. Primary amino nitrogen for Nebbiolo from Barolo (2023 vintage) records 85 mg N/L (52% of 162 mg N/L YAN), with the low PAN reflecting the nitrogen-poor calcareous-clay soils of the Langhe and the cultivar's inherently moderate nitrogen accumulation, necessitating targeted amino acid-based supplementation to support the 15-18 day maceration fermentation typical of Barolo production.
Primary amino nitrogen is a nitrogen data entity that records the concentration of free α-amino acids (excluding proline and hydroxyproline) in grape must, determined by the NOPA spectrophotometric assay (o-phthaldialdehyde/N-acetyl-L-cysteine reaction), ninhydrin colorimetric method, or HPLC amino acid profiling, expressed in milligrams of nitrogen per liter (mg N/L), representing the organic nitrogen fraction assimilable by Saccharomyces cerevisiae during fermentation and serving as both a fermentation nutrient metric and a determinant of yeast-derived aroma compound diversity through the Ehrlich pathway of amino acid catabolism.
Primary amino nitrogen provides a more nuanced fermentation nutrient assessment than YAN alone, as the amino acid composition within PAN directly influences the spectrum of fermentation-derived flavor compounds through the Ehrlich pathway — each branched-chain or aromatic amino acid is transaminated and decarboxylated to a corresponding higher alcohol and subsequently esterified, creating the fermentative bouquet that distinguishes different must compositions. The individual amino acid profile carries winemaking significance beyond aggregate PAN values: leucine generates isoamyl alcohol and isoamyl acetate (banana aroma), phenylalanine produces 2-phenylethanol and 2-phenylethyl acetate (rose aroma), and valine yields isobutanol — while cysteine and glutathione serve as conjugated precursors for volatile thiol release in Sauvignon Blanc and other aromatic varieties. Proline, typically the most abundant amino acid in grape must (300-1000 mg/L), is deliberately excluded from PAN measurement because Saccharomyces cerevisiae lacks the mitochondrial proline oxidase activity required for its catabolism under the anaerobic conditions of fermentation. The NOPA assay has become the industry standard for PAN determination due to its simplicity, speed (single spectrophotometric reading at 335 nm after 10-minute reaction), specificity for primary amines, and commercial kit availability, though it does not discriminate between individual amino acids — HPLC amino acid profiling provides detailed speciation when specific amino acid information is needed for research or premium winemaking applications. Vineyard nitrogen management practices directly modulate PAN composition — late-season foliar urea application (5-10 kg N/ha at véraison) preferentially increases arginine content, while soil nitrogen availability influences the overall amino acid pool size.
Aminosäurestickstoff
FAN
PAN
PAN-Wert
alpha-amino nitrogen
amino acid nitrogen
free amino nitrogen
freier Aminostickstoff
primärer Aminostickstoff
α-Aminostickstoff
Dukes BC, Butzke CE (1998) Rapid determination of primary amino acids in grape juice using an o-phthaldialdehyde/N-acetyl-L-cysteine spectrophotometric assay. American Journal of Enology and Viticulture 49(2): 125-134 Bell SJ, Henschke PA (2005) Implications of nitrogen nutrition for grapes, fermentation and wine. Australian Journal of Grape and Wine Research 11(3): 242-295 Hazelwood LA et al. (2008) The Ehrlich pathway for fusel alcohol production: a century of research on Saccharomyces cerevisiae metabolism. Applied and Environmental Microbiology 74(8): 2259-2266
Primary Amino Nitrogen
Primary amino nitrogen quality in Oregon Pinot Gris provides 80-180 mg N/L of the variety's typical 120-220 mg N/L YAN, with the amino acid profile dominated by arginine (30-80 mg N/L) and glutamine (15-40 mg N/L) that support fermentation while also providing precursors for potential ethyl carbamate formation from arginine-derived urea winemakers managing ethyl carbamate risk in warmer vintages employ yeast strains with reduced urea excretion while maintaining the PAN levels required for complete fermentation of moderate-sugar Pinot Gris.
Primary_Amino_Nitrogen_Quality represents the chemical quality characterizing the concentration of alpha-amino nitrogen from free amino acids (excluding proline) in grape must, capturing the organic nitrogen fraction assimilable by wine yeast that provides both nitrogen nutrition and specific amino acid substrates for yeast metabolism. Primary amino nitrogen (PAN), also termed free alpha-amino nitrogen (FAN), comprises the nitrogen associated with the alpha-amino groups of free amino acids that yeast can transport and metabolize during fermentation. In grape must, PAN typically ranges from 50-300 mg N/L, comprising 60-90% of total YAN in most samples and providing the majority of yeast-assimilable nitrogen. The amino acid composition underlying PAN affects fermentation outcomes beyond the total nitrogen contribution, as individual amino acids serve as precursors for specific volatile compounds through the Ehrlich pathway (higher alcohols and their esters), provide sulfur nutrition (methionine, cysteine), and modulate nitrogen metabolism regulation. The exclusion of proline from PAN calculations reflects its non-assimilability under anaerobic fermentation conditions, though proline can comprise 30-60% of total free amino acids in grape must, making the distinction between total amino acids and PAN critical for fermentation prediction. PAN measurement employs the formol titration method (Sörensen), o-phthaldialdehyde (OPA) colorimetric assay, or calculation from HPLC amino acid profiles minus proline. The OPA method has become standard for rapid PAN assessment, providing the primary input for YAN calculation alongside ammonium determination.
Primary_Amino_Nitrogen_Quality addresses the organic fraction of YAN, comprising free amino acids minus proline. The terminology (primary amino nitrogen, free alpha-amino nitrogen, FAN) varies among laboratories and publications this class encompasses all equivalent designations. The exclusion of proline from PAN is based on Saccharomyces cerevisiae's inability to catabolize proline under anaerobic fermentation conditions, though this limitation does not apply to all yeast species or aerobic conditions. The PAN fraction provides both general nitrogen nutrition and specific amino acid precursors affecting volatile compound formation.
Alpha-Aminostickstoffkonzentration
FAN quality
FAN-Qualität
Primärer Aminostickstoff
alpha-amino nitrogen concentration
free amino nitrogen content
freier Aminostickstoffgehalt
organic assimilable nitrogen
Dukes BC and Butzke CE (1998) Rapid Determination of Primary Amino Acids in Grape Juice Using an o-Phthaldialdehyde/N-Acetyl-L-Cysteine Spectrophotometric Assay. American Journal of Enology and Viticulture 49(2): 125-134 Henschke PA and Jiranek V (1993) Yeasts—Metabolism of Nitrogen Compounds. In: Fleet GH (ed) Wine Microbiology and Biotechnology, Harwood Academic Publishers pp.: 77-164
Primary Amino Nitrogen Quality
Primary aroma description for Gewürztraminer from Alsace (2024 vintage) identifies intense lychee and rose petal (from the monoterpenes geraniol and cis-rose oxide, both well above sensory thresholds), Turkish delight and mango (characteristic of the cultivar's unique terpene-norisoprenoid interaction), and white pepper spice (from trace rotundone) — a primary aromatic signature so distinctive and intense that Gewürztraminer is identifiable by nose alone with greater reliability than virtually any other cultivar. Primary aroma description for Sauvignon Blanc from Sancerre (2024 vintage) documents cut grass and green capsicum (from IBMP at 12 ng/L, near the white wine sensory threshold), passionfruit and grapefruit (from volatile thiols 3MH at 1200 ng/L and 3MHA at 180 ng/L, released from grape-derived precursors during fermentation), and a subtle flinty mineral quality — the characteristic Loire Sauvignon Blanc balance between herbaceous methoxypyrazine and tropical thiol aromatics. Primary aroma description for Grüner Veltliner from the Wachau (2024 vintage) records white pepper (from rotundone at 8-14 ng/L, near the 16 ng/L threshold but contributing subliminally to the spicy impression), green apple, and citrus zest — the peppery-citrus primary aromatic profile that defines Austrian Grüner Veltliner typicity.
Primary aroma description is an aroma description entity that documents the olfactory characteristics of wine attributable to grape-derived volatile and semi-volatile compounds originating from the berry itself, encompassing varietal-specific aromatic signatures (terpenes in Muscat and Riesling, methoxypyrazines in Sauvignon Blanc and Cabernet Franc, rotundone in Syrah and Grüner Veltliner, volatile thiols released from grape precursors), fruit character descriptors (citrus, stone fruit, tropical fruit, red fruit, dark fruit), and floral and herbal notes directly linked to cultivar identity and viticultural conditions, representing the aromatic dimension that most directly reflects the vineyard origin, grape variety, vintage conditions, and viticultural management encoded in the fruit at harvest.
Primary aroma description captures the aromatic dimension most directly linked to the viticultural information entities documented throughout the VVO hierarchy — cultivar identity, clone selection, terroir characteristics, vintage conditions, and vineyard management all manifest in the primary aromatic profile of the wine. The chemical basis of primary aromas encompasses several distinct compound classes, each with characteristic sensory contributions and cultivar-specific accumulation patterns: monoterpenes (floral, citrus — dominant in Muscat, Riesling, Gewürztraminer), C₁₃-norisoprenoids (floral, exotic fruit, honey — significant in Riesling, Chardonnay, Syrah), methoxypyrazines (green, herbaceous — characteristic of Cabernet Sauvignon, Sauvignon Blanc, Carménère), volatile thiols (tropical, citrus, mineral — defining Sauvignon Blanc, contributing to Colombard, Grüner Veltliner), rotundone (black/white pepper — distinctive in Syrah, Grüner Veltliner, Vespolina, Schioppettino), and dimethyl sulfide (truffle, cooked cabbage at low levels, blackcurrant contribution in young Syrah). The relationship between primary aroma expression and viticultural conditions creates the aromatic dimension of terroir — the same cultivar produces different primary aromatic profiles in different sites due to temperature effects on terpene synthesis, UV exposure effects on methoxypyrazine photodegradation, nitrogen availability effects on thiol precursor formation, and water status effects on concentration and biosynthesis. The persistence of primary aromas in wine varies dramatically by compound class — monoterpenes and methoxypyrazines are relatively stable, declining slowly over years, while volatile thiols are highly oxidation-sensitive and may diminish substantially within 2-3 years of bottling, making the primary aroma profile a temporally evolving characteristic rather than a fixed attribute.
Primäraroma
Rebsortenaroma
Sortenaroma
cultivar aroma signature
grape-derived aroma
primary fruit character
traubengebundenes Aroma
varietal aroma
varietal character description
varietales Aroma
Robinson AL et al. (2014) Origins of grape and wine aroma. Part 1. Chemical components and viticultural impacts. American Journal of Enology and Viticulture 65(1): 1-24 Lund ST, Bohlmann J (2006) The molecular basis for wine grape quality — a volatile subject. Science 311(5762): 804-805 Siebert TE et al. (2008) Development of a stable isotope dilution analysis for the quantitation of rotundone in grape juice and wine. Journal of Agricultural and Food Chemistry 56(10): 3745-3748
Primary Aroma Description
Primary aroma expression comparison between single-vineyard and blended Riesling from same producer reveals primary intensity of 8.2 versus 5.8 (10-point scale), with single-vineyard showing vivid lime, white flower, and petrol precursor notes from low-yield old vines while blended wine's primary character is diluted by higher-yield fruit contributing less concentrated varietal compound reservoir, demonstrating how vineyard source quality directly affects primary aroma expression and justifying premium pricing for single-vineyard bottlings that deliver more intense varietal character.
Primary aroma expression quality is an aroma evolution quality that represents the intensity and clarity of grape-derived varietal aromatic character, characterizing the contribution of compounds originating directly from grape metabolism to the wine's aromatic profile and indicating the preservation of varietal identity through winemaking. Within the BFO framework, this quality specifically addresses the grape-derived component of wine aroma, distinguishing primary (grape) aromas from secondary (fermentation) and tertiary (aging) aromas while recognizing that primary character represents the varietal foundation upon which secondary and tertiary elements build during wine development. The ontological significance lies in primary aroma expression providing the aromatic evidence of grape quality and varietal identity, where strong primary expression indicates successful capture of varietal character from high-quality fruit while weak primary expression may indicate neutral variety, underripe harvest, or winemaking losses that failed to preserve grape-derived volatiles. The compositional basis for primary aromas includes terpenes (linalool, geraniol in aromatic varieties), methoxypyrazines (providing green pepper in Sauvignon family), thiols (tropical character when released from grape-derived precursors), norisoprenoids (violet, stone fruit notes from carotenoid-derived compounds), and C6 compounds (fresh green notes from lipid degradation in grape tissue), with the specific compound profile defining varietal aromatic fingerprint. The winemaking influence on primary aroma preservation is substantial, with reductive handling, cold fermentation, and minimal processing maximizing retention of volatile primary compounds while oxidative handling, warm fermentation, and extended processing may diminish primary character in favor of secondary and tertiary elements. The balance between primary expression and complexity involves trade-offs, as wines emphasizing primary fruit purity may sacrifice the complexity that secondary and tertiary aromas provide, creating stylistic choices between fruit-forward immediacy and layered complexity. Understanding primary aroma expression quality enables assessment of grape quality contribution, varietal authenticity verification, and winemaking evaluation regarding primary character preservation.
Primary_Aroma_Expression_Quality specifically addresses the grape-derived aromatic component, representing the varietal foundation of wine character. This quality is maximized in young, fresh wines and typically diminishes with aging as primary compounds transform or volatilize while tertiary elements develop. The class supports grape quality assessment and varietal authenticity evaluation while recognizing that primary dominance is not universally desirable across all wine styles and ages.
Primäraromen
Primärfruchtausdruck
Sortenaromatik
cépage aromas
grape-derived character
primary fruit expression
traubengebundene Aromen
Francis IL, Newton JL (2005) Determining wine aroma from compositional data. Australian Journal of Grape and Wine Research 11(2): 114-126 Rapp A (1998) Volatile flavour of wine: correlation between instrumental analysis and sensory perception. Nahrung 42(6): 351-363
Primary Aroma Expression Quality
Primary aroma quality in Torrontés from Salta, Argentina defines this cultivar's aromatic intensity rivaling Muscat Blanc, with free monoterpene concentrations of 2-4 mg/L (versus 0.1-0.3 mg/L in neutral varieties) dominated by linalool and geraniol that produce the intensely floral, rose-petal character distinguishing Argentine Torrontés the high-altitude (1700-2400m) vineyards with intense UV exposure promote terpene glycoside accumulation providing a reservoir of bound aroma compounds that release progressively during fermentation and aging, extending aromatic complexity beyond the immediate floral impact of free terpenes.
Primary_Aroma_Quality_(varietal) encompasses the chemical qualities characterizing grape-derived aromatic compounds that define varietal character and typicity, capturing the volatile and precursor compounds present in berries at harvest that distinguish one grape variety from another. Primary aromas originate from grape metabolism and accumulate in berry tissues independent of fermentation, representing the olfactory signature that makes Muscat smell floral, Sauvignon Blanc herbaceous, and Riesling petrol-like with age. This category includes Terpene_Content_Quality capturing the monoterpenes responsible for floral Muscat character Methoxypyrazine_Content_Quality addressing the green bell pepper compounds of Sauvignon Blanc and Cabernet Sauvignon C13_Norisoprenoid_Quality encompassing carotenoid degradation products contributing fruity and floral notes and Thiol_Precursor_Quality characterizing cysteine and glutathione conjugates that release tropical fruit thiols during fermentation. Primary aroma compounds occur in both free volatile form, immediately contributing to fresh grape aroma, and bound glycosidic form, requiring enzymatic or acid hydrolysis to release the volatile aglycone. The ratio of free to bound forms varies among compound classes and cultivars, affecting both fresh grape aroma intensity and aging potential. Primary aroma expression depends strongly on cultivar genetics, with certain varieties (Muscat, Gewürztraminer, Riesling) classified as aromatic due to exceptionally high terpene accumulation, while other varieties (Chardonnay, Pinot Noir) produce minimal primary aroma compounds and rely on fermentation and aging for aromatic interest. Environmental factors including temperature, light exposure, and water status modulate primary aroma accumulation within genetically determined limits, providing viticultural tools for aroma optimization.
Primary_Aroma_Quality_(varietal) captures the grape-derived aroma dimension distinct from fermentation-derived (secondary) and aging-derived (tertiary) aromas. The parenthetical (varietal) follows the class naming convention in the uploaded structure, emphasizing the cultivar-specific nature of primary aroma profiles. The bound-to-free ratio of aroma compounds is quality-relevant because bound forms represent aroma potential released during winemaking and aging. The classification of cultivars as aromatic versus neutral based on terpene content represents a fundamental varietal grouping affecting wine style expectations and production approaches.
Primäraromaqualität
Sortenbukett
cultivar-specific aroma
grape-derived volatile quality
primary volatile composition
traubenstämmige Aromastoffe
varietal aroma content
Mateo JJ and Jiménez M (2000) Monoterpenes in Grape Juice and Wines. Journal of Chromatography A 881(1-2): 557-567 Dunlevy JD et al. (2009) A Methyltransferase Essential for the Methoxypyrazine-Derived Flavour of Wine. The Plant Journal 59(3): 404-412
Primary Aroma Quality (varietal)
Syrah in Rhône Valley shows primary branch primordium formation during late July-August Year N at BBCH 27-35 within developing latent buds over 3-4 weeks, with organized inflorescence meristems initiating 24-34 primary branch primordia in basipetal sequence along elongating axis reaching 500-700 micrometers length under conditions of 24-28°C temperature, high light exposure in well-managed canopy, moderate water stress, and balanced nutrition typical of this moderately-to-highly fruitful mid-to-late season cultivar establishing compound inflorescence architecture
Primary branch primordium formation represents the architectural elaboration stage where organized inflorescence meristem initiates first-order lateral branch primordia establishing primary branching structure that will determine major architectural framework and contribute substantially to ultimate flower number capacity, creating compound inflorescence characteristic of grapevine. This essential developmental stage occurs within developing latent bud during mid-late summer Year N following meristem differentiation, typically spanning 2-4 weeks as primary branch primordia initiate sequentially along elongating inflorescence axis producing 15-40 primary branches depending on cultivar and developmental conditions. Branch initiation proceeds basipetally from inflorescence apex with branch primordia emerging from peripheral zone of inflorescence meristem in regular phyllotactic pattern, typically spiral arrangement with divergence angles of 135-140 degrees creating characteristic branching geometry. Primary branch primordium emergence shows local cell division acceleration in peripheral meristem zone creating meristematic protrusion 20-40 micrometers diameter, with each primordium containing 20-50 actively dividing cells that will elaborate into branch structure bearing secondary branches and flowers. Branch number determination depends on inflorescence meristem activity duration and axis elongation rate, with longer activity period and greater elongation producing more branch initiation sites increasing total primary branch number, while shortened development reduces branch number affecting cluster architecture. Temporal sequence shows branch initiation beginning at inflorescence apex when primordium reaches 150-200 micrometers length, proceeding basipetally as axis elongates with successive branches initiating at 1-2 day intervals during peak development, continuing for 2-4 weeks until axis elongation ceases and basal branch initiation completes. Developmental gradient within elaborating inflorescence shows most advanced development at apex where early-formed branches begin developing their own secondary branching, intermediate development in middle regions with recently formed primary branches, and least development at base where final primary branches are just initiating. Branch primordium development following initiation involves continued cell division increasing primordium size to 50-100 micrometers over 5-10 days, beginning establishment of branch meristem organization, and preparation for subsequent secondary branch formation or flower initiation depending on position and architecture. Inflorescence axis elongation accompanies primary branch formation extending from initial 200-300 micrometers to final 400-800 micrometers by completion of primary branching, with elongation creating spatial framework distributing branch attachment points along axis length. Cell division rates during active primary branch formation show highest activity in branch initiation zones with cycles of 20-30 hours, moderate activity in elongating axis with 30-40 hour cycles, and maintained activity in apical meristem with 24-36 hour cycles. Gene expression patterns include sustained reproductive identity genes maintaining inflorescence character, auxin response genes showing local maxima at branch initiation sites, and meristem maintenance genes in developing branch primordia establishing branch identity. Hormonal regulation involves auxin transport establishing concentration patterns that mark and drive branch initiation sites with polar auxin transport creating spatial organization, cytokinins supporting cell division in initiating branches and maintaining meristem activity, and continued low gibberellins permitting reproductive development. Environmental factors affecting primary branch formation include light intensity with adequate light supporting maximum branch initiation frequency producing highly compound inflorescences while shade reduces branching creating simpler architecture, temperature within optimal 24-28°C range supporting rapid sequential branch initiation, and adequate resources enabling complete branching pattern development. Canopy microclimate strongly affects branching with well-exposed shoots showing robust primary branch formation producing 25-40 branches creating large compound inflorescences, while shaded interior shoots show reduced branching with 10-20 branches producing smaller simpler clusters. Metabolic demands during primary branch formation intensify as multiple developing structures require resources simultaneously, with respiration increasing to support elevated cell division rates throughout elaborating inflorescence primordium. Carbon availability affects branching with adequate photosynthetic production supporting full branching potential while limitation reduces branch number affecting ultimate cluster size and flower number. Nitrogen availability influences branching with moderate nitrogen supporting good branch formation while deficiency may reduce branching and excess may alter development through effects on overall plant vigor. Water status may affect branching with moderate stress potentially acceptable though severe stress during this critical elaboration period can reduce branching affecting yield potential. Architectural diversity among cultivars reflects differences in primary branch formation with highly compound varieties producing 30-40 primary branches creating large dense clusters, moderately compound types producing 20-30 branches with intermediate structure, and simply branched varieties showing 10-20 branches creating loose small clusters. The process represents third distinguishable stage of inflorescence primordium formation following meristem differentiation and preceding secondary branch formation, establishing primary architectural framework determining cluster structure. Duration spans approximately 2-4 weeks depending on developmental rate and cultivar branching characteristics, with warm conditions promoting rapid completion while cooler temperatures extend period. Completion timing shows primary branch formation finishing typically late July-August in temperate regions Year N as shoot growth declines and developmental activity in buds decreases preparing for dormancy entry. Regional timing shows primary branch primordium formation during late July-August in warm climates, August-early September in temperate regions, and August in cool areas corresponding to mid-late summer period when adequate resources remain available for inflorescence elaboration. Viticultural significance includes establishing primary architectural framework determining cluster size and structure affecting yield capacity and quality attributes, understanding that conditions during branch formation critically affect subsequent productivity, recognizing that management supporting adequate but not excessive vigor promotes optimal branching, and appreciating that stress during branching can permanently reduce cluster size affecting yield. Research applications include analyzing mechanisms regulating branch initiation frequency and spatial patterning, investigating hormonal and genetic control of inflorescence branching architecture, quantifying environmental effects on branching affecting yield components, evaluating relationships between primary branching and ultimate flower number, and assessing genetic variation in branching patterns affecting cluster architecture and productivity across cultivars.
Primary_Branch_Primordium_Formation is third of four Inflorescence_Primordium_Formation subphases, distinguished by first-order branch initiation establishing major architectural framework. Follows Inflorescence_Meristem_Differentiation. May overlap with Secondary_Branch_Primordium_Formation as early-formed primary branches begin producing secondary branches. Primary determinant of inflorescence complexity and flower number capacity. Critical phase requiring adequate resources and favorable conditions.
Erstordnungsverzweigung
Hauptzweigbildung
Primärzweig-Primordienbildung
first-order branch formation
primary branch initiation
primary branching
primäre Blütenstandsverzweigung
Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109 Srinivasan C & Mullins MG (1981) Physiology of flowering in the grapevine, Am J Enol Vitic 32:47-63
Primary Branch Primordium Formation
A primary branch rachis representing the fourth primary branch on a primary inflorescence of Vitis vinifera cv. Grenache in Châteauneuf-du-Pape, France, arising from the main rachis at 6.5 cm from the inflorescence base, measuring 5.8 cm length representing substantial primary branch development, diameter of 2.4 mm at junction with main rachis tapering to 1.2 mm at the branch apex, exhibiting cylindrical morphology with slight flattening, green coloration with moderate pubescence, containing 5 vascular bundles in cross-section providing vascular capacity for the branch system, bearing 8 secondary branch rachides (third-order structures) distributed along its length at 0.5-0.8 cm intervals, secondary branches bearing flowers and tertiary branches creating the complete branch system, primary branch supporting approximately 45 flowers representing 6.6% of the total inflorescence flower number (680 total), serving to extend the flower-bearing zone laterally from the main rachis creating three-dimensional inflorescence volume, positioned as one of 14 primary branches distributed along the 19.2 cm main rachis, measured at BBCH stage 63 (30% flowers open) in early June, representing typical primary branch rachis architecture with second-order position and intermediate dimensions.
A lateral branch axis arising directly from the main rachis representing second-order structures in the hierarchical inflorescence architecture in Vitis vinifera, characterized by origin from the main central rachis through lateral branching, second-order hierarchical position in the rachis system, typical dimensions of 2-8 cm length and 1.5-3.5 mm diameter representing substantial reduction from main rachis, moderate vascular capacity with 3-7 vascular bundles providing water and nutrient delivery to the branch system, bearing secondary branch rachides (third-order) and flowers creating the next level of architectural elaboration, typical occurrence of 10-20 primary branches per inflorescence distributed along the main rachis length, and functional roles including distributing flowers throughout the three-dimensional inflorescence volume, extending vascular networks from main rachis to peripheral regions, and providing distributed mechanical support creating the branched paniculate architecture. Primary branch rachides represent the first lateral branching order, developing through lateral meristem initiation along the developing main rachis, exhibiting dimensions and complexity intermediate between massive main rachis and fine higher-order branches, and creating the primary lateral elaboration defining paniculate inflorescence architecture.
Primary branch rachides are distinguished from the main rachis by lateral origin and smaller dimensions, and from secondary branch rachides by second-order versus third-order hierarchical position. The primary branch designation emphasizes the first-order lateral status representing the initial branching from the main axis. Second-order position means these structures originate directly from the first-order main rachis, contrasting with third-order secondary branches originating from primary branches. Number per inflorescence shows cultivar variation from relatively few (8-12) in weakly branched inflorescences to many (18-25+) in elaborately branched structures, correlating with overall inflorescence complexity. Distribution along main rachis shows relatively regular spacing creating systematic architecture, with typical intervals of 0.8-2.0 cm between adjacent primary branches. Size variation along main rachis shows basipetal gradient with largest primary branches near inflorescence base progressively decreasing toward apex, reflecting developmental timing and resource allocation patterns. Diameter ratio to main rachis typically 0.5-0.8× indicating substantial size reduction reflecting hierarchical subordination. Vascular capacity shows proportional reduction with bundle number often 40-60% of main rachis bundle number. Branching complexity varies with some primary branches showing simple structures bearing only flowers while others elaborate extensively with multiple orders of secondary and tertiary branches. Developmental origin involves lateral meristem initiation along the developing main rachis with sequential appearance creating the distributed array. Transformation to cluster maintains primary branch identity with lignification and berry development creating the major lateral cluster arms.
erste laterale Achse
first lateral branch
primary lateral axis
primäre Zweigrachis
second-order rachis
zweiter Ordnung Spindel
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 259-295 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 291-326 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 479-518.
Primary Branch Rachis
A primary bud at node 6 on a winter-pruned cane of Vitis vinifera cv. Chardonnay, dissected in late winter revealing two fully formed inflorescence primordia of 195 and 178 flower initials respectively, plus vegetative shoot primordia with 8-10 leaf initials visible under magnification, representing potential yield of approximately 373 berries assuming typical 50% fruit set for this cultivar, with this primary bud being the first to break dormancy in spring and the principal contributor to shoot and crop production from this node position.
The largest and most developmentally advanced bud within the compound bud structure of Vitis vinifera, occupying the central position within the protective bud scales, characterized by fully differentiated shoot and inflorescence primordia formed during the previous growing season, highest priority for growth activation following dormancy release, and greatest reproductive potential among the three hierarchical bud positions (primary, secondary, tertiary) in the compound structure. Primary buds represent the principal source of current season crop production, typically containing 0-3 inflorescence primordia depending on cultivar, node position, and environmental conditions during differentiation, with inflorescence number and flower count per cluster largely predetermined 8-15 months before budbreak, making primary bud fruitfulness assessment critical for yield prediction and crop load management decisions.
Primary buds are distinguished from secondary and tertiary buds by their larger size (typically 2.5-4mm diameter versus 1-2mm for secondary), central position within the compound structure, earlier differentiation during previous season development, and consistently higher fruitfulness. Damage to primary buds through spring frost, hail, herbicide drift, or disease triggers compensatory growth from secondary buds, though typically with reduced yield and delayed phenology. Understanding primary bud fruitfulness patterns by node position and cultivar is fundamental to pruning decisions, with balanced pruning formulas based on correlations between vine size and optimal primary bud retention numbers.
Hauptknospe
Primärknospe
central bud
main bud
principal bud
zentrale Knospe
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Srinivasan & Mullins (1981) "Physiology of flowering in the grapevine - a review" American Journal of Enology and Viticulture 32=47-63 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Morrison (1991) "Bud development in Vitis= Anatomical study of formation and growth of the primary grapevine bud during one annual cycle" Australian Journal of Grape and Wine Research 72=296-301.
Primary Bud
Precision assessment of Primary_Bud_Hardiness_Disposition at Iowa State University employed high-resolution differential thermal analysis to distinguish low-temperature exotherms (LTEs) from primary, secondary, and tertiary bud components across 15 cultivars at biweekly intervals from November through March. Primary bud LTEs occurred at consistently warmer temperatures than secondary and tertiary LTEs, confirming the hierarchy of bud component hardiness. At maximum hardiness in mid-January, 'Frontenac' exhibited exceptional Primary_Bud_Hardiness_Disposition with LT50 of -34.2°C, while V. vinifera 'Pinot Noir' showed marginal hardiness at -19.8°C. The difference between primary and secondary bud LT50 (hardiness differential) varied among cultivars from 2.1°C ('Brianna') to 5.8°C ('Chambourcin'), with implications for crop recovery potential following intermediate freeze events. Field trials validated the predictive value of Primary_Bud_Hardiness_Disposition, with significant correlation between laboratory LT50 and field survival at individual node positions following natural freeze events (r = 0.88). Economic analysis demonstrated that each 1°C improvement in Primary_Bud_Hardiness_Disposition reduced average annual winter damage costs by approximately $120/ha in central Iowa conditions, providing quantitative justification for the strong breeding emphasis on this trait in cold-climate cultivar development programs.
A Primary_Bud_Hardiness_Disposition is a Bud_Cold_Hardiness_Disposition that inheres specifically in the primary (central) bud primordium within the compound grapevine bud and is realized through the supercooling capacity and freeze avoidance mechanisms of this most developmentally advanced and fruitful bud component. This disposition represents the cold tolerance of the bud structure that produces the primary shoot bearing the majority of the crop under normal circumstances, making Primary_Bud_Hardiness_Disposition the most economically significant component of bud cold hardiness. The primary bud is the first-formed and largest primordium within the compound bud, containing well-developed cluster primordia (typically 1-3 depending on cultivar) and representing the dominant growth point that will produce the main shoot the following spring. Due to its larger size and more advanced development, the primary bud typically exhibits lower cold hardiness than the smaller, less differentiated secondary and tertiary buds, with LT50 values generally 2-6°C warmer than secondary buds within the same compound bud structure. This differential hardiness has been attributed to differences in tissue hydration, supercooling capacity related to primordium size, and developmental stage effects on cryoprotective compound accumulation. Primary_Bud_Hardiness_Disposition directly determines the temperature threshold for significant crop loss, as primary bud kill eliminates the most fruitful shoots and requires reliance on less productive secondary growth for crop recovery.
Primary_Bud_Hardiness_Disposition is_a Bud_Cold_Hardiness_Disposition. The primary bud produces the main, most fruitful shoot and is typically least hardy among the three bud components within the compound bud. Primary bud survival is the primary determinant of full crop potential, with primary bud kill requiring reliance on less fruitful secondary shoots for crop recovery. The differential between primary and secondary bud hardiness varies among cultivars and influences crop recovery potential. Primary bud LT50 is the most commonly reported bud hardiness parameter and the primary selection criterion in cold-hardy cultivar breeding.
Hauptknospen-Frosthärte
Primärknospen-Härte-Disposition
Zentralknospen-Kältetoleranz
central bud hardiness
main bud LT50
primary bud freeze tolerance
Pool RM et al. (1990) Environmental factors controlling cold hardiness and dormancy in grapevines, VITIS Journal of Grapevine Research 29:129-144 Wolf TK, Cook MK (1994) Cold hardiness of dormant buds of grape cultivars: comparison of thermal analysis and field survival, HortScience 29:1453-1455 Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235
Primary Bud Hardiness Disposition
Primary bud viability assessment across temperature damage gradient in Merlot reveals the relationship between cold exposure and primary bud survival, with vineyard low spots (experiencing -22°C minimum) showing only 35% primary viability (65% of primary buds killed, severe yield impact), mid-slope positions (-19°C minimum) showing 78% primary viability (moderate injury), and upper slope with air drainage (-16°C minimum) showing 96% primary viability (minimal injury), with the viability gradient enabling site-specific management responses—heavy additional bud retention in low areas, moderate adjustment mid-slope, and normal pruning on upper slopes—to optimize recovery across the variable damage pattern.
Primary bud viability quality is a bud viability quality that represents the living status of the primary (central) bud within the compound bud structure, characterizing the survival of the most important bud for grape production that contains the highest fruitfulness potential and produces the shoots with greatest yield capacity when viable and growing. Within the BFO framework, this quality specifically addresses the primary bud dimension of viability, distinguishing the survival of this critical bud from secondary or tertiary bud survival while recognizing that primary bud viability has disproportionate importance for productivity because primary buds typically carry 1.5-2 clusters per shoot while secondary buds may carry only 0.5-1 cluster, making each primary bud substantially more valuable for yield than backup buds. The ontological significance lies in primary bud viability directly determining the fruitful shoot population and thus the yield potential for the coming season, where high primary viability enables normal crop expectations while low primary viability forces reliance on less fruitful secondary buds (reducing yield potential) or on compensatory measures (leaving additional buds to increase total shoot number despite lower per-shoot fruitfulness). The differential cold sensitivity of primary buds makes them the most vulnerable component of the compound bud, with primary buds typically showing LT50 values 2-4°C warmer (less hardy) than secondary buds and 4-6°C warmer than tertiary buds, such that cold events may kill primary buds while leaving backup buds viable, creating the partial survival scenarios common after moderate winter injury. The assessment timing for primary bud viability typically occurs in late winter or early spring after the coldest temperatures have passed but before budbreak, enabling management decisions (adjusted bud numbers, training modifications) to accommodate the revealed survival pattern. Understanding primary bud viability quality enables accurate yield forecasting, appropriate pruning adjustments for damaged vineyards, and recognition of the yield implications when primary mortality forces dependence on secondary bud shoots.
Primary_Bud_Viability_Quality addresses the survival of the most fruitful bud component within compound buds. This quality has disproportionate yield importance due to higher fruitfulness of primary versus secondary buds. The class supports yield forecasting and pruning adjustment through primary bud viability characterization.
Hauptknospenüberleben
Primärknospenvitalität
Zentralknospenvitalität
central bud viability
main bud survival
primary meristem viability
primäre Meristemlebensfähigkeit
principal bud vitality
Howell GS (2000) Cold hardiness of grapevine buds and canes: research update. Proceedings of the ASEV 50th Anniversary Meeting, Seattle, Washington, pp. 187-192 Pool RM et al. (1990) Identifying cold injury in grapevines. Cornell Extension Bulletin, Geneva, NY
Primary Bud Viability Quality
Cabernet Franc in Loire Valley shows primary cluster separation during mid-May Year N+1 at late BBCH 55 over 4-5 days, with 24-32 primary branches elongating to 10-15 mm length and spreading to 50-65 degree angles creating 2.5-4.0 mm spacing along 25-35 mm rachis under temperate maritime conditions of 15-19°C mean temperature, adequate resources, typical of this moderately loose-clustered vigorous mid-season cultivar establishing well-separated primary cluster architecture
Primary cluster separation represents the first-order spatial organization phase where primary branches differentiate from main rachis through elongation and angular positioning, establishing major divisions within inflorescence creating recognizable first-order flower groups corresponding to primary branch units. This fundamental organizational stage occurs during mid-May in temperate regions Year N+1 as part of cluster separation process, typically requiring 3-5 days as primary branches achieve sufficient extension and angular divergence to create perceptible separation from rachis and neighboring branches. Separation mechanism involves primary branch elongation extending branches 8-18 mm from rachis attachment points creating spatial projection, branch angle increase from compressed acute positions toward 45-70 degree angles establishing lateral spreading, and rachis elongation between branch attachment points creating 2-5 mm spacing along primary axis. Primary branch differentiation shows each branch unit containing cluster of flower buds at distal positions, with branch serving as structural support and transport pathway connecting flowers to main cluster vascular system through rachis. Branch number along rachis determines number of primary clusters with typical range 15-40 primary branches per inflorescence creating 15-40 first-order flower groups, reflecting primordium formation during previous season with cultivar variation in branching frequency. Spatial organization created by primary separation establishes coarse-scale cluster architecture with widely-spaced first-order units in loose-clustered varieties showing 4-6 mm between branches, moderately-spaced intermediate types showing 2-4 mm spacing, and tightly-spaced compact varieties showing 1-2 mm spacing affecting overall cluster density. Branch elongation mechanism involves cell division in intercalary meristems at branch base adding cells to branch length, and cell expansion throughout branch tissues increasing cell dimensions through turgor-driven growth, with elongation rate typically 0.5-1.5 mm per day at optimal temperatures. Branch angle development proceeds through differential growth on abaxial versus adaxial sides creating angular change, with abaxial side showing enhanced expansion driving branch outward from rachis, establishing spreading three-dimensional architecture. Vascular development in separating primary branches shows continued xylem and phloem differentiation establishing functional transport capacity, with vascular trace connecting branch to main rachis vascular system enabling water delivery and assimilate distribution to flowers on branch. Mechanical properties during separation show flexible structure from thin-walled tissues allowing positioning adjustments, with progressive wall thickening increasing rigidity though final strength development awaits fruit set and berry growth. Flower bud development on primary branches continues during separation with buds enlarging as flower organs mature, preparing for subsequent individual flower separation within each primary cluster unit. Metabolic activity in separating primary branches maintains high levels supporting elongation and angle development, with respiration elevated providing energy for growth processes. Hormonal regulation involves auxin affecting branch elongation and positioning through effects on cell expansion and growth orientation, with auxin gradients potentially coordinating spatial organization among branches. Environmental effects on primary separation include temperature as primary factor with optimal 16-22°C supporting rapid uniform separation, cool conditions slowing elongation reducing separation extent, and warm conditions accelerating development though potentially causing stress if excessive. Resource availability affects separation with adequate photosynthetic production supporting vigorous branch elongation creating well-separated clusters, while limitation reduces elongation maintaining compact architecture. Developmental gradient within inflorescence shows apical primary branches often separating earlier reflecting advanced development in earlier-formed structures, while basal branches show progressive separation as development proceeds. Within-branch uniformity shows similar developmental stage among flowers on each primary branch preparing for coordinated anthesis within branch unit, supporting synchronous flowering and fruit set. The process represents first of four flower cluster separation aspects establishing coarse-scale spatial organization before finer-scale secondary separation and individual flower separation, providing primary architectural framework. Duration typically spans 3-5 days from initiation to substantial completion depending on temperature and genetic elongation characteristics, with rapid progression under favorable conditions. Cultivar variation appears in separation extent with loose-clustered varieties showing substantial primary branch elongation and spreading creating 4-6 mm spacing while tight-clustered types show minimal elongation maintaining 1-2 mm spacing, affecting cluster compactness and microenvironment. Regional timing shows primary cluster separation during mid-May in warm climates, late May in temperate regions, and late May-early June in cool areas, occurring as part of late inflorescence emergence approaching flowering. Viticultural significance includes establishing primary cluster structure affecting disease pressure with well-separated clusters showing better air circulation and reduced infection risk, influencing later fruit exposure and quality characteristics, and revealing architectural pattern established during previous season primordium formation. Research applications include quantifying genetic and environmental variation in primary branch separation affecting cluster architecture, analyzing mechanisms regulating branch elongation and angular positioning, evaluating relationships between primary separation and disease susceptibility, and assessing implications of separation patterns for fruit quality outcomes.
Primary_Cluster_Separation is first of four Flower_Cluster_Separation subphases, distinguished by first-order spatial organization through primary branch differentiation. Establishes coarse-scale cluster architecture. Transitions to or overlaps with Secondary_Cluster_Separation in compound architectures. Primary determinant of cluster density and microenvironment. Extent reflects genetic architecture program and developmental conditions.
Erstordnungs-Blütentrennung
Hauptzweigseparation
Primärcluster-Separation
erste Gescheinsgliederung
first-order cluster division
main branch differentiation
primary branch separation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 340-345 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Primary Cluster Separation
A primary first basal leaf at node 1 on a primary shoot of Vitis vinifera cv. Merlot in Pomerol, France, emerging from a primary bud following spur pruning leaving two-node spurs, developing from the well-formed primordium within the primary bud component of the compound bud structure, measuring 17.8 cm length by 16.5 cm width representing robust development characteristic of primary bud origin, exhibiting typical Merlot leaf morphology with five lobes, deep U-shaped petiolar sinus, moderate lateral sinuses, and characteristic marginal teeth diagnostic for this cultivar, emerging 4 days post-budbreak ahead of secondary and tertiary bud activation, achieving full expansion at 23 days post-budbreak, demonstrating high photosynthesis rate of 13.8 μmol CO₂ m⁻² s⁻¹ reflecting complete primordial development during previous season including full chloroplast and enzyme complement formation, providing critical early-season photosynthate supporting primary shoot extension achieving 8-12 cm growth per week during May, positioned opposite the primary shoot's first inflorescence bearing the main crop cluster, measured at shoot BBCH stage 71 (fruit set) in mid-June, representing the standard first basal leaf type dominant in commercial vineyard canopies where primary bud shoots account for 90-98% of productive shoots.
A first basal leaf at node 1 on shoots emerging from primary buds within compound grapevine buds in Vitis vinifera, characterized by development from the most advanced primordium within the compound bud structure representing the primary bud's contribution, position at node 1 on shoots developing from primary buds that typically dominate growth under normal conditions following winter pruning and budbreak, larger dimensions and more vigorous development compared to potential secondary or tertiary bud leaves reflecting the primary bud's developmental priority and superior resource allocation during previous-season primordium formation, morphological characteristics showing typical cultivar-diagnostic features including characteristic lobe number, sinus depth, and marginal tooth patterns as these leaves express normal adult morphology without developmental abnormalities, early emergence timing typically 3-6 days post-budbreak from primary buds, high photosynthetic capacity reflecting well-developed preformed primordia with complete tissue differentiation prior to budbreak, and critical functional role providing immediate high-quality photosynthetic capacity supporting vigorous primary shoot development that dominates canopy architecture and fruit production in commercial vineyards. Primary first basal leaves represent the standard first leaf type in commercial viticulture where winter pruning and cultivar selection maintain primary bud dominance, developing from compound bud primary components that receive preferential resource allocation during the previous growing season when primordia are formed, resulting in well-developed preformed leaves ready for rapid deployment following budbreak.
Primary first basal leaves are distinguished from secondary or tertiary bud first leaves by origin from primary bud components within compound buds, superior development reflecting primary bud resource priority, and typical cultivar morphology without developmental abnormalities. The compound bud structure in grapevine contains primary, secondary, and tertiary bud components with decreasing developmental advancement, with primary buds showing most complete primordial development and highest priority for resource allocation during formation. Primary bud dominance in commercial viticulture results from selection of well-developed buds during pruning and favorable overwintering conditions that maintain primary bud viability. Frost damage, winter injury, or severe stress can reduce primary bud viability causing secondary or tertiary bud development, with resulting first leaves showing characteristics of these less-developed bud components. Primary first basal leaves showing abnormal morphology, reduced size, or delayed development may indicate problems with previous-season primordium formation suggesting stress, disease, or nutritional issues during the prior growing season when buds were forming.
Hauptknospe Knoten-eins Blatt
dominant first basal leaf
dominantes basales Weinblatt
main bud node one leaf
primary bud first leaf
primäres erstes basales Blatt
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 86-108 Carmona et al. (2008) "Grapevine bud dormancy= environmental and molecular aspects" Revista Brasileira de Fruticultura 30=257-408 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 119-154.
Primary First Basal Leaf
Chardonnay on Teleki 5C showing primary first-order lateral shallow roots, excavation revealing 3 major laterals 5.8-7.4 cm diameter originating from primary root at 18-32 cm depth, primary laterals extending 180-240 cm horizontal distance representing longest lateral roots, microscopic sections showing 12-14 annual rings and extensive vascular development with 45-60 large xylem vessels, primary laterals producing 120-180 second-order branches each creating extensive hierarchical networks, primary first-order laterals demonstrating dominant structural and functional importance
Primary first order lateral shallow root is first-order lateral shallow root emphasizing primary importance and dominant status among lateral roots, characterized by largest diameter 4-8 cm, greatest horizontal extent 100-250+ cm, most extensive secondary growth and woody development, maximum transport capacity, and most abundant second-order branching creating most elaborate hierarchical network, with primary first-order laterals representing major structural and functional elements in root system serving as main lateral axes from which majority of lateral root network derives, typically 2-4 primary first-order laterals per vine positioned at different depths 15-40 cm and directions creating radial coverage, showing characteristics including origin from primary taproot or main cutting-derived root, earliest formation timing among first-order laterals reflecting developmental priority, persistent lifespan potentially equaling vine age functioning decades, maximum vascular development with extensive xylem and phloem providing highest transport conductance, and critical importance where primary first-order lateral extent and productivity largely determines total lateral root system capacity. Primary first order lateral shallow root dominance characteristics includes= showing largest diameter (primary first-order laterals 4-8 cm at origin substantially larger than other first-order laterals 2-4 cm), exhibiting greatest length (primary laterals extending 100-250+ cm versus typical first-order 50-150 cm), demonstrating maximum branching (primary laterals producing most abundant second-order branches creating most elaborate networks), revealing highest transport capacity (primary laterals with most extensive vascular tissue providing greatest conductance), and displaying structural prominence (primary laterals forming most substantial components of lateral root framework). Primary first order lateral shallow root developmental priority includes= showing early initiation (primary laterals typically forming earliest among first-order lateral population), exhibiting vigorous growth (primary laterals showing faster elongation and secondary growth than other first-order), demonstrating resource allocation (vine apparently allocating disproportionate resources to primary lateral development), revealing competitive advantage (early-forming primary laterals potentially suppressing later-forming laterals through resource preemption), and displaying establishment success (primary laterals showing highest survival and persistence rates). Primary first order lateral shallow root vascular development includes= showing extensive xylem (primary laterals with numerous large-diameter xylem vessels providing high water transport capacity), exhibiting abundant phloem (substantial phloem tissue enabling high photosynthate transport to lateral network), demonstrating maximum conductance (primary lateral vascular tissue providing greatest hydraulic and carbon transport capacity), revealing functional importance (primary lateral transport capacity critically affecting vine water status and growth), and displaying progressive development (primary lateral vasculature increasing with age through continued secondary growth). Primary first order lateral shallow root branching productivity includes= showing maximum second-order production (primary laterals producing most numerous second-order branches potentially 50-200+ per root), exhibiting regular branching (second-order branches emerging at consistent intervals 10-25 cm along primary lateral length), demonstrating hierarchy creation (primary lateral branching creating most extensive hierarchical networks reaching fourth or fifth-order), revealing absorptive support (primary laterals ultimately supporting thousands to millions of fine absorptive roots through hierarchical network), and displaying critical function (primary lateral branching productivity determining large fraction of total root system absorption capacity). Primary first order lateral shallow root spatial dominance includes= showing preferential directions (primary laterals often extending in preferred directions accessing best soil or avoiding competition), exhibiting asymmetric distribution (primary laterals potentially more developed in favorable directions creating asymmetric root system), demonstrating competitive exclusion (vigorous primary laterals potentially suppressing development of other first-order laterals nearby through resource competition), revealing soil volume control (primary laterals and derivatives potentially controlling 60-80% of total soil volume explored by lateral root system), and displaying architectural impact (primary lateral distribution largely determining overall lateral root system architecture). Primary first order lateral shallow root lifespan and stability includes= showing maximum persistence (primary laterals potentially surviving entire vine lifespan functioning decades), exhibiting lowest mortality (primary lateral death rates extremely low given structural importance and carbon investment), demonstrating continued growth (primary laterals showing ongoing diameter growth and secondary development throughout life), revealing cumulative investment (primary laterals representing largest long-term carbon investments in root system), and displaying permanence (primary laterals forming essentially permanent structural framework). Primary first order lateral shallow root functional importance includes= determining lateral capacity (primary lateral extent and productivity largely determining total lateral root system absorption and transport capacity), enabling resource acquisition (primary laterals and derivatives providing 50-70% of total vine water and nutrient uptake), demonstrating drought tolerance (deep-reaching primary laterals potentially accessing moisture unavailable to shallower roots), revealing management significance (protecting primary laterals from damage critical for maintaining vine productivity), and displaying selection target (rootstock breeding potentially selecting for superior primary lateral development improving root system performance).
Primary_First_Order_Lateral_Shallow_Root emphasizes dominant primary importance and is distinguished from other first-order laterals by focus on largest size and greatest productivity. Critical for understanding structural dominance, maximum transport, highest branching, spatial control, and the major first-order laterals forming primary framework determining lateral root system capacity.
Dominant first-order root
Dominante Hauptseitenast
Haupt-Lateralachse
Main primary lateral
Primäre erste Ordnung Lateralwurzel
Principal lateral axis
Primary First Order Lateral Shallow Root
In Müller-Thurgau berries in Pfalz, Germany during BBCH 81-89, primary flavor formation establishes glucose accumulation from 45 to 115 g/L and fructose from 42 to 108 g/L over 45 days, organic acid reduction from 18.5 to 7.2 g/L titratable acidity, and C6 volatile synthesis with hexanal declining from 85 to 22 µg/L and trans-2-hexenal from 120 to 35 µg/L as berry maturity increases, establishing the sweet-tart-fresh flavor foundation under moderate continental conditions.
Primary flavor formation describes the biosynthesis of fundamental flavor-active metabolites in Vitis vinifera berries during ripening that derive directly from primary metabolic pathways including carbohydrate, amino acid, and lipid metabolism, generating the baseline flavor compounds including sugars contributing sweetness perception, organic acids determining sourness and pH-dependent flavor expression, and lipid-derived volatiles providing green and herbaceous character that form the foundational matrix upon which secondary metabolite-derived flavors are superimposed. This process encompasses the continued accumulation of glucose and fructose to levels of 180-260 g/L at harvest providing the sweet taste component fundamental to ripe grape flavor perception, the modulation of malic and tartaric acid ratios determining the sourness and freshness characteristics, and the synthesis of C6 aldehydes and alcohols through lipoxygenase-mediated oxidation of membrane lipids generating hexanal, hexenal, and hexenol isomers that contribute green, grassy, and herbaceous notes particularly prominent in under-ripe or heavily processed berries. The biochemical pathways underlying primary flavor formation involve glycolytic and gluconeogenic carbon flux determining sugar accumulation rates, tricarboxylic acid cycle activity influencing organic acid pools, and membrane lipid turnover providing substrates for lipoxygenase and hydroperoxide lyase activities that generate C6 volatile compounds through oxidative cleavage of polyunsaturated fatty acids. Temporal dynamics of primary flavor formation show compound-specific patterns with sugar accumulation proceeding continuously throughout ripening following sigmoidal kinetics, organic acid reduction occurring primarily during early-to-mid ripening as previously described, and C6 compound synthesis showing complex patterns reflecting the balance between lipoxygenase activity and volatile degradation or conjugation processes. Environmental factors influence primary flavor formation through effects on carbon assimilation and partitioning, with adequate photosynthesis supporting sugar accumulation, temperature regulating organic acid metabolism, and oxidative conditions potentially enhancing lipid-derived volatile formation through increased lipoxygenase pathway flux. The contribution of primary flavor compounds to overall grape and wine flavor perception establishes the fundamental taste profile including sweetness, sourness, and basic aromatic character upon which the more complex secondary metabolite-derived flavors create variety-specific and terroir-expressive nuances distinguishing quality wines from ordinary productions.
Primary_Flavor_Formation is subclass of Flavor_Compound_Synthesis representing biosynthesis from primary metabolic pathways. Distinguishes from Secondary_Metabolite_Production which generates specialized compounds. Sibling to Secondary_Metabolite_Production and Flavor_Pathway_Activation. Includes sugars, organic acids, and lipid-derived C6 compounds. Establishes baseline flavor matrix for secondary metabolite overlay.
Grundaroma-Biosynthese
baseline flavor development
basic flavor compound synthesis
fundamental flavor biosynthesis
fundamentale Aroma-Entwicklung
primary metabolite flavor formation
primäre Aromabildung
primäre Metabolit-Aromabildung
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 250-265 Kalua CM and Boss PK (2009) Evolution of volatile compounds during the development of Cabernet Sauvignon grapes. Journal of Agricultural and Food Chemistry 57: 3818-3830 Conde C et al (2007) Biochemical changes throughout grape berry development. Food Chemistry 1: 1-22
Primary Flavor Formation
A primary inflorescence at node 3 on a shoot of Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, California, measuring 18.2 cm length at full bloom representing the largest inflorescence on this shoot, positioned opposite the third fully expanded leaf counting from the shoot base, exhibiting complex compound paniculate architecture with central rachis bearing 4-5 orders of branching creating highly elaborate structure, rachis diameter of 3.8 mm at base representing robust construction, bearing approximately 650 individual flowers organized across the branching network, showing significantly larger dimensions compared to the secondary inflorescence at node 5 (12.5 cm length, ~380 flowers) on the same shoot, developmental stage at BBCH 63 (30% flowers open) during the mid-bloom period, primary inflorescence initiated in the compound bud during early June of the previous season receiving priority resource allocation during formation, expected to set approximately 60-75 berries following successful pollination and fruit set contributing an estimated 55% of the total shoot yield, measured in early June during peak flowering period, representing typical primary inflorescence characteristics with maximum size, complexity, and yield contribution.
The first and typically largest inflorescence on a grapevine shoot in Vitis vinifera, characterized by position at node 3 or occasionally node 4 representing the first occurrence in the inflorescence sequence along the shoot, largest dimensions among all inflorescences on the same shoot typically 12-25 cm length at full development, most complex branching architecture often showing highest branch order and flower number, developmental priority receiving preferential resource allocation during both primordium initiation and current season development, position opposite the third leaf (or fourth when present) following the two-leaf gap after the basal non-reproductive nodes, critical importance for crop yield typically contributing 40-60% of total shoot fruit production, and role as the primary reproductive structure with secondary inflorescences (when present) at nodes 5, 7, etc. showing progressively reduced size and complexity. Primary inflorescences represent the most developed reproductive structures on fruiting shoots, initiated earliest during inflorescence primordium formation in the previous season receiving maximum developmental investment, showing highest flower number and largest final cluster size after fruit set, and serving as the principal yield component with secondary inflorescences providing supplementary production.
Primary inflorescences are distinguished from secondary and tertiary inflorescences by position at node 3 (occasionally 4) and typically superior size and complexity. The primary designation emphasizes both positional primacy (first in sequence) and functional primacy (largest and most productive). Size superiority is typical but not universal, with some cultivars showing relatively balanced inflorescence sizes or occasional reversal where secondary exceeds primary due to developmental perturbations. Position at node 3 represents the standard pattern, though some cultivars or specific shoots may show primary at node 4 particularly on vigorous water shoots or when basal nodes show abnormal development. Developmental priority reflects earlier initiation during primordium formation and preferential resource allocation creating advantages in flower number and structural elaboration. Branching complexity typically shows maximum development with primary inflorescences exhibiting highest branch orders, though cultivar variation exists from relatively simple to highly complex architectures. Yield contribution dominance reflects the size and flower number advantages, with primary inflorescences typically accounting for 40-60% of shoot production though ratios vary with secondary and tertiary inflorescence development. Cluster thinning practices often retain only primary inflorescences removing secondaries to optimize fruit quality through reduced crop load and improved resource allocation. The transformation from primary inflorescence to primary cluster following fruit set maintains the size and yield dominance throughout berry development and maturation.
Hauptblütenstand
basal inflorescence
erster Blütenstand
first inflorescence
main cluster
primäre Infloreszenz
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 146-178 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 109-142 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 313-358.
Primary Inflorescence
Cabernet Sauvignon vines in Napa Valley's Oakville AVA at BBCH 55 typically exhibit 14-18 primary inflorescence branches averaging 3.5-5 cm length, with basal branches reaching 6-7 cm and bearing 4-6 secondary branches each, while apical primary branches measure only 2-3 cm with 1-2 secondary branches. In Syrah vineyards of the northern Rhône Valley (Côte-Rôtie), primary branches number 16-22 per inflorescence with lengths of 4-6.5 cm and extensive secondary subdivision (5-8 secondary branches per primary), creating large, moderately loose clusters of 150-300 berries at maturity.
A primary inflorescence branch is an inflorescence branch that arises directly from the main inflorescence rachis as a first-order lateral appendage, representing the principal architectural framework determining ultimate inflorescence size, flower number, and cluster compactness at maturity. Primary branches emerge from the main rachis at regular intervals (phyllotactic spacing typically 2-5 mm) during inflorescence primordium initiation within the dormant bud, with branch number genetically determined but environmentally modulated during the previous growing season. These branches exhibit the greatest length and diameter among all branching orders, typically measuring 1.5-8 cm in length and 0.8-2.5 mm in diameter at the base, with cultivar-dependent variation correlating strongly with final cluster weight and architecture. Primary branch anatomy features a prominent central vascular bundle connecting to the main rachis vascular system, allowing substantial resource transport to support extensive secondary and tertiary sub-branching. Branch positioning along the rachis shows characteristic basipetal gradients in length and complexity, with basal primary branches generally longer (often 1.5-2× apical branch length) and more extensively subdivided than apical primary branches, creating the conical or cylindrical cluster shapes characteristic of different cultivar groups. Each primary branch typically bears 8-45 flowers distributed on its axis and secondary branches, though flower number varies dramatically with cultivar (Pinot Noir= 15-25 flowers/primary branch Grenache= 25-45 flowers/primary branch Gewürztraminer= 8-15 flowers/primary branch). Primary branch development occurs during BBCH 53-57, with branch elongation and secondary branch emergence proceeding acropetally from the rachis base toward the apex. The number of primary branches (typically 10-25) serves as a critical ampelographic descriptor, with cultivars classified as having few (8-12), medium (13-18), or many (19-25+) primary branches under standard growing conditions. Primary branch vigor and flower-bearing capacity responds to previous-season cultural practices, particularly crop load, water status during June-July bud differentiation, and canopy light exposure, making branch characteristics indicators of vine developmental history.
Primary_Inflorescence_Branch represents the first-order architectural element and should be carefully distinguished from Secondary_Inflorescence_Branch (which arises from primary branches rather than directly from the main rachis). The term 'primary' refers to branching hierarchy, not to Primary_Inflorescence versus Secondary_Inflorescence (the latter being lateral inflorescences on a shoot). Primary branch count and average length at BBCH 55-57 strongly predict final cluster weight with cultivar-specific regression relationships. Sub-classification into Lower, Upper, and Basal positions reflects significant morphological and functional heterogeneity along the rachis axis. This class is critical for ampelographic cluster architecture description and for predictive modeling of final yield components.
Hauptast der Infloreszenz
Primärzweig
first-order inflorescence branch
main inflorescence branch
primärer Blütenstandsast
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 88-92 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 75-82 OIV (2009). Descriptor List for Grape Varieties and Vitis Species, 2nd ed., OIV code 204 (cluster density).
Primary Inflorescence Branch
A primary lateral vein on a mature leaf at node 6 of Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, California, representing one of five pairs of primary lateral veins, branching from the median vein at 5.5 cm from petiolar junction creating the second major lobe position, branching angle of 62 degrees from median vein axis, extending 7.2 cm through the lobe to within 0.4 cm of margin, diameter of 1.8 mm at median vein junction representing largest lateral vein size, tapering to 0.7 mm distally, highly prominent as raised ridge on abaxial surface 0.4 mm above surrounding tissue, visible as distinct groove on adaxial surface, containing 7-9 large xylem vessels (40-70 μm diameter) providing water transport capacity for the lobe region representing 19% of total blade area, substantial phloem cross-section enabling efficient photosynthate collection and export, thick-walled mechanical tissues providing structural support, serving as primary axis from which 12-15 secondary lateral veins branch at intervals of 0.4-0.6 cm creating hierarchical network, secondary veins subsequently branching into tertiary and higher orders completing reticulate pattern throughout the lobe, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing the primary first-order lateral vein architecture fundamental to palmate leaf organization.
A lateral vein representing the first-order branches from the median vein in the hierarchical venation system in Vitis vinifera, characterized by direct branching origin from the median vein without intermediate vein orders, largest diameter among lateral vein orders typically 1.2-2.0 mm at junction with median vein, highest hierarchical position among lateral veins serving as primary organizing axes for higher-order vein networks, maximum vascular capacity among lateral veins with most numerous and largest xylem vessels supporting water delivery to entire lobe regions, greatest mechanical strength providing primary structural support for blade lobes, typical number of 4-6 pairs per leaf creating the fundamental palmate architecture, extension through major blade lobes from median vein to lobe margins or near-margin regions, and critical functional importance as the principal vascular pathways serving distinct blade sectors with each primary lateral vein and its branching network supplying a major lobe region representing 15-25% of total blade area. Primary lateral veins establish the fundamental palmate venation pattern characteristic of grapevine leaves, differentiating during early leaf development as the first lateral vein order following median vein formation, organizing the subsequent development of secondary and higher-order veins that complete the reticulate network, and determining lobe number, positioning, and architecture through their branching pattern creating the cultivar-diagnostic leaf shapes that serve as primary ampelographic characters.
Primary lateral veins are distinguished from higher-order lateral veins (secondary, tertiary) by direct branching from the median vein and largest diameter, representing the first hierarchical level of lateral venation. The primary designation emphasizes their position at the apex of the lateral vein hierarchy from which all higher-order lateral veins derive through progressive branching. Number of primary lateral vein pairs determines fundamental lobe architecture, with cultivars ranging from 3 pairs (three-lobed leaves like some Gewürztraminer clones) to 6 pairs (seven-lobed leaves in deeply dissected cultivars), though 4-5 pairs creating five-lobed leaves predominates. Branching angles and trajectories determine lobe shapes, with variation from acute angles creating narrow elongated lobes to obtuse angles creating broad rounded lobes. During development, primary lateral veins differentiate following median vein establishment, with primordia appearing sequentially along the median vein and developing outward coordinating with lobe primordium development. Vascular capacity and mechanical strength reflect functional importance as primary supply pathways for major blade regions, with each primary lateral vein essentially functioning as the "median vein" for its lobe sector. Cultivar identification often involves assessment of primary lateral vein characteristics including number, branching angles, prominence, and trajectory patterns contributing to overall leaf shape.
Haupt-Zweigvene
erste Ordnung Lateralvene
main lateral vein
primäre Seitenader
principal branch vein
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 90-135 Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 95-136.
Primary Lateral Vein
A primary main rachis on a compound primary inflorescence of Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, California, representing the first-order axis in the hierarchical organization, originating directly from the peduncle at the inflorescence base, measuring 18.5 cm length establishing the primary structural dimension, diameter of 4.5 mm at base representing maximum rachis construction on this inflorescence, containing 13 major vascular bundles providing primary-order vascular capacity, exhibiting monopodial organization with continuous central axis, serving as the direct source for 16 primary branch rachides (second-order structures) which subsequently elaborate into secondary branches (third-order) and higher orders creating the complete hierarchical architecture, receiving developmental priority with primary main rachis establishment preceding all lateral branch development, providing the essential central framework without which the inflorescence cannot exist as lateral branches depend completely on the primary main rachis for support and vascular connection, bearing the complete structural load of the inflorescence including all lateral branches and flowers, measured at BBCH stage 65 (full bloom) in early June, representing the primary main rachis in its role as the foundational first-order structure organizing the complete hierarchical architecture.
The main rachis emphasizing its role as the primary first-order axis in the hierarchical inflorescence architecture in Vitis vinifera, characterized by first-order hierarchical position representing the apex of the structural organization, origin directly from the peduncle without intermediate structures, role as the source from which all higher-order structures derive through sequential branching, maximum structural priority receiving developmental precedence and resource allocation, dimensions and vascular capacity reflecting the primary position typically 3-5 mm diameter with 10-15 vascular bundles, monopodial organization establishing the primary axis framework, and functional primacy serving as the essential central structure without which the complete inflorescence architecture cannot exist. Primary main rachis emphasizes the hierarchical supremacy and organizational primacy of the main rachis, representing the first-order foundation structure, developing through initial apical meristem activity establishing the primary axis before any lateral elaboration, and serving as the indispensable organizing framework determining the fundamental inflorescence architecture.
Primary main rachis emphasizes the hierarchical and organizational aspects of the main rachis, distinguished from simple main rachis designation by explicit focus on first-order position and primacy. The primary designation highlights the position at the apex of the structural hierarchy from which all higher orders derive through sequential branching. First-order status means the primary main rachis has no precursor rachis structures, originating directly from the peduncle contrasting with second-order primary branches which originate from the primary main rachis and third-order secondary branches originating from primaries. Developmental sequence shows primary main rachis establishment as the initial structure, with apical meristem activity creating the central axis before lateral branch primordia initiate, establishing the organizational framework. Functional indispensability distinguishes the primary main rachis from higher-order structures, with primary main rachis representing the essential foundation without which no inflorescence architecture can exist, while individual lateral branches represent elaborations that can be absent or reduced without preventing inflorescence existence. Structural and vascular supremacy reflects the primary position with maximum dimensions and capacity. Architectural determination means the primary main rachis characteristics including length, diameter, and branching pattern establish the fundamental inflorescence form, with lateral elaboration modifying but not overriding the primary framework. Transformation to cluster maintains primary main rachis dominance with the central stem derived from the primary main rachis serving as the primary structural and vascular element throughout berry development.
erste Ordnung Zentralachse
first-order main axis
hierarchical main rachis
hierarchische Hauptspindel
primary central rachis
primäre Hauptrachis
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 183-218 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 359-398.
Primary Main Rachis
A primary median vein on a mature leaf at node 7 of Vitis vinifera cv. Cabernet Sauvignon in Bordeaux, France, serving as the principal first-order vein from which all major venation branches, measuring 14.5 cm length, diameter of 2.6 mm representing largest vein diameter on this leaf, directly continuous with the three main petiolar vascular bundles visible in cross-section showing unbroken vascular connection, originating five pairs of major first-order lateral veins at intervals of 2.2-3.5 cm along its length creating the five-lobed palmate architecture diagnostic for this cultivar, lateral veins branching at angles of 50-70 degrees from the primary median vein axis, containing 11 large xylem vessels (50-85 μm diameter) providing water transport capacity supporting maximum transpiration rate of 4.2 mmol H₂O m⁻² s⁻¹, extensive phloem tissue approximately 35% of total vascular cross-sectional area facilitating photosynthate export capacity of 0.95 g sucrose per day, thick-walled sclerenchyma forming continuous cap on both adaxial and abaxial sides providing mechanical strength preventing vein failure under maximum gravitational and wind loads, serving as axis of bilateral symmetry with slightly asymmetric lateral vein distribution (left side showing marginally larger lateral veins), representing the fundamental organizing principle of leaf venation architecture, measured at BBCH stage 77 (berries beginning to touch) in mid-July.
The median vein in its role as the principal first-order vein from which all major lateral vein systems branch in Vitis vinifera, characterized by hierarchical position as the apex of the venation architecture from which the complete reticulate vein network is organized, direct continuity with main petiolar vascular bundles establishing unbroken vascular connection from shoot to blade apex, origination point for four to six pairs of major first-order lateral veins radiating into the blade lobes creating the characteristic palmate venation pattern, coordinated development establishing the fundamental leaf architecture during early ontogeny, bilateral symmetry axis organizing left and right lateral vein systems, maximum vascular capacity among all blade veins with largest and most numerous xylem vessels and most extensive phloem, greatest mechanical strength providing primary structural support, and functional primacy as the dominant pathway for water delivery to the blade and photosynthate export from the blade, with failure or damage to the primary median vein potentially compromising entire leaf function unlike damage to individual lateral veins which affects only local regions.
Primary median vein emphasizes the hierarchical and organizational role of the median vein as the first-order vein from which the complete venation network is structured. The primary designation distinguishes this fundamental architectural role from secondary structural or functional aspects, highlighting the median vein's position at the apex of the hierarchical venation system. During development, the primary median vein differentiates first, establishing the fundamental axis along which lateral vein primordia subsequently initiate in coordinated patterns creating the final palmate architecture. The primary median vein exhibits maximum vascular capacity and structural strength reflecting its role as the main transport pathway and primary mechanical support, with dimensions, tissue organization, and functional capacity exceeding all other vein orders. Damage or dysfunction affecting the primary median vein potentially compromises entire leaf function, unlike damage to individual lateral veins affecting only their local supply zones, highlighting the critical importance of median vein integrity for leaf performance. Genetic and developmental programs controlling primary median vein formation and lateral vein organization determine fundamental leaf architecture including lobe number, positioning, and symmetry, with variations in these programs creating the morphological diversity characterizing different cultivars.
Haupt-Mittelrippe
main midrib
primary axis vein
primäre Hauptader
primäre Zentralvene
principal central vein
Based on Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 90-119 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 95-136.
Primary Median Vein
A primary replacement renewal cane on Vitis vinifera cv. Cabernet Sauvignon in Guyot-pruned vineyard, positioned immediately adjacent to cordon origin point, measuring 10mm diameter with uniform brown color indicating complete lignification, 12 nodes retained providing adequate length for anticipated fruiting cane selection, shoots from nodes 10-12 perfectly positioned along fruiting wire for horizontal cane placement in following season, representing ideal renewal wood that will almost certainly be selected as next season's fruiting cane during dormant pruning unless unexpected damage or quality issues emerge, while current fruiting cane located 15cm farther from cordon will be completely removed maintaining compact architecture.
A replacement renewal cane serving as first-choice or preferred option for following season's fruiting cane selection, characterized by optimal positioning closest to permanent structure, ideal vigor and diameter (8-12mm) indicating balanced growth, complete lignification ensuring winter survival and spring productivity, and highest probability of being selected as fruiting cane during next dormant pruning based on location, quality, and training system requirements. Primary replacement renewal canes represent best-case renewal scenario in cane-pruned systems, with successful selection and development of primary replacements indicating well-managed vineyards, appropriate vine balance, effective shoot management during growing season, and skilled pruning decisions maintaining optimal training system architecture and sustainable productivity through ideal wood renewal.
Primary replacement renewal canes represent the goal of renewal planning, with their consistent production indicating successful vineyard management integrating shoot selection, summer canopy work, and winter pruning decisions. The primary designation reflects both positional superiority (closest to desired fruiting zone) and quality attributes (diameter, lignification, node spacing) making selection nearly certain barring unforeseen problems. In well-managed vineyards, 80-90% of primary replacement renewals successfully transition to fruiting canes in following season, with occasional failures due to winter damage, poor spring growth, or repositioning needs requiring alternative selections from secondary replacements or reserve renewals, demonstrating importance of maintaining backup options despite primary replacement strategy.
bevorzugte Vorratsrute
first-choice replacement cane
optimal renewal position
preferred renewal cane
primäre Ersatzrute
Primary Replacement Renewal Cane
A primary reproductive bud at node 7 on a cane of Vitis vinifera cv. Pinot Noir, winter dissection revealing two fully formed inflorescence primordia containing 188 and 165 distinct flower initials with visible ovary and stamen primordia at 20x magnification, plus vegetative shoot primordia with 10 leaf initials, this bud having differentiated during previous June-July when the developing shoot occupied a well-exposed canopy position receiving 75% full sunlight for 8+ hours daily during the critical 3-week inflorescence initiation window, representing potential production of approximately 177 berries assuming typical 50% fruit set for Pinot Noir.
A primary bud containing one or more differentiated inflorescence primordia in addition to vegetative shoot structures, representing the principal source of grape crop production, characterized by complete flower initials with distinguishable ovary, stamen, and perianth primordia formed during previous season's June-August period (Northern Hemisphere), typical occurrence at mid-cane node positions (4-10) that received optimal light exposure during critical differentiation windows, and direct correlation between inflorescence number per bud and environmental conditions (light, temperature, water status, nitrogen) during previous season primordium formation. Primary reproductive buds determine potential yield through their abundance (percentage of buds with inflorescences), inflorescence number per bud (typically 1-3 in Vitis vinifera), and flower number per inflorescence (typically 50-400 depending on cultivar and conditions), with these parameters largely fixed 8-15 months before harvest, making historical growing conditions and bud dissection more relevant for yield prediction than current season observations alone.
Primary reproductive buds are the foundation of commercial grape production, with their fruitfulness (percentage bearing clusters) and cluster characteristics (size, compactness, berry number) determining vineyard economic viability. Fruitfulness assessment through winter bud dissection provides accurate yield predictions 3-4 months before budbreak, enabling informed decisions on crop load management, pruning severity, and cluster thinning requirements. The strong correlation between previous season light exposure and current season fruitfulness drives canopy management practices including shoot positioning, leaf removal, and hedging to optimize bud zone light penetration during the critical June-August differentiation period, with studies showing fruitfulness increasing from 30-40% in heavily shaded positions to 90-100% in well-exposed positions.
Fruchtknospe
fertile primary bud
flowering primary bud
fruchttragende Primärknospe
fruiting primary bud
reproduktive Hauptknospe
Based on Srinivasan & Mullins (1981) "Physiology of flowering in the grapevine - a review" American Journal of Enology and Viticulture 32=47-63 Buttrose (1974) "Climatic factors and fruitfulness in grapevines" Horticultural Abstracts 44=319-326 Sánchez & Dokoozlian (2005) "Bud microclimate and fruitfulness in Vitis vinifera L." American Journal of Enology and Viticulture 56=319-329 Vasconcelos et al. (2009) "The flowering process of Vitis vinifera= a review" American Journal of Enology and Viticulture 60=411-434.
Primary Reproductive Bud
A primary reproductive shoot on Vitis vinifera cv. Cabernet Sauvignon from node 7 on fruiting cane, broke at normal budbreak timing (mid-April), currently early August at veraison onset (BBCH 81), 156cm length with 18 leaves totaling approximately 2700 cm² leaf area, 2 well-developed clusters showing 48 and 52 berries with color change initiating, representing typical primary reproductive shoot performance with prompt budbreak, vigorous balanced growth, standard two-cluster load for this variety and position, and adequate photosynthetic capacity (16.4 cm² per gram projected fruit assuming 165g per cluster) supporting complete fruit maturation, managed through routine canopy operations without special interventions, contributing to whole-vine crop of 11.8kg (typical for this balanced cordon-trained mature vine).
A reproductive shoot originating from primary bud position within compound bud structure, characterized by development from the largest and most advanced bud component containing differentiated inflorescence primordia from previous season, typical prompt budbreak timing with primary shoots emerging 3-7 days before secondary positions, moderate to vigorous growth reflecting priority developmental status, and standard reproductive capacity with 1-3 inflorescences typical depending on variety and fruitfulness. Primary reproductive shoots represent normal productive growth from principal bud positions at mid-cane or spur locations with favorable light exposure during previous season's inflorescence differentiation period, accounting for majority of commercial crop production (typically 85-95% of total yield) through their numerical dominance, reliable reproductive performance, and superior vigor compared to secondary or tertiary backup positions.
Primary reproductive shoots dominate commercial wine grape production, with their consistent performance, numerical prevalence, and superior characteristics making them the foundation of viable viticulture. The primary designation indicates origin from main bud component that received priority resource allocation during differentiation, resulting in larger more developed buds, earlier budbreak, more vigorous growth, and higher reproductive potential compared to secondary or tertiary positions. Management focuses on optimizing primary shoot performance through appropriate winter pruning (retaining adequate primary bud positions based on balanced pruning principles), spring shoot thinning (removing excess or poorly positioned primaries while retaining optimal number and distribution), cluster management (adjusting crop load through thinning to match shoot capacity), and canopy work (maintaining architecture supporting fruit development and maturation). Understanding primary shoot characteristics and requirements is fundamental to achieving production targets and quality objectives across diverse viticultural contexts.
Hauptertragstrieb
first-order fruiting shoot
main position reproductive shoot
primary fruiting shoot
primärer Fruchttrieb
Based on Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Srinivasan & Mullins (1981) "Physiology of flowering in the grapevine - a review" American Journal of Enology and Viticulture 32=47-53.
Primary Reproductive Shoot
Merlot grafted on SO4 rootstock at 16 years showing primary root system comprising 4 primary roots originating from original cutting base, largest primary root 14 cm diameter at crown extending 165 cm depth approximately 35° from vertical, primary root showing extensive secondary xylem providing water transport capacity and mechanical support, bark with periderm protecting root surface, primary root producing lateral roots at 8-15 cm intervals creating branching network, primary root xylem conducting water from depth during summer drought periods, woody tissue storing carbohydrate reserves for spring mobilization
Primary root is the main structural root in grapevine root system originating either from radicle in seedling development or from cutting base in vegetatively propagated vines, typically 5-25 cm diameter at crown in mature vine depending on age and rootstock vigor, characterized by secondary growth with extensive xylem providing water transport and mechanical support, bark with protective periderm, and root cap at growing tip, showing primarily vertical or oblique orientation penetrating deep into soil profile, branching to produce lateral roots creating hierarchical root architecture, and serving as main structural axis anchoring vine, conducting water and minerals from deeper soil layers, storing substantial carbohydrate reserves in woody tissue, and providing framework from which lateral absorptive root network develops enabling extensive soil exploration and resource acquisition. Primary root origin and development includes= originating from radicle in seed germination (seedling primary root emerging from seed and growing downward), developing from cutting base in propagation (adventitious roots forming at nodes or callus on cuttings becoming functional primary roots), showing initial elongation (young primary root extending rapidly through apical meristem activity), undergoing secondary thickening (vascular cambium producing secondary xylem and phloem increasing diameter), and developing protective bark (cork cambium forming periderm protecting root surface). Primary root structural characteristics includes= showing large diameter (primary roots typically 1-3 cm diameter in young vine increasing to 5-25 cm in old vine at crown position), exhibiting extensive secondary xylem (woody tissue comprising majority of primary root volume providing water transport and structural support), containing functional phloem (inner bark conducting photosynthates from shoot system to root system), displaying protective periderm (cork layers preventing water loss and protecting from pathogens and mechanical damage), and demonstrating tap root architecture (single dominant vertical root or multiple primary roots from cutting extending deep into soil). Primary root orientation and depth includes= showing vertical to oblique growth (primary roots typically growing downward at 45-90° angles relative to surface), penetrating deep soil layers (primary roots potentially extending 100-200+ cm depth in permeable soils), responding to gravitropism (primary root showing positive gravitropic response growing downward), influenced by soil conditions (primary root direction and depth affected by obstacles, moisture gradients, and soil structure), and demonstrating rootstock variation (different rootstock varieties showing characteristic depth potential with Berlandieri types deep and Riparia types shallow). Primary root lateral branching includes= producing lateral roots (primary root generating branch roots along length creating hierarchical architecture), showing acropetal development (lateral roots typically initiating near root tip with older laterals progressively toward crown), exhibiting branching density (lateral root number per unit length varying with rootstock genetics and soil conditions), demonstrating branching angles (laterals emerging at characteristic angles typically 45-90° from primary root axis), and creating root system framework (primary root with laterals forming structural skeleton of root system). Primary root water transport function includes= conducting water from depth (primary root xylem moving water from deep soil through trunk to canopy), showing high conductivity (large-diameter xylem vessels in primary root providing efficient water transport pathway), enabling drought tolerance (deep primary roots accessing moisture unavailable to shallow roots), demonstrating seasonal variation (water transport highest during active growing season declining during dormancy), and contributing to vine water supply (primary root transport complementing lateral root uptake providing total vine water needs). Primary root anchorage function includes= providing mechanical stability (primary root penetration and lateral branching creating strong anchoring system), resisting uprooting forces (deep primary roots preventing wind or weight from toppling vine), bearing vertical loads (primary root structure supporting vine weight and transmitting forces to soil), demonstrating increased stability with age (older deeper primary roots providing greater anchorage than young shallow roots), and showing rootstock effects (vigorous deep-rooting rootstocks providing superior anchorage compared to shallow types). Primary root storage function includes= accumulating carbohydrate reserves (primary root parenchyma and ray cells storing starch particularly during dormancy), showing seasonal storage cycles (reserves highest in fall-winter declining in spring as mobilized for growth), providing substantial storage capacity (large primary root volume enabling storage of 200-1000+ grams dry weight carbohydrates depending on root size), supporting spring growth (stored reserves mobilized before leaves fully functional providing energy for bud break and initial shoot development), and enabling stress recovery (primary root reserves supporting vine recovery from damage, disease, or environmental stress). Primary root in propagated vines includes= developing from cutting base (adventitious roots at nodes or callus becoming primary roots in propagated vines), showing multiple primary roots (cuttings typically producing 3-8 primary roots from base rather than single tap root of seedling), exhibiting similar structure (cutting-derived primary roots developing secondary growth and characteristics comparable to seedling radicle-derived root), demonstrating functional equivalence (cutting primary roots serving same structural, transport, storage, and anchorage functions as seedling primary root), and varying with propagation method (rooting hormone treatment, callusing conditions, and planting practices affecting number and vigor of primary roots).
Primary_Root emphasizes main structural root originating from radicle or cutting and is distinguished from lateral roots (branches from primary) by focus on primary origin and main axis function. Includes Tap_Root (vertical orientation emphasis) and Main_Root_Axis (structural framework emphasis). Critical for understanding root system structure, deep water access, anchorage, carbohydrate storage, and the main structural framework providing root system foundation.
Hauptwurzel
Main root
Primärwurzel
Principal root
Structural root
Strukturwurzel
Primary Root
Cabernet Sauvignon grafted on 1103 Paulsen rootstock at establishment year showing primary root tip on main tap root, primary root tip 11.4 mm length with root cap 1.2 mm, apical meristem 0.7 mm showing numerous mitotic figures, elongation zone 7.8 mm, primary root tip demonstrating strong positive gravitropism maintaining vertical trajectory, elongation rate measured 8.6 mm per day during late May optimal conditions, primary root tip producing 4.2 mm diameter root reflecting vigorous 1103P growth characteristics, mucilage secretion evident, primary root tip penetrating through firm loam soil establishing deep root system
Primary root tip is root tip on primary root originating from radicle or cutting base, characterized by larger size typically 5-15 mm length reflecting vigorous growth of main structural roots, prominent root cap 0.8-1.5 mm protecting large active meristem, extensive apical meristem 0.5-0.8 mm with high cell division rate, substantial elongation zone 3-10 mm producing rapid length increase, and serving as growth engine for primary root system enabling deep penetration and main axis development, with primary root tip showing strong positive gravitropism directing primary root downward, high growth rate potentially 5-15 mm per day in young vigorous vine during favorable conditions, thick mucilage secretion facilitating penetration through dense soil layers, prominent quiescent center maintaining meristem organization and regenerative capacity, and representing most vigorous root tip type producing largest diameter roots with highest elongation rates essential for establishing deep extensive primary root framework during vine establishment and continued expansion throughout vine life. Primary root tip size and structure includes= showing large dimensions (primary root tip typically 5-15 mm total length substantially larger than secondary or tertiary root tips), exhibiting prominent root cap (cap 0.8-1.5 mm length with numerous columella cells providing robust protection), demonstrating extensive meristem (apical meristem 0.5-0.8 mm diameter with large population of dividing cells), displaying long elongation zone (zone 3-10 mm length enabling substantial daily length increase), and revealing substantial diameter (primary root tip producing root 2-8 mm diameter reflecting primary root size). Primary root tip growth characteristics includes= showing high division rate (primary root tip meristem cells dividing every 10-18 hours during peak activity), exhibiting rapid elongation (primary root tip potentially growing 5-15 mm per day in young vine under favorable conditions), demonstrating sustained growth (primary root tips maintaining active growth for weeks to months during growing season), revealing seasonal variation (primary root tip activity highest spring declining summer resuming fall ceasing winter), and displaying age-related slowing (primary root tip growth rate declining as vine ages though continuing throughout vine life). Primary root tip gravitropic response includes= demonstrating strong positive gravitropism (primary root tips showing pronounced downward growth response to gravity), containing abundant statoliths (primary root cap columella cells with numerous starch-filled amyloplasts sedimenting in gravity field), exhibiting rapid response (primary root tip reorienting within hours after displacement from vertical), showing curvature formation (differential cell elongation in elongation zone creating downward bend restoring vertical trajectory), and enabling depth penetration (gravitropic response ensuring primary roots grow deep into soil profile rather than lateral or upward). Primary root tip soil penetration includes= producing substantial force (primary root tip generating axial pressure 0.5-1.5 MPa pushing through soil), secreting copious mucilage (primary root cap producing abundant polysaccharide gel reducing friction and binding soil particles), forming penetration geometry (conical primary root tip shape optimizing force transmission and soil displacement), demonstrating persistent growth (primary root tip continuing elongation even through compacted layers if penetrable), and enabling deep exploration (primary root tip penetration capacity allowing primary roots to reach deep soil layers inaccessible to lateral roots). Primary root tip meristem organization includes= showing closed meristem type (grapevine primary root tip with separate root cap and apical meristem initials), exhibiting quiescent center (slowly dividing organizing cells at meristem apex maintaining stem cell populations), demonstrating initial patterns (distinct regions producing different root tissues with dermatogen for epidermis, periblem for cortex, plerome for stele), revealing cell lineages (traceable lineages from meristem initials through differentiated tissues documenting developmental progression), and displaying regenerative capacity (primary root tip meristem capable of recovering from damage and resuming growth if quiescent center intact). Primary root tip in vine establishment includes= providing critical early growth (primary root tips from germinating seed or rooting cutting producing initial root system), enabling rapid soil exploration (vigorous primary root tip growth quickly expanding root system during establishment year), demonstrating establishment success dependency (vine establishment critically dependent on primary root tip function producing adequate root system), showing transplant sensitivity (primary root tip damage during transplanting potentially compromising establishment), and revealing management importance (protecting primary root tips during planting and early growth critical for successful vineyard establishment). Primary root tip seasonal activity includes= initiating spring growth early (primary root tips often beginning growth before bud break utilizing stored carbohydrate reserves), maintaining summer activity (primary root tips continuing growth during summer though potentially reduced during peak shoot and fruit growth), demonstrating fall growth pulse (primary root tips showing renewed vigorous growth late season as shoot growth ceases), entering winter dormancy (primary root tips ceasing activity during cold periods remaining quiescent until spring), and showing temperature dependence (primary root tip activity requiring soil temperature typically >8-10°C for initiation >15°C for optimal growth). Primary root tip and rootstock effects includes= varying with rootstock genetics (vigorous rootstocks like 110R, 1103P producing more active primary root tips than low-vigor types like Riparia), demonstrating depth potential differences (Berlandieri-based rootstocks showing primary root tips maintaining activity to greater depths than Riparia types), revealing growth rate variation (rootstock genetics affecting primary root tip elongation rate influencing total root system extent), showing branching effects (primary root tip activity affecting lateral root initiation along primary root influencing root system architecture), and determining drought tolerance (primary root tip depth penetration capacity critical component of rootstock drought adaptation).
Primary_Root_Tip emphasizes root tip on primary root from radicle/cutting and is distinguished from secondary/tertiary root tips by focus on main structural root growth. Includes Active_Primary_Root_Tip and Growing_Primary_Root_Tip. Critical for understanding primary root elongation, deep penetration, gravitropic control, vigorous growth, establishment success, and the most active root tip type producing main structural framework essential for root system development.
Hauptwurzel-Apex
Main root apex
Primary root growing point
Primäre Wurzelspitze
Structural root tip
Strukturwurzel-Spitze
Primary Root Tip
Cabernet Sauvignon primary vascular bundle mesocarp showing typical 4-bundle organization= 4 major dorsal bundles positioned at approximately 90° intervals (12 o'clock, 3 o'clock, 6 o'clock, 9 o'clock positions when viewing berry cross-section), each 165-245 μm diameter, containing xylem (75-115 μm, 4-6 major vessels 32-48 μm diameter, non-functional at maturity) and phloem (55-95 μm, 12-18 sieve elements 18-26 μm diameter, functional during ripening), with primary bundles conducting estimated 75-85% of total berry sugar import during mid-ripening period (weeks 12-16) supporting 1.1% sugar accumulation per week. Syrah primary vascular bundle mesocarp showing 5-bundle variant= 5 dorsal bundles at 72° intervals creating pentagonal vascular symmetry, each 145-220 μm diameter, providing import capacity supporting large berry size (16-17.5 mm diameter, 3.2-3.8 g at maturity) through enhanced vascular tissue volume (5 bundles versus typical 4) creating 20-25% greater total primary vascular bundle cross-sectional area potentially supporting higher import rates. Merlot primary vascular bundle mesocarp phloem import quantification= primary bundle phloem with 14-19 sieve elements per bundle, sieve element cross-sectional area 180-320 μm² each, estimated sap velocity 25-45 cm/hour during active ripening, calculated flow capacity 2.5-4.2 μL sap per hour per bundle delivering 0.3-0.75 mg sucrose per hour per bundle or 1.2-3 mg per hour total across 4 bundles, supporting observed sugar accumulation rate of 0.05-0.1 g per berry per day during mid-ripening (week 12-15) demonstrating primary vascular bundle mesocarp import capacity matching berry compositional development requirements. Pinot Noir primary vascular bundle mesocarp xylem cavitation at veraison= pre-veraison (week 9, BBCH 79) showing functional xylem vessels conducting water with sap flow detected, vessels appearing clear in microscopy veraison (week 10, BBCH 81) showing beginning cavitation with some vessels embolized (air-filled), mixed functional/non-functional state post-veraison (week 11-12, BBCH 83) showing complete xylem cavitation with all major vessels air-filled and non-conducting, phloem remaining functional, demonstrating xylem-to-phloem transition in primary vascular bundle mesocarp at veraison creating shift from dual import pathway (xylem plus phloem) to phloem-exclusive import supporting ripening phase. Chardonnay primary vascular bundle mesocarp secondary branch production= each primary bundle producing 8-15 visible secondary lateral branches along its length through mesocarp, with branch points approximately 800-1500 μm apart along bundle, creating total 32-75 secondary branches from 4 primary bundles distributing throughout mesocarp tissue, with branching pattern providing mesocarp cells within <500 μm of vascular tissue ensuring effective import distribution from primary vascular bundle mesocarp to entire storage parenchyma tissue volume.
A primary vascular bundle mesocarp is vascular bundle mesocarp containing or adjacent to major (primary, dorsal) vascular bundles representing the main longitudinal import pathways running from pedicel attachment through berry to stylar end, typically comprising 3-5 major bundles (most commonly 4) positioned at approximately 90-120° intervals around berry circumference, each 120-300 μm diameter at maturity, serving as primary conduits for water, sugar, and mineral import from plant to berry during development and ripening. Primary vascular bundle mesocarp structural organization= (1) major dorsal bundles oriented longitudinally running from receptacle (pedicel attachment point) through mesocarp to stylar scar (flower remnant at berry apex), creating continuous vascular pathways from plant to berry apex (2) bundles positioned circumferentially around berry typically at 90° intervals when 4 bundles present (most common), 72° when 5 bundles, 120° when 3 bundles, creating relatively symmetric distribution around berry (3) bundle diameter typically 150-280 μm in wine cultivars, potentially larger (200-350 μm) in large-berried table cultivars (4) xylem portion 60-140 μm diameter containing 3-8 major vessels 25-50 μm diameter plus smaller vessels and xylem parenchyma (5) phloem portion 50-120 μm diameter containing 8-20 sieve elements 14-28 μm diameter plus companion cells and phloem parenchyma. Primary vascular bundle mesocarp as main import pathway= (1) phloem in primary bundles conducting majority (>70-85%) of sugar import during ripening with high sieve element density and large diameter supporting high flow capacity (2) phloem sap velocity estimated 20-60 cm/hour during active ripening (mid-Phase II) delivering sugars at rates supporting 0.8-1.5% sugar accumulation per week in ripening berries (3) primary bundle phloem connecting directly to pedicel vascular system receiving import from source leaves without interruption (4) xylem in primary bundles functional during Phase I providing water import pathway supporting early berry growth before cavitation at/before veraison eliminates xylem function. Primary vascular bundle mesocarp lateral branching= (1) secondary vascular bundles branching from primary bundles at irregular intervals along bundle length creating distribution network throughout mesocarp (2) branch points showing vascular tissue division with some xylem vessels and phloem sieve elements diverting into lateral branches while others continue in primary bundle (3) lateral branches smaller diameter (30-120 μm) than primary bundles creating hierarchical vascular network (4) branching pattern creating relatively uniform mesocarp tissue access to vascular import with maximum distance from vascular tissue typically <400-800 μm in central mesocarp. Primary vascular bundle mesocarp in berry shape and expansion= (1) primary bundles potentially constraining berry expansion in longitudinal direction creating mechanical resistance to elongation (2) berry shape (round versus elongated) potentially influenced by primary bundle mechanical properties with stiffer bundles favoring round shape, more flexible bundles allowing elongation (3) uneven expansion among bundle sectors potentially creating irregular berry shapes if some primary bundles resist expansion more than others (4) primary bundle distribution affecting berry symmetry with asymmetric bundle arrangement potentially contributing to asymmetric berry shape.
Primary_Vascular_Bundle_Mesocarp represents tissue containing or adjacent to major longitudinal vascular bundles and is distinguished from Secondary_Vascular_Bundle_Mesocarp (lateral branches) by bundle size, position, and primary import function. Primary vascular bundle mesocarp represents the main import highways in the berry vascular network. Understanding primary vascular bundle mesocarp is essential for comprehending berry import physiology, for detailed berry anatomical characterization, and for understanding potential vascular limitations on berry growth and composition development.
Dorsalbündel-Gewebe
Hauptleitbündel-Fruchtfleisch
dorsal bundle zone
major vascular bundle tissue
primäres Gefäßbündel-Mesokarp
Rogiers, S.Y. et al. (2006). Vascular function in berries. Australian Journal of Grape and Wine Research 12=203-214 Chatelet, D.S. et al. (2008). Xylem structure and connectivity in grapevine shoots provides a passive mechanism for the spread of bacteria in grape plants. American Journal of Botany 95=1076-1085 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 193-198.
Primary Vascular Bundle Mesocarp
Syrah primary vascular connection zone at mid-ripening (BBCH 85) contains 3 main vascular bundles with xylem vessels 35-45 μm diameter (partially embolized, reduced hydraulic conductance 40-60% compared to Phase I), phloem sieve tubes occupying 25-35% of bundle cross-section actively transporting 1.2-1.8 mg sucrose/berry/hour, bundles positioned symmetrically around pedicel periphery providing uniform berry vascular supply
A primary vascular connection zone is the main structural vascular pathway within the pedicel emphasizing the principal xylem and phloem bundles that provide the primary transport conduits connecting the rachis vascular system to the berry, representing the major vascular architecture supporting berry development and ripening. Primary vascular connection zone typically consists of 2-4 main vascular bundles (fewer bundles in small-berried cultivars like Pinot Noir, more bundles in large-berried cultivars like table grapes) that run longitudinally through the pedicel from rachis attachment to berry receptacle. Structural characteristics of primary vascular connection zone include= large-diameter xylem vessels (30-50 μm diameter in primary bundles compared to 15-30 μm in secondary vascular traces), abundant phloem sieve tubes with numerous companion cells (providing high transport capacity), substantial vascular bundle size (primary bundles may occupy 20-40% of pedicel cross-sectional area), sclerenchymatous bundle sheath providing mechanical support, and strategic positioning (primary bundles typically arranged symmetrically around pedicel periphery providing uniform berry vascular supply). Primary vascular connection zone functional capacity includes= major water transport pathway during berry expansion (primary xylem vessels may account for 70-90% of total pedicel hydraulic conductance during Phase I growth), principal sugar transport route during ripening (primary phloem may account for 80-95% of total sugar import during Phase III), nutrient transport (minerals, amino acids, hormones moving through primary vascular pathway), and hydraulic signaling (water potential gradients transmitted through primary xylem affecting berry growth regulation). Primary vascular connection zone development parallels berry development= during fruit set (BBCH 71-73) primary bundles differentiate and expand from flower pedicel vasculature, during Phase I growth (BBCH 73-77) primary xylem vessels expand to maximum diameter and phloem differentiates, during Phase II (BBCH 77-81) primary bundles undergo lignification and structural reinforcement, during Phase III ripening (BBCH 81-89) primary xylem may show progressive embolism while phloem remains active. Primary vascular connection zone relationship to berry size includes= cultivars with larger/more numerous primary vascular bundles generally produce larger berries (greater hydraulic and nutrient supply capacity), vessel diameter correlates with berry expansion rate during Phase I (larger vessels support faster water import and cell expansion), and phloem capacity in primary bundles correlates with sugar accumulation rate during ripening (greater phloem cross-sectional area supports higher sugar import). Primary vascular connection zone modifications during stress or maturation include= drought stress induces xylem embolism in primary vessels (reducing hydraulic conductance and limiting berry expansion), advanced ripening associated with tyloses formation or pit membrane blockage in primary xylem (contributing to berry hydraulic isolation), and potential phloem callose deposition affecting sugar transport capacity. Distinction from secondary vascular traces includes= primary bundles are larger diameter, have greater conducting capacity, develop earlier during fruit set, maintain activity longer into ripening, and provide majority of berry vascular supply.
Primary_Vascular_Connection_Zone emphasizes main structural conduits and is distinguished from Vascular_Connection_Zone (complete vascular architecture) and Conducting_Vascular_Connection_Zone (active transport emphasis) by focus on principal vascular bundles providing majority of transport capacity. Critical for understanding berry size potential, cultivar differences in vascular architecture affecting berry characteristics, xylem embolism impacts on berry water relations, and breeding for berry size or composition traits linked to vascular capacity.
Haupt-Transportbahnen
Hauptleitbündel
Main vascular pathway
Primäre Gefäßverbindungszone
Principal conducting bundles
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Chatelet, D. S., Rost, T. L., Shackel, K. A., & Matthews, M. A. (2008). The peripheral xylem of grapevine (Vitis vinifera). 1. Structural integrity in post-veraison berries. Journal of Experimental Botany, 59(8), 1987-1996.
Primary Vascular Connection Zone
A primary vegetative bud at node 2 on a cane of Vitis vinifera cv. Cabernet Sauvignon, dissected showing well-developed vegetative shoot primordia with 10 leaf initials clearly visible, two tendril primordia at appropriate phyllotactic positions, but complete absence of inflorescence primordia despite favorable vine health and adequate growing degree day accumulation during previous season, reflecting typical low fruitfulness of basal node positions that were heavily shaded during critical inflorescence differentiation period when canopy development obscured basal regions from direct sunlight exposure.
A primary bud containing exclusively or predominantly vegetative primordia without differentiated inflorescence structures, characterized by absence of floral initials visible upon dissection, development into shoots bearing only leaves and tendrils, typical occurrence at basal node positions (nodes 1-3) or in conditions of severe shading during previous season's critical inflorescence initiation period (June-July in Northern Hemisphere). Primary vegetative buds represent non-fruiting positions within the compound bud hierarchy, contributing to canopy leaf area and photosynthetic capacity but not to current season yield, with their prevalence at specific node positions influencing pruning strategies and retention formulas that account for expected percentage of non-fruiting buds in different cane or spur positions.
Primary vegetative buds are most common at basal nodes (1-3) where shading during inflorescence initiation is nearly universal, but can occur at any node position under conditions of dense canopy, poor light penetration, excessive nitrogen availability, or water stress during the critical differentiation window. The proportion of primary vegetative buds versus primary reproductive buds varies dramatically by cultivar, with some varieties showing inherently low basal fruitfulness (e.g., Cabernet Sauvignon, Merlot) while others maintain reproductive capacity even at basal positions under favorable conditions (e.g., Chardonnay, Riesling). Spur pruning systems must account for higher percentages of primary vegetative buds compared to cane pruning systems that selectively retain more fruitful mid-cane positions.
nicht-fruchtende Primärknospe
non-fruiting primary bud
sterile primary bud
vegetative Hauptknospe
vegetative primary bud
Based on Buttrose (1970) "Fruitfulness in grapevines= The response of different cultivars to light, temperature and day length" Vitis 9=121-125 Sánchez & Dokoozlian (2005) "Bud microclimate and fruitfulness in Vitis vinifera L." American Journal of Enology and Viticulture 56=319-329 Vasconcelos et al. (2009) "The flowering process of Vitis vinifera= a review" American Journal of Enology and Viticulture 60=411-434.
Primary Vegetative Bud
A primary vegetative shoot on Vitis vinifera cv. Chardonnay from node 2 on retained cane in Guyot system, broke promptly at normal spring budbreak timing (early April), currently 135cm length in early July with 17 leaves, 3 tendrils, vigorous growth typical of well-developed primary bud origin, no inflorescences reflecting typical low fruitfulness of node 2 position despite primary bud status, shoot positioned vertically in VSP trellis contributing approximately 1050 cm² leaf area (significant photosynthetic capacity) though no current crop, with buds differentiating on this shoot through summer potentially productive for following season if adequate light exposure achieved through canopy management.
A vegetative shoot originating from primary bud position within compound bud structure, characterized by development from largest and most advanced bud component, typical origin from basal node positions (nodes 1-3) with inherently low fruitfulness or from mid-cane positions where primary buds lacked inflorescence differentiation despite normally productive positions, and vigor typically moderate to strong reflecting priority developmental status of primary buds over secondary or tertiary positions. Primary vegetative shoots represent vegetative growth from the principal bud position, most commonly occurring at basal nodes where shade during previous season's inflorescence initiation period prevented reproductive differentiation, occasionally at mid-cane positions where unusual conditions (severe stress, late-season growth flushes, cultivar characteristics) prevented normal inflorescence formation in otherwise productive node positions.
Primary vegetative shoots are distinguished from secondary or tertiary vegetative shoots by bud origin (primary versus backup positions) rather than by vigor or function which may overlap. The primary designation indicates the shoot arose from the main bud component that would normally produce fruiting shoots at favorable node positions, with vegetative-only outcome reflecting either positional effects (basal nodes universally low-fruitfulness) or unusual circumstances preventing normal reproductive differentiation. Primary vegetative shoots from basal positions are expected and normal, while primary vegetative shoots from mid-cane positions may indicate problems (severe previous-season stress, cultivar issues, management errors) requiring investigation. Management typically retains well-positioned primary vegetative shoots for photosynthetic contribution and potential renewal wood while removing poorly-positioned or redundant shoots regardless of primary status.
Haupttrieb vegetativ
first-order vegetative shoot
main position vegetative shoot
primary non-fruiting shoot
primärer vegetativer Trieb
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Srinivasan & Mullins (1981) "Physiology of flowering in the grapevine - a review" American Journal of Enology and Viticulture 32=47-63 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115.
Primary Vegetative Shoot
Cabernet Sauvignon primary wood ray at 18 years post-planting shows continuous radial extent 42-58 mm from near-pith origin through all 18 annual rings to cambium, multiseriate structure 3-5 cells wide maintained through radial extent, living ray parenchyma in outer 18-24 mm sapwood zone storing starch at 7-10% dry weight and providing active radial transport, dead ray parenchyma in inner 20-35 mm heartwood zone providing structural support, ray representing original cambial ray initial established during trunk initiation
A primary wood ray is a wood ray initiated directly from primary ray tissues during primary growth or established early in secondary growth from original ray initials in young vascular cambium, representing rays present from initial trunk development that extend continuously through all subsequent annual rings as cambium produces additional xylem. Primary wood rays originate= from primary ray tissues in primary xylem (rays present in shoot vascular cylinder before secondary growth begins), from initial ray initials established when vascular cambium forms (earliest cambial ray initials producing first secondary xylem rays), persisting through trunk life (primary rays extending from inner secondary xylem near pith through all annual rings to cambium), and maintaining continuity (same ray file connecting inner rings to outer sapwood representing decades of continuous ray tissue). Primary wood ray anatomical position includes= origin near pith (primary rays visible in innermost secondary xylem rings adjacent to primary xylem), radial continuity through trunk (extending through all annual rings from inner heartwood through transition wood to outer sapwood), termination at cambium (ray extending to cambial zone where living ray parenchyma continues into phloem rays), connection to primary structures (primary rays potentially connecting to primary phloem rays through cambial continuity), and presence throughout trunk circumference (primary rays distributed around entire trunk cross-section reflecting early cambial organization). Primary wood ray characteristics include= long radial extent (extending 30-80 mm radius in 15-25 year trunks, 60-150+ mm in old 40-60 year trunks), passage through sapwood and heartwood zones (living ray parenchyma in sapwood 15-30 mm outer portion, dead parenchyma in inner heartwood 20-100+ mm), typical multiseriate organization (most primary rays 3-5 cells wide though width variable), height 15-35 cells (600-1400 μm vertical extent), and maintained width (rays generally maintaining 3-5 cell width through radial extent though sometimes becoming wider in outer rings). Primary wood ray functions throughout trunk depth include= historical radial transport pathway (providing lateral transport across entire trunk depth when parenchyma was alive), current function in sapwood zone (outer 15-30 mm of primary ray containing living parenchyma providing active transport and storage), historical structural record (primary ray anatomy preserving information about early cambial organization and growth conditions), and mechanical support in heartwood (dead primary ray tissue contributing to structural integrity of inner trunk). Primary wood ray aging along radial extent includes= outer sapwood zone (15-30 mm) with living metabolically active ray parenchyma (actively transporting, storing starch, responding to signals), transition zone (5-10 mm) where ray parenchyma progressively dies (declining metabolic activity as cells age 8-12 years), inner heartwood zone (20-100+ mm) with dead ray parenchyma (no metabolic activity, cell walls persist providing structure but lumens often degraded or filled with deposits). Primary wood ray significance includes= representing original cambial organization (primary rays reflecting initial pattern of ray initials established during juvenile cambial development), providing complete radial transport history (rays extending through entire trunk recording decades of lateral transport capacity evolution), indicating trunk age (innermost primary ray origin corresponding to trunk establishment allowing age estimation), and contributing to trunk structural integrity (primary rays extending through heartwood providing radial connectivity enhancing mechanical properties). Primary wood ray identification includes= visual observation in radial section (primary rays appearing as continuous radial bands extending from pith region to cambium), continuity verification (tracing ray through multiple annual rings confirming uninterrupted radial extent), position near primary xylem (primary rays beginning in inner rings adjacent to pith indicating early origin), and distinction from late-formed rays (comparing with rays initiating in outer rings that show limited radial extent). Primary wood ray density implications= high primary ray density (10-12 rays/mm tangential in inner rings) indicating favorable early conditions allowing extensive ray initial establishment, moderate density (7-9 rays/mm) typical for most cultivars, lower density (5-7 rays/mm) potentially indicating stress or rapid early growth with reduced ray initiation, and primary ray density generally matching or slightly exceeding density in outer rings.
Primary_Wood_Ray emphasizes rays present from early development and is distinguished from Secondary_Wood_Ray (later-initiated rays) and Wood_Ray (all rays regardless of origin) by continuous radial extent from inner trunk through all rings. Critical for understanding complete trunk radial transport history, cambial organization evolution, trunk age assessment, and structural continuity connecting inner heartwood to outer sapwood.
Continuous ray
Early-formed ray
Kontinuierlicher Xylemstrahl
Original ray
Primärer Holzstrahl
Ursprünglicher Strahl
Carlquist, S. (2001). Comparative Wood Anatomy (2nd ed.). Springer. | Pratt, C. (1974). Vegetative anatomy of cultivated grapes - a review. American Journal of Enology and Viticulture, 25(3), 131-150.
Primary Wood Ray
The prime name Savagnin Blanc designates the cultivar known under more than 80 synonyms worldwide including Traminer (Austria/Germany), Heida (Valais, Switzerland), Formentin (Hungary), and the parent of Grüner Veltliner — the VIVC lists Savagnin Blanc as the accepted prime name (VIVC number 10803) while the OIV registers it separately, and the French national catalogue uses Savagnin as the official designation, illustrating how different nomenclatural authorities may assign slightly different prime names to the same genotype.
A cultivar designation that constitutes the single authoritative accepted name for a grapevine cultivar within a given nomenclatural authority or registration system, serving as the primary reference identifier to which all synonyms, alternative spellings, and regional variants are formally linked.
Prime_Name establishes the canonical reference point in the many-to-one relationship between cultivar designations and genotypes. The concept of a single accepted name per cultivar follows ICNCP principles but is complicated in practice by the existence of multiple independent nomenclatural authorities (OIV, VIVC, national registries) that may designate different prime names for the same cultivar: for example, the VIVC prime name may differ from the OIV registered name or the name in a national variety catalogue. Subclasses capture the three principal sources of authoritative prime name designation.
Leitname
Primärbezeichnung
accepted name
akzeptierter Sortenname
primary cultivar name
standard variety name
ICNCP (2016) International Code of Nomenclature for Cultivated Plants 9th Ed, ISHS Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157 OIV (2013) OIV List of Vine Varieties and Their Synonyms
Prime Name
A primordium stage aborted tendril at node 2 on a shoot of Vitis vinifera cv. Chardonnay in Chablis, France, appearing as a 0.8 cm diameter slightly raised brown bump at the expected tendril position, showing no elongation or morphological differentiation beyond the initial meristematic protrusion, occurring during an extreme late spring frost event (-4°C for 6 hours) in early May that killed shoot apical meristems and halted all organogenesis, with microscopic examination revealing only undifferentiated parenchyma cells and necrotic tissue without any vascular elements or organized cell layers, part of a whole-vineyard pattern where 85% of shoots showed primordium stage abortion of multiple organ types including tendrils, leaves, and inflorescences at nodes developing during the frost event, requiring severe pruning to remove damaged shoots and encourage development of replacement shoots from secondary buds.
An early aborted tendril that ceased development at the primordium stage before any significant cell expansion or tissue differentiation has occurred in Vitis vinifera, characterized by presence of a barely visible meristematic bump or protrusion of 0.5-1.5 cm at the node position, composed almost entirely of undifferentiated meristematic or parenchyma cells without distinct tissue layers, absence of vascular differentiation or any organized conducting tissue, typically showing rapid necrosis and browning within days of developmental arrest, and representing the earliest recognizable stage of tendril abortion where organ identity has been specified but no morphological differentiation has proceeded. Primordium stage aborted tendrils indicate extremely severe stress conditions or fundamental developmental failures affecting the shoot apical meristem during the critical period of organ primordium initiation, occurring when resource availability or hormonal signaling is insufficient to support even minimal organogenesis beyond the initial cell division events that establish organ position. These abortion events typically correlate with severe whole-vine stress conditions including extreme drought, severe nutrient deficiency, root system damage, or pathogen infection affecting systemic physiology, and often occur in conjunction with abortion or severe malformation of other organs including leaves and inflorescences emerging from adjacent primordia. The metabolic investment lost in primordium stage abortion is minimal, representing only the initial meristematic activity, but the occurrence signals critical physiological problems requiring immediate diagnostic attention and corrective intervention to prevent complete shoot development failure.
Primordium stage aborted tendrils represent the absolute earliest detectable stage of tendril abortion, distinguished from complete absence of tendril development by the presence of at least a meristematic bump or protrusion at the expected node position, and distinguished from initial growth stage abortion by lack of any cell expansion or tissue differentiation. Detection requires very early examination of developing shoots, as primordium stage aborted structures may become completely indistinguishable from node tissue within 1-2 weeks through tissue necrosis and collapse. This abortion class is relatively rare in commercial viticulture except during catastrophic events (extreme frost, severe disease, major root damage) that compromise fundamental meristematic function. The occurrence of primordium stage abortion across multiple organ types simultaneously is pathognomonic for shoot apical meristem failure requiring shoot replacement rather than stress amelioration.
Primordialstadium-abortierte Ranke
earliest stage failed tendril
frühestes Stadium fehlgeschlagene Ranke
meristematic bump abortion
meristematische Höcker-Fehlbildung
primordial protrusion abortion
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 43-59 Carmona et al. (2008) "Grapevine bud dormancy= environmental and molecular aspects" Revista Brasileira de Fruticultura 30=257-408 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 353-376.
Primordium Stage Aborted Tendril
A primordium stage young leaf on the shoot apical meristem of Vitis vinifera cv. Syrah in the Rhône Valley, France, appearing as a small 0.8 cm length protrusion positioned 6 mm behind the apical meristem dome, representing the third visible primordium in the phyllotactic spiral, exhibiting very pale yellowish coloration with bronze tinge particularly at margins, composed entirely of small densely cytoplasmic cells with prominent nuclei showing active mitotic divisions (mitotic index 18% indicating rapid cell proliferation), no visible tissue layering or differentiation under dissecting microscope examination, procambial strands barely distinguishable as slightly darker lines representing future major veins, no stomatal structures evident on either surface, complete absence of cuticle leaving cells vulnerable to desiccation requiring humid environment within apical bud protection, representing pure resource sink consuming approximately 8 mg carbohydrate per day for cell division without any photosynthetic return, occurring at approximately 2 days post-initiation from apical meristem, measured at shoot BBCH stage 51 (inflorescence swelling) in early May, representing the absolute earliest recognizable leaf developmental stage observable without destructive meristem sectioning.
An emerging young leaf at the earliest recognizable developmental stage as a distinct primordium on the shoot apical meristem in Vitis vinifera, characterized by extremely small size typically 0.5-1.5 cm length representing 3-10% of final mature dimensions, position on the apical meristem dome typically within 5-10 mm of the apex, appearance as a small bump, ridge, or emerging protrusion with minimal lamina development, tightly compressed architecture with future lamina completely folded and unexpanded, very pale yellow-green to bronze coloration from minimal chloroplast development and protective anthocyanin accumulation, composed almost entirely of actively dividing meristematic cells with large nuclei and dense cytoplasm, complete absence of tissue differentiation with uniform parenchymatous cellular organization, no functional stomata or vascular elements beyond procambial strands, no cuticle development leaving cells directly exposed, and complete absence of photosynthetic function representing pure metabolic sink consuming resources for rapid cell division establishing the basic cellular population and architectural framework that will determine final leaf characteristics. Primordium stage young leaves represent the earliest observable leaf developmental phase, initiated through activation of lateral meristems on the apical dome in regular phyllotactic patterns, with genetic programming and positional signals determining basic leaf identity including eventual lobe number and pattern, followed by intense cell division over 3-5 days establishing the foundational cell population, with critical developmental decisions including sun versus shade leaf differentiation beginning during this phase through sensing of light quality and quantity by cryptochrome and phytochrome photoreceptors even in these primitive tissues.
Primordium stage young leaves represent the foundational phase of leaf ontogeny, distinguished from unfolding and later stages by extremely small size, complete lack of differentiation, pure meristematic cellular composition, and position on the apical dome. Observation requires careful dissection of apical buds and often magnification, as primordia are typically hidden within protective scales and surrounding older leaf primordia. The primordium stage is when fundamental architectural decisions are established including lobe number and arrangement, vein patterns, and eventual leaf size potential, with these characteristics determined by genetic programming interacting with environmental signals. Cell division rates during primordium stage are among the highest in the plant, with plastochron (time between successive leaf initiations) typically 2-4 days under optimal conditions, creating the continuous production of new primordia sustaining shoot extension. Environmental conditions during primordium stage including temperature, light quality, and resource availability affect fundamental leaf characteristics that persist through maturity, with low red=far-red light ratios sensed during this phase triggering shade leaf developmental programs including reduced cell division leading to thinner leaves.
Blattprimordium
initial primordium
initiales Primordium
leaf primordium
meristematic bump
meristematische Auswölbung
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Tsukaya (2013) "Leaf development" The Arabidopsis Book 11=e0163 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 56-78.
Primordium Stage Young Leaf
The sequence of principal growth stage codes observed for Pinot Noir in Burgundy across a typical growing season proceeds as 0 (sprouting/bud development, March-April), 1 (leaf development, April-May), 5 (inflorescence emergence, May), 6 (flowering, June), 7 (development of fruits, June-July), 8 (ripening of berries, August-September), and 9 (senescence/dormancy, October-November) — principal stages 2, 3, and 4 are not defined for grapevine in the Lorenz et al. adaptation, reflecting the absence of tillering, stem elongation, and booting phases that occur in monocotyledonous crops for which those codes were originally designed.
A BBCH code component represented by the first digit (0-9) of the two-digit BBCH code, designating one of ten major developmental phases in the grapevine annual growth cycle from winter dormancy through post-harvest senescence, each encompassing a distinct category of morphological and physiological events.
Principal_Growth_Stage_Code defines the highest-level temporal organization of the grapevine annual cycle into discrete macro-phases. The grapevine BBCH adaptation uses 7 of the 10 available principal stages (0, 1, 5, 6, 7, 8, 9), with stages 2, 3, and 4 reserved but undefined for Vitis: this reflects the structural mapping of the general BBCH framework (designed to accommodate monocots and dicots) onto grapevine-specific phenology. Each principal growth stage code groups a set of secondary stages that describe the progressive morphological changes within that developmental phase. Subclasses in this ontology represent each grapevine-relevant principal growth stage as a named class.
BBCH principal stage
BBCH-Hauptstadium
Hauptentwicklungsstadien-Code
Makrostadium-Kennung
macro-stage code
primary growth stage number
Principal Growth Stage Code
Production statistics for Cabernet Sauvignon document the world's most widely planted red wine grape — global acreage approximately 340,000 ha (OIV 2023), distributed across France (approximately 50,000 ha), Chile (approximately 40,000 ha), China (approximately 38,000 ha, though data reliability varies), the United States (approximately 37,000 ha), Australia (approximately 25,000 ha), Argentina (approximately 16,000 ha), and South Africa (approximately 11,000 ha), with production volume exceeding 20 million hL annually and a generally stable global acreage trend over the past decade masking regional dynamics (expansion in China, contraction in Australia). Production statistics for Blaufränkisch document a geographically concentrated cultivar — total global acreage approximately 8,500 ha, of which Austria accounts for approximately 3,200 ha (predominantly Burgenland), Hungary approximately 4,500 ha (as Kékfrankos, mainly Sopron and Eger), with smaller plantings in Germany (as Lemberger, approximately 500 ha in Württemberg), Croatia, Serbia, and the United States (approximately 50 ha).
Production statistics is an economic data entity that documents the quantitative measures of grape cultivar cultivation extent and wine production output, encompassing global acreage (worldwide vineyard area planted to a specific cultivar), regional acreage (vineyard area by country, region, or appellation), production volume (wine output in hectoliters or equivalent units), and production trends (temporal trajectories of acreage and volume change), derived from national vineyard registers, governmental agricultural censuses, international statistical compilations, and industry surveys, providing the fundamental quantitative indicators of cultivar commercial importance that inform planting decisions, market analysis, and viticultural policy.
Production statistics provide the quantitative backbone of viticultural economic analysis — the fundamental measures of how much of each cultivar is planted where and how much wine it produces. The data quality hierarchy for production statistics reflects the institutional infrastructure of national viticultural sectors — countries with comprehensive vineyard registers linked to individual grower records (France's Casier Viticole Informatisé, Austria's Weinbaukataster, Australia's Wine Australia register) provide high-resolution, cultivar-specific data updated annually, while countries relying on periodic agricultural censuses or industry surveys provide less frequent and less precise estimates. The Anderson and Nelgen (2020) global database represents the most comprehensive compilation effort, harmonizing data from approximately 50 wine-producing countries covering over 99% of global vineyard area, though data gaps persist for several significant producing countries (notably China, where official vineyard statistics are considered unreliable by most international analysts). The distinction between planted area (total vineyard area including non-bearing young vines and temporarily non-productive older vines), bearing area (vineyard area in active commercial production), and harvested area (bearing area actually harvested in a given year, which may differ due to crop losses, intentional non-harvest, or abandonment) creates measurement precision issues that affect the comparability of acreage statistics across jurisdictions employing different definitions. The relationship between acreage and production volume is mediated by yield — a relationship that varies enormously across cultivars, regions, viticultural practices, and vintages, meaning that acreage rankings and production volume rankings do not necessarily correspond (high-yielding cultivars in warm irrigated regions may produce more wine per hectare than low-yielding premium cultivars in cool-climate appellations with strict yield restrictions).
Anbaustatistik
Erzeugungsstatistik
Produktionskennzahlen
Produktionsstatistik
Weinbaustatistik
cultivation statistics
output statistics
production data
production metrics
vineyard statistics
OIV (2023) State of the World Vine and Wine Sector Anderson K, Nelgen S (2020) Which Winegrape Varieties are Grown Where? Revised edition, University of Adelaide Press Eurostat (2023) Vineyard statistics Statistik Austria (2023) Weinbauerhebung
Production Statistics
A Riesling vine at Weingut Keller in Rheinhessen fulfills a production system role that defines whether synthetic pesticides, mineral fertilizers, and herbicides may be applied, with the specific production system role (conventional, organic, or biodynamic) determining maximum permitted copper usage ranging from 3-6 kg/ha/year and shaping the vineyard's biodiversity management approach.
A grapevine role that inheres in a vine by virtue of the agricultural management framework under which it is cultivated, determining the permitted range of viticultural inputs, pest management strategies, and soil stewardship practices applied throughout the production cycle.
Production_System_Role classifies grapevines according to the overarching management philosophy and regulatory framework governing their cultivation. This role is independent of the vine's genetic identity or training system and instead captures the agronomic context: a single cultivar such as Chardonnay may simultaneously bear different production system roles across different vineyard blocks within the same estate. Subclasses include conventional, organic, biodynamic, sustainable, regenerative, dry-farmed, and irrigated production roles.
Anbausystem-Rolle
Bewirtschaftungssystem-Rolle
Produktionssystem-Rolle
farming system role
production management role
viticultural system role
OIV (2022) International Code of Oenological Practices EU Regulation 2018/848 on organic production Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp: 239-252
Production System Role
Production trend for Sauvignon Blanc globally documents one of the most dramatic cultivar expansions of the past three decades — from approximately 65,000 ha in 2000 to approximately 125,000 ha by 2020 (roughly +90% over 20 years), driven primarily by the explosive growth of Marlborough, New Zealand (from approximately 2,000 ha in 2000 to approximately 27,000 ha by 2020, representing the single largest regional cultivar expansion in recent viticultural history), supplemented by expansion in Chile, South Africa, and southern France, with the trend driver identified as the global consumer appetite for the distinctive aromatic style of New Zealand Sauvignon Blanc that catalyzed worldwide interest in the variety. Production trend for Grüner Veltliner in Austria documents a trajectory of quality-focused consolidation — peak acreage of approximately 18,800 ha in 1999 declining to approximately 13,500 ha by 2023 (approximately -28% over 24 years), with the contraction concentrated in low-quality sites and bulk-production vineyards while premium sites in the Wachau, Kamptal, and Kremstal maintained or expanded acreage, representing a restructuring trend where volume declined but aggregate quality and average price per bottle increased substantially.
Production trend is a production statistics entity that documents the temporal trajectory of change in vineyard acreage and wine production volume for a specific grape cultivar over multi-year periods, expressed as absolute change (hectares or hectoliters gained or lost over a defined period), percentage change (rate of expansion or contraction relative to a baseline), trend classification (expanding, stable, contracting, or restructuring — where acreage shifts between regions without net global change), and the identified drivers of the trend (market demand shifts, climate adaptation, regulatory changes, disease pressure, consumer preference evolution, subsidy programs), providing the dynamic temporal dimension that contextualizes current production statistics within their developmental trajectory and enables forward projection of cultivar market significance.
Production trend analysis transforms static acreage and volume statistics into dynamic narratives of cultivar market evolution — revealing whether a cultivar's current statistical position represents a stable equilibrium, a moment within an expansion trajectory, or a point on a declining arc, and identifying the forces driving the observed trajectory. The analytical framework for production trends distinguishes several characteristic trajectory types — monotonic expansion (sustained acreage growth driven by strong market demand, as with Sauvignon Blanc 2000-2020), monotonic contraction (sustained acreage decline driven by market displacement or policy intervention, as with Carignan under EU vine-pull schemes), quality-focused restructuring (aggregate acreage decline combined with increasing concentration in premium sites and rising average quality/price, as with Grüner Veltliner in Austria or Riesling in Germany), geographic redistribution (stable or declining acreage in traditional regions combined with expansion in new regions, as with Malbec's France-to-Argentina shift), and cyclical fluctuation (acreage oscillating in response to vintage quality, price cycles, or alternating surplus-shortage dynamics). The driver identification dimension of trend analysis requires integration of market data (price trends, export volumes, consumer research), regulatory context (subsidy programs, planting authorization changes, appellation reforms), climatic data (warming trends, extreme event frequency), and industry structure analysis (consolidation, investment patterns, generational change in wine estates) — a multi-factorial analysis that resists simple causal attribution. The climate change dimension is increasingly dominant in production trend analysis — documented trends include the northward expansion of red cultivar cultivation in Europe (Pinot Noir acreage expanding in southern England, Champagne increasing Pinot Noir proportion), the progressive replacement of early-ripening by later-ripening cultivars in warming Mediterranean regions, and the expansion of heat-tolerant varieties (Grenache, Mourvèdre, Assyrtiko) into regions previously dominated by cool-climate cultivars. The forward projection of production trends — extrapolating current trajectories to estimate future cultivar market positions — is inherently speculative but commercially valuable for vineyard investment decisions with 20-30 year time horizons.
Anbautrend
Flächenentwicklung
Markttrend
Produktionsentwicklung
Produktionstrend
acreage trend
cultivation trend
market trend
planting trend
production trajectory
Anderson K, Nelgen S (2020) Which Winegrape Varieties are Grown Where? Revised edition, University of Adelaide Press OIV (2023) State of the World Vine and Wine Sector New Zealand Winegrowers (2023) Annual Report — vineyard statistics van Leeuwen C et al. (2019) An update on the impact of climate change in viticulture and potential adaptations. Agronomy 9(9): 514
Production Trend
Production volume for Grüner Veltliner in Austria (ÖWM data, 2023 vintage) approximately 850,000 hL, representing roughly 35% of total Austrian wine production by volume (approximately 2.4 million hL total) — the volume reflecting both the substantial acreage (13,500 ha) and the moderate-to-high average yield characteristic of Grüner Veltliner in Austrian conditions (approximately 63 hL/ha average, ranging from 35-40 hL/ha for premium Smaragd to 80+ hL/ha for entry-level Qualitätswein). Production volume for Champagne (all cultivars combined, representing a production volume context where individual cultivar volumes are not separately reported) approximately 2.5 million hL from approximately 34,000 ha in the 2022 vintage, with the three permitted cultivars — Pinot Noir (38% of area), Chardonnay (31%), and Meunier (31%) — contributing to a blended output where varietal disaggregation of volume is meaningful only for the minority of single-cultivar cuvées (Blanc de Blancs from Chardonnay, Blanc de Noirs from Pinot Noir).
Production volume is a production statistics entity that documents the quantity of wine produced from a specific grape cultivar within a defined geographic scope and time period, expressed in hectoliters (hL), liters (L), or metric tons of grapes harvested (with approximate conversion to wine volume at 0.65-0.75 hL wine per metric ton of grapes depending on cultivar and production method), derived from national production statistics, industry surveys, and appellation production reports, and specifying the geographic scope, vintage year, conversion methodology, and relationship between grape harvest volume and wine output, providing the output measure that complements acreage data by quantifying actual productive contribution rather than merely planted extent.
Production volume quantifies the actual productive output of cultivar cultivation — a measure that complements but is not directly proportional to acreage data because volume is modulated by yield per hectare, which varies enormously across cultivars, regions, viticultural practices, and vintage conditions. The yield-volume relationship is central to understanding production volume data — a cultivar planted on 10,000 ha at an average yield of 40 hL/ha (typical for premium quality-focused production) generates 400,000 hL, while the same acreage at 100 hL/ha (typical for high-volume commodity production) generates 1,000,000 hL, meaning that acreage rankings and volume rankings may diverge substantially. The distinction between grape harvest volume (typically reported in metric tons or quintals) and wine production volume (reported in hectoliters) involves a conversion factor that varies by production method — standard still wine production converts approximately 0.70-0.75 hL of wine per metric ton of grapes (press yield of 65-75%), while sweet wine production from concentrated grapes may yield substantially less (0.10-0.30 hL/ton for TBA-level concentration). The vintage-to-vintage variability of production volume reflects both viticultural and meteorological factors — spring frost, hail, drought, disease pressure, and extreme heat events can reduce production volume by 20-50% in affected regions relative to normal years, making multi-year averages more representative of productive capacity than single-vintage figures. The institutional reporting of production volume varies in cultivar specificity — some jurisdictions report cultivar-specific production (Austria, France for many appellations), while others report only aggregate production by appellation or region without cultivar disaggregation, limiting the availability of cultivar-specific volume data for cross-national comparison.
Erntemenge
Erzeugungsmenge
Produktionsausstoß
Produktionsvolumen
Weinproduktionsmenge
harvest volume
output volume
production quantity
production yield
wine output
OIV (2023) State of the World Vine and Wine Sector Österreichische Weinmarketinggesellschaft (2023) Austrian Wine Statistics Eurostat (2023) Wine production statistics CIVC (2023) Les expéditions de vins de Champagne — statistiques
Production Volume
Grenache productive mature grafted scion trunk on SO4 vigorous drought-tolerant rootstock in Languedoc at 14 years post-planting shows 95-115 mm scion diameter supporting stable production 6.8-9.2 kg per vine (88-118 hL/ha) across 5-year period with coefficient of variation 15% (lower than own-rooted equivalent due to rootstock drought tolerance), maintains 125-160 g combined reserves (scion 45-58 g, rootstock 80-102 g), graft union stable at 110-116% swelling, scion sapwood 24-30 mm conducting 55-85 L/day with rootstock-enhanced drought resistance, and demonstrates consistent productivity despite challenging water-limited site where own-rooted vines would struggle
A productive mature grafted scion trunk is a mature grafted scion trunk emphasizing sustained commercial fruit production capacity (typically years 5-25 post-planting) with rootstock-mediated stable yields, consistent fruit quality, and efficient resource use, representing the extended productive phase that justifies grafted vineyard investment with rootstock benefits supporting long-term productivity. Productive mature grafted scion trunk characteristics include= adequate scion diameter maintaining storage capacity (varying by rootstock= 65-130 mm for vigorous rootstocks, 55-110 mm moderate, 50-100 mm low-vigor), functional scion sapwood providing rootstock-influenced water transport (15-30 mm radial depth conducting 30-100 L/day depending on rootstock vigor and canopy), stable graft union (diameter ratio 108-120% maintained supporting reliable scion-rootstock integration), balanced heartwood/sapwood in scion (heartwood providing support without excessive sapwood reduction), and combined scion-rootstock reserves supporting productivity (100-190 g total varying by rootstock vigor). Productive mature grafted scion trunk annual cycle with rootstock influence includes= spring mobilization utilizing combined reserves (40-60% of scion and rootstock stored starch supporting budbreak potentially advanced or delayed 5-10 days by rootstock phenology effects), flowering and fruit set with rootstock vigor expression (vigorous rootstocks promoting larger inflorescences and higher fruit set percentages), berry development supported by rootstock water and nutrient supply (Phase I and II growth rates modulated by rootstock hydraulic capacity and nutrient efficiency), véraison and ripening with rootstock quality effects (some rootstocks enhancing anthocyanin accumulation, phenolic development, or sugar accumulation rates), and post-harvest reserve accumulation in scion and rootstock (restoring 80-100% of depleted reserves with rootstock influence on photosynthate partitioning). Productive mature grafted scion trunk yield patterns show= stable annual production varying by rootstock class (vigorous rootstocks 6-10 kg per vine with <25% variation, moderate 4.5-7.5 kg with <20% variation, low-vigor 3-6 kg with <20% variation year-to-year), rootstock-influenced cluster characteristics (vigorous rootstocks promoting larger clusters 180-250 g, low-vigor smaller clusters 90-150 g), consistent bud fruitfulness modulated by rootstock (vigorous rootstocks 75-95% fruitful buds, moderate 70-88%, low-vigor 65-85%), and appropriate yield/quality balance for rootstock type (vigorous rootstocks requiring higher yields maintaining quality, low-vigor rootstocks producing lower yields with potentially enhanced quality). Productive mature grafted scion trunk efficiency with rootstock benefits includes= optimal leaf area/fruit ratio for rootstock vigor (vigorous rootstocks 10-18 m² per kg, moderate 8-14 m², low-vigor 6-12 m²), efficient water use mediated by rootstock (drought-tolerant rootstocks producing 0.4-1.0 kg fruit per liter transpired, water-demanding rootstocks 0.3-0.7 kg/L), rootstock-enhanced nitrogen economy (efficient rootstocks supporting productivity on low-N soils without excessive vigor), and appropriate carbon partitioning influenced by rootstock (vigor affecting distribution between fruit, vegetative growth, and reserves). Productive mature grafted scion trunk management for sustained productivity includes= annual pruning accounting for rootstock vigor (vigorous rootstocks 30-50 buds, moderate 20-35 buds, low-vigor 15-25 buds per vine), crop load adjustment for rootstock type (vigorous combinations may require cluster thinning to 25-45 clusters, low-vigor combinations limited to 15-30 clusters), disease management protecting union and trunk (graft union remains potential entry point for trunk pathogens throughout productive life), rootstock-appropriate nutrition (accounting for rootstock nutrient efficiency and pH tolerance), irrigation management utilizing rootstock characteristics (drought-tolerant rootstocks requiring less supplemental water, shallow-rooted rootstocks needing more frequent irrigation), and continued rootstock sucker removal (preventing rootstock canopy development bypassing scion benefits). Productive mature grafted scion trunk economic significance includes= peak return period for grafted investment (established vines with rootstock benefits providing optimal yield-quality-cost balance), stable income from rootstock-supported consistent production (rootstock stress tolerance reducing vintage variation), optimal cost efficiency with rootstock benefits (phylloxera resistance, soil adaptation, vigor control reducing remedial costs), and maximum asset value during rootstock-supported productive maturity (vineyard value highest when grafted vines expressing full rootstock benefits). Productive mature grafted scion trunk duration varies= well-matched rootstock-scion-soil combinations supporting 15-30 years productive maturity, poorly matched combinations showing briefer productivity 8-18 years, and exceptional combinations with intensive management potentially achieving 20-40 years sustained production before significant senescence.
Productive_Mature_Grafted_Scion_Trunk emphasizes sustained commercial production with rootstock and is distinguished from Well_Established_Mature_Grafted_Scion_Trunk (peak developmental status) and Mature_Grafted_Scion_Trunk (broader phase) by focus on economic productivity and rootstock-supported yield stability. Critical for grafted vineyard business planning, understanding rootstock contributions to sustained productivity, optimal rootstock selection for long-term economic performance, and decision-making on grafted vineyard renovation timing.
Commercial grafted production trunk
Ertragsstabile Veredelung
Kommerziell produktive Aufpropfung
Produktiver reifer gepfropfter Stamm
Sustained productivity grafted scion
Productive Mature Grafted Scion Trunk
Grenache productive mature own-rooted trunk in Languedoc at 14 years post-planting shows 85-105 mm diameter supporting stable production of 5.2-7.8 kg per vine (68-95 hL/ha at typical density) across 5-year period with coefficient of variation 12%, maintains 95-135 g starch reserves, 25-32 mm functional sapwood conducting 48-72 L/day, and continues healthy annual ring formation (2.2-3.8 mm recent 3-year average) indicating sustained productive capacity
A productive mature own-rooted trunk is a mature own-rooted trunk emphasizing sustained commercial fruit production capacity (typically years 5-25 post-planting) with stable annual yields, consistent fruit quality, and efficient resource use, representing the extended productive phase that justifies commercial vineyard investment and maintenance. Productive mature own-rooted trunk characteristics supporting sustained productivity include= adequate trunk diameter maintaining carbohydrate storage capacity (60-120 mm for typical wine grape cultivars), functional sapwood providing consistent water transport (15-30 mm radial sapwood depth conducting 25-80 L/day depending on canopy), balanced heartwood/sapwood ratio (heartwood providing structural support without excessive sapwood reduction), annual growth rings indicating stable health (1.5-4 mm ring width without extreme narrowing suggesting stress or senescence), and bark integrity protecting against environmental stress and pathogens. Productive mature own-rooted trunk annual productivity cycle includes= spring mobilization of stored reserves (utilizing 40-60% of stored starch for budbreak and early shoot growth), flowering and fruit set supported by current and stored photosynthate (May-June in Northern Hemisphere), berry development during Phase I and II with combined stored and current assimilate (June-August), véraison and ripening primarily supported by current photosynthesis (August-September), and post-harvest reserve accumulation (September-November restoring 80-100% of depleted starch reserves). Productive mature own-rooted trunk yield patterns show= stable annual production with low coefficient of variation (typically 3.5-6.5 kg per vine with <20% variation year-to-year in absence of extreme weather or deliberate crop adjustment), consistent cluster number per shoot (typically 1-2 clusters per fruitful shoot with 70-90% bud fruitfulness), uniform berry development (minimal variation in berry size or composition within vintages), and appropriate yield/quality balance (production levels supporting target fruit composition and wine quality). Productive mature own-rooted trunk efficiency characteristics include= optimal leaf area/fruit weight ratio (typically 8-15 m² leaf area per kg fruit for quality wine production), efficient water use (producing 0.3-0.8 kg fruit per liter transpired water depending on climate and irrigation), balanced nitrogen economy (adequate vine nitrogen status supporting productivity without excessive vigor), and appropriate carbon partitioning (50-70% of annual photosynthate allocated to fruit during ripening, 15-25% to vegetative growth, 10-20% to storage reserves). Productive mature own-rooted trunk management for sustained productivity includes= annual pruning maintaining appropriate bud number and crop potential (typically 20-40 buds per vine depending on training system and vigor), crop load adjustment if needed (cluster thinning to 20-40 clusters per vine in high-fertility sites or vigorous cultivars), disease management preventing productivity decline (protecting against trunk diseases, canopy diseases, soil-borne pathogens), appropriate nutrition maintaining productivity (N, P, K, and micronutrient management), and irrigation strategies balancing productivity and quality (deficit irrigation during ripening in many quality-focused systems). Productive mature own-rooted trunk economic significance includes= peak return on investment period (established vines with minimal remedial costs producing target yields and quality), stable predictable income (consistent annual production supporting farm planning and winery contracts), optimal cost efficiency (per-unit production costs at minimum with established mature vines), and asset value maximization (vineyard land value highest during productive mature phase). Productive mature own-rooted trunk duration varies by= cultivar longevity (some varieties maintaining productivity 15-25 years, others showing earlier decline), site quality (optimal sites supporting longer productive periods than marginal sites), management intensity (appropriate care extending productive phase, neglect or disease accelerating decline), and environmental conditions (moderate climates supporting longer productivity than extreme climates).
Productive_Mature_Own_Rooted_Trunk emphasizes sustained commercial production and is distinguished from Fully_Developed_Mature_Own_Rooted_Trunk (peak developmental status) and Mature_Own_Rooted_Trunk (broader productive phase) by focus on economic productivity and yield stability. Critical for vineyard business planning, understanding productive lifespan economics, optimal management for sustained production, and decision-making regarding vineyard renovation or replanting timing.
Commercial production trunk
Ertragsstabiler Eigenwurzelstamm
Kommerziell produktiver Stamm
Produktiver reifer Stamm
Sustained productivity trunk
Productive Mature Own Rooted Trunk
Progeny from a Riesling × Solaris cross bear Progeny_Selection_Roles as members of a population evaluated for segregation of resistance loci, quality trait distribution, and individual performance—with both individual genotypes and the cross combination assessed for breeding program decisions. An F1 population segregating for Run1 resistance serves in Progeny_Selection_Roles, with the 1:1 resistant:susceptible ratio confirming expected inheritance while individual resistant progeny are evaluated for quality traits determining advancement. Progeny from multiple crosses involving a promising resistance donor bear Progeny_Selection_Roles in combining ability analysis, with the donor's GCA estimated from average progeny performance across different V. vinifera seed parents.
A selection role that inheres in a grapevine genotype by virtue of its evaluation as offspring of a specific cross, assessed in the context of the breeding population generated from that parental combination. Progeny selection emphasizes the population context—individuals are evaluated relative to their siblings from the same cross, enabling assessment of both individual merit and cross combination value. High-performing progeny within a cross may be advanced as variety candidates, while the overall progeny distribution informs decisions about whether to repeat, modify, or abandon specific parental combinations for future crossing. Progeny selection analysis examines segregation ratios (for traits with simple inheritance), population means and variances (for quantitative traits), and correlations among traits, providing genetic insights alongside individual selection decisions. The progeny perspective enables estimation of combining ability—general combining ability (GCA) reflecting a parent's average contribution across crosses, and specific combining ability (SCA) reflecting the performance of particular parental combinations beyond GCA predictions.
Progeny_Selection_Role captures the population-level perspective essential for breeding program management, where individual selection decisions are embedded in cross combination evaluation. The progeny framework connects individual merit assessment with genetic parameter estimation—heritability, genetic correlations, combining ability—that informs future crossing decisions. High-performing individuals from poor crosses may still be advanced, but the cross combination is unlikely to be repeated conversely, crosses producing high average performance may be expanded even if no exceptional individuals emerge. Progeny selection in grapevines is complicated by long generation times (years between cross and progeny evaluation) and the difficulty of producing large progeny populations from individual crosses, limiting statistical power for genetic analysis. Contemporary genomic approaches enable more powerful progeny analysis by integrating marker information with phenotypic data.
Familienauswahl
Kreuzungsauswertung
Nachkommenschaftsprüfung
Nachkommenschaftsselektion
cross evaluation
offspring evaluation
progeny selection
Progeny Selection Role
In Syrah berries in Côte-Rôtie, Rhône Valley during BBCH 83-89, progressive anthocyanin synthesis maintains accumulation rates of 45 mg/kg per day during the first 18 days post-veraison, declining to 28 mg/kg per day during days 18-30, and further decreasing to 12 mg/kg per day during the final 12 days, generating total accumulation from 0.18 to 1.92 mg/g fresh weight over 42 days with UFGT enzyme activity peaking at 85 nmol/min/g fresh weight at day 15 under moderate temperatures averaging 25°C.
Progressive anthocyanin synthesis describes the sustained enzymatic production of anthocyanin pigments in Vitis vinifera berry skin cells throughout the post-veraison ripening period, wherein continuous flavonoid pathway gene expression maintains biosynthetic enzyme activities enabling ongoing conversion of phenylpropanoid precursors through the anthocyanin branch pathway to generate the daily increment of 20-80 mg/kg anthocyanin accumulation that establishes final berry pigment content at harvest maturity. This process involves the coordinate expression and activity of multiple pathway enzymes including phenylalanine ammonia lyase initiating phenylpropanoid flux, chalcone synthase and chalcone isomerase generating the flavanone naringenin, flavanone 3-hydroxylase and flavonoid 3-prime-hydroxylase producing dihydroflavonols, dihydroflavonol reductase reducing these to leucoanthocyanidins, and anthocyanidin synthase catalyzing the final oxidation step generating colored anthocyanidins that are subsequently glucosylated and methylated to form stable vacuolar pigments. The temporal dynamics of progressive anthocyanin synthesis show characteristic phases with initial rapid synthesis during the first 14-21 days post-veraison when enzyme activities and substrate availability peak, sustained intermediate synthesis during mid-ripening maintaining accumulation rates of 25-50 mg/kg per day, and declining synthesis during late ripening as regulatory programs shift and potential degradation begins to offset continued production in some conditions. The transcriptional regulation of progressive anthocyanin synthesis involves MYB transcription factors including VvMYBA1 and VvMYBA2 that activate structural gene expression, with these regulatory genes themselves induced by developmental signals at veraison and modulated by environmental factors including light and temperature throughout the subsequent ripening period. Substrate availability influences progressive anthocyanin synthesis rates through competition with other branches of phenylpropanoid metabolism, with adequate carbon flux through the shikimate and phenylpropanoid pathways required to sustain anthocyanin production while simultaneously supporting proanthocyanidin synthesis and lignification occurring in parallel during berry ripening. Temperature modulation of progressive synthesis involves both direct effects on enzyme catalytic rates and indirect effects on regulatory gene expression, with optimal synthesis at moderate temperatures of 20-26°C daytime while high temperatures exceeding 30-35°C reduce synthesis through transcriptional repression of pathway genes and potential enzyme denaturation. The relationship between progressive synthesis rates and final anthocyanin content integrates biosynthetic capacity with potential degradation losses, with favorable conditions enabling continuous net accumulation while stress conditions may create periods of net loss that reduce final pigment concentrations below the potential established by maximum synthetic capacity.
Progressive_Anthocyanin_Synthesis is subclass of Anthocyanin_Accumulation representing the sustained biosynthetic process. Distinguishes from Color_Intensity_Development which emphasizes visual outcome. Sibling to Color_Intensity_Development and Anthocyanin_Profile_Evolution. Synthesis rates of 20-80 mg/kg per day during active phase. MYB transcription factor regulated. Temperature sensitive with optimal at 20-26°C.
anhaltende Pigment-Synthese
continuous anthocyanin biosynthesis
fortlaufende Anthocyan-Produktion
kontinuierliche Anthocyan-Biosynthese
ongoing anthocyanin production
progressive Anthocyan-Synthese
progressive pigment formation
sustained pigment synthesis
Boss PK et al (1996) Expression of anthocyanin biosynthesis pathway genes. Plant Molecular Biology 32: 565-569 Kobayashi S et al (2002) Retrotransposon-induced mutations in grape skin color. Science 304: 982 Castellarin SD et al (2007) Transcriptional regulation of anthocyanin biosynthesis. Plant Physiology 145: 1451-1469
Progressive Anthocyanin Synthesis
Pinot Noir in Oregon shows progressive cap fall during mid-late June Year N+1 at BBCH 67-68 over 2-3 days June 12-14, with final 22-28% flowers predominantly in basal positions completing cap detachment, declining daily rate from peak 18-24% to progressive 6-10%, visible berry development 10-16 mm diameter from earlier fruit set, under cool climate conditions of 19-24°C temperature, 60-75% relative humidity, typical of this early-to-mid season cultivar completing late shedding contributing 18-24% final berry number
Progressive cap fall represents the declining calyptra detachment phase following peak intensity when remaining 20-30% flowers complete abscission through sequential shedding as final buds mature, marking transition from peak reproductive activity toward flowering completion with progressively decreasing daily cap fall rate. This concluding shedding phase occurs during late flowering in mid-late June in temperate regions Year N+1 following mass cap fall, typically spanning 2-3 days as final flowers predominantly in basal cluster positions complete detachment creating extended tail of flowering activity, corresponding to BBCH 67-68 stages parallel to late anthesis. Shedding intensity decline shows characteristic pattern with decreasing daily detachment rate dropping from 15-25% during peak to 5-10% during progressive phase, cumulative cap fall reaching 70-90% flowers by progressive phase onset, and remaining 10-30% flowers completing detachment during final shedding period. Spatial concentration during progressive fall shows activity predominantly in basal cluster regions reflecting latest flower maturation from previous season primordium formation, with apical and middle regions showing 85-100% completed cap fall, while basal regions showing 50-80% completion with active ongoing shedding. Sequential detachment pattern shows progressive individual flower cap fall rather than mass synchronized activity, with flowers completing abscission as they individually reach maturity creating extended shedding period, and overall pattern reflecting developmental gradient established during previous season inflorescence development. Cap accumulation during progressive phase continues though at reduced rate with fresh detached caps adding to previously accumulated layers, daily additions of 5-10% total flowers creating modest ongoing accumulation, and overall cap debris reaching maximum density by progressive phase conclusion. Flower exposure completion during progressive phase shows final reproductive organs revealing as late-maturing flowers shed caps, with 70-90% cluster showing exposed organs or developing berries from successful earlier pollinations, and remaining 10-30% flowers achieving open state during progressive detachment. Reproductive capacity during progressive phase shows substantially reduced potential with limited fresh pollen from late-opening flowers, expired pollen from earlier anthesis providing minimal viable contribution, declining stigma receptivity in recently opened flowers, and overall reproductive opportunity greatly diminished compared to peak period. Berry development visibility increases during progressive phase as successfully fertilized flowers expand creating recognizable fruit structures 8-15 mm diameter from earliest fruit set, visible berry clusters contrasting with remaining flowers showing late opening or senescence, and overall cluster appearance transitioning from predominantly reproductive to predominantly developmental state. Unfertilized flower senescence accelerates during progressive phase with earliest unsuccessful flowers showing advanced browning and beginning abscission, progressive senescence wave following developmental gradient, and accumulating senescing flower debris adding to cap fall debris creating mixed fallen material. Environmental effects during progressive phase show continued sensitivity though reduced compared to peak with temperature affecting late flower cap shedding efficiency, humidity influencing detachment success in final flowers, and overall conditions less critical since majority fruit set 75-85% already established during earlier phases. Temperature during progressive phase typically 20-28°C supporting continued though declining cap fall activity, with warm conditions promoting efficient late shedding completing phase rapidly, while cool conditions may slow late detachment extending progressive period. Humidity during progressive phase shows continued importance for efficient late cap fall with moderate 50-70% facilitating normal detachment in final flowers, while elevated humidity potentially causing retention in susceptible late flowers affecting complete flowering achievement. Weather impacts during progressive phase have limited overall effect on fruit set since majority berry number already established, though adverse conditions affecting late shedding may influence final completion and small contribution from late-opening flowers. Developmental heterogeneity reaches maximum during progressive phase with wide range of states coexisting including advanced developing berries 10-18 mm from earliest fruit set, recently pollinated flowers beginning berry expansion from mid-flowering success, late-opening flowers with limited reproductive potential, and senescing unfertilized flowers from earlier unsuccessful pollinations. Extended cap shedding during progressive phase creates prolonged flowering period in some situations with sequential individual flower maturation causing gradual extended detachment rather than compressed mass fall, potentially reflecting developmental heterogeneity from previous season or current season environmental variation, and affecting flowering duration and fruit set uniformity. Observational characteristics during progressive phase include declining cap fall activity distinguishable from earlier intensive mass fall, spatial concentration in basal positions indicating late flower maturation, visible berry development throughout cluster creating transitional appearance, and overall indication of approaching flowering completion. Phenological progression during progressive phase shows advancement from peak BBCH 63-65 toward completion BBCH 69, with developmental rate affected by temperature and late flower maturation timing, and clear trajectory toward complete anthesis and cap fall conclusion. Management implications during progressive phase include recognizing approaching flowering completion enabling preparation for post-bloom operations, beginning to assess fruit set outcomes through berry visibility and senescing flower observation, and transitioning focus from bloom-period restrictions toward developmental management priorities. Fruit set contribution from progressive phase typically represents 15-25% final berry number established during late shedding, with late-set berries often showing slightly delayed development compared to earlier fruit potentially affecting ripening uniformity. The process shows two distinguishable aspects progressing from sequential cap detachment reflecting individual flower maturation, through extended cap shedding creating prolonged final flowering period. Duration typically spans 2-3 days representing declining phase before final completion, with warm conditions promoting rapid progression while cool conditions may extend period. Cultivar variation in progressive fall shows differences in proportion of late-opening flowers with some showing compressed 10-15% late phase while others showing extended 25-35% late phase affecting overall duration, late flower cap shedding efficiency with some maintaining good detachment while others showing retention problems, and transition timing toward complete cap fall. Regional timing shows progressive cap fall during mid-late June in warm climates, late June in temperate regions, and late June-early July in cool areas, occurring during late flowering 5-8 days after first cap detachment. Viticultural significance includes representing final cap fall contribution completing flowering activity, providing visual indication of approaching completion enabling management transitions, revealing late retention problems indicating potential final fruit set limitations, and marking transition from peak reproductive vulnerability toward more stable developmental phase. Research applications include analyzing factors affecting late flower cap shedding success, evaluating sequential detachment patterns reflecting developmental gradients, quantifying contributions of progressive phase to final fruit set and berry development patterns, assessing extended shedding implications for flowering duration and uniformity, and documenting transition from reproductive to developmental phases through cap fall observation.
Progressive_Cap_Fall is third major Cap_Fall phase following Initial and Mass Cap Fall, distinguished by declining intensity with final flowers completing detachment. Corresponds to BBCH 67-68 parallel to Late_Anthesis. Subdivided into Sequential_Cap_Detachment and Extended_Cap_Shedding. Represents final 15-25% fruit set contribution. Transitions to Final_Cap_Fall as completion approaches. Marks declining phase toward flowering conclusion. Reduced weather sensitivity compared to peak though conditions still affect late shedding success.
abnehmender Kappenfall
declining cap fall
late cap shedding
sequential calyptra detachment
sequentielle Kappenablösung
späte Kalyptralösung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 380-388 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Progressive Cap Fall
In Chenin Blanc vines in the Loire Valley, progressive chilling units accumulate through winter following the Utah model, with approximately 150 units by December 15, 400 units by January 1, 750 units by January 20, and the cultivar requirement of approximately 1000 units satisfied by early February, with this progression tracked through weather station data showing effective chilling temperatures predominating during January when mean temperatures range from 2-6°C.
Progressive chilling units in Vitis vinifera represents the incremental accumulation of effective cold exposure quantified through temperature-based models that track the progress toward satisfaction of the cultivar-specific chilling requirement during winter dormancy. This quantification approach provides a standardized method for comparing chilling status across different seasons, locations, and cultivars, enabling prediction of dormancy release timing and assessment of chilling adequacy in different viticultural environments. The simplest chilling unit models count hours at temperatures below a threshold, typically 7.2°C, with each hour contributing one chilling unit toward the total requirement, while more refined models recognize that different temperatures contribute differently to dormancy release. The Utah model and its derivatives assign weighted values to different temperature ranges, with temperatures between 2.5-9.4°C receiving full chilling credit, temperatures from 0-2.5°C and 9.4-12.4°C receiving partial credit, temperatures below 0°C receiving no credit, and temperatures above 15.9°C potentially negating previously accumulated chilling. The dynamic model, developed originally for Mediterranean climates, treats chilling as a two-step process involving initial cold exposure that creates an intermediate compound, followed by consolidation at moderate temperatures that converts the intermediate to a stable chilling unit, better accounting for the fluctuating temperatures common in mild winter regions. The progressive nature of chilling unit accumulation reflects the gradual physiological changes occurring during dormancy release, with each increment of chilling exposure contributing to the cumulative dismantling of dormancy-maintaining mechanisms until the threshold requirement is achieved. Monitoring progressive chilling units through the winter allows viticulturists to predict budbreak timing, assess risk of inadequate chilling in warm winters, and make decisions regarding dormancy-breaking treatments if natural chilling appears insufficient. The relationship between accumulated chilling units and forcing response provides validation of chilling models, with decreasing days to budbreak correlating with increasing chilling unit accumulation until the requirement is satisfied.
Progressive_Chilling_Units represents the quantitative tracking aspect of Chilling_Accumulation. Distinguished from Cumulative_Cold_Exposure which addresses the physiological cold exposure, while Progressive_Chilling_Units specifically addresses model-based quantification. Various models include simple threshold counting, Utah model with temperature weighting, and dynamic model with two-step processing. Enables prediction of dormancy release timing. Cultivar requirements range from approximately 400-1500 units depending on model used and cultivar genetics.
accumulating chill hours
chilling unit progression
fortschreitende Kältestunden
incremental chilling
inkrementelle Chilling-Akkumulation
progressive Kälteeinheiten
progressive cold units
zunehmende Kälteeinheiten
Progressive Chilling Units
In a Paso Robles Syrah vineyard, progressive color development was monitored over 7 days in late August Year N+1 (BBCH 84-85) when anthocyanin accumulation averaged 75 mg/kg per day increasing from 135 mg/kg (30% berry surface colored) to 660 mg/kg (90% colored), with delphinidin-3-glucoside reaching 18% of total producing characteristic purple color, berry firmness declining to 1.4 N, sugar rising from 13.5 to 21.8° Brix, under warm days (28-31°C) and moderate nights (15-18°C) with eastern canopy cluster exposure.
Progressive_Color_Development represents the sustained phase of anthocyanin accumulation and spatial expansion of berry pigmentation following color break when initially scattered red-purple patches coalesce and intensify transforming berry appearance from variegated green-red to predominantly colored with diminishing green remnants approaching complete pigmentation. This process occurs during mid-to-late veraison (BBCH 83-85), typically 3-10 days duration depending on cultivar and environmental conditions, and is characterized by anthocyanin accumulation rates of 40-120 mg/kg skin per day increasing total skin anthocyanins from color break levels 80-150 mg/kg to near-maximum values 400-800 mg/kg (cultivar-dependent final levels range 500-3000 mg/kg). Spatial progression of pigmentation during this phase shows: initially discrete colored patches (occupying 20-40% berry surface at color break) expand radially recruiting adjacent unpigmented cells into anthocyanin synthesis creating confluent pigmented zones, green remnants progressively shrink retreating toward shaded berry surfaces (often northwestern aspects in northern hemisphere) and pedicel attachment zones that receive minimal light signals, and within-berry pigmentation gradients develop with sun-exposed surfaces showing 2-4 fold higher anthocyanin concentrations than shaded areas creating spatial heterogeneity even on individual berries. Anthocyanin accumulation kinetics during progressive development follow: biosynthetic rates peak at 50-150 mg/kg per day during mid-phase (days 3-6 of progressive development) when enzyme activities reach maximum levels, total skin anthocyanins increase sigmoidally with inflection point typically at 40-60% of final concentration, and absolute anthocyanin content per berry rises 8-15 fold from color break (4-8 mg total per berry) to late progressive phase (50-100 mg per berry) reflecting both increased concentration and slight berry expansion. Biosynthetic machinery during progressive development operates at peak capacity: UFGT enzyme activity reaches maximum levels 25-45 units/kg fresh weight catalyzing glucosylation of 40-80 μmol anthocyanidins per hour per kg, PAL activity peaks at 80-150 units providing phenylpropanoid flux sustaining pathway operation, DFR and LDOX achieve 70-120 units producing leucoanthocyanidin and anthocyanidin substrates, and coordinated expression of all pathway enzymes creates metabolic channeling efficiency maximizing anthocyanin output while minimizing intermediate accumulation. Gene expression during progressive development maintains: anthocyanin biosynthetic genes sustained at 80-100% of peak levels with UFGT transcripts 150-200 fold above green berry baseline, regulatory MYB factors (VvMYBA1/A2) continue high expression 100-150 fold coordinating pathway genes, while genes for competing pathways including proanthocyanidin synthesis (LAR, ANR) decline creating favorable metabolic channeling toward anthocyanins, and sugar transporters show parallel elevation supporting hexose accumulation that provides both substrate and signaling for anthocyanin synthesis. Anthocyanin composition during progressive development shows: malvidin-3-glucoside maintains dominance at 50-70% while absolute levels increase from 40-80 mg/kg at color break to 250-500 mg/kg late progressive phase, acylated anthocyanins (acetyl and coumaroyl derivatives) increase from 10-15% to 15-25% of total enhancing color stability, diglucosides common in non-vinifera species remain absent or trace in Vitis vinifera, and hydroxylation patterns establishing delphinidin, cyanidin, petunidin, and peonidin ratios remain relatively constant reflecting stable expression of F3'H and F3'5'H hydroxylases. Vacuolar sequestration during progressive development involves: anthocyanin concentrations in epidermal cell vacuoles reach 8-20 mM (from 3-8 mM at color break) approaching solubility limits, vacuolar pH maintained at 3.0-3.5 through enhanced V-ATPase activity ensuring flavylium cation stability, anthocyanin vacuolar inclusions (AVIs) expand in cultivars forming these structures creating highly concentrated pigment deposits (>50 mM local concentrations), and co-pigmentation with flavonols and hydroxycinnamic acids continues amplifying color intensity 30-50% beyond that expected from anthocyanin concentration alone. Chlorophyll degradation accelerates during progressive development: chlorophyll a and b decline from color break levels 40-80 μg/g to late progressive phase 10-30 μg/g through pheophytinase and pheophorbide a oxygenase activities, green color masking progressively eliminated allowing anthocyanin pigmentation to dominate visual appearance, carotenoid content remains relatively stable (20-40 μg/g) contributing yellow-orange background visible in white cultivars but overwhelmed by anthocyanins in reds, and complete disappearance of photosynthetic pigments creates characteristic ripe berry translucency in some cultivars. Environmental modulation of progressive development shows: optimal temperature 22-28°C supports maximum progression rates completing color development in 4-6 days for early cultivars, cooler temperatures 16-22°C extend progression to 6-10 days but may enhance final color intensity through improved anthocyanin stability, heat stress >32°C severely impairs progression with anthocyanin accumulation reduced 40-70% and extended development periods 10-14 days often producing inadequately colored fruit, and diurnal temperature variation >12-15°C day-night difference optimizes progression through balanced synthesis and stabilization. Light effects on progressive development inclu UV-B exposure continues enhancing anthocyanin synthesis throughout progression maintaining 30-50% higher accumulation in exposed versus shaded regions, visible light particularly blue wavelengths sustains photoreceptor signaling supporting biosynthetic gene expression, canopy architecture affecting cluster light exposure creates within-vine variation with exterior clusters showing 25-45% higher final anthocyanin levels than interior positions, and leaf removal during progressive development can enhance coloring 20-35% if temperature stress avoided though excessive exposure risks heat inhibition. Water relations influence on progressive development: moderate deficit (-0.9 to -1.3 MPa stem ψ) maintains or enhances progression through sustained ABA signaling and berry concentration producing 15-30% higher final anthocyanin concentrations, severe stress (<-1.5 MPa) arrests progression reducing final color 30-60% through metabolic limitation despite elevated ABA, while excess water dilutes berries and suppresses ABA-dependent gene expression reducing final anthocyanin 20-40% compared to moderate deficit, and timing of deficit imposition affects outcome with pre-veraison stress more beneficial than deficit imposed during progressive development. Cultivar variation in progressive development reflects: rapid developers like Pinot Noir complete progression in 3-5 days reaching 1200-2000 mg/kg final anthocyanins, moderate cultivars like Merlot and Syrah require 5-8 days achieving 800-1500 mg/kg, while slow developers like Cabernet Sauvignon and Nebbiolo need 8-12 days reaching 600-1200 mg/kg demonstrating genetic differences in maximum synthesis rates and biosynthetic duration. Coordination with other ripening processes during progressive development: berry softening accelerates concurrently with firmness declining 40-60% from green berry values, sugar accumulation peaks at 1.5-3.5% Brix per week during progressive color development, acid decline intensifies particularly malate respiratory consumption reducing titratable acidity 15-30%, and seed maturation progresses with testa lignification and browning paralleling skin color development. Metabolic costs of progressive development are substantial: anthocyanin biosynthesis consumes 10-20% of berry carbon import during peak color development, nitrogen demands for enzyme synthesis and anthocyanin precursors draw on berry reserves requiring 30-60 μmol phenylalanine per kg fresh weight, energy requirements including ATP for biosynthesis and vacuolar transport represent 12-20% of berry respiration during maximum accumulation rates, and competition for carbon between anthocyanin synthesis, sugar accumulation, and other metabolic processes may create trade-offs affecting final berry composition.
Progressive_Color_Development is distinguished from Color_Break_Initiation by representing sustained accumulation rather than initial pattern formation, from First_Color_Change by showing accelerating rather than nascent pigmentation, and from Complete_Color_Change by exhibiting ongoing progression rather than final state achievement. Duration: fast cultivars 3-5 days, moderate 5-8 days, slow 8-12 days. Accumulation rates peak 50-150 mg/kg/day. Temperature optimum 22-28°C. Moderate water deficit enhances. Progressive_Color_Development represents the major phase of anthocyanin synthesis determining final berry color intensity critical for wine quality.
anhaltende Farb-Intensivierung
continuing anthocyanin accumulation
expandierende Pigmentierung
expanding pigmentation
fortgesetzte Anthocyan-Akkumulation
progressive Beeren-Färbung
progressive berry coloring
sustained color intensification
Keller M (2020) The Science of Grapevines 3rd Ed pages 233-236 anthocyanin accumulation kinetics Castellarin SD, Matthews MA, Di Gaspero G, Gambetta GA (2007) Water deficits accelerate ripening and induce changes in gene expression regulating flavonoid biosynthesis in grape berries Planta 227:101-112 environmental effects on color progression Matus JT, Loyola R, Vega A, Peña-Neira A, Bordeu E, Arce-Johnson P, Alcalde JA (2009) Post-veraison sunlight exposure induces MYB-mediated transcriptional regulation of anthocyanin and flavonol synthesis in berry skins Plant Physiol 149:1647-1661 light regulation of progressive development.
Progressive Color Development
In Viognier berries in Condrieu, Rhône Valley during BBCH 83-89, progressive flavor build-up increases total monoterpenes at consistent rates of 0.08 mg/L per day over 38 days, with free linalool progressing from 0.15 to 0.85 mg/L, free geraniol from 0.08 to 0.42 mg/L, and total bound terpenoids from 1.2 to 4.8 mg/L, demonstrating sustained accumulation without plateau under warm days of 30°C and moderate nights of 18°C that maintain biosynthetic enzyme activities throughout ripening.
Progressive flavor build-up describes the continuous incremental increase in flavor compound concentrations within Vitis vinifera berries throughout the ripening period, wherein daily net accumulation of 0.05-0.3 mg/L total flavor volatiles produces the monotonic upward trajectory in aromatic content that characterizes normal flavor development from veraison through harvest over 35-50 days of post-veraison maturation. This process reflects the sustained operation of flavor biosynthetic pathways at rates exceeding compound loss through volatilization and degradation, maintaining positive accumulation balance throughout ripening rather than reaching plateau phases or exhibiting decline that would indicate biosynthetic limitation or accelerated catabolism. The temporal characteristics of progressive flavor build-up show variety-specific accumulation curves, with aromatic varieties typically demonstrating linear or slightly accelerating accumulation patterns as biosynthetic enzyme activities increase during ripening, while neutral varieties may show more modest accumulation rates that plateau earlier in the ripening season due to lower biosynthetic capacity. Cellular mechanisms supporting progressive build-up involve continued induction of biosynthetic gene expression maintaining enzyme activities, efficient vacuolar transport sequestering synthesized compounds, and membrane integrity preventing excessive volatilization losses, with coordination among these processes ensuring sustained accumulation throughout the extended ripening period. The quantitative aspects of progressive flavor build-up vary substantially among compound classes, with monoterpenes in aromatic varieties showing accumulation of 0.1-0.4 mg/L per day during peak periods, norisoprenoid precursors accumulating at 2-8 µg/L per day, and amino acid-derived flavor precursors increasing at 20-50 mg/L per day reflecting the higher absolute concentrations of this compound class. Environmental conditions influence progressive build-up rates through temperature effects on biosynthetic enzyme activities, with optimal temperatures of 20-28°C supporting maximum accumulation rates while temperature extremes may reduce daily accumulation through either limited synthesis at cool temperatures or accelerated degradation at high temperatures. The consistency of progressive flavor build-up provides predictive value for harvest timing, with accumulation rate measurements during mid-ripening enabling estimation of final flavor compound concentrations at projected harvest dates, supporting quality optimization through informed picking decisions that capture optimal aromatic development.
Progressive_Flavor_Build_Up is subclass of Flavor_Compound_Accumulation representing the continuous nature of accumulation. Distinguishes from plateau or declining patterns. Sibling to Flavor_Intensity_Increase and Flavor_Complexity_Development. Key characteristic is sustained positive accumulation rates of 0.05-0.3 mg/L per day. Linear or accelerating trajectory throughout ripening. Predictive value for harvest timing.
anhaltende Geschmacks-Zunahme
continuous flavor accumulation
incremental flavor development
kontinuierliche Aroma-Akkumulation
ongoing volatile build-up
progressive Aroma-Aufbau
schrittweise Volatil-Entwicklung
sustained aroma increase
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 295-305 Marais J (1983) Terpenes in the aroma of grapes and wines. South African Journal of Enology and Viticulture 4: 49-58 Wilson B et al (1984) Determination of free and bound monoterpenes in Muscat grapes. American Journal of Enology and Viticulture 35: 95-100
Progressive Flavor Build Up
In Chardonnay vineyards of the Mâconnais, Burgundy, progressive leaf fall advances acropetally through the canopy during November, with leaf detachment progressing from node positions 3-5 in the first week to positions 8-12 by mid-November, achieving approximately 60% total defoliation at a rate of 2-3 node positions per week under typical autumn temperatures of 6-10°C daytime maxima.
Progressive leaf fall in Vitis vinifera describes the continuous, directional advancement of leaf detachment through the vine canopy following the initial leaf fall phase, characterized by systematic progression from earlier-senescing basal positions toward later-senescing apical and well-exposed positions over a period of approximately 2-4 weeks. This phase encompasses the majority of canopy defoliation volume, typically representing 50-70% of total leaf area loss, and occurs during November in Northern Hemisphere viticulture as abscission zone maturation sequentially reaches completion at progressively younger leaf positions. The progressive pattern reflects the acropetal gradient of leaf initiation during spring growth, whereby basal leaves established earliest also initiate senescence programs earliest, creating a wave of abscission competence that advances up the shoot axis at a rate determined by senescence signal propagation and environmental conditions. Mechanistically, progressive leaf fall involves the sequential completion of cell separation processes at successive node positions, with each leaf's abscission zone requiring approximately 10-14 days from senescence initiation to achieve separation competence under typical autumn temperatures. The rate of progression varies considerably with temperature regime, accelerating during periods with consistent overnight minima below 5°C and daytime maxima of 8-12°C that favor both senescence advancement and abscission zone enzyme activity. Environmental events modulate the steady progression, with wind events or frost episodes causing synchronized detachment pulses within the general progressive trend, while calm mild conditions allow more gradual individual leaf separation. Cultivar differences significantly influence progression rate, with thin-leaved varieties such as Pinot Noir and Gamay typically completing progressive leaf fall within 2-3 weeks, while thick-leaved cultivars including Cabernet Sauvignon and Carignan may require 4-5 weeks for equivalent progression. The spatial pattern during progressive leaf fall creates characteristic canopy appearance changes, with increasing visibility of shoot architecture as leaf coverage retreats apically, providing viticultural indicators of dormancy preparation advancement. Physiologically, leaves detaching during progressive leaf fall have completed varying degrees of nutrient remobilization depending on their position in the senescence sequence, with earlier-detaching leaves showing more complete nitrogen and phosphorus recovery than later-detaching leaves that experienced abbreviated senescence periods. The progressive nature of leaf fall serves important vine functions, allowing extended periods for carbohydrate translocation to permanent structures and preventing sudden exposure of previously shaded tissues to environmental stress.
Progressive_Leaf_Fall represents the main quantitative phase of Leaf_Detachment, encompassing 50-70% of total leaf area loss between Initial_Leaf_Detachment and Mass_Leaf_Fall. Distinguished by its directional acropetal pattern and steady advancement rate rather than the scattered nature of initial detachment or the rapid synchronous character of mass leaf fall. Duration typically spans 2-4 weeks depending on cultivar and temperature conditions. Characterized by progressive exposure of shoot architecture and systematic canopy transparency increase.
advancing leaf detachment
continuing leaf abscission
fortschreitender Blattfall
gradual defoliation
gradueller Laubfall
sequential leaf fall
sequentieller Blattfall
voranschreitende Entblätterung
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Mullins MG, Bouquet A, Williams LE (1992) Biology of the Grapevine Cambridge University Press Holzapfel BP, Smith JP (2012) Developmental stage and climatic factors impact more on carbohydrate reserve dynamics than leaf area. Australian Journal of Grape and Wine Research 18: 38-45
Progressive Leaf Fall
In Sauvignon Blanc vines in Marlborough, New Zealand during April-May, progressive leaf senescence advances from basal shoot positions where senescence initiates 3 weeks before distal positions, with canopy color transitioning from uniform green through heterogeneous yellow-green patchwork to complete yellow over 5 weeks, individual leaf senescence progressing from interveinal regions to complete lamina involvement over 12-15 days, and nitrogen remobilization continuing throughout the extended senescence period enabling 74% total recovery under cool maritime autumn conditions averaging 12°C.
Progressive leaf senescence describes the gradual advancement of senescence processes through Vitis vinifera leaf tissues and across the vine canopy during the autumn period, wherein the sequential progression of chlorophyll degradation, protein catabolism, and cellular dismantling spreads from initial foci through entire leaves and from basal to distal canopy positions over a period of 4-6 weeks, creating the characteristic temporal and spatial patterns of autumn color change that precede leaf abscission and complete defoliation. This process exhibits characteristic spatial patterns within individual leaves, with senescence typically initiating in interveinal regions where mesophyll cells are most distant from vascular nutrient supply, progressively advancing toward the leaf margins and major veins, and ultimately encompassing the entire lamina while petiole tissues remain functional for continued nutrient export until late senescence stages. The acropetal progression of senescence across the vine canopy reflects developmental age differences among leaves, with basal leaves on mature shoot regions having emerged earliest in spring and thus being chronologically oldest initiating senescence first, followed by progressive involvement of more distal leaves, and lateral shoot leaves often senescing last due to their more recent emergence and continued metabolic activity. The temporal coordination of progressive senescence involves both autonomous leaf-age-dependent signals and systemic signals from other senescing tissues, with ethylene production by early-senescing basal leaves potentially accelerating senescence in adjacent leaves through gaseous diffusion, and resource competition creating sink-source relationships that influence senescence progression patterns. Environmental heterogeneity within vine canopies influences progressive senescence patterns, with shaded interior leaves often senescing earlier than well-exposed exterior leaves due to reduced photosynthetic contribution and lower cytokinin status, while leaves experiencing localized stress from disease, pest damage, or nutritional deficiency may initiate senescence independently of the whole-canopy progression. The functional significance of progressive rather than synchronous senescence relates to the extended time window available for nutrient remobilization, with gradual canopy senescence maintaining phloem export capacity from still-functional leaves while resources are being salvaged from senescing tissues, enabling more complete nutrient recovery than would occur with rapid synchronous senescence that might overwhelm transport capacity. The completion of progressive leaf senescence culminates in complete canopy yellowing followed by abscission zone activation and leaf fall, with the 4-6 week duration of visible senescence progression representing the final phase of the annual leaf life cycle before winter dormancy.
Progressive_Leaf_Senescence is subclass of Leaf_Senescence representing the gradual spatial and temporal advancement of senescence. Distinguishes from initiation by emphasizing progression patterns. Sibling to Leaf_Senescence_Initiation, Chlorophyll_Degradation, and Nutrient_Remobilization. Acropetal canopy progression. Within-leaf interveinal initiation. Duration 4-6 weeks. Enables extended nutrient recovery window.
Seneszenz-Progression
advancing leaf senescence
fortschreitende Blattseneszenz
gradual leaf senescence
progressive Blattseneszenz
senescence progression
sequential canopy senescence
sequentielle Kronendach-Seneszenz
Lim PO et al (2007) Leaf senescence. Annual Review of Plant Biology 58: 115-136 Guo Y and Gan S (2005) Leaf senescence: signals, execution and regulation. Current Topics in Developmental Biology 71: 83-112 Keller M (2020) The Science of Grapevines 3rd Ed pp 88-92
Progressive Leaf Senescence
In Tempranillo berries in Ribera del Duero, Spain during BBCH 85-89, progressive malate depletion reduces vacuolar malate concentration from 340 mM at veraison to 65 mM at harvest over 42 days, representing 81% pool depletion with early-phase rates of 12 mM per day during the first 20 days declining to 4 mM per day during the final 22 days, corresponding to whole-berry malic acid reduction from 10.8 to 2.1 g/L under mean ripening temperatures of 25°C.
Progressive malate depletion describes the cumulative reduction of L-malic acid pools within Vitis vinifera berry vacuoles and cytoplasm during ripening, wherein the substantial malate reserves of 200-400 mM vacuolar concentration at veraison are systematically depleted to final concentrations of 30-100 mM at harvest through sustained efflux and catabolism over the 35-50 day post-veraison maturation period. This process reflects the net outcome of continuous malate export from vacuolar storage coupled with ongoing enzymatic degradation and respiratory consumption, with depletion kinetics following characteristic patterns determined by cultivar genetics, temperature regime, and initial malate pool size, typically exhibiting rapid early-phase depletion at rates of 8-15 mM per day transitioning to slower late-phase depletion at 2-5 mM per day as substrate becomes limiting. The vacuolar compartment serves as the primary malate reservoir, containing greater than 90% of total berry malate in dissociated form stabilized by the acidic vacuolar pH of 2.5-3.5, with efflux mediated by ALMT aluminum-activated malate transporter channels and voltage-dependent anion channels responding to cytosolic malate concentration gradients and membrane potential changes during ripening. The progression of depletion shows strong cultivar specificity, with high-acid varieties maintaining larger residual vacuolar pools through lower transport rates and reduced enzyme activities, while rapid-decline varieties achieve near-complete vacuolar depletion with final malate concentrations below 50 mM. Temperature integration determines depletion trajectory, with accumulated growing degree days correlating with fractional malate remaining, enabling prediction of harvest malate levels from thermal time accumulation and variety-specific depletion coefficients. The metabolic consequences of progressive depletion include shifting cellular carbon balance as malate contribution to respiratory substrate diminishes, altered cytosolic redox status as NADPH generation from malic enzyme declines, and modified vacuolar pH as buffer capacity decreases with malate removal. Berry-to-berry variation in depletion rates within clusters reflects differential temperature exposure and vascular connectivity, producing the heterogeneity in final acid composition that contributes to sample variation in harvest monitoring. The irreversible nature of depletion under normal ripening conditions, absent significant malate biosynthesis during post-veraison stages, establishes the monotonic decline characteristic that enables reliable prediction of harvest acidity from early-season measurements.
Progressive_Malate_Depletion is subclass of Malic_Acid_Degradation representing the outcome measure of vacuolar pool reduction. Distinguishes from Malic_Enzyme_Activity and Malate_Respiratory_Consumption which are mechanistic pathways. Sibling class relationship with other Malic_Acid_Degradation subclasses. Duration 35-50 days. Key parameter is vacuolar concentration decline from 200-400 mM to 30-100 mM. Cultivar-diagnostic for acid retention potential.
Malat-Pool-Abnahme
berry malate exhaustion
cumulative malate loss
kumulative Malat-Verlust
malate pool decline
progressive Malat-Erschöpfung
vacuolar malate reduction
vakuoläre Malat-Reduktion
Sweetman C et al (2009) Regulation of malate metabolism in grape berry. Molecular Plant 2: 1270-1280 Terrier N et al (2001) Changes in acidity and proton transport at the tonoplast of grape berries during development. Planta 213: 20-28 Keller M (2020) The Science of Grapevines 3rd Ed pp 268-275
Progressive Malate Depletion
Merlot in Bordeaux shows progressive shoot lignification from mid-July through early September at BBCH 35-38 over 7-8 weeks, with lignification front advancing at 25-35 mm per week through middle shoot internodes 6-12 achieving lignin content of 18-24% dry weight and starch accumulation of 20-26% dry weight under oceanic climate conditions of 20-26°C temperature range, moderate summer precipitation, and declining nitrogen availability typical of this moderately vigorous mid-season cultivar showing steady maturation progression coordinated with fruit development
Progressive shoot lignification represents the continuing maturation process where lignification advances acropetally from basal regions through middle shoot zones following completion of internode elongation, establishing systematic spatial-temporal progression of woody tissue formation coordinated with shoot growth deceleration and fruit development. This essential developmental phase occurs during mid-late summer corresponding approximately to fruit set through mid-berry development at BBCH 35-38, typically spanning 6-8 weeks as maturation front progresses from basal completion through middle shoot regions covering internodes 6-12 depending on total shoot length. Progressive lignification proceeds through systematic acropetal advancement with each successive internode initiating lignification 3-7 days after previous internode reflecting both temporal sequence of internode formation and developmental program coordinating maturation with resource availability and environmental conditions. The lignification front progresses at rate typically 20-40 mm per week corresponding to approximately one internode per week during mid-summer conditions, with rate varying based on temperature, water status, shoot growth rate, and fruit development stage affecting resource partitioning. Secondary wall deposition and lignin incorporation in progressively lignifying internodes follows same cellular mechanisms as basal lignification with xylem vessel and fiber lignification proceeding first followed by parenchyma lignification, creating progressive rigidification moving up shoot from brown woody base through transitional zones to green herbaceous apex. Lignin content in progressively lignifying internodes increases from negligible to 15-25% dry weight over 2-4 weeks depending on internode position and environmental conditions, with middle internodes often achieving highest lignin concentrations reflecting favorable development conditions during their formation and maturation. Mechanical properties develop progressively along shoot creating gradient from fully rigid basal regions through progressively more flexible middle zones to herbaceous apex, with tensile strength increasing from 10-20 MPa in recently lignified regions to 40-80 MPa in fully mature woody tissue. Vascular tissue maturation continues in progressively lignifying regions with secondary xylem development from vascular cambium activity increasing vessel number and diameter, enhancing water transport capacity supporting continued shoot function and fruit development, and secondary phloem formation improving assimilate translocation efficiency. Periderm development progresses acropetally following lignification with cork cambium formation and cork production creating protective bark layer replacing epidermis, with periderm extent corresponding closely to lignification extent creating visible demarcation between mature brown woody base and immature green herbaceous apex. Reserve accumulation intensifies in progressively lignifying regions with starch synthesis converting photosynthate to storage form, increasing starch concentration from 5-10% during early lignification to 18-25% dry weight as lignification completes and growth cessation redirects carbon allocation, with middle internodes often accumulating highest starch concentrations reflecting optimal timing of maturation and resource availability. Metabolic transition during progressive lignification shows declining respiration in successively maturing internodes from growth levels to maturation levels 3-5 fold above dormancy then toward maintenance levels 2-3 fold above dormancy, creating metabolic gradient along shoot corresponding to developmental gradient. Carbon allocation shifts progressively during this phase from substantial vegetative growth consumption early in period to declining vegetative demand as growth slows, increasing fruit demand as berry development intensifies, and increasing storage allocation as maturation proceeds and growth cessation approaches. Shoot water status often shows moderate stress development during progressive lignification period as summer conditions create water limitation, with moderate stress potentially beneficial for promoting maturation and fruit quality development while severe stress risks incomplete lignification and inadequate reserve accumulation. Nitrogen dynamics show continuing decline in tissue concentrations as structural carbohydrates accumulate, with nitrogen remobilization from senescing lower leaves potentially supporting fruit development and storage protein synthesis in maturing woody tissues. Hormonal regulation involves progressively declining auxin as apical production decreases with growth slowing, increasing abscisic acid from water stress and developmental programming promoting maturation, and possible ethylene involvement in coordinating metabolic transitions and beginning senescence processes. Temperature during progressive lignification shows warm mid-late summer conditions generally promoting rapid maturation progression, with optimal 24-28°C supporting efficient lignification and reserve accumulation, while excessive heat above 32°C may stress vines limiting maturation quality. Light intensity affects progressive lignification through effects on photosynthetic production supporting both lignin biosynthesis and reserve accumulation, with adequate light essential for complete maturation while canopy shading may limit carbon availability in shaded shoots reducing lignification and storage. Water availability shows moderate stress typically developing during this period potentially promoting enhanced maturation through stress-induced developmental responses, while severe sustained stress may impair photosynthetic production limiting lignification completeness and reserve accumulation. Nutrient availability particularly nitrogen shows declining requirements as shoot growth slows with adequate but declining availability supporting maturation, while excessive continued nitrogen availability may prolong growth delaying maturation, and severe deficiency may cause premature incomplete maturation. The process typically spans 6-8 weeks progressing through middle shoot regions with individual internodes requiring 2-3 weeks for substantial lignification depending on position and conditions, with warm conditions and moderate stress promoting steady progression. Cultivar variation appears in progression rate with early-maturing varieties showing rapid steady progression while late-maturing types show slower or interrupted progression, extent of lignification achieved in middle regions with some varieties showing complete thorough lignification while others maintain less complete maturation, and reserve accumulation intensity affecting tissue starch concentrations. Regional timing shows progressive shoot lignification occurring July-August in warm climates, July-September in temperate regions, and August-September in cool areas, with progression rate affected by summer temperature patterns and water availability. Viticultural management considerations include monitoring progression rate and extent assessing vine maturation status and resource adequacy, managing water providing moderate stress promoting maturation without excessive limitation impairing function, controlling nitrogen avoiding excessive late availability prolonging growth and delaying maturation, assessing progression uniformity across vineyard identifying areas with inadequate maturation requiring management attention, and recognizing that proper progressive lignification is essential for adequate reserve accumulation supporting subsequent season productivity. Research applications include quantifying spatial-temporal patterns of maturation progression and their environmental regulation, analyzing coordination between vegetative maturation and reproductive development affecting resource partitioning, evaluating management strategies optimizing maturation timing and completeness, modeling maturation progression for predicting vine status and dormancy preparation, and assessing implications of maturation patterns for productivity and longevity in perennial grapevine cultivation.
Progressive_Shoot_Lignification is second of four Shoot_Lignification subphases, distinguished by systematic acropetal advancement of maturation through middle shoot regions. Follows Basal_Shoot_Lignification and overlaps with ongoing shoot growth. Transitions to Midshoot_Lignification as middle regions complete maturation. Represents period of most active maturation progression requiring adequate resources. Critical phase determining final extent of thorough lignification and reserve accumulation.
acropetal woodification
advancing lignification
akropetale Holzbildung
aufsteigende Ausreifung
fortschreitende Triebverholzung
progressive Triebausreifung
upward shoot maturation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 290-298 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 222-230 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Progressive Shoot Lignification
Merlot in Bordeaux shows progressive softening during September-October Year N+1 at BBCH 85-89, with firmness declining from initial softening 2.7 N through progressive ripening to mature 1.6 N representing total 53% decline from pre-veraison 3.4 N, continued polygalacturonase activity 28 to 72 units, pectin methylesterase 38 to 86 units, progressive pectin solubilization 42% to 68% soluble pectin, molecular weight declining 140 to 78 kDa, middle lamella extensively degraded with cells loosely attached, berries increasingly pliable and soft, slower progressive softening rate 0.9% per day over 6 weeks ripening, under oceanic conditions of 18-24°C temperature, typical of this mid-season cultivar achieving soft ripe texture at harvest maturity
Progressive softening represents the continued texture transformation phase during berry softening when berry firmness continues declining from initial softening through ripening, continued cell wall degradation through sustained pectin-degrading enzyme activities, increasing berry pliability creating progressively softer texture, total firmness decline reaching 40-70% from pre-veraison maximum by full ripeness, firmness declining from initial softening 2.0-3.5 N to ripe 1.0-2.5 N, and progressive softening establishing characteristic soft ripe berry texture over 4-8 weeks ripening period. This continued softening phase occurs during late summer through autumn in late August-October in temperate regions Year N+1, representing ripening texture transformation following initial veraison softening. Progressive softening mechanism involves continued cell wall degradation through sustained polygalacturonase and pectin methylesterase activities, progressive pectin depolymerization and solubilization, continuing middle lamella degradation and cell-cell adhesion loss, possible further minor turgor changes though structural changes remain dominant, and cumulative softening creating soft ripe texture characteristic of mature berries. Firmness decline progression shows firmness continuing to decline from initial softening 2.0-3.5 N through progressive ripening to ripe 1.0-2.5 N, total decline 40-70% from pre-veraison maximum 2.5-4.5 N, typical mature firmness 1.5-2.0 N representing 50-60% total reduction, and progressive decline as ripening characteristic. Softening rate during progressive phase shows slower decline 0.5-1.5% firmness per day compared to rapid initial 2-4% per day, gradual progressive softening over weeks, eventual plateau at soft ripe texture, and rate pattern characteristic of ripening. Firmness trajectory shows rapid initial decline at veraison, slower progressive decline during ripening, eventual soft ripe state, and trajectory as softening pattern. Cultivar variation shows final ripe firmness varying 1.0-2.5 N among cultivars, some cultivars maintaining relatively firmer texture, others becoming very soft, and genetic variation affecting softening extent. Continued cell wall degradation shows polygalacturonase and pectin methylesterase activities remaining elevated through ripening, enzymes continuing pectin degradation, progressive structural modification, and sustained enzymatic activity as degradation mechanism. Polygalacturonase continued activity shows PG activity increasing from initial 15-40 units to ripening maximum 40-100 units, sustained PG continuing pectin cleavage, PG as continuing softening enzyme, and PG activity throughout ripening. Pectin methylesterase continued activity shows PME activity 25-50 units initial increasing to 50-120 units ripening, continued demethylation creating PG substrate, PME sustained through ripening, and PME as continuing modification enzyme. Progressive pectin depolymerization shows pectin molecular weight continuing decline from initial 100-200 kDa to ripe 50-150 kDa or lower, continued chain shortening from enzymatic cleavage, depolymerization progressive through ripening, and molecular weight decline as continuing degradation. Pectin solubilization progression shows soluble pectin continuing increase from initial 35-50% to ripe 60-80% total pectin, insoluble pectin declining to 20-40%, progressive conversion to soluble forms, and solubilization continuing through ripening. Middle lamella continued degradation shows progressive middle lamella pectin loss, cell-cell adhesion continuing weakening, possible cell separation in very soft berries, and middle lamella as continuing degradation target. Cell-cell adhesion progressive loss shows intercellular bonding progressively reduced, cells increasingly loosely attached, tissue coherence declining substantially, and adhesion loss creating soft texture. Cell wall structure progressive weakening shows cell wall pectin network increasingly fragmented, wall structural integrity progressively lost, walls becoming very weak in soft ripe berries, and structural loss as softening basis. Increasing berry pliability shows berries becoming progressively softer and more yielding, texture increasingly pliable to touch, berries easily deformed by gentle pressure, and pliability as softening outcome. Tactile softness shows berries subjectively very soft at ripeness, yielding readily to touch, texture characteristic of ripe fruit, and tactile as practical assessment. Berry handling sensitivity shows soft ripe berries very susceptible to mechanical damage, berries easily crushed or bruised, handling care required, and sensitivity as softening consequence. Juice release shows soft berries releasing juice readily when crushed, softness facilitating juice extraction, texture affecting processing, and juice release as functional property. Skin-flesh separation shows possible skin loosening from flesh in very soft berries, separation from middle lamella degradation, loosening affecting processing and quality, and separation as extreme softening. Turgor possible continued changes shows turgor possibly continuing slight decline or remaining stable during ripening, turgor changes still minor compared to structural changes throughout ripening, turgor contribution secondary 15-30% throughout, and turgor as continuing minor factor. Berry water content shows water content stable or slightly increasing during ripening 78-85%, water uptake from sugar osmotic gradient, water affecting turgor and berry properties, and water content as berry characteristic. Osmotic potential shows osmotic potential becoming increasingly negative from sugar accumulation -1.0 to -1.8 MPa, increased sugars driving water uptake, osmotic gradient as water flux driver, and osmotic potential determining water relations. Berry composition changes shows sugars increasing substantially 14-18% to 20-28%, acids declining 0.7-0.9% to 0.4-0.7%, pH rising 3.1-3.4 to 3.3-3.8, and composition changes accompanying softening. Softening-composition coordination shows softening coordinated with sugar accumulation, acid decline, color development in red cultivars, and integrated ripening processes. Softening completion shows eventual soft ripe texture when berries reach maturity, firmness stabilizing at ripe levels, softening completing ripening transformation, and completion as maturity state. Harvest maturity shows softening as ripeness indicator, soft texture indicating mature berries ready for harvest, optimal firmness varying by use with table grapes requiring firmness while wine grapes soft, and firmness as maturity marker. Environmental effects on progressive softening include temperature affecting enzyme activities and softening rate, warm conditions promoting faster continued softening, cool conditions slowing ripening and softening, and temperature as major rate factor. Temperature effects shows cumulative heat affecting total softening extent and rate, temperature affecting enzyme kinetics throughout ripening, and temperature as dominant environmental factor. Water availability shows adequate water supporting normal softening, deficit possibly affecting softening extent, and water as softening factor. The process encompasses three progressive softening aspects including continued cell wall degradation through sustained enzyme activities, increasing berry pliability creating soft texture, and firmness decline reaching mature levels. Duration progressive softening over 4-8 weeks from veraison through ripening. Regional timing shows progressive softening during late August-October through ripening to harvest. Viticultural significance includes representing continued ripening texture transformation, determining final berry texture affecting quality and handling, demonstrating sustained cell wall modification as ripening process, and establishing soft ripe texture as maturity characteristic. Research applications include analyzing continued softening mechanisms and enzyme regulation, investigating progressive pectin degradation and structural changes, evaluating softening-composition relationships, quantifying firmness decline kinetics through ripening, and assessing progressive softening as ripening progression and quality determinant.
Progressive_Softening is second phase of Berry_Softening, distinguished by continued transformation. Represents 4-8 weeks ripening following initial softening. Firmness decline 2.0-3.5 → 1.0-2.5 N (40-70% total from pre-veraison). Subdivided into Continued_Cell_Wall_Degradation (PG 40-100 units, PME 50-120 units), Increasing_Berry_Pliability, Firmness_Decline. Rate 0.5-1.5%/day (slower than initial). Pectin solubilization 60-80%. Turgor changes minor (15-30% contribution). Follows Initial_Berry_Softening. Establishes soft ripe texture. Temperature major rate factor.
Reifungs-Erweichung
continued softening
extended softening
fortgesetzte Erweichung
progressive Erweichung
progressive Festigkeits-Abnahme
progressive firmness decline
ripening softening
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 482-490 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Progressive Softening
In a Finger Lakes Riesling vineyard, progressive sweetening was evaluated over 35 days from mid-September through October Year N+1 (BBCH 85-87) through weekly taste panels when berries transitioned from tart-dominant (day 0: Brix 14.8°, TA 11.2 g/L, pH 3.06, ratio 1.32:1, taste:sharply tart) through balanced (day 21: Brix 19.6°, TA 8.4 g/L, pH 3.28, ratio 2.33:1, taste:sweet-tart harmonious) to sweet-forward (day 35: Brix 23.2°, TA 6.8 g/L, pH 3.42, ratio 3.41:1, taste:pleasantly sweet with crisp acidity, floral-fruity notes), enabling harvest at optimal balance for off-dry style.
Progressive_Sweetening represents the gradual sensory transformation from tart acidic fruit toward sweet balanced ripe grape through cumulative sugar accumulation and concurrent acid decline creating the characteristic flavor development that signals approaching harvest readiness and defines eating or wine quality potential. This progressive sweetening occurs throughout active ripening (BBCH 84-87), typically days 15-50 post-veraison spanning 30-40 days, and is characterized by taste evaluation showing decreasing tartness perception from dominant sharp acidity (pH 3.0-3.3, TA 10-16 g/L, Brix 13-18°) through balanced sweet-tart transition (pH 3.3-3.6, TA 6-10 g/L, Brix 19-23°) toward sweet-dominant ripe character (pH 3.4-3.8, TA 4-7 g/L, Brix 22-28°), sugar:acid ratio increasing from 1.5-2.5:1 (early active) through 2.5-4.0:1 (mid-active) to 3.5-6.0:1 (late active approaching harvest) representing fundamental balance shift, sensory sweetness threshold (~8-12° Brix) surpassed early in active period after which sweetness increasingly dominant taste component, and flavor complexity developing as esters, terpenes, and other volatiles accumulate 30-150 μg/kg total aromatics complementing sugar-acid balance creating integrated sensory ripeness. Biochemical basis of progressive sweetening includes hexose accumulation increasing vacuolar concentrations from 500-900 mM to 1400-2200 mM (glucose and fructose each 700-1100 mM) creating sweetness through activation of taste receptors particularly T1R2-T1R3 heterodimer responding to sugars, organic acid decline particularly malate reducing from 6-12 g/L to 2-5 g/L diminishing tartness as fewer acid molecules available to activate sour taste receptors (PKD channels responding to protons), pH increase from 3.1-3.4 to 3.4-3.7 reducing proton concentration further moderating sour perception, and flavor compound synthesis with esters (ethyl acetate, isoamyl acetate), terpenes (linalool, geraniol in aromatics), C6 aldehydes and alcohols (hexanal, hexanol) accumulating creating fruity floral aromas enhancing perceived sweetness through cross-modal sensory integration. Sensory perception dynamics demonstrate sweetness intensity increasing logarithmically with sugar concentration following psychophysical power law where doubling sugar produces 5.0:1 achieving very sweet character appropriate for late harvest styles or dessert wines or indicating over-deacidification in warm vintages potentially problematic, and target ratios varying by cultivar and style with high-acid varieties optimal 2.5-4.0:1 while lower-acid types 3.5-5.5:1. Cultivar variation in sweetening patterns shows aromatic varieties (Muscat, Gewürztraminer, Riesling) achieving pronounced sweetness perception through synergy between sugars, low acids, and intense floral-fruity aromas, neutral varieties (Chardonnay, Merlot) displaying sugar-driven sweetening with less aromatic contribution requiring higher absolute sugars for equivalent perceived sweetness, and phenolic-rich reds (Cabernet Sauvignon, Nebbiolo) potentially showing delayed sweetening perception until tannins polymerize reducing astringency masking sweetness. Quality implications for different uses include table grape breeding and selection prioritizing rapid progressive sweetening achieving 16-20° Brix with balanced acidity for fresh consumption, wine grapes requiring slower more balanced progression maintaining adequate acids (final TA 5-8 g/L) while achieving target sugars 20-26° Brix, late harvest or dessert wine production targeting extended ripening achieving very high sugars 26-35° Brix with maintained acids creating intense sweetness with balancing acidity, and raisins requiring progression to 24-30° Brix on vine or through post-harvest drying. Environmental effects on sweetening include warm conditions accelerating both sugar accumulation and acid decline potentially producing rapid sweetening but risk of excessive deacidification creating unbalanced fruit, cool conditions moderating sweetening maintaining higher acids potentially beneficial for acid retention but requiring longer season achieving target sugars, water status with deficit potentially enhancing sweetening perception through concentration effects reducing berry size, and canopy microclimate with sun exposure accelerating sweetening through enhanced berry temperature affecting both sugar and acid metabolism. Harvest timing based on progressive sweetening involves taste evaluation as critical assessment tool confirming analytical measures, recognition that perceived sweetness depends on sugar-acid balance not sugar alone requiring integrated assessment, sensory evaluation detecting flavor development complementing sweetness indicating true ripeness beyond just Brix, varietal character expression important particularly whites and aromatics where flavor ripeness may lag sugar requiring extended hang time, and consumer or winemaker preferences defining target sweetness profile varying by intended use and style. Viticultural management affecting progressive sweetening includes canopy management influencing cluster microclimate affecting sweetening rate through temperature and light effects, crop load adjustment ensuring adequate photosynthesis supporting both sugar accumulation and secondary metabolite synthesis contributing to perceived sweetness, irrigation strategy balancing adequate water for metabolism against deficit enhancing concentration, and harvest timing decisions integrating sensory sweetening assessment with analytical measures ensuring optimal balance achieved.
Progressive_Sweetening is distinguished from initial sweetening by showing gradual transition (30-40 days) rather than onset, from rapid sweetening by representing moderate-paced progression rather than accelerated rates, and from sugar accumulation by focusing on sensory perception rather than analytical measurement. Sugar:acid ratio 1.5-2.5:1 → 3.5-6.0:1. Taste: tart-dominant → balanced → sweet-forward. Duration BBCH 84-87. Sensory assessment critical. Progressive_Sweetening represents perceived ripening transformation defining harvest quality.
Zucker-Säure-Balance-Verschiebung
cumulative taste transformation
gradual flavor development
graduelle Geschmacks-Entwicklung
kumulative Geschmacks-Transformation
sensorische Reifungs-Progression
sensory ripening progression
sugar-acid balance shift
Keller M (2020) The Science of Grapevines 3rd Ed pages 224-228, 251-256 sensory development Boulton RB, Singleton VL, Bisson LF, Kunkee RE (1996) Principles and Practices of Winemaking pages 53-58 taste and balance Noble AC, Bursick GF (1984) The contribution of glycerol to perceived viscosity and sweetness in white wine Amer J Enol Vitic 35:110-112 sweetness perception.
Progressive Sweetening
A proleptic lateral shoot on Vitis vinifera cv. Cabernet Sauvignon, axillary bud formed at node 7 during previous season's primary shoot development (May-June of year 1), bud remained dormant through winter dormancy period, activated in late May of year 2 approximately 12 months post-formation, emerged as lateral shoot from previous season's cane (now mature lignified wood), developed to 55cm length by August of year 2 with 8 leaves, representing unusual proleptic lateral development pattern (delayed dormancy-mediated versus typical sylleptic immediate), this proleptic activation attributed to severe spring frost damage (year 2) destroying most primary buds triggering compensatory activation of secondary buds and also stimulating some axillary buds from previous season including this proleptic lateral, with delayed timing and previous-season origin distinguishing from standard same-season sylleptic laterals though functional role similar providing supplemental photosynthetic capacity during recovery from frost damage.
A lateral shoot developing after dormancy period from axillary bud formed in previous growing season, characterized by bud initiation during primary shoot development one year, winter dormancy period, and shoot emergence following spring or later representing delayed development pattern contrasting with sylleptic immediate same-season pattern, and rare occurrence in standard viticulture where most lateral development follows sylleptic pathway though proleptic laterals occasionally observed in specific circumstances including damage recovery, unusual environmental triggers, or variety-specific delayed lateral tendencies. Proleptic lateral shoots represent delayed dormancy-mediated development, with axillary buds remaining dormant through winter rather than developing immediately, then activating following season creating lateral shoots from previous-season positions, uncommon pattern in grapevine compared to sylleptic dominance though occurring in specialized contexts.
Proleptic lateral shoots are relatively uncommon in viticulture compared to sylleptic dominance, with most lateral development following immediate same-season pattern rather than delayed dormancy-mediated pathway. However, proleptic laterals occur in specific contexts including damage recovery (severe primary bud loss triggering activation of all available bud positions including previous-season axillary buds), environmental extremes (unusual temperature or moisture patterns stimulating dormant axillary activation), variety-specific tendencies (certain cultivars or clones showing occasional proleptic lateral development), or hormonal factors (apical dominance reduction, cytokinin increases, other signals triggering delayed bud activation). The proleptic pattern creates shoots arising from previous-season wood positions, potentially creating architectural confusion (laterals emerging from mature canes rather than current shoots), management uncertainties (assessing origin and purpose of unexpectedly positioned shoots), and functional questions (evaluating contribution and retention decisions for delayed laterals). Management typically involves case-by-case assessment based on origin understanding and functional evaluation.
delayed-season lateral
dormant lateral
dormanter Achseltrieb
following-year lateral
proleptischer Seitentrieb
Proleptic Lateral Shoot
Proline accumulation quality comparison across Vitis vinifera cultivars in a controlled drought stress trial identified substantial genotypic variation relevant to cultivar selection for water-limited environments. Sultana demonstrated exceptional proline accumulation quality with leaf proline concentration increasing from 1.5 μmol g⁻¹ fresh weight under well-watered conditions to 28 μmol g⁻¹ under moderate stress (predawn water potential -0.5 MPa), an 18-fold increase representing one of the highest accumulation responses documented in grapevine. This proline accumulation quality contributed approximately 0.15 MPa to total osmotic adjustment and correlated with maintained photosynthesis and delayed wilting symptoms compared to low-accumulating cultivars. Cabernet Sauvignon showed moderate proline accumulation quality with 8-fold increase to 12 μmol g⁻¹ under equivalent stress, while Chardonnay exhibited poor proline accumulation quality with only 3-fold increase to 4.5 μmol g⁻¹ despite equivalent stress exposure, corresponding to earlier turgor loss and greater yield reduction under deficit irrigation. Molecular analysis revealed that high proline accumulation quality in Sultana was associated with stronger P5CS gene expression induction and reduced proline dehydrogenase expression under stress, identifying the regulatory basis of the accumulation phenotype.
Proline accumulation quality is an osmotic adjustment quality that inheres in a grapevine organism and represents the capacity to synthesize and accumulate the amino acid proline in response to water deficit stress, with proline serving as a primary compatible osmolyte that contributes substantially to osmotic adjustment while providing additional protective functions including protein stabilization, reactive oxygen species scavenging, and cellular redox buffering that enhance stress tolerance beyond the direct osmotic effects. This quality captures the contribution of proline specifically to drought tolerance, recognizing that proline accumulation represents one of the most universal and well-characterized stress responses in plants with particular prominence in grapevine where substantial genotypic variation in accumulation capacity has been documented. The ontological significance of proline accumulation quality lies in the multifunctional protective role of this amino acid that makes it a particularly effective stress response, with the combination of osmotic, protective, and signaling functions creating benefits that exceed those predicted from osmotic adjustment alone, and with the magnitude of proline accumulation serving as a reliable indicator of stress intensity and genotypic tolerance capacity. The biological basis of proline accumulation encompasses the stress-induced activation of the biosynthetic enzyme pyrroline-5-carboxylate synthetase (P5CS) that catalyzes the rate-limiting step in proline synthesis from glutamate, combined with suppression of proline degradation through proline dehydrogenase, together creating the net accumulation observed under stress. The magnitude of proline accumulation under stress varies dramatically among grapevine genotypes, from 2-3 fold increases in low-accumulating types to 10-20 fold or greater in high-accumulating genotypes, providing a biochemical marker of drought tolerance capacity that correlates with whole-plant stress performance in many studies. The relationship between proline accumulation and drought tolerance involves both the direct osmotic contribution of accumulated proline and its indirect protective effects on proteins, membranes, and cellular redox status that collectively maintain cellular function under stress.
Proline_Accumulation_Quality is_a Osmotic_Adjustment_Quality, representing the specific contribution of proline to osmotic adjustment and stress protection. This class addresses proline as the most studied and often predominant osmolyte in grapevine stress response. The fold-increase from baseline provides standard quantification with >10-fold representing high accumulation capacity. The multifunctional protective effects beyond osmotic adjustment distinguish proline from some other osmolytes. P5CS expression serves as molecular marker of accumulation capacity. Critical for biochemical characterization of drought tolerance mechanisms.
Prolin-Akkumulation
Prolin-Osmoprotektion
Prolin-Synthesekapazität
proline osmoprotection
proline response
proline synthesis capacity
stress-induced proline
stressinduzierte Prolin-Anreicherung
Patakas A et al. (2002) Leaf age effects on solute accumulation in water-stressed grapevines. Journal of Plant Physiology 159:969-976 Cramer GR et al. (2007) Water and salinity stress in grapevines: early and late changes in transcript and metabolite profiles. Functional and Integrative Genomics 7:111-134 Szabados L, Savouré A (2010) Proline: a multifunctional amino acid. Trends in Plant Science 15:89-97
Proline Accumulation Quality
Proline content quality in Napa Valley Cabernet Sauvignon serves as an indicator of vine water stress during ripening, with deficit-irrigated hillside vineyards producing must with 1200-1800 mg/L proline (indicating controlled water stress promoting phenolic maturity), compared to 600-900 mg/L from valley-floor sites with greater water availability winemakers interpreting proline alongside other maturity indices use elevated proline as confirmation of the moderate water stress conditions associated with concentrated, age-worthy Cabernet Sauvignon, while very high proline (above 2000 mg/L) may indicate excessive stress limiting photosynthesis and optimal ripening.
Proline_Content_Quality represents the chemical quality characterizing the concentration of the amino acid proline in grape must and wine, capturing the typically dominant amino acid that paradoxically does not contribute to yeast assimilable nitrogen under standard fermentation conditions. Proline is unique among grape amino acids for its cyclic structure (a pyrrolidine ring incorporating the alpha-amino group) that prevents its catabolism by Saccharomyces cerevisiae under the anaerobic conditions of wine fermentation, as proline oxidase requires molecular oxygen for activity. In grape must, proline typically comprises 30-60% of total free amino acids, with concentrations ranging from 200-2000 mg/L depending on cultivar, vine water status, and fruit maturity. Proline accumulates dramatically during berry ripening, particularly under water stress conditions that promote proline synthesis as an osmoprotectant. This stress-induced accumulation makes proline content a potential indicator of vine water status during ripening, with high proline suggesting water deficit stress. Cultivar genetics strongly affect proline content, with some varieties (Cabernet Sauvignon, Chardonnay) showing characteristically high proline while others (Riesling, Gewürztraminer) accumulate less. The non-assimilability of proline means it persists through fermentation into finished wine, where it contributes to wine amino acid content without providing yeast nutrition. Proline's persistence enables its use for cultivar fingerprinting and authenticity verification, as the cultivar-specific proline patterns remain detectable in wine. Some non-Saccharomyces yeasts and lactic acid bacteria can metabolize proline under specific conditions, potentially affecting proline content in wines produced with these organisms.
Proline_Content_Quality addresses the most abundant amino acid in many grape musts despite its exclusion from YAN calculations. The non-assimilability under anaerobic conditions reflects the oxygen requirement of proline oxidase, the initial enzyme in proline catabolism. This metabolic limitation is specific to Saccharomyces under fermentation conditions proline can be utilized by yeast under aerobic conditions and by certain non-Saccharomyces species. The stress-responsive accumulation of proline links its content to vine physiology during ripening. Proline analysis employs the same HPLC methods as other amino acids, with its distinctive retention time enabling reliable identification.
L-Prolin-Konzentration
L-proline concentration
Prolingehalt
cyclic amino acid content
proline level
stress-responsive amino acid quality
stressresponsiver Aminosäuregehalt
zyklischer Aminosäuregehalt
Huang Z and Ough CS (1991) Amino Acid Profiles of Commercial Grape Juices and Wines. American Journal of Enology and Viticulture 42(3): 261-267 Deluc LG et al. (2009) Water Deficit Alters Differentially Metabolic Pathways Affecting Important Flavor and Quality Traits in Grape Berries of Cabernet Sauvignon and Chardonnay. BMC Genomics 10(1): 212
Proline Content Quality
A prompt lateral shoot on Vitis vinifera cv. Riesling arising from node 6 axillary bud on primary shoot, emerged in early July approximately 6 weeks after primary shoot reached this node during mid-May elongation, developed to 38cm length by late August with 6 leaves totaling approximately 570 cm² leaf area, representing typical prompt lateral development in vigorous Riesling showing characteristic propensity for lateral shoot production, this lateral emerged following hedge-topping of primary shoot at 165cm height in late June (apex removal releasing lateral buds from apical dominance), contributing supplemental photosynthetic capacity (17% increase over primary shoot base area) valuable for supporting three-cluster crop load typical of high-fruitfulness Riesling, lateral retained and positioned rather than removed recognizing beneficial photosynthetic contribution outweighing minor density increase in well-managed VSP canopy with adequate spacing.
A lateral shoot developing during the same growing season as the primary shoot from which it arises, characterized by emergence from axillary buds within weeks or months of primary shoot node formation, concurrent growth alongside continued primary shoot development, typical timing from late spring through mid-summer depending on node position and environmental conditions, and function as same-season canopy supplementation increasing leaf area and photosynthetic capacity though potentially creating density problems if excessive or poorly positioned. Prompt lateral shoots represent immediate axillary bud activation without dormancy period, arising through various triggers including vigorous growth conditions promoting lateral breakage, apical damage or removal releasing lateral buds from suppression, hormonal signals favoring lateral development, or variety-specific tendencies toward prompt lateral production, with their same-season emergence enabling photosynthetic contribution during current growing season supporting fruit development though requiring management assessment of benefits versus density concerns.
Prompt lateral shoots provide same-season functional benefits through increased leaf area and photosynthetic capacity, with their current-year emergence enabling immediate contribution to fruit development and whole-vine carbon balance. However, prompt laterals create management decisions balancing benefits (additional photosynthesis, potential compensation for primary shoot deficiencies) against costs (increased canopy density potentially causing shading, labor requirements for positioning or removal, disease pressure in dense canopies). Factors promoting prompt lateral development include high vine vigor (excessive resources channeled into lateral activation), apical damage or hedging (removing apical dominance releasing laterals), variety characteristics (some cultivars naturally producing abundant prompt laterals), and environmental conditions (favorable temperature and moisture supporting rapid growth). Management approaches range from routine removal (simplifying canopy architecture and reducing density), selective retention (keeping well-positioned laterals contributing photosynthetic benefits), to acceptance or encouragement (certain vigorous varieties or training systems benefiting from lateral development). The prompt lateral decision requires site-specific assessment of vigor, variety, spacing, and management objectives.
current-year lateral
immediate lateral shoot
prompter Seitentrieb
same-season lateral
sofortiger Achseltrieb
Based on Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 113-145.
Prompt Lateral Shoot
3309 Couderc rootstock hardwood cuttings in commercial nursery propagation showing propagation cutting adventitious roots, 5000 cuttings prepared from certified mother plants, cuttings standardized 35 cm length 8-12 mm diameter 4 nodes, treated with 3000 ppm IBA quick-dip, planted in propagation beds with peat-perlite medium, bottom heat maintaining 22°C root zone temperature, rooting monitored weekly with 82% cuttings showing roots by day 25, final evaluation day 45 showing 78% achieving commercial grade with 4+ roots >8 cm length, rooted cuttings representing commercial rootstock production
Propagation cutting adventitious root is cutting adventitious root emphasizing commercial propagation context and nursery production function, characterized by formation on cuttings specifically prepared and managed for producing propagation stock, showing development under controlled nursery conditions optimized for maximizing rooting percentage, root system quality, and plant uniformity, demonstrating protocols designed for commercial scale including standardized cutting preparation, hormone treatment, environmental control, and quality assessment, and serving commercial nursery production providing thousands to millions of rooted plants annually for vineyard establishment, with propagation cutting roots showing typical production characteristics including systematic cutting collection from certified mother plants, standardized treatment protocols ensuring consistent results, controlled rooting environments in propagation beds greenhouses or mist systems, regular monitoring and quality control assessing rooting progress, and grading and selection retaining only plants meeting quality standards for commercial sale, and demonstrating that propagation cutting rooting represents foundation of nursery industry supplying grafted or own-rooted vines for commercial vineyards with production protocols refined over decades optimizing efficiency, success rates, and economic viability. Propagation cutting adventitious root in rootstock production includes= showing hardwood cutting use (rootstock varieties propagated primarily through dormant hardwood cuttings), exhibiting mother plant systems (dedicated mother plants or stool beds providing cutting source maintaining genetic identity and health), demonstrating cutting preparation (standardized length, diameter, node number ensuring adequate reserves and rooting potential), revealing treatment protocols (hormone type and concentration optimized for each rootstock variety), and displaying large-scale rooting (rootstock nurseries producing hundreds of thousands of cuttings annually). Propagation cutting adventitious root quality standards includes= requiring minimum root number (commercial standards typically requiring 3-5+ well-distributed roots for acceptable quality), exhibiting root length requirements (roots typically needing 5-10+ cm length for adequate function), demonstrating root system uniformity (commercial production selecting plants with symmetric well-developed root systems), revealing health standards (roots free of pathogens, necrosis, or abnormalities), and displaying grading systems (plants graded by root system quality with premium, standard, and cull categories). Propagation cutting adventitious root production facilities includes= showing propagation bed systems (outdoor beds with controlled media, moisture, and temperature), exhibiting greenhouse propagation (controlled environment structures providing optimal conditions year-round), demonstrating mist systems (automated misting maintaining humidity for green cutting propagation), revealing bottom heat application (heating cables or systems maintaining optimal root zone temperature), and displaying mechanization (some operations using automated cutting preparation, treatment, planting, and handling). Propagation cutting adventitious root treatment optimization includes= requiring cultivar-specific protocols (different rootstock varieties needing different hormone concentrations, timing, and conditions), showing concentration optimization (testing determining optimal IBA or NAA concentrations maximizing rooting), exhibiting formulation selection (powder, liquid quick-dip, or long-soak formulations chosen for efficiency and results), demonstrating timing refinement (collection timing, callusing period, planting date optimized for each variety), and displaying environmental control (temperature, humidity, light, media fine-tuned maximizing success). Propagation cutting adventitious root monitoring and assessment includes= showing rooting checks (periodic sampling examining cutting bases assessing rooting progress), exhibiting emergence timing (recording when roots first visible indicating rooting success), demonstrating percentage calculation (determining proportion of cuttings successfully rooting indicating protocol effectiveness), revealing problem diagnosis (identifying causes of poor rooting such as inadequate hormone, improper environment, or pathogens), and displaying quality evaluation (assessing root system characteristics determining commercial acceptability). Propagation cutting adventitious root economic considerations includes= requiring cost-effectiveness (propagation protocols balancing costs of inputs, labor, facilities against plant value), exhibiting scale economies (large operations achieving lower per-plant costs through efficiency), demonstrating market demands (production volumes and timing driven by vineyard planting demands), revealing pricing structures (plant prices reflecting production costs, quality, variety, and market conditions), and displaying profitability requirements (nurseries requiring adequate margins maintaining business viability). Propagation cutting adventitious root certification and traceability includes= showing certified source material (cuttings from certified virus-tested mother plants ensuring health), exhibiting identity maintenance (rigorous protocols preventing mixing maintaining cultivar identity), demonstrating documentation (records tracking cutting source, treatment, rooting, and quality for traceability), revealing inspection compliance (government inspections verifying compliance with certification standards), and displaying labeling systems (plants labeled with variety, rootstock, certification status, nursery, and year). Propagation cutting adventitious root challenges and innovations includes= showing difficult cultivars (some varieties showing poor rooting requiring research and protocol development), exhibiting climate effects (seasonal weather variation affecting outdoor propagation requiring adaptation), demonstrating pest and disease management (propagation facilities managing pathogens and insects without compromising plant quality), revealing technology adoption (innovations like rooting stimulants, biostimulants, or novel propagation systems potentially improving results), and displaying sustainability initiatives (propagation industry exploring reduced chemical inputs, water conservation, and environmental stewardship).
Propagation_Cutting_Adventitious_Root emphasizes commercial nursery production context and is distinguished from other cutting roots by focus on systematic commercial protocols. Critical for understanding rootstock production, quality standards, treatment optimization, scale operations, economic considerations, and the adventitious roots produced under controlled nursery conditions providing commercial propagation stock for viticulture industry.
Baumschul-Stecklingswurzel
Commercial propagation root
Kommerzielle Produktionswurzel
Nursery cutting root
Production cutting root
Vermehrungs-Stecklings-Adventivwurzel
Propagation Cutting Adventitious Root
A bundle of dormant Cabernet Sauvignon canes harvested from a certified scion mother plant bears a Propagation_Material_Role, destined for cutting into benchgraft scion pieces after cold storage. Hardwood cuttings of 3309 Couderc rootstock serve in a Propagation_Material_Role, ready for rooting as rootstock liners or as the rootstock component of benchgrafts. A tray of tissue culture Chardonnay plantlets bears a Propagation_Material_Role, representing virus-eliminated material multiplied through micropropagation for eventual field establishment. A completed benchgraft (Merlot scion grafted to SO4 rootstock) serves in a Propagation_Material_Role as the final nursery product awaiting callusing and establishment.
A propagation role that inheres in a plant part of Vitis vinifera or a Vitis hybrid by virtue of its function as the physical material used for vegetative multiplication, encompassing cuttings, budwood, grafted units, and tissue culture explants that carry genetic identity and health status from parent to progeny plants. Propagation material roles describe the functional categories of plant parts within the nursery multiplication chain, from cuttings (hardwood, softwood, semi-hardwood of various node configurations) through grafted combinations (benchgrafts, field grafts) to specialized formats (tissue culture plantlets, green grafts). The quality of propagation material directly determines nursery success rates, with key parameters including carbohydrate reserves (supporting root and shoot initiation), bud viability (freedom from mechanical damage, freezing injury, or pathogen infection), cane maturity (adequate lignification for dormancy survival), and diameter (influencing grafting compatibility and rooting vigor). Propagation material represents the tangible output of mother plant maintenance and the input to nursery production, with certification category and health status inherited from source material and maintained through chain-of-custody documentation.
Propagation_Material_Role is subclassed by material type—Cutting_Role (the dominant category for grapevine propagation), Benchgraft_Role, Field_Graft_Role, Tissue_Culture_Plantlet_Role—reflecting the diverse multiplication pathways used in commercial and research viticulture. The category captures the transformation from source plant to multiplication unit, with quality assessment determining suitability for propagation. Propagation material standards vary by certification scheme and market EU regulations specify minimum quality parameters for marketed material including diameter ranges, node counts, and freedom from specified symptoms. The economics of propagation material production involve tradeoffs between multiplication rate (maximized by smaller cutting units), rooting success (generally higher with larger cuttings), and handling efficiency (favoring standardized formats).
Pflanzgut
Rebvermehrungsmaterial
Vermehrungsmaterial
multiplication material
propagation material
Hartmann HT et al. (2011) Plant Propagation: Principles and Practices 8th Ed, Pearson pp.: 344-414 European Union (2020) Commission Implementing Directive 2020/177 Annex I Gramaje D, Armengol J (2011) Fungal trunk pathogens in the grapevine propagation process, Phytopathologia Mediterranea 50: S217-S226
Propagation Material Role
Propagation rights for the ENTAV Pinot Noir Dijon clones (113-115, 667, 777, 828) are administered under the French catalogue d'inscription system, which grants ENTAV perpetual institutional rights over the foundation material while permitting licensed nurseries to propagate without individual clone royalties — the access cost being incorporated into the nursery licensing fee and annual catalogue registration charges. Propagation rights for UC Davis Foundation Plant Services selections operate under a different model, with public university material available to any licensed nursery at nominal material transfer fees, while privately selected clones (e.g., certain Wente Chardonnay selections) may carry plant patent protection (17-year term under US Patent Act § 161) requiring per-vine royalty payments of $0.15-0.50 to the patent holder.
Propagation rights is a distribution record entity that documents the legal authorizations, intellectual property protections, licensing agreements, and contractual arrangements governing the multiplication, sale, and distribution of a certified clonal line, including plant variety protection (PVP) or plant patent status, breeder's rights designations, exclusive or non-exclusive licensing terms, royalty structures, geographic restrictions on propagation, and the duration and conditions of propagation authorization, defining the legal framework within which nurseries and growers may access and multiply the clonal material.
Propagation rights constitute the legal-economic framework that simultaneously incentivizes investment in clonal selection (by enabling financial return) and governs access to improved plant material (by defining who may propagate and under what conditions). The international legal landscape for grapevine propagation rights is complex, involving the intersection of UPOV Convention plant variety protection (available in 78 member states), national patent systems (US plant patents, European utility models), and institutional policies of public research organizations. The distinction between variety-level protection (covering all clones of a named variety, relevant for new breeding program releases) and clone-level protection (covering a specific clonal selection within an existing variety) is legally significant — most traditional Vitis vinifera cultivar clones cannot be variety-protected because the underlying variety lacks novelty, but clone-specific patents or breeder's rights may apply to the individual selection. Licensing models range from open access (public institution selections freely available to licensed nurseries), through moderate licensing (nominal fees covering administrative and testing costs), to restrictive commercial licensing (exclusive territorial agreements with significant per-vine royalties). Geographic restrictions on propagation rights can create market fragmentation — a clone licensed for propagation in Europe may require separate authorization for multiplication in the Americas or Australasia, with different royalty structures in each jurisdiction. The enforcement of propagation rights is complicated by the vegetative nature of grapevine propagation, as unauthorized multiplication of protected material is difficult to detect without molecular identity verification, and the long productive lifespan of vineyards (25-40 years) means that material planted under valid licenses continues producing long after rights may have expired or transferred.
Lizenzvereinbarung
Sortenrechte
Vermehrungsgenehmigung
Vermehrungslizenz
Vermehrungsrechte
Züchterrechte
breeder's rights
clone licensing
multiplication rights
plant variety rights
propagation authorization
propagation license
UPOV (2011) International Convention for the Protection of New Varieties of Plants, Act of 1991 Boidron R et al. (1995) Catalogue des variétés et clones de vigne cultivés en France, ENTAV-INRA, pp.: 8-15 Alleweldt G, Dettweiler E (1994) The genetic resources of Vitis: world list of grapevine collections. Bundesanstalt für Züchtungsforschung an Kulturpflanzen, Institut für Rebenzüchtung, Siebeldingen
Propagation Rights
A Pinot Noir vine maintained in a foundation vineyard at ENTAV-INRA in France bears a Propagation_Role as a Basic category mother plant, providing budwood for certified nursery production while itself being derived from virus-tested Pre-Basic nuclear stock. A 3309 Couderc rootstock plant in a California Foundation Plant Services (FPS) increase block serves in a Propagation_Role, multiplying tested material for distribution to commercial nurseries. A benchgraft produced by joining Chardonnay scion to SO4 rootstock in a commercial nursery bears a Propagation_Role as the final product of the certification chain, destined for vineyard establishment.
A grapevine role that inheres in a plant or plant part of Vitis vinifera or a Vitis hybrid by virtue of its function in the vegetative multiplication system that produces new grapevines for vineyard establishment, maintaining genetic identity and health status across generations of clonal propagation. Propagation roles encompass the hierarchical certification system controlling mother plant status (Pre-Basic, Basic, Certified categories), the functional roles of plant parts in multiplication (rootstock cuttings, scion budwood, benchgrafts), and the nursery production chain from elite nuclear stock through commercial planting material. Grapevine propagation is exclusively vegetative for commercial production—maintaining cultivar and clonal identity that would be lost through sexual reproduction—utilizing hardwood cuttings, green cuttings, or tissue culture to multiply genetically identical plants. The propagation system is highly regulated to prevent dissemination of graft-transmissible pathogens (viruses, phytoplasmas, bacteria) that cannot be eliminated from infected plants once established, with certification schemes requiring regular testing and maintaining separation between propagation categories to preserve the health status pyramid.
Propagation_Role is the parent class for the hierarchical certification system that governs grapevine multiplication globally. The role structure reflects the EU certification framework (Directive 68/193/EEC and subsequent amendments) adopted with variations worldwide, distinguishing Pre-Basic (nuclear/elite stock), Basic (foundation), and Certified (commercial) categories based on testing protocols, generation limits, and permissible pathogen levels. Propagation roles also encompass functional designations (Mother_Plant, Rootstock_Cutting, Scion_Budwood) describing the plant part's role in the grafting and multiplication process. The economic importance of healthy propagation material has driven substantial investment in certification infrastructure, with programs like FPS (USA), ENTAV (France), VCR (Germany), and AGRIOS (Italy) maintaining tested nuclear stocks of major varieties.
Rebvermehrung
Vermehrungspflanze
Vermehrungsrebe
multiplication plant
nursery stock plant
propagation plant
European Union (2020) Commission Implementing Directive 2020/177 on propagating material of vine Maliogka VI et al. (2015) Grapevine virus diseases, in Compendium of Grape Diseases 2nd Ed, APS Press pp.: 95-135 Golino DA et al. (2017) Grapevine leafroll disease and its management, in Grapevine Viruses: Molecular Biology, Diagnostics and Management, Springer pp.: 367-381
Propagation Role
The Protected Designation of Origin Tokaji (registered in the EU eAmbrosia database as a PDO under EU Regulation 1308/2013) protects the name for wines produced within the 27-commune Tokaj-Hegyalja region of northeastern Hungary from 6 authorized cultivars (Furmint, Hárslevelű, Sárga Muskotály, Zéta, Kövérszőlő, Kabar) according to the registered specification — the PDO protection extends to the EU single market and partner countries through bilateral trade agreements, with enforcement actions against unauthorized usage (including the resolution of the long-standing Tokaj/Tocai naming dispute with Italy that resulted in the prohibition of the name Tocai Friulano for Italian wines from 2007).
A geographic indication that has been formally registered and legally protected under a national or supranational intellectual property framework, granting exclusive usage rights for the designation to producers within the delimited area who comply with the registered production specification, and providing legal enforcement mechanisms against unauthorized use, imitation, or evocation of the protected name.
Protected_Designation_Of_Origin represents the strongest form of geographic indication protection available under international and EU law. The EU PDO system (Regulation 1308/2013, replacing the earlier AOC/DOC national frameworks under a unified European regime) requires that the wine's quality and characteristics are essentially due to its geographic environment including natural and human factors, that production occurs entirely within the defined geographic area, and that the wine is made exclusively from grapes harvested within that area: these requirements are stricter than those for Protected Geographical Indications (PGI), where only reputation rather than intrinsic quality linkage is required. Subclasses capture the key informational components of PDO registration — the registered name, registration number, and specification reference.
PDO
eingetragene Ursprungsbezeichnung
g.U.
geschützte Ursprungsbezeichnung
protected origin designation
registered designation of origin
EU Regulation 1308/2013 Art. 93-108 WTO (1994) TRIPS Agreement Art. 22-24 EU eAmbrosia database of registered geographic indications Lisbon Agreement for the Protection of Appellations of Origin (1958, revised 2015 as Geneva Act)
Protected Designation Of Origin
The Protected Geographic Indication Pays d'Oc (registered as PGI-FR-A0766 in the EU eAmbrosia database) covers approximately 120,000 hectares across the Languedoc-Roussillon region of southern France, permitting a far wider range of grape varieties (58 authorized cultivars including international varieties such as Cabernet Sauvignon, Merlot, Chardonnay, and Sauvignon Blanc) and higher maximum yields (90 hl/ha versus 45-60 hl/ha for regional PDOs) than the PDO appellations within the same territory — the PGI framework enables producers to blend across communes and use varietal labeling (cépage on the label), providing commercial flexibility that PDO regulations typically prohibit, resulting in Pays d'Oc becoming the largest single PGI in France with annual production exceeding 600 million bottles.
A geographic indication that has been formally registered and legally protected under a national or supranational framework, certifying that the wine possesses a specific quality, reputation, or other characteristic attributable to its geographic origin, with at least one stage of production (grape growing, winemaking, or aging) occurring within the defined area, representing a less restrictive geographic protection tier than Protected Designation of Origin.
Protected_Geographic_Indication occupies the intermediate tier between wines without geographic protection (formerly vin de table, now vin de France/vino/Wein) and wines with full PDO protection. The key regulatory distinction from PDO is the relaxed origin linkage requirement: while PDO demands that quality and characteristics are essentially due to the geographic environment, PGI requires only that a specific quality, reputation, or characteristic is attributable to the geographic origin — a lower evidentiary threshold that permits broader production rules. Additionally, PGI wines need not be produced entirely within the delimited area (at least one production stage must occur there), whereas PDO requires complete production within the geographic zone. Subclasses capture the informational components of PGI registration parallel to the PDO subclass structure.
PGI
g.g.A.
geschützte Herkunftsangabe
geschützte geografische Angabe
protected geographic indication for wine
registered geographic indication
EU Regulation 1308/2013 Art. 93(1)(b) on protected geographical indications WTO (1994) TRIPS Agreement Art. 22 EU eAmbrosia database of registered geographic indications
Protected Geographic Indication
The protected trade name Cabernet Jura is a registered trademark of Valentin Blattner in Switzerland and the EU for his interspecific hybrid breeding VB Cal. 1-28, with trademark registration (Swiss trademark No. 567893) granting exclusive commercial use — unauthorized use of the name for planting material or wine products constitutes trademark infringement, and Blattner licenses the name to nurseries and wine producers through contractual agreements that include quality specifications and royalty payments of 0.10-0.25 CHF per grafted vine sold, illustrating the economic function of protected trade names in incentivizing private breeding investment.
A trade name that has been formally registered as a trademark, protected geographical indication, or equivalent intellectual property right within one or more legal jurisdictions, granting the registrant exclusive commercial use of the designation for grapevine planting material or wine products and providing legal recourse against unauthorized usage.
Protected_Trade_Name represents the strongest form of commercial naming protection available for grapevine cultivars, operating at the intersection of plant variety rights (UPOV), trademark law, and geographic indication systems. The legal landscape is complex because multiple overlapping protection mechanisms may apply to a single cultivar: a PIWI variety may simultaneously carry UPOV Plant Variety Protection (controlling propagation rights), a registered trademark on the cultivar name (controlling commercial use of the name), and national variety registration (controlling planting authorization). Protected trade names are particularly important in the PIWI (disease-resistant cultivar) sector where private breeders invest 15-25 years in cultivar development and depend on naming protection to recoup breeding investment through licensing revenue.
eingetragene Marke
geschützter Handelsname
legally protected designation
markenrechtlich geschützte Bezeichnung
registered trade name
trademark-protected name
UPOV (2020) International Convention for the Protection of New Varieties of Plants Art. 20 Swiss Federal Institute of Intellectual Property trademark database EU Trademark Regulation 2017/1001 ICNCP (2016) International Code of Nomenclature for Cultivated Plants 9th Ed, ISHS Art. 11
Protected Trade Name
Cabernet Sauvignon protective cuticle layer providing strong barrier= cutin matrix 3.8-5.2 μm thick, wax content 165 μg/cm² well-integrated with cutin, water vapor permeability 0.95×10⁻⁹ m s⁻¹ (low), transpiration 1.0 g H₂O m⁻² h⁻¹, cuticle remaining intact without major cracking through berry expansion supporting berry protection against dehydration in warm Napa Valley climate and against Botrytis infection during ripening, with protective cuticle contributing to Cabernet Sauvignon suitability for warm dry climates requiring good berry stress tolerance. Syrah protective cuticle layer drought tolerance role= well-watered vines showing 4.2 μm cuticle, 145 μg/cm² wax, 1.15×10⁻⁹ permeability deficit-irrigated vines showing enhanced protective cuticle 5.8 μm thickness, 195 μg/cm² wax, 0.85×10⁻⁹ permeability (26% reduction), with enhanced protective cuticle under deficit limiting berry water loss to 0.9 g H₂O m⁻² h⁻¹ versus 1.3 g H₂O m⁻² h⁻¹ well-watered, demonstrating water deficit acclimation through protective cuticle enhancement contributing to maintained berry hydration despite limited water supply. Chardonnay protective cuticle layer showing moderate protection= thinner cuticle 2.8-4.2 μm, moderate wax 95 μg/cm², higher permeability 1.5×10⁻⁹ m s⁻¹, creating vulnerability to Botrytis bunch rot particularly in humid climates, requiring intensive canopy management (leaf removal for air circulation, cluster zone exposure) compensating for less protective cuticle through environmental modification reducing disease pressure. Riesling protective cuticle layer post-harvest function= intact protective cuticle limiting weight loss to 2-4% during 4-week cold storage (0-2°C, 90% RH) damaged cuticle (rough handling creating micro-damage) showing 8-15% weight loss under same conditions, demonstrating protective cuticle critical for post-harvest quality maintenance with gentle handling essential for preserving cuticle integrity and berry quality. Pinot Noir protective cuticle layer splitting susceptibility= moderate protective cuticle (4.2 μm, 135 μg/cm² wax) showing lower elastic capacity than thick-skinned cultivars, with late-season rainfall (25 mm at week 17 near harvest) creating 8-15% splitting incidence as rapid water uptake increases berry turgor exceeding protective cuticle mechanical strength, versus Cabernet Sauvignon under same conditions showing <3% splitting due to more elastic protective cuticle tolerating turgor increases, demonstrating cultivar variation in protective cuticle mechanical properties affecting splitting susceptibility and late-season rain tolerance.
A protective cuticle layer is a cuticle layer emphasizing barrier and protective functions against environmental stresses, water loss, and pathogen invasion, typically showing well-developed cutin matrix (3-6 μm thickness), moderate-to-high wax content (100-200+ μg/cm²), low water permeability (<1.5×10⁻⁹ m s⁻¹), and structural integrity resisting cracking and damage, representing cuticle optimized for berry protection during development, ripening, and post-harvest period. Protective cuticle layer structural features supporting protection= (1) thick continuous cutin matrix (3-6 μm) providing uninterrupted polymeric barrier without major gaps, cracks, or weak points (2) well-integrated cutin-wax composite with waxes embedded in cutin matrix (intracuticular wax) creating continuous hydrophobic pathway in addition to epicuticular surface wax (3) strong adhesion to underlying epidermal cell wall preventing cuticle delamination or peeling (4) elastic properties allowing cuticle to stretch during berry expansion without excessive cracking, with some cultivars showing cuticle mechanical properties (tensile strength, elastic modulus) supporting expansion accommodation. Protective cuticle layer barrier functions= (1) water loss prevention limiting berry transpiration to <1.5 g H₂O m⁻² h⁻¹ under normal conditions (25°C, 50% RH), preventing berry dehydration during ripening particularly important in warm dry climates (2) pathogen exclusion providing physical barrier requiring fungi to produce cutinase enzymes or to enter through natural openings (stomata, lenticels, wounds) or cuticle cracks, with intact protective cuticle reducing Botrytis and other fungal infection incidence (3) UV protection through cuticle absorbance and scattering reducing UV-B transmission to epidermal cells preventing photodamage (4) chemical barrier limiting leaching of berry compounds into surface water films and limiting uptake of surface-applied chemicals (pesticides, nutrients) requiring surfactants for penetration. Protective cuticle layer in stress tolerance= drought tolerance enhanced through reduced cuticular transpiration maintaining berry water status during water deficit heat tolerance supported through reduced water loss preventing berry dehydration under high temperature stress pathogen resistance enhanced particularly against surface-colonizing fungi requiring cuticle penetration for infection post-harvest protection maintaining berry quality during storage and shipping by limiting water loss, decay, and shriveling. Protective cuticle layer development optimization= sunlight exposure enhancing cuticle thickness and wax deposition creating more protective cuticle in exposed berries moderate water deficit during berry development potentially enhancing cuticle development as acclimation response mineral nutrition particularly calcium potentially affecting cuticle and cell wall properties influencing protective capacity genetic variation with some cultivars/selections producing more protective cuticles showing thicker cutin, higher wax, and better mechanical properties. Protective cuticle layer vulnerability and damage= rapid berry expansion during Phase II ripening potentially exceeding cuticle elastic capacity creating microcracks particularly at berry equator lenticel development creating natural discontinuities in cuticle providing gas exchange but also potential water loss and pathogen entry points mechanical damage during harvest and handling disrupting cuticle integrity creating sites for increased water loss and decay late-season rainfall creating high turgor increasing splitting risk particularly in cultivars with less elastic protective cuticles.
Protective_Cuticle_Layer emphasizes barrier and protective functions of the cuticle and is distinguished from Waxy_Cuticle_Layer which emphasizes wax deposition and bloom appearance. Protective cuticle represents the functional perspective on cuticle as primary defense against environmental stresses. Understanding protective cuticle layer properties is essential for assessing cultivar stress tolerance (drought, heat, pathogens), for managing berry protection through cultural practices enhancing cuticle development, and for post-harvest handling protocols maintaining cuticle integrity. The protective cuticle is the first line of defense against water loss and pathogen invasion.
Barriere-Schicht
Schutzüberzug
barrier layer
protective coating
schützende Cuticula-Schicht
Rogiers, S.Y. et al. (2004). Grape berry epicuticular wax and transpiration. American Journal of Enology and Viticulture 55=121-127 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 176-180 Becker, T., Knoche, M. (2011). Water movement through the surfaces of the grape berry and its stem. American Journal of Enology and Viticulture 62=340-350.
Protective Cuticle Layer
Chardonnay grafted on Teleki 5C rootstock showing protective lateral root cap on primary root growing through compacted clay loam, lateral cap 7 cell layers thick comprising 0.18 mm protective barrier, cell walls 1.2 micrometers thick providing mechanical strength, outermost 2 cap layers showing damage from soil abrasion but underlying layers intact protecting meristem, microscopic examination revealing structural integrity despite surface wear, root tip successfully penetrating dense soil demonstrating effective protective function of lateral cap mechanical shield
Protective lateral root cap is lateral root cap emphasizing mechanical protection and shielding function, characterized by structural features optimized for absorbing mechanical impacts and resisting abrasion including thick cell walls 0.5-2 micrometers providing strength, tough mechanical properties resisting deformation and damage, layered organization creating redundant barriers with multiple cell layers protecting meristem, and geometric arrangement distributing mechanical forces broadly preventing concentrated stress, with protective lateral root cap demonstrating defensive capacity including ability to sustain damage without compromising meristem function through expendable design, capacity to absorb soil particle impacts through cell deformation and wall compression, resistance to abrasive wear through robust structure, and rapid renewal replacing damaged cells within days maintaining protective barrier, and representing essential mechanical shield enabling root meristem to function in harsh soil environment where exposed meristem would be quickly damaged compromising root growth. Protective lateral root cap mechanical properties includes= showing wall strength (thick primary walls with high tensile strength resisting mechanical stress), exhibiting cell toughness (lateral cap cells resisting rupture and deformation under compressive and shear forces), demonstrating resilience (lateral cap recovering shape after deformation dissipating mechanical energy), revealing fracture resistance (thick walls and cell turgor preventing catastrophic failure under impact), and displaying wear resistance (lateral cap surface resisting abrasive damage from soil particle friction). Protective lateral root cap layered defense includes= showing multiple barriers (3-8 cell layers creating redundant protection with damage to outer layers not compromising inner layers), exhibiting progressive protection (each layer providing independent protective capacity with cumulative effect enhancing security), demonstrating sacrificial layers (outer layers absorbing damage protecting inner layers and underlying meristem), revealing structural hierarchy (different layers potentially showing different mechanical properties optimizing overall protection), and displaying depth of protection (multiple layers creating substantial barrier 60-200 micrometers thick protecting meristem). Protective lateral root cap force distribution includes= showing broad area distribution (lateral cap geometry spreading point forces from soil particles over large area), exhibiting stress dissipation (cap structure converting concentrated impacts into distributed stresses reducing peak forces on meristem), demonstrating load transfer (mechanical forces transferred through cap layers toward base where attached to root cylinder), revealing geometric optimization (cap curvature and thickness distribution optimizing force distribution patterns), and displaying structural integration (lateral cap functioning as integrated structure rather than independent cells distributing forces efficiently). Protective lateral root cap expendable design includes= showing damage tolerance (lateral cap capable of sustaining cell damage and death without compromising meristem), exhibiting replacement capacity (damaged lateral cap cells continuously replaced by new cells from cap initials maintaining protection), demonstrating cost-effectiveness (expendable design allowing protection through renewable barrier rather than requiring invulnerable structure), revealing strategic sacrifice (lateral cap cells programmed for eventual death creating protection through acceptable losses), and displaying sustainable defense (continuous production and loss creating steady-state protection maintained indefinitely). Protective lateral root cap in soil penetration includes= providing penetration shield (lateral cap protecting meristem during forceful advancement through soil), exhibiting impact absorption (lateral cap cells absorbing kinetic energy from collisions with soil particles or obstacles), demonstrating abrasion resistance (lateral cap surface resisting wear from friction during soil passage), revealing compression resistance (lateral cap resisting compressive forces from soil as root pushes forward), and displaying deformation capacity (lateral cap cells deforming to accommodate irregular soil surfaces without damage). Protective lateral root cap damage response includes= showing damage detection (lateral cap potentially sensing damage through membrane disruption or loss of turgor), exhibiting repair mechanisms (damaged lateral cap potentially undergoing localized repair through enhanced production), demonstrating accelerated replacement (damage potentially stimulating increased cell production replacing damaged tissue faster), revealing compensation (lateral cap potentially increasing thickness or strength in response to severe mechanical stress), and displaying recovery capacity (damaged cap recovering function rapidly through continuous renewal). Protective lateral root cap versus other protective structures includes= contrasting with permanent barriers (lateral cap renewable unlike bark or other permanent protective tissues), showing specialized location (lateral cap protecting growth-critical meristem versus other protective tissues covering mature organs), exhibiting unique renewal (lateral cap continuous replacement versus episodic renewal in other tissues), demonstrating expendable strategy (lateral cap designed for damage and loss versus other tissues minimizing damage), and revealing functional specialization (lateral cap optimized for mechanical protection during growth versus other tissues providing protection for mature structures). Protective lateral root cap functional necessity includes= enabling meristem survival (meristem requires lateral cap protection to survive in soil environment), demonstrating growth requirement (functional lateral cap necessary for normal root elongation and soil penetration), revealing vulnerability consequences (lateral cap damage or absence exposing meristem to potentially lethal mechanical damage), showing evolutionary importance (lateral cap ubiquitous in roots reflecting strong selection for mechanical protection), and displaying management relevance (soil conditions affecting lateral cap damage rates influencing root growth and vine establishment).
Protective_Lateral_Root_Cap emphasizes mechanical shielding function and defensive capacity and is distinguished from other lateral cap aspects by focus on protection mechanism. Critical for understanding mechanical properties, layered defense, force distribution, expendable design, impact absorption, damage tolerance, and the mechanical protection function enabling meristem survival and function in harsh soil environment.
Abschirmende Haube
Defensive lateral cap
Mechanical barrier cap
Mechanische Barrieregewebe
Schützende laterale Wurzelhaube
Shielding cap tissue
Protective Lateral Root Cap
Cabernet Sauvignon grafted on 1103P rootstock at 20 years showing protective outer rootstock bark 4.8 mm thickness providing multiple defense functions, waterproof suberized cork preventing trunk desiccation (moisture loss through bark estimated <5% of unprotected trunk), physical barrier blocking pathogen entry (no trunk diseases detected despite pathogen presence in vineyard), thermal insulation protecting cambium (bark temperature measurements showing 8-12°C moderation of daily fluctuations compared to ambient), mechanical protection evident from minimal damage despite equipment contact, chemical analysis revealing phenolic extractives in cork providing antimicrobial properties
Protective outer rootstock bark is outer rootstock bark emphasizing protective function and role in trunk defense, characterized by suberized cork creating waterproof barrier preventing desiccation, dead tissues providing physical barrier blocking pathogen entry, cork low thermal conductivity insulating cambium from temperature extremes, tough structure resisting mechanical damage from abrasion and impact, and chemical deterrents in suberin and extractives creating hostile environment for decay organisms, serving as essential protective layer maintaining trunk integrity enabling continued vascular cambium function and vine survival under environmental stress, and demonstrating that despite being completely dead tissue, protective bark provides critical defense functions through physical structure and chemical composition accumulated during cell differentiation prior to death. Protective outer rootstock bark waterproofing function includes= preventing water loss through transpiration (suberized cork cells creating impermeable barrier eliminating moisture loss from trunk surface estimated preventing 50-95% potential water loss compared to unprotected surface), maintaining trunk hydration (waterproof bark ensuring water transported through xylem reaches canopy rather than evaporating from trunk), enabling drought tolerance (bark protection critical during water stress preventing additional desiccation stress on vine), showing cultivar variation (some rootstocks with particularly effective waterproof bark showing superior drought tolerance), and demonstrating essential role (experiments removing bark causing rapid desiccation and death confirming protective importance). Protective outer rootstock bark pathogen barrier includes= blocking fungal penetration (cork physical barrier preventing fungal hyphae from reaching cambium and vascular tissues), preventing bacterial entry (dead cork layers creating barrier blocking bacterial invasion into living bark and wood), providing chemical deterrence (suberin and cork extractives creating antimicrobial environment inhibiting pathogen growth), showing vulnerability only when breached (intact bark providing excellent protection but wounds creating entry points bypassing barrier), and demonstrating disease resistance contribution (effective protective bark contributing to overall trunk disease resistance reducing infection frequency). Protective outer rootstock bark temperature insulation includes= protecting cambium from freeze damage (cork low thermal conductivity buffering rapid temperature changes reducing freeze injury risk to sensitive cambium cells), preventing heat damage (bark insulating cambium from excessive sun exposure and high temperatures preventing heat stress or sunburn injury), moderating daily temperature fluctuations (bark dampening diurnal temperature cycles maintaining more stable cambium temperature), showing thickness correlation (thicker bark providing better insulation protecting cambium more effectively from temperature extremes), and enabling climate adaptation (protective bark critical for trunk survival in climates with severe cold winters or hot summers). Protective outer rootstock bark mechanical protection includes= resisting abrasion damage (tough bark protecting trunk from friction damage from equipment, trellising, or wind-blown debris), preventing impact injury (bark cushioning trunk from mechanical impacts reducing bruising, cracking, or cambium damage), blocking animal damage (bark deterring or protecting from browsing, gnawing, or insect attack), showing cultivar differences (some rootstocks with particularly tough bark showing better mechanical protection), and requiring maintenance (damaged bark losing protective capacity necessitating wound care and management). Protective outer rootstock bark chemical defense includes= containing antimicrobial compounds (suberin and cork extractives including phenolics and terpenes inhibiting microbial growth), providing decay resistance (chemical compounds in bark creating unfavorable environment for wood-rotting fungi), showing cultivar variation (rootstock varieties differing in bark extractive content and decay resistance), demonstrating passive effectiveness (chemical defenses present without requiring active synthesis or metabolic maintenance), and requiring initial investment (compounds deposited during cork cell differentiation before death providing long-lasting protection). Protective outer rootstock bark in environmental stress includes= enabling survival in challenging conditions (protective bark critical for trunk function under drought, temperature extremes, or pathogen pressure), reducing stress impacts (effective bark protection buffering environmental stress reducing overall vine stress level), showing damage vulnerability (bark damage or disease compromising protection exacerbating environmental stress effects), demonstrating cultivar suitability (rootstock varieties with effective protective bark better adapted to specific environmental challenges), and requiring management attention (maintaining bark integrity through avoiding mechanical damage and managing diseases critical for maximizing protective function).
Protective_Outer_Rootstock_Bark emphasizes defensive function and is distinguished from Dead_Outer_Rootstock_Bark (death emphasis) by focus on protection roles. Critical for understanding waterproofing, pathogen barriers, temperature insulation, mechanical defense, chemical deterrence, and the essential protective functions of dead outer bark enabling trunk survival and vine productivity.
Barrier bark layer
Barriere-Rindenschicht
Defensive Borke
Defensive outer bark
Protective rhytidome
Schützende äußere Unterlagenrinde
Protective Outer Rootstock Bark
Cabernet Sauvignon protective outer trunk bark at 14 years post-planting shows 5.5-7.8 mm depth of suberized cork layers providing= waterproofing (preventing -3°C), UV shielding (absorbing >95% of 280-320 nm UV-B radiation), and enabling trunk survival through decades of environmental challenges
Protective outer trunk bark is outer trunk bark emphasizing its defensive and insulating functions, comprising suberized impermeable cork layers that prevent water loss, exclude pathogens, buffer temperature extremes, and shield living tissues from mechanical damage and environmental stress, representing the functional barrier safeguarding trunk vitality. Protective outer trunk bark structural basis of protection includes= suberized cell walls (suberin polymer deposition creating waxy hydrophobic barrier impermeable to water and most solutes), multiple cork layers (successive periderms providing redundant protection if outer layers damaged), continuous coverage (complete circumferential bark sheath without gaps), sufficient thickness (2-8 mm depth in mature vines providing adequate barrier), and intact surface (unbroken outer bark preventing pathogen entry and desiccation). Protective outer trunk bark waterproofing function includes= preventing outward water loss (suberized cork preventing desiccation of underlying living phloem and cambium maintaining tissue hydration), blocking inward water entry (during rain or irrigation preventing excessive water uptake that could compromise bark integrity), maintaining trunk water balance (outer bark impermeability enabling vascular tissues to regulate internal water status independent of external moisture), critical during dormancy (winter and summer drought periods when outer bark prevents catastrophic desiccation of dormant tissues), and lenticel-mediated gas exchange (porous regions enabling respiration while maintaining overall impermeability). Protective outer trunk bark pathogen exclusion includes= physical barrier (intact cork preventing fungal spores and bacteria from reaching living susceptible tissues), chemical resistance (suberin and associated waxes inhibiting microbial growth on bark surface), lack of nutrients (dead cork providing no food source for pathogens limiting colonization), exclusion of trunk disease pathogens (outer bark preventing Eutypa, Esca, Botryosphaeria canker fungi from infecting vascular cambium and xylem), and wound vulnerability (breaks in outer bark through pruning, mechanical damage, or cracks providing entry points emphasizing importance of maintaining bark integrity). Protective outer trunk bark mechanical protection includes= impact absorption (compressible cork cushioning blows from equipment, pruning shears, hail, falling debris), abrasion resistance (tough outer bark withstanding rubbing from trellis wires, ties, or wind-blown materials), compression tolerance (cork cells collapsing under pressure without rupturing protecting underlying living tissues), flexibility (allowing trunk bending without bark tearing under moderate wind stress), and buffering biological damage (deterring or mitigating damage from insect boring, animal browsing, bird pecking). Protective outer trunk bark thermal insulation includes= buffering daily temperature fluctuations (outer bark reducing amplitude of temperature cycles experienced by cambium from 20-30°C external range to 8-15°C at cambium), protecting against summer heat (thick bark preventing 35-40°C air temperatures from causing >30°C cambium temperatures preventing heat damage), insulating during winter cold (outer bark moderating freeze rates and reducing minimum temperatures at cambium by 3-8°C compared to exposed tissue), preventing freeze-thaw injury (insulation reducing frequency and severity of freeze-thaw cycles damaging cambial cells), and protecting against sunscald (southwestern sun exposure in winter potentially lethal to thin-barked trunks, adequate outer bark preventing damage). Protective outer trunk bark UV radiation shielding includes= absorbing damaging UV wavelengths (outer bark preventing 280-320 nm UV-B radiation from reaching living tissues), lignin and phenolic UV absorption (compounds in bark cell walls absorbing radiation preventing damage to DNA and proteins in living cells), preventing photo-oxidative damage (outer bark shielding reducing reactive oxygen species formation in living tissues), and enabling sun exposure tolerance (thick outer bark allowing trunks to tolerate direct intense sunlight without sunburn or cell death). Protective outer trunk bark fire resistance includes= low thermal conductivity (air-filled cork cells providing insulation reducing heat transfer to living tissues), ablative protection (outer cork layers burning or charring while insulating inner layers), moisture retention (even dead cork containing some moisture absorbing heat during evaporation), enabling survival of brief fire exposure (thick outer bark allowing trunk to survive grass fires, brief burning of prunings, or flame weeding without cambium death), and limits of protection (sustained intense fire overwhelming outer bark protection killing cambium and causing trunk death). Protective outer trunk bark thickness and protection correlation includes= thin bark (2-4 mm) providing basic protection adequate in benign conditions but vulnerable to severe stress, moderate bark (5-8 mm) offering good protection against typical environmental challenges and minor mechanical damage, thick bark (8-15 mm) providing excellent protection enabling survival of severe temperature extremes, major mechanical damage, and substantial pathogen pressure, and protection thresholds (minimum 3-4 mm outer bark generally necessary for adequate winter cold protection in temperate climates, 5-6 mm beneficial for freeze protection in marginal regions). Protective outer trunk bark vulnerability to damage includes= mechanical wounds breaching protection (pruning cuts, equipment gouges, animal browsing removing outer bark exposing living tissues), sun exposure damage (intense solar radiation potentially killing cambium on southwest trunk exposure particularly in young thin-barked trunks), freeze injury (inadequate bark allowing deep cold penetration killing cambium and inner bark), pathogen entry through cracks (fissures in outer bark potentially allowing disease organism access), and importance of damage prevention (maintaining intact outer bark critical as dead tissue cannot heal requiring underlying living tissue to seal breaches). Protective outer trunk bark cultivar variation in protection includes= thick bark cultivars providing superior protection (enhanced frost tolerance, better mechanical damage resistance, improved pathogen exclusion), thin bark cultivars showing greater vulnerability (requiring careful management to prevent sunscald, freeze damage, mechanical injury), and site matching (thin bark varieties suited to mild climates with minimal mechanical stress, thick bark varieties preferred in harsh environments or mechanized vineyards). Protective outer trunk bark seasonal protection dynamics includes= critical winter protection (outer bark preventing freeze damage to cambium during dormant season), summer protection importance (outer bark preventing desiccation and heat stress during drought periods), protection during active growth (outer bark safeguarding cambium during vulnerable active division periods), and dormancy protection (outer bark maintaining tissue viability through winter dormancy enabling spring reactivation). Protective outer trunk bark significance in trunk longevity includes= enabling decades of survival (protective outer bark allowing trunks to persist 30-80+ years despite environmental challenges), protecting perennial meristems (outer bark safeguarding vascular cambium and cork cambium enabling continuous growth), buffering stress impacts (outer bark reducing severity of freeze, drought, heat, and mechanical stress extending functional lifespan), and determining trunk replacement needs (severe bark damage potentially requiring trunk renewal or vine replanting if protection compromised beyond recovery capacity). Protective outer trunk bark management considerations includes= avoiding unnecessary bark damage (careful pruning, equipment operation, and canopy management minimizing bark injury), protecting young thin bark (trunk wrapping, shade structures, or paint protecting vulnerable young trunks until bark develops), monitoring bark condition (identifying cracks, wounds, or disease symptoms requiring intervention), and understanding protection limitations (recognizing that even thick bark has limits requiring appropriate cultural practices minimizing severe stress).
Protective_Outer_Trunk_Bark emphasizes defensive functions and is distinguished from Outer_Trunk_Bark (anatomical structure), Dead_Outer_Trunk_Bark (cellular status), and Peeling_Outer_Trunk_Bark (sloughing behavior) by focus on barrier and insulation roles. Critical for understanding trunk protection mechanisms, environmental stress tolerance, pathogen exclusion, temperature buffering, bark thickness requirements, and management practices maintaining bark integrity for long-term trunk survival.
Barrier bark
Barriere-Rinde
Defensive bark
Insulating bark
Isolierende Borke
Schützende Außenrinde
Protective Outer Trunk Bark
Syrah protective periderm at 19 years post-planting shows 2.8-4.5 mm thick cork layer providing= waterproofing (-4°C preventing freeze injury), UV shielding (absorbing >96% of damaging 280-320 nm radiation), gas exchange through 6-9 lenticels per cm² (enabling 0.4-0.8 μmol O₂/g bark/hour respiration supporting inner bark metabolism), and enabling trunk survival through decades of environmental stress
Protective periderm is periderm emphasizing its defensive functions including waterproofing, pathogen exclusion, mechanical cushioning, and thermal insulation, representing the functional barrier that safeguards living inner bark and cambium from environmental stress and damage enabling trunk survival and longevity. Protective periderm waterproofing function includes= preventing outward water loss (suberized cork creating impermeable barrier reducing transpiration from inner bark and cambium to 30°C cambium temperatures that would damage or kill cells), winter cold insulation (cork moderating freeze rates and reducing minimum temperatures at cambium by 3-8°C compared to exposed tissue preventing freeze injury), preventing freeze-thaw damage (insulation reducing frequency and severity of freeze-thaw cycles that could disrupt cambial cells and inner bark), and sunscald prevention (southwestern sun exposure in winter potentially lethal to thin-barked trunks, adequate cork layer preventing cambium death from radiant heating and subsequent freezing). Protective periderm UV radiation shielding includes= absorbing damaging wavelengths (cork layers blocking 280-320 nm UV-B radiation preventing DNA and protein damage in living cells), lignin and phenolic compounds (aromatic components in suberin and extractives absorbing UV preventing transmission to sensitive tissues), preventing photo-oxidative stress (cork shield reducing reactive oxygen species formation in living cambium and phloem from UV exposure), and enabling sun tolerance (thick protective periderm allowing trunks to withstand direct intense sunlight without sunburn or cell death). Protective periderm gas exchange accommodation includes= lenticels interrupting impermeability (specialized porous cork regions enabling oxygen diffusion inward and carbon dioxide diffusion outward), maintaining aerobic respiration (lenticel gas exchange supporting living inner bark and cambium metabolic activity requiring oxygen), balancing impermeability and permeability (lenticels comprising 95% waterproof coverage), seasonal lenticel dynamics (lenticels expanding during growing season facilitating increased respiration, partially closing during dormancy reducing water loss), and critical function (without lenticels, impermeable cork would suffocate living tissues, inadequate lenticels causing oxygen limitation and reduced vigor). Protective periderm thickness requirements for adequate protection includes= minimum threshold (2-3 mm total periderm typically necessary for basic protection in benign conditions), moderate protection (4-6 mm providing good resistance to typical environmental challenges and minor mechanical stress), enhanced protection (6-10 mm offering excellent defense against severe temperature extremes, substantial mechanical damage, and major pathogen pressure), cultivar variation (thick-cork cultivars naturally producing adequate protection, thin-cork cultivars potentially vulnerable in harsh conditions), and site matching (thin periderm adequate in mild climates, thick periderm essential in cold climates, mechanized vineyards, or high disease pressure sites). Protective periderm vulnerability and damage includes= mechanical wounds breaching protection (pruning cuts, equipment gouges, animal browsing removing cork exposing living inner bark to desiccation, pathogen entry, and freeze damage), sun exposure injury (intense southwestern winter sun potentially killing cambium through cork on thin-barked young trunks despite insulation), freeze damage (inadequate cork thickness allowing deep cold penetration killing cambium and phloem in severe winters), pathogen penetration through cracks (longitudinal fissures in cork potentially providing entry routes for disease organisms if extending through to living tissues), and importance of integrity maintenance (maintaining unbroken cork barrier critical as breaches potentially leading to localized tissue death, canker development, or catastrophic girdling if extensive). Protective periderm successive formation maintaining protection includes= outer periderm aging and damage (oldest exterior cork layers weathering, cracking, and losing effectiveness over 3-8 years), new deeper periderm formation (cork cambium arising in inner bark beneath aging outer periderm maintaining protection), continuous coverage (new periderm forming before outer periderm completely fails ensuring uninterrupted barrier), progressive interior addition (new periderms adding internally while old periderms deteriorate externally maintaining stable total protective thickness), and enabling trunk expansion (successive periderm formation allowing diameter increase while preserving protective covering as outer cork becomes non-expandable and sloughs). Protective periderm seasonal protection dynamics includes= critical winter protection (cork preventing freeze damage to cambium during dormant season when tissues most vulnerable), summer protection (cork preventing desiccation and heat stress during drought periods when water deficits threaten tissue viability), protection during active growth (cork safeguarding cambium during vulnerable division periods when cells particularly susceptible to stress), and dormancy protection (cork maintaining tissue viability through winter enabling spring reactivation when temperatures rise and growth resumes). Protective periderm cultivar differences in protection includes= thick periderm cultivars (superior protection, enhanced frost tolerance in marginal climates, better mechanical damage resistance in mechanized vineyards, improved pathogen exclusion), thin periderm cultivars (adequate protection in mild climates but requiring careful management, vulnerable to sunscald, freeze injury, and mechanical damage in harsh conditions), and management matching (thin-bark varieties requiring trunk protection measures in exposed sites, thick-bark varieties thriving in challenging environments). Protective periderm significance in trunk longevity includes= enabling decades of survival (protective cork allowing trunks to persist 30-80+ years despite environmental challenges), protecting perennial meristems (cork safeguarding vascular cambium and cork cambium enabling continuous growth and bark renewal), buffering stress impacts (cork reducing severity of freeze, drought, heat, and mechanical stress extending functional lifespan), determining replacement timing (severe cork damage potentially requiring trunk renewal or vine replanting if protection compromised beyond recovery), and foundation for old vine value (thick protective cork enabling vine longevity supporting old vine designations and associated quality characteristics).
Protective_Periderm emphasizes defensive functions and is distinguished from Periderm (anatomical structure), Cork_Layer_Periderm (phellem emphasis), and outer bark (multiple periderms) by focus on barrier and insulation roles. Critical for understanding trunk protection mechanisms, waterproofing and temperature buffering, pathogen exclusion, mechanical damage resistance, thickness requirements for adequate protection, and successive periderm renewal maintaining protection enabling trunk longevity and requiring management practices preserving cork integrity.
Barrier periderm
Barriere-Periderm
Defensive periderm
Insulating periderm
Isolierendes Korkgewebe
Schützendes Periderm
Protective Periderm
Chardonnay grafted on Teleki 5C rootstock at 20 years showing protective rootstock periderm providing multiple defense functions, waterproof cork measured preventing >96% trunk water loss compared to debarked control, pathogen barrier demonstrated by absence of trunk diseases despite pathogen presence in vineyard, thermal monitoring showing cork insulating cambium with 10°C daily temperature moderation, mechanical protection evident from minimal trunk damage despite equipment contact, lenticels enabling adequate gas exchange confirmed by normal tissue respiration rates, protective periderm essential for trunk survival and vine productivity
Protective rootstock periderm is rootstock periderm emphasizing defensive function and role in trunk protection, characterized by integrated three-component system where suberized cork provides waterproof barrier eliminating trunk water loss, dead cork layers create physical barrier blocking pathogen penetration, cork low thermal conductivity insulates cambium from freeze and heat extremes, cork mechanical properties resist damage from abrasion and impact, lenticels enable gas exchange maintaining respiration behind waterproof barrier, phelloderm contributes additional protective layer and potential chemical defense, and cork cambium maintains capacity to produce new protective tissues replacing damaged or aged cork, demonstrating that despite periderm's relatively small volume compared to wood or phloem, protective periderm performs essential trunk defense functions enabling vascular cambium function, vine survival under environmental stress, and continued secondary growth throughout trunk life. Protective rootstock periderm waterproofing function includes= eliminating trunk water loss (suberized cork creating impermeable barrier preventing transpiration from bark surface estimated >95% reduction compared to unprotected trunk), maintaining vascular function (waterproofing ensuring water transported through xylem reaches canopy rather than evaporating from trunk), enabling drought survival (periderm protection critical during water stress preventing additional desiccation), showing effectiveness testing (removing periderm causing rapid desiccation confirming protective importance), and demonstrating cultivar variation (some rootstocks with particularly effective periderm waterproofing showing superior drought tolerance). Protective rootstock periderm pathogen barrier function includes= blocking fungal penetration physically (cork layers preventing hyphal penetration to cambium and vascular tissues), preventing bacterial invasion (dead cork creating barrier blocking bacterial entry to living bark), providing chemical deterrence (suberin and cork extractives creating antimicrobial environment), showing breach vulnerability (intact periderm providing excellent protection but wounds bypassing barrier creating entry points), and contributing to disease resistance (effective periderm reducing trunk disease incidence in field conditions). Protective rootstock periderm thermal insulation function includes= protecting from freeze damage (cork low conductivity buffering rapid temperature drops reducing cambium freeze injury), preventing heat damage (periderm insulating cambium from sun exposure and high air temperatures), moderating diurnal fluctuations (periderm dampening daily temperature cycles maintaining more stable cambium temperature), showing thickness correlation (thicker periderm providing better insulation documented through temperature monitoring), and enabling climate adaptation (protective periderm critical for survival in climates with severe winters or hot summers). Protective rootstock periderm mechanical protection function includes= resisting abrasion (cork protecting trunk from friction damage from equipment, wires, or debris), cushioning impacts (cork compressibility absorbing mechanical shocks reducing bruising or cambium damage), deterring animal damage (tough periderm protecting from browsing, gnawing, or bark stripping), showing elastic properties (cork recovering from compression maintaining protective function after deformation), and demonstrating cultivar differences (some rootstocks with particularly robust periderm showing better mechanical protection). Protective rootstock periderm gas exchange maintenance includes= enabling oxygen diffusion through lenticels (loosely arranged cork cells in lenticels allowing O2 entry necessary for tissue respiration), permitting CO2 release (lenticels enabling CO2 produced by respiration to escape preventing toxic accumulation), maintaining aerobic metabolism (lenticel function ensuring adequate oxygen for living bark and cambium aerobic respiration), showing lenticel density correlation (adequate lenticel number and size critical for maintaining respiratory gas exchange), and demonstrating essential function (blocking lenticels experimentally causing tissue damage confirming necessity). Protective rootstock periderm integrated system function includes= coordinating multiple protections (periderm simultaneously providing waterproofing, pathogen barrier, thermal insulation, and mechanical defense), showing component synergy (cork providing most protection while phelloderm and cambium enabling renewal and adaptation), demonstrating redundancy (multiple periderm layers providing backup protection if outer layer damaged), enabling self-maintenance (cork cambium capacity to produce new tissues allowing periderm repair and replacement), and providing adaptable protection (periderm adjusting thickness and characteristics in response to environmental conditions and stress).
Protective_Rootstock_Periderm emphasizes defensive functions and is distinguished from Rootstock_Periderm (structural emphasis) by focus on protection roles. Critical for understanding waterproofing, pathogen barriers, thermal insulation, mechanical defense, gas exchange maintenance, and the essential protective functions of periderm system enabling trunk survival under environmental stress and pathogen pressure.
Barrier periderm system
Defensives Periderm-System
Schützendes Unterlagen-Periderm
Protective Rootstock Periderm
Cabernet Sauvignon protective seed coat at maturity (BBCH 89) provides mechanical protection through 200-250 μm lignified layers resisting 20-25 N crushing force, chemical protection through 11-14% condensed tannins with antimicrobial and antioxidant properties, physiological protection through controlled imbibition rate 1-2 mg water/seed/hour preventing tissue damage, and temporal protection through dormancy requiring 100-110 day stratification
A protective seed coat is a seed coat emphasizing its multifaceted protective functions including mechanical protection of embryo and endosperm, regulation of water and gas exchange, chemical defense through phenolic compounds, and control of germination timing, representing the functional integration of structural, chemical, and physiological protective mechanisms. Protective functions of the seed coat operate across multiple domains= physical protection (mechanical barrier against crushing, abrasion, and penetration damage from 150-300 μm thick lignified layers), chemical protection (antimicrobial and antioxidant properties of seed coat tannins reducing pathogen invasion and oxidative damage), physiological protection (controlled water uptake preventing tissue damage from rapid imbibition, gas exchange regulation maintaining appropriate embryo oxygen levels), and temporal protection (dormancy mechanisms preventing germination during unfavorable conditions, timing germination for optimal establishment). Structural elements providing protection include= outer cuticle with epicuticular waxes (waterproofing, reducing microbial attachment), epidermis with lignified cell walls (surface barrier), phenolic-rich hypodermal layers (chemical deterrent, antioxidant protection), heavily lignified sclereid layers (mechanical strength, 60-80% of seed coat thickness), and inner integument remnants (additional barrier layer). Chemical protective components include= condensed tannins (proanthocyanidins) comprising 5-15% seed coat dry weight (astringent and antimicrobial properties, oxidative protection), phenolic acids and flavonoids (additional antioxidant capacity), cutin and suberin in outer layers (hydrophobic barrier), and potential presence of germination inhibitors (abscisic acid, phenolic compounds contributing to dormancy). Protective seed coat functions vary with developmental stage= developing seed coats (green, soft, BBCH 75-81) provide minimal protection emphasizing growth over defense, maturing seed coats (tan, hardening, BBCH 83-85) show intermediate protection with progressive lignification and tannin accumulation, and fully mature seed coats (brown to black, hard, BBCH 87-89) provide maximal protection with complete lignification and polymerized tannins. Environmental challenges that protective seed coat resists include= mechanical damage from berry compression in dense clusters or processing operations, pathogen invasion (fungi, bacteria seeking to colonize nutrient-rich embryo and endosperm), oxidative stress (reactive oxygen species potentially damaging embryo cellular components), desiccation damage (protecting embryo from excessive water loss), and premature germination (preventing imbibition and germination during berry development or unfavorable seasons). Protective seed coat compromise has significant consequences= mechanical damage allowing pathogen entry or excessive water uptake, chemical degradation reducing antimicrobial properties, premature softening allowing off-season germination, or developmental failure resulting in seed abortion.
Protective_Seed_Coat emphasizes integrated protective functions and is distinguished from Hard_Seed_Coat (mechanical properties) and Seed_Coat (complete structure) by focus on multifaceted defense mechanisms across physical, chemical, and physiological domains. Critical for understanding seed survival and viability maintenance, germination ecology and dormancy adaptive value, pathogen resistance mechanisms, and wine quality implications where seed coat tannins provide both protection during development and potential astringency if extracted during processing.
Barrier testa
Barriere-Samengewebe
Defensive seed coat
Schutz-Testa
Schützende Samenschale
Cadot, Y., Miñana-Castelló, M. T., & Chevalier, M. (2006). Anatomical, histological, and histochemical changes in grape seeds from Vitis vinifera L. cv Cabernet franc during fruit development. Journal of Agricultural and Food Chemistry, 54(24), 9206-9215. | Bewley, J. D., & Black, M. (1994). Seeds= Physiology of Development and Germination (2nd ed.). Plenum Press.
Protective Seed Coat
Protein content quality in Hunter Valley Semillon presents the stability challenge characteristic of this high-protein variety, with must protein concentrations of 150-250 mg/L requiring substantial bentonite fining (0.5-1.5 g/L) to achieve heat stability for bottle aging the trade-off between protein removal for stability and protein retention for texture leads some premium producers to accept calculated instability risk in wines destined for extended cellaring, where gradual protein precipitation over years produces minimal haze compared to the rapid denaturation from acute temperature abuse.
Protein_Content_Quality represents the chemical quality characterizing the concentration and composition of proteins in grape must, capturing the macromolecular nitrogen fraction that affects wine stability, mouthfeel, and processing requirements. Proteins in grape juice derive primarily from berry tissue (pathogenesis-related proteins, enzymes, structural proteins) with additional contributions from fungal proteins in botrytized fruit. Total protein concentration in grape must typically ranges from 15-250 mg/L, with substantial variation reflecting cultivar genetics, vine nitrogen status, fruit health, and extraction conditions during processing. The quality implications of protein content center on protein haze instability in white wines, where heat-labile proteins (primarily pathogenesis-related proteins) denature and aggregate during storage or temperature fluctuation, producing unacceptable turbidity in bottled wine. This instability drives the widespread use of bentonite fining to remove proteins prior to bottling, with the required bentonite dose correlating with protein content and composition. Protein content also affects wine texture and mouthfeel, with proteins contributing to viscosity and potentially to the perception of body and richness, effects that create tension between stability requirements (favoring protein removal) and textural quality (potentially favoring protein retention). The protein profile varies among cultivars, with some varieties (Sauvignon Blanc, Semillon) showing high pathogenesis-related protein content and correspondingly high haze potential, while others (Chardonnay) typically show lower protein levels. Protein analysis employs Bradford, Lowry, or BCA colorimetric methods for total protein, with SDS-PAGE or HPLC methods providing compositional information about specific protein classes.
Protein_Content_Quality addresses the macromolecular nitrogen fraction distinct from free amino acids and ammonium. Proteins do not contribute to YAN as they require proteolytic degradation before yeast can assimilate the constituent amino acids. The stability implications of protein content distinguish it from other nitrogen forms, making protein management a specific winemaking concern particularly for white wines. The class serves as parent for Pathogenesis_Related_Protein_Quality, the specific protein class most responsible for haze instability.
Gesamtproteingehalt
Proteingehalt
Traubenproteinqualität
grape protein quality
macromolecular nitrogen content
makromolekularer Stickstoffgehalt
protein concentration quality
total protein content
Waters EJ et al. (2005) Preventing Protein Haze in Bottled White Wine. Australian Journal of Grape and Wine Research 11(2): 215-225 Ferreira RB et al. (2002) The Wine Proteins. Trends in Food Science and Technology 13(7): 230-239
Protein Content Quality
Protein cryoprotection quality characterization in cold-acclimating Chardonnay identified the specific protective proteins accumulating during hardiness development and their relationship to freezing tolerance. Proteomic analysis of bud tissues at monthly intervals from October through January quantified the cold-induced protein changes underlying hardiness acquisition. Dehydrin proteins showed the most dramatic accumulation, with DHN1a increasing 18-fold and DHN1b increasing 12-fold from baseline to maximum acclimation, together representing 3.5% of total soluble protein at full hardiness. The temporal pattern of dehydrin accumulation preceded maximum hardiness achievement by 2-3 weeks, consistent with protein accumulation being required for subsequent hardiness expression. Additional cold-responsive proteins included a putative antifreeze protein showing 6-fold increase, a lipid transfer protein increasing 4-fold, and several heat shock proteins providing chaperone function with 3-5 fold increases. Correlation analysis between individual protein levels and LT50 showed dehydrin concentration explaining 82% of hardiness variation, stronger than any other single protein class. Immunolocalization studies revealed dehydrin accumulation concentrated in bud primordia and vascular tissues, consistent with protective function in the cold-sensitive meristematic regions that determine spring growth capacity. The protein cryoprotection quality assessment established dehydrin accumulation as a primary molecular determinant of cold hardiness and identified dehydrin expression level as a potential selection criterion for breeding programs targeting improved cold tolerance.
Protein cryoprotection quality is a cryoprotection quality that inheres in a grapevine organism and represents the capacity to synthesize and accumulate proteins that protect cellular structures during freezing stress, including dehydrins and late embryogenesis abundant (LEA) proteins that stabilize membranes and other proteins during dehydration, antifreeze proteins that modify ice crystal growth to prevent mechanical damage, and cold-regulated proteins that provide various protective functions specific to freezing tolerance. This quality addresses the protein component of cryoprotection that complements sugar accumulation, recognizing that specific proteins are essential for maximum freezing tolerance through protective functions that sugars alone cannot provide, including direct protein-protein interactions that prevent denaturation and specific modifications of ice crystal behavior that reduce mechanical damage to cells. The ontological significance of protein cryoprotection quality lies in the essential contribution of protective proteins to cold hardiness, with the accumulation of dehydrins and related proteins showing strong correlation with freezing tolerance and with genetic variation in protein cryoprotection capacity contributing to differences in maximum hardiness achievable among genotypes. The biological basis of protein cryoprotection encompasses the transcriptional activation of cold-responsive genes encoding protective proteins during acclimation, the translation and post-translational modification of these proteins to produce functional cryoprotectants, and the mechanisms by which the accumulated proteins provide protection including membrane binding by dehydrins, chaperone activity preventing protein aggregation, and ice crystal modification by antifreeze proteins. The dehydrin family represents the most abundant and well-characterized cold-protective proteins, with their highly hydrophilic structure and membrane association enabling stabilization of lipid bilayers during the dehydration stress that accompanies extracellular freezing. The temporal dynamics of protein cryoprotection show characteristic patterns with transcript accumulation beginning within hours of cold exposure, protein accumulation following with maximum levels achieved over weeks of sustained cold, and protein degradation during deacclimation reducing protection as temperatures warm.
Protein_Cryoprotection_Quality is_a Cryoprotection_Quality, representing the accumulation of protective proteins including dehydrins, LEA proteins, and antifreeze proteins. This class addresses the essential protein contribution to maximum freezing tolerance. Dehydrins show 10-20 fold accumulation during acclimation and strong correlation with hardiness (r² > 0.8). The membrane-binding function of dehydrins provides protection distinct from sugar mechanisms. Transcript accumulation (hours) precedes protein accumulation (weeks) during acclimation. Localization in bud primordia consistent with protection of critical meristematic tissues.
Dehydrin-Akkumulation
LEA protein cold protection
LEA-Protein-Kälteschutz
Protein-Kryoprotektion
Schutzprotein-Akkumulation
antifreeze protein function
dehydrin accumulation
protective protein accumulation
Protein Cryoprotection Quality
Provisional status for Touriga Nacional in the Bordeaux appellation documents the 2021 INAO experimental authorization — permitted for trial planting on a maximum of 10% of each property's vineyard area and for blending at a maximum of 5% of the final wine, subject to a 10-year evaluation period during which agronomic performance, wine quality contribution, and appellation stylistic compatibility will be formally assessed, with the possibility of permanent authorization, extension of the provisional period, or revocation depending on the evaluation outcomes. Provisional status for PIWI varieties (pilzwiderstandsfähige — fungus-resistant interspecific hybrids) in Austrian viticulture documents the progressive evaluation of Muscaris, Souvignier Gris, Donauriesling, and Donauveltliner — varieties bred at Klosterneuburg from Vitis vinifera × resistant species crosses, currently authorized for Landwein and Qualitätswein production following successful multi-year evaluation trials, but not yet eligible for DAC designation pending further assessment of wine quality consistency and regional typicity expression.
Provisional status is a classification status entity that documents the temporary or conditional regulatory classification of a grape cultivar that has been permitted for trial cultivation and experimental wine production but has not yet achieved full authorized or recommended status, encompassing experimental planting authorizations (limited-area permits for cultivar evaluation under commercial conditions), provisional catalogue entries (time-limited inclusion in the national variety catalogue pending completion of evaluation trials), climate adaptation trial designations (the emerging regulatory category for cultivars being evaluated for suitability under changing climatic conditions), and breeding program release candidates (new cultivar selections undergoing the pre-authorization evaluation required for commercial classification), specifying the conditions, limitations, duration, and evaluation criteria of the provisional designation.
Provisional status documentation captures the regulatory pipeline through which new or previously unclassified cultivars progress toward full authorization — a process that balances the need for thorough evaluation (ensuring that newly classified cultivars genuinely contribute to regional wine quality) against the urgency of timely access to cultivars that may offer solutions to pressing viticultural challenges (particularly climate adaptation and disease resistance reduction of pesticide dependence). The evaluation period for provisionally classified cultivars typically spans 5-15 years and involves multi-site trial plantings, comparative winemaking studies, sensory evaluation by expert panels, and assessment of consumer and market acceptance — a comprehensive evaluation process that generates the evidence base for permanent classification decisions. The climate change adaptation context has substantially expanded the provisional classification category in recent years — traditional European wine regions that previously had stable, slowly evolving cultivar portfolios are now actively evaluating heat-tolerant, drought-resistant, and early-ripening cultivars for potential permanent inclusion, creating an unprecedented wave of provisional authorizations across France (the INAO "variétés d'intérêt à fin d'adaptation" program), Germany (the Bundessortenamt evaluation of Mediterranean varieties), and Austria (Klosterneuburg's PIWI program and international variety trials). The PIWI (fungus-resistant) variety evaluation represents a particularly significant provisional classification domain — these interspecific hybrids offer dramatic reductions in fungicide application (80-90% fewer treatments) but face regulatory resistance in traditional appellations where Vitis vinifera purity is a fundamental quality principle, and consumer skepticism about hybrid-origin wines, creating a tension between sustainability objectives and cultural tradition that the provisional classification process must navigate. The regulatory consequences of provisional status are deliberately limited — provisionally classified cultivars may typically be used for table wine and regional wine production but are excluded from the most prestigious appellation designations, limiting the economic incentive for widespread commercial planting and ensuring that the cultivar must demonstrate quality before gaining full market access.
Versuchsstatus
bedingte Genehmigung
conditional approval
experimental status
experimentelle Zulassung
provisional classification
provisorische Klassifizierung
temporary classification
trial authorization
vorläufiger Status
INAO (2021) Note d'information — variétés d'intérêt à fin d'adaptation au changement climatique HBLA Klosterneuburg (2023) PIWI-Sorten für den österreichischen Weinbau European Parliament (2013) Regulation (EU) No 1308/2013, Article 81(2) (experimental authorization) Bundessortenamt (2023) Beschreibende Sortenliste Reben
Provisional Status
Syrah on 110 Richter showing proximal first-order lateral shallow roots, excavation revealing laterals originating 25-45 cm from trunk base at 15-28 cm depth, proximal laterals 3.2-4.6 cm diameter extending 40-70 cm horizontal distance, laterals showing 4-6 annual rings, producing second-order branches supporting local absorptive network, proximal first-order laterals demonstrating near-trunk soil exploration complementing more extensive distal lateral network
Proximal first order lateral shallow root is first-order lateral shallow root emphasizing proximal position close to trunk or parent root origin, typically originating 20-60 cm from trunk base, characterized by early emergence during root system development, relatively short horizontal extent 30-80 cm compared to distal laterals, proximity to zone of intensive cultivation or traffic, and serving local soil exploration function accessing resources near trunk while more distal laterals explore farther zones, with proximal first-order laterals showing characteristics including origin from primary root relatively near surface and trunk, horizontal or oblique orientation spreading outward from central zone, moderate diameter 2-5 cm, moderate secondary growth developing some woody tissue, production of second-order branches supporting local absorptive network, and vulnerability to surface disturbances including tillage, traffic compaction, or construction activities affecting near-trunk zone. Proximal first order lateral shallow root positional characteristics includes= showing near-trunk origin (proximal laterals originating 20-60 cm from trunk base relatively close compared to distal laterals 60-150+ cm), exhibiting shallow depth (proximal laterals often at relatively shallow depths 10-30 cm in upper topsoil), demonstrating central zone association (proximal laterals concentrated in zone immediately surrounding trunk), revealing accessibility (proximal laterals easily accessible through excavation or observation in near-trunk area), and displaying exposure (proximal laterals potentially exposed or near-surface particularly on slopes or eroded sites). Proximal first order lateral shallow root length and extent includes= showing limited reach (proximal laterals typically extending 30-80 cm from origin shorter than distal laterals 80-200+ cm), exhibiting local exploration (proximal laterals exploring soil volume immediately surrounding trunk), demonstrating overlap (proximal lateral exploration zones from neighboring vines potentially overlapping in narrow row spacing), revealing competition (proximal laterals potentially experiencing competition from neighboring vine roots in overlapping zones), and displaying termination (proximal lateral growth potentially terminating when encountering competing roots or unfavorable conditions). Proximal first order lateral shallow root development timing includes= showing early formation (proximal laterals often forming earlier than distal laterals during root system establishment), exhibiting developmental priority (young vines potentially emphasizing proximal lateral development establishing near-trunk absorption), demonstrating rapid function (proximal laterals quickly producing second-order branches and fine roots beginning absorption), revealing establishment role (proximal laterals important for young vine establishment enabling early resource capture), and displaying continued production (mature vines continuing to maintain and replace proximal laterals despite also having extensive distal network). Proximal first order lateral shallow root vulnerability includes= showing tillage exposure (proximal laterals highly vulnerable to cultivation damage given shallow depth and proximity to trunk), exhibiting traffic impact (proximal laterals susceptible to compaction from vehicle or foot traffic in near-trunk zone), demonstrating construction risk (vineyard operations or construction potentially damaging proximal laterals during soil disturbance), revealing erosion exposure (soil erosion potentially exposing proximal laterals particularly on slopes), and displaying damage consequences (proximal lateral damage reducing near-trunk resource capture though vine may compensate through distal lateral function). Proximal first order lateral shallow root absorptive function includes= providing near-trunk absorption (proximal laterals and derivatives accessing water and nutrients in soil immediately surrounding trunk), exhibiting local resource capture (proximal laterals exploiting resources in central zone potentially including trunk-applied fertilizers or irrigation), demonstrating supplementary role (proximal laterals supplementing more extensive distal lateral absorption particularly in young vines), revealing competition effects (proximal lateral function potentially reduced in mature vineyards where root zones overlap creating intense competition), and displaying contribution (proximal laterals potentially providing 20-40% of total lateral root system absorption though highly variable). Proximal first order lateral shallow root management considerations includes= requiring tillage caution (cultivation near trunk risking proximal lateral damage suggesting reduced tillage or no-till in near-trunk zone), showing mulch benefits (trunk mulches protecting proximal laterals from disturbance and improving soil conditions), exhibiting irrigation relevance (drip emitters near trunk directly benefiting proximal laterals if placed appropriately), demonstrating fertilizer access (near-trunk fertilizer applications efficiently accessed by proximal laterals), and revealing protection needs (protecting proximal laterals from damage important particularly in young vines where they provide substantial absorption). Proximal first order lateral shallow root in young versus mature vines includes= showing young vine importance (proximal laterals providing large fraction of absorption in young vines before distal laterals extensively developed), exhibiting relative decline (proximal lateral proportional contribution declining as vine matures and distal laterals develop), demonstrating maintained presence (proximal laterals maintained in mature vines though relatively less important than extensive distal network), revealing age-related changes (proximal lateral characteristics potentially changing with vine age through continued secondary growth and branching), and displaying persistence (proximal laterals potentially persisting throughout vine life though some mortality and replacement occurring). Proximal first order lateral shallow root spatial efficiency includes= showing localized exploration (proximal laterals thoroughly exploring limited soil volume near trunk), exhibiting intensive branching (proximal laterals potentially producing dense second-order branching in local zone), demonstrating efficient resource use (proximal laterals providing absorption without requiring extensive lateral elongation), revealing architectural integration (proximal laterals integrating with distal laterals creating complete root system exploring soil from trunk outward), and displaying complementary function (proximal laterals complementing distal laterals together creating effective resource capture system).
Proximal_First_Order_Lateral_Shallow_Root emphasizes near-trunk proximal position and is distinguished from distal laterals by focus on central zone location. Critical for understanding near-trunk absorption, limited extent, tillage vulnerability, young vine importance, and the proximal laterals exploring soil immediately surrounding trunk complementing distal lateral network.
Central-zone first-order root
Near-trunk lateral root
Proximal lateral branch
Proximale erste Ordnung Lateralwurzel
Stamm-nahe Seitenwurzel
Zentral-Lateralast
Proximal First Order Lateral Shallow Root
Merlot proximal seed adjacent tissue (2nd-3rd cell layers from seeds, total 48-78 μm thickness) showing intermediate characteristics= 2nd layer cells 52-62 μm diameter, walls 1.3-1.5 μm, tannins 6.5-7.2 mg/g 3rd layer cells 58-68 μm, walls 1.1-1.4 μm, tannins 5.6-6.5 mg/g positioned 35-95 μm from seed surfaces in mature berries (week 18), showing systematic gradation in cell size (increasing), wall thickness (decreasing), and tannin content (decreasing) with distance from seeds creating smooth transition between immediate seed adjacent maximum and outer endocarp moderate levels. Cabernet Sauvignon proximal seed adjacent tissue tannin gradient demonstrating seed-proximity effect= immediate seed adjacent (innermost layer) 7.8 mg/g tannins (100% of maximum) 2nd layer (proximal inner) 6.9 mg/g (88% of maximum, positioned 38 μm from seeds) 3rd layer (proximal middle) 6.1 mg/g (78% of maximum, 68 μm from seeds) 4th layer (proximal outer/outer endocarp boundary) 5.2 mg/g (67% of maximum, 98 μm from seeds), demonstrating 33% tannin decline over 98 μm distance from immediate seed adjacent maximum through proximal seed adjacent zone creating quantifiable seed-proximity phenolic gradient. Syrah proximal seed adjacent tissue cell size progression showing gradual enlargement= immediate seed adjacent 45-52 μm diameter 2nd layer (proximal inner) 52-62 μm 3rd layer (proximal middle) 60-72 μm 4th layer (proximal outer) 68-82 μm 5th layer (outer endocarp) 78-92 μm, demonstrating systematic cell size increase over 5 layers spanning 140-180 μm distance creating continuous size gradient from innermost minimum to outer endocarp approaching middle mesocarp size without abrupt boundaries between zones. Pinot Noir proximal seed adjacent tissue extraction timing during extended maceration= days 1-8 extracting primarily outer pericarp and beginning middle mesocarp with minimal proximal seed adjacent extraction days 9-14 showing increasing proximal seed adjacent extraction as cells begin breaking down contributing 8-15% of wine skin tannins days 15-20 showing substantial proximal extraction contributing 15-25% of wine skin tannins as extended maceration accesses deeper pericarp layers, demonstrating proximal seed adjacent tissue as intermediate-difficulty extraction target accessible during extended maceration second week. Chardonnay proximal seed adjacent tissue in white cultivar showing graduated structure= 2nd-3rd layers from seeds, 52-82 μm thickness, cells 50-68 μm diameter showing size increase with distance from seeds, walls 1.1-1.4 μm showing thinning with distance, tannins 4.8-6.2 mg/g showing decline with distance creating gradient from innermost maximum (5.5-6.8 mg/g immediate seed adjacent) to outer endocarp moderate levels (3.5-4.8 mg/g), demonstrating proximal seed adjacent tissue gradients present in white cultivars similar to reds though absolute tannin levels lower reflecting cultivar differences in phenolic biosynthetic capacity.
A proximal seed adjacent tissue is seed coat adjacent tissue comprising the 2nd-3rd (sometimes extending to 4th) cell layers from seed surfaces, positioned 30-100 μm from seeds in mature berries, characterized by cells slightly larger than immediate seed adjacent tissue (50-65 μm diameter versus 40-55 μm), moderately thick walls (1.1-1.5 μm versus 1.3-1.8 μm in immediate tissue), and elevated tannin concentration (5.5-7.5 mg/g) lower than immediate tissue maximum (6.5-8.5 mg/g) but substantially higher than outer endocarp (3.5-5.5 mg/g), representing the seed-proximate zone showing declining seed-proximity effects with increasing distance from seed surfaces. Proximal seed adjacent tissue position characteristics= (1) comprising cell layers 2-3 or 2-4 from innermost pericarp boundary depending on total endocarp thickness and classification scheme (2) positioned 30-100 μm from seed surfaces in mature berries (after locular cavity air space development) or 20-90 μm during development (when seeds contact pericarp more closely) (3) experiencing intermediate seed proximity with weaker seed-derived signals or influences than immediate seed adjacent tissue but stronger than distant outer endocarp layers (4) creating transition zone between innermost extreme characteristics (immediate seed adjacent tissue) and more typical endocarp properties (outer endocarp layers). Proximal seed adjacent tissue showing graduated characteristics= (1) cell size intermediate between immediate seed adjacent minimum (40-55 μm) and outer endocarp approaching middle mesocarp (70-90 μm), with proximal tissue 50-65 μm representing gradual size increase (2) wall thickness intermediate between immediate seed adjacent maximum (1.3-1.8 μm) and outer endocarp (0.8-1.2 μm), with proximal tissue 1.1-1.5 μm showing gradual thinning (3) tannin concentration intermediate between immediate seed adjacent maximum (6.5-8.5 mg/g) and outer endocarp moderate levels (3.5-5.5 mg/g), with proximal tissue 5.5-7.5 mg/g showing gradual decline creating tannin gradient (4) all characteristics showing systematic gradation with distance from seeds creating smooth transition rather than abrupt boundary between innermost and outer endocarp zones. Proximal seed adjacent tissue tannin gradient analysis= innermost proximal layer (2nd layer from seeds, 30-50 μm distance) typically showing 6.5-7.5 mg/g tannins (12-20% below immediate seed adjacent maximum) middle proximal layer (3rd layer, 50-80 μm distance) showing 5.8-6.8 mg/g (25-35% below maximum) outermost proximal layer (4th layer, 80-110 μm distance) showing 5-6.2 mg/g (35-45% below maximum), demonstrating systematic tannin decline with distance from seeds creating exponential-like gradient potentially reflecting declining seed-proximity signal strength or position-dependent biosynthetic regulation. Proximal seed adjacent tissue developmental changes= early development showing proximal tissue cells closer to seeds (15-70 μm distance) when berry smaller and seeds fill more of available space expansion during ripening increasing distance from seeds as berry grows faster than seeds creating progressive separation mature berries showing proximal tissue separated from seeds by locular cavity air space plus immediate seed adjacent tissue creating total 40-110 μm distance from seeds late-season potential for slight cell shrinkage reducing total pericarp thickness and slightly reducing seed-proximal distance. Proximal seed adjacent tissue in wine extraction showing intermediate extraction difficulty= more accessible than immediate seed adjacent tissue due to slightly larger cells and thinner walls less accessible than outer endocarp and middle mesocarp due to smaller cells and thicker walls than bulk tissue typically extracting during days 12-18 of extended maceration after outer pericarp and middle mesocarp substantially extracted but before complete immediate seed adjacent tissue extraction contributing 10-18% of wine skin tannins during extended maceration protocols representing significant tannin reservoir accessed through extended maceration duration.
Proximal_Seed_Adjacent_Tissue represents the near-seed endocarp layers slightly more distant than immediate contact and is distinguished from Immediate_Seed_Adjacent_Tissue (innermost 1-2 layers in direct/very close contact) by slightly greater distance and graduated characteristics. Proximal seed adjacent tissue shows intermediate properties between innermost extreme and outer endocarp moderate levels creating transition zone. Understanding proximal seed adjacent tissue is relevant for detailed gradient characterization, for comprehending spatial extent of seed-proximity effects on pericarp structure and composition, and for winemaking extraction strategies where proximal tissue represents intermediate extraction difficulty accessed during extended maceration protocols.
Inneres Endokarp
inner endocarp
nahes Samenkontakt-Gewebe
near seed-contact tissue
proximales samenangrenzedes Gewebe
Proximal Seed Adjacent Tissue
The pruning method description for a Grand Cru Chablis vineyard specifies Guyot simple cane pruning performed manually between January 15 and March 1 (after the coldest period to minimize frost damage to pruning wounds), retaining one fruiting cane of 8 buds selected from the previous season's growth based on internode regularity, cane diameter of 8-10 mm, and position relative to the renewal spur, with one two-bud renewal spur retained on the opposite side of the trunk to ensure alternating cane direction in successive years — total retained bud load of 10 buds per vine at 10,000 vines/ha yielding 100,000 buds/ha, with pruning weight collected and weighed at 0.3-0.5 kg per vine (3,000-5,000 kg/ha) to verify vine balance via the Ravaz index targeting 5-8.
A viticultural practice description that documents the systematic removal of vine wood — canes, spurs, cordons, and non-structural shoots — performed during the dormant season or growing season to regulate vine form, bud number, crop potential, vegetative-reproductive balance, and canopy architecture, encompassing the complete set of decisions and techniques that constitute the single most influential annual viticultural management intervention.
Pruning_Method_Description captures the most consequential annual management decision in viticulture — the selection and retention of specific buds that will produce the following season's shoots, flowers, and fruit. Pruning determines bud number (and therefore potential shoot number and crop load), vine form maintenance (cordon integrity, trunk health, canopy geometry), and disease management (removal of infected wood, wound hygiene): an experienced pruner makes 30-50 individual cut decisions per vine, each affecting the vine's productivity and health for the current and subsequent seasons. The description encompasses both the general pruning system (spur versus cane, bilateral versus unilateral) and the specific implementation parameters (bud load, spur length, cane selection criteria, timing, and pruning weight targets) that translate the system concept into operational practice. Subclasses capture these key specification dimensions.
Rebschnitt-Dokumentation
Schnittmethoden-Beschreibung
Schnittpraxis-Beschreibung
pruning practice documentation
vine pruning record
winter pruning description
Galet P (2000) General Viticulture, Oenoplurimedia pp: 230-295 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 331-365 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 97-130
Pruning Method Description
The pruning system name cordon de Royat designates the spur-pruned bilateral cordon system developed in the 19th century at the agricultural school of Royat near Clermont-Ferrand, specifying a permanent bilateral cordon trained along a single fruiting wire with 5-8 evenly spaced two-bud spurs per cordon arm — this system predominates in warm-climate viticulture (Rhône Valley, southern France, Australia, California) where basal bud fruitfulness is high, while the contrasting pruning system name Guyot (simple or double) designates cane pruning preferred in cooler climates (Burgundy, Champagne, Mosel) where fruitful buds are located at positions 4-8 on the cane rather than at the basal positions retained in spur pruning.
A pruning method description consisting of the standardized or traditional name identifying the fundamental pruning approach applied to the vine, distinguishing primarily between spur pruning (retention of short spurs with 1-3 buds on a permanent cordon), cane pruning (annual selection and retention of one or more long fruiting canes with 6-15 buds), and head pruning (retention of short spurs on a free-standing head without trellis support), with the name serving as the primary reference key for the associated set of pruning protocols.
Pruning_System_Name provides the primary classificatory distinction within pruning methodology. The choice between spur and cane pruning is fundamentally determined by cultivar-specific bud fruitfulness patterns: cultivars with high basal bud fruitfulness (Cabernet Sauvignon, Syrah, Grenache, Merlot) are well-suited to spur pruning because the 1-2 basal buds retained on each spur produce fruitful shoots, while cultivars with low basal bud fruitfulness (Riesling, Sémillon, Sultanina, many table grapes) require cane pruning to access the fruitful buds at positions 5-10 on the cane. Some cultivars (Chardonnay, Pinot Noir) have intermediate fruitfulness patterns that permit either system, with the choice determined by regional tradition, target yield, and training system compatibility. Appellation regulations frequently specify the authorized pruning system — for example, the Châteauneuf-du-Pape cahier des charges mandates gobelet (head-pruned bush vine) or cordon de Royat.
Rebschnittform-Bezeichnung
Schnittart-Name
Schnittsystem-Name
pruning approach name
pruning type designation
vine pruning system designation
Galet P (2000) General Viticulture, Oenoplurimedia pp: 230-260 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 335-350 Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles pp: 25-60
Pruning System Name
The pruning timing specification for Riesling in the Mosel prescribes late dormant pruning between February 15 and March 15 — after the period of maximum winter cold risk (January-early February, when pruning wounds would expose internal tissues to freeze damage at -15 to -20°C) but before sap flow initiation (typically late March in the Mosel, when pruning cuts would cause excessive bleeding of 1-5 liters per vine per day from xylem sap exudation). In contrast, the pruning timing specification for Cabernet Sauvignon in frost-prone Coonawarra recommends delayed pruning in September (Southern Hemisphere spring) to retard budburst by 7-14 days, exploiting the observation that late-pruned vines break bud later and thus escape early spring frosts that damage earlier-pruned vines.
A pruning method description that documents the calendar period or phenological window within which pruning operations are performed, specifying the optimal timing relative to vine dormancy status, winter cold risk, spring frost vulnerability, sap flow initiation, and labor availability that determines the physiological impact of pruning on vine response in the subsequent growing season.
Pruning_Timing_Specification captures the temporal dimension of pruning that significantly influences vine physiology and frost risk management. The timing of dormant pruning relative to the vine's annual carbohydrate cycle affects reserve mobilization and subsequent-season vigor: early pruning (immediately after leaf fall) removes buds before full starch storage is complete, potentially reducing vine reserves, while late pruning (near budburst) delays bud development by 5-14 days — a phenomenon exploited as a frost avoidance strategy in frost-prone regions. The practice of double pruning (initial machine pre-pruning to long spurs in mid-winter followed by final manual pruning to target bud number in late winter) combines the labor efficiency of mechanical pruning with the precision of hand finishing. The relationship between pruning timing and bleeding (xylem sap exudation from cut surfaces) is a persistent concern in some regions though research suggests that bleeding, while visually dramatic, causes minimal vine damage.
Rebschnitt-Terminierung
Schnittfenster-Dokumentation
Schnittzeitpunkt-Spezifikation
pruning calendar description
pruning date range
pruning window specification
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 350-360 Friend AP and Trought MCT (2007) Delayed winter spur-pruning in New Zealand can alter yield components of Merlot grapevines, Australian Journal of Grape and Wine Research 13: 157-164 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 97-115
Pruning Timing Specification
Comparison of Pruning_Weight_Per_Meter_Quality across three Chardonnay training systems in Burgundy reveals equivalent vigor status despite dramatically different per-vine weights: traditional Guyot simple averaging 0.45 kg/m with 0.5 m cane length, bilateral cordon averaging 0.42 kg/m with 1.5 m total cordon, and high-wire minimally pruned plots averaging 0.48 kg/m with 2.0 m canopy extent, demonstrating that spatial normalization enables meaningful comparison across disparate training approaches.
Pruning_Weight_Per_Meter_Quality is a Vigor_Expression_Quality that inheres in a grapevine (Vitis vinifera L.) and represents the mass of one-year-old wood removed during dormant pruning expressed per linear meter of canopy or cordon length, providing a spatially normalized vigor assessment that enables comparison across varying vine spacings, training systems, and canopy configurations. This quality addresses a fundamental limitation of whole-vine pruning weight by accounting for the physical extent of canopy over which vegetative growth is distributed, recognizing that a large pruning weight concentrated in a short canopy section indicates different vigor status than the same weight distributed across a longer canopy extent. In BFO terms, Pruning_Weight_Per_Meter_Quality exists as a derived quality calculated from the ratio of mass to linear distance, with values enabling direct comparison across diverse viticultural systems that would be confounded by raw pruning weight comparisons. The viticultural significance of this normalization is substantial, as modern trellising systems vary widely in cordon length (from 0.5 m single Guyot canes to 4+ m bilateral cordons), vine spacing (0.8-3.0 m within rows), and canopy configuration (single vs. divided), making per-vine pruning weight comparisons misleading without spatial normalization. Target values for balanced wine production typically fall between 0.3-0.6 kg/m, with lower values indicating potential undercropping capacity and higher values suggesting the need for increased crop load, reduced irrigation, or vigor-reducing rootstock adoption. The relationship between pruning weight per meter and canopy density provides insight into the three-dimensional distribution of foliage, with high values indicating dense canopies potentially requiring shoot thinning or training system modification to improve light penetration and air circulation. This normalized measurement facilitates the transfer of research findings across regions and systems, enabling the application of vine balance principles developed in one context to viticultural situations with different spatial configurations.
Pruning_Weight_Per_Meter_Quality enables standardized vigor comparison across the diverse training systems and planting densities encountered in global viticulture. This normalization is essential for translating research findings and best practices across regions with different traditional vine configurations, supporting evidence-based viticultural decision-making independent of local conventions. The measurement requires specification of the linear dimension used (cordon length, row length, canopy extent) for unambiguous interpretation.
Schnittholzdichte
Schnittholzgewicht pro Meter
linear pruning weight
lineare Schnittholzmasse
normalized pruning weight
normiertes Schnittholzgewicht
pruning weight density
pruning weight per meter row
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 28-35 Archer E, Strauss HC (1985) Effect of plant density on root distribution of three-year-old grafted 99 Richter grapevines. South African Journal of Enology and Viticulture 6:25-30 Reynolds AG, Wardle DA, Naylor AP (1996) Impact of training system, vine spacing, and basal leaf removal on Riesling vine performance, berry composition, canopy microclimate, and vineyard labor requirements. American Journal of Enology and Viticulture 47:63-76
Pruning Weight Per Meter Quality
A balanced pruning program for Concord juice grapes in New York's Finger Lakes region adjusts node retention based on Pruning_Weight_Quality, following the formula of 30 nodes for the first pound (454 g) of prunings plus 10 additional nodes for each subsequent pound, such that a vine producing 1.8 kg of prunings would retain 30 + 10 + 10 + 10 = 60 nodes to maintain the yield-to-pruning-weight relationship that maximizes sustainable production and winter hardiness.
Pruning_Weight_Quality is a Vigor_Expression_Quality that inheres in a grapevine (Vitis vinifera L.) and represents the total mass of one-year-old wood (canes) removed during dormant-season pruning, providing the standard viticultural measure of seasonal vegetative biomass accumulation and vine vigor status. This quality serves as the foundation for vine balance assessment, balanced pruning systems, and the calculation of key viticultural indices relating vegetative and reproductive performance. In BFO terms, Pruning_Weight_Quality exists as a mass quality measurable through direct weighing of removed cane material, with values typically expressed per vine, per meter of cordon, or per unit land area depending on the analytical purpose. The physiological basis for pruning weight reflects the integrated carbon balance over the growing season, with photosynthetic carbon fixation partitioned among fruit production, root development, reserve accumulation, and cane biomass, such that pruning weight represents the residual vegetative investment after other demands are satisfied. Typical pruning weights for balanced wine grape production range from 0.3-0.6 kg per vine (or 0.3-0.5 kg per meter of row) for quality-focused production to 1.0-2.0 kg per vine for high-vigor table grape or raisin production, with excessive pruning weights indicating vegetative excess at the expense of fruit quality and insufficient weights suggesting stress or overcropping that compromises vine longevity. The Ravaz Index, calculated as yield divided by pruning weight, represents the most widely used vine balance indicator, with optimal values of 5-10 indicating appropriate balance between crop load and vegetative capacity for quality wine production. Pruning weight shows strong correlations with canopy dimensions, leaf area, and photosynthetic capacity, enabling its use as a surrogate measure for these more labor-intensive assessments. Spatial variation in pruning weight across vineyard blocks forms the basis for precision viticulture zone delineation, with vigor maps derived from remote sensing validated through pruning weight ground-truth measurements that establish the relationship between spectral indices and vine biomass production.
Pruning_Weight_Quality represents the canonical vigor assessment in viticulture, serving as the denominator for vine balance calculations (Ravaz Index) and the basis for balanced pruning systems. The measurement requires complete collection of pruned material, which can be labor-intensive for large-scale assessments, driving interest in proxy measures including cane diameter sampling, shoot counts, and remote sensing approaches. This class relates to vine balance, crop load management, and long-term vineyard sustainability through its integration of seasonal carbon partitioning outcomes.
Anschnittgewicht
Rebholzgewicht
Schnittholzgewicht
Schnittholzmasse
cane weight
dormant pruning weight
pruning mass
vine prunings
wood weight
Ravaz L (1911) L'effeuillage de la vigne. Annales de l'École Nationale d'Agriculture de Montpellier 11:216-244 Bravdo B, Hepner Y, Loinger C, Cohen S, Tabacman H (1984) Effect of crop level on growth, yield and wine quality of a high yielding Carignane vineyard. American Journal of Enology and Viticulture 35:247-252 Kliewer WM, Dokoozlian NK (2005) Leaf area/crop weight ratios of grapevines: influence on fruit composition and wine quality. American Journal of Enology and Viticulture 56:170-181
Pruning Weight Quality
The pruning weight target for a balanced Pinot Noir vineyard in the Côte de Beaune is 0.3-0.5 kg per vine (3,000-5,000 kg/ha at 10,000 vines/ha), yielding a Ravaz index of 5-7 at the target yield of 45-50 hl/ha — pruning weight below 0.2 kg/vine indicates an undervigorous vine requiring reduced bud load or corrective nutrition (particularly nitrogen), while pruning weight above 0.7 kg/vine indicates excessive vigor requiring increased bud load, reduced irrigation or nitrogen, or cover crop competition to redirect vine resources from vegetative growth to fruit ripening. The collection and weighing of pruning wood from representative sample vines (typically 20-50 vines per management block) provides the data for annual vine balance assessment and bud load calibration.
A pruning method description that documents the target mass of one-year-old wood removed during dormant pruning, expressed as kilograms per vine or kilograms per hectare, serving as a retrospective indicator of vine vegetative capacity that enables assessment of vine balance through the Ravaz index (yield-to-pruning-weight ratio) and informs bud load adjustment decisions for the following season.
Pruning_Weight_Target links the annual pruning decision to the vine's long-term productivity balance through the concept of vine capacity — the total amount of vegetative and reproductive growth the vine can support to maturity in a given environment. The Ravaz index (yield divided by pruning weight), proposed by Louis Ravaz in 1903 and refined through subsequent research, provides a dimensionless ratio that normalizes yield against vine capacity: values of 5-10 indicate balanced vines where the available photosynthetic and nutrient resources are appropriately partitioned between fruit ripening and structural growth, values above 10 indicate overcropping (fruit demands exceeding vine capacity, leading to incomplete ripening and depleted reserves), and values below 3 indicate excessive vegetative vigor (shading, poor fruit quality, wasted resources). Modern precision viticulture extends pruning weight assessment through spatially explicit pruning weight mapping using instrumented mechanical pre-pruners that record wood mass with GPS positioning, enabling vine-by-vine or zone-by-zone bud load optimization.
Rebholzmasse-Zielwert
Schnittholzgewicht-Spezifikation
Schnittholzgewicht-Vorgabe
dormant pruning mass target
pruning weight specification
vine capacity indicator
Ravaz L (1911) L'effeuillage de la vigne, Annales de l'École Nationale d'Agriculture de Montpellier 11: 216-244 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 365-380 Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles pp: 91-120 Bramley RGV et al. (2011) Variation in vine vigour, grape yield and vineyard soils and topography as indicators of variation in the chemical composition of grapes, wine and wine sensory attributes, Australian Journal of Grape and Wine Research 17: 217-229
Pruning Weight Target
Pubescent surface quality on Malbec shoot tips (OIV descriptor 004 rating 7-9, very dense prostrate hairs) creates the characteristic white-felted appearance that distinguishes young Malbec growth from the less pubescent Merlot and Cabernet Sauvignon with which it is commonly interplanted in Bordeaux and Argentina this dense pubescence persists on lower leaf surfaces throughout the season (OIV 053 rating 7), contributing to the distinctive silvery-white appearance of Malbec canopy undersides visible when wind lifts the leaves.
Pubescent_Surface_Quality represents the surface texture condition where the external boundary of a grapevine organ bears trichomes (plant hairs) that create a fuzzy, downy, woolly, or bristly tactile character. Pubescence is a prominent and variable surface texture feature in grapevine, particularly on young shoot tips, expanding leaves, and the lower (abaxial) surfaces of mature leaves, providing important ampelographic characters for cultivar identification. Trichome types in Vitis include simple unbranched hairs of varying length and density, and ribbon-like prostrate trichomes that create a woolly or cottony appearance. The degree of pubescence ranges from sparse scattered hairs through moderate coverage to dense woolly indumentum that completely obscures the underlying epidermis. Pubescence is typically most dense on young expanding tissues and decreases as organs mature and expand, diluting trichome density. The functional significance of pubescence includes boundary layer modification that reduces water loss and moderates leaf temperature, physical deterrence of small arthropod herbivores, and alteration of light reflectance affecting photosynthesis and photoprotection. OIV descriptors 004 (density of prostrate hairs on shoot tip) and 053 (density of prostrate hairs between veins on lower surface of blade) capture pubescence characters using categorical scales from absent through very dense. Pubescence assessment employs visual observation of hair density and type, tactile evaluation of surface feel, and microscopic examination for detailed trichome characterization.
Pubescent_Surface_Quality specifically addresses hair-covered surfaces and is complementary to Glabrous_Surface_Quality which describes hairless surfaces. Pubescence varies among organ types, surfaces, and developmental stages, requiring specification of the assessed surface for reproducibility. The trichome types (prostrate versus erect, simple versus branched) provide additional characterization beyond mere presence of pubescence. OIV descriptors employ separate scales for prostrate hairs (creating woolly texture) and erect hairs (creating bristly texture). Dense pubescence may interfere with spray penetration and affect pesticide efficacy.
Pubeszenz
behaarte Oberfläche
downy texture
filzige Textur
flaumige Oberflächenbeschaffenheit
hairy surface
tomentose quality
trichome-covered surface
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 47-54 Galet P (1979) A Practical Ampelography, Cornell University Press pp.: 95-110
Pubescent Surface Quality
Puckering quality assessment comparing wines from same vineyard with different extraction protocols reveals extraction-driven puckering differences, with extended maceration wine showing pronounced puckering (7/10) from seed tannin extraction creating the tight, gripping sensation, while shorter maceration wine displays minimal puckering (3/10) with predominantly skin-derived tannins creating drying without significant contraction, demonstrating how winemaking decisions affect puckering character and enabling style targeting through extraction management.
Puckering quality is a tactile astringency quality that represents the contractile, drawing sensation of oral tissues in response to astringent stimuli, characterizing the reflexive tightening of cheeks and lips that accompanies strong astringent perception and contributes to the gripping, mouth-coating character of tannic wines. Within the BFO framework, this quality specifically addresses the contractile dimension of astringent tactile perception, distinguishing puckering from drying while recognizing that puckering represents a distinct physiological response involving tissue contraction rather than simply reduced lubrication. The ontological significance lies in puckering providing qualitative information about astringency character beyond simple intensity, where the degree and nature of puckering sensation indicates tannin structural properties affecting how astringency is experienced, with certain tannin types creating pronounced puckering while others produce drying without significant contraction. The physiological basis for puckering involves both mechanical and neural components, with the reduced lubrication causing increased surface contact and friction, combined with possible direct effects of polyphenols on epithelial cells creating the sensation of tissue tightening, and the reflexive response of oral musculature contracting in response to the aversive stimulus. The compositional factors affecting puckering character include tannin source (seed tannins typically more puckering than skin tannins), tannin molecular size distribution (intermediate-sized tannins creating maximum puckering), specific phenolic composition (certain compounds more puckering than others at equivalent concentration), and the complex interactions between tannin structure and oral tissue response. The descriptive vocabulary for puckering ranges from absent (no contractile sensation) through mild (slight tightening at finish) to pronounced (obvious contraction of cheeks and lips) to severe (strongly aversive puckering dominating mouthfeel), with appropriate puckering level highly dependent on wine style and regional tradition. Understanding puckering quality enables astringency character assessment beyond intensity, tannin source inference from puckering nature, and quality evaluation incorporating appropriate puckering for wine type.
Puckering_Quality addresses the contractile dimension of tactile astringency distinct from drying. This quality provides information about tannin character and source, with seed tannins typically more puckering than skin tannins. The class supports astringency character assessment and winemaking evaluation through the specific puckering component that indicates tannin structural properties.
Mundverengung
Zusammenziehen
adstringierender Zug
astringent grip
contractile sensation
drawing tightness
kontrahierendes Gefühl
mouth puckering
Puckering Quality
Pulp firmness quality in Autumn Crisp table grapes reaches exceptional levels (4.5-6.0 N puncture force at 3mm depth into pulp) that define this variety's intensely crunchy texture, maintained through the extended California harvest season (August-November) with minimal softening the crisp pulp character distinguishes Autumn Crisp from traditional varieties like Thompson Seedless (pulp firmness 1.5-2.5 N) and commands premium pricing in markets valuing the distinctive textural experience that firm pulp provides.
Pulp_Firmness_Quality represents the physical quality characterizing the mechanical resistance of the internal flesh (mesocarp) tissue of grape berries, capturing the textural property most directly determining eating quality perception of crispness or softness. The pulp comprises the bulk of berry volume and mass, consisting of large, thin-walled parenchyma cells containing vacuoles filled with juice (water, sugars, acids). Pulp firmness reflects the turgor pressure of these cells, the mechanical strength of their thin primary cell walls, and the pectin-mediated adhesion between adjacent cells. During eating, the pulp provides the immediate textural sensation as teeth penetrate the berry after the initial skin resistance, with firm, turgid pulp producing the crisp, crunchy texture valued in premium table grapes while soft, degraded pulp creates the mushy texture associated with overripe or deteriorated fruit. Pulp firmness decreases during ripening through cell wall degradation (pectin solubilization, hemicellulose modification) and turgor changes, with the rate of softening varying substantially among cultivars. Crisp-textured varieties (Autumn Crisp, Sweet Celebration) maintain high pulp firmness through extended ripening, while soft-textured varieties show progressive firmness loss. Pulp firmness measurement requires either puncture testing that penetrates through skin into pulp (with appropriate depth analysis to isolate pulp response) or testing of excised pulp tissue freed from skin constraint. The force-distance profile during puncture through skin-pulp-seed layers can be analyzed to extract pulp-specific parameters, though complete tissue separation provides clearer measurement. Pulp firmness correlates with juice release behavior during crushing, affecting processing efficiency and must characteristics.
Pulp_Firmness_Quality specifically addresses the internal flesh tissue distinct from skin firmness, recognizing the different contributions of these tissues to overall texture and quality. The measurement challenge of isolating pulp properties from skin contribution requires either careful depth-specific analysis of penetration curves or physical tissue separation before testing. Pulp firmness most directly correlates with eating texture perception, making it particularly relevant for table grape quality assessment. The cellular basis of pulp firmness (turgor plus cell wall plus adhesion) provides targets for genetic improvement and storage optimization.
Fruchtfleischfestigkeit
Mesokarp-Textur
Pulpahärte
flesh firmness
innere Gewebefestigkeit
internal tissue firmness
mesocarp texture
pulp hardness
Sato A and Yamada M (2003) Berry Texture of Table, Wine, and Dual-Purpose Grape Cultivars Quantified. HortScience 38(4): 578-581 Rolle L et al. (2012) Instrumental Texture Analysis Parameters as Markers of Table-Grape and Winegrape Quality: A Review. American Journal of Enology and Viticulture 63(1): 11-28
Pulp Firmness Quality
Pulp maturity assessment comparing harvest timing options in Chardonnay reveals the structural and compositional evolution affecting winemaking decisions, with early harvest (22 Brix) showing firm pulp (resistant to crushing, lower free-run yield of 55%, clean juice separation), optimal harvest (24 Brix) showing softened pulp (easy crushing, 68% free-run yield, efficient extraction), and extended hang time (26 Brix) showing very soft pulp (releasing juice easily but with increased solids, potential oxidation from membrane breakdown), demonstrating how pulp maturity affects both the quantity and quality of extractable juice with implications for wine style and processing efficiency.
Pulp maturity quality is a physiological maturity quality that represents the developmental state of berry mesocarp tissue, characterizing the structural and biochemical changes in the flesh that surrounds seeds and underlies skin, including cell wall softening, vacuolar changes, and the accumulation of sugars, acids, and flavor precursors that constitute the bulk of grape juice and provide the primary fermentation substrate. Within the BFO framework, this quality specifically addresses the pulp component of physiological maturity, distinguishing mesocarp development from seed or skin maturation while recognizing that the pulp provides both the volume of fermentable material and the textural characteristics affecting juice release, extraction efficiency, and processing behavior during winemaking. The ontological significance lies in pulp maturity determining the yield and composition of extractable juice, where pulp cell wall breakdown facilitates juice release during pressing, vacuolar integrity affects the purity of released juice, and the accumulated composition of pulp cells provides the sugars for fermentation, the acids for pH balance, and many of the flavor precursors that contribute to wine character. The structural changes during pulp maturation include cell wall pectin solubilization (softening texture), middle lamella breakdown (reducing cell-to-cell adhesion), and changes in cell membrane permeability (affecting solute compartmentalization), with these changes producing the textural softening that accompanies ripening and determines juice extraction behavior. The compositional changes during pulp maturation involve continued sugar accumulation (phloem unloading into pulp cells), organic acid metabolism (malic acid respiration, tartaric acid stability), and flavor precursor development (aroma compound synthesis and glycosylation), with the pulp representing the primary reservoir for these compounds that define must composition. The assessment of pulp maturity involves evaluation of berry texture (firmness to touch, ease of crushing), juice release characteristics (free-run yield, pressing behavior), and the visual appearance of exposed pulp (color, translucency), providing practical indicators complementing compositional analysis. Understanding pulp maturity quality enables assessment of extraction potential, prediction of processing behavior, and interpretation of the textural changes that accompany compositional ripening in the primary juice-bearing tissue.
Pulp_Maturity_Quality addresses the developmental state of berry mesocarp determining juice yield and extraction characteristics. This quality reflects structural and compositional changes affecting processing behavior. The class supports harvest and processing decisions through pulp maturity characterization.
Beerenfleischentwicklung
Fruchtfleischreife
Mesokarp-Reife
Pulpareife
berry flesh development
flesh maturity
mesocarp ripeness
pulp ripeness
Nunan KJ et al. (1998) Changes in cell wall composition during ripening of grape berries. Plant Physiology 118(3): 783-792 Ortega-Regules A et al. (2006) A first approach towards the relationship between grape skin cell-wall composition and anthocyanin extractability. Analytica Chimica Acta 563(1-2): 26-32
Pulp Maturity Quality
Pulp texture quality in Muscat of Alexandria undergoes characteristic transformation during ripening, with pre-véraison berries displaying firm, crisp pulp (penetration force 2.5-3.5 N) that transitions through intermediate softening (1.5-2.0 N at véraison) to the soft, melting texture (0.8-1.2 N) at harvest maturity that enables juice release during gentle pressing while retaining sufficient structure for careful handling over-mature fruit shows pulp degradation (force below 0.5 N) with cellular breakdown creating watery, textureless flesh that produces oxidation-prone must and indicates suboptimal harvest timing.
Pulp_Texture_Quality represents the internal texture quality specifically characterizing the parenchymatous flesh tissue (mesocarp) of grape berries that constitutes the bulk of berry volume and provides the juice for winemaking or the edible portion for fresh consumption. Pulp texture reflects the structural organization and mechanical properties of the mesocarp cells, which are large, thin-walled, highly vacuolated parenchyma cells filled with juice containing sugars, acids, and other solutes. The texture of berry pulp varies from firm and crisp in certain table grape cultivars to soft and melting in most wine grape varieties, with these differences reflecting variation in cell wall rigidity, pectin composition, and intercellular adhesion strength. Crisp pulp texture results from cells that fracture cleanly under pressure, releasing juice while maintaining structural fragments, whereas soft or melting texture results from cells that separate along the middle lamella and deform plastically. Pulp texture changes profoundly during ripening as cell wall-modifying enzymes including polygalacturonase, pectin methylesterase, and expansins alter wall structure and reduce intercellular adhesion. The rate and extent of softening determine the harvest window for table grapes, where excessive softening renders fruit unmarketable, and influence extraction efficiency for wine grapes, where softer pulp releases juice more readily during crushing. Pulp texture assessment employs sensory evaluation of flesh firmness and crispness during chewing, instrumental penetrometry measuring the force required to push a probe through the pulp, and texture profile analysis characterizing multiple mechanical parameters.
Pulp_Texture_Quality specifically addresses the mesocarp tissue that constitutes the berry flesh, distinct from skin texture and seed coat texture. The pulp is the source of juice in winemaking and the primary edible tissue in table grapes. Pulp texture assessment typically involves removing or penetrating the skin to access the underlying flesh. The relationship between pulp texture and juice extractability makes this quality relevant to processing efficiency as well as sensory properties. Cultivar-specific pulp texture profiles (crisp versus melting versus firm-melting) represent important breeding targets for different market segments.
Beerenfleischqualität
Fruchtfleischtextur
Mesokarpstruktur
Pulpabeschaffenheit
fruit flesh quality
pulp consistency
Nunan KJ et al. (1998) Changes in Cell Wall Composition during Ripening of Grape Berries. Plant Physiology 118(3): 783-792 Sato A et al. (2006) Comparison of Berry Texture among Grapevine Cultivars. Journal of the Japanese Society for Horticultural Science 75(1): 11-17
Pulp Texture Quality
Cabernet Sauvignon pulp tissue at maturity comprises 78-82% of berry fresh weight (excluding seeds and skin), consisting of large parenchyma cells averaging 140-165 μm diameter in central region with thin 0.5-0.7 μm walls and vacuoles occupying 92-94% of cell volume, containing 24-26% sugars (12.5-13.5% glucose, 11.5-12.5% fructose), 0.6-0.75% organic acids (tartaric 0.38-0.48%, malic 0.18-0.28%), 78-80% water, 0.4-0.8 mg/g tannins (minimal phenolics), creating soft sweet colorless flesh providing 74-76 L juice per 100 kg berries and determining base wine composition (sugar/alcohol level, acidity) before skin/seed phenolic extraction. Chardonnay white wine pulp tissue showing white wine characteristics= 80-84% of berry fresh weight, cells 145-175 μm diameter, 21-24% sugars, 0.7-0.95% acids (higher acidity than typical red cultivars important for white wine freshness), 76-79% water, yielding 73-77 L juice per 100 kg berries, with pulp tissue juice composition determining complete must composition for white wine production as white winemaking separates juice from skins/seeds immediately after pressing avoiding phenolic extraction. Syrah pulp tissue cell expansion during ripening= veraison (BBCH 81, week 10) cells 72-88 μm diameter, vacuoles 88-91% of volume mid-ripening (week 14) cells 115-145 μm, vacuoles 91-93% maturity (week 18) cells 145-175 μm, vacuoles 92-95%, demonstrating 101-133% cell diameter increase (310-570% volume increase) driving berry enlargement, with pulp tissue cell expansion accounting for 85-92% of total berry volume increase during Phase II ripening. Thompson Seedless table grape pulp tissue showing large-berry characteristics= cells 155-195 μm diameter (large cells creating large berry size 20-24 mm diameter, 5-7 g per berry), 18-21% sugars (moderate for table grapes balancing sweetness with fresh flavor), 4.5-6.5 g/L acidity, 79-82% water, firmness 2-2.8 N (moderately firm maintaining texture during shipping), demonstrating pulp tissue properties determining table grape eating quality (sweetness, juiciness, texture) and shipping durability. Merlot pulp tissue composition gradients showing spatial uniformity= outer pulp (adjacent to outer mesocarp) with 24.2% sugars, 6.4 g/L acids central pulp with 24.8% sugars, 6.2 g/L acids inner pulp (adjacent to inner mesocarp) with 24.5% sugars, 6.5 g/L acids, demonstrating minimal compositional gradients (all within ±3% variation) despite substantial cell size gradients (outer 95-125 μm, central 145-165 μm, inner 105-140 μm), suggesting sugar and acid distribution equilibrates across pulp tissue creating uniform composition for juice/wine production.
A pulp tissue is the soft fleshy parenchymatous tissue comprising the bulk of the berry mesocarp, essentially synonymous with middle mesocarp or mesocarp bulk, typically representing 75-85% of total pericarp volume and 78-88% of berry fresh weight, consisting of large thin-walled parenchyma cells (80-180 μm diameter) with extensive vacuoles (90-95% of cell volume) accumulating sugars, water, and acids, creating the soft juicy edible portion of the berry that provides juice volume and fermentable sugars while containing minimal phenolic compounds. Pulp tissue cellular characteristics= (1) large parenchyma cells representing maximum cell size in berry tissue with central mesocarp cells 110-180 μm diameter at maturity (cultivar-dependent, some table cultivars showing cells up to 200 μm) (2) thin flexible cell walls 0.4-0.8 μm composed primarily of cellulose, hemicellulose, and pectin creating soft texture and allowing dramatic cell expansion during ripening (3) large central vacuoles occupying 90-95% of mature cell volume serving as primary storage compartment for water, sugars, acids, and minerals (4) thin peripheral cytoplasm layer 1-3 μm containing nucleus, plastids, mitochondria with metabolic activity focused on transport and storage (5) minimal intercellular spaces 1-4% of tissue volume providing limited gas diffusion pathways. Pulp tissue composition at maturity= (1) high water content 75-82% creating juicy texture and providing juice yield of 70-78 L per 100 kg berries during pressing (2) high sugar content 18-28% with glucose and fructose in approximately 1=1 ratio accumulated in vacuoles through phloem import during ripening (3) organic acids 0.5-0.9% fresh weight (tartaric acid 0.3-0.6%, malic acid 0.1-0.4%) providing acidity and pH buffering capacity (4) minerals including potassium (1200-2200 mg/kg), calcium (60-150 mg/kg), magnesium (60-120 mg/kg) important for yeast nutrition and wine chemistry (5) minimal phenolic compounds with tannins 0.3-1 mg/g, essentially no anthocyanins (<0.05 mg/g in red cultivars, <0.02 mg/g in whites), creating colorless tissue despite intense skin pigmentation in red/black cultivars. Pulp tissue textural properties= (1) soft texture when cells turgid, yielding readily to compression with firmness typically 1-2.5 N measured by penetrometry (2) juicy texture when cells disrupted releasing aqueous cellular contents (3) dramatic softening during ripening from firm green berry state (4-6 N) to soft ripe state (1-2.5 N) through cell wall modifications and turgor changes (4) potential textural defects including mealiness (excessive cell separation creating dry grainy texture) or mushiness (excessive wall breakdown creating formless texture) when ripening or storage conditions suboptimal. Pulp tissue development= Phase I (BBCH 71-75, weeks 0-6) involving cell division creating pulp cell population (1.2-2.8 million cells representing majority of berry cells) with cells 30-70 μm diameter lag phase (BBCH 77, weeks 6-8) with minimal growth Phase II (BBCH 81-89, weeks 9-18+) involving dramatic cell expansion with cells enlarging from 60-90 μm to 110-180 μm diameter (120-200% diameter increase, 350-700% volume increase per cell) through vacuolar expansion and water accumulation, creating bulk of berry enlargement. Pulp tissue functions= (1) primary storage organ accumulating photosynthates imported via phloem supporting berry growth and composition development (2) juice source providing essentially all grape juice volume for wine, juice, or fresh consumption (3) sugar reservoir providing fermentable sugars for alcoholic fermentation (4) acid reservoir affecting wine pH, titratable acidity, and sensory properties (5) water reservoir maintaining berry hydration and turgor.
Pulp_Tissue is essentially synonymous with Middle_Mesocarp or mesocarp bulk and represents the commercial flesh tissue providing juice and sugars. Sub-classification into Soft, Liquid, and Storage pulp tissue reflects different functional aspects (texture, juice content, storage function). Understanding pulp tissue is fundamental for viticulture and enology as this tissue determines juice yield, sugar levels, acid balance, and base wine composition, representing the primary commercial product component for both wine and table grape production.
Beerenfruchtfleisch
Fruchtmark
Pulpa-Gewebe
flesh tissue
fruit pulp
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 180-190 Coombe, B.G., McCarthy, M.G. (2000). Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6=131-135.
Pulp Tissue
Pulp weight data for Merlot from Pomerol (2024 vintage) records mean pulp weight of 1.05 g (76.1% of 1.38 g total berry weight), with the relatively large pulp fraction producing a generous juice yield of 72% (w/w) but moderate phenolic concentration in the free-run must, informing the winemaker's decision for extended maceration (18-21 days) to maximize skin phenolic extraction into the substantial juice volume. Pulp weight data for Shiraz from McLaren Vale (2023 vintage) documents pulp weight of 0.68 g (71.6% of 0.95 g berry weight) from dry-grown old vines, with the reduced pulp fraction producing proportionally more concentrated must (24.5° Brix, anthocyanins 2.4 mg/g) than irrigated material of the same cultivar with pulp weight of 1.02 g (75.6% of 1.35 g berry weight, 23.8° Brix, anthocyanins 1.5 mg/g). Pulp weight data for Grüner Veltliner from Langenlois (2024 vintage) records 1.22 g (80.3% of 1.52 g berry weight), the high pulp proportion characteristic of white wine varieties where juice yield maximization is a primary processing objective.
Pulp weight data is a berry composition data entity that records the mass of the grape berry mesocarp (pulp/flesh) following removal of the exocarp (skin) and seeds, determined gravimetrically, expressed in grams per berry (g/berry) and as a percentage of total berry weight, representing the tissue fraction that constitutes the primary reservoir of water, fermentable sugars (glucose and fructose), organic acids (tartaric and malic), and the bulk liquid phase that forms grape must upon crushing, and whose volume relative to skin and seed mass fundamentally determines the dilution factor for quality-determining compounds during juice expression and maceration.
Pulp weight data is the volumetric counterpart to skin and seed weight, providing the denominator in the extraction ratio that determines wine concentration — all other factors being equal, berries with less pulp relative to skin produce more concentrated wines because phenolic and aromatic compounds from the skin are dissolved into a smaller liquid volume. The grape berry mesocarp consists of large, thin-walled parenchyma cells that expand dramatically during post-véraison ripening through osmotic water influx driven by sugar accumulation, making pulp weight the most environmentally responsive component of berry composition. Water deficit reduces pulp weight through restricted cell expansion (without substantially affecting skin or seed mass), creating the well-documented mechanism by which regulated deficit irrigation enhances wine concentration. The pulp tissue is not entirely devoid of quality-relevant compounds — hydroxycinnamic acids (caftaric acid, coutaric acid) are present in the pulp and contribute to must browning susceptibility, while some cultivar-specific flavor compounds (such as furaneol/strawberry furanone in certain Muscat and hybrid varieties) may localize in the mesocarp. The juice yield from pulp depends on pressing conditions — gentle whole-cluster pressing extracts primarily vacuolar juice with lower phenolic content, while progressive pressing pressure ruptures cell walls and extracts additional juice enriched in cell wall-associated compounds. For white wine production, pulp weight relative to total berry weight directly determines the maximum juice yield achievable (typically 65-80% of berry weight as expressed juice), while for red wine production, pulp weight determines the liquid volume available for extracting skin and seed components during maceration.
Fruchtfleischanteil
Fruchtfleischgewicht
Mesokarpgewicht
Pulpamasse
Saftfraktionsgewicht
flesh weight
juice fraction weight
mesocarp weight
pulp content data
pulp mass
Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 265-300 Roby G et al. (2004) Berry size and vine water deficits as factors in winegrape composition. Australian Journal of Grape and Wine Research 10(2): 100-107 Coombe BG (1992) Research on development and ripening of the grape berry. American Journal of Enology and Viticulture 43(1): 101-110
Pulp Weight Data
Puncture resistance quality assessment in Cabernet Sauvignon guides winemaking extraction decisions, with skin break force of 0.8-1.2 N (2mm probe) indicating the tough skin structure that requires extended maceration (14-21 days) for optimal color and tannin extraction, compared to Pinot Noir at 0.3-0.5 N where equivalent extraction occurs in 7-12 days winemakers interpreting puncture resistance alongside phenolic ripeness data optimize maceration protocols to match the mechanical properties of each lot, maximizing extraction efficiency while avoiding the over-extraction that extended contact with already-depleted skins can produce.
Puncture_Resistance_Quality represents the physical quality characterizing the resistance of grape tissue to penetration by a probe or point, capturing the firmness dimension relevant to skin integrity assessment, damage from pointed objects, and extraction behavior during crushing. Puncture testing applies force through a small-diameter probe (typically 2-3 mm for berry testing) that concentrates stress at the contact point, generating a force-distance curve as the probe penetrates through sequential tissue layers (skin, pulp, potentially encountering seeds). The puncture response provides more detailed information about tissue structure than compression testing, as the force curve shows distinct features corresponding to skin break, pulp resistance, and seed contact. The initial peak in the puncture curve corresponds to skin break force, the parameter most directly indicating skin mechanical strength independent of internal tissues. Subsequent features reflect pulp firmness as the probe travels through the flesh. Puncture resistance decreases during ripening as both skin strength and pulp firmness decline, with cultivar-specific softening patterns affecting harvest quality. The probe diameter substantially affects results, with smaller probes providing greater discrimination of skin properties while larger probes integrate more pulp response. Test speed affects the viscoelastic response of tissue, requiring standardization for comparable results. Puncture resistance correlates with susceptibility to damage from pointed objects during harvest and handling, and with skin behavior during crushing where seeds and stem fragments create puncture stresses. The skin break force from puncture testing provides the most specific assessment of skin mechanical properties available from whole-berry testing.
Puncture_Resistance_Quality addresses the penetration loading mode that provides tissue-specific information complementing compression testing. The skin break force extracted from puncture curves specifically characterizes skin mechanical properties, making puncture testing particularly valuable for skin integrity assessment. The progressive penetration through tissue layers enables depth-specific analysis distinguishing skin from pulp contribution. The class describes resistance to pointed penetration generally, including during crushing where skin puncture initiates juice release.
Durchstoßwiderstand
Penetrationswiderstand
Punkturfestigkeit
Schalendurchbruchkraft
penetration resistance
probe penetration resistance
puncture firmness
skin break force quality
Letaief H et al. (2008) Assessment of Grape Skin Hardness by a Puncture Test. Journal of the Science of Food and Agriculture 88(9): 1567-1575 Río Segade S et al. (2008) Grape Skin and Seed Hardness Related to the Phenolic Composition. Journal of the Science of Food and Agriculture 88(14): 2452-2458
Puncture Resistance Quality
Purple hue development during Syrah véraison follows a characteristic spatial pattern, with berries on the exposed cluster face transitioning from green through pink to deep purple (CIELAB h° 320-350°) over 7-10 days while shaded interior berries lag by 3-5 days and achieve less intense purple coloration this véraison heterogeneity creates the mottled purple and green appearance of mid-véraison Syrah clusters that gradually homogenizes toward uniform dark purple-black as ripening progresses, with final harvest coloration (h° 350-10°) indicating the red-purple characteristic of ripe Syrah destined for age-worthy wines.
Purple_Hue_Quality represents the chromatic character of grapevine tissues displaying coloration intermediate between red and blue in the visible spectrum, corresponding to wavelengths in the violet region (approximately 380-450 nm) combined with red wavelengths, and CIELAB hue angles of approximately 300-360° (the red-blue transitional region). Purple hue results from anthocyanin pigments present in forms that reflect both red and blue wavelengths, creating the characteristic violet, magenta, and purple colorations associated with young red wines and recently ripened dark grape cultivars. The purple appearance indicates a mixture of anthocyanin forms or a balance between red-reflecting and blue-reflecting pigment states. In grape berries, purple hue characterizes the intermediate ripening stages of colored cultivars during véraison when anthocyanin accumulation produces visible coloration but has not yet reached the intense dark levels that produce blue-black appearance. At harvest maturity, many cultivars retain purple hue in their skins, particularly in the berry shoulder region with greater sun exposure. Purple hue in young wines is associated with monomeric anthocyanins in their native forms, with the progressive shift from purple toward red-brick hues during aging reflecting anthocyanin polymerization and structural changes. Cultivar differences in purple hue expression relate to anthocyanin profile, with delphinidin-rich cultivars displaying stronger blue-purple character and cyanidin-rich cultivars showing red-purple tones. Cool climate conditions and moderate crop loads promote purple hue retention at harvest by maintaining anthocyanin stability and preventing heat-induced degradation.
Purple_Hue_Quality represents the transitional region between red and blue hues, characterized by simultaneous reflection of both red and blue wavelengths. The distinction between purple and red or purple and blue hues is gradational categorical assignment reflects the perceived balance between red and blue color components. Purple coloration is particularly associated with young wines and recently ripened fruit, serving as an indicator of anthocyanin freshness. OIV descriptors for berry skin color include blue-black (category 6) which encompasses intensely colored purple-black fruit. The term violet in wine color description typically indicates a bluer purple than magenta, which has stronger red character.
Magentafärbung
Purpurton
blue-red hue quality
magenta coloration
purple color
purpurne Tönung
violet hue
violette Färbung
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed, John Wiley and Sons pp.: 141-203 OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 83-87
Purple Hue Quality
The breeding program at Hochschule Geisenheim University systematically develops Pyramided_Rpv_Resistance_Disposition genotypes through marker-assisted crossing and selection. A strategic cross between a Rpv1+Rpv3-1 carrier (from Muscadinia introgression line × 'Regent') and a Rpv10+Rpv12 carrier (from V. amurensis-derived material) produces a segregating population where approximately 6.25% (1/16) of offspring inherit all four resistance loci. These elite selections are identified through multiplex genotyping of linked markers and confirmed through phenotypic evaluation with diverse P. viticola isolates including characterized virulent races. Four-gene pyramid genotypes exhibit complete immunity across all tested isolates, with no visible symptoms or microscopic pathogen development even using isolates that partially overcome Rpv3-1 alone. Field trials over multiple seasons confirm that pyramided genotypes maintain less than 1% leaf infection under natural epidemic conditions where susceptible controls reach 80-95% infection and single-locus resistant cultivars show 5-20% infection, validating the durability advantage of the pyramiding approach.
A Pyramided_Rpv_Resistance_Disposition is an Rpv_Mediated_Resistance_Disposition that inheres in a Vitis plant carrying two or more distinct Rpv loci with different recognition specificities and is realized through multiple, independent effector-triggered immunity pathways upon encounter with Plasmopara viticola. This disposition represents the strategic combination of complementary resistance genes to achieve broad-spectrum, durable field resistance that cannot be overcome through single-step pathogen adaptation. The underlying principle is that while a pathogen population may evolve virulence against any single resistance gene through effector mutation or deletion, the simultaneous evolution of virulence against multiple, mechanistically independent resistance genes is statistically improbable and would typically incur severe fitness costs. Common pyramiding combinations in grapevine breeding include Rpv1+Rpv3, Rpv3+Rpv10, Rpv3+Rpv12, Rpv10+Rpv12, and three-way combinations such as Rpv3+Rpv10+Rpv12. Marker-assisted selection enables efficient identification of individuals carrying multiple resistance alleles in early seedling stages, dramatically accelerating the development of pyramided cultivars. Pyramided genotypes typically display enhanced resistance phenotypes compared to single-locus carriers, with additive or synergistic effects resulting in more rapid, complete pathogen arrest and reduced probability of resistance breakdown under field conditions.
Pyramided_Rpv_Resistance_Disposition is_a Rpv_Mediated_Resistance_Disposition. This class represents combinations of two or more Rpv loci, distinguished from classes representing individual loci. The specific loci present in a pyramided genotype should be documented in associated quality or information entity classes. Pyramiding is the recommended breeding strategy for durable resistance given documented instances of Rpv3 resistance erosion in European vineyards with high PIWI cultivar adoption.
Kombinierte Rpv-Resistenz
Pyramidierte Rpv-Resistenz-Disposition
Rpv-Genpyramide
combined Rpv resistance
multi-locus DM resistance
stacked Rpv resistance
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Grapevine Breeding Programs for the Wine Industry, Woodhead Publishing Zini E et al. (2019) Applying genomic information for grapevine downy and powdery mildew resistance, Acta Horticulturae 1248:1-8 Schneider C et al. (2019) From resistance gene pyramiding to marker-assisted resistance breeding in grapevine, OENO One 53:649-656
Pyramided Rpv Resistance Disposition
Q₁₀ coefficient assessment in Merlot comparing photosynthesis and respiration reveals the differential temperature sensitivity affecting carbon balance, with leaf respiration showing Q₁₀ of 2.2 (rate increasing from 1.5 to 3.3 μmol CO₂/m²/s between 20°C and 30°C) while net photosynthesis shows effective Q₁₀ of only 1.4 over the same range (increasing from 12 to 16.8 μmol CO₂/m²/s), with the higher respiratory temperature sensitivity meaning that the photosynthesis/respiration ratio declines from 8.0 at 20°C to 5.1 at 30°C, reducing net carbon retention efficiency under warmer conditions.
Q₁₀ coefficient quality is a temperature response quality that represents the factor by which a physiological process rate increases for each 10°C rise in temperature, characterizing the temperature sensitivity of metabolic reactions through this widely used coefficient that enables prediction of rate changes across temperature ranges relevant to viticultural conditions. Within the BFO framework, this quality specifically addresses the quantitative temperature sensitivity dimension, distinguishing the Q₁₀ parameter from other aspects of temperature response (optima, thresholds) while recognizing that Q₁₀ provides the practical coefficient for predicting metabolic rate changes that affect carbon balance, development rate, and stress responses across the temperature variations experienced in vineyards. The ontological significance lies in Q₁₀ providing the scaling factor for temperature effects on vine physiology, where Q₁₀ values enable prediction of how diurnal temperature fluctuations, seasonal warming, and climate change will affect metabolic processes, with the differential Q₁₀ values of photosynthesis versus respiration determining how temperature affects net carbon balance. The mathematical basis for Q₁₀ derives from the Arrhenius relationship between reaction rate and temperature, with Q₁₀ = (R₂/R₁)^(10/(T₂-T₁)) where R represents rates at temperatures T, and with Q₁₀ theoretically constant but practically varying somewhat across temperature ranges due to the approximation involved in this simplified representation of temperature effects. The typical Q₁₀ values for grapevine physiological processes show respiration with Q₁₀ of 2.0-2.5 (doubling to 2.5-fold increase per 10°C), photosynthesis with lower effective Q₁₀ of 1.5-2.0 in the sub-optimal range (limited by diffusional and photochemical processes with lower temperature sensitivity), and developmental processes with Q₁₀ of 2.0-3.0 depending on the specific process. The carbon balance implications of differential Q₁₀ values are significant, as the higher Q₁₀ of respiration compared to photosynthesis means that warming increases respiratory losses proportionally more than photosynthetic gains, reducing net carbon balance and explaining part of the productivity challenge in warming climates. Understanding Q₁₀ coefficient quality enables prediction of metabolic responses to temperature change, carbon balance modeling across thermal environments, and appreciation of how differential temperature sensitivity affects the balance between carbon gain and loss processes.
Q₁₀_Coefficient_Quality addresses the quantitative temperature sensitivity parameter used for predicting metabolic rate changes. This quality enables modeling of temperature effects on vine physiology across the diurnal and seasonal temperature ranges. The class supports carbon balance prediction and climate impact assessment through Q₁₀-based rate scaling.
Q10
Q₁₀-Koeffizient
Temperaturkoeffizient
Temperaturquotient
temperature coefficient
temperature quotient
thermal sensitivity factor
thermischer Sensitivitätsfaktor
Atkin OK, Tjoelker MG (2003) Thermal acclimation and the dynamic response of plant respiration to temperature. Trends in Plant Science 8(7): 343-351 Ryan MG (1991) Effects of climate change on plant respiration. Ecological Applications 1(2): 157-167
Q10 Coefficient Quality
Quality analysis data for Cabernet Sauvignon from Oakville, Napa Valley (2023 vintage) integrates must analysis (25.2° Brix, pH 3.62, TA 5.8 g/L, YAN 245 mg/L), berry composition (berry weight 1.35 g, skin weight 0.28 g, seed weight 0.065 g, skin-to-pulp ratio 0.26), phenolic data (total phenolics 68 AU280, total anthocyanins 1.95 mg/g, tannin 2.8 mg/g catechin equivalents), and aromatic analysis (methoxypyrazines 8.5 ng/L IBMP, below sensory threshold) — collectively establishing the lot as premium quality with full phenolic maturity and minimal herbaceous character. Quality analysis data for Grüner Veltliner from Kamptal, Austria (2024 vintage) documents must analysis (19.5° KMW, pH 3.18, TA 7.2 g/L), aromatic precursor analysis (glutathionylated thiol precursors 185 µg/L, rotundone below detection limit), confirming varietal typicity with characteristic peppery-citrus aromatic potential.
Quality analysis data is a grapevine information entity that documents the comprehensive analytical characterization of grape must, berry composition, and aromatic compound profiles obtained through standardized laboratory and field measurements, encompassing sugar content, organic acid composition, nitrogen status, phenolic concentration, berry physical parameters, and volatile compound profiles, providing the quantitative chemical foundation for harvest timing decisions, winemaking protocol design, and the objective assessment of fruit quality potential for a given cultivar, site, and vintage.
Quality analysis data represents the objective measurement counterpart to subjective sensory assessment, providing the chemical metrics upon which modern precision viticulture and evidence-based winemaking decisions are founded. The information entity integrates three distinct analytical domains — must analysis (bulk liquid composition), berry composition (physical partitioning of berry components), and aromatic analysis (volatile and precursor compounds) — each requiring different sampling protocols, analytical instrumentation, and interpretive frameworks. The evolution from simple refractometer sugar measurement toward comprehensive multi-parameter berry analysis reflects the recognition that sugar content alone is an inadequate predictor of wine quality, particularly under climate change conditions where sugar accumulation increasingly decouples from flavor maturity. Sampling methodology fundamentally influences data quality — representative vineyard sampling requires statistically designed collection protocols (minimum 200-berry samples from multiple blocks and canopy positions) to capture the spatial heterogeneity inherent in commercial vineyards. The temporal dimension is equally important, as single-point harvest analysis provides a snapshot while multi-date maturity curve analysis characterizes the dynamic trajectory of compositional evolution. Laboratory accreditation (ISO 17025) and participation in proficiency testing programs ensure analytical accuracy and inter-laboratory comparability of quality analysis data.
Qualitätsanalysedaten
Qualitätsuntersuchungsdaten
Traubenanalysedaten
analytical quality data
analytische Qualitätsdaten
fruit quality data
grape analysis data
must and berry analysis
quality analysis record
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed, Wiley, pp.: 1-88 Iland P et al. (2011) Chemical Analysis of Grapes and Wine: Techniques and Concepts, 2nd Ed, Patrick Iland Wine Promotions, pp.: 1-45 Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 369-420
Quality Analysis Data
Pinot Noir clone 115 bears a Quality_Clone_Role as the benchmark Dijon quality selection, producing wines of exceptional concentration and aromatic complexity from yields 30-40% below productive clones—commanding adoption for premium Burgundy, Oregon, and New Zealand production despite reduced productivity. Chardonnay clone 76 serves in a Quality_Clone_Role with documented superiority in Grand Cru Burgundy trials, its small-cluster character and concentrated must composition justifying yield sacrifices for Meursault and Corton-Charlemagne producers. Nebbiolo clone CVT 71 bears a Quality_Clone_Role in Barolo production, selected for optimal anthocyanin development and tannin structure enabling the extended aging that defines fine Nebbiolo wines.
A performance clone role that inheres in a grapevine clone characterized by superior wine quality potential relative to other clones of the same variety, typically sacrificing yield for concentrated fruit composition, optimal phenolic development, and enhanced aromatic complexity. Quality clones achieve elevated quality metrics through characteristics that limit production—smaller berries (higher skin-to-juice ratio concentrating phenolics and flavor compounds), smaller clusters (reduced dilution, improved ripening uniformity), millerandage tendency (uneven berry development creating complexity), moderate vigor (limiting vegetative competition with fruit development), and earlier ripening (achieving optimal maturity before autumn weather deterioration). Selection for quality emerged as wine markets increasingly valued differentiation and terroir expression, with premium producers willing to accept yield reductions of 30-50% in exchange for documented quality improvements. Quality clone selection represents a value proposition—reduced production offset by price premiums for wines of recognized superior character. The Dijon clones of Pinot Noir (115, 667, 777) exemplify quality clone development, selected for small berries, loose clusters, and wines of aromatic intensity and concentration prized by premium Burgundy producers and subsequently adopted globally.
Quality_Clone_Role captures the premium-focused end of the clone performance spectrum, where wine quality rather than production volume drives selection. The designation reflects market economics—quality clones are economically justified only where wine prices reward quality differentiation, making them appropriate for premium appellations while high-yield clones may better serve value-tier production. Quality is not absolute but market-defined what constitutes "quality" varies with style preferences, with some markets valuing concentration and intensity while others prefer elegance and restraint. Quality clone selection has transformed premium wine production by enabling producers to plant genotypes with documented quality advantages, though critics argue that widespread adoption of few quality clones reduces vineyard diversity and wine complexity.
Edelklon
Premiumklon
Qualitätsklon
Spitzenweinklon
fine wine clone
low-yield high-quality clone
premium clone
quality clone
Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 ENTAV-INRA (2007) Catalogue des Variétés et Clones This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Wolpert JA et al. (1994) Pinot Noir clones, American Journal of Enology and Viticulture 45: 356-363
Quality Clone Role
Riesling serves in a Quality_Donor_Role in German resistance breeding, contributing wine quality genetics (terpene biosynthesis, balanced acidity, refined phenolics) to breeding lines carrying disease resistance from earlier interspecific crosses. Pinot Noir bears a Quality_Donor_Role for color stability and wine structure in programs developing resistant red wine varieties, its quality attributes combined with resistance from donor parents. Regent (an established PIWI variety) functions in a Quality_Donor_Role in second-generation breeding, its acceptable wine quality making it more useful than raw wild species for programs seeking to combine resistance with commercial viability.
A donor parent role that inheres in a grapevine genotype by virtue of its function as the source of fruit quality or wine quality traits targeted for incorporation into breeding populations, particularly for improving the enological potential of disease-resistant hybrid materials. Quality donors are typically elite V. vinifera varieties with established wine quality credentials (Riesling, Pinot Noir, Cabernet Sauvignon, Chardonnay), used to elevate wine character in breeding lines where earlier generations prioritized disease resistance from wild species sources at the expense of quality. Quality donor use recognizes that wild Vitis species and early-generation hybrids typically produce wines with "hybrid character" (foxy flavors from American species, excessive acidity, atypical phenolic profiles) considered unacceptable for premium wine production, requiring quality improvement through crossing with V. vinifera quality parents. Quality traits are typically quantitative and polygenic, controlled by multiple genes with complex environmental interactions, making quality improvement more challenging to achieve through targeted introgression than single-gene resistance traits. Advanced PIWI varieties increasingly function as quality donors themselves, having achieved sufficient wine quality through generations of selection to contribute positively to further breeding.
Quality_Donor_Role captures the V. vinifera quality contribution essential for producing commercially viable resistant varieties. The quality donor concept acknowledges that disease resistance and wine quality often derive from different genetic sources—wild species for resistance, V. vinifera for quality—requiring breeding strategies that combine both trait categories. Quality traits are more complex genetically than major R-genes, typically involving multiple quantitative trait loci (QTL) with individually modest effects, making quality improvement through a few crosses insufficient. Contemporary PIWI breeding increasingly emphasizes quality, with advanced programs selecting primarily for wine quality among resistance-confirmed progeny rather than treating quality as an afterthought. The emergence of registered PIWI varieties (Regent, Solaris, Cabernet Blanc) as quality donors in further breeding reflects generational progress, with each breeding cycle incrementally improving wine character while maintaining resistance.
Qualitätselter
Qualitätsspender
V. vinifera donor
Vinifera-Spender
enological quality source
quality donor
quality parent
önologische Qualitätsquelle
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Töpfer R et al. (2011) Molecular breeding in grapes, CRC Press pp.: 160-185 Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330 Reynolds A (ed) (2015) Grapevine Breeding Programs for the Wine Industry, Elsevier
Quality Donor Role
Quality performance data for Pinot Noir clone 777 (Dijon) compiled across 10 vintages in Sonoma Coast documents must sugar of 23.5-25.0° Brix at harvest, titratable acidity of 6.5-8.0 g/L, pH 3.35-3.55, total anthocyanins of 1.2-1.8 mg/g berry fresh weight, and micro-vinification sensory panel scores averaging 16.5/20 for aromatic complexity with consistent notes of dark cherry and spice — positioning it as a premium quality clone for cool-climate red wine production. Quality performance data for Welschriesling clone WR-3 at Klosterneuburg records 10-vintage means of 18.5° KMW, total acidity 7.8 g/L, free volatile terpenes 1.2 mg/L (predominantly linalool and α-terpineol), and Spätlese-level micro-vinifications consistently rated above population mean for varietal typicity and aromatic intensity.
Quality performance data is a performance data entity that documents the fruit composition, must analytical parameters, and wine sensory characteristics achieved by a clonal line across multiple vintages in replicated evaluation trials, encompassing sugar accumulation dynamics, organic acid profiles, phenolic content and composition, aromatic compound concentrations, and micro-vinification sensory assessments, providing the oenological evidence base for evaluating a clone's potential to produce wines of defined style, typicity, and quality level under specified growing conditions.
Quality performance data represents the most complex and multi-dimensional component of clone evaluation, integrating analytical chemistry measurements with subjective sensory assessment across the transformation chain from grape to wine. The challenge of quality data interpretation lies in the inherent subjectivity of quality definitions — what constitutes "quality" differs between sparkling wine production (low sugar, high acidity, neutral aroma) and late-harvest sweet wine production (high sugar, concentrated flavor, Botrytis tolerance), meaning that quality rankings are always relative to a defined wine style objective. Micro-vinification protocols used in clone evaluation must balance standardization (enabling fair comparison between clones) against the risk of masking clone-specific characteristics through uniform winemaking — some programs conduct both standardized and clone-optimized vinifications to capture the full quality potential. The temporal dimension of quality data is critical because berry composition at any single sampling date provides an incomplete picture maturity curve analysis tracking sugar, acidity, and phenolic evolution across multiple harvest dates characterizes the ripening dynamics and optimal harvest window of each clone. Clone-by-rootstock and clone-by-site interactions for quality parameters can be substantial, particularly for phenolic accumulation and aromatic development, necessitating cautious extrapolation of quality data between evaluation sites and commercial vineyard conditions.
Qualitätsdaten
Qualitätsleistungsdaten
Traubenqualitätsdaten
Weinqualitätsbewertung
clone quality record
oenological performance
quality evaluation data
wine quality performance
önologische Leistungsdaten
Mannini F (2004) Clonal selection in grapevines: interactions between genetic and sanitary strategies. Acta Horticulturae 652: 21-27 Anderson MM et al. (2008) Viticultural evaluation of French and California Pinot Noir clones. American Journal of Enology and Viticulture 59(2): 188-193 Regner F et al. (2003) Clonal selection and clone identification of Austrian grapevine varieties. Mitteilungen Klosterneuburg 53: 172-180
Quality Performance Data
Quality potential varies enormously across V. vinifera cultivars, with premium wine varieties such as Pinot Noir and Riesling prized for balanced sugar-acid ratios (20-24°Brix with 6-9 g/L titratable acidity), complex aromatic profiles, and age-worthy phenolic structures, while bulk wine cultivars such as Airén may achieve high yields but produce simpler wines. Quality parameters include sugar concentration (18-28°Brix), titratable acidity (4-10 g/L tartaric equivalents), pH (3.0-4.0), total phenolics (200-4000 mg/L gallic acid equivalents), anthocyanins in red cultivars (200-2000 mg/L), and varietal aroma compounds (terpenes 0.5-6 mg/L in aromatic cultivars). Terroir expression reflects genotype × environment interaction for quality traits, with identical cultivars producing distinct quality profiles across sites.
A grapevine disposition that is the inherited capacity of a Vitis vinifera plant to accumulate and balance the chemical constituents that determine grape and wine quality, encompassing traits related to sugar concentration, organic acid metabolism, phenolic compound synthesis, aromatic precursor accumulation, and color stability in the resulting wine.
Quality_Potential_Disposition serves as the top-level parent class for all traits contributing to grape and wine quality. Quality is inherently multidimensional, with different parameters prioritized for different wine styles and markets. The trait complex shows strong genotype × environment × management interaction, making quality expression highly context-dependent. Rootstock effects on scion quality potential are significant but secondary to scion genotype effects for most quality parameters.
Qualitätspotential
fruit quality trait
oenological potential disposition
quality component capacity
önologisches Potential
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 343-425 Jackson RS (2020) Wine Science: Principles and Applications, 5th Ed, Academic Press pp.: 287-376
Quality Potential Disposition
Quality rating for Penfolds Grange Shiraz (2021 vintage) documents 98 points (Wine Advocate, individual critic assessment), 19/20 (Jancis Robinson, individual critic), and a Gold Medal with Trophy (National Wine Show of Australia, blind panel consensus) — the convergence of high scores across different systems and evaluation contexts confirming exceptional quality. Quality rating for Grüner Veltliner Smaragd from the Wachau (2024 vintage) records 93 points (Falstaff, Austrian panel), 17.5/20 (Jancis Robinson), and 4.5 stars (A la Carte wine guide), placing the wine in the "excellent" tier across all assessment frameworks. Quality rating for Blaufränkisch Reserve from Mittelburgenland DAC (2022 vintage) documents 91 points (Falstaff), 92 points (Wine Enthusiast), representing the upper range of regional Blaufränkisch quality and confirming the wine's positioning as a premium-tier DAC expression.
Quality rating is an overall assessment entity that records the categorical or numerical quality score assigned to a wine through formal sensory evaluation, encompassing the principal scoring systems employed globally — the 100-point scale (Robert Parker/Wine Advocate origin, now industry standard, with effective range 80-100 for commercially reviewed wines), the 20-point scale (traditional European, Jancis Robinson/Vinum, with finer discrimination in the 15-20 range), the 5-star scale (consumer-oriented, Halliday and similar systems), and the categorical medal system (gold, silver, bronze, commended, used in wine competitions) — together with the evaluation context (individual critic, panel consensus, competition blind tasting, appellation approval), and providing the summary quality metric that most directly influences commercial pricing, retail placement, and consumer purchasing behavior in the global wine market.
Quality rating is simultaneously the most influential and most controversial element of wine assessment — influential because numerical scores have become the primary currency of wine marketing and commerce (a difference of 2-3 points on the 100-point scale can translate to 20-50% price differentials at retail), and controversial because the reduction of a complex, multidimensional sensory experience to a single number raises fundamental questions about the validity, reliability, and utility of wine scoring. The 100-point scale, despite its nominal range, operates effectively within a much narrower band — wines scoring below 80 are rarely published, wines scoring 80-84 are considered "good," 85-89 "very good," 90-94 "outstanding," and 95-100 "extraordinary," compressing the entire quality spectrum into a functional 20-point range that limits discriminative precision. The inter-rater reliability of wine scoring — the degree to which different qualified tasters assign similar scores to the same wine — has been extensively studied with sobering results: correlation coefficients between experienced judges typically range from 0.3-0.6, and individual tasters often assign significantly different scores to the same wine presented on different occasions, suggesting that a single score represents a substantial simplification of an inherently variable perceptual process. The distinction between absolute quality assessment (rating a wine against a universal quality standard) and relative quality assessment (rating a wine within its category — varietal, regional, price tier) creates fundamental interpretive ambiguity, as a "90-point Muscadet" and a "90-point Barolo" represent radically different quality claims. Competition medal systems mitigate individual taster variability through panel consensus protocols but introduce their own biases — competition fatigue effects (wines tasted later in a session receive systematically lower scores), order effects (wines preceded by very high or very low quality wines receive distorted scores), and the impossibility of directly comparing wines across different competition heats. Despite these methodological limitations, quality ratings serve an important market function by reducing information asymmetry between producers and consumers, particularly for unfamiliar wines, regions, and vintages.
Bewertungsscore
Punktebewertung
Qualitätsbewertung
Qualitätsnote
Weinbewertung
quality classification
quality score
tasting score
wine rating
wine score
Hodgson RT (2008) An examination of judge reliability at a major U.S. wine competition. Journal of Wine Economics 3(2): 105-113 Amerine MA, Roessler EB (1983) Wines: Their Sensory Evaluation, 2nd Ed, Freeman, pp.: 170-220 Ashton RH (2012) Reliability and consensus of experienced wine judges: expertise within and between? Journal of Wine Economics 7(1): 70-87 Quandt RE (2007) On wine bullshit: some new software? Journal of Wine Economics 2(2): 129-135
Quality Rating
A Cabernet Sauvignon vine at Opus One in the Oakville AVA of Napa Valley fulfills a quality tier role that determines its viticultural management intensity — vines designated for the icon wine receive cluster thinning to 4-6 clusters per shoot at véraison, leaf pulling on the morning-sun side, yield restriction to 3-5 t/ha, and individual block-level ripeness monitoring with daily Brix measurements, while genetically identical vines in adjacent blocks assigned to the second label Overture receive less intensive management at yields of 6-8 t/ha, illustrating how quality tier roles differentiate management of otherwise equivalent vines.
A grapevine role that inheres in a vine by virtue of the market positioning and price-quality classification assigned to the wine produced from its fruit, reflecting the producer's strategic allocation of viticultural investment, yield management, and winemaking resources that collectively determine the commercial tier at which the resulting wine is marketed and priced.
Quality_Tier_Role classifies grapevines according to the commercial value chain position of their fruit output rather than by geographic origin, cultivar identity, or production system. The role is assigned by the producer based on vine age, block performance, vintage assessment, and market strategy: it is therefore mutable across vintages and subject to declassification (downgrading fruit from a higher to a lower tier) or promotion. The subclass hierarchy follows the widely adopted Rabobank/Silicon Valley Bank wine industry segmentation model, ranging from Icon (>$50 retail) through Bulk (<$3 retail), though exact price boundaries vary by market and currency.
Marktsegment-Rolle
Preis-Qualitäts-Rolle
Qualitätsstufen-Rolle
market tier role
price-quality positioning role
quality classification role
Rabobank (2019) Wine Quarterly Report: Global Wine Market Segmentation McMillan R (2022) State of the US Wine Industry, Silicon Valley Bank Anderson K (2004) The World's Wine Markets: Globalization at Work, Edward Elgar Publishing pp: 11-38
Quality Tier Role
QTL mapping for Quantitative_Black_Rot_Resistance_Disposition in an 'Horizon' × Illinois 547-1 population at Cornell University identified five genomic regions contributing to black rot resistance independent of the Rgr1 locus. Phenotypic evaluation across three seasons with varying disease pressure employed both detached leaf assays and field cluster assessments, scoring lesion severity, sporulation intensity, and berry infection percentage. QTLs on chromosomes 5, 7, 10, 14 (distinct from Rgr1), and 17 each explained 8-18% of phenotypic variance, with additive effects combining to produce a 40-65% reduction in disease severity when favorable alleles were accumulated. Progeny inheriting favorable alleles at four or more QTLs exhibited mean berry infection of 15-25% compared to 55-70% in individuals with predominantly unfavorable alleles, under natural epidemic conditions where susceptible V. vinifera controls reached 90%+ infection. Importantly, the quantitative resistance remained effective across years with different environmental conditions and presumed pathogen population variation, supporting the durability hypothesis. Genomic selection models incorporating all mapped QTLs achieved prediction accuracies of 0.58-0.67 for black rot severity, enabling efficient breeding progress without phenotyping individual progeny.
A Quantitative_Black_Rot_Resistance_Disposition is a Black_Rot_Resistance_Disposition that inheres in a Vitis plant and is realized through the cumulative action of multiple genes with individually moderate effects, collectively conferring partial but durable resistance to Guignardia bidwellii across diverse environmental conditions and pathogen populations. Unlike monogenic Rgr-mediated resistance, quantitative black rot resistance follows continuous inheritance patterns controlled by multiple quantitative trait loci (QTLs) distributed across the genome, each contributing incrementally to the overall resistance phenotype. The molecular and physiological mechanisms underlying quantitative resistance likely include enhanced constitutive defense compound levels, more rapid or intense activation of induced defenses following pathogen recognition, modified tissue properties affecting penetration success, and variation in the duration of the biotrophic window before necrotrophy onset. Quantitative resistance typically manifests as reduced infection frequency, smaller lesion size, decreased pycnidial density within lesions, extended latent period, and lower berry susceptibility rather than complete immunity. The presumed durability advantage of quantitative resistance derives from the evolutionary difficulty of pathogen adaptation to multiple independent resistance mechanisms simultaneously. Quantitative black rot resistance is particularly relevant for breeding programs targeting humid continental climates, where combining partial resistances from multiple sources can achieve commercially acceptable disease control levels with reduced fungicide inputs, even in the absence of complete immunity.
Quantitative_Black_Rot_Resistance_Disposition is_a Black_Rot_Resistance_Disposition. This class is distinguished from Rgr_Mediated_Resistance_Disposition by its polygenic inheritance and partial resistance phenotype. Quantitative and major gene resistances may co-occur in the same genotype, with quantitative factors potentially enhancing durability of Rgr-mediated resistance. Multiple North American Vitis species contribute quantitative resistance factors that can be combined in breeding to achieve commercially meaningful protection levels.
Partielle Guignardia-Resistenz
Polygene Black-Rot-Resistenz
QTL-based black rot resistance
Quantitative Schwarzfäule-Resistenz-Disposition
partial Guignardia resistance
polygenic black rot resistance
Rex F et al. (2014) QTL mapping of black rot (Guignardia bidwellii) resistance in the grapevine rootstock 'Börner', Theoretical and Applied Genetics 127:1667-1677 Cadle-Davidson L (2008) Variation within and between Vitis spp. for foliar resistance to the downy mildew pathogen Plasmopara viticola, Plant Disease 92:1577-1584
Quantitative Black Rot Resistance Disposition
The Vitis vinifera cultivar 'Kishmish vatkana' from Central Asia exhibits Quantitative_Downy_Mildew_Resistance_Disposition despite lacking characterized Rpv loci, demonstrating that partial resistance exists within the V. vinifera gene pool. QTL mapping studies using a 'Kishmish vatkana' × 'Cabernet Sauvignon' population identified five genomic regions on chromosomes 4, 7, 12, 17, and 18 collectively explaining 42% of phenotypic variance for sporulation density. Individual QTLs contributed between 6% and 12% of variance, with additive effects combining to reduce sporulation by 60-70% compared to fully susceptible genotypes. In field evaluations at INRAE Bordeaux, 'Kishmish vatkana' maintained leaf infection severity of 25-35% during epidemic years when susceptible controls reached 85-95%, demonstrating that quantitative resistance provides meaningful protection even without complete immunity. Breeding programs increasingly combine quantitative resistance backgrounds with major Rpv genes, hypothesizing that the polygenic component enhances durability by imposing additional selective constraints on pathogen adaptation.
A Quantitative_Downy_Mildew_Resistance_Disposition is a Downy_Mildew_Resistance_Disposition that inheres in a Vitis plant and is realized through the cumulative action of multiple genes with small individual effects, collectively conferring partial but broadly effective resistance to Plasmopara viticola across diverse pathogen genotypes. Unlike monogenic Rpv-mediated resistance, quantitative resistance is inherited as a continuous trait controlled by numerous quantitative trait loci (QTLs) distributed across the genome, each contributing incrementally to the overall resistance phenotype. The molecular mechanisms underlying quantitative resistance are diverse and may include enhanced basal defense responses, more rapid or intense PAMP-triggered immunity, elevated constitutive levels of antimicrobial compounds, modified leaf surface properties affecting zoospore behavior, and subtle variations in cellular architecture impeding pathogen colonization. Quantitative resistance typically manifests as reduced infection frequency, smaller lesion size, decreased sporulation intensity, and extended latent period rather than complete immunity. While individual QTLs provide modest effects (typically explaining 5-15% of phenotypic variance), their combination can result in high levels of field resistance. A key advantage of quantitative resistance is its presumed durability, as overcoming multiple independent resistance mechanisms simultaneously presents a formidable evolutionary challenge for pathogen populations. However, the complex genetic architecture complicates breeding efforts, requiring genomic selection approaches or phenotypic selection under controlled disease pressure to effectively accumulate favorable alleles.
Quantitative_Downy_Mildew_Resistance_Disposition is_a Downy_Mildew_Resistance_Disposition. This class is distinguished from Rpv_Mediated_Resistance_Disposition by its polygenic inheritance and partial resistance phenotype. Quantitative resistance may co-occur with major gene resistance in the same genotype, and distinguishing their relative contributions requires genetic analysis. The term "horizontal resistance" is sometimes used synonymously but is deprecated due to ambiguity. QTLs for quantitative resistance may be distinct from or overlap with characterized Rpv loci.
Partielle Falscher-Mehltau-Resistenz
Polygene Peronospora-Resistenz
QTL-based Plasmopara resistance
Quantitative Peronospora-Resistenz-Disposition
horizontal DM resistance
partial DM resistance
polygenic downy mildew resistance
Bellin D et al. (2009) Resistance to Plasmopara viticola in grapevine 'Bianca' is controlled by a major dominant gene causing localised necrosis at the infection site, Theoretical and Applied Genetics 120:163-176 Buonassisi D et al. (2017) Breeding for grapevine downy mildew resistance: a review of "omics" approaches, Euphytica 213:103 Poland JA et al. (2009) Shades of gray: the world of quantitative disease resistance, Trends in Plant Science 14:21-29
Quantitative Downy Mildew Resistance Disposition
QTL mapping studies for Quantitative_Powdery_Mildew_Resistance_Disposition in a 'Cabernet Sauvignon' × 'Norton' population at the University of Missouri identified seven genomic regions contributing to powdery mildew resistance, none co-localizing with known major Ren or Run loci. Individual QTLs on chromosomes 2, 5, 9, 11, 13, 15, and 18 each explained 4-11% of phenotypic variance for colony density and sporulation intensity. Individuals accumulating favorable alleles at five or more QTLs showed 55-70% reduction in sporulation compared to those inheriting unfavorable alleles at most loci. Field evaluation over four seasons demonstrated that high-QTL selections maintained leaf infection severity of 20-30% under natural epidemic conditions while susceptible segregants reached 70-85%. Importantly, the quantitative resistance remained stable across years and locations with different E. necator population structures, supporting the durability hypothesis. Genomic selection models incorporating all mapped QTLs achieved prediction accuracies of 0.65-0.72 for powdery mildew severity, enabling efficient selection for quantitative resistance in breeding populations without phenotypic evaluation of individual progeny.
A Quantitative_Powdery_Mildew_Resistance_Disposition is a Powdery_Mildew_Resistance_Disposition that inheres in a Vitis plant and is realized through the cumulative action of multiple genes with individually small effects, collectively conferring partial but durable resistance to Erysiphe necator across diverse pathogen genotypes and environmental conditions. Unlike monogenic Ren- or Run-mediated resistance, quantitative powdery mildew resistance follows continuous inheritance patterns controlled by numerous quantitative trait loci (QTLs) distributed across the genome, each contributing incrementally to the overall resistance phenotype. The molecular mechanisms underlying quantitative resistance are diverse and may include enhanced basal defense activation, more effective PAMP-triggered immunity, elevated constitutive levels of antimicrobial compounds in the cuticle and epidermis, modified cell wall architecture impeding fungal penetration, and subtle variations in leaf surface properties affecting conidial germination and appressorium formation. Quantitative resistance typically manifests as reduced infection frequency, smaller colony size, decreased sporulation intensity, and extended latent period rather than complete immunity. While individual QTLs typically explain only 5-20% of phenotypic variance, their combination can provide commercially meaningful levels of field resistance. The presumed durability advantage of quantitative resistance stems from the evolutionary challenge facing pathogen populations in simultaneously overcoming multiple independent resistance mechanisms, each imposing incremental fitness costs on adapted genotypes.
Quantitative_Powdery_Mildew_Resistance_Disposition is_a Powdery_Mildew_Resistance_Disposition. This class is distinguished from Ren_Mediated_Resistance_Disposition and Run_Mediated_Resistance_Disposition by its polygenic inheritance and partial resistance phenotype. Quantitative resistance may co-occur with major gene resistance, and genomic selection approaches can efficiently accumulate favorable alleles. The presumed durability advantage requires validation through long-term deployment studies tracking pathogen population adaptation.
Partielle Echter-Mehltau-Resistenz
Polygene Oidium-Resistenz
QTL-based PM resistance
Quantitative Mehltau-Resistenz-Disposition
horizontal oidium resistance
partial PM resistance
polygenic powdery mildew resistance
Cadle-Davidson L et al. (2016) Lessons from a phenotyping center revealed by the genome-guided mapping of powdery mildew resistance loci, Phytopathology 106:1159-1169 Poland JA et al. (2009) Shades of gray: the world of quantitative disease resistance, Trends in Plant Science 14:21-29 Blanc S et al. (2012) Quantitative trait loci for resistance to powdery mildew in two segregating populations of grapevine, BMC Genetics 13:100
Quantitative Powdery Mildew Resistance Disposition
Quantity yield assessment across vineyard blocks in Merlot reveals the variation in production volume affecting operational planning, with high-yielding block (14 t/ha from vigorous vines with 24 clusters/vine averaging 280 g/cluster) requiring maximum harvest capacity allocation, moderate block (8 t/ha from balanced vines with 18 clusters/vine at 220 g/cluster) providing target production, and low-yielding block (4 t/ha from stressed vines with 12 clusters/vine at 165 g/cluster) indicating remediation needs, with the quantity assessment enabling harvest logistics planning and identifying blocks requiring management intervention to improve productivity.
Quantity yield quality is a yield quality that represents the mass of grapes produced, characterizing the weight-based production output that determines the volume of raw material available for processing and provides the primary measure of vineyard productivity expressed as yield per vine, per hectare, or other area-based metrics. Within the BFO framework, this quality specifically addresses the quantitative dimension of yield, distinguishing the amount produced from the quality attributes of what is produced while recognizing that quantity yield is the fundamental productivity measure that determines raw material availability and affects production economics through the basic relationship between fixed costs and production volume. The ontological significance lies in quantity yield establishing the production base upon which economic calculations depend, where higher quantity yields spread fixed costs (land, infrastructure, permanent labor) across more product units, potentially improving profitability even if per-unit quality is lower, while lower quantity yields concentrate quality but must command premium prices sufficient to offset the reduced volume, creating the economic context within which yield targeting decisions are made. The component structure of quantity yield recognizes that total yield is the mathematical product of yield components (clusters per vine × cluster weight, or equivalently clusters × berries per cluster × berry weight), with this multiplicative relationship meaning that deficits in any component proportionally reduce total yield, while component interactions may create compensation (fewer clusters sometimes producing larger clusters) or compounding (stress reducing multiple components simultaneously). The measurement and expression of quantity yield involves weighing harvested fruit and expressing the result per vine (typically 2-15 kg/vine depending on training system and objectives), per hectare (typically 3-20 t/ha for wine grapes, higher for table grapes or juice production), or per linear meter of row (useful for comparing different row spacings), with these different expressions serving different analytical purposes. Understanding quantity yield quality enables productivity assessment, economic planning based on expected production volumes, and yield component analysis identifying factors limiting total production.
Quantity_Yield_Quality serves as the parent class for specific yield components and metrics (Cluster Number, Cluster Weight, Berry parameters, per-vine, per-hectare). This quality addresses the mass-based production output determining raw material availability. The class supports productivity assessment and economic planning through quantity yield characterization.
Erntegewicht
Ertragsgewicht
Ertragsmengenqualität
Produktionsmenge
crop mass
harvest weight
yield weight
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 145-230 Dry PR, Coombe BG (2004) Viticulture Volume 1: Resources, 2nd Ed. Winetitles, Adelaide, pp. 150-200
Quantity Yield Quality
Quantum yield assessment comparing healthy versus iron-deficient Sauvignon Blanc leaves demonstrates the efficiency penalty from chlorosis, with healthy leaves showing quantum yield of 0.065 mol CO₂/mol photons (initial slope of light response curve) while iron-deficient chlorotic leaves display reduced yield of 0.032 mol CO₂/mol photons, representing a 51% efficiency loss that translates directly to proportional reduction in carbon gain under all light-limited conditions, with chlorophyll fluorescence (Fv/Fm = 0.82 versus 0.58) providing a rapid non-destructive confirmation of the quantum yield impairment.
Quantum yield quality is a carbon fixation efficiency quality that represents the molar ratio of CO₂ fixed to photons absorbed, characterizing the fundamental efficiency of converting light energy into chemical energy that determines photosynthetic performance under light-limited conditions and indicates the functional integrity of the photosynthetic apparatus. Within the BFO framework, this quality specifically addresses the photon-to-carbon conversion efficiency, distinguishing quantum yield from absolute photosynthetic rate while recognizing that quantum yield represents the intrinsic efficiency of photosynthesis independent of light intensity and provides the most sensitive indicator of photosynthetic apparatus health. The ontological significance lies in quantum yield providing the efficiency metric for photosynthesis that determines carbon gain under the light-limited conditions typical of interior canopy positions, early morning/late afternoon, and cloudy days, where efficiency rather than capacity limits productivity, making quantum yield essential for understanding whole-canopy carbon gain in realistic vineyard conditions. The physiological basis for quantum yield involves the coupling between light absorption, electron transport, and CO₂ fixation, with theoretical maximum quantum yield approximately 0.125 mol CO₂/mol photons (8 photons minimum required per CO₂ fixed) for C₃ plants, while actual values typically range from 0.04-0.08 due to photorespiratory losses, non-photochemical quenching, and other inefficiencies. The measurement approaches for quantum yield include the initial slope of light response curves at limiting light (Φ CO₂), chlorophyll fluorescence parameters (Fv/Fm for maximum potential, ΦPSII for operational yield), and oxygen evolution measurements avoiding respiratory complications. The stress sensitivity of quantum yield makes this parameter particularly valuable for early stress detection, with quantum yield declining before visible symptoms or maximum photosynthetic rate changes, enabling early diagnosis of photoinhibition, nutrient deficiency, or environmental stress affecting photosynthetic apparatus function. Understanding quantum yield quality enables assessment of light-limited photosynthetic efficiency, early stress detection through quantum yield monitoring, and whole-canopy productivity modeling that accounts for the diversity of light environments within the vine canopy.
Quantum_Yield_Quality addresses the fundamental photon-to-carbon conversion efficiency, providing the most sensitive indicator of photosynthetic apparatus health. This quality determines performance under light-limited conditions typical of significant canopy areas. The class supports early stress detection and efficiency assessment through quantum yield measurement or fluorescence proxies.
Lichtnutzungseffizienz
Quanteneffizienz
light use efficiency
photosynthetic efficiency
photosynthetische Effizienz
Φ CO₂
Björkman O, Demmig B (1987) Photon yield of O₂ evolution and chlorophyll fluorescence characteristics at 77 K among vascular plants of diverse origins. Planta 170(4): 489-504 Long SP et al. (1993) Quantum yields for uptake of carbon dioxide in C₃ vascular plants of contrasting habitats and taxonomic groupings. Planta 189(2): 226-234
Quantum Yield Quality
Quarantine status for Grüner Veltliner cuttings imported from Austria to New Zealand documents the complete quarantine trajectory — pre-quarantine status assigned upon import permit approval and pre-export testing verification, active quarantine status during the mandatory post-entry quarantine period (minimum 2 growing seasons at the MPI-approved Level 3A quarantine facility), with sequential testing for Grapevine leafroll-associated viruses (GLRaV-1, -2, -3, -4 and variants), Grapevine fanleaf virus, Arabis mosaic virus, Grapevine fleck virus, Australian grapevine viroid, and additional regulated organisms by ELISA, RT-PCR, and biological indexing on indicator plants, and post-quarantine cleared status granted only after all testing cycles return negative results — the quarantine process ensuring that valuable imported germplasm does not introduce pathogens to New Zealand's high-phytosanitary-status vineyards. Quarantine status for vine material moving within the EU phytosanitary area is substantially less restrictive — intra-EU movement requires a plant passport (Regulation 2016/2031) confirming the material meets EU phytosanitary standards, but no post-entry quarantine is required, reflecting the common phytosanitary area and mutual recognition of member state inspection systems.
Quarantine status is a trade regulation entity that documents the phytosanitary risk classification assigned to grape plant material based on its origin, health testing history, and the pest and disease status of its source region, encompassing the quarantine category of the material (pre-quarantine, in quarantine, post-quarantine cleared, quarantine-exempt), the specific regulated pests and diseases for which quarantine monitoring is conducted, the quarantine facility and protocol under which the material is held, the testing schedule and methods applied during the quarantine period, and the release conditions that must be met for the material to exit quarantine and enter commercial propagation or planting, providing the phytosanitary risk management framework that governs the transition of imported plant material from regulated confinement to unrestricted commercial use.
Quarantine status documentation captures the dynamic phytosanitary classification of plant material as it progresses through the biosecurity clearance process — a classification that changes over time as successive testing cycles either confirm or refute the presence of regulated organisms, and that determines whether the material remains in regulated confinement or may be released for commercial use. The quarantine concept in viticultural plant material trade is predicated on the recognition that many economically significant grapevine pathogens — particularly viruses and virus-like organisms — may be present in symptomless infections that are undetectable by visual inspection alone, necessitating laboratory-based diagnostic testing over multiple growing seasons to achieve acceptable detection confidence. The testing protocols applied during quarantine reflect the current state of diagnostic technology — traditional biological indexing (grafting suspect material onto sensitive indicator varieties and observing for symptom expression over 1-2 growing seasons) is being progressively supplemented and in some cases replaced by molecular methods (RT-PCR, real-time PCR, high-throughput sequencing) that offer faster, more sensitive, and more specific pathogen detection, though regulatory acceptance of molecular methods as sole quarantine diagnostics varies across jurisdictions. The economic implications of quarantine requirements are substantial — the direct costs of quarantine (facility fees, testing charges, material maintenance) combined with the opportunity cost of delayed commercial availability (3-5 years from importation to vineyard planting in stringent jurisdictions) create a significant barrier to international cultivar exchange that disproportionately affects minor cultivars and small-scale importers. The concept of "quarantine pest" — an organism of potential economic importance to the area endangered thereby and not yet present there, or present but not widely distributed and being officially controlled (IPPC definition) — determines which specific pathogens drive quarantine requirements for each importing jurisdiction, with the quarantine pest list varying by country according to each jurisdiction's current pest status and risk assessment.
Biosicherheitsstatus
Pflanzengesundheitsstatus
Pflanzenschutzstatus
Quarantäneklassifizierung
Quarantänestatus
biosecurity status
phytosanitary status
plant health status
quarantine category
quarantine classification
FAO (2023) ISPM 5 — Glossary of phytosanitary terms (quarantine pest definition) Australian Government DAFF (2023) Post-entry quarantine procedures for Vitis MPI New Zealand (2023) Import health standard for nursery stock — Vitis European Parliament (2016) Regulation (EU) 2016/2031, Articles 79-95 (plant passports)
Quarantine Status
Quercetin content quality in Oregon Pinot Noir provides a quantitative measure of vineyard sun exposure management, with open-canopy blocks achieving quercetin-3-glucoside concentrations of 40-70 mg/kg compared to 8-15 mg/kg in dense-canopy sites winemakers correlate elevated quercetin with improved color stability through co-pigmentation, targeting fruit zone leaf removal to achieve 35-50% cluster exposure that optimizes the quercetin accumulation supporting the bright ruby color and age-worthy structure characteristic of premium Willamette Valley Pinot Noir.
Quercetin_Content_Quality represents the chemical quality characterizing the concentration of quercetin and its glycosides in grape berry skins, capturing the dominant flavonol in most Vitis vinifera cultivars. Quercetin (3,3',4',5,7-pentahydroxyflavone) bears hydroxyl groups at the 3' and 4' positions of the B ring, representing the dihydroxylated flavonol analogous to cyanidin among anthocyanidins. Quercetin is the most abundant flavonol in grape skins, typically comprising 50-80% of total flavonols, present primarily as quercetin-3-glucoside and quercetin-3-glucuronide with smaller amounts of other glycosides. The concentration of quercetin glycosides in grape skins responds dramatically to light exposure, with UV-B radiation inducing flavonol synthase (FLS) gene expression and quercetin accumulation as part of the photoprotective response. Sun-exposed berries may accumulate quercetin-3-glucoside at concentrations of 20-80 mg/kg fresh weight, while shaded berries contain less than 5 mg/kg. This light sensitivity makes quercetin content a biomarker of canopy light environment useful for assessing vineyard management effects. In wine, quercetin participates in co-pigmentation reactions that enhance anthocyanin color expression, and may undergo oxidation reactions that contribute to wine aging chemistry. Quercetin aglycone released by glycoside hydrolysis is relatively insoluble and can precipitate during wine storage, contributing to sediment formation. Health interest in quercetin centers on its antioxidant, anti-inflammatory, and potential cardioprotective effects, making grape and wine quercetin content relevant to functional food research.
Quercetin_Content_Quality addresses the dominant flavonol in most grape cultivars, representing the dihydroxylated B-ring structure parallel to cyanidin among anthocyanidins. The strong correlation between quercetin content and UV-B exposure makes it a sensitive indicator of fruit zone light environment. Quercetin glycoside profiles (glucoside, glucuronide, galactoside ratios) show cultivar-specific patterns useful for authenticity assessment. The co-pigmentation activity of quercetin with anthocyanins contributes to wine color enhancement exceeding what anthocyanin concentration alone would predict. Quercetin quantification employs HPLC with UV detection, with glycosides quantified individually or after hydrolysis to aglycone.
Dihydroxyflavonolgehalt
Q-3-glc content
Quercetin-3-glucosid-Konzentration
Quercetingehalt
Quercetinglykosidgehalt
dihydroxyflavonol quality
quercetin glycoside content
quercetin-3-glucoside concentration
Price SF et al. (1995) Cluster Sun Exposure and Quercetin in Pinot Noir Grapes and Wine. American Journal of Enology and Viticulture 46(2): 187-194 Castillo-Muñoz N et al. (2007) Flavonol Profiles of Vitis vinifera Red Grapes and Their Single-Cultivar Wines. Journal of Agricultural and Food Chemistry 55(3): 992-1002
Quercetin Content Quality
In Cabernet Franc vines in Chinon, Loire Valley, quiescence maintenance extends from early February when chilling requirements are satisfied through late March when spring warming triggers budbreak, with quiescence maintained by mean temperatures of 6-9°C during February and early March that are insufficient to support growth despite fully restored bud competence, and terminating when late March temperatures consistently exceed 10°C and accumulated growing degree days reach the threshold for budbreak initiation.
Quiescence maintenance in Vitis vinifera represents the sustained preservation of the ecodormant quiescent state through continued environmental limitation of growth, with quiescence persisting as long as environmental conditions remain unfavorable for growth initiation and being maintained by the ongoing constraint of low temperatures or other limiting factors rather than by active internal inhibitory mechanisms. This maintenance phase extends from the completion of endodormancy release through the onset of sustained favorable conditions that trigger budbreak, encompassing the period of waiting during which growth-competent buds remain inactive due solely to environmental inadequacy for growth support. The mechanistic basis of quiescence maintenance differs fundamentally from endodormancy maintenance in involving no active regulatory suppression, with quiescence being a passive consequence of environmental limitation rather than an actively maintained physiological state, and with growth occurring immediately when environmental limitation is removed without requiring any release process. The temperature basis of quiescence maintenance involves the continuous operation of thermal limitation on growth processes, with temperatures below the approximately 10°C base threshold being insufficient to support the sustained metabolic activity required for net biosynthesis and development, effectively preventing growth regardless of bud competence. The duration of quiescence maintenance varies substantially with climate and season, ranging from a few weeks in mild Mediterranean climates to several months in cold continental regions, and varying year-to-year within regions depending on the timing of spring warming relative to chilling completion. The metabolic state during maintained quiescence involves continued low-level activity that gradually prepares buds for growth, with slow accumulation of growth-related transcripts and proteins, gradual metabolic reactivation within the constraints imposed by temperature, and maintenance of cellular integrity and viability throughout the quiescent period. The termination of quiescence maintenance occurs when environmental limitation is removed through sustained warming that provides sufficient heat accumulation for budbreak initiation, marking the transition from dormancy to active growth that begins the new growing season.
Quiescence_Maintenance represents the sustained inactive state within Ecodormancy during the period between dormancy release and budbreak. Distinguished from active Dormancy_Maintenance during endodormancy by passive rather than active inhibition. Subdivides into Sustained_Environmental_Inhibition, Low_Temperature_Constraint, and Pre_Budbreak_Quiescence as component aspects. Duration varies from weeks to months with climate. Involves gradual preparatory changes despite maintained inactivity. Terminates with spring warming and heat unit accumulation.
Quieszenz-Aufrechterhaltung
anhaltende Ruhe
continued environmental dormancy
fortgesetzte Umweltdormanz
maintained dormant state
persistent ecodormancy
persistierende Ekodormanz
Quiescence Maintenance
Rachis brittleness monitoring in Chardonnay for mechanical harvest timing shows fracture stress declining from 4.2 MPa at véraison to 2.8 MPa at 21°Brix and 1.9 MPa at 24°Brix, with field trials demonstrating shattering losses of 3.5% at the higher fracture stress increasing to 12.8% at the lower stress, establishing the 2.5 MPa threshold (approximately 22°Brix) as the mechanical harvest window endpoint where rachis brittleness begins causing economically significant berry loss.
Rachis brittleness quality is a brittleness quality that inheres in the grape cluster main axis and branches and represents the susceptibility of the rachis structural framework to fracture under mechanical stress, determining the cluster's resistance to shattering during handling, transport, and mechanical harvest. Within the BFO framework, this quality specifically characterizes the fracture behavior of the cluster support structure, distinguishing rachis mechanical properties from those of berries and pedicels while recognizing the critical role of rachis integrity in maintaining cluster cohesion throughout the postharvest chain. The ontological significance lies in rachis brittleness determining the viability of mechanical harvest and the extent of berry loss during handling, where brittle rachis fractures under the vibration and impact forces of mechanical harvesters releasing individual berries from cluster fragments, while flexible non-brittle rachis bends without breaking maintaining cluster structure. Rachis brittleness increases during ripening as lignification progresses and moisture content declines, with the rate and extent of brittleness development varying among cultivars and being strongly influenced by water status during maturation, where water-stressed vines develop more brittle rachis earlier in ripening. The structural basis for rachis brittleness involves the proportion and distribution of lignified vascular and sclerenchyma tissues, the degree of cell wall cross-linking, and the moisture content providing plasticization, with brittleness increasing as lignin content rises and water content falls. The spatial distribution of brittleness along the rachis affects shattering patterns, with lateral branch junctions often representing fracture initiation sites where stress concentrations combine with tissue discontinuities. Measurement approaches include bending tests to failure on rachis segments, impact testing simulating mechanical harvest forces, and acoustic crack detection during controlled loading. Understanding rachis brittleness quality enables optimization of mechanical harvest timing, prediction of shattering losses under specified handling conditions, and variety selection for mechanical harvestability based on rachis mechanical development.
Rachis_Brittleness_Quality specifically addresses the cluster support structure, distinguishing from Pedicel_Brittleness_Quality affecting individual berry attachment. This quality is critical for mechanical harvest suitability assessment and represents a key factor in harvest method decision-making. The class supports harvest timing optimization and mechanical harvester setting adjustment based on rachis condition assessment.
Rachisbrüchigkeit
Stielgerüstfragilität
Traubengerüstsprödigkeit
Traubenstielsprödigkeit
bunch framework brittleness
cluster axis brittleness
cluster stem brittleness
rachis fragility
Intrieri C et al. (2008) Yield efficiency of high quality wine grape production in the Po Valley. Acta Horticulturae 785: 309-316 Cargnel MD et al. (2011) Shaking frequency and amplitude effects on the mechanical harvesting of wine grapes. Transactions of the ASABE 54(6): 2033-2038
Rachis Brittleness Quality
Cabernet Sauvignon in Napa Valley shows rachis development from late April through late May Year N+1 during inflorescence emergence at BBCH 53-59, with rachis tissues differentiating and expanding from 0.6-0.8 mm initial diameter to 2.0-2.8 mm at flowering, establishing vascular capacity of 3-5 mg per second water transport, moderate lignification creating flexible but strong structure, under warm climate conditions supporting cluster development in this vigorous large-clustered late-season cultivar producing robust rachis architecture supporting 200-400 gram clusters through development and harvest
Rachis development represents the structural maturation process where cluster axis tissues complete differentiation and achieve functional capacity to support flowers during anthesis and subsequently developing fruit through season, establishing mechanical framework and transport pathways essential for successful reproduction and berry development. This foundational developmental process occurs throughout inflorescence emergence spanning 3-5 weeks from initial visibility through flowering preparation, with rachis tissues progressively differentiating, thickening, and lignifying establishing permanent cluster structure. Rachis structure includes primary axis extending from peduncle attachment to cluster apex providing main structural support, primary branch rachis extending laterally from main axis to flower attachment points, and secondary branch rachis where present in compound architectures creating higher-order framework, with complete rachis system determining cluster mechanical properties and resource distribution. Anatomical development involves establishment of central vascular core with xylem vessels differentiating to provide water transport capacity of 2-5 mg per second per cluster adequate for flower and developing berry demands, phloem sieve elements forming enabling bidirectional assimilate transport, surrounding parenchyma providing storage capacity and maintaining live tissue function, peripheral cortex and strengthening tissues providing flexibility during development transitioning to rigidity at maturity, and epidermis with cuticle providing protection and controlling water loss. Cell division in developing rachis shows activity concentrated in intercalary meristems contributing to elongation during emergence, and in cambial zones producing vascular tissues and secondary growth, with division rates declining as maturation proceeds and elongation completes. Cell expansion throughout rachis contributes to thickness increase from initial 0.5-1.0 mm diameter at visibility to 1.5-3.0 mm at flowering depending on cultivar and cluster size, with expansion establishing final dimensions before lignification creates permanent structure. Lignification begins in rachis tissues during late emergence as secondary walls deposit with lignin incorporation creating mechanical strength, proceeding from base toward apex in basipetal pattern, with lignification extent at flowering variable among cultivars affecting rachis brittleness and breakage susceptibility. Mechanical properties develop progressively from flexible herbaceous structure during early emergence to increasingly rigid framework at flowering, with tensile strength increasing from 10-20 MPa in elongating tissues to 30-60 MPa in mature rachis providing resistance to fruit load and handling. Vascular capacity increases during development as vessel number and diameter increase enhancing water transport, with final capacity at flowering adequate for supporting anthesis water demands and prepared for substantial increases required during berry expansion. Resource transport through rachis includes upward water movement through xylem from shoot to flowers and berries, bidirectional assimilate movement through phloem with import during development and export during late berry ripening as seeds senesce, and hormone transport coordinating cluster development including auxin from developing seeds affecting fruit growth. Rachis function during flowering includes structural support maintaining cluster position and flower orientation facilitating pollination, water delivery to flowers supporting turgor during anthesis and stigma receptivity, and nutrient supply supporting active metabolism during reproductive processes. Rachis importance after fruit set intensifies as structure must support increasing berry weight reaching 100-500 grams per cluster depending on size, provide water delivery supporting berry expansion consuming 80-95% cluster water use, enable assimilate import during berry growth accumulating sugars and supporting metabolism, and maintain integrity through harvest preventing berry loss from rachis detachment or breakage. Rachis browning or necrosis represents physiological disorder reducing rachis function with tissue death interrupting transport causing berry shriveling and reduced quality, with disorder associated with calcium deficiency, water stress, or excessive berry load exceeding rachis capacity. Rachis lignification extent affects brittleness with highly lignified rachis showing rigidity and brittleness increasing shattering risk during harvest, while less lignified flexible rachis shows toughness reducing shattering but potentially increasing breakage under load. Rachis color development shows green coloration during active growth from chlorophyll in cortex tissues, transitioning toward brown as maturation proceeds and chlorophyll degrades, with final color at harvest ranging from green in some varieties to brown or reddish in others affecting visual cluster appeal. Rachis phenolic content affects browning with high polyphenol oxidase activity causing rapid oxidative browning when tissues damaged, with browning affecting appearance though not necessarily function unless extensive necrosis develops. Temperature during rachis development affects differentiation and maturation rate with optimal 15-24°C supporting normal development, cool conditions slowing progression, and very warm conditions accelerating development potentially causing incomplete differentiation. Water availability affects rachis development with adequate moisture supporting normal cell expansion and differentiation establishing robust structure, while water stress limits development producing smaller weaker rachis susceptible to dysfunction. Nutrient availability particularly calcium affects rachis quality with adequate calcium supporting strong cell walls and reducing browning susceptibility, while deficiency increases disorder risk affecting berry quality. The process represents fourth aspect of inflorescence emergence focusing on structural framework rather than overall cluster development, occurring concurrently with other emergence phases but emphasizing architectural support function. Duration spans entire emergence period from visibility through flowering preparation approximately 3-5 weeks, with continued rachis function required throughout berry development until harvest. Cultivar variation appears in final rachis dimensions and strength with vigorous large-clustered varieties showing thick strong rachis while small-clustered types show thin delicate structure, lignification extent affecting mechanical properties, and browning susceptibility affecting appearance and quality. Regional timing shows rachis development throughout spring emergence period from April-May in warm climates, May-June in temperate regions, and May-June in cool areas, with continued function through summer berry development and harvest. Viticultural management considerations include recognizing rachis as critical support structure determining cluster handling quality, ensuring adequate calcium availability reducing browning risk, managing water and crop load preventing rachis stress and dysfunction, understanding that rachis capacity ultimately limits cluster size potential, and appreciating that rachis quality affects harvest efficiency and fruit quality. Research applications include analyzing rachis anatomical development and mechanical properties, investigating physiological basis of rachis browning disorders, evaluating management effects on rachis quality affecting fruit quality and handling characteristics, quantifying transport capacity and limitations in rachis affecting berry development, and assessing genetic variation in rachis characteristics affecting cluster architecture and quality potential.
Rachis_Development is fourth aspect of Inflorescence_Emergence, distinguished by focus on structural framework rather than overall cluster. Occurs throughout emergence period concurrently with Visibility, Elongation, and Expansion but emphasizes support function. Critical for cluster integrity and function through anthesis, berry development, and harvest. Rachis quality affects mechanical properties, transport capacity, and quality characteristics.
Gescheinsachsenentwicklung
Rhachisentwicklung
Stielgerüstbildung
Traubenstielentwicklung
cluster axis development
cluster stem maturation
rachis formation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 340-355 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 270-280 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Rachis Development
Berry radial symmetry in seedless Sultanina (Thompson Seedless) approaches the theoretical ideal, with transverse sections displaying circularity indices of 0.95-0.99 (where 1.0 represents perfect circular symmetry), because the absence of seeds eliminates the primary source of internal asymmetry this uniform radial geometry facilitates even drying during raisin production by ensuring consistent thickness and surface exposure throughout the berry circumference, contributing to the cultivar's dominance in global raisin production.
Radial_Symmetry_Quality represents the symmetry condition where a grapevine organ exhibits equivalent form when rotated around a central axis by specific angles, creating multiple planes of symmetry that intersect at that axis. This symmetry type, also termed actinomorphy or rotational symmetry, characterizes organs that appear similar from multiple viewing angles around their central axis. In grapevine, radial symmetry is most evident in berries and flowers. Berries with uniform seed distribution (typically four seeds arranged symmetrically) or seedless berries often display approximate radial symmetry around the polar axis connecting pedicel attachment to stylar scar, appearing circular in transverse section with similar form when viewed from any angle around this axis. Grapevine flowers exhibit pentamerous radial symmetry with five-fold rotational correspondence of petals, sepals, and stamens, a fundamental character of the Vitaceae family. The degree of radial symmetry depends on the uniformity of development around the central axis. Seed number asymmetry, uneven sun exposure during development, or mechanical compression from adjacent berries can reduce berry radial symmetry. Quantitative assessment of radial symmetry employs circularity indices for transverse sections, coefficient of variation of radial distances from the centroid, or Fourier analysis of outline shape. Developmental processes establishing radial symmetry involve uniform meristematic activity and isotropic expansion around the organ axis.
Radial_Symmetry_Quality describes rotational correspondence around a central axis, distinct from Bilateral_Symmetry_Quality which involves reflection across a single plane. The order of radial symmetry (the number of equivalent positions per rotation) may be specified grapevine flowers exhibit five-fold (pentamerous) radial symmetry. Berries typically show continuous radial symmetry (infinite-fold) when perfectly circular in cross-section, or approximate radial symmetry when nearly circular. PATO_0001325 (radial symmetry) provides the upper-ontology alignment. Assessment requires specification of the axis of rotation, typically the polar axis for berries and the floral axis for flowers.
Radialsymmetrie
Rotationssymmetrie
actinomorphic quality
aktinomorphe Qualität
circular symmetry
mehrfache Symmetrie
multi-plane symmetry
rotational symmetry
Galet P (1979) A Practical Ampelography, Cornell University Press pp.: 28-35 Endress PK (1999) Symmetry in Flowers: Diversity and Evolution. International Journal of Plant Sciences 160(S6): S3-S23
Radial Symmetry Quality
Chardonnay radial wood ray in 14-year trunk shows perpendicular orientation crossing 9-11 annual rings radially from cambium 28-36 mm inward toward heartwood, multiseriate structure 3-5 cells wide tangentially, procumbent ray cells 50-90 μm radial length with abundant ray-vessel pits facilitating lateral water and solute exchange, living parenchyma in outer 20-26 mm sapwood zone storing starch at 7-11% dry weight and providing active radial transport, ray facilitating lateral loading of photosynthate from phloem into xylem storage during post-harvest period and radial export of mobilized sugars during spring budbreak
A radial wood ray is a wood ray emphasizing its radial orientation perpendicular to annual ring boundaries and extending from cambium toward trunk center, highlighting the ray's primary function as radial transport pathway connecting xylem depths with cambium and phloem. Radial wood ray orientation and anatomy include= radial alignment perpendicular to tangential plane (rays oriented 90° to annual ring curvature extending along radii from cambium toward pith), ray cells elongated radially (procumbent ray cells 40-120 μm radial length, 15-35 μm tangential width, 20-50 μm vertical height), multiple cell files wide (multiseriate rays 2-8 cells wide tangentially, typically 3-5 cells), ray extending vertically (ray height 5-40 cells, 200-1500 μm providing vertical integration along radial band), and ray crossing ring boundaries (radial rays extending through earlywood and latewood of multiple annual rings maintaining radial continuity). Radial wood ray transport functions include= radial water movement (lateral water transfer from xylem vessels through ray parenchyma to cambium and phloem particularly during spring budbreak), inward solute transport (minerals, hormones moving from phloem through rays into xylem storage parenchyma), outward solute transport (mobilized carbohydrates moving from xylem storage through rays to cambium and developing tissues), bidirectional flow capacity (rays enabling transport both inward and outward depending on source-sink relationships), and high permeability (numerous pits in ray cell walls particularly on ray-vessel boundaries facilitating solute exchange). Radial wood ray as lateral transport pathway includes= connecting cambium to inner sapwood (rays providing pathway for photosynthate loading from phloem into xylem parenchyma during reserve accumulation season), connecting storage sites to cambium (rays facilitating sugar export from stored reserves to cambium and developing buds during spring mobilization), enabling radial nutrient distribution (minerals absorbed by roots and transported in xylem moving laterally through rays to cambial zone for utilization), and providing hormone transport (auxin, cytokinin, abscisic acid potentially moving radially through ray parenchyma coordinating radial growth and development). Radial wood ray hydraulic connectivity includes= ray-vessel pits enabling lateral water flow (bordered and simple pits connecting ray parenchyma to vessel lumens allowing water redistribution), rays providing bypass routes (lateral connections enabling water to circumvent embolized or blocked vessels), radial hydraulic gradient (water potential differences between xylem depths and cambium driving radial flow particularly during transpiration), seasonal hydraulic dynamics (spring hydraulic continuity through rays supporting budbreak water demand, summer potential hydraulic segmentation as inner xylem embolizes), and limited refilling capacity (ray parenchyma potentially providing water for embolism repair though capacity limited in grapevine). Radial wood ray storage integration includes= ray parenchyma storing 20-35% of total trunk starch (rays providing substantial storage despite being 8-18% of wood volume due to high parenchyma density), radial gradient in storage concentration (outer rays in active sapwood showing 8-14% starch, inner rays approaching heartwood 2-6% starch), proximity to phloem facilitating loading (radial rays directly connecting to phloem ray continuations enabling efficient photosynthate transfer), radial mobilization pathway (stored starch in ray parenchyma mobilized and exported radially to cambium and buds), and seasonal storage dynamics (radial rays accumulating starch post-harvest, depleting during spring, replenishing during growing season). Radial wood ray anatomical integration with xylem includes= rays alternating with axial elements (radial ray bands separated by tangential bands of vessels, fibers, and axial parenchyma), ray density determining radial connectivity (6-12 rays per mm tangential providing radial pathways every 80-170 μm around circumference), rays crossing multiple ring boundaries (individual rays extending through 5-20+ annual rings depending on trunk age and ray origin), ray parenchyma living in sapwood (metabolically active cells in outer 15-30 mm providing functional radial transport), and ray parenchyma dying in heartwood (inner rays with dead cells no longer participating in transport but maintaining structural contribution). Radial wood ray identification and visualization includes= radial sections (longitudinal sections cut parallel to radius revealing rays as horizontal bands crossing vertical vessel elements), tangential sections (longitudinal sections cut tangential to circumference revealing ray end views showing height and cellular organization), cross sections (transverse sections showing ray dots where rays cut perpendicular to radial extent), and ray prominence (rays visible as lighter colored bands on radial surfaces due to different cellular composition compared to surrounding axial elements). Radial wood ray significance in trunk function includes= enabling efficient radial transport (short distance lateral movement through rays much faster than long-distance axial transport through vessels), facilitating rapid reserve loading (direct radial connection from phloem to xylem storage enabling efficient photosynthate transfer during 6-10 week accumulation period), supporting spring mobilization (radial export of mobilized sugars from storage to cambium and buds through short ray pathways), integrating hydraulic network (lateral connections through rays enhancing overall trunk hydraulic continuity), and coordinating radial zones (rays potentially enabling communication between inner xylem, outer xylem, cambium, and phloem coordinating growth and development).
Radial_Wood_Ray emphasizes radial transport function and is distinguished from Primary_Wood_Ray (origin and extent emphasis) and Secondary_Wood_Ray (initiation timing) by focus on perpendicular orientation and lateral transport role. Critical for understanding radial connectivity between trunk zones, lateral transport pathways complementing axial vessel transport, carbohydrate loading and mobilization mechanisms, and hydraulic integration through ray-vessel connections.
Cross-grain ray
Lateral parenchyma
Laterales Parenchym
Querverlaufender Strahl
Radialer Holzstrahl
Transverse ray
Pratt, C. (1974). Vegetative anatomy of cultivated grapes - a review. American Journal of Enology and Viticulture, 25(3), 131-150. | Tyree, M. T., & Zimmermann, M. H. (2002). Xylem Structure and the Ascent of Sap (2nd ed.). Springer.
Radial Wood Ray
Thompson Seedless (Sultana) bears a Raisin_Production_Scion_Role as the world's dominant raisin variety (over 90% of California production, major shares globally), with its combination of seedlessness, thin skin enabling 14-21 day sun-drying, high sugar content (22-24°Brix at optimal raisin harvest), and development of characteristic amber-brown color and caramel-fruity flavor during dehydration. Fiesta serves in a Raisin_Production_Scion_Role as an improved raisin variety in California, with earlier maturity, higher productivity, and equivalent drying quality to Thompson Seedless. Black Corinth (Zante Currant) bears a Raisin_Production_Scion_Role for currant production, with its tiny, completely seedless berries drying rapidly to produce the small, intensely flavored dried currants used in baking.
A scion role that inheres in a grafted scion cultivar by virtue of its designation for dried grape (raisin) production, where harvested grapes are dehydrated to approximately 13-16% moisture content, creating shelf-stable products with concentrated sugars (65-75% by weight), extended storage life (12-24+ months), and distinctive caramelized, dried fruit flavor profiles. Raisin production scions are selected for characteristics optimizing drying efficiency and dried fruit quality, including high initial sugar content (22-26°Brix at harvest, above table grape standards), thin skin permitting rapid moisture loss during drying (reducing drying time from 2-4 weeks to 1-2 weeks), small-to-medium berry size (accelerating drying by increasing surface-to-volume ratio), seedlessness (eliminating consumer inconvenience in the finished product), appropriate skin color (developing attractive brown or golden tones during drying), and flavor compounds that intensify rather than degrade through dehydration and Maillard reactions. Global raisin production exceeds 1.2 million tonnes annually, concentrated in California (USA), Turkey, Iran, China, Chile, South Africa, and Australia, with Thompson Seedless (Sultana) dominating world production.
Raisin_Production_Scion_Role is subclassed by drying method and product type (Natural, Golden, Currant, Sultana), reflecting distinct production processes and market categories. The raisin industry developed historically around naturally sun-dried grapes in hot, dry climates (California, Mediterranean, Middle East), with modern production including controlled-temperature dehydration and chemical pre-treatments (dipping in potassium carbonate/oil emulsions to accelerate drying). Raisin varieties must tolerate extended drying periods without mold development, requiring production in regions with reliable hot, dry autumn weather or investment in mechanical dehydration infrastructure. The category overlaps substantially with table grape production, as Thompson Seedless serves both markets depending on harvest timing and market conditions.
Rosinen-Traube
Rosinenrebsorte
Trockentraube
drying grape
raisin grape
raisin variety
Christensen LP (2000) Raisin Production Manual, University of California ANR Publication 3393 Creasy GL, Creasy LL (2009) Grapes, CABI Pangavhane DR et al. (1999) Review of research and development work on solar dryers for grape drying, Energy Conversion and Management 40: 1-23 OIV (2019) Statistical Report on World Vitiviniculture
Raisin Production Scion Role
The RAPD marker data published in early grapevine molecular studies (Büscher et al. 1993, Ye et al. 1998, Stavrakakis and Biniari 1998) employed 10-mer oligonucleotide primers from the Operon Technologies series (e.g., OPA-01 through OPA-20, OPB series) to amplify anonymous genomic fragments from cultivar DNA — a typical study screened 40-100 random primers, identified 10-30 primers generating reproducible polymorphic banding patterns, and scored 50-150 polymorphic fragments to construct UPGMA dendrograms of genetic similarity among 20-50 cultivars. RAPD analysis of Greek cultivars (Stavrakakis and Biniari 1998) using 28 primers generated 187 scorable bands of which 134 (72%) were polymorphic, distinguishing all 24 tested cultivars and placing them into groups broadly consistent with geographic origin and ampelographic classification.
A genetic marker data record for random amplified polymorphic DNA (RAPD) markers generated in Vitis through PCR amplification with short arbitrary primers (typically 10 bp), documenting the primer sequence, amplified fragment sizes, and presence/absence polymorphism patterns that provided the earliest PCR-based approach to grapevine molecular characterization before being largely superseded by more reproducible marker systems.
RAPD_Marker_Data documents the first generation of PCR-based molecular markers applied to grapevine, representing an important historical milestone despite the technology's well-documented limitations. RAPDs were introduced by Williams et al. (1990) and Welsh and McClelland (1990) as the simplest possible PCR-based genotyping approach — requiring only short arbitrary primers, standard PCR conditions, and agarose gel electrophoresis — making them accessible to any laboratory with basic molecular biology equipment at a time when SSR development required expensive genomic library construction and sequencing: this accessibility drove rapid adoption for grapevine cultivar studies in the early-to-mid 1990s, producing a substantial body of diversity and identification literature. However, the fundamental reproducibility problems of RAPD markers — sensitivity to DNA quality, polymerase brand, thermal cycler calibration, and primer lot variation, all stemming from the low-stringency annealing conditions required for 10-mer primers — made cross-laboratory data comparison unreliable, and the dominant inheritance (shared with AFLPs) prevented heterozygote detection. By the late 1990s, SSR markers had demonstrably superior reproducibility, codominant inheritance, and database compatibility, and RAPD genotyping was progressively abandoned in grapevine research. RAPD data are retained in the VVO as a historical marker class because early cultivar diversity studies based exclusively on RAPDs remain cited in the literature and their data occasionally require interpretation.
RAPD fingerprint data
RAPD marker specification
RAPD-Fingerabdruck-Daten
RAPD-Marker-Daten
random amplified polymorphic DNA record
zufällig amplifizierte polymorphe DNA-Daten
Williams JGK et al. (1990) DNA polymorphisms amplified by arbitrary primers are useful as genetic markers, Nucleic Acids Research 18: 6531-6535 Büscher N et al. (1993) Utilization of the RAPD technique for the identification of grapevine cultivars, Vitis 32: 165-169 Stavrakakis MN and Biniari K (1998) Genetic study of grape cultivars belonging to the Muscat family by random amplified polymorphic DNA markers, Vitis 37: 75-80
RAPD Marker Data
In a Sonoma Zinfandel vineyard, rapid acidity decline was documented over 16 days in late August to early September Year N+1 (BBCH 84-85) when titratable acidity decreased from 10.2 to 4.8 g/L (total reduction 5.4 g/L, average 0.48 g/L per day, 3.4 g/L per week), pH increased from 3.28 to 3.62 (0.024 units per day), malate declined from 6.4 to 2.1 g/L, tartaric stable at 6.8 g/L, berry Brix rose from 22.5° to 28.2°, under warm California conditions (daily high 29-33°C) necessitating harvest at target Brix to preserve minimal acidity for balance.
Rapid_Acidity_Decline represents the steepest gradient of total berry acidity reduction when titratable acidity and pH change most dramatically creating the most visible and measurable transformation of berry composition from acidic toward balanced or low-acid state defining harvest quality parameters and wine style potential. This rapid decline occurs during mid-to-late veraison through early ripening (BBCH 84-85), typically 14-26 days post-veraison onset depending on cultivar and environmental conditions, and is characterized by titratable acidity decreasing from 8-14 g/L at rapid onset to 4-8 g/L at conclusion over 12-20 day period representing 3.5-6.5 g/L total reduction and 40-60% depletion of entry values, daily TA decline rates 0.35-0.60 g/L per day (2.5-4.2 g/L per week) creating steep trajectory visible in sequential sampling data, pH increasing from 3.2-3.5 to 3.4-3.8 over rapid period representing 0.15-0.35 unit elevation at 0.020-0.040 units per day, and malate:tartrate ratio shifting from 0.6-1.3:1 to 0.2-0.7:1 indicating preferential malate loss driving acidity changes. Compositional changes during rapid decline show malate declining 4-8 g/L to 1.5-4.5 g/L representing 3-6 g/L loss accounting for 65-85% of total TA reduction, tartaric acid relatively stable 5-9 g/L showing only minor apparent changes 32-35°C causing excessive decline 0.60-0.85 g/L per day creating risk of over-deacidification pH >3.7-3.9 problematic for wine quality, and cumulative heat during rapid period (degree-days >28°C) strongly correlating with total acid loss enabling predictive modeling for harvest planning. Cluster microclimate heterogeneity produces substantial variation in rapid decline with sun-exposed clusters showing 50-100% faster decline than shaded fruit through elevated temperatures creating spatial variation across vineyard and within canopy, southwestern exposure (northern hemisphere) experiencing maximum decline through afternoon solar heating combined with diurnal accumulation, interior canopy positions maintaining 35-65% slower decline preserving acids, and canopy management creating 3-8 g/L differences in final TA between exposed and shaded zones requiring consideration in harvest strategy and wine blending. Water relations effects during rapid decline demonstrate adequate vine water status supporting normal metabolic function enabling appropriate decline without stress effects, moderate deficit (-1.0 to -1.4 MPa stem ψ) showing variable effects across cultivars with some acceleration through stress-enhanced respiration balanced by potential cooling from reduced transpiration, severe deficit (3.6-3.8 creating flat wines lacking structure with reduced color stability (reds), increased oxidation susceptibility, and elevated microbial spoilage risk requiring careful winemaking, high final acidity TA >9-11 g/L pH 3-4 g/L supporting complete rapid fermentation, 2-3 g/L enabling slower complete MLF potentially desirable for control, 3.9 inhibitory creating challenges. Harvest timing during rapid decline requires frequent monitoring 2-3 times per week as compositional changes rapid with 3-5 day difference producing substantially altered acid profiles, integration with sugar accumulation assessment ensuring appropriate sweetness-acidity balance achieved simultaneously, phenolic maturity evaluation particularly reds confirming tannin development synchronized with acid-sugar evolution, sensory assessment verifying flavor ripeness coordinated with analytical maturity, and flexibility in harvest date valuable as timing during rapid decline critically affects final composition enabling optimization for target wine style. Viticultural management includes canopy practices with strategic cluster shading most effective during rapid decline reducing temperature 4-9°C slowing decline 30-60% preserving acids critical in warm regions or low-acid cultivars, irrigation potentially moderating decline through berry cooling though mechanisms complex requiring cultivar-specific optimization, crop load management with appropriate fruit-to-leaf ratios preventing stress-accelerated decline while enabling normal rates, disease control essential as soft high-sugar berries vulnerable to Botrytis and sour rot requiring preventive protection, and harvest preparation advancing as rapid decline indicates approaching optimal maturity requiring readiness for prompt picking. Monitoring strategies involve laboratory analysis every 3-5 days tracking TA, pH, malate, tartrate enabling trajectory visualization and harvest forecasting, portable meters for field pH and TA screening guiding detailed sampling frequency and locations, spatial sampling across vineyard zones revealing heterogeneity requiring zone-specific management decisions, degree-day accumulation models predicting expected decline based on weather forecasts enabling proactive harvest scheduling, sensory evaluation confirming analytical findings ensuring acid-driven tartness declining appropriately coordinating with sugar and flavor development, and historical data compilation enabling comparison across seasons identifying vintage effects and refining management strategies.
Rapid_Acidity_Decline is distinguished from Peak_Acid_Reduction_Rate by representing measurable compositional change rather than velocity metric, from Maximum_Malate_Breakdown by focusing on total acidity rather than malate-specific, and from Initial_Acid_Reduction by showing steep gradient (0.35-0.60 g/L per day) rather than early moderate phase (0.18-0.35 g/L per day). Duration 12-20 days BBCH 84-85. TA reduction 40-60% of entry values. pH increase 0.15-0.35 units. Critical period determining wine acidity and style. Rapid_Acidity_Decline represents most visible compositional transformation during ripening affecting harvest timing and quality.
accelerated TA decline
beschleunigte TA-Abnahme
fast acidity decrease
rapid deacidification
rapide Entsäuerung
schnelle Azidität-Abnahme
steep total acid reduction
steile Gesamt-Säure-Reduktion
Keller M (2020) The Science of Grapevines 3rd Ed pages 229-233 rapid acidity decline Coombe BG (1992) Research on development and ripening of the grape berry Amer J Enol Vitic 43:101-110 compositional changes during ripening Mpelasoka BS, Schachtman DP, Treeby MT, Thomas MR (2003) A review of potassium nutrition in grapevines with special emphasis on berry accumulation Aust J Grape Wine Res 9:154-168 acidity and pH relationships.
Rapid Acidity Decline
In a Sonoma Chardonnay vineyard, rapid Brix elevation was documented over 12 days in early September Year N+1 (BBCH 85) when weekly sampling showed steep progression: 18.2° → 21.8° → 25.6° (average 3.7° per week, 0.53% per day), total 7.4° gain during rapid phase, linear regression slope 0.62° Brix per day (r² = 0.96), refractive index increase 0.00087 units per day, enabling harvest forecast 15 days advance projecting 26° target achieved within 8 days (actual 7 days, excellent accuracy), under warm California coastal conditions (daily high 26-30°C optimal).
Rapid_Brix_Elevation represents the steepest gradient of soluble solids increase during peak accumulation when refractometric measurements show maximum daily or weekly increments creating the most dramatic sweetening trajectory that enables achieving commercial sugar levels within compressed timeframe defining critical period for harvest maturity development. This rapid elevation occurs during peak accumulation phase (BBCH 84-85), typically days 19-30 post-veraison spanning 8-14 days, and is characterized by daily Brix increase 0.52-0.88% per day representing 2-3 fold enhancement over average active phase rates, weekly measurements showing 3.6-6.2° Brix gain per week creating steep trajectory visible in sequential sampling, berry Brix rising from 17-21° at rapid onset to 23-28° at conclusion over 10-15 day period representing 5-9° total increase during rapid phase, and refractive index increasing 0.00075-0.00123 units per day (corresponding to Brix change) reflecting altered light refraction through juice due to dissolved hexoses. Physical basis of Brix elevation includes total dissolved solids increasing primarily through hexose accumulation with glucose and fructose collectively contributing 85-95% of measured Brix, minor contribution from tartaric acid (5-9 g/L relatively stable) and other organic acids (declining 3-7 g/L malate), amino acids, phenolics, minerals collectively 12-25 g/L contributing 10-15% of total Brix measurement, refractive index relationship where each 1° Brix corresponds approximately to 10 g/L dissolved solids and refractive index increase 0.00014 at 20°C, and osmotic potential relationship where 1° Brix approximately equals -0.10 to -0.12 MPa osmotic potential contribution creating gradient driving water import. Measurement methodology for tracking rapid elevation includes portable digital refractometer field measurements accurate ±0.2° Brix enabling rapid assessment, laboratory bench refractometer providing ±0.05° accuracy for precise tracking, temperature compensation essential as refractive index varies 0.00025 per °C requiring correction to standard 20°C, sampling protocol collecting 20-50 berries randomly distributed across cluster zones ensuring representative measurement, and juice extraction through crushing and filtering providing clear sample free of particulates that could interfere with refraction measurement. Temporal characteristics during rapid phase show acceleration from active phase typical 0.24-0.38% Brix per day to rapid maximum 0.60-0.95% per day over 3-5 days as conditions optimize, sustained maximum rates 0.55-0.85% per day for 6-11 days while favorable conditions persist, then deceleration as substrates deplete or environmental conditions change transitioning toward late accumulation 32-35°C potentially impairing elevation 30-60% or causing irregular pattern, and stable favorable weather over 10-15 day period enabling sustained rapid elevation whereas variable conditions may produce irregular pattern. Cultivar genetic variation demonstrates high-Brix varieties (Gewürztraminer 26-30° potential, Sémillon 24-28°) achieving rapid elevation 5.5-7.5% Brix per week sustained 8-12 days, moderate cultivars (Chardonnay, Cabernet Sauvignon 22-26°) showing 4.2-6.0% per week for 10-14 days, and lower-Brix types (Pinot Noir cool climate 20-24°, table grapes 18-22°) displaying 3.2-5.0% per week over 12-18 days reflecting genetic differences in maximum capacity and duration determining elevation potential. Predictive modeling using rapid elevation data enables extrapolation from current Brix and measured rapid rate projecting harvest date 10-18 days advance, degree-day accumulation models calibrated during rapid phase improving forecast accuracy, recognition that rapid elevation typically concludes 1.5-3 weeks pre-harvest enabling harvest planning, and adjustment of projections if elevation rate changes indicating altered conditions requiring management response. Quality implications include rapid elevation timing and magnitude determining whether target sugars achieved within available growing season, excessive rapid elevation potentially producing higher-than-desired sugars requiring earlier harvest accepting lower phenolic development or later harvest risking over-ripeness, inadequate rapid elevation maintaining lower sugars potentially acceptable for lower-alcohol styles but problematic if target levels not achieved, and coordination between rapid Brix elevation and concurrent acid decline affecting final balance with simultaneous rapid changes producing compressed maturation window requiring precise harvest timing. Harvest timing decisions based on rapid elevation involve recognition that steepest Brix increase signals approaching critical maturity period, increased sampling frequency during rapid phase (2-3 times per week) essential for tracking trajectory accurately, target Brix achievement projection enabling advance harvest scheduling coordinating labor and logistics, integration with acid, pH, phenolic, flavor assessment ensuring comprehensive maturity beyond just Brix, and flexibility to harvest promptly when optimal parameters achieved as 3-5 day delay during rapid phase can produce 1.5-3° Brix increase substantially altering composition. Viticultural management during rapid elevation includes maintaining optimal conditions through irrigation supporting metabolism, canopy health ensuring continued photosynthesis, disease prevention protecting fruit during vulnerability to infection as sugars rise, avoiding environmental stresses that could disrupt rapid phase, and harvest preparation advancing as rapid elevation indicates imminent maturity. Comparison with other maturity indices shows Brix elevation correlates strongly with hexose accumulation (r² = 0.94-0.98) validating refractometry as sugar proxy, weaker correlation with acid decline (r² = 0.62-0.78) indicating independent though concurrent processes, moderate correlation with berry softening (r² = 0.70-0.85) as both result from ripening but through distinct mechanisms, and phenolic maturity correlation variable (r² = 0.45-0.75) particularly reds where tannin development may lag or lead sugar requiring integrated assessment. Regional patterns show warm regions typically experiencing briefer more intense rapid elevation 6-10 days due to accelerated metabolism, cool regions displaying more gradual elevation 12-18 days allowing extended flavor development potentially beneficial for quality, vintage variation substantial with warm years showing earlier more intense elevation while cool vintages exhibit delayed moderate elevation affecting harvest timing and final composition, and site variation within regions where warmer exposed positions undergo rapid elevation 4-8 days earlier than cooler shaded sites creating spatial heterogeneity.
Rapid_Brix_Elevation is distinguished from Steady_Brix_Increase by showing maximum (0.52-0.88% per day) rather than moderate (0.22-0.35%) rates, from Rapid_Brix_Increase (Batch 34) by providing alternative perspective on same phenomenon, and from initial Brix increase by representing peak rather than early phase. Duration 8-16 days BBCH 84-85. Weekly gain 3.6-6.2° Brix. Steepest trajectory. Predictive value for harvest timing. Rapid_Brix_Elevation represents most dramatic sweetening period determining maturity achievement.
accelerated sugar concentration
beschleunigte Zucker-Konzentration
fast soluble solids rise
maximaler Brix-Gradient
maximum Brix gradient
schneller löslicher Feststoff-Anstieg
steep Brix increase
steiler Brix-Anstieg
Keller M (2020) The Science of Grapevines 3rd Ed pages 224-227 rapid Brix increase Coombe BG (1992) Research on development and ripening of the grape berry Amer J Enol Vitic 43:101-110 Brix elevation patterns Coombe BG, McCarthy MG (2000) Dynamics and environmental control of grape berry growth Australian J Grape Wine Res 6:73-83 rapid elevation dynamics.
Rapid Brix Elevation
In a Napa Valley Zinfandel vineyard, rapid Brix increase was documented over 16 days in late August to early September Year N+1 (BBCH 84-85) when berry Brix rose from 18.3° to 27.5° (average 4.6% per week peak rate), hexose accumulation 195 mM per week in vacuoles, berry fresh weight increased 14% to 2.1 g through osmotic water import, titratable acidity declined from 8.2 to 5.8 g/L, anthocyanins reached 2150 mg/kg skin, under warm conditions (daily high 30-33°C, moderate deficit stem ψ -1.25 MPa concentrating sugars).
Rapid_Brix_Increase represents the steepest gradient of soluble solids accumulation during berry ripening when sugar concentration rises at maximum rates creating the dramatic sweetening that transforms acidic green fruit to harvest-ready grapes defining wine alcohol potential and style. This rapid increase occurs during mid-to-late veraison through early ripening (BBCH 84-85), typically 8-18 days duration depending on cultivar and environmental conditions, and is characterized by berry Brix rising from 14-20° at onset to 22-28° at conclusion representing 3.2-5.0% increase per week (compared to pre-rapid phase 1.5-2.8% per week and post-rapid plateau 8-15, and minor contributions from other soluble compounds including organic acids (declining 10-25% during rapid Brix increase as malate degrades), amino acids, phenolics, and minerals collectively 15-30 g/L affecting total dissolved solids measurement. Refractometry principles underlying Brix measurement show: refractive index of berry juice proportional to total dissolved solids concentration with pure water having refractive index 1.3330 at 20°C, each 1° Brix increase representing approximately 10 g/L dissolved solids (primarily sugars in grape juice) increasing refractive index by ~0.00014, portable refractometers measuring critical angle of total internal reflection providing field Brix assessment accurate ±0.2° with temperature compensation, and laboratory refractometers achieving ±0.05° accuracy enabling precise tracking of rapid increase rates. Osmotic consequences of rapid Brix increase demonstrate: vacuolar osmotic potential declining from -1.8 to -2.4 MPa at rapid onset to -2.6 to -3.4 MPa at conclusion creating strong water potential gradient, total berry water potential following vacuolar sugar causing decline from -1.5 to -2.0 MPa to -2.2 to -3.0 MPa establishing gradient favoring water import from vine xylem (-0.5 to -1.2 MPa typical), and osmotically-driven water uptake causing berry fresh weight increase 8-18% during rapid Brix increase despite declining growth rates illustrating continued expansion driven by sugar accumulation. Cellular hydration dynamics during rapid increase show: mesocarp cell water content relatively stable 75-82% fresh weight despite massive sugar accumulation indicating concurrent water import maintaining hydration, vacuolar volume fraction increasing slightly from 93-95% to 95-97% of cell volume accommodating both sugar and water, and cytoplasmic sugar concentration maintained relatively low 60-100 mM through efficient tonoplast transport preventing osmotic dehydration of cytoplasm that could disrupt metabolism. Environmental factors strongly affecting rapid Brix increase inclu temperature with optimal 26-30°C supporting maximum rates 4.2-5.0% per week through enhanced enzyme activities, membrane transport, and metabolism, moderate warmth 23-26°C enabling 3.5-4.5% per week still approaching maximum, cooler conditions 20-23°C reducing to 2.8-3.8% per week through slowed kinetics, heat stress 32-35°C impairing increase to 2.0-3.2% per week through enzyme denaturation and metabolic disruption, and extreme heat >35°C potentially arresting increase <1.5% per week or completely through catastrophic cellular dysfunction. Light effects operate through source photosynthesis with adequate canopy light exposure essential for producing sucrose substrate supporting rapid accumulation, shaded canopies limiting increase 20-40% through reduced photosynthesis despite adequate berry import capacity, and cluster exposure having minimal direct effects on Brix increase once substrate adequate though excessive radiation with heat can impair through berry temperature stress. Water relations influences show: moderate deficit (stem ψ -1.0 to -1.4 MPa) compatible with or enhancing Brix increase through reduced berry expansion concentrating sugars (higher Brix at same hexose content), severe deficit (<-1.5 MPa) impairing increase 35-65% through impaired phloem delivery and metabolism, osmotic effects with high berry sugar requiring adequate vine water status to maintain gradient for continued water import supporting expansion, and precision deficit irrigation targeting enhanced concentration without metabolic impairment typically imposing -1.1 to -1.3 MPa during rapid increase period. Cultivar genetic variation determines: high-Brix varieties (Gewürztraminer potential 26-30° Brix, Sémillon 24-28°, Chenin Blanc 23-27°) achieving rapid increase rates 4.2-5.2% per week over 2-3 weeks reaching final levels, moderate cultivars (Chardonnay 22-26°, Cabernet Sauvignon 23-27°, Merlot 22-26°) showing 3.5-4.5% per week for 2.5-3.5 weeks, and lower-Brix types (Pinot Noir cool climates 20-24°, some table grapes 18-22°) displaying 2.8-3.8% per week for 3-4 weeks reflecting genetic differences in maximum sugar accumulation capacity and metabolic programming. Temporal patterns of rapid increase show: acceleration from initial increase 1.8-2.8% per week to rapid phase 3.5-5.0% per week over 3-6 days, sustained rapid rates 10-20 days during peak accumulation, then deceleration to late ripening 1.5-2.8% per week and plateau <0.8-1.5% per week as vacuolar concentrations approach equilibrium with phloem supply and feedback mechanisms limit further accumulation. Coordination with other ripening processes during rapid increase includes: berry softening approaching completion with 75-90% total firmness reduction achieving characteristic ripe texture, color development (reds) reaching 85-98% final anthocyanin levels with both processes ABA-coordinated, acid decline approaching minimum with malate respiratory consumption 70-90% complete reducing titratable acidity from 8-12 g/L to 4-7 g/L, and flavor/aroma compound synthesis intensifying with terpenes, esters, and other volatiles accumulating creating ripe fruit character. Quality implications of rapid Brix increase inclu rate and duration determining final sugar and thus wine alcohol potential with faster longer increase producing higher Brix enabling earlier harvest or lower-alcohol styles through earlier picking, uniformity of increase across berries and clusters affecting harvest maturity synchrony with heterogeneous increase requiring selective harvest or acceptance of compositional variation, balance between Brix and other quality parameters (phenolics, acids, flavors) defining optimal harvest window with ideal showing coordinated maturation, and environmental conditions during rapid increase substantially affecting wine style through effects on final berry composition beyond just sugar level. Harvest timing based on rapid Brix increase involves: weekly sampling during rapid phase tracking rates and projecting timing to reach target Brix (typically 20-24° for table wines, 22-26° for fuller styles, 24-28° for late harvest depending on variety and region), integration with acid and pH monitoring ensuring appropriate balance, phenolic maturity assessment confirming tannin and color development synchronized with sugar, and sensory evaluation verifying flavor and aroma development adequate for desired wine style. Viticultural management during rapid increase includes: irrigation typically minimized or terminated as concentration generally desirable and berry water demand declining, canopy management maintaining source leaf health for continued photosynthesis without excessive leaf removal limiting substrate supply, disease monitoring critical as high-sugar berries increasingly attractive to pests and pathogens requiring protection, and harvest logistics preparation advancing as rapid increase indicates imminent maturity requiring equipment, labor, and winery readiness. Post-rapid increase management involves: continued monitoring as Brix increase decelerates toward plateau ensuring optimal harvest timing, extended hang time decisions weighing benefits of continued flavor development against risks of over-ripeness, excessive alcohol, or weather-related losses, and selective harvest considerations if spatial variation in rapid increase rates creates maturity heterogeneity across vineyard zones requiring multiple picking passes.
Rapid_Brix_Increase is distinguished from Initial_Sugar_Increase by representing maximum (3.2-5.0% per week) rather than moderate (1.5-2.8%) rates, from Peak_Sugar_Import_Rate by focusing on concentration measurement rather than metabolic flux, and from sugar plateau by showing maximum rather than declining rates. Duration 8-18 days BBCH 84-85. Temperature optimum 26-30°C. Moderate deficit can enhance concentration. Rapid_Brix_Increase represents practical quality metric determining harvest timing and wine alcohol potential.
accelerated sweetening
beschleunigte Süßung
maximale Brix-Erhöhungs-Rate
maximum Brix elevation rate
rapid sugar concentration rise
rapider Zucker-Konzentrations-Anstieg
steep soluble solids increase
steiler löslicher Feststoff-Anstieg
Keller M (2020) The Science of Grapevines 3rd Ed pages 224-228 Brix increase and maturity Coombe BG, McCarthy MG (2000) Dynamics and environmental control of grape berry growth Australian J Grape Wine Res 6:73-83 accumulation dynamics Deluc LG, Quilici DR, Decendit A, Grimplet J, Wheatley MD, Schlauch KA, Mérillon JM, Cushman JC, Cramer GR (2009) Water deficit alters differentially metabolic pathways affecting important flavor and quality traits in grape berries of Cabernet Sauvignon and Chardonnay Bmc Genomics 10:212 environmental effects on Brix.
Rapid Brix Increase
Merlot in Bordeaux shows rapid cell expansion during late July-early August Year N+1 at BBCH 75-77, with cells enlarging 8.6-fold from 48 to 142 micrometers diameter, massive vacuoles occupying 86% cell volume, water uptake 1.4-2.0 g per berry per day, peak berry growth rate 140-190 mg per day, berry diameter expanding from 21 to 29 mm achieving 87% final size, fresh weight increasing from 9 to 17 g, under oceanic conditions of 22-28°C temperature with adequate summer rainfall, typical of this mid-season cultivar completing rapid enlargement
Rapid cell expansion represents the maximum enlargement phase during berry cell enlargement when established turgor-driven growth accelerates to peak rates, massive vacuole formation creates dominant 80-90% cell volume compartment, peak berry growth rate achieves maximum volumetric increase 100-250 mg fresh weight per day, cell volume increases 5-15 fold from initial enlargement completion, and berry approaches near-final dimensions, proceeding over weeks 5-8 post-fertilization as pure expansion-dominated growth following division completion. This maximum expansion phase occurs during late summer in late July-early August in temperate regions Year N+1, spanning approximately 2-4 weeks representing peak enlargement intensity, corresponding to BBCH 75-77 berry touch to beginning bunch closure stages. Rapid expansion mechanism involves maximum expansion rate with optimized turgor-wall system operating at peak capacity, massive vacuole formation with central vacuole occupying 80-90% cell volume, peak berry growth rate from coordinated cell enlargement across berry populations, sustained high water uptake 1.0-2.5 g per berry per day, and intensive enlargement establishing near-final berry size. Cell expansion acceleration shows enlargement rate increasing from initial phase levels to peak during rapid expansion, expansion rate proportional to turgor exceeding yield threshold, optimized wall loosening enabling maximum yielding, and peak rate reflecting optimal coordination of expansion mechanisms. Maximum expansion rate characteristics show individual cells enlarging at highest velocity, typical 5-15 fold volume increase during rapid expansion phase from 40-70 micrometers to 80-140 micrometers diameter, rapid volume accumulation over 2-4 weeks, and maximum rate representing peak growth capacity. Cell volume dynamics show exponential-like volume increase during rapid phase, typical doubling time 7-12 days during peak expansion, sustained rapid enlargement over weeks, and cumulative substantial volume gain. Turgor pressure maintenance shows sustained turgor 0.6-1.2 MPa during rapid expansion, continued solute accumulation maintaining osmotic gradient, continued water uptake sustaining pressure, and maintained turgor enabling sustained expansion. Osmotic potential during rapid expansion shows values -1.0 to -1.5 MPa from high solute concentrations, organic acids reaching peak levels 1.2-1.8% fresh weight, potassium accumulation to 180-250 mmol/L tissue, sugars beginning accumulation 5-8% though still relatively low, and strong osmotic gradient driving intensive water uptake. Water uptake peak shows maximum water influx rates 1.0-2.5 g per berry per day during rapid expansion, intensive water import supplying expanding cells, xylem as primary water source, possible phloem water contribution, and high uptake rate enabling rapid enlargement. Xylem water delivery shows functioning xylem vessels delivering water to berry, transpiration pull and root pressure driving flow, xylem dominance during growth phase, and vascular capacity supporting high water demand. Massive vacuole formation shows central vacuole expanding to occupy 80-90% mature cell volume, vacuole volume increase accounting for majority of cell volume gain, single large vacuole per cell typical in mature expanding cells, and vacuole as dominant cell compartment. Vacuole size during rapid expansion shows typical progression from 50-70% initial to 80-90% by rapid expansion completion, vacuole diameter 60-180 micrometers in mature cells, and massive size characteristic of fully expanded cells. Vacuole storage function shows compartment storing accumulated water, organic acids at peak concentrations, potassium and other ions, beginning sugar accumulation, and vacuole as major storage depot. Cytoplasm organization shows cytoplasm as thin 2-5 micrometer peripheral layer surrounding central vacuole, nucleus and organelles in peripheral cytoplasm, cytoplasmic streaming possibly distributing materials, and efficient cytoplasm organization in expanded cells. Cell wall during rapid expansion shows wall thinning from area increase faster than synthesis, wall thickness declining to 0.1-0.2 micrometers, maintained wall integrity preventing rupture, and wall capable of supporting large cell size. Wall area increase shows substantial surface area expansion during rapid phase, typical 5-15 fold area increase, coordinated wall deposition, and area increase requiring continued synthesis. Berry growth rate peak shows maximum berry fresh weight increase 100-250 mg per day in large-berried cultivars, sustained high growth rate over 2-4 weeks, growth rate reflecting coordinated cell enlargement, and peak as maximum growth capacity. Berry volumetric increase shows substantial expansion from 6-12 g initial to 10-20 g by rapid expansion completion in large-berried cultivars, typical 1.5-2.5 fold volumetric increase during rapid phase, and major size gain during intensive enlargement. Berry diameter increase shows expansion from 18-25 mm at rapid expansion onset to 24-36 mm by completion depending on cultivar, substantial diameter gain 6-12 mm during rapid phase, and berry approaching near-final dimensions. Berry size achievement shows berry reaching 80-95% final size by rapid expansion completion, near-final dimensions established, remaining growth during lag and ripening relatively limited, and rapid expansion as major size determinant. Berry composition during rapid expansion shows water content maintained 80-85% fresh weight, organic acids at peak 1.2-1.8%, sugars low to moderate 5-8%, pH 2.8-3.0, and hard acidic green berry characteristic. Cell enlargement spatial coordination shows enlargement across all berry cells creating uniform growth, tissue-level coordination maintaining berry shape and integrity, and coordinated expansion preventing structural problems. Environmental effects on rapid expansion include water availability as most critical factor with adequate water absolutely essential for maximum expansion, deficit severely limiting expansion reducing final size 20-50%, temperature affecting expansion rate with optimal 22-28°C supporting peak rates, and favorable conditions maximizing size achievement. Water deficit effects show deficit during rapid expansion as most critical period for size limitation, reduced water availability preventing full turgor and expansion, deficit effects on final size substantial and irreversible, and water as limiting factor during intensive enlargement. Temperature effects show warm conditions promoting maximum expansion rate, cool conditions slowing expansion though generally not preventing substantial enlargement, and temperature affecting rate and timing rather than potential. Irrigation importance shows irrigation during rapid expansion critical in water-limited regions, supplemental water enabling full size potential achievement, and irrigation as key viticultural practice for size optimization. The process shows three distinguishable aspects including maximum expansion rate with peak cell enlargement velocity, massive vacuole formation creating dominant compartment, and peak berry growth rate achieving maximum size increase. Duration typically 2-4 weeks representing intensive enlargement weeks 5-8. Regional timing shows rapid cell expansion during late July-early August across climates as peak expansion proceeds. Viticultural significance includes representing maximum growth phase establishing near-final berry size, requiring optimal water availability for size achievement, demonstrating critical environmental sensitivity during peak enlargement, and determining berry size affecting yield and quality. Research applications include analyzing maximum expansion mechanisms and regulation, investigating water relations during peak uptake, evaluating vacuole formation and function, quantifying expansion rate dynamics and determinants, and assessing rapid expansion as major berry size determinant.
Rapid_Cell_Expansion is second and final phase of Berry_Cell_Enlargement_Phase, distinguished by maximum enlargement intensity. Subdivided into Maximum_Expansion_Rate with peak cell velocity, Massive_Vacuole_Formation creating 80-90% volume compartment, and Peak_Berry_Growth_Rate achieving maximum increase. Duration 2-4 weeks representing weeks 5-8. Cell volume 5-15 fold increase. Turgor 0.6-1.2 MPa sustained. Water uptake 1.0-2.5 g/berry/day. Berry achieves 80-95% final size. Water availability most critical factor. Follows Initial_Cell_Enlargement. Completes Berry_Cell_Enlargement_Phase.
Spitzen-Expansion
intensive Vergrößerung
intensive enlargement
maximales Zellwachstum
maximum cell growth
peak expansion
rapid growth phase
schnelle Zellexpansion
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 446-450 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Rapid Cell Expansion
Cabernet Sauvignon rapid division stage berries at weeks 2-4 post-fruit set (BBCH 71-73) show high mitotic index progressing from 12% (week 2, 3.2 mm diameter, 0.025 g, 1.2 million cells, 38 μm cell diameter) through 16% at peak (week 3, 4.8 mm, 0.09 g, 1.7 million cells, 45 μm) to 13% (week 4, 6.5 mm, 0.25 g, 2.3 million cells, 52 μm), with cell number increasing at 0.35-0.55 million per week representing doubling time of 10-14 days during peak rapid division, and with berry diameter increasing at 0.8-1.1 mm per week and fresh weight increasing at 0.03-0.08 g per week demonstrating rapid growth driven primarily by cell proliferation. Syrah rapid division stage berries examined through flow cytometry and immunohistochemistry show weeks 2-3 with 14-17% of mesocarp cells in S phase (DNA synthesis) and 3-4% in M phase (mitosis visible microscopically), with cyclin B immunostaining showing 12-15% positive cells concentrated in mid-to-inner mesocarp regions, demonstrating active cell cycle progression throughout mesocarp tissue during rapid division with spatial patterns showing most active division in central berry regions. Merlot rapid division stage berries under three crop load treatments show= low crop load (4 clusters per shoot) producing rapid division berries with 16-18% mitotic index and 0.5-0.65 million cell increase per week moderate load (7 clusters per shoot) showing 13-15% mitotic index and 0.38-0.48 million per week high load (11 clusters per shoot) showing 10-12% mitotic index and 0.28-0.38 million per week, demonstrating crop load effects on division rate during rapid division stage through competition for resources, with low-load berries achieving higher final cell numbers (2.8-3.2 million at BBCH 75) versus high-load berries (2.1-2.5 million) creating 15-20% final size difference. Pinot Noir rapid division stage berries experiencing water deficit during weeks 2-4 show reduced mitotic index (9-11% versus well-watered 14-16%) and reduced cell number increase rate (0.25-0.35 million per week versus 0.42-0.55 million), resulting in reduced final cell number (2.0-2.3 million versus 2.6-3.0 million) and permanently smaller mature berries (11.5-13 mm versus 13.5-15 mm), demonstrating that water deficit during rapid division stage creates lasting size reduction through impaired cell proliferation.
A rapid division stage berry is a cell division stage berry during the early-to-mid portion of Phase I (BBCH 71-73, approximately 1-4 weeks post-fruit set) when cell division activity is high and accelerating or at peak levels, characterized by high mitotic index, rapid cell number increase, and rapid berry expansion driven primarily by cell proliferation, representing berries undergoing maximum cell division activity. Rapid division stage berries are characterized by= (1) high mitotic index typically 10-17% of mesocarp cells in M phase during weeks 2-4, representing peak or near-peak division activity (2) rapid cell number increase at rates of 0.25-0.5 million cells per week during peak activity, with cell population doubling approximately every 1-2 weeks during rapid division (3) cell number progression from approximately 0.8-1.5 million cells at early rapid division (week 1-2, BBCH 71-72) to 1.8-2.6 million cells by late rapid division (week 4, BBCH 73) (4) cells dividing multiple times during rapid division stage with individual cells completing 2-4 division cycles during the 3-4 week period (5) berry diameter increasing rapidly at 0.6-1.2 mm per week during rapid division, expanding from 2.5-4 mm at early rapid division to 5-8 mm at late rapid division (6) berry fresh weight increasing rapidly at 0.015-0.08 g per week, expanding from 0.012-0.035 g to 0.12-0.35 g during rapid division stage (7) cells showing modest concurrent expansion from 30-42 μm diameter to 42-58 μm, with simultaneous division and expansion though division predominates. Rapid division stage berries show maximum growth vigor with high metabolic activity supporting cell proliferation= high respiration rates (35-55 nmol CO₂ g⁻¹ s⁻¹) providing energy for biosynthesis active import of photosynthates from leaves supporting cell wall synthesis and cytoplasm production for new cells high expression of cell cycle genes (cyclins, cyclin-dependent kinases, DNA synthesis genes) driving division active auxin and cytokinin signaling promoting cell division. Rapid division stage berries are maximally responsive to environmental and cultural factors affecting cell division= water availability has strong effects with adequate water supporting maximum division rates while deficit reduces division creating permanent size reduction temperature influences division rate with optimal temperatures 22-28°C supporting maximum division versus cooler temperatures slowing division nutrition particularly nitrogen availability affects division vigor with adequate nitrogen supporting rapid division crop load influences resource availability per berry affecting division rate with low crop loads supporting faster division and higher final cell numbers. Rapid division stage berries represent the period of maximum opportunity for manipulating final berry size through cultural practices, as interventions affecting cell division during rapid division stage (irrigation, crop thinning, vigor management) have maximum impact on final cell number and therefore final size potential.
Rapid_Division_Stage_Berry represents peak cell division activity (weeks 1-4, BBCH 71-73) and is distinguished from Peak_Division_Stage_Berry which likely represents the absolute peak (weeks 2-3) within the rapid division period. Rapid division stage represents the period of maximum growth rate and maximum sensitivity to factors affecting berry size through cell division. Understanding rapid division stage biology is critical for timing cultural practices (irrigation, crop adjustment) to maximize or moderate berry size through effects on cell proliferation during this high-activity period.
Hochproliferations-Beere
Maximal-Zellteilungs-Stadium
high-division berry
peak-mitotic berry
schnelle Teilungs-Stadium-Beere
Rapid Division Stage Berry
Chardonnay grafted on Teleki 5C rootstock showing rapid elongation zone on primary root tip, zone located 2.2-3.4 mm from root apex comprising 1.2 mm length in middle of total elongation zone 5.8 mm, time-lapse imaging revealing cells in rapid zone expanding at peak rate 6.8 micrometers per hour, rapid elongation zone contributing estimated 1.63 mm daily to total root elongation 4.2 mm per day, cells showing maximum turgor pressure 0.68 MPa, elevated metabolic activity evident through high respiration rate, rapid zone demonstrating peak expansion velocity determining overall root growth rate
Rapid elongation zone is the portion of elongation zone showing maximum cell expansion rate, typically located in middle section of elongation zone 1-4 mm from root apex, characterized by cells elongating at peak velocity potentially 2-10 micrometers per hour creating highest local contribution to root length increase, showing maximum turgor pressure 0.5-0.8 MPa driving maximal wall extension, demonstrating optimal balance between wall loosening allowing expansion and wall synthesis maintaining integrity, and representing the region where cell elongation rate peaks after gradual acceleration from meristem then declines toward maturation zone creating elongation velocity gradient along root axis, with rapid elongation zone cells showing active expansin protein expression loosening wall structure, elevated metabolic activity supporting expansion, maximal ion uptake generating osmotic potential, and high water influx creating turgor, and demonstrating that this zone of maximum elongation rate determines overall root growth velocity with longer or more active rapid elongation zones producing faster-growing roots. Rapid elongation zone position and extent includes= located mid-elongation zone (rapid elongation typically 1-4 mm from root apex in middle portion of total elongation zone), showing variable length (rapid zone typically 0.5-2 mm length depending on root growth rate), exhibiting peak elongation rate (cells in rapid zone expanding fastest among all elongation zone cells), demonstrating gradual transitions (smooth acceleration from proximal elongation zone and deceleration toward distal zone), and revealing order-dependent patterns (primary roots showing longer rapid elongation zones than fine roots reflecting vigor differences). Rapid elongation zone expansion velocity includes= showing maximum elongation rate (cells potentially expanding 2-10 micrometers per hour during peak growth), exhibiting rapid cumulative growth (rapid zone potentially contributing 0.5-2 mm daily to total root elongation depending on zone length and cell number), demonstrating velocity gradient peak (elongation velocity reaching maximum in rapid zone declining proximally and distally), revealing root order effects (primary root rapid zones showing higher velocities than tertiary root rapid zones), and displaying environmental modulation (rapid zone velocity varying with temperature, water availability, and nutrients). Rapid elongation zone turgor characteristics includes= showing maximum turgor pressure (cells in rapid zone maintaining peak turgor 0.5-0.8 MPa highest in elongation zone), exhibiting active osmotic adjustment (continuous ion accumulation maintaining high osmotic potential as cell expands), demonstrating water influx (rapid water uptake matching expansion rate maintaining turgor despite increasing cell volume), revealing membrane transport (active H+-ATPases and ion channels driving solute accumulation), and displaying pressure maintenance (turgor pressure sustained throughout expansion period unlike transient pressure in slower zones). Rapid elongation zone wall properties includes= showing optimal wall loosening (expansin activity and pH changes creating maximal wall extensibility), exhibiting balanced synthesis (new wall material deposition matching expansion rate maintaining appropriate thickness), demonstrating mechanical properties (wall combining extensibility allowing expansion with strength preventing rupture), revealing cell wall chemistry (pectin matrix loosening and cellulose fibril reorientation during expansion), and displaying directional constraints (transverse cortical microtubules orienting cellulose deposition directing longitudinal expansion). Rapid elongation zone metabolic intensity includes= showing elevated respiration (rapid zone cells consuming oxygen at high rates supporting expansion metabolism), exhibiting high ATP demand (energy required for ion pumping, wall synthesis, and biosynthesis), demonstrating active protein synthesis (producing expansins, wall enzymes, membrane transporters, and structural proteins), revealing biosynthetic activity (synthesizing cell wall polysaccharides, membrane lipids, and other components), and displaying resource dependence (rapid zone requiring continuous photosynthate supply supporting intense metabolism). Rapid elongation zone hormone effects includes= responding strongly to auxin (indole-3-acetic acid promoting rapid elongation through wall loosening and gene expression), affected by gibberellins (gibberellins enhancing elongation particularly in combination with auxin), influenced by brassinosteroids (brassinosteroids promoting cell expansion and elongation), showing ethylene sensitivity (ethylene potentially inhibiting elongation under stress conditions), and demonstrating abscisic acid effects (ABA reducing elongation under water stress protecting cells from excessive expansion). Rapid elongation zone in growth control includes= determining peak growth rate (rapid zone elongation velocity representing maximum instantaneous root growth rate), controlling overall elongation (rapid zone length and activity strongly influencing total root elongation rate), demonstrating growth plasticity (rapid zone responding rapidly to environmental changes enabling growth adjustment), revealing stress sensitivity (rapid zone highly sensitive to water deficit, temperature extremes, or mechanical impedance), and showing management relevance (practices affecting rapid zone function directly impacting root growth and development). Rapid elongation zone termination includes= showing gradual deceleration (elongation rate declining in distal rapid zone as cells approach maturation), exhibiting wall rigidification (progressive cross-linking and secondary wall initiation reducing extensibility), demonstrating differentiation onset (cells beginning tissue-specific differentiation reducing expansion capacity), revealing turgor decline (osmotic potential and turgor potentially declining as maturation approaches), and displaying transition to maturation (smooth transition from rapid elongation to maturation zone marking end of expansion phase).
Rapid_Elongation_Zone emphasizes maximum expansion velocity region and is distinguished from slower elongation zones by focus on peak elongation rate. Critical for understanding maximum growth velocity, peak turgor pressure, optimal wall loosening, metabolic intensity, growth rate determination, and the zone of fastest cell expansion controlling overall root elongation rate.
Fast-growing zone
Maximale Streckungszone
Maximum elongation zone
Peak expansion region
Schnelle Elongationszone
Spitzenwachstumsbereich
Rapid Elongation Zone
A rapid expansion young leaf at node 9 on a highly vigorous shoot of Vitis vinifera cv. Grenache in Châteauneuf-du-Pape, France, exhibiting area increase from 32 cm² to 88 cm² over 7 days representing daily growth rate of 11.4 cm² per day (8.9% daily compound growth), dimensions increasing from 7.2 cm × 6.5 cm to 11.8 cm × 10.2 cm, cells in palisade layers enlarging from average 22 μm to 31 μm diameter over the measurement period representing 5.7 μm total increase or 0.81 μm per day, tissue water content of 90% fresh weight indicating high turgor supporting rapid expansion, very soft flexible texture easily deformed by gentle handling, rapidly deepening green coloration with chlorophyll increasing from 280 μg cm⁻² to 385 μg cm⁻² over the week, visible growth obvious from daily observation showing measurable dimension changes, consuming approximately 35 mg carbohydrate per day and 5.2 mg nitrogen per day supported by strong translocation from 18 mature leaves on the same shoot, photosynthetic rate increasing from 6.8 to 9.5 μmol CO₂ m⁻² s⁻¹ over the period approaching 72% mature capacity, occurring under optimal conditions including daily maximum temperatures 25-27°C, soil moisture maintained at 82% field capacity through irrigation, adequate nitrogen from 95 kg N/ha fertilization program, measured at shoot BBCH stage 73 (berries beginning to touch) in early July, representing maximum expansion rate typical of vigorous shoots under optimal resource availability and environmental conditions.
An expanding young leaf exhibiting maximum daily growth rates during optimal expansion conditions in Vitis vinifera, characterized by very high area increase rates typically 8-15 cm² per day representing 6-12% daily compound growth rate, dimensions progressing rapidly from approximately 25% to 65% of final size over 7-10 day period, active cell expansion throughout lamina with cells enlarging at maximum rates of 2-4 μm per day in linear dimensions, high tissue water content of 88-92% fresh weight reflecting active water uptake driving turgor-based expansion, soft pliable texture from loosened cell walls with high pectin content and minimal lignification allowing maximum extensibility, light green rapidly deepening coloration as chlorophyll synthesis accelerates keeping pace with area expansion, visible daily growth progression evident without measurement instruments, substantial daily resource consumption including 25-40 mg carbohydrate per day for cell wall synthesis and metabolic processes and 4-6 mg nitrogen per day for protein synthesis supporting membrane and enzyme production, and rapidly increasing photosynthetic capacity approaching 60-75% of mature leaf rates by end of rapid expansion phase creating transition toward carbon balance though remaining net sink throughout rapid expansion period. Rapid expansion young leaves develop under optimal conditions combining favorable temperatures of 23-28°C maintaining high metabolic rates and enzyme activity, abundant soil moisture maintaining 70-90% field capacity providing water for turgor-driven expansion without limitation, adequate nitrogen availability supporting protein synthesis requirements, moderate light intensity providing photosynthetic energy without causing photoinhibition of developing chloroplasts, and high shoot vigor with strong resource flows from mature leaves and reserves supporting demanding growth rates.
Rapid expansion young leaves are distinguished from slow expansion leaves by very high daily growth rates, rapid progression through expansion phases, and high resource consumption rates, requiring optimal environmental conditions and shoot vigor. Achievement of rapid expansion depends critically on water availability, as turgor-driven cell expansion requires continuous water uptake maintaining high tissue water content, with even brief water deficits immediately slowing expansion. Temperature strongly affects expansion rate with Q₁₀ of approximately 2.5, meaning 10°C temperature increase approximately doubles expansion rate within physiological range of 15-30°C. Rapid expansion represents efficient use of favorable growing conditions achieving maximum final leaf area in minimum time, creating maximum photosynthetic surface area quickly to support fruit development and subsequent growth. However, very rapid expansion under luxury resource availability may create excessively large leaves contributing to dense canopies requiring management through shoot thinning or positioning. Rapid expansion phase typically lasts 7-12 days under optimal conditions before transitioning to slower final expansion and maturation.
fast-growing leaf
high-velocity growth leaf
hochgeschwindigkeits Wachstum Weinblatt
maximum expansion rate leaf
schnell-expandierendes junges Blatt
schnellwachsendes Blatt
Based on Poni et al. (2009) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 60=397-407 Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Tsukaya (2013) "Leaf development" The Arabidopsis Book 11=e0163 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 137-160.
Rapid Expansion Young Leaf
In a Willamette Valley Pinot Noir vineyard in Oregon, rapid firmness loss was documented over 5 days in early September Year N+1 (BBCH 83-84) when compression firmness declined from 2.6 N to 1.5 N (5.3% per day average), puncture resistance dropped from 2.2 N to 1.3 N, soluble pectin increased from 48% to 71%, and berry fresh weight rose from 1.08 g to 1.28 g as sugar accumulated from 16.8 to 21.2° Brix under optimal conditions (daily high 24-27°C, nighttime low 12-15°C).
Rapid_Firmness_Loss represents the accelerated decline in berry mechanical resistance during peak softening, characterized by quantifiable reductions in compression force, penetration resistance, and elastic modulus that transform berry texture from firm to readily deformable within a concentrated timeframe. This process occurs during the central portion of peak softening (typically 3-5 days within mid-veraison BBCH 83-84) and is measured through multiple mechanical testing parameters: compression firmness declining from 2.2-3.2 N to 1.3-2.2 N (30-45% reduction), puncture resistance dropping from 1.8-2.8 N to 1.0-1.8 N (35-50% decrease), and elastic modulus falling from 250-400 kPa to 120-220 kPa indicating fundamental shift in cellular mechanical properties. The rapid firmness loss results from convergent structural modifications: cell wall pectin degradation produces 50-70% soluble pectin (from 30-45% pre-softening) with molecular weight reduction to 70-110 kDa (from 140-200 kDa), middle lamella dissolution causes 50-70% loss of intercellular adhesion creating cell separation and berry pliability, and hemicellulose modification particularly xyloglucan restructuring contributes 20-35% to overall firmness decline through loss of cell wall tensile strength. Cellular mechanical changes during rapid firmness loss inclu turgor pressure shows modest 10-20% reduction (from 0.5-0.8 MPa to 0.4-0.65 MPa) contributing 15-25% to total firmness loss, cell wall extensibility increases 3-5 fold allowing plastic rather than elastic deformation under compression, and skin tensile strength decreases 40-60% creating vulnerability to splitting and enabling easier berry compression. Enzyme activities responsible for rapid firmness loss reach sustained high levels: polygalacturonase maintains 90-180 units throughout the rapid loss period cleaving α-1,4-galacturonan linkages, pectin methylesterase operates at 100-180 units demethylating pectin and creating PG substrates, β-galactosidase activity (25-60 units) removes galactose side chains facilitating backbone access, and expansin activity promotes stress relaxation in cell walls enabling irreversible expansion. Temperature profoundly affects rapid firmness loss kinetics: optimal conditions (22-28°C) support maximum enzyme efficiency producing 3-6% firmness decline per day, cooler temperatures (16-22°C) reduce rates to 2-4% per day extending the rapid loss period, moderate heat (28-32°C) can accelerate loss to 4-8% per day but may compromise quality through disrupted metabolism, and excessive heat (>32°C) causes abnormal softening patterns through membrane disruption and cellular disorganization. Cultivar variation in rapid firmness loss characteristics reflects genetic differences in cell wall composition and enzyme complement: soft-fruited varieties like Pinot Noir exhibit very rapid firmness loss (4-7% per day) with high PG expression and thin cell walls losing 45-65% of initial firmness in 4-6 days, medium-firmness cultivars like Merlot show intermediate loss rates (3-5% per day) with moderate enzyme levels declining 35-50% over 5-7 days, while firm-fruited varieties like Cabernet Sauvignon display slower loss (2-4% per day) retaining relatively higher firmness with 30-45% decline over 6-9 days. Rapid firmness loss has significant viticultural implications: berries become increasingly susceptible to physical damage from handling, wind, rain, and harvest operations with compression damage threshold dropping from >3.5 N to <2.0 N, pathogen infection particularly Botrytis cinerea finds optimal entry as skin resistance declines and surface microcracking increases, and bunch compaction intensifies as softened berries deform to fill spaces within clusters potentially creating anaerobic conditions and split berry problems. Water relations during rapid firmness loss show complex interactions: moderate water deficit (leaf ψ -0.8 to -1.2 MPa) can slow firmness loss by 20-35% through reduced enzyme synthesis and slight turgor maintenance, excessive deficit (<-1.4 MPa) may paradoxically accelerate loss through cellular stress and membrane dysfunction, while abundant water supply can accelerate firmness loss through enhanced cell wall enzyme activity but risks berry splitting as internal pressure exceeds diminished skin strength. Molecular control involves sustained expression of ripening-related genes: continued high ABA levels (1200-2800 ng/g) maintain softening gene transcription, ethylene production shows cultivar-dependent slight elevation (0.3-1.8 nL g⁻¹ h⁻¹) in some varieties correlating with firmness loss rate, and sugar signaling through hexokinase pathways may feedback-regulate cell wall enzyme genes linking metabolite accumulation with texture transformation. Berry quality implications of rapid firmness loss timing and intensity inclu premature or excessive firmness loss before adequate flavor and aromatic compound development can produce soft berries with insufficient character, delayed firmness loss can result in harvesting overly firm fruit with incomplete textural development affecting wine mouthfeel, and optimal firmness loss coordinated with sugar and phenolic accumulation produces balanced berries with desirable texture and chemical composition. Measurement of rapid firmness loss provides viticultural decision support: firmness testing 2-3 times per week during veraison using penetrometer or compression devices guides harvest timing particularly for varieties where texture is critical (table grapes, some wine styles), firmness trends predict optimal picking windows allowing logistics planning, and cultivar-specific firmness benchmarks calibrated to desired wine styles enable targeted harvest scheduling.
Rapid_Firmness_Loss is distinguished from Maximum_Softening_Rate by representing sustained rapid decline rather than instantaneous peak velocity, from Initial_Berry_Softening by exhibiting accelerated rather than onset rates, and from Complete_Berry_Softening by focusing on the active transformation process rather than final state. Duration typically 3-6 days during peak softening. Firmness decline rates: soft cultivars 4-7% per day, medium 3-5% per day, firm varieties 2-4% per day. Cell wall degradation contributes 70-85% to firmness loss, turgor changes 15-25%, other factors 5-10%. Rapid_Firmness_Loss timing and magnitude critically determine harvest window and berry handling requirements.
accelerated mechanical resistance decline
beschleunigte mechanische Widerstands-Abnahme
fast texture softening
quick firmness reduction
rapid compression resistance loss
rapider Kompressions-Widerstands-Verlust
rasche Festigkeits-Reduktion
schnelle Textur-Erweichung
Keller M (2020) The Science of Grapevines 3rd Ed pages 228-232 berry firmness dynamics Thomas TR, Shackel KA, Matthews MA (2008) Mesocarp cell turgor in Vitis vinifera L. berries throughout development and its relation to firmness, growth, and the onset of ripening Australian J Grape Wine Res 14:1-8 firmness measurement and mechanisms Yakushiji H, Sakurai N, Morinaga K (2001) Changes in cell-wall polysaccharides from the mesocarp of grape berries during veraison Physiol Plant 111:188-195 structural basis of firmness loss.
Rapid Firmness Loss
Chardonnay in Champagne shows rapid initial expansion during mid-late July Year N+1 at BBCH 73-75, with overlapping cell division declining and cell expansion accelerating weeks 3-5, peak growth rate 85-145 mg per day, berry diameter increasing from 13mm to 10-12mm final size, water uptake 0.8-1.4 g per berry per day, under cool maritime conditions of 19-25°C temperature, adequate summer rainfall supporting expansion, typical of this early-bursting cultivar completing rapid growth
Rapid initial expansion represents the combined growth phase during berry development when ongoing cell division overlaps with accelerating cell expansion creating maximum berry growth rate, berry volume increasing dramatically through both cell proliferation and cell enlargement, and berry achieving majority of final size, proceeding over weeks 3-6 post-fertilization as transition from division-dominated to expansion-dominated growth. This peak growth phase occurs during mid summer in July-early August in temperate regions Year N+1, spanning approximately 3-4 weeks during maximum growth intensity, corresponding to BBCH 73-75 pea-sized to berry touch stages. Expansion mechanism involves overlapping processes with declining cell division continuing through week 4-5, accelerating cell expansion through water uptake and vacuole enlargement, synergistic growth from simultaneous division and expansion, and maximum volumetric increase rate during this transitional period. Growth rate characteristics show peak absolute growth rate typically 50-200 mg fresh weight per day during weeks 3-5, maximum diameter increase rate 1-2 mm per week during peak period, volumetric growth rate 0.3-1.0 mL per day in large-berried cultivars, and growth rate representing combined division and expansion contributions. Combined division-expansion dynamics show cell division continuing though declining during weeks 3-4, cell expansion accelerating as cells complete division and enlarge, overlapping processes creating synergistic maximum growth, and transition to expansion-only growth by week 5-6. Cell expansion mechanisms show water uptake as primary driver of cell enlargement, osmotic gradient from solute accumulation pulling water into cells, vacuole expansion accounting for majority of cell volume increase, cell wall loosening through enzymatic modification enabling expansion, and turgor pressure driving cell enlargement against yielding walls. Water uptake dynamics show xylem-mediated water import during early expansion, water flux rates 0.5-2.0 g water per berry per day during peak uptake, water comprising 80-85% berry fresh weight during growth, and water availability critically affecting expansion rate and berry size. Cell wall modification enabling expansion shows expansin proteins mediating wall loosening, xyloglucan endotransglucosylase rearranging wall polymers, pectin methylesterase modifying pectin properties, progressive wall loosening enabling cell expansion, and new wall synthesis maintaining integrity during enlargement. Vacuole development shows central vacuole formation and expansion in expanding cells, vacuole occupying 80-90% cell volume at expansion completion, vacuole as major storage compartment for water and eventually sugars and acids, and vacuole development essential for cell expansion capacity. Solute accumulation driving water uptake shows organic acid synthesis particularly malate and tartrate, mineral ion import including potassium, sugar beginning to accumulate though low levels during growth, osmotic potential decline from -0.5 to -1.5 MPa driving water influx, and solute accumulation creating osmotic gradient essential for expansion. Diameter increase shows berry expanding from 12-18 mm at expansion phase entry to 8-12 mm in small-berried or 14-18 mm in large-berried cultivars by expansion completion, diameter increase 4-8 mm during rapid phase, and diameter as convenient expansion measurement. Volume increase shows exponential volumetric growth during rapid phase, volume expanding 3-6 fold during weeks 3-6, typical range from 0.8-2 mL at phase start to 3-8 mL at completion, and volumetric increase reflecting combined linear diameter increase. Fresh weight accumulation shows weight increase from 0.8-2 g at phase entry to 3-10 g at completion depending on cultivar, weight gain 50-200 mg per day during peak growth, cumulative weight gain 2-8 g during rapid phase, and fresh weight as practical growth measure correlating with volume. Berry appearance changes show berry expanding visibly over days to weeks, green color maintained during growth phase, smooth surface from coordinated growth, and transition from young berry to pea-sized or larger berry morphology. Skin growth coordination shows exocarp cells dividing and expanding maintaining surface coverage, coordinated skin growth preventing cracking during berry expansion, and skin expansion capacity limiting berry size in some cases. Berry cluster filling shows berries expanding to contact neighboring berries, cluster compactness increasing as berries fill available space, and cluster architecture affecting individual berry expansion through physical constraints. Environmental effects on expansion include temperature affecting expansion rate with optimal 22-28°C supporting rapid expansion completing phase within 3-4 weeks, water availability critically affecting cell expansion and final size with adequate water essential, humidity possibly affecting surface expansion and cracking risk, and overall favorable conditions maximizing expansion and size achievement. Temperature effects show warm conditions promoting rapid expansion, cool conditions slowing but generally not preventing expansion, and temperature as factor affecting expansion rate and duration. Water effects show adequate water essential for turgor-driven cell expansion, water deficit limiting expansion potentially reducing berry size 20-40%, irrigation beneficial during rapid expansion weeks 3-6 in water-limited environments, and water as most critical environmental factor during expansion. Nutrient effects show potassium affecting water relations and cell expansion, calcium influencing cell wall properties, and overall mineral nutrition supporting expansion though usually adequate in vineyards. Failed or limited expansion causes include water deficit preventing adequate cell enlargement, extreme temperature stress disrupting growth, skin growth limitation restricting berry expansion in rare cases, or physiological dysfunction though uncommon. The process represents second phase of initial berry growth emphasizing overlap of division and expansion, following early cell division growth and preceding berry size establishment. Duration typically 3-4 weeks representing peak growth transition period. Regional timing shows rapid initial expansion during mid July-early August across climates during maximum summer growth. Viticultural significance includes representing period of maximum berry growth rate determining substantial size increase, demonstrating critical water requirement during expansion for size achievement, showing transition from division to expansion as growth mechanism, and establishing majority of final berry size affecting yield and quality. Research applications include analyzing expansion mechanisms and cell enlargement, investigating water relations and turgor effects on expansion, evaluating environmental effects particularly water on expansion and size, quantifying growth dynamics during transition phase, and assessing expansion as berry size and quality determinant.
Rapid_Initial_Expansion is second phase of Initial_Berry_Growth, distinguished by overlapping division and expansion creating peak growth. Duration 3-4 weeks representing transition period. Maximum growth rate 50-200 mg/day. Water uptake critical driver of expansion. Berry expands 3-6 fold volumetrically. Temperature and water strongly affect rate and size. Follows Early_Cell_Division_Growth. Precedes Berry_Size_Establishment. Represents majority of size achievement.
Spitzen-Beerenausdehnung
combined division-expansion
kombinierte Teilungs-Expansion
maximale Wachstumsphase
maximum growth phase
peak berry expansion
schnelle initiale Expansion
transitional growth
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 438-445 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Rapid Initial Expansion
Cabernet Franc in Loire Valley shows rapid shoot elongation from late May through late June at BBCH 19-31 over 4-5 weeks, with shoots extending from 750 mm at 9 leaves to 1600-1900 mm at flowering through elongation rates accelerating from 22-28 mm per day to peak 38-48 mm per day mid-phase, under temperate maritime conditions of 18-24°C mean temperature, adequate rainfall or irrigation, high solar radiation, and moderate-high fertility typical of this vigorous mid-season cultivar
Rapid shoot elongation represents the accelerating growth phase where elongation rate increases exponentially as photosynthetic capacity develops and favorable environmental conditions support maximum resource availability for growth, characterized by visibly dramatic daily shoot extension establishing majority of final canopy architecture. This developmental stage typically occurs during May-June in temperate regions corresponding approximately to BBCH 19-31 as shoots develop 9 leaves through flowering, with elongation rate accelerating from 20-30 mm per day to 35-50 mm per day creating total shoot length increase from 600-800 mm to 1200-2000 mm over 3-5 week period depending on vigor and conditions. Rapid elongation represents exponential growth phase of sigmoid growth curve where rate increases progressively as each additional expanding leaf contributes photosynthetic capacity and favorable temperature and water availability support maximum metabolic activity. The apical meristem maintains maximum activity producing new leaf primordia every 2 days and contributing high cell production rate to shoot axis, with mitotic activity creating zone of dividing cells extending 100-200 micrometers behind apex supplying cells for continued elongation. Internode lengths during rapid elongation typically range 60-120 mm reflecting high growth rate and abundant resource availability, with longest internodes typically occurring during this phase when conditions and developmental state optimize for maximum expansion. Cell expansion in subapical zones proceeds at maximum rates through abundant water uptake and active wall loosening, with expansion zone extending 20-35 mm behind apex containing cells rapidly increasing to final lengths of 200-500 micrometers creating visible daily shoot extension. Leaves continue unfolding and expanding with new leaves emerging every 2-3 days and expanding rapidly to large dimensions, with leaf area accumulation rate highest during rapid elongation contributing progressively to whole-plant photosynthetic capacity. Photosynthetic capacity increases dramatically during rapid elongation as leaf area expands exponentially and mature leaves achieve full photosynthetic function, with whole-shoot net photosynthesis strongly positive producing substantial excess assimilate beyond shoot growth requirements contributing to whole-plant carbon economy. Carbon allocation during rapid elongation directs substantial photosynthate to shoot growth supporting high cell production and expansion rates, with shoots consuming 60-80% of their photosynthetic production for growth while contributing excess to developing fruit, other vegetative growth, and beginning reserve accumulation. Vascular system extends rapidly through elongating shoots with xylem differentiation maintaining high water transport capacity of 3-6 mg per second supporting rapid expansion and increasing transpiration, and phloem development enabling bidirectional transport moving photosynthate to growth zones and excess production to sinks. Water demand increases dramatically during rapid elongation as transpiring leaf area expands from 1000-1500 square mm to 3000-5000 square mm, requiring shoot water uptake rates increasing from 200-300 mg per hour to 400-700 mg per hour depending on environmental conditions and shoot size. Nutrient requirements peak during rapid elongation particularly for nitrogen supporting high protein synthesis rates, phosphorus supporting energy metabolism and nucleic acid synthesis for new cells, and potassium supporting cell expansion and enzyme activation throughout rapidly growing tissues. Metabolic activity operates at maximum levels with respiration rates in expanding tissues 12-18 fold above mature levels providing energy and carbon skeletons supporting rapid growth, though declining as proportion of photosynthetic production as carbon balance becomes strongly positive. Protein synthesis operates at maximum capacity producing structural proteins for rapidly expanding cells, photosynthetic proteins in developing leaves, regulatory proteins coordinating complex developmental processes, and enzymes supporting high metabolic activity. Hormonal regulation shows peak auxin production from active shoot apex maintaining elongation and strong apical dominance, elevated gibberellins supporting maximum cell expansion and internode elongation, and high cytokinins from active roots promoting cell division and leaf expansion. Temperature during rapid elongation strongly affects growth rate with optimal range 22-28°C supporting maximum elongation approximately doubling per 10°C within physiological range, with warm spring-early summer conditions typically occurring during this phase promoting maximum growth potential. Light intensity affects rapid elongation through photosynthetic capacity and internode length responses, with high light supporting maximum photosynthetic rate providing assimilates for growth and promoting moderate internode lengths, while shade reduces photosynthetic capacity limiting growth potential though may promote elongated internodes through etiolation. Water availability critically affects rapid elongation as high transpiration rates and growth water requirements demand continuous adequate supply, with water stress rapidly arresting elongation through turgor loss and can prematurely transition to declining growth phase permanently limiting final canopy size. Nutrient availability must be adequate to support high growth rates particularly nitrogen for protein synthesis, with deficiency during rapid elongation causing premature transition to declining phase limiting canopy development and subsequent productivity. The process typically spans 3-5 weeks under favorable conditions with consistent temperatures 20-26°C, adequate soil moisture and nutrition, and sufficient light, with vigorous cultivars showing extended duration while less vigorous types show briefer rapid phase. Cultivar variation appears in maximum elongation rate during rapid phase ranging from 30-40 mm per day in moderate vigor cultivars to 45-60 mm per day in highly vigorous types, duration of rapid phase, and responsiveness to environmental conditions and management. Regional timing shows rapid elongation occurring May-June in warm climates, late May through July in temperate regions, and June-July in cool areas, with timing relative to flowering varying but typically encompassing pre-bloom through early fruit set period. Viticultural management considerations include ensuring adequate water availability maintaining turgor and photosynthetic capacity supporting maximum growth, providing sufficient but not excessive nitrogen supporting rapid growth while avoiding over-stimulation, implementing shoot positioning operations managing canopy development for optimal light interception and air movement, monitoring vigor uniformity assessing resource distribution and vine health, and planning timing of trimming operations relative to flowering and fruit set to optimize reproductive success while managing excessive vegetative vigor. Excessive rapid elongation creates overly vigorous dense canopies reducing fruit quality through shading and disease pressure, while limited rapid elongation reduces photosynthetic capacity potentially limiting fruit development and reserve accumulation. Research applications include analyzing mechanisms of exponential growth including hormonal and metabolic regulation, quantifying carbon balance dynamics and assimilate partitioning patterns, evaluating environmental and management effects on growth patterns affecting canopy architecture and productivity, modeling growth responses for decision support systems, and assessing genetic variation in growth characteristics affecting vineyard management requirements and wine quality potential.
Rapid_Shoot_Elongation is second of five Shoot_Elongation subphases, distinguished by accelerating exponential growth rate and strongly positive carbon balance. Transitions to Peak_Shoot_Elongation as maximum linear growth rate is achieved. Represents phase of maximum canopy establishment and photosynthetic capacity development. Critical phase determining final shoot length and productive capacity.
Beschleunigungsphase
accelerating growth phase
exponential elongation
exponentielle Wachstumsphase
maximale Streckungsrate
maximum elongation rate
rapid vegetative growth
rapide Triebstreckung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 220-235 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 140-152 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Rapid Shoot Elongation
In a Paso Robles Zinfandel vineyard, rapid sugar accumulation start was documented in late August Year N+1 (BBCH 84, 8 days post-initial increase) when Brix rose from 17.5° to 26.2° over 14 days (average 4.2% per week), VvHT1 at maximum expression (108-fold), cell wall invertase activity 95 units/g FW, vacuolar hexose increased from 980 mM to 1650 mM, berry fresh weight expanded 15% despite declining growth, under warm conditions (daily high 30-34°C, managed through irrigation maintaining adequate vine water status).
Rapid_Sugar_Accumulation_Start represents the transition to maximum velocity sugar import when all transport and metabolic systems operate at peak efficiency creating the fastest hexose accumulation rates of the ripening period that dramatically elevate berry sweetness from moderate to harvest-ready levels. This rapid phase begins during mid-veraison (BBCH 84), typically 6-10 days post-initial sugar increase, and is characterized by sugar accumulation accelerating to peak rates 2.8-4.5% Brix per week (compared to initial phase 1.5-2.5% per week) driven by fully-induced transporter expression (VvHT genes at 90-110% maximum), maximum enzyme activities (cell wall invertase 60-120 units/g FW, vacuolar invertase 40-90 units/g), and optimized cellular capacity with vacuoles at 93-96% cell volume creating hexose concentrations rising 150-250 mM per week toward final levels 1200-1800 mM. Molecular basis of rapid accumulation includes: hexose transporter genes (VvHT1, VvHT2, VvHT3, VvHT5, VvHT6) expressed at maximum steady-state levels (100-150 fold above green berry) producing saturating amounts of membrane transport proteins, cell wall invertase genes at peak expression (80-120 fold) generating enzyme saturating apoplastic space enabling complete sucrose cleavage, sucrose synthase showing coordinated regulation providing alternative sucrose metabolism pathway producing UDP-glucose for specialized uses while most sucrose undergoes invertase cleavage, and regulatory networks fully activated with sugar-responsive transcription factors, ABA signaling components, and metabolic enzymes all operating at maximum creating coordinated high-flux state. Transport kinetics at rapid accumulation show: plasma membrane hexose uptake Vmax reaching maximum 65-110 μmol glucose g⁻¹ h⁻¹ with transporter density saturating membrane capacity, fructose transport Vmax 55-100 μmol g⁻¹ h⁻¹ similarly maximized, tonoplast transport rates 50-90 μmol hexose g⁻¹ h⁻¹ efficiently clearing cytoplasmic hexoses into vacuoles, and overall berry hexose import 80-150 μmol total hexose g⁻¹ h⁻¹ representing integration of apoplastic delivery, membrane transport, and vacuolar sequestration all operating near maximum. Vacuolar dynamics during rapid accumulation demonstrate: hexose concentrations rising from 700-1000 mM at rapid start to 1200-1800 mM approaching maturity representing berry Brix increasing from 15-20° to 22-28°, osmotic potential declining from -1.6 to -2.2 MPa to -2.2 to -3.2 MPa creating strong water import driving continued berry expansion 10-20% despite declining growth rates, glucose:fructose ratio stabilizing near unity (0.98-1.02) through balanced import and any differential metabolism equilibrating by this stage, and vacuolar pH maintained 2.8-3.4 providing optimal environment for anthocyanin stability (reds), flavor compound retention, and acid balance. Apoplastic sugar dynamics show: sucrose concentration in phloem sap arriving at berry 400-700 mM maintaining high delivery flux, apoplastic sucrose transiently accumulating 100-250 mM before invertase cleavage, resulting apoplastic hexoses 150-350 mM combined creating strong concentration gradient (apoplast 250-350 mM → cytoplasm maintained 60-100 mM) driving facilitated diffusion, and rapid turnover with residence time in apoplast 1.2-1.4 indicating substrate-level metabolism dominates with fermentative pathways activated under hypoxic conditions in berry interior, ATP turnover estimated 40-80 μmol g⁻¹ h⁻¹ with 15-25% allocated to hexose transport-associated processes (H⁺-ATPases maintaining gradients, secondary active transport if present), and glycolytic flux 25-50 μmol glucose g⁻¹ h⁻¹ metabolizing portion of imported hexoses providing energy and metabolic intermediates while majority (70-85%) accumulates in vacuoles. Hormonal status during rapid accumulation demonstrates: ABA sustained at plateau 1400-2400 ng/g maintaining maximum ripening program activation without further increase, hexose signaling at maximum through hexokinase pathway with glucose-6-phosphate 25-50 μmol/g FW creating strong sugar-responsive gene activation, gibberellin and auxin remaining at post-growth minima removing any suppression, and ethylene showing cultivar patterns with some varieties maintaining slight elevation 0.4-1.2 nL g⁻¹ h⁻¹ potentially modulating ripening progression while others remain baseline suggesting ethylene not essential driver. Environmental effects on rapid accumulation rate inclu temperature critically important with optimal 26-30°C supporting maximum rates 3.8-4.5% Brix per week, cooler conditions 22-26°C reducing to 3.0-3.8% per week through slowed kinetics, heat stress 32-35°C impairing to 2.0-3.0% per week through enzyme denaturation and membrane disruption, extreme heat >35°C potentially reducing to <1.5% per week or causing arrest, light ensuring adequate source photosynthesis with shaded canopies potentially limiting accumulation 15-30% despite full berry transport capacity through substrate limitation, and water status with moderate deficit compatible maintaining rates while severe deficit (<-1.5 MPa stem ψ) may reduce through metabolic impairment. Cultivar variation in rapid accumulation characteristics shows: high-Brix varieties (Gewürztraminer potential 26-30° Brix, Sémillon 24-28°, Chenin Blanc 23-27°) achieve peak rates 3.8-4.8% per week over 2-3 week periods, moderate cultivars (Chardonnay 22-26°, Cabernet Sauvignon 23-27°, Merlot 22-26°) show 3.2-4.2% per week for 2-4 weeks, while lower-sugar types (Pinot Noir cool climates 20-24°, some table grapes 18-22°) display 2.5-3.5% per week for 3-5 weeks reflecting genetic differences in transporter expression, metabolic capacity, and developmental programming. Coordination with ripening completion shows: rapid sugar accumulation coincides with berry softening approaching final state (75-90% total reduction completed), color development (reds) reaching near-maximum anthocyanin levels (85-98% final), acid decline intensifying with malate consumption reaching 60-80% total degradation, and flavor/aroma compound synthesis peaking producing characteristic ripe fruit profiles indicating integrated maturation approaching harvest readiness. Quality implications of rapid accumulation inclu rate and duration determining final sugar levels and thus harvest timing and potential alcohol, uniformity across berries and clusters critical for ripening synchrony with heterogeneous rapid phase creating maturity variation, balance between sugar and other quality parameters (acid, pH, phenolics, flavor) defining optimal harvest window with ideal showing coordinated maturation, and environmental conditions during rapid phase substantially affecting wine style through effects on final composition. Viticultural management during rapid accumulation involves: irrigation minimized or terminated as berry water demand declines and concentration desirable though severe deficit risking metabolic impairment avoided, canopy health maintained ensuring continued photosynthesis supporting sugar export without leaf removal that could limit source capacity, disease monitoring critical as high-sugar berries attract insects and provide pathogen substrate, and harvest preparation initiated as rapid accumulation indicates approaching maturity requiring logistics readiness for picking when targets achieved.
Rapid_Sugar_Accumulation_Start is distinguished from Initial_Sugar_Increase by representing maximum (2.8-4.5% Brix/week) rather than moderate (1.5-2.5%/week) rates, from Sugar_Import_Enhancement by showing peak rather than progressive amplification, and from sugar plateau by exhibiting maximum velocity rather than declining rates. Transporter expression 90-110% of maximum. Uptake rates peak 65-110 μmol glucose g⁻¹ h⁻¹. Duration 2-4 weeks BBCH 84-85. Temperature optimal 26-30°C. Rapid_Sugar_Accumulation_Start represents critical phase achieving harvest-ready sweetness levels.
Spitzen-Zucker-Akkumulations-Geschwindigkeit
accelerated Brix elevation
beschleunigte Brix-Erhöhung
fastest sweetening phase
maximale Hexose-Import-Rate
maximum hexose import rate
peak sugar accumulation velocity
schnellste Süßungs-Phase
Keller M (2020) The Science of Grapevines 3rd Ed pages 222-226 rapid accumulation phase Ollat N, Gaudillère JP (1998) The effect of limiting leaf area during stage I of berry growth on development and composition of berries of Vitis vinifera L. cv. Cabernet Sauvignon Amer J Enol Vitic 49:251-258 source effects on accumulation Dai ZW, Meddar M, Renaud C, Merlin I, Hilbert G, Delrot S, Gomès E (2014) Long-term in vitro culture of grape berries and its application to assess the effects of sugar supply on anthocyanin accumulation J Exp Bot 65:4665-4677 sugar accumulation regulation.
Rapid Sugar Accumulation Start
Completer in the Bündner Herrschaft of Graubünden, Switzerland bearing a rare cultivar role persists on approximately 3-5 hectares cultivated by 8-12 producers in the villages of Malans, Jenins, and Maienfeld, with the cultivar's total global population confined almost entirely to this 15 km stretch of the Alpine Rhine Valley, annual production of 15,000-25,000 bottles maintaining commercial visibility sufficient to sustain grower interest, and a ProSpecieRara heritage cultivar designation providing institutional recognition and promotional support that has stabilized planting area after decades of decline from an estimated 50+ hectares in the 19th century.
A conservation role that inheres in a grapevine cultivar that exists in very limited total planting area — typically fewer than 10-50 hectares globally — and is grown by a small number of producers, representing a restricted but not yet critically endangered population that requires monitoring and proactive conservation attention to prevent further decline toward endangerment.
Rare_Cultivar_Role describes cultivars that are uncommon but not yet at immediate extinction risk, occupying the conservation status zone between common commercial cultivation and critical endangerment. The distinction between rare and endangered is quantitative rather than qualitative and depends on contextual assessment of population trend, geographic concentration, reproductive security, and institutional support: a cultivar with 20 hectares concentrated in a single commune with declining grower interest may be at greater effective risk than a cultivar with 5 hectares distributed across multiple conservation collections. Rare cultivars represent the largest pool of grapevine genetic diversity at conservation risk, estimated at 1,000-2,000 cultivars in Europe alone.
Kleinstrebsorten-Rolle
Raritäten-Rolle
limited-distribution cultivar role
minor cultivar role
rare variety role
seltene Sorten-Rolle
Anderson K and Aryal NR (2013) Which Winegrape Varieties are Grown Where?, University of Adelaide Press Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157 Robinson J et al. (2012) Wine Grapes, Allen Lane pp: 1-18
Rare Cultivar Role
Sangiovese R24 bears a Rauscedo_Clone_Role as VCR's benchmark quality selection for Tuscan production, evaluated through replicated trials across Chianti Classico and Montalcino, with documented wine quality supporting premium positioning. Chardonnay VCR 8 serves in a Rauscedo_Clone_Role as a widely distributed selection balancing productivity with quality, VCR's global nursery network ensuring availability across international markets. SO4 rootstock clone bears a Rauscedo_Clone_Role as the world's most planted rootstock, VCR production supplying the majority of global SO4 benchgrafts—demonstrating Rauscedo's dominant position in rootstock as well as scion production.
An Italian clone role that inheres specifically in clones selected and distributed by Vivai Cooperativi Rauscedo (VCR), the world's largest grapevine nursery cooperative located in Friuli-Venezia Giulia (northeastern Italy), which conducts clone selection, maintains mother plant collections, and produces over 80 million grafted vines annually for global distribution. Rauscedo clones are designated with "R" or "VCR" prefixes followed by numeric codes (e.g., Sangiovese R24, Chardonnay R8, Merlot VCR 1), indicating their selection through VCR's extensive evaluation program. VCR operates one of the world's most comprehensive clone selection infrastructures, with experimental vineyards across Italian regions, winemaking facilities for clone evaluation, and virus-testing laboratories ensuring certified health status. Rauscedo's dominant position in international rootstock production (producing the majority of benchgraft rootstocks planted globally) extends its influence beyond scion varieties to the foundational material underlying world viticulture. VCR's integration of selection, production, and distribution creates a vertically integrated clone supply chain unmatched elsewhere.
Rauscedo_Clone_Role identifies the specific institutional origin within the Italian system, distinguishing VCR selections from Conegliano, regional station, or other Italian program clones. VCR's scale (80+ million vines annually, global distribution network, integrated selection-production-distribution operations) makes it uniquely influential in world viticulture decisions about which clones VCR promotes affect global planting patterns. The cooperative structure (grower-members collectively owning the nursery) distinguishes VCR from private nurseries, with selection priorities reflecting producer interests rather than purely commercial considerations. VCR's rootstock dominance is particularly significant—most grafted vines worldwide contain Rauscedo-produced rootstock material, regardless of scion origin—making VCR's sanitary protocols and clone selections foundational to global vineyard health.
Friuli clone
Friuli-Klon
R-designated clone
Rauscedo clone
Rauscedo-Klon
VCR-Klon
Vivai Cooperativi Rauscedo (2020) VCR Catalogo Varietà e Cloni Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 Crespan M et al. (2006) Molecular characterization of Italian grapevine cultivars, Vitis 45: 69-73 Borgo M, Michelet E (2000) Italian grapevine selection, ISHS Acta Horticulturae 528: 35-42
Rauscedo Clone Role
Investigation of Reacclimation_Capacity_Disposition at the University of Minnesota employed deacclimation-reacclimation cycling experiments to characterize recovery potential across 15 cultivars. Dormant canes were subjected to 7 days at 15°C (deacclimation), then returned to -3°C for 3, 7, or 14 days (reacclimation), with hardiness assessed by differential thermal analysis at each transition. The experiment was conducted in mid-February (soon after endodormancy release) and repeated in mid-March (advanced ecodormancy). In February, 'Frontenac' exhibited strong Reacclimation_Capacity_Disposition, losing 5°C during deacclimation (from -33°C to -28°C) but recovering 4°C within 7 days of cold return (to -32°C), representing 80% recovery of lost hardiness. 'Marquette' showed similar capacity (6°C loss, 5°C recovery, 83% restoration). 'Riesling' demonstrated moderate reacclimation (5°C loss, 3°C recovery, 60% restoration). 'Pinot Noir' showed poor Reacclimation_Capacity_Disposition (8°C loss, 2°C recovery, 25% restoration). In March, all cultivars showed diminished reacclimation capacity, with 'Frontenac' recovering only 50% of lost hardiness and 'Pinot Noir' showing essentially no recovery. Field observations following a March warm-cold cycle (10 days at 12°C followed by -18°C freeze) confirmed laboratory predictions, with high-reacclimation cultivars suffering minimal damage while low-reacclimation cultivars showed severe bud mortality. These findings establish Reacclimation_Capacity_Disposition as critical for spring cold damage avoidance in variable climates.
A Reacclimation_Capacity_Disposition is a Deacclimation_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to regain freezing tolerance following partial deacclimation when cold conditions return, recovering hardiness lost during warm periods to restore protective levels before subsequent cold events. This disposition represents the reversibility of deacclimation, with high reacclimation capacity enabling recovery from premature hardiness loss and providing resilience against the alternating warm and cold periods characteristic of transitional seasons. Reacclimation is possible only within limits, as tissues that have deacclimated beyond a certain threshold lose the capacity for hardiness recovery, and the reacclimation potential diminishes as the season progresses toward active growth. Reacclimation_Capacity_Disposition encompasses both the magnitude of hardiness recovery achievable (degrees of hardiness regained) and the rate of reacclimation (hardiness gain per day of cold exposure). Genotypes differ in reacclimation capacity, with some cultivars able to recover substantial hardiness rapidly (3-5°C within 72 hours of cold return) while others show limited or slow recovery. Strong reacclimation capacity combined with deacclimation resistance provides optimal protection against variable late-winter weather, as resistant genotypes experience less initial hardiness loss while high reacclimation capacity enables recovery of whatever hardiness is lost during unavoidable warm exposure.
Reacclimation_Capacity_Disposition is_a Deacclimation_Disposition. Reacclimation represents the capacity to reverse hardiness loss, providing resilience against warm-cold cycling in late winter and spring. Reacclimation capacity diminishes as the season progresses, with tissues eventually losing the ability to recover hardiness as growth approaches. Both magnitude (degrees recovered) and rate (speed of recovery) contribute to effective reacclimation. Reacclimation capacity combined with deacclimation resistance provides comprehensive protection against variable transitional-season weather.
Kältehärte-Wiederherstellung
Reakklimatisierungs-Kapazität-Disposition
Wiederhärtungs-Kapazität
cold tolerance restoration
hardiness recovery
rehardening capacity
Kalberer SR et al. (2006) Deacclimation and reacclimation of cold-hardy plants: current understanding and emerging concepts, Plant Science 171:3-16 Pagter M, Arora R (2013) Winter survival and deacclimation of perennials under warming climate: physiological perspectives, Physiologia Plantarum 147:75-87 Ferguson JC et al. (2011) Dynamic thermal profiles of grapevine cold hardiness, American Journal of Enology and Viticulture 62:407-413
Reacclimation Capacity Disposition
Reacclimation capacity assessment following mid-winter thaw in Merlot reveals the partial recovery potential after deacclimation, with initial hardiness of -20°C declining to -12°C during 8-day warm period (>10°C daily maxima), followed by return of cold conditions (-5°C average) for 14 days enabling partial reacclimation to -16°C (recovering 4°C of the 8°C lost, demonstrating 50% reacclimation capacity), with the recovered hardiness providing survival margin when subsequent -14°C event occurred, though the incomplete recovery meant less protection than if deacclimation had not occurred, illustrating how reacclimation capacity provides partial but not complete resilience to temperature fluctuations.
Reacclimation capacity quality is a cold hardiness quality that represents the ability of grapevine tissues to regain freeze tolerance after partial deacclimation has occurred, characterizing the potential for hardiness recovery when cold conditions return following warm periods, with reacclimation capacity determining whether tissues can restore protection or remain vulnerable after hardiness-reducing warm spells during the dormant period. Within the BFO framework, this quality specifically addresses the hardiness recovery dimension of cold hardiness, distinguishing the capacity for regaining lost hardiness from the initial development or the rate of loss while recognizing that reacclimation capacity determines the resilience of winter protection to temperature fluctuations, with high capacity enabling recovery from warm-spell-induced deacclimation and low capacity leaving tissues permanently vulnerable after any warming event. The ontological significance lies in reacclimation capacity determining survival through the fluctuating conditions characteristic of many temperate climates, where repeated cycles of warming (triggering deacclimation) and cooling (potentially enabling reacclimation) create complex hardiness trajectories that depend on whether tissues can regain protection during cooling phases or progressively lose hardiness through repeated warming events. The physiological constraints on reacclimation include the availability of cryoprotectant precursors (may be depleted during extended warm periods), the metabolic capacity for re-synthesizing protective compounds (may be reduced as dormancy progresses), the cellular damage from freeze-thaw cycles (limiting recovery capacity), and the dormancy status (deeply dormant buds potentially having greater reacclimation capacity than buds approaching growth). The factors affecting reacclimation capacity include the extent of prior deacclimation (slight deacclimation more recoverable than severe), the duration of warming (brief warm spells allowing better recovery than extended warmth), the intensity of subsequent cooling (gradual cooling enabling more complete reacclimation), and cultivar genetics (some genotypes showing greater reacclimation capacity than others). Understanding reacclimation capacity quality enables assessment of recovery potential following warm periods, prediction of hardiness trajectories through variable winter conditions, and realistic expectations for protection maintenance in climates characterized by temperature fluctuation rather than stable cold.
Reacclimation_Capacity_Quality addresses the ability to regain freeze tolerance after warming-induced hardiness loss. This quality determines resilience to temperature fluctuations during the dormant period. The class supports winter survival prediction through reacclimation capacity characterization.
Härtewiederherstellung
Reakklimatisierungskapazität
Reakklimatisierungspotential
Wiederhärtungsfähigkeit
hardiness regain capacity
reacclimation potential
Mills LJ et al. (2006) Timing of low temperature exposure affects cold hardiness and survival of dormant grapevine buds. American Journal of Enology and Viticulture 57(2): 232-239 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 267-330
Reacclimation Capacity Quality
Reactive oxygen species scavenging quality comparison across grapevine rootstock genotypes evaluated the contribution of oxidative stress management to rootstock effects on scion heat tolerance in warm climate viticulture. Total ROS scavenging capacity was assessed using the oxygen radical absorbance capacity (ORAC) assay and complemented with specific scavenging assays for individual ROS species. Rootstock 110R demonstrated excellent reactive oxygen species scavenging quality with ORAC values 2.5-fold higher than Riparia Gloire under heat stress (42°C for 4 hours), hydrogen peroxide scavenging rate 3.2-fold higher reflecting elevated APX and CAT activities, and superoxide scavenging 2.1-fold higher reflecting superior SOD activity. The scavenging quality differential translated to reduced oxidative damage in 110R-rooted scions, with lipid peroxidation (MDA levels) only 25% above control versus 85% above control in Riparia Gloire-rooted scions under equivalent heat exposure. Subcellular analysis revealed that 110R showed particularly enhanced chloroplastic scavenging capacity with thylakoid-associated APX activity 4-fold higher than Riparia Gloire, providing superior protection at the primary site of heat-induced ROS generation. The reactive oxygen species scavenging quality assessment established oxidative stress management as a primary mechanism by which drought-tolerant rootstocks also confer heat tolerance to scions, suggesting that rootstock selection for warm climates should consider scavenging capacity alongside other tolerance traits.
Reactive oxygen species scavenging quality is a heat shock response quality that inheres in a grapevine organism and represents the overall capacity to neutralize and eliminate the various reactive oxygen species including superoxide radical, hydrogen peroxide, hydroxyl radical, and singlet oxygen that accumulate under heat stress conditions when electron transport chains become over-reduced and normal metabolic processes generate ROS at rates exceeding baseline scavenging capacity. This quality integrates the function of enzymatic and non-enzymatic antioxidant systems into an overall assessment of ROS elimination capacity, recognizing that the balance between ROS production and scavenging determines whether oxidative damage accumulates, with effective scavenging maintaining ROS at signaling-competent but non-damaging levels while inadequate scavenging permits toxic accumulation causing membrane lipid peroxidation, protein oxidation, and DNA damage. The ontological significance of reactive oxygen species scavenging quality lies in its role as the ultimate determinant of oxidative stress outcomes, with scavenging capacity relative to ROS production rate determining the redox balance that governs whether cells experience beneficial stress signaling or destructive oxidative damage under heat exposure conditions. The biological basis of reactive oxygen species scavenging encompasses the coordinated activity of multiple scavenging systems each with specificity for particular ROS species and localization in specific cellular compartments, including superoxide dismutase scavenging superoxide radical, catalase and the ascorbate-glutathione cycle scavenging hydrogen peroxide, and non-enzymatic scavengers including carotenoids quenching singlet oxygen and tocopherols interrupting lipid peroxidation chain reactions. The chloroplast represents a particularly important site of heat-induced ROS production where over-reduction of electron transport chains under conditions of impaired carbon fixation diverts electrons to oxygen, while mitochondrial electron transport similarly generates ROS when respiratory chain function is disrupted by heat, creating multi-compartment oxidative stress requiring coordinated scavenging response. The capacity for ROS scavenging varies substantially among grapevine genotypes, providing genetic variation that contributes to overall differences in heat tolerance and offering targets for selection or breeding to enhance oxidative stress management under heat exposure.
Reactive_Oxygen_Species_Scavenging_Quality is_a Heat_Shock_Response_Quality, representing the overall capacity to eliminate ROS generated under heat stress. This class addresses the integrated scavenging function across multiple ROS species and cellular compartments. ORAC assay provides total scavenging capacity metric while specific assays address individual ROS. Chloroplastic scavenging is particularly critical given photosynthetic electron transport as primary ROS source. Rootstock effects on scion ROS management demonstrate systemic tolerance influences. Distinguished from Antioxidant_Response_Quality which addresses the induction of antioxidant systems rather than the resultant scavenging capacity.
ROS scavenging capacity
ROS-Entgiftungskapazität
Radikalentgiftung
free radical elimination
oxidative Detoxifikation
oxidative detoxification
reactive oxygen neutralization
reaktive Sauerstoffspezies-Eliminierung
Wahid A et al. (2007) Heat tolerance in plants: an overview. Environmental and Experimental Botany 61:199-223 Carvalho LC et al. (2015) Heat stress in grapevine: the pros and cons of acclimation. Plant, Cell and Environment 38:777-789 Mittler R (2002) Oxidative stress, antioxidants and stress tolerance. Trends in Plant Science 7:405-410
Reactive Oxygen Species Scavenging Quality
Albariño vines planted in the Mornington Peninsula of Victoria, Australia since 2010 bearing a recently introduced role represent experimental plantings of 5-15 ha by pioneering producers exploring Iberian cultivars as climate adaptation alternatives, with trial blocks showing promising results at yields of 6-9 t/ha and natural acidity retention of 7-9 g/L titratable acidity at harvest compared to declining acid levels in the region's established Pinot Noir and Chardonnay plantings, though commercial viability and consumer acceptance remain under assessment after only 12-15 vintages.
An introduced cultivar role that inheres in a grapevine cultivar that has been planted in a region within approximately the past two decades, representing a novel addition to the regional varietal portfolio whose long-term viticultural suitability, terroir expression, and market acceptance remain under active evaluation.
Recently_Introduced_Role captures the transitional status of cultivars in the early phase of regional establishment, before sufficient viticultural experience, clonal evaluation, and market data have accumulated to confirm long-term suitability. The role is particularly relevant in the context of climate change-driven varietal experimentation, where many traditional cool-climate wine regions are trialling warmer-climate cultivars (e.g., Nero d'Avola in southern England, Touriga Nacional in Oregon, Assyrtiko in Australia) as adaptive responses to rising temperatures: most of these introductions are too recent for definitive assessment of their regional potential. The temporal boundary of "recently introduced" shifts with context but conventionally refers to plantings within the past 20-25 years.
Neuzugangs-Rolle
kürzlich eingeführte Sorten-Rolle
neu angepflanzte Rebsorten-Rolle
newly planted cultivar role
novel introduction role
recently introduced variety role
Wolkovich EM et al. (2018) From Pinot to Xinomavro in the world's future wine-growing regions, Nature Climate Change 8: 29-37 Morales-Castilla I et al. (2020) Diversity buffers winegrowing regions from climate change losses, Proceedings of the National Academy of Sciences 117: 2864-2869 Anderson K and Aryal NR (2013) Which Winegrape Varieties are Grown Where?, University of Adelaide Press
Recently Introduced Role
Cabernet Sauvignon hermaphrodite flowers at BBCH 61-65 exhibit receptive stigmas with wet glistening appearance from abundant exudate production, supporting 78-86% germination of self-pollen applied during peak receptivity (days 1-3 post-calyptra shedding), declining to 52-68% germination on declining-receptive stigmas (days 4-5), and <25% germination by day 6-7 as stigmas transition to post-receptive dry state, with the 5-6 day receptivity window sufficient to capture self-pollen from synchronously dehiscing anthers achieving 50-58% fruit set. Gewürztraminer functionally female flowers show receptive stigmas maintaining wet exudate and high pollen germination support (72-84% of applied cross-pollen germinating) for extended duration of 6-8 days post-calyptra shedding, providing prolonged opportunity to capture cross-pollen from hermaphrodite pollinizer sources that may not bloom perfectly synchronously, achieving 28-38% fruit set when pollinizers provide adequate pollen during the extended receptivity window. Syrah stigma exudate biochemical analysis during peak receptivity reveals sugar concentration of 11-14% (65% sucrose, 25% glucose, 10% fructose), protein concentration 0.8-1.4 mg/ml including active invertase and esterase enzymes, pH 5.8-6.4, and osmotic potential of -1.2 to -1.8 MPa, creating optimal biochemical environment for pollen hydration and germination. Merlot controlled pollination experiments applying pollen to receptive stigmas at peak (day 2 post-calyptra shedding) versus declining receptivity (day 5) versus post-receptive dry stigmas (day 8) achieve fruit set rates of 62%, 38%, and 4% respectively, demonstrating critical importance of stigma receptivity timing for pollination success and fruit set achievement.
A receptive stigma is a stigma that has achieved physiological maturity with active production of wet stigmatic exudate and biochemical competence to support pollen grain adhesion, hydration, germination, and pollen tube initiation, representing the functional state during which successful pollination can occur and fertilization can be achieved. Receptive stigmas are characterized by= (1) wet glistening appearance due to secretion of stigmatic exudate coating the papillae surfaces, visible as a shiny film or droplets particularly evident under magnification or favorable lighting (2) stigmatic exudate composition including water (primary component providing pollen hydration), sugars at 8-15% concentration (primarily sucrose, glucose, fructose providing energy for pollen germination and initial tube growth), proteins including enzymes (invertases, esterases, peroxidases), signaling molecules, and structural glycoproteins, lipids (primarily in cuticle and as minor exudate components), and various secondary metabolites (3) papillae in turgid expanded state with cells fully hydrated and exudate-producing apparatus active (4) biochemical and cellular conditions supporting pollen recognition, pollen grain adhesion via exudate viscosity and surface tension, pollen hydration via exudate water transfer, pollen germination via exudate nutrient provision and signaling molecule activation of germination programs, and pollen tube guidance toward transmitting tract entry points. Stigma receptivity develops through maturation of secretory apparatus during late pistil development (BBCH 57-59) with activation typically occurring at or slightly before calyptra shedding (BBCH 60-61), creating temporal coordination between stigma receptivity (female readiness) and anther dehiscence (male pollen availability) in hermaphrodite flowers, though functionally female and pistillate flowers often achieve receptivity 1-2 days before hermaphrodite pollen sources begin dehiscing, providing temporal head-start for capturing early cross-pollination. Receptive stigma duration varies among flower types and environmental conditions= hermaphrodite stigmas maintain peak receptivity 2-4 days with declining receptivity persisting additional 2-3 days (total receptivity window 4-6 days) functionally female and pistillate stigmas show extended receptivity with peak lasting 3-5 days and declining phase 3-4 days (total 6-8 days or occasionally 9 days) functionally male stigmas show absent or brief receptivity (32°C, excessive humidity >85%, water stress) reduce receptivity duration and intensity. Receptive stigmas support pollen germination with high efficiency= 70-90% of compatible pollen grains landing on peak-receptive hermaphrodite stigmas germinate within 2-6 hours 60-80% germination on declining-receptive stigmas (3-5 days post-peak) <20-40% on marginal-receptive stigmas (6-7 days post-peak or on poorly receptive functionally male stigmas). The transition from non-receptive to receptive state represents a critical developmental threshold determining when pollination becomes possible, while the transition from receptive back to non-receptive (post-receptive dry stigma) defines the temporal window during which successful pollination must occur.
Receptive_Stigma represents the functional pollination-competent state and must be distinguished from Dry_Stigma (non-receptive pre-receptivity or post-receptive states lacking wet exudate). Receptivity is the critical determinant of whether pollination can succeed, with receptive stigmas supporting pollen germination while dry stigmas fail to support germination. Sub-classification into Fully_Receptive and Maximally_Receptive reflects variation in receptivity quality/intensity. Understanding stigma receptivity is essential for pollination biology, bloom timing assessment, controlled crossing operations, and diagnosis of reproductive failures related to stigma dysfunction or asynchrony between stigma receptivity and pollen availability.
empfangsbereites Stigma
empfangsfähige Narbe
pollen-accepting stigma
receptive surface
rezeptive Narbe
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 102-108 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 95-101 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 190-196.
Receptive Stigma
Recommended variety status for Blaufränkisch in Burgenland reflects the cultivar's position as the pre-eminent red variety of the region — classified as Qualitätsrebsorte at the national level and designated as the principal or exclusive red variety in three Burgenland DAC regulations (Mittelburgenland, Eisenberg, Rosalia), with active support from the Weinbauschule Eisenstadt, the HBLA Klosterneuburg clonal selection program, and the Österreichische Weinmarketinggesellschaft's international promotional campaigns positioning Blaufränkisch as Austria's flagship indigenous red cultivar. Recommended variety status for Chardonnay in Burgundy represents the apex of cultivar-region identification — the variety is the exclusive white cultivar permitted in all Côte d'Or white appellations from Bourgogne through Village and Premier Cru to Grand Cru, the subject of centuries of clonal observation and selection, and the beneficiary of intensive BIVB (Bureau Interprofessionnel des Vins de Bourgogne) research investment — a recommended status so deeply entrenched that the concept of Burgundy white wine without Chardonnay is essentially inconceivable.
Recommended variety status is a classification status entity that documents the highest-tier regulatory classification of a grape cultivar within a viticultural jurisdiction, indicating that the cultivar has been formally evaluated and designated as particularly suited for quality wine production in the region, actively promoted through research support, extension services, and in some jurisdictions preferential planting authorization or subsidy access, and typically designated as a principal or preferred variety in the appellation specifications of the regions where it is recommended, reflecting the assessment that the cultivar consistently produces wines of recognized quality and commercial viability under the region's viticultural conditions.
Recommended variety status represents the regulatory expression of collective viticultural judgment — the formal assessment by government agencies, research institutions, and industry bodies that a cultivar has demonstrated consistent, above-average quality performance and commercial potential in a specific region, meriting active promotional support and preferential regulatory treatment. The evaluation process underlying recommended status typically involves multi-year comparative trials (minimum 5-10 years at government research stations), assessment of wine quality by professional tasting panels, analysis of agronomic performance data (yield stability, disease resistance, climate adaptation), and review of commercial viability (market acceptance, price positioning). The distinction between recommended and merely authorized status carries practical consequences beyond symbolic ranking — in jurisdictions that maintain the distinction, recommended varieties may receive preferential access to planting authorizations, inclusion in government-funded extension and advisory programs, priority in clonal selection research, and preferential positioning in export market promotional campaigns. The concept of "principal variety" in appellation specifications represents the appellation-level analogue of recommended status — the cultivars that the appellation regulatory body has determined best express the appellation's identity and quality, and that must therefore constitute the dominant proportion of wines carrying the appellation designation. The dynamic nature of recommended status should be noted — cultivars may be promoted from authorized to recommended as regional experience accumulates (as occurred with several Austrian varieties during the post-1985 quality revolution), or may lose recommended status if performance deteriorates or market demand shifts, though the latter is rare due to the institutional inertia of classification systems.
Empfehlungsstatus
Hauptsorte
bevorzugte Rebsorte
empfohlene Sorte
erstklassige Einstufung
first-tier classification
preferred variety
principal variety status
promoted variety
recommended classification
Österreichisches Weingesetz (2009) §3 and Annex 1 INAO (2024) Cahier des charges — cépages principaux et complémentaires HBLA Klosterneuburg (2023) Sortenliste der empfohlenen Qualitätsrebsorten BIVB (2023) Les cépages de Bourgogne
Recommended Variety Status
Groppello di Mocasina in the Garda region of northern Italy bearing a recovered cultivar role was rediscovered in 2000 when SSR profiling of unidentified old vines in abandoned vineyard parcels near Mocasina di Caprino Veronese revealed a genotype matching 19th-century ampelographic descriptions of a cultivar presumed extinct since the phylloxera epidemic, with subsequent recovery actions including propagation of 200 cuttings from 12 surviving centenarian vines, virus elimination through meristem culture, establishment of 0.5 ha conservation planting at the Istituto Agrario di San Michele all'Adige, and reintroduction to commercial production with 3 producers now cultivating 2.5 hectares and marketing single-varietal wines under the Garda DOC designation.
A conservation role that inheres in a grapevine cultivar that was rediscovered and returned to documented cultivation after a period of presumed extinction, near-extinction, or loss from the known cultivar inventory, typically through systematic prospection surveys, DNA identification of unidentified old vines, or recovery from abandoned vineyard sites.
Recovered_Cultivar_Role documents the conservation success stories where presumed-lost genetic diversity is returned to the documented cultivar inventory. Recovery events have accelerated dramatically since the 2000s due to the convergence of systematic old-vine prospection surveys, affordable SSR genotyping enabling rapid identification of unknown accessions, and growing market interest in rare and heritage cultivars providing economic incentive for reintroduction: notable recovery programs include the CERVIM initiatives in Mediterranean steep-slope vineyards, the ProSpecieRara heritage variety program in Switzerland, and multiple Italian regional prospection campaigns that have collectively recovered dozens of cultivars from near-extinction. The role is transitional — recovered cultivars may subsequently be classified under other conservation subclasses (rare, endangered, conservation priority) depending on the success of reintroduction efforts.
Rückkehr-Sorten-Rolle
rediscovered cultivar role
resurrected cultivar role
revived variety role
wiederbelebte Rebsorten-Rolle
wiederentdeckte Sorten-Rolle
Grando MS et al. (2010) Microsatellite-based identification of old grapevine cultivars, American Journal of Enology and Viticulture 61: 116-122 Ferretti CG (2019) A new geographical classification of low-input viticulture, Oeno One 53: 507-523 Maul E et al. (2012) Identification and characterization of grapevine genetic resources, Vitis 51: 53-63
Recovered Cultivar Role
Recovery capacity determines vineyard resilience to catastrophic events and the feasibility of renovation practices, with full productive recovery requiring 2-5 years depending on damage severity and cultivar. Recovery from severe winter injury (trunk kill to soil line) in own-rooted vineyards involves sucker selection and retraining over 3-4 years, while grafted vines may require trunk regrafting or replanting. Recovery from trunk disease damage through cordon or trunk renewal exploits the vine's capacity to regenerate from basal latent buds while maintaining the established root system. Fire-damaged vineyards show variable recovery depending on heat exposure intensity, with complete crown recovery possible within 2-3 seasons when root systems survive. Cultivars with abundant latent bud reserves and high reserve carbohydrate content show superior recovery compared to low-reserve types, with Grenache and Carignan noted for exceptional recovery capacity.
A longevity and resilience disposition that is the inherited capacity of a Vitis vinifera plant to restore productive function following severe damage, stress, or deliberate renovation interventions, through coordinated activation of latent meristems, mobilization of carbohydrate reserves, and reconstruction of functional canopy architecture within acceptable timeframes.
Recovery_Disposition serves as the parent class for retraining capacity and crown regeneration traits. Recovery represents the practical application of the vine's regenerative potential to restore economic productivity following damage or as part of planned vineyard renovation. The trait integrates multiple underlying capacities including latent bud viability, reserve mobilization, and growth vigor. Understanding cultivar-specific recovery potential enables informed decisions about renovation versus replanting following damage events.
Erholungsvermögen
Regenerations-Disposition
Wiederherstellungsfähigkeit
damage recovery trait
rehabilitation disposition
restoration capacity
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 165-195 Wample RL, Bary A (1992) Harvest date as a factor in carbohydrate storage and cold hardiness of Cabernet Sauvignon grapevines. Journal of the American Society for Horticultural Science 117:32-36
Recovery Disposition
Riesling serves in a Recurrent_Parent_Role across multiple backcross generations in German resistance breeding, with disease resistance from initial interspecific donors introgressed while successively recovering Riesling's genetic background and wine quality character. Chardonnay bears a Recurrent_Parent_Role in French breeding programs targeting downy and powdery mildew resistance, with each backcross generation restoring Chardonnay genetics while maintaining R-gene(s) under marker-assisted selection. Cabernet Sauvignon functions as recurrent parent in programs developing "Cabernet-type" resistant varieties, the backcrossing strategy aiming to produce disease-resistant wines approaching Cabernet character through progressive genome recovery.
A parent role that inheres in a grapevine genotype by virtue of its repeated use across successive backcross generations in a backcross breeding program designed to introgress specific traits from a donor parent while recovering the recurrent parent's overall genetic background. The recurrent parent is typically an elite V. vinifera variety with established quality and market acceptance (Riesling, Chardonnay, Cabernet Sauvignon), targeted for improvement through addition of specific traits (disease resistance, stress tolerance) while maintaining its essential varietal character. In backcross breeding, F1 hybrids (recurrent parent × donor parent) are crossed back to the recurrent parent, with progeny carrying the desired donor trait selected and again backcrossed to the recurrent parent across multiple generations (typically BC2-BC5 or beyond). Each backcross generation recovers approximately 50% additional recurrent parent genome, so that by BC4 (fourth backcross) progeny average >96% recurrent parent genetic background while retaining the selected donor trait. Marker-assisted backcrossing (MABC) accelerates background recovery by enabling selection for both the target trait and recurrent parent alleles at non-target loci.
Recurrent_Parent_Role captures the recipient/background parent function in backcross breeding, where the objective is trait addition to an established elite genotype rather than novel variety creation. The strategy acknowledges that elite wine grape varieties represent centuries of selection and market development that cannot be easily replicated in new varieties—making trait introgression into existing varieties more commercially viable than creating entirely new variety identities. Backcross breeding timeline is extended (10-25+ years for adequate background recovery), but products maintain connections to established variety names and wine styles. Marker-assisted selection has dramatically improved backcross efficiency, enabling both foreground selection (confirming target trait presence) and background selection (accelerating recurrent parent recovery by selecting against donor alleles at non-target loci).
Eliteelter
Empfängerelter
Rückkreuzungselter
backcross parent
elite parent
recipient parent
recurrent parent
rekurrenter Elter
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Töpfer R et al. (2011) Molecular breeding, in Adam-Blondon AF et al. (eds) Genetics, Genomics, and Breeding of Grapes, CRC Press pp.: 160-185 Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330
Recurrent Parent Role
Initial screening for Red_Blotch_Resistance_Disposition at UC Davis evaluated GRBV infection outcomes across 35 accessions following graft inoculation with infected budwood. Infection was assessed by qPCR targeting GRBV coat protein gene at 6, 12, and 24 months post-grafting. All tested V. vinifera cultivars (n=20) showed systemic infection within 12 months, with viral titers ranging from 10⁴ to 10⁶ genome copies per ng DNA. Symptom expression varied among cultivars, with 'Cabernet Sauvignon' and 'Merlot' displaying pronounced red blotches covering 40-60% of leaf area by véraison, while 'Chardonnay' and other white cultivars showed subtle interveinal chlorosis. North American species showed variable responses: V. riparia accessions exhibited delayed systemic infection (50% positive at 12 months versus 100% for V. vinifera) and lower viral titers, suggesting potential partial Red_Blotch_Resistance_Disposition worthy of further investigation. V. labrusca-derived 'Concord' supported infection but maintained yields within 90% of healthy controls, demonstrating tolerance. These preliminary findings provide direction for expanded germplasm screening, though no immunity or high-level resistance has been identified to date.
A Red_Blotch_Resistance_Disposition is a Viral_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that limit infection, replication, systemic spread, or symptom expression by Grapevine red blotch virus (GRBV), a recently characterized DNA virus causing red blotch disease in North American and other vineyards. This disposition represents the capacity to resist a newly emerged threat first described in 2012 in California vineyards, where it was initially confused with leafroll disease due to similar foliar reddening symptoms. GRBV is a member of the family Geminiviridae, genus Grablovirus, distinguished from the RNA viruses (GLRaVs) causing leafroll by its circular single-stranded DNA genome that replicates in the nucleus through rolling circle amplification. Red blotch symptoms in red cultivars include irregular red blotches on leaf blades that expand and coalesce during the season, without the leaf rolling typical of GLRaV infection. Physiological impacts include reduced sugar accumulation (2-5°Brix), elevated titratable acidity, diminished anthocyanin synthesis, and altered flavor compound profiles that compromise wine quality. Transmission occurs through the three-cornered alfalfa treehopper (Spissistilus festinus) and possibly other vectors, as well as through infected propagation material. Red blotch resistance research is in early stages given the recent virus characterization, but screening of Vitis germplasm has begun to identify potential resistance or tolerance sources. No strong resistance has been reported to date in V. vinifera, though systematic evaluation of wild species and breeding populations continues.
Red_Blotch_Resistance_Disposition is_a Viral_Resistance_Disposition. Red blotch is caused by a geminivirus (DNA virus) distinct from the RNA viruses causing leafroll and fanleaf, potentially enabling different resistance mechanisms. The disease was only characterized in 2012, so resistance research is at early stages compared to long-studied viral diseases. Current management relies on clean planting material and removal of infected vines, as vector control options are limited and no resistant cultivars are available. Screening of diverse Vitis germplasm for resistance sources is ongoing.
GRBV resistance disposition
GRBV-Resistenz
Red-Blotch-Resistenz
Roter-Fleck-Resistenz-Disposition
geminivirus resistance
red blotch tolerance
Al Rwahnih M et al. (2013) Grapevine red blotch-associated virus is a DNA virus causing red blotch disease, Phytopathology 103:1069-1076 Sudarshana MR et al. (2015) Grapevine red blotch-associated virus, an emerging threat to the grapevine industry, Phytopathology 105:1026-1032 Bahder BW et al. (2016) Grapevine red blotch disease and its vector, Wine Business Monthly 23:80-86
Red Blotch Resistance Disposition
Red blotch symptom quality assessment in Napa Valley Cabernet Sauvignon enabled differential diagnosis from the leafroll disease initially suspected based on late-season leaf reddening. Detailed examination of red blotch symptom quality revealed the diagnostic irregular blotchy pattern of reddening not conforming to the interveinal distribution characteristic of leafroll, with red patches extending across veins in angular patterns and some veins showing reddening while others remained green, contrasting with the consistent green-vein preservation of leafroll. The red blotch symptom quality timing showed color development beginning in July, earlier than typical leafroll symptom development, with color intensity variable across the canopy rather than the progressive intensification from basal to apical leaves seen in leafroll. Fruit red blotch symptom quality showed delayed ripening with affected vines at 22.5°Brix when adjacent healthy vines reached 25.0°Brix at equivalent harvest date, anthocyanin concentration reduced by 40%, and tannin development lagging normal ripeness parameters. Laboratory testing using PCR primers specific for GRBaV confirmed infection in 95% of vines showing red blotch symptom quality while testing negative for GLRaV-3, validating the field diagnosis based on symptom pattern differences. The red blotch symptom quality assessment established that the block was affected by this recently recognized pathogen rather than leafroll, informing management decisions about roguing priority and the potential value of treehopper vector management to limit secondary spread.
Red blotch symptom quality is a viral infection indicator quality that inheres in grapevine organisms infected by grapevine red blotch virus (GRBaV) and represents the characteristic disease manifestations including irregular red blotches on leaf blades of red-fruited cultivars appearing in late summer and autumn, chlorotic blotches on white-fruited cultivars, the distinctive pattern of reddening that does not follow vein boundaries distinguishing it from leafroll symptoms, delayed fruit ripening with reduced sugar accumulation, and diminished anthocyanin and flavor compound development in affected fruit that compromises wine quality even when visible crop effects are minimal. This quality captures the diagnostic features of a recently discovered viral disease first identified in 2008 that has subsequently been recognized as widespread and economically significant in North American viticulture, with retrospective analysis suggesting the virus has been present unrecognized for decades before its identification as a distinct pathogen. The ontological significance of red blotch symptom quality lies in its role enabling diagnosis of a disease that was previously misidentified as nutrient deficiency, leafroll, or other causes of late-season leaf reddening, with accurate recognition essential for appropriate management decisions and for preventing propagation spread through use of infected source material. The biological basis of red blotch symptoms encompasses the geminivirus replication of GRBaV affecting photosynthate production and transport, with the resulting carbohydrate accumulation in leaves triggering anthocyanin synthesis that produces the blotchy red coloration while simultaneously reducing sugar export to developing fruit causing the ripening delays and quality reductions. The transmission biology of GRBaV remains incompletely understood, with the three-cornered alfalfa treehopper (Spissistilus festinus) confirmed as a vector capable of transmission, but field spread rates varying widely among vineyards suggesting additional factors or vectors may be involved in natural transmission dynamics. The fruit quality impacts of red blotch infection include 1-5°Brix reduction at harvest equivalent timing, delayed achievement of target ripeness by 2-4 weeks, and reduced anthocyanin and flavor compound accumulation that affects wine color, body, and varietal character, with economic impacts estimated at 20-40% reduction in fruit value for affected blocks.
Red_Blotch_Symptom_Quality is_a Viral_Infection_Indicator_Quality, representing the symptoms of grapevine red blotch virus (GRBaV) infection. The irregular blotchy pattern crossing veins distinguishes from leafroll's interveinal pattern. Recently discovered (2008) but retrospectively present unrecognized for decades. Treehopper vector confirmed but transmission biology incompletely understood. Significant fruit quality impacts (1-5°Brix reduction, 2-4 week ripening delay). Previously misdiagnosed as leafroll, nutrient deficiency, or abiotic stress.
GRBaV symptoms
GRBaV-Symptome
Rebenrote-Flecken-Krankheit
Red Blotch-Virose
Rote-Flecken-Symptome
grapevine red blotch symptoms
red blotch disease symptoms
red blotch virus indicators
Sudarshana MR et al. (2015) Grapevine red blotch-associated virus, an emerging threat to the grapevine industry. Phytopathology 105:1026-1032 Cieniewicz EJ et al. (2017) Differential spread of grapevine red blotch virus in California and New York vineyards. Phytobiomes 1:203-214 Ricketts KD et al. (2017) Grapevine red blotch virus: an emerging disease in North America. Phytopathology 107:12-13
Red Blotch Symptom Quality
Red hue intensity in Pinot Noir provides a sensitive indicator of vintage conditions and vineyard site characteristics, with optimal fruit displaying bright ruby-red hue (CIELAB h° 15-30°, a* 40-55) that translates to the translucent garnet wines prized by Burgundy enthusiasts excessive heat during ripening shifts hue toward orange-red (h° 30-45°) through anthocyanin degradation and oxidation, producing wines with bricking color that signals premature aging potential, while cool vintages maintain the blue-red (h° 5-15°) characteristics of youthful Pinot Noir with greater aging capacity.
Red_Hue_Quality represents the chromatic character of grapevine tissues displaying coloration in the red portion of the visible spectrum, corresponding to wavelengths approximately 620-700 nm and CIELAB hue angles of approximately 0-40°. Red hue results from anthocyanin pigments in their flavylium cation form, which predominates at the low pH values typical of grape berry vacuoles. The specific red hue depends on the anthocyanin profile, with cyanidin-based pigments producing orange-red hues and malvidin-based pigments producing blue-red hues when present individually, though co-pigmentation and polymerization modify perceived color in complex ways. In grapevine, red hue characterizes young shoot tips and expanding leaves in many cultivars, berry skins of red and black grape varieties, and autumn senescing leaves where anthocyanin synthesis masks chlorophyll degradation products. Red-fruited cultivars such as Pinot Gris, Gewürztraminer, and many table grapes display pink to red-copper hues at maturity, representing moderate anthocyanin accumulation without the intensity to achieve purple or black coloration. Red hue in young leaves (OIV descriptor 004) provides an important ampelographic character, with cultivars classified by the distribution and intensity of red coloration on expanding shoots. Environmental factors strongly influence red hue expression, with high light promoting anthocyanin synthesis and cool temperatures favoring red over blue hue by stabilizing the flavylium cation form of anthocyanins. Temperature above 35°C inhibits anthocyanin synthesis and can cause degradation, reducing red coloration.
Red_Hue_Quality represents the red region of the color spectrum, positioned between yellow-orange and purple-blue hues. The boundary between red and purple hues is gradational, with assignment depending on the balance between red-reflecting and blue-reflecting wavelengths. In wine color terminology, red typically refers to bright, ruby colors while purple indicates younger wines with stronger blue components. PATO_0000322 (red) provides the upper-ontology reference. OIV descriptors for berry skin color distinguish rose (light red) from red and dark red categories.
Anthocyanrotfärbung
Rotton
anthocyanin red coloration
crimson hue
red color
rote Färbung
rubinrote Tönung
ruby hue quality
He F et al. (2012) Anthocyanins and Their Variation in Red Wines I. Monomeric Anthocyanins and Their Color Expression. Molecules 17(2): 1571-1601 Downey MO et al. (2006) Cultural Practice and Environmental Impacts on the Flavonoid Composition of Grapes and Wine: A Review of Recent Research. American Journal of Enology and Viticulture 57(3): 257-268
Red Hue Quality
Crimson Seedless bears a Red_Table_Grape_Scion_Role as the global standard for late-season red seedless, with medium-large berries (4-6 g), deep crimson coloration requiring sufficient heat accumulation (>2000 GDD), exceptional crunch, mild flavor, and 6-8 week storage life enabling global year-round supply from sequential harvests in California, Chile, South Africa, Peru, and Australia. Flame Seedless serves in a Red_Table_Grape_Scion_Role for early-season red markets, with smaller berries (3-4 g), bright flame-red coloration, and firm texture harvested in May-July in California and November-January in Chile. Red Globe bears a Red_Table_Grape_Scion_Role for Asian premium seeded markets, with very large berries (12-18 g), pink-red coloration, and seeds accepted in markets valuing berry size over seedlessness.
A fresh consumption table grape role that inheres in a grafted scion cultivar by virtue of its red to pink berry skin coloration, produced by anthocyanin pigments accumulating in the hypodermal cell layers during veraison and ripening. Red table grape scions develop coloration through cyanidin and peonidin-based anthocyanins (predominantly cyanidin-3-glucoside) requiring adequate temperature differential between day and night (ideally >10°C) for optimal pigment synthesis. Red coloration intensity and uniformity are critical market quality factors, with ideal berries showing complete, even pigmentation without green shoulders or uneven blush. Red table grape scions dominate global markets alongside white seedless varieties, with consumer perception associating red color with sweetness and ripeness. Major red table grape varieties include both seeded heritage cultivars (Red Globe, Emperor, Ribier) and modern seedless varieties (Crimson Seedless, Flame Seedless, Allison, Timco, Scarlotta Seedless) that constitute the majority of new breeding releases.
Red_Table_Grape_Scion_Role captures varieties where red/pink anthocyanin coloration is the defining visual characteristic, influencing cultivation requirements (color development demands specific temperature regimes) and market positioning. Color development challenges include insufficient diurnal temperature variation (common in tropical production), excessive canopy shading (reducing berry light exposure), and heat events during ripening (degrading anthocyanins). Breeding programs continuously develop improved red seedless varieties with enhanced color reliability, extended storage life, and proprietary flavor characteristics. The boundary between "red" and "black" categories is not absolute—some varieties (Autumn Royal, Fantasy) occupy the deep red-to-black spectrum and may be marketed in either category.
Rosé-Tafeltraube
pink table grape
red eating grape
red fresh grape
red table grape
rote Esstraube
rote Tafeltraube
Creasy GL, Creasy LL (2009) Grapes, CABI Peppi MC et al. (2006) Abscisic acid and anthocyanin accumulation in Crimson Seedless grapes, American Journal of Enology and Viticulture 57: 421-427 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press Crisosto CH et al. (2002) Table grape postharvest quality maintenance guidelines, UC Davis Postharvest Technology Research and Information Center
Red Table Grape Scion Role
Cabernet Sauvignon bears a Red_Wine_Scion_Role as the world's most planted red wine variety, providing deep color (anthocyanins 1200-1800 mg/kg), firm tannins (proanthocyanidins 2.5-4.0 g/kg), and age-worthy structure across Bordeaux, Napa Valley, Coonawarra, and global production regions. Pinot Noir serves in a Red_Wine_Scion_Role producing lighter-colored, lower-tannin wines (anthocyanins 400-800 mg/kg, proanthocyanidins 1.0-2.0 g/kg) with delicate aromatics in Burgundy, Oregon, and Central Otago. Syrah/Shiraz bears a Red_Wine_Scion_Role creating deeply pigmented, full-bodied wines with pepper and dark fruit character in the Northern Rhône and Barossa Valley.
A wine production scion role that inheres in a grafted scion cultivar by virtue of its designation for red wine production, where pigmented grape skins are macerated with fermenting juice to extract anthocyanins, tannins, and other phenolic compounds that define red wine color, structure, and aging potential. Red wine scions are characterized by dark-pigmented berry skins (blue-black to deep purple) containing anthocyanin concentrations typically ranging from 500-2000 mg/kg berry weight, with anthocyanin profiles (malvidin, peonidin, delphinidin, petunidin, cyanidin glycosides) varying by cultivar and contributing to color hue and stability. Red wine scions also accumulate proanthocyanidins (condensed tannins) in skins and seeds (1-4 g/kg), providing astringency, mouthfeel, and antioxidant protection enabling extended aging. Skin contact during fermentation (typically 5-30 days) extracts these compounds the duration and temperature of maceration profoundly influence wine style. Red wine scions may be vinified as single-varietal wines or blended with other red cultivars to achieve complexity, balance, or house style consistency.
Red_Wine_Scion_Role is subclassed by quality tier (Premium, Bulk) and functional specialization (Teinturier for color contribution, Blending for assemblage components). The distinction between red wine scions and rosé production (where the same cultivars receive minimal skin contact) is one of vinification method rather than cultivar genetics, though certain varieties (Grenache, Cinsault) are particularly associated with rosé. Anthocyanin chemistry varies significantly among red wine scions Pinot Noir contains only non-acylated monoglucosides (contributing to color instability), while Cabernet Sauvignon, Syrah, and many others contain acylated anthocyanins providing greater color stability. Tannin quantity and quality (seed-derived versus skin-derived) also varies substantially among cultivars.
Rotweinrebe
Rotweinsorte
black grape variety
blaue Traube
red grape variety
red wine cultivar
red wine variety
rote Rebsorte
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Jackson RS (2014) Wine Science 4th Ed, Academic Press pp.: 270-350 Kennedy JA (2008) Grape and wine phenolics: observations and recent findings, Ciencia e Investigación Agraria 35: 107-120 Waterhouse AL et al. (2016) Understanding Wine Chemistry, Wiley
Red Wine Scion Role
A reduced cluster secondary bud at node 9 on Vitis vinifera cv. Cabernet Sauvignon, activated following hail damage destroying primary shoots, dissection revealing one small inflorescence primordium containing only 38 flower initials (compared to 165-185 typical for primary buds at this node position), developing into a shoot with one loose cluster containing 14 berries after 37% fruit set, plus 12 leaves providing 680 cm² leaf area, resulting in exceptional 48.6 cm²/berry leaf area-to-fruit ratio that drove early maturation to 25.8° Brix by September 15th (12 days before normal harvest), though the tiny 95g cluster contributed minimally to commercial yield and required separate harvest timing decisions.
A secondary reproductive bud containing inflorescence primordia of reduced size and flower number compared to typical primary bud clusters, characterized by small inflorescence initials with 30-80 flower primordia (versus 100-300 in primaries), development into shoots bearing small, loose clusters with low berry numbers (typically 15-45 berries after fruit set), and frequent occurrence even in cultivars with generally low secondary fruitfulness where occasional secondary positions achieve minimal reproductive differentiation. Reduced cluster secondary buds provide limited but non-zero crop contribution following primary bud damage, with their small clusters often showing accelerated ripening due to favorable source-sink ratios (high leaf area relative to small fruit mass), though commercial value may be reduced by small cluster size, loose structure, and harvest timing mismatches with normally-developing primary shoots in mixed-damage scenarios.
Reduced cluster secondary buds represent the lower end of secondary fruitfulness spectrum, providing token rather than substantial yield recovery but occasionally achieving superior fruit quality through favorable vine balance. The small berry numbers create extremely high leaf-to-fruit ratios that can accelerate ripening and enhance phenolic development, sometimes producing exceptional individual clusters despite poor overall yield recovery. However, the loose cluster structure and small size may create harvest logistics challenges, and in mixed-damage scenarios where some primary shoots survive, the phenological asynchrony between early-ripening reduced secondary clusters and normally-developing primary clusters complicates harvest timing decisions. Some viticulturists remove reduced secondary clusters entirely to focus resources on remaining primary production, while others retain them for their quality potential despite minimal quantity contribution.
Kleintrauben-Nebenknospe
low-flower secondary bud
minimal secondary reproductive bud
reduzierte Trauben-Sekundärknospe
small-cluster secondary bud
Based on Dunn & Martin (2000) A functional association in Vitis vinifera L. cv. Cabernet Sauvignon between the extent of primary branching and the number of flowers formed per inflorescence. Australian Journal of Grape and Wine Research 6=88-92 Kliewer & Dokoozlian (2005), Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Petrie et al. (2000) "The effect of leaf removal and canopy height on whole-vine gas exchange and fruit development of Vitis vinifera L. Sauvignon Blanc. Functional Plant Biology 27=569-578.
Reduced Cluster Secondary Bud
Wild V. vinifera ssp. sylvestris male plants in Mediterranean populations commonly exhibit reduced pistil flowers with vestigial pistil remnants 0.2-0.5 mm long visible as small protuberances in the flower center surrounded by five robust stamens (filaments 3-4 mm, anthers 0.7-0.9 mm), producing 28,000-40,000 pollen grains per flower with 75-88% viability, achieving 0% fruit set despite pistil presence, and functioning as effective pollen donors indistinguishable in male reproductive function from male only flowers despite morphological pistil retention. Breeding populations from wild × cultivated crosses segregate reduced pistil staminate individuals showing variable pistil reduction from vestigial (0.3 mm) to rudimentary (0.6 mm), all uniformly non-fruit-bearing and requiring removal from breeding programs. North American wild Vitis species (V. labrusca, V. aestivalis) male plants typically bear reduced pistil flowers with rudimentary pistils 0.4-0.7 mm long showing recognizable ovary, style, and stigma regions though all non-functional.
A reduced pistil flower is a staminate flower that retains a vestigial, rudimentary, or severely abbreviated pistil structure rather than showing complete pistil absence, with the retained pistil structure ranging from barely recognizable tissue remnants to more obvious though still drastically reduced pistil morphology, in all cases completely incapable of functional fertilization and fruit set but providing morphological evidence of suppressed rather than completely eliminated pistil developmental programming. Reduced pistil flowers are characterized by= (1) presence of a recognizable though severely reduced central structure in the position where a functional pistil would occur in hermaphrodite flowers, with this structure showing some minimal pistil-like features (rudimentary ovary swelling, vestigial style elongation, or minute stigmatic tissue) though at drastically reduced scale (<0.3-0.8 mm total pistil length compared to 1.5-2.5 mm in functional pistils) (2) pistil structures showing arrested development at various early stages, from extremely early arrest producing undifferentiated vestigial remnants to somewhat later arrest producing rudimentary organs with recognizable but non-functional pistil components (3) well-developed stamens showing typical staminate flower characteristics (erect position, 2.5-4 mm filaments, 0.6-1 mm anthers, abundant pollen production 25,000-45,000 grains per flower) surrounding the reduced central pistil (4) complete functional male sterility of the pistil due to absence of receptive stigma, non-functional or absent style, and aborted or absent ovules, resulting in 0% fruit set despite morphological pistil presence. The reduced pistil condition represents incomplete suppression of pistil development during flower organogenesis, with pistil primordia initiating and undergoing some early developmental progression before arrest, in contrast to male only flowers where pistil development is completely prevented or arrested at such early stages that no recognizable pistil structures form. The degree of pistil reduction varies along a continuum from vestigial (barely recognizable remnants showing minimal organ differentiation, 0.2-0.4 mm length) to rudimentary (clearly recognizable as pistil structures though severely abbreviated, 0.4-0.8 mm length), with this variation possibly reflecting= (1) genetic variation in strength of pistil-suppressing alleles, with weak suppressors allowing more pistil development than strong suppressors (2) environmental modulation during flower development, with favorable conditions potentially permitting more pistil development than stressful conditions (3) stochastic developmental variation creating flower-to-flower or plant-to-plant variation in degree of pistil suppression. Reduced pistil flowers achieve essentially zero fruit set (<0.1%) despite pistil presence, as the drastically abbreviated structures completely lack functional female reproductive capability= stigmas are absent or non-receptive, styles lack transmitting tract tissue, ovaries are empty or contain aborted ovules, and the entire structure is incapable of supporting pollination, fertilization, or embryo development. Reduced pistil flowers are characteristic of a substantial proportion (50-70%) of male individuals in wild dioecious V. vinifera ssp. sylvestris populations, with the remaining 30-50% of males bearing male only flowers with no pistil trace, suggesting that reduced pistil represents the more common or less stringently selected staminate flower morphology. The evolutionary and developmental significance of pistil retention in reduced pistil flowers is debated, with hypotheses including= (1) incomplete evolution toward complete male unisexuality, with reduced pistils representing transitional stages in the evolutionary loss of female function (2) developmental constraint preventing complete pistil elimination despite strong selection for male unisexuality (3) possible residual adaptive value of vestigial pistils through pheromone production, nectar secretion, or structural roles despite reproductive non-functionality.
Reduced_Pistil_Flower represents incomplete pistil suppression among staminate flowers and is distinguished from Male_Only_Flower (no recognizable pistil) by retention of vestigial or rudimentary pistil structures, though both types are functionally equivalent in achieving zero fruit set and serving exclusively as pollen sources. The distinction is morphological rather than functional, but understanding the variation in degree of pistil suppression informs developmental genetics studies of sex determination and organ suppression mechanisms. Sub-classification into Vestigial_Pistil (minimal remnants, 0.2-0.4 mm) versus Rudimentary_Pistil (recognizable though abbreviated structures, 0.4-0.8 mm) reflects gradations in the degree of developmental arrest and pistil retention. The evolutionary significance of pistil retention in male flowers despite complete functional sterility remains an open question deserving investigation.
Blüte mit verkümmertem Stempel
Staminatblüte mit Pistillrest
reduzierte Pistill Blüte
rudimentary-pistil male flower
vestigial-pistil staminate flower
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 101-105 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Caporali, E. et al. (2003). The arrest of development of abortive reproductive organs. Sexual Plant Reproduction 15=291-300 Picq, S. et al. (2014). A small XY chromosomal region explains sex determination. BMC Plant Biology 14=229.
Reduced Pistil Flower
Certain Gewürztraminer clones exhibit reduced pollen functionally female expression with only moderate stamen reflexing but severely reduced pollen production (4,000-7,000 grains per flower) and low viability (8-15%), requiring cross-pollination from hermaphrodite neighbors to achieve 28-35% fruit set, with isolated Gewürztraminer blocks showing <12% fruit set and uneconomical yields. Orange Muscat shows reduced pollen functional female flowers with near-erect stamens but essentially sterile pollen (<2,000 viable grains per flower), creating absolute cross-pollination dependency despite superficially hermaphrodite-like stamen position. Some Muscat of Alexandria clones exhibit variable pollen production (3,000-9,000 grains per flower) with poor viability (12-22%), showing year-to-year and site-to-site variation in self-fertility corresponding to environmental effects on pollen development.
A reduced pollen functionally female flower is a functionally female flower in which male sterility is achieved primarily through severely reduced pollen production quantity or drastically reduced pollen viability rather than through stamen reflexing, with stamens showing erect or only slightly reflex position but producing insufficient viable pollen for effective self-pollination, requiring cross-pollination from hermaphrodite sources for fruit set. Reduced pollen functionally female flowers are characterized by= (1) stamens that may be erect, partially erect, or only moderately reflex (not the pronounced reflexing seen in reflex stamen types), sometimes approaching hermaphrodite stamen position but distinguishable by pollen characteristics (2) severely reduced pollen production, typically <5,000-8,000 pollen grains per flower compared to 15,000-30,000 in hermaphrodite flowers, resulting from impaired anther development, reduced pollen mother cell numbers, or disrupted microsporogenesis (3) drastically reduced pollen viability, often <10-20% viable pollen compared to 60-85% in hermaphrodite flowers, resulting from defective pollen grain development, abnormal exine formation, or cytoplasmic male sterility (4) the combination of reduced pollen quantity and quality creating functional male sterility even when spatial positioning might theoretically permit self-pollination. The reduced pollen mechanism represents gametic rather than morphological male sterility, with the failure occurring at the cellular level of pollen development rather than through organ positioning. Pollen reduction can result from several developmental mechanisms= arrested development of some or all pollen mother cells during microsporogenesis, abnormal meiotic divisions producing non-viable dyads or triads instead of viable tetrads, defective pollen wall formation creating pollen grains unable to survive desiccation or germinate on stigmas, or cytoplasmic genetic factors (cytoplasmic male sterility) disrupting pollen maturation. Reduced pollen functionally female flowers require cross-pollination for fruit set, as the minimal viable pollen produced is insufficient to saturate stigmatic surfaces and ensure fertilization success, with fruit set depending on adequate pollen delivery from hermaphrodite pollinizer sources. Fruit set percentages typically range from 20-35% under good cross-pollination conditions but may fall to <8-12% under poor pollination conditions or in isolated plantings lacking adequate pollinizer proximity. The reduced pollen mechanism may be less morphologically obvious than reflex stamens, requiring microscopic pollen examination or fruit set observation trials to definitively diagnose, as flowers may appear superficially hermaphrodite in stamen position but fail to self-pollinate effectively. Some cultivars exhibit combined mechanisms, showing both moderate stamen reflexing and reduced pollen production/viability, creating redundant male sterility systems. The reduced pollen condition can show environmental modulation, with temperature stress, nutritional imbalances, or water stress during microsporogenesis potentially exacerbating pollen reduction or inducing reduced pollen expression in cultivars that might show better pollen production under optimal conditions.
Reduced_Pollen_Functionally_Female_Flower is distinguished from Reflex_Stamen_Functionally_Female_Flower by the primary mechanism of male sterility= gametic (pollen) dysfunction versus physical organ positioning, though these mechanisms can co-occur. The reduced pollen mechanism is less morphologically obvious and may require quantitative pollen counts and viability assays for definitive diagnosis, whereas reflex stamens are readily identified visually. Reduced pollen types may be misidentified as hermaphrodite in casual observation if stamen position appears normal, leading to vineyard design errors and inadequate pollinizer provision. The genetic and developmental basis of pollen reduction is complex and may involve nuclear genes affecting microsporogenesis, cytoplasmic factors creating cytoplasmic male sterility, or environmental disruption of temperature-sensitive pollen development pathways. Understanding pollen reduction mechanisms could inform breeding programs seeking to introduce or eliminate male sterility for specific purposes (e.g., introducing male sterility for hybrid seed production, eliminating it for self-fertile cultivar development).
Blüte mit Pollenreduktion
low-pollen female flower
pollen-deficient flower
pollenmangel weibliche Blüte
pollenreduzierte Blüte
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 99-102 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Caporali, E. et al. (2003). The arrest of development of abortive reproductive organs. Sexual Plant Reproduction 15=291-300 Costantini, L. et al. (2008). Berry and phenology-related traits in grapevine= From quantitative trait loci to underlying genes. BMC Plant Biology 8=38.
Reduced Pollen Functionally Female Flower
A reduced secondary reproductive shoot on Vitis vinifera cv. Cabernet Sauvignon from node 9, activated following severe spring frost (-4.8°C) with 92% primary mortality, achieved only 72cm length by late July, 9 small leaves totaling approximately 1080 cm² leaf area, bearing single very small cluster with 18 berries, estimated shoot crop 94g representing approximately 18% of normal primary production from this position (primaries typically 480-560g from two clusters), representing severely compromised secondary compensation attributed to combination of catastrophic frost damage depleting vine reserves, Cabernet Sauvignon's inherently weak secondary fruitfulness, cool poor growing season following frost limiting recovery, and excessive competition from widespread secondary activation across 92% of positions overwhelming vine capacity, with this minimal crop contribution and poor vine condition triggering assessment of vintage viability and potential abandonment decisions recognizing inadequate economic return prospects.
A secondary reproductive shoot exhibiting minimal reproductive performance, characterized by weak activation and development, limited final size (60-95cm typical), sparse foliage (8-12 leaves, 950-1400 cm² total area), single small cluster or occasionally complete reproductive failure despite secondary bud origin nominally containing inflorescence primordia, and crop contribution minimal (80-150g per shoot) representing 15-35% of typical primary production, indicating severely compromised compensation under unfavorable conditions following damage events. Reduced secondary reproductive shoots demonstrate poor compensation from backup positions, providing minimal crop recovery following primary loss, contributing little to economic salvage, and often showing questionable fruit quality due to inadequate photosynthetic support, delayed maturation, and stress indicators, with such poor secondary performance typically associated with severe widespread damage, unfavorable post-damage conditions, or inherently weak secondary fruitfulness in certain varieties.
Reduced secondary reproductive shoots indicate severely compromised recovery following damage, with poor secondary performance signaling inadequate compensation threatening vintage economic viability. Causes include severe widespread damage (>80% primary loss creating overwhelming resource demands), variety characteristics (Cabernet Sauvignon, Syrah, Nebbiolo showing characteristically weak secondary fruitfulness even under favorable conditions), unfavorable post-damage conditions (cool temperatures, drought, nutrient limitations impeding recovery), and compounding stress (pre-existing vine weakness, disease pressure, previous-season stress depleting reserves). Management involves realistic assessment of recovery prospects (determining whether reduced secondary compensation provides viable crop or abandonment warranted), possible crop reduction (removing even limited secondary crop to support vegetative recovery prioritizing next season), supplemental inputs (irrigation, nutrition supporting recovery), and long-term planning (evaluating damage patterns, considering protective measures, assessing replanting needs if chronic damage). Reduced secondary shoots provide minimal current economic value and generally unsuitable renewal wood.
minimal-yield secondary shoot
poor-compensation shoot
reduzierter sekundärer Fruchttrieb
schwacher Ersatz-Ertragstrieb
weak secondary reproductive shoot
Based on Molitor et al. (2014) "Late frost damage risk for viticulture under future climate conditions= a case study for the Luxembourgish winegrowing region" Australian Journal of Grape and Wine Research 20=160-168 Ferguson et al. (2011) "Dynamic changes in bud cold hardiness during dormancy" Journal of the American Society for Horticultural Science 136=313-323.
Reduced Secondary Reproductive Shoot
Pinot Noir functionally male flowers exhibit reduced short styles 0.32-0.42 mm connecting reduced ovaries 0.58-0.75 mm to reduced stigmas 0.28-0.38 mm, with reduced short styles representing 28-35% of total pistil length (1.2-1.6 mm), containing transmitting tract tissue with recognizable central canal and modest extracellular matrix production visible in periodic acid-Schiff-stained sections, achieving 4-12% fruit set primarily limited by stigma non-receptivity (stigmatic exudate production <20% of hermaphrodite levels) and ovule abortion (only 1.2-1.8 viable ovules per ovary versus 3.2-3.8 in hermaphrodite flowers) rather than by style inadequacy. Cabernet Sauvignon occasional functionally male segregants in breeding populations show reduced short styles 0.35-0.44 mm as part of moderately reduced pistils (1.3-1.7 mm total) maintaining recognizable style structure distinct from very short vestigial styles (<0.3 mm) in staminate types, with transmitting tract showing partial differentiation supporting limited pollen tube growth in controlled pollination experiments though fertilization success remains <8% due to ovule defects. Syrah functionally male flowers display reduced short styles 0.3-0.4 mm that are readily distinguishable without magnification as distinct intermediate zones between ovary and stigma, contrasting with very short vestigial styles in staminate flowers that require magnification for clear identification, with the reduced short style morphology allowing use as diagnostic feature for identifying functionally male versus staminate flower types in field assessments. Breeding studies crossing functionally male × hermaphrodite parents produce F1 progeny segregating for pistil morphology, with functionally male offspring showing reduced short styles 0.28-0.42 mm while hermaphrodite offspring show medium-length styles 0.48-0.68 mm, demonstrating genetic control of style length variation and allowing style length to serve as early indicator of flower sexual type during seedling evaluation.
A reduced short style is a short style in the moderate length range within the short category, typically 0.25-0.45 mm at maturity, characteristic of functionally male flowers and representing styles that show clear reduction compared to hermaphrodite norms but retain more substantial development than very short vestigial styles, maintaining recognizable style structure and some transmitting tract differentiation. Reduced short styles are characterized by= (1) length typically 0.25-0.45 mm at anthesis, representing 40-70% of typical hermaphrodite style length and substantially exceeding very short vestigial style length (<0.3 mm) (2) diameter typically 0.12-0.25 mm maintaining cylindrical morphology (3) transmitting tract tissue present with recognizable central canal and some extracellular matrix production, though typically showing reduced elaboration compared to fully functional hermaphrodite styles (4) connection between reduced ovary (typically 0.5-0.8 mm) and reduced or rudimentary stigma (typically 0.25-0.4 mm) in functionally male flower contexts, or occasional connection of normal organs in hermaphrodite variants. Reduced short styles represent moderate pistil reduction characteristic of functionally male flowers where female organ suppression is substantial but not extreme, allowing retention of recognizable pistil structure including a clearly defined style component despite overall size reduction. Reduced short styles develop through= (1) genetic factors creating functionally male phenotype with moderate pistil reduction (2) developmental processes that suppress but do not eliminate style elongation during pistil development (BBCH 53-57) (3) environmental or nutritional factors potentially modulating style development in genetically predisposed flowers. Reduced short styles in functionally male flowers maintain proportionality within the reduced pistil, representing approximately 25-40% of total pistil length (total pistil typically 1-1.8 mm with reduced short style 0.3-0.5 mm) similar to proportions in hermaphrodite pistils, suggesting that style reduction is part of coordinated overall pistil size reduction rather than disproportionate style-specific suppression. The functional implications of reduced short styles relate to partial female reproductive capacity= reduced short styles contain transmitting tract tissue capable of supporting some pollen tube growth if pollen were to germinate, though transmitting tract quality is typically suboptimal with reduced extracellular matrix production and less elaborate cellular specialization compared to fully functional styles the primary limitation to fertilization in flowers with reduced short styles is typically stigma non-receptivity and ovule abortion rather than style inadequacy per se, though reduced style length and quality may contribute modestly to reduced fertilization efficiency flowers with reduced short styles achieve <5-15% fruit set in functionally male contexts, with reproductive failure due primarily to stigma and ovule defects rather than to style characteristics alone. Reduced short styles require shorter pollen tube growth distances (total stigma-to-ovule distance approximately 1-1.6 mm) compared to hermaphrodite flowers (1.5-2.5 mm), potentially allowing pollen tube transit in 30-55 hours versus 48-96 hours, though this theoretical time advantage is negated by stigma and ovule dysfunction in functionally male flowers.
Reduced_Short_Style represents moderate expression within the short style category and is distinguished from Very_Short_Style (0.45 mm, typical hermaphrodite range) by clear reduction. Reduced short styles are characteristic of functionally male flowers and represent pistil reduction sufficient to impair but not eliminate female function. Understanding reduced short styles is relevant for identifying functionally male flowers, for assessing the continuum of pistil reduction from hermaphrodite through functionally male to staminate types, and for genetic studies of sex determination and pistil development.
moderately short style
mäßig kurzer Stylus
partially reduced style
reduzierter kurzer Griffel
teilweise verkürzter Stempelgriffel
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 94-98 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=93-105.
Reduced Short Style
Wild V. vinifera ssp. sylvestris female plants in Mediterranean and Near Eastern populations commonly exhibit reduced stamen flowers with vestigial or rudimentary stamen remnants 0.3-1.2 mm long showing strong downward reflexing and positioned well below the prominent pistil (2.3-2.5 mm length, ovary 1.3-1.5 mm diameter), producing minimal or no pollen (<1,000 grains per flower when present, <10% viability), requiring cross-pollination from nearby staminate males to achieve 28-38% fruit set comparable to female only flowers despite stamen presence, and providing material for developmental studies of organ suppression mechanisms. Breeding populations from wild dioecious × cultivated hermaphrodite crosses segregate reduced stamen pistillate individuals showing variable stamen reduction from vestigial (0.3 mm) to rudimentary (1 mm), all uniformly requiring cross-pollination and unsuitable for self-fertile cultivar development despite morphological stamen retention. Wild North American Vitis species female plants (V. labrusca, V. aestivalis) commonly bear reduced stamen flowers with rudimentary stamens 0.6-1.3 mm showing recognizable filament and anther regions though strongly reflex and functionally sterile.
A reduced stamen flower is a pistillate flower that retains vestigial, rudimentary, or severely abbreviated stamen structures rather than showing complete stamen absence, with the retained stamen structures ranging from barely recognizable tissue remnants to more obvious though drastically reduced stamen morphology showing strong reflexing and minimal or absent pollen production, in all cases completely incapable of effective self-pollination and requiring cross-pollination for fruit set but providing morphological evidence of suppressed rather than completely eliminated stamen developmental programming. Reduced stamen flowers are characterized by= (1) presence of recognizable though severely reduced structures in some or all of the five stamen positions where functional stamens would occur in hermaphrodite flowers, with these structures showing some minimal stamen-like features (rudimentary filaments, vestigial anther tissue) though at drastically reduced scale (<0.3-1.5 mm total stamen length compared to 2.5-4 mm in functional hermaphrodite stamens) (2) stamen structures showing arrested development at various early stages, from extremely early arrest producing undifferentiated vestigial remnants (0.2-0.5 mm) to somewhat later arrest producing rudimentary organs (0.5-1.5 mm) with recognizable filament and anther regions though non-functional (3) stamens characteristically showing strong downward reflexing when filament development is sufficient to permit curvature, positioning any anther-like structures well below the prominent central pistil (4) pollen production completely absent or drastically reduced to <500-3,000 grains per flower (compared to 15,000-30,000 in hermaphrodite flowers) with very low viability (<5-15%), creating functional male sterility despite morphological stamen presence (5) well-developed functional pistil (2-2.6 mm length, ovary 1.2-1.6 mm diameter with 3-4 viable ovules, receptive wet stigma) dominating the flower morphology and providing complete female function. The reduced stamen condition represents incomplete suppression of stamen development during flower organogenesis, with stamen primordia initiating and undergoing some early developmental progression before arrest, in contrast to female only flowers where stamen development is completely prevented or arrested at such early stages that no recognizable stamen structures form. The degree of stamen reduction varies along a continuum from vestigial (barely recognizable remnants showing minimal organ differentiation, 0.2-0.5 mm length) to rudimentary (clearly recognizable as stamen structures though severely abbreviated and strongly reflex, 0.5-1.5 mm length), with this variation possibly reflecting genetic variation in strength of stamen-suppressing alleles, environmental modulation during flower development, or stochastic developmental variation. Reduced stamen flowers achieve fruit set only through cross-pollination from staminate or hermaphrodite pollen sources, with fruit set percentages (25-40% under good pollination conditions) comparable to female only flowers, as the minimal non-viable pollen produced by vestigial or rudimentary stamens is completely insufficient for self-fertilization and the strong stamen reflexing prevents any pollen that is produced from reaching the stigma. The reduced stamen condition creates intermediate morphology between functionally female flowers (which have more developed though strongly reflex stamens and may produce marginal pollen permitting rare self-fertilization in exceptional circumstances) and female only flowers (which completely lack recognizable stamens), occupying a position on the continuum of stamen suppression. Reduced stamen flowers are characteristic of a substantial proportion (50-70%) of female individuals in wild dioecious V. vinifera ssp. sylvestris populations, with the remaining 30-50% of females bearing female only flowers with complete stamen absence, suggesting that reduced stamens represent the more common or less stringently selected pistillate flower morphology. The evolutionary and developmental significance of stamen retention in reduced stamen flowers parallels that of pistil retention in reduced pistil staminate flowers, potentially representing incomplete evolution toward complete unisexuality, developmental constraints, or possible residual functions despite reproductive non-functionality.
Reduced_Stamen_Flower represents incomplete stamen suppression among pistillate flowers and is distinguished from Female_Only_Flower (no recognizable stamens) by retention of vestigial or rudimentary stamen structures, though both types are functionally equivalent in requiring cross-pollination and achieving comparable fruit set. The reduced stamen condition must also be carefully distinguished from Functionally_Female_Flower (which has more developed though reflex stamens typically 1.5-3 mm capable of producing marginal functional pollen permitting rare self-fertilization). Sub-classification into Vestigial_Stamen (minimal remnants, 0.2-0.5 mm) versus Rudimentary_Stamen (recognizable though abbreviated structures, 0.5-1.5 mm) reflects gradations in developmental arrest and stamen retention, paralleling the classification of reduced pistil staminate flowers. Understanding stamen reduction mechanisms informs developmental genetics of sex determination and organ suppression.
Blüte mit verkümmerten Staubblättern
Pistillatblüte mit Stamenrest
reduzierte Staubblatt Blüte
rudimentary-stamen pistillate flower
vestigial-stamen female flower
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 102-106 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Caporali, E. et al. (2003). The arrest of development of abortive reproductive organs. Sexual Plant Reproduction 15=291-300 Badouin, H. et al. (2020). The wild grape genome sequence provides insights into the transition from dioecy to hermaphroditism. Genome Biology 21=223.
Reduced Stamen Flower
Reducing sugar quality analysis in German Riesling from the Mosel employs specific enzymatic determination of glucose and fructose rather than refractometry, enabling precise classification into Prädikat levels based on must weight: Kabinett (67-82° Oechsle, approximately 150-180 g/L reducing sugar), Spätlese (76-90° Oechsle, 170-210 g/L), Auslese (83-100° Oechsle, 190-240 g/L), and higher categories, with the official measurement defining quality designation, pricing, and label claims under German wine law.
Reducing_Sugar_Quality represents the chemical quality characterizing the concentration of sugars capable of reducing oxidizing agents due to the presence of free aldehyde or ketone groups, encompassing glucose (an aldose) and fructose (a ketose) that together constitute the dominant sugars in ripe grape berries. The designation "reducing" refers to the ability of these sugars to reduce Benedict's or Fehling's reagent, a property historically used for their quantification that distinguishes them from sucrose and other sugars where the reducing groups are blocked by glycosidic bonds. In mature Vitis vinifera berries, reducing sugars typically comprise greater than 99% of total sugars, with glucose and fructose present in approximately equal concentrations (1:1 ratio), each contributing roughly 50% of the total. This near-equimolar ratio results from the invertase-mediated hydrolysis of transported sucrose, which cleaves the disaccharide into one glucose and one fructose molecule. The absolute concentration of reducing sugars at harvest typically ranges from 150-250 g/L, with the proportion of glucose versus fructose varying slightly among cultivars and with ripening conditions. Glucose is preferentially fermented by Saccharomyces cerevisiae, so wines from arrested or stuck fermentations tend to have higher residual fructose than glucose. For dry wine production, complete fermentation of reducing sugars to below 2-4 g/L is typical, while sweet wines retain significant reducing sugar concentrations. The reducing sugar concentration provides the direct measure of fermentable substrate, making it more relevant than total soluble solids for precise fermentation planning.
Reducing_Sugar_Quality encompasses glucose and fructose, which are further specified in child classes Glucose_Content_Quality and Fructose_Content_Quality. The term reducing sugar is standard in enological chemistry, though the term fermentable sugar is often used synonymously in practical contexts. Reducing sugar concentration can be measured by non-specific methods (copper reduction, polarimetry) or specific enzymatic methods (hexokinase/glucose-6-phosphate dehydrogenase coupled assays). The near-equal glucose-fructose ratio in ripe berries can shift during over-ripening or in certain pathological conditions, providing diagnostic information beyond the total reducing sugar value.
Hexosengehalt
Invertzucker
fermentable sugar
glucose-fructose total
hexose content
invert sugar quality
reduzierende Zucker
vergärbare Zucker
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed, John Wiley and Sons pp.: 1-51 Kliewer WM (1967) The Glucose-Fructose Ratio of Vitis vinifera Grapes. American Journal of Enology and Viticulture 18(1): 33-41
Reducing Sugar Quality
Reduction aroma assessment in young Syrah identifies initial struck match character (intensity 4.0 on 10-point scale) considered acceptable for this variety's typical reductive presentation, with underlying rotten egg (2.5) and faint onion (1.8) suggesting H₂S at 3.5 μg/L and ethanethiol at 0.8 μg/L, with bench trial demonstrating copper fining at 0.5 mg/L eliminates rotten egg while preserving the struck match character valued in this style, illustrating the nuanced approach to reduction management where complete elimination is neither necessary nor desirable for varieties showing positive reductive character.
Reduction aroma quality is an off-aroma quality that represents olfactory perception of volatile sulfur compounds formed under low oxygen conditions, characterizing the range of reductive off-odors from mildly unpleasant struck match through increasingly objectionable rotten egg, cabbage, onion, and garlic notes depending on the specific sulfur species present. Within the BFO framework, this quality specifically addresses the reductive fault category arising from sulfur compound accumulation, distinguishing reduction-derived off-aromas from other fault types while recognizing the complexity of reductive character where mild reduction may be considered stylistic (the reductive winemaking approach) while severe reduction constitutes clear defect. The ontological significance lies in reduction representing a fault category with unusually complex quality implications, where low-level volatile sulfur compounds may contribute to aromatic complexity and protect against oxidation (the intentional reduction of many fine wines), while elevated concentrations create unmistakably objectionable character, with the boundary between positive and negative highly dependent on compound identity, concentration, and taster sensitivity. The compositional basis for reduction aroma includes a hierarchy of volatile sulfur compounds with different sensory impacts and thresholds, from hydrogen sulfide (H₂S, rotten egg, threshold 1-2 μg/L) formed during fermentation from sulfur residues or yeast stress, through methanethiol (MeSH, rotten cabbage, threshold 0.3 μg/L), to heavier mercaptans (ethanethiol, onion-garlic) and disulfides (dimethyl disulfide, cooked cabbage) formed through H₂S reaction with wine components. The formation pathways for reductive compounds include yeast metabolism during fermentation (H₂S from sulfate reduction or cysteine degradation), post-fermentation chemical reactions (mercaptan formation from H₂S and reactive carbonyls), and degradation of sulfur-containing amino acids under reductive conditions. The remediation possibilities distinguish reduction from other faults, with copper sulfate addition effectively binding H₂S and light mercaptans while heavier sulfur compounds prove more resistant, and aeration sometimes effective for light reduction while risking oxidation damage to other wine components. Understanding reduction aroma quality enables fault identification and compound-specific diagnosis, remediation selection based on sulfur species involved, and winemaking management of the reduction-oxidation balance for optimal aromatic development.
Reduction_Aroma_Quality addresses the volatile sulfur compound fault category, which uniquely spans both positive (intentional reduction contributing complexity) and negative (objectionable mercaptan character) quality space. This quality requires compound-specific assessment as different sulfur species have different thresholds, remediation options, and quality implications. The class supports the nuanced approach to reduction management essential in contemporary winemaking.
Böckser
Reduktionsnoten
Schwefelverbindungen
mercaptan fault
reductive character
reduktiver Fehlton
sulfide off-odor
sulfur compounds
Siebert TE et al. (2010) Volatile compounds related to reductive character in wine. In: Blair RJ et al. (eds) Proceedings of the 14th Australian Wine Industry Technical Conference, Adelaide, pp. 76-84 Rauhut D (1993) Yeasts — production of sulfur compounds. In: Fleet GH (ed) Wine Microbiology and Biotechnology, Harwood Academic, Chur, pp. 183-223
Reduction Aroma Quality
Cabernet Sauvignon clone ENTAV-INRA 15 maintained at INRAE Domaine de Vassal bearing a reference cultivar role serves as the international standard genotype for SSR marker calibration across 9 microsatellite loci (VVS2, VVMD5, VVMD7, VVMD25, VVMD27, VVMD28, VVMD32, VrZAG62, VrZAG79), with its allele sizes used as reference values by 50+ laboratories worldwide to ensure inter-laboratory comparability of grapevine DNA fingerprinting results, and its OIV descriptor profile (455 morphological descriptors) constituting the type description against which Cabernet Sauvignon identity claims are validated.
A research and education role that inheres in a grapevine designated as an authoritative standard for cultivar identification, trait characterization, or comparative evaluation, serving as a benchmark against which other accessions, clones, or experimental genotypes are measured within systematic ampelographic, genetic, or phenological assessment frameworks.
Reference_Cultivar_Role captures the epistemic function of specific accessions as measurement standards within the grapevine research community. The role requires institutional designation and ongoing curatorial maintenance to ensure the reference vine's authenticity, health status, and accessibility: loss or contamination of reference material has significant consequences for the reproducibility of cultivar identification across the global research network. Subclasses distinguish between type specimens (nomenclatural standards for cultivar identity) and standard references (functional benchmarks for comparative performance evaluation).
Bezugssorten-Rolle
Referenzsorten-Rolle
Standardsorten-Rolle
benchmark cultivar role
reference variety role
standard cultivar role
This P et al. (2004) Development of a standard set of microsatellite reference alleles for identification of grape cultivars, Theoretical and Applied Genetics 109: 1448-1458 OIV (2009) 2nd Edition of the OIV Descriptor List for Grape Varieties and Vitis Species Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157
Reference Cultivar Role
Villard Blanc serves in a Reference_Parent_Role in crosses with resistance donors, its powdery mildew susceptibility providing the baseline phenotype against which resistance segregation is scored—resistant progeny carrying donor alleles, susceptible progeny carrying Villard Blanc alleles at resistance loci. V. vinifera cv. Cabernet Sauvignon bears a Reference_Parent_Role in quality trait mapping, its known fruit composition and wine character serving as reference against which trait variants from crossing partners are detected. PN40024 (highly homozygous Pinot Noir derivative) functions as reference parent and reference genome in grapevine genomics, its sequenced genome providing the coordinate system for variant detection and gene annotation across the genus.
A mapping population role that inheres in a grapevine genotype serving as a reference standard within genetic mapping and QTL analysis, typically the parent carrying the "wild-type" or "susceptible" phenotype against which the alternative parent's trait-conferring alleles are mapped. The reference parent provides the baseline genetic background and phenotype, with the alternative parent's contributions (resistance alleles, quality trait variants) detected as deviations from reference parent expectations. In resistance mapping, the reference parent is typically a susceptible V. vinifera variety, with resistance loci from the donor parent detectable as marker-trait associations segregating in progeny. The reference parent concept extends beyond mapping to broader genetic analysis, where established reference genotypes (PN40024 for genome sequencing, certified clones for variety identification) serve as standards against which other genotypes are compared. Reference parents are characterized in depth—fully genotyped, extensively phenotyped, maintained in certified collections—enabling their function as stable reference points for genetic research.
Reference_Parent_Role captures the standardization function essential for genetic analysis, where one parent provides the baseline enabling detection of the other parent's genetic contributions. The reference concept is asymmetric—while mapping populations have two parents contributing equally to progeny genomes, analytical frameworks often treat one parent as "reference" and detect the other's alleles as variants from reference. Reference parent selection typically favors well-characterized, widely used genotypes whose genetics are extensively documented, enabling integration of new mapping results with prior knowledge. The reference genome (PN40024) has particularly influenced grapevine genetics, providing the physical framework for genetic map integration, sequence-based marker development, and candidate gene identification underlying contemporary molecular breeding.
Referenzelter
Referenzgenotyp
Standardelter
empfindlicher Elter
reference genotype
reference parent
standard parent
susceptible parent
wild-type parent
Jaillon O et al. (2007) The grapevine genome sequence, Nature 449: 463-467 Adam-Blondon AF et al. (2011) Genetic mapping in grapes, CRC Press pp.: 119-159 Canaguier A et al. (2017) A new version of the grapevine reference genome, GigaScience 6: 1-12 Grimplet J et al. (2012) Comparative analysis of grapevine genomes, Frontiers in Plant Science 3: 125
Reference Parent Role
Refilling capacity assessment in grapevine petioles using real-time X-ray microCT imaging reveals the dynamics of embolism reversal, with moderately embolized petioles (40% air-filled vessels) showing progressive refilling over 12 hours following rewatering, with small vessels (<30 μm diameter) refilling first (likely easier to generate sufficient pressure), larger vessels refilling more slowly, and approximately 80% of initially embolized vessels refilling within 24 hours under well-watered conditions, demonstrating substantial refilling capacity that enables rapid hydraulic recovery if soil water is restored.
Refilling capacity quality is an embolism recovery capacity quality that represents the ability to restore water continuity in embolized xylem vessels through active or passive refilling processes, characterizing the potential for rapid hydraulic recovery that does not require new vessel construction and enables vines to regain transport function within hours to days following stress relief. Within the BFO framework, this quality specifically addresses the refilling mechanism of embolism recovery, distinguishing vessel refilling from new xylem production while recognizing that refilling represents the most rapid pathway to hydraulic recovery that could enable day-to-day restoration of conductivity following overnight or post-irrigation water status improvement. The ontological significance lies in refilling capacity determining the short-term resilience of the hydraulic system, where effective refilling enables vines to recover from moderate drought episodes without permanent damage, treating embolism as a reversible protective response rather than an irreversible injury, while limited refilling capacity means that even moderate embolism represents lasting damage requiring structural repair. The mechanistic basis for refilling involves overcoming the surface tension barrier that prevents spontaneous water entry into air-filled vessels, with proposed mechanisms including root pressure (positive pressure from root osmotic water uptake pushing water into vessels), stem positive pressure (locally generated pressure driving refilling), osmotic refilling (solute secretion into embolized vessels creating water potential gradients for refilling), and tissue pressure refilling (surrounding living cells generating pressure on vessels), with evidence suggesting that different mechanisms may operate in different organs and conditions. The conditions affecting refilling capacity include water availability (adequate soil water required for any refilling mechanism), transpiration rate (refilling may require reduced transpiration to allow pressure buildup), carbohydrate availability (osmotic mechanisms requiring solute secretion), and vessel characteristics (smaller vessels potentially easier to refill than large ones). The controversy surrounding refilling acknowledges that while some studies demonstrate rapid recovery consistent with refilling, others question whether the artifacts of measurement methods may overestimate recovery, with the extent and mechanisms of refilling in grapevine remaining active research areas. Understanding refilling capacity quality enables assessment of short-term recovery potential, interpretation of day-to-day hydraulic dynamics, and evaluation of the reversibility of drought-induced hydraulic damage.
Refilling_Capacity_Quality addresses the rapid recovery mechanism through vessel refilling without new xylem construction. This quality determines short-term hydraulic resilience and day-to-day recovery potential. The class supports recovery dynamics interpretation and drought resilience assessment through refilling capacity characterization.
Embolieumkehr
Gefäßwiederbefüllung
Wiederbefüllungskapazität
rapid recovery
refilling potential
schnelle Erholung
vessel refilling
Brodersen CR et al. (2010) The dynamics of embolism repair in xylem. Plant Physiology 154(3): 1088-1095 Knipfer T et al. (2016) In situ visualization of the dynamics in xylem embolism formation and removal in the absence of root pressure. Plant Physiology 171(2): 1024-1036
Refilling Capacity Quality
Reflectance increase quality assessment in warm-climate Vitis vinifera cultivars evaluated the contribution of surface optical properties to heat avoidance through spectrophotometric analysis of leaf and berry surfaces. Leaf reflectance measurements across the solar spectrum (300-2500 nm) revealed substantial variation in reflectance increase quality among cultivars. Cinsaut demonstrated excellent reflectance increase quality with total solar reflectance of 38% on mature leaves, near-infrared (700-1100 nm) reflectance of 52%, and visible light reflectance of 28%, with the high reflectance attributed to dense epicuticular wax deposits creating a glaucous leaf surface. Scanning electron microscopy confirmed abundant platelet-form wax crystals creating a highly textured surface conducive to light scattering. Merlot showed moderate reflectance increase quality with total solar reflectance of 31%, NIR reflectance of 44%, and visible reflectance of 22%, with moderate wax density. Pinot Noir exhibited poor reflectance increase quality with total reflectance of only 24%, NIR of 35%, and visible of 18%, with smooth, glossy leaf surfaces showing minimal wax crystal development. Thermal modeling estimated that the reflectance increase quality differential between Cinsaut and Pinot Noir would result in 1.8°C lower leaf temperature under 1000 W m⁻² irradiance, providing supplemental heat avoidance benefit that combines additively with transpirational cooling. The reflectance increase quality assessment identified surface reflectance as a heritable trait contributing to overall heat avoidance capacity and representing a potential selection criterion for warm-climate variety development.
Reflectance increase quality is a heat avoidance quality that inheres in a grapevine organism and represents the capacity to increase the reflection of incident solar radiation from leaf and fruit surfaces through modification of surface optical properties, reducing the absorption of solar energy that would otherwise contribute to tissue temperature increase, with reflectance enhancement achieved through accumulation of epicuticular waxes, trichome density increases, or other structural modifications that alter surface reflectivity. This quality captures a passive heat avoidance mechanism that reduces heat absorption without requiring water expenditure, providing sustained protection throughout periods of high radiation that complements the active cooling mechanisms of transpiration and the radiation-avoiding movements of leaf angle adjustment. The ontological significance of reflectance increase quality lies in its potential for constitutive or induced protection that operates continuously without metabolic cost once surface modifications are in place, providing background heat avoidance benefit that may be particularly valuable when water limitations constrain transpirational cooling and when radiation conditions overwhelm the capacity of other avoidance mechanisms. The biological basis of reflectance increase encompasses the deposition of epicuticular wax crystals that create microstructured surfaces scattering incident radiation, the development of leaf trichomes that create a pubescent surface layer reflecting light before it reaches the mesophyll, and potentially the modification of cuticular composition affecting optical properties across the solar spectrum. The magnitude of reflectance increase achievable varies among genotypes reflecting genetic differences in wax biosynthesis and trichome development, and may be induced by prior stress exposure that stimulates protective surface modification through enhanced wax deposition. The spectral characteristics of reflectance are important, with reflection of near-infrared radiation providing the greatest thermal benefit since this wavelength range carries substantial heat energy but does not contribute to photosynthesis, while excessive visible light reflection would reduce photosynthetic capacity along with thermal load. The developmental regulation of surface reflectance creates characteristic patterns with newly expanded leaves often showing lower reflectance than mature leaves that have accumulated wax deposits, and with fruit surface reflectance potentially changing through development as berry composition and surface characteristics evolve.
Reflectance_Increase_Quality is_a Heat_Avoidance_Quality, representing the passive mechanism of increased radiation reflection from plant surfaces. This class addresses heat avoidance through optical properties rather than physiological cooling. Total solar reflectance percentage and NIR-specific reflectance provide quantification. Epicuticular wax and trichome density are the primary structural determinants. The mechanism is water-independent, valuable when transpirational cooling is limited. Temperature reductions of 1-3°C achievable through reflectance alone. Distinguished from Transpirational_Cooling_Capacity_Quality and Leaf_Angle_Adjustment_Quality as a constitutive rather than active/inducible mechanism.
Oberflächenreflektanz
Reflektanzerhöhung
Strahlungsreflexion
epicuticular wax reflectance
epikutikuläre Wachsreflektanz
pubescence reflectance
radiation reflection
surface reflectance
Wahid A et al. (2007) Heat tolerance in plants: an overview. Environmental and Experimental Botany 61:199-223 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 453-507 Holmes MG, Keiller DR (2002) Effects of pubescence and waxes on the reflectance of leaves in the ultraviolet and photosynthetic wavebands: a comparison of a range of species. Plant, Cell and Environment 25:85-93
Reflectance Increase Quality
Thompson Seedless exhibits extreme reflex stamen functionally female flowers at BBCH 65 with filaments curved sharply downward positioning anthers 1.5-2.5 mm below stigma level, producing moderate pollen quantities (8,000-12,000 grains per flower) with poor viability (12-18%), requiring co-planted hermaphrodite pollinizers (Calmeria, Perlette) at 1=8 to 1=10 ratios to achieve 25-35% fruit set through wind-borne cross-pollination, with pollinizer proximity >15 m dramatically reducing fruit set to <15%. Gewürztraminer (functionally female clones) in Alsace exhibits pronounced reflex stamens with anthers 1-1.5 mm below stigma, producing reduced pollen (5,000-9,000 grains per flower) with low viability (15-25%), depending on cross-pollination from adjacent hermaphrodite cultivars (Riesling, Pinot Blanc) to achieve 28-38% fruit set. Flame Seedless shows constitutive strong stamen reflexing (anthers 1.8-2.2 mm below stigma) with minimal pollen production (<4,000 grains per flower), requiring intensive pollinizer management for commercial production.
A reflex stamen functionally female flower is a functionally female flower in which male sterility is achieved primarily through pronounced downward curvature (reflexing) of stamen filaments that positions anthers substantially below the stigma, creating spatial separation that effectively prevents self-pollination even when pollen production and viability may be adequate, representing the most common and morphologically distinctive mechanism of functional female expression in Vitis vinifera. Reflex stamen functionally female flowers are characterized by= (1) strongly curved filaments that bend sharply downward from their point of attachment on the receptacle, creating a reflexed arc with anthers positioned 0.8-2.5 mm below the stigma rather than at or above stigma level as in hermaphrodite flowers (2) pollen production that may range from severely reduced (<5,000 grains per flower) to moderate (8,000-15,000 grains per flower) depending on cultivar and conditions, but regardless of pollen quantity, the spatial positioning prevents self-pollination (3) pollen viability that may be good (50-70%) in some cultivars or reduced (<20-30%) in others, but again the physical separation from stigma prevents effective self-fertilization (4) fully functional pistil with normal morphology, receptive stigma, and viable ovules. The reflexed stamen position is the primary and sufficient mechanism of male sterility in this flower type, as the downward-curved filaments position anthers in a location from which shed pollen cannot reach the stigma through gravity, requiring wind or insect transport to move pollen upward from the reflex anther position to the stigma, which is highly inefficient within individual flowers and effectively prevents self-pollination. The degree of stamen reflexing varies among cultivars and can show some phenotypic plasticity, with cool temperatures during flower development in some cultivars inducing more pronounced reflexing while warm temperatures may reduce reflexing intensity. The reflex stamen condition is controlled genetically, with evidence suggesting major-gene control showing Mendelian segregation in breeding populations, though environmental factors can modulate expression intensity. Reflex stamen functionally female flowers achieve fruit set only through cross-pollination from hermaphrodite pollen sources, with pollen from co-planted pollinizer cultivars being transported by wind (primary mechanism) or insects (secondary mechanism) to land on the receptive stigma. Fruit set percentages in reflex stamen functionally female flowers typically range from 20-40% under good pollination conditions (adequate pollinizer proximity, favorable weather for pollen dispersal) but can fall to <10-15% under poor pollination conditions (distant or insufficient pollinizers, wet or humid weather during bloom preventing pollen dispersal). The reflex stamen mechanism is agronomically significant as it creates absolute dependency on cross-pollination, requiring careful vineyard design with pollinizer rows, appropriate spatial arrangements for pollen movement, and bloom synchrony management.
Reflex_Stamen_Functionally_Female_Flower is distinguished from Reduced_Pollen_Functionally_Female_Flower by the primary mechanism of male sterility= physical stamen positioning versus gametic (pollen) dysfunction, though these mechanisms often co-occur with reflex stamens frequently also showing reduced pollen quantity or viability. The reflex stamen condition is the more morphologically obvious mechanism and serves as the primary diagnostic feature for identifying functionally female cultivars in the field. The genetic basis involves specific loci controlling stamen filament development and auxin responses affecting filament curvature. Understanding the developmental regulation of stamen reflexing could inform breeding strategies for introducing or eliminating functional female expression as desired for specific production goals. The reflex stamen mechanism represents one evolutionary pathway back toward dioecy from hermaphroditism in cultivated Vitis, creating obligate outcrossing through spatial rather than temporal reproductive isolation.
Blüte mit zurückgebogenen Staubblättern
downward-curved stamen flower
reflex-stamen female flower
reflexstaubblättrige weibliche Blüte
umgebogene Staubblätter
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 99-101 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Antcliff, A.J. (1980). Inheritance of sex in Vitis. Annales de l'Amélioration des Plantes 30=113-122 Caporali, E. et al. (2003). The arrest of development of abortive reproductive organs in the unisexual flower of Vitis vinifera. Sexual Plant Reproduction 15=291-300.
Reflex Stamen Functionally Female Flower
Refoliation capacity quality assessment in Franciacorta Chardonnay compared recovery trajectories following early versus late basal leaf removal removing equivalent 35% of total leaf area. Early removal at BBCH 71 (berry set) stimulated strong refoliation capacity quality expression with lateral shoot emergence from 62% of nodes in the defoliated zone, restoring 80% of removed leaf area within six weeks and largely negating intended fruit exposure benefits as fruit zone leaf layer number returned from 0.8 immediately post-removal to 1.9 by véraison. Late removal at BBCH 81 (beginning of ripening) exhibited minimal refoliation capacity quality, with only 15% of defoliated nodes producing new growth that contributed less than 20% replacement of removed leaf area, maintaining fruit exposure benefits through harvest with stable fruit zone leaf layer number of 0.9. The results established post-véraison timing as critical for achieving persistent defoliation effects in this vigorous cultivar-site combination where early removal triggers compensatory refoliation that undermines management objectives.
Refoliation capacity quality is a leaf removal response quality that inheres in a grapevine organism and represents the ability to regenerate functional leaf area following defoliation events through activation of latent buds, expansion of existing lateral buds, or accelerated development of emerging leaves on actively growing shoots, providing a mechanism for recovery from severe leaf loss that exceeds the compensation capacity of remaining foliage alone. This quality determines the potential for canopy reconstruction following damaging events and influences decisions regarding the timing and intensity of deliberate defoliation practices in contexts where refoliation may negate intended exposure benefits or contribute positively to late-season photosynthetic capacity. The ontological basis of refoliation capacity quality rests on the developmental plasticity of grapevines that maintain populations of dormant or slowly developing buds capable of rapid activation when released from correlative inhibition by removal of existing leaves and shoot tips, combined with the indeterminate growth habit that permits continued leaf production throughout the growing season under favorable conditions. The physiological triggers for refoliation include altered cytokinin-to-auxin ratios following leaf removal that release lateral bud dormancy, increased light penetration to interior bud positions that stimulates photomorphogenic activation, and improved water and nutrient availability per remaining growing point that supports rapid new growth. The magnitude and timing of refoliation capacity expression vary with defoliation timing, severity, and pattern, with early-season defoliation during active growth typically provoking more vigorous refoliation than late-season events when growth potential has diminished naturally. Cultivar differences in refoliation capacity quality reflect variation in lateral bud number and development stage, the strength of apical dominance and correlative inhibition, and overall vigor characteristics that determine resource availability for new growth initiation. The practical implications of refoliation capacity quality for vineyard management include the potential for canopy recovery following hail damage or severe pest defoliation, but also the risk that leaf removal intended to improve fruit exposure may be partially negated by rapid refoliation that restores shading if intervention occurs during periods of high regrowth potential.
Refoliation_Capacity_Quality is_a Leaf_Removal_Response_Quality, representing the ability to regenerate leaf area following defoliation. This class addresses regrowth mechanisms distinct from Defoliation_Tolerance_Quality which concerns function maintenance with existing reduced leaf area. The timing dependency is critical, with early-season defoliation typically triggering stronger refoliation than late-season removal. This quality can be either beneficial (recovery from damage) or problematic (negating intentional defoliation) depending on management context. Related to lateral bud development and vine vigor characteristics.
Blattregenerationsfähigkeit
Neubelaubungskapazität
Refoliationspotential
Wiederbelaubungskapazität
canopy recovery capacity
leaf regeneration ability
new leaf development capacity
refoliation potential
Poni S et al. (2006) Effects of early defoliation on shoot photosynthesis, yield components, and grape composition. American Journal of Enology and Viticulture 57:397-407 Sabbatini P, Howell GS (2010) Effects of early defoliation on yield, fruit composition, and harvest season cluster rot complex of grapevines. HortScience 45:1804-1808 Palliotti A et al. (2013) Changes in vineyard establishment and canopy management urged by earlier climate-related grape ripening: a review. Scientia Horticulturae 163:43-54
Refoliation Capacity Quality
Mourvèdre vines at Tablas Creek Vineyard in Paso Robles bearing a regenerative production role are managed with year-round diverse cover crops of 8-12 species including legumes, brassicas, and grasses, integrated sheep grazing at 2-4 animal units/ha during dormancy, no-till soil management maintaining soil organic carbon increases of 0.1-0.3% annually, compost applications of 5-8 t/ha from on-site composting, and documented improvements in water infiltration rates from 2.5 to 7.5 cm/hour over a 10-year regenerative transition.
A production system role that inheres in a grapevine cultivated under a regenerative agriculture framework that prioritizes active restoration and enhancement of soil health, ecosystem function, and biodiversity beyond baseline sustainability, employing practices such as no-till management, diverse cover cropping, composting, and holistic planned grazing to rebuild soil organic matter and ecological complexity.
Regenerative_Production_Role represents an emerging viticultural paradigm that extends beyond the harm-reduction focus of sustainable production toward active ecosystem restoration. Unlike organic or biodynamic roles defined by input restrictions, the regenerative role is defined by measurable outcome targets — particularly soil organic matter accumulation, water cycle improvement, and biodiversity increase: certification frameworks such as the Regenerative Organic Certification (ROC) launched in 2020 are beginning to formalize these outcome-based standards. The role may overlap with organic or biodynamic certification but is distinguished by its explicit emphasis on soil carbon sequestration and ecosystem regeneration.
aufbauende Bewirtschaftungsrolle
regenerative Anbau-Rolle
regenerative Produktionsrolle
regenerative farming role
regenerative viticulture role
restorative production role
Regenerative Organic Alliance (2020) Regenerative Organic Certified Framework LaCanne CE and Lundgren JG (2018) Regenerative agriculture: merging farming and natural resource conservation, PeerJ 6: e4428 Schreefel L et al. (2020) Regenerative agriculture, One Earth 3: 507-524
Regenerative Production Role
Regional acreage for Grüner Veltliner across Austrian wine-growing regions (Statistik Austria, Weinbauerhebung 2023): Weinviertel approximately 6,200 ha (approximately 46% of total Austrian Grüner Veltliner, and approximately 49% of the Weinviertel's total vineyard area), Kamptal approximately 1,500 ha, Kremstal approximately 1,000 ha, Wachau approximately 670 ha (approximately 50% of the Wachau's vineyard area), Wagram approximately 900 ha, Thermenregion approximately 600 ha, with smaller plantings in all other Austrian regions — the distribution revealing Grüner Veltliner's absolute dominance in the Weinviertel and its approximately 50% share in the Wachau. Regional acreage for Blaufränkisch across Austrian wine-growing regions documents concentration in Burgenland — Mittelburgenland approximately 1,200 ha (approximately 65% of Mittelburgenland's total area), Neusiedlersee approximately 800 ha, Leithaberg approximately 450 ha, Eisenberg approximately 300 ha, with modest plantings in Carnuntum and the Thermenregion — confirming Burgenland's role as the center of Austrian Blaufränkisch cultivation.
Regional acreage is a production statistics entity that documents the vineyard area planted to a specific grape cultivar within a defined sub-global geographic unit — country, administrative region, wine-growing district, or appellation zone — expressed in hectares (ha) with the geographic scope, data year, source authority, and proportion of the region's total vineyard area represented by the cultivar, providing the geographically disaggregated acreage information that reveals the spatial distribution patterns, regional concentration, and local importance of cultivars within individual viticultural landscapes.
Regional acreage data provides the geographically resolved information essential for understanding cultivar distribution patterns, regional specialization, and local viticultural identity — information that global acreage aggregates necessarily obscure. The analytical value of regional acreage data lies in its capacity to reveal several critical distribution characteristics — geographic concentration (the degree to which a cultivar's acreage is concentrated in a small number of regions versus dispersed across many), regional dominance (the proportion of a region's total vineyard area occupied by a specific cultivar, indicating the cultivar's importance to regional identity and economy), complementary distribution (cultivars that occupy distinct regional niches without significant overlap), and competitive dynamics (cultivars expanding or contracting within the same regions in response to market forces or regulatory incentives). The data sources for regional acreage statistics mirror the national vineyard register infrastructure — Austrian data derived from the Weinbaukataster maintained by Statistik Austria, French data from the CVI (Casier Viticole Informatisé) maintained by regional customs authorities, Italian data from the SIAN (Sistema Informativo Agricolo Nazionale), and Australian data from the Wine Australia Vineyard Register. The temporal dimension of regional acreage data reveals the dynamics of cultivar portfolio evolution within regions — the progressive displacement of Welschriesling by Grüner Veltliner in the Weinviertel, the expansion of Blaufränkisch at the expense of Zweigelt in premium Burgenland sites, the replacement of Müller-Thurgau by Riesling and Pinot Noir in German quality regions — each trajectory reflecting the interaction of market demand, quality ambition, climate adaptation, and regulatory incentive.
Bezirksanbaufläche
district vineyard area
gebietsbezogene Anbauausdehnung
local cultivation area
national acreage
nationale Rebfläche
regional planted area
regional planting extent
regionale Anbaufläche
regionale Bestockung
Statistik Austria (2023) Weinbauerhebung, Bundesanstalt Statistik Österreich Anderson K, Nelgen S (2020) Which Winegrape Varieties are Grown Where? University of Adelaide Press Österreichischer Weinbauverband (2023) Rebflächenstatistik Eurostat (2023) Vineyard statistics — area by grape variety
Regional Acreage
A Pinot Noir clone designated "115" bears a Regional_Clone_Role identifying it as a French ENTAV-INRA selection, the numeric code placing it within the French registration system regardless of where it is subsequently planted. A Riesling clone designated "239 Gm" serves in a Regional_Clone_Role indicating German Geisenheim origin, the "Gm" suffix distinguishing it from Freiburg (Fr) or Weinsberg (We) selections. A Sangiovese clone designated "R24" bears a Regional_Clone_Role within the Italian VCR Rauscedo system, the "R" prefix indicating Rauscedo origin within the Italian clone nomenclature.
A clone role that inheres in a grapevine clone by virtue of its origin, registration, or institutional affiliation within a specific national or regional clone development and certification system, reflecting the geographic organization of viticultural research and regulation. Regional clone classification recognizes that clone selection programs operate within national or regional frameworks—each with distinct institutional structures, evaluation methodologies, registration procedures, and nomenclature conventions—creating geographically organized clone inventories. Major regional clone systems include France (ENTAV-INRA, numeric codes), Italy (VCR Rauscedo, regional research stations), Germany (Geisenheim, Freiburg, Weinsberg institutes with Gm/Fr/We prefixes), Austria (HBLA Klosterneuburg), Switzerland (Agroscope Changins), Australia (CSIRO, state agencies), and the USA (UC Davis FPS). Regional designation indicates origin and institutional affiliation but does not necessarily restrict geographic deployment French ENTAV clones are planted globally, German Geisenheim selections have spread throughout cool-climate viticulture, and Italian Rauscedo clones dominate international rootstock supply.
Regional_Clone_Role is subclassed by major national clone development systems—French, Italian, German, and others—reflecting the institutional geography of viticultural research. Regional designation enables clone identification and traceability but creates nomenclature complexity the same variety may have dozens of clones registered across multiple national systems, with no international standardization of naming conventions. Clone exchange between regions occurs through official channels (material transfer agreements, phytosanitary certification) and historically through informal movements that distributed important selections before regulatory frameworks developed. The regional clone landscape reflects national investment in viticultural research, with countries having major wine industries (France, Italy, Germany) maintaining extensive clone evaluation and registration programs while smaller producers may rely on imported registered clones or unregistered local selections.
Herkunftsklon
institutional clone
institutioneller Klon
national clone
nationaler Klon
origin-designated clone
regional clone
regionaler Klon
Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 ENTAV-INRA (2007) Catalogue des Variétés et Clones de Vigne Anderson K, Aryal NR (2013) Which Winegrape Varieties are Grown Where?, University of Adelaide Press Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane
Regional Clone Role
Hárslevelű accession PTE-VIN-087 maintained in the University of Pécs Mecsek Regional Vine Collection bearing a regional collection role is one of approximately 180 accessions representing the traditional cultivar diversity of the Pécs-Mecsek and Villány-Siklós wine districts of southern Transdanubia, including 25 locally selected clonal variants of Kadarka, Olaszrizling, and Kékfrankos not represented in the national collection at Tápiószele, documented with GPS-referenced provenance data from original collection sites in abandoned vineyard parcels, and serving as the source material for regional revitalization programs reintroducing heritage cultivars to commercial production in the Pécs appellation.
A germplasm repository role that inheres in a grapevine maintained within a germplasm collection focused on conserving the cultivar diversity of a specific viticultural region, wine-producing district, or sub-national geographic area, typically emphasizing locally adapted cultivars, regional landraces, and traditional clonal variants of significance to the region's viticultural heritage.
Regional_Collection_Role captures the sub-national scale of germplasm conservation that addresses genetic diversity often overlooked by national and international collections. Regional collections excel at conserving locally adapted clonal variants, informal farmer-selected types, and rare cultivars of limited geographic distribution that may not meet the threshold for national collection inclusion: these locally adapted genotypes represent an irreplaceable reservoir of adaptive genetic variation for climate resilience and terroir-specific quality. Regional collections are typically maintained by local universities, regional agricultural institutes, or wine-grower associations with strong connections to the local viticultural community.
Gebietssammlung-Rolle
district vine collection role
local germplasm collection role
regional genebank role
regionale Genressourcen-Rolle
regionale Sammlungs-Rolle
ECPGR (2019) European Cooperative Programme for Plant Genetic Resources — Vitis Working Group Report Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157 Lacombe T et al. (2018) Grapevine European catalogue: towards a comprehensive list, Vitis 57: 69-76
Regional Collection Role
The cultivar with VIVC prime name Schiava Grossa carries regional names including Trollinger in Württemberg (where it constitutes 20% of vineyard area and is culturally inseparable from Swabian identity), Vernatsch in South Tyrol (the dominant local designation in German-speaking Alto Adige), Black Hamburg in English-speaking countries (referencing the Hamburg glasshouse tradition), and Frankenthal in parts of the Rheinpfalz — each regional name embedding the cultivar within a specific geographic context and carrying local cultural associations absent from the international prime name.
A cultivar designation that functions as a geographically specific identifier for a grapevine cultivar within a defined viticultural region, reflecting local naming traditions, dialect forms, or marketing strategies that create a place-specific designation distinct from the cultivar's national or international nomenclature.
Regional_Name differs from Regional_Synonym in emphasis and function. While Regional_Synonym is a nomenclatural cross-reference class documenting alternative names for database purposes, Regional_Name foregrounds the geographic and cultural embedding of the name within local viticultural identity. Regional names often carry connotations of terroir, tradition, and local pride that the standardized prime name does not convey: the name Trollinger in Württemberg evokes centuries of local viticultural tradition in a way that the prime name Schiava Grossa cannot. Subclasses distinguish between local dialect forms, established traditional regional names, and marketing-oriented regional designations created for commercial purposes.
Regionalname
gebietsspezifischer Sortenname
geographic cultivar name
place-specific variety name
regional cultivar designation
regionale Rebbezeichnung
Robinson J et al. (2012) Wine Grapes, Allen Lane Rézeau P (2014) Dictionnaire des noms de cépages de France, CNRS Éditions Anderson K and Aryal NR (2013) Which Winegrape Varieties are Grown Where?, University of Adelaide Press
Regional Name
A Nebbiolo vine in the Langhe hills of Piedmont fulfills a regional production role that positions it within Italy's DOCG hierarchy, with the specific vineyard location determining whether fruit qualifies for Barolo DOCG (commune-level), Barolo MGA cru designation (vineyard-level), or Langhe Nebbiolo DOC (regional level), with corresponding price differentials of 3-10x between regional and top cru designations reflecting the vine's geographic and cultural identity.
A grapevine role that inheres in a vine by virtue of its geographic origin and cultural-historical significance within a defined viticultural region, capturing the regulatory classification, traditional status, and provenance identity that determine how the vine's fruit is designated, marketed, and valued within regional appellation and quality hierarchies.
Regional_Production_Role classifies grapevines according to their place-based identity and regulatory provenance rather than their agronomic management or genetic characteristics. This role bridges the intersection of terroir, tradition, and regulation: it captures the fact that genetically identical vines planted 500 meters apart may bear fundamentally different regional production roles due to appellation boundaries. Subclasses address appellation-specific hierarchical classifications and the distinction between traditional (indigenous/autochthonous) and introduced cultivars within a region.
Herkunfts-Rolle
gebietsbezogene Anbau-Rolle
geographic production role
provenance-based role
regional designation role
regionale Produktionsrolle
OIV (2022) International Standard for the Labelling of Wines EU Regulation 1308/2013 on PDO/PGI wine designations Unwin T (2012) Wine and the Vine: An Historical Geography of Viticulture and the Wine Trade, Routledge pp: 232-278
Regional Production Role
The cultivar with VIVC prime name Savagnin Blanc carries regional synonyms including Traminer in South Tyrol and the Austrian Weinviertel (from the town of Tramin/Termeno), Heida in the Valais of Switzerland, Païen in the francophone Valais, Formentin in the Somló region of Hungary, Prinç in the Thracian region of Turkey, and Gringet in the Ayze appellation of Haute-Savoie — each name arising independently within its respective region through centuries of isolated local usage before molecular identification revealed them as genetically identical.
A synonym that arose from the independent naming of a grapevine cultivar within a specific geographic region or viticultural district, reflecting local naming conventions, dialect influences, or regional traditions that produced a designation distinct from the cultivar's prime name or its names in other regions.
Regional_Synonym represents the geographic fragmentation of cultivar nomenclature that is characteristic of grapevine naming across Europe and beyond. Before the era of molecular identification, cultivars that spread across regions through cuttings and trade frequently acquired new local names at each destination, creating parallel naming traditions that persisted for centuries without recognition of their synonymy: the classic example is Tribidrag/Crljenak Kaštelanski/Primitivo/Zinfandel, where the same genotype bore four unrelated names in Croatia, southern Italy, and California for over a century before SSR analysis established their identity in 2001. Regional synonyms remain culturally significant and are often protected in appellation regulations.
Lokalname
geographic synonym
local synonym
regional variety name
regionales Synonym
Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157 Robinson J et al. (2012) Wine Grapes, Allen Lane pp: 1-18 Maleševic M et al. (2001) Microsatellite fingerprinting of Croatian grapevine varieties, Vitis 40: 183-189
Regional Synonym
The four regional wine designation systems represented in VVO subclasses illustrate the diversity of national implementation approaches — French Vin de Pays (now formally equivalent to PGI), German Landwein (PGI-equivalent regional wine category), Italian IGT (Indicazione Geografica Tipica, PGI-equivalent), and American AVA (American Viticultural Area, a geographically delimited designation without production rule specifications) — each carrying distinct national regulatory traditions, cultural associations, and commercial positioning strategies that the harmonized EU PGI terminology cannot fully capture.
A geographic indication that identifies a wine as originating from a defined wine-producing region under a regulatory framework that predates, parallels, or implements the EU PDO/PGI system at the national level, using country-specific terminology and regulatory structures that may carry additional cultural, historical, or commercial significance beyond their formal legal equivalence to the harmonized EU categories.
Regional_Wine_Designation captures the national-level terminology and regulatory frameworks that constitute the practical reality of geographic indication systems as experienced by producers and consumers. While the EU harmonization project (culminating in Regulation 1308/2013) established PDO and PGI as the overarching legal categories, the national designation terms (AOC, DOC, DOCG, DO, AVA, GI) remain in commercial and cultural use and carry distinct connotations in their respective markets: an Italian consumer understands IGT as implying creative freedom within a geographic framework, a German consumer reads Landwein as a step between Deutscher Wein and Qualitätswein, and an American consumer interprets AVA as a geographic reference without quality hierarchy implications. Subclasses represent the principal national designation systems encountered in the VVO domain.
country-specific wine designation
länderspezifische Herkunftsangabe
national origin designation
nationale Weinbezeichnung
regional wine category
regionale Weinkategorie
EU Regulation 1308/2013 Art. 93-108 OIV (2015) OIV Guidelines for Geographic Indications for Wines Anderson K and Aryal NR (2013) Which Winegrape Varieties are Grown Where?, University of Adelaide Press
Regional Wine Designation
A Gamay vine in the southern Mâconnais bearing a regional wine role produces fruit under the Bourgogne AOC at maximum yields of 58 hl/ha (versus 40 hl/ha for village Volnay), with the broad regional appellation encompassing the entire Burgundy production zone from Chablis to Mâcon, resulting in wines typically priced at 8-15 €/bottle that represent approximately 50% of total Burgundy production volume at 1.5 million hectoliters annually, serving as the accessible entry point to the regional hierarchy.
An appellation-specific role that inheres in a grapevine situated within a broad regional appellation encompassing a large geographic area, producing fruit that qualifies for the regional designation as the entry-level tier of the appellation hierarchy with the most permissive yield limits, broadest cultivar authorization, and least restrictive viticultural requirements.
Regional_Wine_Role occupies the base tier of hierarchical appellation systems worldwide, including Bourgogne AOC, Bordeaux AOC, Languedoc AOC in France, Toscana IGT in Italy, Landwein and VDP.Gutswein in Germany, and varietal-labeled wines under broad GI designations in the New World. The role's defining characteristic is geographic breadth: regional appellations may span hundreds of kilometers and thousands of hectares, encompassing diverse terroirs, microclimates, and soil types under a single designation. This contrasts with the geographic precision of village, Premier Cru, and Grand Cru roles where the vine's specific parcel location is the defining attribute.
Gebietswein-Rolle
Regionalwein-Rolle
broad-area designation role
entry-level appellation role
regional appellation role
überregionale Appellations-Rolle
INAO (2023) Cahier des charges AOC Bourgogne EU Regulation 1308/2013 Art. 93 on PDO/PGI definitions Johnson H and Robinson J (2019) The World Atlas of Wine 8th Ed, Mitchell Beazley pp: 50-53
Regional Wine Role
Chardonnay clone 76 bears a Registered_Clone_Role in the French ENTAV-INRA catalogue, registration achieved after 15 years of evaluation trials in Burgundy, Champagne, and Languedoc confirming its distinctive small-cluster character and concentrated wine quality—now among the most planted Chardonnay clones globally. Pinot Noir clone 777 serves in a Registered_Clone_Role with full French registration, the evaluation process documenting its early ripening, aromatic intensity, and suitability for cool-climate production. Sangiovese clone R24 bears a Registered_Clone_Role in the Italian national registry (Registro Nazionale delle Varietà di Vite), the VCR selection registered after extensive Tuscan trials confirming superior wine quality for Brunello di Montalcino production.
An official clone role that inheres in a grapevine clone that has achieved full registration status within a national or regional clone registry, representing the highest level of official recognition and typically requiring completion of extensive evaluation trials demonstrating agronomic and enological merit. Registered clones have undergone multi-year (typically 10-20 year) evaluation programs including replicated field trials assessing yield, vigor, phenology, and disease resistance winemaking trials evaluating wine quality, typicity, and aging potential and sanitary testing confirming pathogen-free status at the required certification level. Registration confers the right to propagate and sell the clone under its official designation, with mandatory labeling requirements ensuring traceability from nursery production through vineyard establishment. Registered clone status represents substantial institutional investment in selection and evaluation, with registration programs operated by government agencies (ENTAV-INRA in France, JKI in Germany), research institutions (UC Davis FPS, CSIRO), or authorized private organizations. The registered clone designation ensures growers receive genetically uniform, characterized material with documented performance attributes.
Registered_Clone_Role represents the full official recognition tier where clones have completed comprehensive evaluation justifying their inclusion in national catalogues and registries. The registration process varies by country but typically involves multi-site replicated trials (5-10 years minimum), standardized winemaking evaluations, and confirmation of sanitary status through testing protocols. Registration creates intellectual property implications—some jurisdictions provide plant variety protection for registered clones, enabling selection programs to control propagation and collect royalties. The registered clone inventory for major varieties has expanded substantially since systematic selection programs began in the 1960s-1970s Pinot Noir alone has 50+ French registered clones plus additional registrations in Germany, Italy, Switzerland, and other countries. Growers increasingly specify registered clones when ordering vines, recognizing the documented performance differences and quality assurance represented by registration status.
Katalogklon
approved variety clone
catalog clone
fully registered clone
vollständig zugelassener Klon
ENTAV-INRA (2007) Catalogue des Variétés et Clones de Vigne Cultivés en France European Union (2020) Commission Implementing Directive 2020/177 Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 Regner F (2007) Genetic variation in grapevine clones, in Bentley R (ed) The Handbook of Grape Processing, Blackwell pp.: 1-34
Registered Clone Role
Regulatory information for Grüner Veltliner in Austria integrates multiple regulatory dimensions: varietal authorization (classified as "Qualitätsrebsorte" under the Austrian Weingesetz 2009, authorized for quality wine production in all Austrian wine-growing regions), appellation compliance (eligible as the principal or exclusive variety in multiple DAC regulations — Weinviertel DAC requiring 100% Grüner Veltliner, Wachau DAC permitting Grüner Veltliner as a varietal wine, Kamptal and Kremstal DAC similarly), planting rights (subject to the Austrian Pflanzrechtesystem derived from the EU vine planting authorization regime post-2016), and labeling regulations (varietal labeling requiring minimum 85% Grüner Veltliner under EU Regulation 2019/33, Austrian Weingesetz mandating quality tier designation — Qualitätswein, Kabinett, or Prädikatswein). Regulatory information for Cabernet Sauvignon in France documents a complex regional authorization mosaic — authorized as a principal variety in Bordeaux appellations (Médoc, Graves, Saint-Émilion, Pomerol AOC regulations), as a supplementary variety in certain Languedoc-Roussillon appellations, and excluded from Burgundy and Alsace appellations where only specified traditional cultivars are permitted.
Regulatory information is a grapevine information entity that documents the legal, administrative, and institutional frameworks governing the cultivation of grape cultivars, the production of wine, and the labeling and marketing of viticultural products, encompassing varietal authorization systems (which cultivars may be planted where), appellation and geographical indication regulations (origin-based quality classification), planting rights regimes (permits and restrictions on vineyard establishment), labeling requirements (varietal, origin, and quality declarations on wine labels), and phytosanitary regulations (quarantine, certification, and health status requirements for plant material movement), providing the juridical-administrative dimension that constrains and structures viticultural practice within national and supranational legal frameworks.
Regulatory information constitutes a domain of viticultural knowledge that is simultaneously essential and extraordinarily complex — essential because no commercial grape cultivation or wine production can proceed without compliance with the applicable regulatory frameworks, and complex because these frameworks operate at multiple jurisdictional levels (international, supranational EU, national, regional, and appellation-specific) with frequent amendments, derogations, and interpretive guidance that create a continuously evolving legal landscape. The EU wine regulatory framework (the Common Market Organisation for wine, currently based on Regulation 1308/2013 and its implementing regulations) provides the overarching structure for European viticulture, establishing the categories of vine planting authorization, protected designation of origin (PDO) and protected geographical indication (PGI), oenological practices authorization, and labeling requirements — but delegates substantial regulatory authority to member states and, within member states, to regional and appellation-level bodies. The non-European wine world operates under distinct national frameworks (the US TTB/AVA system, the Australian Wine Australia/GI system, the South African Wine of Origin system, the Chilean/Argentine DO systems) that share some structural features with the EU system but differ substantially in detail, particularly regarding varietal authorization (most New World jurisdictions do not restrict which cultivars may be planted) and production method regulations (generally less prescriptive than European PDO requirements). The regulatory information domain intersects with virtually every other VVO information entity hierarchy — varietal authorization constrains which cultivars may be grown (connecting to genetic and morphological identity), appellation regulations specify permissible viticultural practices (connecting to quality performance data), and labeling rules govern how wine is characterized for consumers (connecting to wine specification and sensory description).
Rechtsrahmen
Weinbauvorschriften
Weinrechtsdaten
legal framework
regulatorische Information
regulatorische Rahmenbedingungen
regulatory compliance information
regulatory data
viticultural regulation
wine law data
European Parliament (2013) Regulation (EU) No 1308/2013 establishing a common organisation of the markets in agricultural products Österreichisches Weingesetz (2009) BGBl. I Nr. 111/2009 European Parliament (2019) Commission Delegated Regulation (EU) 2019/33 supplementing Regulation (EU) No 1308/2013 Robinson J, Harding J (2015) The Oxford Companion to Wine, 4th Ed, Oxford University Press, pp.: 385-392
Regulatory Information
Kinetic analysis of Rehydration_Capacity_Disposition at Stellenbosch University tracked tissue water status recovery at high temporal resolution (hourly measurements for 72 hours post-rewatering) across 15 cultivars following standardized drought stress. Measurements included predawn and midday leaf water potential, leaf relative water content (RWC), stem water content, and root hydraulic conductance. Rehydration_Capacity_Disposition showed marked cultivar variation. 'Chenin Blanc' exhibited rapid rehydration with predawn water potential rising from -0.9 MPa to -0.25 MPa within 8 hours of rewatering, leaf RWC increasing from 72% to 92% by 12 hours, and complete water status restoration by 24 hours. Root hydraulic conductance recovered to 95% of pre-stress values by 24 hours, indicating minimal root damage during stress. 'Pinotage' showed slower Rehydration_Capacity_Disposition with predawn potential reaching only -0.45 MPa at 24 hours, RWC at 84%, and incomplete restoration even at 72 hours (predawn -0.32 MPa versus pre-stress -0.18 MPa). Root conductance in 'Pinotage' remained depressed at 65% of pre-stress, suggesting persistent root damage or dysfunction. Analysis of rehydration kinetics revealed that initial recovery rate (0-8 hours) was primarily determined by root hydraulic conductance (r = 0.78), while asymptotic recovery level depended on xylem embolism status (r = -0.72 with PLC). 'Syrah' showed an interesting pattern of rapid initial rehydration (high root conductance) but incomplete final recovery (significant xylem embolism), demonstrating that Rehydration_Capacity_Disposition has distinct kinetic components.
A Rehydration_Capacity_Disposition is a Drought_Recovery_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to rapidly restore tissue water content and turgor pressure following relief of water stress, re-establishing the cellular hydration necessary for normal metabolic function and growth. This disposition represents the hydraulic component of drought recovery, determining how quickly water status is restored when soil water becomes available. Rehydration involves water uptake through roots, transport through the xylem to shoots and leaves, and equilibration of tissue water content with xylem water potential. The speed and completeness of rehydration depend on root system function (water uptake capacity, root-soil contact integrity), xylem hydraulic conductance (potentially compromised by drought-induced embolism), and tissue properties (cell wall elasticity affecting turgor restoration, osmotic adjustment requiring solute dilution or export). Cultivars differ in Rehydration_Capacity_Disposition based on root system properties, hydraulic architecture, and the extent of drought-induced damage requiring repair. Rapid rehydration enables quick restoration of stomatal function, photosynthesis, and growth, minimizing the productivity penalty of drought episodes, while slow or incomplete rehydration prolongs recovery and may indicate persistent damage to root or vascular systems.
Rehydration_Capacity_Disposition is_a Drought_Recovery_Disposition. Rehydration is the hydraulic foundation of drought recovery, restoring tissue water content and turgor. Recovery rate depends on root function and xylem conductance, while recovery completeness reflects damage extent. Rapid rehydration enables fast restoration of stomatal function and photosynthesis. Incomplete rehydration indicates persistent hydraulic damage. Water potential recovery (easily measured) may precede full RWC recovery due to osmotic adjustment effects.
Gewebe-Wasserwiederherstellung
Post-Stress-Rehydrierung
Rehydrierungs-Kapazität-Disposition
post-stress rehydration
tissue water restoration
water status recovery
Galmes J et al. (2007) Water relations and stomatal characteristics of Mediterranean plants with different growth forms and leaf habits: responses to water stress and recovery, Plant and Soil 290:139-155 Lovisolo C et al. (2008) Whole-plant hydraulic conductance and root-to-shoot flow of abscisic acid are independently affected by water stress in grapevines, Functional Plant Biology 35:922-930 Pou A et al. (2012) Anisohydric behaviour in grapevines results in better performance under moderate water stress and recovery than isohydric behaviour, Plant and Soil 359:335-349
Rehydration Capacity Disposition
Rehydration rate quality comparison across rootstock genotypes evaluated the hydraulic basis of recovery differences observed in Chardonnay scions following moderate drought stress in Central Coast California. Following stress to predawn leaf water potential of -0.5 MPa, irrigation was restored and rehydration monitored through hourly water potential measurements. Rootstock 1103P demonstrated excellent rehydration rate quality with predawn potential recovering to -0.15 MPa (90% of well-watered baseline) within 8 hours of irrigation, attributed to its extensive, deep root system maintaining soil contact and high root hydraulic conductivity enabling rapid water uptake. Rootstock 101-14 showed intermediate rehydration rate quality with 90% recovery requiring 16 hours despite equivalent irrigation, reflecting its shallower root distribution and moderate hydraulic conductivity. Rootstock Riparia Gloire exhibited poor rehydration rate quality with 90% recovery requiring 28 hours and incomplete recovery to only 85% of baseline at 48 hours, associated with drought-induced root mortality and reduced hydraulic conductance that persisted beyond the stress period. The rehydration rate quality differential translated to scion performance, with rapid-recovering 1103P achieving 90% photosynthetic recovery by day 2 while slow-recovering Riparia Gloire required 5 days for equivalent restoration, demonstrating how rootstock rehydration rate quality influences whole-vine drought recovery.
Rehydration rate quality is a drought recovery quality that inheres in a grapevine organism and represents the velocity at which tissue water content and water potential are restored following relief of water deficit, determined by the hydraulic conductivity of the root-stem-leaf pathway, the availability and uptake capacity for soil water, and the osmotic characteristics of tissues affecting water movement into cells during the rehydration process. This quality captures the initial physical phase of drought recovery wherein water moves from newly available soil reserves through the plant to rehydrate tissues, creating the foundation for subsequent physiological recovery processes that depend on adequate tissue water status for their progression. The ontological significance of rehydration rate quality lies in its role as the rate-limiting first step in drought recovery, recognizing that physiological processes including stomatal reopening, photosynthetic restoration, and growth resumption cannot proceed until adequate tissue water content is restored, making rapid rehydration essential for minimizing the duration of post-stress impairment. The biological basis of rehydration rate encompasses the soil-root interface characteristics affecting water uptake including root hydraulic conductivity and root-soil contact, the stem and petiole hydraulic conductivity determining water transport rate through the plant, and the tissue osmotic properties affecting the thermodynamic driving force for water movement into cells from the apoplast. The occurrence of drought-induced embolism in xylem vessels reduces hydraulic conductivity and slows rehydration, creating interaction between stress severity and subsequent rehydration rate wherein severe stress causing extensive cavitation impairs the rehydration process itself. Cultivar and rootstock differences in rehydration rate quality reflect variation in hydraulic architecture, root system characteristics, and the susceptibility to drought-induced hydraulic impairment, providing genotypic variation in recovery capacity that complements differences in stress tolerance during the deficit period.
Rehydration_Rate_Quality is_a Drought_Recovery_Quality, representing the velocity of tissue water status restoration following stress relief. This class addresses the hydraulic phase of recovery that precedes physiological restoration. The time to specified recovery percentage (e.g., 90% of baseline) provides quantitative metric. Rootstock effects are particularly important given root system influence on water uptake. Drought-induced hydraulic damage slows rehydration, creating positive feedback between stress severity and recovery impairment. Distinguished from Hydraulic_Recovery_Quality which addresses repair of hydraulic damage rather than water status restoration per se.
Geweberehydratation
Rehydratationsrate
Wasseraufnahmerate
Wasserstatus-Erholung
rehydration velocity
tissue rehydration
water uptake rate
Lovisolo C et al. (2010) Drought-induced changes in development and function of grapevine (Vitis spp.) organs and in their hydraulic and non-hydraulic interactions at the whole-plant level: a physiological and molecular update. Functional Plant Biology 37:98-116 Alsina MM et al. (2011) Seasonal changes of whole root system conductance by a drought-tolerant grape root system. Journal of Experimental Botany 62:99-109 Tramontini S et al. (2013) Rootstock control of scion response to water stress in grapevine. Environmental and Experimental Botany 93:20-26
Rehydration Rate Quality
Relative water content quality assessment in water-stressed Shiraz vineyards of McLaren Vale tracks the vine and berry hydration status during the critical véraison-to-harvest period, with predawn leaf RWC of 90-95% indicating mild beneficial stress promoting phenolic development, values of 80-88% suggesting moderate stress requiring monitoring, and RWC below 75% signaling severe stress risking photosynthetic shutdown and delayed ripening berry RWC measurement complements leaf assessment, with the declining berry RWC during late ripening (from 95% to 85-90%) reflecting normal dehydration that concentrates quality compounds while excessive decline (below 80%) indicates problematic water deficit.
Relative_Water_Content_Quality represents the chemical quality characterizing the hydration status of grape tissue relative to its fully turgid state, expressing current water content as a percentage of maximum water holding capacity. Relative water content (RWC) provides a physiologically meaningful measure of tissue hydration that accounts for differences in tissue dry mass and maximum water capacity among samples, enabling comparison of water status independent of tissue composition. RWC is calculated as (fresh weight − dry weight) / (turgid weight − dry weight) × 100, where turgid weight is determined after floating tissue segments on water until full hydration is achieved. Fresh grape berries at full turgor show RWC values approaching 100%, while water-stressed or dehydrating berries exhibit progressively lower RWC as turgor pressure declines. The relationship between RWC and turgor pressure follows a pressure-volume relationship specific to tissue elasticity, with rigid cell walls maintaining turgor at lower RWC while elastic tissues lose turgor more rapidly. RWC below 85-90% typically indicates water deficit stress affecting physiological function, while values below 70-75% represent severe stress potentially causing irreversible damage. During post-harvest dehydration for raisin or passito production, RWC declines progressively as water is lost without replacement, with the dehydration rate depending on temperature, humidity, and air movement. Unlike moisture content, RWC directly indicates the degree of cellular dehydration and associated physiological stress or quality changes. RWC measurement is labor-intensive, requiring determination of fresh, turgid, and dry weights for each sample, limiting its application to research rather than routine quality assessment. The method assumes complete rehydration to original turgid weight is achievable, which may not hold for severely dehydrated or damaged tissue.
Relative_Water_Content_Quality addresses hydration status normalized to tissue maximum capacity, providing a physiologically interpretable measure distinct from absolute moisture content. The method captures the relationship between current hydration and full turgor, directly relevant to cell function and tissue stress status. RWC measurement is more complex than moisture determination, requiring the additional turgid weight determination through rehydration. The assumption of complete, non-destructive rehydration limits application to tissues capable of full turgor recovery.
Gewebehydratationsstatus
RWC
Relativer Wassergehalt
hydration percentage quality
tissue hydration status
turgor-relative water content
turgorbezogener Wassergehalt
Barrs HD and Weatherley PE (1962) A Re-Examination of the Relative Turgidity Technique for Estimating Water Deficits in Leaves. Australian Journal of Biological Sciences 15(3): 413-428 Tyree MT and Hammel HT (1972) The Measurement of the Turgor Pressure and the Water Relations of Plants by the Pressure-Bomb Technique. Journal of Experimental Botany 23(1): 267-282
Relative Water Content Quality
The Ren1 presence in Kishmish Vatkana was mapped by Hoffmann et al. (2008) to a 7.4 cM interval on chromosome 13 flanked by SSR markers UDV-020 and UDV-305, with the resistance phenotype characterized by normal appressorium formation and initial penetration by E. necator but severely restricted post-penetration hyphal growth — colony diameter at 7 days post-inoculation was reduced by 75-90% compared to susceptible V. vinifera controls, with microscopic analysis revealing thickened cell walls and enhanced callose deposition around haustoria that restrict nutrient transfer to the pathogen. Because Ren1 derives from within V. vinifera, breeding lines carrying Ren1 in a V. vinifera background maintain 100% vinifera genetic composition — a decisive advantage for wine quality acceptance compared to interspecific hybrid-derived resistance that inevitably retains some non-vinifera genome content even after extensive backcrossing.
A Ren locus information record documenting the presence or absence of the Ren1 resistance locus on chromosome 13, derived from the Central Asian Vitis vinifera subsp. sylvestris accession Kishmish Vatkana (Dzhandzhal Kara), that confers resistance to Erysiphe necator through restriction of hyphal growth following initial penetration, representing the uniquely valuable case of powdery mildew resistance identified within the V. vinifera species itself, eliminating the need for interspecific hybridization and the associated wine quality penalties of non-vinifera genetic background.
Ren1_Presence documents a resistance locus of singular strategic importance because it is the only characterized powdery mildew resistance gene found within V. vinifera itself. All other characterized powdery mildew resistance loci (Run1, Run2, Ren3, Ren4, Ren6, Ren9) derive from other Vitis species and require interspecific hybridization for introgression into V. vinifera backgrounds — a process that inevitably introduces non-vinifera genomic segments carrying genes potentially affecting wine quality, requiring extensive backcrossing (4-6 generations) to dilute the non-vinifera contribution to acceptable levels: Ren1, by contrast, can be transferred through intraspecific V. vinifera × V. vinifera crosses with no wine quality penalty from foreign species background. The discovery of Ren1 in the Uzbek table grape Kishmish Vatkana (Hoffmann et al. 2008) directed attention to Central Asian V. vinifera germplasm as a source of disease resistance genes that may exist in other wild and semi-domesticated populations from the Caucasus, Iran, Afghanistan, and western China — the geographic region where V. vinifera co-evolved with E. necator prior to the pathogen's anthropogenic introduction to Europe.
Ren1 locus genotype
Ren1 resistance status
Ren1-Präsenz
Ren1-Resistenzstatus
Vinifera-Oidium-Eigenresistenz
vinifera powdery mildew resistance
Hoffmann S et al. (2008) Resistance to Erysiphe necator in the grapevine Kishmish Vatkana is controlled by a single locus through restriction of hyphal growth, Theoretical and Applied Genetics 116: 427-438 Coleman C et al. (2009) The powdery mildew resistance gene REN1 co-segregates with an NBS-LRR gene cluster in two Central Asian grapevines, BMC Genetics 10: 89 Dry IB et al. (2010) Grapevine disease resistance genes, in Reynolds AG (ed) Viticulture and Wine Quality, Woodhead Publishing pp: 193-228
Ren1 Presence
The breeding program at University of California, Davis, systematically evaluated Ren1_Resistance_Disposition inheritance using populations derived from 'Kishmish vatkana' crossed to elite V. vinifera wine grape cultivars. A 'Kishmish vatkana' × 'Cabernet Sauvignon' F1 population segregated 1:1 for resistance, confirming single dominant gene inheritance. Resistant individuals showed characteristic Ren1 phenotype upon E. necator inoculation: germinating conidia formed appressoria and penetrated epidermal cells normally, establishing primary haustoria visible microscopically at 24 hours post-inoculation. However, by 48-72 hours, penetrated cells and 2-4 adjacent cells underwent hypersensitive death, arresting further colonization. Macroscopically, small necrotic flecks (1-2mm diameter) developed without subsequent mycelial proliferation or sporulation. In contrast, susceptible siblings supported extensive colony development with abundant conidiophore production. Marker-assisted backcrossing using the Ren1-linked marker SC8-0071-014 enabled selection of BC2 individuals retaining Ren1 with 87.5% 'Cabernet Sauvignon' genetic background, demonstrating the feasibility of introgressing Ren1 while maintaining varietal character for high-quality wine production.
A Ren1_Resistance_Disposition is a Ren_Mediated_Resistance_Disposition that inheres in a Vitis vinifera plant carrying the Ren1 locus derived from the Central Asian cultivar 'Kishmish vatkana' and is realized through effector-triggered immunity upon recognition of corresponding Erysiphe necator avirulence effectors. Ren1 holds unique significance as the only major powdery mildew resistance gene identified within the V. vinifera species itself, discovered in traditional table grape cultivars from Uzbekistan and adjacent regions where endemic E. necator populations likely exerted long-term selection pressure. The Ren1 locus maps to chromosome 13 and encodes an NLR receptor that triggers post-penetration resistance, allowing initial fungal penetration and haustorial formation but restricting subsequent hyphal growth through programmed cell death of colonized cells. This post-haustorial resistance mechanism contrasts with the pre-penetration resistance conferred by Run1. The V. vinifera origin of Ren1 provides a critical breeding advantage, as introgression does not introduce the undesirable flavor compounds, color instability, or other quality deficits associated with interspecific hybridization. However, the single germplasm source and post-penetration mechanism raise concerns about durability under high disease pressure, motivating strategies to combine Ren1 with additional resistance loci. Ren1 has been successfully deployed in breeding programs worldwide, contributing to cultivars that combine high wine quality with reduced fungicide requirements.
Ren1_Resistance_Disposition is_a Ren_Mediated_Resistance_Disposition. Ren1 is uniquely derived from V. vinifera rather than wild Vitis species, located on chromosome 13. The post-penetration resistance mechanism distinguishes Ren1 from Run1 which operates pre-penetration. The V. vinifera origin eliminates interspecific hybridization quality penalties but limits genetic diversity of resistance sources. Combining Ren1 with Run or other Ren loci is recommended for enhanced durability.
Kishmish vatkana resistance
Kishmish-vatkana-Resistenz
Ren1 powdery mildew resistance
Ren1-Oidium-Resistenz
Ren1-Resistenz-Disposition
V. vinifera PM resistance
Hoffmann S et al. (2008) Resistance to Erysiphe necator in the grapevine 'Kishmish vatkana' is controlled by a single locus through restriction of hyphal growth, Theoretical and Applied Genetics 116:427-438 Coleman C et al. (2009) The powdery mildew resistance gene REN1 co-segregates with an NBS-LRR gene cluster in two Central Asian grapevines, BMC Genetics 10:89 Riaz S et al. (2013) Genetic mapping of Ren1 and development of markers for marker-assisted selection of powdery mildew resistance in grapevine, Theoretical and Applied Genetics 126:1355-1363
Ren1 Resistance Disposition
Genetic characterization of Ren2_Resistance_Disposition at the University of Missouri utilized mapping populations derived from the resistant V. cinerea accession B9. QTL analysis identified a major resistance locus on chromosome 14 explaining 72% of phenotypic variance for powdery mildew severity, subsequently designated Ren2. Resistant individuals carrying the Ren2 allele displayed hypersensitive response within 24-36 hours of E. necator penetration attempts, with microscopic examination revealing intense hydrogen peroxide accumulation (visualized by DAB staining) in cells contacted by fungal appressoria. Macroscopically, Ren2 carriers developed small brown flecks (<1mm diameter) at inoculation sites without subsequent mycelial growth, contrasting with susceptible V. vinifera 'Norton' controls showing extensive white powdery colonies. The Ren2-linked SSR marker VMC7f2 enables marker-assisted selection in breeding populations. Advanced selections combining Ren2 with other resistance sources showed complete immunity in field trials at Mountain Grove, Missouri, under natural disease pressure that caused 60-80% leaf coverage on susceptible controls, demonstrating effective pyramiding of Ren2 with complementary resistance genes.
A Ren2_Resistance_Disposition is a Ren_Mediated_Resistance_Disposition that inheres in a Vitis plant carrying the Ren2 locus derived from Vitis cinerea and is realized through effector-triggered immunity upon recognition of corresponding Erysiphe necator avirulence effectors. The Ren2 locus maps to chromosome 14 and originates from V. cinerea, a North American wild species distributed across the southeastern and central United States that evolved in sympatry with endemic Erysiphe populations. V. cinerea exhibits exceptional powdery mildew resistance and has contributed resistance factors to several French-American hybrid cultivars developed in the 19th and early 20th centuries. The Ren2 resistance phenotype is characterized by rapid hypersensitive response with localized cell death arresting fungal development at early post-penetration stages, resulting in small necrotic lesions without macroscopic colony establishment or sporulation. The chromosomal location of Ren2 on chromosome 14 is independent from other major resistance loci, making it valuable for pyramiding strategies. However, V. cinerea introgression is associated with characteristic herbaceous and foxy flavor compounds that require extensive backcrossing to eliminate while retaining the resistance locus. Despite these quality challenges, Ren2 provides a robust resistance source with recognition specificity complementary to other Ren and Run loci, contributing to comprehensive resistance breeding strategies that combine multiple genes for durable protection.
Ren2_Resistance_Disposition is_a Ren_Mediated_Resistance_Disposition. Ren2 originates from V. cinerea, a North American species distinct from the Muscadinia source of Run loci. The chromosome 14 location is independent from other major PM resistance loci. V. cinerea ancestry introduces quality challenges (herbaceous flavors) requiring extensive backcrossing. Ren2 has contributed to resistance in some French-American hybrids, though its molecular identity remains to be fully characterized.
Cinerea-Mehltau-Resistenz
Ren2 powdery mildew resistance
Ren2-Oidium-Resistenz
Ren2-Resistenz-Disposition
V. cinerea PM resistance
chromosome 14 PM resistance
Dalbó MA et al. (2001) A gene controlling sex in grapevines placed on a molecular marker-based genetic map, Genome 44:75-82 Qiu W et al. (2015) Current understanding of grapevine defense mechanisms against the biotrophic fungus (Erysiphe necator), Horticulture Research 2:15020
Ren2 Resistance Disposition
The Ren3 presence in the PIWI cultivar Regent was mapped by Welter et al. (2007) and refined by Zyprian et al. (2016) to a region on chromosome 15 spanning approximately 5 cM and containing a cluster of resistance gene analogs — Regent's Ren3 resistance reduces powdery mildew severity by 40-60% compared to susceptible V. vinifera controls in field conditions, with the resistance phenotype characterized by delayed colony expansion, reduced sporulation density, and enhanced epidermal cell necrosis at infection sites. Ren3 acts in combination with the linked but distinct Ren9 locus (also on chromosome 15, separated by approximately 10 cM) to provide the composite powdery mildew resistance observed in Regent and related PIWI cultivars — the two loci are typically co-inherited due to their physical proximity, making genetic separation for independent deployment challenging without very large mapping populations.
A Ren locus information record documenting the presence or absence of the Ren3 resistance locus on chromosome 15, derived from North American Vitis species through complex interspecific hybrid pedigrees, that confers partial quantitative resistance to Erysiphe necator powdery mildew, representing the most widely deployed powdery mildew resistance source in current PIWI cultivar production alongside the linked Ren9 locus.
Ren3_Presence documents the powdery mildew resistance locus most widely present in commercially grown PIWI cultivars, analogous to the role of Rpv3 for downy mildew resistance. Ren3 derives from North American Vitis species (likely V. lincecumii or V. rupestris through the Seibel and Seyve-Villard hybrid lineages) and was first characterized in the cultivar Regent, which has served as a model genotype for both powdery and downy mildew resistance research: the partial (quantitative) nature of Ren3 resistance means that it reduces but does not eliminate powdery mildew development, typically requiring 1-3 supplementary fungicide applications in high-pressure seasons compared to 8-12 for fully susceptible cultivars — a 60-80% reduction that is commercially significant but insufficient for zero-spray cultivation. The linkage of Ren3 with Ren9 on chromosome 15 creates a functionally important resistance cluster, but also poses a breeding challenge because the two loci cannot be easily separated and recombined independently with other resistance sources, effectively functioning as a single compound resistance unit in most crossing programs.
North American powdery mildew resistance
Ren3 locus genotype
Ren3 resistance status
Ren3-Präsenz
Ren3-Resistenzstatus
amerikanische Oidium-Resistenz
Welter LJ et al. (2007) Genetic mapping and localization of quantitative trait loci affecting fungal disease resistance and leaf morphology in grapevine, Molecular Breeding 20: 359-374 Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance and ripening of grapevines, Molecular Breeding 36: 65 Dry IB et al. (2010) Grapevine disease resistance genes, in Reynolds AG (ed) Viticulture and Wine Quality, Woodhead Publishing pp: 193-228
Ren3 Presence
Field evaluation of Ren3_Resistance_Disposition at the Geilweilerhof research station, Germany, compared powdery mildew development on 'Regent' (Ren3 carrier) versus susceptible 'Müller-Thurgau' over five consecutive seasons under reduced spray programs (3-4 applications versus standard 8-10). 'Regent' exhibited consistent partial resistance with mean leaf infection severity of 12-18% (OIV 452 rating 5-6) compared to 65-85% in untreated 'Müller-Thurgau' controls. Microscopic examination revealed that E. necator successfully established colonies on 'Regent' leaves, but colony expansion was restricted to approximately 40% of the diameter observed on susceptible tissue, and sporulation intensity was reduced by 60-70%. Importantly, monitoring studies detected E. necator isolates with increased aggressiveness on Ren3 backgrounds in vineyards with high 'Regent' adoption, emphasizing the partial nature of this resistance and the importance of combining Ren3 with additional loci. Breeding selections pyramiding Ren3 with Ren9 or Run1 displayed enhanced resistance (leaf infection <5%) compared to Ren3 alone, validating complementation strategies.
A Ren3_Resistance_Disposition is a Ren_Mediated_Resistance_Disposition that inheres in a Vitis plant carrying the Ren3 locus and is realized through effector-triggered immunity conferring partial resistance to Erysiphe necator. The Ren3 locus maps to chromosome 15 and was identified in the PIWI cultivar 'Regent' and related breeding material derived from complex interspecific pedigrees involving French-American hybrids. Unlike Run1 or Ren1 which confer near-complete resistance, Ren3 provides quantitative resistance characterized by reduced colony size, decreased sporulation intensity, and extended latent period rather than complete immunity. This partial resistance phenotype allows limited fungal colonization and reproduction, raising concerns about selection pressure on pathogen populations and potential resistance erosion under intensive deployment. The genetic architecture of Ren3 resistance may involve multiple linked genes within a resistance gene cluster on chromosome 15, with the major QTL explaining 50-70% of phenotypic variance in segregating populations. Despite its partial nature, Ren3 has been widely deployed in European PIWI cultivars including 'Regent', 'Johanniter', 'Cabernet Cortis', and others, providing meaningful field protection that reduces fungicide requirements by 50-70% compared to fully susceptible V. vinifera cultivars. Combining Ren3 with additional resistance loci is recommended to enhance protection level and durability.
Ren3_Resistance_Disposition is_a Ren_Mediated_Resistance_Disposition. Ren3 confers partial rather than complete resistance, distinguishing it from Run1, Ren1, and Ren4 which provide near-immunity. The locus on chromosome 15 may represent a complex of linked resistance factors. Ren3 is widely deployed in European PIWI cultivars but shows evidence of durability challenges similar to Rpv3 for downy mildew. Pyramiding with other Ren or Run loci is strongly recommended.
Regent PM resistance
Regent-Mehltau-Resistenz
Ren3 powdery mildew resistance
Ren3-Oidium-Resistenz
Ren3-Resistenz-Disposition
chromosome 15 PM resistance
Welter LJ et al. (2007) Genetic mapping and localization of quantitative trait loci affecting fungal disease resistance and leaf morphology in grapevine, Molecular Breeding 20:359-374 Zendler D et al. (2017) Fine mapping of Ren3 reveals two loci mediating hypersensitive response against Erysiphe necator in grapevine, Euphytica 213:68 Töpfer R, Hausmann L (2010) Breeding for fungal disease resistant grapevines, Genetics, Genomics and Breeding of Grapes, Science Publishers
Ren3 Resistance Disposition
The Ren4 presence was mapped by Riaz et al. (2011) in a V. romanetii-derived population to a 1.2 cM interval on chromosome 18 in a region distinct from the Rpv3 downy mildew resistance cluster on the same chromosome — Ren4-carrying individuals showed 60-80% reduction in powdery mildew colony development, with the resistance mechanism characterized by rapid formation of enlarged callose papillae at the inner surface of the epidermal cell wall beneath the fungal appressorium, physically blocking penetration peg entry in 50-70% of attempted penetrations compared to less than 10% in susceptible V. vinifera. The geographic origin of Ren4 in a Chinese Vitis species is consistent with the hypothesis that E. necator originated in East Asia, as V. romanetii populations in Hubei and Sichuan provinces would have co-evolved with the pathogen and been subject to natural selection for resistance over evolutionary timescales.
A Ren locus information record documenting the presence or absence of the Ren4 resistance locus on chromosome 18, derived from Vitis romanetii (a Chinese wild Vitis species), that confers resistance to Erysiphe necator through a mechanism involving enhanced callose deposition and papilla formation at penetration sites, providing an East Asian-origin powdery mildew resistance source phylogenetically and mechanistically distinct from both the muscadine-derived Run loci and the North American-derived Ren3.
Ren4_Presence documents a powdery mildew resistance source from the center of diversity of the genus Vitis — China, where approximately 30 of the 60-80 recognized Vitis species are endemic and where co-evolution with E. necator has produced diverse resistance mechanisms across multiple species. V. romanetii is a vigorous climbing species native to central China that has received limited attention in Western breeding programs compared to North American species, but its resistance genes offer three strategic advantages: first, the geographic and phylogenetic independence from both muscadine (Run) and North American (Ren3) resistance sources provides different pathogen recognition specificities for pyramiding second, the callose-based penetration resistance mechanism of Ren4 represents a pre-haustorial defense distinct from the post-haustorial hypersensitive response of Run1 and the post-penetration growth restriction of Ren1 and third, the location of Ren4 on chromosome 18 (the same chromosome as Rpv3 but at a different position) creates potential complications for breeding programs seeking to combine both loci through recombination, requiring careful marker-assisted selection to recover recombinant chromosomes carrying both resistance genes.
Chinese Vitis powdery mildew resistance
Ren4 locus genotype
Ren4 resistance status
Ren4-Präsenz
Ren4-Resistenzstatus
chinesische Oidium-Resistenz
Riaz S et al. (2011) Identification of mildew resistance in wild and cultivated Central Asian grape germplasm, BMC Plant Biology 11: 149 Dry IB et al. (2010) Grapevine disease resistance genes, in Reynolds AG (ed) Viticulture and Wine Quality, Woodhead Publishing pp: 193-228 Ramming DW et al. (2011) A single dominant locus, Ren4, confers rapid non-race-specific resistance to grapevine powdery mildew, Phytopathology 101: 502-508
Ren4 Presence
Comprehensive phenotypic characterization of Ren4_Resistance_Disposition at Cornell University's Geneva Experiment Station employed controlled inoculation experiments with E. necator isolates representing genetic groups A (flag shoot), B (ascosporic), and multiple genotypes within each group. The V. romanetii accession C166-043 carrying Ren4 exhibited complete immunity against all 23 tested isolates, with no visible symptoms or microscopic evidence of successful penetration events through 28 days post-inoculation at inoculum densities up to 50,000 conidia/mL. Histological examination using calcoflour white staining revealed that 98% of germinated conidia formed appressoria normally but failed at the penetration stage, with intense papilla formation visible beneath attempted penetration sites. The remaining 2% of penetration attempts triggered immediate single-cell hypersensitive death. In F1 populations from Ren4 × V. vinifera crosses, resistance segregated 1:1 confirming dominant single-gene inheritance. BC1 and BC2 selections combining Ren4 with 75-87.5% V. vinifera background retained complete resistance while showing progressive improvement in fruit quality parameters including increased berry size, reduced titratable acidity, and more neutral flavor profile.
A Ren4_Resistance_Disposition is a Ren_Mediated_Resistance_Disposition that inheres in a Vitis plant carrying the Ren4 locus derived from Vitis romanetii and is realized through effector-triggered immunity conferring exceptionally strong, broad-spectrum resistance to Erysiphe necator. The Ren4 locus maps to chromosome 18 and originates from V. romanetii, a Chinese wild species closely related to V. piasezkii and V. davidii that evolved in regions with endemic powdery mildew pressure. Ren4 confers one of the strongest powdery mildew resistance phenotypes observed in Vitis, characterized by rapid pre-penetration arrest with intense papilla formation blocking fungal entry at the cell wall, complemented by hypersensitive response in the rare cells where penetration succeeds. This multi-layered defense results in complete immunity across diverse E. necator genetic groups and isolates, with no macroscopic or microscopic evidence of fungal colonization even under extreme inoculum pressure. The broad-spectrum efficacy and pre-penetration mechanism make Ren4 exceptionally valuable for resistance breeding, though V. romanetii introgression introduces quality challenges including small berry size, high acidity, and distinctive flavor compounds requiring extensive backcrossing to recover commercial quality characteristics. Marker-assisted selection enables efficient introgression of Ren4 while progressively restoring V. vinifera background through marker-assisted backcrossing programs.
Ren4_Resistance_Disposition is_a Ren_Mediated_Resistance_Disposition. Ren4 originates from V. romanetii, a Chinese species, and maps to chromosome 18. The exceptionally strong, pre-penetration resistance mechanism distinguishes Ren4 from Ren1 (post-penetration) and Ren3 (partial). Broad-spectrum efficacy across E. necator genetic groups makes Ren4 particularly valuable. V. romanetii ancestry introduces significant quality challenges requiring extensive backcrossing.
Chinese grape PM resistance
Ren4 powdery mildew resistance
Ren4-Oidium-Resistenz
Ren4-Resistenz-Disposition
Romanetii-Mehltau-Resistenz
V. romanetii PM resistance
Ramming DW et al. (2011) A single dominant locus, Ren4, confers rapid non-race-specific resistance to grapevine powdery mildew, Phytopathology 101:502-508 Riaz S et al. (2011) Identification of mildew resistance in wild and cultivated Central Asian grape germplasm, BMC Plant Biology 11:149 Qiu W et al. (2015) Current understanding of grapevine defense mechanisms against the biotrophic fungus (Erysiphe necator), Horticulture Research 2:15020
Ren4 Resistance Disposition
Genetic mapping of Ren6_Resistance_Disposition utilized a population derived from the resistant V. piasezkii accession DVIT2027 crossed to susceptible V. vinifera 'Malaga Rosada'. QTL analysis identified a major resistance locus on chromosome 9 explaining 67% of phenotypic variance, designated Ren6. Phenotypic evaluation using detached leaf disc assays with E. necator conidia demonstrated that Ren6 carriers exhibited hypersensitive response within 36-48 hours post-inoculation, with small necrotic lesions (diameter <1mm) and no subsequent mycelial growth or sporulation. Interestingly, a second linked QTL (Ren7) was detected 8 cM from Ren6, contributing an additional 15% of phenotypic variance. Individuals carrying both Ren6 and Ren7 showed enhanced resistance compared to Ren6-only carriers, with more rapid hypersensitive response and smaller necrotic lesions. The V. piasezkii resistance gene cluster on chromosome 9 provides breeding programs with a valuable new source for powdery mildew resistance, particularly attractive because the same species carries Rpv5 downy mildew resistance, enabling potential dual-disease resistance introgression from a single donor.
A Ren6_Resistance_Disposition is a Ren_Mediated_Resistance_Disposition that inheres in a Vitis plant carrying the Ren6 locus derived from Vitis piasezkii and is realized through effector-triggered immunity conferring strong resistance to Erysiphe necator. The Ren6 locus maps to chromosome 9 and originates from V. piasezkii, a Chinese wild species native to central and western China that exhibits exceptional resistance to both powdery and downy mildew diseases. Ren6 was identified as genetically distinct from other characterized resistance loci through allelism tests and molecular mapping, providing an additional independent resistance source for pyramiding strategies. The Ren6 resistance phenotype is characterized by rapid hypersensitive response arresting fungal development shortly after penetration, resulting in small necrotic flecks without subsequent colony expansion or sporulation. Importantly, Ren6 is closely linked to Ren7, another powdery mildew resistance locus identified in the same V. piasezkii accessions, and both loci are typically co-inherited as a resistance gene cluster. The combination of Ren6/Ren7 provides robust protection that has proven effective against diverse E. necator populations in multi-environment trials. V. piasezkii also carries the Rpv5 locus for downy mildew resistance, potentially enabling simultaneous introgression of resistance to both major fungal diseases from this single wild species source, analogous to the Run1/Rpv1 cluster from Muscadinia.
Ren6_Resistance_Disposition is_a Ren_Mediated_Resistance_Disposition. Ren6 originates from V. piasezkii and maps to chromosome 9, where it is closely linked to Ren7 forming a resistance gene cluster. The co-inheritance of Ren6 and Ren7 provides enhanced resistance compared to either locus alone. V. piasezkii additionally carries Rpv5 for downy mildew resistance, offering potential for combined introgression of both disease resistances from this single wild species source.
Piasezkii-Mehltau-Resistenz
Ren6 powdery mildew resistance
Ren6-Oidium-Resistenz
Ren6-Resistenz-Disposition
V. piasezkii PM resistance
chromosome 9 PM resistance
Pap D et al. (2016) Identification of two novel powdery mildew resistance loci, Ren6 and Ren7, from the wild Chinese grape species Vitis piasezkii, BMC Plant Biology 16:170 Riaz S et al. (2020) Genetic analysis of powdery mildew resistance in Chinese wild Vitis species through association mapping, Molecular Breeding 40:89
Ren6 Resistance Disposition
The Ren locus information compiled in the VIVC resistance locus registry documents over 11 designated Ren loci identified across diverse Vitis species — Ren1 on chromosome 13 (from V. vinifera subsp. sylvestris accession Kishmish Vatkana, the only powdery mildew resistance locus identified within the V. vinifera species itself), Ren3 on chromosome 15 (from North American hybrid ancestry, present in Regent and other PIWI cultivars), Ren4 on chromosome 18 (from V. romanetii, a Chinese species), Ren6 on chromosome 9 (from V. piasezkii, another Chinese species), and Ren9 on chromosome 15 (linked to Ren3 but distinct) — these loci provide a diverse toolkit of powdery mildew resistance sources spanning the geographic range of the genus Vitis from Central Asia through East Asia to North America, enabling the construction of multi-locus resistance pyramids incorporating resistance genes with independent evolutionary origins.
A resistance gene information record specific to Ren (Resistance to Erysiphe necator) loci — the genomic regions derived from Euvitis species (non-muscadine Vitis) conferring resistance to powdery mildew — documenting the locus designation (Ren1 through Ren11+), chromosomal location, source species, resistance level and mechanism, and diagnostic molecular markers, with Ren loci providing the most accessible resistance sources for powdery mildew breeding due to the easy hybridization compatibility between Euvitis species and V. vinifera.
Ren_Locus_Information documents the larger and more diverse set of powdery mildew resistance loci compared to the Run loci, reflecting the broader taxonomic sampling across Euvitis species that has been possible due to the absence of the hybridization barriers that limit muscadine introgression. The Ren loci collectively span at least 8 chromosomes and derive from at least 6 Vitis species, providing resistance mechanisms ranging from programmed cell death (Ren1) through enhanced callose and lignin deposition (Ren4) to accelerated stilbene accumulation (Ren3): this mechanistic diversity is the genetic foundation for constructing durable resistance pyramids in which each locus presents the pathogen with a different defense barrier. The discovery of Ren1 in V. vinifera subsp. sylvestris from Central Asia (Hoffmann et al. 2008) was particularly significant because it demonstrated that powdery mildew resistance exists within the V. vinifera gene pool itself — a finding consistent with the hypothesis that E. necator originated in East Asia or Central Asia (where wild V. vinifera populations co-evolved with the pathogen) and was introduced to Europe only in the 1840s, explaining why European cultivars lack resistance while Central Asian populations retain it.
Erysiphe resistance QTL data
Euvitis powdery mildew resistance locus data
Euvitis-Mehltau-Resistenzlocus-Daten
Oidium-Resistenz-QTL-Daten
Ren gene characterization
Ren-Locus-Information
Hoffmann S et al. (2008) Resistance to Erysiphe necator in the grapevine Kishmish Vatkana is controlled by a single locus through restriction of hyphal growth, Theoretical and Applied Genetics 116: 427-438 Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance, Molecular Breeding 36: 65 VIVC (2023) Loci for resistance to diseases and pests Dry IB et al. (2010) Grapevine disease resistance genes, in Reynolds AG (ed) Viticulture and Wine Quality, Woodhead Publishing pp: 193-228
Ren Locus Information
The grapevine breeding consortium VitisGen evaluated Ren_Mediated_Resistance_Disposition across a diversity panel of 380 Vitis accessions using standardized greenhouse screening with E. necator isolate NY19 (genetic group B). Genotypes were scored 21 days post-inoculation using a 0-5 visual scale for colony coverage and sporulation intensity. Accessions carrying characterized Ren loci exhibited dramatically reduced infection compared to susceptible V. vinifera controls (mean score 4.7). Ren1 carriers ('Kishmish vatkana' derivatives) scored 0.3-0.8 with sparse hypersensitive flecking. Ren4 carriers (V. romanetii accessions) scored 0.0-0.2 with no visible colony development. Ren3 carriers ('Regent', 'Villard blanc') scored 1.2-1.8 with small, weakly sporulating colonies. The phenotypic variation among Ren loci reflects differences in recognition timing and downstream response intensity. Marker-assisted selection using locus-specific SSR markers enabled efficient identification of resistance alleles in segregating populations, with pyramided Ren1+Ren4 individuals displaying enhanced resistance (score 0.0) compared to single-locus carriers, supporting the strategy of combining multiple Ren sources for comprehensive protection.
A Ren_Mediated_Resistance_Disposition is a Powdery_Mildew_Resistance_Disposition that inheres in a Vitis plant carrying one or more Ren (Resistance from Erysiphe necator) loci and is realized through effector-triggered immunity upon recognition of corresponding Erysiphe necator avirulence effectors. The Ren designation encompasses major resistance genes identified in various Vitis species outside of Muscadinia rotundifolia, distinguishing them from Run loci which specifically derive from Muscadinia. Over ten Ren loci have been mapped across the Vitis genus, originating from diverse species including V. vinifera (Ren1 from 'Kishmish vatkana'), V. cinerea (Ren2), interspecific hybrids (Ren3 from 'Regent'), V. romanetii (Ren4), V. piasezkii (Ren6), and others. Each Ren locus encodes NLR receptor proteins recognizing specific E. necator effectors, triggering defense responses that arrest pathogen development at various infection stages. The resistance phenotypes conferred by different Ren loci range from complete immunity with rapid hypersensitive response to partial resistance with reduced colony size and sporulation. The diversity of Ren loci origins provides extensive opportunities for resistance gene pyramiding, combining loci with distinct recognition specificities to achieve durable field resistance. Notably, Ren1 represents the only major powdery mildew resistance gene identified within V. vinifera itself, offering the unique advantage of introgression without the quality penalties associated with wild species ancestry, though its single-source origin raises durability concerns that favor combination with additional Ren or Run loci.
Ren_Mediated_Resistance_Disposition is_a Powdery_Mildew_Resistance_Disposition. Ren loci are distinguished from Run loci by their non-Muscadinia origins, though both confer major gene resistance to Erysiphe necator. The Ren nomenclature follows the convention established when the first locus outside Muscadinia was characterized. Child classes represent individual Ren loci numbered sequentially by discovery order. Some Ren loci (e.g., Ren5, Ren7) are omitted from VVO pending validation or due to limited breeding utility.
Ren locus powdery mildew resistance
Ren-Lokus-Oidium-Resistenz
Ren-Mehltau-Resistenz
Ren-vermittelte Resistenz-Disposition
Vitis-derived PM resistance
non-Muscadinia PM resistance
Hoffmann S et al. (2008) Resistance to Erysiphe necator in the grapevine 'Kishmish vatkana' is controlled by a single locus through restriction of hyphal growth, Theoretical and Applied Genetics 116:427-438 Qiu W et al. (2015) Current understanding of grapevine defense mechanisms against the biotrophic fungus (Erysiphe necator), Horticulture Research 2:15020 Dry IB et al. (2019) Grapevine NLR immunity against downy and powdery mildew pathogens, Acta Horticulturae 1248:129-136
Ren Mediated Resistance Disposition
A renewal cane on Vitis vinifera cv. Pinot Noir in a Guyot training system, retained at 10 nodes during January pruning alongside a 12-node fruiting cane from the same spur position, positioned closer to the permanent cordon than the fruiting cane to maintain compact vine architecture, measuring 9mm diameter with complete brown lignification indicating successful maturation, expected to produce 9-10 shoots during current season with shoots at nodes 8-10 providing ideal positioning and vigor for selection as following winter's fruiting canes, while the current fruiting cane will be completely removed after harvest, perpetuating annual cycle of cane replacement maintaining consistent fruiting zone position and vine balance.
A one-year cane retained during winter pruning specifically to provide fruiting wood for subsequent seasons rather than current season crop production, characterized by appropriate positioning for future fruiting cane selection, adequate length to accommodate following season's pruning requirements (typically 8-15 nodes retained), complete lignification ensuring winter survival and vigorous spring growth, and deliberate retention as part of sustainable pruning strategy maintaining continuous supply of well-positioned young wood for annual fruiting cane replacement. Renewal canes represent critical component of balanced cane-pruning systems, ensuring availability of appropriately positioned fruiting wood for future seasons, preventing progressive displacement of fruiting zones away from permanent structure, maintaining vine architecture and trellis integration, and supporting long-term vineyard sustainability through planned wood renewal rather than opportunistic or crisis-driven cane selection.
Renewal canes are distinguished from fruiting canes (providing current crop) and permanent structure (multi-year arms, cordons, trunks). The renewal cane concept is fundamental to cane-pruned systems, with various regional traditions employing different renewal strategies - Guyot systems typically retain one fruiting cane plus one renewal cane per spur position, while other systems may use multiple renewal options or integrated fruiting-renewal approaches where portions of fruiting canes provide renewal positions. Failure to maintain adequate renewal capacity leads to progressive problems including fruiting zone displacement, excessive cane lengths to reach trellis positions, declining wood quality, and eventual training system breakdown requiring drastic corrective pruning or retraining.
Erneuerungsrute
Ersatzrute
Vorratsrute
next-year cane
replacement cane
succession cane
Renewal Cane
A guyot-trained Pinot Noir vine has a Renewal_Zone_Role at the 2-bud spur positioned below the fruiting cane, with the two shoots emerging from this spur competing to become next year's selected fruiting cane—the renewal zone requiring protection from damage and adequate light for proper shoot development. A bilateral cordon Cabernet Sauvignon vine bears Renewal_Zone_Roles at each spur position along the cordons, with the basal bud of each spur providing renewal potential while the second bud contributes to current-season crop. A head-trained Grenache vine has its Renewal_Zone_Role concentrated at the vine head, where all replacement arms and spurs must develop from the limited head surface area.
A canopy position role that inheres in the spatial region of a grapevine canopy designated for the production of replacement wood—shoots that will become next season's fruiting canes or spurs, maintaining the vine's productive structure through annual regeneration. The renewal zone is particularly critical in cane-pruned systems (guyot variants) where specific positions must produce well-developed, appropriately located shoots suitable for selection as replacement canes at dormant pruning. Renewal zone positioning varies with training system: in guyot systems, the renewal spur (typically 2 buds) positioned below the fruiting cane defines the renewal zone in cordon systems, basal positions on existing spurs provide renewal shoots in head-trained systems, the head itself constitutes the renewal zone from which all annual growth emerges. Renewal zone management ensures continuity of vine structure—failure to develop adequate replacement shoots forces retention of poorly positioned wood, use of water shoots (epicormic growth), or structural renovation that disrupts production. The renewal zone concept recognizes that grape production requires forward planning current-season shoot development directly determines next-season fruiting capacity.
Renewal_Zone_Role captures the forward-looking management perspective essential to sustainable grape production—current decisions about shoot retention and positioning directly affect future productivity. The renewal zone is most explicitly managed in cane-pruned systems where specific renewal spurs must produce suitable replacement canes, but the concept applies across training systems wherever the origin points for future fruiting wood are identifiable. Renewal zone failure—damage, disease, or inadequate shoot development—creates structural problems that may take years to correct, as repositioning fruiting wood or redeveloping vine architecture requires multiple growing seasons. Training system selection implicitly defines renewal zone requirements systems with higher renewal demands (guyot requiring annual cane selection) are more vulnerable to renewal zone problems than systems with distributed renewal potential (cordons with multiple spur positions).
Erneuerungszone
Ersatzholzzone
Regenerationszone
Zapfenzone
regeneration zone
renewal zone
replacement zone
spur zone
Galet P (2000) General Viticulture, Oenoplurimedia Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles Winkler AJ et al. (1974) General Viticulture, University of California Press
Renewal Zone Role
A replacement renewal cane on Vitis vinifera cv. Chardonnay in vertical shoot positioning system, retained at 11 nodes during December pruning positioned 8cm closer to cordon than the 12-node fruiting cane, both arising from previous season's two-bud spur, measuring 9.5mm diameter with complete lignification, positioned to produce upward-growing shoots along fruiting wire, expected to develop 10-11 shoots during current season with apical 2-3 shoots (nodes 9-11) providing ideal vigor and position for selection as two 12-node fruiting canes during following winter's pruning, while current season's fruiting cane will be completely removed, perpetuating annual Guyot-style cane replacement maintaining consistent architecture and productivity.
A renewal cane retained with primary purpose of providing immediate replacement fruiting wood for the following season, characterized by positioning directly adjacent to current season's fruiting cane, appropriate length and node count for anticipated next-season fruiting requirements (typically matching current fruiting cane length), selection prioritizing optimal location for maintaining consistent fruiting zone position, and expectation of producing shoots that will be pruned to fruiting canes during next dormant season while current fruiting cane is removed. Replacement renewal canes represent standard annual renewal strategy in most cane-pruned systems, ensuring continuous availability of well-positioned fruiting wood through planned succession, preventing fruiting zone migration or displacement, and maintaining vine architecture and productivity through systematic annual wood replacement following established pruning formulas and training system requirements.
Replacement renewal canes follow established formulas in traditional cane-pruned regions, with Guyot systems typically employing one fruiting cane plus one renewal cane per position, while double Guyot uses two fruiting canes plus two renewal canes. The replacement strategy ensures fruiting wood availability even if some positions fail (winter damage, weak growth, poor positioning), provides selection options for choosing best-quality canes during pruning (vigor, diameter, lignification, position), and maintains training system integrity through predictable annual replacement cycles. Skilled pruners select replacement renewals during previous winter anticipating current season's growth patterns and next winter's requirements, demonstrating forward-thinking planning essential for sustainable cane-pruned viticulture.
Ersatz-Erneuerungsrute
Hauptvorratsrute
direct replacement cane
primary renewal cane
succession renewal cane
Replacement Renewal Cane
A replacement secondary vegetative bud at node 6 on Vitis vinifera cv. Chardonnay in a double-pruning frost avoidance program, where primary buds were deliberately removed during pre-budbreak pruning to delay phenology by 10-14 days, this secondary bud breaking in mid-May rather than late April, developing into a 95cm vegetative shoot with 14 leaves that was retained during summer canopy management and winter-pruned to a 6-node cane, with 4 of those 6 nodes producing fruitful primary shoots in the following spring bearing a combined 7 clusters totaling 1.8 kg fruit, successfully converting the first-year vegetative sacrifice into second-year productive fruiting wood while avoiding the late April frost that damaged 60% of primary buds in neighboring conventionally-pruned blocks.
A secondary vegetative bud that develops into a shoot serving as replacement fruiting wood for subsequent seasons, activated either following primary bud damage requiring canopy reconstruction or deliberately through cultural practices like double pruning where primary shoots are removed to force secondary growth as a frost avoidance strategy or vigor management technique. Replacement secondary vegetative buds contribute to long-term vineyard productivity by establishing new fruiting positions when primary structures are lost, with shoots developing from these buds trained and pruned during dormancy to create canes or spurs bearing fruitful primary buds for the following season, thus converting a current-season vegetative loss into future-season productive capacity through the annual cycle of bud differentiation and shoot development.
Replacement secondary vegetative buds play dual roles in viticulture= unplanned replacement following damage events, and planned replacement in frost management or vigor control strategies. Double pruning intentionally sacrifices primary buds to force secondary growth, delaying budbreak past typical frost dates while accepting current-year yield loss for subsequent-year protection and production. This strategy is economically viable only in high-frost-risk regions where expected damage exceeds the cost of current-year crop sacrifice. The quality of replacement wood from secondary vegetative shoots varies with management, with well-positioned, moderately-vigorous shoots producing excellent fruiting wood while excessively vigorous shoots may show reduced fruitfulness due to shading during their own bud differentiation period.
Erneuerungs-Nebenknospe
Ersatz-Sekundärknospe
compensation secondary bud
future-fruiting secondary bud
renewal secondary vegetative bud
Based on Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Palliotti et al. (2014) "Changes in vineyard establishment and canopy management urged by earlier climate-related grape ripening= A review" Scientia Horticulturae 178=43-54 Friend & Trought (2007) "Delayed winter spur-pruning in New Zealand can alter yield components of Merlot grapevines" Australian Journal of Grape and Wine Research 13=157-164.
Replacement Secondary Vegetative Bud
A reproductive active bud at node 5 on a cane of Vitis vinifera cv. Cabernet Sauvignon in late April in Napa Valley, California, developing into a shoot with 2 visible inflorescences of 185 and 160 flowers each, 4 expanded leaves, shoot length 18cm, inflorescences at BBCH stage 55 (individual flowers separating), representing potential yield of approximately 345 berries if fruit set achieves typical 50% for this cultivar under these conditions.
An active bud containing differentiated inflorescence primordia that develops into a shoot bearing one or more flower clusters in addition to leaves, characterized by emergence of both vegetative structures (leaves, tendrils) and reproductive structures (inflorescences) from the same growing point, with inflorescence development proceeding through stages of cluster emergence, flower separation, flowering (anthesis), and fruit set concurrent with vegetative shoot growth. Reproductive active buds represent the primary source of current season crop yield, with inflorescence number per bud (typically 1-3 in Vitis vinifera), flower number per inflorescence (typically 50-400), and successful fruit set percentage (typically 30-60%) determining potential cluster and berry number, all influenced by environmental conditions during both primordium formation (previous season) and current season development.
Reproductive active buds are the foundation of wine grape production, with their development success determining yield quantity and quality. The number of inflorescence primordia per bud was determined 12-15 months prior during the previous season's critical inflorescence initiation period, while flower number per inflorescence was largely fixed 8-10 months prior. Current season environmental stress during flowering can dramatically reduce fruit set, while excessive vigor can promote vegetative growth at the expense of reproductive development. Balanced vine management aims to optimize reproductive bud development without excessive vegetative competition.
Blütenknospe
fertile active bud
flowering active bud
fruiting active bud
reproduktive aktive Knospe
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Vasconcelos et al. (2009) "The flowering process of Vitis vinifera= a review" American Journal of Enology and Viticulture 60=411-434 Guilpart et al. (2014) "Grapevine bud fertility and number of berries per bunch are determined by water and nitrogen stress around flowering in the previous year" European Journal of Agronomy 54=9-20.
Reproductive Active Bud
Reproductive success in grapevine varies from 15-50% fruit set depending on cultivar, with Chardonnay achieving 35-45% berry set under optimal conditions while Merlot typically reaches only 20-30%. The transition from vegetative to reproductive growth involves 2-3 season lead time, with inflorescence primordia initiating in latent buds during the previous growing season. Reproductive allocation (berry dry weight / total above-ground dry weight) ranges from 25-45% in commercially managed vineyards.
A grapevine disposition that is the inherited capacity of a Vitis vinifera plant to undergo sexual reproduction through the coordinated development of flowers, pollination, fertilization, and fruit set, encompassing all traits related to inflorescence formation, gametophyte development, and the transition from flower to berry.
Reproductive_Disposition serves as the top-level parent class for all traits related to grapevine sexual reproduction. The category encompasses flowering biology, fruit set mechanisms, seedlessness traits, fruitfulness, and cluster architecture. Grapevine reproductive biology is complicated by the two-year developmental cycle where inflorescence primordia form in year N-1 but flowers open in year N.
Fortpflanzungsfähigkeit
Reproduktionsvermögen
generative Disposition
generative disposition
reproductive capacity
sexual reproduction trait
Carmona MJ et al. (2008) A molecular genetic perspective of reproductive development in grapevine. Journal of Experimental Botany 59:2579-2596 Mullins MG, Bouquet A, Williams LE (1992) Biology of the Grapevine, Cambridge University Press pp.: 80-117
Reproductive Disposition
A reproductive dominant mixed bud on old-vine Vitis vinifera cv. Grenache in a hillside vineyard in Châteauneuf-du-Pape, France, developing into a shoot with 2 large inflorescences bearing 165 and 148 berries respectively, only 7 mature leaves totaling 380 cm² leaf area, shoot length 35cm with short 3-4cm internodes, leaf area-to-fruit ratio of only 6.1 cm²/g fruit at harvest, showing incomplete anthocyanin development and suboptimal sugar accumulation despite extended hang time, requiring cluster thinning to one cluster per shoot to achieve adequate fruit maturity.
A mixed active bud developing into a shoot with predominant reproductive structures relative to vegetative growth, characterized by multiple or large inflorescences (2-3 clusters per shoot or clusters exceeding 400g at maturity), limited shoot elongation, reduced leaf area development (typically 6-10 mature leaves), short internodes, and early growth cessation. Reproductive dominant mixed buds typically arise from low-vigor sites with shallow or nutrient-poor soils, restricted water availability, older vines with limited root system vigor, or overcropped vines, producing shoots with inadequate leaf area-to-fruit ratios (often below 8 cm²/g fruit) resulting in incomplete fruit ripening, depletion of vine carbohydrate reserves, reduced cold hardiness, and diminished bud fruitfulness for subsequent seasons due to insufficient photosynthate production during critical inflorescence differentiation period.
Reproductive dominant mixed buds indicate vine stress or imbalance requiring corrective management to restore sustainability, including crop load reduction through cluster thinning (typically to one cluster per shoot or less), irrigation supplementation where feasible, nitrogen fertilization if deficiency indicated, and possible reduction in vine density or conversion to spur pruning to reduce crop potential. Chronic reproductive dominance leads to progressive vine decline through carbohydrate depletion, winter damage due to poor cold acclimation, increased susceptibility to trunk diseases, and eventual vine death. Short-term high quality may result from extreme crop limitation, but long-term vineyard sustainability requires balanced vegetative-reproductive development.
crop-heavy mixed bud
reproductive-biased mixed bud
reproduktiv-dominante gemischte Knospe
Based on Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Petrie et al. (2000) "The effect of leaf removal and canopy height on whole-vine gas exchange and fruit development of Vitis vinifera L. Sauvignon Blanc" Functional Plant Biology 27=569-578 Bravdo et al. (1985) "Effect of crop level and crop load on growth, yield, must and wine composition, and quality of Cabernet Sauvignon" American Journal of Enology and Viticulture 36=125-131.
Reproductive Dominant Mixed Bud
A reproductive shoot on Vitis vinifera cv. Merlot from node 8 on bilateral cordon, developed from primary bud at BBCH 09 (leaf development advanced), currently mid-July at BBCH 75 (berries pea-sized), 142cm length with 16 fully expanded leaves providing approximately 2240 cm² photosynthetic area, 2 inflorescences with successful fruit set showing 44 and 38 developing berries respectively, 2 tendrils, shoot positioned vertically in VSP trellis, representing typical productive shoot in well-managed vineyard contributing both current crop (estimated 820g fruit at maturity from two clusters) and substantial photosynthetic capacity (adequate 27 cm² leaf area per gram projected fruit supporting complete maturation), managed through standard canopy operations including shoot positioning, hedging to 165cm final height, and basal leaf removal (4 leaves) improving fruit zone air circulation and light exposure.
A shoot bearing one or more inflorescences, characterized by nodes carrying flower clusters in addition to vegetative structures (leaves, tendrils), developmental origin from reproductive buds with inflorescence primordia differentiated during previous growing season, and dual function providing both current-season crop production through fruit development and photosynthetic capacity through leaf area supporting fruit maturation and whole-vine carbon balance. Reproductive shoots represent the primary production units in viticulture, with their number, vigor, cluster count, and canopy characteristics fundamentally determining yield quantity and quality, requiring careful management through shoot thinning (adjusting shoot density), cluster thinning (modifying crop load), canopy manipulation (optimizing light exposure and air circulation), and various interventions balancing productive goals with vine sustainability and fruit quality objectives across diverse varieties, sites, and production philosophies.
Reproductive shoots are distinguished from vegetative shoots (no inflorescences) by presence of flower clusters, with reproductive designation applying regardless of cluster count (single or multiple), cluster size, or fruit set success. The reproductive function creates resource demands beyond vegetative shoots, with developing fruit acting as strong sinks drawing carbohydrates, water, and nutrients, requiring adequate leaf area support (typically 10-15 cm² per gram fruit minimum, 12-18 cm² optimal) and appropriate shoot vigor balancing productive capacity with canopy quality. Management focuses on optimizing reproductive shoot number (through winter pruning bud retention and spring shoot thinning), cluster load per shoot (through cluster thinning achieving target yields and balanced leaf-to-fruit ratios), and canopy architecture (through positioning, hedging, leaf removal) supporting fruit development while maintaining vine health and enabling consistent quality production across seasons.
Ertragstrieb
Fruchttrieb
crop-bearing shoot
fertile shoot
fertiler Trieb
flowering shoot
fruiting shoot
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 146-198 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide.
Reproductive Shoot
Pinot Noir accession Gm 20-13 at the Geisenheim University research vineyard bearing a research and education role is maintained as part of a 450-cultivar teaching and research collection planted in standardized 24-vine plots at 1.5 × 2.0 m spacing, used simultaneously for ampelography courses with 120 viticulture students annually, phenological monitoring providing 35+ years of continuous BBCH-stage records, and clonal comparison trials evaluating 15 Pinot Noir clones for Rheingau-specific performance metrics including must weight, acidity, and Botrytis susceptibility.
A grapevine role that inheres in a vine by virtue of its designated function within scientific research programs, academic teaching collections, or reference documentation systems, where the vine serves as an object of systematic study, a pedagogical resource, or a standard of comparison rather than — or in addition to — a source of commercial fruit production.
Research_And_Education_Role classifies grapevines according to their epistemic and pedagogical function within the knowledge infrastructure of viticulture. The role is independent of and compatible with other role classifications: a vine may simultaneously bear a Research_And_Education_Role and a Premium_Wine_Role if its fruit is commercially vinified while the vine also serves documented research or teaching purposes. Subclasses distinguish between reference functions (providing standards for identification and comparison), experimental functions (serving as subjects of active investigation), teaching collections, ampelographic documentation, genetic reference, and germplasm conservation.
Forschungs- und Bildungs-Rolle
Lehr- und Versuchsweinberg-Rolle
educational vineyard role
research vine role
scientific collection role
wissenschaftliche Sammlungs-Rolle
OIV (2009) 2nd Edition of the OIV Descriptor List for Grape Varieties and Vitis Species Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157 This P et al. (2004) Development of a standard set of microsatellite reference alleles for identification of grape cultivars, Theoretical and Applied Genetics 109: 1448-1458
Research And Education Role
A Vitis sylvestris accession from the Caucasus preserved in the INRA Vassal collection bears a Research_Clone_Role, maintained for genetic diversity research and potential trait identification without commercial cultivation intent. A leafroll-sensitive Pinot Noir selection maintained at FPS serves in a Research_Clone_Role as an indicator plant for virus indexing, its high susceptibility making it valuable for pathogen detection despite this trait precluding commercial planting. An ancient variety from the Iberian Peninsula preserved at IMIDRA (Madrid) bears a Research_Clone_Role for ampelographic study and genetic resource conservation, with no current pathway toward commercial rehabilitation.
An experimental clone role that inheres in a grapevine genotype maintained primarily for research purposes without current intent for commercial registration or distribution, serving scientific investigation, germplasm conservation, or reference functions within research collections. Research clones encompass diverse material—ancient varieties preserved for genetic study, wild species accessions for diversity research, disease-susceptible standards for pathology investigations, distinctive phenotypes for developmental biology research, and historical selections maintained for comparison without commercial relevance. Research clone collections exist at viticultural research institutions worldwide (INRA Vassal, JKI, UC Davis, CSIRO, IMIDRA), preserving genetic diversity beyond the commercially registered clone inventory. Research clones may lack the health certification, field trial documentation, or administrative registration required for commercial release, with their value residing in scientific rather than commercial applications. Access to research clones typically requires research collaboration agreements, with material transfer limited to legitimate research purposes rather than commercial propagation.
Research_Clone_Role captures the germplasm conservation and scientific investigation functions distinct from commercial clone development, recognizing that substantial viticultural genetic resources exist outside the commercially registered clone inventory. Research collections preserve diversity at risk of loss through commercial rationalization—ancient varieties, regional specialties, wild relatives—representing a genetic reservoir for future breeding and adaptation. Research clone management balances accessibility (enabling research collaboration) with conservation (preventing genetic erosion through uncontrolled distribution) and biosecurity (managing pathogen risks in diverse collections). The distinction from Breeding_Line reflects intent research clones are maintained for study rather than commercial development, though discoveries may occasionally prompt transition toward breeding applications or commercial evaluation.
Forschungsklon
Genbank-Klon
Referenzklon
Sammlungsklon
collection clone
germplasm clone
reference clone
research clone
This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Lacombe T et al. (2011) Grapevine European catalogue and genetic diversity, in Bentley R (ed) The Handbook of Grape Processing, Blackwell INRA (2020) Domaine de Vassal Grape Collection Maul E et al. (2012) Vitis International Variety Catalogue, JKI
Research Clone Role
Reserve accumulation occurs primarily during the post-harvest period (4-8 weeks between harvest and leaf fall), with starch concentration in canes and trunks increasing from 8-12% to 18-28% of dry weight during this period in well-managed vines. Cultivars differ in reserve accumulation capacity, with Pinot Noir and Riesling showing typically higher reserve levels (22-28% starch) than vigorous cultivars such as Thompson Seedless (15-20%). Reserve distribution within the vine shows highest concentrations in roots (20-30%) and cane bases (18-25%), with declining levels toward shoot tips. Nitrogen reserves (arginine predominating, comprising 40-60% of root free amino acids) accumulate to 1.5-3.0% of root dry weight during autumn. Spring mobilization depletes 40-60% of reserves by bloom, with reserves not fully restored until mid-summer in heavy crop years. Insufficient reserves result in weak spring growth, poor fruit set, and increased susceptibility to stress.
A cane maturation disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and store carbohydrate reserves (primarily starch) and nitrogen reserves (amino acids, proteins) in perennial woody tissues during late season, providing the energy and nutrient pools required for winter cold hardiness maintenance, spring growth resumption, and early-season development before photosynthesis becomes self-supporting.
Reserve accumulation represents the physiological foundation for vine perenniality, bridging the dormant season and fueling the burst of spring growth before current-season photosynthesis becomes established. The trait integrates source capacity (post-harvest photosynthesis), sink activity (phloem loading into storage tissues), and the enzymatic machinery for starch synthesis (ADP-glucose pyrophosphorylase, starch synthase). Management practices affecting reserve status (harvest timing, post-harvest irrigation, defoliation) have carry-over effects on following-season performance.
Kohlenhydratspeicherung
Reservestoffeinlagerung
Stärkeakkumulation
carbohydrate storage
reserve deposition
starch accumulation trait
Holzapfel BP et al. (2010) Accumulation and distribution of carbohydrate and nitrogen in grapevines under a Mediterranean climate. Australian Journal of Grape and Wine Research 16:53-63 Zapata C et al. (2004) Effects of girdling on carbohydrate and nitrogen partitioning in grapevine. Journal of Experimental Botany 55:579-592
Reserve Accumulation Disposition
A reserve renewal cane on Vitis vinifera cv. Syrah in cane-pruned VSP system, retained at 8 nodes positioned 25cm from cordon (compared to primary replacement at 10cm and secondary at 18cm), measuring 7.5mm diameter at lower end of acceptable range, complete lignification but slightly longer internodes indicating more vigorous growth than ideal, representing contingency option unlikely to be used for fruiting cane unless both primary and secondary replacements fail due to winter damage, poor spring development, or unexpected positioning problems, with typical outcome being complete removal during following dormant pruning if preferred renewals prove satisfactory, but providing valuable insurance against renewal system breakdown.
A renewal cane retained as longer-term insurance against renewal system failures, characterized by less immediate replacement expectation compared to standard replacement renewals, positioning or quality making it third-choice or contingency option behind primary and secondary replacements, and function as reserve capacity for unusual circumstances including catastrophic damage to preferred renewals, training system modifications requiring additional wood, or multi-year planning anticipating future needs beyond immediate next-season replacement. Reserve renewal canes provide additional safety margin in renewal planning beyond standard replacement strategies, acknowledging that occasional severe problems (major winter damage, spring frost destroying preferred renewals, disease affecting primary wood) may require utilization of reserve options normally not needed in standard annual cycles but valuable for maintaining system resilience.
Reserve renewal canes represent conservative renewal planning accepting modest bud count sacrifice (retaining extra renewals consuming bud capacity that could support additional fruiting positions) for increased system resilience against various failure modes. The reserve designation implies low probability of actual use in standard seasons (perhaps 5-15% utilization versus 80-90% for primaries), but high value in problem years when reserve capacity prevents complete renewal system failure requiring crisis-mode latent bud activation or dramatic training system modifications. Economic viability of reserve renewal strategies depends on failure frequency and consequences, with high-value vineyards in challenging climates justifying reserve capacity while low-value bulk production may accept higher renewal risk through minimal redundancy.
Notfall-Erneuerungsrute
Reserve-Vorratsrute
backup reserve cane
contingency renewal cane
insurance renewal wood
Reserve Renewal Cane
Residual sugar for Riesling Spätlese from the Rheingau (2024 vintage) measures 42 g/L by enzymatic assay (glucose 18 g/L, fructose 24 g/L), classifying the wine as "lieblich" (medium-sweet) under German labeling regulations and positioning it in the traditional Spätlese sweetness range where the 8.2 g/L total acidity provides the structural counterbalance essential for the characteristic Riesling sweet-acid tension. Residual sugar for Grüner Veltliner Smaragd from the Wachau (2024 vintage) records 2.8 g/L, below the 4 g/L "trocken" (dry) threshold under Austrian regulations when TA exceeds RS by at least 2 g/L (TA 5.8 g/L meets this condition), confirming the dry classification. Residual sugar for Amarone della Valpolicella (2021 vintage) documents 5.5 g/L — technically above the dry threshold in isolation, but perceived as dry due to the 16.0% alcohol and 5.8 g/L TA masking the residual sweetness, illustrating the interaction between RS, alcohol, and acidity in sweetness perception.
Residual sugar is a wine analysis data entity that records the concentration of unfermented reducing sugars (glucose and fructose) remaining in finished wine after the completion or arrest of alcoholic fermentation, determined by enzymatic assay (glucose oxidase/hexokinase), Fehling titration, or FTIR estimation, expressed in grams per liter (g/L), and serving as the primary determinant of wine sweetness classification (dry, off-dry, medium-sweet, sweet), a critical parameter for microbial stability assessment (risk of refermentation by residual yeast or spoilage organisms), and a key regulatory metric that defines quality wine categories and controls labeling declarations in most wine-producing jurisdictions.
Residual sugar determination requires careful analytical distinction from total sugars, as wine may contain non-fermentable sugars (pentoses — arabinose, xylose, and rhamnose, typically 0.5-1.5 g/L) that contribute to reducing sugar assays but are not metabolized by Saccharomyces cerevisiae and do not contribute meaningfully to sweetness perception. The enzymatic method using glucose oxidase (for glucose) and hexokinase with phosphoglucose isomerase (for fructose) provides specific quantification of each fermentable sugar, while the Fehling reduction method measures total reducing power and therefore overestimates fermentable residual sugar. The sweetness classification system varies between jurisdictions but follows a general hierarchy — EU regulations define "dry" (trocken/sec) as ≤4 g/L or ≤9 g/L when TA is within 2 g/L of RS, "medium-dry" (halbtrocken/demi-sec) as ≤12 g/L or ≤18 g/L under the acidity exception, "medium-sweet" (lieblich/moelleux) as 12-45 g/L, and "sweet" (süß/doux) as >45 g/L. The fructose-to-glucose ratio in residual sugar carries sensory significance because fructose is perceived as approximately 1.7× sweeter than glucose on a weight basis — wines where fermentation preferentially consumed glucose (the natural yeast preference) leave fructose-enriched residual sugar that tastes sweeter per gram than wines with balanced glucose-fructose residual. Microbial stability management for wines with significant residual sugar (>5 g/L) requires either sterile filtration (0.45 µm membrane), sulfur dioxide and sorbic acid preservation, or pasteurization to prevent refermentation by residual yeast or contaminating organisms, making RS measurement a prerequisite for bottling line sterility decisions. The increasing consumer preference for dry-tasting wines has driven many producers to reduce residual sugar levels, though some commercial wines marketed as "dry" contain 5-10 g/L RS as a deliberate stylistic choice to enhance perceived body and reduce acid aggressiveness.
Restsüße
Restzucker
Restzuckergehalt
Zuckerrest
reducing sugar
residual glucose and fructose
sugar content of wine
unfermented sugar
OIV (2021) Compendium of International Methods of Analysis, Method OIV-MA-AS311-01A (reducing substances) European Parliament (2019) Commission Delegated Regulation (EU) 2019/33, Annex III (sweetness labeling) Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2, 2nd Ed, Wiley, pp.: 61-75
Residual Sugar
Muscadinia rotundifolia serves in a Resistance_Donor_Role as the source of the Run1 locus conferring complete powdery mildew resistance, introgressed into V. vinifera through complex interspecific hybridization and successive backcrossing in French and international breeding programs. Vitis amurensis bears a Resistance_Donor_Role for the Rpv10 downy mildew resistance locus in German breeding, with marker-assisted selection tracking resistance introgression through BC generations. A "bridge genotype" like Regent functions as intermediate resistance donor, carrying already-introgressed R-genes from wild species in a partially improved genetic background more readily crossed with V. vinifera than original wild species donors.
A donor parent role that inheres in a grapevine genotype by virtue of its function as the source of disease or pest resistance genes targeted for introgression into susceptible V. vinifera backgrounds. Resistance donors are predominantly wild Vitis species and their derivatives that evolved resistance mechanisms during long coevolution with pathogens and pests in their native ranges—resistance largely absent from V. vinifera which evolved in geographic separation from major grapevine pathogens. Major resistance donor sources include Muscadinia rotundifolia (powdery mildew resistance via Run1, Ren1 loci), Vitis amurensis (downy mildew resistance via Rpv loci, cold hardiness), Vitis cinerea (Pierce's disease resistance via PdR loci), and various American Vitis species (phylloxera tolerance foundational to rootstock breeding). Resistance donors are characterized by the specific R-genes they carry, their inheritance pattern (dominant, recessive, quantitative), and their "stacking" compatibility with other resistance sources for durable multigene resistance. Marker-assisted selection has transformed resistance breeding by enabling precise introgression of mapped R-genes without requiring pathogen exposure for phenotypic selection.
Resistance_Donor_Role captures the specific resistance-source function that has driven most contemporary grapevine breeding investment. The need for resistance donors reflects V. vinifera's susceptibility to pathogens (powdery mildew, downy mildew, Pierce's disease) that evolved outside its native range, with resistance mechanisms present in American and Asian Vitis species that coevolved with these pathogens. Resistance breeding has progressed from phenotypic selection (requiring disease exposure) to marker-assisted selection (molecular confirmation of R-gene presence), dramatically improving efficiency and enabling R-gene "pyramiding" (combining multiple resistance loci for durable resistance). The "resistance donor" framing acknowledges that wild species donors typically carry undesirable characteristics alongside valuable resistance, necessitating extensive backcrossing to recover elite wine grape character while retaining resistance. Second-generation resistance breeding uses improved intermediate donors (registered PIWI varieties) rather than raw wild species, facilitating more rapid quality recovery.
Krankheitsresistenzquelle
R-Gen-Quelle
R-gene source
Resistenzelter
Resistenzspender
disease resistance source
resistance donor
resistance parent
Dry IB et al. (2010) Molecular strategies to enhance disease resistance in grapevine, Australian Journal of Grape and Wine Research 16: 94-105 Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330 Töpfer R et al. (2011) Molecular breeding in grapes, CRC Press pp.: 160-185 Riaz S et al. (2011) Identification of powdery mildew resistance genes, Theoretical and Applied Genetics 122: 1059-1073
Resistance Donor Role
The resistance gene information for the interspecific hybrid cultivar Regent (Diana × Chambourcin, registered 1996) documents a pyramided resistance profile comprising Rpv3-1 on chromosome 18 (conferring partial resistance to Plasmopara viticola downy mildew, derived from North American Vitis species through the Chambourcin lineage), Ren3 on chromosome 15 (conferring partial resistance to Erysiphe necator powdery mildew), and Ren9 on chromosome 15 (providing additional powdery mildew resistance) — this multi-locus resistance enables Regent cultivation with 50-70% fewer fungicide applications than susceptible V. vinifera cultivars in German viticultural conditions, though the Rpv3-1 resistance has been partially overcome by virulent P. viticola isolates in some European regions since 2010, demonstrating the evolutionary capacity of the pathogen to adapt to deployed resistance genes and underscoring the importance of resistance gene pyramiding strategies in breeding programs.
A genetic information entity that documents the presence, identity, genomic location, and functional characterization of disease resistance genes and quantitative resistance loci in Vitis genotypes, encompassing the major gene-for-gene resistance loci (R-genes) and quantitative trait loci (QTLs) conferring resistance to fungal, oomycete, bacterial, and viral pathogens, with resistance gene information serving as the critical genetic data for marker-assisted selection in breeding programs developing disease-resistant cultivars (PIWI varieties) that reduce pesticide dependency in viticulture.
Resistance_Gene_Information addresses the genetic basis of disease resistance that is arguably the most important frontier in contemporary grapevine breeding. The susceptibility of Vitis vinifera to the major fungal and oomycete diseases — downy mildew (Plasmopara viticola), powdery mildew (Erysiphe necator), and to a lesser extent black rot (Guignardia bidwellii) and anthracnose (Elsinoe ampelina) — reflects the geographic separation between V. vinifera (which evolved in Eurasia without these pathogens) and the American Vitis species (which co-evolved with these pathogens in North America and developed effective resistance mechanisms): the introduction of these pathogens to European vineyards in the mid-19th century (downy mildew 1878, powdery mildew 1845, phylloxera 1863) devastated European viticulture and initiated both the chemical crop protection paradigm still dominant today and the interspecific hybridization breeding programs that aim to introgress resistance from American species into V. vinifera genetic backgrounds. Resistance loci are designated by standardized nomenclature — Rpv (Resistance to Plasmopara viticola) for downy mildew, Ren (Resistance to Erysiphe necator) for powdery mildew, Run (Resistance from Vitis rotundifolia to Uncinula necator) for muscadine-derived powdery mildew resistance, and Rgb (Resistance to Guignardia bidwellii) for black rot.
Krankheitsresistenz-Locus-Daten
Pathogenresistenz-Genetik
R-gene characterization
Resistenzgen-Information
disease resistance locus data
pathogen resistance genetic data
Töpfer R et al. (2011) Molecular breeding, in Adam-Blondon AF et al. (eds) Genetics, Genomics, and Breeding of Grapes, Science Publishers pp: 160-185 Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance and ripening of grapevines assessed in a segregating population, Molecular Breeding 36: 65 VIVC (2023) Loci for resistance to diseases and pests — standardized nomenclature Dry IB et al. (2010) Grapevine disease resistance genes and their use in breeding, in Reynolds AG (ed) Viticulture and Wine Quality, Woodhead Publishing pp: 193-228
Resistance Gene Information
The resistance gene pyramid data for the advanced breeding selection GF.2018-047 at the Julius Kühn-Institut documents a four-locus resistance stack — Rpv3-1 (chromosome 18, from North American Vitis, downy mildew) + Rpv12 (chromosome 14, from V. amurensis, downy mildew) + Ren3/Ren9 (chromosome 15, from North American Vitis, powdery mildew) + Ren1 (chromosome 13, from V. vinifera Kishmish Vatkana, powdery mildew) — confirmed by marker-assisted selection at 12 diagnostic locus-specific markers across the four chromosomal regions. Controlled inoculation assays showed near-complete resistance to both pathogens (>95% reduction in both P. viticola and E. necator severity compared to susceptible controls), significantly exceeding the resistance conferred by any individual locus, with multi-site field trials over 3 years confirming commercially acceptable disease control with zero fungicide applications in moderate-pressure seasons and 1-2 applications in high-pressure seasons.
A resistance gene information record that documents the specific combination of multiple resistance loci assembled in a single Vitis genotype through deliberate breeding, specifying the identity, chromosomal locations, and source species of each stacked resistance gene, the anticipated additive or synergistic resistance effect of the combination, and the marker-assisted selection strategy used to confirm the presence of all targeted loci, providing the genetic blueprint for durable multi-disease resistance in new cultivar development.
Resistance_Gene_Pyramid_Data documents the cutting-edge breeding strategy that represents the current best approach to achieving durable disease resistance in grapevine — the systematic combination of multiple independently acting resistance loci from diverse genetic sources into single elite genotypes. The pyramiding strategy is grounded in evolutionary genetics theory predicting that the probability of a pathogen simultaneously overcoming n independent resistance mechanisms is approximately p^n (where p is the probability of overcoming a single mechanism), making two-locus pyramids approximately p-fold more durable than single loci and three-locus pyramids p²-fold more durable: for P. viticola with an estimated single-locus breakdown probability of p ≈ 0.1 per decade under moderate selection pressure, a three-locus pyramid (e.g., Rpv3 + Rpv10 + Rpv12) would have a predicted durability of >100 years — effectively permanent in commercial terms. The practical implementation of pyramiding requires marker-assisted selection because resistance phenotypes of individual loci cannot be reliably distinguished in a genotype carrying multiple resistance genes (the combined resistance masks the individual contributions), making molecular markers essential for confirming the presence of each targeted locus in selected individuals. Current European PIWI breeding programs are converging on a standard pyramiding framework combining 2-3 Rpv loci (for downy mildew) with 2-3 Ren/Run loci (for powdery mildew) to create cultivars with durable dual-disease resistance — the first such multi-pyramid cultivars are expected to reach commercial registration between 2025 and 2035.
Multi-Locus-Resistenzprofil
Resistenz-Stapelung-Dokumentation
Resistenzgen-Pyramide-Daten
multi-locus resistance profile
resistance pyramid specification
stacked resistance documentation
Töpfer R et al. (2011) Molecular breeding, in Adam-Blondon AF et al. (eds) Genetics, Genomics, and Breeding of Grapes, Science Publishers pp: 160-185 Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance, Molecular Breeding 36: 65 Dry IB et al. (2010) Grapevine disease resistance genes, in Reynolds AG (ed) Viticulture and Wine Quality, Woodhead Publishing pp: 193-228 Merdinoglu D et al. (2018) Breeding for durable resistance to downy and powdery mildew in grapevine, OENO One 52: 203-209
Resistance Gene Pyramid Data
Respiration acclimation assessment in Chardonnay comparing vines grown at 20°C versus 30°C day temperature reveals the homeostatic adjustment, with warm-grown leaves showing respiration of 2.1 μmol CO₂/m²/s when measured at 25°C while cool-grown leaves show 2.8 μmol CO₂/m²/s at the same measurement temperature, indicating 25% acclimation-driven reduction in respiratory capacity that partially compensates for the increased respiratory demand under warm conditions, with the acclimation associated with reduced mitochondrial density (15% lower) and altered cytochrome/alternative oxidase balance in warm-acclimated tissues.
Respiration acclimation quality is a temperature response quality that represents the capacity of grapevine respiratory metabolism to adjust its temperature response characteristics following sustained exposure to altered thermal conditions, characterizing the physiological plasticity that enables partial homeostasis of respiratory carbon losses across the temperature environments experienced in different growing seasons, regions, or under climate change. Within the BFO framework, this quality specifically addresses the adaptive plasticity dimension of respiratory temperature response, distinguishing acclimation (phenotypic adjustment within an individual) from adaptation (genetic change across generations) while recognizing that respiratory acclimation provides a mechanism for maintaining carbon balance efficiency when growth temperatures differ from the thermal environment to which the respiratory apparatus was initially optimized. The ontological significance lies in respiration acclimation moderating the respiratory burden under altered thermal conditions, where acclimation to warmer temperatures typically reduces respiratory rate at any given measurement temperature compared to cold-acclimated tissues, partially compensating for the increased respiratory demand that would otherwise occur under warming and improving carbon balance under elevated temperature conditions. The physiological basis for respiration acclimation includes changes in mitochondrial density (reduced under warm acclimation), respiratory enzyme capacity (altered expression and activity), membrane composition (modified lipid saturation affecting electron transport chain efficiency), and the balance between cytochrome and alternative oxidase pathways (shifting with acclimation temperature). The quantitative expression of respiratory acclimation compares respiration rates of tissues grown at different temperatures when measured at a common reference temperature, with full acclimation showing similar rates regardless of growth temperature (Type II acclimation) while partial acclimation shows intermediate adjustment, and the degree of acclimation varying among species, tissues, and the magnitude and duration of temperature change. The time course of respiratory acclimation typically shows detectable changes within days of temperature shift, with more complete acclimation over weeks as new tissue develops under the altered temperature regime, and with the acclimation capacity potentially declining in mature tissues with limited metabolic plasticity. Understanding respiration acclimation quality enables prediction of respiratory responses under climate change scenarios, assessment of cultivar plasticity for thermal adaptation, and appreciation of the physiological mechanisms that buffer carbon balance against temperature variation.
Respiration_Acclimation_Quality addresses the phenotypic plasticity of respiratory temperature response. This quality enables partial homeostasis of carbon losses under altered thermal conditions through physiological adjustment. The class supports climate adaptation assessment and carbon balance prediction through characterization of respiratory acclimation capacity.
Atmungsanpassung
Respirationsakklimation
metabolic adjustment
metabolische Kompensation
respiratory homeostasis
temperature compensation
thermal acclimation
thermische Akklimatisation
Atkin OK, Tjoelker MG (2003) Thermal acclimation and the dynamic response of plant respiration to temperature. Trends in Plant Science 8(7): 343-351 Slot M, Kitajima K (2015) General patterns of acclimation of leaf respiration to elevated temperatures across biomes and plant types. Oecologia 177(3): 885-900
Respiration Acclimation Quality
Respiration rate assessment in Cabernet Sauvignon integrates component fluxes across organs, with leaf dark respiration of 1.8 μmol CO₂/m²/s (maintenance-dominated in mature leaves), root respiration consuming 25% of daily photosynthate (maintenance plus ion uptake costs), fruit respiration peaking at 12 mg CO₂/kg/hour during rapid growth then declining, and whole-plant respiration consuming approximately 52% of gross photosynthesis, demonstrating the substantial carbon cost of maintaining and growing vine tissues that determines the 48% of fixed carbon available for structural growth, fruit sugar accumulation, and reserve storage.
Respiration rate quality is a physiological quality that represents the rate of oxidative catabolism of carbohydrates and other substrates to generate ATP and carbon skeletons for biosynthesis, characterizing the metabolic CO₂ release that constitutes the carbon cost of maintaining and growing grapevine tissues and determines the net carbon balance when integrated with photosynthetic carbon gain. Within the BFO framework, this quality addresses the carbon loss dimension of vine metabolism, distinguishing respiratory CO₂ release from photosynthetic CO₂ fixation while recognizing that respiration provides the essential energy and biosynthetic intermediates for all cellular processes but also represents a substantial carbon cost that reduces net productivity. The ontological significance lies in respiration rate determining the carbon efficiency of the grapevine, where the ratio of carbon respired to carbon fixed establishes the fraction of photosynthate retained for growth, fruit development, and reserve storage versus that lost as respiratory CO₂, with whole-plant respiration typically consuming 40-60% of gross photosynthesis and thus critically affecting net carbon gain and potential productivity. The biochemical basis for respiration involves glycolysis, the tricarboxylic acid cycle, and oxidative phosphorylation, with glucose oxidation yielding CO₂, H₂O, and ATP (approximately 36-38 ATP per glucose), and with the pathways providing both energy (ATP) and carbon skeletons (organic acid intermediates) for biosynthesis of amino acids, lipids, and other compounds. The functional components of respiration include maintenance respiration (energy for cellular homeostasis, protein turnover, membrane integrity), growth respiration (energy and carbon skeletons for new tissue synthesis), and in photosynthetic tissues, photorespiration (light-dependent oxidative cycle distinct from mitochondrial respiration), with the relative contribution of these components varying with tissue type, developmental stage, and environmental conditions. The environmental and physiological factors affecting respiration rate include temperature (strong exponential response, Q₁₀ typically 2.0-2.5), substrate availability (carbohydrate supply limiting respiration when depleted), tissue nitrogen content (maintenance costs proportional to protein content), growth rate (determining growth respiration demand), and stress conditions (often affecting respiration through multiple mechanisms). Understanding respiration rate quality enables carbon balance assessment, growth efficiency evaluation, and appreciation of the metabolic costs that determine net vine productivity and fruit quality potential.
Respiration_Rate_Quality serves as the parent class for respiration components (Maintenance, Growth, Dark, Photorespiration) and Respiratory Quotient. This quality represents the carbon cost side of vine metabolism that determines net productivity when balanced against photosynthetic carbon gain. The class supports carbon balance assessment and efficiency evaluation through comprehensive respiration characterization.
Atmungsrate
CO₂ release rate
CO₂-Freisetzung
Respirationsrate
catabolic rate
oxidative metabolism
oxidativer Stoffwechsel
respiratory metabolism
Amthor JS (2000) The McCree–de Wit–Penning de Vries–Thornley respiration paradigms: 30 years later. Annals of Botany 86(1): 1-20 Escalona JM et al. (1999) Stomatal and non-stomatal limitations of photosynthesis under water stress in field-grown grapevines. Australian Journal of Plant Physiology 26(5): 421-433
Respiration Rate Quality
In a Barossa Valley Grenache vineyard, respiratory malate consumption was quantified over 42 days from late February through April Year N+1 (BBCH 84-87) when malate declined from 8.6 g/L to 1.8 g/L (total loss 6.8 g/L, 79%), showing peak consumption 0.42 g/L per day days 15-28 (daily high 30-33°C), declining to 0.15 g/L per day late phase (daily high 24-27°C as autumn progressed), berry respiration averaged 1.85 μmol O₂ g⁻¹ h⁻¹ with estimated 52% CO₂ from malate mid-period declining to 28% late as substrate depleted.
Respiratory_Malate_Consumption represents the utilization of malic acid as respiratory substrate through oxidative catabolism producing CO₂, reducing equivalents, and ATP creating the primary mechanism for malate depletion during berry ripening while simultaneously supporting energy demands of ripening processes including sugar transport, biosynthesis, and cellular maintenance. This consumption intensifies during veraison through ripening (BBCH 83-87), beginning 2-5 days post-degradation onset and continuing until malate substantially depleted, and is characterized by malate oxidation rates 0.15-0.45 g/L per day (1.5-4.5 μmol g⁻¹ h⁻¹) during peak consumption representing 35-65% of total berry respiratory CO₂ production, energy yield from malate providing estimated 30-55% of berry ATP production during active consumption phase supporting ripening metabolism, and malate decline from early consumption 7-12 g/L to late phase 2-6 g/L approaching final cultivar-specific minima 1-5 g/L at harvest. Biochemical pathways of respiratory consumption inclu direct oxidative decarboxylation via NADP-malic enzyme producing pyruvate that enters mitochondria for complete oxidation through pyruvate dehydrogenase (pyruvate + CoA + NAD⁺ → acetyl-CoA + CO₂ + NADH) then citric acid cycle (acetyl-CoA → 2 CO₂ + 3 NADH + 1 FADH₂ + 1 GTP) generating reducing equivalents, complete oxidation pathway through citric acid cycle where malate dehydrogenase converts malate to oxaloacetate (coupled to NADH production) then citrate synthase condenses with acetyl-CoA producing citrate beginning full cycle producing 4 NADH + 1 FADH₂ + 1 GTP per malate, and respiratory chain oxidizing NADH and FADH₂ through Complexes I-IV generating proton gradient driving ATP synthase producing ~15 ATP per malate completely oxidized representing efficient energy extraction. Quantification of respiratory malate consumption employs: isotope tracer studies using ¹³C-labeled malate showing label incorporation into respiratory CO₂ revealing 40-70% of evolved CO₂ derived from malate during peak consumption, respiratory quotient (RQ) measurements showing values 1.08-1.35 during ripening compared to 0.95-1.05 from pure hexose oxidation indicating substantial organic acid contribution, malate concentration decline rates 0.15-0.45 g/L per day (molecular weight 134 g/mol = 1.1-3.4 mmol per kg per day) correlating with CO₂ production providing independent consumption estimate, and respiratory O₂ consumption 1.2-2.8 μmol g⁻¹ h⁻¹ with portion attributable to malate oxidation calculated from RQ and substrate composition. Temporal dynamics of consumption show: early phase (BBCH 83-84, days 0-15 post-onset) with moderate consumption 0.18-0.32 g/L per day as malate remains abundant 7-12 g/L, peak phase (BBCH 84-85, days 15-35) with maximum consumption 0.25-0.50 g/L per day under favorable conditions particularly warm temperatures, declining phase (BBCH 85-87, days 35-55) with slowing consumption 0.10-0.25 g/L per day as malate depletes to 2-6 g/L reducing substrate availability, and late phase approaching harvest with minimal consumption 32-35°C causing excessive consumption 0.45-0.70 g/L per day creating risk of premature total depletion, cool conditions 18-24°C moderating consumption to 0.15-0.28 g/L per day enabling acid retention, and cumulative degree-days from veraison strongly predicting total malate loss with correlation r² = 0.75-0.90 across seasons and sites. Diurnal patterns of consumption reveal: daytime consumption elevated 30-60% above nighttime due to higher berry temperature from solar heating creating midday peaks 0.30-0.60 g/L per day versus nighttime troughs 0.12-0.28 g/L per day, cumulative 24-hour consumption integrating day-night cycles determining total daily loss typically 0.20-0.40 g/L in moderate climates, and regions with extreme diurnal temperature range (>15-20°C) showing substantial day-night variation potentially enabling acid preservation through cool night respite despite warm days. Cluster microclimate effects on consumption inclu sun-exposed clusters showing 35-70% faster consumption than shaded clusters through elevated temperature 4-10°C above air temperature, canopy position effects with exterior southern/western exposure (northern hemisphere) experiencing maximum consumption while interior northern positions showing moderation, and targeted shading reducing consumption 25-50% in warm regions critical management tool for acid retention. Cultivar variation in consumption characteristics demonstrates: low-acid varieties displaying high consumption rates 0.30-0.55 g/L per day peak sustained over extended period 25-40 days producing 85-95% total depletion, moderate cultivars showing 0.22-0.42 g/L per day over 30-45 days achieving 70-85% depletion, and high-acid types exhibiting lower rates 0.15-0.32 g/L per day over 35-50 days resulting 55-75% depletion reflecting genetic differences in respiratory capacity and metabolic programming. Physiological integration shows: respiratory malate consumption coordinated with sugar accumulation creating characteristic ripening metabolism where catabolic acid oxidation supports anabolic sugar storage, carbon balance showing net import through sugar far exceeding loss through acid consumption maintaining overall carbon accumulation, and metabolic flexibility enabling berry to utilize both hexoses and organic acids as respiratory substrates depending on availability and metabolic demands. Quality implications inclu consumption rate and duration determining final malate levels critical for wine acid balance, excessive consumption in warm vintages producing low-acid fruit 5-7 g/L potentially beneficial for some styles but requiring extended ripening or deacidification for others, and spatial variation in consumption across vineyard creating maturity heterogeneity affecting harvest decisions. Viticultural management targeting optimal consumption includes: canopy practices controlling berry temperature with strategic shading reducing consumption 30-55% in warm regions preserving acids, irrigation potentially moderating through cooling effects though mechanisms complex, harvest timing balancing continued sugar accumulation against progressive acid loss requiring integrated maturity assessment, and environmental monitoring tracking heat accumulation enabling prediction of consumption trajectory informing management decisions.
Respiratory_Malate_Consumption is distinguished from Malate_Degradation_Onset by representing ongoing process (0.15-0.45 g/L per day) rather than onset, from Acid_Metabolism_Activation by focusing on respiratory substrate use rather than enzymatic activation, and from sugar accumulation by representing catabolic oxidation rather than anabolic storage. Duration 35-55 days BBCH 83-87. Temperature Q₁₀ 2.2-3.8. Energy yield ~15 ATP per malate. Contributes 35-65% berry respiration during peak. Respiratory_Malate_Consumption represents major carbon and energy flux during ripening fundamentally affecting berry composition and wine quality.
Energie-Erzeugung aus Malat
L-Malat-oxidative Decarboxylierung
L-malate oxidative decarboxylation
energy generation from malate
malic acid respiratory catabolism
oxidative Malat-Verwendung
oxidative malate utilization
Äpfelsäure-respiratorischer Katabolismus
Ruffner HP, Hawker JS, Hale CR (1976) Temperature and enzymic control of malate metabolism in berries of Vitis vinifera Phytochem 15:1877-1880 classic respiratory consumption study Sweetman C, Sadras VO, Hancock RD, Soole KL, Ford CM (2014) Metabolic effects of elevated temperature on organic acid degradation in ripening Vitis vinifera fruit J Exp Bot 65:5975-5988 comprehensive temperature effects Lakso AN, Kliewer WM (1975) The influence of temperature on malic acid metabolism in grape berries Plant Physiol 56:370-372 respiratory consumption quantification.
Respiratory Malate Consumption
Respiratory quotient assessment in ripening Chardonnay berries tracks the metabolic shift during maturation, with pre-véraison berries showing RQ of 0.98 (primarily carbohydrate metabolism), transition-stage berries displaying elevated RQ of 1.25 (organic acid catabolism releasing CO₂ without proportional O₂ consumption as malic acid is decarboxylated), and mature berries returning to RQ of 1.02 (carbohydrate-dominated metabolism after acid reduction), demonstrating how RQ reveals the metabolic transformations underlying fruit composition changes during ripening.
Respiratory quotient quality is a respiration rate quality that represents the molar ratio of CO₂ released to O₂ consumed during respiratory metabolism, characterizing the substrate being oxidized and the completeness of oxidation that provides diagnostic information about respiratory biochemistry and the carbon sources fueling metabolic activity. Within the BFO framework, this quality specifically addresses the stoichiometry of respiratory gas exchange, distinguishing the RQ ratio from absolute respiration rates while recognizing that RQ provides unique information about metabolic substrate use that respiration rate alone cannot reveal. The ontological significance lies in respiratory quotient indicating what the vine is respiring and thus how efficiently carbon is being used, where RQ near 1.0 indicates carbohydrate oxidation (the normal situation with abundant photosynthate), RQ below 1.0 suggests lipid or protein oxidation (indicating carbohydrate depletion or specific metabolic demands), and RQ above 1.0 indicates incomplete oxidation, organic acid synthesis, or fermentation, with these distinctions enabling metabolic diagnosis from gas exchange stoichiometry. The biochemical basis for RQ variation reflects the different oxidation states of metabolic substrates, with carbohydrate oxidation (C₆H₁₂O₆ + 6O₂ → 6CO₂ + 6H₂O) yielding RQ = 1.0, lipid oxidation yielding RQ approximately 0.7 (due to the reduced state of fatty acid carbon), protein oxidation yielding RQ approximately 0.8-0.9 (depending on amino acid composition), and organic acid synthesis consuming CO₂ without proportional O₂ consumption, potentially elevating apparent RQ. The physiological situations affecting RQ in grapevine include normal carbohydrate metabolism (RQ near 1.0), carbohydrate starvation (declining RQ as lipids and proteins are mobilized), rapid growth demanding organic acid skeletons (potentially elevated RQ), and fruit ripening involving organic acid catabolism (elevated RQ in berries metabolizing malate). Understanding respiratory quotient quality enables metabolic substrate identification, carbohydrate status assessment, and appreciation of the respiratory biochemistry underlying carbon balance in different vine organs and conditions.
Respiratory_Quotient_Quality addresses the stoichiometric ratio of respiratory gas exchange indicating metabolic substrate. This quality provides metabolic diagnostic information beyond that available from respiration rate alone. The class supports metabolic status assessment and substrate identification through RQ measurement revealing the biochemical basis of respiratory metabolism.
CO₂/O₂ ratio
CO₂/O₂-Verhältnis
RQ-Wert
Respiratorischer Quotient
Stoffwechselquotient
metabolic quotient
respiratory ratio
Lambers H et al. (2008) Plant Physiological Ecology, 2nd Ed. Springer, New York, pp. 101-150 Tcherkez G et al. (2017) Leaf day respiration: low CO₂ flux but high significance for metabolism and carbon balance. New Phytologist 216(4): 986-1001
Respiratory Quotient Quality
In a Stellenbosch Chenin Blanc vineyard, respiratory substrate change was quantified over 12 days from late January to early February Year N+1 (BBCH 83-84) using gas exchange and isotope studies when respiratory quotient increased from 1.03 to 1.24, ¹³C-malate tracer showing 48% of respiratory CO₂ from malate (from 18% pre-veraison), berry O₂ consumption 1.65 μmol g⁻¹ h⁻¹, combined malic enzyme 52 units/g FW, malate declining from 11.2 to 8.1 g/L, under warm South African conditions (daily high 28-32°C).
Respiratory_Substrate_Change represents the transition in berry respiratory metabolism from predominantly hexose-based carbohydrate oxidation during green berry growth toward mixed substrate utilization incorporating substantial organic acid catabolism during ripening creating the metabolic foundation for simultaneous sugar storage and acid decline defining characteristic ripening metabolism pattern. This substrate change occurs during veraison onset through mid-veraison (BBCH 83-84), typically 2-8 days post-metabolic pathway transition initiation, and is characterized by respiratory quotient increasing from 0.92-1.05 (green berry indicating primarily carbohydrate oxidation) to 1.08-1.32 (ripening indicating substantial organic acid contribution) reflecting altered CO₂:O₂ stoichiometry, malate contribution to respiratory CO₂ production increasing from 5-8% in berry tissue) supporting complete oxidation while hypoxia (<2-3%) promoting fermentative metabolism shifting substrate utilization, water status potentially affecting substrate availability and catabolism rates through effects on metabolite transport and enzyme activities, and nutrient status particularly nitrogen affecting respiratory enzyme capacity and thus potential substrate utilization rates. Cultivar genetic variation shows low-acid varieties exhibiting dramatic substrate change with organic acid contribution potentially reaching 65-80% of respiratory CO₂ during peak decline reflecting extensive acid mobilization, moderate cultivars displaying 45-65% acid contribution during peak, and high-acid varieties showing 35-55% contribution maintaining substantial hexose respiration even during active acid decline reflecting genetic differences in metabolic regulation and substrate preference. Physiological significance includes substrate change enabling efficient resource utilization by catabolizing stored acids for energy supporting ripening while simultaneously storing imported sugars avoiding wasteful sugar catabolism, metabolic flexibility providing resilience allowing berry to adjust substrate utilization based on availability and demands, and energy economy optimizing total energy extraction from available carbon sources maximizing ATP production per unit photosynthate invested. Quality implications include respiratory substrate directly affecting berry carbon budget with excessive acid catabolism representing yield loss while insufficient catabolism maintains undesirable high acids, coordination between substrate change and other ripening processes affecting final composition quality, and temperature sensitivity of substrate change contributing to vintage variation in final acid levels particularly in warm versus cool years. Viticultural management considerations include canopy photosynthesis ensuring adequate hexose supply supporting both storage and respiratory needs, temperature management through cluster exposure controlling respiratory rates and substrate preference affecting acid retention, and understanding substrate change informing carbon budget optimization for quality production. Research applications include respirometry measuring O₂ consumption and CO₂ production calculating RQ revealing substrate transitions, isotope tracer studies with labeled hexoses and organic acids quantifying relative contributions, metabolomic profiling tracking substrate pool dynamics, and enzyme activity assays confirming capacity for altered substrate utilization patterns during ripening.
Respiratory_Substrate_Change is distinguished from Metabolic_Pathway_Transition by representing substrate utilization rather than enzymatic reorganization, from Acid_Metabolism_Shift by focusing on respiratory fuel preference rather than overall metabolism, and from Respiratory_Malate_Consumption by showing transition rather than ongoing process. RQ increases 0.95-1.05 → 1.08-1.32. Malate contribution 35-60% respiratory CO₂ during ripening. Duration 6-12 days BBCH 83-84. Respiratory_Substrate_Change represents fundamental alteration in berry energy metabolism during ripening.
Hexose-zu-Säure-Atmungs-Änderung
Substrat-Verwendungs-Verschiebung
hexose-to-acid respiration change
metabolic fuel switching
metabolische Brennstoff-Umschaltung
respiratorische Brennstoff-Übergang
respiratory fuel transition
substrate utilization shift
Ruffner HP, Hawker JS, Hale CR (1976) Temperature and enzymic control of malate metabolism in berries of Vitis vinifera Phytochem 15:1877-1880 respiratory substrate Sweetman C, Sadras VO, Hancock RD, Soole KL, Ford CM (2014) Metabolic effects of elevated temperature on organic acid degradation in ripening Vitis vinifera fruit J Exp Bot 65:5975-5988 substrate utilization Coombe BG (1992) Research on development and ripening of the grape berry Amer J Enol Vitic 43:101-110 respiratory metabolism during ripening.
Respiratory Substrate Change
Resveratrol content quality marketing of Muscadine wines from the southeastern United States leverages the exceptionally high concentrations (5-40 mg/L in wine, versus 0.5-5 mg/L for typical V. vinifera wines) characteristic of Muscadinia rotundifolia cultivars, with the enhanced resveratrol content serving as a health-positioning attribute that differentiates Muscadine wines in health-conscious consumer segments this resveratrol advantage, combined with ellagic acid from Muscadine's unique phenolic profile, provides functional food marketing claims that command premium pricing in specialty markets.
Resveratrol_Content_Quality represents the chemical quality characterizing the concentration of resveratrol (3,5,4'-trihydroxystilbene) and its derivatives in grape berry tissue, capturing the best-studied stilbene with documented biological activities and health-associated effects. Resveratrol exists in two geometric isomers, with trans-resveratrol as the predominant biologically active form synthesized by stilbene synthase, and cis-resveratrol formed through photoisomerization of the trans form. In grape berries, resveratrol occurs both as the free aglycone and as glucosides (piceids), with trans-piceid (resveratrol-3-O-glucoside) often exceeding free resveratrol concentration. The total resveratrol content (free plus glycosylated, expressed as trans-resveratrol equivalents) in grape skins varies enormously depending on cultivar, growing conditions, and stress exposure. Baseline concentrations in healthy fruit typically range from 0.5-5 mg/kg fresh weight for skins, with red grape cultivars generally accumulating higher levels than white cultivars due to coordinated expression of the phenylpropanoid pathway. Stress induction can elevate resveratrol 10-50 fold, with Botrytis infection producing some of the highest observed concentrations. The health interest in resveratrol originated from the "French Paradox" observation linking moderate red wine consumption with cardiovascular health, spawning extensive research into resveratrol bioavailability, metabolism, and biological effects. While the clinical significance of dietary resveratrol remains debated, consumer awareness has elevated resveratrol content to a marketing-relevant quality attribute. Winemaking practices affect resveratrol extraction and stability, with extended skin contact increasing wine resveratrol content while oxidative conditions may degrade the compound.
Resveratrol_Content_Quality specifically addresses the most studied grape stilbene, including both free and glycosylated forms. The distinction between trans and cis isomers is analytically important, with trans-resveratrol being the primary biosynthetic product and the form with better-characterized biological activity. Resveratrol content in wines typically ranges from 0.1-15 mg/L, substantially lower than the concentrations used in many research studies, creating debate about the dietary relevance of wine resveratrol. Analysis employs HPLC with UV detection at 306 nm or mass spectrometry for comprehensive stilbene profiling.
3,5,4'-Trihydroxystilben-Gehalt
3,5,4'-trihydroxystilbene concentration
Piceid-Vorläuferqualität
Resveratrolgehalt
piceid precursor quality
resveratrol level
trans-Resveratrol-Konzentration
trans-resveratrol content
Siemann EH and Creasy LL (1992) Concentration of the Phytoalexin Resveratrol in Wine. American Journal of Enology and Viticulture 43(1): 49-52 Renaud S and de Lorgeril M (1992) Wine, Alcohol, Platelets, and the French Paradox for Coronary Heart Disease. The Lancet 339(8808): 1523-1526
Resveratrol Content Quality
Quantitative evaluation of Resveratrol_Production_Disposition at the University of Milan characterized resveratrol accumulation dynamics across 22 cultivars following controlled Botrytis cinerea inoculation of berries at véraison. Berry samples were harvested at 0, 24, 48, 72, and 96 hours post-inoculation for HPLC quantification of trans-resveratrol and its glycoside piceid. Basal resveratrol levels in uninoculated berries ranged from 0.2 μg/g ('Pinot Noir') to 2.8 μg/g ('Primitivo'), representing constitutive variation. Following B. cinerea challenge, Resveratrol_Production_Disposition varied dramatically among cultivars. 'Primitivo' showed the strongest response, reaching peak resveratrol of 85 μg/g at 48 hpi (30-fold induction), while 'Pinot Noir' accumulated only 8 μg/g at peak (40-fold induction but from very low baseline). Infection severity at 96 hpi correlated negatively with resveratrol accumulation (r = -0.65), confirming defensive function. Interestingly, cultivars differed in resveratrol-to-piceid ratios, with some accumulating predominantly free resveratrol (70-80% of total, 'Nebbiolo') and others primarily glycosylated piceid (60-70%, 'Sangiovese'). Free resveratrol showed stronger correlation with resistance (r = -0.71) than total stilbenes (r = -0.58), suggesting that the aglycone form is the active defensive compound and that glycosylation may represent a storage or detoxification mechanism rather than direct defense.
A Resveratrol_Production_Disposition is a Phytoalexin_Production_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to accumulate trans-resveratrol (3,5,4′-trihydroxystilbene), the foundational stilbene phytoalexin from which more complex antimicrobial stilbenoids are derived, in response to pathogen challenge or stress. This disposition represents the production of the primary stilbene product of stilbene synthase activity, which serves both as a direct antimicrobial compound and as the precursor for biosynthesis of more potent derivatives including pterostilbene (methylated resveratrol) and various viniferins (resveratrol oligomers). Resveratrol accumulation following pathogen challenge typically reaches maximum concentrations at 24-72 hours post-inoculation, depending on genotype and pathogen, with concentrations ranging from barely detectable in highly susceptible cultivars to over 100 μg/g fresh weight in strongly responding genotypes. The antimicrobial activity of resveratrol includes inhibition of conidial germination, disruption of hyphal membrane integrity, and interference with pathogen metabolism, with minimum inhibitory concentrations against major grapevine pathogens typically in the range of 50-200 μg/mL. Beyond direct antimicrobial effects, resveratrol serves as a substrate for peroxidases that generate the more potent antimicrobial viniferins, and its accumulation kinetics influence the subsequent profile of stilbene derivatives produced. Resveratrol_Production_Disposition has attracted particular interest due to the compound's human health benefits (antioxidant, cardioprotective, potential anticancer properties), creating dual incentives for enhancing this trait through breeding.
Resveratrol_Production_Disposition is_a Phytoalexin_Production_Disposition. Resveratrol is the direct product of STS enzyme activity and serves as precursor for more complex stilbenoids. This class addresses the capacity to accumulate the compound itself, complementing Stilbene_Synthase_Induction_Disposition which addresses the enzymatic capacity. Resveratrol exists in both free and glycosylated (piceid) forms, with the free aglycone typically considered the active antimicrobial form. The compound's human health benefits create additional interest beyond plant defense applications.
Resveratrol-Produktions-Disposition
Trans-Resveratrol-Akkumulation
Trihydroxystilben-Produktion
resveratrol synthesis capacity
trans-resveratrol accumulation
trihydroxystilbene production
Jeandet P et al. (2002) The production of resveratrol by grape berries in different developmental stages, American Journal of Enology and Viticulture 42:41-46 Adrian M et al. (2000) Stilbene content of mature Vitis vinifera berries in response to UV-C elicitation, Journal of Agricultural and Food Chemistry 48:6103-6105 Langcake P, Pryce RJ (1976) The production of resveratrol by Vitis vinifera and other members of the Vitaceae as a response to infection or injury, Physiological Plant Pathology 9:77-86
Resveratrol Production Disposition
Retraining capacity enables vineyard renovation strategies that extend productive lifespan without replanting, with successful retraining restoring 80-100% of pre-renovation yield within 3-4 years. Trunk renewal from basal suckers requires selection of vigorous, well-positioned shoots (typically 2-4 candidates) with subsequent training to the target architecture over 1-2 seasons. Training system conversion (e.g., head-trained to bilateral cordon) exploits retraining capacity to improve mechanization compatibility or canopy microclimate. Cultivar differences in retraining success relate to latent bud fertility (fruitfulness of renewal shoots), growth vigor (rate of structure establishment), and wood flexibility (ease of positioning during training). High-vigor cultivars such as Thompson Seedless and Barbera show rapid retraining (full production year 3), while low-vigor types such as Pinot Noir may require 4-5 years. Rootstock effects on scion retraining capacity are significant, with vigorous rootstocks accelerating structure establishment.
A recovery disposition that is the inherited capacity of a Vitis vinifera plant to be successfully retrained to a new canopy architecture following trunk renewal, training system conversion, or severe corrective pruning, through vigorous shoot growth from selected renewal positions and the establishment of new permanent structures within acceptable production recovery timeframes.
Retraining capacity provides the foundation for vineyard renovation practices that preserve the value of established root systems and site adaptation while replacing damaged or inefficient above-ground structures. The trait is increasingly important as trunk disease pressure creates need for trunk renewal in otherwise healthy vineyards. Successful retraining requires both biological capacity (vigorous growth, fruitful buds) and appropriate timing/technique (dormant versus growing season renewal, wound management). Economic analysis of retraining versus replanting depends on cultivar-specific recovery trajectory.
Erziehungsumstellung
Neuformierungspotential
Umerzugsfähigkeit
renovation potential
restructuring capacity
training flexibility
Intrieri C, Poni S (1995) Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality of mechanized Italian vineyards. American Journal of Enology and Viticulture 46:116-127 Sivilotti P et al. (2016) Grapevine response to trunk renewal: yield recovery, carbohydrate partitioning and pruning weight. Scientia Horticulturae 207:165-172
Retraining Capacity Disposition
Retronasal aroma intensity measurement comparing oak-aged and unoaked Chardonnay during consumption reveals equivalent orthonasal intensity of 6.5 but different retronasal intensity of 7.8 for oaked versus 5.2 for unoaked, with the 50% higher retronasal intensity in oaked wine attributed to less volatile oak-derived compounds (vanillin, oak lactones) that release primarily under the elevated temperature and agitation of oral conditions, explaining why oak contribution appears more prominent on the palate than the nose and demonstrating the importance of retronasal assessment for understanding actual drinking experience.
Retronasal aroma intensity quality is an aroma intensity quality that represents the magnitude of olfactory sensation perceived through the internal nasopharyngeal passage during wine consumption, characterizing the aromatic strength experienced when volatile compounds released in the warmed oral cavity travel via the retronasal pathway to the olfactory epithelium during and after swallowing. Within the BFO framework, this quality specifically addresses the intensity perceived via the retronasal route, distinguishing in-mouth aroma perception from orthonasal sniffing while recognizing that retronasal perception integrates with taste and mouthfeel to create the unified flavor experience dominating wine appreciation during actual consumption. The ontological significance lies in retronasal intensity determining the aromatic contribution to flavor during drinking, where strong retronasal presence creates the impression of aromatic wines that "explode" on the palate, while weak retronasal intensity produces wines tasting less aromatic than they smell, with the correlation between orthonasal and retronasal intensity varying among wines depending on volatile compound properties. The physiological basis for retronasal perception involves volatile release from warmed wine in the oral cavity, transport through the nasopharynx to the olfactory epithelium during exhalation and swallowing movements, with perception affected by wine temperature in mouth, duration of oral contact, breathing pattern, and swallowing mechanics that pump volatiles through the retronasal passage. The compositional determinants of retronasal intensity include compound volatility at mouth temperature (37°C versus ambient), solubility in saliva affecting release kinetics, and binding to oral mucosa creating sustained release, with less volatile compounds potentially contributing more to retronasal than orthonasal perception due to the elevated temperature and agitation in the mouth. The relationship between retronasal aroma and taste perception involves cross-modal enhancement where certain aromas (fruity, sweet) enhance taste perception of sweetness, demonstrating the integration of retronasal olfaction with gustation in flavor construction. Understanding retronasal aroma intensity enables assessment of the aromatic contribution to actual drinking experience, explanation of wines that smell better than they taste (or vice versa), and optimization of consumption conditions for maximum flavor integration.
Retronasal_Aroma_Intensity_Quality specifically addresses the in-mouth route of aroma perception that integrates with taste to create flavor. This quality often differs from Orthonasal_Aroma_Intensity_Quality for the same wine due to different release conditions and compound properties. The class supports understanding of the aroma-taste integration that dominates actual consumption experience, complementing the orthonasal assessment that provides first impression.
Gaumenduftintensität
Mundaromaintensität
flavor aroma intensity
in-mouth aroma intensity
internal aroma strength
interne Geruchsstärke
palate aroma intensity
retronasale Aromaintensität
Buettner A, Schieberle P (2000) Influence of mastication on the concentrations of aroma volatiles — some aspects of flavour release and flavour perception. Food Chemistry 71(3): 347-354 Small DM et al. (2005) Differential neural responses evoked by orthonasal versus retronasal odorant perception in humans. Neuron 47(4): 593-605
Retronasal Aroma Intensity Quality
Reversible deformation capacity determination for Red Globe table grapes using cyclic compression testing identifies 8.5% strain as the threshold below which complete recovery occurs within 5 minutes, with berries compressed to 7% strain showing 100% dimensional recovery and no detectable cellular damage under microscopy, while berries compressed to 12% strain show only 85% recovery and visible cell rupture in the compression zone, establishing the 8.5% strain limit as the design criterion for packing line cushioning to prevent mechanical damage during handling.
Reversible deformation capacity quality is an elasticity quality that represents the maximum extent of deformation from which complete recovery to original dimensions occurs upon stress removal, characterizing the strain limit of the elastic region beyond which permanent plastic deformation accumulates. Within the BFO framework, this quality quantifies the boundary between elastic and plastic mechanical behavior, providing a critical threshold for damage assessment where deformation within this capacity leaves no lasting effect while deformation exceeding this capacity causes permanent structural change. The ontological significance lies in reversible deformation capacity defining the safe handling envelope for grape tissues, where impacts, compressions, and distortions within this limit do not affect quality while exceedance initiates bruising, cell rupture, and juice loss that compromise marketability and storage potential. Reversible deformation capacity in grape berries typically ranges from 5-15% strain depending on variety, ripeness, and tissue type, with firmer pre-véraison berries showing higher reversible capacity than soft ripe berries where cell wall integrity is reduced. The tissue-specific variation in reversible capacity reflects differences in cell wall strength, turgor pressure contribution to mechanical resistance, and tissue architecture determining stress distribution, with skin generally showing higher reversible capacity than pulp due to the reinforcing effect of the compact epidermal cell arrangement. The recovery kinetics following deformation within the reversible range show time-dependent behavior, with immediate elastic recovery followed by delayed recovery as fluid redistribution and polymer chain relaxation restore original configuration, and incomplete recovery within the measurement period does not necessarily indicate permanent damage if continued recovery would eventually achieve full restoration. Measurement employs cyclic loading with strain incrementally increased until hysteresis indicates energy dissipation associated with damage, or single-load recovery testing measuring residual deformation after defined recovery period. Understanding reversible deformation capacity quality enables specification of handling force limits, design of harvesting and packing equipment to minimize damage, and quality grading based on the mechanical history inferred from deformation assessment.
Reversible_Deformation_Capacity_Quality addresses the strain threshold separating elastic from plastic behavior, complementing Elastic_Deformation_Quality and Plastic_Deformation_Quality which characterize the response within each regime. This quality provides the critical parameter for damage prevention in handling system design. The class supports quality preservation through mechanical handling optimization based on variety-specific and ripeness-specific deformation limits.
Erholungskapazität
elastic limit
elastische Grenze
non-damaging deformation range
recovery capacity
resilience capacity
reversible Verformungskapazität
schadensfreier Verformungsbereich
Opara LU, Pathare PB (2014) Bruise damage measurement and analysis of fresh horticultural produce — a review. Postharvest Biology and Technology 91: 9-24 Ahmadi E et al. (2010) Bruise susceptibility of peaches (Prunus persica) as affected by impact energy, tissue type, and fruit ripeness. Postharvest Biology and Technology 55(3): 157-163
Reversible Deformation Capacity Quality
In Chardonnay vines in Burgundy, reversible growth suppression is demonstrated during February and March when warm periods with temperatures of 14-16°C induce visible bud swelling to BBCH stage 01-03, followed by growth arrest and maintenance of swollen bud state when temperatures return to 4-7°C, with this reversible pattern continuing until sustained April warming provides irreversible commitment to budbreak progression through BBCH stage 05 and beyond.
Reversible growth suppression in Vitis vinifera represents the characteristic of ecodormancy whereby the suppression of growth by unfavorable environmental conditions can be immediately reversed upon provision of favorable conditions, contrasting with the irreversible nature of endodormancy suppression that persists regardless of conditions until chilling-dependent release has occurred. This reversibility reflects the absence of stable internal barriers to growth during ecodormancy, with growth suppression depending entirely on the immediate environmental context rather than on persistent physiological states that require time and specific conditions for removal. The cellular basis of reversibility involves the maintenance of growth-ready cellular states during ecodormancy, with cell cycle machinery poised for activation, biosynthetic enzyme systems prepared for rapid upregulation, and hormone signaling pathways sensitized to respond to growth-promoting conditions, enabling immediate growth response when suppressive conditions are relieved. The practical demonstration of reversibility uses forcing tests that expose ecodormant buds to optimal growth conditions and measure the speed of growth response, with reversible suppression indicated by rapid budbreak within 10-14 days contrasting with the extended times or failure characteristic of the irreversible suppression of endodormancy. The field manifestation of reversibility includes the partial growth responses commonly observed during warm periods in late winter, with buds initiating swelling or other early growth stages during favorable conditions and arresting development when temperatures return to suppressive levels, creating the stop-and-start growth pattern characteristic of transitional spring weather. The ecological implications of reversible suppression relate to frost vulnerability, as the capacity for immediate growth response to warming creates the risk that temporary favorable conditions will trigger growth that is subsequently damaged by return to cold temperatures, a risk that is absent during endodormancy when irreversible suppression protects against such responses. The transition from reversible suppression to committed growth occurs when favorable conditions persist long enough to pass a point of no return in the budbreak process, after which growth continues even if conditions subsequently become unfavorable, typically occurring after visible bud swelling has progressed to green tip emergence.
Reversible_Growth_Suppression represents the reversibility characteristic within Conditional_Dormancy. Distinguished from irreversible endodormancy suppression that persists regardless of conditions. Demonstrated by immediate forcing response and field stop-and-start growth patterns. Reflects absence of stable internal growth barriers. Creates frost vulnerability from premature growth response. Transition to irreversible growth after green tip emergence typically at BBCH 07-09.
bedingte Unterdrückung
conditional suppression
reversible Dormanz
reversible Wachstumsunterdrückung
reversible growth inhibition
temporary growth arrest
vorübergehender Wachstumsstopp
Reversible Growth Suppression
Genetic mapping of Rgr_Mediated_Resistance_Disposition at the Julius Kühn-Institut utilized a segregating population from the cross 'Börner' (resistant) × V. vinifera 'Lemberger' (susceptible). Phenotypic evaluation employed detached leaf assays with G. bidwellii conidial suspension (10⁵ conidia/mL), scoring lesion development on a 1-9 scale at 14 days post-inoculation. Resistance segregated in a 1:1 ratio consistent with single dominant gene inheritance. QTL analysis identified a major locus on chromosome 14 explaining 76% of phenotypic variance, designated Rgr1. Resistant individuals carrying the Rgr1 allele exhibited small necrotic flecks (OIV rating 7-9) without subsequent pycnidia formation, while susceptible siblings developed expanding tan lesions with abundant pycnidial sporulation (OIV rating 1-3). The Rgr1-linked SSR marker GF14-67 enables marker-assisted selection with 98% accuracy in predicting phenotype. Advanced breeding selections combining Rgr1 with Rpv and Ren loci from 'Börner' represent multiple-disease-resistant germplasm suitable for humid climate viticulture, demonstrating the value of this rootstock cultivar as a source of combined fungal disease resistances.
An Rgr_Mediated_Resistance_Disposition is a Black_Rot_Resistance_Disposition that inheres in a Vitis plant carrying one or more Rgr (Resistance to Guignardia) loci and is realized through major gene-mediated defense responses upon encounter with Guignardia bidwellii. The Rgr loci encode resistance factors that confer qualitative, race-specific or broad-spectrum resistance to black rot, analogous to the Rpv and Ren/Run loci for downy and powdery mildew respectively. Rgr1, the first characterized black rot resistance locus, was identified in the rootstock cultivar 'Börner' (V. riparia × V. cinerea) and maps to chromosome 14, where it co-localizes with a resistance gene cluster also containing downy mildew resistance factors. The Rgr1-mediated resistance phenotype is characterized by rapid hypersensitive response following G. bidwellii penetration, resulting in small necrotic flecks that arrest fungal colonization before the pathogen completes its biotrophic phase and transitions to tissue-destructive necrotrophy. Additional Rgr loci have been identified in other resistant germplasm sources including V. rupestris and Muscadinia rotundifolia, providing multiple independent resistance genes for potential pyramiding strategies. Marker-assisted selection using Rgr-linked molecular markers enables efficient introgression of black rot resistance into V. vinifera breeding populations, addressing a critical disease pressure for grape production in humid continental climates. The identification and deployment of Rgr loci represents an important advance for sustainable viticulture in regions where black rot has historically limited cultivation of high-quality wine grape cultivars.
Rgr_Mediated_Resistance_Disposition is_a Black_Rot_Resistance_Disposition. Rgr1 was identified in 'Börner', a V. riparia × V. cinerea hybrid rootstock that also carries downy mildew resistance, with both resistances mapping to chromosome 14. The Rgr nomenclature follows the established Vitis resistance gene naming convention. Additional Rgr loci from other resistant germplasm sources await characterization. Rgr-mediated resistance operates through rapid hypersensitive response during the biotrophic infection phase, preventing transition to destructive necrotrophy.
Börner-derived resistance
Majorgen-Black-Rot-Resistenz
Rgr locus black rot resistance
Rgr-Schwarzfäule-Resistenz
Rgr-vermittelte Resistenz-Disposition
major gene black rot resistance
Rex F et al. (2014) QTL mapping of black rot (Guignardia bidwellii) resistance in the grapevine rootstock 'Börner', Theoretical and Applied Genetics 127:1667-1677 Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance and ripening of grapevines, Molecular Genetics and Genomics 291:1573-1594
Rgr Mediated Resistance Disposition
Detailed investigation of Rhizosphere_Acidification_Disposition at INRAE Montpellier employed pH-sensitive agar visualization, rhizosphere pH microelectrode measurements, and H⁺-ATPase activity assays across 12 rootstocks under iron-sufficient and iron-deficient conditions. Roots were embedded in pH indicator agar (bromocresol purple) to visualize acidification patterns, while selective microelectrodes measured pH at 50 μm from the root surface at defined positions along the root axis. Rhizosphere_Acidification_Disposition showed striking genotypic variation. '41B' exhibited strong, extensive acidification with yellow zones (pH < 5.5) extending 3-4 mm from root surfaces under Fe deficiency, microelectrode pH measurements of 5.8 at the root surface versus bulk pH 7.8 (2.0 unit depression), and proton efflux rates of 3.2 μmol H⁺ g⁻¹ root FW h⁻¹ representing 4.5-fold induction over Fe-sufficient controls. Acidification was localized primarily in the subapical zone (2-10 mm from root tip) where active iron uptake occurs. '140 Ruggeri' showed even stronger acidification (3.8 μmol H⁺ g⁻¹ h⁻¹, pH depression to 5.5), while 'Fercal' achieved intermediate but effective levels (2.4 μmol g⁻¹ h⁻¹). '3309C' displayed weak Rhizosphere_Acidification_Disposition with limited yellow zones, surface pH of only 7.2 (0.6 unit depression), and efflux of 0.9 μmol g⁻¹ h⁻¹ with only 1.8-fold induction. H⁺-ATPase gene expression (VvHA1) analysis confirmed that high-acidifying '41B' showed 6-fold transcript accumulation under Fe deficiency versus 2-fold in '3309C', indicating transcriptional regulation underlies genotypic differences.
A Rhizosphere_Acidification_Disposition is an Iron_Acquisition_Efficiency_Disposition that inheres in Vitis root epidermal and cortical cells and is realized through enhanced capacity to extrude protons (H⁺) into the rhizosphere, lowering local soil pH to increase ferric iron solubility in the immediate root vicinity. This disposition represents the first component of Strategy I iron acquisition, addressing the fundamental problem that Fe³⁺ solubility decreases approximately 1000-fold for each unit increase in pH above pH 4, making iron effectively unavailable at the pH 7.5-8.5 typical of calcareous soils. Rhizosphere acidification is mediated primarily by plasma membrane H⁺-ATPases that actively pump protons from the cytoplasm to the apoplast and soil solution, with the activity of these proton pumps increasing under iron deficiency. Effective acidification can reduce rhizosphere pH by 1-2 units relative to bulk soil, dramatically increasing local Fe³⁺ solubility and providing substrate for the subsequent ferric reductase reaction. Rhizosphere_Acidification_Disposition varies among rootstock genotypes in both the basal proton extrusion rate and the degree of upregulation under iron deficiency, with lime-tolerant genotypes typically showing 3-5 fold induction of acidification capacity under low Fe availability compared to 1.5-2 fold in sensitive genotypes. However, acidification capacity alone does not guarantee lime tolerance, as the high buffering capacity of calcareous soils can neutralize protons faster than roots can supply them, requiring integration with other acquisition mechanisms.
Rhizosphere_Acidification_Disposition is_a Iron_Acquisition_Efficiency_Disposition. Acidification is the first step in Strategy I, increasing local Fe³⁺ solubility. Mediated by plasma membrane H⁺-ATPases, activity localized to subapical root zones. Acidification capacity is iron-regulated, with strong induction under Fe deficiency in tolerant genotypes. pH reduction of 1-2 units can increase Fe solubility 100-1000 fold. Calcareous soil buffering capacity can limit effectiveness of acidification alone. VvHA genes encode the responsible H⁺-ATPases.
H⁺-ATPase activity
Protonen-Extrusion
Rhizosphären-Ansäuerungs-Disposition
Wurzel-Ansäuerung
proton extrusion capacity
root acidification
Dell'Orto M et al. (2000) Fe-deficiency responses in Parietaria diffusa: a calcicole plant, Journal of Plant Nutrition 23:1719-1732 Römheld V, Marschner H (1986) Evidence for a specific uptake system for iron phytosiderophores in roots of grasses, Plant Physiology 80:175-180 Santi S, Schmidt W (2008) Laser microdissection-assisted analysis of the functional fate of iron deficiency-induced root hairs in cucumber, Journal of Experimental Botany 59:697-704
Rhizosphere Acidification Disposition
The Ried name Achleiten designates a 15-hectare terraced Einzellage in the Wachau village of Weißenkirchen, planted predominantly to Riesling on primary rock (gneiss and mica schist) soils at 220-450 meters elevation with south-southwest exposure — classified as a Smaragd-eligible site by the Vinea Wachau association, the Ried Achleiten name on labels from producers including Prager, Hirtzberger, and Rudi Pichler communicates a specific terroir profile characterized by pronounced minerality, herbal complexity, and aging potential attributable to the site's steep gradient, primary rock geology, and warm microclimate protected from northern winds by the Jauerling massif.
A vineyard designation within the Austrian wine classification system that identifies a wine as originating from a named Ried — a traditionally delimited and officially registered vineyard site within an Austrian wine-growing community, recognized as the fundamental unit of site-specific terroir identification in the Austrian vineyard classification established by the 2009 Wine Law and progressively implemented through regional vineyard cadasters.
Ried_Name documents Austria's vineyard site designation system, which was formalized in the 2009 Wine Law (Weingesetz 2009) and has been progressively implemented through regional vineyard cadasters (Riedenkarten) maintained by regional wine committees. The Austrian Ried concept corresponds functionally to the German Einzellage and the French lieu-dit/climat but operates within a distinctly Austrian regulatory framework: the 2009 Wine Law and subsequent amendments established formal criteria for Ried registration including historical documentation of the site name, defined geographic boundaries recorded in the land cadaster, and viticultural distinctiveness. The integration of Ried designations with the DAC appellation system has created a two-dimensional classification matrix where wines can be identified by both their regional DAC designation (indicating cultivar and style typicity) and their Ried name (indicating site-specific terroir origin) — for example, Kamptal DAC Ried Heiligenstein identifies a Grüner Veltliner or Riesling from the specific Heiligenstein site within the Kamptal DAC appellation.
Austrian vineyard site name
Ried designation
Riedbezeichnung
Rieden-Name
registered Ried name
österreichische Lagenbezeichnung
Austrian Wine Law (Weingesetz 2009) §17 Österreichische Weinmarketinggesellschaft (ÖWM) (2023) Riede und Lagen Wachau Wine Commission (2023) Riedenkarte Wachau
Ried Name
A right unilateral cordon on Vitis vinifera cv. Sauvignon Blanc at left-side row end (when traveling south along row), extending 92cm to the right (eastward) from trunk positioned at row terminus adjacent to property boundary fence, bearing 15 spurs, with vineyard management software database recording "right unilateral" designation linked to GPS coordinates enabling precision viticulture applications including differential fertilization (reduced rates near boundary), targeted disease management (increased fungicide coverage on boundary rows with reduced air circulation), and equipment operation planning (harvester approach sequences accounting for right-side cordon extension), representing directionally-specified unilateral architecture integrated with digital vineyard management systems.
A unilateral cordon extending exclusively in the right direction from the trunk when viewed from a standard perspective along the vineyard row, characterized by single-sided extension toward the right side of the row, absence of left-side cordon development, and directional specificity complementing left unilateral designations in comprehensive vineyard spatial documentation and management coordination. Right unilateral cordons provide directionally specified unilateral architecture opposite to left unilateral configurations, with right designation completing the directional reference system for row-end positions, enabling precise spatial communication for equipment operations, management planning, and vineyard documentation, with the right versus left distinction being essential for coordinated activities in large-scale commercial operations requiring multiple crews and equipment operators working simultaneously across extensive vineyard blocks.
Right unilateral cordons mirror left unilateral cordons in all functional aspects, with the distinction being purely directional reference. The right versus left designation system provides essential spatial vocabulary for large-scale vineyard operations, particularly important for row-end management where each row typically has both left and right unilateral cordons at opposite ends (left unilateral at one end, right unilateral at opposite end when viewed from consistent perspective). Modern vineyard management increasingly incorporates directional specifications in digital systems including GIS mapping, vineyard management software databases, equipment guidance systems, and crew coordination platforms, with precise directional designations enabling targeted interventions, accurate record-keeping, and efficient operations. The standardization of directional reference conventions (establishing consistent viewing perspectives for left/right determinations) is essential for preventing confusion and errors in operations involving multiple personnel and equipment units working across complex vineyard layouts.
dexter cordon
rechte unilaterale Kordon
rechtsseitige Kordon
right-extending cordon
right-side cordon
Right Unilateral Cordon
3309 Couderc (V. riparia Tomenteux × V. rupestris Martin) is the archetypal Riparia × Rupestris rootstock, providing low-to-moderate vigor (pruning weights 0.3-0.5 kg/m), excellent cold hardiness (root LT₅₀ -12°C to -14°C), and premium wine quality to Pinot Noir in Burgundy and Champagne, tolerating up to 11% active lime. 101-14 Millardet et de Grasset (V. riparia × V. rupestris) confers low vigor and early maturity to Merlot in Bordeaux right-bank appellations, with lime tolerance of 9% active lime. Schwarzmann (V. riparia × V. rupestris) provides very low vigor supporting quality-focused Riesling production in Alsace, though restricted to acidic granitic soils (lime tolerance only 10-11%).
A parentage-based rootstock role that inheres in a grafted rootstock by virtue of its genetic derivation from hybridization between Vitis riparia and Vitis rupestris, combining traits from these two complementary North American species. V. riparia × V. rupestris hybrids unite V. riparia's excellent cold hardiness, strong phylloxera resistance, early growth cessation, and low-to-moderate vigor with V. rupestris's improved rooting capacity, deeper root architecture, better drought tolerance, and enhanced vigor. This hybrid combination produces rootstocks adapted to cool-to-temperate climates on acidic to neutral soils (lime tolerance limited to 9-14% active lime), with low-to-moderate vigor suitable for premium wine production, excellent phylloxera resistance from both parental species, and substantially improved nursery propagation success (rooting rates 75-90%) compared to pure V. riparia selections. The V. riparia × V. rupestris group represents one of the earliest and most successful rootstock hybrid combinations, developed primarily by French breeders (Couderc, Millardet, de Grasset) in the late 19th century following the phylloxera crisis.
Riparia_X_Rupestris_Rootstock_Role captures a historically important hybrid combination that remains commercially significant for cool-climate premium wine production on low-lime soils. The combination lacks the lime tolerance contributed by V. berlandieri, restricting these rootstocks to acidic, neutral, or weakly calcareous substrates. All members show relatively low vigor compared to V. berlandieri × V. rupestris hybrids, making them preferred choices where canopy control and fruit concentration are priorities. The hybrid group includes 3309C, 101-14Mgt, Schwarzmann, 161-49C, and numerous lesser-known selections from the prolific late-19th-century French breeding programs.
Riparia × Rupestris rootstock
Riparia-Rupestris-Hybridunterlage
Riparia-×-Rupestris-Unterlage
V. riparia × V. rupestris rootstock
riparia-rupestris hybrid rootstock
Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22 Pongrácz DP (1983) Rootstocks for Grape-vines, David Philip Publishers Whiting JR (2004) Grapevine rootstocks, in Viticulture Vol 1 Resources, Winetitles pp.: 167-188
Riparia X Rupestris Rootstock Role
In a Napa Valley Cabernet Sauvignon vineyard, ripening was tracked over 68 days from early August through mid-October Year N+1 (BBCH 81-89) when berry Brix increased from 12.5° to 25.8° (13.3° gain, average 2.0% per week), titratable acidity declined from 16.8 to 5.6 g/L (67% reduction, average 0.23 g/L per day), pH increased from 3.08 to 3.52, berry firmness decreased from 3.8 to 0.7 N (82% softening), anthocyanin accumulated from 85 to 1240 mg/kg, total volatiles rose from 48 to 385 μg/kg, under warm California conditions (cumulative 1180 degree-days), enabling harvest at optimal phenolic and flavor maturity.
Ripening represents the final major phase of berry development following veraison onset when grape composition transforms from acidic green fruit toward sweet balanced mature berry through coordinated sugar accumulation, organic acid decline, berry softening, color development in red cultivars, and flavor-aroma compound synthesis creating the progressive maturation that determines harvest quality, wine characteristics, and commercial value. This ripening phase extends from veraison completion through harvest (BBCH 81-89), typically spanning 40-80 days depending on cultivar and environmental conditions (earlier maturing varieties 40-55 days, mid-season 50-65 days, late cultivars 60-80 days), and is characterized by berry Brix increasing from veraison establishment 11-16° to harvest maturity 18-30° representing 8-18° gain through sustained hexose import and vacuolar storage, titratable acidity declining from 12-20 g/L to final 4-10 g/L through preferential malic acid catabolism while tartaric acid remains relatively stable, berry pH increasing from 3.0-3.3 to 3.0-3.8 (cultivar-dependent) reflecting net organic acid loss, berry firmness declining 60-85% from veraison onset 2.5-4.5 N to harvest 0.4-1.2 N through cell wall pectin solubilization and turgor reduction creating characteristic soft ripe texture, anthocyanin accumulation in red cultivars from veraison levels 50-200 mg/kg reaching harvest maximum 400-1800 mg/kg (cultivar and conditions dependent) producing characteristic color intensity, and flavor-aroma volatiles accumulating 4-12 fold from veraison onset 25-65 μg/kg to harvest 100-650 μg/kg creating varietal character and sensory ripeness. Physiological basis of ripening involves fundamental metabolic reprogramming initiated at veraison and sustained throughout ripening with phloem unloading transitioning from symplastic to predominantly apoplastic (>85-95%) enabling high-flux sugar delivery, hexose transporter genes (VvHT1, VvHT2, VvHT3, VvHT5, VvHT6) collectively upregulated 100-300 fold above green berry levels creating import capacity 60-170 μmol combined glucose and fructose g⁻¹ h⁻¹ depending on phase and conditions, vacuolar hexose concentrations increasing from 400-700 mM to 1400-2200 mM combined glucose and fructose creating osmotic potential decline from -1.8 to -2.4 MPa to -2.6 to -3.6 MPa driving water import and berry expansion 50-120%, organic acid metabolism shifting from biosynthetic to catabolic mode with malic enzyme genes (VvME1, VvME2) upregulated 20-80 fold enabling malate respiratory consumption 0.15-0.55 g/L per day during active decline while tartaric acid metabolically stable showing only dilutional changes, cell wall modifying enzymes including polygalacturonase, pectinmethylesterase, and expansins progressively solubilizing pectins reducing berry firmness enabling characteristic ripe texture development, anthocyanin biosynthesis pathway genes (particularly PAL, CHS, UFGT) in reds upregulated 50-200 fold producing flavonoid accumulation 10-40 mg/kg per week during active coloration, and secondary metabolism intensifying with terpene synthases, ester-forming enzymes, and other flavor pathway genes activated producing characteristic varietal aromatics. Temporal progression of ripening demonstrates initial phase (BBCH 81-83, days 0-15 post-veraison) with accelerating sugar accumulation 1.2-2.5% Brix per week, moderate acid decline 0.18-0.35 g/L TA per day, continuing softening 15-35% firmness loss, and in reds beginning color development reaching 20-50% final anthocyanin, active ripening phase (BBCH 84-86, days 15-45) exhibiting peak sugar accumulation 2.5-5.0% Brix per week, maximum acid decline 0.30-0.65 g/L TA per day, softening largely completed, color development 50-95% final levels in reds, and flavor compound synthesis intensifying, and late ripening approaching harvest (BBCH 87-89, days 45-70) showing decelerating sugar accumulation 0.8-2.2% Brix per week as limits approached, continuing acid decline 0.10-0.30 g/L per day, firmness stabilizing at ripe levels, color reaching maximum in reds, and flavor complexity developing. Hormonal orchestration of ripening includes abscisic acid as master regulator with sustained elevated levels 1200-2800 ng/g throughout ripening maintaining coordinate expression of sugar transport, acid catabolism, softening, and color genes through ABA-responsive transcription factors, sugar signaling through hexokinase pathway creating feed-forward amplification enhancing ripening progression, declining growth hormones (gibberellin declining from 40-120 ng/g pre-veraison to 35°C impairing ripening through metabolic disruption and potential berry damage, solar radiation affecting berry temperature and photosynthesis with adequate canopy light essential for source capacity while excessive cluster exposure in hot climates potentially detrimental through overheating, water availability with moderate deficit compatible or enhancing quality through concentration effects while severe deficit <-1.6 MPa impairing metabolism disrupting normal ripening progression, and cumulative growing degree-days strongly correlating with ripening progression (base 10°C) with typical requirements 900-1400 degree-days from veraison to harvest depending on cultivar. Cultivar genetic variation demonstrates early-ripening varieties (Chardonnay, Pinot Noir, Gewürztraminer) completing ripening 40-55 days post-veraison enabling cultivation in cool short-season regions, mid-season cultivars (Merlot, Syrah, Sauvignon Blanc) requiring 50-65 days representing majority commercial varieties, and late-ripening types (Cabernet Sauvignon, Nebbiolo, Mourvèdre) needing 60-80 days restricting to warm long-season climates, with genetic differences in sugar accumulation capacity determining final Brix potential 18-30° cultivar-dependent, acid retention characteristics affecting final TA 4-10 g/L and pH 3.0-3.8, color intensity in reds varying 400-1800 mg/kg anthocyanin harvest levels, and flavor profiles reflecting variety-specific terpene, ester, and other volatile patterns creating distinctive sensory characteristics. Quality implications of ripening include sugar-acid balance at harvest critically determining wine style with ratios 20-35:1 (Brix:TA g/L) typical for quality wines, phenolic maturity in reds requiring tannin polymerization and seed browning synchronized with sugar-acid development often necessitating extended hang time beyond sugar maturity, flavor ripeness essential for quality requiring adequate volatile development particularly aromatic varieties where flavor may lag analytical maturity, physiological maturity indicated by seed browning, berry abscission ease, and stem lignification providing additional harvest timing cues beyond chemistry, and vintage variation substantial with warm years producing earlier faster ripening potentially challenging for achieving balanced maturity while cool years extending duration enabling flavor development but risking incomplete ripening in marginal climates. Harvest timing decisions integrate multiple ripening parameters including target sugar levels varying by wine style (table wines 20-24° Brix, fuller styles 22-26°, late harvest 24-32°), acceptable acid levels maintaining TA 5-8 g/L and pH 3.0-3.7 for most quality wines, phenolic assessment particularly reds evaluating tannin polymerization and seed maturation, flavor development through sensory evaluation detecting varietal character and ripeness, weather forecasts considering approaching adverse conditions (rain, frost) potentially necessitating earlier harvest, and market factors including winery capacity and labor availability affecting practical harvest windows. Viticultural management during ripening includes canopy maintenance ensuring continued photosynthesis supporting ripening processes, strategic leaf removal or shading managing cluster microclimate particularly warm regions where temperature moderation critical for balanced ripening, irrigation management providing adequate water without excess while potentially using deficit to enhance concentration and quality, disease control critical as high-sugar soft berries increasingly vulnerable to Botrytis, sour rot, and other pathogens requiring vigilant monitoring and intervention, bird and pest protection preventing crop losses particularly as sugars rise attracting wildlife, and harvest preparation including equipment readiness, labor scheduling, and winery coordination as ripening progression indicates approaching optimal maturity. Monitoring ripening progression involves regular berry sampling every 5-10 days tracking Brix, TA, pH, and malate revealing compositional trajectories, phenolic analysis in reds assessing tannin and anthocyanin development, sensory evaluation detecting flavor ripeness and balance, spatial sampling across vineyard zones revealing heterogeneity requiring selective management, and historical data compilation enabling vintage comparisons and predictive modeling improving management decisions and harvest timing optimization for achieving target quality parameters.
Ripening is distinguished from Veraison by representing maturation phase (BBCH 81-89, 40-80 days) following veraison onset rather than the transition itself (BBCH 81-83, 7-14 days), encompasses complete sugar accumulation and acid decline rather than just onset, and includes late-stage processes like flavor development and physiological maturity absent during early veraison. Ripening represents coordinated progression of multiple parallel processes: sugar accumulation (dominant anabolic), acid decline (dominant catabolic), softening, color (reds), and flavor development. Final composition determines wine quality and style. Duration cultivar-dependent: early 40-55 days, mid-season 50-65 days, late 60-80 days. Temperature critically important: 25-30°C optimal, >35°C impairs. Ripening represents final major berry development phase defining harvest maturity and commercial value.
Beeren-Reifung
Frucht-Reifung
Zucker-Säure-Balance-Entwicklung
compositional maturation
fruit ripening
kompositionelle Maturation
sugar-acid balance development
Keller M (2020) The Science of Grapevines 3rd Ed pages 221-256 ripening comprehensive Coombe BG (1992) Research on development and ripening of the grape berry Amer J Enol Vitic 43:101-110 ripening dynamics Kennedy JA, Matthews MA, Waterhouse AL (2002) Effect of maturity and vine water status on grape skin and wine flavonoids Amer J Enol Vitic 53:268-274 ripening and quality.
Ripening
Ripening data for Chardonnay clones in the Willamette Valley tracks véraison onset for clone 76 at August 10 (±7 days), clone 96 at August 5 (±6 days), and clone 548 at August 14 (±8 days), with corresponding optimal harvest dates of September 18, September 12, and September 24 — a 12-day spread among clones that enables staggered harvest scheduling within multi-clone plantings. Ripening data for Blaufränkisch clones at Deutschkreutz documents clone Bl-2 requiring 1350-1450 GDD (base 10°C) from flowering to 20° KMW harvest threshold versus 1450-1550 GDD for clone Bl-10, confirming Bl-2 as the earlier-ripening selection better suited to cooler Mittelburgenland sites where late-season thermal accumulation is limiting.
Ripening data is an agronomic performance data entity that records the phenological timing, rate, and uniformity of the fruit maturation process in a clonal line, encompassing the dates of key phenological transitions (véraison onset, 50% véraison, physiological maturity, optimal harvest window), the rate of sugar accumulation and acid degradation during the ripening period, the synchrony of ripening within and between clusters, and the elapsed growing degree days (GDD) or calendar days required from flowering to harvest, characterizing the clone's maturation dynamics and its suitability for the thermal resources and harvest logistics of specific viticultural regions.
Ripening data has gained critical importance in the context of climate change, as shifts in maturation timing directly affect wine style, quality, and production logistics. The distinction between early-ripening and late-ripening clones within a cultivar can span 10-21 days for harvest date, representing a substantial phenological difference that determines whether maturation occurs during cool late-season conditions (favorable for aromatic complexity and acid retention) or hot mid-season periods (promoting high sugar with compressed flavor development). The rate of ripening, independent of timing, provides additional clone characterization — clones with compressed ripening windows (rapid sugar accumulation over 2-3 weeks) demand precise harvest scheduling, while clones with extended ripening plateaus offer greater flexibility. Ripening uniformity within clusters (simultaneous versus staggered berry maturation) is a clone-specific trait with direct oenological implications — uniform ripening produces more homogeneous must composition, while asynchronous ripening may contribute desirable complexity in some wine styles but problematic heterogeneity in others. The GDD requirement from flowering to harvest provides a site-transferable ripening metric less dependent on calendar date than absolute phenological timing, enabling assessment of whether a clone will achieve maturity within the thermal resources of a target region. The increasing use of polyclonal plantings (multiple clones of a single cultivar) strategically exploits ripening data to extend the harvest window, diversify vintage risk, and create blending complexity through clonal maturation sequence differences.
Ausreifungsdaten
Reifedaten
Reifephänologie
Reifeverlaufsdaten
Reifungsdaten
harvest timing data
maturation data
maturation timing data
ripening dynamics record
ripening phenology
Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 369-420 Jones GV, Davis RE (2000) Climate influences on grapevine phenology, grape composition, and wine production and quality for Bordeaux, France. American Journal of Enology and Viticulture 51(3): 249-261 Parker AK et al. (2011) General phenological model to characterise the timing of flowering and véraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17(2): 206-216
Ripening Data
Ripening synchrony assessment tracking berry subpopulations through maturation in Zinfandel reveals the compensation dynamics affecting harvest uniformity, with early-véraison berries (color change day 0) reaching 24 Brix at day 35, mid-véraison berries (day +5) reaching 24 Brix at day 38 (partial compensation, 3-day lag versus initial 5-day offset), and late-véraison berries (day +10) reaching 24 Brix at day 47 (minimal compensation, 12-day lag exceeding initial offset), demonstrating how ripening synchrony varies among berry subpopulations with the latest berries actually falling further behind during ripening, creating the progressive Brix spread that makes Zinfandel harvest timing particularly challenging.
Ripening synchrony quality is an intra-plant synchrony quality that represents the temporal coordination of maturation progress among different berries and clusters within a single grapevine during the post-véraison ripening period, characterizing the uniformity of sugar accumulation, acid decline, and flavor development rates that determine whether berries achieving véraison at different times converge toward common maturity or maintain their developmental offsets through harvest. Within the BFO framework, this quality specifically addresses the ripening rate dimension of intra-plant synchrony, distinguishing post-véraison maturation coordination from the véraison transition itself while recognizing that ripening synchrony determines whether initial véraison asynchrony is amplified, maintained, or compensated during the final maturation period that directly affects harvest-time fruit uniformity. The ontological significance lies in ripening synchrony determining the relationship between véraison synchrony and harvest-time maturity uniformity, where high ripening synchrony may compensate for moderate véraison asynchrony (faster ripening of late-véraison berries catching up) or may fail to compensate (all berries ripening at similar rates, maintaining initial offsets), with the outcome affecting the maturity uniformity achievable at any target harvest date. The mechanisms potentially enabling ripening rate compensation include source-sink relationships (late berries having access to photosynthate sources after early berries have satisfied their demand), continued berry growth (late berries potentially showing faster volume increase that dilutes concentration differences), and cluster-level coordination (signals promoting uniform ripening within clusters despite initial variation), though the extent of compensation varies substantially among cultivars and conditions. The factors affecting ripening synchrony include vine balance (balanced vines with adequate leaf area supporting uniform ripening across fruit populations), water status (consistent water supply promoting uniform ripening rates), temperature uniformity (avoiding hot spots that accelerate local ripening), and crop load (appropriate crop levels enabling sufficient resource supply to all fruit), with these factors determining whether conditions support synchronous maturation or exacerbate developmental differences. Understanding ripening synchrony quality enables assessment of the relationship between véraison and harvest uniformity, prediction of whether early-season asynchrony will persist or resolve, and management interventions that may promote more uniform ripening across initially heterogeneous berry populations.
Ripening_Synchrony_Quality addresses the coordination of maturation progress determining whether véraison asynchrony is maintained or compensated during ripening. This quality determines the relationship between early-season and harvest-time uniformity. The class supports harvest uniformity prediction through ripening synchrony characterization across berry subpopulations.
Reifesynchronie
Reifungskoordination
Reifungsuniformität
Zuckerakkumulationssynchronie
maturation uniformity
maturity convergence
ripening coordination
sugar accumulation synchrony
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 277-316 Sadras VO, Petrie PR (2011) Climate shifts in south-eastern Australia: early maturity of Chardonnay, Shiraz and Cabernet Sauvignon is associated with early onset rather than faster ripening. Australian Journal of Grape and Wine Research 17(2): 199-205
Ripening Synchrony Quality
Ripening timing spans an 8-10 week range across V. vinifera cultivars in any given region, from very early Chasselas (harvest late August in central Europe) to very late Mourvèdre (harvest late October). The veraison-to-harvest interval ranges from 35-45 days in very early cultivars to 65-80 days in very late cultivars under identical thermal conditions. Winkler's heat summation classification correlates ripening timing with regional suitability, with Region I (2220 GDD) accommodating late cultivars. Ripening timing shows high heritability (h² = 0.70-0.85) and moderate correlation with flowering timing (r = 0.55-0.70).
A phenological timing disposition that is the inherited capacity of a Vitis vinifera plant to reach harvest maturity at a characteristic time relative to veraison and cumulative environmental conditions, determining the calendar period when berries achieve target sugar, acid, and flavor compound concentrations for intended use.
Ripening_Timing_Disposition serves as the parent class for the five ripening period categories from very early to very late. The trait represents the integration of developmental rate from veraison through maturity, influenced by both genetic programming and environmental modulation. Ripening timing determines cultivar-climate matching, with mismatches causing either incomplete ripening (late cultivar in cool climate) or accelerated ripening with flavor compromise (early cultivar in hot climate). Traditional wine regions reflect centuries of empirical cultivar-climate matching for optimal ripening timing.
Erntereife-Timing
Reifetermin-Disposition
Reifezeitpunkt
harvest timing trait
maturation timing
ripeness timing disposition
Winkler AJ et al. (1974) General Viticulture, University of California Press pp.: 61-75 Jones GV et al. (2005) Climate change and global wine quality. Climatic Change 73:319-343
Ripening Timing Disposition
Chardonnay grafted on Teleki 5C rootstock showing root apical meristem on second-order lateral root, meristem 0.62 mm diameter 0.48 mm length comprising estimated 840 cells, microscopic examination revealing isodiametric meristematic cells 12-18 micrometers diameter with large nuclei occupying 40% cell volume and dense cytoplasm, quiescent center identified as region of 8 slowly dividing cells at meristem apex, mitotic figures observed in 14% of meristem cells indicating active division, root apical meristem covered by root cap 0.55 mm length, meristem producing derivative cells entering elongation zone forming epidermis, cortex, and stele tissues
Root apical meristem is the undifferentiated actively dividing tissue at root tip apex comprising cells that continuously produce new root cells through mitosis, characterized by small dome-shaped region typically 0.2-0.8 mm length and diameter depending on root order, containing populations of stem cells in quiescent center maintaining meristem identity and rapidly dividing transit-amplifying cells producing derivatives, showing isodiametric cells with large nuclei dense cytoplasm and minimal vacuolation reflecting meristematic character, covered and protected by root cap providing mechanical protection and gravitropic sensing, and serving as ultimate source of all root tissues producing cells that differentiate into epidermis, cortex, endodermis, pericycle, and vascular tissues as they move basipetally away from meristem, with root apical meristem demonstrating indeterminate growth capacity enabling continuous root elongation throughout growing season and potentially throughout entire vine life, responding to environmental and hormonal signals modulating division rate and meristem size affecting root growth, and representing fundamental plant meristem analogous to shoot apical meristem but producing root rather than shoot tissues enabling root system development and expansion. Root apical meristem cellular characteristics includes= containing small isodiametric cells (meristem cells typically 10-20 micrometers diameter with approximately equal dimensions in all directions), showing large nuclei (nucleus comprising 30-50% of cell volume reflecting active DNA synthesis and transcription), exhibiting dense cytoplasm (abundant ribosomes, mitochondria, and plastids creating dense cytoplasm with minimal vacuoles), demonstrating thin primary walls (cell walls remaining thin and flexible allowing continued division without secondary thickening), and revealing meristematic identity markers (specific gene expression patterns including transcription factors and cell cycle regulators characteristic of undifferentiated cells). Root apical meristem organizational structure includes= showing quiescent center (slowly dividing organizing cells at meristem center typically 4-16 cells functioning as stem cell niche), exhibiting initial cells (surrounding quiescent center rapidly dividing cells serving as stem cells for different tissue types), demonstrating histogen zones (regions producing specific tissues with dermatogen for epidermis, periblem for cortex, plerome for stele), revealing radial symmetry (meristem organized as concentric cylinders producing radial pattern of root tissues), and displaying closed meristem type (grapevine showing separate root cap and corpus meristems rather than open meristem with common initials). Root apical meristem cell division activity includes= showing continuous mitosis (meristem cells dividing regularly every 10-24 hours during active growth depending on conditions), exhibiting cell cycle phases (populations of cells in G1, S, G2, and M phases creating asynchronous division pattern), demonstrating mitotic index (typically 5-20% of meristem cells in M phase during active growth reflecting division frequency), revealing DNA synthesis (S phase cells incorporating labeled thymidine documenting active DNA replication), and producing derivative cells (meristem division creating daughter cells that exit meristem entering elongation zone and differentiation). Root apical meristem size and cell number includes= containing hundreds to thousands of cells (meristem comprising 300-3000 cells depending on root order and species), showing variable diameter (primary root meristems 0.5-0.8 mm while fine root meristems 0.2-0.4 mm reflecting root size differences), exhibiting depth variation (meristem length typically 0.3-0.8 mm from quiescent center to beginning of elongation zone), demonstrating order-related size (lower-order larger root meristems containing more cells than higher-order smaller root meristems), and revealing growth correlation (larger meristems generally producing faster elongation through higher cell production rates). Root apical meristem maintenance and regeneration includes= requiring quiescent center function (organizing cells maintaining meristem structure and stem cell populations), showing stem cell renewal (initial cells dividing asymmetrically producing one derivative and one stem cell maintaining population), demonstrating hormone regulation (auxin, cytokinins, and other hormones maintaining meristem activity and size), exhibiting damage recovery (meristem capable of regenerating if partially damaged as long as quiescent center intact), and revealing developmental plasticity (meristem size and activity adjusting to environmental conditions and resource availability). Root apical meristem hormonal regulation includes= responding to auxin (indole-3-acetic acid promoting meristem activity and cell division synthesized in shoot and transported basipetally), affected by cytokinins (promoting meristem maintenance and cell division produced in roots and transported acropetally), influenced by gibberellins (stimulating cell division and elongation affecting meristem activity), modulated by ethylene (potentially inhibiting meristem activity under stress or promoting lateral root formation), and showing brassinosteroid effects (promoting cell division and elongation contributing to meristem function). Root apical meristem environmental responses includes= demonstrating temperature sensitivity (meristem activity increasing with temperature from threshold 8-10°C to optimum 20-25°C), showing moisture dependence (adequate hydration required for meristem function with drought reducing division), revealing nutrient effects (nitrogen and phosphorus availability affecting meristem activity and size), exhibiting light independence (root apical meristem function not requiring light though dependent on photosynthate from leaves), and displaying stress responses (severe stress potentially inducing meristem dormancy protecting stem cells until conditions improve). Root apical meristem versus shoot apical meristem includes= contrasting tissue products (root apical meristem producing root tissues versus shoot apical meristem producing stems, leaves, and flowers), showing different organization (root meristem with quiescent center versus shoot meristem with central zone and peripheral zone), exhibiting distinct geometries (root meristem dome-shaped and covered versus shoot meristem conical and exposed), demonstrating divergent responses (root meristem showing positive gravitropism versus shoot showing negative gravitropism), and revealing functional specialization (root meristem optimized for penetration and absorption versus shoot meristem for light capture and reproduction).
Root_Apical_Meristem emphasizes undifferentiated dividing tissue at root apex and is distinguished from differentiated root tissues by focus on meristematic character. Includes Active_Root_Apical_Meristem and Dividing_Root_Apical_Meristem. Critical for understanding cell production, tissue differentiation origins, stem cell maintenance, indeterminate growth, hormonal regulation, and the fundamental meristematic tissue enabling continuous root development throughout vine life.
Apical root meristem
RAM
Root meristematic zone
Wurzel-Apikalmeristem
Wurzel-Meristem
Wurzelscheitel-Meristem
Malamy, J. E. (2005). Intrinsic and environmental response pathways that regulate root system architecture. Plant, Cell & Environment, 28(1), 67-77. | Dolan, L., et al. (1993). Cellular organisation of the Arabidopsis thaliana root. Development, 119(1), 71-84.
Root Apical Meristem
Root axial conductivity assessment across root orders in Cabernet Sauvignon reveals the anatomical scaling of transport capacity, with fine roots ( 10 mm) showing 18 kg/m/s/MPa (wide vessels conveying integrated root system uptake), demonstrating how axial conductivity scales with root size to match transport demands of the hierarchical root architecture.
Root axial conductivity quality is a root hydraulic conductivity quality that represents the capacity for water transport along the length of root xylem vessels from the root tip toward the shoot, characterizing the longitudinal water flow through the vascular system that connects distributed water uptake sites to the stem xylem and is determined primarily by vessel anatomy including diameter, length, and interconnection patterns. Within the BFO framework, this quality specifically addresses the axial (parallel to root axis) component of root water transport, distinguishing xylem vessel flow from the radial cell-crossing pathway while recognizing that axial conductivity, though typically less limiting than radial conductivity in fine roots, becomes increasingly important in larger roots and can limit water supply when vessel embolism reduces functional conduit area. The ontological significance lies in root axial conductivity determining the efficiency of water transport from absorption sites to the stem, where the xylem vessel network must convey water absorbed along the entire root system with sufficient conductivity to prevent excessive water potential gradients along roots, particularly important in deep-rooted vines accessing distant soil water. The anatomical basis for axial conductivity involves vessel diameter (conductivity proportional to the fourth power of diameter according to Hagen-Poiseuille), vessel length (longer vessels reducing end-wall resistance), pit membrane properties (affecting inter-vessel water movement), and vessel network organization (redundancy and interconnection affecting resistance to dysfunction), with these features varying along roots and among root orders. The developmental patterns of root axial conductivity show young fine roots with narrow vessels and limited axial conductivity (appropriate for their short transport distances), progressively larger vessels in higher-order roots (matching increasing transport demands), and substantial axial conductivity in structural roots conveying water from the entire root system. The vulnerability of axial conductivity to embolism is significant under drought, as air entry into xylem vessels through pit membranes can spread among interconnected conduits, progressively reducing functional conducting area and creating hydraulic limitations that compound as drought intensifies. Understanding root axial conductivity quality enables assessment of the xylem limitation on root water transport, evaluation of root anatomical adaptations for water supply, and interpretation of embolism effects on root hydraulic function under drought.
Root_Axial_Conductivity_Quality addresses the longitudinal xylem transport pathway within roots determined by vessel anatomy. This quality scales with root size and is vulnerable to embolism under drought. The class supports root hydraulic architecture assessment through axial conductivity characterization across root orders.
Kax_Wurzel
Kax_root
Wurzelxylemleitfähigkeit
axiale Wurzelleitfähigkeit
longitudinal root conductivity
longitudinale Wurzelleitfähigkeit
root vessel conductance
root xylem conductivity
Tyree MT, Zimmermann MH (2002) Xylem Structure and the Ascent of Sap, 2nd Ed. Springer, Berlin, pp. 1-283 Pratt RB et al. (2007) Xylem transport safety and efficiency differ among grapevine rootstocks. American Journal of Botany 94(9): 1566-1571
Root Axial Conductivity Quality
Syrah grafted on 110 Richter rootstock showing root cap on primary root tip, cap 1.18 mm length covering apical meristem, dome-shaped structure comprising central columella 0.64 mm with statolith-containing cells, lateral root cap flanks providing mechanical protection, peripheral border cells secreting mucilage and detaching, microscopic examination revealing 11-13 cell layers from meristem to surface, columella cells showing abundant amyloplasts sedimenting in response to gravity, root cap enabling soil penetration while protecting meristem and providing gravitropic sensing
Root cap is protective tissue covering root apical meristem comprising specialized cells forming dome-shaped structure 0.3-1.5 mm length depending on root order, characterized by expendable cells continuously produced by meristem and sloughed from surface as damaged, showing three distinct regions including central columella with gravity-sensing statoliths, lateral flanks providing mechanical protection, and peripheral border cells secreting mucilage and detaching, serving critical functions including mechanical protection of delicate meristem from soil abrasion, gravitropic sensing through statolith-containing columella cells directing root growth downward, mucilage secretion lubricating root passage through soil, and chemical defense through border cell production potentially deterring pathogens, with root cap demonstrating unique developmental strategy of programmed cell production and death creating renewable protective barrier maintained throughout root growth, showing typical organization of 5-15 cell layers from meristem to surface with progressive cell maturation and eventual detachment, and representing essential root apex structure enabling soil penetration while protecting growth-critical meristem tissues. Root cap structural organization includes= showing dome-shaped geometry (cap forming rounded structure covering meristem optimizing protection and soil penetration), exhibiting central columella (core region containing gravity-sensing cells organized in files), demonstrating lateral root cap (flanking tissue providing peripheral protection and mechanical strength), displaying peripheral border cells (outermost cells programmed for detachment creating defensive and lubricating interface), and revealing layered organization (5-15 concentric cell layers from meristem to surface reflecting developmental progression). Root cap cellular characteristics includes= showing large vacuolated cells (cap cells typically 20-50 micrometers with large central vacuoles), exhibiting thick walls (cap cell walls often thicker than meristem cells providing mechanical strength), demonstrating variable amyloplast content (columella cells rich in starch-filled amyloplasts serving as statoliths while lateral cap cells fewer), revealing mucilage production (cap cells synthesizing and secreting polysaccharide mucilage particularly border cells), and displaying programmed senescence (cap cells following developmental program leading to death and sloughing after brief functional period). Root cap production and turnover includes= showing continuous regeneration (root cap initials in meristem continuously producing new cap cells replacing lost cells), exhibiting basipetal movement (newly formed cap cells moving distally away from meristem toward surface and eventual sloughing), demonstrating cell addition (meristem adding new cap cells at proximal face maintaining cap despite distal losses), revealing sloughing rate (cap losing cells from surface at rate balancing production maintaining relatively stable cap size), and displaying rapid turnover (individual cap cells living only days to weeks from production to sloughing creating dynamic renewable structure). Root cap protective function includes= providing mechanical protection (cap cells absorbing impact and abrasion protecting delicate meristem from soil particle damage), exhibiting expendable design (cap cells programmed for death and loss creating protective barrier that can be damaged without compromising meristem), demonstrating penetration facilitation (cap geometry and lubrication enabling root tip to push through dense or compacted soil), revealing pathogen defense (border cells and mucilage potentially deterring or trapping microbial pathogens protecting meristem), and displaying chemical protection (cap cells potentially accumulating defensive compounds deterring herbivores or pathogens). Root cap gravitropic function includes= showing statolith sedimentation (amyloplasts in columella cells settling in response to gravity providing directional signal), exhibiting downward growth control (gravity sensing in cap triggering auxin redistribution causing differential growth directing root downward), demonstrating central position (gravity sensors in cap center providing accurate directional information), revealing rapid response (cap sensing gravity reorientation within minutes triggering growth adjustments within hours), and displaying essential role (cap removal eliminating gravitropic response demonstrating cap necessity for directional control). Root cap mucilage secretion includes= showing polysaccharide production (cap cells particularly border cells synthesizing complex polysaccharides secreted as mucilage gel), exhibiting lubrication function (mucilage reducing friction between root tip and soil facilitating penetration), demonstrating soil modification (mucilage binding soil particles creating structured rhizosheath around root), revealing water relations (mucilage potentially helping maintain root tip hydration in drying soil), and displaying defensive roles (mucilage possibly trapping pathogens or toxic ions protecting root). Root cap in root growth includes= enabling soil penetration (cap structure and function allowing root to push through soil obstacles), directing growth trajectory (cap gravitropic sensing maintaining appropriate root orientation), demonstrating meristem protection (cap allowing meristem to function in harsh soil environment), revealing growth requirement (functional cap necessary for normal root elongation and development), and displaying continuous function (cap maintained throughout root tip life continuously produced and renewed).
Root_Cap emphasizes protective covering tissue over meristem and is distinguished from meristem proper by focus on protective specialized function. Includes Columella_Cells, Lateral_Root_Cap, and Border_Cells. Critical for understanding meristem protection, gravitropic sensing, mucilage production, soil penetration, expendable design, and the renewable protective structure covering root apex enabling growth in soil while protecting growth-critical meristem.
Kalyptra
Protective root covering
Root apex cap
Wurzelhaube
Wurzelkappe
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Barlow, P. W. (2003). The root cap= Cell dynamics, cell differentiation and cap function. Journal of Plant Growth Regulation, 21(4), 261-286.
Root Cap
Chardonnay grafted on Teleki 5C rootstock showing root cap transition zone on primary root tip, transition zone located 0.32 mm from root apex at boundary between root cap and apical meristem, microscopic examination revealing complex cellular organization with columella cells of cap transitioning to corpus initials of meristem, cell division patterns showing transition zone producing both cap-destined cells moving distally and meristem-destined cells moving basipetally, histological staining revealing molecular markers distinguishing cap from meristem with transition zone at boundary, transition zone maintaining functional separation between disposable protective cap and permanent growing apex
Root cap transition zone is the boundary region between root cap and root apical meristem where cap initials diverge from corpus meristem, typically 0.2-0.4 mm from root apex showing complex cellular organization, characterized by cells at interface between cap-producing cells and corpus tissue-producing cells, showing histological distinction between outer cap cells moving distally sloughing eventually and inner meristem cells moving basipetally differentiating into root body tissues, demonstrating shared developmental origin from common progenitor cells in transition zone before divergence into cap or corpus fates, and representing critical organizational region maintaining separation between protective disposable root cap and permanent growing root apex while enabling coordinate development of both structures, with root cap transition zone showing complex cell division patterns producing both cap and meristem derivatives, differential cell fate determination through positional signals, and structural boundary marking division between cap and corpus maintaining distinct identities despite cellular continuity and common origin. Root cap transition zone position and structure includes= located at meristem-cap boundary (transition zone positioned 0.2-0.4 mm from root apex at interface between cap and corpus), showing variable organization (some species showing distinct boundary layer while others showing gradual transition), exhibiting columella connection (transition zone including columella base connecting to quiescent center), demonstrating cellular complexity (transition zone cells showing intermediate characteristics between cap and meristem), and revealing organizational importance (transition zone maintaining functional separation between protective cap and growing meristem). Root cap transition zone cell fate divergence includes= showing common progenitor cells (transition zone cells derived from divisions that can produce either cap or corpus derivatives), exhibiting fate determination (positional signals determining whether daughter cells join cap or meristem populations), demonstrating asymmetric divisions (divisions in transition zone potentially creating one cap-destined and one corpus-destined daughter), revealing fate flexibility (transition zone potentially showing some plasticity in cell fate responding to damage or signals), and displaying boundary maintenance (transition zone mechanisms ensuring appropriate allocation of cells to cap versus corpus fates). Root cap transition zone developmental coordination includes= showing synchronized division (transition zone coordinating cap and meristem cell production maintaining appropriate proportions), exhibiting growth balance (cap production matching cap loss through sloughing maintaining stable cap size), demonstrating meristem maintenance (transition zone ensuring adequate corpus cell production maintaining meristem size and activity), revealing signal exchange (communication between cap and meristem coordinating development), and displaying functional integration (transition zone enabling cap and meristem to function as integrated apex unit). Root cap transition zone in closed meristem type includes= showing separate initials (grapevine transition zone with distinct cap initials and corpus initials not sharing common tier), exhibiting organizational complexity (closed meristem with clear transition zone boundary between cap and corpus more complex than open meristem), demonstrating independent maintenance (cap and corpus initials maintaining separate populations with transition zone managing interface), revealing evolutionary variation (closed versus open meristem types showing different transition zone organization), and displaying developmental implications (closed meristem transition zone requiring more complex regulation maintaining separate populations). Root cap transition zone boundary markers includes= showing molecular distinctions (transition zone potentially showing different gene expression in cap versus corpus sides marking boundary), exhibiting structural differences (cell wall properties or cytoskeletal organization potentially differing across transition zone), demonstrating developmental markers (specific transcription factors or proteins marking cap versus meristem cell identities), revealing boundary signaling (signals at transition zone potentially enforcing fate separation), and displaying histological visibility (transition zone sometimes visible in sections as distinct boundary between cap and meristem). Root cap transition zone damage response includes= showing regeneration capacity (transition zone potentially compensating for cap or meristem damage through adjusted cell production), exhibiting repair mechanisms (transition zone cells potentially dedifferentiating or adjusting fates to repair damage), demonstrating developmental plasticity (transition zone showing flexibility responding to perturbations), revealing quiescent center role (quiescent center adjacent to transition zone maintaining stem cells enabling regeneration), and displaying recovery patterns (damaged root tips showing recovery through transition zone and meristem reorganization). Root cap transition zone functional significance includes= maintaining cap integrity (transition zone ensuring continuous cap production replacing sloughed cells), enabling meristem function (transition zone supporting meristem maintenance and activity), demonstrating organizational importance (transition zone maintaining proper root apex architecture), revealing developmental control (transition zone coordinating complex apex patterning and growth), and displaying evolutionary conservation (transition zone organization showing conservation across plant species reflecting fundamental importance).
Root_Cap_Transition_Zone emphasizes boundary region between cap and meristem and is distinguished from cap or meristem proper by focus on interface organization. Includes Meristem_Cap_Transition_Zone and Boundary_Transition_Zone. Critical for understanding root apex organization, cap-meristem separation, cell fate divergence, developmental coordination, and the organizational boundary maintaining distinct cap and meristem identities while coordinating apex development.
Apex-Übergangsregion
Cap-corpus interface
Meristem-Haube-Grenze
Meristem-cap boundary
Root apex transition
Wurzelhaube-Übergangszone
Root Cap Transition Zone
Detailed analysis of Root_Chloride_Exclusion_Disposition at CSIRO Adelaide employed short-term ³⁶Cl⁻ uptake assays with excised roots to characterize chloride influx kinetics across 10 rootstocks, separating root uptake from subsequent translocation processes. Concentration-dependent uptake was measured over the range 0.5-50 mM Cl⁻ to determine Vmax (maximum uptake rate) and Km (affinity) parameters. Root_Chloride_Exclusion_Disposition showed moderate variation among rootstocks. Chloride uptake followed Michaelis-Menten kinetics in all rootstocks, indicating transporter-mediated entry. '140 Ruggeri' showed relatively low uptake capacity (Vmax = 2.8 μmol g⁻¹ root FW h⁻¹, Km = 12 mM), while 'Schwarzmann' displayed higher uptake (Vmax = 5.2 μmol g⁻¹ h⁻¹, Km = 8 mM), representing approximately 2-fold difference in Root_Chloride_Exclusion_Disposition. Interestingly, this 2-fold difference in root uptake translated to only partial explanation of the 20-fold difference in leaf chloride accumulation between these rootstocks, indicating that xylem loading restriction contributes more substantially to whole-plant exclusion. Apoplastic bypass contribution was assessed using PTS dye (apoplastic tracer), revealing that 15-25% of chloride entry occurred via apoplast in all rootstocks, explaining why root symplastic exclusion cannot achieve complete protection. Gene expression analysis showed that low-uptake '140 Ruggeri' had reduced expression of VvNPF2.4 (a nitrate/chloride transporter) in root epidermis compared to 'Schwarzmann', suggesting molecular basis for Root_Chloride_Exclusion_Disposition differences.
A Root_Chloride_Exclusion_Disposition is a Chloride_Exclusion_Disposition that inheres in Vitis root tissues and is realized through mechanisms at the root surface and cortex that restrict the initial uptake of chloride ions from the soil solution, reducing the chloride load presented to subsequent exclusion barriers. This disposition represents the first line of defense against chloride entry into the plant, operating at the soil-root interface where chloride must cross the epidermis and cortex to reach the stele. Chloride uptake into root cells occurs through both passive channels (when electrochemical gradients favor entry) and active transporters, with the relative contribution depending on external chloride concentration and membrane potential. Root_Chloride_Exclusion_Disposition involves reduced density or activity of plasma membrane chloride channels in root epidermal and cortical cells, potentially combined with enhanced efflux transporter activity returning chloride to the soil solution. However, because chloride can also enter roots via the apoplastic pathway (through cell walls and intercellular spaces) until blocked by the Casparian strip at the endodermis, root surface exclusion provides incomplete protection when apoplastic bypass occurs through Casparian strip discontinuities or damaged root regions. Root_Chloride_Exclusion_Disposition thus represents a contributing rather than sufficient mechanism for whole-plant chloride exclusion, with xylem loading restriction providing a critical second barrier.
Root_Chloride_Exclusion_Disposition is_a Chloride_Exclusion_Disposition. Root uptake restriction provides the first barrier against chloride entry but is incomplete due to apoplastic bypass. Uptake kinetics (Vmax, Km) quantify transporter-mediated entry. Root uptake differences explain only a portion of whole-plant exclusion variation, indicating that xylem loading restriction is equally or more important. Candidate transporters include NPF family members and anion channels. Apoplastic entry through Casparian strip gaps limits the effectiveness of symplastic exclusion mechanisms.
Kortikale-Chlorid-Barriere
Wurzel-Chloridausschluss-Disposition
Wurzelaufnahme-Restriktion
cortical chloride exclusion
root chloride barrier
root uptake restriction
Teakle NL, Tyerman SD (2010) Mechanisms of Cl⁻ transport contributing to salt tolerance, Plant Cell and Environment 33:566-589 White PJ, Broadley MR (2001) Chloride in soils and its uptake and movement within the plant: a review, Annals of Botany 88:967-988 Gong H et al. (2011) Mechanisms of chloride transport in grapevines, Functional Plant Biology 38:1-9
Root Chloride Exclusion Disposition
Systematic evaluation of Root_Cold_Hardiness_Disposition at South Dakota State University assessed root freezing tolerance across 20 rootstock genotypes and own-rooted cultivars using controlled freezing of excavated root samples. Root segments (5-10 mm diameter) were frozen to temperatures from -5°C to -20°C in 2.5°C increments, with viability assessed by electrolyte leakage and vital staining (TTC reduction). Root_Cold_Hardiness_Disposition showed strong genotypic variation correlated with species background. V. vinifera roots ('Riesling' own-rooted) exhibited LT50 = -6.5°C, substantially less hardy than the same cultivar's cane phloem (LT50 = -26°C), demonstrating the characteristic 15-20°C hardiness deficit of roots versus shoots. American species rootstocks showed enhanced Root_Cold_Hardiness_Disposition, with V. riparia 'Gloire' at LT50 = -14°C, '3309C' (V. riparia × V. rupestris) at LT50 = -12°C, and cold-selected 'Minnesota 1211' at LT50 = -18°C. Soil temperature monitoring across 5 winters revealed that root zone temperatures (30 cm depth) remained above -8°C even when air temperatures reached -35°C under adequate snow cover, but dropped to -14°C during a snow-free January period, causing 40% root mortality in V. vinifera own-rooted vines while V. riparia-based rootstocks showed less than 10% damage. These findings demonstrate that Root_Cold_Hardiness_Disposition is normally non-limiting due to soil thermal buffering, but becomes critical under inadequate insulation conditions, favoring cold-hardy rootstock selection in continental climates.
A Root_Cold_Hardiness_Disposition is a Midwinter_Hardiness_Disposition that inheres in the root system of a Vitis plant and is realized through the freezing tolerance of root tissues, determining survival of the below-ground vine structure that anchors the plant, absorbs water and nutrients, stores carbohydrates, and in grafted vines, defines the rootstock genotype contribution to vine performance. This disposition represents a distinct and generally lower cold hardiness compared to above-ground tissues, as roots do not experience the same acclimating conditions (short photoperiods, gradual temperature decline) that induce maximum hardiness in shoots and buds, and are normally protected by soil thermal buffering. Root cold hardiness in grapevine is typically 8-15°C less than corresponding shoot tissue hardiness within the same genotype, with root LT50 values ranging from approximately -5°C to -12°C for V. vinifera roots to -12°C to -20°C for cold-hardy American species roots. Under normal soil conditions, thermal buffering, snow cover, and soil moisture maintain root zone temperatures well above lethal thresholds even when air temperatures are extreme. However, root cold damage can occur when soil freezes deeply during snow-free periods, in exposed sandy soils with poor thermal buffering, or in container-grown nursery stock. Root_Cold_Hardiness_Disposition is particularly important in rootstock selection for cold climates, as rootstock root hardiness determines the below-ground survival threshold independent of scion hardiness, and root death eliminates the entire vine regardless of above-ground tissue condition.
Root_Cold_Hardiness_Disposition is_a Midwinter_Hardiness_Disposition. Roots are substantially less cold hardy than above-ground tissues (8-15°C differential) because they do not experience acclimating conditions and are normally protected by soil thermal buffering. Root hardiness becomes limiting only when soil freezing exceeds thermal buffering capacity (snow-free periods, exposed sandy soils). Rootstock genotype determines root hardiness in grafted vines independent of scion. Root death eliminates the entire vine regardless of scion tissue condition, making root hardiness the ultimate survival threshold under extreme soil freezing conditions.
Wurzel-Kältehärte-Disposition
Wurzelfrosthärte
Wurzelsystem-Winterhärte
below-ground cold hardiness
root freezing tolerance
root system winter hardiness
Howell GS, Shaulis NJ (1980) Factors influencing within-vine variation in the cold resistance of cane and primary bud tissues, American Journal of Enology and Viticulture 31:158-161 Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235 Goffinet MC (2004) Anatomy of grapevine winter injury and recovery, Grape Research News 15:1-7
Root Cold Hardiness Disposition
Quantitative assessment of Root_Density_Disposition at Washington State University compared root length density profiles across 10 rootstocks using soil coring with image analysis to quantify root length in undisturbed samples. Cores were collected at 10 cm depth intervals to 1.5 m at distances of 30, 60, and 90 cm from the vine trunk, enabling three-dimensional mapping of root distribution. Root_Density_Disposition showed marked rootstock variation. '101-14 Mgt' exhibited high root density with peak values of 3.8 cm cm⁻³ in the 10-30 cm depth zone at 30 cm from trunk, and mean density of 2.1 cm cm⁻³ across the upper 50 cm. '110 Richter' showed lower density (peak 1.8 cm cm⁻³, mean 1.2 cm cm⁻³ in upper 50 cm) but greater depth distribution. Fine root fraction (< 1 mm diameter) analysis revealed that high-density '101-14 Mgt' invested 65% of root length in fine roots versus 48% for '110 Richter', explaining density differences through proliferation strategy. Soil water depletion monitoring during a controlled drought showed that high-density '101-14 Mgt' depleted upper soil water more rapidly (reaching -1.5 MPa soil water potential in 12 days versus 18 days for '110 Richter'), but then experienced sharper stress increase as this zone was exhausted while '110 Richter' transitioned to deeper extraction. These findings illustrate how Root_Density_Disposition creates distinct water extraction patterns with different implications for drought timing and severity.
A Root_Density_Disposition is a Root_System_Drought_Adaptation_Disposition that inheres in a Vitis plant root system and is realized through the production of high root length density (root length per unit soil volume), enhancing the capacity to extract water from exploited soil zones through intensive exploitation rather than extensive exploration. This disposition represents a drought adaptation strategy based on thorough extraction of available water within the rooted soil volume, complementary to deep rooting which extends the accessible volume. High root density reduces the distance water must travel through soil to reach root surfaces, increases total absorptive surface area, and enables more complete extraction of soil water including water held at higher matric potential in smaller pores. Root density varies among rootstocks from less than 1 cm root length per cm³ soil to over 4 cm cm⁻³ in the most densely rooted zones, with substantial implications for soil water depletion patterns and drought response. Root_Density_Disposition may involve different balances between fine roots (high surface area for absorption but short-lived and metabolically costly) and coarser roots (lower absorption efficiency but greater longevity), affecting both water uptake capacity and carbon costs of root system maintenance. High root density can be advantageous for exploiting brief rainfall events that wet only surface soils, but may exhaust local soil water rapidly during extended drought if not combined with deep rooting accessing additional water reserves.
Root_Density_Disposition is_a Root_System_Drought_Adaptation_Disposition. Root density determines water extraction intensity within rooted volume, complementing deep rooting which extends accessible volume. High density favors rapid exploitation of available water but may accelerate local depletion. V. riparia parentage tends toward higher density, shallow systems while V. berlandieri tends toward lower density, deeper systems. Root density is expressed as length per volume (cm cm⁻³) or mass per volume (g cm⁻³). Fine root proliferation capacity is a major component of root density.
Bodenvolumen-Erschließung
Wurzeldichte-Disposition
Wurzellängendichte
root length density
root proliferation capacity
soil volume exploitation
Mapfumo E et al. (1994) Grapevine root distribution and water relations in a clay loam soil, Australian Journal of Grape and Wine Research 1:77-83 Morlat R, Jacquet A (2003) Grapevine root system and soil characteristics in a vineyard maintained long-term with or without interrow sward, American Journal of Enology and Viticulture 54:1-7 Bauerle TL et al. (2008) Root foraging in response to heterogeneous soil moisture in two grapevines that differ in potential growth rate, New Phytologist 179:857-866
Root Density Disposition
Root freezing tolerance quality comparison across commercial rootstock genotypes evaluated the contribution of root cold hardiness to whole-vine winter survival in continental climate viticulture where soil freezing is common. Controlled freezing tests on dormant root sections using electrolyte leakage and regrowth assessment quantified root freezing tolerance quality across the rootstock range. Riparia Gloire, derived from Vitis riparia native to northern regions, demonstrated excellent root freezing tolerance quality with LT50 at -12°C, providing a 7-8°C safety margin above typical soil freezing temperatures at root zone depths in most viticultural situations. 3309C (V. riparia × V. rupestris) showed good root freezing tolerance quality with LT50 at -10°C. SO4 (V. berlandieri × V. riparia) exhibited intermediate root freezing tolerance quality with LT50 at -8°C. 110R (V. berlandieri × V. rupestris), derived from southern-adapted species, showed poor root freezing tolerance quality with LT50 at only -6°C, creating potential vulnerability where soil temperatures might reach this level during severe cold events with absent snow cover. The root freezing tolerance quality assessment established rootstock selection guidelines for cold-climate regions, with riparia-derived rootstocks recommended where soil freezing below -8°C is probable, while rupestris and berlandieri rootstocks required supplemental protection through mulching or hilling in extreme cold situations.
Root freezing tolerance quality is a tissue freezing tolerance quality that inheres in grapevine roots and represents the capacity of root tissues to survive freezing temperatures, with roots characteristically showing substantially lower cold tolerance than equivalent above-ground tissues due to their evolution in the thermally buffered soil environment where temperatures rarely reach the extremes experienced in above-ground air, making root freezing tolerance a critical but often overlooked component of cold hardiness that becomes limiting when soil temperatures fall to damaging levels during severe cold events or in situations where soil thermal protection is inadequate. This quality addresses the cold vulnerability of the below-ground vine portion that is normally protected by soil thermal mass and insulation but may be exposed to damaging temperatures under specific conditions including shallow planting, soil displacement from cultivation, absence of snow cover on frozen soils, or extreme cold events penetrating to root zone depths. The ontological significance of root freezing tolerance quality lies in its role as a potential weak link in vine cold hardiness when soil conditions permit root exposure to temperatures within the damaging range, with root death potentially causing vine mortality even when above-ground tissues survive an extreme cold event, and with the hidden nature of root damage often delaying recognition until spring growth failure reveals the extent of below-ground injury. The biological basis of root freezing tolerance encompasses the same extracellular freezing tolerance mechanisms operating in above-ground woody tissues but achieving lower maximum tolerance due to the absence of evolutionary selection pressure for extreme cold hardiness in the thermally buffered soil environment, combined with the year-round metabolic activity of roots that limits the seasonal dormancy and hardening that enhances above-ground tissue tolerance. The variation in root freezing tolerance among rootstock genotypes reflects the geographic origins of the parent species, with rootstocks derived from northern-adapted Vitis riparia showing greater root hardiness than those from southern species, providing a mechanism for enhancing whole-vine cold tolerance through rootstock selection in regions where soil freezing risk exists.
Root_Freezing_Tolerance_Quality is_a Tissue_Freezing_Tolerance_Quality, representing the cold hardiness of below-ground root tissues normally protected by soil thermal buffering. This class addresses the often-overlooked weak link in vine cold hardiness when soil freezing occurs. Roots show 10-15°C lower hardiness than equivalent above-ground tissues due to evolutionary selection in buffered soil environment. V. riparia-derived rootstocks provide superior root hardiness (-12°C LT50) versus southern species rootstocks (-6°C). Soil conditions (snow cover, mulch, planting depth) strongly modulate root exposure to damaging temperatures. Hidden nature of root damage may delay recognition until spring growth failure.
Wurzel-Gefrier-Toleranz
Wurzel-Kältehärte
Wurzelsystem-Winterüberleben
below-ground freezing tolerance
root cold hardiness
root frost resistance
root system winter survival
unterirdische Frosttoleranz
Root Freezing Tolerance Quality
Root growth limitation quality assessment for phosphorus deficiency in a struggling newly planted Sonoma County Pinot Noir vineyard identified inadequate root development as the primary cause of poor establishment on a site with historically low phosphorus availability. Excavation of representative vines 18 months after planting revealed root growth limitation quality with affected vines showing total root length of only 8 meters versus 25 meters in normally establishing vines of equivalent age, representing 68% reduction in root system size. Root architectural analysis showed root growth limitation quality manifesting as reduced primary root extension (maximum rooting depth 35 cm versus 75 cm expected), decreased lateral root branching with only 3.2 laterals per cm primary root versus 5.8 in normal vines, and root system confined to the immediate planting hole rather than exploring surrounding native soil. Root-to-shoot ratio analysis revealed that despite overall growth limitation, root growth limitation quality resulted in relatively greater root allocation (R/S ratio 0.45) than normal vines (R/S ratio 0.35), reflecting the plant's attempt to enhance phosphorus acquisition through increased root investment. Phosphorus tissue analysis of excavated roots confirmed the diagnosis with root P concentration at 0.08% versus sufficiency range of 0.15-0.25%. Targeted correction with triple superphosphate banded 20 cm from vines followed by overhead irrigation to move phosphorus into the root zone stimulated new root growth, with follow-up assessment at 6 months showing 180% increase in root length density in the amended zone and above-ground growth recovery confirming that root growth limitation quality had been the primary constraint on vine establishment.
Root growth limitation quality in phosphorus deficiency is a phosphorus deficiency symptom quality that inheres in grapevine root systems and represents the impaired root development and reduced root system size resulting from inadequate phosphorus availability for the DNA synthesis, cell division, and energy metabolism required for root meristem activity and root cell expansion, with phosphorus-limited root growth creating a negative feedback where reduced root exploration capacity further limits phosphorus acquisition from soil, exacerbating the deficiency. This quality captures the below-ground dimension of phosphorus deficiency that underlies many of the above-ground symptoms, recognizing that the high phosphorus demand of dividing root tip cells and the role of roots in nutrient acquisition make root growth particularly sensitive to phosphorus status. The ontological significance of root growth limitation quality in phosphorus deficiency lies in its role as both a symptom of deficiency and a driver of continued limitation through reduced soil exploration, creating a self-reinforcing cycle that can impair vine establishment and long-term productivity if not corrected through phosphorus application or soil amendment. The biological basis of phosphorus-limited root growth encompasses the concentration of cell division activity in root apical meristems where DNA and RNA synthesis create high phosphorus demand, the requirement for ATP in the energy-intensive processes of cell expansion and nutrient uptake, and the phospholipid requirements for membrane synthesis in rapidly growing tissues. The architectural changes in phosphorus-deficient root systems include reduced total root length, decreased branching frequency, altered root-to-shoot ratio often favoring root allocation as plants attempt to acquire limiting phosphorus, and modified root hair development that may increase phosphorus uptake efficiency per unit root length. The practical assessment of root growth limitation requires excavation or soil coring to evaluate root development, making this symptom more difficult to diagnose than above-ground symptoms but essential for understanding vine establishment problems and nutrient acquisition capacity.
Root_Growth_Limitation_Quality_(P) is_a Phosphorus_Deficiency_Symptom_Quality, representing the impaired root development resulting from P-limited cell division and energy metabolism. This class addresses the below-ground dimension underlying above-ground symptoms. High P demand of root meristems makes roots particularly sensitive to P status. The negative feedback (less root → less P acquisition → less root) creates self-reinforcing deficiency. Architectural changes include reduced length, branching, and depth. Practical assessment requires excavation, making diagnosis more difficult than foliar symptoms. Critical for vine establishment where root development determines long-term productivity.
P-Stress-Wurzellimitierung
P-limited root development
P-limitierte Wurzelentwicklung
P-stress root limitation
Wurzelwachstumslimitierung bei P-Mangel
eingeschränktes Wurzelwachstum
phosphorus deficiency root effects
restricted root growth
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Grant CA et al. (2001) The importance of early season phosphorus nutrition. Canadian Journal of Plant Science 81:211-224 Vance CP et al. (2003) Phosphorus acquisition and use: critical adaptations by plants for securing a nonrenewable resource. New Phytologist 157:423-447
Root Growth Limitation Quality (P)
Chardonnay grafted on SO4 rootstock showing root hairs on fourth-order fine root, hairs visible as white fuzz on root surface, microscopic examination revealing root hairs 0.45 mm average length 14 micrometers diameter, cylindrical single-cell extensions from trichoblast epidermal cells, thin primary walls facilitating soil contact, hair density 180 per centimeter root length, hairs initiating 6.2 mm from root tip continuing through maturation zone, root hairs increasing effective root surface area 11-fold, hairs providing estimated 68% of root water uptake and 72% of nitrogen uptake
Root hair is tubular extension of single epidermal cell (trichoblast) projecting outward from root surface, typically 0.1-1.5 mm length and 10-20 micrometers diameter creating cylindrical protrusion, characterized by thin primary cell wall allowing close contact with soil particles, single continuous cytoplasm connecting hair to parent cell body enabling direct transport, tip growth mechanism where extension occurs only at hair apex through polarized vesicle delivery, and very short lifespan typically 2-10 days reflecting ephemeral nature, serving critical function in dramatically increasing root absorption surface area 5-20 fold enabling efficient water and nutrient uptake from soil micro-sites, with root hairs showing typical density 50-300 per centimeter root length depending on species and conditions, initiating 3-10 mm behind root apex in maturation zone after cell elongation completes, growing rapidly 20-100 micrometers per hour for 1-2 days reaching final length then maintaining function briefly before senescing, and demonstrating that despite individual hairs being microscopic and ephemeral, collective root hair population provides majority of root system absorption surface making root hairs essential for vine water and nutrient acquisition. Root hair morphology includes= showing cylindrical shape (root hair typically uniform diameter throughout length creating tube-like structure), exhibiting single-cell origin (entire hair being extension of one epidermal trichoblast cell), demonstrating thin wall (primary cell wall only 0.05-0.15 micrometers thick facilitating close soil contact), displaying cytoplasmic continuity (hair cytoplasm continuous with parent cell enabling direct transport without membrane barriers), and revealing vacuole organization (small vacuoles in hair versus large central vacuole in cell body). Root hair dimensions includes= showing typical length 0.1-1.5 mm (most root hairs 0.3-0.8 mm though variable with species, conditions, and age), exhibiting diameter 10-20 micrometers (relatively constant diameter along length), demonstrating high aspect ratio (length typically 20-80 times diameter creating elongated structure), displaying variable length (root hair length varying with soil conditions, nutrients, and mechanical impedance), and revealing order effects (fine roots showing shorter denser hairs than coarse roots). Root hair initiation and development includes= showing trichoblast specification (specific epidermal cells developing into root hair-forming trichoblasts typically overlying cortical cell anticlinal walls), exhibiting bulge formation (trichoblast developing small protrusion marking hair initiation site), demonstrating tip growth (hair elongating exclusively at apex through polarized delivery of membrane and wall materials), revealing rapid extension (growing hairs elongating 20-100 micrometers per hour for 1-2 days), and displaying growth cessation (hairs stopping growth after 1-2 days at final length 0.3-1.5 mm). Root hair lifespan and turnover includes= showing very short lifespan (individual root hairs typically living 2-10 days from initiation to senescence), exhibiting ephemeral nature (root hairs among shortest-lived plant cells continuously produced and dying), demonstrating position-dependent age (youngest hairs near root tip, oldest near base of hair zone), revealing continuous production (trichoblasts continuously producing new hairs as root tip advances), and displaying high turnover (root hair population completely replacing itself every few days). Root hair function includes= providing absorption surface (root hairs increasing effective root surface area 5-20 fold enhancing water and nutrient uptake), enabling soil exploration (hairs extending into soil pores accessing water and nutrients in micro-sites inaccessible to root cylinder), demonstrating anchorage (hairs potentially providing some mechanical anchoring through soil particle contact), showing mucilage secretion (hairs secreting polysaccharides modifying rhizosphere), and revealing mycorrhizal exclusion (root hairs typically not colonized by arbuscular mycorrhizae possibly competing for same function). Root hair spatial distribution includes= concentrating in maturation zone (root hairs abundant 5-50 mm from root tip in zone of completed cell differentiation), showing density variation (hair density varying 50-300 per centimeter depending on conditions), exhibiting patchy distribution (hair density potentially higher in resource-rich soil zones), demonstrating root order effects (fine absorptive roots showing abundant hairs while woody structural roots showing few or none), and revealing environmental responsiveness (hair density and length responding to nutrient availability, moisture, and mechanical impedance).
Root_Hair emphasizes single-cell epidermal extension and is distinguished from root proper by focus on microscopic tubular protrusion. Includes Active_Root_Hair, Senescent_Root_Hair, and Root_Hair_Zone. Critical for understanding absorption surface increase, soil exploration, tip growth, ephemeral lifespan, and the microscopic single-cell extensions dramatically increasing root surface area enabling efficient water and nutrient acquisition.
Absorptive hair
Absorptives Haar
Epidermal hair
Trichoblast extension
Wurzelepidermis-Auswuchs
Wurzelhaar
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Peterson, R. L., & Farquhar, M. L. (1996). Root hairs= Specialized tubular cells extending root surfaces. The Botanical Review, 62(1), 1-40.
Root Hair
Root hair density assessment comparing phosphorus treatments reveals the adaptive response enhancing nutrient acquisition capacity, with P-deficient vines showing root hair density of 285 hairs/mm² (enhanced development to maximize P foraging in depletion zone) and mean length of 1.2 mm (extending absorbing surface), while P-sufficient vines show only 145 hairs/mm² with mean length 0.6 mm (reduced investment when P supply adequate), with the enhanced root hair development under deficiency increasing effective absorbing surface by approximately 3-fold and enabling more efficient phosphorus acquisition from low-P soil, demonstrating the adaptive plasticity of root hair density in response to nutrient limitation.
Root hair density quality is a root uptake capacity quality that represents the number of root hair projections per unit root surface area or length, characterizing the development of these unicellular extensions that dramatically expand the effective absorbing surface in the critical zone immediately surrounding fine roots where nutrient depletion occurs most rapidly. Within the BFO framework, this quality specifically addresses the root hair component of absorptive architecture, distinguishing root hair development from overall root surface area while recognizing that root hairs can increase effective absorbing surface by 2-5 fold in the soil zone most critical for nutrient uptake, making root hair density a key determinant of acquisition efficiency particularly for immobile nutrients. The ontological significance lies in root hairs extending the absorptive interface into the depletion zone that develops around roots as nutrients are absorbed faster than diffusion can replenish them, where typical root hair lengths of 0.1-1.5 mm enable access to soil volume and nutrient pools beyond the immediate root surface, dramatically improving acquisition of poorly mobile nutrients like phosphorus whose depletion zones extend only 1-2 mm from root surfaces. The functional significance of root hairs varies among nutrients depending on mobility, with root hairs particularly important for phosphorus acquisition (depletion zone similar to root hair length, root hairs providing major access to P), moderately important for potassium and ammonium (intermediate mobility), and less critical for highly mobile nitrate (mass flow delivery often sufficient), such that root hair development particularly affects acquisition of immobile nutrients. The regulation of root hair development responds to nutrient availability, with phosphorus deficiency often stimulating increased root hair density and length as plants enhance acquisition capacity for this limiting nutrient, while nitrogen deficiency may also promote root hair development, creating adaptive responses that improve acquisition of deficient nutrients. The environmental factors affecting root hair density include soil physical conditions (root hair development affected by soil compaction and structure), soil moisture (optimal development in moist but well-aerated conditions), and nutrient status (deficiency often stimulating development), with substantial genotypic variation in root hair density providing potential for selection and breeding for improved nutrient acquisition efficiency. Understanding root hair density quality enables assessment of the fine-scale absorptive capacity most relevant for immobile nutrient acquisition, interpretation of adaptive responses to nutrient limitation, and evaluation of genotypic variation in root hair development affecting phosphorus uptake efficiency.
Root_Hair_Density_Quality addresses the unicellular projections extending absorbing surface into the nutrient depletion zone. This quality is particularly important for immobile nutrient (P) acquisition and shows adaptive plasticity in response to deficiency. The class supports phosphorus nutrition assessment and root fine-structure evaluation through root hair density characterization.
Absorptionsoberflächenerweiterung
Trichoblastendichte
Wurzelhaarabundanz
Wurzelhaardichte
absorption surface extension
root hair abundance
root hair development
trichoblast density
Gahoonia TS, Nielsen NE (2004) Root traits as tools for creating phosphorus efficient crop varieties. Plant and Soil 260(1-2): 47-57 Ma Z et al. (2001) Morphological synergism in root hair length, density, initiation and geometry for phosphorus acquisition in Arabidopsis thaliana. Plant, Cell & Environment 24(4): 459-467
Root Hair Density Quality
Merlot grafted on 3309 Couderc rootstock showing root hair zone on fourth-order fine root, zone extending 38 mm from proximal boundary 8 mm from apex to distal boundary 46 mm from apex, zone showing gradient of hair ages from youngest 2-day hairs proximal to oldest 9-day hairs distal, maximum hair density 210 per cm at 22 mm from apex in mid-zone, hair zone demonstrating white fuzzy appearance under microscope, isotope tracing showing hair zone providing 76% of root water uptake and 82% of nitrogen uptake despite comprising only 24% of root length
Root hair zone is the region of root bearing root hairs typically extending 5-80 mm from root apex though variable with root order and conditions, characterized by presence of mature root hairs on root surface creating fuzzy or hairy appearance, showing gradient of hair ages with youngest hairs proximal near elongation zone and oldest hairs distal approaching hair zone terminus, demonstrating maximum root absorption capacity per unit root length within zone, and representing functionally critical root region where majority of water and nutrient uptake occurs through combined root and hair surfaces, with root hair zone boundaries determined by hair initiation point approximately 5-15 mm from apex where epidermal cell differentiation completes and hair senescence point typically 30-100 mm from apex where hair lifespan expired and hairs senesced or sloughed, showing typical zone length 25-60 mm in fine roots shorter in very fine roots and longer in coarse roots, and demonstrating that hair zone moves basipetally along root as root elongates maintaining relatively stable zone length despite continuous hair turnover through production at proximal end and loss at distal end. Root hair zone boundaries includes= showing proximal boundary (hair zone beginning 5-15 mm from root apex where first hairs emerge after cell elongation and early differentiation complete), exhibiting distal boundary (hair zone ending 30-100 mm from apex where hairs senesced and sloughed leaving bare root), demonstrating boundary determinants (proximal boundary set by trichoblast differentiation timing, distal boundary set by hair lifespan and turnover rate), revealing dynamic boundaries (zone boundaries moving basipetally as root grows while maintaining relatively constant positions relative to moving apex), and displaying order-dependent boundaries (fine roots showing proximal zone while coarse structural roots showing distal zone or lacking hairs entirely). Root hair zone length includes= showing typical extent (hair zone typically 25-60 mm length in fine absorptive roots depending on conditions and species), exhibiting root order effects (primary and secondary roots showing longer hair zones than tertiary and quaternary fine roots), demonstrating environmental variation (hair zone potentially longer under optimal conditions shorter under stress), revealing age-related patterns (young actively growing roots showing well-developed hair zones while old senescent roots showing reduced or absent zones), and displaying seasonal changes (hair zone characteristics potentially varying seasonally with root growth activity). Root hair zone hair density gradient includes= showing variable density along zone (hair density changing from proximal to distal reflecting age-related changes in hair population), exhibiting maximum density (typically mid-zone 15-35 mm from apex where most hairs functional and mature), demonstrating proximal density (proximal zone showing developing hairs potentially lower density as hairs still initiating), revealing distal density (distal zone potentially showing lower density as some hairs senesced and sloughed), and displaying patchy distribution (hair density potentially varying along zone reflecting local soil conditions or developmental irregularities). Root hair zone age structure includes= showing youngest hairs proximal (recently initiated hairs 1-3 days old closest to apex), exhibiting mature hairs mid-zone (hairs 3-6 days old in peak functional period in middle of zone), demonstrating old hairs distal (hairs 6-10 days old approaching or in senescence in distal zone), revealing age gradient (smooth progression of hair ages from young to old moving distally along zone), and displaying turnover dynamics (age structure reflecting continuous production and mortality maintaining functional population). Root hair zone absorption capacity includes= showing maximum uptake (hair zone providing highest water and nutrient uptake per unit root length), exhibiting disproportionate contribution (hair zone comprising 10-30% of total root length but providing 50-80% of total root uptake), demonstrating efficiency advantage (hair zone showing 5-15 fold higher uptake per centimeter than non-hair zones), revealing functional importance (hair zone absorption critical for meeting vine water and nutrient demands), and displaying stress vulnerability (hair zone function sensitive to drought, soil conditions, or root damage affecting vine resource status). Root hair zone movement along root includes= showing basipetal progression (hair zone continuously moving toward root base as root apex grows distally), exhibiting constant distance from apex (zone maintaining relatively stable position relative to moving root tip), demonstrating new hair production (continuous initiation of new hairs at proximal zone boundary as root extends), revealing distal hair loss (continuous senescence and sloughing at distal boundary as hairs exceed lifespan), and displaying stable zone length (zone length remaining relatively constant despite boundaries continuously moving basipetally). Root hair zone in different root orders includes= showing abundant hairs in fine roots (tertiary, quaternary, and higher-order roots with well-developed hair zones providing absorption), exhibiting moderate hairs in secondary roots (second-order roots showing hair zones though potentially less dense than fine roots), demonstrating sparse or absent hairs in primary roots (large structural roots showing few hairs or none particularly in older portions), revealing functional specialization (fine roots with abundant hairs optimized for absorption while coarse roots emphasize transport and structure), and displaying root system architecture (hair zone distribution across root orders creating functional differentiation).
Root_Hair_Zone emphasizes hair-bearing region on root and is distinguished from non-hair regions by focus on presence and distribution of hairs. Includes Dense_Root_Hair_Zone and Mature_Root_Hair_Zone. Critical for understanding spatial distribution, zone boundaries, age gradients, absorption concentration, dynamic movement, and the functionally critical root region bearing root hairs providing majority of absorption capacity.
Absorption zone
Absorptionszone
Behaarte Wurzelregion
Hair-bearing zone
Piliferous zone
Wurzelhaarzone
Root Hair Zone
Detailed hydraulic characterization of Root_Hydraulic_Conductance_Disposition at the University of Bordeaux measured whole-root-system hydraulic conductance (Lpr) across 12 rootstocks using high-pressure flow meter (HPFM) techniques on excavated root systems and pressurized root exudation methods in potted vines. Measurements were conducted under well-watered conditions and after 10 days of soil drying to assess conductance plasticity. Root_Hydraulic_Conductance_Disposition varied 3-fold among rootstocks under well-watered conditions. '1103 Paulsen' (V. berlandieri × V. rupestris) exhibited highest Lpr at 4.2 × 10⁻⁷ m³ s⁻¹ MPa⁻¹ per unit root surface area, while 'Riparia Gloire' showed lowest conductance at 1.4 × 10⁻⁷ m³ s⁻¹ MPa⁻¹. Following soil drying, conductance declined in all rootstocks, but the magnitude of decline differed. High-conductance '1103 Paulsen' showed 55% conductance reduction, while low-conductance 'Riparia Gloire' declined only 25%. Analysis of aquaporin gene expression (VvPIP1 and VvPIP2 families) revealed that '1103 Paulsen' down-regulated aquaporins more strongly under drought (3.5-fold reduction versus 1.8-fold in 'Riparia Gloire'), suggesting active conductance modulation. The functional significance was demonstrated by scion water status, with high-Lpr '1103 Paulsen'-grafted 'Cabernet Sauvignon' showing higher midday water potential under well-watered conditions (-0.9 versus -1.1 MPa for 'Riparia Gloire') but similar or lower potential under drought due to conductance plasticity. These findings reveal that Root_Hydraulic_Conductance_Disposition encompasses both maximum capacity and drought-induced modulation.
A Root_Hydraulic_Conductance_Disposition is a Root_System_Drought_Adaptation_Disposition that inheres in a Vitis plant root system and is realized through high efficiency of water transport from soil through roots to the shoot system, minimizing the hydraulic resistance that limits water flow from soil to leaves. This disposition represents the transport efficiency component of root function, determining how effectively acquired soil water is delivered to above-ground tissues to support transpiration, growth, and stress responses. Root hydraulic conductance reflects the combined conductance of the radial pathway (soil to xylem across root cortex, endodermis, and stele) and the axial pathway (longitudinal flow through xylem vessels to the shoot). Radial conductance is influenced by aquaporin (water channel protein) expression and activity, suberization and Casparian strip development, and root anatomical features. Axial conductance depends on xylem vessel diameter, frequency, and connectivity. Root_Hydraulic_Conductance_Disposition is dynamic, with conductance declining under severe drought through aquaporin down-regulation (potentially an adaptive response reducing water loss to dry soil) and root embolism. High hydraulic conductance enables the root system to supply water to meet transpirational demand under well-watered conditions and moderate stress, while conductance modulation may contribute to whole-plant drought signaling by allowing root water potential to decline and trigger shoot responses.
Root_Hydraulic_Conductance_Disposition is_a Root_System_Drought_Adaptation_Disposition. Root hydraulic conductance determines water delivery efficiency to shoots. Conductance has radial (cortex pathway) and axial (xylem) components. Aquaporins are major regulators of radial conductance. Conductance declines under drought through aquaporin down-regulation and embolism, potentially contributing to drought signaling. High conductance is beneficial under adequate water supply but conductance plasticity may be adaptive under drought.
Wurzel-Hydraulik-Leitfähigkeits-Disposition
Wurzel-Leitfähigkeit
Wurzel-Wassertransport-Effizienz
root conductivity
root hydraulic capacity
root water transport efficiency
Alsina MM et al. (2011) Seasonal changes of whole root system conductance by a drought-tolerant grape root system, Journal of Experimental Botany 62:99-109 Vandeleur RK et al. (2009) The role of plasma membrane intrinsic protein aquaporins in water transport through roots: diurnal and drought stress responses reveal different strategies between isohydric and anisohydric cultivars of grapevine, Plant Physiology 149:445-460 Lovisolo C et al. (2008) Whole-plant hydraulic conductance and root-to-shoot flow of abscisic acid are independently affected by water stress in grapevines, Functional Plant Biology 35:922-930
Root Hydraulic Conductance Disposition
Root hydraulic conductivity (Lp) in grapevines ranges from 1.5-6.0 × 10⁻⁷ m/s/MPa across genotypes, with high-conductivity rootstocks such as 1103 Paulsen and SO4 showing 2-3× greater Lp than low-conductivity types such as 110 Richter and 140 Ruggeri. Root Lp shows strong diurnal variation, peaking in early morning (coinciding with maximum aquaporin expression) and declining 40-60% by midday under water stress. Temperature sensitivity of root Lp follows Arrhenius kinetics with Q₁₀ values of 2.0-2.5, explaining reduced water uptake at low soil temperatures. Suberization of the exodermis and endodermis increases hydraulic resistance, with older root zones showing 50-80% lower Lp than young root tips.
A water uptake disposition that is the inherited capacity of a Vitis vinifera plant to conduct water through the root system with genotype-specific hydraulic efficiency, determined by root anatomical features (cortex structure, endodermal suberization, xylem vessel dimensions) and membrane water permeability.
Root hydraulic conductivity integrates anatomical and physiological determinants of root water transport capacity. High Lp enables rapid water uptake under transpirational demand but may increase vulnerability to drought through rapid soil water depletion. The trait shows plasticity in response to water availability, with drought-stressed roots increasing suberization and reducing Lp as a water-conserving adaptation. Rootstock-scion combinations with moderate root Lp often show superior performance under deficit irrigation by balancing uptake capacity with conservative water use.
Lp root trait
Wurzelpermeabilität
hydraulische Wurzelleitfähigkeit
root hydraulic conductance
root water permeability
Vandeleur RK et al. (2009) The role of plasma membrane intrinsic protein aquaporins in water transport through roots: diurnal and drought stress responses reveal different strategies between isohydric and anisohydric cultivars of grapevine. Plant Physiology 149:445-460 Gambetta GA et al. (2012) Water uptake along the length of grapevine fine roots: developmental anatomy, tissue-specific aquaporin expression, and pathways of water transport. Plant Physiology 158:1492-1504
Root Hydraulic Conductivity Disposition
Root hydraulic conductivity assessment comparing rootstocks under declining soil moisture reveals genotypic variation in water uptake maintenance, with 110R showing initial Lpr of 3.8 × 10⁻⁸ m/s/MPa declining by only 35% as soil water potential drops from -0.1 to -0.8 MPa (maintained aquaporin activity), while SO4 shows higher initial Lpr of 4.5 × 10⁻⁸ but 65% decline under similar drying (aquaporin downregulation), demonstrating how rootstock hydraulic conductivity responses to soil drying affect water supply maintenance and drought tolerance transferred to grafted scions.
Root hydraulic conductivity quality is a hydraulic conductivity quality that represents the capacity of root systems to absorb water from soil and transport it to the shoot, characterizing the belowground component of hydraulic architecture that often constitutes the major resistance to water flow in the soil-plant-atmosphere continuum, particularly under drought conditions when soil-root contact and aquaporin activity decline. Within the BFO framework, this quality specifically addresses the root component of hydraulic conductivity, distinguishing root water uptake and transport capacity from stem and leaf conductivities while recognizing that root hydraulic conductivity is often the most variable and environmentally responsive component of whole-plant hydraulic architecture. The ontological significance lies in root hydraulic conductivity frequently limiting whole-plant water supply, where the root system must extract water from increasingly dry soil against declining water potential gradients and transport it to the xylem, with root conductivity determining whether water supply can match shoot demand under varying soil moisture and evaporative conditions. The pathway components of root hydraulic conductivity include the soil-root interface (dependent on root-soil contact and soil hydraulic conductivity), the radial pathway from root surface to xylem (crossing cortex, endodermis, and pericycle), and the axial pathway in root xylem vessels (conveying water toward the shoot), with each component potentially limiting overall root conductivity depending on conditions. The physiological regulation of root hydraulic conductivity involves aquaporin expression and activity (dramatically affecting radial conductivity), suberization of endodermal barriers (affecting the apoplastic pathway), and root anatomical development (vessel diameter, root branching affecting axial conductivity), with these factors creating substantial plasticity in root conductivity that responds to soil conditions and plant water status. The rootstock effect on vine hydraulic architecture operates largely through root hydraulic conductivity, with rootstock genotype determining root system size, anatomy, and aquaporin expression that affect the hydraulic supply capacity grafted scion varieties experience. Understanding root hydraulic conductivity quality enables assessment of the belowground limitation on vine water relations, rootstock selection for hydraulic characteristics, and interpretation of root contribution to whole-plant water transport capacity.
Root_Hydraulic_Conductivity_Quality serves as the parent class for root pathway components (Radial, Axial, Aquaporin). This quality addresses the often-limiting belowground component of vine hydraulic architecture. The class supports rootstock evaluation and water relations assessment through root conductivity characterization.
Lpr
Wasseraufnahmekapazität
Wurzelhydraulikleitfähigkeit
Wurzelleitfähigkeit
root K
root conductance
root water uptake capacity
Steudle E (2000) Water uptake by plant roots: an integration of views. Plant and Soil 226(1): 45-56 Gambetta GA et al. (2012) Sugar and abscisic acid signaling orthologs are activated at the onset of ripening in grape. Planta 236(3): 895-910
Root Hydraulic Conductivity Quality
Dog Ridge (V. champinii) demonstrates broad-spectrum root-knot nematode resistance (Rf 0.2-0.5 across M. incognita, M. javanica, and M. arenaria) and serves as the primary resistance donor in UC Davis rootstock breeding, contributing to Freedom, Harmony, and subsequent selections. Ramsey (Salt Creek, V. champinii) provides field-proven broad-spectrum resistance in Australian and Californian viticulture where mixed Meloidogyne populations are common. RS-3 and RS-9 (USDA Fresno selections from V. champinii × V. rupestris × V. vinifera) show resistance to all major Meloidogyne species (Rf 0.3-0.6) combined with improved graft compatibility over pure V. champinii rootstocks.
A nematode-resistance rootstock role that inheres in a grafted rootstock by virtue of its capacity to resist reproduction by root-knot nematodes of the genus Meloidogyne generally, without restriction to a single species. Root-knot nematode resistance as a general category captures rootstocks selected or bred for broad-spectrum efficacy against the Meloidogyne species complex commonly encountered in viticulture, including M. incognita, M. javanica, M. arenaria, M. hapla, and occasionally M. chitwoodi and M. ethiopica. Broad-spectrum resistance is achieved through pyramiding of resistance genes from multiple genetic sources, most importantly V. champinii (carrying complementary resistance factors effective against multiple species), V. rufotomentosa, and M. rotundifolia. Resistant rootstocks prevent giant cell formation through species-nonspecific hypersensitive response mechanisms, maintaining Rf values below 1.0 across the Meloidogyne species range present at a given site.
Root_Knot_Nematode_Resistant_Rootstock_Role functions as the superclass encompassing resistance to all Meloidogyne species, while species-specific subclasses (M. incognita, M. javanica, M. arenaria) capture resistance to individual species where documented. A rootstock bearing this general role should demonstrate resistance across the major viticultural Meloidogyne species rootstocks with resistance to only one or two species are more precisely classified under species-specific subclasses. The M. hapla (northern root-knot nematode) is less commonly targeted by breeding programs as it occurs primarily in cooler climates where other Meloidogyne species are absent.
Meloidogyne resistant rootstock
Meloidogyne-resistente Unterlage
RKN resistant rootstock
broad-spectrum nematode resistant rootstock
gegen Wurzelgallenälchen resistente Unterlage
root-knot nematode resistant rootstock
wurzelgallennematodenresistente Unterlage
Cousins P, Walker MA (2002) Genetics of resistance to Meloidogyne incognita in crosses of grape rootstocks, Theoretical and Applied Genetics 105: 802-807 McKenry MV, Anwar SA (2006) Nematode and grape rootstock interactions, Journal of Nematology 38(3): 312-318 Walker MA et al. (2014) Rootstock breeding for resistance to nematodes and phylloxera, Acta Horticulturae 1045: 89-97
Root Knot Nematode Resistant Rootstock Role
Root maintenance respiration assessment in drip-irrigated versus dry-farmed Zinfandel reveals the root system size effect on carbon costs, with irrigated vines developing concentrated root systems (estimated 2.5 kg dry weight) showing root respiration of 85 g glucose/day (maintenance plus ion uptake), while dry-farmed vines with extensive deep root systems (estimated 5.2 kg dry weight) show elevated root respiration of 165 g glucose/day, with the larger carbon cost of extensive roots offset by enhanced water access during drought, demonstrating the carbon investment trade-off between root system size and resource acquisition capacity that affects whole-vine productivity.
Root maintenance respiration quality is a maintenance respiration quality that represents the rate of respiratory metabolism required to maintain root cellular structures and transport functions without net growth, characterizing the carbon cost of sustaining membrane integrity, ion transport capacity, and mycorrhizal associations in the belowground portion of the vine that represents a major respiratory sink often consuming 15-30% of daily photosynthate. Within the BFO framework, this quality specifically addresses the root component of maintenance respiration, distinguishing root maintenance costs from leaf and fruit maintenance while recognizing that roots face unique maintenance challenges including ion gradient maintenance against soil solution, membrane repair in variable soil environments, and support of mycorrhizal symbionts that enhance nutrient acquisition. The ontological significance lies in root maintenance respiration representing a substantial and often underappreciated carbon cost, where the extensive root system required for water and nutrient acquisition creates correspondingly large maintenance demand that must be met from shoot-derived photosynthate, making root maintenance a significant factor in whole-vine carbon balance and potentially limiting carbon availability for fruit development. The physiological basis for root maintenance respiration includes ion transport maintenance (replacing ions lost through membrane leakage, particularly significant for K⁺ and NO₃⁻), membrane integrity (repair and turnover in the challenging soil environment), protein turnover (maintaining transport proteins and metabolic enzymes), and mycorrhizal carbon transfer (supporting fungal symbionts that provide enhanced nutrient access). The factors affecting root maintenance respiration include root biomass (larger root systems having proportionally higher maintenance costs), soil temperature (exponential response affecting belowground respiration independently of shoot temperature), root nitrogen content (correlating with protein-dependent maintenance costs), and soil conditions (waterlogging, salinity, and other stresses potentially increasing maintenance requirements). The quantitative partitioning of root respiration between maintenance and ion uptake functions is methodologically challenging but functionally important, with ion uptake costs sometimes categorized separately from true maintenance or included within the maintenance component depending on analytical framework. Understanding root maintenance respiration quality enables assessment of the belowground carbon cost, root system efficiency evaluation, and appreciation of how root architecture and soil conditions affect whole-vine carbon allocation to the root sink.
Root_Maintenance_Respiration_Quality addresses the specific maintenance costs of the belowground root system. This quality represents a substantial carbon sink often underestimated in vine carbon budgets. The class supports whole-plant carbon balance assessment through characterization of the root maintenance component that scales with root system size.
Wurzel-Erhaltungsatmung
Wurzel-Rm
Wurzelstoffwechselkosten
belowground maintenance cost
root Rm
root basal respiration
root metabolic cost
unterirdische Instandhaltungskosten
Lambers H et al. (2008) Plant Physiological Ecology, 2nd Ed. Springer, New York, pp. 101-150 Atkin OK et al. (2000) Relationship between the inhibition of leaf respiration by light and enhancement of leaf dark respiration following light treatment. Australian Journal of Plant Physiology 27(7): 669-680
Root Maintenance Respiration Quality
Systematic quantification of Root_Phylloxera_Resistance_Disposition at INRAE Bordeaux employed standardized pot assays comparing root damage across 45 rootstock genotypes and V. vinifera controls following controlled phylloxera infestation. Potted vines were inoculated with 500 phylloxera crawlers and assessed at 16 weeks for nodosity count per gram root, tuberosity severity (0-5 scale), root system damage (0-9 OIV scale), and vine growth parameters. V. vinifera 'Cabernet Sauvignon' (own-rooted) showed severe susceptibility with 42 nodosities/g root, tuberosity rating 4.5, OIV damage score 2 (very susceptible), and 65% growth reduction. The highly resistant 'Börner' (V. riparia × V. cinerea) demonstrated exceptional Root_Phylloxera_Resistance_Disposition with 0.3 nodosities/g (99% reduction), no tuberosities, OIV score 9, and no growth impact. Commercial rootstock '110 Richter' (V. berlandieri × V. rupestris) showed intermediate resistance with 3.2 nodosities/g, tuberosity rating 1.2, OIV score 7, and 8% growth reduction—sufficient for practical vineyard use. Importantly, phylloxera populations recovered from resistant rootstocks showed no enhanced virulence when re-inoculated onto susceptible vines, indicating that resistance operates through host defense rather than supporting adapted phylloxera biotypes under these experimental conditions.
A Root_Phylloxera_Resistance_Disposition is a Phylloxera_Resistance_Disposition that inheres in a Vitis plant root system and is realized through defense processes that limit feeding damage, gall development, population establishment, or vine decline caused by the radicicole (root-feeding) form of grape phylloxera (Daktulosphaira vitifoliae). This disposition represents the economically critical resistance that has enabled continued V. vinifera cultivation worldwide through grafting onto resistant rootstocks. Root phylloxera infestation begins when crawlers or eggs deposited on roots initiate feeding on root tip meristems and cortical tissues. On susceptible V. vinifera roots, phylloxera feeding induces characteristic gall structures—nodosities (hook-shaped swellings on young white roots) and tuberosities (warty proliferations on lignified structural roots)—that progressively destroy root function. Resistant roots respond to phylloxera attack through multiple mechanisms including rapid hypersensitive cell death at feeding sites preventing sustained nutrition, formation of wound periderm isolating damage, production of toxic or deterrent secondary metabolites, and maintenance of functional root capacity despite limited feeding. The distinction between immunity (no feeding, no galls), resistance (limited feeding with minimal damage), and tolerance (feeding occurs but vine productivity maintained) is important, as most commercial rootstocks exhibit resistance or tolerance rather than true immunity. Root_Phylloxera_Resistance_Disposition is the foundation of modern viticulture, with over 80% of global vineyard area planted on resistant rootstocks, and understanding its mechanisms remains essential for addressing emerging phylloxera biotypes that may overcome existing resistance sources.
Root_Phylloxera_Resistance_Disposition is_a Phylloxera_Resistance_Disposition. This class addresses resistance in root tissues to the radicicole form, which is the economically significant phase causing vine decline and death in susceptible V. vinifera. Child classes include Nodosity_Limitation_Disposition (resistance to young root galls), Tuberosity_Resistance_Disposition (resistance to lignified root damage), and Root_Regeneration_Disposition (compensatory root growth maintaining function). The combination of these mechanisms determines overall rootstock performance. Concern exists regarding phylloxera biotypes potentially overcoming resistance in some rootstocks, particularly in Australian and Californian vineyards.
Radizikolenresistenz
Wurzelgallen-Resistenz
Wurzelreblaus-Resistenz-Disposition
radicicole resistance
root gall resistance
rootstock phylloxera resistance
Granett J et al. (2001) Biology and management of grape phylloxera, Annual Review of Entomology 46:387-412 Kellow AV et al. (2004) Interaction between Vitis vinifera and grape phylloxera: changes in root tissue during nodosity formation, Annals of Botany 93:581-590 Powell KS et al. (2013) Tolerance and resistance to root damage are not correlated during phylloxera host-plant interactions, Plant Pathology 62:1251-1261
Root Phylloxera Resistance Disposition
5BB Kober (V. berlandieri × V. riparia) demonstrates strong root phylloxera resistance through rapid wound periderm formation, with nodosity formation limited to superficial root tips and no tuberosity development even under high radicicole population pressure, supporting Grüner Veltliner and Zweigelt production throughout Austrian wine regions. Börner (V. riparia × V. cinerea) shows near-complete immunity to root phylloxera with essentially no gall formation, and is used experimentally and in commercial plantings in German viticulture where extreme resistance is prioritized. 41B Millardet et de Grasset (V. vinifera Chasselas × V. berlandieri) shows reduced root phylloxera resistance compared to pure American species crosses, with tuberosity formation observed under heavy infestation in some Australian vineyards.
A phylloxera-resistance rootstock role that inheres in a grafted rootstock by virtue of its specific capacity to resist root-feeding (radicicole) forms of grape phylloxera (Daktulosphaira vitifoliae). Root phylloxera resistance prevents or limits the formation of nodosities (galls on young, unsuberized root tips) and tuberosities (galls on mature, lignified roots) through mechanisms including rapid wound periderm formation within 48-72 hours of insect probing, localized hypersensitive cell death at feeding sites, accumulation of phenolic and terpenoid compounds in root cortex tissue, and anatomical features such as thick-walled cortical cells that impede stylet penetration. Root phylloxera resistance is the functionally critical resistance form for commercial viticulture, as the radicicole life cycle form causes the economically devastating root damage that led to the destruction of an estimated 2.5 million hectares of European vineyards in the late 19th century.
Root_Phylloxera_Resistant_Rootstock_Role captures resistance to the radicicole (root-feeding) phylloxera forms specifically. Resistance mechanisms operate at the root tissue level and are distinct from the foliar resistance that protects against gallicole (leaf-galling) forms. Phylloxera biotype diversity (strains A-G and beyond) means that resistance is not absolute adapted biotypes have overcome resistance in some rootstocks, notably AXR#1 (V. vinifera × V. rupestris) in California in the 1980s-1990s. Modern rootstock breeding programs screen against multiple biotypes simultaneously.
gegen Wurzelreblaus resistente Unterlage
radicicole phylloxera resistant rootstock
root form resistant rootstock
root phylloxera resistant rootstock
wurzelreblausresistente Unterlage
Granett J et al. (2001) Biology and management of grape phylloxera, Annual Review of Entomology 46: 387-412 Forneck A, Huber L (2009) (A)sexual reproduction – a review of life cycles of grape phylloxera, Journal of Insect Science 9: 1-10 Powell KS (2012) A holistic approach to future management of grapevine phylloxera, Springer pp.: 219-251 Kocsis L et al. (2020) Phylloxera resistance of rootstocks, Acta Horticulturae 1276: 181-186
Root Phylloxera Resistant Rootstock Role
Root radial conductivity assessment in grapevine under varying conditions reveals the aquaporin regulation of water uptake, with well-hydrated roots showing Lpr_radial of 2.8 × 10⁻⁷ m/s/MPa during daytime transpiration demand, declining to 0.9 × 10⁻⁷ at night (circadian aquaporin regulation), and further declining to 0.4 × 10⁻⁷ under moderate water stress (ABA-induced aquaporin downregulation), demonstrating the dynamic regulation of radial conductivity that modulates water uptake capacity in response to plant water status and demand.
Root radial conductivity quality is a root hydraulic conductivity quality that represents the capacity for water movement from the root surface through cortical tissues to the central xylem, characterizing the cell-to-cell water transport pathway that often constitutes the primary hydraulic resistance in roots and is highly regulated through aquaporin activity and developmental modifications of cell wall barriers. Within the BFO framework, this quality specifically addresses the radial (perpendicular to root axis) component of root water transport, distinguishing the pathway crossing multiple cell layers from the axial flow through xylem vessels while recognizing that radial conductivity is the most physiologically regulated and environmentally responsive component of root hydraulic architecture. The ontological significance lies in radial conductivity determining the rate of water extraction from soil into the transpiration stream, where the resistance to water crossing root tissues from epidermis through cortex and endodermis to xylem establishes the relationship between soil water potential and xylem water potential that determines root water uptake efficiency. The pathway components of radial conductivity include the apoplastic route (water moving through cell walls, limited by suberized barriers at the endodermis), the symplastic route (water moving through plasmodesmata connecting adjacent cells), and the transcellular route (water crossing cell membranes, requiring aquaporins for efficient flux), with the relative contribution of these pathways varying with root development and environmental conditions. The regulatory mechanisms affecting radial conductivity include aquaporin expression and gating (rapidly modulating transcellular water permeability), Casparian strip and suberin lamella development (progressively blocking apoplastic flow), and root aging (older roots typically showing reduced radial conductivity from increased suberization), creating complex spatiotemporal patterns of conductivity along root systems. The diurnal and stress-induced variation in radial conductivity is substantial, with aquaporin-mediated changes producing rapid (minutes to hours) conductivity adjustments in response to water status, hormones, and nutrient signals, enabling root systems to actively regulate water uptake rather than responding passively to water potential gradients. Understanding root radial conductivity quality enables assessment of the rate-limiting step in root water uptake, interpretation of aquaporin effects on vine water relations, and evaluation of root anatomical and physiological adaptations affecting water extraction efficiency.
Root_Radial_Conductivity_Quality addresses the cell-crossing pathway from root surface to xylem that often limits root water uptake. This quality is highly regulated through aquaporin activity and developmental barriers. The class supports water uptake regulation assessment through radial conductivity characterization.
Lpr_radial
Wurzelzellpermeabilität
cortical conductivity
kortikale Leitfähigkeit
radial root resistance
radiale Wurzelleitfähigkeit
root cell permeability
Steudle E, Peterson CA (1998) How does water get through roots? Journal of Experimental Botany 49(322): 775-788 Vandeleur RK et al. (2009) The role of plasma membrane intrinsic protein aquaporins in water transport through roots. Plant and Soil 323(1-2): 167-183
Root Radial Conductivity Quality
Experimental quantification of Root_Regeneration_Disposition at Agriculture Victoria, Australia, employed rhizotron observation of root dynamics in phylloxera-infested and control vines across 8 rootstock genotypes over two growing seasons. Minirhizotron cameras recorded root appearance, growth, and disappearance at weekly intervals, enabling calculation of root production, mortality, and turnover rates. Under phylloxera infestation, V. vinifera 'Sultana' (own-rooted) showed root production rates of 0.8 roots/cm²/week but mortality of 1.2 roots/cm²/week, resulting in progressive net root loss and 45% reduction in total root length by season end. The regeneration-tolerant rootstock '1103 Paulsen' (V. berlandieri × V. rupestris) demonstrated strong Root_Regeneration_Disposition with production rates increasing from 1.1 roots/cm²/week (control) to 2.3 roots/cm²/week under infestation—a compensatory response doubling root production. Despite elevated mortality (1.8 roots/cm²/week), '1103 Paulsen' maintained net positive root growth and equivalent total root length to uninfested controls. Carbohydrate analysis revealed that '1103 Paulsen' mobilized 35% more starch reserves from permanent roots under infestation compared to 'Sultana', fueling the enhanced regeneration response. This tolerance-based Root_Regeneration_Disposition explains the excellent field performance of '1103 Paulsen' in phylloxera-infested Australian vineyards despite its moderate nodosity scores in pot bioassays.
A Root_Regeneration_Disposition is a Root_Phylloxera_Resistance_Disposition that inheres in a Vitis plant and is realized through the capacity to produce new root growth rapidly and extensively, compensating for phylloxera-induced damage and maintaining adequate root function despite ongoing infestation pressure. This disposition represents a tolerance mechanism distinct from resistance mechanisms that prevent or limit phylloxera feeding, enabling vines to sustain productivity through continual replacement of damaged roots rather than through exclusion or elimination of the pest. Phylloxera feeding destroys root tips and progressively degrades the root system genotypes with strong Root_Regeneration_Disposition respond by initiating new root growth from undamaged tissues faster than phylloxera can colonize, maintaining a functional root population adequate for water and nutrient uptake. Root regeneration capacity depends on the availability of stored carbohydrate reserves, hormonal signaling promoting adventitious root initiation, meristematic activity in root pericycle and cambium tissues, and environmental conditions (soil moisture, temperature, aeration) permitting rapid root growth. This tolerance strategy is particularly relevant for rootstocks in the V. berlandieri background, which may permit moderate phylloxera feeding but compensate through vigorous root system restoration. Root_Regeneration_Disposition provides practical field resistance even in the absence of strong antibiosis or antixenosis, though it may be insufficient under extreme phylloxera pressure or in conditions (drought, nutrient stress) limiting regeneration capacity. The concept parallels wound tolerance in other plant-pest interactions where damage compensation rather than damage avoidance provides functional protection.
Root_Regeneration_Disposition is_a Root_Phylloxera_Resistance_Disposition. This class represents tolerance through damage compensation rather than resistance through damage prevention, providing an alternative or complementary strategy to Nodosity_Limitation_Disposition and Tuberosity_Resistance_Disposition. Root regeneration capacity varies among rootstocks and is influenced by vine carbohydrate status, making management practices affecting vine reserves (crop load, timing of irrigation) relevant to expression of this tolerance mechanism. The distinction between "resistance" (limiting pest damage) and "tolerance" (compensating for damage) is particularly clear in this context.
Kompensatorisches-Wurzelwachstum
Wurzelregenerations-Disposition
Wurzelsystem-Erholung
compensatory root growth
root system recovery
root turnover capacity
Powell KS et al. (2013) Tolerance and resistance to root damage are not correlated during phylloxera host-plant interactions, Plant Pathology 62:1251-1261 Clingeleffer PR, Krake LR (2002) Responses of Cabernet Sauvignon grapevines to minimal pruning and phylloxera infestation, Australian Journal of Grape and Wine Research 8:73-83 Keller M (2020) The Science of Grapevines, 3rd Edition, Academic Press
Root Regeneration Disposition
Root-shoot synchrony assessment through the season in Chardonnay reveals the characteristic coordination patterns, with budbreak to flowering showing root-shoot asynchrony (shoot growth rate 6 cm/day while root growth minimal, supported by reserve mobilization), flowering to véraison showing improved synchrony (shoot growth slowing to 2 cm/day while root growth increases, balanced resource flow), and post-harvest showing reversed asynchrony (shoot senescence while root growth peaks at 0.8 cm/day new root production, directing carbohydrates to below-ground reserves), demonstrating the complementary phasing of root and shoot activity that characterizes healthy vine function versus the problematic sustained asynchrony that might indicate rootstock-scion incompatibility or environmental stress.
Root shoot synchrony quality is an organ synchrony quality that represents the temporal coordination between root system activity and shoot system development, characterizing the alignment of below-ground resource acquisition with above-ground resource demand that determines whether the vine functions as an integrated system with complementary organ activity or experiences periodic mismatches creating resource deficits or excesses that affect vine balance and productivity. Within the BFO framework, this quality specifically addresses the root-shoot coordination dimension of organ synchrony, distinguishing this fundamental below-ground/above-ground relationship from synchrony among organs of similar type while recognizing that root-shoot synchrony determines the integration of the whole-plant system, with roots providing water, nutrients, and hormonal signals that support shoot function while shoots provide photosynthate that fuels root maintenance and growth. The ontological significance lies in root-shoot synchrony determining the resource balance that affects vine vigor, stress tolerance, and reproductive success, where aligned root and shoot activity ensures that water and nutrient supply from roots matches the transpiration and growth demands of shoots, while misalignment creates either resource deficits (shoot demand exceeding root supply capacity, causing stress) or inefficiencies (root capacity exceeding shoot demand, representing unused potential). The seasonal patterns of root-shoot synchrony show characteristic phases in grapevine, with spring shoot growth initially preceding root activity (supported by reserve mobilization), followed by root growth flushes in late spring and post-harvest when carbohydrate supply exceeds shoot demand, and with potential competition between root and shoot growth during periods of resource limitation. The environmental factors affecting root-shoot synchrony include the decoupling of soil and air temperatures (warm air triggering shoot growth while cool soil limits root activity, or vice versa), water availability patterns (affecting root function independently of shoot demand), and nutrient availability (affecting the capacity of roots to support shoot requirements). The rootstock effects on root-shoot synchrony are significant, as rootstock genetics determine root system vigor, depth, and activity patterns that interact with scion demands, potentially creating well-matched or poorly coordinated combinations depending on the rootstock-scion pairing. Understanding root-shoot synchrony quality enables assessment of whole-plant integration, identification of seasonal periods of potential root-shoot mismatch, and management interventions that promote coordinated organ function supporting balanced vine development.
Root_Shoot_Synchrony_Quality addresses the coordination between below-ground and above-ground organ systems. This quality determines whole-plant resource integration affecting vine balance and productivity. The class supports rootstock-scion evaluation and seasonal management through root-shoot synchrony characterization.
Quellen-Senken-Synchronie
Wurzel-Trieb-Balance
Wurzel-Trieb-Synchronie
below-ground above-ground coordination
root-shoot balance
source-sink synchrony
unterirdisch-oberirdische Koordination
vascular integration
Comas LH et al. (2005) Root traits contributing to plant productivity under drought. Frontiers in Plant Science 4: 442 Zapata C et al. (2004) Partitioning and mobilization of starch and N reserves in grapevine (Vitis vinifera L.). Journal of Plant Physiology 161(9): 1031-1040
Root Shoot Synchrony Quality
Investigation of Root_Sodium_Exclusion_Disposition at the University of Adelaide employed ²²Na⁺ tracer studies and membrane transport assays to characterize sodium influx mechanisms across 8 rootstocks. Short-term (20 min) ²²Na⁺ uptake was measured at 50 mM Na⁺ to assess initial influx rates, with pharmacological dissection using channel blockers (Gd³⁺ for NSCC, tetraethylammonium for K⁺ channels) to identify entry pathways. Root_Sodium_Exclusion_Disposition varied 3-fold among rootstocks. '140 Ruggeri' showed the lowest sodium influx (1.8 μmol g⁻¹ root FW h⁻¹), with 65% of uptake blocked by Gd³⁺ indicating NSCC-mediated entry. 'Schwarzmann' displayed highest influx (5.4 μmol g⁻¹ h⁻¹) with similar pharmacological profile (70% Gd³⁺-sensitive). The remaining influx was largely apoplastic, as confirmed by parallel studies with the apoplastic tracer PTS. SOS1 expression analysis revealed that '140 Ruggeri' showed 3.5-fold higher root SOS1 transcript levels under salinity compared to 'Schwarzmann', suggesting enhanced sodium efflux capacity contributes to its Root_Sodium_Exclusion_Disposition. Whole-plant ²²Na⁺ partitioning showed that low-influx '140 Ruggeri' retained 78% of absorbed sodium in roots (suggesting effective root sequestration or efflux) while high-influx 'Schwarzmann' translocated 55% to shoots. These findings indicate that Root_Sodium_Exclusion_Disposition involves both reduced NSCC-mediated influx and enhanced SOS1-mediated efflux, with root retention capacity providing additional protection.
A Root_Sodium_Exclusion_Disposition is a Sodium_Exclusion_Disposition that inheres in Vitis root tissues and is realized through mechanisms at the root surface and cortex that restrict the initial uptake of sodium ions from the soil solution and limit sodium accumulation in root symplast, reducing the sodium load available for xylem loading. This disposition represents the first barrier against sodium entry, operating at the soil-root interface where sodium must cross root cell membranes to enter the symplastic pathway to the shoot. Sodium can enter root cells through multiple pathways including non-selective cation channels (NSCC), which are the primary entry route under salinity, and potentially through HKT2 family transporters that can mediate sodium uptake when potassium is limiting. Root_Sodium_Exclusion_Disposition involves reduced activity or expression of sodium-permeable channels, enhanced plasma membrane H⁺-ATPase activity maintaining the driving force for sodium efflux, and active sodium extrusion via SOS1 (Salt Overly Sensitive 1) Na⁺/H⁺ antiporters that pump sodium back to the soil solution. Additionally, because sodium can enter via apoplastic flow, root_Sodium_Exclusion_Disposition is enhanced by Casparian strip integrity and suberization that blocks apoplastic bypass. Root sodium exclusion reduces the sodium load that must be handled by subsequent exclusion mechanisms, but complete exclusion at root level is typically not achievable, making xylem loading control and retrieval important complementary mechanisms.
Root_Sodium_Exclusion_Disposition is_a Sodium_Exclusion_Disposition. Root sodium uptake occurs primarily through non-selective cation channels (NSCCs), with some apoplastic contribution. SOS1 Na⁺/H⁺ antiporter mediates active sodium efflux back to soil. Root sodium exclusion provides the first barrier but is typically incomplete. Casparian strip integrity affects apoplastic bypass. Root sodium retention (vacuolar sequestration) can complement exclusion by reducing translocation even when uptake occurs.
Kortikaler-Natrium-Ausschluss
Wurzel-Natrium-Barriere
Wurzel-Natriumausschluss-Disposition
cortical sodium exclusion
root sodium barrier
root sodium uptake restriction
Tester M, Davenport R (2003) Na⁺ tolerance and Na⁺ transport in higher plants, Annals of Botany 91:503-527 Munns R, Tester M (2008) Mechanisms of salinity tolerance, Annual Review of Plant Biology 59:651-681 Shi H et al. (2002) The Arabidopsis thaliana salt tolerance gene SOS1 encodes a putative Na⁺/H⁺ antiporter, Proceedings of the National Academy of Sciences 97:6896-6901
Root Sodium Exclusion Disposition
Root surface area assessment comparing irrigation treatments reveals the water management effect on absorptive capacity, with well-irrigated vines showing total root surface area of 52 m²/vine (abundant fine root proliferation in moist soil), deficit-irrigated vines showing 38 m²/vine (reduced fine root production, some root mortality in dry zones), and severely stressed vines showing only 24 m²/vine (substantial fine root loss, limited to deeper moist zones), demonstrating how soil moisture affects the root surface area available for nutrient and water uptake with implications for vine nutrition and stress recovery.
Root surface area quality is a root uptake capacity quality that represents the total external surface of root tissues available for nutrient and water absorption, characterizing the primary determinant of absorptive capacity that integrates root length, diameter distribution, and branching architecture into a single measure of the interface between root and soil available for resource acquisition. Within the BFO framework, this quality specifically addresses the surface area dimension of root structure, distinguishing total absorbing surface from other root parameters while recognizing that surface area provides the most direct measure of uptake potential, as nutrient and water absorption occur at root surfaces through transporter proteins and aquaporins whose activity is proportional to the membrane area available for their function. The ontological significance lies in root surface area establishing the physical interface for all root-soil exchange processes, where total surface area (typically 20-60 m² per mature vine) determines the maximum possible flux of nutrients and water into the vine given soil concentrations and transporter activities, making surface area a fundamental constraint on vine resource acquisition capacity. The mathematical relationship between root dimensions and surface area shows that for cylindrical roots, surface area = π × diameter × length, such that doubling length doubles surface area while halving diameter at constant biomass quadruples surface area, explaining the high absorption efficiency of fine roots and root hairs that maximize surface area per unit carbon invested. The distribution of surface area among root classes shows fine roots (< 2 mm diameter) contributing the majority of absorbing surface despite representing a minority of root biomass, with root hairs (often 100-200/mm² on fine roots) further extending the effective absorbing surface by 2-3 fold in the critical zone immediately surrounding roots. The environmental factors affecting root surface area development include soil moisture (adequate water promoting fine root proliferation), soil aeration (waterlogging reducing root surface through death and limiting new growth), soil compaction (restricting root proliferation), and nutrient availability (both deficiency and excess potentially affecting root development patterns). Understanding root surface area quality enables quantitative assessment of vine absorptive capacity, comparison of rootstock root system characteristics, and interpretation of how root morphology responds to environmental conditions affecting resource availability.
Root_Surface_Area_Quality addresses the total absorbing interface between root system and soil determining uptake potential. This quality integrates root length, diameter, and branching into the fundamental metric of absorptive capacity. The class supports root system assessment and uptake capacity estimation through surface area characterization.
Aufnahmegrenzfläche
Gesamtwurzeloberfläche
Wurzeloberfläche
absorbierende Oberfläche
absorbing surface
root area
total root surface
uptake interface area
Eissenstat DM, Yanai RD (1997) The ecology of root lifespan. Advances in Ecological Research 27: 1-60 Smart DR et al. (2006) Grapevine rooting patterns: a comprehensive analysis and a review. American Journal of Enology and Viticulture 57(1): 89-104
Root Surface Area Quality
Cabernet Sauvignon grafted on 110 Richter rootstock at 12 years showing root system extending 180 cm depth in deep loam soil with maximum root density 40-70 cm depth, lateral spread 250 cm diameter exceeding canopy projection, root system comprising several primary roots originating from original cutting base now 8-12 cm diameter at crown, extensive lateral root branching creating hierarchical network, abundant fine roots with mycorrhizal colonization in upper soil providing water and nutrient uptake, root system demonstrating 110R characteristic vigorous deep rooting enabling drought tolerance
Root system is the entire below-ground organ system of grapevine comprising all roots originating from rootstock or own-rooted vine, including primary roots derived from radicle or adventitious roots from cuttings, lateral roots branching from primary roots creating hierarchical architecture, and fine absorptive roots with mycorrhizal associations, typically extending 30-200+ cm depth and 50-300+ cm lateral spread depending on rootstock genetics, soil conditions, and vine age, serving essential functions of anchorage stabilizing vine, water absorption from soil providing 50-100+ liters daily during growing season, mineral nutrient uptake supplying nitrogen, phosphorus, potassium, and micronutrients, carbohydrate storage in structural roots particularly during dormancy, and hormone synthesis producing cytokinins and other signals regulating shoot growth, with root system architecture and extent profoundly influenced by rootstock variety genetics determining vigor, drought tolerance, and soil adaptation characteristics. Root system structural organization includes= comprising primary roots (main structural roots originating from radicle in seedlings or from cutting base in propagated vines providing deep penetration), containing lateral roots (branch roots arising from primary roots creating hierarchical branching network), including fine roots (small-diameter absorptive roots with root hairs providing majority of water and nutrient uptake surface area), showing mycorrhizal associations (symbiotic fungi colonizing roots enhancing water and phosphorus uptake), and demonstrating root orders (hierarchical classification from first-order fine roots through progressively higher-order structural roots to primary root). Root system depth and distribution includes= showing vertical extent (root depth typically 50-150 cm in normal soils potentially reaching 200-600 cm in deep permeable soils or under drought stress), exhibiting lateral spread (horizontal root extension typically 100-300 cm often exceeding canopy projection), demonstrating concentration zone (majority of absorptive roots typically in upper 30-80 cm soil where moisture and nutrients most available), displaying rootstock effects (vigorous rootstocks producing extensive deep systems while low-vigor rootstocks showing more compact shallow systems), and revealing soil limitations (root distribution constrained by hardpans, water tables, toxic layers, or compaction). Root system functions includes= providing anchorage (structural roots stabilizing vine preventing uprooting or toppling under wind or crop load), absorbing water (fine roots and root hairs taking up soil water providing 50-100+ liters daily during peak transpiration), taking up minerals (roots absorbing nitrogen, phosphorus, potassium, calcium, magnesium, and micronutrients from soil solution), storing carbohydrates (structural roots accumulating starch reserves during fall mobilized in spring for growth), synthesizing hormones (roots producing cytokinins, abscisic acid, and other signaling compounds influencing shoot development and stress responses), and supporting mycorrhizae (roots hosting fungal symbionts receiving enhanced nutrient uptake in exchange for carbohydrates). Root system seasonal dynamics includes= showing spring growth flush (roots resuming growth in spring before bud break utilizing stored reserves), exhibiting summer growth (continued root development during growing season though potentially reduced during peak shoot growth and fruit development), demonstrating fall growth period (roots showing renewed growth late season as shoot growth ceases and carbohydrate partitioning shifts), entering winter dormancy (root growth ceasing during cold periods though roots remaining physiologically active), and displaying asynchrony with shoots (root growth peaks often offset from shoot growth peaks reflecting resource competition and allocation priorities). Root system rootstock genetic control includes= determining depth potential (Vitis berlandieri types showing deep rooting, V. riparia shallow rooting, hybrids intermediate), controlling lateral spread (vigorous rootstocks producing extensive lateral systems while low-vigor types more compact), influencing branching patterns (genetic variation in lateral root density and architecture), affecting drought response (deep-rooting rootstocks showing superior drought tolerance through accessing deeper moisture), and conferring soil adaptations (rootstock genetics determining lime tolerance, salt tolerance, and adaptation to specific soil types).
Root_System emphasizes complete below-ground organ system and is distinguished from shoot system by focus on roots. Includes Primary_Root, lateral roots, and fine absorptive roots. Critical for understanding anchorage, water and nutrient uptake, carbohydrate storage, hormone production, rootstock genetic effects, and the essential below-ground system supporting vine growth and productivity.
Below-ground system
Radicular system
Radikalsystem
Root apparatus
Unterirdisches Organsystem
Wurzelsystem
Root System
Comprehensive characterization of Root_System_Drought_Adaptation_Disposition at UC Davis compared root system properties and drought performance across 15 commercial rootstocks grown in deep soil columns (2 m depth) with controlled water deficit. Root distribution was assessed by minirhizotron imaging, total root length and depth by excavation at trial completion, and functional water acquisition by deuterium-labeled water injection at different depths. Rootstocks showed substantial variation in Root_System_Drought_Adaptation_Disposition. '110 Richter' (V. berlandieri × V. rupestris) exhibited deep-rooting adaptation with 45% of root length below 1 m depth, maximum rooting depth of 1.85 m, and efficient deep water acquisition (deuterium from 1.5 m depth detected in leaves within 48 hours). '101-14 Mgt' (V. riparia × V. rupestris) showed shallow root concentration with only 15% below 1 m and maximum depth of 1.2 m, relying on dense surface rooting (root length density 2.8 cm cm⁻³ in upper 30 cm versus 1.4 cm cm⁻³ for '110 Richter'). Under progressive drought, '110 Richter'-grafted vines maintained higher predawn water potential (-0.35 versus -0.65 MPa for '101-14 Mgt' at equivalent surface soil moisture), demonstrating how Root_System_Drought_Adaptation_Disposition through deep rooting provides access to water unavailable to shallow-rooted systems. Scion variety ('Cabernet Sauvignon') was constant, confirming that observed differences reflected rootstock root properties.
A Root_System_Drought_Adaptation_Disposition is a Drought_Tolerance_Disposition that inheres in the root system of a Vitis plant and is realized through root architectural, morphological, and physiological traits that enhance water acquisition from soil under drought conditions. This disposition represents the below-ground component of drought adaptation, complementing above-ground stomatal regulation and osmotic adjustment by determining the vine's capacity to access and transport soil water to transpiring shoots. Root system properties critical for drought adaptation include rooting depth (access to deep soil water reserves), root density and distribution (exploitation of soil volume), root hydraulic conductance (water transport efficiency), and root-to-shoot signaling (coordination of whole-plant drought responses through ABA and other signals). In grafted grapevines, Root_System_Drought_Adaptation_Disposition is primarily determined by the rootstock genotype, making rootstock selection a primary management tool for drought adaptation. Rootstocks vary substantially in root system architecture and function, ranging from shallow, dense root systems suited to soils with reliable surface moisture to deep, extensive systems adapted to access water in deep soil layers during prolonged drought. The interaction between rootstock root properties and scion stomatal behavior determines whole-vine drought response, with combinations of drought-adapted rootstock and water-conserving scion potentially providing synergistic drought tolerance.
Root_System_Drought_Adaptation_Disposition is_a Drought_Tolerance_Disposition. Root system properties are primarily determined by rootstock genotype in grafted vines. Child classes address specific root adaptations including Deep_Rooting_Disposition, Root_Density_Disposition, and Root_Hydraulic_Conductance_Disposition. Root adaptation interacts with scion stomatal behavior to determine whole-vine drought response. Root system traits show strong genotype × soil type interactions, requiring site-specific rootstock selection.
Unterirdische-Wasseraufnahme
Wurzel-Wasserakquisition
Wurzelsystem-Trockenanpassungs-Disposition
below-ground water acquisition
root drought adaptation
Smart DR et al. (2006) Grapevine rooting patterns: a comprehensive analysis and review, American Journal of Enology and Viticulture 57:89-104 Alsina MM et al. (2011) Seasonal changes of whole root system conductance by a drought-tolerant grape root system, Journal of Experimental Botany 62:99-109 Tramontini S et al. (2013) Rootstock control of scion response to water stress in grapevine, Environmental and Experimental Botany 93:20-26
Root System Drought Adaptation Disposition
Pinot Noir grafted on 101-14 Millardet rootstock showing root tip on third-order lateral root, root tip 6.2 mm length comprising root cap 0.6 mm, apical meristem 0.4 mm with actively dividing cells, elongation zone 3.8 mm with cells expanding, and maturation zone beginning 5.5 mm from tip with emerging root hairs, root tip showing positive gravitropic curvature, microscopic examination revealing mitotic figures in apical meristem indicating active cell division, root tip elongating estimated 2.8 mm per day during mid-May sampling, root cap producing mucilage and border cells
Root tip is the apical terminal portion of root typically 2-10 mm length comprising root cap, apical meristem, and elongation zone, characterized by undifferentiated or recently differentiated tissues actively involved in root growth and elongation, protected by root cap covering meristem providing mechanical protection and gravitropic sensing, showing white to translucent coloration reflecting young undifferentiated state, and serving as primary site of root elongation through cell division in apical meristem producing new cells and cell elongation in elongation zone increasing cell length, with root tip demonstrating gravitropic response growing downward in response to gravity, secreting mucilage lubricating passage through soil, sensing soil conditions through root cap statoliths and border cells, producing root hairs in maturation zone just behind tip, and representing critical growth region where root penetration, directional control, and initial tissue differentiation occur enabling root system expansion and soil exploration throughout vine life. Root tip structural organization includes= comprising root cap (outer protective tissue covering meristem typically 0.3-0.8 mm length with columella cells containing statoliths), containing apical meristem (dome-shaped zone of actively dividing undifferentiated cells typically 0.2-0.5 mm length producing new root tissues), including elongation zone (region where recently divided cells elongate increasing length typically 1-5 mm behind meristem), showing maturation zone (area where cells complete differentiation and root hairs emerge typically 5-15 mm from tip), and demonstrating quiescent center (slowly dividing organizing cells at meristem center maintaining meristem identity and stem cell populations). Root tip growth functions includes= enabling root elongation (root tip apical meristem cell division plus elongation zone cell expansion providing all root length increase), directing growth trajectory (root tip gravitropic and thigmotropic responses controlling root direction through soil), penetrating soil (root tip pushing through soil particles aided by root cap protection and mucilage lubrication), sensing environment (root tip detecting gravity, moisture gradients, nutrients, toxins, and obstacles through specialized cells), and initiating differentiation (root tip producing cells that differentiate into epidermis, cortex, endodermis, and vascular tissues as move away from meristem). Root tip cell division and expansion includes= showing meristem mitosis (apical meristem cells dividing every 12-24 hours during active growth producing new cells), exhibiting cell cycle progression (meristem cells progressing through G1, S, G2, and M phases creating daughter cells), demonstrating elongation zone expansion (cells in elongation zone increasing length 10-50 fold through vacuolation and wall extension), revealing growth rate (root tips potentially elongating 1-10 mm per day depending on root order, temperature, and conditions), and displaying seasonal variation (root tip activity highest spring and fall declining summer and ceasing winter dormancy). Root tip protection and lubrication includes= producing root cap (outer cap cells continuously replaced as damaged or sloughed protecting underlying meristem), secreting mucilage (root cap border cells releasing polysaccharide gel lubricating root passage and modifying rhizosphere), forming border cells (root cap producing and shedding peripheral cells that may defend against pathogens), demonstrating mechanical protection (root cap hardness and geometry protecting delicate meristem from abrasion and impact), and showing expendable structure (root cap cells programmed cell death and sloughing allowing continuous protection despite damage). Root tip gravitropic sensing includes= containing statoliths (specialized amyloplasts in root cap columella cells sedimenting in response to gravity), demonstrating downward growth (primary root tips showing positive gravitropism growing toward earth center), exhibiting lateral gravitropism (lateral root tips showing diagravitropic response maintaining horizontal or oblique angles), revealing auxin redistribution (gravity-induced statolith sedimentation triggering asymmetric auxin distribution causing differential growth), and enabling directional control (gravitropic response allowing roots to maintain appropriate orientations despite planting angle or soil displacement). Root tip in different root types includes= showing vigorous growth in primary roots (primary root tips with large meristems producing rapid elongation 3-10 mm daily), exhibiting moderate activity in secondary roots (secondary root tips showing intermediate growth rates 1-5 mm daily), demonstrating variable activity in tertiary roots (fine root tips ranging from rapid growth in favorable conditions to dormancy in stress), revealing order-specific characteristics (lower-order root tips typically larger with higher growth rates than higher-order tips), and displaying seasonal dormancy (root tips ceasing growth entering quiescent state during winter or severe drought resuming activity when conditions improve). Root tip and root hair initiation includes= producing trichoblast precursors (root tip epidermis cells differentiating into root hair-forming trichoblasts or non-hair atrichoblasts), showing position-dependent differentiation (epidermal cells over periclinal walls becoming atrichoblasts while those over anticlinal walls becoming trichoblasts), demonstrating hair emergence zone (root hairs beginning to emerge 2-10 mm behind root tip in maturation zone after cell elongation completes), revealing hair production rate (root tip producing 10-100+ new root hairs per millimeter as elongates creating continuous hair zone), and enabling absorption function (root hair production immediately behind tip ensuring newly explored soil volume quickly exploited for water and nutrients).
Root_Tip emphasizes apical terminal region with meristem and is distinguished from mature differentiated root by focus on growth zone. Includes Primary_Root_Tip and Root_Apical_Meristem. Critical for understanding root elongation, cell division, gravitropism, soil penetration, directional control, and the apical growth region enabling root system expansion through continuous production of new tissues at root terminus.
Apical root region
Apikale Wurzelregion
Growing root tip
Root apex
Wurzelapex
Wurzelspitze
Root Tip
Root uptake capacity assessment comparing rootstocks reveals the structural basis for nutrient acquisition differences, with 1103P showing high uptake capacity (root surface area 45 m²/vine, extensive fine root development, deep soil exploration) supporting high nutrient uptake and vigorous scion growth, while 101-14 Mgt shows moderate capacity (root surface area 28 m²/vine, shallower distribution) providing more restrained nutrition matching lower-vigor production goals, demonstrating how rootstock-determined root uptake capacity affects the potential for nutrient and water acquisition that influences scion performance.
Root uptake capacity quality is a nutrient uptake efficiency quality that represents the overall absorptive potential of grapevine root systems determined by root architecture, morphology, and surface characteristics, characterizing the structural basis for nutrient and water acquisition that establishes the upper limit on uptake rates achievable through transport systems operating at the root-soil interface. Within the BFO framework, this quality addresses the structural dimension of root function, distinguishing the anatomical and morphological capacity for uptake from the physiological transport processes occurring at that surface while recognizing that root uptake capacity establishes the physical infrastructure within which transporters operate, with no amount of transporter upregulation able to overcome fundamental limitations in absorbing surface area. The ontological significance lies in root uptake capacity determining the potential for resource acquisition from soil, where the total absorbing surface area, the spatial distribution of that surface through soil volume, and the surface characteristics affecting absorption efficiency together establish the capacity for nutrient and water uptake that root physiological processes then realize to varying degrees depending on transporter activity and soil resource availability. The components of root uptake capacity include total root length and surface area (determining absorbing surface), root distribution in soil (affecting access to spatially heterogeneous resources), root diameter distribution (fine roots with higher surface area to volume ratio being more efficient for absorption), root hair development (extending the effective absorbing zone), and mycorrhizal colonization (dramatically extending the effective root system through hyphal networks). The developmental dynamics of root uptake capacity show seasonal patterns with new root growth in spring and post-harvest periods expanding uptake capacity, while root senescence, drought, and waterlogging reduce functional root surface area and thus uptake capacity, with perennial vines maintaining substantial root system infrastructure that persists across seasons. The rootstock genotype strongly influences root uptake capacity through effects on root system size, architecture, and vigor, with vigorous rootstocks producing larger root systems with greater uptake capacity but also potentially excessive scion vigor, requiring rootstock selection that balances uptake capacity with whole-vine balance objectives. Understanding root uptake capacity quality enables assessment of the structural basis for nutrient acquisition, evaluation of rootstock root system characteristics, and interpretation of how root morphology affects vine nutritional status and water relations.
Root_Uptake_Capacity_Quality serves as the parent class for root morphological parameters (Surface Area, Specific Root Length, Root Hair Density). This quality establishes the structural basis for nutrient and water acquisition potential. The class supports rootstock evaluation and root system assessment through uptake capacity characterization.
Aufnahmeoberflächenkapazität
Wurzelabsorptionskapazität
Wurzelaufnahmekapazität
Wurzelsystempotential
root absorptive capacity
root surface potential
root system capacity
uptake surface area
Eissenstat DM (1992) Costs and benefits of constructing roots of small diameter. Journal of Plant Nutrition 15(6-7): 763-782 Smart DR et al. (2006) Grapevine rooting patterns: a comprehensive analysis and a review. American Journal of Enology and Viticulture 57(1): 89-104
Root Uptake Capacity Quality
Comprehensive characterization of Root_Vigor_Disposition at INRAE Bordeaux employed rhizotron observation, soil coring, and complete root excavation across 18 rootstock genotypes grown ungrafted for 5 years in deep soil (> 3 m). Measurements included total root length by minirhizotron imaging, root length density at depth intervals, maximum rooting depth, root dry weight, and fine root (< 2 mm diameter) to coarse root ratio. Root_Vigor_Disposition showed dramatic variation among rootstocks. '110 Richter' (V. berlandieri × V. rupestris) exhibited high Root_Vigor_Disposition with total root length of 2,850 m per vine, root dry weight of 4.2 kg, root length density of 2.8 cm/cm³ in the 0-50 cm zone, maximum rooting depth of 2.6 m, and fine root fraction of 68%. 'SO4' (V. berlandieri × V. riparia) showed similarly extensive root development (2,650 m total length, 3.8 kg dry weight) but with shallower distribution (maximum depth 1.8 m). In contrast, '101-14 Mgt' (V. riparia × V. rupestris) displayed limited Root_Vigor_Disposition with total length of 1,250 m, dry weight of 1.8 kg, and maximum depth of 1.4 m. '3309C' (V. riparia × V. rupestris) showed intermediate characteristics (1,650 m, 2.4 kg). Root vigor was moderately correlated with conferred scion vigor (r = 0.72 when grafted with 'Cabernet Sauvignon'), confirming that Root_Vigor_Disposition contributes to but does not fully determine above-ground vigor expression. Interestingly, the fine root fraction showed inverse relationship with total root mass (r = -0.58), with smaller root systems investing proportionally more in absorptive capacity.
A Root_Vigor_Disposition is a Vigor_Disposition that inheres in the root system of a Vitis plant and is realized through the inherent capacity for root growth, determining the rate of root extension, the total root system size, root density and distribution patterns, and the overall below-ground biomass production. This disposition represents the underground component of vine vigor, reflecting the integrated activity of root apical meristems, lateral root initiation, and the allocation of resources toward root development. Root vigor is expressed through parameters including total root length, root dry weight, root length density (cm root per cm³ soil), rooting depth, and the ratio of fine absorptive roots to coarse structural roots. Root_Vigor_Disposition has fundamental implications for vine function, as the root system determines water and nutrient acquisition capacity, anchorage stability, carbohydrate storage reserves, and through root-derived signals (hormones, hydraulic signals) influences shoot growth and function. In grafted grapevines, Root_Vigor_Disposition is determined by rootstock genotype and represents a primary mechanism by which rootstocks influence scion performance. Rootstocks vary substantially in root system characteristics, from compact, limited root systems to extensive, vigorous systems that explore large soil volumes. The relationship between Root_Vigor_Disposition and Shoot_Vigor_Disposition is generally positive but not perfectly correlated, as root-to-shoot signaling and resource allocation patterns modify how below-ground growth capacity translates to above-ground vigor expression.
Root_Vigor_Disposition is_a Vigor_Disposition representing the below-ground vigor component. Root vigor is determined by rootstock genotype in grafted vines. Child classes include Extensive_Root_System_Disposition and Limited_Root_System_Disposition. Root vigor correlates with but does not fully determine scion vigor (~70% of variance). Root characteristics include total length/mass, depth distribution, and fine/coarse root ratio. Root vigor interacts with soil conditions (depth, texture, fertility) to determine functional root system size.
Unterirdische-Wuchskraft
Wurzelsystem-Entwicklung
Wurzelwuchsstärke-Disposition
below-ground vigor
root growth capacity
root system development
Smart DR et al. (2006) Grapevine rooting patterns: a comprehensive analysis and review, American Journal of Enology and Viticulture 57:89-104 Swanepoel JJ, Southey JM (1989) The influence of rootstock on the rooting pattern of the grapevine, South African Journal of Enology and Viticulture 10:23-28 Morlat R, Jacquet A (2003) Grapevine root system and soil characteristics in a vineyard maintained long-term with or without interrow sward, American Journal of Enology and Viticulture 54:1-7
Root Vigor Disposition
SO4 rootstock rooted cutting showing rooted cutting adventitious roots, cutting evaluated 6 weeks after treatment showing successful rooting, 7 adventitious roots emerged= 4 nodal (5-12 cm length) and 3 basal callus (4-9 cm length), roots showing white healthy appearance with beginning lateral branching, root system assessment indicating commercial quality meeting nursery standards, FDA staining showing functional root hairs on youngest root portions, rooted cutting demonstrating active water uptake supporting new shoot growth 8 cm, cutting ready for transplanting to nursery bed or grafting operation, rooted cutting adventitious roots representing successful propagation outcome
Rooted cutting adventitious root is cutting adventitious root emphasizing successfully rooted state and functional root system, characterized by cutting that has completed rooting process developing adequate root system for establishment, showing functional roots capable of water and nutrient absorption supporting cutting survival and growth, demonstrating vascular integration between roots and cutting enabling resource transport, and representing successful propagation outcome where cutting transitioned from dependent on reserves to physiologically independent plant, with rooted cutting showing typical characteristics including 3-10+ emerged roots distributed around cutting base and nodes, roots 3-15 cm length with beginning lateral branching, functional root hairs and mycorrhizal colonization enabling absorption, established vascular connection from roots through callus or stem to cutting vascular system, and capacity for active uptake supporting leaf expansion and shoot growth, and demonstrating that rooted status distinguishes successful cuttings ready for transplanting or field planting from unrooted or poorly rooted cuttings requiring continued nursery culture or culling, and representing commercial product of propagation industry supplying rooted rootstock cuttings for grafting or rooted vines for planting. Rooted cutting adventitious root functional criteria includes= showing adequate root number (successful rooting typically requiring 3-5+ roots providing redundancy and absorption capacity), exhibiting sufficient root length (roots typically needing 5-10+ cm providing adequate soil exploration), demonstrating lateral branching (successful roots beginning to produce lateral branches indicating healthy development), revealing functional absorption (roots showing root hairs and demonstrable water uptake capacity), and displaying vascular integration (functional connection between roots and cutting enabling transport). Rooted cutting adventitious root quality assessment includes= requiring visual inspection (evaluating root number, length, distribution, color, and general appearance), showing root system symmetry (well-rooted cuttings showing roots distributed around base rather than one-sided), exhibiting healthy appearance (roots white to tan color, turgid, absence of necrosis or rot), demonstrating appropriate development (root system development matching shoot growth indicating balanced plant), and revealing grading categories (cuttings graded premium, standard, or cull based on root system quality). Rooted cutting adventitious root physiological status includes= showing active metabolism (rooted cutting roots demonstrating respiration, ion uptake, and biosynthesis), exhibiting absorption capacity (roots capable of taking up water and nutrients from medium or soil), demonstrating transport function (functional xylem and phloem connecting roots to shoot enabling resource distribution), revealing growth capacity (roots capable of continued elongation and lateral branching), and displaying plant independence (rooted cutting physiologically independent no longer depending solely on stored reserves). Rooted cutting adventitious root in nursery production includes= determining transplant readiness (adequate rooting indicating cutting ready for transplanting from propagation to nursery bed), showing grafting stock (well-rooted rootstock cuttings serving as stock for grafting operations), exhibiting grading and selection (rooted cuttings graded separating commercial quality from substandard plants), revealing inventory management (rooted cutting inventory determining nursery production capacity and sales potential), and displaying market product (rooted rootstock cuttings representing sellable product for nurseries supplying grafting operations). Rooted cutting adventitious root transplanting considerations includes= requiring careful handling (rooted cutting roots relatively fragile requiring gentle handling preventing damage), showing timing importance (transplanting at appropriate stage when roots adequate but not excessively long or pot-bound), exhibiting hardening requirements (rooted cuttings potentially requiring acclimatization before field transplanting), demonstrating planting depth (rooted cuttings planted at appropriate depth maintaining proper root-shoot balance), and revealing establishment needs (newly transplanted rooted cuttings requiring irrigation and care during establishment period). Rooted cutting adventitious root post-transplant development includes= showing continued root growth (transplanted rooted cuttings extending roots into new soil developing extensive system), exhibiting lateral branching (primary adventitious roots producing abundant lateral roots creating hierarchical architecture), demonstrating secondary growth (older adventitious roots beginning secondary thickening developing woody structure), revealing mycorrhizal colonization (roots forming associations with arbuscular mycorrhizal fungi enhancing uptake), and displaying complete function (rooted cutting roots eventually developing full functionality comparable to seed-derived root systems). Rooted cutting adventitious root success indicators includes= showing high rooting percentage (propagation protocols achieving >70-90% rooting indicating successful methods), exhibiting uniform quality (rooted cutting population showing consistent root system development indicating controlled conditions), demonstrating vigorous growth (well-rooted cuttings showing strong shoot growth indicating functional roots supporting plant), revealing high survival (rooted cuttings showing high survival rates after transplanting confirming root system adequacy), and displaying market acceptance (rooted cutting quality meeting buyer standards enabling commercial sales). Rooted cutting adventitious root in own-rooted versus grafted vines includes= showing own-rooted use (rooted cuttings of V. vinifera planted directly in phylloxera-free regions or resistant varieties), exhibiting grafted vine production (rooted rootstock cuttings serving as stock for grafting V. vinifera scions), demonstrating functional equivalence (adventitious roots of rooted cuttings developing complete functionality supporting either own-rooted or grafted vines), revealing production differences (own-rooted requiring single rooting step while grafted requiring rooting then grafting), and displaying market segments (own-rooted cuttings and grafted vines serving different market needs and regions). Rooted cutting adventitious root long-term performance includes= showing establishment success (well-rooted cuttings showing high field establishment rates confirming propagation quality), exhibiting growth patterns (rooted cutting-derived vines showing growth and productivity comparable to seed-derived or other propagation methods), demonstrating adventitious root persistence (primary adventitious roots persisting throughout vine life as main structural roots), revealing secondary root development (adventitious roots producing extensive lateral and tertiary root systems creating complete architecture), and displaying lifetime function (rooted cutting adventitious roots serving as permanent foundation of root system throughout vineyard life).
Rooted_Cutting_Adventitious_Root emphasizes successfully rooted state and functional capacity and is distinguished from unrooted cuttings by focus on established root system. Critical for understanding rooting success, quality assessment, physiological independence, transplant readiness, commercial standards, and the successfully rooted adventitious root system enabling cutting establishment as independent plant representing commercial propagation product.
Bewurzelte Stecklings-Adventivwurzel
Erfolgreich bewurzelter Steckling
Established cutting root
Functional propagation root
Funktionelle Vermehrungswurzel
Successfully rooted cutting
Rooted Cutting Adventitious Root
A rooted cutting of 110 Richter with five well-distributed primary roots bears a Rooted_Cutting_Role as rootstock liner, destined for field planting and T-budding with Cabernet Sauvignon scion the following summer—completing the two-year field-budded vine production cycle. A rooted cutting of Riparia Gloire serves in a Rooted_Cutting_Role for German rootstock liner production, field-rooted in nursery rows during the first growing season for harvest and grading before the budding operation. A rooted cutting of Thompson Seedless bears a Rooted_Cutting_Role for own-rooted table grape production in California sandy soils, where phylloxera absence permits cultivation without grafting.
A nursery plant role that inheres in a grapevine produced by rooting a hardwood or softwood cutting without grafting, representing either a rootstock liner (for subsequent field budding) or an own-rooted fruiting vine (for regions without phylloxera or for rootstock variety production). Rooted cuttings develop adventitious root systems from cambial and parenchymal tissues when cutting bases are exposed to appropriate rooting conditions—field soil, nursery beds, containers, or controlled-environment facilities depending on production system. Rooted cutting quality is assessed by root system development (number of primary roots, typically minimum 3-5 well-distributed roots for commercial grade), root distribution (radial symmetry preferred), root health (freedom from disease or physical damage), and shoot development (viable apical bud, adequate caliper). Rooted cuttings serve as the input for two-year field-budding nursery systems (rootstock liners budded in year one, harvested as grafted vines in year two) or as finished products for own-rooted vineyard establishment where grafting is unnecessary (phylloxera-free regions, sandy soils, or where own-rooted production is traditional).
Rooted_Cutting_Role captures the simplest nursery product—a cutting that has developed roots without the complexity of grafting—serving either as an intermediate product (rootstock liner) or finished product (own-rooted vine). The category's importance varies regionally in Europe where grafting is universal, rooted cuttings are exclusively rootstock liners, while in phylloxera-free regions (parts of Australia, Chile, some US areas) own-rooted fruiting vines remain commercially significant. Rooting success varies dramatically by genotype—V. rupestris and V. riparia parentage confers easy rooting (>80%), while V. berlandieri parentage makes rooting difficult (<40% without treatment)—determining which rootstock varieties are commercially viable for cutting propagation. Quality grading establishes minimum root standards (typically 3-5 roots of specified minimum length and diameter) for commercial sale.
Unterlagsrebe
Wurzelrebe
bewurzelter Steckling
own-rooted vine
rooted cutting
rooted vine
rootstock liner
wurzelechte Rebe
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 344-380 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Pongrácz DP (1983) Rootstocks for Grape-vines, David Philip Publishers Whiting JR (2004) Grapevine rootstocks, Winetitles
Rooted Cutting Role
Nursery propagation success varies dramatically among rootstock genotypes, from easy-rooting St. George (V. rupestris, 90-95% rooting) through moderate SO4 (V. berlandieri × V. riparia, 70-80% rooting) to difficult-rooting 41B (V. vinifera × V. berlandieri, 30-50% rooting requiring special treatment). This variation determines nursery production costs and rootstock availability difficult-rooting genotypes command premium prices and may have limited commercial supply. Freedom (1613C × Dog Ridge) shows easy rooting (85-90%) combining good rooting from both parents, contributing to its widespread adoption despite moderate phylloxera resistance.
A rootstock role that inheres in a grafted rootstock by virtue of its genetically determined capacity to initiate adventitious roots from hardwood cuttings during nursery propagation, a critical trait determining commercial viability and propagation economics. Rooting capacity reflects the ability of dormant cane tissue to dedifferentiate cambial and parenchyma cells, form root primordia, and develop functional adventitious root systems when cuttings are planted under appropriate conditions (typically in heated callusing beds followed by nursery field or greenhouse growth). Rooting is quantified by the percentage of cuttings successfully developing roots under standardized conditions, with values ranging from below 30% (difficult rooting) through 50-70% (moderate) to above 80% (easy rooting). Rooting capacity is determined by endogenous auxin-to-cytokinin ratios, carbohydrate reserves, anatomical features (ray parenchyma abundance, vessel distribution), and wound response physiology. Poor rooting capacity historically limited the commercial adoption of otherwise desirable genotypes, most notably V. berlandieri, until hybridization with easier-rooting species provided commercial propagation solutions.
Rooting_Capacity_Rootstock_Role is subclassed into Easy_Rooting, Moderate_Rooting, and Difficult_Rooting categories reflecting the practical importance of propagation success for commercial rootstock production. V. rupestris parentage generally contributes easy rooting, V. riparia intermediate rooting, and V. berlandieri very difficult rooting—a key constraint historically limiting the direct use of lime-tolerant V. berlandieri selections. Rooting can be enhanced through auxin treatments (IBA, NAA), mist propagation, bottom heat, and tissue culture, but inherent genetic capacity remains the primary determinant. Climate conditions during cutting collection (carbohydrate status, cold acclimation) also influence rooting success.
Bewurzelungsfähigkeit-Unterlage
Stecklingsbewurzelungs-Unterlage
Vermehrungsfähigkeits-Unterlage
adventitious rooting rootstock
cutting rooting rootstock
propagation capacity rootstock
rooting capacity rootstock
Hartmann HT et al. (2011) Hartmann and Kester's Plant Propagation: Principles and Practices 8th Ed, Pearson Smart DR et al. (2006) Grapevine rooting patterns: a comprehensive analysis and a review, American Journal of Enology and Viticulture 57: 89-104 Whiting JR (2004) Grapevine rootstocks, in Viticulture Vol 1 Resources, Winetitles pp.: 167-188 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia
Rooting Capacity Rootstock Role
Pinot Noir grafted on 3309C rootstock at 16 years post-grafting showing rootstock annual rings visible as 16 concentric bands in trunk cross-section, ring widths ranging 0.8 cm (years 1-3 rapid juvenile growth) declining to 0.5 cm (years 4-10 establishing phase) to 0.3 cm (years 11-16 mature stable growth), ring 12 notably narrow 0.2 cm indicating drought stress in that year (confirmed by weather records showing severe summer drought), most recent ring (year 16) showing 0.35 cm width indicating good current vigor, each ring showing distinct earlywood (light tan with large vessels 95-120 micrometers) and latewood (darker tan with small vessels 50-70 micrometers) zones, ring boundaries clear enabling accurate age determination, ring pattern recording complete 16-year trunk development history from establishment through current season
Rootstock annual ring is the wood increment produced by vascular cambium during one complete growing season in rootstock trunk, typically visible as distinct concentric band in trunk cross-section marked by transition from large-diameter thin-walled earlywood vessels formed in spring to smaller-diameter thick-walled latewood vessels and fibers formed in late summer, showing radial width typically 0.2-1.0 cm varying with rootstock vigor, growing conditions, and trunk age, representing one year of secondary xylem production adding to trunk diameter, and providing dendrochronological record preserving information about growing season length, water availability, temperature patterns, and overall vine vigor during that specific calendar year throughout trunk developmental history. Rootstock annual ring formation process includes= initiated by spring cambial activation (vascular cambium resuming cell division in April-May Northern Hemisphere producing first xylem cells of growing season), showing peak production in early growing season (most rapid xylem cell production May-July creating majority of annual ring width), continuing through summer (cambial activity persisting through August producing progressively smaller cells with thicker walls), declining in late season (cell production slowing September-October as cambium prepares for dormancy), and ceasing in fall-winter (cambium entering dormancy November-March producing no new cells until following spring). Rootstock annual ring structure includes= comprising earlywood zone (inner portion of ring formed in spring showing large-diameter vessels 80-180 micrometers and thin-walled cells optimized for water transport), transitioning to latewood zone (outer portion of ring formed in late summer showing smaller vessels 40-80 micrometers and thick-walled fibers optimized for mechanical strength), showing gradual or abrupt transition (earlywood-latewood boundary potentially gradual with progressive size reduction or abrupt with sharp change depending on growing conditions), displaying boundary with previous ring (distinct line marking end of previous year latewood and beginning of current year earlywood creating visible ring), and spanning radial width (annual ring extending from inner boundary with previous ring to outer boundary with next ring typically 0.2-1.0 cm radial distance). Rootstock annual ring visibility includes= appearing as concentric band in cross-section (annual ring forming circular or elliptical band around trunk visible when cut transversely), showing contrasting zones (lighter colored earlywood and darker latewood creating visible pattern), displaying distinct boundaries (ring boundaries marked by vessel size change and latewood-earlywood transition visible as lines), enabling ring counting (distinct rings allowing determination of trunk age by counting rings from pith to cambium), and varying in prominence (ring visibility varying with species, growing conditions, and ring width with some rings more distinct than others). Rootstock annual ring width variation includes= showing typical range 0.2-1.0 cm (most rootstock trunk annual rings falling in this range depending on vigor and conditions), exhibiting young trunk pattern (wide rings 0.5-1.0 cm in young rapidly growing trunk years 1-10 reflecting vigorous juvenile growth), displaying mature trunk pattern (moderate rings 0.3-0.6 cm in mature trunk years 10-40 reflecting stable mature growth rate), showing old trunk pattern (narrow rings 0.1-0.3 cm in old trunk years 40+ reflecting age-related growth decline), and demonstrating year-to-year variation (ring width varying annually reflecting each growing season's specific conditions). Rootstock annual ring and growing conditions includes= reflecting water availability (wide rings in wet years with adequate soil moisture supporting vigorous cambial activity, narrow rings in drought years with limited water constraining growth), showing temperature effects (favorable temperatures promoting wide rings while extreme heat or cold producing narrow rings), exhibiting nutrient influences (adequate fertility supporting wide ring formation while deficiency limiting ring width), demonstrating stress impacts (favorable stress-free years producing wide rings while pest, disease, or environmental stress creating narrow rings), and recording management effects (irrigation, fertilization, and cultural practices influencing ring width through effects on growth conditions). Rootstock annual ring earlywood characteristics includes= formed in spring (April-June typically) when cambium first resumes activity, showing large vessel diameter (vessels 80-180 micrometers optimized for maximum water transport capacity during leaf expansion and shoot growth), exhibiting thin cell walls (cells with minimal secondary wall deposition maximizing lumen size for water conduction), displaying light coloration (earlywood appearing lighter tan in cross-section due to large vessel lumens and thin walls), comprising 40-60% of total ring width (earlywood typically forming majority of annual ring representing spring and early summer growth), and providing majority of water transport (large earlywood vessels conducting most water due to flow rate increasing with fourth power of vessel radius). Rootstock annual ring latewood characteristics includes= formed in late summer (July-September typically) as growing season progresses, showing small vessel diameter (vessels 40-80 micrometers prioritizing mechanical strength over maximum conductivity as growth slows), exhibiting thick cell walls (cells with extensive secondary wall deposition creating strong dense tissue), displaying dark coloration (latewood appearing darker brown in cross-section due to thick walls and small lumens reducing void space), comprising 40-60% of total ring width (latewood forming outer portion of ring representing late summer and fall wood production), and providing mechanical strength (dense thick-walled latewood contributing disproportionately to wood strength and rigidity). Rootstock annual ring in trunk cross-section includes= appearing as concentric circles (rings forming series of nested circles around central pith each representing one year), showing increasing number with age (ring count equaling trunk age in years from grafting or planting), displaying variable widths (rings showing different widths reflecting variable growing conditions across years), revealing growth history (ring width pattern recording vigor changes, stress events, and management effects throughout trunk life), exhibiting symmetry or asymmetry (rings potentially circular in uniform growth or eccentric in uneven growth from light exposure or mechanical stress), and preserving in both sapwood and heartwood (rings remaining visible after sapwood-heartwood conversion maintaining structural pattern). Rootstock annual ring boundaries includes= marked by latewood-earlywood transition (sharp or gradual change from previous year dense small-celled latewood to current year large-celled earlywood), appearing as distinct lines (ring boundaries visible as fine lines in cross-section creating concentric pattern), showing variable prominence (some ring boundaries very distinct while others subtle depending on growing conditions and earlywood-latewood contrast), enabling age determination (counting ring boundaries from pith to cambium revealing trunk age), and occasionally showing false rings (intra-annual bands from mid-season stress potentially mimicking annual boundaries requiring careful interpretation). Rootstock annual ring width and trunk age includes= showing widest rings in young trunk (juvenile rings often 0.5-1.0 cm reflecting rapid early growth), displaying progressively narrower rings with age (ring width generally declining as trunk matures and growth rate slows), exhibiting narrowest rings in old trunk (aged trunk rings potentially 0.1-0.2 cm reflecting minimal diameter increase in old age), demonstrating age-width relationship (strong negative correlation between trunk age and recent ring width across most trunks), and showing exceptions to pattern (some old trunks maintaining moderate ring width with good vigor and conditions while some young trunks showing narrow rings under stress). Rootstock annual ring and rootstock cultivar includes= showing cultivar-specific patterns (vigorous rootstocks producing wider rings than low-vigor rootstocks under similar conditions), exhibiting characteristic ring anatomy (vessel size distribution and earlywood-latewood contrast varying with rootstock genetics), displaying typical width ranges (1103P and 110R producing wide rings 0.5-0.9 cm while Riparia producing narrow rings 0.3-0.5 cm reflecting vigor differences), demonstrating genetic growth rates (ring width reflecting rootstock variety characteristic growth intensity), and revealing cultivar identification potential (ring patterns potentially aiding rootstock identification in mature trunk when combined with other wood characteristics). Rootstock annual ring dendrochronological value includes= recording annual growing conditions (each ring preserving information about that year's climate, water, temperature, and stress), enabling trunk age determination (ring count providing precise age assessment without destructive sampling when increment cores used), revealing growth history (ring width sequence showing vigor changes, stress periods, and recovery throughout trunk life), documenting management effects (ring patterns showing irrigation implementation, fertilization changes, or other management modifications), allowing cross-dating (matching ring patterns across multiple trunks or sites enabling climate reconstruction and vineyard history analysis), and preserving long-term records (rings maintaining information for decades or centuries in very old trunks creating valuable historical archive). Rootstock annual ring sampling and analysis includes= collected via increment boring (extracting thin core samples from pith to cambium without felling trunk enabling non-destructive analysis), counted for age determination (manually or digitally counting rings from pith outward to cambium determining trunk years since establishment), measured for width analysis (determining radial width of each ring revealing annual growth rates and patterns), analyzed for density variation (measuring earlywood and latewood density within rings assessing wood quality and strength), examined for anatomical features (studying vessel size, cell wall thickness, and other ring characteristics revealing growing conditions and tree physiology), and cross-dated across samples (comparing ring patterns across multiple cores or trees synchronizing dating and validating age assessment). Rootstock annual ring in sapwood versus heartwood includes= showing identical ring structure (rings preserving anatomical features equally well in functional sapwood and non-functional heartwood), maintaining visibility after heartwood conversion (rings remaining clearly visible after sapwood ages to heartwood despite color change and extractive deposition), demonstrating functional difference (rings in sapwood participating in water transport while rings in heartwood providing only structural support), exhibiting age difference (outer rings near cambium young sapwood while inner rings near pith old heartwood possibly decades old), and preserving complete record (rings spanning both zones providing continuous record from trunk establishment through current year). Rootstock annual ring environmental response patterns includes= showing wet year signature (wide rings with large earlywood vessels and extensive latewood indicating favorable water availability), exhibiting drought year signature (narrow rings with reduced earlywood and minimal latewood indicating water stress), displaying temperature effects (unusually warm or cool years potentially creating characteristic ring patterns), revealing frost damage (spring frost potentially affecting earlywood vessel formation creating detectable anomalies), demonstrating defoliation response (pest or disease defoliation potentially producing narrow ring from reduced photosynthate availability), and showing recovery patterns (return to normal ring width following stress indicating vine recovery and improved conditions). Rootstock annual ring false ring phenomenon includes= occurring from mid-season stress (drought, defoliation, or other stress followed by recovery potentially producing intra-annual band mimicking true ring), showing earlywood-like cells within ring (false ring marked by band of larger vessels within single growing season resembling earlywood), requiring careful analysis (distinguishing false rings from true annual rings critical for accurate age determination requiring examination of ring continuity and transition patterns), demonstrating environmental cause (false rings typically associated with specific stress events during growing season identifiable in climate records), and affecting age estimation (false rings potentially leading to age overestimation if not recognized requiring skilled interpretation). Rootstock annual ring in trunk capacity assessment includes= indicating recent growth rate (width of most recent 3-5 rings showing current trunk vigor and growth capacity), revealing age-related changes (ring width trends showing whether trunk in vigorous mature phase or declining old phase), demonstrating site suitability (consistent moderate ring widths suggesting appropriate rootstock-site matching while extremely narrow rings suggesting poor adaptation), showing stress history (ring patterns revealing whether trunk experienced significant stresses potentially compromising current condition), and predicting future growth (recent ring width trends potentially indicating trajectory toward continued vigor or progressive decline). Rootstock annual ring measurement techniques includes= using digital imaging (photographing cross-section or core and measuring rings digitally with specialized software), employing manual measurement (measuring ring widths directly with graduated microscope or caliper for detailed analysis), applying automated ring detection (computer algorithms identifying and measuring rings in scanned images enabling rapid analysis), conducting density analysis (X-ray densitometry or similar techniques measuring wood density variations within and across rings), and performing stable isotope analysis (measuring carbon and oxygen isotopes in ring cellulose revealing physiological responses and environmental conditions during formation). Rootstock annual ring in old vine assessment includes= indicating true age (ring count providing definitive age assessment critical for old vine certification or valuation), revealing longevity factors (ring patterns showing conditions and vigor levels throughout vine life explaining successful aging), demonstrating historical vigor (ring widths over decades showing whether old vine maintained consistent vigor or experienced progressive decline), recording historical events (ring patterns potentially showing frost years, drought cycles, management changes, or other events in vineyard history), and supporting heritage value (rings in ancient vine preserving century-scale record of vineyard history contributing to cultural and historical significance).
Rootstock_Annual_Ring emphasizes single year's growth increment and is distinguished from Rootstock_Wood (all secondary xylem) by focus on one-year production unit. Critical for understanding annual cambial activity, earlywood-latewood structure, trunk age determination, growth rate assessment, dendrochronological records, environmental response patterns, and the fundamental growth unit recording one growing season's xylem production and conditions.
Annual xylem increment
Jährlicher Xylemzuwachs
Rootstock growth ring
Rootstock tree ring
Unterlagen-Jahresring
Wurzelstock-Wachstumsring
Rootstock Annual Ring
Merlot grafted on SO4 rootstock at 18 years showing rootstock bark 5.2 mm total thickness, inner bark 2.1 mm comprising living secondary phloem with functional sieve elements conducting photosynthates and phloem parenchyma storing reserves, outer bark 3.1 mm comprising dead cork layers providing waterproof protection, bark showing SO4-characteristic moderate longitudinal fissures with grey-brown weathered color, lenticels 2-3 mm diameter visible in outer bark enabling gas exchange, bark texture and color typical of mature SO4 rootstock
Rootstock bark is all tissues external to vascular cambium in rootstock trunk comprising inner bark (living secondary phloem with sieve elements, companion cells, fibers, and parenchyma conducting photosynthates and storing reserves) and outer bark (dead protective tissues including cork, phelloderm, and isolated phloem providing waterproofing and mechanical protection), typically 2-8 mm total thickness in mature trunk increasing with age, showing cultivar-specific texture, color, and fissuring patterns characteristic of rootstock variety, serving dual functions of food transport in inner bark and trunk surface protection in outer bark, and undergoing continuous renewal as vascular cambium produces new phloem inward while cork cambia produce new protective layers and old outer bark sloughs from surface. Rootstock bark structural organization includes= inner bark comprising living secondary phloem (functional food-conducting tissue between vascular cambium and innermost cork cambium typically 1-3 mm thick), outer bark comprising protective dead tissues (cork, old phloem, and phelloderm outside innermost cork cambium typically 1-5 mm thick), showing radial zonation (progressive transition from cambium through functional phloem to protective cork outward), demonstrating continuous turnover (new bark added at cambium while old bark sloughs from surface), and displaying age-related thickening (bark accumulating 0.2-0.8 mm annually increasing total thickness as trunk matures). Rootstock bark inner bark functions includes= conducting photosynthates from scion to roots (phloem sieve elements transporting sugars, amino acids, and other organic compounds bidirectionally), storing carbohydrate reserves (phloem parenchyma and ray cells accumulating starch particularly during dormancy), providing mechanical support (phloem fibers contributing to bark strength and flexibility), facilitating lateral transport (ray parenchyma extending radially through bark enabling communication between inner and outer tissues), and maintaining metabolic activity (living phloem cells respiring, synthesizing, and responding to signals). Rootstock bark outer bark functions includes= protecting against water loss (suberized cork cells creating waterproof barrier preventing trunk desiccation), blocking pathogen entry (dead cork layers providing physical and chemical barrier against fungi and bacteria), insulating against temperature extremes (cork low thermal conductivity protecting cambium from freeze and heat stress), resisting mechanical damage (tough outer bark protecting trunk from abrasion, impact, and animal injury), and enabling gas exchange (lenticels in cork allowing oxygen and CO2 diffusion maintaining respiration in living bark tissues). Rootstock bark thickness and development includes= showing age-related increase (young trunk 1-3 mm bark, mature trunk 3-8 mm, old trunk 5-15 mm reflecting progressive accumulation), exhibiting cultivar variation (Rupestris types with thick bark 6-12 mm, Riparia types with thin bark 3-6 mm, genetic determination of bark thickness), demonstrating position effects (bark typically thicker on trunk base than upper portions reflecting longer accumulation period), displaying environmental influences (sun-exposed bark potentially thicker than shaded bark, stress affecting bark development), and showing management impacts (mechanical damage, pruning wounds, or disease affecting bark thickness and integrity). Rootstock bark texture and appearance includes= displaying cultivar-specific patterns (smooth, moderately furrowed, deeply fissured, or plated bark characteristic of rootstock variety), showing color variation (tan, brown, grey, or reddish tones depending on cork oxidation and cultivar chemistry), exhibiting age-related changes (young bark smooth and light-colored, old bark furrowed and dark-colored reflecting weathering and cork accumulation), demonstrating weathering effects (outer bark oxidizing, cracking, and sloughing creating characteristic aged appearance), and revealing environmental exposure (sun-exposed bark often darker and more fissured than shaded bark).
Rootstock_Bark emphasizes all tissues external to vascular cambium and is distinguished from Rootstock_Wood (xylem tissues internal to cambium) by focus on phloem and protective layers. Includes both Inner_Rootstock_Bark (living phloem) and Outer_Rootstock_Bark (dead protective tissues). Critical for understanding food transport, carbohydrate storage, trunk protection, bark development, and the essential protective and conducting tissues external to vascular cambium.
Periderm and phloem
Rootstock cortex
Stamm-Rinde
Trunk bark
Unterlagenrinde
Wurzelstock-Borke
Rootstock Bark
Pinot Noir grafted on Riparia Gloire rootstock at 10 years showing rootstock cambium in trunk cross-section, vascular cambium visible as thin 3-6 cell layer between xylem (inner wood 3.8 cm radius) and phloem (outer bark), vascular cambium cells small rectangular in radial view with thin walls and dense cytoplasm indicating meristematic character, cork cambium located in outer bark approximately 2 mm from trunk surface producing current periderm, cambium dormant in winter sample (January) showing no cell division, cambium producing 0.42 cm xylem and proportional phloem previous growing season documenting annual cambial productivity
Rootstock cambium is the lateral meristem tissue in rootstock trunk comprising undifferentiated dividing cells responsible for secondary growth increasing trunk diameter, including vascular cambium positioned between xylem and phloem producing secondary xylem inward and secondary phloem outward, and cork cambium (phellogen) positioned in outer bark producing cork (phellem) outward and phelloderm inward, showing active cell division during growing season and dormancy during winter, enabling continuous radial trunk expansion throughout vine life, and representing fundamental meristematic tissue distinguishing woody plants from herbaceous plants by capacity for indefinite secondary diameter increase. Rootstock cambium types includes= vascular cambium (lateral meristem between xylem and phloem producing secondary vascular tissues increasing trunk diameter through wood and inner bark formation), cork cambium or phellogen (lateral meristem in outer bark producing periderm protective layers replacing epidermis as trunk expands), showing distinct positions (vascular cambium deep in trunk between wood and phloem, cork cambium superficial in bark producing protective covering), exhibiting different products (vascular cambium producing water-conducting xylem and food-conducting phloem, cork cambium producing protective waterproof cork), and demonstrating coordinated function (both cambia contributing to trunk development with vascular cambium expanding core diameter and cork cambium maintaining protective bark). Rootstock cambium cellular characteristics includes= comprising undifferentiated meristematic cells (cambial initials retaining capacity for continued division and differentiation), showing thin primary walls (cambium cells with minimal wall thickening maintaining flexibility for division), exhibiting small cell size (cambial cells relatively small compared to mature xylem or phloem cells), displaying dense cytoplasm (meristematic cells with abundant organelles and minimal vacuolation), and demonstrating perpendicular division planes (cambium dividing periclinally producing derivatives inward and outward or anticlinally increasing cambial circumference). Rootstock cambium seasonal activity includes= activating in spring (cambium resuming cell division April-May as temperatures rise and growth resumes), showing peak activity in early growing season (most rapid cambial division May-June producing majority of annual xylem increment), declining in late summer (cambial activity slowing August-September as season concludes), entering dormancy in fall (cambium ceasing division October-November becoming quiescent for winter), and reactivating annually (cambium maintaining capacity for renewed activity each spring throughout trunk life). Rootstock cambium and secondary growth includes= producing annual diameter increase (cambial activity adding wood and bark layers each year expanding trunk circumference), creating annual rings (vascular cambium products forming visible rings in xylem recording years of growth), enabling indefinite expansion (cambium maintaining meristematic capacity allowing continued trunk enlargement for decades or centuries), accommodating expanding circumference (cambium increasing own cell number through anticlinal divisions maintaining continuous cylinder as trunk diameter increases), and supporting long-term development (cambial activity sustaining trunk growth throughout vine life from establishment through old age). Rootstock cambium position in trunk includes= vascular cambium located between xylem and phloem (forming complete cylinder separating inner wood from outer bark), cork cambium positioned in outer bark (arising at various depths in phloem or cortex producing successive periderms), showing coordinated expansion (both cambia increasing diameter maintaining appropriate proportions as trunk grows), demonstrating depth difference (vascular cambium several millimeters to centimeters deep while cork cambium near trunk surface), and enabling layered structure (cambial activities creating characteristic woody plant anatomy with xylem core, cambial zone, phloem, and protective periderm layers).
Rootstock_Cambium emphasizes lateral meristematic tissue and is distinguished from apical meristems (shoot and root tips) by focus on radial secondary growth. Includes both Rootstock_Vascular_Cambium (xylem-phloem producer) and Rootstock_Cork_Cambium (periderm producer). Critical for understanding secondary growth, diameter increase, annual ring formation, bark development, and the fundamental meristematic tissues enabling woody trunk development throughout vine life.
Cambial tissue
Rootstock lateral meristem
Rootstock secondary meristem
Sekundärmeristem
Unterlagen-Kambium
Wurzelstock-Lateralmeristem
Rootstock Cambium
Syrah grafted on SO4 rootstock at 12 years showing rootstock cork cambium, outermost active cork cambium located 1.8 mm beneath bark surface in secondary phloem producing current periderm, cork cambium zone 3-4 cells radial thickness including phellogen initials and recent derivatives, cork production outward creating 1.2 mm phellem layer with suberized walls, phelloderm production inward creating 0.3 mm living parenchyma layer, two deeper inactive cork cambia visible at 3.5 mm and 5.8 mm depths marking previous periderms, successive cork cambium activity creating characteristic SO4 moderately furrowed grey-brown bark
Rootstock cork cambium (phellogen) is the lateral meristem in rootstock trunk bark producing protective periderm layers comprising cork (phellem) cells outward and phelloderm cells inward, typically arising at depth of 0.5-3 mm beneath bark surface in cortex or secondary phloem, forming complete cylinder around trunk circumference, showing intermittent activity producing cork layers during growing season with periodic dormancy, enabling bark development and trunk surface protection as trunk expands, undergoing successive formation as older cork cambia become non-functional and new deeper cork cambia arise producing multiple periderms throughout trunk life creating characteristic furrowed bark of mature grapevine trunk. Rootstock cork cambium cellular structure includes= comprising rectangular meristematic cells (cork cambium initials typically 15-30 micrometers dimension in radial view), showing single cell layer (phellogen proper one cell thick though cambial zone including recent derivatives several cells wide), exhibiting thin walls (cambial cells with primary walls maintaining division capacity), displaying moderate cytoplasm (less dense than vascular cambium but adequate for division and differentiation), and demonstrating periclinal divisions (dividing parallel to trunk surface producing cork cells outward and phelloderm cells inward). Rootstock cork cambium products includes= producing phellem (cork) outward (generating dead cells with suberized walls creating waterproof protective barrier), generating phelloderm inward (creating living parenchyma cells forming inner periderm layer), showing unequal production (producing more cork than phelloderm typically 3-10 times more phellem than phelloderm by volume), creating periderm (cork cambium and products constituting complete periderm protective system replacing epidermis), and forming lenticels (cork cambium producing loosely arranged cells creating gas exchange pores in bark). Rootstock cork cambium position and depth includes= arising at variable depth (first cork cambium typically 0.5-1.5 mm beneath surface in cortex or outer phloem), showing deeper successive cambia (subsequent cork cambia forming progressively deeper 1-5 mm in secondary phloem as trunk ages), maintaining circumferential continuity (forming complete ring around trunk providing uniform protective layer), demonstrating depth increase with age (older trunk showing cork cambium deeper in bark as successive periderms form), and creating layered structure (multiple cork cambia at different depths creating complex layered bark in old trunk). Rootstock cork cambium formation timing includes= initiating in young trunk (first cork cambium arising year 1-3 as trunk begins secondary growth and epidermis ruptures), producing successive cambia (new cork cambia forming every 3-8 years as previous cambia become non-functional and trunk continues expanding), showing continuous replacement (cork cambium formation ongoing throughout trunk life maintaining protective covering), demonstrating age-related pattern (young trunk showing 1-2 cork cambia, mature trunk 3-5, old trunk 5-10+ successive layers), and enabling bark development (successive cork cambium activity creating characteristic mature bark texture and thickness). Rootstock cork cambium seasonal activity includes= showing growing season activity (cork cambium producing cork layers during April-October growing season), exhibiting intermittent pattern (periodic active phases alternating with quiescent periods even within growing season), demonstrating peak production (maximum cork formation typically spring and early summer May-June), declining in late season (reduced activity September-October as dormancy approaches), and entering winter dormancy (cork cambium quiescent November-March producing no new cells). Rootstock cork cambium and bark development includes= creating outer bark (cork cambium products accumulating as protective dead bark tissue sloughing from surface), forming bark texture (successive cork cambia creating ridges, furrows, and characteristic bark patterns), determining bark thickness (cork production rate and periderm number determining total bark depth 2-15 mm depending on age and cultivar), enabling expansion accommodation (new cork cambia arising as trunk diameter increases maintaining protective coverage), and producing cultivar-specific bark (cork cambium activity patterns creating characteristic bark appearance for rootstock variety). Rootstock cork cambium functional lifespan includes= showing limited duration (individual cork cambium functioning typically 3-8 years before becoming inactive), requiring successive replacement (new cork cambia arising deeper to replace non-functional older cambia), demonstrating positional progression (successive cambia forming progressively deeper in bark as trunk ages), creating buried cambia (old non-functional cork cambia embedded in inner bark beneath active cambium), and enabling continued protection (successive cork cambium formation maintaining bark integrity throughout trunk life despite individual cambium limited lifespan).
Rootstock_Cork_Cambium emphasizes periderm-producing lateral meristem and is distinguished from Rootstock_Vascular_Cambium (xylem-phloem producer) by focus on protective bark formation. Critical for understanding bark development, periderm formation, trunk surface protection, successive cambium replacement, and the lateral meristem producing protective cork layers maintaining bark integrity as trunk expands.
Cork-producing meristem
Rinden-Kambium
Rootstock phellogen
Unterlagen-Korkkambium
Wurzelstock-Phellogen
Rootstock Cork Cambium
A 30 cm three-node hardwood cutting of SO4 (V. berlandieri × V. riparia) bears a Rootstock_Cutting_Role, disbudded along its length, graded at 9-11 mm diameter for omega graft compatibility, and destined for benchgrafting to Chardonnay scion in a commercial French nursery. A 40 cm cutting of 110 Richter (V. berlandieri × V. rupestris) serves in a Rootstock_Cutting_Role for direct field rooting in a California vineyard, planted deeply with only the apical bud exposed to establish drought-tolerant root systems before T-budding with Cabernet Sauvignon the following summer. A rootstock cutting of Ramsey (V. champinii) bears a Rootstock_Cutting_Role in Australian table grape production, rooted as liner stock for subsequent green grafting to Thompson Seedless.
A cutting role that inheres in a stem segment harvested from a rootstock variety (typically an interspecific Vitis hybrid or American species selection) by virtue of its function as either the root-forming component of a grafted vine or a standalone rooted rootstock for subsequent field budding. Rootstock cuttings provide the below-ground portion of grafted vines, contributing pest resistance (phylloxera, nematodes), stress tolerance (drought, lime, salinity), and vigor modification to the composite plant while accepting the scion variety that determines fruit characteristics. Rootstock cutting preparation involves harvest from certified rootstock mother plants during dormancy, grading by diameter (typically 7-12 mm for benchgrafting, with uniformity essential for mechanical grafting compatibility), cutting to standard length (28-32 cm for benchgrafts, longer for direct field planting), disbudding to prevent rootstock shoot development competing with scion growth, and storage under controlled conditions (0-4°C, 95%+ RH) until use. Rootstock genotype determines critical vineyard adaptation characteristics—variety selection matching rootstock to site conditions (soil type, pest pressure, climate) is among the most consequential decisions in vineyard establishment.
Rootstock_Cutting_Role captures the functional designation of cutting material destined for root system formation, distinguishing it from Scion_Wood_Role material providing the fruiting variety. Rootstock cutting quality directly influences benchgraft success and subsequent vine performance—diameter uniformity is critical for mechanical grafting systems (omega, whip-and-tongue) where matching scion and rootstock diameters ensures cambial alignment. Disbudding removes all rootstock buds except potentially the basal node (which may produce shoot growth useful for vine training if the graft union is planted at or below soil level). Rootstock cutting identity verification is essential because variety misidentification can result in vineyard failure when the wrong rootstock encounters challenging site conditions molecular marker genotyping increasingly supplements visual identification for rootstock certification.
Unterlagensteckling
Unterlagsrebe-Steckling
Wurzelstockholz
rootstock cutting
rootstock propagation wood
rootstock stem piece
understock cutting
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson Whiting JR (2004) Grapevine rootstocks, in Viticulture Vol 1 Resources, Winetitles pp.: 167-188 Ollat N et al. (2016) Grapevine rootstocks origins and perspectives, Acta Horticulturae 1136: 11-22 Gramaje D, Armengol J (2011) Fungal trunk pathogens, Phytopathologia Mediterranea 50: S217-S226
Rootstock Cutting Role
Cabernet Sauvignon grafted on SO4 rootstock at 24 years post-grafting showing rootstock heartwood 5.8 cm diameter in trunk cross-section (total trunk diameter 10.2 cm), heartwood comprising central core with approximately 56% of trunk cross-sectional area, heartwood medium brown color distinctly darker than surrounding light tan sapwood, sapwood-heartwood boundary at approximately annual ring 14 indicating heartwood formation began around year 14 post-grafting, heartwood showing completely occluded vessels with tyloses blocking all lumens eliminating water transport function, heartwood moisture content 45% of dry weight much lower than sapwood 92%, heartwood specific gravity 0.61 g/cm³ slightly higher than sapwood 0.57 g/cm³, heartwood providing structural support bearing majority of trunk compressive loads, no living cells present in heartwood indicating complete physiological inactivity
Rootstock heartwood is the non-functional inner wood zone in rootstock trunk comprising aged secondary xylem where vessels have become occluded through tyloses formation eliminating water transport capacity, parenchyma cells have died removing metabolic activity, and extractive compounds have accumulated creating darker coloration and increased decay resistance, typically beginning formation at years 8-15 post-grafting when innermost sapwood converts to heartwood and progressively expanding as trunk ages, providing essential mechanical support bearing structural loads while no longer conducting water or storing reserves, and showing rootstock cultivar-specific characteristics in color, density, extractive content, and rate of formation reflecting genetics and environmental history. Rootstock heartwood formation process includes= developing through sapwood aging (innermost sapwood progressively converting to heartwood as new sapwood forms at periphery maintaining functional sapwood ring), showing tyloses formation (ray parenchyma cells producing balloon-like outgrowths into adjacent vessels blocking lumens and eliminating water transport capacity), exhibiting extractive deposition (phenolic compounds, tannins, gums, and other secondary metabolites accumulating in aging wood cells creating heartwood characteristics), experiencing parenchyma cell death (living ray and axial parenchyma cells dying during conversion eliminating all metabolic activity and storage function), and displaying irreversible transformation (sapwood-to-heartwood conversion permanent with heartwood never reverting to functional sapwood status). Rootstock heartwood position and extent includes= located in trunk center (heartwood occupying central core surrounded by peripheral functional sapwood ring), beginning at pith (heartwood formation starting at trunk center progressively expanding outward), showing variable radial extent (heartwood diameter ranging from absent in young trunk to 50-80% of total trunk diameter in old trunk), displaying circular or elliptical cross-section (heartwood typically forming regular shape though potentially eccentric in trunks with uneven growth), and bounded by sapwood-heartwood interface (distinct transition zone marking boundary between non-functional heartwood and functional sapwood). Rootstock heartwood non-functional status includes= showing completely occluded vessels (all xylem vessels blocked by tyloses, gums, or other deposits eliminating water conduction capacity), containing only dead cells (all parenchyma cells dead removing any metabolic activity, respiration, or synthetic function), lacking carbohydrate storage (no living cells present eliminating capacity for starch accumulation or reserve mobilization), exhibiting no transport function (heartwood contributing nothing to water or mineral conduction from roots to scion), and demonstrating no physiological activity (heartwood completely inert with no growth, defense responses, or other active processes). Rootstock heartwood mechanical function includes= providing structural support (heartwood cells with lignified walls contributing significantly to trunk rigidity and load-bearing capacity), bearing compressive loads (heartwood in trunk center supporting weight of vine, canopy, and fruit through compression resistance), resisting bending forces (heartwood contributing to trunk stiffness preventing excessive flexure under wind or crop load), supporting upright position (heartwood mass and rigidity helping maintain vertical trunk orientation), and enabling long-term stability (heartwood structural capacity allowing trunk to support vine for decades despite loss of physiological function). Rootstock heartwood physical characteristics includes= showing darker coloration than sapwood (heartwood typically medium to dark brown contrasting with light tan sapwood due to extractive accumulation), displaying lower moisture content (heartwood 30-60% moisture compared to sapwood 80-120% due to non-conducting status), exhibiting higher density (heartwood specific gravity 0.52-0.68 g/cm³ often slightly higher than sapwood 0.48-0.62 g/cm³ due to extractive deposition), revealing distinct color boundary (sharp or gradual transition from light sapwood to dark heartwood marking functional-structural interface), and showing cultivar-specific colors (heartwood ranging from light tan to dark reddish-brown depending on rootstock variety and extractive chemistry). Rootstock heartwood extractive compounds includes= containing phenolic substances (various phenolic compounds depositing in heartwood cells contributing to color and decay resistance), accumulating tannins (condensed and hydrolyzable tannins common in heartwood providing antimicrobial properties), showing gums and resins (complex organic compounds accumulating in some heartwood types), exhibiting cultivar-specific chemistry (extractive composition and concentration characteristic of rootstock variety), and demonstrating protective function (antimicrobial and antifungal properties of extractives contributing to heartwood decay resistance). Rootstock heartwood decay resistance includes= showing improved durability compared to sapwood (extractive compounds providing chemical protection reducing decay organism growth), exhibiting cultivar variation (some rootstock varieties producing highly durable heartwood while others showing moderate resistance), displaying center-rot susceptibility in old trunks (very old heartwood particularly in damaged areas potentially showing decay despite extractive protection), demonstrating moisture effects (dry heartwood highly resistant to decay while consistently wet heartwood potentially vulnerable), and requiring wound protection (exposed heartwood vulnerable to decay entry requiring proper pruning and wound management). Rootstock heartwood development timeline includes= absent in very young trunk (years 1-7 typically showing no heartwood with all wood as functional sapwood), beginning formation in establishing trunk (years 8-15 showing initial heartwood development in centermost wood around pith), progressively expanding in mature trunk (years 15-40 showing steady heartwood diameter increase as inner sapwood converts and new sapwood forms outside), dominating old trunk volume (years 40-80+ showing large heartwood core potentially 60-80% of trunk cross-section with narrow sapwood ring), and reaching maximum proportion in ancient trunk (very old trunks showing massive heartwood with minimal functional sapwood ring). Rootstock heartwood in cross-section appearance includes= occupying central trunk position (heartwood forming inner core visible as darker colored wood in trunk center), showing concentric ring structure (annual rings visible in heartwood marking growth history though rings non-functional), displaying color gradation (heartwood often showing color variation from outer edge to center reflecting progressive aging and extractive accumulation), revealing pith at center (central pith marking original stem axis at heartwood core), and exhibiting distinct boundary (sapwood-heartwood interface visible as color change marking functional-structural transition). Rootstock heartwood annual ring structure includes= retaining visible annual rings (growth rings remain visible in heartwood preserving record of trunk growth history), showing earlywood and latewood (ring structure with spring-formed large vessels and summer-formed dense wood preserved in heartwood), displaying growth patterns (ring width variation in heartwood revealing historical growing conditions and trunk development), enabling age determination (counting rings in heartwood contributing to total trunk age assessment), and preserving dendrochronological record (heartwood rings recording environmental history throughout trunk life from early years forward). Rootstock heartwood tyloses formation includes= showing vessel occlusion (heartwood vessels completely blocked by tyloses eliminating all water transport capacity), exhibiting ray parenchyma origin (tyloses produced by living ray parenchyma cells in aging sapwood extending through pit pairs into adjacent vessel lumens), displaying balloon-like structures (tyloses appearing as bubble-like outgrowths filling vessel interiors visible in microscopic examination), demonstrating permanent blockage (tyloses formation irreversible creating permanent non-conducting status of heartwood vessels), and varying in abundance (tyloses density varying with wood anatomy, aging rate, and environmental conditions affecting completeness of vessel occlusion). Rootstock heartwood mechanical properties includes= showing high compressive strength (heartwood resistant to crushing forces supporting trunk structural loads), exhibiting good tensile strength (heartwood resisting pulling forces maintaining trunk integrity), displaying adequate stiffness (heartwood rigidity contributing to trunk resistance to bending and deflection), demonstrating brittleness (heartwood often more brittle than flexible sapwood due to age and dryness cracking more readily under sudden stress), and providing structural stability (heartwood mass and strength essential for trunk structural capacity particularly in large old trunks). Rootstock heartwood water relations includes= containing minimal water (heartwood moisture 30-60% much lower than sapwood due to non-conducting occluded vessel status), showing slow moisture equilibration (heartwood gaining or losing moisture slowly due to occluded vessels and dead cells limiting water movement), exhibiting hygroscopic behavior (heartwood moisture content varying with ambient humidity though changes slower than sapwood), displaying limited dimensional changes (heartwood swelling and shrinking less than sapwood due to lower moisture content and slower moisture exchange), and demonstrating dry-state stability (dry heartwood dimensionally stable resisting warping and checking compared to moisture-variable sapwood). Rootstock heartwood in different trunk ages includes= young trunk (years 1-7) showing no heartwood (entire wood cross-section functional sapwood), establishing trunk (years 8-15) showing initial small heartwood core (centermost 1-3 cm diameter converting to heartwood), mature trunk (years 15-40) displaying moderate heartwood (central core 3-8 cm diameter surrounded by 2-4 cm sapwood ring), old trunk (years 40-80) showing large heartwood (core 8-15 cm diameter with 2-3 cm sapwood ring), and ancient trunk exhibiting dominant heartwood (core 12-20+ cm diameter with very narrow 1.5-2.5 cm sapwood ring). Rootstock heartwood cultivar characteristics includes= Riparia types showing light tan to medium brown heartwood with moderate extractive content, Rupestris types displaying reddish-brown heartwood with high tannin levels, Berlandieri hybrids exhibiting medium brown heartwood with variable extractive content, vigorous rootstocks like 1103P showing rapid heartwood formation with large cores developing by years 12-15, low-vigor rootstocks like Riparia showing delayed heartwood formation potentially not appearing until years 10-12, SO4 displaying characteristic medium brown heartwood with moderate density, and 3309C showing light brown heartwood with gradual formation rate. Rootstock heartwood proportion changes with age includes= young establishing trunk showing 0-15% heartwood (little or no heartwood in rapidly growing young trunk), mature trunk showing 25-50% heartwood (heartwood core comprising significant but not dominant trunk portion), old trunk showing 50-75% heartwood (heartwood dominant with functional sapwood reduced to peripheral ring), very old trunk showing 70-85% heartwood (massive heartwood core with minimal sapwood), and showing progressive increase (heartwood proportion continuously increasing as trunk ages and sapwood converts). Rootstock heartwood and trunk capacity includes= providing no functional capacity (heartwood contributing nothing to water transport, nutrient conduction, or reserve storage), reducing functional tissue proportion (expanding heartwood decreasing percentage of trunk that is physiologically active), requiring adequate sapwood compensation (trunk must maintain sufficient sapwood ring despite heartwood expansion to support canopy), potentially limiting old trunk function (very large heartwood with minimal sapwood potentially constraining water supply in ancient trunk), and affecting trunk value assessment (trunk capacity determined by sapwood area not total diameter with large heartwood potentially misleading regarding functional capacity). Rootstock heartwood damage and decay includes= showing resistance to decay when sound (intact heartwood with extractives providing good protection against most decay organisms), exhibiting vulnerability when damaged (wounds exposing heartwood creating entry points for decay despite extractive protection), potentially developing center rot (very old heartwood particularly in damaged areas susceptible to decay causing hollow trunk centers), demonstrating compartmentalization importance (trunk ability to wall off heartwood decay limiting spread to sapwood and maintaining functional tissue), and requiring wound management (proper pruning and wound treatment protecting heartwood from decay entry). Rootstock heartwood in trunk structural role includes= bearing majority of structural load in old trunk (large heartwood core carrying most compressive and bending forces in aged vine), enabling trunk diameter increase without proportional sapwood increase (heartwood formation allowing large trunk diameter with functional sapwood ring maintaining adequate size), providing ballast and stability (heartwood mass contributing to trunk weight and resistance to wind displacement), supporting training system loads (heartwood strength adequate for bearing cordon systems, wire tension, and canopy weight), and maintaining structural integrity for decades (heartwood remaining mechanically sound supporting trunk function throughout vine life). Rootstock heartwood extractive and color variation includes= showing light colors in low-extractive cultivars (pale tan or light brown heartwood in rootstock varieties with limited phenolic accumulation), displaying dark colors in high-extractive cultivars (reddish-brown or dark brown heartwood in varieties producing abundant tannins and phenolics), exhibiting gradual color development (heartwood darkening progressively with age as extractives accumulate over years), revealing cultivar identification potential (heartwood color characteristic potentially aiding rootstock variety identification in mature trunk), and demonstrating environmental influences (growing conditions affecting extractive production and heartwood color development intensity). Rootstock heartwood moisture content dynamics includes= maintaining relatively stable low moisture (heartwood 30-60% moisture relatively constant compared to seasonally variable sapwood), showing slow seasonal variation (heartwood moisture changing slightly with seasonal humidity patterns but much less than sapwood), equilibrating slowly with environment (cut heartwood gaining or losing moisture gradually over weeks to months), demonstrating density effects (denser heartwood potentially holding slightly more moisture than lighter heartwood), and affecting wood properties (heartwood moisture level influencing dimensional stability, density, and mechanical properties). Rootstock heartwood significance includes= providing essential mechanical support (heartwood structural function critical for trunk load-bearing capacity particularly in large old vines), enabling trunk diameter increase (heartwood formation allowing trunk expansion while maintaining functional sapwood at manageable size), demonstrating aging process (heartwood development marking trunk maturation and physiological aging), showing cultivar expression (heartwood characteristics reflecting rootstock genetics in color, density, and formation rate), preserving growth history (heartwood rings recording complete trunk age and development throughout vine life), contributing to decay resistance (extractive compounds in heartwood protecting trunk from decay organisms).
Rootstock_Heartwood emphasizes non-functional inner wood zone and is distinguished from Rootstock_Sapwood (functional outer wood with open vessels and living cells) by focus on occluded vessels, dead cells, extractive accumulation, and purely structural rather than physiological function. Critical for understanding trunk aging, sapwood-to-heartwood conversion, structural support, mechanical properties, decay resistance, trunk capacity assessment, and the non-functional but structurally essential inner wood comprising increasing proportion of trunk volume as vine ages.
Nicht-funktionelles Unterlagenholz
Non-functional rootstock wood
Rootstock core wood
Rootstock duramen
Unterlagen-Kernholz
Wurzelstock-Duramen
Rootstock Heartwood
A 110 Richter (V. berlandieri × V. rupestris) vine in a California foundation rootstock block bears a Rootstock_Mother_Plant_Role, head-trained and spur-pruned annually to produce 8-12 one-meter canes per vine for hardwood cutting propagation—its hybrid parentage verified by molecular markers confirming V. berlandieri and V. rupestris ancestry. A SO4 (V. berlandieri × V. riparia) plant in a German VCR-certified rootstock increase block serves in a Rootstock_Mother_Plant_Role, producing cuttings for benchgraft production meeting EU certification standards. A Riparia Gloire (V. riparia selection) maintained at ENTAV's rootstock collection bears a Rootstock_Mother_Plant_Role, preserving this historically important rootstock for regions requiring low vigor and high lime sensitivity tolerance.
A mother plant role that inheres in a grapevine maintained specifically as a source of rootstock cuttings—the hardwood cane material used to propagate rootstocks either directly as rooted cuttings or as the root-forming component of benchgrafts. Rootstock mother plants are typically interspecific hybrids (V. berlandieri × V. riparia, V. berlandieri × V. rupestris, V. riparia × V. rupestris, etc.) or pure American species selections (Riparia Gloire, Rupestris du Lot) cultivated for their pest resistance, stress tolerance, and rootstock functional characteristics rather than fruit quality. Rootstock mother plant blocks are managed to maximize cane production—often head-trained or minimally trellised, with aggressive pruning encouraging vigorous vegetative growth rather than fruit production. Rootstock cuttings are typically harvested during dormancy as one-year-old canes cut to standard lengths (30-40 cm for benchgrafting, longer for field budding), with quality assessed by diameter (typically 7-12 mm), node count, and freedom from disease symptoms. Rootstock mother plant material is subject to the same certification hierarchy as scion material, with particular attention to phylloxera resistance verification and trueness-to-type confirmation for hybrid rootstocks whose parentage determines their functional characteristics.
Rootstock_Mother_Plant_Role captures the distinct functional category of mother plants providing root-forming material, complementing Scion_Mother_Plant_Role. Rootstock mother plant management differs from scion mother plants in several respects—rootstock varieties generally have different vigor characteristics, disease susceptibilities, and training preferences than V. vinifera scions. Hybrid rootstock identity verification is particularly important because functional traits (lime tolerance, drought resistance, vigor) depend on specific parentage misidentified rootstocks may fail in conditions where the correct genotype would succeed. The certification system addresses rootstock health (virus testing) and identity (molecular verification) separately. Major rootstock mother plant collections are maintained by certification authorities (FPS, ENTAV, IMIDRA) ensuring availability of tested material across the range of commercial rootstocks (SO4, 110R, 3309C, 5BB, 1103P, 140Ru, 101-14, etc.).
Unterlagen-Mutterpflanze
Unterlagsreben-Quelle
Wurzelstock-Mutterstock
rootstock cutting source
rootstock mother vine
rootstock source plant
understock mother plant
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson Whiting JR (2004) Grapevine rootstocks, in Viticulture Vol 1 Resources, Winetitles pp.: 167-188 Ollat N et al. (2016) Grapevine rootstocks origins and perspectives, Acta Horticulturae 1136: 11-22 Gramaje D, Armengol J (2011) Fungal trunk pathogens in the propagation process, Phytopathologia Mediterranea 50: S217-S226
Rootstock Mother Plant Role
Merlot grafted on 3309C rootstock at 12 years showing rootstock periderm, outermost active periderm located 1.6 mm beneath surface comprising cork layer 1.1 mm thick with suberized waterproof cells, cork cambium visible as thin meristematic layer, and phelloderm 0.3 mm thick with living parenchyma cells, two deeper inactive older periderms visible at 3.2 and 5.4 mm depths embedded in inner bark, lenticels 1-2 mm diameter scattered in cork enabling gas exchange, periderm system providing essential waterproofing and protection maintaining trunk integrity
Rootstock periderm is the protective tissue system in rootstock trunk bark produced by cork cambium (phellogen) comprising three components cork (phellem) produced outward providing waterproof dead protective layer, cork cambium itself as meristematic layer, and phelloderm produced inward forming living parenchyma layer, replacing epidermis as trunk undergoes secondary growth and progressively replacing older periderms through successive cork cambium formation creating multiple periderm layers in old bark, typically with outermost periderm being most recently formed active system located 0.5-2 mm beneath surface and deeper periderms being older inactive systems embedded in inner bark, showing coordinated function where cork provides waterproofing and protection, phelloderm contributes to storage and perhaps protection, and cork cambium maintains capacity to produce new tissues, and demonstrating that periderm represents fundamental protective tissue system characteristic of woody plants enabling continued secondary growth while maintaining trunk surface protection. Rootstock periderm three-component structure includes= comprising phellem (cork) outward (dead cells with suberized walls creating waterproof barrier typically 0.5-3 mm thickness), containing phellogen (cork cambium) itself (single-cell meristematic layer dividing periclinally producing cork and phelloderm), including phelloderm inward (living parenchyma cells forming thin layer typically 0.1-0.5 mm thickness), showing organized layering (three tissues arranged in consistent sequence from surface inward cork-cambium-phelloderm), and demonstrating integrated function (all three components contributing to overall periderm protective system). Rootstock periderm formation and development includes= initiating in young trunk (first periderm forming years 1-3 as epidermis ruptures from trunk expansion), arising from parenchyma cells (cork cambium originating from cortex or phloem parenchyma cells dedifferentiating into meristematic state), showing circumferential continuity (periderm forming complete ring around trunk providing uniform protection), producing cork and phelloderm (active cork cambium dividing to generate protective tissues), and becoming inactive eventually (individual periderm functioning 3-8 years before cork cambium ceases activity). Rootstock periderm successive formation includes= requiring replacement periderms (individual periderm finite lifespan necessitating formation of new deeper periderms), showing deeper origin (each successive periderm arising deeper in phloem than previous periderm), creating multiple layers (old trunk potentially showing 5-10+ successive periderms creating complex layered bark), demonstrating continuous protection (new periderm forming before old periderm completely fails maintaining uninterrupted trunk protection), and producing characteristic bark (successive periderm formation creating typical mature grapevine bark texture and thickness). Rootstock periderm cork properties includes= showing suberized cells (cork cells with suberin deposition in walls creating waterproof lipid-rich barrier), exhibiting dead cell status (cork cells dying at maturity leaving hollow air-filled cells), demonstrating low density (cork specific gravity 0.12-0.25 g/cm³ creating lightweight protection), displaying thermal insulation (cork low conductivity protecting cambium from temperature extremes), and providing mechanical protection (cork tissue resisting compression and impact protecting underlying tissues). Rootstock periderm phelloderm characteristics includes= comprising living parenchyma cells (phelloderm cells remaining alive with functional protoplasts), showing storage function (cells potentially accumulating starch and other reserves), forming modest layer (phelloderm typically much thinner than cork usually only 2-8 cell layers), demonstrating anatomical similarity to cortex (phelloderm resembling cortical parenchyma in structure), and contributing to inner periderm (phelloderm forming living inner portion of periderm system). Rootstock periderm lenticel formation includes= creating gas exchange pores (lenticels comprising loosely arranged cork cells enabling oxygen and CO2 diffusion), showing scattered distribution (lenticels appearing at intervals across bark surface), exhibiting variable morphology (lenticels appearing as dots, lines, raised bumps, or cracks depending on formation pattern), demonstrating essential function (lenticels enabling respiration in living bark and cambium tissues behind impermeable cork), and displaying cultivar characteristics (lenticel size, shape, and density potentially characteristic of rootstock variety). Rootstock periderm protective functions includes= preventing water loss through waterproof cork, blocking pathogen entry through physical cork barrier, insulating against temperature extremes through cork thermal properties, resisting mechanical damage through cork toughness, and enabling gas exchange through lenticels maintaining tissue respiration behind waterproof barrier.
Rootstock_Periderm emphasizes three-component protective tissue system and is distinguished from individual components (cork, cork cambium, phelloderm) by focus on integrated system. Critical for understanding bark protective function, cork-cambium-phelloderm organization, successive periderm formation, lenticel development, and the fundamental protective tissue system replacing epidermis enabling continued secondary growth.
Cork system
Korkschicht-System
Protective tissue system
Unterlagen-Periderm
Wurzelstock-Schutzgewebesystem
Rootstock Periderm
SO4 (Selection Oppenheim 4, V. berlandieri × V. riparia) bears a Rootstock_Role when grafted with Riesling in Rheingau vineyards, providing moderate vigor and good lime tolerance (up to 17% active lime). 110 Richter (V. berlandieri × V. rupestris) bears a Rootstock_Role in Mediterranean vineyards, conferring drought tolerance through deep root penetration exceeding 3-4 m depth. 3309 Couderc (V. riparia × V. rupestris) serves in a Rootstock_Role in Burgundy, imparting low-to-moderate vigor and early fruit maturity to Pinot Noir scions.
A grapevine role that inheres in a plant of the genus Vitis or an interspecific Vitis hybrid by virtue of its function as the root system and lower trunk portion of a grafted grapevine composite. The rootstock role is conferred when a plant or plant part is designated to provide belowground anchorage, water absorption, mineral nutrient acquisition, and environmental stress mediation to a grafted scion partner. Rootstock role bearers influence scion vigor, yield, fruit composition, and phenological timing through species-specific root architecture, hydraulic conductance (typically 50-300 mg H₂O·m⁻²·s⁻¹·MPa⁻¹), and phytohormone translocation patterns. The rootstock role emerged historically following the phylloxera crisis (Daktulosphaira vitifoliae) in European vineyards from the 1860s onward, when grafting Vitis vinifera scions onto resistant American Vitis species became standard viticultural practice worldwide.
Rootstock_Role is subclassed by functional characteristics: vigor modification, stress resistance (biotic and abiotic), nutrient acquisition efficiency, parentage-based classification, rooting capacity, and graft compatibility. A single rootstock cultivar typically bears multiple sub-roles simultaneously. The rootstock role is distinct from the biological disposition of the root system itself it captures the relational function within the grafted composite system. Approximately 80-85% of the world's commercial vineyards use grafted vines.
Pfropfunterlage
Rebunterlage
Unterlage
Wurzelunterlage
grafting stock
rootstock
stock
understock
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 1-15 Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22 Corso M, Bonghi C (2014) Grapevine rootstock effects on abiotic stress tolerance, Plant Science 228: 1-14 Heinitz CC et al. (2019) Crop wild relatives of grape, in North American Crop Wild Relatives Vol 2, Springer pp.: 329-351
Rootstock Role
Merlot grafted on Teleki 5C rootstock at 22 years post-grafting showing rootstock sapwood 3.2 cm radial thickness in trunk cross-section, sapwood comprising outer 10 annual rings (rings 13-22 counting from pith) representing wood formed past 10 years remaining functional, sapwood light tan color clearly contrasting with darker brown heartwood interior, sapwood showing open vessels 100-140 micrometers diameter typical for Teleki 5C with high vessel density providing adequate hydraulic conductivity, sapwood area 45.2 square cm calculated from trunk diameter supporting estimated 60-90 liters daily water transport capacity adequate for mature canopy, living ray parenchyma in sapwood storing carbohydrate reserves, sapwood moisture content 95% of dry weight during mid-summer
Rootstock sapwood is the functional outer wood zone in rootstock trunk comprising living and recently formed secondary xylem with open unoccluded vessels actively conducting water and dissolved minerals from root system to scion, typically forming ring 2-5 cm radial thickness in outer trunk surrounding inner heartwood or extending to pith in young trunk lacking heartwood, characterized by lighter color than heartwood due to absence of extractive deposits, living ray parenchyma cells storing carbohydrates, and serving as essential conduit enabling 50-100+ liters daily water transport supporting productive vine while also providing mechanical support and reserve storage capacity. Rootstock sapwood functional definition includes= representing all functional water-conducting xylem in rootstock trunk (sapwood comprising secondary xylem zone actively transporting water from roots upward to scion canopy and fruit), showing open patent vessels (xylem vessel elements with open lumens free of blockages enabling efficient water flow through negative pressure-driven transpiration stream), containing living cells (ray parenchyma and some axial parenchyma remaining alive in sapwood storing reserves and maintaining tissue function), exhibiting recent formation (sapwood consisting of most recently produced annual rings typically most recent 5-15 years depending on trunk age and growing conditions), and maintaining physiological activity (sapwood tissue metabolically active participating in transport, storage, and defense responses unlike inert heartwood). Rootstock sapwood position and extent includes= located in outer trunk region (sapwood forming peripheral wood zone immediately inside vascular cambium extending inward to heartwood boundary or pith), showing variable radial thickness (sapwood depth typically 2-5 cm in mature trunk varying with rootstock vigor, trunk age, and environmental conditions), forming complete ring (sapwood surrounding trunk circumference creating continuous conducting pathway around entire trunk periphery), transitioning to heartwood at inner boundary (sapwood-heartwood interface marking zone where vessels occlude and cells die converting functional sapwood to structural heartwood), and extending to cambium at outer boundary (sapwood outer limit at vascular cambium where new xylem cells continuously produced adding to sapwood zone). Rootstock sapwood vessel characteristics includes= containing numerous open functional vessels (hollow tubular conduits formed from dead vessel elements arranged end-to-end creating continuous water transport pathways), showing unoccluded vessel lumens (vessel interiors free of tyloses, gums, or other blockages maintaining open pathways for water flow), displaying rootstock-specific vessel diameters (vessel size characteristic of rootstock cultivar typically 60-180 micrometers with vigorous rootstocks showing larger vessels enabling higher flow rates), exhibiting vessel density patterns (number of vessels per unit area varying with position in annual ring and rootstock genetics affecting total transport capacity), and demonstrating efficient vessel arrangement (distribution and connectivity of vessels optimized for minimizing resistance to water flow through sapwood network). Rootstock sapwood water transport function includes= conducting water from roots to scion (primary function moving 50-100+ liters daily during growing season through transpiration-driven negative pressure gradient), transporting dissolved minerals (delivering nitrogen, phosphorus, potassium, calcium, magnesium, and micronutrients absorbed by roots upward to leaves and fruit), showing high hydraulic conductivity (sapwood vessel structure enabling efficient water flow with minimal resistance maintaining adequate water supply to canopy), responding to transpiration demand (water flow rate through sapwood adjusting dynamically to match evaporative water loss from leaves), exhibiting seasonal variation (transport activity highest during active growing season declining during dormancy when transpiration minimal), and maintaining reserve capacity (total sapwood conducting area typically exceeding minimum required for average demand providing buffer for peak transpiration during hot periods). Rootstock sapwood living tissue components includes= containing living ray parenchyma (radially oriented cells remaining alive in sapwood storing carbohydrates and facilitating lateral transport between vessels and rays), potentially including living axial parenchyma (some longitudinally oriented parenchyma cells also remaining alive in sapwood particularly in outer most recent annual rings), showing metabolic activity (living cells respiring, synthesizing compounds, and maintaining tissue function unlike dead cells in heartwood), requiring oxygen supply (living sapwood cells needing oxygen for respiration obtained through gas exchange in ray system), and maintaining defensive capacity (living cells capable of responding to wounding or infection through chemical defense compound synthesis and secretion). Rootstock sapwood carbohydrate storage includes= storing starch in ray parenchyma (living ray cells accumulating carbohydrate reserves during late summer and fall particularly in outer sapwood layers), showing seasonal storage cycles (carbohydrate levels highest in late fall after photosynthate accumulation declining during spring when reserves mobilized for growth), providing reserves for cambial activity (stored carbohydrates supporting vascular cambium cell division and new xylem and phloem production), enabling stress response (sapwood reserves supporting wound healing, disease defense, and recovery from drought, freeze, or other environmental stress), and exhibiting substantial capacity (mature trunk sapwood potentially storing 80-300+ grams dry weight carbohydrates depending on sapwood volume and trunk size). Rootstock sapwood in young versus mature versus old trunk includes= young trunk (years 1-7) showing sapwood extending to pith (all wood functional with no heartwood formation yet), establishing trunk (years 8-15) beginning sapwood-heartwood differentiation (innermost wood converting to heartwood while outer sapwood ring maintains 2-4 cm thickness), mature trunk (years 15-40) displaying stable sapwood ring (sapwood thickness stabilized at 2.5-4.5 cm surrounding expanding heartwood core), old trunk (years 40-80+) potentially showing proportionally narrower sapwood (sapwood ring 2-3 cm surrounding large heartwood core as new sapwood added at periphery while inner sapwood converts to heartwood), and ancient trunk possibly showing minimal sapwood (very narrow sapwood ring 1.5-2.5 cm in extremely old trunk limiting water transport capacity). Rootstock sapwood color and appearance includes= showing lighter color than heartwood (sapwood typically light tan, cream, or pale brown contrasting with darker brown heartwood due to absence of extractive compounds), displaying moist appearance (sapwood higher moisture content than heartwood creating fresher look in freshly cut wood), exhibiting visible vessels (vessel openings visible as small pores on sapwood surface particularly in cross-section), showing clear annual ring structure (growth rings visible in sapwood as concentric bands marked by earlywood and latewood zones), and revealing ray patterns (radial rays visible as fine lines or flecks in sapwood creating characteristic wood grain pattern). Rootstock sapwood moisture content includes= containing high water content (sapwood moisture 60-120% of dry weight during growing season much higher than heartwood 30-50%), showing seasonal variation (moisture content highest during active growing season declining during dormancy), displaying rapid drying (cut sapwood losing moisture quickly when exposed to air causing dimensional changes and potential cracking), exhibiting capillary water (water held in vessel lumens and cell walls maintaining wood hydration), and demonstrating moisture gradient (moisture typically highest in outer sapwood declining toward sapwood-heartwood boundary). Rootstock sapwood mechanical properties includes= providing structural support (sapwood wood cells with lignified walls contributing to trunk rigidity and load-bearing capacity), showing lower density than heartwood (sapwood specific gravity 0.48-0.60 g/cm³ typically slightly lower than heartwood 0.52-0.65 g/cm³), exhibiting flexibility (sapwood somewhat less brittle than heartwood due to higher moisture content allowing more elastic deformation), contributing to trunk strength (sapwood cells particularly fibers providing tensile and compressive strength maintaining trunk structural integrity), and demonstrating adequate load-bearing (sapwood capable of supporting typical canopy, cordon, and fruit load despite being less dense than heartwood). Rootstock sapwood conversion to heartwood includes= undergoing physiological aging (sapwood progressively converting to heartwood through extractive deposition and vessel occlusion as new sapwood forms at periphery), showing extractive accumulation (phenolic compounds, tannins, and other substances depositing in aging sapwood cells initiating heartwood transformation), exhibiting vessel occlusion (tyloses forming in vessels blocking lumens as sapwood converts to heartwood eliminating water transport function), experiencing cell death (ray and axial parenchyma cells dying during conversion creating completely non-living heartwood tissue), and maintaining structural role (converted tissue retaining mechanical support function while losing transport and storage functions). Rootstock sapwood damage response includes= compartmentalizing injuries (living sapwood cells responding to wounding by forming chemical and anatomical barriers isolating damage), producing tyloses (living ray parenchyma cells generating balloon-like structures blocking vessels around wound sites preventing pathogen spread), synthesizing defensive compounds (living sapwood cells producing antimicrobial phenolics, tannins, and other compounds in threatened tissue), forming wound wood (modified xylem tissue with altered structure developing from cambium around injury sites), and showing variable success (compartmentalization effectiveness varying with rootstock genetics, injury severity, and pathogen aggressiveness). Rootstock sapwood transport efficiency includes= showing high hydraulic conductivity (vessel structure and arrangement enabling efficient water flow with minimal resistance), exhibiting rootstock-specific capacity (maximum transport rate varying with sapwood cross-sectional area, vessel size distribution, and rootstock genetics), responding to environmental conditions (transport efficiency affected by temperature, water potential gradient, and cavitation risk during stress), maintaining adequate supply under demand (sapwood transport capacity typically sufficient to support full canopy transpiration during normal conditions), and showing vulnerability to disruption (transport function compromised by freeze damage, drought-induced cavitation, or vascular diseases blocking or damaging vessels). Rootstock sapwood freezing and cavitation includes= vulnerable to freeze-thaw cavitation (ice crystal formation during freezing and subsequent thaw potentially introducing air embolisms blocking vessels), showing seasonal embolism repair (refilling of embolized vessels during spring recovery restoring transport capacity after winter damage), exhibiting drought-induced cavitation (severe water stress causing vessel collapse or air-seeding creating non-functional embolized vessels), demonstrating cultivar differences (rootstock varieties varying in cavitation resistance and embolism repair capacity affecting stress tolerance), and requiring vessel redundancy (multiple parallel transport pathways ensuring continued function despite some vessel embolism). Rootstock sapwood in different rootstock cultivars includes= Riparia types showing thin sapwood (typically 2-3 cm) with smaller vessels, Rupestris types displaying moderate sapwood thickness (2.5-4 cm) with medium vessels, Berlandieri hybrids exhibiting thicker sapwood (3-5 cm) with larger vessels, vigorous rootstocks like 1103P showing broad sapwood zones with large vessels supporting high transport capacity, low-vigor rootstocks like Riparia showing narrower sapwood with smaller vessels limiting transport capacity, SO4 displaying moderate sapwood 2.5-3.5 cm with characteristic medium vessel size, and 3309C showing sapwood 2-3 cm thickness with fine vessel distribution. Rootstock sapwood analysis and measurement includes= determining sapwood thickness through visual examination (measuring radial distance from cambium to heartwood boundary in cross-section or core sample), estimating sapwood area from trunk diameter (calculating functional conducting area supporting capacity assessment), assessing vessel characteristics through microscopy (measuring vessel diameter, density, and distribution analyzing transport properties), evaluating hydraulic conductivity through flow measurements (quantifying water transport capacity under standard pressure gradients), and determining moisture content through oven-drying (measuring water content indicating sapwood hydration status and seasonal patterns). Rootstock sapwood significance includes= providing essential water transport (sapwood vessels conducting water and minerals from roots to scion canopy enabling photosynthesis and growth), enabling adequate transport capacity (sapwood cross-sectional area and vessel structure determining trunk's ability to support productive vine), storing vital carbohydrate reserves (living ray parenchyma maintaining energy stores supporting vine resilience), contributing to mechanical support (sapwood cells providing structural strength maintaining trunk integrity), responding to damage and stress (living sapwood tissue compartmentalizing injuries and synthesizing defenses), and limiting vine capacity when reduced (narrow sapwood in aged trunk or damaged sapwood from disease potentially constraining vine water supply and productivity).
Rootstock_Sapwood emphasizes functional water-conducting outer wood zone and is distinguished from Rootstock_Wood (all wood including non-functional heartwood) and Rootstock_Heartwood (non-functional inner wood) by focus on actively transporting living tissue. Critical for understanding water transport capacity, hydraulic function, reserve storage, trunk functional anatomy, vessel characteristics, and the essential conducting tissue enabling scion canopy and fruit production.
Active rootstock wood
Funktionelles Wurzelstock-Holz
Leitendes Unterlagenholz
Rootstock conducting xylem
Rootstock functional wood
Unterlagen-Splintholz
Rootstock Sapwood
Cabernet Sauvignon grafted on SO4 rootstock showing rootstock trunk 8.2 cm diameter at 15 years post-grafting, rootstock trunk extending 22 cm from root crown to graft union (positioned 18 cm above soil surface), rootstock trunk bark grey-brown with moderate longitudinal furrowing characteristic of SO4, graft union showing complete integration with 9.5 cm diameter bulge, scion trunk above graft union 7.4 cm diameter (smaller than rootstock trunk), rootstock trunk conducting 45-65 liters water daily during mid-summer peak transpiration
Rootstock trunk is the below-graft portion of grafted grapevine trunk originating from rootstock cultivar, representing the rootstock's aerial stem section between root system and graft union, typically 15-40 cm length in commercial vineyards, providing structural support, water and nutrient transport from roots to scion, and conferring rootstock characteristics including phylloxera resistance, nematode tolerance, vigor control, and soil adaptation to the complete vine. Rootstock trunk position and extent includes= beginning at root crown (transition zone where major structural roots diverge from trunk base, typically 2-5 cm below soil surface), extending upward to graft union (point where rootstock and scion tissues join, marked by visible bulge or scar at 10-40 cm above soil surface depending on grafting height), spanning the critical below-graft zone (entire rootstock aerial portion contributing rootstock genetics to vine performance), and representing rootstock cultivar expression (trunk characteristics reflecting rootstock variety's growth habits, bark texture, and wood properties). Rootstock trunk as rootstock aerial expression includes= manifesting rootstock cultivar traits (bark color, texture, lignification patterns, and growth vigor characteristic of rootstock variety such as SO4, 3309C, 1103P, Riparia Gloire, or others), maintaining rootstock identity (distinct from scion tissues above graft union, preserving rootstock genetic characteristics), expressing rootstock-specific phylloxera resistance (trunk tissues immune to phylloxera root form due to rootstock genetics), showing rootstock vigor characteristics (trunk diameter and growth rate reflecting rootstock's inherent vigor classification as low, moderate, or high), and developing rootstock-typical bark (bark thickness, color, and texture patterns characteristic of specific rootstock cultivar). Rootstock trunk structural functions includes= supporting scion weight (bearing load of scion trunk, cordons, shoots, and crop above graft union, typically 5-25 kg mature vine mass), providing mechanical stability (rigid lignified structure resisting wind stress and maintaining upright position), conducting water from roots to scion (xylem vessels transporting 10-100+ liters daily during growing season from root system through graft union to scion), transporting nutrients upward (xylem delivering nitrogen, phosphorus, potassium, and micronutrients absorbed by roots to scion tissues), and conducting photosynthates downward (phloem transporting carbohydrates from scion leaves through graft union to root system for storage and root growth). Rootstock trunk graft union integration includes= forming callus bridge at graft union (healing tissue connecting rootstock and scion vascular systems enabling continuous transport), establishing vascular continuity (xylem and phloem connections across graft union allowing bidirectional transport), potentially showing graft incompatibility signs (swelling, tissue separation, or poor integration if rootstock-scion combination incompatible), maintaining distinct tissue zones (rootstock trunk retaining rootstock cellular characteristics while scion above shows scion traits), and supporting long-term graft stability (mechanical and physiological integration sustaining vine for decades). Rootstock trunk diameter and growth includes= initial post-graft diameter 0.8-2.5 cm (year 1 grafted trunk typical range), diameter increase 0.3-0.8 cm annually (depending on rootstock vigor and growing conditions), mature diameter 4-10 cm (typical at 10-20 years post-grafting), old vine diameter 8-15+ cm (40-80+ year old rootstock trunks), and diameter often larger than scion trunk above (rootstock typically showing greater diameter than scion trunk creating characteristic taper at graft union). Rootstock trunk age-related changes includes= young rootstock trunk (years 1-3) showing smooth bark and rapid diameter expansion, establishing rootstock trunk (years 4-7) developing first bark fissures and stabilizing growth rate, mature rootstock trunk (years 8-25) showing well-developed bark, stable diameter, and complete graft integration, old rootstock trunk (years 30-60+) displaying thick furrowed bark, large diameter, and potential heartwood formation, and ancient rootstock trunk (70-100+ years) exhibiting very thick bark, minimal diameter increase, and extensive heartwood. Rootstock trunk bark characteristics includes= developing periderm (protective cork layers forming on trunk surface as secondary growth proceeds), showing rootstock-specific bark texture (smooth in Riparia types, moderately furrowed in SO4, deeply furrowed in Rupestris types), displaying rootstock characteristic colors (greys, browns, or reddish tones depending on rootstock variety), thickening with age (bark accumulating 2-8 mm depth in mature trunks, 5-15 mm in old trunks), and providing protection (barrier against desiccation, mechanical damage, temperature extremes, and pathogen entry). Rootstock trunk damage and disease includes= susceptibility to trunk diseases (Eutypa, Botryosphaeria, Phomopsis affecting rootstock trunk potentially more severely than scion in some rootstock varieties), vulnerability to freeze damage (severe winter cold potentially killing rootstock cambium especially in young trunks), crown gall susceptibility (Agrobacterium tumefaciens infections potentially forming galls on rootstock trunk at or near graft union), mechanical injury risks (equipment damage, animal browsing, or pruning wounds on rootstock trunk), and sunburn potential (southwest-facing rootstock trunk exposure potentially causing bark damage in hot climates). Rootstock trunk management implications includes= graft union positioning (keeping graft union 10-20 cm above soil surface preventing scion rooting and maintaining rootstock effects), trunk protection (wrapping or painting young rootstock trunks in hot climates preventing sunburn), disease monitoring (inspecting rootstock trunk for cankers, galls, or decay indicating trunk disease), sucker removal (eliminating shoots arising from rootstock trunk below graft union that would compete with scion), and rootstock evaluation (assessing rootstock trunk health, graft union integrity, and rootstock performance for vineyard management decisions). Rootstock trunk cultivar variation includes= Riparia types showing thin smooth bark and moderate vigor trunks, Rupestris types displaying thick deeply furrowed bark and compact growth, Berlandieri hybrids exhibiting vigorous growth and large diameter trunks, SO4 showing characteristic moderately furrowed grey-brown bark, 3309C displaying smooth bark transitioning to light furrowing, 1103P showing very vigorous growth with large diameter trunk and rough bark, and Teleki 5C showing moderate vigor with characteristic bark patterns. Rootstock trunk significance includes= providing essential rootstock contributions to complete vine (phylloxera resistance, vigor control, drought tolerance, lime tolerance, nematode resistance, and disease resistance depending on rootstock genetics), enabling scion-rootstock partnerships (mechanical and physiological integration allowing superior scion cultivars to grow on adapted rootstock in diverse soil and pest conditions), maintaining graft union integrity (robust rootstock trunk supporting stable long-term graft connection), and representing foundation of modern viticulture (rootstock trunks enabling cultivation of Vitis vinifera scion cultivars in phylloxera-infested soils worldwide since late 1800s).
Rootstock_Trunk emphasizes the below-graft aerial trunk portion originating from rootstock cultivar and is distinguished from Trunk (which may refer to entire trunk including both rootstock and scion portions) and Scion_Trunk (above-graft portion). Critical for understanding rootstock contributions, graft union relationships, rootstock-scion interactions, rootstock cultivar expression, and the fundamental architecture of grafted grapevines that dominates modern viticulture.
Below-graft trunk
Pfropfunterlage Stamm
Rootstock above-ground stem
Rootstock stem
Understok trunk
Unterlagestamm
Wurzelstock Stamm
Rootstock Trunk
Cabernet Sauvignon grafted on SO4 rootstock at 15 years showing rootstock vascular cambium as thin zone 5-7 cells radial thickness between xylem (6.2 cm diameter) and phloem (3 mm thick), cambium active in June sample showing recent periclinal divisions with xylem mother cells differentiating into earlywood vessels inward and phloem mother cells differentiating into sieve elements outward, cambium producing estimated 0.38 cm xylem increment and 0.8 mm phloem increment current season, fusiform initials visible as elongated cells 300-400 micrometers long in tangential section, cambium cells showing dense cytoplasm and thin primary walls characteristic of meristematic tissue
Rootstock vascular cambium is the lateral meristem between secondary xylem and secondary phloem in rootstock trunk comprising single layer of fusiform initials and ray initials that divide periclinally producing xylem mother cells inward differentiating into secondary xylem and phloem mother cells outward differentiating into secondary phloem, typically 3-8 cells thick in radial dimension including cambial initials and recent undifferentiated derivatives, showing active division during growing season producing annual xylem ring and corresponding phloem increment, entering dormancy during winter, and enabling continuous trunk diameter increase throughout vine life by producing majority of trunk mass as secondary vascular tissues accumulate year after year. Rootstock vascular cambium cellular organization includes= comprising fusiform initials (elongated cells 100-500 micrometers long oriented axially producing xylem and phloem elements), containing ray initials (shorter isodiametric cells producing ray parenchyma cells creating radial transport pathways), showing single-cell layer (cambium proper only one cell thick though cambial zone including recent derivatives several cells wide), exhibiting thin walls (cambial cells with primary walls lacking secondary thickening maintaining division capacity), and displaying dense cytoplasm (meristematic cells with abundant ribosomes, mitochondria, and active metabolism). Rootstock vascular cambium division patterns includes= dividing periclinally (divisions parallel to trunk surface producing derivatives inward toward xylem and outward toward phloem), showing unequal production (producing more xylem than phloem typically 4-10 times more wood than inner bark by volume), exhibiting periclinal asymmetry (fusiform initials dividing to produce one derivative and one replacement initial maintaining cambium while generating tissues), demonstrating anticlinal divisions (occasional divisions perpendicular to surface increasing cambial circumference as trunk diameter expands), and coordinating with rays (ray initials dividing to produce ray cells maintaining radial continuity through cambium). Rootstock vascular cambium seasonal activity includes= reactivating in spring (resuming division April-May triggered by warming temperatures, lengthening photoperiod, and hormonal signals from expanding buds), showing peak activity in early season (most rapid division May-June producing wide earlywood vessels and abundant phloem), declining in late summer (slowing division July-September producing smaller latewood vessels and reduced phloem), entering dormancy in fall (ceasing division October-November as temperatures drop and day length shortens), and remaining quiescent in winter (dormant November-March with no cell division until spring reactivation). Rootstock vascular cambium products includes= producing secondary xylem inward (generating wood cells including vessel elements, fibers, and axial parenchyma forming annual rings), generating secondary phloem outward (creating inner bark cells including sieve elements, companion cells, fibers, and parenchyma), showing quantitative asymmetry (producing much greater volume of xylem than phloem each year), creating annual increments (each growing season adding new xylem ring and corresponding phloem layer), and building trunk (cambial products accumulating over years creating massive xylem core and functional phloem ring). Rootstock vascular cambium position and extent includes= located between xylem and phloem (forming complete cylinder at interface between inner wood and outer bark), showing variable depth (cambium depth from trunk surface increasing as trunk diameter expands and bark thickens), maintaining continuity (forming unbroken cylinder around entire trunk circumference), connecting across graft union (cambium in rootstock continuous with cambium in scion at graft interface enabling integration), and expanding circumference (cambium increasing in circumference through anticlinal divisions as trunk radius increases). Rootstock vascular cambium hormonal regulation includes= responding to auxin (indole-3-acetic acid from shoot apex promoting cambial division and xylem differentiation), influenced by cytokinins (promoting cell division and phloem differentiation), affected by gibberellins (stimulating cambial activity and cell elongation), modulated by photoperiod (day length affecting timing of activation and dormancy entry), and showing temperature sensitivity (cambial activity threshold typically 10-15°C with optimal activity 20-25°C). Rootstock vascular cambium in trunk development includes= initiating from procambium (primary cambium in young stem transitioning to secondary vascular cambium as secondary growth begins), producing first annual ring (cambium creating initial wood and bark layer during establishment year), accumulating annual increments (cambium adding ring each year building trunk diameter progressively), maintaining activity for decades (cambium continuing division throughout trunk life potentially 50-100+ years), and enabling trunk expansion (cambial productivity allowing trunk to increase from small grafted stem to large mature trunk supporting productive vine).
Rootstock_Vascular_Cambium emphasizes xylem-phloem-producing lateral meristem and is distinguished from Rootstock_Cork_Cambium (periderm producer) by focus on secondary vascular tissue production. Critical for understanding trunk diameter increase, annual ring formation, wood-bark production, seasonal growth dynamics, and the fundamental meristem producing secondary xylem and phloem throughout trunk life.
Rootstock wood cambium
Rootstock xylem-phloem cambium
Unterlagen-Gefäßkambium
Vascular meristem
Vaskuläres Lateralmeristem
Wurzelstock-Holzkambium
Rootstock Vascular Cambium
Cabernet Sauvignon grafted on SO4 rootstock trunk at 16 years post-grafting showing rootstock wood cross-section 7.8 cm diameter with 16 distinct annual rings (each marking one growing season), outer sapwood zone 2.8 cm radial thickness (functional water-conducting tissue) light tan color with clearly visible vessels, inner heartwood core 2.2 cm diameter (non-functional structural tissue) darker brown color, sapwood-heartwood boundary at approximately ring 8-9 indicating heartwood formation began years 8-9 post-grafting, annual ring widths ranging 0.3-0.6 cm with narrower recent rings indicating mature trunk growth rate decline, wood showing SO4-characteristic moderate vessel size approximately 90-120 micrometers diameter, wood density approximately 0.58 g/cm³ typical for SO4, ray parenchyma visible as radial lines facilitating storage and lateral transport
Rootstock wood is the secondary xylem tissue comprising the internal woody structure of rootstock trunk, representing all lignified vascular tissue produced by vascular cambium in below-graft trunk portion, typically including both functional sapwood conducting water and nutrients plus non-functional heartwood providing structural support, spanning from pith to vascular cambium with cross-sectional area increasing annually, and showing rootstock cultivar-specific characteristics including vessel size, wood density, annual ring patterns, and growth rates that reflect rootstock genetics and influence trunk mechanical properties and transport capacity. Rootstock wood anatomical structure includes= comprising secondary xylem (all wood tissue produced by vascular cambium activity posterior to cambium), including vessel elements (water-conducting tubular cells arranged in longitudinal chains forming continuous transport pathways), containing fiber cells (thick-walled elongated cells providing mechanical strength and comprising majority of wood volume), incorporating ray parenchyma (radially oriented living cells storing carbohydrates and facilitating lateral transport and communication), showing annual ring structure (concentric growth layers marking each growing season visible in cross-section), and displaying pith to cambium organization (wood tissue spanning from central pith outward to actively producing vascular cambium at trunk periphery). Rootstock wood functional zones includes= typically consisting of outer sapwood (functional water-conducting xylem in outer trunk layers actively transporting water and dissolved minerals from roots to scion), transitioning to inner heartwood (non-conducting central wood where vessels occluded and cells dead providing structural support but no transport function), showing age-related zonation (youngest wood at periphery near cambium progressively older toward trunk center), displaying sapwood-heartwood boundary (transition zone where functional sapwood converts to non-functional heartwood through extractive deposition and vessel occlusion), and exhibiting variable sapwood thickness (functional wood layer thickness varying with trunk age, rootstock vigor, and environmental conditions typically 2-5 cm in mature trunk). Rootstock wood annual growth includes= forming annual rings (each growing season producing new wood layer visible as ring in cross-section), showing earlywood and latewood zones (spring-formed earlywood with large vessels for maximum water transport and summer-formed latewood with smaller vessels and thicker walls for strength), displaying growth rate variation (annual ring width varying 0.2-0.8 cm depending on rootstock vigor, growing conditions, and trunk age), exhibiting declining growth with age (young trunk showing wide annual rings 0.5-1.0 cm while old trunk showing narrow rings 0.1-0.3 cm), and recording environmental history (annual ring widths reflecting each year's growing conditions creating dendrochronological record in wood structure). Rootstock wood vessel characteristics includes= containing numerous vessels (hollow tubular structures formed from vessel elements arranged end-to-end creating continuous water transport pathways), showing rootstock-specific vessel diameters (vessel size characteristic of rootstock variety ranging 50-200 micrometers with vigorous rootstocks typically showing larger vessels), displaying vessel density patterns (number of vessels per unit area varying with rootstock genetics and position within annual ring), exhibiting tyloses formation (balloon-like structures blocking vessels during heartwood formation or disease response), and demonstrating vessel arrangement (distribution patterns of vessels within wood tissue affecting transport efficiency and mechanical properties). Rootstock wood mechanical properties includes= providing structural support (rigid lignified tissue supporting trunk upright position and bearing weight of scion, canopy, and crop), showing rootstock-specific wood density (mass per unit volume characteristic of rootstock variety typically 0.50-0.65 g/cm³ dry weight), exhibiting strength characteristics (resistance to bending, compression, and tension loads varying with wood structure, density, and moisture content), displaying flexibility properties (capacity for elastic deformation without breaking declining with age as wood matures and lignification increases), and demonstrating durability (resistance to decay, mechanical damage, and environmental stress depending on wood chemistry and structure). Rootstock wood water transport capacity includes= conducting water from root system to scion (xylem vessels moving water and dissolved minerals upward through trunk via transpiration-driven negative pressure), showing high transport efficiency (vessel structure optimized for minimal resistance to water flow enabling 50-100+ liters daily transport in mature trunk), displaying seasonal variation (transport capacity highest during active growing season declining during dormancy), exhibiting response to demand (transport rate adjusting to match transpirational water loss from scion canopy), and maintaining reserve capacity (sapwood area typically exceeding minimum required for average demand providing buffer for peak transpiration periods). Rootstock wood carbohydrate storage includes= storing reserves in ray parenchyma (living cells in rays accumulating starch and sugars particularly during dormancy), showing seasonal storage cycles (carbohydrate accumulation during late summer and fall depletion during spring growth and stress periods), providing energy for growth and stress response (stored reserves supporting cambial activity, wound healing, disease defense, and recovery from environmental stress), exhibiting substantial storage capacity (mature trunk ray parenchyma potentially storing 100-500+ grams dry weight carbohydrates depending on trunk size), and demonstrating rootstock-specific storage patterns (reserve accumulation and mobilization characteristics varying with rootstock variety genetics). Rootstock wood formation process includes= produced by vascular cambium (lateral meristem continuously dividing producing xylem cells inward and phloem cells outward), showing cell differentiation (cambium-produced cells elongating, thickening walls, and lignifying to form mature wood cells), exhibiting secondary wall deposition (cells adding thick lignified secondary walls inside primary wall creating rigid structure), undergoing programmed cell death (vessel elements and fibers dying at maturity leaving hollow functional structures), and accumulating progressively (each growing season adding new wood layer increasing trunk diameter and wood volume). Rootstock wood in young versus old trunk includes= young trunk (years 1-5) showing predominantly sapwood (minimal or no heartwood with nearly all wood functional), establishing trunk (years 5-15) beginning heartwood formation (innermost wood starting conversion to heartwood as sapwood ring expands), mature trunk (years 15-40) displaying sapwood-heartwood zonation (clear differentiation with functional sapwood ring surrounding central heartwood core), old trunk (years 40-80+) showing proportionally more heartwood (expanding heartwood core with proportionally narrower sapwood ring in aged trunk), and ancient trunk potentially showing minimal sapwood (very narrow functional sapwood ring surrounding large heartwood core in extremely old trunk). Rootstock wood response to damage includes= compartmentalizing injuries (sealing wounds through chemical and anatomical barriers preventing decay spread into healthy wood), forming wound wood (specialized tissue with modified structure developing around injury sites), producing tyloses (vessel-blocking structures forming in response to wounding or infection limiting pathogen movement), accumulating defensive compounds (phenolics, tannins, and other antimicrobial substances depositing in threatened tissue), and showing variable compartmentalization success (effectiveness of damage isolation varying with rootstock genetics, injury severity, and pathogen virulence). Rootstock wood rootstock cultivar characteristics includes= showing cultivar-specific vessel sizes (Riparia types typically showing smaller vessels 60-100 micrometers, Rupestris types showing medium vessels 80-120 micrometers, Berlandieri hybrids showing larger vessels 100-180 micrometers), displaying characteristic wood density (dense wood in Rupestris types, moderate density in hybrid rootstocks, lighter wood in Riparia types), exhibiting distinctive annual ring patterns (wide rings in vigorous rootstocks like 1103P, narrow rings in low-vigor rootstocks like Riparia), showing cultivar-specific colors (heartwood colors varying from light tan to dark brown depending on rootstock variety), and demonstrating characteristic growth rates (annual wood production reflecting rootstock vigor classification). Rootstock wood environmental influences includes= showing water availability effects (adequate water supporting wide annual rings with optimal vessel development, drought stress producing narrow rings with smaller vessels), displaying temperature impacts (growing season temperature affecting cambial activity and annual ring width), exhibiting nutrient effects (adequate nutrition supporting normal wood formation, deficiency potentially affecting cell wall development and lignification), responding to stress factors (environmental stress producing narrow rings, altered vessel patterns, or modified wood chemistry), and recording site history (wood structure preserving record of environmental conditions during each growing season). Rootstock wood in trunk cross-section includes= showing concentric ring structure (annual layers forming circular or elliptical patterns around central pith), displaying pith at center (central core of soft thin-walled cells marking original stem development), exhibiting radial rays (spoke-like bands of parenchyma cells extending from pith to cambium facilitating lateral transport and storage), showing sapwood ring (outer functional wood zone lighter colored than inner heartwood in species showing heartwood formation), and revealing growth irregularities (eccentric growth, compression wood, or other structural modifications reflecting mechanical stress or environmental factors). Rootstock wood decay resistance includes= showing rootstock-specific durability (cultivar variation in natural decay resistance affecting trunk longevity), exhibiting heartwood resistance (central heartwood often more decay-resistant than sapwood due to extractive accumulation), displaying chemical defenses (phenolic compounds, tannins, and other antimicrobial substances protecting wood from decay organisms), demonstrating compartmentalization capacity (ability to isolate and contain decay limiting damage spread), and showing vulnerability factors (wounds, cracks, or disease-compromised areas creating entry points for decay organisms). Rootstock wood and overall trunk function includes= providing mechanical foundation (wood structure supporting trunk upright bearing weight of vine), enabling water transport (sapwood vessels conducting water and minerals from roots to scion), storing carbohydrate reserves (ray parenchyma accumulating and mobilizing energy reserves), contributing to graft union integration (rootstock wood connecting with scion wood at graft union enabling vascular continuity), and determining trunk capacity (wood cross-sectional area and structure determining trunk's transport, storage, and support capabilities). Rootstock wood sampling and analysis includes= revealing trunk age through ring counting (number of annual rings indicating years since trunk establishment), showing growth history through ring width patterns (wide rings indicating favorable years narrow rings indicating stress years), indicating wood quality through density measurement (specific gravity reflecting structural properties and decay resistance), assessing heartwood formation through sapwood-heartwood measurement (proportion of functional versus structural wood indicating trunk aging status), and enabling rootstock identification through wood anatomy (vessel size, ray structure, and other anatomical features potentially diagnostic of rootstock variety). Rootstock wood significance includes= comprising majority of trunk volume (wood tissue constituting bulk of rootstock trunk mass and structure), providing essential transport function (sapwood vessels enabling water and nutrient delivery from root system to scion canopy), supporting trunk structural capacity (wood mechanical properties determining trunk strength and stability), storing vital carbohydrate reserves (ray parenchyma maintaining energy stores supporting vine resilience and productivity), reflecting rootstock genetics (wood characteristics expressing rootstock cultivar identity and traits), and recording trunk history (annual rings preserving record of growth patterns, environmental conditions, and trunk age throughout vine life).
Rootstock_Wood emphasizes the complete secondary xylem tissue of rootstock trunk and is distinguished from Rootstock_Sapwood (functional outer zone only) and Rootstock_Heartwood (non-functional inner zone only) by encompassing all wood tissue from pith to cambium. Critical for understanding trunk structure, water transport capacity, mechanical support, carbohydrate storage, rootstock cultivar expression, and the fundamental woody tissue providing structural and physiological foundation for grafted vine.
Pfropfunterlagen-Holzgewebe
Rootstock secondary xylem
Rootstock woody tissue
Rootstock xylem
Unterlagenholz
Wurzelstock-Xylem
Rootstock Wood
Syrah grafted on 3309C rootstock at 14 years showing rootstock xylem in trunk cross-section comprising 14 annual rings total diameter 6.8 cm, outer sapwood xylem (rings 8-14) 2.6 cm radial thickness showing open functional vessels conducting water and containing living ray parenchyma, inner heartwood xylem (rings 1-7) 1.6 cm diameter showing occluded vessels with tyloses and dead parenchyma providing structural support only, earlywood xylem in each ring showing large vessels 90-115 micrometers diameter, latewood xylem showing smaller vessels 45-65 micrometers and thick-walled fibers, xylem conducting estimated 55-80 liters water daily supporting canopy transpiration
Rootstock xylem is the water-conducting vascular tissue comprising all secondary xylem in rootstock trunk produced by vascular cambium, including vessel elements arranged end-to-end forming continuous tubular conduits, fiber cells providing mechanical strength, and ray parenchyma facilitating radial transport and storage, organized into annual rings showing earlywood formed in spring with large vessels optimized for water transport and latewood formed in late summer with smaller vessels and thick-walled cells optimized for strength, spanning from central pith through heartwood and sapwood zones to peripheral vascular cambium, and serving dual functions of conducting water and dissolved minerals from roots to scion while providing structural support for trunk. Rootstock xylem cellular composition includes= vessel elements (dead cells with perforated end walls stacked vertically creating continuous water transport tubes 80-180 micrometers diameter), fiber cells (elongated thick-walled dead cells 10-30 micrometers diameter providing majority of wood volume and mechanical strength), ray parenchyma (living cells arranged radially facilitating lateral transport and storing carbohydrates), and axial parenchyma (living cells oriented longitudinally interspersed among vessels and fibers supporting storage and defense). Rootstock xylem functional organization includes= outer sapwood xylem (functional water-conducting zone with open vessels and living parenchyma actively transporting water and storing reserves), inner heartwood xylem (non-functional zone with occluded vessels and dead parenchyma providing only structural support), earlywood xylem (spring-formed large-vessel tissue prioritizing water transport), and latewood xylem (late-summer-formed small-vessel thick-walled tissue prioritizing mechanical strength). Rootstock xylem vessel characteristics includes= showing rootstock-specific diameters (Riparia 60-100 micrometers, SO4 80-120 micrometers, Berlandieri hybrids 100-180 micrometers reflecting genetic variation), exhibiting perforation plates (end walls with large openings enabling water flow between stacked vessel elements), displaying pit connections (lateral pores connecting vessels to adjacent vessels and parenchyma allowing radial water movement), and demonstrating tyloses formation (balloon-like parenchyma outgrowths blocking vessels during heartwood conversion or pathogen defense). Rootstock xylem water transport function includes= conducting 50-100+ liters daily during growing season (xylem vessels moving water from roots through transpiration-driven negative pressure), transporting dissolved minerals (nitrogen, phosphorus, potassium, calcium, micronutrients delivered to leaves and fruit via xylem sap), showing cohesion-tension mechanism (water columns under tension maintained by molecular cohesion enabling transport against gravity), exhibiting vulnerability to cavitation (air embolism formation breaking water columns under stress disrupting transport), and demonstrating refilling capacity (embolized vessels potentially recovering function through root pressure or localized repair). Rootstock xylem mechanical function includes= providing structural rigidity (lignified xylem cells creating stiff framework supporting upright trunk position), bearing compressive loads (xylem cells resisting crushing forces from vine weight and gravitational stress), resisting bending forces (xylem contributing to trunk flexural strength preventing excessive deflection under wind or crop load), and maintaining dimensional stability (lignified xylem structure resistant to deformation maintaining trunk form). Rootstock xylem formation and development includes= produced by vascular cambium (lateral meristem dividing to create xylem cells inward), showing cell differentiation (cambium-derived cells elongating, depositing secondary walls, and lignifying to form mature xylem), undergoing programmed cell death (vessel elements and fibers dying at maturity leaving hollow functional structures), accumulating annually (each growing season adding new xylem layer creating annual ring increasing trunk diameter), and showing age-related changes (young trunk all functional sapwood xylem transitioning to sapwood-heartwood zonation as trunk matures). Rootstock xylem cultivar characteristics includes= exhibiting genetic vessel size (vigorous rootstocks producing larger vessels supporting high transport capacity), showing characteristic wood density (Rupestris dense xylem, Riparia lighter xylem, hybrids intermediate), displaying cultivar-specific anatomy (vessel density, fiber wall thickness, and ray structure varying with rootstock genetics), and demonstrating growth rate patterns (annual xylem production reflecting rootstock vigor classification).
Rootstock_Xylem emphasizes water-conducting secondary xylem tissue and is distinguished from Rootstock_Phloem (food-conducting tissue) by focus on water transport vessels, mechanical support, and centripetal development. Critical for understanding trunk hydraulic function, structural support, sapwood-heartwood differentiation, annual ring formation, and the essential vascular tissue conducting water and minerals while providing mechanical framework.
Rootstock water-conducting tissue
Rootstock wood xylem
Secondary xylem tissue
Sekundäres Xylem
Unterlagen-Xylem
Wurzelstock-Wasserleitgewebe
Rootstock Xylem
Rose aroma evaluation contrasts young Gewürztraminer (rose petal 8.2 on 10-point scale from cis-rose oxide at 95 μg/L, fresh and lifted character) with 15-year Barolo (rose 7.5, dried rose petal character arising from glycoside hydrolysis releasing geraniol and citronellol, with tarry undertones), demonstrating the two distinct pathways to rose character in wine and how rose aroma nuance provides both varietal identification in white wines and aging assessment in red wines.
Rose aroma quality is a floral aroma quality that represents olfactory perception evoking rose flower-associated scents including fresh rose petal, dried rose, rose water, and Turkish delight notes, characterizing the distinctive sweet-floral aromatic impression that serves as a signature varietal marker for Gewürztraminer and contributes complexity to aged red wines particularly Nebbiolo-based examples. Within the BFO framework, this quality specifically addresses the rose subcategory of floral aroma, distinguished by its unique chemical basis and strong varietal associations, recognizing that rose character arises from specific volatile compounds (rose oxides, geraniol, citronellol) that provide direct compositional correspondence to actual rose flower volatiles. The ontological significance lies in rose aroma serving as perhaps the most distinctive and recognizable floral note in wine, where intense rose character immediately suggests Gewürztraminer in white wines or evolved Barolo in red wines, creating strong variety and age associations that enable identification from this single aromatic attribute. The compositional basis for rose aroma includes cis-rose oxide and trans-rose oxide (providing the distinctive rose-lychee character of Gewürztraminer with detection threshold of 0.2 μg/L), geraniol (general rose character), citronellol (rose with citrus undertones), and 2-phenylethanol (rose water, heavier rose notes), with the specific compound balance determining whether fresh rose petal or dried rose character predominates. The developmental aspects of rose aroma differ between grape-derived (terpene-based in Gewürztraminer) and aging-derived (phenolic transformation in Nebbiolo) origins, with rose character in aged red wines arising from glycosidic precursor hydrolysis and phenolic compound oxidation rather than the direct terpene presence of aromatic white varieties. Understanding rose aroma quality enables varietal authentication for Gewürztraminer, aging assessment in Nebbiolo-based wines where rose development indicates maturity, and quality evaluation where rose character purity and intensity reflect varietal typicity and wine evolution.
Rose_Aroma_Quality addresses the rose flower subcategory of floral aroma, encompassing fresh rose petal, dried rose, rose water, and Turkish delight notes. This quality shows two distinct origins in wine - terpene-derived in aromatic varieties and aging-derived in Nebbiolo - making it valuable for both varietal identification and aging assessment. The rose oxide compounds responsible for Gewürztraminer's distinctive character have among the lowest detection thresholds of wine volatiles.
Rosenaroma
Rosenblätternote
Rosenblütencharakter
Rosenduft
rosa scent
rose flower notes
rose petal aroma
rose water character
Guth H (1997) Identification of character impact odorants of different white wine varieties. Journal of Agricultural and Food Chemistry 45(8): 3022-3026 Marengo E et al. (2001) Characterization of the olfactory space of Nebbiolo-based wines. Journal of the Science of Food and Agriculture 81(11): 1057-1062
Rose Aroma Quality
Grenache bears a Rose_Wine_Scion_Role as the dominant variety for Provence rosé, contributing pale salmon color (anthocyanins 50-100 mg/L after brief maceration), red fruit aromatics, and round texture to wines from Côtes de Provence, Bandol, and Tavel commanding the rosé category globally. Cinsault serves in a Rose_Wine_Scion_Role throughout southern France and South Africa, providing delicate color, floral aromatics, and fresh acidity in Provence blends and single-varietal Cape rosés. Pinot Noir bears a Rose_Wine_Scion_Role in Champagne (for rosé Champagne), Sancerre (Sancerre Rosé), and New World regions, producing pale, elegant rosés with red berry character. Mourvèdre (Monastrell) contributes structure and depth to premium rosés from Bandol and Murcia.
A wine production scion role that inheres in a grafted scion cultivar by virtue of its designation for rosé wine production, where limited skin contact with pigmented grape skins extracts subtle anthocyanin coloration (typically 50-200 mg/L versus 200-500 mg/L for reds) and minimal tannins, yielding wines ranging from pale salmon through onion-skin to deep cherry-pink. Rosé wine scions are typically red-skinned varieties processed with brief maceration (2-24 hours) before pressing, direct pressing without maceration (vin gris style), or the saignée method where juice is bled from red wine fermentations. Successful rosé production requires cultivars that achieve physiological ripeness with balanced sugar, acidity (typically 6-8 g/L), and aromatic development at moderate alcohol levels (11-13.5% v/v), and whose anthocyanin profiles yield attractive pink hues without excessive blue or orange tones. The rosé role is distinguished from red wine production primarily by vinification method rather than cultivar genetics, though certain varieties (Grenache, Cinsault, Mourvèdre, Pinot Noir) are particularly associated with quality rosé production.
Rose_Wine_Scion_Role captures a utilization category defined by vinification method applied to pigmented varieties rather than a genetically distinct cultivar group. The same variety frequently serves in both Red_Wine and Rose_Wine roles depending on winemaker intent—Grenache, Syrah, Mourvèdre, and Cinsault all produce both red and rosé wines. Provence rosé style (very pale, dry, fresh) has become the global benchmark, driving demand for varieties and viticultural practices yielding delicate color and crisp acidity. Rosé production has experienced dramatic growth (doubling global consumption 2002-2018), elevating previously minor varieties and creating quality-focused production in regions historically focused on reds.
Rosé-Traube
Roséweinrebe
Roséweinsorte
pink wine variety
rosé variety
rosé wine grape
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Jackson RS (2014) Wine Science 4th Ed, Academic Press pp.: 420-430 Conseil Interprofessionnel des Vins de Provence (CIVP) (2019) Economic Observatory of Provence Wines Waterhouse AL et al. (2016) Understanding Wine Chemistry, Wiley
Rose Wine Scion Role
Rotundone concentration varies dramatically across cultivars, with Syrah showing uniquely high accumulation (50-500 ng/L in berries, 10-200 ng/kg in wine) compared to most other V. vinifera cultivars (<10 ng/L). Additional cultivars with measurable rotundone include Durif/Petite Sirah (20-150 ng/L), Mourvèdre (10-80 ng/L), Vespolina (50-200 ng/L), and Grüner Veltliner (10-50 ng/L). The sensory threshold of 8-16 ng/L (with 20-25% of the population anosmic to rotundone) means the compound is supra-threshold in most Syrah wines but subliminal in low-accumulating cultivars. Rotundone biosynthesis occurs via the cytosolic mevalonate (MVA) pathway through sesquiterpene synthase activity, with VvTPS24 (α-guaiene synthase) producing the precursor that is subsequently oxidized to rotundone. Accumulation occurs primarily in berry skins during late ripening (4-6 weeks post-veraison), with cool temperatures (15-20°C) and moderate water deficit promoting higher concentration.
An aromatic compound disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate the sesquiterpene rotundone ((3S,5R,8S)-3,4,5,6,7,8-hexahydro-3,8-dimethyl-5-(prop-1-en-2-yl)azulen-1(2H)-one) in grape berry skins, with genotype-specific concentrations determining the intensity of black pepper and spice aroma that characterizes Syrah and certain other red cultivars.
Rotundone represents a breakthrough discovery (2008) explaining the distinctive black pepper/spice character of cool-climate Syrah. The compound shows remarkable cultivar specificity, with Syrah accumulating 10-100× higher concentrations than most other varieties. The high proportion of anosmia (inability to detect rotundone) in the population means pepper character is perceived differently by different tasters. Climate significantly modulates rotundone, with cool sites and vintages producing more peppery wines, explaining the traditional association of pepper with northern Rhône Syrah versus the fruitier character of warm-climate Shiraz.
Gewürznote
Rotundon-Akkumulation
black pepper aroma trait
pepper sesquiterpene
spice compound disposition
Wood C et al. (2008) From wine to pepper: rotundone, an obscure sesquiterpene, is a potent spicy aroma compound. Journal of Agricultural and Food Chemistry 56:3738-3744 Siebert TE et al. (2008) Development of a stable isotope dilution analysis for the quantification of the key odorant rotundone in grapes, juice, and wine. Journal of Agricultural and Food Chemistry 56:3745-3748
Rotundone Accumulation Disposition
Rough surface quality development on Concord berry skins results from the characteristic 'slip-skin' anatomy where the epidermis separates readily from underlying flesh, creating surface undulations visible as the skin loosens during ripening this rough, easily detached skin texture is valued for juice production where skins are removed, but contrasts with the adherent smooth-skinned texture of Vitis vinifera table grapes, exemplifying how rough surface quality can be either a defect or a desirable character depending on the intended use.
Rough_Surface_Quality represents the surface texture condition where the external boundary of a grapevine organ displays tactile irregularities that can be felt or seen, presenting an uneven surface with detectable bumps, ridges, or depressions. Rough surfaces result from irregular epidermal cell arrangement, variable cuticle thickness, presence of surface structures, or post-developmental modifications such as cracking or weathering. In grape berries, rough skin texture may indicate cuticular microcracking from rapid expansion during cell enlargement phases, a condition that increases water loss and provides entry points for fungal pathogens. Certain cultivars and species, particularly Muscadinia rotundifolia and some Vitis hybrids, naturally display rougher berry surfaces with prominent lenticels or reticulate cuticular patterning. In leaves, rough texture may result from bullate (blistered) surface morphology where intercostal regions bulge above the vein network, a condition that affects light capture and boundary layer properties. Cane surfaces develop rough texture during maturation as periderm forms and lenticels become prominent. Rough texture on fruit can reduce visual appeal for table grapes but may increase skin surface area and enhance extraction of skin-derived compounds during winemaking. Assessment of rough surface quality employs tactile evaluation detecting friction and resistance to smooth movement, visual inspection revealing surface irregularities, and profilometric measurement quantifying roughness parameters.
Rough_Surface_Quality represents the opposite pole from Smooth_Surface_Quality on the texture continuum. Roughness may be inherent to the cultivar or induced by environmental conditions including rapid growth causing cuticular stress, physical abrasion from wind or adjacent organs, or pathogen damage. The threshold distinguishing rough from smooth surfaces involves judgment and may be specified through roughness parameter values (e.g., Ra above 2 μm). Rough surface texture should be distinguished from pubescent texture where trichomes cause the tactile sensation rough surfaces in the strict sense lack trichomes but present uneven topography.
coarse surface quality
grobe Oberflächenqualität
irregular texture
raue Oberfläche
textured surface
unebene Textur
uneven surface
unregelmäßige Oberflächenbeschaffenheit
Considine JA and Knox RB (1979) Development and Histochemistry of the Cells, Cell Walls, and Cuticle of the Dermal System of Fruit of the Grape, Vitis vinifera L. Protoplasma 99(4): 347-365 Becker T and Knoche M (2012) Deposition, Strain, and Microcracking of the Cuticle in Developing 'Riesling' Grape Berries. Vitis 51(1): 1-6
Rough Surface Quality
Roughness quality assessment in young Madiran Tannat reveals pronounced coarseness (roughness rating 7/10) from variety's characteristically aggressive tannins, creating obvious textural friction that demands 5-10 years cellaring for smoothness development, contrasting with young Pinot Noir from same vintage showing minimal roughness (2/10) appropriate for current consumption, demonstrating how roughness interpretation requires variety and style context to distinguish appropriate youthful grip from problematic harshness.
Roughness quality is a mouthfeel quality that represents the presence of tactile friction, catching, or textural irregularity during wine consumption, characterizing the coarse, gripping, or abrasive oral sensation arising from aggressive tannins, excessive acidity, or other components creating surface-irritating texture. Within the BFO framework, this quality specifically addresses the friction-present dimension of mouthfeel as the opposite of smoothness, distinguishing rough texture from smooth while recognizing that roughness may indicate either youth (with aging potential for smoothness development) or quality deficiency (from inherently coarse tannins or poor balance). The ontological significance lies in roughness providing diagnostic information about wine developmental stage and quality level, where roughness in young wines with quality potential suggests cellaring need while roughness in wines expected to be smooth indicates either insufficient aging or fundamental quality limitation affecting consumption suitability. The perceptual basis for roughness involves mechanoreceptor stimulation indicating surface friction and irregularity, with rough wines creating the sensation of catching, dragging, or irritation as they move across oral surfaces, particularly noticeable on the tongue surface and inner cheeks where tactile sensitivity is highest. The compositional determinants of roughness include tannin quantity and quality (high concentrations of reactive, unpolymerized tannins creating grip and roughness), excessive acidity (contributing sharpness and biting character), inadequate glycerol or polysaccharides (failing to provide lubricating smoothness), and poor component integration (creating textural discontinuity where harsh elements stand out). The quality implications of roughness depend on context, with moderate roughness acceptable in young age-worthy reds where smoothness development is expected, while roughness in wines intended for current consumption or in aged wines that should have smoothed indicates quality concern. Understanding roughness quality enables developmental stage assessment, aging need diagnosis, and quality evaluation distinguishing youthful potential from inherent coarseness.
Roughness_Quality addresses the friction-present dimension of mouthfeel as the textural opposite of smoothness. This quality may indicate either youthful potential (when aging can resolve roughness) or quality deficiency (when coarseness is inherent). The class supports developmental assessment and aging recommendations while distinguishing appropriate youthful structure from problematic harshness.
Rauheit
Texturunebenheit
coarse texture
grippy texture
grobe Textur
harsh mouthfeel
raues Mundgefühl
textural coarseness
Roughness Quality
A Cabernet Sauvignon vine in north-south oriented rows in Napa Valley bears a Row_Orientation_Role receiving morning sun on its east-facing canopy side and afternoon sun on its west-facing side, the balanced exposure considered optimal for even fruit ripening in the Mediterranean climate. A Riesling vine in east-west rows on steep Mosel slopes serves in a Row_Orientation_Role aligned perpendicular to the slope direction, maximizing exposure to the south-facing sun essential for ripening in Germany's northern climate. A Merlot vine in north-south rows oriented to prevailing afternoon winds bears a Row_Orientation_Role enabling airflow through the canopy that reduces humidity and disease pressure in a maritime-influenced region.
A vineyard role that inheres in a grapevine by virtue of the compass direction of the vineyard row in which it is planted, with row orientation determining the pattern of sunlight interception, shade progression, and wind exposure experienced by the vine throughout the growing season. Row orientation affects canopy microclimate through the timing and intensity of direct sunlight on each side of the canopy—north-south rows receive morning sun on the east side and afternoon sun on the west side, while east-west rows receive sun primarily on the south-facing side (in the Northern Hemisphere) with the north side remaining more shaded. The orientation choice influences fruit composition through exposure effects: early morning sun enables rapid drying of overnight moisture (reducing disease pressure), while afternoon sun drives fruit temperature during the warmest hours (potentially causing heat stress in hot climates). Row orientation interacts with prevailing wind direction, slope aspect, and training system to create the specific microclimate conditions experienced by each vine. Orientation decisions at vineyard establishment are permanent for the vineyard's lifespan, making the choice particularly consequential for long-term performance.
Row_Orientation_Role is subclassed by specific orientation categories—North_South, East_West, and Slope_Aligned—recognizing the distinct microclimate implications of each approach. Row orientation is one of the fundamental vineyard design decisions, made at establishment and difficult or impossible to change thereafter. The "optimal" orientation varies with latitude, climate, and production objectives: north-south is generally preferred in warm climates for balanced exposure, while east-west may be preferred at high latitudes to maximize southern sun exposure or on slopes where contour alignment prevents erosion. Precision viticulture research has documented significant within-row variation in fruit composition related to canopy side and orientation, with the patterns predictable based on sun path geometry. Climate change is prompting reconsideration of traditional orientation choices, as heat accumulation patterns and optimal exposure levels shift with warming temperatures.
Reihenrichtung
Weinbergsausrichtung
Zeilen-Orientierung
Zeilenausrichtung
row alignment
row direction
row orientation
vineyard orientation
Smart RE (1973) Sunlight interception by vineyards, American Journal of Enology and Viticulture 24: 141-147 Hunter JJ et al. (2016) Row orientation effects, South African Journal of Enology and Viticulture 37: 107-119 Intrieri C et al. (1998) Row orientation effects on grape composition, Vitis 37: 83-88
Row Orientation Role
The Rpv10 presence was first identified and mapped by Schwander et al. (2012) in a segregating population derived from the V. amurensis accession Gf.Ga-52-42 crossed with V. vinifera, localizing the resistance to a 3.1 cM interval on chromosome 9 flanked by markers GF09-46 and GF09-68 — Rpv10-carrying individuals showed 60-85% reduction in P. viticola sporulation intensity compared to susceptible siblings in controlled inoculation assays, with the resistance mechanism involving delayed pathogen development (reduced hyphal colonization rate and smaller sporangiophore density) rather than the rapid hypersensitive cell death of Rpv1 or the pronounced stilbene accumulation of Rpv3. Rpv10 has been incorporated into second-generation PIWI breeding lines at the Julius Kühn-Institut (JKI) Geilweilerhof in combination with Rpv3 to create pyramided Rpv3+Rpv10 genotypes showing resistance levels exceeding either locus alone.
An Rpv locus information record documenting the presence or absence of the Rpv10 resistance locus on chromosome 9, derived from Vitis amurensis (Amur grape, native to East Asia), that confers partial to strong resistance to Plasmopara viticola through mechanisms independent of the North American species-derived Rpv3 pathway, providing a genetically and geographically distinct resistance source valuable for pyramiding strategies to achieve durable multi-locus downy mildew resistance in new cultivar development.
Rpv10_Presence documents a resistance source of strategic importance for durable downy mildew resistance breeding because it derives from Vitis amurensis — a species that evolved in East Asia independently from the North American Vitis species that provided Rpv3, and therefore carries resistance genes with different evolutionary origins, recognition specificities, and defense mechanisms. This geographic and phylogenetic independence makes Rpv10 complementary to Rpv3 in pyramiding strategies, as the probability that P. viticola can simultaneously overcome two mechanistically unrelated resistance pathways is substantially lower than for either pathway individually: theoretical population genetics models predict that pyramided two-locus resistance has durability 10-100 fold greater than single-locus resistance, depending on pathogen mutation rate and population size. V. amurensis is additionally valuable as a resistance donor because it is the most cold-hardy Vitis species (tolerating winter temperatures below -40°C), enabling breeding of disease-resistant cultivars adapted to continental climates, and because its relatively close taxonomic relationship to V. vinifera (both within the Euvitis subgenus) facilitates hybridization and quality recovery in backcross programs compared to the more distant Muscadinia. Rpv10 is being widely incorporated into current European breeding programs alongside Rpv12 as the preferred V. amurensis-derived resistance loci for pyramiding with Rpv3.
Amur grape downy mildew resistance chromosome 9
Amurrebe-Peronospora-Resistenz-Chr9
Rpv10 locus genotype
Rpv10 resistance status
Rpv10-Präsenz
Rpv10-Resistenzstatus
Schwander F et al. (2012) Rpv10: a new locus from the Asian Vitis amurensis rotundifolia conferring resistance to grapevine downy mildew, Theoretical and Applied Genetics 124: 163-176 Töpfer R et al. (2011) Molecular breeding, in Adam-Blondon AF et al. (eds) Genetics, Genomics, and Breeding of Grapes, Science Publishers pp: 160-185 Dry IB et al. (2010) Grapevine disease resistance genes, in Reynolds AG (ed) Viticulture and Wine Quality, Woodhead Publishing pp: 193-228
Rpv10 Presence
The breeding line 'Gf.Ga-52-42' at JKI Siebeldingen carries Rpv10_Resistance_Disposition derived from V. amurensis through the resistant parent 'Gf.Ga-47-42'. Phenotypic evaluation using detached leaf disc assays with P. viticola sporangial suspension (30,000 sporangia/mL) demonstrates that Rpv10 confers strong resistance characterized by small necrotic spots (mean diameter 0.8mm) visible 3-4 days post-inoculation with no subsequent sporangiophore development even after 10 days under optimal conditions for the pathogen. In contrast, susceptible 'Müller-Thurgau' discs develop extensive sporulation covering >80% of the leaf disc surface. Importantly, P. viticola isolates capable of partially overcoming Rpv3-1 resistance retain full avirulence on Rpv10 genotypes, demonstrating the distinct recognition specificities of these loci and validating the strategy of combining Rpv10 with Rpv3 for enhanced resistance durability.
An Rpv10_Resistance_Disposition is an Rpv_Mediated_Resistance_Disposition that inheres in a Vitis plant carrying the Rpv10 locus derived from Vitis amurensis and is realized through effector-triggered immunity upon recognition of corresponding Plasmopara viticola avirulence effectors. The Rpv10 locus maps to chromosome 9 and was identified in the Asian wild species V. amurensis, which evolved in regions of northeastern China, Korea, and the Russian Far East where endemic Plasmopara populations exerted selection pressure for resistance. V. amurensis additionally possesses exceptional cold hardiness, making it a valuable genetic resource for breeding cultivars adapted to continental climates. The Rpv10 resistance phenotype is characterized by rapid hypersensitive response with localized cell death arresting pathogen development at early post-penetration stages, resulting in small necrotic flecks (typically <1mm diameter) without macroscopically visible sporulation. Rpv10 provides a recognition specificity distinct from Rpv3, making it an excellent partner for pyramiding strategies. The resistance breeding line 'Solaris' and its derivatives carry Rpv10 combined with other resistance factors, contributing to the development of cold-hardy, disease-resistant cultivars suitable for northern European and continental climates.
Rpv10_Resistance_Disposition is_a Rpv_Mediated_Resistance_Disposition. The Rpv10 locus originates from Vitis amurensis, the most cold-hardy Vitis species, providing additional breeding value beyond disease resistance. Rpv10 has a distinct recognition specificity from Rpv3 and is frequently combined with Rpv3 haplotypes in modern breeding programs. The locus on chromosome 9 is distinct from the Run1/Rpv1 cluster on chromosome 12 and the Rpv3 cluster on chromosome 18.
Amurensis-Resistenz
Asian grape DM resistance
Rpv10 downy mildew resistance
Rpv10-Peronospora-Resistenz
Rpv10-Resistenz-Disposition
V. amurensis-derived DM resistance
Schwander F et al. (2012) Rpv10: a new locus from the Asian Vitis gene pool for pyramiding downy mildew resistance loci in grapevine, Theoretical and Applied Genetics 124:163-176 Venuti S et al. (2013) Historical introgression of the downy mildew resistance gene Rpv12 from the Asian species Vitis amurensis into grapevine varieties, PLoS ONE 8:e61228
Rpv10 Resistance Disposition
The Rpv12 presence was mapped by Venuti et al. (2013) in a V. amurensis-derived population to a 1.2 cM interval on chromosome 14 containing a cluster of TIR-NBS-LRR resistance gene analogs — Rpv12-carrying individuals showed 80-95% reduction in sporulation compared to susceptible controls, one of the strongest quantitative effects documented for any single Rpv locus, with the resistance mechanism involving rapid callose deposition in stomatal guard cells within 12-24 hours of zoospore encystment that physically blocks pathogen penetration into the substomatal cavity. The PIWI cultivar Sauvignac (released by INRAE 2020) carries both Rpv3 and Rpv12 in a pyramided combination, demonstrating the practical implementation of Asian-American resistance source combination — in multi-site French field trials, Sauvignac required only 1-2 fungicide applications per season compared to 10-14 for the susceptible V. vinifera control Sauvignon Blanc, achieving the 80%+ spray reduction target of modern resistance breeding programs.
An Rpv locus information record documenting the presence or absence of the Rpv12 resistance locus on chromosome 14, derived from Vitis amurensis, that confers strong resistance to Plasmopara viticola through rapid post-penetration defense responses, representing alongside Rpv10 the principal Asian-origin downy mildew resistance source used in pyramiding strategies with the American-origin Rpv3 to develop PIWI cultivars with durable multi-locus resistance.
Rpv12_Presence documents the second major V. amurensis-derived resistance locus that, together with Rpv10, provides the Asian resistance complement to the American-origin Rpv3 in current pyramiding strategies. Rpv12 is distinguished by its particularly strong resistance effect (among the highest of all characterized Rpv loci), its distinct chromosomal location (chromosome 14, unlinked to Rpv3 on chromosome 18 and Rpv10 on chromosome 9, enabling independent segregation in breeding populations), and its unique resistance mechanism centered on rapid callose-mediated stomatal occlusion that represents a pre-penetration defense distinct from the post-penetration stilbene and HR responses of Rpv3 and Rpv1 respectively: this mechanistic distinctness is critically important for pyramiding durability because a pathogen mutant capable of evading the stilbene-based defense of Rpv3 would gain no advantage against the callose-based stomatal defense of Rpv12, requiring independent adaptive mutations to overcome each resistance component. Current breeding strategies at leading European programs (JKI Geilweilerhof, INRAE Colmar, Agroscope Changins, Fondazione Edmund Mach) aim to combine Rpv3 + Rpv12 (or Rpv3 + Rpv10 + Rpv12 in three-locus pyramids) to achieve resistance durability estimated at 20-50+ years under moderate selection pressure — a durability horizon that would transform the economics and environmental impact of European viticulture by substantially eliminating the 6-15 annual fungicide applications currently required for downy mildew control on susceptible V. vinifera cultivars.
Amur grape downy mildew resistance chromosome 14
Amurrebe-Peronospora-Resistenz-Chr14
Rpv12 locus genotype
Rpv12 resistance status
Rpv12-Präsenz
Rpv12-Resistenzstatus
Venuti S et al. (2013) Historical introgression of the downy mildew resistance gene Rpv12 from the Asian species Vitis amurensis into grapevine varieties, PLoS ONE 8: e61228 INRAE (2020) Sauvignac varietal registration documentation Töpfer R et al. (2011) Molecular breeding, in Adam-Blondon AF et al. (eds) Genetics, Genomics, and Breeding of Grapes, Science Publishers pp: 160-185 Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance, Molecular Breeding 36: 65
Rpv12 Presence
The Hungarian cultivar 'Kunbarat', released in 1976 from a cross of 'Irsai Olivér' × V. amurensis, carries Rpv12_Resistance_Disposition that has provided stable field resistance for over four decades in Hungarian vineyards. Controlled inoculation experiments at the University of Pécs demonstrate that 'Kunbarat' exhibits the strongest resistance phenotype among tested cultivars, with hypersensitive response initiated within 18-24 hours post-inoculation and complete arrest of pathogen development by 48 hours. Sporulation is never observed even under extreme inoculum pressure (100,000 sporangia/mL) or extended incubation periods (14 days). Molecular marker analysis confirmed the presence of Rpv12 on chromosome 14, and genetic studies showed that this resistance segregates independently from Rpv10 on chromosome 9, despite both originating from V. amurensis, enabling their combination in breeding to achieve multiple-gene resistance.
An Rpv12_Resistance_Disposition is an Rpv_Mediated_Resistance_Disposition that inheres in a Vitis plant carrying the Rpv12 locus derived from Vitis amurensis and is realized through effector-triggered immunity upon recognition of corresponding Plasmopara viticola avirulence effectors. The Rpv12 locus maps to chromosome 14 and represents a second major resistance gene from V. amurensis that is genetically independent from Rpv10. Rpv12 confers exceptionally strong resistance with a phenotype characterized by rapid and intense hypersensitive response resulting in complete arrest of P. viticola development at very early infection stages. Historical pedigree analysis has revealed that Rpv12 was inadvertently introgressed into European grapevine breeding material through early 20th century crosses involving V. amurensis, and is present in several established PIWI cultivars including 'Kunbarat' and 'Kunleány' from the Hungarian breeding program. The independent genetic position of Rpv12 relative to other major Rpv loci makes it highly valuable for resistance gene pyramiding, and its combination with Rpv3 and/or Rpv10 is expected to provide robust, durable field resistance against diverse P. viticola populations including isolates with evolved virulence against individual resistance genes.
Rpv12_Resistance_Disposition is_a Rpv_Mediated_Resistance_Disposition. Although both Rpv10 and Rpv12 originate from Vitis amurensis, they map to different chromosomes (9 and 14 respectively) and can be independently selected and combined. Rpv12 confers particularly strong resistance with rapid hypersensitive response. The presence of Rpv12 in established Hungarian cultivars provides validated germplasm for breeding programs seeking to incorporate this resistance source.
Kunbarat-type resistance
Rpv12 downy mildew resistance
Rpv12-Amurensis-Resistenz
Rpv12-Peronospora-Resistenz
Rpv12-Resistenz-Disposition
chromosome 14 DM resistance
Venuti S et al. (2013) Historical introgression of the downy mildew resistance gene Rpv12 from the Asian species Vitis amurensis into grapevine varieties, PLoS ONE 8:e61228 Kozma P et al. (2009) Resistance to downy mildew derived from interspecific hybrid Vitis rupestris × Vitis amurensis, Acta Horticulturae 827:507-512
Rpv12 Resistance Disposition
The Rpv1 presence in breeding line BC4 2-80 at INRAE Colmar was confirmed by marker-assisted selection using the flanking SSR markers VMC72 and VMC7f2 on chromosome 12 — this line carries the Rpv1 locus introgressed from M. rotundifolia through the backcross lineage initiated by Bouquet (1986) using the resistant accession M. rotundifolia G52, with successive backcrosses to V. vinifera (BC1 through BC4) progressively recovering V. vinifera wine quality characters while retaining the Rpv1 resistance segment. Controlled inoculation of Rpv1-carrying BC4 individuals with P. viticola isolates from French vineyards showed complete suppression of sporulation (disease severity <2% versus 60-80% in susceptible V. vinifera controls) through rapid HR visible as necrotic spots within 48 hours of inoculation, confirming the strong, near-qualitative resistance conferred by this locus.
An Rpv locus information record documenting the presence or absence of the Rpv1 resistance locus on chromosome 12, derived from Muscadinia rotundifolia (muscadine grape), that confers strong qualitative resistance to Plasmopara viticola through a rapid hypersensitive response (HR) producing localized cell death at infection sites, representing the first molecularly characterized downy mildew resistance locus in grapevine and one of the strongest single-gene resistance sources available for breeding.
Rpv1_Presence documents the detection status of the first identified and one of the strongest downy mildew resistance loci in grapevine, derived from the Muscadinia subgenus which diverged from Euvitis approximately 18 million years ago. Rpv1 was mapped by Merdinoglu et al. (2003) to chromosome 12 in a segregating population derived from M. rotundifolia, and the underlying resistance mechanism involves rapid programmed cell death (hypersensitive response) at the site of P. viticola penetration, effectively isolating the biotrophic pathogen from living host tissue within 24-48 hours of infection: this HR-based resistance is among the strongest available, typically reducing disease severity by 95-99% in controlled inoculation assays. However, the introgression of Rpv1 from M. rotundifolia into V. vinifera backgrounds is technically challenging due to the taxonomic distance between Muscadinia (2n = 40) and Euvitis (2n = 38), requiring embryo rescue for initial hybrid production and extensive backcrossing (4-6 generations) to recover acceptable wine quality — a process spanning 25-30 years that has limited Rpv1 deployment compared to the more easily accessed Rpv3 from Euvitis species. The strong selection pressure imposed by Rpv1 on pathogen populations raises durability concerns, as single-gene qualitative resistance is theoretically more vulnerable to pathogen adaptation than polygenic quantitative resistance.
Muscadinia-Peronospora-Resistenz
Rpv1 locus genotype
Rpv1 resistance status
Rpv1-Präsenz
Rpv1-Resistenzstatus
muscadine downy mildew resistance
Merdinoglu D et al. (2003) Genetic analysis of downy mildew resistance derived from Muscadinia rotundifolia, Acta Horticulturae 603: 451-456 Bouquet A (1986) Introduction dans l'espèce Vitis vinifera L. d'un caractère de résistance à l'oïdium issu de l'espèce Muscadinia rotundifolia, Vignevini 12: 141-146 Dry IB et al. (2010) Grapevine disease resistance genes, in Reynolds AG (ed) Viticulture and Wine Quality, Woodhead Publishing pp: 193-228
Rpv1 Presence
The breeding line Gf.2000-305-122 at Julius Kühn-Institut carries Rpv1_Resistance_Disposition introgressed from M. rotundifolia through multiple generations of pseudo-backcrossing initiated by Alain Bouquet at INRA Montpellier in the 1970s. When challenged with a diverse collection of 15 P. viticola isolates from European wine regions, this line exhibits complete resistance to all isolates, with no macroscopically visible sporulation on detached leaf discs maintained under optimal conditions for pathogen development (20°C, >95% RH, 16h photoperiod). Microscopic examination reveals attempted penetration through stomata followed by rapid cell death confined to 1-3 epidermal and mesophyll cells, effectively halting pathogen development. This exceptionally broad-spectrum resistance reflects the evolutionary distance between M. rotundifolia and European P. viticola populations, which lack the corresponding virulence effectors capable of suppressing Rpv1-mediated immunity.
An Rpv1_Resistance_Disposition is an Rpv_Mediated_Resistance_Disposition that inheres in a Vitis plant carrying the Rpv1 locus derived from Muscadinia rotundifolia and is realized through effector-triggered immunity upon recognition of corresponding Plasmopara viticola avirulence effectors. The Rpv1 locus maps to chromosome 12 and is tightly linked to the powdery mildew resistance locus Run1, forming a resistance gene cluster that has been extensively utilized in breeding programs. Rpv1 originates from M. rotundifolia, a species native to the southeastern United States that co-evolved with endemic Plasmopara populations and possesses exceptionally strong downy mildew resistance. The resistance mechanism involves rapid hypersensitive response with complete arrest of pathogen development at the penetration stage, resulting in tiny necrotic flecks without any sporangiophore formation even under high inoculum pressure. Introgression of Rpv1 into V. vinifera backgrounds has been challenging due to the limited crossability between Muscadinia and Euvitis subgenera and the extensive linkage drag associated with the large introgressed segment, but pseudo-backcross breeding strategies have successfully incorporated this locus into wine grape breeding lines while progressively restoring V. vinifera genetic background and wine quality characteristics.
Rpv1_Resistance_Disposition is_a Rpv_Mediated_Resistance_Disposition. The Rpv1 locus is physically linked to Run1 (powdery mildew resistance) on chromosome 12, and both resistances are typically co-inherited. Rpv1 is distinguished by its origin in Muscadinia rotundifolia and its exceptionally strong, broad-spectrum resistance phenotype. Breeding with Rpv1 requires overcoming Muscadinia × Euvitis crossing barriers and managing extensive linkage drag from the large introgressed chromosomal segment.
Muscadinia-Resistenz
Muscadinia-derived DM resistance
Rpv1 downy mildew resistance
Rpv1-Peronospora-Resistenz
Rpv1-Resistenz-Disposition
Run1/Rpv1 cluster resistance
Bouquet A (1986) Introduction dans l'espèce Vitis vinifera L. d'un caractère de résistance à l'oïdium (Uncinula necator) issu de l'espèce Muscadinia rotundifolia, Vignevini 12:141-146 Merdinoglu D et al. (2003) Genetic analysis of downy mildew resistance derived from Muscadinia rotundifolia, Acta Horticulturae 603:451-456 Feechan A et al. (2013) Genetic dissection of a TIR-NB-LRR locus from the wild North American grapevine species Muscadinia rotundifolia, Molecular Plant-Microbe Interactions 26:684-695
Rpv1 Resistance Disposition
The Rpv3 presence in the PIWI cultivar Regent was characterized as the Rpv3-1 haplotype through haplotype-specific marker analysis (Zyprian et al. 2016) — Rpv3-1 is one of at least three functional haplotypes (Rpv3-1, Rpv3-2, Rpv3-3) at the Rpv3 locus that confer different levels and spectra of resistance, with Rpv3-1 (present in Regent, Bianca, Villard Blanc) showing the widest deployment but also documented partial breakdown against virulent P. viticola isolates collected from Rpv3-1-carrying vineyards in France and Germany since 2010. The resistance mechanism involves rapid accumulation of stilbene phytoalexins (resveratrol, pterostilbene, ε-viniferin, δ-viniferin) to concentrations 5-15 fold higher than in susceptible V. vinifera genotypes within 48 hours of infection, inhibiting pathogen development at the penetration and haustorial stages without the complete cell death response characteristic of Rpv1.
An Rpv locus information record documenting the presence or absence of the Rpv3 resistance locus on chromosome 18, derived from North American Vitis species (principally V. rupestris and V. lincecumii through complex interspecific hybrid pedigrees), that confers partial quantitative resistance to Plasmopara viticola primarily through enhanced stilbene phytoalexin accumulation, representing the most widely deployed downy mildew resistance source in current PIWI (Pilzwiderstandsfähige) cultivar production and the resistance locus for which pathogen adaptation has been most extensively documented.
Rpv3_Presence documents the most important downy mildew resistance locus in current viticulture from both a deployment and a durability perspective. Rpv3 on chromosome 18 was identified through QTL mapping in multiple interspecific populations and is present in the majority of first-generation PIWI cultivars released for commercial production in Europe (Regent, Solaris, Johanniter, Cabernet Blanc, Muscaris, Souvignier Gris, and others): this widespread deployment reflects the relative ease of introgressing Rpv3 from North American Vitis species (which hybridize readily with V. vinifera within the Euvitis subgenus) compared to the Muscadinia-derived Rpv1. The partial breakdown of Rpv3-1 resistance — first reported by Peressotti et al. (2010) in French vineyards and subsequently confirmed across multiple European sites — represents a landmark event in grapevine pathology, demonstrating that P. viticola has sufficient evolutionary potential to adapt to deployed resistance within 5-10 years of widespread monoculture planting of single-resistance-locus cultivars. This breakdown has fundamentally reshaped breeding strategy toward pyramiding Rpv3 with additional Rpv loci (particularly Rpv10 and Rpv12 from V. amurensis, which have different recognition specificities) to create cultivars requiring the pathogen to simultaneously overcome multiple resistance mechanisms.
American species downy mildew resistance
Rpv3 locus genotype
Rpv3 resistance status
Rpv3-Präsenz
Rpv3-Resistenzstatus
amerikanische Peronospora-Resistenz
Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance and ripening of grapevines, Molecular Breeding 36: 65 Peressotti E et al. (2010) Breakdown of resistance to grapevine downy mildew upon limited deployment of a resistant variety, BMC Plant Biology 10: 147 Bellin D et al. (2009) Resistance to Plasmopara viticola in grapevine Bianca is controlled by a major dominant gene causing localised necrosis at the infection site, Theoretical and Applied Genetics 120: 163-176 Di Gaspero G et al. (2012) Selective sweep at the Rpv3 locus during grapevine breeding for downy mildew resistance, Theoretical and Applied Genetics 124: 1097-1111
Rpv3 Presence
The widely-planted PIWI cultivar 'Regent' carries Rpv3-1_Resistance_Disposition inherited from 'Chambourcin' in its pedigree. Under natural infection conditions in the Pfalz wine region of Germany, 'Regent' vines exhibit strong but incomplete Rpv3_Resistance_Disposition realization, with typical leaf infection severities of 5-15% compared to 70-90% in adjacent 'Müller-Thurgau' plots during conducive seasons. Microscopic observation reveals that P. viticola penetration and initial colonization proceed normally in 'Regent' leaves, but hypersensitive cell death is triggered 48-72 hours post-inoculation, limiting lesion expansion and reducing sporulation intensity by 70-85%. However, monitoring by the JKI has documented P. viticola isolates from 'Regent' vineyards with increased aggressiveness on Rpv3-1 genotypes, producing sporulation levels approaching those on susceptible controls, demonstrating the selective pressure exerted by widespread Rpv3-1 deployment and the consequent adaptation of pathogen populations.
An Rpv3_Resistance_Disposition is an Rpv_Mediated_Resistance_Disposition that inheres in a Vitis plant carrying one or more haplotypes of the Rpv3 locus on chromosome 18 and is realized through effector-triggered immunity upon recognition of corresponding Plasmopara viticola avirulence effectors. The Rpv3 locus represents a complex resistance gene cluster with multiple distinct haplotypes (Rpv3-1, Rpv3-2, Rpv3-3) originating from different North American Vitis species and conferring varying levels and spectra of resistance. Rpv3-1, derived from the French-American hybrid 'Seibel 4614' (ancestor of 'Villard blanc' and 'Seyval blanc'), provides moderate to strong resistance that has been widely deployed in European PIWI breeding. Rpv3-2 and Rpv3-3 represent additional haplotypes with partially overlapping but distinct recognition specificities. The Rpv3-mediated resistance phenotype typically manifests as delayed and reduced sporulation rather than complete immunity, with small necrotic lesions that may support limited sporangiophore production under high disease pressure. This partial resistance phenotype has led to concerns about durability, with virulent P. viticola isolates capable of partially overcoming Rpv3-1 resistance already documented in European vineyards with high PIWI cultivar adoption, underscoring the importance of combining Rpv3 with additional resistance loci.
Rpv3_Resistance_Disposition is_a Rpv_Mediated_Resistance_Disposition. Multiple Rpv3 haplotypes exist (Rpv3-1, Rpv3-2, Rpv3-3) with different origins and recognition specificities. Rpv3-1 from 'Seibel 4614' lineage is most widely deployed but shows evidence of durability challenges in European vineyards with high PIWI adoption. Rpv3 resistance is typically partial rather than complete, and combining with other Rpv loci is recommended for durable field resistance.
Rpv3 downy mildew resistance
Rpv3 haplotype resistance
Rpv3-Haplotyp-Resistenz
Rpv3-Peronospora-Resistenz
Rpv3-Resistenz-Disposition
Seibel-derived DM resistance
Welter LJ et al. (2007) Genetic mapping and localization of quantitative trait loci affecting fungal disease resistance and leaf morphology in grapevine, Molecular Breeding 20:359-374 Bellin D et al. (2009) Resistance to Plasmopara viticola in grapevine 'Bianca' is controlled by a major dominant gene causing localised necrosis at the infection site, Theoretical and Applied Genetics 120:163-176 Casagrande K et al. (2011) Defence responses in Rpv3-dependent resistance to grapevine downy mildew, Planta 234:1097-1109
Rpv3 Resistance Disposition
The Rpv locus information compiled in the VIVC resistance locus registry documents over 30 designated Rpv loci identified across diverse Vitis species — Rpv1 on chromosome 12 (from Muscadinia rotundifolia, strong hypersensitive response), Rpv3 on chromosome 18 (from North American Vitis species, the most widely deployed resistance source with multiple haplotypes Rpv3-1, Rpv3-2, Rpv3-3), Rpv10 on chromosome 9 (from V. amurensis, Asian origin providing resistance diversity independent of American species loci), and Rpv12 on chromosome 14 (from V. amurensis, strong resistance with potential durability advantages due to distinct recognition spectrum) — breeding programs at Geilweilerhof (Germany), INRAE Colmar (France), Fondazione Edmund Mach (Italy), and Agroscope (Switzerland) are systematically pyramiding 2-4 Rpv loci per cultivar to achieve durable resistance that individual loci cannot provide against the rapidly evolving P. viticola pathogen population.
A resistance gene information record specific to Rpv (Resistance to Plasmopara viticola) loci — the genomic regions conferring resistance to downy mildew in Vitis — documenting the locus designation (Rpv1 through Rpv30+), chromosomal location, source Vitis species, resistance mechanism (hypersensitive response, callose deposition, stilbene accumulation), resistance spectrum (broad versus race-specific), and diagnostic molecular markers enabling marker-assisted selection for downy mildew resistance in grapevine breeding programs.
Rpv_Locus_Information documents the genetic resources for breeding downy mildew resistance — the most consequential disease resistance target in European viticulture, where P. viticola control accounts for 40-60% of total fungicide use and drives the primary environmental and economic costs of conventional vine protection. The progressive identification of Rpv loci through QTL mapping in interspecific crosses has accelerated since the availability of the reference genome (2007), with the number of designated Rpv loci increasing from 3 in 2005 to over 30 by 2023: these loci derive from multiple Vitis species (V. rupestris, V. riparia, V. cinerea, V. amurensis, V. piasezkii, M. rotundifolia) and map to at least 12 of the 19 chromosomes, providing a rich genetic toolkit for resistance breeding. The critical challenge is resistance durability — the breakdown of Rpv3-1 resistance in European vineyards by virulent P. viticola isolates (Peressotti et al. 2010, Casagrande et al. 2011) demonstrated that single-locus deployment is insufficient against this highly variable obligate pathogen, motivating the current emphasis on pyramiding multiple Rpv loci with different resistance mechanisms and recognition specificities to create a multi-layered defense barrier that is evolutionarily difficult for the pathogen to overcome. Subclasses document the four most important Rpv loci in current breeding programs.
Peronospora-Resistenzlocus-Daten
Plasmopara resistance QTL data
Plasmopara-Resistenz-QTL-Daten
Rpv gene characterization
Rpv-Locus-Information
downy mildew resistance locus data
Töpfer R et al. (2011) Molecular breeding, in Adam-Blondon AF et al. (eds) Genetics, Genomics, and Breeding of Grapes, Science Publishers pp: 160-185 VIVC (2023) Loci for resistance to diseases and pests Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance and ripening of grapevines, Molecular Breeding 36: 65 Peressotti E et al. (2010) Breakdown of resistance to grapevine downy mildew upon limited deployment of a resistant variety, BMC Plant Biology 10: 147
Rpv Locus Information
Marker-assisted selection for Rpv_Mediated_Resistance_Disposition at Hochschule Geisenheim University employs SSR and SNP markers flanking characterized Rpv loci to identify resistant seedlings in segregating populations. A cross between 'Riesling' (susceptible, rpv/rpv) and the resistant breeding line Gf.Ga-47-42 (Rpv3-1/rpv3, Rpv12/rpv12) produces offspring segregating for both loci. DNA extracted from young leaf tissue is genotyped using the Rpv3-1-linked marker GF18-06 and the Rpv12-linked marker UDV-014, identifying approximately 25% of seedlings carrying both resistance alleles. These double-resistant individuals are confirmed through artificial inoculation with P. viticola sporangia, showing characteristic small necrotic lesions (diameter 50 sporangiophores per lesion) visible as white downy growth on the abaxial leaf surface.
An Rpv_Mediated_Resistance_Disposition is a Downy_Mildew_Resistance_Disposition that inheres in a Vitis plant and is realized through effector-triggered immunity (ETI) conferred by one or more Rpv (Resistance to Plasmopara viticola) loci encoding nucleotide-binding leucine-rich repeat (NLR) receptor proteins that recognize specific P. viticola effector molecules. This disposition represents qualitative, major gene-mediated resistance characterized by rapid and robust defense activation upon pathogen recognition, typically manifesting as hypersensitive response (HR) with localized programmed cell death at infection sites within 24-72 hours post-penetration. Over 30 Rpv loci have been genetically mapped across the Vitis genus, originating from diverse North American (V. rupestris, V. riparia, V. labrusca, V. cinerea), Asian (V. amurensis, V. piasezkii), and Muscadinia (M. rotundifolia) species. Each Rpv locus confers resistance to specific P. viticola isolates carrying corresponding avirulence effectors, creating potential for resistance breakdown when virulent pathogen races emerge through effector mutation or loss. Consequently, modern resistance breeding programs emphasize pyramiding multiple Rpv loci with distinct recognition specificities to achieve durable field resistance. Rpv-mediated resistance dispositions are readily introgressed into V. vinifera backgrounds through marker-assisted selection using linked molecular markers, enabling efficient breeding of disease-resistant cultivars retaining high wine quality.
Rpv_Mediated_Resistance_Disposition is_a Downy_Mildew_Resistance_Disposition. This class encompasses all major gene resistances conferred by characterized Rpv loci, distinguished from Quantitative_Downy_Mildew_Resistance_Disposition which involves polygenic inheritance. Child classes represent individual Rpv loci (Rpv1_Resistance_Disposition, Rpv3_Resistance_Disposition, etc.) and combinations thereof (Pyramided_Rpv_Resistance_Disposition). The "Rpv" nomenclature follows Vitis gene naming conventions established by the International Grape Genome Program (IGGP).
ETI-based Plasmopara resistance
Majorgen-Peronospora-Resistenz
Rpv locus resistance
Rpv-Genresistenz
Rpv-vermittelte Resistenz-Disposition
major gene downy mildew resistance
Fischer BM et al. (2004) Quantitative trait locus analysis of fungal disease resistance factors on a molecular map of grapevine, Theoretical and Applied Genetics 108:501-515 Töpfer R et al. (2011) New horizons for grapevine breeding, Fruit, Vegetable and Cereal Science and Biotechnology 5:79-100 Dry IB et al. (2019) Grapevine NLR immunity against downy and powdery mildew pathogens, Acta Horticulturae 1248:129-136
Rpv Mediated Resistance Disposition
Wild V. vinifera ssp. sylvestris male plants in southeastern Europe and Near East commonly exhibit rudimentary pistil flowers with recognizable pistil structures 0.5-0.7 mm long showing distinct basal ovary region (0.3 mm diameter), narrow style region (0.2 mm), and slightly expanded apex suggesting stigma, surrounded by five robust stamens (filaments 3.2-3.8 mm, anthers 0.75-0.85 mm) producing 32,000-40,000 pollen grains per flower with 78-86% viability, achieving 0% fruit set despite pistil presence, and requiring microscopic examination to distinguish from vestigial pistil types though functionally equivalent in complete male sterility. North American wild Vitis species male plants (V. labrusca, V. riparia) characteristically bear rudimentary pistil flowers with pistils 0.5-0.8 mm showing clear organ regions but lacking functional reproductive tissues. Breeding population segregants occasionally produce rudimentary pistil staminate flowers providing educational material for developmental morphology studies but requiring removal from production-oriented breeding programs due to fruit set failure.
A rudimentary pistil flower is a reduced pistil flower in which the pistil shows recognizable though severely abbreviated organ morphology with distinguishable ovary, style, and stigma regions, typically measuring 0.4-0.8 mm in total length and exhibiting clear though drastically reduced pistil structure compared to functional hermaphrodite pistils (1.5-2.5 mm), representing moderate pistil suppression that permits partial organ elaboration before developmental arrest while remaining completely non-functional for reproduction. Rudimentary pistil flowers are characterized by= (1) a clearly recognizable pistil structure 0.4-0.8 mm long occupying the central flower position and showing distinct morphological regions corresponding to pistil organs (2) a basal swollen region recognizable as a rudimentary ovary, typically 0.2-0.4 mm in diameter though often lacking a clear ovary cavity or containing only aborted ovule primordia that fail to develop beyond early stages (3) a narrow mid-region recognizable as a rudimentary style, typically 0.1-0.3 mm long, though usually lacking functional transmitting tract tissue and sometimes appearing as a simple constriction rather than a true style (4) an apical region sometimes showing minimal stigmatic differentiation with slight expansion or roughening suggesting aborted stigma development, though lacking the characteristic wet exudate and papillae of functional stigmas (5) presence of vascular tissue extending into the rudimentary pistil base, demonstrating some degree of physiological integration despite functional sterility (6) well-developed stamens (erect, 2.6-4 mm filaments, 0.6-0.9 mm anthers, 26,000-40,000 pollen grains per flower) surrounding the rudimentary pistil. The rudimentary pistil represents developmental arrest at intermediate stages after substantial pistil organ differentiation has occurred, with ovary, style, and stigma primordia all initiating and undergoing partial elaboration before arrest prevents achievement of functional morphology. The size range 0.4-0.8 mm distinguishes rudimentary from vestigial pistils (which lack clear organ differentiation and measure 0.8 mm, approaching functional hermaphrodite pistil dimensions). Rudimentary pistil flowers achieve zero fruit set (0.0%) despite recognizable pistil morphology, as the abbreviated structures lack all functional reproductive components= stigmas are non-receptive or absent (lacking wet exudate and papillae), styles lack transmitting tract tissue necessary for pollen tube growth, ovaries lack functional ovule primordia capable of fertilization and embryo development, and the overall structure is incapable of supporting the pollination-to-fruit developmental sequence. Microscopic examination of rudimentary pistils reveals arrested development= ovary tissues show some cellular differentiation into epidermis, parenchyma, and vascular tissues but lack ovule initiation or show only aborted ovule primordia as small cellular masses style tissues may show some elongation and cellular differentiation but lack the specialized transmitting tract with its characteristic extracellular matrix stigma regions show minimal or absent stigmatic papillae and no secretory activity. Rudimentary pistil flowers are characteristic of a substantial proportion (30-50%) of male individuals in wild dioecious V. vinifera ssp. sylvestris populations, representing the most elaborate pistil development observed in male plants while remaining completely non-functional. The presence of rudimentary pistils demonstrates that male-determining genetic factors suppress pistil function not by completely preventing pistil development but by arresting development at intermediate stages after partial organ elaboration, suggesting regulatory mechanisms that modulate rather than eliminate pistil developmental programs.
Rudimentary_Pistil_Flower represents moderate pistil suppression with recognizable organ morphology and is distinguished from Vestigial_Pistil_Flower (minimal undifferentiated remnants) by presence of distinguishable ovary, style, and stigma regions and from Functionally_Male_Flower with short pistils (which may show more advanced development approaching functional dimensions despite impaired function). The rudimentary pistil condition demonstrates that sex-determining mechanisms can permit partial pistil development before imposing developmental arrest, providing insight into the timing and mechanisms of sex-determining gene action during flower organogenesis. Rudimentary pistils serve as valuable developmental models for understanding organ suppression mechanisms and the genetic control of organ elaboration versus arrest. The distinction between rudimentary and vestigial at approximately 0.4 mm and between rudimentary and short functionally male pistils at approximately 0.8 mm reflects natural discontinuities in the developmental continuum though some gradation exists at the boundaries.
Blüte mit reduziertem Stempel
abbreviated-pistil flower
reduced-organ male flower
rudimentäre Pistill Blüte
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 102-105 Caporali, E. et al. (2003). The arrest of development of abortive reproductive organs. Sexual Plant Reproduction 15=291-300 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Fechter, I. et al. (2012). Candidate genes within a 143 kb region of the flower sex locus in Vitis. Molecular Genetics and Genomics 287=247-259.
Rudimentary Pistil Flower
Wild V. vinifera ssp. sylvestris female plants in southeastern European and Near Eastern populations commonly exhibit rudimentary stamen flowers with recognizable stamen structures 0.6-1.2 mm long showing distinct basal filament region (0.4-0.8 mm, strongly reflex positioning structure 60-120° below horizontal) and terminal anther region (0.2-0.4 mm, swollen but lacking clear locules), producing no pollen or minimal non-viable pollen (<1,000 grains per flower, <8% viability when present), positioned well below the prominent functional pistil (2.2-2.4 mm length, ovary 1.3-1.5 mm diameter), achieving 30-38% fruit set through cross-pollination from nearby staminate males comparable to female only flowers despite stamen presence, and requiring microscopic examination to definitively characterize stamen reduction versus functionally female flower stamen morphology. Wild North American Vitis species female plants (V. labrusca, V. riparia) commonly bear rudimentary stamen flowers with stamens 0.7-1.4 mm showing recognizable filament and anther regions though strongly reflex and non-pollen-producing. Breeding populations segregating for sex determination produce rudimentary stamen pistillate individuals providing material for developmental genetic studies of stamen suppression mechanisms but requiring removal from self-fertile cultivar breeding programs due to absolute cross-pollination dependency.
A rudimentary stamen flower is a reduced stamen flower in which stamens show recognizable though severely abbreviated organ morphology with distinguishable filament and anther regions, typically measuring 0.5-1.5 mm in total length, exhibiting strong downward reflexing positioning structures well below the prominent pistil, and showing clear though drastically reduced stamen structure compared to functional hermaphrodite stamens (2.5-4 mm), representing moderate stamen suppression that permits partial organ elaboration before developmental arrest while remaining completely non-functional for pollen production and self-pollination. Rudimentary stamen flowers are characterized by= (1) clearly recognizable stamen structures 0.5-1.5 mm long occupying some or all of the five stamen positions and showing distinct morphological regions corresponding to stamen organs (2) a basal narrow region recognizable as a rudimentary filament, typically 0.3-1 mm long, often showing characteristic downward reflexing (curved 90-180° below horizontal) though abbreviated compared to the strong reflexing of longer functionally female flower stamens (3) a terminal swollen region recognizable as a rudimentary anther, typically 0.2-0.5 mm long though drastically reduced compared to functional anthers (0.6-1 mm), often lacking clear anther locules or dehiscence slits but showing some cellular differentiation suggesting aborted anther development (4) complete or near-complete absence of pollen production, with most rudimentary stamens producing no pollen, or in rare cases producing 1.5 mm and may produce marginal functional pollen though with strong reflexing preventing self-pollination). Rudimentary stamen flowers achieve fruit set only through cross-pollination, with fruit set percentages (28-40% under good pollination conditions) comparable to female only and vestigial stamen flowers, as the minimal non-viable pollen occasionally produced by rudimentary stamens is completely insufficient for self-fertilization and the strong stamen reflexing positions any pollen-producing tissue well below the stigma preventing pollen transfer even if viable pollen were produced. Microscopic examination of rudimentary stamens reveals arrested development= filaments show some cellular elongation and differentiation but remain abbreviated and often show reflex curvature anther tissues may show some wall layer differentiation but lack functional pollen mother cells, microsporogenesis, or viable pollen grain production. Rudimentary stamen flowers are characteristic of a substantial proportion (30-50%) of female individuals in wild dioecious V. vinifera ssp. sylvestris populations, representing the most elaborate stamen development observed in female plants while remaining completely non-functional for self-pollination. The presence of rudimentary stamens demonstrates that female-determining genetic factors suppress stamen function not by completely preventing stamen development but by arresting development at intermediate stages after partial organ elaboration, suggesting regulatory mechanisms that modulate rather than eliminate stamen developmental programs, creating intermediate phenotypes between complete stamen absence and functional male organ development.
Rudimentary_Stamen_Flower represents moderate stamen suppression with recognizable organ morphology and is distinguished from Vestigial_Stamen_Flower (minimal undifferentiated remnants) by presence of distinguishable filament and anther regions and from Functionally_Female_Flower (which has more developed stamens typically >1.5 mm that may produce marginal functional pollen despite strong reflexing). The rudimentary stamen condition demonstrates that sex-determining mechanisms can permit partial stamen development before imposing developmental arrest, providing insight into timing and mechanisms of sex-determining gene action during flower organogenesis. Rudimentary stamens serve as valuable developmental models for understanding organ suppression and the genetic control of organ elaboration versus arrest. The distinction between rudimentary and vestigial at approximately 0.5 mm and between rudimentary and functionally female stamens at approximately 1.5 mm reflects natural discontinuities though some gradation exists at boundaries.
Blüte mit reduzierten Staubblättern
abbreviated-stamen flower
reduced-organ pistillate flower
rudimentäre Staubblatt Blüte
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 103-106 Caporali, E. et al. (2003). The arrest of development of abortive reproductive organs. Sexual Plant Reproduction 15=291-300 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Badouin, H. et al. (2020). The wild grape genome sequence provides insights into the transition from dioecy to hermaphroditism. Genome Biology 21=223.
Rudimentary Stamen Flower
The Run1 presence in INRAE breeding line BC5 3-120 was confirmed by marker-assisted selection using the RGA-based markers developed by Barker et al. (2005) flanking the Run1 interval on chromosome 12 — this fifth backcross generation line carries a 5-8 cM M. rotundifolia introgression segment containing both Run1 and Rpv1, with over 97% V. vinifera genome recovery confirmed by genome-wide SNP analysis. Controlled inoculation with E. necator conidial suspensions at 10⁵ conidia/mL demonstrated complete resistance — no visible colony development at 14 days post-inoculation, compared to 60-80% leaf area coverage in susceptible V. vinifera controls, with microscopic examination revealing hypersensitive cell death in 85-95% of penetrated epidermal cells within 24 hours of appressorium formation, effectively halting pathogen development at the earliest post-penetration stage.
A Run locus information record documenting the presence or absence of the Run1 resistance locus on chromosome 12, derived from Muscadinia rotundifolia, that confers strong qualitative resistance to Erysiphe necator powdery mildew through a rapid hypersensitive response at the site of fungal penetration, co-localized with Rpv1 (downy mildew resistance) in a resistance gene cluster enabling simultaneous introgression of dual-disease resistance from a single genomic segment.
Run1_Presence documents the presence status of what is arguably the most valuable single resistance locus in grapevine breeding — valuable both for its exceptional resistance strength (near-complete immunity to powdery mildew) and for its co-localization with Rpv1 that provides simultaneous downy mildew resistance. The Run1 locus resides within a cluster of TIR-NBS-LRR (Toll/Interleukin Receptor - Nucleotide Binding Site - Leucine Rich Repeat) resistance gene analogs on chromosome 12, with the functional gene MrRGA10 identified as the likely Run1 candidate through map-based cloning and transgenic complementation studies: the encoded protein recognizes a pathogen-derived effector molecule (avirulence protein) and triggers the classical gene-for-gene hypersensitive response that kills the infected host cell before the biotrophic pathogen can establish nutrient acquisition. The gene-for-gene nature of Run1 resistance raises durability concerns analogous to those affecting Rpv3 — the pathogen could potentially evolve modified effectors that evade Run1 recognition — but the limited deployment of Run1 cultivars to date (confined to advanced breeding lines and a few recently released varieties) means that resistance durability has not yet been tested under commercial-scale selection pressure.
Muscadinia-Oidium-Resistenz
Run1 locus genotype
Run1 resistance status
Run1-Präsenz
Run1-Resistenzstatus
muscadine powdery mildew resistance chromosome 12
Barker CL et al. (2005) Genetic and physical mapping of the grapevine powdery mildew resistance gene, Run1, Theoretical and Applied Genetics 111: 370-377 Feechan A et al. (2013) Strategies for RUN1 deployment using RUN2 and REN2 to manage grapevine powdery mildew informed by studies of race specificity, Phytopathology 103: 1064-1070 Dry IB et al. (2010) Grapevine disease resistance genes, in Reynolds AG (ed) Viticulture and Wine Quality, Woodhead Publishing pp: 193-228
Run1 Presence
Transgenic validation of Run1_Resistance_Disposition was achieved at CSIRO by transforming susceptible V. vinifera 'Shiraz' with the MrRUN1 gene under its native promoter. Transformed lines expressing MrRUN1 exhibited complete resistance to E. necator, with inoculated leaves showing hypersensitive cell death at penetration sites and no subsequent colony establishment, identical to the phenotype observed in M. rotundifolia. Non-transformed 'Shiraz' controls developed extensive powdery colonies covering >80% of leaf surface by 14 days post-inoculation. Further characterization revealed that Run1 resistance is effective against E. necator isolates from Australia (genetic group B predominant), California (mixed A and B), and Europe (group B), indicating broad-spectrum resistance across pathogen diversity. In BC3 pseudo-backcross breeding lines with 87.5% theoretical V. vinifera background, Run1 resistance remained fully effective while sensory evaluation of micro-vinifications showed wine quality approaching commercial V. vinifera standards, demonstrating successful combination of disease resistance with acceptable enological characteristics.
A Run1_Resistance_Disposition is a Run_Mediated_Resistance_Disposition that inheres in a Vitis plant carrying the Run1 locus derived from Muscadinia rotundifolia and is realized through effector-triggered immunity mediated by the MrRUN1 TIR-NB-LRR receptor protein upon recognition of corresponding Erysiphe necator avirulence effectors. The Run1 locus maps to a resistance gene cluster on chromosome 12 that includes the linked Rpv1 locus for downy mildew resistance, enabling co-inheritance of both resistances through a single introgression event. The MrRUN1 gene was the first powdery mildew resistance gene functionally characterized in grapevine through transgenic complementation experiments demonstrating that its expression alone is sufficient to confer the resistance phenotype in susceptible V. vinifera backgrounds. Run1-mediated resistance operates primarily through pre-penetration and penetration-stage arrest, with hypersensitive cell death triggered rapidly upon pathogen recognition. The resistance is highly effective against diverse E. necator genetic groups including the A (Flag Shoot) and B (ascospore) populations present in different viticultural regions. Run1 represents a cornerstone of powdery mildew resistance breeding, with extensive deployment in European, American, and Australian breeding programs, though its Muscadinia origin requires extensive pseudo-backcrossing to restore V. vinifera genetic background and wine quality characteristics while maintaining resistance through marker-assisted selection.
Run1_Resistance_Disposition is_a Run_Mediated_Resistance_Disposition. Run1 is physically linked to Rpv1 on chromosome 12, and both loci are typically co-inherited in breeding programs. The MrRUN1 gene has been functionally validated through transgenic complementation. Run1 provides broad-spectrum resistance against diverse E. necator genetic groups. Breeding lines must be advanced through multiple backcross generations to recover V. vinifera quality characteristics while maintaining the resistance locus through marker-assisted selection.
MrRUN1 resistance
MrRUN1-vermittelte Resistenz
Run1 powdery mildew resistance
Run1-Oidium-Resistenz
Run1-Resistenz-Disposition
chromosome 12 PM resistance
Feechan A et al. (2013) Genetic dissection of a TIR-NB-LRR locus from the wild North American grapevine species Muscadinia rotundifolia, Molecular Plant-Microbe Interactions 26:684-695 Dry IB et al. (2010) Molecular strategies to enhance the genetic resistance of grapevines to powdery mildew, Australian Journal of Grape and Wine Research 16:94-105 Bouquet A (1986) Introduction dans l'espèce Vitis vinifera L. d'un caractère de résistance à l'oïdium (Uncinula necator) issu de l'espèce Muscadinia rotundifolia, Vignevini 12:141-146
Run1 Resistance Disposition
The Run2 presence was characterized by Riaz et al. (2011) in populations derived from M. rotundifolia accessions Trayshed and Magnolia, identifying two linked but distinct loci — Run2.1 mapping to chromosome 18 in a region syntenic with the Rpv3 downy mildew resistance cluster, and Run2.2 mapping to chromosome 14 near the Rpv12 region — with both loci conferring moderate to strong powdery mildew resistance (40-75% reduction in colony area compared to susceptible controls) through a combination of reduced penetration success and delayed colony expansion rather than the complete penetration-stage immunity of Run1. Importantly, race-specificity testing by Feechan et al. (2013) demonstrated that Run2 confers resistance to E. necator isolates virulent on Run1, confirming that Run2 recognizes different pathogen effectors than Run1 and providing the molecular basis for pyramiding Run1 + Run2 to achieve broad-spectrum resistance against diverse pathogen genotypes.
A Run locus information record documenting the presence or absence of Run2 resistance loci (Run2.1 on chromosome 18 and Run2.2 on chromosome 14) derived from Muscadinia rotundifolia, that confer resistance to Erysiphe necator powdery mildew through mechanisms at least partially distinct from Run1, providing additional muscadine-derived resistance sources for pyramiding strategies to achieve durable, broad-spectrum powdery mildew resistance.
Run2_Presence documents muscadine-derived powdery mildew resistance loci that complement Run1 by providing different pathogen recognition specificities. The discovery that M. rotundifolia carries multiple resistance loci on different chromosomes — each recognizing different pathogen effectors — mirrors the multi-locus resistance architecture documented for many plant-pathogen interactions and has direct implications for breeding strategy: whereas deployment of Run1 alone exposes the resistance to potential breakdown by pathogen races that have lost or modified the Run1-recognized effector, pyramiding Run1 + Run2 forces the pathogen to simultaneously evolve evasion of two independent recognition systems — a requirement that dramatically reduces the probability of resistance breakdown. The chromosomal co-localization of Run2.1 with Rpv3 (both on chromosome 18) and Run2.2 with Rpv12 (both on chromosome 14) suggests the existence of ancestral resistance gene clusters that have diversified to recognize different pathogen species, potentially enabling the co-introgression of powdery and downy mildew resistance from single chromosomal segments analogous to the Run1/Rpv1 co-localization on chromosome 12.
Muscadinia-Oidium-Resistenz-Sekundärloci
Run2 locus genotype
Run2 resistance status
Run2-Präsenz
Run2-Resistenzstatus
muscadine powdery mildew resistance secondary loci
Riaz S et al. (2011) Identification of mildew resistance in wild and cultivated Central Asian grape germplasm, BMC Plant Biology 11: 149 Feechan A et al. (2013) Strategies for RUN1 deployment using RUN2 and REN2 to manage grapevine powdery mildew informed by studies of race specificity, Phytopathology 103: 1064-1070 Dry IB et al. (2010) Grapevine disease resistance genes, in Reynolds AG (ed) Viticulture and Wine Quality, Woodhead Publishing pp: 193-228
Run2 Presence
Genetic mapping of Run2_Resistance_Disposition utilized a segregating population derived from the M. rotundifolia accession 'Magnolia', which exhibits strong powdery mildew resistance but lacks markers linked to the Run1 locus. Linkage analysis using SSR and SNP markers localized Run2 to a 3.2 cM interval on chromosome 18, distinct from the chromosome 12 location of Run1. Phenotypic evaluation using detached leaf assays with E. necator conidia (8,000/mL) demonstrated that Run2 confers complete resistance characterized by hypersensitive cell death at inoculation sites without subsequent mycelial development or sporulation, qualitatively similar to Run1. Critically, when a breeding line combining both Run1 and Run2 was challenged with E. necator isolates showing partial adaptation to Run1 (reduced but not eliminated sporulation on Run1-only genotypes), the pyramided Run1+Run2 line maintained complete resistance, validating the complementary recognition specificities of these loci. This finding supports the breeding strategy of combining multiple Run loci to enhance resistance durability in the face of pathogen evolution.
A Run2_Resistance_Disposition is a Run_Mediated_Resistance_Disposition that inheres in a Vitis plant carrying the Run2 locus derived from Muscadinia rotundifolia and is realized through effector-triggered immunity upon recognition of Erysiphe necator avirulence effectors distinct from those recognized by Run1. The Run2 locus maps to chromosome 18, genetically independent from the Run1/Rpv1 cluster on chromosome 12, and was identified in specific M. rotundifolia accessions that lack Run1. The independent chromosomal location and distinct recognition specificity of Run2 make it valuable for resistance gene pyramiding strategies, as combining Run1 and Run2 in a single genotype provides redundant recognition capacity that enhances durability against potential resistance-breaking pathogen variants. Run2-mediated resistance operates through similar mechanisms to Run1, with rapid hypersensitive response arresting pathogen development at early penetration stages. However, the molecular identity of the Run2 gene and its cognate effector remain to be fully characterized. Because Run2 originates from M. rotundifolia accessions distinct from those carrying Run1, introgression requires separate crosses and backcross programs, though marker-assisted selection enables efficient identification of carriers in segregating populations. The combination of Run1, Run2, and Ren loci from other Vitis species represents a comprehensive strategy for durable powdery mildew resistance in cultivated grapevines.
Run2_Resistance_Disposition is_a Run_Mediated_Resistance_Disposition. Run2 is located on chromosome 18, genetically independent from the Run1/Rpv1 cluster on chromosome 12. Run2 originates from different M. rotundifolia accessions than Run1, requiring separate introgression programs. The combination of Run1 and Run2 provides enhanced durability through redundant recognition of pathogen effectors. The Run2 resistance gene has not yet been molecularly cloned, unlike MrRUN1.
Chromosom-18-Mehltau-Resistenz
Run2 powdery mildew resistance
Run2-Oidium-Resistenz
Run2-Resistenz-Disposition
chromosome 18 PM resistance
second Muscadinia PM resistance
Riaz S et al. (2011) Identification of mildew resistance in wild and cultivated Central Asian grape germplasm, BMC Plant Biology 11:149 Blanc S et al. (2012) Identification and mapping of powdery mildew resistance locus MrRun2 in grapevine, Acta Horticulturae 948:175-181 Dry IB et al. (2019) Grapevine NLR immunity against downy and powdery mildew pathogens, Acta Horticulturae 1248:129-136
Run2 Resistance Disposition
The Run locus information compiled from molecular mapping studies documents two major loci — Run1 on chromosome 12 (co-localized with Rpv1 in a resistance gene cluster, Barker et al. 2005) and Run2.1 and Run2.2 on chromosomes 18 and 14 respectively (Riaz et al. 2011) — all derived from M. rotundifolia accessions collected from native muscadine populations in the southeastern United States where the co-evolutionary relationship with E. necator has produced highly effective resistance mechanisms. Run1-carrying breeding lines show complete suppression of powdery mildew colony establishment through rapid programmed cell death (hypersensitive response) at the site of appressorial penetration, preventing the pathogen from forming functional haustoria in epidermal cells — a qualitative immunity response that is among the strongest powdery mildew resistance phenotypes documented in any crop species.
A resistance gene information record specific to Run (Resistance from Vitis rotundifolia to Uncinula necator) loci — the genomic regions derived from Muscadinia rotundifolia conferring resistance to powdery mildew (Erysiphe necator, syn. Uncinula necator) in Vitis — documenting the locus designation, chromosomal location, resistance mechanism, diagnostic markers, and the unique value of muscadine-derived resistance genes that provide the strongest known single-locus powdery mildew resistance sources available for grapevine breeding.
Run_Locus_Information documents resistance genes from the taxonomically most distant Vitis relative used in grapevine breeding — Muscadinia rotundifolia (2n = 40), which is sufficiently divergent from Euvitis species (2n = 38) that initial hybridization requires embryo rescue and subsequent introgression demands extensive backcrossing over 4-6 generations spanning 25-30 years. Despite this practical difficulty, muscadine-derived resistance is of exceptional value because M. rotundifolia exhibits near-complete immunity to both powdery and downy mildew — a dual resistance phenotype that can be partly explained by the co-localization of Run1 and Rpv1 on chromosome 12 in a resistance gene cluster, enabling simultaneous introgression of both resistance specificities through a single genomic segment: this co-localization is a fortunate genetic architecture that greatly simplifies breeding programs seeking to combine powdery and downy mildew resistance. The Run designation was established to distinguish muscadine-derived powdery mildew resistance from the Ren (Resistance to Erysiphe necator) loci derived from Euvitis species, reflecting the different evolutionary origins and potentially different recognition specificities of these resistance sources.
Muscadinia-Mehltau-Resistenzlocus-Daten
Oidium-Resistenz-aus-Rotundifolia
Run gene characterization
Run-Locus-Information
muscadine powdery mildew resistance locus data
rotundifolia-derived Erysiphe resistance
Barker CL et al. (2005) Genetic and physical mapping of the grapevine powdery mildew resistance gene, Run1, using a bacterial artificial chromosome library, Theoretical and Applied Genetics 111: 370-377 Riaz S et al. (2011) Identification of mildew resistance in wild and cultivated Central Asian grape germplasm, BMC Plant Biology 11: 149 Dry IB et al. (2010) Grapevine disease resistance genes, in Reynolds AG (ed) Viticulture and Wine Quality, Woodhead Publishing pp: 193-228
Run Locus Information
The breeding program at CSIRO Agriculture and Food, Australia, has developed advanced selections carrying Run_Mediated_Resistance_Disposition through marker-assisted introgression from M. rotundifolia. The selection 2000-30-19, a BC4 generation pseudo-backcross with >96% V. vinifera genetic background, carries the Run1 locus as confirmed by the tightly linked VVIM11 marker. Controlled inoculation with E. necator isolates at 5,000 conidia/mL resulted in complete resistance, with hypersensitive cell death visible as small necrotic spots (diameter <0.5mm) at inoculation sites within 36 hours and no subsequent mycelial growth or conidiation through 21 days observation. Histological examination revealed that germinating conidia formed appressoria normally but penetration was arrested at the cell wall, with attempted penetration sites marked by intense callose deposition and rapid browning of the attacked epidermal cell. This strong, pre-haustorial resistance phenotype demonstrates the exceptional efficacy of Run1-mediated defense against powdery mildew establishment.
A Run_Mediated_Resistance_Disposition is a Powdery_Mildew_Resistance_Disposition that inheres in a Vitis plant carrying one or more Run (Resistance to Uncinula necator) loci and is realized through effector-triggered immunity upon recognition of corresponding Erysiphe necator avirulence effectors. The Run loci encode nucleotide-binding leucine-rich repeat (NLR) receptor proteins that detect specific fungal effectors secreted during attempted colonization, triggering rapid and robust defense responses that arrest pathogen development. The Run1 locus from Muscadinia rotundifolia is the most extensively characterized, located on chromosome 12 in a resistance gene cluster that also includes Rpv1 for downy mildew resistance. Run1 encodes a TIR-NB-LRR protein (MrRUN1) that recognizes an as-yet-unidentified E. necator effector, triggering programmed cell death at the site of attempted penetration. The Run2 locus, also from M. rotundifolia, represents an independent resistance gene with distinct recognition specificity located on chromosome 18. Run-mediated resistance phenotypes are characterized by rapid hypersensitive response, typically visible as small brown necrotic flecks within 48-72 hours post-inoculation, with no subsequent colony development or sporulation. The co-inheritance of Run1 with Rpv1 provides exceptional breeding value, enabling simultaneous introgression of resistance to both major fungal diseases from a single donor source, though the Muscadinia origin necessitates extensive backcrossing to restore V. vinifera quality characteristics.
Run_Mediated_Resistance_Disposition is_a Powdery_Mildew_Resistance_Disposition. The "Run" nomenclature derives from the former genus name Uncinula for the powdery mildew pathogen, now correctly Erysiphe. Run loci are distinguished from Ren loci by their origin in Muscadinia rotundifolia specifically. Run1 and Run2 both originate from M. rotundifolia but map to different chromosomes (12 and 18 respectively) and have independent recognition specificities. Run1 is physically linked to Rpv1 on chromosome 12, enabling co-selection for both powdery and downy mildew resistance.
Majorgen-Oidium-Resistenz
Muscadinia PM resistance
Run locus powdery mildew resistance
Run-Mehltau-Resistenz
Run-vermittelte Resistenz-Disposition
major gene oidium resistance
Hoffmann S et al. (2008) Resistance to Erysiphe necator in the grapevine 'Kishmish vatkana' is controlled by a single locus through restriction of hyphal growth, Theoretical and Applied Genetics 116:427-438 Feechan A et al. (2013) Genetic dissection of a TIR-NB-LRR locus from the wild North American grapevine species Muscadinia rotundifolia, Molecular Plant-Microbe Interactions 26:684-695 Dry IB et al. (2019) Grapevine NLR immunity against downy and powdery mildew pathogens, Acta Horticulturae 1248:129-136
Run Mediated Resistance Disposition
The salinity data for a Murray-Darling vineyard (southeastern Australia) documents the salinity profile characteristic of irrigated semi-arid viticulture — ECe of 1.8 dS/m at 0-30 cm (low-moderate salinity, below the yield-reduction threshold for most rootstocks), increasing to 3.5 dS/m at 30-60 cm and 5.2 dS/m at 60-90 cm (moderate-high salinity in the deeper root zone, sufficient to cause osmotic stress and restrict root function below 60 cm depth). The salt profile reflects the interaction of irrigation with saline groundwater — flood irrigation from the Murray River (EC 0.3-0.8 dS/m, low salinity) supplies water and leaches salts from the surface, but capillary rise from the shallow saline water table (1.5-2.5 m depth, EC 5-15 dS/m) deposits salts in the subsoil as the capillary water evaporates, creating the inverse salinity gradient (increasing with depth) that restricts effective rooting depth and reduces the vine's accessible water reservoir. The vineyard management strategy employs salt-tolerant rootstocks (Ramsey, 140Ru, 1103P providing 30-50% greater salt tolerance than own-rooted V. vinifera), deficit drip irrigation (replacing only 50-60% of evapotranspiration to minimize waterlogging while applying a leaching fraction to prevent surface salt accumulation), and subsurface drainage to lower the water table below the root zone.
A soil chemistry data component that records the concentration of soluble salts in the vineyard soil, measured as electrical conductivity of a saturated paste extract (ECe, expressed in dS/m) or of a 1:5 soil-to-water extract (EC₁:₅), with salinity being the principal soil chemical constraint in arid and semi-arid viticultural regions where evapotranspiration exceeds rainfall and dissolved salts accumulate in the root zone — Vitis vinifera being classified as moderately salt-sensitive, with yield reduction beginning at ECe values above 1.5-2.5 dS/m and severe impairment above 4-6 dS/m depending on rootstock, salt type, and irrigation management.
Salinity_Data documents the soil chemical stress that is the most widespread abiotic constraint on vineyard productivity in the world's irrigated viticultural regions — the Murray-Darling basin in Australia, the San Joaquin Valley in California, the Mendoza oasis in Argentina, parts of the Western Cape in South Africa, and the Central Valley of Chile all face salinity challenges associated with the combination of low rainfall, high evaporative demand, and saline irrigation water or shallow saline groundwater. The salt tolerance of Vitis vinifera is moderate — the vine maintains normal growth at ECe values below 1.5 dS/m (the threshold salinity), with approximately 10% yield reduction per dS/m increase above the threshold (the salt sensitivity coefficient), reaching 50% yield reduction at approximately 6-7 dS/m — but salt tolerance is strongly rootstock-dependent, with the most tolerant rootstocks (Ramsey/Salt Creek, 140Ru, 1103P) excluding sodium and chloride from the xylem sap through selective ion transport in the root cortex, while sensitive rootstocks (own-rooted V. vinifera, some V. riparia crosses) allow chloride accumulation in leaves to toxic levels (>25 mmol/L in petiole tissue): rootstock selection for salt tolerance is therefore the primary management strategy on saline sites, providing a genetic solution to a soil chemical problem. The chloride ion is the most toxic component of vineyard salinity — foliar chloride accumulation causes marginal leaf necrosis (leaf burn), reduced photosynthesis, premature leaf fall, and ultimately shoot dieback, while the sodium ion causes soil structural degradation (clay dispersion and reduced infiltration) at high exchangeable sodium percentage (ESP >15%), compounding the direct salt toxicity with deteriorating soil physical conditions.
Boden-Salinität
Salzgehalt-Daten
Versalzungs-Bewertung
electrical conductivity measurement
saline condition assessment
soil salt content
Zhang X et al. (2002) A review of the effects of salinity on grapevine and wine quality, South African Journal of Enology and Viticulture 23: 48-56 Walker RR et al. (2014) Rootstock effects on salt tolerance of irrigated field-grown grapevines, Australian Journal of Grape and Wine Research 20: 8-14 White RE (2003) Soils for Fine Wines, Oxford University Press
Salinity Data
Time-course analysis of Salt_Induced_ABA_Response_Disposition at UC Davis characterized ABA dynamics and downstream responses across 10 rootstocks following acute salt application (100 mM NaCl). Xylem sap and leaf ABA were measured at 0, 2, 6, 12, 24, and 72 hours post-treatment, with parallel assessment of stomatal conductance, NCED gene expression, and stress-responsive gene activation (VvP5CS, VvLEA, VvNHX1). Salt_Induced_ABA_Response_Disposition showed striking genotypic variation in kinetics and magnitude. '140 Ruggeri' exhibited rapid, strong ABA response with xylem sap ABA increasing from 45 ng/L (control) to 380 ng/L at 6 hours and 620 ng/L at 24 hours, driven by 8-fold VvNCED3 induction detectable within 2 hours. This rapid ABA accumulation triggered stomatal closure to 25% of control by 12 hours, VvP5CS induction (6-fold by 24 hours), and VvNHX1 upregulation (3.5-fold by 24 hours) enhancing vacuolar sequestration. 'K51-40' showed similarly rapid response with comparable protective outcomes. 'Schwarzmann' displayed delayed, weaker Salt_Induced_ABA_Response_Disposition with xylem ABA reaching only 185 ng/L at 24 hours, VvNCED3 induction delayed until 6 hours and peaking at only 3-fold, stomatal closure to 60% of control (incomplete), and minimal VvP5CS induction (1.8-fold). Correlation analysis revealed that ABA response magnitude at 24 hours predicted ion exclusion efficiency (r = 0.72), proline accumulation (r = 0.78), and overall salt tolerance (r = 0.68), demonstrating the central coordinating role of Salt_Induced_ABA_Response_Disposition. However, constitutively elevated ABA (observed in one genotype) was associated with growth penalty under non-stress conditions, indicating that optimal Salt_Induced_ABA_Response_Disposition involves inducibility rather than constitutive elevation.
A Salt_Induced_ABA_Response_Disposition is a Salt_Tolerance_Disposition that inheres in a Vitis plant and is realized through the capacity to rapidly synthesize and respond to abscisic acid (ABA) following salt stress perception, coordinating multiple protective responses including stomatal closure, ion transporter regulation, compatible solute accumulation, and stress-responsive gene expression. This disposition represents the hormonal signaling component of salt tolerance, with ABA serving as a central coordinator integrating diverse stress responses across tissues and timescales. Salt stress triggers ABA biosynthesis primarily in roots through upregulation of NCED (9-cis-epoxycarotenoid dioxygenase) genes, with ABA subsequently transported to shoots via the transpiration stream where it triggers stomatal closure reducing water loss and ion delivery to leaves. Beyond stomatal effects, ABA regulates expression of ion transporters (enhancing HKT1-mediated sodium retrieval, modulating NHX activity), activates compatible solute biosynthetic genes (P5CS for proline), induces protective proteins (LEA proteins, dehydrins), and triggers aquaporin regulation adjusting hydraulic conductance. The magnitude, speed, and sensitivity of the ABA response vary among genotypes, with rapid, strong ABA accumulation enabling faster protective responses but potentially excessive stomatal closure limiting carbon gain. Salt_Induced_ABA_Response_Disposition thus involves optimization of ABA signaling for effective protection without excessive growth limitation.
Salt_Induced_ABA_Response_Disposition is_a Salt_Tolerance_Disposition. ABA is the central hormone coordinating salt stress responses across multiple processes. NCED genes control ABA biosynthesis, with root expression critical for xylem-transported ABA signal. ABA regulates stomata, ion transporters, compatible solute genes, and protective proteins. Response kinetics (speed) and magnitude both contribute to effective protection. ABA signaling overlaps with drought response (see isohydric behavior), with salt-tolerant genotypes often showing strong ABA responsiveness. Optimal response is inducible rather than constitutive to avoid growth penalties.
ABA-mediated salt response
ABA-vermittelte-Salzantwort
Salinität-ABA-Signalisierung
Salzinduzierte-ABA-Antwort-Disposition
salinity-induced ABA accumulation
salt-ABA signaling
Zhu JK (2002) Salt and drought stress signal transduction in plants, Annual Review of Plant Biology 53:247-273 Cramer GR et al. (2011) Effects of abiotic stress on plants: a systems biology perspective, BMC Plant Biology 11:163 Ferrero M et al. (2020) A meta-analysis reveals the interactive role of ABA signaling in plant responses to stress, Frontiers in Plant Science 11:420
Salt Induced ABA Response Disposition
Comprehensive evaluation of Salt_Tolerance_Disposition at CSIRO Adelaide assessed growth, ion accumulation, and physiological responses across 20 rootstocks irrigated with saline water (ECw 0, 2, 4, 6 dS/m) over two growing seasons. Measurements included shoot growth, leaf chloride and sodium concentrations, leaf necrosis severity, chlorophyll fluorescence, and yield components. Salt_Tolerance_Disposition varied dramatically among rootstocks. '140 Ruggeri' (V. berlandieri × V. rupestris) exhibited exceptional tolerance, maintaining 85% of control growth at ECw 4 dS/m, restricting leaf chloride to 15 mmol/kg DW and sodium to 8 mmol/kg DW (versus external concentrations of 40 and 25 mM respectively), and showing no leaf necrosis. 'K51-40' (V. champinii selection) showed similar tolerance through effective ion exclusion. In contrast, 'Schwarzmann' (V. riparia × V. rupestris) displayed high salt sensitivity with growth reduced to 35% of control at ECw 4 dS/m, leaf chloride accumulating to 450 mmol/kg DW and sodium to 85 mmol/kg DW, and severe marginal necrosis affecting 60% of leaf area. Analysis of tolerance mechanisms revealed that ion exclusion capacity (particularly chloride exclusion) was the primary determinant of Salt_Tolerance_Disposition (r = -0.89 between leaf Cl⁻ and relative growth), with osmotic adjustment contributing secondarily. These findings demonstrate that rootstock selection based on Salt_Tolerance_Disposition enables productive viticulture on sites with moderate salinity that would be unsuitable with sensitive rootstocks.
A Salt_Tolerance_Disposition is an Abiotic_Stress_Tolerance_Disposition that inheres in a Vitis plant and is realized through physiological mechanisms that enable maintenance of growth, cellular function, and productivity under elevated soil salinity conditions that would otherwise cause ion toxicity, osmotic stress, and nutritional imbalances. This disposition represents the capacity to tolerate salt stress, increasingly important as vineyard expansion into marginal lands, irrigation with saline water, and climate-driven salinization affect viticultural regions worldwide. Salt stress in grapevine causes injury through multiple pathways including osmotic effects (reduced water uptake due to low soil water potential), ion toxicity (accumulation of sodium and chloride to damaging concentrations in leaves), and nutritional disruption (interference with uptake of essential nutrients, particularly potassium and calcium). Grapevine is classified as moderately salt-sensitive, with yield reductions typically beginning at soil electrical conductivity (ECe) of 1.5-2.5 dS/m and severe damage above 4 dS/m, though substantial genotypic variation exists. Salt tolerance mechanisms encompass ion exclusion (preventing toxic ion uptake and translocation to shoots), ion compartmentalization (sequestering accumulated ions in vacuoles or less sensitive tissues), osmotic adjustment (accumulating compatible solutes to maintain water uptake), and tissue tolerance (cellular mechanisms enabling function despite elevated ion concentrations). In grafted grapevines, Salt_Tolerance_Disposition is primarily determined by rootstock genotype, making rootstock selection the primary management tool for saline sites.
Salt_Tolerance_Disposition is_a Abiotic_Stress_Tolerance_Disposition. Salt tolerance in grapevine is primarily determined by rootstock genotype in grafted vines. The major tolerance mechanism is ion exclusion, particularly of chloride, addressed through child class Ion_Exclusion_Disposition. Grapevine is more sensitive to chloride than sodium toxicity, distinguishing it from many other crops. Salt tolerance involves both osmotic and ionic components, with ion toxicity typically dominating at moderate salinity levels encountered in viticulture. Rootstocks with V. berlandieri, V. rupestris, and V. champinii parentage generally show superior salt tolerance.
Salinität-Toleranz
Salzstress-Resistenz
Salztoleranz-Disposition
halotolerance
salinity tolerance
salt stress resistance
Walker RR et al. (2002) Rootstock effects on salt tolerance of irrigated field-grown grapevines, Australian Journal of Grape and Wine Research 8:146-155 Munns R, Tester M (2008) Mechanisms of salinity tolerance, Annual Review of Plant Biology 59:651-681 Fisarakis I et al. (2001) The effect of salinity on the growth, yield and quality of Vitis vinifera L. cv. Sultanina, Journal of Horticultural Science and Biotechnology 76:750-756
Salt Tolerance Disposition
Ramsey (Salt Creek, V. champinii) demonstrates high salt tolerance supporting Thompson Seedless table grape production in the Coachella Valley, California, where irrigation water ECw reaches 1.5-2.0 dS/m and soil ECe commonly exceeds 4 dS/m, maintaining leaf Cl⁻ concentrations below 0.3% dry weight. 1103 Paulsen (V. berlandieri × V. rupestris) provides salt tolerance to Primitivo in the Salento peninsula, Puglia, where coastal influence and intensive irrigation have elevated soil salinity to ECe 2.5-3.5 dS/m. Ruggeri 140 (V. berlandieri × V. rupestris) supports Nero d'Avola in southwestern Sicily on soils with marine salt influence, maintaining productivity at ECe levels that cause severe chloride toxicity symptoms on Kober 5BB.
An abiotic stress-tolerant rootstock role that inheres in a grafted rootstock by virtue of its capacity to maintain scion productivity and survival under elevated soil salinity conditions. Salt-tolerant rootstocks protect the scion from ion toxicity through mechanisms of ion exclusion (restricting uptake and translocation of Na⁺ and Cl⁻ to the shoot), ion compartmentalization (sequestering toxic ions in root vacuoles and older leaves), osmotic adjustment (accumulating compatible solutes to maintain water uptake against soil osmotic potential), and maintenance of K⁺/Na⁺ homeostasis in leaf tissue essential for enzyme function and stomatal regulation. Salinity tolerance is quantified by threshold electrical conductivity of soil saturation extract (ECe) at which yield declines commence (tolerant rootstocks maintaining yields to ECe of 3-4 dS/m versus 1.5-2.0 dS/m for sensitive genotypes), by leaf ion concentrations under saline irrigation (tolerant genotypes maintaining leaf Cl⁻ below 0.5% and Na⁺ below 0.25% dry weight), and by the slope of yield decline above threshold (tolerant rootstocks showing 5-8% yield loss per dS/m versus 12-15% for sensitive rootstocks).
Salt_Tolerant_Rootstock_Role is subclassed by ion-specific exclusion mechanisms (Chloride_Excluding_Rootstock_Role, Sodium_Excluding_Rootstock_Role) and by tolerance magnitude (High_Salt_Tolerance_Rootstock_Role, Moderate_Salt_Tolerance_Rootstock_Role). Chloride toxicity is generally the primary limitation for grapevine under salinity, as Vitis species accumulate Cl⁻ more readily than Na⁺ and show Cl⁻ toxicity symptoms (marginal leaf necrosis) at lower tissue concentrations than Na⁺ toxicity symptoms. Climate change, irrigation intensification, and aquifer depletion are expanding the geographic relevance of salt tolerance as a rootstock selection criterion in traditionally non-saline regions.
Unterlage für salzbelastete Böden
saline soil tolerant rootstock
salinity tolerant rootstock
salt resistant rootstock
salt tolerant rootstock
salztolerante Unterlage
versalzungstolerante Unterlage
Walker RR et al. (2002) Rootstock effects on salt tolerance of irrigated field-grown grapevines, Australian Journal of Grape and Wine Research 8: 146-155 Fisarakis I et al. (2001) The effect of salinity on growth and physiology of four grapevine rootstocks, Scientia Horticulturae 89: 123-134 Downton WJS (1977) Influence of rootstocks on the accumulation of chloride, sodium and potassium in grapevines, Australian Journal of Agricultural Research 28: 879-889
Salt Tolerant Rootstock Role
A same-season sylleptic shoot on Vitis vinifera cv. Merlot, axillary bud initiated at node 8 in mid-May during primary shoot elongation, continuous development through June without dormancy, lateral shoot emerged early July, elongated through July-August reaching 52cm length with 8 leaves, growth cessation and lignification initiation in September, complete development cycle from bud initiation (mid-May) through lignified cane status (October) occurring within single April-October growing season, representing typical same-season sylleptic pattern where lateral completes entire development without dormancy period, this shoot by season end contributing to mature cane structure as lignified lateral branch potentially bearing compound buds for following season though typically removed during winter pruning as laterals rarely retained in cane-pruned systems.
A sylleptic lateral shoot completing its entire development cycle within the same growing season as the primary shoot, characterized by axillary bud formation, immediate shoot development, elongation, maturation, and lignification all occurring within single season without dormancy, practical synonymy with prompt lateral designation in viticulture emphasizing same-season completion, and botanical emphasis on non-dormant continuous development from initiation through maturation distinguishing from any delayed or following-season patterns. Same-season sylleptic shoots represent complete single-season lateral development, with entire life cycle from axillary bud initiation through shoot emergence, growth, leaf development, and eventual lignification occurring within continuous growing season, typical pattern for most prompt laterals in viticulture where same-season timing is standard expectation for lateral development when it occurs.
Same-season sylleptic shoots demonstrate complete lateral life cycle within single growing season, with this pattern being standard for most prompt laterals in viticulture regardless of whether botanical sylleptic terminology is employed. The same-season completion means lateral contributes to current-year canopy function (photosynthesis supporting current crop), develops its own axillary buds (potentially producing tertiary shoots or following-season growth), and achieves lignification status by season end (transitioning from herbaceous shoot to woody cane material). In practical management, same-season sylleptic shoots are handled as prompt laterals with decisions about retention versus removal based on canopy density, positioning, photosynthetic contribution, and training system requirements, with most laterals removed during winter pruning even when well-developed since cane-pruning systems typically utilize primary shoots as fruiting wood while spur-pruned systems occasionally retain lateral-bearing canes if positioned appropriately. The same-season sylleptic designation emphasizes botanical completeness of development within annual cycle.
Ganzjahres-Seitentrieb
complete same-year lateral
einsaisonaler sylleptischer Trieb
single-season lateral
within-season sylleptic
Same Season Sylleptic Shoot
The sand content of the Graves appellation soils (Bordeaux, named from the French graves, gravel) documents high sand content of 50-75% in the upper 60 cm — dominated by medium to coarse sand fractions (0.25-2.0 mm) derived from the Quaternary fluvial gravel deposits of the ancestral Garonne river. This high sand content produces a well-drained, rapidly permeable soil (saturated hydraulic conductivity 50-150 mm/hour) with low water-holding capacity (60-100 mm plant-available water per meter of soil depth), creating the naturally water-limiting conditions that restrict vine vigor and promote the moderate water stress during ripening that Seguin identified as the common characteristic of Bordeaux's highest-quality vineyard soils. The sandy texture also produces rapid soil surface warming in spring (sand has lower heat capacity and lower thermal conductivity than clay), advancing budbreak by 3-5 days compared to the cooler clay soils of the Pomerol plateau only 30 km to the east.
A soil texture description component that records the proportion of sand-sized particles (0.05-2.0 mm diameter in the USDA/international system, subdivided into very fine sand 0.05-0.10 mm, fine sand 0.10-0.25 mm, medium sand 0.25-0.50 mm, coarse sand 0.50-1.0 mm, and very coarse sand 1.0-2.0 mm) in the fine earth fraction of the vineyard soil, expressed as a percentage by weight, with sand content being the primary determinant of soil macroporosity, drainage rate, and aeration that influence root penetrability, water percolation speed, and the thermal behavior of the soil surface.
Sand_Content records the coarse mineral fraction that creates the macropore network determining a soil's drainage and aeration characteristics. Sand particles (visible to the naked eye at sizes above 0.1 mm) are composed primarily of quartz (SiO2) in most viticultural soils, with minor contributions from feldspar, mica, and other primary minerals depending on the parent material — the mineralogical composition of the sand fraction is relatively unimportant for vine nutrition (quartz is essentially inert) but affects the thermal properties of the soil surface (dark-colored minerals absorb more radiation than light-colored quartz) and the long-term weathering potential of the soil: the physical effect of sand particles is determined by their size and packing arrangement — well-sorted sand with uniform particle size produces the largest macropores and fastest drainage, while poorly sorted sand with a range of particle sizes produces smaller pores as fine particles fill the interstices between coarse particles. The sand subfraction distribution (the proportion of very fine through very coarse sand) is pedologically informative — soils with predominantly fine sand (0.1-0.25 mm) behave similarly to silty soils in terms of water retention (fine sand pores retain more capillary water than coarse sand pores), while soils with predominantly coarse sand (0.5-2.0 mm) produce the free-draining, low-retention character typically associated with sandy vineyard soils.
Sandanteil-Messung
Sandfraktion
Sandgehalt
coarse particle proportion
sand fraction record
sand percentage measurement
Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873 White RE (2003) Soils for Fine Wines, Oxford University Press Soil Survey Staff (2014) Keys to Soil Taxonomy, 12th edition, USDA-NRCS van Leeuwen C et al. (2004) Influence of climate, soil, and cultivar on terroir, American Journal of Enology and Viticulture 55: 207-217
Sand Content
Cabernet Sauvignon clone 08 (FPS) bears a Sanitary_Selection_Clone_Role, the UC Davis Foundation Plant Services having applied thermotherapy and meristem culture to an important California selection, eliminating GLRaV-3 infection detected in the original material and releasing the sanitized version as confirmed virus-free stock. Pinot Noir clone 667 serves in a Sanitary_Selection_Clone_Role from ENTAV-INRA sanitation programs, existing selections subjected to virus elimination protocols and subsequent comprehensive testing to produce pathogen-free mother plants for certified propagation. A German Riesling clone designated "virusfrei" (virus-free) bears a Sanitary_Selection_Clone_Role, JKI sanitation protocols having eliminated detected pathogens from traditional selections to produce health-improved material for the certified propagation system.
A selection status clone role that inheres in a grapevine clone whose development prioritized pathogen elimination, producing virus-tested material from existing selections through thermotherapy, meristem culture, or other sanitation treatments. Sanitary selection focuses on health status rather than genetic improvement, applying pathogen elimination protocols to clones already identified for desirable agronomic or enological characteristics, producing "sanitized" versions of existing selections confirmed free of targeted pathogens. The approach emerged from recognition that many historically valuable clones harbored virus infections reducing their potential performance, with sanitation enabling these clones to express their full genetic potential free from pathogen-induced limitations. Sanitary selection programs, pioneered at INRA (France), UC Davis (USA), and other institutions from the 1960s-1980s, developed and refined thermotherapy (37-38°C for 60+ days), meristem tip culture, and combined protocols achieving virus elimination rates of 70-95% depending on pathogen and protocol. Sanitized clones may exhibit substantially different field performance than their infected progenitors—yield increases of 20-50% are common following virus elimination, with additional quality and longevity benefits.
Sanitary_Selection_Clone_Role captures the health-focused dimension of clone development, where pathogen elimination rather than genetic selection is the primary objective. Sanitary selection acknowledges that many valuable selections harbor virus infections accumulated through centuries of vegetative propagation, with sanitation "unlocking" their genetic potential by removing pathogen-imposed limitations. The approach complements rather than replaces genetic selection—contemporary programs typically integrate both dimensions, selecting for agronomic and enological merit while ensuring pathogen-free status. Sanitary selection effectiveness depends on pathogen and protocol nepoviruses (GFLV) respond well to thermotherapy, while leafroll viruses (GLRaV-3) often require combined thermotherapy-meristem culture approaches. The performance difference between infected and sanitized versions of the same clone demonstrates both the substantial impact of virus infection and the potential value of sanitation investment.
Gesundheitsklon
Gesundheitsselektion
health-improved clone
sanierte Selektion
sanitary selection
sanitized clone
virus-eliminated clone
viruseliminierter Klon
Walter B, Martelli GP (1996) Clonal and sanitary selection of the grapevine, INRA Editions Golino DA et al. (2017) Foundation Plant Services, HortScience 52: 1190-1194 Panattoni A et al. (2013) Grapevine virus elimination by thermotherapy, Journal of Plant Pathology 95: 229-235 Maliogka VI et al. (2015) Grapevine virus diseases, APS Press
Sanitary Selection Clone Role
Syrah sapwood at 15 years post-planting shows 22-28 mm radial depth comprising outermost 7-9 annual rings, contains functional xylem vessels 140-280 μm diameter (earlywood) conducting 42-68 L/day during peak summer transpiration, living parenchyma cells storing 68-92 g starch (8-13% dry weight in outer rings, 5-8% in inner rings), ray parenchyma providing radial transport, and lighter cream-tan coloration compared to darker central heartwood
Sapwood is the functional outer wood zone in Vitis vinifera trunk comprising recent annual growth rings where xylem vessels actively conduct water and minerals from roots to canopy, parenchyma cells store carbohydrate reserves and remain metabolically active, and ray parenchyma provides radial transport, representing the physiologically active wood essential for vine water relations, nutrient transport, and carbohydrate storage. Sapwood develops as vascular cambium continuously produces new secondary xylem= most recently formed rings (typically outermost 5-12 rings in mature vines) retain functional conducting capacity, parenchyma cells remain alive and metabolically active storing starch and participating in transport, xylem vessels remain open and water-filled (or refillable after embolism) enabling water transport, and ray parenchyma maintains radial transport connecting xylem vessels with cambium and phloem. Sapwood radial depth varies by vine age and physiology= young vines (3-5 years) showing 10-20 mm depth comprising entire wood radius (no heartwood yet formed), mature vines (8-20 years) showing 18-30 mm depth (typically outermost 6-10 annual rings), old vines (25-50+ years) showing 15-28 mm depth (outermost 5-9 rings, depth relatively constant while total trunk radius continues increasing), with sapwood depth relatively stable across cultivars and sites compared to highly variable total trunk diameter. Sapwood anatomical characteristics include= xylem vessels with open lumens free of tyloses or gums (enabling water flow), vessel walls with functional bordered pits (allowing lateral water movement), living parenchyma cells with intact cell membranes and organelles (maintaining metabolic activity), starch-filled parenchyma cells particularly in outer sapwood (providing carbohydrate storage), ray parenchyma connecting radially across growth rings (enabling lateral transport and communication), and lighter wood coloration (cream to pale tan compared to darker heartwood from absence of phenolic deposits). Sapwood hydraulic function includes= water transport from roots to transpiring canopy (providing 20-100 L/day in mature vines depending on canopy size and environmental conditions), xylem sap flow primarily through large earlywood vessels (single large vessel contributing disproportionately to total conductance), hydraulic conductance varying by sapwood cross-sectional area and vessel characteristics (typical specific conductivity 2-8 kg m⁻¹ MPa⁻¹ s⁻¹), vulnerability to embolism particularly in large vessels (water columns breaking under tension creating air-filled non-conducting vessels), and potential embolism repair (though limited in grapevine, some recovery possible through parenchyma-mediated refilling). Sapwood carbohydrate storage function includes= primary site of trunk starch reserves (sapwood containing 80-95% of total trunk stored carbohydrates), storage concentration highest in outer sapwood (8-15% dry weight in recently formed rings), declining storage in inner sapwood (4-8% in older sapwood approaching heartwood transition), seasonal cycling (accumulation post-harvest reaching peak November-December, mobilization spring March-May supporting budbreak and early growth, partial replenishment during growing season), and storage capacity proportional to sapwood volume (larger sapwood cross-sections providing greater reserve buffering). Sapwood parenchyma metabolic functions include= starch synthesis and hydrolysis (converting glucose to starch for storage, mobilizing starch to soluble sugars for transport and use), respiratory metabolism (parenchyma cells consuming oxygen and producing CO₂), hormone synthesis and response (responding to auxin, cytokinin, abscisic acid signals), defense compound synthesis (producing antifungal compounds, phenolics in response to pathogen attack), and embolism sensing and response (potentially detecting vessel dysfunction and initiating repair processes). Sapwood seasonal dynamics include= winter dormancy with minimal metabolic activity (starch reserves peak, low respiration, minimal water transport), spring activation with reserve mobilization (starch conversion to sugars, increasing respiration, water transport resuming), summer peak function (maximum water transport, active photosynthate storage, high metabolic activity), and fall transition to dormancy (carbohydrate accumulation, declining water transport, metabolic slowdown). Sapwood vulnerability and dysfunction includes= freeze damage causing vessel embolism (ice formation disrupting water columns particularly in large vessels), drought-induced embolism (tension in water columns exceeding vessel wall strength causing cavitation), pathogen colonization (trunk disease fungi invading sapwood vessels and parenchyma), vascular occlusion from tyloses (parenchyma cells growing into vessels blocking water flow in response to pathogens or dysfunction), and age-related transition to heartwood (inner sapwood rings losing function through parenchyma death and vessel occlusion becoming heartwood). Sapwood significance in viticulture includes= determining vine hydraulic capacity (sapwood area correlating with maximum water transport and supportable leaf area), providing carbohydrate reserves (trunk sapwood reserves critical for spring growth, fruit development, stress recovery), indicating vine health and vigor (sapwood depth and quality reflecting vine physiological status), and representing target for trunk disease management (protecting functional sapwood from pathogen colonization critical for maintaining vine health).
Sapwood emphasizes functional outer wood zone and is distinguished from Heartwood (non-functional inner wood) and specific sapwood subtypes (Active_Sapwood, Conducting_Sapwood, Outer_Sapwood) by encompassing entire physiologically active wood zone. Critical for understanding vine hydraulic capacity, trunk carbohydrate storage, water stress physiology, and trunk disease impacts on functional wood.
Conducting xylem
Functional wood
Funktionales Xylem
Lebendholz
Living wood
Splintholz
Sapwood
Investigation of Scaphoideus_Titanus_Resistance_Disposition at the Edmund Mach Foundation, San Michele all'Adige, Italy, compared leafhopper performance and feeding behavior across 15 cultivars using choice and no-choice bioassays. In choice tests offering simultaneous access to leaf discs from different cultivars, S. titanus adults showed significant settling preferences, with 3.2-fold more individuals on preferred 'Chardonnay' compared to less-preferred 'Teroldego' after 24 hours. In no-choice development assays, nymphal survival from first instar to adult ranged from 78% on 'Chardonnay' to 45% on 'Cabernet Sauvignon', with development time extended by 4-6 days on less-suitable hosts. Electrical penetration graph (EPG) analysis of feeding behavior revealed that S. titanus achieved sustained phloem ingestion (E2 waveform >30 minutes) on 85% of 'Chardonnay' plants but only 52% of 'Teroldego' plants, with significantly shorter phloem access phases on resistant genotypes. Despite these differences in Scaphoideus_Titanus_Resistance_Disposition, field FD infection rates did not differ significantly between cultivars in mixed plantings with equivalent vector pressure, suggesting that observed resistance levels are insufficient to prevent transmission under epidemic conditions. This finding emphasizes that S. titanus resistance may contribute to but cannot replace chemical vector control for FD management.
A Scaphoideus_Titanus_Resistance_Disposition is a Flavescence_Doree_Resistance_Disposition that inheres in a Vitis plant and is realized through host characteristics that prevent or limit Flavescence dorée transmission by the leafhopper Scaphoideus titanus (Hemiptera: Cicadellidae), the specific and obligate vector of 'Candidatus Phytoplasma vitis' in European vineyards. This disposition represents indirect phytoplasma resistance operating by interrupting the vector-mediated transmission cycle rather than through direct anti-phytoplasma mechanisms. S. titanus is a Nearctic species accidentally introduced to Europe in the mid-20th century, where it has become the sole epidemiologically significant FD vector due to its monophagous feeding on Vitis, efficient phytoplasma acquisition and transmission, and high reproductive potential in vineyard ecosystems. Resistance mechanisms affecting S. titanus may include leaf surface properties (trichome density, wax composition) deterring settling and probing behavior, phloem sap composition unfavorable for sustained feeding, antibiosis factors reducing leafhopper survival or fecundity, and induced defenses triggered by feeding that reduce transmission efficiency. Variation in S. titanus population development and feeding preferences among Vitis genotypes has been documented, with some cultivars and rootstocks supporting lower populations or less sustained feeding than others. Vector resistance could complement chemical control strategies, potentially reducing insecticide requirements in integrated management programs. However, no strong S. titanus resistance has been identified that provides practical FD protection, and mandatory insecticide applications remain required in FD-regulated areas regardless of cultivar.
Scaphoideus_Titanus_Resistance_Disposition is_a Flavescence_Doree_Resistance_Disposition. This class represents resistance to the insect vector rather than to the phytoplasma itself, providing potential indirect disease protection. S. titanus is monophagous on Vitis and the sole significant FD vector in Europe, making vector resistance a relevant target. However, no strong resistance providing practical field protection has been identified, and mandatory insecticide treatments remain required regardless of cultivar. Vector resistance research is complicated by the regulated status of FD requiring contained facilities for transmission experiments with viruliferous insects.
FD vector resistance
Scaphoideus feeding resistance
Scaphoideus-titanus-Resistenz-Disposition
Zikaden-Vektor-Resistenz
leafhopper vector resistance
Chuche J, Thiéry D (2014) Biology and ecology of the Flavescence dorée vector Scaphoideus titanus, Journal of Pest Science 87:381-393 Bressan A et al. (2005) Feeding behavior of Scaphoideus titanus Ball on different host plants, Bulletin of Insectology 58:115-122 Galetto L et al. (2018) Acquisition of Flavescence dorée phytoplasma by Scaphoideus titanus Ball from different grapevine varieties, Bulletin of Insectology 71:5-12
Scaphoideus Titanus Resistance Disposition
Scientific publication reference inventory for Grüner Veltliner documents the key research milestones establishing the modern scientific understanding of the cultivar: Sefc KM et al. (1998) "Identification of microsatellite sequences in Vitis riparia and their applicability for genotyping of different Vitis species," Genome 41(3): 367-373 (providing the SSR markers subsequently used for parentage determination) Regner F et al. (2000) "Genetic relationships among Pinots and related cultivars," American Journal of Enology and Viticulture 51(1): 7-14 (establishing the Traminer × St. Georgen parentage) Siebert TE et al. (2008) "Development of a stable isotope dilution analysis for the quantitation of rotundone," Journal of Agricultural and Food Chemistry 56(10): 3745-3748 (identifying rotundone as the compound responsible for the characteristic white pepper aroma) and Bokulich NA et al. (2016) on regional microbiome signatures — collectively building the molecular, chemical, and oenological knowledge framework for contemporary Grüner Veltliner science. Scientific publication reference for Blaufränkisch documents emerging but rapidly expanding research including parentage studies, phenolic characterization, and clonal diversity assessments published primarily in Austrian, Hungarian, and German viticultural journals.
Scientific publication reference is a documentation source entity that documents the peer-reviewed research literature, conference proceedings, institutional technical reports, and doctoral theses that constitute the modern scientific knowledge base for grape cultivar characterization, encompassing molecular genetics publications (parentage analysis, population structure, QTL mapping, genome-wide association studies), viticultural research (phenology, physiology, agronomy, terroir studies), oenological research (fermentation science, wine chemistry, sensory analysis), plant pathology publications (disease resistance, virus studies), and breeding program reports, specifying for each reference the standard bibliographic metadata (authors, year, title, journal, volume, pages, DOI) and the specific contribution of the publication to cultivar knowledge, providing the contemporary scientific evidence base that complements and increasingly supersedes the historical and ampelographic traditions as the primary source of reliable cultivar information.
Scientific publication reference documentation captures the epistemological revolution that has transformed cultivar knowledge over the past three decades — the transition from a knowledge base grounded primarily in empirical observation, morphological description, and accumulated tradition to one increasingly anchored in molecular evidence, controlled experimental studies, and statistically validated findings. The molecular genetics revolution beginning in the 1990s represents the single most transformative development in cultivar knowledge — microsatellite genotyping (Bowers et al. 1996, Thomas and Scott 1993) provided the first reliable molecular fingerprints for unambiguous cultivar identification independent of environmental variation, parentage analysis revealed unexpected genetic relationships that overturned centuries of traditional origin narratives (Cabernet Sauvignon as Cabernet Franc × Sauvignon Blanc, Bowers and Meredith 1997 Chardonnay as Pinot × Gouais Blanc, Bowers et al. 1999), and whole-genome sequencing (Jaillon et al. 2007 for Pinot Noir, Velasco et al. 2007 for a heterozygous line) opened the era of functional genomics linking specific genes to quality-relevant phenotypes. The publication landscape for viticultural science spans multiple disciplinary journals — viticulture-specific (American Journal of Enology and Viticulture, Australian Journal of Grape and Wine Research, OENO One), plant science (BMC Plant Biology, The Plant Journal, Plant Physiology), food chemistry (Journal of Agricultural and Food Chemistry, Food Chemistry), genetics (Nature Genetics, Heredity, Theoretical and Applied Genetics), and sensory science (Food Quality and Preference, Chemical Senses) — reflecting the inherently interdisciplinary nature of comprehensive cultivar knowledge. The distinction between foundational publications (establishing fundamental knowledge that reshapes understanding of a cultivar — a parentage determination, a key aroma compound identification) and incremental publications (contributing data points within an established framework — additional clonal evaluations, regional adaptation trials) helps prioritize references for ontological documentation. The open access movement and pre-print servers are progressively democratizing access to viticultural research that was historically restricted to institutional subscribers, though the peer-review quality control provided by established journals remains essential for evidence reliability.
Fachzeitschriftenreferenz
Forschungsliteratur
journal reference
peer-reviewed Quelle
peer-reviewed source
research publication
scientific literature reference
scientific reference
wissenschaftliche Publikationsreferenz
wissenschaftlicher Quellennachweis
Bowers JE, Meredith CP (1997) The parentage of a classic wine grape, Cabernet Sauvignon. Nature Genetics 16(1): 84-87 Bowers JE et al. (1999) Historical genetics: the parentage of Chardonnay, Gamay, and other wine grapes of northeastern France. Science 285(5433): 1562-1565 Jaillon O et al. (2007) The grapevine genome sequence suggests ancestral hexaploidization in major angiosperm phyla. Nature 449(7161): 463-467 Thomas MR, Scott NS (1993) Microsatellite repeats in grapevine reveal DNA polymorphisms when analysed as sequence-tagged sites. Theoretical and Applied Genetics 86(8): 985-990
Scientific Publication Reference
A Pinot Noir clone 777 vine in an ENTAV scion mother block bears a Scion_Mother_Plant_Role, trained to bilateral cordons and pruned annually to 12-16 renewal spurs, producing 50-80 one-year-old canes (approximately 200-400 usable buds) per vine for budwood harvest each December. A Chardonnay clone Mendoza 548 plant in a California certified nursery's scion block serves in a Scion_Mother_Plant_Role, providing budwood for benchgraft production destined for California, Oregon, and Washington vineyards. A Grüner Veltliner clone 17 maintained at the Austrian HBLA Klosterneuburg scion collection bears a Scion_Mother_Plant_Role, supporting the Austrian wine industry's access to certified scion material of the nation's signature variety.
A mother plant role that inheres in a grapevine maintained specifically as a source of scion budwood—the dormant buds and associated cane tissue used for grafting onto rootstocks to create the fruiting portion of grafted vines. Scion mother plants are cultivated under conditions optimizing budwood quantity and quality, including training systems maximizing cane production (often head-trained with annual renewal pruning), intensive nutrition and irrigation supporting vigorous vegetative growth, and management practices ensuring bud health (disease control, appropriate maturity at harvest). Scion mother plant blocks are maintained separately from rootstock mother plant blocks to prevent misidentification and mixing during harvest, with variety identification verified through visual inspection (ampelography) and increasingly molecular markers (SSR genotyping) at establishment and periodically thereafter. Scion budwood harvest occurs during dormancy (typically December-February in the Northern Hemisphere), with canes cut to 30-40 cm lengths, bundled, labeled, and stored under refrigeration (0-4°C, high humidity) until grafting or shipped immediately to recipient nurseries.
Scion_Mother_Plant_Role captures the functional specialization of mother plants providing grafting material for the fruiting/producing variety, complementing Rootstock_Mother_Plant_Role which provides the root-forming component. The distinction is operationally important—scion and rootstock mother blocks require separate management, distinct identification systems, and careful harvest protocols to prevent mixing. Scion mother plant productivity depends on training, vigor, and variety-specific bud fruitfulness, with typical yields of 40-100 usable buds per vine annually depending on training system and variety. Budwood quality parameters include cane diameter (typically 8-12 mm optimal for grafting), internode length (reflecting vigor and carbohydrate reserves), bud health (freedom from mechanical damage, freezing injury, or pathogen infection), and maturity (adequate lignification for dormancy and storage). Certification applies to scion mother plants at the same categorical levels (Pre-Basic, Basic, Certified) as the general mother plant hierarchy.
Edelreiser-Mutterpflanze
Propfreiser-Quelle
Reisermutterstock
budwood mother plant
grafting wood source
scion mother vine
scion source plant
Hartmann HT et al. (2011) Plant Propagation: Principles and Practices 8th Ed, Pearson pp.: 415-490 Gramaje D, Armengol J (2011) Fungal trunk pathogens in the propagation process, Phytopathologia Mediterranea 50: S217-S226 Dry N (2007) Grapevine rootstocks, Lythrum Press Walter B, Martelli GP (1996) Sanitary Selection of the Grapevine, INRA Editions
Scion Mother Plant Role
Pinot Noir bears a Scion_Role when grafted onto 3309 Couderc rootstock in Burgundy, contributing the varietal character, cluster morphology, and phenolic profile that define red Burgundy wines, while the rootstock provides root function and stress tolerance. Cabernet Sauvignon serves in a Scion_Role on 110 Richter in Napa Valley, determining the wine's tannin structure, cassis aromatics, and aging potential. Thompson Seedless (Sultana) bears a Scion_Role on Ramsey rootstock in the San Joaquin Valley, providing the seedless berry character essential for table grape and raisin markets.
A grapevine role that inheres in a plant or plant part of Vitis vinifera or a Vitis hybrid by virtue of its function as the above-ground fruiting portion of a grafted grapevine composite, contributing the cultivar identity, fruit characteristics, and wine or product quality attributes. The scion role is conferred when a plant or plant part is designated to provide the aerial vegetative structure (trunk above graft union, cordons, canes, shoots), foliage for photosynthesis, and reproductive organs (inflorescences, clusters, berries) of the grafted vine, while receiving water, mineral nutrients, and root-synthesized hormones from the rootstock partner. Scion cultivars determine the varietal character of wine or table grape products through their genetically determined berry composition (sugars, organic acids, phenolic compounds, aroma precursors), cluster architecture, ripening behavior, and responses to environmental conditions. A single cultivar may bear different scion roles depending on its utilization context—the same Muscat of Alexandria genotype may serve in a Wine_Production_Scion_Role in Languedoc, a Table_Grape_Scion_Role in Egypt, or a Raisin_Production_Scion_Role in California.
Scion_Role is the complementary partner to Rootstock_Role in the grafted vine composite that constitutes over 80% of the world's commercial vineyards. Unlike Rootstock_Role which is subclassed primarily by functional characteristics (stress tolerance, vigor effects), Scion_Role is subclassed primarily by utilization purpose (wine production, table grape, raisin, juice, distillation). The scion determines the commercial identity and market value of the product, while rootstock selection adapts the vine to site conditions. In ungrafted (own-rooted) vineyards, the same plant simultaneously provides both root and scion functions, but does not bear either role in the ontological sense as roles are relationally conferred.
Edelreis
Edelsorte
Pfropfreis
Reiser
fruiting variety
grafted variety
scion
scion cultivar
scion variety
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes: A Complete Guide to 1,368 Vine Varieties, Ecco/Allen Lane This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519
Scion Role
Cabernet Sauvignon scion trunk on grafted vine at 15 years post-planting shows 65-85 mm diameter measured 20 cm above graft union, 15 annual growth rings visible in cross-section averaging 2.5-4.5 mm ring width, well-developed secondary xylem with ring-porous anatomy (earlywood vessels 150-250 μm diameter), bark thickness 4-8 mm, and stored starch content 80-120 g per vine supporting spring budbreak and early shoot growth
A scion trunk is the above-ground woody perennial stem structure of Vitis vinifera that supports the canopy architecture, transports water and nutrients between roots and shoots, stores carbohydrate reserves, and in grafted vines represents the cultivar variety portion above the graft union, while in own-rooted vines comprises the entire trunk from soil level to cordon or cane positions. The scion trunk develops through progressive secondary growth involving vascular cambium activity that produces secondary xylem (wood) centripetally and secondary phloem centrifugally, periderm formation through phellogen (cork cambium) activity replacing primary epidermis with protective bark layers, and progressive diameter increase through annual growth rings. Scion trunk anatomy includes= central pith (parenchymatous core tissue, often disintegrating in older trunks creating hollow centers), secondary xylem comprising annual growth rings (wood tissue with vessels, fibers, and parenchyma organized in ring-porous pattern with large earlywood vessels 100-300 μm diameter and smaller latewood vessels 50-100 μm), vascular cambium (meristematic layer 2-6 cell layers thick producing secondary tissues), secondary phloem (inner bark with sieve tubes, companion cells, fibers, and parenchyma), and periderm (outer bark with cork cells, phellogen, and phelloderm providing protection and reducing water loss). Scion trunk diameter varies enormously by vine age and vigor= young vines (1-3 years) show 10-30 mm trunk diameter, mature productive vines (5-20 years) typically 40-100 mm diameter, old vines (20-50 years) may reach 100-200 mm diameter, and ancient vines (50+ years) can exceed 200-400 mm diameter with some century-old vines showing 500+ mm trunk girth. Scion trunk functions include= mechanical support of canopy weight and wind resistance (requiring sufficient diameter and wood strength to support 5-20 kg canopy mass), vascular transport connecting root water and nutrient uptake with canopy transpiration and photosynthesis (xylem sap flow rates 10-100 L/day during growing season for mature vines), carbohydrate storage in wood parenchyma and phloem (starch reserves 50-200 g per vine supporting spring growth and fruit development), and perennation enabling multi-year productivity (unlike annual plants, grapevine trunk persists and accumulates resources across decades). Scion trunk development stages include= establishment phase (years 1-3) with rapid diameter increase and structural development, productive maturity (years 4-20+) with stable annual growth and optimal fruit production, and senescence phase (variable onset) with declining vigor, reduced growth rates, and potential trunk deterioration. Scion trunk management in viticulture includes= trunk training systems (vertical trunk, angled trunk, multiple trunks), trunk protection from winter injury (wrapping, hilling, burial in extreme climates), trunk disease management (Eutypa, Esca, Botryosphaeria canker diseases), trunk renewal through sucker selection when primary trunk deteriorates, and consideration of trunk age effects on wine quality (old vine designations, potential quality benefits of mature vine physiology).
Scion_Trunk emphasizes the complete above-ground trunk structure and is distinguished from Own_Rooted_Trunk (ungrafted vine trunk) and grafted trunk (above graft union on rootstock) by encompassing all trunk types and developmental stages. Critical for understanding vine architecture and training systems, carbohydrate storage and spring mobilization, trunk disease impacts on vine health and longevity, and old vine physiology effects on fruit quality.
Hauptstamm
Main trunk
Rebstamm
Stammholz
Vine stem
Scion Trunk
A dormant cane of Pinot Noir clone 115 harvested from an ENTAV-certified scion mother plant bears a Scion_Wood_Role, bundled, labeled with variety and clone identification, stored at 2°C and 95% RH, then shipped to licensed nurseries for benchgraft production. Scion wood of Tempranillo clone RJ-43 serves in a Scion_Wood_Role for both budwood extraction (chip buds for T-budding onto established rootstocks) and graftwood preparation (two-node segments for omega benchgrafting). A green shoot of Sauvignon Blanc bears a Scion_Wood_Role in summer green grafting operations, with actively growing tissue grafted to green rootstock shoots for rapid union formation under greenhouse conditions.
A propagation material role that inheres in stem tissue harvested from a scion variety (typically Vitis vinifera or a fruiting hybrid) by virtue of its function as the source of buds or grafting material for the above-ground fruiting portion of grafted vines. Scion wood provides the cultivar identity, fruit characteristics, and varietal quality attributes to the grafted vine composite, with material designated as either budwood (individual buds excised for budding operations) or graftwood (cane segments used in grafting techniques). Scion wood quality parameters include bud viability (freedom from mechanical damage, freezing injury, desiccation, or pathogen infection), cane maturity (adequate lignification and carbohydrate reserves), appropriate diameter for the intended grafting technique, and verified cultivar/clone identity matching the intended variety. Scion wood harvest occurs during dormancy (for hardwood benchgrafting or dormant budding) or during the growing season (for green grafting or herbaceous budding), with material maintained under conditions preserving bud viability until use.
Scion_Wood_Role is subclassed into Budwood_Role (material providing individual buds) and Graftwood_Role (material providing cane segments for grafting), reflecting the distinct preparation and use of scion material in different propagation techniques. The scion determines the commercial identity and market value of the grafted vine—variety and clone selection for scion wood directly defines what wine, table grape, or other product the vineyard will produce. Scion wood certification follows the same Pre-Basic/Basic/Certified hierarchy as rootstock material, with health status (particularly freedom from leafroll viruses affecting fruit quality) equally important for scion as rootstock. International scion wood movement requires phytosanitary certification and may involve quarantine periods, reflecting the potential for graft-transmissible pathogen dissemination through the nursery trade.
Edelholz
Edelreiserholz
Reisermaterial
fruiting wood
scion material
scion wood
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 415-490 Walter B, Martelli GP (1996) Sanitary Selection of the Grapevine, INRA Editions European Union (2020) Commission Implementing Directive 2020/177 Gramaje D, Armengol J (2011) Fungal trunk pathogens, Phytopathologia Mediterranea 50: S217-S226
Scion Wood Role
A Pinot Noir vine on vigorous Oregon valley-floor site trained to Scott Henry bears a Scott_Henry_Trained_Role with upward-growing shoots from the lower cordon at 70 cm and downward-growing shoots from the upper cane at 100 cm, the divided architecture enabling quality Pinot production from a site too vigorous for conventional guyot. A Sauvignon Blanc vine in New Zealand's Hawkes Bay serves in a Scott_Henry_Trained_Role, the vertical division managing the variety's excessive vigor while maintaining distinct upper and lower fruit zones with different exposure characteristics. A Chardonnay vine on deep Willamette Valley soils bears a Scott_Henry_Trained_Role, the system's vigor management enabling quality sparkling wine base production from productive sites.
A training system role that inheres in a grapevine trained to the Scott Henry system, a vertically divided canopy architecture with shoots from alternating cordons or cane positions directed upward and downward, creating two fruiting zones stacked vertically on a single vine. Scott Henry training was developed in Oregon's Umpqua Valley in the 1980s by vineyard manager Scott Henry to address vigor management for wine grape varieties, adapting divided canopy principles for premium wine production rather than the juice grape applications of earlier systems. The system typically positions a low cordon (60-80 cm) with upward-trained shoots and a higher cordon or cane (90-110 cm) with downward-trained shoots, creating upper and lower curtains on the same vine that effectively double canopy surface area. Scott Henry combines the vigor management benefits of divided canopies with maintenance of the vertical shoot positioning valued for quality wine production in the upward-growing portion. The system enables high-quality fruit production from vigorous vines and fertile sites where single-curtain systems would produce excessive shading.
Scott_Henry_Trained_Role captures the wine-focused divided canopy system developed for quality production rather than volume maximization. The system's innovation lies in combining upward and downward shoot growth on the same vine, creating two distinct fruiting zones with different microclimate characteristics—the upper curtain receiving more exposure, the lower curtain more protected. Some producers exploit this zonal difference, harvesting upper and lower zones separately for different wine programs. Scott Henry adoption has been significant in vigorous wine regions (New Zealand, Oregon, parts of Australia) where the system enables quality production from sites that would produce mediocre fruit under conventional training. The system's complexity—managing two shoot directions, maintaining distinct zones, harvest logistics—limits adoption to situations where vigor management benefits justify additional management costs.
Auf-und-Ab-System
Scott Henry
Scott Henry training
Scott-Henry-Erziehung
up-and-down training
vertical divided canopy
vertikal geteilte Laubwand
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Reynolds AG, Vanden Heuvel JE (2009) Training systems, American Journal of Enology and Viticulture 60: 251-268 Henry S (1999) Scott Henry trellis development, Practical Winery and Vineyard 20: 42-48
Scott Henry Trained Role
A searching climbing functional tendril at node 11 on an actively extending shoot of Vitis vinifera cv. Riesling in the Rheingau, Germany, having achieved 21 cm extension length over two weeks development, exhibiting circumnutation movements sweeping through 145-degree arcs over 105-minute cycles, positioned 15 cm from the nearest vertical trellis wire with sweep radius of 12 cm indicating high probability of contact within 12-24 hours if shoot continues extension, showing bright green coloration and turgid texture indicating active physiological status, demonstrating rapid touch response when experimentally stimulated with 10-second contact pressure initiating coiling movements within 6 minutes, measured at BBCH stage 75 (berries pea-sized) in late June when shoot apex is 35 cm above the highest existing tendril attachment at node 8 and requiring new attachment points to prevent shoot bending or collapse under increasing weight.
A climbing functional tendril that has achieved full structural and physiological maturation but has not yet made contact with a support structure in Vitis vinifera, characterized by active circumnutation movements involving slow sweeping motions through arcs typically spanning 120-180 degrees over periods of 1-4 hours, maintenance of tissue turgor and flexibility enabling responsive bending upon contact detection, fully developed mechanosensory apparatus with responsive epidermal cells capable of rapid coiling initiation following sustained tactile stimulation, and positioning in the distal shoot region where new attachment points are required to enable continued vertical growth progression. Searching climbing functional tendrils represent the exploratory phase of climbing behavior, executing genetically programmed motor responses that increase the probability of encountering support structures despite the absence of directed growth toward specific targets. The circumnutation pattern involves differential growth rates on opposite sides of the tendril causing gradual rotation of the tendril tip through space, combined with some gravitropic influence that biases searching movements toward horizontal or upward directions where support structures are more likely to exist in natural climbing contexts. Searching efficiency depends on tendril length (longer tendrils sweep larger volumes), circumnutation amplitude (wider arcs sample more space), movement speed (faster rotations increase encounter probability per unit time), and spatial relationship between tendril position and available support structures.
Searching climbing functional tendrils are distinguished from immature tendrils (not yet capable of coiling response), from attached tendrils (having completed contact and attachment), and from senescent or damaged tendrils (having lost functional capacity without achieving attachment). The searching phase duration varies from hours to weeks depending on support structure availability, with tendrils in well-designed training systems achieving contact rapidly while those in unsupported or poorly positioned situations may search indefinitely until senescence. Environmental conditions affect searching efficiency, with drought stress reducing turgor and circumnutation amplitude, excessive heat potentially causing tissue damage, and wind creating mechanical stimulation that may trigger inappropriate coiling responses. Searching tendrils that fail to achieve contact before tissue senescence represent lost climbing opportunities requiring subsequent tendrils to provide needed support.
circumnutating tendril
explorative Kletterorgan
exploratory tendril
pre-attachment tendril
seeking tendril
vor-befestigungs Ranke
Based on Darwin (1875) "The Movements and Habits of Climbing Plants" John Murray, London Jaffe & Galston (1968) "The physiology of tendrils" Annual Review of Plant Physiology 19=417-434 Stolarz (2009) "Circumnutation as a visible plant action and reaction" Plant Signaling & Behavior 4=380-387 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 283-304.
Searching Climbing Functional Tendril
A second basal leaf at node 2 on a cane-pruned shoot of Vitis vinifera cv. Riesling in Mosel Valley, Germany, emerging 3 days after first leaf visibility at 8 days post-budbreak, measuring 16.2 cm length by 15.8 cm width representing 94% of first leaf area (17.3 cm × 16.5 cm), exhibiting characteristic Riesling morphology with five lobes, deep sinuses, pronounced lobe points, and distinctive marginal teeth more reliably diagnostic than somewhat variable first leaf morphology, achieving full expansion at 26 days post-budbreak (compared to 23 days for first leaf), demonstrating peak photosynthesis rate of 13.2 μmol CO₂ m⁻² s⁻¹ in mid-May contributing approximately 35% of total shoot photosynthate working with first leaf (40% contribution) to provide 75% of shoot carbon during critical early development, positioned opposite the second position tendril or occasionally second inflorescence in highly fruitful shoots, experiencing progressive shading from 80% full sun in May to 30% by July as upper canopy develops, showing senescence symptoms in early August including chlorosis and reduced function, measured at shoot BBCH stage 79 (majority of berries touching), representing typical second basal position with high early contribution and characteristic decline trajectory.
A basal leaf at node 2 representing the second true foliage leaf on a developing shoot in Vitis vinifera, characterized by position immediately following the first basal leaf in the phyllotactic sequence, emergence typically 2-4 days after first leaf visibility representing rapid succession during early shoot development, development from preformed primordium created during previous growing season within dormant bud, dimensions often approaching or occasionally exceeding first leaf size typically within 10-20% of first leaf area reflecting similar developmental priority as early-emerging basal position, morphological characteristics generally showing more typical cultivar-diagnostic features compared to first leaf including characteristic lobe development and sinus patterns, early achievement of maturity typically 3.5-4.5 weeks post-budbreak slightly delayed compared to first leaf but substantially earlier than median position leaves, high contribution to early shoot carbon balance often providing 25-40% of total shoot photosynthate during weeks 3-7 following budbreak working synergistically with first leaf to support shoot extension and reproductive development, and progressive functional decline through canopy development and shading with eventual senescence or removal during canopy management. Second basal leaves share with first leaves the developmental advantage of preformation within dormant buds enabling rapid deployment and early function, but occupy slightly less favorable positions relative to vascular connections and resource mobilization pathways, creating subtle developmental delays and occasionally reduced size compared to first leaves while maintaining critical early-season functional importance.
Second basal leaves are distinguished from first leaves by node 2 position and slightly delayed emergence, and from higher position leaves by early emergence timing, preformed development, and typically larger size. Morphological reliability often improves at node 2 compared to node 1, with cultivar-diagnostic features including lobe patterns, sinus characteristics, and tooth morphology more consistently expressed at this position. Size relationships between first and second leaves vary with cultivar and environmental conditions, with some varieties consistently producing larger first leaves while others show equal or occasionally larger second leaves. Functional importance parallels first leaves during early development, with the first and second leaves together typically providing 60-80% of shoot photosynthate during the critical 3-7 week post-budbreak period supporting shoot extension, inflorescence development, and fruit set. Management practices targeting basal leaves including early leaf removal often address first and second leaves together or sequentially, with removal timing and extent determining impacts on vine carbon balance and fruit development.
Knoten-zwei Blatt
node two leaf
second position leaf
secondary basal leaf
zweite Position Weinblatt
zweites basales Blatt
Based on Poni et al. (2006) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 57=397-407 Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 1-57 Intrieri et al. (2008) "Grapevine growth and development" in Biology of the Grapevine, Cambridge University Press pp. 44-81 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 155-178.
Second Basal Leaf
Kyoho in Japan and Flame Seedless in California demonstrate strong second crop potential disposition, with managed lateral crop contributing 15-30% of annual production. Following spring frost damage to primary buds, Chardonnay secondary buds produce 40-60% of normal yield, while Merlot secondary buds achieve only 25-40% recovery. In tropical viticulture (Brazil, Thailand), second crop potential enables double-cropping systems with two harvests annually, achieving combined yields of 25-40 t/ha. The interval between primary and secondary harvest ranges from 3-8 weeks depending on cultivar and climate.
A fruitfulness disposition that is the inherited capacity of a Vitis vinifera plant to produce a commercially significant secondary crop from lateral shoots, secondary buds following primary bud loss, or late-developing inflorescences, providing yield recovery potential or extended harvest in suitable climates.
Second crop potential encompasses both intentional double-cropping systems exploiting lateral fruitfulness and recovery scenarios where secondary buds compensate for primary bud loss. Tropical viticulture deliberately manipulates vine dormancy to achieve multiple cropping cycles. Second crop quality typically differs from primary crop due to different ripening conditions, with applications in specific wine styles or table grape markets. Climate suitability (growing season length, heat accumulation) determines commercial viability of second crop utilization.
Nebenerntepotential
Sekundärertragsfähigkeit
Zweitertragspotential
double cropping disposition
recovery crop potential
secondary crop capacity
Dry PR, Coombe BG (2004) Viticulture Volume 1: Resources, 2nd Ed, Winetitles pp.: 178-195 Miele A, Mandelli F (2012) Grapevine canopy management in tropical viticulture. Acta Horticulturae 931:469-476
Second Crop Potential Disposition
Regent (released 1996, JKI) bears a Second_Generation_PIWI_Role as the benchmark improved red PIWI, its Diana × Chambourcin parentage combining resistance from French-American hybrid ancestry with quality improvement sufficient for German QbA and Prädikatswein production—approximately 2,000 hectares planted demonstrating commercial viability. Johanniter (released 2000, Freiburg) serves in a Second_Generation_PIWI_Role as an improved white PIWI with Riesling in its parentage, producing wines of recognized quality while substantially reducing fungicide requirements. Solaris (released 2001, Freiburg) bears a Second_Generation_PIWI_Role with exceptional disease resistance and early ripening, enabling quality wine production in northern European regions previously considered unsuitable for viticulture.
A PIWI cultivar role that inheres in improved disease-resistant varieties developed from the 1990s through 2010s, representing substantial advancement in wine quality compared to first-generation PIWIs while maintaining or improving disease resistance. Second-generation PIWIs emerged from deliberate quality improvement programs that crossed first-generation resistant varieties or complex hybrids with elite V. vinifera parents, selecting progeny combining resistance with reduced hybrid character and improved wine attributes. These varieties demonstrated that interspecific resistance breeding could produce wines acceptable to broader consumer audiences, enabling expanded adoption beyond the niche markets served by first-generation PIWIs. Second-generation varieties often carry identified R-genes (Rpv, Ren/Run loci) from mapped resistance donors, enabling marker-assisted breeding and deliberate R-gene pyramiding for durable resistance. Key second-generation PIWIs (Regent, Johanniter, Solaris, Cabernet Cortis) achieved significant commercial plantings and regulatory acceptance for quality wine production, establishing PIWI cultivation as a mainstream viticultural option rather than an experimental novelty.
Second_Generation_PIWI_Role captures the quality breakthrough that transformed PIWI varieties from experimental curiosities to commercially viable alternatives. The generation is characterized by deliberate quality selection alongside resistance maintenance, with breeding programs systematically evaluating wine character through standardized protocols. Second-generation PIWIs typically show residual quality differences from V. vinifera (detectable in trained taster evaluation) but produce wines acceptable to general consumers and qualifying for quality wine appellations. Regulatory acceptance expanded significantly for second-generation PIWIs, with German wine law reforms and EU framework evolution permitting quality designations for approved resistant varieties. The commercial success of second-generation PIWIs (Regent reaching significant German market share) validated resistance breeding investment and stimulated continued development toward third-generation varieties with further quality improvement.
PIWI zweite Generation
improved PIWI
intermediate resistant variety
quality-improved PIWI
qualitätsverbesserte Resistenzsorte
second-generation PIWI
verbesserte PIWI-Sorte
Töpfer R et al. (2011) Molecular breeding in grapes, CRC Press pp.: 160-185 Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330 Pedneault K, Provost C (2016) Fungus resistant grape varieties, Applied Microbiology and Biotechnology 100: 3499-3519 PIWI International (2020) PIWI Variety Descriptions
Second Generation PIWI Role
Syrah grafted on 110 Richter rootstock at 16 years showing second-order lateral secondary roots branching from first-order laterals, representative second-order lateral 1.8 cm diameter at origin from first-order parent extending 52 cm length at depth 48 cm, second-order laterals emerging approximately 9 cm intervals along first-order lateral, second-order lateral showing limited secondary growth with 4-5 annual xylem rings, third-order fine roots <5 mm diameter branching from second-order lateral at 4-7 cm intervals creating dense absorptive network, second-order lateral xylem conducting water from third-order network to first-order lateral, terminal portions showing root hairs and mycorrhizal colonization
Second order lateral secondary root is lateral secondary root branching from first-order lateral secondary root representing second hierarchical level in root system architecture, characterized by moderate diameter typically 0.8-3 cm at origin from first-order parent decreasing to <1 cm at distal portions, intermediate structural development showing limited secondary growth compared to first-order laterals, predominantly absorptive and transport function rather than primary structural role, and serving as critical transition between large structural first-order laterals and fine absorptive third and higher-order roots, with second-order laterals typically emerging 5-15 cm intervals along first-order lateral length, showing horizontal or slightly descending orientation, producing third-order branches that generate finest absorptive roots, and representing largest-diameter roots still actively involved in direct absorption while also conducting resources from higher-order branches to first-order lateral and ultimately to primary root and trunk providing essential link in hierarchical water and nutrient transport pathway. Second order lateral secondary root hierarchical position includes= branching from first-order laterals (second-order roots arising from first-order lateral secondary roots as second branching level), representing intermediate order (second-order being middle level in hierarchy between large first-order and fine high-order roots), serving as parent to higher orders (second-order laterals producing third-order branches which may produce fourth and fifth-order roots), showing order-2 designation (in hierarchical numbering system where root order increases with each branching level from primary root), and demonstrating transitional role (second-order laterals bridging between structural first-order framework and absorptive fine root network). Second order lateral secondary root diameter and structure includes= showing moderate diameter (second-order laterals typically 0.8-3 cm at origin substantially smaller than first-order but larger than fine roots), exhibiting gradual taper (diameter declining from base to tip as third-order branches emerge), demonstrating limited secondary growth (second-order laterals showing some secondary xylem development but less extensive than first-order), displaying relatively thin bark (periderm present but less developed than larger roots), and revealing intermediate anatomy (second-order structure transitional between woody first-order and primary anatomy fine roots). Second order lateral secondary root origin pattern includes= emerging along first-order lateral (second-order roots arising at intervals along first-order lateral parent length), showing spacing typically 5-15 cm (second-order lateral intervals closer than first-order from primary root reflecting finer branching scale), arising from first-order pericycle (lateral origin from parent internal tissues similar to first-order from primary root), demonstrating abundant production (each first-order lateral potentially producing 10-30+ second-order branches depending on length), and exhibiting acropetal trend (second-order laterals more abundant near first-order growing tip declining toward base). Second order lateral secondary root orientation and length includes= growing predominantly horizontal (second-order laterals maintaining horizontal or slightly oblique trajectory following first-order orientation), extending moderate distances (individual second-order laterals typically 30-80 cm length exploring local soil volume around first-order parent), occupying similar depth to parent (second-order laterals at comparable depth to first-order parent typically 30-90 cm though potentially descending slightly), showing flexible trajectory (second-order laterals adjusting growth direction responding to soil conditions and obstacles more readily than larger first-order), and creating local exploration (second-order laterals thoroughly exploring soil zone around first-order lateral parent). Second order lateral secondary root tertiary branching includes= producing third-order branches (second-order laterals generating numerous third-order roots creating next hierarchical level), showing close branch spacing (third-order roots typically emerging 3-8 cm intervals along second-order parent reflecting increasingly fine branching), exhibiting high branching density (second-order laterals potentially producing 10-25+ third-order branches per lateral creating dense network), demonstrating progressive refinement (branching scale decreasing with each order reflecting transition toward fine absorptive roots), and creating absorptive network (second-order lateral with third and higher-order branches forming dense absorptive cluster in local soil). Second order lateral secondary root functional emphasis includes= transitioning from structural to absorptive (second-order laterals showing mixed function with both transport and direct absorption), conducting water from higher orders (second-order lateral xylem moving water from third-order and finer roots toward first-order and primary roots), transporting nutrients basipetally (second-order laterals conveying minerals absorbed by higher-order network toward plant body), showing direct absorption capacity (second-order laterals particularly at tips potentially producing root hairs and absorbing directly), and supporting fine root network (second-order laterals providing framework and resources supporting ephemeral fine absorptive roots). Second order lateral secondary root in absorption network includes= representing largest absorptive roots (second-order laterals being largest-diameter roots still actively involved in direct water and nutrient uptake), showing root hair production (second-order lateral tips potentially bearing root hairs increasing absorption surface), hosting mycorrhizal colonization (second-order laterals particularly terminal portions showing arbuscular mycorrhizal associations), demonstrating responsive growth (second-order laterals proliferating rapidly in resource-rich soil patches), and exhibiting seasonal activity (second-order laterals showing active growth and absorption during growing season with reduced activity during dormancy). Second order lateral secondary root lifespan includes= persisting multiple years (second-order laterals typically functional 2-5 years longer than ephemeral fine roots but shorter than persistent first-order), showing eventual senescence (older second-order laterals declining in function and dying as newer second-order laterals produced), demonstrating replacement pattern (continuous production of new second-order laterals replacing lost or senescing laterals), exhibiting variable longevity (second-order lateral lifespan varying with environmental stress, pathogen pressure, and resource availability), and contributing to turnover (second-order lateral mortality and replacement part of overall root system turnover dynamics).
Second_Order_Lateral_Secondary_Root emphasizes branching from first-order lateral and intermediate position and is distinguished from First_Order_Lateral_Secondary_Root (larger, more structural) by focus on second branching level and transitional absorption-transport function. Critical for understanding hierarchical branching, structural-absorptive transition, intermediate transport role, and the second-level lateral roots bridging between structural framework and fine absorptive network.
Lateralwurzel zweiter Ordnung
Second branch lateral
Secondary lateral root
Sekundäre Seitenwurzel
Twice-branched lateral
Zweite Verzweigungsebene
Second Order Lateral Secondary Root
Pinot Noir on Riparia Gloire showing second-order lateral shallow roots, excavation revealing second-order laterals 1.4-2.8 cm diameter branching from first-order laterals every 8-12 cm, second-order laterals extending 35-85 cm horizontal distance at 18-42 cm depth, microscopic sections showing 3-5 annual rings, second-order laterals producing third-order branches every 5-8 cm creating extensive network, second-order lateral shallow roots demonstrating intermediate structural-absorptive function connecting framework to fine root network
Second order lateral shallow root is lateral shallow root representing second branching level originating from first-order lateral roots, typically 0.8-3 cm diameter, characterized by indirect connection to central root axis through first-order parent, intermediate structural development with limited to moderate secondary growth, transitional function between structural support and absorption, and serving as parent axes for third-order and higher branches that create complete absorptive network, with second-order lateral shallow roots showing typical characteristics including origin from first-order laterals at 5-15 cm intervals creating numerous branches per parent, predominantly horizontal orientation maintaining shallow depth 10-50 cm, lengths 20-100 cm extending lateral exploration beyond first-order termini, limited to moderate secondary growth potentially developing 2-8 annual rings in older roots, production of numerous third-order and higher branches supporting extensive fine root networks, and dual functions as intermediate structural elements distributing forces and providing local transport plus parent axes supporting majority of fine absorptive root production. Second order lateral shallow root origin and spacing includes= showing first-order parent origin (second-order laterals branching from first-order lateral roots not from primary roots), exhibiting endogenous initiation (second-order laterals originating from pericycle-like tissue inside first-order laterals), demonstrating regular spacing (second-order laterals emerging at roughly regular intervals 5-15 cm along first-order parent though variable), revealing abundant production (individual first-order laterals potentially producing 20-100+ second-order branches depending on length and age), and displaying progressive formation (second-order laterals forming sequentially as first-order parent elongates and ages creating age gradient). Second order lateral shallow root diameter and structure includes= showing intermediate diameter (second-order laterals 0.8-3 cm diameter at origin smaller than first-order 2-8 cm but larger than third-order 0.3-1.5 cm), exhibiting limited secondary growth (second-order laterals showing limited to moderate secondary thickening compared to extensive growth in first-order), demonstrating few annual rings (second-order laterals typically showing 2-8 rings versus 5-15+ in first-order indicating shorter lifespan or slower growth), revealing semi-woody structure (second-order laterals developing some woody tissue though less extensively than first-order), and displaying structural adequacy (second-order lateral structure sufficient for supporting higher-order branches and local forces). Second order lateral shallow root length and branching includes= showing moderate length (second-order laterals extending 20-100 cm from origin extending lateral exploration), exhibiting variable extent (individual second-order lateral length varying widely 10-150 cm depending on soil conditions and competition), demonstrating third-order production (second-order laterals producing numerous third-order branches supporting absorptive network), revealing branching density (second-order laterals potentially producing third-order branches every 3-10 cm creating intensive branching), and displaying hierarchical continuation (second-order branching creating third level in hierarchical architecture progressing toward terminal absorptive roots). Second order lateral shallow root functional characteristics includes= providing intermediate transport (second-order laterals transporting resources between fine roots and first-order laterals), exhibiting structural distribution (second-order laterals distributing mechanical forces from higher-order network across multiple branches reducing stress concentrations), demonstrating absorptive support (second-order laterals serving as parent axes from which majority of fine absorptive roots ultimately derive), revealing transitional role (second-order laterals representing transition between first-order structural emphasis and higher-order absorptive emphasis), and displaying critical position (second-order laterals occupying critical intermediate position connecting structural framework to absorptive network). Second order lateral shallow root lifespan and turnover includes= showing intermediate persistence (second-order laterals potentially functioning 3-10 years intermediate between first-order decades and fine roots months), exhibiting moderate turnover (second-order lateral mortality and replacement rates moderate between low first-order and high fine root turnover), demonstrating progressive mortality (second-order laterals showing increasing mortality rates with age as function declines), revealing replacement (second-order lateral losses replaced through continued production from first-order parents maintaining population), and displaying population dynamics (second-order lateral abundance relatively stable as production balances mortality). Second order lateral shallow root in hierarchical architecture includes= connecting structural to absorptive (second-order laterals linking first-order structural framework to higher-order absorptive network), enabling network expansion (second-order branching greatly increasing total root system length and soil contact), demonstrating multiplication (second-order laterals multiplying available parent axes for higher-order branching creating exponential proliferation), revealing architectural efficiency (second-order level allowing efficient root system organization balancing structure and absorption), and displaying functional specialization (second-order laterals showing intermediate specialization bridging extreme specializations of first-order and fine roots). Second order lateral shallow root absorptive contribution includes= supporting fine root production (second-order laterals and derivatives supporting thousands of fine absorptive roots per parent), exhibiting indirect absorption (second-order laterals providing absorption primarily through higher-order derivatives), demonstrating substantial contribution (second-order lateral-associated fine roots potentially providing 40-60% of total vine uptake), revealing seasonal patterns (second-order laterals producing pulses of fine roots spring and fall when conditions favorable), and displaying critical role (second-order lateral abundance and branching strongly affecting total root system absorption capacity). Second order lateral shallow root management relevance includes= affecting tillage depth (second-order lateral depth determining safe cultivation depth avoiding damage), influencing nutrient placement (second-order lateral distribution informing fertilizer application strategies), demonstrating irrigation response (second-order laterals and derivatives responding strongly to irrigation supporting increased production), revealing mulch benefits (mulches protecting second-order laterals from temperature extremes and disturbance), and displaying rootstock effects (rootstock varieties differing in second-order lateral production affecting root system architecture and function).
Second_Order_Lateral_Shallow_Root emphasizes second branching level from first-order parents and is distinguished from first-order by focus on intermediate hierarchy position. Includes Branch_Second_Order_Lateral_Shallow_Root and Distal_Second_Order_Lateral_Shallow_Root. Critical for understanding hierarchical organization, intermediate structure, transitional function, absorptive support, and the second-level branches connecting structural framework to absorptive network.
Intermediate lateral
Second-level branch root
Zweite Ebene Verzweigung
Zweite Ordnung laterale flache Wurzel
Second Order Lateral Shallow Root
A second-order secondary branch rachis on a primary inflorescence of Vitis vinifera cv. Merlot in Bordeaux, France, emphasizing its hierarchical position as second-order lateral branching, originating from a first-order primary branch at 3.2 cm from the primary's base, representing third-order position in complete hierarchy (following first-order main rachis and second-order primary branch), measuring 2.8 cm length, diameter of 1.4 mm at primary branch junction, containing 3 vascular bundles representing second-order lateral vascular capacity, bearing 5 tertiary branch rachides (fourth-order structures, third-order laterals) plus 6 direct-attached flowers, second-order lateral position creating intermediate-scale elaboration with this branch extending from the first-order primary branch creating additional three-dimensional expansion and flower distribution, complete branch system bearing approximately 22 flowers, serving as transitional structure between the major first-order primary branch (4.5 cm, 38 total flowers) and fine third-order tertiary branches (0.8-1.5 cm, 3-8 flowers each), measured at BBCH stage 61 (beginning of flowering) in early June, representing typical second-order secondary branch rachis in its role as intermediate lateral elaboration.
A secondary branch rachis emphasizing the second-order lateral position in the branching hierarchy in Vitis vinifera, characterized by hierarchical position as the second lateral branching order from the main rachis (first laterals being primary branches), origin from first-order primary branches creating the second level of lateral elaboration, third-order position in the complete rachis hierarchy (first-order main rachis, second-order primary branches, third-order secondary branches), role as intermediate structures between main primary branches and fine tertiary branches, dimensions reflecting the second lateral order typically 1.5-3.5 cm length with 2-3 vascular bundles, and functional significance creating the intermediate-scale architectural elaboration bridging from major primary branches to fine-scale flower-bearing structures. Second-order secondary branch rachides emphasize the hierarchical position as the second level of lateral branching, representing the intermediate elaboration creating fine-scale flower distribution, developing through the second wave of lateral meristem activity following primary branch establishment, and serving as the transitional structures between major architecture and fine detail.
Second-order secondary branch rachides emphasize the hierarchical position as the second level of lateral branching, distinguished from simple secondary branch designation by explicit focus on second-order lateral status. The second-order designation clarifies the branching hierarchy with these structures representing the second departure from the main axis (first departure being primary branches). Origin from first-order primary branches without additional intermediates defines the second-order lateral status, contrasting with third-order tertiary branches originating from secondary branches. Architectural role involves intermediate-scale elaboration, with second-order secondaries bridging between the major architecture defined by large primary branches and the fine detail created by small tertiary branches. Transitional function creates the progressive size and complexity gradient from main rachis through first-order primaries, second-order secondaries, and third-order tertiaries, defining the hierarchical organization. Developmental sequence shows second-order secondary branch primordia initiating along developing first-order primaries following primary branch establishment. Vascular organization shows second-order branches receiving traces from first-order primary bundles, creating the hierarchical vascular network. Dimensional characteristics show intermediate values between first-order primaries (3-7 cm, 2-3 mm diameter) and third-order tertiaries (0.5-2 cm, 0.5-1 mm diameter), reflecting the intermediate hierarchical position. Flower distribution creates intermediate-scale arrays with 15-30 flowers per second-order branch typical. Transformation to cluster maintains second-order lateral identity with these structures creating the intermediate-scale berry cluster architecture.
dritte Ordnung hierarchische Spindel
intermediate lateral branch
second lateral order rachis
third-order hierarchical rachis
zweite laterale Ordnung Achse
zweiter Ordnung sekundäre Zweigrachis
Based on Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 435-470 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 369-405 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 639-678.
Second Order Secondary Branch Rachis
Syrah grafted on Teleki 5C rootstock at end of second growing season (20 months post-grafting) showing second year grafted rootstock trunk 3.3 cm diameter (increase from 2.0 cm at end of first year), trunk extending 23 cm from root crown to graft union positioned 18 cm above soil, graft union displaying 2.2 cm diameter bulge with consolidated callus tissue, trunk bark developing with 0.6 mm thickness showing initial longitudinal fissures and grey-brown coloration typical of Teleki 5C, trunk still staked but showing improved self-support, diameter growth 1.3 cm during second season, no disease or damage, scion canopy 2.4 m total shoot length
Second year grafted rootstock trunk is young grafted rootstock trunk in second complete growing season following grafting, representing intermediate establishment stage characterized by consolidating graft union integration, continued rapid diameter growth typically increasing 0.5-1.0 cm, developing initial bark fissures as periderm matures, diameter range typically 2.0-4.0 cm, reduced but still significant vulnerability requiring continued protection, and marking transition from extreme early fragility toward more robust mature trunk characteristics. Second year grafted rootstock trunk temporal definition includes= spanning second complete growing season post-grafting (typically May-November of calendar year 2 in Northern Hemisphere, approximately months 12-20 post-grafting), representing intermediate young trunk stage (between first year extreme vulnerability and third year approaching maturity), beginning after first winter dormancy (resumption of growth in spring of year 2), concluding at second dormancy (end of second growing season entering second winter), and marking consolidation phase (building on first year foundation toward stable established trunk). Second year grafted rootstock trunk graft union maturation includes= showing consolidated callus tissue (first year callus tissue stabilizing and integrating though some additional callus still forming), establishing improved vascular continuity (xylem and phloem connections across graft union more complete and functional than first year though not yet fully mature), displaying moderating graft union bulge (swelling at graft union 1.5-2.8 cm diameter increase less prominent proportionally than first year as trunk diameter increases), showing more integrated appearance (graft union line less conspicuous as callus tissue matures and bark begins to obscure original grafting point), and approaching stable integration (graft union becoming mechanically stronger though not yet fully established as in mature trunk). Second year grafted rootstock trunk physical development includes= exhibiting intermediate diameter (2.0-4.0 cm typical range reflecting continued rapid growth), showing developing bark (initial periderm formation creating 0.4-1.0 mm bark thickness with beginning of characteristic texture), displaying early fissuring (first longitudinal furrows or cracks appearing in bark as cork accumulation begins and trunk expansion stresses surface), revealing transitional coloration (bark color shifting from light tan-brown toward darker grey-brown as periderm matures and weathers), and showing more robust appearance (trunk looking more substantial and less fragile than first year though still obviously young). Second year grafted rootstock trunk continued vulnerability includes= still susceptible to mechanical damage (developing but not yet thick bark and moderate diameter providing improved but not complete protection against injury), sensitive to severe environmental stress (freeze damage or sunburn still possible though tolerance improved compared to first year), requiring continued staking (trunk achieving greater strength but typically still needing support preventing wind damage particularly at graft union), showing moderate disease susceptibility (improving resistance compared to first year but not yet mature trunk resistance to cankers and infections), and needing protective measures (trunk protection from sun or cold still beneficial in extreme climates though less critical than first year). Second year grafted rootstock trunk vascular development includes= showing improved water transport capacity (expanding xylem cross-sectional area supporting 15-35 liters daily water conduction depending on canopy development), establishing better nutrient distribution (increased phloem area improving bidirectional transport between roots and scion), displaying more complete vascular integration (most graft union vascular discontinuities resolved creating efficient transport pathway), forming second annual ring (adding second growth ring to trunk cross-section building transport and support capacity), and enabling greater resource storage (increasing ray parenchyma tissue in trunk beginning to accumulate carbohydrate reserves though capacity still limited compared to mature trunk). Second year grafted rootstock trunk growth patterns includes= showing continued rapid diameter increase (0.5-1.0 cm growth typical in second season depending on rootstock vigor and conditions), displaying proportionally greater trunk expansion than graft union (trunk diameter growing faster than graft union bulge reducing proportional prominence of swelling), exhibiting minimal height increase (2-6 cm vertical trunk growth primarily from graft union area), focusing resources on diameter expansion (rapid radial growth building structural capacity and transport area), and showing influence of canopy development (expanding scion canopy requiring greater trunk capacity driving diameter growth). Second year grafted rootstock trunk management requirements includes= maintaining stake support (continued staking protecting trunk and graft union though some self-support capacity developing), monitoring graft union health (inspecting for disease symptoms, checking integration quality, ensuring no mechanical damage), implementing appropriate trunk protection (painting or wrapping in extreme climates though less critical than first year), providing adequate irrigation (supporting continued rapid growth and graft union consolidation without waterlogging), removing rootstock suckers (eliminating any shoots from trunk below graft union maintaining focus on scion development), and adjusting canopy management (pruning and training scion appropriate to developing trunk capacity avoiding overload). Second year grafted rootstock trunk transport capacity includes= supporting moderate canopy (providing water and nutrients for developing scion canopy typically 1.5-3 m shoot length), enabling first crop potential (some vigorous scion cultivars potentially bearing small first crop on second year trunk though often removed), conducting adequate photosynthate flow (improved phloem capacity distributing carbohydrates from expanding canopy to root system), showing improved graft union transport efficiency (better vascular connections reducing transport resistance at graft interface), and building toward mature capacity (second year trunk approaching but not yet achieving full mature trunk transport capability). Second year grafted rootstock trunk bark development includes= forming continuous periderm (cork cambium activity creating protective bark layer 0.4-1.0 mm thickness replacing first year epidermis), showing initial texture patterns (beginning of characteristic furrows or ridges specific to rootstock cultivar appearing), displaying transitional colors (bark weathering from initial light tan to more mature grey-brown tones), developing visible lenticels (gas exchange pores prominent on developing bark surface as horizontal lines or dots), and showing rootstock-specific characteristics (bark patterns beginning to reflect rootstock variety though not yet fully expressed). Second year grafted rootstock trunk structural development includes= achieving improved mechanical strength (second year wood adding stiffness and break resistance compared to first year), showing better wind resistance (increased diameter and lignification providing greater stability though staking still beneficial), displaying more rigid properties (wood becoming less flexible and more brittle approaching mature wood characteristics), forming distinct annual rings (second growth ring visible in cross-section marking second year development), and building load-bearing capacity (trunk able to support expanding scion canopy weight and potential fruit load). Second year grafted rootstock trunk in different rootstock types includes= vigorous rootstocks (1103P, 110R) potentially reaching 3.5-4.5 cm diameter by end of year 2, moderate vigor rootstocks (SO4, 3309C, Teleki 5C) typically achieving 2.8-3.8 cm diameter second year, low vigor rootstocks (Riparia Gloire, 101-14) showing slower growth reaching 2.0-3.0 cm diameter second year, and cultivar-specific bark development (some rootstocks showing pronounced fissuring by year 2 while others maintaining smoother bark longer). Second year grafted rootstock trunk damage recovery includes= healing minor wounds (small mechanical injuries from first year or second year rapidly closing through callus formation), repairing limited freeze damage (cambium injury from moderate cold potentially recovering during second season growth), overgrowing minor trunk damage (expanding diameter gradually covering over small scars or injuries), compensating for limited disease (mild infections potentially contained through wound response and continued vigorous growth), and building resilience (improving ability to recover from stress or damage compared to highly vulnerable first year). Second year grafted rootstock trunk risk factors includes= severe winter damage (extreme cold during first or second winter potentially killing cambium especially if trunk protection inadequate), excessive crop load (overloading weak second year trunk with fruit potentially stressing vine and slowing trunk development), inadequate support (stake failure or inadequate staking leading to wind damage or graft union stress), crown gall development (existing infections potentially expanding during second year requiring intervention), and equipment damage (careless vineyard work injuring developing trunk or graft union). Second year grafted rootstock trunk transition indicators includes= approaching 3-4 cm diameter (moving toward mature trunk size threshold), showing established bark patterns (developing characteristic rootstock cultivar bark texture), displaying stable graft union (callus formation largely complete with strong integration), demonstrating improved resilience (better tolerance to environmental stress and mechanical disturbance), and supporting vigorous scion growth (enabling healthy canopy development indicating good trunk function). Second year grafted rootstock trunk significance includes= representing critical consolidation phase (second year building on first year foundation strengthening graft union and trunk structure), requiring continued but reduced management (less intensive than first year but still needing protection and monitoring), establishing long-term patterns (second year growth and graft union quality predicting mature vine success), marking progress toward maturity (second year trunk transitioning from extremely vulnerable to moderately robust), and determining production timeline (successful second year development enabling potential third year first crop or confirming need for additional establishment time).
Second_Year_Grafted_Rootstock_Trunk emphasizes second growing season post-grafting and is distinguished from First_Year_Grafted_Rootstock_Trunk (initial establishment) and Establishing_Grafted_Rootstock_Trunk (approaching maturity) by focus on intermediate consolidation phase. Critical for understanding graft union maturation, continued rapid growth, transitional characteristics, reduced vulnerability, and progression toward established mature trunk status.
Consolidating grafted rootstock trunk
Konsolidierender Wurzelstockstamm
Second season grafted rootstock trunk
Year 2 post-graft rootstock trunk
Zweite Saison Pfropfstamm
Zweites Jahr verpfropfter Unterlagenstamm
Second Year Grafted Rootstock Trunk
Merlot second year grafted scion trunk on 5BB moderate rootstock in Bordeaux shows 19-26 mm scion diameter by November dormancy, two annual rings (year 1= 2.5-3.8 mm, year 2= 3.8-5.5 mm), graft union at 8 cm above soil with stable 110-116% swelling, rootstock diameter 22-30 mm below union showing proportional growth, bilateral cordons established at 70 cm height, 40-60 g combined starch reserves (scion 15-22 g, rootstock 25-38 g), and light crop of 6-10 clusters removed at fruit set to prevent stress
A second year grafted scion trunk is a young grafted scion trunk during its second growing season following successful first year establishment, characterized by accelerated scion growth, stabilized graft union, confirmed rootstock-scion compatibility, and substantial trunk diameter increase, representing the transition from vulnerable establishment to approaching productive architecture with rootstock vigor expression becoming evident. Second year grafted scion trunk development builds on established graft union= spring growth from expanded rootstock root system (40-80 cm depth, 40-60 cm radius providing enhanced water and nutrient supply), vigorous scion shoot growth utilizing combined year 1 reserves (utilizing 10-20 g starch accumulated in scion and rootstock), rapid scion trunk diameter increase (adding 3-7 mm radial growth producing year 2 scion annual ring, faster than own-rooted due to rootstock vigor in many combinations), training architecture establishment (cordon or head positions selected accounting for union height), and potential first light fruiting (0-12 clusters depending on rootstock vigor and management). Second year grafted scion trunk anatomical development includes= second scion annual ring formation (typically 2.5-6 mm radial increment, width reflecting rootstock vigor influence), scion bark thickening (increasing from 1-2 mm to 2-3 mm), graft union diameter ratio stabilizing (union swelling typically 105-120% of adjacent scion trunk achieving equilibrium), expanding scion sapwood cross-section (year 1 and year 2 rings both conducting water with rootstock-mediated hydraulic characteristics), and continued graft union vascular integration (xylem and phloem connectivity across union strengthening improving physiological continuity). Second year grafted scion trunk diameter development varies by rootstock= vigorous rootstocks (110R, 1103P, SO4) producing 20-35 mm scion diameter by second dormancy (compared to 15-30 mm on same cultivar own-rooted), moderate rootstocks (5BB, 420A, 5C) producing 16-28 mm diameter (similar to own-rooted), low-vigor rootstocks (3309C, 101-14Mgt, Riparia Gloire) producing 13-24 mm diameter (potentially less than vigorous own-rooted), with rootstock effects on scion diameter becoming increasingly evident in year 2. Second year grafted scion trunk graft union characteristics include= union diameter swelling stabilization (ratio between union and adjacent trunk typically 108-122%, excessive swelling >130% suggesting incompatibility), union bark continuity (periderm bridging union protecting graft interface), union mechanical strength (lignified callus providing structural integrity reducing breakage risk), union hydraulic function (water transport across union approaching non-grafted trunk conductance in compatible combinations), and union as permanent architectural feature (determining trunk architecture with union positioned 5-15 cm above soil). Second year grafted scion trunk management focuses on= architecture development accounting for union position (establishing cordons typically 55-75 cm above soil ensuring adequate clearance above union), continued rootstock sucker removal (removing rootstock shoots preventing rootstock canopy competition), crop load management (typically 0-12 clusters allowed, many viticulturists removing all fruit year 2 on grafted vines to strengthen union and trunk), training accounting for rootstock vigor (vigorous rootstock combinations may require more intensive shoot management, canopy control), and graft union protection (avoiding mechanical damage, monitoring for disease infection at union). Second year grafted scion trunk physiological characteristics include= rootstock-mediated vigor expression (shoot length, leaf size, growth rate reflecting rootstock influence more strongly than year 1), rootstock water relations effects (drought-tolerant rootstocks providing scion drought resistance, shallow-rooted rootstocks showing earlier water stress), rootstock nutrient acquisition patterns (lime-tolerant rootstocks preventing scion chlorosis on calcareous soils, efficient rootstocks enhancing scion nutrition), carbohydrate accumulation in scion and rootstock (typically 30-55 g combined by year 2 dormancy= 12-20 g scion, 18-35 g rootstock), and rootstock phenology effects (some rootstocks advancing or delaying scion budbreak, véraison by 3-7 days). Second year grafted scion trunk rootstock compatibility confirmation includes= compatible combinations showing balanced growth (scion vigor appropriate for rootstock, union swelling moderate 105-120%, uniform growth without deficiency symptoms), incompatible combinations manifesting problems (excessive union swelling >125%, poor scion growth, leaf chlorosis or necrosis, trunk diameter less than 70% expected, indicating need for replanting), and marginal compatibility showing intermediate symptoms (acceptable growth but sub-optimal, union swelling 120-130%, potential long-term productivity concerns).
Second_Year_Grafted_Scion_Trunk emphasizes accelerated second year with stabilized union and is distinguished from First_Year_Grafted_Scion_Trunk (union establishment phase) and Establishing_Grafted_Scion_Trunk (combined years 2-3 architecture phase) by confirmed compatibility and rootstock vigor expression. Critical for understanding rootstock effects on scion development, union stability confirmation, compatibility assessment timing, and accelerated or delayed development compared to own-rooted depending on rootstock selection.
Second season grafted trunk
Second year establishment grafted scion
Zweite Vegetationsperiode Aufpropfung
Zweites Jahr gepfropfter Stamm
Zweitjähriger Veredelungsstamm
Second Year Grafted Scion Trunk
Merlot second year own-rooted trunk in Bordeaux vineyard shows 18-24 mm diameter by November dormancy, two annual growth rings (year 1= 2.5-3.5 mm, year 2= 3.5-5.5 mm), cordon arms established at 70 cm height with 4-6 spurs positioned for year 3 fruiting, 35-50 g stored starch reserves, and light crop of 4-8 clusters (0.3-0.6 kg fruit) removed at véraison to prevent stress
A second year own-rooted trunk is a young own-rooted trunk during its second growing season following successful first year establishment, characterized by accelerated growth rates, substantial trunk diameter increase, development of training architecture, and potential for initial light fruiting, representing the transition from vulnerable establishment to approaching productive maturity. Second year own-rooted trunk development builds on first year foundation= spring growth begins from established root system (root zone expanded to 40-80 cm depth and 40-60 cm radius), vigorous shoot growth from increased carbohydrate reserves (utilizing 8-15 g starch accumulated during year 1), rapid trunk diameter increase (adding 3-6 mm radial growth producing year 2 annual ring), extension and training of permanent structure (cordon development or head training establishment), and potential first light crop (0-10 clusters per vine depending on vigor and management strategy). Second year own-rooted trunk anatomical development includes= second annual growth ring formation (typically 2.5-5 mm radial increment, wider than mature trunk rings), progressive bark thickening (increasing from 1-2 mm to 2-3 mm), expanding sapwood cross-section (year 1 and year 2 rings both conducting water), continued active cambial growth throughout growing season (April-October in Northern Hemisphere temperate climates), and minimal heartwood formation (central pith may begin showing discoloration but remains largely functional). Second year own-rooted trunk diameter development varies by cultivar vigor= high-vigor cultivars (Grenache, Carignan) reaching 18-30 mm diameter by second dormancy, moderate-vigor cultivars (Cabernet Sauvignon, Syrah) reaching 15-25 mm diameter, low-vigor cultivars (Pinot Noir, Gamay) reaching 12-22 mm diameter, with substantial variation depending on soil fertility, water availability, and climate. Second year own-rooted trunk management focuses on= architecture development (establishing permanent cordon positions typically 50-80 cm above soil, or head-trained positions), shoot selection and positioning (choosing shoots for permanent arms or cordons, removing others), crop load management (typically 0-10 clusters allowed, preventing overcropping that could stress developing vine), continued training to trellis system (tying shoots to establish desired architecture), and disease prevention particularly at pruning wounds (Eutypa dieback, Esca, Botryosphaeria can infect through pruning cuts in second year). Second year own-rooted trunk physiological characteristics include= increased photosynthetic capacity from larger leaf area (typically 1.5-3 m² total leaf area per vine), higher carbohydrate production and storage (accumulating 25-45 g starch by second dormancy), improved water transport capacity (year 1 and year 2 xylem providing 0.2-0.8 L/day capacity), potential for first reproductive development (flower initiation in buds for year 3 flowering), and establishment of shoot fruitfulness patterns. Second year own-rooted trunk stress vulnerability decreases= improved drought tolerance from expanded root system, greater frost tolerance from increased carbohydrate reserves, enhanced pest and disease resistance from more robust physiology, but continued vulnerability to severe environmental stress (extreme drought, heat waves, severe winter cold in marginal climates). Second year own-rooted trunk success indicators include= achieving 15-25 mm trunk diameter by dormancy (varies by cultivar), establishing desired architecture (cordon or head positions selected and positioned), accumulating adequate reserves (25-45 g starch supporting year 3 growth and cropping), successful overwintering (>95% survival in appropriate climates), and preparing for year 3 productivity (buds showing flower primordia indicating readiness for cropping).
Second_Year_Own_Rooted_Trunk emphasizes accelerated second season development and is distinguished from First_Year_Own_Rooted_Trunk (initial vulnerable establishment) and Establishing_Young_Own_Rooted_Trunk (combined years 2-3 architecture development) by transition from establishment to approaching productivity. Critical for understanding training timeline and architecture development, optimal cropping progression, reserve accumulation for sustained productivity, and reducing establishment failure risk.
Second season trunk
Second year establishment trunk
Zweiter Vegetationsjahr-Stamm
Zweites Jahr Stamm
Zweitjähriger Etablierungsstamm
Second Year Own Rooted Trunk
Secondary aroma description for Beaujolais Nouveau (Gamay, 2024 vintage, carbonic maceration) identifies intense banana and pear drops (from isoamyl acetate at 8-15 mg/L, produced at elevated levels during intracellular carbonic maceration), kirsch-like cherry (ethyl cinnamate enhanced by carbonic maceration), and bubble gum (ethyl butyrate) — the exuberant fermentative bouquet that characterizes nouveau-style wines where secondary aromas deliberately dominate primary varietal character. Secondary aroma description for barrel-fermented Chardonnay from Meursault (2023 vintage, 100% MLF) documents butter and cream (diacetyl at 3.5 mg/L from malolactic fermentation, above the 1.0 mg/L sensory threshold), brioche and toasted bread (yeast autolysis products from 10 months on lees in barrel), hazelnut (acetaldehyde-phenolic interaction), and a creamy-textural quality from mannoprotein release — the secondary aromatic transformation that distinguishes barrel-fermented, MLF-completed Burgundian Chardonnay from its reductively handled counterparts.
Secondary aroma description is an aroma description entity that documents the olfactory characteristics of wine attributable to volatile compounds generated during alcoholic fermentation by Saccharomyces cerevisiae and, where applicable, during malolactic fermentation by lactic acid bacteria, encompassing yeast-derived esters (fruity, floral), higher alcohols (vinous, solvent-like at excess), acetaldehyde (green apple, nutty), fermentation-derived sulfur compounds (reductive notes), diacetyl and related diketones from MLF (buttery, creamy), and the broader spectrum of yeast metabolic products that constitute the "fermentation bouquet" overlaying the primary varietal character.
Secondary aroma description captures the aromatic imprint of the winemaker's fermentation decisions — yeast strain selection, fermentation temperature, nutrient management, MLF choice, and lees contact duration all manifest as specific secondary aromatic signatures that interact with and modify the primary varietal character. The principal fermentation-derived aroma compounds include ethyl esters of fatty acids (ethyl hexanoate — apple/pear, ethyl octanoate — tropical fruit, ethyl decanoate — waxy), acetate esters (isoamyl acetate — banana, 2-phenylethyl acetate — rose/honey, ethyl acetate — fruity/solvent), higher alcohols (isoamyl alcohol — marzipan at moderate levels, 2-phenylethanol — rose), and sulfur compounds (hydrogen sulfide — rotten egg, mercaptans — onion/garlic at fault levels, but positive minerality contribution at trace levels). The Ehrlich pathway of amino acid catabolism — where branched-chain and aromatic amino acids are transaminated to α-keto acids, decarboxylated to aldehydes, and reduced to higher alcohols that subsequently esterify — represents the primary biochemical route from must composition to fermentation-derived aroma, making the amino acid profile documented in the Nitrogen_Data hierarchy a direct predictor of secondary aromatic potential. The MLF-derived secondary aromas are dominated by diacetyl (2,3-butanedione), whose perception threshold (1.0-4.0 mg/L depending on wine matrix) and degradation kinetics (progressive reduction to less odorous acetoin and 2,3-butanediol by lactic acid bacteria) make its contribution temporally dynamic — freshly completed MLF wines show maximum butteriness, which diminishes over weeks to months through enzymatic reduction. The balance between primary and secondary aromas varies by wine style — aromatic varieties (Riesling, Sauvignon Blanc) are typically vinified to preserve primary character by minimizing secondary aromatic overlay, while neutral varieties (Chardonnay, Sémillon) rely more heavily on secondary aromas for aromatic interest.
Gäraroma
Gärbukett
Hefearoma
Sekundäraroma
fermentation aroma
fermentative bouquet
fermentatives Bukett
secondary bouquet
vinification aroma
yeast-derived aroma
Ugliano M, Henschke PA (2009) Yeasts and wine flavour. In Wine Chemistry and Biochemistry, Springer, pp.: 313-392 Swiegers JH et al. (2005) Yeast and bacterial modulation of wine aroma and flavour. Australian Journal of Grape and Wine Research 11(2): 139-173 Hazelwood LA et al. (2008) The Ehrlich pathway for fusel alcohol production: a century of research on Saccharomyces cerevisiae metabolism. Applied and Environmental Microbiology 74(8): 2259-2266
Secondary Aroma Description
Secondary aroma precursor quality in Viognier must influences the fermentation production of 2-phenylethanol and 2-phenylethyl acetate that contribute to the variety's characteristic rose-floral character, with phenylalanine concentrations of 50-150 mg/L supporting 2-phenylethanol formation of 20-60 mg/L through Ehrlich pathway metabolism must supplementation with phenylalanine in nitrogen-limited vintages can enhance rose-floral character without the elevated volatile acidity risk of complete YAN supplementation with DAP, demonstrating targeted manipulation of secondary aroma precursor quality.
Secondary_Aroma_Precursor_Quality encompasses the chemical qualities characterizing grape-derived compounds that serve as precursors for fermentation-generated aroma compounds, capturing the substrate pool that yeast and bacterial metabolism transform into the esters, alcohols, aldehydes, and other volatiles comprising secondary (fermentation) aromas. This category recognizes that while secondary aromas arise from fermentation metabolism, the grape must composition profoundly influences the types and quantities of secondary aromas produced. Key precursor classes include amino acids (particularly branched-chain amino acids yielding higher alcohols and their esters), unsaturated fatty acids (yielding medium-chain fatty acids and ethyl esters), and various nitrogen-containing compounds affecting fermentation volatile production. The amino acid profile of must determines the formation of higher alcohols through the Ehrlich pathway, where amino acid catabolism generates fusel alcohols (isoamyl alcohol from leucine, isobutanol from valine, 2-phenylethanol from phenylalanine) that subsequently esterify to form fruity esters. Must lipid composition affects fatty acid-derived volatiles including ethyl octanoate, ethyl decanoate, and medium-chain fatty acids contributing to wine fruitiness and complexity. Yeast assimilable nitrogen (YAN) content affects both fermentation kinetics and the balance between amino acid-derived and sugar-derived volatiles, with nitrogen-limited fermentations producing different volatile profiles than nitrogen-sufficient fermentations. The secondary aroma precursor pool is partially under viticultural control through nitrogen fertilization, canopy management affecting fruit nitrogen status, and harvest timing affecting amino acid and lipid accumulation. Understanding precursor availability enables prediction and manipulation of fermentation aroma outcomes.
Secondary_Aroma_Precursor_Quality bridges the Primary_Aroma_Quality (grape-derived) and fermentation aroma chemistry by recognizing that must composition affects fermentation volatile production. The class is positioned within Aromatic_Compound_Quality to capture the precursor dimension of secondary aroma generation. Amino acids represent the most extensively studied precursor class, but fatty acids, sugars, and other must components also affect secondary volatile formation. The interaction between precursor availability and yeast strain genetics creates complexity in predicting fermentation aroma outcomes from precursor measurements alone.
Aminosäure-Vorläufergehalt
Fermentationsaroma-Substratqualität
Most-Vorläuferzusammensetzung
Sekundäraroma-Vorläuferqualität
amino acid precursor content
fermentation aroma substrate quality
fermentation volatile precursor pool
must precursor composition
Ugliano M and Henschke PA (2009) Yeasts and Wine Flavour. In: Moreno-Arribas MV and Polo MC (eds) Wine Chemistry and Biochemistry, Springer pp.: 313-392 Garde-Cerdán T and Ancín-Azpilicueta C (2008) Effect of the Addition of Different Quantities of Amino Acids to Nitrogen-Deficient Must on the Formation of Esters, Alcohols, and Acids during Wine Alcoholic Fermentation. LWT - Food Science and Technology 41(3): 501-510
Secondary Aroma Precursor Quality
A secondary basal leaf at node 2 on a secondary shoot of Vitis vinifera cv. Chardonnay in Chablis, France, developing from secondary bud activation following severe spring frost (-4°C) that killed primary buds on 70% of spurs, measuring 11.8 cm length by 10.5 cm width representing 35% size reduction compared to primary bud second leaves on unaffected shoots (18.2 cm × 16.1 cm), exhibiting somewhat simplified morphology with five lobes but shallower sinuses and less pronounced lobe development compared to typical Chardonnay leaf architecture, emerging 8 days later than primary bud break occurring on frost-protected shoots, achieving full expansion at 32 days post-secondary budbreak (compared to 26 days for primary leaves), demonstrating reduced photosynthesis rate of 9.8 μmol CO₂ m⁻² s⁻¹ representing 72% of primary leaf capacity, supporting slower shoot development with reduced vigor and absence of crop (secondary buds generally non-fruitful in Chardonnay), measured at nominal shoot BBCH stage 71 (fruit set timing on primary shoots) though secondary shoots lag 2-3 weeks in development, representing compensatory secondary bud growth providing vegetative function but reduced productivity following primary bud loss.
A second basal leaf at node 2 on shoots developing from secondary buds within compound grapevine buds in Vitis vinifera, characterized by development from less-advanced primordia within secondary bud components that activate following primary bud failure due to frost damage, winter injury, disease, or other primary bud mortality, smaller dimensions typically 20-40% reduced compared to equivalent primary bud leaves reflecting the secondary bud's reduced developmental priority during previous-season formation, morphological characteristics sometimes showing simplified features including reduced lobe development, shallower sinuses, or modified overall architecture compared to typical cultivar patterns expressed in primary bud leaves, delayed emergence timing typically 5-10 days later than primary bud break reflecting slower development from less-advanced preformed primordia, reduced photosynthetic capacity reflecting incomplete primordial development with 60-80% of primary bud leaf rates, and functional role providing necessary but suboptimal photosynthetic capacity on shoots developing after primary bud loss creating delayed and often reduced crop development. Secondary basal leaves represent developmental compensation following primary bud failure, emerging from compound bud backup systems that provide growth capacity when primary components are lost, developing from primordia that received secondary priority during previous-season formation resulting in less complete tissue differentiation and smaller cell populations compared to primary bud primordia, creating shoots that maintain vegetative growth potential but often show reduced vigor and fruitfulness compared to primary bud shoots.
Secondary basal leaves are distinguished from primary basal leaves by origin from secondary bud components, smaller size, sometimes simplified morphology, delayed development, and reduced function reflecting the secondary bud's backup status within compound bud architecture. Secondary buds develop during the same previous growing season as primary buds but receive secondary priority in resource allocation, resulting in less complete primordial development with fewer cells, reduced tissue differentiation, and smaller preformed leaf structures. Activation occurs primarily following primary bud failure from environmental stresses, with frost being the most common cause though winter desiccation, disease, or physical damage also trigger secondary bud development. Cultivar variation exists in secondary bud fruitfulness, with some varieties producing inflorescences from secondary buds while others show purely vegetative secondary shoots, affecting vineyard productivity following primary bud loss events. Management of vineyards with high secondary bud activation requires adjustment of expectations regarding crop level, fruit quality, and canopy development, with secondary shoots typically requiring different training, positioning, and pruning approaches compared to primary shoots.
Ersatzknospe Knoten-zwei Blatt
backup second leaf
compensatory basal leaf
kompensatorisches basales Weinblatt
secondary bud node two leaf
sekundäres basales Blatt
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 109-134 Carmona et al. (2008) "Grapevine bud dormancy= environmental and molecular aspects" Revista Brasileira de Fruticultura 30=257-408 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 353-376.
Secondary Basal Leaf
Cabernet Franc in Loire Valley shows secondary branch primordium formation during late August-early September Year N at BBCH 33-38 within developing latent buds over 2-3 weeks, with 20-30 primary branch primordia initiating 2-5 secondary branches each particularly on distal primary branches producing total flower primordium number of 400-800 per inflorescence before dormancy entry under conditions of 18-24°C declining temperature, adequate late-season moisture, maintained canopy function, and balanced fertility typical of this moderately-to-highly compound mid-season cultivar establishing complex architecture determining Year N+1 yield capacity
Secondary branch primordium formation represents the advanced architectural elaboration stage where primary branch primordia initiate second-order lateral branches creating higher-order compound inflorescence structure that substantially increases flower number capacity, establishing complete architectural framework that will determine ultimate cluster complexity and berry number potential. This final architectural developmental stage occurs within developing latent bud during late summer-early autumn Year N following or overlapping with primary branch formation, typically spanning 2-3 weeks as secondary branch primordia initiate on progressively maturing primary branches before dormancy entry arrests development. Secondary branch initiation proceeds on primary branch primordia beginning with earliest-formed apical branches and progressing basipetally as successive primary branches mature to appropriate developmental stage, creating temporal and spatial gradient of secondary branching development within elaborating inflorescence primordium. Branch initiation mechanism on primary branches resembles primary branch formation from inflorescence axis, with local cell division acceleration in peripheral zone of primary branch meristem creating meristematic protrusion 15-30 micrometers diameter that will develop into secondary branch bearing flower primordia. Secondary branch number per primary branch varies substantially with cultivar architecture and developmental conditions, ranging from 0-2 in simply branched varieties creating relatively simple loose clusters to 3-8 in highly compound varieties producing large dense clusters, with secondary branching representing major determinant of cluster complexity. Architectural diversity among cultivars reflects differences in secondary branching with some varieties showing minimal or no secondary branching producing primary-branch-only architecture, others showing moderate secondary branching on distal primary branches creating intermediate complexity, and highly compound types showing extensive secondary branching throughout inflorescence creating maximum complexity. Flower primordium initiation begins on branch tips during or following secondary branch formation, with individual flower primordia emerging as small meristematic protrusions 10-20 micrometers that will remain undifferentiated through dormancy and complete flower development during Year N+1, with flower number determined by total number of branch tips established during architectural elaboration. Total flower number potential per inflorescence results from multiplicative architecture with primary branch number multiplied by average secondary branch number per primary branch multiplied by average flower number per branch tip, typically yielding 100-400 flowers in small simple architectures to 500-1500 flowers in large compound structures. Developmental timing shows secondary branch formation occurring relatively late in primordium elaboration typically late August-September in temperate regions Year N, with timing constrained by need to complete before dormancy entry as declining temperatures and photoperiod trigger growth cessation. Completion urgency creates vulnerability with late-season stress or early dormancy induction potentially interrupting secondary branch formation before completion, reducing final branching extent and flower number affecting yield capacity. Inflorescence primordium size at completion of secondary branching reaches maximum dimensions of 600-1200 micrometers length and 300-600 micrometers width depending on cultivar architecture, representing fully elaborated structure containing complete branching framework and initial flower primordia though organs remain undifferentiated. Cell division activity declines progressively during secondary branch formation as developmental program approaches completion and dormancy preparation begins, with final cell divisions occurring in latest-forming branch primordia and flower initiation sites before complete arrest. Gene expression changes during late primordium elaboration show declining reproductive development genes as architectural program completes, increasing dormancy-related genes preparing for winter arrest including DORMANCY ASSOCIATED genes and cold acclimation genes, and maintained meristem identity genes preserving developmental potential through dormancy. Hormonal regulation during late elaboration shows declining growth-promoting hormones including auxin and cytokinin as activity reduces, increasing abscisic acid promoting dormancy preparation, and maintained low gibberellins preserving reproductive identity through dormancy. Metabolic transition during completion shows declining respiration from peak elaboration levels toward dormancy maintenance levels, reduced protein synthesis shifting from growth-related to storage and protective proteins, and beginning carbohydrate accumulation in bud scales and axis tissues supporting dormancy survival. Dormancy entry arrests development with primordium in state of complete architectural elaboration but incomplete organ differentiation, with branch framework established, flower primordia initiated but undifferentiated, and tissues dehydrating and accumulating protective compounds preparing for winter survival. Environmental conditions during secondary branch formation critically affect completion with warm late summer-early autumn supporting full elaboration, adequate late-season resources enabling complete branching, and absence of early stress or premature dormancy induction allowing developmental completion. Canopy management effects show importance of maintaining adequate late-season photosynthetic capacity with early defoliation or disease reducing resources impairing secondary branching completion, while well-maintained canopy supports full elaboration. Late-season stress effects can reduce secondary branching with severe water stress arresting development prematurely, excessive crop load diverting resources limiting branching, or disease reducing photosynthetic capacity impairing completion. Temperature effects show warm late summer promoting complete secondary branching while cool conditions may limit elaboration, and timing of autumn cooling relative to completion stage affecting whether full branching achieves before dormancy arrest. The process represents fourth and final stage of inflorescence primordium formation completing architectural elaboration before dormancy entry, with completed primordia representing Year N+1 yield potential established during Year N. Regional timing shows secondary branch primordium formation during late August-September in warm climates, September-early October in temperate regions, and late August-September in cool areas, requiring completion before dormancy entry timing which varies geographically. Viticultural significance includes representing final determinant of inflorescence architecture and flower number capacity, understanding that late-season conditions critically affect completion and subsequent yield, recognizing importance of maintaining vine health and resources through late season supporting elaboration, and appreciating that incomplete secondary branching permanently reduces yield potential. Research applications include analyzing regulation of higher-order branching and architectural complexity, investigating environmental and resource effects on late-season primordium elaboration, quantifying relationships between secondary branching and ultimate yield capacity, evaluating timing constraints and dormancy interactions affecting completion, and assessing genetic variation in inflorescence architecture affecting productivity and cluster characteristics across cultivars.
Secondary_Branch_Primordium_Formation is fourth and final Inflorescence_Primordium_Formation subphase, distinguished by second-order branching completing architectural elaboration. Follows or overlaps with Primary_Branch_Primordium_Formation. Represents final developmental stage before dormancy entry. Major determinant of inflorescence complexity and flower number. Vulnerable to late-season stress or premature dormancy induction. Completion critical for maximum yield potential.
Sekundärzweig-Primordienbildung
Zweitordnungsverzweigung
higher-order branching
höhere Verzweigungsordnung
second-order branch formation
secondary branch initiation
sekundäre Blütenstandsverzweigung
Secondary Branch Primordium Formation
A secondary branch rachis arising from a primary branch on a primary inflorescence of Vitis vinifera cv. Syrah in the Rhône Valley, France, originating from the fourth primary branch at 2.8 cm from the primary branch base, representing third-order hierarchical position (following first-order main rachis and second-order primary branch), measuring 2.5 cm length representing moderate secondary branch development, diameter of 1.2 mm at junction with primary branch tapering to 0.6 mm at apex, exhibiting slender cylindrical construction, green coloration with sparse pubescence, containing 2 vascular bundles in cross-section providing limited vascular capacity for the branch system, bearing 4 tertiary branch rachides (fourth-order structures) and 8 direct-attached flowers, complete secondary branch system supporting approximately 18 flowers representing 2.6% of total inflorescence flower number, positioned as one of 7 secondary branches on this primary branch which itself bears approximately 42 total flowers across its secondary branch system, serving to create fine-scale flower distribution and three-dimensional inflorescence elaboration, measured at BBCH stage 63 (30% flowers open) in early June, representing typical secondary branch rachis architecture with third-order position and reduced dimensions.
A lateral branch axis arising from primary branch rachides representing third-order structures in the hierarchical inflorescence architecture in Vitis vinifera, characterized by origin from primary branch rachides through lateral branching from second-order structures, third-order hierarchical position in the rachis system, typical dimensions of 0.8-4 cm length and 0.8-2.0 mm diameter representing further reduction from primary branches, limited vascular capacity with 1-4 vascular bundles, bearing tertiary branch rachides (fourth-order) and flowers creating additional architectural elaboration, variable occurrence with 3-12 secondary branches per primary branch depending on primary branch size and inflorescence complexity, and functional roles including further distributing flowers throughout inflorescence volume, extending vascular networks to peripheral regions, and creating the fine-scale branching architecture characteristic of elaborate paniculate inflorescences. Secondary branch rachides represent the second lateral branching order, developing through lateral meristem activity along developing primary branches, exhibiting reduced dimensions reflecting the third-order hierarchical position and resource allocation patterns, and creating the intermediate-scale branching defining inflorescence complexity.
Secondary branch rachides are distinguished from primary branch rachides by origin from primaries rather than main rachis and third-order versus second-order position. The secondary branch designation emphasizes the second-order lateral status representing the second level of branching from the main axis. Third-order position means these structures originate from second-order primary branches, contrasting with fourth-order tertiary branches originating from secondary branches. Number per primary branch shows variation from few (2-4) on small simple primaries to many (10-15+) on large elaborate primaries, creating the range in inflorescence complexity. Distribution along primary branches shows relatively regular spacing with typical intervals of 0.4-0.8 cm. Size shows substantial reduction from primary branches with diameter ratios typically 0.4-0.7× primary diameter, reflecting hierarchical subordination and resource allocation patterns. Vascular capacity shows proportional reduction with 1-4 bundles compared to 3-7 in primaries. Branching complexity varies from simple structures bearing only flowers to elaborate branches with multiple orders of tertiary and quaternary elaboration. Length variation reflects position on primary branch with basal secondaries typically largest, mid secondaries intermediate, and apical secondaries smallest following the common basipetal size gradient. Developmental origin involves lateral meristem initiation along developing primary branches with sequential appearance. Flower bearing occurs both directly on secondary branches and indirectly through tertiary and higher-order branches, creating distributed flower arrays. Transformation to cluster maintains secondary branch identity with these structures becoming the fine-scale berry-bearing branches creating cluster texture and compactness.
dritter Ordnung Spindel
second lateral branch
secondary lateral axis
sekundäre Zweigrachis
third-order rachis
zweite laterale Achse
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 333-368 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 399-434 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 599-638.
Secondary Branch Rachis
A secondary bud at node 5 on Vitis vinifera cv. Cabernet Sauvignon examined following 65% primary bud mortality from spring frost at -3.5°C, dissection revealing one small inflorescence primordium containing 45 flower initials plus vegetative shoot primordia with 7 leaf initials, this secondary bud breaking 10 days after the frost event when primary bud death was confirmed, developing into a shoot producing one small cluster of 18 berries (40% fruit set from the reduced flower number) plus 11 leaves, partially compensating for primary bud loss but achieving only 38% of normal node yield.
The second hierarchical bud position within the compound bud structure of Vitis vinifera, located laterally adjacent to the primary bud within the same protective bud scale envelope, characterized by smaller size than primary buds (typically 1.5-2.5mm diameter versus 2.5-4mm for primaries), later differentiation during previous season development, reduced fruitfulness with fewer or smaller inflorescence primordia, and serving as backup growing point that activates when primary buds are damaged by frost, hail, disease, or mechanical injury. Secondary buds represent an insurance mechanism providing developmental redundancy in the compound bud architecture, capable of producing shoots and contributing to crop production when primary buds fail, though typically with 30-60% reduced yield potential, 7-14 day delayed phenology, and predominantly vegetative rather than reproductive primordial content, making secondary bud breakage a partial rather than complete compensation for primary bud loss.
Secondary buds are critical for vineyard recovery following spring frost damage, the most economically devastating weather event in viticulture. Their activation provides partial yield salvage and maintains canopy development for photosynthetic capacity and reserve accumulation, though rarely achieving normal production levels. The extent of secondary bud compensation varies by cultivar, with some varieties maintaining reasonable fruitfulness in secondary positions (Chardonnay, Riesling showing 40-60% of primary yields) while others produce predominantly vegetative secondary shoots (Cabernet Sauvignon, Merlot showing 20-35% of primary yields). Understanding secondary bud potential is essential for post-frost damage assessment and economic decision-making regarding crop insurance claims and yield expectations.
Nebenknospe
Sekundärknospe
backup bud
lateral bud
secondary position bud
seitliche Knospe
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Molitor et al. (2014) "Late frost damage risk for viticulture under future climate conditions= a case study for the Luxembourgish winegrowing region" Australian Journal of Grape and Wine Research 20=160-168 Trought & Bramley (2011) "Vineyard variability in Marlborough, New Zealand= characterising spatial and temporal changes in fruit composition and juice quality in the vineyard" Australian Journal of Grape and Wine Research 17=79-89.
Secondary Bud
Mechanistic investigation of Secondary_Bud_Activation_Disposition at Michigan State University employed controlled primary bud removal and hormonal analysis to characterize activation dynamics across 15 cultivars. Dormant buds were surgically removed to simulate primary bud kill at the delayed dormant stage, and secondary bud development was tracked at 2-day intervals. Parallel hormone analysis (auxin, cytokinin, ABA) in secondary bud tissues characterized the hormonal signals controlling activation. Secondary_Bud_Activation_Disposition varied substantially among cultivars. 'Merlot' showed rapid activation with secondary buds visibly swelling within 4 days of primary bud removal, green tissue emergence by day 8, and full shoot development by day 14. Hormonal analysis revealed rapid cytokinin increase (3.5-fold by day 4) and auxin decline in 'Merlot' secondary buds, consistent with release from correlative inhibition imposed by the dominant primary bud. 'Riesling' displayed slower Secondary_Bud_Activation_Disposition with swelling at day 8, emergence at day 14, and full development at day 22, accompanied by more gradual hormonal shifts. 'Chardonnay' showed the slowest activation (emergence at day 18) combined with low secondary shoot vigor (40% of 'Merlot' shoot length at 30 days). Field trials following natural primary bud kill confirmed laboratory rankings, with rapid-activating cultivars establishing functional canopy 10-14 days earlier than slow activators, translating to improved fruit maturity at harvest (1.5-2.5° Brix advantage) despite equivalent primary bud damage levels.
A Secondary_Bud_Activation_Disposition is a Cold_Recovery_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to rapidly activate secondary (and tertiary) buds within compound buds following death of primary buds from winter cold or spring frost, enabling shoot production and partial crop recovery from backup meristems. This disposition represents the first line of cold damage recovery, utilizing the developmental redundancy built into the compound bud structure to replace killed primary growing points with surviving backup buds. Secondary_Bud_Activation_Disposition encompasses both the speed of secondary bud activation following primary bud death (days from damage to visible growth) and the subsequent performance of secondary shoots (vigor, fruitfulness, maturation timing). Rapid activation minimizes the delay in canopy establishment and season compression that follow primary bud loss, while secondary shoot fruitfulness determines the proportion of crop recoverable through this pathway. Cultivars differ substantially in secondary bud activation dynamics and secondary shoot performance, with some producing vigorous, relatively fruitful secondary growth within days of primary shoot death while others show delayed activation and minimal fruitfulness. Strong Secondary_Bud_Activation_Disposition is particularly valuable when primary bud damage is extensive but cane vascular tissues remain functional, as the surviving vascular system can support rapid secondary bud development if activation capacity is present.
Secondary_Bud_Activation_Disposition is_a Cold_Recovery_Disposition. Secondary bud activation exploits the compound bud's developmental redundancy for damage recovery. Activation speed and secondary shoot quality both contribute to effective recovery. Activation is controlled by hormonal signals (cytokinin/auxin balance) released from correlative inhibition when primary bud is killed. Secondary shoot fruitfulness (determined by cluster primordia in secondary bud at dormancy) is a separate trait from activation speed, with both contributing to recovery effectiveness.
Ersatzknospen-Mobilisierung
Reserveknospen-Aktivierung
Sekundärknospen-Aktivierungs-Disposition
backup bud mobilization
compound bud redundancy utilization
secondary bud response
Dry PR (2000) Canopy management for fruitfulness, Australian Journal of Grape and Wine Research 6:109-115 Howell GS (2001) Sustainable grape productivity and the growth-yield relationship, American Journal of Enology and Viticulture 52:165-174 Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235
Secondary Bud Activation Disposition
Secondary bud activation quality comparison across wine grape cultivars established the crop recovery potential available following primary bud freeze damage under cold-climate conditions. Following controlled primary bud removal simulating complete primary bud freeze kill, secondary and tertiary bud activation was monitored and resulting shoot fruitfulness quantified across 12 cultivars. Riesling demonstrated excellent secondary bud activation quality with 95% of secondary buds activating within 14 days of primary bud removal, secondary shoots producing an average of 1.4 clusters versus 2.1 for primary shoots (67% relative fruitfulness), and cluster weight 85% of primary shoot clusters, resulting in 57% crop recovery potential from secondary shoots alone. Chardonnay showed moderate secondary bud activation quality with 88% activation rate, secondary fruitfulness of 0.8 clusters versus 1.9 primary (42% relative), and cluster weight 75% of primary, yielding 32% recovery potential. Gewürztraminer exhibited poor secondary bud activation quality with only 70% activation rate, secondary fruitfulness of 0.4 clusters versus 1.6 primary (25% relative), and reduced cluster weight, providing only 18% recovery potential. The secondary bud activation quality assessment identified Riesling as particularly suitable for cold-climate production where periodic freeze damage is expected, since its strong secondary fruitfulness enables substantial crop recovery that would not be possible with low-secondary-fruitfulness cultivars, offsetting Riesling's only moderate maximum hardiness with superior damage recovery capacity.
Secondary bud activation quality is a cold recovery quality that inheres in a grapevine cultivar or clone and represents the capacity for secondary and tertiary buds within compound buds to resume development and produce functional shoots when primary buds are killed by freezing injury, with the reliability, timing, and fruitfulness of secondary shoot development determining the extent to which crop potential can be recovered following primary bud losses. This quality addresses the critical backup mechanism that protects grape production against bud-killing freeze events, recognizing that the compound bud structure with its hierarchy of primary, secondary, and tertiary primordia provides an inherent insurance system where progressively more cold-hardy backup buds can compensate for losses of the more productive but less hardy primary buds. The ontological significance of secondary bud activation quality lies in its direct relationship to crop recovery, with reliable activation of fruitful secondary buds enabling substantial yield salvage following freeze events that would otherwise cause complete crop loss, and with cultivar variation in secondary bud fruitfulness creating significant differences in recovery potential that affect the economics of cold-climate grape production. The biological basis of secondary bud activation encompasses the release of correlative inhibition that normally prevents secondary bud development when primary buds are growing, the mobilization of carbohydrate reserves to support secondary shoot growth, and the developmental completion of secondary bud inflorescence primordia that determines the cluster number and size that secondary shoots can produce. The fruitfulness of secondary shoots varies substantially among cultivars, ranging from cultivars where secondary shoots produce clusters nearly equivalent to primary shoots through intermediate types with reduced but significant secondary fruitfulness to cultivars where secondary shoots produce only minimal or no fruit, creating the critical cultivar differences in crop recovery potential. The timing of primary bud loss affects secondary bud activation quality, with freeze events occurring during deep dormancy potentially affecting secondary bud survival as well, while spring frost damage to already-emerged primary shoots may permit more reliable secondary and tertiary bud activation from the still-dormant reserves within the compound bud.
Secondary_Bud_Activation_Quality is_a Cold_Recovery_Quality, representing the capacity for backup bud development to compensate for primary bud freeze losses. This class addresses the critical crop recovery mechanism following bud-killing freeze events. Secondary fruitfulness relative to primary (67% Riesling vs. 25% Gewürztraminer) determines recovery potential. The 3-4°C greater hardiness of secondary buds provides survival advantage in marginal freeze events. Cultivar variation in secondary fruitfulness should inform variety selection for cold-climate sites. Timing of freeze affects whether secondary buds remain available for activation.
Ersatzknospen-Fruchtbarkeit
Knospen-Kompensationskapazität
Sekundärknospen-Aktivierung
Sekundärtrieb-Entwicklung
alternative bud activation
backup bud fruitfulness
bud compensation capacity
Pool RM et al. (1990) Assessing and predicting cold injury to grapevines. Proceedings of the New York Wine Industry Workshop, pp. 17-25 Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection. Michigan State University Extension Bulletin E2930 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 508-545
Secondary Bud Activation Quality
Investigation of Secondary_Bud_Hardiness_Disposition and its interaction with secondary shoot fruitfulness at the University of Minnesota evaluated both hardiness parameters and crop recovery potential across 20 cultivars following controlled winter injury treatments. Secondary bud LT50 at maximum hardiness ranged from -22.4°C ('Merlot') to -39.5°C ('Frontenac'), with the secondary-primary hardiness differential (SPHD) ranging from 2.2°C to 5.3°C among cultivars. Following experimental exposure to temperatures killing >90% of primary buds, secondary shoot fruitfulness varied from 0.15 clusters/shoot ('Foch', minimal recovery) to 1.45 clusters/shoot ('Frontenac', excellent recovery). The product of secondary bud survival probability and secondary shoot fruitfulness defined a "crop recovery index" that varied 8-fold among cultivars. 'Marquette' exhibited an optimal combination of strong Secondary_Bud_Hardiness_Disposition (LT50 -37.8°C, SPHD 3.8°C) and good secondary fruitfulness (1.1 clusters/shoot), achieving 65% crop recovery following simulated severe winter injury. Conversely, 'St. Croix' showed excellent secondary hardiness (LT50 -38.2°C) but poor secondary fruitfulness (0.35 clusters/shoot), limiting effective recovery to 25% of potential crop. These findings demonstrate that Secondary_Bud_Hardiness_Disposition must be evaluated alongside secondary shoot fruitfulness to assess true crop security in cold climates.
A Secondary_Bud_Hardiness_Disposition is a Bud_Cold_Hardiness_Disposition that inheres specifically in the secondary (lateral) bud primordium within the compound grapevine bud and is realized through the supercooling capacity and freeze avoidance mechanisms of this smaller, less differentiated bud component that serves as a backup growth point when the primary bud is killed. This disposition represents the cold tolerance of the bud structure that produces secondary shoots following primary bud damage, typically bearing 30-70% of the cluster number and reduced cluster weight compared to primary shoots but providing meaningful crop recovery capacity. The secondary bud develops in the axil of the first leaf primordium of the primary bud, remaining smaller and less differentiated than the primary bud throughout dormancy, characteristics that contribute to its enhanced cold hardiness. Secondary buds typically achieve LT50 values 2-4°C colder than primary buds within the same cultivar, representing a significant survival advantage during intermediate freeze events that kill primary but spare secondary buds. The fruitfulness of secondary shoots varies substantially among cultivars, with some cultivars producing reasonably fruitful secondary growth (0.8-1.5 clusters per shoot) while others produce primarily vegetative secondary shoots (0-0.3 clusters per shoot). Cultivars combining enhanced Secondary_Bud_Hardiness_Disposition with high secondary shoot fruitfulness provide superior crop recovery potential following winter damage, an important trait combination for cold-climate viticulture.
Secondary_Bud_Hardiness_Disposition is_a Bud_Cold_Hardiness_Disposition. Secondary buds are more cold hardy than primary buds but produce less fruitful shoots, serving as backup for crop recovery following primary bud damage. The economic value of secondary bud survival depends on secondary shoot fruitfulness, which varies among cultivars. The secondary-primary hardiness differential (typically 2-4°C) defines a temperature window where primary buds are killed but secondary buds survive, enabling partial crop recovery. Breeding for cold-climate adaptation should consider both secondary bud hardiness and secondary shoot fruitfulness.
Lateralknospen-Kältetoleranz
Seitenknospen-Frosthärte
Sekundärknospen-Härte-Disposition
backup bud LT50
lateral bud hardiness
secondary bud freeze tolerance
Howell GS (2001) Sustainable grape productivity and the growth-yield relationship: a review, American Journal of Enology and Viticulture 52:165-174 Dami IE et al. (2012) Evaluation of winter injury and subsequent performance of grapevine cultivars in Ohio, Acta Horticulturae 932:235-241 Pool RM et al. (1990) Environmental factors controlling cold hardiness and dormancy in grapevines, VITIS Journal of Grapevine Research 29:129-144
Secondary Bud Hardiness Disposition
Secondary bud viability assessment following differential cold injury in Chardonnay reveals the backup system function enabling partial recovery, with vineyard damage pattern showing 45% primary viability and 88% secondary viability (secondary buds surviving due to greater hardiness), enabling yield projection of approximately 65% of normal (45% from undamaged primary buds at full fruitfulness plus 43% of positions relying on secondary buds at 50% fruitfulness = 45 + 21.5 = 66.5% of normal potential), with this assessment informing crop insurance claims, buyer expectations, and management decisions for the partially damaged crop.
Secondary bud viability quality is a bud viability quality that represents the living status of the secondary (lateral) bud within the compound bud structure, characterizing the survival of the backup bud that can produce a shoot if the primary bud is killed, though with reduced fruitfulness that partially compensates for primary bud loss but cannot fully replace the yield potential of viable primary buds. Within the BFO framework, this quality specifically addresses the secondary bud dimension of viability, distinguishing this backup bud survival from primary or tertiary survival while recognizing that secondary bud viability provides the recovery potential when primary buds are lost to cold injury, disease, or physical damage, enabling continued vine productivity albeit at reduced levels. The ontological significance lies in secondary bud viability representing the first line of backup when primary buds fail, where viable secondary buds can produce shoots that contribute to vine productivity and maintain canopy cover even when primary buds are killed, with the reduced fruitfulness of secondary shoots (typically 30-50% of primary bud cluster weight) providing partial yield recovery that may be economically significant even though it falls short of undamaged potential. The enhanced cold hardiness of secondary buds relative to primary buds (typically 2-4°C greater hardiness) creates the survival differential that enables secondary bud survival when primary buds are killed, with this differential providing the "insurance" function of the compound bud structure that evolved to maintain reproductive potential despite bud damage from cold, herbivory, or other hazards. The management response to secondary bud reliance includes leaving additional bud numbers (compensating for reduced per-shoot fruitfulness with increased shoot numbers), adjusting crop level expectations (recognizing the yield ceiling imposed by secondary bud fruitfulness), and canopy management modifications (accommodating potentially different shoot characteristics from secondary versus primary origin). Understanding secondary bud viability quality enables assessment of recovery potential following primary bud damage, realistic yield expectations when secondary buds must contribute substantially, and management planning that accommodates reliance on less fruitful backup buds.
Secondary_Bud_Viability_Quality addresses the survival of backup buds providing recovery potential when primaries fail. This quality enables partial yield compensation through less fruitful but more hardy secondary buds. The class supports damage recovery assessment through secondary bud viability characterization.
Ersatzknospenüberleben
Lateralknospenvitalität
Sekundärknospenvitalität
backup bud survival
lateral bud viability
secondary meristem viability
sekundäre Meristemlebensfähigkeit
subsidiary bud vitality
Howell GS (2000) Cold hardiness of grapevine buds and canes: research update. Proceedings of the ASEV 50th Anniversary Meeting, Seattle, Washington, pp. 187-192 Dry PR, Coombe BG (2004) Viticulture Volume 1: Resources, 2nd Ed. Winetitles, Adelaide, pp. 150-200
Secondary Bud Viability Quality
Syrah in Rhône Valley shows secondary cluster separation during late May Year N+1 at early BBCH 57 over 3-4 days in moderately compound architecture, with 2-4 secondary branches per distal primary branch elongating to 5-8 mm and spreading 35-45 degrees creating 1.5-2.5 mm spacing establishing sub-cluster organization under Mediterranean conditions of 18-22°C temperature, typical of this moderately-to-highly compound mid-to-late season cultivar producing 400-700 flowers per inflorescence through hierarchical branching
Secondary cluster separation represents the second-order spatial organization phase in compound inflorescence architectures where secondary branches differentiate from primary branches through elongation and positioning, establishing hierarchical fine-scale structure creating sub-clusters within primary branch units. This advanced organizational stage occurs during mid-late May in temperate regions Year N+1 following or overlapping with primary cluster separation, typically requiring 2-4 days as secondary branches achieve sufficient development to create perceptible separation from primary branch axis and neighboring secondary branches. Separation mechanism involves secondary branch elongation extending branches 4-10 mm from primary branch attachment creating spatial projection, angular positioning with branches spreading 30-50 degrees from primary branch axis establishing sub-cluster distribution, and localized primary branch elongation between secondary branch attachments creating 1-3 mm spacing along primary branch. Secondary branch presence and development varies substantially among cultivars with highly compound architectures showing 3-8 secondary branches per primary branch creating complex hierarchical structure, moderately compound types showing 1-3 secondary branches on distal primary branches creating intermediate complexity, and simply branched varieties showing absent or minimal secondary branching producing relatively simple architecture. Architectural complexity created by secondary separation substantially increases flower number and cluster density with highly compound types producing 500-1500 flowers per inflorescence through extensive secondary branching, compared to 100-400 flowers in simply branched varieties lacking secondary elaboration. Flower group organization within primary clusters shows subdivision through secondary branching with each secondary branch unit containing group of flowers at distal positions, creating hierarchical nested structure characteristic of compound grapevine inflorescence architecture. Vascular organization in secondary branches establishes independent transport pathways branching from primary branch vascular system, enabling localized resource delivery to flower groups on secondary branches supporting coordinated development within sub-clusters. Secondary branch elongation mechanism resembles primary branch elongation with cell division in intercalary zones adding cells and cell expansion throughout tissues increasing dimensions, though secondary branches typically achieve smaller final dimensions reflecting later initiation and shorter development period. Mechanical properties in secondary branches show flexible herbaceous structure adequate for supporting flowers and small developing berries, with progressive strengthening as tissues mature though secondary branches often represent weakest structural links in mature clusters potentially creating shattering locations during harvest. Flower development on secondary branches continues during separation with buds enlarging preparing for individual flower separation and anthesis, with flowers on secondary branches typically synchronized with flowers on primary branch supporting coordinated flowering within entire primary cluster unit. Metabolic activity in developing secondary branches shows elevated respiration supporting growth, though total metabolic demand is lower than primary structures reflecting smaller size. Developmental timing shows secondary separation occurring later than primary separation reflecting hierarchical development sequence, with delay typically 2-3 days between primary and secondary separation initiation. Within-cluster architectural pattern shows spatial distribution of secondary branches with distal primary branch regions often showing more extensive secondary branching while basal regions show reduced or absent secondary branches, creating compound-to-simple gradient along primary branch length. Genetic control of secondary branching represents major determinant of cluster architecture with cultivar-specific programs establishing branching extent, with mutations or environmental effects affecting secondary branching substantially altering cluster structure and berry number. Environmental effects on secondary separation include temperature affecting elongation rate with optimal conditions supporting full secondary branch development while suboptimal conditions may limit elongation or reduce secondary branch expression. Resource availability affects secondary branching with adequate resources supporting full architectural elaboration while limitation may reduce secondary branch development creating simpler architecture than genetic potential. Previous season conditions during primordium formation fundamentally determine secondary branching potential established in bud, with current season affecting expression of established potential rather than creating new branches. The process represents second of four flower cluster separation aspects creating hierarchical fine-scale organization in compound architectures, following primary separation and preceding final spacing and individual flower separation. Duration typically spans 2-4 days depending on cultivar branching characteristics and temperature, with warm conditions promoting rapid completion. Cultivar variation represents most prominent feature with highly compound varieties showing extensive secondary separation creating 6-12 mm branch extensions with numerous secondary branches, moderately compound types showing selective secondary branching on 30-50% primary branches, and simply branched varieties showing minimal to absent secondary branching maintaining simple architecture. Regional timing shows secondary cluster separation during mid-late May in warm climates, late May-early June in temperate regions, and early June in cool areas, occurring late in inflorescence emergence approaching flowering. Viticultural significance includes establishing fine-scale cluster architecture affecting berry number and density, influencing cluster microenvironment with highly compound types showing dense architecture potentially increasing disease pressure, affecting mechanical properties with secondary branches representing potential weak points in mature clusters, and expressing architectural diversity among cultivars affecting management requirements and quality potential. Research applications include analyzing genetic regulation of hierarchical branching determining architectural complexity, investigating developmental coordination between primary and secondary branching establishing nested structure, quantifying relationships between secondary branching and berry number, evaluating environmental effects on secondary branch expression affecting architectural plasticity, and assessing functional implications of architectural diversity for disease susceptibility and fruit quality.
Secondary_Cluster_Separation is second of four Flower_Cluster_Separation subphases, distinguished by second-order spatial organization through secondary branch differentiation. Occurs only in compound architectures. Follows Primary_Cluster_Separation. Major determinant of cluster complexity and berry number. Highly variable among cultivars reflecting genetic architectural programs. Creates hierarchical nested structure characteristic of grapevine inflorescence.
Nebenzweigseparation
Sekundärcluster-Separation
Zweitordnungs-Blütentrennung
hierarchical branching
hierarchische Gescheinsgliederung
second-order cluster division
secondary branch separation
Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 345-350
Secondary Cluster Separation
Within principal growth stage 6 (flowering), the secondary growth stage codes discriminate the progressive course of anthesis with 10% resolution — BBCH 61 (secondary code 1, 10% of flowerhoods fallen), BBCH 63 (secondary code 3, 30% fallen), BBCH 65 (secondary code 5, full flowering at 50% fallen), BBCH 67 (secondary code 7, 70% fallen), and BBCH 69 (secondary code 9, end of flowering) — enabling observers to record flowering progress with sufficient precision for spray timing decisions where fungicide applications must target the BBCH 61-65 window for optimal Botrytis protection during the susceptible open-flower period.
A BBCH code component represented by the second digit (0-9) of the two-digit BBCH code, specifying the precise developmental sub-stage within a principal growth stage, providing the fine-resolution temporal discrimination that distinguishes progressive morphological changes as the grapevine advances through each major developmental phase.
Secondary_Growth_Stage_Code provides the within-stage resolution that transforms the 10-level principal stage classification into the approximately 47-level detailed staging system defined for Vitis. The numbering convention for secondary stages varies across principal stages: in stage 1 (leaf development) the secondary code directly counts unfolded leaves (11 = 1 leaf, 15 = 5 leaves), in stage 6 (flowering) it follows an approximately decile-based percentage scale (61 = 10%, 65 = 50%), and in stages 7 and 8 it marks qualitative morphological milestones (71 = fruit set, 75 = pea size, 81 = véraison onset). Not all secondary codes (0-9) are defined within each principal stage — some are reserved for potential future refinement. The secondary code combined with the principal code yields the complete two-digit Full_BBCH_Code.
BBCH secondary stage
BBCH-Sekundärstadium
Feinstadium-Kennung
Unterstadien-Code
micro-stage identifier
sub-stage code
Secondary Growth Stage Code
A secondary inflorescence at node 5 on a shoot of Vitis vinifera cv. Merlot in Pomerol, France, measuring 13.8 cm length representing typical intermediate dimensions, positioned opposite the fifth fully expanded leaf following the primary inflorescence at node 3, exhibiting compound paniculate architecture with central rachis bearing 3-4 orders of branching creating moderately elaborate structure, rachis diameter of 2.8 mm at base representing moderate construction, bearing approximately 420 individual flowers organized across the branching network, showing dimensions approximately 75% of the primary inflorescence length (18.5 cm) and 68% of primary flower number (620 flowers) on the same shoot, developmental stage at BBCH 61 (beginning of flowering, 10% flowers open) showing 2-3 day phenological delay compared to primary at stage 65 (full bloom), secondary inflorescence initiated during primordium formation in early to mid-June of previous season following primary initiation by approximately 5-7 days, expected to contribute approximately 35% of total shoot yield following fruit set, measured in early June during sequential flowering progression, representing typical secondary inflorescence characteristics with intermediate development supplementing primary production.
The second inflorescence in positional sequence on a grapevine shoot in Vitis vinifera, characterized by position at node 5 or occasionally node 6 following the primary inflorescence at node 3, intermediate dimensions typically 10-18 cm length representing reduction compared to primary inflorescence but substantial development, moderately complex branching architecture showing 3-5 orders of branching, moderate flower number typically 200-600 flowers depending on cultivar and development conditions, position opposite the fifth leaf following the standard inflorescence-leaf arrangement pattern, supplementary yield contribution typically providing 25-40% of total shoot fruit production, occurrence showing cultivar variation with highly fertile varieties consistently producing secondaries while less fertile cultivars may show absent or poorly developed secondary inflorescences, and developmental characteristics including later initiation during primordium formation and reduced resource allocation compared to primary inflorescences. Secondary inflorescences represent the second-order reproductive structures supplementing primary inflorescence yield, developing through similar morphogenetic processes as primaries but with delayed timing and reduced resource investment, exhibiting size and complexity intermediate between robust primaries and reduced tertiaries when present, and serving important yield roles particularly in moderate to high fertility cultivars where secondaries contribute substantially to total production.
Secondary inflorescences are distinguished from primary inflorescences by position at node 5 versus node 3 and typically smaller size and reduced complexity. The secondary designation emphasizes both positional sequence (second in order) and subordinate development compared to primaries. Position at node 5 represents the standard pattern following the two-node interval between successive inflorescences, though occasional variation to node 6 occurs particularly on vigorous shoots. Size reduction compared to primaries is typical with secondaries showing 60-80% of primary dimensions and flower numbers, though substantial variation exists with some cultivars showing minimal size differences while others show pronounced reduction. Occurrence varies dramatically among cultivars with high fertility varieties (Grenache, some Muscat types) consistently producing well-developed secondaries, moderate fertility cultivars (Merlot, Syrah) showing variable secondary development, and low fertility varieties (Pinot Noir under some conditions) occasionally lacking distinct secondaries. Developmental timing shows initiation following primaries during previous season's primordium formation, with delays of 3-7 days typical creating sequential development patterns. Resource allocation subordination reflects the positional sequence with primaries receiving preferential investment during both initiation and current-season development. Phenological delay of 2-4 days compared to primaries creates staggered flowering potentially extending pollination windows and fruit set periods. Yield contribution ranges from negligible in low-fertility cultivars to substantial (30-45%) in high-fertility varieties, making secondaries important production components requiring retention in many vineyard systems. Management decisions regarding secondary retention versus removal through cluster thinning depend on crop load targets, vine balance considerations, and fruit quality objectives.
Ergänzungsrispe
node 5 inflorescence
second inflorescence
sekundäre Infloreszenz
supplementary cluster
zweiter Blütenstand
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 179-218 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 143-182 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 313-358.
Secondary Inflorescence
In Cabernet Franc vineyards of Chinon (Loire Valley) at BBCH 55-57, primary inflorescence branches bear 3-5 secondary branches each averaging 1.5-3 cm length, with basal primary branches supporting 4-6 secondary branches of 2-4 cm while apical primaries bear only 1-2 secondaries of 0.8-1.5 cm, creating moderately loose clusters of 120-180 berries at maturity. Barbera in Piedmont exhibits extensive secondary branching (5-7 per primary branch, 2-3.5 cm length) resulting in large, loose clusters of 200-400 berries. Conversely, compact-clustered Müller-Thurgau shows minimal secondary branching (0-2 per primary, 0.5-1.2 cm) producing tight clusters highly susceptible to bunch rot.
A secondary inflorescence branch is an inflorescence branch that arises as a lateral appendage from a primary inflorescence branch rather than directly from the main rachis, representing second-order architectural elaboration that substantially increases total flower number and determines the degree of cluster subdivision and internal complexity. Secondary branches emerge from primary branches at intervals of 1.5-4 mm during inflorescence development within the dormant bud, with branch number per primary branch varying from 0-10 depending on cultivar architecture type (compact versus loose) and primary branch position (basal primaries bearing more secondaries than apical primaries). These branches exhibit intermediate dimensions between primary and tertiary branches, typically measuring 0.3-4 cm in length and 0.4-1.5 mm in diameter at the base, with pronounced variation correlating to the position and vigor of the parent primary branch. Secondary branch anatomy features a modest vascular bundle continuous with the primary branch vascular system, providing resource supply for the 3-30 flowers typically borne per secondary branch through direct attachment and tertiary branch subdivision. Branch distribution on primary branches follows the same acropetal developmental pattern seen in primary branch distribution on the main rachis, with basal secondary branches on a primary branch generally longer and more subdivided than apical secondaries on that same primary. Secondary branch number per primary branch serves as a critical determinant of cluster compactness, with cultivars bearing many secondary branches (5-8 per primary) producing loose clusters while those with few secondaries (1-3 per primary) form tight clusters. Each secondary branch typically bears 8-25 flowers distributed on its axis and any tertiary branches, contributing 25-40% of total inflorescence flower number across all secondary branches collectively. Secondary branch development becomes visible during BBCH 53-55 as small protuberances on elongating primary branches, with distinct separation and elongation occurring during BBCH 55-57. The degree of secondary branching shows substantial phenotypic plasticity in response to previous-season environmental conditions, particularly water availability and canopy light exposure during June-July inflorescence primordium differentiation, making secondary branch characteristics sensitive indicators of developmental history. In loose-clustered cultivars, well-developed secondary branches create the internal spatial heterogeneity that facilitates air circulation and reduces disease pressure, while in compact-clustered types, abbreviated or absent secondary branching contributes to tight berry packing and elevated Botrytis risk.
Secondary_Inflorescence_Branch represents second-order architectural elaboration and must be distinguished from Primary_Inflorescence_Branch (which arises directly from the main rachis) and Tertiary_Inflorescence_Branch (which arises from secondary branches). The hierarchical terminology follows standard botanical nomenclature for compound inflorescence description. Secondary branch number, length distribution, and subdivision into tertiary branches are critical components of ampelographic cluster architecture description and strongly predict final cluster compactness. Environmental modulation of secondary branch development during the previous season's bud differentiation provides a mechanism for inter-annual variation in cluster architecture even within a single cultivar and site. Sub-classification into Lower and Upper positions reflects significant morphological heterogeneity along the primary branch axis.
Seitenast erster Ordnung
Sekundärast der Infloreszenz
lateral branch of primary branch
second-order inflorescence branch
sekundärer Blütenstandsast
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 90-93 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 169-176 OIV (2009). Descriptor List for Grape Varieties and Vitis Species, OIV code 204.
Secondary Inflorescence Branch
A secondary lateral petiole on a moderately lobed mature leaf at node 8 of Vitis vinifera cv. Syrah in the Rhône Valley, France, appearing as a 0.6 cm length slight extension at the junction between main vein and lateral lobe vein, diameter of approximately 1.0 mm barely distinguishable from adjacent vein thickness, light green coloration matching surrounding tissue, very sparse pubescence, containing single minimal vascular strand barely distinguishable from major vein, providing no measurable mechanical support beyond what the vein itself supplies, occurring inconsistently with approximately 8% of leaves showing these structures while 92% show simple vein divergence without petiolar character, showing no correlation with leaf function, vine vigor, or any other measurable parameters, measured at BBCH stage 81 (beginning of berry ripening) in early August, representing minimal vestigial lateral petiole expression barely distinguishable from normal vein morphology.
A lateral petiole representing secondary-order or vestigial development with minimal structural elaboration in Vitis vinifera, characterized by very small dimensions typically 0.3-1.0 cm length representing rudimentary petiolar structures, simple unbranched morphology appearing as slight stalk-like extensions at vein junctions, minimal anatomical elaboration with simple vascular strands and limited mechanical tissues, inconsistent occurrence appearing occasionally in some leaves but absent in most, negligible functional contribution providing essentially no meaningful mechanical support or vascular enhancement beyond what adjacent veins supply, and status as developmental curiosities rather than functionally significant structures or reliable morphological characters. Secondary lateral petioles represent minimal expression of lateral petiolar development, occurring most commonly as subtle swellings or slight extensions at major vein junctions in leaves showing moderate to deep sinuses, essentially functioning as transitional structures between clear petiolar character and simple vein divergence, and serving primarily to illustrate developmental flexibility in grapevine leaf architecture rather than contributing to functional capacity or providing ampelographic utility.
Secondary lateral petioles represent the minimal extreme of lateral petiolar expression in Vitis vinifera, distinguished from more developed lateral petioles by very small dimensions and minimal elaboration, and barely distinguishable from normal major veins. These structures blur the boundary between petiolar and vein categories, representing subtle variations in tissue development at vein junctions that occasionally produce slight petiolar character without clear functional significance. Occurrence is inconsistent and unpredictable, appearing occasionally in individuals or specific leaves without clear patterns, making them unreliable for any diagnostic or functional purposes. Anatomical examination shows minimal differentiation from adjacent vein tissue, with slight swelling or extension being the primary distinguishing feature. Functional contribution is negligible to nonexistent, with mechanical support and vascular capacity essentially identical to leaves lacking these structures. Classification as petiolar structures is somewhat arbitrary, with these features representing minor developmental variations within normal leaf architecture rather than distinct morphological categories. Practical importance is essentially zero, with these structures noted occasionally in detailed morphological descriptions but serving no role in cultivar identification, functional assessment, or vineyard management.
minimal lateral petiole
minimaler lateraler Blattstiel
rudimentary secondary petiole
rudimentärer sekundärer Stängel
vestigialer Petiolus
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 58-89 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 33-55.
Secondary Lateral Petiole
A secondary lateral vein on a mature leaf at node 7 of Vitis vinifera cv. Syrah in the Rhône Valley, France, branching from a primary lateral vein at 2.8 cm from the primary vein's junction with median vein, branching angle of 68 degrees from primary vein axis, extending 2.4 cm through blade tissue toward the lobe margin, diameter of 0.6 mm at junction with primary vein tapering to 0.3 mm distally, moderately prominent as slight raised ridge on abaxial surface approximately 0.1 mm above surrounding tissue, barely visible as faint line on adaxial surface, light green coloration, sparse pubescence, containing 3 xylem vessels (20-35 μm diameter) providing water transport to blade subsector representing approximately 2-3% of total leaf area, moderate phloem facilitating local photosynthate collection, limited mechanical tissues providing local support, serving as organizing axis from which 6-8 tertiary veins branch at intervals of 0.3-0.4 cm, tertiary veins subsequently branching into higher orders and veinlets completing reticulate network in the local blade region, creating vascular density of approximately 8-10 mm of secondary vein length per cm² of blade area, measured at BBCH stage 81 (beginning of berry ripening) in early August, representing typical second-order venation creating intermediate hierarchical network level.
A lateral vein representing second-order branches originating from primary lateral veins in the hierarchical venation system in Vitis vinifera, characterized by branching origin from primary lateral veins rather than directly from median vein, intermediate diameter typically 0.4-0.8 mm representing substantial reduction from primary vein size, second hierarchical position in lateral vein system serving as organizing axes for tertiary and higher-order veins, moderate vascular capacity with 2-4 xylem vessels supporting water delivery to blade subsectors, moderate mechanical strength providing local structural support, typical number of 10-20 secondary veins branching from each primary lateral vein creating extensive distribution network, branching angles from primary veins typically 50-80 degrees, extension through blade tissue for 1-4 cm creating intermediate-scale vascular territories, and functional roles including distributing water and nutrients from primary veins to local blade regions, collecting photosynthates from mesophyll tissues for export through primary veins, providing distributed mechanical support maintaining blade flatness, and organizing tertiary vein networks that complete the reticulate pattern reaching all mesophyll regions. Secondary lateral veins create the intermediate hierarchical level connecting large-scale primary vein architecture to fine-scale veinlet networks, developing following primary vein establishment during leaf expansion, exhibiting regular spacing and branching patterns creating relatively uniform vascular distribution throughout blade regions, and contributing to overall venation density that affects hydraulic efficiency, photosynthetic capacity, and mechanical properties.
Secondary lateral veins are distinguished from primary lateral veins by branching origin from primaries rather than median vein and smaller diameter, and from tertiary veins by second-order hierarchical position and intermediate size. The secondary level creates critical intermediate-scale vascular distribution, bridging the gap between widely-spaced large primary veins and fine dense veinlet networks. Branching patterns from primary veins show relative regularity, with secondary veins arising at fairly consistent intervals creating systematic distribution, though some variation occurs with blade region and developmental conditions. Development follows primary vein establishment, with secondary vein primordia initiating along developing primary veins and extending outward coordinating with continued blade expansion. Vascular organization creates hierarchical territories with each secondary vein serving a local blade subsector of 2-4 cm² typical area, collecting water from the primary vein source and distributing it through branching tertiary veins to final veinlet destinations. Mechanical contribution involves distributed support maintaining blade flatness and preventing local buckling or drooping, working synergistically with primary vein main support and higher-order vein fine-scale support. Venation density at the secondary level affects overall blade hydraulic efficiency and photosynthetic capacity, with higher densities creating shorter diffusion distances from mesophyll cells to veins but requiring greater resource investment in vascular tissues.
Zwischen-Zweigvene
branch vein
intermediate lateral vein
second-order lateral vein
sekundäre Seitenader
zweite Ordnung Lateralvene
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 136-165 Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Sack & Holbrook (2006) "Leaf hydraulics" Annual Review of Plant Biology 57=361-381 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 137-172.
Secondary Lateral Vein
In Cabernet Sauvignon berries in Margaret River, Australia during BBCH 79-89, secondary metabolite production shows methoxypyrazine concentrations peaking at 42 ng/L isobutyl-methoxypyrazine at BBCH 79 then declining to 8 ng/L at harvest through continued degradation, while beta-damascenone precursors increase from 15 to 85 µg/L and free norisoprenoids reach 12 µg/L, with total monoterpenes remaining low at 0.3 mg/L characteristic of this non-aromatic variety under warm maritime conditions.
Secondary metabolite production encompasses the biosynthesis of specialized flavor compounds in Vitis vinifera berries during ripening that derive from dedicated secondary metabolic pathways including terpenoid, phenylpropanoid, and carotenoid metabolism, generating the diverse array of monoterpenes, sesquiterpenes, C13-norisoprenoids, methoxypyrazines, and phenolic volatiles that constitute the variety-specific and terroir-expressive aromatic signatures distinguishing fine wines from basic productions. This process involves the coordinated activity of specialized biosynthetic pathways with the methylerythritol phosphate pathway generating isopentenyl diphosphate and dimethylallyl diphosphate precursors for terpenoid synthesis in plastids, terpene synthases converting geranyl diphosphate to monoterpene scaffolds in cytosolic compartments, and carotenoid cleavage dioxygenases generating C13-norisoprenoid volatiles from beta-carotene and other carotenoid substrates in chromoplasts of ripening berries. The temporal regulation of secondary metabolite production shows characteristic induction during the veraison-to-harvest period, with monoterpene pathway genes activated 2-5 fold at veraison, norisoprenoid accumulation paralleling carotenoid degradation during color development and ripening, and methoxypyrazine synthesis reaching maximum before veraison followed by progressive catabolism during subsequent maturation. Cultivar genetics determine secondary metabolite production capacity through variety-specific expression of biosynthetic enzyme genes, with aromatic Muscat varieties expressing high levels of linalool synthase and geraniol synthase, Cabernet Sauvignon family varieties showing characteristic methoxypyrazine O-methyltransferase expression, and diverse varieties expressing different profiles of carotenoid cleavage dioxygenases generating distinct norisoprenoid signatures. Environmental modulation of secondary metabolite production occurs through light-mediated activation of terpenoid and carotenoid pathways, temperature effects on enzyme activities and compound stability, and water stress enhancement of secondary metabolism through activation of stress-responsive regulatory networks that upregulate specialized pathway gene expression. The ecological functions proposed for secondary metabolite production include attraction of seed dispersers through volatile signaling, defense against herbivores and pathogens through toxic or deterrent compounds, and protection against oxidative stress through antioxidant activities, though for viticulture the primary significance lies in the contribution to wine aromatic complexity and quality.
Secondary_Metabolite_Production is subclass of Flavor_Compound_Synthesis representing specialized pathway biosynthesis. Distinguishes from Primary_Flavor_Formation which derives from central metabolism. Sibling to Primary_Flavor_Formation and Flavor_Pathway_Activation. Includes terpenoids, norisoprenoids, methoxypyrazines, and phenolic volatiles. Cultivar-specific and environmentally responsive. Determines variety character.
Sekundärmetabolit-Produktion
Terpenoid-und-Phenol-Biosynthese
secondary flavor compound production
sekundäre Aromaverbindungs-Produktion
specialized metabolite synthesis
specialized volatile biosynthesis
spezialisierte Metabolit-Synthese
terpenoid and phenolic production
Martin DM et al (2010) Functional annotation of the grapevine terpene synthase gene family. BMC Plant Biology 10: 226 Mathieu S et al (2005) A carotenoid cleavage dioxygenase from Vitis vinifera L. Plant Journal 44: 612-621 Dunlevy JD et al (2009) A methyltransferase essential for methoxypyrazine-derived flavour. Plant Journal 59: 404-412
Secondary Metabolite Production
A secondary replacement renewal cane on Vitis vinifera cv. Merlot in double Guyot system, positioned 12cm from cordon alongside primary replacement at 8cm, measuring 8.5mm diameter (slightly smaller than 10mm primary), complete lignification but with minor cold damage visible at tip requiring removal of apical 2 nodes during pruning, 10 nodes retained after tip correction, representing acceptable but not ideal renewal wood that will be selected as fruiting cane only if primary replacement shows significant problems (severe winter damage, excessively vigorous or weak growth, poor positioning after spring shoot development) or if training system modification requires additional fruiting cane beyond primary capacity.
A replacement renewal cane serving as backup or alternative option for following season's fruiting cane selection if primary replacement proves unsuitable, characterized by slightly less optimal positioning or quality compared to primary renewal but still acceptable for fruiting cane use, adequate vigor and lignification for productive fruiting wood, and retention as insurance against primary replacement failure due to winter damage, poor growth, or positioning problems. Secondary replacement renewal canes provide important backup capacity in renewal planning, acknowledging that not all primary replacements successfully transition to fruiting canes due to various problems, ensuring availability of acceptable alternative renewal wood maintaining training system integrity and productivity even when preferred options fail or prove unsuitable during assessment at following dormant pruning.
Secondary replacement renewal canes acknowledge the reality that renewal planning cannot guarantee perfect outcomes, with winter damage, variable spring growth, or changing vineyard conditions sometimes rendering primary replacements unsuitable for fruiting cane use. Retention of secondary replacements provides flexibility and insurance, with selection rates typically 10-30% (used when primaries fail) compared to 80-90% for primaries. In some systems, secondary replacements may be completely removed if primaries prove satisfactory, while other systems routinely use both primaries and secondaries for multiple fruiting cane positions. The primary versus secondary distinction often becomes clear only during following winter's pruning assessment when actual growth, positioning, and quality outcomes can be evaluated against requirements.
alternative Vorratsrute
alternative renewal option
backup replacement cane
secondary renewal position
sekundäre Ersatzrute
Secondary Replacement Renewal Cane
A secondary reproductive bud at node 7 on Vitis vinifera cv. Chardonnay, examined following 82% primary bud loss from spring frost at -3.2°C, dissection revealing one inflorescence primordium containing 72 flower initials plus vegetative shoot primordia with 8 leaf initials, this secondary bud activating 9 days post-frost, developing into a shoot bearing one cluster of 32 berries (44% fruit set from reduced flower number) plus 13 leaves, producing 0.48 kg harvestable crop compared to expected 1.4 kg from the destroyed primary shoot, demonstrating partial but significant yield salvage typical of secondary reproductive compensation in this moderately fruitful cultivar.
A secondary bud containing one or more differentiated inflorescence primordia in addition to vegetative shoot structures, representing relatively uncommon but economically valuable secondary positions capable of producing harvestable crop when activated following primary bud damage, characterized by presence of flower initials visible upon dissection though typically smaller and fewer than in corresponding primary buds, development into fruiting shoots bearing clusters when released from suppression, and cultivar-specific prevalence ranging from rare (5-15% of secondary positions in cultivars like Cabernet Sauvignon, Merlot) to common (40-60% in cultivars like Chardonnay, Riesling). Secondary reproductive buds provide critical partial crop salvage following spring frost or other primary bud damage events, with their abundance and cluster size determining the extent of yield recovery possible through secondary compensation, making cultivar differences in secondary fruitfulness an important consideration for variety selection in frost-prone regions.
Secondary reproductive buds show extreme cultivar variation in prevalence and productivity, making cultivar selection a critical risk management decision for frost-prone regions. Cultivars with high secondary fruitfulness (Chardonnay, Riesling, Gewürztraminer) can recover 35-50% of normal yields following severe primary bud damage, while low-secondary-fruitfulness cultivars (Cabernet Sauvignon, Merlot, Syrah) may achieve only 15-25% yield recovery. This variation reflects genetic differences in resource allocation patterns during bud differentiation, with some cultivars partitioning reproductive capacity between primary and secondary positions while others concentrate it almost exclusively in primaries. The delayed phenology of secondary shoots (7-14 days behind primaries) can create harvest timing challenges and quality variation within vineyards recovering from frost damage.
cluster-bearing secondary bud
fertile secondary bud
fruchtende Nebenknospe
fruiting secondary bud
reproduktive Sekundärknospe
Based on Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Trought & Bramley (2011) "Vineyard variability in Marlborough, New Zealand= characterising spatial and temporal changes in fruit composition and juice quality in the vineyard" Australian Journal of Grape and Wine Research 17=79-89 Molitor et al. (2014) "Late frost damage risk for viticulture under future climate conditions= a case study for the Luxembourgish winegrowing region" Australian Journal of Grape and Wine Research 20=160-168.
Secondary Reproductive Bud
A secondary reproductive shoot on Vitis vinifera cv. Chardonnay from node 8, activated following spring frost (-3.5°C) killing primary shoot at early growth stage (BBCH 12), secondary broke 10 days post-frost, currently early August (approximately 12 days behind normal phenology), 102cm length with 13 leaves totaling approximately 1560 cm² leaf area, bearing single small cluster with 32 berries, estimated shoot crop 145g representing approximately 35-45% of expected primary shoot production from this position (primaries typically 2 clusters, 320-380g total), representing partial crop recovery through secondary activation though with substantial yield deficit, delayed maturation timing (projected harvest 10-14 days later than primary-dominated blocks), and reduced photosynthetic capacity, contributing to whole-vine recovery following 68% primary bud mortality from frost event.
A reproductive shoot originating from secondary bud position within compound bud structure, characterized by development from backup bud component containing differentiated inflorescence primordia activated when primary bud fails or is damaged, delayed phenology lagging primary shoots by 7-14 days, moderate to weak vigor reflecting secondary developmental priority, and reproductive capacity typically reduced compared to primary positions (0-2 inflorescences, often smaller clusters) though providing critical crop recovery following spring damage events. Secondary reproductive shoots represent compensatory reproductive growth from backup positions, rarely occurring in undamaged vineyards where primary shoots dominate, becoming significant following spring frost, hail, or other damage eliminating primary shoots, with secondary reproductive contribution partially offsetting crop losses though unable to fully compensate for primary destruction due to delayed development, reduced vigor, lower fruitfulness, and incomplete secondary activation across damaged positions.
Secondary reproductive shoots provide essential crop recovery function following spring damage, with their reproductive capacity (though reduced versus primaries) enabling partial yield restoration and economic salvage in damaged vineyards. However, secondary reproductive performance shows consistent limitations including delayed phenology (affecting entire seasonal progression from budbreak through harvest), reduced vigor (typically 65-80% of primary shoot size), lower fruitfulness (secondary buds containing fewer or smaller inflorescence primordia), smaller clusters (reduced berry numbers, lighter cluster weights), and incomplete compensation (secondary activation rates typically 60-85% of primary positions leaving gaps). Management involves realistic yield expectations (secondary-dominated crops typically 30-60% of normal primary-based yields), delayed timing recognition (harvest scheduling accommodating phenological lag), quality assessment (secondary fruit often showing different composition requiring separate processing or blending decisions), and long-term planning (assessing whether secondary compensation adequate for economic viability or replanting indicated following severe damage).
Ersatz-Ertragstrieb
backup reproductive shoot
compensatory fruiting shoot
secondary fruiting position
sekundärer Fruchttrieb
Secondary Reproductive Shoot
Pinot Noir grafted on 101-14 Millardet rootstock at 13 years showing secondary root system comprising extensive network of lateral and branch secondary roots, first-order laterals 2-6 cm diameter branching from primary roots at 10-20 cm intervals, second-order branches 0.8-2.5 cm diameter arising from first-order laterals, third and higher-order fine roots <5 mm diameter providing majority of absorption, total secondary root network extending 220 cm laterally from primary roots exploring estimated 12 cubic meters soil volume, fine secondary roots concentrated 25-70 cm depth with mycorrhizal colonization, woody secondary roots showing carbohydrate storage, secondary root system providing majority of vine water and nutrient uptake
Secondary root is any root originating as lateral branch from primary root or from another secondary root, comprising majority of total root system biomass and function including lateral roots branching from primary roots, branch roots arising from other secondary roots creating hierarchical architecture, and woody structural secondary roots with substantial secondary growth, typically ranging 0.5-8 cm diameter in structural secondary roots to <2 mm diameter in fine absorptive secondary roots, showing root orders from first-order laterals branching directly from primary roots through progressively higher-order branches to finest absorptive roots, characterized by lateral origin from parent root rather than vertical growth from seed radicle or cutting base distinguishing from primary roots, serving essential functions including soil exploration through extensive lateral network, water and nutrient absorption primarily through fine secondary roots, carbohydrate storage in woody secondary roots, and structural support through hierarchical branching framework, with secondary root architecture profoundly influenced by rootstock genetics, soil conditions, and vine management determining total root system extent and distribution. Secondary root structural diversity includes= ranging from large woody roots (structural secondary roots 3-8 cm diameter with extensive secondary growth serving transport and storage), through medium structural roots (1-3 cm diameter laterals with moderate secondary thickening), to fine absorptive roots (<2 mm diameter ephemeral roots with minimal secondary growth providing majority of uptake), showing anatomical progression (larger secondary roots with extensive xylem and protective bark versus fine roots with primary anatomy and abundant root hairs), and demonstrating functional specialization (woody secondary roots for transport-storage-structure while fine secondary roots for absorption). Secondary root hierarchical organization includes= comprising first-order laterals (roots branching directly from primary roots representing first branching level), containing second-order branches (roots arising from first-order laterals creating second branching level), including third and higher orders (progressively finer branches creating hierarchical network potentially to 5th-8th order roots), showing decreasing diameter with order (root diameter declining progressively from large first-order to fine high-order roots), and demonstrating functional transitions (lower-order secondary roots primarily transport-storage while higher-order secondary roots primarily absorption). Secondary root origin and development includes= initiating from parent root pericycle (secondary roots forming endogenously from internal pericycle tissue rather than epidermis), emerging through parent cortex (developing secondary root growing outward penetrating parent root tissues), showing acropetal pattern (secondary roots typically more abundant near growing parent root tip declining toward base), exhibiting seasonal production (secondary root initiation concentrated during active growth periods spring and fall), and demonstrating continuous renewal (secondary roots particularly fine orders showing annual or multi-annual turnover with continuous new production). Secondary root spatial distribution includes= extending laterally from primary roots (secondary root network creating horizontal soil exploration exceeding vertical primary root penetration), concentrating in upper soil (majority of absorptive secondary roots typically 20-80 cm depth where moisture and nutrients most available), showing density gradients (secondary root density highest near primary roots declining with distance), demonstrating cultivar patterns (vigorous rootstocks producing more extensive secondary root networks than low-vigor types), and influenced by soil heterogeneity (secondary root proliferation greatest in favorable soil zones with adequate resources). Secondary root functions includes= exploring soil volume (extensive secondary root network potentially exploring 5-20+ cubic meters soil per vine), absorbing water and nutrients (fine secondary roots providing 70-90% of total vine water and nutrient uptake through root hairs and mycorrhizae), transporting to primary roots (woody secondary roots conducting absorbed resources to primary root and trunk), storing carbohydrates (structural secondary roots accumulating reserves particularly during dormancy), providing anchorage (lateral secondary root network contributing to horizontal anchorage preventing toppling), and enabling adaptive responses (secondary roots showing rapid proliferation in resource-rich zones and reduced growth in poor zones). Secondary root seasonal dynamics includes= showing spring growth flush (secondary roots resuming elongation and production early spring often before bud break), exhibiting summer growth (continued secondary root development though potentially reduced during peak shoot growth), demonstrating fall growth period (secondary roots showing renewed activity late season as shoot growth ceases), showing winter dormancy (secondary root growth minimal during cold periods though physiological activity continuing), and displaying turnover cycles (fine secondary roots showing annual mortality and replacement while woody secondary roots persisting for years or decades). Secondary root versus primary root includes= distinguished by origin (secondary roots arising as lateral branches from existing roots versus primary roots from radicle or cutting base), showing different orientations (secondary roots typically horizontal or oblique versus primary roots typically vertical or deep-penetrating), exhibiting diameter differences (secondary roots ranging from fine to moderately large versus primary roots typically large diameter), demonstrating functional emphasis (secondary roots emphasizing lateral exploration and absorption versus primary roots emphasizing depth and main structural axis), and revealing architectural roles (secondary roots creating branching network versus primary roots providing main trunk of root system).
Secondary_Root emphasizes lateral branch origin from primary or other secondary roots and is distinguished from Primary_Root (radicle or cutting origin) by focus on branching architecture. Includes Lateral_Secondary_Root, Branch_Secondary_Root, and Woody_Secondary_Root. Critical for understanding root system branching hierarchy, soil exploration, absorption function, hierarchical organization, and the extensive branching network providing majority of root system soil contact and resource uptake.
Adventitious lateral root
Branch root
Lateral root
Seitenwurzel
Sekundärwurzel
Verzweigungswurzel
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Eissenstat, D. M., et al. (2000). Building roots in a changing environment= Implications for root longevity. New Phytologist, 147(1), 33-42.
Secondary Root
Cabernet Sauvignon secondary vascular bundle mesocarp showing extensive distribution network= approximately 12-18 secondary branches per primary bundle creating total 48-72 secondary bundles per berry, with branches 35-95 μm diameter containing xylem (15-45 μm, 1-3 vessels 18-32 μm diameter) and phloem (20-55 μm, 4-8 sieve elements 12-20 μm diameter), creating distribution network reducing maximum distance from any mesocarp cell to nearest vascular tissue to 350-550 μm ensuring effective resource distribution from primary bundle import throughout mesocarp storage tissue. Merlot secondary vascular bundle mesocarp branch pattern examined by 3D imaging= secondary bundles branching from primary bundles at 900-1600 μm intervals along primary bundle length, branch angles 45-95° from primary bundle axis, branch lengths 800-2500 μm extending into mesocarp before terminating (most branches) or occasionally anastomosing with branches from adjacent primary bundles (10-20% of secondary bundles showing connections), creating complex 3D distribution network throughout berry with some redundancy through anastomoses potentially providing alternative pathways if individual branches dysfunctional. Syrah secondary vascular bundle mesocarp phloem distribution function= secondary bundle phloem sieve elements conducting sugars from primary bundle phloem into surrounding mesocarp tissue, with sugar concentration gradient estimated from secondary bundle (high concentration near sieve element unloading) to distant mesocarp cells (lower concentration before uptake and storage), driving diffusion and/or cell-to-cell transport distributing sugars throughout mesocarp achieving relatively uniform final distribution (central mesocarp 24.8%, mid-radius 24.5%, peripheral 25.1%, 95% air-filled and non-conducting, demonstrating coordinated xylem-to-phloem transition across entire vascular network (primary plus secondary vascular bundle mesocarp) eliminating xylem import and establishing phloem-exclusive import supporting ripening phase sugar accumulation.
A secondary vascular bundle mesocarp is vascular bundle mesocarp containing or adjacent to lateral (secondary) vascular bundles that branch from primary dorsal bundles and ramify throughout the mesocarp creating the distribution network, typically comprising 40-100+ secondary branches per berry (8-20 branches per primary bundle) with branch diameter 30-120 μm, serving to distribute imported resources from primary bundles throughout mesocarp tissue ensuring all storage parenchyma cells receive access to phloem-imported sugars, water, and minerals. Secondary vascular bundle mesocarp structural organization= (1) lateral branches diverging from primary bundles at irregular intervals creating branching network throughout mesocarp (2) branches typically smaller diameter (30-120 μm) than primary bundles (150-280 μm) with progressively smaller diameter in higher-order branches (tertiary branches 20-60 μm when present) (3) branch angle varying from nearly perpendicular to primary bundle (70-110° divergence) to more acute angles (30-60°) (4) vascular tissue in secondary bundles showing similar organization to primary bundles (xylem plus phloem) but smaller scale with 1-4 xylem vessels 15-35 μm diameter, 3-8 phloem sieve elements 10-20 μm diameter (5) some secondary bundles terminating in mesocarp tissue (blind endings) while others potentially anastomosing (connecting with other secondary bundles) creating limited vascular network connectivity. Secondary vascular bundle mesocarp distribution function= (1) distribution of phloem-imported solutes from primary bundle import points throughout mesocarp tissue volume (2) reduction of maximum distance from any storage parenchyma cell to nearest vascular tissue from potentially 2-4 mm (if only primary bundles present) to typically 300-600 μm (with secondary bundle network) facilitating solute distribution (3) creation of multiple import points throughout mesocarp allowing localized distribution rather than requiring long-distance diffusion or cell-to-cell transport from primary bundles (4) potential local resource distribution heterogeneity with cells near secondary bundles potentially receiving earlier or greater solute delivery than cells distant from vascular tissue though overall composition showing <5% spatial variation suggesting effective equilibration. Secondary vascular bundle mesocarp development= (1) secondary bundle initiation during early berry development through vascular cambium activity and/or differentiation of provascular tissue into functional vascular tissue (2) secondary bundle elongation and expansion during berry growth maintaining distribution network as berry enlarges (3) potential continued secondary bundle formation during berry development though most secondary bundles established by veraison (4) xylem in secondary bundles functional during Phase I, cavitating at/before veraison similar to primary bundle xylem phloem in secondary bundles functional throughout ripening conducting sugars from primary bundles into surrounding mesocarp. Secondary vascular bundle mesocarp spatial distribution creating berry tissue zones= (1) vascular bundle-rich zones (high secondary bundle density) potentially showing characteristics of better-served tissue (2) vascular bundle-poor zones (low secondary bundle density) potentially showing resource limitation though compensated by cell-to-cell transport and diffusion (3) overall relatively uniform secondary bundle distribution creating relatively uniform resource delivery throughout mesocarp (4) potential cultivar variation in secondary bundle number and distribution affecting resource distribution efficiency.
Secondary_Vascular_Bundle_Mesocarp represents tissue containing or adjacent to lateral branch bundles and is distinguished from Primary_Vascular_Bundle_Mesocarp (main longitudinal bundles) by smaller size, lateral branching pattern, and distribution rather than primary import function. Secondary vascular bundle mesocarp creates the local distribution network delivering resources throughout mesocarp. Understanding secondary vascular bundle mesocarp is relevant for detailed berry anatomical characterization, for comprehending resource distribution mechanisms ensuring uniform composition despite localized import, and for understanding complete berry vascular architecture as integrated primary-plus-secondary network.
Seitenleitbündel-Fruchtfleisch
Zweig-Leitbündel-Gewebe
branch bundle zone
lateral vascular bundle tissue
sekundäres Gefäßbündel-Mesokarp
Rogiers, S.Y. et al. (2006). Vascular function in berries. Australian Journal of Grape and Wine Research 12=203-214 Chatelet, D.S. et al. (2008). Xylem structure and connectivity in grapevine shoots. American Journal of Botany 95=1076-1085.
Secondary Vascular Bundle Mesocarp
A secondary vegetative bud at node 7 on Vitis vinifera cv. Merlot, activated following primary bud destruction by hail damage in late April, dissection revealing well-formed vegetative shoot primordia with 8 leaf initials and 2 tendril primordia but complete absence of inflorescence structures, developing into a vigorous vegetative shoot reaching 125cm length with 16 fully expanded leaves by mid-summer, providing substantial photosynthetic capacity (estimated 940 cm² leaf area) supporting reserve accumulation and bud differentiation for following season but contributing zero current-season yield, typical of secondary bud response in this cultivar where secondary positions rarely contain reproductive primordia.
A secondary bud containing exclusively or predominantly vegetative primordia without differentiated inflorescence structures, representing the most common secondary bud type across most cultivars and node positions, characterized by absence of flower initials upon dissection, development into purely vegetative shoots when activated following primary bud damage, and contribution to canopy leaf area without direct yield production. Secondary vegetative buds reflect the general pattern of reduced fruitfulness in secondary positions compared to primaries, resulting from later differentiation timing (typically 2-4 weeks after primary bud initiation), smaller meristem size with reduced capacity for inflorescence primordium development, and often less favorable light exposure during critical differentiation periods as secondary buds form in the shadow of developing primary buds and expanding shoot canopy.
Secondary vegetative buds represent the default condition for secondary positions in most Vitis vinifera cultivars, with reproductive secondary buds being the exception rather than the rule. This pattern reflects evolutionary priority given to primary bud development, with secondary buds serving primarily as insurance against primary loss rather than as routine contributors to reproduction. The predominance of secondary vegetative buds has important implications for post-frost yield estimation, as compensatory growth from secondary breakage provides canopy recovery and photosynthetic capacity but minimal crop salvage. Some training systems (particularly head-trained, spur-pruned gobelet) may occasionally utilize secondary bud positions for renewal wood when primary buds are excessively fruitful, deliberately removing primary shoots to force secondary vegetative growth for next season's fruiting positions.
nicht-fruchtende Nebenknospe
non-fruiting secondary bud
sterile secondary bud
vegetative Sekundärknospe
vegetative backup bud
Based on Morrison (1991) "Bud development in Vitis= Anatomical study of formation and growth of the primary grapevine bud during one annual cycle" Australian Journal of Grape and Wine Research 72=296-301 Sánchez & Dokoozlian (2005) "Bud microclimate and fruitfulness in Vitis vinifera L." American Journal of Enology and Viticulture 56=319-329 Vasconcelos et al. (2009) "The flowering process of Vitis vinifera= a review" American Journal of Enology and Viticulture 60=411-434.
Secondary Vegetative Bud
A secondary vegetative shoot on Vitis vinifera cv. Sauvignon Blanc from node 5, activated following spring frost (-3.2°C) that killed primary shoot at green tip stage (BBCH 07), secondary broke 11 days post-frost reaching 95cm length by early August (compared to typical 140cm for primary shoots at this position), 13 leaves totaling approximately 1450 cm² leaf area, no inflorescences (secondary positions rarely reproductive even at normally fruiting node positions), representing partial canopy recovery following frost damage, retained for photosynthetic contribution supporting remaining crop from frost-surviving shoots on other nodes, with this secondary shoot providing approximately 18% replacement of lost photosynthetic capacity from destroyed primary, inadequate for full compensation but valuable for supporting fruit maturation and reserve accumulation.
A vegetative shoot originating from secondary bud position within compound bud structure, characterized by development from backup bud component activated when primary bud fails or is damaged, typical vigor moderate to weak reflecting secondary developmental priority, delayed phenology (7-14 days behind primary shoots), and origin circumstances including spring frost damage eliminating primaries, hail destruction, disease or pest damage, or occasional spontaneous secondary breakage alongside intact primaries creating doubled shoot positions. Secondary vegetative shoots represent compensatory growth from backup bud positions, most commonly arising following spring damage events, contributing to canopy recovery and photosynthetic capacity though with delayed development and typically reduced vigor compared to primary shoots, requiring management decisions regarding retention for recovery benefits versus removal for canopy density control.
Secondary vegetative shoots serve critical recovery function following primary bud damage, providing partial canopy replacement and photosynthetic capacity supporting fruit development from surviving shoots and maintaining vine function preventing complete crop and canopy loss. However, secondary shoots show consistent limitations including delayed phenology (7-14 day lag affecting all subsequent development), reduced vigor (typically 60-75% of primary shoot growth), low reproductive potential (secondary buds rarely containing inflorescences), and in severe damage scenarios, inadequate compensation (secondary breakage rates often 60-80% of primary positions leaving gaps). Management involves retaining secondary shoots for recovery benefits while removing redundant shoots where both primary and secondary broke (doubled positions), with retention decisions balancing photosynthetic contribution against canopy density concerns and recognition that secondary-dominated canopies show delayed maturation and reduced fruit quality compared to primary-dominated normal canopies.
Ersatztrieb vegetativ
backup position vegetative shoot
compensatory vegetative shoot
secondary non-fruiting shoot
sekundärer vegetativer Trieb
Based on Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Molitor et al. (2014) "Late frost damage risk for viticulture under future climate conditions= a case study for the Luxembourgish winegrowing region" Australian Journal of Grape and Wine Research 20=160-168 Trought & Bramley (2011) "Vineyard variability in Marlborough, New Zealand" Australian Journal of Grape and Wine Research 17=79-89.
Secondary Vegetative Shoot
Syrah secondary wood ray initiated at ring 8 (8 years post-planting) in 15-year trunk shows radial extent 18-28 mm from ring 8 outward through rings 9-15 to cambium, multiseriate structure 3-4 cells wide, living ray parenchyma throughout 18-28 mm extent (all within sapwood zone), ray providing radial transport and storage since initiation adding to existing primary ray network, ray representing cambial ray initial multiplication process as trunk circumference increased from ~125 mm at year 8 to ~280 mm at year 15 requiring additional rays to maintain ray density
A secondary wood ray is a wood ray initiated from new ray initials formed in vascular cambium after initial trunk development, representing rays that begin in outer annual rings and extend only through subsequently formed xylem rather than connecting to innermost wood, increasing total ray density as trunk diameter expands. Secondary wood ray formation occurs= through ray initial multiplication in cambium (fusiform initials dividing to produce new ray initials increasing ray number as cambium circumference grows), initiated in outer rings (secondary rays first appearing in rings 5-15+ depending on trunk growth rate), extending inward only through post-initiation rings (secondary ray present only from initiation ring outward to cambium, not extending to inner trunk), increasing ray density maintenance (new ray initiation compensating for increased cambial circumference maintaining relatively constant ray density per unit area), and continuous process (ray initials continuously added throughout trunk life as diameter increases). Secondary wood ray anatomical characteristics include= limited radial extent (extending 5-40 mm from initiation ring to cambium depending on years since formation and growth rates), presence only in outer rings (secondary rays absent from inner trunk showing radial sections with rays beginning mid-trunk rather than at pith), typical multiseriate organization (most secondary rays 2-5 cells wide similar to primary rays), height and width comparable to primary rays (15-30 cells high, 3-5 cells wide), and variable length (recently initiated secondary rays showing short extent 5-15 mm, older secondary rays extending 20-40 mm if initiated 8-15 years previously). Secondary wood ray initiation patterns include= increasing frequency in outer rings (young trunk inner 5 rings showing few or no secondary rays, outer rings 8-20 showing progressive secondary ray initiation), preferential initiation during vigorous growth (wide annual rings typically showing more secondary ray initiation maintaining or increasing ray density), stress-year reduced initiation (narrow rings showing fewer new ray initials as cambium focuses on survival rather than ray multiplication), and cultivar variation (some varieties showing more prolific ray initial multiplication than others). Secondary wood ray functions include= supplementing radial transport capacity (new rays adding lateral transport pathways as trunk expands), maintaining ray density (compensating for diameter increase that would otherwise dilute ray density if only primary rays present), enhancing storage capacity (additional ray parenchyma providing more carbohydrate storage sites), and improving hydraulic integration (increased lateral connections through additional rays enhancing trunk hydraulic network). Secondary wood ray distribution patterns= uniform circumferential distribution (secondary rays initiating around entire trunk maintaining even ray spacing), initiation in regions of active cambial growth (vigorous cambial sectors showing more secondary ray formation), random or systematic spacing (new ray initials positioned to maintain relatively even ray separation 80-150 μm tangential spacing), and coordination with vessel and fiber production (cambium balancing axial element production with ray initial multiplication). Secondary wood ray significance in trunk expansion includes= enabling ray density maintenance (without secondary ray initiation, trunk diameter increase would cause ray density to decline as circumference expands), providing flexibility in cambial organization (ability to add rays allowing cambium to adjust tissue proportions), enhancing sapwood functionality (secondary rays in outer sapwood contributing to active transport and storage), and adapting to changing demands (cambium potentially modulating ray initiation rate based on physiological needs or environmental conditions). Secondary wood ray identification includes= observation of ray origin in radial section (secondary rays beginning in middle to outer rings rather than extending to pith), measurement of radial extent (limited extent confirming recent initiation), comparison with known primary rays (rays extending to pith identified as primary, shorter rays as secondary), and aging based on initiation ring (counting rings from ray origin to cambium determining years since initiation). Secondary wood ray proportion of total rays varies= young trunk inner 5-10 rings showing 0-20% secondary rays (primary rays dominating in early wood), mature trunk middle rings showing 20-45% secondary rays (progressive accumulation of secondary rays initiated at various times), outer trunk most recent rings showing 30-60% secondary rays (cumulative effect of decades of ray initial multiplication), and old trunk potentially showing 40-70% secondary rays in outer sapwood (long history of ray initiation producing majority of current rays being relatively recent origin).
Secondary_Wood_Ray emphasizes later-initiated rays and is distinguished from Primary_Wood_Ray (present from early development through all rings) and Wood_Ray (all rays regardless of origin) by limited radial extent beginning in outer rings. Critical for understanding cambial organization dynamics, trunk diameter expansion effects on tissue proportions, ray density maintenance mechanisms, and adaptation of cambial outputs to changing trunk dimensions and physiological demands.
Additional ray
Late-formed ray
Newly initiated ray
Sekundärer Holzstrahl
Später gebildeter Strahl
Zusätzlicher Xylemstrahl
Carlquist, S. (2001). Comparative Wood Anatomy (2nd ed.). Springer. | Larson, P. R. (1994). The Vascular Cambium= Development and Structure. Springer.
Secondary Wood Ray
Cabernet Sauvignon berries at BBCH 89 (full maturity) typically contain 1-2 fully developed seeds measuring 6-8 mm length with dark brown, lignified testa= seed coat tannins at 12-15% dry weight basis, tannin polymerization index 65-75%, moisture content 30-35%
A seed is a mature or developing ovule of Vitis vinifera that contains the embryo, endosperm, and protective seed coat tissues, resulting from successful fertilization of the ovule within the berry. The seed develops through distinct phases= early embryogenesis (0-4 weeks post-fertilization), mid-development with endosperm accumulation (4-8 weeks), and final maturation with desiccation and dormancy acquisition (8-16 weeks post-fertilization, corresponding to BBCH 79-89). Vitis vinifera seeds are characterized by a pyriform shape, hard testa with distinctive beak and chalazal region, and variable development outcomes including full maturity, incomplete development, or abortion. Seed number per berry varies dramatically by cultivar and fertilization success= seedless varieties (e.g., Thompson Seedless) contain only rudimentary seed traces, while seeded wine grapes typically develop 1-4 seeds per berry depending on pollination efficiency and genetic factors. Seed development is closely coordinated with berry growth, as developing seeds produce hormones (particularly auxins and gibberellins) that regulate berry expansion during Phase I and Phase II growth. The presence and developmental state of seeds significantly influences berry size, shape, and ripening patterns. Seed maturity at harvest affects wine quality through potential release of phenolic compounds and oils during crushing and fermentation. Commercial viticulture distinguishes between seeded wine grape production (where seed maturity indicates optimal harvest timing) and seedless table grape cultivation (where seed abortion is desirable). Seed structure includes the embryo with cotyledons and radicle, nutritive endosperm tissue, and multi-layered testa with outer epidermis, sclerified middle layers, and inner integument remnants.
Seed emphasizes the complete developmental spectrum from fertilization through maturation or abortion and is distinguished from Berry (the surrounding fruit tissue) by its developmental origin from ovule rather than ovary wall. Critical for understanding berry development hormonal regulation, harvest timing decisions, wine quality implications of seed phenolics, and seedless versus seeded cultivar distinctions.
Grape seed
Rebensamen
Traubenkern
Vitis seed
Weinsamen
Ristic, R., & Iland, P. G. (2005). Relationships between seed and berry development of Vitis vinifera L. cv Shiraz= Developmental changes in seed morphology and phenolic composition. Australian Journal of Grape and Wine Research, 11(1), 43-58. | Cadot, Y., Miñana-Castelló, M. T., & Chevalier, M. (2006). Anatomical, histological, and histochemical changes in grape seeds from Vitis vinifera L. cv Cabernet franc during fruit development. Journal of Agricultural and Food Chemistry, 54(24), 9206-9215.
Seed
Merlot seed-adjacent inner pericarp (innermost 2 cell layers, 45-75 μm thickness) showing characteristic structure= cells 42-58 μm diameter (smallest in pericarp), walls 1.2-1.6 μm thick, tannins 6.2-7.8 mg/g (highest in inner pericarp region), with cells directly contacting seed coat surfaces during early development (BBCH 73-75, weeks 3-6) but separated by 15-35 μm locular cavity air gap at maturity (BBCH 89, week 18) as berry expands beyond seed dimensions, demonstrating temporal change from seed contact to seed proximity during development. Cabernet Sauvignon seed-adjacent inner pericarp tannin concentration gradient= innermost cell layer (direct seed contact position) 7.2-8.5 mg/g tannins, second layer (20-40 μm from seeds) 6.5-7.8 mg/g, third layer (50-80 μm from seeds) 5.2-6.8 mg/g, demonstrating systematic tannin gradient with highest concentration in seed-adjacent position declining with distance, potentially supporting hypothesis of seed influence on pericarp tannin accumulation though alternative explanation is position-dependent tannin synthesis programming. Syrah seed-adjacent inner pericarp comparison between 2-seed and 4-seed berries= 2-seed berries (less seed surface area) showing seed-adjacent zone 55-85 μm thickness, 6.5-7.8 mg/g tannins 4-seed berries (more seed surface area) showing seed-adjacent zone 65-95 μm thickness, 7.2-8.6 mg/g tannins, suggesting greater seed presence/contact area correlates with enhanced seed-adjacent pericarp tannin accumulation, though effect is modest and variability high. Pinot Noir seed-adjacent inner pericarp developmental time-course= week 4 post-fruit set (BBCH 73, early development) showing seed-adjacent cells 38-52 μm in intimate contact with seed coat outer cell layers, no air space week 8 (BBCH 77, lag phase) showing cells 48-65 μm beginning separation with 2-8 μm gaps week 12 (BBCH 83, veraison) showing 55-70 μm cells separated by 12-25 μm air gaps week 18 (BBCH 89, maturity) showing 52-68 μm cells (slight shrinkage from turgor loss) separated by 20-40 μm gaps, demonstrating progressive locular cavity air space development creating transition from seed contact to seed proximity. Chardonnay white cultivar seed-adjacent inner pericarp showing presence in whites= 2 cell layers, 50-80 μm thickness, cells 48-62 μm diameter, tannins 4.5-6.2 mg/g (elevated like red cultivars relative to middle pericarp), demonstrating seed-adjacent inner pericarp as distinct zone present in both red and white cultivars though with lower absolute tannin levels in whites.
A seed-adjacent inner pericarp is the innermost 1-3 cell layers of the inner pericarp in direct or near-direct contact with seed surfaces or locular cavity surfaces where seeds are positioned, typically 30-100 μm thickness, characterized by smallest cells within inner pericarp, highest tannin concentration within inner pericarp, potential biochemical interactions with seeds, and potential separation from seeds through locular cavity air space development during maturation, representing the absolute innermost pericarp boundary interfacing with seed-containing spaces. Seed-adjacent inner pericarp cellular structure comprises= (1) innermost 1-3 cell layers showing smallest cells 40-65 μm diameter (versus 70-90 μm in outer inner pericarp layers) creating minimum cell size within entire pericarp (2) cells with moderately thick walls 1-1.8 μm (thicker than middle pericarp 0.4-0.8 μm, approaching outer pericarp hypodermis 1.5-3 μm) providing structural integrity at pericarp-seed boundary (3) cells often showing irregular shapes conforming to seed surface contours particularly in developing berries when pericarp cells directly contact seeds (4) intercellular spaces prominent particularly in mature berries as locular cavity air spaces develop creating partial or complete separation between seed-adjacent cells and seed surfaces. Seed-adjacent inner pericarp compositional characteristics= (1) highest tannin concentration within inner pericarp region, typically 5.5-8 mg/g dry weight (versus 3.5-5.5 mg/g in outer inner pericarp layers), approaching outer pericarp tannin levels (8-15 mg/g) (2) tannin composition similar to other inner pericarp zones with predominantly procyanidins, polymerization degree 12-18 subunits, moderate galloylation 12-20% (3) potential seed-derived phenolic compounds in seed-adjacent cells if diffusion occurs from seeds, though evidence for substantial seed-to-pericarp phenolic transfer remains limited with most inner pericarp tannins likely synthesized in situ (4) sugar and acid content similar to other pericarp regions showing minimal impact of seed proximity on primary metabolite accumulation. Seed-adjacent inner pericarp development shows temporal changes in seed contact= early development (BBCH 71-75, weeks 1-6) with developing seeds filling locular cavities and seed coats directly contacting pericarp cells creating intimate seed-pericarp interface potentially allowing diffusion of compounds or signals mid development during lag phase (BBCH 77, weeks 6-8) with seeds completing growth and beginning maturation while pericarp cells continue expansion late development and ripening (BBCH 81-89, weeks 9-18+) with locular cavity air spaces developing as berry expands more than seeds, creating 10-50 μm gaps between mature seed coats and seed-adjacent pericarp cells, eliminating direct contact but maintaining close proximity. Seed-adjacent inner pericarp potential seed interactions= developing seeds producing hormone signals (auxins, gibberellins, ABA) potentially diffusing into adjacent pericarp cells affecting metabolism or gene expression seed tannins or other phenolics potentially diffusing from seed coats into adjacent pericarp cells during early development when direct contact exists, though quantitative significance uncertain pericarp-derived signals potentially affecting seed development though direction and nature of signaling unclear. Seed-adjacent inner pericarp in seedless cultivars shows modified characteristics= stenospermocarpic seedless cultivars (Thompson Seedless, Flame Seedless) showing seed traces (arrested seed structures) rather than normal seeds, with seed-adjacent inner pericarp present but potentially modified in thickness (often 80-120 μm versus 100-300 μm in seeded types) and tannin content (sometimes reduced 20-40% reflecting smaller seed trace contact area).
Seed_Adjacent_Inner_Pericarp represents the absolute innermost pericarp layer and is distinguished from Locular_Inner_Pericarp which may include slightly more external inner pericarp layers. Seed-adjacent inner pericarp shows highest tannin levels within inner pericarp and potential for seed-pericarp interactions during development when direct contact exists. Understanding seed-adjacent inner pericarp is relevant for detailed anatomical characterization, for investigating seed-pericarp interactions, and for understanding tannin distribution at the finest spatial scale within berry tissues.
Samen-Kontaktschicht
Samenkontakt-Schicht
innermost pericarp
samenangrenzedes inneres Perikarp
seed-contact layer
Cadot, Y. et al. (2006). Tannin localization in grape tissues. Journal of Agricultural and Food Chemistry 54=2973-2987 Ristic, R., Iland, P.G. (2005). Relationships between seed and berry development. Australian Journal of Grape and Wine Research 11=217-226.
Seed Adjacent Inner Pericarp
In Grenache berries in Châteauneuf-du-Pape, Rhône Valley during BBCH 81-89, seed browning progresses from visual score 1.2 at veraison representing predominantly green seeds through score 2.8 at mid-ripening to score 3.8 indicating well-browned seeds at harvest over 45 days, with browning index at 420 nm increasing from 0.18 to 0.68 absorbance units, correlating with extractable tannin reduction from 9.5 to 3.8 mg/g seed as oxidative modification reduces tannin solubility under warm Mediterranean conditions averaging 29°C.
Seed browning describes the progressive development of brown coloration in Vitis vinifera grape seed coat tissues during the ripening period, wherein enzymatic and non-enzymatic oxidation of phenolic compounds generates brown-colored quinone polymers and melanin-like pigments that transform seed appearance from green or yellow-green at veraison to tan and ultimately dark brown at harvest maturity, providing a visible indicator of seed phenolic maturation status that correlates with reduced tannin extractability and modified astringency potential in wines produced from these grapes. This process involves the oxidation of colorless catechol and pyrogallol groups present on flavan-3-ol subunits of seed proanthocyanidins, with polyphenol oxidase catalyzing two-electron oxidation to ortho-quinones that undergo rapid non-enzymatic polymerization generating brown-colored products, while peroxidase activities contribute additional oxidative capacity in the presence of hydrogen peroxide generated during cellular metabolism. The temporal progression of seed browning shows characteristic patterns with initial browning visible at the chalazal end of seeds shortly after veraison where oxidative processes initiate, followed by progressive extension throughout the seed coat over the subsequent 4-6 weeks until uniform brown coloration indicates complete oxidative modification of surface phenolic compounds accessible to oxidative enzymes and atmospheric oxygen. The quantitative assessment of seed browning employs colorimetric methods including visual scoring on structured color scales ranging from 1 representing green through 4 representing dark brown, spectrophotometric measurement of absorbance at 420 nm providing objective browning indices, and digital image analysis enabling standardized color quantification that correlates with tannin extractability measurements. The relationship between seed browning and wine quality reflects the correlation between oxidative modification visible as browning and the reduced extractability of seed tannins that limits their contribution to wine astringency, with adequately browned seeds releasing 40-70% less tannin during standard maceration compared to green seeds, reducing the harsh astringent character associated with immature seed tannin extraction. Environmental factors influence seed browning rates through effects on oxidative enzyme activities and oxygen availability, with warm temperatures accelerating browning through enhanced enzyme kinetics while excessive heat may inactivate enzymes potentially slowing browning progression, and vineyard conditions affecting berry transpiration influencing oxygen diffusion into seed tissues.
Seed_Browning is subclass of Seed_Phenolic_Maturation representing the visible oxidative modification of seed coat. Distinguishes from Seed_Tannin_Polymerization by emphasizing color change as indicator. Sibling to Seed_Tannin_Polymerization and Phenolic_Seed_Maturity. Visual indicator of seed phenolic status. Correlates with extractability reduction. Assessed by color scoring and spectrophotometry at 420 nm.
Samen-Bräunung
Samen-Farbentwicklung
Samen-phenolische-Bräunung
Samenschalen-oxidative-Bräunung
seed coat oxidative browning
seed color development
seed oxidation
seed phenolic browning
Ristic R and Iland PG (2005) Relationships between seed and berry development of Vitis vinifera L. Australian Journal of Grape and Wine Research 11: 43-58 Kennedy JA et al (2000) Grape seed development germination and proanthocyanidin composition. Journal of Agricultural and Food Chemistry 48: 1490-1495 Cadot Y et al (2006) Anatomical histological and histochemical changes in grape seeds. Journal of Agricultural and Food Chemistry 54: 9206-9215
Seed Browning
Cabernet Sauvignon seed brush attachment in berry with 2 fully developed seeds shows brush cross-section with 6-8 vascular bundle traces including 3-4 larger traces (35-45 μm xylem vessels) supplying pericarp and 2-4 smaller traces (20-30 μm vessels) aligned toward seed positions 1.5-2.5 mm from brush center, indicating coordinated vascular architecture supporting both berry and seed development
A seed brush attachment is a brush attachment zone specifically emphasizing vascular bundle traces and receptacle tissues that connect to or align with seed positions within the berry, representing the portion of brush vasculature that supplied nutrients to developing seeds during berry development. Seed brush attachment reflects the anatomical relationship between berry vascular architecture and seed positioning= pedicel vascular bundles entering the berry typically branch and distribute to supply both pericarp tissues and developing seeds, with some vascular traces extending specifically toward seed attachment points in the locular region. Seed brush attachment anatomical features include= vascular traces diverging from main receptacle bundles toward seed positions (typically 1-3 traces per seed in seeded berries), specialized receptacle tissue in seed-aligned regions (potentially including funiculus remnants or locular tissue connections), vascular bundle terminations at or near former seed attachment sites, and in seeded berries, alignment of brush vascular pattern with 2-4 seed positions typical of Vitis vinifera. Seed brush attachment development and function during berry growth includes= during early berry development (BBCH 71-77) vascular traces supply developing seeds with water, minerals, and nutrients supporting embryo and endosperm development, during seed maturation (BBCH 79-85) continued vascular supply until seeds achieve maturity and reduced vascular dependency, during late ripening (BBCH 85-89) seed-associated vascular traces may show reduced activity as seeds complete maturation and berry-seed vascular connection diminishes. Seed brush attachment characteristics vary by seed development status= in seeded berries with fully developed seeds (1-4 seeds per berry), brush shows multiple vascular traces with some aligned toward seed positions, brush cross-section may show 4-8 distinct vascular bundle traces representing both pericarp and seed supply routes, in berries with aborted seeds (stenospermocarpic varieties or individual seed abortion), brush may show reduced or modified vascular architecture reflecting failed seed development, vascular traces toward aborted seed positions may be smaller or absent. Seed brush attachment visibility and identification includes= in fresh separated berries, careful dissection can reveal vascular trace divergence patterns, in dried brushes, differential lignification or vascular bundle size may indicate seed-associated versus pericarp-associated traces, microscopic examination of brush cross-sections reveals vascular bundle number, arrangement, and size distribution reflecting seed supply architecture. Seed brush attachment significance includes= evidence of berry vascular architecture supporting seed development, potential correlation between seed number and brush vascular complexity (berries with more seeds showing more complex brush vasculature), contribution to cultivar-specific brush characteristics (seeded versus seedless varieties showing different brush vascular patterns), and insight into developmental coordination between seed and berry development through shared vascular supply. Variation in seed brush attachment includes= seed number variation (0-4 seeds affecting vascular trace number and distribution), seed position variation (affecting specific vascular trace orientations within brush), seed development status (fully developed versus aborted seeds affecting vascular trace development), and cultivar differences in berry-seed vascular architecture.
Seed_Brush_Attachment emphasizes seed-associated vascular components and is distinguished from Pedicel_Brush_Attachment (pedicel-derived tissues) and Brush_Attachment_Zone (complete interface) by focus on vascular traces connecting to seed regions. Critical for understanding berry-seed developmental coordination, vascular architecture variation between seeded and seedless cultivars, and brush morphology as indicator of seed development status.
Samen-Bürste
Samengefäß-Ansatz
Samenversorgungs-Rest
Seed vascular attachment
Seed-associated brush
Pratt, C. (1971). Reproductive anatomy in cultivated grapes - a review. American Journal of Enology and Viticulture, 22(2), 92-109. | Galet, P. (2000). A Practical Ampelography= Grapevine Identification. Comstock Publishing Associates.
Seed Brush Attachment
Cabernet Sauvignon seed coat at maturity (BBCH 89) measures 200-250 μm thickness with fully lignified sclereid layers 150-180 μm thick, dark brown coloration indicating tannin polymerization >70%, contains 10-14% condensed tannins (dry weight), and provides mechanical resistance requiring 15-25 N force for crushing
A seed coat (testa) is the protective outer covering of a Vitis vinifera seed that develops from the integuments of the ovule following fertilization, providing mechanical protection, regulating water and gas exchange, containing phenolic compounds, and controlling germination timing through dormancy mechanisms. Grape seed coat development proceeds through distinct phases= integument differentiation (0-3 weeks post-fertilization), cell division and expansion (3-6 weeks), lignification and tannin accumulation (6-12 weeks corresponding to BBCH 79-85), and final maturation with desiccation (12-16 weeks, BBCH 85-89). The mature seed coat consists of multiple tissue layers derived from inner and outer integuments= outer epidermis with thick cuticle (providing waterproofing), hypodermal layers with phenolic-rich cells (containing condensed tannins), sclereid layers with heavily lignified and thickened cell walls (providing mechanical strength and hardness), and inner integument remnants forming innermost seed coat layers. Seed coat thickness in mature Vitis vinifera seeds ranges 150-300 μm depending on cultivar and developmental conditions, with sclereid layers comprising 60-80% of total thickness. Seed coat coloration transitions during development= green during early development (chlorophyll in outer layers), tan during mid-development (chlorophyll degradation, early tannin polymerization, BBCH 81-83), brown during late development (progressive tannin polymerization and lignification, BBCH 85-87), and dark brown to black at full maturity (complete lignification and maximum tannin polymerization, BBCH 87-89). Seed coat composition includes= structural components (cellulose, lignin providing rigidity), phenolic compounds (condensed tannins, particularly proanthocyanidins, comprising 5-15% seed coat dry weight), cutin and waxes in outer cuticle (reducing water permeability), and various secondary metabolites. Seed coat functions include= mechanical protection of embryo and endosperm from physical damage, regulation of water uptake during imbibition (hard seed coat delays imbibition requiring stratification for gradual softening), gas exchange control (oxygen and carbon dioxide diffusion affecting embryo respiration), chemical protection through phenolic compounds (antimicrobial and antioxidant properties), and dormancy maintenance through mechanical constraint and chemical inhibitors. Seed coat hardness and impermeability in Vitis vinifera contributes to physiological dormancy= hard lignified seed coat provides mechanical resistance to embryo expansion, phenolic compounds may inhibit germination, and reduced permeability delays imbibition requiring stratification for progressive seed coat softening and increased permeability.
Seed_Coat emphasizes the complete protective covering and is distinguished from Hard_Seed_Coat (mechanical strength emphasis) and Protective_Seed_Coat (protective function emphasis) by encompassing all developmental, structural, compositional, and functional aspects. Critical for understanding seed maturity assessment, dormancy mechanisms, germination requirements, wine quality implications of seed coat tannin extraction, and breeding for seed coat traits affecting germination or processing characteristics.
Samenschale
Schutzgewebe des Samens
Seed covering
Testa
Seed Coat
Merlot seed coat adjacent tissue (innermost 2-3 endocarp cell layers, total 50-85 μm thickness) showing characteristic structure= cells 42-58 μm diameter (smallest in pericarp), walls 1.2-1.6 μm thick (among thickest except epidermis 3-6 μm), tannins 6.2-7.8 mg/g (highest in endocarp region), with cells directly contacting seed coat surfaces during early development (BBCH 73-75, weeks 3-6) but separated by 15-35 μm locular cavity air gap at maturity (BBCH 89, week 18) as berry expansion created separation, demonstrating temporal change from seed contact to seed proximity during development affecting potential for direct seed-pericarp interactions. Cabernet Sauvignon seed coat adjacent tissue tannin concentration gradient= innermost cell layer (direct seed contact position) 7.2-8.5 mg/g tannins second layer (20-40 μm from seeds) 6.5-7.8 mg/g third layer (50-80 μm from seeds) 5.2-6.8 mg/g fourth layer (outer endocarp, 90-120 μm from seeds) 4.2-5.5 mg/g, demonstrating systematic exponential-like tannin decline with distance from seeds creating sharp phenolic gradient over 80-120 μm distance potentially supporting hypothesis of seed influence on pericarp tannin accumulation though alternative explanation remains position-dependent tannin biosynthetic regulation. Syrah seed coat adjacent tissue seed number comparison= berries with 2 seeds showing seed coat adjacent tissue zone 55-85 μm thickness with 6.5-7.8 mg/g tannins (lower seed surface area) berries with 4 seeds showing 65-95 μm thickness with 7.2-8.6 mg/g tannins (higher seed surface area creating more extensive seed-adjacent zone), suggesting greater seed presence correlates with enhanced seed coat adjacent tissue development and tannin accumulation though effect modest and variable with biological variability high. Pinot Noir seed coat adjacent tissue developmental time-course examined by microscopy= week 4 post-fruit set (BBCH 73, early Phase I) showing seed coat adjacent tissue cells 38-52 μm in intimate contact with developing seed coats, no air space visible week 8 (BBCH 77, lag phase) showing cells 48-65 μm beginning separation with 2-8 μm gaps appearing week 12 (BBCH 83, veraison) showing 55-70 μm cells separated by 12-25 μm air gaps week 18 (BBCH 89, maturity) showing 52-68 μm cells (slight shrinkage from turgor changes) separated by 20-40 μm gaps, demonstrating progressive locular cavity air space development creating transition from direct seed contact to seed proximity as berry development proceeds. Chardonnay white cultivar seed coat adjacent tissue showing presence in whites= 2-3 cell layers, 55-90 μm thickness, cells 48-62 μm diameter, tannins 4.5-6.2 mg/g (elevated relative to middle mesocarp 0.4-0.9 mg/g similar to red cultivars), demonstrating seed coat adjacent tissue as distinct innermost pericarp zone present in both red and white cultivars with tannin enrichment characteristic though absolute tannin levels lower in whites than reds reflecting overall cultivar differences in phenolic biosynthetic capacity.
A seed coat adjacent tissue is endocarp tissue in direct contact with or immediately adjacent to seed surfaces, comprising the innermost 1-4 cell layers of the endocarp typically 40-120 μm total thickness, characterized by smallest cells within endocarp (40-65 μm diameter), moderately thick to thick walls (1-1.8 μm), highest tannin concentration within endocarp region (5-8 mg/g), and position creating potential for seed-pericarp biochemical interactions during development when seeds contact tissue before locular air space formation, representing the absolute innermost pericarp boundary interfacing with seeds. Seed coat adjacent tissue anatomical characteristics= (1) innermost 1-4 cell layers (typically 2-3 layers in most cultivars) positioned immediately adjacent to seed surfaces or to locular cavity air spaces where seeds are located (2) cells showing smallest dimensions within entire pericarp with diameter 40-65 μm (versus 70-90 μm in outer endocarp, 110-180 μm in middle mesocarp) creating minimum cell size at pericarp-seed boundary (3) walls moderately thick to thick 1-1.8 μm approaching or exceeding outer mesocarp wall thickness (0.8-1.5 μm) and substantially thicker than middle mesocarp walls (0.4-0.8 μm) (4) cells often showing irregular shapes conforming to seed surface contours particularly during development when pericarp directly contacts seeds (5) intercellular spaces sometimes prominent particularly in mature berries as locular cavity development creates partial or complete separation between seed coat adjacent tissue and seed surfaces. Seed coat adjacent tissue compositional characteristics= (1) highest tannin concentration within endocarp region typically 5.5-8 mg/g dry weight (versus 3.5-5.5 mg/g in outer endocarp layers), approaching outer pericarp skin tannin levels (8-15 mg/g) (2) tannin composition predominantly procyanidins with polymerization degree 12-18 subunits and galloylation 12-20% similar to other skin tannins (3) sugar and acid content similar to other pericarp regions (18-26% sugars, 0.5-0.9% acids) showing primary metabolite distribution equilibrates throughout pericarp despite secondary metabolite gradients (4) potential trace seed-derived compounds if diffusion occurs from adjacent seeds during development though quantitative significance uncertain with most seed coat adjacent tissue phenolics likely synthesized in situ rather than imported from seeds. Seed coat adjacent tissue developmental changes in seed proximity= early development (BBCH 71-75, weeks 1-6 post-fruit set) with developing seeds filling locular cavities and seed coats directly contacting seed coat adjacent tissue creating intimate interface potentially allowing compound diffusion or signaling mid development during lag phase (BBCH 77, weeks 6-8) with seeds completing maturation and beginning lignification while pericarp cells continue some expansion late development and ripening (BBCH 81-89, weeks 9-18+) with locular cavity air spaces developing as berry expansion exceeds seed dimensions creating 10-50 μm gaps between mature seeds and innermost seed coat adjacent tissue eliminating direct contact but maintaining close proximity. Seed coat adjacent tissue potential seed interactions= developing seeds producing hormone signals (auxins, gibberellins, ABA) potentially diffusing into adjacent pericarp tissue affecting cell metabolism or gene expression seed phenolic compounds potentially diffusing from seed coats into adjacent pericarp during early development when direct tissue contact exists though evidence limited pericarp-derived signals potentially affecting seed development though mechanisms and significance unclear spatial correlation between high seed coat adjacent tissue tannin levels and seed proximity suggesting possible seed influence on pericarp phenolic accumulation though alternative explanation is position-dependent developmental programming. Seed coat adjacent tissue in wine extraction= representing deepest extractable pericarp tannin reservoir requiring extended vigorous maceration (typically >12-18 days with elevated temperature and frequent cap management) for substantial extraction contributing 8-15% of total wine skin tannins during extended maceration when seed coat adjacent tissue cells break down releasing phenolic contents tannins showing high polymerization and moderate astringency similar to other skin tannins contributing to wine structure and complexity during extended maceration protocols.
Seed_Coat_Adjacent_Tissue represents the innermost endocarp layers in contact with or very close to seeds and is distinguished from more external endocarp layers by position, smallest cell size, and highest endocarp tannin levels. Sub-classification into Immediate (direct contact) and Proximal (very close but not contacting) seed adjacent tissue reflects proximity gradations. Understanding seed coat adjacent tissue is relevant for detailed anatomical characterization, for investigating potential seed-pericarp interactions during development, for comprehending finest-scale spatial tannin distribution affecting wine extraction, and for understanding endocarp as innermost functional boundary of berry flesh.
Innerstes Endokarp
Samenkontakt-Gewebe
Samenschalenangrenzedes Gewebe
innermost endocarp
seed-contact tissue
Cadot, Y. et al. (2006). Tannin localization and characterization in grape skins and seeds. Journal of Agricultural and Food Chemistry 54':2973-2987 Ristic, R., Iland, P.G. (2005). Relationships between seed and berry development of Vitis vinifera. Australian Journal of Grape and Wine Research 11':217-226.
Seed Coat Adjacent Tissue
Cabernet Sauvignon in Napa Valley shows seed coat formation during July-October Year N+1, with ovule integuments differentiating into seed coat following fertilization, outer integument cells developing into testa with radial elongation and secondary wall deposition 2.8-4.2 micrometers thickness, progressive lignification weeks 8-16 creating rigid structure, brown pigmentation developing from tan to dark brown, inner integument forming tegmen 25-45 micrometers thickness, mature seed coat 85-135 micrometers total thickness providing mechanical protection and impermeability with controlled permeability at hilum, hard smooth surface by week 16, under warm climate conditions of 24-30°C temperature, typical of this late-season cultivar completing seed coat maturation
Seed coat formation represents the protective layer development process during seed development when ovule integuments differentiate into seed coat comprising outer testa and inner tegmen layers, progressive lignification creates hard impermeable brown protective structure, and coat maturation produces mechanical protection for embryo and endosperm while regulating water and gas exchange, proceeding from early integument development through lignification to hardened mature coat, spanning approximately 12-16 weeks coordinating with embryo and endosperm development completing by seed maturation. This protective structure formation process occurs during summer to early autumn in July-October in temperate regions Year N+1, representing seed coat development from fertilization through mature seed. Seed coat formation mechanism involves integument development with ovule integuments differentiating into seed coat layers, testa lignification with outer integument lignifying creating hard outer layer, seed coat hardening with progressive structural maturation, pigmentation developing creating brown color, and mature coat providing protection and regulating seed physiology. Integument foundation shows ovule containing two integuments surrounding nucellus and embryo sac, outer integument 3-5 cell layers thick, inner integument 2-4 cell layers thick, integuments as precursors for seed coat development, and integument differentiation initiating following fertilization. Outer integument characteristics shows outer integument giving rise to testa outer seed coat, integument positioned at ovule exterior, cells undergoing differentiation and lignification, and outer layer as primary protective structure. Inner integument characteristics shows inner integument developing into tegmen inner seed coat layer, integument between testa and endosperm, cells differentiating though less lignified than testa, and inner layer as secondary coat component. Seed coat layer structure shows mature seed coat with distinct testa and tegmen layers, testa as outer brown lignified layer providing mechanical protection, tegmen as inner layer with less lignification, and two-layer structure characteristic of grapevine seeds. Testa development shows outer integument cells elongating radially, cells depositing secondary cell walls, lignin accumulating in walls creating rigidity, cells eventually dying leaving lignified walls as protective layer, and testa as major protective component. Testa cell wall thickening shows substantial secondary wall deposition, walls 2-5 micrometers thickness in mature testa, cellulose microfibrils embedded in lignin matrix, and thick walls providing mechanical strength. Lignification process shows phenolic precursors including monolignols synthesized, monolignols polymerized into lignin, lignin deposited in cell walls creating rigid structure, progressive lignification increasing wall rigidity, and lignification as key protective mechanism. Lignin characteristics shows complex phenolic polymer, lignin types including guaiacyl and syringyl lignin, lignin cross-linking with cell wall polysaccharides, and lignin providing impermeability and strength. Lignification timing shows lignification initiating during mid-seed development weeks 8-12, intensive lignification weeks 12-16, completion by seed maturation, and timing coordinating with embryo maturation. Testa pigmentation shows brown pigments including oxidized phenolics accumulating during lignification, pigments creating characteristic brown seed coat color, color darkening with maturation from tan to dark brown, and color as maturation indicator. Pigment development shows phenolic compounds oxidizing during maturation, enzymatic and non-enzymatic oxidation producing brown pigments, pigmentation correlating with lignification and hardening, and color indicating coat maturity. Tegmen development shows inner integument differentiating less extensively than outer, tegmen cells with primary and thinner secondary walls, tegmen less lignified and pigmented than testa, tegmen as inner protective and regulatory layer, and tegmen completing development during maturation. Tegmen function shows possible role in regulating water and solute movement to embryo and endosperm, tegmen as barrier though less impermeable than testa, and inner layer contributing to coat structure. Seed coat thickness shows mature seed coat typically 50-150 micrometers total thickness, testa accounting for 60-80% thickness as major layer, tegmen thinner 20-40% thickness, and coat thickness providing protection while enabling eventual germination. Coat impermeability shows lignified testa creating largely impermeable barrier to water and gases, impermeability preventing pathogen entry and regulating imbibition, controlled permeability at hilum and micropyle regions enabling eventual water uptake, and impermeability as protective function. Hilum characteristics shows hilum as seed attachment scar where funiculus detached, hilum region with less lignification enabling preferential water entry, hilum as water entry point during germination imbibition, and hilum as structural feature. Micropyle characteristics shows micropyle as small opening where pollen tube entered, micropyle region enabling gas exchange, possible water entry point though less than hilum, and micropyle as seed structural feature. Mechanical protection shows hard lignified coat protecting embryo and endosperm from mechanical damage, coat resisting compression and impact, protection essential for seed dispersal and storage, and mechanical strength as major coat function. Pathogen protection shows impermeable coat preventing fungal and bacterial entry, lignified structure resistant to degradation, coat protecting internal tissues during storage and until germination, and pathogen resistance as protection function. Desiccation protection shows coat reducing water loss rate from internal tissues, coat enabling controlled desiccation during seed maturation, coat maintaining low seed moisture during storage, and desiccation control as physiological function. Seed coat maturation shows progressive structural changes during final seed development weeks 12-18, lignification completing, coat hardening increasing rigidity, color darkening to mature brown, and maturation coordinating with embryo and endosperm maturation. Hardening process shows coat becoming increasingly rigid and hard, hardness from lignification and structural changes, mature coat requiring substantial force to fracture, and hardness as maturation outcome. Coat texture shows mature seed coat hard and smooth, coat surface with characteristic pattern from cell organization, texture affecting seed handling, and surface as seed characteristic. Environmental effects on seed coat formation include temperature affecting development rate with optimal 22-26°C supporting normal coat development, nutrient availability particularly phenolic precursors affecting lignification, water availability affecting normal development, and conditions affecting coat quality. Temperature effects shows warm conditions promoting lignification and maturation, cool conditions potentially extending development, and temperature as rate factor. The process encompasses three sequential phases including integument development with ovule integuments differentiating into coat layers, testa lignification with outer layer lignifying and pigmenting, and seed coat hardening completing protective structure. Duration approximately 12-16 weeks representing coat development coordinating with seed maturation. Regional timing shows seed coat formation during July-October as protective layer develops. Viticultural significance includes representing protective structure formation essential for seed viability, enabling seed storage and dispersal, regulating seed hydration and germination, and determining seed physical characteristics. Research applications include analyzing seed coat development and regulation, investigating lignification mechanisms and regulation, evaluating coat permeability and protection functions, quantifying coat structure and composition, and assessing seed coat formation as seed protection and germination regulator.
Seed_Coat_Formation represents protective layer development within Seed_Development. Subdivided into Integument_Development with ovule integuments differentiating (outer → testa, inner → tegmen), Testa_Lignification with outer layer lignifying and pigmenting brown, and Seed_Coat_Hardening completing structure. Duration 12-16 weeks. Mature coat 50-150 μm (testa 60-80%, tegmen 20-40%). Lignified, impermeable (except hilum/micropyle), hard brown. Mechanical and pathogen protection. Coordinates with embryo/endosperm development. Temperature 22-26°C optimal.
Integument-Differenzierung
Samenbedeckungs-Entwicklung
Samenschale-Bildung
Schutzschicht-Formation
coat development
integument differentiation
protective layer formation
seed covering development
Seed Coat Formation
Chardonnay in Champagne shows seed coat hardening during September-October Year N+1, with testa completing lignification reaching 28-34% dry weight lignin, coat rigidity increasing substantially achieving 68-92 N penetration resistance mature hardness, impermeability maximized with controlled permeability at hilum, brown pigmentation reaching mature dark brown color, tegmen completing development 22-38 micrometers thickness with 8-14% lignin, coat desiccating to 12-16% moisture compacting tissues, mature integrated coat 68-118 micrometers total thickness providing maximum mechanical and pathogen protection, under cool maritime conditions of 16-22°C temperature, typical of this early-bursting cultivar completing seed coat maturation by week 16-18
Seed coat hardening represents the final protective structure maturation phase during seed coat formation when lignified testa completes structural maturation, coat rigidity increases to maximum hardness, impermeability finalizes creating fully protective barrier, brown pigmentation reaches mature dark coloration, tegmen inner layer completes development, and integrated coat structure achieves mature protective state by weeks 14-18 enabling seed storage and eventual germination. This final hardening and maturation phase occurs during early-mid autumn in September-October in temperate regions Year N+1, representing seed coat maturation completion coordinating with overall seed maturation. Hardening mechanism involves continued lignin cross-linking increasing structural integrity, possible additional lignin deposition completing accumulation, cell wall maturation and stabilization, coat desiccation removing moisture increasing density and hardness, structural compaction from water loss, and mature coat as maximally protective structure. Lignification completion shows lignin accumulation reaching final levels 25-40% testa dry weight, lignin polymerization completing creating fully cross-linked matrix, lignin distribution throughout testa uniform, and completed lignification as hardening foundation. Lignin cross-linking shows continued cross-link formation between lignin and wall polysaccharides, cross-linking increasing wall rigidity and integration, mature wall as highly cross-linked structure, and cross-linking as hardening mechanism. Cell wall maturation shows walls achieving final structural state, wall components stabilized in mature configuration, walls maximally rigid from lignification and cross-linking, and mature walls as protective structures. Coat rigidity increase shows progressive hardening as maturation proceeds, rigidity increasing substantially during final maturation weeks 14-18, mature coat requiring high force to fracture typically >50-100 N penetration resistance, and maximum rigidity as maturation outcome. Hardness measurement shows penetrometer or compression testing quantifying hardness, hardness increasing 3-5 fold from early to mature hardening, mature seed coat hard and resistant to deformation, and hardness as quantitative maturation indicator. Coat impermeability finalization shows impermeability maximized in mature coat, water permeability very low except at hilum, gas permeability minimal, impermeability preventing uncontrolled water uptake and pathogen entry, and impermeability as protection function. Impermeability testing shows water uptake measurements showing very low permeability, coat preventing rapid imbibition except through hilum, controlled permeability enabling eventual germination, and impermeability as coat characteristic. Hilum characteristics shows hilum region as controlled permeability site, hilum as seed attachment scar, hilum with less lignified cells enabling preferential water entry during germination, and hilum as functional feature. Micropyle characteristics shows micropyle as small pore region, possible gas exchange through micropyle, micropyle contributing to controlled permeability, and micropyle as structural feature. Brown pigmentation completion shows pigmentation reaching maximum intensity, mature seed coat medium to dark brown depending on cultivar, color from accumulated oxidized phenolics, color stable in mature dry seed, and dark brown as mature characteristic. Color stability shows mature pigmentation stable during seed storage, color indicating maturation state, color variation among cultivars from medium brown to nearly black, and color as seed identification feature. Tegmen maturation shows inner integument-derived tegmen completing development, tegmen as inner coat layer 15-40 micrometers thickness, tegmen less lignified than testa 5-15% lignin, tegmen cells possibly remaining alive in mature seed or dying, and tegmen as secondary protective layer. Tegmen structure shows tegmen cells smaller than testa cells, tegmen walls thinner with less secondary wall deposition, tegmen positioned between testa and endosperm, and tegmen as inner boundary. Coat layer integration shows testa and tegmen forming integrated two-layer structure, layers adhering through middle lamella, integrated coat functioning as unified protective structure, and layer integration as structural feature. Total coat thickness shows mature seed coat 50-150 micrometers total thickness varying with cultivar and position, testa accounting for 30-100 micrometers 60-80% total, tegmen 20-50 micrometers 20-40%, and thickness providing protection while enabling germination. Coat desiccation shows coat water content declining during final maturation, water loss increasing coat density, desiccated coat harder and more compact, coat moisture 10-20% in mature dry seed, and desiccation as hardening factor. Structural compaction shows coat tissues compacting as water removed, cell wall spaces reduced, coat density increasing, compaction contributing to hardness, and compaction as maturation change. Coat texture shows mature coat surface hard and smooth, surface with fine texture from cell organization, surface possibly glossy from lipids or cuticle, and texture as seed characteristic. Surface features shows possible surface sculpture or patterns from cell shapes, surface relatively smooth in grapevine, surface affecting seed handling and identification, and surface as morphological feature. Mechanical properties shows mature coat with high tensile strength, compression resistance, impact resistance, mechanical properties enabling seed survival during handling and storage, and mechanical protection as primary function. Protective function completion shows mature coat providing maximum protection from mechanical damage, desiccation, pathogens, and environmental stresses, protection enabling long-term seed storage and survival until germination, and protection as coat primary function. Coat maturation timing shows maturation completing by weeks 14-18 coordinating with embryo and endosperm maturation, timing varying slightly with cultivar and environment, mature coat in mature ready-to-disperse seed, and timing as developmental endpoint. Maturation completion indicators shows coat dark brown color, maximum hardness, complete impermeability, and indicators documenting maturation. Mature seed coat characteristics shows coat as hard rigid brown structure, coat 50-150 micrometers thickness, coat impermeable except at hilum, coat protecting embryo and endosperm, and coat enabling seed viability and storage. Environmental effects on hardening include temperature affecting maturation rate with optimal 20-26°C supporting normal hardening, humidity affecting desiccation with low humidity promoting faster drying and hardening, and environmental conditions affecting final coat properties. Temperature effects shows warm conditions accelerating final maturation, cool conditions potentially extending hardening period, and temperature as rate factor. Humidity effects shows low humidity promoting coat desiccation and hardening, high humidity slowing desiccation, and humidity affecting hardening dynamics. The process represents third and final phase of seed coat formation completing protective structure, following integument development and testa lignification. Duration approximately 4 weeks representing final hardening weeks 14-18. Regional timing shows seed coat hardening during September-October as seed matures. Viticultural significance includes representing seed coat maturation completion, establishing maximum protective structure for seed storage and dispersal, enabling long-term seed viability, and determining final seed physical characteristics affecting handling and germination. Research applications include analyzing hardening mechanisms and regulation, investigating coat mechanical properties and protective functions, evaluating maturation dynamics and completion, quantifying coat structure and composition, and assessing seed coat hardening as seed protection and viability determinant.
Seed_Coat_Hardening is third and final phase of Seed_Coat_Formation, distinguished by final maturation. Represents weeks 14-18. Lignification completion 25-40% DW testa. Rigidity maximum (>50-100 N penetration resistance). Impermeability finalized (except hilum/micropyle). Dark brown pigmentation. Tegmen completion 15-40 μm (5-15% lignin). Coat desiccation 10-20% moisture, compaction. Mature coat 50-150 μm (testa 60-80%, tegmen 20-40%). Follows Testa_Lignification. Completes Seed_Coat_Formation. Maximum protection established.
Samenschale-Verhärtung
Schalen-Reifungs-Vollendung
Schutzstruktur-Vollendung
coat finalization
coat maturation completion
final hardening
finale Verhärtung
protective structure completion
Seed Coat Hardening
The seed color of Merlot at optimal harvest maturity (24-26°Brix, pH 3.5-3.7) documents fully brown to dark brown testa (the desired state for quality red winemaking) — the outer seed coat has completed lignification, the tannin within the testa cells has polymerized to higher molecular weight forms with reduced astringency, and the endosperm tissue has partially desiccated, reducing the diffusion rate of tannin from the seed into the wine during fermentation. The assessment of seed color progression from green (at véraison) through yellow, tan, and brown to dark brown (at full maturity) provides viticulturists with a visual indicator of phenolic maturity that complements the chemical maturity indicators (sugar, acid, pH) — in cool vintages or late-ripening blocks where sugar accumulation outpaces tannin maturation, green seed remnants at harvest indicate incomplete phenolic ripeness that may produce wines with harsh, bitter seed tannin character despite adequate sugar levels.
A seed description component that records the color of the outer seed coat (testa) at harvest maturity, scored on a scale from green-yellow (immature) through yellow-brown, brown, dark brown, to black-brown (fully mature), with seed testa color serving as a practical indicator of seed maturity (lignification status) that determines the extractability and sensory quality of seed tannin — green, immature seeds releasing harsh, aggressive tannins during winemaking, while fully brown-black, mature seeds releasing rounder, less astringent tannins — making seed color assessment a routine harvest decision tool alongside sugar, acid, and berry skin color evaluation.
Seed_Color documents a character that functions both as a cultivar-diagnostic descriptor (the testa color at full maturity varies between cultivars, with some producing uniformly dark brown-black seeds and others retaining lighter brown tones even at full ripeness) and as a ripeness indicator (the progressive browning of the testa during maturation from green through brown to dark brown reflects the sequential biochemical processes of tannin oxidation, tannin polymerization, and testa cell lignification that together determine the sensory quality of extractable seed tannin). The color change results from the oxidation and browning of proanthocyanidin (condensed tannin) polymers within the testa cells — the initial green-yellow color reflects the presence of monomeric and short-chain proanthocyanidins in non-lignified cells, the transition to brown reflects the progressive oxidative polymerization of these tannins to longer-chain, less astringent forms combined with the deposition of lignin in the testa cell walls, and the final dark brown-black color indicates complete lignification and tannin polymerization: this process is not synchronous across all seeds within a berry or across berries within a cluster, necessitating assessment of multiple seeds from multiple berries to determine the overall seed maturity status. The concept of phenolic maturity — the stage at which skin anthocyanin extraction is maximal, skin tannin extraction is optimal, and seed tannin extraction is minimal (achieved when seeds are fully brown and lignified, reducing tannin diffusion) — has become a standard harvest decision criterion in premium red wine production, with seed color being the most accessible visual indicator of the seed tannin maturity component.
Kernfarbe
Samen-Reifefarbe
Testa-Pigmentierung
pip maturity color
seed coat color record
testa pigmentation assessment
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Ristic R and Iland PG (2005) Relationships between seed and berry development of Vitis vinifera L. cv Shiraz, Australian Journal of Grape and Wine Research 11: 43-58 Kennedy JA et al. (2000) Changes in grape seed polyphenols during fruit ripening, Phytochemistry 55: 77-85
Seed Color
Pinot Noir in Burgundy shows seed compartment cell division during early-mid July Year N+1 at BBCH 72-73, with endocarp cells adjacent to 1.4-2.0 developing seeds undergoing limited proliferation creating 2-3 cell layer seed cavity lining, seed compartment cell generation accounting for 3.8-4.6% total berry cells, 1-2 seed cavities 5-7 mm diameter accommodating developing seeds, under continental conditions of 22-27°C temperature, typical of this early-to-mid season cultivar establishing seed-berry interface structure
Seed compartment cell division represents the localized proliferation phase during peak division activity when endocarp cells adjacent to developing seeds and surrounding seed cavity undergo division creating seed compartment structure, accounting for 3-5% total berry cell generation, possibly responding to local auxin maxima from nearby seeds, and establishing specialized tissue region interfacing between seeds and mesocarp. This regionalized division phase occurs during mid summer in early-mid July in temperate regions Year N+1, proceeding during peak division weeks 2-3 post-fertilization, corresponding to BBCH 72-73 young fruit stages. Seed compartment division mechanism involves endocarp inner pericarp layer cells adjacent to seeds undergoing limited proliferation, localized division creating 2-4 cell layer seed cavity lining, possible enhanced division from local auxin maximum near seeds though evidence limited, specialized cell differentiation potentially accompanying division, and seed compartment establishment creating functional interface. Endocarp localization shows tissue positioning at pericarp inner surface directly adjacent to developing seeds, endocarp surrounding individual seeds in seed cavity, tissue contact with seed integuments, and positioning enabling seed-pericarp communication. Endocarp cell division characteristics show limited division compared to abundant mesocarp proliferation, typical 2-4 cell layer thickness from restricted proliferation, periclinal divisions increasing radial thickness, and cell generation accounting for 3-5% total berry cells. Seed cavity formation shows localized space development surrounding each seed, cavity lined by endocarp tissue, cavity size accommodating seed growth, and cavity structure from coordinated endocarp division and mesocarp displacement. Seed cavity architecture shows typically 2-4 cavities per berry corresponding to seed number, cavity diameter 4-8 mm accommodating 4-6 mm mature seeds, endocarp lining providing protective interface, and cavity organization reflecting seed positioning and development. Local auxin effects hypothesis shows seeds as auxin source with highest concentration in immediate vicinity, possible auxin gradient with maximum at seed surface declining into mesocarp, local auxin potentially stimulating nearby endocarp division though direct evidence limited, and gradient possibly affecting regional proliferation though mesocarp also highly proliferative. Endocarp versus mesocarp proliferation comparison shows mesocarp 20-30 fold greater cell generation than endocarp, mesocarp extensive 15-25 cell layer thickness versus endocarp limited 2-4 layers, and differential proliferation creating tissue-specific cellularity. Endocarp cell specialization shows potential differentiation into transfer cells or modified parenchyma, possible structural specialization for seed-pericarp interface function, cell wall modifications possibly enabling nutrient or signal exchange, and specialized cells providing functional interface though poorly characterized in grapevine. Seed development coordination shows seeds developing in parallel with berry division phase, seed size increase correlating with berry growth, seed maturation largely completing by division phase end, and seed-berry coordination ensuring developmental synchronization. Seed size during division phase shows seeds growing from 2-3 mm at division onset to 4-6 mm by completion, seed expansion coordinated with berry growth, and seed cavity accommodating seed enlargement. Endocarp-seed interface functions hypothetically include nutrient transfer from pericarp to developing seed, hormone communication from seed to berry affecting growth, mechanical support and protection for seeds, and signal integration coordinating seed and berry development. Cell division near funiculus shows possible enhanced division near vascular connection, funiculus providing nutrient and hormone transport, and funiculus region potentially showing specialized division patterns. Endocarp division timing shows division coinciding with peak mesocarp proliferation weeks 2-3, coordinated timing across pericarp regions, and endocarp completion concurrent with overall division phase end. Division completion and maturation shows endocarp division completing by week 4-5, cells transitioning to expansion and differentiation, mature endocarp as thin protective layer around seeds, and completed endocarp structure persisting through berry development and ripening. Environmental effects on seed compartment division include temperature affecting division rate similarly to other pericarp regions, seed development providing proliferative stimulus through auxin production, and overall favorable conditions supporting endocarp division though representing small fraction of total proliferation. Seed number effects show more seeds creating more seed compartments and endocarp tissue, total endocarp cellularity correlating with seed number, and multiple seeds increasing localized proliferation though each compartment similar. Seed compartment assessment methods include histological sectioning showing endocarp tissue adjacent to seeds, cell counting in seed cavity regions, seed cavity size and structure measurement, and endocarp architecture analysis. The process represents third and final aspect of peak division activity emphasizing seed region specialization, following maximum cell number generation and pericarp cell proliferation. Duration concurrent with peak division weeks 2-3 representing localized seed compartment proliferation. Regional timing shows seed compartment cell division during early-mid July across climates during peak berry division phase. Viticultural significance includes establishing seed compartment structure providing seed-berry interface, representing specialized regional proliferation creating tissue heterogeneity, accounting for small but potentially functionally important cell population, and demonstrating coordinated tissue-specific division patterns. Research applications include analyzing endocarp division regulation and specialization, investigating seed-berry communication and interface functions, evaluating local auxin effects on regional proliferation, quantifying seed compartment architecture and cellularity, and assessing endocarp as specialized berry tissue with potential functional significance.
Seed_Compartment_Cell_Division is third and final aspect of Peak_Division_Activity, distinguished by localized endocarp proliferation near seeds. Represents specialized regional division accounting for 3-5% total cells. Limited 2-4 cell layer endocarp thickness contrasting with abundant mesocarp. Possibly responding to local seed auxin though poorly characterized. Creates seed cavity structure and seed-berry interface. Duration concurrent with peak weeks 2-3. Completes Peak_Division_Activity documentation. Specialized tissue-specific proliferation.
Endokarp-Proliferation
Samenfach-Zellteilung
Samenkavität-Bildung
Samenregion-Zellteilung
endocarp proliferation
localized seed area division
seed cavity formation
seed region cell division
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 437-438
Seed Compartment Cell Division
Seed density analysis in Merlot reveals values of 1.089±0.021 g/mL at véraison increasing to 1.147±0.018 g/mL at harvest, with the 5.3% density increase correlating with seed coat browning from chlorophyll degradation and tannin oxidation, providing winemakers with a visual proxy for seed tannin extractability where browner, denser seeds release less harsh tannins during fermentation.
Seed density quality is a tissue density quality that inheres in grape seeds and represents the mass per unit volume of this reproductive structure characterized by high density resulting from extensive lignification of the seed coat, accumulation of storage reserves including oils and proteins in the endosperm, and the compact cellular organization required for embryo protection and long-term viability. Within the BFO framework, this quality distinguishes seeds as the highest-density component of the grape berry, typically ranging from 1.10-1.20 g/mL compared to 1.05-1.10 g/mL for pulp tissue, reflecting the fundamentally different developmental program and biochemical composition of reproductive versus vegetative berry tissues. The ontological significance extends to seed density serving as a maturity indicator independent of sugar accumulation, where progressive lignification during berry development increases seed coat density while internal reserves consolidate, providing a structural marker of physiological seed maturity. Seed density stabilizes relatively early in berry development compared to ongoing pulp density changes, with maximum density typically achieved by véraison as testa lignification completes, though internal reserve redistribution may continue through maturation. Measurement approaches for seed density require careful extraction without damage to the seed coat, followed by pycnometry, density gradient flotation, or calculated approaches from mass and geometric volume estimation. The contribution of seeds to whole-berry density depends on seed number (typically 0-4 per berry with cultivar variation), individual seed mass (25-75 mg), and the volume proportion of seeds relative to total berry volume (2-6%). Understanding seed density supports interpretation of whole-berry flotation behavior and provides insight into seed maturation timing relative to pulp ripening progression.
Seed_Density_Quality addresses the reproductive structure specifically, recognizing its distinct developmental program and compositional profile compared to pericarp tissues. This quality relates to seed maturity assessment and tannin management in winemaking, where seed density and color together indicate extraction potential. The class excludes seedless (stenospermic) varieties where rudimentary seed traces have different density characteristics.
Kernendichte
Saatgutdichte
Samendichte
Traubenkernendichte
grape seed compactness
grape seed specific gravity
pip density
seed mass density
Ristic R, Iland PG (2005) Relationships between seed and berry development of Vitis vinifera L. cv Shiraz: developmental changes in seed morphology and phenolic composition. Australian Journal of Grape and Wine Research 11(1): 43-58 Cadot Y et al. (2006) Anatomical, histological, and histochemical changes in grape seeds from Vitis vinifera L. cv Cabernet franc during fruit development. Journal of Agricultural and Food Chemistry 54(24): 9206-9215
Seed Density Quality
The seed description of Cabernet Sauvignon documents 1-3 well-developed seeds per berry (mean 1.8), with individual seeds measuring 6-7 mm length and 4-5 mm width, pyriform (pear-shaped) outline with a prominent pointed beak (rostrum) of 1.5-2.0 mm, dark brown to black testa color at harvest maturity indicating full lignification of the outer seed coat, and a deeply grooved ventral chalazal depression — the fully mature, darkly colored seeds contribute 20-40% of the total wine tannin through extraction during fermentation, with seed tannin character (shorter-chain, more astringent proanthocyanidins dominated by catechin subunits) becoming more extractable as the testa lignifies and the surrounding endosperm tissue desiccates during the final weeks of berry ripening.
A berry description component that documents the morphological characteristics of the grape seeds (pips) contained within the berry — the mature ovules consisting of the outer and inner seed coats (testa), the endosperm, and the embryo — recording number, size, shape, color, surface texture, and the beak (rostrum) morphology that provide cultivar-diagnostic characters and determine the seed tannin contribution to wine structure and astringency.
Seed_Description addresses the reproductive structures within the berry that have dual significance — biologically, the seeds are the products of sexual reproduction containing the embryo and the genetic recombination that generates the diversity from which new cultivar genotypes can be selected enologically, the seeds are a major source of tannin (proanthocyanidins) that contributes to wine structure, astringency, and aging potential. Each grape berry contains a maximum of four seeds (two per carpel in the bicarpellate ovary), but the actual seed number per berry varies from zero (in seedless cultivars or parthenocarpic berries) to four, with the mean seed number cultivar-specific and influenced by pollination conditions: the seed number directly affects berry size (each developing seed produces growth hormones that promote surrounding tissue development, so more seeds produce larger berries) and wine composition (more seeds increase the potential tannin extraction per berry, but larger berries dilute the skin-derived compounds). The morphological characters of the seed — shape, size, beak prominence, testa color, and surface texture — are among the most genetically stable ampelographic characters because the seed is protected within the berry from environmental modification, and seed morphology has been used since antiquity for cultivar identification in archaeological contexts where seed remains in waterlogged or charred deposits are the only evidence of cultivar identity.
Kern-Beschreibung
Samen-Phänotyp-Dokumentation
Traubenkern-Morphologie
grape seed morphology
pip characterization
seed phenotype documentation
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Ristic R and Iland PG (2005) Relationships between seed and berry development of Vitis vinifera L. cv Shiraz, Australian Journal of Grape and Wine Research 11: 43-58
Seed Description
Cabernet Sauvignon in Napa Valley shows seed development during July-October Year N+1, with double fertilization initiating development late June, embryo progressing through globular stage weeks 1-3, heart stage weeks 3-7, torpedo stage weeks 7-12, mature embryo weeks 12-16 with 3.8-4.6 mm length, endosperm developing through free nuclear proliferation weeks 0-2 and cellularization weeks 2-4, storage reserves accumulating 32-38% lipids and 26-32% proteins in cotyledons, seed desiccating from 58% to 12-16% moisture, seed coat lignifying creating hard dark brown protective layer, mature seed 5.2-6.4 mm length weighing 32-46 mg by 14-16 weeks post-fertilization, 1.8-2.4 seeds per berry average, under warm climate conditions coordinating seed maturation with berry ripening, typical of this late-season cultivar achieving complete seed development
Seed development represents the comprehensive reproductive maturation process in grapevine when fertilized ovule develops into mature viable seed containing embryo and endosperm within protective seed coat, proceeding from double fertilization through embryo morphogenesis, endosperm development, storage reserve accumulation, and seed maturation including desiccation and dormancy acquisition, spanning approximately 90-120 days from fertilization to mature seed ready for dispersal, with seed development coordinating with berry development ensuring reproductive success before fruit ripening. This complete seed maturation process occurs during summer to early autumn in Year N+1 from fertilization in late June-July through maturation in September-October in temperate regions, encompassing entire seed development from zygote through mature seed, representing critical reproductive phase establishing next generation. Seed development encompasses sequential coordinated phases including embryo development with zygote division initiating embryogenesis, embryo morphogenesis creating organized seedling structures through globular, heart, and torpedo stages, and embryo maturation completing development with storage reserve accumulation, alongside parallel endosperm development with endosperm proliferation through free nuclear and cellular phases, nutrient accumulation establishing storage tissues, and endosperm maturation supporting embryo development and providing germination reserves. Embryo development characteristics show zygote undergoing first division 24-72 hours post-fertilization producing two-cell proembryo, embryo morphogenesis proceeding through globular stage weeks 1-3 with undifferentiated cell mass, heart stage weeks 3-7 with cotyledon primordia emergence, torpedo stage weeks 7-12 with elongated embryonic axis and cotyledon development, and mature embryo stage weeks 12-18 with fully formed structures including radicle, hypocotyl, cotyledons, and shoot apex. Endosperm development characteristics show triploid endosperm 3n=57 initiating free nuclear division immediately post-fertilization, multinucleate syncytium forming through rapid nuclear divisions without cytokinesis, cellularization occurring weeks 2-4 creating cellular endosperm, and endosperm tissue providing nutritive support for embryo development and accumulating storage reserves. Seed coat development shows integuments differentiating into seed coat layers, outer integument forming testa outer seed coat layer, inner integument forming tegmen inner layer, progressive lignification creating hard protective coat, and brown pigmentation developing indicating maturation. Double fertilization foundation shows two sperm cells from pollen tube fusing independently with egg cell creating 2n=38 diploid zygote and with central cell creating 3n=57 triploid endosperm, double fertilization as angiosperm characteristic providing embryo and nutritive tissue, and fertilization initiating coordinated seed development. Developmental timeline shows fertilization at 0 days establishing starting point, early embryo development weeks 1-3 with rapid cell divisions, mid embryo morphogenesis weeks 3-12 with structure formation, late embryo maturation weeks 12-18 with reserve accumulation, and seed maturation completing by 90-120 days depending on cultivar and environment. Embryo growth pattern shows initial slow growth during early divisions, rapid growth during morphogenesis weeks 3-12, growth deceleration during maturation weeks 12-18, and final mature embryo size 3-5 mm length within seed. Endosperm temporal dynamics shows rapid initial proliferation weeks 0-4 establishing tissue, nutrient accumulation weeks 4-12 building reserves, possible partial endosperm consumption by embryo in some species though extent unclear in grapevine, and mature endosperm as storage tissue surrounding embryo. Storage reserve accumulation shows lipids accumulating primarily in cotyledons reaching 30-40% dry weight, proteins accumulating 25-35% dry weight, minor carbohydrates, and reserves supporting germination and seedling establishment. Lipid accumulation shows triacylglycerols as major storage lipids, oleic and linoleic acids as predominant fatty acids, lipid bodies forming in cotyledon cells, and lipids providing energy reserves for germination. Protein accumulation shows storage proteins including albumins and globulins accumulating in protein bodies, proteins providing amino acids for seedling growth, and protein as major reserve component. Seed maturation processes show progressive desiccation reducing moisture from 50-70% mid-development to 10-20% mature seed, metabolic activity declining during desiccation, desiccation tolerance developing enabling seed survival, dormancy possibly imposed preventing premature germination, and hard dry seed as mature state. Seed desiccation characteristics show water loss through seed coat and funiculus, progressive dehydration over final weeks of development, desiccation tolerance mechanisms including protective proteins and sugars, and low moisture enabling storage and dormancy. Seed coat maturation shows lignification creating hard impermeable brown coat, mechanical protection for embryo and endosperm, coat impermeability preventing pathogen entry and regulating water uptake, and mature coat as protective structure. Seed coat color shows progression from green immature to brown maturing to dark brown mature seed, color change from pigment deposition and oxidation, brown color indicating lignification and maturation, and color as convenient maturity indicator. Seed size variation shows cultivar variation with typical mature seed 4-7 mm length and 3-5 mm width, seed fresh weight 20-50 mg depending on development success, within-berry variation from differential ovule fertilization and development, and size affecting germination vigor. Seed number per berry shows 0-4 ovules per flower with typically 0-3 developing into mature seeds, variable seed number from fertilization success and developmental survival, seed number affecting berry development through hormone production particularly during early growth, and average seed number 1.5-2.5 per berry across populations. Seed-berry developmental coordination shows seeds developing within berry throughout berry growth and ripening, seed maturation completing during or shortly before berry ripening, mature seeds in ripe berry ready for dispersal, and coordinated development ensuring reproductive success with fruit development. Hormonal signaling from seeds shows seeds producing auxin during early development stimulating berry growth, auxin as critical signal preventing berry abscission and promoting expansion, hormone production declining as seeds mature, and seed hormones coordinating seed-berry development. Seed maturation timing shows maturation completing by late lag phase to early ripening weeks 12-18 post-fertilization, timing varying with cultivar and environment, mature seeds enabling berry progression through lag phase and veraison, and maturation as potential developmental checkpoint. Seed viability shows mature seeds capable of germination if separated and provided appropriate conditions, germination requiring stratification cold treatment for dormancy release in many cultivars, viable seeds representing reproductive success, and viability as seed development outcome. Seed dispersal preparation shows mature seed in ripe berry ready for consumption by frugivores, seed coat protecting embryo during passage through digestive tract, animal dispersal as primary natural mechanism, and seed-fruit syndrome as coevolved dispersal strategy. Environmental effects on seed development include temperature affecting development rate with warm conditions accelerating and cool conditions slowing progression, water availability important for normal development particularly during growth phases, nutrient supply supporting storage reserve accumulation, and overall favorable conditions ensuring complete seed development. Temperature effects show heat accumulation driving developmental progression, growing degree days determining development timing, optimal ranges 22-28°C for normal development, and temperature as major rate determinant. Seed development failure shows some ovules failing to develop into mature seeds from fertilization failure, early embryo abortion, inadequate resources, or developmental abnormalities, failed ovule frequency creating variable seed number 0-3 per berry, and seed failure affecting berry development and size. Seedless berries shows parthenocarpic seedless cultivars developing berries without fertilization or with aborted seed traces, seedless development lacking hormonal stimulation from seeds requiring exogenous hormones for berry development, and seedless berries as special case. The process encompasses two major parallel developmental programs including embryo development with zygote division, embryo morphogenesis, and embryo maturation creating seedling structures, and endosperm development with endosperm proliferation, nutrient accumulation, and endosperm maturation providing nutritive support. Duration typically 90-120 days from fertilization to mature seed varying by cultivar and environment. Regional timing shows seed development during July-October in temperate regions coordinating with berry development. Viticultural significance includes representing reproductive success ensuring next generation, coordinating with berry development through hormonal signaling particularly during early berry growth, mature seeds possibly influencing berry developmental progression through lag phase, and seed development as fundamental reproductive process. Research applications include analyzing seed developmental regulation and coordination, investigating embryo and endosperm developmental programs, evaluating seed-berry communication and coordination, quantifying storage reserve accumulation and composition, and assessing seed development as reproductive success indicator and berry development coordinator.
Seed_Development represents complete reproductive maturation from fertilization to mature seed. Subdivided into Embryo_Development with Zygote_Division, Embryo_Morphogenesis (globular, heart, torpedo stages), and Embryo_Maturation with reserve accumulation and Endosperm_Development with Endosperm_Proliferation (free nuclear, cellular), Nutrient_Accumulation, and Endosperm_Maturation. Duration 90-120 days. Mature seed 4-7 mm, 20-50 mg. Storage reserves 30-40% lipids, 25-35% proteins. Desiccation 50-70% to 10-20% moisture. Seed coat lignified brown. Coordinates with berry development. Critical reproductive process.
Samenentwicklung
Samenreifung
ovule to seed development
reproductive development
reproduktive Entwicklung
seed maturation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 438-442 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Seed Development
Pinot Noir in Burgundy shows seed development focus during late August-September Year N+1 at BBCH 81, with seeds completing maturation during 2.5 week minimal growth period, seed coat lignification creating hard dark brown protective layer, embryo achieving full developmental maturity with viable embryonic axis, seed moisture declining from 42% to 14%, final mature seeds 4.8-6.2 mm length and 3.2-4.6 mm width weighing 28-42 mg, 1.6-2.2 mature seeds per berry average, under continental conditions of 20-26°C temperature, typical of this early-to-mid season cultivar completing seed maturation during lag
Seed development focus represents the seed maturation completion phase during minimal growth period when seeds finalize developmental maturation achieving fully viable state, seed coat completes lignification creating hard protective brown layer, embryo and endosperm complete development achieving germination capacity, seed desiccation proceeds reducing moisture content, and mature seeds possibly signal berry developmental readiness for veraison progression, occurring during lag phase as critical reproductive maturation process. This seed maturation phase occurs during late summer to early autumn in late August-September in temperate regions Year N+1, proceeding over 1-3 weeks during minimal growth period, corresponding to BBCH 81 beginning of ripening stage with seed maturation completing. Seed focus mechanism involves seeds completing final developmental processes during berry growth quiescence, developmental resources allocated to reproductive maturation, seed maturation as prerequisite for berry ripening progression, mature seeds possibly producing hormonal or developmental signals enabling veraison, and seed completion as critical lag phase function. Seed developmental state during lag shows seeds at advanced maturity at lag phase initiation, completing final maturation during minimal growth period, achieving full developmental maturity by mid-to-late lag phase, and mature seeds ready for dispersal if separated from berry. Seed coat maturation shows lignification completing during minimal growth period, lignin deposition creating hard protective layer, seed coat color progression from brown to dark brown to black indicating maturation, coat impermeability increasing preventing water and pathogen entry, and mature coat as seed protection. Lignification process shows phenolic compounds polymerizing into lignin, cross-linking creating rigid structure, coat hardening progressively during lag, and lignification as major maturation process. Seed coat color changes shows progression from green immature seeds to brown early maturation to dark brown late maturation, color darkening correlating with lignification and maturation degree, final dark brown to black color indicating full maturity, and color as convenient maturation indicator. Embryo maturation shows embryonic axis completing development during lag phase, cotyledons filling with storage reserves, embryo achieving developmental capacity for germination, viable embryo in mature seed ready for germination if environmental conditions suitable, and embryo maturation as reproductive completion. Embryo developmental completion shows radicle, hypocotyl, cotyledons fully formed, meristems established in embryonic axis, developmental program complete awaiting germination activation, and structural maturation achieved. Storage reserve accumulation shows cotyledons accumulating proteins, lipids, some carbohydrates as energy reserves, typical seed reserves 30-40% lipids, 25-35% proteins, stored compounds supporting germination and seedling establishment, and reserve accumulation as maturation function. Endosperm maturation shows endosperm tissue completing development during lag, nutrient storage in endosperm, possible partial endosperm consumption by developing embryo in some cases, and endosperm contribution to seed maturation and germination capacity. Seed desiccation shows progressive moisture loss during maturation, water content declining from 50-70% immature to 10-20% mature seeds, desiccation tolerance developing enabling seed survival, reduced metabolism in desiccated state, and desiccation as maturation characteristic. Seed moisture decline shows progressive dehydration during lag phase, water loss through seed coat and funiculus, eventual low moisture stable state, and moisture decline indicating maturation progression. Seed texture changes shows softening hard texture progression during maturation, immature seeds soft and pliable, maturing seeds progressively hardening, mature seeds hard and resistant to deformation, and texture as maturation indicator. Seed size during lag shows seeds at or approaching final mature dimensions, typical mature seed size 4-7 mm length and 3-5 mm width, seed fresh weight 20-50 mg depending on seed development success, and size established during earlier berry development maintained during lag. Seed number per berry shows typical 0-3 mature seeds per berry depending on fertilization success and development, seed number affecting berry development through hormone production during growth but less critical during lag, and variable seed number characteristic of grapevine. Seed maturation completion timing shows seeds achieving full maturity by mid-to-late lag phase typically 1-2 weeks into minimal growth period, maturation timing varying with cultivar and environment, and completion enabling berry developmental progression. Seed maturity indicators shows hard dark brown to black seed coat, low moisture content 10-20%, germination capacity if tested, and indicators documenting mature state. Seed-berry developmental coordination shows seed maturation possibly signaling berry readiness for ripening, hormonal signals from seeds potentially regulating berry development, abscisic acid from seeds possibly contributing to veraison triggering, and coordination ensuring reproductive and dispersal preparation. Seed hormone production shows mature seeds possibly producing or reducing hormone levels affecting berry, abscisic acid possibly increasing from maturing seeds, gibberellin possibly declining, and hormonal signals as coordination mechanism. Seed maturation as veraison prerequisite shows hypothesis that seed maturation completion enables veraison progression, immature seeds possibly preventing premature ripening, mature seeds permitting developmental progression, and seed maturity as gating factor though evidence partially correlational. Seed maturation in seedless berries shows seedless parthenocarpic berries lack seed maturation but still undergo lag and veraison, seed maturation not absolutely required though may influence timing, seedless berry development showing seed-independent progression, and complexity of seed-berry coordination. Berry resources during seed focus shows minimal berry growth enabling metabolic resources for seed completion, berry quiescence not requiring growth resources, coordination as resource allocation strategy, and functional interpretation of lag phase. Environmental effects on seed maturation include temperature affecting maturation rate with warm conditions promoting faster completion and cool conditions extending maturation period, water availability affecting seed development though less critical than during earlier phases, and environmental factors influencing maturation timing. Temperature effects show heat accumulation advancing seed maturation, growing degree days correlating with completion timing, and temperature as maturation rate determinant. Seed maturation assessment shows seed coat color observation, hardness testing, moisture content measurement, germination testing, and assessments characterizing maturation state. The process represents second aspect of minimal growth period emphasizing seed maturation completion, between near-zero growth rate and pre-veraison metabolism. Duration proceeding over 1-3 weeks during minimal growth period. Regional timing shows seed development focus during late August-September across climates as seeds mature during lag. Viticultural significance includes representing critical reproductive maturation completing seed development, possibly gating berry developmental progression to ripening, demonstrating seed-berry developmental coordination, and ensuring reproductive success before fruit ripening and dispersal. Research applications include analyzing seed maturation processes and regulation, investigating seed-berry communication and coordination, evaluating seed maturation as veraison prerequisite or correlate, quantifying maturation timing and environmental effects, and assessing seed development focus as functional explanation for lag phase quiescence.
Seed_Development_Focus is second aspect of Minimal_Growth_Period, distinguished by seed maturation completion. Represents final lignification (hard dark brown coat), embryo maturation (viable germination capacity), desiccation (50-70% to 10-20% moisture). Mature seeds 4-7 mm, 20-50 mg. Possible hormonal signaling to berry. Maturation as potential veraison prerequisite. Duration 1-3 weeks during lag. Between Near_Zero_Growth_Rate and Pre_Veraison_Metabolism. Critical reproductive completion.
Samenentwicklungs-Abschluss
Samenentwicklungs-Fokus
Samenreifungs-Vollendung
reproductive maturation
reproduktive Reifung
seed development completion
seed finalization
seed maturation completion
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 456-460 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Seed Development Focus
Cabernet Sauvignon in Napa Valley shows seed dormancy acquisition during late September-October Year N+1, with embryo developing deep dormancy preventing immediate germination, abscisic acid accumulating to 420-680 ng/g dry weight in mature embryo during weeks 14-18, germination inhibitor accumulation and metabolic blocks establishing, mature seed requiring 85-110 days cold stratification at 2-4°C for complete dormancy release, unstratified seeds showing <5% germination while stratified seeds germinating 78-92%, dormancy as adaptation preventing autumn germination ensuring winter passage before spring germination, under warm climate conditions of 22-28°C maturation temperature, typical of this late-season cultivar establishing dormancy for germination timing control
Seed dormancy acquisition represents the germination control mechanism establishment phase during seed maturation when embryo develops dormancy preventing immediate germination after maturation, dormancy requiring cold stratification typically 60-120 days at 1-5°C for release, abscisic acid possibly accumulating in embryo imposing or maintaining dormancy, metabolic blocks preventing germination established, dormancy as adaptation preventing premature germination in unfavorable conditions, and dormancy acquisition occurring during late maturation weeks 14-18 completing seed maturation and establishing germination control. This germination control establishment phase occurs during early-mid autumn in late September-October in temperate regions Year N+1, representing embryo dormancy development. Dormancy acquisition mechanism involves embryo physiological state changes preventing germination, possible abscisic acid accumulation imposing dormancy, germination inhibitor accumulation or germination promoter deficiency, metabolic blocks preventing embryo growth, developmental arrest maintaining dormancy, and integrated dormancy mechanisms preventing premature germination. Seed dormancy definition shows inability to germinate under normally favorable conditions despite being viable, dormancy as block preventing germination even when water, oxygen, temperature adequate, dormancy requiring specific treatments for release, and dormancy as regulated developmental state. Dormancy types shows embryo dormancy as primary type in grapevine with embryo unable to germinate, coat-imposed dormancy possible but generally secondary from hard impermeable coat, physiological dormancy as classification with embryo physiological block, and dormancy type affecting release requirements. Embryo dormancy characteristics shows embryo in dormant non-germinative state in mature seed, embryo morphologically complete and viable but unable to grow, dormancy residing in embryo tissue not primarily coat, and embryo dormancy as grapevine characteristic. Dormancy depth shows dormancy intensity varying among cultivars, deep dormancy requiring extended stratification 90-120 days, intermediate dormancy requiring moderate stratification 60-90 days, weak dormancy with limited requirements 30-60 days or germinating readily, and dormancy depth as genetic variation. Cultivar variation shows different grapevine cultivars exhibiting varying dormancy depths, Vitis vinifera typically showing intermediate to deep dormancy, wild Vitis species possibly showing different patterns, and genetic control of dormancy intensity. Abscisic acid role shows ABA as hormone regulating dormancy in many species, ABA possibly accumulating during seed maturation in grapevine, ABA levels in mature seed possibly 200-800 ng/g dry weight, ABA imposing or maintaining dormancy through metabolic effects, ABA inhibiting germination-related processes, and ABA as major dormancy regulator though evidence in grapevine incomplete. ABA accumulation timing shows ABA synthesis possibly increasing during late maturation, ABA content rising weeks 14-18, peak ABA in mature dormant seed, and timing coordinating with dormancy establishment. ABA biosynthesis shows ABA synthesized through carotenoid pathway, zeaxanthin epoxidase and 9-cis-epoxycarotenoid dioxygenase NCED as key enzymes, biosynthesis in embryo and possibly other seed tissues, and synthesis as ABA accumulation mechanism. ABA effects on embryo shows ABA inhibiting embryo axis growth, ABA suppressing germination-related gene expression, ABA maintaining metabolic quiescence, and ABA as germination inhibitor. Germination inhibitor accumulation shows possible accumulation of other inhibitory compounds in embryo or seed, inhibitors preventing germination until removed or inactivated, and inhibitors as potential dormancy factors though less studied than ABA. Germination promoter deficiency shows germination potentially requiring gibberellins, cytokinins, other promoters, dormant embryo deficient in promoters, promoter deficiency contributing to germination block, and promoter balance as dormancy mechanism. Metabolic blocks shows metabolic pathways required for germination blocked in dormant embryo, blocks including respiratory limitations, biosynthetic capacity limitations, cell cycle progression blocks, and metabolic blocks preventing embryo growth. Respiratory capacity shows dormant embryo with limited respiratory capacity, respiration increasing during stratification and germination, respiratory limitation as potential dormancy mechanism, and respiration as metabolic indicator. Cell division block shows dormant embryo unable to resume cell division and growth, cell cycle arrest maintaining dormancy, release enabling cell cycle progression, and division block as dormancy mechanism. Developmental arrest shows embryo development paused in mature state, arrest preventing germination, arrest reversible through dormancy release, and arrest as dormancy state. Dormancy acquisition timing shows dormancy developing during late maturation weeks 14-18, dormancy established by seed maturation completion, dormant state in mature seed at dispersal, and timing as developmental endpoint. Stratification requirement shows cold stratification as primary dormancy release treatment, stratification typically 60-120 days at 1-5°C for complete dormancy release, cold treatment enabling subsequent germination at warm temperature, and stratification as release mechanism. Stratification effects shows stratification causing ABA decline and gibberellin increase, metabolic capacity increasing, embryo transitioning from dormant to germinative state, and stratification enabling subsequent germination. After-ripening possible role shows dry storage after-ripening possibly contributing to dormancy release in some seeds, after-ripening less important than stratification in grapevine, and after-ripening as additional factor. Coat-imposed dormancy possible shows hard seed coat possibly contributing to dormancy through mechanical resistance or impermeability, coat effects generally secondary to embryo dormancy in grapevine, scarification or coat removal not typically required though may accelerate germination, and coat as minor dormancy factor. Dormancy adaptive significance shows dormancy preventing germination immediately after seed maturation and dispersal in autumn, dormancy ensuring germination delayed until favorable spring conditions after winter cold, cold requirement ensuring winter passage before germination, and dormancy as adaptation to temperate climate seasonality. Ecological role shows dormancy synchronizing germination with spring after winter, dormancy preventing germination in autumn before harsh winter, and dormancy as survival mechanism. Dormancy assessment shows germination testing determining dormancy state, dormant seeds failing to germinate under favorable conditions, stratified seeds germinating readily, and germination as dormancy indicator. Germination after stratification shows stratified seeds germinating 70-95% under favorable conditions 20-25°C with moisture, germination demonstrating dormancy release, and germination as functional test. Environmental effects on dormancy acquisition include temperature during seed maturation possibly affecting dormancy depth, maturation conditions influencing ABA accumulation, and environmental factors affecting dormancy establishment. Temperature effects shows warm maturation possibly promoting dormancy development, temperature affecting dormancy depth, and temperature as maturation factor. The process represents second aspect of seed maturation establishing germination control, following embryo desiccation tolerance and preceding seed ripening completion. Duration dormancy acquisition during weeks 14-18 representing germination control establishment. Regional timing shows seed dormancy acquisition during late September-October as maturation completes. Viticultural significance includes representing germination timing control preventing premature germination, demonstrating adaptation to temperate seasonality, requiring stratification for seed propagation, and affecting germination success in viticulture and breeding. Research applications include analyzing dormancy mechanisms and regulation, investigating hormonal control particularly abscisic acid and gibberellins, evaluating stratification effects and release mechanisms, quantifying dormancy depth variation, and assessing dormancy acquisition as germination control determinant.
Seed_Dormancy_Acquisition is second aspect of Seed_Maturation, distinguished by germination control establishment. Represents weeks 14-18. Embryo physiological dormancy (primary type). ABA accumulation possibly 200-800 ng/g. Metabolic blocks preventing germination. Requires stratification 60-120 days at 1-5°C for release. Dormancy depth varies by cultivar (weak to deep). Coat-imposed dormancy secondary. Adaptive significance: prevents premature germination, ensures spring germination. Follows Embryo_Desiccation_Tolerance. Precedes Seed_Ripening_Completion. Critical germination timing control.
Dormanz-Entwicklung
Keimungs-Blockierung
Samen-Dormanz-Erwerb
dormancy development
germination block
germination control
Seed Dormancy Acquisition
Pinot Noir in Burgundy shows seed maturation during September-October Year N+1, with embryo completing reserve accumulation 33-39% lipids and 28-34% proteins in cotyledons, endosperm completing 22-27% proteins and 14-20% lipids, LEA proteins accumulating providing desiccation tolerance, protective sugars sucrose 8-13% and raffinose 2-4% dry weight, progressive desiccation reducing water content from 58% to 12-16%, respiratory rate declining from 1.8 to <0.2 μmol O₂ g⁻¹ h⁻¹, embryo dormancy established requiring 75-100 days stratification, abscisic acid 340-620 ng/g, seed coat hardening completing, mature seed 4.2-5.8 mm length weighing 24-38 mg, viable and ready for dispersal, under continental conditions of 18-24°C temperature, typical of this early-to-mid season cultivar completing seed maturation coordinating with berry ripening
Seed maturation represents the final seed development phase when embryo and endosperm complete reserve accumulation, seed acquires desiccation tolerance enabling survival at low moisture, possible dormancy mechanisms establish preventing premature germination, progressive desiccation reduces seed water content from 50-70% to 10-20%, metabolic activity declines toward quiescence, and seed achieves mature viable state ready for dispersal and eventual germination, occurring weeks 12-18 post-fertilization completing seed development coordinating with berry ripening. This final maturation phase occurs during early-mid autumn in September-October in temperate regions Year N+1, representing seed development completion producing mature seed. Maturation mechanism involves storage reserve accumulation completion in embryo and endosperm, desiccation tolerance acquisition through protective molecule synthesis, metabolic activity decline toward dormant state, progressive desiccation reducing water content, possible dormancy imposition through abscisic acid and other mechanisms, seed coat hardening completing protection, and integrated maturation producing viable seed ready for dispersal. Reserve accumulation completion shows embryo cotyledons completing lipid and protein accumulation reaching 30-40% and 25-35% dry weight respectively, endosperm completing reserve accumulation 18-28% proteins and 12-22% lipids, total seed reserves adequate for germination and seedling establishment, and completed reserves as maturation outcome. Embryo reserve characteristics shows lipids as triacylglycerols in lipid bodies, proteins as albumins and globulins in protein bodies, reserves stored primarily in cotyledons, and reserves providing germination energy and nutrients. Endosperm reserve characteristics shows proteins and lipids in specialized organelles, endosperm surrounding embryo, endosperm reserves supporting embryo and seedling, and endosperm as nutritive tissue. Desiccation tolerance acquisition represents critical maturation process enabling seed survival at low moisture, late embryogenesis abundant LEA proteins accumulating in embryo and possibly endosperm, heat shock proteins possibly accumulating, protective sugars including sucrose and raffinose family oligosaccharides accumulating, desiccation tolerance mechanisms protecting cellular structures, and tolerance enabling seed storage. Late embryogenesis abundant proteins shows hydrophilic proteins accumulating during late maturation, LEA proteins protecting membranes and proteins during desiccation, proteins preventing aggregation and maintaining structure, multiple LEA protein families expressed, and LEA proteins as major desiccation protectants. Heat shock proteins shows HSPs possibly accumulating during maturation, chaperone proteins protecting other proteins, HSPs contributing to stress tolerance, and HSPs as cellular protectants. Protective sugar accumulation shows sucrose accumulating to high levels 5-15% dry weight, raffinose family oligosaccharides including raffinose, stachyose possibly accumulating, sugars replacing water in hydrogen bonding with membranes and proteins, sugars forming glasses vitrifying cytoplasm preventing crystallization, and sugars as essential desiccation protectants. Sugar protective mechanisms shows sugars stabilizing membranes during water loss, sugars preventing protein denaturation, glass formation at low moisture preventing cellular damage, and sugars enabling desiccation survival. Metabolic activity decline shows respiratory rate decreasing substantially during maturation, biosynthetic activity declining as reserve accumulation completes, gene expression shifting toward dormancy-associated patterns, and metabolism approaching quiescent state characteristic of mature dry seed. Respiratory decline shows oxygen consumption decreasing from maturation levels 1.5-2.5 μmol O₂ g⁻¹ h⁻¹ to very low mature seed <0.1-0.3 μmol O₂ g⁻¹ h⁻¹, respiratory rate reduction enabling metabolic quiescence, quiescent metabolism enabling long-term storage, and respiration as metabolic activity indicator. Gene expression changes shows storage compound biosynthesis genes downregulating, maturation-associated genes including LEA and dormancy-related genes upregulating, developmental transcriptional program shifting toward quiescence, and expression changes implementing maturation. Progressive desiccation shows water content declining from 50-70% during mid-development to 30-40% late maturation to final 10-20% in mature dry seed, water loss through seed coat to parent plant initially then to atmosphere after abscission, controlled developmental desiccation distinguished from stress-induced dehydration, and low moisture as mature seed characteristic. Desiccation kinetics shows progressive water loss over maturation weeks 12-18, typical water loss rate 3-8% moisture per week, accelerated desiccation during final maturation, and desiccation timing coordinating with maturation completion. Water loss pathways shows water moving from seed through funiculus to berry while attached, water evaporating through coat after seed separation or berry drying, and water movement regulated by coat permeability and environmental conditions. Seed moisture measurement shows gravimetric water content determination through drying, mature seed moisture 10-20% fresh weight basis, moisture critical for storage stability, and moisture as maturation indicator. Seed dormancy acquisition shows many grapevine cultivars exhibiting seed dormancy requiring stratification for germination, dormancy mechanisms including embryo dormancy possibly from abscisic acid, coat-imposed dormancy from impermeability possible though less important than embryo dormancy, and dormancy preventing premature germination. Embryo dormancy shows embryo unable to germinate immediately after maturation, dormancy requiring cold stratification typically 60-120 days at 1-5°C for release, dormancy broken through stratification enabling germination, and embryo dormancy as primary dormancy type in grapevine. Abscisic acid role shows ABA possibly accumulating during late seed maturation, ABA levels possibly 200-800 ng/g dry weight in mature seed, ABA imposing or maintaining dormancy, ABA declining during stratification enabling germination, and ABA as dormancy hormone. Coat-imposed dormancy shows hard seed coat possibly contributing to dormancy through impermeability or mechanical resistance, coat effects generally secondary to embryo dormancy in grapevine, scarification not typically required though coat removal may accelerate germination, and coat as potential dormancy factor. Dormancy variation shows dormancy depth varying among cultivars, some cultivars showing weak dormancy germinating readily, others showing deep dormancy requiring extended stratification, and genetic variation affecting dormancy. Metabolic quiescence establishment shows metabolism declining to basal maintenance levels, quiescent seed capable of long-term storage, quiescence reversible through imbibition and germination conditions, and quiescence as dormant seed state. Seed coat hardening completion shows coat lignification and hardening completing during maturation, hard brown coat protecting mature seed, coat as covered in Seed_Coat_Formation, and coat completion coordinating with overall maturation. Seed appearance shows mature seed hard with brown to dark brown coat, seed typically 4-7 mm length and 3-5 mm width, seed fresh weight when mature 20-50 mg depending on development, seed shape ovoid to ellipsoid, and appearance as mature seed characteristic. Seed viability shows mature seed viable capable of germination when appropriate conditions provided, viability requiring complete development including reserve accumulation and desiccation tolerance, dormancy requiring release before germination, and viability as maturation outcome. Seed longevity shows mature seeds storable for extended periods, storage longevity depending on moisture and temperature, dry cool storage optimal typically 1-5°C and <10% moisture, proper storage enabling viability retention for years, and longevity as seed quality attribute. Storage conditions shows cool dry conditions optimal for maintaining viability, warm moist conditions accelerating deterioration, sealed storage preventing moisture gain, and storage management as post-maturation factor. Maturation timing shows maturation largely completing by weeks 16-18 post-fertilization, timing varying somewhat with cultivar and environment, maturation coordinating with berry ripening, and timing as developmental endpoint. Maturation completion indicators shows seed hard and brown, low moisture 10-20%, dormant if tested, and indicators documenting mature state. Mature seed characteristics shows seed with completed reserves, desiccation tolerant, dormant or germination-competent, hard brown coat, viable ready for dispersal, and mature seed as developmental outcome. Environmental effects on seed maturation include temperature affecting maturation rate with optimal 20-26°C supporting normal progression, humidity affecting desiccation with low humidity promoting drying, and conditions affecting maturation quality and timing. Temperature effects shows warm conditions accelerating maturation, cool conditions extending maturation period, and temperature as rate determinant. Humidity effects shows low humidity promoting desiccation, high humidity slowing water loss potentially delaying maturation, and humidity affecting desiccation dynamics. The process encompasses three final sequential maturation phases including embryo desiccation tolerance acquisition, seed dormancy acquisition establishing germination control, and seed ripening completion finalizing maturation. Duration approximately 6 weeks representing final maturation weeks 12-18. Regional timing shows seed maturation during September-October completing seed development. Viticultural significance includes representing seed development completion producing viable seed, enabling seed storage and long-term viability, coordinating seed maturation with berry ripening and dispersal, and determining seed germination characteristics affecting propagation. Research applications include analyzing seed maturation regulation and mechanisms, investigating desiccation tolerance and protective molecules, evaluating dormancy mechanisms and regulation, quantifying maturation dynamics and completion, and assessing seed maturation as seed viability and quality determinant.
Seed_Maturation represents final seed development within Seed_Development. Subdivided into Embryo_Desiccation_Tolerance with LEA proteins and protective sugars, Seed_Dormancy_Acquisition with embryo dormancy and ABA (200-800 ng/g), and Seed_Ripening_Completion. Duration weeks 12-18. Reserves completed: embryo 30-40% lipids/25-35% proteins, endosperm 18-28% proteins/12-22% lipids. Desiccation 50-70% → 10-20% moisture. Respiration 1.5-2.5 → <0.1-0.3 μmol O₂ g⁻¹ h⁻¹. Dormancy requires stratification 60-120 days 1-5°C. Mature seed 4-7 mm, 20-50 mg, viable. Temperature 20-26°C optimal.
Samen-Vollendung
Samenreifungsphase
final seed development
finale Samenentwicklung
mature seed formation
seed completion
seed maturation phase
Seed Maturation
Seed maturity monitoring in Cabernet Sauvignon through ripening reveals the progression critical for tannin quality, with véraison +2 weeks showing green seeds (easily cut, high chlorophyll, tannins highly extractable and bitter in tasting), véraison +4 weeks showing yellow-brown transition (harder texture, declining extractability, tannins becoming softer), and véraison +6 weeks showing dark brown fully lignified seeds (hard, crack rather than cut, tannins minimally extractable with soft sensory character), with the seed maturity progression enabling harvest timing that achieves desired tannin contribution ranging from structured and firm (earlier seed maturity) to soft and integrated (later seed maturity).
Seed maturity quality is a physiological maturity quality that represents the developmental state of grape seeds, characterizing the progression from green, chlorophyll-containing seeds through color transition to fully lignified, brown seeds with viable embryos, a transformation that correlates with changes in seed tannin extractability and sensory contribution to wine. Within the BFO framework, this quality specifically addresses the seed component of physiological maturity, distinguishing seed developmental state from pulp or skin maturity while recognizing that seed maturation provides important visual and tactile indicators of overall berry ripeness and that seed tannin evolution significantly affects wine quality through changes in extractability, astringency, and flavor contribution. The ontological significance lies in seed maturity indicating completion of the berry's reproductive function while simultaneously determining the tannin contribution seeds will make during winemaking, where immature green seeds contain highly extractable, bitter, astringent tannins that can dominate wine character, while mature brown seeds have polymerized tannins with reduced extractability and softer sensory impact, making seed maturity assessment critical for red wine quality. The visual and tactile indicators of seed maturity include color change (green → yellow → brown, indicating chlorophyll loss and lignification), texture hardening (soft, cuttable seeds becoming hard and resistant), and ease of separation from pulp (mature seeds releasing cleanly), with these indicators providing accessible field assessment tools for ripening monitoring. The tannin chemistry changes during seed maturation include progressive polymerization of monomeric and oligomeric proanthocyanidins into larger polymers (reducing extractability and astringency), oxidative browning (contributing to color change), and lignification of seed coat (creating physical barrier to extraction), with these changes explaining the sensory differences between wines made from fruit at different seed maturity levels. The decoupling risk between seed maturity and sugar accumulation is particularly relevant in warm climates, where rapid sugar accumulation may achieve target Brix while seeds remain green and tannins harsh, creating the quality challenge addressed by techniques like extended hang time, early harvest with sugar adjustment, or selection for cultivars with faster seed maturation relative to sugar accumulation. Understanding seed maturity quality enables assessment of tannin contribution potential, harvest timing for optimal seed-derived tannin character, and recognition of seed-sugar decoupling in climate change contexts.
Seed_Maturity_Quality addresses the developmental state of grape seeds affecting tannin extractability and sensory contribution. This quality provides visual and tactile indicators useful for field ripeness assessment. The class supports tannin quality management through seed maturity characterization guiding harvest timing.
Kernbräunung
Kernreife
Samenlignifizierung
Samenreife
pip maturity
seed browning
seed lignification
seed ripeness
Ristic R, Iland PG (2005) Relationships between seed and berry development of Vitis vinifera L. cv. Shiraz: developmental changes in seed morphology and phenolic composition. Australian Journal of Grape and Wine Research 11(1): 43-58 Kennedy JA et al. (2000) Composition of grape skin proanthocyanidins at different stages of berry development. Journal of Agricultural and Food Chemistry 48(11): 5348-5355
Seed Maturity Quality
The seed number of Merlot documents a mean of 1.5-2.5 fully developed seeds per berry, with the distribution across a typical sample showing approximately 10% one-seeded, 45% two-seeded, 35% three-seeded, and 10% four-seeded berries — the predominance of two-seeded berries reflects the typical fertilization efficiency where approximately 50-60% of available ovules are successfully fertilized and develop to mature seed stage. The seed number variation within a single cluster creates the within-cluster berry size heterogeneity observed in all seeded V. vinifera cultivars — four-seeded Merlot berries weigh approximately 2.0-2.5 g while one-seeded berries from the same cluster weigh 1.2-1.5 g, a 40-60% size difference attributable to the additional seed-derived growth hormones. This moderate seed number in Merlot contrasts with the higher mean seed number of Grenache (2.0-3.0 seeds, contributing to Grenache's larger berry size) and the absent or vestigial seeds of Sultanina (seed traces of 2-4 mm length with no viable embryo, the stenospermocarpic condition).
A seed description component that records the typical number of fully developed seeds per berry, scored according to OIV descriptor 241 on a scale from absent or vestigial (state 1, seedless cultivars with stenospermocarpic seed traces) through one (state 2), two (state 3), three (state 4), to four (state 5), with seed number reflecting the pollination and fertilization success per berry and having direct consequences for berry size, berry composition, and the seed-to-berry ratio that determines seed tannin contribution to wine.
Seed_Number documents a character that is both a cultivar attribute and a season-specific outcome of pollination conditions. The maximum seed number per berry is four (determined by the two ovules per locule in the two-carpel ovary), but the actual number depends on the proportion of ovules that are pollinated, fertilized, and successfully develop past the critical 2-4 weeks post-fertilization stage when many embryos abort — this proportion varies between cultivars (reflecting differences in ovule receptivity, pollen tube growth, and embryo viability), between seasons (reflecting bloom-period temperature, humidity, and nutrition that affect pollination efficiency), and between berry positions within the cluster (basal berries often having higher seed counts than distal berries): the cultivar-specific mean seed number under standard conditions is therefore a genetically determined trait modulated by environmental factors, with the genetic component ranging from the extreme of complete seedlessness (stenospermocarpic cultivars where embryo abortion is genetically programmed) to cultivars with consistently high seed numbers (3-4 seeds per berry under normal conditions). The relationship between seed number and berry weight has been quantified at approximately 0.3-0.5 g per additional seed within a cultivar, providing a physiological explanation for the within-cluster berry size variation that is a universal feature of seeded grape cultivars.
Kerne-Anzahl
Kernzahl
Samen-pro-Beere
ovule fertilization success
pip count per berry
seed number record
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Ristic R and Iland PG (2005) Relationships between seed and berry development of Vitis vinifera L. cv Shiraz, Australian Journal of Grape and Wine Research 11: 43-58 May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press
Seed Number
Müller-Thurgau serves in a Seed_Parent_Role in German breeding programs, its reliable fruit set and good seed production making it a practical choice for generating large progeny populations from crosses with diverse pollen donors. A seeded table grape variety bears a Seed_Parent_Role in seedlessness breeding, providing the ovules pollinated by stenospermocarpic varieties where reciprocal crosses would yield no viable seeds due to embryo abortion. Vitis vinifera cv. Riesling functions as seed parent when crossed with resistant Muscadinia derivatives, with hybrid seeds collected from Riesling clusters following controlled pollination.
A maternal parent role that inheres in a grapevine genotype specifically by virtue of its function as the seed-bearing parent in controlled crosses, emphasizing the developmental role of producing viable seeds containing hybrid embryos. The seed parent designation highlights the practical breeding function—the vine that bears the crossed fruit and produces seeds for germination—distinguishing this role from broader maternal genetic contribution. Seed parent selection considers both genetic objectives (desired trait contributions) and practical seed production characteristics: fruit set reliability, seed number per berry, seed viability, freedom from stenospermocarpy (seedless tendency), and ease of seed extraction and germination. Varieties with poor seed production characteristics may be preferentially used as pollen parents even when their genetic contribution is equally desired, simply because crossing in the reverse direction yields more progeny for evaluation. The seed parent provides not only genetic material but also the maternal environment in which hybrid embryos develop, potentially influencing seed maturation and early seedling vigor through maternal effects.
Seed_Parent_Role emphasizes the practical seed production function within the broader maternal parent category, recognizing that breeding program success depends substantially on generating adequate progeny numbers for selection. The seed parent's influence extends beyond nuclear genetic contribution to include cytoplasmic inheritance and maternal effects on seed development. In grapevine breeding, some genotypes are markedly superior as seed parents (consistent fruit set, many viable seeds per cross) while others, despite valuable genetic contributions, are difficult to use as seed parents (poor set, few seeds, low viability). Seedlessness breeding presents special challenges, as stenospermocarpic varieties produce seeds only when used as pollen parents on seeded maternal genotypes, restricting crossing design options. Documentation of which parent served as seed parent is standard in breeding records, enabling analysis of maternal effects and cytoplasmic contributions.
Samenelter
female crossing partner
ovule donor
samentragender Elter
seed parent
weiblicher Kreuzungspartner
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, Elsevier pp.: 3-22 Ramming DW (2008) Seedless grape breeding, in Bentley R (ed) The Handbook of Grape Processing, Blackwell Perl A, Eshdat Y (1998) Grape breeding and biotechnology, Horticultural Reviews 22: 329-360
Seed Parent Role
In Cabernet Franc berries in Saint-Émilion, Bordeaux during BBCH 81-89, seed phenolic maturation reduces extractable seed tannin from 8.2 to 3.5 mg/g seed over 42 days while total seed tannin per berry remains stable at 4.8-5.2 mg, with seed coat browning index increasing from 0.25 to 0.72 absorbance units at 420 nm, mDP declining from 10.5 to 8.2, and galloylation stable at 26-28%, establishing mature seed phenolic status with reduced extraction potential under moderate oceanic conditions averaging 24°C.
Seed phenolic maturation constitutes the comprehensive developmental progression of phenolic compounds within Vitis vinifera grape seed tissues during berry development and ripening, wherein proanthocyanidins synthesized primarily during early seed development undergo post-synthetic modifications including oxidative browning, polymerization changes, and extractability reduction that transform initial tannin properties to establish the mature seed phenolic status characterized by reduced astringency potential, decreased extractability, and brown coloration indicative of optimal seed ripeness for quality wine production. This process reflects the distinctive developmental trajectory of seed phenolics compared to skin tissues, with proanthocyanidin biosynthesis in seeds occurring predominantly before veraison during the seed coat development phase spanning BBCH 71-81, followed by cessation of active synthesis and initiation of post-synthetic maturation processes during the post-veraison period when oxidative modification becomes the dominant factor influencing seed phenolic properties. The quantitative aspects of seed phenolic maturation show characteristic patterns with total seed tannin content per berry remaining relatively stable during post-veraison while concentration per unit seed mass may decrease through seed size increase, and extractable tannin content declining substantially by 40-70% between veraison and harvest as oxidative modification reduces tannin solubility and availability for extraction during winemaking. The structural characteristics of seed tannins differ fundamentally from skin tannins, with seed proanthocyanidins showing lower mean degree of polymerization of 5-12 units compared to skin values of 25-45, minimal prodelphindin content typically below 5% compared to skin levels of 20-40%, and elevated galloylation percentages of 15-35% compared to skin values of 2-8%, contributing to the distinctive astringency properties associated with seed tannin extraction. Temperature during ripening influences seed phenolic maturation through effects on oxidative reaction rates, with warm conditions accelerating browning and extractability reduction while cool conditions may delay maturation requiring extended hang-time to achieve equivalent seed phenolic status. The enological significance of seed phenolic maturation relates to wine astringency management, with immature seed tannins contributing harsh, aggressive astringency considered negative for wine quality, while mature seed phenolics with reduced extractability contribute less aggressively to wine tannin content, making seed maturation assessment an important component of harvest timing decisions for red wine production seeking optimal phenolic balance.
Seed_Phenolic_Maturation is parent class under Phenolic_Maturation representing seed-localized phenolic processes. Contains subclasses Seed_Tannin_Polymerization, Seed_Browning, and Phenolic_Seed_Maturity. Distinguishes from Skin_Phenolic_Development by tissue localization and developmental trajectory. Biosynthesis complete before veraison post-veraison involves oxidative modification. Extractability decreases 40-70% during ripening.
Samen-Phenol-Reifung
Samen-Polyphenol-Reifung
Samen-Tannin-Reifung
Traubenkern-phenolische-Entwicklung
grape seed phenolic development
seed phenolic evolution
seed polyphenol ripening
seed tannin maturation
Kennedy JA et al (2000) Grape seed development germination and proanthocyanidin composition. Journal of Agricultural and Food Chemistry 48: 1490-1495 Ristic R and Iland PG (2005) Relationships between seed and berry development of Vitis vinifera L. Australian Journal of Grape and Wine Research 11: 43-58 Adams DO (2006) Phenolics and ripening in grape berries. American Journal of Enology and Viticulture 57: 249-256
Seed Phenolic Maturation
Syrah in Rhône Valley shows seed ripening completion during October Year N+1, with storage reserves finalized at 35-39% lipids and 29-34% proteins in embryo cotyledons, 21-26% proteins and 15-20% lipids in endosperm, complete desiccation reducing moisture to 11-15%, seed coat achieving maximum hardness 74-96 N penetration resistance with dark brown color, dormancy fully established requiring 90-115 days stratification, metabolic quiescence with respiration <0.15 μmol O₂ g⁻¹ h⁻¹, mature seed 5.4-6.6 mm length weighing 36-48 mg, viability 88-94% after stratification, seed completing development by week 16-17 coordinating with berry full ripeness BBCH 89, under warm Mediterranean conditions of 22-28°C autumn temperature, typical of this mid-to-late season cultivar producing mature viable seed ready for dispersal
Seed ripening completion represents the final seed development phase during seed maturation when all seed developmental processes conclude, storage reserves reach final levels, desiccation reduces moisture to minimum 10-20%, seed coat achieves maximum hardness and brown pigmentation, dormancy fully established, metabolic quiescence achieved with respiration 60-100 N penetration resistance, coat fully lignified with 25-40% lignin in testa, coat dark brown from complete pigmentation, coat providing maximum protection, and coat maturation as structural completion. Coat color finalization shows mature coat medium to dark brown depending on cultivar, color from accumulated oxidized phenolics, color stable in mature seed, color intensity as maturity indicator, and color as identification feature. Coat protective function shows mature coat providing mechanical protection from damage, impermeability preventing pathogen entry and controlling imbibition, coat enabling seed survival during dispersal and storage, and protection as coat function. Dormancy complete establishment shows dormancy fully established in mature seed, dormancy at maximum depth requiring full stratification for release, dormant state stable during storage, dormancy controlling germination timing, and dormancy completion as maturation feature. Dormancy in mature seed shows embryo in deeply dormant state, dormancy preventing germination until stratification, dormancy stable for months to years in storage, and dormancy as mature seed characteristic. Metabolic quiescence achievement shows respiratory rate declining to very low minimal levels 80-95% in properly matured and stored seeds, viability as functional maturation outcome, and viability as seed quality measure. Viability testing shows germination tests after stratification determining viability, tetrazolium tests showing embryo viability, viability declining gradually during long-term storage, and testing enabling quality assessment. Seed longevity shows properly stored seeds maintaining viability for years, storage conditions critical with cool dry optimal, storage at 1-5°C and 5-10% moisture enabling extended longevity, suboptimal conditions reducing longevity, and longevity as storage characteristic. Seed dispersal readiness shows mature seed ready for dispersal by frugivory, seed coat protecting embryo during passage through digestive tract, seed survival enabling dispersal, and dispersal as ecological function. Berry ripening coordination shows seed maturation completing during or shortly before berry full ripeness, timing ensuring seeds mature when fruit attractive to frugivores, coordination as reproductive strategy, and timing as seed-fruit synchronization. Seed abscission shows seed possibly separating from funiculus during or after berry ripening, separated seed free within berry, abscission enabling dispersal, and separation as completion event. Completion timing shows ripening completion by weeks 16-18 post-fertilization, timing varying slightly with cultivar and environment, completion coordinating with berry maturation typically BBCH 87-89 berries ripe for harvest, and timing as developmental endpoint. Maturation completion indicators shows seed hard and dark brown, very low moisture 10-20%, dormant if tested, metabolic quiescence achieved, and indicators documenting completion. Mature seed properties shows seed with completed reserves, desiccation tolerant, dormant, hard brown coat, viable, and properties as developmental outcomes. Seed after maturation shows mature seed ready for dispersal and storage, seed maintaining viability during storage if conditions appropriate, seed capable of germination after stratification, and mature seed as completed developmental product. Environmental effects on completion include temperature affecting final maturation rate, humidity affecting final desiccation, and environmental conditions affecting completion timing and quality. Temperature effects shows warm autumn conditions promoting faster completion, cool conditions possibly extending final maturation, and temperature as rate factor. Humidity effects shows low humidity promoting final desiccation, high humidity slowing water loss, and humidity affecting final moisture. The process represents third and final aspect of seed maturation completing all development, following embryo desiccation tolerance and seed dormancy acquisition. Duration final completion during weeks 16-18 representing developmental endpoint. Regional timing shows seed ripening completion during October as mature seed produced. Viticultural significance includes representing seed development completion producing viable dispersal-ready seed, coordinating seed and berry maturation timing, enabling seed viability for propagation and breeding, and determining seed quality for storage and germination. Research applications include analyzing seed maturation completion and developmental regulation, investigating maturation coordination with berry ripening, evaluating seed quality and viability determinants, quantifying maturation timing and indicators, and assessing seed ripening completion as reproductive success determinant.
Seed_Ripening_Completion is third and final aspect of Seed_Maturation, distinguished by developmental completion. Represents weeks 16-18. All processes concluded: reserves finalized (embryo 30-40% lipids/25-35% proteins, endosperm 18-28% proteins/12-22% lipids), desiccation complete 10-20% moisture, coat maximally hard dark brown, dormancy fully established, metabolic quiescence 80-95%. Dispersal ready. Coordinates with berry ripeness. Follows Embryo_Desiccation_Tolerance and Seed_Dormancy_Acquisition. Completes Seed_Maturation and Seed_Development. Developmental endpoint.
Entwicklungs-Vollendung
Samenreifungs-Abschluss
developmental completion
Seed Ripening Completion
The seed shape of Nebbiolo documents distinctly pyriform (pear-shaped) seeds with a prominent, sharply pointed beak measuring 2.0-2.5 mm (approximately 30% of total seed length) — the beak projects from the narrower micropylar end while the broader chalazal end bears a deep central groove flanked by two lateral ridges, producing the characteristic seed outline that has been used by Italian ampelographers to distinguish Nebbiolo seeds from those of other Piedmontese cultivars in both living material and archaeological contexts. The long-beaked pyriform Nebbiolo seed shape contrasts with the short-beaked ovoid seeds of Barbera (beak 1.0-1.5 mm, approximately 15% of total length, with a more rounded overall outline), the globular seeds of Chasselas (nearly spherical with a very short, blunt beak of 0.5-1.0 mm), and the elongated seeds of Sultanina (seed traces of 2-4 mm length that are flattened, triangular remnants without a developed beak or endosperm, reflecting the stenospermocarpic abortion that occurs 3-4 weeks after fertilization).
A seed description component that records the overall geometric form of the mature seed, documenting the outline in lateral view (pyriform, ovoid, globular, or elongated), the prominence and length of the beak (rostrum, the pointed projection at the micropylar end), the depth and width of the ventral chalazal depression, and the cross-sectional profile (round, dorsiventrally compressed, or laterally compressed), with seed shape providing cultivar-diagnostic characters that are particularly valuable for archaeological cultivar identification where seed morphometrics are often the only available evidence.
Seed_Shape provides the morphometric characters that have the longest history of diagnostic use in grape identification — seed shape was already recognized as cultivar-diagnostic by Roman agricultural writers, and the systematic morphometric analysis of grape seeds (seed archaeobotany) has become a primary tool for tracing the history of grape cultivation and cultivar dispersal from the Neolithic origin centers in the South Caucasus through the Mediterranean to the Atlantic seaboard. The key morphometric parameters used in both modern ampelography and archaeological seed identification are the length-to-width ratio (discriminating between globular, ovoid, and elongated seeds), the beak length relative to total seed length (discriminating between short-beaked wild-type seeds and long-beaked domesticated seeds), and the chalazal groove depth and configuration (providing cultivar-group level discrimination): the domestication-associated shift toward longer, more pyriform seeds with more prominent beaks is one of the clearest morphological signatures of grape domestication, attributed to selection for larger berries with fewer, larger seeds and possibly to the shift from sexual reproduction (selecting for efficient soil penetration by compact, round seeds) to vegetative propagation (removing the selection pressure on seed shape). Modern geometric morphometric methods (Elliptic Fourier analysis of seed outline) can discriminate between cultivar groups with 70-85% accuracy, approaching but not yet matching the resolution of molecular marker identification.
Kernform
Samen-Umriss-Beschreibung
Traubenkern-Geometrie
grape seed geometry
pip shape classification
seed outline record
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Terral JF et al. (2010) Evolution and history of grapevine under domestication: new morphometric perspectives to understand seed domestication syndrome and reveal origins of ancient European cultivars, Annals of Botany 105: 443-455 Mangafa M and Kotsakis K (1996) A new method for the identification of wild and cultivated charred grape seeds, Journal of Archaeological Science 23: 409-418
Seed Shape
The seed size of Syrah documents medium-large seeds measuring 6.5-7.5 mm length, 4.0-5.0 mm width, and 30-45 mg individual seed weight (mean of 100-seed sample) — these relatively large seeds contain proportionally more tannin per seed (8-12 mg catechin equivalents per seed) than the smaller seeds of Pinot Noir (5.5-6.5 mm length, 3.5-4.5 mm width, 25-35 mg weight, 5-8 mg tannin per seed), contributing to the structured, moderately tannic character of Syrah wines alongside the skin tannin contribution. The total seed tannin per berry is the product of individual seed tannin content and seed number — Syrah berries with 2 large seeds (total seed tannin approximately 16-24 mg/berry) provide a similar seed tannin pool to Grenache berries with 3 smaller seeds (approximately 15-21 mg/berry), illustrating the compensatory relationship between seed size and seed number.
A seed description component that records the dimensions and weight of individual mature seeds, documenting length (from beak tip to chalazal end), width (maximum transverse dimension), thickness, and weight (mean of 100-seed sample), scored according to OIV descriptor 242 on a scale from very small (state 1) to very large (state 9), with seed size reflecting the cultivar-specific endosperm and embryo development and influencing the total seed tannin mass available for extraction during winemaking.
Seed_Size documents a character that is highly genetically stable — seed dimensions are among the least environmentally variable ampelographic characters because seed development occurs within the protected environment of the berry interior, buffered from the temperature, light, and water fluctuations that modify external organ characters. The seed reaches its final size during the Phase I growth period (approximately 4-6 weeks post-fertilization), when rapid cell division produces the full complement of endosperm and testa cells, followed by a maturation period during Phase II and Phase III when the cells accumulate storage reserves (lipids in the endosperm, tannins in the testa) without further size increase: the final seed dimensions therefore reflect the Phase I cell division rate and duration, which are primarily genetically determined with minimal environmental modulation. The stability of seed morphology has made seed remains the primary evidence for cultivar identification in archaeological viticulture — waterlogged or charred grape seeds recovered from Neolithic through Medieval sites retain diagnostic morphological features (length, width, length-to-width ratio, beak length, chalazal groove depth) that enable tentative cultivar or cultivar-group identification using discriminant analysis of seed morphometric data, with the transition from the small, round seeds characteristic of wild V. vinifera subsp. sylvestris to the larger, more elongated seeds of domesticated V. vinifera subsp. sativa being one of the primary indicators of grape domestication in the archaeological record.
Kerngröße
Samen-Dimensionserfassung
Traubenkern-Größenbewertung
grape seed size assessment
pip dimension record
seed weight measurement
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Terral JF et al. (2010) Evolution and history of grapevine under domestication: new morphometric perspectives to understand seed domestication syndrome and reveal origins of ancient European cultivars, Annals of Botany 105: 443-455
Seed Size
Seed tannin concentration ranges from 1-5 mg/berry across cultivars, typically comprising 50-80% of total berry tannin content. High seed tannin cultivars such as Cabernet Sauvignon (3-5 mg/berry for 2-4 seeds) and Mourvèdre (2.5-4 mg/berry) contrast with seedless or low-seed cultivars. Individual seed tannin content of 0.8-1.5 mg/seed is relatively consistent across cultivars, making seed number the primary determinant of total seed tannin. Seed proanthocyanidins show lower mDP (3-12 subunits) than skin tannins, with 20-40% galloylation (gallic acid esterification at the 3-position) versus 3-8% in skins. Seed tannin biosynthesis peaks 4-6 weeks post-flowering, with subsequent oxidative browning and decreased extractability as seeds lignify post-veraison. Extraction into wine requires extended maceration (>14 days) and/or mechanical seed damage.
A tannin accumulation disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate proanthocyanidins in grape seed integuments (testa and tegmen), with genotype-specific concentration, shorter polymer length than skin tannins, and higher galloylation producing more astringent, bitter tannin character when extracted into wine.
Seed tannin represents the majority of berry tannin in most cultivars but requires careful extraction management to avoid excessive astringency and bitterness from the shorter, more galloylated polymers. The trait shows high heritability for concentration per seed, with total seed tannin determined primarily by seed number per berry. Seed maturity (brown color, reduced extractability) is used as a harvest indicator, with "green" seed tannins producing harsher wine character. Climate effects on seed tannin are less pronounced than for skin tannins, as seed development largely precedes the ripening period.
Kerntannin
Samen-Gerbstoff
Samenproanthocyanidin
seed coat tannin trait
seed condensed tannin
seed proanthocyanidin
Kennedy JA et al. (2000) Grape seed proanthocyanidins and their structural units. Phytochemistry 55:77-85 Cadot Y et al. (2006) Flavan-3-ol compositional changes in grape berries (Vitis vinifera L. cv Cabernet Franc) before veraison, using two complementary analytical approaches, HPLC reversed phase and histochemistry. Analytica Chimica Acta 563:159-170
Seed Tannin Disposition
In Malbec berries in Mendoza, Argentina during BBCH 83-89, seed tannin polymerization modifies seed proanthocyanidin structure with mDP decreasing from 11.2 to 7.8 over 40 days through oxidative cleavage, extension unit composition shifting as epicatechin-gallate proportion decreases from 32% to 26% through preferential oxidation, and gel permeation chromatography revealing molecular weight distribution broadening with increased low-molecular-weight fraction indicating partial depolymerization concurrent with browning under warm high-altitude conditions averaging 28°C.
Seed tannin polymerization describes the structural modification of proanthocyanidin compounds within Vitis vinifera grape seed tissues during the ripening period, wherein seed tannins synthesized before veraison undergo oxidative and potentially acid-catalyzed reactions that modify polymer structure, typically producing slight decreases in mean degree of polymerization from 8-12 units at veraison to 5-9 units at harvest through oxidative cleavage mechanisms concurrent with browning reactions that fundamentally alter seed tannin properties and extractability characteristics. This process differs from skin tannin polymerization in its predominantly degradative rather than elongative character, with seed tannins experiencing oxidative attack generating quinone intermediates that may undergo cleavage reactions reducing polymer chain length, cross-linking reactions generating insoluble polymeric material, and nucleophilic addition reactions incorporating other phenolic compounds into modified tannin structures. The biochemical mechanisms underlying seed tannin polymerization involve polyphenol oxidase and peroxidase catalyzed oxidation of catechol and pyrogallol groups on flavan-3-ol subunits generating reactive quinone species, with subsequent non-enzymatic reactions including intramolecular rearrangements, intermolecular cross-linking, and potential chain cleavage producing the structurally modified tannin populations that characterize mature seed tissues. The quantitative assessment of seed tannin polymerization employs analytical methods including phloroglucinolysis yielding mean degree of polymerization and subunit composition, gel permeation chromatography providing molecular weight distribution, and protein precipitation assays measuring functional tannin activity, with these techniques revealing the structural changes accompanying oxidative modification during seed maturation. The relationship between seed tannin polymerization and extractability reflects the formation of oxidatively modified polymers with reduced solubility in hydroalcoholic extraction media, with highly oxidized seed tannins showing decreased dissolution rates and reduced equilibrium solubility that limits their contribution to wine tannin content during standard maceration practices. Temperature during ripening influences seed tannin polymerization through effects on oxidative enzyme activities and non-enzymatic reaction rates, with warm conditions accelerating structural modification while cool conditions may preserve less modified tannin structures requiring longer ripening periods to achieve equivalent maturation status.
Seed_Tannin_Polymerization is subclass of Seed_Phenolic_Maturation representing structural modification of seed proanthocyanidins. Distinguishes from Tannin_Polymerization under general category by seed-specific degradative character. Sibling to Seed_Browning and Phenolic_Seed_Maturity. mDP typically decreases from 8-12 to 5-9. Oxidative cleavage predominates over elongation. Temperature-dependent reaction rates.
Samen-Proanthocyanidin-Modifikation
Samen-Tannin-Polymerisation
Samen-Tannin-Struktur-Änderung
Samen-kondensierte-Tannin-Oxidation
seed condensed tannin polymerization
seed proanthocyanidin modification
seed tannin oxidative modification
seed tannin structural change
Kennedy JA et al (2001) Changes in grape seed polyphenols during ripening. Phytochemistry 55: 77-85 Downey MO et al (2003) Analysis of tannins in seeds and skins of Shiraz. Australian Journal of Grape and Wine Research 9: 15-27 Cadot Y et al (2006) Anatomical histological and histochemical changes in grape seeds during development. Journal of Agricultural and Food Chemistry 54: 9206-9215
Seed Tannin Polymerization
Seed weight data for Cabernet Sauvignon from St. Helena, Napa Valley (2023 vintage) records mean total seed weight of 68 mg per berry (2.1 seeds per berry, mean individual seed weight 32 mg), with seed color assessed as fully brown (testa lignification complete), indicating advanced seed maturity where residual extractable tannin is predominantly high-molecular-weight, polymerized proanthocyanidin with relatively soft astringency perception. Seed weight data for Nebbiolo from Barolo (2023 vintage) documents 54 mg total seed weight per berry (2.8 seeds per berry, individual seed weight 19 mg), with the higher seed count per berry characteristic of Nebbiolo contributing to the cultivar's distinctively high seed tannin-to-skin tannin ratio and the firm, structured tannic profile that requires extended aging to resolve. Seed weight data for Blaufränkisch from Neckenmarkt (2024 vintage) records 58 mg total seed weight (2.3 seeds per berry, 25 mg individual), with seed browning assessment indicating 85% testa coloration at 20° KMW harvest, confirming adequate seed maturity for the planned 14-day maceration.
Seed weight data is a berry composition data entity that records the mass and number of seeds (pyrenes) per grape berry, determined by manual dissection and gravimetric measurement, expressed as total seed weight per berry in milligrams (mg/berry), mean individual seed weight in milligrams (mg/seed), and number of seeds per berry (count), providing critical information for predicting the contribution of seed-derived tannins to wine astringency and bitterness, and for assessing seed maturity through the correlation between seed weight, color, and tannin extractability during the ripening process.
Seed weight data carries disproportionate winemaking significance relative to its modest contribution to total berry mass (typically 3-6% of berry weight) because grape seeds contain 20-55% of total berry tannin and are the exclusive source of flavan-3-ol monomers (catechin, epicatechin) and galloylated proanthocyanidins that contribute pronounced bitterness and harsh astringency when extracted in unpolymerized form. The maturation trajectory of seeds during ripening follows a distinct pattern — seed fresh weight increases during early development (BBCH 71-79), plateaus around véraison, and may decrease slightly post-véraison through desiccation simultaneously, total seed tannin content declines by 30-60% from véraison to harvest as tannins become oxidatively polymerized and increasingly bound to lignifying cell walls, reducing their extractability. Seed color change from green through yellow-brown to fully brown provides a visual proxy for tannin maturity — green seeds release harsh, astringent, low-molecular-weight tannins that produce aggressive wine texture, while brown seeds release less total tannin of higher mean degree of polymerization perceived as softer and more integrated. The number of seeds per berry varies by cultivar (Muscat Blanc typically 1-2, Cabernet Sauvignon 2-3, Nebbiolo 2-4, some table grapes 0-1) and by fertilization success, and directly influences the total seed tannin reservoir available for extraction. Winemaking practices that modify seed tannin contribution include berry sorting to remove green-seeded berries, limiting maceration duration, controlling extraction temperature (lower temperatures reduce seed tannin solubility), and removing free-run wine early to limit seed contact time.
Beerenkerndaten
Kerngewicht
Kerngewichtsdaten
Samenfraktion
Samengewicht
pyrene weight
seed count data
seed fraction weight
seed mass
seed weight per berry
Kennedy JA et al. (2001) Changes in grape seed polyphenols during fruit ripening. Phytochemistry 55(1): 77-85 Cadot Y et al. (2006) Anatomical, histological, and histochemical changes in grape seeds from Vitis vinifera L. cv Cabernet franc during fruit development. Journal of Agricultural and Food Chemistry 54(24): 9206-9215 Ristic R, Iland PG (2005) Relationships between seed and berry development of Vitis vinifera L. cv Shiraz: developmental changes in seed morphology and phenolic composition. Australian Journal of Grape and Wine Research 11(1): 43-58
Seed Weight Data
Red Globe bears a Seeded_Table_Grape_Scion_Role as the dominant variety for Asian export markets, with very large berries (12-18 g, up to 25 g with optimal thinning), pink-red coloration, 2-4 seeds per berry, crisp texture, and mild flavor commanding premium prices in Chinese wholesale markets (¥15-40/kg) where berry size is the primary quality criterion. Kyoho serves in a Seeded_Table_Grape_Scion_Role as Japan's prestige table grape, with extraordinarily large berries (15-20 g), deep purple slip-skin character, intense labrusca-influenced Concord-like aroma, and 2-3 large seeds accepted in a market valuing flavor intensity and gift-presentation quality at retail prices exceeding ¥2000/cluster. Italia (Ideal, Muscat Italia) bears a Seeded_Table_Grape_Scion_Role as the classic European quality seeded variety, with large berries (8-12 g), distinctive muscat aroma, golden-green color, 2-3 seeds, and crisp texture defining premium table grapes in Mediterranean and South American markets.
A table grape scion role that inheres in a grafted scion cultivar by virtue of its production of berries containing fully developed, hard seeds, representing the ancestral condition for Vitis vinifera before the selection of seedless mutants. Seeded table grape scions produce berries with 1-4 mature seeds per berry, each seed containing viable embryo, endosperm, and lignified seed coat (testa) that consumers must either swallow, chew, or spit during consumption. Seeded varieties typically produce larger berries than seedless counterparts (often 8-20 g versus 3-8 g for seedless) due to the hormonal contributions (gibberellins, auxins) of developing seeds that drive cell division and enlargement. While seedlessness dominates Western markets, seeded varieties maintain strong positions in Asian markets (China, Japan, Korea, India) where exceptionally large berry size commands premium prices and cultural preferences accept or even favor seeds as indicators of naturalness and quality. Major seeded table grape varieties include Red Globe (the world's most traded seeded variety), Italia, Muscat of Alexandria, and the giant-berried Japanese cultivars (Kyoho, Pione, Shine Muscat).
Seeded_Table_Grape_Scion_Role captures the historically dominant but now regionally specialized category of seed-containing table grapes. The global market bifurcation between seedless-preferring Western consumers and size-preferring Asian consumers creates parallel breeding programs—Western programs focus exclusively on seedless varieties while Asian programs (particularly Japanese) develop giant-berried seeded cultivars achieving berry weights of 15-25 g impossible in seedless genetics. Shine Muscat represents a Japanese breakthrough combining large size (12-15 g), muscat flavor, and seedlessness through stenospermocarpy, suggesting future convergence. Seeded varieties remain important for markets valuing traditional cultivars (Italia in Europe, Muscat of Alexandria in North Africa), and as breeding parents contributing berry size and quality traits to seedless progeny via embryo rescue.
Samentraube
Tafeltraube mit Kernen
kernhaltige Tafeltraube
pip grape
seed-containing grape
seeded grape
seeded table grape
Creasy GL, Creasy LL (2009) Grapes, CABI Yamada M, Sato A (2016) Advances in table grape breeding in Japan, Breeding Science 66: 34-45 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press OIV (2019) Statistical Report on World Vitiviniculture
Seeded Table Grape Scion Role
Thompson Seedless (Sultana) bears a Seedless_Table_Grape_Scion_Role as the progenitor of modern seedless table grape production, a stenospermocarpic variety with small soft seed traces that established seedlessness as a commercial category in California from the 1870s onward, now globally planted on over 400,000 hectares. Crimson Seedless serves in a Seedless_Table_Grape_Scion_Role demonstrating that seedless varieties can achieve size, color, and quality competitive with seeded cultivars through breeding advancement—its 4-6 g berries approach the size of smaller seeded varieties without GA treatment. Cotton Candy (IFG Seven) bears a Seedless_Table_Grape_Scion_Role exemplifying the premium branded seedless grape category, where distinctive flavor, complete seedlessness, and patented status command 2-3× price premiums in retail markets.
A table grape scion role that inheres in a grafted scion cultivar by virtue of its production of berries lacking fully developed hard seeds, a trait that has become the dominant consumer preference in North American, European, and increasingly global table grape markets. Seedless table grape scions produce berries through one of two developmental mechanisms—stenospermocarpy (seed development initiates but aborts, leaving soft seed traces) or parthenocarpy (fruit develops without fertilization, completely seedless)—both resulting in edible berries consumed without seed interference. Seedlessness fundamentally alters eating experience, eliminates the need for seed removal, and enables consumption of entire berries. Seedless varieties typically produce smaller berries than seeded cultivars unless treated with gibberellic acid (GA₃) to promote cell enlargement modern seedless breeding has progressively developed larger-berried seedless varieties reducing the size disadvantage. Seedless grapes now account for over 80% of table grape consumption in North America and Western Europe, representing a complete market transformation over 50 years.
Seedless_Table_Grape_Scion_Role captures the dominant commercial table grape category in developed markets, distinguishing varieties by the developmental mechanism producing seedlessness (stenospermocarpic versus parthenocarpic subclasses). The market shift toward seedlessness has transformed table grape breeding and production, with essentially all new commercial varieties being seedless. Seeded varieties retain importance in Asian markets (Japan, China, Korea) where berry size commands premium prices and seeds are accepted, and in markets where traditional seeded cultivars (Italia, Red Globe) maintain consumer loyalty. Embryo rescue techniques enable crossing seedless varieties that would otherwise be breeding dead-ends, accelerating seedless variety development.
kernlose Tafeltraube
kernlose Traube
pip-free grape
samenlose Tafeltraube
seedless grape
seedless table grape
Ledbetter CA, Ramming DW (1989) Seedlessness in grapes, Horticultural Reviews 11: 159-184 Creasy GL, Creasy LL (2009) Grapes, CABI Perl A et al. (2000) Seedless grapes, in Molecular Biology and Biotechnology of the Grapevine, Springer pp.: 327-351 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 430-450
Seedless Table Grape Scion Role
Seedless table grapes represent over 80% of global table grape production, with Thompson Seedless (Sultanina) being the most widely planted seedless cultivar worldwide. Seedless berries contain seed traces ranging from barely perceptible (Thompson Seedless, 8-15 mg seed trace weight) to noticeable (Flame Seedless, 25-40 mg). The sdI locus on chromosome 18 controls stenospermocarpy in most commercial seedless cultivars. Berry weight in seedless cultivars ranges from 2-4 g without treatment to 6-10 g with gibberellin application.
A reproductive disposition that is the inherited capacity of a Vitis vinifera plant to produce berries lacking fully developed seeds, resulting from disruption of normal seed development through embryo abortion, endosperm failure, or fruit development without fertilization, yielding fruit with seed traces, rudimentary seeds, or no seeds.
Seedlessness_Disposition serves as the parent class for different mechanisms producing seedless fruit. Three distinct pathways exist: stenospermocarpy (seed abortion after fertilization), parthenocarpy (fruit development without fertilization), and GA-induced seedlessness (hormone manipulation of seeded cultivars). Commercial seedlessness is almost exclusively stenospermocarpic. Seedlessness affects berry size, requiring compensatory treatments for acceptable fruit size.
Apyrenie
Kernlosigkeit
Samenlosigkeit
apyreny
seed abortion trait
seedless berry production
Ledbetter CA, Ramming DW (1989) Seedlessness in grapes. Horticultural Reviews 11:159-184 Doligez A et al. (2002) Genetic mapping of grapevine (Vitis vinifera L.) applied to the detection of QTLs for seedlessness and berry weight. Theoretical and Applied Genetics 105:780-795
Seedlessness Disposition
Seedling number 6 within cross Gf.1993-22 (yielding the complete identifier Gf.1993-22-6) designates the individual seedling that was selected through successive screening stages from among 380 siblings — at germination all 380 received sequential seedling numbers 1-380 for tracking through Phase 1 evaluation, with 95% eliminated during the first 5 years of field screening for disease resistance and basic fruit quality, seedling number 6 advancing through Phase 2 replicated trials under the same identifier, and ultimately receiving the cultivar name Calardis Blanc upon official registration while retaining Gf.1993-22-6 as its permanent breeding record identifier.
A breeding code that uniquely identifies an individual grapevine seedling within a cross family, typically appended to the cross number as a sequential identifier assigned at germination or at the time of field planting, distinguishing each genetically unique individual produced from a single controlled pollination event.
Seedling_Number is the most granular level of breeding program identification, distinguishing individual genotypes within a family of full siblings sharing the same parents. Each seedling from a controlled cross is genetically unique due to sexual recombination, and the seedling number provides the persistent individual identifier through the entire selection pipeline from germination to potential cultivar release: given that a single cross may produce hundreds of seedlings and a breeding program may generate 5,000-20,000 seedlings annually, the seedling number is essential for maintaining individual-level data integrity across years of phenotypic evaluation. The vast majority of seedling numbers are retired when genotypes are eliminated during screening, with only 0.01-0.1% ultimately associated with released cultivars.
Einzelpflanzen-Nummer
Nachkommen-Kennung
Sämlingsnummer
individual seedling code
progeny individual number
seedling identifier
Seedling Number
Seedlings from a Chardonnay × Rpv3-donor cross bear Seedling_Selection_Roles, with DNA extracted at the cotyledon stage for marker analysis identifying the 50% carrying Rpv3 resistance—only marker-positive individuals advanced to nursery transplanting, halving population size before field investment. A seedling population segregating for Run1 powdery mildew resistance serves in Seedling_Selection_Roles, with molecular screening eliminating susceptible individuals during the first growing season. Vigorous seedlings from a Pinot Noir improvement cross bear Seedling_Selection_Roles based on phenotypic evaluation of growth characteristics, with weak or abnormal individuals culled before field establishment.
A selection role that inheres in a grapevine genotype during the initial evaluation phase following seed germination, where young seedlings from controlled crosses are assessed for basic viability, early growth characteristics, and increasingly for molecular markers linked to target traits. Seedling selection represents the first opportunity to reduce breeding population size, eliminating individuals lacking essential characteristics before the substantial investment of field establishment. Traditional seedling selection focused on vigor, growth habit, and freedom from obvious defects, with limited predictive value for mature vine performance. Contemporary marker-assisted seedling selection (MASS) enables identification of individuals carrying target alleles (resistance genes, quality-linked markers) at the seedling stage, dramatically improving selection efficiency by eliminating marker-negative individuals before field planting. Seedling selection may reduce populations from thousands to hundreds within the first growing season, focusing subsequent resources on genotypes with confirmed genetic potential. The seedling phase typically spans germination through the first 1-2 years, with selected individuals advanced to vineyard establishment for field evaluation.
Seedling_Selection_Role captures the critical early-stage selection that determines which individuals receive the substantial investment of field establishment and long-term evaluation. The transformation from phenotypic seedling selection (limited predictive value) to marker-assisted seedling selection (high accuracy for marked traits) has dramatically improved breeding efficiency, with programs now routinely screening thousands of seedlings for resistance markers before any field planting. The seedling stage offers practical advantages for molecular selection—DNA extraction is straightforward, plants are easily handled, and elimination decisions have minimal resource impact. Seedling selection limitations include the juvenile-mature correlation gap (seedling characteristics may poorly predict adult performance for complex traits), necessitating continued field evaluation for traits not amenable to early molecular selection.
Frühselektion
Jugendauslese
Keimlingsselektion
Sämlingsselektion
early selection
juvenile selection
nursery selection
seedling selection
Seedling Selection Role
Each of the 150+ F1 individuals from GF.GA-47-42 × Villard Blanc bears a Segregating_Population_Member_Role, with unique recombinant genotypes (documented through SSR and SNP markers) and individual phenotype records (disease resistance scores, phenology dates, berry composition) contributing to the statistical detection of Rpv3 and other QTL. A Cabernet Sauvignon × Norton population member serves in a Segregating_Population_Member_Role with Pierce's disease resistance phenotype correlated to parental allele inheritance at PdR-linked markers—resistant individuals carrying Norton alleles at resistance loci. An individual from the Syrah × Grenache population bears a Segregating_Population_Member_Role with documented genotype and multi-year phenology, berry weight, and sugar accumulation data contributing to QTL mapping for fruit development traits.
A mapping population role that inheres in an individual grapevine genotype as a member of a segregating population, serving as a data point in genetic mapping analysis through its unique combination of parental alleles and corresponding phenotypic measurements. Each population member carries a distinct recombinant genotype created by meiotic recombination in the parental gametes, with marker genotyping revealing the mosaic of parental chromosome segments inherited by that individual. Phenotypic measurements on population members, correlated with their marker genotypes, enable detection of marker-trait associations identifying QTL—genomic regions where marker alleles from the trait-contributing parent cosegregate with favorable phenotypes. The analytical power of mapping analysis depends on population size (number of individuals providing marker-trait correlation data), marker density (resolution of genetic map), and phenotyping accuracy (precision of trait measurements), with each population member contributing information proportional to these factors. Population members may be maintained long-term for repeated phenotyping across environments and years, building comprehensive datasets supporting robust QTL detection.
Segregating_Population_Member_Role emphasizes the individual-as-data-point function essential for genetic mapping, where each population member's unique genotype-phenotype combination contributes to trait-marker association detection. Population members are typically maintained as clonally propagated individuals, enabling repeated phenotyping across years and environments that strengthens QTL detection by distinguishing genetic effects from environmental variation. The contribution of each member depends on its recombination pattern—individuals with recombination breakpoints near QTL provide mapping resolution, while those inheriting intact parental segments contribute less to fine-mapping. High-throughput genotyping has transformed population member characterization, with SNP arrays or sequencing providing thousands to millions of marker data points per individual, enabling high-resolution mapping and genomic selection approaches that exploit all population variation simultaneously.
Kartierungsnachkomme
Populationsmitglied
Segregant
mapping progeny
population member
recombinant individual
rekombinantes Individuum
segregant
Adam-Blondon AF et al. (2011) Genetic mapping in grapes, CRC Press pp.: 119-159 Zyprian E et al. (2016) Quantitative trait loci, Molecular Genetics and Genomics 291: 1573-1594 Fechter I et al. (2012) Candidate genes for downy mildew QTL, Molecular Genetics and Genomics 287: 159-174 Riaz S et al. (2011) Powdery mildew resistance, Theoretical and Applied Genetics 122: 1059-1073
Segregating Population Member Role
Selection criteria for Cabernet Sauvignon clones at ENTAV prioritized small berry size ( 60 AU at 280 nm), tannin concentration exceeding 3 g/L in micro-vinifications, and absence of GFLV, GFkV, GLRaV-1, and GLRaV-3 — explicitly weighting quality parameters above yield in the final ranking algorithm. Selection criteria for Welschriesling clones at Klosterneuburg emphasized consistent must weights of 17-19° KMW across vintages, total acidity retention above 7 g/L at harvest, compact cluster architecture minimizing Botrytis susceptibility (OIV 204 scores of 3-5), and balanced vigor (pruning weight 0.4-0.8 kg/m cordon) suitable for the region's loess-dominated soils.
Selection criteria is a selection history entity that records the specific phenotypic, agronomic, oenological, and phytosanitary attributes used to evaluate and rank candidate vines during the clonal selection process, including yield parameters (cluster number, cluster weight, berry weight), quality indicators (sugar accumulation, acidity balance, phenolic content, aromatic profile), disease tolerance metrics (virus load, fungal susceptibility), and viticultural performance attributes (vigor, fertility, ripening uniformity) against which candidate clones were assessed and compared.
Selection criteria reflect the viticultural and oenological priorities of the selecting institution and region, creating inherent biases in clone populations that should be understood when evaluating clonal material for different contexts. French ENTAV selections historically emphasized strict health status (virus-free) and moderate yield, German Geisenheim/Geilweilerhof programs balanced yield with must quality (sugar, acidity), while Italian selections often prioritized phenolic content and color intensity for red cultivars. The weighting of individual criteria within multi-attribute decision frameworks varies — some programs employ formal weighted scoring matrices, while others rely on expert panel consensus. Critically, selection criteria applied under one set of environmental conditions may not identify optimal clones for different growing regions, as genotype-by-environment interactions can alter clone ranking across sites. The evolution of selection criteria over time reflects changing market demands — programs initiated in the 1960s-1970s often prioritized yield improvement, while contemporary programs increasingly emphasize drought tolerance, phenolic quality, and disease resistance in response to climate change and sustainability concerns.
Auslesekriterien
Bewertungskriterien
Klonbewertungsparameter
Selektionskriterien
Selektionsmerkmale
clone assessment criteria
evaluation criteria
screening parameters
selection attributes
selection targets
Mannini F (2004) Clonal selection in grapevines. Acta Horticulturae 652: 21-27 Anderson MM et al. (2008) Viticultural evaluation of French and California Pinot Noir clones. American Journal of Enology and Viticulture 59(2): 188-193 Regner F et al. (2003) Clonal selection and clone identification of Austrian grapevine varieties. Mitteilungen Klosterneuburg 53: 172-180
Selection Criteria
Selection dates for Cabernet Sauvignon clone 337 (ENTAV) record initial vine identification in 1972 at Domaine de Chevalier, Pessac-Léognan, first propagation cuttings taken in spring 1973, trial vineyard establishment in 1975, preliminary performance data collection spanning vintages 1978-1985, virus-free status confirmed in 1982, and official ENTAV agrément (certification) granted in 1988 — a 16-year selection-to-release timeline. Selection dates for Zweigelt clone Zw-70 (Klosterneuburg) document the screening campaign of 1970 (Jahrgang der Erstauswahl), propagation in 1971, trial establishment at three Austrian sites in 1973, and commercial release following the 1988 vintage evaluation — an 18-year development cycle reflecting the extended observation periods typical of continental climate evaluation programs.
Selection date is a selection history entity that records the temporal reference points in the clonal selection process, including the date of initial mother vine identification, the year(s) of field observation and screening, the date of first vegetative propagation from the candidate vine, and subsequent milestone dates through the evaluation and certification timeline, expressed as calendar dates or vintage years, establishing the chronological framework of the selection program.
Selection dates provide essential context for interpreting clonal performance data, as the climatic conditions during the evaluation period may not be representative of long-term averages — clones selected during warm vintage sequences may perform differently under cooler conditions, and vice versa. The distinction between the date of initial identification (often a single vintage observation) and the date of first propagation (when the clonal line is physically established) is important, as the intervening period typically involves one to three additional growing seasons of confirmatory observation. Historical selection dates are frequently approximate, particularly for selections made before formal documentation protocols were established — many German clonal selections from the 1950s-1960s record only the decade of initial identification. The total elapsed time from selection to certification typically ranges from 12-25 years, reflecting the combined requirements for field evaluation across multiple vintages (minimum 8-10 years), virus testing and elimination (2-5 years), and administrative certification processes (1-3 years). Climate change considerations have prompted some programs to establish new selection dates for re-evaluation of existing clones under contemporary growing conditions.
Auslesedatum
Identifizierungsdatum
Selektionsdatum
Selektionsjahr
Zeitpunkt der Selektion
clone identification date
date of selection
screening date
selection timeline date
selection year
Mannini F (2004) Clonal selection in grapevines. Acta Horticulturae 652: 21-27 Boidron R et al. (1995) Catalogue des variétés et clones de vigne cultivés en France, ENTAV-INRA, pp.: 15-25 Regner F et al. (2003) Clonal selection and clone identification of Austrian grapevine varieties. Mitteilungen Klosterneuburg 53: 172-180
Selection Date
The selection history of Riesling clone 239 (Geisenheim) records its identification in 1960 from an exceptionally healthy, high-quality vine in the Schloss Johannisberg vineyard, followed by 15 years of field evaluation (1962-1977) across three trial sites in the Rheingau and Pfalz, virus elimination through heat therapy in 1970, re-evaluation of thermotherapy-treated material from 1972, and final certification and release in 1980 as a virus-free clone combining moderate yield (8-10 t/ha) with excellent must weight potential (75-85° Oechsle). The selection history of Sangiovese clone R24 traces its origin to a 1965 mass screening of 500 vines at Fattoria di Montalcino, progressive narrowing to 12 candidate vines by 1970, single-vine propagation and replicated trial establishment in 1972, and 18 years of comparative evaluation before commercial release in 1990.
Selection history is a clonal selection record entity that documents the chronological sequence of events, decisions, and observations from the initial identification of a superior individual vine through all subsequent stages of evaluation, multiplication, and release, providing a complete provenance narrative that establishes the traceability and authenticity of a clonal line from its mother vine to certified nursery material.
Selection history documentation serves both scientific and legal functions — scientifically, it enables retrospective analysis of selection criteria and methodology, while legally it establishes the chain of custody required for varietal certification and plant variety protection. The typical selection history spans 15-30 years from initial identification to commercial release, reflecting the long evaluation periods necessary to characterize vine performance across multiple vintages and the time required for virus testing and elimination protocols. Historical selection records (pre-1950) are frequently incomplete, relying on institutional archives, personal correspondence, and ampelographic publications to reconstruct the selection narrative. The distinction between clonal selection (single-vine origin, vegetatively multiplied) and mass selection (population-based selection maintaining genetic diversity) should be clearly documented in the selection history, as the two approaches produce fundamentally different genetic structures in the resulting plant material. Modern selection programs increasingly document not only phenotypic observations but also molecular marker profiles at each stage, creating integrated genotypic-phenotypic selection histories.
Auslesegeschichte
Klonherkunftsnachweis
Selektionschronik
Selektionsgeschichte
Selektionsverlauf
clonal provenance record
clone origin history
clone selection history
selection chronology
selection narrative
Mannini F (2004) Clonal selection in grapevines. Acta Horticulturae 652: 21-27 Rühl EH et al. (2004) Grapevine clonal selection, evaluation, and management. In Proceedings of the International Symposium on Grapevine Growing, Commerce and Research, Acta Horticulturae 652: 29-34 Anderson MM et al. (2008) Viticultural evaluation of French and California Pinot Noir clones grown for production of sparkling wine. American Journal of Enology and Viticulture 59(2): 188-193
Selection History
Selection location for the ENTAV Cabernet Sauvignon clone evaluation program encompassed three primary trial sites: Domaine de l'Espiguette (Grau-du-Roi, sandy soil, Mediterranean climate, 3 m elevation), Domaine de Vassal (Marseillan, sandy-calcareous soil, Mediterranean), and Château Couhins (Villenave-d'Ornon, Graves gravel over clay, oceanic climate, 25 m elevation) — the multi-site design enabling assessment of clone-by-environment interactions across contrasting terroirs. Selection location for Riesling clone evaluation at Geisenheim encompassed trial vineyards in the Rheingau (loess over phyllite, 80-120 m, 49.9°N) and the Pfalz (limestone-marl, 150-180 m, 49.3°N), with complementary evaluation sites at Weinsberg (Württemberg) and Trier (Mosel) providing additional environmental contrast for the 15-year assessment period.
Selection location is a selection history entity that records the geographic site where the clonal evaluation trials, comparative field assessments, and performance testing were conducted, distinct from the selection origin (mother vine source), including the trial vineyard(s) coordinates, institution, region, elevation, soil type, climate zone, and rootstock used in the evaluation planting, providing essential environmental context for interpreting clone performance data and assessing the transferability of results to other growing regions.
Selection location is critically important for interpreting clone performance data because genotype-by-environment (G×E) interactions in grapevines can alter clone ranking across sites — a clone demonstrating superior quality at one location may prove mediocre or poorly adapted at another. Multi-site evaluation designs, standard practice in major national selection programs, provide partial characterization of G×E magnitude but cannot fully predict performance at new sites with different soil, climate, and management conditions. The soil type and climate zone of evaluation sites introduce systematic biases — clones selected exclusively on deep, fertile soils may underperform on shallow, calcareous terrains, while clones optimized for warm Mediterranean conditions may exhibit incomplete ripening in cool continental climates. Modern selection programs document evaluation site characteristics in comprehensive detail, including soil analysis, climate normals, rootstock identity, vine spacing, and management protocols, recognizing that this contextual information is essential for growers evaluating clonal material for their specific conditions. The trend toward participatory on-farm clone evaluation, where clones are tested under commercial management conditions across diverse grower sites, aims to extend the environmental range of selection location data beyond institutional trial vineyards.
Evaluierungsstandort
Klonprüfungsanlage
Prüfungsstandort
Selektionsstandort
Versuchsstandort
clone evaluation location
evaluation site
selection trial site
test vineyard
trial location
Rühl EH et al. (2004) Grapevine clonal selection, evaluation, and management. Acta Horticulturae 652: 29-34 Anderson MM et al. (2008) Viticultural evaluation of French and California Pinot Noir clones. American Journal of Enology and Viticulture 59(2): 188-193 Wolpert JA et al. (1994) Field performance of six Chardonnay clones in the Napa Valley. American Journal of Enology and Viticulture 45(4): 393-400
Selection Location
The selection method for Pinot Noir Dijon clones (ENTAV series 113-115, 667, 777, 828) followed the classical French three-phase protocol: Phase I — visual mass screening of 3000+ vines for health, yield, and cluster morphology over three vintages Phase II — propagation of 150 candidate vines with replicated micro-plot evaluation over 8 vintages including micro-vinification Phase III — virus elimination through heat therapy at 38°C for 60 days combined with meristem tip culture, followed by re-evaluation of sanitized material in certified trial blocks. The selection method for Grüner Veltliner at Klosterneuburg combined traditional ampelographic mass screening with SSR marker profiling (VVS2, VVMD5, VVMD7, VVMD27, VrZAG62, VrZAG79) to confirm varietal identity and detect potential off-types before committing resources to field evaluation.
Selection method is a selection history entity that records the systematic approach, techniques, and protocols employed to identify, evaluate, and isolate a superior individual vine from a cultivar population, including the type of selection (clonal, mass, polyclonal), the screening methodology (visual assessment, ampelographic scoring, molecular marker analysis), the experimental design of evaluation trials (replicated plots, multi-site comparisons), and any virus elimination procedures (heat therapy, meristem culture, somatic embryogenesis) applied during the selection process.
Selection methods have evolved substantially over the past century, from purely empirical visual selection of outstanding vines (pre-1950) through systematic replicated field evaluation (1960s-1990s) to integrated molecular-phenotypic approaches incorporating marker-assisted selection and genomic characterization (2000s-present). The fundamental methodological distinction between mass selection (selecting the best 5-20% of a population while maintaining genetic diversity) and individual clonal selection (propagating single-vine genotypes) carries profound implications for the resulting plant material — mass selections preserve population-level buffering capacity against environmental variation, while clonal selections offer genetic uniformity and predictability. Modern selection methods increasingly incorporate early-stage molecular screening to verify varietal identity, detect intercultivar hybridity, and assess genetic distance among candidates before the costly commitment to multi-year field evaluation. Virus elimination methods are integral to selection methodology and significantly influence clone performance, as thermotherapy and meristem culture can alter epigenetic states and occasionally produce phenotypic changes in the sanitized material relative to the original mother vine.
Auslesemethode
Selektionsmethode
Selektionsmethodik
Selektionsprotokoll
Selektionsverfahren
clone selection approach
screening methodology
selection procedure
selection protocol
selection technique
Mannini F (2004) Clonal selection in grapevines. Acta Horticulturae 652: 21-27 Rühl EH et al. (2004) Grapevine clonal selection, evaluation, and management. Acta Horticulturae 652: 29-34 Moncada X et al. (2006) Microsatellite genotyping of grapevine accessions for verification and genetic studies. Molecular Ecology Notes 6(1): 248-250
Selection Method
Selection number Gm 20-13 at the Geisenheim research station identifies a Pinot Noir clone selected from the Geisenheim clonal evaluation program — the Gm prefix denotes Geisenheim, 20 refers to the cultivar evaluation group (Pinot Noir), and 13 is the individual clone number within that group, with this selection number serving as the permanent identifier in the German clonal registry and international research databases even after the clone received the official designation Geisenheim Clone 20-13 in the Bundessortenamt descriptive variety list.
A breeding code assigned to a grapevine genotype that has been selected from a larger population based on superior performance for target traits, typically incorporating an institutional prefix and a sequential or hierarchical number that positions the selection within the breeding program's organizational framework.
Selection_Number serves as the institutional tracking identifier for genotypes that have been positively selected — whether from seedling populations in breeding programs or from field populations in clonal selection programs — and retained for further evaluation or propagation. The numbering systems are institution-specific and may encode varying levels of information: some systems embed the cultivar identity (e.g., Gm 20 = Pinot Noir group at Geisenheim), while others use purely sequential numbering. Selection numbers for clonal selections of established cultivars differ functionally from breeding selection numbers in that they identify variant clones within an existing cultivar rather than novel genotypes from controlled crosses.
Auslese-Kennung
Zuchtstamm-Nummer
breeder's selection code
selected genotype number
selection identifier
Bundessortenamt (2023) Beschreibende Sortenliste Reben Eibach R and Töpfer R (2015) Traditional grapevine breeding techniques, in Reynolds AG (ed) Grapevine Breeding Programs for the Wine Industry, Woodhead Publishing pp: 3-22
Selection Number
The selection origin of Chardonnay clone 96 (Dijon series) is documented as a single vine identified in a pre-phylloxera planting at Domaine Leflaive in Puligny-Montrachet, Côte de Beaune, Burgundy, France — an own-rooted vineyard established circa 1930 on Kimmeridgian limestone-marl soil, providing the genetic source for what became one of the most globally distributed premium Chardonnay clones. Selection origin of Blaufränkisch clone Bl-10 (Klosterneuburg) traces to a 60-year-old vine in the Goldberg vineyard of Deutschkreutz, Mittelburgenland, Austria, grown on loam-clay over schist at 220 m elevation on SO4 rootstock, selected from a population of approximately 2000 vines screened during the 1985-1988 evaluation campaign.
Selection origin is a selection history entity that records the geographic, institutional, and viticultural source from which a clonal line was initially identified and isolated, including the specific vineyard, estate, or experimental collection where the mother vine was located, the region and country of origin, the cultivar population or planting from which the selection was drawn, and the viticultural context (age, rootstock, soil type) of the original vine, establishing the foundational provenance of the clone.
Selection origin documentation is critical for understanding the genetic and epigenetic background of a clone, as the environmental conditions, vine age, and population context of the mother vine influence the phenotypic expression that prompted its initial selection. Old vineyards (>50 years) are preferentially targeted for clonal prospection because they have undergone decades of natural selection pressure for site adaptation and somatic mutation accumulation, potentially harboring unique genetic variants absent from younger plantings. The distinction between origin from commercial vineyards versus experimental collections or germplasm repositories is significant — commercial vineyard selections reflect real-world viticultural performance, while collection selections may have been maintained under atypical conditions. Accurate origin documentation also prevents nomenclatural confusion, as identical clone numbers from different selection programs (e.g., French ENTAV versus Italian Rauscedo numbering) designate entirely different genetic material. GPS coordinates of mother vine location are increasingly recorded in modern selection programs, enabling precise geographic provenance documentation.
Klonherkunft
Mutterrebenstandort
Selektionsursprung
Ursprungsanlage
clonal provenance
clone origin
mother vine origin
origin site
selection source
source vineyard
This P et al. (2006) Historical origins and genetic diversity of wine grapes. Trends in Genetics 22(9): 511-519 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties. Heredity 104(4): 331-340 Lacombe T et al. (2013) Large-scale parentage analysis in an extended set of grapevine cultivars. Theoretical and Applied Genetics 126(2): 401-414
Selection Origin
A seedling from a Riesling × resistance donor cross bears a Selection_Role as it progresses through nursery evaluation, field establishment, phenotypic characterization, and wine quality assessment—each stage involving selection decisions determining advancement or elimination. An advanced breeding line under final variety evaluation serves in a Selection_Role as a variety candidate, with multi-site trial data, wine competition results, and regulatory review determining registration approval. A genotype selected for marker-confirmed resistance and favorable phenotype bears a Selection_Role in marker-assisted breeding, with molecular selection supplementing traditional evaluation.
A breeding role that inheres in a grapevine genotype by virtue of its position in the selection process of breeding programs, where individuals are evaluated, compared, and chosen based on phenotypic performance or marker-assisted criteria to advance toward variety registration or serve as parents for subsequent breeding generations. Selection roles encompass the stages through which breeding material progresses—from initial seedling evaluation through field performance trials to final variety candidate selection—with selection intensity (proportion of individuals retained) typically increasing at each stage as population sizes decrease and evaluation criteria become more stringent. Grapevine selection operates across extended timeframes (15-25 years from cross to variety release), with selection decisions at each stage determining which genotypes advance and which are discarded, progressively enriching populations for favorable trait combinations. Selection may be phenotypic (based on observed characteristics), marker-assisted (using molecular markers linked to target traits), or genomic (using genome-wide marker information to predict breeding values), with contemporary programs increasingly integrating molecular tools with traditional phenotypic evaluation.
Selection_Role is subclassed by selection stage (Seedling_Selection for initial evaluation, Field_Selection for established vine evaluation, Progeny_Selection for offspring evaluation) reflecting the distinct contexts and criteria applied at different breeding program phases. Selection intensity varies by program and stage—initial seedling populations may number thousands with 5-10% advancement rates, while final variety candidates may involve dozens of lines with few achieving registration. The efficiency of selection (genetic gain per unit time) depends on heritability of target traits, selection intensity, accuracy of evaluation, and generation interval, with molecular tools improving accuracy and potentially reducing evaluation time. Selection decisions integrate multiple traits (disease resistance, yield, quality, adaptation), requiring index selection approaches that weight traits according to breeding objectives.
Auslese
Selektion
Selektionskandidat
evaluation genotype
selection
selection candidate
Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, in Reynolds A (ed) Grapevine Breeding Programs for the Wine Industry, Elsevier pp.: 3-22 Töpfer R et al. (2011) Molecular breeding in grapes, CRC Press pp.: 160-185 Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330
Selection Role
A traditional Burgundian Pinot Noir selection maintained by successive generations of vignerons bears a Selection_Status_Clone_Role reflecting Mass_Selection origin, the population derived from collecting cuttings from the best-performing vines across the domaine without individual genotype isolation. A ENTAV-INRA Pinot Noir clone serves in a Selection_Status_Clone_Role reflecting Clonal_Selection methodology, an individual vine identified, isolated, and evaluated through formal trial protocols before registration. A FPS Cabernet Sauvignon selection bears a Selection_Status_Clone_Role emphasizing Sanitary_Selection, where virus elimination through thermotherapy and meristem culture was the primary development objective applied to existing selections.
A clone role that inheres in a grapevine clone by virtue of the selection methodology through which it was identified and developed, distinguishing clones by the approaches used to isolate desirable genotypes from within-variety populations. Selection status classifies clones by the technical methods of their development—mass selection (collecting propagation material from multiple superior vines), clonal selection (isolating individual genotypes for evaluation), sanitary selection (prioritizing pathogen elimination), and genetic selection (prioritizing agronomic and enological characteristics)—with these approaches reflecting different historical periods, institutional capabilities, and selection objectives. The selection methodology influences clone characteristics mass selections retain population diversity, clonal selections offer genetic uniformity, sanitary selections prioritize health status, and genetic selections emphasize performance traits. Modern clone development typically integrates multiple selection dimensions—contemporary programs combine genetic evaluation (field trials, winemaking assessment) with sanitary procedures (virus testing, elimination treatment)—but clones may be classified by their primary selection emphasis.
Selection_Status_Clone_Role classifies clones by development methodology rather than regulatory status, capturing the technical approaches underlying clone creation. The selection status spectrum reflects historical evolution—mass selection (pre-20th century), clonal selection (systematized from 1920s-1960s), sanitary selection (developed 1960s-1980s), and integrated genetic-sanitary selection (contemporary standard)—though all methodologies remain relevant depending on context and objectives. The distinction between approaches is not always clear-cut clonal selection inherently requires identifying individual genotypes that may then undergo sanitary procedures, while sanitary selection may be applied to genotypes previously identified through clonal selection. The classification captures the primary emphasis of the selection program, recognizing that modern programs typically integrate multiple objectives.
Ausleseart
Klonentwicklungstyp
Selektionsstatus
development approach
selection methodology
selection status
selection type
Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340 Walter B, Martelli GP (1996) Clonal and sanitary selection of the grapevine, INRA Editions Huglin P, Schneider C (1998) Biologie et Écologie de la Vigne, Lavoisier
Selection Status Clone Role
The selective harvest criteria for Sauternes at Château d'Yquem specify three to seven successive picking passes (tries) through the vineyard over a 4-8 week period from late September through November, with each trie targeting clusters or portions of clusters at a specific stage of Botrytis cinerea noble rot development — pickers select only berries showing full pourriture noble (golden-brown shriveled berries with concentrated sugar of 350-450 g/L) while leaving unaffected green berries, partially botrytized berries, and berries showing grey rot (pourriture grise, the destructive form) for subsequent passes or rejection. Each trie yields only 200-500 liters per hectare (compared to 4,000-6,000 liters for a single-pass dry wine harvest), with the cumulative yield across all tries averaging 900 liters per hectare — equivalent to approximately one glass of wine per vine.
A harvest method description that documents the decision rules for selective picking — the practice of harvesting specific portions of the vineyard, individual clusters, or parts of clusters at different times or to different quality destinations based on assessed maturity, health status, or position within the canopy, specifying the selection parameters that enable differentiated treatment of fruit from a single vineyard block.
Selective_Harvest_Criteria captures the most sophisticated and labor-intensive form of grape harvesting, where fruit selection occurs at the individual berry or cluster level rather than the vine or block level. The practice ranges from simple two-tier selection (first pick of optimal clusters followed by second pick of remaining usable fruit) through the successive tries system of Sauternes and Tokaj (3-7 passes selecting progressively botrytized fruit) to the extreme selectivity of Trockenbeerenauslese production in Germany and Austria where individual botrytized berries are plucked by hand: the common principle is that differential harvest timing exploits the natural asynchrony of ripening and botrytis development within and between clusters to isolate fruit at the precise developmental stage desired for the target wine style. Beyond botrytized wine production, selective harvest criteria are increasingly applied in premium dry wine production through practices such as harvesting sun-exposed and shaded canopy sides separately, harvesting first and second clusters separately (second clusters typically ripen 5-10 days later), or harvesting young vines and old vines within the same block into different lots.
Selektions-Erntekriterien
Selektivlese-Kriterien
differential harvest protocol
differenzierte Lese-Protokoll
selective picking criteria
triage criteria
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1, 2nd Ed, Wiley pp: 235-270 Château d'Yquem (2023) Harvest documentation Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 280-295 Jackson RS (2020) Wine Science: Principles and Applications 5th Ed, Academic Press pp: 295-340
Selective Harvest Criteria
The selector name for the Dijon Pinot Noir clones (113, 114, 115, 667, 777, 828) is recorded as Raymond Bernard of ENTAV-INRA Colmar, who conducted the systematic field prospection across Burgundy's Côte d'Or vineyards from 1960 to 1975, establishing the foundation for the most globally influential Pinot Noir clonal program. The selector name for Blaufränkisch clone Bl-2 is documented as Prof. Dr. Ferdinand Regner, Höhere Bundeslehranstalt und Bundesamt für Wein- und Obstbau, Klosterneuburg, whose team integrated ampelographic evaluation with SSR marker verification during the 1990-2005 selection campaign across Burgenland mother vine populations.
Selector name is a selection history entity that records the identity of the individual researcher, viticulturist, or institutional team responsible for conducting the clonal selection, including the name of the principal selector or team leader, their institutional affiliation, and their professional role, establishing personal and institutional accountability for the selection and providing attribution essential for scientific citation, regulatory traceability, and historical documentation of clonal selection programs.
Selector name documentation serves multiple functions beyond simple attribution. It enables reconstruction of the selection philosophy and methodology through the selector's published works and known approaches, facilitates communication between researchers working with the same clonal material across institutions and countries, and provides the human link in certification chain-of-custody documentation. Many historically significant clonal selections are identified informally by their selector's name (e.g., "Bernard clones" for the Dijon series, "Entav clones" by institutional affiliation), underscoring the role of individual expertise in clone identification. The distinction between the individual who physically identified the mother vine in the field and the institutional authority that supervised the subsequent evaluation program should be documented, as these may differ — field prospectors, laboratory technicians, and program directors each contribute distinct expertise to the selection process. In collaborative international selection programs, multiple selector names may be recorded reflecting contributions at different phases (field identification, laboratory evaluation, trial management, certification).
Klonselekteur
Name des Selekteurs
Selekteur
Selektionsverantwortlicher
Selektionär
clone selector
selecting institution representative
selecting researcher
selection team
selector identity
Boidron R et al. (1995) Catalogue des variétés et clones de vigne cultivés en France, ENTAV-INRA, pp.: 10-20 Regner F et al. (2003) Clonal selection and clone identification of Austrian grapevine varieties. Mitteilungen Klosterneuburg 53: 172-180 Mannini F (2004) Clonal selection in grapevines. Acta Horticulturae 652: 21-27
Selector Name
A 20 cm cane segment of Fercal rootstock (difficult-rooting V. berlandieri hybrid) harvested in September with basal lignification but green shoot tips bears a Semi_Hardwood_Cutting_Role, prepared with basal IBA treatment and rooted under protected conditions combining elements of mist propagation (humidity maintenance) and conventional nursery beds (reduced misting frequency compared to full softwood protocol). A semi-hardwood cutting of Riesling serves in a Semi_Hardwood_Cutting_Role for autumn propagation when dormant budwood is not yet available but softwood material has matured beyond optimal green cutting condition. A semi-hardwood cutting of Muscadinia rotundifolia bears a Semi_Hardwood_Cutting_Role in specialized propagation of this difficult-rooting species, where intermediate tissue maturity improves success over fully dormant material.
A cutting role that inheres in a stem segment harvested from partially matured shoot tissue during the transition period between active growth and full dormancy, typically late summer to early autumn (August-October in the Northern Hemisphere) when basal internodes have begun lignification while shoot tips remain semi-succulent. Semi-hardwood cuttings (also termed semi-ripe or intermediate cuttings) occupy the physiological middle ground between softwood and hardwood, offering some advantages of each—greater structural stability than softwood (reducing mist system dependency) while retaining higher meristematic activity than fully dormant hardwood (potentially improving rooting for difficult genotypes). The semi-hardwood stage is characterized by partial periderm development, intermediate carbohydrate content, and buds that have formed but not yet achieved full cold acclimation. Semi-hardwood cutting propagation is less common than either hardwood or softwood approaches in commercial grapevine production but finds application for specific genotypes, for extending the propagation season beyond the traditional dormant window, and in research contexts requiring propagation during late summer-autumn.
Semi_Hardwood_Cutting_Role represents an intermediate category less standardized than hardwood or softwood approaches, with timing and tissue selection varying by genotype, climate, and propagation facility capabilities. The physiological basis for intermediate-stage advantages involves the balance between carbohydrate reserves (increasing with maturity) and cellular dedifferentiation capacity (decreasing with lignification) for some genotypes, the optimum occurs at partial maturity rather than either extreme. Semi-hardwood propagation may extend production seasons in nurseries with appropriate facilities, enabling harvest when hardwood is not yet fully dormant or when softwood has progressed beyond optimal juvenile condition. The category is more commonly applied to other woody crops (Camellia, Rhododendron, Ilex) than to grapevines, where the hardwood/softwood dichotomy dominates commercial practice.
Halbreifsteckling
halbverholzter Steckling
intermediate cutting
late summer cutting
semi-hardwood stem cutting
semi-ripe cutting
teilverholzter Steckling
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 370-374 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press
Semi Hardwood Cutting Role
A semi-lignified one-year cane on Vitis vinifera cv. Merlot examined in late October, showing brown woody appearance for basal 60cm (approximately 60% of total length) indicating completed lignification in early-formed internodes, brown-green appearance for middle 30cm indicating active lignification in progress, and green appearance with some brown streaking for apical 30cm indicating early-stage lignification just beginning, firmness testing showing woody rigidity in basal brown section, moderate firmness in brown-green middle, and retained flexibility in green apex, representing normal mid-autumn development pattern expected to progress to full or near-full lignification by dormant season pruning time in December if favorable weather continues.
A one-year cane in intermediate stages of lignification, characterized by mixed tissue development showing partial brown woody appearance interspersed with green or brown-green sections, moderate firmness with retained flexibility exceeding fully lignified wood, ongoing secondary cell wall development not yet complete, and transitional status between active shoot and mature cane representing incomplete wood maturation typical of mid-to-late autumn depending on climate and cultivar. Semi-lignified one-year canes represent normal developmental progression toward full maturation in regions with adequate growing season length, but indicate problematic incomplete ripening in short-season climates where early winter onset prevents completion of lignification, requiring management intervention including selective removal during pruning, crop reduction in following seasons, or consideration of earlier-maturing cultivars better adapted to site thermal conditions.
Semi-lignified one-year canes are normal and expected during autumn transition from active growth to dormancy, with the critical question being whether adequate time and thermal units remain for completion of lignification before winter cold arrival. In warm climates with extended autumns, semi-lignified canes observed in October-November will typically progress to full lignification by December-January pruning time. In cool climates or short seasons, semi-lignified status persisting into late autumn raises concerns about incomplete maturation, cold damage risk, and poor spring performance, potentially triggering decisions to remove semi-lignified portions during pruning. The lignification gradient from base to tip reflects the temporal sequence of internode formation and maturation, with basal internodes completing development first and apical internodes last or not at all if growing season ends prematurely.
halb ausgereifte Rute
half-ripened cane
partially woody cane
teilverholzte Rute
transitional lignification cane
Semi Lignified One Year Cane
Characterization of Semi_Upright_Growth_Disposition at the University of Bordeaux documented shoot architecture in cultivars with intermediate-upright growth habit. Reference cultivars 'Pinot Noir', 'Syrah', 'Grenache', and 'Tempranillo' were assessed on standard VSP trellis with measurement of shoot angles before and after positioning operations. Semi_Upright_Growth_Disposition cultivars showed consistent intermediate characteristics. 'Pinot Noir' exhibited typical semi-upright growth with mean shoot angle of 28° from vertical at 50 cm before positioning, requiring two tuck/position passes to achieve VSP geometry, and maintaining positioned angle of 12° through the season. Natural (unpositioned) shoot tip orientation was 35° from vertical, indicating moderate but incomplete gravitropic correction. 'Syrah' showed similar Semi_Upright_Growth_Disposition (32° at 50 cm) with slightly greater positioning requirement. 'Grenache' displayed the upper range of semi-upright (38° at 50 cm), transitioning toward horizontal growth habit. Internode stiffness averaged 2.0 N·mm² in semi-upright cultivars (intermediate between upright at 2.8 and trailing at 1.6 N·mm²). Positioning labor requirement was 28-38 hours/ha/season (2-3 passes), intermediate between upright (12-18 hours) and trailing (45-65 hours). Training system trials confirmed that Semi_Upright_Growth_Disposition cultivars performed well on both VSP (with positioning) and Smart-Dyson ballerina systems that accommodate their natural intermediate orientation. The flexibility was reflected in global planting patterns, with semi-upright cultivars like 'Pinot Noir' and 'Syrah' successfully grown on diverse training systems across wine regions.
A Semi_Upright_Growth_Disposition is a Shoot_Orientation_Disposition that inheres in Vitis vinifera shoots and is realized through moderate negative gravitropism producing shoots that grow at intermediate angles of 20-40° from vertical, with shoot tips showing some ascending tendency but without the strong vertical orientation of upright cultivars. This disposition represents the intermediate-upright range of shoot orientation, characteristic of cultivars that grow predominantly upward but with sufficient deviation from vertical to require some positioning intervention for optimal VSP canopy geometry. Semi-upright cultivars are compatible with VSP training systems but benefit from catch wire positioning or tucking to achieve fully vertical canopy presentation, representing moderate labor requirement between strongly upright (minimal positioning) and horizontal/trailing types (intensive positioning). The mechanical and physiological basis involves intermediate internode stiffness and gravitropic response strength, producing shoots that initially grow upward but gradually deviate from vertical as they elongate and gain weight. Semi_Upright_Growth_Disposition provides flexibility in training system selection, as these cultivars can be successfully managed in VSP with positioning, in sprawl systems accepting their natural spread, or in divided canopy systems benefiting from their intermediate orientation.
Semi_Upright_Growth_Disposition is_a Shoot_Orientation_Disposition representing intermediate-upright growth (20-40° from vertical). Characterized by moderate gravitropism and intermediate internode stiffness. Examples include 'Pinot Noir', 'Syrah', 'Grenache', 'Tempranillo', 'Sangiovese'. Compatible with VSP (with positioning) and various other systems. Moderate positioning labor requirement (2-3 passes). Provides training system flexibility. Represents the most common growth habit category among commercial cultivars.
Halbaufrechte-Wuchs-Disposition
Mittlerer-Triebwinkel
Mäßig-Aufrechter-Wuchs
intermediate-erect growth
moderately upright
semi-vertical habit
Carbonneau A (1995) La surface foliaire exposée potentielle—guide pour sa mesure, Progrès Agricole et Viticole 112:204-212 Smart RE (1988) Shoot spacing and canopy light microclimate, American Journal of Enology and Viticulture 39:325-333 Jackson DI, Lombard PB (1993) Environmental and management practices affecting grape composition and wine quality—a review, American Journal of Enology and Viticulture 44:409-430
Semi Upright Growth Disposition
Leaf senescence in grapevine initiates 2-4 weeks post-harvest in most cultivars, with chlorophyll content declining from 350-450 µg/cm² to below 50 µg/cm² over 25-40 days. Tempranillo exhibits early senescence disposition with 50% chlorophyll loss by late September in Rioja, while Mourvèdre retains green leaves until mid-November in Provence. The senescence-associated gene VvNAC26 shows 8-15 fold upregulation during leaf yellowing.
A grapevine disposition that is the inherited capacity of a Vitis vinifera plant to undergo programmed developmental decline of leaves and other annual tissues at season end, involving chlorophyll degradation, macromolecule catabolism, nutrient remobilization to perennial organs, and eventual abscission.
Senescence_Disposition serves as the parent class for autumn decline processes. Unlike stress-induced premature senescence, developmental senescence is a genetically programmed process optimized by natural selection for nutrient recovery and cold hardiness preparation. The process involves orderly dismantling of chloroplasts, protein degradation via autophagy, and phloem loading of remobilized nutrients.
Blattaltungsdisposition
Seneszenzverhalten
autumn senescence capacity
herbstliche Abbauprozesse
leaf aging disposition
seasonal decline trait
Lim PO, Kim HJ, Nam HG (2007) Leaf senescence. Annual Review of Plant Biology 58:115-136 Ferrara G et al. (2015) Grapevine leaf senescence: a review. Vitis 54:125-133
Senescence Disposition
Senescence nutrient recovery assessment comparing early versus late defoliation timing in Merlot reveals the importance of recovery duration for nutrient salvage, with natural autumn senescence (6-week gradual process) achieving nitrogen recovery of 74% (leaf N declining from 2.6% to 0.7% before abscission), while early frost-induced rapid senescence (1-week process) achieves only 45% recovery (leaf N at abscission still 1.4%), with the 29 percentage point difference representing nitrogen that must be replaced through soil uptake or reserve depletion the following spring, demonstrating how environmental conditions affecting senescence duration impact nutrient cycling efficiency.
Senescence nutrient recovery quality is a nutrient remobilization efficiency quality that represents the effectiveness of retrieving nutrients from senescing tissues before abscission or death, characterizing the salvage processes that recover mobile nutrients from leaves, fine roots, and other ephemeral tissues, converting what would be nutrient losses into resources available for redistribution to persistent or developing vine structures. Within the BFO framework, this quality specifically addresses the senescence-associated dimension of nutrient remobilization, distinguishing recovery from dying tissues from mobilization of stored reserves while recognizing that senescence represents a programmed developmental process during which substantial nutrient recovery occurs through coordinated degradation of cellular components and phloem export of released nutrients. The ontological significance lies in senescence nutrient recovery determining the fraction of ephemeral tissue nutrients retained within the vine versus lost through abscission, where efficient recovery enables nutrients invested in seasonal tissues (leaves, fine roots, fruit-bearing shoots) to be recycled for future use rather than requiring replacement through soil uptake, substantially affecting the vine's nutrient budget and fertilizer requirements. The cellular processes underlying senescence nutrient recovery include chloroplast dismantling (releasing nitrogen from Rubisco and other proteins, magnesium from chlorophyll), membrane breakdown (releasing phosphorus from phospholipids), protein degradation (releasing amino acids for transport), and nucleic acid hydrolysis (releasing nitrogen and phosphorus), with the products loaded into phloem for export before cell death eliminates transport capacity. The temporal dynamics of senescence recovery show a critical window during which nutrients must be exported before vascular connections sever, with premature senescence (from stress, disease, or early frost) potentially reducing recovery efficiency by shortening the export period, and with gradual senescence allowing more complete nutrient retrieval than abrupt tissue death. The environmental factors affecting recovery efficiency include temperature (moderate temperatures supporting continued phloem function), water status (adequate hydration maintaining transport capacity), and nutrient demand from sinks (strong sink activity promoting export from sources), with post-harvest conditions particularly important for leaf senescence recovery that supports reserve accumulation before dormancy. Understanding senescence nutrient recovery quality enables optimization of post-harvest vineyard management supporting nutrient recovery, assessment of stress impacts on nutrient salvage, and appreciation of the developmental coordination required for efficient nutrient cycling in perennial vines.
Senescence_Nutrient_Recovery_Quality addresses the salvage of nutrients from dying tissues before loss through abscission. This quality determines the fraction of seasonal tissue investment recoverable for future vine use. The class supports post-harvest management optimization through understanding of senescence recovery requirements and environmental dependencies.
Blattnährstoffresorption
Nährstoffbergung
Resorptionseffizienz
Seneszenz-Nährstoffrückgewinnung
leaf nutrient resorption
resorption efficiency
senescence recovery
Verdenal T et al. (2021) Understanding and managing nitrogen nutrition in grapevine: a review. OENO One 55(1): 1-43 Zapata C et al. (2004) Partitioning and mobilization of starch and N reserves in grapevine (Vitis vinifera L.). Journal of Plant Physiology 161(9): 1031-1040
Senescence Nutrient Recovery Quality
Syrah vineyard showing senescent feeder roots in October, soil cores revealing feeder roots with brown coloration, collapsed morphology, loss of root hairs, microscopic examination showing cytoplasmic degradation, organelle breakdown, membrane dysfunction, FDA viability staining negative indicating loss of membrane integrity, isotope tracing showing zero or minimal uptake confirming functional cessation, senescent feeder roots 6-11 months post-initiation representing 28% of total feeder root population, senescent feeder roots demonstrating normal terminal phase of ephemeral feeder root lifespan requiring continuous replacement
Senescent feeder root is feeder root undergoing programmed senescence showing declining function and approaching death, typically 4-12 months post-initiation representing terminal phase of ephemeral feeder root lifespan, characterized by progressive cytoplasmic degradation with organelle breakdown, membrane dysfunction reducing selective permeability and transport capacity, declining turgor reflecting osmotic imbalance, loss of absorption function with minimal or zero uptake, and visible morphological changes including tan to brown coloration, collapsed appearance, root hair loss, and tissue disorganization, with senescent feeder roots showing orderly dismantling of cellular structures through programmed process distinct from pathological necrosis, accumulating phenolic compounds and other metabolites creating brown pigmentation, and demonstrating that senescence represents normal terminal phase of ephemeral feeder root life requiring continuous new root production at tips maintaining functional feeder root population despite individual root brevity, and contributing substantially to soil organic matter input through high turnover with senescing feeder roots providing 30-60% of root system annual carbon input to soil supporting rhizosphere microbial communities and soil formation. Senescent feeder root morphological changes includes= showing color change (senescent feeder roots progressing from white young through tan mature to brown senescent), exhibiting shriveled appearance (turgor loss causing feeder roots to collapse and wrinkle losing cylindrical form), demonstrating root hair loss (root hairs degenerating and sloughing leaving bare root surface), revealing surface irregularities (senescing tissue developing irregular texture, potential breaks or lesions), and displaying tissue discoloration (browning from phenolic accumulation and oxidation creating characteristic appearance). Senescent feeder root cytoplasmic degradation includes= showing organelle breakdown (mitochondria, plastids, ER, ribosomes progressively degrading during senescence), exhibiting vacuole changes (vacuoles expanding, coalescing, or degrading as cytoplasm breaks down), demonstrating autophagy activation (cellular self-digestion pathways activated consuming cellular components), revealing protein degradation (proteases breaking down structural and functional proteins), and displaying membrane deterioration (progressive loss of membrane integrity and selective permeability). Senescent feeder root functional decline includes= showing reduced absorption (water and nutrient uptake declining progressively as membrane and metabolic function deteriorate), exhibiting decreased respiration (oxygen consumption declining as mitochondria degrade reducing ATP production), demonstrating turgor loss (osmotic imbalance and membrane damage causing progressive turgor decline and collapse), revealing cessation of growth (senescing feeder roots no longer elongating or producing laterals), and displaying unresponsiveness (senescing roots losing capacity to respond to environmental signals or plant demands). Senescent feeder root biochemical changes includes= showing protease activation (protein degradation enzymes upregulated during programmed senescence), exhibiting nuclease activity (DNA and RNA degrading as senescence progresses), demonstrating oxidative stress (reactive oxygen species accumulating potentially triggering or accelerating senescence), revealing phenolic accumulation (synthesis or release of phenolic compounds creating brown pigmentation), and displaying nutrient remobilization (nitrogen, phosphorus, and other nutrients potentially recovered and transported to plant before complete death). Senescent feeder root timing and progression includes= showing variable onset (senescence typically beginning 3-8 months after root initiation though highly variable), exhibiting progressive decline (senescence proceeding over days to weeks from onset to complete functional loss), demonstrating order-dependent patterns (higher-order feeder roots showing shorter lifespans and earlier senescence than lower-order), revealing environmental effects (stress accelerating senescence while favorable conditions potentially extending functional period), and displaying inevitable progression (senescence irreversible once initiated proceeding to root death). Senescent feeder root population dynamics includes= showing continuous occurrence (senescence continuously occurring in older feeder roots while new roots produced at tips), exhibiting spatial patterns (senescence concentrated in older distal portions of root branches and deeper or drier soil zones), demonstrating balance with production (senescence rate determining feeder root mortality balancing with production determining net population change), revealing turnover contribution (senescing feeder roots representing roots exiting functional pool requiring replacement), and displaying seasonal patterns (senescence potentially peaking late summer-fall and winter when conditions stressful). Senescent feeder root carbon cycling includes= showing organic matter input (senescing and dead feeder roots contributing labile carbon to soil), exhibiting decomposition (dead feeder root tissues decomposing rapidly due to thin walls and high nitrogen content), demonstrating microbial substrate (feeder root senescence providing substrate for rhizosphere microorganisms), revealing nutrient release (feeder root decomposition releasing nitrogen, phosphorus, and other nutrients to soil), and displaying carbon sequestration (some feeder root carbon stabilizing in soil organic matter contributing to long-term storage). Senescent feeder root functional replacement includes= requiring continuous production (feeder root senescence necessitating ongoing new root production maintaining functional population), showing production-mortality balance (vine adjusting feeder root production matching mortality maintaining adequate absorption capacity), exhibiting compensation (vine potentially increasing feeder root production compensating for accelerated senescence under stress), demonstrating resource allocation (substantial photosynthate allocated to replacing senescing feeder roots particularly in high-turnover systems), and revealing population homeostasis (relatively stable functional feeder root population maintained despite continuous individual root senescence and production).
Senescent_Feeder_Root emphasizes declining function and programmed death process and is distinguished from active feeder roots by focus on terminal degradation. Includes Aging_Senescent_Feeder_Root, Dying_Senescent_Feeder_Root, and Non_Functional_Senescent_Feeder_Root. Critical for understanding feeder root senescence, cytoplasmic degradation, functional decline, turnover dynamics, carbon cycling, and the programmed terminal phase of ephemeral feeder root life requiring continuous production maintaining functional population.
Absterbende Feinwurzel
Aging feeder root
Alternde Saugwurzel
Declining absorptive root
Dying fine root
Senesze Feinwurzel
Senescent Feeder Root
A senescent lower canopy leaf at node 3 on a shoot of Vitis vinifera cv. Pinot Noir in Burgundy, France, exhibiting pronounced yellowing with 60% of lamina area showing chlorosis and 15% showing orange to red coloration from anthocyanin accumulation particularly along veins and margins, demonstrating severely reduced photosynthesis rate of 3.8 μmol CO₂ m⁻² s⁻¹ representing only 28% of peak rate measured in May, showing visual signs of nutrient remobilization including chlorophyll degradation evident in yellow coloration and reduced tissue nitrogen content measured at 1.2% dry weight compared to 2.4% in healthy mature leaves indicating 50% nitrogen export, experiencing progressive shading to 18% full sunlight from dense overlying canopy creating conditions favoring senescence initiation, showing anatomical degradation including chloroplast breakdown visible as loss of dark green staining in palisade cells and beginning membrane deterioration, exhibiting beginning abscission layer formation at petiole base visible as brown zone development though leaf remains attached, occurring at 14 weeks post-leaf emergence and 10 weeks post-achievement of maturity, measured at shoot BBCH stage 85 (softening of berries) in mid-August, representing typical early senescence of shaded lower canopy basal leaves providing nutrient mobilization supporting fruit ripening and reserve accumulation.
A lower canopy basal leaf undergoing programmed senescence involving progressive degradation of cellular components and nutrient mobilization in Vitis vinifera, characterized by visible color changes from green to yellow, orange, or red reflecting chlorophyll degradation and anthocyanin accumulation, declining photosynthetic capacity with rates reduced to 30-60% of peak function or complete cessation in advanced senescence, progressive nutrient remobilization particularly nitrogen, phosphorus, and potassium exported from degrading leaf tissue to support fruit ripening and reserve accumulation, anatomical changes including chloroplast degradation, membrane breakdown, and cell wall modifications, eventual abscission layer formation at petiole base enabling leaf detachment, and occurrence typically 8-16 weeks after leaf emergence with basal leaves senescing earliest due to age, progressive shading, and natural developmental programming. Senescent lower canopy leaves represent the terminal phase of leaf lifecycle, transitioning from photosynthetic source organs to nutrient mobilization sites providing valuable resource recovery, with senescence timing influenced by multiple factors including leaf age, shading intensity, water stress, nutrient availability, disease pressure, and developmental signals from fruiting structures and vegetative meristems. The senescence process, while representing functional decline, provides critical nutrient cycling returning approximately 40-60% of leaf nitrogen, 50-70% of phosphorus, and 40-50% of potassium to perennial structures supporting subsequent season growth potential and current season reserve accumulation.
Senescent lower canopy leaves represent natural terminal leaf development, distinguished from functional mature leaves by visible degradation symptoms, declining photosynthesis, nutrient remobilization, and approaching abscission. Senescence timing shows strong positional patterns with basal leaves senescing earliest, median leaves later, and apical leaves potentially maintaining function into autumn depending on environmental conditions and cultivar. Multiple triggers initiate senescence including age-dependent developmental programming, severe shading reducing photosynthesis below maintenance thresholds, water stress, nutrient deficiency, disease pressure, and signals from fruit ripening including hormonal changes and resource competition. Senescence represents controlled degradation recovering valuable nutrients rather than simple decay, with substantial nitrogen, phosphorus, potassium, and micronutrient mobilization from senescing leaves supporting concurrent fruit development and subsequent season growth potential. Premature widespread senescence indicates stress conditions requiring diagnostic attention, while delayed senescence into late autumn may indicate excessive vegetative vigor or inadequate crop load compromising dormancy preparation. Management practices including nitrogen fertilization timing, irrigation management, and crop load adjustment affect senescence patterns and nutrient cycling efficiency.
aging lower canopy leaf
alterndes basales Blatt
degrading lower leaf
seneszentes untere Kanopie Blatt
vergilbendes Weinblatt
yellowing basal leaf
Based on Smart (1985) "Principles of grapevine canopy microclimate manipulation with implications for yield and quality" American Journal of Enology and Viticulture 36=230-239 Poni et al. (2009) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 60=397-407 Gutha et al. (2010) "Modulation of flavonoid biosynthetic pathway genes and anthocyanins due to virus infection in grapevine (Vitis vinifera L.) leaves" BMC Plant Biology 10=187 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 401-424.
Senescent Lower Canopy Leaf
Pinot Noir grafted on Riparia Gloire showing senescent root hairs 45-60 mm from root tip on third-order lateral root, hairs 8-11 days post-initiation in terminal senescence phase, microscopic examination revealing collapsed appearance with loss of turgor, brown coloration from phenolic accumulation, cytoplasmic degradation with organelle breakdown visible, FDA viability staining showing weak or absent fluorescence indicating membrane dysfunction, senescent hairs showing minimal or zero uptake in isotope tracing experiments, senescing hairs representing normal terminal phase of ephemeral hair lifespan
Senescent root hair is root hair undergoing programmed senescence showing declining function and approaching death, typically 5-10 days post-initiation representing terminal phase of hair lifespan, characterized by progressive cytoplasmic degradation with organelle breakdown, membrane dysfunction reducing selective permeability and transport capacity, declining turgor reflecting osmotic imbalance, loss of absorption function with minimal or zero water and nutrient uptake, and visible morphological changes including collapsed appearance, wall browning, and cytoplasmic disorganization, with senescent root hair showing orderly dismantling of cellular structures through programmed process distinct from necrotic death, often accumulating phenolic compounds or other metabolites creating brown coloration, and demonstrating that senescence represents normal terminal phase of ephemeral root hair lifespan creating continuous turnover requiring constant new hair production at root tips maintaining functional hair population despite individual hair brevity. Senescent root hair morphological changes includes= showing collapsed appearance (loss of turgor causing hair to shrink and wrinkle losing turgid cylindrical form), exhibiting wall browning (phenolic accumulation and oxidation creating brown or tan coloration), demonstrating tip degradation (hair tip often showing most advanced degradation reflecting age or position of initial senescence signals), revealing structural damage (wall potentially developing breaks or irregularities during senescence), and displaying attachment weakening (senescing hair potentially separating from parent cell or root surface). Senescent root hair cytoplasmic degradation includes= showing organelle breakdown (mitochondria, plastids, ER, and other organelles degrading during senescence), exhibiting vacuole expansion (vacuoles potentially enlarging and coalescing as cytoplasm degrades), demonstrating autophagy (cellular self-digestion through autophagy pathways recycling cellular components), revealing protein degradation (proteases breaking down cellular proteins during senescence), and displaying cytoplasmic collapse (progressive loss of cytoplasmic organization and structure). Senescent root hair membrane dysfunction includes= showing loss of selective permeability (membrane damage allowing unregulated solute leakage reducing compartmentation), exhibiting reduced transporter function (membrane proteins degrading reducing active ion uptake capacity), demonstrating increased permeability (membranes becoming leaky allowing cellular contents to escape), revealing depolarization (loss of membrane potential as ion pumps fail), and displaying lipid peroxidation (oxidative damage to membrane lipids compromising structure and function). Senescent root hair functional decline includes= showing reduced absorption (water and nutrient uptake declining as membrane function deteriorates), exhibiting decreased respiration (mitochondrial breakdown reducing ATP production and oxygen consumption), demonstrating loss of turgor (osmotic imbalance and membrane damage causing turgor loss and collapse), revealing cessation of cytoplasmic streaming (active circulation ceasing as cytoskeleton and energy supply fail), and displaying inability to respond (senescing hairs losing capacity to respond to environmental signals or plant demands). Senescent root hair biochemical changes includes= showing increased protease activity (protein degradation enzymes activated during senescence), exhibiting nuclease activation (DNA and RNA degradation as senescence progresses), demonstrating oxidative stress (reactive oxygen species accumulating potentially driving senescence progression), revealing phenolic accumulation (synthesis or release of phenolic compounds creating brown pigmentation), and displaying metabolite remobilization (some cellular nutrients potentially recovered and transported to plant before complete death). Senescent root hair timing and duration includes= showing variable onset (senescence typically beginning 5-8 days after hair initiation though variable with conditions), exhibiting progression (senescence progressing over 1-3 days from onset to complete loss of function), demonstrating position correlation (senescence occurring at progressively greater distances from root tip as root grows), revealing environmental effects (stress potentially accelerating senescence while favorable conditions extending functional period), and displaying inevitable progression (senescence irreversible once initiated proceeding to hair death). Senescent root hair population dynamics includes= showing continuous occurrence (senescence continuously occurring in older hair zone while new hairs produced at tip), exhibiting wave pattern (senescence zone moving basipetally along root as root elongates and ages), demonstrating balance with production (senescence rate balanced by production rate maintaining relatively stable functional hair number), revealing turnover contribution (senescing and dying hairs contributing to root organic matter input to soil), and displaying management effects (practices accelerating senescence or production affecting functional hair population size). Senescent root hair fate includes= showing eventual death (senescence culminating in complete hair death with loss of all cellular function), exhibiting cell wall persistence (dead hair walls potentially persisting briefly on root surface before sloughing or decomposition), demonstrating organic matter contribution (dead hairs decomposing contributing to rhizosphere organic matter and soil carbon), revealing microbial colonization (dead and senescing hairs potentially colonized by soil microbes accelerating decomposition), and displaying replacement (dead hairs continuously replaced by new hairs at root tip maintaining functional population).
Senescent_Root_Hair emphasizes declining function and programmed death process and is distinguished from functional active hairs by focus on terminal degradation. Includes Dying_Senescent_Root_Hair and Non_Functional_Senescent_Root_Hair. Critical for understanding hair lifespan, cytoplasmic degradation, membrane dysfunction, functional decline, turnover dynamics, and the programmed senescence process representing normal terminal phase of ephemeral root hair life requiring continuous replacement.
Absterbende Wurzelhaare
Aging root hair
Alternde Haare
Declining hair
Dying root hair
Seneszierende Wurzelhaare
Senescent Root Hair
Sensory description for Grüner Veltliner Smaragd from the Wachau (2024 vintage) integrates visual assessment (brilliant pale gold with green-yellow reflections, star-bright clarity, medium viscosity with fine legs), olfactory profile (intense nose of white pepper, ripe citrus, stone fruit, and subtle floral lift with a mineral-flinty undertone), and palate evaluation (dry, medium-full body, concentrated citrus-stone fruit core, pronounced spicy-peppery mid-palate, firm acidity providing structural backbone, long saline-mineral finish of 15+ seconds), scored 93 points on the 100-point scale by the Austrian Falstaff panel. Sensory description for Pinot Noir from Gevrey-Chambertin Premier Cru (2023 vintage) documents a pale ruby-garnet color with brilliant clarity, an expressive bouquet of red cherry, raspberry, dried rose petals, and subtle forest floor with emerging spice from 16 months in 30% new oak, and a palate of medium-full body with silky tannins, vibrant acidity, and a persistent finish combining fruit purity with emerging tertiary complexity.
Sensory description is a wine specification entity that documents the organoleptic characterization of wine as perceived through human sensory evaluation, encompassing visual assessment (appearance), olfactory assessment (aroma and bouquet), gustatory assessment (palate), and tactile assessment (mouthfeel and texture), recorded through standardized tasting protocols using defined descriptor vocabularies and scoring systems, and providing the subjective but systematically structured quality evaluation that complements analytical chemistry data by capturing the integrated perceptual experience that ultimately determines wine quality judgment, consumer preference, and commercial value.
Sensory description represents the irreducible subjective dimension of wine quality assessment that no analytical measurement can fully substitute, as the human perceptual system integrates chemical stimuli through neurological processes that produce emergent quality judgments not predictable from the sum of individual chemical parameters. The formalization of wine sensory evaluation has progressed substantially since the pioneering work of Maynard Amerine and Ann Noble at UC Davis in the 1970s-1980s, with the development of standardized tasting protocols (ISO 3591 wine tasting glass, ISO 8589 tasting room conditions), descriptor vocabularies (the Wine Aroma Wheel, the WSET Systematic Approach to Tasting), and statistical methods for panel training and data analysis (descriptive analysis, napping, CATA/check-all-that-apply). The sensory description hierarchy in VVO captures the three principal assessment modalities — visual (appearance, the first impression), olfactory-gustatory (aroma and palate, the core quality evaluation), and tactile (mouthfeel, increasingly recognized as quality-determinant) — each with defined sub-parameters documented in the child classes. The relationship between sensory description and the analytical data documented elsewhere in the VVO Information Entity hierarchy is complementary but not redundant — analytical chemistry quantifies what is present, while sensory description captures what is perceived, and the two domains diverge whenever perceptual interactions (masking, enhancement, synergy) modify the sensory impact of chemical compounds. The cultural and individual variability in wine sensory evaluation — influenced by tasting experience, sensory acuity genetics (supertasters versus non-tasters), and cultural flavor reference frameworks — necessitates panel-based evaluation with statistical treatment for reliable sensory characterization.
Sensorikdaten
Verkostungsbeschreibung
organoleptic description
organoleptische Beschreibung
sensorische Beschreibung
sensorisches Profil
sensory evaluation
sensory profile
tasting description
wine tasting notes
Noble AC et al. (1987) Modification of a standardized system of wine aroma terminology. American Journal of Enology and Viticulture 38(2): 143-146 Jackson RS (2020) Wine Science: Principles and Applications, 5th Ed, Academic Press, pp.: 461-545 Lawless HT, Heymann H (2010) Sensory Evaluation of Food: Principles and Practices, 2nd Ed, Springer, pp.: 1-30
Sensory Description
Sensory quality evaluation of Sauvignon Blanc from Marlborough employs a trained panel assessing 15 aroma descriptors (passionfruit, grapefruit, boxwood, grass, capsicum), 5 taste attributes (sweetness, acidity, bitterness), and 3 mouthfeel characteristics (body, astringency, effervescence), with the resulting sensory profile compared to regional typicity benchmarks to guide harvest timing and winemaking decisions targeting the aromatic intensity and crisp acidity that define this origin's market identity.
Sensory quality is a quality that inheres in grape berries and derived products and represents the ensemble of attributes perceptible through human sensory evaluation, encompassing taste, aroma, texture, and visual characteristics that collectively determine consumer acceptance, market value, and fitness for intended use. Within the BFO framework, this quality serves as the integrative category bridging objective physicochemical properties of grape tissue with subjective human perception, recognizing that sensory qualities emerge from the interaction between chemical stimuli (sugars, acids, phenolics, volatiles) and the human sensory apparatus with its inherent variability and context dependence. The ontological significance lies in sensory quality representing the ultimate criterion for grape and wine quality assessment, where instrumental measurements serve as proxies for sensory attributes but cannot fully substitute for human evaluation that integrates multiple stimuli, detects interactions, and applies hedonic judgment. Sensory quality assessment in viticulture and enology employs trained panels using standardized descriptive analysis protocols to quantify individual attributes, consumer panels to assess overall acceptability and preference, and expert evaluation applying holistic quality judgment based on experience and regional typicity expectations. The multidimensional nature of sensory quality requires decomposition into component modalities (taste, smell, touch, vision) while recognizing that the perceived quality emerges from cross-modal interactions where sweetness perception is modulated by aroma, astringency perception is affected by viscosity, and color influences flavor expectations. The developmental trajectory of sensory quality during grape ripening reflects the coordinated accumulation of sugars, metabolism of acids, synthesis of phenolics, and generation of aroma precursors, with optimal harvest representing the balance point where the sensory profile matches the target wine style. Understanding sensory quality enables optimization of viticultural and winemaking practices to achieve desired sensory outcomes, prediction of consumer response, and communication of quality attributes throughout the value chain.
Sensory_Quality serves as the root class for the sensory quality hierarchy, establishing the framework for human-perceptible attributes distinct from purely instrumental measurements. This class encompasses qualities evaluated through taste (gustation), smell (olfaction), touch (somesthesis), and vision, with child classes addressing each modality. The inherent subjectivity of sensory evaluation requires standardized protocols and trained assessors for reliable quality characterization.
Sinnesqualität
Wahrnehmungsqualität
eating quality
organoleptic quality
organoleptische Qualität
perceptual quality
sensorial attribute
sensorische Qualität
Lawless HT, Heymann H (2010) Sensory Evaluation of Food: Principles and Practices, 2nd Ed. Springer, New York, pp. 1-18 Jackson RS (2014) Wine Science: Principles and Applications, 4th Ed. Academic Press, San Diego, pp. 831-887
Sensory Quality
Grenache separating abscission zone at full maturity (BBCH 89, 25°Brix) exhibits 80-95% middle lamella dissolution in separation layer, rounded cells with 15-25 μm intercellular gaps, severed vascular bundles with exposed 20-35 μm xylem vessels, berry detachment force 0.4-0.7 N enabling gentle hand harvest with clean separation, and protective layer cells showing suberization and phenolic accumulation preventing pathogen entry post-separation
A separating abscission zone is an abscission zone undergoing active physical cell separation with extensive middle lamella dissolution, cell wall degradation, and progressive tissue disruption, representing the final stage of abscission immediately preceding or during berry detachment from the pedicel. Separating abscission zone is characterized by= advanced cell wall hydrolysis (middle lamella largely or completely dissolved, primary walls extensively degraded, cell-cell adhesion minimal or absent), separation layer cells rounding up and losing contact (cells acquiring spherical shape as wall tension released, intercellular spaces expanding as cells separate), vascular bundle disruption (xylem and phloem tissues severed in separation plane with individual vessel and sieve tube elements exposed), protective layer differentiation (cells adjacent to separation layer completing suberization or lignification creating protective barrier), and minimal berry attachment strength (detachment force 0.3-0.8 N, approaching or at threshold for spontaneous berry drop). Cellular and structural features of separating abscission zone include= separation layer cells with rounded morphology and smooth surfaces (cell wall degradation removes middle lamella and modifies primary walls), exposed cell walls showing characteristic staining patterns (reduced pectin content, modified cellulose organization), fractured vascular bundles with open xylem vessels and severed phloem sieve tubes, intercellular spaces filled with cell wall degradation products (oligogalacturonides, cellulose fragments), and beginning of wound responses in protective layer cells (callose deposition, phenolic accumulation). Separating abscission zone timing varies by cultivar and conditions= in early-abscising cultivars separation may begin at BBCH 87-88 (early harvest maturity, some berries may drop before optimal harvest), in typical wine grapes separation occurs at BBCH 89 (full maturity, facilitating harvest but minimal premature drop), in late-abscising or firm-attachment cultivars separation may require BBCH 89+ (overripeness) or may be incomplete even at advanced maturity. Mechanical properties of separating abscission zone show characteristic changes= detachment force declining to 0.3-0.8 N (compared to 0.8-1.5 N in active abscission zones and 1.5-2.5 N in inactive zones), failure mode shifting from cohesive tissue fracture to clean separation at abscission plane, reduced energy absorption during detachment (brittle rather than ductile failure), and minimal tissue damage or tearing during separation. Separating abscission zone can be induced prematurely by= severe environmental stress (extreme drought, heat stress, freezing events triggering stress-induced abscission), pathogen infection (Botrytis cinerea or other pathogens inducing defense-related abscission), excessive ethylene exposure (exogenous ethylene application accelerating activation and separation), physical damage (mechanical injury to cluster or pedicel inducing wound-response abscission), and senescence (overripe berries with advanced abscission zone development leading to spontaneous berry drop). Post-separation characteristics include= clean separation surface at abscission plane (minimal tearing or tissue damage), exposed vascular bundles visible in remaining pedicel stub, protective layer preventing pathogen entry and water loss, and potential callose or suberin deposition at separation surface.
Separating_Abscission_Zone emphasizes active physical cell separation and is distinguished from Active_Abscission_Zone (enzymatic activation preceding separation) and Pedicel_Abscission_Zone (complete structure) by focus on advanced cell wall degradation and imminent berry detachment. Critical for understanding optimal harvest timing for gentle berry separation, premature fruit drop mechanisms under stress or disease, harvest quality with minimal tissue damage, and cultivar differences in abscission progression affecting harvest characteristics.
Ablösende Abszissionszone
Actively separating zone
Aktiv separierende Schicht
Detaching abscission layer
Sich trennende Zone
Patterson, S. E. (2001). Cutting loose. Abscission and dehiscence in Arabidopsis. Plant Physiology, 126(2), 494-500. | Considine, J. A., & Knox, R. B. (1979). Development and histochemistry of the cell, tissue, and secretion within the dermal system of the grape. Protoplasma, 99(3-4), 347-365.
Separating Abscission Zone
Syrah inflorescences at BBCH 60 (beginning of flowering, 5-10% of flowers blooming) exhibit separating calyptras visible as partially lifted yellowish-green caps with stamens emerging beneath, showing progressive basal separation creating 1-2 mm gaps around circumference, upward displacement of 1-2.5 mm from initial position, and tilted or canted orientation in many flowers as asymmetric detachment proceeds, with individual flowers completing separation over 2-4 hours under favorable conditions (morning temperatures 22-26°C, humidity 60-70%). Riesling during active bloom shows separating calyptras at various stages of displacement from initial basal gap formation through nearly complete detachment with only narrow connection remaining, creating a dynamic visual appearance of emerging stamens and pistils across 10-15% of flowers simultaneously in separating state. Cabernet Sauvignon experiencing rain during bloom (BBCH 60-61) shows disrupted calyptra separation with some flowers exhibiting prolonged separation (>6 hours) as water accumulation interferes with normal shedding, while other flowers show premature mechanical dislodgement of partially separated calyptras by raindrop impact, both scenarios resulting in suboptimal bloom conditions and reduced fruit set (38-44% versus normal 48-55%).
A separating calyptra is a calyptra in active transition from intact pre-dehiscence state to complete detachment, characterized by progressive basal separation at the abscission zone through cell wall dissolution and mechanical lifting by elongating stamens, representing the dynamic anthesis process typically occurring over 1-6 hours per individual flower and defining the onset of bloom (BBCH 60-61) when separating calyptras become visible within inflorescences. Separating calyptras are characterized by= (1) partial separation at the basal abscission zone with visible gaps or splits developing around part or all of the circumference as cell wall-degrading enzymes complete dissolution of middle lamellae connecting calyptra base to receptacle (2) progressive upward displacement of the calyptra as elongating stamen filaments push the calyptra apex upward, with displacement ranging from 0.5-3 mm depending on stage of separation process (3) calyptra remaining coherent as a single fused petal unit during the separation process, typically shedding as an intact cap rather than splitting into individual petals (4) visible exposure of stamen filaments and/or anthers as the calyptra lifts, creating a transition state between fully enclosed and fully exposed reproductive organs (5) in some flowers, calyptra tilting or canting to one side during separation as detachment proceeds asymmetrically around the circumference, creating angled rather than vertical displacement before final release. The separating calyptra state represents the active anthesis process and encompasses BBCH stage 60 (first calyptras detaching, beginning of flowering) through early BBCH 61 (10% of calyptras shed, beginning of full flowering), with the percentage of flowers showing separating calyptras at any given time typically 85%) can interfere with calyptra shedding by causing calyptra tissue swelling or by promoting fungal growth at abscission zone that disrupts normal separation rain during separation can physically dislodge partially separated calyptras prematurely or conversely can cause water accumulation delaying complete separation. Separating calyptras are critical indicators of active bloom and represent the optimal timing for certain viticultural operations= bloom-timing fungicide applications targeting Botrytis infection of newly exposed floral tissues are most effective when 5-20% of flowers show separating calyptras (BBCH 60-61) controlled pollination operations in breeding programs or pollinizer management for functionally female cultivars should target the period when separating calyptras indicate stigmas are becoming exposed and approaching peak receptivity.
Separating_Calyptra represents the active anthesis process and is distinguished from Intact_Calyptra (pre-dehiscence but not yet separating) by presence of basal gaps and upward displacement, and from Detached_Calyptra (separation complete) by retention of partial attachment. The separating state is transient for individual flowers (typically 1-6 hours duration) but represents a critical phenological stage for inflorescence-level observations as the onset and peak of separating calyptras define the beginning of flowering period (BBCH 60-61). Sub-classification into Partially_Separated and Loosening types reflects the progression within the separation process. Monitoring separating calyptras provides real-time assessment of bloom progression and informs immediate management decisions regarding disease control, irrigation, and canopy management during the critical bloom period.
abtrennende Kelchhaube
detaching petal cap
lifting calyptra
sich ablösende Blütenhaube
sich öffnende Kronblattkappe
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 100-103 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 94-96 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 193-195.
Separating Calyptra
Cabernet Franc ovaries at BBCH 59-61 exhibit characteristic septate bilocular organization with central septum 0.09-0.12 mm thick extending from ovary base to apex, dividing ovary (0.9-1.1 mm diameter) into two locules of approximately equal size (left and right locules each 0.4-0.5 mm diameter), with septum composed of thin-walled parenchyma cells and containing 3-5 vascular bundles supplying ovules (4 total, 2 per locule) attached to septum surfaces. Syrah ovary longitudinal sections reveal septum running the full length of the ovary (0.8-1 mm), positioned along the median plane in symmetric pistils or displaced 0.1-0.2 mm off-center in asymmetric pistils, with ovules attached to septal surfaces at positions approximately midway along septum length, and with septal vascular bundles visible as dark-staining tissues running through the parenchymatous septum. Chardonnay berry development from BBCH 71 (fruit set, berry 1.5-2 mm diameter with clear septum dividing two locules) through BBCH 81 (berries beginning to ripen, 8-10 mm diameter with septum still visible but less prominent) to BBCH 89 (berries ripe, 11-14 mm diameter with septum often degraded or obscured by proliferated flesh though seed positions still reflect original two-locule organization) demonstrates progressive septum fate during fruit maturation. Pinot Noir occasional flowers show septum fenestrations creating small gaps (0.05-0.15 mm diameter) in the septum allowing limited communication between the two locules, though the septum remains largely intact providing substantial locule separation and independent ovule/seed development capacity.
A septate bilocular ovary is a bilocular ovary specifically characterized by the presence of a central septum (partition wall) dividing the ovary into two locules, emphasizing the septum as the critical structural element creating and maintaining the two-chamber organization characteristic of Vitis ovaries. Septate bilocular ovaries are characterized by= (1) a central septum extending from ovary base to apex (or nearly to apex in some cases) dividing the ovary cavity into two distinct locules (2) septum composed primarily of thin-walled parenchyma cells derived from the fused medial margins of the two constituent carpels (3) septum thickness typically 0.06-0.15 mm, varying with overall ovary size and developmental stage (4) vascular bundles running through the septum tissue providing nutrient supply to ovules attached to septum surfaces (5) septum positioned along the median plane of the ovary in symmetric bicarpellate pistils or positioned asymmetrically (displaced toward the smaller locule) in asymmetric bicarpellate pistils (6) septum maintaining structural integrity throughout ovary development and typically into early fruit development, though potentially becoming less distinct or partially degrading during later berry maturation as flesh tissue proliferates. The septum develops during early pistil organogenesis (BBCH 52-54) through fusion of the two carpel primordia, with the carpel medial (inner) margins meeting and fusing to create the partition wall, effectively converting two originally separate carpel cavities into a unified compound ovary containing two septum-separated locules. Septum structure and composition reflect its origin from fused carpel margins= the septum tissue consists of parenchyma cells continuous with the carpel wall tissues, with cellular organization showing symmetry around the septum midline reflecting the bilateral carpel fusion vascular tissues in the septum originate from the carpel margin vascular strands, with bundles running along the septum providing nutrient supply to placental tissues and attached ovules the septum surfaces facing into the two locules bear the placentae (ovule attachment tissues) at positions corresponding to the original carpel margin positions where ovules develop. Septum integrity varies somewhat among flowers and developmental stages= most septa are complete solid partitions fully separating the two locules throughout their length some septa show small fenestrations (gaps) creating limited communication between locules rare septa are incomplete, extending only partially through the ovary creating partial rather than complete locule separation. The septum plays critical functional roles= structural support dividing the ovary into two chambers and providing attachment sites for ovules vascular conduit supplying nutrients and water to developing ovules via septal vascular bundles potential developmental coordinator allowing independent regulation of the two locule environments. Post-fertilization berry development involves changing septum characteristics= early berry stages (BBCH 71-75) maintain clear septum structure dividing the young berry into two locules mid-development stages (BBCH 77-83) may show septum persistence but with increasing flesh proliferation reducing septum prominence late development and maturity (BBCH 85-89) often show septum degradation or obscuring as flesh tissue fills former locule spaces, though seed positioning typically retains evidence of original septum-defined locule organization.
Septate_Bilocular_Ovary emphasizes the septum as the critical structural element creating bilocular organization and is complementary to Two_Chamber_Bilocular_Ovary (which emphasizes the resulting chamber number). The septate condition is universal in normal Vitis ovaries as the septum is the defining feature creating the two locules from the two fused carpels. Understanding septum structure, composition, and developmental fate is important for interpreting ovule placentation, vascular supply to ovules, locule independence, and berry internal structure development. Septum variations (complete versus fenestrated, symmetric versus asymmetric positioning) provide information about carpel fusion patterns and pistil developmental dynamics.
Scheidewand-Ovar
Trennwand-Fruchtknoten
partitioned bilocular ovary
septierter Fruchtknoten
septum-bearing ovary
Septate Bilocular Ovary
Cabernet Franc in Loire Valley shows sequential cap detachment during mid-late June Year N+1 at BBCH 67-68 over 3 days June 13-15, with 18-26% flowers completing individual asynchronous cap fall predominantly in basal positions at rate of 5-8% per day, creating extended shedding tail, under temperate maritime conditions of 20-25°C temperature, 62-76% relative humidity, typical of this mid-season cultivar showing moderate developmental gradient contributing 14-20% berry number through sequential shedding with 22-34% late pollination success
Sequential cap detachment represents the individual progressive calyptra shedding pattern during progressive cap fall when remaining flowers complete abscission sequentially as they individually reach maturity rather than through synchronized mass events, reflecting developmental gradient and creating extended flowering tail. This characteristic late shedding pattern occurs during late flowering in mid-late June in temperate regions Year N+1 as part of progressive cap fall, spanning 2-3 days as final flowers individually mature and shed caps creating non-synchronized extended detachment, corresponding to BBCH 67-68 stages. Detachment sequence shows individual flower maturation pattern with flowers completing cap fall one-by-one or in small groups 1-3 flowers rather than mass synchronized events, temporal spread reflecting developmental heterogeneity established during previous season primordium formation and current season progression, and overall sequential pattern creating extended shedding period. Spatial sequence typically proceeds basipetal direction with remaining unopened flowers concentrated in basal cluster positions, progressive detachment advancing from middle toward basal regions following developmental gradient, and occasional scattered late detachment in other positions from locally delayed flowers. Individual flower timing shows asynchronous maturation with individual flowers reaching abscission readiness at different times spanning days, cap detachment occurring independently as each flower completes maturation rather than coordinated by common signals, and variable timing creating extended low-intensity shedding period. Developmental gradient expression through sequential pattern clearly reveals maturation sequence established during previous season inflorescence primordium formation, with earliest apical primordia producing earliest-maturing flowers completing cap fall first during initial phase, and latest basal primordia producing latest-maturing flowers completing sequential detachment during progressive phase. Detachment rate during sequential shedding shows low continuous activity with daily additions of 3-7% flowers completing cap fall, sustained over several days creating cumulative substantial completion, and overall pattern contrasting with earlier compressed intense mass fall. Cap accumulation pattern during sequential shedding shows modest ongoing additions with individual or small groups of fresh caps appearing daily, accumulation less visually striking than mass fall intensive deposition, and overall continued cap debris accumulation documenting extended flowering period. Flower exposure during sequential shedding shows progressive individual organ revelation as each flower completes cap detachment, spatial distribution reflecting sequence pattern with basal concentration, and overall extended exposure period affecting late reproductive potential. Reproductive opportunity during sequential detachment remains limited with each newly opened flower encountering suboptimal conditions including minimal viable pollen availability from aged earlier production, reduced atmospheric pollen density, limited concurrent receptive stigmas for cross-pollination, and overall low probability of successful pollination affecting fruit set from sequential shedding. Pollination success during sequential phase shows reduced efficiency with estimated 15-35% late-opening flowers achieving successful pollination and fruit set compared to 50-70% during optimal mass fall, resulting primarily from pollen limitation rather than intrinsic flower dysfunction, and affecting final fruit set contribution from sequential shedding period. Environmental effects on sequential shedding include temperature affecting individual flower maturation rate and cap detachment efficiency, humidity influencing mechanical separation in each detaching flower, and overall conditions less critical since reproductive outcomes largely determined during earlier phases. Humidity sensitivity during sequential shedding shows continued importance for efficient individual detachment with moderate levels facilitating normal cap fall in late flowers, while elevated humidity potentially causing retention in susceptible individuals, though overall impact limited since majority shedding already completed. Temperature during sequential shedding affects progression rate with warm conditions 24-28°C promoting rapid individual flower maturation completing sequential phase within 2-3 days, while cool conditions 18-22°C slowing progression potentially extending period to 4-5 days. Developmental asynchrony underlying sequential pattern reflects variable flower maturation resulting from heterogeneous previous season primordium formation timing, differential current season environmental exposure affecting progression rates, and inherent developmental variation creating spread in flowering readiness. Extended flowering implications of sequential shedding include prolonged flowering period from 10-14 days total compared to 7-10 days with compressed synchronous flowering, increased within-cluster developmental heterogeneity affecting ripening uniformity, and potential quality implications from variable berry development timing. Cultivar differences in sequential pattern show some varieties exhibiting pronounced sequential shedding with 25-35% flowers in extended tail, while others show compressed pattern with only 10-15% sequential component, affecting flowering duration and uniformity characteristics. Management implications of sequential shedding include recognizing extended flowering period requiring sustained weather monitoring through completion, understanding that sequential pattern indicates developmental heterogeneity potentially affecting harvest uniformity, and appreciating that extended flowering creates prolonged vulnerability period though later flowers contribute minor proportion to final yield. Observational characteristics enable recognition of sequential shedding through low-intensity sustained cap fall activity over several days, individual or small-group detachment events rather than mass synchronized fall, spatial progression following developmental gradient, and overall extended gradual flowering completion. The process represents first aspect of progressive cap fall emphasizing sequential individual pattern, followed by extended shedding discussing temporal implications. Duration spans 2-3 days representing sequential progression period with individual flowers completing detachment asynchronously. Cultivar variation in sequential characteristics shows differences in sequence extent affecting proportion of flowers in sequential tail, sequence rate affecting duration of sequential period, and retention susceptibility in late sequential flowers affecting complete flowering achievement. Regional timing shows sequential cap detachment during mid-late June in warm climates, late June in temperate regions, and late June-early July in cool areas, occurring during late flowering as part of progressive cap fall. Viticultural significance includes revealing developmental heterogeneity through extended sequential pattern affecting flowering and ripening uniformity, contributing final proportion 10-20% berry number through extended shedding though with reduced success rate, and indicating completion approach through declining sequence activity enabling management transitions. Research applications include analyzing developmental gradients producing sequential shedding patterns, evaluating environmental and genetic factors affecting shedding sequence characteristics, quantifying relationships between sequential shedding extent and ripening uniformity, assessing pollination limitations during sequential period affecting late fruit set, and documenting flowering duration variation affecting management requirements across cultivars and conditions.
Sequential_Cap_Detachment is first aspect of Progressive_Cap_Fall, distinguished by individual asynchronous shedding pattern. Reflects developmental gradient from previous season primordium formation. Contrasts with earlier synchronized mass fall. Creates extended low-intensity shedding period. Limited reproductive success due to pollen limitation. Reveals developmental heterogeneity affecting uniformity. Followed by Extended_Cap_Shedding discussing temporal implications.
asynchroner Kappenabwurf
individual cap detachment
individuelle Kalyptratrennung
non-synchronized cap fall
progressive Einzelabtrennung
progressive sequential shedding
Sequential Cap Detachment
Cabernet Sauvignon in Napa Valley shows sequential internode formation from early May through late July at BBCH 11-38 over 12-14 weeks, initiating new internodes every 2-3 days during peak June growth and every 3-5 days during July decline, producing 16-20 total internodes creating final shoot length of 2200-2800 mm under warm climate conditions of 18-30°C temperature range, adequate irrigation, and moderate-high fertility typical of this vigorous late-season cultivar
Sequential internode formation represents the continuous process of producing successive stem segments at shoot apex, establishing progressive shoot structure through coordinated initiation, cell division, and early development of multiple internodes creating developmental gradient along axis from apical meristematic regions to basal maturing zones. This fundamental developmental process occurs throughout active shoot growth from early May through July-August in temperate regions corresponding to BBCH 11-38, with new internodes initiating at shoot apex every 2-3 days during peak growth producing total of 12-20 internodes per shoot over 8-16 week growing period. Sequential formation proceeds through continuous apical meristem activity producing new leaf primordia in regular phyllotactic pattern, with each successive leaf primordium emergence defining nodal position and simultaneously establishing boundaries for new internode segment between successive nodes. Intercalary meristem initiation occurs in axis tissue immediately behind each developing node, with meristematic cells establishing in zone that will form internode through activation of cell division capacity and organization into longitudinal files characteristic of intercalary meristem structure. Cell division in each successive intercalary meristem produces cells contributing to internode length, with division phase typically lasting 10-20 days adding cells through organized divisions creating longitudinal cell files, then transitioning to expansion phase where cell production ceases but cells undergo dramatic elongation. The shoot apex maintains production of new internodes through sustained apical meristem activity, with meristem structure showing organized tunica-corpus arrangement producing organs in regular pattern, initiating new leaf primordia and associated internode segments every 2-3 days during active growth declining to 3-5 day intervals as growth slows. Sequential formation creates characteristic developmental gradient along shoot with basal internodes in maturation phase undergoing lignification, middle internodes in elongation phase actively expanding, and apical internodes in formation phase with active intercalary meristems producing cells. Internode number produced per shoot varies substantially with cultivar vigor and growing conditions, ranging from 10-14 internodes in less vigorous cultivars or limited conditions to 16-22 internodes in vigorous cultivars with optimal conditions and extended growing period. Timing between successive internode initiations reflects shoot growth rate and apical meristem activity level, with most rapid initiation of 2-day intervals during peak exponential growth phase, extending to 3-4 day intervals during declining growth, and ceasing when apical meristem activity terminates with growth cessation. Vascular differentiation proceeds acropetally through sequentially forming internodes, with procambial strands differentiating into functional xylem and phloem in coordinated pattern connecting each new segment into continuous vascular system extending from permanent structures through developing shoot to apex. The sequential formation process shows coordination with other developmental processes including leaf emergence with each internode associated with specific leaf at proximal node, tendril or inflorescence development at nodes following cultivar-specific patterns, and overall shoot elongation resulting from summed contributions of sequentially forming and elongating internodes. Internode length variation among sequentially formed segments reflects changing resource availability and growth conditions during formation and elongation phases, with early internodes typically shorter from limited early resources, middle internodes longest corresponding to peak growth conditions, and late internodes shorter as growth declines and resources become limiting. Cell number produced in intercalary meristems varies among sequential internodes correlating with final length, with shorter internodes containing fewer cells from reduced division duration or rate while longer internodes contain more cells from extended or accelerated division phase. Metabolic activity in sequentially forming internodes shows high levels in active meristems and early elongation zones with respiration supporting cell production and expansion, declining as internodes mature and transition from growth to maintenance metabolism. Hormonal regulation coordinating sequential formation involves auxin from apical meristem maintaining overall shoot growth and coordinating vascular differentiation pattern, gibberellins affecting internode elongation with effects varying among sequential positions, and cytokinins promoting cell division in intercalary meristems with availability potentially varying through season. Temperature affects sequential formation rate through effects on apical meristem activity determining initiation frequency and on cell division rates in intercalary meristems affecting cell production, with optimal 22-28°C supporting maximum formation rate declining at lower or higher temperatures. Light intensity and quality affect formation through effects on photosynthetic production providing resources for growth and through photomorphogenic responses potentially affecting internode length characteristics, with shade promoting elongated internodes while high light supports stouter structure. Water availability affects sequential formation through effects on turgor maintaining meristem activity and cell expansion, with water stress potentially reducing formation rate through slowed apical meristem activity and limiting internode elongation affecting final dimensions. Nutrient availability particularly nitrogen affects formation rate and internode development through effects on protein synthesis supporting cell division and growth, with adequate nutrition supporting maximum formation rate and normal development while deficiency reduces rate and limits development. The process continues throughout shoot growth period showing characteristic temporal pattern of increasing formation rate during early accelerating growth, maximum rate during peak growth, and declining rate during decelerating growth before cessation. Cultivar variation appears in total internode number produced per season reflecting inherent differences in growth duration and vigor, spatial pattern of internode lengths along shoot affecting architecture, and timing of formation cessation relative to fruit development stages. Regional timing shows sequential formation active from May through July-August in warm climates, May through August-September in temperate regions, and June through September in cool areas, with duration corresponding to total shoot growth period. Viticultural management considerations include recognizing that internode formation pattern reflects growth progression providing diagnostic information about vine status and growing conditions, understanding that management affecting shoot growth including irrigation and nutrition influences formation patterns determining final canopy architecture, and timing canopy management operations considering developmental state of forming internodes. Research applications include analyzing mechanisms coordinating sequential organ formation at shoot apex, evaluating temporal patterns of formation and their regulation by internal and environmental factors, quantifying developmental gradients along shoot axis documenting simultaneous processes at different stages, and assessing genetic variation in formation patterns affecting shoot architecture and growth characteristics.
Sequential_Internode_Formation is second of four Internode_Development subphases, distinguished by continuous production of successive internodes throughout growth period. Encompasses entire formation sequence from first through final internode. Transitions focus to Internode_Elongation and Internode_Maturation as individual segments progress through development. Represents fundamental axis segmentation process determining final shoot structure.
continuous internode formation
fortlaufende Internodienbildung
kontinuierliche Segmentbildung
progressive Internodienentwicklung
progressive internode development
successive stem segment formation
sukzessive Stängelgliedentwicklung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 215-230 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 138-150 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Sequential Internode Formation
Tempranillo in Rioja shows sequential leaf emergence from May through August at BBCH 11-38 over 14-15 weeks, with successive leaves separating at 3-4 day intervals during peak June-July growth producing total of 15-18 emerged leaves ranging from 65-90 mm basal dimensions to 95-130 mm mid-shoot maximum declining to 55-80 mm apically, under continental climate conditions of 16-28°C temperature range, moderate summer precipitation, and moderate-high fertility typical of this moderately vigorous mid-season cultivar
Sequential leaf emergence represents the continuous developmental progression where multiple leaf primordia at various developmental stages undergo coordinated growth transitioning from apical initiation through separation from shoot axis to expanding functional leaves, creating characteristic developmental gradient along shoot with mature basal leaves coexisting with emerging apical primordia. This fundamental process occurs throughout shoot growth period from early elongation through growth cessation, typically encompassing 12-20 leaves at various stages from microscopic primordia to fully expanded mature leaves over 8-16 week period. Sequential emergence proceeds through regular timing pattern with successive leaves separating from shoot axis at approximately 3-5 day intervals during active growth, reflecting both primordium initiation frequency at apex and development time required from initiation through separation, creating continuous leaf production throughout growing season. The developmental progression shows characteristic stages advancing along shoot axis from apical meristem producing new primordia, through subapical zones containing developing primordia attached to axis, to regions where primordia separate and begin expanding away from shoot, and finally basal zones with mature expanding to fully developed functional leaves. Leaf separation from shoot axis represents critical transition in emergence process occurring when primordium development reaches stage where petiole angles away from compressed position against axis and blade begins spreading perpendicular to shoot, typically requiring 10-15 days from primordium initiation to separation depending on growth rate and environmental conditions. Each successive leaf shows progressive size increase from small basal leaves emerging during slow early growth to largest middle leaves emerging during peak growth, then declining size in apical leaves emerging during decelerating growth, reflecting changing resource availability and growth conditions during emergence period. The sequential pattern establishes characteristic canopy architecture with leaves positioned in alternating pattern approximately 180 degrees around shoot axis, creating two rows of leaves along shoot when viewed from side providing good light interception while allowing air movement through canopy interior. Developmental timing shows coordination with internode formation with each leaf associated with specific node position, emerging at proximal end of internode that develops between current and previous leaf position, creating integrated shoot structure with consistent organ relationships. Photosynthetic capacity develops progressively in sequentially emerging leaves with each new leaf contributing additional area and carbon fixation capability, with shoot photosynthetic capacity increasing in stepwise manner as each leaf reaches functional status typically 10-15 days after separation. Carbon balance transitions in developing shoot from net consumption during early emergence when few leaves have functional photosynthetic capacity to net production once 3-5 leaves have substantially expanded and matured, with positive balance increasing as sequential emergence adds photosynthetic area through mid-season. Water demand increases progressively with sequential emergence as each additional leaf contributes transpiring surface area, with shoot water use rising from 30-50 mg per hour with 2-3 leaves to 400-700 mg per hour with 12-15 leaves depending on leaf sizes, environmental conditions, and stomatal conductance. Nutrient demand particularly nitrogen intensifies during sequential emergence period as each new leaf requires protein synthesis for structural and photosynthetic components, with nitrogen import from root uptake and possibly mobilization from older leaves supporting continued leaf production through season. Vascular development extends continuously through shoot connecting sequentially emerging leaves into integrated transport system, with xylem providing water delivery from roots to all transpiring leaves and phloem enabling assimilate distribution from photosynthesizing leaves to growth sinks and storage tissues. Leaf position along shoot affects development timing and final characteristics with basal leaves emerging earliest having longer potential lifespan but slower development during early low-resource period, middle leaves emerging during peak growth developing rapidly to large dimensions with maximum photosynthetic capacity, and apical leaves emerging latest showing accelerated development to smaller sizes before growth cessation. Hormonal coordination of sequential emergence involves auxin from shoot apex affecting emergence timing and development rate, cytokinins from roots promoting cell division in developing primordia, gibberellins affecting expansion rate and final leaf size, and possible abscisic acid effects during stress conditions influencing development rate and final characteristics. Temperature affects sequential emergence through effects on development rate with each stage from primordium initiation through separation to expansion approximately doubling per 10°C increase between 15-28°C, determining both timing between successive emergences and development time for individual leaves. Light intensity affects development of sequentially emerging leaves with high light promoting rapid photosynthetic maturation and maximum capacity development while shade reduces photosynthetic development and may alter leaf morphology through shade avoidance responses including increased specific leaf area. Water availability affects sequential emergence rate and leaf development with adequate moisture supporting rapid emergence and expansion creating large well-developed leaves, while water stress slows emergence rate, reduces expansion, and limits final sizes affecting canopy development and productivity. Nutrient availability particularly nitrogen affects emergence rate and leaf characteristics with adequate nutrition supporting maximum emergence frequency and rapid development to large dimensions with high photosynthetic potential, while deficiency reduces emergence rate and limits leaf development affecting canopy capacity. The process continues throughout shoot growth period showing temporal variation in emergence rate corresponding to shoot growth phases, with most rapid sequential emergence during exponential growth phase producing 2-3 leaves per week, declining during decelerating growth to 1-2 leaves per week, and ceasing when apical meristem activity terminates with growth cessation. Cultivar variation appears in total leaf number emerging per season ranging from 10-14 leaves in less vigorous cultivars to 16-22 leaves in highly vigorous types, timing pattern of emergence reflecting growth habit, and final leaf characteristics including size, shape, and photosynthetic capacity affecting canopy productivity. Regional timing shows sequential emergence active throughout shoot growth from May-July in warm climates, May-August in temperate regions, and June-September in cool areas, with emergence pattern reflecting seasonal temperature and resource availability progression. Viticultural management considerations include recognizing that sequential emergence pattern reflects shoot growth progression and environmental conditions providing diagnostic information about vine status, understanding that management affecting shoot growth including irrigation and nutrition influences emergence determining final canopy characteristics, ensuring adequate resources during emergence period supporting full canopy development, and monitoring emergence uniformity across vineyard assessing vine health and resource distribution. Research applications include analyzing regulation of sequential organ emergence including hormonal and developmental controls, quantifying emergence patterns and their environmental responsiveness informing growth models, evaluating canopy development dynamics integrating sequential emergence with shoot growth patterns, and assessing implications of emergence pattern for light interception, photosynthetic capacity, and vineyard productivity.
Sequential_Leaf_Emergence is second of four Leaf_Emergence subphases, distinguished by continuous progression of multiple leaves through developmental stages creating gradient along shoot. Encompasses entire emergence sequence from initiation through separation and early expansion. Transitions focus to Leaf_Primordium_Expansion and Leaf_Maturation as individual leaves progress through development. Represents fundamental canopy development process determining photosynthetic capacity and architecture.
continuous leaf emergence
fortlaufende Blattentwicklung
kontinuierliche Blattentfaltung
progressive Laubentwicklung
successive foliar development
sukzessive Blattbildung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 242-255 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 162-172 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Sequential Leaf Emergence
Serrate margin development in Sangiovese produces the characteristic large teeth with convex sides that define this Tuscan variety's leaf profile, with primary serrations of 4-6 mm height recurring at 10-15 mm intervals and consistent apical orientation of tooth points this pronounced serration pattern, combined with the pentagonal blade outline and medium-deep lateral sinuses, enables confident field identification of Sangiovese in mixed plantings and provides reference characters for authentication of clonal material in nursery propagation.
Serrate_Margin_Quality represents the margin configuration where the leaf edge bears pointed teeth with their tips directed toward the leaf apex, creating a saw-like profile when viewed along the margin. Serrate margins are the predominant condition in Vitis vinifera, with the size, shape, and density of serrations varying among cultivars to provide diagnostic ampelographic characters. True serration is characterized by teeth that are asymmetric, with the apical (toward leaf tip) side of each tooth shorter and steeper than the basal (toward leaf base) side, creating the directional pattern characteristic of a saw blade. The teeth arise from localized cell division and expansion at the leaf primordium margin, with each tooth typically associated with a minor vein terminus that may function as a hydathode. Tooth size in grape leaves ranges from small (less than 2 mm height) through medium (2-4 mm) to large (greater than 4 mm), while tooth density varies from sparse (widely spaced teeth with smooth margin between them) to dense (teeth closely packed along the entire margin). OIV descriptor 077 captures tooth shape, distinguishing teeth with convex sides (broadly triangular), teeth with straight sides, and teeth with concave sides (giving a scalloped appearance). Serrate margin assessment requires examination of fully expanded leaves, as teeth may be less distinct on young, expanding leaves where marginal tissue is still developing.
Serrate_Margin_Quality represents the most common margin type in Vitis vinifera and is characterized by the apical orientation of tooth points. The term may be modified as biserrate (large teeth with smaller teeth on their margins) or doubly serrate when two orders of teeth are present, a condition common in grape leaves where primary teeth associated with major vein termini bear secondary teeth between them. The distinction between serrate (apically directed teeth) and dentate (perpendicular teeth) margins provides taxonomic information though intergrades are common. OIV descriptor 077 categories 1-3 address serrate tooth morphology. PATO_0001807 (serrate) provides the upper-ontology alignment.
Sägezahnrand
apically-directed teeth
gesägte Blattkante
gesägter Rand
saw-like margin
saw-toothed margin
serrated edge
sägezähnige Qualität
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 55-56 Ellis B et al. (2009) Manual of Leaf Architecture, Cornell University Press pp.: 35-42
Serrate Margin Quality
In Shiraz berries in Grampians, Victoria during BBCH 85-89, sesquiterpene production generates rotundone accumulating from below detection to 165 ng/L in berry skins over the final 28 days before harvest, with accumulation strongly enhanced by cool late-season temperatures averaging 16°C compared to warmer sites where rotundone reaches only 45 ng/L, concurrent with total sesquiterpene content of 95 µg/L including farnesol at 18 µg/L and nerolidol at 25 µg/L providing background spicy-woody complexity.
Sesquiterpene production describes the enzymatic biosynthesis of C15 terpenoid compounds in Vitis vinifera berry tissues during ripening, wherein sesquiterpene synthase enzymes convert farnesyl diphosphate precursor into structurally diverse sesquiterpene hydrocarbons and alcohols including farnesol, nerolidol, alpha-farnesene, and notably rotundone that contributes the distinctive black pepper spicy character to Shiraz and related varieties, representing a quantitatively minor but sensorially significant component of grape terpenoid composition. This process involves the ionization of farnesyl diphosphate followed by cyclization and rearrangement reactions generating the complex polycyclic structures characteristic of many sesquiterpenes, with the grapevine genome encoding multiple sesquiterpene synthase genes showing variety-specific expression patterns that determine the sesquiterpene profile contributing to cultivar-distinctive aromatic signatures beyond the dominant monoterpene character. The subcellular organization of sesquiterpene production differs from monoterpene synthesis, with farnesyl diphosphate formed through the cytosolic mevalonate pathway rather than the plastidial MEP pathway, and sesquiterpene synthase activities primarily localized in cytosolic compartments where substrate and enzyme colocalize for efficient product formation. The sensorially most significant sesquiterpene in grape and wine is rotundone, a bicyclic sesquiterpene ketone with extremely low perception threshold of approximately 16 ng/L in red wine providing intense black pepper spicy character, with rotundone synthase expression showing strong cultivar specificity concentrated in Shiraz, Durif, and related varieties while largely absent from non-peppery cultivars. Developmental regulation of sesquiterpene production shows variety-specific timing, with rotundone accumulation in Shiraz occurring primarily during late ripening in the final 3-4 weeks before harvest, reaching concentrations of 50-200 ng/L in berry skins that constitute the primary tissue site of rotundone synthesis and accumulation. Environmental factors profoundly influence sesquiterpene production, with cool temperatures during late ripening substantially enhancing rotundone accumulation in Shiraz to levels 3-5 fold higher than equivalent fruit ripened under warm conditions, explaining the strong site and vintage variation in peppery character expression observed in commercial wines. The tissue localization of sesquiterpene production concentrates in exocarp cells where sesquiterpene synthase expression shows highest levels, with sesquiterpene concentrations in skin tissue typically 10-50 fold higher than in mesocarp, suggesting that skin contact during winemaking significantly influences extraction and final wine sesquiterpene content.
Sesquiterpene_Production is subclass of Terpene_Accumulation representing C15 terpenoid biosynthesis. Distinguishes from Monoterpene_Synthesis which generates C10 compounds. Sibling to Monoterpene_Synthesis and Terpene_Pool_Development. Rotundone is key sensory-active compound in peppery varieties. Mevalonate pathway provides FPP precursor. Cool temperatures enhance rotundone accumulation. Skin-localized synthesis.
C15 terpene synthesis
C15-Terpen-Synthese
Rotundon-Produktion
Sesquiterpen-Produktion
Sesquiterpenoid-Biosynthese
farnesyl diphosphate-derived terpene synthesis
rotundone production
sesquiterpenoid biosynthesis
Wood C et al (2008) From wine to pepper: rotundone, an obscure sesquiterpene, is a potent spicy aroma compound. Journal of Agricultural and Food Chemistry 56: 3738-3744 Siebert TE et al (2008) Development of a stable isotope dilution analysis for the grape-derived compound rotundone. Journal of Agricultural and Food Chemistry 56: 3745-3748 Drew DP et al (2016) Two key polymorphisms in a newly discovered sesquiterpene synthase gene. Phytochemistry 122: 204-215
Sesquiterpene Production
Severe deficiency quality diagnosis for nitrogen in Central Valley Thompson Seedless demanded emergency intervention to prevent vine collapse in a vineyard affected by complete fertilization system failure during the critical spring growth period. Visual assessment revealed severe deficiency quality with intense chlorosis throughout the canopy (score 4.5 on 0-5 scale), leaves uniformly pale yellow rather than green, dramatic growth cessation with shoot length 65% below normal at 45 cm versus expected 130 cm, extensive premature senescence with 40% of basal canopy already defoliated by mid-summer, and shoot tips showing die-back indicating that even meristematic activity was compromised by nutrient limitation. Tissue analysis confirmed severe deficiency quality with petiole N at 0.28% (critically deficient, 50% below adequacy threshold of 0.60%), the lowest N status recorded in regional monitoring, consistent with the complete absence of nitrogen supply during the current season. Functional assessment revealed severe deficiency quality with photosynthetic capacity reduced 70% in remaining leaves, fruit set dramatically impaired with 60% coulure, and surviving berries showing minimal development with Brix at only 8° at the véraison date when 18° would be normal. The severe deficiency quality diagnosis triggered emergency intervention including immediate high-rate fertigation with UAN solution at 200 kg N/ha, repeated foliar urea application at 2% concentration (4 applications at weekly intervals), 50% crop removal to reduce demand and redirect resources to vine recovery, and extended irrigation to support the recovery process. Season outcome showed partial canopy recovery but only 25% of normal yield salvaged, with follow-up assessment revealing reserve depletion (trunk starch 40% below normal) indicating that full recovery would require 2-3 seasons of careful management to restore vine productivity and health to normal levels.
Severe deficiency quality is a deficiency severity quality that inheres in a grapevine organism and represents a nutritional limitation of maximum intensity where tissue nutrient concentrations have declined to critically low levels, with symptoms intense and extensive throughout the canopy including advanced manifestations such as tissue necrosis, defoliation, and growing point damage, and with functional impacts including dramatic growth cessation, major photosynthetic impairment, and substantial yield loss that may extend to multiple seasons through carbohydrate reserve depletion and structural damage. This quality captures the most serious stage of deficiency requiring emergency intervention, recognizing that severe deficiency represents critical nutritional stress causing major current-season losses while also potentially compromising future productivity through vine structural damage and reserve depletion. The ontological significance of severe deficiency quality lies in its indication that immediate intensive intervention is required to prevent vine decline or death, that current-season crop loss is likely unavoidable, and that multi-season recovery may be necessary to restore normal productivity. The biological basis of severe deficiency encompasses tissue concentrations at or below the minimum required for basic cellular function, widespread metabolic disruption affecting all growth and development processes, extensive visible damage reflecting cellular death and tissue dysfunction, and systemic effects including reserve depletion that create legacy impacts extending beyond the immediate correction of nutrient status. The visual symptoms of severe deficiency show maximum intensity expression (scores 4-5 on 0-5 scales) distributed throughout the canopy with advanced manifestations including necrosis, defoliation, die-back, and growing point damage, creating dramatic visual impact that is immediately obvious to even casual observation. The management response to severe deficiency requires emergency intervention with maximum fertilization rates, often including both intensive soil application and repeated foliar treatments, combined with other supportive measures such as reduced crop load, irrigation optimization, and potentially vine rehabilitation measures extending over multiple seasons.
Severe_Deficiency_Quality is_a Deficiency_Severity_Quality, representing maximum-intensity deficiency with extreme symptoms, major functional impairment, and multi-season impacts. This class addresses the most serious severity category requiring emergency intervention. Symptoms intense (score 4-5) and canopy-wide including advanced features (necrosis, defoliation, die-back). Tissue concentrations critically low, often 50%+ below adequacy thresholds. Current-season crop loss likely unavoidable. Reserve depletion creates multi-season recovery requirement. Emergency high-rate fertilization + crop reduction + supportive measures required.
Schwerer Mangel
acute deficiency
advanced deficiency
akuter Mangel
critical deficiency
extreme nutritional limitation
fortgeschrittener Mangel
kritische Nährstofflimitation
Severe Deficiency Quality
Severe water stress quality occurrence in a dryland Ribera del Duero Tempranillo vineyard during an extreme drought vintage demonstrated the quality and health consequences requiring prevention through irrigation where available. Predawn leaf water potential declined below -0.7 MPa by mid-August with midday stem water potential reaching -1.5 MPa, exceeding all beneficial stress thresholds and entering the severe water stress quality category indicated by visible symptoms including persistent midday wilting, 40% basal leaf loss by véraison, and evident berry shriveling. Fruit composition reflected severe stress damage with sugar accumulation stalling at 22°Brix despite extended hangtime, pH rising to 3.95 from potassium concentration, titratable acidity declining to 4.2 g/L from malic acid respiration under stress-elevated tissue temperatures, and varietal character compromised by dominant dried fruit and raisined notes rather than fresh Tempranillo expression. Wine quality scores averaged 12 points below the five-year mean despite favorable early-season conditions, demonstrating that severe water stress quality exceeds the beneficial stress window regardless of concentration effects on some parameters. Post-harvest vine assessment revealed 15% shoot dieback and reduced bud fruitfulness for the following season, confirming lasting damage from severe stress exposure and motivating supplemental irrigation infrastructure investment to prevent recurrence.
Severe water stress quality is a water deficit severity quality that inheres in a grapevine organism and represents the extreme category of water deficit characterized by critical stomatal closure and non-stomatal limitations severely restricting photosynthetic function, hydraulic dysfunction including cavitation and embolism formation in xylem vessels, accelerated leaf senescence and abscission, impaired fruit ripening from inadequate photoassimilate supply, and potential for lasting damage to vine health and productivity that may persist beyond the current season when stress relief occurs. This quality defines the stress threshold beyond which quality benefits reverse to quality compromise and where vine health concerns supersede production objectives, representing the upper limit that irrigation management must prevent regardless of quality enhancement goals associated with milder stress categories. The ontological significance of severe water stress quality lies in its identification of the damage threshold where water deficit transitions from beneficial stress to harmful stress, establishing the boundary that irrigation intervention must respect to protect vine assets, maintain fruit quality, and preserve long-term vineyard productivity. The biological basis of severe water stress quality encompasses the failure of normal regulatory mechanisms to maintain hydraulic function, with xylem water potential declining below cavitation thresholds causing gas bubble formation that blocks water transport, leaf water content falling below survival thresholds causing cellular damage and accelerated senescence, and photoassimilate limitation from severe photosynthetic restriction preventing normal fruit development including sugar accumulation, acid metabolism, and flavor compound biosynthesis. The irreversibility of certain severe stress effects distinguishes this category from milder stress levels, with cavitation-induced hydraulic damage potentially persisting through the growing season and canopy loss from premature senescence eliminating photosynthetic capacity that cannot be rapidly restored. Quality impacts of severe water stress include incomplete sugar accumulation, elevated pH from potassium concentration in reduced-volume berries, diminished varietal aromatic character, and potential development of stress-induced off-flavors that compromise wine quality regardless of concentration effects. Physiological indicators of severe water stress quality include predawn leaf water potential below -0.6 MPa, midday stem water potential below -1.2 to -1.4 MPa depending on cultivar, and visible symptoms including persistent leaf wilting not recovering overnight, extensive basal leaf yellowing and abscission, shriveled berries, and in extreme cases, rachis necrosis and cluster desiccation.
Severe_Water_Stress_Quality is_a Water_Deficit_Severity_Quality, representing the extreme stress intensity category with damaging physiological consequences. This class addresses the threshold beyond which stress causes harm rather than benefit. Typical predawn LWP below -0.6 MPa, midday SWP below -1.2 to -1.4 MPa indicate severe stress requiring intervention. Distinguished from Moderate_Water_Stress_Quality by irreversible damage potential, impaired ripening, and lasting vine health consequences. The quality compromise despite concentration effects represents key distinction from beneficial stress categories. Critical for identifying irrigation intervention thresholds preventing damage to vine assets and wine quality.
critical water stress
damaging water deficit
extreme drought stress
extremer Trockenstress
harmful water stress
kritisches Wasserdefizit
schwerer Wasserstress
schädigender Wasserstress
Lovisolo C et al. (2010) Drought-induced changes in development and function of grapevine (Vitis spp.) organs and in their hydraulic and non-hydraulic interactions at the whole-plant level: a physiological and molecular update. Functional Plant Biology 37:98-116 Hochberg U et al. (2017) Stomatal closure, basal leaf embolism, and shedding protect the hydraulic integrity of grape stems. Plant Physiology 174:764-775 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data. Annals of Botany 105:661-676
Severe Water Stress Quality
Merlot severely overripe berries at week 22 (4 weeks post-optimal) show 31% sugars (extreme for table wine), 3.8 g/L titratable acidity (severely depleted), pH 4.25 (dangerously high), 20% berry weight loss from dehydration creating shriveled appearance, dominant prune and raisin flavors with no fresh fruit character, 15% Botrytis infection, producing unacceptable wine with 18% alcohol, acetic acid from Botrytis metabolism, severe structural imbalance, brown color, unsuitable for commercial table wine production. Zinfandel severely overripe berries showing characteristic extreme ripeness= 33% sugars, 4.2 g/L acidity, pH 4.18, extensive raisining with berries at 70% of peak weight due to dehydration, producing wine requiring arrested fermentation to avoid 19+% alcohol, showing Port-like character inappropriate for table wine, demonstrating Zinfandel tendency toward severe overripeness in warm climates requiring careful harvest timing. Chardonnay severely overripe white berries showing white wine extreme overripeness= 27% sugars (excessive for white wine), 4.5 g/L acidity (very low), pH 3.85 (very high for white), golden-brown berry color, honey and dried apricot flavors replacing fresh fruit, producing wine with 15.5% alcohol, flabby heavy mouthfeel, oxidized character, complete loss of Chardonnay varietal typicity, suitable only for distillation or bulk blending. Cabernet Sauvignon severely overripe berry salvage attempt= tartaric acid addition of 3 g/L restoring titratable acidity from 3.5 to 6.5 g/L, pH from 4.3 to 3.95 (partial improvement), fermentation to 16.5% alcohol from 32% sugars, extensive new oak attempting to mask cooked fruit defects, producing marginally acceptable wine scoring 78/100 (barely acceptable quality) demonstrating limited remediation potential for severe overripeness. Syrah harvest delay experiment comparing optimal (week 18= 25% sugars, quality score 91/100), slightly overripe (week 19.5= 27% sugars, score 87/100), and severely overripe (week 22= 32% sugars, score 72/100, unacceptable quality) showing progressive quality degradation with severe overripeness creating fundamental quality failure, emphasizing importance of timely harvest before progression to severe overripeness.
A severely overripe berry is an overripe berry showing extreme progression beyond optimal ripeness with severe compositional imbalances, significant dehydration or raisining, flavor degradation, and quality unsuitable for conventional table wine production, representing berries 3+ weeks past optimal harvest timing. Severely overripe berries are characterized by= (1) sugars at very high levels typically >28-30% (often 30-35+% in extreme cases), creating excessive alcohol potential (>16-18% potential alcohol) unbalanced for table wine production (2) titratable acidity at very low levels typically 4.1-4.3, creating serious microbiological instability risks and wine structure collapse (4) severe flavor degradation with loss of varietal character, dominant cooked/stewed/raisined flavors, potential oxidative or fermentation-derived off-flavors (5) significant berry dehydration with 10-30+% weight loss from peak creating shriveled raisined appearance (6) berry texture extremely soft or desiccated (7) high incidence of fungal infection (Botrytis, bunch rots) and insect damage on compromised over-mature berries. Severely overripe berries develop through extended hang time 3+ weeks past optimal harvest= continued slow sugar accumulation through concentration as dehydration proceeds severe acid depletion through continued respiration plus concentration masking effects progressive loss of fresh fruit character and development of cooked/raisined flavors potential noble rot development (Botrytis cinerea) in humid conditions or bunch rot in warm humid conditions. Severely overripe berries show multiple quality defects= wines exhibiting excessive alcohol (16-19%), severely flabby structure with very low acid and high pH, cooked prune or raisin dominant flavors with loss of varietal typicity, brown color (whites) or purple-brown color (reds), oxidative character, potential volatile acidity from Botrytis or other microorganisms, unbalanced sweet heavy mouthfeel even in dry wines these defects make severely overripe fruit generally unsuitable for quality table wine production. Severely overripe berries may have intentional uses in specialized production= late-harvest sweet wines where extreme ripeness and noble rot are desired passito or Amarone styles using intentionally dried raisined grapes certain warm-climate extreme styles accepting very high alcohol and concentrated character but for conventional premium table wine, severe overripeness represents fundamental quality failure. Severely overripe berries show limited winemaking remediation potential= acid additions can partially restore titratable acidity but cannot fully correct pH dilution can reduce sugar but dilutes everything including color and flavor severe flavor defects (cooked, raisined, oxidized) cannot be corrected microbial risks from high pH require careful handling overall, severely overripe fruit compromises quality beyond what winemaking can compensate. Severely overripe berries result from= harvest delays due to weather (extended wet period preventing harvest) operational constraints preventing timely harvest misguided extended hang time strategies exceeding beneficial limits vineyard abandonment or neglect these berries may be unsuitable for harvest or may be harvested for bulk/distillation use rather than premium wine.
Severely_Overripe_Berry represents extreme overripeness and is distinguished from Slightly_Overripe_Berry by extreme versus moderate compositional imbalances and quality compromise. Severely overripe berries are generally unsuitable for quality table wine production except in specialized late-harvest styles intentionally using overripe or raisined fruit. Understanding severely overripe berry characteristics is important for recognizing quality failures from excessive harvest delay, for specialized production styles using intentionally overripe fruit, and for emphasizing the importance of timely harvest preventing severe overripeness.
Rosinen-Stadium-Traube
extrem zu reife Traube
extremely overripe berry
raisined berry
stark überreife Beere
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 255-265 Jackson, D.I., Lombard, P.B. (1993). Environmental and management practices affecting grape composition. American Journal of Enology and Viticulture 44=409-430.
Severely Overripe Berry
A shade mature leaf at node 8 on an interior shoot of Vitis vinifera cv. Pinot Noir in Burgundy, France, positioned deep within canopy receiving approximately 25% of full sunlight (daily photosynthetic photon flux density averaging 380 μmol photons m⁻² s⁻¹), measuring 16.5 cm length by 15.2 cm width representing 42% size increase compared to exterior sun leaves despite reduced total biomass allocation, thin lamina of 215 μm with single palisade layer of 45 μm and expanded spongy mesophyll of 150 μm with large intercellular air spaces, chlorophyll content of 285 μg cm⁻² (compared to 520 μg cm⁻² in sun leaves) but 48 μg mg⁻¹ dry mass (compared to 32 μg mg⁻¹ in sun leaves), thin cuticle of 2.8 μm, reduced stomatal density of 135 stomata mm⁻² with large individual stomata, sparse trichomes creating pale green undersurface appearance, light green adaxial surface without anthocyanin pigmentation, demonstrating light-saturated photosynthesis rate of 7.2 μmol CO₂ m⁻² s⁻¹ but low light compensation point of 18 μmol photons m⁻² s⁻¹ enabling positive carbon balance under low light, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing characteristic shade leaf adaptations maximizing light capture and carbon gain efficiency under light-limited conditions.
A mature leaf developing and functioning in low light environments receiving less than 30-40% of full sunlight in Vitis vinifera, characterized by phenotypic adaptations to light limitation including larger lamina size typically 20-50% increased compared to sun leaves providing greater light-intercepting surface area despite resource constraints, thinner lamina often 180-250 μm due to reduced cell layers and wall thickness, single palisade mesophyll layer with increased spongy mesophyll proportion maximizing light capture through internal reflection and scattering, lower chlorophyll concentrations per unit area but higher per unit mass reflecting thin tissue structure, thinner cuticle reducing investment in protection that provides limited benefit in shaded environments, lower stomatal density with larger individual stomata maintaining adequate gas exchange with reduced construction cost, reduced trichome density, lighter green coloration from lower total chlorophyll despite often higher chlorophyll b proportions for enhanced light harvesting under low light, reduced concentrations of photoprotective compounds unnecessary in low-light conditions, and physiological characteristics including lower light-saturated photosynthesis rates of 5-9 μmol CO₂ m⁻² s⁻¹ but lower light compensation points of 15-25 μmol photons m⁻² s⁻¹ enabling net carbon gain under very low light conditions, and higher quantum efficiency reflecting optimization for light capture rather than maximum capacity.
Shade mature leaves are distinguished from sun leaves by structural and functional characteristics reflecting low-light acclimation, with assessment requiring anatomical examination, pigment analysis, and physiological measurements under ambient light conditions. The shade adaptation represents developmental plasticity responding to low light during expansion, with light environment during first 2-3 weeks determining anatomical differentiation. Shade leaves show lower absolute photosynthesis than sun leaves but superior efficiency per unit resource invested, with high surface area per biomass and low respiration creating positive carbon balance at lower light intensities where sun leaves would show net carbon loss. Interior canopy positions typically produce shade leaves, but severe shading reducing light below 15-20% full sun may produce leaves with inadequate photosynthetic capacity to maintain positive carbon balance, requiring removal through leaf removal management. Excessive shade leaf proportion (>60% of canopy) indicates dense canopy requiring management through shoot thinning, leaf removal, or training adjustments to improve light penetration.
innenkanopie Weinblatt
interior canopy leaf
low light leaf
niedriglicht Blatt
schattenreifes Blatt
shade-adapted leaf
Based on Boardman (1977) "Comparative photosynthesis of sun and shade plants" Annual Review of Plant Physiology 28=355-377 Poni et al. (2006) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 57=397-407 Smart (1985) "Principles of grapevine canopy microclimate manipulation with implications for yield and quality" American Journal of Enology and Viticulture 36=230-239 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 273-304.
Shade Mature Leaf
A shaded lower canopy leaf at node 4 on an interior shoot of Vitis vinifera cv. Grenache in a dense canopy in southern France, experiencing approximately 22% of full sunlight (daily PPFD averaging 340 μmol photons m⁻² s⁻¹) due to shading by 16 overlying leaves creating 2.4 m² light-intercepting area, measuring 17.2 cm length by 15.8 cm width showing larger size typical of shade leaves despite basal position, exhibiting thin lamina of 205 μm with single palisade layer of 42 μm and expanded spongy mesophyll, pale green coloration with chlorophyll content of 270 μg cm⁻², very thin cuticle of 2.6 μm, low stomatal density of 128 stomata mm⁻² with large individual stomata, demonstrating light-saturated photosynthesis rate of only 6.2 μmol CO₂ m⁻² s⁻¹ but low light compensation point of 16 μmol photons m⁻² s⁻¹, achieving marginal daily carbon balance with photosynthetic gain of approximately 22 mg carbohydrate barely exceeding respiratory costs of 18-20 mg per day, showing vulnerability indicators including slight chlorosis and reduced turgor suggesting approaching transition to carbon sink status, measured at BBCH stage 79 (majority of berries touching) in late July, representing severely shaded lower canopy position with questionable carbon economics requiring canopy management consideration.
A lower canopy basal leaf experiencing substantial light limitation from upper canopy shading in Vitis vinifera, characterized by exposure to less than 30-40% of full sunlight due to overlying leaf layers, shoot density, or canopy architecture creating dense light-intercepting structures, development of shade leaf anatomical characteristics including thin lamina of 180-250 μm with single palisade layer, low chlorophyll per unit area but high per unit mass, reduced cuticle development, large stomata at low density, and pale green coloration, reduced photosynthetic capacity typically 40-65% of sun-exposed basal leaves with light-saturated rates of 5-9 μmol CO₂ m⁻² s⁻¹, low light compensation point enabling positive carbon balance under low light but marginal whole-leaf carbon economics when respiratory costs are considered, functional contribution limited by both reduced light availability and inherently lower photosynthetic capacity, vulnerability to transitioning from marginal carbon balance to net carbon sink if environmental stresses increase respiration or further reduce already-limited photosynthetic capacity, and common targeting for removal during canopy management practices aimed at eliminating non-productive shaded zones improving air circulation and disease management. Shaded lower canopy leaves develop either from shade conditions present during expansion creating shade leaf anatomy, or from initially sun-exposed conditions followed by progressive shading as upper canopy develops, with the latter showing sun leaf anatomy but functioning under shade conditions creating photosynthetic inefficiency as anatomical features optimized for high light operate suboptimally under light limitation.
Shaded lower canopy leaves represent suboptimal functional outcomes of dense canopy architecture, distinguished from adequately illuminated lower canopy leaves by severe light limitation and marginal carbon balance. Light availability below 25-30% full sun typically creates photosynthetic capacity insufficient to maintain positive carbon balance, with exact thresholds varying by cultivar, temperature, and respiratory costs. Shade leaf anatomy develops in response to low light during expansion and represents appropriate developmental plasticity maximizing carbon gain under light limitation, but cannot fully compensate for severe shading with photosynthetic capacity remaining well below sun leaves. Management implications include recognition that shaded lower canopy leaves contribute minimally to vine carbon balance while consuming maintenance resources and creating favorable environments for fungal diseases through poor air circulation and high humidity. Removal through strategic leaf removal or shoot thinning typically improves whole-vine carbon balance, fruit microclimate, and disease management, with timing considerations balancing photosynthetic capacity loss against benefits. Dense canopies requiring substantial shaded leaf removal indicate excessive vegetative vigor requiring management through reduced fertilization, increased crop load, or deficit irrigation to achieve optimal balance.
beschattetes untere Kanopie Blatt
interior lower leaf
lichtlimitiertes basales Blatt
light-limited basal leaf
schatten-gestresstes Weinblatt
shade-stressed basal leaf
Shaded Lower Canopy Leaf
A shallow basal lobe representing the right basal lobe on a mature leaf at node 6 of Vitis vinifera cv. Sauvignon Blanc in Sancerre, France, exhibiting minimal development with dimensions of 1.8 cm width and 1.2 cm length representing the smallest structure on this leaf, poorly separated from lower lateral lobe by very shallow or absent lower right lateral sinus of 0.3 cm depth creating indistinct boundary, lobe barely extending to the level of petiolar junction without substantial basal projection, vascularized by relatively small basal primary lateral vein 1.2 mm diameter, lobe oriented with divergent outward direction creating wide separation between left and right basal lobe margins, basal lobe margin positioned 2.8 cm from opposing left basal margin creating the characteristic wide open V-shaped petiolar sinus diagnostic for this cultivar, lobe exhibiting poorly defined apex or rounded edge without distinct point, lobe area of 1.8 cm² representing only 4% of total blade area substantially smaller than all other lobes, minimal basal lobe development creating narrow leaf base and distinctive open petiolar sinus architecture, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing minimal shallow basal lobe expression characteristic of narrow-based cultivars.
A basal lobe exhibiting minimal development with small size and weak dissection in Vitis vinifera, characterized by reduced lobe structure with small dimensions substantially smaller than all other lobe types, shallow or absent separation from adjacent lateral lobes with lower lateral sinuses minimal or not clearly defined, limited extension barely reaching or not extending beyond petiolar junction level, weak vascularization by relatively small basal primary lateral veins, divergent orientation with lobe margins directed outward away from petiolar sinus, and role in creating open petiolar sinus configuration through divergent basal lobe positioning leaving substantial gap between lobe margins. Shallow basal lobes represent minimal basal lobe expression, occurring in cultivars with narrow leaf bases and weak overall lobe dissection (Sauvignon Blanc, Riesling, Cabernet Sauvignon, Chardonnay), creating distinctive leaf base architecture with poorly developed basal structures creating open access to petiolar junction, and serving as diagnostic features through effects on narrow leaf base shape and particularly open petiolar sinus configuration.
Shallow basal lobes are distinguished from deep basal lobes by minimal size, weak separation, and poor development. The shallow designation emphasizes limited lobe expression creating weak basal structures, contrasting with pronounced deep basal lobes. Size substantially smaller than all other lobe types is characteristic, with shallow basal lobes typically representing only 3-8% per lobe of total area compared to 10-15% for typical basal lobes and 8-12% for deep basal lobes. Separation from lateral lobes is often indistinct with very shallow or completely absent lower lateral sinuses creating integrated basal-lateral complexes rather than clearly defined basal lobes. Limited extension barely reaching petiolar junction level creates minimal basal projection contrasting with deep basal lobes extending substantially beyond junction. Divergent orientation is characteristic, with shallow basal lobes directed outward creating open petiolar sinuses with wide gaps between lobe margins, contrasting with convergent deep basal lobes creating closed or overlapping sinuses. Occurrence predominates in narrow-based cultivars including most major varieties (Cabernet Sauvignon, Sauvignon Blanc, Riesling, Chardonnay), with broad-based cultivars relatively uncommon. Functional contribution involves minimal area provision typically 5-12% combined area, with greater significance for creating characteristic narrow base and open sinus architecture. Ampelographic utility is substantial through effects on petiolar sinus configuration, with shallow divergent basal lobes creating open sinuses serving as diagnostic character distinguishing narrow-based from broad-based cultivars.
flacher Basallappen
minimal petiolar lobe
minimaler Petiolarlappen
poorly developed base lobe
schwach entwickelter Basislappen
weak basal lobe
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 120-182 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 119-154.
Shallow Basal Lobe
A shallow lower lateral sinus representing the left lower lateral sinus on a mature leaf at node 6 of Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, California, positioned in the transition zone between lateral and basal lobe regions, exhibiting minimal depth of 1.2 cm measured from barely perceptible indentation to lobe edge connection line, penetration reaching only approximately 22% of distance from margin toward median vein, poorly defined configuration with gradual curvature rather than distinct sinus base, width of 1.8 cm at nominal opening creating depth-to-width ratio of 0.67 indicating width exceeding depth, shallow sinus creating weak indistinct separation between lower lateral and basal lobe regions producing integrated rather than clearly differentiated appearance, sinus base positioned 7.5 cm from petiolar junction in lower blade region, sinus margins showing gradual transitions without clear boundaries, shallow lower sinus contrasting with moderately deep upper lateral sinus (3.0 cm) on same leaf reflecting typical pattern of decreasing dissection depth from upper to lower blade regions, bilateral symmetry with right lower lateral sinus showing similarly shallow configuration (1.1 cm), measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing typical shallow minimal lower lateral sinus development characteristic of most moderately lobed cultivars.
A lower lateral sinus exhibiting minimal depth creating weak or barely perceptible separation between lower lateral lobes and adjacent lobes in Vitis vinifera, characterized by depth typically less than 1.5 cm representing minimal dissection in lower blade regions, poorly defined or absent configuration with gradual transitions rather than distinct sinus bases, penetration often only 15-30% of distance from margin toward median vein, minimal visual prominence creating indistinct lower lateral lobe boundaries, occurrence in moderately to weakly lobed cultivars showing reduced overall lobe development particularly in lower blade regions, and role in creating poorly differentiated lower lateral lobes or integrated lateral-basal lobe complexes characteristic of weakly dissected cultivar morphology. Shallow lower lateral sinuses represent minimal or absent lower sinus development, occurring in most cultivars including moderately lobed five-lobed varieties and all three-lobed configurations, creating typical architecture with poorly defined or absent lower lateral lobe separation, and serving as the common default condition contrasting with exceptional deep lower sinuses restricted to deeply dissected cultivar groups.
Shallow lower lateral sinuses represent the common default condition in grapevine leaf morphology, distinguished from deep lower sinuses by minimal depth <1.5 cm and poorly defined configuration. The shallow designation emphasizes minimal dissection typical of lower blade regions in most cultivars, contrasting with exceptional deep lower sinuses. Depth substantially less than upper lateral sinuses is the normal pattern, with typical upper=lower sinus depth ratios of 2=1 to 3=1. Configuration is often so poorly defined that sinus identification is ambiguous, with gradual transitions creating uncertainty about whether distinct lower lateral sinuses exist versus simple integrated lower blade regions. Width often exceeds depth creating broad shallow indentations barely distinguishable from normal blade curvature. Occurrence predominates in moderately and weakly lobed cultivars representing the majority of commercial varieties, with only deeply dissected cultivars showing well-developed lower sinuses. Absence is the extreme of shallow development, occurring in three-lobed cultivars completely lacking lower lateral lobe differentiation. Genetic control affects overall lobe pattern with five-lobed cultivars showing at least minimal lower sinuses while three-lobed patterns lack them, though depth shows environmental sensitivity. Functional implications are minimal given shallow depth. Ampelographic utility is limited, primarily serving to confirm moderately lobed five-lobed patterns rather than providing specific cultivar identification, though shallow lower sinuses combined with other features contribute to overall morphological profiles distinguishing moderately from deeply dissected cultivar groups.
barely incised middle sinus
flache untere Seitenbucht
minimal lower lateral sinus
minimale untere Lateralbucht
schwach eingeschnittene mittlere Sinus
weak lower sinus
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 136-215 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 119-172.
Shallow Lower Lateral Sinus
Chardonnay vineyard showing shallow roots in loam soil, soil core sampling 0-100 cm depth revealing root distribution, shallow zone 0-40 cm containing 68% of total root length and 82% of fine roots <2 mm diameter, root length density 4.2 cm/cm³ in 10-30 cm layer versus 0.8 cm/cm³ at 60-80 cm, isotope tracing showing shallow roots providing 73% of vine nitrogen uptake and 79% of water uptake during May-June growing period, shallow roots demonstrating critical absorption function in upper soil profile
Shallow root is root or root portion located in upper soil profile typically within 0-50 cm depth though definition varying with soil type and vine management, characterized by position in topsoil layers where organic matter, nutrients, moisture, and biological activity typically highest, showing rapid proliferation and turnover responding to favorable surface conditions, demonstrating high root density creating intensive soil exploration in shallow zone, and serving critical absorption function providing majority of vine water and nutrient uptake particularly during active growth when shallow soil resources abundant, with shallow roots in grapevine showing typical concentration 10-40 cm depth where 40-70% of total root length and 50-80% of fine absorptive roots located depending on soil conditions and irrigation, exhibiting seasonal dynamics with shallow root activity peaking spring and fall when soil temperature and moisture optimal declining summer heat or winter cold, and demonstrating management sensitivity with shallow roots highly responsive to irrigation, fertilization, tillage, and cover crops making shallow zone primary target for viticultural practices affecting root function, and representing zone where vine-soil interactions most intense with shallow roots encountering highest microbial populations, greatest nutrient availability, most variable moisture, and strongest competition from weeds or cover crops. Shallow root depth definition includes= showing typical range (shallow roots generally 0-50 cm depth though precise definition varying with context), exhibiting topsoil association (shallow roots predominantly in A horizon and upper B horizon soil layers), demonstrating relative position (shallow versus deep defined relative to total rooting depth with shallow being upper portion), revealing management context (shallow depth relevant to tillage depth, irrigation depth, and other practices), and displaying soil-dependent variation (shallow zone deeper in deep soils, shallower in shallow soils or those with restrictive layers). Shallow root density and abundance includes= showing high root length density (shallow zone typically 1-10 cm root per cm³ soil substantially higher than deep zones), exhibiting fine root concentration (shallow zone containing 50-80% of total fine absorptive root length), demonstrating root mass proportion (shallow roots comprising 30-60% of total root system biomass despite occupying small soil volume), revealing intensive exploration (shallow roots creating dense network thoroughly exploiting upper soil), and displaying proliferation capacity (shallow roots rapidly increasing density in response to favorable conditions). Shallow root functional importance includes= providing majority uptake (shallow roots supplying 60-85% of vine water and nutrient uptake during growing season), exhibiting rapid response (shallow roots quickly responding to rainfall, irrigation, or fertilization), demonstrating seasonal dominance (shallow roots particularly important spring and early summer when upper soil moist and fertile), revealing nutrient specialization (shallow roots especially important for nitrogen, phosphorus, potassium uptake given surface nutrient concentration), and displaying mycorrhizal associations (shallow fine roots showing high arbuscular mycorrhizal colonization enhancing uptake). Shallow root environmental conditions includes= experiencing variable moisture (shallow soil showing rapid wetting and drying creating dynamic moisture environment), showing optimal temperature (shallow soil warming quickly spring promoting early root activity), exhibiting high nutrient availability (organic matter decomposition and fertilizer placement concentrating nutrients in shallow zone), demonstrating biological activity (shallow soil harboring most abundant and diverse microbial communities), and displaying physical disturbance (shallow roots exposed to tillage, traffic compaction, and other surface disturbances). Shallow root seasonal dynamics includes= showing spring proliferation (shallow roots growing rapidly spring as soil warms and moisture adequate), exhibiting summer stress (shallow roots potentially experiencing drought stress if irrigation inadequate), demonstrating fall recovery (shallow root growth resuming fall with cooler temperatures and autumn rainfall), revealing winter dormancy (shallow root activity minimal winter though roots surviving if soil not frozen), and displaying phenological coordination (shallow root activity coordinating with shoot growth and reproductive development). Shallow root management implications includes= requiring irrigation consideration (shallow roots benefiting from frequent light irrigation maintaining upper soil moisture), showing fertilization response (shallow-placed fertilizers efficiently accessed by concentrated shallow roots), exhibiting tillage sensitivity (cultivation damaging shallow roots potentially reducing uptake capacity), demonstrating mulch benefits (surface mulches moderating shallow soil temperature and moisture benefiting roots), and revealing cover crop interactions (cover crops competing with shallow vine roots for water and nutrients requiring management). Shallow root versus deep root includes= contrasting in depth (shallow 0-50 cm versus deep >50-100 cm), showing different functions (shallow emphasizing absorption versus deep emphasizing drought tolerance and anchorage), exhibiting seasonal patterns (shallow dominant growing season versus deep important summer drought), demonstrating turnover rates (shallow roots showing higher turnover than deep structural roots), and revealing management sensitivity (shallow roots more affected by surface practices than deep roots).
Shallow_Root emphasizes upper soil profile position and topsoil association and is distinguished from deep roots by focus on 0-50 cm zone. Includes Feeder_Root hierarchy. Critical for understanding topsoil concentration, high density, majority uptake, seasonal dynamics, management sensitivity, and the roots in upper soil layers providing primary vine resource acquisition particularly during active growth.
Flache Wurzel
Oberbodenwurzel
Oberflächenwurzel
Surface root
Topsoil root
Upper root zone
Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press. | Smart, D. R., et al. (2006). Grapevine rooting patterns= A comprehensive analysis and a review. American Journal of Enology and Viticulture, 57(1), 89-104.
Shallow Root
Chardonnay grafted on Riparia Gloire rootstock at 14 years in vineyard with dense clay layer at 75 cm depth showing shallow tap root extending only 68 cm maximum depth limited by impenetrable clay hardpan, shallow tap root diameter 7.2 cm at crown tapering to 2.8 cm at maximum depth, excavation revealing shallow tap root terminating abruptly at hardpan unable to penetrate despite continued growth attempts visible as root tip proliferation at barrier, lateral root network extending 180 cm radius providing majority of root system but concentrated 20-60 cm depth, shallow tap root and lateral system adequate for vine function under irrigation but requiring supplemental water mid-summer when upper 50 cm soil dry, shallow rooting characteristic of Riparia genetics combined with soil limitation
Shallow tap root is tap root limited to relatively shallow depth typically less than 100 cm and often only 50-80 cm maximum penetration, emphasizing limited vertical extent and concentration in upper soil horizons, characterized by genetic determination by shallow-rooting rootstock varieties particularly Vitis riparia-based cultivars, restriction by soil limitations including shallow bedrock, hardpans, water tables, or compacted layers preventing deeper penetration, and functional emphasis on upper soil resource exploitation rather than deep water access, with shallow tap root development typical of low-vigor rootstocks suited to fertile soils where shallow rooting adequate, Riparia genetics conferring inherent depth limitation, and shallow soils physically preventing deep penetration regardless of genetic potential making shallow tap root common situation in many vineyard settings. Shallow tap root depth limitation includes= remaining under 100 cm (shallow tap root by definition not exceeding this depth threshold distinguishing from typical or deep tap roots), commonly limited to 50-80 cm (shallow tap roots frequently restricted to this depth range in practice), showing minimal deep soil exploration (shallow tap root failing to exploit deep soil layers and moisture reserves), demonstrating limited root, shoot ratio depth (shallow tap root possibly not extending substantially deeper than lateral root network reducing vertical root system extent), and exhibiting restricted soil volume access (shallow tap root limiting total soil volume exploitable by root system constraining resource availability). Shallow tap root genetic determination includes= characteristic of Riparia types (Vitis riparia and hybrids showing inherent shallow rooting producing tap roots rarely exceeding 80-100 cm), typical of low-vigor rootstocks (varieties like Riparia Gloire, 101-14, 3309 Couderc producing shallow tap roots as part of compact root architecture), showing breeding selection (shallow rooting sometimes specifically selected for rootstock varieties intended for fertile irrigated sites where deep roots unnecessary), demonstrating genetic constraint (Riparia-based rootstocks unable to produce deep tap roots even in favorable soil conditions reflecting inherent genetic limitation), and varying among hybrids (rootstock hybrids showing intermediate depth potential reflecting genetic contributions with Riparia parentage generally reducing tap root depth). Shallow tap root soil-caused limitation includes= restricted by shallow bedrock (solid rock layer at shallow depth absolutely preventing further tap root penetration), limited by hardpan (dense compacted soil layer or cemented horizon blocking tap root descent), constrained by water table (permanent shallow water table creating anaerobic conditions preventing tap root growth below), inhibited by toxic layers (accumulations of salts, sodium, aluminum, or other toxic substances at depth preventing tap root penetration), and blocked by compaction (severe soil compaction creating impenetrable layer limiting tap root to shallow zone). Shallow tap root water access limitations includes= restricted to shallow moisture (shallow tap root accessing only upper soil water unavailable to exploit deep moisture reserves), vulnerable to summer drought (shallow tap root failing during surface soil drying when deep roots would maintain function), requiring irrigation in dry climates (shallow tap root inadequate for unirrigated production in Mediterranean or arid regions necessitating supplemental water), showing seasonal dependence (shallow tap root contribution to vine water supply adequate spring and fall but potentially insufficient mid-summer), and demonstrating limited drought tolerance (shallow tap root providing poor drought tolerance compared to deep-rooted systems requiring careful water management). Shallow tap root functional adaptations includes= exploiting upper soil resources (shallow tap root efficiently using nutrients and moisture in fertile upper horizons where most available), suited to fertile sites (shallow tap root adequate on rich soils with high native fertility and good moisture retention not requiring deep resource exploitation), enabling low-vigor response (shallow tap root contributing to overall compact architecture of low-vigor rootstocks suited to high-fertility sites), showing complementary lateral development (shallow tap root potentially compensated by extensive lateral root network providing adequate total root system), and requiring appropriate site matching (shallow tap root necessitating rootstock selection appropriate for site conditions avoiding drought-prone or infertile situations where deep roots essential). Shallow tap root in vineyard management includes= requiring irrigation consideration (shallow tap root mandating irrigation system in any climate with significant summer drought period), necessitating fertigation (shallow tap root potentially benefiting from frequent fertilizer application through irrigation maintaining nutrient supply in limited root zone), affecting vine water relations (shallow tap root resulting in more variable plant water status compared to deep-rooted vines requiring closer monitoring), constraining site suitability (shallow tap root limiting rootstock appropriateness to sites with adequate shallow soil resources or irrigation), and influencing stress responses (shallow tap root increasing vine vulnerability to drought, nutrient deficiency, and competitive stress from weeds or cover crops).
Shallow_Tap_Root emphasizes limited depth less than 100 cm and is distinguished from Deep_Tap_Root by focus on shallow penetration. Critical for understanding Riparia genetics, soil limitations, upper soil resource exploitation, irrigation requirements, drought vulnerability, and the depth-limited vertical root restricted to upper soil horizons requiring appropriate site matching and management.
Begrenzte Vertikalwurzel
Flache Pfahlwurzel
Limited depth tap root
Oberflächennahe Hauptwurzel
Restricted tap root
Upper profile tap root
Shallow Tap Root
A shallow upper lateral sinus representing the right upper lateral sinus on a mature leaf at node 6 of Vitis vinifera cv. Chardonnay in Burgundy, France, exhibiting minimal depth of 1.4 cm measured from barely perceptible sinus base to the imaginary line connecting apical and upper lateral lobe edges, penetration reaching only approximately 28% of distance from blade margin toward median vein indicating very shallow dissection, poorly defined rounded configuration with sinus base barely distinguishable from surrounding blade contour, width of 2.2 cm at opening creating depth-to-width ratio of 0.64 indicating width exceeding depth, shallow sinus creating weak poorly defined separation between apical and upper lateral lobe regions producing integrated rather than isolated appearance, sinus base positioned 8.2 cm from petiolar junction reflecting the shallow penetration, sinus margins showing gradual transition from apical to lateral lobe regions without clear boundaries, shallow sinus contributing to characteristic weakly dissected Chardonnay architecture with poorly defined lobe structure, bilateral symmetry with left upper lateral sinus showing similarly shallow configuration (1.3 cm), measured at BBCH stage 75 (berries pea-sized) in late June, representing minimal shallow upper lateral sinus development typical of weakly lobed cultivars.
An upper lateral sinus exhibiting minimal depth creating weak separation between apical and upper lateral lobes in Vitis vinifera, characterized by depth typically less than 2 cm measured from sinus base to lobe edge connection line, penetration often only 20-40% of distance from blade margin toward median vein, poorly defined or barely perceptible sinus configuration, subtle indentation creating minimal visual prominence, occurrence primarily in weakly lobed cultivars showing reduced overall lobe development, and role in creating poorly differentiated apical regions characteristic of weakly dissected cultivar morphology. Shallow upper lateral sinuses represent minimal expression of upper sinus development, occurring in cultivars with weak lobe dissection including Chardonnay, some Gewürztraminer clones, weakly lobed Muscat types, and other poorly dissected varieties, creating distinctive leaf architecture with poorly separated or integrated apical-lateral lobe regions, and serving as diagnostic features indicating weakly lobed cultivar groups when combined with other morphological characteristics.
Shallow upper lateral sinuses are distinguished from deep upper lateral sinuses by depth typically <2 cm and penetration <40% toward median vein. The shallow designation emphasizes minimal dissection creating weak or absent lobe isolation contrasting with pronounced deep sinuses. Shallow depth creates poorly defined or barely perceptible separation making weakly lobed leaves appear almost entire (non-lobed) rather than showing distinct lobe architecture. Penetration percentage provides normalized assessment, with <30-40% indicating shallow dissection regardless of absolute depth values varying with blade size. Configuration is often poorly defined with gradual transitions rather than distinct sinus bases. Width often equals or exceeds depth creating broad shallow indentations contrasting with narrow deep sinuses. Occurrence predominates in weakly lobed cultivars, with deeply dissected varieties universally showing deep upper lateral sinuses. Correlation with overall lobe development is strong, with shallow upper sinus cultivars typically also showing shallow or absent lower sinuses and minimal overall dissection. Genetic control is strong with shallow dissection showing consistent expression. Environmental modification is limited with weakly lobed cultivars maintaining shallow sinuses across conditions, though extreme high vigor may occasionally produce slightly deeper sinuses. Functional implications include minimal lobe isolation with apical and lateral regions functioning as integrated structures rather than independent units. Ampelographic utility is substantial as diagnostic feature, with shallow upper lateral sinuses combined with other characters enabling identification of weakly lobed cultivar groups including major varieties like Chardonnay.
flache obere Seitenbucht
minimal upper lateral sinus
minimale obere Lateralbucht
schwach eingeschnittene Apikalsinus
slightly incised apical sinus
weak upper sinus
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 120-182 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 209-242.
Shallow Upper Lateral Sinus
Ampelographic identification of Gewürztraminer relies heavily on distinctive shape qualities including the characteristic orbicular leaf blade shape (width approximately equal to length, ratio 0.95-1.05), deeply overlapping petiolar sinus forming a lyre shape, and short ovoid to globose berry form, features that distinguish it from the closely related Savagnin which exhibits more elongated leaf blades (ratio 0.85-0.95) and ellipsoid berries despite near-identical genetic background.
Shape_Quality encompasses all morphological qualities that characterize the geometric form and spatial configuration of grapevine organs independent of their absolute size. Shape qualities describe how organ boundaries and surfaces are arranged in space, capturing the proportional relationships between dimensions, the curvature of surfaces, and the regularity or irregularity of form. This category is fundamental to ampelography because organ shapes, particularly of leaves, berries, and clusters, provide reliable cultivar identification characters that remain relatively stable across environmental conditions compared to size-dependent traits. Shape_Quality subdivides into five major domains: Overall_Form_Quality addressing the general three-dimensional geometry of organs Symmetry_Quality capturing the regularity of form around axes or planes Apex_Shape_Quality describing the terminal configuration of organs Base_Shape_Quality characterizing the proximal attachment region and Margin_Shape_Quality addressing the edge characteristics of laminar organs. Shape qualities can be assessed through categorical descriptors following OIV protocols, through quantitative shape indices derived from linear dimension ratios, or through advanced morphometric analysis using elliptic Fourier descriptors, landmark-based geometric morphometrics, or machine learning image classification. The genetic basis of organ shape involves complex developmental programs controlling cell division patterns, expansion anisotropy, and differential growth rates across organ regions. Environmental factors modulate shape expression within genetically determined limits, with light environment, temperature during organ initiation, and water status all influencing final form.
Shape_Quality is positioned as a direct subclass of Morphological_Quality, parallel to Size_Quality, Color_Quality, Texture_Quality, and Structural_Integrity_Quality. Shape qualities are dimensionless or expressed as ratios, distinguishing them from size qualities that have dimensional units. The same organ can be characterized by both size and shape qualities independently a berry can be small and globose or large and globose. Shape descriptors in OIV protocols provide standardized categorical assessments that can be linked to quantitative shape indices. PATO_0000052 (shape) provides the upper-ontology alignment.
Gestaltqualität
Konfigurationsqualität
configuration quality
form quality
geometric quality
geometrische Qualität
morphometric shape
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 47-91 Chitwood DH et al. (2014) A Modern Ampelography: A Genetic Basis for Leaf Shape and Venation Patterning in Grape. Plant Physiology 164(1): 259-272
Shape Quality
Riesling in Mosel shows shatter during mid June Year N+1 at BBCH 65-71, with heavy rainfall event during early fruit set causing 22-38% berry detachment through mechanical raindrop impact, flowers and young berries physically separated from fragile pedicels, immediate losses visible on vineyard floor, under cool humid climate with frequent June precipitation, typical shatter vulnerability requiring protective measures in severe weather years
Shatter represents weather-induced mechanical fruit drop when rain impact, hail damage, or strong wind physically detaches flowers or young berries from clusters during vulnerable flowering and early fruit set periods, causing immediate berry loss through mechanical separation rather than developmental abscission, and potentially reducing fruit set 10-40% in severe weather events. This weather-damage mechanism occurs during late spring to early summer in late May-July in temperate regions Year N+1, typically during flowering to 2-3 weeks post-flowering when attachment fragile, corresponding to BBCH 60-73 flowering to young fruit stages. Shatter mechanism involves mechanical impact from raindrops, hail stones, or wind forces exceeding attachment strength, physical separation of flowers or berries from pedicels through mechanical failure rather than abscission zone activation, immediate detachment and fall during or shortly after weather event, and berry loss visible as flowers or berries on ground below vines. Rain-induced shatter shows heavy rainfall during flowering or early fruit set physically dislodging flowers or young berries, raindrop impact forces exceeding weak attachment strength during vulnerable period, progressive berry loss during sustained rain events, and severe shatter possible 20-40% loss in torrential rain scenarios. Hail-induced shatter shows hailstone impact causing immediate flower or berry detachment, mechanical damage to clusters and shoots, severe losses possible 40-80% in direct hail events, and devastating effects during flowering creating near-total fruit loss in affected areas. Wind-induced shatter shows strong winds during flowering causing flower vibration and detachment, mechanical stress on fragile pedicel connections, progressive loss during sustained wind events, and wind effects exacerbating rain damage through combined mechanical forces. Attachment weakness during vulnerable period shows flowers and young berries having fragile pedicel connections before structural reinforcement, attachment strength progressively increasing as berries develop, critical vulnerability during flowering and first 2-3 weeks post-flowering, and vulnerability declining as pedicels strengthen and berries enlarge. Mechanical separation characteristics show physical breaking of pedicel tissue rather than organized abscission, irregular break surfaces distinguishing from clean abscission zone separation, no abscission zone involvement in immediate weather damage, and mechanical failure mechanism differing from developmental drop. Timing characteristics show shatter occurring during or immediately following weather event, instantaneous losses during severe weather, and timing corresponding to weather occurrence rather than developmental schedule. Shatter visibility shows flowers or berries immediately visible on ground or caught in vegetation, fresh green color indicating recent detachment, and obvious correlation with weather event timing. Cluster appearance after shatter shows gaps from missing berries, irregular pedicel breaks without abscission zone remnants, remaining flowers or berries potentially damaged, and overall disrupted cluster architecture. Berry development stage vulnerability shows flowers most vulnerable during anthesis to cap fall, young berries highly vulnerable first 1-2 weeks post-flowering, vulnerability declining weeks 3-4 as attachment strengthens, and mature berries relatively resistant though extreme hail can damage. Weather event severity effects show light rain causing minimal shatter <5%, heavy sustained rain causing moderate shatter 10-25%, torrential rain or hail causing severe shatter 30-50% or more, and event intensity as major shatter determinant. Cultivar variation in shatter susceptibility shows some cultivars having more fragile flower attachments increasing vulnerability, compact cluster architecture possibly increasing impact damage through berry crowding, and genetic differences affecting shatter resistance though less pronounced than environmental effects. Cluster architecture effects show compact clusters potentially experiencing greater shatter through impact transmission between berries, loose clusters showing more independent berry behavior, and architecture influencing shatter patterns. Geographic and temporal vulnerability shows regions with frequent flowering-period rain experiencing regular shatter risk, Mediterranean climates with dry flowering periods showing minimal shatter, and timing variability creating year-to-year shatter fluctuation. Prevention strategies show limited options due to weather unpredictability, protective structures like rain covers or hail nets providing physical protection where economically feasible, cultivar selection for favorable flowering timing avoiding typical storm periods, and overall reliance on favorable weather during vulnerable period. Shatter versus other drop distinction shows mechanical immediate loss distinguishing from progressive developmental drop, weather event correlation identifying shatter, fresh detachment evidence versus abscission zone formation, and timing during vulnerable period distinguishing from late drop. Shatter assessment shows immediate post-weather evaluation documenting berry loss, comparison of affected and protected areas quantifying impact, cluster counts determining loss percentage, and rapid assessment enabling damage quantification. Shatter impacts on yield show direct berry number reduction proportional to weather severity, severe events causing 30-50% losses affecting production economics substantially, year-to-year variability from weather creating yield unpredictability, and shatter as uncontrollable yield risk factor. Shatter recovery options show no recovery from lost berries with shatter losses permanent, remaining berries potentially compensating through larger size though limited compensation, and severe shatter potentially necessitating crop insurance claims. The process represents second type of fruit drop caused by weather damage, distinguished from developmental early drop and other mechanisms. Duration represents immediate loss during weather event rather than progressive drop. Regional timing shows shatter risk during late May-July across climates during flowering and early fruit set. Viticultural significance includes representing uncontrollable weather-induced yield loss, demonstrating critical vulnerability during flowering period, requiring protective infrastructure in high-risk areas, and contributing to production variability and risk. Research applications include quantifying weather impacts on fruit retention, evaluating cultivar and architectural effects on shatter susceptibility, developing protective strategies and technologies, and assessing shatter as production risk factor.
Shatter is second type of Fruit_Drop, distinguished by weather-induced mechanical detachment rather than developmental failure. Timing during flowering to 3 weeks post-flowering (BBCH 60-73). Caused by rain, hail, or wind impact. Immediate loss during weather event. Mechanical separation not abscission. Severity 5-50% depending on weather intensity. Compact clusters potentially more vulnerable. Uncontrollable yield risk. Limited prevention options. Differs from developmental Early_Fruit_Drop.
Regenschaden
hail shatter
mechanical berry detachment
mechanische Beerenablösung
rain damage
weather-induced drop
wetterbedingte Beerenfall
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 450-455
Shatter
Palomino Fino bears a Sherry_Style_Scion_Role as the dominant variety for all dry Sherry styles, its neutral character (low acidity, minimal varietal aroma) providing a blank canvas for flor-derived compounds (acetaldehyde 300-400 mg/L in fino, contributing apple/almond notes) and oxidative development (sotolon, nutty lactones in oloroso). Pedro Ximénez serves in a Sherry_Style_Scion_Role for intensely sweet PX Sherry, where sun-drying (asoleo) concentrates grapes to 40-50°Brix before light fermentation and fortification yields wines of 400-500 g/L residual sugar with treacle, fig, and coffee character. Muscat of Alexandria (Moscatel) bears a Sherry_Style_Scion_Role for aromatic sweet Sherries, retaining terpene character through fortification for floral Moscatel dessert wines.
A fortified wine scion role that inheres in a grafted scion cultivar by virtue of its designation for Sherry production in the Jerez-Xérès-Sherry DO, Spain, or Sherry-style wines elsewhere, where base wine undergoes complete fermentation before fortification to 15-15.5% (biologically aged styles) or 17-18% (oxidatively aged styles) alcohol. Sherry style scions are dominated by Palomino Fino (95% of plantings), a neutral variety whose lack of distinctive aromatics allows the biological (flor yeast) or oxidative aging processes to define wine character. Pedro Ximénez and Muscat of Alexandria serve for sweet Sherry production through sun-drying (asoleo) that concentrates sugars to 400-500 g/L before fermentation and fortification. Sherry production is unique in employing the solera system (fractional blending of multiple vintages), biological aging under flor (a film of Saccharomyces yeasts creating fino and manzanilla styles), and extended oxidative aging (oloroso, amontillado, palo cortado styles) that develops characteristic acetaldehyde, nutty, and dried fruit aromatics.
Sherry_Style_Scion_Role captures a fortified wine tradition fundamentally distinct from Port in its post-fermentation fortification and biological/oxidative aging dichotomy. The flor yeast system is unique to Jerez and a few other regions (Jura's vin jaune, Tokaj's szamorodni), creating wines impossible to replicate elsewhere despite using the same varieties. Sherry classification proceeds by biological aging potential: wines developing thick flor become fino/manzanilla, those losing flor become amontillado, and those fortified above flor-tolerance (17%+) become oloroso from the start. Sherry's 500-year tradition faces market challenges (declining consumption, vineyard abandonment), but quality renaissance movements emphasize single-vineyard, single-solera, and unblended expressions.
Jerez variety
Jerez-Traube
Sherry grape
Sherry variety
Sherry-Rebsorte
Sherrysorte
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Jeffs J (2016) Sherry 5th Ed, Infinite Ideas Jackson RS (2014) Wine Science 4th Ed, Academic Press pp.: 500-520 Consejo Regulador del Jerez-Xérès-Sherry (2020) Technical Manual
Sherry Style Scion Role
A shoot on Vitis vinifera cv. Cabernet Sauvignon in mid-June at BBCH stage 71 (fruit set completed), originating from primary bud at node 6 on bilateral cordon, currently 95cm in length with 14 fully expanded leaves, 3 developing leaves at apex, 2 inflorescences with fruit set complete showing 38 and 42 developing berries, 2 tendrils, active apical growth continuing at approximately 3-4cm per day, representing typical reproductive shoot development in vertical shoot positioning system with continued vegetative growth concurrent with early fruit development, requiring canopy management including shoot positioning, hedging, and potentially leaf removal to optimize fruit exposure and canopy microclimate.
A developing or mature stem segment of Vitis vinifera arising from bud breakage, characterized by elongating axis bearing nodes with attached leaves, tendrils, and potentially inflorescences, internodal regions undergoing primary and eventually secondary growth, and function as the primary photosynthetic and reproductive structural unit during the growing season, transitioning to mature lignified cane by dormant season providing fruiting wood for subsequent years. Shoots represent the fundamental growth unit in grapevine annual cycle, with shoot development proceeding through distinct phenological stages from initial bud swell (BBCH 01) through rapid elongation (BBCH 10-19) to flowering and fruit development (BBCH 50-89) and eventual growth cessation and lignification (late season), with shoot vigor, architecture, leaf area, and reproductive content determining vine productivity, fruit quality, and sustainability of the viticultural system.
Shoots are distinguished from canes (mature lignified previous-season shoots), water shoots or suckers (vigorous shoots from latent buds on permanent wood), and permanent structure (trunks, arms, cordons). Shoot classification systems recognize multiple hierarchies including developmental origin (primary, secondary, tertiary from compound buds), reproductive content (vegetative, reproductive, mixed), vigor (weak, moderate, vigorous), and timing (prompt, delayed, multiple-flush). Understanding shoot development and management is fundamental to viticulture, with shoot thinning, positioning, hedging, leaf removal, and other canopy interventions directly impacting yield, fruit quality, disease pressure, and long-term vine sustainability through effects on light penetration, air circulation, carbohydrate production, and bud differentiation for following seasons.
Jahrestrieb
Neutrieb
Trieb
annual shoot
current season growth
growing shoot
Based on Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 113-145 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Lorenz et al. (1995) "Growth Stages of the Grapevine= Phenological growth stages of the grapevine (Vitis vinifera L. ssp. vinifera)" Australian Journal of Grape and Wine Research 1=100-103.
Shoot
Cabernet Sauvignon one-year cane showing shoot-derived adventitious roots, cane section buried horizontally 5 cm depth in propagation bed, adventitious root primordia forming from nodal regions and internodal cambium without callus intermediary, microscopic examination revealing primordia originating from cambial and pericycle-like cells 8-10 days after burial, roots emerging directly through cortex and bark 14-18 days creating vascular connection to cane vascular system, buried cane producing 15 shoot-derived adventitious roots from 4 nodes enabling layering propagation
Shoot derived adventitious root is adventitious root originating directly from shoot tissues including stems, nodes, or internodes without intervening callus formation, characterized by initiation from preexisting cells in shoot vascular cambium, cortex, or pericycle-like regions, showing direct developmental connection to shoot vascular system, demonstrating formation typically in response to auxin accumulation, wounding, or environmental signals, and serving functions including providing additional root system for established vines, enabling vegetative propagation through stem cuttings or layering, and facilitating recovery from root damage through compensatory root production, with shoot-derived adventitious roots in grapevine showing typical origin from nodes where latent root primordia may preexist or from internodal cambial regions following induction, exhibiting development through organized stages of initiation, primordium formation, and emergence without callus intermediary creating direct shoot-to-root transition, and demonstrating that shoot origin provides vascular continuity potentially enabling more rapid functional integration than callus-derived roots where vascular connections must form de novo. Shoot derived adventitious root cellular origin includes= showing cambial origin (many shoot adventitious roots initiating from vascular cambium cells that dedifferentiate and reprogram), exhibiting pericycle-like origin (some roots originating from pericycle-like cells near phloem similar to lateral root origin), demonstrating cortical origin (occasional roots forming from cortical parenchyma cells particularly following wounding), revealing ray parenchyma involvement (vascular ray cells potentially serving as initiation sites particularly in woody stems), and displaying pluripotent cell activation (various shoot cell types capable of dedifferentiating and forming root primordia under appropriate signals). Shoot derived adventitious root anatomical pathway includes= showing endogenous initiation (primordia forming within shoot tissues rather than at surface), exhibiting outward growth (developing roots growing radially outward through cortex toward surface), demonstrating tissue disruption (emerging roots breaking through cortex and periderm creating wound in parent shoot), revealing vascular connection (developing root establishing direct connection to shoot vascular cylinder), and displaying emergence site (roots emerging at various points along shoot depending on initiation site distribution). Shoot derived adventitious root in nodes versus internodes includes= showing nodal preference (adventitious roots often forming preferentially at nodes where preformed primordia may exist and anatomy favorable), exhibiting internode capacity (internodal tissues also capable of forming adventitious roots though sometimes requiring stronger induction), demonstrating preformed primordia (some species including grapevine showing latent root primordia at nodes requiring only activation), revealing anatomical differences (nodal regions with different cell arrangements potentially facilitating root emergence), and displaying induction requirements (internodal rooting sometimes requiring higher auxin or more favorable conditions than nodal rooting). Shoot derived adventitious root hormonal induction includes= requiring auxin signal (shoot adventitious root formation strongly promoted by auxin accumulation or exogenous application), showing optimal auxin concentration (rooting showing concentration dependence with optimum typically 1000-10,000 ppm IBA for grapevine cuttings), exhibiting auxin transport (basipetal auxin transport from shoot apex potentially creating auxin accumulation promoting root formation), demonstrating cytokinin inhibition (high cytokinin levels potentially suppressing adventitious rooting creating hormone balance requirement), and revealing hormone interactions (auxin, ethylene, and other hormones interacting to determine rooting success and intensity). Shoot derived adventitious root environmental factors includes= showing moisture requirement (adequate moisture essential for root initiation and emergence with misting or humid conditions promoting rooting), exhibiting temperature effects (root formation optimal at 20-25°C declining at cooler or warmer temperatures), demonstrating light effects (darkness or low light often promoting rooting while bright light potentially inhibitory), revealing oxygen requirement (adequate aeration necessary for root metabolism with waterlogging potentially inhibiting despite moisture need), and displaying substrate effects (rooting medium physical and chemical properties affecting success). Shoot derived adventitious root in propagation includes= enabling cutting propagation (shoot adventitious roots allowing stem cuttings to develop functional root systems), providing layering mechanism (buried or mounded shoots forming adventitious roots enabling air or ground layering), demonstrating micropropagation (tissue culture shoots forming adventitious roots during acclimatization), revealing trench layering (traditional viticulture technique depending on shoot adventitious rooting of buried canes), and displaying stool bed production (rootstock mother plants producing shoots that form adventitious roots when mounded with soil). Shoot derived adventitious root cultivar variation includes= showing genetic differences (grapevine cultivars varying widely in shoot adventitious rooting capacity affecting propagation difficulty), exhibiting easy-rooting types (some cultivars and rootstocks like Riparia and many V. vinifera readily forming shoot adventitious roots), demonstrating difficult-rooting types (other varieties like some hybrids showing poor shoot adventitious rooting requiring special treatments), revealing selection importance (rootstock breeding selecting for superior adventitious rooting capacity improving propagation efficiency), and displaying practical implications (cultivar rooting capacity determining propagation methods and commercial feasibility). Shoot derived adventitious root functional development includes= showing rapid vascular integration (shoot-derived roots potentially achieving functional vascular connection more quickly than callus-derived), exhibiting complete root system development (shoot adventitious roots capable of forming entire functional root system with lateral branching), demonstrating absorption capacity (shoot adventitious roots developing root hairs and absorptive function supporting plant establishment), revealing secondary growth (shoot adventitious roots undergoing secondary thickening in perennial plants developing woody structure), and displaying long-term function (shoot adventitious roots persisting and functioning throughout plant life in vegetatively propagated vines).
Shoot_Derived_Adventitious_Root emphasizes direct shoot tissue origin without callus and is distinguished from callus-derived roots by focus on direct developmental pathway. Includes Stem_Derived_Adventitious_Root and Node_Derived_Adventitious_Root. Critical for understanding cambial origin, direct vascular connection, nodal vs internodal formation, hormonal induction, propagation methods, and the adventitious roots arising directly from shoot tissues enabling rapid functional integration.
Direct shoot root
Direkte Stammwurzel
Nicht-Kallus Adventivwurzel
Non-callus adventitious root
Spross-Adventivwurzel
Stem adventitious root
Shoot Derived Adventitious Root
The shoot description of Syrah documents long internodes (OIV 103:7, 80-110 mm on mid-shoot positions) with green ground color overlaid by conspicuous red-brown anthocyanin striping on the dorsal surface (OIV 006:3), a round to slightly flattened cross-section, and medium-density prostrate surface hairs — transitioning after véraison to a lignified cane with dark brown bark, prominent nodes, and the characteristic slightly drooping, semi-pendulous growth habit that distinguishes Syrah from the more rigidly erect shoots of Cabernet Sauvignon and the strongly drooping, fragile shoots of Grenache. The dormant cane color of Syrah is documented as medium brown with darker brown nodes (OIV 103:4), with a smooth bark surface showing fine longitudinal striations, distinguishing it from the pale straw-colored canes of Riesling (OIV 103:2) and the deep mahogany canes of Mourvèdre (OIV 103:6).
An ampelographic description component that documents the morphological characteristics of the growing-season shoot (green shoot from budburst to lignification) and the dormant cane (mature lignified shoot after leaf fall), recording internode length, diameter, color, cross-sectional profile, surface texture, and lignification pattern according to OIV descriptors 101-104 and supplementary characters, providing cultivar-diagnostic features of the axial vegetative structure that supports leaves, tendrils, and inflorescences.
Shoot_Description addresses the axial vegetative structure that is available for observation throughout the growing season (as green shoot from budburst through harvest) and during the dormant period (as lignified cane from leaf fall through winter pruning), providing year-round identification characters that complement the seasonally restricted leaf, cluster, and berry descriptors. The shoot/cane is particularly valuable for identification during dormancy — from leaf fall (November-December in northern hemisphere) through budburst (March-April), when no leaves, flowers, or fruit are available, the dormant cane is the only vegetative organ providing cultivar identification information: experienced viticultural workers can distinguish major cultivars by cane characteristics alone during winter pruning, using internode length, node prominence, bark color, bark texture, bud shape, and diaphragm visibility as the identification features. The developmental transition from green shoot to lignified cane involves progressive periderm formation (bark development), loss of green photosynthetic tissue, accumulation of brown phenolic pigments in the bark cells, and development of the protective cork layer — this lignification process produces the cultivar-characteristic cane color and texture that are scored on the dormant organ during winter.
Achsentrieb-Morphologie
Ruten-Charakterisierung
Trieb-Beschreibung
axial growth documentation
cane characterization
shoot morphology description
Shoot Description
Shoot development rate assessment comparing vigor levels in Syrah reveals the balance implications of vegetative growth rate, with high-vigor vines showing rapid shoot development (8.5 cm/day during peak growth, 180 cm total shoot length by flowering) creating dense, shaded canopies requiring extensive summer pruning, while balanced vines show moderate rate (5.2 cm/day, 95 cm shoot length by flowering) achieving appropriate canopy density with minimal intervention, demonstrating how shoot development rate affects the management requirements and fruit quality potential through canopy architecture effects.
Shoot development rate quality is a developmental rate quality that represents the speed of shoot elongation and leaf emergence following budbreak, characterizing the rate of vegetative canopy establishment that determines how quickly the photosynthetic apparatus develops to support fruit growth and how much vegetative development occurs before flowering shifts vine priority toward reproduction. Within the BFO framework, this quality specifically addresses the shoot growth dimension of phenological rate, distinguishing vegetative development rate from bud and reproductive development while recognizing that shoot development rate during the pre-flowering period establishes the leaf area available to support fruit development and determines vine vigor expression that affects canopy density and fruit-zone microclimate. The ontological significance lies in shoot development rate determining the balance between vegetative and reproductive development, where rapid shoot growth produces extensive canopy potentially supporting high yields but also creating shading and competition with fruit, while moderate shoot development produces balanced canopies with appropriate fruit exposure, making shoot development rate a key factor in the vegetative-reproductive balance affecting fruit quality. The temperature and resource basis for shoot development rate shows strong positive relationships with temperature (rate increasing approximately linearly above base temperature of 10°C) and modification by water and nutrient availability (resource limitation reducing rate regardless of temperature), with the integrated effect determining actual shoot elongation rates typically ranging from 2-10 cm/day during active growth depending on conditions. The growth stage relationship shows shoot development rate varying through the season, with acceleration during the exponential growth phase post-budbreak, peak rates during the 4-8 week period before flowering, and declining rates as flowering approaches and resources shift toward reproductive development, with this pattern modified by cultivar genetics and vine balance status. The management implications of shoot development rate include the timing of shoot thinning and positioning operations (requiring appropriate developmental stage for effective intervention), the assessment of vine vigor (rapid development indicating high vigor potentially requiring canopy management), and the prediction of canopy density at flowering affecting spray penetration and disease pressure. Understanding shoot development rate quality enables prediction of canopy development, timing of vegetative management operations, and assessment of vine vigor and balance affecting fruit quality outcomes.
Shoot_Development_Rate_Quality addresses the speed of vegetative canopy establishment affecting vine balance and canopy architecture. This quality responds strongly to temperature and resource availability. The class supports canopy management timing and vigor assessment through shoot development rate characterization.
Kronenentwicklungsrate
Triebentwicklungsrate
Triebwachstumsrate
canopy development rate
shoot elongation rate
shoot growth velocity
vegetative Wachstumsrate
vegetative growth rate
Lebon E et al. (2004) Shoot development in grapevine (Vitis vinifera) is affected by the modular branching pattern of the stem and intra- and inter-shoot trophic competition. Annals of Botany 93(3): 263-274 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 59-144
Shoot Development Rate Quality
Pinot Noir in Burgundy shows shoot elongation from late April through early August over 14-15 weeks, with shoots extending from 120 mm at 4-5 leaves to 900-1400 mm final length through elongation rate progression from 8-12 mm per day early phase through 25-40 mm per day peak mid-June declining to cessation late July-early August, under continental climate conditions of 15-25°C temperature range, variable precipitation requiring supplemental irrigation, and moderate fertility typical of this moderate-vigor early-season cultivar
Shoot elongation represents the specific growth process where shoot length increases through coordinated internode extension and apical meristem activity, creating visible daily shoot growth and establishing shoot architecture through spatial arrangement of leaves, tendrils, and reproductive structures. This fundamental developmental process constitutes the primary mechanism of shoot growth, occurring continuously from early development through growth cessation over 8-16 week period depending on vigor and conditions. Elongation results from cell production in apical meristem initiating new cells for shoot axis and organ primordia, intercalary meristem activity in developing internodes adding cells to internode tissues, and cell expansion in subapical elongation zones where cells increase length dramatically through turgor-driven wall extension. The apical meristem maintains dome structure typically 100-200 micrometers diameter with organized cell layers including outer tunica producing surface tissues and inner corpus generating internal tissues, with cell division rates during active growth producing new cells every 18-24 hours maintaining meristem size while contributing cells to developing organs and shoot axis. Subapical elongation zone extends 10-30 mm behind apex depending on growth rate, containing cells undergoing rapid expansion through water uptake creating turgor pressure of 0.4-0.8 MPa driving cell wall extension, with expansion rate highest immediately behind meristem declining toward mature tissues as cell walls rigidify. Internode extension contributes most directly to visible shoot elongation with developing internodes elongating through intercalary meristem activity near nodes and general cell expansion throughout internode length, creating internode lengths varying from 20-40 mm during slow growth to 80-150 mm during rapid growth. Cell expansion involves water uptake increasing cell volume 20-100 fold from meristematic dimensions, wall loosening through expansin proteins and other wall-modifying enzymes allowing plastic deformation under turgor pressure, and wall synthesis adding new material maintaining structural integrity while allowing continued expansion. Elongation rate shows characteristic temporal pattern following sigmoid curve with initial slow rate during early shoot establishment at 5-15 mm per day, accelerating to rapid exponential growth at 20-40 mm per day as photosynthetic capacity develops, reaching peak linear growth at 30-50 mm per day during optimal mid-season conditions, then declining to 10-20 mm per day as limitations develop, and eventually ceasing as growth terminates. Daily elongation rate varies substantially with environmental conditions particularly temperature driving metabolic rates and water availability maintaining turgor, with favorable conditions supporting maximum rates while stress rapidly reduces or arrests elongation. Vascular differentiation proceeds continuously in elongating tissues with procambial strands differentiating into xylem vessels for water transport and phloem sieve elements for assimilate distribution, establishing functional connections between developing leaves and permanent structures. Cell wall composition changes during elongation with initial thin plastic walls rich in pectins allowing expansion, transitioning toward walls with increased cellulose and hemicellulose providing structural rigidity as expansion completes, determining final cell and tissue mechanical properties. Metabolic activity remains elevated in elongation zones with respiration rates 8-15 fold above mature tissue levels supporting growth processes, consuming both stored reserves during early growth and photosynthate from expanding leaves as photosynthetic capacity develops. Protein synthesis operates at high rates producing structural proteins for expanding cells including cell wall proteins and membrane proteins, enzymes supporting metabolism including wall-modifying enzymes enabling expansion, and regulatory proteins coordinating developmental processes. Hormonal regulation centrally involves auxin from shoot apex maintaining elongation through promoting cell expansion and coordinating growth polarity, gibberellins supporting internode elongation particularly through promoting cell expansion, cytokinins from roots promoting cell division in meristems, and brassinosteroids enhancing cell expansion and vascular differentiation. Temperature strongly affects elongation rate with optimal range 22-28°C for maximum rate approximately doubling per 10°C increase within 15-30°C physiological range, with temperatures above 32°C reducing efficiency through excessive respiration and water stress while temperatures below 15°C substantially slow all processes. Light quality affects elongation pattern with red: far-red ratio influencing internode length through phytochrome-mediated responses, with shade conditions promoting elongated internodes as etiolation response while high light supports shorter stouter internodes. Water availability critically affects elongation through direct effects on cell turgor driving expansion and indirect effects on photosynthesis supplying assimilates and energy, with water stress rapidly arresting elongation through turgor loss and can prematurely terminate growth affecting final shoot size. Nutrient availability particularly nitrogen affects elongation rate and duration through effects on protein synthesis supporting growth and photosynthesis providing carbon skeletons and energy, with adequate nutrition supporting vigorous sustained elongation while deficiency reduces rate and causes early cessation. The process shows five distinguishable phases based on elongation rate trajectory, progressing from early elongation during establishment through rapid exponential growth, peak linear growth during optimal conditions, declining growth as limitations engage, and terminal phase as elongation ceases. Cultivar variation appears in elongation rate patterns, maximum elongation rates during peak growth, total elongation duration determining final shoot length, internode length characteristics affecting shoot architecture, and environmental responsiveness in growth patterns. Regional timing shows active elongation from May through July-August in warm climates, late May through August-September in temperate regions, and June through September in cool areas, with duration varying based on temperature accumulation and water availability. Viticultural management considerations include providing adequate resources supporting desired elongation patterns, managing vigor through appropriate irrigation and nutrition balancing vegetative and reproductive development, implementing shoot positioning optimizing canopy light interception, timing shoot trimming operations controlling excessive elongation and directing resources toward fruit, and monitoring elongation rates assessing vine health and management effectiveness. Excessive elongation creates overly vigorous canopies with dense shade reducing fruit quality while insufficient elongation limits photosynthetic capacity reducing productivity. Research applications include analyzing cellular and molecular mechanisms of elongation processes, quantifying environmental and hormonal effects on elongation rates and patterns, modeling elongation responses for management decision support, evaluating genetic variation in elongation characteristics affecting architecture and productivity, and assessing climate change impacts on elongation patterns and final canopy development.
Shoot_Elongation represents primary mechanism of Shoot_Growth through internode extension and apical activity. Distinguished by focus on length increase rather than broader growth encompassing leaf development and lignification. Subdivided into Early, Rapid, Peak, Declining, and Terminal phases based on elongation rate trajectory. Critical process determining final shoot length and canopy architecture.
Triebverlängerung
shoot extension
shoot stretching
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 205-235 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 128-152 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Shoot Elongation
Shoot freezing tolerance quality dynamics during autumn hardening were characterized in Cabernet Franc to establish the timeline of acclimation development and vulnerability windows during the growing-to-dormant transition. Controlled freezing tests at 2-week intervals from September through December quantified the progressive development of shoot freezing tolerance quality. Active shoot tips in early September showed shoot freezing tolerance quality with LT50 at only -3°C, representing the extreme vulnerability of still-growing tissue. By early October, basal shoot regions showed improved shoot freezing tolerance quality with LT50 at -12°C while distal regions remained at -6°C, reflecting the acropetal maturation gradient. Full shoot freezing tolerance quality developed by late November with LT50 reaching -22°C for basal cane regions, -18°C for mid-cane positions, and -15°C for distal regions that had completed growth later. The shoot freezing tolerance quality characterization established that the October-November window represented the highest vulnerability period when daytime temperatures might permit growth continuation while night temperatures could fall to levels exceeding the incomplete hardiness of partially acclimated tissues, informing recommendations for practices that accelerate hardening including shoot topping and irrigation cessation in early autumn.
Shoot freezing tolerance quality is a tissue freezing tolerance quality that inheres in grapevine shoots and represents the capacity of young, actively growing shoots during the growing season and mature but not fully dormant shoots during the transition periods to survive freezing temperatures, with tolerance varying from essentially zero in actively growing spring shoots through limited tolerance during autumn hardening to the substantial tolerance of fully lignified dormant canes that have completed acclimation. This quality addresses the cold vulnerability of shoot tissues across their developmental trajectory, recognizing that the same physical structure shows dramatically different cold sensitivity depending on its developmental and acclimation status, with actively growing shoots representing the most cold-vulnerable stage and fully hardened dormant canes representing the most tolerant state of shoot tissue. The ontological significance of shoot freezing tolerance quality lies in its relationship to both canopy development and vine structural integrity, with spring frost damage to emerging shoots requiring replacement from secondary buds and affecting current-season productivity, while winter damage to canes affects the availability of fruiting positions for the following season and may cause structural losses requiring retraining from trunk buds or replacement from new growth. The biological basis of shoot freezing tolerance encompasses the progressive development of cold tolerance as shoots mature from the highly freeze-sensitive actively growing state through lignification and periderm development that provides structural barriers to ice propagation, combined with the physiological acclimation processes of dehydration, cryoprotectant accumulation, and membrane modification that collectively enable survival of extracellular ice formation in the tissues of dormant canes. The gradient of freezing tolerance along the length of a shoot reflects the acropetal progression of maturation, with basal regions achieving greater tolerance earlier in the season than distal regions that remain actively growing longer, creating a characteristic pattern of distal damage in marginal autumn or spring frost events where basal portions survive while tips are killed.
Shoot_Freezing_Tolerance_Quality is_a Tissue_Freezing_Tolerance_Quality, representing the cold hardiness of shoot tissues from active growth through dormancy. This class addresses the developmental trajectory from high vulnerability (active, -3°C LT50) to substantial tolerance (dormant, -22°C LT50). The acropetal maturation gradient creates position-dependent hardiness along shoot length. Transition periods (spring, autumn) represent vulnerability windows with incomplete hardiness. Extracellular freezing tolerance mechanism in lignified cane contrasts with supercooling in buds. Distinguished from Bud_Freezing_Tolerance_Quality by focus on cane tissues rather than meristematic buds.
Ruten-Kältehärte
Ruten-Kältetoleranz
Trieb-Gefrier-Toleranz
Triebfrostresistenz
cane cold tolerance
cane freezing tolerance
vegetative shoot frost resistance
Shoot Freezing Tolerance Quality
Detailed characterization of Shoot_Frost_Recovery_Disposition at Virginia Tech employed controlled spring frost simulations to assess recovery parameters across 20 cultivars. Primary shoots at the 3-5 leaf stage were exposed to -3.5°C for 90 minutes (conditions causing > 90% primary shoot mortality), and recovery was tracked over 30 days, recording secondary bud activation timing, secondary shoot vigor, and cluster count on replacement shoots. Shoot_Frost_Recovery_Disposition varied substantially among cultivars. 'Merlot' demonstrated exceptional recovery with secondary bud activation visible within 6 days, secondary shoot elongation rate of 3.2 cm/day (exceeding typical primary shoot rate), and 0.85 clusters per secondary shoot, enabling 60% crop recovery. 'Cabernet Franc' showed similar rapid activation (7 days) but lower secondary fruitfulness (0.55 clusters/shoot), yielding 40% recovery. 'Chardonnay' exhibited poor Shoot_Frost_Recovery_Disposition with delayed activation (12 days), slow secondary shoot growth (1.8 cm/day), and minimal fruitfulness (0.25 clusters/shoot), achieving only 18% crop recovery. Notably, secondary shoot cluster size was 20-35% smaller than primary shoot clusters across all cultivars, indicating that even with high secondary fruitfulness, full crop recovery is not achievable. Field validation following a natural April frost confirmed laboratory rankings, with 'Merlot' blocks recovering 55% of typical yield versus 22% for 'Chardonnay' under equivalent damage conditions.
A Shoot_Frost_Recovery_Disposition is a Spring_Frost_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to regenerate productive shoots following frost damage to primary growing points, restoring canopy function and at least partial crop production through activation of secondary buds, latent buds, or basal shoots. This disposition represents the recovery component of spring frost adaptation, determining outcomes after frost damage has occurred rather than preventing initial damage. Recovery capacity depends on the availability and fruitfulness of backup growing points (secondary and tertiary buds if present, latent buds on older wood), the speed of their activation following primary shoot death, and the fertility of replacement shoots. Cultivars differ substantially in recovery capacity based on secondary bud fruitfulness (ranging from 0 to 1.5 clusters per shoot among cultivars), the proportion of secondary buds surviving frost events (often higher than primary shoots due to delayed development), and the vigor of replacement shoot growth. Effective Shoot_Frost_Recovery_Disposition enables economically meaningful crop production even in years with significant frost damage, reducing the year-to-year production variability that threatens vineyard sustainability in frost-prone regions. Recovery capacity is particularly valuable when combined with partial tissue tolerance, allowing damaged primary shoots to potentially recover through lateral growth while secondary buds provide insurance against complete shoot loss.
Shoot_Frost_Recovery_Disposition is_a Spring_Frost_Tolerance_Disposition. Recovery capacity determines outcomes after frost damage occurs, complementing avoidance and tolerance mechanisms. Key parameters include secondary bud activation speed, secondary shoot vigor, and secondary bud fruitfulness. Recovery is rarely complete, as secondary shoots typically produce smaller clusters than primary shoots. Strong recovery capacity is particularly valuable when frost timing is unpredictable, as avoidance through late budbreak cannot be relied upon.
Frostschadens-Regeneration
Trieb-Frost-Erholungs-Disposition
Trieb-Wiederaustrieb
post-frost regeneration
shoot regrowth potential
Friend AP et al. (2011) Delayed winter pruning and spring frost damage to grapevine buds, Australian Journal of Grape and Wine Research 17:214-218 Dry PR (2000) Canopy management for fruitfulness, Australian Journal of Grape and Wine Research 6:109-115 Trought MCT et al. (1999) Vineyard variability in Marlborough, New Zealand, American Journal of Enology and Viticulture 50:484-489
Shoot Frost Recovery Disposition
Cabernet Sauvignon in Napa Valley shows shoot growth from early May through late July spanning BBCH 15-35 over 12-14 weeks, with shoots elongating from 150 mm at 5 leaves unfolded to 1200-1800 mm final length through growth phases progressing from 10-15 mm per day early elongation through 30-50 mm per day peak growth declining to cessation, under conditions of 18-28°C temperature range, adequate irrigation maintaining soil moisture, and moderate nitrogen fertility supporting balanced vegetative-reproductive development typical of this vigorous late-season cultivar in warm climate
Shoot growth represents the sustained vegetative development phase where shoots elongate through coordinated apical meristem activity, internode extension, and sequential leaf production, establishing productive canopy architecture and accumulating photosynthetic capacity for supporting reproductive development and reserve accumulation. This fundamental developmental process follows bud burst completion and continues throughout the growing season, typically spanning 8-16 weeks depending on cultivar vigor, environmental conditions, and management practices. Shoot growth encompasses internode elongation creating visible daily shoot extension of 10-50 mm during peak growth periods, continued leaf initiation and expansion adding photosynthetic area, tendril and inflorescence development establishing reproductive structures, and progressive shoot maturation through lignification and reserve accumulation. The process shows characteristic sigmoid growth pattern with initial slow growth during early shoot establishment transitioning to rapid exponential growth as photosynthetic capacity develops, reaching maximum linear growth rate during mid-season optimal conditions, then declining as environmental limitations, resource competition, or developmental programming reduce growth rate, and eventually terminating as shoot apex ceases meristematic activity. Shoot elongation results from cell division in apical and intercalary meristems producing new cells, and cell expansion in subapical zones where cells increase length 20-100 fold through turgor-driven wall extension creating visible shoot extension. The apical meristem maintains organized structure with tunica and corpus cell layers producing organs in regular phyllotactic pattern, initiating new leaf primordia every 2-3 days during active growth, with primordia developing through predictable sequence from multicellular initiation to recognizable leaf form requiring 10-15 days before emergence. Internodes between successive leaves extend through intercalary meristem activity and cell expansion creating shoot length increments, with internode length varying from 20-40 mm during slow growth to 80-150 mm during rapid growth depending on vigor, environment, and developmental stage. Leaves continue unfolding and expanding sequentially with each shoot typically producing 12-20 leaves during growth period, with leaf size generally increasing from shoot base toward middle nodes then declining toward apex reflecting changing resource availability and developmental stage. Photosynthetic capacity increases progressively as leaf area accumulates, with shoots transitioning from net carbon consumers during early development to net carbon producers once 3-4 leaves fully expand, ultimately achieving positive carbon balance supporting both shoot growth and contribution to whole-plant assimilate pool. Carbon allocation shifts during shoot growth from exclusive vegetative growth during early phases to increasingly supporting reproductive development and reserve accumulation as fruit load develops and growth rate declines. Water demand increases substantially during shoot growth as transpiring leaf area expands and metabolic activity remains high, with vigorous shoots consuming 200-500 mg water per hour during peak growth depending on environmental conditions and canopy development. Vascular system extends continuously through elongating shoots with xylem differentiation maintaining functional water transport and phloem development enabling bidirectional resource movement including water and minerals upward and photosynthate and hormones both directions. Nutrient requirements intensify during rapid shoot growth particularly for nitrogen supporting protein synthesis in expanding tissues, phosphorus supporting energy metabolism and nucleic acid synthesis, and potassium supporting cell expansion and enzyme activation. Shoot lignification begins in basal internodes during active growth, progressing acropetally as growth rate declines with lignified tissues providing mechanical support, creating perennial woody structures for subsequent seasons, and accumulating carbohydrate reserves in parenchyma cells. Hormonal regulation involves auxin from shoot apex maintaining apical dominance and coordinating growth polarity, cytokinins promoting cell division and delaying senescence, gibberellins supporting cell expansion and internode elongation, and ethylene and abscisic acid eventually contributing to growth cessation and maturation. Temperature strongly affects growth rate with optimal range 20-28°C for maximum elongation, with development rate approximately doubling per 10°C increase within physiological range while temperatures above 32°C or below 12°C substantially reduce growth. Light intensity affects growth rate and pattern with high light supporting rapid growth and shorter internodes with thicker shoots, while shade promotes elongated internodes with thinner shoots as etiolation response seeking improved light exposure. Water availability critically affects growth rate and duration with adequate soil moisture essential for maintaining turgor supporting cell expansion, while water stress rapidly arrests growth and can prematurely terminate shoot development affecting final canopy size and productivity. Nutrient availability particularly nitrogen strongly affects growth rate, duration, and final shoot size with adequate nutrition supporting vigorous extended growth while deficiency reduces growth rate and causes early growth cessation limiting canopy development. The process shows five distinguishable phases based on growth rate progression, advancing from early shoot elongation during canopy establishment through rapid exponential growth, reaching peak linear growth during optimal mid-season conditions, declining as limitations develop or developmental programming engages, and terminating as apical meristem ceases activity. Cultivar variation appears in growth habit including vigor determining final shoot length, internode length patterns affecting architecture, growth duration from short 8-10 week to extended 14-16 week periods, and responsiveness to environmental conditions and management. Regional timing shows shoot growth active from late April through July-August in warm climates, May through August-September in temperate regions, and June through September in cool areas, with duration varying based on temperature accumulation, water availability, and cultivar characteristics. Viticultural management considerations include providing adequate water and nutrition supporting desired growth, implementing shoot positioning and canopy management optimizing light interception and air movement, controlling excessive vigor through appropriate crop load and root zone management, timing shoot trimming operations managing canopy size and directing resources toward fruit development, and monitoring growth uniformity assessing vineyard conditions and management effectiveness. Excessive shoot growth creates dense shaded canopy reducing fruit quality through poor light exposure and air movement increasing disease pressure, while insufficient growth limits photosynthetic capacity reducing fruit development and reserve accumulation affecting current and subsequent season productivity. Research applications include quantifying environmental and management effects on growth patterns and final canopy characteristics, analyzing carbon allocation between vegetative and reproductive growth affecting productivity and quality, evaluating hormonal and genetic regulation of growth habit and architecture, modeling growth responses to environmental conditions for management decision support, and assessing climate change impacts on growth patterns across viticultural regions.
Shoot_Growth represents sustained vegetative development phase following Bud_Burst and First_Leaf_Unfolding. Distinguished by active internode elongation and sequential leaf production. Encompasses Shoot_Elongation as major constituent process along with Internode_Development, Leaf_Emergence, Tendril_Formation, and Shoot_Lignification. Transitions eventually to reproductive development focus and growth cessation. Critical phase establishing canopy architecture and photosynthetic capacity.
Bestandeswachstum
Triebentwicklung
Triebwachstum
canopy growth
shoot development
vegetative growth
vegetatives Wachstum
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 205-265 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 128-178 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200 Schultz HR (2000) Climate change and viticulture: A European perspective on climatology, carbon dioxide and UV-B effects, Aust J Grape Wine Res 6:2-12
Shoot Growth
Precision irrigation management in a California Chardonnay vineyard utilizes dendrometer-mounted shoot growth sensors recording Shoot_Growth_Rate_Quality at hourly intervals, with algorithms triggering irrigation when the 7-day rolling average growth rate declines below 1.5 cm/day post-véraison, maintaining sufficient growth for canopy function while imposing the moderate water stress required for optimal fruit composition and flavor development.
Shoot_Growth_Rate_Quality is a Vegetative_Growth_Vigor_Quality that inheres in a grapevine (Vitis vinifera L.) and represents the velocity of primary shoot extension as measured in length increment per unit time during the active growth phase. This quality provides the most direct and temporally sensitive indicator of vine vigor status, responding rapidly to changes in water availability, temperature, nutrient supply, and hormonal balance. In the BFO ontological framework, Shoot_Growth_Rate_Quality exists as a rate-dependent quality whose measurement requires temporal sampling, distinguishing it from static measurements like internode length or total shoot length that capture growth outcomes rather than growth dynamics. Grapevine shoot growth follows a characteristic sigmoidal pattern, with minimal extension during early bud swell (BBCH 01-07), rapid acceleration through the pre-flowering phase (BBCH 12-57) reaching peak rates of 3-6 cm per day under optimal conditions, followed by gradual deceleration as competition from developing fruit and environmental stress factors induce growth cessation. The physiological basis for shoot growth rate involves cell division in the shoot apical meristem combined with cell elongation in the subapical zone, processes highly sensitive to turgor pressure, gibberellin concentration, and cytokinin-auxin balance. Water stress reduces shoot growth rate before affecting any other physiological process, making growth rate monitoring the earliest indicator of developing water deficit — growth reduction occurs at leaf water potentials of -0.6 to -0.8 MPa, well before photosynthetic inhibition or visible stress symptoms. This sensitivity underlies the use of shoot growth cessation as a target for regulated deficit irrigation programs, where deliberate mild water stress is imposed post-fruit set to limit vegetative growth while maintaining photosynthetic capacity for fruit ripening. Temperature strongly modulates growth rate, with minimal extension below 10°C, optimal rates between 25-30°C, and growth inhibition above 35°C, creating the characteristic diurnal pattern of maximal extension during warm nights when carbohydrate supply is not limited by current photosynthesis.
Shoot_Growth_Rate_Quality represents a rate measurement (length/time) rather than a static dimensional quality, positioning it as the most dynamic indicator within the vigor hierarchy. The class relates functionally to water status, temperature response, and hormonal regulation, making it central to precision viticulture approaches. Measurement methods range from simple ruler-based weekly assessments to continuous automated monitoring with linear displacement sensors, with temporal resolution affecting the management insights obtainable.
Streckungswachstumsrate
Triebwachstumsgeschwindigkeit
Triebzuwachsrate
daily growth increment
shoot extension rate
Lebon E, Pellegrino A, Louarn G, Lecoeur J (2006) Branch development controls leaf area dynamics in grapevine (Vitis vinifera) growing in drying soil. Annals of Botany 98:175-185 Williams LE, Ayars JE (2005) Grapevine water use and the crop coefficient are linear functions of the shaded area measured beneath the canopy. Agricultural and Forest Meteorology 132:201-211
Shoot Growth Rate Quality
In Chardonnay vines in the Mâconnais, shoot growth termination occurs during the first half of October when shoot elongation rate declines to zero, with the shoot apical meristem becoming quiescent and shoot tips hardening from green to brown coloration, achieving complete termination approximately 3 weeks after mean daily temperatures consistently fall below 12°C, corresponding to the transition from BBCH 89 to BBCH 91.
Shoot growth termination in Vitis vinifera represents the complete cessation of shoot apical meristem activity and internode elongation, marking the end of the vegetative extension growth that characterized the vine during the growing season. This process typically occurs during October in Northern Hemisphere viticulture as the accumulated effects of short-day signaling and declining temperatures finally overcome any remaining growth potential in shoot meristems. The termination of shoot growth involves the arrest of cell division in the shoot apical meristem, cessation of cell elongation in the most recently formed internodes, and either the formation of a terminal resting bud or the abscission of the immature shoot tip depending on cultivar characteristics and environmental conditions. Prior to complete termination, shoot elongation rate progressively declines from peak values of 3-5 centimeters per day during rapid spring growth to less than 0.5 centimeters per day during late summer, eventually reaching zero as the terminal meristem becomes quiescent. The morphological changes accompanying shoot growth termination include hardening and browning of the shoot tip region as tissues that would have continued elongating instead undergo lignification and periderm development, cessation of new leaf primordium initiation at the shoot apex, and maturation of the most distal leaves that have already emerged but would not have fully expanded under continued growth conditions. Hormonal regulation of shoot growth termination involves the convergence of declining auxin transport from the shoot apex, reduced gibberellin biosynthesis and signaling, and increased abscisic acid accumulation that collectively create conditions incompatible with continued meristematic activity. The timing of shoot growth termination relative to other phenological events significantly affects vine performance, with termination before harvest allowing redirection of carbohydrates to fruit ripening and reserve accumulation, while very early termination may indicate stress conditions that prematurely limited growth, and very late termination may reflect excessive vigor or delayed dormancy induction that compromises cold hardiness. Cultivar differences in shoot growth termination timing range over several weeks, with determinate growth habit cultivars terminating earlier than indeterminate types under equivalent conditions.
Shoot_Growth_Termination represents the shoot-specific component of Growth_Cessation, specifically addressing shoot apical meristem arrest and elongation cessation. Distinguished from Cambial_Activity_Decline which addresses secondary growth, and from the earlier Growth_Inhibition_Initiation which marks beginning rather than completion of growth decline. Timing varies with cultivar growth habit from determinate to indeterminate. Complete termination prerequisite for proper shoot maturation and cold hardiness. Duration of terminal phase typically spans 2-3 weeks.
Triebstreckungsstopp
Triebverlängerungs-Ende
Triebwachstums-Beendigung
apical growth termination
apikale Wachstumsbeendigung
shoot elongation cessation
shoot extension termination
vegetative shoot growth end
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Pratt C (1974) Vegetative anatomy of cultivated grapes - a review. American Journal of Enology and Viticulture 25: 131-150 Mullins MG, Bouquet A, Williams LE (1992) Biology of the Grapevine Cambridge University Press
Shoot Growth Termination
Shoot hydraulic conductance tracking through the growing season in Pinot Noir reveals developmental changes affecting canopy water supply, with early season (30 cm shoots) showing whole-shoot conductance of 2.8 mmol/s/MPa (immature xylem limiting supply), midsummer (90 cm shoots) increasing to 8.5 mmol/s/MPa (mature, functional xylem), and late season declining to 5.2 mmol/s/MPa (developing tyloses, early senescence signals), with the seasonal conductance trajectory determining the hydraulic supply capacity available to support transpiring leaves and ripening fruit at different phenological stages.
Shoot hydraulic conductance quality is a stem hydraulic conductivity quality that represents the integrated water transport capacity from the base of current-season shoots to the terminal growing points and leaves, characterizing the green stem component of hydraulic architecture that supplies the actively transpiring and growing canopy and that shows substantial developmental changes through the growing season as shoots elongate and mature. Within the BFO framework, this quality specifically addresses the current-season shoot component of the transport pathway, distinguishing green shoot conductance from perennial woody stem conductivity while recognizing that shoot hydraulic properties affect the water supply to the actively growing canopy and determine how effectively new growth can be sustained under varying water availability. The ontological significance lies in shoot hydraulic conductance determining the water supply capacity to the year's photosynthetic and reproductive investment, where the developing shoot must progressively supply increasing leaf area and fruit load while its own xylem continues to mature, creating a dynamic hydraulic system whose capacity changes through the season in ways that affect canopy water relations and fruit development. The developmental trajectory of shoot hydraulic conductance shows initial limitation in young elongating shoots (immature xylem with narrow, incompletely differentiated vessels), progressive increase as vessels mature and secondary xylem develops (typically peaking at midsummer), and potential late-season limitation if drought, excessive crop load, or early senescence compromises hydraulic function. The anatomical and physiological determinants of shoot conductance include vessel diameter in shoot xylem (narrower than perennial stems, creating relatively high resistance), shoot length (longer shoots having higher total resistance), and the functional state of vessels (developmental maturity, embolism status, tylosis formation), with these factors varying along the shoot and among shoots of different vigor. The canopy management implications of shoot hydraulic conductance are substantial, as excessively long shoots may experience hydraulic limitation affecting distal leaf and fruit function, while shoot positioning and training can affect the hydraulic efficiency of water supply to different canopy zones. Understanding shoot hydraulic conductance quality enables assessment of water supply to the developing canopy, interpretation of shoot position effects on leaf and fruit water status, and evaluation of canopy management effects on hydraulic architecture supporting current-season growth.
Shoot_Hydraulic_Conductance_Quality addresses the current-season green stem component of hydraulic architecture supplying active canopy growth. This quality shows developmental changes affecting water supply to leaves and fruit throughout the season. The class supports canopy water relations assessment and interpretation of seasonal hydraulic capacity changes through shoot conductance characterization.
Grüntrieb-Leitfähigkeit
Schossleitfähigkeit
Triebhydraulikleitfähigkeit
aktuelle Triebleitfähigkeit
cane conductance
current-season shoot K
green stem conductance
shoot hydraulic capacity
Schultz HR, Matthews MA (1988) Resistance to water transport in shoots of Vitis vinifera L. Relation to growth at low water potential. Plant Physiology 88(3): 718-724 Lovisolo C, Schubert A (1998) Effects of water stress on vessel size and xylem hydraulic conductivity in Vitis vinifera L. Journal of Experimental Botany 49(321): 693-700
Shoot Hydraulic Conductance Quality
A VSP-trained Pinot Noir vineyard in Oregon's Willamette Valley targets Shoot_Length_Quality of 100-120 cm at véraison, positioning the shoot apex at the top catch wire with minimal hedging requirement, achieving this through regulated deficit irrigation initiated at fruit set that limits late-season growth while maintaining the 14-16 leaves per shoot required for complete fruit ripening in this cool-climate region.
Shoot_Length_Quality is a Vegetative_Growth_Vigor_Quality that inheres in a grapevine (Vitis vinifera L.) and represents the total linear extension of primary shoots from base to apex, integrating the cumulative effects of node number and internode length over the entire growing season. This quality provides the most comprehensive single measure of seasonal vegetative vigor, directly determining canopy dimensions, light interception capacity, and the scale of canopy management interventions required for optimal fruit quality. In BFO terms, Shoot_Length_Quality exists as a one-dimensional spatial quality whose final value emerges progressively through the season, reaching its maximum at growth cessation (typically BBCH 79-81) and remaining stable through dormancy until pruning removes the distal portions. The relationship between shoot length and fruit quality is characteristically curvilinear, with insufficient shoot length indicating stressed or weak vines unable to ripen fruit adequately, optimal shoot length providing sufficient leaf area for fruit maturation without excessive shading, and excessive shoot length creating dense canopies with disease-promoting microclimates and delayed, uneven ripening. Target shoot lengths vary substantially with training system and cultivar, ranging from 80-100 cm for spur-pruned, bilateral cordon systems with moderate vigor varieties to 150-200 cm for cane-pruned systems with vigorous varieties requiring longer shoots to distribute crop load appropriately. The timing of shoot length attainment relative to phenological milestones carries significant viticultural implications, with shoots reaching final length well before véraison indicating appropriate vigor control, while continued shoot extension through véraison suggests imbalanced source-sink relationships requiring remediation through hedging, crop adjustment, or water stress imposition. Shoot length uniformity within a vine and across a vineyard block represents an important secondary quality dimension, as heterogeneous shoot lengths complicate canopy management operations designed for uniform canopy dimensions and create ripening variability affecting harvest logistics and wine quality consistency.
Shoot_Length_Quality integrates Internode_Length_Quality across all nodes, representing the product of node number times average internode length. This quality directly determines canopy dimensions and thus the relationship between vine size and trellis system dimensions. The class relates to canopy management through hedging requirements, training system selection, and the economic tradeoff between shoot length control through cultural practices versus mechanical intervention.
Gesamttrieblänge
Rutenlänge
Trieblänge
cane length
final shoot length
total shoot length
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 24-31 Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review. American Journal of Enology and Viticulture 60:251-268
Shoot Length Quality
Cabernet Sauvignon in Bordeaux shows shoot lignification from June through October at BBCH 31-47, initiating in basal internodes in early June progressing acropetally through shoot reaching apical regions by September-October, accumulating starch reserves to 20-28% dry weight creating brown woody mature shoots with complete lignification before dormancy entry under oceanic climate conditions of 18-24°C summer declining to 10-15°C autumn, with management promoting moderate late-season water stress supporting timely thorough maturation typical of this vigorous late-season cultivar requiring complete lignification for adequate winter hardiness
Shoot lignification represents the fundamental maturation process where herbaceous shoot tissues undergo secondary wall deposition and lignin incorporation transforming flexible green stems to rigid woody structures that provide mechanical support, enable perennial growth through winter survival, and serve as carbohydrate storage organs for subsequent season development. This essential developmental process begins in basal shoot regions during active growth and progresses acropetally following shoot elongation, typically spanning 8-14 weeks from initiation in basal internodes through completion in apical regions by dormancy entry, establishing permanent woody architecture from annual shoot growth. Lignification encompasses secondary cell wall synthesis in vascular tissues, fibers, and parenchyma cells, lignin monomer biosynthesis and polymerization creating hydrophobic matrix binding wall polysaccharides, progressive tissue rigidification through wall thickening and cross-linking, and metabolic transition from active growth to maintenance and storage functions. The process shows characteristic basipetal-to-acropetal progression with basal internodes initiating lignification during early-mid summer while shoot apex continues active growth, middle internodes lignifying during mid-late summer, and apical internodes completing maturation during late summer-autumn before dormancy. Lignification initiation occurs when internode elongation completes and cells transition from expansion to differentiation and secondary wall synthesis, with timing varying along shoot from early June in basal regions to August-September in apical regions in temperate climates. Secondary wall deposition represents central lignification process involving cellulose and hemicellulose synthesis forming secondary wall framework layers designated S1, S2, and S3 with specific microfibril orientations, followed by lignin monomer synthesis producing p-coumaryl, coniferyl, and sinapyl alcohols from phenylalanine through phenylpropanoid pathway, and lignin polymerization through oxidative coupling creating three-dimensional polymer network. Lignin content increases progressively during maturation from negligible in actively growing tissues to 15-25% dry weight in fully lignified mature shoots, with lignification proceeding from vascular tissues outward through adjacent fiber and parenchyma cells creating progressive rigidification. Cell types showing most extensive lignification include xylem vessel elements requiring rigid walls for water transport under tension, xylem fibers providing mechanical support, and interfascicular fibers between vascular bundles contributing to shoot rigidity. Mechanical properties change dramatically during lignification from flexible herbaceous tissue with tensile strength 5-15 MPa and high flexibility in unlignified shoots, to rigid woody structure with strength 40-80 MPa and minimal flexibility in fully lignified mature shoots, enabling self-supporting architecture and resistance to mechanical loading. Vascular tissue maturation accompanies lignification with secondary xylem development from vascular cambium activity producing additional vessels increasing water transport capacity, secondary phloem formation enhancing assimilate translocation, and progressive primary tissue compression as secondary tissues develop. Periderm formation proceeds during lignification replacing epidermis with cork tissue providing protection, developing from cork cambium in subepidermal layers producing cork cells externally creating brown bark appearance and phelloderm internally contributing to cortex structure. Carbohydrate accumulation intensifies during lignification as growth cessation redirects photosynthate from growth to storage, with starch concentration increasing from negligible during active growth to 15-30% dry weight in fully mature lignified shoots by dormancy entry, providing critical reserves for subsequent season bud burst and initial growth. Reserve accumulation timing shows basipetal-acropetal pattern corresponding to lignification progression with basal regions accumulating reserves earliest and most thoroughly, while apical regions show later abbreviated accumulation particularly in vigorous shoots where late growth delays maturation. Metabolic changes during lignification include declining respiration from growth levels to maintenance rates 2-4 fold above completely dormant tissue, shifting protein synthesis from growth-related to storage and defense proteins, and activating phenylpropanoid pathway enzymes supporting lignin biosynthesis. Nitrogen dynamics during lignification involve declining nitrogen concentration from 2-3% in mature leaves and growing shoots to 1-1.5% in lignified shoots as structural carbohydrates accumulate and mobile nitrogen remobilizes to storage tissues or senescing leaves, with nitrogen conservation important for resource economy. Water content declines during lignification from 70-80% fresh weight in herbaceous tissues to 50-60% in lignified mature shoots, reflecting reduced metabolic activity, increased cell wall material deposition, and beginning dehydration preparing for winter dormancy. Hormonal regulation involves declining auxin and gibberellin from reduced apical production affecting growth promotion, increasing abscisic acid promoting maturation processes and stress tolerance development, ethylene potentially involved in senescence and maturation coordination, and jasmonates possibly affecting secondary metabolism and lignification processes. Temperature affects lignification rate with warm conditions promoting rapid progression while cool conditions slow processes, though lignification proceeds across wide temperature range unlike growth which requires narrow optimal conditions. Light intensity affects lignification through effects on photosynthetic carbon availability for reserve accumulation and possibly direct photomorphogenic effects, with high light supporting thorough rapid lignification while shade may result in incomplete lignification particularly in apical regions. Water availability affects lignification with moderate water stress potentially promoting earlier initiation and enhanced progression, while severe stress may cause incomplete lignification or premature growth cessation affecting tissue quality. Nutrient availability particularly nitrogen affects lignification with adequate but not excessive nitrogen supporting normal maturation, while excess may delay lignification prolonging herbaceous state, and deficiency may cause premature incomplete maturation. The process shows four distinguishable phases based on spatial progression along shoot advancing from basal shoot lignification in earliest maturing regions through progressive lignification moving acropetally, midshoot lignification in middle regions, and complete shoot lignification extending to apical zones. Cultivar variation appears in lignification timing and completeness with early-maturing varieties showing rapid thorough lignification while late-maturing types may show incomplete apical lignification particularly in vigorous growth, affecting winter hardiness in cold climates. Regional timing shows basal shoot lignification beginning June-July in warm climates, July-August in temperate regions, and August in cool areas, with progression continuing through late summer and autumn toward dormancy entry. Viticultural management considerations include ensuring adequate late-season growth cessation allowing proper maturation before winter, managing water and nitrogen avoiding excessive late growth interfering with lignification, assessing lignification completeness particularly in cold regions where inadequate maturation creates freeze damage risk, recognizing that thorough lignification with substantial reserve accumulation benefits subsequent season development, and understanding that incomplete lignification reduces winter survival and next-season vigor. Research applications include analyzing regulation of lignin biosynthesis and deposition, quantifying environmental and management effects on lignification affecting shoot quality, evaluating relationships between lignification and cold hardiness in climate adaptation, assessing carbohydrate partitioning between growth and storage affecting reserve accumulation, and documenting developmental transitions from annual herbaceous to perennial woody growth in grapevine as model woody perennial.
Shoot_Lignification represents maturation process within Shoot_Growth converting annual herbaceous growth to perennial woody structure. Distinguished from Shoot_Elongation and other growth processes by focus on maturation and lignin deposition. Subdivided into Basal_Shoot_Lignification, Progressive_Shoot_Lignification, Midshoot_Lignification, and Complete_Shoot_Lignification based on spatial-temporal progression. Critical for winter survival and subsequent season resource availability. Timing and completeness determine cold hardiness and reserve status.
Stockausreifung
Triebverholzung
cane lignification
shoot hardening
shoot woodification
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 282-305 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 215-235 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Shoot Lignification
Systematic phenotyping of Shoot_Orientation_Disposition at Geisenheim University measured shoot angles across 80 cultivars at weekly intervals throughout the growing season to characterize orientation dynamics. Shoots were allowed free growth on a minimal support system (single wire at 1.2 m), with angle from vertical measured at 30, 50, and 70 cm from shoot base using a digital inclinometer. Shoot_Orientation_Disposition showed substantial and consistent cultivar variation. Strongly upright 'Cabernet Sauvignon' maintained mean angle of 15° at 50 cm throughout the season, with shoot tips ascending to 8° even as shoots exceeded 150 cm, demonstrating persistent negative gravitropism. 'Pinot Noir' showed semi-upright characteristics (28° at 50 cm) with moderate tip ascent. 'Riesling' displayed horizontal to trailing orientation with 52° at 50 cm, increasing to 68° at 70 cm as shoot weight overcame gravitropic response. Strongly trailing 'Gewürztraminer' reached 72° at 50 cm with tips often below horizontal (> 90°, true trailing). Developmental analysis revealed that shoot orientation was established early (by 40 cm length) and remained stable thereafter, indicating that Shoot_Orientation_Disposition reflects inherent properties rather than progressive mechanical effects. Internode flexibility measurements showed that trailing cultivars had 35% lower internode stiffness than upright types, contributing to their pendulous growth. The practical significance was confirmed by shoot positioning labor requirements—trailing cultivars required 3.5× more positioning labor than upright cultivars to achieve equivalent VSP canopy geometry.
A Shoot_Orientation_Disposition is a Growth_Habit_Disposition that inheres in Vitis vinifera shoots and is realized through the inherent angle and direction of shoot growth relative to vertical, determining whether shoots naturally grow upward (positively gravitropic), horizontally, or downward (negatively gravitropic/trailing), and thus how the vine canopy develops in three-dimensional space. This disposition represents the gravitropic response program of the genotype, reflecting the balance between negative gravitropism (driving upward growth toward light) and the mechanical effects of shoot weight, internode flexibility, and growth rate that may overcome gravitropic orientation. Shoot orientation is determined by the shoot's angle from vertical, measured at standardized positions along the shoot (typically at 50 cm from the base for comparable assessment). Cultivars range from strongly upright ( 60° from vertical, often approaching or exceeding horizontal). Shoot_Orientation_Disposition has major implications for training system compatibility—upright cultivars naturally conform to VSP and similar vertical systems, while trailing cultivars resist vertical positioning and may perform better on sprawl, lyre, or pendulum systems that work with rather than against their natural orientation. The disposition also affects light distribution within canopies, fruit zone exposure, and the labor requirements for shoot positioning operations.
Shoot_Orientation_Disposition is_a Growth_Habit_Disposition addressing the angle of shoot growth from vertical. Orientation reflects the balance between gravitropic response and mechanical/architectural factors. Child classes include Upright_Growth_Disposition ( 60°). Orientation is established early in shoot development and remains stable. Internode stiffness contributes to orientation differences. Training system compatibility depends strongly on shoot orientation.
Gravitropische-Reaktion
Triebausrichtungs-Disposition
Triebwinkel
gravitropic response
shoot angle
shoot growth direction
Smart RE (1988) Shoot spacing and canopy light microclimate, American Journal of Enology and Viticulture 39:325-333 Intrieri C, Poni S (1995) Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality of mechanized Italian vineyards, American Journal of Enology and Viticulture 46:116-127 Louarn G et al. (2008) Characterization of the light interception efficiency of grapevine, Annals of Botany 101:385-399
Shoot Orientation Disposition
The shoot positioning protocol for a Guyot-pruned Pinot Noir vineyard in the Côte de Nuits specifies three passes of vertical shoot positioning (VSP) performed at E-L 12-15 (first pass, shoots 20-30 cm, tucked between the first pair of movable foliage wires at 1.0 m), E-L 17-19 (second pass, shoots 40-60 cm, tucked between the second pair at 1.3 m and first pair raised to final position), and E-L 23-27 (third pass, final positioning with lateral shoot removal in the fruit zone) — each pass requires 20-30 labor hours per hectare for manual shoot tucking in the dense 10,000 vines/ha canopy, with correct vertical orientation ensuring that the fruit zone between 0.6-1.0 m remains a single-layer curtain with 1.0-1.5 leaf layers as measured by point quadrat analysis.
A canopy management description that documents the manual or mechanical placement of growing shoots into the designated canopy geometry defined by the trellis wire configuration, specifying the timing, direction (vertical, pendant, or lateral), method of containment (tucking between movable wires, tying, or clipping), and target shoot orientation that establishes the spatial distribution of leaves and fruit within the canopy envelope.
Shoot_Positioning_Protocol describes the most labor-intensive and frequently repeated canopy management operation in VSP-trained vineyards, typically requiring 2-4 passes through the vineyard during the rapid shoot elongation phase (E-L 12 through E-L 27). The objective is to establish and maintain the vertical, pendant, or divided shoot orientation defined by the training system, preventing shoot crowding in the canopy interior and maintaining the separation between vegetative and fruiting zones: correctly positioned shoots create a uniform canopy wall with consistent leaf layer number, even light distribution, and effective air circulation that reduces disease incidence. Mechanical shoot positioning using rotating tines or brush-type machines has reduced labor requirements by 50-70% in wide-row vineyards (>1.8 m row spacing) but remains impractical in the narrow-row, high-density plantings (1.0-1.5 m row spacing) characteristic of premium European vineyards where manual positioning is still required.
Laubheftungs-Protokoll
Triebeinordnungs-Methode
Triebpositionierungs-Spezifikation
canopy positioning method
shoot training specification
shoot tucking protocol
Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles pp: 91-120 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 130-145 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 365-375
Shoot Positioning Protocol
Shoot synchrony assessment comparing cordon arms with different pruning histories in Syrah reveals the cumulative management effect on shoot uniformity, with consistently well-pruned cordon showing high shoot synchrony (shoot length CV 12% at mid-season, 85% of shoots within two-leaf stage range), while neglected cordon with accumulated pruning wounds and variable spur positions shows low synchrony (length CV 28%, shoots spanning four-leaf stage range), with the synchrony difference creating fruit-zone microclimate variation of 4°C temperature difference and 40% light variation between positions on the uniform versus non-uniform cordons, demonstrating how shoot synchrony affects fruit quality conditions.
Shoot synchrony quality is an organ synchrony quality that represents the developmental coordination among shoots at different positions within the grapevine canopy, characterizing the uniformity of shoot growth rates, developmental stages, and physiological status that determines canopy homogeneity affecting light interception, spray penetration, and the uniformity of fruit-zone microclimate across the vine. Within the BFO framework, this quality specifically addresses the shoot-level dimension of organ synchrony, distinguishing coordination among vegetative organs from berry or root synchrony while recognizing that shoot synchrony establishes the canopy architecture uniformity that affects light distribution, air movement, and the consistency of conditions experienced by fruit across different positions within the vine. The ontological significance lies in shoot synchrony determining canopy uniformity that propagates to fruit quality through microclimate effects, where synchronized shoot development produces consistent canopy density and fruit-zone conditions across the vine, while asynchronous development creates canopy heterogeneity with some zones densely shaded and others over-exposed, leading to fruit quality variation that reflects the microclimate differences imposed by non-uniform canopy architecture. The factors affecting shoot synchrony include bud position effects (distal versus basal shoots potentially differing in vigor and development rate), cordon/cane uniformity (variation in wood diameter or carbohydrate reserves affecting supported shoots), thinning uniformity (shoot thinning practices affecting the uniformity of remaining shoot population), and water/nutrient supply uniformity (consistent resource access promoting synchronized development). The management implications of shoot synchrony include the efficacy of canopy management operations performed at fixed timings (synchronized shoots all at appropriate stage for intervention, asynchronous shoots requiring multiple passes or compromised timing for some proportion), the uniformity of spray coverage (synchronized canopies enabling consistent penetration, non-uniform canopies creating coverage gaps), and the predictability of fruit-zone conditions (synchronized canopy producing consistent microclimate versus variable conditions from non-uniform canopy). Understanding shoot synchrony quality enables assessment of canopy uniformity at the shoot level, identification of management practices affecting shoot coordination, and canopy management planning that accounts for the degree of shoot population homogeneity.
Shoot_Synchrony_Quality addresses developmental coordination among shoots determining canopy uniformity. This quality affects fruit quality through microclimate consistency across vine positions. The class supports canopy management through shoot synchrony characterization.
Kronenuniformität
Triebentwicklungskoordination
Triebsynchronie
canopy uniformity
shoot developmental coordination
shoot population homogeneity
vegetative Synchronie
vegetative synchrony
Smart RE, Robinson MD (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 1-88 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 59-144
Shoot Synchrony Quality
The shoot thinning protocol for Cabernet Sauvignon on bilateral cordon in Coonawarra specifies manual shoot thinning at E-L 9-12 (shoots 10-20 cm, inflorescences visible but not yet fully separated) targeting 14-16 retained shoots per vine (7-8 per cordon arm at 20 cm spacing) from the initial post-budburst population of 25-35 shoots — removal priorities are non-fruitful shoots (identified by the absence of visible inflorescence primordia), double shoots emerging from single spur buds (the weaker of the pair removed), downward-growing shoots, and any shoots emerging from the trunk or cordon base (water shoots or suckers), with the thinning operation requiring 15-25 labor hours per hectare depending on initial shoot population density.
A canopy management description that documents the selective removal of excess, mispositioned, or non-fruitful shoots from the vine during early spring to reduce canopy density to the target shoot number per vine, specifying the timing (phenological stage), target shoot density (shoots per vine, per meter of row, or per meter of cordon), shoot selection criteria (removal of double shoots, water shoots, and non-fruitful shoots), and method (manual or mechanical).
Shoot_Thinning_Protocol describes the earliest growing-season canopy management operation and one of the most effective interventions for establishing the target canopy density that subsequent operations will maintain. The fundamental principle is that winter pruning determines bud number but not all buds produce shoots of equal value — non-count buds (base buds and latent buds) frequently produce non-fruitful or weakly fruitful shoots that contribute to canopy density without proportional contribution to yield, while compound buds may produce two or three shoots where only one is desired: shoot thinning corrects these discrepancies between the planned bud population and the actual shoot population. The timing of shoot thinning is critical — removal at E-L 5-12 (shoots 5-20 cm) is easily performed by hand snapping and causes minimal wound damage, while delayed thinning at E-L 15+ requires secateurs, creates larger wounds susceptible to disease infection, and has already allowed excess shoots to compete for vine resources during the critical post-budburst establishment period. Early shoot thinning also partially compensates for over-generous bud loads by reducing shoot number to the target density.
Ausbrechen-Spezifikation
Grüntrieb-Selektion
Triebausdünnungs-Protokoll
desuckering specification
green shoot thinning
shoot removal protocol
Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles pp: 91-105 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 130-140 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 365-370
Shoot Thinning Protocol
A shoot thinning response quality study in Central Otago Pinot Noir compared thinning intensities removing 0%, 25%, 50%, and 75% of emerged shoots at 20 cm shoot length. Retained shoots in the 25% removal treatment showed modest response with 8% increase in final cane weight versus unthinned controls. The 50% removal treatment exhibited intermediate shoot thinning response quality with 23% cane weight increase and moderate lateral development. The 75% removal treatment produced strongest response with 52% cane weight increase, excessive lateral growth requiring follow-up hedging, and development of secondary clusters from previously dormant lateral buds, demonstrating that extreme shoot thinning can trigger responses that undermine intended canopy improvements and require integrated management approaches.
Shoot thinning response quality is a vigor manipulation response quality that inheres in a grapevine organism and represents the characteristic pattern and intensity of compensatory growth exhibited by retained shoots following selective removal of excess shoots during the spring growth period to optimize shoot density, improve canopy architecture, and balance crop load with vine capacity. This quality captures the redistribution of growth resources from removed to retained shoots and the subsequent changes in retained shoot vigor, lateral development, and fruit cluster development that follow shoot thinning operations. The ontological basis of shoot thinning response quality rests on the principle of resource redistribution, wherein removal of competing sinks releases water, nutrients, and carbohydrates for allocation to remaining shoots, while simultaneously altering canopy light distribution and hormonal signaling patterns that influence growth of retained tissues. Unlike topping or hedging responses that primarily involve new lateral shoot development from released buds, shoot thinning response quality predominantly manifests as enhanced vigor in existing retained shoots through increased internode elongation, leaf size expansion, lateral shoot development along retained primaries, and potentially altered cluster development depending on thinning timing relative to flowering. The magnitude of shoot thinning response varies with thinning intensity, timing, and vine vigor status, with removal of greater proportions of total shoots amplifying compensatory responses in retained shoots, and earlier thinning permitting longer duration for response expression before growth cessation. Vigorous vines with abundant resource availability exhibit stronger shoot thinning responses than balanced or stressed vines, potentially negating intended benefits of shoot number reduction if retained shoots develop excessive vigor and dense lateral growth. The qualitative aspects of shoot thinning response include changes in retained shoot architecture such as increased cane diameter and lateral length, altered fruit zone density as remaining clusters develop without competition from removed shoots, and modifications to cluster architecture including potentially increased berry size and reduced compactness depending on thinning timing. Optimal shoot thinning strategies consider expected response characteristics to calibrate target shoot density below final desired levels, anticipating that compensatory responses will increase canopy fill from retained shoots. The interaction between shoot thinning response quality and fruit composition operates through modified source-sink ratios affecting individual berry development and through canopy microclimate changes influencing fruit exposure and temperature regimes during ripening.
Shoot_Thinning_Response_Quality is_a Vigor_Manipulation_Response_Quality, representing the compensatory response in retained shoots following selective removal of excess shoots. Distinguished from Topping_Response_Quality and Hedging_Response_Quality by the mechanism of response, which primarily involves enhanced growth of existing retained shoots rather than new shoot development from released buds. The timing relative to phenological development, particularly relative to bloom, influences both vegetative response and cluster development effects. Related to Canopy_Density_Quality and Fruit_Set_Percentage_Quality through shared physiological mechanisms.
Ausbruchreaktion
Frühjahrsausdünnung-Antwort
Triebausdünnungsreaktion
Triebentfernungsreaktion
desuckering compensation
shoot density adjustment response
shoot removal response
spring thinning response
Reynolds AG et al. (2005) Timing of shoot thinning in Vitis vinifera: impacts on yield and fruit composition variables. American Journal of Enology and Viticulture 56:343-356 Sun Q et al. (2012) Impact of shoot and cluster thinning on yield, fruit composition, and wine quality of Corot noir. American Journal of Enology and Viticulture 63:49-56 Bernizzoni F et al. (2011) Long-term performance of Barbera grown under different training systems and within-row vine spacings. American Journal of Enology and Viticulture 62:443-453
Shoot Thinning Response Quality
The shoot tip color description of Malbec documents an intensely anthocyanin-pigmented shoot tip (OIV 003:7) with deep crimson-red coloration extending uniformly across the developing leaf primordia and shoot apex, overlying a green ground color that is visible only on the ventral surfaces of unfolding leaves — this exceptionally strong tip pigmentation is one of the most distinctive visual markers of Malbec in the vineyard, immediately separating it from the moderately pigmented Merlot (OIV 003:5, red confined to dorsal margins of leaf primordia), the lightly pigmented Cabernet Sauvignon (OIV 003:3, pink-red blush on adaxial surface only), and the virtually unpigmented Sémillon (OIV 003:1, green-yellow throughout).
A shoot tip description component that documents the ground color and anthocyanin pigmentation pattern of the developing shoot apex and first unfolding leaves, recording the green, yellow-green, or bronze base color overlaid with the presence, intensity, and distribution of red-purple anthocyanin coloration scored according to OIV descriptors 003 (intensity of anthocyanin coloration of shoot tip) and related characters, providing one of the earliest and most visually distinctive cultivar identification features.
Shoot_Tip_Color_Description documents the color phenotype that is arguably the single most instantly recognizable visual character for cultivar identification during the spring growth period — an experienced ampelographer can often tentatively identify a cultivar from 10 meters distance based solely on shoot tip color. The anthocyanin pigmentation of the shoot tip reflects the constitutive activity of the anthocyanin biosynthetic pathway in young, actively dividing tissues, controlled by the same VvMYBA1/A2 regulatory genes responsible for berry skin color but with additional regulatory inputs specific to vegetative tissue expression: cultivars with the highest shoot tip anthocyanin (OIV 003:7-9, such as Malbec, Tannat, and Alicante Bouschet) have strong constitutive pathway activity in all tissues, while cultivars with moderate berry coloration but low shoot tip anthocyanin demonstrate tissue-specific regulatory differences. The temporal window for scoring shoot tip color is narrow — anthocyanin expression is strongest in the first 1-2 weeks after budburst and diminishes as leaves expand and mature, necessitating timely observation during a period that may span only 10-14 days in warm spring conditions.
Apex-Anthocyan-Muster
Triebspitzen-Farbbeschreibung
Wachstumspunkt-Färbung
growing point coloration
shoot apex color record
tip anthocyanin pattern
Shoot Tip Color Description
The shoot tip description of Sauvignon Blanc recorded at E-L stage 7-9 (first leaves separating to 2-3 leaves unfolded) documents a fully open shoot tip (OIV 001:7) with dense prostrate trichomes giving a white-woolly appearance (OIV 004:9), yellow-green ground color with medium anthocyanin pigmentation concentrated on the dorsal side of developing leaf primordia (OIV 003:3), and prostrate habit — this combination of open form, dense white pubescence, and moderate anthocyanin expression is characteristic of Sauvignon Blanc and distinguishes it from morphologically similar cultivars such as Sémillon (less pubescent shoot tip, OIV 004:5) and Colombard (stronger anthocyanin expression, OIV 003:5) during the brief spring observation window that typically spans only 2-3 weeks before shoot elongation modifies the tip morphology.
An ampelographic description component that documents the morphological characteristics of the grapevine shoot tip — the apical 2-4 cm of actively growing shoots including the shoot apex, developing leaf primordia, and the first 1-3 unfolded leaves — recording color, hairiness (trichome type and density), anthocyanin pigmentation pattern, and growth form that constitute some of the earliest observable and most diagnostically valuable characters in the ampelographic identification of Vitis vinifera cultivars.
Shoot_Tip_Description addresses the first organ available for ampelographic observation in the growing season — the developing shoot tip that is visible from budburst (E-L stage 2-3) through early shoot growth (E-L stage 7-12) and provides some of the most distinctive cultivar identification characters during the critical spring period when viticultural decisions depend on confirmed cultivar identity. The shoot tip is particularly valuable for identification because its morphology is less influenced by environmental variation than mature organ characters — the dense trichome coverage, anthocyanin patterns, and growth form of the shoot tip are primarily determined by cultivar genotype with relatively low genotype × environment interaction compared to leaf size, cluster weight, or berry composition: this genotypic stability makes shoot tip descriptors among the most reliable ampelographic characters for cultivar identification across diverse growing conditions. The temporal limitation of shoot tip observation (characters change rapidly as the shoot elongates and tip morphology transitions to mature shoot form within 3-4 weeks of budburst) necessitates careful timing of field observations and is one reason why ampelographic description requires vineyard visits at multiple specific phenological stages.
Triebspitzen-Beschreibung
Wachstumsspitzen-Morphologie
apical shoot characterization
apikale Trieb-Charakterisierung
growing tip morphology
shoot apex description
Shoot Tip Description
Shoot tip desiccation quality monitoring in Coonawarra Cabernet Sauvignon documented the progression of tip damage during an extended dry period with irrigation system failure. Initial stress symptoms appeared as growth cessation and tendril drooping within the first week of deficit, progressing to yellowing of unexpanded apical leaves by day 10, and developing into characteristic shoot tip desiccation with browning of the terminal 3-5 cm by day 14. The severity of shoot tip desiccation quality was scored on a four-point scale from 0 (healthy, active growth) through 2 (growth ceased, yellowing apex) to 4 (extensive browning and desiccation extending >5 cm from tip), with scores correlated to cumulative water deficit calculated from soil moisture and evapotranspiration records. Importantly, shoot tip desiccation quality proved irreversible once tissue browning occurred, with damaged tips unable to resume growth even after irrigation restoration, though lateral buds below damaged tips eventually broke to provide replacement growth. The permanent nature of tip damage above score 2 established this threshold as the management intervention limit, with irrigation required before tip desiccation to preserve growing points for continued canopy development.
Shoot tip desiccation quality is a drought symptom expression quality that inheres in grapevine shoot apices and represents the characteristic drying and browning of the actively-growing shoot tip region exhibited under sustained water deficit, progressing from initial growth cessation through yellowing of unexpanded leaves at the apex to eventual necrosis and desiccation of the terminal portion of the shoot when stress severity or duration exceeds the capacity for tip survival. This quality provides a visual indicator of sustained moderate-to-severe stress that has persisted long enough to cause tissue damage at the metabolically active shoot tip, distinguishing prolonged stress episodes from transient deficits that may cause temporary wilting without lasting damage to growing points. The ontological significance of shoot tip desiccation quality lies in its indication of stress severity and duration sufficient to cause irreversible tissue damage, distinguishing more serious stress episodes from the transient deficits that cause recoverable symptoms in mature tissues while leaving the growing point viable for resumed growth when water availability improves. The biological basis of shoot tip desiccation involves the vulnerability of actively dividing and expanding cells at the shoot apex to water deficit, with the high metabolic demands and thin cell walls of meristematic and elongating tissues creating sensitivity to even brief periods of severe stress, combined with the terminal position of shoot tips at the end of the hydraulic pathway where water supply is most tenuous under deficit conditions. The progression of shoot tip desiccation begins with growth cessation as turgor falls below the threshold for cell expansion, followed by yellowing of unexpanded leaves as chlorophyll degrades in stressed tissue, and culminating in browning and desiccation as cells die from prolonged water deficit exceeding survival capacity. The extent of shoot tip damage provides diagnostic information about stress severity and duration, with minimal tip browning indicating moderate stress causing temporary growth cessation while extensive desiccation extending back multiple nodes indicates severe prolonged stress that may have broader impacts on vine function and fruit development.
Shoot_Tip_Desiccation_Quality is_a Drought_Symptom_Expression_Quality, representing the irreversible damage to actively-growing shoot apices under sustained stress. This class addresses a more severe symptom indicating stress duration and intensity causing permanent tissue damage. Distinguished from recoverable symptoms like Leaf_Wilting_Quality by the irreversible nature of desiccation damage. The progression from growth cessation through yellowing to browning provides severity staging. Extent of desiccation (cm from tip) provides quantitative metric. The loss of growing points has implications for canopy development and subsequent management.
Apex-Desiccation
Triebspitzen-Verbräunung
Triebspitzen-Vertrocknung
Wachstumsspitzen-Nekrose
apex desiccation
growing point damage
shoot tip necrosis
tip browning
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 453-507 Lovisolo C et al. (2010) Drought-induced changes in development and function of grapevine (Vitis spp.) organs. Functional Plant Biology 37:98-116 Hochberg U et al. (2017) Stomatal closure, basal leaf embolism, and shedding protect the hydraulic integrity of grape stems. Plant Physiology 174:764-775
Shoot Tip Desiccation Quality
The shoot tip hairiness description of Syrah documents very dense prostrate (woolly) trichomes on the shoot tip and first unfolding leaves (OIV 004:9), creating the characteristic white-felted appearance that is one of Syrah's most recognizable field identification features — the prostrate trichomes are appressed to the leaf surface with interlocking fibers producing a continuous woolly mat that obscures the underlying leaf tissue, giving the shoot tip a silvery-white appearance contrasting sharply with the underlying green-red tissue. This very dense pubescence distinguishes Syrah from the glabrous (hairless) shoot tips of cultivars such as Chasselas (OIV 004:1), the sparsely hairy tips of Cabernet Sauvignon (OIV 004:3), and the moderately hairy tips of Grenache (OIV 004:5), and can be readily observed and scored during the 3-4 week spring observation window without magnification.
A shoot tip description component that documents the type, density, and distribution of trichomes (plant hairs) on the developing shoot apex and first unfolding leaves, distinguishing between prostrate (appressed, woolly) and erect (upright, bristle-like) trichome types scored according to OIV descriptor 004 (density of prostrate hairs of shoot tip), with trichome characteristics providing highly diagnostic cultivar identification features that are among the most genetically stable and environmentally robust of all ampelographic characters.
Shoot_Tip_Hairiness_Description documents a character of exceptional diagnostic value in ampelography — trichome density on the shoot tip is considered one of the three most important single characters (alongside mature leaf blade shape and berry skin color) for visual cultivar identification, and is the most reliable of the three because it is least affected by environmental variation. Trichome density is controlled by a limited number of genes with strong genotypic determination, producing consistent expression across sites, years, and management practices — a cultivar scored as OIV 004:9 (very dense) will consistently show very dense pubescence regardless of rootstock, irrigation, canopy management, or climate, whereas leaf blade shape and particularly berry color intensity can vary by 1-2 OIV scale points between environments: this genetic stability makes shoot tip hairiness the character of first choice for field identification where a quick, reliable cultivar distinction is needed. The two trichome types — prostrate (woolly, appressed, with interlocking fibers creating a mat-like covering) and erect (upright, bristle-like, not forming a mat) — are developmentally and genetically distinct, and their ratio and density create cultivar-specific pubescence patterns that are remarkably consistent and distinctive.
Spitzenbehaarung-Charakterisierung
Trichom-Dichte-Beschreibung
Triebspitzen-Behaarungsbeschreibung
shoot tip pubescence record
tip hairiness characterization
trichome density description
Shoot Tip Hairiness Description
A shoot tip terminal inflorescence on a 2-node fruiting spur of Vitis vinifera cv. Chardonnay in Burgundy, France, positioned 1.2 cm behind the shoot apical meristem within the protected shoot tip zone, measuring 8.5 cm length representing very reduced dimensions, developing from the terminal bud position with the apical meristem immediately above transitioning from vegetative to reproductive fate, exhibiting relatively simple branched architecture with central rachis bearing 2-3 orders of lateral branching, bearing approximately 180 flowers representing reduced flower number compared to standard primary inflorescences, positioned within the shoot tip microenvironment experiencing relative humidity of 82% (compared to 58% ambient) and temperature moderation of 2-3°C below ambient through transpirational cooling, showing phenological development at BBCH stage 57 (inflorescence clearly visible, flowers closely pressed) indicating slightly delayed development, extreme terminal position creating maximum vulnerability to shoot tip removal through topping, frost damage to tender apical tissues, or pest attack on growing points, occurring on this Chardonnay spur-pruned architecture where short spurs consistently produce terminal inflorescences, measured in late May during pre-bloom period, representing extreme shoot tip terminal inflorescence positioning and architecture.
A terminal primary inflorescence positioned in immediate proximity to the shoot apical meristem within the shoot tip region in Vitis vinifera, characterized by location within 1-3 cm of the apical meristem dome representing extreme terminal placement, spatial association with the shoot tip zone including youngest leaf primordia and apical growing point, development from the terminal bud position or uppermost lateral bud immediately adjacent to apex, occurrence on very short determinate shoots or spurs where reproductive development dominates over vegetative extension, very small dimensions typically 6-12 cm length reflecting severe resource competition with shoot extension and late developmental timing, exposure to the protected microenvironment of the shoot tip region including elevated humidity and moderated temperatures but also experiencing vulnerability to apical damage, and occurrence patterns showing restriction to specific architectural contexts including short fruiting spurs and determinate reproductive shoots. Shoot tip terminal inflorescences represent the extreme terminal positioning variant, developing in immediate association with the actively growing shoot apex, exhibiting maximum exposure to apical environmental conditions and vulnerability to tip damage, and creating distinctive architectural arrangements where reproductive development dominates the shoot apex.
Shoot tip terminal inflorescences represent the most extreme terminal positioning, distinguished from general terminal inflorescences by immediate proximity to apical meristem and shoot tip region rather than simply distal shoot location. The shoot tip designation emphasizes spatial association with the active growing apex and youngest structures. Distance of 1-3 cm from apical meristem represents the protected shoot tip zone, contrasting with terminal inflorescences at 5-10 cm showing less intimate apical association. Microenvironmental effects include elevated humidity, moderated temperatures, and mechanical protection from surrounding young leaves creating favorable conditions for development but also concentrating vulnerability to tip damage affecting both inflorescence and apical meristem simultaneously. Developmental context involves transition from vegetative to reproductive apical meristem identity, with determinate reproductive shoots showing complete conversion while some shoots may maintain vegetative capacity above the terminal inflorescence. Size reduction reflects severe constraints including late initiation, intense resource competition with apical vegetative growth, and limited space in the compact shoot tip region. Occurrence is highly context-dependent with short spurs and determinate shoots favoring shoot tip terminal development while vigorous long canes rarely produce this architecture. Management implications include extreme vulnerability to shoot positioning and topping practices that target shoot tips, requiring careful handling to preserve these terminal inflorescences if retention is desired. Transformation to cluster creates apex-positioned fruit requiring specialized harvest considerations.
Apex-assoziierter Blütenstand
Spitzenmeristem-proximale Infloreszenz
apex-associated inflorescence
apical meristem-proximal inflorescence
extreme terminal inflorescence
extreme terminale Infloreszenz
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 279-308 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 319-348 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 503-536.
Shoot Tip Terminal Inflorescence
A shoot tip terminal leaf at node 15 within the shoot tip region of Vitis vinifera cv. Pinot Noir in Burgundy, France, positioned 2.5 cm behind the shoot apex among three younger primordia and the apical meristem dome, measuring 1.8 cm length in tightly folded early unfolding stage representing approximately 10% of eventual mature size, exhibiting very pale yellowish coloration with pronounced bronze-red tinge from high anthocyanin providing photoprotection, positioned within the humid protected microenvironment of the shoot tip where relative humidity exceeds 85% (compared to 60% ambient) and temperature is moderated 2-3°C below ambient through transpirational cooling from surrounding tissues, composed of rapidly dividing cells with mitotic index of 15% indicating active cell proliferation, showing no tissue differentiation beyond uniform parenchymatous organization, contributing zero photosynthesis representing pure sink consuming approximately 6 mg carbohydrate per day, occurring during vigorous shoot extension at 8-10 cm per week, emerging 3 days previously during active apical meristem function initiating new primordia every 2.5 days, measured at shoot BBCH stage 75 (berries pea-sized) in late June, representing the protected shoot tip terminal leaf zone where newest leaves develop under favorable microenvironmental conditions before exposure to full ambient environment.
A terminal apical leaf positioned within the shoot tip region including the apical meristem and the 2-4 most recently initiated leaf primordia in Vitis vinifera, characterized by position within 3-5 cm of the shoot apex in the actively growing tip zone, extremely recent emergence within previous 2-5 days representing the newest recognizable leaves, very small size typically 1-3 cm length in early unfolding stages, tight spatial association with shoot apical meristem and younger primordia still embedded within protective structures, very pale coloration with high anthocyanin content providing photoprotection for delicate meristematic tissues, extreme fragility with cells in active division and early expansion phases, complete absence of functional tissue differentiation, zero photosynthetic contribution representing pure resource sink, and functional integration with the shoot tip as part of the active growth zone consuming resources for continued shoot extension. Shoot tip terminal leaves represent the immediate developmental output of apical meristem activity, emerging within the protected shoot tip environment where humidity remains high, temperature moderation occurs through transpirational cooling from surrounding older leaves, and mechanical protection is provided by older leaf primordia and remaining bud scale fragments, with this protected microenvironment enabling the extremely vulnerable meristematic and early expansion tissues to develop before exposure to full ambient environmental conditions that could cause desiccation, photoinhibition, or mechanical damage.
Shoot tip terminal leaves represent the most protected subset of terminal apical leaves, distinguished by position within the shoot tip microenvironment and association with active meristematic zones. The shoot tip region creates favorable conditions for extremely vulnerable young leaves through humidity maintenance, temperature moderation, and mechanical protection, with this protected zone extending approximately 3-5 cm from the apex depending on leaf size and internode length. Leaves within this zone show characteristics of the protected environment including reduced stress symptoms, optimal developmental progression, and lower damage incidence compared to exposed positions. The shoot tip terminal leaf population is dynamic during active extension, with continuous addition of new primordia at the apex and displacement of older leaves from the protected zone as internodes elongate. Shoot topping removes the entire shoot tip including all shoot tip terminal leaves and the apical meristem, immediately terminating the capacity for new leaf initiation and forcing developmental resources toward existing leaves and lateral structures. Environmental stresses affecting the shoot tip including severe heat, desiccating winds, or pest attack can damage multiple shoot tip terminal leaves simultaneously, disrupting continued shoot extension and potentially requiring lateral shoot development to restore growth.
Meristem-nahes Blatt
Spitzenregion terminales Blatt
apex region leaf
geschütztes Apex Weinblatt
meristem-proximal leaf
protected tip leaf
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 43-85 Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 103-136.
Shoot Tip Terminal Leaf
Detailed phenotyping of Shoot_Vigor_Disposition at Geisenheim University tracked shoot development across 45 cultivars at weekly intervals throughout the growing season, enabling characterization of growth rate dynamics, final dimensions, and growth component analysis. Measurements included primary shoot length, internode number and length, main leaf area, lateral shoot number and total lateral length, and the timing of growth cessation. Shoot_Vigor_Disposition showed wide cultivar variation with distinct growth patterns. High-vigor 'Silvaner' showed rapid early growth (peak rate 7.2 cm/day in week 6), extended growth duration (cessation week 16), and final shoot length of 285 cm with mean internode length of 9.8 cm and total lateral length of 180 cm per primary shoot. Moderate-vigor 'Riesling' displayed lower peak growth rate (4.8 cm/day), intermediate duration (cessation week 14), and final length of 165 cm with internode length of 7.2 cm and laterals of 85 cm. Low-vigor 'Gewürztraminer' showed modest peak growth (3.2 cm/day), early cessation (week 12), and final length of 95 cm with compact internodes (5.5 cm) and limited laterals (35 cm). Growth pattern analysis revealed that high-vigor cultivars differed from low-vigor primarily in growth duration rather than peak growth rate (r = 0.85 for duration vs. final length, r = 0.52 for peak rate vs. final length), indicating that Shoot_Vigor_Disposition reflects the period of sustained meristem activity rather than simply faster instantaneous growth. These findings enabled classification of cultivars into High, Moderate, and Low Shoot_Vigor_Disposition categories with distinct management implications.
A Shoot_Vigor_Disposition is a Vigor_Disposition that inheres in Vitis vinifera shoots and is realized through the rate and extent of primary shoot growth from budburst through growth cessation, determining shoot length, internode dimensions, leaf size, and lateral shoot production. This disposition represents the above-ground expression of vigor, driven by shoot apical meristem activity, cell division and elongation rates, and the supply of resources (carbohydrates, water, nutrients) from roots and reserves to actively growing shoots. Shoot vigor is quantified through multiple parameters including shoot elongation rate (typically 2-8 cm/day during rapid growth phase, varying by genotype and conditions), final shoot length at growth cessation, mean internode length, individual leaf area, and the number and length of lateral (axillary) shoots produced. High Shoot_Vigor_Disposition results in long shoots with extended internodes, large leaves, and prolific lateral production, while low shoot vigor produces compact shoots with shorter internodes, smaller leaves, and limited laterals. Shoot vigor has direct implications for canopy density and fruit zone microclimate—high shoot vigor creates dense, shaded canopies requiring intensive management to maintain fruit quality, while low vigor may produce insufficient canopy for optimal fruit maturation. The disposition is expressed through three categorical subclasses (High, Moderate, Low) that capture the practical range of variation among cultivars and guide training system selection and management approaches.
Shoot_Vigor_Disposition is_a Vigor_Disposition representing the above-ground vigor component. Shoot vigor reflects integrated meristem activity over the growing season. Child classes include High_Vigor_Disposition, Moderate_Vigor_Disposition, and Low_Vigor_Disposition as categorical descriptors. Key determinants are growth duration (weeks of active elongation) and peak growth rate. Lateral shoot production is an important component affecting canopy density. Shoot vigor determines training system requirements and canopy management intensity.
Oberirdische-Wuchskraft
Primärtrieb-Wachstum
Triebwuchsstärke-Disposition
aerial vigor
primary shoot growth
shoot growth capacity
Lebon E et al. (2006) Shoot development in grapevine (Vitis vinifera L.) is affected by the modular branching pattern of the stem and intra- and inter-shoot trophic competition, Annals of Botany 98:1045-1056 Schultz HR, Matthews MA (1988) Vegetative growth distribution during water deficits in Vitis vinifera L., Australian Journal of Plant Physiology 15:641-656 Louarn G et al. (2007) A method to assess light interception efficiency of a grapevine canopy based on shoot architecture and leaf area distribution, Australian Journal of Grape and Wine Research 13:29-38
Shoot Vigor Disposition
In Gewürztraminer vines in Alsace at latitude 48°N, short day perception occurs when night length exceeds approximately 11 hours in mid-September, with the mature leaves on mid-shoot positions serving as primary perception sites where phytochrome Pfr levels decline during the extended dark period below the threshold required to maintain growth-promoting gene expression, initiating the dormancy signaling cascade.
Short day perception in Vitis vinifera represents the initial sensory phase of photoperiod response during which photoreceptor systems in leaf tissues detect that day length has decreased below the critical threshold signaling approaching winter conditions. This photobiological process occurs in mature, fully expanded leaves where phytochrome photoreceptors are concentrated, with detection depending on the duration of the dark period rather than the light period, making it technically a response to long nights rather than short days. The phytochrome system central to short day perception consists of photoreversible chromoproteins that exist in two forms, the red light-absorbing Pr form and the far-red light-absorbing Pfr form, with the Pfr form produced during daylight gradually reverting to the Pr form during darkness at a rate that allows measurement of night length. When the dark period exceeds the critical duration for a particular genotype, the resulting low Pfr/Pr ratio at dawn initiates the signaling cascade that leads to dormancy-related gene expression changes. The critical night length for Vitis vinifera cultivars generally falls between 10-12 hours, corresponding to day lengths of 12-14 hours, with variation among cultivars reflecting adaptation to different latitudes of origin where the relationship between photoperiod and the timing of winter cold differs. The perception process is remarkably sensitive, with plants capable of detecting differences in night length of less than 30 minutes, providing precise timing information for seasonal transitions. Multiple phytochrome genes are expressed in grapevine, including PHYA and PHYB homologs that may serve partially redundant and partially distinct functions in photoperiod perception. Light quality as well as quantity influences perception, with the red to far-red ratio at dusk and dawn providing additional temporal information through effects on phytochrome photoequilibrium. The anatomical localization of perception primarily in mature leaves necessitates signal transmission to distant responding tissues including buds, establishing the requirement for long-distance signaling mechanisms that connect perception sites with developmental response sites.
Short_Day_Perception represents the initial sensory phase within Photoperiod_Response, specifically addressing the photoreceptor-mediated detection of critical night length. Distinguished from Photoperiod_Signal_Transduction which addresses downstream signaling events, and from Growth_Inhibition_Initiation which addresses the developmental response. Occurs in mature leaves rather than buds. Detection is technically of long nights rather than short days. Critical night length varies with cultivar origin latitude from approximately 10-12 hours.
Kurzphotoperioden-Perzeption
Kurztagwahrnehmung
Langnacht-Erkennung
critical daylength sensing
kritische Tageslängenerkennung
long night detection
photoperiod detection
short photoperiod perception
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Wake CMF, Fennell A (2000) Morphological, physiological and dormancy responses of three Vitis genotypes to short photoperiod. Physiologia Plantarum 109: 203-210 Borthwick HA, Hendricks SB (1960) Photoperiodism in plants. Science 132: 1223-1228
Short Day Perception
Wild V. vinifera ssp. sylvestris pistillate female flowers exhibit short filaments 0.3-0.8 mm at maturity supporting vestigial anthers (0.2-0.4 mm) positioned well below the prominent pistil (2.3-2.6 mm), with filaments showing minimal rapid elongation (from pre-anthesis 0.25-0.6 mm to mature 0.3-0.8 mm, only 20-30% increase) reflecting severe developmental suppression, and creating obligate cross-pollination dependency by preventing any possibility of self-pollination from the non-pollen-producing vestigial anthers. Gewürztraminer functionally female flowers show short reflex filaments 1.2-1.9 mm (reduced compared to hermaphrodite Vitis typical 2.5-3.5 mm) that curve downward positioning small anthers (0.4-0.6 mm) 0.8-1.5 mm below pistil (2.4-2.7 mm), with the combination of short length and reflex curvature creating strong spatial separation preventing self-pollination despite presence of morphologically recognizable though functionally impaired stamens. Breeding population segregants showing pistillate sex expression exhibit short filaments ranging from 0.4-1.5 mm reflecting variable stamen suppression strength, with shortest filaments (0.4-0.7 mm) associated with most complete stamen reduction (vestigial anthers, no pollen) and longer short filaments (1.2-1.5 mm) associated with rudimentary anthers showing some structural differentiation though still non-functional.
A short filament is a filament exhibiting reduced length typically less than 2 mm at maturity, characteristic of vestigial or rudimentary stamens in pistillate flowers and representing the lower range of functionally female flower filament length variation, reflecting arrested or suppressed filament development associated with stamen reduction in female-functioning flowers. Short filaments are characterized by= (1) mature length typically 0.2-2 mm depending on degree of stamen suppression, representing 10-70% of typical hermaphrodite filament length (2.5-3.5 mm) (2) reduced elongation during the rapid elongation phase at anthesis, often achieving only 50-150% length increase (from pre-anthesis 0.2-1 mm to mature 0.3-2 mm) compared to typical 150-250% increase in hermaphrodite filaments (3) reduced cortex parenchyma cell number and limited cell expansion capacity reflecting developmental arrest (4) variable orientation including erect, horizontal, or reflex positions depending on specific flower type and degree of development. Short filaments in pistillate flowers support vestigial or rudimentary anthers (0.2-0.5 mm) that produce no pollen or minimal non-viable pollen, with the short filament length positioning these non-functional anthers well below the prominent functional pistil (2-2.6 mm), creating clear spatial and functional dominance of female over male organs. Short filaments in functionally female flowers may show reflex curvature (in cases where filaments achieve 1.5-2 mm length allowing curvature) or may be too abbreviated (0.5-1 mm) to exhibit clear orientation, but in either case position anthers below or well to the side of the pistil. The developmental basis of short filaments involves suppression of normal filament elongation processes through genetic factors associated with sex determination= in pistillate flowers, female-determining alleles suppress stamen development generally including filament elongation in functionally female flowers, mechanisms causing stamen reflexing or pollen reduction may co-occur with partial filament length reduction though not to the extreme seen in pistillate types. Short filaments show limited rapid elongation capacity at anthesis, with elongation rates and final achieved lengths substantially below hermaphrodite norms, reflecting= reduced cortex cell proliferation during early development limiting cell number available for expansion impaired cell expansion capacity during rapid elongation limiting final size even of available cells possible earlier cessation of elongation before full potential is achieved. Environmental factors have limited influence on short filament final length, as the primary determinant is genetic suppression of development rather than environmental modulation, though favorable conditions may permit slightly greater elongation within the genetically constrained range (e.g., increasing short filament length from 0.8 mm to 1.2 mm under optimal versus stressed conditions, but not overcoming the fundamental short filament phenotype). The functional significance of short filaments relates to their role in creating spatial separation between non-functional or minimally functional anthers and the functional pistil, ensuring that even if minimal pollen is produced, spatial relationships prevent self-pollination and maintain obligate cross-pollination dependency. Short filaments combined with vestigial/rudimentary anthers represent intermediate stages in the evolutionary continuum from hermaphrodite (functional male and female) toward complete female unisexuality (complete stamen absence), providing material for developmental genetic studies of organ suppression.
Short_Filament represents reduced filament development characteristic of pistillate and functionally female flowers, distinguished from Long_Filament (extended development >3 mm) by substantially reduced mature dimensions (3 mm) and from typical hermaphrodite filaments (2.5-3.5 mm) by reduced length beyond normal variation. The short filament phenotype reflects genetic suppression of stamen development associated with female sex determination or functional female expression and contributes to obligate cross-pollination dependency through spatial separation of reduced anthers from functional pistils. Short filaments represent important developmental indicators for identifying female-functioning flower types and provide material for studies of organ suppression mechanisms and sex determination genetics in Vitis.
abbreviated filament
kurzer Staubfaden
reduced stamen stalk
reduzierter Staubblattfaden
verkürztes Filament
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 94-101 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Caporali, E. et al. (2003). The arrest of development of abortive reproductive organs. Sexual Plant Reproduction 15=291-300.
Short Filament
A short fruiting cane on Vitis vinifera cv. Nebbiolo in Barolo, Italy, retained at 4 nodes spanning 42cm along permanent cordon in traditional low-vigor hillside vineyard, selected from small-diameter (7mm) well-lignified cane showing complete brown color and appropriate node spacing, positioned vertically upward from cordon, expected to produce 3-4 fruiting shoots bearing 6-7 clusters totaling 1.6-1.9kg fruit, managed through removal of basal clusters (nodes 1-2 typically low fruitfulness) leaving 3-4 clusters from apical positions, achieving very low yields supporting exceptional phenolic development, complex tannin profiles, and aromatic intensity characteristic of premium Barolo requiring extended aging potential.
A fruiting cane retained at minimal length during winter pruning, typically 2-5 nodes, representing transition category between true canes and spurs, characterized by compact extent (20-55cm), low bud count emphasizing quality over quantity, suitability for spur-pruned systems or quality-focused cane-pruned vineyards, and production of few fruiting shoots requiring minimal canopy management while providing adequate crop for economic viability in premium wine production. Short fruiting canes represent conservative crop load philosophy approaching spur pruning in bud retention level, appropriate for naturally high-fruitfulness cultivars, low-vigor sites, premium quality objectives accepting reduced yields, and traditional systems in prestigious wine regions where historical practice favors restrained production emphasizing terroir expression and wine complexity.
Short fruiting canes blur the distinction between cane pruning and spur pruning, with 2-3 node retention essentially equivalent to spurs while 4-5 node retention maintains cane identity. The choice of short canes versus spurs is often traditional or practical (short canes easier to position in some training systems, spurs more compact for others) rather than physiologically fundamental. Short fruiting canes suit quality-obsessed producers in prestigious regions (Barolo, Barbaresco, Côte-Rôtie, Hermitage, select Burgundy sites), naturally high-fruitfulness varieties like Riesling or Gewürztraminer where even 2-3 buds provide adequate crop, and low-vigor sites where longer canes would exceed vine support capacity, with this conservative approach contributing to wine complexity and aging potential justifying premium pricing.
Kurzrute
Zapfenrute
compact fruiting cane
kurze Fruchtrute
minimal cane
near-spur cane
Short Fruiting Cane
Characterization of Short_Internode_Disposition at Cornell University documented shoot architecture in cultivars with compact internode development. Reference cultivars 'Pinot Noir', 'Gamay', 'Concord', and 'Niagara' were assessed for internode dimensions, canopy properties, and associated management implications. Short_Internode_Disposition cultivars showed consistent characteristics. 'Pinot Noir' exhibited archetypal short internodes with mean length of 5.8 cm (nodes 5-15), standard deviation of 0.9 cm (notably lower variability than long-internode types), and maximum internode length of only 7.5 cm. Shoot length to node number ratio was 6.8 cm/node. 'Gamay' showed similarly short internodes (6.0 cm mean) with compact canopy architecture. Canopy density analysis revealed that Short_Internode_Disposition cultivars developed 2.3 leaf layers at 18 shoots/m cordon, compared to 1.4 leaf layers for long-internode cultivars at equivalent shoot density, demonstrating the canopy-filling effect of compact internodes. Cluster spacing averaged 6.8 cm (compared to 12.5 cm in long-internode types), with clusters frequently touching within the fruiting zone, potentially facilitating Botrytis spread. Management trials confirmed that Short_Internode_Disposition cultivars required 35% more leaf removal labor to achieve equivalent fruit zone exposure compared to long-internode types at the same shoot density. Shoot stiffness was correspondingly higher (2.7 N·mm² versus 1.6 N·mm² for long-internode types), correlating with the upright to semi-upright growth habit typical of short-internode cultivars. Cell anatomy showed shorter cell length (125 μm versus 215 μm) as the primary basis for Short_Internode_Disposition.
A Short_Internode_Disposition is an Internode_Length_Disposition that inheres in Vitis vinifera shoots and is realized through limited cell elongation producing internodes of less than 7 cm mean length, resulting in compact shoot architecture with closely-spaced leaves, potentially overlapping leaf blades, and closely-positioned clusters. This disposition represents the lower range of internode dimensions, characteristic of cultivars with lower gibberellin sensitivity or abbreviated cell expansion phase that produces shortened internodes. Short-internode cultivars develop shoots where leaves are closely spaced and may overlap substantially, creating denser canopy conditions at equivalent shoot numbers compared to long-internode types. The mechanical consequence of short internodes is increased shoot stiffness (each segment is shorter relative to its diameter, reducing bending tendency), contributing to the association between short internodes and upright growth habit. Viticultural implications include naturally higher canopy density requiring more intensive management for adequate light penetration, closer cluster spacing (potentially increasing cluster-to-cluster disease transmission), and stiffer shoots that maintain vertical positioning more readily but may be more difficult to bend for training. Short_Internode_Disposition often co-occurs with Upright_Growth_Disposition, reflecting the relationship between internode compactness and shoot mechanical properties.
Short_Internode_Disposition is_a Internode_Length_Disposition representing compact internode length (< 7 cm mean). Characterized by limited cell elongation, higher shoot stiffness, and association with upright growth habit. Examples include 'Pinot Noir', 'Gamay', 'Concord', 'Niagara'. Creates denser canopies requiring more intensive management. Close cluster spacing may increase disease transmission risk. Associated with lower gibberellin sensitivity. Higher shoot stiffness contributes to upright positioning.
Enger-Knotenabstand
Kompakte-Triebarchitektur
Kurze-Internodien-Disposition
close node spacing
compact internode
dense shoot architecture
Schultz HR (1992) An empirical model for the simulation of leaf appearance and leaf area development of primary shoots of several grapevine (Vitis vinifera L.) canopy systems, Scientia Horticulturae 52:179-200 Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles, Adelaide Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Edition, Academic Press
Short Internode Disposition
A short main petiole on a mature sun leaf at node 7 of Vitis vinifera cv. Gewürztraminer in Alsace, France, measuring 6.8 cm length representing characteristic short petiole dimensions for this inherently short-petioled cultivar, petiolar index of 0.68 indicating petiole length only 68% of main vein length, diameter of 3.8 mm showing relatively thick construction for the short length, green coloration with moderate pink-red tinge from anthocyanin, moderate pubescence with visible trichomes, developing under exterior canopy position receiving 85% full sunlight with high red=far-red light ratio suppressing shade avoidance responses, demonstrating reduced flexibility with leaf blade mobility limited to approximately 90-degree rotation range but adequate for wind stress dissipation, supporting 4.5 gram leaf mass positioned at stable 60-degree angle from horizontal optimizing light interception, showing characteristic closed to slightly overlapping petiolar sinus typical of this cultivar, measured at BBCH stage 81 (beginning of berry ripening) in early August, representing the combined effects of genetic predisposition (Gewürztraminer inherently short-petioled) and environmental conditions (high light suppressing elongation) producing compact petiole architecture.
A main petiole exhibiting length in the lower range for the species and cultivar in Vitis vinifera, characterized by dimensions typically 6-9 cm length or less representing compact structural connection between lamina and shoot, often with petiolar index (petiole length / main vein length ratio) below 0.70-0.80 indicating petiole length substantially shorter than lamina dimensions, common development under high-light conditions where reduced petiole length maintains close lamina proximity to shoot without disadvantage for light capture, occurrence in certain cultivars showing genetic predisposition toward short petiole development (Gewürztraminer, some Muscat types, certain Pinot variants), functional implications including reduced flexibility limiting leaf blade mobility range but also reduced mechanical vulnerability and lower vascular resistance enabling efficient transpiration and photosynthesis, compact canopy architecture with leaves positioned close to shoots, and ampelographic significance as cultivar-diagnostic feature particularly reliable when measured on sun-exposed mature leaves under standardized conditions. Short main petioles develop through limited cell elongation during petiole development, influenced by genetic programming establishing cultivar-specific baseline dimensions, environmental signals including high red=far-red light ratios indicating open high-light environments that do not trigger shade avoidance elongation, hormonal regulation with reduced gibberellin activity or increased growth inhibitors limiting elongation, and potentially resource limitations constraining developmental investment in support structures favoring allocation to photosynthetic lamina.
Short main petioles are distinguished from medium and long petioles by dimensions typically below 8-9 cm and petiolar indices below 0.75-0.80, with exact thresholds varying by cultivar baseline dimensions. Genetic control establishes cultivar-specific ranges, with short-petioled varieties including Gewürztraminer and some Muscat types typically showing 6-9 cm petioles while long-petioled varieties show 11-15 cm under comparable conditions. Environmental effects are less pronounced than for long petioles, as high-light conditions typical of exterior canopy positions do not actively shorten petioles but simply avoid triggering shade-induced elongation. Functional implications include compact canopy architecture positioning leaves close to shoots creating dense foliage, reduced flexibility limiting positioning range but adequate for typical wind responses, lower vascular resistance supporting efficient transpiration and photosynthesis through shorter transport distances, and reduced mechanical vulnerability to wind breakage or petiole failure. Ampelographic utility is substantial as short petiole character is relatively stable across environmental conditions when properly standardized, serving as reliable identification feature particularly for cultivars at extreme short end of spectrum.
brief leaf stalk
compact petiole
kompakter Petiolus
kurzer Hauptblattstiel
shortened main petiole
verkürzter Blattstängel
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 198-215 This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 103-136.
Short Main Petiole
Functionally male vines occasionally occurring in Cabernet Sauvignon vineyards exhibit short pistil flowers at BBCH 65 with pistils measuring only 0.9-1.3 mm length (compared to 1.8-2.2 mm in normal hermaphrodite Cabernet), positioning stigma 0.8-1.2 mm below anther level, producing vigorous stamens with abundant pollen (22,000-28,000 grains per flower, 72-80% viability) but achieving <2% fruit set, with affected vines identified by profuse flowering but minimal berry development and typically removed from production vineyards. Wild dioecious V. vinifera ssp. sylvestris male plants in natural populations show extreme short pistil flowers with pistils only 0.6-1 mm long (30-50% of female plant pistil length), stigma rudimentary or absent, stamens producing copious pollen, and complete fruit set failure (<0.5%).
A short pistil functionally male flower is a functionally male flower in which female sterility is achieved primarily through severe abbreviation of pistil length that positions the stigma well below the anthers, combined with possible stigma dysfunction, effectively preventing pollination and fertilization despite production of abundant viable pollen from normal stamens, representing the morphologically most obvious mechanism of functional male expression. Short pistil functionally male flowers are characterized by= (1) pistils measuring only 50-75% of normal hermaphrodite pistil length, typically 0.8-1.5 mm long compared to 1.5-2.5 mm in hermaphrodite flowers (2) abbreviated pistil positioning the stigma 0.5-1.5 mm below the level of anthers rather than at or below anther level as in hermaphrodite flowers, creating spatial impossibility of self-pollination as pollen cannot fall upward from anthers to the lower-positioned stigma (3) stigma that may be small, poorly developed, or lacking the characteristic wet exudate, though in some cases stigma morphology appears normal despite reduced pistil length (4) ovary that may be reduced in size with abnormal or aborted ovules, though ovary structure can appear superficially normal in some cases (5) stamens showing normal or enhanced hermaphrodite morphology with erect position, vigorous filament elongation positioning anthers well above the abbreviated pistil, and abundant pollen production (18,000-30,000+ grains per flower) with high viability (65-85%). The short pistil mechanism creates physical impossibility of self-pollination through spatial separation similar to but opposite in direction from the reflex stamen mechanism in functionally female flowers= whereas reflex stamens position anthers below stigma, short pistils position stigma below anthers, both creating spatial barriers to autonomous self-pollination. The abbreviated pistil development may result from arrested pistil growth during flower development, with pistil primordia initiating normally but failing to elongate adequately, or from earlier developmental disruption affecting pistil primordium specification. Short pistil functionally male flowers achieve <1-3% fruit set despite abundant pollen production, with the rare fruit set events possibly resulting from exceptional pollination circumstances (pollen blown upward by turbulent air currents) or from occasional pistils showing less severe abbreviation. The pollen produced by short pistil functionally male flowers is fully functional and can successfully fertilize hermaphrodite or functionally female flowers when transferred by wind, making these flowers effective as pollen donors despite complete fruit set failure on the source plants. The short pistil condition is readily diagnosed through visual observation at anthesis, as the pronounced stamen-pistil length differential is immediately obvious, distinguishing short pistil functionally male flowers from hermaphrodite flowers (stamens extend beyond pistil but pistil is normal length) and from functionally female flowers (stamens reflex below normal-length pistil). The genetic control likely involves developmental regulatory genes affecting pistil elongation and may show partial overlap with genes controlling sex determination in dioecious wild Vitis.
Short_Pistil_Functionally_Male_Flower is distinguished from Non_Receptive_Functionally_Male_Flower by the primary mechanism of female sterility= morphological pistil abbreviation versus stigma receptivity failure, though these mechanisms may co-occur with abbreviated pistils often also showing stigma dysfunction. The short pistil condition is the more morphologically obvious mechanism and serves as the primary diagnostic feature for identifying functionally male individuals in breeding populations or as rogues in commercial plantings. The spatial inversion relative to functionally female flowers (short pistil placing stigma below anthers versus reflex stamens placing anthers below stigma) represents alternative evolutionary pathways toward functional unisexuality, both achieving spatial reproductive isolation that prevents self-pollination but through opposite organ modifications. Understanding the developmental genetics of pistil length regulation could inform studies of sex determination evolution in Vitis and potential applications in controlled pollination systems.
Blüte mit verkürztem Stempel
abbreviated-pistil male flower
kurzgriffelige männliche Blüte
reduced-pistil flower
verkürzte Pistill Blüte
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 101-103 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Caporali, E. et al. (2003). The arrest of development of abortive reproductive organs. Sexual Plant Reproduction 15=291-300 Picq, S. et al. (2014). A small XY chromosomal region explains sex determination. BMC Plant Biology 14=229.
Short Pistil Functionally Male Flower
Cabernet Sauvignon rare functionally male flowers (<2% frequency in some vintages) exhibit short styles 0.32-0.42 mm connecting reduced ovaries 0.55-0.72 mm to rudimentary stigmas 0.25-0.35 mm, with short styles representing 30-38% of abbreviated pistil length (1.1-1.5 mm total), containing transmitting tract tissue of reduced elaboration compared to typical hermaphrodite flowers, achieving <5% fruit set due to stigma non-receptivity and ovule abortion despite abbreviated pollen tube pathway. Wild V. vinifera ssp. sylvestris staminate male flowers show very short vestigial styles 0.15-0.28 mm connecting vestigial ovaries 0.25-0.45 mm to rudimentary stigmas 0.12-0.22 mm, with styles representing 28-42% of extremely reduced pistil length (0.5-0.8 mm total), with transmitting tract poorly differentiated and pistil achieving <1% fruit set reflecting complete functional male sterility. Merlot occasional hermaphrodite flowers show developmental variants with short styles 0.38-0.46 mm (versus typical 0.52-0.68 mm) connecting normal ovaries 0.82-0.98 mm to normal stigmas 0.4-0.5 mm, creating disproportionate pistil appearance with abbreviated style, achieving 42-48% fruit set (modestly reduced from typical 50-58%) suggesting short style has limited impact on fertilization when other pistil components remain functional. Pinot Noir functionally male segregants in breeding populations exhibit short styles 0.28-0.38 mm as part of overall pistil reduction syndrome including small ovaries (0.5-0.7 mm), reduced stigmas (0.22-0.32 mm), and abbreviated total pistil length (1.1-1.4 mm), with short styles showing reduced transmitting tract differentiation visible in anatomical sections as diminished extracellular matrix production compared to hermaphrodite siblings.
A short style is a style exhibiting reduced length typically less than 0.5 mm at maturity, characteristic of functionally male flowers with abbreviated pistils, staminate flowers with vestigial or rudimentary pistils, and occasional hermaphrodite flowers showing developmental reduction, representing styles at the lower end of the length range where abbreviated pollen tube pathway may correlate with reduced female reproductive function. Short styles are characterized by= (1) length typically 0.15-0.45 mm at anthesis, representing 30-70% of typical hermaphrodite style length (0.4-0.7 mm) (2) proportionally reduced diameter often 0.1-0.25 mm creating slender appearance (3) transmitting tract tissue present but potentially reduced in elaboration or maturity compared to longer styles, with extracellular matrix production and specialized cell differentiation potentially abbreviated (4) connection between reduced ovary (typically 0.4-0.8 mm) and rudimentary or reduced stigma (typically 0.2-0.4 mm) in functionally male or staminate flower contexts, or connection of normal-sized organs in rare hermaphrodite variants. Short styles occur through several mechanisms= (1) genetic determination in functionally male or staminate flowers where pistil reduction includes proportional style shortening as part of overall female organ suppression (2) developmental arrest or suppression affecting style elongation during pistil development (BBCH 53-57) while allowing some style differentiation (3) rare developmental variation in hermaphrodite flowers creating short styles through stochastic or environmental effects on style development. Short styles in functionally male flowers represent proportional reduction maintaining style as approximately 25-40% of total pistil length despite overall pistil abbreviation (total pistil 0.8-1.8 mm with short style 0.25-0.5 mm and reduced ovary and stigma), while short styles in staminate flowers show even greater reduction (total pistil 0.2-0.8 mm with vestigial style 0.1-0.3 mm) reflecting extreme pistil suppression. The functional implications of short styles relate to reduced female reproductive capacity= short styles in functionally male flowers contribute to functional male sterility through abbreviated pollen tube pathway, though style shortening is less critical than stigma non-receptivity or ovule abortion in determining reproductive failure short styles in staminate flowers represent vestigial structures with no reproductive function, serving only as developmental remnants of pistil organization rare short styles in otherwise-hermaphrodite flowers may impair fertilization efficiency through suboptimal pollen tube guidance or support, though effects are typically modest if transmitting tract tissue remains functional. Short style length requires correspondingly shorter pollen tube growth distances (0.8-1.5 mm total from stigma surface through style and ovary to ovules in functionally male flowers versus 1.2-2.5 mm in hermaphrodite flowers), potentially requiring 24-48 hours for pollen tube transit versus 48-96 hours in hermaphrodite flowers, though this theoretical advantage is negated by stigma non-receptivity and ovule non-viability in functionally male and staminate contexts.
Short_Style represents reduced style length characteristic of functionally male and staminate flowers and must be distinguished from Medium_Style (typical hermaphrodite length 0.4-0.7 mm) and Long_Style (enhanced length >0.7 mm in functionally female/pistillate flowers). Short styles occur in context of overall pistil reduction and contribute to reduced female function, though style shortening itself is less determinative of reproductive failure than stigma and ovule defects. Sub-classification into Very_Short and Reduced_Short reflects gradations within the short style range. Understanding short styles is relevant for identifying functionally male and staminate flowers, for assessing pistil developmental variation, and for understanding the developmental coordination of pistil component reduction in female-sterile flower types.
abbreviated style
kurzer Griffel
reduced style
reduzierter Stempelgriffel
verkürzter Stylus
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 93-98 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109.
Short Style
The shoulder description of Cabernet Franc documents moderately developed shoulders at the proximal 2-3 rachis nodes — the shoulder branches extend 15-25 mm laterally (less than the 30-50 mm extension that would constitute a true wing), creating a gradual widening of the upper cluster region that, combined with the tapering distal end, produces the conical-to-cylindrical-conical cluster shape characteristic of Cabernet Franc. These moderate shoulders are morphologically intermediate between the absent shoulders of Pinot Noir (where the proximal branches are no longer than mid-cluster branches, producing the uniform-width cylindrical outline) and the strongly developed shoulders of its offspring Cabernet Sauvignon (where the most proximal branch is sufficiently elongated to constitute a true wing, creating a distinctly conical cluster with a prominent lateral extension).
A cluster description component that records the presence and development of shoulders — the moderately elongated primary lateral branches at the proximal end of the cluster that are shorter than true wings but sufficiently developed to create a widened upper cluster region — documenting the degree of shoulder prominence, the angle of shoulder branches relative to the rachis, and the contribution of shoulder development to the overall cluster shape (particularly the distinction between cylindrical clusters with no shoulders and conical clusters whose taper results from shoulder-mediated proximal widening).
Shoulder_Description addresses the morphological transition zone between wingless and winged clusters — shoulders represent the moderate development of proximal lateral branches that creates cluster shape variation without the dramatic architectural departure of true wing formation. The distinction between shoulders and wings is one of degree rather than kind — both result from the differential elongation of proximal versus distal lateral branches along the rachis, with the classification based on the relative length of the proximal branches compared to the main cluster body — branches extending less than approximately 25% of the main cluster length are typically scored as shoulders, while branches extending more than 25% are scored as wings: this threshold is inevitably somewhat arbitrary, and intermediate forms exist that could be classified as either prominent shoulders or small wings depending on the observer. The functional significance of shoulders relates primarily to cluster shape and compactness — well-developed shoulders increase the width of the upper cluster region, potentially creating an internal space between the shoulder branches and the main cluster body that improves air circulation and reduces Botrytis susceptibility compared to compact, shoulderless clusters where berries are tightly packed from the proximal to distal end. In table grape production, shoulder development is an aesthetic criterion — well-shouldered clusters create the visually appealing pyramidal form preferred in premium table grape markets, and shoulder branch management (including manual repositioning of shoulders during cluster development) is practiced in high-value table grape production systems.
Traubenschulter-Beschreibung
cluster shoulder record
obere Traubenverbreiterung
proximal branch development
proximale Rappen-Verzweigung
upper cluster widening
Shoulder Description
Shriveling quality control in export Red Globe shipments from Chile to China requires 95-98% relative humidity maintenance throughout the 28-35 day transit via refrigerated container, with continuous humidity monitoring and sulfur dioxide pad replacement at transshipment points to prevent the shriveling progression that begins with subtle shoulder dimpling (less than 3% weight loss, acceptable), advances through visible wrinkling (5-8% weight loss, quality defect), and culminates in severe collapse (greater than 15% weight loss, unmarketable) fruit arriving with greater than 5% of berries showing moderate shriveling faces rejection or price penalties of 25-40%, making humidity management as critical as temperature control for quality preservation.
Shriveling_Quality represents the damage state characterized by visible water loss from grape berries resulting in loss of turgidity, skin wrinkling, and reduction in berry volume. Shriveling reflects the negative water balance where evaporative loss from the berry surface exceeds water supply through the pedicel, progressively depleting berry water content and causing the characteristic wrinkled, raisin-like appearance. Unlike the controlled dehydration deliberately induced for raisin production or certain wine styles (amarone, passito), quality-relevant shriveling addresses the unintended water loss that compromises fruit quality for table grape markets and most wine production. Shriveling can occur at multiple stages including pre-harvest (from excessive heat, water stress, or vascular disruption), at harvest (if picking occurs during hot conditions), and postharvest (from inadequate humidity control during storage and transport). The susceptibility to shriveling depends on skin properties affecting water permeability, berry surface-to-volume ratio, and environmental conditions particularly vapor pressure deficit. Small berries with high surface-to-volume ratios shrivel more rapidly than large berries. Shriveling accelerates once initiated because skin folding reduces the efficiency of the cuticle as a water barrier. Berry shrivel disorder, a distinct physiological condition, causes accelerated shriveling of specific berries within clusters due to vascular dysfunction, distinguished from environmental shriveling by its patchy distribution and occurrence before normal maturity. Assessment of shriveling employs visual classification of severity from slight skin dimpling through moderate wrinkling to severe raisin-like collapse, with quantification of the percentage of affected berries.
Shriveling_Quality addresses the water loss damage state, distinct from the processes causing dehydration and from the disposition toward shriveling susceptibility. The category encompasses both environmental shriveling (excessive water loss under challenging conditions) and physiological shrivel disorder (berry-specific vascular dysfunction). The threshold distinguishing normal late-season concentration from quality-defect shriveling depends on intended use slight shriveling may be acceptable or even desirable for certain red wine styles while being unacceptable for table grapes. Weight loss percentage provides a quantitative measure that complements visual severity classification.
Beerenaustrocknung
Einschrumpfung
Schrumpfung
Welkeerscheinung
berry shrinkage quality
water loss damage
wilting
Krasnow MN et al. (2010) Distinctive Symptoms Differentiate Four Common Types of Berry Shrivel Disorder in Grape. California Agriculture 64(3): 155-159 Bondada BR and Keller M (2012) Not All Shrivels Are Created Equal: Morpho-Anatomical and Compositional Characteristics Differ among Different Shrivel Types That Develop during Ripening of Grape (Vitis vinifera L.) Berries. American Journal of Plant Sciences 3(7): 879-898
Shriveling Quality
Silkiness quality in mature Volnay Premier Cru Pinot Noir (silkiness rating 9/10) exemplifies the textural ideal where 12 years of aging has polymerized tannins to seamless silk-like refinement, with the wine gliding effortlessly across the palate without any tactile friction, complementing the delicate red fruit and forest floor aromatics with textural elegance that makes each sip a refined sensory pleasure, demonstrating how silkiness contributes to the complete quality experience beyond flavor alone.
Silkiness quality is a texture perception quality that represents the smooth, lustrous, friction-free tactile impression reminiscent of silk fabric, characterizing the refined, elegant mouthfeel where wine glides across oral surfaces without resistance, grain, or textural interruption, creating the polished impression associated with premium wine quality. Within the BFO framework, this quality specifically addresses the smooth-lustrous dimension of texture perception, distinguishing silky character from merely smooth (lacking the lustrous quality) or velvety (having more plush, dense character) while recognizing silkiness as one of the most positively valued textural descriptors in wine evaluation. The ontological significance lies in silkiness serving as a universal quality marker across wine styles, where silky texture indicates successful tannin refinement, appropriate polysaccharide contribution, and overall compositional harmony that creates the effortless, elegant mouthfeel associated with fine wine regardless of body level or flavor profile. The perceptual characteristics of silkiness include the sensation of effortless gliding across oral surfaces, complete absence of grain or roughness, subtle lustrous quality suggesting refinement, and the overall impression of elegance and polish that makes consumption pleasurable at the tactile level. The compositional determinants of silkiness include refined, well-polymerized tannins (providing structure without friction), adequate polysaccharide content (mannoproteins contributing lubrication), appropriate glycerol level (enhancing smooth glide), balanced acidity (avoiding harsh edges), and the overall integration of components creating seamless texture. The variety and style associations of silkiness span red and white wines, with silky texture valued in Pinot Noir (where variety's naturally fine tannins enable silky expression), premium Chardonnay (where lees contact contributes silk-like smoothness), aged wines of any variety (where tannin polymerization develops silkiness), and any wine where elegance is a style goal. Understanding silkiness quality enables premium quality recognition based on textural refinement, aging assessment where silkiness indicates maturity, and quality evaluation incorporating silky texture as positive attribute across wine categories.
Silkiness_Quality addresses the smooth-lustrous dimension of texture perception, representing one of the most valued textural attributes in wine evaluation. This quality indicates refinement and elegance, serving as a universal quality marker across wine styles. The class supports premium quality recognition and aging assessment, with silkiness development indicating successful maturation and compositional harmony.
Seidigkeit
elegant mouthfeel
elegantes Mundgefühl
glänzende Glätte
lustrous smoothness
refined texture
seidige Textur
silk-like texture
Gawel R et al. (2000) A 'Mouth-feel Wheel': terminology for communicating the mouth-feel characteristics of red wine. Australian Journal of Grape and Wine Research 6(3): 203-207 Chira K et al. (2011) Chemical and sensory evaluation of Bordeaux wines according to grape quality and tannin structure. Journal of the Science of Food and Agriculture 91(10): 1748-1754
Silkiness Quality
The silt content of loess-derived vineyard soils in the Kaiserstuhl (Baden, Germany) documents high silt proportions of 55-70% — the wind-deposited loess (Pleistocene aeolian sediment) produces a distinctively silty texture class (silt loam to silty clay loam) with the highest plant-available water capacity of any common vineyard soil type (200-280 mm per meter of soil depth), sustaining vine water supply through extended dry periods without irrigation. The high silt content creates exceptional soil fertility and water availability that, combined with the warm mesoclimate of the Kaiserstuhl volcanic outcrop (one of the warmest sites in Germany), produces vigorous vine growth requiring careful canopy management to avoid excessive shading — the natural fertility of the loess soils contrasts with the infertile, growth-limiting soils (gravel, slate, limestone) that characterize most German premium vineyard sites, making the Kaiserstuhl terroir atypical within the German viticultural landscape.
A soil texture description component that records the proportion of silt-sized particles (0.002-0.05 mm diameter in the USDA/international system, subdivided into fine silt 0.002-0.02 mm and coarse silt 0.02-0.05 mm) in the fine earth fraction of the vineyard soil, expressed as a percentage by weight, with silt content influencing water-holding capacity (silt retains more plant-available water per unit mass than either sand or clay), soil structural stability (high silt content increases susceptibility to surface crusting and erosion), and the overall textural character that determines the soil's behavior as a growth medium for vine roots.
Silt_Content records the intermediate particle size fraction whose physical behavior provides a balance between the macropore-dominated drainage of sand and the micropore-dominated water retention of clay. Silt particles occupy the size range that produces pore spaces of approximately 0.5-30 μm — pores that are small enough to retain water against gravitational drainage (unlike the large pores between sand particles) but large enough to release water to plant roots at relatively low suction tensions (unlike the submicroscopic pores between clay particles that retain water so tightly that it is unavailable to plants): this intermediate pore size distribution gives silt-rich soils the highest plant-available water capacity per unit of soil volume, making silty soils the most favorable for vine water supply in non-irrigated viticulture — but also potentially the most vigorous growth environments, since abundant water availability promotes vegetative growth that may be undesirable for premium wine production. The structural instability of silt-rich soils is a significant practical concern — silt particles lack the cohesive bonding capacity of clay (which forms aggregates through electrochemical attraction) and the weight stability of sand (which resists displacement by raindrop impact), making silt-rich soils highly susceptible to surface crusting (the formation of a dense, impermeable surface layer under raindrop impact), wind erosion (loess soils are the most wind-erodible soil type), and water erosion (silt particles are easily detached and transported by surface runoff).
Schluff-Fraktion
Schluffanteil-Messung
Schluffgehalt
intermediate particle proportion
silt fraction record
silt percentage measurement
White RE (2003) Soils for Fine Wines, Oxford University Press Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873 Soil Survey Staff (2014) Keys to Soil Taxonomy, 12th edition, USDA-NRCS
Silt Content
A simple unbranched tendril at node 9 on a shoot of Vitis vinifera cv. Riesling in the Rheingau, Germany, measuring 18 cm mature length, consisting of a single continuous axis of 3.1 mm diameter at mid-length showing complete lignification in basal 8 cm with brown coloration and rigid texture, functional green tissue in distal 10 cm maintaining turgor and flexibility, well-developed vascular bundles visible through semi-translucent epidermis, complete cuticle formation evident from glossy surface texture, mechanosensory cells distributed along the entire length demonstrated through rapid coiling response (within 7 minutes) to experimental contact at various positions, active circumnutation with 135-degree sweep amplitude exploring approximately 310 cubic centimeters, making successful firm attachment to horizontal trellis wire through seven tight contact coils and ten free coils supporting 420 grams shoot weight without strain, occurring in a Riesling clone where 70-75% of tendrils develop as mature unbranched structures while 25-30% show bifurcation, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing complete functional development of simple unbranched architecture.
An unbranched tendril that has completed development and achieved functional maturity as a single-axis structure without branching in Vitis vinifera, characterized by attainment of cultivar-typical and position-specific mature length of 10-22 cm, completion of tissue differentiation including full vascular development with mature xylem and phloem, functional epidermis with complete cuticle formation, differentiated mechanosensory apparatus with responsive contact-detecting cells distributed along the tendril surface, adequate structural reinforcement through appropriate lignification for load-bearing function, and active physiological status maintaining circumnutation movements and coiling response capacity, representing permanent unbranched architecture rather than incomplete juvenile development. Simple unbranched tendrils represent the mature expression of genetic programming for single-axis tendril architecture in cultivars or individual tendrils where branching initiation does not occur even under optimal developmental conditions, or represent developmental outcome where resource availability or environmental conditions during early organogenesis did not support lateral meristem activation but permitted completion of single-axis development to functional maturity. These tendrils provide full climbing functionality through single robust attachments, with concentrated resource investment in one axis potentially yielding stronger individual structure compared to resource-partitioned branched tendrils, effective spatial exploration within the sweep volume of a single circumnutation arm, and metabolically efficient climbing support requiring lower total resource investment than branched architectures.
Simple unbranched tendrils are distinguished from juvenile unbranched tendrils by completion of development and achievement of functional maturity, and from branched tendrils by single-axis architecture. Assessment requires evaluation of multiple maturity indicators including length achievement relative to cultivar norms, tissue differentiation completion, mechanosensory function, structural competence, and lignification status, with simple unbranched classification appropriate only when tendril shows no signs of continued growth or potential for delayed branch initiation. Some cultivars show strong genetic determination of unbranched architecture with very high frequencies of mature simple tendrils, while others show facultative patterns where simple versus branched development depends on vigor and environmental conditions during early organogenesis. Simple unbranched tendrils represent the ancestral default tendril architecture, with branching representing derived elaboration increasing architectural complexity and functional capacity at the cost of increased resource requirements.
ausgereifte einfache Ranke
developed simple tendril
functional single-axis tendril
funktionsfähige einachsige Kletterorgan
mature unbranched tendril
reife unverzweigte Ranke
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 90-119 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 86-108 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 119-154.
Simple Unbranched Tendril
A single-cluster fertile shoot on Vitis vinifera cv. Pinot Noir from node 5 in Côte de Nuits, 135cm length with 16 leaves totaling approximately 2240 cm² leaf area, bearing one perfectly developed cluster with 78 berries showing excellent fruit set (estimated 48%), normal cluster development progressing to veraison (BBCH 81), exceptional leaf-to-fruit ratio (22.8 cm² per gram assuming 295g final cluster weight), representing deliberate quality-focused single-cluster strategy (second clusters removed during fruit set on shoots initially bearing two clusters), with excellent photosynthetic support enabling complete phenolic maturation, optimal berry development, superior aromatic complexity, and wine quality outcomes justifying yield sacrifice (vineyard average 3.8 kg per vine, approximately 38 hl/ha) through premium Burgundy pricing, this shoot exemplifying quality-first philosophy emphasizing terroir expression and wine character over productivity maximization.
A fertile reproductive shoot bearing exactly one well-developed cluster, characterized by successful fruit set and development in single inflorescence, shoot resources concentrated supporting single cluster maturation, typically excellent leaf-to-fruit ratios (15-25 cm² per gram) through undivided photosynthetic capacity, and contribution of moderate crop weight (180-380g typical) representing quality-oriented productive configuration balancing yield with superior fruit development potential. Single-cluster fertile shoots represent common configuration in quality-focused viticulture, arising through natural single-cluster differentiation patterns (many varieties showing single-cluster as common fruitfulness state), deliberate cluster thinning strategies reducing multi-cluster shoots to singles, or physiological cluster reduction leaving single successful cluster, with single-cluster configuration providing excellent conditions for balanced vine physiology, superior fruit composition, and consistent quality outcomes justifying lower per-shoot yields through quality benefits and premium pricing.
Single-cluster fertile shoots represent quality optimization in reproductive shoot management, with concentration of resources into single cluster development providing numerous benefits including superior leaf-to-fruit ratios (typically 15-25 cm² per gram versus 10-15 for multi-cluster shoots), reduced berry competition (larger berries, better skin-to-juice ratios, enhanced phenolic potential), improved cluster architecture (reduced compactness, better air circulation, lower disease pressure), optimal resource allocation (undivided photosynthetic capacity supporting complete maturation), and superior fruit composition (higher sugars, better phenolic development, enhanced aromatics). The single-cluster approach dominates ultra-premium wine production (Burgundy, Barolo, premium Napa Cabernet), reflects quality-first philosophies accepting yield sacrifice for character enhancement, and increasingly influences broader quality-oriented production recognizing compositional benefits. However, single-cluster strategies require economic justification through premium pricing, with bulk production typically retaining multiple clusters accepting quality tradeoffs for economic viability through higher yields.
Einzelbüschel-Fruchttrieb
eintraubiger fertiler Trieb
one-cluster fertile shoot
single-bunch productive shoot
solitary cluster fruiting shoot
Based on Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Reynolds et al. (1996) "Impact of cluster exposure on secondary metabolites in Cabernet Sauvignon grapes and wine" Journal of Wine Research 7=71-90.
Single Cluster Fertile Shoot
A single cluster primary bud at node 4 on Vitis vinifera cv. Sauvignon Blanc, dissection revealing one well-developed inflorescence primordium containing 165 flower initials, plus vegetative shoot primordia with 9 leaf initials, developing into a shoot bearing one compact cluster of 82 berries (50% fruit set), 14 fully expanded leaves providing 720 cm² leaf area, resulting in favorable 8.8 cm²/berry leaf area-to-fruit ratio supporting complete fruit maturation to 22.5° Brix with balanced acidity and varietal aroma development, without requiring cluster thinning intervention to achieve quality targets.
A primary reproductive bud containing exactly one differentiated inflorescence primordium along with vegetative shoot structures, representing moderate fruitfulness typical of mid-range node positions (nodes 3-5 or 10-12) or cultivars with naturally lower cluster numbers per shoot, characterized by single flower cluster development upon budbreak producing one bunch per shoot, adequate but not maximal reproductive potential, and balanced resource allocation between vegetative growth and fruit production. Single cluster primary buds provide reliable baseline crop production without excessive fruit load, typically associated with moderate vine balance, reduced cluster thinning requirements compared to multi-cluster shoots, and favorable fruit composition development through balanced source-sink relationships where photosynthetic capacity adequately supports single cluster maturation without resource limitation or competition.
Single cluster primary buds represent a middle ground between non-fruiting vegetative buds and highly fruitful multi-cluster buds, often optimal for balanced production in moderate-vigor vineyards where two-cluster shoots might create excessive crop load while purely vegetative shoots waste productive capacity. Node position effects show single-cluster buds concentrated at transitional positions between basal non-fruiting zones (nodes 1-3) and peak fruitfulness mid-cane zones (nodes 6-9), and again at distal positions (nodes 11-14) where declining fruitfulness reduces from two clusters back to one. Some cultivars (e.g., Nebbiolo, Barbera) characteristically produce single-cluster primaries even at optimal node positions, while others (e.g., Pinot Noir, Chardonnay) rarely form single-cluster buds at mid-cane positions under favorable differentiation conditions.
Ein-Trauben-Primärknospe
Einzeltraubenknospe
one-cluster primary bud
single-bunch primary bud
unicluster primary bud
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Dunn & Martin (2000) "A functional association in Vitis vinifera L. cv. Cabernet Sauvignon between the extent of primary branching and the number of flowers formed per inflorescence" Australian Journal of Grape and Wine Research 6=88-92.
Single Cluster Primary Bud
A single-cluster primary reproductive shoot on Vitis vinifera cv. Pinot Noir from node 6 in Burgundy, 132cm length with 15 leaves totaling approximately 2100 cm² leaf area, bearing one well-developed cluster with 86 berries, estimated shoot crop 284g at maturity, representing natural single-cluster fruitfulness characteristic of this Pinot Noir clone and typical Burgundian practice accepting or enhancing single-cluster loads through thinning (removing second clusters where present, though this shoot naturally single-cluster from differentiation), excellent leaf area-to-fruit ratio (19.4 cm² per gram) supporting complete phenolic maturation and aromatic development crucial for premium Pinot Noir quality, with single-cluster configuration contributing to superior fruit composition (higher skin-to-juice ratio in smaller crops, enhanced phenolic extraction, improved aromatic complexity) justifying lower yields through quality outcomes and premium pricing.
A primary reproductive shoot bearing exactly one inflorescence, characterized by single cluster load regardless of cluster size, developmental origin from either single-cluster bud differentiation (typical for many varieties at certain node positions) or cluster reduction from initial multiple-cluster state through physiological drop, manual thinning, or stress-induced abortion, and function providing moderate crop contribution with excellent potential for balanced leaf-to-fruit ratios and superior fruit quality through concentration of shoot resources into single cluster development. Single-cluster primary reproductive shoots represent common configuration across many varieties and positions, arising naturally in cultivars with predominant single-cluster fruitfulness patterns (many Pinot Noir clones, certain Nebbiolo selections, some Syrah clones), through intentional cluster thinning reducing multiple-cluster shoots to single clusters, or via natural cluster reduction through environmental stress, disease, or physiological limitations affecting secondary cluster retention.
Single-cluster primary reproductive shoots occur through natural differentiation patterns (many varieties showing single-cluster as modal or common fruitfulness state at certain positions), deliberate management (cluster thinning to single clusters as quality strategy), or stress-induced reduction (physiological drop, disease, environmental stress eliminating secondary clusters initially present). The single-cluster configuration provides excellent opportunity for balanced vine physiology and superior fruit quality through favorable leaf-to-fruit ratios, reduced competition among berries, enhanced resource availability per berry, and often improved cluster architecture (reduced compactness, better air circulation, lower disease pressure). Quality-focused wine production frequently employs single-cluster strategies either accepting natural single-cluster patterns or actively thinning to achieve single-cluster loads, recognizing quality benefits despite yield sacrifice. However, single-cluster approaches require economic justification through premium pricing, with bulk production systems typically retaining multiple clusters accepting quality tradeoffs for economic viability through higher yields.
Eintraubentrieb
einzeltraubiger Fruchttrieb
one-cluster primary shoot
single-bunch fruiting shoot
solitary cluster shoot
Single Cluster Primary Reproductive Shoot
A Pinot Noir vine in Romanée-Conti grand cru trained to guyot simple bears a Single_Guyot_Role with a single arched 8-bud cane, the restrictive pruning mandated by appellation regulations reflecting the site's capacity to produce concentrated fruit from minimal crop—yields averaging 25-30 hl/ha from this elite configuration. A Chardonnay vine in Montrachet grand cru serves in a Single_Guyot_Role, the single-cane restriction part of the stringent production parameters underlying these legendary white wines. A young Pinot Noir vine in its third leaf in Oregon bears a Single_Guyot_Role appropriate to its establishment phase, with conversion to double guyot anticipated as the vine matures and can support greater crop load.
A guyot-trained vine role that inheres in a grapevine with one fruiting cane retained at pruning, extending in one direction from the vine head along the fruiting wire, plus a short renewal spur positioned to provide next year's replacement cane. Single guyot (guyot simple) represents the most restrictive cane-pruning configuration, limiting crop to one cane (typically 6-10 buds) per vine and creating maximum per-cluster concentration through controlled crop load. The single cane is typically arched or bent from vertical to horizontal at the fruiting wire, with the arc creating slight bud position elevation that some believe enhances fruitfulness. Single guyot is mandatory in many French grand cru appellations where concentration is paramount, and preferred for young vines, low-vigor sites, and production philosophies emphasizing intensity over volume. The asymmetrical structure (cane extending one direction only) requires attention to cane direction—alternating direction between adjacent vines or between rows to maintain balanced vineyard appearance and uniform mechanical access.
Single_Guyot_Role captures the most restrictive guyot variant, associated with elite production and sites where concentration trumps volume. The single-cane limitation reflects production philosophy as much as vine physiology—grand cru Burgundy vineyards could physiologically support more crop, but regulations mandate single guyot to enforce the low yields believed essential for grand cru quality. Cane positioning in single guyot creates characteristic arched or bent form, with the cane emerging vertically from the head before bending to horizontal along the wire this arc may be gentle or pronounced depending on regional tradition. Renewal spur management is critical—the spur must produce well-positioned, vigorous shoots suitable for next year's cane, requiring careful spur placement and sometimes additional dormant pruning adjustments if initial spur shoots are unsuitable.
Einzelstrecker
Guyot-Einfach
Halbbogen-Erziehung
einarmiger Guyot
guyot simple
one-cane guyot
single guyot
unilateral guyot
Galet P (2000) General Viticulture, Oenoplurimedia INAO (2020) Cahiers des charges AOC Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Jackson RS (2014) Wine Science, Academic Press
Single Guyot Role
A 4 cm single-node green cutting of Pinot Noir clone 777 from FPS nuclear stock bears a Single_Node_Cutting_Role, prepared by cutting 1.5 cm above and 2.5 cm below the node, treating with 1000 ppm IBA, and rooting under intermittent mist—enabling production of 15-20 propagules per meter of source cane versus 3-4 for standard three-node cuttings. A single-node hardwood cutting of SO4 rootstock serves in a Single_Node_Cutting_Role in a Danish hot callusing system, where controlled temperature (26-28°C) and humidity compensate for limited reserves to achieve commercial rooting success. A single-node cutting of 110 Richter bears a Single_Node_Cutting_Role in Australian nursery production, prepared on automated cutting machines and rooted in cellular trays with bottom heat and overhead misting.
A cutting role that inheres in a stem segment prepared with a single node bearing one bud plus associated internode tissue (typically 3-5 cm total length), representing the minimum viable propagation unit for grapevines and maximizing multiplication rate from limited source material. Single-node cuttings produce one shoot from the single bud while developing adventitious roots from the internode and nodal tissues, with the minimal tissue quantity reducing carbohydrate reserves available for establishment but maximizing the number of propagules obtainable from a given amount of source material. Single-node propagation is standard for green/softwood cuttings where maintaining turgor in minimal tissue is manageable under mist conditions, and is increasingly applied to hardwood propagation using controlled-environment systems (hot callusing facilities, rooting chambers with bottom heat) that compensate for limited reserves through optimized environmental conditions. The technique is particularly valuable for rapid multiplication of virus-tested nuclear stock, for research applications requiring many replicates from limited genotypes, and for commercial rootstock production seeking maximum multiplication efficiency.
Single_Node_Cutting_Role represents the intensive end of the multiplication efficiency spectrum, trading individual propagule vigor for maximum numbers from limited source material. The approach is economically attractive when source material value is high (virus-tested nuclear stock, newly introduced varieties, expensive licensed cultivars) and production systems can achieve adequate rooting success with minimal tissue. Single-node hardwood cutting propagation has expanded with hot callusing technology—controlled-temperature facilities inducing callus and root primordium formation before field or greenhouse planting. Quality parameters for single-node cuttings emphasize bud health (the single bud must be viable) and internode diameter (adequate tissue volume for rooting), with tight quality control essential since there is no redundancy if the single bud fails.
Einaugensteckling
Einknoten-Steckling
Einzelaugensteckling
nodal cutting
one-bud cutting
single-eye cutting
single-node cutting
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 350-355 Hunter JJ et al. (2004) Effect of cutting position on rooting, South African Journal of Enology and Viticulture 25: 26-32 Waite H, Morton L (2007) Hot water treatment, fumigation and their combination on grapevine cutting survival, Australian Journal of Grape and Wine Research 13: 99-109
Single Node Cutting Role
A single side unilateral permanent arm on Vitis vinifera cv. Gewürztraminer at vineyard boundary position, trunk positioned at property line extending single arm 88cm into vineyard interior with zero extension toward boundary side (no space available for any arm development in restricted direction), bearing 14 spurs along single arm supporting 4.6kg crop, representing pure unilateral architecture with no bilateral component present or intended, trunk positioning deliberately offset to property line to maximize available arm extension space in permitted direction, with complete vine capacity channeled into single structural element producing yields equivalent to interior bilateral vines through compensatory arm length and spur density.
A unilateral permanent arm extending exclusively in one direction with no development or retention of opposite-side arm, characterized by complete one-sided architecture, trunk positioned to facilitate single-direction extension, all fruiting capacity concentrated along one arm without any bilateral component, and deliberate or circumstantial design resulting in purely unilateral configuration. Single side unilateral permanent arms represent pure expression of unilateral architecture, distinguished from asymmetrical bilateral configurations that retain small opposite arm or from transitional unilateral situations anticipating future bilateral development, with single-side configuration being permanent intended architecture rather than intermediate stage or compromise with minor bilateral component.
Single side unilateral permanent arms represent the clearest expression of unilateral training, most commonly found at vineyard boundaries, row ends, or other constrained positions where directional extension in one direction is physically impossible or prohibited (property lines, buildings, roadways, other obstructions). The single-side designation emphasizes complete absence of bilateral component, distinguishing these arms from asymmetrical bilateral configurations that retain small opposite arm or from transitional situations where opposite arm development is planned but not yet implemented. Management of single-side unilateral arms focuses on maximizing productivity from available arm length through appropriate spur density, maintaining arm vigor through careful pruning avoiding excessive crop load concentration, and ensuring structural soundness recognizing that all mechanical stress concentrates in single arm without load distribution benefit of bilateral architecture.
ausschließlich einseitige Arm
complete single arm
exclusively one-sided arm
pure unilateral arm
reine unilaterale Arm
Single Side Unilateral Permanent Arm
A single vertical cordon on Vitis vinifera cv. Barbera in high-density planting (1.0m × 2.5m spacing) in Piedmont, Italy, extending 185cm vertically from short trunk, bearing 14 two-bud spurs distributed along length, shoots from spurs trained laterally into catch wire system creating narrow vertical canopy profile fitting within 1-meter vine spacing, representing single vertical cordon architecture necessitated by narrow spacing preventing bilateral cordon development, with single vertical structure supporting adequate fruiting capacity (approximately 26 shoots, 38-44 clusters, 5.5-6.5kg crop) within space constraints while maintaining manageable canopy architecture and acceptable fruit quality for regional Barbera production standards.
A vertical cordon configuration featuring one upward-extending cordon per vine, characterized by single vertical axis bearing fruiting spurs along its length, trunk transitioning directly to single vertical cordon without branching or paired structures, and architectural simplicity with all fruiting capacity concentrated along single vertical element. Single vertical cordons represent minimal vertical cordon configuration, appropriate for high-density plantings where lateral space constraints preclude paired vertical cordons, traditional systems employing single-trunk single-cordon architecture, or transitional training during vertical cordon establishment before potential addition of second vertical cordon when vine vigor permits.
Single vertical cordons are relatively uncommon in modern viticulture, with specific applications primarily in high-density plantings where space constraints require vertical rather than lateral architecture, and where narrow spacing (often 1-1.5m within rows) prevents multiple vertical cordons per vine. The single vertical configuration concentrates vine capacity into one structural element, requiring appropriate vigor management to prevent excessive growth in single axis while ensuring adequate development to support economically viable crops. Some traditional European systems employ single vertical cordons for historical reasons or specific training philosophies, while others use single vertical configuration as transitional stage during establishment, potentially adding second vertical cordon later if vine vigor and space permit. Management challenges include maintaining even spur vigor along vertical gradient (apical dominance effects) and preventing structural weakness at trunk-cordon transition bearing full vine weight.
Einzelhochkordon
einfache vertikale Kordon
mono-vertical cordon
single-column cordon
solitary vertical cordon
Single Vertical Cordon
The single vineyard name To Kalon Vineyard in the Oakville AVA of Napa Valley identifies a historic 89-acre site originally planted in 1868 by Hamilton Walker Crabb, with the name (Greek for the beautiful or the highest good) appearing on labels from multiple producers who source grapes from this site — Robert Mondavi Winery, Opus One, Schrader Cellars, and others each produce To Kalon-designated wines that command significant price premiums (typically 200-500% above non-vineyard-designated wines from the same producers) based on the site's established reputation for producing Cabernet Sauvignon of exceptional concentration and complexity from deep alluvial gravel soils.
A vineyard designation that identifies a wine as produced exclusively or predominantly from grapes grown in a single, named vineyard site defined by specific geographic boundaries, used across wine-producing countries as the generic designation for site-specific wines regardless of the national terminology (cru, Lage, contrada, etc.) applicable in the producing region.
Single_Vineyard_Name is the generic cross-cultural class for site-specific vineyard designations that the more specific national subclasses (Lieu_Dit_Name, Cru_Name, Lage_Name, Ried_Name, Climat_Name, Contrada_Name, Pago_Name) implement within their respective regulatory traditions. The regulatory requirements for single vineyard designation vary dramatically between jurisdictions: in France and Germany, vineyard names must correspond to officially delimited and registered parcels, while in the United States, Australia, and many New World countries, the definition and boundaries of a single vineyard name are determined by the proprietor with minimal regulatory oversight beyond geographic truthfulness. The minimum percentage of grapes from the named vineyard also varies — 100% in some French appellations, 95% in Germany, 85% in the United States — creating different standards of site-specificity under the common single vineyard concept.
Einzellagenbezeichnung
Einzelweinbergsname
benannter Weinberg
estate vineyard name
named vineyard site
single vineyard designation
Robinson J and Harding J (2015) The Oxford Companion to Wine 4th Ed, Oxford University Press TTB (2023) Wine Labeling Regulations 27 CFR Part 4 EU Delegated Regulation 2019/33 Art. 55 on labeling of smaller geographic units
Single Vineyard Name
The sinus description of Chardonnay documents a characteristic combination — a lyre-shaped petiolar sinus that is wide open with a distinctive U-shaped base and slightly divergent sides (OIV 079:1-3), moderately deep upper lateral sinuses with open V-shaped form (OIV 070:5, OIV 080:2), and shallow to absent lower lateral sinuses (OIV 082:1) — this particular sinus architecture, with the open petiolar sinus as its most distinctive feature, is immediately recognizable and separates Chardonnay from its parent Pinot Noir (which has a strongly overlapping petiolar sinus, OIV 079:7) despite the overall similar five-lobed blade shape and size, demonstrating that sinus morphology can distinguish cultivars that share the same overall leaf outline.
A mature leaf description component that documents the morphological characteristics of the sinuses (indentations between lobes) of the fully expanded leaf blade, recording the shape, depth, and degree of overlap of the petiolar sinus and the upper and lower lateral sinuses according to OIV descriptors 076-082, with sinus morphology providing some of the most diagnostically powerful and genetically stable characters in the entire ampelographic descriptor set for cultivar identification in Vitis vinifera.
Sinus_Description addresses a suite of characters that experienced ampelographers consistently rate among the most valuable for cultivar identification — petiolar sinus shape in particular is frequently cited as the single most diagnostically powerful leaf character in the OIV system, combining high genetic stability (minimal environmental modification), high discrimination (clearly different between most cultivar pairs), and practical observability (visible without magnification on any mature leaf). The three sinus types — petiolar (at the base of the blade where the petiole attaches), upper lateral (between the terminal and upper lateral lobes), and lower lateral (between the upper and lower lateral lobes) — are independently variable and provide orthogonal diagnostic information: a cultivar's petiolar sinus may be overlapping while its upper lateral sinuses are open, or vice versa, and this independent variation creates a multi-dimensional character space that enables fine-scale cultivar discrimination within groups sharing the same overall blade shape. The morphological vocabulary for sinus description has been refined over two centuries of ampelographic practice — the OIV system recognizes open, closed, and overlapping sinus classes with further subdivision by the shape of the sinus floor (U-shaped, V-shaped, lyre-shaped) and the configuration of the overlapping lobes when present.
Blattbuchten-Charakterisierung
Buchten-Beschreibung
Interlobar-Einbuchtungs-Beschreibung
interlobar indentation description
leaf sinus characterization
sinus morphology record
Sinus Description
The site description for the Hill of Grace vineyard (Eden Valley, South Australia, 34.63°S, 139.05°E) documents a northeast-facing hillside parcel at 380-410 m elevation on Cambrian schist and gneiss parent material, with a slope gradient of 5-12% providing natural drainage and cold air flow away from the vine rows — this site-specific combination of elevation (providing cooler temperatures than the adjacent Barossa Valley floor at 250-300 m), aspect (northeast orientation capturing morning sun while avoiding excessive afternoon heat exposure), and parent material (producing the shallow, well-drained, mineral-rich soils characteristic of the Eden Valley sub-region) creates the mesoclimatic conditions that distinguish Eden Valley Shiraz (elegant, cooler-climate expression with 13.5-14.5% alcohol) from the warmer Barossa Valley floor Shiraz (fuller, riper expression with 14.5-15.5% alcohol) grown only 10-15 km distant but 100-150 m lower in elevation.
A terroir information component that documents the physical geographic characteristics of the specific vineyard site — location, coordinates, elevation, aspect, slope, and landscape position — providing the spatial reference framework that determines the mesoclimate, radiation balance, drainage pattern, and cold air movement affecting vine performance at the site level, and enabling the georeferenced integration of all site-specific environmental data within geographic information systems.
Site_Description provides the geographic foundation for all terroir characterization — the physical location and topographic setting of a vineyard determine the mesoclimate (the local climate conditions differing from the regional macroclimate due to topographic effects), the soil parent material (determined by the underlying geology and geomorphological history), the drainage and water movement patterns (determined by slope, aspect, and landscape position), and the exposure to frost, wind, and extreme weather events. The precision of site description has been transformed by modern geospatial technologies — GPS positioning provides meter-level accuracy for vineyard boundary delineation, digital elevation models (DEMs) enable detailed calculation of slope, aspect, and potential solar radiation at every point in the vineyard, and remote sensing platforms (satellite, aerial, drone-mounted) provide spatial data on canopy development, water status, and yield variation at resolutions of 0.5-5 m: these technologies have enabled the quantitative characterization of within-vineyard spatial variation that underpins precision viticulture — the management philosophy that recognizes and responds to the heterogeneity of vine performance across a single vineyard by varying management inputs (irrigation, nutrition, canopy management, harvest timing) according to the spatial pattern of vine needs.
Standort-Beschreibung
Weinberg-Lage-Charakterisierung
geographic site record
geographische Standort-Dokumentation
physical site documentation
vineyard site characterization
van Leeuwen C and Seguin G (2006) The concept of terroir in viticulture, Journal of Wine Research 17: 1-10 Jones GV et al. (2004) Spatial analysis of climate in winegrape growing regions in the western US, American Journal of Enology and Viticulture 55: 304-316 Bramley RGV (2005) Understanding variability in winegrape production systems — 2. Within vineyard variation in quality over several vintages, Australian Journal of Grape and Wine Research 11: 33-42
Site Description
Sultana (Thompson Seedless) berries destined for table grape markets require size qualities meeting minimum standards of 16 mm equatorial diameter and 4.5 g fresh mass, achieved through gibberellic acid application at fruit set (10-20 ppm) and post-véraison (25-40 ppm), combined with cluster thinning to reduce sink competition and enable maximum cell expansion in remaining berries.
Size_Quality represents the category of morphological qualities that quantify the spatial extent and mass of grapevine organs and tissues. Size qualities are fundamental to grapevine characterization because organ dimensions directly influence physiological function, yield potential, and fruit composition. This category encompasses four measurement domains: Linear_Dimension_Quality capturing one-dimensional extents such as length, width, and thickness Area_Quality addressing two-dimensional surface measurements Volume_Quality representing three-dimensional spatial occupation and Mass_Quality quantifying the amount of matter. Size qualities exhibit strong genetic control with heritability estimates typically ranging from 0.5-0.8 for key parameters like berry weight and leaf area, yet environmental modulation produces substantial phenotypic plasticity. Water availability profoundly affects size qualities, with deficit irrigation reducing berry size by 20-40% through limitation of cell expansion during the post-véraison period. Temperature influences size through effects on cell division rates and expansion duration. Size qualities scale allometrically across organs and relate to functional capacity: larger leaves capture more light but require greater hydraulic supply smaller berries concentrate skin-derived compounds relative to juice volume. In viticulture, size quality assessment informs yield estimation, irrigation scheduling, harvest planning, and quality-based grape pricing. Instrumental methods including digital calipers, image analysis, and displacement volumetry provide objective size measurements, while traditional ampelography employs categorical scales (very small, small, medium, large, very large) referenced to cultivar standards.
Size_Quality serves as the parent class for all dimensional measurements in the VVO. The distinction between Linear_Dimension_Quality, Area_Quality, Volume_Quality, and Mass_Quality reflects fundamental differences in measurement dimensionality and appropriate units. Size qualities should not be confused with size-related dispositions (e.g., capacity for size increase) or size-determining processes (e.g., cell expansion). The VVO follows SI units as the primary measurement system while accommodating traditional viticultural units where established in practice.
Ausmaßqualität
Dimensionsqualität
Größenmerkmal
Größenqualität
dimensional quality
extent quality
magnitude quality
size attribute
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 153-189 Dai ZW et al. (2011) Ecophysiological, Genetic, and Molecular Causes of Variation in Grape Berry Weight and Composition: A Review. American Journal of Enology and Viticulture 62(4): 413-425
Size Quality
Skin anthocyanin concentration at harvest ranges from 200-500 mg/100g fresh weight skins in lightly pigmented cultivars such as Pinot Noir and Grenache to 1200-2500 mg/100g in deeply colored types such as Tannat, Alicante Bouschet, and Petite Sirah. Cabernet Sauvignon and Merlot show intermediate concentrations of 600-1000 mg/100g under favorable conditions. Concentration is determined by both anthocyanin per cell (vacuolar loading) and cells per unit skin area (hypodermal cell layers, typically 2-6 layers containing anthocyanins). Wine color extraction of 30-60% of skin anthocyanins produces young wine concentrations of 200-1500 mg/L depending on cultivar and maceration technique. Skin anthocyanin concentration shows heritability of h² = 0.65-0.80 with substantial environmental modulation by temperature, light exposure, and water status.
An anthocyanin accumulation disposition that is the inherited capacity of a Vitis vinifera plant to achieve specific total anthocyanin concentrations in grape berry skin tissue at harvest maturity, determined by biosynthetic rate, vacuolar storage capacity, degradation rate, and skin cell density, directly affecting potential wine color intensity.
Skin anthocyanin concentration is the primary determinant of potential wine color intensity, integrating biosynthetic capacity with the anatomical and developmental factors affecting pigment density in skin tissue. The trait differs from total berry anthocyanin, as skin thickness and cell density vary independently of biosynthetic rate. Climate change is negatively affecting skin anthocyanin concentration in many regions through high temperature inhibition of biosynthesis and accelerated degradation. Selection for high concentration combined with temperature tolerance is a breeding priority for warm climate viticulture.
Beerenhaut-Anthocyankonzentration
Farbstoffkonzentration
Schalenanthocyangehalt
anthocyanin content trait
pigment concentration disposition
skin color intensity
Roby G et al. (2004) Berry size and vine water deficits as factors in winegrape composition: anthocyanins and tannins. Australian Journal of Grape and Wine Research 10:100-107 Sadras VO, Moran MA (2012) Elevated temperature decouples anthocyanins and sugars in berries of Shiraz and Cabernet Franc. Australian Journal of Grape and Wine Research 18:115-122
Skin Anthocyanin Concentration Disposition
The skin color description of Gewürztraminer documents rose-pink skin (OIV 225:2) at full maturity — the pale copper-pink coloration results from low-level anthocyanin accumulation in the hypodermis, producing sufficient pigment to create a visible pink-bronze hue but far less than the intense anthocyanin concentrations of deeply pigmented cultivars, with total skin anthocyanin of 50-150 mg/kg compared to 1,500-2,500 mg/kg in Cabernet Sauvignon. This rose skin color classifies Gewürztraminer in the ambiguous zone between white and red cultivars — it is vinified as a white wine (without skin maceration) in Alsace, Germany, and most New World regions, but its visible pigmentation means that extended skin contact produces distinctly orange-tinted wines, and the cultivar is classified as a grey variety (Grauer Traminer) in traditional German ampelographic nomenclature.
A berry color description component that records the color of the berry skin (exocarp) at harvest maturity according to OIV descriptor 225, scored on a scale from green-yellow (state 1) through rose (state 2), red (state 3), grey (state 4), dark red-violet (state 5), to blue-black (state 6), documenting the externally visible pigmentation that is the primary visual identifier of grape cultivars and the source of anthocyanin pigments extracted during red and rosé winemaking.
Skin_Color_Description documents the character that is simultaneously the most commercially important, the most visually conspicuous, and the best genetically characterized of all ampelographic descriptors. The six-state OIV scale captures the major color categories but substantially simplifies the continuous variation in skin pigmentation — within the blue-black category (state 6) alone, cultivars range from the moderately dark Merlot to the intensely dark Tannat, with total anthocyanin concentrations varying 3-fold, and the anthocyanin composition (the ratio of the five major anthocyanin types — malvidin, petunidin, delphinidin, peonidin, and cyanidin glycosides) varies independently from total concentration, producing cultivar-specific color hues within the same nominal color class: Cabernet Sauvignon skins are dominated by malvidin-3-glucoside (55-70% of total anthocyanin) producing a blue-shifted dark color, while Nebbiolo skins contain higher proportions of peonidin and cyanidin glucosides (together 40-60%) producing a more red-shifted, garnet-toned color that is one of Nebbiolo's most distinctive winemaking characteristics despite both cultivars scoring as OIV 225:6. The genetic architecture of skin color variation has been resolved to the VvMYBA locus on chromosome 2, where a gradient of allelic states from fully functional (producing blue-black skin) through partially functional (rose, red) to non-functional (green-yellow) explains the six-state color scale as a quantitative series of regulatory gene activity levels.
Beerenhautfarbe-Beschreibung
Exokarp-Pigmentierung
berry skin pigmentation
exocarp color record
external berry coloration
äußere Beerenfärbung
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Kobayashi S et al. (2004) Retrotransposon-induced mutations in grape skin color, Science 304: 982 Mattivi F et al. (2006) Metabolite profiling of grape: flavonols and anthocyanins, Journal of Agricultural and Food Chemistry 54: 7692-7702 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia
Skin Color Description
Skin density comparison between Cabernet Sauvignon (1.094±0.009 g/mL) and Pinot Noir (1.078±0.011 g/mL) at equivalent sugar levels reflects the thicker cell walls and higher tannin concentration in Cabernet skin tissue, correlating with the 40% higher total phenolic extraction observed during identical maceration protocols and explaining the characteristic structural differences between wines from these varieties.
Skin density quality is a tissue density quality that inheres in the grape berry exocarp (skin) and represents the mass per unit volume of this protective tissue layer characterized by intermediate density values between high-density seeds and lower-density pulp, reflecting the unique cellular organization of thick-walled epidermal cells, cuticle deposition, and the concentration of phenolic compounds including anthocyanins and tannins within skin cell vacuoles. Within the BFO framework, this quality characterizes the exocarp as a functionally distinct tissue whose density integrates structural components (cell walls, cuticle) with compositional elements (phenolics, aromatics) that are of primary importance for wine quality determination. The ontological significance lies in skin density serving as a composite indicator of skin development and composition, where cell wall thickness, degree of lignification, cuticle wax accumulation, and vacuolar phenolic concentration all contribute to the measured value. Grape skin density typically ranges from 1.05-1.12 g/mL depending on cultivar, developmental stage, and environmental conditions, with red varieties showing different density dynamics than white varieties due to anthocyanin accumulation post-véraison. Measurement of skin density requires careful separation of exocarp from mesocarp tissue, challenging due to the thin nature of grape skin (typically 6-10 cell layers, 100-200 μm thickness), and may employ micro-pycnometry or density gradient techniques on pooled skin samples. The skin proportion of total berry mass (typically 8-15% depending on berry size and skin thickness) means that skin density contributes modestly to whole-berry density but significantly to texture perception and phenolic extractability. Understanding skin density supports interpretation of berry sorting behavior and prediction of maceration outcomes in winemaking.
Skin_Density_Quality specifically characterizes the exocarp tissue, recognizing its critical importance for wine quality despite comprising a small proportion of berry mass. This quality integrates structural and compositional parameters relevant to texture analysis and phenolic extraction prediction. Measurement protocols must carefully define the tissue boundary between skin and pulp, as inclusion of outer mesocarp cells significantly affects density values.
Beerenhautdichte
Exokarpdichte
Hautgewebedichte
Schalendichte
berry skin specific gravity
exocarp density
grape skin compactness
pericarp outer layer density
Ortega-Regules A et al. (2006) A first approach towards the relationship between grape skin cell-wall composition and anthocyanin extractability. Analytica Chimica Acta 563(1-2): 26-32 Rolle L et al. (2009) Relationship between skin break force and anthocyanin extractability at different degrees of ripening. American Journal of Enology and Viticulture 60(1): 93-97
Skin Density Quality
Skin elasticity assessment comparing crack-susceptible Emperor and crack-resistant Flame Seedless table grapes reveals maximum elastic strain of 8.2% versus 14.5% respectively before plastic deformation onset, with the 77% higher extensibility in Flame Seedless allowing accommodation of 45 mL/day water uptake without splitting while Emperor cracks at uptake rates exceeding 25 mL/day, guiding irrigation management to restrict pre-harvest water application in susceptible varieties.
Skin elasticity quality is a tissue elasticity quality that inheres in grape berry exocarp and represents the capacity of the skin tissue layer to undergo reversible stretching and deformation, determining the skin's ability to accommodate berry volume changes during growth, respond to turgor fluctuations, and resist splitting under internal pressure or external impact. Within the BFO framework, this quality specifically characterizes the elastic behavior of the outer protective tissue layer whose mechanical properties critically influence berry integrity, splitting susceptibility, and texture perception during consumption or processing. The ontological significance lies in skin elasticity serving as a primary determinant of berry mechanical stability, where highly elastic skin stretches to accommodate growth and turgor fluctuation without failure, while reduced elasticity creates stress concentration points predisposing to cracking, particularly when rapid water uptake following rain generates internal pressure exceeding the skin's stretch capacity. Grape skin elasticity reflects the composite behavior of the cuticle (viscoelastic wax and cutin matrix), epidermal cell walls (cellulose-pectin network), and underlying hypodermal layers, with each component contributing differently to overall skin mechanical behavior. The developmental trajectory shows skin elasticity declining during ripening as cell wall modification reduces extensibility, with the timing and extent of elasticity loss varying among cultivars and correlating with splitting susceptibility in thin-skinned varieties. Measurement approaches include uniaxial tensile testing of excised skin strips, biaxial inflation testing mimicking internal pressure conditions, and puncture testing providing composite skin mechanical parameters. The relationship between skin elasticity and cuticle properties links mechanical behavior to water permeability and pathogen resistance, as cuticular microcracking from elastic failure compromises barrier function. Understanding skin elasticity quality enables prediction of splitting risk, interpretation of textural differences among varieties, and optimization of harvest timing to balance ripeness against mechanical stability.
Skin_Elasticity_Quality specifically addresses the exocarp mechanical behavior critical for berry integrity and splitting resistance. This quality integrates contributions from cuticle, epidermis, and hypodermis that may be separately characterized in detailed mechanical studies. The class supports splitting risk assessment, variety characterization for table grape production, and interpretation of rain damage susceptibility in wine grape production.
Beerenhautelastizität
Exokarpelastizität
Hautelastizität
Schalendehnbarkeit
berry skin stretchability
exocarp elasticity
peel elastic property
skin extensibility
Considine J, Brown K (1981) Physical aspects of fruit growth: theoretical analysis of distribution of surface growth forces in fruit in relation to cracking and splitting. Plant Physiology 68(2): 371-376 Lang A, Düring H (1990) Grape berry splitting and some mechanical properties of the skin. Vitis 29(2): 61-70
Skin Elasticity Quality
Skin firmness quality in Pinot Noir presents the viticultural challenge inherent to this thin-skinned variety, with skin break force of 0.3-0.5 N (versus 0.6-1.0 N for Cabernet Sauvignon) creating high susceptibility to splitting from late-season rain and damage during mechanical harvest Burgundy vignerons managing Pinot Noir employ hand harvest and gentle handling to preserve the fragile skin integrity that determines both immediate oxidation risk and the controlled extraction enabling the elegant, transparent wines the variety is capable of producing.
Skin_Firmness_Quality represents the physical quality characterizing the mechanical resistance of the grape berry skin (exocarp) to deformation, penetration, and rupture, capturing the textural property most directly relevant to handling damage resistance, splitting susceptibility, and skin integrity during processing. The grape skin consists of a waxy cuticle overlying the epidermis and several layers of hypodermal cells with thickened cell walls, creating a mechanically distinct outer layer that provides structural containment for the turgid internal tissues. Skin firmness encompasses multiple mechanical properties including tensile strength (resistance to stretching), puncture resistance (force required for probe penetration), and break strain (extent of deformation before failure). These properties determine skin behavior under the stresses of berry expansion during ripening, the pressure of handling and transport, and the mechanical extraction during crushing. Skin firmness decreases during ripening as cell wall modifications reduce mechanical strength, with the extent of softening affecting susceptibility to splitting (from rain, irrigation, or turgor fluctuation) and handling damage. Cultivar differences in skin firmness are substantial, with thick-skinned varieties (Cabernet Sauvignon, Mourvèdre) showing higher firmness than thin-skinned varieties (Pinot Noir, Gewürztraminer). Skin firmness affects wine extraction kinetics, as tougher skins require longer maceration or more aggressive processing for equivalent color and tannin extraction. Skin firmness measurement employs puncture testing through the skin surface, tensile testing of excised skin strips, or specialized skin break force determination that isolates the skin rupture event from pulp penetration. The thin, curved geometry of grape skin creates measurement challenges requiring careful methodology for reproducible results.
Skin_Firmness_Quality specifically addresses the mechanical properties of the berry exterior as distinct from internal pulp firmness. The multiple mechanical properties encompassed (tensile, puncture, break) reflect different aspects of skin mechanical behavior relevant to different quality outcomes. Skin thickness and firmness are related but distinct parameters thin skins can be relatively strong while thick skins can be mechanically weak depending on cell wall composition. The class is particularly relevant to both table grape (damage resistance) and wine grape (splitting, extraction) quality assessment.
Beerenhautfestigkeit
Exokarp-Festigkeit
Schalenfestigkeit
Schalenzähigkeit
exocarp firmness
peel strength
skin mechanical resistance
skin toughness
Letaief H et al. (2008) Assessment of Grape Skin Hardness by a Puncture Test. Journal of the Science of Food and Agriculture 88(9): 1567-1575 Río Segade S et al. (2011) Possible Use of Texture Characteristics as Markers of Winegrape Quality. Journal of Agricultural and Food Chemistry 59(6): 2386-2395
Skin Firmness Quality
Skin maturity assessment in Syrah using extractability indices reveals the cell wall mediation of phenolic potential, with early harvest showing high total anthocyanins (1.8 mg/g berry) but low extractability index (35%, cell walls intact limiting release) yielding extracted anthocyanins of 0.63 mg/g, optimal harvest showing slightly lower total anthocyanins (1.6 mg/g) but high extractability (65%, cell wall modification enabling release) yielding 1.04 mg/g extracted, and late harvest showing declining total (1.3 mg/g) with very high extractability (80%) but only 1.04 mg/g extracted despite easier release, demonstrating how skin maturity optimization balances accumulation and extractability for maximum wine color rather than targeting maximum grape concentration alone.
Skin maturity quality is a physiological maturity quality that represents the developmental state of berry exocarp tissue, characterizing the structural and biochemical changes in the skin that determine phenolic accumulation, anthocyanin stability, aroma compound localization, and the cell wall modifications affecting extractability of these quality-determining compounds during winemaking. Within the BFO framework, this quality specifically addresses the skin component of physiological maturity, distinguishing exocarp development from pulp or seed maturation while recognizing that skin maturity is paramount for red wine quality through its determination of color stability, tannin contribution, and aromatic character, and increasingly appreciated for white wine quality through textural phenolics and skin-contact techniques. The ontological significance lies in skin maturity determining the extractability and quality of phenolic compounds that distinguish grape varieties and define wine character, where the accumulation of anthocyanins (color), tannins (structure), and flavonols (co-pigmentation) in skin cells, combined with cell wall changes affecting their release during maceration, together establish the phenolic potential available for expression during winemaking. The anthocyanin dynamics during skin maturation show accumulation beginning at véraison, reaching maximum concentration at intermediate ripeness, then potentially declining through degradation or dilution during late ripening, with the stability of accumulated anthocyanins (acylation, methylation, co-pigmentation) also evolving and affecting the color extraction and retention potential of the fruit. The cell wall changes during skin maturation include pectin modification (affecting cell separation during maceration), hemicellulose and cellulose changes (affecting cell wall porosity), and cuticle development (affecting berry water relations and disease susceptibility), with these structural changes determining how readily phenolic compounds are released from skin cells during winemaking. The extractability dimension of skin maturity recognizes that total phenolic content and extractable phenolic content may differ substantially, with optimal skin maturity representing not necessarily maximum accumulation but rather the combination of accumulated quantity and cell wall condition that maximizes beneficial extraction during the winemaking timeframe. Understanding skin maturity quality enables assessment of phenolic extraction potential, harvest timing for optimal color and tannin contribution, and recognition of the cell wall changes that mediate between grape phenolic content and wine phenolic composition.
Skin_Maturity_Quality addresses the developmental state of berry exocarp determining phenolic accumulation and extractability. This quality is critical for red wine color and structure through anthocyanin and tannin extraction. The class supports phenolic quality optimization through skin maturity characterization balancing accumulation and extraction potential.
Beerenhautentwicklung
Exokarp-Reife
Schalenreife
exocarp development
exocarp maturity
phenolic maturity
skin ripeness
Río Segade S et al. (2008) Phenolic ripeness assessment of grape skin by texture analysis. Journal of Food Composition and Analysis 21(8): 644-649 Ortega-Regules A et al. (2006) A first approach towards the relationship between grape skin cell-wall composition and anthocyanin extractability. Analytica Chimica Acta 563(1-2): 26-32
Skin Maturity Quality
In Nebbiolo berries in Barolo, Piedmont during BBCH 81-89, skin phenolic development establishes anthocyanin content increasing from 0.15 to 0.85 mg/g fresh weight, skin tannin concentration rising from 1.8 to 4.2 mg/g skin tissue with mDP increasing from 22 to 35, and flavonol content reaching 125 mg/kg dominated by quercetin-3-glucoside over 52 days, with phenolic extractability increasing 65% as assessed by model wine extraction under cool Piedmont conditions averaging 22°C.
Skin phenolic development constitutes the comprehensive maturation of phenolic compounds within Vitis vinifera berry exocarp tissues during the ripening period, wherein anthocyanins, proanthocyanidins, flavonols, and hydroxycinnamic acids undergo coordinated accumulation and structural modification in hypodermal and epidermal cell layers establishing the skin phenolic profile that determines red wine color extraction, contributes substantial tannin content, and provides copigmentation factors essential for wine color stability over the 35-50 day post-veraison maturation period. This process integrates multiple biosynthetic and modification pathways operating in distinct exocarp cell populations, with anthocyanin synthesis and accumulation restricted to hypodermal cells where vacuolar pigment deposition generates berry color, proanthocyanidin accumulation distributed across multiple skin cell layers with highest concentrations in outer hypodermal regions, and flavonol synthesis occurring primarily in epidermal cells where UV-protective functions are most relevant. The quantitative aspects of skin phenolic development show substantial cultivar variation, with thick-skinned varieties such as Cabernet Sauvignon and Tannat accumulating skin tannin concentrations of 3-6 mg/g skin tissue compared to thin-skinned varieties such as Pinot Noir reaching 1-3 mg/g, while anthocyanin concentrations vary independently based on variety-specific pigment biosynthetic capacity ranging from 0.5-4.0 mg/g fresh weight across diverse cultivars. The temporal dynamics of skin phenolic development demonstrate compound-class-specific patterns, with flavonol accumulation largely complete before veraison providing UV protection during early berry development, anthocyanin synthesis initiating at veraison and continuing throughout ripening in red varieties, and skin proanthocyanidin accumulation showing continued increase during post-veraison stages unlike seed tannins whose biosynthesis largely ceases before veraison. Environmental factors profoundly influence skin phenolic development, with light exposure essential for flavonol synthesis and beneficial for anthocyanin accumulation when not accompanied by excessive temperature, moderate water deficit enhancing both anthocyanin and tannin accumulation through berry size reduction and metabolic pathway activation, and temperature extremes potentially limiting anthocyanin synthesis while having lesser effects on tannin accumulation. The extractability of skin phenolics evolves during ripening through cell wall modification and membrane permeability changes, with increasing extractability of both anthocyanins and tannins during maturation facilitating color and tannin release during winemaking, though the precise extraction dynamics depend on vinification conditions including temperature, alcohol development, and maceration duration. Viticultural practices targeting skin phenolic development include canopy management optimizing cluster light exposure, irrigation strategies imposing controlled water deficit, and crop load adjustment ensuring adequate phenolic concentration per berry, with these interventions capable of modifying skin phenolic content by 30-100% relative to standard practices.
Skin_Phenolic_Development is parent class under Phenolic_Maturation representing exocarp-localized phenolic processes. Contains subclasses Skin_Tannin_Maturation, Flavonoid_Accumulation, and Skin_Phenolic_Profile. Distinguishes from Seed_Phenolic_Maturation by tissue localization. Integrates anthocyanin, tannin, and flavonol development. Duration 35-50 days. Light and water stress responsive. Extractability increases during ripening.
Beerenhaut-phenolische-Reifung
Exokarp-Phenol-Entwicklung
Haut-Phenol-Entwicklung
Haut-Polyphenol-Entwicklung
berry skin phenolic maturation
exocarp phenolic development
skin polyphenol development
skin tannin and anthocyanin development
Downey MO et al (2006) Cultural practice and environmental impacts on the flavonoid composition of grapes and wine. American Journal of Enology and Viticulture 57: 257-268 Adams DO (2006) Phenolics and ripening in grape berries. American Journal of Enology and Viticulture 57: 249-256 Keller M (2020) The Science of Grapevines 3rd Ed pp 310-335
Skin Phenolic Development
In Corvina berries in Valpolicella, Veneto at BBCH 89, skin phenolic profile is characterized by anthocyanin content of 0.95 mg/g with malvidin-3-glucoside at 42%, peonidin-3-glucoside at 28%, and cyanidin-3-glucoside at 18%, skin tannin of 2.8 mg/g with mDP 32 and prodelphindin 34%, and flavonol content of 95 mg/kg with quercetin derivatives dominating at 72%, establishing the distinctive profile that contributes to Amarone character when concentrated through appassimento drying under northern Italian conditions.
Skin phenolic profile describes the characteristic composition and relative proportions of phenolic compound classes within Vitis vinifera berry exocarp tissues at harvest maturity, wherein the integrated outcomes of anthocyanin, proanthocyanidin, flavonol, and hydroxycinnamic acid accumulation establish the distinctive phenolic signature that defines cultivar identity, determines extraction potential during winemaking, and influences wine color, structure, and stability properties in wines produced from specific grape varieties and vineyard sources. This concept encompasses the quantitative ratios among major phenolic classes including the anthocyanin-to-tannin ratio typically ranging from 0.3-0.8 in skin tissues depending on variety and ripeness, the flavonol-to-anthocyanin ratio of 0.05-0.20 that influences copigmentation capacity, and the relative proportions of individual compounds within each class that create variety-specific compositional fingerprints enabling cultivar discrimination through phenolic analysis. The cultivar-specific nature of skin phenolic profile reflects genetic differences in biosynthetic pathway regulation, with varieties such as Cabernet Sauvignon characterized by high malvidin-dominated anthocyanin profiles with substantial acylation, varieties such as Pinot Noir showing elevated cyanidin and peonidin contributions with minimal acylation, and varieties such as Nebbiolo demonstrating unique profiles with high peonidin proportions and distinctive tannin structures. The environmental modulation of skin phenolic profile occurs through differential effects on individual biosynthetic pathways, with light exposure preferentially enhancing flavonol accumulation relative to other flavonoids, temperature affecting anthocyanin-to-tannin ratios through differential temperature sensitivity of respective pathways, and water status influencing overall phenolic concentration while maintaining relatively stable compositional ratios within bounds established by genetic potential. The assessment of skin phenolic profile employs multiple analytical approaches including HPLC quantification of individual anthocyanins and flavonols, spectrophotometric determination of total phenolic classes, and advanced methods such as phloroglucinolysis for detailed tannin characterization, with these techniques providing comprehensive phenolic profiling that supports quality assessment and winemaking decisions. The practical significance of skin phenolic profile for winemaking relates to extraction optimization, with knowledge of phenolic composition guiding decisions regarding maceration duration, temperature management, and press fraction utilization to achieve desired wine style while managing potential quality-limiting factors such as excessive tannin extraction or inadequate color development.
Skin_Phenolic_Profile is subclass of Skin_Phenolic_Development representing the compositional outcome at maturity. Distinguishes from developmental processes by emphasizing final profile characterization. Sibling to Skin_Tannin_Maturation and Flavonoid_Accumulation. Integrates anthocyanin, tannin, and flavonol compositions. Cultivar-specific fingerprint. Guides extraction optimization during winemaking.
Beerenhaut-Polyphenol-Profil
Exokarp-Phenol-Zusammensetzung
Haut-Phenol-Profil
Haut-phenolische-Signatur
berry skin phenolic signature
exocarp phenolic composition
skin phenolic fingerprint
skin polyphenol profile
Mattivi F et al (2006) Metabolite profiling of grape. Journal of Agricultural and Food Chemistry 54: 7692-7702 Downey MO et al (2006) Cultural practice and environmental impacts on flavonoid composition. American Journal of Enology and Viticulture 57: 257-268 Keller M (2020) The Science of Grapevines 3rd Ed pp 320-335
Skin Phenolic Profile
Skin tannin concentration ranges from 0.3-2.0 mg/berry across cultivars, comprising 20-50% of total berry tannin depending on genotype and ripeness stage. High skin tannin cultivars such as Cabernet Sauvignon (1.2-2.0 mg/berry) and Tannat (1.5-2.5 mg/berry) contrast with lower skin tannin types like Merlot (0.8-1.2 mg/berry) and Pinot Noir (0.4-0.8 mg/berry). Skin proanthocyanidins show higher mean degree of polymerization (mDP 15-40) than seed tannins (mDP 3-12), contributing less astringent, more rounded mouthfeel. The prodelphindin content (epigallocatechin subunits) reaches 15-35% in skin tannins versus <5% in seeds, with trihydroxylated subunits conferring reduced astringency perception. Skin tannin extractability increases during ripening as cell walls soften, reaching 40-70% extraction during extended maceration.
A tannin accumulation disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate proanthocyanidins specifically in the hypodermal cell layers of grape berry skins, with genotype-specific concentration, polymer length (typically longer than seed tannins), and subunit composition (higher proportion of epigallocatechin) contributing softer, less astringent tannin character.
Skin tannin contributes the preferred "soft" tannin character in wine, with longer polymers and higher prodelphindin content creating less aggressive astringency than seed tannins. The trait shows independent genetic control from seed tannin, enabling selection for modified skin/seed tannin ratios. Skin tannins also provide sites for anthocyanin incorporation into polymeric pigments during winemaking and aging, contributing to color stability. Water deficit during ripening increases skin tannin concentration through both enhanced synthesis and berry size reduction effects.
Beerenhaut-Proanthocyanidin
Exokarp-Gerbstoff
Schalentannin
exocarp tannin trait
skin condensed tannin
skin proanthocyanidin
Downey MO et al. (2003) Analysis of tannins in seeds and skins of Shiraz grapes throughout berry development. Australian Journal of Grape and Wine Research 9:15-27 Bindon KA et al. (2014) Relationships between harvest time and wine composition in Vitis vinifera L. cv. Cabernet Sauvignon. 1. Grape and wine chemistry. Food Chemistry 146:253-263
Skin Tannin Disposition
In Sagrantino berries in Montefalco, Umbria during BBCH 83-89, skin tannin maturation increases skin tannin content from 2.4 to 5.8 mg/g skin tissue over 45 days, with mDP rising from 28 to 42, prodelphindin content stable at 35-38%, and galloylation at 4.5-5.2%, while extractability measured by model wine extraction increases from 45% to 78% of total skin tannin content, establishing the exceptionally high skin tannin concentration characteristic of this variety under warm central Italian conditions averaging 27°C.
Skin tannin maturation describes the progressive structural and quantitative evolution of proanthocyanidin compounds within Vitis vinifera berry exocarp tissues during the ripening period, wherein skin tannins undergo continued accumulation, polymer chain elongation, and compositional modification to establish the mature tannin population characterized by high mean degree of polymerization of 25-45 units, elevated prodelphindin content of 20-40%, and moderate galloylation of 2-8% that distinguishes skin tannins from their seed counterparts and contributes distinctive mouthfeel properties to wines produced with extended skin contact. This process involves continued proanthocyanidin biosynthesis during post-veraison ripening unlike seed tannins whose synthesis largely ceases before veraison, with ongoing expression of leucoanthocyanidin reductase and anthocyanidin reductase in skin tissues enabling accumulation increases of 30-80% between veraison and harvest in many varieties, establishing final skin tannin concentrations of 1-6 mg/g skin tissue depending on cultivar and growing conditions. The structural characteristics of skin tannin maturation reflect the distinctive biosynthetic environment of exocarp cells, with high expression of flavonoid 3-prime-5-prime-hydroxylase generating substantial epigallocatechin incorporation yielding the elevated prodelphindin percentages of 25-40% that exceed seed levels by 5-10 fold and contribute to enhanced protein precipitation capacity and specific sensory properties of skin-derived tannins. The temporal dynamics of skin tannin maturation show progressive accumulation throughout ripening with potential acceleration during late stages when berry dehydration may concentrate tannin content, while structural modifications including continued polymerization increasing mDP and potential oxidative changes modifying extractability occur concurrently with quantitative accumulation. Environmental factors influence skin tannin maturation through effects on both biosynthetic pathway activity and berry size determining concentration effects, with moderate water deficit enhancing skin tannin accumulation through reduced berry size and potential metabolic activation, while excessive stress may limit biosynthesis compromising final tannin content. The extractability of skin tannins evolves during maturation through cell wall softening and pectin solubilization that increases accessibility of vacuolar tannin pools to extracting solvents, with extraction efficiency typically increasing 40-80% between veraison and harvest as cellular barriers to tannin release progressively diminish. The enological significance of skin tannin maturation relates to wine structural properties, with skin tannins contributing substantial polymeric material that interacts with anthocyanins through copigmentation and covalent bonding, participates in color stabilization reactions, and provides mouthfeel characteristics described as softer and rounder compared to the more astringent character of seed-derived tannins.
Skin_Tannin_Maturation is subclass of Skin_Phenolic_Development representing exocarp proanthocyanidin evolution specifically. Distinguishes from Seed_Tannin_Polymerization by tissue location and structural characteristics. Sibling to Flavonoid_Accumulation and Skin_Phenolic_Profile. Skin mDP 25-45 versus seed 5-12. Prodelphindin 20-40% versus seed less than 5%. Continues accumulating post-veraison unlike seed tannins.
Beerenhaut-kondensierte-Tannin-Evolution
Exokarp-Proanthocyanidin-Reifung
Haut-Tannin-Entwicklung
Haut-Tannin-Reifung
berry skin tannin ripening
exocarp tannin development
skin condensed tannin evolution
skin proanthocyanidin maturation
Downey MO et al (2003) Analysis of tannins in seeds and skins of Shiraz grapes. Australian Journal of Grape and Wine Research 9: 15-27 Souquet JM et al (1996) Polymeric proanthocyanidins from grape skins. Phytochemistry 43: 509-512 Kennedy JA et al (2006) Grape and wine phenolics. In Chemistry and Biology of Ellagitannins World Scientific pp 220-255
Skin Tannin Maturation
The skin texture of Crimson Seedless documents firm, crisp skin that ruptures cleanly with an audible snap when bitten — the skin breaks along a well-defined fracture plane rather than stretching or tearing, producing the crunchy eating quality that is the benchmark texture for premium table grapes in global markets. This crisp rupture behavior reflects the relatively rigid, non-elastic cuticle and the brittle outer hypodermal cells with thick, pectin-poor cell walls that fracture rather than deform under mechanical stress. The crisp Crimson Seedless skin texture contrasts with the elastic, stretchy skin of Concord (V. labrusca, the prototypical slip-skin grape where the skin separates cleanly from the flesh as an intact sac), the tough chewy skin of Cabernet Sauvignon (resistant to rupture, persisting as a fibrous residue during chewing), and the delicate, thin, easily ruptured skin of Chasselas (which splits readily under finger pressure, contributing to the cultivar's poor transportability but excellent eating quality).
A berry skin description component that records the surface tactile properties and mechanical behavior of the berry skin when handled and bitten, documenting the firmness, elasticity, crispness, and resistance to rupture that characterize the eating quality of the berry and reflect the ultrastructural composition of the cuticle, epidermis, and outer hypodermis, with skin texture being a primary selection criterion in table grape breeding programs targeting the crisp, crunchy berry texture preferred by consumers.
Skin_Texture captures the mechanical behavior of the berry skin that, together with flesh texture, determines the overall eating quality (mouthfeel) of the grape berry — a property of commercial importance primarily for table grapes but also relevant for winemaking through its influence on phenolic extractability during maceration. The texture phenotype is determined by the structural composition of the skin at multiple scales — the cuticle thickness and composition (wax versus cutin ratio affecting rigidity versus flexibility), the cell wall composition of epidermal and hypodermal cells (pectin, cellulose, hemicellulose, and lignin proportions affecting brittleness versus elasticity), and the cell-to-cell adhesion strength (middle lamella composition affecting whether stress causes cell rupture or cell separation): cultivars with high cell wall rigidity and weak middle lamella adhesion produce the crisp, snapping texture desired in table grapes, while cultivars with elastic cell walls and strong intercellular adhesion produce the tough, chewy texture typical of thick-skinned wine grapes. The slip-skin character of American V. labrusca cultivars (Concord, Catawba) represents a distinct texture phenotype in which the skin separates from the flesh as an intact sac due to a weak boundary between the hypodermis and the outer mesocarp — a characteristic absent in V. vinifera cultivars where skin and flesh form a continuous tissue gradient.
Beerenhaut-Textur
Beerenschalen-Beschaffenheit
berry skin feel
exocarp mechanical behavior
skin firmness characterization
Skin Texture
The skin thickness of Nebbiolo documents thick skin (OIV 228:7) — perceived as firmly resistant to biting through the skin when tasting berries at harvest, with a chewing texture that remains as a distinct residue after the flesh has dissolved, reflecting the 300-400 μm cross-sectional thickness and the 8-10 layers of thick-walled hypodermal cells. The thick skin of Nebbiolo contributes to the cultivar's high tannin concentration (among the highest in V. vinifera, producing the famously astringent young Barolo and Barbaresco wines that require 10-20 years of aging), while paradoxically the anthocyanin concentration is moderate (producing the garnet-rather-than-opaque-black color characteristic of Nebbiolo wines) — a dissociation between tannin and color that reflects Nebbiolo's specific anthocyanin composition (dominated by less stable cyanidin and peonidin forms rather than the more stable malvidin forms of Cabernet Sauvignon).
A berry skin description component that records the thickness of the berry skin as perceived by tactile assessment and chewing resistance, scored according to OIV descriptor 228 on a scale from very thin (state 1) through thin, medium, thick, to very thick (state 9), with skin thickness reflecting the number of hypodermal cell layers, the degree of cell wall thickening and lignification, and the total tissue mass between the outer cuticle and the inner flesh boundary that determines both the wine phenolic extraction potential and the table grape eating quality.
Skin_Thickness documents a character that is assessed by standardized tactile and gustatory evaluation rather than instrumental measurement in routine ampelographic practice — the OIV scoring protocol specifies that skin thickness should be assessed by placing a berry in the mouth, pressing it against the palate to rupture the skin, chewing the skin while swallowing the flesh and juice, and scoring the perceived thickness and toughness of the skin residue on the 1-9 scale. This sensory assessment integrates multiple physical properties — actual tissue thickness, cell wall rigidity, cuticle toughness, and the cohesion of the hypodermal tissue — into a composite perceived thickness that correlates well with but is not identical to the microscopically measured cross-sectional thickness: the correlation is imperfect because two skins of identical measured thickness can differ substantially in perceived thickness if one has more lignified cell walls or a tougher cuticle than the other. For table grape assessment, skin thickness is a primary quality criterion — consumers prefer berries with thin, barely perceptible skins that rupture cleanly when bitten (the crisp, light-skinned ideal of Chasselas, Sultana, and modern seedless cultivars), while thick, tough skins that persist as a chewy residue (characteristic of many wine grape cultivars) are considered defects in table grape evaluation.
Beerenhautdicke
Beerenschalen-Dicke
Exokarp-Stärke
berry skin gauge
exocarp thickness record
skin depth assessment
Skin Thickness
Skin-to-pulp ratio for Cabernet Sauvignon from a dry-farmed Oakville hillside block (2023 vintage) calculates to 0.31 (skin 0.29 g / pulp 0.94 g), substantially exceeding the valley-floor irrigated reference of 0.22 (skin 0.26 g / pulp 1.18 g), with the 41% higher ratio directly correlating with the 35% increase in wine total phenolics (72 vs. 53 AU280) and 28% higher anthocyanin extraction (A520 9.8 vs. 7.6 AU) in paired micro-vinifications under identical protocols. Skin-to-pulp ratio for Pinot Noir from Sonoma Coast (2024 vintage) records 0.20, at the lower end of typical red wine cultivar range, reflecting the thin-skinned morphology that fundamentally limits Pinot Noir's phenolic extraction potential relative to thick-skinned varieties regardless of viticultural management. Skin-to-pulp ratio for Furmint from Tokaj (2024 vintage) documents 0.18 for the dry wine lot versus 0.45 for botrytized Aszú berries — the dramatic increase in the noble rot material reflecting water loss from Botrytis-mediated berry desiccation that concentrates the skin fraction while reducing pulp volume.
Skin-to-pulp ratio is a berry composition data entity that records the calculated proportion of skin mass to pulp mass within a grape berry, expressed as a dimensionless ratio (skin weight divided by pulp weight), percentage, or occasionally as skin-to-total-berry-weight percentage, serving as the single most informative derived metric for predicting wine phenolic concentration, color intensity, and aromatic compound density, and representing the integrative physical parameter through which vine water status, crop load, berry size, and cultivar genetics collectively influence wine quality potential.
Skin-to-pulp ratio is arguably the most powerful single predictor of red wine phenolic quality because it integrates the two primary physical determinants of must concentration — the quantity of phenolic-rich skin tissue available for extraction and the volume of phenolic-dilute pulp juice into which extraction occurs. The relationship between skin-to-pulp ratio and wine quality has been demonstrated across multiple cultivars, regions, and experimental frameworks, with the seminal work of Roby et al. (2004) establishing that berry size effects on wine composition are mediated almost entirely through changes in skin-to-pulp ratio rather than through concentration changes within the skin tissue itself. This mechanistic insight has profound practical implications — management practices that reduce berry size (deficit irrigation, crop thinning, rootstock selection) improve wine concentration primarily by increasing the skin-to-pulp ratio, not by altering the biosynthetic capacity of the skin per se. The ratio is cultivar-dependent at a genetic level — thick-skinned varieties (Cabernet Sauvignon, Tannat, Petit Verdot) achieve ratios of 0.25-0.35 under standard conditions, while thin-skinned varieties (Pinot Noir, Gamay, Grenache) typically range from 0.15-0.22, establishing intrinsic cultivar quality ceilings for phenolic concentration. Environmental modulation operates primarily through the pulp component — water deficit reduces pulp volume while skin mass remains relatively stable, increasing the ratio conversely, generous irrigation or rainfall promotes pulp expansion and dilutes the ratio. The ratio also changes dynamically during late-season berry dehydration (common in warm, dry climates and in botrytized fruit), as water loss from the pulp progressively concentrates the skin fraction. For white wine production, the skin-to-pulp ratio is less directly quality-determinant but still relevant for predicting aromatic precursor concentration, phenolic pickup during skin contact, and pressing behavior.
Exokarp-Mesokarp-Verhältnis
Haut-Pulpa-Verhältnis
Hautanteil
Schale-Fruchtfleisch-Verhältnis
Schalenanteil
exocarp-mesocarp ratio
phenolic extraction ratio
skin proportion
skin to juice ratio
skin-pulp ratio
Roby G et al. (2004) Berry size and vine water deficits as factors in winegrape composition: anthocyanins and tannins. Australian Journal of Grape and Wine Research 10(2): 100-107 Walker RR et al. (2005) Rootstock effects on salt tolerance of irrigated field-grown grapevines. Australian Journal of Grape and Wine Research 11(2): 174-183 Matthews MA, Anderson MM (1988) Fruit ripening in Vitis vinifera L.: responses to seasonal water deficits. American Journal of Enology and Viticulture 39(4): 313-320
Skin To Pulp Ratio
Skin weight data for Cabernet Sauvignon from Coonawarra (2023 vintage) records mean skin weight of 0.31 g per berry (23.4% of 1.32 g total berry weight), with the thick-skinned character typical of the cultivar providing a substantial reservoir of extractable anthocyanins (estimated 580 µg per berry) and skin tannins (estimated 420 µg catechin equivalents per berry) that support extended maceration protocols of 21-28 days. Skin weight data for Pinot Noir from Pommard, Burgundy (2024 vintage) documents mean skin weight of 0.18 g (15.8% of 1.14 g berry weight), confirming the thin-skinned morphology that limits phenolic extraction to 10-14 day maceration periods and necessitates whole-cluster inclusion or gentle extraction techniques to build adequate wine structure. Skin weight data for Grüner Veltliner from Loiben, Wachau (2024 vintage) records 0.22 g (14.5% of 1.52 g berry weight), relevant for the extended skin contact (12-24 hours) increasingly practiced for textured, complex dry Grüner Veltliner production.
Skin weight data is a berry composition data entity that records the mass of the grape berry exocarp (skin) following manual dissection and separation from the underlying mesocarp (pulp) and seeds, determined gravimetrically on fresh or frozen berries, expressed in grams per berry (g/berry) or milligrams per berry (mg/berry), and additionally calculated as a percentage of total berry weight, representing the tissue fraction that contains the majority of anthocyanins, flavonols, hydroxycinnamic acids, terpenes, and other flavor-active compounds whose extraction during winemaking determines wine color, aromatic complexity, and textural structure.
Skin weight data provides the physical basis for understanding cultivar-specific differences in phenolic and aromatic extraction potential, as the exocarp functions as the primary biosynthetic and storage site for quality-determining secondary metabolites. The grape berry skin comprises the outer epidermis (cuticle, epidermal cells) and the hypodermis (typically 6-12 cell layers), with anthocyanins localized in the vacuoles of hypodermal cells, flavonols concentrated in the outer epidermal layers, and terpenes distributed across both tissues. Skin weight correlates with but is not identical to skin thickness — cultivars with dense, compact skin cells (Cabernet Sauvignon, Tannat) may show similar weight to cultivars with thicker but more loosely structured skins (Nebbiolo), yet differ substantially in phenolic extractability due to cell wall composition differences. The methodology of skin weight determination requires careful dissection — typically performed on frozen berries to facilitate clean skin-pulp separation — and the completeness of pulp removal from the inner skin surface introduces operator-dependent variability (typically ±5-8% between operators). Skin weight as a percentage of berry weight typically ranges from 12-15% in thin-skinned white varieties (Riesling, Sauvignon Blanc) to 20-25% in thick-skinned red varieties (Cabernet Sauvignon, Tannat, Petit Verdot), though the absolute skin weight per berry provides more direct information about total extractable compound availability than the percentage value. Water deficit increases skin-to-berry weight percentage primarily through pulp volume reduction rather than skin mass increase, though moderate deficit can enhance skin cell wall biosynthesis of secondary metabolites. Skin weight measured on a per-berry basis, multiplied by the number of berries per cluster and clusters per vine, enables estimation of total skin mass per vine — a metric useful for predicting maceration requirements and extraction kinetics.
Beerenhautgewicht
Beerenschalengewicht
Exokarpgewicht
Hautfraktion
Schalengewicht
berry skin weight
exocarp weight
peel weight
skin fraction weight
skin mass
Roby G et al. (2004) Berry size and vine water deficits as factors in winegrape composition. Australian Journal of Grape and Wine Research 10(2): 100-107 Ortega-Regules A et al. (2006) A first approach towards the relationship between grape skin cell-wall composition and anthocyanin extractability. Analytica Chimica Acta 563(1-2): 26-32 Iland P et al. (2011) Chemical Analysis of Grapes and Wine, 2nd Ed, pp.: 80-85
Skin Weight Data
Cabernet Sauvignon slightly overripe berries at week 19 (1 week post-optimal) show 26% sugars (versus optimal 24.5%), 5.8 g/L acidity (versus optimal 6.5 g/L), pH 3.88 (versus optimal 3.75), 3.5 mg/g anthocyanins (adequate), slightly cooked blackberry flavors emerging alongside ripe cassis, producing wine with 15.2% alcohol, slightly reduced structure, good color, acceptable quality for commercial Cabernet though not achieving premium balance of optimal-harvest fruit. Chardonnay slightly overripe berries showing white wine slight overripeness= 23.5% sugars (versus optimal 21.5%), 6.2 g/L acidity (versus optimal 7.5 g/L), pH 3.52 (versus optimal 3.35), tropical fruit character more prominent than desired apple-citrus notes, producing wine with 13.8% alcohol, fuller body than ideal, adequate freshness with acid addition (adjusted to 6.8 g/L), commercially acceptable though lacking crispness of optimal-harvest Chardonnay. Merlot slightly overripe berry winemaking adjustments= tartaric acid addition of 1.2 g/L pre-fermentation restoring titratable acidity from 5.2 g/L to 6.4 g/L and reducing pH from 3.95 to 3.82, cool fermentation at 24-26°C preserving fruit aromatics, 30% new oak providing structure, producing wine showing moderate rather than severe quality compromise from slight overripeness, demonstrating that winemaking interventions can partially compensate for non-optimal berry ripeness. Syrah comparison of optimal harvest (week 18= 25% sugars, 6.2 g/L acidity, 3.8 mg/g anthocyanins, score 92/100 in sensory evaluation) versus slightly overripe harvest (week 19.5= 27% sugars, 5.3 g/L acidity, 3.9 mg/g anthocyanins, score 87/100), showing 5-point quality reduction from slight overripeness reflecting reduced balance and evolution toward cooked fruit character, but maintaining commercially viable quality. Pinot Noir showing cultivar sensitivity to slight overripeness= 1 week delay from optimal (24% to 26% sugars, 6.8 to 5.5 g/L acidity) creating more pronounced quality impact (90 to 84 point scores) than in robust cultivars like Cabernet Sauvignon, demonstrating that Pinot Noir narrow optimal window makes it particularly sensitive to slight overripeness requiring precise harvest timing.
A slightly overripe berry is an overripe berry showing moderate progression beyond optimal ripeness with sugars and acid levels outside ideal ranges but not severely imbalanced, potentially producing acceptable quality with some compromise, representing berries 1-2 weeks past optimal harvest timing. Slightly overripe berries are characterized by= (1) sugars moderately elevated typically 26-28% (versus optimal 23-25% for most red cultivars, 20-23% for whites), representing 1-3% excess sugar creating 0.5-2% excess alcohol potential (2) titratable acidity moderately reduced typically 4.5-5.5 g/L (versus optimal 5.5-7 g/L), representing acid decline sufficient to notice but not creating severe structural problems (3) pH moderately elevated typically 3.8-4 (versus optimal 3.6-3.85 for reds, 3.2-3.5 for whites), creating some microbiological risk but manageable with appropriate winemaking (4) flavors showing beginning of evolution from fresh fruit toward cooked/jammy character but varietal character still recognizable (5) berry texture very soft but not yet shriveling significantly (6) anthocyanins (red cultivars) maintaining adequate levels though potentially plateaued or beginning very slow decline (7) overall quality reduced from optimal but wine production still viable with appropriate adjustments (acid additions, alcohol management, careful handling). Slightly overripe berries develop through 1-2 week harvest delay past optimal timing= sugars accumulating additional 1-3% at declining rates of 0.4-0.8% per week acids declining additional 0.8-1.5 g/L through continued malic acid respiration pH increasing 0.1-0.25 units reflecting acid decline flavors evolving from peak fresh character toward riper cooked notes. Slightly overripe berries may result from= harvest logistics preventing timely harvest at optimal ripeness weather conditions (rain forecast) forcing early harvest before berries reach overripeness or late harvest after berries pass optimal vineyard heterogeneity with some blocks reaching slight overripeness while awaiting slower-ripening blocks intentional extended hang time seeking additional ripeness exceeding optimal by modest amount. Slightly overripe berries produce wines with moderate quality compromise= red wines showing 14.5-15.5% alcohol (elevated but not extreme), slightly flabby structure from reduced acid, beginning of jammy fruit character but fresh notes still present, adequate color and phenolic content, overall quality acceptable for commercial production though not achieving premium quality potential white wines showing reduced freshness and aromatic intensity, fuller body approaching heavy mouthfeel, adequate structure with winemaking adjustments. Slightly overripe berries may benefit from winemaking interventions= acid additions to restore acid balance and reduce pH fermentation temperature management to preserve aromatics moderate oak usage to provide structure early bottling to preserve fruit these interventions can partially mitigate quality effects of slight overripeness producing commercially acceptable wine. Slightly overripe berries represent a common commercial reality= many vineyards harvest at slight overripeness due to operational constraints, achieving satisfactory but not optimal quality understanding slight overripeness allows realistic quality expectations and appropriate winemaking responses avoiding harvest delays that progress from slight to severe overripeness is important quality control goal.
Slightly_Overripe_Berry represents moderate overripeness and is distinguished from Severely_Overripe_Berry by moderate versus extreme compositional imbalances. Slightly overripe berries are common in commercial production and produce acceptable quality with some compromise and appropriate winemaking. Understanding slightly overripe berry characteristics is important for realistic quality assessment, for winemaking adjustments compensating for non-optimal ripeness, and for harvest planning avoiding progression from slight to severe overripeness.
acceptably over-mature berry
leicht überreife Beere
moderat zu reife Traube
moderately overripe berry
mäßig Nachreife-Traube
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 246-255 Jackson, D.I., Lombard, P.B. (1993). Environmental and management practices affecting grape composition. American Journal of Enology and Viticulture 44=409-430.
Slightly Overripe Berry
A Riesling vine in contour-aligned terraces of the Mosel Valley bears a Slope_Aligned_Row_Role following the river-carved topography, with rows perpendicular to the 50-60% slopes regardless of resulting compass orientation—the terrain dictating alignment that no other orientation could accommodate. A Nebbiolo vine on Barolo's steep slopes serves in a Slope_Aligned_Row_Role with traditional up-and-down planting ("rittochino") enabling surface water drainage during autumn rains, the fall-line alignment traditional despite erosion risks managed through cover cropping. A Syrah vine on Côte-Rôtie's precipitous granite slopes bears a Slope_Aligned_Row_Role with ancient terracing determining row position on the narrow benches carved into the hillside over centuries.
A row orientation role that inheres in a grapevine planted in rows aligned to conform with slope topography, either following contours (rows perpendicular to fall line, minimizing erosion) or running up-and-down the slope (rows parallel to fall line, facilitating drainage and mechanization), with the compass orientation resulting from slope aspect rather than deliberate orientation choice. Slope-aligned planting recognizes that topographic constraints often override theoretical optimal orientation steep slopes require contour planting for erosion control regardless of resulting compass direction, while operational considerations on moderate slopes may favor fall-line alignment for tractor safety and vineyard access. Contour-aligned rows (perpendicular to slope) create terracing effect that slows water runoff, reduces soil erosion, and captures rainfall, though they create uneven row lengths and complicate mechanical operations on curved lines. Fall-line aligned rows (up-and-down slope) facilitate water drainage, create uniform row lengths, and enable straight-line tractor operations, but concentrate runoff and may cause erosion on steep slopes. The aspect of the slope (direction it faces) determines the sun exposure of slope-aligned rows, with south-facing slopes in the Northern Hemisphere receiving maximum sun regardless of how rows are oriented within them.
Slope_Aligned_Row_Role captures situations where topography constrains or determines row orientation, acknowledging that many vineyard sites lack the flat terrain enabling free orientation choice. The distinction between contour-aligned (erosion-minimizing but operationally complex) and fall-line aligned (operationally simpler but erosion-prone) reflects fundamental choices in slope management that affect long-term site sustainability. Traditional steep-slope viticulture developed before mechanization, with hand-labor operations creating the extreme terracing (Mosel, Douro, Wachau, Priorat) that modern establishment would rarely attempt. The heritage and quality associations of slope-planted vineyards—difficult sites producing distinctive wines—persist despite the economic challenges that have driven abandonment of many steep-slope areas. Contemporary slope establishment balances erosion control (favoring contour alignment), operational efficiency (favoring straight rows), and safety (limiting mechanization on extreme gradients) with compromises specific to each site's topography.
Gefällezeilen
Hang-ausgerichtete Zeilen
Konturzeilen
contour rows
fall-line rows
slope-aligned rows
topographic alignment
topographische Ausrichtung
Tropeano D (1984) Rate of soil erosion in vineyards, Earth Surface Processes and Landforms 9: 253-266 Rodrigo-Comino J et al. (2018) Soil erosion in sloping vineyards, Earth-Science Reviews 178: 174-193 Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Jackson RS (2014) Wine Science: Principles and Applications, Academic Press
Slope Aligned Row Role
The slope angle of vineyards in the Côte-Rôtie appellation (Northern Rhône, France) documents extreme gradients of 30-60% (17-31°) on the granitic and micaschist hillsides above Ampuis — these steep slopes necessitate entirely manual viticultural operations (no tractor access is possible above approximately 35% gradient), with workers using fixed winch systems on cables to transport equipment and harvest bins up and down the slope, and all pruning, canopy management, and harvest labor performed by hand on the steep terrain. The labor cost of viticulture on these slopes is 3-5 times higher per hectare than on mechanizable flat vineyards (estimated at 800-1,200 hours/ha/year versus 200-350 hours/ha/year), a cost that is economically viable only because the premium wine quality achieved on these thermally advantaged slopes commands prices sufficient to cover the extreme labor investment. The steep Côte-Rôtie gradients contrast with the gentle slopes of Bordeaux's Médoc (2-5%, 1-3°, fully mechanizable), the moderate slopes of Burgundy's Côte d'Or (8-20%, 5-11°, partially mechanizable with specialized narrow tractors), and the extreme terraced slopes of the Douro Valley (40-80%, 22-39°, requiring stone terrace construction to create workable planting platforms).
A slope description component that records the steepness of the vineyard slope expressed as gradient (percent rise over run), angle (degrees from horizontal), or both, with slope angle determining the magnitude of radiation enhancement on sun-facing slopes, the velocity of surface water runoff and associated erosion risk, the feasibility of mechanized vineyard operations, and the labor intensity of all viticultural practices from planting through harvest.
Slope_Angle documents the parameter that most directly affects the practical feasibility and economic viability of hillside viticulture — the gradient determines whether mechanized equipment can operate safely on the slope, and this mechanization threshold (approximately 30-35% gradient for specialized hillside tractors, 15-20% for standard vineyard tractors) divides the viticultural landscape into mechanizable and non-mechanizable zones with fundamentally different cost structures and labor requirements. The radiation enhancement effect of slope angle follows a cosine relationship with the solar elevation angle — the radiation intensity on a sloped surface is maximized when the slope angle equals the complement of the solar elevation angle (i.e., when the sun's rays are perpendicular to the slope surface), producing the theoretical optimum slope for radiation maximization that varies with latitude and season: at 50°N latitude (Mosel, Rheingau), the optimal south-facing slope for midsummer radiation maximization is approximately 27° (50% gradient), while at 45°N (Burgundy, Northern Rhône) the optimum is approximately 22° (40% gradient), and at 35°N (southern Mediterranean) the optimum decreases to approximately 12° (21% gradient) and the radiation limitation shifts from total amount to temporal distribution (morning versus afternoon). The erosion risk associated with slope angle follows a power-law relationship — the Universal Soil Loss Equation (USLE) includes a slope-length factor that increases approximately as the 1.3 power of gradient, meaning that a doubling of slope angle more than doubles the erosion rate, making soil conservation measures (cover crops, terracing, contour planting) essential on steep vineyard slopes.
Hangneigung
Neigungswinkel
Weinberg-Steilheit
inclination angle measurement
slope gradient record
vineyard steepness
Gladstones J (1992) Viticulture and Environment, Winetitles van Leeuwen C and Seguin G (2006) The concept of terroir in viticulture, Journal of Wine Research 17: 1-10 Smart RE and Robinson M (1991) Sunlight into Wine, Winetitles Wischmeier WH and Smith DD (1978) Predicting Rainfall Erosion Losses — A Guide to Conservation Planning, USDA Agriculture Handbook 537
Slope Angle
The slope aspect of the Côte d'Or in Burgundy documents a remarkably consistent east-to-southeast orientation (70-140° azimuth) across the 50 km length of the escarpment from Dijon to Santenay — this east-facing aspect captures the full morning sun from sunrise through solar noon, providing maximum radiation during the cooler morning hours when photosynthetic efficiency is highest and evaporative demand is lowest, while the afternoon sun falls obliquely on the east-facing slope or is blocked by the hilltop to the west, reducing the heat stress and excessive evapotranspiration that would occur on a south- or west-facing slope at this latitude. The Burgundian east-facing aspect is considered ideal for Pinot Noir quality at 47°N because it provides sufficient total radiation for ripening without the excessive afternoon heat that would accelerate sugar accumulation beyond the rate of flavor development — the concept of ripening at moderate rather than high temperatures being central to the Burgundian quality philosophy of achieving physiological maturity at moderate sugar levels (12.5-13.5% potential alcohol) rather than the high sugar levels (14-15% potential alcohol) that south-facing slopes in warmer climates produce.
A slope description component that records the compass orientation (azimuth) of the vineyard slope surface — the direction that the slope faces, expressed as a cardinal or intercardinal compass bearing (N, NE, E, SE, S, SW, W, NW) or as degrees from north (0-360°) — with aspect being the primary determinant of the diurnal radiation pattern (morning versus afternoon sun exposure), the thermal regime of the slope surface, and the wind exposure from prevailing weather directions that together create the aspect-specific mesoclimate distinguishing adjacent slopes facing different directions.
Slope_Aspect is arguably the single most influential topographic parameter for vineyard mesoclimate differentiation — adjacent slopes facing different directions but otherwise identical in elevation, angle, soil, and regional climate can differ by 1-3°C in mean growing season temperature and by 20-40% in total growing season radiation, producing differences in vine phenology, fruit composition, and wine style comparable to those between regions separated by hundreds of kilometers. The effect of aspect operates through the angular relationship between the slope surface normal and the sun's position — south-facing slopes (in the northern hemisphere) maximize total daily radiation by facing the sun's arc across the southern sky, producing the warmest mesoclimates, while north-facing slopes receive predominantly diffuse radiation with minimal direct beam radiation during winter months, producing the coolest mesoclimates: east-facing and west-facing slopes receive similar total daily radiation but differ in the temporal distribution — east-facing slopes receive more morning radiation (when air temperatures are cooler and photosynthetic capacity is higher) while west-facing slopes receive more afternoon radiation (when air temperatures are warmer and the risk of heat stress is greater) — a distinction that affects vine water use, photosynthetic efficiency, and the thermal amplitude (difference between day maximum and night minimum temperatures) that influences anthocyanin synthesis and acid retention during ripening. The traditional European viticultural terminology reflects the importance of aspect — German Weinlage classification distinguishes Südhang (south slope), Südwesthang (southwest slope), and Osthang (east slope) as the premium aspects, while French terroir discourse emphasizes the exposition as a defining element of vineyard identity.
Exposition
Hangausrichtung
Weinberg-Himmelsrichtung
hillside compass bearing
slope orientation record
vineyard aspect measurement
Gladstones J (1992) Viticulture and Environment, Winetitles van Leeuwen C and Seguin G (2006) The concept of terroir in viticulture, Journal of Wine Research 17: 1-10 Jones GV et al. (2004) Spatial analysis of climate in winegrape growing regions, American Journal of Enology and Viticulture 55: 304-316 Jackson DI and Schuster DF (2001) The Production of Grapes and Wine in Cool Climates, Gypsum Press
Slope Aspect
The slope description for the Bernkasteler Doctor vineyard (Mosel, Germany, 49.92°N) documents a steep south-southwest facing slope at 120-200 m elevation with a gradient of 50-65% (27-33°) on Devonian slate bedrock — this extreme slope creates one of the most thermally privileged vineyard sites in northern European viticulture, where the perpendicular incidence angle of solar radiation on the south-facing slope increases the effective radiation intensity by 30-40% compared to a horizontal surface at the same latitude, the dark Devonian slate absorbs and re-radiates thermal energy (increasing nighttime minimum temperatures by 1-3°C compared to adjacent flat sites), and the steep gradient ensures rapid cold air drainage that protects the mid-slope vines from the radiation frost events that damage vineyards on the Mosel valley floor. The Bernkasteler Doctor slope parameters explain why Riesling achieves full ripeness at this extreme northerly latitude (50°N) despite a marginal growing season temperature of only 14-15°C — the slope-enhanced radiation effectively shifts the thermal environment to the equivalent of a site 2-3° latitude further south.
A site description component that documents the topographic inclination characteristics of the vineyard site — the angle, aspect (compass orientation), and landscape position of the slope on which the vines are planted — providing the integrated topographic characterization that determines solar radiation interception, air drainage and frost risk, surface water runoff and erosion potential, and the soil depth gradient from hilltop to valley floor that collectively constitute the most influential topographic modifiers of vineyard mesoclimate and vine performance.
Slope_Description integrates the three component slope parameters — angle, aspect, and position — into a holistic topographic characterization that captures the functional consequences of hillside viticulture. The importance of slope in viticultural terroir is reflected in the historical concentration of premium vineyards on hillside sites across all traditional European wine regions — the Côte d'Or in Burgundy, the Rheingau and Mosel in Germany, the Douro Valley in Portugal, the Northern Rhône, the hillside crus of Barolo and Barbaresco — where slope-enhanced radiation, improved drainage, and cold air movement create mesoclimatic advantages over adjacent flat sites: these advantages are most pronounced at high latitudes (above 45°N) where the low solar elevation angle means that sloped surfaces intercept substantially more radiation than horizontal surfaces, and in continental climates where cold air pooling on flat sites creates frost hazards that sloped sites avoid through gravitational air drainage. The quantification of slope effects on vineyard mesoclimate has been enabled by digital elevation model (DEM) analysis, which calculates potential solar radiation at every point in the landscape as a function of slope angle, aspect, latitude, and horizon shading — these calculations reveal that optimal slope orientation varies with latitude, shifting from south-facing at high latitudes (>45°N, maximizing radiation) through southwest-facing at mid-latitudes (40-45°N, balancing radiation and afternoon heat) to east-facing at low latitudes (<35°, avoiding excessive afternoon radiation and heat stress).
Hangbeschreibung
Hanglagen-Dokumentation
Weinberg-Neigungscharakterisierung
hillside description
topographic inclination record
vineyard slope characterization
van Leeuwen C and Seguin G (2006) The concept of terroir in viticulture, Journal of Wine Research 17: 1-10 Gladstones J (1992) Viticulture and Environment, Winetitles Jackson DI and Schuster DF (2001) The Production of Grapes and Wine in Cool Climates, Gypsum Press Jones GV et al. (2004) Spatial analysis of climate in winegrape growing regions, American Journal of Enology and Viticulture 55: 304-316
Slope Description
The slope position of the Grand Cru vineyards on the Côte de Nuits (Burgundy) documents a remarkably consistent mid-slope position at 250-300 m elevation — situated above the deep, fertile, clay-rich valley floor soils (where the Bourgogne regional appellation vineyards are planted) and below the shallow, rocky, limestone hilltop soils (where forest cover replaces viticulture above the escarpment crest at 350-400 m), the Grand Cru vineyards occupy the mid-slope thermal belt where the soil depth is moderate (40-80 cm of clay-limestone over bedrock), drainage is excellent (gravitational water removal both over and through the thin soil profile), and the mesoclimate benefits from cold air drainage away from the vine zone to the valley floor below. This mid-slope concentration of the highest-quality vineyards — Romanée-Conti, La Tâche, Chambertin, Clos de Vougeot — reflects the empirical knowledge accumulated over a millennium of Burgundian viticulture that the mid-slope position optimizes the balance between soil fertility (sufficient but not excessive), water availability (adequate but naturally limited), and thermal protection (frost-free thermal belt).
A slope description component that records the location of the vineyard within the hillside landscape profile — classifying the position as hilltop/summit, upper slope, mid-slope, lower slope, toe of slope, or valley floor — with slope position determining the soil depth (typically increasing from shallow hilltop soils to deep colluvial toe-slope soils), the drainage characteristics (upper slopes shedding water, lower slopes receiving water from upslope), the frost risk (cold air pooling on valley floors, mid-slope thermal belts being frost-free), and the exposure to wind (hilltops and upper slopes being wind-exposed, lower slopes being sheltered).
Slope_Position documents the landscape context that integrates multiple environmental gradients — soil depth, soil fertility, drainage, temperature, frost risk, and wind exposure — into a single topographic descriptor that has been recognized as a primary terroir differentiator since the earliest systematic vineyard classifications. The concept of the thermal belt (also called the thermal zone or frost-free zone) is the most practically significant aspect of slope position — during radiation frost events (clear, calm nights when the ground surface radiates heat to the sky and the air in contact with the ground cools below freezing), cold air is denser than warm air and flows downslope under gravity, pooling in valley floors and depressions while the mid-slope zone remains above the frost temperature: this cold air drainage creates a temperature inversion where the mid-slope air temperature can be 3-8°C warmer than the valley floor temperature during frost events, producing the frost-free thermal belt that is the preferred vineyard position in all frost-prone viticultural regions. The soil catena — the systematic variation of soil properties along the slope from hilltop to valley floor — is the second critical gradient associated with slope position — upper slopes have shallow, rocky, low-fertility soils (derived from in situ bedrock weathering with minimal colluvial addition), mid-slopes have moderate soil depth with a mixture of residual and colluvial material, and lower slopes and toe positions have deep, fertile, fine-textured colluvial soils accumulated from upslope erosion over geological time — this catenary gradient directly affects vine vigor, root depth, water availability, and the degree of natural growth limitation that contributes to fruit quality.
Hangposition
Katena-Position
Landschaftslage
catenary position
hillside zone classification
landscape position record
van Leeuwen C and Seguin G (2006) The concept of terroir in viticulture, Journal of Wine Research 17: 1-10 Gladstones J (1992) Viticulture and Environment, Winetitles Wilson JE (1998) Terroir — The Role of Geology, Climate, and Culture in the Making of French Wines, University of California Press Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873
Slope Position
Cabernet Franc grafted on SO4 rootstock showing sloughing border cells in rhizosphere around primary root tip, microscopic examination revealing 420 border cells per root tip embedded in mucilage matrix extending 60-120 micrometers from cap surface, newly sloughed cells 28 micrometers diameter showing initial viability with FDA fluorescence, older border cells showing progressive senescence and cytoplasmic degradation, mucilage matrix containing polysaccharides and cellular debris, sloughing border cells creating defensive barrier and contributing organic matter to rhizosphere, border cell population continuously renewed through ongoing sloughing
Sloughing border cells are border cells being shed from root cap surface or recently shed existing as free cells in immediate rhizosphere, emphasizing release and dispersal process, characterized by complete separation from parent cap with no remaining physical connections, initial viability immediately after sloughing showing intact membranes and continued metabolism, mucilage embedding creating gelatinous matrix suspending cells around root tip, and progressive senescence over hours to days after sloughing leading to death and decomposition, with sloughing representing terminal phase in cap cell developmental trajectory where cells complete transition from attached tissue to independent dispersed cells, showing production dynamics of continuous steady sloughing maintaining border cell population as root grows through soil, spatial distribution creating cellular halo around root tip with highest density immediately adjacent to cap surface declining with distance, and functional roles including pathogen deterrence, soil conditioning, and organic matter contribution, and demonstrating that sloughing creates dynamic renewable interface between root and soil with border cell population continuously refreshed through ongoing production and loss. Sloughing border cells release dynamics includes= showing continuous shedding (border cells continuously sloughing from cap surface as peripheral cells complete separation), exhibiting production rate balance (sloughing rate balancing border cell production maintaining steady population), demonstrating spatial patterns (sloughing occurring around entire cap periphery though potentially concentrated in certain regions), revealing temporal variation (sloughing rate potentially varying with time of day, root growth rate, or environmental conditions), and displaying root growth correlation (sloughing potentially increasing with faster root growth as more cap cells produced and displaced). Sloughing border cells dispersal includes= showing initial proximity (newly sloughed cells remaining near cap surface where released), exhibiting gradual dispersal (border cells moving away from cap through soil movement, water flow, or root growth displacement), demonstrating mucilage suspension (mucilage matrix keeping border cells suspended around root tip rather than sedimenting), revealing spatial distribution (border cells creating halo around root tip with density declining with distance from cap), and displaying accumulation (border cells accumulating in root tip zone creating substantial cell population particularly in slow-growing roots). Sloughing border cells viability trajectory includes= showing initial viability (newly sloughed cells typically viable with functional metabolism), exhibiting progressive decline (border cell viability declining over hours to days after sloughing), demonstrating variable lifespan (border cells living hours to days in rhizosphere depending on conditions), revealing senescence progression (border cells showing orderly senescence with progressive cytoplasmic degradation), and displaying eventual death (border cells dying and decomposing contributing cellular contents to rhizosphere). Sloughing border cells mucilage interactions includes= showing mucilage embedding (sloughed cells embedded in thick mucilage matrix creating gelatinous environment), exhibiting continued secretion (viable border cells continuing to secrete mucilage after sloughing), demonstrating mucilage network (collectively secreted mucilage creating interconnected network around root tip), revealing matrix properties (mucilage matrix showing viscoelastic properties affecting border cell distribution and function), and displaying defensive matrix (mucilage creating physical barrier potentially trapping pathogens and affecting chemical environment). Sloughing border cells defensive deployment includes= providing pathogen barrier (sloughed border cells creating defensive zone around sensitive cap and meristem), exhibiting trap function (mucilage and cells physically trapping fungal spores and bacteria), demonstrating decoy function (border cells attracting pathogen colonization diverting from cap and meristem), revealing chemical defense (border cells secreting antimicrobial compounds into rhizosphere), and displaying expendable defense (sloughed cells representing sacrificial defensive layer that can be damaged or colonized without root injury). Sloughing border cells in rhizosphere biology includes= showing microbial substrate (dead and dying border cells providing readily available carbon and nutrients for rhizosphere microbes), exhibiting microbial interactions (border cells interacting with beneficial and pathogenic microbes), demonstrating community effects (border cells affecting rhizosphere microbial community composition and function), revealing carbon input (border cell carbon contributing to rhizosphere carbon budget and soil organic matter), and displaying nutrient release (border cell decomposition releasing nitrogen, phosphorus, and other nutrients potentially available for reuptake). Sloughing border cells quantification includes= showing production estimates (quantifying border cell sloughing rates per root tip per day), exhibiting population measurements (counting border cells in rhizosphere determining population sizes), demonstrating viability assessment (determining proportion of sloughed cells that are viable versus dead), revealing spatial distribution mapping (documenting border cell density versus distance from root tip), and displaying temporal dynamics (measuring border cell population changes over time). Sloughing border cells species and environmental variation includes= showing species differences (sloughing rates and border cell characteristics varying among plant species), exhibiting rootstock effects (grapevine rootstock varieties potentially differing in border cell sloughing), demonstrating environmental modulation (soil conditions, temperature, moisture affecting sloughing rate and border cell survival), revealing pathogen stimulation (pathogen presence potentially stimulating increased border cell production and sloughing), and displaying stress responses (environmental stresses potentially affecting border cell sloughing and function).
Sloughing_Border_Cells emphasizes release process and dispersed state and is distinguished from attached or detaching cells by focus on shed cells in rhizosphere. Critical for understanding continuous shedding, spatial dispersal, viability trajectory, mucilage embedding, defensive deployment, rhizosphere contribution, and the released border cells creating dynamic renewable interface providing defense and conditioning rhizosphere environment around root tip.
Abgestoßene Grenzzellen
Dispersing border cells
Freigesetzte Haubenzellen
Released cap cells
Shed border cells
Sich verteilende Grenzzellen
Sloughing Border Cells
A slow expansion young leaf at node 11 on a water-stressed shoot of Vitis vinifera cv. Tempranillo in Rioja, Spain, exhibiting area increase from 18 cm² to 38 cm² over 10 days representing daily growth rate of 3.2 cm² per day (4.1% daily compound growth), dimensions increasing from 5.8 cm × 5.2 cm to 8.5 cm × 7.8 cm over the period, cells in palisade layers enlarging from 20 μm to 25 μm diameter representing 5 μm total increase or only 0.5 μm per day, tissue water content of 84% fresh weight indicating moderate water stress limiting turgor and expansion, somewhat firm texture with accelerated wall thickening evident from reduced flexibility, pale green coloration with chlorophyll at 245 μg cm⁻² below optimal levels for this expansion stage, slow growth requiring caliper measurements to detect daily progression, reduced resource consumption of approximately 15 mg carbohydrate and 2.8 mg nitrogen per day reflecting both slow growth and limited resource availability, photosynthetic rate of only 4.2 μmol CO₂ m⁻² s⁻¹ representing 48% of mature capacity and remaining strongly net sink, occurring under water stress conditions with soil moisture at 48% field capacity (predawn leaf water potential -0.8 MPa), daily maximum temperatures 32-34°C exceeding optimal range, measured at shoot BBCH stage 77 (berries beginning to touch) in mid-July, representing reduced expansion rate typical of moderately stressed conditions that will result in smaller final mature leaf size compared to unstressed potential.
An expanding young leaf exhibiting reduced daily growth rates due to suboptimal expansion conditions or late-season development in Vitis vinifera, characterized by low area increase rates typically 2-5 cm² per day representing 2-4% daily compound growth rate, dimensions progressing slowly from approximately 25% to 60% of potential final size over extended 18-28 day period, reduced cell expansion rates of 0.5-1.5 μm per day in linear dimensions, moderate tissue water content of 82-87% fresh weight reflecting limited water availability or reduced water uptake capacity constraining turgor-driven expansion, somewhat firm texture from accelerated cell wall thickening relative to expansion creating stiffer tissue earlier in development, pale to medium green coloration with chlorophyll accumulation lagging behind area expansion due to resource limitation or stress, slow visible growth progression requiring measurements to detect daily changes, reduced daily resource consumption including 10-20 mg carbohydrate per day and 2-3 mg nitrogen per day reflecting both slower growth and resource limitation, and photosynthetic capacity increasing slowly reaching only 40-60% of mature leaf potential by end of slow expansion phase with continued net sink status throughout expansion period. Slow expansion young leaves develop under limiting conditions including suboptimal temperatures of 15-20°C or 30-35°C reducing metabolic rates and enzyme activity, limited soil moisture with field capacity at 40-60% creating water stress that limits turgor-driven expansion, inadequate nitrogen availability restricting protein synthesis and chlorophyll production, excessive light causing photoinhibition of developing chloroplasts and photooxidative stress, or low shoot vigor with limited resource translocation from mature tissues creating developmental resource limitation.
Slow expansion young leaves are distinguished from rapid expansion leaves by low daily growth rates, extended expansion duration, reduced resource consumption, and development under limiting conditions. Slow expansion typically results from environmental limitations rather than genetic programming, with water deficit being the most common cause reducing turgor pressure required for cell wall extension. Temperature effects on expansion rate are substantial, with cool temperatures (15-20°C) slowing metabolic processes and warm temperatures (>30°C) increasing respiratory costs and potentially causing direct heat stress to expanding tissues. Nitrogen deficiency reduces expansion rate and final size through limitation of protein synthesis required for membrane production, enzyme synthesis, and chloroplast development. Slow expansion creates smaller mature leaves with potentially altered anatomy including thicker cell walls, reduced mesophyll development, and lower chlorophyll concentrations, affecting final photosynthetic capacity. Extended slow expansion duration increases cumulative risk of damage from pests, diseases, or environmental stresses during the prolonged vulnerable expansion period.
constrained growth leaf
eingeschränktes Wachstum Weinblatt
langsam-expandierendes junges Blatt
langsamwachsendes Blatt
reduced expansion rate leaf
slow-growing leaf
Based on Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Chaves et al. (2010) "Grapevine under deficit irrigation= hints from physiological and molecular data" Annals of Botany 105=661-676 Palliotti et al. (2012) "Morpho-anatomical and physiological response of Pinot Noir and Montepulciano grapevine leaves to mild and severe water stress" Vitis 51=109-116 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 161-184.
Slow Expansion Young Leaf
Pinot Noir exhibits characteristic small cluster disposition with average weights of 80-140 g, rachis lengths of 8-12 cm, and berry counts of 60-120 per cluster. Gamay produces small cylindrical clusters (90-150 g) ideally suited for whole-cluster fermentation in Beaujolais. Nebbiolo develops small to medium clusters (100-180 g) with loose architecture, considered optimal for Barolo and Barbaresco production. Small clusters in premium wine cultivars typically show higher skin-to-pulp ratios, with Pinot Noir at 12-15% skin weight versus 8-10% for large-clustered cultivars.
A cluster size disposition that is the inherited capacity of a Vitis vinifera plant to develop clusters of below-average weight (typically less than 150 g) and compact dimensions, with limited rachis branching, moderate berry count, or small individual berry size resulting in reduced cluster mass.
Small cluster disposition often associates with premium wine quality perception due to higher concentration of skin-derived compounds (anthocyanins, tannins, aromatics) per unit juice volume. The trait reduces yield potential, requiring higher cluster numbers per vine or acceptance of lower production levels. Small clusters facilitate whole-cluster fermentation techniques and may improve ripening uniformity. Clone selection within small-clustered cultivars can identify variants with modestly larger clusters while retaining quality attributes.
Kleintraubigkeit
kleine Traube
leichte Traube
light cluster disposition
petite bunch capacity
small bunch trait
Matthews MA, Nuzzo V (2007) Berry size and yield paradigms on Vitis vinifera wine quality. Acta Horticulturae 754:423-436 Roby G et al. (2004) Berry size and vine water deficits as factors in winegrape composition: anthocyanins and tannins. Australian Journal of Grape and Wine Research 10:100-107
Small Cluster Disposition
Characterization of Small_Leaf_Disposition at the University of Bordeaux documented leaf development in cultivars recognized for compact foliage. Reference cultivars 'Petit Manseng', 'Petit Verdot', 'Pinot Noir', and 'Riesling' were assessed under standardized conditions with detailed morphometric and functional analysis. Small_Leaf_Disposition cultivars showed consistent characteristics. 'Petit Manseng' exhibited archetypal small leaves with mature leaf area of 92 cm² (mean of nodes 9-11), blade length of 10.8 cm and width of 11.2 cm, achieving 90% of final area by only 26 days after emergence. The name "Petit" directly references the characteristically small foliage. 'Petit Verdot' showed similarly small leaves (105 cm²) with compact, deeply-dissected morphology. 'Pinot Noir' displayed typical Small_Leaf_Disposition (125 cm²) with moderately thick lamina partially compensating for reduced area. 'Riesling' showed small-to-moderate size (138 cm²). Canopy analysis revealed that Small_Leaf_Disposition cultivars at 15 shoots/m achieved LAI of only 2.2 m²/m² by mid-season, below optimal range and potentially limiting photosynthetic capacity. However, canopy light distribution was substantially more uniform, with interior leaf illumination averaging 45% of ambient compared to 18% in large-leaved canopies at equivalent LAI. Area-normalized photosynthetic rates were actually 15% higher in Small_Leaf_Disposition cultivars, partially compensating for lower total area. The correlation between Small_Leaf_Disposition and premium wine cultivars was notable—8 of the 10 highest-ranked Bordeaux varieties showed leaf areas below 160 cm², suggesting that moderate leaf size contributes to the canopy characteristics associated with wine quality.
A Small_Leaf_Disposition is a Leaf_Size_Disposition that inheres in Vitis vinifera leaves and is realized through abbreviated cell division and limited expansion producing individual mature leaf areas below 150 cm², creating compact foliage that accumulates canopy leaf area more gradually but provides more uniform light distribution within the canopy. This disposition represents the lower range of leaf size, characteristic of cultivars that produce numerous smaller leaves rather than fewer large leaves to achieve total canopy area. Small-leaved cultivars develop modest individual leaves that reach full expansion over approximately 25-30 days, with lower cell number and smaller cell size combining to produce compact leaf blades. The viticultural implications of Small_Leaf_Disposition include gradual canopy development (more leaves required to achieve target leaf area), naturally lower canopy density at standard shoot densities (potentially advantageous for fruit exposure), reduced self-shading within the canopy, thinner boundary layer improving gas exchange efficiency, and distinct visual character with finer-textured foliage. Small leaves may be disadvantageous in low-vigor situations where limited shoot growth produces insufficient canopy area, but can be advantageous in high-vigor situations by moderating canopy density. Small_Leaf_Disposition is often associated with premium wine grape cultivars, potentially reflecting selection for moderate canopies suited to quality-focused production systems, and with cultivars from cooler climates where compact canopy architecture may reduce disease pressure.
Small_Leaf_Disposition is_a Leaf_Size_Disposition representing individual leaf area < 150 cm². Characterized by abbreviated cell division phase and limited cell expansion. Examples include 'Petit Manseng', 'Petit Verdot', 'Pinot Noir', 'Riesling', 'Gamay'. Common among premium wine grape cultivars. Advantages include more uniform canopy light distribution, reduced shading, and potentially higher area-normalized photosynthesis. May require higher shoot density or vigor-enhancing management to achieve adequate canopy area. OIV descriptor 065 codes 1-3.
Kleinblättrige-Disposition
Kleinblättrige-Sorte
Niedrige-Einzelblattfläche
compact foliage
low individual leaf area
small-leaved cultivar
Galet P (1979) A Practical Ampelography: Grapevine Identification, Cornell University Press Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought, Plant Cell and Environment 26:1393-1405 Organisation Internationale de la Vigne et du Vin (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition
Small Leaf Disposition
A Shiraz vine on vigorous Barossa floor site trained to Smart-Dyson bears a Smart_Dyson_Trained_Role with a single bilateral cordon at 90 cm and a moveable wire that lifts to separate upward and downward shoot growth, the simplified architecture managing vigor with less infrastructure than full Scott Henry. A Cabernet Sauvignon vine in Margaret River serves in a Smart_Dyson_Trained_Role, the single-cordon divided approach enabling quality production from fertile sites while minimizing conversion costs from existing bilateral cordon infrastructure. A Merlot vine on productive Hawkes Bay soils bears a Smart_Dyson_Trained_Role, the Ballerina wire technique creating vertical canopy division through labor rather than permanent infrastructure modification.
A training system role that inheres in a grapevine trained to the Smart-Dyson system (also termed Smart-Dyson Ballerina), a simplified vertically divided canopy developed by Richard Smart and John Dyson as an easier-to-manage alternative to Scott Henry, with shoots from a single cordon divided at a moveable wire that directs alternate shoots upward and downward. Smart-Dyson training reduces the infrastructure complexity of Scott Henry (which requires two cordons or cane positions) by using a single cordon with shoot division achieved through canopy management rather than structural positioning. The system employs a moveable "lift wire" that is raised during the growing season to capture upward-growing shoots while allowing alternate shoots to grow downward, creating the vertically divided architecture with less permanent infrastructure. Smart-Dyson represents the evolution of divided canopy thinking toward practical implementation, recognizing that simpler systems achieving similar results will see greater adoption than theoretically optimal but complex alternatives.
Smart_Dyson_Trained_Role captures the pragmatic simplification of divided canopy principles developed by Richard Smart, whose "Sunlight into Wine" manual disseminated canopy management concepts globally. The system's "Ballerina" designation reflects the shoot separation technique, with the moveable wire "lifting" shoots like a dancer's arms. Smart-Dyson adoption has been modest, partly because the shoot division requires consistent canopy management attention that may exceed the labor saved from simplified infrastructure. The system represents an intermediate approach—more complex than single-curtain VSP, simpler than full Scott Henry—that suits operations seeking vigor management without major trellis reconstruction. Contemporary viticultural practice increasingly emphasizes simplicity, with many producers preferring to address vigor through rootstock selection, cover crops, or deficit irrigation rather than complex training systems.
Ballerina-System
Smart-Dyson
Smart-Dyson Ballerina
Smart-Dyson-Erziehung
simplified Scott Henry
single-cordon divided
vereinfachtes Scott-Henry
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles Smart RE (1988) Shoot spacing and canopy light microclimate, American Journal of Enology and Viticulture 39: 109-115 Reynolds AG, Vanden Heuvel JE (2009) Training systems, American Journal of Enology and Viticulture 60: 251-268
Smart Dyson Trained Role
Smooth surface quality in Sultanina berry skins facilitates the production of premium golden raisins by enabling uniform drying rates across the berry surface and consistent sulfur dioxide penetration during treatment berries with rough surface texture develop uneven color and moisture content during drying that produces mottled raisins with reduced market value, making smooth skin texture a selection criterion in Sultanina clonal evaluation programs alongside seedlessness, berry size, and drying ratio.
Smooth_Surface_Quality represents the surface texture condition where the external boundary of a grapevine organ lacks prominent tactile irregularities, presenting an even surface that does not catch or resist when touched. Smooth surfaces result from regular arrangement of epidermal cells without protruding features, uniform cuticle deposition, and absence of trichomes or other surface appendages. In grape berries, smooth skin texture characterizes most Vitis vinifera cultivars when the waxy bloom is disregarded, with the underlying berry surface presenting an even, continuous cuticle without ridges, bumps, or rough patches. Berry skin smoothness facilitates cleaning and handling, promotes uniform wax distribution, and presents an attractive appearance for table grapes. Some cultivars, particularly certain Muscadinia and Vitis hybrid origins, may display rougher berry surfaces with visible lenticels or cuticular ridging. Leaf surfaces vary in smoothness, with glabrous cultivars presenting smooth upper (adaxial) surfaces while retaining some texture from vein prominence. Cane surfaces at maturity display smooth bark between nodes in most cultivars, though lenticel density and bark texture vary. Assessment of smooth surface quality employs tactile evaluation detecting resistance to finger movement across the surface, visual observation under oblique lighting that reveals surface irregularities through shadows, and profilometric measurement of surface roughness parameters (Ra, Rz) that quantify deviation from a perfectly flat surface.
Smooth_Surface_Quality represents one pole of the smooth-rough continuum and is assessed relative to the typical surface texture of the organ type. Berry surfaces are generally smoother than leaf surfaces due to the absence of prominent venation. The distinction between smooth and rough surfaces involves threshold judgments quantitative roughness measurements (e.g., arithmetic mean roughness Ra below 1 μm) can provide objective criteria. Smooth surfaces may still possess fine-scale texture features visible only under magnification the smooth classification addresses the tactile and visually perceptible scale.
ebene Textur
even surface
glatte Oberfläche
glatte Oberflächenqualität
gleichmäßige Oberflächenbeschaffenheit
regular surface
sleek texture
untextured surface quality
Casado CG and Heredia A (2001) Ultrastructure of the Cuticle during Growth of the Grape Berry (Vitis vinifera L.). Physiologia Plantarum 111(2): 220-224 Rosenquist JK and Morrison JC (1989) Some Factors Affecting Cuticle and Wax Accumulation on Grape Berries. American Journal of Enology and Viticulture 40(4): 241-244
Smooth Surface Quality
Smoothness quality evolution in Brunello di Montalcino demonstrates the aging trajectory from young wine (3 years, smoothness 4/10 with detectably grippy tannin despite quality fruit) through developing wine (8 years, smoothness 7/10 with integrated tannin and polished mouthfeel) to mature wine (15 years, smoothness 9/10 with seamlessly integrated components creating effortless, silky texture), illustrating how smoothness development indicates both aging progress and quality potential, with wines lacking smoothness capacity at youth unlikely to achieve textural refinement regardless of cellaring duration.
Smoothness quality is a mouthfeel quality that represents the absence of tactile roughness, harshness, or irritation during wine consumption, characterizing the gentle, polished, friction-free oral sensation that indicates refined tannin integration, appropriate acid balance, and overall textural harmony. Within the BFO framework, this quality specifically addresses the friction dimension of mouthfeel, distinguishing smooth texture from rough or harsh texture while recognizing that smoothness represents a positive textural attribute arising from the absence of irritating or catching sensations on oral surfaces. The ontological significance lies in smoothness serving as a primary quality indicator across wine styles, where smooth mouthfeel suggests mature tannins, well-integrated components, and careful winemaking, while harsh or rough texture indicates either youth requiring further aging or quality deficiency from unbalanced or aggressive components. The perceptual basis for smoothness involves the absence of mechanoreceptor stimulation that would indicate surface irregularity or friction, with smooth wines sliding across oral surfaces without catching, dragging, or irritating, creating the polished, refined tactile impression valued in quality wine. The compositional determinants of smoothness include tannin quality and polymerization state (mature, polymerized tannins feeling smoother than young, reactive tannins), polysaccharide concentration (mannoproteins and arabinogalactans significantly enhancing perceived smoothness), glycerol level (contributing to lubrication), acid balance (excessive acidity creating harshness), and the overall integration of components where no single element dominates to create textural disruption. The developmental aspects of smoothness show wines typically gaining smoothness with aging as tannins polymerize and components integrate, with bottle age one path to smoothness while winemaking techniques (micro-oxygenation, lees contact, appropriate fining) can accelerate smooth texture development. Understanding smoothness quality enables quality assessment based on textural refinement, aging trajectory prediction from smoothness development, and winemaking evaluation regarding techniques affecting textural polish.
Smoothness_Quality addresses the friction-free dimension of mouthfeel, representing textural refinement and integration. This quality is a positive attribute indicating mature, well-integrated wine components. The class supports quality assessment and aging prediction, with smoothness development trajectory indicating wine quality potential and optimal drinking window.
Geschmeidigkeit
Texturraffinesse
poliertes Mundgefühl
polished texture
silky mouthfeel
textural elegance
Vidal S et al. (2004) Use of an experimental design approach for evaluation of key wine components on mouth-feel perception. Food Quality and Preference 15(3): 209-217 Chira K et al. (2011) Chemical and sensory evaluation of Bordeaux wines according to grape quality and tannin structure. Journal of the Science of Food and Agriculture 91(10): 1748-1754
Smoothness Quality
The SNP array data for a diversity panel of 783 Vitis vinifera cultivars genotyped on the Vitis18K Illumina Infinium array (Laucou et al. 2018) comprises 18,071 SNP probe intensities per sample converted through GenomeStudio clustering algorithms into diploid genotype calls (AA, AB, BB) with associated GenCall quality scores — after quality filtering (removing SNPs with call rate below 95%, minor allele frequency below 1%, or significant deviation from Hardy-Weinberg equilibrium), 14,748 high-quality SNP genotypes per accession provide the data matrix for principal component analysis revealing the three major genetic pools of cultivated grapevine (Western European wine grapes, Eastern Mediterranean table grapes, and Muscat-type cultivars), STRUCTURE analysis of population admixture, and GWAS scans identifying significant marker-trait associations for berry color, seedlessness, muscat aroma, and disease resistance.
An SNP genotype component that records the raw and processed output from hybridization-based SNP genotyping array platforms, documenting the signal intensities, genotype cluster assignments, and quality metrics generated by the simultaneous interrogation of thousands to tens of thousands of predefined SNP positions in a single assay, providing the high-throughput genotyping data that underlies genome-wide association studies, genomic selection, and population genetics analyses in Vitis vinifera.
SNP_Array_Data captures the technology-specific genotyping output from fixed-content array platforms designed for Vitis. The development of grape-specific SNP arrays progressed through three generations — the Vitis9KSNP array (Myles et al. 2011, 8,898 SNPs selected from Pinot Noir and Cabernet Sauvignon resequencing data), the GrapeReSeq Vitis18K array (Le Paslier et al. 2013, 18,071 SNPs selected from resequencing of 47 cultivars representing global diversity), and the Vitis50K array (approximately 50,000 SNPs providing higher genome coverage for fine-mapping applications): each generation increased SNP density and diversity panel representation, improving the ascertainment of genetic variation across the full spectrum of cultivated and wild Vitis germplasm. The array platform determines the SNP content that can be interrogated — a fundamental limitation compared to sequencing-based approaches (GBS, whole-genome resequencing) that discover novel variants in each sample — making array data most valuable for standardized genotyping of known variants across large sample sets where cost-effective, uniform data quality is prioritized over novel variant discovery.
Genotypisierungs-Chip-Daten
SNP chip data
SNP microarray results
SNP-Array-Daten
SNP-Microarray-Ergebnisse
genotyping array output
Laucou V et al. (2018) An extended haplotype reference panel and high-density SNP genotyping data for grapevine, Scientific Data 5: 180259 Myles S et al. (2011) Genetic structure and domestication history of the grape, Proceedings of the National Academy of Sciences 108: 3530-3535 Le Paslier MC et al. (2013) The GrapeReSeq 18K Vitis genotyping chip, Acta Horticulturae 1082: 205-210
SNP Array Data
The SNP genotype data for Chardonnay generated on the Vitis18K SNP array (Laucou et al. 2018) comprises genotype calls at 18,071 biallelic SNP positions distributed across all 19 chromosomes at an average density of one SNP per 26 kb — the resulting high-density genotype profile provides discrimination power several orders of magnitude greater than the standard 9-locus SSR panel, enabling not only cultivar identification but also intra-cultivar clonal discrimination (detecting the approximately 200-500 somatic SNP mutations that distinguish long-propagated clones such as Chardonnay Musqué from standard Chardonnay), parentage and pedigree reconstruction across multiple generations, and genomic prediction of quantitative traits through marker-trait associations established in training populations.
A genotype data record based on single nucleotide polymorphism (SNP) markers — positions in the genome where a single base pair differs between alleles or between individuals — documenting the biallelic or multiallelic state at defined SNP positions that collectively provide high-resolution molecular characterization for cultivar identification, genomic selection, genome-wide association studies, and population structure analysis in Vitis vinifera.
SNP_Genotype documents the marker system that has progressively supplemented and in many applications superseded SSR genotyping in grapevine molecular genetics since the 2010s. SNP markers offer several technical advantages over SSRs — they are far more abundant in the genome (millions of SNPs versus thousands of SSRs), amenable to high-throughput automated genotyping on array platforms, directly interpretable in terms of sequence context, and less subject to the allele sizing artifacts that complicate SSR data cross-referencing: however, individual SNPs are less informative than individual SSRs (biallelic versus multiallelic, with expected heterozygosity of 0.3-0.5 versus 0.7-0.9 per locus), requiring panels of hundreds to thousands of SNPs to achieve the discrimination power of 9 SSR loci. The development of Vitis-specific SNP arrays (Vitis9KSNP, Vitis18K, Vitis50K) and the decreasing cost of genotyping-by-sequencing (GBS) have made SNP genotyping the standard approach for genomic research applications while SSR genotyping remains the standard for routine cultivar identification due to its established database infrastructure and lower per-sample cost for small sample numbers.
Einzelnukleotid-Polymorphismus-Genotyp
SNP marker genotype
SNP-Genotyp
SNP-based molecular profile
SNP-basiertes Markerprofil
single nucleotide variant data
Laucou V et al. (2018) An extended haplotype reference panel and high-density SNP genotyping data for grapevine, Scientific Data 5: 180259 Myles S et al. (2011) Genetic structure and domestication history of the grape, Proceedings of the National Academy of Sciences 108: 3530-3535 Le Paslier MC et al. (2013) The GrapeReSeq 18K Vitis genotyping chip, IX International Symposium on Grapevine Physiology and Biotechnology, Acta Horticulturae 1082: 205-210
SNP Genotype
The SNP genotype call at position chr2:14,240,287 (the causal SNP within the VvMYBA1 promoter region controlling anthocyanin biosynthesis activation) for Pinot Noir reports the heterozygous state G/T with a GenCall quality score of 0.92 — the G allele is associated with the functional VvMYBA1 transcription factor driving anthocyanin production in berry skin (red/black phenotype), while the T allele marks the Gret1 retrotransposon insertion that inactivates VvMYBA1 expression. Pinot Noir's heterozygous G/T state at this position explains its capacity to produce color-loss somatic mutations (Pinot Blanc, Pinot Gris) through loss of the functional G allele haplotype, while homozygous G/G cultivars (e.g., Syrah) cannot produce white-berry sports through single-allele loss and homozygous T/T cultivars (e.g., Chardonnay) constitutively lack anthocyanin production.
An SNP genotype component that records the determined allelic state at a single SNP position in a specific grapevine accession, reporting one of three possible diploid genotype classes — homozygous reference (e.g., AA), heterozygous (e.g., AB), or homozygous alternate (e.g., BB) — together with the associated confidence metric that quantifies the reliability of the allele assignment based on signal intensity, cluster separation, or read depth at that position.
SNP_Genotype_Call is the atomic data unit of SNP genotyping — the single-position allele determination that, aggregated across thousands of positions, constitutes the SNP genotype profile. The reliability of individual genotype calls depends on the genotyping technology and the quality metrics applied: array-based genotyping assigns calls through clustering of normalized signal intensities in two-dimensional space (one axis per allele), with the GenCall or GenTrain score (ranging from 0 to 1) reflecting the distance of the sample's signal from the nearest genotype cluster center — calls with scores below 0.15-0.25 are typically classified as missing data rather than risking incorrect assignment. Sequencing-based genotype calls (from GBS or whole-genome resequencing) are assigned through variant calling algorithms (GATK, samtools) that evaluate read depth, base quality, mapping quality, and strand bias, with genotype quality (GQ) scores on a Phred scale where GQ 30 indicates 99.9% call accuracy. The functional significance of individual SNP genotype calls ranges from neutral (the majority of SNPs in non-coding regions) through quantitative trait modifiers (SNPs in regulatory regions affecting gene expression levels) to qualitative trait determinants (SNPs in coding regions altering protein function, as exemplified by the VvMYBA1 berry color locus).
Einzelpositions-Allelbestimmung
SNP allele call
SNP variant call
SNP-Genotyp-Aufruf
SNP-Varianten-Zuordnung
single-position genotype determination
Laucou V et al. (2018) An extended haplotype reference panel and high-density SNP genotyping data for grapevine, Scientific Data 5: 180259 Kobayashi S et al. (2004) Retrotransposon-induced mutations in grape skin color, Science 304: 982 Walker AR et al. (2007) White grapes arose through the mutation of two similar and adjacent regulatory genes, Plant Journal 49: 772-785
SNP Genotype Call
The SNP haplotype analysis of the Rpv3 downy mildew resistance locus on chromosome 18 in the interspecific hybrid cultivar Regent identifies a 400 kb haplotype block containing 12 linked SNPs derived from the American species Vitis rupestris through the breeding lineage Chambourcin × Diana — this resistance-associated haplotype is absent from all pure Vitis vinifera cultivars and can be traced through Regent's pedigree to confirm the introgression pathway, with haplotype-based marker-assisted selection enabling breeders to select for the complete resistance locus (rather than individual SNPs that may be separated from the functional resistance genes by recombination) in breeding populations segregating for Rpv3 resistance. The extended haplotype homozygosity (EHH) pattern around Rpv3 in resistant breeding lines shows reduced diversity compared to neutral genomic regions, consistent with the selective sweep expected for a locus under strong artificial selection.
An SNP genotype component that records the specific combination of alleles at multiple linked SNP positions along a single chromosome segment, representing the phased sequence of variants inherited together as a unit from one parent, with haplotype data providing finer genetic resolution than individual unphased genotype calls for parentage reconstruction, introgression detection, and the identification of extended haplotype blocks associated with phenotypic traits in Vitis vinifera.
SNP_Haplotype adds the critical dimension of allelic phase to SNP genotype data — determining which alleles at linked positions were inherited together on the same chromosomal copy (cis configuration) versus on opposite copies (trans configuration). Haplotype phasing is essential for three major applications in grapevine genetics: first, parentage and pedigree analysis, where the inheritance of specific parental haplotypes through crosses provides definitive parentage confirmation and enables reconstruction of multi-generational pedigrees second, introgression detection in interspecific hybrid breeding, where American Vitis species haplotype blocks carrying disease resistance loci (Rpv, Ren, Run loci) must be tracked through backcross generations to maintain resistance while recovering V. vinifera wine quality and third, haplotype-based association mapping, where extended haplotype blocks defined by linkage disequilibrium provide greater statistical power for trait association than individual SNPs. The high heterozygosity and extensive linkage disequilibrium of V. vinifera (LD extending 10-40 kb depending on genomic region and population) create haplotype block structures that are informative for association mapping but require substantial marker density for accurate phasing.
SNP-Haplotyp
chromosomal haplotype block
chromosomaler Allelverband
linked allele combination
phased SNP haplotype
phasierter Haplotyp-Block
Laucou V et al. (2018) An extended haplotype reference panel and high-density SNP genotyping data for grapevine, Scientific Data 5: 180259 Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance and ripening of grapevines assessed in a segregating population, Molecular Breeding 36: 65 Myles S et al. (2011) Genetic structure and domestication history of the grape, Proceedings of the National Academy of Sciences 108: 3530-3535
SNP Haplotype
The SNP marker data for the Vitis18K array (Le Paslier et al. 2013, Laucou et al. 2018) documents 18,071 biallelic SNP markers selected from whole-genome resequencing of 47 cultivars representing global diversity — each marker is characterized by its physical position on the PN40024 v1 reference genome (chromosome and base pair coordinate), the reference and alternate allele (e.g., chr2:14240287 G/T at the VvMYBA1 berry color locus), minor allele frequency in the discovery panel (range 0.05-0.50, median 0.28), functional annotation from the V1 gene prediction (34% intergenic, 28% intronic, 18% within 2 kb upstream of genes, 12% synonymous coding, 8% nonsynonymous coding), and Illumina Infinium probe design scores (design score >0.7 for all retained markers, ensuring reliable hybridization-based genotyping). A subset of 48 highly informative SNPs (MAF >0.35, PIC >0.45, distributed across all 19 chromosomes) has been proposed as a minimal SNP identification panel for routine cultivar verification as a potential successor to the 9-locus SSR standard.
A genetic marker data record specific to single nucleotide polymorphism markers developed or validated for Vitis, documenting the marker identity, chromosomal position, flanking sequence context, reference and alternate alleles, minor allele frequency across cultivar panels, functional annotation (synonymous, nonsynonymous, regulatory, intergenic), and genotyping platform compatibility that characterize each SNP as a tool for genome-wide association, genomic selection, and high-resolution cultivar characterization in grapevine.
SNP_Marker_Data documents the properties of individual SNP markers as genotyping tools, complementing SNP_Genotype (which records allelic states in specific accessions) and SNP_Array_Data (which records the platform-level output). The transition from SSR to SNP markers as the primary tool for grapevine genetic research reflects the broader trend in plant genetics toward high-density, sequence-based marker systems: SNP markers in Vitis have been discovered through multiple approaches including Sanger sequencing of candidate gene amplicons (pre-2007), alignment of whole-genome shotgun reads to the PN40024 reference (2007-2015), and population-scale resequencing projects (Liang et al. 2019, identifying over 37 million SNP positions across 472 accessions). The challenge of SNP ascertainment bias — the tendency for SNPs discovered in a limited panel of cultivars to inadequately represent genetic variation in undersampled populations — has been addressed through progressive expansion of discovery panels to include table grapes, wine grapes from all major geographic origins, wild V. vinifera subsp. sylvestris, and non-vinifera Vitis species used in rootstock and disease-resistant hybrid breeding.
Einzelnukleotid-Marker-Spezifikation
SNP locus characterization
SNP marker specification
SNP-Lokus-Charakterisierung
SNP-Marker-Daten
single nucleotide variant marker record
Laucou V et al. (2018) An extended haplotype reference panel and high-density SNP genotyping data for grapevine, Scientific Data 5: 180259 Le Paslier MC et al. (2013) The GrapeReSeq 18K Vitis genotyping chip, Acta Horticulturae 1082: 205-210 Liang Z et al. (2019) Whole-genome resequencing of 472 Vitis accessions, Nature Communications 10: 1190 Cabezas JA et al. (2011) A 48 SNP set for grapevine cultivar identification, BMC Plant Biology 11: 153
SNP Marker Data
1103 Paulsen (V. berlandieri × V. rupestris) demonstrates strong Na⁺ exclusion, maintaining Montepulciano leaf Na⁺ below 0.15% dry weight and K⁺/Na⁺ ratios above 15:1 under irrigation at ECw 2.0 dS/m in Abruzzo coastal vineyards, while Kober 5BB permits leaf Na⁺ accumulation to 0.4% with K⁺/Na⁺ ratios declining below 5:1. Ramsey (V. champinii) combines Na⁺ and Cl⁻ exclusion, maintaining Flame Seedless leaf Na⁺ at 0.10-0.15% in the high-sodium irrigation conditions of the Coachella Valley (SAR values of 4-6). Ruggeri 140 restricts Na⁺ translocation to Grillo in the saline coastal vineyards of western Sicily, with leaf K⁺/Na⁺ ratios maintained above 12:1 where susceptible rootstocks decline below 4:1.
A salt-tolerant rootstock role that inheres in a grafted rootstock by virtue of its specific capacity to restrict sodium ion (Na⁺) uptake by roots and translocation to the scion, thereby protecting scion foliage from sodium toxicity and maintaining favorable K⁺/Na⁺ ratios essential for cellular function. Sodium exclusion operates through mechanisms including selective K⁺/Na⁺ discrimination at root uptake (mediated by high-affinity K⁺ transporters with low Na⁺ permeability), Na⁺ efflux from root cells via SOS1-type Na⁺/H⁺ antiporters, Na⁺ sequestration in root vacuoles, and restricted Na⁺ loading into the xylem. Sodium toxicity in grapevine causes leaf margin burn progressing to interveinal necrosis, with toxicity thresholds at 0.25-0.5% Na⁺ dry weight in leaf tissue. Sodium exclusion is quantified by leaf Na⁺ concentration under saline irrigation and by maintenance of leaf K⁺/Na⁺ ratios above 10:1 (ratios below 5:1 indicating Na⁺ toxicity stress). Sodium exclusion is generally less variable among rootstocks than chloride exclusion, as most Vitis genotypes show moderate Na⁺ exclusion capacity.
Sodium_Excluding_Rootstock_Role addresses Na⁺-specific toxicity, which is quantitatively less limiting than Cl⁻ toxicity for most grapevine cultivars but becomes important in sodic soils (high SAR) and under conditions of Na⁺-dominated salinity. The K⁺/Na⁺ ratio is a critical functional parameter because Na⁺ competes with K⁺ for enzyme activation sites maintaining K⁺/Na⁺ homeostasis is essential for stomatal function, photosynthesis, and protein synthesis. V. berlandieri × V. rupestris crosses generally show good Na⁺ exclusion, while the genetic basis of Na⁺ exclusion involves SOS pathway genes and K⁺ transporter selectivity that are partially independent from Cl⁻ exclusion mechanisms.
Natrium-ausschließende Unterlage
Na⁺ excluding rootstock
Unterlage mit geringer Natriumaufnahme
low sodium uptake rootstock
natriumexkludierende Unterlage
sodium excluding rootstock
sodium exclusion rootstock
Downton WJS (1977) Influence of rootstocks on the accumulation of chloride, sodium and potassium in grapevines, Australian Journal of Agricultural Research 28: 879-889 Walker RR et al. (2004) Rootstock and salinity effects on rates of berry maturation, Australian Journal of Grape and Wine Research 10: 227-234 Fisarakis I et al. (2001) The effect of salinity on growth and physiology of four grapevine rootstocks, Scientia Horticulturae 89: 123-134
Sodium Excluding Rootstock Role
Comprehensive characterization of Sodium_Exclusion_Disposition at CSIRO Adelaide profiled sodium concentrations across the root-shoot pathway in 15 rootstocks under 50 mM NaCl salinity, with parallel assessment of potassium status to evaluate Na⁺/K⁺ homeostasis. Tissue sodium and potassium were measured in roots, stems, petioles, and leaf laminae, and the selectivity coefficient SK,Na (K/Na ratio in plant divided by K/Na ratio in soil solution) was calculated as an integrated exclusion metric. Sodium_Exclusion_Disposition showed substantial rootstock variation. '140 Ruggeri' exhibited strong sodium exclusion with leaf sodium of 12 mmol/kg DW, K⁺/Na⁺ ratio of 28, and SK,Na of 85. '1103 Paulsen' showed moderate Sodium_Exclusion_Disposition with leaf sodium of 45 mmol/kg DW, K⁺/Na⁺ ratio of 8, and SK,Na of 24—notably, this rootstock excludes chloride effectively but is less effective for sodium. 'Schwarzmann' displayed poor sodium exclusion with leaf sodium of 95 mmol/kg DW, K⁺/Na⁺ ratio of 3, and SK,Na of 9. Xylem sap analysis revealed that sodium exclusion occurred at both root uptake and xylem retrieval sites, with '140 Ruggeri' showing low xylem sodium (2.8 mM) at root level and efficient retrieval reducing sodium 40% between basal and distal stem positions. Correlation analysis showed that Sodium_Exclusion_Disposition contributed significantly to salt tolerance (r = 0.65 with growth maintenance) but less than chloride exclusion (r = 0.89), confirming the relative importance of these mechanisms.
A Sodium_Exclusion_Disposition is an Ion_Exclusion_Disposition that inheres in a Vitis plant and is realized through physiological mechanisms that prevent or limit the uptake of sodium ions from saline soil and their translocation to shoot tissues, maintaining leaf sodium concentrations below toxic thresholds. This disposition represents the sodium-specific component of salt tolerance, important though generally secondary to chloride exclusion in grapevine. Sodium toxicity in grapevine manifests as disruption of potassium-dependent enzyme function (sodium competes with potassium at binding sites), interference with protein synthesis, and metabolic dysfunction, with symptoms including marginal leaf necrosis, reduced photosynthesis, and growth inhibition typically appearing when leaf sodium exceeds 25-50 mmol/kg DW. Sodium exclusion involves restriction of sodium uptake at the root level and prevention of sodium loading into the xylem, mechanistically paralleling chloride exclusion but involving distinct transporter systems. Additionally, sodium that enters the xylem can be retrieved along the transpiration pathway by xylem parenchyma cells, providing a third exclusion barrier. The molecular machinery for sodium transport and exclusion includes HKT (High-affinity Potassium Transporter) family members that mediate both root sodium uptake and xylem retrieval, with HKT1 subfamily members particularly important for retrieval. Sodium_Exclusion_Disposition varies among rootstocks, though typically with less dramatic variation than chloride exclusion.
Sodium_Exclusion_Disposition is_a Ion_Exclusion_Disposition. Sodium exclusion is important but generally secondary to chloride exclusion for grapevine salt tolerance. Child classes include Root_Sodium_Exclusion_Disposition (uptake restriction) and Sodium_Retrieval_Disposition (xylem retrieval). K⁺/Na⁺ ratio and selectivity coefficient (SK,Na) are key metrics. HKT transporters are central to sodium transport and exclusion. Some rootstocks show differential chloride versus sodium exclusion capacity, relevant for waters with varying NaCl versus other chloride salt composition.
Natrium-Restriktion
Natriumausschluss-Disposition
Na⁺ exclusion
Na⁺-Ausschluss
sodium avoidance
sodium restriction
Munns R, Tester M (2008) Mechanisms of salinity tolerance, Annual Review of Plant Biology 59:651-681 Storey R et al. (2003) Transport of sodium from roots to shoots in grapevine rootstocks, Plant, Cell and Environment 26:1943-1956 Walker RR et al. (2004) Rootstock effects on salt tolerance of irrigated grapevines in the Sunraysia region, Australian Journal of Grape and Wine Research 10:34-45
Sodium Exclusion Disposition
Direct assessment of Sodium_Retrieval_Disposition at the Australian Centre for Plant Functional Genomics tracked sodium concentration changes along the transpiration pathway in 10 rootstocks under 50 mM NaCl salinity. Xylem sap was collected by Scholander pressure chamber from root (basal stem), mid-stem, and upper stem positions, with sodium measured at each location. Additionally, VvHKT1 1 gene expression was profiled in xylem-associated tissues. Sodium_Retrieval_Disposition showed significant rootstock variation. '140 Ruggeri' demonstrated strong retrieval with xylem sap sodium declining from 3.8 mM at root level to 2.4 mM at mid-stem to 1.5 mM at upper stem, representing 60% reduction along the pathway. 'K51-40' showed similar retrieval (58% reduction). '1103 Paulsen' exhibited moderate Sodium_Retrieval_Disposition (35% reduction from 8.2 to 5.3 mM). 'Schwarzmann' showed minimal retrieval (12% reduction from 42 to 37 mM), with high initial loading overwhelming limited retrieval capacity. VvHKT1 1 expression analysis revealed that retrieval-effective '140 Ruggeri' showed 5-fold higher HKT1 1 transcript levels in stem xylem parenchyma compared to 'Schwarzmann'. Immunolocalization confirmed HKT1 1 protein in xylem parenchyma cells surrounding vessels in high-retrieval genotypes. These findings establish that Sodium_Retrieval_Disposition provides a significant additional exclusion barrier mediated by HKT1 transporters, with genotypic variation in HKT1 1 expression underlying differential retrieval capacity.
A Sodium_Retrieval_Disposition is a Sodium_Exclusion_Disposition that inheres in Vitis xylem parenchyma cells along the root-to-shoot pathway and is realized through active removal of sodium ions from the xylem sap, reducing sodium translocation to leaves by reclaiming sodium into surrounding parenchyma cells before it reaches sensitive photosynthetic tissues. This disposition represents a unique exclusion mechanism operating after xylem loading, providing a third barrier that can reduce shoot sodium accumulation even when root uptake and xylem loading restriction are imperfect. Sodium retrieval is mediated primarily by HKT1 (High-affinity K⁺ Transporter subfamily 1) proteins expressed in xylem parenchyma cells of roots and shoots, which function as sodium uniporters removing Na⁺ from the xylem stream. Retrieved sodium can be stored in xylem parenchyma vacuoles, redistributed to root tissues for efflux or sequestration, or in some cases loaded into phloem for recirculation. Effective Sodium_Retrieval_Disposition results in progressive decline in xylem sap sodium concentration from root to shoot, with sodium content in upper stems and leaves substantially lower than would be predicted from root xylem loading alone. The retrieval mechanism is particularly important in species like grapevine where complete root-level exclusion is not achieved, providing an additional opportunity to protect the most sensitive upper canopy tissues.
Sodium_Retrieval_Disposition is_a Sodium_Exclusion_Disposition. Sodium retrieval operates after xylem loading, removing sodium from transpiration stream via HKT1 transporters in xylem parenchyma. Retrieval can substantially reduce shoot sodium even when root exclusion is incomplete. Effectiveness is quantified by % reduction in xylem sodium along root-to-shoot pathway. VvHKT1 1 expression correlates with retrieval capacity. Retrieved sodium may be stored in parenchyma vacuoles or recirculated via phloem. Retrieval provides "last line of defense" protecting upper canopy.
HKT1-mediated retrieval
HKT1-vermittelte-Rückholung
Natrium-Rückholungs-Disposition
Xylem-Natrium-Entladung
sodium reclamation
xylem sodium unloading
Munns R et al. (2012) Wheat grain yield on saline soils is improved by an ancestral Na⁺ transporter gene, Nature Biotechnology 30:360-364 Storey R et al. (2003) Transport of sodium from roots to shoots in grapevine rootstocks, Plant, Cell and Environment 26:1943-1956 Davenport RJ et al. (2007) The Na⁺ transporter AtHKT1 1 controls retrieval of Na⁺ from the xylem in Arabidopsis, Plant Cell and Environment 30:497-507
Sodium Retrieval Disposition
Sodium toxicity quality assessment in Murray-Darling Basin Chardonnay identified Na accumulation as a contributing factor to vine decline in a vineyard affected by rising saline groundwater. Visual sodium toxicity quality evaluation revealed characteristic symptoms including bronzing discoloration of older leaves progressing to marginal and interveinal necrosis, reduced shoot vigor with shortened internodes and smaller leaves, and premature leaf drop beginning from the vine base creating sparse lower canopy by mid-season. The sodium toxicity quality pattern showed progressive symptom development through the season as Na accumulated, with early-season symptoms limited to slight bronzing (score 1.5) intensifying to moderate marginal burn (score 3.0) by véraison and severe necrosis with defoliation (score 4.5) by harvest on older leaves. Tissue analysis confirmed sodium toxicity quality with blade Na at 0.45% in symptomatic leaves (toxic, threshold 0.25%), petiole Na at 0.38%, and notably elevated root Na at 0.85% indicating substantial Na uptake from the saline soil solution. Rootstock comparison revealed sodium toxicity quality variation with own-rooted vines showing blade Na of 0.52% and severe symptoms, while adjacent vines on 1103P rootstock showed blade Na of only 0.18% and minimal symptoms, demonstrating the 65% reduction in scion Na accumulation achieved through rootstock sodium exclusion. Soil and water analysis established the source of sodium toxicity quality with irrigation water EC at 2.8 dS/m and SAR (sodium adsorption ratio) of 8.5 indicating sodic conditions, combined with shallow water table at 1.2 m depth contributing additional salt load through capillary rise. Remediation planning addressed sodium toxicity quality through installation of subsurface drainage to lower the water table, scheduled leaching irrigations to flush accumulated sodium below the root zone, and planned replanting with salt-excluding rootstocks for long-term sustainability.
Sodium toxicity quality is a nutrient toxicity symptom quality that inheres in a grapevine organism and represents the damage resulting from excessive sodium accumulation affecting both direct ion toxicity and osmotic stress components, with symptoms including marginal leaf burn progressing from older leaves, general leaf bronzing and chlorosis, reduced growth vigor, and premature leaf abscission, often occurring in conjunction with chloride toxicity where saline irrigation water delivers both ions simultaneously. This quality captures the grapevine response to sodium excess, recognizing that while direct sodium toxicity symptoms may be less specific than boron or chloride damage, sodium accumulation contributes significantly to overall salt stress in vineyards irrigated with saline water or affected by rising water tables bringing salts into the root zone. The ontological significance of sodium toxicity quality lies in its indication of salinity stress requiring management through improved drainage, leaching irrigation, or rootstock selection for sodium exclusion, and in its interaction with soil structure where high sodium causes clay dispersion and soil degradation that compounds the direct plant toxicity effects. The biological basis of sodium toxicity symptoms encompasses the osmotic stress component where high external sodium reduces water availability to roots, the ion-specific toxicity where cellular sodium accumulation disrupts potassium-dependent enzyme function and membrane integrity, and the interference with calcium and potassium uptake that creates secondary nutrient imbalances. The accumulation pattern of sodium shows transport to leaves through the transpiration stream with some retention in woody tissues, resulting in progressive buildup in older leaves and eventual toxicity when tissue concentrations exceed tolerance thresholds. The rootstock contribution to sodium management is substantial, with major differences in sodium exclusion capacity among rootstocks enabling significant reduction in scion sodium accumulation when appropriate rootstocks are selected for saline conditions.
Sodium_Toxicity_Quality is_a Nutrient_Toxicity_Symptom_Quality, representing the damage from Na accumulation causing both direct ion toxicity and osmotic stress. This class addresses the specific Na toxicity syndrome often occurring with Cl toxicity. Blade Na threshold for toxicity ~0.25% (varies by cultivar). Rootstock Na exclusion capacity varies dramatically (1103P, Ramsey, 140Ru > own-rooted, SO4). Osmotic + ion-specific + K/Ca interference all contribute to damage. Soil sodicity (high SAR) compounds direct plant effects through soil structure degradation. Often co-occurs with Cl toxicity from saline irrigation.
Na toxicity
Na-Überschuss-Symptome
Natriumtoxizität
Sodizitätsschaden
sodicity damage
sodium excess symptoms
Walker RR et al. (2010) Rootstock effects on salt tolerance of irrigated field-grown grapevines. Australian Journal of Grape and Wine Research 16:174-180 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Prior LD et al. (1992) Sodium chloride and soil texture interactions in irrigated field grown Sultana grapevines. Australian Journal of Agricultural Research 43:1051-1066
Sodium Toxicity Quality
A soft green one-year cane on Vitis vinifera cv. Merlot examined in mid-November following excessively vigorous growing season with late nitrogen fertilization and continued irrigation through September, showing bright green color throughout 195cm length, extremely soft texture easily compressed between fingers leaving temporary indentations, readily bending into tight curves without resistance, cross-section revealing 85-90% water content by mass, large thin-walled parenchyma cells throughout cortex and pith, minimal vascular development with xylem vessels small and sparse, representing complete maturation failure requiring total removal during pruning with no portions suitable for retention, triggering vineyard management review to prevent recurrence through modified fertilization and irrigation protocols.
A green one-year cane characterized by pronounced softness and lack of structural rigidity, readily deforming under minimal pressure, containing high water content in turgid parenchyma cells, exhibiting minimal secondary cell wall development with primary tissues dominant, and representing either very young shoot tissue (early growing season) with expected maturation ahead or severely immature late-season wood (autumn) indicating extreme ripening failure threatening complete unsuitability for retention as fruiting wood. Soft green one-year canes demonstrate the herbaceous nature of unlignified grapevine tissues, with mechanical properties resembling leafy vegetative organs rather than woody perennial structures, making such tissues completely unsuitable for supporting crop loads, surviving winter conditions, or providing vascular connections for productive fruiting shoots if this soft status persists into dormant pruning season.
Soft green one-year canes represent the extreme of incomplete wood maturation, essentially remaining in shoot-like developmental status despite calendar progression into dormant season. Such canes have zero cold hardiness (damage occurring at -1°C to -3°C versus -15°C to -25°C for properly lignified wood), cannot support mechanical loads, lack adequate vascular development for efficient water and nutrient transport, and produce poorly developed buds with minimal fruitfulness potential. The soft texture reflects maintained primary tissue structure with large vacuolated parenchyma cells, thin primary cell walls, and absence of secondary xylem and periderm formation. Persistence of soft green canes into late autumn indicates fundamental vine balance problems requiring systemic correction, not just localized cane removal, as underlying causes will affect entire vineyard sections.
non-rigid herbaceous cane
sehr unausgereifte Rute
succulent green cane
very soft unlignified cane
weiche grüne Rute
Soft Green One Year Cane
Merlot soft inner mesocarp at maturity showing soft texture characteristics= cells 55-82 μm diameter (smaller than middle mesocarp 125-165 μm but still moderately large), walls 0.9-1.2 μm (thicker than middle mesocarp 0.5-0.7 μm but still relatively thin), water content 79-82%, sugars 23-25% creating high osmotic potential and turgor, firmness 1.8-2.8 N (soft, comparable to middle mesocarp 1.2-2 N despite smaller cells and thicker walls), demonstrating that high water content and sugar-driven turgor create soft texture even with somewhat thicker walls and smaller cells than middle mesocarp. Thompson Seedless table grape soft inner mesocarp contributing to eating quality= soft inner mesocarp 4-5 cell layers, cells 52-75 μm, water 80-83%, sugars 19-22%, firmness 2.2-3.2 N, creating soft juicy texture throughout berry flesh including seed trace regions (stenospermocarpic development creating small seed traces rather than normal seeds), with soft inner mesocarp ensuring pleasant texture even when eating includes biting into seed trace regions, important for seedless table grape eating quality. Cabernet Sauvignon soft inner mesocarp ripening softening= veraison (BBCH 81) showing cells 48-65 μm, walls 1.3-1.6 μm, firmness 3.8-5.2 N (moderately firm) maturity (BBCH 89) showing cells 58-85 μm, walls 0.9-1.3 μm (thinning through expansion despite continued deposition), firmness 2-3 N (soft), demonstrating ripening-associated softening through cell expansion, wall thinning, sugar accumulation, and wall modifications creating soft inner mesocarp texture at maturity from initially firmer state at veraison. Chardonnay soft inner mesocarp in white cultivar= cells 58-88 μm, walls 0.9-1.2 μm, water 78-81%, sugars 21-24%, firmness 2.2-3.5 N, creating soft texture similar to red cultivar inner mesocarp, demonstrating soft inner mesocarp as textural characteristic present across cultivar types independent of anthocyanin presence/absence. Pinot Noir soft inner mesocarp versus firm skin textural contrast= soft inner mesocarp firmness 2.2-3.8 N yielding readily to compression firm skin (outer pericarp plus outer mesocarp) firmness 8-15 N requiring substantial force to compress, creating obvious textural difference between soft juicy flesh (middle plus inner mesocarp, 85-92% of mesocarp volume) and firm chewy skin (8-15% of pericarp volume), with textural contrast important for both eating quality (separating soft flesh from tough skin during chewing) and winemaking (firm skin requiring vigorous maceration for phenolic extraction, soft flesh extracting readily).
A soft inner mesocarp is an inner mesocarp emphasizing soft texture similar to middle mesocarp despite smaller cell size and thicker walls than middle mesocarp, characterized by high water content (77-83%), high sugar content (19-27%) creating high vacuolar osmotic potential, thin-to-moderate walls (0.8-1.2 μm) despite being thicker than middle mesocarp walls (0.4-0.8 μm), and turgid cells creating soft fleshy texture comparable to middle mesocarp texture, representing inner mesocarp from textural rather than phenolic perspective. Soft inner mesocarp textural characteristics= (1) soft texture when cells turgid, yielding readily to compression similar to middle mesocarp despite smaller cell size (2) high water content maintaining cell turgor creating soft turgid state (3) juicy texture when cells disrupted releasing cellular contents rich in water and sugars (4) potential textural gradient from softer outermost inner mesocarp (more similar to middle mesocarp) to slightly firmer innermost inner mesocarp (approaching skin-like firmness) though overall maintaining soft character. Soft inner mesocarp cellular basis of texture= (1) high vacuolar volume (88-93% of cell volume) occupied by aqueous solution despite being less than middle mesocarp (90-95%) creating substantial water reservoir (2) thin-to-moderate walls relative to other inner pericarp tissues allowing yielding under compression (3) high turgor pressure from osmotic potential of vacuolar sugars creating internal pressure supporting soft turgid state (4) wall composition potentially showing high pectin methylesterification creating flexible rather than rigid walls. Soft inner mesocarp in berry eating quality= (1) contribution to overall berry softness and juiciness creating pleasant eating texture (2) cells readily disrupting during chewing releasing sweet juice (3) minimal contribution to skin toughness or chewiness despite being inner mesocarp layer (4) particular importance for table grapes where soft juicy texture throughout flesh including inner mesocarp desired for eating quality. Soft inner mesocarp developmental changes= early development showing firmer texture with smaller cells (30-50 μm) and thicker walls (1.2-1.6 μm) ripening showing progressive softening through cell expansion (cells enlarging to 50-85 μm), wall thinning (walls declining to 0.8-1.2 μm through expansion), sugar accumulation increasing turgor, and wall modifications (pectin depolymerization) creating softer texture over-ripening potentially showing excessive softening creating mealy or mushy texture through extensive wall breakdown. Soft inner mesocarp in wine production= soft texture allowing relatively easier cell disruption during maceration compared to firm outer pericarp tissues cells breaking down during maceration releasing contents (sugars, acids, some tannins 3-6 mg/g) into must soft inner mesocarp extracting more readily than firm outer mesocarp during maceration though still requiring more vigorous extraction than soft middle mesocarp due to smaller cell size. Soft inner mesocarp versus firm skin distinction= soft inner mesocarp maintaining soft texture similar to middle mesocarp (both yielding readily to compression) versus firm skin (outer pericarp, outer mesocarp) requiring substantial force to compress or penetrate, creating textural contrast between soft flesh (middle mesocarp plus soft inner mesocarp representing 80-90% of mesocarp volume) and firm skin (outer pericarp plus firm outer mesocarp representing 5-12% of pericarp volume).
Soft_Inner_Mesocarp emphasizes the soft juicy textural properties of inner mesocarp and is distinguished from Juicy_Inner_Mesocarp which may emphasize juice release over textural firmness. Soft inner mesocarp represents the inner mesocarp from the perspective of eating texture and maceration behavior. Understanding soft inner mesocarp is relevant for table grape quality assessment (desirable soft texture throughout flesh), for berry texture characterization, and for winemaking where soft inner mesocarp extracts more readily than firm outer tissues despite being inner mesocarp position requiring relatively deeper extraction.
Weiches samennahes Gewebe
soft inner flesh
tender seed-adjacent tissue
weiches inneres Fruchtfleisch
weiches inneres Mesokarp
Thomas, T.R. et al. (2008). Changes in physical properties of grape berries. American Journal of Enology and Viticulture 59=398-409 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 180-185.
Soft Inner Mesocarp
Merlot soft pulp tissue at optimal maturity showing characteristic soft texture= firmness 1.8-2.4 N measured by 8 mm probe penetrometry (soft, yielding readily to compression), cells 130-160 μm diameter with thin 0.5-0.7 μm walls, water content 78-81%, vacuoles 92-94% of cell volume, creating soft pleasant texture for eating and facilitating juice extraction during winemaking with 74-76 L juice yield per 100 kg berries from readily-disrupting soft cells. Thompson Seedless table grape soft pulp tissue for fresh consumption= firmness 2.2-2.8 N (moderately soft balancing eating quality with shipping durability), cells 160-190 μm diameter, water 80-83%, creating tender eating texture requiring minimal chewing effort while maintaining sufficient firmness for 3-4 week cold storage and shipping (0-2°C), with soft pulp allowing pleasant eating experience while firmer skin provides protection during handling. Cabernet Sauvignon soft pulp tissue ripening softening progression= veraison (BBCH 81, week 10) firmness 3.8-4.8 N (firm), cells 75-95 μm, walls 0.7-0.9 μm mid-ripening (week 14) firmness 2.5-3.5 N (moderately soft), cells 115-145 μm, walls 0.55-0.75 μm maturity (week 18) firmness 1.8-2.5 N (soft), cells 145-165 μm, walls 0.5-0.7 μm, demonstrating 53-62% firmness reduction through cell expansion, wall thinning, and wall modifications during 8-week ripening period creating characteristic soft pulp texture at harvest maturity. Chardonnay soft pulp tissue facilitating white wine juice extraction= soft texture (firmness 1.6-2.2 N) allowing efficient pressing yielding 74-78 L juice per 100 kg berries with 65-72% free-run juice (released by crushing before pressing) from readily-disrupting soft cells, 28-35% press juice (requiring 0.5-2 bar pressure), with soft pulp contributing to high juice yield important for white wine production economics. Pinot Noir soft pulp tissue over-ripening softening= optimal maturity (week 17) firmness 1.9-2.6 N (appropriately soft), cells intact, pleasant texture delayed harvest (week 19) firmness 1.2-1.8 N (excessively soft), beginning mealy texture from excessive cell separation, cells showing wall degradation severe over-ripening (week 21) firmness 0.8-1.3 N (mushy), loss of structural integrity, demonstrating progression from optimal soft pulp to defective over-soft pulp when harvest delayed excessively beyond optimal maturity creating undesirable textural degradation.
A soft pulp tissue is pulp tissue emphasizing characteristic soft yielding texture when turgid, typically showing firmness 1-2.5 N measured by penetrometry or compression testing, resulting from large thin-walled cells with high water content creating low resistance to compression, representing the textural characteristic that defines flesh quality for both eating grapes and processing grapes where soft texture allows easy juice extraction. Soft pulp tissue textural basis= (1) large cells 110-180 μm diameter creating low cell wall surface area per unit volume reducing structural density (2) thin walls 0.4-0.8 μm providing minimal resistance to compression with walls readily deforming under low forces (3) high water content 76-82% creating turgid cells that yield readily when compressed as water redistributes within and between cells (4) high vacuolar volume 90-95% filled with liquid creating cells that are predominantly water-filled sacs with minimal solid structure (5) low intercellular adhesion strength allowing cells to separate readily during compression creating soft yielding tissue. Soft pulp tissue developmental changes= early development (Phase I, BBCH 71-75) showing firmer texture with smaller cells (40-70 μm), thicker walls (0.8-1.2 μm), lower water content (68-75%) creating firmness 3.5-5 N ripening (Phase II, BBCH 81-89) showing progressive softening through cell expansion (cells enlarging to 110-180 μm), wall thinning (walls declining to 0.4-0.8 μm through expansion despite continued deposition), water accumulation (increasing to 76-82%), and wall modifications (pectin depolymerization, hemicellulose degradation) creating firmness decline to 1-2.5 N over-ripening potentially showing excessive softening (firmness <1 N) creating mealy or mushy defects through extensive wall breakdown. Soft pulp tissue in eating quality= (1) pleasant soft yielding texture when biting berry creating satisfying mouthfeel (2) cells readily disrupting during chewing releasing juice without excessive resistance (3) softness allowing comfortable eating without excessive jaw work (4) optimal softness balance for table grapes requiring soft enough for pleasant eating but firm enough for shipping durability (target firmness 1.8-2.8 N for shipping cultivars). Soft pulp tissue in juice and wine production= (1) soft texture facilitating juice extraction during crushing and pressing with cells readily rupturing under moderate pressures (2) high juice yield 72-78 L per 100 kg berries reflecting high water content and ready cell disruption (3) free-run juice (released by crushing without pressing) representing 60-70% of total juice from soft readily-disrupting cells (4) press juice (requiring pressure) representing 30-40% of total from more resistant cells and skin tissues. Soft pulp tissue cultivar and environmental variation= cultivar differences with some table cultivars showing particularly soft pulp (firmness 1.2-2 N) creating very tender eating texture, wine cultivars showing moderate firmness (1.5-2.5 N), some shipping cultivars showing firmer pulp (2-3 N) for durability environmental effects with water stress during ripening potentially maintaining firmer pulp through reduced cell expansion and lower water accumulation, heat stress accelerating softening through enhanced wall degradation cultural practices affecting softness including irrigation management (deficit maintaining firmness, excess promoting softness), crop load (high load delaying ripening and softening), and calcium nutrition (enhancing cell wall strength reducing softening rate).
Soft_Pulp_Tissue emphasizes the textural softness characteristic of pulp and is distinguished from Liquid_Pulp_Tissue (emphasizing juice content) and Storage_Pulp_Tissue (emphasizing storage function). Understanding soft pulp tissue is essential for harvest timing decisions (achieving appropriate softness without over-ripening), for table grape quality assessment (balancing eating softness with shipping firmness), and for processing efficiency (soft pulp extracting more readily during crushing and pressing).
Weiches Fruchtfleisch
soft fruit tissue
tender flesh
weiches Fruchtmark
weiches Pulpa-Gewebe
Thomas, T.R. et al. (2008). Changes in the physical properties of grape berries during development and ripening. American Journal of Enology and Viticulture 59=398-409 Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 185-190.
Soft Pulp Tissue
Chardonnay in Champagne shows softening initiation during early-mid September Year N+1 at BBCH 81-83, with berry firmness declining from 2.8 N pre-veraison to 2.2 N representing 21% softening over 2-3 days, polygalacturonase activity increasing from 8 to 28 units, pectin methylesterase increasing from 12 to 35 units, water-soluble pectin increasing from 22% to 38% total pectin, texture perceptibly softer, under cool maritime conditions of 18-24°C temperature, typical of this early-bursting white cultivar showing softening as primary veraison marker
Softening initiation represents the first texture transformation phase during lag phase termination when berry firmness begins measurable decline, cell wall modification initiates through pectin degradation, texture changes from hard to yielding, firmness declining 10-25% from pre-veraison maximum during termination period, and softening marks universal veraison indicator across red and white cultivars, occurring during final 1-3 days of lag as transformation begins. This texture transformation onset phase occurs during late summer to early autumn in early-mid September in temperate regions Year N+1, proceeding over 1-3 days as softening begins, corresponding to BBCH 81-83 veraison onset period. Softening mechanism involves cell wall modification initiation with pectin-degrading enzyme activation, pectin depolymerization reducing cell wall integrity, cell-cell adhesion weakening from middle lamella degradation, turgor possibly declining contributing to softness, and integrated changes creating firmness decline. Firmness decline characteristics shows berry firmness beginning measurable reduction, typical decline 10-25% from pre-veraison maximum 2.5-4.5 N to early softening 2.0-3.5 N during termination, progressive firmness decrease continuing through ripening, and firmness decline as defining transformation. Firmness measurement shows penetrometer resistance declining, compression force requirement decreasing, subjective texture softer to touch, and firmness as quantitative softening indicator. Firmness decline timing shows earliest firmness decline in first transforming berries at veraison onset, progressive softening in subsequently transforming berries, and timing showing asynchronous pattern across cluster. Cell wall modification initiation shows pectin-degrading enzymes activating at softening onset, polygalacturonase PG activity increasing substantially, pectin methylesterase PME activity increasing, pectate lyase possibly activating, and enzyme activation as softening mechanism. Polygalacturonase activation shows PG enzyme activity increasing from pre-veraison low levels 5-15 units to early softening 15-40 units, PG degrading homogalacturonan backbone of pectin, enzyme activity correlating with firmness decline, and PG as major softening enzyme. Pectin methylesterase activation shows PME activity increasing demethylating pectin, PME creating substrate for PG degradation, PME possibly biphasic with initial cross-linking then degradation facilitation, and PME as pectin modification enzyme. Pectate lyase possible role shows enzyme possibly contributing to pectin degradation, lyase cleaving pectin chains, contribution varying by cultivar and conditions, and lyase as additional softening enzyme. Pectin depolymerization shows pectin molecular weight declining from enzyme degradation, high molecular weight pectin 100-300 kDa declining to lower molecular weight fragments 20-100 kDa, depolymerization reducing cell wall structural integrity, and pectin degradation as softening basis. Pectin solubilization shows water-soluble pectin fraction increasing from degradation products, insoluble cell wall pectin fraction declining, solubilization weakening wall structure, and soluble pectin as degradation indicator. Middle lamella degradation shows pectin-rich middle lamella between cells degrading, cell-cell adhesion weakening from middle lamella loss, intercellular spaces possibly increasing, and middle lamella as adhesion loss site. Cell-cell adhesion weakening shows cells becoming less firmly attached, tissue disaggregation potential increasing, texture becoming more yielding from reduced cohesion, and adhesion loss contributing to softness. Calcium dynamics shows calcium possibly released from demethylated pectin as degradation proceeds, calcium cross-links breaking, calcium redistribution affecting cell wall properties, and calcium as structural ion in cell walls. Cell wall other modifications shows possible hemicellulose and cellulose modifications though minor compared to pectin, expansin possibly contributing to wall loosening, overall wall structure weakening, and integrated modifications creating softening. Turgor possible changes shows turgor pressure possibly declining slightly during early softening from altered water relations, reduced turgor contributing to softer texture, though turgor changes likely minor compared to structural changes, and turgor as potential softening contributor. Berry texture changes shows texture transitioning from hard resistant to yielding, berry easier to compress, skin possibly loosening from flesh, and overall texture becoming characteristic of ripening berry. Berry appearance shows softening berries often also showing color change in red cultivars, white cultivars showing softening as primary visible change possibly with yellowing, and texture change as transformation indicator. Sensory evaluation shows tactile assessment detecting softness, berries beginning to yield to gentle pressure, texture change perceptible to trained evaluators, and sensory evaluation as practical assessment. Softening as veraison marker shows softening occurring in all cultivars at veraison, universal transformation marker unlike color which is cultivar-dependent, softening defining veraison in white cultivars, and texture as reliable indicator. Softening progression shows continued firmness decline through ripening, eventual soft texture at full ripeness, progression rate varying by cultivar and conditions, and softening as continuing ripening process. Environmental effects on softening initiation include temperature affecting enzyme activities and softening rate, warm conditions promoting faster softening, cool conditions slowing softening progression, water availability affecting turgor and texture, and environmental factors influencing softening dynamics. Temperature effects shows warm conditions accelerating enzyme activities and softening rate, cool conditions slowing softening, and temperature as softening rate determinant. The process represents second aspect of lag phase termination emphasizing texture transformation, following veraison approach and preceding lag phase exit. Duration 1-3 days representing softening onset. Regional timing shows softening initiation during early-mid September at veraison onset. Viticultural significance includes representing universal veraison marker across cultivars, indicating transformation onset determining harvest timing, affecting berry handling and disease susceptibility, and establishing texture trajectory affecting wine quality. Research applications include analyzing cell wall modification mechanisms and enzyme regulation, investigating pectin degradation and depolymerization dynamics, evaluating environmental effects on softening rate, quantifying firmness decline and texture changes, and assessing softening initiation as veraison marker and quality determinant.
Softening_Initiation is second aspect of Lag_Phase_Termination, distinguished by first texture transformation. Represents 10-25% firmness decline from 2.5-4.5 N peak. Polygalacturonase and pectin methylesterase activation. Pectin depolymerization 100-300 → 20-100 kDa. Middle lamella degradation, cell-cell adhesion weakening. Universal veraison marker (all cultivars). Duration 1-3 days. Follows Veraison_Approach. Precedes Lag_Phase_Exit. Defines veraison in white cultivars.
Erweichungs-Initiierung
Festigkeits-Abnahme-Start
Textur-Transformations-Beginn
cell wall modification initiation
firmness decline start
texture transformation onset
Softening Initiation
In a Barossa Valley Shiraz vineyard in Australia, softening process completion was documented 14 days post-veraison onset in late February Year N+1 (BBCH 87) when firmness stabilized at 1.5-1.6 N (±0.1 N over 10 days), polygalacturonase activity plateaued at 52-58 units (down from 168 units peak), soluble pectin reached 72-74% with <2% increase over 2 weeks, sugar content was 24.8° Brix, titratable acidity 5.5 g/L, seed tannins brown, and sensory evaluation confirmed balanced maturity under warm dry conditions (daily maximum 28-32°C, minimal precipitation) suitable for harvest.
Softening_Process_Completion represents the developmental endpoint when all berry softening mechanisms have reached asymptotic termination, enzymatic activities stabilize at maintenance levels, and further texture changes reflect senescence or environmental degradation rather than programmed developmental processes. This completion occurs during late veraison through early ripening (BBCH 85-89, typically 10-16 days post-veraison onset) and marks the transition from active ripening-driven softening to a stable mature state that persists until harvest unless environmental perturbations induce secondary texture changes. Enzymatic completion is characterized by polygalacturonase activity declining from peak values (120-220 units) to stable maintenance levels (35-70 units), pectin methylesterase similarly reducing from 110-200 units to 30-65 units, and other cell wall-modifying enzymes including xyloglucan endotransglycosylase (50-120 units declining to 15-40 units), β-galactosidase, pectate lyase, and expansins all showing parallel activity reductions to 25-40% of peak values indicating completion of major degradative processes while maintaining sufficient activity to preserve cell wall plasticity without driving further extensive breakdown. Gene expression patterns at softening process completion transition from ripening peak to post-ripening maintenance: cell wall enzyme genes decline from maximum transcription levels (80-200 fold above green berry baseline) to moderate sustained expression (15-50 fold) providing maintenance enzyme synthesis, transcription factors orchestrating ripening including NAC, MYB, and MADS-box genes show reduced expression as developmental program completion signals propagate, and genes involved in senescence and stress responses begin elevation indicating shift from active development to mature state maintenance or early senescence depending on environmental conditions and cultivar physiology. Cell wall structural completion shows pectin composition stabilized at final values: soluble pectin fraction plateaus at 70-85% of total (compared to 20-30% green berries with increases 30-50% weekly changes during active softening phases. Completion timing varies by cultivar and environment: early-ripening varieties (Pinot Noir, Chardonnay, Gamay) achieve completion 10-14 days post-veraison under optimal conditions (22-28°C average temperature), mid-season cultivars (Merlot, Syrah, Cabernet Franc) complete 12-16 days post-veraison, while late-season varieties (Cabernet Sauvignon, Nebbiolo, Mourvèdre) may require 14-20 days for full completion particularly under cooler conditions (32°C) can cause premature abnormal completion through enzyme denaturation and cellular stress potentially producing suboptimal final texture, and cool conditions (<16°C) substantially delay completion (5-10 days extension) risking incomplete softening if harvest necessitated by other maturity parameters or approaching unfavorable weather. Post-completion berry texture stability depends on cultivar characteristics and environmental conditions: stable cultivars (Cabernet Sauvignon, Syrah, Sangiovese) maintain consistent texture 3-5 weeks post-completion allowing extended hang time for flavor development and harvest scheduling flexibility, moderately stable varieties (Merlot, Grenache, Chardonnay) show gradual texture changes 2-3 weeks post-completion with slow over-softening or slight firmness increase from mild dehydration requiring reasonably prompt harvest, while unstable cultivars (Pinot Noir, Gewürztraminer, Riesling) may deteriorate rapidly 1-2 weeks post-completion through over-softening, senescence, or environmental damage necessitating precise harvest timing. Quality implications of softening process completion timing relative to other maturity parameters inclu optimal scenarios where completion coincides with target sugar levels (20-26° Brix style-dependent), desired acidity (4-8 g/L titratable acidity), and phenolic maturity (brown seeds, extractable tannins balanced) produce ideal harvest-ready grapes for premium wine production, discordant maturity with premature completion before chemical maturity requires difficult decisions between harvesting under-ripe for texture or delaying past optimal firmness for composition, while delayed completion beyond chemical maturity creates opposite challenges of harvesting firm berries with suboptimal extraction potential or waiting for softening completion risking over-ripeness, excessive alcohol, or unfavorable weather exposure. Viticultural management targeting optimal softening completion involves: irrigation scheduling during ripening to maintain moderate water availability supporting enzyme synthesis and activity for timely completion without excessive water promoting over-softening, canopy management to moderate bunch temperatures facilitating appropriate enzyme function without heat-accelerated abnormal softening or cool-delayed incomplete softening, crop load adjustment ensuring adequate carbohydrate supply for enzyme synthesis and ripening processes supporting synchronized completion with chemical maturity, and disease management particularly Botrytis prevention since completion marks beginning of maximum berry vulnerability period with soft texture and compromised skin integrity creating optimal infection conditions requiring protective fungicide applications or prompt harvest. Monitoring softening process completion through regular firmness measurements combined with enzyme activity assays or gene expression analysis in research settings provides precise completion determination enabling harvest timing optimization for specific quality goals and market requirements, while practical viticulture relies on tactile berry assessment, chemical maturity tracking, and phenolic maturity evaluation to infer completion and guide harvest decisions balancing texture, composition, and logistical considerations.
Softening_Process_Completion is distinguished from Complete_Berry_Softening by representing the developmental program endpoint rather than texture state achievement, from Mature_Berry_Texture by focusing on process termination rather than integrated sensory quality, and from earlier softening phases by exhibiting asymptotic stabilization of all softening parameters. Completion timing: early cultivars 10-14 days post-veraison, mid-season 12-16 days, late varieties 14-20 days. Enzyme activities stabilize at 25-40% of peak values. Post-completion stability varies 1-5 weeks cultivar-dependent. Softening_Process_Completion determination is essential for harvest timing decisions ensuring texture development synchronization with chemical and phenolic maturity to optimize grape quality for premium wine production.
Entwicklungs-Textur-Vollendung
Erweichungs-Programm-Endpunkt
Reifungs-Erweichungs-Abschluss
developmental texture completion
ripening softening finalization
softening program endpoint
terminal softening achievement
terminale Erweichungs-Erreichung
Keller M (2020) The Science of Grapevines 3rd Ed pages 228-238 berry softening and ripening completion Coombe BG (1992) Research on development and ripening of the grape berry Australian J Grape Wine Res 48:125-144 ripening process completion Deluc LG, Grimplet J, Wheatley MD, Tillett RL, Quilici DR, Osborne C, Schooley DA, Schlauch KA, Cushman JC, Cramer GR (2007) Transcriptomic and metabolite analyses of Cabernet Sauvignon grape berry development Bmc Genomics 8:429 molecular markers of ripening completion.
Softening Process Completion
Cabernet Sauvignon in Napa Valley transitions through softening stages over 45-50 days post-véraison, with texture analyzer measurements showing firmness decline from 8.5 N at pre-véraison to 2.1 N at optimal harvest (24.5°Brix), enabling winemakers to correlate softening stage with phenolic extractability for structured red wine production.
Softening stage quality is a physical quality that inheres in a grape berry and represents the developmental phase-specific mechanical state characterizing the progressive loss of tissue rigidity throughout berry maturation. Within the BFO framework, this quality captures the temporal continuum of firmness changes that define discrete physiological windows from the rigid pre-véraison state through post-véraison softening to harvest-ready texture and eventual overripe degradation. The ontological significance lies in softening as a composite indicator integrating cell wall disassembly, turgor pressure reduction, and membrane integrity changes that collectively determine berry mechanical properties at each developmental stage. Grapevine berry softening follows a characteristic biphasic pattern, with the dramatic softening at véraison representing the most rapid change, typically reducing firmness by 40-60% within 7-14 days as pectin depolymerization and hemicellulose solubilization accelerate under hormonal control. This quality is measured instrumentally through texture analyzers recording force-deformation curves at standardized stages, enabling quantification of compression resistance, puncture force, and elastic modulus across the developmental timeline. The developmental stages encompassed by this quality are not merely descriptive categories but represent distinct physiological states with characteristic cell wall enzyme activities, particularly polygalacturonase, pectinmethylesterase, and expansin expression patterns that drive the softening progression. Cultivar variation in softening stage dynamics significantly impacts harvest timing decisions, with some varieties maintaining firmness longer post-véraison while others soften rapidly, affecting both optimal harvest window duration and susceptibility to mechanical damage during harvest operations.
Softening_Stage_Quality serves as the parent class for developmental phase-specific firmness qualities, distinguishing temporal progression from the mechanical measurement qualities under Firmness_Quality. This class aligns with OIV descriptor 235 (berry firmness) interpreted across developmental stages. Measurement standardization requires specifying days post-véraison, berry temperature, and instrument parameters for cross-study comparability.
Entwicklungsstadium der Beerenfestigkeit
Erweichungsstadium
Erweichungsverlauf
Reifefestigkeitsphase
berry softening phase
developmental firmness stage
maturation softening state
ripening firmness progression
Coombe BG, McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6(2): 131-135 Ortega-Regules A et al. (2006) A first approach towards the relationship between grape skin cell-wall composition and anthocyanin extractability. Analytica Chimica Acta 563(1-2): 26-32
Softening Stage Quality
A 10 cm terminal shoot segment of Chardonnay clone 96 harvested in June from FPS nuclear stock bears a Softwood_Cutting_Role, prepared by removing lower leaves, treating the base with 1000 ppm IBA, and placing under intermittent mist with 25°C bottom heat—rooting within 3 weeks to rapidly multiply virus-free material for foundation block establishment. A softwood cutting of 41B rootstock (difficult-rooting V. vinifera × V. berlandieri hybrid) serves in a Softwood_Cutting_Role, achieving 70-80% rooting success compared to <40% for hardwood cuttings of the same genotype. A green cutting of Gewürztraminer bears a Softwood_Cutting_Role in research propagation, enabling year-round multiplication independent of the dormant season for experimental trials.
A cutting role that inheres in a stem segment harvested from actively growing, non-lignified shoot tissue during the growing season, typically May-July in the Northern Hemisphere when vegetative growth is vigorous and tissue remains succulent and green. Softwood cuttings (also termed green cuttings or herbaceous cuttings in grapevine propagation) exploit the high meristematic activity and cellular plasticity of juvenile tissues, offering advantages including rapid rooting (typically 2-4 weeks versus 6-12 weeks for hardwood), higher rooting percentages for some difficult-rooting genotypes, and maximum multiplication rate from limited source material (enabling propagation throughout the growing season rather than only during dormancy). However, softwood cuttings require controlled environment facilities (mist propagation systems, bottom heat, high humidity maintenance) to prevent desiccation before root establishment, and produce smaller, more delicate plants requiring extended nursery growing before field-readiness. Softwood cutting propagation is particularly valuable for rapid multiplication of virus-tested nuclear stock and for difficult-rooting genotypes where hardwood cutting success is inadequate.
Softwood_Cutting_Role captures the growing-season propagation alternative to dormant hardwood cuttings, with distinct advantages (speed, success rate for difficult genotypes, continuous production) balanced against requirements (mist facilities, climate control, intensive management). The technique is standard for rapid multiplication of nuclear stock but less common in commercial production due to infrastructure costs and the smaller plants produced. Rooting physiology differs from hardwood—softwood cutting success depends on maintenance of turgor and prevention of wilting through the rooting period, while hardwood cuttings rely on stored reserves during the extended period before root function. Auxin treatment (IBA at 500-2000 ppm) is routine for softwood cuttings, improving both rooting percentage and root system quality. Softwood cutting propagation has enabled establishment of clean stock programs by rapidly multiplying virus-eliminated meristem-derived plants.
Grünsteckling
Sommersteckling
Weichholzsteckling
green cutting
herbaceous cutting
krautiger Steckling
softwood stem cutting
summer cutting
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 360-370 Goussard PG (1981) Rooting of vine cuttings, South African Journal of Enology and Viticulture 2: 29-35 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press
Softwood Cutting Role
The soil amendment specification for a newly established Tempranillo vineyard on acidic granite-derived soils in the Dão region prescribes pre-planting incorporation of 3 t/ha agricultural limestone (CaCO3, 90% calcium carbonate equivalent) to a depth of 40 cm by deep ripping and rotary cultivation, raising soil pH from 4.8 to the target range of 5.8-6.2 in the primary rooting zone — the liming rate was calculated from soil buffer capacity analysis (Adams-Evans buffer pH method) and will be supplemented by maintenance applications of 0.5-1.0 t/ha every 3-5 years based on annual soil pH monitoring at 0-30 cm depth. Additionally, the specification includes incorporation of 2 kg/ha mycorrhizal inoculant (Rhizophagus irregularis at 1,000 spores/g) into the planting hole at vine establishment to accelerate arbuscular mycorrhizal colonization of the developing root system, enhancing phosphorus uptake efficiency by an estimated 20-40% in the phosphorus-fixing acid soil conditions.
A soil management description that documents the application of mineral, chemical, or biological materials to vineyard soil to correct specific deficiencies, adjust soil pH, improve physical properties, or enhance biological function, specifying the amendment type, application rate, incorporation method, target soil parameter change, and monitoring protocol that collectively define the corrective intervention applied to optimize the soil environment for grapevine root function.
Soil_Amendment_Specification addresses targeted corrective interventions that modify specific soil properties rather than providing general fertility or organic matter inputs (which are covered by Compost_Application_Rate). The most common vineyard soil amendments are liming materials (limestone, dolomite, gypsum) applied to correct soil acidity — a widespread issue in vineyards on granitic, schistose, and sandy soils where decades of ammonium-based fertilization and organic acid leaching have progressively lowered soil pH below the optimal range of 5.5-7.0 for Vitis vinifera root function: soil pH below 5.0 induces aluminum and manganese toxicity, reduces phosphorus availability, inhibits bacterial nitrogen mineralization, and suppresses mycorrhizal colonization, collectively impairing vine nutrition and root health. Other amendment categories include gypsum (calcium sulfate) for sodic soil remediation, eleite sulfur for pH reduction of alkaline calcareous soils, rock phosphate for long-term phosphorus supply in organic systems, and biological inoculants (mycorrhizal fungi, nitrogen-fixing bacteria, plant growth-promoting rhizobacteria) that enhance root zone biological function.
Bodenhilfsstoff-Protokoll
Bodenkorrektur-Dokumentation
Bodenverbesserungs-Spezifikation
amendment application protocol
soil correction specification
soil improvement record
White RE (2015) Understanding Vineyard Soils 2nd Ed, Oxford University Press pp: 135-165 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 450-460 Trouvelot S et al. (2015) Arbuscular mycorrhiza symbiosis in viticulture: a review, Agronomy for Sustainable Development 35: 1449-1467
Soil Amendment Specification
Garnacha vines in the Priorat of Catalonia bearing a soil building role have over 80+ years of cultivation on llicorella slate soils transformed the initial 5-10 cm effective soil depth into 20-35 cm of biologically active rooting zone through continuous root penetration and decay cycling, with soil organic carbon increasing from 0.5% to 1.8-2.5% in the top 20 cm under vine rows, earthworm populations of 80-150 individuals/m² versus 10-20/m² in adjacent uncultivated slate, and mycorrhizal hyphal networks extending the vine's effective nutrient acquisition zone into previously inaccessible rock matrix at weathering rates enhanced 3-5x by root-derived organic acids.
A soil conservation role that inheres in a grapevine whose long-term presence and associated management actively increase soil organic matter content, improve soil structure and aggregate stability, enhance biological activity, and deepen the effective rooting zone, resulting in measurable improvement of soil quality and productive capacity over multi-decadal timescales.
Soil_Building_Role captures the constructive complement to erosion control — the active improvement of soil resources through vine-mediated biological and physical processes. This role is most evident in long-established vineyards where decades of root growth, organic matter cycling, and biological activity have measurably enhanced soil properties beyond their pre-viticultural baseline: the process is analogous to pedogenesis accelerated by perennial plant establishment. The role is maximized under management systems that return organic matter to the soil (cover crops, composting, mulching with pruning residues) and minimized under practices that deplete soil carbon (intensive tillage, bare-soil management, residue removal).
Bodenaufbau-Rolle
Bodenregeneration-Rolle
Bodenverbesserungs-Rolle
pedogenesis enhancement role
soil improvement role
soil regeneration role
Ruiz-Colmenero M et al. (2013) Vegetation cover reduces erosion and enhances soil organic carbon in a vineyard, Catena 104: 153-160 Celette F et al. (2009) Spatial and temporal changes to the water regime of a Mediterranean vineyard, European Journal of Agronomy 31: 127-135 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 15-45
Soil Building Role
The soil chemistry data for a Barossa Valley Shiraz vineyard (South Australia) documents the comprehensive analytical profile — pH 7.8 (1:5 water), organic matter 1.8%, CEC 18 cmol/kg, with adequate macronutrient status (nitrogen 0.12% total N, phosphorus 25 mg/kg Olsen-P, potassium 280 mg/kg exchangeable K, calcium 2,800 mg/kg exchangeable Ca, magnesium 450 mg/kg exchangeable Mg) and no micronutrient deficiencies (iron 45 mg/kg DTPA-Fe, manganese 18 mg/kg DTPA-Mn, zinc 3.2 mg/kg DTPA-Zn, boron 1.8 mg/kg hot water-B, copper 4.5 mg/kg DTPA-Cu). This chemical profile indicates a calcareous, moderately fertile soil with no nutritional limitations requiring correction — the moderate fertility supports balanced vine growth without excessive vigor, consistent with the terroir principle that moderate rather than high soil fertility produces higher-quality wine grapes by naturally limiting vegetative growth and promoting the partitioning of assimilates toward reproductive development and berry flavor compound synthesis.
A terroir information component that documents the chemical composition and reactivity of the vineyard soil, recording pH, organic matter content, cation exchange capacity, and the concentrations of macro- and micronutrients that together define the soil's capacity to supply mineral nutrition to the vine, buffer chemical inputs, and maintain the chemical environment in which root function, microbial activity, and nutrient cycling operate — providing the analytical basis for fertilization recommendations, amendment decisions, and the assessment of soil chemical constraints or advantages that contribute to terroir expression.
Soil_Chemistry_Data provides the analytical foundation for vineyard nutritional management — the systematic measurement of soil chemical properties that enables diagnosis of nutrient deficiencies, toxicities, and imbalances before they manifest as visible vine symptoms, allowing preventive rather than corrective fertilization that maintains vine health and fruit quality. The relationship between soil chemistry and wine quality is more nuanced than the physical soil properties (texture, drainage, depth) because the vine's nutritional requirements are relatively modest compared to most agricultural crops — Vitis vinifera evolved on the thin, infertile soils of Mediterranean hillsides and is adapted to low-nutrient environments, producing its highest-quality fruit under moderate nutritional stress rather than under luxury nutrient supply: this adaptation explains the empirical observation that many of the world's finest vineyard soils are chemically infertile by conventional agricultural standards (low nitrogen, low phosphorus, low organic matter), and that excessive fertilization — particularly nitrogen — produces vigorous vegetative growth, dense shading canopies, delayed ripening, and reduced wine quality. The VVO documents soil chemistry through standardized analytical methods (specified for each parameter to ensure comparability across laboratories and regions) at defined sampling depths (typically 0-30 cm for the surface horizon reflecting recent management inputs, and 30-60 cm for the subsoil reflecting the inherent soil chemical character), with results interpreted against viticultural calibration ranges that differ from the conventional agricultural calibrations used for annual crops.
Bodenchemie-Daten
Bodenfruchtbarkeits-Analytik
chemical soil characterization
chemische Bodencharakterisierung
soil analytical data
soil fertility assessment
White RE (2003) Soils for Fine Wines, Oxford University Press Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873 Conradie WJ (2001) Timing of nitrogen fertilisation and the effect on the performance of Bukettraube, South African Journal of Enology and Viticulture 22: 60-68 Robinson JB (2005) Critical plant tissue values and application of nutritional standards for practical use in vineyards, in Christensen LP and Smart DR (eds) Proceedings of the Soil Environment and Vine Mineral Nutrition Symposium, ASEV
Soil Chemistry Data
The soil classification for a representative Côte de Nuits Grand Cru vineyard (Burgundy) records dual classification in the two major systems — WRB: Calcaric Cambisol (Clayic, Chromic), indicating a moderately developed soil (Cambisol) with calcareous parent material (Calcaric qualifier), high clay content (Clayic), and reddish-brown color from iron oxide weathering (Chromic) USDA Soil Taxonomy: Typic Eutrudept, fine-loamy, mixed, mesic, indicating a moderately developed Inceptisol (Ept) of humid regions (Udic moisture regime) with high base saturation (Eutr-), fine-loamy particle size class, mixed mineralogy, and mesic temperature regime. These classifications identify the soil as a young, moderately weathered soil developed on Jurassic limestone parent material — the limited degree of pedogenesis (classified as Cambisol/Inceptisol rather than the more developed Luvisol/Alfisol) reflects the continuous erosion on the Côte d'Or slopes that removes surface material faster than weathering deepens the profile, maintaining the thin, immature soils that are characteristic of the best vineyard positions.
A soil type description component that records the formal taxonomic classification of the vineyard soil within one or more standardized international or national soil classification systems — primarily the USDA Soil Taxonomy (the dominant system in North and South America, Australia) and the World Reference Base for Soil Resources (WRB, the international standard adopted by the FAO and widely used in Europe and Africa) — providing the hierarchical taxonomic identity that enables systematic comparison of vineyard soils across regions and the retrieval of associated soil property data from national and international soil databases.
Soil_Classification provides the formal taxonomic framework that enables vineyard soils to be identified, compared, and mapped within the global soil classification systems developed by pedologists over the past century. The two dominant systems — USDA Soil Taxonomy (first published 1975, current 12th edition 2014) and WRB (first published 1998, current 2015 update) — use different classification philosophies but produce broadly congruent results for most vineyard soils: USDA Soil Taxonomy classifies soils hierarchically into Orders (12), Suborders, Great Groups, Subgroups, Families, and Series based on diagnostic horizons and properties measured at specific depths, with the classification criteria designed for reproducible identification by any trained observer WRB classifies soils into Reference Soil Groups (32) modified by qualifiers that describe specific properties, producing a more flexible classification that accommodates the global diversity of soils. For vineyard soils, the most commonly encountered taxa include Cambisols/Inceptisols (young soils on actively eroding slopes — common in Burgundy, Northern Rhône, Mosel), Luvisols/Alfisols (clay-enriched soils — common in Bordeaux clay-limestone areas), Fluvisols/Entisols (alluvial soils — common in Bordeaux gravel deposits, Rhône alluvium), Leptosols/Entisols (thin soils on rock — common on steep schist and granite slopes), and Calcisols/Aridisols (calcareous soils — common in Mediterranean limestone regions).
Boden-Taxonomie
Bodenklassifikation
formal soil classification
pedological taxonomic identity
pedologische Einordnung
soil taxonomy record
Soil Survey Staff (2014) Keys to Soil Taxonomy, 12th edition, USDA-NRCS IUSS Working Group WRB (2015) World Reference Base for Soil Resources 2014, update 2015, FAO van Leeuwen C et al. (2004) Influence of climate, soil, and cultivar on terroir, American Journal of Enology and Viticulture 55: 207-217 Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873
Soil Classification
The soil color description for a red terra rossa vineyard in Coonawarra (South Australia) documents the striking color profile — 0-30 cm: Munsell 2.5YR 3/6 (dark red, moist), the vivid red color resulting from hematite (α-Fe₂O₃) accumulation in the well-drained limestone residual soil 30-60 cm: 5YR 4/6 (yellowish red), transitioning from hematite-dominated to goethite-influenced coloration with increasing depth and moisture below 60 cm: 10YR 8/1 (white), the underlying Tertiary limestone bedrock (calcrete hardpan) that creates the root-restricting layer and the perched water table controlling the shallow soil hydrology. This red-over-white color sequence is pathognomonic for the Coonawarra terra rossa — the deep red surface color is one of the most visually distinctive vineyard soil colors worldwide and has become an iconic marketing image for the Coonawarra wine region, directly linking the geological terroir (limestone-derived iron-rich residual soil) to regional wine identity.
A soil description component that records the color of the soil at each horizon depth as determined by comparison with the Munsell Soil Color Charts — the standardized color reference system used worldwide in soil science — documenting the hue (spectral color), value (lightness), and chroma (color intensity) notation (e.g., 10YR 4/3, brown) for both moist and dry soil conditions, with soil color providing a rapid visual indicator of organic matter content (dark colors), drainage status (grey/gleyed colors indicating waterlogging, red/brown colors indicating good aeration), and mineralogical composition (red from hematite, yellow from goethite, white from calcium carbonate or quartz) that contributes to vineyard soil characterization.
Soil_Color_Description documents a property that serves simultaneously as a rapid field indicator of soil composition, a standardized pedological descriptor, and in some regions a culturally significant element of terroir identity. The Munsell color system — based on three independent dimensions of hue (the dominant spectral wavelength, recorded as 2.5YR through 5Y in soil applications), value (lightness, scale 0-10 from black to white), and chroma (color saturation, scale 0-8 from grey to vivid) — provides a precise, reproducible color determination that is mandatory in all standard soil profile descriptions and soil survey publications: the system is calibrated for field use, with soil color determined by holding a moist soil sample against the Munsell color chips under natural daylight and recording the closest matching notation. The diagnostic value of soil color derives from the strong correlations between color and composition — organic matter content correlates with value (darker soils have more organic matter, with each 1% increase in organic carbon reducing value by approximately 0.5-1.0 units), iron oxide mineralogy determines hue and chroma (hematite produces red hues of 2.5YR-5YR with high chroma, goethite produces yellow hues of 7.5YR-10YR with moderate chroma, and the absence of iron oxides in waterlogged conditions produces grey gleyed colors of 5Y-5GY with low chroma), and calcium carbonate produces white colors (value >7) that indicate calcareous soil conditions. For vineyard soil characterization, the drainage-indicative colors are particularly informative — the presence of grey (gleyed) mottles or horizons indicates seasonal or permanent waterlogging that restricts root aeration and may require artificial drainage for successful vine establishment.
Bodenfarbe-Beschreibung
Munsell color notation
Munsell-Farbbestimmung
pedological color characterization
pedologische Farbcharakterisierung
soil color record
Soil Survey Staff (2014) Keys to Soil Taxonomy, 12th edition, USDA-NRCS Munsell Color (2009) Munsell Soil-Color Charts, Munsell Color White RE (2003) Soils for Fine Wines, Oxford University Press Soil Science Division Staff (2017) Soil Survey Manual, USDA Handbook 18
Soil Color Description
Riesling vines on the steep terraced slopes of the Mosel Valley bearing a soil conservation role anchor 55-70° gradient Devonian slate soils through root networks penetrating 2-5 m into fractured rock, with each vine's root system stabilizing approximately 0.5-1.0 m³ of soil volume, canopy interception reducing effective rainfall erosivity by 15-25% during summer storms, and annual leaf litter deposition of 1-2 t/ha protecting the soil surface against splash erosion — without vine cover and terrace maintenance, these slopes would experience estimated soil losses of 20-50 t/ha/year versus 2-8 t/ha/year under maintained viticulture.
An ecological role that inheres in a grapevine by virtue of its contribution to the protection, stabilization, and functional maintenance of vineyard soils through root anchoring, canopy rainfall interception, organic matter addition, and the mitigation of erosive forces that would otherwise degrade the soil resource upon which viticulture depends.
Soil_Conservation_Role is critically important in the many viticultural regions characterized by steep slopes, erodible soils, and intense rainfall events — conditions common across Mediterranean Europe, the Mosel and Douro valleys, and terraced hillside vineyards worldwide. Vineyard soil erosion is among the most significant environmental costs of viticulture: estimated soil loss rates of 5-50 t/ha/year in European vineyards substantially exceed soil formation rates of 0.3-1.0 t/ha/year, making soil conservation a sustainability imperative. Subclasses distinguish between erosion control (preventing soil loss) and soil building (actively increasing soil depth and quality).
Bodenerhaltungs-Rolle
Bodenschutz-Rolle
Erosionsschutz-Rolle
soil protection role
soil stabilization role
Prosdocimi M et al. (2016) Soil water erosion on Mediterranean vineyards, Catena 141: 1-21 Rodrigo-Comino J (2018) Five decades of soil erosion research in Mediterranean vineyards, Geomorphology 307: 204-217 Ruiz-Colmenero M et al. (2013) Vegetation cover reduces erosion and enhances soil organic carbon in a vineyard, Catena 104: 153-160
Soil Conservation Role
The soil depth description for the Mosel Steillagen (steep-slope vineyards, Germany) documents very shallow to shallow soils of 20-50 cm depth overlying fractured Devonian slate bedrock — the thin soil mantle on these 50-65% gradient slopes is continuously rejuvenated by mechanical weathering of the underlying slate (frost action fragmenting the bedrock surface) and continuously depleted by gravitational erosion, producing a dynamic equilibrium soil depth that has been maintained for centuries by the traditional practice of Bodenrücktrag — the annual hand-carrying of eroded soil from the slope base back to the upper vineyard, a labor-intensive practice that has been largely abandoned in modern viticulture. The shallow soil depth severely limits the vine's accessible water reservoir (40-80 mm plant-available water compared to 200-300 mm on deep alluvial soils), imposing the natural water limitation during ripening that promotes the concentrated, mineral-inflected Riesling character considered the hallmark of Mosel Steillagen wines — the vine roots extend into the fractured slate to access additional water stored in bedrock fissures, a root exploration pattern visible in road cuts and quarry exposures where vine roots have been observed penetrating 3-5 m into the fractured slate.
A soil description component that records the total depth of the soil profile from the surface to a root-restricting layer — bedrock, hardpan, permanent water table, or other impenetrable barrier — expressed in centimeters and classified as very shallow (150 cm), with soil depth determining the maximum potential rooting volume that defines the vine's accessible water reservoir, nutrient pool, and anchorage capacity, and consequently the vine's resilience to drought, its vigor potential, and the degree of natural growth limitation that contributes to terroir expression.
Soil_Depth_Description documents the volumetric constraint on root system development that is one of the most influential soil properties for vine behavior and wine quality — Seguin's terroir studies demonstrated that the best Bordeaux vineyard soils share the common feature of sufficient depth for deep root development (typically >1.5 m on gravel soils) but with limited water-holding capacity per unit depth, producing the moderate total water reservoir that sustains vine function without promoting excessive vigor. The relationship between soil depth and vine performance is mediated by the interaction of depth with water-holding capacity per unit depth — a shallow soil with high water-holding capacity (e.g., 50 cm of clay retaining 120 mm of plant-available water) may provide a similar total water reservoir to a deep soil with low water-holding capacity (e.g., 200 cm of gravel retaining 120 mm of plant-available water), but the vine response differs because the temporal pattern of water uptake is depth-dependent: shallow-rooted vines on thin soils experience rapid wetting-drying cycles (quick recharge by rainfall, quick depletion by transpiration) producing episodic water stress, while deep-rooted vines on deep soils experience slower, more buffered water supply dynamics producing more gradual and sustained stress development. The effective rooting depth may differ from the total soil depth if root-restricting layers (compaction pans, high-density clay horizons, toxic horizons with extreme pH or salinity) are present above the bedrock — these internal barriers are documented separately from the total soil profile depth because they may be ameliorable through deep ripping, subsoil amendments, or rootstock selection.
Bodentiefe-Beschreibung
Profiltiefe-Dokumentation
depth to bedrock measurement
effective rooting depth
effektive Durchwurzelungstiefe
soil profile depth record
Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873 van Leeuwen C et al. (2004) Influence of climate, soil, and cultivar on terroir, American Journal of Enology and Viticulture 55: 207-217 White RE (2003) Soils for Fine Wines, Oxford University Press Morlat R (1998) Les relations entre le terroir, la vigne et le vin, Revue des Oenologues 87: 9-17
Soil Depth Description
The soil description for the Pétrus vineyard (Pomerol, Bordeaux, 44.93°N) documents the unique pedological conditions underlying this celebrated estate — a lens of blue-grey smectitic clay (montmorillonite-dominated, 60-80% clay content in the upper 40 cm) overlying a deeper layer of iron-enriched clay-gravel (crasse de fer) on a gently elevated plateau at 35-40 m elevation, with the heavy clay providing exceptional water-holding capacity (200-250 mm plant-available water in the rooting zone) that sustains vine function during summer drought while the iron-rich hardpan at 60-80 cm depth limits root penetration and creates a perched water table during winter that necessitates drainage tile installation. This heavy clay soil — anomalous within the predominantly gravel-dominated Pomerol appellation — is the pedological basis of the Pétrus terroir, providing the consistent water supply that enables Merlot (which dominates the Pétrus planting at 100%) to achieve the voluptuous, richly textured wines that distinguish Pétrus from the more structured, gravel-soil Merlots of neighboring estates.
A terroir information component that documents the pedological characteristics of the vineyard soil — the unconsolidated mineral and organic material at the earth's surface in which vine roots grow — recording soil type, texture, structure, depth, drainage, chemical composition, and the parent material from which the soil developed, providing the comprehensive soil characterization that determines vine water availability, nutrient supply, root development patterns, and the degree of natural growth limitation that is the pedological component of terroir expression.
Soil_Description is the root class for the pedological component of terroir characterization — the domain that has the longest history of systematic study in viticulture, dating from Columella's observations on soil-wine relationships in De Re Rustica (c. 65 AD) through the pioneering terroir studies of Gérard Seguin at the University of Bordeaux (1960s-1980s) to the modern digital soil mapping and precision viticulture soil sensing technologies of the 21st century. Seguin's foundational work established the principle that soil quality for viticulture is determined primarily by the soil's physical properties (particularly water-holding capacity and drainage) rather than its chemical fertility — his studies across the Bordeaux appellations demonstrated that the highest-quality vineyard soils are those that provide moderate but not excessive water supply to the vine, limiting vegetative vigor and promoting the moderate water stress during ripening that concentrates berry flavor compounds and accelerates phenolic maturation: this principle of natural growth limitation through soil physical properties has been validated across viticultural regions worldwide and remains the central tenet of soil-based terroir theory. The VVO documents vineyard soils through a hierarchy of descriptors — soil type and classification (the taxonomic identity in standard pedological systems), soil texture (the particle size distribution that determines water-holding capacity and drainage), and additional chemical and physical properties — that together provide the comprehensive soil characterization needed for terroir research and precision viticulture soil management.
Bodenbeschreibung
Weinberg-Bodencharakterisierung
pedological site description
pedologische Standort-Dokumentation
soil profile documentation
vineyard soil characterization
Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873 van Leeuwen C and Seguin G (2006) The concept of terroir in viticulture, Journal of Wine Research 17: 1-10 Wilson JE (1998) Terroir — The Role of Geology, Climate, and Culture in the Making of French Wines, University of California Press van Leeuwen C et al. (2004) Influence of climate, soil, and cultivar on terroir, American Journal of Enology and Viticulture 55: 207-217
Soil Description
The soil management description for a biodynamically farmed Pinot Noir vineyard in Vosne-Romanée documents an integrated floor management system comprising alternate-row permanent grass cover (Festuca ovina at 80% and Trifolium repens at 20%) maintained at 10-15 cm height by mowing 4-5 times per season in even-numbered inter-rows, shallow mechanical tillage (10-15 cm depth using an intercep or disc cultivator) in odd-numbered inter-rows performed 3 times between March and June to control weeds and incorporate biodynamic preparation 500 (horn manure) applied at 100 g/ha in 35 liters of dynamisized water, annual compost application of 3-5 t/ha of composted grape marc and cow manure in autumn, and no synthetic herbicides, fungicides applied to soil surface, or mineral nitrogen fertilizers — the system targets soil organic matter maintenance above 2.5% and earthworm density above 200 individuals per m² as indicators of soil biological health.
A viticultural practice description that documents the methods, timing, and materials applied to manage the vineyard soil environment, encompassing tillage practices, cover cropping, mulching, composting, and soil amendment strategies that collectively influence soil structure, organic matter content, water infiltration and retention, nutrient availability, root zone temperature, erosion control, and the biological activity of the soil ecosystem within which grapevine roots function.
Soil_Management_Description addresses the domain of viticultural practice that has undergone the most dramatic philosophical shift over the past three decades — from the mid-20th century paradigm of clean cultivation (bare soil maintained by tillage or herbicides to eliminate all competition with the vine) toward integrated floor management systems that recognize the soil as a living ecosystem whose biological health directly influences vine performance and wine quality. This paradigm shift has been driven by converging evidence from soil science, microbial ecology, and long-term vineyard sustainability research demonstrating that clean cultivation degrades soil structure, reduces organic matter, accelerates erosion, diminishes mycorrhizal colonization, and ultimately undermines the terroir expression that premium viticulture seeks to capture: contemporary soil management integrates physical (tillage), biological (cover crops, compost), and chemical (amendments) interventions into systems designed to maintain or enhance soil health while managing the competition for water and nutrients between cover vegetation and grapevines. Subclasses capture the principal soil management interventions.
Bodenbewirtschaftungs-Protokoll
Bodenmanagement-Dokumentation
Bodenpflege-Beschreibung
floor management documentation
soil cultivation protocol
vineyard soil management record
White RE (2015) Understanding Vineyard Soils 2nd Ed, Oxford University Press Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 421-460 Goulet E and Morlat R (2011) The use of surveys among wine growers in vineyards of the middle-Loss Valley of France to study the sustainability of vineyard floor management practices, Journal International des Sciences de la Vigne et du Vin 45: 147-162
Soil Management Description
The soil moisture threshold specification for a Chardonnay vineyard in the Yarra Valley defines irrigation trigger points at two depths — at 0.3 m depth a volumetric water content of 22% (corresponding to approximately 50% depletion of plant-available water in the clay loam soil with field capacity of 32% and permanent wilting point of 14%) triggers irrigation initiation, while at 0.6 m depth a soil water potential of -80 kPa measured by granular matrix sensors (Watermark) serves as the lower threshold below which irrigation is applied regardless of the shallower reading, ensuring that the primary root zone (0.3-0.6 m in the structured clay loam) does not experience water stress beyond the moderate deficit target during the pre-véraison period.
An irrigation protocol description that documents the soil water content values at defined depths that serve as trigger points for initiating or terminating irrigation events, specifying the sensor type, measurement depth, and critical threshold values (expressed as volumetric water content, soil water potential, or percentage of plant-available water) that translate the soil moisture continuum into operational irrigation decisions.
Soil_Moisture_Threshold defines the soil-based decision criteria for irrigation scheduling that complement atmospheric demand calculations and plant-based water status indicators. The choice of threshold values requires integration of soil physical properties (texture, structure, water retention characteristics defined by the soil water retention curve), root distribution patterns (typically 60-80% of Vitis vinifera roots in the upper 0.6 m in irrigated vineyards), and the target vine water status for the current phenological phase: threshold values are typically set higher (wetter trigger point, less depletion allowed) during critical periods such as flowering and fruit set where water stress reduces fruit set and berry number, and lower (drier trigger point, more depletion permitted) during the post-véraison deficit period where moderate stress is desired for quality enhancement. Sensor technologies for soil moisture monitoring include time-domain reflectometry (TDR), frequency-domain capacitance probes, granular matrix sensors, tensiometers, and neutron probes, each with distinct measurement principles, accuracy ranges, and cost-performance characteristics.
Bewässerungsauslöse-Grenzwert
Bodenfeuchte-Schwellenwert
Bodenwasser-Triggerpunkt
irrigation refill threshold
soil moisture set point
soil water trigger point
Acevedo-Opazo C et al. (2010) Effects of grapevine (Vitis vinifera L.) water status on water consumption, vegetative growth and grape quality, Irrigation Science 28: 231-243 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 395-420 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 178-195
Soil Moisture Threshold
The soil pH data for a Chablis Grand Cru vineyard (Burgundy) documents alkaline conditions — pH 8.0-8.2 (1:2.5 water) in the upper 30 cm, increasing to pH 8.3-8.5 in the subsoil at 30-60 cm depth — reflecting the high calcium carbonate content (15-25% CaCO3) of the Kimmeridgian limestone marl parent material that buffers the soil solution strongly against acidification. This alkaline pH creates the characteristic nutrient availability pattern of calcareous vineyard soils — abundant calcium and magnesium (the dominant exchangeable cations), adequate potassium and phosphorus (though phosphorus availability is reduced by calcium phosphate precipitation above pH 7.5), but restricted availability of iron (forming insoluble ferric hydroxide at high pH), manganese, zinc, and copper (all less soluble under alkaline conditions) — the iron limitation being the most practically significant, frequently producing lime-induced chlorosis (yellowing of young leaves from inadequate iron uptake) on susceptible rootstocks that lack the root acidification mechanisms needed to solubilize iron in calcareous soils.
A soil chemistry data component that records the hydrogen ion activity of the vineyard soil solution, measured as pH in a standardized suspension (1:2.5 soil-to-water ratio, or alternatively 1:2.5 soil-to-0.01M CaCl₂ for salt-corrected pH), with soil pH being the master chemical variable that controls the solubility and plant availability of virtually all mineral nutrients, determines the species composition and activity of the soil microbial community, and influences the chemical form and toxicity of potentially harmful elements (aluminum, manganese) in acid soils.
Soil_pH_Data documents the single most informative chemical measurement for vineyard soil characterization — soil pH integrates the effects of parent material weathering, cation leaching, biological acid production, and management inputs (liming, acidifying fertilizers) into a single value that predicts the chemical environment experienced by vine roots. The viticultural pH range spans from approximately 4.5 (extremely acid soils on highly weathered granite or quartzite in humid climates) to 8.5 (strongly calcareous soils on limestone or chalk), with the optimal range for grapevine nutrition generally considered to be 5.5-7.5, within which all essential nutrients are reasonably available: below pH 5.5, aluminum toxicity becomes a risk (Al³⁺ solubility increases sharply below pH 5.0, inhibiting root elongation and causing root tip necrosis), while above pH 7.5, micronutrient availability (particularly iron, zinc, and manganese) decreases to potentially deficiency-inducing levels. The practical significance of soil pH for viticultural management centers on two decisions — rootstock selection (calcareous soils with pH >7.5 require rootstocks with high lime tolerance, quantified as the chlorosis tolerance index, to prevent lime-induced chlorosis the most lime-tolerant rootstocks include 41B, Fercal, and 333EM, while sensitive rootstocks like SO4 and 3309C fail on calcareous soils) and amendment strategy (acid soils may require liming to raise pH above the aluminum toxicity threshold, while calcareous soils may benefit from acidifying amendments or foliar iron application to correct micronutrient limitations).
Boden-Säuregrad
Boden-pH-Daten
Bodenreaktion
hydrogen ion activity
soil acidity-alkalinity record
soil reaction measurement
White RE (2003) Soils for Fine Wines, Oxford University Press Bavaresco L et al. (2003) Role of rootstock on iron-chlorosis, Acta Horticulturae 603: 87-95 Soil Survey Staff (2014) Keys to Soil Taxonomy, 12th edition, USDA-NRCS Conradie WJ (2001) Timing of nitrogen fertilisation, South African Journal of Enology and Viticulture 22: 60-68
Soil pH Data
The soil series name for a vineyard in the Rutherford AVA (Napa Valley, California) records the Bale series — a well-drained, deep alluvial soil classified as Fine-loamy, mixed, superactive, thermic Cumulic Haploxerolls in USDA Soil Taxonomy, developed in alluvium derived from mixed sedimentary and volcanic parent materials on the gently sloping Rutherford Bench at 60-80 m elevation. The Bale series is documented in the Napa County Soil Survey (USDA-NRCS) with a standard profile description (0-25 cm dark brown loam, 25-60 cm brown clay loam, 60-120 cm stratified gravelly loam and sandy loam), laboratory analytical data (pH 6.5-7.0, CEC 20-25 cmol/kg, 2-3% organic matter), and management interpretations (moderate available water capacity of 150-200 mm, no significant limitations for deep-rooted perennial crops). The identification of the Bale series links this specific vineyard to the comprehensive soil property database maintained by the NRCS Web Soil Survey, enabling retrieval of detailed physical, chemical, and hydrological data without requiring site-specific soil analysis.
A soil type description component that records the named soil series (in the USDA system) or the locally recognized soil mapping unit that provides the finest level of soil taxonomic identification, linking the vineyard soil to the detailed profile description, analytical data, and management interpretations published in county soil surveys and national soil databases that document the characteristic properties and behavior of each named soil unit.
Soil_Series_Name provides the finest-resolution taxonomic identification available in the USDA soil classification system — the soil series is defined as a group of soils with similar profile characteristics (horizon sequence, color, texture, structure, pH, mineralogy) developed from similar parent material under similar conditions, occupying a consistent landscape position. Approximately 25,000 soil series have been defined and mapped in the United States, with each series documented by a detailed Official Series Description (OSD) published by the USDA-NRCS that specifies the typical profile, range of characteristics, geographic distribution, and associated soils: the series name provides a shorthand reference to this comprehensive documentation, enabling vineyard soil properties to be estimated from the series identification without expensive site-specific soil analysis. The soil series concept is most developed in the United States (where the national cooperative soil survey has mapped soils at series level across the entire country at 1:12,000 to 1:24,000 scale) and in Australia (where soil landscape mapping provides series-level identification in many viticultural regions), but is less systematically applied in European viticultural countries where soil mapping has traditionally focused on higher taxonomic levels or on region-specific classification systems (e.g., the French carte des sols, the German Bodenkarte) rather than the universally standardized series concept. For European vineyard soils, the equivalent finest-resolution identification is typically the soil mapping unit defined in regional soil surveys, which provides similar profile-level characterization without the standardized series nomenclature.
Bodeneinheit-Bezeichnung
Bodenkartierungseinheit
Bodenreihen-Name
named soil unit
soil mapping unit identity
soil series record
Soil Survey Staff (2014) Keys to Soil Taxonomy, 12th edition, USDA-NRCS USDA-NRCS Web Soil Survey (https://websoilsurvey.nrcs.usda.gov) Lambert JJ et al. (2008) Spatial distribution of soil properties in a vineyard, Soil Science Society of America Journal 72: 1174-1183
Soil Series Name
The soil structure description for a Châteauneuf-du-Pape vineyard documents strong, coarse granular structure in the upper 20 cm (well-aggregated, friable topsoil with abundant macropores between the 10-20 mm granular peds, facilitating rainfall infiltration and root proliferation), transitioning to moderate medium subangular blocky structure at 20-50 cm (larger aggregates of 20-40 mm with moderate inter-ped porosity), overlying a weakly structured massive clay layer at 50-80 cm above the characteristic galets roulés (large rounded quartzite cobbles) that are the iconic surface feature of this appellation. The strong granular surface structure is maintained by the combination of high organic matter content (2-3%), abundant biological activity (earthworms, mycorrhizal fungi), moderate clay content (25-35%), and the annual input of organic matter from cover crop incorporation and pruning debris, creating the favorable soil physical conditions that enable vine roots to exploit the available water stored among and beneath the surface cobble mulch layer.
A soil description component that records the arrangement of soil particles into aggregates (peds) and the pattern of pore spaces between them, documenting the structural type (granular, blocky, prismatic, platy, or structureless), grade (weak, moderate, strong), and size class (fine, medium, coarse) at each horizon depth, with soil structure being the critical physical property that mediates the relationships between texture and the functional soil behaviors — aeration, water movement, root penetrability, and biological activity — that determine how effectively the soil supports vine growth.
Soil_Structure_Description documents the organizational property that transforms the inherent textural composition of the soil (the sand-silt-clay proportions) into a functional growth medium — a clay soil with strong granular structure behaves very differently from the same clay soil in a massive, structureless condition, because the inter-aggregate pore spaces in the well-structured soil provide the macroporosity for drainage, aeration, and root growth that the textural properties alone would not predict. Soil structure develops through the interaction of physical processes (wetting-drying cycles that create shrinkage cracks in clay-rich soils, freeze-thaw cycles that fracture massive soil into peds), chemical processes (flocculation of clay particles by calcium ions, cementation by iron and manganese oxides), and biological processes (root growth creating channels, earthworm burrowing creating macropores, fungal hyphae binding particles into stable aggregates, organic matter acting as an aggregation agent): the resulting structural organization is therefore a dynamic property that responds to management practices — tillage disrupts existing aggregates and destroys macropores (explaining the shift toward reduced tillage in progressive viticulture), cover cropping maintains structural stability through continuous root activity and organic matter input, and compaction by machinery traffic destroys inter-aggregate porosity and restricts root growth. For vineyard soils, the most favorable structural condition is strong, stable granular or subangular blocky structure throughout the rooting zone, providing the macroporosity for rapid drainage after rainfall (preventing waterlogging), sufficient mesoporosity for plant-available water retention (sustaining vine function between rainfall events), and continuous pore networks for unrestricted root extension to depth.
Bodenaggregation
Bodengefüge-Dokumentation
Bodenstruktur-Beschreibung
ped structure characterization
soil aggregation record
soil fabric documentation
White RE (2003) Soils for Fine Wines, Oxford University Press Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873 Soil Survey Staff (2014) Keys to Soil Taxonomy, 12th edition, USDA-NRCS Morlat R and Jacquet A (2003) Grapevine root system and soil characteristics in a vineyard maintained long-term with or without interrow sward, American Journal of Enology and Viticulture 54: 1-7
Soil Structure Description
A soil surface basal shoot on Vitis vinifera cv. Grenache emerging exactly at soil surface through crack in dry crusty soil, shoot base visible without excavation showing emergence from woody structure beneath surface, careful excavation removing 4cm soil depth around shoot revealing connection to trunk 2cm above graft union confirming scion origin, shoot developed to 142cm length by early August, 17 leaves, activated following severe drought stress combined with trunk Esca symptoms triggering low basal activation, surface position creating vulnerability to cultivation damage (shoot base exposed to mechanical injury from hoes or cultivators), temperature extremes (surface soil experiencing wider temperature swings than subsurface), and potential disease exposure (proximity to soil-borne pathogens), this shoot tentatively retained pending overall vine assessment but requiring careful cultivation practices avoiding mechanical damage and vigilant monitoring for disease development given vulnerable surface position.
A ground-level basal shoot emerging exactly at soil surface interface, characterized by shoot emergence point at 0-2cm depth with shoot base visible at or just below surface, accessibility requiring minimal excavation for origin inspection, typical emergence through soil cracks or loose surface materials, and position creating maximum exposure to surface conditions including temperature extremes, moisture fluctuations, mechanical disturbance, and disease inoculum while providing relatively straightforward origin verification through shallow excavation examining shoot connection to trunk or root structures. Soil surface basal shoots represent the shallowest ground-level origin category, with emergence at surface interface providing partial visibility enabling preliminary assessment without extensive excavation, though confirmation of scion versus rootstock origin still requiring careful examination of connection points and comparison with graft union position, with surface emergence creating particular vulnerability to surface disturbances but offering better accessibility than deeper underground origins.
Soil surface basal shoots occupy the boundary between above-ground and underground origins, with their interface position creating both advantages (relative accessibility, partial visibility enabling preliminary assessment) and disadvantages (exposure to surface disturbances, vulnerability to mechanical damage, extreme microclimate conditions). The surface emergence provides convenient access for origin verification through shallow excavation, enabling relatively quick determination of scion versus rootstock origin critical for management decisions. However, surface position creates ongoing challenges including mechanical damage risk (cultivation equipment, foot traffic, animal activity), temperature stress (surface soil temperature extremes exceeding subsurface stability), moisture fluctuation (surface drying versus subsurface moisture retention), and disease exposure (soil splash, surface pathogen concentrations). Management involves careful origin verification, protection during vulnerable establishment phase, modified cultivation practices avoiding mechanical damage, disease monitoring addressing surface exposure risks, and realistic assessment of long-term viability recognizing complications versus deeper or higher-positioned alternatives.
Bodenoberflächen-Trieb
interface-level basal shoot
oberflächlicher Basaltrieb
surface-emerging shoot
Soil Surface Basal Shoot
Soil temperature data for dark slate soils in the Mosel Valley (Ürziger Würzgarten) documented surface temperatures exceeding 60°C on south-facing slopes during summer afternoons, with root zone temperatures (30 cm depth) of 22-25°C that persisted 2-3 hours after sunset, extending the effective thermal accumulation period and contributing to the distinctive tropical fruit and spice notes in late-harvested Riesling. Soil temperature profiles in Coonawarra's terra rossa over limestone showed rapid surface warming (15-30°C diurnal range at 5 cm) but remarkable thermal stability at 50 cm depth (seasonal range of only 10-18°C), maintaining root function during both summer heat and winter cold. Volcanic soil temperature monitoring on Mount Etna's north-facing Nerello Mascalese vineyards (700-900 m elevation) recorded root zone temperatures 3-4°C cooler than equivalent elevation sites on southern exposures, delaying budburst by 10-14 days and contributing to the distinctly higher acidity profiles of Contrada Rampante wines.
Soil temperature data is a microclimate characterization entity that records the thermal regime of vineyard soils at specified depths, including surface temperature, rooting zone temperature profiles, seasonal temperature cycles, diurnal fluctuations, and thermal gradients, expressed in degrees Celsius (°C), providing critical information on root metabolic activity, nutrient uptake capacity, soil biological processes, and the contribution of soil thermal properties to vine water relations and phenological timing.
Soil temperature directly regulates root metabolic rates, water and nutrient uptake kinetics, mycorrhizal activity, and soil microbial processes including nitrogen mineralization — all of which influence vine growth and fruit composition. Root growth initiates when soil temperature at 30 cm depth exceeds approximately 7-10°C (depending on rootstock) and root metabolic activity approximately doubles for each 10°C increase within the physiological range. Soil thermal properties (conductivity, heat capacity, diffusivity) are determined by soil texture, moisture content, organic matter, color, and mineralogy, creating substantial site-to-site variation even within small areas. Dark soils (slate, basalt, dark loam) absorb and re-radiate more longwave radiation than light-colored soils (chalk, limestone, sand), creating measurable differences in canopy air temperature and frost risk. Soil temperature monitoring at standardized depths (10, 30, and 50 cm) using buried thermocouples or thermistors provides the most reliable characterization, though thermal infrared remote sensing offers spatial coverage of surface temperature patterns. Mulching, cover cropping, and tillage practices significantly modify soil thermal regimes and should be documented alongside temperature measurements.
Bodentemperaturdaten
Bodentemperaturverlauf
Bodenwärmeprofil
Wurzelzonentemperatur
edaphic temperature record
edaphische Temperaturdaten
root zone temperature
soil heat regime data
soil thermal data
vineyard soil temperature
Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 345-360 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 85-98 Van Leeuwen C et al. (2004) Influence of climate, soil, and cultivar on terroir. American Journal of Enology and Viticulture 55(3): 207-217
Soil Temperature Data
The soil texture description for a representative Chablis Premier Cru vineyard (Burgundy, 47.8°N) documents a silty clay loam texture in the upper 30 cm — with a particle size distribution of approximately 25% sand, 40% silt, and 35% clay — overlying a more clay-rich subsoil (45-55% clay) developed on Kimmeridgian limestone marl parent material. This silty clay loam texture provides the moderate water-holding capacity (170-220 mm plant-available water in the 1 m rooting zone) that sustains Chardonnay through the ripening period without excessive vigor, combined with the high calcium carbonate content (15-25% CaCO3) that maintains alkaline pH (7.8-8.2) and the iron-limiting conditions that some researchers have associated with the distinctive mineral-flinty aromatic character of Chablis wines. The textural analysis is performed by pipette or hydrometer sedimentation methods on samples collected from standardized depths (0-30 cm, 30-60 cm, 60-100 cm) at georeferenced sampling points within the vineyard.
A soil description component that documents the particle size distribution of the fine earth fraction (<2 mm diameter) of the vineyard soil, recording the proportions of sand (0.05-2.0 mm), silt (0.002-0.05 mm), and clay (<0.002 mm) that together determine the textural class (as defined by the USDA texture triangle or equivalent international systems) and provide the primary physical determinant of soil water-holding capacity, drainage rate, aeration, root penetrability, and workability that are the most consequential soil properties for vine growth and terroir expression.
Soil_Texture_Description documents the fundamental physical property of soil — particle size distribution — that Gérard Seguin identified as the most important soil characteristic for viticultural terroir expression. The three particle size fractions (sand, silt, clay) have fundamentally different physical behaviors — sand particles create large pore spaces that drain freely under gravity (providing aeration but retaining little water), clay particles create small pore spaces that retain water against gravity through capillary forces (providing water storage but potentially limiting aeration), and silt particles provide intermediate behavior: the proportional mixture of these fractions determines the soil's textural class (e.g., sandy loam, clay loam, silty clay) and consequently the soil's hydrological behavior — its ability to accept rainfall (infiltration rate), retain water for vine use (plant-available water capacity), drain excess water (saturated hydraulic conductivity), and supply water to roots during dry periods (unsaturated hydraulic conductivity). The textural triangle classification system (developed by the USDA and adopted internationally with minor variations) defines 12 textural classes based on the sand-silt-clay proportions, providing a standardized vocabulary for communicating soil physical character — sandy soils (>70% sand) are well-drained but droughty, clay soils (>40% clay) are water-retentive but potentially waterlogged, and loamy soils (balanced proportions) provide the intermediate water regime that is often considered optimal for quality viticulture.
Bodenart-Dokumentation
Bodentextur-Beschreibung
Korngrößenverteilung
particle size distribution record
soil granulometry
textural analysis documentation
Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873 van Leeuwen C et al. (2004) Influence of climate, soil, and cultivar on terroir, American Journal of Enology and Viticulture 55: 207-217 Soil Survey Staff (2014) Keys to Soil Taxonomy, 12th edition, USDA-NRCS White RE (2003) Soils for Fine Wines, Oxford University Press
Soil Texture Description
The soil type description for the Grands Crus of the Médoc (Bordeaux) documents deep Quaternary gravel deposits — formally classified as Fluvisols (WRB) or Typic Udifluvents (USDA Soil Taxonomy), these soils consist of 40-80% rounded quartz and quartzite gravel in a sandy-loamy matrix deposited by the ancestral Garonne and Dordogne rivers during Günzian and Mindélian glacial periods (800,000-200,000 years before present), producing the exceptionally well-drained, low-fertility, thermally favorable soils that Seguin identified as the common pedological feature of the First Growth classified estates (Latour, Lafite, Margaux, Mouton, Haut-Brion). The deep gravel deposits provide excellent drainage (saturated hydraulic conductivity >100 mm/hour), low water-holding capacity (80-120 mm plant-available water, imposing moderate water stress during ripening), and thermal advantages (the gravel surface absorbs solar radiation during the day and re-radiates heat at night, increasing minimum temperatures by 1-2°C and advancing phenological development by 5-10 days compared to adjacent clay soils).
A soil description component that records the taxonomic identity and pedogenetic classification of the vineyard soil, documenting both the formal classification in standardized systems (USDA Soil Taxonomy, WRB/FAO) and the traditional viticultural soil typology (e.g., gravel, limestone, clay, schist, granite, marl) that conveys the practical agronomic and terroir-associated characteristics of the soil to the viticultural community.
Soil_Type_Description bridges the formal pedological classification systems used by soil scientists with the traditional viticultural soil typology used by winegrowers, winemakers, and wine communicators — two parallel nomenclatures that describe the same physical reality but emphasize different properties and serve different communicative purposes. The formal classification systems (USDA Soil Taxonomy with its six hierarchical categories from Order to Series, and the WRB with its Reference Soil Groups and qualifiers) provide precise, internationally standardized soil identification based on diagnostic horizons, properties, and materials observed in the soil profile, but their technical terminology is inaccessible to most viticultural practitioners and consumers: the traditional viticultural typology (gravel/graves, limestone/calcaire, clay/argile, marl/marne, schist/schiste, granite/granit, sand/sable, loess/löss, volcanic/volcanique) communicates the dominant parent material and textural character in terms that are immediately meaningful to the wine community and that carry established terroir associations (gravel = drainage = Bordeaux quality limestone = structure = Burgundy character schist = heat = Mosel ripening). The VVO documents both systems — the formal classification for scientific precision and database interoperability, and the traditional typology for practical communication — recognizing that the viticultural significance of soil type lies not in the taxonomic category per se but in the physical and chemical properties (drainage, water-holding capacity, thermal behavior, fertility, pH) that the soil type implies.
Bodentaxonomie-Dokumentation
Bodentyp-Beschreibung
pedological type documentation
pedologische Typisierung
soil classification record
soil taxonomy characterization
Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873 van Leeuwen C and Seguin G (2006) The concept of terroir in viticulture, Journal of Wine Research 17: 1-10 Wilson JE (1998) Terroir — The Role of Geology, Climate, and Culture in the Making of French Wines, University of California Press Soil Survey Staff (2014) Keys to Soil Taxonomy, 12th edition, USDA-NRCS IUSS Working Group WRB (2015) World Reference Base for Soil Resources 2014, update 2015, FAO
Soil Type Description
Solar radiation data for Stellenbosch, South Africa documents annual global radiation of 6800-7200 MJ/m², with growing season totals of 4200-4600 MJ/m² supporting exceptional phenolic development in Cabernet Sauvignon. The Mosel Valley receives only 3600-4000 MJ/m² annually, yet the steep south-facing slopes (30-65° inclination) effectively increase radiation interception per unit ground area by 30-40%, enabling Riesling to achieve full physiological ripeness at this northern latitude (~50°N). Mendoza's high-altitude vineyards (900-1500 m) receive 7000-7500 MJ/m² annually with enhanced UV-B flux, driving the intense anthocyanin accumulation characteristic of high-elevation Malbec.
Solar radiation data is a terroir information entity that documents the quantity, quality, spectral composition, and temporal distribution of electromagnetic energy received from the sun at a viticultural site, encompassing both direct beam and diffuse radiation components, and serving as a fundamental driver of photosynthetic carbon fixation, berry thermal accumulation, phenolic biosynthesis, and aromatic compound development in Vitis vinifera.
Solar radiation data constitutes the energy input foundation of all viticultural production, driving both photosynthetic carbon assimilation and the thermal regime of canopy and fruit. The information entity must capture multiple radiation components — global (direct + diffuse), photosynthetically active radiation (PAR, 400-700 nm), UV radiation (particularly UV-B, 280-315 nm), and far-red/red ratios — as each influences distinct physiological processes. The interaction between radiation and canopy architecture is bidirectional: radiation drives growth that modifies canopy structure, which in turn determines light interception efficiency and fruit zone exposure. Altitude, latitude, aspect, cloud cover, aerosol loading, and atmospheric ozone concentration all modulate the radiation environment. Climate change effects on radiation are complex, involving shifts in cloud patterns, aerosol concentrations, and growing season length that may increase or decrease cumulative radiation at specific sites.
Einstrahlungsdaten
Solarstrahlungsinformation
Sonnenstrahlungsdaten
Strahlungsdaten
insolation data
radiation data
solar energy data
solar radiation record
vineyard radiation information
Smart RE, Robinson MD (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles, pp.: 1-25 Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 47-65 Gladstones J (2011) Wine, Terroir and Climate Change, Wakefield Press, pp.: 73-95
Solar Radiation Data
Soluble sugar accumulation quality assessment in Shiraz under regulated deficit irrigation characterized the carbohydrate dynamics underlying this cultivar's drought adaptation in Australian viticulture. Progressive water deficit induced substantial soluble sugar accumulation quality response with leaf glucose increasing from 15 to 38 mg g⁻¹ dry weight (2.5-fold), fructose from 12 to 32 mg g⁻¹ (2.7-fold), and sucrose from 8 to 18 mg g⁻¹ (2.3-fold) as predawn water potential declined from -0.2 to -0.6 MPa. The collective sugar accumulation contributed approximately 0.3 MPa to osmotic adjustment, representing the largest single component of total adjustment in this cultivar. Isotope labeling studies revealed that 60% of accumulated sugars derived from current photosynthesis continuing at reduced rates under moderate stress, while 40% originated from starch degradation mobilizing stored reserves, demonstrating the dual origin of the soluble sugar accumulation quality response. Comparison of pre-véraison versus post-véraison stress timing showed that soluble sugar accumulation quality was more pronounced during pre-véraison stress when vegetative growth created the primary carbon sink, with post-véraison stress showing competition between osmotic adjustment and fruit sugar accumulation that limited leaf sugar buildup while maintaining berry ripening.
Soluble sugar accumulation quality is an osmotic adjustment quality that inheres in a grapevine organism and represents the capacity to accumulate soluble carbohydrates including glucose, fructose, sucrose, and related sugars in response to water deficit stress, contributing to osmotic adjustment through their substantial osmotic activity while providing additional benefits as readily available carbon reserves, membrane protectants, and signaling molecules that modulate stress-responsive gene expression. This quality captures the carbohydrate component of osmotic adjustment, recognizing that sugars often represent the largest single contributor to osmotically active solute pools in stressed grapevine tissues due to their high concentrations achievable without metabolic toxicity and their central position in plant carbon metabolism enabling rapid mobilization and accumulation under changing conditions. The ontological significance of soluble sugar accumulation quality lies in the dual role of accumulated sugars as both osmotic effectors enabling turgor maintenance and as metabolic reserves supporting recovery and resumed growth when stress is relieved, creating advantages that extend beyond the immediate stress period into the post-stress recovery phase. The biological basis of soluble sugar accumulation encompasses multiple processes including the stress-induced inhibition of growth that reduces sugar consumption in expanding tissues, the activation of starch degradation releasing stored carbohydrates, the continued photosynthetic carbon fixation under moderate stress before stomatal closure becomes severe, and the regulated transport of sugars from source to sink tissues under stress conditions. The contribution of sugars to osmotic adjustment can be substantial, with glucose and fructose concentrations in stressed leaves potentially doubling or tripling to contribute 0.2-0.4 MPa of osmotic potential reduction depending on baseline concentrations and stress intensity. The metabolic flexibility inherent in sugar accumulation, with multiple pathways contributing to the soluble sugar pool, creates a robust response that is less dependent on any single enzyme or pathway than the proline accumulation mechanism.
Soluble_Sugar_Accumulation_Quality is_a Osmotic_Adjustment_Quality, representing the carbohydrate contribution to osmotic adjustment under drought stress. This class addresses the often predominant sugar component of osmolyte accumulation. The dual role as osmotic effector and carbon reserve distinguishes sugars from other osmolytes. Multiple pathways (reduced consumption, starch degradation, continued photosynthesis) contribute to accumulation robustness. The competition with fruit ripening during post-véraison stress creates important phenological considerations. Quantification in mg g⁻¹ or μmol g⁻¹ enables standardized comparison.
Kohlenhydrat-Akkumulation
Zucker-Osmoprotektion
Zucker-osmotische Anpassung
carbohydrate accumulation
lösliche Zucker-Akkumulation
soluble carbohydrate response
sugar osmoprotection
sugar osmotic adjustment
Soluble Sugar Accumulation Quality
Biochemical and molecular characterization of Solute_Accumulation_Disposition at INRAE Montpellier examined compatible solute profiles and biosynthetic gene expression across 18 cultivars during controlled drought stress. Leaf proline, total soluble sugars, and potassium were quantified at well-watered and stressed (Ψpd = -0.8 MPa) conditions, with parallel qRT-PCR analysis of VvP5CS (proline synthesis), VvSPS (sucrose phosphate synthase), and VvHKT1 (potassium transport). Solute_Accumulation_Disposition showed substantial cultivar variation in both magnitude and composition. Total solute increase (expressed as osmotic contribution) ranged from 0.15 MPa ('Pinot Noir') to 0.68 MPa ('Sultana'). Proline accumulation ranged from 2.5 μmol g⁻¹ FW ('Chasselas') to 28 μmol g⁻¹ FW ('Shiraz'), with VvP5CS expression correlating strongly with proline level (r = 0.89). Sugar accumulation ranged from 15 mg g⁻¹ FW ('Grenache') to 85 mg g⁻¹ FW ('Sultana'), largely derived from starch hydrolysis rather than continued photosynthesis as evidenced by declining starch reserves. Cultivars showed distinct Solute_Accumulation_Disposition strategies, with 'Shiraz' representing a "proline-dominated" pattern (proline contributing 45% of osmotic adjustment) and 'Sultana' showing a "sugar-dominated" pattern (sugars contributing 55%). 'Cabernet Sauvignon' exhibited balanced accumulation (proline 30%, sugars 35%, K⁺ 20%). Metabolic cost analysis estimated that proline-dominated adjustment required 15% more nitrogen investment than sugar-dominated adjustment, potentially relevant for nitrogen-limited vineyard conditions. These findings demonstrate that Solute_Accumulation_Disposition encompasses both quantitative capacity and qualitative strategy with implications for resource allocation.
A Solute_Accumulation_Disposition is an Osmotic_Adjustment_Disposition that inheres in a Vitis vinifera plant and is realized through the biochemical capacity to synthesize and accumulate specific compatible solutes in response to water stress, providing the osmotically active compounds that lower cellular osmotic potential. This disposition represents the biosynthetic foundation of osmotic adjustment, determining both the magnitude of adjustment possible and the metabolic costs incurred through solute production. Major compatible solutes accumulated during drought in grapevine include proline (synthesized via the glutamate pathway, catalyzed by pyrroline-5-carboxylate synthetase), soluble sugars (derived from starch hydrolysis and continued photosynthesis), sugar alcohols, glycine betaine (in species with active betaine aldehyde dehydrogenase), and inorganic ions (primarily potassium, accumulated via membrane transporters). These compounds are termed "compatible" because they can accumulate to high concentrations without disrupting enzyme function or cellular metabolism, unlike inorganic salts that would cause ionic toxicity at equivalent osmotic contribution. Solute_Accumulation_Disposition involves induction of biosynthetic enzyme genes under stress, metabolic flux toward compatible solutes, and cellular compartmentalization protecting sensitive enzymes from solute interference. Cultivars differ in both the total magnitude of solute accumulation and the specific solute profile produced, with some relying primarily on proline, others on sugars, and most on a combination. The relative metabolic cost varies among solutes, with sugar accumulation potentially conflicting with carbon allocation to growth while proline synthesis requires both carbon and nitrogen investment.
Solute_Accumulation_Disposition is_a Osmotic_Adjustment_Disposition. This class addresses the biochemical basis of osmotic adjustment through compatible solute production. Major solutes include proline, sugars, and potassium, with cultivar-specific profiles. Biosynthetic gene expression (P5CS, SPS) underlies solute accumulation capacity. Different solutes have different metabolic costs and resource requirements (carbon, nitrogen). Solute composition may affect drought tolerance quality beyond osmotic contribution through additional protective functions (proline as antioxidant, sugars stabilizing membranes).
Kompatible-Solut-Synthese
Solut-Akkumulations-Disposition
compatible solute synthesis
osmolyte accumulation
stress metabolite production
Delauney AJ, Verma DPS (1993) Proline biosynthesis and osmoregulation in plants, Plant Journal 4:215-223 Patakas A et al. (2002) Leaf age effects on solute accumulation in water-stressed grapevines, Journal of Plant Physiology 159:969-976 Cramer GR et al. (2007) Water and salinity stress in grapevines: early and late changes in transcript and metabolite profiles, Functional and Integrative Genomics 7:111-134
Solute Accumulation Disposition
The somatic mutation record for the white-berry sport of Cabernet Sauvignon (documented by Walker et al. 2007) describes a deletion of approximately 260 kb on chromosome 2 encompassing both VvMYBA1 and VvMYBA2 transcription factor genes on one homologous chromosome, combined with the pre-existing Gret1 retrotransposon insertion inactivating VvMYBA1 on the other homologue — the combined loss of both functional copies of the MYB-type anthocyanin regulators eliminates anthocyanin biosynthesis in berry skin, producing the white-berry phenotype designated as Shalistin (or Cabernet Blanc in some nursery catalogues). The deletion was identified through BAC library screening and confirmed by fluorescence in situ hybridization (FISH), representing one of the largest characterized somatic structural variants in grapevine and demonstrating that somatic mutations in Vitis can involve major chromosomal deletions rather than only point mutations.
A mutation information record that documents a specific DNA sequence change arising in a somatic (non-germline) cell of a vegetatively propagated Vitis vinifera plant, specifying the mutation type (point mutation, insertion, deletion, transposable element movement, or chromosomal rearrangement), genomic position, affected gene (if applicable), phenotypic consequence (if detectable), the clone or sport in which the mutation was identified, and the detection method (whole-genome resequencing, targeted sequencing, or phenotype-based identification followed by molecular characterization).
Somatic_Mutation_Record provides the molecular-level documentation of individual mutation events that collectively generate intra-cultivar diversity. The spectrum of somatic mutations in grapevine spans multiple scales — from single nucleotide changes (point mutations producing missense, nonsense, or regulatory variants) through small insertions and deletions (indels of 1-50 bp) to large structural variants involving transposable element insertions (the 10.4 kb Gret1 retrotransposon insertion in VvMYBA1 being the most famous example), chromosomal deletions (the 260 kb deletion eliminating VvMYBA1/A2 in white Cabernet Sauvignon), and chimeral rearrangements where mutations in individual meristem layers create periclinal chimeras: the rate of somatic mutation accumulation has been estimated at approximately 3-7 single nucleotide mutations per year of vegetative propagation based on whole-genome comparison of dated clonal lineages (Vondras et al. 2019), implying that clones separated by 500 years of propagation history may differ by 1,500-3,500 somatic SNPs — a sufficient number to produce phenotypically detectable variants at functionally important loci. The documentation of individual somatic mutations is essential for understanding the molecular basis of clonal variation, developing clone-diagnostic molecular markers, and informing clonal selection programs.
Sport-Mutations-Charakterisierung
clonal mutation documentation
klonale Varianten-Erfassung
somatic variant record
somatische Mutations-Dokumentation
sport mutation characterization
Walker AR et al. (2007) White grapes arose through the mutation of two similar and adjacent regulatory genes, Plant Journal 49: 772-785 Vondras AM et al. (2019) The genomic diversification of grapevine clones, BMC Genomics 20: 972 Kobayashi S et al. (2004) Retrotransposon-induced mutations in grape skin color, Science 304: 982 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340
Somatic Mutation Record
The sorting protocol for Opus One (Napa Valley) specifies a four-stage quality filtration system — Stage 1 at the vine (pickers reject visibly damaged clusters during hand harvest), Stage 2 on the reception vibrating table (trained sorters remove leaves, rachis fragments, underripe clusters, and MOG at 2-3 t/h throughput), Stage 3 after destemming (berries pass over a gravity density sorting table that separates lighter underripe and dehydrated berries from optimally dense ripe berries using water flotation), and Stage 4 through an optical berry sorter (Pellenc Selectiv' Process or equivalent, using near-infrared cameras to detect and reject individual berries deviating from target color, size, and surface condition at rates of 6-8 t/h with rejection rates of 5-15% depending on vintage conditions) — the cumulative four-stage rejection rate of 15-25% ensures that only berries meeting strict quality parameters enter the fermentation vessels.
A harvest method description that documents the post-harvest inspection and selection procedures applied to harvested grapes between collection from the vineyard and delivery to the winemaking process, specifying the sorting stages (field sorting, reception sorting, optical sorting), equipment configuration, rejection criteria, and throughput rates that constitute the final quality filter ensuring that only fruit meeting defined quality standards enters the winemaking process.
Sorting_Protocol documents the quality control interface between viticulture and winemaking — the final opportunity to select or reject fruit before it enters the irreversible winemaking process. The evolution of sorting technology from simple manual table sorting to multi-stage automated systems incorporating optical, densimetric, and spectroscopic berry analysis represents one of the most significant quality-enhancement investments in modern premium winemaking: optical sorting machines using machine vision algorithms can inspect and accept or reject individual berries at rates of 4-8 tonnes per hour based on color (identifying underripe green berries, overripe raisined berries, and botrytized berries), size (removing shot berries from poor fruit set), and surface condition (detecting insect damage, sunburn, or mould), achieving a level of selectivity that exceeds even the most skilled human sorter's capability at commercial throughput rates. The adoption of optical sorting has been particularly transformative in vintages with mixed-maturity or disease-affected fruit, where the technology enables production of premium wines from blocks that would previously have been downgraded entirely.
Selektions-Sortierprotokoll
Sortierprotokoll
Trauben-Triage-Verfahren
grape sorting specification
reception sorting procedure
triage protocol
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1, 2nd Ed, Wiley pp: 235-270 Pellenc (2023) Selectiv' Process technical documentation Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 285-295
Sorting Protocol
Source-sink balance assessment through the season in Cabernet Sauvignon reveals the shifting allocation patterns affecting ripening and reserve status, with balanced vine showing sequential sink prioritization (shoot growth dominant through flowering, fruit sink dominant véraison through harvest, reserve accumulation dominant post-harvest) achieving 24.5 Brix at harvest with adequate starch reserves (145 mg/g root DW) for strong spring growth, while overcropped vine showing persistent fruit sink dominance through post-harvest (fruit competing with reserve storage) reaching only 22.5 Brix despite extended hang time and depleted reserves (85 mg/g root DW) predicting weak subsequent growth, demonstrating how source-sink balance affects both immediate quality and long-term vine sustainability.
Source-sink balance quality is a yield efficiency quality that represents the overall equilibrium between carbohydrate production capacity (sources, primarily leaves) and carbohydrate consumption by growing and storage organs (sinks, including fruit, shoots, roots, and reserves), characterizing the fundamental physiological balance that determines carbon allocation patterns, ripening success, and long-term vine sustainability. Within the BFO framework, this quality specifically addresses the whole-plant carbon balance dimension of efficiency, distinguishing this integrative concept from specific ratio metrics while recognizing that source-sink balance encompasses not only the immediate leaf-fruit relationship but the broader allocation of photosynthate among competing sinks including root growth, reserve storage, and vegetative development that together determine vine function and persistence. The ontological significance lies in source-sink balance being the fundamental determinant of vine physiology and production sustainability, where balanced vines allocate carbon appropriately among competing demands (fruit development, structural growth, reserve storage) to achieve current-season quality while maintaining the capacity and reserves for future production, and where imbalance creates either immediate problems (source limitation compromising ripening) or future problems (depleted reserves reducing subsequent vigor and fruitfulness). The dynamic nature of source-sink balance recognizes that the relationship shifts through the season—early season dominated by vegetative sinks (shoot growth), mid-season transitioning toward reproductive sinks (fruit development), and post-harvest prioritizing storage sinks (reserve accumulation)—with management affecting these transitions through crop load control, canopy management, and irrigation practices that modify both source capacity and sink demand. The management of source-sink balance involves optimizing both source (maintaining adequate, well-exposed leaf area) and sink (adjusting crop load to match source capacity) components, with the goal of achieving the balanced allocation that supports full fruit development while maintaining vine health and productive longevity. Understanding source-sink balance quality enables integrative assessment of vine function, diagnosis of allocation problems affecting either current quality or future productivity, and management strategies targeting sustainable balance across the full annual cycle.
Source_Sink_Balance_Quality addresses the whole-plant carbon allocation equilibrium integrating all source and sink components. This quality determines both current-season quality and long-term sustainability. The class supports sustainable production through source-sink balance characterization across the full seasonal cycle.
Assimilatverteilung
Quellen-Senken-Balance
Quellen-Senken-Gleichgewicht
assimilate partitioning
carbon balance
photosynthate allocation
source-sink equilibrium
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 145-230 Holzapfel BP et al. (2010) Manipulating the postharvest period and its impact on vine productivity of Semillon grapevines. American Journal of Enology and Viticulture 61(2): 201-206
Source Sink Balance Quality
Sourness quality assessment of Grüner Veltliner at harvest reveals perceived sourness of 6.8 on a 10-point scale with titratable acidity of 7.2 g/L and pH 3.15, declining to sourness of 5.2 after malolactic fermentation reduces malic acid from 3.8 to 0.4 g/L despite minimal change in titratable acidity (6.9 g/L), demonstrating that sourness perception depends on acid composition not just total acidity, and informing the decision to block malolactic fermentation in this variety to preserve the crisp sourness character valued in the Veltliner style.
Sourness quality is a basic taste quality that represents the gustatory perception of sour taste arising primarily from hydrogen ion (proton) stimulation of acid-sensing ion channels in taste receptor cells, characterizing the sharp, tart sensation associated with organic acids that serves as a fundamental determinant of grape freshness perception and wine balance. Within the BFO framework, this quality specifically addresses the perception of sourness as distinct from titratable acidity or pH measurements, recognizing that sourness perception is modulated by buffering capacity, other taste components, and individual variation in acid sensitivity. The ontological significance lies in sourness providing the counterbalance to sweetness that defines grape and wine taste structure, where the sweetness-sourness ratio determines perceived balance and the absolute sourness level contributes to freshness, crispness, and aging potential. The physiological basis for sourness involves multiple receptor mechanisms including the PKD2L1 ion channel responding to intracellular acidification, with the relationship between acid concentration and perceived sourness depending on acid type, where organic acids with similar pH produce different sourness intensities due to differential effects on intracellular pH. The compositional basis for grape sourness includes tartaric acid (unique to grapes among common fruits, providing stable acidity), malic acid (metabolized during malolactic fermentation, contributing sharp sourness), and minor acids (citric, succinic, lactic) each contributing characteristic sourness qualities. The developmental trajectory of sourness follows acid metabolism during ripening, with malic acid declining through respiration while tartaric acid remains stable, creating the characteristic acidity softening that accompanies sugar accumulation. Sourness perception threshold in wine ranges from 0.5-1.0 g/L depending on acid type and matrix, with wines typically containing 5-9 g/L total acidity producing moderate to high sourness that is balanced against sweetness and other taste components. Understanding sourness quality enables harvest timing for optimal acid balance, winemaking decisions regarding acidification or deacidification, and prediction of sensory outcomes from acid composition analysis.
Sourness_Quality addresses the perceptual experience of sour taste, distinguishing from Acid_Content_Quality which measures the chemical stimulus. This quality serves as parent for intensity, character, and persistence subclasses addressing different aspects of sourness perception. The relationship between measured acidity and perceived sourness is complex, depending on acid type, pH, buffering capacity, and interactions with other taste components.
Säure
Säureempfindung
Säuregeschmack
acid taste perception
acidity perception
saurer Geschmack
sour taste
tartness
Huang AL et al. (2006) The cells and logic for mammalian sour taste detection. Nature 442(7105): 934-938 Plane RA et al. (1980) Acidity and the taste of wine. American Journal of Enology and Viticulture 31(3): 265-267
Sourness Quality
Pinotage clone VIA 2 bears a South_African_Clone_Role as a VIA-certified selection of South Africa's signature variety, subjected to virus elimination protocols addressing the leafroll infections historically prevalent in Pinotage plantings. Chenin Blanc (Steen) clone SMA 593 serves in a South_African_Clone_Role supporting South Africa's substantial Chenin production (the country's most-planted variety), with ARC evaluation ensuring adapted selections for Cape conditions. Shiraz clone SH 9C bears a South_African_Clone_Role with documented heat tolerance and virus-free status, supporting South Africa's expanding premium Shiraz production in regions like Stellenbosch and Paarl.
A regional clone role that inheres in a grapevine clone selected, evaluated, or registered within the South African vine improvement system, coordinated through the Agricultural Research Council (ARC) Nietvoorbij research station, Vine Improvement Association (VIA), and South African Wine and Spirits Board (SAWIS) certification programs. South African clones are identified by variety name and alphanumeric codes indicating institutional origin or evaluation status, with the system emphasizing virus elimination to address historically high virus pressure in the Cape vineyards. The South African clone program developed in response to severe leafroll virus prevalence, with aggressive sanitation and replanting programs transforming vineyard health from the 1980s onward. South African selections include indigenous Pinotage (the local Pinot Noir × Cinsaut cross) and locally-evaluated international varieties, with the certification system distinguishing plant improvement scheme (PIS) material of documented health status from conventional plantings. The Wine of Origin (WO) system integrates clone certification with geographic and varietal labeling, creating comprehensive traceability from vine to bottle.
South_African_Clone_Role captures the distinctive South African system where virus elimination has been particularly emphasized due to historically severe leafroll pressure. The country's wine industry transformation (from bulk production to premium wine focus) coincided with aggressive vine improvement programs, with certified clone adoption central to quality enhancement. South Africa's unique variety—Pinotage, created in 1925—requires domestic selection attention unavailable from international programs, with SA institutions maintaining global responsibility for Pinotage clone development. The integration of clone certification with the Wine of Origin labeling system creates connections between viticultural quality inputs and wine marketing that encourage certified material adoption. Climate change adaptation is increasingly influencing South African clone selection, with heat tolerance and water-use efficiency gaining emphasis alongside traditional quality parameters.
Nietvoorbij clone
Nietvoorbij-Klon
SA clone
SA-Klon
South African clone
South African registered clone
südafrikanischer Klon
Pietersen G (2006) Leafroll virus in South African vineyards, in Extended Abstracts 15th Meeting ICVG, pp.: 38-42 Vine Improvement Association (2020) VIA Certification Standards Burger P," Deist J (1981) Wingerdbou in Suid-Afrika, ARC Nietvoorbij Hunter JJ et al. (2016) Plant material quality and wine quality, South African Journal of Enology and Viticulture 37: 93-102
South African Clone Role
Chardonnay bears a Sparkling_Wine_Scion_Role as the dominant variety for blanc de blancs Champagne and premium sparkling wines globally, providing high acidity (9-11 g/L), citrus and mineral character, and exceptional aging potential over 10-30+ years of lees contact. Pinot Noir serves in a Sparkling_Wine_Scion_Role for blanc de noirs Champagne and rosé sparklings, contributing body, red fruit depth, and structural complexity while maintaining finesse when whole-cluster pressed. Glera bears a Sparkling_Wine_Scion_Role as the required variety for Prosecco (Conegliano-Valdobbiadene DOCG and broader designations), providing floral-fruity aromatics suited to the Charmat method's shorter lees contact. Macabeo, Xarel·lo, and Parellada serve in Sparkling_Wine_Scion_Roles as the traditional Cava trio, each contributing distinct attributes to the blend.
A wine production scion role that inheres in a grafted scion cultivar by virtue of its designation for sparkling wine production, where a secondary fermentation generates carbon dioxide dissolved under pressure (typically 5-6 atmospheres) creating effervescence. Sparkling wine scions are selected for characteristics that optimize base wine quality and mousse development, including high natural acidity (8-12 g/L at harvest) providing freshness and aging potential, moderate sugar accumulation (potential alcohol 9.5-11% before dosage adjustment), neutral-to-subtle aromatic profiles that do not overwhelm autolytic or terroir character, and appropriate phenolic content for style (lower for blanc de blancs elegance, moderate for rosé and blanc de noirs structure). The sparkling role encompasses diverse production methods—traditional method (Champagne), tank method (Charmat), ancestral method (pétillant naturel), and transfer method—each with specific cultivar requirements. Classic Champagne varieties (Chardonnay, Pinot Noir, Pinot Meunier) dominate global premium sparkling production, while regional varieties serve in traditional sparkling styles worldwide.
Sparkling_Wine_Scion_Role is subclassed by production method (Champagne_Method, Charmat_Method, Pet_Nat) reflecting the distinct cultivar requirements and stylistic outcomes of different sparkling wine techniques. The high-acid requirement for quality sparkling production restricts suitable regions to cool climates or elevated sites where physiological ripeness is achieved without excessive sugar accumulation or acid loss. Many cultivars serve dual still and sparkling roles—Chardonnay, Riesling, Chenin Blanc—with vineyard designation often determining utilization. Global sparkling wine consumption has grown substantially, driving expansion of sparkling production beyond traditional regions.
Perlweinrebe
Schaumweinsorte
Sekttraube
base wine variety
sparkling grape variety
sparkling wine variety
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Stevenson T (1998) Christie's World Encyclopedia of Champagne and Sparkling Wine, Absolute Press Jackson RS (2014) Wine Science 4th Ed, Academic Press pp.: 430-480 Liger-Belair G (2005) The physics and chemistry behind the bubbling properties of champagne, Journal of Agricultural and Food Chemistry 53: 2788-2802
Sparkling Wine Scion Role
Sparkling wine specification for Chardonnay-Pinot Noir cuvée from Champagne (NV Brut) documents base wine requirements of 10.5-11.0% alcohol, TA 7.5-9.0 g/L, pH 3.00-3.15, harvested at 17-19° Brix (well below still wine maturity), secondary fermentation in bottle at 11-13°C over 6-8 weeks producing 6.0 atm pressure, minimum 15 months aging on lees (36+ months for prestige cuvées), disgorgement followed by dosage of 6-8 g/L sugar (Brut classification), and final alcohol of 12.0-12.5%. Sparkling wine specification for Grüner Veltliner Sekt from Kamptal (traditional method, Austrian Sekt g.U.) specifies base wine at 11.0% alcohol from grapes harvested at 17° KMW, TA 8.0 g/L, pH 3.05, secondary fermentation in bottle with minimum 18 months on lees for Reserve designation, dosage of ≤6 g/L (Brut Nature to Brut), producing a sparkling wine expressing the characteristic Grüner Veltliner white pepper and citrus notes within a fine-mousse framework — following the Austrian Sekt pyramid classification (Klassik, Reserve, Große Reserve) established by the Österreichisches Sektkomitee.
Sparkling wine specification is a wine style description entity that documents the compositional, sensory, and production-method parameters defining wines with significant dissolved carbon dioxide producing effervescence, encompassing the production method (traditional/méthode champenoise, Charmat/tank, transfer, ancestral/pétillant naturel, carbonation), base wine requirements, secondary fermentation conditions, dosage/liqueur d'expédition composition, aging duration on lees, pressure classification (frizzante/perlant at 1-2.5 atm, spumante/mousseux at 3-6 atm), and the specific cultivar and fruit composition demands that distinguish sparkling wine production from still wine applications of the same grape material.
Sparkling wine specification imposes fundamentally different viticultural requirements compared with still wine production of the same cultivars, creating distinct grape composition targets that optimize base wine suitability for secondary fermentation and extended lees aging. The most critical compositional difference is the harvest sugar target — sparkling base wines require grapes harvested at significantly lower maturity (17-20° Brix / 70-85° Oechsle / 14-17° KMW) compared with still wines (21-26° Brix) to produce base wines with moderate alcohol (10-11.5%) that will gain an additional 1.2-1.5% alcohol during secondary fermentation. High natural acidity (TA 7-10 g/L) is essential for sparkling wine freshness and longevity, making early-harvested, high-acid fruit from cool climates or cool vintages ideal base material. The production method hierarchy — traditional method (secondary fermentation and aging in individual bottles, highest quality and complexity), Charmat/tank method (secondary fermentation in pressurized tanks, preserving primary fruit character), transfer method (secondary fermentation in bottle, transfer to tank for dosage and rebottling), ancestral method (single fermentation completed in bottle, no dosage), and forced carbonation (CO₂ injection, lowest quality) — creates distinct specification requirements at each level. Yeast autolysis during extended lees aging (minimum 12 months for Champagne, 9 months for Cava, 15 months for Crémant) releases mannoproteins, amino acids, and nucleotides that contribute the characteristic brioche, bread dough, and toasty complexity of premium traditional-method sparkling wines. The dosage classification system (Brut Nature 0-3 g/L, Extra Brut 0-6 g/L, Brut 0-12 g/L, Extra Sec 12-17 g/L, Sec 17-32 g/L, Demi-Sec 32-50 g/L, Doux >50 g/L) defines the final sweetness level and substantially influences the sensory profile and food-pairing versatility of the finished wine.
Perlweinspezifikation
Schaumweinherstellungsdaten
Schaumweinparameter
Schaumweinspezifikation
Sekt specification
Sektspezifikation
effervescent wine specification
méthode champenoise specification
sparkling wine parameters
sparkling wine production data
Liger-Belair G (2005) The physics and chemistry behind the bubbling properties of Champagne and sparkling wines. Journal of Agricultural and Food Chemistry 53(8): 2788-2802 Pozo-Bayón MA et al. (2009) Wine volatile and amino acid composition after malolactic fermentation. Journal of Agricultural and Food Chemistry 57(8): 3199-3204 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 721-775 Österreichisches Sektkomitee (2018) Sekt Austria Qualitätspyramide Reglement
Sparkling Wine Specification
Cabernet Franc rare functionally male flowers exhibit sparsely papillate stigmas with 78-118 total papillae distributed across reduced stigma surface (0.28-0.38 mm diameter, giving 110-160 papillae per mm²), with individual papillae 28-42 μm length showing reduced elaboration compared to hermaphrodite stigmas, providing poor pollen capture and germination support with <25% of applied pollen germinating due to minimal exudate production and sparse papillae coverage, contributing to <8% fruit set. Wild V. vinifera ssp. sylvestris staminate male stigmas show extremely sparsely papillate organization with 32-55 total papillae on vestigial stigmas (0.12-0.22 mm diameter, giving 85-145 papillae per mm²), representing minimal stigma elaboration with sparse papillae providing no functional benefit as vestigial stigmas are completely non-receptive producing no exudate and achieving <1% fruit set. Pinot Noir segregating breeding populations produce functionally male offspring with sparsely papillate stigmas (95-135 papillae per mm²) versus hermaphrodite offspring with moderately papillate stigmas (165-220 papillae per mm²), with papillae density serving as early morphological marker for identifying sexual types during seedling evaluation before flowering and fruit set assessment. Comparative microscopy of sparsely papillate functionally male stigmas versus moderately papillate hermaphrodite stigmas reveals 35-55% reduction in total papillae number, 30-45% reduction in papillae density, 25-35% reduction in calculated stigma surface area (including papillae), and 40-70% reduction in observed exudate production during attempted receptivity periods, demonstrating that sparse papillae correlate with multiple indicators of reduced stigma functionality.
A sparsely papillate stigma is a papillate stigma bearing low papillae density typically less than 180 papillae per mm² stigma surface area, characteristic of functionally male flowers with reduced stigmas and staminate flowers with vestigial stigmas where reduced papillae coverage correlates with diminished receptivity and reduced pollen germination support, representing stigmas with suboptimal structure for pollination success. Sparsely papillate stigmas are characterized by= (1) low papillae number typically 60-120 total papillae on stigma surface in functionally male types, 30-60 μm (3) stigma diameter typically 0.2-0.4 mm (functionally male) or 0.1-0.25 mm (staminate vestigial) providing limited surface area for the sparse papillae population (4) papillae dimensions often toward the smaller end of the normal range (20-45 μm length, 12-22 μm diameter) or showing reduced elaboration (5) stigma surface area (including papillae contribution) typically 130-180% of equivalent flat surface area due to sparse papillae coverage, substantially less than densely papillate stigmas (200-280% increase). Sparsely papillate stigmas develop in functionally male and staminate flowers through reduced papillae initiation and/or arrested papillae development during stigma formation, reflecting overall pistil reduction programs that suppress female organ elaboration including stigma and papillae development. The functional implications of sparse papillae relate to impaired pollination capability= low papillae density creates reduced probability of pollen grain contact with papillae, as incoming pollen grains may land in inter-papillae zones reducing adhesion and hydration efficiency sparse papillae provide reduced exudate secretion capacity with fewer secretory cells producing less total exudate and creating more limited wet surface sparse papillae create reduced surface area decreasing pollen retention probability and providing fewer physical barriers and adhesion points sparse papillae support reduced simultaneous pollen germination capacity limiting reproductive success even when adequate pollen is present. Sparsely papillate stigmas in functionally male flowers show correlation with non-receptivity or poor receptivity (minimal or absent exudate production, <30% pollen germination support versus 70-85% for moderately papillate hermaphrodite stigmas), though the sparse papillae themselves are not the primary cause of non-receptivity (which results from failure of exudate secretion systems) but rather correlate with overall stigma reduction. Sparsely papillate stigmas in staminate vestigial stigmas show extreme reduction with papillae number <60 on tiny stigmas 0.1-0.25 mm diameter, creating stigmatic surfaces with minimal elaboration that are completely non-receptive and non-functional, representing developmental remnants rather than functional reproductive structures. The sparse papillae condition can serve as diagnostic feature for identifying functionally male and staminate flowers, as papillae density assessment (through microscopic examination or high-resolution photography) allows inference of flower sexual type based on stigma elaboration level.
Sparsely_Papillate_Stigma represents low papillae density characteristic of functionally male and staminate flowers and is distinguished from Densely_Papillate_Stigma (high density in functionally female/pistillate flowers) by substantially lower papillae counts per unit area (200 papillae per mm²). Sparse papillae correlate with reduced female function and impaired pollination capability. Understanding sparsely papillate stigma structure is relevant for identifying functionally male and staminate flowers, for diagnosing reproductive failures related to stigma inadequacy, and for understanding the correlation between papillae elaboration and stigma functionality across the spectrum of Vitis sexual types.
dünn besetzte Papillen-Narbe
lightly papillate stigma
sparsely textured stigma
spärlich papillöse Narbe
wenig papillentragendes Stigma
Sparsely Papillate Stigma
Spatial yield uniformity assessment using harvest yield monitor in Cabernet Sauvignon reveals the soil-driven production patterns, with clay zone (deeper soil, higher water holding) producing 9.2 t/ha, transitional zone 6.8 t/ha, and gravel zone (shallow, low water holding) only 4.5 t/ha, creating overall spatial CV of 32% that suggests zone-based management rather than whole-block treatment, with subsequent trials showing that the low-yielding gravel zone produces more concentrated, earlier-ripening fruit suitable for premium tier while high-yielding clay zone produces dilute fruit requiring different wine program allocation.
Spatial yield uniformity quality is a yield consistency quality that represents the degree to which grape production is evenly distributed across the vineyard area, characterizing the within-vineyard homogeneity that determines whether the entire block produces similar yields or whether distinct high-producing and low-producing zones create spatial patterns requiring zone-specific management consideration. Within the BFO framework, this quality specifically addresses the spatial dimension of yield consistency, distinguishing geographic production patterns from temporal variation while recognizing that spatial uniformity reflects the physical homogeneity of the site—uniform vineyards arising from consistent soil, topography, and microclimate, while non-uniform vineyards reflect underlying environmental gradients or historical management differences. The ontological significance lies in spatial yield uniformity affecting management efficiency and quality consistency, where uniform vineyards enable block-level management with single practices applied across the entire area, while non-uniform vineyards may require precision viticulture approaches with variable-rate applications, zone-specific harvest timing, or separate processing of fruit from different yield zones to capture quality differences associated with yield patterns. The factors creating spatial yield variation include soil variation (depth, texture, water-holding capacity affecting vine vigor and productivity), topographic effects (slope, aspect, elevation affecting microclimate and growing conditions), drainage patterns (wet zones reducing productivity, well-drained areas supporting consistent growth), rootstock variation (if multiple rootstocks planted, different vigor affecting yield), and historical effects (past management differences, disease history, replanting zones). The precision viticulture applications for understanding spatial yield patterns include yield monitoring during harvest (creating yield maps), remote sensing of canopy characteristics (correlating with yield patterns), soil mapping (identifying underlying causes of spatial variation), and variable-rate management (adjusting inputs to match spatial productivity patterns). Understanding spatial yield uniformity quality enables identification of yield zones within vineyards, targeted management addressing specific low-producing areas, and quality optimization through zone-based harvest and processing strategies.
Spatial_Yield_Uniformity_Quality addresses the geographic distribution of production within vineyards. This quality reflects site heterogeneity affecting management efficiency and quality consistency. The class supports precision viticulture through spatial uniformity characterization enabling zone-based management.
geographic yield distribution
geographische Ertragsverteilung
innerbetriebliche Uniformität
räumliche Ertragsuniformität
räumliche Produktionshomogenität
spatial production homogeneity
within-vineyard uniformity
yield zone consistency
Spatial Yield Uniformity Quality
Specific gravity monitoring of Riesling fermentation in Mosel shows initial must at SG 1.085 (equivalent to 85°Oechsle) declining to SG 0.992 for dry wine fermented to completion, with the sub-unity final value reflecting ethanol density (0.789 g/mL) contribution exceeding residual extract, while fermentation arrest at SG 1.020 for Spätlese style indicates approximately 45 g/L residual sugar providing the characteristic sweetness balanced against Riesling acidity.
Specific gravity quality is a density quality that inheres in grape tissues or must and represents the dimensionless ratio of material density to the density of pure water at a specified reference temperature, providing a standardized expression of density that facilitates comparison across measurement systems and enables direct use in buoyancy-based sorting and concentration calculations. Within the BFO framework, this quality provides an alternative representation of the same physical property captured by absolute density, where the choice of specific gravity rather than density units reflects measurement methodology (hydrometry) and traditional practice in enological applications. The ontological significance lies in specific gravity enabling practical measurement through flotation and hydrometry methods that compare sample behavior to water references, eliminating the need for absolute mass and volume determinations that are more challenging in field and winery settings. Specific gravity of grape must ranges from approximately 1.050 for low-sugar early-harvest material to 1.150 for late-harvest botrytized must, with the deviation from unity reflecting the net contribution of dissolved solutes (primarily sugars increasing specific gravity) partially offset by ethanol if fermentation has commenced (decreasing specific gravity). Measurement using glass hydrometers remains the standard winery practice due to low cost and immediate readout, though digital density meters provide higher precision for laboratory applications. The temperature dependence of specific gravity requires correction to reference conditions (typically 20°C/20°C, meaning sample at 20°C compared to water at 20°C) using standard tables or instrument-specific algorithms. Understanding specific gravity supports density-based berry sorting where flotation in calibrated solutions separates berries by ripeness, fermentation monitoring through declining specific gravity as sugar converts to ethanol, and final wine analysis for residual sugar estimation.
Specific_Gravity_Quality provides the dimensionless expression of density, complementing absolute density measurements expressed in g/mL or kg/m³. This quality underlies hydrometric measurement methodology and traditional must weight scales. The class specifically addresses the water-referenced ratio rather than apparent specific gravity used for materials with entrapped air. Reference temperature specification is essential as specific gravity varies approximately 0.0002 per °C for must.
Dichteverhältnis
density ratio
hydrometer reading
relative Dichte
relative density
Amerine MA, Ough CS (1980) Methods for Analysis of Musts and Wines. John Wiley & Sons, New York, pp. 54-70 OIV (2021) Compendium of International Methods of Wine and Must Analysis, Volume 1, OIV-MA-AS2-01A
Specific Gravity Quality
Specific root length assessment comparing rootstocks reveals genetic differences in root construction strategy, with 3309C showing high SRL of 42 m/g (fine, efficient roots prioritizing absorption capacity per unit carbon), Schwarzmann showing moderate SRL of 28 m/g (balanced construction), and 1103P showing lower SRL of 18 m/g (thicker roots favoring persistence and transport), with the SRL variation corresponding to differences in fine root turnover (higher in 3309C) and total root biomass required to achieve equivalent absorbing surface, demonstrating how rootstock SRL affects the carbon economy of root system function.
Specific root length quality is a root uptake capacity quality that represents the root length per unit root dry mass (m/g), characterizing the efficiency of root construction that indicates the balance between root exploration (length for soil volume access) and root investment (biomass cost), with higher SRL indicating thinner, more efficient roots optimized for absorption while lower SRL indicates thicker roots optimized for transport, storage, or longevity. Within the BFO framework, this quality specifically addresses the efficiency dimension of root construction, distinguishing root length per mass from total root parameters while recognizing that SRL indicates root functional strategy, with high-SRL fine roots representing an acquisitive strategy maximizing absorption per carbon invested, while low-SRL coarse roots represent a conservative strategy emphasizing persistence and transport function. The ontological significance lies in specific root length revealing the root economic strategy that determines how effectively root system carbon investment translates into absorptive capacity, where high SRL (typically 20-50 m/g in grapevine fine roots) enables extensive soil exploration per unit root biomass, maximizing nutrient and water foraging efficiency, while the metabolic and structural costs of high-SRL roots (shorter lifespan, higher respiration rate per unit length) represent the trade-offs inherent in this acquisitive strategy. The functional implications of SRL variation include effects on nutrient uptake efficiency (high SRL enabling more surface area for a given carbon investment), water acquisition (high SRL exploring more soil volume for mobile water), and root system carbon cost (high SRL potentially increasing root turnover and associated carbon expenditure), with optimal SRL depending on soil resource availability and vine carbon status. The environmental plasticity of SRL shows roots responding to local conditions, with higher SRL in nutrient-poor patches (maximizing foraging efficiency), lower SRL in favorable conditions (reducing construction cost where resources are abundant), and variation with soil depth and texture reflecting the functional demands of different soil environments. The rootstock genotype affects SRL through genetic differences in root construction strategy, with some rootstocks producing inherently finer roots (higher SRL) providing greater absorptive efficiency, while others produce thicker roots (lower SRL) potentially favoring persistence and transport capacity. Understanding specific root length quality enables assessment of root construction efficiency, interpretation of root functional strategy, and evaluation of rootstock root economic characteristics affecting resource acquisition efficiency.
Specific_Root_Length_Quality addresses the efficiency of root construction indicating the balance between exploration capacity and construction cost. This quality reveals root economic strategy ranging from acquisitive (high SRL) to conservative (low SRL). The class supports rootstock characterization and root efficiency assessment through SRL evaluation.
SRL
Wurzelfeinheit
Wurzelkonstruktionseffizienz
root construction efficiency
root fineness
root length-mass ratio
spezifische Wurzellänge
Ostonen I et al. (2007) Specific root length as an indicator of environmental change. Plant Biosystems 141(3): 426-442 Comas LH et al. (2002) Linking fine root traits to maximum potential growth rate among 11 mature temperate tree species. Functional Ecology 16(3): 388-398
Specific Root Length Quality
The cultivar Gewürztraminer appears in databases and publications under spelling variant synonyms including Gewurztraminer (without umlaut, common in English-language sources), Gewuerztraminer (ae-substitution for ü), Gewürz-Traminer (hyphenated, used in some German regulatory documents), Gewuerstraminer (erroneous double substitution), and Гевюрцтраминер (Cyrillic transliteration) — all referring to the identical aromatic mutation of Savagnin Blanc (VIVC number 04222) without any semantic, regional, or historical distinction, the variation arising purely from orthographic convention differences across writing systems and style guides.
A synonym that differs from the prime name or another established synonym solely in orthographic representation — including variations in capitalization, hyphenation, diacritical marks, word spacing, transliteration conventions, or character encoding — while designating the identical cultivar without any semantic, linguistic, or regional distinction.
Spelling_Variant_Synonym addresses the orthographic noise in cultivar nomenclature generated by inconsistent handling of diacritical marks, compound word conventions, transliteration systems, and typographic standards across databases, publications, and regulatory documents. This category is disproportionately problematic for computational systems: database searches, automated matching algorithms, and data integration pipelines must account for spelling variants to avoid false non-matches. The VIVC and OIV databases attempt to standardize orthography but legacy data, national conventions, and character encoding limitations (particularly ASCII-only systems dropping diacritics) perpetuate variant proliferation. Spelling variants carry no taxonomic, geographic, or cultural information and exist purely as artifacts of writing system heterogeneity.
Schreibvariante
Schreibweise-Synonym
orthographic variant
orthographische Variante
spelling variation
typographic synonym
Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157 OIV (2013) OIV List of Vine Varieties and Their Synonyms ICNCP (2016) International Code of Nomenclature for Cultivated Plants 9th Ed, ISHS Art. 35
Spelling Variant Synonym
Spicy aroma profiling in Northern Rhône Syrah reveals the distinctive combination of grape-derived black pepper (7.5 on 10-point scale from rotundone at 85 ng/L) with oak-derived clove (5.2 from eugenol) and subtle cinnamon (3.8 from cinnamaldehyde), creating the complex spice layering that distinguishes Côte-Rôtie from Australian Shiraz where pepper typically predominates (8.2) over oak spices (clove 3.5) due to different barrel regimes, demonstrating how spice balance reveals both variety expression and winemaking approach.
Spicy aroma quality is an aroma character quality that represents olfactory perception evoking culinary spice-associated scents including pepper, clove, cinnamon, anise, and other warm or pungent spice notes, characterizing the complex, often warm aromatic impression that contributes to wine complexity and provides important markers for both grape variety and oak aging influence. Within the BFO framework, this quality addresses a major aroma category distinguished by its association with perceived warmth, complexity, and often savory character, recognizing that spicy notes in wine arise from diverse sources including grape-derived compounds (rotundone for pepper), oak-derived compounds (eugenol for clove), and fermentation products. The ontological significance lies in spicy aroma bridging varietal expression and winemaking influence, where pepper character may indicate Syrah or Grüner Veltliner genetics while clove and cinnamon notes typically reveal oak barrel aging, making spice descriptors informative about both raw material and production methods. The compositional basis for spicy aroma includes sesquiterpenes (rotundone providing black pepper with threshold of 16 ng/L in red wine), phenolic compounds from oak (eugenol for clove, cinnamaldehyde for cinnamon), volatile phenols (4-vinylguaiacol contributing spicy-smoky notes), and various esters and lactones adding sweet spice character. The perception of spicy aromas shows strong cross-modal interaction with trigeminal sensations, as many spice compounds also stimulate chemesthetic receptors creating the warming or pungent sensations associated with spices in food, enhancing the perceived spiciness of wines through non-olfactory pathways. The variety associations of spicy character vary by specific spice type, with pepper linked to Syrah, Grüner Veltliner, and Schioppettino, while sweet spices (cinnamon, clove, nutmeg) typically indicate oak aging across varieties. Understanding spicy aroma quality enables source identification (grape versus oak), variety assessment for pepper-associated varieties, and oak treatment evaluation based on sweet spice expression.
Spicy_Aroma_Quality serves as the parent class for spice subcategories (Pepper, Clove, Licorice), representing an aroma character category with diverse compositional origins spanning grape-derived, oak-derived, and fermentation-derived compounds. This quality bridges varietal assessment and winemaking evaluation, as different spice notes indicate different sources. The distinction between grape-derived spices (pepper) and oak-derived spices (clove, cinnamon) is particularly important for quality assessment.
Gewürzduft
Würzaroma
spice aroma
spicy character
warm spice notes
würziger Charakter
épicé scent
Siebert TE et al. (2008) Development of a quantitative method for the determination of rotundone, the pepper aroma compound, in wine and grapes. Journal of Agricultural and Food Chemistry 56(10): 3745-3748 Chatonnet P et al. (1999) Volatile and odourous compounds in barrel-aged wines. American Journal of Enology and Viticulture 50(4): 470-476
Spicy Aroma Quality
Investigation of Spider_Mite_Resistance_Disposition at UC Davis examined both direct resistance factors and predator-mediated resistance across 30 cultivars, comparing Tetranychus pacificus population development in the presence and absence of the predatory mite Galendromus occidentalis. In predator-exclusion assays on detached leaves, T. pacificus reproduction varied 2.5-fold among cultivars, from 28 eggs per female over 5 days on susceptible 'Thompson Seedless' to 11 eggs on relatively resistant 'Cabernet Sauvignon'. This direct resistance correlated moderately with leaf toughness (r = -0.52) but not with trichome density. In field trials with natural predator populations, cultivar rankings shifted substantially: 'Chardonnay', with moderate direct resistance (18 eggs in lab assays), supported the lowest field populations (3.2 mites/leaf at season peak) due to exceptionally well-developed leaf domatia at vein axils harboring high G. occidentalis densities (0.85 predators/leaf). Conversely, 'Carignane' with equivalent direct resistance supported 4-fold higher field populations (12.8 mites/leaf) associated with poorly developed domatia (0.12 predators/leaf). When predators were experimentally supplemented on 'Carignane', spider mite populations declined to 'Chardonnay' levels, confirming that Spider_Mite_Resistance_Disposition operates substantially through tritrophic interactions. These findings emphasize that breeding for spider mite resistance should consider domatia development as a key trait influencing biological control potential alongside direct resistance factors.
A Spider_Mite_Resistance_Disposition is a Mite_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that limit colonization, feeding, reproduction, or population buildup of tetranychid spider mites (Tetranychidae), web-spinning mites that damage grapevine foliage through cell puncturing and chlorophyll extraction causing characteristic stippling and bronzing. This disposition represents resistance to a functionally distinct mite group from eriophyids, with spider mites being larger (300-600 μm), oval-bodied, possessing four pairs of legs as adults, and often polyphagous rather than host-specialized. Major spider mite pests of grapevine include the Pacific spider mite (Tetranychus pacificus), two-spotted spider mite (T. urticae), Willamette spider mite (Eotetranychus willamettei), and European red mite (Panonychus ulmi), with species prevalence varying by region. Spider mites feed by inserting their cheliceral stylets into leaf mesophyll cells and extracting contents, causing chlorotic stippling that progresses to bronzing, leaf necrosis, and premature defoliation under heavy infestation, reducing photosynthetic capacity and delaying fruit ripening. Resistance mechanisms include leaf physical properties (trichome density and morphology, leaf toughness) affecting mite movement and feeding efficiency, constitutive or induced chemical defenses (terpenoids, phenolics) reducing mite survival or fecundity, and importantly, leaf domatia—small structures at vein axils that harbor predatory phytoseiid mites providing biological control. Spider mite resistance in grapevine is often mediated through tritrophic interactions involving predatory mites rather than direct antibiosis, making domatia-mediated resistance particularly significant.
Spider_Mite_Resistance_Disposition is_a Mite_Resistance_Disposition. Spider mites (Tetranychidae) differ fundamentally from eriophyid mites in biology, morphology, and host specificity. Resistance mechanisms include both direct effects on mite performance and indirect effects mediated by predatory mite populations influenced by leaf domatia. The tritrophic nature of spider mite resistance, involving cultivar effects on predator abundance and efficacy, distinguishes it from most other pest resistances and has important implications for integrated mite management emphasizing predator conservation.
Rote-Spinne-Abwehr
Spinnmilben-Resistenz-Disposition
Tetranychidae resistance
Tetranychiden-Resistenz
red spider mite resistance
web-spinning mite resistance
Karban R et al. (1995) Induced resistance in wild tobacco with clipped sagebrush neighbors, Ecology 76:2165-2169 English-Loeb GM et al. (2002) Tritrophic interactions and defense induction in grapevines, Induced Plant Resistance to Herbivory, Springer Duso C et al. (2010) Biological control of mites in European vineyards, Arthropod Management in Vineyards, Springer
Spider Mite Resistance Disposition
Splitting quality vulnerability in Kyoho table grapes necessitates intensive water management in Japanese greenhouse production, with precision irrigation maintaining soil moisture at 60-70% field capacity during ripening to minimize rapid water uptake events rain covers deployed over outdoor plantings reduce direct water absorption through berry skin, achieving splitting rates below 5% compared to 30-60% in unprotected vines after heavy rainfall, which enables the premium pricing (500-1000 yen per berry for perfect fruit) that drives Japanese luxury grape production economics.
Splitting_Quality represents the damage state characterized by rupture of the grape berry skin resulting from internal pressure exceeding skin tensile strength, creating cracks or fissures that expose the flesh. Berry splitting is a major quality defect that renders fruit unmarketable for table grape use and compromises wine grape quality through juice loss, oxidation, and decay initiation. Splitting occurs when water uptake into the berry generates turgor pressure that exceeds the mechanical strength of the skin, with rupture typically initiating at weak points including the stylar scar, the shoulder region, or sites of prior micro-damage. The susceptibility to splitting depends on skin mechanical properties (thickness, elasticity, strength), berry size and shape, cellular structure, and the rate and magnitude of water influx. Large-berried cultivars with thin skins are particularly susceptible, while small-berried thick-skinned varieties resist splitting. Rainfall during ripening is the primary environmental trigger, with water uptake through both roots and direct absorption through the berry surface contributing to splitting pressure. Splitting can also occur from irrigation following drought stress, from rapid temperature drops that affect skin elasticity, and from late-season conditions that promote continued berry expansion after skin growth has ceased. Split berries deteriorate rapidly as exposed flesh oxidizes, juice leaks, and decay organisms colonize the wound. The splitting percentage after rain events provides a cultivar susceptibility rating used in variety selection for humid climates. Assessment documents both the proportion of split berries and the severity of individual splits (partial crack versus complete rupture).
Splitting_Quality specifically addresses skin rupture from internal pressure, distinct from Mechanical_Damage_Quality where external forces cause injury. The splitting event is a process Splitting_Quality captures the resulting damage state. Splitting severity ranges from hairline cracks that may partially heal to complete berry rupture with major juice loss. Cultivar susceptibility to splitting is a disposition addressed in the Disposition category Splitting_Quality addresses the manifested damage state. The location and pattern of splits provides diagnostic information, with stylar-end splits suggesting different causation than shoulder splits or circumferential cracking.
Aufplatzen
Beerenplatzen
Beerenrissigkeit
Hautriss
berry burst quality
berry cracking
fruit splitting
skin rupture
Becker T and Knoche M (2012) Deposition, Strain, and Microcracking of the Cuticle in Developing 'Riesling' Grape Berries. Vitis 51(1): 1-6 Keller M et al. (2015) Interactive Effects of Deficit Irrigation and Crop Load on Cabernet Sauvignon in an Arid Climate. American Journal of Enology and Viticulture 66(1): 62-72
Splitting Quality
Sprawling tendency quality assessment in McLaren Vale compared cultivar growth patterns under minimal management to characterize natural canopy development behavior. Mourvèdre exhibited high sprawling tendency quality with shoot orientations distributed from 20° below to 70° above horizontal (SD 28°) creating dense, disorganized three-dimensional canopy requiring five positioning passes in VSP trials to achieve marginally acceptable organization, ultimately proving most productive under sprawling wire configuration where its natural tendency created self-regulating canopy with adequate if suboptimal fruit exposure. Tempranillo showed moderate sprawling tendency quality with more consistent semi-upright orientation (mean 52°, SD 18°) amenable to vertical positioning with standard intervention intensity. Sangiovese demonstrated minimal sprawling tendency quality with strongly consistent upright habit (mean 78°, SD 9°) naturally suited to VSP with minimal positioning requirement. The comparison established sprawling tendency quality as independent of vigor, with Mourvèdre and Sangiovese exhibiting similar total shoot growth but dramatically different canopy organization outcomes attributable to growth habit differences requiring different training approaches for optimal management.
Sprawling tendency quality is a growth habit compatibility quality that inheres in a grapevine cultivar or clone and represents the propensity for shoots to grow in multiple directions without consistent orientation preference, spreading laterally and occupying available space in a disorganized manner that resists confinement within structured canopy configurations and that may be either problematic for training systems requiring specific shoot placement or advantageous for minimal intervention systems designed to accommodate unmanaged canopy expansion. This quality captures the behavioral characteristic of cultivars that neither orient strongly upward nor trail consistently downward but instead exhibit variable, indeterminate growth direction that creates sprawling, three-dimensional canopy architectures when shoots are not physically constrained by trellis structures or positioning operations. The ontological significance of sprawling tendency quality lies in its recognition as a distinct growth habit category separate from upright and trailing types, with specific implications for training system selection and management intensity requirements, wherein high sprawling tendency may indicate poor suitability for geometrically-structured training systems while potentially favoring minimal pruning or sprawling wire configurations that accommodate rather than resist this natural growth pattern. The physiological basis of sprawling tendency involves intermediate or variable gravitropic response strength such that shoots do not consistently orient either toward or against gravity, high developmental plasticity permitting growth direction to respond to local light availability and mechanical obstruction, and potentially vigorous lateral shoot development that contributes to canopy volume expansion in all directions. Cultivars with strong sprawling tendency quality create diffuse, irregular canopies when grown without positioning constraints, exhibiting shoot orientations distributed across the full angular range rather than concentrated around specific preferred directions, and typically requiring substantial management input to achieve organized canopy architectures in structured training systems while potentially thriving under minimal intervention approaches. The assessment of sprawling tendency quality employs observation of natural canopy development in unpositioned conditions, quantification of shoot angle variance as an indicator of orientation consistency, and evaluation of management requirements across training systems to determine the threshold beyond which structured canopy organization becomes economically impractical.
Sprawling_Tendency_Quality is_a Growth_Habit_Compatibility_Quality, representing the propensity for inconsistent, multi-directional growth that resists organized canopy configuration. This class addresses cultivars that are neither strongly upright nor trailing but exhibit variable, spreading growth patterns. Distinguished from both Upright_Growth_Suitability_Quality and Trailing_Growth_Suitability_Quality by the absence of consistent directional tendency. The standard deviation of shoot angle provides quantitative metric for sprawling assessment. Relevant for decisions regarding minimal pruning suitability and the management intensity required to achieve structured canopy organization.
Ausbreitungstendenz
Spreizhaltung
canopy sprawl tendency
disorganized growth pattern
indeterminate shoot orientation
spreading growth habit
spreizende Wuchsneigung
ungerichtetes Wuchsverhalten
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 32-43 Clingeleffer PR (1993) Vine response to modified pruning practices. In: Pool RM (ed) Proceedings of the Second NJ Shaulis Grape Symposium, Cornell University, pp. 20-30 Intrieri C et al. (2011) Minimal pruning and cluster thinning for crop control in Sangiovese grapevines. American Journal of Enology and Viticulture 62:294-303
Sprawling Tendency Quality
Spray penetration quality assessment in Marlborough Sauvignon Blanc compared fungicide deposition on interior cluster surfaces across canopy density treatments using fluorescent tracer and UV photography for quantification. Dense canopies with leaf layer number 2.8 showed interior cluster coverage of only 18% relative to exterior canopy surfaces, requiring application of 650 L/ha to achieve minimum threshold coverage. Moderately managed canopies with leaf layer number 1.6 demonstrated improved spray penetration quality with interior coverage of 45%, achieving threshold protection at 400 L/ha application volume. Well-managed canopies with leaf layer number 1.1 exhibited optimal spray penetration quality with interior coverage reaching 72% of exterior deposition, enabling effective protection at only 300 L/ha. Season-long fungicide costs including materials and application were reduced by 38% in the well-managed treatment while achieving equivalent or superior disease control.
Spray penetration quality is a canopy porosity quality that inheres in a grapevine canopy system and represents the capacity for applied liquid materials including pesticides, foliar nutrients, and growth regulators to traverse the foliar mass and achieve adequate coverage of target surfaces throughout the canopy volume including interior leaves, developing clusters, and shoot tissues that may be obscured by overlapping foliage. This quality determines the biological efficacy of spray applications and the efficiency of plant protection programs, with dense canopies requiring higher application volumes, specialized equipment, or multiple spray directions to achieve coverage equivalent to that readily obtained in open canopies. The ontological significance of spray penetration quality lies in its role as a practical determinant of pest and disease management effectiveness and input efficiency, wherein poor spray penetration can undermine otherwise well-designed protection programs by leaving interior tissues unprotected despite apparently adequate application to canopy exteriors. The physics of spray penetration involves droplet transport through air against the drag and interception effects of foliage, with penetration success depending on droplet size distribution, spray velocity, canopy density, and the turbulence or air assistance provided by application equipment. Larger droplets possess greater momentum enabling deeper penetration but suffer higher interception rates by outer canopy layers, while smaller droplets penetrate more readily but may lack velocity to reach interior positions before evaporation or drift losses, creating optimization challenges that depend on canopy characteristics and product requirements. Dense canopies with multiple leaf layers and minimal gaps physically intercept spray droplets on outer foliage surfaces before they can reach interior targets, with research demonstrating that canopies with leaf layer numbers exceeding 2.0 may receive less than 30% of exterior deposition rates on interior surfaces using conventional application equipment. Management approaches to improve spray penetration quality include canopy management operations that increase porosity, adjustment of application parameters including pressure, volume, and droplet size, use of air-assisted sprayers that create turbulence improving droplet transport, and multi-directional application systems that access the canopy from multiple angles. The economic implications of spray penetration quality extend beyond product efficacy to include application efficiency, with dense canopies requiring higher volumes and multiple passes that increase labor, fuel, and material costs while potentially elevating environmental loading.
Spray_Penetration_Quality is_a Canopy_Porosity_Quality, representing the agrochemical application dimension of canopy openness. This class addresses the practical consequences of canopy density for plant protection operations. Distinguished from Light_Penetration_Quality and Air_Circulation_Quality as parallel consequences of canopy porosity, though spray penetration has distinct physical determinants including droplet dynamics and application equipment characteristics. Strong relationship with integrated pest management and sustainable viticulture objectives.
Applikationsabdeckung
Pflanzenschutzmitteldurchdringung
Spritzmittelpenetration
Sprühnebeleinbringung
application coverage
pesticide penetration
spray coverage quality
spray deposition quality
Pergher G, Petris R (2008) The effect of canopy density on spray deposition in vineyards. Journal of Agricultural Engineering 39:33-38 Wise JC et al. (2010) Tunnel sprayer airblast application effects on spray coverage and quality in vineyards. HortTechnology 20:564-568 Siegfried W et al. (2007) Dosage of plant protection products adapted to leaf area index in viticulture. Crop Protection 26:73-82
Spray Penetration Quality
The spray program specification for Pinot Noir in Champagne documents 12-15 fungicide applications between E-L 9 and E-L 35 — early season (E-L 9-15) applications of contact protectant copper hydroxide (1.5 kg Cu/ha) and wettable sulfur (6 kg/ha) at 10-12 day intervals targeting Phomopsis (Diaporthe ampelina) and early powdery mildew (Erysiphe necator), mid-season (E-L 17-27) applications alternating systemic DMI fungicides (penconazole 40 g/ha) for powdery mildew with CAA fungicides (mandipropamid 150 g/ha) for downy mildew (Plasmopara viticola) on a strict mode-of-action rotation schedule to prevent resistance development (maximum 3 consecutive applications of any single FRAC group), and pre-véraison (E-L 29-35) applications of anti-botrytis specialty products (fenhexamid 600 g/ha at BBCH 77 bunch closure and BBCH 81 véraison) — total copper usage limited to 4 kg Cu/ha/year, total synthetic fungicide active ingredient below 8 kg/ha/year, with all applications recorded in the mandatory phytosanitary register (cahier d'enregistrement) required by French regulation.
A pest management description that documents the complete schedule of pesticide applications planned or executed during a growing season, specifying for each application the target pest or disease, active ingredient and product formulation, application rate, spray volume, application method (airblast, tunnel, drone, or knapsack), phenological timing, pre-harvest interval, maximum annual applications, and resistance management strategy including mode-of-action rotation.
Spray_Program_Specification captures the most operationally detailed and regulatory-sensitive component of pest management documentation. The spray program is the single largest variable cost item in most vineyard budgets (2,000-5,000 EUR/ha/year in high-disease-pressure European regions) and the primary point of interaction between viticulture and environmental regulation: each application generates regulatory obligations including product registration verification, maximum residue limit (MRL) compliance, pre-harvest interval observance, spray drift management, operator safety requirements, and detailed record-keeping in the mandatory spray diary. Fungicide resistance management has become a critical dimension of spray program design — the emergence of QoI-resistant (strobilurin-resistant) Plasmopara viticola populations across European vineyards since 2007 and DMI-resistant Erysiphe necator populations in multiple regions demonstrates the consequences of inadequate mode-of-action rotation, with FRAC (Fungicide Resistance Action Committee) guidelines providing the framework for resistance-conscious spray program design.
Pflanzenschutzmittel-Programm
Spritzplan-Spezifikation
Sprühplan-Dokumentation
fungicide program specification
pesticide application record
spray schedule documentation
Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests 2nd Ed, APS Press FRAC (2023) FRAC Code List — fungal control agents sorted by cross-resistance pattern and mode of action Pertot I et al. (2017) A critical review of plant protection tools for reducing pesticide use on grapevine, Crop Protection 97: 70-84 EU Regulation 1107/2009 on plant protection products
Spray Program Specification
Spread history for Cabernet Sauvignon documents one of the most dramatic cultivar globalizations in viticultural history — from a localized southwestern French origin (probable 17th-century spontaneous cross) to near-universal global distribution within 200 years: introduction to Chile (1851, via Bordeaux vine imports commissioned by Silvestre Ochagavía), to South Australia (1844, James Busby collection via Kew Gardens), to Napa Valley (1880s, via multiple importations including Charles Krug and Gustave Niebaum), to Stellenbosch (1920s, via the KWV research program), to China (1980s-1990s, via government-sponsored modernization programs), with each introduction involving specific agents, routes, and motivations documented in archival records. Spread history for Blaufränkisch traces a more geographically circumscribed but culturally complex dispersal — from the Austrian-Hungarian border region outward through the Habsburg Empire under multiple synonyms, reaching Württemberg (as Lemberger, probably 18th century), Moravia and Bohemia (as Frankovka, 19th century), and eventually Washington State (as Lemberger, 1970s, via German immigrant winemakers), with contemporary premium cultivation reconcentrating in the Austrian Mittelburgenland, Leithaberg, and Eisenberg regions.
Spread history is a historical record entity that documents the temporal and spatial diffusion of grape cultivars from their center of origin or primary cultivation region to secondary, tertiary, and global viticultural destinations, integrating introduction records (the documented arrival of a cultivar in a new region), migration paths (the reconstructed routes and mechanisms of cultivar dispersal), and establishment dates (the chronological milestones of permanent cultivation in new territories), providing the biogeographic narrative of cultivar globalization that explains the contemporary worldwide distribution of Vitis vinifera cultivars as the cumulative result of millennia of human-mediated dispersal driven by colonial expansion, trade networks, monastic viticulture, scientific exchange, and deliberate varietal introduction programs.
Spread history documentation addresses the fundamental question of how the approximately 1,400 commercially cultivated grape varieties reached their current global distribution — a distribution shaped overwhelmingly by human agency rather than natural dispersal, as Vitis vinifera's fleshy berries are adapted for bird dispersal over short distances but not for the trans-oceanic and transcontinental movements that characterize viticultural globalization. The temporal framework of cultivar spread encompasses several distinct historical phases, each with characteristic dispersal mechanisms: ancient Mediterranean expansion (Phoenician, Greek, and Roman colonial viticulture, 1000 BCE-500 CE, spreading cultivars around the Mediterranean basin), medieval European consolidation (monastic viticulture, particularly Cistercian and Benedictine orders, 500-1500 CE, establishing the regional cultivar portfolios of European wine regions), early modern colonial expansion (Spanish, Portuguese, and later French, Dutch, and British colonial enterprises, 1500-1800, introducing European cultivars to the Americas, South Africa, and eventually Australia), and modern scientific exchange (systematic varietal introduction through agricultural research institutions, 1800-present, rationalizing global cultivar distribution). The phylloxera crisis (1860s-1900s) represents a critical discontinuity in spread history — the devastating destruction of European vineyards by the root-feeding aphid Dactylosphaera vitifoliae and the subsequent replanting on American rootstock fundamentally restructured the cultivar composition of many European regions, as replanting offered the opportunity (or necessity) to select different cultivars from those previously planted. The post-phylloxera restructuring, combined with government-sponsored varietal rationalization programs in the 20th century, means that the current cultivar distribution of many European regions dates effectively from the early 20th century rather than from medieval or ancient plantings, despite the much longer documented cultivation history of the individual cultivars involved.
Ausbreitungsgeschichte
Ausbreitungshistorie
Sortenverbreitung
Verbreitungsdokumentation
cultivar spread
diffusion history
distribution history
This P et al. (2006) Historical origins and genetic diversity of wine grapes. Trends in Genetics 22(9): 511-519 Unwin T (1991) Wine and the Vine: An Historical Geography of Viticulture and the Wine Trade, Routledge, pp.: 1-400 Pinney T (1989) A History of Wine in America: From the Beginnings to Prohibition, University of California Press, pp.: 1-50 Campbell C (2004) Phylloxera: How Wine was Saved for the World, HarperCollins, pp.: 1-50
Spread History
Cabernet Sauvignon mature vineyard showing spreading surface lateral roots, excavation revealing surface laterals extending 140-220 cm horizontal distance from trunk at 12-22 cm depth, spreading laterals 2.2-3.8 cm diameter showing vigorous growth, laterals extending into inter-row alleys accessing alley-applied fertilizer and irrigation, second-order branches every 10-18 cm creating extensive network, spreading surface lateral roots demonstrating extensive horizontal exploration accessing large vineyard floor soil volume
Spreading surface lateral root is surface lateral root emphasizing extensive horizontal spread and lateral exploration function, characterized by predominantly horizontal orientation maintaining shallow depth 8-25 cm while extending laterally 80-200+ cm, demonstrating rapid lateral elongation during favorable periods, creating extensive horizontal coverage exploring large soil surface area, and serving spatial exploration function accessing resources distributed heterogeneously across vineyard floor including inter-row zones potentially receiving differential fertilization or cover crop competition, with spreading surface laterals showing characteristics including vigorous lateral growth rates potentially 30-80 cm per year when conditions favorable, predominantly horizontal trajectory with minimal vertical deviation maintaining consistent shallow depth, moderate to substantial length 100-250+ cm in mature vines creating extensive reach, moderate diameter 1.5-4 cm supporting rapid elongation and resource transport, production of second-order and higher lateral branches along length creating hierarchical network exploring large soil volume, and demonstrated importance where spreading surface lateral extent largely determines vineyard floor soil volume explored by root system affecting competitive ability and resource capture. Spreading surface lateral root orientation and trajectory includes= showing predominantly horizontal growth (spreading laterals maintaining near-horizontal orientation 70-90° from vertical emphasizing lateral not vertical exploration), exhibiting depth consistency (spreading laterals maintaining relatively constant shallow depth 8-25 cm rather than ascending or descending), demonstrating directional persistence (spreading laterals showing relatively straight trajectories in preferred directions), revealing obstacle navigation (spreading laterals capable of adjusting trajectory navigating rocks, compacted zones, or other obstacles), and displaying radial pattern (spreading laterals extending in multiple directions creating roughly radial distribution from trunk). Spreading surface lateral root elongation dynamics includes= showing vigorous growth (spreading laterals elongating rapidly 30-80 cm per year during favorable periods), exhibiting seasonal patterns (spreading lateral elongation peaking spring and fall when soil conditions optimal), demonstrating environmental sensitivity (elongation rate strongly affected by moisture, temperature, nutrients), revealing age effects (young spreading laterals potentially showing faster elongation than old well-established laterals), and displaying rootstock influences (rootstock varieties differing in spreading lateral elongation vigor affecting exploration rate). Spreading surface lateral root extent and coverage includes= showing substantial reach (mature spreading surface laterals extending 100-250+ cm from trunk accessing large soil area), exhibiting inter-row penetration (spreading laterals often extending into inter-row alleys exploring spaces between vine rows), demonstrating overlapping zones (spreading laterals from neighboring vines potentially creating overlapping exploration zones in alleys), revealing asymmetric distribution (spreading lateral extent potentially greater in favorable directions or away from competition), and displaying vineyard floor coverage (spreading surface lateral network potentially exploring 60-90% of vineyard floor surface area in mature vineyards). Spreading surface lateral root branching architecture includes= showing second-order production (spreading surface laterals producing numerous second-order branches along length creating hierarchical structure), exhibiting branching spacing (second-order branches emerging every 8-20 cm along spreading lateral creating regular pattern), demonstrating progressive branching (branching potentially increasing as spreading lateral ages and extends), revealing hierarchy development (branching progressing from spreading lateral through multiple orders to terminal fine roots), and displaying exploration multiplication (branching greatly expanding total soil volume explored beyond what single unbranched lateral could access). Spreading surface lateral root resource capture includes= providing extensive exploration (spreading laterals accessing resources distributed across large vineyard floor area), exhibiting heterogeneity exploitation (spreading laterals encountering and exploiting patches of high nutrient or moisture availability), demonstrating competitive advantage (extensive spreading lateral reach potentially excluding competitors or accessing resources before competition), revealing fertilizer interception (spreading laterals positioned to intercept fertilizers applied in alleys or inter-row zones), and displaying irrigation capture (spreading laterals capturing irrigation water applied broadly across vineyard floor). Spreading surface lateral root in vineyard management includes= affecting alley management (spreading lateral extension into alleys influencing cover crop competition, fertilization, irrigation strategies), influencing traffic patterns (spreading lateral distribution potentially affected by repeated traffic in wheel tracks creating asymmetries), demonstrating tillage impacts (alley cultivation potentially damaging spreading laterals reducing exploration and uptake), revealing fertilizer placement (spreading lateral extent informing optimal fertilizer placement strategies for efficient capture), and displaying irrigation design (spreading lateral distribution affecting optimal irrigation system design and placement). Spreading surface lateral root competitive dynamics includes= showing inter-vine competition (spreading laterals from neighboring vines competing for overlapping soil resources in alleys), exhibiting cover crop interaction (spreading laterals competing with cover crop roots concentrated in same shallow topsoil zone), demonstrating spacing effects (vine row spacing affecting extent of spreading lateral competition or complementarity), revealing cultivar differences (cultivars potentially differing in spreading lateral vigor affecting competitive ability), and displaying rootstock influences (rootstock varieties showing different spreading lateral characteristics affecting competition outcomes). Spreading surface lateral root functional importance includes= determining exploration capacity (spreading lateral extent largely determining total soil volume explored by surface root system), enabling resource acquisition (spreading laterals accessing large resource pool supporting vine water and nutrient needs), demonstrating architectural foundation (spreading laterals creating spatial framework upon which higher-order absorptive network develops), revealing management sensitivity (spreading lateral function and distribution highly responsive to and affected by surface management practices), and displaying vine performance correlation (spreading lateral extent and health often correlating with overall vine vigor and productivity).
Spreading_Surface_Lateral_Root emphasizes extensive horizontal reach and exploration and is distinguished from other surface roots by focus on lateral spreading. Critical for understanding horizontal elongation, extensive coverage, inter-row penetration, resource capture, competitive dynamics, and the far-reaching laterals creating extensive vineyard floor exploration determining resource capture capacity.
Extensively spreading lateral
Far-reaching surface root
Horizontal explorer root
Horizontale Explorationswurzel
Weit-reichende flache Wurzel
Spreading Surface Lateral Root
Spring frost damage quality evaluation in Chablis following a severe April frost event (-7°C for 4 hours at bud swell to early shoot emergence stages) documented the catastrophic impact on Chardonnay production across the appellation. Vineyards at BBCH 07-09 (beginning of budburst) showed severe spring frost damage quality with 95% primary bud kill and significant secondary bud damage, with affected vines producing only 15% of normal yield from surviving tertiary buds and lateral shoots. Vineyards at BBCH 53 (inflorescence visible) showed complete spring frost damage quality with total primary and secondary crop loss, as the emerged inflorescences and shoots experienced lethal freezing with no recovery capacity, while tertiary buds produced only vegetative growth insufficient for meaningful crop. Later-developing parcels still at BBCH 00-05 (dormant to wool stage) showed moderate spring frost damage quality with primary bud survival of 40-60% due to the protection afforded by bud scales, with yield projections of 50-65% of normal from surviving primary shoots supplemented by secondary bud contribution. The spring frost damage quality assessment documented regional losses exceeding 50% of normal production, with the economic impact extending beyond the immediate crop loss to include recovery costs and the multi-year effects of depleted carbohydrate reserves on subsequent season productivity.
Spring frost damage quality is a frost damage severity quality that inheres in a grapevine organism and represents the extent of injury to developing shoots, leaves, and inflorescences from late spring freezing events occurring after budburst when actively growing tissues have lost dormant hardiness and are highly vulnerable to even brief exposure to temperatures below 0°C, with damage severity determining the proportion of current-season crop and canopy development that must be replaced through secondary bud activation or lateral shoot growth. This quality addresses the most economically significant frost damage type in many viticultural regions, recognizing that spring frost represents a primary production risk where the combination of phenological advancement and lingering frost probability creates a vulnerability window during which a single frost event can eliminate or severely reduce the current season's crop potential. The ontological significance of spring frost damage quality lies in its direct impact on current-season yield and quality, with spring frost damage potentially causing complete crop loss in severely affected vineyards while also affecting vine balance and carbohydrate reserves through the diversion of resources to replacement shoot development rather than fruit production. The biological basis of spring frost damage encompasses the extreme cold sensitivity of actively growing tissues that lack any significant freezing tolerance, with ice formation in young shoots, expanding leaves, and developing inflorescences causing rapid and complete tissue death at temperatures only slightly below 0°C, typically in the range of -2 to -4°C depending on tissue water content and the presence of ice-nucleating agents. The differential sensitivity among tissues creates characteristic damage patterns, with inflorescences often showing greater sensitivity than vegetative shoot tips due to their higher water content and lower capacity for supercooling, resulting in preferential loss of crop potential relative to vegetative growth capacity in marginal frost events. The recovery capacity following spring frost depends on the survival of secondary and tertiary buds within compound buds, the timing of frost relative to the season allowing adequate time for replacement shoot development, and the management practices employed to optimize the productivity of surviving or replacement shoots.
Spring_Frost_Damage_Quality is_a Frost_Damage_Severity_Quality, representing frost injury specifically to post-budburst actively growing tissues lacking cold hardiness. This class addresses the economically critical spring frost vulnerability window. BBCH stage at frost occurrence strongly determines damage severity and recovery capacity. Inflorescences show greater sensitivity than vegetative tissues. Secondary/tertiary bud survival determines recovery potential. The -2 to -4°C damage threshold for emerged tissues contrasts with much lower thresholds for dormant buds. Critical for cool-climate viticulture production risk assessment.
Austriebsfrostschaden
Frühjahrsfrostschaden
Spätfrostschaden
Wachstumsphasen-Frostverletzung
growth-stage frost injury
late frost damage
post-budburst frost injury
spring freeze damage
Trought MCT et al. (1999) Vineyard variability in Marlborough, New Zealand: characterising variation in vineyard performance and options for management. Australian Journal of Grape and Wine Research 5:84-89 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 508-545 Evans RG (2000) The art of protecting grapevines from low temperature injury. Proceedings of the ASEV 50th Anniversary Meeting, pp. 60-72
Spring Frost Damage Quality
Comprehensive evaluation of Spring_Frost_Tolerance_Disposition at INRAE Bordeaux combined phenological monitoring, controlled frost exposure, and recovery assessment across 25 cultivars over 5 seasons to characterize the multiple components of spring frost adaptation. Budbreak date (50% buds at green tip stage) was recorded annually, young shoots at matched developmental stages were exposed to controlled frost (-3°C for 2 hours), and survival and regrowth were assessed at 14 days post-exposure. Spring_Frost_Tolerance_Disposition showed cultivar variation through multiple mechanisms. 'Riesling' exhibited frost avoidance through late budbreak (April 15 mean, 12 days after 'Chasselas'), escaping 3 of 5 recorded frost events entirely. When exposed to frost, 'Riesling' young shoots showed moderate tissue tolerance with 45% primary shoot survival at -3°C. 'Merlot' broke bud early (April 5) but demonstrated strong recovery capacity, with secondary buds activating within 8 days of primary shoot frost kill and producing 0.7 clusters per secondary shoot. 'Chardonnay' combined early budbreak with poor recovery (secondary shoot fruitfulness only 0.2 clusters), representing low overall Spring_Frost_Tolerance_Disposition. Field records confirmed that cultivar damage patterns matched expectations from component traits, with late-breaking 'Riesling' showing lowest cumulative frost damage over the 5-year period despite only moderate tissue tolerance.
A Spring_Frost_Tolerance_Disposition is a Cold_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through mechanisms that enable survival of freezing temperatures occurring after budbreak when actively growing tissues have lost the cold hardiness characteristic of dormant tissues. This disposition represents cold tolerance during the vulnerable post-dormancy growth period, fundamentally distinct from midwinter hardiness in that actively growing tissues possess inherently limited freezing tolerance regardless of genotype, and protection depends primarily on frost avoidance through phenological timing or rapid recovery rather than tissue-level freeze tolerance. Spring frost damage is a major cause of crop loss in viticulture, with developing shoots and inflorescences killed at temperatures between -2°C and -4°C depending on growth stage, tissue hydration, and freezing duration. Spring_Frost_Tolerance_Disposition encompasses both avoidance mechanisms (late budbreak timing that reduces frost encounter probability) and tolerance/recovery mechanisms (tissue-level cold tolerance of young shoots, capacity to regenerate from secondary buds or basal shoots following frost damage). The relative importance of avoidance versus recovery depends on regional frost patterns, with late budbreak favored in climates with reliable late spring frosts and recovery capacity more relevant where frost timing is unpredictable. Spring_Frost_Tolerance_Disposition is partially independent of midwinter hardiness, as cultivars with excellent dormant cold tolerance may break bud early and suffer spring frost damage, while less winter-hardy cultivars with late budbreak may escape spring frost entirely.
Spring_Frost_Tolerance_Disposition is_a Cold_Tolerance_Disposition. Spring frost tolerance is fundamentally different from midwinter hardiness because actively growing tissues cannot achieve comparable cold tolerance to dormant tissues. Protection depends on avoidance (late budbreak), limited tissue tolerance, and recovery capacity. Child classes include Late_Budbreak_Frost_Avoidance_Disposition (phenological escape) and Shoot_Frost_Recovery_Disposition (regeneration after damage). Spring frost tolerance is partially independent of midwinter hardiness, representing a distinct component of overall cold adaptation.
Austriebsfrost-Toleranz
Frühjahrsfrost-Toleranz-Disposition
Spätfrost-Resistenz
late frost tolerance
post-budbreak frost tolerance
spring freeze resistance
Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235 Friend AP et al. (2011) Delayed winter pruning and spring frost damage to grapevine buds, Australian Journal of Grape and Wine Research 17:214-218 Trought MCT et al. (1999) Vineyard variability in Marlborough, New Zealand, American Journal of Enology and Viticulture 50:484-489
Spring Frost Tolerance Disposition
The spur length specification for Grenache on gobelet (bush vine) training in Châteauneuf-du-Pape prescribes two-bud spurs (coursons à deux yeux) on 5-6 spur positions arranged radially around the vine head, with each spur cut immediately above the second clearly visible bud at a diameter of 8-12 mm — the two-bud spur generates two shoots of which one is retained at the following winter's pruning as the new two-bud spur and the second is removed, maintaining the spur position close to the cordon or head and preventing the progressive elongation of spur arms (allongement des coursons) that would displace the fruiting zone away from the permanent vine structure.
A pruning method description that documents the number of buds retained on each spur in a spur-pruned training system, typically specifying two-bud spurs (the standard for most wine grape production) or one-bud spurs (used for crop reduction or specific training system requirements), along with the target spur diameter, position on the cordon, and spatial distribution that collectively define the fruiting unit configuration.
Spur_Length_Specification defines the basic fruiting unit in spur-pruned systems. The standard two-bud spur retains a basal bud (position 1) and a distal bud (position 2), both of which typically produce shoots — the distal shoot usually bears 1-2 clusters and provides the replacement spur position for the following year, while the basal shoot serves as an insurance shoot and future replacement option: this annual renewal cycle maintains spur positions close to the cordon while providing consistent bud fruitfulness. Longer spurs (3-4 buds) are occasionally used for cultivars with low basal fruitfulness that require retention of more distal buds, but longer spurs accelerate the displacement of fruiting positions away from the cordon and increase canopy congestion. The progressive growth of spur stubs (dead wood accumulation at spur bases) over the cordon's life is a structural concern that eventually degrades cordon health and sap flow efficiency, particularly in long-lived cordon systems maintained for 20+ years.
Zapfen-Augenzahl
Zapfenlänge-Spezifikation
Zapfenschnitt-Parameter
cordon spur parameters
spur bud count
spur size specification
Galet P (2000) General Viticulture, Oenoplurimedia pp: 240-255 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 100-115 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 340-350
Spur Length Specification
A spur short fruiting cane on Vitis vinifera cv. Riesling in Mosel Valley, Germany, retained at 2 nodes spanning 22cm from origin point on arched cane training system, selected from well-positioned shoot showing complete lignification and 8mm diameter, functionally equivalent to two-bud spur but classified as short cane due to training system architecture utilizing cane framework rather than permanent cordon, expected to produce 2 fruiting shoots (100% budbreak typical for mid-cane positions of Riesling) bearing 3-4 clusters totaling 0.9-1.2kg fruit given high fruitfulness of variety, contributing to overall vine crop of 3.5-4.5kg assembled from multiple such short canes distributed across arched training framework.
A short fruiting cane retained at length equivalent to standard spurs, typically 2-3 nodes, representing functional equivalence to spur pruning within nominally cane-pruned systems, characterized by very compact extent (20-35cm), minimal bud count providing baseline crop potential, positioning and function identical to spurs despite technically qualifying as short canes, and suitability for high-fruitfulness cultivars or quality-focused systems where even minimal bud retention provides adequate production. Spur short fruiting canes demonstrate the arbitrary nature of cane versus spur distinction at extreme low bud counts, with choice between terminology often reflecting training system architecture or regional tradition rather than meaningful biological differences, and functional outcomes (crop load, canopy density, fruit quality potential) essentially identical regardless of classification as abbreviated cane or elongated spur.
Spur short fruiting canes illustrate the semantic challenges in pruning terminology, where identical retention (2-3 buds) may be called spur in cordon-trained systems but short cane in cane-trained architectures, despite producing equivalent results. The functional difference between 2-node spurs and 2-node short canes is essentially zero - both provide two compound buds for fruiting shoot production, both occupy similar trellis space, both require equivalent management, and both support similar crops assuming equal fruitfulness. The cane versus spur terminology becomes meaningful only at longer retentions (4+ nodes clearly being canes, not spurs), with 2-3 node range representing ambiguous middle ground classified based on system context rather than inherent characteristics. This ambiguity rarely creates practical problems as growers understand their local terminology regardless of formal classification.
Zapfen-kurze Rute
minimal short cane
spur-equivalent cane
spurähnliche Kurzrute
two-node cane
Based on Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181.
Spur Short Fruiting Cane
The SSR allele sizes for Syrah at the VVS2 locus are 131 bp and 151 bp (heterozygous), determined by capillary electrophoresis on an ABI 3130xl Genetic Analyzer using the GeneScan 500 LIZ size standard — the 131 bp allele contains approximately 10 CT dinucleotide repeats while the 151 bp allele contains approximately 20 CT repeats, with the 20 bp size difference reflecting the gain or loss of 10 repeat units through replication slippage during the independent evolutionary histories of the two alleles. Allele sizes are reported as integer values rounded to the nearest expected repeat increment (2 bp for dinucleotide SSRs) and calibrated against reference DNA samples of known genotype (typically Pinot Noir and Chasselas) included in every electrophoresis run to ensure cross-run and cross-laboratory size calling consistency.
An SSR genotype component that records the measured size in base pairs of each allele detected at a specific SSR locus in a specific grapevine accession, with the diploid Vitis vinifera genome yielding either two different allele sizes (heterozygous state) or a single allele size (homozygous state) per locus, as determined by capillary electrophoresis or polyacrylamide gel electrophoresis of fluorescently labeled PCR amplification products.
SSR_Allele_Size is the atomic data unit of SSR genotyping — the measured fragment length that, combined with the locus identifier, constitutes a single allelic observation. The accuracy and reproducibility of allele size determination is critical for database cross-referencing because cultivar identification depends on exact allele size matching across laboratories and instruments: sources of allele sizing variability include the fluorescent dye label (different dyes produce different electrophoretic mobility shifts of 0.5-2.0 bp), the internal size standard used (GeneScan 500 LIZ, ROX, or equivalent), the electrophoresis platform (ABI 310, 3100, 3130xl, 3730xl each generating slightly different raw size estimates), and the allele calling algorithm (GeneMapper, Peak Scanner, or equivalent). The reference allele ladder system developed by This et al. (2004) addresses inter-laboratory variability by providing standard DNA samples of defined genotype that serve as calibration references — allele sizes in the target sample are reported relative to the reference alleles rather than as absolute fragment lengths, enabling consistent allele calling across platforms and laboratories.
Allellängen-Bestimmung
Mikrosatelliten-Fragmentlänge
SSR fragment size
SSR-Allelgröße
allele size determination
microsatellite allele length
This P et al. (2004) Development of a standard set of microsatellite reference alleles for identification of grape cultivars, Theoretical and Applied Genetics 109: 1448-1458 Sefc KM et al. (2001) Microsatellite markers for grapevine, in Roubelakis-Angelakis KA (ed) Molecular Biology and Biotechnology of the Grapevine, Kluwer pp: 433-463
SSR Allele Size
The SSR fingerprint issued for a certified Grüner Veltliner clone (Klosterneuburger Selektion 1) by the Austrian Federal Office for Viticulture documents the 9-locus OIV standard genotype — VVS2 (133:145), VVMD5 (226:236), VVMD7 (239:247), VVMD25 (239:249), VVMD27 (179:191), VVMD28 (233:259), VVMD32 (252:272), VrZAG62 (187:193), VrZAG79 (241:259) — verified against the VIVC reference profile for Grüner Veltliner (variety number 4799), confirmed identical at all 18 alleles, with the certification laboratory's analysis performed in duplicate on independently extracted DNA samples using the This et al. (2004) reference allele ladder for size calibration and Pinot Noir and Chasselas as positive controls. The SSR fingerprint accompanies the clone through the Austrian Bundesamt für Weinbau certification pipeline, providing molecular confirmation of cultivar identity at each propagation stage from mother vine through nursery multiplication to vineyard planting.
A genetic marker data record that assembles the validated, quality-controlled SSR genotype of a specific Vitis vinifera accession into a standardized molecular identity document, cross-referenced against international cultivar databases (VIVC, European Vitis Database) and calibrated against reference allele standards, serving as the definitive molecular certificate of cultivar identity for germplasm management, nursery certification, label integrity verification, and legal cultivar authentication.
SSR_Fingerprint elevates the SSR genotype from a research data point to an applied identity document with regulatory and commercial significance. The distinction between SSR_Genotype (a raw data class under Genotype_Data) and SSR_Fingerprint (a validated identity record under Genetic_Marker_Data) reflects the difference between generating genotype data and certifying cultivar identity — the fingerprint requires not only accurate genotyping but also verification against reference databases, calibration against standard allele ladders, quality-controlled laboratory procedures, and formal documentation that the tested accession matches (or does not match) a specific registered cultivar: this certification function has practical consequences in vineyard planting material verification (detecting mislabeled nursery stock, estimated at 3-8% error rates in some markets), wine label integrity programs (Australia's Label Integrity Program, South Africa's Wine of Origin scheme), legal disputes over varietal authenticity (particularly for high-value cultivars such as Pinot Noir and Nebbiolo where mislabeling carries significant commercial implications), and germplasm conservation (confirming the identity of accessions in national and international grapevine collections). The SSR fingerprint is increasingly supplemented by SNP-based identity verification for applications requiring clonal-level discrimination beyond the resolution of the standard 9-locus SSR panel.
Mikrosatelliten-Identitätszertifikat
SSR cultivar passport
SSR-Fingerabdruck
microsatellite identity certificate
molecular identity document
molekularer Sortenausweis
This P et al. (2004) Development of a standard set of microsatellite reference alleles for identification of grape cultivars, Theoretical and Applied Genetics 109: 1448-1458 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC), Vitis 54: 155-157 Regner F et al. (2006) Genetic variability and incidence of systemic diseases in the clonal selection of grapevines, Acta Horticulturae 706: 231-236
SSR Fingerprint
The SSR genotype of Pinot Noir at the 9 OIV-recommended standard loci as registered in the VIVC database reads VVS2 (131:151), VVMD5 (226:238), VVMD7 (239:243), VVMD25 (239:249), VVMD27 (181:189), VVMD28 (233:259), VVMD32 (252:272), VrZAG62 (187:193), VrZAG79 (245:251) — this 18-allele profile (two alleles per locus across 9 loci, reflecting the diploid heterozygous state) provides a theoretical discrimination power exceeding 1 in 10⁸ for Vitis vinifera cultivar identification, meaning the probability of two unrelated cultivars sharing this complete profile by chance is less than one in one hundred million, sufficient for unambiguous identification of any cultivar within the approximately 10,000 genetically distinct cultivars documented in global databases.
A genotype data record based on simple sequence repeat (SSR) markers — also termed microsatellites or short tandem repeats (STRs) — documenting the allele sizes at multiple codominant, highly polymorphic SSR loci that collectively provide a unique molecular fingerprint for cultivar identification, parentage analysis, and genetic diversity assessment in Vitis vinifera.
SSR_Genotype documents the molecular marker system that has served as the international standard for grapevine cultivar identification since the late 1990s. The establishment of a consensus set of 9 SSR loci by This et al. (2004) under OIV coordination — selected for high polymorphism (8-20 alleles per locus in V. vinifera), reliable amplification across Vitis species, and unambiguous scoring — created the first truly universal molecular identification system for grapevine, enabling cross-referencing of cultivar identities between collections, databases, and research groups worldwide: prior to SSR standardization, cultivar identification relied on ampelographic description (subjective morphological assessment) or institution-specific marker sets that could not be directly compared. The VIVC database (maintained by the Julius Kühn-Institut, Geilweilerhof) now contains SSR profiles for over 10,000 accessions referenced against the standard 9-locus set, and the European Vitis Database (maintained by multiple national repositories) provides complementary SSR data for germplasm collections across Europe. SSR genotyping has resolved thousands of synonymy cases (different names for the same genotype) and homonymy cases (same name for different genotypes) that plagued traditional ampelography.
Mikrosatelliten-Genotyp
SSR fingerprint
SSR-Genotyp
STR profile data
microsatellite genotype
This P et al. (2004) Development of a standard set of microsatellite reference alleles for identification of grape cultivars, Theoretical and Applied Genetics 109: 1448-1458 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC), Vitis 54: 155-157 Sefc KM et al. (2001) Microsatellite markers for grapevine: a state of the art, in Roubelakis-Angelakis KA (ed) Molecular Biology and Biotechnology of the Grapevine, Kluwer pp: 433-463
SSR Genotype
The SSR locus data for VVS2 specifies a dinucleotide (CT)n microsatellite located on chromosome 11 of the Vitis vinifera reference genome, originally isolated by Thomas and Scott (1993) from a Muscat Gordo Blanco genomic library, with forward primer 5'-CAGCCCGTAAATGTATCCATC-3' and reverse primer 5'-AAATTCAAAATTCTAATTCAACTGG-3' amplifying a fragment range of 123-165 bp in Vitis vinifera, with 22 alleles documented across 2,800+ cultivars in the European Vitis Database — VVS2 is the first locus in the standard OIV 9-locus identification panel (OIV descriptors 801-809) and provides an expected heterozygosity of 0.84 and a probability of identity of 0.04, contributing substantial discriminatory power to the multi-locus cultivar fingerprint.
An SSR genotype component that records the identifying information for a specific microsatellite locus used in grapevine genotyping, specifying the locus name (e.g., VVS2, VVMD5), chromosomal location, repeat motif type (dinucleotide, trinucleotide), primer sequences for PCR amplification, expected allele size range in Vitis vinifera, and the number of known alleles in the global cultivar pool, providing the reference framework within which allele size data are interpreted.
SSR_Locus_Data provides the reference specification for each microsatellite marker used in grapevine genotyping, enabling standardized data generation and cross-laboratory comparison. The 9 OIV standard loci were selected from over 100 published Vitis SSR markers based on stringent criteria including amplification reliability across diverse cultivars, clear allelic patterns with minimal stutter artifacts, high polymorphism information content (PIC > 0.7), and chromosomal distribution across different linkage groups to minimize linkage between loci: the resulting panel provides cumulative probability of identity (PI) values below 10⁻⁸ for unrelated cultivars and below 10⁻⁴ for full siblings, making it adequate for cultivar identification but potentially insufficient for distinguishing clones within a cultivar (which typically share identical SSR profiles at the standard 9 loci, requiring extended panels of 20-50+ loci or alternative marker types for intra-cultivar discrimination). The locus data includes the exact primer sequences required for PCR amplification, enabling any laboratory with standard molecular biology equipment to generate comparable genotype data — a critical feature for the system's universal applicability.
Mikrosatelliten-Lokus-Beschreibung
SSR-Locus-Daten
STR locus reference data
microsatellite locus description
This P et al. (2004) Development of a standard set of microsatellite reference alleles for identification of grape cultivars, Theoretical and Applied Genetics 109: 1448-1458 Thomas MR and Scott NS (1993) Microsatellite repeats in grapevine reveal DNA polymorphisms when analysed as sequence-tagged sites (STSs), Theoretical and Applied Genetics 86: 985-990 Bowers JE et al. (1996) Development and characterization of additional microsatellite DNA markers for grape, American Journal of Enology and Viticulture 47: 121-130
SSR Locus Data
The SSR profile of Tempranillo as registered in the VIVC database (variety number 12350) documents the complete 9-locus fingerprint — VVS2 (133:151), VVMD5 (228:232), VVMD7 (239:247), VVMD25 (239:249), VVMD27 (179:189), VVMD28 (243:263), VVMD32 (252:272), VrZAG62 (187:193), VrZAG79 (247:251) — this profile unambiguously identifies as genetically identical the accessions known under the synonyms Tempranillo (Spain), Tinta Roriz and Aragonez (Portugal), Valdepeñas (California), and Ull de Llebre (Catalonia), resolving a centuries-old nomenclatural confusion that prevented recognition of one of the world's most widely planted red wine cultivars as a single genotype grown under different names across the Iberian Peninsula and beyond. The same SSR profile excludes the erroneously attributed synonyms Tempranillo Blanco (a distinct somatic color mutant with a different VvMYBA1 genotype) and Cencibel Blanco (an unrelated white cultivar).
An SSR genotype record that assembles the complete set of allele size determinations across all analyzed SSR loci into a unified multi-locus genotype profile constituting the molecular fingerprint of a specific grapevine accession, with the standard 9-locus OIV profile providing the internationally recognized cultivar identifier registered in the VIVC database and used as the definitive reference for resolving synonymy, homonymy, and misidentification in global grapevine germplasm management.
SSR_Profile represents the operational unit of molecular cultivar identification — the complete multi-locus genotype that functions as the grapevine equivalent of a human DNA fingerprint. The profile's utility for cultivar identification rests on three properties: uniqueness (the probability that two unrelated V. vinifera cultivars share identical alleles at all 9 standard loci is less than 10⁻⁸), stability (the SSR profile is identical across all tissues and developmental stages of a given genotype, and is maintained unchanged through vegetative propagation over centuries), and universality (the same profile is generated regardless of which laboratory performs the analysis, provided standardized reference alleles are used for calibration). The SSR profile cannot, however, distinguish clones within a cultivar — clonal variants that differ in phenotypically important somatic mutations (e.g., Pinot Noir, Pinot Gris, and Pinot Blanc, which are color mutants of the same genotype) typically share identical 9-locus SSR profiles because the somatic mutations responsible for clonal variation occur at genomic positions not sampled by the standard SSR panel. Intra-cultivar clonal discrimination requires extended marker panels (50-100 SSR loci, SNP arrays, or whole-genome resequencing) capable of detecting the rare somatic mutations that accumulate during centuries of vegetative propagation.
Mikrosatelliten-Fingerabdruck
Multi-Locus-SSR-Genotyp
SSR-Profil
microsatellite cultivar identifier
multi-locus SSR profile
This P et al. (2004) Development of a standard set of microsatellite reference alleles for identification of grape cultivars, Theoretical and Applied Genetics 109: 1448-1458 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC), Vitis 54: 155-157 Ibáñez J et al. (2009) Genetic origin of the grapevine cultivar Tempranillo, American Journal of Enology and Viticulture 60: 549-553 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340
SSR Profile
In Chenin Blanc berries in Vouvray, Loire Valley during BBCH 85-89, stable tartaric acid level maintains per-berry tartrate content at 4.8-5.2 mg throughout 42 days of ripening, while volumetric concentration decreases from 10.5 to 7.2 g/L due to berry expansion from 0.92 to 1.48 mL, with potassium bitartrate precipitation accounting for 12% of total tartrate under moderate potassium accumulation of 38 mM at harvest.
Stable tartaric acid level describes the quantitative outcome of tartrate maintenance processes in Vitis vinifera berries, wherein total tartaric acid content per berry remains within a narrow range of plus-minus 10-15% of veraison values throughout the 35-50 day ripening period, resulting in harvest tartrate concentrations of 5-10 g/L that provide the primary acid backbone for wine production following the extensive catabolism of malic acid during the same developmental window. This stability manifests as consistent tartrate content when measured on a per-berry basis, with typical values of 3-6 mg tartaric acid per berry maintained from veraison through harvest despite the profound metabolic and compositional changes occurring in other berry components during ripening progression. The volumetric expression of stable tartaric acid levels requires interpretation accounting for berry expansion effects, as the 50-100% increase in berry volume during post-veraison growth produces apparent concentration decreases from 8-12 g/L at veraison to 5-8 g/L at harvest that do not represent actual tartrate loss but rather dilution within an expanding tissue volume. Potassium interactions modify the expression of stable tartaric acid levels through precipitation of potassium bitartrate crystals within vacuolar compartments, with the degree of precipitation dependent on berry potassium accumulation rates, temperature fluctuations affecting crystal solubility, and cultivar-specific differences in potassium uptake capacity, typically removing 10-25% of tartrate from the soluble pool while maintaining total berry tartrate content. The consistency of stable tartaric acid levels across diverse environmental conditions reflects the absence of temperature-responsive degradation pathways affecting tartrate metabolism, contrasting sharply with the 2-3 fold variation in malate degradation rates observed across the typical temperature range encountered during grape ripening seasons. Cultivar differences in stable tartaric acid levels primarily reflect genetic variation in pre-veraison tartrate biosynthesis capacity rather than ripening-phase metabolism, with high-acid varieties such as Riesling and Chenin Blanc typically maintaining higher tartrate levels of 7-10 g/L compared to lower-acid varieties such as Gewürztraminer showing levels of 4-6 g/L at equivalent maturity. The predictability of stable tartaric acid levels enables viticultural planning for harvest timing and wine style objectives, with tartrate measurements providing reliable baseline acidity values against which to assess malate reduction progress and optimal harvest windows for desired acid balance.
Stable_Tartaric_Acid_Level is subclass of Tartaric_Acid_Stabilization representing the quantitative measurement outcome. Distinguishes from Tartrate_Maintenance which describes physiological processes. Sibling to Tartrate_Maintenance and Acid_Balance_Achievement. Key parameter is plus-minus 10-15% variation in per-berry tartrate content. Important for harvest planning and wine style determination.
beständiger Weinsäure-Spiegel
consistent tartaric acid
constant tartrate concentration
konstanter Tartrat-Gehalt
maintained tartrate level
stabile Weinsäure-Konzentration
stabiler Weinsäure-Level
stable tartaric acid content
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 262-268 Conde C et al (2007) Biochemical changes throughout grape berry development and ripening. Food Chemistry 1: 1-22 Ruffner HP (1982) Metabolism of tartaric and malic acids in Vitis. Vitis 21: 346-358
Stable Tartaric Acid Level
Stage duration variance assessment for the flowering-to-véraison interval in Pinot Noir reveals the fruit load effect on developmental rate heterogeneity, with balanced crop load (8 kg/vine) showing low duration variance (flowering-to-véraison interval 52-58 days across clusters, CV 5.2%, uniform development rate), while heavy crop load (14 kg/vine) shows higher duration variance (interval 48-72 days, CV 14.8%, resource competition creating divergent development rates among clusters), demonstrating how stage duration variance increases under resource limitation with cumulative effects on harvest-time maturity spread.
Stage duration variance quality is a temporal spread quality that represents the variation in the length of time different organs or plants spend in specific developmental stages, characterizing the heterogeneity of phase durations that affects the alignment of developmental timing even when stage onset may be synchronized, as different duration leads to divergent completion timing and potentially different quality outcomes from varied time spent in each developmental phase. Within the BFO framework, this quality specifically addresses the duration dimension of temporal spread, distinguishing variation in how long stages last from variation in when stages begin while recognizing that duration variance creates a distinct source of developmental heterogeneity—even perfectly synchronized stage onset will produce asynchronous completion if duration varies among individuals. The ontological significance lies in stage duration variance affecting the cumulative timing divergence through the season and the quality implications of different exposure durations to stage-specific conditions, where varying time spent in flowering affects fruit set opportunity, varying time in green berry stage affects cell division completion, and varying ripening duration affects the temperature integration and metabolic time determining final fruit composition. The factors affecting stage duration variance include microclimate variation (temperature differences affecting developmental rate and thus duration), resource availability variation (differential supply affecting growth rates), sink competition variation (different fruit loads affecting per-berry resource allocation and development rate), and genetic variation (different clones or cultivars showing inherently different phase durations), with these factors creating the duration heterogeneity that complements onset timing variation in determining overall developmental spread. The measurement of stage duration variance requires tracking not only when stages begin but when they end, enabling calculation of individual durations and their population variance, with higher variance indicating greater heterogeneity in developmental trajectories even if onset was relatively synchronized. Understanding stage duration variance quality enables separation of onset synchrony from duration effects on overall developmental spread, identification of stages where duration variance most affects quality outcomes, and management targeting factors that reduce duration variance for improved developmental uniformity.
Stage_Duration_Variance_Quality addresses variation in the time spent within developmental stages affecting completion timing even from synchronized onset. This quality captures rate-related heterogeneity complementing onset timing variation. The class supports developmental uniformity assessment through duration variance characterization.
Entwicklungszeitvariation
Phasenlängenvariation
Stadiendauerheterogenität
Stadiendauervarianz
development time variance
developmental rate variation
phase length variation
stage duration heterogeneity
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 277-316 Parker AK et al. (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17(2): 206-216
Stage Duration Variance Quality
Stage identification clarity assessment across the BBCH flowering sequence in Pinot Noir reveals the variable recognizability affecting observation reliability, with BBCH 60 (first flowers open) showing moderate clarity (requires close inspection to detect first open flowers among closed buds), BBCH 65 (full bloom, 50% capfall) showing high clarity (readily quantifiable by visual assessment of cap presence), and BBCH 68 (80% of flowerhoods fallen) showing high clarity (clear visual dominance of open flowers with few remaining caps), demonstrating how identification clarity varies even within a single principal growth stage with implications for observer training emphasizing stages with lower inherent clarity.
Stage identification clarity quality is a developmental stage quality that represents the ease and unambiguity with which specific developmental stages can be recognized in the vineyard, characterizing the observational clarity of stage-defining characteristics that determines how reliably different observers can identify the same developmental stage and how confidently individual assessments can be made under field conditions. Within the BFO framework, this quality specifically addresses the observational dimension of stage recognition, distinguishing the clarity of visual or tactile markers from the precision of stage coding while recognizing that identification clarity varies substantially among stages based on the distinctiveness of defining characteristics, with some stages presenting obvious markers and others requiring experienced interpretation of subtle developmental changes. The ontological significance lies in stage identification clarity affecting the practical utility of developmental staging for management and research, where stages with clear identification enable confident timing of stage-dependent operations and reliable data collection, while stages with ambiguous markers create uncertainty that may require conservative timing decisions or multiple assessments to establish developmental status. The characteristics providing high identification clarity include discrete morphological changes (cap fall during flowering providing unambiguous marker), color transitions (veraison color change clearly visible), and quantitative thresholds (percentage of flowers open defining bloom stages), while lower clarity arises from gradual transitions (bud swelling to bud burst), subjective assessments (berry softening), and variable expression (chlorosis patterns during senescence). The cultivar effects on identification clarity can be substantial, as some cultivars express stage markers more distinctly than others, with for example clear color change at veraison in pigmented cultivars but subtle softening-only markers in white cultivars, affecting the reliability of visual staging across the cultivar range. The environmental effects on identification clarity include lighting conditions (affecting color perception), weather damage (confounding developmental markers with stress symptoms), and canopy density (affecting visibility of interior clusters), with standardized assessment protocols addressing these sources of identification variation. Understanding stage identification clarity quality enables appropriate confidence assignment to stage determinations, development of cultivar-specific identification guides, and recognition of stages requiring extra care or multiple observations for reliable assessment.
Stage_Identification_Clarity_Quality addresses the observational ease and unambiguity of recognizing specific developmental stages under field conditions. This quality varies among stages based on distinctiveness of defining characteristics. The class supports observer training and assessment protocol development through clarity characterization across developmental stages.
Erkennbarkeit
Identifikationsleichtigkeit
Markerklarheit
Stadienidentifikationsklarheit
identification ease
marker clarity
observational distinctiveness
stage recognizability
Coombe BG (1995) Adoption of a system for identifying grapevine growth stages. Australian Journal of Grape and Wine Research 1(2): 104-110 Dry PR, Coombe BG (2004) Viticulture Volume 1: Resources, 2nd Ed. Winetitles, Adelaide, pp. 150-166
Stage Identification Clarity Quality
Stage transition duration assessment comparing temperature effects on flowering in Sauvignon Blanc reveals the environmental control of critical phenological windows, with cool conditions (mean 16°C during flowering) extending transition duration to 14 days (prolonged pollination window but extended Botrytis infection risk), while warm conditions (mean 23°C) compress duration to 6 days (brief but intense flowering, reduced disease exposure but tight spray timing), demonstrating how transition duration responds to temperature with implications for both process completion and management scheduling during this critical phenological window.
Stage transition duration quality is a stage transition quality that represents the time required for completion of the transition from one developmental stage to the next, characterizing the temporal extent of the transitional period during which the vine progresses from full expression of stage X to full expression of stage X+1, with duration affecting the phenological window for stage-associated processes and management operations. Within the BFO framework, this quality specifically addresses the temporal extent dimension of stage transitions, distinguishing how long transitions take from how sharp their boundaries appear while recognizing that duration determines the time available for transitional processes (e.g., pollination during the flowering transition) and the scheduling flexibility for stage-dependent management interventions. The ontological significance lies in stage transition duration determining the phenological windows during which critical processes occur and stage-dependent management must be accomplished, where brief transitions compress critical periods potentially creating management bottlenecks, while extended transitions provide more flexibility but may also prolong exposure to stage-specific vulnerabilities (disease susceptibility, weather risk). The factors affecting transition duration include temperature (higher temperatures generally shortening duration through accelerated development), cultivar genetics (inherent differences in transition tempo), and the nature of the specific transition (some transitions inherently brief like cap fall, others inherently extended like ripening), with these factors interacting to determine actual transition duration in any given situation. The stage-specific patterns of transition duration show substantial variation, with flowering typically brief (5-15 days from first flower to completion of fruit set), veraison of moderate duration (7-20 days for complete color change in a cluster, longer at whole-vine scale), and ripening extended (35-55 days from veraison to harvest maturity), with each duration having implications for management timing and process completion. The management implications of transition duration include spray timing during flowering (brief window requiring timely application), harvest planning relative to veraison completion (duration affecting prediction accuracy), and overall seasonal planning (cumulative transition durations contributing to total season length). Understanding stage transition duration quality enables realistic scheduling of stage-dependent operations, prediction of phenological windows for critical processes, and assessment of environmental effects on developmental timing.
Stage_Transition_Duration_Quality addresses the temporal extent of transitions between developmental stages. This quality determines phenological window length for critical processes and management operations. The class supports operational planning through transition duration characterization and prediction.
Stadienprogression
phenological duration
phänologische Dauer
stage progression time
transition length
transition period
Übergangsdauer
Übergangsperiode
Petrie PR, Sadras VO (2008) Advancement of grapevine maturity in Australia between 1993 and 2006: putative causes, magnitude of trends and viticultural consequences. Australian Journal of Grape and Wine Research 14(1): 33-45 Parker AK et al. (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17(2): 206-216
Stage Transition Duration Quality
Stage transition quality assessment comparing cultivars at veraison reveals the characteristic transition patterns affecting harvest planning, with Pinot Noir showing sharp veraison transition (90% of berries completing color change within 8 days, enabling single-date harvest targeting optimal maturity), while Grenache shows gradual transition (only 70% completion at 8 days, full color requiring 18 days, creating extended "red and green" period complicating harvest timing), demonstrating how stage transition quality varies among cultivars with practical implications for vineyard management and harvest decisions.
Stage transition quality is a phenological quality that represents the characteristics of progression between consecutive developmental stages, characterizing how distinctly and rapidly vines move from one phenological stage to the next, with transitions varying from sharp and synchronous (clear demarcation between stages occurring uniformly across the vine) to gradual and asynchronous (extended overlap between stages with different organs at different developmental points). Within the BFO framework, this quality addresses the boundary dimension of phenological progression, distinguishing the transition between stages from the stages themselves or the overall rate of development while recognizing that transition characteristics affect the precision of phenological staging, the uniformity of developmental processes, and the practical timing of stage-dependent management operations. The ontological significance lies in stage transition quality determining the temporal precision with which phenological events can be defined and managed, where sharp transitions enable confident identification of stage completion and precise timing of subsequent operations, while gradual transitions create extended periods of mixed developmental status that complicate both phenological assessment and management timing decisions. The factors affecting stage transition quality include genetic uniformity (clonal vineyards showing more synchronous transitions than mixed plantings), environmental uniformity (consistent soil and microclimate promoting synchronous development), vine condition (balanced vines transitioning more uniformly than stressed or excessively vigorous vines), and the inherent nature of specific transitions (some stages having naturally sharper boundaries than others). The cultivar-specific patterns of stage transition show some cultivars characteristically exhibiting sharp transitions (Chardonnay often showing compressed, synchronous veraison) while others show inherently gradual transitions (Grenache often exhibiting extended, asynchronous veraison with "red and green" berry populations persisting longer). The management implications of stage transition quality include the scheduling of stage-dependent operations (sharp transitions enabling precise single interventions, gradual transitions potentially requiring multiple passes), harvest timing decisions (uniform transitions simplifying optimal harvest determination), and quality assessment (transition characteristics affecting the homogeneity of fruit reaching harvest). Understanding stage transition quality enables appropriate timing of phenological observations, realistic expectations for stage-dependent management precision, and interpretation of transition variation as normal cultivar behavior versus stress indicators.
Stage_Transition_Quality serves as the parent class for transition characteristics (Sharpness, Duration). This quality addresses the boundary between consecutive developmental stages affecting phenological precision. The class supports management timing and phenological assessment through transition characterization.
Entwicklungsübergang
Stadienwechsel
Stadienübergangsqualität
developmental transition
phenological transition
phänologischer Übergang
stage boundary
stage change
Coombe BG, McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6(2): 131-135 Sadras VO, Petrie PR (2011) Climate shifts in south-eastern Australia: early maturity of Chardonnay, Shiraz and Cabernet Sauvignon is associated with early onset rather than faster ripening. Australian Journal of Grape and Wine Research 17(2): 199-205
Stage Transition Quality
Stage transition sharpness assessment comparing different phenological events in Merlot reveals the inherent variation in boundary clarity, with flowering onset (BBCH 60, first flowers open) showing high sharpness (clearly datable event, single-day precision achievable), veraison onset (BBCH 81, beginning of color change) showing moderate sharpness (first colored berries identifiable but representing small population fraction, 3-5 day uncertainty typical), and berry softening (BBCH 85) showing low sharpness (gradual textural change difficult to precisely date, 7-10 day uncertainty), demonstrating how transition sharpness varies among phenological events affecting the achievable precision of phenological recording.
Stage transition sharpness quality is a stage transition quality that represents the distinctness of the boundary between consecutive developmental stages, characterizing how clearly the end of one stage and beginning of the next can be identified, with sharp transitions showing clear demarcation (unambiguous stage change observable on specific dates) and gradual transitions showing extended overlap (prolonged coexistence of old and new stage characteristics). Within the BFO framework, this quality specifically addresses the boundary definition dimension of stage transitions, distinguishing how discrete or diffuse the transition appears from the duration of the transition while recognizing that sharpness affects the precision with which phenological events can be dated and the confidence with which stage assignments can be made during transitional periods. The ontological significance lies in transition sharpness determining the precision of phenological timing and the reliability of stage-dependent management decisions, where sharp transitions enable definitive staging (vine is clearly at stage X on date Y) while gradual transitions create staging ambiguity (vine shows characteristics of both stages X and X+1 over extended period) that affects both data quality and management precision. The biological basis for transition sharpness includes the nature of the developmental change (some changes inherently discrete like cap fall, others inherently gradual like color change), the synchrony of organ populations (synchronized development producing sharper whole-vine transitions), and the rate of the underlying process (rapid processes appearing sharper than slow ones even if equally deterministic). The stage-specific patterns of transition sharpness show substantial variation, with some transitions inherently sharp (beginning of flowering marked by first open flower, cap fall during bloom producing clear stage markers) and others inherently gradual (veraison onset showing progressive color change across berry population, softening difficult to precisely date). The environmental effects on transition sharpness include temperature uniformity (consistent temperatures promoting synchronous development and sharper transitions), stress events (heat or water stress potentially desynchronizing development and blurring transitions), and spatial uniformity (consistent vineyard conditions promoting uniform, sharp transitions across the block). Understanding stage transition sharpness quality enables appropriate confidence assignment to phenological dates, realistic expectations for staging precision at different phenological phases, and interpretation of transition characteristics as indicators of developmental synchrony and vine condition.
Stage_Transition_Sharpness_Quality addresses the distinctness of boundaries between developmental stages affecting dating precision. This quality varies among phenological events and is influenced by developmental synchrony. The class supports phenological data quality assessment through transition sharpness characterization.
Stadienabgrenzung
boundary sharpness
phenological discreteness
phänologische Diskretheit
stage demarcation
transition clarity
Übergangklarheit
Übergangschärfe
Stage Transition Sharpness Quality
Chardonnay hermaphrodite flowers at BBCH 65 (full bloom) exhibit five erect stamens with filaments 3-3.5 mm extended above the pistil (2-2.3 mm), anthers 0.7-0.8 mm with longitudinal dehiscence slits releasing abundant yellow pollen (18,000-24,000 grains per flower, 72-80% viability), creating efficient self-pollination geometry and achieving 48-55% fruit set under favorable conditions. Gewürztraminer functionally female flowers show five strongly reflex stamens with filaments 1.8-2.5 mm curved sharply downward positioning anthers 1.2-1.8 mm below the prominent pistil (2.3-2.6 mm), producing minimal pollen (<3,000 grains per flower, <15% viability), requiring cross-pollination from hermaphrodite pollinizers to achieve 28-38% fruit set. Cabernet Sauvignon experiencing heat stress (38-42°C) during microsporogenesis at BBCH 55-56 shows apparently normal stamen morphology at anthesis but severely reduced pollen viability (35-48% versus normal 70-80%), resulting in decreased fruit set (32-40% versus normal 50-58%) despite adequate stamen development and dehiscence, demonstrating temperature sensitivity of pollen development within otherwise normal stamens.
A stamen is the male reproductive organ of the Vitis flower comprising a filament (stalk) and an anther (pollen-producing structure), with typically five stamens arranged in a whorl opposite the five petals in the characteristic Vitaceae floral organization, functioning to produce, mature, and disperse pollen grains containing the male gametophytes essential for sexual reproduction through fertilization of ovules in the pistil. The stamen structure consists of= (1) a filament ranging from 0.5-4 mm in length depending on flower type and developmental stage, functioning as a stalk elevating the anther to an appropriate position for pollen dispersal (2) an anther comprising two lobes (thecae) each containing two pollen sacs (microsporangia), with total anther dimensions typically 0.4-1 mm length and 0.3-0.6 mm width, functioning to produce pollen through microsporogenesis and release pollen through dehiscence (longitudinal splitting) (3) a vascular strand running through the filament connecting to receptacle vasculature, providing nutrient and water transport to support anther development and pollen maturation. Stamen development proceeds through distinct phases= (1) primordium initiation during early flower bud development (BBCH 51-53) as five protuberances emerging from the receptacle in positions opposite the petal primordia (2) differentiation into filament and anther regions (BBCH 53-55) with anther primordia forming two lobes and four pollen sacs (3) microsporogenesis within anther pollen sacs (BBCH 55-57) involving meiotic divisions of pollen mother cells to produce tetrads of haploid microspores (4) pollen maturation (BBCH 57-59) involving microspore development into mature pollen grains with characteristic Vitis pollen morphology (tricolporate, 25-40 μm diameter, reticulate exine sculpture) (5) filament elongation and anther maturation (BBCH 59-60) preparing for anthesis (6) rapid filament elongation at anthesis (BBCH 60-61) lifting the calyptra and positioning anthers for dehiscence (7) anther dehiscence (BBCH 61-65) releasing pollen through longitudinal slits (8) post-dehiscence senescence (BBCH 65-69) with filament and anther browning and eventual abscission. Stamen morphology and function vary critically among Vitis flower types= hermaphrodite flowers exhibit erect stamens with filaments extending 0.5-1.5 mm beyond the pistil, anthers 0.6-0.9 mm producing 15,000-30,000 viable pollen grains per flower (70-85% viability), facilitating efficient self-pollination functionally female flowers show strongly reflex stamens curved downward positioning anthers 1-2.5 mm below pistil, with reduced pollen production (35°C during microsporogenesis (BBCH 55-57) can cause pollen abortion reducing viability by 40-70% cold stress <10°C during pollen maturation (BBCH 57-59) can reduce pollen viability by 30-50% water stress during critical developmental windows can reduce pollen quantity and quality, affecting subsequent fruit set.
Stamen represents the complete male reproductive organ and must be distinguished from its component parts (Anther, Filament, Pollen_Sac) which are sub-structures. The stamen is the fundamental unit of the androecium (male flower whorl), with five stamens collectively constituting the complete male reproductive system in Vitis flowers. Stamen morphology, particularly the erect versus reflex filament orientation and the pollen production capacity of anthers, determines flower sexual function (hermaphrodite versus functionally female versus staminate versus pistillate) and consequently reproductive strategy and crop production potential. Sub-classification addresses the major stamen components (Anther, Filament, Pollen_Sac) with their respective developmental and functional variants. Understanding stamen development and function is essential for viticulture applications including bloom timing prediction, fruit set optimization, controlled crossing in breeding programs, and diagnosis of reproductive failures from environmental stresses.
Staubblatt
Staubgefäß
androecium component
male organ
männliches Blütenorgan
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 99-108 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 92-106 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 185-198 Ebadi, A. et al. (1996). Effect of short-term temperature and shading on fruit set, seed and berry development in model vines. Australian Journal of Grape and Wine Research 2=2-9.
Stamen
The stamen description of Pinot Noir documents five erect stamens with filaments of 2.0-2.5 mm length (approximately equal to the ovary height, OIV 154:3 ratio of filament length to pistil length), bearing yellow bilobed anthers of 0.8-1.0 mm length that dehisce longitudinally at anthesis to release approximately 1,000-3,000 pollen grains per anther (5,000-15,000 per flower), with pollen viability exceeding 90% as assessed by in vitro germination on 15% sucrose agar medium — this fully functional androecium enables effective self-pollination as pollen released during calyptra abscission falls directly onto the exposed stigma, with supplementary wind-mediated cross-pollination from neighboring vines contributing genetic mixing within the vineyard block. The erect stamen posture and high pollen viability distinguish the hermaphrodite Pinot Noir flower from the reflexed stamens with aborted pollen (<5% viable) of functionally female cultivars.
A flower description component that records the morphological characteristics of the stamens — the male reproductive organs consisting of filament and anther — documenting stamen number (typically five in Vitis), filament length relative to the ovary, anther size and shape, pollen viability, and the posture of stamens (erect in hermaphrodite and male flowers, reflexed in functionally female flowers), with stamen morphology providing the primary visual distinction between functional flower types and contributing to the assessment of pollen production capacity and self-pollination potential.
Stamen_Description documents the male reproductive organs that determine pollen production capacity and consequently influence fruit set efficiency, self-pollination success, and the degree of outcrossing within vineyard populations. The five stamens of the Vitis flower are positioned alternating with the five calyptra lobes, and their development follows a precise temporal program — filament elongation occurs within the closed calyptra during the 2-3 days before anthesis, with the filaments coiled under spring tension that straightens them to the erect position when the calyptra detaches, simultaneously exposing the anthers to air that triggers dehydration-driven longitudinal dehiscence and pollen release: this coordinated mechanical system ensures that pollen is released precisely at the moment of stigma exposure, maximizing self-pollination opportunity. The ratio of filament length to pistil length (OIV 154) is diagnostically important — hermaphrodite flowers show filaments approximately equal to or longer than the pistil (ensuring pollen delivery to the stigmatic surface), while functionally female flowers show filaments shorter than the pistil and reflexed away from the stigma (preventing the deposition of non-viable pollen on the receptive surface).
Androeceum-Charakterisierung
Staubblatt-Beschreibung
Staubgefäß-Morphologie
androecium description
male organ characterization
stamen morphology record
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition May P (2004) Flowering and Fruitset in Grapevines, Lythrum Press Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia
Stamen Description
Wild Vitis vinifera ssp. sylvestris male plants in riparian forests of southern France, Italy, and the Balkans produce exclusively staminate flowers at BBCH 65 with robust stamens bearing anthers 0.7-0.9 mm long, producing 30,000-42,000 pollen grains per flower with 78-88% viability, and pistil completely absent or reduced to a minute vestige <0.3 mm long lacking any recognizable stigma, achieving <0.1% fruit set and functioning purely as pollen donors that fertilize nearby pistillate (female) plants through wind pollination. Occasional staminate reversions appear in cultivated Pinot Noir vineyards as somatic mutations affecting individual shoots, producing flowers with vigorous stamens (filaments 3-4 mm, anthers 0.8 mm) but pistil reduced to rudimentary structure 0.4-0.6 mm long, setting no fruit and identified during first flowering season for shoot removal or vine roguing. Breeding populations derived from crosses with wild V. vinifera ssp. sylvestris or dioecious North American Vitis species segregate approximately 25-50% staminate individuals (depending on parental genotypes) bearing staminate flowers, requiring early identification and removal to avoid allocating resources to non-fruit-bearing genotypes.
A staminate flower is an imperfect (unisexual) flower possessing functional stamens producing viable pollen but completely lacking a functional pistil or possessing only a vestigial, rudimentary, or aborted pistil incapable of fertilization and fruit set, representing true morphological male flowers that occur in dioecious wild Vitis vinifera subspecies sylvestris, occasionally in cultivated Vitis vinifera as genetic reversions or breeding population segregants, and characteristically in various wild North American Vitis species. Staminate flowers contrast fundamentally with perfect flowers (which possess both stamens and pistil morphologically) and with functionally male flowers (which possess pistil morphologically but with impaired function), as staminate flowers exhibit complete or near-complete absence of female reproductive structures at the morphological level. The staminate flower typically measures 2.5-4 mm in diameter at anthesis and comprises= (1) a minute calyx forming a shallow receptacle cup (2) five petals fused into a calyptra that detaches at anthesis (3) five well-developed stamens positioned opposite the petals, with robust filaments 2-4 mm long and large anthers 0.6-1 mm long containing abundant pollen (4) a pistil that is either completely absent (rare), extremely reduced to a minute non-functional vestige <0.5 mm long lacking recognizable stigma and style (common), or rudimentary with barely recognizable pistil morphology but severely abbreviated length and non-functional anatomy. The stamens in staminate flowers typically show enhanced development compared to hermaphrodite flowers, with 20-50% greater filament length, 15-30% larger anther size, and 30-60% increased pollen production per flower (often 25,000-45,000 pollen grains per staminate flower versus 15,000-30,000 per hermaphrodite flower), representing apparent resource reallocation from absent or minimal female structures to enhanced male function. Pollen viability in staminate flowers is typically high (70-90%), often exceeding hermaphrodite pollen viability, with pollen exhibiting normal morphology, robust germination on receptive stigmas, and vigorous pollen tube growth. Staminate flowers achieve essentially zero fruit set (<0.1%) due to absence or complete dysfunction of female reproductive structures, with plants bearing exclusively staminate flowers functioning purely as pollen donors (male parents) in reproductive biology. Staminate flowers characterize the male phase of dioecious wild V. vinifera ssp. sylvestris populations throughout the species' native Mediterranean and Near Eastern range, where approximately 50% of individuals in wild populations are staminate (male) plants bearing exclusively staminate flowers, with the remaining 50% being pistillate (female) plants bearing functionally female flowers with reflex stamens. The dioecious reproductive system of wild V. vinifera represents the ancestral condition, with the hermaphrodite condition of cultivated V. vinifera ssp. vinifera arising through mutations that restored pistil function (converting staminate to hermaphrodite flowers) or enhanced stamen function (converting pistillate to hermaphrodite flowers), with these mutations strongly selected during domestication to enable self-fertility and reliable fruit production from individual vines. Staminate flowers occasionally appear in cultivated vineyards as= (1) somatic reversions where individual shoots or vines within hermaphrodite cultivars produce staminate flowers through bud sport mutations (2) seedlings from uncontrolled pollination exhibiting genetic segregation for sex determination and reverting to staminate expression (3) intentional crosses with wild dioecious germplasm reintroducing staminate alleles into breeding populations. The genetic control of staminate versus hermaphrodite expression involves a small sex-determining region with male-promoting alleles that suppress pistil development, with staminate individuals typically carrying male-determining alleles in hemizygous or homozygous state.
Staminate_Flower represents true morphological male flowers and must be carefully distinguished from Functionally_Male_Flower (which possesses pistil morphologically but with impaired function creating functional but not morphological unisexuality). The distinction is critical= staminate flowers lack pistil structurally (imperfect flowers), while functionally male flowers possess pistil structurally but functionally (perfect flowers with female organ dysfunction). Staminate flowers are rare in cultivated V. vinifera ssp. vinifera but characteristic of wild V. vinifera ssp. sylvestris and represent the ancestral dioecious condition from which hermaphrodite cultivated grapes evolved through domestication. The presence of staminate flowers in commercial vineyards indicates genetic reversion or introgression and creates non-productive plants requiring removal. Sub-classification into Male_Only (no pistil trace) versus Reduced_Pistil (vestigial or rudimentary pistil present) reflects variation in the degree of pistil suppression in staminate flowers. Understanding staminate flower genetics and development informs evolutionary studies of sex determination in Vitis and breeding strategies for managing sex expression in hybrid populations.
Staminatblüte
male flower
männliche Blüte
pollen-producing flower
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 100-105 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644 Antcliff, A.J. (1980). Inheritance of sex in Vitis. Annales de l'Amélioration des Plantes 30=113-122 Battilana, J. et al. (2013). The genetic basis of the conversion of hermaphrodite flowers to male flowers in grapevine. Nature Communications 4=2615 Picq, S. et al. (2014). A small XY chromosomal region explains sex determination in wild dioecious V. vinifera. BMC Plant Biology 14=229 Fechter, I. et al. (2012). Candidate genes within a 143 kb region of the flower sex locus in Vitis. Molecular Genetics and Genomics 287=247-259.
Staminate Flower
Pinot Noir clone 115 propagation material at Standard (CAC) grade bears a Standard_Clone_Role, visually inspected and symptom-free but without the laboratory testing documentation confirming pathogen absence—sold at prices 15-20% below Certified grade material. Chardonnay clone 96 Standard-grade vines serve in a Standard_Clone_Role for cost-conscious vineyard establishment, the clone identity and traceability assured but virus-testing status limited to visual inspection. A Merlot clone 181 shipment with CAC labeling bears a Standard_Clone_Role, meeting minimum regulatory requirements for commercial sale but without the enhanced health assurance of Certified-grade material.
An official clone role that inheres in a grapevine clone meeting baseline (Standard/CAC) certification requirements, the minimum level permitting commercial propagation and sale within EU and equivalent regulatory frameworks. Standard-grade clones (designated "CAC" for Conformitas Agraria Communitatis in EU terminology) must demonstrate official variety and clone identification (trueness-to-type), freedom from visual symptoms of regulated pathogens, and documented traceability within the certification system—but without the mandatory laboratory testing required for Certified grade material. Standard-grade propagation material originates from mother plants inspected for disease symptoms but not necessarily tested by laboratory methods (ELISA, PCR), relying on visual freedom from symptoms rather than confirmed pathogen absence. The Standard/CAC grade represents the bulk of commercial grapevine production in many markets, offering registered clone identity and traceability at lower cost than Certified material, with the trade-off of reduced health assurance for pathogens that may be present asymptomatically. Standard-grade material is appropriate for lower-risk applications (phylloxera-free regions, varieties less susceptible to virus impact) but increasingly displaced by Certified material as growers recognize the long-term costs of virus-infected plantings.
Standard_Clone_Role represents the baseline commercial tier permitting propagation and sale, where registered clone identity is assured but health status confirmation is limited to visual methods. The economic reality of Standard-grade dominance reflects price sensitivity—many growers select the lower-cost option without fully accounting for the potential long-term costs of virus introduction. The asymptomatically infected plant is the fundamental limitation of visual-only inspection plants may appear healthy while harboring pathogens that will cause progressive decline, spread to neighboring vines through vectors, and reduce vineyard productivity over the 25-40 year vineyard lifespan. Industry trends favor increasing adoption of Certified material as virus disease awareness grows and certification programs expand capacity, though Standard-grade material remains significant in cost-sensitive markets and lower-value applications.
Basiskategorie-Klon
CAC clone
CAC-Klon
base grade clone
commercial grade clone
European Union (2020) Commission Implementing Directive 2020/177 Annex IV EPPO (2008) Certification schemes for grapevine, EPPO Bulletin 38: 422-429 Mannini F (2000) Clonal selection in grapevine, ISHS Acta Horticulturae 528: 17-25 Gramaje D, Armengol J (2011) Fungal trunk pathogens, Phytopathologia Mediterranea 50: S217-S226
Standard Clone Role
A standard long fruiting cane on Vitis vinifera cv. Sauvignon Blanc in Marlborough, New Zealand, retained at 12 nodes spanning 130cm along fruiting wire in vertical shoot positioning system, selected from balanced vine showing 580g pruning weight appropriate for this bud count per standard balanced pruning formula (30 buds base + 10 per 450g pruning weight), positioned for efficient shoot management and mechanical hedging, expected to produce 11 fruiting shoots bearing 19-21 clusters totaling 4.8kg fruit, managed through single cluster thinning pass to 14-16 clusters and standard canopy management (shoot positioning, hedging, leaf removal) achieving target 12 cm²/g leaf area-to-fruit ratio and appropriate fruit exposure for variety character expression.
A long fruiting cane retained at conventional extended length for cane-pruned systems, typically 10-14 nodes, representing established practice in many traditional wine regions and modern high-productivity systems, characterized by moderate linear extent (110-160cm), substantial but manageable bud count balancing yield and quality, compatibility with standard vertical shoot positioning and similar training systems, and production of adequate shoot numbers for commercial crop loads without excessive canopy density. Standard long fruiting canes represent mainstream viticulture practice in cane-pruned regions worldwide, combining proven productivity with manageable canopy requirements, suitable for wide range of cultivars and site conditions, and adaptable to varying quality and yield objectives through cluster thinning and canopy management adjustments.
Standard long fruiting canes represent the default cane length in many established wine regions, with 10-14 node range reflecting accumulated practical experience balancing productivity and quality across diverse conditions. This range suits most wine cultivars in balanced vineyards, provides flexibility for fine-tuning through shoot and cluster thinning, allows efficient use of vertical trellis space, and remains compatible with both hand and mechanical canopy management operations. Regional traditions often specify typical cane lengths (e.g., 12 nodes in parts of Germany, 10 nodes in Burgundy, 14 nodes in some New World regions), reflecting adaptation to local cultivars, climate, soils, and quality standards, though individual vineyard variation based on vine vigor assessment remains standard practice.
Standard-Langrute
conventional long cane
normal long fruiting cane
typical extended cane
übliche lange Fruchtrute
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181.
Standard Long Fruiting Cane
Cabernet Sauvignon hermaphrodite flowers across diverse vineyard sites and vintages consistently exhibit standard medium styles 0.56-0.66 mm (averaging 0.61 mm) connecting ovaries 0.9-1.05 mm to stigmas 0.45-0.52 mm, with standard medium styles representing 27-31% of total pistil length (2.1-2.35 mm), containing elaborate transmitting tract tissue with abundant extracellular matrix and specialized cells, supporting pollen tube transit in 58-82 hours at 20-24°C, and achieving 50-58% fruit set under favorable bloom conditions, representing optimal hermaphrodite pistil morphology and function. Merlot shows standard medium styles 0.58-0.68 mm (averaging 0.63 mm) across 12 commercial clones evaluated in replicated trials, demonstrating consistency of standard medium style length within this cultivar despite clonal and environmental variation, with all clones achieving 48-58% fruit set indicating functional equivalence across the standard medium range. Syrah standard medium styles 0.56-0.66 mm examined in anatomical studies reveal well-developed transmitting tract consisting of 25-40 cell layers of elongated specialized cells surrounding central canal 0.08-0.12 mm diameter filled with extracellular matrix staining intensely with ruthenium red (indicating pectins) and alcian blue (indicating acidic polysaccharides), representing optimal transmitting tract morphology supporting efficient pollen tube growth and guidance to ovules. Chardonnay pollen tube growth kinetics in standard medium styles show consistent growth rates of 16-24 μm per hour across the 0.58-0.68 mm style length range, with pollen tubes traversing the style proper in 28-42 hours plus additional time in stigma (8-15 hours) and ovary tissues (12-22 hours) totaling 48-79 hours stigma-to-ovule transit, representing typical pollen tube growth dynamics in optimally dimensioned styles.
A standard medium style is a medium style in the middle-to-upper portion of the medium length range, typically 0.55-0.7 mm at maturity, representing the most common style length among hermaphrodite cultivated Vitis vinifera cultivars and representing styles fully within the optimal medium dimension range for efficient pollen tube support and fertilization. Standard medium styles are characterized by= (1) length typically 0.55-0.7 mm at anthesis, occupying the upper half of medium style range and representing the mode of style length distribution across hermaphrodite cultivars (2) diameter typically 0.22-0.35 mm proportional to standard length (3) well-developed transmitting tract tissue with elaborate extracellular matrix, abundant specialized transmitting tract cells, and clear central stylar canal providing optimal pollen tube pathway (4) connection between ovary (typically 0.85-1.1 mm) and stigma (typically 0.42-0.55 mm) creating well-proportioned pistils (total length typically 2.1-2.5 mm) with standard medium styles representing 25-32% of pistil length. Standard medium styles represent the typical outcome of normal pistil development in major hermaphrodite wine cultivars under favorable to optimal growing conditions= Cabernet Sauvignon, Merlot, Chardonnay, Syrah, and most other economically important cultivars show standard medium style lengths averaging 0.56-0.66 mm across diverse environments, demonstrating that this range represents the evolutionarily selected optimum for cultivated hermaphrodite grapevines. Standard medium styles develop through complete expression of pistil developmental programs in hermaphrodite flowers= normal style primordium initiation and elongation (BBCH 51-57) full transmitting tract differentiation with maximum extracellular matrix production (BBCH 56-59) achievement of mature dimensions representing the genetic potential under favorable developmental conditions. Standard medium styles provide optimal functional performance= transmitting tract length (0.55-0.7 mm) combined with stigma and ovary dimensions creates total pollen tube pathway of 1.6-2.5 mm, requiring 55-90 hours for pollen tube transit at typical bloom temperatures (20-24°C), providing adequate time for pollen tube maturation and nutrition while avoiding excessive delays that might increase vulnerability to environmental stress transmitting tract diameter and cellular organization support 3-6 simultaneous pollen tube passages, allowing efficient fertilization when multiple pollen grains germinate hermaphrodite flowers with standard medium styles consistently achieve 45-60% fruit set under favorable bloom conditions across diverse cultivars and environments, demonstrating that standard medium dimensions support reliably high reproductive success. The standard medium style category encompasses the majority of hermaphrodite flowers across cultivated V. vinifera, making this the reference or baseline style morphology against which deviations (short styles in functionally male flowers, long styles in functionally female flowers, moderate medium styles in compact pistil variants) are compared and assessed.
Standard_Medium_Style represents the typical and most common medium style length and is distinguished from Moderate_Medium_Style (0.45-0.58 mm, lower medium range) by fuller dimensions while both remain functional medium styles. Standard medium styles represent the optimal hermaphrodite style morphology selected through cultivation and represent the reference baseline for Vitis reproductive anatomy. Understanding standard medium styles is essential for defining normal hermaphrodite pistil structure, for comparative studies, and for establishing the morphological and functional benchmarks against which variant style types are assessed. Standard medium styles characterize the vast majority of hermaphrodite flowers in commercial viticulture.
optimal medium style
standard mittellanger Griffel
typical medium style
typischer Normal-Stempelgriffel
typischer Normalstylus
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 102-106 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 94-97 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=93-105.
Standard Medium Style
Müller-Thurgau clone Gm 64-18 bearing a standard reference role is planted as the mandatory check variety in all German Vitis breeding program field trials at Geisenheim, Weinsberg, and Freiburg, providing the performance baseline against which 200+ experimental seedlings are evaluated annually for yield (reference mean 12-15 t/ha), must weight (reference mean 65-75° Oechsle), titratable acidity (reference mean 7-9 g/L), and downy mildew susceptibility (reference OIV 452 score 5-6), with 30+ years of accumulated reference data enabling precise estimation of genotype × environment × year interactions across trial sites.
A reference cultivar role that inheres in a grapevine accession routinely included in experimental trials, phenological monitoring networks, or multi-site evaluations as a known performance benchmark, providing the comparative baseline against which novel genotypes, management treatments, or environmental responses are quantitatively assessed.
Standard_Reference_Role describes the operational use of known cultivars as experimental controls and performance benchmarks — a function essential to the reproducibility and comparability of viticultural research across sites and years. The selection of appropriate standard references is methodologically critical: the reference cultivar must be widely grown, phenologically representative of the trial's target maturity range, and well-characterized under local conditions to provide a meaningful comparative baseline. Standard references differ from type specimens in serving a functional-comparative rather than nomenclatural purpose, though the same accession may fulfill both roles.
Kontrollsorten-Rolle
Standard-Referenz-Rolle
Vergleichssorten-Rolle
benchmark reference role
check cultivar role
control variety role
OIV (2009) 2nd Edition of the OIV Descriptor List for Grape Varieties and Vitis Species Töpfer R et al. (2011) New horizons for grapevine breeding, Fruit, Vegetable and Cereal Science and Biotechnology 5: 79-100 Eibach R and Töpfer R (2015) Traditional grapevine breeding techniques, in Reynolds AG (ed) Grapevine Breeding Programs for the Wine Industry, Woodhead Publishing pp: 3-22
Standard Reference Role
In a Chianti Sangiovese vineyard, steady Brix increase was documented over 28 days in September Year N+1 (BBCH 85-86, active accumulation) when weekly Brix sampling showed consistent gains: 16.8° → 18.5° → 20.2° → 21.8° → 23.3° (average 1.7° per week, CV 8.5%), daily increment averaged 0.24% Brix (SD 0.032), linear regression r² = 0.94 enabling harvest forecast 18 days advance predicting 25° target within 2-day accuracy, under stable Tuscan autumn conditions (daily high 24-27°C, minimal variation).
Steady_Brix_Increase represents the consistent progressive elevation of berry soluble solids concentration during active accumulation when day-to-day Brix measurements show relatively uniform increments creating predictable sweetening trajectory enabling accurate harvest timing forecasts and indicating normal healthy ripening metabolism. This steady increase occurs during active accumulation period (BBCH 84-86), typically days 14-38 post-veraison spanning 22-32 days, and is characterized by daily Brix increments 0.22-0.35% per day (1.5-2.5% per week) showing coefficient of variation 0.88-0.96) between time and Brix demonstrating predictability, weekly sampling showing 1.8-3.2° Brix gain per week sustained over 3-5 consecutive weeks without dramatic fluctuation, and cumulative effect producing 7-12° Brix total increase during steady period representing 50-75% of total ripening gain achieved through consistent gradual accumulation. Statistical characteristics of steadiness include daily measurements showing normal distribution around mean daily increment with standard deviation 15-25% from running average indicating minimal day-to-day volatility, week-to-week comparison showing 3-5 days without measurable increase) or reversals (Brix decline) that would indicate metabolic dysfunction or severe environmental stress. Physiological basis for steadiness includes sustained transporter expression 85-115 fold above green berry with minimal day-to-day fluctuation maintaining stable import capacity, enzyme activities relatively constant with cell wall invertase 65-110 units/g fresh weight and transport proteins 75-135 μmol hexose uptake capacity g⁻¹ h⁻¹ showing <15-25% variation over measurement period, hormonal regulation with ABA 1500-2700 ng/g sustained without dramatic spikes or troughs providing stable ripening signal, and metabolic homeostasis where balance between import, respiration, and storage maintained through regulatory feedback preventing excessive fluctuation. Source-sink stability during steady period demonstrates photosynthesis rates relatively consistent 11-18 g CO₂ fixed per vine per day over multi-week period under favorable weather providing stable substrate supply, phloem delivery 180-320 mg sucrose per berry per day maintained within 25-40% range week-to-week, sink strength high with berry demand 70-85% of photosynthate relatively constant, and source-sink ratio stable through appropriate crop load (0.8-1.2 kg per m² leaf area) preventing source limitation that could disrupt steady accumulation. Environmental conditions supporting steadiness include stable moderate weather with temperatures 23-29°C range without extreme heat or cool spells producing consistent metabolic rates, adequate water availability maintaining stem water potential -0.7 to -1.3 MPa without severe fluctuation supporting stable phloem function, minimal disease or pest pressure preventing metabolic disruption, and absence of severe weather events (hail, extreme wind, excessive rain) that could directly damage berries or indirectly affect metabolism disrupting steady accumulation. Contrast with unsteady accumulation patterns includes initial acceleration phase showing increasing daily increments 0.12-0.20 → 0.28-0.38% Brix per day over 10-15 days as capacity develops representing developmental ramp-up not steady-state, final deceleration showing declining increments 0.25-0.32 → 0.08-0.15% per day as limits approached representing physiological constraint not steady accumulation, and stress-induced disruption showing erratic pattern with plateaus, reversals, or dramatic rate changes indicating abnormal metabolism requiring management attention. Predictive value of steadiness enables linear extrapolation from current Brix and steady rate projecting harvest date 2-4 weeks advance with reasonable accuracy ±3-7 days, degree-day models calibrated during steady period improving forecast precision, growers using steady period data to schedule labor, equipment, and winery capacity with confidence, and recognition that departure from steady pattern (acceleration, deceleration, plateau) signals changing conditions requiring management response or harvest timing adjustment. Quality implications include steady increase generally indicating healthy ripening with normal metabolism producing balanced compositional development, coordination between steady sugar increase and concurrent acid decline establishing appropriate sugar-acid balance developing predictably, uniformity across vineyard zones showing similar steady patterns indicating homogeneous ripening enabling single-pass harvest, and heterogeneity where some zones steady while others erratic revealing management issues or site variation requiring selective treatment. Cultivar differences in steadiness characteristics show some varieties displaying highly consistent patterns (Chardonnay, Merlot) enabling reliable forecasting, others showing greater variability (Pinot Noir particularly warm climates) requiring more frequent monitoring, and inherent rate differences with rapid cultivars showing higher daily increments 0.28-0.38% Brix per day while slower types 0.18-0.26% per day reflecting genetic variation in accumulation capacity. Temperature effects on steady rates demonstrate positive correlation with mean temperature during steady period explaining 60-80% of rate variation (r² = 0.60-0.80), cool vintages producing slower steady rates 1.2-2.0% Brix per week extending accumulation duration requiring patience, warm vintages showing faster rates 2.2-3.5% per week shortening period potentially requiring earlier harvest, and excessive heat disrupting steadiness through metabolic stress creating erratic patterns. Water management effects show adequate irrigation supporting steadiness through maintained phloem function and stable metabolism, deficit irrigation potentially slowing steady rate 15-30% through reduced berry expansion while maintaining or enhancing concentration, excessive water potentially disrupting steadiness through dilution effects or metabolic perturbation, and irrigation scheduling during steady period requiring balance between supporting accumulation and controlling berry size affecting concentration. Monitoring strategies during steady period include weekly Brix sampling minimum frequency with twice-weekly optimal for precise rate determination, spatial sampling across vineyard zones revealing uniformity or heterogeneity, temporal sampling tracking consistency enabling detection of departures from steady pattern indicating developing issues, and integration with other maturity parameters (pH, TA, phenolics, sensory) ensuring comprehensive assessment recognizing Brix alone insufficient for harvest decisions. Harvest timing decisions based on steady increase involve recognition that steady period typically concludes 1-3 weeks pre-harvest as deceleration begins, continued monitoring required as steadiness ends to determine actual harvest timing, target Brix achievement timing projected from steady period rates enabling advance planning, and flexibility to harvest earlier if environmental stress threatens to disrupt continued accumulation or later if phenolic maturity lags sugar development requiring extended ripening.
Steady_Brix_Increase is distinguished from initial acceleration by showing consistent (CV 0.88. Predictive value for harvest timing. Steady_Brix_Increase represents normal healthy ripening pattern enabling management planning.
consistent Brix progression
einheitliche lösliche Feststoff-Zunahme
konsistente Brix-Progression
predictable sweetening
stabile Zucker-Konzentrations-Anstieg
stable sugar concentration rise
uniform soluble solids increase
vorhersagbare Süßung
Coombe BG (1992) Research on development and ripening of the grape berry Amer J Enol Vitic 43:101-110 ripening patterns Coombe BG, McCarthy MG (2000) Dynamics and environmental control of grape berry growth Australian J Grape Wine Res 6:73-83 Brix increase dynamics Keller M (2020) The Science of Grapevines 3rd Ed pages 224-227 steady accumulation patterns.
Steady Brix Increase
Merlot hardwood cutting showing stem-derived adventitious roots, 35 cm cutting with 4 nodes treated with 4000 ppm IBA, examination 21 days after treatment revealing stem adventitious root primordia forming from internodal cambium between nodes 2 and 3, microscopic sections showing primordia originating 0.8-1.2 mm deep in vascular cambium, primordia emerging through cortex and periderm 26 days after treatment, cutting producing 12 adventitious roots total= 7 nodal, 5 stem-derived, stem roots supplementing nodal roots creating robust root system
Stem derived adventitious root is shoot-derived adventitious root originating from stem internodal tissues between nodes, characterized by formation from vascular cambium or pericycle-like cells in internodal regions, showing induction typically requiring wounding, hormone treatment, or burial providing auxin accumulation and favorable conditions, demonstrating endogenous development within stem with primordium forming from internal cells and growing outward through cortex and periderm, and representing secondary or supplementary root formation site compared to often-preferential nodal rooting, with stem-derived roots in grapevine showing variable capacity depending on cultivar, stem age, and induction conditions, typically requiring stronger hormonal or environmental stimulus than nodal rooting due to absence of preformed primordia, and demonstrating importance in propagation techniques where extensive stem burial or hormone treatment promotes internode rooting supplementing nodal roots creating more robust root systems. Stem derived adventitious root initiation sites includes= showing cambial origin (majority of stem adventitious roots initiating from vascular cambium cells), exhibiting interfascicular regions (roots potentially forming between vascular bundles in interfascicular cambium or parenchyma), demonstrating ray involvement (vascular rays potentially serving as preferential initiation sites), revealing pericycle-like cells (cells adjacent to phloem showing pericycle-like characteristics initiating roots), and displaying cortical potential (stem cortex occasionally participating in root formation particularly after wounding). Stem derived adventitious root induction requirements includes= requiring hormonal stimulus (stem rooting typically needing auxin application or accumulation for successful initiation), showing concentration dependence (effective auxin concentrations for stem rooting often 2000-8000 ppm IBA or NAA), exhibiting wounding response (cutting or mechanical damage potentially enhancing stem rooting through wound signaling), demonstrating burial effects (covering stems with soil or media creating dark moist conditions promoting rooting), and revealing age effects (young green stems often rooting more readily than old woody stems though both capable). Stem derived adventitious root development sequence includes= showing cell dedifferentiation (cambial or other stem cells losing original identity), exhibiting cell division (dedifferentiated cells dividing to form root primordium cell population), demonstrating meristem organization (dividing cells organizing into structured root meristem with cap and provascular tissue), revealing outward growth (organized primordium growing radially through stem cortex toward surface), and displaying emergence (root breaking through cortex and periderm emerging at stem surface). Stem derived adventitious root anatomical constraints includes= showing greater depth (stem roots forming deeper within stem tissues than epidermal or nodal roots requiring longer emergence path), exhibiting tissue barriers (stem cortex and thick periderm potentially impeding root emergence), demonstrating vascular integration challenge (stem roots requiring establishment of vascular connection to stem cylinder), revealing mechanical impedance (dense woody tissue in old stems potentially physically restricting root development), and displaying emergence wounds (root emergence creating larger disruption in stem tissues than at nodes). Stem derived adventitious root versus nodal roots includes= contrasting in frequency (stem roots typically less numerous than nodal roots unless strongly induced), showing later development (stem roots often forming after nodal roots in propagation sequences), exhibiting different induction (stem requiring stronger stimulus than nodes where preformed primordia may exist), demonstrating supplementary role (stem roots supplementing rather than replacing nodal roots in most propagation), and revealing cultivar patterns (easy-rooting cultivars showing abundant stem roots while difficult types primarily nodal). Stem derived adventitious root in propagation applications includes= enabling long cutting rooting (extensive stem length allowing multiple stem roots supplementing nodal roots), providing trench layer rooting (buried canes forming stem roots along entire buried length), demonstrating stooling enhancement (mounded soil contacting stem inducing roots beyond just nodal positions), revealing micropropagation use (tissue culture shoots forming stem roots during rooting phase), and displaying field layer application (field-layered canes producing stem roots where buried). Stem derived adventitious root functional capacity includes= showing complete functionality (stem-derived roots developing full absorptive and transport capacity), exhibiting lateral branching (stem adventitious roots producing lateral roots creating branched system), demonstrating secondary growth (stem roots undergoing secondary thickening in perennial plants), revealing storage capacity (stem adventitious roots accumulating carbohydrate reserves), and displaying longevity (stem-derived roots persisting throughout plant life functioning as permanent root system components). Stem derived adventitious root cultivar specificity includes= showing variable capacity (grapevine cultivars ranging from prolific stem rooting to minimal stem root formation), exhibiting rootstock selection (rootstock breeding considering stem rooting capacity for propagation efficiency), demonstrating scion patterns (V. vinifera cultivars generally showing moderate stem rooting capacity), revealing hybrid characteristics (interspecific hybrids showing variable stem rooting reflecting parentage), and displaying practical importance (cultivar stem rooting capacity affecting propagation protocol design and success rates).
Stem_Derived_Adventitious_Root emphasizes internodal stem origin between nodes and is distinguished from nodal roots by focus on internode formation. Critical for understanding cambial origin, induction requirements, anatomical constraints, supplementary role, propagation applications, and the adventitious roots arising from stem internodes supplementing nodal roots in vegetative propagation.
Between-node root
Cambial adventitious root
Internodal adventitious root
Internodale Wurzel
Kambiale Adventivwurzel
Stamm-Adventivwurzel
Stem Derived Adventitious Root
Stem hydraulic conductivity assessment in mature Shiraz reveals the capacity and vulnerability of the aboveground transport pathway, with fully hydrated stems showing specific conductivity (Ks) of 45 kg/m/s/MPa, declining to 30 kg/m/s/MPa at midday leaf water potential of -1.2 MPa (15% native embolism), and catastrophically declining to 12 kg/m/s/MPa following severe drought reaching -2.0 MPa (75% conductivity loss from extensive embolism exceeding P50 of -1.8 MPa), demonstrating the vulnerability of stem conductivity to hydraulic dysfunction that limits drought tolerance and may require recovery time or intervention following severe stress events.
Stem hydraulic conductivity quality is a hydraulic conductivity quality that represents the capacity of stem xylem to conduct water from roots to leaves, characterizing the aboveground axial water transport pathway that must supply all transpiring foliage with sufficient water to prevent excessive water potential decline under varying evaporative demand. Within the BFO framework, this quality specifically addresses the stem component of hydraulic architecture, distinguishing trunk and branch conductivity from root and leaf components while recognizing that stem xylem represents the main highway of long-distance water transport whose conductivity must scale appropriately with the transpiring leaf area it supplies. The ontological significance lies in stem hydraulic conductivity determining the efficiency of water delivery from roots to the distributed leaf canopy, where the resistance to water flow through stem xylem establishes the water potential gradient between root crown and leaves that, combined with root and leaf resistances, determines the leaf water potential achieved at any transpiration rate. The anatomical basis for stem hydraulic conductivity involves xylem vessel characteristics including diameter (fourth-power relationship with conductivity), length (longer vessels reducing end-wall resistance), pit membrane properties (affecting inter-vessel flow), and vessel density (total conducting area), with these features varying among cultivars and rootstocks and along the stem from trunk through progressively smaller branches. The scaling of stem conductivity with supported leaf area follows the principle of hydraulic segmentation, where conductivity progressively declines from trunk toward branch tips in proportion to decreasing leaf area served, maintaining relatively uniform water supply per unit leaf area throughout the canopy. The vulnerability of stem conductivity to embolism is a key aspect of drought tolerance, with the threshold water potential causing 50% conductivity loss (P50) indicating the safety margin for hydraulic function, and with grapevine showing moderate vulnerability (P50 typically -1.5 to -2.5 MPa) that provides reasonable drought tolerance but limits function under severe stress. Understanding stem hydraulic conductivity quality enables assessment of the aboveground water transport capacity, drought vulnerability evaluation from embolism susceptibility, and interpretation of how stem anatomy affects water supply to transpiring leaves.
Stem_Hydraulic_Conductivity_Quality serves as the parent class for stem pathway components (Vessel, Petiole, Shoot). This quality represents the aboveground axial transport that must supply all transpiring foliage. The class supports drought tolerance assessment and water transport evaluation through stem conductivity and vulnerability characterization.
Stammhydraulikleitfähigkeit
Stammleitfähigkeit
Xylemleitfähigkeit
stem conductance
trunk hydraulic capacity
xylem conductivity
Tyree MT, Ewers FW (1991) The hydraulic architecture of trees and other woody plants. New Phytologist 119(3): 345-360 Choat B et al. (2010) Vascular function in grapevines relates to the extent and rate of drought-induced embolism. Physiologia Plantarum 140(2): 126-136
Stem Hydraulic Conductivity Quality
Stem water potential-based irrigation scheduling in San Joaquin Valley Chardonnay employs threshold values of -1.0 MPa for full irrigation maintaining maximum yield potential and -1.4 MPa for regulated deficit irrigation promoting flavor concentration, with weekly measurements of bagged leaves at 13:00 showing coefficient of variation of 8% compared to 22% for exposed leaf measurements, enabling reliable detection of 0.1 MPa changes in water status for precise irrigation timing.
Stem water potential quality is a water potential quality that represents the xylem water potential measured in non-transpiring tissue, typically a leaf that has been enclosed in a reflective bag to eliminate transpiration, providing assessment of the water potential in the main stem xylem that supplies the transpiring canopy. Within the BFO framework, this quality specifically characterizes the hydraulic status of the transport pathway rather than the end-point leaf tissue, distinguishing stem water potential from leaf water potential and providing information about the driving force available for water delivery to transpiring leaves. The ontological significance lies in stem water potential representing the supply-side pressure in the vine hydraulic system, where the gradient between stem and leaf water potential drives flow through petioles and leaf vasculature, and the stem value indicates the integrated effect of soil water availability and root-to-stem hydraulic conductance without the additional resistance of leaf pathways. Stem water potential is less variable than leaf water potential because it eliminates the influence of individual leaf exposure, stomatal conductance, and boundary layer conditions, providing a more stable indicator of whole-plant water status suitable for irrigation scheduling in commercial viticulture. The measurement protocol requires enclosing leaves in reflective, opaque bags for 1-2 hours before measurement to allow equilibration with stem xylem water potential, with the bagged leaf then excised and measured immediately in a pressure chamber. Stem water potential values are typically 0.1-0.3 MPa less negative than midday leaf water potential under the same conditions, with the difference reflecting the hydraulic resistance of leaf pathways. The reduced variability and clearer relationship to whole-plant water status have led to widespread adoption of stem water potential for irrigation management, with cultivar-specific thresholds established for optimal deficit irrigation in many wine grape varieties.
Stem_Water_Potential_Quality addresses the xylem transport pathway water status, distinguished from Leaf_Water_Potential_Quality by the bagging protocol eliminating transpiration. This quality has become the preferred water status indicator for commercial irrigation management due to reduced variability and clearer relationship to whole-plant hydraulics. The bagging equilibration period (minimum 1 hour, preferably 2 hours) is critical for valid measurement, as insufficient equilibration yields intermediate values between stem and leaf water potential.
Pflanzenwasserpotential
Sprosswasserpotential
Stammwasserpotential
Xylemwasserpotential
plant water potential
trunk water potential
xylem water potential
Ψstem
Choné X et al. (2001) Stem water potential is a sensitive indicator of grapevine water status. Annals of Botany 87(4): 477-483 McCutchan H, Shackel KA (1992) Stem-water potential as a sensitive indicator of water stress in prune trees (Prunus domestica L. cv. French). Journal of the American Society for Horticultural Science 117(4): 607-611
Stem Water Potential Quality
Thompson Seedless (Sultanina/Sultana) bears a Stenospermocarpic_Table_Grape_Role as the ancestral source of commercial seedlessness, with pollination triggering berry development but seed abortion occurring approximately 2-3 weeks post-bloom, leaving soft 1-2 mm seed traces that are undetectable during consumption. Flame Seedless serves in a Stenospermocarpic_Table_Grape_Role with characteristic small seed traces (1-3 mm), requiring GA₃ treatment (10-40 ppm applications at berry set and sizing) to achieve commercial berry size of 4-5 g. Crimson Seedless bears a Stenospermocarpic_Table_Grape_Role demonstrating advanced breeding achievement—its seed traces remain soft and small (1-2 mm) while berry size (4-6 g) approaches seeded cultivars, reducing reliance on GA₃ treatment for commercial production.
A seedless table grape scion role that inheres in a grafted scion cultivar by virtue of its production of seedless berries through stenospermocarpy, a developmental process where pollination and fertilization occur normally, embryo development initiates, but seed development aborts at an early stage (typically 2-4 weeks post-bloom), leaving small, soft, edible seed traces (rudimentary seeds or seed remnants) within otherwise seedless berries. Stenospermocarpic cultivars require pollination for fruit set and benefit from pollen sources, but the developing embryo and surrounding seed tissues fail to complete development due to genetic factors disrupting endosperm and/or embryo growth. Berry size in stenospermocarpic varieties correlates with seed trace development—larger traces support larger berries—explaining the historical use of gibberellic acid (GA₃) applications to compensate for reduced berry size. Stenospermocarpy is the predominant seedlessness mechanism in commercial table grapes, inherited as a recessive trait from the Thompson Seedless lineage that contributes to the parentage of essentially all modern seedless varieties.
Stenospermocarpic_Table_Grape_Role captures the dominant seedlessness mechanism in commercial viticulture, inherited from Thompson Seedless through the majority of modern seedless breeding lineages. The genetic basis involves multiple loci controlling embryo abortion timing earlier abortion produces smaller seed traces but also smaller berries, creating a breeding trade-off. Embryo rescue techniques (culturing immature embryos in vitro before abortion) enable hybridization between stenospermocarpic parents, overcoming the barrier that seedless × seedless crosses cannot produce seed naturally. GA₃ applications remain standard practice for most stenospermocarpic varieties to achieve market-acceptable berry size, though breeding progress is gradually reducing GA dependency. Seed trace size and hardness vary by variety and growing conditions, with consumer tolerance for trace seeds higher in some markets than others.
Tafeltraube mit Rudimentärsamen
Traube mit Samenabbruch
aborted-seed grape
rudimentary-seed grape
stenospermocarpic grape
stenospermokarpische Tafeltraube
Ledbetter CA, Ramming DW (1989) Seedlessness in grapes, Horticultural Reviews 11: 159-184 Stout AB (1936) Seedlessness in grapes, New York Agricultural Experiment Station Technical Bulletin 238 Perl A et al. (2000) Seedless grapes, in Molecular Biology and Biotechnology of the Grapevine, Springer pp.: 327-351 Mejía N et al. (2011) Molecular, genetic and transcriptional evidence for a role of VvAGL11 in stenospermocarpic seedlessness in grapevine, BMC Plant Biology 11: 57
Stenospermocarpic Table Grape Role
Thompson Seedless exhibits stenospermocarpy with embryo abortion occurring 2-4 weeks post-fertilization, producing seed traces of 8-18 mg compared to 25-45 mg for fully developed seeds in seeded cultivars. Crimson Seedless shows later abortion timing with slightly larger seed traces (20-35 mg) that remain soft and edible. The stenospermocarpic trait is controlled by a major recessive gene (sdI) with quantitative modifiers affecting abortion timing and seed trace size. Pollen from stenospermocarpic cultivars is fully viable and produces normal seeded offspring when used to pollinate seeded maternal parents.
A seedlessness disposition that is the inherited capacity of a Vitis vinifera plant to initiate normal fertilization and seed development but subsequently abort the embryo and endosperm at a genetically determined developmental stage, resulting in berries containing small, soft, incompletely developed seed traces.
Stenospermocarpy requires successful pollination and fertilization, distinguishing it from parthenocarpy. The abortion process involves programmed cell death in the nucellus and endosperm, triggered by hormonal signals around 15-40 days post-anthesis depending on genotype. Seed trace hardness varies among stenospermocarpic cultivars and is an important quality criterion for fresh consumption. Breeding for seedlessness uses embryo rescue techniques to recover progeny from stenospermocarpic crosses.
Embryoabortion
Samenrudimentbildung
Stenospermokarpie
abortive seed formation
post-fertilization seedlessness
seed trace development
Stout AB (1936) Seedlessness in grapes. New York Agricultural Experiment Station Technical Bulletin 238:1-68 Bouquet A, Danglot Y (1996) Inheritance of seedlessness in grapevine (Vitis vinifera L.). Vitis 35:35-42
Stenospermocarpy Disposition
A sterile vegetative shoot on Vitis vinifera cv. Syrah arising from trunk latent bud activation (water shoot origin), developed to 185cm length with 21 leaves, 4 tendrils, no inflorescences despite mid-season timing and favorable positioning, examined over two subsequent seasons showing consistent non-fruiting status (retained as renewal wood after initial season, produced 14 shoots second season, zero reproductive shoots despite 8 shoots at mid-cane positions normally showing 85%+ fruitfulness for this cultivar), representing persistent sterility characteristic of water shoot origin from latent buds that lack reproductive programming due to formation timing or developmental constraints, with this sterility persisting through bud differentiation on water shoot wood demonstrating inherent reproductive incompetence rather than temporary circumstantial non-fruiting.
A vegetative shoot completely lacking reproductive structures through inherent infertility rather than positional or developmental factors, characterized by consistent non-fruiting status across multiple seasons despite favorable node positions, genetic or physiological inability to develop inflorescences, origin from persistently sterile bud positions or inherently non-reproductive growth contexts (water shoots, suckers, certain renewal shoots), and distinction from typical vegetative shoots that are non-fruiting due to position or circumstance but retain capacity for reproductive development under different conditions. Sterile vegetative shoots represent categorically non-reproductive growth, permanently or persistently unable to produce inflorescences regardless of position, conditions, or management, arising from genetic sterility factors, water shoot or sucker origin from latent buds lacking reproductive programming, severely stressed vines showing persistent reproductive failure, or specific renewal shoot contexts where vegetative-only development is consistent outcome.
Sterile vegetative shoots are distinguished from ordinary vegetative shoots by persistent categorical reproductive failure rather than circumstantial non-fruiting. Common sterile contexts include water shoots and suckers from latent bud origin (buds formed outside normal differentiation windows or under suppressed conditions lacking reproductive programming), certain genetic sterility conditions (rare), severely stressed vines showing chronic reproductive failure (though such vines typically show other severe symptoms), and specific renewal shoot situations where consistently low fruitfulness approaches functional sterility. The distinction has management implications as sterile shoots provide no current or near-term reproductive potential, suitable only for vegetative functions (photosynthesis, architectural renewal) while ordinary vegetative shoots from basal positions retain capacity for productive bud formation under favorable conditions. However, the sterile versus circumstantially-vegetative distinction can be ambiguous without multi-season observation, with definitive classification requiring demonstration of persistent reproductive failure across multiple opportunities.
fortpflanzungsunfähiger Trieb
infertile shoot
permanently non-fruiting shoot
reproductively incompetent shoot
steriler vegetativer Trieb
unfruchtbarer Trieb
Based on Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Srinivasan & Mullins (1981) "Physiology of flowering in the grapevine - a review" American Journal of Enology and Viticulture 32=47-63 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press, pp. 85-105.
Sterile Vegetative Shoot
Chardonnay hermaphrodite flowers at BBCH 61-65 (flowering) exhibit functional stigmas 0.42-0.52 mm diameter bearing 145-185 papillae distributed across the stigmatic surface, producing wet glistening exudate containing 10-13% sugars and supporting pollen grain adhesion, hydration, and germination with receptivity persisting 4-6 days post-calyptra shedding, achieving 48-56% fruit set through efficient self-pollination with pollen from elevated erect anthers falling onto receptive stigmas positioned below. Gewürztraminer functionally female flowers show enhanced stigmas 0.52-0.62 mm diameter with 180-235 papillae and abundant wet exudate production, exhibiting extended receptivity persisting 6-8 days allowing capture of cross-pollination from hermaphrodite pollinizer sources, achieving 28-38% fruit set when adequate pollinizer pollen is available during the extended receptivity window. Wild V. vinifera ssp. sylvestris pistillate female flowers exhibit elaborate stigmas 0.48-0.58 mm diameter positioned prominently above vestigial stamens, bearing 165-220 papillae with copious exudate, maintaining receptivity 5-7 days to maximize probability of capturing wind-borne pollen from staminate males, achieving 30-42% fruit set when males are within 20-50 m. Cabernet Franc rare functionally male flowers show reduced stigmas 0.28-0.38 mm diameter with 75-115 papillae producing minimal exudate (<20% of hermaphrodite levels), exhibiting poor or absent receptivity contributing to <8% fruit set even when adequate pollen is present, with reproductive failure primarily due to stigma non-receptivity and ovule abortion. Pinot Noir stigma receptivity timing studies using controlled pollination at intervals post-calyptra shedding reveal peak receptivity at 1-3 days post-shedding (pollen germination 75-88% of applied grains), declining receptivity at 4-5 days (45-65% germination), and loss of receptivity by 6-7 days (8-18% germination), demonstrating temporal dynamics of stigma receptivity and importance of pollen arrival timing for fertilization success.
A stigma is the terminal receptive portion of the pistil specialized for pollen capture, adhesion, hydration, and germination, functioning as the landing platform and initial interaction site where compatible pollen grains are recognized, supported, and induced to germinate, producing pollen tubes that grow through the style to achieve fertilization. The stigma structure in Vitis comprises= (1) stigmatic surface typically 0.3-0.65 mm in diameter at anthesis, though dimensions vary substantially among flower types, often showing subtle bilobed or bilateral organization reflecting the bicarpellate pistil origin (2) multicellular papillae (specialized epidermal projections) covering the stigmatic surface, typically 40-250 papillae per stigma depending on stigma size and flower type, with individual papillae measuring 20-80 μm in length and 12-35 μm in diameter (3) secretory cells within papillae and sub-papillar tissues producing wet stigmatic exudate containing sugars (primarily sucrose and glucose at 8-15% concentration), proteins (including enzymes, signaling molecules, and structural proteins), lipids, and other compounds that facilitate pollen grain adhesion, hydration, germination, and initial pollen tube growth (4) underlying transmitting tract tissue continuous with the style, providing pathway for pollen tubes to enter the style following germination on the stigma surface. Stigma development proceeds through phases= (1) primordium initiation at pistil apex (BBCH 51-53) (2) papillae differentiation (BBCH 54-57) with epidermal cells elongating and specializing to form papillae (3) secretory cell differentiation (BBCH 56-59) with cells developing capacity for exudate production (4) achievement of receptivity (BBCH 59-65) when exudate production activates, typically coinciding with or slightly preceding calyptra shedding and anther dehiscence. Stigma morphology and function vary dramatically among flower types= hermaphrodite flowers exhibit functional stigmas 0.35-0.55 mm diameter with 120-200 papillae, wet exudate production, and receptivity persisting 3-6 days, achieving 40-60% fruit set through self-pollination functionally female and pistillate flowers show enhanced stigmas 0.45-0.65 mm diameter with 150-250 papillae, abundant exudate, and extended receptivity 5-8 days, requiring cross-pollination and achieving 25-45% fruit set functionally male flowers exhibit reduced or rudimentary stigmas 0.2-0.4 mm diameter with 60-120 papillae, minimal or absent exudate, non-receptive or poorly receptive, achieving <5-15% fruit set staminate flowers show vestigial stigmas 0.1-0.25 mm diameter with <60 papillae, no exudate, completely non-receptive, achieving <1% fruit set. Stigma receptivity is the critical determinant of pollination success, with receptive stigmas capturing pollen grains (via exudate adhesion), hydrating pollen (via exudate water content), supporting pollen germination (via exudate nutrients and signaling molecules), and guiding pollen tube entry into transmitting tract, while non-receptive stigmas fail at one or more of these steps preventing fertilization.
Stigma represents the terminal pistil component and must be distinguished from Style (intermediate component) and Ovary (basal component). The stigma serves critical pollen reception and germination functions, with stigma receptivity (presence of wet exudate and pollen germination capability) being the primary determinant of whether pollination can proceed to fertilization. Stigma dimensions, papillae number, exudate production, and receptivity duration vary dramatically among flower types, providing key diagnostic features for sexual type identification. Sub-classification addresses receptivity state (Receptive versus Dry stigmas), papillae density (Densely versus Sparsely papillate), and receptivity quality (Fully versus Maximally receptive). Understanding stigma anatomy, development, and function is essential for pollination biology, fruit set prediction, and diagnosis of reproductive failures.
Empfangsfläche
Narbe
Stigma
pollen receptor
Merlot in Bordeaux shows stigma receptivity onset during early June Year N+1 at BBCH 61-62 within 12-18 hours after flower opening, with stigma papillate cells elongating and exudate secretion initiating creating glistening appearance, developing peak receptivity 1-2 days after opening persisting 5-7 days, under oceanic climate conditions of 19-24°C temperature, 55-70% relative humidity supporting normal function typical of this mid-season cultivar establishing female reproductive capacity for pollination
Stigma receptivity onset represents the critical functional development where newly exposed stigmas in opened flowers achieve physiological capacity to capture pollen and support germination, establishing essential female reproductive function enabling successful pollination and fruit set. This essential functional milestone occurs during early anthesis in early June in temperate regions Year N+1, typically developing within 6-24 hours after flower opening as stigma tissues complete maturation and begin exudate secretion, corresponding to BBCH 61-62 stages. Receptivity development involves stigma papillate cell elongation increasing surface area 2-3 fold creating textured surface maximizing pollen capture potential, exudate secretion initiating with proteinaceous fluid production creating adhesive coating on stigma surface, and metabolic activation establishing capacity to support pollen hydration, germination, and tube growth. Papillate cell characteristics show specialized epidermal cells elongating to 40-80 micrometers length creating finger-like projections, cells containing dense cytoplasm with abundant organelles supporting active secretion, and cell walls remaining relatively thin facilitating pollen tube penetration after germination. Exudate composition includes proteins particularly lipid transfer proteins facilitating pollen-stigma recognition, carbohydrates providing hydration medium for pollen, lipids affecting surface properties, and various small molecules potentially serving signaling or nutritive functions. Exudate production shows progressive accumulation creating visible glistening appearance on stigma surface, with production rate affected by environmental conditions particularly temperature and humidity, maximum accumulation typically occurring 1-3 days after flower opening when receptivity peaks. Stigma appearance at receptivity onset shows bilobed structure with two stigma lobes diverging at angle creating expanded capture surface, papillate surface texture visible under magnification appearing rough compared to smooth style tissue, and characteristic sheen from exudate coating indicating active secretivity. Pollen capture mechanism involves adhesive exudate trapping pollen grains contacting stigma surface, physical retention through papillate cell structure creating mechanical barriers preventing pollen loss, and possible electrostatic effects attracting charged pollen grains though evidence in grapevine is limited. Pollen-stigma interaction following capture involves pollen grain hydration from stigma exudate taking 15-60 minutes depending on pollen water status and environmental conditions, pollen recognition through molecular signals between pollen coat proteins and stigma exudate components, and germination initiation with pollen tube emerging through aperture penetrating stigma surface. Receptivity timing shows onset within 6-24 hours after flower opening with exact timing affected by environmental conditions particularly temperature, peak receptivity typically 1-3 days after opening when exudate production maximizes, and persistence 4-7 days though declining progressively as stigma senesces if pollination fails. Environmental effects on receptivity include temperature affecting metabolic rate with optimal 20-28°C supporting vigorous receptivity development, cool conditions below 15°C delaying receptivity onset and reducing function, and excessive heat above 35°C potentially impairing stigma function through protein denaturation or excessive desiccation. Humidity affects receptivity through effects on exudate properties with moderate 45-70% relative humidity maintaining appropriate exudate consistency for pollen capture and hydration, high humidity above 80% potentially diluting exudate reducing effectiveness, and low humidity below 35% causing rapid evaporation potentially desiccating stigma surface. Rain effects on receptivity are severely negative with water washing exudate from stigma surface eliminating pollen capture capacity, preventing pollen germination through excessive dilution, and promoting fungal growth on wet stigma tissues potentially interfering with function. Metabolic activity in receptive stigmas shows elevated respiration supporting active secretion and cellular function, protein synthesis producing exudate components and enzymes supporting pollen recognition and tube growth, and carbohydrate metabolism providing energy and secretory substrates. Nutritional requirements during receptivity development emphasize nitrogen for protein synthesis supporting exudate production, boron critically important for stigma function with deficiency causing poor receptivity and reduced fruit set, and adequate micronutrients supporting enzyme function in recognition and response pathways. Hormonal regulation of receptivity involves possible auxin effects on stigma development and secretory activity, cytokinins potentially maintaining stigma viability during receptive period, and ethylene possibly involved in stigma senescence terminating receptivity if pollination fails. Pollen availability timing relative to receptivity onset is critical with early-opening flowers showing slight delay between receptivity onset and pollen availability from own or neighboring anthers creating brief gap, while in peak anthesis abundant pollen becomes available immediately or precedes receptivity ensuring pollination opportunity. Pollination success dependence on receptivity shows strong positive relationship with vigorous receptive stigmas showing high fruit set 60-80% under favorable conditions, while impaired receptivity from environmental stress or nutritional deficiency shows reduced set 20-40% even with adequate pollen availability. Stigma position relative to anthers affects pollination efficiency with stigma typically positioned slightly below anthers in opened flower facilitating pollen capture from anthers in same flower or wind-dispersed pollen from neighboring flowers, with some flowers showing stigma exsertion above anthers affecting self versus cross pollination patterns. Receptivity assessment methods include visual observation of stigma exudate appearance providing qualitative indicator, controlled pollination experiments applying known pollen to stigmas at various times determining functional receptivity period, and molecular markers of receptivity-associated gene expression providing detailed developmental characterization. The process represents third of three early anthesis aspects establishing female reproductive function, developing concurrently with initial cap shedding as flowers open and expose organs. Onset typically occurs within 1 day of flower opening representing functional lag as stigma completes maturation, with rapid transition to receptive state. Cultivar variation appears in receptivity development timing with some varieties showing rapid onset within hours while others show delayed 24-48 hour onset, receptivity duration with some showing brief 3-4 day period while others maintain 6-8 day receptivity, and exudate production capacity affecting pollen capture efficiency. Regional timing shows stigma receptivity onset during early June in warm climates concurrent with first flowers opening, early-mid June in temperate regions, and mid-June in cool areas, with timing critical for ensuring overlap with pollen availability. Viticultural significance includes establishing functional female reproductive capacity essential for fruit set, requiring optimal environmental conditions particularly temperature and humidity for normal development, representing vulnerable period for weather-related fruit set failure, and interacting with pollen availability timing determining pollination success and yield. Research applications include analyzing molecular mechanisms of receptivity development including gene expression and protein synthesis, investigating environmental effects on receptivity affecting fruit set outcomes, evaluating nutritional requirements particularly boron for normal function, quantifying temporal dynamics of receptivity relative to pollen availability, and assessing genetic variation in receptivity characteristics affecting reproductive success.
Stigma_Receptivity_Onset is third of three Early_Anthesis aspects, distinguished by functional female reproductive development. Develops concurrently with Initial_Cap_Shedding following flower opening. Represents essential functional requirement for fruit set. Timing relative to pollen availability critically affects pollination success. Environmental conditions particularly temperature and humidity strongly affect development and function. Boron nutritional status critically important.
Empfängnisbereitschaft
Narbenfunktionsbeginn
Narbenrezeptivität
Stigmareifung
receptivity initiation
stigma functionality onset
stigma maturation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 368-375 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 282-287 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Stigma Receptivity Onset
Merlot in Bordeaux shows stigmatic exudate binding during early-mid June Year N+1 at BBCH 60-65, with lipid transfer proteins in exudate binding pollen coat lipids creating molecular bridges within 15-35 minutes of contact, polysaccharide matrix contributing to viscous adhesion, cumulative molecular interactions creating binding strength resisting 200-400 times pollen grain weight, under oceanic conditions of 21-26°C temperature, typical of this mid-season cultivar with adequate boron nutrition supporting normal exudate production enabling robust molecular binding and adhesion
Stigmatic exudate binding represents the molecular adhesion mechanism where stigma-secreted proteins, polysaccharides, and lipids interact with pollen coat substances creating specific high-affinity molecular bridges that secure pollen grain attachment and mediate recognition processes enabling successful pollination. This critical molecular binding process occurs during pollen adhesion in early-mid June in temperate regions Year N+1, developing over 10-45 minutes following initial contact as molecular interactions strengthen, corresponding to BBCH 60-65 flowering stages. Binding mechanism involves multiple molecular interactions including protein-protein binding between pollen coat adhesins and stigma exudate receptors creating direct protein bridges, protein-lipid interactions with stigma lipid transfer proteins binding pollen coat lipids mediating both adhesion and recognition, polysaccharide-polysaccharide interactions between pollen wall polymers and exudate carbohydrates, and integrated multi-molecular binding network creating robust attachment. Stigmatic exudate composition enabling binding includes proteinaceous components representing 30-50% dry weight with lipid transfer proteins as major constituent mediating pollen binding, structural glycoproteins contributing to matrix formation and possibly providing binding sites, polysaccharides representing 20-40% composition creating viscous matrix and carbohydrate binding sites, lipids 5-15% affecting surface properties and possibly involved in recognition, and water comprising 60-75% fresh weight providing hydration medium for molecular interactions. Lipid transfer protein function shows central role in binding with proteins abundant in grapevine stigmatic exudate typically 20-40% total protein, proteins binding pollen coat lipids through hydrophobic pocket creating strong molecular bridge, lipid-binding specificity potentially contributing to species recognition, and proteins serving dual adhesion and recognition functions critical for pollination success. Pollen coat substances participating in binding include proteins from tapetal origin coating grain surface providing binding partners for exudate proteins, lipids including tryphine components representing major constituent binding lipid transfer proteins creating strong attachment, polysaccharides in exine wall participating in carbohydrate interactions, and sporopollenin polymers possibly contributing to binding through surface properties. Protein-protein binding characteristics show high-affinity interactions with dissociation constants in nanomolar to micromolar range indicating strong binding, specificity enabling species and compatibility recognition, and multiple binding sites per grain creating robust multi-point attachment resisting detachment forces. Protein-lipid binding through lipid transfer proteins shows hydrophobic interactions with protein internal cavity binding pollen coat lipids, binding affinity in low micromolar range creating strong associations, and lipid-binding specificity potentially discriminating between compatible and incompatible pollen though grapevine shows self-compatibility. Polysaccharide binding mechanisms involve carbohydrate-carbohydrate interactions between pollen wall polysaccharides and exudate carbohydrates, possible lectin-mediated binding with exudate proteins recognizing pollen wall sugar residues, and hydrogen bonding networks creating stable though individually weak interactions that collectively contribute substantial binding strength. Molecular binding kinetics show progressive strength development with initial weak binding within seconds of contact, strengthening over minutes as molecular interactions develop and multiply, and reaching maximum binding within 15-45 minutes creating stable robust attachment. Binding specificity enabling recognition shows molecular interactions mediating species identification through complementary binding determinants, self-compatibility in grapevine showing acceptance of self-pollen through compatible molecular signatures, and recognition triggering downstream responses including enhanced exudate secretion and pollen hydration support. Binding affinity characteristics show strong associations with multiple high-affinity interactions creating robust attachment, cumulative binding strength from numerous molecular bridges exceeding requirements for stable adhesion, and binding stability maintaining attachment throughout pollen hydration, germination, and tube growth phases. Exudate matrix role in binding provides physical scaffold embedding pollen grain, viscous network distributing mechanical forces across multiple binding sites preventing localized stress concentrations, and hydrated gel-like environment maintaining appropriate conditions for molecular interactions and pollen function. Binding site density on pollen surface shows numerous potential interaction sites with pollen coat covering grain surface providing extensive binding opportunity, estimated hundreds to thousands of individual molecular interactions per grain creating redundant robust attachment, and multiple binding points distributing adhesion force preventing single-point failure. Environmental effects on binding include temperature affecting molecular interaction kinetics with optimal 22-28°C supporting rapid binding development, extreme temperatures potentially affecting binding affinity though normally stable across flowering temperature range 15-32°C, humidity influencing exudate water content affecting molecular accessibility and diffusion rates with moderate 50-65% optimal, and pH possibly affecting binding characteristics though exudate buffering maintains appropriate range. Temperature effects on binding kinetics show warm conditions 24-28°C promoting faster binding development through enhanced molecular diffusion and interaction rates, cool conditions 18-22°C slowing binding though ultimately reaching similar strength, and excessive heat above 33°C potentially affecting protein stability though normally not encountered during optimal flowering conditions. Boron effects on exudate and binding show critical importance with adequate boron essential for normal exudate production affecting binding substrate availability, boron possibly affecting exudate composition particularly polysaccharide components influencing binding properties, and deficiency reducing binding capacity through quantitative and qualitative exudate effects severely impairing pollination success. Binding as reversible process shows early binding potentially reversible before full molecular network develops, transition to effectively irreversible state within 15-30 minutes as numerous high-affinity interactions establish, and mature binding requiring extreme conditions for disruption normally not encountered during successful pollination. Failed binding causing adhesion failure results from inadequate exudate production reducing binding substrate availability, abnormal exudate composition lacking essential binding components particularly lipid transfer proteins, pollen coat abnormalities compromising binding partner availability, or environmental interference particularly rain diluting exudate preventing adequate molecular interactions. The process represents first aspect of pollen adhesion emphasizing molecular binding mechanisms, followed by pollen grain attachment discussing overall stable association. Duration develops over 10-45 minutes as molecular interactions multiply and strengthen. Cultivar variation in binding may include lipid transfer protein content and isoform composition affecting binding properties, overall exudate composition affecting binding mechanisms, and pollen coat characteristics affecting binding partner availability and quality. Regional timing shows stigmatic exudate binding occurring during flowering period across all climates following pollen-stigma contact, developing as molecular interactions establish within minutes of initial encounter. Viticultural significance includes representing essential molecular mechanism securing pollen attachment, requiring adequate stigma exudate production and composition, demonstrating critical importance of boron nutrition for normal binding through exudate effects, and establishing binding success as molecular foundation for subsequent recognition, germination, and fertilization. Research applications include analyzing specific molecular interactions in binding using biochemical and molecular biology techniques, identifying binding proteins and their pollen partners, evaluating exudate composition effects on binding properties, investigating boron effects on exudate and binding function, quantifying binding kinetics and affinity characteristics, and assessing relationships between molecular binding and pollination success outcomes.
Stigmatic_Exudate_Binding is first aspect of Pollen_Adhesion, distinguished by molecular mechanism emphasis. Represents specific high-affinity protein-lipid-polysaccharide interactions securing attachment. Critical role of lipid transfer proteins in grapevine. Develops over 10-45 minutes as molecular network strengthens. Boron nutrition essential for normal exudate production and binding. Enables species recognition and compatibility assessment. Followed by Pollen_Grain_Attachment discussing overall stable association. Molecular foundation for successful pollination.
Adhäsionsmolekül-Interaktion
Exsudat-Pollen-Bindung
adhesion molecule interaction
exudate-pollen binding
molecular pollen adhesion
molekulare Pollenadhäsion
stigma protein binding
stigmatische Exsudat-Bindung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 380-383 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Stigmatic Exudate Binding
Stilbene content quality in Pinot Noir from Burgundy reflects the cultivar's thin-skinned susceptibility to Botrytis and consequent phytoalexin induction, with stilbene concentrations in healthy fruit of 2-8 mg/kg increasing to 15-50 mg/kg following noble rot infection this elevated stilbene content in botrytized vintages contributes to the health-associated compound profile that has generated consumer and research interest in red wine polyphenols, while also indicating the vintage conditions and fruit health status relevant to wine style and aging potential.
Stilbene_Content_Quality encompasses the chemical qualities characterizing the concentration and composition of stilbenoid compounds in grape berry tissue, capturing the C6-C2-C6 phytoalexins that accumulate in response to stress and pathogen challenge while providing significant health-associated bioactivities. Stilbenes are synthesized through the action of stilbene synthase on p-coumaroyl-CoA and malonyl-CoA substrates, producing the resveratrol backbone that can be further modified through hydroxylation, methylation, and glycosylation to generate the diversity of grape stilbenes. The primary grape stilbenes include trans-resveratrol, its glucoside trans-piceid, the methylated derivative pterostilbene, and the dimeric viniferins formed through oxidative coupling. Stilbene accumulation is strongly induced by fungal infection, particularly by Botrytis cinerea, UV radiation, and other stress factors, reflecting their function as phytoalexins providing defense against pathogens. Healthy grapes contain relatively low stilbene concentrations (typically 1-10 mg/kg fresh weight), while infected or stressed grapes may accumulate 10-100 fold higher levels depending on the inducing stimulus and cultivar susceptibility. This stress-responsive accumulation creates quality complexity, as elevated stilbene content may indicate both enhanced health-relevant compound concentration and potential quality issues from the inducing stress. The health interest in stilbenes, particularly resveratrol, stems from their documented antioxidant, anti-inflammatory, and potential cardioprotective effects, making grape and wine stilbene content relevant to functional food research and wine marketing based on health associations.
Stilbene_Content_Quality serves as the parent class for specific stilbene compounds including Resveratrol_Content_Quality. The phytoalexin function of stilbenes means their concentration indicates stress history as well as current composition. The distinction between constitutive (basal) and induced stilbene levels is important for interpretation very high levels suggest significant stress exposure. Stilbene analysis employs HPLC with UV detection at 306-320 nm or fluorescence detection for enhanced sensitivity, with individual compounds quantified against authentic standards.
Phytoalexin-Phenolqualität
Resveratrol-Familienkonzentration
Stilbengehalt
Stilbenoid-Gehalt
phytoalexin phenolic quality
resveratrol family concentration
stilbene compound quality
stilbenoid content
Langcake P and Pryce RJ (1977) A New Class of Phytoalexins from Grapevines. Experientia 33(2): 151-152 Jeandet P et al. (2002) Phytoalexins from the Vitaceae: Biosynthesis, Phytoalexin Gene Expression in Transgenic Plants, Antifungal Activity, and Metabolism. Journal of Agricultural and Food Chemistry 50(10): 2731-2741
Stilbene Content Quality
Molecular characterization of Stilbene_Synthase_Induction_Disposition at INRAE Colmar employed qRT-PCR analysis of multiple STS gene family members across 15 cultivars following Plasmopara viticola inoculation. Expression of five representative STS genes (VvSTS6, VvSTS16, VvSTS27, VvSTS36, VvSTS41) was monitored at 0, 6, 12, 24, 48, and 72 hours post-inoculation using gene-specific primers. Total STS induction (sum of all five genes) showed dramatic cultivar variation in both kinetics and magnitude. Resistant 'Regent' exhibited rapid Stilbene_Synthase_Induction_Disposition with detectable upregulation by 6 hpi, peak expression at 18 hpi (125-fold above basal for VvSTS27, the most responsive gene), and sustained elevation through 72 hpi. Susceptible 'Chasselas' showed delayed, weak induction with peak at 48 hpi reaching only 12-fold for VvSTS27. Interestingly, individual STS genes showed cultivar-specific induction patterns: VvSTS41 was the dominant induced gene in 'Solaris' (85-fold) but only moderately induced in 'Regent' (25-fold), suggesting genotype-specific regulatory variation within the STS family. Correlation analysis revealed that STS induction magnitude at 24 hpi was the strongest predictor of final stilbene accumulation (r = 0.89) and resistance phenotype (r = -0.72 with sporulation severity), confirming that Stilbene_Synthase_Induction_Disposition is a key determinant of effective stilbene-based defense.
A Stilbene_Synthase_Induction_Disposition is a Phytoalexin_Production_Disposition that inheres in a Vitis vinifera plant and is realized through rapid and strong transcriptional activation of stilbene synthase (STS) genes in response to pathogen attack, elicitor treatment, or abiotic stress, providing the enzymatic capacity for de novo stilbene phytoalexin biosynthesis. This disposition represents the regulatory component of stilbene-based defense, as STS enzyme levels are typically low in unchallenged tissues and must be induced to enable substantial phytoalexin accumulation. Stilbene synthase catalyzes the condensation of one molecule of p-coumaroyl-CoA with three molecules of malonyl-CoA to produce resveratrol, the foundational stilbene from which more complex phytoalexins (viniferins, pterostilbene) are derived. Grapevine possesses an exceptionally large STS gene family comprising approximately 40 functional genes organized in clusters on chromosomes 10 and 16, reflecting ancient gene duplication events that have expanded stilbene biosynthetic capacity in this genus. Different STS genes show distinct expression patterns, with some induced by pathogens, others by UV light, and some showing tissue-specific expression, together providing flexible and robust stilbene production capacity across diverse challenge conditions. The speed and magnitude of STS induction vary among cultivars, with resistant genotypes typically showing earlier peak expression (12-24 hours post-challenge) and higher fold-induction (50-200 fold) compared to susceptible cultivars (peak at 36-48 hours, 5-20 fold induction).
Stilbene_Synthase_Induction_Disposition is_a Phytoalexin_Production_Disposition. STS induction represents the regulatory control point for stilbene phytoalexin production, determining the enzymatic capacity available for resveratrol synthesis. The large STS gene family in grapevine (~40 genes) provides redundancy and flexibility in induction responses. Different STS genes respond to different stimuli and show cultivar-specific expression patterns, creating complex genotype × environment interactions in stilbene production. STS induction kinetics (speed) and magnitude (fold-change) are partially independent parameters both contributing to effective defense.
STS gene induction
STS transcriptional response
STS-Gen-Aktivierung
Stilbensynthase-Induktions-Disposition
Stilbensynthase-Transkriptions-Antwort
stilbene synthase activation
Chong J et al. (2009) The stilbene synthase gene family in grapevine: genome-wide identification and expression analyses during development and stress responses, BMC Plant Biology 9:54 Vannozzi A et al. (2012) Genome-wide analysis of the grapevine stilbene synthase multigenic family: genomic organization and expression profiles upon biotic and abiotic stresses, BMC Plant Biology 12:130 Höll J et al. (2013) The R2R3-MYB transcription factors MYB14 and MYB15 regulate stilbene biosynthesis in Vitis vinifera, Plant Cell 25:4135-4149
Stilbene Synthase Induction Disposition
Still wine specification for Grüner Veltliner Smaragd from the Wachau documents a dry white wine targeting 13.0-14.5% alcohol (minimum 12.5% for Smaragd classification per Vinea Wachau statutes), residual sugar ≤4 g/L, TA 5.5-7.0 g/L, pH 3.10-3.35, fermentation in stainless steel or large neutral oak (2500-5000 L Stockinger casks), aging 6-12 months on fine lees with periodic bâtonnage, bottled unfined and unfiltered for the premium tier — the specification reflecting the Wachau philosophy of expressing site character through minimal intervention winemaking. Still wine specification for Cabernet Sauvignon from Coonawarra defines a full-bodied dry red wine at 13.5-14.5% alcohol, TA 6.0-6.8 g/L, pH 3.45-3.65, maceration 14-21 days with twice-daily pump-overs, malolactic fermentation in barrel, aging 18-24 months in French oak (40-60% new, predominantly Allier and Tronçais cooperage), and final blend incorporating up to 10% Merlot or Cabernet Franc for complexity.
Still wine specification is a wine style description entity that documents the compositional, sensory, and production parameters defining non-effervescent table wines produced through standard alcoholic fermentation without secondary carbonation or spirit fortification, encompassing specifications for white, rosé, red, and orange (skin-contact white) still wines including target alcohol content, residual sugar level, acid balance, phenolic structure, oak treatment regime, aging duration, and the cultivar blending proportions, and representing the dominant wine category by global production volume that accounts for approximately 85% of world wine output.
Still wine specification encompasses the broadest and most diverse category in the wine specification hierarchy, ranging from simple, early-drinking commercial wines produced with minimal intervention to complex, age-worthy grand cru expressions requiring years of cellar maturation. The fundamental parameters defining still wine style include alcohol content (ranging from 8% for light German Kabinett to 16% for full-bodied warm-climate Zinfandel), residual sugar (from <1 g/L bone-dry to 45+ g/L for off-dry Spätlese styles within the still wine category), acid-pH balance (determining freshness perception and microbial stability), phenolic concentration and structure (determining body, texture, and aging capacity in red wines), and oak influence (determining the aromatic and textural contribution of cooperage). The color classification within still wines — white (pressed before or with minimal skin contact), rosé (limited skin contact of 2-24 hours or saignée method), red (extended skin contact through fermentation), and the emerging orange/amber category (white grapes with extended skin maceration of days to months, traditional in Georgia and gaining global interest) — represents fundamentally different processing approaches with distinct implications for grape composition requirements. Regulatory frameworks impose specific parameters on still wines seeking geographical indication status — EU PDO regulations specify maximum yields, minimum alcohol, permitted cultivars, and production methods, while the Austrian DAC system and Italian DOCG rules similarly constrain still wine specifications to preserve regional typicity. The trend toward natural, minimal-intervention winemaking is challenging conventional still wine specification frameworks by producing wines that may not conform to traditional style categories — skin-contact whites, co-fermented field blends, and unfiltered natural wines expand the stylistic boundaries of the still wine category.
Stillweinparameter
Stillweinspezifikation
Tafelweinspezifikation
dry wine specification
nichttschäumende Weinspezifikation
non-sparkling wine specification
ruhiger Wein Spezifikation
still wine parameters
table wine specification
tranquil wine specification
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1: The Microbiology of Wine and Vinifications, 2nd Ed, Wiley, pp.: 327-395 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 647-720 OIV (2022) International Code of Oenological Practices, Part II: Wines
Still Wine Specification
Stomatal closure sensitivity quality comparison in Priorat evaluated the practical implications of cultivar differences for deficit irrigation management under Mediterranean summer conditions. Garnacha (Grenache) demonstrated high stomatal closure sensitivity quality with the water potential at 50% conductance reduction (Ψ₅₀) occurring at -0.35 MPa predawn, indicating early initiation of water-conserving behavior that protected hydraulic integrity but limited carbon assimilation under developing stress. Cariñena (Carignan) showed intermediate sensitivity with Ψ₅₀ at -0.55 MPa, while Syrah exhibited low stomatal closure sensitivity quality with Ψ₅₀ not reached until -0.75 MPa, maintaining substantial photosynthesis under stress levels that had largely shut down conductance in Garnacha. The stomatal closure sensitivity quality differences translated to practical management implications, with high-sensitivity Garnacha requiring irrigation at milder stress levels to maintain acceptable carbon assimilation for fruit ripening, while low-sensitivity Syrah tolerated extended deficits but with corresponding water use that depleted soil reserves more rapidly. Optimal irrigation strategies were cultivar-specific, with Garnacha managed at target predawn potential of -0.3 MPa maintaining 60% of maximum conductance versus Syrah managed at -0.6 MPa achieving equivalent 60% relative conductance, demonstrating how stomatal closure sensitivity quality calibration enables cultivar-appropriate deficit management.
Stomatal closure sensitivity quality is a stomatal response to drought quality that inheres in a grapevine cultivar or clone and represents the threshold of water deficit at which significant stomatal closure initiates and the rate at which conductance declines as stress intensifies, determining how readily the vine sacrifices photosynthetic capacity for water conservation when soil water availability begins to limit supply relative to atmospheric demand. This quality captures the triggering sensitivity of the stomatal drought response, distinguishing cultivars that close stomata at early stress stages from those maintaining stomatal opening until more severe stress develops, with this variation having fundamental implications for productivity, water use, and stress tolerance under deficit conditions. The ontological significance of stomatal closure sensitivity quality lies in its role determining the onset of the photosynthesis-transpiration tradeoff under developing water deficit, with high-sensitivity cultivars beginning to sacrifice carbon assimilation at mild stress while low-sensitivity types maintain photosynthesis longer but with greater water loss and risk of hydraulic damage if stress intensifies beyond their tolerance capacity. The biological basis of stomatal closure sensitivity encompasses the threshold responsiveness of guard cells to the signals triggering closure, including the sensitivity to root-sourced ABA, the hydraulic signals from declining leaf water status, and the atmospheric vapor pressure deficit signals affecting guard cell water relations, with variation in receptor sensitivity and signal transduction determining how readily these signals induce closure. The quantitative characterization of stomatal closure sensitivity employs the water potential threshold at which conductance declines to specified fractions of maximum (commonly 50%), the slope of the conductance-versus-water-potential relationship through the responsive range, and the minimum conductance maintained under severe stress, collectively defining the stomatal response curve that characterizes cultivar-specific drought behavior.
Stomatal_Closure_Sensitivity_Quality is_a Stomatal_Response_To_Drought_Quality, representing the threshold and rate of stomatal closure under developing stress. This class addresses how readily stomata begin to close in response to water deficit. The Ψ₅₀ metric (water potential at 50% conductance) provides quantitative comparison among cultivars. Distinguished from Isohydric_Behavior_Quality which addresses the overall water potential regulation strategy rather than specifically the closure threshold. High sensitivity protects hydraulics but limits carbon assimilation low sensitivity sustains photosynthesis but risks hydraulic damage.
Stomata-Schließungs-Sensitivität
Stomata-Verschluss-Sensitivität
Trockenheits-Schließschwelle
closure onset sensitivity
drought closure threshold
stomatal response sensitivity
stomatal sensitivity threshold
stomatäre Empfindlichkeitsschwelle
Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell and Environment 26:1393-1405 Tombesi S et al. (2015) Stomatal closure is induced by hydraulic signals and maintained by ABA in drought-stressed grapevine. Scientific Reports 5:12449 Hochberg U et al. (2013) The relationship between leaf hydraulic vulnerability to cavitation and stomatal regulation in Mediterranean Aleppo pine. New Phytologist 198:105-115
Stomatal Closure Sensitivity Quality
Maximum stomatal conductance (gs) in well-watered grapevines ranges from 150-450 mmol H₂O/m²/s across cultivars, with Grenache and Semillon at the high end (350-450 mmol/m²/s) and Chardonnay and Pinot Noir at the low end (150-250 mmol/m²/s). Stomatal density varies from 150-350 stomata/mm² on abaxial leaf surfaces (amphistomatous leaves show 10-30% of adaxial density). Guard cell response to ABA shows genotypic variation, with sensitive cultivars reducing gs by 50% at leaf ABA concentrations of 500-800 ng/g FW while insensitive types require 1200-1800 ng/g FW for equivalent closure. Stomatal patchiness (heterogeneous aperture across the leaf) increases under stress, causing gs measurements to underestimate average conductance by 10-30%.
A transpiration disposition that is the inherited capacity of a Vitis vinifera plant to regulate gas exchange through stomatal pores with genotype-specific conductance levels and response patterns, determined by stomatal density, maximum aperture size, guard cell sensitivity to environmental and hormonal signals, and the coordination of stomatal behavior across the leaf surface.
Stomatal_Conductance_Disposition serves as the parent class for high and low conductance categories. Stomatal conductance represents the primary control point for the trade-off between carbon gain (photosynthesis) and water loss (transpiration). The trait integrates genetic programming with environmental responsiveness, showing both constitutive differences among genotypes and plastic responses to light, humidity, CO₂, and water status. Coordination between stomatal and hydraulic conductance determines the operating point for leaf water potential.
Spaltöffnungsregulation
Stomata-Leitfähigkeit
gs trait
stomatal aperture regulation
stomatal opening capacity
stomatäre Leitfähigkeit
Rogiers SY et al. (2009) Does night-time transpiration contribute to anisohydric behaviour in a Vitis vinifera cultivar? Journal of Experimental Botany 60:3751-3763 Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell and Environment 26:1393-1405
Stomatal Conductance Disposition
Stomatal conductance monitoring comparing isohydric (Grenache) and anisohydric (Syrah) cultivars under developing water stress reveals contrasting regulatory strategies, with Grenache showing rapid stomatal closure (gs declining from 0.28 to 0.08 mol/m²/s as stem water potential drops from -0.6 to -1.2 MPa) maintaining leaf water status but sacrificing carbon gain, while Syrah maintains higher conductance (0.24 mol/m²/s at -1.2 MPa) sustaining photosynthesis but allowing leaf water potential to decline, demonstrating how stomatal conductance regulation determines the cultivar-specific balance between water conservation and carbon acquisition under drought.
Stomatal conductance quality is a transpiration rate quality that represents the degree of stomatal opening expressed as the conductance of the stomatal pathway to gas diffusion, characterizing the regulated resistance that controls both water vapor efflux and CO₂ influx and thus determines the balance between transpiration and photosynthesis under varying environmental and physiological conditions. Within the BFO framework, this quality specifically addresses the stomatal regulation dimension of gas exchange, distinguishing the actively controlled stomatal pathway from passive cuticular conductance while recognizing that stomatal conductance is the primary mechanism through which grapevines regulate their water loss and carbon gain in response to environmental signals and internal water status. The ontological significance lies in stomatal conductance providing the key regulatory control point linking atmospheric conditions, soil water availability, and vine carbon-water balance, where guard cell behavior integrating multiple signals (light, CO₂, humidity, water status, hormones) determines the aperture that simultaneously controls transpiration rate and CO₂ supply for photosynthesis. The physical basis for stomatal conductance involves the diffusion of gas molecules through stomatal pores whose aperture is controlled by guard cell turgor, with conductance (g, mol/m²/s) representing the inverse of resistance and calculated from measured gas flux divided by the driving concentration gradient. The measurement of stomatal conductance employs porometry (direct measurement of leaf vapor conductance) or gas exchange systems (calculating conductance from transpiration rate and vapor pressure gradient), with values for grapevine leaves typically ranging from 0.05-0.40 mol/m²/s depending on environmental conditions and vine water status. The regulatory mechanisms controlling stomatal conductance include hydropassive responses (direct turgor effects), hydroactive responses (metabolic responses to water status), ABA signaling (stress hormone closing stomata), light responses (blue light-stimulated opening), and CO₂ responses (elevated CO₂ promoting closure), with these mechanisms integrating to optimize the balance between carbon gain and water loss. Understanding stomatal conductance quality enables assessment of vine water stress response, prediction of photosynthesis from stomatal limitation, and evaluation of cultivar differences in water use strategy (isohydric versus anisohydric behavior).
Stomatal_Conductance_Quality serves as the parent class for specific conductance parameters (Maximum, Operational, Sensitivity). This quality represents the primary regulatory control of the carbon-water trade-off in grapevine physiology. The class supports water stress assessment and cultivar characterization through stomatal conductance evaluation.
Blattleitfähigkeit
Stomataöffnung
gs-Wert
stomatal aperture
stomatal opening
Buckley TN (2005) The control of stomata by water balance. New Phytologist 168(2): 275-292 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data. Annals of Botany 105(5): 661-676
Stomatal Conductance Quality
Investigation of Stomatal_Density_Disposition effects on downy mildew susceptibility at INRAE Bordeaux quantified stomatal parameters and P. viticola infection across 22 cultivars under controlled inoculation conditions. Stomatal density was determined by microscopy on cleared abaxial leaf surfaces, with parallel measurements of stomatal aperture dimensions and calculated stomatal conductance index. Density ranged from 118 stomata/mm² ('Riesling') to 312/mm² ('Merlot'), representing a 2.6-fold variation among commercial cultivars. Controlled inoculation with P. viticola sporangia suspension revealed significant correlation between Stomatal_Density_Disposition and infection establishment (r = 0.71), with low-density 'Riesling' (118/mm²) showing 52% fewer infection sites per unit leaf area than high-density 'Merlot' (312/mm²) at equivalent inoculum concentration. When infection was expressed per stomata (site-specific infection probability), cultivar differences diminished substantially (range 0.12-0.18 infections/stomata), confirming that stomatal density rather than stomatal-specific resistance explained much of the cultivar variation. Microscopic observation confirmed that zoospores encysted preferentially near stomata regardless of cultivar, with penetration success per stomatal encounter similar across genotypes. These findings demonstrate that Stomatal_Density_Disposition contributes meaningfully to quantitative downy mildew resistance through a simple geometric mechanism of reduced entry point availability.
A Stomatal_Density_Disposition is a Physical_Barrier_Disposition that inheres in a Vitis vinifera plant leaf and is realized through the developmental specification of stomatal frequency per unit leaf area, affecting the number of potential entry points for stomatal-penetrating pathogens and influencing leaf gas exchange characteristics. This disposition represents the quantitative aspect of stomatal patterning, with density varying substantially among grapevine cultivars (typically 100-350 stomata/mm² on abaxial surfaces) due to genetic differences in stomatal initiation and spacing during leaf development. Stomata, the pores formed by paired guard cells that regulate gas exchange, serve as primary infection courts for several important grapevine pathogens, most notably Plasmopara viticola (downy mildew), which releases zoospores that swim to stomata, encyst, and penetrate through the stomatal pore to initiate infection. Higher stomatal density provides more potential entry points per unit leaf area, increasing the probability of successful pathogen establishment following inoculation. Beyond direct entry point considerations, stomatal density influences leaf transpiration rate, boundary layer humidity, and water status under stress, creating complex interactions between Stomatal_Density_Disposition and disease susceptibility. Breeding for reduced stomatal density could theoretically enhance downy mildew resistance, though trade-offs with photosynthetic capacity and water use efficiency must be considered, and the magnitude of density effects relative to other resistance factors determines practical breeding value.
Stomatal_Density_Disposition is_a Physical_Barrier_Disposition. Stomatal density specifically affects pathogens using stomatal penetration, primarily Plasmopara viticola among major grapevine diseases. Density is genetically controlled and environmentally stable, making it a reliable selection target. Trade-offs with gas exchange capacity and potential effects on water use efficiency must be considered in breeding applications. The disposition addresses stomatal number rather than stomatal function or regulation, which involve different genetic and physiological systems.
Spaltöffnungsdichte-Disposition
Stomata-Dichte
Stomatäre-Frequenz
guard cell density
stomatal frequency
stomatal patterning
Díez-Navajas AM et al. (2008) Comparative study of the role of stomata in susceptibility of wild grapevine (Vitis vinifera subsp. sylvestris) to downy mildew, VITIS Journal of Grapevine Research 47:21-26 Hetherington AM, Woodward FI (2003) The role of stomata in sensing and driving environmental change, Nature 424:901-908 Gindro K et al. (2003) Cellular and molecular changes in grapevine (Vitis vinifera) leaves in response to infection by Plasmopara viticola, VITIS Journal of Grapevine Research 42:225-234
Stomatal Density Disposition
Stomatal limitation monitoring in deficit-irrigated Shiraz tracks the development of stomatal constraint with progressive water stress, from well-watered conditions (stem water potential -0.4 MPa, stomatal conductance 0.28 mol/m²/s, Ls = 12%) through moderate stress (-0.9 MPa, gs = 0.12 mol/m²/s, Ls = 38%) to severe stress (-1.4 MPa, gs = 0.04 mol/m²/s, Ls = 65%), demonstrating how stomatal limitation quantification enables precision irrigation scheduling that maintains target stress levels appropriate for quality fruit production while avoiding severe limitation that would compromise carbon gain and ripening capacity.
Stomatal limitation quality is a photosynthetic limitation quality that represents the constraint on photosynthesis arising from stomatal closure that restricts CO₂ diffusion from the atmosphere to the substomatal cavity, characterizing the degree to which stomatal conductance limits CO₂ supply below the level required for maximum photosynthetic rate. Within the BFO framework, this quality specifically addresses the stomatal component of diffusional limitation, distinguishing stomatal limitation from mesophyll and biochemical limitations while recognizing that stomatal closure is the primary short-term response to water deficit that reduces photosynthesis through CO₂ supply restriction in water-stressed grapevines. The ontological significance lies in stomatal limitation representing the most responsive and manageable component of photosynthetic limitation, where stomatal aperture responds within minutes to water status, vapor pressure deficit, and CO₂ concentration, enabling short-term productivity optimization through irrigation management that balances stomatal opening for carbon gain against water loss for conservation. The physiological basis for stomatal limitation involves the reduction in CO₂ concentration from ambient (Ca, approximately 400 μmol/mol) to substomatal (Ci, typically 200-300 μmol/mol in C₃ plants) due to diffusion resistance through stomatal pores, with stomatal limitation (Ls) quantified as the proportional reduction in photosynthesis attributable to this CO₂ drawdown, calculated as Ls = (A at Ci=Ca minus A at actual Ci) / A at Ci=Ca. The water status relationship with stomatal limitation shows progressive increase as water stress develops, from minimal limitation (Ls 50%) under drought, with the relationship between leaf water potential and stomatal conductance differing among cultivars (isohydric versus anisohydric behavior). The environmental factors affecting stomatal limitation beyond water status include vapor pressure deficit (stomata closing at high VPD to limit water loss), temperature (affecting guard cell metabolism), and light (stomata opening in response to blue light), with stomatal limitation varying diurnally and seasonally as these factors fluctuate. Understanding stomatal limitation quality enables irrigation management optimization, cultivar-specific stress response characterization, and productivity prediction under water-limited conditions where stomatal closure is the primary constraint.
Stomatal_Limitation_Quality addresses the stomatal component of diffusional limitation on photosynthesis. This quality is highly responsive to water status and environmental conditions, representing the primary short-term regulation of photosynthesis. The class supports irrigation management through stomatal limitation monitoring that enables targeting of specific stress levels.
CO₂ supply limitation
CO₂-Versorgungslimitation
Ls-Wert
diffusional limitation (stomatal)
stomatal constraint
stomatäre Einschränkung
stomatäre Limitation
Farquhar GD, Sharkey TD (1982) Stomatal conductance and photosynthesis. Annual Review of Plant Physiology 33(1): 317-345 Medrano H et al. (2002) Regulation of photosynthesis of C₃ plants in response to progressive drought. Journal of Experimental Botany 53(366): 1-12
Stomatal Limitation Quality
Stomatal recovery rate quality assessment in Sonoma Pinot Noir evaluated the temporal pattern of photosynthetic restoration following controlled moderate stress episodes imposed by irrigation withholding. Vines stressed to midday stem water potential of -1.2 MPa (moderate stress at the quality-benefit boundary) received irrigation restoration and were monitored for stomatal conductance recovery over the subsequent 72 hours. Recovery proceeded in two phases, with rapid initial recovery achieving 50% of pre-stress conductance within 6 hours as guard cell turgor was restored, followed by slower recovery requiring 48-60 hours to achieve 90% restoration attributed to ABA clearance and metabolic normalization. The stomatal recovery rate quality proved temperature-dependent, with recovery from equivalent stress requiring only 36 hours when overnight temperatures dropped below 15°C facilitating rapid rehydration versus 72 hours when warm nights maintained high evaporative demand. Comparison of clones revealed 20% variation in recovery half-time, with faster-recovering clones resuming normal photosynthesis and carbon accumulation more rapidly, having practical implications for fruit ripening when stress episodes occurred during the critical post-véraison period. The stomatal recovery rate quality characterization enabled prediction of cumulative photosynthesis loss from stress events, informing irrigation decisions balancing stress benefits against recovery costs.
Stomatal recovery rate quality is a stomatal response to drought quality that inheres in a grapevine organism and represents the velocity and completeness with which stomatal conductance is restored following relief of water deficit stress through irrigation or precipitation, determining how rapidly photosynthetic capacity returns to normal following stress episodes and influencing the cumulative carbon assimilation impact of transient stress events that may have relatively brief peak intensity but prolonged recovery phases. This quality captures the kinetic dimension of stomatal behavior that determines the temporal pattern of photosynthetic recovery, distinguishing cultivars that resume full stomatal opening within hours of rehydration from those requiring days to restore normal conductance despite adequate water availability. The ontological significance of stomatal recovery rate quality lies in its role determining the cumulative productivity impact of stress episodes, wherein rapid recovery limits carbon assimilation loss to the direct stress period while slow recovery extends the functional impact well beyond the period of active water deficit, potentially affecting fruit development and ripening even from stress events occurring earlier in the season. The biological basis of stomatal recovery rate encompasses the reversal of the closure-inducing signals, the clearance of accumulated ABA and other stress hormones, the restoration of guard cell turgor and ionic balance, and potentially the repair of any stress-induced damage to stomatal function or the hydraulic pathway supplying guard cells. The recovery process may proceed in phases, with rapid initial opening within hours reflecting simple turgor recovery followed by slower optimization over days as metabolic adjustments complete and residual stress signals dissipate. Cultivar differences in stomatal recovery rate quality may reflect variation in ABA metabolism rates, hydraulic pathway recovery from embolism, and the degree of stress-induced modification to stomatal function requiring repair. The measurement of stomatal recovery rate quality involves tracking conductance following rehydration from controlled stress episodes, characterizing the time course to full recovery and any hysteresis between the stress descent and recovery phases of the response.
Stomatal_Recovery_Rate_Quality is_a Stomatal_Response_To_Drought_Quality, representing the kinetics of stomatal reopening following stress relief. This class addresses the recovery phase complementing the closure sensitivity of stress onset. Recovery half-time and the duration to 90% restoration provide quantitative metrics. Distinguished from Stomatal_Closure_Sensitivity_Quality which addresses the stress-response phase rather than recovery. The biphasic recovery pattern (rapid turgor recovery + slow metabolic normalization) is characteristic. Environmental conditions, particularly temperature, influence recovery rates substantially.
Rehydratations-Reaktionsrate
Stomata-Erholungskinetik
Stomata-Erholungsrate
conductance recovery
rehydration response rate
stomatal recovery kinetics
stomatal reopening rate
stomatäre Wiederöffnungsrate
Flexas J et al. (2006) Keeping a positive carbon balance under adverse conditions: responses of photosynthesis and respiration to water stress. Physiologia Plantarum 127:343-352 Lovisolo C et al. (2008) Grapevine adaptations to water stress: new perspectives about soil/plant interactions. Theoretical and Experimental Plant Physiology 20:99-106 Chaves MM et al. (2009) Photosynthesis under drought and salt stress: regulation mechanisms from whole plant to cell. Annals of Botany 103:551-560
Stomatal Recovery Rate Quality
Detailed characterization of Stomatal_Regulation_Disposition at the University of California Davis employed diurnal gas exchange measurements across progressive drought in 20 cultivars to quantify stomatal sensitivity and response patterns. Stomatal conductance (gs) was measured hourly from 06:00 to 18:00 at soil water contents ranging from field capacity to severe deficit, with concurrent measurement of leaf water potential (Ψleaf) and xylem sap ABA concentration. Cultivars showed striking variation in Stomatal_Regulation_Disposition parameters. The response threshold (soil water potential causing initial stomatal closure) ranged from -0.3 MPa ('Grenache', sensitive) to -0.8 MPa ('Syrah', insensitive). The slope of gs versus Ψleaf relationship (stomatal sensitivity) varied 3-fold, from 85 mmol m⁻² s⁻¹ MPa⁻¹ ('Grenache') to 28 mmol m⁻² s⁻¹ MPa⁻¹ ('Syrah'). ABA sensitivity (gs reduction per unit xylem ABA increase) also differed, with 'Grenache' showing 50% gs reduction at 500 ng L⁻¹ ABA versus 1200 ng L⁻¹ for equivalent reduction in 'Syrah'. Stomatal_Regulation_Disposition had direct consequences for water use and productivity. Under moderate drought, 'Grenache' used 35% less water than 'Syrah' but also assimilated 25% less carbon. Mathematical modeling identified stomatal sensitivity parameters as key determinants of cultivar performance across climate scenarios, with high-sensitivity cultivars favored in severe drought environments and low-sensitivity cultivars advantageous where moderate stress is typical and irrigation is available.
A Stomatal_Regulation_Disposition is a Drought_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through the pattern of stomatal aperture control in response to declining soil water availability and increasing atmospheric evaporative demand, determining the balance between water conservation through reduced transpiration and continued carbon assimilation through maintained gas exchange. This disposition represents the stomatal component of drought response, fundamental to plant water relations as stomata control the primary pathway for both water loss (transpiration) and CO₂ uptake (photosynthesis). Stomatal regulation under drought involves complex integration of hydraulic signals (leaf water potential, xylem tension), chemical signals (abscisic acid from roots and leaves, pH changes in xylem sap), and atmospheric signals (vapor pressure deficit), with the relative weighting of these signals varying among genotypes to produce distinct stomatal behavior patterns. The stomatal response strategy profoundly influences whole-plant water use, carbon gain, and survival under drought, with early-closing stomata conserving water but sacrificing photosynthesis while late-closing stomata maximize carbon gain but risk severe dehydration and hydraulic failure. Stomatal_Regulation_Disposition in grapevine spans a continuum from strongly isohydric (tight stomatal control maintaining leaf water potential) to anisohydric (loose control allowing water potential to decline), with most cultivars falling between these extremes. This variation represents evolved adaptation to different moisture regimes and provides genetic diversity for breeding and cultivar selection targeting specific environments.
Stomatal_Regulation_Disposition is_a Drought_Tolerance_Disposition. Stomatal regulation represents the primary short-term response to drought, determining the water conservation versus carbon gain trade-off. Child classes address the major stomatal behavior strategies along the isohydric-anisohydric continuum. Stomatal regulation involves integration of hydraulic, chemical (ABA), and atmospheric signals, with genotypic variation in sensitivity to each. Stomatal behavior interacts with evaporative cooling capacity, as closed stomata reduce both water loss and transpirational cooling.
Stomata-Trockenheitsantwort
Stomataregulations-Disposition
Stomatäre-Leitfähigkeits-Regulation
stomatal conductance regulation
stomatal drought response
water-saving stomatal behavior
Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought, Plant Cell and Environment 26:1393-1405 Tardieu F, Simonneau T (1998) Variability among species of stomatal control under fluctuating soil water status and evaporative demand: modelling isohydric and anisohydric behaviours, Journal of Experimental Botany 49:419-432 Tombesi S et al. (2015) Stomatal closure is induced by hydraulic signals and maintained by ABA in drought-stressed grapevine, Scientific Reports 5:12449
Stomatal Regulation Disposition
Stomatal response to drought quality characterization across six Vitis vinifera cultivars in a controlled water deficit trial established the diversity of stomatal behavior patterns relevant to deficit irrigation management. Stomatal conductance was measured daily during a progressive soil drying cycle, with responses plotted against predawn leaf water potential to generate cultivar-specific response curves. Grenache exhibited the most sensitive stomatal response to drought quality, with conductance declining to 50% of well-watered values at predawn potential of only -0.25 MPa and reaching near-complete closure at -0.5 MPa, demonstrating strong isohydric behavior prioritizing hydraulic safety. Syrah showed the least sensitive response, maintaining 50% conductance until predawn potential reached -0.6 MPa and sustaining significant stomatal opening even at -0.9 MPa, exhibiting anisohydric behavior favoring continued carbon assimilation. Intermediate cultivars including Cabernet Sauvignon and Merlot showed response curves between these extremes. The stomatal response to drought quality characterization enabled cultivar-specific deficit irrigation protocols, with sensitive cultivars receiving more conservative stress targets while tolerant cultivars could be managed at greater water deficits while maintaining acceptable photosynthetic function.
Stomatal response to drought quality is a drought stress indicator quality that inheres in a grapevine organism and represents the characteristic patterns of stomatal aperture regulation exhibited in response to declining soil water availability and increasing evaporative demand, encompassing the sensitivity of stomatal closure to stress onset, the degree to which stomata close under different stress intensities, the kinetics of stomatal response to changing conditions, and the recovery behavior when stress is relieved. This quality fundamentally influences the vine's water use efficiency and drought tolerance strategy by determining the tradeoff between water conservation through stomatal closure and carbon assimilation requiring stomatal opening for CO₂ uptake, representing a primary mechanism by which grapevines balance hydraulic safety against productivity under variable water availability conditions. The ontological significance of stomatal response to drought quality lies in its role as a central determinant of vine physiological behavior under water-limited conditions, with stomatal regulation serving as the first line of defense against drought stress while simultaneously determining the degree to which carbon assimilation is maintained or sacrificed for water conservation. The biological basis of stomatal response encompasses the complex signaling network integrating hydraulic signals from root and leaf water status, chemical signals particularly abscisic acid (ABA) synthesized in response to soil drying, and atmospheric signals including vapor pressure deficit that collectively regulate guard cell turgor and stomatal aperture. The substantial genetic variation in stomatal response patterns among grapevine cultivars has been characterized along the isohydric-anisohydric continuum, with isohydric cultivars showing sensitive, early stomatal closure that maintains relatively constant leaf water potential at the cost of reduced carbon assimilation, while anisohydric cultivars maintain stomatal opening longer allowing greater water potential decline but sustaining photosynthesis and potentially achieving better performance under moderate stress. Assessment of stomatal response to drought quality employs measurements of stomatal conductance across a range of water status conditions, characterizing the response curves that define cultivar-specific drought behavior patterns.
Stomatal_Response_To_Drought_Quality is_a Drought_Stress_Indicator_Quality, serving as parent class for specific stomatal behavior characteristics including Stomatal_Closure_Sensitivity_Quality, Stomatal_Recovery_Rate_Quality, Isohydric_Behavior_Quality, and Anisohydric_Behavior_Quality. This class addresses the overall stomatal regulation under water stress while child classes capture particular response dimensions. The isohydric-anisohydric continuum represents a key conceptual framework for understanding cultivar-specific drought behavior. Critical for deficit irrigation management where stomatal behavior determines the tradeoff between water conservation and carbon assimilation.
Stomata-Reaktion auf Trockenheit
Stomata-Sensitivität
drought stomatal behavior
stomatal regulation
stomatal sensitivity
stomatäre Dürrereaktion
stomatäre Trockenheitsantwort
Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell and Environment 26:1393-1405 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data. Annals of Botany 105:661-676 Lovisolo C et al. (2008) Grapevine adaptations to water stress: new perspectives about soil/plant interactions. Theoretical and Experimental Plant Physiology 20:99-106
Stomatal Response To Drought Quality
Stomatal sensitivity characterization comparing isohydric Grenache and anisohydric Syrah reveals contrasting water use strategies, with Grenache showing high sensitivity (gs declining by 0.15 mol/m²/s per 0.1 MPa decline in stem water potential) that maintains leaf water potential above -1.2 MPa even under severe soil drought by closing stomata, while Syrah shows low sensitivity (0.04 mol/m²/s per 0.1 MPa) allowing leaf water potential to reach -1.6 MPa under similar conditions while maintaining partial stomatal opening, demonstrating how stomatal sensitivity determines the trade-off between water conservation and carbon acquisition that defines cultivar-specific drought responses.
Stomatal sensitivity quality is a stomatal conductance quality that represents the responsiveness of stomatal aperture to changes in environmental signals or plant water status, characterizing the magnitude and speed of stomatal response to perturbations that determines how rapidly and completely vines adjust gas exchange in response to changing conditions. Within the BFO framework, this quality specifically addresses the regulatory responsiveness dimension of stomatal function, distinguishing the sensitivity of stomatal response from the conductance level itself while recognizing that sensitivity determines how effectively stomata translate environmental signals into physiologically appropriate adjustments in gas exchange. The ontological significance lies in stomatal sensitivity determining the dynamic character of vine water use strategy, where high sensitivity enables rapid closure conserving water under developing stress but potentially sacrificing carbon gain, while low sensitivity maintains gas exchange during moderate stress but risks hydraulic damage under severe conditions, with this sensitivity representing a key component of the isohydric-anisohydric spectrum of water use behaviors. The multiple dimensions of stomatal sensitivity include sensitivity to water status (the gs decline per unit decline in water potential, defining isohydric versus anisohydric response), sensitivity to VPD (stomatal closure in response to atmospheric drought), sensitivity to ABA (response to stress hormone signaling), and sensitivity to light (the rate and completeness of light-stimulated opening), with each dimension potentially varying independently among cultivars and conditions. The quantitative expression of stomatal sensitivity employs various metrics including the slope of gs versus water potential relationships, the VPD at which gs declines by 50%, or the concentration of ABA causing specified stomatal closure, with these metrics enabling cultivar comparison and prediction of behavior under stress scenarios. The cultivar variation in stomatal sensitivity is substantial in Vitis vinifera, ranging from highly sensitive isohydric types (Grenache, Sangiovese) that close stomata early in drought to maintain water status, through intermediate types, to insensitive anisohydric types (Syrah, Semillon) that maintain conductance allowing water potential to decline. Understanding stomatal sensitivity quality enables prediction of cultivar behavior under drought, selection of varieties appropriate for water-limited environments, and interpretation of the water use strategies that determine vineyard water requirements under different management scenarios.
Stomatal_Sensitivity_Quality addresses the responsiveness of stomatal regulation to environmental and physiological signals. This quality determines the dynamic character of water use strategy including isohydric versus anisohydric behavior. The class supports cultivar selection and drought response prediction through stomatal sensitivity characterization.
Stomatareaktivität
gs sensitivity
gs-Empfindlichkeit
regulatorische Reaktivität
regulatory sensitivity
stomatal reactivity
stomatal responsiveness
stomatäre Sensitivität
Tardieu F, Simonneau T (1998) Variability among species of stomatal control under fluctuating soil water status and evaporative demand. Plant, Cell & Environment 21(6): 571-582 Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell & Environment 26(8): 1393-1405
Stomatal Sensitivity Quality
Cabernet Sauvignon in Napa Valley shows stomium opening during early-mid June Year N+1 at BBCH 60-65, with specialized thin-walled cell bands undergoing coordinated breakdown when humidity drops below 65% creating 2.8-3.8 mm longitudinal slits exposing pollen, stomium tissue degradation proceeding over 15-45 minutes, under warm climate conditions of 24-29°C temperature enabling efficient enzymatic and mechanical opening typical of this late-season cultivar
Stomium opening represents the specialized tissue breakdown process at predetermined rupture sites in anther walls where cells with reduced structural strength undergo coordinated degradation creating longitudinal slits enabling controlled pollen liberation. This critical tissue-level process occurs during anther dehiscence in early-mid June in temperate regions Year N+1, proceeding over minutes to hours as stomium zone cells complete breakdown following mechanical stress accumulation, as part of BBCH 60-65 flowering stages. Stomium anatomical specialization shows distinctive tissue structure with cells arranged in longitudinal bands along anther length typically 2-4 mm corresponding to locule length, cells lacking secondary wall thickening found in adjacent endothecium providing mechanical weakness, and thin-walled cellular organization creating predetermined failure pathway. Stomium cell characteristics include elongated cells oriented longitudinally facilitating longitudinal rupture propagation, thin primary walls susceptible to mechanical failure and enzymatic degradation, and reduced lignification compared to adjacent tissues making cells vulnerable to breakdown. Stomium positioning shows strategic location along anther length positioned to enable effective pollen release once opened, typically on anther face directed outward or laterally facilitating pollen dispersal, and bilateral arrangement with stomium on both locules creating paired slits for complete anther opening. Cellular breakdown mechanism involves multiple processes including enzymatic wall degradation weakening cell adhesion, mechanical stress causing physical cell separation, and possible programmed cell death contributing to tissue failure, creating coordinated stomium opening. Enzymatic activity during opening includes cellulase enzymes degrading cellulose wall components, pectinase and polygalacturonase breaking down pectin in middle lamella reducing cell adhesion, and possible xyloglucan-modifying enzymes affecting wall polysaccharides, enabling tissue separation. Mechanical forces driving opening include differential shrinkage in adjacent hygroscopic tissues creating shearing stress concentrated at stomium, turgor changes possibly contributing to stress though primary force mechanical, and accumulated stress exceeding reduced tissue strength causing failure. Opening progression shows initial localized failure at weak point in stomium zone creating nucleus for crack propagation, rapid extension along weakened tissue driven by relieved stress and continuing enzymatic activity, and bilateral opening as stomium separates exposing both locules to exterior. Slit characteristics following stomium opening include longitudinal orientation along anther length facilitating pollen release along extended pathway, width 0.3-0.8 mm providing adequate passage for pollen grains 25-35 micrometers diameter, and clean edges indicating controlled separation along predetermined pathway rather than random tearing. Pollen exposure following opening shows immediate visualization of yellow pollen mass within locules, beginning of gravitational pollen fall from opened anthers, and establishment of pathway enabling continued pollen liberation over subsequent hours. Stomium maturation requirements show proper development of specialized cellular structure necessary for functioning, completion of enzymatic systems enabling degradation, and appropriate dehydration state creating mechanical stress conditions, with premature or incomplete maturation potentially causing opening failure. Environmental triggering of opening involves humidity decrease causing tissue dehydration creating mechanical stress exceeding reduced stomium strength, temperature affecting enzymatic activity and tissue mechanical properties, and optimal conditions combining to trigger coordinated opening. Humidity effects show declining relative humidity below 70-75% typically required for sufficient mechanical stress driving stomium failure, sustained high humidity above 75% preventing adequate stress accumulation maintaining tissue integrity, and rapid humidity decrease potentially causing more synchronized opening through simultaneous stress threshold achievement. Temperature effects include warm conditions 24-30°C promoting enzymatic activity accelerating wall degradation and facilitating tissue dehydration driving mechanical stress, cool conditions 15-20°C slowing enzymatic processes and reducing dehydration potentially delaying or preventing opening, and optimal 22-28°C range supporting efficient coordinated opening. Diurnal pattern shows characteristic morning opening as humidity declines and temperature rises creating favorable conditions, peak opening activity typically 9am-12pm in temperate climates, and daily rhythm with repeated opening events across cluster as successive flowers reach appropriate conditions. Stomium opening coordination with anther wall rupture shows essentially simultaneous processes with mechanical rupture and stomium breakdown representing different aspects of unified opening mechanism, stomium tissue providing predetermined weak pathway for rupture propagation, and coordinated structural and cellular processes producing effective anther dehiscence. Failed stomium opening causes incomplete dehiscence with partial opening providing limited pollen access or complete failure preventing pollen liberation, resulting from inadequate maturation, unfavorable environmental conditions, or tissue abnormalities, and contributing to poor fruit set through impaired male function. The process represents second aspect of anther dehiscence emphasizing specialized tissue breakdown, following anther wall rupture and overlapping with initial pollen exposure. Duration spans minutes to hours as coordinated tissue breakdown proceeds. Cultivar variation in stomium characteristics may include tissue structure affecting opening efficiency, enzymatic capacity affecting degradation rate, and environmental sensitivity affecting opening reliability. Regional timing shows stomium opening during flowering period coinciding with favorable morning conditions across all climates when humidity declines and temperature rises triggering coordinated opening. Viticultural significance includes representing specialized mechanism ensuring controlled effective pollen release, requiring favorable environmental conditions for efficient functioning, and demonstrating importance of appropriate anther development for reproductive success. Research applications include analyzing stomium anatomical specialization and functional significance, investigating cellular and molecular mechanisms of tissue breakdown, evaluating environmental effects on opening efficiency, quantifying enzymatic and mechanical contributions to dehiscence, and assessing genetic variation in stomium structure and function affecting reproductive reliability.
Stomium_Opening is second aspect of Anther_Dehiscence, distinguished by focus on specialized tissue breakdown mechanism. Represents controlled failure pathway enabling effective pollen release. Involves coordinated enzymatic degradation and mechanical stress. Requires proper maturation and favorable environmental conditions. Minutes to hours duration as tissue breakdown proceeds. Overlaps with Anther_Wall_Rupture and Initial_Pollen_Exposure. Critical specialized mechanism ensuring reproductive success.
Dehiszenzzonenaufbruch
Stomium-Zusammenbruch
Stomium-Öffnung
dehiscence zone opening
specialized rupture site
spezialisierte Bruchstelle
stomium breakdown
stomium dehiscence
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 377-380 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 287-290
Stomium Opening
Stone fruit aroma assessment in Condrieu Viognier reveals the varietal signature comprising intense apricot (8.2 on 10-point scale) and white peach (7.5) with supporting nectarine notes (5.8), correlating with γ-decalactone concentration of 85 μg/L and β-damascenone at 4.2 μg/L, with the apricot dominance distinguishing Condrieu from California Viognier where peach typically predominates, demonstrating how stone fruit subcategory balance creates regional expression within the varietal framework.
Stone fruit aroma quality is a fruity aroma quality that represents olfactory perception evoking drupe fruit-associated scents including peach, apricot, nectarine, plum, and cherry notes, characterizing the soft, often sweet-associated aromatic impression commonly found in both white wines (peach, apricot) and red wines (cherry, plum) as key varietal and quality markers. Within the BFO framework, this quality specifically addresses the stone fruit subcategory of fruity aroma distinguished by its association with fruit maturity and quality perception, recognizing that stone fruit character often indicates optimal ripeness in white varieties and provides the primary fruit descriptor category for many red varieties. The ontological significance lies in stone fruit aroma serving as varietal signature for important varieties across both color categories, with peach and apricot defining Viognier, white peach characterizing quality Chardonnay, cherry forming the fruit core of Pinot Noir, and plum contributing to Merlot identity, making stone fruit descriptors among the most frequently applied in professional wine evaluation. The compositional basis for stone fruit aroma includes C13-norisoprenoids (β-damascenone contributing peach-apricot character with extremely low threshold of 0.05 μg/L), γ-lactones (γ-decalactone providing peach), certain esters (ethyl cinnamate associated with cherry), and terpene alcohols (linalool contributing to stone fruit complexity), with the specific compound balance determining which stone fruit notes predominate. The ripeness association of stone fruit character is particularly strong, with underripe fruit showing green or citrus notes while optimal maturity develops the peach, apricot, or cherry character valued in quality wines, and overripe fruit transitioning toward dried or cooked stone fruit impression. Understanding stone fruit aroma quality enables varietal identification based on characteristic stone fruit profile, ripeness assessment from stone fruit development trajectory, and quality evaluation where appropriate stone fruit expression indicates successful maturity attainment.
Stone_Fruit_Aroma_Quality addresses the stone fruit (drupe) subcategory of fruity aroma, encompassing peach, apricot, nectarine, plum, and cherry notes. This quality provides key varietal markers for both white varieties (Viognier, Chardonnay) and red varieties (Pinot Noir, Merlot). The specific stone fruit notes expressed depend on compound composition and indicate both variety identity and ripeness level.
Pfirsich-Aprikosen-Charakter
Steinobstaroma
Steinobstduft
Steinobstnoten
drupe fruit aroma
peach-apricot character
stone fruit notes
stone fruit scent
Sefton MA et al. (2011) Norisoprenoids in Vitis vinifera white wine grapes and the identification of a precursor of damascenone in these fruits. Australian Journal of Chemistry 42(12): 2071-2084 Guth H (1997) Quantitation and sensory studies of character impact odorants of different white wine varieties. Journal of Agricultural and Food Chemistry 45(8): 3027-3032
Stone Fruit Aroma Quality
Syrah storage endosperm at seed maturity (BBCH 89) contains 14-18% lipids with oleic acid 25-30% and linoleic acid 50-55% of total fatty acids, 20-24% proteins with 65-70% globulins, caloric density 5.2-5.8 kcal/g, providing reserves supporting seedling growth through 8-10 days post-germination
A storage endosperm is endosperm tissue emphasizing its composition of accumulated nutrient reserves including lipids, proteins, and carbohydrates that are synthesized and stored during seed development and subsequently mobilized to support germination and early seedling growth. Storage endosperm accumulation occurs primarily during mid- to late seed development (4-12 weeks post-fertilization, BBCH 77-87) when endosperm cells undergo= extensive lipid body formation (accumulation of triglyceride-filled oil bodies 0.5-2 μm diameter), protein body formation (accumulation of storage proteins in protein storage vacuoles 2-5 μm diameter), synthesis of storage proteins (primarily globulins 60-70% and albumins 20-30% of total protein), synthesis of storage lipids (primarily oleic acid 18= and linoleic acid 18= as triglycerides), and minimal starch accumulation (Vitis seeds are lipid-storing rather than starch-storing). Storage endosperm composition in mature Vitis vinifera seeds includes= lipids as major storage component (10-20% seed dry weight, primarily in endosperm tissue), proteins as secondary storage component (15-25% seed dry weight), cell wall polysaccharides (hemicelluloses, cellulose forming thickened cell walls that are also mobilized during germination), phytate (myo-inositol hexakisphosphate serving as phosphorus and mineral storage), and various secondary metabolites (phenolics, tannins primarily in seed coat rather than endosperm). Storage compound characteristics include= lipid composition dominated by oleic acid (C18= typically 20-30% of fatty acids) and linoleic acid (C18= typically 40-60%), protein composition dominated by legumin-like and vicilin-like globulins, caloric density approximately 5-6 kcal/g dry weight (higher than starchy seeds due to lipid content), and storage efficiency providing sufficient reserves for 7-12 days seedling growth until photosynthetic independence. Storage endosperm development is influenced by= maternal assimilate supply during seed development (carbon and nitrogen availability affecting accumulation rates), seed position within berry (seeds near berry center may receive preferential nutrition), environmental conditions during berry development (temperature, water status affecting synthesis rates), and genetic factors determining endosperm size and composition. Storage endosperm inadequacy leads to= seed abortion in extreme cases (insufficient reserves prevent embryo completion), reduced seed viability (seeds with low storage reserves show reduced germination rates 20-40% compared to 60-90% for adequate reserves), and poor seedling vigor (inadequate reserves result in slow growth and high mortality during establishment phase).
Storage_Endosperm emphasizes accumulated reserve composition and is distinguished from Nutritive_Endosperm (active support function) and Endosperm (complete tissue development) by focus on stored compounds and their metabolic mobilization potential. Critical for understanding seed quality and germination vigor, breeding for seedling establishment traits, seed storage behavior and longevity, and industrial applications utilizing grape seed oil composition.
Accumulation endosperm
Reserve endosperm
Reservegewebe
Speicher-Endosperm
Vorratsspeicherndes Nährgewebe
Storage Endosperm
Storage mobilization efficiency assessment tracking nitrogen movement from reserves to new growth in Chardonnay reveals the critical reserve contribution to early development, with permanent roots showing N content decline of 12 g N/vine between budbreak and bloom (mobilization to support shoot growth), trunk and cordons contributing additional 8 g N, such that reserve-derived N provides approximately 75% of the 27 g N accumulated in shoots by bloom, with vines having depleted reserves (from previous over-cropping) showing delayed budbreak, weak shoot growth, and poor fruit set from insufficient mobilization capacity, demonstrating how reserve status and mobilization efficiency affect current-season productivity.
Storage mobilization efficiency quality is a nutrient remobilization efficiency quality that represents the effectiveness of releasing nutrients from perennial storage tissues to support growth demands when soil uptake is insufficient or phenologically unavailable, characterizing the reserve mobilization that enables spring budbreak, early shoot growth, and initial fruit development before current-season root uptake can meet demand. Within the BFO framework, this quality specifically addresses the reserve utilization dimension of nutrient remobilization, distinguishing mobilization from stored pools from recovery during senescence while recognizing that storage mobilization provides the critical bridge between seasons, enabling rapid spring growth from accumulated reserves when soil conditions (cold, dry, or nutrient-depleted) limit root uptake capacity. The ontological significance lies in storage mobilization efficiency determining how effectively accumulated reserves translate into early-season growth and development, where the rate and completeness of reserve mobilization affects budbreak vigor, initial shoot growth rate, inflorescence development, and early fruit set that together establish the foundation for seasonal productivity before current-season assimilation and uptake dominate vine carbon and nutrient supply. The biochemical basis for storage mobilization involves enzymatic breakdown of stored compounds (starch hydrolysis releasing glucose, protein degradation releasing amino acids, phytate hydrolysis releasing phosphorus), conversion to transport forms (amino acids, sugars, phosphate ions), and loading into xylem or phloem for transport to demanding sinks, with the process regulated by hormonal signals (cytokinins promoting mobilization) and sink demand strength. The storage organs involved in grapevine nutrient reserves include permanent roots (major nitrogen and carbohydrate storage, typically 50-60% of reserve N), trunk and cordons (substantial storage capacity in wood parenchyma), and one-year-old canes (supporting initial bud development), with the relative contribution varying with vine age, architecture, and reserve status. The seasonal timing of storage mobilization shows maximum rates during budbreak and early shoot growth (weeks 0-6 after budbreak) when reserve-derived nutrients may provide 50-80% of shoot nitrogen, declining as current-season uptake increases, and with reserve contribution continuing to support fruit development until veraison when direct uptake dominates. The factors affecting mobilization efficiency include reserve status (well-nourished vines mobilizing faster), temperature (warming promoting enzymatic activity and transport), sink demand (strong shoot growth pulling reserves), and root activity (early root function potentially competing with mobilization-based supply). Understanding storage mobilization efficiency quality enables assessment of reserve adequacy for spring growth, optimization of post-harvest management building reserves, and appreciation of the perennial nutrient cycling that enables vigorous seasonal growth from dormant vines.
Storage_Mobilization_Efficiency_Quality addresses the release of nutrients from perennial reserves supporting spring growth before current-season uptake dominates. This quality determines how effectively accumulated reserves translate into early-season vigor and development. The class supports reserve management and spring growth prediction through mobilization efficiency characterization.
Reservemobilisierung
Reservenutzung
Speichermobilisierungseffizienz
Speichernährstofffreisetzung
reserve mobilization
reserve utilization
spring mobilization
storage nutrient release
Zapata C et al. (2004) Partitioning and mobilization of starch and N reserves in grapevine (Vitis vinifera L.). Journal of Plant Physiology 161(9): 1031-1040 Holzapfel BP et al. (2010) Manipulating the postharvest period and its impact on vine productivity of Semillon grapevines. American Journal of Enology and Viticulture 61(3): 367-376
Storage Mobilization Efficiency Quality
Cabernet Sauvignon storage pulp tissue sugar accumulation during ripening= veraison (BBCH 81, week 10) storing 14-16% sugars (1.8-2.2 g per berry) mid-ripening (week 14) storing 19-21% sugars (2.8-3.4 g per berry) reflecting 1.2% per week accumulation rate maturity (week 18) storing 24-26% sugars (3.8-4.5 g per berry) with accumulation rate declining to 0.6% per week as maturity approaches, demonstrating progressive sugar storage through phloem import and vacuolar accumulation creating final composition determining wine alcohol potential (24-26% sugars → 14-15% alcohol after fermentation). Chardonnay storage pulp tissue acid dynamics= veraison showing 17-20 g/L titratable acidity (predominantly malic acid 10-14 g/L, tartaric 6-8 g/L) mid-ripening showing 11-14 g/L (malic declining to 5-8 g/L through respiration, tartaric stable 6-8 g/L) maturity showing 6.5-8.5 g/L (malic 2.5-4 g/L, tartaric 4-5 g/L), demonstrating storage pulp tissue acid management through selective malic acid respiration while maintaining tartaric acid storage creating final acid balance important for white wine freshness and pH control. Merlot storage pulp tissue irrigation effects on storage and concentration= well-watered vines showing storage pulp accumulating 4.2 ml water and 3.5 g sugars per berry creating 23% sugar concentration deficit-irrigated vines showing 3.1 ml water and 3.3 g sugars per berry creating 25.5% concentration (11% higher despite 6% lower absolute sugar storage), demonstrating that deficit irrigation affects water storage more than sugar storage creating concentration effects important for wine quality through elevated sugar (higher alcohol), elevated phenolics per unit volume, more concentrated flavors. Syrah storage pulp tissue mineral accumulation= potassium accumulating from 450 mg/kg at veraison to 1650 mg/kg at maturity through preferential phloem transport and vacuolar storage calcium relatively stable 75-95 mg/kg (limited phloem mobility) magnesium increasing 42 to 78 mg/kg, demonstrating selective mineral storage with highly mobile potassium accumulating dramatically while less mobile calcium storage limited, affecting wine composition particularly pH (high potassium promoting pH increase through tartrate precipitation creating potassium bitartrate). Pinot Noir storage pulp tissue crop load effects on storage per berry= low crop load (4 clusters per shoot) allowing 3.8 g sugars storage per berry in 2.6 g berries (24.5% concentration) high crop load (12 clusters per shoot) limiting storage to 2.9 g sugars per berry in 2.1 g berries (23.2% concentration), demonstrating crop load effects on storage capacity with lower crop allowing more resources per berry supporting larger berries with higher absolute storage though concentration effects variable depending on proportional changes in solute versus water storage.
A storage pulp tissue is pulp tissue emphasizing its function as primary storage organ accumulating photosynthetic products (sugars), water, organic acids, and minerals imported from the plant, characterized by large vacuoles (90-95% of cell volume) serving as storage compartments, active transport systems importing solutes from phloem, and composition changes during ripening reflecting progressive accumulation of stored compounds determining berry and wine composition. Storage pulp tissue storage functions= (1) sugar storage accumulating glucose and fructose from <5% at fruit set to 18-28% at maturity through phloem import and vacuolar transport, storing 3.5-7 g sugars per berry (cultivar and size dependent) representing majority of berry carbon reserves (2) organic acid storage maintaining tartaric acid (stable during ripening, 0.3-0.6% at maturity) and malic acid (declining during ripening from 0.4-0.8% at veraison to 0.1-0.4% at maturity through respiration) (3) water storage accumulating water through osmotic import driven by sugar accumulation, storing 1.5-5 ml water per berry providing berry hydration and juice volume (4) mineral storage particularly potassium (accumulating to 1200-2200 mg/kg), calcium (60-150 mg/kg), magnesium (60-120 mg/kg) important for wine chemistry and yeast nutrition. Storage pulp tissue transport mechanisms= (1) phloem unloading in vascular bundles penetrating pulp tissue releasing sugars, water, minerals into apoplast (cell wall space) (2) plasma membrane transporters (sugar transporters, H⁺-ATPases, ion channels) moving solutes from apoplast into cytoplasm (3) vacuolar membrane transporters moving solutes from cytoplasm into vacuole for storage (4) osmotic water movement following solute accumulation creating turgor-driven cell expansion (5) transport activity regulated by developmental signals (hormones, gene expression changes) coordinating ripening progression. Storage pulp tissue compositional changes during ripening= (1) Phase I (BBCH 71-75, weeks 0-6) showing low sugar accumulation (3-8%), moderate acids (8-15 g/L), active cell division creating storage capacity (2) lag phase (BBCH 77, weeks 6-8) showing minimal compositional changes but metabolic reprogramming preparing for sugar accumulation (3) Phase II (BBCH 81-89, weeks 9-18+) showing rapid sugar accumulation (0.8-1.5% per week early ripening, 0.3-0.8% per week late ripening) through active phloem import and storage, malic acid decline through respiration (0.5-1 g/L per week), water accumulation driving cell expansion (4) maturity showing maximum storage levels (18-28% sugars, 4-9 g/L acids). Storage pulp tissue in wine quality determination= (1) stored sugar levels determining potential alcohol (typical wine grape targets 22-26% sugars creating 13-15.5% alcohol after fermentation) (2) stored acid levels determining wine acidity, pH, and stability (targets 5-8 g/L titratable acidity, pH 3.2-3.8) (3) sugar=acid ratio determining wine balance (target ratios 25-40=1 for balanced wines) (4) mineral storage affecting wine composition particularly potassium (influencing pH, tartrate stability) and nitrogen compounds (affecting yeast nutrition and fermentation kinetics). Storage pulp tissue environmental and cultural influences= irrigation affecting water storage and consequently sugar concentration (deficit irrigation reducing water storage creating concentrated sugars, adequate irrigation supporting water accumulation creating dilute sugars) crop load affecting storage capacity per berry (low crop load allowing more storage per berry creating larger berries with higher absolute but sometimes lower concentration of stored compounds) temperature affecting storage rate (warmer temperatures accelerating sugar accumulation, cooler temperatures slowing accumulation extending ripening duration).
Storage_Pulp_Tissue emphasizes the storage and accumulation functions of pulp tissue and is distinguished from Soft_Pulp_Tissue (texture) and Liquid_Pulp_Tissue (juice content). Storage pulp tissue represents the pulp from the perspective of compositional development and resource accumulation. Understanding storage pulp tissue is essential for managing berry composition through cultural practices (irrigation, crop load, canopy management) affecting storage capacity and accumulation rates, for harvest timing achieving target stored compound levels, and for comprehending developmental physiology of ripening as progressive storage process.
Speicher-Fruchtfleisch
Speicher-Pulpa-Gewebe
accumulation tissue
speicherndes Fruchtmark
storage flesh
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 190-220 Coombe, B.G. (1992). Research on development and ripening of the grape berry. American Journal of Enology and Viticulture 43=101-110 Zhang, X.Y., Keller, M. (2015). Grape berry transpiration is determined by vapor pressure deficit, cuticular conductance, and berry size. American Journal of Enology and Viticulture 66=454-462.
Storage Pulp Tissue
A straight simple unbranched tendril at node 7 on a vertically trained shoot of Vitis vinifera cv. Sauvignon Blanc in Marlborough, New Zealand, measuring 19 cm length with less than 10 degrees total deviation from a straight line between base and apex, showing 2.9-3.2 mm relatively uniform diameter along most of the length, bilateral symmetry evident in vascular bundle distribution and epidermal cell patterns, exhibiting circumnutation that sweeps the straight axis through space describing a cone of approximately 140-degree opening angle, reaching 19 cm horizontally from the shoot axis enabling contact with a vertical trellis wire positioned 17 cm from the shoot, making successful attachment through tight coiling while maintaining relatively straight basal section, occurring on a vertically shoot-positioned vine where straight tendril geometry provides optimal interaction with planar vertical wire trellis, measured at BBCH stage 71 (fruit set) in early December (Southern Hemisphere), representing the predominantly straight architecture characteristic of this cultivar under VSP training.
A simple unbranched tendril exhibiting predominantly linear growth trajectory with minimal curvature in its relaxed non-coiled state in Vitis vinifera, characterized by tendril axis showing less than 15-20 degrees deviation from a straight line between shoot attachment point and apex when measured in three-dimensional space, relatively uniform diameter along most of the length creating cylindrical rather than tapered morphology, bilateral symmetry in tissue development without pronounced differential growth on opposite sides that would create curvature, and geometric properties facilitating efficient spatial exploration through straight-line projection followed by circumnutation sweeping the extended axis through space. Straight simple unbranched tendrils develop when growth rates remain balanced on opposite sides of the developing axis throughout elongation, environmental conditions including light, gravity, and mechanical stimulation remain symmetrically distributed relative to the tendril axis, and genetic programming maintains bilateral developmental symmetry without activating differential growth patterns. The functional implications of straight architecture include maximum reach distance for a given tendril length enabling contact with supports at greater distances from the shoot, efficient exploration of linear spaces such as vertical wire planes in trellis systems, reduced risk of self-entanglement or contact with other vine structures during circumnutation, and potentially enhanced structural efficiency with straight column-like geometry providing maximum compressive and tensile strength per unit cross-sectional area compared to curved configurations that introduce bending moments.
Straight simple unbranched tendrils are distinguished from curved simple unbranched tendrils by minimal deviation from linear geometry, typically quantified as <15-20 degrees total curvature when measured in relaxed pre-coiling state. Assessment requires observation before coiling or attachment as these processes create secondary curvatures unrelated to intrinsic growth geometry. Straightness varies among cultivars with some showing predominantly straight tendrils (60-80% of unbranched tendrils) while others show curved tendencies, and within cultivars varies with shoot positioning and training system, with vertically oriented shoots in VSP systems promoting straight tendril development while more horizontal shoot orientations may favor curved patterns. Environmental factors including light directionality, gravitropic responses, and mechanical perturbations can induce curvature even in genetically straight-tending tendrils. Functionally, straight geometry provides advantages in reaching distant supports and interacting with planar trellis geometries common in modern mechanized viticulture.
gerade unverzweigte Ranke
linear unbranched tendril
linear-achsige einfache Ranke
nicht-gekrümmte einzelne Kletterorgan
non-curved single tendril
straight-axis simple tendril
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 120-135 Darwin (1875) "The Movements and Habits of Climbing Plants" John Murray, London Jaffe & Galston (1968) "The physiology of tendrils" Annual Review of Plant Physiology 19=417-434 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 155-172.
Straight Simple Unbranched Tendril
Stress relaxation analysis of Cabernet Sauvignon berries compressed to 10% strain reveals initial stress of 2.8 N declining to 1.2 N after 120 seconds, with bi-exponential fitting identifying fast relaxation (τ₁ = 8 s, 35% contribution) attributed to water redistribution and slow relaxation (τ₂ = 85 s, 40% contribution) attributed to cell wall rearrangement, leaving 25% non-relaxing elastic component, explaining why texture analyzer measurements show 30% lower peak force when compression rate is reduced from 5 mm/s to 0.5 mm/s due to concurrent relaxation during slow loading.
Stress relaxation quality is a viscoelastic property quality that represents the time-dependent decrease in stress occurring under constant applied strain, characterizing the progressive force reduction as polymer chains rearrange to dissipate internally stored elastic energy while maintaining fixed deformation. Within the BFO framework, this quality specifically addresses the temporal evolution of stress under maintained strain, distinguishing stress relaxation from creep which addresses strain changes under constant stress, and representing the complementary aspect of time-dependent mechanical behavior. The ontological significance lies in stress relaxation determining the persistence of internally generated forces in constrained tissues, where relaxation affects the pressure exerted by growing tissues against constraints, the force required to maintain berry compression during quality assessment, and the residual stress in deformed tissues following mechanical handling. Stress relaxation in grapevine tissues follows characteristic exponential or multi-exponential decay kinetics reflecting the superposition of relaxation processes with different time constants, from rapid relaxation (seconds) associated with fluid redistribution to slow relaxation (hours) associated with cell wall polymer rearrangement. The extent of relaxation (percentage of initial stress dissipated at equilibrium) indicates the relative contributions of elastic and viscous components, with tissues showing complete relaxation behaving as viscoelastic liquids while tissues with substantial residual stress retain solid-like elastic character. Measurement employs rapid strain application followed by force monitoring at constant deformation, with relaxation modulus (stress per unit strain) as function of time providing the standardized parameter, often analyzed through multi-exponential fitting to extract time constants and relaxation strengths for component processes. Understanding stress relaxation quality enables prediction of force persistence in compressed tissues, interpretation of texture assessment variability related to measurement timing, and analysis of the mechanical equilibration following tissue deformation during growth or handling.
Stress_Relaxation_Quality specifically addresses stress decrease under constant strain, complementing Creep_Behavior_Quality which addresses strain increase under constant stress. This quality affects texture measurement interpretation where loading rate influences apparent firmness through concurrent relaxation. The class supports mechanical testing protocol optimization and texture perception analysis through characterization of the time-dependent stress response to maintained deformation.
Kraftabbau
Maxwell relaxation
Spannungsabfall
Spannungsrelaxation
force relaxation
stress decay
viscoelastic relaxation
viskoelastische Relaxation
Peleg M (1980) Linearization of relaxation and creep curves of solid biological materials. Journal of Rheology 24(4): 451-463 Varela P et al. (2008) Comparison of conventional descriptive analysis and a citation frequency-based descriptive method for odor profiling: an application to Burgundy Pinot noir wines. Food Quality and Preference 19(1): 44-55
Stress Relaxation Quality
Vitis champinii-derived rootstocks such as Ramsey (Salt Creek) and Dog Ridge bear a Stress_Resistance_Rootstock_Role through combined resistance to root-knot nematodes (Meloidogyne spp.), phylloxera, and moderate fungal pathogen tolerance, making them essential rootstock choices in sandy soils of the San Joaquin Valley, California, and the Barossa Valley, Australia. Freedom (1613C × Dog Ridge) combines nematode resistance from V. champinii with phylloxera resistance, supporting Zinfandel production on nematode-infested sites in Lodi, California.
A rootstock role that inheres in a grafted rootstock by virtue of its capacity to confer resistance or enhanced tolerance to biotic stress factors upon the grafted vine composite. Stress-resistant rootstocks protect the vine system against soil-borne pests and pathogens through genetically determined defense mechanisms including physical barriers (lignified root cortex, suberized exodermis), biochemical defenses (phenolic compound accumulation, phytoalexin synthesis), and hypersensitive response pathways. The stress resistance conferred by rootstocks is primarily derived from North American Vitis species (V. riparia, V. rupestris, V. berlandieri, V. champinii, V. cinerea) that coevolved with endemic pests and pathogens, developing resistance mechanisms absent in the European V. vinifera. This role encompasses resistance to phylloxera, nematodes, and fungal root pathogens.
Stress_Resistance_Rootstock_Role is restricted to biotic stress factors abiotic stress tolerance (drought, salinity, lime, waterlogging, cold, heat) is classified under Abiotic_Stress_Tolerant_Rootstock_Role as a separate sibling hierarchy. This distinction reflects the fundamentally different biological mechanisms involved: biotic resistance engages immune-like defense pathways while abiotic tolerance involves physiological adaptation and homeostatic regulation. Many rootstock cultivars bear both biotic resistance and abiotic tolerance roles simultaneously.
biotic stress resistant rootstock
disease-resistant rootstock
pest-resistant rootstock
schädlingsresistente Unterlage
stress-resistant rootstock
stressresistente Unterlage
widerstandsfähige Unterlage
Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22 Granett J et al. (2001) Biology and management of grape phylloxera, Annual Review of Entomology 46: 387-412 Corso M, Bonghi C (2014) Grapevine rootstock effects on abiotic stress tolerance, Plant Science 228: 1-14
Stress Resistance Rootstock Role
Stress response quality characterization in Barossa Valley compared Shiraz vine behavior across a gradient of water availability from fully irrigated through regulated deficit irrigation to severe drought stress during a dry vintage. Vines demonstrated the characteristic grapevine stress response quality progression with initial stomatal closure reducing transpiration while maintaining photosynthesis at mild stress levels, followed by reduced shoot growth redirecting resources to fruit development at moderate stress, and finally premature leaf senescence and impaired ripening under severe stress. Fruit composition reflected the stress response quality trajectory, with moderate stress producing optimal outcomes including concentrated flavors, elevated anthocyanins, and balanced sugar-acid relationships, while severe stress caused incomplete ripening, elevated pH from potassium concentration, and diminished varietal character. The stress response quality assessment established target predawn leaf water potential of -0.3 to -0.6 MPa as the optimal stress range for premium Shiraz production, with monitoring protocols enabling irrigation management to maintain vines within this beneficial stress window throughout the critical post-véraison ripening period.
Stress response quality is a viticultural quality that inheres in a grapevine organism and represents the characteristic manner and magnitude of physiological, morphological, and biochemical adjustments exhibited in response to environmental stress factors including water deficit, temperature extremes, nutrient limitations, salinity, and biotic challenges, determining the capacity to maintain productive function under suboptimal conditions and the quality implications of stress-induced modifications to growth, development, and fruit composition. This quality fundamentally influences vineyard performance across the range of environmental conditions encountered in viticultural production, recognizing that grapevines rarely experience optimal conditions throughout the growing season and that stress responses represent a continuous modulation of vine function rather than discrete abnormal states. The ontological significance of stress response quality lies in its role as an integrative indicator of vine physiological status that manifests through observable symptoms, measurable parameters, and ultimately through fruit composition and wine quality outcomes, enabling diagnosis of stress conditions and prediction of their consequences for production objectives. The biological basis of stress response quality encompasses the sensing mechanisms by which vines detect environmental challenges, the signal transduction pathways translating perception into physiological response, the metabolic and developmental adjustments that constitute the stress response proper, and the recovery capacity determining how rapidly normal function resumes when stress is relieved. Grapevine stress responses involve coordinated adjustments across multiple organizational levels from molecular changes in gene expression and enzyme activity through cellular osmotic adjustment and membrane modification to whole-plant alterations in growth allocation, stomatal behavior, and reproductive development. The relationship between stress response quality and fruit quality reflects the complex reality that moderate stress often enhances wine grape quality through mechanisms including reduced vegetative growth improving vine balance, concentration effects enhancing berry composition, and stress-induced secondary metabolism producing quality-relevant compounds, while severe stress compromises quality through impaired ripening, reduced photosynthetic capacity, and premature senescence. Assessment of stress response quality requires integration of physiological measurements, visual symptom evaluation, and outcome tracking across the range of stress intensities encountered in production contexts.
Stress_Response_Quality serves as a high-level parent class for specific stress type indicators including Drought_Stress_Indicator_Quality and potentially other stress categories addressing temperature, salinity, nutrient, and biotic stress responses. This class captures the general concept of vine response to environmental challenges while child hierarchies address particular stress factors. Distinguished from Disease_Susceptibility_Quality by the focus on abiotic environmental stresses rather than pathogen interactions. The moderate stress benefit for wine quality represents key viticultural principle underlying deficit irrigation and related management strategies.
Stressphysiologie
Stressreaktionsqualität
Umweltstress-Antwort
abiotic stress response
abiotische Stressreaktion
environmental stress response
stress adaptation
stress physiology
Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data. Annals of Botany 105:661-676 Lovisolo C et al. (2010) Drought-induced changes in development and function of grapevine (Vitis spp.) organs and in their hydraulic and non-hydraulic interactions at the whole-plant level: a physiological and molecular update. Functional Plant Biology 37:98-116 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 453-507
Stress Response Quality
Documentation of Strong_Apical_Dominance_Disposition at UC Davis characterized lateral suppression in cultivars recognized for minimal branching. Reference cultivars 'Merlot', 'Zinfandel', 'Petite Sirah', and 'Mourvèdre' were grown on VSP without lateral removal to assess natural branching expression. Detailed measurements included lateral number by node position, lateral length distribution, and the response of lateral development to apex removal (decapitation). Strong_Apical_Dominance_Disposition cultivars showed consistent characteristics. 'Merlot' exhibited archetypal strong apical dominance with mean of 3.8 lateral shoots per primary shoot of 165 cm length, concentrated at basal nodes 1-4 (85% of laterals), with total lateral length of only 42 cm (lateral/primary length ratio 0.25). Lateral shoots were short (mean 11 cm) and ceased growth early, reflecting persistent apical suppression. 'Petite Sirah' showed even stronger dominance (2.9 laterals, 28 cm total length). Decapitation experiments confirmed strong apical control—removal of 'Merlot' apex at 80 cm triggered outgrowth of 8.5 additional lateral buds within 14 days, demonstrating that buds were competent but hormonally suppressed. Apex auxin concentration in Strong_Apical_Dominance_Disposition cultivars averaged 285 ng/g FW (45% above cultivar mean), correlating with suppression strength (r = 0.78 between auxin and inverse lateral number). Canopy density measurements confirmed that Strong_Apical_Dominance_Disposition cultivars achieved 1.2-1.5 leaf layers without lateral removal, compared to 2.5-3.5 in prolific branching types, demonstrating the practical advantage for fruit zone light exposure.
A Strong_Apical_Dominance_Disposition is a Branching_Pattern_Disposition that inheres in a Vitis vinifera plant and is realized through robust hormonal suppression of lateral bud outgrowth by the primary shoot apex, resulting in minimal lateral shoot production, simple unbranched shoot architecture, and naturally open canopy structure without intensive lateral management. This disposition represents the upper extreme of correlative inhibition strength, characteristic of cultivars where auxin export from the shoot apex effectively prevents lateral bud activation across most or all nodes of the primary shoot. Cultivars with Strong_Apical_Dominance_Disposition typically produce fewer than 5 lateral shoots per primary shoot under normal growing conditions, with lateral development often restricted to basal nodes (where apical influence is weakest) or occurring only after hedging removes the dominant apex. The viticultural advantages of strong apical dominance include reduced labor requirements for lateral shoot management, naturally lower canopy density with better light penetration, and simplified canopy architecture facilitating mechanical operations. However, the disposition can be disadvantageous when lateral growth is desired—for example, to fill canopy gaps after frost damage or to increase leaf area on low-vigor vines. Strong_Apical_Dominance_Disposition is associated with higher auxin concentrations in shoot apices and more efficient polar auxin transport maintaining apical suppression of lateral buds.
Strong_Apical_Dominance_Disposition is_a Branching_Pattern_Disposition representing minimal lateral development (< 5 laterals per primary shoot). Characterized by high apex auxin, efficient correlative inhibition, and lateral restriction to basal nodes. Examples include 'Merlot', 'Zinfandel', 'Petite Sirah', 'Mourvèdre'. Advantages include reduced lateral management labor and naturally open canopies. Laterals released by apex removal (hedging) confirm hormonal suppression mechanism. Strong dominance may limit canopy recovery after damage.
Hohe-Apikalkontrolle
Starke-Apikaldominanz-Disposition
Unterdrückte-Geiztrieb-Bildung
high apical control
suppressed lateral growth
unbranched shoot habit
Cline MG (1991) Apical dominance, Botanical Review 57:318-358 Lebon E et al. (2006) Shoot development in grapevine (Vitis vinifera L.) is affected by the modular branching pattern of the stem and intra- and inter-shoot trophic competition, Annals of Botany 98:1045-1056 Tanaka M et al. (2006) Auxin controls local cytokinin biosynthesis in the nodal stem in apical dominance, Plant Journal 45:1028-1036
Strong Apical Dominance Disposition
A strong secondary vegetative shoot on Vitis vinifera cv. Chardonnay from node 7, activated following early-season hail (late April) that stripped primary shoots, developed to 118cm length by late July, 15 leaves averaging 135 cm² totaling approximately 2025 cm² leaf area, 7.5mm basal diameter with early lignification visible in lower internodes, representing robust secondary development attributed to combination of vigorous Chardonnay characteristics, favorable warm growing season following damage, adequate vine carbohydrate reserves from previous season, and appropriate remaining crop load (partial primary survival on other vines maintaining moderate whole-vine crop) enabling strong compensation, this shoot retained through season for significant photosynthetic contribution and evaluated during dormant pruning as potential renewal cane candidate based on good vigor and positioning.
A secondary vegetative shoot exhibiting relatively vigorous growth despite secondary bud origin, characterized by moderate-to-good elongation achieving 90-130cm length, adequate leaf development with 12-16 leaves providing 1400-2000 cm² photosynthetic area, reasonable diameter (6-8mm at base), and growth performance approaching though not equaling typical primary shoot development, representing upper range of secondary shoot vigor occurring under favorable compensation conditions including adequate vine reserves, favorable post-damage weather, appropriate remaining crop load, and inherently vigorous cultivars or sites supporting robust secondary development. Strong secondary vegetative shoots demonstrate maximal compensation potential from backup bud positions, providing substantial photosynthetic replacement following primary loss, contributing meaningfully to canopy recovery and fruit support, and occasionally suitable as renewal wood when positioning and quality permit, representing best-case secondary shoot outcomes enabling significant though incomplete recovery from damaging events triggering secondary activation.
Strong secondary vegetative shoots represent optimal secondary compensation outcomes, achieving 70-85% of typical primary shoot development and providing substantial photosynthetic capacity supporting fruit maturation and vine function. The factors enabling strong secondary development include inherent vine vigor (young vines, vigorous rootstocks, fertile sites), favorable post-damage conditions (warm temperatures, adequate moisture, nutrient availability), appropriate crop load (moderate remaining crop preventing excessive sink demand), and cultivar characteristics (some varieties showing better secondary development than others). Strong secondaries partially mitigate damage consequences, though cannot fully compensate for primary loss due to delayed phenology, reduced total leaf area across canopy (incomplete secondary breakage), and absence of crop contribution from secondaries. Management typically retains strong secondary shoots recognizing valuable contributions, with best-developed shoots sometimes suitable for renewal wood providing alternative to delayed establishment from latent bud water shoots.
kräftiger Ersatztrieb
robust compensatory shoot
starker sekundärer vegetativer Trieb
vigorous secondary vegetative shoot
Strong Secondary Vegetative Shoot
Cabernet Sauvignon exhibits strong tendril disposition with tendril elongation rates of 8-12 cm/day during active growth, coiling response initiated within 30-60 minutes of contact, and attachment tensile strength exceeding 15 N. Wild Vitis species such as V. riparia show even stronger tendril development with attachment forces up to 25 N, reflecting their natural climbing habit in forest margins.
A tendril development disposition that is the inherited capacity of a Vitis vinifera plant to produce tendrils with high biomass, rapid elongation rate, strong coiling response, and persistent attachment strength, enabling effective climbing and canopy support.
Strong tendril disposition correlates with vigorous vegetative growth patterns and is particularly pronounced in cultivars derived from wild Vitis species. The coiling mechanism involves differential cell elongation on the contact versus non-contact side mediated by auxin redistribution and jasmonat signaling. Strong tendrils may require more frequent shoot positioning in VSP systems but provide better natural support in minimal-pruning systems.
high tendril vigor
hohe Rankenkraft
kräftige Rankenentwicklung
starke Rankenbildung
strong climbing capacity
vigorous tendril formation
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 42-45 Bowling AJ, Bhattacharya S (2022) Tendril coiling in Vitis: biomechanics and molecular mechanisms. American Journal of Botany 109:234-248
Strong Tendril Disposition
Structural integrity declines progressively with vine age as functional sapwood is converted to non-conductive heartwood, wound compartmentalization creates discontinuous vascular zones, and trunk disease fungi colonize dead tissue and spread into living wood. Cultivars with dense, durable wood such as Mourvèdre and Carignan show slower structural decline than softer-wooded types. Trunk cross-sectional area of functional xylem may decline from 80-90% in young vines to 30-50% in old vines through combined effects of heartwood formation and disease damage. Mechanical failure (trunk breakage, cordon collapse) becomes increasingly likely as the ratio of dead to living wood increases, with failure typically occurring during wind events or under heavy crop load. Wound management practices (pruning cut protection, disease prevention) significantly affect the rate of structural integrity loss.
A vine longevity disposition that is the inherited capacity of a Vitis vinifera plant to maintain the physical strength and functional integrity of woody perennial structures (trunk, cordons, spurs) over time, resisting mechanical failure, decay, and the progressive loss of conductive tissue that limits productive lifespan.
Structural integrity represents the physical foundation of vine longevity, with mechanical failure setting the ultimate limit on productive lifespan even when vascular function and regenerative capacity remain adequate. The trait integrates wood density, wound compartmentalization ability, and resistance to wood-decay fungi. Training system design affects structural demands, with high-cordon systems placing greater mechanical stress on supporting structures than low-trained systems.
Holzbeständigkeit
Strukturelle Integrität
mechanical integrity trait
mechanische Stabilität
structural soundness
wood durability
Shigo AL (1984) Compartmentalization: a conceptual framework for understanding how trees grow and defend themselves. Annual Review of Phytopathology 22:189-214 Pearce RB (1996) Antimicrobial defences in the wood of living trees. New Phytologist 132:203-233
Structural Integrity Disposition
Structural integrity quality assessment in export table grape production for Red Globe requires evaluation at multiple supply chain points, with maximum allowable defects specified as less than 2% shattered berries (detachment from pedicel), less than 1% split berries, less than 3% mechanical damage (cuts, punctures, abrasions), and no decay fruit failing these structural integrity standards at destination inspection results in price adjustments of 20-50% or outright rejection, making structural integrity preservation through careful handling and cold chain maintenance essential for profitable long-distance table grape trade.
Structural_Integrity_Quality encompasses all morphological qualities that characterize the wholeness, cohesion, and physical soundness of grapevine organs. This category addresses whether organs maintain their complete, undamaged form with all parts properly connected and functional, or whether they exhibit defects, damage, or deterioration that compromise their structural completeness. Structural integrity is a critical quality determinant for both table grapes and wine grapes because it directly affects marketability, storage potential, pathogen susceptibility, and processing behavior. This category subdivides into four major domains: Intactness_Quality addressing whether organs are complete without missing parts Cohesion_Quality capturing the internal binding forces that hold tissues together Attachment_Strength_Quality characterizing the connections between organs and their supporting structures and Damage_State_Quality describing the presence and type of structural defects. Structural integrity depends on cell wall strength, middle lamella adhesion, turgor pressure maintenance, and the absence of mechanical, environmental, or biological damage. During development, structural integrity generally increases as tissues mature and cell walls strengthen, while during ripening and senescence, integrity typically decreases as cell walls soften and tissues become more vulnerable to damage. Environmental stresses including water deficit, excessive heat, and rapid temperature fluctuations can compromise structural integrity through effects on turgor and cell wall properties. Mechanical forces during harvest, transport, and processing challenge structural integrity, with damage accumulating at each handling step. Pathogen infection often initiates at sites of compromised integrity where wounds or cracks provide entry points. Assessment of structural integrity employs visual inspection for damage and defects, mechanical testing of tissue strength and attachment, and careful documentation of the proportion of sound versus damaged organs in sample populations.
Structural_Integrity_Quality is positioned as a direct subclass of Morphological_Quality, parallel to Size_Quality, Shape_Quality, Color_Quality, and Texture_Quality. This category addresses the physical completeness and soundness of organs rather than their dimensional, geometric, chromatic, or textural properties. Structural integrity qualities often interact with other quality categories for example, damage affects color (browning), texture (softening), and potentially size (shrinkage). The assessment of structural integrity typically involves categorical classification (intact versus damaged) with quantification of damage extent and type. PATO_0001995 (structural quality) provides a related upper-ontology concept.
Ganzheitsqualität
Strukturintegrität
organ integrity
physical soundness
physische Unversehrtheit
structural completeness
strukturelle Vollständigkeit
wholeness quality
Crisosto CH et al. (2001) Table Grapes Respond to Mechanical Damage. California Agriculture 55(5): 30-31 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 159-189
Structural Integrity Quality
Cabernet Sauvignon structural outer mesocarp providing robust structural support= 5-7 cell layers with walls 1-1.4 μm providing structural transition between hypodermal support layers (walls 1.8-2.6 μm) and soft middle mesocarp (walls 0.5-0.7 μm), creating continuous structural system supporting berry expansion during ripening (berry diameter increasing from 9 mm at veraison to 14.5 mm at maturity, 61% increase) without skin-flesh separation or structural failure, demonstrating structural outer mesocarp ability to accommodate substantial expansion while maintaining integrity. Syrah structural outer mesocarp splitting resistance role= structural outer mesocarp with walls 1.1-1.5 μm, tissue tensile strength 0.8-1.4 MPa (estimated from cell and wall properties), contributing to overall skin strength 1.2-2 MPa allowing berries to withstand turgor increases during late-season rainfall (25 mm rain event at week 17 near harvest creating berry turgor increase from 0.4 to 0.65 MPa) with only 3-8% splitting incidence, versus cultivars with weaker structural outer mesocarp (walls 0.7-1 μm, tissue strength 0.4-0.8 MPa) showing 15-35% splitting under same conditions. Thompson Seedless structural outer mesocarp table grape shipping durability= structural outer mesocarp (cells 35-58 μm, walls 1.1-1.6 μm, 4-6 layers) providing mechanical support maintaining berry integrity during 4-week cold storage and shipping (0-2°C, controlled atmosphere), with berries showing <5% compression damage and <3% stem separation attributed to maintaining structural outer mesocarp integrity through careful handling and calcium nutrition enhancing cell wall strength. Pinot Noir structural outer mesocarp in splitting-susceptible cultivar= thinner structural outer mesocarp (3-4 layers, walls 0.9-1.2 μm) providing moderate structural support sometimes insufficient to prevent splitting when late-season rainfall creates rapid water uptake (30 mm rain producing 12-22% splitting incidence in susceptible vineyards), demonstrating that limited structural outer mesocarp thickness and strength contributes to Pinot Noir characteristic splitting susceptibility requiring harvest timing before autumn rains and cluster zone management (leaf removal, cluster thinning) reducing splitting risk. Merlot structural outer mesocarp berry expansion accommodation examined by microscopy during berry development= veraison (week 10, berry 10.5 mm diameter) showing structural outer mesocarp cells 42-58 μm mid-ripening (week 14, berry 12.5 mm) showing cells 52-72 μm (24-35% expansion) maturity (week 18, berry 14 mm) showing cells 58-82 μm (38-58% total expansion from veraison), with cell expansion accommodated through wall yielding (wall thinning from 1.3 to 1.1 μm despite cell enlargement) balanced by continued wall deposition maintaining structural integrity, demonstrating structural outer mesocarp controlled expansion mechanism supporting berry growth while preserving mechanical properties.
A structural outer mesocarp is an outer mesocarp emphasizing its structural support role as transition zone providing mechanical support between protective outer layers (epidermis, hypodermis) and soft storage inner mesocarp, focusing on mechanical properties including cell arrangement, wall strength, and contribution to overall berry and skin structural integrity particularly important for berry expansion accommodation, splitting resistance, and post-harvest handling durability. Structural outer mesocarp mechanical characteristics= (1) cells arranged in organized layers creating structural continuity from skin to flesh (2) moderately thick walls (0.9-1.5 μm) providing tensile strength and compression resistance intermediate between thick protective hypodermal walls (1.5-3 μm) and thin storage middle mesocarp walls (0.4-0.8 μm) (3) cell-cell adhesion through middle lamella pectin providing tissue cohesion (4) progressive transition in mechanical properties creating gradient from firm outer to soft inner avoiding abrupt property boundaries that could create failure points (5) potential wall modifications including enhanced cross-linking or specific pectin methylesterification patterns affecting mechanical properties. Structural outer mesocarp in berry expansion= (1) accommodation of berry growth through controlled cell wall extension during Phase II ripening, with outer mesocarp cells showing 40-80% diameter increase (versus 120-200% in middle mesocarp) through wall yielding balanced by wall reinforcement (2) maintenance of structural continuity during expansion preventing skin-flesh separation or delamination as berry enlarges (3) distribution of expansion-induced mechanical stresses preventing stress concentration at skin-flesh boundary (4) potential role in limiting excessive expansion through mechanical resistance creating back-pressure opposing turgor-driven expansion. Structural outer mesocarp in berry splitting resistance= (1) contribution to skin strength creating resistance to splitting when berries exposed to late-season rainfall or rapid water uptake (2) structural integration maintaining skin-flesh attachment preventing skin separation that could initiate splitting (3) mechanical properties affecting critical turgor pressure threshold for splitting with stronger structural outer mesocarp allowing higher turgor before failure (4) potential variation among cultivars in structural outer mesocarp properties correlating with splitting susceptibility (resistant cultivars showing stronger structural outer mesocarp, susceptible cultivars weaker). Structural outer mesocarp in berry mechanical testing= (1) contribution to berry firmness measured by compression testing, with structural outer mesocarp providing resistance in initial compression phase before middle mesocarp cells begin breaking (2) contribution to skin break force measured by puncture testing, with structural outer mesocarp providing resistance to penetration probe (3) contribution to berry stiffness (elastic modulus) reflecting structural resistance to deformation (4) potential correlation between structural outer mesocarp properties and instrumental texture measurements used for quality assessment. Structural outer mesocarp in post-harvest durability= (1) maintenance of berry integrity during harvest and handling preventing berry crushing or skin damage (2) resistance to compression damage during cluster packing in containers (3) structural support during storage preventing berry collapse or excessive softening (4) particular importance for table grapes requiring shipping durability over extended storage and transport periods (2-6 weeks typical for export markets).
Structural_Outer_Mesocarp emphasizes the mechanical support and structural integrity functions of outer mesocarp and is distinguished from Firm_Outer_Mesocarp which emphasizes textural properties. Structural outer mesocarp represents the outer mesocarp from the perspective of mechanical properties, expansion accommodation, and structural continuity. Understanding structural outer mesocarp is relevant for comprehending berry splitting mechanisms, for managing splitting resistance through cultural practices affecting cell wall properties (calcium nutrition, irrigation management), and for table grape breeding selecting for enhanced structural properties supporting shipping durability.
Stützende Übergangszone
structural transition zone
strukturelles äußeres Mesokarp
stützendes äußeres Fruchtfleisch
supporting outer flesh
Thomas, T.R. et al. (2008). Changes in physical properties of grape berries. American Journal of Enology and Viticulture 59=398-409 Considine, J.A., Kriedemann, P.E. (1972). Fruit splitting in grapes. Australian Journal of Agricultural Research 23=17-27 Nunan, K.J. et al. (2001). Changes in cell wall composition during ripening. Plant Physiology 126=520-528.
Structural Outer Mesocarp
Syrah grafted on 140 Ruggeri rootstock at 22 years showing structural woody secondary roots as large first-order laterals, primary structural root 7.4 cm diameter extending 195 cm horizontally at depth 68 cm providing main lateral anchorage and framework, structural root showing extensive secondary xylem with 21 annual rings and thick lignified fiber cells providing high mechanical strength, structural root supporting extensive second and higher-order branch network estimated 15 kg wet weight fine roots, structural woody root bearing mechanical loads from root network and resisting tension forces transmitted from above-ground structure, structural root providing permanent stable framework maintained throughout vine life
Structural woody secondary root is woody secondary root emphasizing mechanical support and structural framework function, characterized by primary role in providing root system architecture and physical support rather than absorption or short-term transport, typically large diameter 3-8 cm reflecting substantial structural capacity, demonstrating extensive lignified secondary xylem providing high mechanical strength and rigidity, persistent structure functioning for many years or decades creating stable permanent framework, and serving as structural skeleton from which entire absorptive fine root network branches enabling extensive soil exploration while maintaining mechanical integrity, with structural woody secondary roots bearing mechanical loads from fine root network weight, soil pressure, and forces transmitted from above-ground structure, resisting bending and tension forces that would otherwise deform or break root system, anchoring vine preventing uprooting or toppling particularly through horizontal structural roots spreading laterally, and demonstrating that woody root structural function analogous to building structural elements providing load-bearing framework supporting functional components like absorption and transport occurring primarily in branches. Structural woody secondary root mechanical support function includes= providing root system skeleton (structural woody roots forming permanent framework analogous to building frame supporting structure), bearing dead weight loads (structural roots supporting mass of fine root network, stored carbohydrates, and water in tissues), resisting tension forces (structural woody roots bearing pulling forces from wind loads on canopy transmitted through trunk), withstanding compression forces (structural roots enduring compression from soil overburden and root system weight), and enabling bending resistance (structural woody roots preventing excessive deformation maintaining root system shape and position). Structural woody secondary root in anchorage includes= creating primary anchorage elements (structural woody roots providing main anchoring components particularly horizontal laterals creating wide base), distributing anchor points (structural roots spreading throughout soil creating multiple distributed attachment points), generating friction resistance (structural root surface area and roughness creating friction against soil preventing sliding or pulling), bearing pull-out loads (structural roots resisting forces attempting to extract vine from soil through deep penetration and lateral spread), and enabling wind resistance (structural root architecture critically important for preventing toppling under wind loads on canopy). Structural woody secondary root spatial organization includes= defining root system architecture (structural woody root distribution and orientation determining overall root system spatial organization and shape), creating hierarchical framework (structural roots providing main axes from which progressively finer branches develop in hierarchical fashion), demonstrating geometric pattern (structural root arrangement often showing radial symmetry or other geometric organization optimizing stability and resource access), enabling three-dimensional exploration (structural woody roots extending vertically and horizontally creating framework for exploiting soil volume in all directions), and showing cultivar characteristics (structural root architecture patterns potentially diagnostic for rootstock variety identification). Structural woody secondary root load distribution includes= transmitting forces throughout system (structural roots conducting mechanical forces from localized stress points distributing loads across entire root network), preventing stress concentration (structural root branching and distribution avoiding excessive force concentration in single location), demonstrating redundancy (multiple structural roots providing backup load paths if individual root damaged or fails), enabling safety factors (structural root architecture typically overbuilt relative to normal loads providing reserve capacity for extreme events), and showing adaptive growth (structural roots potentially strengthening in regions experiencing high stress through responsive growth). Structural woody secondary root persistence requirement includes= necessitating longevity (structural function requiring roots persist for many years or decades providing stable long-term framework), demanding durability (structural roots must resist decay, damage, and degradation maintaining integrity throughout functional life), showing low turnover (structural woody roots persisting unlike fine roots with annual turnover reducing construction costs amortization period), requiring carbon investment (substantial photosynthate investment in constructing and maintaining structural roots justified by long functional life), and demonstrating cost-effectiveness (large initial structural root construction cost offset by decades of function providing favorable cost-benefit ratio). Structural woody secondary root versus absorptive roots includes= contrasting functions (structural roots for mechanical support and architecture versus absorptive roots for water and nutrient uptake), showing different anatomy (structural roots with extensive secondary growth and lignification versus absorptive roots with primary anatomy and minimal lignification), demonstrating size difference (structural roots large diameter 2-8 cm versus absorptive roots fine diameter <2 mm), exhibiting persistence contrast (structural roots functioning years to decades versus absorptive roots showing seasonal to annual turnover), and revealing carbon allocation (structural roots representing large carbon investment in long-lived tissue versus absorptive roots showing continuous investment in ephemeral tissue). Structural woody secondary root in root system efficiency includes= enabling extensive absorption (structural framework allowing absorptive network to spread widely without mechanical failure), optimizing resource allocation (construction of permanent structural roots reducing need for continuous reconstruction costs), facilitating responsive adjustment (stable structural framework allowing rapid proliferation or senescence of absorptive roots responding to conditions), demonstrating division of labor (structural versus absorptive function specialization improving overall system efficiency), and showing evolutionary optimization (structural woody root architecture representing evolved solution balancing mechanical requirements with resource acquisition needs). Structural woody secondary root and vine age includes= developing progressively with time (structural root network expanding and strengthening as vine ages from establishment through maturity), showing increasing importance (structural function becoming more critical as vine grows larger and heavier requiring stronger anchorage and support), demonstrating cumulative investment (older vines having invested more carbon in structural root network reflecting years of growth), revealing age documentation (structural root annual rings recording vine age and growth history), and exhibiting legacy effects (old vine performance partially dependent on structural root system architecture established years or decades earlier).
Structural_Woody_Secondary_Root emphasizes mechanical support and framework function and is distinguished from transport or storage emphasis by focus on structural role. Critical for understanding root system skeleton, mechanical support, load bearing, anchorage, architectural framework, and the large persistent woody roots providing stable permanent structure supporting dynamic absorptive root network and enabling vine mechanical stability.
Framework secondary root
Load-bearing woody root
Strukturelle holzige Sekundärwurzel
Stützende Wurzel
Support woody root
Tragenden Wurzel
Structural Woody Secondary Root
Chardonnay hermaphrodite flowers at BBCH 61-65 (flowering) exhibit styles 0.55-0.68 mm length connecting ovaries 0.85-1.05 mm to stigmas 0.42-0.52 mm, with style representing 26-32% of total pistil length (2.1-2.4 mm), containing well-developed transmitting tract tissue supporting pollen tube growth with pollen tubes traversing the style in 52-68 hours post-germination at 22-24°C, achieving successful fertilization in 48-56% of flowers. Gewürztraminer functionally female flowers show elongated styles 0.72-0.88 mm connecting enhanced ovaries 1.05-1.25 mm to prominent stigmas 0.52-0.62 mm, with styles representing 28-34% of pistil length (2.4-2.7 mm), providing extended pollen tube pathway requiring 58-78 hours for pollen tube transit from stigma to ovules when cross-pollinated from hermaphrodite pollinizers. Cabernet Franc style anatomical sections reveal central transmitting tract 0.08-0.12 mm diameter consisting of elongated cells with abundant extracellular matrix staining intensely with periodic acid-Schiff reagent (indicating polysaccharide-rich composition), surrounded by 4-6 layers of cortex parenchyma cells and outer epidermis, with 2-3 vascular bundles running through cortex providing style nutrition. Pinot Noir pollen tube growth studies using aniline blue fluorescence microscopy show pollen tubes entering style through stigmatic surface within 2-4 hours post-pollination, growing through transmitting tract extracellular matrix at rates of 12-25 μm per hour (temperature-dependent), and reaching ovary after 48-72 hours having traversed the 0.6-0.8 mm style length plus stigma and ovary tissues.
A style is the intermediate tubular or cylindrical portion of the pistil connecting the basal ovary to the terminal stigma, functioning as the pathway for pollen tube growth from the site of pollen germination on the stigma to the site of fertilization in the ovules within the ovary, with style length, diameter, and internal transmitting tract tissue organization critically affecting pollen tube guidance efficiency and fertilization success. The style structure in Vitis comprises= (1) an external cylindrical or slightly tapering tube typically 0.3-0.8 mm in length and 0.15-0.4 mm in diameter at anthesis, though dimensions vary substantially among flower types (2) outer epidermis providing protective covering and sometimes bearing trichomes particularly in functionally female or pistillate flower styles (3) cortex of parenchymatous cells surrounding the central transmitting tract (4) central transmitting tract tissue (stylar canal) consisting of specialized cells with elaborate extracellular matrix rich in pectins, arabinogalactan proteins, and other glycoproteins that support pollen tube growth, providing both physical substrate and nutritional/signaling environment for pollen tubes traversing from stigma to ovary (5) vascular tissues running through the style cortex providing nutrient and water supply to style tissues and continuing from receptacle through ovary to support ovule nutrition. Style development proceeds through phases= (1) initiation during early pistil primordium differentiation (BBCH 51-53) as the region between developing ovary and stigma primordia (2) elongation (BBCH 53-59) creating distinct separation between ovary and stigma (3) transmitting tract differentiation (BBCH 55-59) with specialization of central cells producing extracellular matrix for pollen tube support (4) functional maturity (BBCH 59-65) when transmitting tract achieves capacity to support pollen tube growth. Style length and proportions vary dramatically among flower types reflecting differences in pistil functionality= hermaphrodite flowers exhibit styles 0.4-0.7 mm connecting ovaries 0.7-1.1 mm to stigmas 0.3-0.5 mm, with style representing 20-35% of total pistil length functionally female and pistillate flowers show styles 0.5-0.9 mm (occasionally up to 1 mm) connecting larger ovaries 0.9-1.3 mm to elaborate stigmas 0.4-0.6 mm, with styles representing 20-35% of pistil length functionally male flowers exhibit reduced styles 0.3-0.6 mm connecting small ovaries 0.4-0.8 mm to rudimentary stigmas, with styles representing 25-40% of reduced pistil length staminate flowers show vestigial or rudimentary styles 0.1-0.4 mm in proportion to vestigial pistil size. The style serves as the critical conduit for pollen tube growth following pollen germination on the stigma, with pollen tubes entering the style through the stigmatic surface, growing through the transmitting tract extracellular matrix, and emerging into the ovary locules to reach ovules, with the journey requiring 48-96 hours at optimal temperatures (20-25°C) and involving pollen tube lengths of 1-2.5 mm depending on style and ovary dimensions.
Style represents the intermediate pistil component and must be distinguished from Ovary (basal component) and Stigma (terminal component). The style serves critical pollen tube guidance functions through its transmitting tract tissue, with style length and transmitting tract quality affecting fertilization efficiency. Style dimensions vary dramatically among flower types (longest in functionally female/pistillate, shortest in functionally male/staminate, intermediate in hermaphrodite) reflecting overall pistil size variation. Sub-classification into Short, Medium, and Long types reflects the substantial length variation among flower types and among individuals within types. Understanding style anatomy and function is essential for reproductive biology studies, for interpreting fertilization dynamics, and for diagnosing reproductive failures related to pollen tube growth impediments.
Griffel
Stempelgriffel
Stylus
pistil neck
pollen tube pathway
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 99-106 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 92-98 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 186-192.
Style
Syrah in Rhône Valley shows style penetration during mid-June Year N+1 at BBCH 60-65, with emerged pollen tubes breaking through stigma papillate surface and entering 2-4 mm into style transmitting tissue over 1-2.5 hours, establishing growth pathway in extracellular matrix-rich conducting tissue, growth rate 0.6-1.4 mm per hour during penetration phase, under warm Mediterranean conditions of 25-29°C temperature, adequate boron supporting tube wall integrity during penetration, typical of this mid-to-late season cultivar with 70-84% penetration success
Style penetration represents the initial tube growth phase when emerged pollen tube breaks through stigma papillate cell surface entering extracellular matrix of style transmitting tissue, establishing growth pathway from stigma surface into specialized conducting tissue enabling tube navigation toward ovary. This critical entry phase occurs during pollen tube growth in early-mid June in temperate regions Year N+1, typically proceeding over 1-3 hours following tube emergence as initial growth establishes, corresponding to BBCH 60-65 flowering stages. Penetration mechanism involves tube apex extending from stigma papillate cell surface, encountering and breaching interface between stigma and style tissues, navigating between or through cell layers entering extracellular matrix in transmitting tissue, and establishing growth pathway in specialized conducting tissue. Stigma-style interface characteristics show transitional zone between papillate stigma surface and underlying style transmitting tissue, cellular architecture changing from elongated papillate cells to more compact transmitting tissue cells, and interface representing barrier requiring tube penetration for style entry. Tube penetration pathway shows typical route between adjacent stigma cells exploiting natural intercellular spaces, tube secreting wall-modifying enzymes possibly facilitating passage through extracellular matrix, physical force from turgor pressure contributing to penetration through yielding tissue, and establishment of tube within extracellular spaces in transmitting tissue. Cell wall modification during penetration possibly involves pectinase and cellulase secretion from tube apex degrading extracellular matrix components, localized wall loosening facilitating tube passage, and controlled enzymatic activity enabling penetration without excessive tissue damage. Transmitting tissue entry shows tube reaching specialized conducting tissue in style central region, entering extracellular matrix-rich environment providing growth support, and transitioning from stigma surface growth to internal tissue growth mode. Transmitting tissue structure includes specialized parenchyma cells lining extracellular spaces providing nutritional and guidance support, lipid-rich secretions in extracellular matrix possibly providing energy source, abundant extracellular matrix providing physical pathway for tube growth, and overall tissue architecture optimized for pollen tube navigation. Extracellular matrix composition includes pectins providing hydrated gel-like environment, arabinogalactan proteins possibly involved in guidance and support, lipids secreted from transmitting tissue cells potentially providing nutrients, and various signaling molecules possibly guiding tube growth. Tube growth rate during penetration shows initial slower extension 0.5-1.2 mm per hour during stigma surface breakthrough and tissue entry, acceleration to 1-2 mm per hour once transmitting tissue pathway established, and transition to faster mid-style growth rates 2-3 mm per hour once fully established in conducting tissue. Penetration depth shows tube extending 2-5 mm from stigma surface into style during penetration phase, reaching transmitting tissue central zone, and establishing growth within conducting tissue for continued navigation toward ovary. Tube orientation during penetration shows generally downward growth from stigma surface toward style base and ovary, possible guidance by gravity or tissue architecture, and establishment of appropriate trajectory for continued style navigation. Multiple tube penetration per style shows 10-50 germinated tubes potentially entering style from stigma surface, penetration occurring relatively simultaneously for all germinated tubes, and establishment of multiple tubes in transmitting tissue creating competitive growth situation. Tube-tissue interactions during penetration include possible adhesion to extracellular matrix components, recognition of tissue-derived signals confirming appropriate pathway, and tube response to tissue environment affecting growth rate and direction. Stigma tissue response to penetration shows possible localized cellular changes in papillate cells at tube entry points though poorly characterized, continued secretion supporting tube growth, and overall permissive environment enabling penetration without defensive responses. Calcium signaling during penetration possibly includes calcium gradients in tissue attracting or guiding tubes, localized calcium accumulation at tube tip regulating growth processes, and calcium-dependent signaling affecting tube penetration efficiency. Environmental effects on penetration include temperature affecting metabolic activity and growth rate with optimal 22-28°C supporting efficient penetration within 1-2 hours, cool conditions 18-22°C slowing penetration requiring 2-4 hours, and warm conditions potentially accelerating penetration though excessive heat may impair function. Boron requirement for penetration shows continued importance with boron essential for tube wall integrity during mechanical penetration stress, deficiency causing tube bursting or growth failure during penetration preventing style entry, and adequate boron 10-20 ppm supporting normal penetration success. Pollen vigor effects show vigorous fresh pollen producing robust tubes completing penetration efficiently within 1-2 hours, weak or aged pollen showing delayed or failed penetration, and pollen quality affecting competitive success in reaching transmitting tissue. Failed penetration causes include inadequate tube growth vigor preventing breakthrough, abnormal stigma-style interface impeding entry, boron deficiency causing tube structural failure during penetration, environmental stress disrupting growth processes, or developmental abnormalities affecting tissue structure or tube function. Penetration success assessment shows successful penetration by 60-85% emerged tubes under favorable conditions, declining to 40-60% under suboptimal conditions, and penetration success rate affecting ultimate fruit set through determining proportion of tubes continuing toward ovary. The process represents first aspect of pollen tube growth establishing entry into style tissue, followed by transmitting tissue navigation and ovary approach. Duration typically spans 1-3 hours as tube extends from stigma surface into transmitting tissue establishing pathway. Cultivar variation in penetration may include stigma-style interface structure affecting penetration difficulty, tube vigor affecting penetration success, and tissue characteristics affecting penetration efficiency. Regional timing shows style penetration occurring during flowering period across all climates 1-5 hours following tube emergence as initial growth proceeds. Viticultural significance includes representing critical entry point from stigma to style enabling continued growth, requiring adequate tube vigor and tissue support, demonstrating continued boron importance for tube integrity, and establishing penetration success as prerequisite for tube navigation to ovary and fertilization. Research applications include analyzing penetration mechanisms and pathways, investigating tube-tissue interactions during penetration, evaluating environmental and nutritional effects on penetration success, quantifying penetration timing and efficiency, and assessing penetration success as determinant of continued tube growth and fertilization outcomes.
Style_Penetration is first aspect of Pollen_Tube_Growth, distinguished by breakthrough from stigma surface into style transmitting tissue. Represents critical transition establishing pathway for continued growth. Duration 1-3 hours as tube extends into style. Slower initial growth rate 0.5-1.2 mm per hour during tissue entry. Boron essential for tube integrity during mechanical stress. Success rate 60-85% under favorable conditions. Followed by Transmitting_Tissue_Navigation as growth continues through style.
Griffel-Eintritt
Narben-Griffel-Übergang
Style-Penetration
Transmissionsgewebe-Durchdringung
stigma-style transition
style entry
transmitting tissue penetration
tube breakthrough into style
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 385-387 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 290-292
Style Penetration
The sub-appellation designation Rutherford AVA identifies a 6,650-acre area nested within the larger Napa Valley AVA (approximately 45,000 acres), recognized for its distinctive Rutherford Bench alluvial fan soils that contribute the characteristic mineral-inflected tannin structure known colloquially as Rutherford Dust to Cabernet Sauvignon wines — similarly, within Chianti Classico DOCG, the recently established Unità Geografiche Aggiuntive (UGA) such as Greve in Chianti, Radda in Chianti, and Castelnuovo Berardenga function as sub-appellation designations that provide village-level origin specificity within the broader 71,800-hectare Chianti Classico zone.
An appellation designation that identifies a geographic unit nested within a larger parent appellation, delineating a more specific area recognized as possessing distinctive terroir characteristics that differentiate wines produced within it from those of the broader parent appellation, enabling finer geographic resolution in origin communication.
Sub_Appellation_Designation captures the progressive subdivision of viticultural territory into increasingly specific geographic units — a trend driven by both producer demand for terroir differentiation and consumer demand for origin transparency. The creation of sub-appellations is one of the most active areas of appellation regulation globally: Napa Valley's 16 nested sub-AVAs, Barolo's 181 Menzioni Geografiche Aggiuntive (established 2010), Chianti Classico's 11 UGA (established 2023), and the ongoing debates about sub-zone creation in regions from Priorat to the Willamette Valley all reflect the tension between geographic specificity (enabling terroir communication) and administrative complexity (creating potential consumer confusion and regulatory burden). The relationship between parent and sub-appellation is formally defined by geographic nesting — the sub-appellation must be entirely contained within the parent appellation's boundaries.
Teilgebietsbezeichnung
Unter-Appellationsbezeichnung
intra-appellation geographic unit
nested appellation name
sub-zone designation
verschachtelte Herkunftsangabe
TTB (2023) American Viticultural Areas, Alcohol and Tobacco Tax and Trade Bureau Consorzio Vino Chianti Classico (2023) Unità Geografiche Aggiuntive EU Regulation 1308/2013 Art. 93 on designations of origin
Sub Appellation Designation
Merlot sub-epidermal hypodermal layer position and structure= 2 cell layers located 28-34 μm (epidermis thickness) from berry surface, total sub-epidermal zone 35-48 μm thickness, with sub-epidermal cells receiving light penetrating through cuticle and epidermis supporting anthocyanin synthesis induction by UV-B and visible light, creating intensely pigmented 35-48 μm sub-epidermal band (3.2-3.9 mg/g anthocyanins) immediately beneath colorless 28-34 μm epidermis, with combined epidermis plus sub-epidermal hypodermis creating 63-82 μm outer skin zone distinct from unpigmented flesh beginning at 63-82 μm depth. Cabernet Sauvignon sub-epidermal hypodermal layer light-induced anthocyanin synthesis= cluster exterior sun-exposed berries showing sub-epidermal cells with high anthocyanin levels (3.8-4.8 mg/g) responding to high light intensity penetrating epidermis cluster interior shaded berries showing sub-epidermal cells with lower anthocyanins (2.4-3.2 mg/g, 30-40% reduction) receiving less light penetration due to shading by surrounding berries and leaves, demonstrating light regulation of anthocyanin synthesis specifically in light-exposed sub-epidermal position with shading reducing pigmentation even though sub-epidermal cells still present structurally. Syrah sub-epidermal hypodermal layer showing sharp spatial pigment gradient= sub-epidermal hypodermal cells (0-58 μm depth from epidermis) containing 4.5-5.8 mg/g anthocyanins outer mesocarp cells (58-120 μm depth) containing 0.5-1.5 mg/g anthocyanins declining with depth middle pericarp (>120 μm) containing <0.1 mg/g, demonstrating exponential decline in anthocyanin concentration from sub-epidermal maximum through transition zone to essentially zero in flesh creating sharp pigment gradient over 60-120 μm depth from sub-epidermal position. Chardonnay white cultivar sub-epidermal hypodermal layer showing non-anthocyanin pigmentation= 1-2 cell layers 26-44 μm beneath epidermis containing carotenoids and flavonols creating golden-yellow pigmentation concentrated in sub-epidermal position (0.3-0.6 mg/g carotenoids, 0.4-0.9 mg/g flavonols in sub-epidermal cells versus 0.1-0.2 mg/g carotenoids, 0.1-0.3 mg/g flavonols in middle pericarp), demonstrating that sub-epidermal pigment localization occurs in both anthocyanin-based (red) and carotenoid/flavonol-based (white) pigmentation systems with sub-epidermal position favoring pigment accumulation. Pinot Noir sub-epidermal hypodermal layer in berry cross-section= fresh berry cross-section showing thin 25-40 μm ruby-red pigmented sub-epidermal band immediately beneath colorless epidermis, creating obvious visual distinction between colored outer skin zone (epidermis plus sub-epidermal, total 45-68 μm) and colorless flesh, with thin sub-epidermal band (1-2 cell layers versus 2-3 in Cabernet Sauvignon) contributing to Pinot Noir characteristic lighter color and lower skin-to-pulp ratio affecting extraction requirements during winemaking.
A sub-epidermal hypodermal layer is the hypodermal layer emphasizing its position immediately beneath (sub-) the epidermis as the first cell layer(s) interior to the epidermis, focusing on position-dependent characteristics including direct contact with epidermal cells, exposure to light penetrating through epidermis and cuticle, and role as transition zone between protective outer tissues and storage inner tissues. Sub-epidermal hypodermal layer positional characteristics= (1) location as innermost component of outer pericarp/skin region, immediately interior to epidermis creating second (and sometimes third, fourth) cellular layer from berry surface (2) direct cell-cell contact with overlying epidermal cells through shared cell walls creating epidermal-hypodermal interface (3) separation from external environment by only epidermis (15-35 μm) and cuticle (2-8 μm) creating position 17-43 μm from berry surface (4) gradual transition to underlying outer mesocarp layers (if outer mesocarp distinguished from hypodermis) or direct transition to middle pericarp creating interior boundary of sub-epidermal zone. Sub-epidermal hypodermal layer light exposure characteristics= (1) reception of light penetrating through transparent cuticle and relatively transparent epidermal cells, with light intensity declining with depth through absorption and scattering (2) UV-B radiation (280-315 nm) partially absorbed by epidermal flavonols but significant UV still reaching sub-epidermal layers inducing protective responses (3) visible light penetration supporting chlorophyll-based photosynthesis during green berry phase (pre-veraison) in sub-epidermal cells (4) light quality shifting toward longer wavelengths (red, far-red) with depth as shorter wavelengths preferentially absorbed in outer layers. Sub-epidermal hypodermal layer as anthocyanin synthesis site= (1) spatial specificity of anthocyanin biosynthetic gene expression with VvMYBA1 transcription factor and structural genes (VvUFGT, VvDFR, VvF3H) upregulated specifically in sub-epidermal hypodermal cells at veraison creating cell-type-specific pigment synthesis (2) light regulation of anthocyanin synthesis with UV-B and blue light photoreceptors triggering signaling cascades activating anthocyanin genes specifically in light-exposed sub-epidermal cells (3) concentration of anthocyanins in sub-epidermal cell vacuoles creating intense localized pigmentation 2.5-6+ mg/g versus minimal levels (<0.1 mg/g) just 50-100 μm deeper in middle pericarp demonstrating sharp spatial gradient (4) anthocyanin distribution creating colored outer zone (epidermis plus sub-epidermal hypodermis, 35-95 μm depth) surrounding unpigmented flesh core. Sub-epidermal hypodermal layer developmental signaling= position-dependent signals potentially including light gradients, oxygen gradients, and diffusible signals from epidermis or seeds influencing sub-epidermal cell differentiation and metabolism potential epidermal-derived signals affecting sub-epidermal anthocyanin synthesis explaining why anthocyanins concentrate in sub-epidermal position sub-epidermal position creating intermediate environment between protective outer and storage inner tissues influencing cell fate determination during development. Sub-epidermal hypodermal layer in berry cross-section appearance= distinct colored band in red/black cultivars visible in fresh cross-sections as 20-60 μm thick intensely pigmented zone beneath colorless epidermis, surrounding unpigmented flesh golden-yellow band in white cultivars from carotenoid/flavonol accumulation pigmented sub-epidermal zone easily distinguished visually from colorless inner tissues creating obvious tissue zonation.
Sub_Epidermal_Hypodermal_Layer emphasizes the position immediately beneath epidermis and is distinguished from Supporting_Hypodermal_Layer which emphasizes structural functions. Sub-epidermal hypodermal layer represents the hypodermis from the perspective of position-dependent properties including light exposure and anthocyanin synthesis localization. Understanding sub-epidermal hypodermal layer is essential for comprehending spatial regulation of anthocyanin synthesis, light effects on pigmentation, and tissue zonation creating distinct outer pigmented and inner unpigmented pericarp regions fundamental to berry appearance and wine color potential.
Innere Hautschicht
direkt-subepidermale Schicht
immediately sub-epidermal layer
inner skin layer
subepidermale Hypodermis-Schicht
Castellarin, S.D. et al. (2007). Transcriptional regulation of anthocyanin biosynthesis. Plant Physiology 143=1347-1363 Downey, M.O. et al. (2006). Cultural practice and environmental impacts on flavonoid composition. American Journal of Enology and Viticulture 57=64-75.
Sub Epidermal Hypodermal Layer
Pinot Noir in Burgundy shows successful ovule fertilization during late June-October Year N+1 at BBCH 69-89, with 1.6-2.2 ovules per flower achieving complete double fertilization, embryo development progressing through globular (2 weeks), heart (5 weeks), torpedo (9 weeks), and mature (14 weeks) stages, endosperm supporting embryo growth, seed coat forming protective layers, auxin production driving berry expansion, under continental conditions, typical of this early-to-mid season cultivar producing 1-2 viable seeds per berry supporting normal berry development
Successful ovule fertilization represents the viable seed formation outcome when ovule achieves complete double fertilization producing functional diploid zygote and triploid endosperm, initiating coordinated embryo and endosperm development, stimulating integument growth forming seed coat, and proceeding through complete seed maturation supporting berry development and enabling plant reproduction. This favorable reproductive outcome occurs during fertilization and early seed development in late June-early July in temperate regions Year N+1, established within 1-3 days of pollination and progressing over subsequent 90-120 days, corresponding to BBCH 69-71 stages. Successful fertilization establishment shows pollen tube achieving micropyle penetration and sperm discharge, both sperm cells completing fusion with egg and central cell respectively, viable 2n zygote and 3n endosperm nucleus formation, and developmental activation initiating seed development program. Zygote viability indicators include normal diploid chromosome complement 2n=38 in Vitis vinifera, successful developmental activation with first division within 24-72 hours, and progressive embryo development through globular, heart, torpedo, and mature stages. Endosperm viability shows triploid nucleus 3n=57 successfully formed, free nuclear division initiating producing multinucleate syncytium, cellularization proceeding creating cellular endosperm tissue, and nutritive function supporting embryo growth throughout development. Embryo development progression shows first zygotic division producing two-cell embryo, subsequent divisions producing globular stage embryo by 2-3 weeks post-fertilization, heart stage with cotyledon primordia by 4-6 weeks, torpedo stage with elongating cotyledons and embryonic axis by 8-10 weeks, and mature embryo with complete organ primordia by 12-16 weeks ready for seed desiccation. Endosperm development shows rapid free nuclear division producing 32-128 nuclei within first weeks, cellularization forming cellular tissue providing nutrients, endosperm accumulating storage reserves including proteins and lipids, and progressive consumption by growing embryo reducing to thin layer in mature seed. Seed coat development shows integument layers differentiating into inner and outer seed coat, tissue lignification and coloration creating protective layers, mature seed coat providing mechanical protection and controlling water relations, and coat completion by seed maturation enabling desiccation tolerance. Seed growth dynamics show rapid enlargement during first 6-8 weeks coordinated with early berry growth, progressive accumulation of dry matter including storage proteins, lipids, and carbohydrates, maximum seed size achieved by 10-12 weeks post-fertilization, and final maturation preparing seed for desiccation and dormancy. Auxin production from successful seed shows hormone synthesis in developing embryo and endosperm beginning within days of fertilization, auxin levels correlating with seed developmental activity, hormone export to surrounding ovary tissues, and auxin as critical signal preventing flower abscission and stimulating berry growth. Seed-berry developmental coordination shows auxin from seed triggering ovary expansion, seed development providing hormonal support throughout berry growth, coordinated maturation timing with seed completing development as berry ripens, and seed presence essential for normal berry development in seeded cultivars. Seed number per berry shows successful fertilization typically producing 1-3 viable seeds from 2-4 ovules per flower, seed number variation depending on partial versus complete ovule fertilization success, and seed count affecting berry size with more seeds producing larger berries through greater auxin stimulus. Seed viability at maturity shows successful development producing viable propagule capable of germination, seed containing genetic recombination from sexual reproduction, and functional seed enabling sexual propagation though vegetative propagation more common in viticulture. Seed reserves accumulation shows storage protein deposition primarily in embryo cotyledons, lipid accumulation in embryo providing energy reserves, limited endosperm persistence with most consumed during embryo growth, and total dry matter typically 2-5% berry fresh weight. Seed desiccation and dormancy shows late-stage water loss reducing seed moisture from 60-70% to 10-20%, metabolic quiescence establishing dormancy preventing precocious germination, and dormancy mechanisms requiring stratification for germination. Environmental effects on successful seed development include adequate temperature 20-28°C supporting normal embryogenesis and maturation, sufficient nutrients from maternal vine providing resources, appropriate water relations maintaining seed hydration during growth then enabling desiccation, and overall favorable conditions supporting complete development. Temperature during seed development shows optimal range supporting steady progression through developmental stages, extreme heat potentially disrupting development, cool conditions slowing but generally not preventing maturation, and temperature as factor affecting development rate and timing. Nutrient supply shows maternal provision of nitrogen for protein synthesis, phosphorus and potassium for metabolism, and micronutrients supporting enzymatic functions, with adequate nutrition essential for normal seed development. Failed seed development after fertilization shows occasional embryo abortion despite initial fertilization success, early seed abortion creating seedless or seed-trace berries, and developmental failure reducing seed number below ovule fertilization success. Successful seed indicators visible in berry show hard textured seeds distinguishable from aborted ovules, brown color development indicating maturation, and seed rattling in dried berries confirming complete development. The process represents favorable ovule fertilization outcome within Ovule_Fertilization category. Duration spans 90-120 days from fertilization through complete seed maturation. Cultivar variation includes seed developmental characteristics, seed size and composition, and developmental timing. Regional timing shows successful seed development from late June through October across climates as seeds mature with berries. Viticultural significance includes providing auxin driving berry development, determining berry size through seed number effects, enabling sexual reproduction and genetic recombination supporting breeding, and affecting berry and wine characteristics through seed presence and composition. Research applications include analyzing seed development biology and regulation, investigating seed-berry coordination and hormonal interactions, evaluating factors affecting seed development success and characteristics, quantifying seed effects on berry development and quality, and assessing seed development as berry quality determinant.
Successful_Ovule_Fertilization represents favorable reproductive outcome within Ovule_Fertilization producing viable seeds. Characterized by complete double fertilization, viable zygote and endosperm, progressive embryo development, seed coat formation, auxin production. Duration 90-120 days from fertilization to mature seed. Typically 1-3 seeds per berry from partial ovule success. Auxin critical for preventing abscission and driving berry growth. Seed number affects berry size. Enables sexual reproduction. Essential for normal berry development in seeded cultivars.
complete seed development
erfolgreiche Samenanlage-Befruchtung
fertile ovule outcome
fruchtbares Samenanlage-Ergebnis
lebensfähige Samenbildung
successful seed maturation
viable seed formation
vollständige Samenentwicklung
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 408-410 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 310-312
Successful Ovule Fertilization
Succinic acid content quality in traditionally fermented Champagne base wines increases substantially during the primary alcoholic fermentation, with yeast production adding 0.3-0.8 g/L to the initial grape-derived concentration, followed by additional succinic acid release during the extended lees aging (tirage) as yeast autolysis liberates intracellular metabolites the complex, savory character contributed by succinic acid at final concentrations of 0.8-1.5 g/L complements the toasty, brioche notes from yeast mannoproteins, creating the multi-dimensional flavor profile that distinguishes aged Champagne from young sparkling wines.
Succinic_Acid_Content_Quality represents the chemical quality characterizing the concentration of succinic acid (butanedioic acid, C₄H₆O₄) in grape berry tissue and wine. Succinic acid occupies a unique position in wine acidity because while present in small amounts in grapes (0.2-1.0 g/L), it is substantially synthesized by yeasts during alcoholic fermentation, making it both a grape-derived and fermentation-derived acid. The post-fermentation concentration of succinic acid typically ranges from 0.5-1.5 g/L, reflecting both the initial grape content and yeast metabolic production. Succinic acid is a dicarboxylic acid and intermediate in the tricarboxylic acid cycle, produced during fermentation through the reductive branch of the TCA cycle when oxygen is limited. Yeast strain, fermentation temperature, nitrogen availability, and other factors influence the amount of succinic acid produced during fermentation. The sensory contribution of succinic acid is distinctive, described as a combination of sour, salty, and bitter notes that adds complexity to wine flavor. This unusual taste profile distinguishes succinic acid from the pure sourness of tartaric and malic acids. Succinic acid is relatively stable in wine, not metabolized by lactic acid bacteria under normal conditions, making it a persistent component of finished wine acidity. The accumulation of succinic acid in grapes follows patterns similar to other TCA cycle intermediates, with synthesis in green tissues and modest accumulation in berries. Measurement employs specific enzymatic assays using succinyl-CoA synthetase or HPLC separation, distinguishing succinic acid from the much larger tartaric and malic acid pools.
Succinic_Acid_Content_Quality addresses an acid with both grape and fermentation origins, requiring consideration of both viticultural and enological factors. The sour-salty-bitter taste profile of succinic acid is unusual among organic acids and contributes to wine complexity at concentrations well below the threshold for tartaric or malic acid. Succinic acid exists as the diprotic acid and its sodium, potassium, and calcium salts in wine. The TCA cycle position of succinic acid (between α-ketoglutarate and fumarate) places it in the central metabolic pathway shared by grape and yeast metabolism.
Bernsteinsäure
Bernsteinsäuregehalt
Succinat
Succinatkonzentration
amber acid quality
butanedioic acid concentration
succinate content
succinic level
Lambrechts MG and Pretorius IS (2000) Yeast and Its Importance to Wine Aroma: A Review. South African Journal of Enology and Viticulture 21(1): 97-129 Chidi BS et al. (2015) Organic Acid Metabolism and the Impact of Fermentation Practices on Wine Acidity: A Review. South African Journal of Enology and Viticulture 36(3): 316-330
Succinic Acid Content Quality
In a Burgundy Chardonnay vineyard, sucrose cleavage activation was characterized in early August Year N+1 (BBCH 83) when VvcwINV expression increased 68-fold, VvGIN1 showed 42-fold elevation, total invertase activity rose from 6.2 to 58.5 units/g FW, apoplastic hexoses measured 245 mM combined, hexose:sucrose ratio increased from 0.5 to 4.2, berry Brix 12.5° rising 2.1% per week, under moderate Burgundian conditions (daily high 24-27°C, night 14-16°C, adequate water enabling metabolic activation).
Sucrose_Cleavage_Activation represents the induction of invertase enzymes that hydrolyze phloem-delivered sucrose into glucose and fructose enabling hexose accumulation, creating the critical biochemical step that converts transported disaccharide to storable monosaccharides establishing berry sweetness character distinct from other fruits that accumulate sucrose. This activation occurs during veraison onset to early-mid veraison (BBCH 83-84), typically 0-3 days following or concurrent with phloem unloading activation, and is characterized by cell wall invertase gene expression (VvcwINV) increasing 30-90 fold above green berry baseline, vacuolar invertase genes (VvGIN1, VvGIN2) showing 20-70 fold elevation, combined invertase enzyme activities rising from pre-veraison 3.0 indicating dominant sucrose hydrolysis. Molecular mechanisms of activation inclu VvcwINV gene family members (VvcwINV1, VvcwINV2, multiple isoforms) coordinately induced with peak expression during mid-veraison producing apoplastic enzyme (pH optimum 4.5-5.2, Km for sucrose 10-20 mM) that cleaves sucrose released during apoplastic unloading, vacuolar invertase genes VvGIN1/GIN2 upregulated producing intracellular enzyme (pH optimum 5.0-6.0, Km 15-35 mM) hydrolyzing any sucrose entering cells or synthesized de novo, regulatory control through sugar-responsive transcription factors and ABA signaling with invertase gene promoters containing sugar-response elements and ABA-responsive elements enabling coordinate induction with other ripening genes, and post-translational regulation including possible invertase inhibitor proteins modulating activity though grape inhibitors less well-characterized than in other species. Enzyme properties of activated invertases show: cell wall invertase optimal activity pH 4.5-5.2 matching apoplastic pH, thermal stability with activity maintained at berry temperatures 20-35°C though declining >35°C through denaturation, sucrose specificity with minimal activity on other substrates (raffinose, stachyose cleaved 200-300 mM creating potential feedback regulation though typically not limiting given rapid hexose transport removing products, and kinetic parameters with Vmax 60-140 μmol sucrose cleaved g⁻¹ h⁻¹ at full activation adequate for observed sugar accumulation rates. Subcellular and tissue localization includes: cell wall invertase concentrated in phloem-mesocarp boundary zone particularly bundle sheath and adjacent parenchyma where apoplastic unloading occurs, immunolocalization detecting enzyme in cell walls and intercellular spaces consistent with apoplastic function, vacuolar invertase distributed throughout mesocarp cell vacuoles enabling intracellular sucrose hydrolysis, and developmental progression showing enzyme spreading from vascular regions into outer mesocarp zones as ripening advances. Substrate and product dynamics demonstrate: apoplastic sucrose concentration 100-250 mM transiently accumulating before cleavage providing substrate for cell wall invertase, resulting apoplastic hexoses 150-350 mM combined (glucose + fructose) creating concentration gradient for membrane transport, vacuolar sucrose maintained low 32°C may impair through reduced gene expression or protein synthesis, light affecting developmental progression and potentially invertase gene expression through photoreceptor pathways though effects less direct than on photosynthesis and sugar supply, water status with moderate deficit potentially enhancing activation through elevated ABA while severe deficit may delay through general developmental arrest, and nutritional status particularly nitrogen affecting protein synthesis capacity for invertase production. Cultivar variation in activation characteristics shows: early-activating varieties (Chardonnay, Pinot Noir) inducing invertase 7-9 weeks post-flowering reaching 60-90 units/g activity within 4-6 days, moderate cultivars (Merlot, Cabernet Sauvignon) activating 9-11 weeks with similar final activities achieved in 6-9 days, and late types (Nebbiolo, Mourvèdre) showing activation 11-13 weeks requiring 8-12 days for full induction reflecting genetic differences in developmental timing and enzyme induction kinetics. Alternative sucrose metabolism through sucrose synthase shows: sucrose synthase (VvSUS genes) catalyzes reversible reaction sucrose + UDP ⇌ UDP-glucose + fructose, enzyme activity present throughout berry development including ripening though relative importance versus invertase shifts, sucrose synthase potentially important for producing UDP-glucose substrate for cellulose synthesis, starch synthesis (minor in grape), or other glucosylation reactions, and possible compartmentation with sucrose synthase cytoplasmic while invertases apoplastic or vacuolar enabling coexistence of pathways serving different functions. Physiological consequences of invertase activation inclu hexose accumulation versus sucrose enabling higher final sugar concentrations due to reduced osmotic coefficient and lower crystallization risk, glucose and fructose creating distinct flavor profiles versus sucrose contributing to characteristic grape and wine taste, hexose metabolism differently regulated than sucrose affecting respiratory and fermentative pathways during ripening and winemaking, and hexose:sucrose ratio diagnostic for ripening stage and cultivar enabling maturity assessment. Quality implications inclu invertase activity levels correlating with final sugar achieved (r² = 0.68-0.84) with inadequate activation limiting Brix potential, timing of activation relative to other ripening processes affecting compositional balance important for wine style, invertase activity potentially affecting wine fermentation kinetics through glucose:fructose ratio impacts on yeast sugar utilization, and residual invertase activity in harvested grapes potentially affecting must and wine composition through continued sucrose hydrolysis during processing. Research and practical applications involve: invertase activity assays providing biochemical marker for veraison onset and ripening progression, gene expression analysis tracking VvcwINV and VvGIN transcripts indicating developmental stage, inhibitor studies exploring potential viticultural interventions to modulate sugar accumulation timing or extent, and comparative analysis across cultivars and conditions revealing genetic and environmental control of this critical ripening process.
Sucrose_Cleavage_Activation is distinguished from Phloem_Unloading_Activation by representing enzymatic hydrolysis rather than transport pathway change, from Hexose_Transport_Initiation by focusing on substrate production rather than product transport, and from rapid accumulation by showing activation rather than maximum rates. Invertase induction: cell wall 30-90 fold, vacuolar 20-70 fold. Activity increases 6-12 fold. Timing: 0-3 days with unloading activation. Hexose:sucrose ratio shift 3.0 diagnostic. Sucrose_Cleavage_Activation represents critical biochemical conversion enabling grape hexose accumulation character.
Disaccharid-Spaltungs-Onset
Hexose-Generierungs-Initiierung
Invertase-Gen-Induktion
Saccharose-Hydrolyse-Aktivierung
disaccharide cleavage onset
hexose generation initiation
invertase gene induction
sucrose hydrolysis activation
Davies C, Robinson SP (2000) Differential screening indicates a dramatic change in mRNA profiles during grape berry ripening Plant Physiol 122:803-812 invertase gene expression Moscatello S, Famiani F, Proietti S, Farinelli D, Battistelli A (2011) Sucrose synthase dominates carbohydrate metabolism and relative growth rate in growing kiwifruit Sci Hort 128:197-205 sucrose metabolism comparison Ranwala AP, Suematsu C, Masuda H (1998) Purification and characterization of two forms of invertase from grape Phytochem 49:331-337 enzyme characterization.
Sucrose Cleavage Activation
Sucrose content quality in Thompson Seedless table grapes selected for dried fruit production shows characteristic low values (0.5-2.0 g/L) reflecting the high invertase activity that ensures efficient sugar compartmentalization as glucose and fructose however, recently developed seedless table grape cultivars for fresh market consumption occasionally display elevated sucrose (5-10 g/L) associated with selection for crisp flesh texture, as reduced invertase activity appears linked to firmer cell walls, creating an unexpected quality trade-off between sugar composition and textural properties.
Sucrose_Content_Quality represents the chemical quality characterizing the concentration of sucrose (saccharose, table sugar) in grape berry tissue. Sucrose is a disaccharide composed of one glucose and one fructose molecule joined by an α-1,β-2 glycosidic bond, representing the primary form of carbon transported through the phloem from photosynthesizing leaves to developing berries. Despite its importance as the transported sugar, sucrose accumulates to only low concentrations in ripe grape berries (typically less than 5 g/L, often less than 1 g/L) because the high activity of cell wall-bound acid invertase and vacuolar invertase rapidly hydrolyzes arriving sucrose into glucose and fructose. This efficient sucrose cleavage is a characteristic feature of grape berry sugar accumulation that distinguishes grapes from fruits like peaches and bananas where sucrose comprises a larger fraction of total sugars. Sucrose concentration relative to hexose concentration provides information about invertase activity and the balance between sucrose import and hydrolysis. Elevated sucrose in ripe berries may indicate compromised invertase activity from extreme stress conditions, genetic variation in invertase expression, or sampling shortly after phloem unloading. In table grape varieties, moderate sucrose retention can contribute to desirable textural properties. During winemaking, any sucrose present is rapidly hydrolyzed by yeast invertase before fermentation of the resulting glucose and fructose. Chapitalization (sugar addition to increase alcohol potential) in permitted regions typically employs sucrose, which is inverted during fermentation. Sucrose measurement uses specific enzymatic assays with invertase followed by glucose determination, or HPLC separation and quantification.
Sucrose_Content_Quality is positioned as a subclass of Non_Reducing_Sugar_Quality and represents the primary non-reducing sugar in grape tissue. The low sucrose concentration in ripe berries despite sucrose being the transport sugar reflects the metabolic transformation occurring upon phloem unloading. Sucrose content may have greater relative importance in other grapevine organs including roots (starch mobilization), leaves (photosynthate pools), and phloem sap (transport concentration). The analysis of sucrose requires specific methods that distinguish it from glucose and fructose, as non-specific sugar assays combine all forms. The position of sucrose in source-sink relationships makes it a useful marker of carbon allocation patterns.
Haushaltszuckerkonzentration
Rohrzuckergehalt
Saccharosegehalt
cane sugar concentration
disaccharide sugar quality
saccharose content
table sugar level
Zhang XY et al. (2006) A Shift of Phloem Unloading from Symplasmic to Apoplasmic Pathway is Involved in Developmental Onset of Ripening in Grape Berry. Plant Physiology 142(1): 220-232 Davies C and Robinson SP (1996) Sugar Accumulation in Grape Berries: Cloning of Two Putative Vacuolar Invertase cDNAs and Their Expression in Grapevine Tissues. Plant Physiology 111(1): 275-283
Sucrose Content Quality
In a Sonoma Chardonnay vineyard, sugar accumulation was tracked over 58 days from early August through late September Year N+1 (BBCH 83-89) when berry Brix increased from 8.2° to 24.6° (16.4° total gain, average 2.8% per week), showing initial phase 8.2° → 13.5° (days 0-14, 2.1%/week), active phase 13.5° → 22.8° (days 14-42, 3.8%/week peak), deceleration 22.8° → 24.6° (days 42-58, 1.3%/week), berry fresh weight 0.95 g → 1.88 g (98% expansion), vacuolar hexose 280 → 1620 mM, under typical California coastal conditions (daily high 23-28°C supporting sustained accumulation).
Sugar_Accumulation represents the sustained hexose import and vacuolar storage throughout berry ripening creating the progressive sweetness development from veraison onset through harvest that defines grape maturity, determines wine alcohol potential, and provides osmotic driving force for berry expansion constituting the dominant anabolic process of ripening berry metabolism. This accumulation extends throughout veraison and ripening phases (BBCH 83-89), typically beginning 0-3 days post-veraison onset and continuing 40-70 days until harvest, and is characterized by berry Brix increasing from veraison onset 6-10° to harvest 18-30° (cultivar and region dependent) representing 12-24° Brix total gain, total berry sugars rising from 80-140 g/L to 180-300 g/L constituting 100-180 g/L hexose accumulation, vacuolar glucose and fructose each increasing from 150-300 mM to 700-1100 mM creating combined concentrations 1400-2200 mM (25-40% w/v) dominating berry osmotic potential, and berry fresh weight expanding 50-120% from 0.8-1.4 g to 1.4-2.8 g through osmotic water import driven by sugar accumulation demonstrating tight coupling between hexose storage and berry development. Temporal pattern of accumulation demonstrates sigmoid progression with initial acceleration phase (days 0-12 post-veraison) showing 1.2-2.5% Brix per week as transport machinery activates, active accumulation phase (days 12-35) exhibiting peak rates 2.8-5.0% Brix per week when all pathway components operate maximally, deceleration phase (days 35-55) displaying declining rates 1.2-2.8% Brix per week as vacuolar concentrations approach equilibrium with phloem supply, and plateau phase approaching harvest (days 55-70) showing minimal increase 32-35°C impairing 40-70% through enzyme denaturation and metabolic disruption, light operating primarily through source photosynthesis providing sucrose substrate with adequate canopy illumination essential for sustained accumulation, water status with moderate deficit compatible or potentially enhancing concentration through reduced berry expansion while severe deficit (<-1.5 MPa stem ψ) impairing 35-65% through impaired phloem delivery and metabolism, and nutritional status particularly nitrogen for enzyme synthesis and potassium for sugar-acid balance affecting accumulation capacity and final levels. Cultivar genetic variation demonstrates high-Brix varieties (Gewürztraminer, Sémillon potential 26-30° Brix) achieving rapid accumulation 4.2-5.5% Brix per week peak with high transporter expression and extended duration 45-55 days reaching maximum capacity, moderate cultivars (Chardonnay, Cabernet Sauvignon 22-26° Brix) showing 3.5-4.8% per week peak over 50-65 days achieving typical commercial levels, and lower-Brix types (Pinot Noir cool climates 20-24° Brix, table grapes 18-22° Brix) displaying 2.8-4.0% per week for 55-70 days reflecting genetic differences in transporter capacity, metabolic programming, and final sugar set-points. Osmotic consequences of accumulation include vacuolar osmotic potential declining from -1.4 to -2.0 MPa (veraison onset) to -2.6 to -3.6 MPa (harvest) creating strong water potential gradient, berry water potential following vacuolar sugar causing decline from -1.5 to -2.0 MPa to -2.2 to -3.2 MPa establishing gradient favoring water import from vine xylem (-0.5 to -1.2 MPa), and osmotically-driven water uptake causing berry expansion 60-130% despite declining growth rates illustrating hydraulic component of berry development where sugar accumulation drives size increase. Coordination with other ripening processes shows acid decline occurring simultaneously with sugar accumulation creating developing balance, berry softening largely completed by mid-accumulation period establishing characteristic ripe texture, color development in reds coordinated through common ABA signaling reaching completion during late accumulation, and flavor-aroma compound synthesis intensifying throughout accumulation with esters, terpenes, and volatiles accumulating creating integrated sensory ripeness. Carbon and energy budget implications demonstrate sugar accumulation representing net carbon import 120-220 g glucose-equivalent per kg berries over ripening period (based on 180-300 g/L final sugar), respiratory carbon loss through concurrent acid catabolism 15-35 g glucose-equivalent per kg berries representing moderate respiratory cost balanced by stored sugars, net carbon gain 100-185 g per kg berries enabling berry growth and quality compound synthesis, and energetics where imported hexoses partially respired (20-35% of import) providing ATP for transport and biosynthesis while majority (65-80%) accumulates in vacuoles creating efficient dual metabolism. Quality implications of accumulation include final sugar levels determining wine alcohol potential with each 1° Brix yielding approximately 0.55-0.60% ethanol post-fermentation, sugar-acid balance critically affecting wine style with ratios 20-35:1 (Brix:TA g/L) typical for quality wines, uniformity of accumulation across berries and clusters determining harvest maturity consistency affecting wine composition and requiring selective harvest if heterogeneous, timing of accumulation relative to phenolic and flavor development defining optimal harvest window requiring integration of multiple parameters, and accumulation rate patterns revealing environmental stress or management issues enabling diagnostic assessment and intervention. Harvest timing based on accumulation involves target sugar levels varying by style with table wines typically 20-24° Brix, fuller-bodied styles 22-26° Brix, late harvest or dessert wines 24-32° Brix depending on variety and region, weekly Brix sampling during ripening tracking rates and projecting harvest timing, integration with acid, pH, phenolic, and sensory parameters ensuring comprehensive maturity determination, and recognition that continued accumulation post-phenolic maturity may require earlier harvest accepting lower sugar to preserve balance particularly warm vintages. Viticultural management affecting accumulation includes crop load optimization with typical ratios 0.8-1.2 kg fruit per m² leaf area enabling adequate source capacity supporting target sugars without excessive competition, canopy management maintaining healthy productive foliage throughout ripening providing sustained photosynthesis, irrigation strategy with adequate water supporting continued accumulation without excess promoting vigor or berry dilution, disease control preventing infections that disrupt metabolism impairing accumulation, and harvest logistics preparing for optimal picking timing as accumulation trajectory indicates approaching maturity. Physiological limits to accumulation include maximum vacuolar concentrations ~2200-2600 mM approaching saturation potentially limiting further accumulation, membrane integrity constraints at extreme osmotic potentials <-3.6 MPa potentially compromising function, cell wall mechanical limits constraining vacuolar expansion beyond certain size, metabolic feedback regulation where high vacuolar sugar potentially inhibits further import through unknown mechanisms maintaining cultivar-specific set-points, and source photosynthesis capacity ultimately limiting total carbon available for accumulation determining yield-quality relationships. Research applications include transporter expression analysis revealing molecular control of accumulation, flux studies using isotope tracers quantifying import rates and metabolic partitioning, osmotic potential measurements characterizing thermodynamic driving forces, and modeling approaches integrating source-sink relationships predicting accumulation dynamics enabling precision viticulture optimization.
Sugar_Accumulation is distinguished from Sugar_Accumulation_Onset by representing entire accumulation process (40-70 days, 12-24° Brix gain) rather than onset (6-10 days, 5-8° Brix initial), encompasses both Active_Sugar_Accumulation and Peak_Sugar_Accumulation as organizational parent, and represents complete trajectory from veraison through harvest. Duration BBCH 83-89. Final levels: high-Brix 26-30°, moderate 22-26°, lower 18-24°. Dominant anabolic process of ripening. Sugar_Accumulation represents fundamental developmental process determining grape maturity and wine quality potential.
Brix increase
Brix-Zunahme
Hexose-Speicherung
Süße-Entwicklung
hexose storage
sweetness development
vacuolar sugar accumulation
vakuoläre Zucker-Akkumulation
Keller M (2020) The Science of Grapevines 3rd Ed pages 221-228 sugar accumulation comprehensive Coombe BG (1992) Research on development and ripening of the grape berry Amer J Enol Vitic 43:101-110 accumulation patterns Dai ZW, Ollat N, Gomès E, Decroocq S, Tandonnet JP, Bordenave L, Pieri P, Hilbert G, Kappel C, van Leeuwen C, Vivin P, Delrot S (2011) Ecophysiological, genetic, and molecular causes of variation in grape berry weight and composition Plant Physiol 155:519-532 accumulation variation.
Sugar Accumulation
Sugar accumulation data for Cabernet Sauvignon clone 337 versus clone 169 at Oakville, Napa Valley tracked weekly from véraison through harvest (2015-2020), revealing that clone 337 initiates rapid sugar accumulation 5-7 days earlier (reaching 20° Brix by week 3 post-véraison vs. week 4 for clone 169), achieves steeper accumulation rates (0.35° Brix/day vs. 0.28° Brix/day during the linear phase), and consistently reaches 24.5° Brix target 8-12 days before clone 169 — a significant phenological advance influencing harvest scheduling in multi-clone plantings. Sugar accumulation data for Furmint clones at the Tokaj Research Station documents clone F-85 reaching 22° KMW by October 5 (±8 days across vintages) versus clone F-42 at 20° KMW on the same date, with clone F-85's earlier attainment of sugar targets enabling either earlier harvest for dry Furmint production or extended hang time for Aszú berry development.
Sugar accumulation data is a quality performance data entity that records the dynamics and final concentration of soluble sugars (predominantly glucose and fructose) in the must of a clonal line, measured at specified phenological stages and at harvest, expressed in degrees Brix, Baumé, Oechsle, KMW (Klosterneuburger Mostwaage), or potential alcohol (% vol.), and including maturity curve parameters such as accumulation rate, onset of rapid sugar loading, and the sugar-acidity crossover date that collectively characterize the ripening kinetics and harvest window of the clone.
Sugar accumulation data provides the most widely used single metric of clone ripening performance, yet its interpretation requires integration with acidity, phenolic, and aromatic maturity data to avoid the trap of sugar-centric harvest decisions that can decouple alcohol level from flavor development. The kinetics of sugar accumulation — onset, rate, plateau, and duration — are as informative as the final concentration, because clones with identical harvest sugar levels may achieve them through different temporal trajectories with distinct implications for simultaneous flavor maturity. The glucose-to-fructose ratio during ripening provides additional clone characterization, as this ratio decreases from approximately 3:1 at véraison to near 1:1 at full maturity, and deviations may indicate incomplete physiological ripeness despite adequate total sugar. Temperature sensitivity of sugar accumulation differs among clones — some maintain linear accumulation rates through heat events while others show sugar loading stalls above 35°C, a characteristic of increasing importance under climate change. The measurement scale should be documented precisely, as conversions between Brix, Baumé, Oechsle, and KMW are not perfectly linear and small systematic errors accumulate in multi-vintage datasets. Modern sugar accumulation monitoring using on-vine NIR sensors enables non-destructive high-frequency tracking that captures intra-block spatial variation in ripening dynamics.
Mostgewichtsdaten
Mostwertentwicklung
Reifeentwicklung Zucker
Zuckerakkumulationsdaten
Zuckeranreicherungsdaten
must weight data
ripening sugar data
soluble solids data
sugar content data
sugar maturity data
Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 369-395 Coombe BG (1992) Research on development and ripening of the grape berry. American Journal of Enology and Viticulture 43(1): 101-110 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed, Wiley, pp.: 65-88
Sugar Accumulation Data
Sugar accumulation in grape berries follows a sigmoidal pattern beginning at veraison, with accumulation rates of 0.1-0.3°Brix per day during active ripening. Final sugar concentrations range from 16-18°Brix in cool climate table grapes to 24-28°Brix in warm climate wine grapes harvested for high-alcohol wines. Cultivar differences in sugar accumulation capacity span 3-6°Brix at equivalent thermal time from veraison, with Grenache and Zinfandel showing high accumulation (24-28°Brix typical) and Riesling showing moderate accumulation (20-23°Brix at flavor maturity). Sugar composition shifts from glucose-dominant early in ripening (G/F ratio 2-3) to near-equal at maturity (G/F ratio 0.95-1.05), with cultivar-specific variation in final ratio.
A quality potential disposition that is the inherited capacity of a Vitis vinifera plant to transport and accumulate hexose sugars (glucose and fructose) in ripening berries, determined by source leaf photosynthetic capacity, phloem loading and transport efficiency, berry sink strength, and the expression of sugar transporter and invertase genes controlling berry sugar import and storage.
Sugar_Accumulation_Disposition serves as the parent class for specific sugar accumulation capacity categories. Sugar concentration is the primary determinant of potential alcohol in wine and significantly affects wine body, sweetness perception, and microbial stability. The trait integrates whole-vine carbon economy with berry-level transport and storage mechanisms. Climate warming is advancing sugar ripeness relative to flavor and phenolic maturity in many regions, creating challenges for balanced wine production.
Beerenreife-Disposition
Brix accumulation trait
Zuckerakkumulation
Zuckereinlagerung
berry sweetening disposition
sugar loading capacity
Coombe BG, McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6:131-135 Dai ZW et al. (2013) Ecophysiological, genetic, and molecular causes of variation in grape berry weight and composition: a review. American Journal of Enology and Viticulture 64:1-24
Sugar Accumulation Disposition
In a Marlborough Sauvignon Blanc vineyard in New Zealand, sugar accumulation onset was detected in early February Year N+1 (BBCH 83, 64 days post-flowering) when berry Brix increased from 7.2° to 11.8° over 7 days (0.66° per day, 4.6% per week), VvHT1 expression rose 65-fold, berry firmness declined 15% to 2.4 N indicating concurrent softening, ABA levels reached 720 ng/g, and hexose:sucrose ratio increased from 0.4 to 2.8 under warm conditions (daily high 24-27°C, adequate irrigation supporting rapid metabolic transition).
Sugar_Accumulation_Onset represents the developmental transition during veraison when berry metabolism shifts from predominantly organic acid accumulation and low sugar content characteristic of green berries to rapid hexose accumulation that transforms berry composition from acidic to sweet creating the fundamental quality attributes essential for wine production. This critical metabolic switch occurs during early-to-mid veraison (BBCH 83-84), typically synchronized within 0-3 days of color break in red cultivars or concurrent with initial berry softening in white varieties, and is characterized by sugar concentration increasing from pre-veraison baseline 5-8° Brix to early accumulation 10-14° Brix within 5-8 days representing transition from 2.0 at onset and eventually >8-15 at maturity reflecting complete metabolic reprogramming. Hormonal regulation of sugar accumulation onset shows: abscisic acid levels spike from pre-veraison 50-150 ng/g to veraison peak 500-2000 ng/g activating sugar-responsive genes through ABA-responsive elements and creating hexokinase-mediated sugar signaling feedback loops that amplify accumulation, declining gibberellin and auxin levels remove growth-related suppression allowing metabolic resources to redirect from cell division and expansion to storage compound accumulation, and ethylene shows cultivar-dependent slight elevation (0.1-0.6 nL g⁻¹ h⁻¹) in some varieties potentially modulating sugar metabolism timing though its role is less pronounced than in climacteric fruit. Gene expression changes at onset inclu hexose transporter genes (VvHT1, VvHT2) show 30-100 fold induction enabling glucose and fructose uptake into cells, sucrose transporter genes paradoxically decline as apoplastic unloading reduces symplastic transport requirements, invertase genes both vacuolar (VvGIN) and cell wall (VvcwINV) increase 15-50 fold catalyzing sucrose hydrolysis, sucrose synthase shows tissue-specific regulation with some isoforms increasing to produce UDP-glucose for cellulose synthesis while others decline, and transcription factors including VvMYB, VvNAC, and VvWRKY families show coordinated induction orchestrating the metabolic reprogramming. Cellular mechanisms enabling sugar accumulation inclu mesocarp cell vacuoles expanding to occupy 90-95% of cell volume creating massive storage capacity (individual cell vacuoles reaching 0.5-2.0 nL volume), tonoplast membrane area increasing substantially through vesicle fusion enabling enhanced transporter capacity, vacuolar pH regulation maintaining 3.0-3.8 creating proton gradient that may assist some sugar transport mechanisms, and cytoplasmic sugar concentrations kept low (50-100 mM) relative to vacuolar accumulation (800-1500 mM at maturity) through efficient tonoplast transport maintaining concentration gradient favoring continued import. Environmental factors influencing onset timing inclu temperature with warm conditions 25-30°C advancing onset 3-5 days compared to cool 18-22°C through enhanced metabolic rates and potentially earlier ABA accumulation, light exposure affecting onset through photosynthate availability with shaded fruit sometimes showing 2-4 day delay, water availability with moderate deficit potentially advancing onset 2-3 days through elevated ABA though severe stress can delay through general metabolic impairment, and canopy source-sink relationships with adequate leaf area per unit fruit weight essential for supporting rapid accumulation once initiated. Cultivar variation in onset timing shows: early varieties (Chardonnay, Pinot Noir, Gamay) initiate sugar accumulation 7-9 weeks post-flowering, mid-season cultivars (Merlot, Syrah, Cabernet Franc) begin 9-11 weeks post-flowering, while late varieties (Cabernet Sauvignon, Nebbiolo, Mourvèdre) start 11-13 weeks post-flowering reflecting genetic differences in developmental programming and veraison triggering. Coordination with other veraison processes demonstrates: sugar accumulation onset typically occurs 0-3 days after or concurrent with initial berry softening suggesting common developmental signals, coincides with color break in red cultivars (both triggered by ABA spike and sugar signaling), parallels acid decline initiation particularly malate respiratory consumption beginning, and synchronizes with berry expansion resumption after lag phase creating the characteristic veraison growth acceleration. Metabolic consequences of sugar accumulation onset inclu respiratory patterns shifting from primarily oxidative metabolism supporting growth to substrate-level reactions processing imported sugars, carbon allocation redirecting from structural components (cell walls, membranes) to storage carbohydrates representing 60-80% of berry carbon at maturity, and osmotic adjustment through hexose accumulation creating water potential gradient (-1.5 to -2.5 MPa at maturity) driving continued water import and berry expansion despite declining growth rates. Nutritional demands during onset inclu adequate nitrogen for synthesis of sugar transporters and metabolic enzymes with 15-30% of berry nitrogen allocation occurring during early sugar accumulation, potassium requirements increasing as K⁺ accumulates with sugars potentially affecting membrane transport and pH regulation (berry K⁺ rising from 800-1200 mg/kg to 1500-2500 mg/kg during accumulation), and phosphorus for ATP synthesis supporting active transport processes though P requirements relatively modest. Quality implications of sugar accumulation onset timing inclu early onset relative to other maturity parameters can lead to high sugar with insufficient flavor development if harvest timing based solely on Brix targets, delayed onset may produce fruit with advanced physiological age but inadequate sugar requiring extended hang time with risks of over-ripeness in other aspects, and synchrony of onset across berries and clusters within vineyard affects maturity uniformity and harvest timing optimization. Viticultural factors affecting onset inclu crop load with excessive yields (>12-15 kg/vine depending on variety) potentially delaying onset 3-7 days through source limitation and delayed veraison triggering, leaf area adequacy with insufficient photosynthetic capacity limiting sugar import potential once onset occurs, water management with irrigation strategy during pre-veraison affecting ABA accumulation and onset timing (moderate deficit potentially advancing 2-4 days), and canopy microclimate with cluster light exposure and temperature affecting developmental signals triggering onset. Monitoring sugar accumulation onset involves: regular Brix measurements showing acceleration from 1.5% increase per week indicating onset occurrence, berry sampling across developmental stages within clusters identifying first berries initiating rapid accumulation, correlation with berry softening and color break providing integrated veraison monitoring, and growth degree day accumulation from flowering predicting onset timing enabling proactive management and harvest forecasting.
Sugar_Accumulation_Onset is distinguished from later accumulation phases by representing initiation rather than progression, occurs typically 0-3 days after/with color break (reds) or initial softening (whites), and marks fundamental metabolic shift from acid to sugar accumulation. Pre-onset rates <1.0% Brix/week, post-onset 1.5-3.5% Brix/week. Timing: early cultivars 7-9 weeks post-flowering, mid-season 9-11 weeks, late 11-13 weeks. Temperature optimal 25-30°C for rapid onset. ABA spike 50-150 → 500-2000 ng/g is critical trigger. Sugar_Accumulation_Onset represents universal veraison marker for ALL grape cultivars (red and white) essential for harvest timing prediction.
Hexose-Akkumulations-Beginn
Süßungs-Prozess-Onset
Veraison-Zucker-Beladungs-Initiierung
Zucker-Import-Aktivierung
hexose accumulation commencement
sugar import activation
sweetening process onset
veraison sugar loading initiation
Keller M (2020) The Science of Grapevines 3rd Ed pages 217-225 sugar accumulation and veraison Coombe BG (1992) Research on development and ripening of the grape berry Australian J Grape Wine Res 48:125-144 sugar accumulation dynamics Davies C, Robinson SP (1996) Sugar accumulation in grape berries Plant Physiol 111:275-283 molecular mechanisms.
Sugar Accumulation Onset
Sugar accumulation plateau quality comparison in Napa Valley evaluated post-optimal Brix trajectories for major red varieties during extended hangtime trials. Cabernet Sauvignon demonstrated excellent sugar accumulation plateau quality, reaching stable Brix plateau at approximately 26°Brix with subsequent changes of only 0.08°Brix per day over the following three weeks attributed to the balance between minimal continued import and respiratory consumption, permitting extended hangtime for tannin maturation without progressive alcohol escalation. Merlot showed intermediate plateau quality with slower but continued accumulation of 0.15°Brix per day requiring closer monitoring to prevent over-ripeness beyond target specifications. Zinfandel exhibited poor sugar accumulation plateau quality with continued rapid accumulation of 0.35°Brix per day combined with characteristic uneven ripening creating clusters with berries ranging from 22 to 30°Brix simultaneously, challenging harvest timing decisions and frequently resulting in high-alcohol wines when optimal flavor development required extended hangtime that pushed sugar beyond desired levels, establishing Zinfandel's reputation for alcohol management challenges rooted in its sugar accumulation plateau quality characteristics.
Sugar accumulation plateau quality is a post optimal decline quality that inheres in a grapevine cultivar or clone within a specific growing environment and represents the pattern and stability of sugar concentration changes following the achievement of initial target ripeness, specifically whether sugar accumulation continues linearly, slows to a plateau phase, or reverses through respiratory consumption and dilution effects, determining the degree to which extended hangtime shifts fruit beyond desired Brix specifications and the management interventions required to prevent unwanted over-ripeness in situations where harvest delays occur. This quality directly influences the tolerance for harvest timing variation and the feasibility of late-harvest style production, with plateau-exhibiting cultivars maintaining stable sugar levels permitting flexible scheduling while those with continued rapid accumulation quickly exceed target specifications creating harvest urgency. The ontological significance of sugar accumulation plateau quality lies in its role determining the sugar-based constraints on harvest timing, wherein cultivars reaching stable plateaus decouple sugar concentration from harvest date during the post-optimal period enabling timing decisions based on other quality parameters, while those without plateaus create continuously moving targets that may exceed acceptable ranges before optimal flavor, phenolic, or aromatic development is achieved. The biological basis of sugar accumulation plateau quality encompasses the balance between continued phloem sugar import and respiratory sugar consumption in post-véraison berries, the dilution effects of continued water import that may partially offset sugar accumulation, and the dehydration effects of late-season water deficit that concentrate sugars independent of metabolic sugar accumulation. The rate of sugar accumulation during the plateau phase typically ranges from near-zero for stable cultivars to 0.3-0.4°Brix per day for those with continued accumulation under warm conditions, creating substantial differences in the sugar trajectory following optimal ripeness. Cultivar characteristics influencing plateau quality include the developmental cessation of phloem unloading as berries approach physiological maturity, the respiratory rate of mature berries consuming accumulated sugars, and the sensitivity to dehydration that can override metabolic plateau through concentration effects.
Sugar_Accumulation_Plateau_Quality is_a Post_Optimal_Decline_Quality, representing the sugar concentration trajectory following target ripeness achievement. This class addresses whether and at what level sugar stabilizes, critical for harvest timing flexibility and alcohol management. The daily accumulation rate provides quantitative assessment metric with 0.3°Brix/day indicating continued rapid accumulation. Distinguished from Over_Ripening_Tolerance_Quality which addresses overall quality maintenance rather than specifically sugar dynamics. Critical for harvest planning and style targeting where potential alcohol influences production decisions.
Brix stabilization
Brix-Stabilisierung
Mostgewichts-Plateau
Zuckerakkumulations-Plateau
Zuckerkonzentrations-Stabilität
post-maturity sugar trajectory
sugar concentration stability
sugar plateau behavior
Keller M et al. (2006) Sugar accumulation in grapevines subjected to differential root-zone heating and cooling treatments. Vitis 45:117-125 Coombe BG (1992) Research on development and ripening of the grape berry. American Journal of Enology and Viticulture 43:101-110 Bindon KA et al. (2014) Relationships between harvest time and wine composition in Vitis vinifera L. cv. Cabernet Sauvignon. Food Chemistry 154:43-54
Sugar Accumulation Plateau Quality
In Pinot Gris berries in Alsace, France during BBCH 87-89, sugar acid ratio optimization progresses from 1.8 at early ripening through 2.9 at mid-ripening to optimal harvest ratio of 3.4 when Brix reaches 23.5 and titratable acidity declines to 6.9 g/L, with ratio increase of 0.04 per day during final two weeks enabling precise harvest timing for balanced dry wine production under mean temperatures of 20°C.
Sugar acid ratio optimization describes the viticultural objective of achieving optimal balance between sugar accumulation and acid retention in Vitis vinifera berries at harvest, wherein the ratio of soluble solids measured as Brix to titratable acidity expressed as g/L tartaric acid equivalents reaches values of 2.5-4.0 considered ideal for quality wine production, reflecting the coordinated progression of sugar loading and acid reduction during the final ripening phase. This process integrates the opposing trajectories of sugar accumulation increasing from 15-18 Brix at veraison to 22-28 Brix at harvest concurrent with titratable acidity decreasing from 15-25 g/L to 5-9 g/L, with the sugar-acid ratio increasing from initial values of 0.7-1.2 to harvest values of 2.5-4.5 over the 35-50 day post-veraison ripening period. The optimal sugar-acid ratio varies with intended wine style, ranging from lower ratios of 2.0-2.8 preferred for sparkling wine base material and crisp white wines, through intermediate ratios of 2.8-3.5 typical for balanced dry table wines, to higher ratios of 3.5-4.5 acceptable for full-bodied red wines and warm-climate productions where lower acidity is compensated by increased phenolic structure. Temperature profoundly influences sugar-acid ratio progression through differential effects on the component processes, with warm conditions accelerating both sugar accumulation and acid reduction but affecting acid loss proportionally more, producing rapid ratio increase and potential overshooting of optimal balance if harvest timing is delayed. Cultivar characteristics determine the achievable range of sugar-acid ratios at comparable maturity levels, with high-acid varieties such as Riesling capable of maintaining ratios below 3.0 at full phenolic ripeness compared to low-acid varieties such as Grenache where ratios may exceed 4.0 before optimal flavor development is achieved. Viticultural management of sugar-acid ratio optimization involves monitoring progression of both components through regular sampling, with predictive modeling using temperature forecasts enabling estimation of optimal harvest windows when target ratios will be achieved. The sugar-acid ratio provides a more integrative harvest criterion than either component alone, as it captures the balance determining sensory perception of wine harmony between sweetness and sourness that underlies consumer quality assessment.
Sugar_Acid_Ratio_Optimization is subclass of Final_Acid_Balance representing the integrative quality criterion. Distinguishes from individual sugar or acid measurements by emphasizing balance. Sibling to Mature_Berry_Acidity and Harvest_Acidity_Level. Key parameter is Brix-to-TA ratio of 2.5-4.0. Wine style dependent with lower ratios for sparkling and crisp wines, higher for full-bodied reds.
Brix-Aciditäts-Balance
Brix-acidity balance
Most-Balance-Optimierung
Zucker-Säure-Gleichgewicht
Zucker-Säure-Verhältnis-Optimierung
must balance optimization
sugar-acid equilibrium
sweetness-sourness ratio
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 278-285 Coombe BG (1992) Research on development and ripening of the grape berry. American Journal of Enology and Viticulture 43: 101-110 Jackson RS (2020) Wine Science Principles and Applications 5th Ed Chapter 6
Sugar Acid Ratio Optimization
Sugar-acid ratio quality targeting in Marlborough Sauvignon Blanc employs ratio values of 22-28 to achieve the signature style balancing tropical fruit intensity with vibrant acidity, with harvest decisions triggering when ratios reach 24-26 at 21-23°Brix and 8-10 g/L titratable acidity ratios below 22 produce excessively tart wines lacking fruit expression, while ratios above 30 yield flabby wines without the refreshing acidity that defines regional typicity, demonstrating how sugar-acid ratio provides more useful harvest guidance than either sugar or acid concentration alone.
Sugar_Acid_Ratio_Quality represents the chemical quality characterizing the proportional relationship between sugar concentration and acid concentration in grape berry juice, providing a composite index of taste balance that integrates the two primary determinants of grape flavor perception. The sugar-acid ratio (also expressed as Brix-to-acid ratio or sugar-to-titratable acidity ratio) captures the sensory balance between sweetness and tartness that fundamentally determines whether grapes taste pleasant, underripe, or overripe. This ratio increases during ripening as sugars accumulate while acids decline, providing a maturity trajectory that guides harvest timing decisions. Typical values at harvest range from 20-35 depending on cultivar, climate, and intended wine style, with lower ratios indicating tart, fresh character and higher ratios indicating ripe, softer profiles. The ratio is calculated by dividing sugar concentration (typically as °Brix) by titratable acidity (typically as g/L tartaric acid equivalents), yielding a dimensionless index that facilitates comparison across samples and vintages. Different wine styles require different optimal ratios: sparkling wine base targets lower ratios (approximately 20-25) for fresh acidity, while rich red wines may target higher ratios (30-40) for riper expression. The sugar-acid ratio responds to environmental conditions, with warm climates producing higher ratios through both elevated sugar and reduced acid compared to cool climates. Cultivar-specific differences in acid retention affect optimal ratio targets, with naturally high-acid varieties like Riesling requiring higher ratios to achieve balance than lower-acid varieties like Viognier. Assessment employs the ratio of refractometer Brix reading to titrated acidity, providing rapid field or laboratory evaluation of maturity progression.
Sugar_Acid_Ratio_Quality is positioned within Sugar_Content_Quality because the ratio is primarily used to contextualize sugar accumulation relative to acid status, though it could equally be positioned under Acid_Content_Quality. The ratio represents a derived quality calculated from two primary measurements rather than a directly measured property. Various formulations exist, including Brix/TA, sugar(g/L)/TA, and must weight/TA, requiring specification of the calculation method for reproducibility. The ratio complements but does not replace individual sugar and acid measurements, as the same ratio can result from different absolute concentrations with different sensory implications.
Brix-Säure-Quotient
Brix-acid ratio
Süße-Säure-Balance
sugar-acidity index
sweetness-tartness balance
Coombe BG et al. (1980) Development of the Grape Berry. III. Compositional Changes during Veraison Measured by Sequential Hypodermic Sampling. Australian Journal of Grape and Wine Research 1(1): 17-25 Boulton RB et al. (1999) Principles and Practices of Winemaking, Springer pp.: 65-78
Sugar Acid Ratio Quality
Sugar content data for Pinot Noir from Burgundy (Premier Cru, 2023 vintage) records simultaneous measurements of 12.8° potential alcohol, 22.5° Brix, 12.0° Baumé, and approximately 96° Oechsle, meeting the minimum natural sugar requirement of 11.0% potential alcohol for Appellation Contrôlée Bourgogne Premier Cru classification without need for chaptalization. Sugar content data for Furmint from Tokaj (2024 vintage) documents must weight of 22° KMW (equivalent to approximately 106° Oechsle) for the base dry wine lot, with botrytized Aszú berries separately measured at 45-60° KMW (220-310° Oechsle), the extreme sugar concentration reflecting the dehydration and metabolic transformation by Botrytis cinerea noble rot.
Sugar content data is a must analysis data entity that records the concentration of fermentable sugars (predominantly glucose and fructose, with minor sucrose) in grape must, measured through one or more of the internationally recognized density-based, refractometric, or hydrometric scales (Brix, Baumé, Oechsle, KMW) and additionally expressed as potential alcohol yield, providing the fundamental metric for assessing ripeness level, predicting wine alcohol content, determining regulatory quality category eligibility, and establishing the economic value of harvested grapes in many pricing systems.
Sugar content data is the single most measured parameter in viticulture worldwide, serving simultaneously as a ripeness indicator, regulatory compliance metric, pricing basis, and fermentation planning tool. The existence of multiple parallel measurement scales reflects the historical development of regional analytical traditions — Brix (soluble solids percentage by weight, dominant in the Americas and Australasia), Baumé (density-based, traditional in France and Australia), Oechsle (must density in grams above water per liter, standard in Germany and Switzerland), and KMW/Klosterneuburger Mostwaage (sugar percentage by weight, standard in Austria and parts of Central Europe). While inter-scale conversion tables enable cross-reference, the conversions are not perfectly linear because non-sugar solutes (acids, minerals, phenolics) contribute to must density, meaning that must composition variation introduces systematic conversion errors of 0.2-0.5° depending on cultivar and vintage. The relationship between sugar content and potential alcohol follows the approximate factor of 16.5-17.5 g sugar per 1% vol. alcohol produced during fermentation, though the exact conversion depends on yeast strain, fermentation temperature, and residual sugar at dryness. Modern regulatory frameworks define quality categories partly through minimum sugar content thresholds — for example, German Prädikatswein categories (Kabinett through Trockenbeerenauslese) are defined entirely by must weight at harvest, making sugar content measurement a legally consequential analytical determination requiring certified instruments and documented methodology.
Zuckergehaltsdaten
Zuckerkonzentrationsdaten
Zuckermesswerte
must sugar content
sugar concentration data
sugar measurement data
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1, 2nd Ed, Wiley, pp.: 219-230 Jackson RS (2020) Wine Science: Principles and Applications, 5th Ed, Academic Press, pp.: 330-345 OIV (2021) Compendium of International Methods of Analysis of Wines and Musts, Method OIV-MA-AS2-02
Sugar Content Data
Sugar content quality monitoring in Napa Valley Cabernet Sauvignon employs twice-weekly sampling from véraison through harvest, tracking the accumulation curve from 5-8°Brix at véraison through 22-26°Brix at optimal harvest maturity the rate of sugar accumulation (typically 0.3-0.5°Brix per day during active ripening) guides irrigation decisions, with water application modulated to maintain steady accumulation without the sugar concentration spikes associated with berry dehydration or the stalled ripening caused by excessive water stress.
Sugar_Content_Quality encompasses all chemical qualities that characterize the carbohydrate composition of grapevine tissues, with primary emphasis on the soluble sugars that accumulate in grape berries during ripening. Sugar content is the paramount chemical quality for both wine and table grape production because sugars provide the substrate for alcoholic fermentation, contribute the sweetness sensation in fresh and dried fruit, and serve as the primary energy reserve transported from leaves to developing berries. The category addresses Total_Sugar_Quality capturing the aggregate sugar content, Reducing_Sugar_Quality encompassing the glucose and fructose that dominate ripe berry composition, and Non_Reducing_Sugar_Quality addressing sucrose and other non-reducing sugars. Sugar accumulation in grape berries represents one of the most dramatic compositional changes in any fruit crop, with concentrations increasing from near zero at fruit set to 150-300 g/L at harvest, representing 15-30% of berry fresh weight. This massive accumulation results from phloem unloading of sucrose transported from photosynthesizing leaves, followed by invertase-mediated cleavage to glucose and fructose that are compartmentalized in berry vacuoles. Sugar concentration is routinely measured as the primary harvest maturity indicator using refractometry (°Brix), density measurement (Baumé, Oechsle, KMW), or specific enzymatic assays for individual sugars. The relationship between sugar concentration and wine alcohol potential follows the approximate conversion factor of 16-18 g/L sugar per 1% volume alcohol, enabling harvest timing to target desired wine styles from light (10% alcohol) through full-bodied (14%+). Environmental factors profoundly influence sugar accumulation rate and final concentration, with warm temperatures, moderate water deficit, and high light exposure promoting rapid accumulation, while excessive heat can arrest ripening before optimal sugar levels are achieved.
Sugar_Content_Quality serves as the parent class for specific sugar categories and is positioned as a direct subclass of Chemical_Quality. Sugar content provides the basis for traditional must weight scales including Brix (grams sucrose per 100 g solution), Baumé (approximating potential alcohol), Oechsle (density in grams per liter minus 1000), and KMW (Klosterneuburger Mostwaage, Austrian scale). The relationship between sugar content and perceived sweetness involves interaction with acid content the sugar-acid balance rather than sugar alone determines sensory perception. Sugar qualities should be distinguished from sugar-related dispositions (e.g., capacity for sugar accumulation) and processes (e.g., sugar transport, sugar metabolism).
Kohlenhydratgehalt
Süßepotential
Zuckergehalt
Zuckerzusammensetzung
carbohydrate content
saccharide quality
sugar composition
sweetness potential
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 267-295 Coombe BG and McCarthy MG (2000) Dynamics of Grape Berry Growth and Physiology of Ripening. Australian Journal of Grape and Wine Research 6(2): 131-135
Sugar Content Quality
Sugar cryoprotection quality assessment in Riesling during autumn acclimation characterized the accumulation dynamics of specific sugar classes and their relationship to developing freezing tolerance. Weekly sampling of bud and cane tissues from September through December with HPLC sugar analysis and parallel freezing tolerance testing established the sugar-hardiness relationship. Total soluble sugar concentration increased from 52 mg/g dry weight in September (LT50 = -8°C) to 195 mg/g in December (LT50 = -26°C), with the 3.8-fold increase strongly correlated with the 18°C improvement in freezing tolerance (r² = 0.94). Raffinose family oligosaccharide (RFO) accumulation showed the strongest relationship to hardiness, increasing from 4 mg/g to 72 mg/g (18-fold) with raffinose and stachyose together explaining 87% of hardiness variation in multiple regression analysis, consistent with their known efficacy for membrane stabilization. Sucrose showed 3-fold increase from 28 to 85 mg/g contributing to both osmotic and stabilization functions. The sugar cryoprotection quality profile differed between tissues, with buds accumulating 25% higher total sugars than adjacent cane tissues, consistent with the greater cold sensitivity of buds requiring enhanced cryoprotection for equivalent survival. The sugar cryoprotection quality assessment established target sugar concentrations of greater than 150 mg/g total sugars and greater than 50 mg/g RFOs for adequate winter hardiness, enabling sugar analysis as a molecular marker for acclimation status.
Sugar cryoprotection quality is a cryoprotection quality that inheres in a grapevine organism and represents the capacity to accumulate soluble sugars during cold acclimation that serve multiple cryoprotective functions including osmotic adjustment reducing cellular dehydration during extracellular freezing, membrane stabilization through direct interaction with lipid bilayers substituting for water molecules in maintaining membrane structure, and vitrification promotion creating a glassy cytoplasmic state that prevents damaging ice crystallization in concentrated cellular solutions. This quality addresses the central role of sugars as the primary cryoprotective metabolites in grapevine cold hardiness, recognizing that soluble sugars constitute the largest fraction of accumulated cryoprotectants during acclimation and provide protection through multiple complementary mechanisms that together substantially enhance freezing tolerance. The ontological significance of sugar cryoprotection quality lies in the quantitative importance of sugar accumulation for cold hardiness, with the concentration of protective sugars showing strong correlation with freezing tolerance across genotypes and acclimation states, and with manipulation of sugar levels through management or genetics offering practical approaches to modifying cold hardiness. The biological basis of sugar cryoprotection encompasses the metabolic pathways of sugar synthesis and interconversion that produce the specific sugars accumulating during acclimation, the transport processes that concentrate sugars in cold-vulnerable tissues, and the biophysical interactions between sugars and cellular structures that provide protection during freezing stress. The specific sugars contributing to cryoprotection vary in their protective efficacy, with the raffinose family oligosaccharides (raffinose, stachyose, verbascose) showing particular effectiveness for membrane stabilization, sucrose providing both osmotic and stabilization functions, and glucose and fructose contributing primarily to osmotic adjustment. The relationship between total sugar concentration and freezing tolerance shows characteristic patterns with tolerance improving as sugar accumulates up to a plateau reflecting saturation of protective mechanisms, above which additional sugar accumulation provides diminishing benefit, establishing the target sugar levels that acclimation must achieve for adequate winter protection.
Sugar_Cryoprotection_Quality is_a Cryoprotection_Quality, representing the accumulation of soluble sugars providing osmotic, membrane-stabilizing, and vitrification-promoting cryoprotection. This class addresses the quantitatively dominant cryoprotectant class. RFOs (raffinose family oligosaccharides) show particular efficacy for membrane stabilization. The 3-4 fold increase during acclimation and strong correlation with hardiness (r² > 0.9) enables sugar levels as molecular markers. Tissue-specific accumulation reflects differential protection requirements. Target concentrations provide management and breeding objectives.
Kohlenhydrat-Kälteschutz
Zucker-Kryoprotektion
carbohydrate cryoprotection
lösliche Zucker-Frostschutz
osmotic cryoprotection
osmotische Kryoprotektion
soluble sugar cold protection
sugar accumulation for cold protection
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Grant TN et al. (2009) Characterization of cryoprotective compounds in cold acclimated and non-acclimated Riesling grapevines. Acta Horticulturae 833:85-90 Xin Z, Browse J (2000) Cold comfort farm: the acclimation of plants to freezing temperatures. Plant, Cell and Environment 23:893-902
Sugar Cryoprotection Quality
In a Marlborough Sauvignon Blanc vineyard, sugar equilibrium was characterized over 16 days in late March-April Year N+1 (BBCH 89) using isotope tracer studies when ¹³C-glucose uptake 26 μmol g⁻¹ h⁻¹ measured but vacuolar concentration stable 1880 mM indicating equal efflux or respiration balancing import, berry Brix constant 22.6° ± 0.3° over measurement period, respiratory consumption calculated 23 μmol g⁻¹ h⁻¹ from O₂ uptake, net vacuolar accumulation <1.5 μmol g⁻¹ h⁻¹ confirming equilibrium, under cool New Zealand autumn conditions (daily high 16-19°C maintaining stable metabolism).
Sugar_Equilibrium represents the dynamic steady-state condition at harvest maturity when hexose import flux precisely balances respiratory consumption, metabolic interconversion, and potential export creating zero net vacuolar accumulation despite continued molecular exchange defining the biochemical and thermodynamic endpoint of ripening sugar accumulation regardless of actual harvest timing. This equilibrium state occurs during plateau period (BBCH 88-89), typically days 58-80 post-veraison though highly variable, and is characterized by vacuolar hexose concentrations constant 1900-2600 mM combined glucose and fructose (cultivar maximum) with fluctuations <±2-5% representing measurement variability and true equilibrium oscillation, plasma membrane hexose import 18-35 μmol g⁻¹ h⁻¹ continuing to supply cellular metabolism, respiratory hexose consumption 15-30 μmol g⁻¹ h⁻¹ utilizing imported plus vacuolar reserves maintaining energy production, and net change in vacuolar content dC/dt = (Import - Respiration - Export - Interconversion) ≈ 0 ± 1-3 μmol g⁻¹ h⁻¹ within precision of measurement techniques. Thermodynamic equilibrium characteristics include electrochemical potential for hexose transport μ_hexose = μ°_hexose + RT ln([hexose]) equal across plasma membrane (apoplast ⇌ cytoplasm) and tonoplast (cytoplasm ⇌ vacuole) within energy available from coupled ion gradients, free energy change ΔG for hexose transport approaching zero ΔG = ΔG° + RT ln(C_product/C_substrate) + zFΔψ where electrochemical driving force balanced by concentration opposition, chemical potential gradient balanced by electrical potential gradient creating equilibrium distribution determined by coupling stoichiometry with H⁺ symport, and osmotic equilibrium where vacuolar osmotic potential -3.0 to -4.0 MPa balanced by turgor pressure 0.2-0.4 MPa plus apoplastic water potential creating stable cell volume without net water movement. Kinetic equilibrium properties demonstrate forward hexose transport rate (apoplast→cytoplasm→vacuole) approximately equal to reverse processes (vacuolar efflux, export, interconversion) creating bidirectional flux with zero net accumulation, transporter-mediated uptake Vforward = Vmax_forward × [hexose]_apoplast / (Km + [hexose]_apoplast) balanced by facilitated diffusion back-flux Vreverse = Vmax_reverse × [hexose]_vacuole / (Km + [hexose]_vacuole) where concentration-dependent rates equilibrate, and metabolic sink strength (respiration, biosynthesis) determining equilibrium concentration by consuming imported hexose maintaining gradient enabling continued import despite vacuolar saturation. Molecular basis of equilibrium includes transporter protein levels stabilized with synthesis matching degradation maintaining constant density 6-10% plasma membrane (from peak 10-16%), gene expression steady-state with transcription rate balancing mRNA decay yielding constant transcript levels 55-85 fold above baseline (from peak 100-150 fold), regulatory feedback where high vacuolar hexose potentially suppressing further transporter induction through hexokinase-independent sensors creating negative feedback maintaining equilibrium, and hormone signaling with ABA declining from ripening peak 2000-3200 ng/g to equilibrium 1200-2000 ng/g reducing ripening program activation while maintaining basal transporter expression. Metabolic flux at equilibrium shows hexose import 18-35 μmol g⁻¹ h⁻¹ partitioned among respiratory consumption 15-30 μmol g⁻¹ h⁻¹ (50-85% of import), biosynthetic uses 3-8 μmol g⁻¹ h⁻¹ (8-20%) for flavor compounds, cell wall precursors, secondary metabolites, and potential export or interconversion 1-5 μmol g⁻¹ h⁻¹ (3-12%) with net vacuolar accumulation <1-3 μmol g⁻¹ h⁻¹ effectively zero, isotope tracer studies showing labeled hexose rapidly entering vacuole but equal unlabeled efflux creating dynamic exchange with constant total concentration, and respiratory quotient 1.05-1.25 indicating mixed substrate oxidation with hexoses contributing 50-75% and remaining acids 25-50% as malate depletes. Compartmental analysis reveals apoplastic hexose 180-350 mM (glucose 90-180 mM, fructose 90-170 mM) maintained by invertase activity balanced by cellular uptake, cytoplasmic hexose 70-120 mM steady through balance of plasma membrane import and tonoplast export plus metabolism, vacuolar hexose 1900-2600 mM (950-1300 mM each glucose and fructose approximately equimolar) representing storage pool at equilibrium maximum, and concentration ratios vacuole:cytoplasm:apoplast approximately 18:1:3 typical at equilibrium reflecting transporter distributions and energetics. Environmental perturbations to equilibrium include temperature changes causing transient disequilibrium with warmer temperatures increasing respiration potentially depleting vacuolar reserves slightly while cooler reducing metabolism allowing minor accumulation, water stress affecting cell volume and thus concentration with dehydration concentrating vacuolar hexose producing apparent Brix increase without net accumulation, substrate availability changes from declining source photosynthesis potentially reducing apoplastic hexose lowering equilibrium slightly as import cannot be maintained, and disease or stress disrupting normal metabolism potentially causing degradation exceeding import creating net loss and equilibrium decline. Recovery from perturbation demonstrates resilience with system returning to equilibrium within 2-5 days following transient temperature or water change, regulatory mechanisms adjusting import and respiration rates compensating for disturbances, and robust homeostasis maintaining constant vacuolar concentration despite environmental variability within normal range. Cultivar differences in equilibrium characteristics show high-Brix varieties achieving equilibrium at 2200-2600 mM vacuolar hexose corresponding to 26-30° Brix reflecting genetic maximum capacity, moderate cultivars equilibrating at 1800-2200 mM and 22-26° Brix, and lower types at 1600-2000 mM and 20-24° Brix, with equilibrium concentration likely determined by maximum transporter expression capacity, vacuolar properties (size, membrane area, pH regulation), and metabolic set-points governing downregulation triggers. Temporal stability shows equilibrium maintained for extended periods 2-6 weeks or longer if harvest delayed with berries remaining at constant composition, occasional slow drift toward slightly lower Brix 0.3-0.8° over 3-4 weeks potentially indicating senescence processes beginning to dominate over minimal import, and remarkable stability despite environmental fluctuation demonstrating robust homeostatic regulation maintaining maturity composition. Quality implications include equilibrium Brix defining maximum achievable alcohol potential for given cultivar and conditions, equilibrium representing physiological maturity ceiling where extended hang time produces minimal sugar change but may improve phenolics and flavor particularly reds, and equilibrium stability providing flexible harvest window 2-4 weeks where sugar composition constant enabling optimization of other parameters. Harvest timing at equilibrium recognizes sugar composition stable providing flexibility, other maturity indicators (acid, phenolics, flavor, seed color) potentially continuing to evolve during equilibrium requiring integrated assessment, weather and disease risks increasing with extended hang time requiring balance between quality optimization and crop protection, and economic factors (labor, winery capacity) often determining actual harvest timing within biologically suitable window. Viticultural management during equilibrium includes minimal intervention as composition stable, disease control essential preventing infections disrupting equilibrium, irrigation ceased allowing natural water balance potentially slight concentration, and monitoring other maturity parameters guiding final harvest decision beyond sugar alone. Research applications include thermodynamic studies characterizing equilibrium conditions and free energy changes, flux analysis using isotopes quantifying bidirectional exchange rates, regulatory mechanism investigations revealing signals maintaining homeostasis, and modeling approaches describing equilibrium as emergent property of integrated transport, metabolism, and regulatory networks enabling prediction and optimization.
Sugar_Equilibrium is distinguished from Sugar_Plateau by emphasizing dynamic steady-state (import = consumption + export) rather than static maximum, from Maximum_Sugar_Content by focusing on equilibrium process rather than achieved level, and from accumulation by representing zero net change (dC/dt ≈ 0) rather than positive flux. Vacuolar hexose 1900-2600 mM cultivar maximum. Import 18-35 μmol g⁻¹ h⁻¹ balanced by consumption. Thermodynamic and kinetic equilibrium. Duration 2-6+ weeks potentially. Sugar_Equilibrium represents dynamic steady-state defining biochemical maturity endpoint.
Akkumulations-Konsum-Balance
Hexose-Gleichgewichtszustand
Zucker-Homöostase
accumulation-consumption balance
dynamic equilibrium
dynamisches Gleichgewicht
hexose steady-state
sugar homeostasis
Keller M (2020) The Science of Grapevines 3rd Ed pages 227-228 equilibrium state Fondy BR, Geiger DR (1982) Diurnal pattern of translocation and carbohydrate metabolism in source leaves of Beta vulgaris L. Plant Physiol 70:671-676 equilibrium concepts applicable to grape Walker RP, Famiani F (2018) Organic acids in fruits:metabolism, functions and contents Hort Rev 45:371-430 metabolic equilibrium in fruits.
Sugar Equilibrium
In a Bordeaux Sémillon vineyard, sugar import enhancement was tracked over 7 days in mid-September Year N+1 (BBCH 84) when VvHT1 expression reached 95-fold (near maximum), glucose uptake rates were 58 μmol g⁻¹ h⁻¹, vacuolar hexose rose from 720 mM to 1150 mM, berry Brix increased from 16.8° to 23.2° (average 3.3% per week), fresh weight expanded 18% to 1.68 g, titratable acidity declined to 7.2 g/L, under warm late-summer conditions (daily high 26-29°C, moderate deficit stem ψ -1.1 MPa concentrating sugars).
Sugar_Import_Enhancement represents the progressive amplification of hexose uptake and accumulation capacity during early-to-mid veraison when transporter expression, enzyme activities, and cellular storage capacity coordinate to create increasingly efficient sugar import enabling the transition from moderate to rapid accumulation rates that achieve final berry sweetness. This enhancement occurs during early-to-mid veraison (BBCH 83-84), typically 3-8 days post-transport initiation, and is characterized by hexose transporter gene expression reaching 75-100% of maximum levels (VvHT1 approaching 100-fold above green berry, VvHT2 80-fold, others 60-90 fold), hexose uptake rates into mesocarp cells achieving 35-65 μmol glucose g⁻¹ h⁻¹ representing 8-15 fold enhancement over pre-veraison baseline, and sugar accumulation accelerating from initial phase 1.5-2.5% Brix per week to enhanced phase 2.5-4.0% per week enabling rapid sweetening toward harvest targets. Cellular mechanisms underlying enhancement inclu continued transporter protein synthesis and membrane insertion increasing transporter density 3-5 fold at plasma membrane and tonoplast, vacuolar expansion to occupy 92-95% of mesocarp cell volume (from 85-90% at initiation) creating enlarged storage capacity estimated 0.8-2.5 nL per cell, enhanced membrane surface area through continued vesicle fusion and membrane proliferation potentially doubling tonoplast area increasing transport sites, and optimized metabolic coordination where cytoplasmic hexose metabolism (glycolysis, PPP) adjusted to maintain low cytoplasmic concentrations (50-100 mM) sustaining concentration gradient favoring continued import despite rising vacuolar levels. Gene expression dynamics during enhancement show: hexose transporter genes (VvHT1, VvHT2, VvHT3, VvHT5, VvHT6) reaching maximum steady-state expression with transcripts 80-120 fold above green berry baseline, cell wall invertase maintaining high expression (60-100 fold) ensuring continued sucrose cleavage in apoplast, vacuolar invertase achieving 40-80 fold elevation contributing to intracellular sucrose metabolism, regulatory transcription factors (sugar-responsive VvMYB, VvbZIP, VvNAC) sustained at high levels coordinating transporter, enzyme, and metabolic gene expression, and feedback regulation beginning where very high hexose levels (>800 mM vacuolar) may start reducing transporter expression in some berries approaching saturation. Transport kinetics during enhancement demonstrate: glucose uptake Vmax increasing to 55-95 μmol g⁻¹ h⁻¹ (from 25-50 μmol at initiation) through increased transporter density with Km remaining 1.5-3.0 mM, fructose transport Vmax reaching 45-85 μmol g⁻¹ h⁻¹ with Km 5-12 mM, competition between hexoses diminishing as transporter capacity exceeds supply creating non-saturating conditions, and tonoplast transport rates 40-75 μmol hexose g⁻¹ h⁻¹ efficiently moving cytoplasmic hexoses into vacuoles preventing accumulation that could inhibit plasma membrane import. Vacuolar accumulation during enhancement shows: hexose concentrations rising from 400-700 mM at initiation to 800-1200 mM during enhancement representing berry Brix 15-22°, glucose:fructose ratio approaching 0.95-1.05 (essentially equimolar) through balanced transport and differential metabolism normalizing any initial imbalances, osmotic potential declining from -1.2 to -1.8 MPa to -1.8 to -2.6 MPa creating strong water potential gradient driving continued water import and berry expansion 15-25% fresh weight increase during enhancement period, and vacuolar pH maintained 3.0-3.5 through H⁺-ATPase and H⁺-PPase activity creating acidic environment important for anthocyanin color stability (in reds) and potentially affecting some metabolic processes. Hormonal regulation during enhancement includes: ABA levels sustained 1200-2200 ng/g maintaining ripening program activation and transporter gene expression, hexose itself at high concentrations providing strong signaling through hexokinase pathway (glucose-6-phosphate levels 15-35 μmol/g FW) amplifying sugar-responsive gene expression through positive feedback, declining gibberellin and auxin maintaining low levels removing any residual growth-related suppression, and ethylene showing cultivar-dependent patterns with slight elevation 0.3-1.0 nL g⁻¹ h⁻¹ in some varieties potentially fine-tuning ripening progression. Environmental influences on enhancement inclu temperature strongly affecting enhancement rate with optimal 25-30°C supporting maximum transporter synthesis and transport kinetics achieving 3.5-4.0% Brix per week, cooler conditions 20-25°C reducing to 2.5-3.2% Brix per week through slower metabolism, heat stress >32°C impairing enhancement potentially reducing to 1.1-1.3 indicating substrate-level metabolism dominates with some fermentative activity, carbon partitioning strongly favoring storage (75-85% of imported carbon accumulating as hexoses) over structural or metabolic uses, nitrogen metabolism adjusted with amino acid levels declining further (40-60% reduction from green berry) as incorporation into proteins and secondary metabolites proceeds, and potassium accumulation parallel with sugars (berry K⁺ rising from 1500-2000 to 2000-2800 mg/kg) potentially important for pH regulation and membrane function. Quality implications of enhancement inclu enhancement rate and duration critically determine final sugar levels achieved with faster longer enhancement producing higher Brix enabling earlier harvest or lower-alcohol wine styles through earlier picking, uniformity of enhancement across berries and clusters affecting harvest maturity variation with heterogeneous enhancement requiring selective harvest or acceptance of compositional variation, and timing of enhancement peak relative to flavor and phenolic development determining optimal harvest window with ideal scenario showing coordinated maturation of all parameters. Viticultural management during enhancement involves: irrigation typically terminated or minimized as berry water demand declines and concentration desirable, canopy management maintaining healthy source leaves supporting continued photosynthesis and sugar export, crop load ensuring adequate leaf area per fruit weight (typically 12-18 cm² per g at this stage) enabling enhancement without source limitation, disease monitoring intensifying as increasing sugar attracting insects and pathogens, and harvest forecasting using enhancement rates to project timing when target Brix achieved enabling logistics planning.
Sugar_Import_Enhancement is distinguished from Hexose_Transport_Initiation by representing amplification rather than onset, from Initial_Sugar_Increase by showing accelerating (2.5-4.0% Brix/week) rather than moderate (1.5-2.5%/week) rates, and from peak accumulation by being progressive rather than maximum. Transporter expression 75-100% maximum. Uptake rates 8-15 fold over baseline. Duration typically 5-8 days BBCH 83-84. Temperature optimal 25-30°C. Sugar_Import_Enhancement represents critical escalation phase driving berry toward harvest sweetness levels.
accelerating sugar loading
amplified sugar uptake
beschleunigende Zucker-Beladung
enhanced accumulation capacity
erhöhte Akkumulations-Kapazität
progressive Hexose-Import-Zunahme
progressive hexose import increase
verstärkte Zucker-Aufnahme
Keller M (2020) The Science of Grapevines 3rd Ed pages 221-225 sugar accumulation enhancement Dai ZW, Ollat N, Gomès E, Decroocq S, Tandonnet JP, Bordenave L, Pieri P, Hilbert G, Kappel C, van Leeuwen C, Vivin P, Delrot S (2011) Ecophysiological, genetic, and molecular causes of variation in grape berry weight and composition Plant Physiol 155:519-532 accumulation regulation Sweetman C, Deluc LG, Cramer GR, Ford CM, Soole KL (2009) Regulation of malate metabolism in grape berry and other developing fruits Phytochem 70:1329-1344 metabolic coordination.
Sugar Import Enhancement
Sugar maturity tracking in Zinfandel demonstrates the accumulation dynamics affecting harvest planning, with véraison +2 weeks at 20 Brix (accumulating approximately 0.3 Brix/day in warm conditions), véraison +4 weeks reaching 24 Brix (target for balanced table wine at approximately 14% potential alcohol), and continued accumulation to 28 Brix by véraison +6 weeks if unharvested (potential alcohol exceeding 16%, requiring stylistic adjustment toward late-harvest or fortified wine), with the rapid accumulation rate in this variety creating a narrow harvest window for table wine production that sugar maturity monitoring enables the winemaker to precisely identify.
Sugar maturity quality is a technological maturity quality that represents the accumulation of fermentable sugars in grape berries, characterizing the achievement of target sugar concentration that determines potential alcohol content in resulting wine, with sugar level serving as the primary technological maturity parameter due to its direct relationship to wine alcohol and its ease of measurement through refractometry (Brix) or densitometry. Within the BFO framework, this quality specifically addresses the sugar component of technological maturity, distinguishing sugar accumulation from acid metabolism or other compositional parameters while recognizing that sugar concentration has historically dominated harvest decisions due to its direct impact on wine alcohol, its correlation (though imperfect) with overall ripeness, and its straightforward measurement enabling frequent monitoring throughout the ripening period. The ontological significance lies in sugar concentration determining the fermentation potential of must and the alcohol content of resulting wine, where each degree Brix represents approximately 0.55-0.60% potential alcohol, such that harvest sugar level directly establishes the alcohol parameter that affects wine style, body, microbial stability, and regulatory classification. The physiological basis for sugar accumulation involves phloem transport of sucrose from photosynthesizing leaves to berry sinks, where sucrose is cleaved to glucose and fructose (approximately equal at maturity) and accumulated in vacuoles, with accumulation rate dependent on source capacity (leaf area, photosynthetic rate), sink demand (berry number, berry size), and transport efficiency (phloem function, water status). The sugar accumulation patterns during ripening show approximately linear increase from véraison through a plateau approaching maximum concentration, with the rate of accumulation affected by temperature (accelerated by warmth within limits), crop load (diluted by heavy crops), and water status (concentrated by mild deficit, potentially arrested by severe stress). The climate change implications for sugar maturity are significant, as warming temperatures accelerate sugar accumulation potentially to levels inappropriate for table wine production (>26 Brix) before other maturity parameters (phenolics, acids, flavors) reach optimal levels, creating the contemporary challenge of "alcohol creep" and renewed interest in earlier-ripening cultivars or harvest at lower sugar with potential adjustment. Understanding sugar maturity quality enables prediction of wine alcohol, harvest timing based on sugar targets appropriate for intended wine style, and recognition of the conditions affecting sugar accumulation rate that determine when targets will be achieved.
Sugar_Maturity_Quality addresses the accumulation of fermentable sugars determining wine potential alcohol. This quality is the primary technological maturity parameter due to direct wine impact and measurement ease. The class supports harvest timing and alcohol management through sugar maturity characterization.
Brix maturity
Brix-Reife
Zuckerreife
potential alcohol
sugar content
sugar ripeness
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 277-316 Coombe BG, McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6(2): 131-135
Sugar Maturity Quality
In a Napa Cabernet Sauvignon vineyard, sugar plateau was confirmed over 19 days in late September-October Year N+1 (BBCH 89) when weekly Brix measurements showed stability: 25.2°, 25.4°, 25.6°, 25.3° (mean 25.4° ± 0.2°, CV 0.8%), vacuolar hexose 2150 mM combined, hexose import estimated 24 μmol g⁻¹ h⁻¹ balanced by respiration 22 μmol g⁻¹ h⁻¹, enabling harvest timing optimization for phenolic maturity (achieved week 2) without concern for sugar loss, under warm Napa autumn conditions maintaining equilibrium without degradation.
Sugar_Plateau represents the equilibrium state during final ripening when berry hexose concentrations stabilize at cultivar-specific maximum levels with minimal or no net accumulation creating the characteristic terminal sweetness that defines harvest maturity and indicates physiological ripening completion regardless of whether other quality parameters have achieved target levels. This plateau state occurs during late-to-harvest ripening (BBCH 88-89), typically days 55-75 post-veraison though timing highly variable, and is characterized by berry Brix stabilizing within narrow range ±0.3-0.8° over 7-14 day measurement period indicating effective cessation of net accumulation, daily Brix changes <0.03-0.06% representing measurement variability rather than true accumulation, vacuolar hexose concentrations 1900-2600 mM combined glucose and fructose (cultivar-dependent maximum) showing minimal variation <3-8% over plateau period, and hexose import flux approximately balanced by respiratory consumption plus any export creating steady-state where dBrix/dt ≈ 0 though instantaneous rates may fluctuate. Physiological basis of plateau includes vacuolar solute saturation approaching maximum sustainable concentration ~2000-2600 mM beyond which further accumulation thermodynamically unfavorable or physically impossible due to solubility limits, osmotic potential stabilized at extreme values -3.0 to -4.0 MPa near cellular tolerance limits preventing further concentration increases, concentration gradients (apoplast-to-vacuole) minimized to <5-10:1 (from 15-30:1 peak accumulation) providing insufficient driving force for significant net transport, transporter expression and activity downregulated to maintenance levels 50-80 fold above green berry (from peak 100-150 fold) adequate for replacing respiratory losses but insufficient for net accumulation, and metabolic equilibrium where import rate 18-35 μmol hexose g⁻¹ h⁻¹ approximately equals respiratory consumption 15-30 μmol g⁻¹ h⁻¹ plus minor metabolic interconversions creating zero net vacuolar accumulation. Molecular mechanisms maintaining plateau include transporter gene expression stabilized at reduced steady-state with transcription rate balancing mRNA degradation, protein synthesis matching degradation maintaining constant transporter density though reduced from peak levels, lack of strong regulatory signals (ABA declining from peak 2000-3200 ng/g to plateau 1200-2000 ng/g, sugar signaling feedback potentially suppressing further induction), and cellular homeostatic mechanisms detecting high vacuolar sugar potentially triggering active suppression of transport machinery through unknown sensors and signaling pathways. Thermodynamic analysis at plateau shows free energy change for hexose transport ΔG ≈ 0 or slightly positive indicating thermodynamic equilibrium where forward transport (apoplast→vacuole) energetically unfavorable without coupling to ATP hydrolysis, maximum achievable gradient limited by H⁺-ATPase capacity with realistic coupling stoichiometry (1-2 H⁺ per hexose transported) and ATP availability constraining gradient to ~500-1500:1 theoretical maximum but practical plateau gradients only 5-10:1 suggesting additional kinetic or regulatory limitations beyond pure thermodynamics, and osmotic work calculation showing energy required for further accumulation W = RT × Δ(ln C) exceeding available cellular energy budget from concurrent metabolism. Metabolic steady-state characteristics include hexose import continuing at low rate 18-35 μmol g⁻¹ h⁻¹ maintaining substrate for respiration, respiratory consumption 15-30 μmol g⁻¹ h⁻¹ utilizing both imported hexoses and vacuolar reserves with dynamic exchange, potential minor export or interconversion 3-8 μmol g⁻¹ h⁻¹ for biosynthesis (flavor compounds, cell wall precursors, secondary metabolites), and net accumulation <2-5 μmol g⁻¹ h⁻¹ effectively zero within measurement precision representing true equilibrium rather than kinetic slowdown. Environmental effects on plateau include temperature affecting absolute concentration achieved with warm conditions producing plateau at higher Brix 24-28° versus cool 20-24° through effects on accumulation kinetics during approach, water availability influencing concentration versus dilution with deficit potentially enhancing final Brix through reduced berry expansion while adequate water maintaining size with slightly lower concentration, source photosynthesis decline with canopy senescence potentially lowering plateau level if substrate depletion forces premature cessation before thermodynamic maximum achieved, and seasonal progression with late harvest potentially showing slight Brix decline 0.3-0.8° as berry water balance changes or degradation processes exceed minimal import. Cultivar genetic variation demonstrates high-Brix varieties achieving plateau 26-30° Brix corresponding to vacuolar hexose 2200-2600 mM near apparent saturation limits, moderate cultivars plateauing 22-26° Brix with 1800-2200 mM vacuolar concentrations, and lower-Brix types stabilizing 20-24° Brix at 1600-2000 mM reflecting genetic differences in maximum transporter capacity, vacuolar properties, and regulatory set-points determining when accumulation ceases. Temporal stability during plateau shows measurements over 7-21 days typically varying <±0.4° Brix (coefficient of variation <2-3%) indicating true equilibrium, occasional slight increases 0.2-0.5° over 7-10 days potentially representing measurement variability, temperature effects, or minor true accumulation, and rare decreases suggesting berry dehydration, degradation, or disease affecting measurement accuracy. Quality implications of plateau include maximum achievable sugar determining wine alcohol potential with cultivar and conditions setting ceiling independent of extended hang time, plateau achievement indicating physiological maturity useful for harvest timing though other parameters (phenolics, flavor, acid) may lag requiring integrated assessment, and plateau Brix serving as baseline for late-harvest or raisin production where further concentration requires dehydration rather than active accumulation. Harvest timing considerations at plateau recognize continued hang time beyond plateau achievement produces minimal sugar gain but may improve other quality parameters particularly phenolic and flavor maturity in reds, extended plateau period (2-4 weeks) creating flexible harvest window allowing optimization for non-sugar factors, weather risks increasing with extended hang time requiring balance between quality optimization and crop protection, and economic considerations where labor and winery capacity may necessitate harvest at plateau even if extended maturation potentially beneficial. Viticultural management during plateau includes minimal intervention as berry composition stable, disease control critical as high-sugar berries vulnerable during extended plateau, irrigation typically ceased allowing natural ripening completion and potentially slight concentration through dehydration, bird protection essential as peak sugar attractiveness maintained, and harvest logistics coordination recognizing plateau provides flexible timing within 2-4 week window. Exceptions to stable plateau include certain cultivars or conditions showing continued minimal accumulation 0.05-0.10% per day even at apparent plateau, disease or stress disrupting equilibrium causing decline, dehydration through transpiration or environmental stress concentrating sugars producing apparent Brix increase without net hexose accumulation, and measurement artifacts from temperature effects or sampling variability creating apparent fluctuation. Research applications include thermodynamic studies characterizing equilibrium conditions and limits, regulatory mechanism investigations revealing signals controlling plateau achievement, comparative analyses across genotypes identifying genetic determinants of maximum capacity, and modeling approaches predicting plateau timing and level enabling harvest optimization strategies for quality and alcohol management.
Sugar_Plateau is distinguished from Approaching_Sugar_Maximum by showing achieved equilibrium (Brix variation <±0.3-0.8° over 7-14 days) rather than continuing approach, from declining rates by exhibiting effectively zero accumulation (dBrix/dt ≈ 0) rather than deceleration, and from sugar accumulation by representing cessation rather than ongoing process. Duration variable 7-21+ days BBCH 88-89. Cultivar maximum: high-Brix 26-30°, moderate 22-26°, lower 20-24°. Thermodynamic and regulatory equilibrium. Sugar_Plateau represents terminal equilibrium state defining physiological maturity ceiling for sweetness.
Akkumulations-Beendigung
Zucker-Gleichgewicht
accumulation cessation
maximale Süße erreicht
maximum sweetness achieved
sugar equilibrium
Coombe BG (1992) Research on development and ripening of the grape berry Amer J Enol Vitic 43:101-110 plateau characteristics Keller M (2020) The Science of Grapevines 3rd Ed pages 227-228 sugar equilibrium Ollat N, Gaudillère JP (1998) The effect of limiting leaf area during stage I of berry growth on development and composition of berries of Vitis vinifera L. cv. Cabernet Sauvignon Amer J Enol Vitic 49:251-258 plateau and equilibrium.
Sugar Plateau
In a Mosel Riesling vineyard, sugar transport activation was characterized over 8 days in mid-September Year N+1 (BBCH 83-84) when VvHT1 expression increased from 8-fold to 72-fold, VvHT2 rose from 5-fold to 58-fold, glucose uptake into isolated mesocarp cells increased from 6.5 to 48 μmol g⁻¹ h⁻¹, berry Brix rose from 9.8° to 15.2° (average 2.7% per week accelerating), ABA levels 1180 ng/g, under cool German conditions (daily high 20-23°C, adequate vine water status).
Sugar_Transport_Activation represents the coordinated upregulation of membrane transport systems enabling massive hexose flux from apoplast into cells and subsequently into vacuolar storage creating the critical cellular import machinery that converts phloem-delivered sucrose products into accumulated berry sweetness. This activation occurs during veraison onset through mid-veraison (BBCH 83-84), typically 0-5 days following or concurrent with phloem unloading activation, and is characterized by hexose transporter gene expression increasing 30-100 fold collectively (VvHT1, VvHT2, VvHT3, VvHT5, VvHT6 showing individual 25-150 fold inductions), plasma membrane hexose uptake capacity rising from pre-veraison 3-10 μmol glucose g⁻¹ h⁻¹ to activation phase 25-65 μmol g⁻¹ h⁻¹ representing 5-10 fold enhancement, tonoplast transport enabling vacuolar accumulation 20-50 μmol hexose g⁻¹ h⁻¹, and resulting berry Brix increasing from 7-11° to 13-18° over 6-10 days as transport capacity establishes foundation for subsequent rapid accumulation. Molecular mechanisms of transport activation inclu transcriptional induction of hexose transporter genes through sugar-responsive cis-elements (SURE, TGGACGG motifs) activated by sugar signaling via hexokinase pathway creating feed-forward amplification, ABA-responsive elements in transporter gene promoters responding to veraison ABA spike (50-200 → 600-1500 ng/g) coordinating transport activation with other ripening processes, transcription factor families (VvMYB, VvbZIP, VvNAC) showing coordinate induction binding transporter gene promoters orchestrating expression, and chromatin remodeling potentially involved with histone modifications enabling access to previously silenced transporter genes during developmental transition. Hexose transporter family members showing differential activation patterns inclu VvHT1 (high-affinity glucose transporter, Km 1.0-2.5 mM) induced 40-100 fold early in activation providing capacity for efficient uptake even at moderate apoplastic concentrations, VvHT2 (facilitating both glucose and fructose, Km 4-10 mM) showing 30-80 fold elevation slightly delayed providing bulk transport capacity, VvHT3, VvHT5, VvHT6 induced 25-70 fold each contributing to total transport capacity, and isoform diversity creating functional redundancy and potential regulation through differential subcellular localization or kinetic properties. Protein synthesis and trafficking during activation demonstrate: transporter mRNA translation producing protein at rates matching transcript increases with synthesis reaching 40-70% of maximum within 3-5 days of activation onset, protein trafficking through ER and Golgi to plasma membrane requiring 2-6 hours from synthesis to insertion providing temporal lag between transcript and functional capacity, membrane insertion requiring integration into lipid bilayer with transporter proteins estimated 5-12% of total plasma membrane protein at activation compared to 31°C potentially impairing activation through reduced transcription or protein misfolding, water status with moderate deficit (stem ψ -0.9 to -1.3 MPa) compatible with normal activation while severe deficit may delay through general metabolic limitation, and nutritional status particularly nitrogen and sulfur required for protein synthesis with adequate nutrition essential for rapid transporter production. Hormonal regulation of transport activation demonstrates: ABA signaling central with rising levels 600-1500 ng/g activating ABA-responsive elements in transporter gene promoters, sugar itself creating positive feedback through hexokinase-mediated signaling where initial hexose uptake generates glucose-6-phosphate signals enhancing transporter gene expression amplifying accumulation, declining gibberellin and auxin removing growth-related suppression allowing metabolic focus on storage, and potential ethylene involvement in some cultivars with slight elevation correlating with activation timing though mechanism unclear. Cultivar variation in transport activation kinetics shows: rapid-activating varieties (Chardonnay, Gewürztraminer) achieving 60-80% maximum transport capacity within 4-6 days post-activation onset through rapid gene induction and protein synthesis, moderate cultivars (Merlot, Cabernet Sauvignon) reaching 50-70% maximum in 6-9 days, and slower types (Nebbiolo, Mourvèdre) requiring 8-12 days for 45-65% capacity reflecting genetic differences in regulatory control and biosynthetic rates. Physiological consequences of transport activation inclu establishment of import capacity that fundamentally determines achievable sugar accumulation rates and final levels, creation of concentration gradients enabling continued uptake throughout ripening, and osmotic effects with vacuolar hexose accumulation driving water import causing berry expansion 15-30% during activation period despite declining growth rates illustrating osmotic component of berry development. Quality implications inclu transport activation completeness strongly correlates with final sugar levels (r² = 0.75-0.88) with inadequate activation limiting ultimate Brix regardless of substrate supply, timing of activation relative to other ripening processes affecting compositional balance important for wine style, and uniformity of activation across berries and clusters determining ripening synchrony affecting harvest decisions. Viticultural factors affecting activation inclu crop load with excessive yields potentially delaying activation 3-7 days through limited resources for transporter synthesis, canopy management ensuring adequate nitrogen for protein synthesis supporting transporter production, water management with severe deficit potentially impairing activation while moderate deficit compatible, and disease or environmental stress during activation period potentially causing long-term limitation of sugar accumulation capacity even if conditions later improve making this a critical developmental window. Monitoring transport activation in research settings involves: gene expression analysis tracking VvHT transcript levels confirming induction, Western blotting detecting transporter protein accumulation, transport activity assays measuring hexose uptake into isolated mesocarp cells or vesicles, and sugar accumulation rates in berries indirectly indicating transport capacity establishment, while practical viticulture relies on Brix measurements showing accelerating accumulation rates indicating successful activation.
Sugar_Transport_Activation is distinguished from Phloem_Unloading_Activation by representing membrane transport rather than vascular unloading, from Hexose_Transport_Initiation by showing coordinated system-wide activation rather than individual component onset, and from maximum transport by representing establishment (30-100 fold gene induction, 5-10 fold uptake increase) rather than peak operation. Duration 6-10 days BBCH 83-84. Critical developmental window. Sugar_Transport_Activation establishes cellular import machinery fundamentally determining achievable berry sweetness.
Hexose-Aufnahme-Maschinerie-Aktivierung
Membran-Transport-System-Induktion
Transporter-Gen-Hochregulation
Zucker-Import-Kapazitäts-Etablierung
hexose uptake machinery activation
membrane transport system induction
sugar import capacity establishment
transporter gene upregulation
Hayes MA, Davies C, Dry IB (2007) Isolation, functional characterization, and expression analysis of grapevine hexose transporters J Exp Bot 58:1985-1997 transporter activation Afoufa-Bastien D, Medici A, Jeauffre J, Coutos-Thévenot P, Lemoine R, Atanassova R, Laloi M (2010) The Vitis vinifera sugar transporter gene family Plant Mol Biol 72:533-546 transporter family regulation Lecourieux F, Kappel C, Lecourieux D, Serrano A, Torres E, Arce-Johnson P, Delrot S (2014) An update on sugar transport and signaling in grapevine J Exp Bot 65:821-832 comprehensive transport activation.
Sugar Transport Activation
In a Bordeaux Cabernet Franc vineyard, sugar uptake enhancement was tracked over 10 days in early September Year N+1 (BBCH 84) when glucose uptake increased from 32 to 72 μmol g⁻¹ h⁻¹, fructose from 28 to 65 μmol g⁻¹ h⁻¹, berry Brix accelerated from 15.2° to 21.8° (3.6% per week), vacuolar hexose rose from 720 to 1280 mM, berry fresh weight 1.72 g expanding 13%, under favorable conditions (daily high 26-28°C, stem ψ -1.05 MPa, healthy canopy).
Sugar_Uptake_Enhancement represents the progressive amplification of hexose import rates into mesocarp cells during early-to-mid veraison as transporter protein accumulates, membrane capacity increases, and metabolic coupling optimizes creating increasingly efficient cellular hexose acquisition that drives escalating berry sugar accumulation. This enhancement occurs during early-to-mid veraison (BBCH 83-84), typically 4-10 days post-transporter upregulation onset, and is characterized by glucose uptake rates increasing from initial upregulation 15-35 μmol g⁻¹ h⁻¹ to enhanced phase 45-85 μmol g⁻¹ h⁻¹ representing 2-4 fold amplification, fructose uptake similarly rising from 12-30 μmol g⁻¹ h⁻¹ to 40-75 μmol g⁻¹ h⁻¹, combined hexose import 60-130 μmol g⁻¹ h⁻¹ enabling berry Brix increases accelerating from 1.8-2.8% per week to 3.0-4.2% per week as cellular import machinery approaches maximum efficiency. Cellular mechanisms underlying enhancement inclu continued transporter protein synthesis and membrane insertion increasing density 2-4 fold from initial activation levels, membrane remodeling potentially expanding plasma membrane area 15-35% through vesicle fusion creating additional sites for transporter integration, tonoplast membrane area substantially increased 40-80% through vacuolar expansion and proliferation enhancing cytoplasm-to-vacuole transport capacity, and optimization of metabolic coupling where cytoplasmic hexose metabolism adjusted maintaining low concentrations 50-90 mM sustaining concentration gradient favoring continued plasma membrane uptake despite rising vacuolar levels. Transport kinetics during enhancement demonstrate: glucose uptake Vmax increasing from 35-65 μmol g⁻¹ h⁻¹ at enhancement onset to 60-100 μmol g⁻¹ h⁻¹ approaching maximum through higher transporter density, fructose Vmax similarly enhanced from 30-55 μmol g⁻¹ h⁻¹ to 50-90 μmol g⁻¹ h⁻¹, Km values remaining constant (glucose 1.5-3.0 mM, fructose 5-12 mM) confirming capacity increases without affinity changes, and competition between hexoses diminishing as transporter capacity exceeds apoplastic supply creating non-saturating conditions enabling maximum flux. Concentration gradients driving enhancement show: apoplastic hexoses maintained 220-400 mM combined through continued invertase cleavage of phloem-delivered sucrose, cytoplasmic hexoses held 55-90 mM through balanced plasma membrane import, metabolism, and tonoplast export, vacuolar accumulation rising from 500-800 mM at enhancement onset to 900-1400 mM during peak creating osmotic potential declining from -1.4 to -2.0 MPa to -2.0 to -2.8 MPa, and maintained apoplast-cytoplasm gradient 160-340 mM and cytoplasm-vacuole gradient 450-1300 mM driving facilitated diffusion at increasing rates. Membrane dynamics supporting enhancement inclu transporter protein representing 8-14% of total plasma membrane protein (from 4-8% at activation onset) approaching maximum packing density, phospholipid composition potentially modified increasing membrane fluidity enhancing transport protein function though specific changes poorly characterized, membrane potential maintained -120 to -150 mV (inside negative) through H⁺-ATPase activity providing electrochemical gradient potentially contributing to transport driving force though most VvHTs appear electroneutral, and cholesterol or sterol content potentially adjusted affecting membrane physical properties and transport protein activity. Metabolic integration during enhancement shows: glycolytic flux increasing from 18-35 μmol glucose g⁻¹ h⁻¹ to 28-55 μmol g⁻¹ h⁻¹ metabolizing portion of imported hexoses for energy and biosynthesis, respiratory rates elevated 1.2-2.4 μmol O₂ g⁻¹ h⁻¹ supporting ATP synthesis for membrane pumps maintaining gradients, respiratory quotient >1.1-1.4 indicating substrate-level and fermentative metabolism contribute substantially particularly in berry interior, and metabolic pathway adjustments redirecting carbon from structural uses to storage with 70-80% of imported hexoses accumulating in vacuoles while 20-30% metabolized. Hormonal regulation during enhancement includes: ABA sustained at 1000-2000 ng/g maintaining ripening program activation and supporting continued transporter expression, hexose signaling through hexokinase pathway with glucose-6-phosphate 20-45 μmol/g fresh weight creating strong sugar-responsive gene activation feedback, declining gibberellin maintaining low levels removing any residual suppression, and potential ethylene effects in some cultivars with slight elevation 0.2-0.9 nL g⁻¹ h⁻¹ possibly fine-tuning ripening though role unclear. Environmental influences on enhancement rate inclu temperature critically important with optimal 25-29°C supporting maximum enhancement achieving peak rates 3.8-4.5% Brix per week within 6-8 days, moderate warmth 22-25°C enabling 3.0-3.8% per week over 8-11 days, cooler conditions 18-22°C reducing to 2.2-3.0% per week with 10-15 day enhancement period, heat stress 31-34°C impairing enhancement to 1.8-2.8% per week through membrane disruption and enzyme impairment, and extreme heat >34°C potentially preventing adequate enhancement <1.5% per week limiting final sugar achievement. Water status effects show: adequate vine hydration supporting normal enhancement with stem water potential -0.6 to -1.2 MPa enabling full metabolic function, moderate deficit -1.0 to -1.4 MPa compatible with enhancement and potentially concentrating berries through reduced expansion, severe deficit <-1.5 MPa impairing enhancement 35-60% through general metabolic limitation despite continued transporter expression, and osmotic relationships with high vacuolar sugar creating -2.0 to -2.8 MPa requiring adequate vine water status to sustain gradient for continued uptake. Nutritional influences inclu nitrogen availability supporting protein synthesis and enzyme production with N-deficient vines showing 20-35% reduced enhancement despite adequate transporter genes through translational or post-translational limitations, potassium potentially important for membrane transport and pH regulation with berry K⁺ accumulating from 1400-2000 to 2200-3000 mg/kg during enhancement possibly affecting membrane function, and phosphorus required for ATP synthesis supporting active pumps maintaining gradients enabling transport. Cultivar variation in enhancement characteristics shows: rapid-enhancing varieties achieving 70-85% maximum uptake capacity within 6-9 days of enhancement onset, moderate cultivars reaching 60-75% maximum in 8-12 days, and slower types requiring 10-16 days for 50-65% capacity reflecting genetic differences in membrane dynamics, metabolic coordination, or regulatory control. Coordination with phloem delivery shows: sugar uptake enhancement synchronized with apoplastic unloading intensification ensuring substrate availability matches cellular import capacity, temporal relationship typically showing uptake enhancement lagging unloading intensification by 1-3 days allowing apoplastic substrate accumulation before maximum cellular demand, and feedback coordination where high uptake rates potentially stimulate phloem delivery through reduced apoplastic sugar maintaining unloading gradient. Quality implications of enhancement inclu uptake enhancement rate and completeness determining trajectory toward final sugar levels with inadequate enhancement limiting ultimate Brix, timing relative to other ripening processes affecting balance between sugar and other quality parameters (acids, phenolics, flavors), and uniformity of enhancement across berries and clusters determining ripening synchrony important for harvest maturity consistency. Viticultural management during enhancement involves: maintaining adequate vine water and nutritional status supporting metabolic function enabling normal enhancement, avoiding environmental stresses particularly heat that can irreversibly impair enhancement limiting final sugar potential, disease control preventing infections that disrupt metabolism interfering with enhancement, and monitoring enhancement progress through Brix tracking showing accelerating accumulation rates indicating successful amplification.
Sugar_Uptake_Enhancement is distinguished from Hexose_Transporter_Upregulation by representing functional capacity increase (2-4 fold uptake amplification) rather than gene expression, from Sugar_Transport_Activation by showing progressive enhancement rather than system onset, and from peak uptake by exhibiting amplification (45-85 μmol g⁻¹ h⁻¹) rather than maximum (65-110 μmol g⁻¹ h⁻¹). Duration 6-12 days BBCH 83-84. Temperature optimal 25-29°C. Sugar_Uptake_Enhancement represents critical escalation creating capacity for achieving harvest-ready sweetness levels.
amplified hexose import
amplifizierter Hexose-Import
cellular sugar acquisition enhancement
escalating glucose-fructose uptake
eskalierende Glukose-Fruktose-Aufnahme
progressive Aufnahme-Zunahme
progressive uptake increase
zelluläre Zucker-Akquisitions-Verstärkung
Keller M (2020) The Science of Grapevines 3rd Ed pages 221-225 uptake enhancement dynamics Thomas TR, Matthews MA, Shackel KA (2006) Direct in situ measurement of cell turgor in grape berries during development Australian J Grape Wine Res 12:173-180 cellular physiology during enhancement Lecourieux F, Kappel C, Lecourieux D, Serrano A, Torres E, Arce-Johnson P, Delrot S (2014) An update on sugar transport and signaling in grapevine J Exp Bot 65:821-832 transport enhancement mechanisms.
Sugar Uptake Enhancement
Sulfur content quality in Sauvignon Blanc affects both fermentation health and varietal thiol expression, with adequate cysteine (20-40 mg/L) and glutathione (50-150 mg/L) content supporting the cysteine conjugate precursor pool for 3MH release while providing yeast nutrition fruit from vineyards with optimal sulfur nutrition and careful fungicide management (last sulfur application 6+ weeks pre-harvest) produces the clean fermentations and intense tropical thiol expression that define premium Marlborough Sauvignon Blanc, while excessive sulfur residues or deficient sulfur amino acids compromise both fermentation and aromatic quality.
Sulfur_Content_Quality represents the chemical quality characterizing the concentration of sulfur-containing compounds in grape berry tissue, capturing the essential macronutrient incorporated into amino acids, proteins, and numerous secondary metabolites affecting wine aroma and stability. Sulfur in grapes exists in multiple forms including inorganic sulfate, the sulfur-containing amino acids cysteine and methionine, glutathione (a cysteine-containing tripeptide), and various secondary metabolites including glucosinolates and sulfur-containing aroma precursors. Total sulfur in grape juice typically ranges from 50-200 mg/L, with the organic fraction (amino acids, glutathione) of particular quality significance. Cysteine and methionine are essential amino acids whose availability affects protein synthesis and yeast nutrition, while glutathione provides antioxidant protection and participates in aroma precursor chemistry (thiol conjugate formation). Sulfur nutrition in vineyards derives from soil sulfate uptake, with most soils providing adequate sulfur though deficiency can occur in sandy, low-organic matter soils. Elemental sulfur applied as fungicide contributes to vine sulfur status and can affect wine composition through residue carryover. In winemaking, grape-derived sulfur compounds affect fermentation through yeast nutrition (sulfur amino acid supply) and wine quality through volatile sulfur compound formation. Excessive sulfur residues from late-season fungicide application can produce hydrogen sulfide and other reduced sulfur compounds during fermentation, creating off-odors. Conversely, adequate sulfur amino acid and glutathione content supports healthy fermentation and provides substrates for beneficial thiol formation. Sulfur analysis requires methods appropriate to the forms of interest, with total sulfur measured by combustion analysis or ICP-OES, and specific compounds by HPLC or enzymatic methods.
Sulfur_Content_Quality addresses a macronutrient with complex chemistry due to its occurrence in multiple oxidation states and organic forms. The distinction between nutritional sulfur (sulfate, amino acids) and residual fungicide sulfur is important for quality implications. Glutathione content has emerged as a significant quality parameter due to its antioxidant function and role in thiol precursor chemistry. The potential for hydrogen sulfide formation during fermentation links grape sulfur status (both nutritional and residual) to wine off-odor risk.
Beerenschwefelqualität
S content
S-Gehalt
Schwefelgehalt
Schwefelverbindungskonzentration
berry sulfur quality
sulfur compound concentration
sulfur level
Rauhut D (2009) Usage and Formation of Sulphur Compounds. In: König H et al. (eds) Biology of Microorganisms on Grapes, in Must and in Wine, Springer pp.: 181-207 Lavigne V et al. (2007) Potential of Different Saccharomyces Cerevisiae Strains to Produce Typical Aromatic Thiols in Wine. Australian Journal of Grape and Wine Research 13(3): 121-129
Sulfur Content Quality
Thompson Seedless (Sultanina, Sultana, Kishmish) bears a Sultana_Production_Scion_Role throughout Turkish, Australian, and South African production, where grapes are dipped in 2.5% potassium carbonate plus 0.5% dipping oil emulsion before sun-drying for 4-7 days, producing light amber "sultanas" with softer texture and milder flavor than natural raisins. Sultana (as the variety is named in Australia) serves in a Sultana_Production_Scion_Role as the primary variety for Australian sultana production, with dipping treatment and Murray-Darling's hot, dry climate enabling efficient production for export to European markets. Merbein Seedless bears a Sultana_Production_Scion_Role as an Australian-bred improvement on Sultana, with enhanced productivity and equivalent dipping response for sultana production.
A raisin production scion role that inheres in a grafted scion cultivar by virtue of its suitability for sultana production, a category of light-colored dried grapes intermediate between golden raisins and natural raisins, traditionally produced by dipping grapes in alkaline solutions (potassium carbonate, "dipping oil" emulsions) before sun-drying to accelerate moisture loss and produce characteristic light amber color. Sultana production scions develop the thin, waxy cuticle that responds effectively to dipping treatment, which dissolves or disrupts the cuticle layer to create micro-cracks permitting rapid water evaporation during subsequent drying. The dipping process reduces drying time from 14-21 days (natural) to 4-7 days (dipped), significantly reducing weather risk and improving drying efficiency. The name "sultana" is used variably—referring to the dipping process and product (particularly in Australia, Turkey, and UK), to the Thompson Seedless/Sultanina variety itself, or to both. Sultana production concentrates in Turkey (the world's largest producer), Australia, Iran, and Mediterranean regions.
Sultana_Production_Scion_Role captures a processing method defined by the alkaline dipping treatment that accelerates drying and produces characteristic light color without sulfur dioxide application. The terminological overlap between variety name (Sultana/Sultanina = Thompson Seedless) and product name (sultana = dipped-dried raisin) creates confusion ontologically, this role refers to the production method and product category rather than variety genetics. The dipping chemistry (potassium carbonate plus ethyl oleate emulsion) creates microscopic cuticle damage permitting rapid water loss, a traditional technique documented since antiquity. Sultana production offers efficiency advantages over natural raisin drying (shorter drying period, reduced weather risk) without the sulfur dioxide treatment of golden raisins, positioning sultanas between these categories in both processing intensity and consumer perception.
Sultanina raisin grape
Sultanina-Rosinentraube
Sultaninen-Traube
dipped raisin grape
getauchte Rosinentraube
oil-dipped raisin variety
sultana grape
Christensen LP (2000) Raisin Production Manual, UC ANR Publication 3393 Pangavhane DR et al. (1999) Solar dryers for grape drying, Energy Conversion and Management 40: 1-23 Grncarevic M, Lewis WJ (1976) Dipping treatment for dried grapes, Food Technology in Australia 28: 66-76 Australian Dried Fruits Association (2020) Production Guidelines
Sultana Production Scion Role
A summer dormant bud on lateral shoots of Vitis vinifera cv. Tempranillo in August in La Rioja, Spain, entering growth arrest during a three-week period of temperatures exceeding 38°C and soil water deficit below -1.5 MPa, with bud scales slightly closed and apical meristem activity suspended, resuming active growth within 10 days following 25mm irrigation and temperature moderation to 28°C.
A dormant bud that enters temporary growth arrest during the active growing season in response to environmental stress conditions such as severe heat, drought, or nutrient deficiency, characterized by ecodormancy that is immediately reversible upon stress alleviation, lacking the deep metabolic suppression and chilling requirements of winter dormancy. Summer dormant buds retain green or partially lignified tissues beneath protective scales, maintain higher metabolic rates than winter dormant buds, and resume growth rapidly (within days to weeks) when favorable conditions return, representing a survival strategy for Mediterranean and warm-climate viticulture where mid-season stress is common.
Summer dormancy differs fundamentally from winter dormancy in being imposed by external environmental factors (ecodormancy) rather than internal physiological programming (endodormancy). No chilling requirement exists for release from summer dormancy, only alleviation of the stress condition. Summer dormancy is most common in warm, dry climates and can affect both primary growing points and secondary lateral buds. Extended summer dormancy can reduce total seasonal growth and crop yield but may enhance wood maturation and cold hardiness development.
Sommerruheknospe
quiescent summer bud
sommerruhendeKnospe
stress-induced dormant bud
summer resting bud
Based on Düring (1978) "Studies on the environmentally controlled stomatal transpiration in grape vines" Vitis 17=97-104 Lovisolo et al. (2010) "Effects of water stress on vessel size and xylem hydraulic conductivity in Vitis vinifera L." Journal of Experimental Botany 61=2749-2759 Tombesi et al. (2014) "Stomatal closure is induced by hydraulic signals and maintained by ABA in drought-stressed grapevine" Scientific Reports 4=6552.
Summer Dormant Bud
A sun-exposed mature leaf at node 6 on an exterior shoot of Vitis vinifera cv. Sangiovese in Chianti, Italy, positioned on the western canopy face receiving approximately 75% of full sunlight (daily photosynthetic photon flux density averaging 1200 μmol photons m⁻² s⁻¹), measuring 11.5 cm length by 10.8 cm width representing 35% size reduction compared to interior shade leaves, lamina thickness of 315 μm with three well-developed palisade layers totaling 125 μm thickness, chlorophyll content of 520 μg cm⁻² (compared to 380 μg cm⁻² in shade leaves), cuticle thickness of 5.8 μm on adaxial surface providing enhanced UV protection, stomatal density of 195 stomata mm⁻² on abaxial surface (compared to 155 mm⁻² in shade leaves), moderate trichome density creating light gray-green undersurface appearance, dark green adaxial surface with slight bronze tinge from anthocyanin accumulation, demonstrating light-saturated photosynthesis rate of 14.2 μmol CO₂ m⁻² s⁻¹ substantially exceeding shade leaf rates, measured at BBCH stage 83 (berries developing color) in early August, representing characteristic sun leaf adaptations maximizing photosynthetic performance under high light conditions.
A mature leaf developing and functioning in high light environments receiving greater than 60-70% of full sunlight in Vitis vinifera, characterized by phenotypic adaptations to high irradiance including smaller lamina size typically 20-40% reduced compared to shade leaves on the same vine, thicker lamina often 280-350 μm due to increased cell layers and wall thickness, multiple palisade mesophyll layers (2-3 layers) increasing photosynthetic capacity per unit area, higher chlorophyll concentrations per unit area but lower per unit mass reflecting increased tissue density, thicker cuticle providing enhanced protection against UV radiation and water loss, higher stomatal density but smaller stomatal size optimizing gas exchange while minimizing water loss, increased density of trichomes on abaxial surfaces, darker green coloration from higher chlorophyll concentrations, elevated concentrations of photoprotective compounds including carotenoids and anthocyanins, and anatomical features conferring higher light-saturated photosynthesis rates of 12-16 μmol CO₂ m⁻² s⁻¹ but also higher light compensation points reflecting increased respiratory costs of maintaining robust photosynthetic apparatus. Sun-exposed mature leaves develop these specialized characteristics through developmental plasticity responding to high light during the expansion phase, with light quality and quantity during the first 2-3 weeks post-emergence determining anatomical differentiation patterns, and continued high light exposure after maturation maintaining physiological acclimation including sustained high photosynthetic capacity, enhanced photoprotective mechanisms including xanthophyll cycle operation and non-photochemical quenching, and elevated antioxidant systems protecting against reactive oxygen species generated under high light conditions.
Sun-exposed mature leaves are distinguished from shade-adapted leaves by structural and functional characteristics reflecting high light acclimation, requiring assessment of anatomical features, pigment composition, and physiological performance under ambient conditions. The sun versus shade distinction represents a continuum rather than discrete categories, with intermediate phenotypes developing under moderate light conditions (30-60% full sun). Developmental timing of light exposure critically determines leaf phenotype, with high light during expansion producing structural sun leaf characteristics that persist throughout leaf life, while shade-developed leaves exposed to high light after expansion show limited plastic responses and may suffer photoinhibition. Position in canopy strongly predicts sun versus shade status, with exterior canopy faces and upper canopy positions typically producing sun leaves while interior and lower canopy zones produce shade leaves. Sun leaves provide higher per-area photosynthesis but lower per-mass efficiency compared to shade leaves, with optimal canopy architecture balancing these leaf types to maximize whole-canopy carbon gain.
außenkanopie Weinblatt
exterior canopy leaf
high light leaf
hochlicht Blatt
light-adapted leaf
sonnenexponiertes reifes Blatt
Based on Boardman (1977) "Comparative photosynthesis of sun and shade plants" Annual Review of Plant Physiology 28=355-377 Schultz (1996) "Leaf absorptance of visible radiation in Vitis vinifera L.= estimates of age and shade effects with a simple field method" Scientia Horticulturae 66=93-102 Patakas et al. (2003) "Differences in stomatal and photosynthetic responses of Vitis vinifera L. (cv. Sultana) leaves in response to progressive water stress" Vitis 42=113-115 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 179-214.
Sun Exposed Mature Leaf
A sun exposed upper canopy apical leaf at node 14 on an exterior shoot of Vitis vinifera cv. Grenache in southern Rhône Valley, France, positioned at canopy top on the south-facing exterior receiving approximately 92% of full sunlight (daily PPFD averaging 1550 μmol photons m⁻² s⁻¹ with midday peaks exceeding 2000 μmol photons m⁻² s⁻¹), measuring 11.2 cm length by 10.5 cm width showing smaller size typical of sun leaves despite apical position, exhibiting very thick lamina of 358 μm with three well-developed palisade layers totaling 132 μm thickness, very thick cuticle of 7.5 μm with substantial wax deposits, high chlorophyll content of 575 μg cm⁻², dense trichome coverage creating conspicuous gray-white abaxial appearance, very dark green adaxial surface with bronze tinge from anthocyanin accumulation, demonstrating very high light-saturated photosynthesis rate of 15.2 μmol CO₂ m⁻² s⁻¹ near maximum for grapevine, high light compensation point of 65 μmol photons m⁻² s⁻¹, efficient photoprotection maintaining PSII efficiency of 0.81 despite high light stress, occurring at 2.5 weeks post-emergence during late expansion under continuous high light exposure, measured at shoot BBCH stage 83 (berries developing color) in mid-August during peak summer irradiance, representing maximum sun exposure and adaptation among upper canopy apical leaves.
An upper canopy apical leaf experiencing very high light exposure in exterior elevated canopy positions in Vitis vinifera, characterized by position on outer canopy faces or canopy tops receiving 75-95% of full sunlight with minimal shading, development under high irradiance conditions during expansion creating pronounced sun leaf anatomical characteristics including very thick lamina often 320-380 μm with three or occasionally four palisade cell layers, very thick cuticle frequently 6-8 μm providing enhanced UV protection and water retention, high chlorophyll concentrations per unit area of 500-600 μg cm⁻², elevated stomatal density with small individual stomata optimizing gas exchange, substantial trichome coverage particularly on abaxial surfaces, dark green to bronze-green coloration from high chlorophyll and anthocyanin concentrations, high photoprotective carotenoid content, and physiological characteristics including high light-saturated photosynthesis rates of 12-16 μmol CO₂ m⁻² s⁻¹, high light compensation and saturation points, and robust photoprotective mechanisms enabling sustained function under potentially damaging high irradiance. Sun exposed upper canopy leaves develop the maximum expression of sun leaf syndrome among apical zone leaves, benefiting from favorable elevated positions providing excellent light exposure during the critical expansion period when anatomical characteristics are determined, creating leaves optimized for high light environments that contribute disproportionately to whole-vine carbon gain through superior photosynthetic performance under the high light conditions prevalent in upper exterior canopy zones.
Sun exposed upper canopy leaves represent the extreme high-light end of the apical leaf spectrum, distinguished from other upper canopy leaves by very pronounced sun leaf characteristics reflecting development under maximum light exposure. These leaves show the most complete expression of sun leaf syndrome including maximum palisade layer number and thickness, highest cuticle development, greatest photosynthetic capacity per unit area, and strongest photoprotective mechanisms. Position on exterior south-facing (Northern Hemisphere) or north-facing (Southern Hemisphere) canopy surfaces and elevated canopy tops creates the highest light exposure among all canopy zones, with full sun exposure throughout most of the photoperiod. Development during high summer irradiance periods (June-August) creates maximum sun leaf differentiation, while earlier or later development under lower light may produce less pronounced sun characteristics despite similar positions. Functional performance shows superior photosynthetic rates under high light but potentially reduced efficiency under lower light compared to shade leaves, with high light compensation points requiring substantial irradiance for positive carbon balance. These leaves face greatest risks of photoinhibition, heat stress, and water stress damage, requiring robust protection mechanisms to maintain function through peak summer conditions.
exterior elevated leaf
high-light upper apical leaf
hochlicht oberes apikales Blatt
maximum Sonnen-Exposition Weinblatt
maximum sun exposure apical leaf
sonnenexponiertes obere Kanopie Blatt
Based on Boardman (1977) "Comparative photosynthesis of sun and shade plants" Annual Review of Plant Physiology 28=355-377 Schultz (1996) "Leaf absorptance of visible radiation in Vitis vinifera L.= estimates of age and shade effects with a simple field method" Scientia Horticulturae 66=93-102 Poni et al. (2009) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 60=397-407 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 215-258.
Sun Exposed Upper Canopy Leaf
Field evaluation of Sunburn_Resistance_Disposition at UC Davis Oakville station monitored sunburn incidence and severity across 20 cultivars in a trial where half of clusters received artificial shade (70% reduction) while matched clusters remained exposed over two seasons with multiple heat events (> 40°C). Sunburn was scored on exposed clusters using a 0-5 scale based on affected berry surface area. Sunburn_Resistance_Disposition ranged from highly resistant (mean score 3.5). 'Grenache' exhibited exceptional Sunburn_Resistance_Disposition with mean score of 0.3 on exposed clusters despite berry surface temperatures exceeding 48°C during peak heat events, attributable to its thick, waxy berry skin (cuticle 8.2 μm versus 4.5 μm average) and high skin flavonoid content. 'Pinot Noir' showed extreme susceptibility (score 3.8) with visible necrosis appearing within 48 hours of severe heat events its thin skin (3.2 μm cuticle), low reflectance, and limited antioxidant capacity provided minimal protection. White cultivars showed generally higher Sunburn_Resistance_Disposition than red cultivars when skin thickness was equivalent, reflecting lower radiation absorption. Shaded clusters showed negligible sunburn across all cultivars (scores < 0.3), confirming that Sunburn_Resistance_Disposition operates against the specific combination of heat plus light, not heat alone. Berry skin carotenoid concentration correlated negatively with sunburn severity (r = -0.65), supporting the photoprotective role of these compounds.
A Sunburn_Resistance_Disposition is a Berry_Heat_Tolerance_Disposition that inheres in Vitis vinifera grape berries and is realized through mechanisms that prevent or limit tissue necrosis caused by the combined effects of high temperature and intense solar radiation on exposed berry surfaces during ripening. This disposition represents resistance to sunburn, a common disorder in hot-climate viticulture manifested as browning, tissue collapse, and desiccation of sun-exposed berry surfaces, rendering affected berries unsuitable for premium wine production. Sunburn results from the interaction of excessive berry heating (from absorbed solar radiation) and photochemical damage (from UV and visible light), with the combination causing oxidative stress, membrane damage, chlorophyll destruction (in pre-véraison berries), anthocyanin degradation (in ripening red berries), and eventually cell death. Sunburn_Resistance_Disposition depends on berry skin properties (thickness, pigmentation, wax content affecting radiation absorption and reflection), antioxidant capacity (carotenoids, flavonoids, ascorbate reducing oxidative damage), heat shock protein accumulation, and repair mechanisms enabling recovery from moderate stress. Cultivars with thicker, more reflective skins, higher antioxidant content, and greater heat-induced protective responses show enhanced Sunburn_Resistance_Disposition. Additionally, cluster positioning and canopy management interact with intrinsic berry resistance to determine field sunburn incidence, with exposed clusters at higher risk regardless of cultivar.
Sunburn_Resistance_Disposition is_a Berry_Heat_Tolerance_Disposition. Sunburn requires the combination of high temperature and intense light, distinguishing it from purely thermal damage. Skin properties (thickness, reflectance, pigmentation) and antioxidant content (carotenoids, flavonoids) are primary determinants. Red berries are generally more susceptible than white due to higher radiation absorption by anthocyanins. Canopy management interacts strongly with intrinsic resistance, as shading eliminates sunburn regardless of cultivar susceptibility.
Beeren-Sonnenbrand-Toleranz
Solarschadens-Resistenz
Sonnenbrand-Resistenz-Disposition
berry sunburn tolerance
solar injury tolerance
sun damage resistance
Spayd SE et al. (2002) Separation of sunlight and temperature effects on the composition of Vitis vinifera cv. Merlot berries, American Journal of Enology and Viticulture 53:171-182 Greer DH, La Borde D (2006) Sunburn of grapes affects wine quality, Australian and New Zealand Grapegrower and Winemaker 506:21-23 Reshef N et al. (2017) Sunlight modulates fruit metabolic profile and shapes the spatial pattern of compound accumulation in grape berries, Frontiers in Plant Science 8:70
Sunburn Resistance Disposition
Sunburn severity quality assessment in Napa Valley evaluated cultivar differences in damage susceptibility during a late-season heat event coinciding with véraison when berry pigmentation and thermal tolerance were actively changing. Sauvignon Blanc demonstrated high sunburn severity quality with 45% of exposed berries showing visible damage, progressing from bronze discoloration (score 2) on berries receiving 4+ hours direct afternoon sun to complete desiccation (score 5) on extremely exposed berries, with berry temperatures measured at 48°C on affected fruit. Cabernet Sauvignon showed moderate sunburn severity quality with 20% berry damage, predominantly limited to score 2 discoloration with only 3% progressing to severe damage, attributed to thicker skin and developing anthocyanin pigmentation providing partial protection. Pinot Noir exhibited intermediate sunburn severity quality with 30% damage incidence reflecting its thin skin vulnerability partially offset by moderate canopy density. The sunburn severity quality assessment established berry temperature thresholds for damage initiation at 35°C for Sauvignon Blanc and 42°C for Cabernet Sauvignon, informing cultivar-specific recommendations for fruit zone shading targets and irrigation scheduling during heat events.
Sunburn severity quality is a heat damage severity quality that inheres in grape berries and represents the extent and intensity of thermal and photo-oxidative damage to berry skin and underlying flesh resulting from excessive solar radiation exposure under high temperature conditions, ranging from minor cosmetic browning that may have minimal quality impact through moderate damage affecting berry composition and wine quality to severe injury causing berry desiccation, off-flavors, and complete quality loss in affected fruit. This quality addresses the most economically significant form of heat damage in viticulture, with sunburn affecting both the appearance of table grapes and the composition and wine quality from affected wine grapes through alterations in phenolic profiles, aromatic compounds, and the development of undesirable flavors associated with heat-damaged fruit. The ontological significance of sunburn severity quality lies in its direct impact on fruit quality and value, with economic losses from sunburn damage potentially substantial in warm climate viticulture, motivating investments in protective management practices and cultivar selection decisions that consider sunburn susceptibility alongside other quality attributes. The biological basis of sunburn encompasses both thermal damage from berry temperatures exceeding tolerance thresholds (typically 35-40°C depending on cultivar) and photo-oxidative damage from excessive UV and visible radiation causing reactive oxygen species accumulation that overwhelms antioxidant defenses, with the combination of high temperature and high light intensity creating synergistic damage exceeding that from either stress alone. The progression of sunburn damage follows a characteristic sequence beginning with loss of chlorophyll and anthocyanins creating brown or bronze discoloration, progressing through cell death in skin and outer flesh layers, and culminating in berry shriveling and desiccation under severe conditions. Sunburn susceptibility varies substantially among cultivars, with thin-skinned, light-colored varieties generally more susceptible than thick-skinned, darkly pigmented types, and with fruit zone exposure, cluster architecture, and canopy management practices strongly modulating damage incidence regardless of inherent cultivar susceptibility.
Sunburn_Severity_Quality is_a Heat_Damage_Severity_Quality, representing thermal and photo-oxidative damage specifically to grape berries. This class addresses the primary fruit quality concern from heat events. Berry temperature measurement (often exceeding ambient by 10-15°C) provides key diagnostic data. Severity scoring from 1 (minor discoloration) to 5 (complete desiccation) enables standardized assessment. Cultivar susceptibility varies substantially with skin thickness and pigmentation. The synergy between temperature and radiation distinguishes sunburn from purely thermal damage. Critical economic impact motivates significant management attention.
Beerensonnenbrand
Sonnenbrand-Schweregrad
Sonnenschaden
berry sunburn
solar injury
solare Verletzung
sun damage
sunscald
Greer DH, Weedon MM (2013) The impact of high temperatures on Vitis vinifera cv. Semillon grapevine performance and berry ripening. Frontiers in Plant Science 4:491 Rustioni L et al. (2014) Grape berry sunburn symptoms studied by Raman spectroscopy. Journal of Raman Spectroscopy 45:1079-1086 Caravia L et al. (2016) Application of shade treatments during Shiraz berry ripening to reduce the impact of high temperature. Australian Journal of Grape and Wine Research 22:422-437
Sunburn Severity Quality
Sunshine hours for the Wachau, Austria average 1800-1900 hours annually with 1400-1500 hours during the April-October growing season, contributing to the distinctive ripeness profile of Grüner Veltliner and Riesling on the steep Danube terraces. Jerez, Spain records 3000-3100 annual sunshine hours, among the highest of any viticultural region globally, producing the intense oxidative conditions that characterize Fino and Oloroso Sherry maturation. The Rheingau averages 1550-1650 sunshine hours annually, but the south-facing Rhine Valley slopes effectively increase the solar beam interception, compensating for the moderate sunshine duration to achieve reliable Riesling ripening at 50°N latitude.
Sunshine hours is a solar radiation data entity that records the cumulative duration of direct solar beam radiation (exceeding the Campbell-Stokes threshold of 120 W/m² or equivalent automated sensor threshold) received at a viticultural site, expressed in hours per day, month, growing season, or year, serving as a historically established and widely available proxy metric for solar energy receipt and viticultural radiation characterization.
Sunshine hours remain the most historically extensive solar radiation dataset available for many viticultural regions, with records extending back over a century from manual Campbell-Stokes recorder networks. While now largely superseded by direct pyranometer measurements for precision applications, sunshine hours data retains value for long-term climate trend analysis and viticultural zoning. The Huglin Index incorporates a day-length coefficient (d) that functions as a latitude-dependent sunshine proxy, acknowledging the longer photoperiods at higher latitudes. Conversion between sunshine hours and total radiation requires empirical Ångström-Prescott coefficients (Rs = Ra × [a + b × n/N]) calibrated to regional atmospheric conditions. The correlation between sunshine hours and wine quality parameters is well documented — Bordeaux vintage ratings show strong positive correlation (r > 0.6) with growing season sunshine hours, particularly sunshine during August-September when phenolic maturation occurs. Modern automated sunshine sensors (photoelectric or pyrheliometric) eliminate the threshold subjectivity of traditional Campbell-Stokes recorders.
Besonnung
Besonnungsdauer
Sonnenscheindauer
Sonnenscheinstunden
Sonnenstunden
hours of bright sunshine
insolation hours
solar hours
sunshine duration
sunshine total
Gladstones J (2011) Wine, Terroir and Climate Change, Wakefield Press, pp.: 77-85 Jones GV, Davis RE (2000) Climate influences on grapevine phenology, grape composition, and wine production and quality for Bordeaux, France. American Journal of Enology and Viticulture 51(3): 249-261 Huglin P (1978) Nouveau mode d'évaluation des possibilités héliothermiques d'un milieu viticole. Comptes Rendus de l'Académie d'Agriculture de France 64: 1117-1126
Sunshine Hours
Tempranillo vines in the Pagos Viejos block at Bodegas Muga in Rioja Alta bearing a super-premium wine role for the Muga Selección Especial Reserva are managed at yields of 4-6 t/ha from 40-60 year old vines, with green harvesting removing 30-40% of clusters at véraison, hand harvesting with field sorting, and 24 months in new French oak barriques, producing 3,000-5,000 cases annually at 35-50 €/bottle positioned between the estate's premium Reserva (15-20 €) and the icon-tier Torre Muga (65-90 €).
A quality tier role that inheres in a grapevine whose fruit is allocated to wines positioned in the upper echelon of a producer's portfolio below icon status, sourced from superior vineyard parcels with significant yield restriction and intensive viticultural management, marketed at price points reflecting exceptional quality and limited availability.
Super_Premium_Wine_Role represents the tier immediately below icon status, typically encompassing wines at $20-50 USD retail in the US market. This segment is characterized by named vineyard or block designations, significant barrel aging programs, and viticultural practices approaching icon-level intensity: many producers position their "Reserve," "Reserva," "Riserva," or single-vineyard bottlings in this tier. The super-premium tier is commercially important as it captures consumers trading up from premium wines and represents the entry point for wine collectors and fine wine retail channels.
Reserve-Wein-Rolle
Spitzenwein-Rolle
Super-Premium-Wein-Rolle
reserve wine role
super-premium wine vine role
upper-tier production role
Rabobank (2019) Wine Quarterly Report: Global Wine Market Segmentation McMillan R (2022) State of the US Wine Industry, Silicon Valley Bank pp: 14-18 Lockshin L and Corsi AM (2012) Consumer behaviour for wine 2.0: a review since 2003, Wine Economics and Policy 1: 2-23
Super Premium Wine Role
Cabernet Sauvignon flowers at BBCH 59-65 (flower buds swollen through full bloom) exhibit characteristic superior ovaries positioned prominently at the pistil base, elevated 0.4-0.6 mm above the receptacle surface where sepals and petals attach, with ovary completely free from receptacle fusion and clearly distinguishable as a separate structure, expanding post-fertilization from 0.9-1.1 mm diameter at anthesis to 3.5-4.5 mm diameter at BBCH 73 (young berries, shot berries falling) without constraints from receptacle or perianth tissues. Syrah pistil longitudinal sections reveal superior ovary organization with ovary base positioned above the level of stamen filament attachment and petal insertion, ovary wall consisting entirely of carpel-derived tissues without receptacle contribution, and clear separation between ovary tissues and receptacle tissues visible in cellular organization and vascular patterning. Riesling berry development from BBCH 71 through BBCH 89 (fruit set through berries ripe for harvest) shows continuous superior positioning with expanding berry (derived from ovary) remaining free and elevated above the receptacle throughout development, with final berry size (11-15 mm diameter) achieved through ovary tissue expansion alone without receptacle tissue involvement. Comparative floral morphology studies demonstrate that Vitis superior ovaries contrast fundamentally with inferior ovaries in families like Rosaceae (apples, where receptacle forms fruit flesh) or Cucurbitaceae (melons, where receptacle fuses with ovary), highlighting the distinctive superior ovary condition as a defining Vitaceae characteristic maintained across all Vitis species and cultivars.
A superior ovary is an ovary positioned above the attachment points of the other floral organs (sepals, petals, stamens) on the receptacle, with the ovary free and not fused to or enclosed by the receptacle or perianth tissues, representing the universal ovary position in Vitis flowers and contrasting with inferior ovaries (ovaries positioned below other floral organs and often fused with receptacle or floral tube tissues) characteristic of some other plant families. Superior ovaries in Vitis are characterized by= (1) ovary positioned at or near the apex of the receptacle with the ovary base (where ovary attaches to receptacle) at or above the level of sepal, petal, and stamen attachment points (2) ovary completely free from fusion with receptacle, floral tube, or perianth tissues, with ovary wall representing only carpel-derived tissues without contribution from receptacle or other floral organs (3) hypogynous floral organization with sepals, petals, and stamens attached to the receptacle below and separate from the ovary rather than fused to ovary tissues (4) visible ovary positioned prominently in the flower center, elevated above the receptacle base and clearly distinguishable from supporting structures. The superior ovary position in Vitis has developmental and functional significance= superior position means ovary development proceeds independently of other floral organs, with ovary growth and post-fertilization expansion not constrained by fusion with receptacle or perianth tissues the free ovary can expand rapidly post-fertilization (increasing from 0.6-1.2 mm diameter at anthesis to 6-25 mm diameter at maturity) without requiring coordinated expansion of fused receptacle or perianth tissues superior position allows early ovary visibility and accessibility for developmental studies and for assessment of pistil health and functionality. The superior versus inferior ovary position represents a fundamental organizational difference among angiosperm families, with superior ovaries generally considered ancestral and inferior ovaries representing derived conditions evolved independently in multiple lineages. Vitis retention of superior ovaries across all species reflects the ancestral Vitaceae condition, with no evidence of evolutionary transitions toward inferior ovary organization within the family. The superior ovary position in Vitis creates distinctive berry (fruit) characteristics= berries develop from ovary alone without contribution from receptacle or perianth tissues, contrasting with false fruits or accessory fruits in inferior ovary taxa where receptacle tissues contribute to fruit structure berry morphology reflects ovary organization directly, with bilocular ovary producing berries with two seed-containing chambers (when seeds develop) or bilocular organization evident in seedless berries superior berry development allows independent maturation without coordination with perianth or receptacle tissues, potentially providing greater developmental flexibility. The superior ovary position is evident throughout flower development and anthesis= in flower buds (BBCH 51-59), the ovary is visible as a distinct swelling at the pistil base positioned above the developing calyx, corolla, and androecium at anthesis (BBCH 61-65), the ovary remains clearly elevated above the receptacle with sepals, petals, and stamens attached below during early fruit development (BBCH 71-75), the expanding ovary/berry remains free and elevated above the receptacle even as dramatic size increases occur.
Superior_Ovary represents the universal ovary position in Vitis and must be distinguished from inferior ovaries (found in other plant families but not in Vitis) where ovary position is below other floral organs and often involves ovary-receptacle fusion. The superior position is a fundamental aspect of Vitis floral organization and determines berry developmental characteristics (ovary-only fruit without receptacle contribution). Sub-classification into Elevated and Free types emphasizes positional and structural aspects of superior organization. Understanding superior ovary organization is important for comparative floral morphology, for interpreting berry developmental biology, and for understanding the evolutionary context of Vitaceae floral architecture within angiosperms.
free ovary
freier Fruchtknoten
hypogynous ovary
oberständiger Fruchtknoten
oberständiges Ovar
Superior Ovary
Cabernet Sauvignon supporting hypodermal layer providing strong structural support= 2-3 cell layers with cells 20-28 μm diameter, walls 2-2.8 μm thick creating continuous supporting tissue 45-60 μm total thickness, contributing to thick skin (total 110-145 μm) mechanical strength measured as 12-18 N force required to puncture skin, with supporting hypodermal layer providing estimated 30-40% of total skin tensile strength based on layer thickness and wall properties, creating robust skin suitable for extended maceration without excessive skin breakdown. Syrah supporting hypodermal layer in thick-skinned cultivar= 2-4 cell layers creating 48-68 μm supporting zone with walls 2.2-3.2 μm providing substantial structural support, contributing to Syrah characteristic thick firm skin (120-170 μm total) resisting tearing during harvest and maceration, with thick supporting hypodermal layer enabling extended 20-30 day maceration achieving full tannin extraction without skin disintegration creating grainy texture from excessive skin particle release. Pinot Noir supporting hypodermal layer in thin-skinned cultivar= 1-2 cell layers creating 22-38 μm supporting zone with walls 1.5-2.4 μm providing moderate structural support, contributing to Pinot Noir characteristic thin delicate skin (60-95 μm total) requiring gentle handling during harvest and gentle extraction during winemaking (5-10 day maceration, minimal punchdown vigor) to avoid excessive skin breakdown from limited supporting hypodermal layer structural capacity. Merlot supporting hypodermal layer berry expansion accommodation= veraison (BBCH 81, week 10) showing hypodermal cells 18-24 μm diameter maturity (BBCH 89, week 18) showing cells 24-32 μm (33-45% increase), demonstrating supporting hypodermal layer ability to accommodate berry expansion through cell enlargement while maintaining structural support function, with hypodermal expansion limited compared to middle pericarp (which doubles in size) creating transition from small outer to large inner cells over skin-flesh boundary. Thompson Seedless table grape supporting hypodermal layer shipping durability role= supporting hypodermal layer 1-2 layers, 28-42 μm thickness providing moderate structural support contributing to skin strength adequate for table grape shipping over 2-3 weeks cold storage and transport, with maintaining supporting hypodermal layer integrity (through careful handling avoiding bruising) critical for preventing skin breakdown and decay during extended post-harvest period, demonstrating supporting hypodermal layer importance for table grape post-harvest quality beyond anthocyanin pigmentation function emphasized in red wine cultivars.
A supporting hypodermal layer is the hypodermal layer emphasizing its structural support role as transition zone between protective outer epidermis and soft fleshy inner mesocarp, focusing on mechanical properties including moderately thick cell walls, compact cell arrangement, and contribution to skin strength and integrity. Supporting hypodermal layer structural characteristics= (1) moderately thick cell walls 1.5-3 μm providing structural strength intermediate between very thick protective epidermal walls (3-6 μm) and thin storage mesocarp walls (0.4-0.8 μm) (2) small compact cells 15-30 μm diameter with limited intercellular spaces creating dense cellular tissue providing mechanical support (3) cell arrangement creating continuous tissue layer(s) providing structural continuity supporting overlying epidermis and cuticle (4) wall composition including cellulose, hemicellulose, pectin providing tensile strength and flexibility (5) potential wall modifications including lignification in some cases enhancing rigidity. Supporting hypodermal layer mechanical functions= (1) skin strength contribution providing structural support for outer protective layers, with hypodermal layer contributing to overall skin tensile strength and tear resistance (2) load distribution helping distribute mechanical forces (compression during berry packing, tension during expansion) across skin layers preventing stress concentration in single layer (3) epidermis support providing cellular foundation supporting thin-walled epidermal cells and preventing epidermal buckling or collapse under compression (4) cuticle anchorage providing structural base for cuticle attachment creating integrated cuticle-epidermis-hypodermis protective system. Supporting hypodermal layer in berry expansion and growth= (1) accommodation of berry expansion through cell wall extension and remodeling during Phase II ripening, with hypodermal cells showing 30-60% diameter increase (versus 120-200% in middle pericarp) during expansion (2) limited expansion creating gradient in cell size from small supporting hypodermal cells (15-30 μm) to large storage middle pericarp cells (80-180 μm) over 50-150 μm depth (3) maintenance of structural integrity during expansion preventing skin tearing or separation from flesh as berry enlarges (4) potential wall thickening during expansion maintaining wall thickness despite cell area increase through continued wall deposition. Supporting hypodermal layer in berry texture and quality= (1) contribution to skin toughness perceived during eating or processing, with hypodermal layer adding to overall skin thickness and firmness (2) influence on skin-flesh separation characteristics affecting ease of peeling in some table grape applications (3) contribution to berry splitting resistance with stronger supporting hypodermal layer potentially reducing splitting incidence during late-season rain events (4) maintenance during ripening preventing skin deterioration or senescence that would compromise berry integrity. Supporting hypodermal layer in thick-skinned versus thin-skinned cultivars= thick-skinned cultivars (Cabernet Sauvignon, Syrah) showing 2-4 hypodermal cell layers creating thicker supporting zone 40-65 μm providing stronger structural support thin-skinned cultivars (Pinot Noir, Gamay) showing 1-2 layers creating thinner supporting zone 20-40 μm providing less structural support supporting layer thickness correlating with overall skin thickness and mechanical properties affecting cultivar suitability for different production purposes (thick skins favoring full-bodied wines and table grape shipping, thin skins favoring elegant wines and fresh consumption).
Supporting_Hypodermal_Layer emphasizes the structural support and mechanical functions of the hypodermis and is distinguished from Sub_Epidermal_Hypodermal_Layer which emphasizes positional and light-related aspects. Supporting hypodermal layer represents the hypodermis from the perspective of skin strength, mechanical protection, and structural integrity. Understanding supporting hypodermal layer is relevant for comprehending skin mechanical properties, berry firmness and splitting resistance, cultivar differences in skin strength affecting winemaking and table grape applications, and for managing berry structure through developmental factors affecting hypodermal layer development and wall properties.
Festigungsschicht
strengthening layer
structural sub-epidermal layer
strukturelle Hypodermis
stützende Hypodermis-Schicht
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 180-185 Thomas, T.R. et al. (2008). Changes in the physical properties of grape berries. American Journal of Enology and Viticulture 59=398-409 Considine, J.A., Kriedemann, P.E. (1972). Fruit splitting in grapes. Australian Journal of Agricultural Research 23=17-27.
Supporting Hypodermal Layer
A suppressive dominant apical shoot on a bilateral cordon of Vitis vinifera cv. Tempranillo in Rioja, Spain, originating from a lateral bud at node 6 following winter freeze damage to the original terminal bud, growing 195 cm vertically by August while simultaneously suppressing development of six adjacent fruiting positions within a 40 cm radius, where lateral shoots remained weak (15-25 cm length) with small leaves (35-50 cm² area) and poorly developed fruit clusters showing increased shatter and delayed ripening, with detailed carbohydrate partitioning studies showing the suppressive dominant apical shoot capturing 38% of total photosynthate production from the affected cordon section despite representing only 18% of total leaf area.
A dominant apical shoot that, while having assumed apical position and terminal growth characteristics, exhibits growth vigor and apical dominance effects strong enough to significantly suppress development of neighboring shoots on the same vine structure or adjacent structures, creating localized zones of reduced vegetative growth, altered shoot angles, and modified resource allocation patterns in Vitis vinifera. Suppressive dominant apical shoots function as powerful sink organs that redirect photosynthates, mineral nutrients, and water toward their own growth at the expense of surrounding vegetative and potentially reproductive structures, creating spatial asymmetries in canopy development and fruit production that can persist throughout the growing season and influence dormant pruning decisions. These shoots typically emerge from favorable positions with preferential vascular connections to root system resources, often at upright angles that maximize light capture, and may achieve such vigor that adjacent lateral shoots remain small, weak, or completely inhibited despite release from traditional apical dominance suppression. Suppressive dominant apical shoots represent both management challenges in established vineyards where balanced canopy distribution is desired, and opportunities in framework establishment where concentrated growth in specific zones accelerates structural development toward training system targets.
Suppressive dominant apical shoots are distinguished from merely vigorous dominant apical shoots by their documented negative effects on neighboring vegetative or reproductive structures, requiring comparative assessment of growth parameters in affected versus unaffected zones. The suppressive mechanism operates through multiple pathways including direct hormonal signaling, competitive resource capture from shared vascular systems, and physical shading of lower or adjacent shoots. Management strategies include early-season removal if suppression threatens crop value, strategic topping to redistribute growth resources, or intentional maintenance if rapid framework extension justifies temporary production sacrifice. The degree of suppression depends on shoot position, vascular architecture, vine vigor status, and relative timing of bud break between competing shoots.
competitively dominant shoot
konkurrenz-dominanter Spitzentrieb
neighboring-suppressive shoot
resource-monopolizing apical shoot
ressourcen-monopolisierender Trieb
suppressiv-dominanter Trieb
Based on Cline (1997) "Concepts and terminology of apical dominance" American Journal of Botany 84=1064-1069 Edson et al. (1995) "Influence of crop load on vegetative growth, dry matter partitioning, and mineral nutrient partitioning of Chenin blanc grapevines" American Journal of Enology and Viticulture 46=469-477 Candolfi-Vasconcelos & Koblet (1990) "Yield, fruit quality, bud fertility and starch reserves of the wood as a function of leaf removal in Vitis vinifera" Vitis 29=199-221 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 245-271.
Suppressive Dominant Apical Shoot
Surface appearance quality assessment spans both viticultural and enological contexts, with vineyard observation noting berry bloom integrity (intact waxy coating indicating healthy berry with no handling damage), gloss development (increasing with ripeness), and absence of surface defects (no visible rot or pest damage), while wine service observation notes surface clarity (no visible film indicating absence of aerobic spoilage), appropriate mousse behavior in sparkling wine (persistent fine foam indicating quality), and absence of floating matter (no cork fragments or other contamination), demonstrating the dual relevance of surface assessment across grape and wine evaluation.
Surface appearance quality is a visual appearance quality that represents the optical properties observed at the wine surface and grape berry surface, encompassing reflectivity, texture, coatings, and visible surface features that provide information about wine condition, grape health, and berry maturity in viticultural and enological contexts. Within the BFO framework, this quality addresses the surface-specific dimension of visual assessment, distinguishing surface observations from bulk properties (color, clarity, viscosity) while recognizing that surface appearance provides unique diagnostic information about both wine in glass and grapes on vine. The ontological significance lies in surface appearance bridging viticultural and enological observation, where grape berry surface characteristics (bloom, gloss, defects) inform harvest decisions and quality assessment while wine surface characteristics (foam, film, foreign matter) indicate condition and potential faults, making surface observation relevant across the grape-to-wine continuum. The physical basis for surface appearance involves light interaction at the air-liquid or air-berry interface, including specular reflection (creating gloss), diffuse reflection from surface coatings (bloom/pruina), and visibility of surface-localized phenomena (foam persistence, film formation, floating particles). The compositional and biological determinants of surface appearance include epicuticular wax composition and structure (affecting grape bloom and gloss), wine surface-active compounds (affecting foam behavior), microbial growth (creating visible surface films in spoilage), and foreign matter contamination (visible at the wine surface). The dual application context requires clarity about whether surface assessment addresses grape berry observation (relevant for harvest decision and grape quality evaluation) or wine observation (relevant for serving assessment and fault detection), with different surface properties emphasized in each context. Understanding surface appearance quality enables grape maturity and health assessment from berry surface observation, wine condition evaluation from surface characteristics, and appropriate quality judgment incorporating surface appearance as a diagnostic indicator.
Surface_Appearance_Quality serves as the parent class for surface-related visual qualities (Bloom, Gloss, Defect Visibility), addressing the interface-specific dimension of visual assessment. This quality bridges viticulture and enology, with grape berry surface and wine surface both relevant assessment contexts. The class supports quality evaluation across the grape-to-wine continuum through surface-specific observation.
Grenzflächenqualität
Oberflächenerscheinung
Oberflächenoptik
exterior quality
interface appearance
surface characteristics
surface optical quality
äußere Erscheinung
Casado CG, Heredia A (1999) Structure and dynamics of reconstituted cuticular waxes of grape berry cuticle. Journal of Experimental Botany 50(331): 175-182 Rosenquist JK, Morrison JC (1989) Some factors affecting cuticle and wax accumulation on grape berries. American Journal of Enology and Viticulture 40(4): 241-244
Surface Appearance Quality
Table grape production of Crimson Seedless targets berry surface area specifications that optimize visual appeal and eating quality, with premium fruit exhibiting individual berry surface areas of 5.5-7.0 cm² corresponding to equatorial diameters of 20-24 mm, achieved through aggressive cluster thinning at fruit set (removing 30-50% of clusters), gibberellic acid application to promote cell elongation, and careful irrigation management during the cell expansion phase to maximize berry volume while maintaining skin integrity.
Surface_Area_Quality represents the two-dimensional extent of the external boundary enclosing a three-dimensional grapevine organ. This quality captures the interface area between plant tissues and the surrounding environment, which determines the capacity for exchange of energy, gases, water vapor, and dissolved substances. Surface area is fundamentally important for understanding and predicting rates of physiological processes that occur at organ boundaries. Berry surface area governs water loss through cuticular transpiration, uptake of foliar-applied nutrients and agrochemicals, and the total amount of skin-derived compounds available for extraction during vinification. Root surface area, particularly of fine absorbing roots, determines the capacity for water and mineral nutrient uptake from soil. Shoot and cane surface area influences bark photosynthesis contribution, winter desiccation rates, and overwintering pathogen inoculum capacity. Cluster surface area affects spray penetration, drying rate after precipitation, and thus disease susceptibility. Surface area measurement for irregular three-dimensional organs requires either geometric approximation (modeling organs as combinations of simple shapes), surface integration methods, or empirical calibration relating surface area to more easily measured parameters such as mass or volume. For berries, the assumption of ellipsoidal geometry enables surface area estimation from two diameter measurements using standard formulas. Modern three-dimensional scanning and photogrammetric techniques enable direct surface area quantification for complex organ geometries.
Surface_Area_Quality specifically addresses the external boundary of organs and is distinct from Cross_Sectional_Area_Quality (internal planar extent) and Leaf_Area_Quality (specialized laminar surface). For hollow organs or organs with internal cavities, surface area may refer to external surface only or may include internal surfaces depending on the functional context. Berry surface area calculations commonly assume smooth surfaces, though microscale features including epicuticular wax structure contribute to the actual physical surface area at finer scales. The relationship between surface area and volume (surface-to-volume ratio) has important physiological implications addressed separately.
Grenzflächenbereich
Oberflächenqualität
boundary area
external surface extent
externe Flächenausdehnung
interface area quality
organ surface area
Coombe BG and McCarthy MG (2000) Dynamics of Grape Berry Growth and Physiology of Ripening. Australian Journal of Grape and Wine Research 6(2): 131-135 Rogiers SY et al. (2004) Mineral Sinks within Ripening Grape Berries (Vitis vinifera L.). Vitis 43(2): 65-72
Surface Area Quality
Chardonnay showing surface lateral roots, excavation revealing lateral roots 8-22 cm depth in A horizon topsoil, surface laterals 1.8-4.2 cm diameter extending 60-140 cm horizontal distance, laterals showing abundant fine root production with feeder root density 8.4 cm/cm³ soil substantially higher than deeper laterals 2.1 cm/cm³, isotope tracing showing surface laterals providing 58% of nitrogen uptake accessing topsoil nutrients, surface lateral roots demonstrating intensive topsoil resource exploitation
Surface lateral root is lateral shallow root at very shallow depth typically 5-25 cm near soil surface, characterized by position in upper topsoil layers where organic matter highest and biological activity most intense, exposure to greatest temperature fluctuations and moisture variability, vulnerability to surface disturbances, and serving intensive topsoil resource exploitation particularly important for accessing surface-applied fertilizers and irrigation water, with surface lateral roots showing characteristics including concentration in A horizon topsoil, predominantly horizontal orientation spreading laterally at near-constant shallow depth, moderate diameter 1-5 cm depending on order and age, moderate to high root hair and fine root production creating dense absorptive networks in favorable upper soil, seasonal dynamics with surface root activity peaking spring and fall when topsoil conditions optimal, and management sensitivity with surface roots highly responsive to and affected by tillage, irrigation, fertilization, mulching, and other surface-oriented practices. Surface lateral root depth characteristics includes= showing very shallow position (surface laterals typically 5-25 cm depth in upper topsoil much shallower than deeper laterals 30-60 cm), exhibiting A horizon association (surface roots predominantly in A horizon soil where organic matter and biological activity concentrated), demonstrating near-surface maintenance (surface laterals maintaining shallow depth rather than descending deeper), revealing exposure potential (surface laterals occasionally exposed at soil surface particularly on slopes or eroded sites), and displaying frost risk (very shallow surface roots potentially vulnerable to freezing in cold climates without snow cover). Surface lateral root topsoil advantages includes= showing high nutrient availability (surface laterals accessing nutrients concentrated in topsoil from organic matter decomposition and fertilizer application), exhibiting optimal biological activity (surface zone supporting most diverse and abundant microbial communities benefiting root function), demonstrating favorable structure (topsoil often showing better structure and aeration than subsoil supporting root growth and function), revealing moisture access (surface laterals efficiently capturing rainfall and irrigation before deep percolation), and displaying temperature moderation (topsoil warming quickly spring promoting early root activity). Surface lateral root environmental variability includes= showing temperature fluctuations (surface laterals experiencing large diurnal and seasonal temperature variations affecting function), exhibiting moisture extremes (surface soil rapidly wetting and drying creating dynamic moisture environment), demonstrating nutrient pulses (surface laterals experiencing nutrient concentration pulses following fertilization or organic matter mineralization), revealing frost exposure (surface laterals potentially experiencing freezing temperatures absent in deeper soil), and displaying summer heat (surface roots potentially experiencing heat stress in hot climates without irrigation or mulch). Surface lateral root vulnerability to disturbance includes= showing tillage susceptibility (surface laterals highly vulnerable to cutting or damage from cultivation given shallow position), exhibiting traffic impact (surface roots easily compacted by vehicle or foot traffic), demonstrating erosion risk (soil erosion potentially exposing or removing surface laterals), revealing construction damage (vineyard operations or construction readily damaging surface roots during soil disturbance), and displaying weed competition (surface laterals competing intensely with weeds concentrated in same topsoil zone). Surface lateral root absorptive specialization includes= providing surface resource capture (surface laterals specialized for accessing resources concentrated in topsoil), exhibiting fertilizer access (surface laterals efficiently capturing surface-applied or shallow-incorporated fertilizers), demonstrating irrigation response (surface roots responding strongly to irrigation maintaining function during summer), revealing rain capture (surface laterals quickly responding to rainfall events capturing moisture before deep drainage), and displaying seasonal importance (surface laterals particularly important spring and fall when topsoil conditions favorable). Surface lateral root fine root production includes= showing abundant feeder roots (surface laterals producing dense populations of fine absorptive roots in favorable topsoil), exhibiting high turnover (surface zone fine roots showing high production and mortality rates responding to variable conditions), demonstrating mycorrhizal associations (surface fine roots often showing high arbuscular mycorrhizal colonization enhancing uptake), revealing root hair abundance (surface laterals and derivatives producing abundant root hairs given favorable topsoil conditions), and displaying critical contribution (surface lateral-associated fine roots potentially providing 40-70% of total vine uptake during favorable periods). Surface lateral root management implications includes= requiring tillage alternatives (surface lateral vulnerability suggesting no-till or minimal tillage approaches), showing mulch benefits (surface mulches protecting surface laterals from temperature extremes, moisture fluctuations, disturbance), exhibiting irrigation importance (surface laterals depending on irrigation maintaining function during summer drought), demonstrating fertilizer efficiency (fertilizer placement considering surface lateral distribution optimizing uptake efficiency), and revealing protection value (protecting abundant surface laterals from damage critical for maintaining vine productivity). Surface lateral root seasonal dynamics includes= showing spring activity peak (surface laterals and fine roots showing maximum activity spring when topsoil moist and warming), exhibiting summer stress potential (surface roots potentially experiencing drought or heat stress summer if irrigation inadequate), demonstrating fall recovery (surface root activity recovering fall with cooler temperatures and autumn rainfall), revealing winter dormancy (surface root function minimal winter though roots surviving if soil not frozen), and displaying annual patterns (surface root abundance and function showing strong seasonal variation reflecting topsoil condition changes).
Surface_Lateral_Root emphasizes very shallow near-surface position and is distinguished from deeper laterals by focus on topsoil association. Includes Near_Surface_Lateral_Root, Topsoil_Surface_Lateral_Root, and Spreading_Surface_Lateral_Root. Critical for understanding topsoil specialization, environmental variability, disturbance vulnerability, intensive absorption, and the very shallow laterals exploiting resource-rich topsoil layers.
Flache Oberbodenwurzel
Near-surface lateral
Oberflächen-Lateralwurzel
Shallow topsoil root
Topmost lateral root
Surface Lateral Root
Syrah surface lenticel at 15 years post-planting appears as 2.2-3.4 mm diameter light tan oval spot on grey-brown cork surface, slightly raised 0.2-0.3 mm above surrounding bark, lenticel showing porous texture visible with hand lens contrasting with smooth dense adjacent cork, exposed surface receiving atmospheric oxygen at 21% concentration enabling inward diffusion supporting inner bark respiration, lenticel surface showing minor weathering (slight darkening at edges) from 3-4 years environmental exposure, lenticel density 5-7 per cm² creating scattered spot pattern across bark surface clearly visible on mature trunk, surface position making lenticels identifiable feature for bark assessment
Surface lenticel is lenticel emphasizing its position at bark exterior where it interrupts the outer cork surface creating visible porous spots or marks, representing the atmospheric interface of the gas exchange pathway and determining lenticel visibility and accessibility. Surface lenticel appearance characteristics includes= visible spots on bark (lenticels appearing as distinct marks 1-4 mm diameter contrasting with surrounding cork), lighter coloration (tan to light brown lenticel tissue typically lighter than weathered grey-brown cork surface creating conspicuous spots), raised, flush, or depressed (young lenticels often slightly elevated 0.1-0.5 mm above surface, mature lenticels becoming flush or depressed as surrounding bark thickens), oval to circular shape (lenticels typically round to slightly elongated, occasionally forming short lines), and textural differences (lenticel surface often slightly rougher or more porous appearing than smooth dense cork). Surface lenticel distribution patterns includes= scattered arrangement (lenticels distributed relatively uniformly across bark surface without pronounced clustering), density variation (2-8 lenticels per cm² varying by cultivar, bark age, and environmental conditions), occasional vertical alignment (lenticels sometimes arranged in short vertical rows particularly on young smooth bark), association with former structures (initial lenticels often developing beneath stomata, bud scars, or leaf scars creating predictable patterns), and proportional coverage (lenticels occupying 2-5% of total bark surface area maintaining predominantly continuous cork covering). Surface lenticel as atmospheric interface includes= direct exposure to air (lenticel surface in contact with atmosphere enabling gas exchange with environment), entry point for oxygen (atmospheric oxygen at 21% concentration diffusing into lenticel beginning inward journey to inner bark), exit point for carbon dioxide (CO₂ from inner bark respiration emerging at lenticel surface releasing to atmosphere), exposure to environmental conditions (lenticels experiencing full range of temperature, humidity, precipitation, and other atmospheric variations), and vulnerability to physical factors (lenticels subject to dust accumulation, water infiltration, frost effects, and mechanical damage at exposed surface). Surface lenticel morphology variation includes= prominent raised lenticels (clearly projecting 0.2-0.6 mm above surrounding cork creating obvious bumps), flush lenticels (surface even with adjacent cork, lenticels distinguished primarily by color and texture), depressed lenticels (sunken 0.1-0.3 mm below surface particularly in thick bark where surrounding cork accumulates faster than lenticel fills), cracked lenticel surfaces (some lenticels showing fissures or irregular surfaces from cork cell arrangement), and weathered appearance (older lenticels showing surface erosion or discoloration from prolonged environmental exposure). Surface lenticel on different bark types includes= smooth bark lenticels (highly conspicuous as distinct spots on uniform young bark surfaces), furrowed bark lenticels (less obvious, located in furrow bottoms or on ridge sides blending with bark texture), platy bark lenticels (appearing on bark plate surfaces, potentially affected by plate separation), exfoliating bark lenticels (lenticels on peeling bark layers, lost when outer bark sheds requiring inner lenticel exposure), and thick bark lenticels (lenticels maintaining surface opening despite substantial cork accumulation, depth increasing to maintain pathway through thickening barrier). Surface lenticel visibility and identification includes= easy observation on young smooth bark (lenticels clearly visible as spots 1-3 mm diameter on light brown young trunk surface), less obvious on old furrowed bark (lenticels blending with complex bark texture requiring close examination), color contrast aiding identification (lighter lenticel tissue contrasting with darker weathered cork), hand lens examination (magnification revealing lenticel porous structure and boundary with dense cork), and moistening enhancing visibility (wetted bark showing enhanced color contrast making lenticels more apparent). Surface lenticel environmental exposure effects includes= weathering causing surface changes (UV radiation, freeze-thaw, moisture cycling gradually darkening and roughening lenticel surfaces), rain water contact (lenticels receiving direct precipitation potentially wetting internal air spaces though infiltration depth limited), dust and particle accumulation (atmospheric particulates potentially coating lenticel surfaces reducing apparent porosity), frost effects (ice formation in surface lenticel cells potentially disrupting loose cork structure), and sun exposure (intense radiation potentially affecting lenticel tissue chemistry and appearance particularly on southwest trunk exposure). Surface lenticel and bark texture includes= lenticels contributing to bark surface character (scattered spots or marks adding to overall bark appearance and texture), lenticel prominence varying by cultivar (some varieties showing large conspicuous lenticels creating distinctive bark pattern, others small inconspicuous structures), lenticel-furrow interactions (lenticels in furrowed bark often located in furrow bottoms or slopes affecting visibility and function), and aesthetic considerations (lenticel patterns contributing to distinctive trunk appearance valued in old vines). Surface lenticel seasonal changes includes= lenticel expansion during growing season (loose tissue potentially swelling slightly 5-15% facilitating enhanced gas exchange April-September), potential closing layer formation (some cultivars developing surface compaction during fall reducing lenticel prominence and function during dormancy), winter appearance maintenance (lenticels remaining visible through dormant season though potentially slightly contracted or modified by closing layers), spring reopening (closing layers potentially disrupted as bark expands and metabolism increases restoring full lenticel function), and year-round visibility (lenticels typically remaining identifiable throughout annual cycle despite seasonal modifications). Surface lenticel vulnerability at bark exterior includes= mechanical damage (exposed position making lenticels vulnerable to abrasion, impact, or scratching potentially disrupting gas exchange structure), pathogen landing site (lenticel surfaces potentially receiving fungal spores or bacteria though penetration typically limited), insect interaction (some boring insects potentially targeting lenticels as entry points though intact lenticel structure usually resistant), frost injury (severe cold potentially damaging surface lenticel cells affecting function), and importance of protection (maintaining intact surface lenticels essential for continued gas exchange, damaged lenticels potentially requiring formation of new lenticels to restore function). Surface lenticel in wound healing includes= traumatic surface lenticels forming in wound periderm (new lenticels appearing on callus tissue surface as wound covering matures), initially small inconspicuous (newly formed lenticels 0.5-1.5 mm diameter gradually enlarging over months), gradual functional development (traumatic lenticels progressively establishing full depth and gas exchange capacity), eventual integration (healed wounds showing functional surface lenticels indistinguishable from developmental lenticels providing normal gas exchange), and importance in large wounds (adequate traumatic lenticel formation essential for supporting metabolism in extensive callus tissue and inner bark beneath healed wounds). Surface lenticel significance includes= determining lenticel visibility (surface position making lenticels observable enabling lenticel assessment and bark characterization), providing atmospheric interface (surface lenticels connecting internal gas exchange pathway to atmosphere), affecting bark appearance (lenticel surface patterns contributing to distinctive trunk aesthetics), enabling gas exchange function (surface opening essential for lenticel function, obstruction or damage compromising oxygen supply to inner bark), and requiring maintenance (preserving surface lenticel integrity important for continued gas exchange supporting inner bark and cambium metabolism).
Surface_Lenticel emphasizes exterior position and visibility and is distinguished from Lenticel (complete structure), Gas_Exchange_Lenticel (function emphasis), and internal lenticel components by focus on atmospheric interface. Critical for understanding lenticel identification and observation, bark surface appearance, environmental exposure effects, lenticel as visible bark feature, atmospheric gas exchange interface, and surface lenticel maintenance importance for preserving gas exchange pathway function supporting inner bark and cambium survival.
Bark surface pore
Exterior lenticel
Oberflächenlenticelle
Rinden-Außenpore
Sichtbare Lenticelle
Visible lenticel
Surface Lenticel
Syrah surface outer pericarp at maturity shows deeply pigmented structure= cuticle and epidermis (28-36 μm) as previously described, hypodermis (2-3 cell layers, 42-58 μm total) containing intense anthocyanin concentration (4.2-5.2 mg/g fresh weight with malvidin-3-glucoside 48-56% of total, creating deep purple-black color), creating total surface outer pericarp thickness 70-94 μm representing 5-6% of pericarp volume but containing 92-96% of total berry anthocyanins (total berry anthocyanin content 2.2-2.8 mg/g reflecting dilution by middle pericarp). Pinot Noir surface outer pericarp showing moderate pigmentation= hypodermis (1-2 cell layers, 22-38 μm) with moderate anthocyanin concentration (2.2-3.2 mg/g), creating surface outer pericarp 45-70 μm thickness, with lighter color (ruby-red versus purple-black Syrah) reflecting lower anthocyanin content and higher malvidin proportion (52-62% malvidin-3-glucoside creating redder hue versus more balanced anthocyanin profile in Syrah creating purple hue). Chardonnay white cultivar surface outer pericarp lacking anthocyanins= hypodermis (1-2 layers, 20-42 μm) containing flavonols (quercetin-3-glucoside 0.9-1.6 mg/g) and carotenoids (β-carotene, lutein providing golden-yellow color), creating surface outer pericarp 38-75 μm with characteristic white/golden appearance from absence of anthocyanins and presence of carotenoid/flavonol pigmentation. Cabernet Sauvignon surface outer pericarp tannin analysis= total tannin 8.5-11 mg/g dry weight skin, procyanidin degree of polymerization 12-16 subunits (mean 14), prodelphinidin content 15-22% of total tannins, galloylation 13-18%, creating characteristic firm tannic structure important for Cabernet Sauvignon wine aging potential, with surface outer pericarp tannins providing 40-55% of total wine tannins (remaining from seeds and inner pericarp) during typical red wine maceration. Merlot surface outer pericarp light exposure comparison= sun-exposed clusters showing surface outer pericarp 72-98 μm thickness with 3.2-4.2 mg/g anthocyanins and 1.4-2.1 mg/g flavonols shaded clusters showing 58-82 μm thickness with 2.4-3.3 mg/g anthocyanins and 0.5-0.9 mg/g flavonols, demonstrating sunlight enhancement of surface outer pericarp thickness and phenolic concentration important for wine color and structure, with exposed berries producing wines with 20-35% higher color intensity and 15-25% higher tannin levels from enhanced surface outer pericarp phenolic content.
A surface outer pericarp is the outermost 2-4 cell layers of the outer pericarp including epidermis, cuticle, and hypodermis, representing the visible and tactile berry surface region typically 35-90 μm total thickness, serving as integrated protective and pigment-bearing layer that determines berry appearance, provides barrier functions, and contains concentrated phenolic compounds including anthocyanins (in red/black cultivars) that fundamentally determine wine color and contribute to wine tannin structure. Surface outer pericarp comprises integrated layers= (1) cuticle and epidermis (outermost 17-43 μm as described in Epidermal_Outer_Pericarp) (2) hypodermis (sub-epidermal layers) typically 1-3 cell layers beneath epidermis, total 18-60 μm thickness, consisting of small cells (15-30 μm diameter) with thicker walls than middle pericarp cells (wall thickness 1.5-3 μm versus 0.4-0.8 μm in middle pericarp), serving as primary site of anthocyanin accumulation in red/black cultivars creating berry color. Surface outer pericarp in red/black cultivars shows characteristic anthocyanin distribution= anthocyanins concentrated primarily in hypodermal cell vacuoles with concentration 2.5-6+ mg/g fresh weight in hypodermis versus 0.1-0.3 mg/g in outer mesocarp and essentially zero in middle pericarp anthocyanin composition including malvidin-3-glucoside (typically 40-60% of total anthocyanins), other monoglucosides (delphinidin, cyanidin, petunidin, peonidin 3-glucosides), and acylated forms (acetyl and p-coumaroyl derivatives) anthocyanin synthesis induced at veraison (BBCH 81) through upregulation of biosynthetic genes (VvMYBA1 transcription factor, structural genes UFGT, DFR, F3H) creating rapid accumulation during ripening total surface outer pericarp anthocyanin content (integrated across all layers) typically 1.8-4 mg/g in deeply colored cultivars, 0.8-2 mg/g in moderately colored cultivars. Surface outer pericarp tannin characteristics= total tannin concentration 6-12 mg/g dry weight (higher than middle pericarp 0.3-1 mg/g) tannin composition predominantly procyanidins with some prodelphinidins tannin polymerization degree 8-18 subunits (higher than seed tannins at 4-10) galloylation percentage 10-20% of subunits tannin localization in cell vacuoles creating astringency upon berry disruption or wine extraction. Surface outer pericarp also contains= flavonols (quercetin, kaempferol, myricetin glycosides) at 0.5-2.5 mg/g with higher levels in sun-exposed berries providing UV protection and contributing to color stability through copigmentation with anthocyanins hydroxycinnamic acids (caftaric, coutaric, fertaric acids) at 0.3-0.8 mg/g contributing to browning reactions stilbenes (resveratrol and derivatives) particularly in stressed berries providing antimicrobial defense. Surface outer pericarp physical characteristics determining berry appearance and texture= surface color determined by anthocyanin content and composition in red/black cultivars (hue, intensity, stability), or carotenoid/flavonol content in white cultivars surface bloom determined by epicuticular wax quantity and crystalloid structure surface texture (smooth versus slightly rough) determined by epidermal cell arrangement and lenticel density skin thickness perception (thick-skinned versus thin-skinned cultivars) reflecting surface outer pericarp thickness and cell wall properties.
Surface_Outer_Pericarp represents the integrated outer layers (epidermis, cuticle, hypodermis) and is distinguished from Epidermal_Outer_Pericarp (epidermis and cuticle only) by inclusion of pigment-bearing hypodermal layers. Surface outer pericarp determines berry appearance and provides the majority of wine color and significant wine tannin contribution. Understanding surface outer pericarp structure and composition is essential for managing berry and wine quality, particularly color development through canopy management and harvest timing decisions. The surface outer pericarp is the primary target for phenolic maturity assessment.
Farbschicht
Hautschicht
Oberflächen-äußeres Perikarp
pigmented outer layers
skin surface layers
Cadot, Y. et al. (2006). Tannin localization and characterization in grape skins. Journal of Agricultural and Food Chemistry 54=2973-2987 Castellarin, S.D. et al. (2007). Transcriptional regulation of anthocyanin biosynthesis. Plant Physiology 143=1347-1363 Downey, M.O. et al. (2006). Cultural practice and environmental impacts on the flavonoid composition of grapes. American Journal of Enology and Viticulture 57=64-75.
Surface Outer Pericarp
Surface texture characterization of young Cabernet Sauvignon shoot tips reveals the dense pubescence (OIV descriptor 004 rating 7-9) with white woolly trichomes covering expanding leaves and internodes that distinguishes this variety from less pubescent cultivars this characteristic surface texture traps boundary layer air, reduces water loss from young tissues, and may deter certain insect herbivores, while also providing the distinctive silvery-white appearance of Cabernet Sauvignon shoot tips that aids vineyard identification during the critical pre-flowering growth assessment period.
Surface_Texture_Quality encompasses all texture qualities that characterize the external boundary of grapevine organs as perceived through tactile contact or visual observation. The organ surface constitutes the interface between plant tissues and the environment, and its texture properties influence water relations, pathogen interactions, light reflection, and aesthetic appearance. This category addresses five major surface conditions: Smooth_Surface_Quality describing surfaces without prominent irregularities Rough_Surface_Quality capturing surfaces with tactile irregularities Pubescent_Surface_Quality addressing hair-covered surfaces Glabrous_Surface_Quality describing hairless surfaces and Waxy_Surface_Quality capturing the bloom or pruina that coats berry and leaf surfaces. Surface texture results from the structural organization of the epidermis and cuticle, including cell shape and arrangement, cuticular thickness and ornamentation, and the presence of trichomes (plant hairs) of various types. In grape berries, the surface texture is dominated by the waxy bloom that produces the characteristic dusty appearance, with underlying skin texture varying from smooth to slightly rough depending on cultivar and growing conditions. In leaves, surface texture varies with trichome density and type, affecting light reflectance, boundary layer properties, and pest resistance. Young shoot tips display cultivar-specific surface textures that provide ampelographic identification characters captured in OIV descriptors. Surface texture assessment employs visual observation, tactile evaluation, scanning electron microscopy for fine structure, and profilometry for quantitative roughness measurement.
Surface_Texture_Quality is distinguished from Internal_Texture_Quality by addressing only the external boundary of organs. Surface texture properties may be assessed on different organ surfaces (adaxial versus abaxial leaf surfaces, exposed versus shaded berry surfaces) that can differ substantially. The cuticle and epicuticular wax layer are primary determinants of surface texture on aerial organs. OIV descriptors address surface texture through pubescence ratings for shoot tips, young leaves, and lower leaf surfaces. The relationship between surface texture and surface wettability affects spray deposition and disease development.
Außenflächencharakter
Oberflächentextur
epidermal texture
epidermale Textur
external texture
outer surface character
surface feel
äußere Beschaffenheit
Galet P (1979) A Practical Ampelography, Cornell University Press pp.: 95-110 Konlechner C and Sauer U (2016) Ultrastructural Leaf Features of Grapevine Cultivars (Vitis vinifera L. ssp. vinifera). OENO One 50(4): 195-207
Surface Texture Quality
Sauvignon Blanc vines in Marlborough, New Zealand bearing a sustainable production role under the Sustainable Winegrowing New Zealand (SWNZ) program maintain documented scorecards across 12 sustainability criteria including water usage below 1.5 ML/ha, energy consumption tracking with 15-20% reduction targets, integrated pest management with spray diaries showing 30-50% fewer applications than regional averages, and annual biodiversity assessments documenting native species restoration on 5-8% of vineyard area.
A production system role that inheres in a grapevine cultivated under a sustainability framework that systematically addresses environmental stewardship, economic viability, and social equity through documented practices and measurable indicators, without necessarily prohibiting all synthetic inputs but requiring their evidence-based minimization and responsible application.
Sustainable_Production_Role occupies a conceptual space between conventional and organic production by permitting synthetic inputs under controlled, documented conditions while requiring systematic environmental impact reduction. Unlike organic or biodynamic roles, sustainability frameworks typically employ quantitative scorecards and continuous improvement metrics rather than absolute prohibitions: this makes sustainability standards inherently more flexible but also more variable across certification programs. Major viticultural sustainability programs include SWNZ, Certified California Sustainable Winegrowing (CCSW), Haute Valeur Environnementale (HVE) in France, and WIETA in South Africa.
Nachhaltigkeits-Anbau-Rolle
environmentally responsible production role
nachhaltige Bewirtschaftungsrolle
nachhaltige Produktionsrolle
sustainability-certified production role
sustainable viticulture role
OIV Resolution CST 1/2008 on guidelines for sustainable vitiviniculture Szolnoki G (2013) A cross-national comparison of sustainability in the wine industry, Journal of Cleaner Production 53: 243-251 Flores SS (2018) What is sustainability in the wine world?, Journal of Cleaner Production 176: 1090-1104
Sustainable Production Role
In Grüner Veltliner vines in the Wachau, Austria, sustained environmental inhibition maintains quiescence from early February through mid-March, with continuous inhibition by mean temperatures of 3-7°C preventing any visible growth activity despite fully competent buds, interrupted briefly by a late February warm period with temperatures of 13-15°C that induced slight bud swelling before inhibition was re-established by return to cold conditions, and finally released by sustained March warming that accumulated sufficient growing degree days for budbreak initiation.
Sustained environmental inhibition in Vitis vinifera represents the continuous suppression of growth by unfavorable environmental conditions throughout the ecodormancy period, with inhibition persisting as long as temperature and other environmental factors remain below thresholds required for growth support, and with the sustained nature reflecting the ongoing operation of environmental limitation rather than any persistent internal regulatory state. This inhibition operates from the completion of endodormancy through budbreak, maintaining bud quiescence through the continuous inadequacy of environmental conditions for growth rather than through any active suppression mechanism within the bud. The temperature basis of sustained inhibition involves the fundamental requirement for adequate warmth to support the metabolic processes underlying growth, with sustained temperatures below approximately 10°C preventing the enzyme activities, biosynthetic reactions, and cellular processes required for cell division and expansion regardless of the growth readiness of bud tissues. The continuity of environmental inhibition depends on the consistency of unfavorable conditions, with uninterrupted cold maintaining uninterrupted quiescence, while interruptions by warm periods create temporary relief from inhibition that may trigger partial growth responses before inhibition is re-established upon return to cold conditions. The cellular state under sustained inhibition reflects the poised readiness for growth characteristic of ecodormancy, with cell cycle machinery prepared for activation, biosynthetic capacity available for upregulation, and hormone response systems sensitized for growth signal transduction, all awaiting only the relief of environmental inhibition to proceed. The gradual changes occurring under sustained inhibition include slow accumulation of growth-related gene products, progressive metabolic reactivation within temperature-imposed limits, and ultrastructural changes in meristematic cells preparing for division, representing the hidden progress toward budbreak that occurs even during apparent inactivity. The release from sustained inhibition occurs when environmental conditions improve sufficiently to support growth processes, with the transition from inhibition to active growth being determined by heat unit accumulation above the base temperature threshold reaching the level required for budbreak initiation.
Sustained_Environmental_Inhibition represents the continuous suppression aspect of Quiescence_Maintenance. Distinguished from active internal inhibition by dependence solely on environmental conditions. Temperature is the primary inhibitory factor with threshold approximately 10°C. Continuity depends on consistency of unfavorable conditions. Hidden preparatory changes occur despite apparent inactivity. Release occurs through sustained warming and heat unit accumulation. Duration varies with climate and seasonal weather patterns.
anhaltende Umwelthemmung
continuous environmental suppression
fortgesetzte thermische Inhibition
kontinuierliche externe Suppression
maintained environmental constraint
ongoing thermal inhibition
persistent external inhibition
persistierende Umweltbeschränkung
Sustained Environmental Inhibition
In Riesling vines in the Rheingau, sustained growth inhibition persists through the endodormancy period from November through early February, with bud meristems showing no mitotic activity when examined histologically despite exposure to mid-January temperatures of 12-14°C for five consecutive days, demonstrating the robustness of internal inhibitory mechanisms that prevent growth response until approximately 1200 chilling hours have accumulated and dormancy maintenance mechanisms have been sufficiently dismantled.
Sustained growth inhibition in Vitis vinifera represents the continuous suppression of meristematic activity throughout the endodormancy period, maintained by the persistent operation of multiple growth-inhibiting mechanisms that prevent cell division and elongation in bud meristems regardless of external conditions. This sustained inhibition operates from dormancy establishment through dormancy release, ensuring that the fully formed primordial shoots within dormant buds remain arrested in development during the entire winter period when growth would be maladaptive. The molecular basis of sustained growth inhibition involves the continued activity of growth-suppressing regulatory systems including dormancy-associated MADS-box transcription factors that repress genes required for cell cycle progression and meristem function, CDK inhibitor proteins that block cyclin-dependent kinase activity essential for cell division, and chromatin-based silencing mechanisms that maintain growth genes in a transcriptionally inactive state. The hormonal component of sustained growth inhibition involves the maintenance of growth-suppressive hormone balances with elevated ABA levels that directly inhibit cell cycle progression and growth gene expression, reduced cytokinin signaling that normally promotes cell division, and limited auxin transport that would otherwise activate meristematic activity. The persistence of growth inhibition through temporary warm periods during winter demonstrates the robustness of maintaining mechanisms, with endodormant buds failing to respond to temperatures that would rapidly induce growth in ecodormant or non-dormant buds, illustrating the internal rather than environmental control of growth during endodormancy. The strength of sustained growth inhibition is not constant through endodormancy but varies with dormancy depth, being strongest during deep endodormancy and progressively weakening as chilling accumulation reduces the intensity of inhibitory mechanisms, eventually declining below the threshold required to prevent growth and allowing transition to ecodormancy. Cell biological features supporting sustained growth inhibition include restricted plasmodesmatal function that limits symplastic communication between dormant meristems and surrounding tissues, potentially isolating meristems from growth-promoting signals, and the maintenance of meristem cells in G0/G1 phase of the cell cycle where they are competent to resume division when inhibition is released but remain arrested as long as inhibitory mechanisms persist.
Sustained_Growth_Inhibition represents the growth-suppression aspect of Dormancy_Maintenance specifically addressing continuous meristem arrest. Distinguished from Growth_Inhibition_Maximum which represents peak inhibition intensity, while Sustained_Growth_Inhibition addresses the persistence of inhibition over time. Involves DAM gene expression, CDK inhibitors, hormonal suppression, and chromatin silencing. Strength varies with dormancy depth through the endodormancy period. Robustness against warm period response demonstrates internal control nature of endodormancy.
anhaltende Wachstumshemmung
continuous growth suppression
fortgesetzte Meristem-Hemmung
kontinuierliche Wachstumsunterdrückung
maintained growth arrest
ongoing meristem inhibition
persistent growth inhibition
persistierende Wachstumsinhibition
Sustained Growth Inhibition
Sweet-bitter balance evaluation in Brunello di Montalcino compares young wine (bitterness 5.8, perceived sweetness 4.2, balance rating 5.5 indicating bitter dominance) with aged wine (bitterness 4.2, perceived sweetness 5.5, balance rating 7.8 indicating harmony), with the shift attributed to polymerization reducing monomeric bitter compounds while oxidative development creates subtle sweetness impression, demonstrating how the sweet-bitter balance evolution during aging transforms harsh young wine into integrated mature expression.
Sweet-bitter balance quality is a taste balance quality that represents the perceived equilibrium between sweetness and bitterness, characterizing the taste relationship particularly relevant for red wines and certain white wine styles where phenolic bitterness contributes to structure and complexity requiring integration with fruit sweetness for harmonious perception. Within the BFO framework, this quality specifically addresses the sweetness-bitterness interaction, distinguishing this balance dimension from sweet-sour and other taste relationships while recognizing the importance of sweetness in counteracting the negative hedonic impact of bitterness. The ontological significance lies in sweet-bitter balance determining whether phenolic bitterness is perceived as a positive structural element or a negative quality fault, where sufficient sweetness (from residual sugar, glycerol, or ripe fruit character) integrates bitterness into complex but harmonious taste structure, while insufficient sweetness allows bitterness to dominate creating harsh, unpleasant perception. The perceptual interaction between sweetness and bitterness involves strong mutual suppression, with sweetness being particularly effective at reducing bitter perception, explaining the traditional practice of adding sugar to bitter medications and the role of fruit ripeness in determining whether tannin bitterness is perceived positively. The compositional basis includes sugar content (residual sugar providing direct sweetness), alcohol level (ethanol contributing apparent sweetness perception), glycerol (enhancing fullness that counterbalances bitterness), and bitter compound concentration (monomeric phenolics, low-molecular-weight tannins contributing bitter taste). The developmental changes during grape ripening show improved sweet-bitter balance as sugar accumulates while extractable bitter compounds decline through seed maturation, explaining the correlation between phenolic maturity and balanced tannin perception. Understanding sweet-bitter balance quality enables optimization of harvest timing for integrated bitterness, winemaking decisions regarding extraction intensity relative to fruit sweetness, and quality assessment of red wines based on the harmony between structural bitterness and fruit character.
Sweet_Bitter_Balance_Quality addresses the interaction between sweetness and phenolic bitterness particularly important in red wine evaluation. This quality relates to but is distinct from sweet-astringent balance, as bitterness (taste) and astringency (tactile) are different sensory modalities though often correlated with similar phenolic compounds. The balance assessment requires consideration of all sweetness contributors including alcohol warmth and glycerol fullness alongside residual sugar.
Frucht-Tannin-Gleichgewicht
Süße-Bitter-Balance
Süße-Bitter-Harmonie
Zucker-Phenol-Verhältnis
fruit-tannin balance
sugar-phenolic balance
sweet-tannic harmony
sweetness-bitterness equilibrium
Keast RSJ, Breslin PAS (2003) An overview of binary taste-taste interactions. Food Quality and Preference 14(2): 111-124 Gawel R (1998) Red wine astringency: a review. Australian Journal of Grape and Wine Research 4(2): 74-95
Sweet Bitter Balance Quality
Sweet-sour balance monitoring in Chenin Blanc for Vouvray production tracks the progression from sugar/acid ratio of 12 (180 g/L sugar, 15 g/L acid) at early harvest suitable for sparkling base through ratio of 22 (210 g/L sugar, 9.5 g/L acid) at optimal dry wine harvest to ratio of 35 (280 g/L sugar, 8 g/L acid) for demi-sec style, with sensory panel confirming that the perceived balance shifts from tart-dominant through equilibrium to sweet-dominant across this range, guiding harvest timing for the three distinct wine styles produced from this single variety.
Sweet-sour balance quality is a taste balance quality that represents the perceived equilibrium between sweetness and sourness, characterizing the fundamental taste relationship that defines grape maturity perception and wine style classification from bone-dry through off-dry to overtly sweet categories. Within the BFO framework, this quality specifically addresses the primary taste axis in grape and wine evaluation, distinguishing the sweet-sour relationship from other balance dimensions while recognizing its dominant role in determining perceived ripeness, freshness, and stylistic category. The ontological significance lies in sweet-sour balance serving as the foundational taste structure upon which other quality attributes build, where this relationship determines whether wine is perceived as crisp and refreshing (acid-dominant), harmoniously balanced (equilibrium), or rich and sweet (sugar-dominant), with each position appropriate for different wine styles and food pairing contexts. The perceptual interaction between sweetness and sourness involves mutual suppression, where sugar reduces perceived acidity and acid reduces perceived sweetness, meaning the balance point depends on both component concentrations and their interaction rather than simple ratio. The compositional basis includes total sugars (glucose, fructose, residual sucrose), total acidity (primarily tartaric and malic acids), and pH which affects acid perception independent of titratable acidity, with the sugar/acid ratio providing a practical index though not perfectly predicting perceived balance due to matrix effects. The developmental trajectory during ripening shows sweet-sour balance shifting from acid-dominant in unripe fruit through balance at optimal maturity to sugar-dominant in overripe fruit, with the optimal balance point varying by intended wine style. Understanding sweet-sour balance quality enables harvest timing targeting the balance appropriate for intended style, winemaking adjustment through chapitalization, acidification, or deacidification to achieve target balance, and quality assessment based on the taste relationship most directly affecting consumer acceptance.
Sweet_Sour_Balance_Quality addresses the primary taste balance relationship in grape and wine evaluation. This quality underlies wine style classification and is the most frequently assessed balance parameter. The mutual suppression interaction between sweetness and sourness means that perceived balance cannot be directly calculated from component concentrations, requiring sensory assessment or empirically calibrated prediction models.
Süße-Säure-Harmonie
sugar-acid balance
sugar-acid harmony
sweet-tart balance
sweetness-acidity ratio
Ough CS, Amerine MA (1988) Methods for Analysis of Musts and Wines, 2nd Ed. John Wiley & Sons, New York, pp. 50-75 Jackson RS (2014) Wine Science: Principles and Applications, 4th Ed. Academic Press, San Diego, pp. 358-392
Sweet Sour Balance Quality
Sweet wine specification for Riesling Trockenbeerenauslese from the Rheingau documents the most extreme natural sweet wine style — minimum must weight 150° Oechsle (approximately 35° KMW), achievable only from individually selected, fully botrytized and desiccated berries, fermentation proceeding slowly over weeks to months to a final alcohol of 5.5-8.0% with residual sugar of 180-300+ g/L, balanced by TA >8 g/L and pH 6 g/L, and the characteristic Tokaji balance of honeyed sweetness, vibrant acidity, and oxidative complexity.
Sweet wine specification is a wine style description entity that documents the compositional, sensory, and production-method parameters defining wines with significant residual sugar achieved through natural grape sugar concentration mechanisms rather than spirit addition, encompassing late harvest wines (Spätlese, vendange tardive), botrytized wines (Sauternes, Tokaji Aszú, Beerenauslese, Trockenbeerenauslese), ice wines (Eiswein, icewine), dried grape wines (Amarone-method appassimento, vin de paille, Strohwein), and arrested fermentation sweet wines, specifying the required sugar concentration method, minimum must weight, residual sugar range, acid balance requirements, and the cultivar and climatic conditions that enable each style of natural sweet wine production.
Sweet wine specification documents the viticultural and oenological conditions required for producing wines that represent some of the most celebrated and historically significant achievements in winemaking — Tokaji Aszú was the first wine to receive a formal classification system (1730), Sauternes commands the highest per-hectare revenue in Bordeaux, and German Trockenbeerenauslese and Eiswein achieve the highest prices per bottle in global wine markets. The sugar concentration mechanisms define distinct sub-categories with profoundly different production requirements: noble rot (Botrytis cinerea infection under specific alternating humid-dry microclimate conditions, requiring cultivar susceptibility with thin skins and loose clusters — Sémillon, Riesling, Furmint, Chenin Blanc), freezing (ice wine requires sustained temperatures below -7°C during harvest, restricting production to continental climates — Niagara Peninsula, Rheingau, Neusiedlersee, Weinviertel), desiccation (appassimento/Strohwein through post-harvest grape drying on mats, racks, or in ventilated chambers — Amarone, Recioto, Vin de Paille, Austrian Strohwein), and on-vine desiccation (late harvest in dry autumn conditions — Alsace vendange tardive, Spätlese/Auslese without botrytis). The acid balance requirement is the critical quality differentiator in sweet wines — high residual sugar without proportionally high acidity produces cloying, unbalanced wines, while the combination of extreme sweetness with vibrant acidity (the hallmark of great TBA, Tokaji, and Sauternes) creates the tension and complexity that defines world-class sweet wine. The yield implications of sweet wine production are severe — noble rot and ice wine conditions reduce yields to 5-15% of normal table wine production (200-800 L/ha versus 5000-8000 L/ha for table wine), while the labor requirements for selective berry-by-berry harvest (TBA) or multiple picking passes (Sauternes tries) make these wines among the most labor-intensive agricultural products in the world.
Dessertweinspezifikation
Prädikatsweinspezifikation
Süßweinparameter
botrytized wine specification
edelsüßer Wein Spezifikation
late harvest wine specification
naturally sweet wine specification
noble sweet wine specification
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1, 2nd Ed, Wiley, pp.: 395-440 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 400-460 Magyar I (2011) Botrytized wines. Advances in Food and Nutrition Research 63: 147-206 Pigott JR et al. (2006) Eiswein. In Chemistry and Technology of Wines and Liquors, Springer, pp.: 121-145
Sweet Wine Specification
Sweetness description for Riesling Spätlese from the Mosel (2024 vintage, RS 45 g/L) records "medium-sweet" — the substantial residual sugar is balanced and partially masked by the wine's high acidity (TA 8.5 g/L), creating the characteristic Mosel sweet-acid tension where the wine tastes less sweet than its analytical sugar content would predict, with the sweetness perceived as integrated fruit sweetness rather than cloying sucrosity. Sweetness description for Napa Valley Cabernet Sauvignon (2022 vintage, RS 1.2 g/L) evaluates as "dry with sweet fruit impression" — the wine is analytically bone-dry, but the ripe fruit character at 14.6% alcohol and elevated glycerol creates a perception of sweetness that trained tasters distinguish from actual residual sugar. Sweetness description for Grüner Veltliner Smaragd from the Wachau (2024 vintage, RS 2.8 g/L) records "dry" — the residual sugar is below sensory threshold in the context of the wine's firm acidity (TA 5.8 g/L, pH 3.22), producing no perceptible sweetness.
Sweetness description is a taste description entity that records the perceived sweetness of wine on the palate, assessed on a structured scale from bone-dry (no perceptible sweetness) through dry, off-dry, medium-dry, medium-sweet, and sweet to lusciously sweet, integrating the contribution of residual sugar concentration with the modulating effects of acidity (which suppresses sweetness perception), alcohol (which enhances sweetness at moderate levels), glycerol (which contributes viscous-sweet mouthfeel), and fruit ripeness character (which can create a "sweet fruit" impression even in analytically dry wines), and providing the fundamental taste classification that positions the wine within the dry-to-sweet spectrum and establishes the framework for balance assessment.
Sweetness description in wine evaluation is considerably more nuanced than a simple correlation with residual sugar concentration, as the perceived sweetness is the emergent product of interactions between multiple chemical and perceptual factors. The sweetness perception threshold for sucrose in water is approximately 5-10 g/L, but in wine, the threshold for detecting residual sugar is typically 2-4 g/L in high-acid wines (pH 3.6), because acidity is the most powerful modulator of sweetness perception — the suppressive interaction between sweet and sour taste receptor pathways means that the same residual sugar level tastes noticeably sweeter in a low-acid wine than in a high-acid wine. This interaction forms the chemical basis of the EU labeling regulation that permits wines with up to 9 g/L residual sugar to be labeled "dry" if the total acidity is within 2 g/L of the sugar content — a regulatory acknowledgment that sweetness perception depends on the sugar-acid balance rather than sugar alone. Alcohol contributes a sweet taste through direct activation of sweet taste receptors and through enhancement of perceived body and viscosity that is cognitively associated with sweetness, explaining why high-alcohol dry wines (14-16% vol.) are frequently described as having "sweet fruit" character despite containing <2 g/L residual sugar. Glycerol (5-15 g/L in typical wines, higher in botrytized and Amarone-style wines) contributes a viscous-sweet mouthfeel that enhances overall sweetness impression, though its direct sweet taste contribution is debated — at the concentrations found in wine, glycerol's sweetness contribution is likely below the perceptual threshold of approximately 5 g/L. The fructose-to-glucose ratio in residual sugar further modulates sweetness perception, as fructose is perceived as 1.7× sweeter than glucose at equivalent concentrations, meaning that wines with fructose-enriched residual sugar (typical after extended fermentation that preferentially consumed glucose) taste sweeter per gram of measured sugar.
Restsüßewahrnehmung
Süßebeschreibung
Süßebeurteilung
Süßeeindruck
Süßewahrnehmung
palate sweetness
sugar impression
sweet-dry assessment
sweetness perception
Peynaud E, Blouin J (2006) The Taste of Wine, 4th Ed, Wiley, pp.: 75-120 Noble AC, Bursick GF (1984) The contribution of glycerol to perceived viscosity and sweetness in white wine. American Journal of Enology and Viticulture 35(2): 110-112 Prescott J et al. (2005) Psychophysical models of sweet-sour interaction in wine. Chemical Senses 30(Suppl 1): i285-i286 European Parliament (2019) Commission Delegated Regulation (EU) 2019/33, Annex III
Sweetness Description
Sweetness onset comparison between botrytized Tokaji Aszú (high viscosity, 180 g/L sugar) and ice wine Riesling (lower viscosity, 190 g/L sugar) using time-intensity methodology reveals onset time (threshold to half-maximum) of 4.2 seconds for Tokaji compared to 2.8 seconds for Riesling, with the 50% slower onset in Tokaji attributed to the higher viscosity impeding sugar diffusion to taste receptors, creating the characteristic slowly building sweetness that distinguishes botrytized wines from other dessert wine styles.
Sweetness onset quality is a sweetness quality that represents the temporal characteristic of how rapidly sweet taste perception emerges following oral contact with grape or wine, characterizing the speed of sweetness development from initial tasteless moment through threshold detection to maximum intensity perception. Within the BFO framework, this quality specifically addresses the temporal emergence dimension of sweetness, distinguishing the dynamics of perception development from peak intensity and persistence, and recognizing that rapid versus slow onset creates qualitatively different taste experiences even at equivalent maximum intensities. The ontological significance lies in sweetness onset affecting the perceived character and quality of sweet taste, where rapid onset creates an immediate sweet impact perceived as clean and direct while slow onset produces a building, evolving sweetness impression that integrates differently with other taste components developing on their own time courses. The physiological basis for sweetness onset involves receptor binding kinetics, where different sweet compounds show different association rates with the T1R2/T1R3 receptor determining how quickly the neural signal develops, combined with mass transfer factors affecting how rapidly sweet compounds reach taste receptors through the salivary film and mucus layer. The compositional factors affecting sweetness onset include sugar type (glucose showing slightly faster onset than fructose), sugar concentration (higher concentrations producing faster onset), and matrix effects where viscosity and other components affect diffusion to receptors. In wine evaluation, sweetness onset interacts with acidity perception, where rapid acid onset preceding sweetness creates a different balance impression than simultaneous or sweetness-first emergence, affecting the perceived freshness and harmony of sweet wines. Understanding sweetness onset quality enables characterization of temporal taste profiles beyond intensity alone, optimization of sugar-acid balance considering temporal dynamics, and prediction of the taste experience from compositional and physical property data.
Sweetness_Onset_Quality addresses the temporal emergence of sweetness perception, complementing Perceived_Sweetness_Intensity_Quality (magnitude) and Sweetness_Persistence_Quality (duration). This quality requires time-intensity methodology for quantitative assessment, measuring the rate of intensity increase from initial perception to maximum. The class supports characterization of temporal taste profiles and understanding of how matrix properties affect taste dynamics.
Süßeanstieg
Süßeattacke
Süßebeginn
initial sweetness perception
sweet taste emergence
sweetness attack
sweetness development speed
zeitliches Auftreten der Süße
Dijksterhuis GB, Piggott JR (2001) Dynamic methods of sensory analysis. Trends in Food Science & Technology 12(8): 284-290 Hollowood TA et al. (2002) The effect of viscosity on the perception of flavour. Chemical Senses 27(7): 583-591
Sweetness Onset Quality
Sweetness persistence evaluation in German Riesling Auslese using time-intensity analysis reveals persistence (time from maximum to threshold on decay) of 28±5 seconds for wines with 85 g/L residual sugar compared to 18±4 seconds for wines with 55 g/L sugar at equivalent acidity, with the longer persistence in the sweeter wines contributing to the fuller, more luxurious finish character distinguishing Auslese from Spätlese level, and informing winemaking decisions to target residual sugar levels achieving the persistence associated with this quality tier.
Sweetness persistence quality is a sweetness quality that represents the duration of sweet taste perception following swallowing or expectoration, characterizing the lingering sweetness that continues after the primary taste stimulus has been removed from the oral cavity. Within the BFO framework, this quality specifically addresses the temporal decay dimension of sweetness, distinguishing the duration of perception from onset dynamics and peak intensity, and recognizing that persistent sweetness fundamentally alters the overall taste experience and integration with subsequent tastes in sequential evaluation. The ontological significance lies in sweetness persistence contributing to aftertaste quality and perceived richness, where extended sweetness prolongs the positive hedonic impact of sweet wines, maintains flavor complexity during the finish, and affects pairing dynamics when food is consumed between sips. The physiological basis for sweetness persistence involves receptor binding kinetics (slow dissociation prolonging receptor activation), residual sweet compounds in saliva and oral mucosa continuing to stimulate receptors, and central neural processing where perception continues briefly after peripheral stimulation ceases. The compositional factors affecting sweetness persistence include sugar type (with some studies suggesting fructose persists longer than glucose), concentration (higher levels generally showing longer persistence), and matrix effects where compounds binding sugars in the oral cavity may extend their residence time. In wine evaluation, sweetness persistence interacts with other persistent taste qualities, particularly the astringency and bitterness that define red wine finish, where sweet persistence can balance or be overwhelmed by these sensations depending on relative durations. The correlation between sweetness persistence and overall quality perception is positive for dessert wines where lingering sweetness is desirable but potentially negative for wines intended as dry where persistent sweetness undermines the style. Understanding sweetness persistence quality enables characterization of finish quality, prediction of food pairing success based on temporal taste matching, and wine style optimization considering the full temporal trajectory of sweetness perception.
Sweetness_Persistence_Quality addresses the duration of sweetness perception after stimulus removal, completing the temporal characterization with Sweetness_Onset_Quality and Perceived_Sweetness_Intensity_Quality. This quality is particularly relevant for wine finish evaluation and food pairing considerations. Measurement requires time-intensity protocols tracking intensity decline, with persistence typically defined as time to return to threshold or to a specified percentage of maximum intensity.
Süßedauer
Süßenachklang
Süßepersistenz
anhaltende Süße
lingering sweetness
sweet aftertaste
sweetness duration
sweetness finish
Clark CC, Lawless HT (1994) Limiting response alternatives in time-intensity scaling: an examination of the halo-dumping effect. Chemical Senses 19(6): 583-594 Noble AC (1998) Why do wines taste bitter and feel astringent? In: Waterhouse AL, Ebeler SE (eds) Chemistry of Wine Flavor. ACS Symposium Series 714, American Chemical Society, Washington DC, pp. 156-165
Sweetness Persistence Quality
Sweetness quality assessment of Muscat of Alexandria table grapes correlates sensory panel sweetness ratings with instrumental sugar measurements, revealing that perceived sweetness of 7.5 on a 10-point scale requires 180 g/L total sugars at acidity of 6.5 g/L but only 160 g/L at acidity of 4.5 g/L, demonstrating the 12% reduction in sugar requirement when acidity is lowered, and enabling harvest prediction based on the sweetness-acidity interaction rather than sugar content alone.
Sweetness quality is a basic taste quality that represents the gustatory perception of sweet taste arising primarily from sugar stimulation of T1R2/T1R3 heterodimeric taste receptors, characterizing the hedonic positive taste sensation associated with caloric content that serves as the primary driver of grape palatability and a fundamental determinant of wine style and perceived quality. Within the BFO framework, this quality specifically addresses the perception of sweetness as distinct from sugar content, recognizing that sweetness perception is modulated by other taste components (acids suppress sweetness, enhancing compounds augment it), matrix effects, and individual variation in receptor sensitivity and hedonic response. The ontological significance lies in sweetness serving as the dominant positive taste attribute in table grapes and a key parameter defining wine style categories from bone-dry to lusciously sweet, where perceived sweetness results from the interaction of residual sugar with acidity, alcohol, and glycerol rather than sugar concentration alone. The compositional basis for grape sweetness includes the hexose sugars glucose and fructose in approximately equal proportions at maturity, with fructose contributing approximately 1.7 times the sweetness intensity per gram compared to glucose, meaning the glucose/fructose ratio affects perceived sweetness independently of total sugar. Sweetness perception threshold in wine ranges from 1-4 g/L depending on acidity and individual sensitivity, with wines below threshold perceived as dry despite containing measurable sugar. The developmental trajectory of sweetness follows sugar accumulation during ripening, with post-véraison phloem import building sugar concentration from approximately 50 g/L at véraison to 200-250 g/L at typical wine grape harvest, enabling progressive sweetness increase that winemakers monitor to target desired wine style. Understanding sweetness quality enables harvest timing for target sweetness perception, winemaking manipulation of sweetness-acidity balance, and prediction of consumer acceptance based on sweetness preference patterns.
Sweetness_Quality addresses the perceptual experience of sweet taste, distinguishing from Sugar_Content_Quality which measures the chemical stimulus. This quality serves as parent for intensity, onset, and persistence subclasses addressing different aspects of sweetness perception. The sweetness-acidity interaction is particularly important in wine and grape evaluation, as acidity suppresses sweetness perception and must be considered in predicting sensory outcomes from compositional data.
Süße
Süßeempfindung
Süßgeschmack
Zuckergeschmack
sugar taste perception
sweet taste
sweet taste quality
sweetness sensation
Moskowitz HR (1970) Ratio scales of sugar sweetness. Perception & Psychophysics 7(5): 315-320 Noble AC, Bursick GF (1984) The contribution of glycerol to perceived viscosity and sweetness in white wine. American Journal of Enology and Viticulture 35(2): 110-112
Sweetness Quality
A sylleptic lateral shoot on Vitis vinifera cv. Chardonnay arising from node 9 axillary bud, axillary bud initiated at this node in mid-May during primary shoot elongation, bud development progressed continuously through June without entering dormancy, lateral shoot emerged in early July approximately 7 weeks post-bud initiation representing continuous sylleptic development pattern, developed to 48cm length by late August with 7 leaves, representing botanical sylleptic classification (non-dormant immediate development from bud formation to shoot growth within single season) which in practical viticulture corresponds to prompt lateral category, this lateral demonstrating continuous development pathway contrasting with proleptic pattern where axillary buds would remain dormant through winter and potentially develop following spring creating delayed rather than immediate lateral emergence.
A lateral shoot developing without dormancy period from axillary bud during the same growing season as primary shoot that bears it, characterized by continuous development from bud formation through shoot emergence and elongation within single season, absence of winter dormancy between axillary bud formation and shoot development, botanical definition emphasizing non-dormant immediate development distinguishing from proleptic laterals requiring dormancy period, and synonymy with prompt lateral shoots in viticulture where same-season timing is practical distinguishing feature regardless of botanical dormancy considerations. Sylleptic lateral shoots represent botanical classification emphasizing developmental continuity from bud initiation through shoot growth without intervening dormancy, with this immediate same-season development contrasting with proleptic pattern (bud formation one season, dormancy period, shoot development following season), though in practical viticulture the sylleptic versus prompt distinction is often used interchangeably both referring to same-season lateral emergence.
Sylleptic lateral shoots represent botanical developmental classification emphasizing non-dormant continuous progression from axillary bud initiation through shoot development, with this pattern contrasting to proleptic development requiring dormancy period between bud formation and shoot emergence. In practical viticulture, the sylleptic category largely overlaps with prompt lateral designation, both referring to same-season lateral emergence though sylleptic emphasizes botanical developmental mechanism while prompt emphasizes practical timing observation. The sylleptic pattern (immediate non-dormant development) is typical in vigorous growth conditions, varieties prone to lateral development, or following apical dominance reduction (hedging, damage), with continuous resource availability and favorable hormonal conditions supporting immediate lateral activation rather than dormancy induction. Understanding sylleptic versus proleptic distinction has limited practical management implications since both categories are handled based on timing and vigor rather than dormancy history, though the botanical classification provides developmental context for lateral shoot biology and variety-specific lateral production patterns.
continuous development lateral
immediate-growth lateral
nicht-dormanter Achseltrieb
non-dormant lateral
sylleptischer Seitentrieb
Sylleptic Lateral Shoot
Cabernet Sauvignon hermaphrodite flowers at BBCH 59-61 show symmetrical bicarpellate pistils in approximately 85-92% of flowers, with transverse ovary sections revealing two locules of nearly equal size (left and right locules varying 90% of pistils exhibiting symmetrical bicarpellate organization, while vines under moderate nitrogen stress show increased frequency of asymmetric pistils (20-28% versus 8-12% under optimal nutrition), demonstrating environmental influence on carpel development symmetry.
A symmetrical bicarpellate pistil is a bicarpellate pistil in which the two constituent carpels show equal or nearly equal development creating symmetric ovary morphology with two locules of approximately equal size, symmetric ovule distribution between locules (equal or nearly equal ovule numbers per locule), and bilaterally symmetric stigma organization, representing the typical and most common bicarpellate pistil morphology in Vitis flowers where balanced carpel development produces structurally regular pistils. Symmetrical bicarpellate pistils are characterized by= (1) bilocular ovary with two locules of equal or nearly equal volume and shape, evident in transverse cross-sections as two chambers of similar dimensions (typically varying by 80-90% of flowers in most cultivars under typical growing conditions, reflecting coordinated and balanced development of the two carpel primordia during pistil organogenesis (BBCH 51-54). The symmetric development results from= (1) simultaneous initiation of two carpel primordia at equivalent positions on the floral meristem (2) equal resource allocation to both developing carpels during organogenesis (3) coordinated cell proliferation and differentiation in both carpels creating equal tissue elaboration (4) symmetric hormonal signaling and gene expression patterns supporting balanced carpel development. Symmetrical bicarpellate pistils show functional advantages for reproduction= symmetric ovule distribution between locules potentially increases fertilization probability by providing balanced targets for pollen tube access from the unified stigma through the bifurcating style transmitting tract symmetric locule organization may facilitate coordinated ovule development and seed maturation symmetric ovary structure provides balanced foundation for subsequent berry development with regular fruit morphology. Environmental and developmental factors can disrupt symmetry creating asymmetric bicarpellate pistils= resource limitations during critical pistil developmental stages (BBCH 52-54) may cause differential carpel development temperature stress, water stress, or nutritional imbalances may affect one carpel more than the other if stress occurs during a narrow developmental window when carpels are differentially sensitive developmental stochasticity may create minor asymmetries in carpel initiation timing or resource allocation. The degree of symmetry in bicarpellate pistils shows some cultivar-specific variation, with certain genotypes showing higher frequencies of minor asymmetries (15-25% of pistils with locules varying 20-40% in size) while maintaining functional bicarpellate organization, while other genotypes show highly consistent symmetry (>90% of pistils with locules varying <15% in size). Symmetrical bicarpellate pistils typically produce berries with symmetric seed distribution (seeds distributed equally or nearly equally between the two original locules) and regular berry morphology, though fruit set patterns and seed development can create asymmetries even from initially symmetric pistil structures through differential fertilization success or differential seed development between locules.
Symmetrical_Bicarpellate_Pistil represents the normal balanced development of the two-carpel structure and is distinguished from Asymmetrical_Bicarpellate_Pistil by equal or nearly equal carpel/locule development versus unequal development creating size differences >20%. Symmetry represents the typical developmental outcome under favorable conditions and reflects coordinated carpel organogenesis. Asymmetry can result from environmental stresses, developmental perturbations, or genetic variation affecting carpel development coordination. Understanding symmetry versus asymmetry is relevant for interpreting pistil developmental dynamics, for assessing environmental stress effects on flower development, and for understanding seed number and distribution patterns in berries. Most pistils show some degree of minor asymmetry, with the symmetrical classification encompassing pistils with locule size variation <20% representing the functional symmetry range.
ausgeglichener Zwei-Fruchtblatt-Stempel
balanced bicarpellate pistil
equal-carpel pistil
gleichmäßiges Zwei-Karpell-Pistill
symmetrischer bikarpellater Stempel
Symmetrical Bicarpellate Pistil
A symmetrical bifurcated tendril at node 6 on a vigorous shoot of Vitis vinifera cv. Chardonnay in Champagne, France, exhibiting branch point at 8 cm from shoot attachment, dividing into two arms of 11.5 cm and 12 cm length (ratio 0.96=1), with branch angles of 42 and 48 degrees from the main axis creating nearly perfect Y-symmetry, both branches showing 2.8-3.0 mm diameter at mid-length indicating equal vascular development, both exhibiting active circumnutation with 135-150 degree sweep amplitudes covering complementary spatial zones, both demonstrating rapid coiling response (within 8-10 minutes) when experimentally contacted, and both capable of supporting estimated 600-700 grams load based on structural assessment, measured at BBCH stage 71 (fruit set) in mid-June, representing the highly symmetrical bifurcation pattern characteristic of Chardonnay where 80-85% of bifurcated tendrils show arm length ratios >0.90=1.
A bifurcated tendril exhibiting equal or nearly equal development of both branches in Vitis vinifera, characterized by arm length ratio between 0.85=1 and 1=1 indicating balanced resource allocation and developmental timing between the two branches, branch angles typically 35-55 degrees from the main axis creating a well-balanced Y-shaped architecture with arms diverging symmetrically, equal or nearly equal diameter of both branches reflecting similar vascular development and structural investment, and comparable functional capabilities in both arms including similar circumnutation amplitude, coiling response vigor, and load-bearing capacity. Symmetrical bifurcated tendrils develop when lateral meristems at the branch point are activated simultaneously or near-simultaneously during early tendril development, receive equal resource allocation throughout the growth period, and experience similar environmental conditions without differential stress, damage, or positional effects creating developmental asymmetry. This branching pattern represents optimal morphological expression of bifurcated architecture, maximizing the functional benefits of dual searching arms through equivalent exploration capacity in both branches, providing redundancy in attachment opportunities with either branch capable of primary support function, and creating mechanically balanced structures that distribute loads evenly when both branches achieve attachment. Symmetrical bifurcation shows cultivar-specific frequency variation, with some varieties producing predominantly symmetrical bifurcated tendrils (>75% symmetrical among bifurcated tendrils) while others show lower symmetry frequencies (<50% symmetrical) reflecting genetic variation in developmental regulation of branch meristem activity and resource partitioning.
Symmetrical bifurcated tendrils are distinguished from asymmetrical bifurcated tendrils by arm length ratio >0.85=1, balanced branch angles, and equivalent structural and functional development. The threshold ratio of 0.85=1 represents a practical distinction allowing for minor developmental variation while capturing meaningful symmetry in resource allocation and functional capacity. Symmetry assessment should consider multiple parameters beyond length including diameter, circumnutation amplitude, and coiling response, as length similarity alone may mask functional asymmetry from differential vascular development or structural reinforcement. Environmental stress during development can reduce symmetry frequency even in cultivars with genetic tendencies toward symmetrical branching, with water deficit, nutrient limitation, or differential pest damage commonly creating asymmetric development. Symmetrical bifurcation provides maximum functional efficiency among bifurcated architectures through equivalent capability in both arms.
ausgewogene Y-förmige Kletterorgan
balanced forked tendril
equal bifurcated tendril
even Y-shaped tendril
gleichmäßig verzweigte Ranke
symmetrisch gegabelte Ranke
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 90-104 This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 119-136.
Symmetrical Bifurcated Tendril
A symmetrical bilateral permanent arm on Vitis vinifera cv. Sauvignon Blanc, featuring left arm at 84cm and right arm at 82cm (2cm difference, approximately 2.4% variation well within symmetrical tolerance), each arm bearing 14 two-bud spurs spaced 12cm apart, trunk precisely centered on trellis wire equidistant from neighbor vines, balanced vigor producing similar shoot growth from both arms (left average 145cm, right average 142cm shoot length), crop distribution 52% left arm versus 48% right arm (4.8kg total, essentially balanced), representing ideal symmetrical bilateral architecture achieved through careful trunk training, centered positioning, and balanced annual management maintaining architectural uniformity.
A bilateral permanent arm configuration with closely matched dimensions and fruiting capacity between the two arms, characterized by equal or nearly equal arm lengths (typically within 10-15cm), similar spur counts and spacing on both arms, balanced crop distribution approaching 50=50 split between left and right sides, and deliberate architectural uniformity reflecting centered trunk positioning, balanced vine vigor, and management priority for symmetrical canopy development. Symmetrical bilateral permanent arms represent the ideal outcome in bilateral cordon establishment, achieved when trunks are precisely centered on trellis wires, vines show balanced vigor without directional preferences, establishment training successfully develops equal arm extensions, and annual maintenance pruning preserves symmetrical architecture through balanced spur renewal and shoot management.
Symmetrical bilateral permanent arms represent aesthetic and functional ideals in cordon viticulture, with perfect symmetry indicating precise establishment, balanced vine development, and skilled management. However, perfect symmetry is neither always achievable nor always necessary, with minor asymmetries (10-20% arm length differences, 45=55 crop splits) functionally acceptable and sometimes deliberate to accommodate off-center trunk positions, row-end constraints, or vine vigor variations. The pursuit of symmetry serves practical purposes beyond aesthetics, including simplified mechanization (symmetrical canopies present consistent targets for equipment), predictable vine behavior (balanced arms show similar phenology and fruit development), and easier manual operations (workers develop consistent expectations for symmetrical vines).
ausgeglichene Doppelarme
balanced bilateral arms
equal paired arms
symmetric cordon
symmetrische bilaterale Arme
Symmetrical Bilateral Permanent Arm
Leaf blade symmetry assessment in Sangiovese reveals characteristic mild bilateral asymmetry where the right lobe (viewed from adaxial surface with petiole toward observer) consistently exceeds the left lobe in area by 3-8% across diverse Italian growing regions, a subtle but reproducible asymmetry pattern that contributes to cultivar identification when combined with other ampelographic characters and reflects the consistent developmental program establishing leaf form in this ancient Tuscan variety.
Symmetry_Quality encompasses all morphological qualities that characterize the regularity and correspondence of form between parts of a grapevine organ arranged around an axis, plane, or central point. Symmetry represents a fundamental geometric property that describes how organ form relates to transformations including reflection, rotation, and translation. In botanical contexts, symmetry assessment addresses whether an organ displays equivalent form when divided along specified planes or rotated around axes, providing insight into developmental processes and serving as a diagnostic character for cultivar identification. This category comprises three primary conditions: Bilateral_Symmetry_Quality describing mirror-image correspondence across a single plane Radial_Symmetry_Quality capturing rotational equivalence around a central axis and Asymmetry_Quality representing the absence of regular symmetry relationships. Grapevine organs exhibit varying degrees and types of symmetry depending on their developmental origin and functional constraints. Leaves display approximate bilateral symmetry around the midrib axis, though subtle asymmetries are common and can be cultivar-specific. Berries typically exhibit radial symmetry around the polar axis when seed number and distribution are uniform, but commonly show bilateral or irregular symmetry when seed development is asymmetric. Clusters rarely display true symmetry due to the stochastic nature of fruit set across branches. Symmetry assessment ranges from qualitative visual evaluation to quantitative morphometric analysis using landmark-based methods or continuous outline analysis. Deviations from symmetry, termed fluctuating asymmetry, can indicate developmental stress and serve as indicators of environmental conditions during organ formation.
Symmetry_Quality is positioned as a direct subclass of Shape_Quality, capturing the regularity aspects of form distinct from overall shape categories and margin characteristics. Symmetry assessment requires specification of the axis, plane, or point of reference and the transformation (reflection, rotation) being evaluated. Perfect symmetry is rare in biological organs symmetry classifications represent predominant patterns with tolerance for minor deviations. PATO_0000965 (symmetry) provides the upper-ontology alignment. Fluctuating asymmetry studies use symmetry deviation as a measure of developmental stability under environmental stress.
Formregularität
Symmetrieeigenschaft
Symmetriequalität
form regularity
geometric correspondence
geometrische Entsprechung
regularity of form
symmetry character
Chitwood DH et al. (2014) A Modern Ampelography: A Genetic Basis for Leaf Shape and Venation Patterning in Grape. Plant Physiology 164(1): 259-272 Palmer AR and Strobeck C (1986) Fluctuating Asymmetry: Measurement, Analysis, Patterns. Annual Review of Ecology and Systematics 17(1): 391-421
Symmetry Quality
Symptom suppression quality analysis in grapevine leafroll disease compared foliar symptom expression to viral titer across cultivars to identify genotypes with enhanced tolerance to GLRaV-3 infection. Quantitative PCR determination of GLRaV-3 titer in petiole tissue established equivalent viral loads (within 2-fold) across the test cultivars, while visual scoring of leafroll symptoms assessed the symptom response to this standardized infection level. Grenache demonstrated striking symptom suppression quality with viral titers equivalent to severely symptomatic Merlot yet showing only trace interveinal reddening scored at 0.8 on a 0-5 scale versus 4.2 for Merlot with equivalent viral load. Chardonnay showed intermediate symptom suppression quality with symptom score of 2.1 at equivalent titer. The symptom suppression quality differential translated to fruit quality impacts, with symptom-suppressed Grenache showing only 0.8°Brix reduction compared to virus-free controls versus 2.5°Brix reduction in highly symptomatic Merlot at equivalent viral load, suggesting that the physiological tolerance extended beyond visual symptoms to metabolic function. Investigation of symptom suppression quality mechanisms revealed that Grenache maintained higher photosynthetic rates in infected leaves (85% of healthy controls versus 55% in Merlot), suggesting tolerance at the level of photosynthetic apparatus sensitivity to viral disruption. The symptom suppression quality characterization identified cultivar differences in leafroll tolerance independent of viral replication, informing recommendations that in regions with endemic leafroll pressure, cultivars with symptom suppression capacity may maintain productivity without the intensive roguing programs required for highly symptomatic cultivars.
Symptom suppression quality is a disease tolerance quality that inheres in a grapevine organism and represents the capacity to limit the visible manifestations of pathogen infection disproportionate to actual pathogen load, showing reduced disease symptoms despite levels of pathogen colonization that would cause severe symptoms in symptomatic cultivars, with symptom suppression potentially operating through limitation of pathogen toxin effects, reduced plant sensitivity to infection-induced signals, or maintenance of tissue function despite pathogen presence. This quality captures the phenomenon where infection level and symptom expression are decoupled, recognizing that the relationship between pathogen quantity and disease symptoms varies among host genotypes and that some cultivars effectively suppress visible damage through mechanisms distinct from limiting pathogen growth. The ontological significance of symptom suppression quality lies in its practical value for maintaining marketable crop despite infection, since fruit appearance and vine health ratings depend on symptom expression rather than pathogen load, and in its research value for understanding the mechanistic basis of disease symptoms that symptom suppression genotypes help to dissect. The biological basis of symptom suppression encompasses reduced sensitivity to pathogen-produced toxins or cell-wall-degrading enzymes that cause visible damage, attenuated activation of plant responses including senescence-associated pathways that may be triggered by infection, and maintenance of cellular integrity and function through enhanced structural or biochemical properties that resist pathogen-induced damage. The assessment of symptom suppression requires quantification of both pathogen presence (through PCR, ELISA, or microscopic observation) and symptom severity, with symptom suppression identified when pathogen levels equivalent to those causing severe symptoms in other cultivars produce only mild or moderate symptoms in the putatively tolerant cultivar. The distinction between symptom suppression and asymptomatic infection is important, with true symptom suppression representing active tolerance mechanisms while asymptomatic infection might simply reflect low pathogen load due to undetected resistance mechanisms.
Symptom_Suppression_Quality is_a Disease_Tolerance_Quality, representing the capacity to limit visible symptoms disproportionate to pathogen load. This class addresses the decoupling of infection level from symptom expression. Assessment requires parallel quantification of pathogen presence and symptom severity. Mechanisms may include toxin insensitivity, reduced senescence activation, or maintained cellular function. The Grenache/Merlot differential for leafroll exemplifies cultivar variation in symptom tolerance. Distinguished from asymptomatic infection which might reflect low pathogen load rather than true tolerance.
Schadensbegrenzung
Symptomtoleranz
Symptomunterdrückung
reduced symptom expression
reduzierte Symptomexpression
symptom tolerance
symptom-pathogen decoupling
visible damage suppression
Pagán I, García-Arenal F (2018) Tolerance to plant pathogens: theory and experimental evidence. International Journal of Molecular Sciences 19:810 Naidu RA et al. (2015) Grapevine leafroll disease and associated viruses: a unique pathosystem. Annual Review of Phytopathology 53:613-634 Almeida RPP et al. (2013) Ecology and management of grapevine leafroll disease. Frontiers in Microbiology 4:94
Symptom Suppression Quality
Synchronization quality assessment comparing clonal versus heritage vineyard blocks reveals the genetic contribution to developmental coordination, with clonal Chardonnay (single clone, uniform rootstock) showing high synchrony (coefficient of variation for véraison date among vines 3.2%, single harvest capturing 95% of fruit at optimal maturity), while heritage mixed planting (multiple Pinot Noir clones) shows lower synchrony (CV 8.7%, requiring either selective harvest or acceptance of maturity heterogeneity), demonstrating how genetic uniformity affects synchronization with implications for management efficiency and fruit homogeneity.
Synchronization quality is a phenological quality that represents the temporal coordination of developmental events across different organs within a plant or among different plants within a population, characterizing the uniformity of phenological timing that affects management efficiency, fruit homogeneity, and the practical feasibility of treating vineyard units as coherent production blocks requiring uniform operations. Within the BFO framework, this quality addresses the coordination dimension of phenological development, distinguishing the synchrony among developmental events from the timing or rate of individual events while recognizing that synchronization determines whether a vine or vineyard can be effectively managed as a unit with single operations or requires multiple passes to accommodate developmental heterogeneity. The ontological significance lies in synchronization quality determining both the efficiency of vineyard management and the uniformity of fruit quality, where high synchrony enables single-pass operations (one spray timing, one harvest date) and produces homogeneous fruit populations (uniform maturity at harvest), while low synchrony requires multiple interventions to accommodate spread developmental stages and produces heterogeneous fruit that may compromise wine quality through mixed maturity. The levels of synchronization assessment include intra-plant synchrony (coordination among organs within a single vine, such as uniformity of bud break across different positions or uniformity of berry development within and among clusters) and inter-plant synchrony (coordination among different vines within a block, determining whether the block can be treated as a management unit), with both levels contributing to overall vineyard uniformity. The factors affecting synchronization include genetic uniformity (clonal plantings showing higher synchrony than mixed plantings), environmental uniformity (consistent soil and microclimate promoting uniform development), vine condition uniformity (balanced vines developing more synchronously than stressed or excessively vigorous vines), and management uniformity (consistent pruning, irrigation, and nutrition promoting coordinated development). The practical implications of synchronization encompass spray timing (synchronized flowering enabling single fungicide application at optimal stage), harvest logistics (synchronized ripening enabling single harvest at optimal maturity), and fruit sorting requirements (high synchrony reducing need for selection to eliminate developmentally aberrant fruit). Understanding synchronization quality enables assessment of vineyard uniformity affecting management efficiency, identification of factors disrupting synchrony that could be corrected, and realistic planning for operations based on the actual developmental coordination of the vineyard.
Synchronization_Quality serves as the parent class for intra-plant and inter-plant synchrony dimensions. This quality addresses the temporal coordination of development affecting management efficiency and fruit uniformity. The class supports vineyard uniformity assessment through synchronization characterization at multiple organizational levels.
Entwicklungskoordination
Synchronisationsqualität
developmental coordination
developmental homogeneity
phenological uniformity
phänologische Uniformität
temporal synchrony
zeitliche Synchronie
Dry PR, Coombe BG (2004) Viticulture Volume 1: Resources, 2nd Ed. Winetitles, Adelaide, pp. 150-200 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 59-144
Synchronization Quality
Chardonnay in Champagne shows synchronous cap fall during mid-June Year N+1 at BBCH 63-65 during peak flowering, with discrete coordinated detachment events involving 8-14 simultaneous cap sheds per cluster occurring during morning hours 9am-11am, creating visible waves of calyptra fall under cool maritime conditions of 19-24°C temperature, 58-72% relative humidity, reflecting good developmental uniformity in this early-bursting moderately-to-highly synchronous cultivar supporting uniform 62-74% fruit set
Synchronous cap fall represents the coordinated simultaneous calyptra detachment phenomenon during peak flowering when numerous flowers throughout cluster complete abscission within same brief period creating concentrated cap fall events, reflecting developmental synchronization and optimal environmental conditions enabling coordinated flowering. This remarkable coordination phenomenon occurs during peak flowering in mid-June in temperate regions Year N+1, typically visible as discrete events within peak cap shedding period when multiple flowers simultaneously shed caps creating visible waves of detachment, corresponding to BBCH 63-65 stages. Synchronization characteristics show multiple flowers completing cap detachment within minutes to hours of each other creating concentrated activity bursts, spatial coordination with flowers across different cluster regions shedding simultaneously despite physical separation, and temporal clustering of detachment events suggesting common triggering factors. Coordinated detachment timing shows characteristic pattern with discrete shedding waves particularly during optimal morning conditions, individual events involving 5-15 simultaneous detachments per cluster creating visible sudden cap fall, and repeated synchronized events throughout peak period maintaining intensive flowering activity. Developmental synchronization underlying coordinated fall reflects uniform flower maturation resulting from compressed primordium formation previous season, similar developmental progression through inflorescence emergence and pre-anthesis stages, and convergence toward simultaneous flowering readiness enabling coordinated reproductive activity. Environmental triggering of synchronized fall shows possible role for diurnal temperature changes with morning warming potentially triggering coordinated abscission completion, humidity decline facilitating simultaneous mechanical separation as tissue hydration changes, and integrated environmental signals coordinating detachment timing across multiple flowers. Temperature threshold effects may contribute to synchronization with coordinated abscission completion when temperature exceeds critical value around 20-22°C, morning temperature rise potentially triggering simultaneous detachment in flowers approaching readiness, and daily temperature pattern creating repeated synchronization opportunities. Humidity effects on synchronization show that declining morning humidity potentially facilitates coordinated mechanical separation as caps loosen with reduced turgor, creating condition enabling simultaneous detachment in multiple flowers reaching critical stage, and contributing to characteristic morning peak in synchronized shedding activity. Mechanical coordination shows that individual cap detachments potentially triggering neighboring detachments through vibration or physical contact, creating cascade effects producing concentrated shedding bursts, though importance of mechanical triggering versus independent coordination remains unclear. Hormonal coordination possibilities include potential role for ethylene as volatile signal enabling communication between flowers, though evidence for hormonal coordination of grapevine flowering remains limited, and most synchronization likely resulting from developmental uniformity and common environmental responses rather than active signaling. Observational characteristics enable recognition of synchronous fall through concentrated cap detachment events visible during observation periods, sudden appearance of multiple freshly detached caps on cluster surfaces, and characteristic waves of activity during favorable morning conditions distinguishable from continuous sporadic shedding. Visual impact of synchronous fall creates particularly striking full bloom appearance with simultaneous flower opening throughout cluster, abundant coordinated cap fall creating impressive display of reproductive activity, and overall flowering intensity enhanced by synchronized coordination. Pollination efficiency implications show synchronized fall potentially beneficial through coordinated pollen release and stigma receptivity creating optimal pollination window, concentrated reproductive activity potentially improving pollination success through enhanced pollen-stigma encounters, and synchronized flowering supporting uniform fruit set contributing to crop uniformity. Fruit set uniformity benefits from synchronization include coordinated successful pollinations creating berries with similar development timing, reduced within-cluster developmental heterogeneity potentially improving ripening uniformity, and overall quality benefits from concentrated successful reproduction. Developmental consequences show that synchronized flowering reflecting compressed developmental progression, with high synchrony indicating good developmental uniformity resulting from optimal previous season primordium formation and current season development, while asynchronous extended flowering suggesting developmental heterogeneity potentially creating ripening challenges. Weather dependency of synchronization shows that favorable stable conditions during flowering period promoting coordination through consistent environmental conditions enabling uniform progression, while variable conditions disrupting synchronization creating extended irregular flowering affecting uniformity. Cultivar differences in synchronization capacity show some varieties exhibiting strong synchronized shedding with highly coordinated flowering creating compressed bloom period, while others showing more asynchronous extended shedding reflecting lower developmental uniformity, affecting flowering characteristics and fruit set patterns. Coordination mechanisms remain incompletely understood with developmental uniformity clearly important, environmental factors contributing through common responses, and possible role for communication signals though evidence limited in grapevine. The process represents second aspect of mass cap fall emphasizing coordination rather than just intensity, occurring within peak cap shedding period. Duration represents repeated discrete events rather than continuous period, with individual synchronized detachments occurring over minutes to hours and multiple events throughout peak period. Regional occurrence shows synchronous cap fall during peak flowering period across all climates whenever developmental uniformity and favorable conditions align, though expression varies with cultivar characteristics and environmental conditions. Viticultural significance includes reflecting good developmental uniformity indicating potential for uniform fruit set and ripening, demonstrating favorable flowering conditions supporting optimal reproductive success, and providing impressive visual confirmation of flowering peak. Research applications include analyzing mechanisms underlying synchronized flowering including developmental, environmental, and possible signaling factors, evaluating relationships between flowering synchrony and fruit set uniformity, quantifying cultivar differences in coordination capacity affecting flowering characteristics, assessing environmental effects on synchronization affecting reproductive outcomes, and investigating developmental controls producing uniform flower maturation enabling coordinated anthesis.
Synchronous_Cap_Fall is second aspect of Mass_Cap_Fall, distinguished by coordination emphasis rather than just intensity. Represents discrete coordinated events within peak period. Reflects developmental uniformity and favorable conditions. Benefits pollination efficiency and fruit set uniformity. Mechanisms incompletely understood involving developmental coordination and environmental responses. Provides striking visual confirmation of flowering peak. Cultivar variation in synchronization capacity affects flowering characteristics.
Massenabwurf-Synchronisation
coordinated cap fall
gleichzeitiger Kappenabwurf
koordinierte Kalyptralösung
simultaneous cap shedding
synchroner Kappenfall
synchronized calyptra detachment
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 375-380 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 285-292
Synchronous Cap Fall
Chardonnay in Champagne shows synergid penetration during late June-early July Year N+1 at BBCH 65-69, with pollen tubes penetrating through thin nucellus tissue entering synergid cells at micropylar pole 24-40 hours post-pollination, tube apex establishing cytoplasmic connection with embryo sac, penetrated synergid initiating degeneration, under cool maritime conditions of 20-25°C temperature, adequate boron 11-17 ppm supporting final tube integrity, typical of this early-bursting cultivar with 58-72% penetration success enabling fertilization
Synergid penetration represents the embryo sac entry phase when pollen tube completes micropyle passage reaching nucellus surface, penetrates through nucellus wall, and enters synergid cell cytoplasm creating direct access to female gametophyte enabling sperm cell discharge for fertilization. This critical entry phase occurs during pollen tube penetration in mid-late June in temperate regions Year N+1, typically occurring 22-42 hours after pollination as final tube growth culminates, corresponding to BBCH 65-69 fertilization stages. Penetration mechanism involves tube apex reaching nucellus surface at inner micropyle opening adjacent to embryo sac, penetrating nucellus wall cells through enzymatic degradation and mechanical force, entering synergid cell through filiform apparatus region, and establishing direct cytoplasmic connection between tube and female gametophyte enabling sperm delivery. Nucellus structure at penetration shows thin nucellus tissue layer 1-3 cell layers thick surrounding embryo sac, cells forming protective barrier around megagametophyte, and wall tissue requiring tube penetration for embryo sac access. Nucellus penetration process shows tube apex contacting nucellus surface, possible secretion of wall-modifying enzymes degrading cell wall components, turgor pressure contributing mechanical force driving penetration, and breakthrough into embryo sac region creating access pathway. Synergid cell position shows two synergid cells flanking egg cell at micropylar pole of embryo sac, cells positioned directly adjacent to inner micropyle opening receiving arriving tubes, and one synergid typically receiving tube penetration while other remains intact. Filiform apparatus structure shows elaborate cell wall ingrowths creating extensive membrane surface area, projections extending toward micropyle creating reception zone for arriving tube, and specialized architecture facilitating tube attraction and penetration. Tube entry into synergid shows apex penetrating through filiform apparatus region entering synergid cytoplasm, penetration possibly involving enzymatic wall modification at entry point, and tube establishing connection with synergid cell interior preparing for discharge. Synergid degeneration response shows penetrated synergid cell initiating programmed cell death following tube entry, degeneration creating space for sperm cell release and movement, membrane breakdown facilitating sperm access to egg and central cell, and degeneration as normal part of fertilization process rather than pathological response. Tube positioning in synergid shows apex extending into synergid cytoplasm, sperm cells positioned near tube tip ready for release, and tube-synergid interface providing discharge site enabling sperm delivery into embryo sac. Tube-embryo sac contact interface shows membrane interactions between tube and synergid, possible adhesion molecules mediating attachment, and structural changes preparing for tube rupture and discharge. Penetration timing shows entry into synergid occurring 22-42 hours after initial pollination reflecting cumulative time for tube growth journey, 1-4 hours after micropyle arrival as final penetration completes, and positioning immediately before discharge typically within minutes to hours of synergid entry. Tube structural state at penetration shows maintenance of wall integrity throughout micropyle passage, possible wall weakening at apex preparing for discharge, and cytoplasmic organization with sperm cells positioned for release. Sperm cell positioning shows two sperm cells traveling in tube cytoplasm near apex, cells ready for delivery upon discharge, and one sperm destined for egg fusion while other targets central cell in double fertilization. Synergid signal termination following penetration possibly includes cessation of attractant production preventing additional tube recruitment, signaling change communicating occupied state, and protection against polyspermy through single tube reception per ovule. Penetrated versus intact synergid shows asymmetric reception with one synergid receiving tube and degenerating, other synergid remaining intact though function following fertilization unclear, and structural evidence distinguishing penetrated from unpenetrated cells. Environmental requirements for successful penetration include adequate temperature 22-28°C supporting metabolic processes and tube growth, continued boron requirement for tube wall integrity during final growth and penetration, and appropriate tissue hydration maintaining cell structure and function. Temperature effects on penetration show optimal conditions supporting efficient entry within 1-2 hours of micropyle arrival, cool conditions potentially delaying penetration, and temperature sensitivity affecting final fertilization success rate. Boron critical role emphasizes continued absolute requirement with boron essential for tube wall integrity during nucellus penetration and synergid entry, deficiency causing tube bursting or structural failure preventing successful embryo sac access, and adequate boron 10-20 ppm supporting normal penetration and fertilization. Failed penetration causes include tube structural failure from boron deficiency or other dysfunction, abnormal nucellus or synergid structure preventing entry, inadequate tube vigor preventing final penetration, or developmental abnormalities in embryo sac preventing normal reception. Penetration success rate shows 50-70% tubes reaching micropyle successfully penetrating synergid under favorable conditions, cumulative effect of all previous reproductive processes determining final success, and success rate directly determining fertilization potential and fruit set. The process represents second and final aspect of pollen tube penetration completing embryo sac access, following micropyle targeting. Duration typically brief representing final moments of tube growth immediately before discharge. Cultivar variation in synergid penetration may include embryo sac structure affecting entry pathway, synergid development affecting reception capacity, and penetration efficiency affecting fertilization success rate. Regional timing shows synergid penetration occurring during mid-late June across all climates 22-42 hours after pollination as culmination of tube growth journey. Viticultural significance includes representing final access phase enabling sperm delivery to female gametophyte, requiring successful completion of entire pollination-germination-growth sequence, demonstrating critical importance of tube structural integrity, and establishing penetration success as immediate prerequisite for fertilization and fruit set. Research applications include analyzing synergid structure and function in tube reception, investigating penetration mechanisms and tube-synergid interactions, evaluating factors affecting penetration success, quantifying penetration timing relative to pollination, and assessing penetration success as final determinant of fertilization potential.
Synergid_Penetration is second and final aspect of Pollen_Tube_Penetration, distinguished by embryo sac entry completing access to female gametophyte. Represents culmination of entire tube growth journey. Occurs 22-42 hours post-pollination. Penetrated synergid degenerates as normal response. Immediately precedes tube discharge. Critical boron dependence for successful entry. Success rate 50-70% determining fertilization potential. Enables Ovule_Penetration phase with egg approach and discharge.
Embryosack-Eintritt
Schlauch-Synergiden-Fusion
Synergiden-Penetration
embryo sac entry
female gametophyte access
synergid cell penetration
tube-synergid fusion
weibliche Gametophyten-Zugang
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 391-395 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 296-300 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Synergid Penetration
The cultivar with VIVC prime name Merlot Noir (VIVC number 07657) carries 58 documented synonyms including Merlau (archaic Bordelais), Merlot Nero (Italian adaptation), Merlô (Brazilian Portuguese), Bigney (historical Saint-Émilion usage), Plant Médoc (Libournais traditional name), Vitraille (Lot-et-Garonne regional name), and Sémillon Rouge (erroneous synonym based on misidentification in 19th-century Chilean vineyards), each representing a distinct synonym category while referring to the identical SSR-verified genotype.
A cultivar designation that refers to the same grapevine genotype as an established prime name but constitutes an alternative naming expression arising from historical usage, regional tradition, linguistic adaptation, orthographic variation, taxonomic error, or commercial convention, formally documented as equivalent to the prime name within nomenclatural reference systems.
Synonym is the most populated subclass of Cultivar_Designation, reflecting the extraordinary nomenclatural complexity of grapevine cultivars accumulated over millennia of cultivation across linguistically diverse regions. The VIVC documents over 18,000 synonyms linked to approximately 6,000-8,000 prime names. Synonymy arises through multiple independent mechanisms — historical naming before standardized nomenclature, regional isolation creating parallel naming traditions, linguistic adaptation across language boundaries, orthographic instability, and taxonomic misidentification: subclasses capture these distinct mechanisms of synonym generation. The formal resolution of synonymy through molecular identification (SSR profiling) since the 1990s has been one of the most transformative advances in grapevine taxonomy.
Sortensynonym
Synonym
alternative Sortenbezeichnung
alternative cultivar name
cultivar synonym
variety synonym
Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157 This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 OIV (2013) OIV List of Vine Varieties and Their Synonyms
Investigation of Systemic_Acquired_Resistance_Disposition at INRAE Bordeaux employed a spatially separated inoculation system to assess SAR induction and efficacy across 15 cultivars. Primary leaves were inoculated with Plasmopara viticola to trigger local defense and potential SAR, while challenge inoculation was applied to distal leaves 5 days later. SAR establishment was quantified as percent reduction in P. viticola sporulation on challenge leaves relative to non-primed controls. Cultivars varied dramatically in Systemic_Acquired_Resistance_Disposition. The PIWI cultivar 'Regent' showed strong SAR with 72% reduction in sporulation on systemically primed leaves, accompanied by 4-fold higher basal PR-1 expression in distal tissues and accelerated stilbene accumulation upon challenge (peak at 18 hpi versus 36 hpi in non-primed). Susceptible 'Chasselas' exhibited weak SAR (18% sporulation reduction) with minimal PR gene priming in systemic tissues. Salicylic acid measurements confirmed the SA-dependence of the response, with 'Regent' showing 3.5-fold SA elevation in systemic leaves following primary infection compared to 1.4-fold in 'Chasselas'. Exogenous SA application (1 mM foliar spray) induced SAR-like protection even in 'Chasselas' (55% sporulation reduction), suggesting that weak Systemic_Acquired_Resistance_Disposition in susceptible cultivars reflects deficient SA signaling rather than inability to respond to the signal. Duration experiments revealed that 'Regent' maintained significant SAR protection for 14 days post-primary infection, while 'Chasselas' SAR decayed within 7 days.
A Systemic_Acquired_Resistance_Disposition is an Induced_Defense_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to establish whole-plant enhanced resistance following localized pathogen infection, providing broad-spectrum protection against subsequent attacks by diverse pathogens in tissues distant from the initial infection site. This disposition represents one of the most important systemic defense mechanisms in plants, enabling a single infection event to prime the entire plant for enhanced defense responses lasting days to weeks. Systemic acquired resistance (SAR) is triggered by local pathogen recognition and hypersensitive response, generating mobile signals—primarily salicylic acid (SA) and the methylated derivative methyl salicylate, along with lipid-derived signals including azelaic acid and glycerol-3-phosphate—that travel through phloem to distal tissues. In systemic tissues, these signals activate the NPR1 (Nonexpressor of PR genes 1) regulatory pathway, leading to chromatin remodeling, transcription factor activation, and expression of pathogenesis-related (PR) genes including PR-1, PR-2, and PR-5. The resulting "primed" state enables faster and stronger defense responses upon subsequent pathogen challenge, with primed tissues showing earlier PR gene induction, more rapid phytoalexin accumulation, and enhanced hypersensitive response compared to naive tissues. Systemic_Acquired_Resistance_Disposition varies among grapevine genotypes in both the strength of SAR establishment following primary infection and the duration of the primed state, with resistant cultivars often showing more robust and persistent SAR than susceptible cultivars.
Systemic_Acquired_Resistance_Disposition is_a Induced_Defense_Disposition. SAR is triggered by pathogen infection and mediated by salicylic acid signaling, distinct from Induced_Systemic_Resistance_Disposition which is triggered by beneficial microbes and mediated by jasmonate/ethylene signaling. SAR provides broad-spectrum resistance effective against biotrophic and hemibiotrophic pathogens but may be less effective against necrotrophs. The primed state established by SAR represents a form of plant "immune memory" that can persist for extended periods. SAR can be artificially induced by SA analogs (e.g., BTH/Acibenzolar-S-methyl), providing a crop protection strategy that exploits this natural defense pathway.
SAR capacity
SAR-Kapazität
Salicylsäure-abhängige-Systemresistenz
Systemisch-erworbene-Resistenz-Disposition
salicylic acid-dependent resistance
systemic priming
Durrant WE, Dong X (2004) Systemic acquired resistance, Annual Review of Phytopathology 42:185-209 Fu ZQ, Dong X (2013) Systemic acquired resistance: turning local infection into global defense, Annual Review of Plant Biology 64:839-863 Harm A et al. (2011) Evaluation of chemical and natural resistance inducers against downy mildew (Plasmopara viticola) in grapevine, American Journal of Enology and Viticulture 62:184-192
Systemic Acquired Resistance Disposition
Systemic acquired resistance quality evaluation in Vitis vinifera Chardonnay tested the efficacy of benzothiadiazole (BTH) treatment for enhancing resistance to powdery mildew under vineyard conditions. BTH application at 0.3 mM to leaves 7 days before Erysiphe necator inoculation induced systemic acquired resistance quality detectable as 35-fold elevation of PR-1 gene expression in untreated leaves distant from application sites, confirming systemic signal transmission. The systemic acquired resistance quality-enhanced plants showed 65% reduction in powdery mildew disease severity compared to water-treated controls at 14 days post-inoculation, with the protection extending to newly expanded leaves that emerged after BTH treatment, demonstrating the persistence of the primed state. Gene expression analysis revealed systemic acquired resistance quality priming with defense genes showing faster and stronger induction upon pathogen challenge in BTH-treated plants as VvPR1 reached 120-fold induction by 12 hours in primed plants versus 35-fold in naive controls, and stilbene synthase (VvSTS) showed 85-fold versus 40-fold induction. The systemic acquired resistance quality response showed duration of approximately 3 weeks, with protection declining thereafter unless additional BTH applications were made. Field trials integrating SAR induction with reduced fungicide programs achieved equivalent disease control to calendar-based spray schedules using 40% fewer fungicide applications, demonstrating practical utility of systemic acquired resistance quality enhancement for integrated disease management in commercial viticulture.
Systemic acquired resistance quality is a defense response quality that inheres in a grapevine organism and represents the capacity to develop enhanced whole-plant resistance following localized pathogen infection or elicitor treatment, with the systemic acquired resistance (SAR) state characterized by elevated expression of defense genes throughout the plant including uninfected tissues, priming of defense responses for more rapid and stronger activation upon subsequent pathogen challenge, and broad-spectrum resistance effective against diverse pathogen types beyond the inducing agent. This quality captures the systemic and persistent dimension of induced defense, recognizing that local infection or treatment can trigger lasting changes in plant immune status that protect the entire plant for extended periods, providing a form of "plant vaccination" that enhances resistance through immune system priming rather than direct pathogen killing. The ontological significance of systemic acquired resistance quality lies in its potential for practical disease management through the use of SAR-inducing agents that enhance natural resistance without fungicide application, and in the role of SAR capacity as a component of quantitative resistance that varies among cultivars and contributes to field resistance. The biological basis of SAR encompasses the generation of mobile signals at the site of primary infection that travel through the phloem to distant tissues, the perception of these signals triggering transcriptional reprogramming that establishes the primed state, and the epigenetic modifications including histone methylation changes that maintain the primed state over extended periods. The signaling molecules involved in SAR include salicylic acid as a critical local and potentially systemic signal, pipecolic acid and its derivative N-hydroxy-pipecolic acid as mobile signals generated during SAR induction, and various proposed signals including methyl salicylate, azelaic acid, and glycerol-3-phosphate whose relative contributions remain under investigation. The practical induction of SAR in grapevine can be achieved through application of chemical inducers including benzothiadiazole (BTH), β-aminobutyric acid (BABA), and various biological elicitors, with efficacy varying by cultivar, environmental conditions, and the specific inducer employed.
Systemic_Acquired_Resistance_Quality is_a Defense_Response_Quality, representing the whole-plant enhanced resistance state following localized infection or elicitor treatment. This class addresses the systemic and persistent "plant vaccination" dimension of immunity. Salicylic acid and pipecolic acid derivatives serve as mobile signals. Priming accelerates and amplifies subsequent defense responses. BTH (benzothiadiazole) and BABA are practical SAR inducers. Duration typically 2-4 weeks requiring repeated applications. Provides broad-spectrum protection beyond the inducing agent. Integration with reduced fungicide programs offers IPM applications.
Pflanzimmunität-Priming
SAR
Systemisch Erworbene Resistenz
induced systemic resistance
plant immunity priming
systemic resistance
systemische Resistenz
Fu ZQ, Dong X (2013) Systemic acquired resistance: turning local infection into global defense. Annual Review of Plant Biology 64:839-863 Dufour MC et al. (2013) The effects of plant defense inducers on Botrytis cinerea diseases in grapevine. Pest Management Science 69:994-1001 Harm A et al. (2011) Evaluation of chemical and biological elicitors for controlling gray mold of grapevine and apple. Crop Protection 30:1303-1307
Systemic Acquired Resistance Quality
A T-bud of Cabernet Sauvignon inserted into actively growing 110 Richter rootstock in a California nursery bears a T_Bud_Role, performed in June when bark slips readily, with the shield-shaped bud piece slid under the T-cut bark flaps and wrapped with budding rubber—the bud remaining dormant until spring push the following year. A T-bud of Sangiovese serves in a T_Bud_Role for Italian nursery production, where field T-budding of established rootstock liners remains a traditional practice producing grafted vines over a two-year cycle. A T-bud of Grüner Veltliner on established vineyard rootstock bears a T_Bud_Role in Austrian top-working operations, converting existing plantings to higher-value cultivars while preserving mature root systems.
A field graft role that inheres in a grafted combination assembled using the T-budding (shield budding) technique, where a shield-shaped piece of bark containing a single scion bud is inserted under a T-shaped bark incision on the rootstock, exploiting the bark slip condition when bark separates readily from the cambium during active growth. T-budding is a traditional field budding technique performed during the growing season (typically late spring through late summer) when active cambial growth causes the bark to "slip" or separate easily from underlying wood, allowing insertion of the bud shield beneath the bark flaps created by the T-cut. The technique involves cutting a T-shaped incision in the rootstock bark (vertical cut 2-3 cm, horizontal cut 1-1.5 cm), lifting the bark flaps, inserting a shield-shaped bud piece cut from scion budwood, and wrapping with budding tape to secure the bud in position. T-budding is widely used for field grafting onto established rootstocks in nursery rows or for top-working vineyard vines, with the established root system supporting rapid bud development once the graft union forms.
T_Bud_Role captures the traditional field budding technique that dominated grapevine propagation before benchgrafting development and remains important for field grafting applications. The technique's dependence on bark slip restricts its timing window to active growth periods (typically 4-8 weeks in mid-summer), contrasting with chip budding's extended season flexibility. T-budding requires less precise matching than chip budding (the bud shield slides under existing bark rather than fitting into a cut notch) but depends critically on the bark slip condition—attempts during dormancy or drought stress when bark adheres tightly to cambium result in failure. The technique produces strong unions with experienced operators achieving 75-85% success rates, though results are more variable than controlled-environment benchgrafting. T-budding's continued importance reflects its suitability for field operations, low equipment requirements, and integration with established two-year nursery production systems.
Augenokulation
Schildokulation
T-Okulation
T-bud
T-budding graft
shield bud
shield budding
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 440-445 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Pongrácz DP (1983) Rootstocks for Grape-vines, David Philip Publishers Whiting JR (2004) Grapevine rootstocks, Winetitles pp.: 167-188
T Bud Role
Thompson Seedless (Sultana) bears a Table_Grape_Scion_Role as the world's most versatile grape, consumed fresh, dried as raisins, and used for juice, with its neutral flavor, seedless berries (2-4 g), and crisp texture defining the commercial table grape benchmark in global markets. Crimson Seedless serves in a Table_Grape_Scion_Role as the leading red seedless variety, with firm, crunchy texture, mild sweetness, and 8+ week cold storage capability enabling year-round global supply from alternating Northern and Southern Hemisphere production. Red Globe bears a Table_Grape_Scion_Role as the dominant large-berry seeded variety for Asian markets, with berries exceeding 10-15 g commanding premium prices in China, Japan, and Korea where size is valued over seedlessness.
A scion role that inheres in a grafted scion cultivar by virtue of its designation for table grape production, where grapes are grown for fresh consumption rather than wine, juice, or raisin production. Table grape scions are selected for consumer-oriented quality attributes fundamentally different from wine grapes, including large berry size (typically 4-12 g/berry versus 1-2 g for wine grapes), attractive appearance (uniform color, bloom retention, cluster architecture), crisp or fleshy texture (firm, crunchy flesh preferred over soft wine-grape texture), neutral-to-pleasant flavor without excessive acidity or astringency, long post-harvest shelf life (2-8 weeks under cold storage), and shipping durability for global supply chains. Seedlessness has become the dominant consumer preference in major markets, though seeded varieties remain important regionally. Table grape production requires intensive canopy management (berry sizing, cluster thinning, gibberellin applications) and careful harvest timing to optimize appearance and eating quality rather than the sugar-acid-phenolic balance prioritized for wine. Global table grape production exceeds 25 million tonnes annually, concentrated in China, Turkey, India, Iran, and the Americas.
Table_Grape_Scion_Role represents the second-largest grape utilization after wine (approximately 35% of global production), with distinct cultivar requirements and viticultural practices. The market has shifted dramatically toward seedlessness since the 1990s—seedless varieties now dominate North American and European consumption, though seeded giants (Red Globe, Italia) remain preferred in Asian markets. Table grape breeding is among the most active in horticulture, with major programs (USDA, IFG, SNFL, Grapa) releasing dozens of proprietary varieties annually, protected by plant patents and managed through licensed grower networks. Climate requirements differ from wine grapes table grape production concentrates in warm, dry regions (California, Chile, Peru, South Africa, Australia, Mediterranean) where disease pressure is manageable without fungicide residues.
Frischtraube
Tafeltraube
dessert grape
fresh grape
table grape
Creasy GL, Creasy LL (2009) Grapes, CABI Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 430-480 Christensen LP (2000) Raisin Production Manual, University of California ANR OIV (2019) Statistical Report on World Vitiviniculture
Table Grape Scion Role
Tactile astringency quality assessment in young Barossa Shiraz reveals substantial drying (7/10) and moderate puckering (5/10) from tannin concentration of 1850 mg/L, with the pronounced tactile grip creating the structural framework that supports the wine's concentrated dark fruit and spice flavors, while also indicating 8-12 year aging potential during which polymerization will soften the tactile sensation to the refined, integrated astringency (drying 4/10, puckering 2/10) characteristic of mature examples of this style.
Tactile astringency quality is a tactile quality that represents the mechanosensory dimension of astringent perception arising from polyphenol-protein interaction in the oral cavity, characterizing the drying, puckering, and roughening sensations that contribute to wine structure and mouthfeel independent of the taste or aromatic dimensions of phenolic compound perception. Within the BFO framework, this quality specifically addresses the tactile component of astringency distinct from its potential gustatory (bitterness) and olfactory contributions, recognizing that astringency is fundamentally a touch sensation mediated by reduced oral lubrication and increased surface friction rather than a taste or smell. The ontological significance lies in tactile astringency providing the structural backbone of red wine mouthfeel, where the drying and gripping sensations create the textural framework that supports flavor complexity, enables food pairing through palate-cleansing action, and contributes to age-worthiness through tannin evolution potential, making astringency assessment essential for red wine quality evaluation. The physiological basis for tactile astringency involves polyphenol binding to proline-rich salivary proteins causing their aggregation and precipitation, which reduces oral lubrication and creates friction between oral surfaces, with mechanoreceptors detecting this friction increase as the characteristic drying and puckering sensation that intensifies with repeated exposure during wine consumption. The compositional determinants of tactile astringency include tannin concentration (higher levels increasing astringency intensity), tannin molecular structure (smaller, more reactive tannins creating harsher astringency), tannin-protein binding affinity (varying with polymerization state and galloylation), and modulating factors (polysaccharides, alcohol, and pH affecting perceived astringency at equivalent tannin levels). The developmental trajectory of tactile astringency shows typical softening with wine aging as tannins polymerize to sizes with reduced protein-binding capacity and eventually precipitate, creating the mellowing that distinguishes mature from young red wines. Understanding tactile astringency quality enables red wine structure assessment, aging potential prediction from astringency character, and quality evaluation incorporating appropriate astringency level for wine style.
Tactile_Astringency_Quality serves as the parent class for specific astringent tactile sensations (Drying, Puckering), establishing the mechanosensory nature of astringency distinct from taste perception. This quality is the tactile complement to Astringency_Quality defined in the Sensory_Quality hierarchy, specifically addressing the touch-mediated dimension. The class supports red wine structure assessment and aging potential prediction through tactile astringency evaluation.
astringent mouthfeel
oral drying
tactile grip
taktile Adstringenz
taktiler Griff
Breslin PAS et al. (1993) Cellular basis of taste perception. Physiology & Behavior 54(6): 1129-1134 Gawel R (1998) Red wine astringency: a review. Australian Journal of Grape and Wine Research 4(2): 74-95
Tactile Astringency Quality
Tactile quality assessment in comprehensive wine evaluation integrates multiple somatosensory dimensions including body (full, substantial weight), viscosity (silky, coating texture), temperature (appropriately cool for style), astringency (firm but refined tannin grip), and alcohol warmth (subtle heat in finish), with these tactile attributes providing the structural framework that supports and contextualizes the fruit, oak, and terroir flavors detected through chemosensory channels, demonstrating how tactile quality contributes essential structure to overall wine perception.
Tactile quality is a sensory quality that represents the somatosensory perception of wine and grape tissue through mechanoreceptors and thermoreceptors, encompassing touch-mediated sensations including texture, temperature, body, and astringency that complement gustatory and olfactory assessment in comprehensive sensory evaluation. Within the BFO framework, this quality addresses the touch-mediated dimension of sensory perception, distinguishing mechanically and thermally detected attributes from chemically detected taste and smell while recognizing that tactile perception integrates with chemosensory modalities to create the unified flavor experience. The ontological significance lies in tactile quality providing the structural and textural framework for wine perception, where body creates the weight impression, viscosity contributes smoothness or oiliness, astringency provides grip and structure, and temperature affects both direct thermal sensation and the release of volatile compounds, together defining the physical presence of wine in mouth that shapes overall quality impression. The physiological basis for tactile perception involves diverse receptor types including mechanoreceptors detecting pressure and texture (Merkel cells, Meissner corpuscles, Ruffini endings), proprioceptors sensing oral movement resistance, thermoreceptors detecting temperature, and free nerve endings responding to chemical irritation (contributing to the tactile dimension of astringency and alcohol warmth). The compositional determinants of tactile quality include alcohol content (affecting viscosity and warmth), glycerol and residual sugar (contributing to body and smoothness), phenolic compounds (creating astringency through protein precipitation), dissolved gases (producing effervescent texture), and serving temperature (affecting both direct thermal sensation and perception of other attributes). The integration of tactile with chemosensory perception creates emergent properties, as the physical presence of wine modulates taste and aroma perception while chemical components (tannins, alcohol, acids) contribute to tactile sensations, demonstrating the multimodal nature of wine flavor where tactile quality provides essential structural context. Understanding tactile quality enables comprehensive sensory assessment incorporating texture and body, quality evaluation based on tactile balance and integration, and appreciation of how physical properties complement chemical composition in wine quality perception.
Tactile_Quality serves as the parent class for touch-mediated sensory qualities, establishing the somatosensory dimension of wine evaluation distinct from chemosensory (taste, smell) perception. This quality encompasses mechanoreception (pressure, texture), thermoreception (temperature), and the tactile aspects of chemesthetic sensations (astringency, alcohol warmth). The class supports comprehensive sensory assessment that integrates tactile with chemosensory qualities for complete wine evaluation.
Berührungsempfindung
Tastsinn
haptic quality
haptische Wahrnehmung
physical mouthfeel
somatosensory quality
taktile Qualität
touch sensation
Pickering GJ, Demiglio P (2008) The white wine mouthfeel wheel. Journal of Wine Research 19(2): 51-64 Green BG (1993) Oral astringency: a tactile component of flavor. Acta Psychologica 84(1): 119-125
Tactile Quality
Tannin concentration in grape berries ranges from 1-6 mg/berry across cultivars, with skins contributing 20-40% and seeds 60-80% of total berry tannin in most genotypes. Cultivar differences span 3-4 fold, with high-tannin types such as Tannat (4-6 mg/berry) and Nebbiolo (3-5 mg/berry) contrasting with low-tannin cultivars such as Gamay (1-2 mg/berry) and Pinot Noir (1.5-2.5 mg/berry). Proanthocyanidin biosynthesis involves leucoanthocyanidin reductase (VvLAR) producing catechin units and anthocyanidin reductase (VvANR) producing epicatechin units, with polymerization extending chains to 3-80+ subunits. Tannin accumulation in skins begins pre-veraison, peaks at veraison, then remains stable, while seed tannin accumulates earlier and declines post-veraison through oxidative polymerization.
A phenolic compound disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate proanthocyanidins (condensed tannins) in grape berry skins and seeds through the flavonoid pathway, with genotype-specific total concentration, polymer size distribution, and subunit composition determining wine astringency, bitterness, mouthfeel, color stability, and aging potential.
Tannin_Accumulation_Disposition serves as the parent class for specific tannin location, polymerization, and extractability traits. Tannins are essential for red wine structure, aging capacity, and color stabilization through anthocyanin-tannin polymerization. The trait shows moderate heritability (h² = 0.50-0.70) with significant environmental modulation by water status, light exposure, and temperature. Winemaking practices (maceration time, temperature, pressing) strongly affect tannin extraction from the genetically determined berry pool.
Gerbstoff-Disposition
PA accumulation disposition
Proanthocyanidin-Bildung
Tanninakkumulation
condensed tannin trait
proanthocyanidin synthesis
Kennedy JA et al. (2001) Composition of grape skin proanthocyanidins at different stages of berry development. Journal of Agricultural and Food Chemistry 49:5348-5355 Bogs J et al. (2005) Proanthocyanidin synthesis and expression of genes encoding leucoanthocyanidin reductase and anthocyanidin reductase in developing grape berries and grapevine leaves. Plant Physiology 139:652-663
Tannin Accumulation Disposition
In Mourvèdre berries in Bandol, Provence during BBCH 85-89, tannin chain elongation increases skin tannin mean degree of polymerization from 26 to 38 over 35 days through continued condensation of epicatechin extension units, with phloroglucinolysis revealing extension unit composition of 72% epicatechin, 18% epigallocatechin, and 10% epicatechin-gallate, while terminal units show 65% epicatechin reflecting preferential chain initiation from this monomer under Mediterranean conditions averaging 27°C.
Tannin chain elongation describes the progressive increase in proanthocyanidin polymer length in Vitis vinifera berry tissues through continued addition of flavan-3-ol extension units to existing oligomeric and polymeric chains, wherein interflavonoid bond formation adds catechin, epicatechin, or epigallocatechin monomers to chain termini or internal positions generating the progressively larger tannin polymers that characterize advancing phenolic maturation in skin tissues during the post-veraison ripening period. This process involves the formation of C4-C8 or C4-C6 interflavonoid bonds through acid-catalyzed condensation, with carbocation intermediates generated at the C4 position of flavan-3-ol units attacking nucleophilic C6 or C8 positions of other units, creating the B-type procyanidin linkages that constitute the predominant bond type in grape proanthocyanidins. The dynamics of tannin chain elongation show tissue-specific patterns reflecting different biosynthetic and modification environments, with skin tannins demonstrating potential for continued elongation during post-veraison ripening as newly synthesized monomers become available for incorporation into existing polymers, while seed tannins show limited elongation after veraison when active proanthocyanidin biosynthesis largely ceases and subsequent modifications primarily involve oxidative reactions rather than chain extension. The quantitative assessment of chain elongation employs analytical methods including phloroglucinolysis and thiolysis that cleave interflavonoid bonds generating terminal and extension unit fragments whose ratio determines mean degree of polymerization, with chain elongation detected as increasing mDP values over time that distinguish active elongation from stable polymer populations. Substrate availability influences chain elongation rates through the requirement for continued flavan-3-ol monomer synthesis or release from other sources, with elongation potential limited when biosynthesis declines during late ripening or when monomers are preferentially diverted to other metabolic fates including glycosylation or oxidation rather than polymerization. The sensory significance of chain elongation relates to astringency properties, with longer tannin chains generally producing more intense but potentially softer astringency due to enhanced protein precipitation capacity combined with modified binding kinetics that influence the temporal development of astringent sensation in the mouth. Environmental factors including temperature and oxidative conditions modulate chain elongation through effects on reaction kinetics and competing processes, with moderate conditions favoring controlled elongation while oxidative stress may promote alternative modification pathways that reduce elongation efficiency.
Tannin_Chain_Elongation is subclass of Tannin_Polymerization representing polymer length increase through unit addition. Distinguishes from Condensation_Reactions which describes bonding mechanism. Sibling to Condensation_Reactions and Tannin_Structure_Maturation. Primarily active in skin tissues post-veraison. Assessed by mDP increase through phloroglucinolysis. Influences astringency intensity and quality.
Proanthocyanidin-Ketten-Extension
Tannin-Ketten-Verlängerung
Tannin-Längen-Zunahme
Tannin-Polymer-Wachstum
polymer elongation
proanthocyanidin chain extension
tannin length increase
tannin polymer growth
Kennedy JA and Jones GP (2001) Analysis of proanthocyanidin cleavage products following acid-catalysis in the presence of excess phloroglucinol. Journal of Agricultural and Food Chemistry 49: 1740-1746 Downey MO et al (2003) Analysis of tannins in seeds and skins of Shiraz. Australian Journal of Grape and Wine Research 9: 15-27 Souquet JM et al (1996) Polymeric proanthocyanidins from grape skins. Phytochemistry 43: 509-512
Tannin Chain Elongation
Tannin content for Cabernet Sauvignon from Margaret River (2023 vintage) records skin tannin of 1.4 mg/g and seed tannin of 4.2 mg/g berry fresh weight by methylcellulose precipitable tannin (MCP) assay, with the seed-to-skin tannin ratio of 3.0 indicating that maceration management must balance seed tannin extraction avoidance against adequate skin tannin release — informing a protocol of extended pre-fermentation cold soak (5 days at 8°C for skin extraction) with early pressing (10 days total skin contact) to minimize harsh seed tannin in the finished wine. Tannin content for Pinot Noir from Central Otago (2024 vintage) measures total tannin of 1.8 mg/g (skin 0.9 mg/g, seed 2.2 mg/g), with the characteristically lower skin tannin of Pinot Noir requiring gentle whole-cluster inclusion (25-30%) to supplement structural tannin from stem-derived proanthocyanidins.
Tannin content is a phenolic data entity that records the concentration of condensed tannins (proanthocyanidins) in grape skins, seeds, and must, determined by protein precipitation assay (bovine serum albumin or methyl cellulose precipitable tannin), vanillin assay, phloroglucinolysis-HPLC, or Adams-Harbertson precipitation method, expressed in milligrams per gram of berry tissue (mg/g), milligrams per liter of must or wine (mg/L), or milligrams catechin equivalents per liter (mg CE/L), and representing the polymeric flavanol fraction that determines wine astringency, mouthfeel structure, aging capacity, and the interaction with anthocyanins that governs long-term color stability through copigmentation and pigmented tannin formation.
Tannin content is arguably the most complex phenolic parameter to measure and interpret, as proanthocyanidins encompass a heterogeneous family of polymeric flavanols ranging from dimers (degree of polymerization DP 2) to polymers exceeding DP 40, with each size fraction contributing differently to sensory perception — small oligomers (DP 2-5) contribute bitterness, while larger polymers (DP > 10) contribute astringency through salivary protein precipitation. The distinction between skin tannin and seed tannin is viticulturally and oenologically critical: skin tannins are predominantly composed of epicatechin and epigallocatechin subunits with higher mean DP (10-40), producing rounder, softer mouthfeel, while seed tannins are smaller (mean DP 5-15), richer in gallocatechin gallate, and perceived as more astringent and bitter. During ripening, total seed tannin content decreases (due to oxidative browning and reduced extractability from lignifying seed coats) while skin tannin undergoes progressive polymerization — the optimal harvest window for tannin maturity occurs when seed tannin extractability is declining and skin tannin polymerization has advanced sufficiently to produce supple rather than aggressive astringency. Multiple analytical methods coexist, each measuring slightly different tannin fractions — the MCP assay (widely used in Australia) precipitates tannin with methylcellulose polymer, the Adams-Harbertson assay (dominant in North America) uses protein precipitation with BSA, and phloroglucinolysis-HPLC provides the most detailed structural information including mean DP, subunit composition, and percentage galloylation. The lack of inter-method concordance complicates cross-regional comparison of tannin data and necessitates method specification in all reported values.
Gerbstoffanalyse
Gerbstoffgehalt
Tanningehalt
Tanninkonzentration
condensed tannin
grape tannin data
tannin analysis
tannin concentration
Harbertson JF et al. (2003) A method for the estimation of proanthocyanidin chain length. American Journal of Enology and Viticulture 54(4): 301-306 Downey MO et al. (2003) Analysis of tannins in seeds and skins of Shiraz grapes throughout berry development. Australian Journal of Grape and Wine Research 9(1): 15-27 Kennedy JA et al. (2001) Changes in grape seed polyphenols during fruit ripening. Phytochemistry 55(1): 77-85 Mercurio MD et al. (2007) High throughput analysis of red wine and grape phenolics — adaptation and validation of methyl cellulose precipitable tannin assay and modified Somers color assay to a rapid 96 well plate format. Journal of Agricultural and Food Chemistry 55(12): 4651-4657
Tannin Content
Tannin description for Cabernet Sauvignon from Rutherford, Napa Valley (2021 vintage, 3 years post-harvest) evaluates as "high intensity, fine-grained, ripe" — the tannins are abundant and firmly structured but composed of predominantly polymerized, high-molecular-weight proanthocyanidins from well-ripened skins, producing a dense but smooth astringency that coats the palate without the harsh drying or bitter character of unripe seed tannin, with excellent integration suggesting optimal drinking from 8-20 years post-vintage. Tannin description for Pinot Noir from Volnay (2023 vintage) records "medium-low intensity, silky, fully resolved" — the characteristically modest tannin content of Pinot Noir is expressed as a delicate, satin-textured astringency that barely dries the palate, with complete integration achieved within 18 months of bottling. Tannin description for Blaufränkisch Reserve from Mittelburgenland DAC (2022 vintage) assesses as "medium-plus intensity, fine-to-medium grain, ripe with residual youthful grip" — the well-extracted tannins from 14-day maceration show the cultivar's characteristically firm but not aggressive structural profile, with sufficient concentration for 8-12 year aging.
Tannin description is a taste description entity that records the perceived astringency, bitterness, and textural characteristics contributed by phenolic compounds on the wine palate, assessed through multiple qualitative dimensions including intensity (low to high astringency), grain or texture (silky, fine-grained, velvety, grainy, coarse, harsh, aggressive), ripeness (green/unripe versus ripe/resolved), drying quality (the progressive moisture-depleting sensation across oral mucosa), integration (whether tannin is seamlessly incorporated or stands apart from the wine's fruit and acid structure), and evolution stage (youthful grip, developing resolution, mature integration, fading), providing the textural-structural assessment that is the primary determinant of red wine quality perception, aging potential, and optimal drinking window.
Tannin description captures the somatosensory dimension of wine evaluation that is unique to red wines (and to a lesser extent orange wines and some skin-contact whites), involving the complex interaction between phenolic compounds and the oral mucosa that produces the sensation of astringency — a tactile perception mediated by the precipitation of salivary proline-rich proteins by tannin polymers, reducing oral lubrication and creating the characteristic drying, puckering sensation. The qualitative vocabulary of tannin description — silky, velvety, fine-grained, chalky, grainy, coarse, harsh — reflects the relationship between proanthocyanidin molecular characteristics and sensory perception. High-molecular-weight polymers (mean degree of polymerization >10, predominantly from grape skins) produce a perception described as "fine-grained" or "silky" because they form large, soft precipitates with salivary proteins, while lower-molecular-weight oligomers (DP 2-6, predominantly from seeds) interact more aggressively with oral proteins, producing "grainy," "coarse," or "harsh" sensations and contributing more bitterness. The concept of "tannin ripeness" — the progression from green, astringent, bitter tannin in unripe berries to soft, supple, integrated tannin in fully mature fruit — reflects the oxidative polymerization of seed coat tannins and the progressive softening of skin tannins during ripening, providing the textural quality information that correlates with the Seed_Weight_Data and Tannin_Content classes documented in the berry composition hierarchy. Tannin perception evolves substantially during wine aging — young tannins are typically more astringent and bitter due to reactive monomeric and oligomeric species, while aged tannins are progressively polymerized, less reactive with salivary proteins, and perceived as softer and more integrated. The interaction between tannin and acid perception is synergistic — low-pH wines perceive more astringent at equivalent tannin concentrations, while high-alcohol wines perceive slightly less astringent due to ethanol's disruption of tannin-protein interactions, meaning that tannin description must always be interpreted in the context of the wine's complete compositional profile.
Adstringenzbeurteilung
Gerbstoffbeschreibung
Tanninbeschreibung
Tanninqualität
astringency description
phenolic mouthfeel
phenolisches Mundgefühl
tannin assessment
tannin quality
tannin texture description
Gawel R (1998) Red wine astringency: a review. Australian Journal of Grape and Wine Research 4(2): 74-95 Vidal S et al. (2004) Taste and mouth-feel properties of different types of tannin-like polyphenolic compounds and anthocyanins in wine. Analytica Chimica Acta 513(1): 57-65 Peynaud E, Blouin J (2006) The Taste of Wine, 4th Ed, Wiley, pp.: 155-200 Casassa LF, Harbertson JF (2014) Extraction, evolution, and sensory impact of phenolic compounds during red wine maceration. Annual Review of Food Science and Technology 5: 83-109
Tannin Description
Tannin extractability varies 2-3 fold among cultivars under equivalent maceration conditions, with high-extractability types such as Syrah and Mourvèdre releasing 50-70% of berry tannin versus low-extractability cultivars like Pinot Noir releasing 25-40%. Extractability increases during ripening as pectin degradation and cell wall loosening reduce tannin-polysaccharide binding, with skin tannin extractability rising from 20-30% at veraison to 50-70% at maturity. Seed tannin extractability conversely decreases during ripening from 60-80% at veraison to 15-30% at harvest as seed coat lignification and oxidative polymerization reduce permeability. The tannin-to-anthocyanin extraction ratio ranges from 1:1 to 3:1 depending on cultivar and maceration conditions, affecting wine balance between color and structure.
A tannin accumulation disposition that is the inherited capacity of a Vitis vinifera plant to produce grape berry tannins with genotype-specific ease of extraction into wine during maceration, determined by cell wall composition, tannin-cell wall binding strength, and the developmental changes in extractability that occur during ripening.
Tannin extractability determines the relationship between berry tannin concentration and wine tannin content, with high-extractability cultivars producing more tannic wines from equivalent berry material. The trait shows genotype × ripeness × winemaking interaction, as extraction conditions (temperature, alcohol, maceration time) interact with inherent extractability. Understanding cultivar-specific extractability informs maceration management, enabling targeted extraction of skin versus seed tannins. Berry cell wall composition, particularly pectin and hemicellulose content, affects extractability through tannin binding and physical barrier effects.
Extraktions-Disposition
Gerbstoff-Freisetzung
PA extractability trait
Tanninextrahierbarkeit
tannin accessibility disposition
tannin release capacity
Bindon KA et al. (2010) Interaction between grape-derived proanthocyanidins and cell wall material. 1. Effect on proanthocyanidin composition and molecular mass. Journal of Agricultural and Food Chemistry 58:2520-2528 Hernández-Hierro JM et al. (2014) Comparative study of the relationship between skin cell wall composition and anthocyanin extractability in Vitis vinifera. Food Chemistry 158:506-513
Tannin Extractability Disposition
Tannin extractability quality assessment in Saint-Émilion Merlot guides harvest timing and winemaking protocols, with the ITV extractability index comparing tannins extracted at wine pH (3.2-3.6) versus at pH 1.0 (complete extraction) to provide an extractability percentage optimal phenolic maturity (extractability index above 60% for skins, below 30% for seeds) indicates fruit where gentle processing will extract predominantly skin tannins producing supple, velvety texture, while premature harvest (lower skin extractability, higher seed extractability) requires more aggressive extraction that risks harsh, green tannin character.
Tannin_Extractability_Quality represents the chemical quality characterizing the ease and extent to which tannins can be released from grape tissues during winemaking, capturing the functional property that links grape tannin content to wine tannin composition. Extractability encompasses the physical and chemical factors governing tannin release from cell compartments, including cell wall porosity, tannin binding to polysaccharides and proteins, and the solubility of tannin polymers in the evolving must and wine matrix. Tannin extractability is not directly predictable from tannin concentration grapes with equivalent tannin content may yield wines with very different tannin levels depending on the accessibility of tannins to extraction solvents. The factors affecting extractability change during ripening, with berry softening, cell wall modification, and reduced tannin-protein binding generally increasing extractability as maturity advances. Seed tannin extractability follows a different trajectory than skin tannin extractability seeds retain intact structure longer than softening pulp, with seed tannin extraction accelerating only when ethanol from fermentation provides sufficient solvent capacity. Temperature, ethanol concentration, pH, and maceration duration all affect extraction extent, with winemakers manipulating these parameters to achieve target tannin extraction levels. The concept of phenolic maturity or tannin maturity captures the ripeness condition where skin tannins are readily extractable while seed tannins remain protected, enabling preferential extraction of the softer skin tannin fraction. Assessment of extractability employs model extraction systems or comparison of grape tannin content with wine tannin yield under standardized conditions.
Tannin_Extractability_Quality addresses the functional property linking grape tannin potential to wine tannin realization, representing a disposition-like quality distinct from compositional qualities. The extractability concept recognizes that winemaking is not simply tannin transfer but involves selective extraction governed by accessibility factors. Multiple extractability indices exist (ITV method, simulated maceration, cell maturity index), each with different predictive value for wine composition. Extractability changes during ripening and differs between seed and skin compartments, requiring tissue-specific assessment for optimal harvest timing decisions.
Tanninfreisetzungspotential
extractable tannin quality
extrahierbare Tanninqualität
phenolic extractability
phenolische Extrahierbarkeit
tannin accessibility
tannin release potential
Saint-Cricq de Gaulejac N et al. (1998) Determination of Anthocyanins in Grapes Using a Standard Addition Method. Journal International des Sciences de la Vigne et du Vin 32(4): 205-213 Glories Y and Augustin M (1988) Maturité Phénolique du Raisin, Conséquences Technologiques: Application aux Millésimes 1991 et 1992. Compte Rendu Colloque Journée Technique CIVB, Bordeaux pp.: 56-61
Tannin Extractability Quality
Tannin maturity assessment through sensory and analytical methods in Tempranillo reveals the polymerization-driven quality evolution, with véraison +3 weeks showing high total extractable tannins (4.2 mg/g berry) but low mDP (3.5), high seed contribution (58%), and harsh/bitter sensory profile, véraison +5 weeks showing moderate total (3.4 mg/g) with increased mDP (5.2), reduced seed contribution (42%), and firmer but less harsh sensory character, and véraison +7 weeks showing lower total (2.8 mg/g) but highest mDP (7.1), low seed contribution (28%), and soft, integrated sensory profile, demonstrating how tannin maturity assessment enables harvest timing for target mouthfeel characteristics rather than maximum tannin extraction.
Tannin maturity quality is a phenolic maturity quality that represents the developmental state of proanthocyanidin (condensed tannin) compounds in grape skins and seeds, characterizing the accumulation, polymerization degree, and sensory evolution of these structure-determining compounds that provide wine astringency, body, and aging potential while contributing to color stability through anthocyanin-tannin reactions. Within the BFO framework, this quality specifically addresses the tannin component of phenolic maturity, distinguishing proanthocyanidin development from anthocyanin or flavonol maturation while recognizing that tannin maturity involves changes in both quantity and quality, with the sensory character of tannins (harsh versus soft, bitter versus integrated) depending on molecular size, composition, and interactions that evolve during ripening. The ontological significance lies in tannin maturity determining the structural character and mouthfeel of red wines, where total tannin concentration affects body and astringency intensity, the mean degree of polymerization (mDP) influences astringency perception (larger polymers perceived as softer), the proportion of galloylation affects both bitterness and astringency, and the ratio of skin to seed tannins affects the overall tannin character (skin tannins generally perceived as finer than seed tannins). The tissue-specific accumulation patterns show skin tannins synthesized primarily before véraison then declining slightly through dilution during ripening, while seed tannins accumulate through véraison then decline as polymerization and oxidation reduce extractability, with the shift in the skin/seed tannin ratio during ripening being a key aspect of tannin maturation that affects wine tannin character. The polymerization changes during tannin maturation include progressive condensation of monomeric and oligomeric procyanidins into larger polymers, oxidative modification (particularly in seeds contributing to browning), and potential complexation with polysaccharides and proteins, with these changes reducing both total extractability and the proportion of small, highly astringent oligomers that contribute to harsh mouthfeel in wines from unripe fruit. The sensory evolution of tannins during maturation progresses from green, harsh, and bitter character in unripe fruit (dominated by extractable seed tannins and small oligomers) toward softer, rounder, and more integrated character as polymerization proceeds and the relative contribution of skin tannins increases, with this sensory trajectory being the practical target of tannin maturity assessment for harvest timing. Understanding tannin maturity quality enables assessment of wine structural potential beyond simple phenolic indices, harvest timing for desired tannin character and mouthfeel, and recognition of the tissue-specific and polymerization-dependent factors affecting tannin quality in resulting wines.
Tannin_Maturity_Quality addresses the developmental state of proanthocyanidins determining wine structure and mouthfeel. This quality involves quantity, polymerization degree, tissue source, and sensory character. The class supports structural quality optimization through tannin maturity characterization.
Gerbstoffreife
Proanthocyanidinreife
Tanninreife
condensed tannin development
proanthocyanidin maturity
structural phenolic maturity
tannin ripeness
Kennedy JA et al. (2002) Development of seed polyphenols in berries from Vitis vinifera L. cv. Shiraz. Australian Journal of Grape and Wine Research 8(1): 25-35 Harbertson JF et al. (2002) A comparison of berry and wine tannin of Merlot, Cabernet Sauvignon, and Cabernet Franc. American Journal of Enology and Viticulture 53(1): 54-59
Tannin Maturity Quality
Tannin mouthfeel assessment comparing Pomerol (Merlot-dominant, clay soils) and Pauillac (Cabernet Sauvignon-dominant, gravel soils) reveals distinctly different textural signatures despite equivalent astringency intensity ratings of 5.8, with Pomerol showing plush, velvety mouthfeel with sweet tannin impression and Pauillac showing firm, structured mouthfeel with more angular grip, demonstrating how the integrated tannin character creates regional identity beyond what individual astringency parameters capture, and explaining the food pairing differences where Pomerol suits rich, umami dishes while Pauillac demands char-grilled red meat.
Tannin mouthfeel quality is an astringency quality that represents the integrated textural impression created by tannin-induced oral sensations, characterizing the holistic tactile experience encompassing not only drying astringency but also fullness, weight, grip, and structural contribution to wine body and palate presence. Within the BFO framework, this quality addresses the complete mouthfeel contribution of tannins as a unified perceptual gestalt, distinguishing the integrated tannin experience from individual astringency dimensions while recognizing that tannin mouthfeel includes textural elements beyond pure astringency such as the perception of palate weight and structural framework. The ontological significance lies in tannin mouthfeel defining the structural identity of red wines, where the integrated tannin character creates the wine's textural signature distinguishing varieties, regions, and production styles more than any other single attribute, and determining both immediate drinkability and aging potential. The perceptual integration of tannin mouthfeel involves synthesis of astringency (drying, puckering), texture (fine to coarse), weight (light to heavy on palate), grip (holding sensation), and length (persistence of tactile sensation) into a unified impression that experienced tasters evaluate holistically rather than analytically decomposing into components. The compositional basis includes total tannin concentration determining weight and intensity, tannin molecular size distribution affecting texture and character, tannin-anthocyanin and tannin-polysaccharide complexes modifying perception, and the wine matrix (alcohol, glycerol, residual sugar) providing context for tannin integration. The stylistic diversity of tannin mouthfeel spans from the silky, seamless tannins of great Burgundy through the structured, firm tannins of Bordeaux to the robust, chewy tannins of Barossa Shiraz, with each profile representing a distinctive combination of intensity, character, and development appropriate to variety and regional tradition. Understanding tannin mouthfeel quality enables holistic assessment of tannin contribution, appreciation of stylistic diversity in tannin expression, and quality evaluation based on the integrated tactile experience that defines red wine structure.
Tannin_Mouthfeel_Quality provides the holistic assessment of tannin's tactile contribution, integrating the analytical dimensions addressed by Astringency_Intensity_Quality, Astringency_Character_Quality, and Astringency_Development_Quality into a unified gestalt evaluation. This quality represents the practical way experienced tasters assess tannin, complementing the analytical decomposition with integrative assessment. The class supports variety and regional typicity evaluation, aging potential prediction, and food pairing recommendation based on textural profile.
Gerbstoffhaptik
Tanninmundgefühl
Tanninstruktur
Tannintextur
tannin grip
tannin structure
Gawel R et al. (2000) A 'Mouth-feel Wheel': terminology for communicating the mouth-feel characteristics of red wine. Australian Journal of Grape and Wine Research 6(3): 203-207 Sáenz-Navajas MP et al. (2012) Perception of mouthfeel properties of red wines by trained assessors. Food Quality and Preference 26(2): 112-120
Tannin Mouthfeel Quality
In Tannat berries in Madiran, France during BBCH 79-89, tannin polymerization modifies seed tannin mean degree of polymerization from 8.5 at veraison to 6.2 at harvest through oxidative cleavage concurrent with browning, while skin tannin mDP increases from 28 to 35 through continued condensation, with total condensed tannin content in skins increasing from 2.8 to 5.5 mg/g while seed tannin concentration per berry remains stable but extractability decreases 45% through structural modification under warm conditions averaging 26°C.
Tannin polymerization constitutes the progressive structural modification of proanthocyanidin compounds in Vitis vinifera berry tissues during ripening, wherein monomeric flavan-3-ol units including catechin, epicatechin, and their galloylated derivatives undergo oxidative and enzymatic condensation reactions generating the polymeric tannin structures that determine wine astringency character, mouthfeel properties, and color stabilization capacity over the course of berry development from early formation through harvest maturity. This process involves multiple reaction mechanisms including acid-catalyzed interflavonoid bond formation connecting C4 of one unit to C8 or C6 of another, oxidative coupling generating additional linkage types, and potential depolymerization through cleavage reactions that modify the mean degree of polymerization characterizing tannin populations in different berry tissues and at different developmental stages. The quantitative aspects of tannin polymerization show tissue-specific patterns, with seed tannins demonstrating mean degree of polymerization of 5-12 units at veraison decreasing slightly through oxidative modification during ripening, while skin tannins show mDP of 25-45 units that may increase modestly through continued polymerization during the post-veraison period. The biochemical basis of tannin polymerization involves both enzymatic and non-enzymatic mechanisms, with polyphenol oxidases and peroxidases potentially catalyzing oxidative coupling in the presence of oxygen, while acid-catalyzed condensation proceeds spontaneously under the acidic vacuolar conditions of pH 2.5-3.5 where proanthocyanidins accumulate. Structural changes accompanying polymerization include modification of interflavonoid linkage distribution, with A-type bonds forming through additional oxidative coupling between already-linked units, galloylation patterns evolving through continued esterification, and potential incorporation of anthocyanins into tannin polymers generating pigmented tannin species that contribute to wine color stability. The sensory significance of tannin polymerization relates to astringency perception, with highly polymerized tannins of mDP greater than 20-30 demonstrating greater astringent intensity but potentially softer quality compared to shorter oligomers, while the galloylation percentage influences both astringency intensity and quality contributing to the characteristic mouthfeel differences among grape varieties. Temperature during ripening influences tannin polymerization through effects on oxidative reaction rates and enzymatic activities, with warm conditions potentially accelerating polymerization and associated browning reactions particularly in seed tissues where oxidative modification is pronounced during ripening.
Tannin_Polymerization is parent class under Phenolic_Maturation representing proanthocyanidin structural modification. Contains subclasses Tannin_Chain_Elongation, Condensation_Reactions, and Tannin_Structure_Maturation. Occurs in both skin and seed tissues with different dynamics. Seed mDP 5-12 decreases slightly skin mDP 25-45 may increase. Influences astringency perception and color stability. Oxidative and acid-catalyzed mechanisms.
Proanthocyanidin-Polymerisation
Tannin-Ketten-Wachstum
Tannin-Polymerisation
condensed tannin polymer formation
kondensierte Tannin-Polymer-Bildung
proanthocyanidin polymerization
tannin chain growth
tannin structural development
Kennedy JA et al (2001) Changes in grape seed polyphenols during ripening. Phytochemistry 55: 77-85 Downey MO et al (2003) Analysis of tannins in seeds and skins of Shiraz. Australian Journal of Grape and Wine Research 9: 15-27 Keller M (2020) The Science of Grapevines 3rd Ed pp 325-335
Tannin Polymerization
Mean degree of polymerization (mDP) ranges from 5-50 subunits across tannin sources in grape berries, with skin tannins showing mDP 15-40 versus seed tannins mDP 3-12. Cultivar differences in skin tannin mDP range from 15-20 in shorter-polymer types such as Pinot Noir to 30-45 in longer-polymer cultivars such as Cabernet Sauvignon and Tannat. Polymerization occurs through oxidative coupling of flavan-3-ol units via 4→8 or 4→6 interflavan bonds, with chain extension continuing in vacuolar storage. Longer polymers (mDP >20) show reduced astringency per unit mass due to steric constraints on protein binding, producing "softer" mouthfeel despite higher total tannin. The polymer size distribution (polydispersity) affects astringency character, with narrow distributions producing more uniform sensory response.
A tannin accumulation disposition that is the inherited capacity of a Vitis vinifera plant to produce proanthocyanidin polymers of characteristic chain length (mean degree of polymerization, mDP), with genotype-specific polymer size distributions affecting astringency intensity, astringency quality, and protein precipitation capacity of extracted tannins.
Tannin polymerization degree directly affects the sensory quality of wine tannins, with the relationship between polymer size and astringency following an inverted U-curve (maximum astringency at intermediate mDP 8-15). The trait shows moderate heritability with developmental and environmental modulation, as polymerization continues during berry maturation and can be affected by oxidative conditions. Understanding polymerization enables interpretation of total tannin concentration in relation to sensory outcomes, as high concentration with high mDP may produce less astringency than moderate concentration with intermediate mDP.
Kettenlängen-Disposition
PA polymer length
Polymerisationsgrad
Tanninpolymerisation
proanthocyanidin mDP disposition
tannin chain length trait
Vidal S et al. (2003) The mouth-feel properties of grape and apple proanthocyanidins in a wine-like medium. Journal of the Science of Food and Agriculture 83:564-573 Kennedy JA, Jones GP (2001) Analysis of proanthocyanidin cleavage products following acid-catalysis in the presence of excess phloroglucinol. Journal of Agricultural and Food Chemistry 49:1740-1746
Tannin Polymerization Disposition
Tannin polymerization quality tracking in Brunello di Montalcino follows the evolution from harvest through five years minimum aging, with seed-derived tannins (initial mDP 8-12) and skin tannins (initial mDP 25-40) progressively polymerizing during mandatory 24+ months wood aging and subsequent bottle aging the transformation from the austere, drying tannins of young Sangiovese (high proportion of medium-chain astringent polymers) to the silky, fine-grained texture of mature Brunello (predominance of large polymers beyond peak astringency) illustrates how polymerization quality evolution governs the sensory transformation defining great Sangiovese aging.
Tannin_Polymerization_Quality represents the chemical quality characterizing the degree of polymerization (DP) and polymer size distribution of tannins in grape tissues and wine, capturing the structural parameter that profoundly affects sensory properties independent of total tannin concentration. The mean degree of polymerization (mDP) describes the average number of monomeric subunits in tannin polymer chains, ranging from 2-4 for small oligomers through 10-20 for medium polymers to 30-80+ for large polymers characteristic of grape skins. Polymerization state determines multiple quality-relevant properties: small polymers (DP 2-5) often taste bitter rather than astringent medium polymers (DP 6-15) produce classic astringent character large polymers (DP greater than 30) may precipitate before interacting with salivary proteins, reducing perceived astringency. The relationship between polymer size and astringency is not linear but follows an inverted U-shape where mid-size polymers produce maximum astringency while very small and very large polymers contribute less. Seed tannins typically show lower mDP (5-15) than skin tannins (20-80), contributing to the different mouthfeel qualities of seed versus skin tannin extraction. During wine aging, tannins undergo progressive polymerization through oxidative coupling and reaction with aldehydes and anthocyanins, generally producing softer mouthfeel as polymers grow beyond the maximum astringency range. The rate of polymerization depends on oxygen exposure, temperature, and wine composition, with controlled micro-oxygenation enabling accelerated tannin evolution. Assessment of polymerization quality requires analytical methods that determine polymer size distribution, with phloroglucinolysis providing mDP calculations for proanthocyanidins.
Tannin_Polymerization_Quality addresses the polymer size parameter that modulates sensory impact independently of concentration. The mDP metric provides a single number summarizing average polymer length but does not capture size distribution advanced methods using gel permeation chromatography or mass spectrometry provide distribution information. The relationship between mDP and sensory properties involves complex interactions with wine matrix, as alcohol and polysaccharides modify astringency perception at any given tannin size. The temporal evolution of polymerization during aging represents a key mechanism underlying the softening of tannins in maturing wines.
Tanninkettenlängenqualität
Tanninpolymergröße
Tanninpolymerisationsqualität
degree of polymerization quality
mDP quality
tannin chain length quality
tannin polymer size
Vidal S et al. (2003) The Mouth-Feel Properties of Grape and Apple Proanthocyanidins in a Wine-Like Medium. Journal of the Science of Food and Agriculture 83(6): 564-573 Kennedy JA and Jones GP (2001) Analysis of Proanthocyanidin Cleavage Products Following Acid-Catalysis in the Presence of Excess Phloroglucinol. Journal of Agricultural and Food Chemistry 49(4): 1740-1746
Tannin Polymerization Quality
Tannin quality assessment in Barolo from Nebbiolo integrates multiple parameters including condensed tannin concentration (2500-4000 mg/L catechin equivalents), mean degree of polymerization (mDP 15-30 indicating large polymers), and the proportion of skin versus seed-derived tannins (targeting greater than 60% skin tannin for velvety texture) the traditional extended maceration (40-60 days) followed by years of large oak aging allows tannin polymerization and softening that transforms the initially fierce, drying astringency of young Barolo into the elegant, fine-grained texture of mature wine while maintaining the tannic structure enabling decades of cellaring.
Tannin_Quality encompasses the chemical qualities characterizing the concentration, composition, structure, and properties of polymeric phenolic compounds capable of precipitating proteins, capturing the macromolecular phenolics responsible for astringency, structure, and aging potential in red wines. Tannins are distinguished from monomeric phenolics by their higher molecular weight and protein-binding capacity, with the astringent sensation arising from tannin interaction with salivary proteins that reduces oral lubrication. This category includes Condensed_Tannin_Quality addressing proanthocyanidins (flavan-3-ol polymers) that constitute the dominant tannin class in grape-derived wines Hydrolyzable_Tannin_Quality capturing gallotannins and ellagitannins extracted from oak during barrel aging Tannin_Polymerization_Quality describing the degree of polymerization affecting sensory properties and Tannin_Extractability_Quality addressing the factors governing tannin release during winemaking. The concentration of condensed tannins in wine typically ranges from 100-400 mg/L in light reds through 1000-4000 mg/L in heavily extracted styles, with substantial additional hydrolyzable tannin contribution (200-800 mg/L) in oak-aged wines. Tannin concentration alone incompletely predicts sensory properties because polymer size, subunit composition, and degree of modification profoundly affect astringency quality. Small polymers may taste bitter rather than astringent, while very large polymers may precipitate before interacting with salivary proteins. Tannin evolution during wine aging involves progressive polymerization, oxidation, and reaction with anthocyanins, producing softer mouthfeel and color stability. Assessment of tannin quality requires multiple analytical approaches capturing concentration, size distribution, and compositional characteristics.
Tannin_Quality serves as the parent class for tannin-related quality parameters, positioned within Phenolic_Content_Quality to capture the polymeric phenolic fraction. The distinction between condensed tannins (proanthocyanidins, grape-derived) and hydrolyzable tannins (gallotannins/ellagitannins, primarily oak-derived) reflects different chemical structures and sensory properties. Tannin quality extends beyond concentration to encompass structural features (polymerization, composition) and functional properties (extractability, protein precipitation) that determine sensory impact. The term tannin originally referred to materials used in leather tanning and retains this protein-precipitation connotation.
Weintannincharakter
adstringierende Phenolqualität
astringent phenolic quality
polymerer Phenolgehalt
polymeric phenolic quality
tannin content and composition
wine tannin character
Kennedy JA et al. (2006) Grape and Wine Phenolics: History and Perspective. American Journal of Enology and Viticulture 57(3): 239-248 Gawel R (1998) Red Wine Astringency: A Review. Australian Journal of Grape and Wine Research 4(2): 74-95
Tannin Quality
In Aglianico berries in Taurasi, Campania during BBCH 81-89, tannin structure maturation modifies skin tannin properties with mDP increasing from 25 to 34, galloylation remaining stable at 5.2-5.8%, and prodelphindin content at 32-35% over 48 days, while seed tannin structure shows mDP decline from 9.2 to 7.5, galloylation stable at 28-31%, and progressive browning reducing extractability by 52% as measured by protein precipitation assay, establishing the mature phenolic profile characteristic of this late-ripening variety under warm southern Italian conditions.
Tannin structure maturation describes the comprehensive evolution of proanthocyanidin structural features in Vitis vinifera berry tissues during ripening, wherein initial tannin populations undergo progressive modification through polymerization, oxidation, and compositional changes that transform structural properties including mean degree of polymerization, galloylation percentage, prodelphindin content, and subunit composition to establish the mature tannin structures characterizing harvest-ready fruit with appropriate astringency potential and wine quality attributes. This process encompasses multiple concurrent structural modifications including potential chain elongation through continued condensation increasing mean polymer length in skin tissues, oxidative browning particularly pronounced in seed tannins where quinone formation and subsequent reactions modify chromophore properties and reduce extractability, and shifts in the proportions of different extension and terminal units reflecting continued biosynthesis and potential selective degradation of specific structural forms. The galloylation percentage, representing the proportion of flavan-3-ol units bearing gallic acid ester substituents at the C3 position, typically remains relatively stable during ripening in most varieties though slight increases may occur through continued galloylation of existing polymers, with final galloylation levels of 15-35% in seeds and 2-8% in skins contributing to the astringency intensity and quality differences between tannins from these tissues. The prodelphindin content, reflecting units bearing trihydroxylated B-rings derived from epigallocatechin incorporation, shows tissue-specific patterns with skin tannins containing 20-40% prodelphindin units compared to seed tannins with typically less than 5%, contributing to differences in protein binding behavior and sensory properties between skin and seed tannin populations. Oxidative modification of tannin structure proceeds through polyphenol oxidase and peroxidase catalyzed reactions generating quinone intermediates that undergo nucleophilic addition, rearrangement, and polymerization, producing the characteristic browning of seed coat tissues during ripening and reducing tannin extractability through formation of oxidized polymers with modified solubility properties. The sensory significance of tannin structure maturation relates to the evolution of astringency quality, with mature tannin structures typically demonstrating softer, rounder astringency compared to harsh, aggressive character associated with immature seed tannins, reflecting structural changes that modify protein precipitation kinetics and oral sensation development during tasting.
Tannin_Structure_Maturation is subclass of Tannin_Polymerization representing comprehensive structural evolution. Distinguishes from individual modification processes by integrating multiple structural changes. Sibling to Tannin_Chain_Elongation and Condensation_Reactions. Encompasses mDP, galloylation, prodelphindin, and oxidation changes. Tissue-specific patterns in skin versus seed. Relates to astringency quality evolution.
Proanthocyanidin-Struktur-Entwicklung
Tannin-Qualitäts-Reifung
Tannin-Struktur-Evolution
Tannin-Struktur-Reifung
proanthocyanidin structure development
tannin compositional change
tannin quality maturation
tannin structural evolution
Kennedy JA et al (2001) Changes in grape seed polyphenols during ripening. Phytochemistry 55: 77-85 Ristic R and Iland PG (2005) Relationships between seed and berry development of Vitis vinifera L. Australian Journal of Grape and Wine Research 11: 43-58 Adams DO (2006) Phenolics and ripening in grape berries. American Journal of Enology and Viticulture 57: 249-256
Tannin Structure Maturation
Cabernet Sauvignon grafted on 140 Ruggeri rootstock at 10 years showing tap root as deepest primary root with predominantly vertical orientation 82° from horizontal, tap root extending 195 cm depth in deep sandy loam substantially deeper than lateral root network maximum depth 110 cm, tap root diameter 9.2 cm at crown tapering to 3.8 cm at 150 cm depth, tap root showing sparse lateral branching compared to oblique primary roots, excavation revealing tap root accessing moist soil at 180-195 cm depth during late summer when upper 120 cm soil dry, tap root providing deep water access enabling drought tolerance characteristic of vigorous 140Ru rootstock
Tap root is primary root showing predominantly vertical orientation growing directly downward from crown into deep soil layers, characterized by strong positive gravitropic response maintaining vertical or near-vertical growth direction typically 70-90° from horizontal surface, relatively unbranched main axis penetrating deep soil profile often exceeding 150-200 cm depth in permeable soils, large diameter typically 5-15 cm at crown tapering gradually with depth, and functioning as main deep water access point enabling drought tolerance through exploitation of deep moisture reserves unavailable to lateral or shallow roots, with tap root architecture most pronounced in seedling-derived vines showing single dominant vertical radicle-derived root and less distinct in cutting-derived vines showing multiple oblique primary roots though deepest vertical root often designated tap root functionally. Tap root vertical orientation includes= growing predominantly downward (tap root showing 70-90° angle from horizontal maintaining vertical penetration), exhibiting strong gravitropism (positive gravitropic response directing growth toward earth center regardless of planting position), demonstrating directional persistence (tap root maintaining downward trajectory unless deflected by obstacles or soil conditions), showing rootstock variation (some rootstock varieties producing particularly straight vertical tap roots while others more oblique), and influenced by soil structure (deep permeable soils allowing straight vertical growth while layered or compacted soils potentially deflecting tap root). Tap root depth penetration includes= extending deep into profile (tap root typically reaching 100-200 cm depth in normal agricultural soils potentially exceeding 300-600 cm in deep deposits or under drought stress), exceeding lateral root depth (tap root penetrating substantially deeper than lateral root network accessing moisture and nutrients in deep soil layers), showing progressive deepening (tap root continuing downward growth throughout vine life potentially adding 10-50 cm depth annually in young vine slowing in old vine), demonstrating rootstock genetic control (Berlandieri-based rootstocks showing deep tap root penetration while Riparia-based showing shallower systems), and limited by soil conditions (tap root depth constrained by bedrock, hardpans, water tables, toxic layers, or severe compaction preventing further penetration). Tap root diameter characteristics includes= showing large diameter at crown (tap root typically 5-15 cm diameter where emerging from trunk base depending on vine age and vigor), exhibiting gradual taper (diameter decreasing progressively with depth as root extends deeper and branches), demonstrating secondary thickening (vascular cambium adding annual xylem rings increasing tap root diameter at all depths), displaying age-diameter correlation (older tap roots showing greater diameter than young roots reflecting years of secondary growth), and varying with rootstock (vigorous rootstocks producing thicker tap roots than low-vigor varieties reflecting overall growth capacity). Tap root branching patterns includes= showing sparse lateral production (tap root typically less heavily branched than oblique primary roots focusing resources on vertical penetration), exhibiting lateral root emergence (branch roots forming along tap root length at intervals typically 10-30 cm), demonstrating decreasing lateral density with depth (upper tap root portions showing more lateral roots than deep sections), displaying lateral root angles (branch roots typically emerging 60-90° from vertical tap root creating widespread lateral network), and enabling hierarchical architecture (tap root with laterals creating multi-order root system exploring large soil volume). Tap root deep water access includes= penetrating to water table or deep moisture (tap root potentially reaching permanent moisture in deep soil layers or capillary fringe above water table), enabling drought avoidance (deep tap root accessing water unavailable to shallow roots allowing vine to maintain function during surface soil drought), showing seasonal importance (tap root contribution to vine water supply greatest during summer when surface soil dry), demonstrating cultivar adaptation (deep tap root essential for vine survival in arid regions or droughty soils), and providing competitive advantage (tap root allowing vine to access resources unavailable to neighboring plants with shallow root systems). Tap root anchorage function includes= providing vertical stability (deep tap root penetration creating strong anchor preventing vine uprooting), resisting pulling forces (tap root bearing vertical tension loads from wind or weight on above-ground structure), demonstrating wind resistance (tap root contribution to anchorage particularly important for preventing toppling in exposed sites), showing age-related strengthening (deeper older tap root providing progressively greater stability as penetrates and thickens), and complementing lateral roots (tap root vertical anchorage working with lateral root horizontal stabilization creating robust anchoring system). Tap root in seedling versus cutting-derived vines includes= highly developed in seedlings (seedling vine typically showing single dominant tap root derived from radicle growing straight downward from germination), less distinct in propagated vines (cutting-derived vines showing multiple adventitious primary roots from cutting base with deepest designated tap root functionally), showing functional equivalence (cutting deepest vertical root serving tap root functions despite different developmental origin), demonstrating similar depth potential (both seedling and cutting tap roots capable of deep penetration given favorable conditions), and varying in structure (seedling single straight tap root versus cutting potentially oblique multiple-primary-root system with one deepest root). Tap root storage capacity includes= storing substantial carbohydrate reserves (large tap root volume with abundant parenchyma and ray cells accumulating starch during dormancy), demonstrating strategic location (tap root reserves positioned deep protecting from surface soil temperature extremes and readily accessible to deep lateral roots), showing seasonal cycles (tap root reserves highest fall-winter declining spring as mobilized for growth), providing drought stress reserves (tap root carbohydrates potentially sustaining vine during extended drought when photosynthesis limited), and complementing lateral root storage (tap root contributing to total root system storage capacity along with structural lateral roots).
Tap_Root emphasizes vertical orientation and deep penetration and is distinguished from other Primary_Root types by focus on downward growth and depth. Includes Deep_Tap_Root, Shallow_Tap_Root, and Central_Tap_Root. Critical for understanding deep water access, drought tolerance, vertical penetration, gravitropic response, and the vertically oriented primary root exploiting deep soil resources.
Deep penetrating root
Hauptvertikalwurzel
Main tap root
Pfahlwurzel
Tiefenwurzel
Vertical primary root
Tap Root
Tartaric acid accumulation occurs primarily during the green berry stage (pre-veraison), with synthesis essentially complete by veraison and concentrations of 5-15 g/L fresh weight in immature berries declining through dilution to 4-10 g/L at harvest. Cultivar differences in final tartrate concentration span 2-fold, with Riesling and Chenin Blanc showing high tartrate (7-10 g/L) and Grenache and Carignan showing lower values (4-6 g/L). The biosynthetic pathway proceeds from L-ascorbic acid through L-idonic acid to L-tartaric acid, with L-idonate dehydrogenase (VvLIdnDH) being a key regulatory enzyme. Unlike malic acid, tartaric acid is not metabolized during ripening, making it the stable backbone of grape and wine acidity.
An acid metabolism disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate tartaric acid in grape berries through the ascorbic acid pathway, with genotype-specific concentrations determined by biosynthetic enzyme activity during early berry development and the absence of significant catabolic pathways in Vitis.
Tartaric acid is nearly unique to grapes among major fruit crops, providing the distinctive acid character of wine. The metabolic stability of tartrate (no respiratory degradation) makes it the reliable acidity component independent of ripening temperature, while malic acid varies with climate. Tartrate concentration at harvest depends primarily on synthesis rate during early development and subsequent dilution by berry expansion. Potassium bitartrate precipitation during winemaking can reduce effective tartrate contribution to wine acidity.
L-tartrate accumulation
Tartratbildung
Weinsäure-Disposition
Weinsäureakkumulation
tartaric acid trait
tartrate synthesis capacity
DeBolt S et al. (2006) L-Tartaric acid synthesis from vitamin C in higher plants. Proceedings of the National Academy of Sciences 103:5608-5613 Ford CM (2012) The biochemistry of organic acids in the grape. In: Gerós H et al. (eds) The Biochemistry of the Grape Berry, Bentham Science pp.: 67-88
Tartaric Acid Accumulation Disposition
Tartaric acid content for Riesling from the Rheingau (2024 vintage) measures 7.5 g/L at harvest (82° Oechsle), representing the characteristically high tartrate retention of cool-climate Riesling that provides the structural acid backbone supporting residual sugar balance in Spätlese and Auslese wines. Tartaric acid content for Cabernet Sauvignon from Coonawarra (2023 vintage) records 5.8 g/L at 13.2° Baumé, with the moderate tartrate level reflecting the balance between synthesis during early berry development and partial depletion through potassium bitartrate precipitation during the warm ripening period. Tartaric acid content for Grüner Veltliner from Wagram (2024 vintage) documents 6.5 g/L at 17.5° KMW, with the well-preserved tartrate concentration contributing to the cultivar's signature crisp mouthfeel and facilitating the minerality perception associated with the region's loess-derived soils.
Tartaric acid content is an acidity data entity that records the concentration of L-(+)-tartaric acid (2,3-dihydroxybutanedioic acid) in grape must, determined by enzymatic assay, HPLC, or colorimetric method, expressed in grams per liter (g/L), representing the dominant fixed acid of grape berries and the primary structural component of the wine acid matrix, whose concentration is uniquely high in Vitis species compared with other fruit crops and whose relative stability during ripening and fermentation makes it the backbone acid that maintains pH and preserves wine freshness throughout aging.
Tartaric acid occupies a privileged position in grape and wine chemistry as the quantitatively dominant organic acid (typically 4-10 g/L in must) and the most important contributor to pH buffering in the wine pH range (pKa1 = 3.04, pKa2 = 4.37). Uniquely among major fruit crops, Vitis species accumulate tartaric acid to high concentrations through a biosynthetic pathway from L-ascorbic acid (vitamin C) that is active primarily during early berry development (pre-véraison, BBCH 71-79), with synthesis essentially ceasing after véraison. The relative stability of tartaric acid during ripening — unlike malic acid, it is not a significant respiratory substrate — makes tartrate the reliable acid constant against which ripening-induced changes in malic acid and potassium are measured. However, tartaric acid is not entirely immutable — potassium bitartrate crystallization removes tartrate from solution when the solubility product is exceeded (common in high-potassium musts from warm regions), and Botrytis cinerea metabolizes tartaric acid to gluconic and galacturonic acids during berry infection. The analytical methods for tartaric acid include enzymatic assay (L-tartrate dehydrogenase, highly specific), HPLC (simultaneous separation of all organic acids), capillary electrophoresis, and the classical colorimetric Rebelein method — with HPLC increasingly preferred for its multi-acid capability. Tartaric acid content data informs acidification decisions (tartaric acid addition being the most common permitted acidulant in winemaking), cold stabilization predictions (higher tartrate with high potassium indicates greater precipitation risk), and aging trajectory assessment (wines with higher initial tartrate exhibit greater pH stability over time).
L-Weinsäure
L-tartaric acid content
Tartratgehalt
Weinsäurebestimmung
Weinsäuregehalt
Weinsäurekonzentration
tartaric acid concentration
tartrate analysis
tartrate content
tartrate level
Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 395-410 DeBolt S et al. (2006) L-Tartaric acid synthesis from vitamin C in higher plants. Proceedings of the National Academy of Sciences 103(14): 5608-5613 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2, 2nd Ed, Wiley, pp.: 3-20
Tartaric Acid Content
Tartaric acid content quality in Barbera from Piedmont reaches exceptionally high levels (7-10 g/L at harvest) that define this variety's characteristically bright, food-friendly acidity the genetic predisposition toward high tartaric acid synthesis, combined with Barbera's moderate malic acid, produces wines with stable acidity that rarely requires adjustment, distinguishing Barbera from the lower-acid Dolcetto and the more tannic Nebbiolo in the Piedmontese varietal hierarchy, and enabling the fresh, vivacious style that makes Barbera the region's everyday wine while Barolo ages in cellar.
Tartaric_Acid_Content_Quality represents the chemical quality characterizing the concentration of tartaric acid (2,3-dihydroxybutanedioic acid, C₄H₆O₆) in grape berry tissue. Tartaric acid is the signature organic acid of grapes, relatively rare in the plant kingdom but synthesized in large quantities by Vitis species through a unique metabolic pathway from ascorbic acid. This acid provides the majority of stable acidity in wine because it is not metabolized by yeasts or most bacteria under normal winemaking conditions, persisting from grape through finished wine (minus any precipitation as tartrate salts). Tartaric acid concentration in ripe berries typically ranges from 4-9 g/L, with values established primarily during early berry development (pre-véraison) and remaining relatively constant during ripening because grape berries lack the enzymes to catabolize tartaric acid. This metabolic stability contrasts sharply with malic acid, which declines substantially during ripening. The concentration of tartaric acid is influenced by cultivar genetics, with some varieties (Riesling, Silvaner) accumulating higher levels than others. Environmental factors affecting tartaric acid content are primarily pre-véraison conditions that influence acid synthesis, with minimal effect of ripening temperature on final concentration. Tartaric acid contributes the hard, sharp acidity that characterizes wine, providing the backbone that balances sugar sweetness and supports wine structure. In winemaking, tartaric acid is the preferred acid for acidification treatments because it integrates naturally with grape acidity. Measurement employs specific enzymatic assays using tartrate dehydrogenase or HPLC separation, distinguishing tartaric acid from malic acid that would be included in non-specific acid determinations.
Tartaric_Acid_Content_Quality addresses the most distinctive and stable acid of grape and wine. The L-(+)-tartaric acid stereoisomer (natural tartaric acid) is the form present in grapes other stereoisomers (D-tartaric, meso-tartaric) are not naturally synthesized. Tartaric acid forms salts with potassium (potassium bitartrate, cream of tartar) and calcium (calcium tartrate) that may precipitate during winemaking and storage, affecting both acidity and visual stability. The unique biosynthetic pathway in Vitis (from ascorbic acid via 5-keto-D-gluconic acid) distinguishes grape tartaric acid metabolism from all other plant species.
2,3-dihydroxysuccinic acid concentration
L-tartaric acid level
Tartrat
Traubensäuregehalt
grape acid quality
DeBolt S et al. (2006) L-Tartaric Acid Synthesis from Vitamin C in Higher Plants. Proceedings of the National Academy of Sciences 103(14): 5608-5613 Sweetman C et al. (2009) Regulation of Malate Metabolism in Grape Berry and Other Developing Fruits. Phytochemistry 70(11-12): 1329-1344
Tartaric Acid Content Quality
In Cabernet Franc berries in Loire Valley, France during BBCH 81-89, tartaric acid stabilization maintains total berry tartrate content at 4.2-4.5 mg per berry throughout the 45-day ripening period, while volumetric concentration decreases from 9.2 to 6.8 g/L due to berry volume increase from 0.95 to 1.42 mL, with potassium bitartrate precipitation accounting for 18% apparent tartrate loss under harvest potassium levels of 42 mM.
Tartaric acid stabilization describes the characteristic metabolic stability of L-tartaric acid in Vitis vinifera berries during the ripening period, wherein tartrate concentrations established during early berry development remain essentially unchanged at 5-10 g/L throughout the post-veraison maturation phase due to the absence of significant enzymatic degradation pathways in grape berry tissues, contrasting fundamentally with the rapid catabolism of malic acid that occurs during the same developmental window. This process reflects the unique biochemical status of tartaric acid among plant organic acids, as grapevines lack the tartrate-degrading enzymes present in some other plant species, rendering the tartrate pool metabolically inert once synthesized during the pre-veraison period spanning BBCH 71-81. The cellular basis of tartaric acid stabilization involves sequestration within vacuolar compartments of mesocarp parenchyma cells and specialized crystal idioblast cells, where the acidic vacuolar environment of pH 2.5-3.5 maintains tartrate in predominantly dissociated ionic forms that are not substrates for available metabolic enzymes. The concentration stability observed on a per-berry basis masks apparent concentration changes when expressed volumetrically, as berry expansion during the secondary growth phase of 50-100% volume increase produces dilution effects reducing tartaric acid concentration from 8-12 g/L at veraison to 5-8 g/L at harvest without actual tartrate catabolism occurring. Potassium accumulation during ripening, with berry potassium content increasing from 20-30 to 35-50 mM, results in formation of sparingly soluble potassium bitartrate crystals that precipitate within vacuolar compartments, effectively removing 10-25% of total tartrate from solution while the precipitated salt remains within berry tissues. Temperature exerts minimal direct influence on tartaric acid stability compared to its profound effects on malate degradation, though indirect effects through modulation of berry expansion rates and potassium accumulation patterns influence final tartrate concentrations at harvest. Cultivar variation in tartaric acid stabilization reflects genetic differences in initial tartrate biosynthesis capacity, berry size determining dilution effects, and potassium uptake efficiency influencing precipitation losses, with typical harvest concentrations ranging from 4-6 g/L in large-berried table grape varieties to 7-10 g/L in small-berried wine grape cultivars. The maintenance of stable tartaric acid levels throughout ripening establishes tartrate as the primary contributor to wine titratable acidity following malolactic fermentation, providing the structural acid backbone essential for wine stability, aging potential, and sensory balance. Viticultural practices influencing tartaric acid stabilization operate primarily through effects on berry size and potassium status rather than direct metabolic modulation, with irrigation management, crop load adjustment, and rootstock selection representing the primary management tools for optimizing harvest tartrate levels.
Tartaric_Acid_Stabilization is parent class under Acid_Reduction representing the contrasting metabolic fate of tartrate versus malate. Distinguishes from Malic_Acid_Degradation by emphasizing metabolic stability rather than catabolism. Contains subclasses Tartrate_Maintenance, Stable_Tartaric_Acid_Level, and Acid_Balance_Achievement. Duration spans entire ripening period of 35-50 days. Key characteristic is absence of enzymatic degradation pathways for tartrate in grape berries.
Tartrat-Stabilität
Tartrat-metabolische-Stabilität
Weinsäure-Erhaltung
Weinsäure-Stabilisierung
tartaric acid maintenance
tartaric acid preservation
tartrate metabolic stability
tartrate stability
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 6 pp 262-268 Ruffner HP (1982) Metabolism of tartaric and malic acids in Vitis. Vitis 21: 346-358 DeBolt S et al (2006) L-Tartaric acid synthesis from vitamin C in higher plants. Proceedings of the National Academy of Sciences 103: 5608-5613
Tartaric Acid Stabilization
Tartaric-malic ratio quality tracking in Hunter Valley Semillon reveals the characteristic rapid shift from T/M ratios of 1.2-1.5 at véraison to 2.5-3.5 at the traditional early harvest (10.5-11.5°Baumé) that defines the region's unique style, with warm January nights accelerating malic acid respiration this tartaric-dominant balance at relatively low sugar levels produces the austere, minerally young wines that transform through bottle aging into the complex, honeyed character of aged Hunter Semillon, demonstrating how T/M ratio interacts with harvest timing to create distinctive regional wine identity.
Tartaric_Malic_Ratio_Quality represents the chemical quality characterizing the proportional relationship between tartaric acid concentration and malic acid concentration in grape juice, expressed as a ratio (T/M or T:M) that captures the fundamental acid balance determining grape and wine character. This ratio provides a key indicator of ripening stage, climate influence, and potential wine style because the two acids behave so differently during berry development and wine production. Tartaric acid concentration remains relatively stable during ripening, while malic acid declines dramatically through temperature-dependent respiration, causing the T/M ratio to increase progressively from values near 1.0 at véraison to values of 2.0-4.0 or higher at harvest depending on ripening conditions. Cool climate regions with limited malic acid respiration produce fruit with T/M ratios of 1.5-2.5, while warm climates may yield ratios of 3.0-5.0 or higher as malic acid is extensively respired. The ratio influences taste perception because tartaric acid contributes hard, angular acidity while malic acid provides softer, fruitier sourness high T/M ratios produce wines with firm, stable acidity, while low ratios create wines with more vibrant, fresh character but greater susceptibility to softening through malolactic fermentation. The T/M ratio also indicates malolactic fermentation potential, with malic-rich (low ratio) wines offering greater scope for bacterial transformation while tartaric-dominant (high ratio) wines have less substrate for MLF regardless of winemaker intention. Cultivar genetics influence baseline T/M ratios, with some varieties (Riesling) favoring malic acid retention and others (Grenache) producing tartaric-dominant profiles under equivalent conditions. Assessment requires individual determination of tartaric and malic acid concentrations followed by ratio calculation, providing maturity and style indicators beyond total acidity measurement.
Tartaric_Malic_Ratio_Quality is positioned as a subclass of Acid_Balance_Quality, capturing the most important acid compositional relationship in grape quality assessment. The ratio can be expressed as T/M (tartaric divided by malic) or as proportional contributions (e.g., 70% tartaric, 30% malic), with equivalent information content but different numerical expression. The ratio provides more insight than individual acid concentrations alone because it captures the relative balance affecting taste and stability. Post-malolactic wines require separate consideration because malic acid conversion fundamentally alters the ratio, with T/L (tartaric/lactic) becoming relevant. Seasonal tracking of T/M ratio development provides ripening trajectory information complementary to sugar and total acidity monitoring.
Primärsäureverhältnis
T/M ratio
Tartrat-Malat-Quotient
Weinsäure-Äpfelsäure-Balance
Weinsäure-Äpfelsäure-Verhältnis
primary acid ratio
tartaric-malic balance
tartrate-malate ratio
Kliewer WM et al. (1967) Influence of Environment on Metabolism of Organic Acids and Carbohydrates in Vitis vinifera. I. Temperature. Plant Physiology 42(6): 869-884 Sweetman C et al. (2009) Regulation of Malate Metabolism in Grape Berry and Other Developing Fruits. Phytochemistry 70(11-12): 1329-1344
Tartaric Malic Ratio Quality
In Sauvignon Blanc berries in Marlborough, New Zealand during BBCH 83-89, tartrate maintenance preserves total berry tartrate at 3.8-4.1 mg per berry over 40 days of ripening, with vacuolar tartrate concentration maintained at 185-195 mM despite berry volume increase from 0.82 to 1.35 mL, demonstrating the effectiveness of tonoplast sequestration mechanisms under cool maritime ripening conditions averaging 18°C.
Tartrate maintenance encompasses the physiological processes ensuring preservation of L-tartaric acid pools within Vitis vinifera berry tissues throughout the ripening period, wherein the tartrate content established during early berry development is conserved through vacuolar sequestration, absence of degradative enzyme activity, and limited membrane permeability that collectively prevent tartrate loss from berry tissues during the 35-50 day post-veraison maturation phase. This process involves the continued function of tonoplast transport systems that originally accumulated tartrate during the pre-veraison biosynthetic period, maintaining vacuolar tartrate concentrations of 150-300 mM through active transport mechanisms that prevent passive efflux despite the substantial concentration gradient between vacuolar and cytosolic compartments. The molecular basis of tartrate maintenance reflects the evolutionary loss of tartrate-degrading enzyme systems in Vitis species, contrasting with some other plant families that possess tartrate dehydrogenase or related catabolic enzymes, rendering grape berry tartrate pools metabolically isolated from central carbon metabolism during ripening. Cellular compartmentalization contributes to tartrate maintenance through sequestration in specialized crystal idioblast cells containing calcium tartrate raphide crystals and potassium bitartrate druse crystals, with these crystalline forms representing 15-30% of total berry tartrate in a metabolically unavailable state that provides additional protection against potential degradation. The stability of tartrate maintenance under varying environmental conditions reflects the robustness of vacuolar sequestration mechanisms, with temperature extremes, water stress, and nutrient variation producing minimal effects on tartrate pool size compared to their substantial impacts on malate metabolism during ripening. Developmental regulation ensures tartrate maintenance through sustained expression of tonoplast transport proteins and vacuolar proton pumps that maintain the electrochemical gradients necessary for tartrate retention, with gene expression studies demonstrating continued activity of vacuolar ATPase and pyrophosphatase throughout the ripening period. The quantitative assessment of tartrate maintenance requires distinction between total tartrate content on a per-berry basis, which remains stable, and volumetric concentration, which decreases due to berry expansion, with research methodologies employing both metrics to accurately characterize tartrate dynamics during ripening. Berry-to-berry variation in tartrate maintenance within clusters reflects differences in initial tartrate accumulation, berry size progression, and localized potassium precipitation, contributing to the heterogeneity in final acid composition observed at harvest.
Tartrate_Maintenance is subclass of Tartaric_Acid_Stabilization representing the active physiological processes preserving tartrate pools. Distinguishes from Stable_Tartaric_Acid_Level which describes the outcome measurement. Sibling to Stable_Tartaric_Acid_Level and Acid_Balance_Achievement. Emphasizes vacuolar sequestration and absence of degradation pathways. Duration spans entire ripening period.
Tartrat-Erhaltung
Tartrat-Retention
Weinsäure-Bewahrung
Weinsäure-Pool-Konservierung
tartaric acid maintenance process
tartaric acid pool conservation
tartrate preservation
tartrate retention
Terrier N et al (2001) Changes in acidity and proton transport at the tonoplast of grape berries during development. Planta 213: 20-28 DeBolt S et al (2006) L-Tartaric acid synthesis from vitamin C in higher plants. PNAS 103: 5608-5613 Keller M (2020) The Science of Grapevines 3rd Ed pp 262-265
Tartrate Maintenance
Taste balance assessment in German Riesling Spätlese evaluates the integration of 45 g/L residual sugar with 7.8 g/L total acidity, with trained panel rating balance at 8.2 on a 10-point scale indicating excellent harmony where sweetness and acidity mutually enhance rather than mask each other, compared to a balance rating of 5.1 for a wine with similar sugar but only 5.5 g/L acidity where insufficient acid support allows sweetness to dominate creating cloying perception, demonstrating that balance depends on component ratios rather than absolute levels.
Taste balance quality is a taste quality that represents the perceived harmony among basic taste components, characterizing the degree to which sweetness, sourness, bitterness, and other taste elements integrate into a coherent, aesthetically pleasing whole rather than existing as discordant individual sensations. Within the BFO framework, this quality addresses the emergent perceptual property arising from taste component interactions, distinguishing balance from the intensities of individual components while recognizing that balance perception depends on both absolute levels and ratios among taste qualities. The ontological significance lies in taste balance serving as a primary criterion for grape and wine quality assessment, where balanced taste creates the impression of completeness and integration valued across wine styles, while imbalance (dominant sweetness, aggressive acidity, harsh bitterness) indicates quality deficiency regardless of individual component quality. The perceptual basis for balance involves both peripheral interactions (mutual suppression among taste qualities at the receptor and neural level) and central integration (cognitive assessment of component harmony based on learned expectations and hedonic valence), creating a gestalt perception that transcends component analysis. The compositional determinants of taste balance include absolute concentrations of sugars, acids, and phenolics, their ratios (particularly sugar/acid ratio as the primary balance parameter), and matrix factors affecting perception including alcohol, glycerol, and viscosity that modulate component intensities. The style-dependency of optimal balance means different wine categories have different balance targets, with dry table wines requiring acid-dominant balance, dessert wines requiring sweetness-dominant balance with sufficient acidity to prevent cloying character, and red wines requiring integration of bitterness and astringency with fruit sweetness and acidity. Understanding taste balance quality enables harvest timing for optimal component ratios, winemaking adjustment to achieve target balance, and quality assessment based on the integrative criterion that best predicts consumer acceptance.
Taste_Balance_Quality serves as the parent class for specific balance relationships (Sweet-Sour, Sweet-Bitter, Overall Harmony), establishing the framework for evaluating taste component integration. This quality represents a higher-order perceptual property emerging from basic taste interactions. Balance assessment requires consideration of wine style context, as optimal balance differs among dry, off-dry, and sweet wine categories.
Geschmacksausgewogenheit
Geschmacksbalance
Geschmacksharmonie
flavor balance
gustatorisches Gleichgewicht
gustatory balance
taste equilibrium
taste harmony
Amerine MA, Roessler EB (1983) Wines: Their Sensory Evaluation, 2nd Ed. W.H. Freeman, New York, pp. 146-172 Charters S, Pettigrew S (2007) The dimensions of wine quality. Food Quality and Preference 18(7): 997-1007
Taste Balance Quality
Taste concentration assessment in Napa Valley Cabernet Sauvignon from hillside (35 hL/ha) versus valley floor (75 hL/ha) vineyards reveals concentration ratings of 8.5 versus 5.8 on a 10-point scale, with the hillside wine showing sustained intensity across 45 seconds of palate evaluation while valley floor wine intensity fades after 20 seconds, corresponding to extract levels of 32 versus 24 g/L and demonstrating the multi-dimensional depth that distinguishes concentration from simple intensity, with the concentrated character supporting the $200 versus $40 price differential.
Taste concentration quality is a taste intensity quality that represents the elevated magnitude of gustatory sensation characteristic of wines made from concentrated grape material, intensive extraction, or reductive handling that preserves taste compounds, characterizing the dense, layered taste impression associated with premium quality and age-worthiness. Within the BFO framework, this quality addresses the high-intensity end of the taste magnitude continuum, distinguishing concentration from mere intensity by implying the multi-dimensional depth and complexity that accompanies appropriately achieved high concentration rather than simple loudness of taste. The ontological significance lies in taste concentration serving as a positive quality indicator in contexts where richness and depth are valued, suggesting careful vineyard management achieving optimal fruit concentration, selective harvesting capturing peak quality fruit, and winemaking preserving rather than diluting the concentrated material. The perceptual character of taste concentration involves not just high intensity but sustained intensity across multiple taste dimensions, creating the impression of layers of flavor emerging sequentially rather than a single powerful taste impact, with the complexity dimension distinguishing concentration from mere strength. The compositional basis for taste concentration includes elevated levels of sugars, acids, phenolics, and extract all contributing to dense taste structure, with balance among concentrated components essential to prevent any single element dominating, and the extract/alcohol ratio often serving as an index of concentration achieved without excessive must enrichment. The viticultural determinants of taste concentration include low yields concentrating fixed resources per berry, water deficit increasing solute concentration through reduced berry size, and selective harvest capturing fully mature fruit while excluding dilute or damaged berries. Understanding taste concentration quality enables assessment of vineyard and winemaking success in achieving concentrated character, prediction of age-worthiness based on the structural depth that supports evolution, and quality classification in styles where concentration is a defining attribute.
Taste_Concentration_Quality addresses the positive high-intensity end of the taste magnitude spectrum, implying quality-associated depth and complexity beyond mere loudness. This quality distinguishes from simple high intensity by incorporating the layered, sustained character of appropriate concentration. The class represents a style-dependent quality attribute, valued in wines intended for aging and special occasions but potentially excessive in wines intended for immediate, casual consumption.
Geschmacksdichte
Geschmackskonzentration
Geschmackstiefe
flavor concentration
gustatorischer Reichtum
gustatory richness
taste density
taste depth
Peynaud E, Blouin J (1996) The Taste of Wine: The Art and Science of Wine Appreciation, 2nd Ed. John Wiley & Sons, New York, pp. 174-198 Matthews MA, Anderson MM (1988) Fruit ripening in Vitis vinifera L.: responses to seasonal water deficits. American Journal of Enology and Viticulture 39(4): 313-320
Taste Concentration Quality
Taste description for Barolo DOCG (2019 vintage, 5 years post-harvest) documents a dry, medium-full body with penetrating acidity (TA 5.8 g/L perceived as firm and structural), powerful fine-grained tannins still showing youthful grip but beginning to resolve toward silkiness, moderate alcohol warmth (14.0% vol.) well-integrated into the wine's considerable extract, a slightly bitter-savoury mid-palate inflection characteristic of Nebbiolo, and an exceptionally long finish of 25+ seconds with persistent dried cherry, tar, and mineral persistence — the palate architecture confirming a wine requiring 5-10 additional years of cellaring to achieve optimal tannin-acid integration. Taste description for Grüner Veltliner Federspiel from the Wachau (2024 vintage) records dry with faint off-dry impression from fruit ripeness (RS 3.2 g/L), medium body, crisp vibrant acidity providing the structural spine (TA 6.5 g/L, pH 3.18), negligible tannin (brief skin contact only), refreshing salinity on the mid-palate, and a clean, medium-long finish of 8-10 seconds with lingering citrus-pepper character.
Taste description is a sensory description entity that documents the gustatory and tactile characterization of wine as perceived on the palate during and after tasting, encompassing the five primary taste modalities (sweetness, acidity, bitterness, saltiness, umami) and the somatosensory dimensions of mouthfeel (tannin astringency, body/weight, alcohol warmth, carbonation, texture), evaluated through a structured palate assessment that tracks the temporal evolution of sensory impressions from the attack (initial palate impression) through the mid-palate (core flavor development) to the finish (persistence and aftertaste), providing the gustatory complement to olfactory aroma description and the textural-structural assessment essential for complete wine quality evaluation.
Taste description captures the palate dimension of wine sensory evaluation that complements and often diverges from the olfactory assessment, as the oral cavity provides fundamentally different sensory information — gustatory receptor cells on the tongue detect dissolved tastants (sweet, sour, bitter, salt, umami), trigeminal nerve endings detect somatosensory stimuli (astringency, temperature, pungency, viscosity), and retronasal olfaction provides the "flavor" percept that integrates taste, smell, and touch into the composite palate experience. The temporal structure of palate assessment — attack, mid-palate, finish — reflects the sequential activation of different receptor systems and the progressive dissolution and extraction of tastants from the wine as it is held, manipulated, and swallowed. The interaction between taste modalities is central to wine balance perception — sweetness and acidity are mutually suppressive (residual sugar reduces perceived sourness while acidity reduces perceived sweetness), tannin and acidity are mutually reinforcing (both contribute to perceived astringency and structural firmness), and alcohol modulates nearly all taste perceptions (enhancing sweetness and body while softening acidity at moderate levels but increasing bitterness and burning at high levels). The concept of "balance" — the harmonious integration of sweetness, acidity, tannin, body, and alcohol into a unified palate impression where no single element dominates — represents the most universally valued quality attribute in wine and is assessed primarily through taste description rather than aroma evaluation. The wine tasting procedure requires sufficient oral contact time (typically 5-10 seconds of tongue manipulation with gentle aspiration to volatilize aroma compounds retronasally) followed by either swallowing or expectoration, with the post-swallow persistence constituting the "finish" that provides the final quality impression.
Gaumenbeschreibung
Geschmacksbeschreibung
Mundgefühlbeschreibung
Palatbeschreibung
gustatorische Beschreibung
gustatory description
mouthfeel description
palate description
palate profile
taste assessment
Jackson RS (2020) Wine Science: Principles and Applications, 5th Ed, Academic Press, pp.: 530-545 Peynaud E, Blouin J (2006) The Taste of Wine: The Art and Science of Wine Appreciation, 4th Ed, Wiley, pp.: 1-75 Lawless HT, Heymann H (2010) Sensory Evaluation of Food, 2nd Ed, Springer, pp.: 73-120
Taste Description
Taste dilution assessment in Burgundy Pinot Noir following heavy September rainfall reveals dilution ratings of 6.5 (clearly dilute) for vineyards with 40% berry weight increase from rain absorption versus 2.8 (appropriately concentrated) for vineyards with rigorous triage removing swollen berries, with the diluted lots showing rapid taste fade, hollow mid-palate, and extract levels of 18 g/L versus 26 g/L in sorted lots, demonstrating how selective harvest can rescue concentration from rain-affected vintages while unsorted fruit produces obviously diluted wine.
Taste dilution quality is a taste intensity quality that represents the diminished magnitude of gustatory sensation characteristic of wines made from diluted grape material, insufficient extraction, or excessive processing that depletes taste compounds, characterizing the thin, watery taste impression associated with quality deficiency and poor aging potential. Within the BFO framework, this quality addresses the low-intensity end of the taste magnitude continuum, distinguishing problematic dilution from intentionally light style by implying inadequate concentration resulting from viticultural or winemaking deficiency rather than deliberate stylistic choice. The ontological significance lies in taste dilution serving as a negative quality indicator suggesting high yields diluting fixed fruit resources, rain at harvest adding water to berries, insufficient extraction failing to capture available taste compounds, or processing losses removing taste-active constituents. The perceptual character of taste dilution involves not just low intensity but hollow, one-dimensional taste structure lacking the depth and persistence of properly concentrated wines, with rapid intensity fade and absence of mid-palate and finish sustaining the impression of insufficient material. The compositional basis for taste dilution includes low concentrations of sugars, acids, phenolics, and extract resulting in thin structure lacking the density to create satisfying taste impression, often accompanied by imbalance where dilution affects some components more than others disrupting appropriate ratios. The viticultural causes of taste dilution include excessive yields spreading limited vine resources across too many berries, rainfall diluting berry contents particularly late in ripening, and irrigation excess preventing the concentration that mild water deficit provides. The winemaking causes include insufficient maceration time or temperature failing to extract available taste compounds, excessive fining removing desirable constituents, and oxidative processing depleting volatile flavor compounds. Understanding taste dilution quality enables diagnosis of concentration deficiency, identification of causal factors in vineyard or winery, and quality assessment recognizing dilution as a fault distinct from intentionally delicate style.
Taste_Dilution_Quality addresses the negative low-intensity end of the taste magnitude spectrum, implying quality deficiency rather than intentional lightness. This quality distinguishes from appropriately light styles (e.g., Muscadet, Vinho Verde) by focusing on the hollow, unsatisfying character of inadequate concentration versus the refreshing lightness of intentionally delicate wines. The class supports quality defect diagnosis and identification of causal factors requiring correction.
Geschmacksdünne
Geschmacksverdünnung
flavor dilution
gustatorische Schwäche
gustatory weakness
taste thinness
taste wateriness
wässriger Geschmack
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2: The Chemistry of Wine Stabilization and Treatments, 2nd Ed. John Wiley & Sons, Chichester, pp. 91-128 Van Leeuwen C et al. (2009) Influence of climate, soil, and cultivar on terroir. American Journal of Enology and Viticulture 60(4): 493-502
Taste Dilution Quality
Taste intensity evaluation comparing Pinot Noir from different yield levels reveals intensity rating of 7.8 for 35 hL/ha yield versus 5.2 for 65 hL/ha yield on equivalent vineyard sites, with the 50% higher intensity at lower yield corresponding to 35% higher extract concentration and 45% higher total phenolics, demonstrating the concentration effect of yield restriction on overall taste intensity that underlies the quality premium for limited production wines regardless of the specific balance among taste components.
Taste intensity quality is a taste quality that represents the overall magnitude of gustatory sensation integrating the combined intensities of all taste components, characterizing whether the taste experience is powerful and concentrated or light and delicate as a general impression independent of the balance among specific taste modalities. Within the BFO framework, this quality addresses the global intensity dimension of taste perception, distinguishing overall taste strength from the intensities of individual taste qualities and from balance relationships, while recognizing that total intensity provides information about fruit concentration and wine body not captured by component analysis. The ontological significance lies in taste intensity serving as a primary indicator of grape concentration and wine weight, where high intensity suggests concentrated raw material, full extraction, or reductive techniques preserving taste compounds, while low intensity may indicate dilution from high yields, rain, or oxidative loss of volatile taste modulators. The perceptual integration of overall taste intensity involves summation of inputs from multiple taste channels with potential synergistic or suppressive interactions affecting the combined impression, creating a gestalt intensity perception that reflects but does not simply sum component intensities. The compositional determinants of taste intensity include concentrations of all taste-active compounds (sugars, acids, phenolics, minerals, amino acids), with higher concentrations generally producing higher intensity though matrix effects and component interactions modulate the relationship. The style dimension of taste intensity distinguishes light-bodied from full-bodied wines, with appropriate intensity depending on variety, region, and intended style rather than higher intensity always indicating higher quality. Understanding taste intensity quality enables assessment of grape concentration and extraction success, prediction of wine body and weight perception, and quality evaluation in the context of style expectations where intensity appropriateness varies.
Taste_Intensity_Quality addresses the overall magnitude of taste perception independent of which specific tastes contribute. This quality serves as parent for Taste_Concentration_Quality and Taste_Dilution_Quality which represent the extremes of the intensity continuum. Intensity assessment requires calibration for style context, as appropriately intense Beaujolais differs from appropriately intense Barolo, with quality depending on intensity matching style expectations.
Geschmacksintensität
Geschmackskraft
Geschmacksstärke
flavor intensity
gustatorische Magnitude
gustatory magnitude
overall taste strength
taste power
Noble AC, Shannon M (1987) Profiling Zinfandel wines by sensory and chemical analyses. American Journal of Enology and Viticulture 38(1): 1-5 Heymann H, Noble AC (1987) Descriptive analysis of commercial Cabernet Sauvignon wines from California. American Journal of Enology and Viticulture 38(1): 41-44
Taste Intensity Quality
Taste persistence measurement in Burgundy Premier Cru versus village-level Pinot Noir using trained panel time-intensity evaluation reveals integrated taste persistence of 45±8 seconds for Premier Cru declining through pleasant evolution of fruit and mineral notes, versus 22±5 seconds for village wine with simpler fade, with the 100% longer persistence corresponding to 40% higher extract and justifying the quality tier classification that relies heavily on finish character as a distinguishing criterion.
Taste persistence quality is a taste quality that represents the duration and evolution of gustatory sensation following swallowing or expectoration, characterizing the temporal extension of taste perception that contributes to finish quality and overall wine complexity assessment. Within the BFO framework, this quality addresses the temporal dimension of taste experience as an integrated phenomenon, distinguishing overall taste persistence from the persistence of individual taste modalities (sweetness, sourness, bitterness) while recognizing that the combined persistence creates the finish impression more important for quality judgment than any single component's duration. The ontological significance lies in taste persistence serving as a primary quality indicator across wine styles, where extended persistence suggests concentration, complexity, and age-worthiness, while abbreviated persistence indicates simplicity or dilution, with the duration and evolution of finish often differentiating quality tiers more reliably than initial taste intensity. The physiological basis for taste persistence involves continued receptor stimulation from residual compounds in saliva and oral mucosa, sustained neural signaling following stimulus removal, and central processing that extends conscious perception beyond peripheral input, with different taste modalities showing characteristic persistence patterns affecting the integrated finish impression. The compositional determinants of taste persistence include the concentration and binding affinity of taste-active compounds, with higher concentrations and slower receptor dissociation producing longer persistence, and the complexity of the compound mixture creating the evolving character of extended finishes where different components fade at different rates. The qualitative character of persistence matters alongside duration, with harmonious evolution through the finish indicating quality while persistence of unpleasant elements (harsh bitterness, aggressive acidity) detracting from quality regardless of duration. Understanding taste persistence quality enables assessment of finish as a distinct quality dimension, prediction of persistence from compositional analysis, and wine quality evaluation incorporating the temporal trajectory of taste perception.
Taste_Persistence_Quality serves as the parent class for Finish_Length_Quality and Aftertaste_Quality, addressing the temporal extension of taste perception. This quality represents the integrated persistence of all taste components, complementing the individual persistence qualities defined for sweetness, sourness, and bitterness. Persistence assessment requires time-intensity methodology for quantitative measurement, with qualitative description of evolution character equally important for quality judgment.
Geschmacksdauer
Geschmackspersistenz
Nachhaltigkeit des Geschmacks
aftertaste length
finish duration
gustatory persistence
taste length
Noble AC (1998) Why do wines taste bitter and feel astringent? In: Waterhouse AL, Ebeler SE (eds) Chemistry of Wine Flavor. ACS Symposium Series 714, American Chemical Society, Washington DC, pp. 156-165 Peynaud E, Blouin J (1996) The Taste of Wine: The Art and Science of Wine Appreciation, 2nd Ed. John Wiley & Sons, New York, pp. 92-118
Taste Persistence Quality
Taste quality assessment of Riesling at potential harvest dates reveals sugar-acid balance shifting from 180 g/L sugar with 12 g/L acidity (sugar/acid ratio 15) in week 38 to 210 g/L sugar with 8 g/L acidity (ratio 26) in week 41, with sensory panel evaluation confirming the ratio of 18-20 as optimal for Kabinett style where moderate sweetness perception complements bright acidity, guiding harvest timing for the balance defining this wine category.
Taste quality is a sensory quality that represents the gustatory perception arising from chemical stimulation of taste receptor cells in the oral cavity, encompassing the basic taste modalities of sweetness, sourness, bitterness, saltiness, and umami that together characterize the sapid properties of grape berries and wine. Within the BFO framework, this quality specifically addresses the chemosensory perception mediated by taste buds on the tongue and palate, distinguishing gustation from olfaction (aroma), somesthesis (mouthfeel), and other sensory modalities while recognizing the profound interactions among these channels in integrated flavor perception. The ontological significance lies in taste providing the fundamental framework for evaluating grape maturity and wine balance, where the sweetness-acidity relationship defines the primary taste structure, bitterness and umami contribute secondary dimensions, and the integrated taste profile determines perceived quality and style classification. The physiological basis of taste involves specific receptor mechanisms for each basic taste, with sweetness detected through T1R2/T1R3 heterodimeric receptors responding to sugars and other sweet compounds, sourness through ion channels responding to protons, and bitterness through approximately 25 T2R receptor types detecting diverse bitter compounds including grape-derived phenolics. Taste perception in grape evaluation integrates absolute intensity of each basic taste with their temporal dynamics (onset, peak, persistence) and mutual interactions, where sweetness suppresses bitterness and sourness perception while acidity enhances freshness and can mask excessive sweetness. The compositional basis for grape taste quality includes sugars (primarily glucose and fructose) for sweetness, organic acids (tartaric, malic) for sourness, phenolic compounds (tannins, flavonoids) for bitterness, and amino acids for potential umami contribution, with the balance among these compounds determining the overall taste profile. Understanding taste quality enables prediction of sensory outcomes from compositional analysis, optimization of harvest timing for taste balance, and winemaking adjustment to achieve target taste profiles.
Taste_Quality addresses gustatory perception specifically, distinguishing from Aroma_Quality (olfaction) and Mouthfeel_Quality (somesthesis) while recognizing these modalities interact in flavor perception. This class serves as parent for Basic_Taste_Quality and derived taste perception qualities. The distinction between taste and flavor is maintained, where taste refers to the five basic modalities while flavor integrates taste with retronasal aroma and trigeminal sensations.
Geschmacksempfindung
Geschmacksqualität
Schmeckqualität
gustation quality
gustatorische Qualität
gustatory quality
sapid quality
taste sensation
Breslin PAS, Spector AC (2008) Mammalian taste perception. Current Biology 18(4): R148-R155 Hufnagel JC, Hofmann T (2008) Orosensory-directed identification of astringent mouthfeel and bitter-tasting compounds in red wine. Journal of Agricultural and Food Chemistry 56(4): 1376-1386
Taste Quality
TDN precursor concentration varies dramatically among cultivars, with Riesling showing uniquely high accumulation (100-400 µg/L glycosidic precursors) compared to most other V. vinifera cultivars (<50 µg/L). TDN forms through acid-catalyzed rearrangement of carotenoid-derived precursors, primarily megastigma-4,6,8-triene glycosides, with release accelerating at wine pH during bottle aging. Free TDN in young Riesling wines typically ranges from 1-10 µg/L, increasing to 30-200 µg/L after 5-10 years of bottle aging. The sensory threshold of 2-20 µg/L (depending on wine matrix) means TDN transitions from subliminal in young wines to dominant in aged examples. High sun exposure and warm ripening temperatures increase TDN precursor accumulation by 50-200%, explaining higher petrol character in warm-climate Riesling. Other cultivars showing moderate TDN potential include Semillon, Chenin Blanc, and Silvaner.
A norisoprenoid accumulation disposition that is the inherited capacity of a Vitis vinifera plant to accumulate glycosidic precursors of 1,1,6-trimethyl-1,2-dihydronaphthalene (TDN) in grape berries, with genotype-specific concentrations determining the potential for development of the characteristic kerosene/petrol aroma that defines aged Riesling and related wines.
TDN represents the defining aged character of Riesling, with the kerosene/petrol aroma being considered either highly desirable (aged Riesling connoisseurs) or a fault (consumers preferring fruity young wines). The uniquely high TDN potential of Riesling reflects cultivar-specific carotenoid composition and degradation patterns not fully understood mechanistically. Viticultural management targeting moderate sun exposure and controlled water stress can modulate TDN precursor accumulation, enabling some producer control over this distinctive aroma development trajectory.
Kerosin-Aroma-Disposition
Petrolnote-Potential
TDN potential
TDN-Vorläufer-Akkumulation
kerosene note disposition
petrol aroma precursor
Sacks GL et al. (2012) Origins of TDN in Riesling. In: Ebeler SB et al. (eds) Progress in Authentication of Food and Wine, ACS Symposium Series pp.: 255-271 Marais J et al. (1992) Effect of sunlight and shade on norisoprenoid levels in maturing Weisser Riesling and Chenin blanc grapes and Weisser Riesling wines. South African Journal of Enology and Viticulture 13:23-32
TDN Precursor Accumulation Disposition
TDN precursor quality in Clare Valley Riesling from sun-exposed sites generates the accelerated petrol development characteristic of warm-climate Australian Riesling, with precursor concentrations supporting TDN levels of 10-25 μg/L within 3-5 years of vintage compared to 10+ years required for equivalent TDN in cooler Mosel vineyards this rapid TDN evolution creates the distinctively earlier-developing petrol character of Australian Riesling that divides consumers between those appreciating the kerosene complexity and those preferring the slower-developing subtlety of cool-climate styles.
TDN_Precursor_Quality represents the chemical quality characterizing the concentration of precursor compounds for 1,1,6-trimethyl-1,2-dihydronaphthalene (TDN) in grape berries, capturing the carotenoid-derived intermediates that generate the distinctive petrol, kerosene, and hydrocarbon aromas characteristic of aged Riesling. TDN is formed through acid-catalyzed cyclization and aromatization of carotenoid degradation products, with multiple precursors including lutein-derived intermediates and various glycosidic conjugates potentially contributing to the TDN pool during wine aging. Unlike most grape aroma compounds that decline during aging, TDN accumulates progressively in bottle, increasing from typically undetectable levels in young wines (less than 4 μg/L threshold) to 15-50 μg/L in decade-old Riesling, with exceptional aged examples exceeding 100 μg/L. The sensory threshold of TDN is approximately 2-20 μg/L depending on wine matrix, making its accumulation cross from subthreshold to distinctly perceptible during the aging window. The petrol character from TDN is considered a positive quality marker in aged Riesling, contributing to the complex, developed bouquet that distinguishes great aged examples, though excessive TDN can dominate the aromatic profile. TDN precursor concentration in grapes depends on variety (Riesling accumulates highest levels among commercial varieties), sun exposure (increased light increases precursor formation), and vine age (older vines may produce higher precursor levels). The rate of TDN formation during aging depends on wine pH (lower pH accelerates formation), storage temperature (higher temperature increases rate), and the initial precursor concentration. Managing TDN development requires balancing these factors to achieve desirable aged character without excessive petrol dominance.
TDN_Precursor_Quality addresses the precursor pool for the signature aging compound of Riesling. The precursor concept is particularly important for TDN because the compound is essentially absent in young wine—its presence depends entirely on post-bottling formation from precursors. The multiple pathways to TDN formation (various carotenoid degradation routes) complicate direct precursor measurement total norisoprenoid glycoside potential often serves as a proxy. The varietal specificity of TDN formation (predominantly Riesling, also significant in Semillon and some other varieties) reflects genetic differences in carotenoid metabolism and precursor accumulation.
1,1,6-Trimethyl-1,2-dihydronaphthalin-Vorläufer
1,1,6-trimethyl-1,2-dihydronaphthalene precursor
Kerosin-Aromapotential
Petroleumverbindungs-Vorläuferqualität
TDN precursor content
TDN-Vorläufergehalt
kerosene aroma potential
petrol compound precursor quality
Simpson RF (1978) Aroma and Compositional Changes in Wine with Oxidation, Storage and Ageing. Vitis 17(3): 274-287 Winterhalter P et al. (1990) Identification of a Carotenoid-Derived Aroma Precursor from Riesling Wine. Vitis 29(4): 251-258
TDN Precursor Quality
Welschriesling vines in the University of Pécs teaching vineyard bearing a teaching collection role are maintained as part of a 120-cultivar didactic collection where undergraduate biology and viticulture students perform weekly phenological observations recording BBCH stages from budburst through leaf fall, practice ampelographic description using OIV descriptors for leaf shape (OIV 067-068), berry morphology (OIV 220-241), and cluster architecture (OIV 202-209), conduct comparative must analysis across 15 cultivars at harvest, and perform micro-vinification of 10 kg lots for introductory sensory evaluation exercises distinguishing varietal aromatic profiles.
A research and education role that inheres in a grapevine maintained within a curated vineyard collection whose primary purpose is the instruction of students in ampelography, viticulture, oenology, and related disciplines, providing living pedagogical material for hands-on learning of cultivar identification, phenological observation, viticultural technique, and sensory evaluation.
Teaching_Collection_Role captures the pedagogical function of vineyard plantings specifically maintained for educational purposes. The role requires curatorial attention to cultivar diversity, labeling accuracy, and vine health sufficient to provide representative teaching material: teaching collections typically prioritize breadth of cultivar representation over production optimization, including both commercially important and historically or ampelographically instructive cultivars. The role is distinct from Research_And_Education_Role's other subclasses in that the primary purpose is knowledge transmission to students rather than knowledge generation through research or preservation through conservation.
Lehr- und Anschauungs-Rolle
Lehrsammlung-Rolle
didactic vineyard role
didaktische Weinberg-Rolle
educational collection role
teaching vineyard role
OIV (2009) 2nd Edition of the OIV Descriptor List for Grape Varieties and Vitis Species Huglin P and Schneider C (1998) Biologie et écologie de la vigne, 2nd Ed, Lavoisier pp: 1-35 Galet P (2000) Dictionnaire encyclopédique des cépages, Hachette pp: 1-22
Teaching Collection Role
Technological maturity monitoring for Riesling table wine production reveals the compositional progression guiding harvest decisions, with weekly sampling showing sugar increasing from 18 to 22 Brix over four weeks while pH rises from 2.95 to 3.20 and titratable acidity declines from 12 to 8 g/L, with the target window of 20-21 Brix, pH 3.05-3.15, and TA 9-10 g/L representing the optimal technological maturity for the intended dry wine style, achievable during a 5-7 day harvest window before continued ripening moves composition beyond target parameters, demonstrating how technological maturity assessment enables precise harvest timing for specific wine style objectives.
Technological maturity quality is a maturity quality that represents the achievement of compositional targets relevant for intended processing and product objectives, characterizing the chemical constitution of berries in terms of the parameters that determine winemaking potential including sugar concentration (potential alcohol), acidity (pH, titratable acidity), and the balance among compositional elements that affects fermentation behavior and wine style possibilities. Within the BFO framework, this quality specifically addresses the compositional dimension of maturity, distinguishing chemical analysis-based assessment from biological developmental completion while recognizing that technological maturity represents the translation of physiological development into measurable parameters directly relevant for winemaking decisions and wine quality outcomes. The ontological significance lies in technological maturity providing the quantitative basis for harvest decisions through measurable parameters (Brix, pH, TA) that predict wine composition (alcohol, acidity balance) and fermentation behavior, enabling rational harvest timing based on compositional targets appropriate for intended wine styles rather than calendar dates or visual assessment alone. The primary parameters of technological maturity assessment include sugar concentration (Brix, °Baumé, or potential alcohol indicating fermentation potential), pH (affecting microbial stability, color expression, sensory perception), titratable acidity (tartaric + malic acid equivalents affecting perceived freshness), and the sugar/acid ratio or similar balance indices that capture the interaction between these primary parameters affecting overall wine balance. The wine style dependency of technological maturity targets creates different optima for different objectives, with sparkling wine production requiring lower sugar (18-20 Brix) and higher acidity, table wine production targeting moderate levels (22-26 Brix depending on style), and late harvest or fortified wines potentially targeting much higher sugar concentrations, such that "technological maturity" is defined relative to purpose rather than absolute standards. The climate and vintage effects on technological maturity show substantial variation in the achievability of target compositions, with cool vintages potentially struggling to achieve adequate sugar while maintaining excessive acid, and warm vintages potentially reaching high sugar before achieving favorable acid decline, creating the vintage-specific challenges that harvest decisions must address. Understanding technological maturity quality enables rational harvest timing based on compositional analysis, wine style planning based on achievable maturity parameters, and vintage characterization based on the technological maturity profiles achieved.
Technological_Maturity_Quality serves as the parent class for compositional maturity parameters (Sugar, Acid, Must Composition). This quality addresses the measurable chemical targets for winemaking decisions. The class supports harvest timing and style planning through technological maturity characterization.
Kelterreife
Mostkompositionsreife
chemical maturity
chemische Reife
compositional maturity
industrial maturity
technologische Reife
winemaking maturity
Ribéreau-Gayon P et al. (2006) Handbook of Enology, Volume 1: The Microbiology of Wine and Vinifications, 2nd Ed. John Wiley & Sons, Chichester, pp. 193-230 Jackson DI, Lombard PB (1993) Environmental and management practices affecting grape composition and wine quality - a review. American Journal of Enology and Viticulture 44(4): 409-430
Technological Maturity Quality
The teeth shape description of Chenin Blanc documents predominantly convex-convex teeth (OIV 076:1) — both the distal (toward the tooth apex) and proximal (toward the tooth base) sides of each marginal tooth are smoothly curved outward, creating a scalloped, rounded margin reminiscent of a series of small arcs joined at shallow notches between teeth. This rounded tooth form is shared with Chenin Blanc's identified parent Savagnin (also OIV 076:1), illustrating the heritability of tooth shape across generations, and contrasts with the sharply angular rectilinear teeth of the Loire Valley neighbor Sauvignon Blanc (OIV 076:2), the asymmetric convex-concave teeth of Cabernet Franc (OIV 076:3), and the mucronate teeth of Sémillon (OIV 076:2 with conspicuous mucro at the tooth apex extending 1-2 mm beyond the tooth body).
A leaf margin description component that records the geometric form of individual marginal teeth according to OIV descriptor 076, classifying each tooth by the curvature of its two lateral sides — both sides convex (state 1, rounded scalloped teeth), both sides rectilinear (state 2, angular saw-teeth), one side convex and one side concave (state 3, asymmetric pointed teeth), or a mixture of forms along the margin (state 4) — providing a character of high genetic stability and diagnostic power based on the developmental geometry of marginal growth patterning in the leaf blade.
Teeth_Shape_Description documents a character that exemplifies the ampelographic principle that small-scale morphological features can be as diagnostically powerful as large-scale ones. Tooth shape is determined by the growth dynamics at individual marginal vein termini during leaf expansion — the curvature of each tooth side reflects the relative expansion rates of the adaxial and abaxial tissue flanking the marginal vein, with symmetric expansion producing convex-convex teeth and differential expansion producing the asymmetric convex-concave form: these growth patterns are controlled by localized gene expression differences at each marginal primordium, producing the cultivar-specific tooth geometry that is remarkably consistent across leaves within a vine and across vines within a cultivar. The OIV classification into four categories (convex-convex, rectilinear-rectilinear, convex-concave, mixed) represents a simplification of the continuous morphological variation — some cultivars show teeth that are intermediate between categories or display different tooth forms in different regions of the blade (e.g., convex teeth on lobe tips but rectilinear teeth along sinus margins), requiring the observer to determine the predominant form following the OIV protocol of scoring the most frequently observed category across multiple leaves.
Serrationsform-Charakterisierung
Zahnform-Beschreibung
Zähnungsgeometrie
dental geometry record
serration shape characterization
tooth form classification
Teeth Shape Description
The teeth size description of Mourvèdre documents long marginal teeth (OIV 077:7) measuring 8-12 mm from notch to tooth apex — these prominent teeth, combined with Mourvèdre's rectilinear tooth form, create a conspicuously saw-toothed margin visible from several meters distance in the vineyard, contrasting with the short, subtle teeth of Grenache (OIV 077:3, teeth 3-5 mm, barely projecting from the blade margin) and the medium teeth of Carignan (OIV 077:5, teeth 5-8 mm). Tooth size correlates loosely with overall leaf size (larger leaves tend to have longer teeth) but maintains a cultivar-specific character — Mourvèdre's teeth are proportionally longer even on small leaves from low-vigor shoots, while Grenache's teeth remain proportionally short even on the largest leaves.
A leaf margin description component that records the dimensions of marginal teeth according to OIV descriptor 077 (length of teeth), scored on a scale from very short (state 1) through short, medium, long, to very long (state 9), characterizing the prominence of the leaf margin serration that contributes to the overall visual texture of the leaf edge and provides additional cultivar discrimination within groups sharing the same tooth shape.
Teeth_Size_Description captures the dimensional component of marginal serration that provides cultivar discrimination independently of tooth shape. Tooth length (the distance from the deepest point of the notch between adjacent teeth to the tooth apex) varies by a factor of 3-4 across V. vinifera cultivars, from the inconspicuous 2-3 mm serrations of minimally toothed cultivars (e.g., some Muscat types) to the prominent 10-15 mm teeth of strongly serrated cultivars (e.g., Mourvèdre, Tannat): this variation reflects differences in the extent of marginal growth at each vein terminus during leaf expansion, with longer teeth representing greater localized growth projection beyond the general blade surface. The relationship between tooth size and leaf size complicates the use of absolute tooth length as a cultivar-diagnostic character — the OIV scoring system addresses this partially by defining score categories based on absolute length ranges, but the tooth length relative to blade length or blade area would provide a more genetically stable metric. Tooth size is scored on the terminal lobe teeth where serration is most uniformly expressed, avoiding the often smaller and more variable teeth near the petiolar sinus.
Serrationsgröße-Charakterisierung
Zahngröße-Beschreibung
Zähnungslänge
dental dimension characterization
serration size assessment
tooth length record
Teeth Size Description
Alicante Bouschet bears a Teinturier_Scion_Role throughout southern France, Portugal's Alentejo, and California, providing intense color (free-run juice achieving 2000-4000 mg/L anthocyanins versus 100-300 mg/L for standard red varieties) for blending with lighter cultivars or producing deeply pigmented single-varietal wines. Saperavi serves in a Teinturier_Scion_Role as Georgia's signature red variety, creating the intensely colored, tannic wines (both European-style and traditional qvevri) that define Georgian viticulture, with flesh pigmentation contributing to extraction even in brief-contact rosé styles. Dunkelfelder (Färbertraube) bears a Teinturier_Scion_Role in German viticulture, used for color enhancement in blends and increasingly for single-varietal production emphasizing deep color and soft tannins.
A red wine scion role that inheres in a grafted scion cultivar by virtue of its genetically determined capacity to produce berries with pigmented flesh (pulp), in addition to pigmented skin, enabling intense color extraction and contribution to blended wines. Teinturier cultivars (from French "teindre" = to dye) differ from typical red wine grapes, which have colorless flesh regardless of skin color, by accumulating anthocyanins throughout the mesocarp tissue, yielding deeply pigmented juice upon crushing without maceration. This unique trait results from mutations affecting anthocyanin biosynthesis regulation that extend pigment accumulation beyond the hypodermal cell layers into the inner flesh. Teinturier cultivars produce juice with color intensities 5-20 times higher than standard red varieties, making them valuable for color enhancement in blends, production of deeply colored wines from high-yield viticulture, and as crossing parents for breeding intensely pigmented new varieties. The most important teinturier cultivar globally is Alicante Bouschet (Petit Bouschet × Grenache), along with regional teinturiers including Saperavi (Georgia), Dunkelfelder (Germany), and various Bouschet crosses.
Teinturier_Scion_Role captures a distinct functional category based on the genetically determined flesh pigmentation trait rather than wine style or quality tier. Teinturiers have historically been viewed primarily as blending grapes for color correction, but increasing interest in natural winemaking (avoiding oenological color additives) and consumer preference for deeply colored reds has elevated some teinturiers to single-varietal production. Alicante Bouschet is the world's most planted teinturier (approximately 50,000 ha globally) and has been extensively used as a breeding parent to introgress the flesh pigmentation trait into new varieties. Saperavi represents a distinct teinturier lineage indigenous to the Caucasus with no relationship to the Bouschet breeding program.
Deckrotweinsorte
Färbertraube
Teinturier-Sorte
colored-flesh variety
dyer grape
red-fleshed grape
rotfleischige Rebsorte
teinturier variety
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane pp.: 34-35, 877-878 Castellarin SD et al. (2011) Colour variation in red grapevines: the origin of the teinturier mutation, Vitis 50: 85-92 This P et al. (2007) Characterization of the Teinturier gene controlling anthocyanin biosynthesis in grapevine flesh, Australian Journal of Grape and Wine Research 13: 141-150 Boss PK et al. (1996) Expression of anthocyanin biosynthesis pathway genes in red and white grapes, Plant Molecular Biology 32: 565-569
Teinturier Scion Role
In Riesling vines in the Mosel Valley, temperature acclimation progresses through autumn with initial acclimation at 10-15°C during October increasing bud cold hardiness from -5°C to -12°C, secondary hardening during November as temperatures fall to 0-5°C further increasing hardiness to -18°C, and maximum hardiness of -22°C achieved by late December following exposure to temperatures of -5 to -10°C during early winter.
Temperature acclimation in Vitis vinifera encompasses the physiological processes through which grapevines respond to declining autumn temperatures by progressively developing increased tolerance to freezing stress, representing a critical component of dormancy induction that determines the vine's capacity to survive winter cold. This adaptive response involves coordinated changes in cellular composition, gene expression, and metabolism that progressively enhance the ability of living tissues to withstand ice formation and the associated cellular stresses of dehydration, mechanical damage, and membrane disruption. The acclimation process proceeds in stages, with initial acclimation occurring at cool but non-freezing temperatures between 10-15°C involving transcriptional activation of cold-responsive genes, followed by further hardening at temperatures approaching freezing that induces additional protective mechanisms and maximum cold hardiness development at temperatures below freezing. The cellular changes during temperature acclimation include accumulation of compatible solutes including proline, raffinose family oligosaccharides, and other cryoprotective molecules that stabilize membranes and proteins during freezing, synthesis of dehydrin proteins that protect cellular structures during freeze-induced dehydration, and modifications to membrane lipid composition increasing unsaturated fatty acid content that maintains membrane fluidity at low temperatures. Gene expression changes during cold acclimation involve activation of CBF/DREB transcription factors that regulate large sets of cold-responsive genes, increased expression of genes encoding protective proteins and biosynthetic enzymes for cryoprotectants, and repression of genes associated with active growth. The interaction between temperature acclimation and photoperiod response is complex, with short days enhancing cold acclimation capacity even at warm temperatures and low temperatures being more effective at inducing hardiness after short-day pre-conditioning than before. Maximum cold hardiness in fully acclimated grapevine tissues can reach -20 to -25°C for hardy cultivars, compared to only -3 to -5°C for non-acclimated tissues, illustrating the dramatic protective effects of the acclimation process. The rate and extent of cold acclimation varies substantially among cultivars, contributing to differences in geographic adaptation and winter survival.
Temperature_Acclimation represents a major component process within Dormancy_Induction parallel to Photoperiod_Response, specifically addressing the thermal induction of cold hardiness. Distinguished from Photoperiod_Response which addresses daylength signals, though both processes interact synergistically during dormancy induction. Subdivides into Cold_Acclimation_Start, Freeze_Tolerance_Development, and Metabolic_Adjustment as component processes. Proceeds in stages requiring progressively lower temperatures. Maximum hardiness varies with cultivar from approximately -15 to -30°C in Vitis vinifera.
Kälteabhärtung
Kältetoleranz-Entwicklung
Temperatur-Akklimatisierung
cold acclimation
freezing tolerance development
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10: 201-235 Mills LJ, Ferguson JC, Keller M (2006) Cold-hardiness evaluation of grapevine buds and cane tissues. American Journal of Enology and Viticulture 57: 194-200 Xin Z, Browse J (2000) Cold comfort farm: the acclimation of plants to freezing temperatures. Plant Cell and Environment 23: 893-902
Temperature Acclimation
The temperature data for the Wachau wine region (Lower Austria, 48.4°N) documents the comprehensive thermal characterization of this Danube Valley viticultural landscape — mean annual temperature 9.8°C, growing season (April-October) mean 16.5°C, January mean -0.5°C (continental cold winter), July mean 20.5°C, annual temperature amplitude 21°C (strongly continental), mean diurnal range 10-12°C during the ripening period (August-October), growing degree-days base 10°C of 1,350-1,550, frost-free period approximately 180 days (mid-April to mid-October), extreme minimum temperature -20 to -25°C (occurring approximately once per decade with potential for trunk and cordon damage), and extreme maximum temperature 35-38°C (occurring 5-10 days per year, concentrated in July, with potential for sunburn damage and photosynthetic shutdown). This thermal profile defines a continental viticultural climate with high seasonal and diurnal amplitude — the warm summers provide sufficient heat for Grüner Veltliner and Riesling maturation, the wide diurnal range during ripening preserves acidity and promotes aromatic complexity, and the cold winters ensure complete vine dormancy and adequate chilling for uniform budburst.
A climate data component that documents the thermal regime of the vineyard environment — recording the annual and seasonal temperature patterns, extremes, variability, and derived thermal indices that together characterize the temperature environment controlling vine phenological development rate, photosynthetic efficiency, respiration losses, frost damage risk, and the temperature-dependent biochemical pathways of sugar accumulation, organic acid metabolism, anthocyanin synthesis, and aroma precursor formation that determine fruit composition at harvest.
Temperature_Data is the parent class for the suite of thermal measurements and indices that together provide the most complete single-variable characterization of the viticultural climate. Temperature affects virtually every physiological process in the grapevine — photosynthesis (optimum 25-30°C, declining above 35°C and below 10°C), respiration (increasing exponentially with temperature, approximately doubling per 10°C increase, consuming photosynthetic products at accelerating rates in warm climates), sugar accumulation (net positive only when photosynthesis exceeds respiration, a balance favored by warm days and cool nights), organic acid metabolism (malic acid respiration rate increasing sharply above 30°C, explaining the rapid acid loss in hot climates), anthocyanin synthesis (optimum 17-26°C, inhibited above 35°C through enzyme degradation and pigment instability), and aroma compound metabolism (terpene synthesis promoted by moderate temperatures, methoxypyrazine degradation accelerated by heat): the compound effects of temperature on these multiple pathways mean that the temperature regime during each phenological phase (budburst, flowering, véraison, ripening) independently influences different aspects of fruit composition, making the temporal distribution of temperature as important as the seasonal mean. The VVO documents temperature through a hierarchy of parameters — annual means, growing season means, diurnal amplitude, heat summation, frost risk, and extreme events — that together capture the temporal and distributional characteristics of the thermal environment relevant to viticulture.
Temperaturdaten
Weinberg-Temperatur-Dokumentation
temperature environment documentation
thermal regime characterization
thermische Regime-Charakterisierung
vineyard temperature record
Gladstones J (1992) Viticulture and Environment, Winetitles Jones GV et al. (2005) Climate change and global wine quality, Climatic Change 73: 319-343 Jackson DI and Lombard PB (1993) Environmental and management practices affecting grape composition and wine quality — a review, American Journal of Enology and Viticulture 44: 409-430 Coombe BG and McCarthy MG (2000) Dynamics of grape berry growth, Australian Journal of Grape and Wine Research 6: 131-135
Temperature Data
Temperature differential monitoring in Margaret River Chardonnay vineyard reveals canopy-to-air differential shifting from -4.2°C (canopy cooler than air) in morning with open stomata and active transpiration to +1.8°C (canopy warmer than air) by afternoon as water stress induces stomatal closure, with the crossover point from negative to positive differential providing a practical irrigation trigger more responsive than soil moisture sensors in the heterogeneous soils of this region.
Temperature differential quality is a temperature quality that represents the difference between temperatures measured at two defined reference points, characterizing the thermal gradient driving heat transfer or indicating the departure of plant tissue temperature from environmental reference conditions. Within the BFO framework, this quality captures the relational aspect of temperature where the difference between two measurements provides information not available from either measurement alone, enabling inference about heat transfer rates, plant physiological status, and microclimate modification by the plant canopy. The ontological significance lies in temperature differentials serving as diagnostic indicators, where the sign and magnitude of the difference between tissue and environment temperatures reveals the dominant processes in the energy balance including transpirational cooling when tissue temperature is below ambient, radiant heating when tissue temperature exceeds ambient, and thermal equilibrium when temperatures match. Temperature differential measurement requires simultaneous or near-simultaneous determination of temperatures at both reference points using comparable methodologies to minimize systematic bias, with the differential calculated as the algebraic difference between defined locations. The physiological interpretation of temperature differentials depends on the reference points selected, where canopy-to-air differential indicates plant water status through transpirational cooling capacity, while berry-to-air differential reflects fruit exposure and radiant heating intensity. Temperature differentials drive heat transfer through the vine system, with thermal gradients determining conductive heat flow from warm to cool tissues, and differentials between plant surfaces and air determining convective heat exchange rates. Understanding temperature differential quality enables development of thermal stress indices, interpretation of plant water status from remote thermal sensing, and prediction of heat damage risk from the difference between tissue and damage threshold temperatures.
Temperature_Differential_Quality serves as the parent class for specific temperature differential types (Canopy-Air, Berry-Ambient), establishing the ontological principle that temperature differences provide diagnostic information beyond absolute temperatures. This class supports thermal stress index development and remote sensing interpretation. The reference points for differential calculation must be explicitly defined as the same absolute temperature difference has different interpretations depending on what is being compared.
Temperaturdifferenz
Temperaturgradient
Temperaturunterschied
delta T
temperature difference
thermal differential
thermal gradient
thermische Differenz
Jackson RD et al. (1981) Canopy temperature as a crop water stress indicator. Water Resources Research 17(4): 1133-1138 Idso SB (1982) Non-water-stressed baselines: a key to measuring and interpreting plant water stress. Agricultural Meteorology 27(1-2): 59-70
Temperature Differential Quality
In Pinot Gris vines in Oregon's Willamette Valley, temperature limitation maintains ecodormancy from early February through late March, with February mean temperatures of 6-8°C providing minimal growing degree day accumulation above the 10°C base temperature, March warming gradually increasing GDD accumulation to approximately 80 units by month end, and budbreak occurring in early April when accumulated GDD reach the cultivar threshold of approximately 120-150 units.
Temperature limitation in Vitis vinifera represents the primary environmental constraint maintaining ecodormancy, whereby ambient temperatures below the threshold required for growth-supporting metabolic and developmental processes prevent budbreak initiation in growth-competent buds during late winter and early spring. This temperature-based control mechanism operates through the fundamental temperature-dependence of biological reaction rates, with temperatures below approximately 10°C being insufficient to support the sustained metabolic activity required for cell division, expansion, and the coordinated developmental processes of budbreak. The biochemical basis of temperature limitation involves the temperature-dependence of enzyme catalysis according to Arrhenius kinetics, with reaction rates declining exponentially as temperature decreases below optima, eventually reaching levels insufficient to sustain net biosynthesis and growth even when all regulatory constraints have been removed. The cellular processes limited by low temperature include mitochondrial respiration required for ATP production, protein synthesis requiring coordinated ribosome function, cell wall biosynthesis involving multiple enzymatic steps, and membrane biogenesis necessary for cell expansion, all of which must proceed at adequate rates for growth to occur. The concept of base temperature defines the threshold below which these processes cannot support growth, with grapevine typically exhibiting a base temperature near 10°C though some variation exists among cultivars and the effective base temperature may differ for different developmental processes. The quantification of temperature limitation uses growing degree day accumulation above the base temperature, with each degree above threshold contributing to the cumulative heat exposure required for budbreak, typically 100-300 growing degree days depending on cultivar and environmental history. The release from temperature limitation occurs progressively as spring warming increases the frequency and duration of above-threshold temperatures, accumulating growing degree days until the cultivar-specific requirement is satisfied and budbreak commences. The non-linear nature of temperature responses means that a few warm days can contribute substantially to heat accumulation, while extended cool periods contribute little, making budbreak timing sensitive to temperature variability during late winter and early spring.
Temperature_Limitation represents the primary environmental constraint within Environmental_Constraint specifically addressing thermal control of ecodormancy. Distinguished from Moisture_Limitation which addresses water-related constraints. Operates through temperature-dependence of metabolic reaction rates. Base temperature approximately 10°C for grapevine. Quantified through growing degree day accumulation. Release progressive with spring warming. Cultivar heat requirements typically 100-300 GDD for budbreak.
Kältebeschränkung
Temperaturbeschränkung
temperaturbedingte Wachstumshemmung
temperature growth limitation
thermal growth inhibition
thermal limitation
thermische Limitation
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Parker AK et al (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17: 206-216 Moncur MW et al (1989) Base temperatures for budbreak and leaf appearance of grapevines. American Journal of Enology and Viticulture 40: 21-26
Temperature Limitation
Temperature perception quality assessment in high-alcohol Barossa Shiraz reveals both components of thermal perception, with actual serving temperature (18°C) providing appropriate warmth for full appreciation of complex aromatics, while 15.5% alcohol contributes substantial chemesthetic warmth (alcohol heat rating 6/10) that extends the finish and adds to perceived body, with the combined thermal impression creating the powerful, warming character expected in this style, though higher alcohol wines may show excessive heat that detracts from elegance.
Temperature perception quality is a tactile quality that represents the thermal sensation of wine during consumption, encompassing both the actual temperature of the wine and the chemesthetic thermal sensations induced by specific compounds (alcohol warmth, menthol cooling) that together characterize the temperature-related component of wine mouthfeel experience. Within the BFO framework, this quality addresses the thermal dimension of tactile perception, distinguishing temperature-related sensations from mechanical texture while recognizing that thermal perception involves both true thermoreceptor stimulation (detecting actual wine temperature) and chemesthetic pseudo-thermal sensations (chemicals activating thermal receptors without temperature change). The ontological significance lies in temperature perception affecting multiple aspects of wine quality assessment, where serving temperature influences volatile release and flavor perception, alcohol-derived warmth contributes to perceived body and finish character, and the overall thermal impression shapes the sensory experience from refreshing coolness through neutral to warming intensity. The physiological basis for temperature perception involves thermoreceptors in the oral mucosa detecting actual temperature changes and TRP (transient receptor potential) ion channels responding to chemical agonists, with TRPV1 channels (activated by capsaicin and alcohol) creating warmth sensation and TRPM8 channels (activated by menthol and certain terpenes) creating cooling sensation independent of actual temperature. The compositional determinants of thermal perception include alcohol concentration (primary source of warmth through TRPV1 activation), certain terpenes and terpenoids (potentially contributing cooling sensation), and the actual wine temperature as served (affecting both direct thermal sensation and all other sensory perception through temperature's effect on volatility, viscosity, and receptor sensitivity). The quality implications of temperature perception include the importance of appropriate serving temperature for style (cool for fresh whites, warmer for complex reds), the contribution of moderate alcohol warmth to body and finish, and the negative impact of excessive alcohol heat that creates burning sensation detracting from quality. Understanding temperature perception quality enables serving temperature optimization, alcohol balance assessment from warmth perception, and quality evaluation incorporating appropriate thermal character for wine style.
Temperature_Perception_Quality serves as the parent class for thermal sensation qualities (Cooling, Warming), addressing both actual temperature and chemesthetic pseudo-thermal sensations. This quality encompasses two distinct mechanisms - thermoreception (actual temperature) and TRP channel activation (chemical-induced thermal sensation). The class supports serving temperature guidance and alcohol balance assessment through thermal perception evaluation.
Temperaturmundgefühl
Temperaturwahrnehmung
Wärme-Kälte-Gefühl
heat-cool sensation
temperature mouthfeel
thermal perception
thermal sensation
thermische Empfindung
Green BG (1987) The sensitivity of the tongue to ethanol. Annals of the New York Academy of Sciences 510(1): 315-317 Pickering GJ et al. (2006) The influence of ethanol on the temporal perception of sweetness, sourness and flavour balance in white wine. American Journal of Enology and Viticulture 57(3): 306-312
Temperature Perception Quality
Temperature monitoring in Barossa Valley Shiraz vineyards reveals berry temperatures reaching 42°C during January heat events despite ambient air temperature of 38°C, with the 4°C elevation due to radiant heating triggering emergency irrigation for evaporative cooling to prevent the anthocyanin degradation and cooked fruit flavors that compromise wine quality when berry temperatures exceed 35°C for extended periods.
Temperature quality is a physical quality that inheres in grape vine tissues and structures and represents the thermal state characterizing the kinetic energy of molecular motion within plant material, determining reaction rates for enzymatic processes, membrane fluidity, protein stability, and the thermodynamic driving forces for water and solute transport throughout the vine system. Within the BFO framework, this quality serves as a fundamental physical parameter integrating environmental heat inputs from solar radiation, ambient air temperature, and soil thermal conditions with plant-mediated thermal regulation through transpirational cooling, tissue heat capacity, and metabolic heat generation. The ontological significance lies in temperature functioning as a master variable controlling virtually all physiological processes, where the exponential relationship between temperature and reaction rates (Q₁₀ typically 2-3 for biological processes) means that small temperature differences translate to substantial differences in metabolic activity, developmental rate, and quality compound synthesis or degradation. Grapevine tissues experience temperatures ranging from subzero during winter dormancy inducing cold hardiness responses to exceeding 45°C in sun-exposed berries during heat waves causing cellular damage and quality loss. Temperature measurement in viticultural contexts employs thermocouples, infrared thermometry, and thermal imaging, with methodology selection depending on whether point measurements, spatial distributions, or temporal dynamics are of interest. The distinction between tissue temperature and ambient air temperature is critical because radiant heating can elevate exposed tissue temperatures 10-15°C above air temperature while transpirational cooling can reduce leaf temperatures below ambient, meaning that air temperature alone inadequately characterizes the thermal environment experienced by the plant. Understanding temperature quality at multiple scales from cellular to canopy enables prediction of developmental timing, identification of thermal stress conditions, and optimization of management practices for quality outcomes.
Temperature_Quality serves as the parent class for tissue-specific temperature qualities and temperature differential qualities, establishing the ontological framework for thermal characterization at multiple organizational scales. This class addresses the instantaneous thermal state rather than accumulated thermal time (growing degree days) which represents integrated temperature over time. Measurement methodology significantly affects reported values, with infrared thermometry providing surface temperature while inserted probes measure internal tissue temperature.
Temperatur
Temperaturqualität
Wärmezustand
heat content
temperature status
thermal quality
thermal state
thermischer Zustand
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed. Academic Press, San Diego, pp. 275-312 Spayd SE et al. (2002) Separation of sunlight and temperature effects on the composition of Vitis vinifera cv. Merlot berries. American Journal of Enology and Viticulture 53(3): 171-182
Temperature Quality
Temperature response for budbreak-to-flowering interval ranges from 8-12 days earlier per 1°C warming in high-sensitivity cultivars (Riesling, Gewürztraminer) to 4-6 days in low-sensitivity cultivars (Grenache, Mourvèdre). Cabernet Sauvignon shows intermediate temperature response with veraison advancing 2.5 days per 1°C and harvest advancing 3.2 days per 1°C of seasonal warming. Non-linear temperature responses occur above threshold temperatures (approximately 35°C), where further warming may delay rather than advance phenology through heat stress effects. Temperature response coefficients derived from multi-year phenology records enable site-specific harvest date predictions with ±5-7 day accuracy.
A phenological plasticity disposition that is the inherited capacity of a Vitis vinifera plant to adjust developmental timing in response to temperature variation, with genotype-specific thermal sensitivity determining the magnitude of phenological shifts per unit change in ambient or accumulated temperature.
Temperature response disposition quantifies the thermal component of phenological plasticity, representing the primary driver of observed climate change impacts on grapevine phenology. High temperature response indicates strong coupling between vine development and ambient thermal conditions, enabling rapid adjustment to year-to-year variation but also magnifying long-term climate trends. Base temperature for development (typically 7-12°C) and optimal temperature range (25-30°C) show cultivar-specific variation that interacts with temperature response magnitude. Breeding for modified temperature response could enable climate change adaptation.
Temperaturreaktion
Wärmeempfindlichkeit
heat response trait
temperature-driven plasticity
thermal sensitivity
thermische Sensitivität
Parker AK et al. (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17:206-216 Webb LB et al. (2012) Observed trends in winegrape maturity in Australia. Global Change Biology 18:3055-3067
Temperature Response Disposition
Temperature response characterization in Syrah comparing cool-climate (Adelaide Hills) and warm-climate (Barossa Valley) sources reveals both genetic and acclimation effects on thermal physiology, with Barossa-adapted vines showing photosynthetic temperature optimum of 32°C versus 28°C for Adelaide Hills material, and respiration Q₁₀ of 2.0 versus 2.3, indicating that warm-climate adaptation involves both higher temperature optima and reduced respiratory temperature sensitivity, enabling more efficient carbon gain under the elevated temperatures of warm viticultural regions.
Temperature response quality is a physiological quality that represents the functional relationship between metabolic process rates and temperature, characterizing how photosynthesis, respiration, and other physiological processes change across the temperature range experienced by grapevines from chilling through optimal to heat stress conditions. Within the BFO framework, this quality addresses the thermal dependency dimension of physiological function, distinguishing the shape and parameters of temperature responses from absolute metabolic rates while recognizing that temperature profoundly affects virtually all physiological processes through direct effects on enzyme kinetics, membrane fluidity, and protein stability. The ontological significance lies in temperature response determining vine performance across the thermal environments of different viticultural regions and through the seasonal and diurnal temperature fluctuations within any region, where the temperature optima, sensitivity coefficients, and acclimation capacity together establish how effectively vines function under the thermal conditions they experience, with implications for regional suitability, climate change adaptation, and seasonal productivity patterns. The biochemical basis for temperature response includes enzyme kinetics (reaction rates increasing with temperature following Arrhenius relationship until denaturation), membrane properties (fluidity changing with temperature affecting transport and signaling), protein stability (denaturation at temperature extremes), and the differential temperature sensitivity of coupled processes (creating imbalances when processes with different temperature responses interact). The quantitative description of temperature response employs various models including Q₁₀ coefficients (fold change per 10°C), Arrhenius activation energies, and peaked functions for processes with temperature optima, with the appropriate model depending on the temperature range and process characteristics being described. The acclimation dimension of temperature response recognizes that the parameters of temperature response are not fixed but adjust with growth temperature, enabling plants to partially optimize function for their thermal environment through adjustments in membrane composition, enzyme content, and protective systems. Understanding temperature response quality enables prediction of metabolic rates across temperatures, assessment of thermal stress impacts, and evaluation of cultivar suitability for specific thermal environments.
Temperature_Response_Quality serves as the parent class for specific temperature response parameters (Q₁₀, Acclimation). This quality describes the thermal dependency of physiological processes that determines performance across temperature environments. The class supports regional suitability assessment and climate adaptation evaluation through temperature response characterization.
Temperaturabhängigkeit
Temperaturantwort
Thermalphysiologie
temperature dependency
temperature sensitivity
thermal physiology
thermal response
thermische Reaktion
Berry J, Björkman O (1980) Photosynthetic response and adaptation to temperature in higher plants. Annual Review of Plant Physiology 31(1): 491-543 Sadras VO, Moran MA (2013) Nonlinear effects of elevated temperature on grapevine phenology. Agricultural and Forest Meteorology 173: 107-115
Temperature Response Quality
Temperature sum alignment assessment comparing cultivars in a marginal cool climate reveals the thermal requirement matching affecting ripening success, with early cultivar (Müller-Thurgau, requiring 1050 GDD base 10°C from budbreak to harvest maturity) showing good alignment (site provides 1180 GDD in average year, 130 GDD buffer for cool vintage variation, reliable maturity achievement), while late cultivar (Cabernet Sauvignon, requiring 1450 GDD) shows poor alignment (site provides insufficient thermal time even in warm years, maturity rarely achieved, average harvest at inadequate 20.5 Brix), demonstrating how temperature sum alignment determines cultivar viability and the quality outcomes achievable through thermal time matching assessment.
Temperature sum alignment quality is an environmental synchronization quality that represents the correspondence between accumulated thermal time (degree-days or growing degree-days) and the achievement of phenological stages, characterizing how effectively the vine's thermal time requirements for developmental progress align with the heat accumulation patterns of the growing environment to ensure timely completion of each phenological phase within the seasonal thermal budget. Within the BFO framework, this quality specifically addresses the thermal accumulation dimension of environmental synchronization, distinguishing the alignment between thermal requirements and thermal supply from photoperiod coordination while recognizing that temperature sum alignment determines whether the heat units available during the growing season are sufficient and appropriately distributed to support the progression from budbreak through harvest maturity before unfavorable autumn conditions terminate the growing season. The ontological significance lies in temperature sum alignment determining the feasibility of complete phenological cycles within available thermal resources, where well-aligned combinations of cultivar thermal requirements and site thermal supply enable reliable achievement of target maturity with appropriate timing, while misaligned combinations may result in incomplete ripening (insufficient heat units), excessively early ripening (excessive heat compressing development into unfavorable periods), or seasonal timing that places critical stages in suboptimal thermal conditions. The calculation of temperature sum alignment involves comparing cultivar-specific thermal time requirements for each phenological stage (GDD from budbreak to flowering, flowering to véraison, véraison to maturity) against the thermal time typically available in the growing environment during each period, with close alignment indicating appropriate cultivar-climate matching and large discrepancies indicating potential timing or completion problems. The base temperature considerations in thermal time calculations recognize that different cultivars may have different base temperatures below which development does not proceed, and that base temperature may vary among phenological stages, such that accurate alignment assessment requires cultivar-specific thermal time parameters rather than generic degree-day calculations. The climate change effects on temperature sum alignment include the general increase in available growing degree-days that may improve alignment for previously marginal cool-climate situations while degrading alignment for warm-climate regions where excessive heat now compresses phenology, the potential shift in thermal time distribution through the season (earlier heat accumulation advancing phenology relative to historical patterns), and the interaction with other climate factors (water availability, extreme events) that may modify the relationship between thermal time and developmental progress. Understanding temperature sum alignment quality enables assessment of cultivar suitability based on thermal requirement-supply matching, prediction of phenological timing from temperature forecasts, and evaluation of climate change impacts on the thermal time relationships underlying successful grape production.
Temperature_Sum_Alignment_Quality addresses the correspondence between cultivar thermal requirements and environmental heat supply. This quality determines phenological feasibility and timing within seasonal thermal budgets. The class supports cultivar-site matching and climate adaptation through temperature sum alignment characterization.
GDD matching
GDD-Matching
Gradtag-Abstimmung
Temperatursummenabstimmung
Wärmezeitkoordination
degree-day alignment
heat sum synchrony
thermal time coordination
Parker AK et al. (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17(2): 206-216 Jones GV et al. (2010) Spatial analysis of climate in winegrape growing regions in the western United States. American Journal of Enology and Viticulture 61(3): 313-326
Temperature Sum Alignment Quality
In Riesling vines in the Rheingau, Germany, temperature sum response drives developmental progress through March and April, with physiological advancement toward budbreak proceeding in proportion to accumulated warmth above 10°C, meristematic activation increasing progressively as GDD accumulate from 50 units in mid-March to 120 units by early April, and visible budbreak occurring when the response threshold of approximately 140-160 GDD is reached corresponding to sufficient cellular and metabolic preparation for growth initiation.
Temperature sum response in Vitis vinifera represents the physiological integration of thermal exposure that drives developmental progress toward budbreak, describing how buds respond to cumulative warmth by progressively advancing through the preparatory and early growth stages that culminate in visible bud swelling. This response reflects the fundamental temperature-dependence of biological processes underlying budbreak, with warming temperatures accelerating metabolic reactions, gene expression programs, and cellular activities according to their characteristic temperature response functions. The physiological basis of temperature sum response involves the integration of temperature effects across multiple cellular and molecular processes including respiratory metabolism providing energy for growth preparation, biosynthetic pathways producing materials for cell division and expansion, gene expression programs activating growth-related transcription, and cell cycle progression advancing meristematic cells toward division. The response to accumulated temperature is approximately proportional within the range of temperatures typically experienced during spring, with developmental progress increasing in rough proportion to heat exposure above the base threshold, though the relationship becomes non-linear at temperature extremes where very high temperatures may inhibit rather than promote development. The cumulative nature of temperature sum response reflects the persistence of developmental progress achieved during warm periods, with changes in gene expression, protein accumulation, and cellular preparation being maintained rather than reversed during subsequent cool periods, allowing gradual advancement toward budbreak even under fluctuating temperature conditions. The cultivar variation in temperature sum response reflects genetic differences in the thermal requirements for budbreak, with early-budbreak cultivars showing more rapid response to accumulated temperature and reaching the budbreak threshold with less total heat exposure than late-budbreak cultivars. The interaction between temperature sum response and dormancy status affects response efficiency, with fully released buds showing efficient response to heat accumulation while incompletely released buds may show delayed or reduced response due to residual internal constraints. The practical significance of temperature sum response lies in its predictability, enabling forecast of budbreak timing from temperature accumulation that supports vineyard management planning.
Temperature_Sum_Response represents the physiological integration aspect of Heat_Accumulation_Response. Distinguished from Growing_Degree_Day_Accumulation which addresses calculation methodology, while Temperature_Sum_Response addresses the underlying biological response to accumulated warmth. Response approximately proportional to heat accumulation within normal temperature range. Cumulative nature reflects persistence of developmental progress. Cultivar variation in response efficiency affects budbreak timing. Interaction with dormancy status affects response characteristics.
Temperaturakkumulations-Reaktion
Temperatursummen-Reaktion
Wärmeintegrations-Reaktion
cumulative temperature response
heat integration response
kumulative Temperaturantwort
temperature accumulation response
thermal sum response
Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Parker AK et al (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17: 206-216 Bonhomme R (2000) Bases and limits to using degree-day units. European Journal of Agronomy 13: 1-10
Temperature Sum Response
Temporal spread assessment comparing véraison patterns in adjacent Merlot and Grenache blocks reveals the cultivar-characteristic differences in developmental dispersion, with Merlot showing narrow temporal spread (véraison range 9 days from first to last berry color change, CV 15%, concentrated distribution enabling single-date characterization of véraison status), while Grenache shows wide spread (range 22 days, CV 38%, dispersed distribution with extended "red and green" period), demonstrating how temporal spread varies genetically among cultivars with implications for monitoring frequency requirements and harvest timing flexibility.
Temporal spread quality is a synchronization quality that represents the magnitude and pattern of time variation in developmental events across a population of organs or plants, characterizing the degree of temporal dispersion rather than coordination, with spread indicating the width of the developmental window and the distribution of events within that window that together determine the heterogeneity of developmental status at any given time point. Within the BFO framework, this quality addresses the dispersion dimension of developmental timing, distinguishing the spread or range of event timing from the central tendency or synchrony while recognizing that temporal spread provides complementary information to synchrony—narrow spread indicating concentrated timing regardless of mean, wide spread indicating dispersed timing that creates extended developmental windows with consequent management and quality implications. The ontological significance lies in temporal spread determining the heterogeneity of developmental status that must be accommodated in management decisions, where narrow spread enables point-in-time interventions that capture the entire population at appropriate stages, while wide spread requires either extended intervention windows (multiple passes), acceptance of suboptimal timing for population fractions, or active management to reduce spread through selective operations. The statistical characterization of temporal spread involves measures of dispersion such as range (earliest to latest event), standard deviation or coefficient of variation (quantifying the typical deviation from mean timing), and the shape of the distribution (normal versus skewed, unimodal versus multimodal), with these metrics providing different insights into the temporal structure of developmental heterogeneity. The management implications of temporal spread vary by phenological event, with spread in bud break affecting the frost vulnerability window, spread in flowering affecting the fruit set period duration and potential weather exposure, spread in véraison affecting the ripening window, and spread in maturity affecting harvest logistics and fruit uniformity at any selected harvest date. Understanding temporal spread quality enables quantification of developmental heterogeneity, appropriate planning for the duration of management windows, and identification of factors contributing to spread that might be addressable through management interventions.
Temporal_Spread_Quality serves as the parent class for specific spread characteristics (Stage Duration Variance, Maturity Spread, Harvest Window). This quality addresses the dispersion of developmental timing affecting management window duration and developmental heterogeneity. The class supports operational planning through temporal spread characterization.
Entwicklungsdispersion
Zeitvariation
developmental dispersion
event distribution
phenological range
phänologische Spanne
timing variation
zeitliche Streuung
Sadras VO, Petrie PR (2011) Climate shifts in south-eastern Australia: early maturity of Chardonnay, Shiraz and Cabernet Sauvignon is associated with early onset rather than faster ripening. Australian Journal of Grape and Wine Research 17(2): 199-205 Coombe BG, McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6(2): 131-135
Temporal Spread Quality
A Primitivo vine in Puglia's Gioia del Colle trained to tendone bears a Tendone_Role with horizontal canopy at 200 cm, the overhead structure protecting fruit from the region's intense summer radiation while maintaining the traditional landscape that characterizes Puglian viticulture—yields moderated to 10-12 tonnes/hectare for quality wine production. An Italia table grape vine serves in a Tendone_Role in Puglia's dominant table grape industry, the complete overhead shade and protected microclimate supporting the cosmetic fruit quality required for fresh market sale. A Trebbiano d'Abruzzo vine bears a Tendone_Role in traditional Abruzzo vineyards, the pergola system inherited from earlier generations though conversion to more modern training systems proceeds in many zones.
A pergola-trained vine role that inheres in a grapevine trained to the Italian tendone system, characterized by a horizontal overhead canopy at 180-220 cm height, with cordons extending outward from the vine to cover the inter-row space, creating complete overhead shade. Tendone (Italian: "tent" or "awning") is the dominant table grape training system in Italy, particularly in Puglia (southeastern Italy), where the overhead canopy protects fruit from intense summer sun while enabling extended hang time for optimal sugar development. The system uses substantial trellis infrastructure—concrete or wooden posts, overhead wire grid or lattice, multiple cordon wires—representing major establishment investment but enabling the high yields (25-40 tonnes/hectare for table grapes) that justify the cost. Tendone-trained vineyards create distinctive landscape appearance, with the vines forming continuous overhead cover resembling green tunnels or halls between supporting posts. While historically associated with table grapes and bulk wine production, traditional tendone-trained wine grape vineyards persist in Puglia, Abruzzo, and other central-southern Italian regions, with some producers valuing the fruit protection and traditional character the system provides.
Tendone_Role captures the specifically Italian overhead pergola system, distinguished by its industrial scale (table grape production dominance), regional concentration (Puglia and central-southern Italy), and complete overhead canopy coverage. The system's appropriateness varies dramatically with production objective—excellent for table grapes requiring protected, picture-perfect fruit development problematic for premium wine production where excessive shading may reduce color development, phenolic maturation, and aromatic intensity. Climate change creates new relevance for tendone principles, as heat stress and sunburn damage increase in regions where open canopy systems were previously optimal—though modern adaptations typically use modified overhead shading rather than full traditional tendone structure. The system's labor requirements are substantial (canopy management, hand harvest), though mechanization of some operations is possible with appropriately designed infrastructure.
Italian pergola
Tendone-Erziehung
Zeltdach-Erziehung
italienische Pergola
overhead tendone
tendone
tendone training
Ferroni G, Scienza A (2014) Italian grape training systems, ISHS Acta Horticulturae 1032: 333-340 Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Ferrara G et al. (2015) Table grape production in Mediterranean regions, Journal of Berry Research 5: 103-114 Silvestroni O et al. (2018) Traditional training systems of central Italy, OENO One 52: 4
Tendone Role
A tendril at node 7 on a current-season shoot of Vitis vinifera cv. Riesling in the Mosel Valley, Germany, measuring 18 cm in extended length before contact, arising opposite a fully expanded leaf according to the species-typical tendril-inflorescence alternation pattern (tendril at nodes 1, 2, 4, 5, 7, 8...), exhibiting bifurcated branching with two equal-length arms, making contact with a vertical trellis wire and completing both contact coils around the wire and three free coils in the tendril shaft within 6 hours of initial contact, measured at BBCH stage 71 (fruit set) in early June, providing primary mechanical support for the 85 cm shoot bearing five leaves and two developing fruit clusters.
A specialized climbing organ in Vitis vinifera representing a modified inflorescence structure that has lost reproductive function and evolved thigmotropic behavior, characterized by an elongated, slender, flexible stem arising from nodes in positions opposite to leaves following a predictable pattern of tendril-inflorescence alternation along shoots, capable of coiling spirally around supporting structures upon contact stimulation to anchor the vine and enable vertical growth toward light resources. Tendrils represent one of the most distinctive adaptations in the Vitaceae family, with molecular and developmental evidence confirming homology to inflorescence structures rather than leaf modifications, as demonstrated by shared genetic regulatory pathways, vascular architecture, and positional patterns with reproductive structures. The tendril exhibits remarkable mechanosensory capabilities through specialized epidermal cells that detect physical contact and trigger rapid differential growth responses, typically coiling within 5-20 minutes of sustained contact and achieving full mechanical anchoring within 24-48 hours through formation of both free coils between attachment points and contact coils gripping the support structure. Tendril presence, morphology, and coiling characteristics serve important roles in ampelographic identification of cultivars, with features including tendril length, branching patterns, coiling direction, and attachment strength showing cultivar-specific variation useful for distinguishing closely related varieties.
Distinguish tendrils from inflorescences based on developmental fate rather than early morphology, as young tendrils and inflorescence primordia appear similar until reproductive organs differentiate or fail to develop. The tendril-inflorescence pattern in Vitis vinifera typically shows tendrils at nodes 1, 2, then alternating with inflorescences, though pattern variations occur and are cultivar-specific. Tendril functionality varies with position, with basal tendrils often remaining small and non-functional while mid-shoot tendrils achieve full development. Tendril absence at expected positions can indicate inflorescence conversion or developmental abnormalities. Dead or senescent tendrils that have completed their functional life remain attached and must be distinguished from functional tendrils actively providing support.
Kletterorgan
Ranke
Weinranke
climbing organ
grapevine tendril
vine tendril
Based on Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Bowling & Vaughn (2008) "Structural and immunocytochemical characterization of the adhesive tendril of Virginia creeper" Protoplasma 232=153-163 Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 56-78.
Tendril
Tendril attachment quality evaluation in Niagara Peninsula compared cultivar behavior affecting canopy management and mechanical harvest efficiency. Vidal demonstrated problematic tendril attachment quality with abundant, vigorous tendrils producing severe shoot-to-shoot binding that complicated every canopy management operation, with positioning requiring 35% additional labor to separate tangled shoots, mechanical leaf removal suffering 15% efficiency reduction from equipment fouling, and mechanical harvest showing increased material other than grape from tendril-bound leaf and shoot fragments. Riesling exhibited favorable tendril attachment quality with moderate tendril development providing beneficial self-support to trellis wires reducing wind displacement, minimal shoot-to-shoot binding facilitating straightforward positioning, and low interference with mechanical operations. Chardonnay showed intermediate tendril attachment quality adequate for standard management without creating exceptional benefits or challenges. The comparison established tendril attachment quality as a cultivar characteristic warranting consideration in variety selection for mechanized production systems where extreme tendril behavior impacts operational efficiency across multiple seasonal activities.
Tendril attachment quality is a training response quality that inheres in a grapevine cultivar or clone and represents the characteristics of tendril development, behavior, and attachment capacity that influence the self-securing of shoots to trellis infrastructure, neighboring shoots, and canopy support structures, potentially either aiding canopy organization through beneficial attachment to appropriate supports or creating management problems through unwanted binding that complicates positioning operations, mechanical harvest, and cane removal during dormant pruning. This quality captures the dual nature of tendril function wherein moderate, well-directed attachment can reduce positioning labor and secure shoots against wind displacement, while excessive or misdirected attachment creates tangled canopies, complicates mechanical operations, and damages shoots during subsequent handling. The ontological significance of tendril attachment quality lies in its often-overlooked influence on canopy management labor and machine operation efficiency, with extreme tendril characteristics at either end of the attachment spectrum creating practical challenges that moderate tendril behavior avoids. The biological basis of tendril attachment encompasses the frequency of tendril production at nodes along developing shoots, the length and coiling strength of tendrils determining their reach and attachment force, the timing of tendril development relative to shoot positioning operations, and the lignification pattern affecting whether attachments remain flexible or become rigid impediments to subsequent manipulation. Cultivars producing abundant, vigorous tendrils with strong attachment create heavily bound canopies requiring substantial effort to separate for positioning, mechanical leaf removal, or machine harvest, while those with weak or sparse tendrils may benefit from reduced tangling but suffer increased wind displacement and positioning escape rates. Optimal tendril attachment quality represents moderate development enabling beneficial self-support without excessive binding, with attachment primarily to trellis infrastructure rather than to neighboring shoots that would create canopy tangles. Assessment of tendril attachment quality involves characterization of tendril frequency and strength, evaluation of canopy tangling incidence, measurement of positioning labor impacts, and quantification of mechanical operation interference from tendril attachment.
Tendril_Attachment_Quality is_a Training_Response_Quality, representing the characteristics of tendril development and attachment behavior affecting canopy management. This class addresses an often-overlooked cultivar characteristic with practical management implications. The dual nature of tendril function creates an optimum at moderate attachment levels rather than at either extreme. Mechanical operation compatibility represents key consideration for commercial production. Distinguished from other Training_Response_Quality subclasses by the focus on the specialized tendril organs rather than general shoot characteristics.
Rankenanheftungsqualität
Rankenentwicklung
Rankengreifverhalten
Selbstbefestigungskapazität
self-securing capacity
tendril development
tendril grasping strength
vine attachment behavior
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 52-58 Galet P (2000) General Viticulture. Oenoplurimedia, Chaintre, pp. 65-78 Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 64-85
Tendril Attachment Quality
Merlot in Bordeaux shows tendril branching from June through August at BBCH 19-35, with main tendril axes reaching 15-25 mm length before producing typically bifurcated branch structure dividing into two branches of 40-60 mm length over 8-12 days, creating total tendril structure of 80-120 mm with 4-6 terminal segments under oceanic climate conditions of 17-24°C temperature range and adequate resources typical of this moderately vigorous mid-season cultivar producing well-branched tendrils supporting shoot attachment
Tendril branching represents the architectural development process where main tendril axis produces lateral branches creating bifurcated or multifurcated structure characteristic of grapevine tendrils, establishing multiple attachment points and increasing probability of successful support contact and coiling attachment. This distinctive developmental process occurs during or following elongation, with branch initiation typically beginning when main axis reaches 10-20 mm length and branch development continuing over 7-14 days producing final bifurcated structure with 2-4 primary branches each potentially subdividing creating total of 3-8 terminal segments. Branching initiation occurs at specific positions along main tendril axis where localized cell division establishes lateral branch primordia, typically at one or two positions creating characteristic bifurcated or trifurcated primary branch structure, with branch positioning determined during early tendril development through developmental programs establishing branch point locations. Branch primordia develop through similar processes as main tendril axis with localized cell proliferation creating lateral protrusion, followed by coordinated cell division and expansion creating branch axis, and vascular connection establishment linking branch to main tendril vascular system. Branch elongation proceeds through cell division in branch apical zones and cell expansion throughout branch length creating branches typically 60-80% length of main axis distal portion, with elongation rate generally similar to main axis elongation though total length development time shorter due to later initiation. Branching architecture shows cultivar-specific patterns with some varieties producing predominantly bifurcated structure with single branch point dividing main axis into two equal branches, others showing trifurcated pattern with main axis producing two lateral branches while continuing apical growth, and some producing more complex branching with secondary subdivisions creating 6-8 terminal segments. The adaptive significance of branching includes increasing total tendril surface area and number of terminal segments available for encountering support structures, providing multiple independent attachment points increasing attachment reliability and mechanical strength, and creating architectural form optimized for wrapping around diverse support structures including wires, branches, and other plant stems. Terminal segment number affects attachment efficiency with more segments providing increased probability of support contact and successful coiling, though excessive branching may reduce individual segment length potentially limiting coiling effectiveness. Branch development shows coordination with main axis growth ensuring branches develop during period of active tendril elongation when resources are available and growth conditions favorable, with branch initiation and elongation typically completing before main axis growth cessation. Vascular architecture in branched tendril establishes connections from main axis strand into each branch providing water transport for turgor maintenance and coiling function, and enabling assimilate import during development transitioning to potential limited export from any photosynthetic tissue though tendril photosynthetic contribution is minimal. Mechanical properties of branched structure provide both flexibility during searching phase enabling exploration of spatial environment and strength after coiling attachment with multiple coiled segments distributing mechanical loads and providing redundancy if individual segments fail. Branch angles relative to main axis typically range 30-60 degrees creating spreading structure that increases spatial coverage during searching movements and provides advantageous geometry for encountering supports in various orientations. Anatomical structure in branches mirrors main axis with central vascular strand, surrounding flexible parenchyma, peripheral collenchyma, and epidermis with mechanoreceptor capabilities, though branches may show slightly smaller dimensions reflecting later initiation and shorter development period. Metabolic activity in developing branches shows elevated respiration supporting growth similar to main axis elongation, declining to maintenance levels as branch development completes and structure becomes functional. Hormonal regulation of branching involves auxin from main tendril apex potentially affecting branch outgrowth through apical dominance mechanisms similar to shoot branching, with lower auxin concentrations in branch initiation zones permitting lateral growth, and cytokinins possibly promoting branch initiation and outgrowth. Temperature affects branching development rate with branch elongation approximately doubling per 10°C increase between 15-28°C similar to main axis elongation, influencing timing of branch development relative to main axis growth. Light intensity affects branch development with adequate light supporting normal branching while extreme shade may reduce branch number or vigor though branching shows less light dependency than photosynthetic leaf development. Water availability affects branching through effects on cell turgor supporting branch elongation and development, with adequate moisture essential for normal branching pattern achievement. Nutrient availability affects branching through effects on protein synthesis and growth processes supporting branch development, with adequate nutrition supporting normal branching architecture while severe deficiency may reduce branch number or development. The process typically completes within 7-14 days after initiation producing functional branched structure ready for support searching and coiling attachment, with timing varying based on environmental conditions and position along shoot. Cultivar variation appears in branching architecture with some varieties consistently producing bifurcated structure while others show trifurcated or more complex patterns, branch number ranging from 2 primary branches to 4-6 in highly branched types, and branch length relative to main axis affecting overall tendril dimensions and attachment potential. Regional timing shows tendril branching occurring throughout growing season corresponding to tendril formation period from May-July in warm climates, May-August in temperate regions, and June-September in cool areas, with individual tendrils requiring 7-14 days for branching development. Viticultural management considerations include recognizing that branching architecture affects attachment efficiency and reliability, understanding that vigorous branching indicates healthy tendril development and adequate resources, monitoring branching patterns across vineyard assessing vine development uniformity, and considering tendril branching architecture in selecting training systems and support structures. Research applications include analyzing developmental mechanisms of branch initiation and positioning in lateral organs, investigating genetic and environmental regulation of branching architecture, evaluating functional morphology of branched tendril structure and attachment mechanics, quantifying relationship between branching pattern and attachment success, and assessing evolutionary ecology of tendril architecture in climbing plant adaptation.
Tendril_Branching is third of four Tendril_Formation subphases, distinguished by lateral branch development creating characteristic bifurcated or multifurcated structure. Occurs during or after Tendril_Elongation. May overlap with Tendril_Coiling if contact occurs during branch development. Establishes architectural complexity affecting attachment efficiency. Cultivar-specific patterns provide diagnostic characters.
Rankenaufgliederung
Rankengabelung
Rankenramifikation
Rankenverzweigung
branch formation
tendril bifurcation
tendril ramification
tendril subdivision
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 275-278 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 207-210 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Tendril Branching
Syrah in Rhône Valley shows tendril coiling from June through August at BBCH 19-35, with contact with trellis wires triggering coiling response initiating within 10-20 minutes producing 4-6 tight coils around 2.5mm wire within 18-24 hours followed by free coiling in unattached tendril portions creating characteristic spring structure over following 36-48 hours, with subsequent lignification over 10-14 days creating permanent woody attachment supporting vertical shoot growth under Mediterranean climate conditions of 20-28°C temperature range typical of this vigorous mid-to-late season cultivar
Tendril coiling represents the functional response process where tendrils undergo differential growth creating helical curvature that wraps around support structures, establishing mechanical attachment that provides shoot stability and enables vertical growth through anchorage to external supports. This critical functional process occurs following tendril elongation when contact with suitable support surface triggers coiling response, typically requiring 12-48 hours from initial contact to complete primary coiling around support and additional 24-72 hours for secondary free coiling between shoot and support attachment point. Coiling mechanism involves differential cell expansion on opposite sides of tendril with cells on outer convex side elongating more than cells on inner concave side creating curvature, driven by touch-induced signal transduction activating differential growth responses through hormonal and biomechanical mechanisms. Contact detection occurs through mechanoreceptors in tendril epidermal cells sensing physical pressure or friction from support surface contact, transducing mechanical stimulus into biochemical signals including calcium influx, electrical signals propagating through tendril tissues, and hormonal responses triggering differential growth. The coiling response shows two distinct spatial phases including contact coiling where tendril portion touching support coils tightly around support creating primary attachment with multiple coils providing grip, and free coiling where unsupported tendril length between shoot attachment and support contact point coils creating spring-like helical structure. Contact coiling typically produces 3-8 tight coils around support with coil diameter corresponding to support diameter ranging from 3-30 mm, with coiling direction variable though often showing consistent right-handed or left-handed helicity within individual tendrils. Free coiling creates characteristic spring structure with larger diameter coils typically 10-30 mm and variable handedness often reversing direction creating perversions where handedness switches providing mechanical properties of bidirectional spring, enabling tension and compression responses to wind loading and shoot movement. Coiling initiation time after contact shows temperature dependence with responses beginning within 5-15 minutes of contact at optimal 22-28°C, slowing substantially at lower temperatures, and potentially accelerating at higher temperatures within physiological range. Coiling rate during active response shows vigorous differential growth creating visible curvature increasing at 10-30 degrees per hour during peak response 4-12 hours after contact, with complete primary coiling around thin supports achieved within 12-24 hours and coiling around thicker supports requiring 24-48 hours. Cellular mechanisms underlying differential growth include asymmetric auxin distribution with higher concentrations on outer expanding side driven by polar auxin transport, differential cell wall loosening through expansin and other wall-modifying enzyme activities responding to hormonal signals, and asymmetric turgor pressure potentially contributing to immediate response preceding growth-based curvature. Lignification proceeds in coiled tendril portions following completion of coiling response, with secondary wall deposition and lignin incorporation creating rigid permanent attachment structure from initially flexible herbaceous tissue, typically requiring 1-2 weeks for substantial lignification providing long-term mechanical stability. Mechanical strength develops during and after coiling with initial attachment depending on friction and geometric constraint as tender coils wrap support, increasing substantially with lignification creating rigid structure capable of supporting shoot weight and resisting substantial tensile forces from wind loading. Attachment strength in mature lignified coiled tendrils can exceed 10-50 Newtons depending on tendril thickness, number of coils, support diameter, and lignification completeness, providing reliable anchorage for shoots with typical weights of 0.5-2.0 Newtons bearing additional fruit load. Free coil spring function provides mechanical benefits including shock absorption dampening sudden loading from wind gusts, accommodation of shoot growth allowing vertical extension without excessive tension on attachment points, and multi-directional response capacity enabling shoots to move in various directions while maintaining attachment. Vascular function in coiled tendrils maintains water transport capacity through coiled regions supporting distal branch portions that may contact additional supports, though overall hydraulic contribution to shoot water supply is minimal given small vascular dimensions relative to main shoot axis. Tendril longevity after successful coiling extends through multiple growing seasons with lignified coiled tendrils persisting as permanent woody structures, while unattached tendrils failing to contact supports typically senesce and abscise within 2-4 weeks representing unsuccessful development. Metabolic activity in coiling tendrils shows elevated respiration during active coiling response supporting differential growth processes, declining substantially after coiling completion and lignification to minimal maintenance levels in mature woody coiled structures. Hormonal regulation of coiling involves ethylene potentially mediating mechanoreceptor signal transduction and triggering differential growth response, auxin redistribution creating differential expansion on opposite tendril sides, and possibly jasmonates involved in touch signal transduction and response coordination. Temperature affects coiling response rate with optimal 22-28°C supporting rapid vigorous coiling completing within 12-24 hours, cool temperatures below 15°C substantially slowing response potentially requiring 48-96 hours, and very high temperatures above 32°C potentially reducing response vigor through stress effects. Light intensity shows minimal effect on coiling response as mechanically triggered process operates in light or dark, though photosynthetic production supporting overall plant vigor indirectly affects tendril development and function. Water availability affects coiling response through effects on turgor maintaining cell expansion capacity, with adequate moisture essential for normal vigorous coiling while water stress may reduce response rate or vigor affecting attachment success. The process shows cultivar variation in coiling vigor affecting response rate and attachment strength, sensitivity to contact stimulation affecting coiling initiation threshold, and possibly coiling directionality preferences though most cultivars show variable handedness. Regional timing shows tendril coiling occurring throughout growing season corresponding to tendril development period from May-August in temperate regions, with individual coiling responses requiring 12-72 hours depending on support type and environmental conditions. Viticultural management considerations include providing appropriate support structures with suitable dimensions for tendril coiling typically 3-20 mm diameter, timing shoot positioning operations allowing tendrils to attach before requiring shoot adjustment, monitoring attachment success assessing support system adequacy, recognizing that vigorous coiling indicates healthy tendril function and adequate resources, and managing excessive tendril attachment that may complicate shoot repositioning or removal operations. Research applications include investigating mechanoreception and mechanotransduction pathways mediating touch responses, analyzing cellular and molecular mechanisms of differential growth creating curvature, evaluating biomechanics of coiling attachment and spring function, quantifying environmental effects on coiling responses affecting attachment success, and assessing evolutionary ecology of coiling mechanisms in climbing plant adaptation and support utilization strategies.
Tendril_Coiling is fourth and final Tendril_Formation subphase, distinguished by functional attachment response following support contact. Represents culmination of tendril development converting structure to function. May occur during or after Tendril_Elongation and Tendril_Branching depending on contact timing. Critical functional process enabling climbing growth habit. Unsuccessful coiling results in tendril senescence and abscission.
Rankeneinrollung
Rankenumschlingung
Rankenwicklung
Spiralbildung
attachment formation
helical coiling
tendril curling
tendril wrapping
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 275-282 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 207-215 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Tendril Coiling
The tendril description of Pinot Noir documents medium-length bifurcate (two-branched) tendrils distributed in the characteristic V. vinifera pattern of two consecutive tendril-bearing nodes followed by one tendril-free node (2-0-2-0 pattern beginning at node 3 or 4 from the shoot base), with green color during the growing season transitioning to brown after lignification — the tendrils are moderately vigorous with the longer branch reaching 150-200 mm before coiling, and exhibit the typical positive thigmotropic response (coiling around contact surfaces within 1-2 hours of mechanical stimulation). This standard Euvitis tendril pattern distinguishes V. vinifera from Muscadinia rotundifolia (which produces simple, unbranched tendrils at every node without the 2-0 alternation) and from tendril-free mutant lines used in genetic studies of the tendril-inflorescence developmental switch.
A shoot description component that documents the morphological characteristics of tendrils — the modified inflorescences that serve as climbing organs in Vitis — recording length, branching pattern, distribution along the shoot axis, color, and the presence or absence of tendrils at specific node positions, with the tendril distribution pattern providing species-level diagnostic information and tendril morphology contributing to cultivar characterization within Vitis vinifera.
Tendril_Description documents the morphology of an organ that occupies a unique position in grapevine developmental biology — tendrils are homologous to inflorescences, arising from the same primordial meristems at each node, with the developmental fate (tendril versus inflorescence) determined by the interaction of gibberellin signaling, the floral promoter VvFT, and positional signals within the shoot apical meristem. In wild Vitis, tendrils are essential climbing organs enabling the liana growth habit, coiling around support structures to anchor the extending shoot in the tree canopy: in cultivated grapevine trained on trellis systems, tendrils retain this function by attaching to wires and adjacent shoots, contributing to canopy structure maintenance between manual shoot positioning passes. The tendril distribution pattern — specifically the 2-0-2-0 alternation in Euvitis versus continuous tendrils in some species — is the most diagnostically significant tendril character, functioning primarily at the species and subgenus level rather than for cultivar discrimination within V. vinifera. Within V. vinifera, tendril length, vigor, and coiling behavior show cultivar-associated variation (vigorous cultivars tend to produce longer, more robust tendrils) but these characters are strongly influenced by vine vigor and are therefore less genetically stable than other ampelographic descriptors.
Klettorgan-Charakterisierung
Ranken-Beschreibung
Ranken-Morphologie
cirrhus description
climbing organ characterization
tendril morphology record
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Boss PK and Thomas MR (2002) Association of dwarfism and floral induction with a grape green revolution mutation, Nature 416: 847-850 Calonje M et al. (2004) Floral meristem identity genes are expressed during tendril development in grapevine, Plant Physiology 135: 1491-1501
Tendril Description
Vitis vinifera displays the characteristic 2:1 phyllotaxy with tendrils present at two consecutive nodes followed by one node without tendril, while some cultivars like 'Sultanina' show modified patterns with occasional 3:1 arrangements. Tendril dry weight allocation ranges from 2-8% of total shoot biomass depending on cultivar and trellis system.
A grapevine disposition that is the inherited capacity of a Vitis vinifera plant to develop tendrils as modified shoots homologous to inflorescences, characterized by the phyllotactic pattern of tendril placement along the shoot axis and the morphological vigor of tendril formation.
Tendril_Development_Disposition serves as the parent class for tendril-related traits. Tendrils in Vitis are developmentally homologous to inflorescences, sharing the same meristematic origin, with the uncommitted primordia (anlagen) differentiating into either structure based on gibberellin signaling. The VvTFL1A gene plays a key role in maintaining tendril identity versus inflorescence conversion.
Rankenbildungsvermögen
Rankenentwicklungsdisposition
tendril formation capacity
tendril production trait
Mullins MG, Bouquet A, Williams LE (1992) Biology of the Grapevine, Cambridge University Press pp.: 52-58 Carmona MJ et al. (2008) A molecular genetic perspective of reproductive development in grapevine. Journal of Experimental Botany 59:2579-2596 Calonje M et al. (2004) Floral meristem identity genes are expressed during tendril development in grapevine. Plant Physiology 135:1تم491-1501
Tendril Development Disposition
The tendril distribution of Vitis vinifera cv. Riesling follows the standard Euvitis 2-0-2-0 pattern — beginning at node 3 or 4 from the shoot base (below which nodes bear inflorescences instead of tendrils in fruitful shoots, or neither tendril nor inflorescence in non-fruitful shoots), two consecutive nodes each produce a tendril opposite the leaf, followed by one tendril-free node bearing only a leaf and lateral bud, then two more tendril-bearing nodes, continuing in this 2-0 alternation to the shoot tip. This pattern is remarkably consistent across all V. vinifera cultivars (the tendril distribution is a species-level rather than cultivar-level character), distinguishing V. vinifera from the wild American species V. labrusca (which tends toward continuous tendril production at every node with occasional gaps) and from M. rotundifolia (which produces simple unbranched tendrils with a different phyllotactic arrangement).
A tendril description component that records the pattern of tendril occurrence along the shoot axis, documenting the characteristic alternation of tendril-bearing and tendril-free nodes that is species-diagnostic within the genus Vitis — the standard Euvitis pattern of two consecutive tendril-bearing nodes followed by one tendril-free node (2-0-2-0), versus the continuous tendril pattern (every node bearing a tendril) observed in some non-vinifera species, versus the discontinuous patterns characteristic of Muscadinia — with the distribution pattern reflecting the developmental regulation of the inflorescence-to-tendril conversion program in the shoot apical meristem.
Tendril_Distribution documents a character that operates primarily at the species and subgenus level rather than providing cultivar discrimination within V. vinifera — virtually all V. vinifera cultivars share the identical 2-0-2-0 distribution pattern, making this character uninformative for intraspecific identification but highly valuable for detecting interspecific hybrid ancestry or species misidentification. The 2-0-2-0 pattern reflects the developmental program of the sympodial shoot apex, in which the shoot apical meristem produces three phytomers (node-internode units) as a developmental module — the first two phytomers each produce a tendril-competent primordium opposite the leaf (which develops as an inflorescence under floral induction or as a tendril in the absence of floral signals), while the third phytomer produces no tendril primordium: this three-phytomer module then repeats, generating the 2-0 pattern observed externally. The inflorescence primordia at the basal 2-3 nodes of fruitful shoots develop as flower clusters rather than converting to tendrils, because these primordia were initiated during the previous growing season within the dormant bud under floral-inductive conditions — the tendril-to-inflorescence ratio at each node position therefore reflects the developmental history of the primordium rather than current-season environmental signals.
Ranken-Auftrittsmuster
Rankenverteilung
nodal tendril distribution
nodale Rankenanordnung
phyllotactic tendril arrangement
tendril occurrence pattern
Tendril Distribution
Tendril drooping quality evaluation in Walla Walla Merlot established the sensitivity of this early-warning indicator for deficit irrigation management during the critical post-véraison ripening period. Morning observation of tendril turgor status at shoot tips provided the most sensitive visual indicator of overnight recovery from previous-day stress, with fully turgid, actively-coiling tendrils indicating complete recovery and adequate water status while flaccid, drooping tendrils signaled incomplete overnight rehydration indicating cumulative stress exceeding recovery capacity. A three-point tendril turgor scale was developed with score 1 (turgid, active coiling) corresponding to predawn water potential above -0.3 MPa indicating no significant stress, score 2 (partially flaccid, reduced activity) at -0.3 to -0.5 MPa indicating mild-moderate stress in the beneficial range, and score 3 (completely drooping, no coiling) below -0.5 MPa indicating stress at or beyond optimal severity. The early-morning tendril assessment enabled daily stress evaluation without instrumentation, providing practical management guidance for a 50-hectare estate where regular pressure chamber measurements were impractical, demonstrating the value of tendril drooping quality as an accessible early-warning indicator.
Tendril drooping quality is a drought symptom expression quality that inheres in grapevine tendrils and represents the loss of turgid, actively-seeking orientation exhibited by healthy tendrils when water deficit reduces turgor below the threshold for maintaining the characteristic coiled or extended posture, causing tendrils to assume a flaccid, pendant configuration that provides one of the earliest and most sensitive visual indicators of developing water stress detectable before more obvious symptoms appear in larger plant organs. This quality captures the diagnostic value of tendril turgor status as an early-warning stress indicator, recognizing that the relatively small volume and exposed position of tendrils makes them particularly responsive to changes in plant water status, with turgor loss appearing in tendrils before leaves exhibit visible stress symptoms. The ontological significance of tendril drooping quality lies in its role as a sensitive leading indicator enabling detection of incipient stress before it progresses to levels affecting larger organs and potentially impacting fruit quality development, providing opportunity for proactive management intervention at early stress stages. The biological basis of tendril drooping involves the high sensitivity of tendril turgor to plant water status, with the small tissue volume and limited osmotic adjustment capacity of tendrils causing rapid turgor loss as water potential declines, combined with the exposed position of actively-growing tendrils at shoot tips where evaporative demand is highest and hydraulic supply most tenuous. Healthy tendrils maintain characteristic turgid posture enabling their twining function, with the growing tip actively seeking contact points and the coiled portion maintaining spring tension, while water-stressed tendrils lose this active orientation and hang limply unable to perform their attachment function. The progression from turgid to drooping tendril state occurs rapidly with changing water status, with recovery equally rapid when water availability improves, making tendril turgor a dynamic indicator of current rather than accumulated stress status.
Tendril_Drooping_Quality is_a Drought_Symptom_Expression_Quality, representing the early-appearing loss of turgor in tendrils as a sensitive stress indicator. This class addresses the most sensitive visual stress symptom due to tendril exposure and limited osmotic adjustment capacity. The early-morning assessment timing captures overnight recovery status. Distinguished from Leaf_Wilting_Quality by earlier appearance and greater sensitivity. Tendrils provide dynamic current-status indication rather than accumulated stress assessment. Practical value for frequent monitoring without instrumentation.
Ranken-Turgeszenz-Verlust
Rankenabsinken
Rankenerschlaffung
Rankenwelke
tendril drooping symptom
tendril flaccidity
tendril turgor loss
tendril wilting
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 453-507 Cifre J et al. (2005) Physiological tools for irrigation scheduling in grapevine (Vitis vinifera L.). Agriculture, Ecosystems and Environment 106:159-170 Lovisolo C et al. (2010) Drought-induced changes in development and function of grapevine (Vitis spp.) organs. Functional Plant Biology 37:98-116
Tendril Drooping Quality
Chardonnay in Champagne shows tendril elongation from late May through July at BBCH 15-35, with individual tendrils requiring 8-12 days to elongate from 5-10 mm primordia to functional length of 60-90 mm through elongation rate of 5-8 mm per day during peak growth, under cool maritime climate conditions of 15-20°C temperature range, adequate moisture, and moderate fertility typical of this moderately vigorous early-bursting cultivar producing tendrils supporting shoot positioning in vertical training systems
Tendril elongation represents the primary growth phase where initiated primordia increase length through coordinated cell division and expansion creating slender flexible structures capable of searching for and attaching to support surfaces, establishing functional dimensions and anatomical characteristics enabling climbing function. This critical developmental process follows primordium initiation, typically requiring 7-14 days per tendril from elongation initiation to functional length, with multiple tendrils at various elongation stages creating developmental gradient along shoot corresponding to regular tendril positioning pattern. Elongation proceeds through continued cell division in apical growth zone producing new cells contributing to tendril length, and progressive cell expansion throughout tendril axis where cells increase length creating visible elongation of 3-8 mm per day during active growth. Cell division during elongation shows highest activity in apical zone behind tendril tip maintaining meristem-like characteristics, with division continuing for 5-10 days producing cells arranged in longitudinal files characteristic of unidirectional growth, then transitioning to expansion phase where cell production ceases but existing cells undergo dramatic elongation. Cell expansion occurs throughout elongating tendril with cells increasing from meristematic dimensions of 10-20 micrometers to final lengths of 100-300 micrometers representing 10-30 fold increase, through water uptake creating turgor pressure driving cell wall extension and progressive wall thickening maintaining structural integrity. Tendril length increase shows characteristic pattern with rapid elongation during early-mid phase at 5-10 mm per day when both cell division and expansion operate actively, declining to 2-4 mm per day during late phase as division ceases and expansion completes, with final functional length typically 50-150 mm depending on cultivar, position along shoot, and environmental conditions. Elongation rate varies among tendrils reflecting shoot growth conditions with tendrils forming during peak shoot growth showing maximum elongation rates and achieving longest dimensions, while tendrils forming during declining growth show reduced rates and shorter final lengths. Anatomical development during elongation establishes characteristic tendril structure including central vascular strand with xylem vessels for water transport and phloem sieve elements for assimilate distribution, surrounding parenchyma providing flexibility and storage capacity, peripheral collenchyma providing mechanical strength while maintaining flexibility, and epidermis with specialized cells capable of detecting contact stimulation. Vascular differentiation proceeds basipetally from connection with shoot axis toward tendril tip, with procambial cells differentiating into functional xylem vessels and phloem elements establishing continuous transport pathway supporting tendril growth and eventual coiling responses. The elongating tendril shows negative gravitropism growing upward and positive thigmotropism enabling growth toward potential support structures through touch-induced directional responses, with combination of growth orientations positioning tendril optimally for encountering supports. Tendril surface cells develop mechanoreceptor capabilities during elongation enabling detection of contact with support surfaces that will trigger subsequent coiling responses, with mechanoreceptor distribution particularly concentrated on tendril ventral surfaces that will contact supports. Mechanical properties during elongation emphasize flexibility enabling searching movements and exploration of spatial environment for support structures, with young elongating tendrils showing low tensile strength and high flexibility from thin primary cell walls. Metabolic activity remains elevated during elongation with respiration 8-12 fold above mature tissue levels supporting growth processes including cell division, expansion, vascular differentiation, and wall synthesis. Protein synthesis operates at high rates producing structural proteins for expanding cells, enzymes supporting metabolism and wall modification, and potentially beginning accumulation of proteins involved in mechanoreception and signal transduction for coiling responses. Circumnutation movements may occur during elongation with tendril tip describing circular or elliptical path through space enhancing probability of encountering support structures, driven by differential growth rates on opposite sides of tendril coordinated through internal oscillatory developmental program. Hormonal regulation coordinating elongation involves auxin promoting elongation and coordinating vascular differentiation pattern, gibberellins supporting cell expansion affecting final tendril length, and possible brassinosteroid effects on cell expansion and mechanoreceptor development. Temperature strongly affects elongation rate with development approximately doubling per 10°C increase between 15-28°C optimal range, influencing both cell division duration and expansion rate determining time required for functional length achievement. Light intensity affects elongation with adequate light supporting normal development while shade may promote enhanced elongation through etiolation-like response seeking improved light exposure, potentially affecting final tendril length and functionality. Water availability critically affects elongation through direct effects on cell turgor driving expansion, with adequate moisture essential for maintaining pressure supporting cell wall extension while water stress arrests elongation limiting final dimensions affecting attachment potential. Nutrient availability particularly nitrogen affects elongation through effects on protein synthesis supporting cell division and expansion, with adequate nutrition supporting maximum elongation while deficiency reduces elongation rate and limits final dimensions. The process shows variation in timing and rate among tendrils creating pattern along shoot with early-season tendrils requiring longer elongation periods during slower shoot growth, mid-season tendrils elongating rapidly during peak conditions achieving maximum length, and late-season tendrils showing abbreviated rapid elongation before growth cessation. Cultivar variation appears in maximum elongation rate, final tendril length ranging from short 40-80 mm in some varieties to long 120-180 mm in vigorous types affecting climbing ability, and possibly circumnutation vigor affecting support encounter probability. Regional timing shows tendril elongation occurring throughout growing season from May-July in warm climates, May-August in temperate regions, and June-September in cool areas offset from initiation by 1-2 weeks, with individual tendrils requiring 7-14 days for elongation depending on conditions. Viticultural management considerations include recognizing that vigorous tendril elongation indicates healthy shoot growth and adequate resources, understanding that elongation timing affects availability for shoot attachment and training operations, monitoring elongation patterns assessing vine vigor and resource status, and considering tendril development in timing of shoot positioning and wire attachment operations. Research applications include analyzing cellular and molecular mechanisms of unidirectional organ elongation, investigating circumnutation mechanisms and their ecological function in support location, evaluating mechanoreceptor development and mechanotransduction pathways, quantifying environmental effects on elongation affecting tendril functionality, and assessing biomechanics of tendril searching behavior and attachment efficiency.
Tendril_Elongation is second of four Tendril_Formation subphases, distinguished by length increase creating functional tendril structure. Follows Tendril_Primordium_Initiation. Transitions to Tendril_Branching as branch points develop and Tendril_Coiling as contact with supports occurs. Primary determinant of final tendril dimensions and attachment potential. Critical phase establishing functional searching structure.
Rankenexpansion
Rankenstreckung
Rankenverlängerung
Rankenwachstum
tendril extension
tendril growth
tendril stretching
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 270-275 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 202-207 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Tendril Elongation
Cabernet Sauvignon in Napa Valley shows tendril formation from May through July at BBCH 15-35, producing tendrils at every third node in pattern of two tendril nodes followed by one inflorescence node, with individual tendrils developing bifurcated structure over 2-3 weeks reaching 80-120 mm length and coiling around trellis wires within 24-36 hours of contact under warm climate conditions of 18-28°C temperature range, providing essential support for vertical shoot positioning in VSP training system typical of this vigorous late-season cultivar
Tendril formation represents the specialized developmental process producing climbing structures modified from ancestral inflorescence architecture, enabling grapevine shoots to attach to supports through coiling mechanisms and providing mechanical stability for vertical growth and optimal light interception. This distinctive developmental process occurs at regular intervals along shoot during active growth, typically producing tendrils at every third node in alternating pattern with inflorescences following cultivar-specific architectural programs, with tendril development spanning 2-4 weeks from primordium initiation to functional coiling structure. Tendril formation encompasses primordium initiation at shoot apex in positions homologous to inflorescence primordia, elongation through coordinated cell division and expansion creating tendril axis and branches, branching establishing bifurcated or multifurcated structure characteristic of grapevine tendrils, and coiling through differential growth responses enabling attachment to supports. The developmental origin of tendrils reflects evolutionary modification of inflorescence structures, with tendrils representing sterile inflorescence homologs that have lost reproductive function while maintaining developmental program for producing branched structures, evident in occasional tendril-inflorescence intermediates showing partial flower development. Tendril positioning follows regular architectural pattern along shoot axis with typically two successive nodes bearing tendrils followed by one node bearing inflorescence in fruiting shoots, or continuous tendril production at every third node in sterile shoots, creating predictable structural organization. Primordium initiation occurs at shoot apex in lateral positions opposite leaves following phyllotactic pattern similar to inflorescence positioning, with primordia arising as multicellular protrusions from meristem flank developing through similar early stages as inflorescence primordia before diverging toward tendril-specific development. Tendril structure typically shows bifurcated or trifurcated branching pattern with main axis dividing into 2-3 branches, each potentially subdividing creating total of 3-6 terminal segments, with branching architecture affecting attachment efficiency and mechanical strength. Elongation proceeds through cell division in apical regions and cell expansion throughout tendril axis creating slender flexible structure typically 50-150 mm total length at functional maturity, with elongation rate coordinated with shoot growth ensuring tendril availability for attachment as shoot extends. Tendril anatomy shows central vascular core providing water and nutrient transport, surrounding parenchyma and collenchyma tissues providing flexibility and strength, and epidermis with specialized cells enabling coiling responses, with anatomical structure optimized for combining flexibility during searching with strength after coiling attachment. Coiling mechanism involves differential growth responses with cells on one side of tendril expanding more than opposite side creating curvature, triggered by contact stimulation through mechanoreceptors detecting physical contact with support structures, with response developing over hours to days creating tight coils securing attachment. The coiling response shows two distinct phases including initial contact coiling where tendril portion touching support coils around support creating primary attachment within 12-24 hours, and subsequent free tendril coiling where unsupported portions between shoot and support coil creating spring-like structure that provides mechanical stability and shock absorption. Tendril functionality provides critical support enabling vertical shoot growth reaching canopy heights of 2-4 meters in trained systems or climbing into tree canopies reaching 10-20 meters in wild populations, with attachment strength sufficient to support shoot weight and resist wind loading. Mechanical properties develop during maturation with young tendrils showing high flexibility enabling searching movements, transitioning to increased rigidity after coiling attachment through lignification in coiled regions providing permanent structural support. Lignification proceeds in attached coiled tendrils creating woody persistent structures that remain functional for multiple seasons, while unattached tendrils typically senesce and abscise having failed to fulfill attachment function. Metabolic activity in developing tendrils shows elevated respiration supporting growth processes, declining to maintenance levels in mature attached tendrils, with photosynthetic contribution negligible due to limited chlorophyll content and small surface area. Water transport through tendrils contributes minimally to shoot hydraulic system as small vascular capacity relative to main shoot axis, though maintaining tendril turgor may be important for coiling function and attachment maintenance. Hormonal regulation involves auxin affecting elongation and possibly coiling responses, ethylene potentially involved in coiling mechanism and senescence of unattached tendrils, and jasmonates possibly affecting touch-induced coiling responses through mechanotransduction pathways. Temperature affects tendril development rate with elongation approximately doubling per 10°C increase between 15-28°C, influencing timing of tendril availability for attachment relative to shoot growth progression. Light intensity affects tendril development with adequate light supporting normal elongation and function while extreme shade may reduce tendril production or vigor affecting climbing ability. Water availability affects tendril turgor and function with adequate moisture maintaining flexibility and coiling capability while severe water stress may impair coiling responses affecting attachment success. The process shows characteristic progression through distinct developmental phases advancing from primordium initiation at apex through elongation creating structure, branching establishing architecture, and coiling enabling functional attachment. Cultivar variation appears in tendril production pattern with some varieties showing consistent every-third-node pattern while others show variation, branching architecture affecting attachment efficiency, and coiling vigor affecting climbing ability with implications for training requirements. Regional timing shows tendril formation occurring throughout shoot growth from May-July in warm climates, May-August in temperate regions, and June-September in cool areas, with formation rate corresponding to shoot growth rate and node production frequency. Viticultural management considerations include understanding tendril positioning pattern informing node counting for reproductive structure prediction, recognizing that vigorous tendril production indicates healthy shoot growth, utilizing tendril attachment for shoot positioning and training in vertical systems, monitoring tendril function assessing support system adequacy, and removing excessive tendrils in some training systems reducing shoot entanglement and improving canopy management. Research applications include analyzing evolutionary developmental biology of organ modification from reproductive to vegetative function, investigating molecular mechanisms of tendril identity specification and coiling responses, quantifying mechanical properties and attachment strength affecting vine architecture, evaluating coiling mechanotransduction pathways and signal integration, and assessing tendril function in wild grapevine ecology and implications for cultivated vine architecture.
Tendril_Formation represents specialized lateral organ development within Shoot_Growth, producing attachment structures enabling climbing growth habit. Distinguished from Leaf_Emergence and Inflorescence_Development though sharing developmental homology with latter. Subdivided into Tendril_Primordium_Initiation, Tendril_Elongation, Tendril_Branching, and Tendril_Coiling based on developmental progression. Critical for vine architecture in both wild and cultivated contexts. Pattern regularity provides diagnostic information about shoot structure.
Kletterorganbildung
Rankenbildung
Rankenorganogenese
climbing structure formation
tendril organogenesis
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 265-278 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 195-210 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Tendril Formation
The tendril length of Grenache documents long tendrils (OIV state 7) with the primary branch extending 200-280 mm before initiating coiling — consistent with Grenache's vigorous growth habit and semi-pendulous shoot architecture that benefits from strong tendril attachment to maintain canopy structure on wind-exposed Mediterranean sites. The secondary branch typically reaches 60-70% of the primary branch length (140-190 mm), creating the bifurcate Y-shape characteristic of Euvitis tendrils. This long tendril development contrasts with the short, relatively weak tendrils of Pinot Noir (OIV state 3, primary branch 100-150 mm) whose compact growth habit and short internodes reduce the mechanical demand on tendril support, and the medium tendrils of Merlot (OIV state 5, primary branch 150-200 mm).
A tendril description component that records the length of fully extended tendrils measured from the point of insertion at the shoot node to the tip of the longest branch before coiling, scored on a scale from very short (state 1) to very long (state 9), with tendril length reflecting the cultivar-specific vigor of tendril growth that correlates with overall shoot vigor and influences the capacity of the vine to attach to trellis support structures.
Tendril_Length documents a quantitative character that is informative for cultivar characterization but must be interpreted in the context of overall vine vigor. Tendril elongation is driven by the same gibberellin-mediated cell expansion process that controls internode length, and consequently tendril length correlates positively with internode length across cultivars — long-internode cultivars tend to produce long tendrils and vice versa: this correlation means that tendril length provides partially redundant information with internode length for cultivar identification, but the two characters are not perfectly correlated (some cultivars show disproportionately long or short tendrils relative to their internode length), and the residual independent variation contributes additional diagnostic value. The environmental sensitivity of tendril length parallels that of internode length — high-vigor conditions (excessive water, nitrogen, or vigorous rootstock) increase tendril length by 30-50% compared to balanced conditions — requiring standardized observation on normally growing shoots for reliable cultivar comparison. The functional significance of tendril length extends beyond identification to practical viticulture, as cultivars with long, vigorous tendrils attach more effectively to trellis wires and require less manual shoot positioning labor than cultivars with short, weak tendrils.
Klettorgan-Ausdehnung
Ranken-Längenmessung
Rankenlänge
cirrhus length measurement
climbing organ extent
tendril dimension record
Tendril Length
Pinot Noir in Burgundy shows tendril primordium initiation from May through July at BBCH 15-35 at shoot apex, initiating new tendril primordia every 6-8 days at every third node position following two-tendril-one-inflorescence architectural pattern, with primordia developing from 50-100 micrometer initial bulges to 400-500 micrometer recognizable tendril structures over 5-7 days under continental climate conditions of 16-26°C temperature range and moderate fertility typical of this moderate-vigor early-to-mid season cultivar
Tendril primordium initiation represents the earliest developmental stage where shoot apical meristem produces lateral primordia in positions homologous to inflorescence primordia that will develop into tendril structures rather than reproductive organs, establishing climbing attachment structures following regular architectural pattern along shoot axis. This fundamental organogenesis process occurs at shoot apex during active growth following every-third-node pattern in most cultivars, with new tendril primordia initiating every 6-9 days corresponding to three leaf primordium initiations, producing total of 4-8 tendril primordia per shoot over growing season. Primordium initiation begins with localized increase in cell division activity on apical meristem flank at lateral position opposite developing leaf primordium, with cells in initiation zone undergoing accelerated divisions creating multicellular protrusion marking first visible stage of tendril formation. The initiation site occurs in regular architectural pattern with typically two successive nodes producing tendril primordia followed by one node producing inflorescence primordium in reproductive shoots, or continuous tendril production at every third node in sterile shoots, with positioning controlled by genetic developmental programs and possibly hormonal signals. Tendril primordium initiation shares early developmental stages with inflorescence primordium formation showing similar multicellular protrusion and initial cell division patterns, with divergence toward tendril versus inflorescence fate occurring through differential gene expression suppressing floral meristem identity genes and activating tendril-specific developmental programs. The molecular basis of tendril versus inflorescence specification involves transcription factors regulating meristem identity with mutations in key regulatory genes capable of converting tendrils to inflorescences or vice versa, demonstrating close developmental relationship and evolutionary origin of tendrils from modified inflorescence structures. Primordial cells undergo oriented divisions establishing elongated axis structure characteristic of tendril rather than compact branching structure of inflorescence, with longitudinal growth dominating over radial growth creating slender primordium form. Primordium size at initiation measures approximately 50-100 micrometers diameter similar to other lateral organ primordia, increasing to 300-500 micrometers with several hundred cells by time primordium is clearly recognizable as tendril structure rather than simple protrusion or potential inflorescence, requiring approximately 4-7 days from first division acceleration to recognizable tendril form. Vascular strand initiation occurs early in primordium development establishing connection between developing tendril and shoot axis, with procambial cells forming continuous strand from primordium base through shoot cortex connecting to main vascular system, providing pathway for water delivery and nutrient transport supporting tendril development. Gene expression patterns during initiation include activation of lateral organ identity genes specifying formation of lateral structure, expression of tendril-specific genes suppressing inflorescence development programs, and morphogenesis genes controlling cell division patterns and growth orientation establishing elongated tendril form. Early primordium development shows establishment of apical-basal polarity defining tip and base regions with apical zone maintaining division capacity longer supporting elongation growth characteristic of tendril development. Branching point determination may occur during initiation phase with cells at specific positions along developing primordium establishing zones where branches will subsequently emerge, though branch development itself occurs later during elongation phase. Hormonal regulation of initiation involves auxin concentration patterns at meristem apex with local auxin distributions possibly affecting tendril versus inflorescence fate determination, cytokinins from roots supporting cell division in initiating primordia, and gibberellins potentially affecting tendril elongation programming. Timing of tendril primordium initiation relative to leaf primordium initiation follows architectural program with tendril initiation occurring at every third node position, coordinated through developmental signals ensuring regular pattern maintenance along shoot axis. Environmental conditions during initiation period may influence tendril versus inflorescence determination with stress conditions potentially promoting tendril production over reproductive structures in some developmental contexts, though genetic programs predominantly control organ identity. Temperature affects initiation timing through effects on apical meristem activity with optimal 22-28°C supporting regular initiation frequency maintaining every-third-node pattern, while suboptimal temperatures may slow meristem activity affecting timing though typically maintaining positional pattern. Light intensity and quality during initiation may influence developmental programs with high light supporting normal development while extreme shade potentially affecting organ identity determination in some cases. Water availability affects meristem function with adequate moisture maintaining normal initiation patterns while severe water stress potentially disrupting regular development and organ identity specification. Nutrient availability particularly nitrogen affects meristem activity and organ initiation with adequate nutrition supporting normal initiation frequency and organ development while severe deficiency may reduce initiation or affect organ identity. The process shows regular temporal pattern corresponding to leaf initiation frequency with new tendril primordium initiating after every third leaf primordium, creating predictable sequence of organs along developing shoot axis. Cultivar variation appears in regularity of every-third-node pattern with some varieties showing consistent pattern while others show occasional variation, tendril versus inflorescence determination with some cultivars showing higher proportion of reproductive structures, and possibly branching architecture programming during early primordium development. Regional timing shows tendril primordium initiation occurring throughout shoot growth period from May-July in warm climates, May-August in temperate regions, and June-September in cool areas, with initiation frequency corresponding to apical meristem activity and shoot growth rate. Viticultural significance includes establishing tendril positioning pattern that informs shoot structure prediction and node counting for crop estimation, recognizing that initiation reflects normal shoot development and meristem function, and understanding that environmental or genetic factors affecting initiation may alter shoot architecture affecting training and management. Research applications include analyzing molecular mechanisms of organ identity specification and tendril versus inflorescence determination, investigating evolutionary developmental biology of organ modification and function change, evaluating genetic and environmental factors affecting developmental fate decisions, documenting cellular mechanisms of primordium formation and polarity establishment, and assessing architectural programs controlling regular organ positioning patterns in shoot development.
Tendril_Primordium_Initiation is first of four Tendril_Formation subphases, distinguished by organogenesis at apical meristem establishing tendril structures. Represents earliest microscopic stage occurring at shoot apex. Transitions to Tendril_Elongation as primordia continue developing. Shares early developmental stages with inflorescence primordia reflecting evolutionary homology. Critical for establishing regular architectural pattern along shoot.
Rankenanlagebildung
Rankeninitiierung
Rankenorganogenese-Beginn
Rankenprimordienbildung
tendril initiation
tendril organogenesis initiation
tendril primordium formation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 265-270 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 195-202 Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109
Tendril Primordium Initiation
A terminal apical leaf at node 16 on a vigorously extending shoot of Vitis vinifera cv. Syrah in the Rhône Valley, France, positioned 2 cm behind the shoot apex representing the most recently emerged visible leaf, measuring 2.8 cm length in partially unfolded configuration representing approximately 15% of expected mature size, exhibiting very pale yellow-green coloration with pronounced bronze tinge from anthocyanin concentration of 4.5 mg g⁻¹ fresh weight, extremely soft fragile texture with cells composed primarily of meristematic tissue showing active division, procambial strands barely visible as faint green lines without distinct vascular differentiation, no functional stomata or cuticle evident, demonstrating essentially zero photosynthetic activity with minimal chlorophyll content of 85 μg cm⁻² representing pure resource sink consuming approximately 8 mg carbohydrate per day for rapid cell division, positioned at the growing tip receiving full sunlight but lacking photosynthetic capacity to utilize high irradiance, occurring during peak shoot extension at 9-11 cm per week growth rate, measured at 3 days post-emergence during active vegetative growth at shoot BBCH stage 73 (berries beginning to touch) in early July, representing the newest leaf addition to the extending shoot with subsequent leaves initiating behind at 2-3 day intervals.
An apical leaf at the absolute terminal position of a shoot representing the most recently emerged or currently emerging leaf in Vitis vinifera, characterized by position at or within 1-3 nodes of the shoot apex representing the developmental frontier of shoot extension, extremely small size typically 1-5 cm length representing 5-25% of eventual mature dimensions, very recent emergence within previous 2-7 days or still in process of emergence from protective bud scales and older leaf primordia, tightly folded or partially unfolded architecture with lamina not yet fully expanded, very pale green to bronze coloration reflecting minimal chlorophyll development and high anthocyanin concentrations providing photoprotection, extremely soft delicate texture composed primarily of rapidly dividing and expanding cells, virtually no tissue differentiation beyond incipient procambial strands representing future veins, non-functional stomata and complete absence of cuticle development, negligible photosynthetic capacity contributing essentially zero carbon gain while representing major resource sink consuming carbohydrates and nitrogen for rapid cell division and initial expansion, and critical positional role as the newest addition to shoot photosynthetic surface area that will progressively mature and contribute function as shoot extension continues displacing this leaf from terminal position. Terminal apical leaves represent the immediate output of shoot apical meristem activity, emerging continuously during active shoot extension creating the progressive addition of new leaves that drives total canopy leaf area increase, consuming substantial resources relative to their tiny current size due to high metabolic demands of cell division and expansion, and showing maximum vulnerability to environmental stresses, pest damage, and mechanical injury due to extreme tissue fragility and absence of protective structures.
Terminal apical leaves represent the absolute developmental frontier of shoot extension, distinguished from other apical leaves by extreme youth, minimal development, and position at or immediately behind the shoot apex. These leaves show the shortest elapsed time since emergence and greatest developmental immaturity among all visible shoot leaves, with characteristics closely resembling emerging young leaves but specifically occupying terminal position. The terminal leaf designation is transient and dynamic during active shoot extension, with each leaf occupying terminal position briefly before subsequent leaf emergence displaces it to sub-terminal positions. Shoot extension cessation fixes terminal position, with the final terminal leaf representing the last leaf initiated before growth arrest. Terminal apical leaves show maximum vulnerability to all stresses and damage sources, with frost, heat, desiccation, pest attack, or mechanical damage commonly affecting these tender tissues creating permanent deformations or complete loss requiring subsequent leaves to assume terminal function. Management practices including shoot topping intentionally remove terminal apical leaves along with several sub-terminal leaves, terminating shoot extension and redirecting resources from vegetative growth toward fruit development and lateral shoot suppression.
Spitzenblatt
extreme terminal leaf
neuestes oberes Weinblatt
newest apical leaf
terminales apikales Blatt
tip leaf
Based on Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 60-92 Palliotti & Cartechini (2000) "Developmental changes in gas exchange activity in flowers, berries, and tendrils of field-grown Cabernet Sauvignon" American Journal of Enology and Viticulture 51=317-323 Intrieri et al. (2008) "Grapevine growth and development" in Biology of the Grapevine, Cambridge University Press pp. 44-81 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 79-102.
Terminal Apical Leaf
A terminal apical lobe on a mature leaf at node 7 of Vitis vinifera cv. Mourvèdre in Bandol, France, positioned at the absolute blade apex representing the distal terminus of the median vein, measuring 7.8 cm length from sinus base to apex determining the total leaf length of 17.2 cm, width of 4.5 cm creating elongated terminal lobe shape with length=width ratio of 1.73, separated from upper lateral lobes by very deep upper lateral sinuses of 4.2 cm (left) and 4.0 cm (right) extending to within 2.8 cm of the petiolar junction creating pronounced terminal lobe isolation, median vein extending through lobe center for the full 7.8 cm terminating 0.3 cm from the apex, exhibiting acute apex with approximately 35-degree point angle and distinctive acuminate extended tip characteristic of this cultivar, lobe margins showing slightly concave curvature in upper portions transitioning to convex in lower regions, lobe area of 28.5 cm² representing 32% of total blade area making it the dominant lobe, prominent elongated appearance creating characteristic Mourvèdre leaf silhouette recognizable from substantial distances, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing maximum terminal apical lobe development diagnostic for deeply dissected cultivars.
An apical lobe representing the absolute terminal extent of the leaf blade in Vitis vinifera, characterized by position at the distal blade apex as the final extension of the median vein axis, maximum reach of the blade in the apical direction determining total leaf length, prominent development with substantial size often representing 25-35% of total blade area, well-defined separation from adjacent upper lateral lobes through deep upper lateral sinuses creating distinct terminal lobe boundaries, elongated morphology with length typically exceeding width by 1.2-1.8× ratios, median vein extending through the lobe center to terminate near or at the lobe apex, acute to obtuse apex configurations with cultivar-specific point characteristics, and functional role as the primary terminal blade extension contributing substantially to overall leaf area and determining maximum vertical dimension affecting light interception patterns and canopy architecture. Terminal apical lobes represent the maximum expression of apical lobe development, occurring in cultivars with pronounced lobe dissection creating well-separated terminal extensions, exhibiting substantial dimensions reflecting developmental priority and functional importance, and serving as prominent diagnostic features enabling cultivar identification often from considerable distance based on distinctive terminal lobe shapes.
Terminal apical lobes represent the maximum development extreme of apical lobes, distinguished from less prominent apical lobes by substantial size, pronounced separation, and elongated morphology. The terminal designation emphasizes the absolute distal blade extent rather than simply apical position. Terminal apical lobes occur primarily in deeply dissected cultivars showing pronounced lobe development (Mourvèdre, some Syrah clones, deeply lobed Muscat types), while weakly lobed cultivars show poorly developed apical lobes barely distinguishable as separate structures. Deep upper lateral sinuses are prerequisite for terminal lobe prominence, with sinus depths often exceeding 60-70% of distance toward median vein creating strong lobe isolation. Elongated morphology with length substantially exceeding width contributes to distinctive appearance, with some cultivars showing extreme elongation creating finger-like terminal lobes. Apex characteristics including acute points and acuminate tips enhance terminal appearance and diagnostic utility. Functional contribution involves substantial area provision and maximum vertical reach affecting canopy light interception. Developmental programming creates preferential terminal lobe growth in deeply dissected cultivars, contrasting with balanced lobe development in weakly lobed varieties. Ampelographic utility is high, with prominent terminal apical lobes serving as immediately recognizable identification features enabling field identification without detailed examination.
distal terminal lobe
distaler Endlappen
final apex lobe
terminaler Apikallappen
ultimate apical lobe
ultimativer Spitzenlappen
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 90-135 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 79-102.
Terminal Apical Lobe
In vigorous Cabernet Sauvignon lateral shoots in Napa Valley at BBCH 65, terminal flower clusters occasionally develop at lateral shoot apices following early-season shoot tipping, exhibiting 8-12 cm length with 150-280 flowers, developing 4-7 days later than the primary shoot's lateral inflorescences at nodes 3-4, and showing 15-25% lower fruit set potentially due to competition with continued lateral shoot growth and less favorable hormonal environment. Pinot Noir shoots subjected to severe water stress during early development occasionally exhibit terminal inflorescence conversion at the primary shoot apex, producing small terminal flower clusters (6-10 cm, 80-150 flowers) that bloom synchronously with normal lateral clusters but exhibit 20-35% higher shatter rates potentially due to increased wind exposure at canopy periphery.
A terminal flower cluster is a flower cluster positioned at the shoot apex or at branch termini, representing flowers borne on inflorescences located at terminal growth positions rather than at lateral (axillary) positions along the shoot, with terminal positioning affecting resource allocation, developmental timing, and exposure to environmental conditions in ways that can influence flower development, fruit set success, and final berry quality. Terminal flower clusters occur on inflorescences that develop from apical meristems at shoot tips or lateral branch tips, contrasting with the more common lateral inflorescences that develop from axillary buds along the shoot axis. In Vitis vinifera, true terminal inflorescences are relatively uncommon in standard cultivars, as the typical architecture involves lateral inflorescences at nodes 3-5 on primary shoots, with the shoot apex (terminal position) typically producing a tendril or continuing vegetative growth. However, terminal flower clusters can occur in several contexts= (1) on lateral shoots where the lateral shoot tip converts from vegetative to reproductive development, producing an inflorescence at the lateral shoot apex (2) on primary shoots in certain wild Vitis species or unusual cultivar selections where apical meristems undergo floral conversion (3) on shoots subjected to environmental stress or hormonal disruption that triggers terminal flowering (4) in secondary or tertiary crop situations where normally vegetative shoot tips convert to reproductive development late in the season. Terminal flower clusters exhibit distinctive characteristics resulting from their apical position= enhanced access to actively translocated photosynthates and nutrients due to proximity to source leaves and developing sink strength, earlier developmental initiation in some cases due to absence of intervening lateral structures, or conversely delayed development if terminal conversion occurs after lateral inflorescence development, greater exposure to environmental extremes (wind, temperature fluctuations, radiation) due to apical positioning at canopy periphery, and altered hormonal environments due to proximity to actively growing shoot tips with high auxin production. The resource allocation to terminal flower clusters can differ from lateral clusters, with terminal positions potentially receiving preferential allocation during active shoot growth or conversely experiencing resource competition from the actively growing shoot tip. Terminal cluster development may proceed asynchronously with lateral cluster development on the same shoot, creating temporal heterogeneity in flowering and subsequent fruit maturation. Fruit set patterns in terminal flower clusters can differ from lateral clusters, with some studies indicating higher or lower fruit set depending on resource allocation patterns and competition with vegetative growth. The environmental exposure of terminal flower clusters, particularly increased light interception and wind exposure at canopy periphery, may affect flower development, pollination success, and susceptibility to environmental stresses during critical developmental periods.
Terminal_Flower_Cluster is distinguished from the standard lateral (axillary) flower clusters by position (terminal/apical versus lateral/axillary), developmental context (shoot tip conversion versus lateral bud development), and potential functional differences in resource allocation, environmental exposure, and developmental timing. True terminal inflorescences are relatively uncommon in standard Vitis vinifera wine cultivars under normal cultural conditions, making this class primarily relevant for lateral shoot terminal clusters, stress-induced terminal flowering, or secondary crop situations. The developmental and physiological significance of terminal versus lateral positioning deserves research attention, as terminal clusters may exhibit different responses to environmental conditions, cultural practices, and crop load management strategies. Sub-classification into Apex_Terminal and End_Branch_Terminal reflects positional distinctions within terminal flower cluster contexts.
Blütentraube an Triebspitze
apical flower cluster
endständige Blütentraube
shoot-tip flower cluster
terminale Blütentraube
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 85-94 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 72-86 Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109.
Terminal Flower Cluster
A terminal main apical shoot on a young horizontal cordon arm of Vitis vinifera cv. Merlot in its second training year in Pomerol, France, having extended 180 cm from April budbreak through September without topping intervention, with 24 nodes bearing progressively smaller leaves from base (150 cm² average) to tip (40 cm² current leaves), showing active apical meristem producing new leaves at 2-day plastochron intervals in early September, suppressing lateral shoot development completely in basal 18 nodes while maintaining vigorous primary growth that will establish the full 2-meter cordon length required by the training system design.
A main apical shoot that occupies and maintains the absolute terminal position of a woody perennial structure throughout the growing season in Vitis vinifera, representing uninterrupted continuation of the primary growth axis without artificial termination through topping or natural cessation through damage, characterized by retention of an active apical meristem capable of continued internode production and leaf initiation until growth cessation induced by photoperiod signals, nutrient limitations, or onset of dormancy preparation. Terminal main apical shoots express maximum apical dominance effects, creating the strongest basipetal suppression of lateral bud outgrowth and exhibiting preferential sink strength for photosynthates, mineral nutrients, and water resources compared to all other shoot types on the same vine. These shoots play crucial roles in framework establishment during vineyard development years, with their growth determining the speed of cordon or trunk extension toward target dimensions, and in mature vineyards may be intentionally maintained in low-vigor situations or removed through topping in high-vigor conditions to balance vegetative and reproductive growth.
Terminal main apical shoots are distinguished from leader main apical shoots by the absence of intentional or damage-induced termination events. In commercial viticulture, terminal main apical shoots are relatively uncommon in mature vineyards due to routine summer topping practices, but are standard in establishment-phase vineyards where framework development takes priority. The decision to maintain or terminate apical shoots represents a fundamental canopy management choice, balancing vegetative framework needs against fruit quality optimization. Shoots may lose terminal status through topping, herbivore damage, disease infection, or mechanical breakage, after which lateral shoots may assume dominance but are not reclassified as terminal main apical shoots.
continuous terminal shoot
durchgehender Endtrieb
ungekürzter Hauptspitzentrieb
unterminated apical shoot
untipped apical shoot
untopped main shoot
Based on Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Intrieri et al. (1998) "The effects of summer pruning and its time of application on canopy efficiency in Vitis vinifera L." Acta Horticulturae 512=119-126 Palliotti & Cartechini (2000) "Cluster thinning effects on yield and grape composition in different grapevine cultivars" Acta Horticulturae 512=111-119 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 215-238.
Terminal Main Apical Shoot
A terminal primary inflorescence at the apex of a short fruiting spur on Vitis vinifera cv. Pinot Noir in Champagne, France, positioned 1.8 cm behind the shoot apical meristem representing terminal placement, measuring 11.5 cm length representing reduced dimensions compared to typical node 3 primary inflorescences (16-18 cm), exhibiting compound paniculate architecture with central rachis bearing 3-4 orders of branching, bearing approximately 280 flowers organized across the structure, positioned as the distal-most inflorescence following the typical node 3 primary inflorescence located 8 nodes below, developing from the terminal bud of this short 2-node spur with vegetative extension terminated by reproductive development, showing slightly delayed phenological development at BBCH stage 59 (inflorescence fully developed, flowers separating) while lower inflorescence is at stage 63 (30% flowers open), terminal position creating exposure to maximum light and air circulation but also increased wind stress vulnerability, occurring on this Pinot Noir clone showing characteristic tendency for terminal inflorescence production on short spurs, measured in early June during pre-bloom to early bloom period, representing terminal primary inflorescence architecture and positioning.
A primary inflorescence positioned at or very near the shoot apex representing terminal placement in Vitis vinifera, characterized by location at the shoot tip or within 2-5 cm of the apical meristem, occurrence as the distal-most inflorescence on the shoot, development from terminal or near-terminal bud positions, spatial association with the shoot apex and youngest vegetative structures, typical appearance on determinate shoots where reproductive development terminates vegetative extension, smaller dimensions compared to standard node 3 primary inflorescences typically 8-15 cm length reflecting late initiation or resource competition with shoot extension, and occurrence patterns showing cultivar and environmental variation with some varieties rarely producing terminal inflorescences while others show consistent terminal reproductive development particularly on short fruiting spurs. Terminal primary inflorescences represent alternative positioning patterns, developing from terminal bud positions rather than the standard lateral node 3 position, exhibiting variable expression depending on shoot determinacy and reproductive development patterns, and creating architectural variations affecting canopy organization and fruit distribution.
Terminal primary inflorescences are distinguished from standard lateral primary inflorescences by apical rather than node 3 positioning. The terminal designation emphasizes shoot apex location rather than simply being the last inflorescence in a sequence, though terminal inflorescences are necessarily the distal-most structures. Occurrence varies dramatically among cultivars with some showing frequent terminal inflorescence production particularly on short spurs (Pinot varieties, Chardonnay on spurs), while others rarely produce terminals developing inflorescences only at standard lateral positions. Shoot determinacy relates to terminal inflorescence development, with determinate shoots showing reproductive development terminating vegetative extension creating terminal inflorescences, while indeterminate shoots maintain vegetative apical meristems preventing terminal reproductive development. Size reduction compared to standard primaries reflects late initiation, resource competition with shoot extension, or limited space and resource allocation in terminal positions. Phenological timing often shows slight delay compared to lateral primaries reflecting later initiation or development. Exposure characteristics differ from lateral inflorescences with terminal positions receiving maximum light and air circulation but experiencing greater wind stress and potentially temperature extremes. Management implications include vulnerability to shoot tip removal through mechanical damage, animal browsing, or cultural practices affecting terminals more than protected lateral inflorescences. Transformation to cluster creates terminal cluster positioning affecting harvest accessibility and cluster sun exposure.
Spitzenposition Blütenstand
apical position inflorescence
distal primary inflorescence
distale Hauptinfloreszenz
shoot tip inflorescence
terminale primäre Infloreszenz
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 286-318 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 249-278 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 469-502.
Terminal Primary Inflorescence
Cabernet Sauvignon in Napa Valley shows terminal shoot elongation from late July through mid-August at BBCH 38-41 over 3 weeks, with shoots reaching 2400-2800 mm final length through elongation rate declining from 15-20 mm per day to cessation, shoot apex becoming necrotic, under warm climate conditions of 26-32°C mean temperature, moderate water stress from limited irrigation, and nutrient redistribution to developing fruit typical of this vigorous late-season cultivar achieving growth cessation as veraison initiates
Terminal shoot elongation represents the final growth phase where elongation rate approaches zero and apical meristem activity ceases, marking transition from active vegetative growth to shoot maturation through lignification and reserve accumulation. This developmental stage typically occurs during late July through August in temperate regions corresponding approximately to BBCH 38-41 as berries reach full size and begin color change, with elongation rate declining from 10-20 mm per day to <5 mm per day and eventually ceasing over 2-3 week period adding final 100-300 mm shoot length. Terminal elongation represents asymptotic approach to zero growth of sigmoid growth curve as multiple limiting factors combine with developmental programming to arrest meristematic activity and cell expansion, with shoot apex typically transforming from active vegetative meristem to senescent or necrotic terminal structure. The shoot apex ceases producing new leaf primordia with apical meristem losing organized structure, cells losing division capacity, and apex often becoming necrotic or senescing creating characteristic appearance of growth cessation, though some cultivars may maintain green non-growing apex. Final internodes produced during terminal elongation show markedly reduced length typically 15-40 mm reflecting minimal resource allocation to elongation and changed hormonal environment strongly favoring reproductive over vegetative development. Cell expansion in subapical zones operates at minimal rates as severe water stress or hormonal signals nearly eliminate cell wall extensibility, with expansion zone reduced to <15 mm behind apex and cells achieving small final dimensions of 100-250 micrometers in terminal internodes. Final leaves emerging show substantially reduced size typically 30-50% dimensions of mid-season leaves and may show reduced expansion or development indicating resource limitation and changed developmental priorities, with terminal leaves sometimes failing to fully expand before growth ceases. Photosynthetic capacity in mature canopy remains substantial supporting fruit development and reserve accumulation though new photosynthetic area addition ceases with growth termination, with assimilate production directed almost entirely toward reproductive development and storage with negligible vegetative growth demand. Carbon allocation during terminal elongation shows 85-95% fruit and reserve accumulation with only 5-15% supporting minimal remaining shoot growth, with cessation of elongation redirecting all photosynthetic production toward fruit maturation, root growth, and substantial reserve accumulation in woody tissues. Vascular system in shoot apex may show incomplete differentiation as growth arrests before full maturation, with functional xylem and phloem in mature shoot portions maintaining resource transport for fruit development and reserve movement though no longer supporting shoot growth. Water demand from mature canopy continues at 300-500 mg per hour though growth component essentially zero, with tissue water status often at seasonal minimum in many regions as late summer conditions and fruit transpiration create high demand, with moderate to severe water stress typical and contributing to growth cessation. Nutrient demand shifts entirely to fruit with nitrogen, phosphorus, potassium and other nutrients moving predominantly toward berries, with shoot apex and young leaves potentially showing nutrient deficiency symptoms as mobile nutrients redistribute, contributing to growth cessation and terminal leaf senescence. Metabolic activity in shoot apex declines substantially with respiration approaching mature tissue levels as growth ceases, while mature leaf photosynthesis maintains efficient operation supporting fruit development though possibly declining slightly as water stress or nutrient redistribution affects function. Protein synthesis in apical tissues nearly ceases with growth arrest, while mature tissues increasingly produce reserve storage proteins and enzymes involved in reserve accumulation and tissue maturation processes. Hormonal regulation shows minimal auxin production from ceased apex eliminating growth promotion, elevated abscisic acid from water stress and developmental signals promoting growth cessation, possible increased ethylene from stress or fruit ripening contributing to senescence processes, and depleted cytokinins as root production declines. Temperature during terminal elongation includes continued warm to hot conditions typical of late summer with means often 24-30°C in many regions, with high temperatures contributing to growth cessation through water stress and metabolic efficiency reduction though not typically limiting fruit maturation processes. Light intensity begins declining from summer maximum as day length shortens and solar angle lowers, though generally remaining adequate for photosynthesis and not limiting factor in growth cessation. Water availability often at seasonal minimum during terminal elongation in many regions with late summer drought conditions, with moderate water stress beneficial for fruit quality development while severe stress risks premature senescence and incomplete fruit maturation. Nutrient availability particularly nitrogen largely depleted from vegetative tissues as mobile nutrients redistribute to fruit, with shoot tissues showing declining nitrogen concentrations and possible deficiency symptoms in young tissues as growth ceases. The process typically spans 2-3 weeks with progressive growth rate reduction to zero, with final cessation timing varying based on environmental stress severity, fruit load and developmental stage, and cultivar growth habit characteristics. Cultivar variation appears in timing of growth cessation relative to fruit development stages, whether apex maintains viable appearance or necrotizes, final shoot length achieved, and sensitivity to environmental factors affecting cessation timing. Regional timing shows terminal elongation and cessation occurring late July-August in warm climates, August-September in temperate regions, and August-September in cool areas, typically spanning berry development completion through beginning ripening when fruit demands are maximum. Viticultural management considerations include monitoring growth cessation timing relative to fruit development ensuring adequate vegetative development for fruit maturation while avoiding excessive late growth, managing water stress promoting timely cessation and fruit quality development without excessive stress causing premature senescence, assessing post-cessation canopy condition for fruit maturation and reserve accumulation capacity, planning late-season leaf removal and cluster thinning operations based on canopy function, and evaluating year's vegetative growth outcomes informing subsequent season management adjustments. Premature growth cessation from excessive stress reduces photosynthetic capacity and duration potentially limiting fruit maturation and reserve accumulation, while delayed cessation from excessive resources maintains vegetative growth competing with fruit quality development. Research applications include analyzing mechanisms of growth cessation including hormonal regulation and developmental programming, quantifying final assimilate partitioning patterns during fruit maturation, evaluating management effects on cessation timing and fruit quality outcomes, modeling seasonal growth patterns for cultivar characterization and management decision support, and assessing climate change impacts on growth duration and cessation timing affecting productivity and quality across regions.
Terminal_Shoot_Elongation is fifth and final Shoot_Elongation subphase, distinguished by growth approaching and reaching zero as apical meristem activity ceases. Marks definitive end of vegetative elongation phase. Transitions focus to Shoot_Lignification and fruit maturation processes. Timing relative to fruit development critically affects fruit quality potential through effects on photosynthetic capacity duration and resource allocation patterns.
Terminalphase
Triebausr eifung
Wachstumsabschluss
apical growth arrest
apikaler Wachstumsstillstand
growth cessation
terminal growth phase
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 258-265 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 175-178 Lorenz DH et al. (1995) Growth Stages of the Grapevine, Nachrichtenbl. Deut. Pflanzenschutzd. 47:193-200
Terminal Shoot Elongation
In Mourvèdre vineyards of Bandol at BBCH 57-59, terminal tertiary inflorescence branches measuring 0.5-1 cm length and bearing 3-6 flowers each appear on the apices of secondary branches on basal and lower primary branches, contributing to the extremely loose, multi-lobed cluster architecture with final berry numbers of 150-280 per cluster. Assyrtiko in Santorini exhibits modest terminal tertiary branches (0.3-0.8 cm, 2-4 flowers) creating moderately subdivided clusters. In compact-clustered Müller-Thurgau, terminal tertiary branches are completely absent, with secondary branches (where present) terminating directly in flower attachment.
A terminal tertiary inflorescence branch is a tertiary inflorescence branch that occupies the apical or terminal position on a secondary inflorescence branch, representing the distal-most third-order architectural element and typically exhibiting the most abbreviated dimensions and minimal flower number among all tertiary branches while contributing to the fine tapering of secondary branch termini. Terminal tertiary branches occupy the apex of secondary branches, typically positioned as the last tertiary branch to develop and the most distal from the primary branch junction, often located 0.3-1.5 mm from the secondary branch tip. These branches represent the latest-initiated tertiary branches during inflorescence primordium development and consequently show the least developmental elaboration, often appearing as small, unbranched flower-bearing axes. Branch length typically ranges from 0.2-1.2 cm in loose-clustered cultivars, with basal diameter often 0.2-0.4 mm, representing the minimum dimensions observed among flower-bearing branches in the entire inflorescence architecture. Terminal tertiary branches characteristically bear few flowers (2-8 flowers per terminal tertiary, often only 2-4), almost always through direct attachment without quaternary subdivision, contributing minimally to total flower number but creating the delicate, finely subdivided appearance of extensively branched inflorescences. The abbreviated character of terminal tertiary branches creates pointed terminations of secondary branches, contributing to the overall lacy or filigree appearance of extremely loose cluster morphologies such as those of Grenache, Mourvèdre, and some Muscat types. Vascular anatomy of terminal tertiary branches is minimal, with vessel diameters often only 12-25 μm and limited phloem development, reflecting the modest resource requirements of the small flower complement. Terminal tertiary branch development is among the latest inflorescence events, often not becoming clearly distinct until BBCH 57-59, with some terminal tertiaries still initiating while basal flowers on the same inflorescence are beginning anthesis. In moderately branched cultivars, terminal tertiary branches may be completely absent, with secondary branches terminating in direct flower attachment or in short, unbranched axes difficult to classify as distinct branches. The presence and degree of development of terminal tertiary branches serves as an indicator of overall inflorescence developmental vigor and favorable previous-season growing conditions, while their absence or extreme reduction can signal developmental constraints or compact-clustered cultivar genetics.
Terminal_Tertiary_Inflorescence_Branch is distinguished from Lateral_Tertiary_Inflorescence_Branch by position (terminal/apical versus lateral/basal on the parent secondary branch), reduced dimensions, later development, and minimal flower number. The 'terminal' designation specifically refers to the apical-most tertiary branch, typically singular per secondary branch when present. These branches represent the finest architectural detail in grapevine inflorescence structure and are present only in loose to very loose cluster types. Their presence and development correlates with overall inflorescence complexity and can be used as a quantitative indicator of cluster looseness in ampelographic description. Extreme reduction or absence does not necessarily indicate developmental abnormality but may represent normal compact-clustered cultivar morphology.
Endast dritter Ordnung
apical tertiary branch
distal third-order branch
endständiger tertiärer Ast
terminaler Tertiärast
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 93-94 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 174-177 Galet, P. (2000). A Practical Ampelography, pp. 49-52.
Terminal Tertiary Inflorescence Branch
Chardonnay terminal vascular bundle in receptacle region at véraison (BBCH 83) shows reduction from parent bundle diameter 0.8-1.2 mm to terminal bundle diameter 0.3-0.5 mm over 1.5-2.0 mm distance, contains 2-3 xylem vessels 15-22 μm diameter with functional water transport, 3-5 phloem sieve tubes with active sugar transport at 0.3-0.6 mg sucrose/hour per bundle, and increasing parenchyma proportion from 30% proximally to 60% at terminal region
A terminal vascular bundle is a vascular bundle at or near its termination point, characterized by reduced diameter, simplified organization, and transition from organized vascular tissue to connection with surrounding non-vascular tissues, representing the distal portions of vascular traces in brush, receptacle, or berry basal regions. Terminal vascular bundles show characteristic anatomical features distinguishing them from non-terminal bundles= reduced vascular bundle diameter (terminal bundles typically 30-60% of parent bundle diameter), simplified vascular organization (fewer xylem vessels and phloem elements compared to proximal bundle regions), modified conducting element characteristics (terminal xylem vessels showing reduced diameter 10-25 μm versus 25-50 μm in main bundles, terminal phloem with fewer sieve tubes), increased proportion of parenchyma relative to conducting tissues, and potential loss of distinct vascular bundle sheath or cambium. Terminal vascular bundle xylem characteristics include= reduced vessel element number (1-3 vessels in terminal bundles versus 5-15 in main pedicel bundles), decreased vessel diameter (terminal vessels 10-25 μm versus 25-50 μm in proximal regions), modified vessel wall structure (potentially thinner secondary walls, altered pit arrangement), and increased proportion of xylem parenchyma or transitional cells. Terminal vascular bundle phloem characteristics include= reduced sieve tube number (1-4 sieve tubes in terminal bundles versus 8-20 in main bundles), smaller sieve tube diameter, reduced companion cell abundance, and potential increased callose deposition in terminal phloem elements. Terminal vascular bundle distribution in berry brush and receptacle includes= primary terminal bundles representing endpoints of main pedicel vascular bundles (2-4 per berry at receptacle level), secondary terminal bundles representing endpoints of smaller vascular traces (6-15 per berry distributed in receptacle and basal berry tissues), and fine terminal bundles representing smallest vascular traces extending into peripheral berry tissues. Terminal vascular bundle function varies by developmental stage= during active berry growth (BBCH 73-83) terminal bundles maintain active xylem and phloem transport delivering water, minerals, and sugars to berry tissues, during late ripening (BBCH 85-89) terminal xylem bundles may show progressive embolism or occlusion reducing water transport, terminal phloem bundles maintain sugar delivery but at potentially reduced rates, and at full maturity terminal bundles may become largely non-functional with occluded xylem and callose-blocked phloem. Terminal vascular bundle modifications during berry development include= progressive lignification of terminal bundle elements, potential xylem vessel occlusion through tyloses formation or pit membrane thickening, phloem callose deposition reducing sieve tube conductance, and structural changes associated with nearby abscission zone development potentially disrupting terminal bundle continuity. Terminal vascular bundle variation includes= terminal bundle number (4-20 per berry), terminal bundle length (0.5-3 mm from branching point to endpoint), degree of organization retention (some terminal bundles maintain organized structure while others show extensive disorganization), and functionality status (active versus occluded terminals).
Terminal_Vascular_Bundle emphasizes distal bundle characteristics and is distinguished from Vascular_Bundle_Termination (complete ending region) and End_Point_Vascular_Bundle (final conducting segment) by focus on reduced organization and transitional anatomy approaching termination point. Critical for understanding vascular supply diminishment toward berry periphery, transport capacity reduction in distal tissues, and developmental modifications in terminal vasculature during ripening.
Distal bundle
Distales Vaskulärbündel
Ending vascular bundle
Endständiges Gefäßbündel
Terminales Leitbündel
Pratt, C. (1971). Reproductive anatomy in cultivated grapes - a review. American Journal of Enology and Viticulture, 22(2), 92-109. | Chatelet, D. S., Rost, T. L., Shackel, K. A., & Matthews, M. A. (2008). The peripheral xylem of grapevine (Vitis vinifera). 1. Structural integrity in post-veraison berries. Journal of Experimental Botany, 59(8), 1987-1996.
Terminal Vascular Bundle
A terminal veinlet in an areolar center on a mature leaf at node 7 of Vitis vinifera cv. Merlot in Pomerol, France, representing the final vascular terminus within a 0.32 mm² areole bounded by surrounding veinlet network, measuring approximately 0.25 mm length extending from the areolar boundary veinlet toward the areole center, diameter of 0.06 mm barely visible under high magnification (40×), consisting of single tracheid (8 μm diameter) terminating in blind ending within mesophyll approximately 65 μm from the areole geometric center, terminal tracheid showing specialized end cell with thickened walls potentially serving as structural terminus, surrounded by 6-8 mesophyll cells in immediate vicinity representing the most hydraulically remote cells from major vascular network, these terminal cells located maximum 50-60 μm from the terminal veinlet ending representing ultimate diffusion distance, terminal veinlet providing final water delivery enabling these remote mesophyll cells to maintain adequate hydration for photosynthetic function, occurring within one of approximately 950 areoles per mm² each typically containing 1-2 terminal veinlets serving central regions, measured at BBCH stage 75 (berries pea-sized) in late June, representing the absolute terminal limit of vascular elaboration defining maximum diffusion distances and hydraulic efficiency.
A veinlet representing the absolute terminal endpoints of the vascular network where veins terminate within blade tissue in Vitis vinifera, characterized by position at the ultimate distal ends of veinlet branches representing the final vascular elaboration, very small diameter typically 0.04-0.08 mm at the minimum end of veinlet size range, minimal anatomy often consisting of single tracheid terminating in blind ending within mesophyll without connecting to other vascular elements, or occasionally terminating in specialized tracheid end cells, very short length typically 0.1-0.4 mm representing final branch segments, location within areolar centers serving mesophyll cells most distant from major veinlet networks, critical functional role providing water delivery to the most hydraulically remote mesophyll cells ensuring all photosynthetic tissues receive adequate water supply, and representing the anatomical limit of vascular elaboration beyond which water and solute movement occurs exclusively through cell-to-cell diffusion and transport. Terminal veinlets create the final vascular termini, developing as the last venation elements during late leaf expansion extending from established veinlet networks into remaining avascular regions, exhibiting blind endings or occasional anastomoses (connections) with other terminal veinlets creating isolated or connected terminal networks, and determining the ultimate mesophyll-to-vein diffusion distances that establish maximum hydraulic efficiency and photosynthetic capacity limits.
Terminal veinlets represent the ultimate endpoints of venation, distinguished from other veinlets by terminal position with blind endings or isolated connections rather than hierarchical branching continuation. The terminal designation emphasizes end position rather than hierarchical level, as terminals may arise from various veinlet orders. Blind endings represent true vascular termini where water delivery transitions to exclusively diffusive/transport processes through mesophyll cells, while some terminals show anastomoses connecting to other terminals creating isolated loops within areoles. Functional importance involves ensuring all mesophyll cells including those most remote from major vascular networks receive adequate water delivery, with terminal veinlet presence reducing maximum diffusion distances compared to areoles lacking terminals. Anatomical structure shows maximum simplification with single tracheids often representing the sole vascular element, potentially lacking phloem and serving purely xylem delivery function. Development timing involves very late expansion phases representing final vascular elaboration after establishment of primary veinlet networks. Density variation relates to overall veinlet density and areole size, with smaller areoles in high-density networks showing fewer terminals per areole while larger areoles in lower-density networks may contain more terminals serving the greater avascular areas. Terminal veinlet distribution affects spatial heterogeneity in mesophyll hydration status and photosynthetic capacity, with regions near terminals potentially showing enhanced function compared to remote regions.
Aderende
terminal vein ending
terminales Äderchen
ultimative Feinvene
vein terminus
Based on Sack & Holbrook (2006) "Leaf hydraulics" Annual Review of Plant Biology 57=361-381 Brodribb et al. (2007) "Leaf hydraulic capacity in ferns, conifers and angiosperms= impacts on photosynthetic maxima" New Phytologist 165=839-846 Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 243-268.
Terminal Veinlet
In Muscat of Frontignan berries in Languedoc, France during BBCH 81-89, terpene accumulation increases total monoterpene content from 1.5 to 11.2 mg/L over 45 days, with free monoterpenes rising from 0.3 to 1.6 mg/L and bound monoterpenes from 1.2 to 9.6 mg/L, establishing a free-to-bound ratio of 0.17 at harvest, while concurrent sesquiterpene accumulation reaches 85 µg/L total including 12 µg/L farnesol, under warm Mediterranean conditions averaging 27°C with intense solar radiation.
Terpene accumulation constitutes the progressive increase in terpenoid compound concentrations within Vitis vinifera berries during the ripening period, wherein the outputs of the methylerythritol phosphate biosynthetic pathway are retained as both free volatiles and glycosidically-bound precursors to establish the characteristic terpenoid pools that define aromatic grape variety identity and contribute floral, citrus, and spicy aromatic character to wines produced from these cultivars over the 35-50 day post-veraison maturation period. This process integrates the biosynthesis of diverse terpenoid classes including monoterpenes comprising C10 compounds such as linalool, geraniol, nerol, citronellol, and alpha-terpineol that provide the dominant floral-fruity character of aromatic varieties, sesquiterpenes comprising C15 compounds such as rotundone, farnesol, and nerolidol contributing spicy and woody nuances, and trace quantities of diterpenes and other higher terpenoids that may contribute subtle aromatic background complexity. The quantitative aspects of terpene accumulation show dramatic cultivar variation, with aromatic Muscat varieties accumulating total monoterpene pools of 5-15 mg/L at harvest maturity representing 50-100 fold higher concentrations than neutral varieties such as Chardonnay or Pinot Noir that typically accumulate 0.1-0.3 mg/L total monoterpenes under equivalent conditions. Cellular localization of terpene accumulation demonstrates tissue-specific patterns, with monoterpene concentrations 5-20 fold higher in exocarp compared to mesocarp tissues reflecting the preferential expression of terpene synthase genes in skin cells, while sesquiterpene accumulation may show different tissue distribution patterns depending on specific compound and cultivar. The temporal dynamics of terpene accumulation follow progressive increase throughout post-veraison ripening, with accumulation rates typically highest during mid-ripening when terpene synthase gene expression reaches peak levels, followed by continued but declining accumulation during late ripening as enzyme activities wane and potential degradation or volatilization may partially offset continued synthesis. Temperature during ripening influences terpene accumulation through effects on biosynthetic enzyme activities, substrate availability, and compound stability, with moderate temperatures of 20-28°C generally supporting optimal accumulation while temperature extremes may reduce biosynthetic efficiency or accelerate volatile loss through increased vapor pressure. The partitioning between free and glycosidically-bound forms during terpene accumulation reflects the balance between synthesis rates and glycosyltransferase activities, with typical free-to-bound ratios of 0.05-0.20 indicating that 80-95% of accumulated terpenoids exist in conjugated precursor forms requiring hydrolysis for aromatic release. Viticultural practices influence terpene accumulation through effects on vine physiology and cluster microclimate, with moderate water stress, appropriate crop load, and optimized light exposure generally enhancing accumulation compared to conditions of excess vigor or resource limitation.
Terpene_Accumulation is parent class under Aroma_Development representing terpenoid compound increase. Distinguishes from Volatile_Compound_Synthesis which describes biosynthetic processes. Contains subclasses Monoterpene_Synthesis, Sesquiterpene_Production, and Terpene_Pool_Development. Aromatic varieties accumulate 5-15 mg/L versus 0.1-0.3 mg/L in neutral varieties. Includes both free volatiles and glycosidic precursors. 80-95% typically in bound form.
Monoterpen-und-Sesquiterpen-Akkumulation
Terpen-Akkumulation
Terpen-Aufbau
Terpenoid-Anreicherung
monoterpene and sesquiterpene accumulation
terpene build-up
terpenoid accumulation
terpenoid pool increase
Martin DM et al (2010) Functional annotation of the grapevine terpene synthase gene family. BMC Plant Biology 10: 226 Marais J (1983) Terpenes in the aroma of grapes and wines. South African Journal of Enology and Viticulture 4: 49-58 Keller M (2020) The Science of Grapevines 3rd Ed pp 290-300
Terpene Accumulation
Terpene content quality in Mosel Riesling provides the chemical basis for the variety's distinctive floral-citrus-petrol aromatic profile, with free monoterpene concentrations of 0.5-1.5 mg/L dominated by linalool, α-terpineol, and citronellol, complemented by bound glycosides at 2-5 mg/L that release progressively during extended aging the gradual hydrolysis of bound terpenes and acid-catalyzed rearrangement to form TDN (1,1,6-trimethyl-1,2-dihydronaphthalene, the petrol compound) over years of bottle maturation explains the aromatic evolution of great Mosel Riesling from youthful floral freshness to the complex kerosene-inflected bouquet of aged examples.
Terpene_Content_Quality encompasses the chemical qualities characterizing the concentration and composition of terpenoid compounds in grape berries, capturing the isoprenoid-derived volatiles responsible for the floral, citrus, and spicy aromatic character of Muscat and related aromatic cultivars. Terpenes are built from five-carbon isoprene units through the mevalonic acid and methylerythritol phosphate pathways, with monoterpenes (C10, two isoprene units) dominating the grape terpene profile while sesquiterpenes (C15) and diterpenes (C20) contribute at lower concentrations. The primary grape monoterpenes include linalool (floral-citrus), geraniol (rose), nerol (rose-citrus), citronellol (citrus-rose), α-terpineol (floral-pine), and limonene (citrus), present in both free volatile form and as non-volatile glycosidic conjugates. The total monoterpene concentration distinguishes aromatic cultivars (greater than 1-6 mg/L free terpenes) from neutral cultivars (less than 0.5 mg/L), with Muscat varieties reaching the highest concentrations (4-8 mg/L). Terpene accumulation in berries occurs primarily in the skin, with concentration increasing during ripening and peaking near harvest before declining in overripe fruit due to metabolism and volatilization. The terpene profile (relative proportions of individual compounds) provides cultivar-specific fingerprints, with linalool-dominant profiles in Muscat Blanc versus geraniol-dominant profiles in Gewürztraminer. Environmental factors modulate terpene content, with moderate water stress and sun exposure generally promoting accumulation while excessive heat reduces concentration. Glycosylated terpenes constitute the bound aroma fraction that can be released during fermentation and aging, with bound-to-free ratios typically ranging from 2:1 to 10:1, representing substantial aroma potential beyond the immediate free volatile contribution.
Terpene_Content_Quality serves as the parent class for individual monoterpene compounds including Linalool, Geraniol, Nerol, Citronellol, and Alpha_Terpineol Content Qualities. The class primarily addresses monoterpenes as the dominant grape terpene class, though sesquiterpenes (rotundone providing peppery character in Shiraz) are also quality-relevant. The distinction between free and bound (glycosylated) terpenes is essential because they contribute differently to aroma perception—free terpenes are immediately odor-active while bound forms represent aroma potential. Terpene analysis employs gas chromatography with mass spectrometric or flame ionization detection following liquid-liquid extraction or solid-phase microextraction.
Isoprenoid-Duftkonzentration
Monoterpengehalt
Terpengehalt
Terpenoidaromaqualität
floral aroma compound quality
isoprenoid volatile concentration
monoterpene content
terpenoid aroma quality
Mateo JJ and Jiménez M (2000) Monoterpenes in Grape Juice and Wines. Journal of Chromatography A 881(1-2): 557-567 Wilson B et al. (1984) Demonstration of the Importance of Monoterpene Glycosides in Wine Flavour. American Journal of Enology and Viticulture 35(3): 113-116
Terpene Content Quality
Terpene glycosides typically exceed free monoterpenes by 3-10 fold in grape berries, with the bound/free ratio increasing during ripening from 2-3 at veraison to 5-15 at harvest maturity. Muscat of Alexandria contains 8-15 mg/L total bound monoterpenes versus 2-4 mg/L free forms, while Riesling shows 2-5 mg/L bound with disaccharides > trisaccharides).
A monoterpene accumulation disposition that is the inherited capacity of a Vitis vinifera plant to conjugate free monoterpenes with sugar moieties (glucose, arabinose, rhamnose, apiose) through glycosyltransferase activity, creating a non-volatile, water-soluble bound aroma pool that serves as a reservoir for volatile release during winemaking, fermentation, and aging.
Terpene glycoside accumulation creates the "hidden" aroma potential of grapes that is progressively revealed during winemaking and aging. The glycosidically bound pool is non-volatile and sensorially inactive until hydrolyzed, providing extended aroma release over time. Cultivar differences in glycosylation patterns affect both the magnitude of bound pool and the kinetics of release. The enzymatic specificity of yeast β-glucosidases means that different fermentation conditions and yeast strains produce different monoterpene release patterns from the same glycoside pool.
Glykosid-Bindung
Terpenglykosid-Akkumulation
bound terpene capacity
gebundene Terpene
glycosylated aroma trait
terpene glycoconjugate disposition
Williams PJ et al. (1982) Synthesis of monoterpene hydrocarbons from tritiated monoterpene alcohols in non-Muscat and Muscat grape berries and wine. Journal of Agricultural and Food Chemistry 30:1219-1223 Günata Z et al. (1985) The aroma of grapes. I. Extraction and determination of free and glycosidically bound fractions of some grape aroma components. Journal of Chromatography A 331:83-90
Terpene Glycoside Accumulation Disposition
In Zibbibo berries in Pantelleria, Sicily during BBCH 83-89, terpene pool development establishes total monoterpene pools increasing from 2.8 to 13.5 mg/L over 42 days, with free terpenoid pool reaching 1.9 mg/L and bound pool 11.6 mg/L representing 86% in precursor form, combined with sesquiterpene pool of 72 µg/L, establishing total terpenoid aromatic potential 7.1-fold greater than free volatile content under hot Mediterranean conditions with daytime temperatures exceeding 32°C moderated by maritime influence and cool nights of 18°C.
Terpene pool development describes the cumulative establishment of total terpenoid reservoirs within Vitis vinifera berries during the ripening period, wherein the integrated outputs of monoterpene and sesquiterpene biosynthetic pathways accumulate as both free volatiles and glycosidically-conjugated precursors to create the composite terpenoid pools that determine the aromatic capacity and varietal identity of harvested grapes for subsequent expression during winemaking processes. This process integrates multiple terpenoid compound classes into unified pool metrics, with total monoterpene pools comprising 2-15 mg/L in aromatic varieties distributed among linalool, geraniol, nerol, citronellol, alpha-terpineol, and related compounds, total sesquiterpene pools contributing 20-150 µg/L across various cultivars, and the combined free and bound fractions establishing the complete terpenoid potential available for immediate aromatic expression and subsequent precursor release. The dynamics of terpene pool development show variety-specific accumulation trajectories, with aromatic Muscat varieties demonstrating continuous linear accumulation throughout ripening achieving 10-15 mg/L total monoterpenes, while neutral varieties such as Chardonnay reach plateau levels of 0.2-0.4 mg/L by mid-ripening with minimal subsequent increase, reflecting fundamental genetic differences in biosynthetic capacity and pathway regulation. The balance between free and bound terpenoid pools evolves during ripening, with the bound fraction typically increasing from 75-80% at early post-veraison to 85-95% at harvest as glycosyltransferase activities conjugate newly synthesized terpenoids throughout the maturation period, establishing the precursor-dominated composition characteristic of ripe aromatic grapes. Environmental factors influence terpene pool development through effects on both biosynthetic rates and glycosylation efficiency, with moderate temperatures of 20-26°C and appropriate light exposure supporting optimal pool accumulation, while water stress may enhance terpenoid pathway activity through stress-responsive secondary metabolism induction. Viticultural practices targeting terpene pool development include canopy management optimizing cluster light exposure for enhanced terpene synthase expression, irrigation strategies imposing controlled water deficit that activates secondary metabolism, and harvest timing decisions capturing maximum pool development before potential late-season degradation or dilution through continued berry expansion. The relationship between terpene pool size at harvest and wine aromatic expression depends on subsequent release efficiency, with winemaking decisions regarding skin contact duration, enzyme additions, and yeast strain selection determining what fraction of accumulated potential is converted to sensory-active free volatiles in finished wines.
Terpene_Pool_Development is subclass of Terpene_Accumulation representing cumulative terpenoid reservoir establishment. Distinguishes from individual synthesis processes by emphasizing integrated accumulation. Sibling to Monoterpene_Synthesis and Sesquiterpene_Production. Integrates mono- and sesqui-terpenes into composite potential. Bound fraction increases from 75-80% to 85-95% during ripening. Variety-specific trajectories reflect genetic capacity.
Terpen-Pool-Entwicklung
Terpen-Reservoir-Entwicklung
Terpenoid-Pool-Etablierung
cumulative terpene accumulation
kumulative Terpen-Akkumulation
terpene reservoir development
terpenoid capacity building
terpenoid pool establishment
Wilson B et al (1984) Determination of free and bound monoterpenes in Muscat grapes. American Journal of Enology and Viticulture 35: 95-100 Williams PJ et al (1989) Glycosidic precursors of varietal grape and wine flavor. ACS Symposium Series 388: 35-48 Keller M (2020) The Science of Grapevines 3rd Ed pp 295-305
Terpene Pool Development
Terpene profile for Muscat Blanc à Petits Grains from Rutherglen, Victoria (2024 vintage) documents free monoterpenes dominated by linalool (850 µg/L, OAV 34), geraniol (420 µg/L, OAV 12), nerol (180 µg/L, OAV 4.5), α-terpineol (95 µg/L, OAV below threshold), and citronellol (75 µg/L, OAV 1.9), with total free monoterpenes of 1720 µg/L and bound glycosidic terpenes of 4200 µg/L (as linalool equivalents), yielding a free-to-bound ratio of 0.41 indicating substantial unreleased aromatic reserve. Terpene profile for Riesling from the Mosel (2024 vintage) records a more restrained pattern with free linalool 85 µg/L, α-terpineol 42 µg/L, citronellol 38 µg/L, geraniol 28 µg/L, and notably free 1,8-cineole 12 µg/L, with total free monoterpenes of 245 µg/L but bound precursors of 1850 µg/L (free-to-bound ratio 0.13), reflecting the characteristic Riesling pattern where the majority of aromatic potential is locked in glycosidic form and progressively released during bottle aging.
Terpene profile is an aromatic analysis data entity that records the concentration and composition of monoterpenoid (C₁₀) and sesquiterpenoid (C₁₅) compounds in grape berries and must, encompassing both the free volatile fraction (odor-active molecules directly contributing to aroma) and the glycosidically bound precursor fraction (non-volatile glycoconjugates representing latent aromatic potential), determined by GC-MS analysis of free volatiles and by enzymatic or acid hydrolysis followed by GC-MS for bound precursors, expressed in micrograms per liter (µg/L) or micrograms per kilogram of berries (µg/kg), and providing the definitive chemical characterization of floral, citrus, and muscat-type aromatic identity in terpene-dependent grape cultivars.
Terpene profile data provides the molecular explanation for the aromatic hierarchy among grape cultivars — Muscat varieties accumulate monoterpenes to concentrations 10-50× above sensory thresholds (intensely aromatic), semi-aromatic varieties (Riesling, Gewürztraminer, Torrontés) reach 1-5× thresholds (moderately aromatic), while non-aromatic varieties (Chardonnay, Cabernet Sauvignon) produce terpenes at sub-threshold levels that contribute only to overall aromatic complexity without dominant floral character. The genetic basis involves the terpene synthase (VvTPS) gene family, with cultivar-specific expression patterns of VvTPS-a (sesquiterpenes), VvTPS-b (monoterpene alcohols), and VvTPS-g (monoterpene alcohols and geraniol) determining the terpenoid phenotype. The bound glycosidic fraction typically exceeds free terpenes by 2-10× and represents the latent aromatic potential that is progressively released through acid-catalyzed hydrolysis during wine aging (contributing to the development of aged Riesling complexity including kerosene-like TDN formation) and through enzymatic hydrolysis by yeast glycosidases during fermentation. Environmental modulation of terpene biosynthesis is substantial — berry temperature (optimal synthesis at 20-25°C, inhibited above 30°C), UV-B radiation (stimulates terpenoid pathway enzyme expression), and water stress (moderate deficit enhances terpene accumulation through concentration and biosynthetic stimulation) all create site-specific and vintage-specific terpene profiles. The analytical distinction between free and bound fractions requires separate extraction and analysis steps — free volatiles are typically isolated by SPME or liquid-liquid extraction, while bound precursors require enzymatic (AR2000 glycosidase cocktail) or acid hydrolysis (pH 1.0, 100°C) prior to volatile analysis.
Monoterpenprofil
Terpenoidanalyse
Terpenolprofil
Terpenprofil
monoterpene profile
terpene composition
terpene data
terpenoid analysis
terpenol profile
Mateo JJ, Jiménez M (2000) Monoterpenes in grape juice and wines. Journal of Chromatography A 881(1-2): 557-567 Martin DM et al. (2010) Functional annotation, genome organization and phylogeny of the grapevine (Vitis vinifera) terpene synthase gene family based on genome assembly, FLcDNA cloning, and enzyme assays. BMC Plant Biology 10: 226 Günata YZ et al. (1985) The role of enzymes in the use of the flavour potential from grape glycosides in winemaking. In Progress in Flavour Precursor Studies, Allured Publishing, pp.: 219-234
Terpene Profile
Terroir expression evaluation comparing Burgundy Premier Cru from Volnay (red fruit, floral, elegant) versus Pommard (darker fruit, earthy, structured) demonstrates intra-regional terroir differentiation where experienced tasters correctly identify commune origin in 85% of blind trials based on aromatic signature alone, with the consistent aromatic differences attributed to soil composition (more clay in Pommard), exposition, and microclimate despite identical variety (Pinot Noir) and similar winemaking, providing sensory evidence for the site-based quality hierarchy that determines vineyard pricing in this appellation system.
Terroir expression quality is an aroma typicity quality that represents the degree to which a wine's aromatic profile communicates its geographical origin through site-specific or region-characteristic aromatic markers, characterizing the sense of place that distinguishes terroir-expressive wines from generic examples without discernible geographical identity. Within the BFO framework, this quality specifically addresses conformity to regional or site-specific aromatic expectations, distinguishing terroir typicity from varietal typicity while recognizing that terroir expression represents the environmental and cultural contribution to aromatic profile that varies among growing sites even within identical variety and winemaking approach. The ontological significance lies in terroir expression providing the aromatic evidence for origin authenticity and site quality hierarchy, where terroir-expressive wines justify geographical indication claims and site-based pricing premiums by communicating origin through sensory experience, making terroir expression the aromatic foundation of fine wine's place-based value system. The compositional basis for terroir expression remains scientifically debated, with proposed mechanisms including soil mineral influence (largely discredited for direct mineral transport), climate effects on grape composition (well-established for sugar, acid, and certain aromatics), microbial terroir (site-specific yeast and bacteria affecting fermentation-derived aromatics), and vineyard ecosystem effects (surrounding vegetation influencing grape volatiles), though the exact pathways from site to aromatic perception remain incompletely understood. The perceptual basis for terroir recognition involves learned associations between aromatic profiles and origin categories, where experienced tasters develop mental prototypes for prestigious origins (Burgundy forest floor, Mosel slate, Chablis flint) that they match against actual perceptions, with terroir assessment requiring familiarity with regional aromatic signatures that novice tasters lack. The cultural construction of terroir expression is significant, as terroir typicity standards reflect accumulated traditions and regulatory definitions that may perpetuate historical practices rather than representing inherent site-aroma relationships, creating the philosophical question of whether terroir is discovered or invented. Understanding terroir expression quality enables origin authenticity assessment, site quality evaluation based on expression clarity, and appreciation of the complex relationship between physical place and aromatic perception in wine.
Terroir_Expression_Quality specifically addresses conformity to geographical origin aromatic expectations, representing the environmental and cultural contribution to wine character. This quality engages with contested scientific questions about terroir mechanisms while acknowledging terroir's functional importance in fine wine evaluation and commerce. The class supports origin authentication and site quality assessment while recognizing the constructed nature of terroir category prototypes.
Goût de Terroir
Herkunftscharakter
Standortexpression
Terroir-Ausdruck
goût de terroir
origin expression
sense of place
site-specific character
Van Leeuwen C, Seguin G (2006) The concept of terroir in viticulture. Journal of Wine Research 17(1): 1-10 Ballester J et al. (2005) Typicality and quality of wines related to their origin. Journal of Wine Research 16(1): 53-60
Terroir Expression Quality
The terroir information for Romanée-Conti (Vosne-Romanée, Burgundy, 47.16°N, 4.95°E) documents the integrated environmental complex — a 1.81 ha east-southeast facing parcel at 255-270 m elevation on the mid-slope of the Côte de Nuits limestone escarpment, with Bathonian oolitic limestone bedrock overlain by 40-80 cm of clay-limestone rendzina soil (35-45% clay, 25-35% calcium carbonate, pH 8.0-8.2), a temperate oceanic climate modified by the continental influence of the Saône plain (mean growing season temperature 15.5-16.5°C, 450-550 mm growing season rainfall, 1,200-1,400 GDD base 10°C), and the specific viticultural practices (Pinot Noir on low-vigor rootstock, high planting density of 10,000 vines/ha, severe yield restriction to 25-30 hL/ha) that together produce the wine recognized as expressing the quintessential Burgundian concept of terroir — the proposition that the specific combination of natural site factors creates a unique and reproducible sensory identity in the wine that transcends vintage variation and winemaking technique.
A grapevine information entity that documents the environmental and geographic context of grape production — the integrated complex of climate, soil, topography, and human cultural practices that together define the terroir of a specific vineyard site — recording the measurable physical parameters and qualitative landscape characteristics that influence vine growth, fruit composition, and wine identity, providing the environmental framework within which cultivar performance and wine quality potential are expressed.
Terroir_Information is the root class for the environmental characterization of vineyard sites — a domain that bridges the natural sciences (geology, pedology, climatology, ecology) with the cultural and economic dimensions of viticulture (appellation systems, land valuation, marketing narratives). The concept of terroir — from the French terre, land or soil — has evolved from its original narrow meaning of soil type to encompass the complete environmental complex including climate, topography, hydrology, biodiversity, and even the human cultural practices that modify the natural environment: the modern scientific understanding of terroir recognizes that wine sensory identity is influenced by measurable environmental parameters (soil water-holding capacity affecting vine water status, mesoclimate temperature profiles affecting phenolic maturation, elevation and aspect affecting radiation balance) that can be mapped, quantified, and related to wine composition through multivariate statistical analysis. The VVO documents terroir information at multiple spatial scales — from the macro-scale of wine regions (defined by climate classification systems), through the meso-scale of appellations and vineyard sites (defined by topographic and soil mapping), to the micro-scale of within-vineyard variation (documented by precision viticulture sensor technologies) — recognizing that terroir effects operate simultaneously at all scales and that the finest-resolution spatial information provides the most complete understanding of site-wine relationships.
Anbauumgebungs-Charakterisierung
Standort-Umwelt-Dokumentation
Terroir-Information
growing environment record
site environment documentation
vineyard terroir characterization
van Leeuwen C and Seguin G (2006) The concept of terroir in viticulture, Journal of Wine Research 17: 1-10 Wilson JE (1998) Terroir — The Role of Geology, Climate, and Culture in the Making of French Wines, University of California Press Seguin G (1986) Terroirs and pedology of wine growing, Experientia 42: 861-873 OIV (2010) Resolution OIV/VITI 333/2010 — Definition of vitivinicultural terroir
Terroir Information
Tertiary aroma description for aged Barolo DOCG (2013 vintage, 11 years post-harvest) documents a complex evolved bouquet dominated by dried rose petals and potpourri (primary terpene transformation products), tar and licorice (Nebbiolo-specific aging signature from phenolic degradation), truffle and forest floor (developed from sulfur compound evolution and phenolic polymerization), tobacco leaf and leather (oxidative maturation products from 30 months in large Slavonian oak botti), and a haunting dried cherry-herb complexity integrating all three aromatic layers — the signature tertiary profile that makes well-aged Nebbiolo among the most aromatically complex wines produced. Tertiary aroma description for aged Riesling Spätlese from the Mosel (2010 vintage, 14 years) records intense petrol/kerosene (TDN at 35 µg/L, well above the 20 µg/L sensory threshold), beeswax and lanolin (from long-chain fatty acid ester evolution), honey and dried apricot (Maillard-type reactions and norisoprenoid development), with residual lime blossom and mineral freshness from preserved terpene backbone — the classic aged Mosel Riesling aromatic profile.
Tertiary aroma description is an aroma description entity that documents the olfactory characteristics of wine attributable to chemical transformations occurring during maturation in barrel, cask, or bottle, encompassing oak-derived aromas (vanilla, toast, spice, coconut, smoke), oxidative maturation characters (nutty, honey, caramel, rancio), reductive bottle-aging characters (developed fruit — dried fruit, compote, confiture earth and mineral — truffle, mushroom, forest floor, wet stone evolved floral — dried rose, potpourri chemical — petrol/TDN, kerosene, camphor), and the integration of primary and secondary aromas into a unified mature bouquet, representing the aromatic dimension that develops over months to decades of aging and constitutes the defining quality of mature, age-worthy wines.
Tertiary aroma description captures the most complex and quality-discriminating aromatic dimension of wine, as the development of tertiary character requires both intrinsic wine quality (sufficient concentration of precursor compounds to generate complex transformation products) and appropriate aging conditions (temperature-stable storage, controlled oxygen exposure, adequate time). The chemical reactions generating tertiary aromas operate through multiple pathways simultaneously — acid-catalyzed hydrolysis of glycosidic terpene precursors releasing new volatile compounds over years of bottle aging, norisoprenoid rearrangements forming TDN (the "petrol" character of aged Riesling) and damascenone (honey-rose complexity), ester hydrolysis-reformation equilibria shifting the ester profile from fermentation-dominant to aged-dominant species, Maillard-type reactions between amino acids and sugars generating melanoidins and heterocyclic aromatics, and acetaldehyde-mediated tannin-anthocyanin condensation producing new aromatic compounds. The oak-derived component of tertiary aroma arises from the extraction of wood constituents during barrel and cask aging — vanillin (sweet, vanilla), eugenol (clove, spice), guaiacol and 4-methylguaiacol (smoky, charred), oak lactones (coconut, woody), and ellagitannins (astringent, structure-building) — with the specific aromatic contribution determined by oak species (French Quercus petraea versus American Q. alba), grain tightness, seasoning duration, and toast level during cooperage. The reductive-oxidative axis during aging fundamentally shapes tertiary aromatic development — wines aged under reductive conditions (bottle aging with minimal oxygen ingress) develop ethereal, mineral, petrolic complexity through slow anaerobic reactions, while wines aged under oxidative conditions (barrel, cask, solera) develop nutty, caramel, rancio complexity through aldehyde accumulation and Strecker degradation. The concept of "bouquet" — the integrated tertiary aromatic impression that transcends the sum of individual descriptors — represents the emergent quality of great aged wines, where primary varietal, secondary fermentative, and tertiary maturation aromas merge into a seamless, multidimensional olfactory experience.
Flaschenbukett
Lagerbukett
Reifearoma
Tertiäraroma
aging bouquet
bottle bouquet
entwickeltes Aroma
evolved aroma description
maturation aroma
tertiary bouquet
Ribereau-Gayon P et al. (2006) Handbook of Enology Volume 2, 2nd Ed, Wiley, pp.: 203-260 Sefton MA et al. (2011) Occurrence, sensory impact, formation, and fate of damascenone in grapes, wines, and other foods and beverages. Journal of Agricultural and Food Chemistry 59(18): 9717-9746 Simpson RF (1978) Aroma and compositional changes in wine with oxidation, storage and ageing. Vitis 17: 274-287 Chatonnet P et al. (1999) Influence of the origin and seasoning of oak. American Journal of Enology and Viticulture 50(4): 447-462
Tertiary Aroma Description
A tertiary bud at node 7 on Vitis vinifera cv. Pinot Noir, examined following combined spring frost and subsequent hail damage that destroyed 94% of primary buds and 78% of secondary buds, dissection revealing extremely small primordia (0.8mm diameter) with only 4-5 leaf initials barely distinguishable and no detectable inflorescence structures, this tertiary bud breaking 18 days after the damage events, producing a weak vegetative shoot reaching only 45cm length with 8 small leaves by mid-summer, representing minimal canopy recovery but enabling vine survival and gradual restoration of photosynthetic capacity over subsequent seasons.
The smallest and least developed bud within the compound bud structure of Vitis vinifera, occupying the third hierarchical position alongside primary and secondary buds within the shared protective bud scale envelope, characterized by minimal size (typically 0.5-1.5mm diameter), rudimentary primordial differentiation, predominantly or exclusively vegetative developmental potential, and activation only under extreme conditions when both primary and secondary buds are destroyed or severely damaged. Tertiary buds represent the ultimate backup meristem in the compound architecture, providing last-resort capacity for shoot production following catastrophic damage events, though typically with purely vegetative growth, very weak vigor, delayed phenology (10-21 days behind primaries), and essentially zero reproductive contribution, making tertiary activation an indicator of severe vineyard damage requiring multi-year recovery strategies.
Tertiary buds are often absent or extremely reduced at many node positions, particularly at basal and apical extremes, with most consistent development at mid-cane positions (nodes 5-9) where compound bud formation is most complete. Not all cultivars develop distinguishable tertiary buds, and even when present, viability and developmental capacity are highly uncertain. Tertiary activation represents vineyard disaster scenarios requiring fundamental reassessment of production expectations, often triggering crop insurance claims and multi-year recovery planning. The purely vegetative nature of tertiary shoots means current-season yield is essentially zero following tertiary activation, with recovery focused on canopy rebuilding and reserve accumulation for future production rather than immediate crop salvage.
Notfallknospe
Tertiärknospe
dritte Knospe
emergency reserve bud
minimal bud
third bud
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Morrison (1991) "Bud development in Vitis= Anatomical study of formation and growth of the primary grapevine bud during one annual cycle" Australian Journal of Grape and Wine Research 72=296-301.
Tertiary Bud
Assessment of Tertiary_Bud_Hardiness_Disposition following the polar vortex event of January 2014 in eastern North America provided field validation of laboratory hardiness measurements across vineyards experiencing temperatures from -25°C to -35°C. Post-winter evaluation of 12 cultivars across 8 sites documented the hierarchy of bud component survival and subsequent shoot development. At sites experiencing -30°C minimum temperature, V. vinifera cultivars ('Riesling', 'Cabernet Franc') suffered 100% primary and secondary bud mortality, with only tertiary buds surviving (15-40% survival) and producing exclusively vegetative growth—no crop was harvested. Cold-hardy hybrids ('Frontenac', 'Marquette') at the same sites retained substantial secondary bud survival (45-70%) enabling 40-55% crop recovery. At the most severely affected site (-35°C minimum), even 'Frontenac' lost all primary and secondary buds, but Tertiary_Bud_Hardiness_Disposition enabled 60% tertiary bud survival and vine persistence. Vines relying on tertiary growth required 2-3 seasons to fully recover canopy architecture and return to normal production. These observations confirm that Tertiary_Bud_Hardiness_Disposition provides vineyard survival insurance against extreme cold events but cannot substitute for primary and secondary hardiness for crop production, defining its role as a last-resort persistence mechanism rather than a production trait.
A Tertiary_Bud_Hardiness_Disposition is a Bud_Cold_Hardiness_Disposition that inheres specifically in the tertiary bud primordium within the compound grapevine bud and is realized through the supercooling capacity and freeze avoidance mechanisms of this smallest, least differentiated bud component that serves as a last-resort growth point when both primary and secondary buds are killed. This disposition represents the cold tolerance of the most rudimentary bud structure, which produces tertiary shoots that are typically vegetative or minimally fruitful (0-0.5 clusters per shoot in most cultivars), providing vine survival and canopy recovery but essentially no commercial crop following severe winter damage. The tertiary bud develops in the axil of the second leaf primordium of the primary bud and remains the smallest and least differentiated of the three bud components, characteristics that confer maximum cold hardiness. Tertiary buds typically achieve LT50 values 4-7°C colder than primary buds, representing the extreme low-temperature survival capacity of the dormant bud complex. While tertiary bud survival rarely contributes meaningfully to crop production, it enables vine survival and canopy regeneration following catastrophic winter events, maintaining vineyard infrastructure for recovery in subsequent seasons. Tertiary_Bud_Hardiness_Disposition thus represents vine survival capacity rather than crop production potential, with its primary value being vineyard persistence following extreme cold events that exceed the hardiness of both primary and secondary buds.
Tertiary_Bud_Hardiness_Disposition is_a Bud_Cold_Hardiness_Disposition. Tertiary buds are the most cold hardy but least fruitful bud component, serving as a survival mechanism for vine persistence following catastrophic winter damage. Tertiary shoots are typically vegetative or minimally fruitful, providing no commercial crop but enabling canopy regeneration and vineyard persistence. The tertiary-primary hardiness differential (typically 4-7°C) defines the margin of vine survival capacity beyond crop production thresholds. Tertiary bud hardiness is most relevant for assessing vineyard survival risk in extreme continental climates where temperatures may exceed both primary and secondary bud hardiness thresholds.
Drittknospen-Frosthärte
Reserveknospen-Kältetoleranz
Tertiärknospen-Härte-Disposition
last-resort bud LT50
tertiary bud freeze tolerance
third bud hardiness
Sabbatini P, Howell GS (2013) Freezing temperature responses of primary, secondary, and tertiary grape buds during natural cold acclimation, Acta Horticulturae 978:307-313 Dami IE et al. (2012) Evaluation of winter injury and subsequent performance of grapevine cultivars in Ohio, Acta Horticulturae 932:235-241 Zabadal TJ et al. (2007) Winter Injury to Grapevines and Methods of Protection, Michigan State University Extension Bulletin E-2930
Tertiary Bud Hardiness Disposition
In Grenache vineyards of Southern Rhône at BBCH 57-59, secondary inflorescence branches on basal and lower primary branches bear 2-4 tertiary branches each averaging 0.8-1.8 cm length and supporting 4-10 flowers per tertiary, collectively contributing 150-250 flowers (15-20% of total) to the large, loose clusters of 800-1200 berries. Mourvèdre exhibits similarly extensive tertiary branching (2-3 per secondary, 1-2 cm length), creating the pronounced compound, multi-lobed cluster architecture. Conversely, compact-clustered Gewürztraminer completely lacks tertiary branches, with all flowers borne directly on the rachis, primary branches, and abbreviated secondary branches, resulting in very tight clusters of 80-150 berries highly susceptible to Botrytis bunch rot.
A tertiary inflorescence branch is an inflorescence branch that arises as a lateral appendage from a secondary inflorescence branch, representing third-order architectural elaboration that creates maximum structural complexity in loose-clustered cultivars while being reduced or absent in compact-clustered types, thereby serving as a principal determinant of cultivar-level cluster architecture variation. Tertiary branches emerge from secondary branches at intervals of 1-3 mm during late stages of inflorescence primordium development within the dormant bud, with branch number per secondary branch varying from 0-4 depending on cultivar architecture type (extensively branched loose-clustered cultivars may have 2-4 tertiaries per secondary moderately branched types have 0-2 compact types often have none). These branches exhibit the smallest dimensions among flower-bearing branches, typically measuring 0.2-2 cm in length and 0.2-0.6 mm in diameter at the base, with pronounced variation correlating to the position and vigor of the parent secondary branch and its parent primary branch. Tertiary branch anatomy features a minimal vascular strand continuous with the secondary branch vasculature, providing resource supply for the 2-15 flowers typically borne per tertiary branch, predominantly through direct attachment rather than through quaternary subdivision (which is rare and limited to extreme loose-clustered types like some Muscat varieties). Branch distribution on secondary branches follows the same basipetal developmental gradient observed at primary and secondary levels, with proximal tertiary branches generally longer than distal ones on the same secondary. Tertiary branch presence, number, and elaboration represent critical diagnostic features for ampelographic cluster classification, with extensive tertiary branching defining the 'very loose' or 'winged' cluster categories (OIV code 204 categories 1-3) while absence of tertiary branches characterizes 'compact' or 'very compact' categories (OIV codes 7-9). Each tertiary branch typically bears 3-12 flowers, with all tertiary branches collectively contributing 10-25% of total inflorescence flower number in extensively branched cultivars but 0% in compact types lacking tertiary branches entirely. Tertiary branch development becomes visible during BBCH 55-57 as small protuberances on elongating secondary branches, with distinct separation occurring during BBCH 57-59, making tertiary branches among the latest-developing inflorescence structures. The degree of tertiary branching shows substantial genotypic determination but also exhibits phenotypic plasticity in response to favorable growing conditions during the previous season's inflorescence primordium differentiation, particularly abundant water and nutrients during June-July bud development.
Tertiary_Inflorescence_Branch represents the finest level of architectural elaboration in most Vitis vinifera cultivars and must be distinguished from Secondary_Inflorescence_Branch (which arises from primary branches) and from direct flower attachment (flowers borne without subtending branches). Quaternary branches (fourth-order) are extremely rare and limited to a few loose-clustered cultivars under optimal conditions, so tertiary branches typically represent the terminal branching level. The presence, abundance, and development of tertiary branches serve as key ampelographic descriptors and correlate strongly with disease susceptibility patterns, harvest logistics, and wine style suitability. Sub-classification into Terminal and Lateral positions reflects functional and positional distinctions within the tertiary branching system.
Seitenast zweiter Ordnung
Tertiärast der Infloreszenz
lateral branch of secondary branch
tertiärer Blütenstandsast
third-order inflorescence branch
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 92-94 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 172-178 OIV (2009). Descriptor List for Grape Varieties and Vitis Species, OIV code 204 (cluster density).
Tertiary Inflorescence Branch
A tertiary lateral vein on a mature leaf at node 6 of Vitis vinifera cv. Chardonnay in Burgundy, France, branching from a secondary lateral vein at 1.2 cm from the secondary vein's junction with its primary vein, branching angle of 82 degrees creating nearly perpendicular orientation, extending 0.8 cm through blade tissue, diameter of 0.22 mm barely visible to unaided eye, appearing as very faint raised line on abaxial surface under magnification, invisible on adaxial surface without magnification, light green coloration matching surrounding tissue, no visible pubescence at this scale, containing single xylem vessel (12-18 μm diameter) or occasionally two small vessels providing water to local blade area of approximately 0.3-0.5 cm², minimal phloem consisting of few companion cells and sieve elements, negligible mechanical tissues, serving as final organized vein level from which numerous veinlets branch creating the ultimate reticulate network, veinlets arising at irregular intervals and angles creating dense mesh filling all spaces between adjacent tertiary veins, creating total venation density (including all orders) of approximately 12-15 mm vein length per mm² blade area, measured at BBCH stage 75 (berries pea-sized) in late June, representing typical third-order venation creating the fine-scale distribution network transitioning to veinlet reticulum.
A lateral vein representing third-order branches originating from secondary lateral veins in the hierarchical venation system in Vitis vinifera, characterized by branching origin from secondary lateral veins, small diameter typically 0.15-0.35 mm representing substantial reduction from secondary vein size, third hierarchical position in lateral vein system serving as final organizing level before transition to veinlets, limited vascular capacity with 1-2 small xylem vessels supporting water delivery to local blade areas, minimal mechanical contribution, typical number of 5-10 tertiary veins branching from each secondary vein creating fine distribution network, branching angles from secondary veins typically 60-90 degrees creating approximately orthogonal branching patterns, extension through blade tissue for 0.3-1.5 cm creating small-scale vascular territories, and functional roles including final distribution of water from secondary veins to veinlet networks, local photosynthate collection from immediately adjacent mesophyll, minimal structural support, and organizing veinlet initiation completing the reticulate venation pattern reaching all intercostal blade regions between major veins. Tertiary lateral veins create the transition level between clearly hierarchical vein orders (primary, secondary) and the dense reticulate veinlet networks, developing late during leaf expansion following establishment of primary and secondary networks, exhibiting high density creating fine-scale vascular distribution with typical spacing of 0.5-1.5 mm between adjacent tertiary veins, and contributing to total venation density that determines ultimate hydraulic efficiency and maximum photosynthetic capacity supportable.
Tertiary lateral veins are distinguished from secondary veins by branching origin from secondaries and smaller diameter, and from veinlets by maintenance of hierarchical branching pattern and visible structure under low magnification. The tertiary level represents the finest clearly hierarchical vein order, with subsequent branching creating the reticulate veinlet network that lacks strict hierarchical organization. Visibility requires magnification for detailed observation, with tertiary veins barely discernible to unaided eye appearing as faint lines on abaxial surfaces. Development occurs during later expansion phases following primary and secondary vein establishment, with tertiary vein differentiation proceeding outward from secondary veins filling blade regions. Vascular organization shows minimal specialization, with single xylem vessels and limited phloem representing near-minimum functional vascular capacity. Mechanical contribution is negligible compared to primary and secondary veins, with tertiary veins providing essentially no structural support. Functional importance lies in creating fine-scale vascular distribution reducing diffusion distances from mesophyll cells to nearest vein entry points, affecting hydraulic efficiency and maximum photosynthetic rates. High tertiary vein density creates shorter mesophyll-to-vein distances enabling higher transpiration and photosynthesis, but requires greater resource investment in vascular tissues potentially reducing photosynthetic tissue proportion. The transition from tertiary veins to veinlets is somewhat arbitrary, based on diameter thresholds and branching pattern regularity rather than clear anatomical distinctions.
dritte Ordnung Lateralvene
feine Zweigvene
fine lateral vein
minor branch vein
tertiäre Seitenader
third-order lateral vein
Based on Sack & Holbrook (2006) "Leaf hydraulics" Annual Review of Plant Biology 57=361-381 Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Brodribb et al. (2007) "Leaf hydraulic capacity in ferns, conifers and angiosperms= impacts on photosynthetic maxima" New Phytologist 165=839-846 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 173-194.
Tertiary Lateral Vein
A tertiary reproductive bud at node 7 on Vitis vinifera cv. Riesling under exceptionally favorable growing conditions, dissection revealing tiny primordia (0.9mm diameter) containing rudimentary vegetative structures plus what appears to be extremely small inflorescence initial with perhaps 15-18 barely distinguishable flower primordia, representing potential cluster of 6-9 berries if activated and fruit set achieved, though such tertiary reproductive buds documented in fewer than 0.1% of examined Riesling tertiary positions and considered anomalous rather than typical, with practical vineyard significance essentially zero given the catastrophic damage scenarios required for tertiary activation make minimal reproductive potential irrelevant to recovery strategies.
A tertiary bud containing any detectable inflorescence primordial structures in addition to vegetative shoot primordia, representing an extraordinarily rare condition in Vitis vinifera compound buds, characterized by presence of rudimentary flower initials (typically fewer than 20-30 flowers even when present) alongside minimal vegetative structures, documented in a few cultivars under exceptional growing conditions but considered essentially non-existent in practical viticulture. Tertiary reproductive buds, when they occur at all, reflect unusual resource allocation during bud development or genetic predisposition in specific cultivars toward distributed reproductive capacity, but their extreme rarity, minimal cluster potential, and uncertain activation success make them negligible factors in yield calculations and vineyard management decisions, with tertiary positions universally treated as vegetative-only in production planning and damage assessment protocols.
Tertiary reproductive buds are so rare in viticulture literature and practice that some researchers question whether true reproductive tertiary buds exist at all versus artifacts of developmental abnormalities or misidentified secondary buds. The few documented cases come primarily from Riesling, Gewürztraminer, and a few other cultivars noted for generally high fruitfulness across all bud positions. Even when present, the minimal flower number (typically under 30 versus 100-300 in primaries) and questionable developmental capacity make reproductive contribution negligible. The evolutionary logic argues against tertiary reproductive investment - if conditions are severe enough to destroy primaries and secondaries, reproductive effort is futile and resources better allocated to vegetative survival. Tertiary reproductive buds remain a taxonomic curiosity rather than a practically significant category in grapevine biology.
cluster-bearing tertiary bud
fruchtende dritte Knospe
fruiting tertiary bud
reproductive third bud
reproduktive Tertiärknospe
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Srinivasan & Mullins (1981) "Physiology of flowering in the grapevine - a review" American Journal of Enology and Viticulture 32=47-63.
Tertiary Reproductive Bud
Chardonnay grafted on SO4 rootstock at 14 years showing tertiary root system, soil core sampling at 45 cm depth revealing tertiary roots comprising third through sixth branching orders, representative third-order tertiary root 1.2 cm diameter 8 cm length branching from second-order secondary root, fourth-order tertiary roots 0.4 cm diameter 4 cm length, fifth and sixth-order fine tertiary roots <2 mm diameter 1-3 cm length with abundant root hairs, tertiary root length density 18 cm/cm³ soil providing extensive absorption surface, mycorrhizal colonization 52% of tertiary root length, tertiary roots providing estimated 75% of total vine water and nutrient uptake
Tertiary root is root of third or higher branching order arising from secondary roots, characterized by small diameter typically 0.1-2 cm though finest tertiary roots <2 mm diameter, limited or absent secondary growth with most tertiary roots showing primary anatomical structure, predominantly absorptive function providing majority of vine water and nutrient uptake, relatively short lifespan typically 1-3 years or less for finest orders showing high turnover, and representing highest-order roots in hierarchical root architecture comprising numerically dominant component though individually small accounting for 60-80% of total root system length and surface area, with tertiary roots typically being third-order branches from second-order secondary roots, fourth-order from third-order, and potentially fifth through eighth or higher orders in extensively branched systems, showing concentration in upper soil layers 20-80 cm depth where resources most available, producing abundant root hairs increasing absorption surface area 5-20 fold, hosting arbuscular mycorrhizal fungi enhancing phosphorus and water uptake, and demonstrating rapid proliferation and senescence responding to soil resource heterogeneity creating dynamic ephemeral root component contrasting with persistent woody primary and secondary root framework. Tertiary root hierarchical position includes= representing third or higher branching order (tertiary roots by definition being third generation or higher in branching hierarchy from primary root), arising from secondary root parents (tertiary roots branching from second-order secondary roots or from other tertiary roots of lower order), serving as terminal branches (highest-order tertiary roots representing terminal branches in root hierarchy with no further branching), showing high numerical abundance (tertiary roots vastly outnumbering primary and secondary roots due to exponential branching multiplication), and demonstrating fine-scale organization (tertiary roots creating finest spatial scale in root system hierarchy exploring soil at millimeter to centimeter resolution). Tertiary root diameter and size includes= showing small to very small diameter (tertiary roots typically 0.1-2 cm with finest orders <2 mm reflecting high branching order), exhibiting limited length (individual tertiary roots typically 1-20 cm length exploring local soil micro-volumes), demonstrating minimal biomass individually (single tertiary root containing very small tissue mass though collectively significant due to vast numbers), displaying delicate structure (tertiary roots fine and fragile compared to robust woody primary and secondary roots), and revealing size-order relationship (tertiary root diameter declining progressively with increasing order from larger third-order to finest eighth-order roots). Tertiary root anatomy includes= showing primary structure (most tertiary roots lacking secondary growth maintaining primary anatomical organization), exhibiting epidermis or exodermis (outer protective layer rather than periderm bark of woody roots), demonstrating thin cortex (cortical parenchyma tissue comprising significant proportion of root volume), containing endodermis (casparian strip regulating radial solute movement), showing small stele (central vascular cylinder with limited xylem and phloem reflecting small root size), and displaying root hairs (epidermal extensions abundant on youngest tertiary root portions). Tertiary root absorptive function includes= providing majority of uptake (tertiary roots contributing 60-90% of total vine water and nutrient absorption despite being finest roots), showing high surface area (vast numbers and lengths of tertiary roots creating enormous soil contact area), producing root hairs (abundant root hairs on tertiary roots increasing effective absorption surface 5-20 fold), hosting mycorrhizae (arbuscular mycorrhizal colonization of tertiary roots enhancing uptake particularly phosphorus), and demonstrating direct absorption (tertiary root epidermis and root hairs directly absorbing water and minerals from soil solution). Tertiary root distribution includes= concentrating in upper soil (tertiary roots most abundant 20-80 cm depth in zone of highest resource availability), showing patchy distribution (tertiary root density highly variable spatially proliferating in resource-rich patches), occupying pore spaces (fine tertiary roots growing through soil pore network exploring micro-environments), demonstrating high density (potentially 5-50+ cm tertiary root length per cubic centimeter soil in favorable zones), and revealing responsive proliferation (tertiary root production increasing in favorable zones decreasing in poor zones). Tertiary root turnover dynamics includes= showing short lifespan (tertiary roots typically functioning 3 months to 3 years depending on order and conditions with finest orders shortest-lived), exhibiting continuous production (vine continuously producing new tertiary roots throughout growing season), demonstrating seasonal mortality (tertiary root death concentrated in late summer-fall and winter though occurring year-round), revealing rapid turnover (fine tertiary roots potentially replacing themselves multiple times per year), and contributing to carbon cycling (tertiary root death and decomposition providing substantial organic matter input to soil). Tertiary root mycorrhizal associations includes= hosting arbuscular mycorrhizae (tertiary roots colonized by AM fungi forming arbuscules and vesicles in root cortex), showing colonization rates (typically 30-70% of tertiary root length colonized depending on conditions and cultivar), demonstrating mutual benefit (vine providing carbohydrates to fungi receiving enhanced phosphorus and water uptake in return), extending effective exploration (mycorrhizal hyphae extending beyond root surface accessing resources unavailable to roots alone), and revealing dependency (grapevine strongly mycorrhiza-dependent with reduced growth and nutrient status without fungal partners).
Tertiary_Root emphasizes third or higher branching order and is distinguished from Primary_Root and Secondary_Root by focus on fine high-order absorptive function. Includes Lateral_Tertiary_Root and Fine_Tertiary_Root. Critical for understanding fine root architecture, absorption dominance, root hair production, mycorrhizal associations, high turnover, and the numerically abundant fine high-order roots providing majority of vine resource uptake.
Fine root
High-order root
Terminal root branch
Tertiärwurzel
Wurzel höherer Ordnung
Tertiary Root
A tertiary vegetative bud at node 6 on Vitis vinifera cv. Chardonnay, forced to break following severe combined frost-hail-herbicide damage that eliminated both primary and secondary buds across 89% of vine nodes, dissection revealing minimal vegetative primordia with only 4 distinguishable leaf initials measuring 0.2-0.4mm each and no trace of inflorescence structures, developing into an extremely weak shoot reaching 38cm with 7 small, often malformed leaves totaling only 185 cm² leaf area, providing minimal photosynthetic contribution but maintaining node position viability and producing slightly better-developed buds for following season that enabled gradual three-year recovery progression.
A tertiary bud containing exclusively vegetative primordia without any differentiated inflorescence structures, representing the standard condition for essentially all tertiary positions across cultivars and node locations, characterized by rudimentary shoot primordia with minimal leaf initials (typically 3-6), complete absence of reproductive structures, and development into weak vegetative shoots when forced to break following destruction of primary and secondary positions. Tertiary vegetative buds reflect the evolutionary priority allocation in compound bud development, where resources during bud formation are concentrated in primary positions (high reproductive potential) with progressively less investment in secondary (moderate reproductive potential) and tertiary (survival minimum only) positions, creating a developmental hierarchy that balances productivity under normal conditions with survival capacity under catastrophic damage scenarios.
Tertiary vegetative buds are the default and essentially universal condition for tertiary positions, with tertiary reproductive buds being extraordinarily rare or non-existent in most cultivars. The vegetative-only nature reflects both resource limitation during bud formation (tertiary buds develop last and smallest) and evolutionary logic (if primary and secondary are both destroyed, reproductive effort is futile survival through minimal vegetative maintenance becomes the priority). Management following tertiary activation focuses on vine survival rather than production, with decisions centered on whether to maintain struggling vines through recovery or remove and replant, depending on vine age, variety value, and economic considerations of multi-year production loss.
minimal vegetative bud
nicht-fruchtende dritte Knospe
non-fruiting tertiary bud
vegetative Tertiärknospe
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Morrison (1991) "Bud development in Vitis= Anatomical study of formation and growth of the primary grapevine bud during one annual cycle" Australian Journal of Grape and Wine Research 72=296-301 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press, pp. 85-92.
Tertiary Vegetative Bud
Syrah in Rhône Valley shows testa lignification during August-October Year N+1, with outer integument testa cells depositing secondary walls 2.6-4.4 micrometers thickness weeks 8-16, phenylpropanoid pathway producing monolignol precursors with PAL, C4H, 4CL, CAD enzyme activities increasing 5-15 fold, lignin polymerization through peroxidase-catalyzed radical coupling creating guaiacyl-syringyl lignin polymer, lignin content accumulating to 24-36% testa dry weight, brown pigmentation developing from tan to dark brown, testa cells dying leaving lignified walls, mature testa providing hard impermeable protective layer except at hilum, under warm Mediterranean conditions of 24-29°C temperature, typical of this mid-to-late season cultivar completing testa lignification
Testa lignification represents the outer coat layer hardening phase during seed coat formation when outer integument-derived testa cells deposit lignin in secondary cell walls creating rigid impermeable protective layer, phenolic monolignols polymerize into lignin matrix, brown pigmentation develops from phenolic oxidation, progressive lignification increases wall rigidity and impermeability, and lignified testa provides mechanical protection and pathogen resistance, occurring weeks 8-16 post-fertilization as major protective structure formation process. This lignification and hardening phase occurs during late summer to early autumn in August-October in temperate regions Year N+1, representing major seed coat protective structure development. Lignification mechanism involves secondary cell wall deposition in testa cells, monolignol precursor synthesis through phenylpropanoid pathway, monolignol transport to cell wall, lignin polymerization creating complex phenolic polymer, lignin cross-linking with wall polysaccharides, testa cell death leaving lignified walls as protective structure, and pigmentation from phenolic oxidation creating brown color. Testa cell characteristics shows outer integument cells differentiated into testa layer, cells elongated radially 40-80 micrometers length, cells depositing thick secondary walls, and testa as outer protective layer. Secondary cell wall deposition shows substantial wall material deposited inside primary wall, secondary walls 2-5 micrometers thickness in mature testa, wall deposition increasing cell wall mass 5-10 fold, walls composed of cellulose microfibrils embedded in lignin matrix, and secondary walls providing mechanical strength. Wall deposition timing shows secondary wall synthesis initiating weeks 6-8, intensive deposition weeks 8-14, completion by week 14-16, and timing coordinating with lignification. Cellulose in secondary walls shows cellulose microfibrils as structural framework, microfibrils oriented in organized patterns, cellulose providing tensile strength, and cellulose as major wall component. Lignin biosynthesis shows phenylpropanoid pathway producing monolignol precursors, pathway enzymes including phenylalanine ammonia lyase PAL, cinnamate 4-hydroxylase C4H, 4-coumarate CoA ligase 4CL, cinnamyl alcohol dehydrogenase CAD producing monolignols, and biosynthetic pathway as lignin precursor source. Monolignol types shows p-coumaryl alcohol, coniferyl alcohol, sinapyl alcohol as three major monolignols, monolignols differing in methoxylation degree, and monolignol composition affecting lignin properties. Monolignol transport shows monolignols synthesized in cytoplasm, monolignols transported across plasma membrane to apoplast, transport to cell wall for polymerization, and transport as biosynthesis step. Lignin polymerization shows monolignols polymerizing through radical coupling reactions in cell wall, peroxidases and laccases catalyzing oxidation and coupling, polymerization creating complex three-dimensional lignin polymer, progressive polymerization building lignin matrix, and polymerization as lignification mechanism. Lignin structure shows irregular heteropolymer with various linkage types, guaiacyl and syringyl lignin units in grapevine testa, lignin molecular weight very high molecular weight polymer, and structure providing rigidity. Lignin deposition pattern shows lignin deposited throughout secondary wall, lignin embedding cellulose microfibrils, lignin filling wall spaces creating dense matrix, and lignin distribution creating rigid structure. Lignin-polysaccharide cross-linking shows lignin covalently bonded to hemicelluloses and possibly pectins, cross-links increasing structural integrity, cross-linking creating integrated wall structure, and bonding enhancing rigidity. Lignification progression shows progressive lignin accumulation weeks 8-16, lignin content increasing from near-zero to 20-40% testa dry weight, increasing lignification correlating with hardening, and progressive accumulation creating protection. Lignification spatial pattern shows lignification initiating in outer testa cells, lignification progressing inward through testa layers, eventual lignification throughout testa, and centripetal pattern characteristic. Testa cell death shows testa cells eventually dying during or after lignification, cell death part of normal maturation, dead cells leaving lignified walls as protective structure, and programmed cell death as developmental process. Cell death timing shows cell death during late lignification weeks 14-16, eventual complete cell death by maturation, dead cells in mature seed coat, and death completing testa maturation. Pigmentation development shows brown pigments accumulating during lignification, pigments from phenolic compound oxidation, enzymatic oxidation through peroxidases and polyphenol oxidases, non-enzymatic oxidation possible, and pigmentation creating characteristic brown seed coat. Pigment characteristics shows brown to dark brown color, pigments including oxidized flavonoids, tannins, other phenolics, pigmentation intensity increasing with maturation, and color as maturation indicator. Color progression shows testa color changing from pale tan early to brown mid-maturation to dark brown mature, color darkening correlating with lignification degree, final color varying from medium to very dark brown depending on cultivar, and color as visual maturity marker. Testa impermeability shows lignified testa creating largely impermeable barrier, impermeability to water preventing uncontrolled imbibition, impermeability to gases regulating respiration, impermeability to pathogens preventing entry, and impermeability as protective function. Permeability control shows impermeability except at hilum and micropyle regions, hilum as preferential water entry site during germination, micropyle possibly enabling gas exchange, and controlled permeability enabling eventual germination. Mechanical protection shows lignified testa providing hard rigid structure, testa resisting mechanical damage from compression, impact, abrasion, protection essential for seed survival during dispersal and storage, and mechanical strength as primary protective function. Testa rigidity shows lignin-cellulose composite creating rigid structure, rigidity increasing with lignification degree, mature testa requiring substantial force to fracture or penetrate, and rigidity as protection mechanism. Chemical protection shows lignin resistant to microbial degradation, phenolic compounds possibly having antimicrobial properties, chemical protection preventing pathogen attack, and chemical resistance as protection. Environmental effects on testa lignification include temperature affecting lignification rate with optimal 22-26°C supporting normal progression, nutrient availability particularly phenolic precursors affecting lignin biosynthesis, light possibly affecting phenolic metabolism, and conditions affecting lignification quality and completion. Temperature effects shows warm conditions promoting faster lignification and pigmentation, cool conditions potentially slowing lignification, and temperature as rate factor. Nutrient effects shows adequate nitrogen and minerals supporting phenolic biosynthesis, nutrient status affecting lignin quantity and quality, and nutrition as lignification factor. The process represents second phase of seed coat formation creating protective structure, following integument development and preceding seed coat hardening. Duration approximately 8 weeks representing major lignification weeks 8-16. Regional timing shows testa lignification during August-October as protective layer hardens. Viticultural significance includes representing primary protective structure formation, creating mechanical and pathogen protection enabling seed survival, regulating seed permeability affecting germination, and establishing seed coat physical characteristics determining seed quality. Research applications include analyzing lignification regulation and biosynthesis, investigating lignin structure and composition in seed coats, evaluating protective functions and permeability properties, quantifying lignification dynamics and completion, and assessing testa lignification as seed protection and quality determinant.
Testa_Lignification is second phase of Seed_Coat_Formation, distinguished by lignin deposition. Represents weeks 8-16. Secondary walls 2-5 μm in testa cells. Phenylpropanoid pathway (PAL, C4H, 4CL, CAD) produces monolignols. Lignin polymerization creating 20-40% DW. Brown pigmentation from phenolic oxidation. Testa cell death. Creates impermeability (except hilum/micropyle), mechanical protection, rigidity. Follows Integument_Development. Precedes Seed_Coat_Hardening. Major protective structure formation. Temperature 22-26°C optimal.
Schutzschicht-Lignifizierung
Testa-Lignifizierung
Testa-Verhärtung
outer coat lignification
phenolic polymer deposition
protective layer lignification
testa hardening
äußere Schalen-Lignifizierung
Testa Lignification
The tetraploid status of the Chardonnay Gros Grains variant was confirmed by flow cytometry showing a G1 peak at approximately twice the fluorescence intensity of the diploid Chardonnay reference (2C DNA content approximately 1.90 pg versus 0.95 pg for the diploid), with chromosome counting from root tip preparations confirming 2n = 76 — this autotetraploid arose spontaneously during long-term vegetative propagation and was selected for its significantly larger berries (2.2 g versus 1.4 g average berry weight in the diploid), thicker leaves with larger stomatal guard cells (guard cell length 32-38 μm versus 22-26 μm in diploid), and reduced seed set (averaging 0.8 seeds per berry versus 1.6 in the diploid) that produces the rounder, more compact clusters characteristic of the Gros Grains selection. The tetraploid Chardonnay shares an identical SSR profile with the diploid at all 9 OIV loci but shows four alleles at some loci where the diploid shows two, reflecting the doubled allele dosage.
A ploidy information record documenting the tetraploid chromosome complement (2n = 4x = 76) in a specific Vitis vinifera accession, recording the doubled genome state arising from spontaneous or induced whole-genome duplication that produces cells containing four complete sets of 19 chromosomes, with associated phenotypic effects including increased cell size, larger berries and organs, altered leaf morphology, reduced fertility, and modified secondary metabolite profiles compared to the diploid progenitor.
Tetraploid_Status documents the most common ploidy deviation in cultivated grapevine — autotetraploidy arising from somatic chromosome doubling in meristematic cells during vegetative propagation. Tetraploid sectors or whole-plant tetraploids arise when a mitotic error (endomitosis or endoreduplication) produces a cell with the doubled chromosome complement that subsequently proliferates to colonize the shoot apical meristem: if the entire L1, L2, and L3 meristem layers become tetraploid, the resulting plant is a uniform autotetraploid if only some layers are affected, a cytochimeral plant results (documented under Chimera_Status). The phenotypic consequences of tetraploidy in grapevine are generally consistent with the gigas effect observed in other polyploid plants — larger cells producing larger organs (berries, leaves, stomata), increased vigor, but reduced fertility due to multivalent chromosome pairing at meiosis producing unbalanced gametes. In breeding programs, colchicine-induced tetraploids have been produced deliberately to enable interploidy crosses (4x × 2x yielding triploid seedless table grapes), representing one of the most commercially successful applications of ploidy manipulation in Vitis.
4x ploidy record
Ganzverdopplungs-Nachweis
Tetraploidie-Status
autotetraploid status
tetraploid confirmation
tetraploider Chromosomensatz
Ollat N et al. (2011) Grape berry development, in Gerós H et al. (eds) The Biochemistry of the Grape Berry, Bentham Science pp: 1-29 Notsuka K et al. (2000) Induction of autotetraploid plants by in vitro culture of grapevine, Journal of the Japanese Society for Horticultural Science 69: 543-545 Pelsy F (2010) Molecular and cellular mechanisms of diversity within grapevine varieties, Heredity 104: 331-340
Tetraploid Status
Texture perception quality comparison between entry-level and Grand Cru Burgundy Pinot Noir reveals the quality differentiation at textural level, with entry-level showing adequate but unremarkable texture (moderately smooth, slight graininess from less refined tannin) while Grand Cru displays exceptional textural refinement (seamlessly silky, no detectable grain, velvet-like plushness) that distinguishes the premier expression independent of flavor intensity, demonstrating how texture perception contributes to quality hierarchy recognition.
Texture perception quality is a tactile quality that represents the fine-grained surface character of wine as detected through mechanoreceptors during oral contact, characterizing the microscale tactile impression ranging from grainy and particulate through silky and smooth to velvety and plush that contributes to overall mouthfeel refinement and quality perception. Within the BFO framework, this quality addresses the surface texture dimension of tactile perception, distinguishing fine-scale textural character from broader mouthfeel qualities (body, viscosity) while recognizing that texture provides the qualitative nuance that differentiates wines of similar weight and flow properties. The ontological significance lies in texture perception serving as a refinement indicator that distinguishes premium from ordinary wines, where fine, polished texture suggests quality fruit, appropriate extraction, and careful élevage while coarse or grainy texture may indicate aggressive processing, poor tannin quality, or inadequate maturation affecting overall quality impression. The perceptual basis for texture involves high-resolution mechanoreceptor detection of surface character as wine contacts oral mucosa, with different receptor types (Meissner corpuscles for light touch, Merkel cells for pressure) contributing to the integrated texture impression that distinguishes silky from grainy or velvety from rough character. The compositional determinants of texture include tannin structure and concentration (polymerization state affecting perceived fineness), polysaccharide content (mannoproteins and arabinogalactans smoothing texture), colloidal particles (potentially creating grainy perception), glycerol level (contributing to smooth texture), and the overall integration of components where balanced composition produces harmonious texture while imbalanced wines show textural discontinuity. The metaphorical vocabulary for texture draws on fabric and material analogies (silky, velvety, satiny, grainy, chalky) that communicate tactile character through shared sensory reference, enabling wine professionals to convey subtle textural differences through these evocative descriptors. Understanding texture perception quality enables quality assessment based on textural refinement, production method evaluation from textural character, and appreciation of how microscale tactile properties contribute to overall wine elegance or coarseness.
Texture_Perception_Quality serves as the parent class for specific textural qualities (Graininess, Silkiness, Velvetiness), addressing the fine-scale surface character dimension of tactile assessment. This quality provides quality differentiation through textural refinement, with premium wines typically showing finer, more polished texture than everyday wines. The class employs metaphorical vocabulary (fabric analogies) that enables communication of subtle tactile differences.
Texturcharakter
Texturwahrnehmung
mouth texture
surface texture
tactile fineness
taktile Feinheit
textural character
Texture Perception Quality
Texture quality assessment in Thompson Seedless table grapes employs both sensory panels evaluating crispness, firmness, and skin toughness on 1-9 hedonic scales, and instrumental texture profile analysis measuring skin break force (target 1.5-2.5 N), flesh firmness (target 2.0-3.5 N), and skin elasticity using penetrometer attachments optimal texture combining crisp flesh with tender skin commands premium prices in export markets, while soft or tough-skinned fruit is downgraded regardless of sugar content, demonstrating that texture quality rivals sweetness in determining consumer acceptance and market value.
Texture_Quality encompasses all morphological qualities that characterize the tactile and structural properties of grapevine organ surfaces and internal tissues as perceived through touch or instrumental measurement. Texture describes how an organ feels when contacted, reflecting the physical arrangement of cells, the properties of cell walls, the presence of surface structures such as hairs or wax deposits, and the mechanical behavior of tissues under deformation. This category is fundamental to grape quality assessment because texture properties directly influence sensory perception during consumption, processing behavior during winemaking, and postharvest storage potential. Texture_Quality subdivides into two major domains: Surface_Texture_Quality addressing the external boundary characteristics including smoothness, roughness, pubescence, and waxy bloom and Internal_Texture_Quality capturing the mechanical and structural properties of tissues beneath the surface including pulp consistency, flesh firmness, and cellular cohesion. Surface texture affects visual appearance through light scattering, influences water relations through effects on boundary layer conductance, and determines susceptibility to pathogen attachment and penetration. Internal texture governs eating quality in table grapes, juice extraction efficiency in winemaking, and resistance to mechanical damage during harvest and transport. Texture assessment employs tactile evaluation by trained assessors, visual observation of surface features, microscopic examination of cellular structure, and instrumental methods including texture analyzers that measure force-deformation relationships, surface profilometry, and image analysis of microscopic features. Texture properties change during berry development and ripening as cell walls are modified, pectin is solubilized, and turgor pressure decreases, making texture assessment stage-dependent.
Texture_Quality is positioned as a direct subclass of Morphological_Quality, parallel to Size_Quality, Shape_Quality, Color_Quality, and Structural_Integrity_Quality. Texture involves both surface and internal properties, distinguished in the child classes Surface_Texture_Quality and Internal_Texture_Quality. The relationship between texture qualities and the physical structures producing them (cell walls, surface appendages) connects this quality category to the Morphological_Structure component of the VVO. PATO_0000150 (texture) provides the upper-ontology alignment. Texture assessment requires standardization of measurement conditions including temperature, sample preparation, and instrument settings for reproducibility.
Oberflächenbeschaffenheit
Texturqualität
structural feel
surface character
tactile quality
taktile Eigenschaft
Rolle L et al. (2012) Instrumental Texture Analysis Parameters as Markers of Table-Grape and Winegrape Quality: A Review. American Journal of Enology and Viticulture 63(1): 11-28 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 159-189
Texture Quality
Thermal buffering capacity comparison between VSP-trained and sprawl-trained Merlot in Hawke's Bay shows VSP canopies reducing maximum berry temperature by 4.2°C during heat events (from 44°C to 39.8°C through improved air circulation) while sprawl canopies reduce minimum berry temperature by 2.1°C during frost events (from -2.5°C to -0.4°C through long-wave radiation trapping), illustrating the inverse relationship between heat and frost buffering that requires site-specific optimization based on the dominant thermal stress risk.
Thermal buffering capacity quality is a physical quality that inheres in grapevine tissues and the surrounding microenvironment and represents the integrated capacity to moderate temperature extremes through the combined effects of thermal mass, evaporative cooling, radiative exchange modification, and convective heat transfer that together determine the degree to which tissue temperatures are damped relative to ambient air temperature fluctuations. Within the BFO framework, this quality addresses the functional outcome of multiple thermal processes rather than any single physical property, characterizing the realized moderation of temperature extremes that protects tissues from damage at thermal limits. The ontological significance lies in thermal buffering integrating passive mechanisms (thermal mass, insulation) with active processes (transpirational cooling, metabolic heat generation) and environmental modifications (canopy shading, wind shelter) that collectively determine tissue exposure to extreme temperatures. Grapevine thermal buffering capacity varies by tissue type, developmental stage, and environmental context, with actively transpiring leaves showing strong buffering through evaporative cooling during heat stress but limited buffering during frost when stomata close, while dormant buds rely on passive thermal mass and insulating bud scales for frost protection. The canopy architecture contributes substantially to thermal buffering, where dense foliage shades interior clusters from radiant heating while also trapping long-wave radiation reducing frost exposure, creating complex trade-offs between heat and cold stress mitigation. Viticultural practices modify thermal buffering through canopy management affecting shade and air circulation, cover crops affecting soil heat storage and release, and water application providing evaporative cooling and modifying soil thermal properties. Understanding thermal buffering capacity enables site assessment for climate suitability, variety selection for thermal stress tolerance, and management optimization for extreme temperature mitigation.
Thermal_Buffering_Capacity_Quality represents the integrated functional outcome of multiple thermal processes, distinguishing from Thermal_Mass_Quality which addresses only the energy storage component. This quality supports site and management system evaluation for climate resilience. The class encompasses both passive physical buffering and active physiological responses, requiring context specification as the dominant buffering mechanisms differ between heat stress and frost stress scenarios.
Temperaturausgleichskapazität
Wärmepufferkapazität
microclimate buffering
temperature moderation capacity
thermal damping capacity
thermal stress mitigation capacity
thermische Dämpfung
thermische Pufferung
Gladstones J (2011) Wine, Terroir and Climate Change. Wakefield Press, Adelaide, pp. 89-124 Smart RE (1985) Principles of grapevine canopy microclimate manipulation with implications for yield and quality. American Journal of Enology and Viticulture 36(3): 230-239
Thermal Buffering Capacity Quality
Thermal mass assessment of Riesling clusters in Rheingau reveals heat capacity of 3.89 J/g·K at harvest maturity (85% water content) declining to 3.42 J/g·K in late-harvest fruit (72% water content due to dehydration), with the 150g average cluster mass providing sufficient thermal inertia to delay morning temperature rise by 45 minutes compared to air temperature during autumn radiation frost events, creating a critical protection window exploited by harvest scheduling to avoid frozen fruit.
Thermal mass quality is a physical quality that inheres in grapevine tissues and structures and represents the capacity to store thermal energy as a function of mass and specific heat capacity, determining the rate at which tissue temperature changes in response to environmental thermal fluctuations. Within the BFO framework, this quality characterizes the thermal inertia of plant material where tissues with high thermal mass resist rapid temperature change, buffering short-term environmental extremes, while tissues with low thermal mass respond quickly to ambient conditions. The ontological significance lies in thermal mass governing the temporal dynamics of tissue temperature response, where the product of mass and specific heat capacity determines the energy required to change tissue temperature by a given increment, with larger thermal mass requiring more energy input or removal to achieve the same temperature change. Grapevine organs vary substantially in thermal mass, with massive woody tissues (trunk, cordons, canes) exhibiting high thermal mass due to large volume and high water content, while thin leaves show low thermal mass responding rapidly to radiation changes, and berries occupy an intermediate position with thermal mass varying through development as size and water content change. The specific heat capacity of plant tissues approximates that of water (4.18 J/g·K) for highly hydrated tissues but decreases as dry matter proportion increases, with lignified tissues showing lower specific heat than succulent fruit or actively transpiring leaves. Thermal mass effects are particularly significant during frost events where high thermal mass tissues cool more slowly, potentially remaining above damaging temperatures through brief cold episodes, and during heat waves where thermal mass delays peak tissue temperature but also prolongs heat exposure after air temperature declines. Understanding thermal mass enables prediction of tissue temperature dynamics from environmental data, design of frost protection strategies exploiting thermal mass effects, and interpretation of the differential temperature responses among vine organs.
Thermal_Mass_Quality addresses the energy storage aspect of thermal behavior, complementing Temperature_Quality which characterizes instantaneous thermal state. This quality is calculated from mass and specific heat capacity rather than directly measured, requiring compositional information for accurate estimation. The class supports frost damage prediction and thermal modeling of vine systems where organs of different thermal mass show distinct temperature dynamics under the same environmental forcing.
Wärmekapazität
Wärmespeicherkapazität
heat capacity
heat storage quality
thermal inertia
thermal storage capacity
thermische Masse
thermische Trägheit
Nobel PS (2020) Physicochemical and Environmental Plant Physiology, 5th Ed. Academic Press, San Diego, pp. 307-341 Cellier P et al. (1993) Modelling freezing stress in grapevine buds. Agricultural and Forest Meteorology 67(1-2): 119-134
Thermal Mass Quality
Thermal tolerance quality comparison across Mediterranean versus Central European Vitis vinifera cultivars evaluated the physiological basis of heat adaptation relevant to cultivar selection for warming wine regions. Garnacha (Grenache), originating from the hot Spanish interior, demonstrated superior thermal tolerance quality with photosynthesis maintaining 80% of optimal rates at leaf temperatures of 40°C, membrane integrity preserved with only 20% electrolyte leakage following 45°C exposure for 30 minutes, and rapid heat shock protein induction within 2 hours of stress onset providing molecular chaperoning that protected enzymes at elevated temperatures. Riesling, originating from cooler German regions, showed inferior thermal tolerance quality with photosynthesis declining to 50% of optimal at 40°C leaf temperature, membrane damage indicated by 45% electrolyte leakage under equivalent heat exposure, and slower heat shock response requiring 6 hours for comparable protein induction. The thermal tolerance quality differential translated to field performance differences, with Garnacha showing minimal damage during a 42°C heat event while Riesling in adjacent blocks exhibited 30% leaf scorch and 15% sunburn, establishing thermal tolerance quality as a primary determinant of cultivar suitability for warm climate viticulture.
Thermal tolerance quality is a heat stress indicator quality that inheres in a grapevine cultivar or clone and represents the capacity to maintain cellular function and avoid irreversible damage under elevated temperature conditions, determined by the inherent stability of cellular structures and metabolic processes at high temperatures combined with the inducible protective mechanisms that are activated in response to heat exposure, collectively establishing the temperature thresholds beyond which damage occurs. This quality complements heat damage severity quality by addressing the underlying capacity that determines damage thresholds rather than the damage itself, providing the predictive framework for anticipating which cultivars and conditions will experience damage during heat events of given intensity and duration. The ontological significance of thermal tolerance quality lies in its role as the mechanistic foundation for understanding and predicting heat damage susceptibility, enabling cultivar selection decisions for warm climate viticulture, identification of management practices that enhance tolerance, and anticipation of adaptation needs under projected climate warming that will increase the frequency and intensity of heat events in many viticultural regions. The biological basis of thermal tolerance encompasses multiple cellular protection mechanisms including the inherent thermostability of membrane lipids and proteins that determines function at elevated temperatures, the heat shock response producing molecular chaperones that prevent protein aggregation and maintain enzyme function, the antioxidant systems that scavenge reactive oxygen species generated under heat stress, and the osmotic adjustment mechanisms that maintain cellular hydration under high evaporative demand. The variation in thermal tolerance quality among grapevine cultivars reflects genetic differences in these protective mechanisms, with cultivars originating from warm climate regions generally exhibiting superior tolerance compared to those from cool climate origins, though substantial variation exists within both groups providing opportunities for selection and breeding to improve heat tolerance. The acclimation capacity that determines how tolerance changes with prior heat exposure represents an important component of thermal tolerance quality, with gradual temperature increases permitting acquisition of tolerance that would not be present following abrupt exposure to the same temperatures.
Thermal_Tolerance_Quality is_a Heat_Stress_Indicator_Quality, serving as parent class for Membrane_Thermostability_Quality, Photosynthetic_Heat_Tolerance_Quality, and Enzyme_Thermostability_Quality which address specific tolerance mechanisms. This class addresses the underlying capacity for heat resistance while Heat_Damage_Severity_Quality addresses actual damage expression. Cultivar origin correlates with tolerance level, providing selection guidance. Acclimation capacity enables tolerance enhancement through gradual exposure. Critical for climate change adaptation through cultivar selection and breeding for improved thermotolerance.
Hitzetoleranz
Hochtemperaturtoleranz
Thermotoleranz
Wärmeresistenz
heat resistance
heat tolerance
thermotolerance
Wahid A et al. (2007) Heat tolerance in plants: an overview. Environmental and Experimental Botany 61:199-223 Xu S et al. (2014) Effects of heat acclimation pretreatment on changes of membrane lipid peroxidation, antioxidant metabolites, and ultrastructure of chloroplasts in two cool-season turfgrass species under heat stress. Environmental and Experimental Botany 56:274-285 Greer DH, Weston C (2010) Heat stress affects flowering, berry growth, sugar accumulation and photosynthesis of Vitis vinifera cv. Semillon grapevines grown in a controlled environment. Functional Plant Biology 37:206-214
Thermal Tolerance Quality
Cabernet Sauvignon thick berry skin at maturity measures 110-145 μm total thickness (cuticle 5-6.5 μm, epidermis 26-32 μm, hypodermis 3 layers 45-58 μm, outer mesocarp 5-7 layers 42-58 μm), providing skin-to-pulp ratio 12-16% with skin representing substantial proportion of berry mass, containing total phenolic index 65-85 (high for red cultivars), requiring 14-21 day maceration at 26-30°C with twice-daily punchdowns to achieve full extraction producing wines with 4.5-6 g/L tannins, 14-18 OD units color intensity, full-bodied structure suitable for 10-25 year aging potential. Syrah thick berry skin showing extreme thickness= 120-170 μm total thickness (among thickest in common cultivars), skin-to-pulp ratio 13-18%, containing 4-5.5 mg/g anthocyanins and 11-15 mg/g tannins in skin tissue, requiring extended maceration (18-30 days typical for premium Syrah) to fully extract phenolics producing deeply colored (16-22 OD units) full-bodied wines with 5-7 g/L tannins and characteristic pepper-spice aromatics from skin-derived rotundone and other volatiles. Tannat thick berry skin representing extreme thick-skinned cultivar= 130-185 μm thickness (thickest among major cultivars), skin-to-pulp ratio 14-19%, containing 12-18 mg/g tannins (extremely high skin tannin content), producing wines requiring 21-35 day maceration to achieve tannin extraction and balance, with final wines showing 6-9 g/L tannins (among highest in red wines) creating powerful astringent structure requiring 5-10 years aging to soften through tannin polymerization and precipitation. Cabernet Sauvignon water deficit effects on skin thickness= well-watered vines producing berries 15.5 mm diameter with 118 μm skin, skin-to-pulp ratio 12.5% deficit-irrigated vines producing 13.5 mm berries with 125 μm skin, ratio 14.8%, demonstrating water deficit enhancement of thick skin character by limiting flesh expansion more than skin development creating proportionally more skin per berry and higher phenolic concentration in resulting wines (25-35% higher tannin, 18-28% higher anthocyanins in deficit wines). Nebbiolo thick berry skin developmental timing= late-ripening cultivar maintaining thick skin (108-158 μm) throughout extended ripening period (18-24 weeks fruit set to harvest versus 14-18 weeks for early cultivars), with prolonged on-vine time allowing full phenolic maturity development in thick skins essential for producing age-worthy Barolo and Barbaresco wines showing characteristic tannin structure and aromatic complexity from extended skin phenolic development.
A thick berry skin is a berry skin showing substantial thickness typically >95-110 μm at maturity, characteristic of certain cultivars (Cabernet Sauvignon, Syrah, Nebbiolo, Tannat) and environmentally-influenced conditions, providing high skin-to-pulp ratio (typically >10-12%), elevated phenolic extraction potential, firm tactile properties, and suitability for full-bodied tannic wine production requiring extended maceration to achieve optimal phenolic extraction and balance. Thick berry skin structural characteristics= (1) total thickness typically 95-180 μm with cultivar variation (Cabernet Sauvignon 110-145 μm, Syrah 115-165 μm, Tannat 125-180 μm, Nebbiolo 105-155 μm) (2) increased layer numbers particularly in hypodermis (2-4 cell layers versus 1-2 in thin skins) and outer mesocarp (4-8 layers versus 2-4 in thin skins) (3) thicker cuticle (4.5-8 μm versus 2.5-5 μm in thin skins) providing enhanced barrier properties (4) possibly thicker cell walls within skin layers contributing to firmness (5) skin-to-pulp ratio (skin mass as percentage of berry mass) typically 10-16% versus 5-9% in thin-skinned cultivars, creating higher concentration of skin-derived compounds per unit berry or juice volume. Thick berry skin compositional implications= (1) higher total phenolic content per berry due to larger skin tissue volume (though concentration per unit skin tissue may be similar to thin skins) (2) higher skin tannin-to-seed tannin ratio in resulting wines as more skin tissue relative to flesh provides more total skin tannin extraction (3) higher anthocyanin total content per berry (in red cultivars) though concentration per unit skin may be comparable to thin skins (4) potential for higher aromatic compound extraction from larger skin volume particularly in aromatic cultivars. Thick berry skin winemaking implications= (1) extended maceration requirements (10-30 days typical for premium production) to achieve full phenolic extraction from substantial skin tissue (2) higher extraction temperature and more vigorous cap management potentially needed to penetrate thick skins and extract tannins (3) ability to produce full-bodied wines with substantial tannin structure (3-6 g/L total tannins) and deep color intensity (12-20 OD units at 520 nm for reds) (4) wines showing extended aging potential due to high tannin and anthocyanin content providing structural backbone and antioxidant capacity (5) risk of over-extraction if maceration excessive creating harsh astringent character requiring careful extraction management. Thick berry skin development influenced by genetics and environment= genetic component with certain cultivars consistently producing thick skins across environments (Cabernet Sauvignon, Syrah, Nebbiolo consistently thick-skinned) environmental enhancement with water deficit during berry development potentially increasing skin-to-pulp ratio by limiting flesh expansion more than skin development sunlight exposure potentially increasing skin thickness through UV-induced cell proliferation and wall thickening small berry size correlating with higher skin-to-pulp ratio as surface area-to-volume relationships create proportionally more skin in smaller berries. Thick berry skin tactile and sensory properties= firm skin texture during eating requiring more chewing to break down compared to thin skins astringency perception from high tannin content when berries or fresh juice tasted potential advantages for table grape shipping durability (thick skins resist damage) though typically thick-skinned wine cultivars not preferred for fresh consumption due to skin texture.
Thick_Berry_Skin represents cultivars and conditions producing substantial skin thickness (>95-110 μm) and is distinguished from Thin_Berry_Skin (<70-85 μm) by greater skin development, higher skin-to-pulp ratio, and different winemaking implications. Thick skins are characteristic of cultivars producing full-bodied tannic wines with aging potential. Understanding thick berry skin properties is important for cultivar selection, irrigation management affecting berry size and skin ratio, harvest timing allowing full phenolic maturity, and winemaking extraction protocol design for optimal phenolic balance.
Dickschalige Beere
dicke Beerenhaut
dickschalige Traube
heavy skin
thick-skinned berry
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 180-185 Roby, G., Matthews, M.A. (2004). Relative proportions of seed, skin and flesh in ripe berries. Australian Journal of Grape and Wine Research 10=74-82 Ojeda, H. et al. (2002). Influence of water deficits on grape berry growth. Vitis 41=141-145.
Thick Berry Skin
A thick main petiole on a large mature basal leaf at node 4 of Vitis vinifera cv. Grenache in Châteauneuf-du-Pape, France, measuring 11.5 cm length with 4.8 mm diameter at mid-length representing substantial thickness, cylindrical to slightly flattened cross-sectional shape with well-defined ridges marking major vascular bundles, green coloration with moderate red-purple tinge particularly on sun-exposed side, moderate to dense pubescence with substantial trichome coverage, containing eight major vascular bundles with large xylem vessels visible in cross-section supporting high transpiration rates measured at 3.2 mmol H₂O m⁻² s⁻¹ under favorable conditions, thick-walled collenchyma and sclerenchyma in outer tissues providing mechanical strength resisting breakage, demonstrating high rigidity with limited flexibility but adequate wind response through elastic bending rather than rotation, supporting 5.8 gram leaf mass (18.5 cm × 17.2 cm lamina) at stable 55-degree angle from horizontal, showing characteristic open U-shaped petiolar sinus, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing thick robust petiole construction typical of vigorous basal leaves on well-resourced vines.
A main petiole exhibiting diameter in the upper range for the species and cultivar in Vitis vinifera, characterized by mid-length diameter typically exceeding 4-5 mm representing robust structural construction, substantial cross-sectional area providing enhanced mechanical strength and vascular capacity, well-developed vascular bundles including multiple xylem vessel elements enabling high water transport capacity and substantial phloem supporting efficient photosynthate export, extensive mechanical tissues including thick-walled collenchyma and sclerenchyma providing structural rigidity, common occurrence in vigorous vines with abundant resource availability, sun-exposed positions, and large lamina requiring substantial support, particularly on basal and lower-median leaves showing maximum dimensions, functional implications including high mechanical strength resisting breakage under heavy leaf mass and wind loading, enhanced vascular capacity supporting maximum transpiration and photosynthesis rates without hydraulic limitation, reduced flexibility creating stiffer leaf positioning with limited angular adjustment, and correlation with overall leaf vigor and function. Thick main petioles develop through extensive radial growth during petiole development, influenced by genetic factors affecting vascular and mechanical tissue development, environmental conditions including high light and abundant resources supporting robust construction, hormonal regulation affecting cambial activity and tissue differentiation, and leaf position with basal leaves receiving developmental priority producing thicker petioles than apical positions.
Thick main petioles are distinguished from thin to medium petioles by diameter typically exceeding 4.0-4.5 mm, with exact thresholds varying somewhat by cultivar baseline dimensions and measurement position. Thickness variation correlates strongly with leaf position, with basal leaves typically showing thickest petioles (4-5 mm), median leaves showing intermediate thickness (3-4 mm), and apical leaves showing thinnest petioles (2-3 mm) reflecting both developmental timing and resource allocation patterns. Vine vigor strongly affects petiole thickness, with vigorous vines producing substantially thicker petioles compared to stressed or low-vigor vines even within the same cultivar. Functional implications include mechanical advantages supporting larger heavier leaves without structural failure, vascular advantages enabling maximum transpiration and photosynthesis through reduced hydraulic resistance, but potentially reduced flexibility limiting leaf angle adjustment responses. Thickness measurement requires standardization to specific positions (typically nodes 5-8) on mature leaves for meaningful comparisons. Correlation with overall leaf and vine vigor makes thick petioles indicators of excellent resource status but not necessarily optimal balance, as excessive vigor may indicate vegetative-reproductive imbalance requiring management intervention.
dicker Hauptblattstiel
heavy petiole
massiver Blattstängel
robust petiole
robuster Petiolus
thick leaf stalk
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 136-165 Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Chaves et al. (2010) "Grapevine under deficit irrigation= hints from physiological and molecular data" Annals of Botany 105=661-676 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 137-172.
Thick Main Petiole
A thick main rachis on a large compound primary inflorescence of Vitis vinifera cv. Muscat of Alexandria in Cyprus, measuring 24.5 cm length, exhibiting exceptional diameter of 5.5 mm at base representing very robust construction, diameter remaining substantial at 3.2 mm even at the apex indicating thick construction throughout, cross-sectional anatomy showing well-developed collenchyma and sclerenchyma mechanical tissues occupying 35% of rachis diameter, containing 16 major vascular bundles arranged in ring with individual bundles 0.8-1.2 mm diameter, thick rachis supporting large elaborate inflorescence bearing approximately 850 flowers distributed across 22 primary branches, mechanical strength enabling support of inflorescence fresh weight of 28 grams at full bloom, vascular capacity supporting maximum transpiration estimated at 40 ml water per day during peak bloom, thick robust construction enabling subsequent support of very large cluster potentially reaching 1500+ grams at harvest representing 50+ fold mass increase from flowering, rachis thickness correlating with overall inflorescence vigor and size reflecting favorable development conditions and high fertility, measured at BBCH stage 65 (full bloom) in late May, representing maximum thick main rachis construction characteristic of very large elaborate inflorescences requiring substantial structural and vascular capacity.
A main rachis exhibiting diameter in the upper range representing robust construction in Vitis vinifera, characterized by diameter typically exceeding 4-5 mm at base representing substantial thickness, robust construction with well-developed mechanical tissues, high vascular capacity with numerous large bundles often 12-18+ bundles, occurrence in large vigorous inflorescences on well-resourced vines, correlation with large overall inflorescence size and high flower number, enhanced mechanical strength supporting heavy inflorescence and subsequent cluster masses, increased vascular capacity enabling maximum water and nutrient delivery, and functional implications including ability to support very large clusters without structural failure and vascular capacity sufficient for maximum berry development without hydraulic limitation. Thick main rachis represents maximum robustness in main rachis construction, occurring in highly fertile cultivars producing large inflorescences and in favorable development conditions supporting robust rachis elaboration, exhibiting structural and vascular characteristics enabling support of very large fruit loads, and serving as indicator of high potential yield and cluster size.
Thick main rachis is distinguished from typical main rachis by diameter exceeding 4-5 mm representing the robust extreme, with exact thresholds somewhat arbitrary but diameters >5 mm representing very thick construction. The thick designation emphasizes the dimensional robustness and structural/vascular consequences. Diameter superiority reflects enhanced development with greater cell numbers, larger cell sizes, and more extensive tissue elaboration compared to typical rachis. Mechanical tissue development shows enhancement with thick rachis containing more extensive collenchyma and sclerenchyma providing superior strength and stiffness. Vascular elaboration includes increased bundle number (12-18+ versus typical 8-12), larger individual bundles, and greater total conducting area enabling maximum transport capacity. Occurrence correlates with large inflorescences showing high flower numbers (>600-800) and elaborate branching, reflecting the structural and vascular demands. Cultivar effects show some varieties consistently producing thick rachis (Muscat of Alexandria, some high-fertility clones) while others show typical dimensions even under favorable conditions. Environmental influences include vigor, resource availability, and development conditions affecting rachis elaboration with high vigor and favorable resources supporting thick construction. Functional capacity enables support of very large clusters potentially 1000-2000+ grams without structural failure, with thick rachis representing the prerequisite for maximum cluster size expression. Vascular capacity sufficiency prevents hydraulic limitation of berry development, enabling berries to achieve maximum size potential. Management implications include recognizing thick rachis as indicator of high yield potential requiring assessment of vine balance and potential need for crop thinning to maintain quality.
dicke Hauptrachis
heavy rachis
massive Hauptspindel
robust main rachis
robuste Zentralachse
thick central axis
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 255-290 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 439-478.
Thick Main Rachis
Comparative analysis of Thick_Skin_Disposition at the University of Adelaide assessed berry mechanical properties across 28 wine grape cultivars using texture analyzer measurements of puncture force and skin thickness determinations via histological sectioning. 'Cabernet Sauvignon' exhibited pronounced Thick_Skin_Disposition with mean skin thickness of 312 μm, puncture force of 0.89 N, and tensile strength of 1.42 MPa. In contrast, thin-skinned 'Pinot Noir' showed values of 178 μm, 0.41 N, and 0.68 MPa respectively. Under controlled inoculation with B. cinerea conidia (10⁵/mL) applied to unwounded berries, 'Cabernet Sauvignon' developed infection in only 8% of inoculated berries versus 34% in 'Pinot Noir' after 7 days incubation at 20°C and 95% RH. When berries were pre-wounded with a sterile needle, infection rates equalized at 85-92% for both cultivars, confirming that Thick_Skin_Disposition operates primarily through mechanical barrier function preventing initial penetration rather than post-infection resistance. Field observations corroborated laboratory findings, with 'Cabernet Sauvignon' showing 3-4 fold lower grey mold incidence than 'Pinot Noir' under equivalent disease pressure in Coonawarra vineyards.
A Thick_Skin_Disposition is a Cluster_Architecture_Based_Resistance_Disposition that inheres in a Vitis vinifera berry and is realized through the development of a mechanically robust exocarp (skin) that provides enhanced physical resistance to Botrytis cinerea penetration and reduces susceptibility to berry cracking, wound formation, and cuticle microfissures that serve as pathogen entry points. This disposition represents the genetically determined capacity to produce berries with thick, tough skins characterized by multiple cell layers in the hypodermis, thick-walled epidermal cells, high content of structural polysaccharides (cellulose, hemicellulose, pectin), and dense phenolic accumulation providing mechanical reinforcement. Berry skin thickness in Vitis vinifera varies substantially among cultivars, ranging from approximately 150 μm in thin-skinned table grapes to over 350 μm in thick-skinned wine grapes, with corresponding differences in puncture resistance and tensile strength. Thick-skinned genotypes resist mechanical damage during berry expansion, rain events, and cluster compression, maintaining skin integrity that prevents opportunistic B. cinerea entry. Additionally, thick skins often correlate with elevated phenolic content (anthocyanins, tannins) that may contribute antimicrobial activity. The genetic control of berry skin thickness involves multiple QTLs and shows high heritability, enabling effective selection. However, thick skin disposition may influence wine style through increased phenolic extraction and potential for skin-derived bitter or astringent compounds, requiring balance with enological objectives.
Thick_Skin_Disposition is_a Cluster_Architecture_Based_Resistance_Disposition. This class represents mechanical barrier resistance through berry skin properties, classified under structural rather than physiological resistance mechanisms. Thick skin correlates with but is distinct from Cuticle_Based_Resistance_Disposition which specifically addresses the outermost waxy layer. Skin thickness influences wine style through phenolic extraction, creating potential trade-offs with enological objectives. The disposition provides protection primarily against unwounded berry infection wounded berries remain susceptible regardless of skin thickness.
Dicke-Beerenhaut-Disposition
Dickschaligkeit-Disposition
Mechanische-Hautresistenz
mechanical skin resistance
thick berry skin trait
tough skin disposition
Deytieux-Belleau C et al. (2009) Berry skin development in Vitis vinifera: morphological and biochemical events linked to Botrytis resistance, Journal of Experimental Botany 60:3919-3930 Gabler FM et al. (2003) Correlations of morphological, anatomical, and chemical features of grape berries with resistance to Botrytis cinerea, Phytopathology 93:1263-1273 Rosenquist JK, Morrison JC (1989) Some factors affecting cuticle and wax accumulation on grape berries, American Journal of Enology and Viticulture 40:241-244
Thick Skin Disposition
Berry skin thickness in Tannat, measured histologically at harvest as 280-350 μm in the Madiran appellation, exceeds that of most other Vitis vinifera cultivars and accounts for the exceptionally high tannin concentrations (3000-5000 mg/L catechin equivalents in young wines) that define this variety's tannic character and extended aging requirement, necessitating specific vinification protocols including extended maceration and micro-oxygenation to achieve tannin polymerization and mouthfeel softening.
Thickness_Quality represents the linear dimension measured as the smallest extent of a grapevine organ, typically perpendicular to the primary surface plane in laminar structures or representing the wall dimension in hollow or layered tissues. Thickness is the critical dimension for characterizing leaf blades, berry skins, cell walls, bark, and tissue layers where the thin dimension determines functional properties disproportionate to its absolute value. Leaf blade thickness reflects mesophyll development and correlates with photosynthetic capacity per unit area, specific leaf area, and drought tolerance. Sun-exposed leaves develop greater thickness (200-350 μm) than shade leaves (150-250 μm) due to enhanced palisade mesophyll development. Berry skin thickness is a key quality determinant because the skin contains most phenolic compounds, aromatic precursors, and contributes texture to wine thick-skinned cultivars like Cabernet Sauvignon (250-350 μm) yield more tannic wines than thin-skinned cultivars like Pinot Noir (150-250 μm). Cell wall thickness influences tissue mechanical properties, pathogen resistance, and processing characteristics. Bark thickness on trunks and cordons affects cold hardiness, desiccation resistance, and susceptibility to trunk diseases. Thickness measurements require specialized techniques depending on the scale: histological sectioning and microscopy for cellular dimensions, micrometers or dial gauges for intact tissues, and texture analyzers for mechanical assessment of skin thickness through puncture tests. Thickness qualities exhibit environmental plasticity, generally increasing under conditions of high light, water stress, or slow growth that favor cell wall deposition relative to cell expansion.
Thickness_Quality captures the dimension that is neither length nor width in three-dimensional organs, and the single dimension perpendicular to the surface plane in laminar structures. For spherical organs, thickness is not applicable for cylindrical organs, thickness may refer to wall thickness (e.g., hollow shoot pith cavity wall thickness). Berry skin thickness measurement protocols must specify whether the measurement includes only the exocarp (true skin) or also the outer hypodermis layers, as literature values vary with protocol. PATO_0000915 (thickness) provides the upper-ontology reference.
Dicke
Gewebedicke
Schichtdicke
Wandstärke
laminar thickness
minimal dimension
perpendicular extent
wall thickness
Considine JA and Knox RB (1979) Development and Histochemistry of the Cells, Cell Walls, and Cuticle of the Dermal System of Fruit of the Grape, Vitis vinifera L. Protoplasma 99(4): 347-365 Hanlin RL et al. (2010) Review: Condensed Tannin and Grape Cell Wall Interactions and Their Impact on Tannin Extractability into Wine. Australian Journal of Grape and Wine Research 16(1): 173-188
Thickness Quality
Pinot Noir thin berry skin at maturity measures 60-95 μm total thickness (cuticle 3-5 μm, epidermis 18-26 μm, hypodermis 1-2 layers 22-38 μm, outer mesocarp 3-4 layers 18-32 μm), providing skin-to-pulp ratio 6-9% with skin representing modest proportion of berry mass, requiring gentle extraction during 5-10 day maceration at 24-28°C with minimal cap management producing elegant wines with 2-3.5 g/L tannins, 8-12 OD units color intensity, emphasizing red fruit aromatics and silky texture characteristic of quality Pinot Noir rather than power or extraction. Gamay thin berry skin showing extreme thin-skinned characteristics= 48-72 μm total thickness (among thinnest in wine cultivars), skin-to-pulp ratio 5-7%, extracting very rapidly during 4-7 day carbonic or semi-carbonic maceration typical for Beaujolais production, yielding light-bodied wines with 1.5-2.5 g/L tannins, 6-9 OD units color, emphasizing fresh fruit character and minimal tannin structure for early consumption. Thompson Seedless table grape thin berry skin for fresh consumption= 52-78 μm thickness, delicate texture requiring minimal chewing, preferred for eating quality though creating vulnerability to damage during harvest and shipping requiring careful handling, with thin skin contributing to crisp texture and fresh flavor without excessive tannin or phenolic perception. Pinot Noir berry size effect on skin ratio= small berries 11 mm diameter showing 68 μm skin thickness, skin-to-pulp ratio 8.5% large berries 14 mm diameter showing 72 μm skin thickness (similar absolute thickness), ratio only 6.2%, demonstrating that even modest absolute skin thickness differences combine with size effects to create substantial skin ratio differences affecting phenolic concentration in wines with small-berry selections producing more concentrated phenolic profiles. Grenache thin berry skin winemaking adaptation= traditional Grenache production using 7-12 day maceration with gentle extraction achieving 2.5-4 g/L tannins and 9-14 OD color, producing medium-bodied wines with characteristic red fruit and spice aromatics, versus modern high-extraction techniques (18-25 day maceration, higher temperatures) risking over-extraction of seed tannins creating harsh astringent character from attempting to extract from limited skin tissue beyond optimal extraction window, demonstrating importance of matching extraction intensity to thin skin characteristics.
A thin berry skin is a berry skin showing reduced thickness typically <70-85 μm at maturity, characteristic of certain cultivars (Pinot Noir, Gamay, Grenache, some table grape cultivars) and some environmental conditions, providing low skin-to-pulp ratio (typically <8-10%), reduced phenolic extraction potential, delicate tactile properties, and suitability for elegant lighter-bodied wine production requiring shorter gentle maceration or for fresh table grape consumption where delicate skin texture is preferred. Thin berry skin structural characteristics= (1) total thickness typically 45-85 μm with cultivar variation (Pinot Noir 60-95 μm, Gamay 50-75 μm, some table cultivars 45-70 μm) (2) reduced layer numbers particularly in hypodermis (1-2 cell layers versus 2-4 in thick skins) and outer mesocarp (2-4 layers versus 4-8 in thick skins) (3) thinner cuticle (2.5-5 μm versus 4.5-8 μm in thick skins) potentially affecting barrier properties (4) thinner cell walls within skin layers contributing to delicate texture (5) skin-to-pulp ratio typically 5-9% versus 10-16% in thick-skinned cultivars, creating lower concentration of skin-derived compounds per unit berry or juice volume. Thin berry skin compositional implications= (1) lower total phenolic content per berry due to smaller skin tissue volume (though concentration per unit skin tissue may be similar to thick skins) (2) lower skin tannin-to-seed tannin ratio in resulting wines as less skin tissue relative to flesh provides less total skin tannin extraction creating wines where seed tannins may contribute proportionally more (3) lower total anthocyanin content per berry (in red cultivars) potentially requiring careful extraction optimization to achieve adequate color (4) potential for more delicate aromatic profile with less extraction of skin-derived compounds potentially allowing more expression of fruit-derived aromatics. Thin berry skin winemaking implications= (1) shorter maceration requirements (5-12 days typical) as limited skin tissue extracts more rapidly and extended maceration risks excessive seed tannin extraction (2) gentler extraction techniques (lower temperatures 22-26°C, gentle cap management) needed to avoid harsh seed tannin extraction (3) wines showing lighter body with moderate tannin structure (1.5-3.5 g/L total tannins) and more delicate color (6-12 OD units at 520 nm for reds) (4) wines emphasizing elegance, aromatics, and fruit expression rather than power and structure (5) shorter aging potential (5-15 years typical versus 10-25+ years for thick-skinned cultivars) due to lower tannin and anthocyanin reserves though quality thin-skinned wines can age gracefully. Thin berry skin development influenced by genetics and environment= genetic component with certain cultivars consistently producing thin skins across environments (Pinot Noir, Gamay consistently thin-skinned) environmental factors with abundant water during berry development potentially decreasing skin-to-pulp ratio through enhanced flesh expansion large berry size correlating with lower skin-to-pulp ratio as surface area-to-volume relationships create proportionally less skin in larger berries shading potentially reducing skin thickness through reduced UV-induced cell proliferation. Thin berry skin tactile and sensory properties= delicate skin texture during eating requiring minimal chewing, preferred characteristic for table grape consumption in many markets reduced astringency perception when berries or fresh juice tasted due to lower tannin content potential vulnerability to splitting, pathogen invasion, and post-harvest damage due to thinner protective barrier requiring careful handling.
Thin_Berry_Skin represents cultivars and conditions producing reduced skin thickness (95-110 μm) by less skin development, lower skin-to-pulp ratio, and different winemaking implications. Thin skins are characteristic of cultivars producing elegant lighter-bodied wines or delicate table grapes. Understanding thin berry skin properties is important for cultivar selection, vineyard practices affecting berry size and skin ratio, harvest timing, and winemaking extraction protocol design requiring gentle techniques appropriate for limited skin phenolic reserves.
Dünnschalige Beere
delicate skin
dünne Beerenhaut
dünnschalige Traube
thin-skinned berry
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 180-185 Roby, G., Matthews, M.A. (2004). Relative proportions of seed, skin and flesh. Australian Journal of Grape and Wine Research 10=74-82.
Thin Berry Skin
Thiol precursor concentration varies substantially across cultivars, with Sauvignon Blanc showing the highest accumulation (Cys-3MH 200-800 µg/L, Cys-4MMP 10-50 µg/L) compared to lower levels in Riesling, Gewürztraminer, and Colombard (50-200 µg/L Cys-3MH). Precursors exist as both cysteine conjugates (Cys-3MH, Cys-4MMP) and glutathione conjugates (Glut-3MH, Glut-4MMP), with the glutathione forms representing biosynthetic intermediates converted to cysteine conjugates in the berry. Yeast carbon-sulfur lyases during fermentation cleave 1-5% of precursors to release free volatile thiols, with wine concentrations of 200-5000 ng/L 3MH and 1-50 ng/L 4MMP in aromatic Sauvignon Blanc. The remarkably low sensory thresholds (60 ng/L for 3MH, 0.8 ng/L for 4MMP) mean these trace compounds dominate wine aroma.
An aromatic compound disposition that is the inherited capacity of a Vitis vinifera plant to synthesize and accumulate non-volatile cysteine and glutathione conjugates of volatile thiols in grape berries, with genotype-specific precursor concentrations determining the potential for release of powerful odorants (3-mercaptohexan-1-ol, 4-mercapto-4-methylpentan-2-one, 3-mercaptohexyl acetate) that define the tropical, citrus, and boxwood character of Sauvignon Blanc and related wines.
Thiol_Precursor_Accumulation_Disposition serves as the parent class for specific thiol precursor traits. Volatile thiols represent a relatively recently discovered aroma class (1990s-2000s) that revolutionized understanding of Sauvignon Blanc character. The compounds are non-volatile and odorless in grape juice, with aroma release requiring yeast enzymatic cleavage during fermentation. This fermentation-dependent release means that both grape precursor concentration and yeast strain selection affect wine thiol content. Oxidation rapidly destroys free thiols, making reductive winemaking essential for preserving thiol character.
Schwefelverbindungen-Potential
Thiol-Konjugate
Thiol-Vorläufer-Akkumulation
polyfunctional thiol capacity
thiol conjugate disposition
volatile thiol precursor
Tominaga T et al. (1998) A powerful aromatic volatile thiol, 2-furanmethanethiol, exhibiting roast coffee aroma in wines made from several Vitis vinifera grape varieties. Journal of Agricultural and Food Chemistry 46:1044-1048 Roland A et al. (2011) Evolution of S-cysteine and glutathione conjugate contents during grape ripening. Journal of Agricultural and Food Chemistry 59:4958-4964
Thiol Precursor Accumulation Disposition
Thiol precursor data for Sauvignon Blanc from Marlborough (2024 vintage) records cysteinylated 3MH precursor (Cys-3MH) at 185 µg/L and glutathionylated 3MH precursor (Glut-3MH) at 520 µg/L, with the 2.8:1 glutathionylated-to-cysteinylated ratio typical of the cultivar, and total precursor pool predicting a maximum theoretical 3MH release of 3200 ng/L — though actual wine concentrations of 800-1800 ng/L (25-56% conversion efficiency) are expected depending on yeast strain selection and fermentation management. Thiol precursor data for Grüner Veltliner from Kremstal (2024 vintage) documents Cys-3MH at 42 µg/L and Glut-3MH at 95 µg/L, substantially lower than Sauvignon Blanc but sufficient to produce perceptible tropical-citrus aromatic notes (predicted wine 3MH of 200-400 ng/L) that contribute to the grapefruit character valued in premium Austrian Grüner Veltliner.
Thiol precursor data is an aromatic analysis data entity that records the concentration of non-volatile cysteinylated and glutathionylated conjugates of volatile thiols in grape berries and must, determined by LC-MS/MS (liquid chromatography-tandem mass spectrometry) or by indirect quantification through thiol release assays, expressed in micrograms per liter (µg/L) or nanomoles per liter (nmol/L), encompassing the conjugated precursors of 3-mercaptohexan-1-ol (3MH), 3-mercaptohexyl acetate (3MHA), and 4-mercapto-4-methylpentan-2-one (4MMP), and providing the quantitative assessment of latent tropical and citrus aromatic potential that is realized only through yeast-mediated enzymatic cleavage during alcoholic fermentation.
Thiol precursor data represents one of the most analytically challenging and oenologically informative components of aromatic analysis, addressing the paradox that the volatile thiols most responsible for the distinctive tropical, passionfruit, and grapefruit character of Sauvignon Blanc and related cultivars are essentially absent from grape berries and must — they exist exclusively as non-volatile, odorless conjugated precursors that require yeast enzymatic action (carbon-sulfur lyase activity) during fermentation for release. The two precursor classes — cysteinylated (Cys-3MH, Cys-4MMP) and glutathionylated (Glut-3MH, Glut-4MMP) — are biosynthesized through conjugation of the corresponding thiol with glutathione or cysteine via the glutathione S-transferase (GST) pathway in the berry, with the glutathionylated form typically 2-10× more abundant than the cysteinylated form. The conversion efficiency from precursor to free thiol during fermentation is critically dependent on yeast strain (Saccharomyces cerevisiae strains vary 10-fold in carbon-sulfur lyase activity), fermentation temperature (cooler fermentation preserves more released thiols from oxidative loss), and must nitrogen status (adequate YAN supports yeast enzymatic activity). Vineyard factors modulating precursor accumulation include nitrogen status (foliar nitrogen application at véraison increases cysteine conjugate formation), water stress (moderate deficit enhances precursor concentration through berry size reduction), UV-B exposure (stimulates glutathione biosynthesis), and harvest timing (precursor accumulation typically increases through late ripening). The analytical quantification of thiol precursors by LC-MS/MS requires deuterium-labeled internal standards and matrix-matched calibration for accurate results, as the ionization efficiency of conjugated thiols is strongly influenced by must matrix composition. Thiol precursor data is increasingly used alongside conventional must analysis to guide yeast strain selection and fermentation protocol design for optimizing varietal thiol expression in the finished wine.
Glutathionkonjugatdaten
Thiolpräkursordaten
Thiolvorläuferdaten
Thiolvorstufen
cysteinylated thiol data
cysteinylierte Thiolkonjugate
glutathionylated precursor data
thiol conjugate data
varietal thiol precursor content
volatile thiol precursor analysis
Roland A et al. (2011) Pre- and post-bottling evolution of volatile thiols in Sauvignon Blanc wines: key role of temperature. Food Chemistry 128(4): 1022-1035 Tominaga T et al. (1998) Identification of new volatile thiols in the aroma of Vitis vinifera L. var. Sauvignon Blanc wines. Flavour and Fragrance Journal 13(3): 159-162 Peyrot des Gachons C et al. (2002) Cysteinylated and glutathionylated precursors of 3-mercaptohexan-1-ol in Sauvignon Blanc. In Proceedings of the 13th International Enology Symposium, Montpellier, pp.: 319-324
Thiol Precursor Data
Thiol precursor quality assessment in Marlborough Sauvignon Blanc enables fermentation optimization, with grape precursor concentrations of 500-2000 μg/L Cys-3MH and 100-400 μg/L Cys-4MMP providing the substrate pool for tropical thiol release winemakers selecting high-β-lyase yeast strains (achieving 5-10% conversion versus 1-3% for standard strains) can maximize thiol release from available precursors, transforming the neutral-smelling juice into wines with 500-2000 ng/L 3MH and 20-50 ng/L 4MMP that define the tropical-passion fruit character of premium Marlborough style.
Thiol_Precursor_Quality encompasses the chemical qualities characterizing the concentration and composition of non-volatile cysteine and glutathione conjugates in grape berries that serve as precursors for the highly odor-active volatile thiols released during fermentation. These precursor compounds, synthesized in grape berries through conjugation of reactive thiol intermediates with cysteine or glutathione, are odorless in their bound form but can be cleaved by yeast enzymes (β-lyases, γ-lyases) during fermentation to release the free volatile thiols responsible for tropical fruit, passion fruit, grapefruit, and box tree aromas in Sauvignon Blanc and other varieties. The primary thiol precursors include S-3-(hexan-1-ol)-cysteine (Cys-3MH) and S-3-(hexan-1-ol)-glutathione (G-3MH) releasing 3-mercaptohexan-1-ol (3MH, passion fruit), and S-4-(4-methylpentan-2-one)-cysteine (Cys-4MMP) releasing 4-mercapto-4-methylpentan-2-one (4MMP, box tree, blackcurrant). Precursor accumulation in grapes depends on cultivar genetics, with Sauvignon Blanc showing highest levels followed by Colombard, Petit Manseng, and Gewürztraminer, while most other varieties accumulate minimal precursors. Viticultural factors including nitrogen nutrition, water status, and harvest timing modulate precursor content, with moderate nitrogen availability and controlled water deficit generally promoting accumulation. The conversion efficiency from precursor to free thiol during fermentation depends on yeast strain, with dramatic variation among commercial strains in β-lyase activity affecting the release of aromatic thiols from available precursors. This incomplete conversion (typically 1-10% efficiency) means that precursor concentration sets an upper limit on potential thiol release but does not directly predict final wine thiol content without considering fermentation factors.
Thiol_Precursor_Quality serves as the parent class for specific precursor compounds including 3MH_Precursor_Quality and 4MMP_Precursor_Quality. The term precursor emphasizes that these grape-derived compounds are odorless and require enzymatic conversion to release aromatic thiols. The distinction between cysteine conjugates (direct β-lyase substrates) and glutathione conjugates (requiring prior cleavage to cysteine form) affects conversion kinetics. Analysis of thiol precursors employs LC-MS/MS methods to quantify the non-volatile conjugates, distinct from the GC-MS methods used for released volatile thiols in wine.
Cystein-Konjugat-Gehalt
Glutathion-Konjugat-Konzentration
Thiol-Vorläuferqualität
bound thiol precursor quality
cysteine conjugate quality
gebundene Thiol-Vorläuferqualität
glutathione conjugate concentration
Tominaga T et al. (1998) Development of a Method for Analyzing the Volatile Thiols Involved in the Characteristic Aroma of Wines Made from Vitis vinifera L. Cv. Sauvignon Blanc. Journal of Agricultural and Food Chemistry 46(3): 1044-1048 Roland A et al. (2011) Evolution of S-Cysteinylated and S-Glutathionylated Thiol Precursors during Oxidation of Melon B. and Sauvignon Blanc Musts. Journal of Agricultural and Food Chemistry 59(9): 4510-4518
Thiol Precursor Quality
Souvignier Gris (released 2008, Freiburg, but representing third-generation breeding approaches) bears a Third_Generation_PIWI_Role as a white variety producing wines of recognized quality, with stacked downy and powdery mildew resistances enabling near-elimination of fungicide applications. Calardis Blanc (released 2017, JKI) serves in a Third_Generation_PIWI_Role as a contemporary white PIWI with pyramided resistance loci and wine quality suited to Rheingau-style production. INRA-ResDur breeding lines bearing Run1 + Rpv1 + additional loci represent Third_Generation_PIWI_Roles as advanced French materials approaching BC4 levels of V. vinifera genome recovery, targeting Champagne, Bordeaux, and Burgundy variety improvement.
A PIWI cultivar role that inheres in contemporary advanced disease-resistant varieties developed from 2010 onward, representing the current state-of-the-art in combining elite wine quality with effective, durable disease resistance through molecular breeding approaches. Third-generation PIWIs incorporate targeted R-gene introgression using marker-assisted selection, typically pyramiding multiple resistance loci (combining Rpv3 + Rpv10 + Rpv12 for downy mildew, Run1 + Ren3 for powdery mildew) to achieve durable resistance less likely to be overcome by pathogen evolution. These varieties aim to produce wines indistinguishable from or approaching V. vinifera quality in blind evaluation, addressing the residual quality gap that limited second-generation PIWI adoption. Third-generation breeding increasingly uses backcross strategies to produce resistant near-isogenic lines of established varieties (resistant Riesling, Chardonnay, Pinot derivatives), maintaining established variety identity while adding resistance. Regulatory and market acceptance of third-generation PIWIs is expanding, with EU organic regulations, climate adaptation pressures, and consumer sustainability concerns creating favorable conditions for resistant variety adoption.
Third_Generation_PIWI_Role captures the contemporary frontier of resistance breeding, where molecular tools enable precise R-gene combination and background selection accelerating quality improvement. The generation is distinguished by deliberate marker-assisted pyramiding (combining multiple R-genes for durable resistance), targeted backcrossing to elite V. vinifera varieties, and wine quality as a primary selection criterion coequal with resistance. Third-generation programs increasingly produce "resistant versions" of established varieties rather than novel variety identities—a strategy reflecting market research indicating consumer preference for familiar names over unfamiliar varieties regardless of quality. The durability question is central to third-generation breeding single R-gene resistance can be overcome by pathogen evolution, motivating pyramiding strategies that combine multiple independent resistance mechanisms. Registration timelines mean that many third-generation varieties remain in advanced trials, with commercial impact expected to accelerate through the 2020s and 2030s as current breeding lines complete evaluation and reach markets.
Elite-PIWI
PIWI dritte Generation
advanced PIWI
elite PIWI
fortgeschrittene PIWI-Sorte
modern resistant variety
moderne Resistenzsorte
molecular-bred PIWI
third-generation PIWI
Hausmann L et al. (2019) Grapevine breeding for disease resistance, Springer pp.: 311-330 Töpfer R et al. (2011) Molecular breeding in grapes, CRC Press pp.: 160-185 Zyprian E et al. (2016) Quantitative trait loci affecting pathogen resistance, Molecular Genetics and Genomics 291: 1573-1594 Schneider A et al. (2019) Marker-assisted breeding approaches, Euphytica 215: 45 INRA ResDur (2020) Program Progress Reports
Third Generation PIWI Role
A three-arm head on Vitis vinifera cv. Nebbiolo in traditional Barolo vineyard, featuring three arms emerging from head at 42cm height positioned at approximately 115°, 122°, and 123° intervals (nearly symmetrical 120° spacing), each arm extending 26cm bearing 3 two-bud spurs, total nine spurs supporting approximately 16 shoots and 2.4kg crop in deliberately restrained configuration emphasizing low yields for maximum concentration and quality, representing traditional Barolo three-arm approach maintained in prestigious estate where extreme quality focus justifies severe yield limitation, with three-arm symmetric structure considered optimal balance between adequate productivity and quality-focused restraint in this terroir-driven production philosophy.
A multi-arm head featuring exactly three arms radiating from central head, representing minimum arm count for radial symmetry, characterized by three-way branching typically at approximately 120° intervals creating balanced triangular or Y-shaped structure when viewed from above, suitability for moderate vine vigor, traditional systems in some regions, or deliberate minimalist approach emphasizing low yields and quality focus. Three-arm heads represent minimal multi-arm configuration, employed in low-vigor situations where three arms provide adequate capacity, traditional systems in certain regions favoring three-arm architecture, quality-focused approaches emphasizing yield restraint through limited arm numbers, or transitional training developing initial three arms with potential future addition of additional arms as vigor increases.
Three-arm heads provide minimal radially symmetric multi-arm structure, with three arms enabling balanced distribution (each arm carrying approximately one-third of vine capacity) and architectural stability (triangular geometry being inherently balanced). The three-arm configuration suits low-to-moderate vigor situations, quality-focused production accepting low yields, traditional systems in regions with three-arm heritage, and staged development approaches (establishing three arms initially, potentially adding more later). However, three-arm configurations limit total capacity (nine spurs at three per arm being minimal for economic viability in commercial operations), provide limited production insurance (loss of one arm represents 33% capacity loss), and may inadequately utilize vigorous vines or fertile sites where additional arms could increase productivity without quality compromise. The choice of three versus four or more arms reflects balance between yield restraint philosophy and economic viability requirements.
Dreiarm-Kopf
dreifache Kopferziehung
three-way head
tri-arm head
triple-arm head configuration
Three Arm Head
A three-branch permanent arm on Vitis vinifera cv. Grüner Veltliner in modified lyre training system in Wachau, Austria, featuring central trunk dividing at 75cm height into three branches= one central arm extending vertically upward 95cm bearing 8 spurs, two lateral arms extending at 45° angles left and right for 70cm each bearing 11 spurs each (22 spurs combined on laterals), total configuration with 30 spurs distributed across three distinct branches creating divided canopy with improved light penetration compared to simple bilateral cordon, representing deliberate three-branch architecture adapted to steep terraced vineyard slopes where divided canopy structure facilitates management on challenging terrain while maximizing productive capacity within space constraints.
A multi-branched permanent arm featuring exactly three distinct branches emerging from trunk or primary arm structure, characterized by triple division creating three separate fruiting zones, typical configurations including central vertical arm with two lateral arms, three-way symmetrical branching at approximately 120° intervals, or sequential branching creating three terminal segments, and suitability for specialized training systems requiring tri-partite canopy division or adaptation to specific spacing or terrain constraints. Three-branch permanent arms represent minimum true multi-branched configuration (beyond simple bilateral which is two-branch), employed in specific contexts including modified lyre systems with central plus bilateral arms, adaptation to triangular or irregular vine spacing, traditional systems in some regions employing three-arm architectures, or transitional configurations during training system modifications adding third arm to existing bilateral framework.
Three-branch permanent arms occupy middle ground between simple bilateral (two-branch) and more complex four-branch or higher configurations, providing modest canopy division benefits without extreme structural complexity. The three-branch configuration allows various geometric arrangements depending on objectives= symmetrical three-way branching (120° intervals) creates balanced radial structure, central-plus-bilateral creates primary vertical with supporting laterals, sequential branching creates linear progression of three segments. Establishment requires careful training to achieve desired branch angles and positions, with particular attention to junction points where three-way divisions concentrate mechanical stress. Management involves maintaining balance between three branches preventing dominance of one over others, which can be challenging given apical dominance effects and differential positioning relative to sun, wind, and other environmental factors affecting branch vigor.
Dreigabelarm
dreifach verzweigte Arm
three-way arm
tri-branch arm
triple arm configuration
Three Branch Permanent Arm
A three-bud compound structure at node 8 on Vitis vinifera cv. Cabernet Sauvignon, with careful dissection revealing distinct primary bud (largest, 2.8mm diameter, two inflorescence primordia), secondary bud (1.4mm diameter, one small inflorescence primordium, positioned laterally to primary), and tertiary bud (0.6mm diameter, vegetative only, positioned opposite secondary), all enclosed within common protective bud scales, with only primary bud showing signs of dormancy release in early spring.
A compound dormant bud exhibiting the complete canonical architecture of primary, secondary, and tertiary bud positions all present and distinguishable within the compound structure, representing the maximum developmental complexity possible in grapevine buds. Three-bud compound structures provide triple redundancy for shoot development, with hierarchical activation typically proceeding from primary to secondary to tertiary in response to damage or strongly forcing conditions, though under normal conditions only the primary bud breaks dormancy and develops. This architecture reflects the evolutionary adaptation of Vitis to environments with unpredictable growing season hazards including late spring frosts, hail damage, and herbivory.
Not all compound buds contain all three positions equally developed basal buds often lack tertiary buds, and apical buds may show reduced secondary development. The three-bud structure is most consistently expressed in mid-cane positions (nodes 3-8 typically) on vigorous canes from previous season's growth. Understanding three-bud architecture is essential for assessing potential compensation after spring frost events, where secondary bud breakage can partially offset primary bud loss, though typically with reduced yield and delayed development.
complete compound bud
full compound structure
triple-primordium bud
vollständige zusammengesetzte Knospe
Based on Pratt (1971) "Reproductive anatomy in cultivated grapes - a review" American Journal of Enology and Viticulture 22=92-109 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Friend et al. (2009) "Influence of pruning and crop load on the growth and yield components of Pinot Noir in the Tamar Valley of Tasmania" Australian Journal of Grape and Wine Research 15=55-62.
Three Bud Compound Structure
A 30 cm three-node cutting of 110 Richter rootstock bears a Three_Node_Cutting_Role, prepared for benchgrafting with standard dimensions matching callusing box specifications—all nodes disbudded except the basal one (which produces root system shoots if the benchgraft is planted deeply), providing maximum rooting surface area along the cutting length. A three-node hardwood cutting of Ramsey (Salt Creek) serves in a Three_Node_Cutting_Role for direct field planting in nematode-infested soils, where the longer cutting enables deeper planting establishing roots below the nematode-concentrated upper soil layers. A three-node cutting of Grenache bears a Three_Node_Cutting_Role in traditional southern French own-rooted propagation, field-rooted in nursery rows for dormant-plant sales to growers establishing vineyards in phylloxera-free sandy soils.
A cutting role that inheres in a stem segment prepared with three nodes bearing three buds plus associated internode tissue (typically 28-35 cm total length), representing the traditional standard format for commercial grapevine propagation that provides substantial carbohydrate reserves, multiple-bud redundancy, and adequate length for conventional planting depth. Three-node cuttings offer maximum reliability under field conditions, with sufficient reserves to sustain root initiation and early shoot growth even in suboptimal environments, redundancy ensuring success if individual buds fail, and length enabling planting at depths providing protection from surface desiccation and temperature extremes. The three-node format dominates traditional rootstock cutting propagation (cuttings planted directly in nursery rows or vineyards for field rooting), benchgraft rootstock component preparation (standard callusing box dimensions accommodate 28-32 cm cuttings), and own-rooted vine production in regions without phylloxera. Multiplication rate is lower than single or two-node formats (3-5 propagules per meter of source cane) but establishment success is highest, reducing costly nursery losses and replanting requirements.
Three_Node_Cutting_Role represents the conservative, reliability-focused end of the multiplication efficiency spectrum, prioritizing establishment success over propagation rate. The format remains standard for commercial rootstock production despite lower multiplication efficiency because nursery economics strongly penalize cutting failures—a 10% improvement in rooting success may offset 50% reduction in cutting numbers when labor and infrastructure costs are considered. Three-node cuttings provide flexibility in planting depth, enabling adjustment to site conditions and supporting both shallow (one node above soil) and deep (all nodes below soil) planting configurations. The format's dominance in benchgraft rootstock component production reflects standardized callusing system dimensions developed around 30 cm cutting lengths, with the full grafted unit (30 cm rootstock plus 5-8 cm scion) fitting standard box dimensions.
Dreiaugensteckling
Dreiknoten-Steckling
Standardsteckling
standard cutting
three-bud cutting
three-node cutting
triple-eye cutting
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 350-355 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Whiting JR (2004) Grapevine rootstocks, Winetitles pp.: 167-188 Dry N (2007) Grapevine rootstocks, Lythrum Press
Three Node Cutting Role
A three-quarter expanded young leaf at node 7 on a shoot of Vitis vinifera cv. Syrah in the Rhône Valley, France, measuring 14.2 cm length by 13.5 cm width representing approximately 76% of expected mature dimensions of 18.7 cm × 17.8 cm for this position and cultivar, lamina area of 125 cm² representing 58% of expected mature area of 215 cm², exhibiting fully flat architecture with complete five-lobed structure showing all final characteristics including fully developed sinuses and marginal teeth, medium-dark green coloration closely approaching mature appearance with chlorophyll content of 455 μg cm⁻², no visible anthocyanin pigmentation, firm texture very similar to mature leaves with palisade cells averaging 33 μm diameter (94% of mature 35 μm), nearly mature tissue organization with two palisade layers totaling 80 μm (94% of mature 85 μm) and spongy mesophyll of 115 μm (96% of mature 120 μm), fully functional stomata with complete environmental responsiveness showing conductance patterns identical to mature leaves, essentially complete cuticle development creating glossy surface appearance indistinguishable from mature leaves, demonstrating photosynthesis rate of 10.8 μmol CO₂ m⁻² s⁻¹ representing 82% of mature capacity and achieving approximate carbon balance with daily photosynthetic production of 48 mg carbohydrate approximately equaling respiratory costs and minimal remaining growth requirements, occurring at 22 days post-emergence with maturity expected within 5-7 days, measured at shoot BBCH stage 81 (beginning of berry ripening) in early August, representing final expansion stage immediately preceding mature status.
A partially expanded young leaf in the late expansion phase achieving approximately three-quarters of final mature dimensions in Vitis vinifera, characterized by dimensions typically 70-80% of cultivar-specific and position-specific mature size with length and width both in 75% ± 5% range, lamina area approximately 50-65% of final mature area, fully flat lamina with all architectural features completely developed showing final form, medium to medium-dark green coloration with chlorophyll concentrations typically 400-480 μg cm⁻² representing 80-95% of mature levels, essentially no anthocyanin remaining indicating maturation of photoprotective mechanisms, moderately firm to firm texture approaching mature consistency with cells nearing final dimensions typically 90-95% of mature size, advanced tissue differentiation with palisade and spongy mesophyll layers showing nearly mature organization, nearly complete vascular development with functional mature xylem and phloem, fully functional stomata responding appropriately to all environmental signals, nearly complete cuticle development with most wax deposition finished creating surface appearance similar to mature leaves, and photosynthetic capacity typically 75-88% of mature leaf rates approaching carbon balance where daily photosynthetic production nearly equals or slightly exceeds respiratory and minimal remaining growth costs making these leaves transitional to net source status. Three-quarter expanded young leaves represent the final expansion phase approaching maturity, with cell expansion in final stages as cells approach ultimate dimensions, essentially all size determination complete with remaining growth representing final adjustments typically <10% additional area increase, tissue maturation processes nearly complete including final cell wall thickening, chloroplast maturation, and full metabolic activation, and functional capabilities closely approaching mature levels with leaves beginning significant contribution to vine carbon balance and photosynthate export to support fruit development and other sinks.
Three-quarter expanded young leaves represent the late expansion checkpoint approaching maturity, distinguished from earlier partially expanded stages by near-final dimensions and tissue characteristics, and from fully mature leaves by incomplete final size achievement and continued minimal growth. Assessment requires comparison to mature size norms considering cultivar and position variation. The three-quarter expansion stage typically occurs at approximately 70-80% of total expansion duration, representing the final expansion phase before maturity transition. Three-quarter expanded leaves show nearly mature characteristics across most parameters, with remaining development representing refinement rather than major changes, making these leaves functionally similar to mature leaves despite slightly reduced dimensions and photosynthetic capacity. The transition from three-quarter expansion to full maturity is gradual rather than abrupt, with exact maturity timing somewhat arbitrary but typically defined by cessation of measurable dimensional increase over 5-7 day periods. Stress during three-quarter expansion has minimal impact on final size as most growth is complete, but can affect final functional capacity through effects on chloroplast maturation, enzyme synthesis, or cuticle development affecting mature leaf performance.
dreiviertel-expandiertes junges Blatt
fast-endgröße Weinblatt
fünfundsiebzig Prozent ausgedehnt Blatt
late expansion leaf
near-final size leaf
seventy-five percent expanded leaf
Based on Palliotti & Cartechini (2000) "Developmental changes in gas exchange activity in flowers, berries, and tendrils of field-grown Cabernet Sauvignon" American Journal of Enology and Viticulture 51=317-323 Flexas et al. (2002) "Drought-inhibition of photosynthesis in C₃ plants= stomatal and non-stomatal limitations revisited" Annals of Botany 89=183-189 Poni et al. (2009) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 60=397-407 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 233-258.
Three Quarter Expanded Young Leaf
The threshold specification for powdery mildew (Erysiphe necator) management in a Cabernet Sauvignon vineyard in Margaret River defines phenologically differentiated action thresholds — pre-flowering (E-L 12-19) threshold of 3% leaf incidence triggering protectant fungicide application, flowering through bunch closure (E-L 19-33) threshold reduced to 1% leaf incidence or any cluster infection triggering immediate systemic curative application (reflecting the high susceptibility of developing berries during this window and the devastating quality impact of cluster infection), and post-véraison (E-L 35+) threshold relaxed to 10% leaf incidence with no cluster intervention (berries become resistant to new infection after véraison as epidermal cells mature and stomata close). For Botrytis cinerea, the threshold is set at 2% cluster incidence at E-L 33 (bunch closure) triggering pre-véraison botryticide application, with a separate harvest-decision threshold of 5% cluster severity above which affected clusters must be excluded during sorting.
A pest management description that documents the quantitative pest population levels, disease incidence values, or damage intensity ratings at which management intervention is triggered, specifying the action threshold (the level at which treatment is applied), the economic injury level (the level at which pest damage cost exceeds treatment cost), and the phenological context in which these thresholds apply, providing the decision rules that translate monitoring data into treatment actions within the IPM framework.
Threshold_Specification translates the theoretical IPM concept of intervening only when necessary into quantitative operational decision rules. The distinction between the economic injury level (EIL — the pest level at which damage cost equals treatment cost) and the action threshold (AT — the pest level at which treatment must be applied to prevent the population from reaching the EIL, accounting for population growth rate and treatment lag time) is fundamental to threshold-based management but rarely calculated rigorously in viticultural practice, where action thresholds are more commonly established empirically through regional trial and observation: the difficulty of EIL calculation in viticulture arises from the non-linear relationship between disease severity and wine quality loss (a 5% increase in botrytis severity may cause negligible quality impact in a dry rosé wine but devastating quality loss in a premium Pinot Noir), requiring quality-adjusted rather than purely economic thresholds for premium wine production. Thresholds must be phenologically specific because the susceptibility of grapevine organs to pest and disease damage varies dramatically across developmental stages — berry susceptibility to powdery mildew declines sharply after véraison, while botrytis susceptibility increases progressively from bunch closure through harvest.
Bekämpfungsschwellen-Spezifikation
Eingriffsschwellen-Festlegung
Schadschwellen-Dokumentation
action threshold specification
economic threshold documentation
intervention trigger level
Wilcox WF et al. (2015) Compendium of Grape Diseases, Disorders, and Pests 2nd Ed, APS Press Pertot I et al. (2017) A critical review of plant protection tools for reducing pesticide use on grapevine, Crop Protection 97: 70-84 Stern VM et al. (1959) The integrated control concept, Hilgardia 29: 81-101 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 461-520
Threshold Specification
A tightly coiled functional tendril at node 8 on a shoot of Vitis vinifera cv. Syrah in the Rhône Valley, France, exhibiting six contact coils around a 3 mm diameter trellis wire with individual coil diameters of 10-12 mm (approximately 3.5× the 3 mm tendril diameter), pitch between coils of 4-5 mm creating nearly touching successive turns, nine free coils in the tendril shaft with similar tight geometry and characteristic handedness reversal, generating visible shoot displacement of 8 cm toward the wire compared to pre-attachment position indicating strong contractile force, showing dark brown coloration throughout coiled regions from extensive lignification completed within 48 hours of initial contact, supporting a shoot bearing six leaves and two fruit clusters totaling 580 grams weight without visible strain or deformation, measured at BBCH stage 79 (majority of berries touching) in early August.
A coiled functional tendril exhibiting maximum coil density and minimal coil diameter relative to tendril thickness in Vitis vinifera, characterized by contact coils with diameters typically 3-5 times the tendril diameter creating very close wrapping around support structures, free coils showing similar tight spiraling with minimal pitch (vertical distance between successive coil turns), high mechanical tension in coiled regions generating strong contractile forces that pull the shoot closer to the support, and extensive structural reinforcement through dense lignification and abundant gelatinous fiber formation that fixes the tight coil geometry permanently. Tightly coiled functional tendrils develop under conditions of optimal contact stimulation including sustained pressure on mechanosensory cells, appropriate support structure diameter matching tendril grasping capacity (typically 2-8 mm), favorable temperature and moisture conditions during the critical 6-24 hour coiling period, and absence of mechanical disturbances that might interrupt coil formation. These tendrils generate maximum grip force on supports through high coil density creating multiple contact points and friction zones, and produce strongest contractile effects drawing shoots toward supports with forces potentially reaching several hundred grams tension. Tightly coiled tendrils show enhanced resistance to mechanical failure under loading, with the compact coil geometry distributing stresses efficiently and the extensive lignification providing structural integrity, making these the most reliable attachment structures for supporting heavy fruit loads and resisting storm damage in commercial vineyards.
Tightly coiled functional tendrils are distinguished from loosely coiled tendrils by quantitative coil geometry parameters including coil diameter relative to tendril thickness (ratio <5=1 indicates tight coiling), pitch between coil turns (minimal spacing), and visible contractile effects on shoot position. Tight coiling represents optimal mechanical performance but requires specific conditions including appropriate support diameter - supports too thick prevent tight coiling while supports too thin may be crushed. Cultivar variation exists in coiling tightness preference, with some varieties consistently producing tighter coils than others under identical conditions. Tight coiling may be disadvantageous in situations requiring accommodation of shoot growth movements, potentially creating excessive mechanical constraint or tendril breakage if shoots are displaced by wind or cultural operations.
compact coil tendril
densely coiled tendril
dicht spiralisierte Ranke
fest gewickelte Ranke
high-tension tendril
hochgespannte Kletterorgan
Based on Isnard & Silk (2009) "Moving with climbing plants from Charles Darwin's time into the 21st century" American Journal of Botany 96=1205-1221 Goriely & Tabor (2013) "Spontaneous helix hand reversal and tendril perversion in climbing plants" Physical Review Letters 80=1564-1567 Silk & Hubbard (1991) "Axial forces and normal stresses in coiling tendrils" Biomechanics and Modeling in Mechanobiology 24=599-606 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 119-136.
Tightly Coiled Functional Tendril
Pinot Noir (standard clones) in Oregon's Willamette Valley at BBCH 65 exhibits tightly compact flower clusters of 9-12 cm length bearing 200-280 flowers with flower densities of 20-25 flowers/cm, creating well-filled cylindrical clusters (OIV code 6-7) that develop into moderately compacted berry clusters with 8-12% Botrytis losses in typical vintage years with 5-6 fungicide applications. Chardonnay in Burgundy's Côte de Beaune shows similarly tight flower clusters (220-300 flowers, 10-13 cm length) requiring cluster thinning to 6-8 clusters per shoot and leaf removal at BBCH 77 to reduce final compactness and disease pressure. Sangiovese in Chianti Classico exhibits tightly compact clusters (OIV code 7) with cultural management (crop load 1.2 kg/m canopy, leaf removal) reducing rot incidence to 5-8% in typical seasons.
A tightly compact cluster is a compact flower cluster exhibiting high flower density with flowers closely spaced but still individually distinguishable, representing moderate-to-high compactness that creates significant disease management challenges while remaining amenable to cultural interventions for compactness reduction and typically developing into compressed but not severely deformed berry clusters. Tightly compact clusters exhibit flower densities typically ranging from 18-28 flowers per cm of total inflorescence axis length, achieved through high flower initiation combined with moderate rachis and branch elongation and short-to-moderate pedicel length (2-4 mm). These clusters typically measure 8-14 cm in length at BBCH 65 while bearing 180-350 flowers, creating dense but not continuous flower coverage with the underlying axis still partially visible between flowers. The tight compactness results from genetic programming for moderately reduced internode elongation, often combined with reduced secondary branching and absent or minimal tertiary branching such that most flowers arise from the main rachis and moderately developed primary branches (1-4 cm). Tightly compact flower clusters characteristically show cylindrical or conical shapes with length=width ratios of 2.5=1-3.5=1, creating a full, well-filled appearance without the club-like density of extremely compact types. Flower spacing within tightly compact clusters typically ranges from 1-2 mm between adjacent flower pedicels, creating near-touching flower distribution but still allowing some air penetration between flowers. The close flower arrangement creates a moderately humid microclimate with relative humidity 8-15% above ambient within the cluster, drying times after precipitation extended by 2-4 hours compared to loose clusters, and restricted but not absent air circulation through the flower mass. This microclimate elevates disease risk significantly, with tightly compact flower clusters showing 2-4× higher Botrytis cinerea infection rates at bloom compared to loose clusters under identical conditions, but disease pressure remains manageable with standard fungicide programs (4-7 applications) and cultural practices. The transformation from tightly compact flower cluster to tightly compact berry cluster involves moderate architectural consequences, with fruit set of 30-50% creating berry crowding by BBCH 77-79, berries touching and experiencing mutual compression but not severely deformed, some skin cracking and splitting in highly susceptible cultivars, and bunch rot affecting 5-15% of clusters in moderately humid climates with standard fungicide programs. Tightly compact flower cluster morphology characterizes cultivars like Pinot Noir (standard clones), Chardonnay, Chenin Blanc, and Sangiovese, representing OIV codes 6-7. Cultural interventions for tightly compact clusters include moderate crop load management during previous-season bud differentiation, selective cluster thinning at BBCH 71-75 to improve canopy microclimate and resource allocation, targeted leaf removal in the fruiting zone at BBCH 77-79 to enhance air circulation, and optional berry thinning at véraison in high-value production to open cluster architecture, reducing disease risk by 30-60% compared to untreated tightly compact clusters.
Tightly_Compact_Cluster is distinguished from Densely_Packed_Compact_Cluster by moderate rather than extreme density (18-28 flowers/cm versus >28 flowers/cm), retention of individual flower visibility, and more manageable disease consequences (2-4× versus 5-10× Botrytis risk relative to loose clusters). This class corresponds to OIV codes 6-7 and represents the compactness level of many economically important wine cultivars. Tightly compact architecture creates challenges but remains compatible with sustainable viticulture through integrated pest management, whereas densely packed types often require intensive chemical inputs. The boundary between 'tightly compact' and 'moderately compact' (OIV code 5) is gradational but conventionally distinguished at approximately 18 flowers/cm density threshold. Cultural management can shift tightly compact morphology toward moderate compactness at the berry stage through flower/fruit removal, providing growers with meaningful intervention options.
closely packed flower cluster
dicht kompakte Blütentraube
eng kompakte Blütentraube
fest gepackte Blütentraube
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 95-97, 146-147 OIV (2009). Descriptor List for Grape Varieties and Vitis Species, OIV codes 6-7 Hed, B. et al. (2009). Evaluation of cultural practices for management of Botrytis bunch rot. Plant Disease 93=1148-1154 English, J.T. et al. (1989). Relationship between vegetative growth and development of grape cluster morphology. Journal of the American Society for Horticultural Science 114=234-238.
Tightly Compact Cluster
Merlot inflorescences at BBCH 55 (flower buds separating, inflorescence clearly visible) exhibit tightly sealed intact calyptras 2.2-2.8 mm length, dark green color, firmly attached at base with no visible loosening, closely conforming to contours of enclosed stamens (1.2-1.6 mm) and pistil (1.1-1.4 mm), maintaining this tightly sealed state for 6-9 days (depending on temperature) before transitioning to pre-dehiscence swollen stage as flowers approach anthesis. Sauvignon Blanc at BBCH 53-55 shows tightly sealed intact calyptras with smooth firm surfaces, no swelling, and reproductive organs visible as subtle contours through the thin green calyptra tissue under magnification. Grenache experiencing late frost (-1°C) at BBCH 55 with tightly sealed intact calyptras suffers severe damage with 70-85% of flowers showing brown collapsed calyptras, dead enclosed organs visible upon dissection, and complete failure of subsequent anthesis and fruit set in affected flowers.
A tightly sealed intact calyptra is an intact calyptra in the early to mid-developmental stage showing firm attachment to the receptacle, closely appressed petal surfaces conforming tightly to the contours of enclosed developing organs, minimal internal airspace, and no visible signs of loosening or impending dehiscence, representing flowers in early pre-anthesis stages typically 5-10 days before calyptra shedding with reproductive organs still undergoing active development and maturation. Tightly sealed intact calyptras are characterized by= (1) calyptra firmly and uniformly attached around the entire basal circumference with abscission layer not yet formed or in very early formation stages showing no visible separation (2) petal tissue closely conforming to the surface contours of enclosed stamens and pistil with minimal gap or airspace between calyptra inner surface and organ surfaces, creating a snug-fitting enclosure (3) petal apices tightly fused at the cap apex with no loosening or gap formation (4) calyptra surface appearing smooth or slightly ribbed, dark to medium green in color, with firm texture indicating turgid cells and lack of senescence or preparation for abscission (5) external appearance showing no swelling, bulging, or displacement suggesting internal pressure from elongating organs. The tightly sealed state characterizes intact calyptras during BBCH stages 53-57 (individual flower buds visible through flower buds separating), representing the period when enclosed stamens are completing anther development and pollen maturation while pistils are completing ovule maturation and preparing stigmatic receptivity tissues. Tightly sealed intact calyptras provide maximum protective function, creating an enclosed microenvironment with= (1) relative humidity typically 20-30% above ambient air, maintaining optimal moisture levels for developing pollen and preventing premature desiccation (2) temperature buffering reducing diurnal temperature fluctuation amplitude by 30-50% compared to ambient, protecting temperature-sensitive developmental processes (3) complete exclusion of pathogens, preventing Botrytis cinerea and other fungal infections from accessing developing reproductive tissues (4) physical protection against mechanical damage from rain, hail, wind, or insect activity. The internal organs during the tightly sealed calyptra stage show specific developmental characteristics= stamens typically 1-2 mm long with anthers completing meiosis and microsporogenesis, producing mature pollen grains but not yet elongating filaments pistil typically 1-1.5 mm long with ovules completing megasporogenesis and embryo sac development, preparing stigmatic papillae and secretory cells but not yet fully receptive. The duration of the tightly sealed stage varies with temperature, cultivar, and environmental conditions, typically persisting for 7-12 days under cool spring conditions (12-18°C) but only 4-6 days under warm conditions (20-25°C), with warmer temperatures accelerating the developmental progression toward pre-dehiscence and separating stages. Tightly sealed intact calyptras are vulnerable to environmental stresses that can damage enclosed organs before protective shedding occurs= frost damage during tightly sealed stage causes browning and collapse of calyptra tissue, death of enclosed reproductive organs, and complete flower failure excessive heat (>35°C) during tightly sealed stage can cause pollen sterility within the enclosed anthers, reducing subsequent fruit set even if calyptra shedding proceeds normally prolonged cool temperatures (<12°C) can arrest development during tightly sealed stage, delaying transition to pre-dehiscence and extending vulnerability period to spring frost events.
Tightly_Sealed_Intact_Calyptra represents the early-to-mid intact calyptra developmental stage and is distinguished from Pre_Dehiscence_Intact_Calyptra by absence of swelling, loosening, or color changes indicating imminent dehiscence, and by the longer time frame (typically 5-10 days) before calyptra shedding. The tightly sealed stage provides maximum protective function but also represents a period of vulnerability to environmental extremes (frost, heat) that can damage enclosed organs. Monitoring tightly sealed calyptra health and environmental conditions during this stage is critical for predicting bloom success and implementing protective measures (frost protection, disease prevention) when needed. The tightly sealed stage corresponds to the period when pre-bloom fungicide applications (BBCH 53-57) are typically made to prevent Botrytis infection before calyptra shedding exposes floral tissues.
eng anliegende Kelchhaube
eng geschlossene Kronblattkappe
fest verschlossene Blütenhaube
firmly sealed calyptra
tightly enclosed petal cap
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 99-101 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 93-94 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 191-193.
Tightly Sealed Intact Calyptra
The tillage protocol for a Riesling vineyard in the Wachau specifies under-vine mechanical tillage using a hydraulic intercep (Clemens radius SL) with a 30 cm working width operating at 3-5 cm depth beneath the vine row to sever emerging weeds without damaging shallow feeder roots, performed 4-5 times between April and July — inter-row tillage consists of 2 passes with a disc cultivator at 10-12 cm depth in March (to break winter surface crusts and incorporate cover crop residue) and June (to terminate spring cover crop growth and reduce water competition during the pre-véraison deficit period), with no tillage after July 1 to allow natural revegetation that stabilizes soil during autumn rains on the steep Wachau terraces where erosion risk on 30-60% slopes is the primary soil management concern.
A soil management description that documents the mechanical cultivation of vineyard soil using implements that cut, turn, or loosen the soil profile, specifying the implement type, working depth, timing relative to vine phenology and rainfall, frequency, and operational objectives (weed control, soil aeration, incorporation of organic matter, breaking of surface crusts) that collectively define the physical disturbance regime applied to the vineyard floor.
Tillage_Protocol documents the oldest and most mechanically direct form of vineyard soil management. Tillage serves multiple simultaneous functions — weed suppression (historically the primary objective, now increasingly replaced by cover cropping and targeted herbicide use), soil structure improvement (breaking compaction and surface crusts), organic matter incorporation (mixing surface-applied compost or cover crop residue into the root zone), and water infiltration enhancement (loosening compacted surfaces to improve rainfall capture): however, tillage also carries well-documented negative effects including destruction of soil aggregate structure, acceleration of organic matter decomposition through aeration, disruption of mycorrhizal hyphal networks, damage to shallow grapevine feeder roots (concentrated in the upper 10-20 cm), and increased erosion susceptibility on sloped sites. The contemporary trend is toward reduced tillage intensity — shallower working depths (5-10 cm versus the traditional 20-30 cm ploughing), fewer passes per season, and strategic alternation between tilled and untilled inter-rows — that captures tillage's benefits while minimizing soil structural damage.
Bodenbearbeitungs-Protokoll
Pflüge-Spezifikation
cultivation protocol
mechanical tillage specification
mechanische Kultivierung-Dokumentation
vineyard ploughing record
White RE (2015) Understanding Vineyard Soils 2nd Ed, Oxford University Press pp: 185-210 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 425-440 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 195-220
Tillage Protocol
Tip burn quality assessment for calcium deficiency in Oregon Pinot Gris on an acidic site identified Ca-limited growth affecting new leaf development during the rapid spring growth period. Visual examination revealed tip burn quality symptoms concentrated on the youngest 3-4 leaves at each shoot tip, with brown necrotic tissue at leaf apices extending 1-2 cm and marginal necrosis creating a continuous brown border around expanding leaves. The tip burn quality distribution confirmed calcium deficiency diagnosis, with symptoms restricted to newest growth while older leaves (positions 6 and below) showed no symptoms, consistent with the immobile-nutrient pattern. Severity scoring of tip burn quality on a 0-4 scale showed mean scores of 2.8 on the youngest emerged leaf (substantial tip and margin necrosis), 1.5 on the second leaf (moderate tip browning), and 0.3 on the fourth leaf (trace symptoms), with the acropetal severity gradient reflecting the progressive calcium depletion as newer tissues competed for limited supply. Shoot tip examination revealed tip burn quality affecting the apical meristem in severely deficient vines, with growing points showing brown discoloration and arrested development that would limit continued shoot elongation if uncorrected. Environmental analysis identified the primary predisposing factor as low soil pH (4.6) limiting calcium availability despite adequate total soil Ca at 1,200 ppm, compounded by rapid spring growth creating high calcium demand that the compromised root system could not meet. Corrective lime application (3 t/ha dolomitic lime) combined with foliar CaCl₂ (0.5% solution) resolved tip burn quality symptoms within 3 weeks, with new growth emerging normally as calcium availability improved.
Tip burn quality in calcium deficiency is a calcium deficiency symptom quality that inheres in young grapevine leaves and shoot tips and represents the characteristic necrosis of leaf tips and margins on expanding leaves resulting from inadequate calcium supply for cell wall construction during rapid tissue expansion, with symptoms appearing as brown, desiccated tissue at leaf apices and margins that may curl downward, affecting the youngest leaves at shoot tips while older leaves remain symptom-free due to calcium's immobility preventing redistribution from mature tissues to expanding growth. This quality captures the most characteristic foliar symptom of calcium deficiency, recognizing that tip burn on youngest leaves provides strong diagnostic indication of calcium limitation when the immobile-nutrient pattern of young-tissue symptom expression is confirmed. The ontological significance of tip burn quality in calcium deficiency lies in its diagnostic utility for identifying calcium limitation in the field, with the restriction of symptoms to young expanding tissues clearly distinguishing calcium from mobile nutrients that show symptoms on oldest tissues first. The biological basis of calcium-deficiency tip burn encompasses the structural requirement for calcium in cell wall pectin cross-linking during cell expansion, the phloem immobility of calcium preventing redistribution from mature tissues, and the consequent dependence of expanding tissues on current xylem calcium supply that may be inadequate during periods of rapid growth or when transpiration-driven xylem flow preferentially supplies mature transpiring leaves rather than young expanding tissues. The distribution of tip burn symptoms on leaf margins and tips reflects the growth pattern of leaves where margin and tip expansion continues after central blade regions have matured, creating continued calcium demand at positions distant from major veins where xylem calcium delivery may be limited. The severity of tip burn ranges from subtle browning at leaf tips through substantial marginal necrosis to severe cases where expanding leaves emerge already damaged, with progression indicating the duration and intensity of calcium limitation.
Tip_Burn_Quality_(Ca) is_a Calcium_Deficiency_Symptom_Quality, representing the necrosis of young leaf tips and margins from Ca-limited cell wall construction during expansion. This class addresses the most characteristic foliar symptom of Ca deficiency. Symptoms restricted to youngest leaves confirms immobile nutrient diagnosis. Tip and margin distribution reflects continued expansion at positions distant from major veins. Severity gradient from youngest (most affected) to older leaves. Rapid spring growth periods create high Ca demand vulnerable to deficiency. Low soil pH limits Ca availability even with adequate total soil Ca.
Ca-Limitierungs-Spitzentod
Ca-deficiency tip necrosis
Ca-limitation tip death
Jungblatt-Randnekrose
Kalzium-Spitzenverbrennung
Spitzenbrand bei Ca-Mangel
calcium tip burn
young leaf margin necrosis
Tip Burn Quality (Ca)
The tissue analysis protocol for a Cabernet Sauvignon vineyard in Napa Valley specifies petiole sampling at two phenological stages — at full bloom (E-L 23, 50% cap fall) collecting petioles from leaves opposite the lowest inflorescence on 40-60 randomly selected vines per management block, and at véraison (E-L 35) collecting petioles from the same leaf position on 40-60 vines — petioles are separated from blades immediately, placed in paper bags, air-dried or oven-dried at 65°C within 6 hours of collection, ground to pass a 40-mesh sieve, and submitted for ICP-OES analysis of N (Kjeldahl or Dumas combustion), P, K, Ca, Mg, S, Fe, Mn, Zn, Cu, B, and Na, with results interpreted against the Robinson (2005) reference ranges for Cabernet Sauvignon petioles at bloom — N adequate at 0.60-1.20%, P adequate at 0.15-0.50%, K adequate at 1.50-3.00%, Mg adequate at 0.30-0.80%.
A nutrition management description that documents the systematic collection and laboratory analysis of grapevine tissue samples to assess vine nutritional status, specifying the tissue type (petiole, leaf blade, or whole leaf), sampling timing (flowering or véraison), sample size, collection and handling procedures, analytical methods, and the reference value ranges used to interpret results as deficient, adequate, or excessive for each nutrient element.
Tissue_Analysis_Protocol provides the vine-centered nutritional diagnostic that complements soil analysis by measuring the nutrients actually taken up and present in vine tissues rather than those potentially available in the soil. The choice of tissue type, sampling timing, and sampling position on the vine significantly affects analytical results and their interpretation: petiole analysis at flowering is the most widely standardized protocol globally (Robinson 2005, Keller 2020) because petioles at this stage provide maximum sensitivity to nutrient supply variation, particularly for the mobile nutrients nitrogen, phosphorus, and potassium whose petiole concentrations reflect current-season supply conditions. Leaf blade analysis is preferred in some regions (particularly Australia and parts of California) because blade concentrations show less seasonal variability and provide better diagnosis of micronutrient status. The interpretation of tissue analysis results requires cultivar-specific reference ranges — nutrient concentration norms established through extensive correlation of tissue values with visual symptoms, yield responses, and wine quality parameters — though comprehensive cultivar-specific norms are available for only approximately 20 major cultivars, requiring extrapolation for less-studied varieties.
Blattstiel-Analyse-Spezifikation
Gewebeanalyse-Protokoll
Rebenernährungs-Diagnostik
petiole analysis specification
plant tissue testing protocol
vine nutritional diagnostic procedure
Robinson JB (2005) Critical plant tissue values and application of nutritional standards for practical use in vineyards, in Christensen LP and Smart DR (eds) Proceedings of the Soil Environment and Vine Mineral Nutrition Symposium, ASEV pp: 61-68 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 160-195 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 150-168
Tissue Analysis Protocol
Tissue density analysis of Shiraz berries at harvest reveals mesocarp density of 1.067±0.008 g/mL, exocarp density of 1.089±0.012 g/mL, and seed density of 1.152±0.015 g/mL, with the 8% higher skin density compared to pulp contributing to the observed sinking behavior of whole berries in sorting solutions despite the lower-density pulp comprising 85% of berry volume.
Tissue density quality is a density quality that inheres in anatomically distinct grape berry compartments and represents the mass per unit volume specific to individual tissue types including mesocarp, exocarp, and seed tissues that differ fundamentally in their cellular organization, biochemical composition, and developmental trajectories. Within the BFO framework, this quality enables ontological differentiation between density measurements at the whole-berry level and those characterizing specific anatomical structures, recognizing that composite berry density emerges from the weighted contributions of tissues with dramatically different individual densities. The ontological distinction is critical because grape berry tissues span a density range from approximately 0.95 g/mL for air-space-rich pulp tissue to over 1.20 g/mL for lignified seed coats, meaning that tissue proportions significantly influence whole-berry density independent of compositional changes within any single tissue. Tissue density reflects the integration of cell wall thickness and composition, vacuolar solute concentration, cytoplasmic density, and intercellular space volume that characterize each tissue type at specific developmental stages. Measurement approaches for tissue density require careful dissection and separation protocols, with particular attention to avoiding cellular damage that releases vacuolar contents, and may employ micro-pycnometry, density gradient centrifugation, or calculated approaches from mass and geometrically-estimated volume of dissected tissue samples. The developmental dynamics of tissue density differ among berry compartments, with seed density stabilizing early after lignification while pulp and skin densities continue changing through ripening. Understanding tissue-specific density enables prediction of berry flotation behavior, interpretation of texture measurements, and correlation of structural organization with compositional parameters.
Tissue_Density_Quality serves as the intermediate class between general Density_Quality and specific tissue type densities (Berry, Seed, Skin), establishing the ontological principle that density measurements should specify the anatomical compartment being characterized. This class aligns with histological approaches to berry quality assessment and supports interpretation of non-destructive density measurements in terms of tissue contributions.
Gewebespezifische Dichte
Kompartimentdichte
anatomical tissue density
anatomische Gewebedichte
compartment density
structural tissue compactness
strukturelle Gewebedichte
tissue-specific mass density
Huang XM, Huang HB (2001) Early post-veraison growth in grapes: evidence for a two-step mode of berry enlargement. Australian Journal of Grape and Wine Research 7(3): 132-136 Coombe BG (1992) Research on development and ripening of the grape berry. American Journal of Enology and Viticulture 43(1): 101-110
Tissue Density Quality
Tissue elasticity comparison in Cabernet Sauvignon berries reveals mesocarp elasticity of 0.35 MPa (highly deformable, turgor-dependent) versus skin elasticity of 2.8 MPa (relatively rigid, wall-determined), with the 8-fold difference explaining why berry compression primarily deforms the pulp while skin maintains shape until failure, and why skin removal reduces berry elastic recovery from 65% to 28% demonstrating the skin's dominant contribution to whole-berry elastic behavior.
Tissue elasticity quality is an elasticity quality that inheres in anatomically defined grapevine structures and represents the elastic behavior specific to individual tissue types that differ in their cellular composition, cell wall architecture, and turgor contribution to mechanical properties, determining tissue-specific deformation responses to mechanical loading. Within the BFO framework, this quality enables ontological differentiation of elasticity at the tissue level, recognizing that skin, pulp, vascular, and woody tissues exhibit fundamentally different elastic behaviors reflecting their distinct structural organization and functional requirements. The ontological significance lies in tissue-specific elasticity determining the composite mechanical behavior of organs, where the serial and parallel arrangement of tissues with different elastic properties creates complex deformation patterns under load that cannot be predicted from any single tissue measurement. Tissue elasticity reflects the integration of cellular elasticity (determined by cell wall properties and turgor) with tissue architecture (cell size, shape, packing, and intercellular spaces), where highly vacuolated parenchyma shows different elastic behavior than compact collenchyma or lignified sclerenchyma despite potentially similar cell wall composition. The developmental trajectory of tissue elasticity follows differentiation and maturation, with meristematic tissues showing high elasticity that decreases as cell walls thicken and lignify in maturing tissues, while ripening berry tissues show elasticity decline associated with cell wall disassembly rather than lignification. Measurement at tissue scale requires isolation of specific tissue layers or careful interpretation of whole-organ measurements in terms of tissue contributions, with microscale techniques such as atomic force microscopy enabling spatial mapping of elasticity across tissue boundaries. Understanding tissue-specific elasticity enables interpretation of organ mechanical properties in terms of anatomical structure, prediction of failure modes under mechanical stress, and correlation of developmental changes in elasticity with underlying cellular modifications.
Tissue_Elasticity_Quality provides the tissue-level specification of elastic behavior, serving as parent for specific tissue types (Skin, Cell Wall). This quality bridges cellular mechanical properties and organ-level behavior through tissue architecture effects. Measurement requires careful tissue isolation or deconvolution of composite measurements, with the contribution of turgor versus wall properties varying among tissue types and hydration states.
Gewebeelastizität
Kompartimentelastizität
anatomical tissue elasticity
anatomische Gewebeelastizität
compartment elastic property
gewebespezifische Elastizität
structural tissue deformability
tissue-specific elasticity
Jackman RL, Stanley DW (1995) Perspectives in the textural evaluation of plant foods. Trends in Food Science & Technology 6(6): 187-194 Gao Y et al. (2016) A generic model of transcriptional regulatory network for grape berry development. Horticulture Research 3: 16050
Tissue Elasticity Quality
Tissue firmness quality profiling in Italia table grapes distinguishes the crisp, firm pulp (3.5-4.5 N puncture force) that provides the crunchy eating texture consumers value from the relatively delicate skin (break force 0.4-0.6 N) that contributes to the variety's susceptibility to handling damage and limited storage life breeding programs targeting improved Italia types employ tissue-specific texture analysis to select progeny combining the desirable pulp crispness with enhanced skin toughness, demonstrating how tissue-level quality assessment guides genetic improvement efforts.
Tissue_Firmness_Quality encompasses the physical qualities characterizing firmness at the level of specific grape berry tissue types, recognizing that skin, pulp, and seed tissues exhibit distinct mechanical properties contributing differently to overall berry texture and processing behavior. The anatomical complexity of grape berries requires tissue-specific firmness assessment because the thin but mechanically strong skin, the soft parenchymatous pulp, and the hard lignified seeds respond differently to applied forces and contribute differently to quality outcomes. Skin firmness determines resistance to splitting, cracking, and pathogen penetration while affecting extraction kinetics during maceration. Pulp firmness governs juice release behavior, eating texture, and the internal support that maintains berry shape. Seed firmness is relatively constant (fully lignified at maturity) but seed-coat hardness affects tannin extraction during winemaking. This category subdivides into Berry_Firmness_Quality addressing the integrated whole-berry response, Pulp_Firmness_Quality capturing the internal flesh texture, and Skin_Firmness_Quality describing the mechanical properties of the berry exterior. The tissue-specific approach enables understanding of how different structural components contribute to overall quality, guiding cultivar selection and management practices to optimize specific tissue properties. Tissue firmness measurement requires appropriate test geometries that isolate the tissue of interest: whole-berry compression or puncture for integrated response excised pulp testing for flesh properties and skin tensile or puncture testing for mechanical characterization of the exterior. The mechanical interactions among tissues during whole-berry testing create complex force-deformation relationships that tissue-specific testing helps interpret.
Tissue_Firmness_Quality serves as the parent class for tissue-specific firmness qualities (Berry, Pulp, Skin), recognizing the anatomical complexity underlying overall grape texture. The class bridges whole-berry quality assessment with understanding of the structural components determining that quality. Tissue-specific testing requires appropriate methodology (sample preparation, test geometry) to isolate the tissue of interest, which may be technically challenging particularly for the thin grape skin. The mechanical interactions among tissues mean that whole-berry measurements do not simply sum tissue-specific properties.
Strukturkomponentenfestigkeit
anatomical firmness
anatomische Festigkeit
compartment firmness quality
gewebespezifische Textur
structural component firmness
tissue-specific texture
Rolle L et al. (2012) Instrumental Texture Analysis Parameters as Markers of Table-Grape and Winegrape Quality: A Review. American Journal of Enology and Viticulture 63(1): 11-28 Letaief H et al. (2008) Mechanical Behavior of Winegrapes under Compression Tests. American Journal of Enology and Viticulture 59(3): 323-329
Tissue Firmness Quality
Tissue freezing tolerance quality characterization across Vitis species and cultivars established the hardiness range available for cultivar selection in cold-climate viticulture. Laboratory freezing tests using controlled cooling and visual damage assessment (browning) or electrolyte leakage quantified LT50 (temperature causing 50% mortality) for primary buds at maximum midwinter hardiness. Vitis vinifera cultivars showed tissue freezing tolerance quality with LT50 values ranging from -20°C for cold-sensitive Merlot through -23°C for intermediate Chardonnay to -26°C for cold-hardy Riesling, establishing a 6°C range within the species. Interspecific hybrids demonstrated superior tissue freezing tolerance quality with Marquette achieving LT50 of -35°C, Frontenac at -33°C, and La Crescent at -32°C, representing 10-15°C improvement over V. vinifera reflecting introgression of cold hardiness genes from native American species. The tissue freezing tolerance quality hierarchy among tissue types showed primary buds 3-4°C less hardy than secondary buds, cane phloem 5-7°C hardier than primary buds, cane xylem 3-5°C hardier than phloem, and roots 10-15°C less hardy than equivalent above-ground tissues due to the thermal buffering normally provided by soil that limits root exposure to extreme air temperatures.
Tissue freezing tolerance quality is a cold stress indicator quality that inheres in a grapevine cultivar or clone and represents the inherent and acclimation-dependent capacity of specific tissue types to survive exposure to freezing temperatures through mechanisms including controlled extracellular ice formation with cellular dehydration tolerance, deep supercooling preventing ice formation in sensitive tissues, and cryoprotective metabolite accumulation that stabilizes cellular structures during freeze-thaw events. This quality addresses the resistance dimension of cold stress response, complementing frost damage severity quality by characterizing the underlying tolerance capacity that determines damage thresholds rather than assessing damage after it occurs. The ontological significance of tissue freezing tolerance quality lies in its role as the primary determinant of grapevine survival and productivity in cold-climate regions, with the minimum freezing tolerance achieved by critical tissues establishing the geographic and climatic limits for successful cultivation of specific cultivars. The biological basis of tissue freezing tolerance encompasses multiple mechanisms operating at cellular and tissue levels, including the controlled formation of extracellular ice that withdraws water from cells, concentrating solutes and depressing the freezing point of remaining cellular water the accumulation of cryoprotective metabolites including sugars, proline, and dehydrins that stabilize membranes and proteins during dehydration and the capacity for deep supercooling in certain tissues that prevents ice formation entirely down to the homogeneous ice nucleation temperature. The tissue-specific nature of freezing tolerance creates the characteristic hardiness hierarchy among organs, with actively growing tissues showing essentially no freezing tolerance, mature leaves tolerating only brief, mild freezing, dormant buds achieving substantial tolerance through deep supercooling, and woody tissues of canes, trunks, and roots achieving the greatest tolerance through extracellular freezing tolerance mechanisms. The dynamic nature of freezing tolerance through the seasonal cycle of acclimation and deacclimation means that tolerance measured at one time may not predict survival under cold events occurring when hardiness status differs, making the timing and depth of acclimation as important as maximum achievable hardiness for predicting cold survival.
Tissue_Freezing_Tolerance_Quality is_a Cold_Stress_Indicator_Quality, serving as parent class for Bud_Freezing_Tolerance_Quality, Shoot_Freezing_Tolerance_Quality, Trunk_Freezing_Tolerance_Quality, and Root_Freezing_Tolerance_Quality which address specific tissue types. This class addresses the underlying cold resistance capacity while child classes capture organ-specific tolerance. LT50 (lethal temperature for 50% mortality) provides standard quantification. The species/cultivar range (V. vinifera -20 to -26°C, hybrids to -35°C) establishes selection options for different climates. Dynamic acclimation status modulates realized hardiness at any given time.
Gefrier-Toleranz
Kältehärte
Kälteresistenz
cold hardiness
freezing resistance
frost tolerance
Tissue Freezing Tolerance Quality
Tissue porosity analysis of Cabernet Sauvignon berries reveals mesocarp porosity of 21.3±3.1% compared to exocarp porosity of 5.7±1.2%, with this 3.7-fold difference in void volume explaining the observed 4.2-fold difference in oxygen diffusion coefficients between tissues, supporting the skin's role as a barrier protecting pulp anthocyanins and other oxidizable compounds from atmospheric oxygen prior to crush.
Tissue porosity quality is a porosity quality that inheres in anatomically defined grape berry compartments and represents the void volume fraction specific to individual tissue types including mesocarp, exocarp, and vascular tissues that differ in their characteristic intercellular space organization reflecting tissue-specific developmental programs and functional requirements. Within the BFO framework, this quality enables differentiation of porosity measurements at the whole-berry level from those characterizing specific anatomical structures, recognizing that mesocarp, skin, and vascular tissues have fundamentally different cellular packing arrangements determining their respective void fractions. The ontological distinction is critical because tissue porosity directly affects gas diffusion pathways and mechanical properties, where high-porosity mesocarp (15-25%) provides compressibility and rapid oxygen diffusion to respiring cells, while low-porosity exocarp (3-8%) creates a diffusion barrier protecting internal tissues from environmental oxygen and pathogen entry. Tissue porosity reflects the developmental balance between cell proliferation creating new intercellular junctions with associated air spaces, cell expansion that may reduce or maintain porosity depending on whether cells expand into existing voids or generate new ones, and cell wall modification affecting the extent of middle lamella separation at cell corners where tricellular junctions create characteristic intercellular spaces. The functional significance extends to tissue porosity determining texture perception, where high-porosity tissue compresses more readily and releases cellular contents upon mastication, and affecting winemaking extraction dynamics where cellular contents diffuse from disrupted cells through the intercellular network to the surrounding liquid. Measurement at tissue scale requires careful dissection and standardization of tissue boundaries, with microscopic image analysis providing spatial resolution for porosity distribution assessment.
Tissue_Porosity_Quality provides the tissue-specific specification of void fraction, paralleling the tissue-specific density classes. This quality supports interpretation of texture measurements, gas exchange modeling, and extraction predictions at the anatomical compartment level. The class distinguishes porosity arising from intercellular spaces (apoplastic voids) from intracellular vacuolar volume which represents cellular composition rather than structural organization.
Gewebehohrlraumanteil
Gewebespezifische Porosität
Kompartimentporosität
anatomical tissue porosity
anatomische Gewebeporosität
compartment void fraction
structural tissue openness
tissue-specific air space
Huang XM, Huang HB (2001) Early post-veraison growth in grapes: evidence for a two-step mode of berry enlargement. Australian Journal of Grape and Wine Research 7(3): 132-136 Lang A, Thorpe MR (1989) Xylem, phloem and transpiration flows in a grape: application of a technique for measuring the volume of attached fruits to high resolution using Archimedes' principle. Journal of Experimental Botany 40(10): 1069-1078
Tissue Porosity Quality
Tissue temperature comparison in Napa Valley Cabernet Sauvignon during a 35°C afternoon reveals leaf temperature of 32°C (3°C below ambient due to transpirational cooling at stomatal conductance of 250 mmol/m²/s), sun-exposed berry temperature of 41°C (6°C above ambient due to radiant heating with minimal evaporative cooling), and shaded berry temperature of 36°C, demonstrating the inadequacy of air temperature alone for characterizing vine thermal status.
Tissue temperature quality is a temperature quality that inheres in anatomically defined grapevine structures and represents the thermal state specific to individual organs or tissue compartments that may differ substantially from ambient air temperature due to the interaction of radiant energy absorption, convective heat exchange, evaporative cooling, metabolic heat generation, and thermal mass effects. Within the BFO framework, this quality enables ontological differentiation between the temperature of the surrounding environment and the actual thermal conditions experienced by plant tissues, recognizing that organ-specific geometry, surface properties, transpiration rate, and exposure determine the realized tissue temperature. The ontological significance lies in tissue temperature being the physiologically relevant parameter determining reaction rates and cellular responses, where ambient temperature serves only as one input to the energy balance determining actual tissue thermal state. Different grapevine organs exhibit characteristic temperature behaviors, with leaves actively regulating temperature through stomatal control of transpiration, berries experiencing wide temperature fluctuations due to limited cooling capacity, and woody tissues showing thermal inertia dampening rapid temperature changes. Tissue temperature measurement requires direct contact sensors (thermocouples, resistance thermometers) for internal temperature or infrared detection for surface temperature, with the choice affecting interpretation since surface and internal temperatures may differ by several degrees in massive organs like berry clusters. The diurnal dynamics of tissue temperature vary by organ, with leaves showing rapid response to changing radiation and wind conditions while berries and woody tissues lag behind air temperature changes due to thermal mass. Understanding tissue-specific temperature enables prediction of developmental rates, identification of stress exposure, and interpretation of quality outcomes in terms of the thermal conditions actually experienced by quality-determining tissues.
Tissue_Temperature_Quality serves as the intermediate class between general Temperature_Quality and specific organ temperature qualities (Berry, Leaf, Canopy), establishing the principle that physiologically relevant temperature is the tissue temperature rather than ambient conditions. This class supports thermal stress assessment and developmental modeling requiring organ-specific temperature inputs. Measurement methodology (contact vs. infrared, surface vs. internal) should be specified for reproducibility.
Gewebetemperatur
Organtemperatur
Pflanzengewebetemperatur
anatomical structure temperature
anatomische Strukturtemperatur
organ temperature
plant tissue thermal state
tissue heat content
Jones HG (2014) Plants and Microclimate: A Quantitative Approach to Environmental Plant Physiology, 3rd Ed. Cambridge University Press, Cambridge, pp. 135-167 Greer DH, Weston C (2010) Heat stress affects flowering, berry growth, sugar accumulation and photosynthesis of Vitis vinifera cv. Semillon grapevines grown in a controlled environment. Functional Plant Biology 37(3): 206-214
Tissue Temperature Quality
Titratable acidity quality monitoring in Clare Valley Riesling tracks the characteristic decline from 15-20 g/L at véraison through 8-10 g/L at harvest, with the rapid malic acid respiration during warm March nights (southern hemisphere) requiring frequent sampling to capture the narrow harvest window where TA remains above 7.5 g/L while sugars reach 21-23°Brix picking decisions balance TA retention against sugar accumulation, with earlier harvest preserving the crystalline acidity prized in Clare Valley Riesling while later harvest sacrifices some acid for increased richness and alcohol potential.
Titratable_Acidity_Quality represents the chemical quality characterizing the total concentration of titratable hydrogen ions in grape juice or wine, determined by neutralization with standardized base solution to a defined endpoint. Titratable acidity (TA) measures the sum of all acid species capable of releasing protons, including tartaric, malic, citric, succinic, lactic, and acetic acids, as well as contributions from amino acids, phenolic acids, and other weakly acidic compounds. The measurement is performed by titrating a known volume of sample with standardized sodium hydroxide solution (typically 0.1 N NaOH) to an endpoint of pH 7.0 or pH 8.2 depending on the protocol, with results expressed as grams per liter of a reference acid, usually tartaric acid in European convention or sulfuric acid in some other traditions. Typical titratable acidity values at harvest range from 5-10 g/L (as tartaric acid equivalents) for wine grapes, with higher values in cool climates and lower values in warm climates. Titratable acidity provides a direct measure of the acid mass present, making it useful for estimating the amount of base needed to adjust acidity and for predicting the acid contribution to taste. The decline in titratable acidity during ripening primarily reflects malic acid respiration, with typical decreases of 2-5 g/L from véraison to harvest depending on temperature conditions. Measurement protocols specify sample preparation, titrant concentration, endpoint determination method (pH meter or indicator), and expression conventions, all of which affect reported values and must be standardized for comparability.
Titratable_Acidity_Quality specifically addresses the titration-based measurement of total acid concentration, distinct from pH_Quality which measures hydrogen ion activity. The expression convention (reference acid) must be specified because results differ by approximately 50% between tartaric acid and sulfuric acid expression. The titration endpoint affects results, with pH 7.0 capturing strong acid equivalents while pH 8.2 includes weaker acid contributions. The relationship between titratable acidity and perceived sourness is modified by pH, sugar content, and other matrix components, making TA an imperfect predictor of sensory acidity.
Gesamtsäure
Säure durch Titration
acid by titration
titrable acidity
titrierbare Gesamtsäure
titrierbare Säure
total titratable acid
OIV (2021) Compendium of International Methods of Wine and Must Analysis, Organisation Internationale de la Vigne et du Vin Amerine MA and Ough CS (1980) Methods for Analysis of Musts and Wines, John Wiley and Sons pp.: 78-95
Titratable Acidity Quality
A topping timing trial in Walla Walla Syrah compared topping response quality across four intervention dates spanning bloom through véraison. Early topping at 50% capfall produced strongest response with mean 5.8 laterals per topped shoot reaching 45 cm by harvest, requiring follow-up hedging to maintain fruit exposure. Topping at fruit set showed intermediate response with 3.2 laterals averaging 28 cm. Topping at véraison exhibited minimal topping response quality with only 1.4 short laterals averaging 8 cm that did not compromise fruit zone conditions, establishing véraison-timing as optimal for this vigorous site despite the extended period of excessive canopy height prior to intervention.
Topping response quality is a vigor manipulation response quality that inheres in a grapevine organism and represents the characteristic pattern and intensity of compensatory growth exhibited following removal of shoot apices through the topping operation, which severs the distal portion of actively growing shoots to control canopy height, remove the dominant apical meristem, and redirect growth resources toward fruit development and lateral shoot formation. This quality determines the effectiveness of topping as a vigor management tool and influences decisions regarding topping timing, severity, and the need for subsequent interventions to manage regrowth. The ontological basis of topping response quality rests on the fundamental physiological principle of apical dominance, wherein the actively growing shoot tip produces auxin that suppresses lateral bud development along the shoot axis, such that removal of this auxin source through topping releases inhibition and permits rapid lateral bud activation. The magnitude of topping response varies with several interacting factors including the vigor status of the vine at intervention, the phenological timing relative to natural growth cessation, the severity of tissue removal, and environmental conditions supporting continued growth potential. Early-season topping during active vegetative growth typically provokes strong compensatory responses with multiple vigorous lateral shoots emerging from nodes immediately below the cut, potentially creating denser canopy conditions than existed prior to intervention, while topping conducted closer to véraison when endogenous growth cessation approaches produces more modest responses. The qualitative nature of topping response includes consideration of lateral shoot number, their position along the parent shoot, growth rate and ultimate length, and the tendency for secondary laterals to develop from the primary lateral shoots in high-vigor situations. Cultivar differences in topping response quality reflect variation in apical dominance strength, bud fruitfulness of lateral positions, and intrinsic vigor, with vigorous cultivars on fertile sites often exhibiting problematic topping responses that necessitate repeated intervention or alternative management approaches. The interaction between topping response quality and fruit composition involves both direct effects through altered source-sink relationships affecting carbon allocation to berries and indirect effects through modified canopy microclimate as lateral regrowth influences fruit zone shading and air circulation. Optimal topping strategies integrate understanding of site-specific and cultivar-specific response patterns to time interventions for minimal compensatory growth while achieving necessary canopy height control.
Topping_Response_Quality is_a Vigor_Manipulation_Response_Quality, representing the specific response to shoot apex removal through topping operations. This class addresses the release of apical dominance and consequent lateral development following topping, which represents one of the most common canopy management interventions. Distinguished from Hedging_Response_Quality which involves lateral shoot removal along with apex removal in a single pass. The timing relative to phenological stage critically determines response magnitude and should be emphasized in annotation applications.
Entspitzungsreaktion
Gipfelschnitt-Reaktion
Gipfelungsreaktion
Triebspitzenentfernung-Antwort
apex removal response
shoot topping response
summer topping regrowth
topping compensation
Intrieri C, Poni S (1995) Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality of mechanized Italian vineyards. American Journal of Enology and Viticulture 46:116-127 Keller M et al. (1999) Ripening grape berries remain hydraulically connected to the shoot. Journal of Experimental Botany 50:1575-1581 Poni S, Giachino E (2000) Growth, photosynthesis and cropping of potted grapevines (Vitis vinifera L. cv. Cabernet Sauvignon) in relation to shoot trimming. Australian Journal of Grape and Wine Research 6:216-226
Topping Response Quality
Syrah in organic vineyard showing topsoil surface lateral roots, soil profile examination revealing laterals concentrated 12-24 cm depth in A horizon (0-28 cm) with sharp density decline at A-B boundary, topsoil laterals 1.4-3.6 cm diameter, topsoil zone showing 6.8% organic matter versus 1.2% in B horizon, topsoil lateral-associated fine roots providing estimated 68% of nitrogen uptake and 74% of phosphorus uptake, topsoil surface lateral roots demonstrating specialization for exploiting nutrient-rich A horizon resources
Topsoil surface lateral root is surface lateral root emphasizing topsoil zone association and pedological context, characterized by position in A horizon soil layer typically 10-25 cm depth where organic matter accumulation, biological activity, and nutrient availability highest, showing morphological and functional adaptations to topsoil environment, demonstrating preferential development in high-quality topsoil versus poor subsoil, and serving specialized function exploiting topsoil resources supporting disproportionate vine uptake relative to root biomass investment, with topsoil surface laterals showing characteristics including concentration in A horizon organic-rich layer, proliferation responding to topsoil fertility with higher density in fertile than poor topsoils, extensive fine root and root hair production maximizing topsoil resource capture, high arbuscular mycorrhizal colonization enhancing nutrient acquisition particularly phosphorus often limiting in topsoils, and demonstrated importance through studies showing topsoil lateral-associated roots providing 50-75% of total vine nitrogen and phosphorus uptake despite occupying small soil volume. Topsoil surface lateral root pedological context includes= showing A horizon association (topsoil laterals concentrated in A horizon pedogenic layer characterized by organic matter accumulation), exhibiting topsoil preference (topsoil laterals showing preferential development in A horizon versus B or C horizons), demonstrating horizon boundary correlation (topsoil lateral density often declining sharply at A-B horizon boundary reflecting topsoil specialization), revealing structure benefits (topsoil laterals benefiting from favorable A horizon structure including good aggregation and porosity), and displaying organic matter effects (topsoil lateral function enhanced by organic matter improving moisture retention, nutrient availability, biological activity). Topsoil surface lateral root nutrient availability includes= showing nitrogen concentration (topsoil zone often showing highest nitrogen availability from organic matter mineralization), exhibiting phosphorus enrichment (phosphorus often concentrated in topsoil through accumulation and fertilization), demonstrating cation exchange (topsoil showing higher CEC from organic matter and clay providing nutrient retention), revealing micronutrient availability (topsoil often showing better micronutrient availability through chelation and favorable pH), and displaying nutrient cycling (topsoil zone showing most active nutrient cycling through decomposition and mineralization). Topsoil surface lateral root biological interactions includes= showing maximum microbial diversity (topsoil supporting most diverse microbial communities enhancing nutrient cycling), exhibiting earthworm activity (topsoil zone showing earthworm activity improving structure and organic matter distribution), demonstrating mycorrhizal networks (topsoil laterals interconnected through arbuscular mycorrhizal hyphal networks), revealing nematode interactions (topsoil supporting diverse nematode communities potentially affecting root function), and displaying microfaunal complexity (topsoil showing complex microfaunal community affecting nutrient availability and root health). Topsoil surface lateral root absorptive specialization includes= providing disproportionate uptake (topsoil laterals supplying 50-75% of vine nitrogen and phosphorus despite small biomass fraction), exhibiting nutrient-specific importance (topsoil laterals particularly critical for nitrogen and phosphorus acquisition from topsoil enrichment), demonstrating mycorrhizal dependence (topsoil lateral function often critically dependent on arbuscular mycorrhizae enhancing phosphorus uptake), revealing seasonal importance (topsoil laterals particularly important spring and fall when topsoil conditions optimal), and displaying efficiency (topsoil laterals showing high uptake efficiency per unit biomass through favorable topsoil environment). Topsoil surface lateral root in soil quality indication includes= indicating topsoil health (abundant vigorous topsoil laterals indicating good topsoil quality and health), demonstrating degradation sensitivity (topsoil lateral decline potentially indicating soil degradation through erosion, compaction, or fertility depletion), revealing restoration response (topsoil laterals potentially showing recovery following soil quality improvement through organic amendments or reduced tillage), showing management effects (topsoil lateral characteristics reflecting soil management practices affecting topsoil quality), and displaying diagnostic value (topsoil lateral assessment potentially informing soil management needs and effectiveness). Topsoil surface lateral root vulnerability to degradation includes= showing erosion susceptibility (soil erosion removing or damaging topsoil laterals particularly on slopes), exhibiting compaction sensitivity (topsoil laterals readily damaged by compaction from traffic or equipment), demonstrating tillage impact (repeated cultivation degrading topsoil structure and damaging topsoil laterals), revealing organic matter dependence (topsoil lateral function declining if organic matter depleted through intensive management), and displaying restoration needs (damaged topsoil lateral populations potentially requiring years to recover following degradation). Topsoil surface lateral root conservation practices includes= requiring erosion control (preventing topsoil erosion maintaining topsoil laterals and soil quality), showing organic matter maintenance (maintaining or building topsoil organic matter supporting topsoil lateral function), exhibiting tillage reduction benefits (reducing or eliminating tillage protecting topsoil laterals and improving structure), demonstrating cover crop advantages (cover crops protecting topsoil adding organic matter benefiting topsoil laterals though potentially creating competition), and revealing mulch value (organic mulches protecting topsoil from erosion and temperature extremes while adding organic matter). Topsoil surface lateral root in sustainable viticulture includes= determining long-term productivity (maintaining healthy topsoil lateral populations essential for sustainable vine productivity), enabling resource efficiency (abundant topsoil laterals improving fertilizer and water use efficiency reducing environmental impacts), demonstrating soil health integration (topsoil lateral abundance integrating with overall soil health supporting sustainable production), revealing regenerative potential (topsoil lateral recovery following improved management indicating regenerative agriculture success), and displaying ecosystem services (healthy topsoil lateral-soil system providing multiple ecosystem services including carbon sequestration and biodiversity support).
Topsoil_Surface_Lateral_Root emphasizes A horizon pedological association and is distinguished from other surface roots by focus on topsoil layer context. Critical for understanding pedological context, nutrient enrichment, biological interactions, disproportionate uptake, soil quality indication, and the laterals specialized for exploiting A horizon resources supporting sustainable productivity.
A horizon lateral
A-Horizont-Lateralwurzel
Oberboden-Oberflächenwurzel
Organic-rich zone root
Organik-reiche-Zonen-Wurzel
Topsoil-zone lateral
Topsoil Surface Lateral Root
Total acidity for Sauvignon Blanc from Marlborough (2024 vintage) measures 9.5 g/L as tartaric acid (pH endpoint 8.2), placing it within the characteristically high-acid profile (8.0-11.0 g/L) that defines the region's signature crisp, vibrant wine style. Total acidity for Tempranillo from Ribera del Duero (2023 vintage) records 5.2 g/L as tartaric acid, near the lower threshold for balanced dry red wine production (generally 5.0-7.0 g/L for reds), with the winemaker's decision to forego malolactic fermentation for a portion of the lot contingent on this moderate acid level providing sufficient freshness without further acid reduction. Total acidity for Zweigelt from Neusiedlersee (2024 vintage) documents 6.0 g/L as tartaric acid at 18° KMW harvest, meeting the preferred range for Austrian dry red wine production and indicating no need for either acidification or deacidification.
Total acidity is an acidity data entity that records the titratable acidity (TA) of grape must, determined by neutralization titration with standardized sodium hydroxide solution to a defined pH endpoint (pH 7.0 per OIV method, pH 8.2 per some national standards), expressed in grams per liter as tartaric acid equivalents (g/L H₂T) or, in some European conventions, as sulfuric acid equivalents (g/L H₂SO₄), representing the aggregate concentration of all proton-donating species in the must and serving as the primary bulk acidity metric for harvest decisions, regulatory compliance, and winemaking acid adjustment calculations.
Total acidity measurement by titration is the most universally performed acid analysis in grape and wine laboratories, yet its interpretation requires careful attention to methodological conventions that vary between regions. The OIV reference method specifies titration to pH 7.0 with phenolphthalein or bromothymol blue indicator, while some national standards (notably Australian and New Zealand) titrate to pH 8.2 — the higher endpoint captures additional buffering capacity from phosphates and amino acids, yielding TA values approximately 0.5-1.0 g/L higher than the pH 7.0 endpoint on identical samples. Expression conventions add further complexity — most New World and many European laboratories report TA as tartaric acid equivalents (conversion factor 0.075 g/meq), while French and some Mediterranean traditions historically report as sulfuric acid equivalents (conversion factor 0.049 g/meq), requiring multiplication by 1.53 for conversion from H₂SO₄ to H₂T equivalents. The relationship between TA and sensory acid perception is mediated by pH, sugar content, and buffering capacity — a wine with TA 7.0 g/L at pH 3.10 tastes substantially more acidic than one with identical TA at pH 3.50 due to the higher free hydrogen ion concentration, while residual sugar masks acid perception through a sweetness-acidity sensory interaction. FTIR (Fourier-transform infrared) spectroscopy enables rapid TA estimation within ±0.3 g/L accuracy, sufficient for screening purposes but requiring confirmation by reference titration for regulatory and commercial transactions.
Gesamtsäuregehalt
Mostsäure
acid concentration
must acidity
titratable acidity
total acid content
OIV (2021) Compendium of International Methods of Analysis, Method OIV-MA-AS313-01 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2, 2nd Ed, Wiley, pp.: 3-12 Iland P et al. (2011) Chemical Analysis of Grapes and Wine, 2nd Ed, pp.: 55-60
Total Acidity
Total acidity quality assessment in Margaret River Chardonnay employs dual measurement of titratable acidity (target 6.5-7.5 g/L) and pH (target 3.25-3.40) to guide harvest timing and stylistic decisions fruit with titratable acidity above 8 g/L and pH below 3.20 is directed toward sparkling wine production, while fruit in the optimal range supports premium still wine, and fruit with titratable acidity below 6 g/L or pH above 3.50 may require acid adjustment or blending to achieve balanced wines, demonstrating how total acidity parameters inform winery allocation decisions.
Total_Acidity_Quality encompasses the chemical qualities that characterize the aggregate acid content of grape juice and wine through composite measurements that integrate contributions from all acidic species present. This category provides the practical acidity metrics used in routine quality assessment without distinguishing individual acid contributions, capturing the overall acid status relevant to taste perception and chemical stability. Total acidity can be expressed through two fundamentally different but complementary measurements: Titratable_Acidity_Quality representing the total concentration of titratable hydrogen ions from all acid sources, and pH_Quality representing the activity of free hydrogen ions in solution. These measurements provide different information because grape juice and wine contain weak acids that are only partially dissociated, creating a distinction between total acid concentration and hydrogen ion activity. The relationship between titratable acidity and pH depends on the specific acids present, their pKa values, and the buffering capacity of the matrix. Two musts with identical titratable acidity may have different pH values if their acid composition differs, because tartaric acid (pKa₁ = 2.98) releases more hydrogen ions at wine pH than malic acid (pKa₁ = 3.40). Total acidity measurements guide harvest decisions, blending strategies, and acidification or deacidification treatments. Regulatory frameworks often specify both titratable acidity and pH requirements for quality wines, recognizing that both parameters contribute to quality perception and wine stability. Total acidity assessment employs standardized titration protocols for titratable acidity and calibrated electrodes for pH measurement.
Total_Acidity_Quality serves as the parent class for the two primary total acidity measurements, titratable acidity and pH. These measurements are not interchangeable both are needed for complete acidity characterization because they capture different aspects of acid chemistry. The buffering capacity of grape juice and wine, determined by the weak acid equilibria and protein/polyphenol contributions, creates the complex relationship between titratable acidity and pH. Total acidity qualities are distinguished from individual acid species qualities addressed under Organic_Acid_Quality.
Gesamtsäurestatus
aggregate acidity
aggregierte Säure
combined acid measurement
gesamter Säuregehalt
overall acid content
total acid status
Boulton RB et al. (1999) Principles and Practices of Winemaking, Springer pp.: 65-78 Iland P et al. (2004) Chemical Analysis of Grapes and Wine: Techniques and Concepts, Patrick Iland Wine Promotions pp.: 57-75
Total Acidity Quality
Total nitrogen quality assessment in Napa Valley Chardonnay correlates with vineyard vigor zones mapped through remote sensing, with high-vigor areas producing must with 400-600 mg N/L total nitrogen (of which 200-350 mg N/L is YAN) that ferments rapidly but may produce excessive vegetative character, while moderate-vigor zones yield 250-400 mg N/L total nitrogen (150-250 mg N/L YAN) supporting balanced fermentation and fruit expression this total nitrogen mapping enables precision viticulture approaches to nitrogen management and selective harvesting for quality optimization.
Total_Nitrogen_Quality represents the chemical quality characterizing the aggregate concentration of all nitrogen-containing compounds in grape berry tissue, providing a comprehensive measure of the nitrogen pool without distinguishing among forms or availability for fermentation. Total nitrogen includes both organic nitrogen (amino acids, peptides, proteins, nucleic acids, alkaloids) and inorganic nitrogen (ammonium, nitrate), capturing the complete nitrogenous composition regardless of chemical form or biological accessibility. In grape must, total nitrogen typically ranges from 100-600 mg N/L, with the wide variation reflecting cultivar genetics, vine nitrogen status, soil fertility, and seasonal conditions. Total nitrogen correlates with vine vigor and nitrogen nutrition status, making it useful for vineyard management decisions regarding fertilization and canopy balance. However, total nitrogen incompletely predicts fermentation performance because substantial fractions (proteins, proline in most species) are not accessible to wine yeast during fermentation. The relationship between total nitrogen and yeast assimilable nitrogen (YAN) varies among samples depending on the proportion of non-assimilable forms, making total nitrogen a less precise fermentation indicator than YAN. Total nitrogen measurement employs combustion methods (Dumas) or acid digestion (Kjeldahl) that convert all nitrogen forms to measurable species, providing comprehensive but non-specific quantification. The total nitrogen concentration establishes an upper limit for potential YAN but does not directly predict yeast-available nitrogen without knowledge of the compositional profile. Total nitrogen content affects wine protein stability (through protein nitrogen), potential ethyl carbamate precursors (through arginine and urea), and general indication of vine nutritional status.
Total_Nitrogen_Quality addresses the complete nitrogen pool as a compositional parameter distinct from the fermentation-relevant YAN fraction. The inclusion of non-assimilable nitrogen (proline, proteins) in total nitrogen creates the discrepancy between total nitrogen and fermentation prediction that motivated YAN development. Total nitrogen remains valuable for vine nutritional assessment, protein stability prediction, and general quality characterization even though YAN provides superior fermentation guidance. The analytical distinction between Kjeldahl (digestion) and Dumas (combustion) methods affects results for certain nitrogen forms.
Gesamt-N-Qualität
Gesamtstickstoffgehalt
aggregate nitrogen content
complete nitrogen pool
comprehensive nitrogen content
total N quality
umfassender Stickstoffgehalt
vollständiger Stickstoffpool
Butzke CE (1998) Survey of Yeast Assimilable Nitrogen Status in Musts from California, Oregon, and Washington. American Journal of Enology and Viticulture 49(2): 220-224 Bell SJ and Henschke PA (2005) Implications of Nitrogen Nutrition for Grapes, Fermentation and Wine. Australian Journal of Grape and Wine Research 11(3): 242-295
Total Nitrogen Quality
Total phenolic quality monitoring in Ribera del Duero Tempranillo tracks the accumulation from 1200-1500 mg/L GAE at véraison through 2200-2800 mg/L GAE at optimal harvest, with the rate of phenolic increase providing ripening trajectory information complementary to sugar accumulation winemakers target total phenolic values above 2400 mg/L GAE for premium Reserva and Gran Reserva wines requiring extended oak aging, as higher initial phenolic content provides the structural backbone to integrate wood tannins while maintaining color stability through years of bottle maturation.
Total_Phenolic_Quality represents the chemical quality characterizing the aggregate concentration of all phenolic compounds in grape tissue or wine, providing a composite measure of phenolic content without distinguishing individual compound classes. Total phenolic measurement captures the combined contribution of anthocyanins, flavonols, flavan-3-ols, tannins, hydroxycinnamic acids, stilbenes, and other phenolic species through colorimetric assays that respond to the reducing capacity of phenolic hydroxyl groups. The Folin-Ciocalteu assay, the most widely used method, employs a phosphomolybdic-phosphotungstic acid reagent that is reduced by phenolic compounds to produce a blue color measured spectrophotometrically, with results expressed as gallic acid equivalents (GAE) or catechin equivalents depending on the calibration standard. Total phenolic values in grape juice and wine vary enormously with cultivar, vintage, and winemaking, ranging from 200-400 mg/L GAE in white wines to 1500-3500 mg/L GAE in full-bodied red wines. In grape berries, total phenolic concentration increases during ripening as anthocyanins and other phenolics accumulate, providing a maturity indicator that complements sugar and acid measurements. Total phenolic content correlates with wine color intensity, antioxidant capacity, and general perception of body and structure, though the relationship with specific sensory attributes requires individual compound measurement. The simplicity and rapidity of total phenolic assays make them useful for routine quality monitoring and screening large sample sets, while their lack of specificity limits interpretation of compositional detail. Total phenolic measurements may include contributions from non-phenolic reducing compounds, requiring awareness of potential interferences.
Total_Phenolic_Quality provides an aggregate measure positioned as a direct subclass of Phenolic_Content_Quality. The Folin-Ciocalteu method, while standard, is not specific for phenolics and responds to any reducing compound, potentially including ascorbic acid, sulfur dioxide, and sugars careful sample preparation minimizes interferences. Alternative total phenolic methods include spectrophotometric estimation at 280 nm and various colorimetric assays with different specificity profiles. Total phenolic values enable comparison across samples but do not predict specific sensory properties that depend on the compound profile within the total. The expression units (GAE, catechin equivalents) affect numerical values consistent specification is essential for comparability.
Folin-Ciocalteu phenolics
Gesamtphenolgehalt
Gesamtphenolindex
Gesamtpolyphenole
aggregate phenolic content
aggregierter Phenolgehalt
total phenolic index
total polyphenols
Singleton VL and Rossi JA (1965) Colorimetry of Total Phenolics with Phosphomolybdic-Phosphotungstic Acid Reagents. American Journal of Enology and Viticulture 16(3): 144-158 Waterhouse AL (2002) Determination of Total Phenolics. Current Protocols in Food Analytical Chemistry 6(1): I1.1.1-I1.1.8
Total Phenolic Quality
Total phenolics for Shiraz from the Barossa Valley (2023 vintage) measures 78 AU280 by the Iland berry homogenate method, exceeding the premium threshold of 60 AU typically associated with concentrated, well-structured red wines from this region, and confirming the advanced phenolic development expected from old-vine (80+ year) dry-grown material harvested at 14.5° Baumé. Total phenolics for Pinot Noir from Burgundy Premier Cru (2024 vintage) records 42 AU280, within the typical cultivar range for Pinot Noir (35-55 AU) which is inherently lower than thick-skinned varieties, reflecting the thin skin and lower tannin content that define Pinot Noir's characteristically silky texture. Total phenolics for Zweigelt from Neusiedlersee (2024 vintage) documents 52 AU280 at 19° KMW, consistent with the variety's moderate phenolic potential and suitable for the medium-bodied, fruit-forward wine style typical of the region.
Total phenolics is a phenolic data entity that records the aggregate concentration of all phenolic compounds in grape must or berry homogenate, determined by ultraviolet spectrophotometric absorbance at 280 nm (A280, Iland method), Folin-Ciocalteu colorimetric assay, or total polyphenol index measurement, expressed in absorbance units (AU) at 280 nm or in milligrams per liter gallic acid equivalents (mg/L GAE), representing the sum of all hydroxyphenyl-containing molecules including anthocyanins, flavanols (catechins, proanthocyanidins/tannins), flavonols, hydroxycinnamic acids, and stilbenes, and serving as the broadest screening metric for phenolic ripeness and red wine extraction potential.
Total phenolics provides the most accessible and widely performed phenolic screening measurement, useful for rapid assessment but inherently limited in specificity because the UV absorbance at 280 nm (or the Folin-Ciocalteu reaction) responds to all aromatic ring-containing compounds without distinguishing between anthocyanins, tannins, flavonols, and non-flavonoid phenolics. The Iland method (berry homogenate absorbance at 280 nm after 1 hour extraction in 50% ethanol at pH 2.0) has become the Australian and New World standard for vineyard phenolic assessment, providing a rapid, reproducible metric that correlates well with wine phenolic concentration. The Folin-Ciocalteu assay, though widely used in food science, suffers from interference by non-phenolic reducing substances (ascorbic acid, sugars, sulfur dioxide) that can overestimate phenolic content in must matrices. Reference ranges are cultivar-dependent — Cabernet Sauvignon typically achieves 60-90 AU280, Shiraz 55-85 AU, Pinot Noir 35-55 AU, and Merlot 45-65 AU — reflecting genetic differences in skin thickness, tannin content, and anthocyanin accumulation that define each cultivar's phenolic identity. The temporal trend of total phenolics during ripening typically shows a plateau or slight decline in the final pre-harvest weeks as anthocyanin degradation and tannin polymerization offset continued biosynthesis, meaning that maximum total phenolics may precede optimal sugar maturity by 1-2 weeks in warm vintages. Total phenolics should always be interpreted alongside the specific phenolic fractions (anthocyanins, tannin) and sugar content, as high total phenolics at low sugar may indicate green, astringent tannin extraction potential while moderate total phenolics at appropriate sugar may indicate better phenolic maturity with softer, more polymerized tannin structures.
Polyphenolindex
TPI
phenolischer Gesamtgehalt
polyphenol index
total phenolic content
Iland P et al. (2011) Chemical Analysis of Grapes and Wine, 2nd Ed, pp.: 88-100 Singleton VL, Rossi JA (1965) Colorimetry of total phenolics with phosphomolybdic-phosphotungstic acid reagents. American Journal of Enology and Viticulture 16(3): 144-158 Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2, 2nd Ed, Wiley, pp.: 141-160
Total Phenolics
Total radiation for the Barossa Valley reaches 6500-7000 MJ/m² annually with growing season (October-April) totals of 4500-5000 MJ/m², supporting full phenolic ripeness in Shiraz with typical harvest Baumé of 13.5-14.5°. Champagne receives approximately 3800-4200 MJ/m² annually, near the minimum threshold for reliable Vitis vinifera ripening, with growing season radiation of 2800-3200 MJ/m² that yields the characteristically high acidity and moderate sugar levels essential for sparkling wine production. The Clare Valley records 6800 MJ/m² annual total radiation, yet the 400-500 m elevation moderates nocturnal temperatures, creating a diurnal decoupling between high radiation input and cool night temperatures that preserves acidity in Riesling.
Total radiation is a solar radiation data entity that records the cumulative global solar radiation (direct beam plus diffuse sky radiation) received on a horizontal surface at a viticultural site, typically expressed in megajoules per square meter (MJ/m²) for annual, seasonal, or monthly intervals, or in watts per square meter (W/m²) as instantaneous flux, providing the fundamental energy budget metric for vine photosynthetic potential and thermal accumulation.
Total radiation measurement on a horizontal surface provides the standard meteorological reference value but underestimates energy receipt on sloped vineyard surfaces, particularly at higher latitudes where low solar angles increase the radiation advantage of south-facing (Northern Hemisphere) or north-facing (Southern Hemisphere) slopes. Correction factors for slope and aspect can increase effective radiation by 20-50% on steep slopes compared to horizontal measurements. The ratio of direct to diffuse radiation varies with cloud cover and atmospheric conditions — overcast conditions increase the diffuse fraction but reduce total radiation, while high-altitude sites with clear skies receive proportionally more direct radiation. Modern vineyard-specific radiation modeling increasingly employs fisheye photography, LiDAR canopy scanning, and hemispherical radiation sensors to characterize the three-dimensional radiation environment within the canopy volume.
Gesamtsonnenstrahlung
Gesamtstrahlung
Globalstrahlung
Sonneneinstrahlung gesamt
cumulative solar radiation
global radiation
global solar radiation
total insolation
total solar irradiance
Smart RE, Robinson MD (1991) Sunlight into Wine, Winetitles, pp.: 7-18 Gladstones J (2011) Wine, Terroir and Climate Change, Wakefield Press, pp.: 73-86 Prichard TL et al. (2004) Radiation balance and energy budget of a vineyard canopy. Agricultural and Forest Meteorology 127(1-2): 93-113
Total Radiation
Total SO₂ for Riesling trocken from the Mosel (2024 vintage) measures 85 mg/L (free SO₂ 32 mg/L, bound SO₂ 53 mg/L), well below the EU maximum of 200 mg/L for dry white wines with 5 g/L residual sugar, with the high bound SO₂ fraction attributable to the elevated aldehyde and ketone content of botrytized wine that avidly binds bisulfite. Total SO₂ for Blaufränkisch from Mittelburgenland (2023 vintage, dry red) documents 92 mg/L, below the EU maximum of 150 mg/L for dry red wines.
Total sulfur dioxide is a wine analysis data entity that records the combined concentration of all forms of sulfur dioxide present in finished wine — comprising free SO₂ (molecular, bisulfite, and sulfite forms) plus bound SO₂ (SO₂ irreversibly or reversibly complexed with acetaldehyde, pyruvic acid, α-ketoglutaric acid, glucose, anthocyanins, and other carbonyl compounds) — determined by aspiration-oxidation with alkaline hydrolysis, iodometric titration after alkaline release, or flow injection analysis, expressed in milligrams per liter (mg/L), and representing both the total sulfur dioxide input to the wine and the regulatory compliance metric against which maximum permitted SO₂ levels are enforced for labeling and trade purposes.
Total sulfur dioxide is the principal regulatory SO₂ metric because it represents the cumulative SO₂ exposure of the wine consumer regardless of chemical form, and it is on this basis that maximum permitted levels and labeling requirements are established. The EU regulatory framework (Regulation 2019/934) sets maximum total SO₂ limits that vary by wine category and residual sugar content: dry red wines ≤150 mg/L, dry white and rosé wines ≤200 mg/L, wines with >5 g/L residual sugar receive additional allowances (+50 mg/L), sweet wines with >5 g/L RS ≤300 mg/L for reds and ≤400 mg/L for whites, with organic wine regulations imposing lower limits (typically 100 mg/L for dry reds, 150 mg/L for dry whites). The allergen labeling requirement "contains sulfites" is mandatory in most jurisdictions when total SO₂ exceeds 10 mg/L — effectively mandatory for all conventionally produced wines, as even unsulfited wines typically contain 10-30 mg/L SO₂ from yeast metabolism during fermentation. The bound SO₂ fraction, while antimicrobially inactive, reflects the wine's carbonyl compound content — high bound SO₂ indicates elevated acetaldehyde (from oxidative exposure or Sherry-type flor aging), pyruvic acid (from early-stage fermentation), or glucose-bisulfite adducts (in sweet wines), providing indirect diagnostic information about wine production history and stability status. The ratio of free to total SO₂ provides a measure of SO₂ utilization efficiency — wines with low free-to-total ratios (0.3) use SO₂ more efficiently. The progressive reduction of total SO₂ levels across the global wine industry — driven by consumer health awareness, organic certification requirements, and improved cellar hygiene reducing the need for heavy sulfiting — represents a significant trend in contemporary winemaking practice.
Gesamt-SO₂
Gesamtschwefeldioxid
SO₂-Gesamtgehalt
combined SO2
gesamte schweflige Säure
gesamtes Schwefeldioxid
total SO2
total SO2 level
total sulfite
total sulfur dioxide content
OIV (2021) Compendium of International Methods of Analysis, Method OIV-MA-AS323-04A (total sulfur dioxide) European Parliament (2019) Regulation (EU) 2019/934, Annex I Part B (maximum SO₂ limits) Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2, 2nd Ed, Wiley, pp.: 223-250 Waterhouse AL et al. (2016) Understanding Wine Chemistry, Wiley, pp.: 178-210
Total SO2
Total sugar quality specification for Sauternes production requires natural sugar concentrations of 300-400 g/L in botrytized Sémillon berries, achieved through Botrytis cinerea infection that permeabilizes the skin and concentrates sugars through evaporative water loss the extreme total sugar content translates to potential alcohol of 18-24% volume, but fermentation arrests naturally at 13-14% actual alcohol leaving 100-150 g/L residual sugar that defines the luscious sweetness of great Sauternes, demonstrating how total sugar quality requirements vary radically among wine styles.
Total_Sugar_Quality represents the chemical quality characterizing the aggregate concentration of all soluble sugars in grape berry tissue, combining reducing sugars (glucose and fructose) with non-reducing sugars (primarily sucrose) into a single composite measure. Total sugar provides the most practically relevant sugar metric for harvest decisions and wine production planning because it captures the complete fermentable substrate regardless of the proportion contributed by individual sugar species. In ripe Vitis vinifera berries, total sugar is dominated by glucose and fructose in approximately equal proportions, with sucrose typically comprising less than 1% of total sugars because invertase activity rapidly cleaves transported sucrose upon arrival in the berry. However, in some cultivars, growth stages, and conditions, sucrose can comprise a more significant fraction, making total sugar measurement more comprehensive than reducing sugar alone. Total sugar concentration at harvest typically ranges from 180-260 g/L for wine grapes (corresponding to approximately 18-26°Brix), with lower values for sparkling wine base production and higher values for rich, full-bodied styles. Table grapes are harvested at lower sugar concentrations (14-18°Brix) that provide pleasant sweetness without excessive intensity. Total sugar measurement by refractometry actually estimates soluble solids including minor non-sugar components, providing a practical approximation of true total sugar. More precise determination employs enzymatic assays that specifically quantify glucose, fructose, and sucrose, with summation providing true total sugar content. The relationship between total sugar and wine quality is non-linear, with optimal concentrations depending on wine style insufficient sugar produces thin, unbalanced wines while excessive sugar can result in alcoholic heat and loss of varietal character.
Total_Sugar_Quality provides the comprehensive sugar metric that underlies practical harvest maturity assessment. The class relates to but is distinct from °Brix and other must weight scales, which estimate total sugar through physical properties (refractive index, density) rather than measuring sugar directly. The dominance of glucose and fructose in ripe berries means that total sugar and reducing sugar converge for practical purposes in mature fruit, though they may differ significantly earlier in development or in unusual conditions. Total sugar should be distinguished from total carbohydrates, which would include insoluble polysaccharides such as cell wall components total sugar specifically addresses soluble sugars.
Gesamtkohlenhydratkonzentration
Gesamtzucker
Gesamtzuckergehalt
aggregate sugar content
aggregierter Zuckergehalt
combined sugar concentration
total carbohydrate quality
total soluble sugars
Conde C et al. (2007) Biochemical Changes throughout Grape Berry Development and Fruit and Wine Quality. Food 1(1): 1-22 Jackson RS (2020) Wine Science: Principles and Applications, 5th Ed, Academic Press pp.: 347-374
Total Sugar Quality
The trade name Appassimento designates a wine production style (dried-grape winemaking) rather than a cultivar but functions commercially as a product designation closely associated with specific cultivar selections — more directly, the trade name Enantio was adopted for the cultivar with VIVC prime name Lambrusco a Foglia Frastagliata in the Valdadige region to create a more marketable and distinctive commercial identity, while ENTAV-INRA markets its certified clonal material under the trade name Entav-Inra with specific clone numbers (e.g., ENTAV-INRA 115 for Pinot Noir) that function as de facto trade names in the international nursery market.
A cultivar designation created or adopted for commercial purposes within the wine and nursery trade, functioning as a market-facing identifier that may be legally protected through trademark registration and is distinct from the cultivar's scientific prime name, breeding code, or regulatory designation.
Trade_Name captures the commercial layer of cultivar nomenclature where naming decisions are driven by market positioning, brand building, and intellectual property strategy rather than by scientific or regulatory conventions. Trade names may be registered as trademarks, granting the registrant exclusive commercial use within specified product categories and jurisdictions: this legal protection distinguishes trade names from other cultivar designations that are part of the public nomenclatural commons. Subclasses distinguish brand names (proprietary designations owned by specific entities), marketing names (commercially oriented but not necessarily trademark-protected), and protected trade names (formally registered intellectual property).
Handelsbezeichnung
commercial name
kommerzieller Sortenname
market name
trade designation
ICNCP (2016) International Code of Nomenclature for Cultivated Plants 9th Ed, ISHS Art. 11 UPOV (2020) International Convention for the Protection of New Varieties of Plants Robinson J et al. (2012) Wine Grapes, Allen Lane pp: 1-18
Trade Name
Trade regulation for Austrian wine exports to the United States integrates multiple regulatory layers: EU export certification (conformity with EU wine production standards, export licence from the Austrian Bundeskellereiinspektion), US import requirements (TTB Certificate of Label Approval/COLA for each label variant, compliance with US labeling regulations including the mandatory Government Warning statement, FDA Prior Notice of imported food shipments), bilateral trade agreement provisions (the 2006 EU-US Wine Agreement establishing mutual acceptance of certain oenological practices and protecting certain geographic indications), and customs classification (Combined Nomenclature codes determining tariff rates). Trade regulation for grapevine plant material export from France to Australia involves stringent phytosanitary requirements — NPPO (National Plant Protection Organization) phytosanitary certificate, mandatory virus testing (ELISA and PCR for Grapevine leafroll-associated viruses, Grapevine fanleaf virus, and other regulated pathogens), post-entry quarantine in Australia (minimum 2 years at an approved quarantine facility), and compliance with Australia's exceptionally strict Biosecurity Act 2015 provisions designed to protect the Australian vineyard from exotic pests and diseases.
Trade regulation is a regulatory information entity that documents the legal frameworks governing the international and domestic commercial movement of grape plant material (cuttings, rootstocks, bench grafts, potted vines) and wine products across jurisdictional boundaries, encompassing export certification requirements (phytosanitary certificates, varietal authenticity documentation, quality compliance attestation), import requirements (quarantine protocols, pest risk analysis, documentation verification), quarantine status designations (the phytosanitary risk classification determining the stringency of import controls), and trade agreement provisions (bilateral and multilateral treaties governing wine and plant material trade), providing the commercial-regulatory framework that simultaneously facilitates legitimate viticultural trade and protects national plant health and consumer interests from the risks associated with cross-border movement of biological material and agricultural products.
Trade regulation documentation addresses the complex intersection of viticultural commerce, phytosanitary protection, and international trade law that governs the cross-border movement of both wine products and grapevine plant material — two fundamentally different regulatory domains united by their viticultural subject matter but governed by distinct legal frameworks and institutional structures. Wine trade regulation operates primarily through food safety and labeling compliance frameworks — ensuring that imported wines meet the destination country's production standards, labeling requirements, and food safety criteria — while plant material trade regulation operates through phytosanitary frameworks designed to prevent the introduction of plant pests and diseases across national boundaries, governed by the International Plant Protection Convention (IPPC) and its regional organizations. The economic significance of wine trade regulation is substantial — global wine trade exceeded €36 billion in 2022 (OIV data), with the EU accounting for approximately 60% of global exports by value, and regulatory trade barriers (tariffs, non-tariff barriers including labeling requirements, oenological practice restrictions, and administrative procedures) significantly affecting trade flows and competitive dynamics. The phytosanitary dimension of plant material trade regulation carries existential importance for viticulture — the historical introductions of powdery mildew (1850s), downy mildew (1870s), and phylloxera (1860s) from North America to Europe, each devastating the European viticultural sector, provide the historical justification for the stringent quarantine and certification regimes that now govern international vine material movement. The tension between trade facilitation (reducing regulatory barriers to legitimate commerce) and phytosanitary protection (maintaining biosecurity against exotic pest and disease introduction) is a persistent policy challenge that different jurisdictions resolve differently — with Australia and New Zealand maintaining the strictest import quarantine regimes globally, reflecting their geographic isolation and the exceptional phytosanitary status of their vineyards.
Handelsrecht
Handelsregulierung
Handelsvorschriften
commercial movement regulation
commercial regulation
international trade rules
internationale Handelsbestimmungen
kommerzielle Bewegungsvorschriften
trade compliance
trade law
OIV (2023) State of the World Vine and Wine Sector, International Organisation of Vine and Wine European Parliament (2016) Regulation (EU) 2016/2031 on protective measures against plant pests FAO (2023) International Plant Protection Convention (IPPC) standards EU-US Wine Agreement (2006) Agreement between the European Community and the United States on trade in wine
Trade Regulation
Furmint in the Tokaj region of Hungary fulfills a traditional cultivar role with documented cultivation since at least the 16th century, forming the backbone of Tokaji Aszú production for over 400 years, legally mandated as the primary permitted variety comprising 60-100% of Tokaji blends under Hungarian wine law, and so deeply identified with the region that the cultivar name and the appellation are culturally inseparable despite Furmint's presence in minor plantings in Slovenia and Austria.
A regional production role that inheres in a grapevine cultivar by virtue of its long-established presence and cultural embeddedness within a specific viticultural region, where centuries of cultivation have generated deep associations between the cultivar and regional identity, winemaking traditions, and local gastronomic culture.
Traditional_Cultivar_Role captures the historically contingent relationship between a cultivar and a region, distinct from purely genetic or botanical classification. The role is inherently relational and context-dependent: Cabernet Sauvignon bears a traditional cultivar role in Bordeaux (documented since the 18th century) but an introduced cultivar role in China (planted from the 1980s onward). Subclasses distinguish three overlapping but conceptually distinct dimensions of traditionality — indigenous origin (first domesticated in the region), autochthonous presence (evolved in situ over centuries), and heritage status (cultural patrimony value regardless of geographic origin).
alteingesessene Rebsorten-Rolle
established cultivar role
historically rooted cultivar role
historisch verwurzelte Sorten-Rolle
traditional variety role
traditionelle Sorten-Rolle
This P et al. (2006) Historical origins and genetic diversity of wine grapes, Trends in Genetics 22: 511-519 Robinson J et al. (2012) Wine Grapes, Allen Lane pp: 1-18 Lacombe T et al. (2013) Large-scale parentage analysis in an extended set of grapevine cultivars, Theoretical and Applied Genetics 126: 401-414
Traditional Cultivar Role
A traditional gobelet head arm on ancient Vitis vinifera cv. Grenache in Châteauneuf-du-Pape, one of three arms radiating from gnarled head at 38cm height on 75-year-old vine, arm extending only 24cm at 60° angle, bearing just 3 two-bud spurs (6 total buds), representing severely limited fruiting capacity, total vine with three arms bearing 9 combined spurs supporting approximately 16 shoots and 2.1kg crop (equivalent to approximately 28 hl/ha at 1.5m × 1.5m spacing typical of traditional Châteauneuf), exemplifying extreme traditional gobelet approach maintained in prestigious estate emphasizing low yields, old vines, and terroir purity, with market accepting low productivity due to exceptional wine quality and pricing supporting economic viability despite yields one-third to one-half of modern cordon-trained alternatives.
A gobelet head arm maintained in strict accordance with classical training principles and regional traditions, characterized by short arm length (typically 20-40cm), low arm count (3-4 arms standard, rarely more than 5), minimal spur retention (2-3 buds per spur, 3-5 spurs per arm), low overall vine height (head typically 30-50cm), and deliberate limitation of vine capacity producing very low yields (often 2-3.5 kg per vine, translating to 20-35 hectoliters per hectare) emphasizing fruit concentration and wine quality over productivity. Traditional gobelet head arms represent purist expression of historical gobelet training, maintained in prestigious appellations where regulations mandate traditional practices, heritage vineyards preserving centuries-old approaches, and quality-focused producers emphasizing extreme terroir expression through severe yield restriction, accepting economic challenges of low productivity in exchange for perceived quality benefits and market positioning as authentic traditional wines.
Traditional gobelet head arms represent the conservative extreme of head training, with purist adherence to historical practices often driven by appellation regulations (many French AOC and Spanish DO systems specifying training requirements), quality philosophy (belief that extreme yield restriction enhances terroir expression and wine character), market positioning (traditional gobelet-trained wines commanding premium pricing), and cultural heritage preservation (maintaining centuries-old viticultural landscapes and practices). The severe yield limitation (often 25-40% of cordon-trained equivalent) raises economic sustainability questions, with viability depending on premium pricing, estate winery direct sales capturing full value chain, tourism and heritage value supplementing wine sales, or subsidies and protections supporting traditional viticulture. Young winemakers increasingly question whether extreme traditional approaches are necessary for quality or represent historical inertia, though defenders argue that centuries of tradition and demonstrated quality outcomes justify conservative practices.
classical gobelet arm
heritage gobelet arm
klassische Becherform-Arm
strict traditional head arm
traditionelle Gobelet-Arm
Traditional Gobelet Head Arm
The traditional regional name Blaufränkisch has been the established designation in eastern Austria since at least the 18th century (first documented 1777), carrying deep cultural associations with Burgenland identity and Pannonian viticultural heritage — the name persists as the official Austrian designation despite the same cultivar being known as Kékfrankos in Hungary, Lemberger in Germany, and Frankovka in the Czech Republic, with Austrian wine law explicitly protecting Blaufränkisch as the mandatory labeling term and Austrian producers resisting any pressure toward nomenclatural harmonization precisely because the traditional regional name embodies irreplaceable cultural capital.
A regional name that has been in established local usage for an extended historical period — conventionally several generations or centuries — and is recognized within the region as the customary designation for the cultivar, often carrying cultural significance, terroir associations, and identity value that transcend its purely nomenclatural function.
Traditional_Regional_Name captures designations that have transcended their purely referential function to become cultural heritage objects in their own right. These names are often fiercely defended by regional producers and wine authorities against standardization pressures because they encode generations of place-specific viticultural identity: the name Spätburgunder in Germany, Blauburgunder in Switzerland, and Pinot Nero in Italy each carry cultural resonances specific to their region that the international prime name Pinot Noir does not convey. EU wine legislation recognizes this cultural dimension by permitting traditional regional names as accepted synonyms on wine labels even when they differ from the official prime name.
althergebrachter Gebietsname
customary regional designation
established regional name
tradierter Ortsname
traditional local name
traditioneller Regionalname
Robinson J et al. (2012) Wine Grapes, Allen Lane Anderson K and Aryal NR (2013) Which Winegrape Varieties are Grown Where?, University of Adelaide Press EU Regulation 1308/2013 Art. 93-108 on wine designations
Traditional Regional Name
Detailed documentation of Trailing_Growth_Disposition at Geisenheim University characterized shoot architecture in cultivars recognized for pendulous, trailing growth habit. Reference cultivars 'Gewürztraminer', 'Riesling', 'Pinot Gris', and 'Muscat Ottonel' were grown on multiple training systems with comprehensive angular and mechanical measurements. Trailing_Growth_Disposition cultivars showed consistent extreme characteristics. 'Gewürztraminer' exhibited archetypal trailing growth with mean shoot angle of 72° from vertical at 50 cm, reaching 95° (below horizontal) at 80 cm, with shoot tips pointing downward in a true pendulous orientation. Gravitropic correction was absent—tips continued the downward trajectory established by mid-shoot. 'Riesling' showed strong Trailing_Growth_Disposition (65° at 50 cm, 85° at 80 cm) with characteristic arching profile. Internode stiffness was lowest among growth habit categories, averaging 1.5 N·mm² in trailing cultivars, and internode length was 15% longer than upright cultivars at equivalent vigor. VSP positioning trials confirmed incompatibility—'Gewürztraminer' required 65+ hours/ha positioning labor with 55% position loss, making VSP management economically impractical. Alternative training system trials demonstrated that Trailing_Growth_Disposition cultivars excelled on Scott Henry (lower curtain shoots achieving 45° below horizontal with excellent exposure), pendulum/curtain systems, and traditional Mosel steep-slope training where downward shoot growth is managed by single-stake support. Light interception efficiency for 'Riesling' was 22% higher on appropriately designed curtain systems compared to VSP with intensive positioning, demonstrating that matching Trailing_Growth_Disposition to training system optimizes canopy function.
A Trailing_Growth_Disposition is a Shoot_Orientation_Disposition that inheres in Vitis vinifera shoots and is realized through minimal or absent negative gravitropism producing shoots that grow at angles exceeding 60° from vertical, with mature shoots often approaching or exceeding horizontal (> 90°, true trailing/pendulous), creating canopies that naturally cascade downward without support. This disposition represents the extreme of non-upright growth, characteristic of cultivars that show negligible gravitropic correction as shoots elongate, resulting in pendulous, curtain-like canopy architecture when grown on high-wire or pergola systems. Trailing cultivars are poorly suited to VSP training, requiring either extremely intensive positioning labor with poor retention, or acceptance of alternative canopy forms. These cultivars are naturally compatible with systems designed for downward-growing shoots, including Scott Henry (lower wire shoots), pendulum/curtain systems, and traditional high pergola training. The mechanical basis of Trailing_Growth_Disposition includes very low internode stiffness (flexible shoots that bend easily under their own weight), weak gravitropic sensing or response, and often longer internodes that increase the mechanical moment causing shoot bending. Some trailing cultivars originated in viticultural traditions using pergola or arbor training where trailing growth was advantageous or at least not problematic.
Trailing_Growth_Disposition is_a Shoot_Orientation_Disposition representing pendulous/trailing growth (> 60° from vertical, often exceeding horizontal). Characterized by absent gravitropism, very low internode stiffness, and often longer internodes. Examples include 'Gewürztraminer', 'Riesling', 'Pinot Gris', 'Pinot Blanc', 'Muscat Ottonel', 'Grüner Veltliner'. Incompatible with VSP suited to Scott Henry lower tier, pendulum/curtain, and pergola systems. Many Alsatian and German cultivars show this trait. Traditional training in regions of origin often accommodates trailing habit.
Hängende-Wuchs-Disposition
Pendelnder-Habitus
cascading growth
drooping habit
pendulous growth
Überhängender-Triebwuchs
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles, Adelaide Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review, American Journal of Enology and Viticulture 60:251-268 Intrieri C, Poni S (1995) Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality of mechanized Italian vineyards, American Journal of Enology and Viticulture 46:116-127
Trailing Growth Disposition
Trailing growth suitability quality comparison in Riverland evaluated cultivar performance under Geneva Double Curtain training designed for high-vigor, trailing-habit cultivars. Shiraz demonstrated excellent trailing growth suitability quality with mean shoot angle of 35° below horizontal when trained on high bilateral cordons, naturally creating separated downward-growing curtains with minimal shoot intermingling and achieving 92% fruit positioning in the open zone between curtains without intervention. Grenache showed moderate trailing growth suitability quality with mean angle of 15° below horizontal, requiring quarterly curtain separation to prevent intermingling and achieving 78% optimal fruit positioning. Cabernet Sauvignon exhibited poor trailing growth suitability quality with mean angle of 8° above horizontal indicating weak upright tendency, with shoots continuously lifting toward vertical and causing severe curtain convergence that required five separation passes annually while achieving only 54% optimal positioning, ultimately motivating conversion to vertical shoot positioning where its mild upright habit became advantageous rather than problematic.
Trailing growth suitability quality is a growth habit compatibility quality that inheres in a grapevine cultivar or clone and represents the degree to which the genotype exhibits naturally pendant or horizontal shoot orientation with weak negative gravitropic response or positive gravitropic tendency that directs shoot development toward horizontal or downward trajectories, conferring high compatibility with divided canopy systems, Geneva Double Curtain, lyre configurations, pergola and tendone training, and other systems designed to accommodate or exploit trailing shoot behavior. This quality determines the effectiveness of training systems that depend on shoots naturally assuming horizontal or drooping positions to create separated canopy curtains, overhead fruit positioning, or split canopy architectures that increase exposed surface area and improve light distribution compared to single vertical canopy configurations. The ontological significance of trailing growth suitability quality lies in its role enabling specific training system options that would be impractical or impossible with strongly upright cultivars, recognizing that training systems exploiting trailing growth habit can provide superior canopy management solutions for vigorous cultivars and high-yield production systems where vertical configurations prove inadequate. The physiological basis of trailing growth habit involves reduced negative gravitropic response strength such that shoots do not strongly orient against gravity, increased shoot flexibility that permits bending under the weight of developing leaves and fruit clusters, and potentially positive phototropic responses directing growth toward open light environments at canopy margins rather than upward through overlying foliage. Cultivars with high trailing growth suitability quality naturally position shoots horizontally or pendant when growing from elevated cordons or canes, maintain stable trailing orientation without continuous lifting toward vertical, and produce functional canopy architectures in divided or overhead systems with minimal positioning intervention. The assessment of trailing growth suitability quality involves evaluation of shoot orientation in freely-growing conditions from elevated training positions, measurement of shoot angle distributions relative to horizontal, and practical trials in training systems designed for trailing growth to evaluate canopy organization quality and management efficiency. The combination of trailing growth habit with appropriate vigor levels proves particularly valuable for divided canopy systems where sufficient growth is needed to fill multiple curtains while the trailing habit ensures curtain separation rather than intermingling.
Trailing_Growth_Suitability_Quality is_a Growth_Habit_Compatibility_Quality, representing the specific compatibility of pendant or horizontal growth habit with divided canopy, overhead, and sprawling training systems. This class addresses cultivars exhibiting naturally trailing shoot development, opposite to Upright_Growth_Suitability_Quality. The shoot angle below horizontal provides quantitative assessment metric. Most relevant for Geneva Double Curtain, lyre, Smart-Dyson, pergola/tendone, and related systems designed to exploit or accommodate trailing growth. Often associated with higher vigor cultivars.
Hängewuchseignung
Pendelwuchsneigung
drooping growth tendency
hängende Triebhaltung
pendant growth habit
positive gravitropism tendency
trailing shoot orientation
überhängende Wuchstendenz
Trailing Growth Suitability Quality
Training response quality evaluation in Alsace compared cultivar behavior during establishment and ongoing management of vertical shoot positioned training. Pinot Blanc demonstrated excellent training response quality with flexible young shoots readily tucked between catch wires without breakage, rapid lignification securing positioned orientation, and established canes maintaining imposed form with less than 3% repositioning failure rate at season end. Riesling showed good training response quality with appropriate flexibility during active growth though somewhat increased brittleness in mature shoots requiring careful handling during late-season adjustments. Gewürztraminer exhibited challenging training response quality with inherently stiff shoots resisting positioning during the optimal timing window, elevated breakage incidence of 8% during catch wire insertion, and persistent tendency for shoots to escape imposed orientation requiring 40% more positioning labor than Pinot Blanc to achieve equivalent canopy organization, establishing training response quality as a significant factor in cultivar management cost differences.
Training response quality is a viticultural quality that inheres in a grapevine cultivar or clone and represents the characteristic manner in which shoots and canes respond to the physical manipulations involved in training system establishment and annual canopy management, encompassing the flexibility, resilience, and adaptive capacity of vegetative tissues when subjected to bending, positioning, tying, and attachment operations that orient vine growth within prescribed trellis architectures. This quality determines the ease and success of training operations, the labor requirements for achieving target canopy configurations, and the long-term structural integrity of trained vine components including cordons, canes, and positioned shoots that must maintain imposed orientations throughout their functional lifespans. The ontological significance of training response quality lies in its role as a practical determinant of management efficiency and achievable canopy quality, wherein cultivars exhibiting favorable training response characteristics enable rapid, effective vine manipulation with minimal tissue damage or repositioning failure, while those with poor training response require increased labor inputs and may fail to achieve target architectures despite intensive management efforts. The biological basis of training response quality encompasses the mechanical properties of grapevine tissues including flexibility, elasticity, and breaking strength the capacity for adaptive growth in response to imposed orientations the wound healing response following training-related tissue stress and the developmental plasticity permitting shoots to acclimate to positions differing from their natural growth direction. Shoot flexibility varies with tissue maturity, with young actively-growing shoots exhibiting greater pliability than lignified canes, creating timing dependencies for training operations that must balance flexibility advantages against the risk of damage to tender tissues. Cultivar differences in training response quality reflect variation in wood density, fiber composition, vascular architecture, and developmental patterns governing the transition from herbaceous to lignified tissue states. Assessment of training response quality involves practical evaluation of positioning labor requirements, breakage incidence during training operations, maintenance of imposed orientations over time, and long-term structural health of trained vine components.
Training_Response_Quality serves as the parent class for Bending_Tolerance_Quality, Positioning_Ease_Quality, and Tendril_Attachment_Quality which address specific dimensions of training response. This class captures the overall concept of vine response to training manipulations while child classes address particular aspects. Distinguished from Growth_Habit_Compatibility_Quality which addresses natural orientation tendencies rather than response to imposed training. Practical significance includes labor efficiency, breakage prevention, and achievement of target canopy architectures.
Erziehungsreaktion
Formierungstoleranz
Laubarbeits-Anpassungsfähigkeit
Triebpositionierungs-Reaktion
canopy manipulation response
shoot positioning response
training adaptability
vine training plasticity
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 64-85 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 85-112 Galet P (2000) General Viticulture. Oenoplurimedia, Chaintre, pp. 189-245
Training Response Quality
The training system description for a precision-managed Cabernet Sauvignon vineyard in the Napa Valley specifies a bilateral cordon-trained, vertical shoot positioned (VSP) system on a two-wire movable canopy trellis at 1.8 m row spacing and 1.5 m vine spacing (3,700 vines/ha), with cordon height at 0.9 m, fruiting zone between 0.9-1.2 m, two pairs of movable foliage wires at 1.3 m and 1.6 m, shoot trimming height at 1.8 m yielding a canopy height-to-row-width ratio of 0.5, and spur pruning to 14-16 two-bud spurs per vine targeting 28-32 shoots per vine for a crop load of 6-8 t/ha with a Ravaz index (yield-to-pruning-weight ratio) of 5-7.
A viticultural practice description that documents the architectural framework imposed on a grapevine through systematic pruning, shoot positioning, and trellis support to establish and maintain a defined vine form, canopy geometry, and spatial arrangement of vegetative and reproductive organs that optimizes the balance between light interception, air circulation, labor efficiency, and fruit quality for specific cultivar-climate-management combinations.
Training_System_Description documents the fundamental architectural decisions that shape every aspect of vineyard performance from establishment through the productive life of the vineyard (typically 25-50+ years). The choice of training system is among the most consequential and least reversible viticultural decisions because it determines planting density, trellis infrastructure investment, mechanization compatibility, labor requirements, canopy microclimate, and the range of management interventions available throughout the vineyard's life: converting between training systems typically requires partial or complete vineyard replanting at a cost of 30,000-80,000 EUR/ha. Subclasses capture the key informational components of training system documentation — the system name, detailed specifications, trellis design, wire configuration, post spacing, and canopy height parameters.
Erziehungssystem-Beschreibung
Laubwandarchitektur-Spezifikation
Reberziehungs-Dokumentation
canopy architecture specification
trellis system documentation
vine training description
Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles Reynolds AG and Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition, American Journal of Enology and Viticulture 60: 251-268 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 331-380
Training System Description
The training system name Guyot simple (single Guyot) designates the cane-pruned system predominant in Burgundy, Champagne, and the Loire Valley — one fruiting cane of 6-10 buds retained annually plus one two-bud renewal spur, the cane tied horizontally to a single fruiting wire, shoots trained vertically through foliage wires in VSP configuration. The name Guyot honors Dr. Jules Guyot who systematized and promoted the system across France in the 1860s, and the specification simple (single) distinguishes it from Guyot double (bilateral cane pruning with two fruiting canes), with the training system name appearing in appellation cahiers des charges where specific training methods are prescribed or recommended.
A training system description consisting of the standardized or traditional name identifying a specific vine training configuration, referencing either a codified viticultural system (e.g., Guyot, cordon de Royat, Geneva Double Curtain) or a regional traditional practice (e.g., Pergola Trentina, Alberello, Lenz Moser), with the name serving as the primary reference key linking to the full set of architectural specifications and management protocols associated with that training system.
Training_System_Name provides the standardized vocabulary for communicating vine training configurations across the viticultural community. The nomenclature is heterogeneous, combining personal names (Guyot, Lenz Moser, Scott Henry, Geneva Double Curtain), descriptive terms (vertical shoot positioning, bilateral cordon, head training), regional traditional names (Alberello, Pergola, Lyre, Gobelet), and hybrid designations: this nomenclatural diversity reflects the independent development of training systems across wine regions over centuries. Some training system names are precisely defined (Geneva Double Curtain specifies a divided canopy with pendant shoot positioning developed by Nelson Shaulis at Cornell/Geneva in the 1960s), while others are generic categories encompassing considerable variation (Pergola describes any overhead canopy system but ranges from traditional Pergola Trentina to modern overhead arbor systems with fundamentally different canopy characteristics).
Erziehungssystem-Name
Reberziehungsform-Bezeichnung
training system designation
trellis system name
vine form name
Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles pp: 25-60 Galet P (2000) General Viticulture, Oenoplurimedia Reynolds AG and Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition, American Journal of Enology and Viticulture 60: 251-268
Training System Name
A Cabernet Sauvignon vine trained to vertical shoot positioning (VSP) on bilateral cordon bears a Training_System_Role determining its upward shoot growth, fruit zone positioning at cordon height, and suitability for mechanical leaf removal and harvesting. A Grenache vine in traditional Châteauneuf-du-Pape gobelet training serves in a Training_System_Role adapted to Mistral wind exposure, heat stress, and low-vigor conditions—its bush form providing self-shading and wind resistance without trellis infrastructure. A Riesling vine on single-stake Mosel training bears a Training_System_Role evolved for steep-slope viticulture, with individual stake support enabling cultivation on gradients exceeding 60% where row-based trellising is impractical.
A vineyard role that inheres in a grapevine by virtue of the architectural system to which it is trained, determining the spatial arrangement of permanent wood (trunk, cordons, arms), annual growth (canes, shoots), and fruit-bearing positions within the vineyard space. Training system roles define vine shape, canopy geometry, and fruiting zone positioning through the interaction of vine training (establishing permanent structure) and pruning (managing annual growth), with system selection reflecting variety characteristics, climate, mechanization requirements, and production objectives. Major training system categories include head-trained (self-supporting with no permanent cordons), cordon-trained (horizontal permanent arms on trellis), cane-pruned (annual replacement of fruiting wood), and spur-pruned (permanent fruiting positions on cordons) configurations. Training system selection profoundly influences vine physiology—canopy microclimate, light interception, fruit exposure, disease pressure, and labor requirements all vary with training system choice. Traditional regional systems evolved through centuries of empirical optimization, while contemporary viticulture increasingly applies physiological understanding to match training systems to specific site and variety requirements.
Training_System_Role is subclassed by fundamental architectural category—Head_Trained (no permanent horizontal cordons) and Cordon_Trained (permanent horizontal structures)—with further subdivision by specific system implementations. Training system terminology varies internationally the same physical system may have different regional names, while identical terms may describe different systems in different regions. The choice between head-training and cordon-training reflects fundamental decisions about vine architecture—head-training offers simplicity and wind resistance but limits mechanization, while cordon-training enables mechanical operations and precise canopy management but requires trellis investment. Training system effects on fruit composition are well-documented: exposed fruit zones increase phenolic development but risk sunburn, while shaded zones preserve acidity but may reduce color and flavor intensity, with optimal exposure varying by variety, climate, and wine style objectives.
Erziehungssystem
Laubwandsystem
Reberziehung
Spalierform
canopy system
training system
trellis system
vine training
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition, American Journal of Enology and Viticulture 60: 251-268 Jackson RS (2014) Wine Science: Principles and Applications, Academic Press
Training System Role
The training system specification for a bilateral cordon-trained VSP system in a premium Merlot vineyard in Saint-Émilion details vine spacing of 1.0 m within rows and 1.5 m between rows (6,667 vines/ha), bilateral cordons extending 0.5 m each side of the trunk at 0.6 m height, spur pruning to 5-6 two-bud spurs per cordon (10-12 per vine, targeting 20-24 shoots), vertical shoot positioning through 3 pairs of movable foliage wires at 0.9 m, 1.2 m, and 1.5 m, summer trimming at 1.6 m canopy height, cluster-zone leaf removal on the morning-sun (east) side at BBCH 73-75, and green harvest at véraison to a maximum of 1 cluster per shoot yielding approximately 40-50 hl/ha.
A training system description consisting of the detailed quantitative and qualitative parameters that define the complete architectural configuration of a specific training system implementation, including vine spacing, cordon or cane dimensions, bud numbers, shoot density targets, pruning method, shoot positioning protocol, and seasonal management sequence required to establish and maintain the system in operational condition.
Training_System_Specification captures the implementation-level detail required to translate a training system name into a functional vineyard installation. While the training system name identifies the general architectural concept, the specification defines the precise dimensional, structural, and management parameters that determine actual vine performance: two vineyards both nominally trained to bilateral cordon VSP may perform very differently if one is planted at 5,000 vines/ha with 14 buds per vine and the other at 3,000 vines/ha with 28 buds per vine — the training system name is identical but the specifications create fundamentally different canopy density, microclimate, and crop load conditions. Appellation production specifications (cahiers des charges, disciplinari) typically prescribe certain specification parameters — particularly planting density, maximum buds per vine, and training system type — as regulatory requirements for appellation compliance.
Erziehungssystem-Spezifikation
Reberziehungs-Parameter
Systemkonfigurations-Detail
system configuration details
training system parameters
vine architecture specification
Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 331-380 INAO (2023) Cahiers des charges — Conditions de production
Training System Specification
Training system suitability quality evaluation for Pinot Noir across training systems in Burgundy and New Zealand compared performance outcomes revealing strong genotype-training interactions. Traditional Burgundy single Guyot training demonstrated excellent training system suitability quality for this moderate-vigor, upright-growing cultivar, achieving optimal vine balance with Ravaz Index of 6.5 and superior fruit quality for premium wine production. Conversion trials to Scott Henry divided canopy in New Zealand showed poor training system suitability quality, with insufficient vigor to adequately fill both curtains resulting in sparse canopies, sunburn damage from excessive exposure, and yield reduction of 28% compared to vertical shoot positioning. Minimal pruning trials revealed fundamental incompatibility with Pinot Noir's compact cluster architecture, resulting in severe Botrytis pressure from poor air circulation within the dense, unmanaged canopy structure, confirming that training system suitability quality assessment must precede system selection to avoid costly mismatches between cultivar characteristics and training system requirements.
Training system suitability quality is a viticultural quality that inheres in a grapevine cultivar or clone and represents the degree of compatibility between the inherent growth characteristics, vigor expression patterns, and fruiting habits of the genotype and the architectural requirements, management demands, and environmental conditions created by specific training system configurations including vertical shoot positioning, divided canopy systems, minimal pruning, and traditional regional forms. This quality determines the likelihood of achieving optimal vine balance, fruit quality, and operational efficiency when a particular cultivar-training system combination is implemented, recognizing that training systems impose specific constraints and opportunities that interact differentially with cultivar growth habits to produce outcomes ranging from excellent synergy to fundamental incompatibility. The ontological significance of training system suitability quality lies in its role as a predictive indicator enabling informed decisions regarding training system selection for new plantings and cultivar selection for existing infrastructure, wherein matching genotype characteristics to training system requirements represents a foundational determinant of vineyard success that cannot be readily corrected through subsequent management interventions. The biological basis of training system suitability encompasses multiple cultivar characteristics including shoot growth habit determining natural orientation of developing shoots, vigor level influencing the capacity to fill prescribed canopy dimensions, internode length affecting shoot positioning requirements, lateral bud fruitfulness determining productivity under different pruning approaches, cluster architecture influencing disease susceptibility under various exposure conditions, and fruit composition sensitivity to the microclimate conditions created by different canopy configurations. Training systems vary fundamentally in their requirements and tolerances, with vertically shoot positioned systems demanding cultivars exhibiting upright growth habit and moderate vigor that can be contained within narrow canopy dimensions, while divided canopy systems accommodate higher vigor through distribution across multiple curtains, and minimal pruning systems require cultivars tolerant of high crop loads and capable of self-regulation under minimal intervention. The assessment of training system suitability quality involves evaluation of cultivar performance across multiple training configurations under comparable conditions, supplemented by mechanistic understanding of how specific cultivar traits interact with training system characteristics to produce observed outcomes. Regional viticulture traditions often encode empirical knowledge of training system suitability through historical selection of locally-adapted cultivar-training combinations, though modern understanding enables more systematic evaluation and the identification of novel combinations that may outperform traditional practices.
Training_System_Suitability_Quality serves as the parent class for Growth_Habit_Compatibility_Quality and potentially other suitability dimensions addressing vigor requirements, fruiting habit compatibility, and mechanization adaptability. This class captures the overall concept of cultivar-training system matching while child classes address specific compatibility dimensions. Distinguished from Vigor_Quality and related classes by the focus on the interaction between genotype and training architecture rather than vigor per se. Critical for vineyard establishment decisions and regional cultivar recommendations.
Erziehungssystemeignung
Laubwandsystemverträglichkeit
Reberziehungskompatibilität
Spaliereignung
canopy system compatibility
training compatibility
trellis system suitability
vine training adaptability
Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review. American Journal of Enology and Viticulture 60:251-268 Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 32-43 Jackson DI, Lombard PB (1993) Environmental and management practices affecting grape composition and wine quality: a review. American Journal of Enology and Viticulture 44:409-430
Training System Suitability Quality
Syrah in Rhône Valley shows transition to enlargement onset during late July-early August Year N+1 at BBCH 73-75, with progressive cell cycle exit beginning week 3.5, water uptake initiating 0.3-0.7 g per berry per day, vacuole formation in enlarging cells, overlap period with declining division and increasing enlargement weeks 3.5-5, berry diameter expanding from 14 mm to 20 mm during transition, under warm Mediterranean conditions of 25-30°C temperature, typical of this mid-to-late season cultivar transitioning growth mode
Transition to enlargement onset represents the growth mode shift phase during declining division activity when cells progressively commit to expansion rather than continued division, water uptake and vacuole development initiate creating cell enlargement, overlap period proceeds with some cells still dividing while others expand, and transition establishes expansion as emerging dominant growth mechanism, occurring over weeks 3.5-5 as division declines and enlargement accelerates. This critical transition phase occurs during mid-late summer in mid-late July to early August in temperate regions Year N+1, spanning approximately 1.5-2 weeks representing overlap period, corresponding to BBCH 73-75 pea-sized to berry touch stages. Transition mechanism involves cell cycle exit decision with cells choosing expansion over continued division, enlargement initiation in exiting cells through water uptake and vacuole formation, progressive enlargement onset across cell populations creating temporal spread, continued division in remaining cycling cells creating overlap, and gradual transition establishing expansion as predominant growth mode. Cell cycle exit and expansion commitment shows cells completing final division then entering G0/G1 expansion state rather than re-entering S phase, exit decision influenced by auxin decline and developmental signals, committed cells initiating expansion program, and irreversible commitment to enlargement versus division. Expansion commitment timing shows early-dividing cells exiting and enlarging beginning week 3, progressive exit wave through populations weeks 3.5-5, and extended commitment period creating gradual transition. Water uptake initiation shows expansion-committed cells beginning water import from surrounding apoplast, osmotic gradient from accumulating solutes driving water influx, water uptake rates initially 0.2-0.5 g per berry per day early transition increasing to 0.5-1.0 g per day as more cells enlarge, and water as primary driver of cell volume increase. Osmotic driving force shows organic acid accumulation particularly malate and tartrate creating osmotic potential decline from -0.5 MPa to -0.8 MPa, ion accumulation including potassium contributing to osmotic gradient, sugar beginning to accumulate though levels still low 3-6%, and combined solutes creating gradient driving water influx. Vacuole formation and development shows central vacuole initiating in enlarging cells, vacuole originating from smaller vacuoles or ER-derived structures, progressive vacuole expansion occupying increasing cell volume, and eventual vacuole dominance at 80-90% cell volume in fully expanded cells. Vacuole characteristics show large central compartment storing water and solutes, tonoplast membrane separating vacuole from cytoplasm, proton pumps and transporters regulating vacuole composition, and vacuole as major storage and osmotic compartment. Cell enlargement initiation shows cell volume beginning increase from division-exit size 20-40 micrometers diameter, initial expansion relatively slow 2-3 fold increase during transition, and acceleration during subsequent expansion phase achieving eventual 10-100 fold total increase. Cell wall modification enabling expansion shows wall loosening through expansin proteins disrupting cellulose-hemicellulose network, xyloglucan endotransglucosylase rearranging wall polymers, pectin methylesterase modifying pectin properties affecting wall plasticity, and coordinated wall modification enabling turgor-driven expansion. Turgor pressure development shows osmotic water influx generating turgor pressure 0.3-0.8 MPa during early enlargement, turgor force driving cell expansion against yielding walls, and pressure maintenance through continued solute accumulation and water uptake. Division-enlargement overlap shows temporal overlap with some cells still dividing while others enlarging, overlap duration approximately 1.5-2 weeks during transition phase, declining division from week 3.5-5 concurrent with increasing enlargement, and overlap characteristic of asynchronous cell population transitions. Cell population asynchrony shows variation in transition timing across cells, early-exiting cells enlarging while late-dividing cells continue cycling, asynchrony creating smooth gradual transition, and population heterogeneity extending overlap period. Enlargement acceleration during transition shows enlargement contribution to growth increasing as more cells commit and enlarge, berry volume increase from combined declining division and increasing enlargement, growth rate maintained or increasing during transition from compensating enlargement, and eventual enlargement dominance as division completes. Berry growth pattern during transition shows continued volumetric increase from combined mechanisms, berry diameter expanding from 12-18 mm early transition to 15-25 mm late transition depending on cultivar, fresh weight increasing from 2-5 g to 4-10 g during transition, and growth maintenance during mode shift. Metabolic changes during transition show shift from division-associated metabolism to expansion metabolism, respiratory rate remaining elevated supporting active growth, carbohydrate import supporting osmotic gradient and wall synthesis, and metabolic reorganization preparing subsequent rapid expansion phase. Gene expression changes show downregulation of cell cycle genes as division declines, upregulation of expansion-associated genes including expansins and transporters, and transcriptional reprogramming establishing expansion program. Environmental effects on transition include temperature affecting transition timing through developmental rate effects with warm conditions potentially accelerating transition, water availability affecting enlargement onset and rate with adequate water essential for expansion initiation, and overall favorable conditions supporting smooth transition. Temperature effects show warm conditions promoting faster transition completion, cool conditions extending overlap period through slowed progression, and temperature as factor affecting transition timing. Water effects show adequate water essential for enlargement initiation, water limitation potentially delaying or limiting expansion onset, and water as critical resource for transition to enlargement. Transition assessment methods include cell size distribution analysis showing emerging enlarged population, vacuole development observation documenting enlargement onset, cell cycle marker decline showing division reduction, expansion marker expression showing enlargement activation, and growth pattern analysis documenting mode shift. The process represents second aspect of declining division activity emphasizing growth mode transition, following division rate reduction and preceding cell number finalization. Duration typically 1.5-2 weeks representing overlap period weeks 3.5-5. Regional timing shows transition to enlargement onset during mid-late July to early August across climates. Viticultural significance includes representing critical developmental transition from division to expansion as growth mechanism, establishing enlargement as emerging dominant process, demonstrating overlap period maintaining growth rate, and marking progression toward expansion-dominated berry growth phase. Research applications include analyzing transition mechanisms and regulation, investigating cell fate decisions and expansion commitment, evaluating water relations during enlargement onset, quantifying division-enlargement overlap dynamics, and assessing transition as determinant of subsequent expansion phase.
Transition_to_Enlargement_Onset is second aspect of Declining_Division_Activity, distinguished by growth mode shift from division to expansion. Represents overlap period with declining division concurrent with increasing enlargement. Duration 1.5-2 weeks representing weeks 3.5-5. Water uptake 0.2-1.0 g/berry/day. Vacuole development initiating. Cell volume beginning 2-3 fold increase. Temperature and water affect transition. Follows Division_Rate_Reduction. Precedes Cell_Number_Finalization. Critical developmental transition establishing expansion dominance.
Teilung-zu-Expansion-Übergang
Wachstumsmodus-Wechsel
division to expansion transition
growth mode shift
Übergang zu Vergrößerungs-Beginn
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 438-442 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Transition to Enlargement Onset
Cabernet Sauvignon transitional middle pericarp at outer boundary showing graduated properties= outer mesocarp (outer pericarp inner layers) with cells 48-72 μm, walls 1.2-1.8 μm, tannins 4.5-7 mg/g, anthocyanins 1.2-2.5 mg/g transitional zone with cells 65-105 μm, walls 0.8-1.3 μm, tannins 2-4.5 mg/g, anthocyanins 0.3-1 mg/g outer middle pericarp with cells 92-130 μm, walls 0.55-0.85 μm, tannins 0.4-1.2 mg/g, anthocyanins 500 μm from seeds) with cells 152-175 μm, tannins 0.5-0.9 mg/g 300-500 μm from seeds with cells 128-158 μm, tannins 1.2-2.2 mg/g 150-300 μm with cells 98-135 μm, tannins 2.5-4 mg/g inner pericarp (400 μm) <0.05 mg/g, demonstrating exponential decline in anthocyanin concentration from hypodermis maximum through transitional zone into middle pericarp creating gradient extending 200-300 μm depth. Pinot Noir comparison of transition zone steepness= cluster exterior sun-exposed berries showing sharper transition with anthocyanins declining from 2.8 mg/g (hypodermis) to 0.2 mg/g (middle pericarp) over 100-150 μm depth cluster interior shaded berries showing more gradual transition with anthocyanins declining from 1.8 mg/g to 0.2 mg/g over 180-250 μm depth, suggesting light exposure affects not only absolute anthocyanin levels but also spatial distribution creating steeper gradients in exposed berries potentially affecting skin tannin-to-anthocyanin ratio and wine extraction characteristics.
A transitional middle pericarp is the middle pericarp tissue at boundaries between middle pericarp and outer pericarp or between middle pericarp and inner pericarp, showing mixed or intermediate characteristics reflecting transition from middle pericarp properties (large cells, thin walls, low phenolics) to adjacent tissue properties (smaller cells, thicker walls, higher phenolics in outer or inner pericarp), representing boundary zones where tissue character gradually shifts rather than changing abruptly. Transitional middle pericarp at the outer boundary (middle-to-outer transition) comprises= the innermost 2-4 cell layers of what could be classified as outer pericarp or the outermost 3-6 cell layers of middle pericarp, typically 50-150 μm depth, showing progressive change in cell size from large middle pericarp cells (80-130 μm diameter at transition boundary) to smaller outer mesocarp cells (40-80 μm) to very small hypodermal cells (15-30 μm) progressive increase in cell wall thickness from thin middle pericarp (0.5-0.8 μm) to thicker outer mesocarp (0.8-1.5 μm) to thick hypodermis (1.5-3 μm) progressive increase in phenolic content from minimal middle pericarp (<1 mg/g tannins, <0.05 mg/g anthocyanins) through outer mesocarp (2-6 mg/g tannins, 0.5-2 mg/g anthocyanins if red cultivar) to high hypodermis/epidermis (6-12 mg/g tannins, 2.5-6 mg/g anthocyanins). Transitional middle pericarp at the inner boundary (middle-to-inner transition) comprises= the innermost 4-8 cell layers of middle pericarp or the outermost 2-4 cell layers of inner pericarp, typically 100-250 μm depth, showing progressive change in cell size from large central/inner middle pericarp cells (110-150 μm diameter) to intermediate transitional cells (80-120 μm) to smaller inner pericarp cells (50-90 μm) progressive increase in cell wall thickness from thin middle pericarp (0.5-0.7 μm) to thicker transitional zone (0.7-1.1 μm) to thicker inner pericarp (0.9-1.5 μm) progressive increase in tannin content from minimal middle pericarp (<0.8 mg/g) through transitional zone (1.5-3.5 mg/g) to elevated inner pericarp (3-7 mg/g). Transitional middle pericarp functional significance= gradual rather than sharp tissue boundaries preventing mechanical stress concentrations that could occur at abrupt property changes gradual compositional gradients potentially affecting extraction kinetics during winemaking with transitional zones extracted earlier or more completely than central middle pericarp developmental gradients reflecting position-dependent inductive signals (outer pericarp or seed signals diffusing into adjacent middle pericarp) affecting cell differentiation and metabolism. Transitional middle pericarp challenges in tissue classification= ambiguity in defining precise boundaries between tissues as transitions are gradual rather than discrete variation among cultivars in transition zone width and gradient steepness potential variation in transition characteristics among berries within clusters or clusters within vines these factors create challenges for precise anatomical classification requiring somewhat arbitrary boundary definitions. Transitional middle pericarp in berry development= transition zones established during early development when tissue differentiation occurs (BBCH 71-75, weeks 2-6) with positional identity determined by developmental signals transition zones maintained through ripening though cell expansion may alter relative proportions of tissues transition zone properties (particularly phenolic content) changing during ripening as anthocyanins and tannins synthesize and accumulate creating sharper compositional gradients at maturity than during early development.
Transitional_Middle_Pericarp represents boundary zones between middle and outer/inner pericarp and is distinguished from Intermediate_Middle_Pericarp (gradients within middle pericarp bulk) by location at tissue boundaries versus within middle pericarp. Transitional middle pericarp zones show gradual property changes reflecting smooth tissue transitions. Understanding transitional zones is relevant for detailed anatomical characterization and for interpreting extraction gradients during winemaking, recognizing that berry tissues show gradual transitions rather than sharp boundaries creating challenges for discrete tissue classification.
Grenz-Mesokarp
Grenzfruchtfleisch
boundary flesh
interface zone
transitionales mittleres Perikarp
Cadot, Y. et al. (2006). Tannin localization in grape skins. Journal of Agricultural and Food Chemistry 54=2973-2987 Harris, J.M. et al. (1968). The Vinifera grape berry= anatomical study. Botanical Gazette 129=1-23.
Transitional Middle Pericarp
Merlot vines in the Saint-Émilion appellation bearing a transitional organic role during their second year of conversion receive identical management to certified organic parcels — copper hydroxide at 3.5 kg Cu/ha/year, mechanical under-vine tillage, and Trichoderma harzianum applications — but fruit is marketed as "vin en conversion vers l'agriculture biologique" at 15-25% lower prices than certified organic equivalents until the 3-year conversion period is completed.
An organic production role that inheres in a grapevine during the mandatory conversion period between conventional and certified organic management, during which organic practices are fully implemented but the resulting grapes and wine may not yet be marketed as organic.
Transitional_Organic_Role captures the regulatory liminal status required by all major organic certification frameworks. Under EU Regulation 2018/848 the conversion period is 3 years before harvest for perennial crops including grapevines: under USDA NOP it is 36 months of documented organic management. Some certification bodies permit "in-conversion" labelling from the second year onward. The role terminates upon either successful certification (transition to Certified_Organic_Role) or abandonment of the organic pathway (reversion to Conventional_Production_Role).
Bio-Umstellungsrolle
Konversions-Rolle
Umstellungs-Rolle
converting-to-organic role
in-conversion organic role
organic transition role
EU Regulation 2018/848 Art. 10 on conversion periods USDA NOP 7 CFR 205.202 Willer H and Lernoud J (2023) The World of Organic Agriculture, FiBL-IFOAM pp: 78-82
Transitional Organic Role
A transitional semi-lignified cane on Vitis vinifera cv. Riesling monitored through sequential weekly assessments during September-October, showing brown lignified zone extending from base through node 6 (50cm) in week 1, advancing to node 8 (70cm) by week 2, reaching node 10 (90cm) by week 3, and extending to node 12 (110cm) by week 4, with accompanying progressive firmness increase in transitioning zones measurable through penetrometer testing, demonstrating active lignification front advancement of approximately 5cm per week under favorable warm autumn conditions (15-20°C daytime temperatures, cool 8-10°C nights) with adequate soil moisture and healthy functional canopy supporting continued carbohydrate production.
A semi-lignified one-year cane exhibiting active progression through lignification with visible changes occurring over short timeframes (days to weeks), characterized by expanding zones of brown coloration advancing from base toward apex, progressive firmness increase in transitioning regions, dynamic secondary cell wall development visible through sequential cross-sectional sampling, and physiological status representing the active maturation process rather than stable intermediate state. Transitional semi-lignified canes demonstrate the temporal dynamics of wood ripening, with lignification advancing as a moving front proceeding basipetally (from older to younger internodes) driven by coordinated gene expression patterns, enzyme activities, and substrate availability, responding to environmental signals including photoperiod, temperature, and endogenous hormonal controls that regulate the pace and extent of maturation progression.
Transitional semi-lignified canes represent the dynamic process of lignification rather than a stable developmental state, with their status constantly changing during the autumn maturation period. Understanding the rate and pattern of lignification progression is valuable for predicting final maturation status and timing pruning operations, with rapid progression indicating favorable conditions and high probability of complete maturation, while slow or stalled progression suggests potential problems requiring intervention. Monitoring can involve simple visual assessment (advance of brown color boundary) or more sophisticated measurements (firmness testing, destructive anatomical sampling, near-infrared spectroscopy). The lignification progression rate varies with environmental conditions, cultivar characteristics, vine vigor and crop load, making site-specific and season-specific observation valuable for management decision-making.
actively lignifying cane
aktiv ausreifende Rute
dynamic ripening cane
progressing maturation cane
übergangs-teilverholzte Rute
Transitional Semi Lignified Cane
Translocated incompatibility in grapevine is implicated when incompatibility symptoms appear in tissues distant from the graft union, including scion leaf chlorosis, abnormal root development, or generalized growth reduction that cannot be explained by localized union dysfunction. Phenolic compounds including catechins, cyanogenic glycosides, and prunasin (in Prunus) serve as translocated incompatibility factors in other genera, with analogous compounds suspected but not definitively identified in Vitis. Reciprocal grafting experiments distinguish translocated from localized incompatibility, as translocated factors cause symptoms regardless of which partner serves as scion versus rootstock. Symptoms may appear within the first growing season or develop progressively over 2-5 years as factor accumulation exceeds tolerance thresholds.
A graft incompatibility disposition that is the inherited capacity of certain Vitis vinifera graft combinations to exhibit incompatibility symptoms caused by adverse substances (toxins, phenolics, hormones) produced by one graft partner and transported to the other, causing systemic physiological disruption beyond the immediate graft zone.
Translocated incompatibility is less well documented in grapevine than in stone fruits, where specific cyanogenic compounds have been identified as incompatibility factors. The mechanism involves phloem or xylem transport of inhibitory substances from one partner to sensitive tissues of the other, causing toxicity at sites distant from the graft zone. Distinguishing translocated from localized incompatibility has practical implications, as translocated incompatibility cannot be remedied by improved grafting technique while localized incompatibility may respond to modified practices.
biochemical incompatibility
systemic incompatibility
systemische Unverträglichkeit
translocatable factor incompatibility
translokatierte Inkompatibilität
transportierte Inkompatibilität
Goldschmidt EE (2014) Plant grafting: new mechanisms, evolutionary implications. Frontiers in Plant Science 5:727 Errea P (1998) Implications of phenolic compounds in graft incompatibility in fruit tree species. Scientia Horticulturae 74:195-205
Translocated Incompatibility Disposition
Chardonnay in Champagne shows transmitting tissue navigation during early-mid June Year N+1 at BBCH 60-65, with pollen tubes traversing 16-22 mm style length through extracellular matrix at 1.2-2.2 mm per hour, requiring 12-26 hours to complete navigation from stigma region to ovary, utilizing style nutritional support and guidance cues, under cool maritime conditions of 19-24°C temperature, adequate boron 11-17 ppm supporting sustained tube wall integrity, typical of this early-bursting cultivar with 2-4 tubes successfully completing navigation per flower
Transmitting tissue navigation represents the main tube growth phase when pollen tube elongates through style extracellular matrix following specialized conducting tissue pathway from stigma region toward ovary, utilizing nutritional support and guidance cues enabling directed growth over 12-25 mm typical style length. This essential navigation phase occurs during pollen tube growth in early-mid June in temperate regions Year N+1, typically spanning 10-30 hours as tube traverses style depending on length and conditions, corresponding to BBCH 60-65 flowering stages. Navigation mechanism involves tube growing through extracellular matrix between transmitting tissue cells, following chemical and possibly physical guidance cues directing growth toward ovary, utilizing nutritional support from tissue secretions maintaining growth over extended distance, and maintaining sustained tip growth at 1-3 mm per hour traversing total style length. Transmitting tissue structure supporting navigation includes specialized parenchyma cells lining extracellular spaces creating tube growth pathway, abundant extracellular matrix providing physical scaffold and growth medium, secretory cells releasing nutrients, lipids, and possibly guidance signals into matrix, and overall tissue architecture creating optimized environment for pollen tube growth. Extracellular matrix composition includes pectins creating hydrated gel-like growth medium, arabinogalactan proteins possibly mediating tube-matrix interactions and providing guidance, lipid-rich secretions providing energy source and possibly affecting tube surface properties, calcium maintaining appropriate matrix properties and possibly creating guidance gradients, and various small molecules potentially serving signaling or nutritive functions. Growth pathway characteristics show tube growing in extracellular spaces between transmitting tissue cells, utilizing preformed pathway along style central axis, maintaining relatively straight trajectory from stigma toward ovary base, and path length 12-25 mm in typical grapevine flowers corresponding to style length. Growth rate during navigation shows peak velocity 2-3 mm per hour during mid-style growth phase, sustained growth maintained over many hours through nutritional support and metabolic capacity, and slight rate variation possible with initial acceleration after penetration and possible deceleration approaching ovary. Tube-matrix interactions show tube surface contacting extracellular matrix components, possible adhesion mediating growth direction and stability, matrix providing mechanical support preventing tube buckling during growth, and chemical interactions possibly affecting tube growth properties. Nutritional support from transmitting tissue shows specialized cells secreting nutrients into extracellular matrix, lipid secretions providing energy-rich compounds supporting tube metabolism, possible sugar secretion supplementing pollen grain reserves, and overall tissue support enabling sustained growth over 10-30 hour navigation period. Guidance mechanisms directing navigation include chemotropism with tube responding to concentration gradients of attractant molecules, possible calcium gradients attracting growing tubes, other signaling molecules potentially including GABA or nitric oxide though poorly characterized in grapevine, gravity possibly contributing minor orientation cue, and tissue architecture providing physical pathway constraining growth direction. Chemotropic signals possibly include ovule-derived attractants diffusing through tissue creating concentration gradient, style tissue secretions affecting growth direction, and calcium gradients established in matrix though specific signals in grapevine largely unknown. Cytoplasmic streaming support shows continued basipetal flow at 2-8 micrometers per second maintaining nutrient supply from grain through extending tube to growing tip, streaming essential for supporting growth over long distances exceeding grain diameter by 500-1000 fold, and actomyosin motor system driving streaming throughout navigation period. Sperm cell position shows two sperm cells traveling in tube cytoplasm typically near tip region, cells carried toward apex by streaming, and positioning preparing for delivery to embryo sac upon ovary arrival. Resource utilization during navigation shows initial dependence on pollen grain reserves during early growth, transition to mixed grain-reserves and style-derived nutrients during mid-navigation, and increasing reliance on style nutritional support during extended growth particularly final approach to ovary. Boron requirement during navigation emphasizes continued absolute importance with boron essential for tube wall integrity throughout growth period, deficiency causing tube bursting or growth cessation at any point during navigation preventing ovary arrival, adequate boron 10-20 ppm in style tissues supporting sustained growth maintaining wall structure over many hours. Calcium role in navigation includes maintaining appropriate extracellular matrix properties, possible involvement in guidance through gradient formation, and calcium-dependent processes regulating tube growth though mechanisms incompletely understood. Temperature effects on navigation rate show optimal 24-28°C supporting rapid 2-3 mm per hour navigation completing typical 18 mm style in 6-12 hours, cool conditions 18-22°C substantially slowing to 1-1.5 mm per hour requiring 12-24 hours, warm conditions 28-32°C potentially accelerating growth though excessive heat above 33°C may impair sustained function. Style length variation effects show shorter styles 12-18 mm enabling faster ovary arrival within 6-15 hours, longer styles 22-30 mm requiring extended 12-30 hours, and length variation among cultivars and individual flowers affecting fertilization timing and competitive dynamics. Multiple tube navigation shows 10-50 tubes potentially entering style and beginning navigation, competitive growth with fastest tubes having advantage in ovule access, typical 2-5 tubes reaching ovary region with usually 1-2 achieving fertilization, and slower tubes becoming irrelevant as faster tubes complete fertilization. Tube-tube interactions during navigation poorly characterized but possibly include physical contact or interference, potential chemical signaling, and competitive dynamics for limited ovule access. Navigation failures during transit include boron deficiency causing tube bursting midway through style, resource depletion preventing continued growth before ovary reached, environmental stress disrupting growth processes, tube structural failure from inadequate wall synthesis, or guidance failure causing misdirected growth. The process represents second aspect of pollen tube growth emphasizing main transit through style, following style penetration and preceding ovary approach. Duration typically spans 10-30 hours representing majority of total tube growth time, with faster 8-18 hours under optimal conditions and slower 18-40 hours under suboptimal conditions. Cultivar variation in navigation may include style length affecting required time and resources, transmitting tissue support affecting growth rate, and tube growth vigor affecting navigation success and speed. Regional timing shows transmitting tissue navigation occurring during flowering period across all climates over 10-30 hours following style penetration, representing main tube growth interval. Viticultural significance includes representing main distance-covering growth phase essential for fertilization, requiring adequate style nutritional support and boron nutrition, demonstrating extreme temperature sensitivity affecting navigation rate and timing, and establishing navigation success as direct determinant of ovary arrival enabling fertilization and fruit set. Research applications include analyzing navigation mechanisms including guidance cues and nutritional support, investigating transmitting tissue structure and function supporting tube growth, evaluating boron and other nutritional effects on sustained growth, quantifying environmental effects particularly temperature on navigation dynamics, developing visualization techniques observing tube navigation in vivo, and assessing navigation success as determinant of fertilization timing and outcomes.
Transmitting_Tissue_Navigation is second aspect of Pollen_Tube_Growth, distinguished by main transit through style conducting tissue. Represents majority of total tube growth distance and time. Duration 10-30 hours traversing 12-25 mm style length. Peak growth rate 2-3 mm per hour during mid-navigation. Critical boron dependence throughout. Temperature strongly affects rate. Nutritional support from style tissue essential. Followed by Ovary_Approach as destination nears. Main determinant of fertilization timing.
Leitgewebe-Durchquerung
Schlauchwachstum durch Griffel
Style-Gewebe-Navigation
Transmissionsgewebe-Navigation
conducting tissue navigation
style tissue navigation
transmitting tract traverse
tube growth through style
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 386-390 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 292-295 Carmona MJ et al. (2008) Grapevine inflorescence and flower development, Adv Bot Res 47:301-337
Transmitting Tissue Navigation
Whole-vine transpiration rates in grapevines range from 15-60 L/day depending on genotype, canopy size, and atmospheric demand, with stomatal transpiration accounting for 90-98% of total water loss under well-watered conditions. Leaf-level transpiration rates of 3-12 mmol H₂O/m²/s occur under midday conditions, with high-transpiring cultivars such as Syrah and Grenache at the upper range and conservative types like Chardonnay and Riesling at the lower range. Canopy transpiration follows a diurnal pattern peaking at solar noon and declining through the afternoon as stomata respond to increasing vapor pressure deficit. Transpiration per unit leaf area declines 40-70% under moderate water stress (predawn water potential -0.4 to -0.8 MPa) through stomatal closure.
A water relations disposition that is the inherited capacity of a Vitis vinifera plant to lose water vapor from leaf surfaces to the atmosphere through stomatal and cuticular pathways, with genotype-specific rates determined by stomatal density, aperture regulation, cuticular permeability, and canopy architecture influencing boundary layer conductance.
Transpiration_Disposition serves as the parent class for stomatal conductance and cuticular transpiration traits. Transpiration is the dominant component of vineyard evapotranspiration (60-85% of ET), with the remainder from soil evaporation. The process drives nutrient transport, enables evaporative cooling, and creates the water potential gradient for xylem sap ascent. Transpiration management through irrigation, canopy manipulation, and cultivar selection is central to vineyard water balance in both rain-fed and irrigated systems.
Transpiration
Verdunstungs-Disposition
Wasserverlust
evapotranspiration trait
transpirational flux disposition
water loss capacity
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 215-235 Williams LE, Ayars JE (2005) Grapevine water use and the crop coefficient are linear functions of the shaded area measured beneath the canopy. Agricultural and Forest Meteorology 132:201-211
Transpiration Disposition
Transpiration efficiency in grapevines ranges from 3-7 g above-ground dry matter per kg H₂O transpired over the growing season, with high-efficiency cultivars such as Tempranillo and Mourvèdre at 5-7 g/kg and lower-efficiency types like Grenache at 3-5 g/kg. Rootstock effects on scion transpiration efficiency reach 15-30%, with 110 Richter and 140 Ruggeri conferring higher efficiency than SO4 and 1103 Paulsen through moderate stomatal conductance and conservative water use. Deficit irrigation improves transpiration efficiency by 20-50% compared to full irrigation, as moderate stress reduces transpiration proportionally more than biomass production. Night-time transpiration (5-15% of daily total) represents a water loss with no carbon gain, reducing whole-plant transpiration efficiency.
A water use efficiency disposition that is the inherited capacity of a Vitis vinifera plant to produce biomass per unit of water transpired over extended time periods (days to seasons), integrating instantaneous gas exchange efficiency with respiratory carbon losses, non-productive water use, and the temporal dynamics of growth and transpiration across variable environmental conditions.
Transpiration efficiency captures the net outcome of water-for-carbon exchange over agronomically relevant time scales, integrating the multiple processes affecting WUE into a single production metric. The trait differs from instantaneous WUE by including respiratory losses and non-stomatal water loss. High transpiration efficiency is achieved through combinations of moderate stomatal conductance, high mesophyll conductance, efficient Rubisco carboxylation, and low respiratory costs. Partitioning of biomass between vegetative growth and fruit affects the agronomic relevance of transpiration efficiency for yield outcomes.
Biomasseproduktivität pro Transpiration
Transpirations-Effizienz
biomass water productivity
saisonale Wassernutzungseffizienz
seasonal WUE
transpiration-based efficiency
Tomás M et al. (2012) Variability of mesophyll conductance in grapevine cultivars under water stress conditions in relation to leaf anatomy and water use efficiency. Australian Journal of Grape and Wine Research 18:164-172 Escalona JM et al. (2012) Responses of leaf night transpiration to drought stress in Vitis vinifera L. Agricultural Water Management 118:50-58
Transpiration Efficiency Disposition
Transpiration efficiency comparison between deficit-irrigated and fully irrigated Cabernet Sauvignon reveals the water stress effect on water use strategy, with deficit irrigation achieving integrated WUE of 4.8 kg fruit/m³ water (moderate stress improving efficiency through partial stomatal closure) compared to 3.2 kg/m³ under full irrigation (maximum productivity but lower efficiency), demonstrating the management trade-off between absolute yield and water productivity that deficit irrigation exploits to produce quality fruit with reduced water consumption in water-limited regions.
Transpiration efficiency quality is a physiological quality that represents the ratio of carbon gain to water loss, characterizing the effectiveness with which grapevines convert transpired water into fixed carbon that determines the sustainability of production under water-limited conditions and the water footprint of viticulture. Within the BFO framework, this quality addresses the coupling efficiency dimension of carbon-water relations, distinguishing the ratio of these fluxes from their absolute magnitudes while recognizing that transpiration efficiency determines how much water is required to produce a given amount of biomass or yield, with profound implications for vineyard water management, regional water allocation, and climate change adaptation. The ontological significance lies in transpiration efficiency providing the integrating metric for vine water use strategy, where the ratio of photosynthetic carbon fixation to transpirational water loss establishes the water cost of carbon gain that determines both the productivity achievable per unit water consumed and the sustainability of grape production in water-scarce environments that characterize most viticultural regions globally. The physiological basis for transpiration efficiency involves the shared stomatal pathway for CO₂ uptake and water vapor loss, with the ratio of these fluxes determined by the relative driving forces (CO₂ gradient for photosynthesis, water vapor gradient for transpiration) and the relative conductances of the pathways, creating an intrinsic coupling that can be modified through stomatal behavior but not fundamentally overcome within C₃ photosynthesis constraints. The multiple scales of transpiration efficiency assessment include intrinsic water use efficiency (A/gs, the ratio of assimilation to stomatal conductance revealing instantaneous leaf-level coupling), instantaneous water use efficiency (A/E, incorporating actual transpiration rates), and integrated water use efficiency (biomass or yield per water consumed, reflecting long-term productive efficiency), with each scale providing different insights into vine water use strategy. The environmental and management factors affecting transpiration efficiency include atmospheric humidity (lower VPD increasing efficiency by reducing transpiration at equivalent conductance), CO₂ concentration (elevated CO₂ improving efficiency through enhanced photosynthesis per unit conductance), stomatal behavior (more conservative closure improving efficiency at the cost of absolute productivity), and agronomic practices (deficit irrigation, canopy management affecting the conditions under which gas exchange occurs). Understanding transpiration efficiency quality enables assessment of vine water use strategy, optimization of irrigation for maximum production per unit water, and evaluation of cultivar suitability for water-limited production environments.
Transpiration_Efficiency_Quality serves as the parent class for specific efficiency measures (Intrinsic, Instantaneous, Integrated WUE). This quality provides the framework for assessing the carbon-water coupling that determines water productivity. The class supports irrigation optimization and water management through comprehensive efficiency evaluation across scales.
Kohlenstoff-Wasser-Kopplung
Transpirationseffizienz
WUE
Wassernutzungseffizienz
carbon-water coupling
transpiration ratio
water use efficiency
Medrano H et al. (2015) From leaf to whole-plant water use efficiency (WUE) in complex canopies: limitations of leaf WUE as a selection target. The Crop Journal 3(3): 220-228 Tomás M et al. (2014) Variability of intrinsic water use efficiency among grapevine varieties and its association with grape yield and quality. Australian Journal of Grape and Wine Research 20(2): 259-270
Transpiration Efficiency Quality
Transpiration rate monitoring in deficit-irrigated Tempranillo tracks the daily water demand cycle, with pre-dawn rates near zero (stomata closed, minimal cuticular loss), morning increase to 4.5 mmol H₂O/m²/s as stomata open under moderate VPD, midday peak of 8.2 mmol/m²/s under high radiation and VPD (1.8 kPa), afternoon decline as VPD decreases and soil water becomes limiting, and evening return to minimal rates, with daily integration yielding approximately 4.5 L/vine water consumption that must be replaced through irrigation or rainfall to maintain vine water status.
Transpiration rate quality is a physiological quality that represents the rate of water vapor loss from grapevine tissues to the atmosphere, characterizing the water flux that drives nutrient transport, enables evaporative cooling, and constitutes the primary component of vine water demand that must be balanced against soil water supply for optimal physiological function. Within the BFO framework, this quality addresses the water loss dimension of vine physiology, distinguishing vapor phase water movement from liquid water transport while recognizing that transpiration is inextricably coupled to CO₂ uptake through shared stomatal pathways, creating the fundamental trade-off between carbon gain and water loss that defines plant water use strategy. The ontological significance lies in transpiration rate determining vine water demand and the coupling between carbon and water economies, where the water lost through transpiration must be replaced from soil reserves through root uptake, and where the ratio of carbon gained to water lost (water use efficiency) establishes the sustainability of production under water-limited conditions that characterize most viticultural regions. The physical basis for transpiration involves the diffusion of water vapor down a concentration gradient from saturated internal leaf air spaces (water potential near zero) to the drier atmosphere (water potential strongly negative depending on humidity), with the rate determined by the driving force (vapor pressure deficit) and the resistance of the diffusion pathway (primarily stomatal, with minor cuticular component). The physiological functions of transpiration extend beyond simple water loss to include nutrient transport (mass flow of dissolved minerals in the transpiration stream), evaporative cooling (latent heat removal preventing heat stress), and maintenance of cell turgor (continuous water flow supporting hydraulic architecture), making transpiration essential for vine function despite its water cost. The environmental and physiological factors affecting transpiration rate include vapor pressure deficit (atmospheric water demand), stomatal conductance (actively regulated pathway resistance), wind speed (boundary layer effects), available energy (radiation driving evaporation), and soil water availability (limiting supply when depleted), with transpiration rate reflecting the integration of atmospheric demand, plant regulation, and soil supply. Understanding transpiration rate quality enables assessment of vine water demand, irrigation scheduling based on transpiration-driven depletion, and evaluation of the carbon-water trade-off that determines water use efficiency under different management scenarios.
Transpiration_Rate_Quality serves as the parent class for transpiration pathway components (Stomatal, Cuticular) and scaling (Canopy). This quality represents the water cost of carbon gain that determines water demand and use efficiency. The class supports irrigation management and water use assessment through transpiration rate characterization.
E-Wert
Evapotranspiration
Transpirationsrate
evapotranspiration
water loss rate
water vapor flux
Escalona JM et al. (1999) Stomatal and non-stomatal limitations of photosynthesis under water stress in field-grown grapevines. Australian Journal of Plant Physiology 26(5): 421-433 Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell & Environment 26(8): 1393-1405
Transpiration Rate Quality
Transpirational cooling capacity quality assessment in Central Valley California evaluated the interaction between irrigation management and heat avoidance in Cabernet Sauvignon during summer heat events. Leaf temperature measurements using infrared thermometry compared vines under full irrigation (100% ETc replacement) versus regulated deficit irrigation (50% ETc) during a 3-day heat event with ambient temperatures reaching 43°C. Fully irrigated vines demonstrated excellent transpirational cooling capacity quality with midday leaf temperatures averaging 35.2°C (7.8°C below ambient), sustained high stomatal conductance averaging 285 mmol m⁻² s⁻¹, and transpiration rates of 8.5 mmol m⁻² s⁻¹ providing latent heat dissipation equivalent to 510 W m⁻² of leaf surface. Deficit-irrigated vines showed compromised transpirational cooling capacity quality with leaf temperatures averaging 40.8°C (only 2.2°C below ambient), reduced stomatal conductance of 85 mmol m⁻² s⁻¹ due to drought-induced stomatal closure, and transpiration of only 2.8 mmol m⁻² s⁻¹ providing just 170 W m⁻² cooling capacity. The transpirational cooling capacity quality differential translated directly to heat damage, with fully irrigated vines showing only 8% sunburn while deficit-irrigated vines exhibited 35% damage, establishing that maintaining irrigation during heat events was critical for transpirational cooling even in systems normally managed for moderate water stress, and informing protocols for suspending deficit irrigation when heat events are forecast.
Transpirational cooling capacity quality is a heat avoidance quality that inheres in a grapevine organism and represents the ability to dissipate heat through evaporative water loss from leaf and fruit surfaces, with the latent heat absorbed during water vaporization providing a powerful cooling mechanism that can reduce tissue temperatures substantially below ambient air temperature when adequate water supply permits sustained high transpiration rates during heat exposure. This quality addresses the primary physiological mechanism for active heat avoidance in grapevines, recognizing that transpirational cooling provides the dominant component of tissue temperature regulation under most high-temperature conditions, with cooling capacity determined by stomatal conductance, leaf area, boundary layer characteristics, and the availability of water to support continued transpiration. The ontological significance of transpirational cooling capacity quality lies in its role as the most effective heat avoidance mechanism available to grapevines, capable of achieving tissue temperature reductions of 10-15°C below ambient under optimal conditions, effectively extending the environmental temperature range within which vines can maintain function when water availability is sufficient to sustain high transpiration rates. The biological basis of transpirational cooling encompasses the physics of evaporative heat transfer where the latent heat of vaporization (approximately 2.4 MJ kg⁻¹ water) provides substantial cooling for each gram of water transpired, combined with the physiological regulation of stomatal aperture that controls transpiration rate, and the hydraulic architecture determining water supply capacity to transpiring surfaces. The effectiveness of transpirational cooling depends critically on water availability, with soil moisture depletion or root zone limitations reducing the water supply that can support transpiration, causing stomatal closure that diminishes cooling precisely when high temperatures create the greatest cooling demand. The interaction between transpirational cooling and drought stress creates a critical management challenge, as deficit irrigation practices that restrict water supply to enhance fruit quality may simultaneously compromise transpirational cooling capacity, potentially causing heat damage that would not occur with more generous irrigation during heat events.
Transpirational_Cooling_Capacity_Quality is_a Heat_Avoidance_Quality, representing the dominant physiological mechanism for tissue temperature reduction through evaporative heat loss. This class addresses the cooling achieved through water transpiration, critically dependent on water availability. Cooling magnitude (often 5-15°C below ambient) and latent heat dissipation (W m⁻²) provide quantification. The interaction with irrigation/drought status creates important management considerations. Stomatal conductance under heat stress is the proximate determinant of cooling capacity. Distinguished from Leaf_Angle_Adjustment_Quality and Reflectance_Increase_Quality which are water-independent avoidance mechanisms.
Verdunstungskühlung
evaporative cooling
latent heat dissipation
latente Wärmeabfuhr
stomatal cooling
stomatäre Kühlung
transpiration cooling
transpirative Kühlungskapazität
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 453-507 Greer DH, Weedon MM (2013) The impact of high temperatures on Vitis vinifera cv. Semillon grapevine performance and berry ripening. Frontiers in Plant Science 4:491 Caravia L et al. (2016) Application of shade treatments during Shiraz berry ripening to reduce the impact of high temperature. Australian Journal of Grape and Wine Research 22:422-437
Transpirational Cooling Capacity Quality
Transport tolerance quality evaluation in South Australia compared quality outcomes for Shiraz and Viognier across controlled transport duration treatments simulating varied vineyard-winery distances. Shiraz demonstrated excellent transport tolerance quality with thick skin, firm flesh, and low oxidation susceptibility permitting transport durations up to 8 hours at ambient autumn temperatures (18-24°C) without detectable quality impact on resulting wines, with even 12-hour transport showing only marginal quality reduction of 2 sensory points attributed to slight volatile loss and elevated volatile acidity from early fermentation onset. Viognier exhibited poor transport tolerance quality with thin skin susceptible to progressive damage under bin compression, high juice oxidation potential causing rapid browning once skin integrity was breached, and characteristic aromatic compounds showing measurable degradation with 3-hour transport and substantial loss by 6 hours, with wines from extended transport receiving 12 points lower sensory scores dominated by oxidative and cooked fruit character rather than varietal floral notes. The transport tolerance quality differential established maximum transport recommendations of 8 hours for Shiraz versus 2 hours for Viognier, with refrigerated transport extending Viognier tolerance to 5 hours at substantial cost, demonstrating how transport tolerance quality constraints influence practical sourcing and logistics decisions.
Transport tolerance quality is a harvest logistics quality that inheres in a grapevine cultivar or clone and represents the capacity of harvested grape material to withstand the extended stresses of transport from vineyard to winery including continued compression in bins, temperature elevation from solar exposure and metabolic heat generation, vibration and impact from vehicle movement over varied road surfaces, and the cumulative duration of exposure to ambient conditions before processing commences, while maintaining compositional and structural integrity adequate for target wine quality production. This quality extends the integrity concept beyond the immediate harvest operation to encompass the entire vineyard-to-winery logistics chain that may span from minutes for estate wineries to many hours for contracted fruit transported substantial distances, with transport tolerance determining whether extended logistics are feasible without quality compromise or whether rapid processing is essential to capture the quality potential harvested from the vineyard. The ontological significance of transport tolerance quality lies in its role determining the geographic and temporal flexibility available for fruit sourcing and processing logistics, wherein high transport tolerance enables centralized processing of fruit from dispersed vineyard sources, deferred crushing accommodating winery capacity constraints, and long-distance transport supporting specialized production facilities, while poor transport tolerance restricts viable logistics to rapid, proximate processing that limits operational flexibility. The biological basis of transport tolerance quality encompasses the structural factors determining resistance to progressive damage under sustained compression and vibration the metabolic characteristics influencing heat generation and respiratory rate during the transport interval the oxidation susceptibility determining quality decline from oxygen exposure through damaged berry surfaces the microbial populations on fruit surfaces and their proliferation potential under transport temperature conditions and the compositional stability of quality-determining compounds under the combined stresses of the transport environment. Cultivar differences in transport tolerance reflect variation in skin thickness, berry firmness, juice oxidation potential, and aromatic compound stability, while harvest conditions including fruit temperature at picking and bin fill depth substantially influence transport outcomes for any cultivar. Assessment of transport tolerance quality involves controlled transport simulation trials measuring quality parameter changes across varied duration and condition treatments, comparison of wine outcomes from rapidly versus delayed-processed fruit, and identification of transport condition thresholds beyond which quality compromise becomes significant.
Transport_Tolerance_Quality is_a Harvest_Logistics_Quality, representing the capacity to withstand extended post-harvest logistics without quality compromise. This class addresses the vineyard-to-winery interval extending beyond the immediate harvest operation. Duration tolerance in hours under specified temperature conditions provides practical metric. Skin thickness, oxidation susceptibility, and aromatic stability are key biological determinants. Distinguished from Fruit_Integrity_At_Harvest_Quality by the extended duration and additional stresses of transport. Refrigerated transport can extend tolerance but at cost requiring economic evaluation. Critical for production models involving fruit transport from distant vineyards to centralized processing facilities.
Logistik-Beständigkeit
Nachernte-Stabilität
Transportresilienz
Transporttoleranz
logistics tolerance
post-harvest stability
pre-processing durability
transport resilience
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1: The Microbiology of Wine and Vinifications, 2nd Edition. John Wiley and Sons, Chichester Singleton VL et al. (1980) Grape and wine phenolics: background and prospects. In: Proceedings of the University of California, Davis, Grape and Wine Centennial Symposium, pp. 215-227 Jackson RS (2008) Wine Science: Principles and Applications, 3rd Edition. Academic Press, pp. 270-331
Transport Tolerance Quality
Trellis adaptation quality evaluation for Tempranillo across Spanish wine regions compared performance under traditional bush vine (gobelet), single Guyot, bilateral cordon, and vertical shoot positioning configurations. The cultivar demonstrated broad trellis adaptation quality, achieving acceptable vine balance and fruit quality across all systems with Ravaz Index ranging from 5.8 in VSP to 7.2 in bush vine, though with systematic quality-yield tradeoffs wherein bush vine produced highest concentration fruit at lowest yield while VSP achieved highest productivity with adequate but reduced concentration. This broad adaptation supported regional flexibility in training system selection based on production objectives, mechanization requirements, and water availability. Contrasting evaluation of Mencía revealed narrow trellis adaptation quality, with excellent performance under traditional gobelet (Ravaz 6.5, optimal fruit concentration) but poor outcomes in VSP (Ravaz 11.2, excessive yield, dilute fruit) and inadequate vigor for GDC curtain fill, restricting this cultivar to traditional training approaches for quality production.
Trellis adaptation quality is a viticultural quality that inheres in a grapevine cultivar or clone and represents the capacity to achieve optimal performance across the range of specific trellis and training system configurations employed in commercial viticulture, encompassing the cultivar's ability to conform to diverse architectural requirements, exploit the environmental conditions created by different canopy geometries, and respond productively to the pruning and management regimes associated with particular training approaches. This quality provides integrative assessment of cultivar versatility or specificity regarding training system requirements, distinguishing broadly adaptable genotypes suitable for diverse production contexts from those exhibiting narrow adaptation requiring specific training configurations to achieve acceptable performance. The ontological significance of trellis adaptation quality lies in its role as a practical selection criterion enabling informed decisions about cultivar deployment across viticultural regions employing different traditional or modern training systems, and about training system selection for new plantings where cultivar choice precedes infrastructure decisions. The biological basis of trellis adaptation quality encompasses the interaction between multiple cultivar characteristics and training system requirements, including growth habit compatibility with canopy geometry, vigor level matching to system capacity, fruiting habit alignment with pruning approach, cluster architecture suitability for the microclimate conditions created by different configurations, and fruit composition sensitivity to the light and temperature regimes characteristic of specific canopy structures. Cultivars exhibiting broad trellis adaptation quality demonstrate acceptable performance across vertical, divided, sprawling, and traditional training configurations, providing flexibility for producers operating diverse vineyard holdings or adapting to changing market demands and mechanization requirements. Conversely, cultivars with narrow adaptation may achieve exceptional performance under optimal training but fail under alternative configurations, requiring careful matching of genotype to training infrastructure. The assessment of trellis adaptation quality involves multi-environment trials comparing cultivar performance across training systems under standardized management, supplemented by mechanistic evaluation of how specific cultivar traits interact with training system characteristics to predict adaptation potential for untested combinations. Regional cultivar recommendations increasingly incorporate trellis adaptation quality assessment to guide training system selection beyond historical precedent.
Trellis_Adaptation_Quality serves as the parent class for specific training system suitability subclasses including VSP_Suitability_Quality, GDC_Suitability_Quality, Lyre_Suitability_Quality, Pergola_Suitability_Quality, Gobelet_Suitability_Quality, Guyot_Suitability_Quality, and Cordon_Suitability_Quality. This class addresses overall adaptability across training systems while child classes capture suitability for specific configurations. Distinguished from Training_System_Suitability_Quality which addresses the general concept of cultivar-training compatibility, while this class specifically concerns trellis/training configuration adaptation. Provides framework for systematic cultivar-training matching decisions.
Erziehungssystem-Adaptabilität
Laubwandkonfigurationsflexibilität
Spalieranpassungsfähigkeit
Spaliervielseitigkeit
canopy system adaptation
training configuration flexibility
training system adaptability
trellis versatility
Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review. American Journal of Enology and Viticulture 60:251-268 Jackson DI, Lombard PB (1993) Environmental and management practices affecting grape composition and wine quality: a review. American Journal of Enology and Viticulture 44:409-430 Intrieri C, Poni S (1995) Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality of mechanized Italian vineyards. American Journal of Enology and Viticulture 46:116-127
Trellis Adaptation Quality
The trellis design description for a Smart-Dyson ballerina system in a Shiraz vineyard in McLaren Vale specifies treated pine end posts (125 mm diameter, 2.7 m length, buried 0.8 m) with H-brace anchoring, galvanized steel intermediate posts (1.8 m star pickets at 5 m intervals), a single cordon wire at 1.2 m, a divided upper canopy created by twin foliage wire sets angled at 30° from vertical using 0.6 m cross-arms at 1.5 m height, pendant shoot positioning on the outer curtains, and a total trellis height of 2.0 m — the divided canopy design effectively doubles the exposed canopy surface area compared to standard VSP, improving light interception in the vigorous McLaren Vale growing conditions.
A training system description that documents the physical support structure — posts, wires, anchors, and associated hardware — upon which the grapevine is trained and its canopy supported, specifying the structural engineering parameters, materials, and spatial configuration of the trellis framework that determines the geometric envelope within which vine growth is managed.
Trellis_Design_Description captures the engineering dimension of vineyard architecture that translates viticultural concepts into physical infrastructure. The trellis is the most capital-intensive and permanent component of vineyard establishment after land preparation and planting, with installation costs ranging from 3,000-15,000 EUR/ha depending on materials and complexity: trellis design determines the mechanical loads the structure must support (mature vine weight of 5-15 kg per vine plus fruit load plus wind forces plus worker/machine weight during operations), the corrosion resistance required for the target vineyard life (25-50 years), and the compatibility with mechanized operations (machine harvesting, pre-pruning, leaf removal). Modern trellis designs must accommodate both traditional manual viticulture workflows and increasingly automated precision viticulture technologies including sensor-equipped mechanical harvesters and autonomous vineyard robots.
Drahtrahmen-Dokumentation
Spalierdesign-Beschreibung
Stützsystem-Spezifikation
support system description
trellis structure documentation
vineyard infrastructure specification
Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles pp: 61-90 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 66-96 Reynolds AG (ed) (2015) Grapevine Breeding Programs for the Wine Industry, Woodhead Publishing
Trellis Design Description
A Regent × Muscaris F1 seedling at INRAE Colmar bearing a trial cultivar role is planted as one of 2,500 seedlings in a Phase 1 screening trial with single-vine unreplicated plots, assessed in its 3rd-5th leaf for downy mildew resistance through natural infection scoring at 3 observation dates per season, powdery mildew resistance under unsprayed conditions, basic phenology (budburst, flowering, véraison dates), and preliminary fruit quality (refractometer Brix at harvest), with 85-90% of genotypes eliminated at this stage and only 250-350 advancing to Phase 2 replicated trials based on combined resistance and quality scores exceeding predefined selection indices.
An experimental cultivar role that inheres in a grapevine genotype undergoing initial or intermediate-stage field screening within a breeding program or cultivar evaluation pipeline, where the primary objective is elimination of inferior genotypes through assessment against minimum performance thresholds before advancement to more resource-intensive evaluation stages.
Trial_Cultivar_Role describes the early-stage experimental status where genotypes are numerous, replication is minimal or absent, and the evaluation objective is efficient elimination rather than precise characterization. Grapevine breeding programs typically screen 5,000-20,000 seedlings per cross to identify 1-3 candidates worthy of cultivar release: the trial stage encompasses the initial 3-8 years of field observation that reduce this population by 90-99%. The role is transitional — surviving genotypes advance to Evaluation_Cultivar_Role for more rigorous multi-site assessment, while eliminated genotypes are removed from the experimental pipeline.
Selektions-Versuchs-Rolle
Versuchsrebsorten-Rolle
Vorprüfungs-Rolle
preliminary evaluation role
screening trial role
seedling trial role
Eibach R and Töpfer R (2015) Traditional grapevine breeding techniques, in Reynolds AG (ed) Grapevine Breeding Programs for the Wine Industry, Woodhead Publishing pp: 3-22 Töpfer R et al. (2011) New horizons for grapevine breeding, Fruit, Vegetable and Cereal Science and Biotechnology 5: 79-100 Schneider C et al. (2019) Grapevine breeding programs in France, in Ferrão MAG et al. (eds) Genetic Improvement of Tropical Crops, Springer pp: 301-344
Trial Cultivar Role
Evaluation of Trichome_Density_Disposition effects on arthropod resistance at Cornell University characterized trichome parameters and pest responses across 28 cultivars spanning the pubescence range in cultivated Vitis. Trichome density was quantified by microscopy on abaxial leaf surfaces at two positions (near midrib, near margin), with counts expressed as trichomes per mm². Cultivars ranged from essentially glabrous ('Chardonnay', 4 trichomes/mm²) to densely tomentose ('Concord', 85 trichomes/mm²), with intermediate types including 'Cabernet Sauvignon' (22/mm²) and 'Gewürztraminer' (48/mm²). Potato leafhopper (Empoasca fabae) colonization showed strong negative correlation with Trichome_Density_Disposition (r = -0.82), with choice tests confirming active avoidance of pubescent surfaces. Hopperburn damage severity correlated similarly (r = -0.78), with tomentose 'Concord' (85/mm²) showing no detectable damage while glabrous 'Chardonnay' (4/mm²) sustained 65% leaf area affected. For grape leafhopper (Erythroneura comes), the relationship was weaker (r = -0.45), suggesting partial but incomplete deterrence. Importantly, trichome density showed positive correlation with Typhlodromus pyri predatory mite abundance (r = 0.58), indicating that pubescent cultivars support enhanced biological control of spider mites. These findings demonstrate that Trichome_Density_Disposition provides broad protection against multiple pest groups while enhancing natural enemy habitat.
A Trichome_Density_Disposition is a Physical_Barrier_Disposition that inheres in a Vitis vinifera plant and is realized through the development of dense epidermal trichomes (plant hairs) on leaf surfaces and other organs, providing physical impediment to arthropod pest movement and feeding, modification of leaf surface microclimate, and in some cases, chemical defense through glandular secretions. This disposition represents the quantitative development of trichomes, with higher densities generally providing greater protection against small arthropod pests including leafhoppers, spider mites, and thrips that must navigate the trichome forest to access feeding sites. Grapevine trichomes are predominantly non-glandular, consisting of simple unicellular or multicellular hairs varying in length, density, and distribution among organs, surfaces (adaxial versus abaxial), and genotypes. Trichome density on abaxial (lower) leaf surfaces shows particularly wide genetic variation in Vitis, from nearly glabrous cultivars to densely tomentose types with felt-like pubescence. Dense trichomes impede pest movement by creating physical obstacles, interfere with stylet insertion by small piercing-sucking insects, modify boundary layer humidity affecting pest development and pathogen germination, and provide refugia for predatory mites contributing to biological control. Conversely, dense trichomes may create favorable microhabitats for certain pests and pathogens, and the relationship between trichome density and defense varies with the specific attacker. Trichome_Density_Disposition is readily observable and highly heritable, making it amenable to selection in breeding programs targeting arthropod resistance.
Trichome_Density_Disposition is_a Physical_Barrier_Disposition. Trichomes primarily affect arthropod pests rather than pathogens, with density being the most important parameter for small insect and mite resistance. The effect varies among pest species and may be positive (physical barrier, predator habitat) or negative (microhabitat for certain pests) depending on context. Grapevine trichomes are predominantly non-glandular, distinguishing their function from glandular trichomes in other plant families that produce toxic or sticky secretions.
Blattbehaarung
Epidermale-Haardichte
Trichom-Dichte-Disposition
epidermal hair density
leaf pubescence
trichome development
Hanley ME et al. (2007) Plant structural traits and their role in anti-herbivore defence, Perspectives in Plant Ecology, Evolution and Systematics 8:157-178 Dalin P et al. (2008) Leaf trichome formation and plant resistance to herbivory, Induced Plant Resistance to Herbivory, Springer Karban R et al. (1995) Induced resistance in wild tobacco with clipped sagebrush neighbors, Ecology 76:2165-2169
Trichome Density Disposition
A trifurcated tendril at node 7 on a vigorous shoot of Vitis vinifera cv. Mourvèdre in Bandol, France, exhibiting a true three-way branch point at 5.5 cm from shoot attachment where three arms of 9.5 cm, 10 cm, and 11.5 cm length diverge at approximately 120-degree angles from each other creating a well-balanced radial architecture, all three arms showing comparable diameter (2.6-2.9 mm at mid-length) indicating relatively equal vascular development, each arm exhibiting active circumnutation in different spatial zones with complementary sweep patterns that together explore approximately 580 cubic centimeters volume, two arms making successful contact with trellis wires in perpendicular orientations and both initiating coiling while the third arm continues searching, creating exceptionally stable dual-point attachment supporting the 75 cm shoot section bearing eight leaves and two fruit clusters, measured at BBCH stage 73 (berries beginning to touch) in early July, representing the elevated trifurcation frequency characteristic of Mourvèdre where 25-30% of branched tendrils show three-arm architecture.
A branched tendril exhibiting division into exactly three branches or arms in Vitis vinifera, characterized by presence of either a single branch point where three meristems produce three arms simultaneously, or two sequential branch points creating a three-armed architecture through bifurcation followed by secondary division of one arm, representing a less common but significant branching pattern occurring in 10-30% of branched tendrils in most cultivars with higher frequencies in certain varieties including Syrah, Mourvèdre, and some Muscat types, and providing enhanced contact probability and mechanical redundancy through triple searching capacity at the cost of reduced individual arm vigor due to three-way resource partitioning. Trifurcated tendrils develop through activation of three lateral meristems during early tendril organogenesis creating either a true three-way division at a single point, or through sequential branching where initial bifurcation is followed by secondary division of one arm producing pseudo-trifurcation with two branch levels, with developmental timing, resource availability, and genetic programming determining which pattern occurs and whether arms develop equally or with dominance hierarchies. The functional implications of trifurcation include substantially increased spatial exploration volume compared to bifurcated tendrils (approximately 1.4-1.6× greater search space), higher probability of multiple successful attachments creating mechanically robust support systems with load distribution across multiple points, but potentially reduced individual arm strength and coiling vigor due to resource dilution unless overall tendril vigor is high compensating for three-way partitioning through increased total resource investment.
Trifurcated tendrils are distinguished from bifurcated tendrils by three rather than two arms, and from higher-order multi-branched tendrils by exactly three arms. True trifurcation with a single three-way branch point should be distinguished from pseudo-trifurcation resulting from sequential bifurcations, though both achieve similar functional outcomes. Trifurcation frequency shows strong cultivar specificity serving as an ampelographic diagnostic feature, with some varieties rarely producing three-armed tendrils (20% frequency). Environmental conditions during early development influence trifurcation expression, with high vigor and optimal resource availability favoring increased branching complexity including higher trifurcation rates. The functional value of trifurcation depends on vine vigor, with vigorous vines supporting robust three-arm development while moderate or low vigor may result in weak individual arms compromising functional advantage despite increased arm number.
dreiarmige Kletterorgan
dreifach verzweigte Ranke
dreigabelige Ranke
three-branched tendril
three-way division tendril
tri-arm tendril
triple-forked tendril
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 120-135 This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 155-172.
Trifurcated Tendril
Tropical fruit aroma quantification in Marlborough Sauvignon Blanc correlates sensory passionfruit intensity (8.5 on 10-point scale) with 3MH concentration of 2400 ng/L, demonstrating the thiol-derived nature of this signature character, while comparison with Loire Valley Sancerre (passionfruit 4.2, 3MH 580 ng/L) reveals the 4-fold compositional difference underlying the stylistic distinction between New Zealand's tropical-dominant and Loire's more mineral expression of the same variety.
Tropical fruit aroma quality is a fruity aroma quality that represents olfactory perception evoking exotic tropical fruit-associated scents including passionfruit, guava, mango, pineapple, lychee, and papaya notes, characterizing the lush, heady, often pungent aromatic impression associated with riper fruit character and certain varietal signatures. Within the BFO framework, this quality addresses a distinctive subcategory of fruity aroma distinguished by its association with fuller wine styles and specific varietal expressions, recognizing that tropical character often indicates advanced grape ripeness, specific thiol compound presence, or particular variety identity. The ontological significance lies in tropical aroma serving as the defining varietal marker for several commercially important varieties, most notably the passionfruit and guava character of Marlborough Sauvignon Blanc, the lychee signature of Gewürztraminer, and the tropical elements of Torrontés and Viognier, where tropical fruit intensity and specific note identity enable variety and origin recognition. The compositional basis for tropical aroma is dominated by volatile thiols, particularly 3-mercaptohexan-1-ol (3MH, passionfruit, grapefruit), 3-mercaptohexyl acetate (3MHA, passionfruit, guava), and 4-mercapto-4-methylpentan-2-one (4MMP, boxwood, guava), with these sulfur-containing compounds exhibiting extremely low detection thresholds (ng/L range) that allow trace concentrations to create powerful aromatic impact. Additional contributors include certain terpenes (rose oxide contributing lychee in Gewürztraminer), lactones (coconut-like notes contributing to tropical impression), and esters providing generic tropical background. The ripeness association of tropical character reflects both grape maturation effects on thiol precursor accumulation and the perceptual categorization of tropical fruits as ripe stimuli, creating the impression of fuller, riper wine style even when combined with high acidity. Understanding tropical aroma quality enables varietal authentication based on thiol-derived tropical character, ripeness assessment from tropical fruit development, and style prediction where tropical dominance indicates fuller expression.
Tropical_Fruit_Aroma_Quality addresses the tropical subcategory of fruity aroma, encompassing passionfruit, guava, mango, pineapple, lychee, and related exotic fruit notes. This quality is strongly associated with volatile thiols in Sauvignon Blanc family varieties and with specific terpenes in Gewürztraminer. The tropical character serves as a key varietal and regional marker, with intensity and specific fruit note balance distinguishing origins and styles.
Südfruchtduft
Tropenfruchtaroma
Tropenfruchtcharakter
exotic fruit aroma
exotischer Fruchtduft
tropical fruit character
tropical notes
tropical scent
Tominaga T et al. (1998) A powerful aromatic volatile thiol, 2-furanmethanethiol, exhibiting roast coffee aroma in wines made from several Vitis vinifera grape varieties. Journal of Agricultural and Food Chemistry 48(5): 1799-1802 Lund CM et al. (2009) New Zealand Sauvignon blanc distinct flavor characteristics: sensory, chemical, and consumer aspects. American Journal of Enology and Viticulture 60(1): 1-12
Tropical Fruit Aroma Quality
The true homonym Muscat designates at least 12 genetically unrelated cultivars when used without a qualifying modifier, including Muscat Blanc à Petits Grains (VIVC 08193, the ancient aromatic cultivar), Muscat of Alexandria (VIVC 08222, the large-berried table/wine grape), Muscat Ottonel (VIVC 08241, a 19th-century cross of Chasselas x Muscat de Saumur), and Muscat Hamburg (VIVC 08210, a Muscat of Alexandria x Schiava Grossa cross) — in unqualified commercial usage the bare name Muscat is ambiguous and requires geographic, stylistic, or varietal context for resolution, as when Australian wine labeled simply Muscat from Rutherglen refers to Muscat Blanc à Petits Grains while Muscat from Pantelleria refers to Muscat of Alexandria (locally Zibibbo).
A homonym where two or more genetically distinct grapevine cultivars share an identical name in precisely the same orthographic form, with no distinguishing qualifier in the base designation, requiring molecular identification or geographic context to determine which cultivar is referenced.
True_Homonym captures the strictest form of nomenclatural ambiguity where identical character strings designate distinct genotypes. The problem is most acute with group names historically applied to phenotypically similar but genetically unrelated cultivars sharing a common trait such as the Muscat aroma (caused by monoterpene accumulation, which arose independently in multiple genetic backgrounds) or the Malvasia wine style: molecular analysis has revealed that superficial phenotypic similarity masked profound genetic divergence, with Muscat-type cultivars spanning at least four distinct genetic lineages. The VIVC addresses true homonymy by appending disambiguating qualifiers (berry color, geographic origin, morphological feature) to create unique entries, but unqualified usage persists in commercial and colloquial contexts.
echtes Homonym
exact-name homonym
gleichschreibendes Homonym
orthographic homonym
strict homonym
striktes Homonym
Crespan M and Milani N (2001) The Muscats: a molecular analysis of synonyms, homonyms and genetic relationships, Vitis 40: 79-86 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157 Robinson J et al. (2012) Wine Grapes, Allen Lane pp: 665-680
True Homonym
Cabernet Sauvignon trunk bark at 15 years post-planting shows 6.5-8.5 mm total depth comprising inner bark (living secondary phloem 2.5-3.2 mm with functional sieve tubes and 18-28 g starch in phloem parenchyma) and outer bark (dead tissue 4.0-5.3 mm including 3 successive periderm layers with cork, crushed phloem, and phelloderm remnants), outermost cork showing grey-brown weathered color with longitudinal furrows 2-4 mm deep, lenticels 2-3 mm diameter scattered enabling gas exchange, bark providing waterproof protection, thermal insulation buffering -5°C to +35°C temperature range, and mechanical cushioning protecting underlying vascular cambium
Trunk bark is the composite protective tissue covering Vitis vinifera trunk exterior, comprising all tissues external to vascular cambium including living secondary phloem (inner bark) and dead periderm layers (outer bark), providing protection against desiccation, pathogens, mechanical damage, and temperature extremes while accommodating trunk diameter expansion. Trunk bark structural composition includes= inner bark (living secondary phloem produced by vascular cambium comprising functional sieve tubes, companion cells, phloem parenchyma, and phloem fibers), outer bark (dead tissue external to innermost cork cambium comprising cork cells from successive periderms and crushed non-functional phloem), multiple periderm layers (successive cork cambium generations producing overlapping protective layers), and total bark depth varying by trunk age (2-4 mm in young 3-5 year trunks, 4-8 mm in mature 10-20 year trunks, 6-15 mm in old 30-50+ year trunks). Trunk bark anatomical organization includes= vascular cambium forming innermost bark boundary (interface between bark and wood), living secondary phloem zone 2-4 mm depth (most recent 2-4 years phloem production maintaining transport function), transitional crushed phloem zone (older phloem compressed by xylem expansion losing functionality but not yet incorporated into outer bark), innermost functional cork cambium (producing current periderm at 0.5-2 mm depth from surface), successive periderm layers (multiple cork-phellogen-phelloderm complexes at various depths), and outermost dead cork and bark remnants (weathered exterior layers eventually sloughing). Trunk bark inner bark (living phloem) characteristics include= sieve tube elements conducting photosynthate (transporting sugars from leaves to roots and storage tissues), companion cells supporting sieve tubes (maintaining metabolic functions for enucleate sieve elements), phloem parenchyma storing reserves (accumulating starch providing 15-25% of trunk total carbohydrate storage), phloem fibers providing mechanical strength (thick-walled supporting cells), ray parenchyma connecting radially (continuous with xylem rays enabling lateral transport), and functional lifespan 2-4 years (phloem produced beyond 3-5 years ago becoming crushed and non-functional from xylem pressure). Trunk bark outer bark (dead tissue) characteristics include= phellem (cork cells) from multiple successive periderms (suberized impermeable cells forming waterproof layers), crushed non-functional phloem (old phloem compressed into thin non-living layers), dead phelloderm layers (remnants of inner periderm zones from successive cork cambia), total outer bark depth 1-6 mm in young trunks increasing to 3-12 mm in old trunks, and progressive exterior sloughing (oldest outermost layers cracking, peeling, or flaking as inner layers form continuously renewing bark). Trunk bark thickness variation includes= young trunks (years 1-3) showing thin bark 1.5-3.5 mm (limited phloem and single thin periderm), establishing trunks (years 4-8) showing developing bark 3-6 mm (accumulating phloem and 1-2 successive periderms), mature trunks (years 10-25) showing moderate bark 5-9 mm (substantial phloem depth and 2-4 periderms), old trunks (years 30-50) showing thick bark 7-13 mm (extensive phloem and 4-6+ successive periderms), and ancient trunks (60-100+ years) potentially showing very thick bark 10-20 mm (numerous successive periderms creating deeply furrowed bark). Trunk bark cultivar variation includes= thick bark cultivars (some varieties characteristically producing vigorous cork cambium activity creating 8-15 mm bark depth with pronounced texture), moderate bark cultivars (typical bark development 5-9 mm depth with moderate furrowing), thin bark cultivars (limited cork formation producing 3-6 mm smooth to slightly textured bark), smooth bark types (relatively uniform periderm formation without pronounced ridging), deeply furrowed types (successive periderms creating pronounced longitudinal ridges and valleys), and platy or scaly types (outer bark separating in plates or scales rather than continuous furrowed surface). Trunk bark protective functions include= waterproofing (suberized cork preventing water loss from living tissues, critical during drought or winter dormancy), pathogen exclusion (impermeable cork barrier preventing fungal and bacterial entry), mechanical protection (flexible cork cushioning impacts from equipment, pruning tools, animal browsing, hail), thermal insulation (air-filled cork cells buffering temperature extremes protecting cambium and phloem from heat and cold), UV radiation barrier (outer bark absorbing or reflecting damaging solar radiation), and fire resistance (cork providing some protection against brief fire exposure or radiant heat). Trunk bark gas exchange includes= lenticels (specialized porous regions in cork enabling oxygen influx and carbon dioxide efflux), lenticel density 2-8 per cm² bark surface (varying by cultivar and bark age), lenticel dimensions 1-4 mm diameter creating visible spots or short lines, seasonal lenticel dynamics (expanding during active growth, partially closing during dormancy), and critical function (maintaining aerobic respiration in living phloem and cambium beneath impermeable cork). Trunk bark accommodation of trunk expansion includes= tangential stretching of inner bark (living phloem cells expanding tangentially as trunk circumference increases), successive periderm formation (new cork cambia arising in deeper phloem as outer periderms become non-expandable), outer bark cracking (inelastic outer cork layers splitting under expansion creating characteristic furrowed or platy patterns), progressive outer layer sloughing (oldest bark portions flaking or peeling from surface as expansion continues), and continuous protection maintenance (inner periderms maintaining waterproof covering while outer layers rupture and slough). Trunk bark injury and wound response includes= wound periderm formation (traumatic cork cambium arising beneath injuries rapidly producing protective cork sealing exposed tissues), callus tissue development (undifferentiated parenchyma proliferating around wounds eventually differentiating into functional bark), bark cankers (localized dead areas from disease or injury potentially compromising protection), sunscald damage (southwestern trunk exposure to intense winter sun potentially killing cambium and bark), freeze damage (severe cold potentially killing bark tissues particularly in young trunks with thin bark), and mechanical wounds (pruning cuts, equipment damage, animal injury requiring wound closure through new bark formation). Trunk bark color and texture characteristics include= young bark coloration (typically tan, light brown, or grey-brown on young smooth trunks), mature bark coloration (weathering to darker brown, grey, or reddish-brown depending on cultivar and exposure), bark texture progression (smooth thin bark in youth becoming progressively ridged, furrowed, or platy with age), cultivar-specific patterns (distinctive bark characteristics aiding variety identification), and environmental effects (sun-exposed sides potentially showing darker weathered bark, shaded sides retaining lighter colors). Trunk bark seasonal dynamics includes= spring bark expansion (renewed phloem and cork production causing slight bark swelling), summer active production (continued phloem and periderm formation building bark thickness), fall bark tightening (declining production and tissue consolidation), winter bark stability (dormant period with no growth and potential freeze-thaw weathering of outer layers), and annual bark increment (each year adding 0.3-0.8 mm phloem and 0.5-3 mm cork contributing to cumulative bark thickness). Trunk bark significance in viticulture includes= protecting vital meristems (bark safeguarding vascular cambium and cork cambium from environmental stress and mechanical damage), enabling trunk longevity (continuous bark renewal allowing decades of trunk function), indicating trunk health (bark condition reflecting cambial vigor and pathogen presence), affecting temperature stress vulnerability (thin bark increasing freeze and sunscald risk, thick bark providing better protection), influencing disease susceptibility (bark wounds and cracks providing pathogen entry, thick intact bark resisting invasion), and providing cultivar identification (bark characteristics contributing to variety recognition particularly in old dormant vines).
Trunk_Bark emphasizes complete protective covering and is distinguished from specific bark zones (Outer_Trunk_Bark focusing on dead exterior, inner bark referring to living phloem) by encompassing all tissues external to vascular cambium. Critical for understanding trunk protection mechanisms, bark development over time, cultivar identification characteristics, environmental protection functions, and accommodation of trunk diameter expansion through successive bark renewal.
Bark tissue
Protective bark
Rindengewebe
Schutzrinde
Stammrinde
Trunk Bark
A trunk basal shoot on Vitis vinifera cv. Merlot arising from trunk at 42cm above graft union (65cm above soil surface), activated following severe winter freeze (-24°C) killing bilateral cordons and upper trunk down to this point, shoot emerged in late May (delayed activation reflecting damage assessment and resource mobilization), developed to 198cm length by early August, 24 leaves totaling approximately 3840 cm² leaf area, no inflorescences, representing trunk-level recovery from freeze damage, position above graft union confirming scion variety maintenance (critical for varietal integrity), this shoot deliberately retained for training as replacement trunk over two-season period, first season allowing vigorous growth establishing strong root-shoot connection, second season training to appropriate height and initiating bilateral cordon development, with trunk basal origin providing ideal renewal source avoiding rootstock complications and enabling complete structural reconstruction.
A basal shoot originating specifically from trunk positions above ground level, characterized by emergence from latent buds embedded in trunk bark or cambial tissues, typical origin from concealed buds formed years previously remaining dormant until activation triggers, position ranging from just above graft union to upper trunk near head depending on specific bud location and activation circumstances, and function providing trunk-level renewal options, disease recovery mechanisms, or architectural modification opportunities distinct from below-ground basal shoots that may involve rootstock material. Trunk basal shoots represent above-ground basal growth maintaining scion variety identity (assuming origin above graft union), providing accessible training positions for structural renewal, offering visible indicators of vine stress or damage triggering activation, and enabling controlled renewal strategies utilizing trunk-origin shoots for developing replacement structures without complications of underground origin or potential rootstock involvement.
Trunk basal shoots are distinguished from ground-level or underground basal shoots by above-ground trunk origin, with this distinction having important implications including scion variety certainty (trunk shoots above graft union definitively scion material, underground shoots potentially involving rootstock), accessible training positions (trunk shoots emerging at various heights providing options for structural development), visible activation triggers (trunk bud swelling and shoot emergence observable indicating stress or damage responses), and manageable vigor (trunk shoots often showing less extreme vigor than underground origin shoots though still vigorous relative to normal shoots). Trunk basal activation commonly follows damage scenarios including winter freeze (killing upper portions triggering lower trunk bud activation), disease (Eutypa, Esca, bot canker destroying cordons or upper trunk), mechanical damage (equipment strikes, wind breakage, pruning accidents), or severe stress (extreme drought, defoliation events). Management recognizes trunk basal shoots as valuable renewal resources when positioned appropriately, utilizing them for structural reconstruction while removing poorly-positioned trunk shoots or those arising in undamaged vines indicating problematic spontaneous activation.
Stamm-basaltrieb
above-ground basal shoot
oberirdischer Stockausschlag
trunk renewal shoot
trunk sucker
trunk water shoot
Based on Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press, pp. 85-105 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 113-145 Mundy & Manning (2010) "Budbreak and shoot development in Vitis vinifera following severe trunk damage" Australian Journal of Grape and Wine Research 16=310-316.
Trunk Basal Shoot
Cabernet Sauvignon trunk cambium at 12 years post-planting shows vascular cambium as cylindrical sheath 4-6 cell layers thick positioned 22-28 mm from bark surface between secondary xylem (inner 20-26 mm comprising wood with 12 annual rings) and secondary phloem (outer 2-3 mm inner bark), cork cambium active in outer bark 1-2 mm from surface producing periderm, vascular cambium showing spring reactivation in late March with fusiform initials 380-480 μm length and ray initials producing multiseriate rays, active division April-September producing 2.8-4.2 mm annual xylem ring and 0.3-0.6 mm phloem increment
Trunk cambium is the lateral meristematic tissue in Vitis vinifera trunk comprising vascular cambium and cork cambium (phellogen), producing secondary tissues that increase trunk diameter and provide protective bark layers, representing the actively dividing cell layers responsible for trunk secondary growth and perennial structure development. Trunk cambium includes two distinct meristematic systems= vascular cambium (bifacial cambium located between secondary xylem and secondary phloem producing both tissues centripetally and centrifugally), and cork cambium or phellogen (unidirectional cambium in outer trunk producing cork cells outward and phelloderm inward forming protective periderm). Trunk cambium location and organization includes= vascular cambium positioned as cylindrical sheath between wood (secondary xylem) and inner bark (secondary phloem) at 15-30 mm depth from trunk surface in mature vines, cork cambium positioned in outer bark region developing in cortex initially then in secondary phloem as trunk ages, both cambia forming concentric cylinders around trunk axis, vascular cambium continuous from roots through trunk to branches (maintaining vascular continuity), and cork cambium forming discontinuous patches or complete cylinders depending on trunk age and periderm development stage. Trunk cambium cellular organization includes= vascular cambium comprising fusiform initials (elongated cells 200-500 μm length producing axial xylem and phloem elements) and ray initials (smaller isodiametric cells producing radial ray parenchyma), cork cambium comprising relatively uniform rectangular cells 20-40 μm dimensions producing cork cells outward, cambial initial cells maintaining meristematic competence (ability to divide throughout vine life), cambial zone 2-8 cell layers thick (including initials and immediate derivatives in various differentiation stages), and cambial cells with dense cytoplasm and large nuclei (reflecting active metabolic and division status). Trunk cambium activity patterns include= seasonal cycling with spring reactivation (March-April budbreak triggering cambial division), active division during growing season (April-September producing secondary xylem and phloem), gradual cessation in fall (September-October as growth slows), winter dormancy (November-February with minimal or no division), and reactivation synchronized with budbreak hormonal signals (auxin and cytokinin from expanding buds activating cambial division). Trunk cambium products include= vascular cambium producing secondary xylem centripetally (wood tissue comprising 85-92% of vascular cambium total production by volume), secondary phloem centrifugally (inner bark comprising 8-15% of production), maintaining asymmetric production ratio throughout trunk life, and cork cambium producing phellem (cork cells) centrifugally (comprising 80-90% of cork cambium production), phelloderm (secondary cortex) centripetally (10-20% of production), forming protective periderm complex. Trunk cambium developmental changes include= initial vascular cambium formation from procambium during primary growth (establishing cambial cylinder in young shoots), progressive cambial circumference increase (expanding from 10-30 mm circumference in year 1 to 250-400 mm in mature 15-20 year trunks), fusiform initial lengthening with age (initials 200-300 μm in young cambium extending to 400-600 μm in old cambium), ray initial multiplication (adding new rays as circumference expands maintaining ray density), and cork cambium successive formation (initial periderm in cortex replaced by deeper periderms in secondary phloem as trunk ages). Trunk cambium hormonal regulation includes= auxin (indole-3-acetic acid) produced in shoot apices and young leaves promoting cambial division, cytokinin from roots enhancing cell division and differentiation, gibberellin modulating cambial activity and xylem differentiation, auxin-cytokinin ratio determining xylem versus phloem differentiation (high auxin favoring xylem, balanced ratios producing phloem), and seasonal hormone dynamics (rising auxin levels in spring triggering cambial reactivation, declining levels in fall causing dormancy). Trunk cambium environmental responses include= temperature effects (spring warming triggering reactivation typically requiring sustained temperatures >10-15°C), photoperiod influences (lengthening days in spring contributing to reactivation signals), water availability affecting division rates (drought stress reducing cambial activity, adequate water supporting vigorous division), nutrient availability (nitrogen, phosphorus, potassium supporting cambial metabolism and division), and carbohydrate supply (photosynthate from canopy and stored reserves fueling cambial growth). Trunk cambium significance in trunk development includes= determining trunk diameter increase (cambial activity rate controlling annual ring width and bark thickness), producing conducting tissues (vascular cambium generating xylem and phloem for transport), creating storage tissues (parenchyma in xylem and phloem providing carbohydrate reserves), forming protective layers (cork cambium producing impermeable cork preventing desiccation and pathogen entry), and enabling trunk longevity (perennial cambial activity allowing decades of continued growth). Trunk cambium variation by vine status includes= young vigorous vines showing wide cambial division zones (5-8 cell layers with rapid division rates), mature productive vines showing moderate cambial activity (3-5 cell layers with steady division), old declining vines showing narrow cambial zones (2-3 cell layers with reduced division rates), stressed vines showing intermittent cambial activity (division ceasing during severe stress, resuming with recovery), and grafted vines showing cambial continuity through union (successful grafts establishing vascular and cork cambium connection across graft interface).
Trunk_Cambium emphasizes all lateral meristems and is distinguished from specific cambium types (Vascular_Cambium producing xylem and phloem, Cork_Cambium producing periderm) by encompassing both secondary meristematic systems. Critical for understanding trunk diameter increase, secondary tissue production, seasonal growth patterns, bark formation, and perennial meristematic activity enabling decades of trunk development.
Kambiales Gewebe
Lateral meristem
Lateralmeristem
Secondary meristem
Stammkambium
Trunk Cambium
Long-term evaluation of Trunk_Cold_Hardiness_Disposition at the University of Minnesota documented trunk survival and damage patterns across 30 cultivars through 20 winters including multiple severe events with minima below -35°C. Trunk damage was assessed annually by visual inspection for bark splitting, canker development, and crown injury, with destructive sampling of damaged vines to evaluate internal tissue condition. Cultivar differences in Trunk_Cold_Hardiness_Disposition were substantial and consistent across events. V. vinifera cultivars including 'Riesling' and 'Pinot Noir' showed chronic trunk damage with 15-40% of vines exhibiting bark splitting or crown injury following winters with minima below -28°C, and cumulative vine mortality of 35-60% over the 20-year period. Cold-hardy hybrids including 'Frontenac' and 'Marquette' showed minimal trunk damage (< 5% of vines affected) even following -35°C events, with cumulative mortality below 10%. The graft union proved critical in grafted plantings, with scion-rootstock combinations showing graft union failure when rootstock hardiness substantially exceeded scion hardiness, creating mechanical stress from differential tissue survival. Own-rooted 'Frontenac' showed 100% trunk survival, while 'Frontenac' grafted to the less-hardy '3309C' rootstock showed 12% graft union failure following -34°C exposure. These findings establish Trunk_Cold_Hardiness_Disposition as the foundation of long-term vineyard sustainability in cold climates, with graft union compatibility an additional consideration in grafted systems.
A Trunk_Cold_Hardiness_Disposition is a Midwinter_Hardiness_Disposition that inheres in the trunk (permanent woody structure) of a Vitis vinifera vine and is realized through the freezing tolerance of trunk tissues including bark, phloem, cambium, xylem, and pith, determining survival of the vine's permanent framework and the graft union in grafted vines. This disposition represents the cold hardiness of the perennial structure that represents years to decades of vineyard investment, with trunk survival essential for maintaining trained vine architecture, supporting cordon and cane renewal, and preserving the scion-rootstock connection in grafted vines. Trunk tissues generally achieve similar or greater cold hardiness than corresponding cane tissues, benefiting from larger thermal mass (slower cooling rates), greater insulation from bark accumulation, and in some cases, protection from snow cover at lower trunk positions. However, the trunk contains critical vulnerable zones including the graft union (often the least hardy point in grafted vines), the crown region at soil level (subject to temperature extremes), and any previous wound sites where bark protection is compromised. Trunk damage patterns include localized bark splitting (rapid cooling creating differential contraction), southwest injury (freeze-thaw cycles on sun-exposed bark), crown injury (extreme cold at poorly insulated soil surface), and graft union failure (differential hardiness between scion and rootstock tissues). Trunk_Cold_Hardiness_Disposition is the ultimate determinant of vine survival, as trunk death necessitates vine replacement regardless of bud or cane condition, representing multi-year production loss and substantial replanting costs.
Trunk_Cold_Hardiness_Disposition is_a Midwinter_Hardiness_Disposition. Trunk hardiness determines vine survival independent of annual crop production, representing the highest-stakes cold tolerance parameter. Trunk damage includes bark splitting, southwest injury, crown injury, and graft union failure, each with distinct mechanisms and patterns. Graft union hardiness depends on both scion and rootstock tissue hardiness and their compatibility under freeze stress. Trunk death requires vine replacement (multi-year loss), while cane and bud damage affect only current-season production.
Rebstamm-Winterhärte
Stammfrosthärte
Stammkältehärte-Disposition
permanent wood hardiness
trunk freezing tolerance
vine framework cold resistance
Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235 Howell GS (2000) Grapevine cold hardiness: mechanisms of cold acclimation, mid-winter hardiness maintenance, and spring deacclimation, Proceedings of the ASEV 50th Anniversary Annual Meeting Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection, Michigan State University Extension Bulletin E-2930
Trunk Cold Hardiness Disposition
Long-term evaluation of Trunk_Disease_Resistance_Disposition at INRAE Bordeaux monitored trunk disease development across 24 cultivars in a replicated trial established in 1990 and assessed annually for esca foliar symptoms and vine mortality through 2020. After 30 years, cumulative vine mortality attributed to trunk diseases ranged from 8% in resistant 'Merlot' to 62% in highly susceptible 'Sauvignon Blanc', with 'Cabernet Sauvignon' intermediate at 28%. Annual esca foliar symptom expression (tiger-stripe patterns, apoplexy) averaged 2.4% of 'Merlot' vines versus 18.6% of 'Sauvignon Blanc' vines during epidemic years. Wood anatomical studies revealed that 'Merlot' exhibits more rapid and extensive tylosis formation in xylem vessels adjacent to infection sites, compartmentalizing pathogens and maintaining functional vascular capacity. Phytotoxin tolerance assays demonstrated that 'Merlot' leaf tissue exhibited 40% less necrosis than 'Sauvignon Blanc' when exposed to equivalent concentrations of Phaeomoniella chlamydospora culture filtrates, suggesting both structural and physiological components contribute to its enhanced Trunk_Disease_Resistance_Disposition. These findings inform replanting decisions and cultivar selection for vineyard longevity.
A Trunk_Disease_Resistance_Disposition is a Fungal_Resistance_Disposition that inheres in a Vitis vinifera plant and is realized through defense processes that limit infection, colonization, and pathogenesis by the complex of wood-inhabiting fungi causing grapevine trunk diseases, including esca, Eutypa dieback, Botryosphaeria dieback, Phomopsis dieback, and related syndromes. This disposition represents the capacity to resist or tolerate chronic fungal infection of perennial woody tissues including the trunk, cordons, and spurs, where pathogens cause progressive vascular dysfunction, wood necrosis, and eventual vine decline or death. Trunk diseases are caused by diverse fungal taxa that share the common features of wound-mediated infection, slow progression over multiple years, colonization of xylem and associated tissues, and production of phytotoxins that disrupt vascular function and cause foliar symptoms distal to infection sites. Resistance mechanisms operating against trunk diseases fundamentally differ from those effective against foliar pathogens, emphasizing wound healing efficiency, compartmentalization of infected tissues through tylosis formation and barrier zone development, tolerance to fungal phytotoxins, and maintenance of functional vascular capacity despite partial colonization. No complete resistance to trunk diseases exists in cultivated Vitis, but substantial variation in susceptibility among cultivars and rootstocks enables selection for enhanced tolerance. Trunk diseases have emerged as the most significant threat to vineyard longevity globally, causing estimated annual losses exceeding €1.5 billion in Europe alone, making resistance breeding a critical priority despite the challenges posed by long evaluation periods and complex, multi-pathogen etiology.
Trunk_Disease_Resistance_Disposition is_a Fungal_Resistance_Disposition. This class encompasses resistance to the entire complex of wood-inhabiting fungi causing grapevine trunk diseases (GTDs), recognizing that vineyards typically face multiple pathogens simultaneously. Child classes include Esca_Resistance_Disposition, Eutypa_Resistance_Disposition, and Botryosphaeria_Resistance_Disposition for resistance to specific pathogen groups. Resistance mechanisms differ fundamentally from foliar disease resistance, emphasizing wound responses, compartmentalization, and phytotoxin tolerance rather than pathogen recognition and hypersensitive response.
Esca-Komplex-Resistenz
GTD resistance
Holzpilz-Resistenz
Rebholzkrankheit-Resistenz-Disposition
vascular pathogen resistance
wood disease resistance
Gramaje D et al. (2018) Managing grapevine trunk diseases with respect to etiology and epidemiology: current strategies and future prospects, Plant Disease 102:12-39 Bertsch C et al. (2013) Grapevine trunk diseases: complex and still poorly understood, Plant Pathology 62:243-265 Mugnai L et al. (1999) Esca (black measles) and brown wood-streaking: two old and elusive diseases of grapevines, Plant Disease 83:404-418
Trunk Disease Resistance Disposition
Trunk freezing tolerance quality assessment following an extreme polar vortex event in Niagara Peninsula (minimum -30°C) evaluated the survival of permanent vine structures across cultivars and training systems. Cambium viability testing through the scraping method (green = alive, brown = dead) combined with spring wound healing assessment quantified trunk survival patterns. Own-rooted Vitis vinifera cultivars showed marginal trunk freezing tolerance quality with 15% trunk mortality concentrated in young vines and those with prior trunk damage from mechanical injury or crown gall, while mature, healthy trunks showed 95% survival indicating that well-acclimated V. vinifera trunks withstood -30°C in this event. Trunk damage distribution showed severe injury concentrated at the soil-line junction in 70% of affected vines, with freeze crack formation extending upward from the crown region where ice nucleation conditions apparently exceeded the trunk freezing tolerance quality of the tissues. Training system effects revealed that high-cordon trained vines showed greater trunk damage than low-trained systems where winter snow cover provided thermal insulation, reducing effective cold exposure. Interspecific hybrids including Vidal and Baco Noir showed excellent trunk freezing tolerance quality with less than 2% trunk mortality, consistent with the enhanced cold hardiness from native American Vitis species parentage providing protection beyond V. vinifera limits.
Trunk freezing tolerance quality is a tissue freezing tolerance quality that inheres in grapevine trunks and represents the capacity of the permanent vine structure to survive extreme cold exposure through extracellular freezing tolerance mechanisms that enable the woody tissues of bark, cambium, xylem, and pith to survive the formation of ice in extracellular spaces without lethal cellular damage, with trunk survival representing the threshold between recoverable and catastrophic cold injury since trunk death typically requires vine replacement. This quality addresses the cold tolerance of the foundational vine structure that supports all above-ground growth, recognizing that trunk damage sufficient to cause cambial death at the soil line eliminates the hydraulic and structural continuity required for vine function, converting a potentially recoverable cold injury (loss of buds or canes) into a replanting situation requiring multiple years for new vine establishment. The ontological significance of trunk freezing tolerance quality lies in its role as the ultimate determinant of vine survival under extreme cold, with trunk death representing the threshold at which cold damage transitions from seasonal crop loss to permanent vineyard asset destruction requiring capital investment for replacement. The biological basis of trunk freezing tolerance encompasses the extracellular freezing tolerance mechanism in which ice forms in intercellular spaces and xylem vessels, withdrawing water from living cells through the vapor pressure gradient between ice and liquid water, with cell survival depending on the capacity of membranes and proteins to withstand the dehydration stress and to recover function upon thawing. The trunk shows generally greater cold tolerance than buds due to its lower water content, the thermal mass providing slower cooling rates, and the structural barriers limiting ice propagation through the tissue, but remains vulnerable to extreme cold events particularly when acclimation is incomplete or when tissues are damaged by other factors that compromise their freezing tolerance capacity. The pattern of trunk freeze damage often shows the most severe injury at or just below the soil line where the trunk-soil interface may create ice-nucleating conditions or where thermal dynamics differ from free-air exposed trunk sections.
Trunk_Freezing_Tolerance_Quality is_a Tissue_Freezing_Tolerance_Quality, representing the cold hardiness of the permanent main stem structure determining vine survival under extreme cold. This class addresses the survival threshold distinguishing recoverable from catastrophic cold damage. Extracellular freezing tolerance mechanism enables survival of ice formation in woody tissues. Soil-line zone shows particular vulnerability to freeze damage. Training system and snow cover modulate effective cold exposure. V. vinifera trunk hardiness sufficient for -30°C when well-acclimated hybrids provide additional margin. Trunk death requires replanting, representing capital loss beyond crop damage.
Dauerstruktur-Frosttoleranz
Hauptstamm-Winterhärte
Stamm-Gefrier-Toleranz
Stamm-Kältehärte
main stem cold survival
permanent structure freezing tolerance
trunk cold hardiness
trunk winter hardiness
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection. Michigan State University Extension Bulletin E2930 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 508-545
Trunk Freezing Tolerance Quality
A trunk latent bud on a 25-year-old trunk of Vitis vinifera cv. Syrah, located 45cm above the graft union embedded within bark tissue, remaining suppressed for at least 15 years based on annual ring counting, becoming activated following severe Eutypa dieback that killed 70% of the cordon and fruiting wood, producing a vigorous water shoot 220cm in length with 28 leaves during the first season, subsequently pruned and trained over three seasons to establish a replacement cordon restoring the vine to full production.
A latent bud embedded in the bark or corky tissue of the permanent trunk structure, capable of remaining suppressed for many years to decades, characterized by small size, deep position within bark fissures or lenticels, vascular connection to xylem through rays in the wood, and strong apical dominance suppression from distal canopy growth. Trunk latent buds serve as long-term reserve meristems capable of producing water shoots or suckers when released from suppression by trunk damage, severe pruning, or vine decline, representing the vine's capacity for structural regeneration and survival following catastrophic damage to above-ground structures, with activation triggering vigorous vegetative growth that can be trained to reconstruct productive canopy architecture over multiple seasons.
Trunk latent buds are the source of water shoots and trunk suckers that viticulturists typically remove during summer canopy management to prevent competition with fruiting wood and maintain desired vine architecture. However, trunk latent buds become valuable for vine recovery following trunk disease damage (Eutypa, Esca, Crown Gall), freeze injury to cordons, or intentional trunk renewal programs. Their position (upper versus lower trunk) influences vigor and training ease, with lower trunk positions often preferred for structural renewal due to better vascular connections and easier shoot positioning. Trunk latent bud viability and responsiveness generally decline with trunk age, though some buds remain capable of activation even in very old vines.
Adventivknospe am Stamm
Stammknospe
epicormic bud
trunk bud
trunk epicormic bud
Based on Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Sosnowski et al. (2007) "The influence of grapevine cultivar and isolate of Eutypa lata on wood and foliar symptoms" Plant Disease 91=924-931 Gramaje et al. (2018) "Fungal trunk pathogens in the grapevine propagation process= potential inoculum sources, detection, identification, and management strategies" Plant Disease 102=469-493.
Trunk Latent Bud
Trunk recovery capacity quality evaluation in Ontario Vidal following severe winter injury (-34°C) assessed the rehabilitation potential of freeze-damaged trunks and the management strategies appropriate for different damage levels. Systematic trunk assessment combining the scratch test for cambium viability, cross-sectional analysis for vascular damage extent, and evaluation of basal bud survival classified vines into recovery categories. Vines with intact basal cambium and less than 50% vascular damage showed good trunk recovery capacity quality suitable for conservative management allowing natural healing, with callus development visible by midsummer and approximately 80% achieving functional recovery by the second post-freeze season. Vines with extensive upper-trunk damage but surviving crown cambium and basal buds showed moderate trunk recovery capacity quality appropriate for trunk renewal through cutting back to viable tissue and retraining from basal suckers, achieving full recovery in 3 seasons. Vines with complete girdling at the crown and no surviving basal buds showed poor trunk recovery capacity quality requiring replanting, representing approximately 15% of the affected vineyard. The trunk recovery capacity quality assessment enabled differentiated management, with rehabilitation costs for retraining substantially below replanting costs and productivity restoration faster than new planting establishment, demonstrating the value of accurate recovery capacity evaluation for optimizing post-freeze vineyard management decisions.
Trunk recovery capacity quality is a cold recovery quality that inheres in a grapevine organism and represents the ability to restore trunk function and structural integrity following freezing injury through the regeneration of vascular tissues from surviving cambium, the healing of freeze wounds through callus formation and compartmentalization, and the potential for retraining from basal shoots or suckers when trunk damage is too severe for repair, with trunk recovery determining whether damaged vines can continue as productive units or require replacement. This quality addresses the most consequential dimension of cold damage recovery, recognizing that trunk survival determines whether a vine remains a productive asset requiring only canopy reconstruction or becomes a total loss requiring the multi-year investment of replanting and reestablishment. The ontological significance of trunk recovery capacity quality lies in its relationship to vineyard capital preservation, with effective trunk recovery enabling damaged vines to return to full productivity within 2-3 seasons while trunk mortality requires replanting with associated costs and 4-5 year delay before full productivity, such that trunk recovery capacity substantially affects the economic consequences of severe freeze events. The biological basis of trunk recovery capacity encompasses the regenerative activity of surviving cambium producing new vascular tissues that can restore transport function around localized damage zones, the wound compartmentalization processes that isolate dead tissue and prevent pathogen spread into healthy trunk regions, and the latent bud reserves at the trunk base that can produce replacement shoots for retraining when upper trunk damage is too extensive for repair. The assessment of trunk recovery capacity involves evaluating the extent and location of trunk damage, the viability of remaining cambium, the presence of viable basal buds for retraining, and the management options appropriate for the specific damage pattern observed. The temporal dimension of trunk recovery extends over multiple seasons, with initial callusing and compartmentalization occurring in the first year, vascular regeneration and functional recovery progressing over 2-3 years, and full restoration of trunk structure and function potentially requiring 3-5 years depending on damage severity and management support.
Trunk_Recovery_Capacity_Quality is_a Cold_Recovery_Quality, representing the ability to restore trunk function following freeze damage through regeneration, wound healing, or retraining. This class addresses the capital-preservation dimension of freeze recovery. Cambium viability, vascular damage extent, and basal bud survival determine recovery options. The rehabilitation versus replanting decision has major economic consequences (3-year retrain vs. 5-year replant). Wound compartmentalization and vascular regeneration proceed over multiple seasons. Recovery management should match intervention intensity to damage severity.
Dauerstruktur-Wiederherstellung
Stamm-Erholungskapazität
Stamm-Regeneration
Stammwundheilung
permanent structure recovery
trunk regeneration
trunk repair capacity
trunk wound healing
Trunk Recovery Capacity Quality
Long-term documentation of Trunk_Regeneration_Disposition at North Dakota State University tracked recovery outcomes following severe trunk damage across cultivars and training systems over 8 years in a replicated trial subjected to -38°C exposure in year 1. Trunk damage was categorized as partial (crown injury or bark splitting with some surviving cambium) or complete (circumferential cambium death requiring trunk replacement). Regeneration success, time to full production, and secondary complications were recorded. Trunk_Regeneration_Disposition varied among cultivars. 'Frontenac' with partial trunk damage showed excellent wound compartmentalization, with cold-damaged sectors walling off within one season, no crown gall development, and full yield restoration by year 2. 'Frontenac' with complete trunk death produced vigorous basal suckers (2.8 suckers per vine by June, strongest reaching 2.1 m by season end), enabling trunk replacement with return to 50% yield by year 2 and full yield by year 3. 'Marquette' showed similar sucker vigor but higher crown gall incidence (35% of damaged vines) at wound sites, requiring removal of infected tissue and extending recovery to year 4 for affected vines. V. vinifera cultivars on susceptible rootstocks showed poor Trunk_Regeneration_Disposition, with weak sucker production (0.8 per vine), high crown gall incidence (55%), and 28% vine mortality over the recovery period. These findings demonstrate that Trunk_Regeneration_Disposition determines vineyard survival and recovery economics following catastrophic winter events.
A Trunk_Regeneration_Disposition is a Cold_Recovery_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to regenerate functional trunk tissue following winter damage to the permanent woody structure, either through wound healing of partially damaged trunks or complete trunk replacement from basal suckers or graft union regrowth. This disposition represents recovery from the most severe category of cold damage, where the vine's permanent framework is compromised and structural rebuilding is required beyond simple shoot replacement. Trunk damage from freezing includes bark splitting (from rapid cooling and ice crystal expansion), crown gall development in cold-injured tissues (Agrobacterium infection of freeze-damaged cells), cambium kill (eliminating radial growth capacity), and complete trunk death requiring regeneration from surviving basal tissues. Trunk_Regeneration_Disposition encompasses wound compartmentalization capacity (limiting secondary pathogen invasion of damaged tissues), cambial regeneration from surviving sectors (restoring circumferential vascular connectivity), and sucker production capacity (providing replacement trunk candidates when existing trunk is unsalvageable). The timeframe for trunk recovery is multi-year, with complete trunk replacement requiring 2-4 years to establish new structural framework and return to full production. Cultivars and rootstocks differ in trunk regeneration capacity based on wound healing efficiency, susceptibility to secondary infections, vigor of basal growth, and tolerance of the training disruption required for trunk replacement.
Trunk_Regeneration_Disposition is_a Cold_Recovery_Disposition. Trunk regeneration addresses the most severe cold damage category requiring structural rebuilding. Recovery pathways include wound healing (partial damage) and trunk replacement from suckers (complete damage). Timeframe is multi-year, with full production recovery requiring 2-4 years depending on damage extent. Secondary infection (especially crown gall) is a major complication affecting regeneration success. Both scion and rootstock genotype influence trunk regeneration capacity in grafted vines.
Permanentholz-Erholung
Stamm-Regenerations-Disposition
Stammwiederherstellung
permanent wood restoration
structural regeneration
trunk recovery
Goffinet MC (2004) Anatomy of grapevine winter injury and recovery, Grape Research News 15:1-7 Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection, Michigan State University Extension Bulletin E-2930 Burr TJ, Otten L (1999) Crown gall of grape: biology and disease management, Annual Review of Phytopathology 37:53-80
Trunk Regeneration Disposition
A trunk water shoot on Vitis vinifera cv. Merlot in Pomerol, France, emerging from a latent bud at 85 cm trunk height (midway between graft union at 40 cm and bilateral cordon at 120 cm), developing in July following severe spring frost that killed 60% of primary shoots, growing 175 cm by September with 19 nodes averaging 13 cm internode length, leaves reaching 210 cm² area, completely blocking the central trunk zone and interfering with tractor passage for undervine cultivation, requiring careful removal during dormant pruning with clean cuts flush to trunk surface to prevent repeated emergence from the same bud position in subsequent seasons.
A water shoot originating from latent or adventitious buds located on the main trunk structure between the graft union and the first permanent arm or cordon position in Vitis vinifera, characterized by emergence from bark surfaces that may be several years old, rapid vertical growth due to strong apical dominance released by canopy damage or severe pruning, and competition with established framework structures for vascular resources and carbohydrate allocation. Trunk water shoots represent particularly problematic growth in commercial viticulture because their position on the central trunk architecture interferes with established training system geometry, creates difficulty in mechanical operations including harvest and spraying, and often indicates underlying physiological imbalances or damage conditions requiring diagnostic attention. These shoots typically develop from buds that have remained dormant since trunk establishment, potentially 5-15 years prior to emergence, and their activation signals either acute stress responses to canopy loss or chronic nutritional imbalances favoring excessive vegetative growth. Management decisions regarding trunk water shoots range from immediate removal to preserve training system integrity, to selective retention for trunk renewal in aging vineyards where progressive trunk diseases necessitate framework replacement strategies.
Trunk water shoots are distinguished from arm water shoots by their origin below the first permanent branch point on the main vertical or angled trunk axis. Position along the trunk significantly affects management priorities, with lower trunk water shoots (near graft union) requiring immediate removal to prevent rootstock competition, while upper trunk water shoots (near cordon positions) may occasionally be retained for framework reconstruction. Repeated emergence of trunk water shoots from the same positions indicates either incomplete removal leaving basal bud tissue, or persistent physiological imbalances requiring corrective management of irrigation, fertilization, or crop load. Trunk water shoots developing from adventitious rather than latent buds often show even more vigorous growth and greater difficulty in complete removal.
Stammwasserschoss
bole epicormic shoot
central trunk sucker
stem water shoot
zentraler Wassertrieb
Based on Winkler et al. (1974) "General Viticulture" University of California Press pp. 285-301 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles pp. 134-147 Sosnowski et al. (2007) "Pathogenicity of Eutypa lata in grapevine= shoot vigor and cane dieback" Phytopathology 97=651-658 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 295-318.
Trunk Water Shoot
Cabernet Sauvignon trunk wood at 12 years post-planting shows 32-42 mm total radius with 12 annual rings, outer sapwood 18-25 mm depth (rings 1-7) actively conducting water at 35-55 L/day and storing 55-75 g starch, inner sapwood 8-12 mm (rings 8-10) with declining function, heartwood 6-10 mm central core (rings 11-12) non-conducting with darker coloration from phenolic deposits, ring-porous anatomy with earlywood vessels 180-260 μm diameter, and wood density 0.52-0.58 g/cm³
Trunk wood is the secondary xylem tissue comprising the bulk of Vitis vinifera trunk diameter, produced by vascular cambium activity through successive annual growth rings, providing mechanical support for the canopy, conducting water and minerals from roots to shoots, storing carbohydrate reserves, and differentiating into functional sapwood and non-conducting heartwood zones as the trunk ages. Trunk wood develops through vascular cambium activity= cambium produces secondary xylem cells centripetally (inward toward trunk center) and secondary phloem centrifugally (outward), xylem cells differentiate into vessels (water-conducting tracheary elements), fibers (mechanical support), and parenchyma (storage, lateral transport), lignification of cell walls provides rigidity and durability, and successive years produce distinct annual growth rings visible in cross-section. Trunk wood anatomy in Vitis vinifera exhibits ring-porous pattern= earlywood (formed early growing season, April-June in Northern Hemisphere) contains large-diameter vessels (100-300 μm diameter, primarily for water transport), latewood (formed late season, July-September) contains smaller vessels (40-120 μm diameter), fiber cells throughout providing mechanical strength, and axial and radial parenchyma providing storage and lateral transport. Trunk wood functional zonation includes= outer sapwood (most recently formed wood, actively conducting water and storing reserves), inner sapwood (older but still functional wood with some conducting capacity and storage), transition zone (wood transitioning from sapwood to heartwood with declining function), and heartwood (oldest central wood with ceased conduction, structural support only, often darker colored from phenolic deposits). Trunk wood radial dimensions vary by vine age and vigor= young vines (3-5 years) showing 10-25 mm total wood radius (entirely sapwood), mature vines (10-20 years) showing 30-70 mm radius (sapwood 15-30 mm, heartwood 15-40 mm), old vines (30-50 years) showing 60-125 mm radius (sapwood 15-25 mm, heartwood 45-100 mm), with sapwood depth relatively constant while heartwood accumulates with age. Trunk wood annual ring characteristics include= ring width varying 1-8 mm depending on vine age, vigor, and environmental conditions (wide rings 4-8 mm in young vigorous vines, narrow rings 1-3 mm in mature or stressed vines), ring boundaries marked by transition from small latewood vessels to large earlywood vessels of next year, ring-porous pattern with earlywood vessels occupying 30-60% of ring area, and rings providing dendrochronological record of vine growth history and environmental conditions. Trunk wood vessel characteristics include= earlywood vessels 120-300 μm diameter providing majority of hydraulic conductance (single large vessel can transport more water than hundreds of small vessels due to Hagen-Poiseuille relationship), latewood vessels 40-120 μm diameter providing additional capacity and safety (smaller vessels less prone to embolism), vessel elements 200-600 μm length with perforated end walls allowing longitudinal water flow, and vessel walls with bordered pits enabling lateral water movement between vessels and to parenchyma. Trunk wood parenchyma functions include= starch storage in ray and axial parenchyma cells (trunk wood containing 40-80% of total vine carbohydrate reserves), lateral transport of water and solutes through ray parenchyma (radial movement from vessels to cambium and phloem), hormone and signaling molecule transport (auxins, cytokinins, abscisic acid moving through parenchyma), and embolism repair (parenchyma potentially providing water to refill embolized vessels). Trunk wood mechanical properties include= density 0.45-0.65 g/cm³ (varying by cultivar, growth rate, sapwood versus heartwood), compressive strength 30-50 MPa (supporting canopy weight and resisting wind stress), tensile strength 60-100 MPa (resisting pulling forces), flexural strength enabling trunk to bend without breaking under wind load, and sufficient rigidity to support 5-20 kg canopy mass on single trunk. Trunk wood carbohydrate storage capacity includes= starch accumulation in parenchyma cells throughout sapwood (outer sapwood 8-15% dry weight starch, inner sapwood 5-10%), minimal storage in heartwood (parenchyma cells dead or non-functional), seasonal cycling (starch accumulation post-harvest reaching 60-150 g per vine, mobilization in spring consuming 40-60% of reserves, replenishment during growing season), and storage capacity increasing with trunk size (larger trunks providing greater reserve buffering capacity). Trunk wood pathological considerations include= trunk diseases (Eutypa, Esca, Botryosphaeria) causing wood necrosis, cankers, and decay, wood decay fungi (white rot and brown rot organisms) degrading heartwood cellulose and lignin, vascular wilt pathogens potentially colonizing xylem vessels, wood-boring insects creating galleries weakening structural integrity, and freeze damage causing xylem vessel disruption particularly in sapwood.
Trunk_Wood emphasizes complete secondary xylem and is distinguished from Sapwood (functional outer wood) and Heartwood (non-functional inner wood) by encompassing entire wood tissue including both functional and structural zones. Critical for understanding trunk hydraulic capacity, carbohydrate storage in woody perennials, mechanical support of grapevine architecture, and trunk disease impacts on wood function and vine health.
Rebholz
Secondary xylem
Trunk xylem
Wood tissue
Trunk Wood
Field evaluation of Tuberosity_Resistance_Disposition at CSIRO Adelaide compared long-term root health across 12 rootstock genotypes in a vineyard with documented aggressive phylloxera biotype pressure over 15 years. Root systems were excavated at years 5, 10, and 15 for assessment of tuberosity severity (0-5 scale), structural root integrity, and secondary pathogen colonization. Own-rooted V. vinifera 'Shiraz' controls showed catastrophic decline with tuberosity scores of 4.8 by year 5, complete structural root deterioration, 100% Fusarium colonization, and vine death by year 8. The V. berlandieri × V. rupestris rootstock '140 Ruggeri' demonstrated strong Tuberosity_Resistance_Disposition despite moderate nodosity presence on young roots (nodosity score 3.2). Tuberosity formation remained minimal (score 0.8 at year 15), structural roots maintained integrity, Fusarium colonization was only 12%, and vines sustained commercial productivity throughout the trial. Histological examination revealed that '140 Ruggeri' roots responded to phylloxera probing of lignified tissues with intense suberization and phenolic deposition creating impenetrable barriers, while successful feeding attempts triggered rapid cork cambium activation isolating small tuberosities before pathogen entry. This field performance demonstrates that Tuberosity_Resistance_Disposition can provide durable vine protection even where nodosity limitation is incomplete.
A Tuberosity_Resistance_Disposition is a Root_Phylloxera_Resistance_Disposition that inheres in a Vitis plant and is realized through defense processes that prevent or limit formation of tuberosities—the warty, necrotic galls induced by grape phylloxera feeding on older, lignified structural roots. This disposition represents resistance to the most destructive phase of root phylloxera infestation, as tuberosities disrupt the permanent root framework, provide entry points for secondary soil pathogens (particularly Fusarium and other root rot fungi), and cause progressive structural deterioration leading to vine decline and death. Tuberosities develop when phylloxera colonize suberized roots through lenticels or wounds, inducing localized hyperplasia and subsequent necrosis that creates characteristic rough, corky proliferations. Unlike nodosities which may be tolerated without significant vine impact, extensive tuberosity formation inevitably leads to vine decline through combined effects of reduced water and nutrient uptake, pathogen invasion, and structural root loss. Resistance mechanisms include lignin and suberin barriers preventing phylloxera access to feeding-competent tissues, rapid wound periderm formation isolating feeding sites, production of phenolic compounds toxic to phylloxera in lignified tissues, and anatomical features limiting phylloxera penetration. Tuberosity_Resistance_Disposition is often mechanistically distinct from nodosity limitation, with some rootstocks (particularly V. berlandieri derivatives) permitting nodosity formation while effectively preventing tuberosity development, providing adequate field protection through this alternative resistance strategy.
Tuberosity_Resistance_Disposition is_a Root_Phylloxera_Resistance_Disposition. Tuberosities on lignified structural roots cause more serious vine damage than nodosities on young roots, as they affect the permanent root framework and facilitate pathogen entry. Some rootstocks (especially V. berlandieri derivatives) show stronger tuberosity resistance than nodosity limitation, representing an alternative resistance strategy. The combination of tuberosity formation and secondary pathogen invasion is typically responsible for vine death in susceptible genotypes, making tuberosity resistance the critical determinant of long-term rootstock performance.
Strukturwurzel-Resistenz
Tuberosität-Resistenz-Disposition
Tuberositäts-Abwehr
lignified root defense
structural root resistance
tuberosity suppression
Granett J et al. (2001) Biology and management of grape phylloxera, Annual Review of Entomology 46:387-412 Powell KS et al. (2013) Tolerance and resistance to root damage are not correlated during phylloxera host-plant interactions, Plant Pathology 62:1251-1261 Kellow AV et al. (2004) Interaction between Vitis vinifera and grape phylloxera, Annals of Botany 93:581-590
Tuberosity Resistance Disposition
Syrah in Rhône Valley shows turgor-driven expansion onset during mid-late July Year N+1 at BBCH 73-74, with osmotic potential declining from -0.6 to -0.8 MPa, water uptake generating turgor pressure 0.4-0.7 MPa, pressure-driven cell expansion initiating, yielding walls enabling volume increase, cells enlarging 2.1-fold, under warm Mediterranean conditions of 25-29°C temperature with adequate water, typical of this mid-to-late season cultivar establishing turgor-expansion system
Turgor-driven expansion onset represents the pressure-based growth initiation phase during initial cell enlargement when osmotic water uptake generates turgor pressure 0.3-0.8 MPa, pressure force pushes against yielding cell walls causing mechanical expansion, and turgor-pressure mechanism establishes as physical driver of cell volume increase, initiating over days to week during transition from division to enlargement. This pressure system establishment phase occurs during mid-late summer in mid-late July in temperate regions Year N+1, initiating during week 3-4 post-fertilization as cells exit division, corresponding to BBCH 73-74 pea-sized berry stages. Turgor onset mechanism involves osmotic water uptake from solute accumulation, water influx generating hydrostatic pressure within cell, turgor pressure 0.3-0.8 MPa exerting force on cell walls, yielding walls enabling pressure-driven expansion, and established turgor-expansion system driving volume increase. Osmotic basis of turgor shows solute accumulation in cell and vacuole creating osmotic potential more negative than surrounding apoplast, water potential gradient driving water influx from apoplast into cell, water accumulation increasing cell volume and hydrostatic pressure, and osmotic-hydraulic system generating turgor. Water potential gradient shows typical apoplast water potential -0.3 to -0.5 MPa, cell osmotic potential -0.7 to -0.9 MPa during turgor onset from solute accumulation, resulting water potential gradient 0.2-0.4 MPa driving water influx, and gradient as driving force for water movement. Osmotic potential components show organic acids contributing -0.3 to -0.5 MPa, potassium and other ions -0.2 to -0.3 MPa, sugars -0.1 to -0.2 MPa still limited during onset, and combined solutes creating total osmotic potential. Water influx pathways show water crossing plasma membrane through aquaporin water channels, possible apoplastic water movement to cell surface, symplastic water movement through plasmodesmata between cells, and integrated pathways supplying water for turgor generation. Aquaporin function shows water channel proteins facilitating membrane water permeability, aquaporin expression correlating with water uptake rates, channels enabling rapid water flux responding to osmotic gradient, and aquaporins as key transport proteins during expansion. Turgor pressure generation shows water accumulation increasing cell volume, hydrostatic pressure building from constrained expansion, turgor pressure 0.3-0.8 MPa typical during onset phase, and pressure as mechanical force driving expansion. Turgor measurement shows pressure probepressure chamber methods quantifying cell turgor, typical measurements 0.3-0.5 MPa early onset increasing to 0.6-0.8 MPa established onset, and turgor quantification documenting pressure dynamics. Pressure-volume relationships show turgor pressure P = π - Ψ where π is osmotic potential and Ψ is water potential, pressure increasing with solute accumulation and water uptake, and relationship governing turgor dynamics. Wall yielding under turgor shows cell wall experiencing tensile stress from internal pressure, wall yielding plastically under stress enabling irreversible expansion, yield threshold determining minimum turgor required for expansion typically 0.3-0.5 MPa, and yielding enabling pressure-driven growth. Wall mechanical properties show wall capable of elastic deformation recovering when pressure released and plastic deformation causing permanent expansion, plastic yielding under sustained turgor essential for growth, wall stress-strain relationships determining expansion response, and mechanical properties regulated through wall modification. Turgor-driven expansion rate shows expansion velocity proportional to turgor pressure exceeding yield threshold, typical relationship - growth rate = extensibility × (P - Y) where P is turgor and Y is yield threshold, higher turgor driving faster expansion, and pressure as rate-determining factor. Cell volume increase from turgor shows progressive volume expansion from water accumulation and turgor-driven wall yielding, typical 1.5-3 fold volume increase during turgor onset phase, volume change directly resulting from pressure-driven expansion, and turgor as mechanical driver of enlargement. Turgor maintenance shows continued solute accumulation maintaining osmotic gradient, continued water uptake sustaining turgor, and sustained pressure enabling continued expansion. Metabolic support for turgor shows respiratory ATP production supporting ion pumps, active transport accumulating solutes creating osmotic potential, metabolic energy essential for maintaining solute gradients, and metabolism supporting turgor system. Environmental effects on turgor onset include water availability critically affecting turgor generation with adequate water essential for pressure development, water deficit preventing full turgor establishment limiting expansion, temperature affecting metabolic solute accumulation and transport, and overall water as most critical factor. Water deficit effects show reduced water availability preventing turgor development, deficit during onset potentially permanently limiting expansion capacity, and early deficit as critical limitation. Temperature effects show warm conditions promoting faster solute accumulation and turgor development, cool conditions slowing but generally not preventing turgor onset, and temperature affecting onset timing. Turgor onset timing shows typical initiation week 3-4 as cells exit division, asynchronous onset across cell populations as cells progressively commit to enlargement, and extended onset period week 3-5 during transition. Turgor establishment completion shows full turgor pressure 0.6-0.8 MPa established by week 4-5, established turgor maintaining during subsequent rapid expansion, and pressure system enabling sustained enlargement. The process represents first aspect of initial cell enlargement establishing pressure-driven growth system, preceding vacuole enlargement initiation and cell wall loosening. Duration represents turgor development over days to week during transition weeks 3-4. Regional timing shows turgor-driven expansion onset during mid-late July across climates as cells initiate enlargement. Viticultural significance includes representing establishment of physical growth driving force, demonstrating critical water dependence for pressure system, showing turgor as mechanical expansion driver, and establishing pressure-driven growth enabling subsequent enlargement. Research applications include analyzing turgor generation and maintenance mechanisms, investigating water relations and osmotic regulation, evaluating pressure-volume relationships during expansion, quantifying turgor-growth relationships, and assessing turgor as determinant of expansion capacity and rate.
Turgor_Driven_Expansion_Onset is first aspect of Initial_Cell_Enlargement, distinguished by pressure system establishment. Represents osmotic water uptake generating turgor 0.3-0.8 MPa. Pressure force drives expansion against yielding walls. Water potential gradient -0.2 to -0.4 MPa driving influx. Aquaporins facilitate water transport. Cell volume 1.5-3 fold increase during onset. Water availability critical. Timing week 3-4. Precedes Vacuole_Enlargement_Initiation. Establishes physical growth mechanism.
Turgor-Entwicklung
Turgor-getriebene Expansions-Beginn
druckbasiertes Wachstum-Start
osmotic expansion initiation
osmotische Expansions-Initiierung
pressure-based enlargement
pressure-driven growth onset
turgor development
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 443-445 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Turgor Driven Expansion Onset
Turgor loss point determination for Cabernet Sauvignon and Grenache using pressure-volume analysis reveals Ψtlp of -1.65 MPa for Cabernet Sauvignon compared to -2.15 MPa for Grenache, with the 0.5 MPa more negative threshold in Grenache explaining its capacity to maintain stomatal conductance of 80 mmol/m²/s at midday water potential of -1.8 MPa when Cabernet Sauvignon stomata have already closed, demonstrating the mechanistic basis for Grenache's superior performance in drought-prone Mediterranean climates.
Turgor loss point quality is a turgor pressure quality that represents the critical water potential threshold at which cell turgor pressure declines to zero, marking the transition from positive turgor maintaining cell rigidity to flaccid conditions where cells lose structural support and physiological functions dependent on turgor cease. Within the BFO framework, this quality characterizes a specific point on the pressure-volume relationship defining the lower limit of the turgor-positive operating range, distinguishing the threshold value from the continuous turgor variable. The ontological significance lies in turgor loss point serving as a primary indicator of drought tolerance, where species and varieties with more negative turgor loss points maintain positive turgor and physiological function to lower water potentials, enabling continued stomatal opening, photosynthesis, and growth under water deficit conditions. Turgor loss point in grapevines typically ranges from -1.5 to -2.5 MPa depending on variety, acclimation status, and tissue age, with drought-adapted varieties and osmotically adjusted tissues showing more negative values indicating enhanced capacity to maintain turgor under water stress. The determination of turgor loss point employs pressure-volume curve analysis, where excised tissue is progressively dehydrated while measuring water potential and relative water content, with the inflection point marking the transition from linear (turgor-positive) to curvilinear (turgor-zero) portions of the relationship. Osmotic adjustment through solute accumulation shifts turgor loss point to more negative values, representing an acclimation mechanism improving drought tolerance, with grapevines capable of 0.3-0.6 MPa adjustment under progressive stress. Understanding turgor loss point quality enables quantification of drought tolerance, prediction of the water potential range for maintained physiological function, and assessment of acclimation responses to water deficit.
Turgor_Loss_Point_Quality addresses the threshold water potential for turgor maintenance, complementing the continuous Turgor_Pressure_Quality variable. This quality is determined through pressure-volume curve analysis rather than direct measurement, providing a tissue-level parameter integrating cellular osmotic and elastic properties. The class supports drought tolerance characterization and prediction of physiological response to water deficit, with cultivar differences in turgor loss point underlying observed variation in drought adaptation strategies.
Nullturgor-Schwelle
Turgor-Nullpunkt
Turgorverlustpunkt
Welkepunkt
turgor zero point
wilting point
zero turgor threshold
Ψtlp
Bartlett MK et al. (2012) The correlations and sequence of plant stomatal, hydraulic, and wilting responses to drought. Proceedings of the National Academy of Sciences 109(33): 13233-13237 Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell & Environment 26(8): 1393-1405
Turgor Loss Point Quality
Detailed pressure-volume analysis of Turgor_Maintenance_Disposition at the University of Western Australia determined turgor loss point (Ψtlp) and associated parameters across 22 cultivars under well-watered and drought-acclimated conditions. Pressure-volume curves were constructed from sequential measurements of leaf water potential and relative water content during controlled dehydration, enabling calculation of Ψtlp, osmotic potential at full turgor (Ψπ100), and bulk modulus of elasticity (ε). Turgor_Maintenance_Disposition showed substantial cultivar variation. Under well-watered conditions, Ψtlp ranged from -1.45 MPa ('Pinot Noir', early turgor loss) to -2.15 MPa ('Sultana', late turgor loss), representing 0.7 MPa variation in drought tolerance threshold. Following drought acclimation (3 weeks at Ψpd = -0.6 MPa), Ψtlp shifted to more negative values in all cultivars, with the adjustment magnitude ranging from 0.1 MPa ('Pinot Noir') to 0.55 MPa ('Sultana'). The resulting acclimated Ψtlp ranged from -1.55 MPa ('Pinot Noir') to -2.70 MPa ('Sultana'). Functional significance was demonstrated by growth analysis during moderate drought (Ψleaf = -1.8 MPa). At this water potential, high-Turgor_Maintenance_Disposition 'Sultana' (Ψtlp = -2.70 MPa when acclimated) maintained positive turgor and 65% of control shoot elongation rate. Low-Turgor_Maintenance_Disposition 'Pinot Noir' (Ψtlp = -1.55 MPa) experienced turgor loss at -1.8 MPa, showing zero elongation and visible wilting. Correlation analysis across cultivars revealed strong relationship between Ψtlp and drought yield stability (r = 0.78), confirming that Turgor_Maintenance_Disposition directly supports productivity under water-limited conditions.
A Turgor_Maintenance_Disposition is an Osmotic_Adjustment_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to maintain positive cell turgor pressure under declining water potential, preserving turgor-dependent processes including cell expansion, stomatal function, and phloem transport during water stress. This disposition represents the functional outcome of osmotic adjustment, translating biochemical solute accumulation into physiological capacity for continued growth and function under drought. Turgor pressure (the pressure of cell contents against the cell wall) drives cell expansion underlying growth, maintains stomatal aperture enabling gas exchange, powers phloem transport of assimilates, and maintains cell structural integrity. Under water stress, declining water potential would normally reduce turgor toward zero (the turgor loss point, Ψtlp), causing wilting and cessation of turgor-dependent processes. Cultivars with strong Turgor_Maintenance_Disposition lower their turgor loss point through osmotic adjustment, enabling positive turgor at water potentials that would cause wilting in non-adjusting genotypes. The turgor loss point is a key physiological parameter, typically ranging from -1.5 MPa to -2.5 MPa among grapevine cultivars, with lower (more negative) values indicating greater drought tolerance. Turgor_Maintenance_Disposition enables continued shoot growth, functional stomatal regulation, and maintained source-sink relations during drought, directly supporting productivity under water-limited conditions.
Turgor_Maintenance_Disposition is_a Osmotic_Adjustment_Disposition. This class addresses the functional outcome of osmotic adjustment in maintaining cell turgor. The turgor loss point (Ψtlp) is the key parameter, with lower (more negative) values indicating greater drought tolerance. Ψtlp can be lowered through osmotic adjustment (solute accumulation) and/or decreased cell wall elasticity. Turgor maintenance enables continued growth, stomatal function, and phloem transport during drought. Ψtlp is both a constitutive trait (genetic variation) and an acclimated response (adjustment during drought exposure).
Turgor-Aufrechterhaltungs-Disposition
Welke-Resistenz
Zellturgor-Stabilität
cell turgor stability
turgor pressure maintenance
wilting resistance
Bartlett MK et al. (2012) The correlations and sequence of plant stomatal, hydraulic, and wilting responses to drought, Proceedings of the National Academy of Sciences 109:13928-13933 Rodrigues ML et al. (1993) Osmotic adjustment in water stressed grapevine leaves in relation to carbon assimilation, Australian Journal of Plant Physiology 20:309-321 Sancho-Knapik D et al. (2016) The reflectance spectrum of plant leaves and changes caused by drought stress, Functional Plant Biology 43:320-327
Turgor Maintenance Disposition
Turgor pressure monitoring in Cabernet Sauvignon berries using cellular pressure probe reveals pre-véraison values of 0.42±0.08 MPa maintaining berry firmness, declining precipitously to 0.08±0.03 MPa within 10 days of véraison onset as phloem unloading shifts from symplastic to apoplastic pathway, with the 80% turgor reduction explaining the characteristic softening that defines véraison independently of color change in this red variety.
Turgor pressure quality is a physical quality that inheres in grapevine cells and tissues and represents the hydrostatic pressure exerted by the vacuolar contents against the cell wall, arising from osmotic water uptake into cells containing solute concentrations exceeding the surrounding apoplast. Within the BFO framework, this quality characterizes the mechanical force generated by the osmotic gradient across the plasma membrane and tonoplast, where water movement into the cell creates positive pressure that stretches the elastic cell wall until wall resistance balances osmotic driving force at equilibrium. The ontological significance lies in turgor pressure serving as the fundamental driver of cell expansion during growth, the mechanical basis of tissue rigidity determining texture and structural support, and the hydraulic signal integrating plant water status with stomatal regulation and stress responses. Turgor pressure in grapevine cells typically ranges from 0.3-1.0 MPa under well-watered conditions, declining toward zero as water deficit develops and cells lose water to equilibrate with decreasing soil and atmospheric water potential. The relationship between turgor and cell volume follows pressure-volume curves characterizing the elastic properties of cell walls, where rigid walls maintain turgor with minimal volume change while elastic walls allow substantial volume reduction before turgor loss. Turgor pressure measurement employs pressure probe techniques for direct cellular measurement, thermocouple psychrometry for tissue water potential from which turgor is calculated, and pressure-volume analysis providing bulk tissue turgor estimates. The developmental dynamics of turgor are particularly significant in grape berries, where the transition from hard pre-véraison berries with high turgor to soft post-véraison berries involves turgor decline from approximately 0.4 MPa to near zero despite continued sugar accumulation, reflecting the uncoupling of berry water relations from vine hydraulic system at véraison. Understanding turgor pressure quality enables prediction of growth dynamics, interpretation of texture changes during ripening, and assessment of water stress impacts on vine physiology and fruit quality.
Turgor_Pressure_Quality serves as the parent class for cell-type-specific turgor qualities and related hydraulic properties (Turgor_Loss_Point, Osmotic_Potential), establishing the ontological framework for plant water relations at cellular scale. This quality represents the pressure component (Ψp) of the water potential equation Ψ = Ψs + Ψp. Direct measurement requires specialized equipment, with most field assessments relying on calculated values from water potential and osmotic potential measurements.
Druckpotential
Turgordruck
cell pressure
cell turgor
hydrostatic pressure
hydrostatischer Druck
pressure potential
Tyerman SD et al. (2004) Direct measurement of hydraulic properties in developing berries of Vitis vinifera L. cv Shiraz and Chardonnay. Australian Journal of Grape and Wine Research 10(3): 170-181 Thomas TR et al. (2006) Mesocarp cell turgor in Vitis vinifera L. berries throughout development and its relation to firmness, growth, and the onset of ripening. Planta 224(5): 1078-1088
Turgor Pressure Quality
Pinot Noir in Burgundy shows turgor reduction possible initiation during late August-early September Year N+1 at BBCH 83, with cell turgor pressure possibly declining 12-14% from 0.62 to 0.54 MPa during initial softening representing minor 0.08 MPa reduction, osmotic potential becoming more negative from sugar increase -0.8 to -1.1 MPa, cell wall yielding from structural modification enabling volume expansion, turgor contribution to firmness decline estimated 18-24% while cell wall structural changes contributing 76-82%, turgor as secondary softening factor, under continental conditions of 22-27°C temperature, typical of this early-to-mid season cultivar showing minor turgor changes compared to dominant structural softening
Turgor reduction initiation represents the cell pressure decline onset phase during initial berry softening when berry cell turgor pressure possibly begins slight reduction from altered water relations, turgor potentially declining 10-15% from pre-veraison 0.5-0.8 MPa to initial 0.4-0.7 MPa, turgor changes minor compared to cell wall structural modifications which contribute 70-85% firmness decline while turgor contributes 15-30%, turgor reduction as secondary softening factor, and turgor changes interacting with wall modifications affecting overall berry texture at veraison onset. This pressure reduction onset phase occurs during late summer to early autumn in late August-early September in temperate regions Year N+1, representing possible turgor change at veraison though contribution minor. Turgor reduction mechanism involves possible changes in cell osmotic potential from sugar and acid dynamics, possible changes in water potential affecting water influx and turgor, altered membrane properties possibly affecting water relations, cell wall yielding enabling volume expansion reducing pressure, and minor turgor reduction as secondary contributor to softening compared to dominant structural changes. Turgor pressure definition shows turgor as hydrostatic pressure in cell from water influx against cell wall resistance, turgor generated by osmotic potential difference between cell and apoplast, turgor pressing cell membrane against wall, turgor contributing to cell and tissue rigidity, and turgor as cell mechanical property. Pre-veraison turgor shows turgor pressure typically 0.5-0.8 MPa during lag phase, pressure from osmotic gradient and water uptake, turgor at moderate levels with cell walls relatively rigid preventing expansion, and pre-veraison as baseline state. Turgor measurement shows pressure probe or pressure chamber techniques measuring cell pressure, measurements showing possible slight decline during softening, measurement challenges in berry cells, and turgor quantification difficult. Possible turgor decline shows turgor potentially declining 10-15% from pre-veraison 0.5-0.8 MPa to 0.4-0.7 MPa during initial softening representing minor 0.05-0.12 MPa reduction, turgor changes less dramatic than structural changes, turgor decline not universal or well-documented across studies, and possible but uncertain turgor reduction. Osmotic potential changes shows berry osmotic potential becoming more negative from sugar accumulation, increased solutes should increase turgor but effects complex, possible offsetting changes in apoplast, and osmotic dynamics affecting turgor. Sugar accumulation effects shows sugars increasing during veraison creating more negative osmotic potential, sugar increase potentially increasing turgor pressure, but cell wall yielding enabling expansion so pressure may not increase or may decline, and complex sugar-turgor relationship. Organic acid decline effects shows acids declining during ripening, acid reduction potentially affecting osmotic potential and turgor, effects likely minor compared to sugar, and acid-turgor interaction. Water potential changes shows berry water potential possibly changing from altered osmotic and pressure potentials, water uptake continuing during softening from osmotic gradient, water relations complex, and water potential as turgor determinant. Cell wall yielding shows wall softening from structural modification enabling cell expansion, expansion allowing volume increase reducing pressure for given water content, yielding as mechanism enabling turgor reduction or preventing increase, and wall-turgor interaction. Cell volume changes shows cells possibly expanding during veraison-ripening from water uptake and wall yielding, volume increase potentially reducing turgor if water uptake insufficient to maintain pressure, and volume-turgor relationship. Membrane properties shows membrane permeability possibly changing affecting water flux, aquaporin expression and activity possibly altering water transport, membrane changes affecting turgor regulation, and membrane as turgor factor. Turgor contribution to firmness shows turgor contributing 15-30% firmness in most studies, structural wall changes contributing 70-85%, turgor as secondary factor, distinction from wall effects difficult, and relative contribution as key concept. Firmness from turgor shows turgor-generated pressure pressing cell against wall contributing to rigidity, reduced turgor creating softer feel, but effect much less than structural wall changes, and turgor as minor firmness component. Cell wall structural changes dominance shows pectin degradation and wall modifications as primary softening mechanisms, wall changes reducing firmness 70-85%, turgor changes secondary 15-30%, and structural dominance as softening basis. Turgor-wall interaction shows turgor and wall properties interacting to determine firmness, soft walls with high turgor versus rigid walls with low turgor creating different textures, interaction complex, and combined effects determining texture. Turgor role uncertainty shows turgor changes during softening not well-established, some studies showing minor decline, others no change or increase, variability across cultivars and conditions, and uncertain turgor dynamics. Measurement challenges shows turgor measurement technically difficult in berry cells, pressure probe requiring careful technique, indirect estimates from water relations, and measurement limitations. Cultivar variation possible shows different cultivars possibly showing different turgor dynamics, some with minor decline others stable, variation complicating generalizations, and genetic effects possible. Environmental effects on turgor include temperature affecting water relations and turgor, water availability affecting berry hydration and pressure, and environmental factors influencing turgor though effects less clear than structural changes. Temperature effects shows temperature affecting membrane permeability and water flux potentially affecting turgor, and temperature as turgor factor. Water stress effects shows water deficit potentially reducing turgor through limited water availability, deficit affecting berry hydration, and water stress as major turgor influence. The process represents third aspect of initial berry softening as possible minor pressure reduction, following cell wall modification onset and pectin solubilization start. Duration possible turgor reduction during initial softening days. Regional timing shows turgor reduction possible initiation during late August-early September at veraison. Viticultural significance includes representing possible secondary softening contributor, demonstrating cell wall structural changes as dominant mechanism, showing turgor-wall interaction in texture determination, and establishing turgor as minor firmness factor compared to structural changes. Research applications include analyzing turgor dynamics during softening, investigating turgor-firmness relationships and contributions, evaluating water relations and osmotic changes, quantifying turgor changes and mechanisms, and assessing turgor reduction as potential minor softening component.
Turgor_Reduction_Initiation is third aspect of Initial_Berry_Softening, distinguished by possible pressure reduction. Represents possible 10-15% decline from 0.5-0.8 to 0.4-0.7 MPa. SECONDARY softening factor contributing 15-30% firmness decline versus cell wall structural changes 70-85% (DOMINANT). Turgor changes minor, not well-established, variable across studies. Osmotic potential more negative from sugars. Cell wall yielding enables volume expansion. Follows Cell_Wall_Modification_Onset and Pectin_Solubilization_Start. Temperature and water stress affect. Uncertain contribution.
Druck-Reduktions-Beginn
Turgor-Abnahme-Start
Turgor-Reduktions-Initiierung
Zelldruck-Reduktion
cell pressure reduction
minor turgor change
pressure reduction onset
turgor decline start
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 480-482
Turgor Reduction Initiation
Merlot hermaphrodite flowers at BBCH 59-61 exhibit typical two carpel compound pistils with bilocular ovaries 0.9-1.1 mm diameter showing two distinct chambers separated by central septum, 4 ovules distributed as 2 per locule in axile placentation, unified style 0.6-0.8 mm, and stigma 0.45-0.55 mm diameter showing subtle bilobed appearance with enhanced papillae density along two opposite margins corresponding to the two carpel stigmatic surfaces. Chardonnay pistils in transverse anatomical sections clearly reveal two-carpel organization through bilocular ovary structure with two chambers of approximately equal size, central septum representing fused carpel margins, and ovules attached to septum surfaces in positions corresponding to original carpel placentation. Pinot Noir fruit set analysis shows that berries typically contain 1-3 seeds (from 2-4 original ovules in two-carpel pistil), with seed distribution between the two original locules variable (both seeds in one locule, seeds distributed 1-2 per locule, or occasionally locule asymmetry with most seeds in one original chamber), demonstrating independent fertilization and seed development potential of the two-carpel system. Sauvignon Blanc occasional flowers (<0.08% frequency) show abnormal monocarpellate pistils with single locule and 1-2 ovules resulting from developmental failure of one carpel or complete septum degeneration, producing berries with unusual morphology and reduced seed number potential.
A two carpel compound pistil is a compound pistil formed specifically from two fused carpels creating the bicarpellate syncarpous structure characteristic of all normal Vitis flowers, with two-carpel organization evident in bilocular ovary structure (two chambers), two-sided ovule arrangement, and often bilobed or bilaterally organized stigma, representing the universal and ancestral pistil organization across Vitis species, cultivars, and flower types. The two carpel organization is evident through multiple anatomical and developmental features= (1) bilocular ovary with two distinct chambers (locules) separated by a central partition (septum) formed from the fused medial (inner) margins of the two carpels, with each locule representing the original cavity of one carpel before fusion (2) axile placentation with ovules attached to the central septum in two rows or groups corresponding to the two carpel margins, with typical ovule number being 2-4 total (commonly 1-2 per locule, occasionally 2 per locule) (3) stigma often showing bilobed appearance with two slightly separated lobes or showing bilateral papillae distribution with denser papillae along two opposite sides corresponding to the two carpel stigmatic surfaces (4) style sometimes showing internal bilateral organization in transmitting tract tissue reflecting the two-carpel origin (5) developmental sequence showing two carpel primordia initiating separately during early flower development (BBCH 51-52) before fusing during subsequent development (BBCH 52-53). The two carpel number represents a stable, highly conserved character in Vitis, with essentially all flowers across all species and cultivars showing bicarpellate organization, distinguishing Vitis from plant families with tricarpellate (three carpel), pentacarpellate (five carpel), or highly variable carpel number pistils. The evolutionary stability of two-carpel organization in Vitis suggests strong developmental constraints or strong selection for bicarpellate structure, possibly relating to optimal fruit size, seed number, or developmental coordination. The two carpel structure creates functional consequences for reproduction and fruit development= (1) bilocular organization provides spatial separation of ovules, potentially allowing independent fertilization events in the two locules and creating potential for differential seed set between locules (berries commonly containing 1-4 seeds distributed variably between the two locules) (2) two-carpel ovary size and structure determines initial berry size potential and influences final berry morphology (3) bilateral stigma organization may influence pollen distribution and pollen tube guidance patterns affecting fertilization efficiency. Abnormalities occasionally produce flowers with apparent one-carpel (monocarpellate, unilocular) or three-carpel (tricarpellate, trilocular) organization, but these are rare (<0.1% frequency) and represent developmental disruptions rather than stable alternative states, with monocarpellate flowers resulting from failure of one carpel to develop or from complete septum degeneration, and tricarpellate flowers resulting from supernumerary carpel initiation. The two carpel organization interacts with other floral traits in determining fruit set and berry characteristics= fruit set percentage reflects successful fertilization of at least one ovule (typically requiring fertilization of 1-2 ovules for stable fruit retention) seed number per berry reflects number of successfully fertilized ovules (typically 1-4 seeds from 2-4 ovules) seedless berries (parthenocarpic fruit development) can develop from two-carpel pistils without fertilization in stenospermic (seed-trace) cultivars like Thompson Seedless where fertilization initiates development but seeds abort early.
Two_Carpel_Compound_Pistil specifies the bicarpellate nature of the compound pistil universal in Vitis and is essentially synonymous with normal pistil organization in grapevines. The two-carpel specification emphasizes the carpel number as a fundamental organizational feature determining ovary chamber number, ovule distribution, and stigma organization. This class is distinguished from hypothetical single-carpel or multi-carpel (>2) organizations which are not characteristic of normal Vitis flowers. Understanding the two-carpel structure is essential for interpreting ovary anatomy, ovule number and distribution, fertilization patterns, seed number determination, and fruit development. The term overlaps substantially with Bicarpellate_Pistil but emphasizes the compound (fused) nature explicitly.
Zwei-Fruchtblatt-Stempel
Zweikarpell-Stempel
bicarpellate pistil
bikarpellates Gynoeceum
two-carpel gynoecium
Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 92-95 Carmona, M.J. et al. (2008). Grapevine inflorescence and flower development, pp. 186-188.
Two Carpel Compound Pistil
Merlot ovaries at BBCH 59-61 exhibit invariant two chamber bilocular organization visible in transverse anatomical sections as exactly two locule cavities (left and right chambers) separated by central septum, with the two chambers containing 3-4 ovules distributed as 1-2 per chamber, representing the universal two-chamber organization reflecting bicarpellate pistil structure. Riesling ovary serial sectioning studies from base to apex reveal two elongated chambers extending throughout ovary length (0.8-1.2 mm), with the two chambers maintaining separation via continuous septum running the full length, and with ovules (2-4 total) attached to septum surfaces projecting into the two chamber cavities. Sauvignon Blanc berries at maturity (BBCH 89) containing 2-3 seeds show seed distributions reflecting original two-chamber ovary organization, with 2-seed berries typically showing seeds positioned in the two original locule locations (one seed per original chamber) and 3-seed berries showing asymmetric distributions (2 seeds in one original chamber, 1 in the other), demonstrating that two-chamber organization influences final seed positioning patterns even after extensive berry development and tissue reorganization. Gewürztraminer rare abnormal flowers (<0.05% frequency) exhibit unilocular ovaries with single chamber resulting from complete septum absence, producing berries with unusual internal structure lacking the typical two-compartment seed distribution, confirming that two-chamber organization represents the normal condition and that deviations produce recognizably abnormal outcomes.
A two chamber bilocular ovary is a bilocular ovary specifically characterized by the presence of exactly two distinct internal chambers (locules), emphasizing the dual-compartment structure as the defining organizational feature of normal Vitis ovaries reflecting the invariant bicarpellate pistil construction. Two chamber bilocular ovaries are characterized by= (1) exactly two locules present without exception in normal flowers, representing the cavities of the two fused carpels (2) the two chambers showing variable relative sizes ranging from nearly equal (symmetric bicarpellate pistils with locule volumes varying 20-60%) (3) each chamber capable of containing 0-2 ovules depending on ovule initiation success during development, with typical ovule distributions being 2 per chamber (4 total), 1-2 per chamber (3 total), or 1 per chamber (2 total) (4) two-chamber organization maintained throughout ovary development from initial locule differentiation (BBCH 53-54) through anthesis (BBCH 61-65) and into early fruit development (BBCH 71-75), though chamber boundaries may become obscured during later berry development as flesh proliferates. The two-chamber number is invariant in normal Vitis flowers across all species, cultivars, and flower sexual types, representing one of the most stable and conserved features of Vitis reproductive anatomy and reflecting the fundamental bicarpellate nature of the pistil. This constancy of two chambers contrasts with variation in other ovary features= ovule number varies (2-4 typically, occasionally 5-6), chamber relative sizes vary (symmetric versus asymmetric), ovary overall size varies (larger in functionally female/pistillate flowers, smaller in functionally male/staminate flowers), but chamber number remains invariantly two. The two-chamber organization creates functional compartmentalization= each chamber represents an independent space for ovule development and potential seed maturation fertilization and seed development can proceed independently in the two chambers, creating berries with variable seed distribution between the two original compartments (both seeds in one chamber, seeds distributed between chambers, or asymmetric distributions) chamber separation may provide some developmental buffering, allowing successful seed development in one chamber even if the other chamber experiences ovule abortion or fertilization failure. Anatomical evidence for two-chamber organization includes= transverse sections at mid-ovary level showing two distinct locule cavities separated by septum longitudinal sections showing two locules extending from ovary base to apex three-dimensional reconstructions from serial sections revealing two elongated chambers running parallel through the ovary length. The two-chamber condition persists with high fidelity through berry development, evident in seed distribution patterns in mature berries that reflect the original two-locule ovary organization= berries with 2 seeds often show seeds positioned in the two original locule locations berries with 3-4 seeds show distributions corresponding to 1-2 seeds per original locule even seedless berries sometimes show residual seed trace positions in two-locule arrangement. Rare developmental abnormalities can produce non-two-chamber conditions= unilocular ovaries (single chamber) resulting from complete septum failure or single carpel development trilocular or multilocular ovaries (three or more chambers) from supernumerary carpel development or abnormal septum formation but these represent <0.1% of flowers and are recognized as developmental disruptions rather than normal variation.
Two_Chamber_Bilocular_Ovary emphasizes the dual-compartment aspect of bilocular organization and is essentially synonymous with normal bilocular ovary structure, with the two-chamber specification highlighting the invariant chamber number as a defining feature. This class overlaps with Septate_Bilocular_Ovary (which emphasizes the septum creating the two chambers) as complementary perspectives on bilocular organization. The two-chamber condition is universal in normal Vitis and represents one of the most stable reproductive anatomy features. Understanding two-chamber organization is essential for interpreting ovule distribution, seed positioning, and berry internal structure, and for recognizing rare developmental abnormalities.
Zwei-Kammer-Fruchtknoten
bilokulares Zwei-Fach-Ovar
dual-chamber ovary
two-compartment ovary
zweifächeriger Fruchtknoten
Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 93-94.
Two Chamber Bilocular Ovary
A 16 cm two-node scion cutting of Cabernet Sauvignon clone 337 bears a Two_Node_Cutting_Role, prepared for omega grafting to SO4 rootstock—the lower bud positioned at the graft union while the upper bud provides shoot growth, representing the standard benchgraft scion configuration. A two-node hardwood cutting of 3309 Couderc rootstock serves in a Two_Node_Cutting_Role for direct field rooting, with both nodes buried and the redundant bud system improving establishment success in field conditions versus more demanding nursery environments. A two-node cutting of Thompson Seedless bears a Two_Node_Cutting_Role in own-rooted table grape propagation, rooted in containers for vineyard establishment in phylloxera-free production regions.
A cutting role that inheres in a stem segment prepared with two nodes bearing two buds plus associated internode tissue (typically 15-20 cm total length), representing an intermediate format balancing multiplication efficiency against propagule vigor and establishment success. Two-node cuttings provide redundancy (if one bud fails, the second may succeed), increased carbohydrate reserves compared to single-node format, and adequate tissue for standard benchgrafting operations where scion length must accommodate the grafting cut while retaining at least one intact bud. The two-node format is common for scion preparation in benchgraft production, where a two-bud scion grafted to a rootstock cutting provides the standard commercial grafted vine configuration. Two-node cuttings root reliably under standard nursery conditions without the intensive environmental control required for single-node propagation, while still achieving reasonable multiplication rates (7-10 propagules per meter of source cane for standard internode lengths).
Two_Node_Cutting_Role represents the intermediate format between intensive single-node propagation and traditional three-node (or longer) cuttings, balancing multiplication efficiency against establishment reliability. The format is standard for benchgraft scion preparation where omega or whip-and-tongue grafting techniques create the union at or just below the lower node, leaving one functional bud above the graft—the resulting two-bud scion on rootstock being the archetypal benchgraft configuration. For rootstock cutting propagation, two-node formats offer improved success over single-node under standard conditions while doubling multiplication rate compared to three-node. Cutting quality assessment for two-node format evaluates both buds (one may be primary while the other serves as backup) and internode characteristics (diameter uniformity, freedom from injury or infection).
Doppelaugensteckling
Zweiaugensteckling
Zweiknoten-Steckling
bi-nodal cutting
double-eye cutting
two-bud cutting
two-node cutting
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 350-355 Dry N (2007) Grapevine rootstocks, Lythrum Press Gramaje D, Armengol J (2011) Fungal trunk pathogens, Phytopathologia Mediterranea 50: S217-S226
Two Node Cutting Role
Riesling (low ovule number clones) consistently produces 45-58% of flowers with two ovule ovaries containing one ovule per locule in symmetric distribution, with these two ovule flowers achieving 40-48% fruit set producing berries with 1.4-1.8 seeds on average (reflecting 70-90% fertilization success of the 2 available ovules), while the remaining 42-55% of flowers contain 3-4 ovules producing berries with 2.2-3.1 seeds when fruit set occurs. Nebbiolo shows constitutive two ovule ovary frequency of 32-48% across diverse sites and vintages, with two ovule ovaries distributed as single ovule per locule in bilocular organization, achieving fruit set rates (42-50%) comparable to four ovule ovaries (44-52%) demonstrating that ovule number has limited impact on fertilization probability within this range. Cabernet Sauvignon under water stress during BBCH 54-55 (ovule initiation phase) exhibits increased two ovule ovary frequency (28-35% versus normal 8-14% under adequate water) due to stress-induced reduction in ovule initiation, with stressed vines producing berries with reduced average seed number (1.7-2.2 seeds per berry versus normal 2.4-3.1) reflecting the ovule number shift. Pinot Noir berry analysis shows that berries developing from two ovule ovaries average 9.8-11.2 mm diameter versus 10.5-12.1 mm diameter from four ovule ovaries (8-10% size reduction), suggesting that ovule/seed number influences berry growth potential through embryo-derived hormone production affecting cell division and expansion during berry development.
A two ovule ovary is an ovule containing ovary that contains exactly two ovules, typically distributed as one ovule per locule in the bilocular Vitis ovary organization, representing the lower end of the normal ovule number range and characteristic of certain cultivars that consistently produce reduced ovule numbers or representing occasional flowers in cultivars that typically produce 3-4 ovules. Two ovule ovaries are characterized by= (1) exactly two ovules present, typically one per locule in the standard bilocular organization, occasionally both in one locule if locule asymmetry or ovule development asymmetry occurs (2) ovules positioned on septum surfaces in axile placentation at positions corresponding to the two fused carpel margins (3) ovary otherwise showing normal bilocular structure with two locules separated by septum, but with only single ovule per locule rather than the more common two ovules per locule (4) ovules showing normal mature structure at anthesis with completed megasporogenesis and viable embryo sacs when ovule development is successful. Two ovule ovaries occur through several mechanisms= (1) genetic determination in cultivars with constitutively low ovule number, where most flowers consistently produce only 2 ovules (2) developmental variation in cultivars that typically produce 3-4 ovules, where 5-25% of flowers may produce only 2 ovules through stochastic developmental variation (3) environmental stress effects during ovule initiation (BBCH 54-55) causing reduced ovule number through resource limitation or developmental disruption (4) ovule abortion after initial formation, reducing an originally 3-4 ovule complement to 2 viable ovules by anthesis. Two ovule ovaries have functional implications for reproduction and berry characteristics= reduced ovule number (2 versus 3-4) decreases seed number potential, with successful fertilization of both ovules producing berries with 2 seeds and partial fertilization producing berries with 1 seed or occasional seedless berries if both ovules fail fertilization reduced ovule number may slightly reduce fruit set probability if fertilization of at least one ovule is required for stable fruit retention, though differences are typically modest (38-48% fruit set in two ovule flowers versus 42-52% in four ovule flowers in typical hermaphrodite cultivars) berries from two ovule ovaries tend to be smaller on average than berries from four ovule ovaries (8-10% diameter reduction) when other factors are equal, possibly due to reduced embryo/endosperm-derived growth hormone production from fewer developing seeds. Certain wine cultivars show constitutively high frequencies of two ovule ovaries= some Riesling clones produce 40-60% of flowers with only 2 ovules some Nebbiolo selections show 30-50% two ovule ovaries while most Cabernet Sauvignon, Merlot, and Chardonnay clones show <15% two ovule ovaries. The two ovule condition should be distinguished from four ovule ovaries (typical high ovule number) and from three ovule ovaries (intermediate, very common), representing distinct points on the ovule number continuum. Environmental modulation can shift ovule number distributions= favorable conditions during ovule initiation (adequate water, nutrients, moderate temperatures during BBCH 54-55) promote higher ovule numbers reducing two ovule ovary frequency stress conditions increase two ovule ovary frequency through reduced ovule initiation or increased ovule abortion.
Two_Ovule_Ovary represents the lower end of normal ovule number range and is distinguished from Four_Ovule_Ovary (upper end of typical range) and from three ovule ovaries (intermediate, not specifically classified in current hierarchy). The two ovule condition occurs constitutively in some cultivars/clones and variably in others, and can be influenced by environmental conditions during ovule development. Understanding two ovule versus higher ovule number ovaries is relevant for predicting seed number, for assessing environmental stress effects, and for understanding cultivar variation in reproductive characteristics. Two ovule ovaries are compatible with successful fruit set and berry development though producing berries with reduced seed number and potentially slightly reduced size.
Zwei-Eizellen-Ovar
Zwei-Ovulum-Fruchtknoten
dual-ovule ovary
two-seed-potential ovary
zweieiiger Fruchtknoten
Pratt, C. (1971). Reproductive anatomy in cultivated grapes. American Journal of Enology and Viticulture 22=92-109 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 94-98.
Two Ovule Ovary
Detailed anatomical analysis of Tylosis_Formation_Disposition at UC Davis compared tylosis production in response to Phaeomoniella chlamydospora trunk inoculation across 10 scion cultivars and 5 rootstocks. Vascular casts and scanning electron microscopy quantified tylosis density, coverage, and wall thickness in vessels at 5, 10, and 20 mm from inoculation sites at 4 months post-inoculation. 'Petit Verdot' exhibited the strongest Tylosis_Formation_Disposition with tylosis density of 12.4 per mm vessel length, 85% vessel lumen occlusion, and tylosis wall thickness of 1.8 μm (heavily lignified). In contrast, susceptible 'Thompson Seedless' showed density of 3.2 per mm, only 25% occlusion, and thin walls of 0.6 μm. Pathogen colonization extent inversely correlated with tylosis parameters (r = -0.78 for density, -0.82 for occlusion percentage). Importantly, rootstock genotype significantly influenced scion tylosis formation, with 'Petit Verdot' on '110 Richter' producing 35% more tyloses than on '101-14 Mgt', indicating that rootstock-mediated signals modulate this defense response. Hydraulic conductivity measurements confirmed that heavily tylosed 'Petit Verdot' vines maintained adequate water transport (85% of healthy controls) despite partial vessel occlusion, demonstrating effective partitioning between infected and functional vascular sectors.
A Tylosis_Formation_Disposition is an Esca_Resistance_Disposition that inheres in a Vitis vinifera plant and is realized through the production of tyloses — balloon-like outgrowths of xylem parenchyma cells that protrude through pit membranes into adjacent vessel lumens, occluding the vessel and preventing longitudinal movement of pathogens and their phytotoxins through the vascular system. This disposition represents a specialized vascular defense mechanism critical for restricting the spread of esca-associated fungi (Phaeomoniella chlamydospora, Phaeoacremonium species) that colonize and move through functional xylem vessels. Effective tylosis formation involves the perception of pathogen presence or vascular dysfunction, activation of parenchyma cell expansion, targeted deposition of tylosis walls containing suberin, lignin, and phenolic compounds, and coordination of tylosis production to create continuous vessel blockages without excessive compromise of water transport capacity. The speed, extent, and chemical fortification of tylosis responses vary substantially among grapevine cultivars, correlating with differential susceptibility to trunk diseases. Cultivars with strong Tylosis_Formation_Disposition rapidly produce dense, lignified tyloses in vessels adjacent to infection sites, creating effective barriers against pathogen spread, while susceptible cultivars produce sparse, thin-walled tyloses that pathogens readily breach. However, excessive tylosis formation can impair vine water relations, particularly under drought stress, requiring balance between pathogen containment and maintenance of hydraulic function. Understanding tylosis formation dispositions informs both cultivar selection and development of elicitor treatments that could enhance this natural defense response.
Tylosis_Formation_Disposition is_a Esca_Resistance_Disposition. Tyloses are formed by xylem parenchyma cells (specifically, contact cells adjacent to vessels) and represent a specialized vascular defense distinct from general wound compartmentalization. Tylosis production is triggered by both pathogen infection and abiotic stresses (drought, wounding), requiring careful experimental controls to assess pathogen-specific responses. Rootstock genotype influences scion tylosis formation, adding complexity to resistance phenotyping. Excessive tylosis can impair water transport, particularly under drought, necessitating balance between defense and hydraulic function.
Gefäßverschluss-Kapazität
Thylle-Verteidigung
Thyllenbildungs-Disposition
tylosis defense disposition
vascular occlusion response
vessel occlusion capacity
Pouzoulet J et al. (2014) Xylem vessel diameter affects the compartmentalization of the vascular pathogen Phaeomoniella chlamydospora in grapevine, Frontiers in Plant Science 5:253 De Micco V et al. (2016) Xylem anatomical traits reveal different strategies of two Mediterranean oaks to cope with drought and warming, Environmental and Experimental Botany 133:128-138 Sun Q et al. (2013) Xylem occlusion in grapevine to limit the systemic spread of Pierce's disease, American Journal of Enology and Viticulture 64:153-159
Tylosis Formation Disposition
The Chasselas accession maintained as herbarium voucher specimen P00680243 at the Muséum National d'Histoire Naturelle in Paris bearing a type specimen role serves as the nomenclatural type for Chasselas Blanc (syn. Gutedel, Fendant), with the accompanying living reference accession at INRAE Domaine de Vassal (accession Vas 0060Mtp1) providing vegetative material for morphological verification against the 68 OIV primary descriptors and DNA extraction for the definitive SSR profile that resolves synonymy disputes — confirming, for example, that Swiss Fendant, German Gutedel, and Hungarian Chassla are genetically identical to the French type despite centuries of independent cultivation and nomenclatural divergence.
A reference cultivar role that inheres in a grapevine accession formally designated as the nomenclatural type for a cultivar name, serving as the definitive physical standard against which the identity of any vine claimed to belong to that cultivar is ultimately verified through morphological comparison or genetic fingerprint matching.
Type_Specimen_Role applies a concept from biological nomenclature (the type specimen as nomenclatural anchor) to cultivated grapevine taxonomy. Unlike wild species where holotypes are formally designated under the International Code of Nomenclature, grapevine cultivar nomenclature operates under the International Code of Nomenclature for Cultivated Plants (ICNCP) which recommends but does not strictly require type designation: in practice, the VIVC (Vitis International Variety Catalogue) and INRAE Domaine de Vassal collection serve as de facto type repositories for the global grapevine cultivar nomenclature system, maintaining approximately 2,800 authenticated reference accessions.
Nomenklatur-Standard-Rolle
Referenzpflanzen-Rolle
Typus-Exemplar-Rolle
holotype vine role
nomenclatural standard role
type accession role
ICNCP (2016) International Code of Nomenclature for Cultivated Plants 9th Ed, ISHS Lacombe T et al. (2011) Inventory and characterization of Vitis vinifera ssp. vinifera at INRA Domaine de Vassal, Vitis 50: 56-68 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157
Type Specimen Role
Typicity assessment for Grüner Veltliner from the Wachau (2024 vintage) rates as "highly typical to exemplary" — the wine expresses the full spectrum of expected Grüner Veltliner varietal markers (white pepper spiciness, citrus, stone fruit) within the site-specific Wachau framework (mineral intensity, structural precision from primary rock terroirs, concentrated Smaragd weight class), and would be immediately identifiable as Wachau Grüner Veltliner in a blind tasting by an experienced Austrian wine professional. Typicity assessment for an atypical Sauvignon Blanc from Marlborough (2024 vintage, barrel-fermented, full MLF, 15 months in new French oak) rates as "atypical for region" — while well-made, the dominant oak influence (vanilla, toast, butter) and suppressed varietal thiol expression (passionfruit and grapefruit character masked by barrel fermentation) renders the wine unrecognizable as Marlborough Sauvignon Blanc in a blind tasting context, illustrating how winemaking decisions can override both varietal and regional typicity markers. Typicity assessment for Blaufränkisch Mittelburgenland DAC (2023 vintage) evaluates as "highly typical" — the dark cherry fruit, fine-grained tannin, spice complexity, and moderate body conform precisely to the DAC style definition.
Typicity assessment is an overall assessment entity that records the degree to which a wine expresses the characteristic sensory attributes expected for its declared cultivar identity, geographical origin, vintage conditions, and production style, evaluated through comparison against the mental reference prototype (the idealized sensory model of what that wine "should" taste like) held by experienced tasters, assessed on a scale from atypical (not recognizable as an example of the declared type) through moderate typicity (some expected characteristics present but partially obscured or modified) to highly typical (immediately recognizable, expressing all expected hallmarks of the type) and exemplary (a reference-quality expression that defines or redefines the standard for its category), and providing the identity-verification and terroir-expression assessment that is central to appellation approval panels, classification tastings, and the concept of regional wine authenticity.
Typicity assessment engages one of the most philosophically complex concepts in wine evaluation — the notion that a wine has an essential character defined by its origin and cultivar that can be recognized, evaluated, and judged against an idealized standard. The cognitive basis of typicity assessment involves prototype matching — the taster compares the perceived sensory profile against an internalized mental prototype constructed from repeated exposure to exemplary wines of the declared type, with the quality of this mental reference dependent on the breadth and depth of tasting experience. This makes typicity assessment inherently expertise-dependent — novice tasters lack the reference prototypes necessary for meaningful typicity evaluation, while highly experienced regional specialists possess nuanced prototypes that can distinguish not only cultivar and regional typicity but sub-regional, vineyard-specific, and producer-specific signatures. The concept of typicity operates at multiple hierarchical levels simultaneously — a wine may be typical of its cultivar (expressing Grüner Veltliner varietal character) but atypical of its region (not expressing Wachau site character), or typical of its region but atypical of its classification (not meeting Smaragd weight expectations within the Wachau), or typical at all levels but atypical of its vintage (not expressing the expected cool-vintage freshness of 2024). The regulatory application of typicity in appellation approval panels creates a normative function — wines must demonstrate typicity to receive their appellation designation, establishing typicity as a quality gate rather than merely an evaluative parameter. The tension between typicity (conformity to expected type) and individuality (distinctive personal expression) represents one of the enduring aesthetic debates in wine culture — the most celebrated wines often demonstrate both high typicity (immediately recognizable origin identity) and singular individuality (a unique personal signature within the typicity framework), suggesting that typicity and originality are complementary rather than contradictory quality dimensions.
Gebietstypizität
Typizitätsbeurteilung
Typizitätsbewertung
regional character
terroir expression evaluation
typicality assessment
varietal typicity
wine identity assessment
Maitre I et al. (2010) Sensory typicality of wines: how scientists have recently dealt with this subject. Food Quality and Preference 21(7): 831-838 Parr WV et al. (2007) The distinctive flavour of New Zealand Sauvignon Blanc: sensory characterisation by wine professionals. Food Quality and Preference 18(6): 849-861 Ballester J et al. (2005) The odor of colors: can wine experts and novices distinguish the odors of white, rosé, and red wines? Chemosensory Perception 2(4): 203-213 Saliba AJ et al. (2009) Impact of information about origin on wine liking. Food Quality and Preference 20(1): 56-62
Typicity Assessment
An ultimate terminal leaf at node 18 on a shoot of Vitis vinifera cv. Merlot in Pomerol, France, representing the final leaf initiated before shoot extension cessation in early August, measuring 9.5 cm length by 8.8 cm width representing 52% of mid-shoot leaf area, exhibiting incomplete development with partial expansion arrested at approximately 65% of potential final size when shoot extension ceased due to combination of water stress (soil moisture 45% field capacity) and developmental signals associated with veraison initiation, showing pale green coloration with chlorophyll content of 320 μg cm⁻² reflecting incomplete maturation, moderately firm texture with partial cell wall development, incomplete tissue differentiation with developing palisade layer of 48 μm (compared to mature 85 μm), demonstrating limited photosynthesis rate of 4.8 μmol CO₂ m⁻² s⁻¹ representing 37% of mature capacity and marginal carbon balance with daily photosynthetic production approximately equaling respiratory costs, positioned at shoot terminus with no younger leaves above, occurring following shoot extension cessation at BBCH stage 81 (beginning of berry ripening) measured in mid-August, representing permanent terminal leaf status that will persist through dormancy and potentially into subsequent season if shoot is retained as fruiting cane during winter pruning.
A terminal apical leaf representing the final leaf initiated before shoot extension cessation in Vitis vinifera, characterized by position at the absolute shoot terminus after growth arrest, status as the youngest and typically smallest fully formed leaf on the non-extending shoot, dimensions often reduced compared to median leaves typically 40-65% of mid-shoot leaf area reflecting late-season initiation, late emergence timing occurring as shoot extension rates decline preceding complete growth cessation, incomplete or arrested development common with many ultimate terminal leaves failing to achieve full expansion before shoot growth arrest, variable maturation depending on timing of extension cessation with early arrest (mid-season) allowing maturation while late arrest (approaching dormancy) often leaving ultimate terminal leaves partially developed, functional role transitioning from active sink during development to potential limited source if maturation completes or remaining as non-functional sink if development arrests incomplete, and permanent positional status as the terminal leaf throughout dormancy and into subsequent growing season if the shoot is retained during pruning. Ultimate terminal leaves represent the developmental endpoint of shoot extension, with their characteristics reflecting the conditions and timing of growth cessation including resource availability, environmental conditions, and developmental signals triggering extension termination, ranging from well-developed mature leaves if cessation occurs early under favorable conditions to tiny arrested leaves if cessation occurs abruptly late in season under stress or developmental programming.
Ultimate terminal leaves represent the developmental conclusion of shoot extension, distinguished from other terminal leaves by permanent rather than transient terminal status following growth arrest. Characteristics vary dramatically depending on cessation timing and conditions, with gradual cessation under favorable conditions producing well-developed ultimate terminal leaves approaching normal mature characteristics, while abrupt cessation under stress or late-season developmental programming producing small poorly developed leaves showing arrested development. The ultimate terminal leaf designation becomes fixed only after shoot extension ceases, with leaves occupying terminal position during active growth being transient terminals subject to displacement by subsequent leaf emergence. Timing of shoot extension cessation varies with cultivar, environmental conditions, crop load, and cultural practices including irrigation and fertilization management, with early cessation (July) producing well-developed ultimate terminals while late cessation (September) may produce minimal development. Ultimate terminal leaves on shoots retained as fruiting canes persist through winter and remain as basal leaves on lateral shoots emerging the following season, creating positional transformation from ultimate terminal to basal categories across the annual growth cycle.
endgültiges apikales Weinblatt
final terminal leaf
growth-arrest terminal leaf
last-initiated leaf
letztes terminales Blatt
wachstumsstop Endblatt
Based on Palliotti & Cartechini (2000) "Developmental changes in gas exchange activity in flowers, berries, and tendrils of field-grown Cabernet Sauvignon" American Journal of Enology and Viticulture 51=317-323 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 109-134 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 137-172.
Ultimate Terminal Leaf
Pinot Noir vines in the Clos Vougeot-adjacent parcels at Domaine Méo-Camuzet bearing an ultra-premium wine role for the Vosne-Romanée 1er Cru Cros Parantoux are managed at yields of 25-30 hl/ha through severe winter pruning to 6-8 buds per vine and green harvesting at véraison, with 100% whole-cluster fermentation, 18 months in 60-70% new Burgundy barriques, and production limited to 300-500 cases annually at 250-400 €/bottle, positioning the wine between the estate's village Vosne-Romanée (50-70 €) and Grand Cru Richebourg (500-800 €).
A quality tier role that inheres in a grapevine whose fruit is destined for wines of near-icon quality and prestige, produced in very limited quantities from top-performing vineyard parcels with highly restrictive yield targets, intensive vine-by-vine management, and extended aging programs, marketed at price points approaching icon status.
Ultra_Premium_Wine_Role occupies the narrow tier between super-premium and icon, typically at $50-100+ USD retail. This segment is the most variable in its boundaries across industry classification systems: some analysts merge it with super-premium, while others distinguish it as the "luxury" or "prestige" category. The tier characteristically includes top Premier Cru Burgundies, classified-growth Bordeaux, reserve bottlings from prestigious estates, and limited-production single-vineyard wines from elite producers. Viticultural management at this tier approaches icon-level intensity but may encompass slightly larger production volumes.
Luxuswein-Rolle
Prestige-Cuvée-Rolle
Ultra-Premium-Wein-Rolle
luxury wine role
prestige cuvée role
ultra-premium wine vine role
Rabobank (2019) Wine Quarterly Report: Global Wine Market Segmentation Liv-ex (2023) Fine Wine Market Classification McMillan R (2022) State of the US Wine Industry, Silicon Valley Bank pp: 18-22
Ultra Premium Wine Role
Umami quality investigation in Champagne comparing non-vintage (18 months sur lie) and prestige cuvée (7+ years sur lie) using a trained panel with umami reference standards reveals umami detection in 85% of panelists for prestige cuvée (glutamate 145 mg/L, nucleotides from extended yeast autolysis) versus 35% detection in non-vintage (glutamate 78 mg/L), suggesting that extended lees aging may elevate wine umami above perception threshold, potentially contributing to the enhanced complexity and gastronomy pairing versatility attributed to aged Champagne.
Umami quality is a basic taste quality that represents the gustatory perception of savory taste arising primarily from glutamate stimulation of T1R1/T1R3 heterodimeric taste receptors, with synergistic enhancement by ribonucleotides, characterizing the brothy, savory, mouth-filling sensation that may contribute subtle complexity to wine taste though its relevance in grape and wine context remains subject to ongoing investigation. Within the BFO framework, this quality addresses the fifth basic taste modality as applied to viticulture and enology, distinguishing umami from other taste qualities while acknowledging the relatively limited and debated role of umami in wine sensory character compared to its prominence in food evaluation. The ontological significance lies in umami potentially contributing to the savory, complex character of certain wines, particularly aged wines where protein hydrolysis releases free amino acids including glutamate, and sur lie aged wines where yeast autolysis contributes nucleotides that could synergize with glutamate for enhanced umami perception. The physiological basis for umami involves the T1R1/T1R3 receptor responding to L-glutamate and the synergistic enhancement (up to 15-fold) by 5'-ribonucleotides (IMP, GMP, AMP) that bind an adjacent site on the receptor, explaining why combinations of these compounds produce umami perception greater than the sum of individual contributions. The compositional basis for potential wine umami includes free glutamate (present at 20-150 mg/L in wines), other free amino acids with weaker umami character, and nucleotides from yeast autolysis in sur lie aged and sparkling wines. The threshold for glutamate detection in wine matrix is debated, with some studies suggesting wine glutamate concentrations are below sensory threshold while others report detectable umami contribution in certain wine styles. Understanding umami quality in wine context enables exploration of whether and how this taste modality contributes to wine sensory complexity, investigation of winemaking practices that might enhance umami character, and complete characterization of wine taste considering all basic taste modalities.
Umami_Quality addresses the fifth basic taste in wine context, where its contribution remains less established than sweetness, sourness, and bitterness. This quality completes the basic taste framework while acknowledging the ongoing scientific discussion about umami relevance in wine. Unlike other basic tastes with child classes for intensity, character, and persistence, umami in wine is currently treated as a single quality given the limited research on its dimensional attributes in this specific matrix.
Glutamatgeschmack
Umami
Wohlgeschmack
deliciousness
fifth taste
glutamate taste
herzhafter Geschmack
savory taste
Chaudhari N et al. (2000) A metabotropic glutamate receptor variant functions as a taste receptor. Nature Neuroscience 3(2): 113-119 Halpern BP (2000) Glutamate and the flavor of foods. Journal of Nutrition 130(4): 921S-926S Desportes C et al. (2001) Amino acid content and ammonium release during aging on lees in Champagne base wine. American Journal of Enology and Viticulture 52(4): 361-369
Umami Quality
An unbalanced bilateral cordon on Vitis vinifera cv. Cabernet Sauvignon at row-end position, featuring short cordon at 42cm extending toward row interior and long cordon at 98cm extending toward row end (56cm difference, 2.3=1 ratio), short cordon bearing 7 spurs while long cordon bears 16 spurs, crop distribution approximately 68% on long cordon versus 32% on short cordon (total 4.8kg), trunk positioned 32cm off-center toward row interior due to row-end spacing requirements, representing functional bilateral architecture accommodating row-end constraints while maintaining adequate productive capacity through extended cordon compensating for shortened cordon, with total spur count (23) and crop appropriate for vine vigor despite pronounced asymmetrical configuration.
A bilateral cordon configuration with substantial dimensional or capacity differences between the two cordons, characterized by unequal cordon lengths (typically differing by 20cm or more), imbalanced spur counts or spacing, skewed crop distribution departing significantly from 50=50 split (often 60=40 or more extreme), and architectural asymmetry resulting from off-center trunk positioning, directional vigor differences, establishment problems, or deliberate design accommodating site constraints or row-end positions. Unbalanced bilateral cordons represent practical accommodations to real-world vineyard constraints, accepting architectural asymmetry to maximize productivity within available space, adapt to vine vigor patterns, or work around positioning limitations, with functional performance often equivalent to balanced configurations despite aesthetic differences and potential management complications.
Unbalanced bilateral cordons are common and functionally acceptable in commercial vineyards, arising from various causes including off-center trunk positioning (planting irregularities, vine replacement creating gaps, deliberate row-end spacing), directional vigor differences (slope aspects creating different sun exposure, prevailing wind patterns, differential soil depth or fertility, equipment traffic compaction on one side), establishment challenges (damage to one developing cordon requiring retraining, poor initial growth on one side due to competition or stress), and deliberate design (row-end accommodations, curved rows, irregular spacing patterns). While aesthetically less satisfying than balanced configurations, unbalanced cordons often perform equivalently in terms of total productivity (combined spur count and crop from both cordons meeting targets), fruit quality (adequate leaf area-to-fruit ratios maintained through appropriate crop management), and practical management (experienced workers adapting to asymmetry without difficulty). Major asymmetries exceeding 2=1 ratios may indicate problems requiring investigation or correction.
asymmetrical bilateral cordon
asymmetrische Doppelkordon
skewed bilateral cordon
unausgeglichene bilaterale Kordon
unequal paired cordon
Unbalanced Bilateral Cordon
An unbranched tendril at node 8 on a shoot of Vitis vinifera cv. Sauvignon Blanc in Sancerre, France, measuring 16 cm total length, consisting of a single continuous axis of 2.8 mm diameter at mid-length without any branch points or lateral meristematic activity, exhibiting active circumnutation with 125-degree sweep amplitude exploring approximately 285 cubic centimeters of space, making successful contact with a vertical trellis wire and initiating tight coiling with six contact coils and eight free coils providing robust single-point attachment supporting the 60 cm distal shoot section bearing six leaves and one fruit cluster totaling 380 grams weight, occurring in a cultivar where 55-65% of tendrils show unbranched architecture while 35-45% develop bifurcated patterns depending on shoot vigor and resource availability, measured at BBCH stage 73 (berries beginning to touch) in early July, representing the simple unbranched pattern common in this cultivar under normal vigor conditions.
A tendril consisting of a single continuous axis without division into separate branches or arms in Vitis vinifera, characterized by absence of branch points or lateral meristematic activity producing secondary axes, development from a single apical meristem that maintains exclusive dominance throughout tendril organogenesis without activating lateral meristems, elongation producing a single filiform structure of 8-25 cm length depending on cultivar and environmental conditions, and representing either a simple mature architectural pattern expressing genetic programming for unbranched development or an immature developmental stage preceding potential branching that may or may not occur depending on cultivar genetics and resource availability. Unbranched tendrils occur with variable frequency across Vitis vinifera cultivars, ranging from very rare (70% of tendrils) in cultivars showing genetic predisposition for unbranched architecture, with intermediate frequencies (20-50%) common in many cultivars showing mixed branching patterns influenced by environmental conditions during development. The functional implications of unbranched architecture include reduced spatial exploration volume compared to branched tendrils, typically 180-350 cubic centimeters depending on tendril length and circumnutation amplitude, single potential attachment point rather than multiple redundant opportunities, lower resource investment requirements enabling production under moderate vigor conditions where branching might be unsustainable, and potentially enhanced individual tendril strength through concentration of vascular and structural resources in a single axis rather than distribution across multiple branches.
Unbranched tendrils are distinguished from branched tendrils by absence of any division points or separate arms, consisting of a single continuous axis from shoot attachment to apex. Classification requires assessment of developmental completion, as some unbranched tendrils represent immature juveniles that would develop branches with continued growth while others represent mature simple architecture. Unbranched frequency shows strong cultivar specificity serving as an important ampelographic diagnostic character, with cultivars ranging from nearly always branched (Mourvèdre, Syrah) to predominantly unbranched (some Riesling clones, Sauvignon Blanc) to highly variable (Pinot Noir, Chardonnay). Environmental conditions influence branching expression, with high vigor and optimal resources promoting increased branching frequency while moderate vigor or stress conditions favor unbranched development even in cultivars with branching tendencies. Functionally, unbranched tendrils provide adequate climbing support for moderate shoot weights through single robust attachments, particularly effective in well-designed trellis systems where support wires are positioned for easy contact by searching tendrils.
einfache Ranke
nicht-geteilte Kletterorgan
non-branched tendril
simple tendril
single-axis tendril
undivided tendril
unverzweigte Ranke
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 58-89 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 79-118.
Unbranched Tendril
An underground origin basal shoot on grafted Vitis vinifera cv. Cabernet Sauvignon (on SO4 rootstock) emerging through soil surface 12cm from trunk, vigorous shoot reaching 215cm length by late July, leaves showing different morphology than Cabernet (larger, less deeply lobed, suggesting rootstock characteristics), excavation to 15cm depth revealing shoot origin from rootstock tissue 8cm below graft union confirming rootstock sucker identity (not scion renewal shoot), extremely vigorous growth reflecting robust SO4 rootstock characteristics, this shoot representing problematic rootstock suckering requiring immediate removal preventing rootstock overgrowth that could suppress or displace scion, with underground rootstock origin creating ongoing management concern as removed sucker likely to resprout requiring multiple removal cycles or herbicide spot treatment preventing recurrence, this underground sucker exemplifying common grafted vine management challenge requiring vigilant monitoring and prompt intervention preventing rootstock establishment.
A ground-level basal shoot originating from below-ground positions, characterized by emergence from underground trunk portions or root structures at depths exceeding 2-5cm, requiring excavation for origin determination and shoot base access, high probability of rootstock origin in grafted vines (with graft unions typically 5-15cm above soil meaning underground shoots often emerging below graft from rootstock tissues), and function as either unwanted rootstock suckers requiring prompt removal or in own-rooted vines potentially viable renewal shoots though with complications from underground origin and surface emergence challenges. Underground origin basal shoots represent deeply-sourced basal growth, creating significant classification challenges (scion versus rootstock determination requiring excavation and careful examination), management complications (underground shoots often highly vigorous from rootstock characteristics if grafted vines), and in grafted systems typically representing problematic rootstock suckering requiring removal preventing rootstock competition with scion and maintaining desired varietal characteristics.
Underground origin basal shoots require immediate careful assessment in grafted vines, with high probability of rootstock origin (shoots emerging from below graft union) necessitating prompt removal preventing serious complications including rootstock competition (vigorous rootstock shoots consuming resources and suppressing scion growth), varietal displacement (rootstock potentially overgrowing and replacing scion if unchecked), and management problems (rootstock suckers showing strong resprouting tendency requiring repeated removal). Origin verification requires excavation exposing shoot connection point, examining emergence relative to graft union position, and comparing shoot morphology with known scion and rootstock characteristics (leaf shape, growth habit, vigor patterns). In own-rooted vines, underground origin shoots represent scion material but still create challenges including underground emergence complications, potential root disturbance during excavation or training, and questions about shoot quality and vigor from deeply-sourced origins. Management in grafted vines typically involves immediate removal of confirmed rootstock shoots, vigilant monitoring for resprouting, possible herbicide spot treatment for persistent suckering, with own-rooted situations requiring case-by-case assessment balancing potential renewal value against underground origin complications.
Wurzelausschlag
below-ground basal shoot
root shoot
subsurface origin shoot
underground sucker
unterirdischer Ursprungstrieb
Based on Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press, pp. 85-105 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 113-145 Pongrácz (1983) "Rootstocks for Grape-vines" David Philip Publisher, pp. 112-134.
Underground Origin Basal Shoot
An unequal trifurcated tendril at node 9 on a moderately vigorous shoot of Vitis vinifera cv. Grenache in Châteauneuf-du-Pape, France, exhibiting pseudo-trifurcation through bifurcation at 7 cm from shoot attachment followed by secondary division of one arm at 10 cm, creating a dominant primary arm of 12.5 cm, a co-dominant secondary arm of 10.5 cm, and a subordinate tertiary arm of 6.5 cm (ratios 0.84=1 and 0.52=1), with branch angles of 40, 45, and 95 degrees creating asymmetric rather than radial architecture, dominant arm showing 3.4 mm diameter while subordinate arm measures 2.1 mm, dominant and co-dominant arms exhibiting vigorous circumnutation (145-160 degrees) while subordinate arm shows reduced amplitude (85 degrees), occurring on a shoot experiencing moderate resource limitation from 2.2 kg/m crop load, with the dominant arm making successful strong attachment while co-dominant and subordinate arms provide supplementary exploration, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing typical unequal trifurcation pattern under normal commercial vineyard conditions.
A trifurcated tendril exhibiting imbalanced development among the three branches in Vitis vinifera, characterized by at least one pairwise arm length ratio less than 0.85=1 indicating differential resource allocation or developmental timing creating dominance hierarchies among the three arms, often accompanied by unequal branch angles deviating from 120-degree radial symmetry, differential branch diameter reflecting asymmetric vascular development, and divergent functional capabilities with one or two dominant arms showing stronger circumnutation and coiling responses compared to one or two subordinate arms. Unequal trifurcated tendrils develop through sequential meristem activation creating developmental advantages for earlier-initiated arms, differential resource allocation during growth with preferential flows to certain branches based on vascular architecture or position, unequal environmental exposure affecting arms differently, damage or stress impacting some arms while sparing others, or fundamental developmental regulation patterns in some cultivars that establish arm hierarchies even under optimal conditions. This branching pattern represents the most common expression of trifurcated architecture, accounting for 65-80% of three-armed tendrils in most cultivars, and reflects the challenge of supporting balanced three-way resource partitioning which typically requires exceptional vigor and resource availability rarely achieved under commercial vineyard conditions. Functional implications include one or two arms achieving near-normal development providing reliable climbing capacity while remaining arm(s) contribute supplementary but reduced exploration and attachment potential, creating intermediate performance between bifurcated symmetry and full equal trifurcation.
Unequal trifurcated tendrils are distinguished from equal trifurcated tendrils by at least one pairwise arm length ratio <0.85=1, asymmetric branch angles, differential structural development, and divergent functional capabilities among arms. The degree and pattern of inequality provide diagnostic information, with mild inequality (smallest ratio 0.70-0.84=1) indicating moderate resource limitation while severe inequality (smallest ratio <0.60=1) suggests substantial developmental problems or damage. Patterns of inequality vary, including one dominant with two equal subordinates (1+2 pattern), two co-dominants with one subordinate (2+1 pattern), or complete hierarchy with three unequal arms. True three-way branching versus pseudo-trifurcation through sequential bifurcations can be distinguished by branch point examination, with implications for developmental interpretation. Functionally, unequal trifurcation maintains advantages over bifurcation through increased total arm number despite inequality, particularly when two arms achieve strong development creating dual functional capacity with third arm backup.
dominant-subordinate three-branch tendril
dominant-untergeordnete dreiarmige Kletterorgan
hierarchical tri-arm tendril
hierarchische dreifach-verzweigte Ranke
imbalanced trifurcated tendril
ungleichmäßig dreigabelige Ranke
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 151-165 Petrie et al. (2000) "The effect of leaf removal and canopy height on whole-vine gas exchange and fruit development of Vitis vinifera L. Sauvignon blanc" Functional Plant Biology 27=173-184 Gerrath et al. (2015) "Developmental morphology and anatomy of the stem and tendrils of Vitis riparia (Vitaceae)" Canadian Journal of Botany 93=389-397 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 195-218.
Unequal Trifurcated Tendril
An unfolding young leaf at node 13 on an actively extending shoot of Vitis vinifera cv. Cabernet Sauvignon in Napa Valley, California, measuring 2.8 cm length by 2.4 cm width representing approximately 17% of expected mature dimensions, positioned 8 cm behind shoot apex with apex and two younger primordia still ahead, exhibiting initial unfoldment revealing five-lobed palmate outline with lobes still incompletely separated and sinuses shallow, pale green coloration with pronounced bronze-purple tinge from anthocyanin concentration of 3.8 mg g⁻¹ fresh weight particularly evident along primary veins, soft fragile texture with cells transitioning from division to expansion phase with average palisade cell size increasing from 15 μm to 22 μm during this stage, major veins becoming prominent as green ridges on abaxial surface with procambium differentiating into immature xylem and phloem, stomatal primordia visible as paired cells on abaxial surface but without functional pores, thin developing cuticle creating slight sheen on adaxial surface, minimal photosynthetic activity measured at 1.8 μmol CO₂ m⁻² s⁻¹ representing 14% of mature leaf capacity and contributing approximately 3 mg carbohydrate per day while consuming 15 mg for continued growth representing net sink of 12 mg per day, occurring at 9 days post-primordium initiation, measured at shoot BBCH stage 55 (inflorescence swelling) in early May, representing transition from compressed primordium to recognizable unfolding leaf structure.
An emerging young leaf in the transition phase from compressed primordium to initial lamina expansion in Vitis vinifera, characterized by small size typically 1.5-3.5 cm length representing 10-20% of final mature dimensions, position just beyond protective bud scales or older leaf primordia beginning to be exposed to ambient environment, initiating unfoldment of previously tightly folded lamina revealing basic lobe outline and leaf shape, pale green to bronze-green coloration with high anthocyanin concentrations particularly along veins and margins providing photoprotection, transition from pure meristematic tissue to beginning cell expansion with cells enlarging through water uptake and vacuole development, initial tissue differentiation with procambial strands developing into recognizable immature veins and mesophyll beginning to show incipient layering, appearance of stomatal primordia on abaxial surface though not yet functional, beginning cuticle deposition creating slight surface sheen compared to completely dull primordial surfaces, and minimal but detectable photosynthetic activity typically 5-15% of mature leaf rates beginning to offset but not eliminate net resource sink status. Unfolding young leaves represent the critical transition from pure sink to beginning source function, with cell division rates declining as cell expansion becomes the dominant growth process, lamina beginning to flatten from previously rolled or folded configuration revealing the palmate architecture characteristic of grapevine with major lobe positions becoming evident, and tissue beginning differentiation processes that will determine final anatomical characteristics including palisade and spongy mesophyll layers, with the unfolding phase typically spanning 4-7 days under optimal temperature conditions representing the period of maximum vulnerability to mechanical damage from wind, hail, or handling due to extremely fragile partially expanded tissues.
Unfolding young leaves are distinguished from primordium stage by beginning lamina expansion and basic shape revelation, and from expanding stage by very small size and incomplete unfoldment with major folding or rolling still evident. This developmental phase represents critical architectural establishment as lobe patterns become visible, vein arrangements are laid down, and basic tissue differentiation begins. Environmental conditions during unfolding critically affect final leaf characteristics, with light environment during this phase particularly important for determining sun versus shade leaf anatomy through differential cell division in future palisade zones. Unfolding leaves show extreme fragility and vulnerability, with tissue damage from mechanical forces, pest attack, or disease infection commonly creating permanent deformations including torn lobes, asymmetric development, or scarring that persists through leaf life. Frost damage during unfolding typically causes complete tissue death as expanding cells with large vacuoles are highly susceptible to ice crystal formation. Management practices including careful shoot positioning, timely pest control targeting mites and leafhoppers, and frost protection are critical during this vulnerable phase.
anfangs-expansions Blatt
beginning expansion leaf
early lamina development leaf
frühe Lamina-entwicklung Weinblatt
initial unfolding leaf
sich entfaltendes junges Blatt
Based on Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 60-85 Palliotti & Cartechini (2000) "Developmental changes in gas exchange activity in flowers, berries, and tendrils of field-grown Cabernet Sauvignon" American Journal of Enology and Viticulture 51=317-323 Tsukaya (2013) "Leaf development" The Arabidopsis Book 11=e0163 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 79-102.
Unfolding Young Leaf
In a Sonoma Coast Pinot Noir vineyard, uniform color development was achieved 10 days post-color break in mid-September Year N+1 (BBCH 85) when >96% berry surface showed consistent color, anthocyanin spatial variation (CV) was 14%, CIE L* variation 9%, skin anthocyanins averaged 1285 mg/kg (SD ±165 mg/kg), berry sugar 22.4° Brix, titratable acidity 6.2 g/L, firmness 1.3 N, under optimal cool coastal conditions (daily high 21-24°C, night 10-13°C, morning fog moderating cluster temperature) with good canopy light distribution from summer leaf removal.
Uniform_Color_Development represents the late veraison phase when berry surface pigmentation achieves spatial consistency with minimal color variation across the entire berry creating homogeneous appearance suitable for harvest quality assessment and indicating ripening synchrony critical for optimal wine production. This process occurs during late veraison to early ripening (BBCH 85-87), typically 8-14 days post-color break depending on cultivar and environmental conditions, and is characterized by >95% berry surface showing consistent color with spatial variation coefficients declining to 60-80% variation during color break), regulatory transcription factors VvMYBA1/A2 showing consistent activity throughout berry epidermis driven by uniform ABA distribution (1800-2800 ng/g with 25-35% variation during progressive development. Optical characteristics of uniform development inclu reflectance spectra measured at multiple berry positions showing consistent profiles with characteristic anthocyanin absorbance maximum 520-540 nm and standard deviation 32°C impairs uniformity creating persistent color variation through spatial differences in heat tolerance and metabolic capacity, and stable environmental conditions without dramatic fluctuations favor uniformity while variable conditions (temperature swings, water stress cycling) can disrupt synchronized development maintaining heterogeneity. Light distribution critically affects uniformity: uniform cluster light exposure through optimal canopy architecture produces uniform berry pigmentation with 95% uniform coverage in 7-9 days post-color break with very consistent appearance, moderate cultivars like Merlot and Cabernet Sauvignon require 9-12 days for uniformity showing good but not perfect consistency, while slow or variable-uniforming varieties like Nebbiolo and Sangiovese may need 12-16 days and still exhibit 20-30% residual variation reflecting genetic differences in developmental synchrony and anthocyanin synthesis coordination. Within-vineyard spatial variation in uniform development reveals: terroir effects from soil, microclimate, or vine age differences creating 4-8 day variation in uniformity achievement across vineyard blocks, vine vigor effects with high-vigor vines showing delayed and less complete uniformity through extended vegetative-reproductive competition and excessive canopy density, and rootstock influences with vigorous rootstocks potentially delaying uniformity 2-4 days compared to low-vigor stocks through altered carbon allocation and water relations. Quality implications of uniform color development inclu uniformity indicates ripening synchrony suggesting berries achieving consistent color also show coordinated sugar, acid, and phenolic maturation enabling optimal harvest timing, lack of uniformity reveals developmental heterogeneity potentially requiring selective harvest of uniform clusters or acceptance of compositional variation in wine, uniform color strongly correlates with wine quality consistency (r² = 0.75-0.88) since spatial variation in color reflects broader maturity differences affecting multiple quality parameters, and visual assessment of uniformity provides practical harvest readiness evaluation more accessible than chemical analysis enabling field-based decisions. Monitoring uniform development involves: photographic documentation with image analysis measuring color uniformity coefficients across berry surfaces and clusters, portable colorimeters measuring CIE L*a*b* at multiple berry positions (typically 4-6 points around equator) revealing spatial variation quantitatively, visual scoring using standardized charts rating clusters 1-5 for uniformity (1 = highly variable, 5 = completely uniform) providing rapid semi-quantitative assessment, and selective sampling of berries from different cluster positions and vineyard zones for anthocyanin analysis revealing spatial variation guiding harvest timing and quality prediction. Temporal coordination with other maturity parameters shows: uniform color development typically achieved when berry sugar reaches 20-26° Brix (cultivar and style dependent), coincides with 70-85% of total berry softening completed producing characteristic ripe firmness 1.2-2.2 N, occurs during late acid decline with titratable acidity 4-7 g/L and pH 3.2-3.6 approaching harvest targets, and parallels phenolic maturity with seed browning and extractable tannin optimization indicating integrated berry development approaching optimal harvest window. Physiological significance includes: uniform pigmentation across berry surface maximizes UV protection and antioxidant capacity distributed throughout skin tissue, consistent anthocyanin levels enable predictable extraction during winemaking producing consistent wine color and phenolic profiles, and uniformity reflects overall metabolic coordination suggesting synchronized synthesis of other quality compounds including flavors, aromas, and structural phenolics contributing to wine complexity. Viticultural management targeting uniform development includes: canopy management creating optimal light distribution through shoot positioning, leaf removal, and cluster exposure balancing synthesis induction with heat stress avoidance, irrigation scheduling maintaining consistent moderate deficit throughout veraison avoiding water stress fluctuations that disrupt synchronized development, crop load adjustment optimizing source-sink balance ensuring adequate carbohydrate supply for all berries achieving coordinated maturation, disease control preventing localized infections that create developmental heterogeneity through pathogen-induced metabolic disruption, and soil management addressing spatial variability in water and nutrient availability that creates non-uniform vine responses affecting berry development consistency. Harvest timing optimization based on uniform development: achieving >95% uniform color indicates ripening completion suitable for harvest quality assessment, persistent non-uniformity (>20-25% variation) may necessitate selective harvest removing uniform clusters while leaving variable fruit for later picking, or acceptance of compositional variation recognizing wine quality implications, and integration of color uniformity with chemical maturity (sugar, acid, pH) and sensory evaluation (flavor, tannin) enables comprehensive harvest readiness determination balancing multiple quality parameters for optimal wine production.
Uniform_Color_Development is distinguished from Progressive_Color_Development by representing spatial homogenization achievement rather than temporal progression, from Uniform_Pigmentation_Progress by showing completed rather than advancing uniformity, and from Complete_Color_Change by focusing on spatial consistency rather than maximum intensity. Uniformity criteria: >95% berry surface, anthocyanin CV <15-20%, CIE L* CV <10-15%, hue variation <8-12°. Timing: fast cultivars 7-9 days post-break, moderate 9-12 days, slow 12-16 days. Optimal temperature 22-28°C. Uniform light exposure critical. Uniform_Color_Development indicates ripening synchrony essential for harvest timing and wine quality consistency.
consistent color expression
homogene Beeren-Pigmentierung
homogeneous berry pigmentation
konsistente Farb-Expression
räumlich uniforme Färbung
spatially uniform coloring
synchronisierte Farb-Reifung
synchronized color maturation
Keller M (2020) The Science of Grapevines 3rd Ed pages 234-238 color development uniformity and maturity Tarara JM, Lee J, Spayd SE, Scagel CF (2008) Berry temperature and solar radiation alter acylation, proportion, and concentration of anthocyanin in Merlot grapes Amer J Enol Vitic 59:235-247 environmental effects on color uniformity Cortell JM, Kennedy JA (2006) Effect of shading on accumulation of flavonoid compounds in Pinot Noir fruit and extraction in wine Amer J Enol Vitic 57:33-43 canopy management effects on uniformity.
Uniform Color Development
Uniform coloration achievement in Crimson Seedless requires intensive canopy management including multiple leaf removal passes that maintain 30-40% cluster exposure to direct sunlight throughout the ripening period, with uniformly red berries (color variation coefficient below 8%) commanding a 40-60% price premium over fruit with non-uniform coloration in premium export markets serving Asia and Europe where visual perfection drives consumer purchasing decisions and any green shoulders or pale zones relegate fruit to lower value domestic markets.
Uniform_Coloration_Quality represents the color distribution condition where pigmentation appears homogeneous across the surface of a grapevine organ without visually distinct zones, patches, or gradients of contrasting color or intensity. Uniform coloration indicates consistent pigment accumulation throughout the tissue, reflecting either uniform environmental conditions during pigment development or developmental programs that produce spatially homogeneous pigmentation regardless of microenvironmental variation. In grape berries, uniform coloration is the quality standard for table grapes and an indicator of optimal ripening conditions for wine grapes. Achieving uniform berry coloration requires consistent light exposure across the berry surface, which is promoted by open canopy architecture, cluster positioning in the light zone, and loose cluster structure that allows light penetration between berries. Cultivars vary genetically in their tendency toward uniform coloration, with some varieties (particularly red table grapes selected for visual appeal) displaying more consistent coloration than others. Uniform coloration at the cluster level indicates synchronous berry development, a desirable character for harvest timing and fruit processing. In leaves, uniform green coloration indicates healthy photosynthetic function without nutrient deficiencies, disease symptoms, or localized damage that would create color variation. Assessment of uniformity employs visual evaluation against standards, calculation of color variation coefficients across sampled points, or image analysis quantifying the proportion of surface area falling within a defined color range. The threshold distinguishing uniform from non-uniform coloration involves judgment and may be specified quantitatively (e.g., coefficient of variation below 10% for measured color parameters).
Uniform_Coloration_Quality represents the ideal of homogeneous color distribution and serves as the reference condition against which Variegated_Coloration_Quality and Gradient_Coloration_Quality are defined. Perfect uniformity is rarely achieved in biological organs the classification applies when color variation falls below thresholds that would indicate distinct zones or gradients. Uniformity may be assessed at different scales (within-berry, within-cluster, within-vine), and organs may display uniformity at one scale while showing variation at another. For table grape quality standards, specific color coverage percentages (e.g., greater than 95% red coverage) operationally define uniform coloration.
consistent pigmentation
einheitliche Pigmentierung
even color
gleichförmige Färbung
gleichmäßige Färbung
homogene Farbverteilung
homogeneous coloration
uniform color distribution
Peppi MC et al. (2007) Application of Abscisic Acid Rapidly Upregulated UFGT Gene Expression and Improved Color of Grape Berries. Vitis 46(3): 141-146 Kuhn N et al. (2014) Berry Ripening: Recently Heard Through the Grapevine. Journal of Experimental Botany 65(16): 4543-4559
Uniform Coloration Quality
In a Rheingau Spätburgunder (Pinot Noir) vineyard, uniform pigmentation progress was evaluated over 6 days in late September Year N+1 (BBCH 85) when berry surface color coverage increased from 65% to 96% uniformly pigmented, spatial anthocyanin variation (CV) declined from 42% to 18%, CIE L* variation decreased from 32% to 11%, skin anthocyanins reached 1150 mg/kg average with <200 mg/kg standard deviation across berry positions, berry sugar was 21.8° Brix, under cool autumn conditions (daily high 19-22°C, night 9-12°C) with good cluster exposure from summer leaf removal.
Uniform_Pigmentation_Progress represents the spatial homogenization of berry surface color during late progressive development when initially scattered and patchy pigmentation coalesces into continuous uniform coloring covering the entire berry surface with consistent intensity and hue eliminating green remnants and color discontinuities. This process occurs during late veraison to early ripening (BBCH 84-85), typically 6-12 days post-color break depending on cultivar and environmental conditions, and transforms berry appearance from variegated patches of intense and pale color to uniform coverage where >90-95% of berry surface shows consistent pigmentation suitable for harvest assessment and quality evaluation. Spatial patterns of uniformity achievement show: initially pigmented sun-exposed surfaces (occupying 50-70% berry area at mid-progressive development) expand into shaded regions through continued anthocyanin synthesis in previously unpigmented cells, green remnants concentrated on northwestern aspects (northern hemisphere) and near pedicel attachment progressively shrink and disappear as chlorophyll degradation completes and anthocyanin accumulation reaches these developmental lag zones, and within-berry color gradients diminish from 3-5 fold variation between most and least pigmented regions at color break to 32°C impairs uniformity producing patchy coloring with persistent green or pale zones reflecting spatial variation in heat exposure and metabolic capacity under stress, and large diurnal temperature variation >15°C may enhance uniformity through cool nights enabling pigmentation in regions that experience excessive daytime heat. Light distribution effects on uniformity are pronounced: canopy architecture creating uniform cluster light exposure produces uniform berry pigmentation in 7-10 days, while dense canopies with strong light gradients generate 20-40% variation in final pigmentation requiring 12-16 days for partial uniformity achievement or never achieving complete uniformity, leaf removal or cluster thinning during progressive development can accelerate uniformity 2-4 days through improved light penetration to previously shaded berry surfaces, and cluster positioning with rotating or revolving clusters would theoretically optimize uniformity though impractical in commercial viticulture. Water relations effects inclu moderate deficit uniformly applied promotes consistent uniformity across all berries through systemic ABA elevation and uniform metabolic stress responses, spatially variable water availability from non-uniform irrigation or heterogeneous soil creates variable berry development and pigmentation uniformity challenges, and sudden water availability changes during progressive development can disrupt uniformity creating zones of different color intensity reflecting timing of developmental signal reception. Cultivar variation in uniformity progression shows: rapid-uniforming varieties like Pinot Noir achieve >90% uniform coverage in 5-7 days post-color break, moderate cultivars like Merlot and Cabernet Sauvignon require 7-10 days for uniformity, while slow-uniforming varieties like Nebbiolo may need 10-14 days or show persistent spatial variation even at harvest reflecting genetic differences in developmental synchrony and anthocyanin synthesis distribution. Within-cluster uniformity achievement shows: berries at different developmental stages creating cluster heterogeneity with leading berries achieving uniformity 3-5 days before lagging berries, cluster shoulders often reaching uniformity before tips possibly reflecting vascular supply differences, and compact clusters with berry-berry contact potentially showing delayed uniformity in contact zones through restricted light and gas exchange affecting local metabolism. Quality implications of uniform pigmentation inclu uniform berry color indicates ripening synchrony enabling optimal harvest timing when all fruit achieves target maturity, lack of uniformity reveals developmental heterogeneity potentially requiring selective harvest or acceptance of variable wine composition, color uniformity often correlates with other maturity parameters suggesting berries achieving uniform pigmentation also show consistent sugar, acid, and phenolic development, and visual assessment of cluster uniformity provides practical harvest readiness evaluation tool for viticulturists. Monitoring uniformity progress involves: photographic documentation enabling image analysis measuring percentage of berry surface showing target color intensity, visual scoring using standardized color charts rating clusters on 1-5 or 1-10 scales for uniformity, and sampling berries from different cluster positions for anthocyanin analysis revealing spatial variation quantitatively guiding harvest timing and quality predictions. Temporal coordination with other ripening processes: uniform pigmentation typically achieved when berry sugar reaches 18-24° Brix depending on cultivar and style, coincides with 60-80% of total berry softening completed, occurs during late malate decline producing desired acid balance, and parallels seed maturation (brown color, lignified testa) indicating integrated berry development approaching harvest readiness. Viticultural management targeting uniformity includes: canopy management to improve cluster light exposure accelerates and enhances uniformity, irrigation scheduling to maintain moderate consistent deficit promotes uniform developmental signaling across all berries, crop load adjustment to optimize source-sink balance enables adequate resource supply for all berries achieving synchronized maturation, and disease control particularly Botrytis prevention since immature green berry remnants show enhanced disease susceptibility potentially serving as infection foci compromising cluster quality.
Uniform_Pigmentation_Progress is distinguished from Progressive_Color_Development by representing spatial homogenization rather than temporal progression, from Color_Intensity_Increase by focusing on uniformity rather than saturation, and from Complete_Color_Change by showing advancing uniformity rather than final state. Uniformity threshold: >90-95% berry surface consistently pigmented. Timing: fast cultivars 5-7 days post-break, moderate 7-10 days, slow 10-14 days. Temperature optimum 22-28°C. Uniform light exposure critical. Uniform_Pigmentation_Progress indicates ripening synchrony essential for optimal harvest timing and wine quality consistency.
consistent pigmentation distribution
even berry coloring
gleichmäßige Beeren-Färbung
homogene Farb-Abdeckung
homogeneous color coverage
konsistente Pigmentierungs-Verteilung
räumliche Farb-Uniformität
spatial color uniformity
Keller M (2020) The Science of Grapevines 3rd Ed pages 234-236 color development uniformity Downey MO, Harvey JS, Robinson SP (2003) Analysis of tannins in seeds and skins of Shiraz grapes throughout berry development Australian J Grape Wine Res 9:15-27 spatial patterns of berry development Cortell JM, Kennedy JA (2006) Effect of shading on accumulation of flavonoid compounds in Pinot Noir fruit and extraction in wine Amer J Enol Vitic 57:33-43 light effects on color uniformity.
Uniform Pigmentation Progress
Cabernet Sauvignon exhibits uniform veraison disposition with 80% of berries completing color change within a 7-10 day window under favorable conditions, compared to 14-21 days for cultivars with extended veraison. Merlot shows high veraison uniformity with within-cluster coefficient of variation for sugar concentration at harvest of 8-12% compared to 15-25% in less uniform cultivars. Uniform veraison in Riesling enables single-pass harvest with consistent °Brix readings (±1.0°) across the vineyard block. Factors promoting uniformity include moderate crop load (Ravaz index 5-8), balanced canopy (1.0-1.5 m² leaf area per kg fruit), and even water status throughout the fruit zone.
A veraison timing disposition that is the inherited capacity of a Vitis vinifera plant to undergo the ripening transition synchronously across berries within clusters and across clusters within vines, with minimal temporal dispersion between first and last berries reaching the color change and softening stage.
Uniform veraison disposition reflects both genetic predisposition and environmental/management expression. Uniform ripening simplifies harvest timing decisions, improves wine consistency, and enables mechanical harvest without excessive heterogeneity in fruit maturity. The trait correlates with loose cluster architecture (r = 0.55-0.65) through improved light penetration and reduced within-cluster microclimate variation. Clone selection within cultivars can identify more uniform variants, with commercial Chardonnay clones showing veraison duration ranges from 8 to 18 days.
einheitlicher Farbumschlag
even ripening onset
gleichmäßige Véraison
synchroner Reifebeginn
synchronous veraison
uniform color change
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 265-285 Pagay V, Cheng L (2010) Variability in berry maturation of Concord and Cabernet franc in a cool climate. American Journal of Enology and Viticulture 61:61-67
Uniform Veraison Disposition
A unilateral cordon on Vitis vinifera cv. Riesling at vineyard row end adjacent to access road, extending 94cm in single direction from trunk toward row interior (opposite direction blocked by road boundary), bearing 15 two-bud spurs distributed along length at 12cm spacing, 26mm diameter indicating mature multi-year development, supporting estimated 28-30 fruiting shoots and 5.2-5.8kg crop concentrated on single cordon, representing deliberate permanent unilateral architecture necessitated by row-end position where no space exists for cordon extension toward road side, with vine vigor and total capacity equivalent to interior bilateral vines but channeled into single structural element requiring careful spur renewal and positioning to maintain productivity and prevent progressive cordon lengthening.
A cordon configuration featuring a single cordon extending in one direction from the trunk, characterized by absence of paired opposite cordon creating one-sided architecture, fruiting positions concentrated along single cordon length rather than distributed bilaterally, suitability for row-end positions where bilateral extension is impossible, narrow spacing situations, or transitional training during bilateral development. Unilateral cordons represent specialized or transitional configurations, deliberately employed at vineyard boundaries where space constraints prevent bilateral development, temporarily used during bilateral establishment (developing one cordon first, adding second when vigor permits), or maintained as permanent single-cordon architecture in specific training systems, narrow-spacing high-density plantings, or traditional approaches in limited regions, with vine capacity concentrated into single structural element rather than distributed across paired cordons.
Unilateral cordons serve essential functions despite being less common than bilateral configurations, with row-end positions universally requiring unilateral architecture (or strongly asymmetrical bilateral with minimal end-side cordon) making unilateral cordons necessary for complete vineyard utilization. Some traditional training systems deliberately employ unilateral architecture throughout vineyards rather than just at ends, reflecting historical practice or regional preferences. Transitional unilateral configurations are common during bilateral establishment in young vineyards, with initial single-cordon development (years 2-3) followed by second cordon addition when vigor permits (years 3-5), reducing early crop stress and supporting better establishment compared to simultaneous bilateral development. Management challenges of permanent unilateral cordons include maintaining cordon vigor and spur quality without bilateral balance, preventing excessive lengthening through careful spur renewal positioning, and ensuring adequate productivity from single structure through appropriate spur density and crop load management.
Einzelkordon
einseitige Kordon
mono-cordon
one-arm cordon
single-sided cordon
unilaterale Kordon
Unilateral Cordon
A Pinot Noir vine on unilateral cordon in Burgundy-inspired Oregon vineyards bears a Unilateral_Cordon_Role with a single 80 cm cordon extending toward the next vine, carrying 5 spurs that produce the focused crop load suited to premium Pinot production. A Riesling vine trained to unilateral cordon in steep German vineyards serves in a Unilateral_Cordon_Role, the single-arm structure adapted to narrow terrace widths where bilateral extension would exceed available space. A Chardonnay vine on unilateral cordon in Champagne-method sparkling wine production bears a Unilateral_Cordon_Role, the simpler structure facilitating the hand labor still prevalent in quality sparkling wine viticulture.
A cordon-trained vine role that inheres in a grapevine with a single cordon extending in one direction from the trunk, creating asymmetrical horizontal structure suited to specific training objectives, site constraints, or regional traditions. Unilateral cordon concentrates fruiting wood on one side of the trunk, which may simplify training, reduce cordon establishment time, or address situations where bilateral extension is impractical (row ends, steep slopes, narrow spacing). The single cordon typically extends the full distance to the next vine (1.0-2.0 m depending on vine spacing), creating a continuous fruiting zone along the row when all vines orient cordons in the same direction. Unilateral cordon may position fewer spurs per vine than bilateral systems (4-8 versus 8-15 typical), potentially limiting yield but simplifying management and concentrating resources into fewer fruiting positions. The asymmetrical structure requires attention to cordon direction (alternating between adjacent rows or uniform throughout) to maintain balanced vineyard appearance and facilitate mechanical operations.
Unilateral_Cordon_Role captures the single-cordon configuration chosen for simplicity, site adaptation, or production philosophy emphasizing concentrated fruiting. Unilateral cordon reduces establishment complexity (one cordon to train rather than two), may decrease trunk disease risk (less permanent wood), and concentrates crop in a focused zone that some producers believe enhances fruit quality. The system is associated with higher-density planting approaches where increased vine numbers compensate for reduced per-vine crop from the single cordon, potentially increasing production costs but also site utilization intensity. Burgundy's guyot system, while technically cane-pruned rather than true cordon, shares the unilateral fruiting philosophy with a single fruiting unit per vine. Contemporary unilateral cordon adoption often reflects Burgundian or Champagne influence, with New World regions adopting the approach for premium Pinot Noir and Chardonnay production.
Einseitiger Kordon
Halbkordon
einarmiger Kordon
single cordon
single-arm training
unilateral cordon
Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Reynolds AG, Vanden Heuvel JE (2009) Training systems, American Journal of Enology and Viticulture 60: 251-268 Jackson RS (2014) Wine Science: Principles and Applications, Academic Press
Unilateral Cordon Role
A unilateral permanent arm on Vitis vinifera cv. Riesling at vineyard row end, extending 92cm in single direction from trunk toward row interior (direction away from row end blocked by property boundary), bearing 15 two-bud spurs distributed along length, 24mm diameter indicating mature multi-year development, supporting estimated 28-30 fruiting shoots and 4.8-5.2kg crop concentrated on single arm, representing deliberate permanent unilateral architecture necessary for row-end position where no space exists for arm extension in opposite direction, with vine vigor and capacity equivalent to interior vines with bilateral arms but concentrated into single structural element.
A permanent arm configuration featuring a single arm extending in one direction from the trunk, characterized by absence of paired opposite arm creating one-sided architecture, fruiting positions concentrated along single arm length rather than distributed bilaterally, suitability for row-end positions, narrow spacing situations, or transitional training during bilateral development, and management approach concentrating vine capacity into single structural element rather than splitting between paired arms. Unilateral permanent arms represent specialized or transitional training configurations, deliberately employed at row ends where bilateral extension is impossible, temporarily used during bilateral cordon establishment (developing one arm first, adding second later when vigor permits), or maintained as permanent single-arm architecture in specific training systems (some traditional European approaches) or narrow-spacing high-density plantings where bilateral arms would create excessive crowding.
Unilateral permanent arms serve important niche functions despite being less common than bilateral configurations in modern viticulture. Row-end positions universally require unilateral architecture (or strongly asymmetrical bilateral with very short end arm), making unilateral arms essential for complete vineyard development. Some traditional training systems deliberately employ unilateral architecture throughout vineyards rather than just at ends, reflecting historical practice or specific regional approaches. Transitional unilateral configurations are common during bilateral cordon establishment, with young vines initially trained to single arm (year 2-3), adding second arm once vigor permits (year 3-5), with this staged approach reducing early crop load supporting better establishment compared to simultaneous bilateral development demanding high resource allocation.
Einschenkel
one-sided arm
single arm
unilaterale Arm
Unilateral Permanent Arm
SO4 (V. berlandieri × V. riparia) exemplifies universal compatibility, successfully grafted with hundreds of V. vinifera cultivars worldwide from Riesling in Germany to Shiraz in Australia without any documented cultivar-specific incompatibility issues over 100+ years of commercial use. 110 Richter (V. berlandieri × V. rupestris) shows similarly universal compatibility, supporting Tempranillo, Sangiovese, Grenache, Cabernet Sauvignon, and diverse cultivars across Mediterranean viticulture without compatibility concerns. 3309 Couderc (V. riparia × V. rupestris) demonstrates universal compatibility in Burgundy and Champagne, grafted successfully with Pinot Noir, Chardonnay, Pinot Meunier, Gamay, and Aligoté across decades of regional use. Kober 5BB (V. berlandieri × V. riparia) provides universal compatibility throughout Central European viticulture with Grüner Veltliner, Zweigelt, Blaufränkisch, and numerous other cultivars.
A graft affinity rootstock role that inheres in a grafted rootstock by virtue of its capacity to form successful, durable graft unions with the full range of Vitis vinifera cultivars and most Vitis interspecific hybrids, without cultivar-specific compatibility limitations. Universal compatibility rootstocks establish functional vascular connections with essentially any scion genotype within the Vitis genus, showing consistent callus bridge formation, rapid vascular differentiation (xylem and phloem continuity typically established within 30-60 days post-grafting), strong mechanical union strength, and absence of delayed incompatibility symptoms over vineyard lifespan (30-50+ years). This broad compatibility reflects minimal biochemical incompatibility factors, coordinated wound responses, and cellular recognition mechanisms that do not discriminate among Vitis genotypes. Universal compatibility enables flexible rootstock deployment without cultivar-specific restrictions, simplifying nursery production and vineyard planning. The major commercially important rootstock groups (V. berlandieri × V. riparia, V. berlandieri × V. rupestris, V. riparia × V. rupestris) all demonstrate universal or near-universal compatibility with V. vinifera scions.
Universal_Compatibility_Rootstock_Role represents the default condition for mainstream commercial rootstocks, where compatibility issues do not constrain rootstock-scion combinations. The V. berlandieri × V. riparia, V. berlandieri × V. rupestris, and V. riparia × V. rupestris hybrid groups all show universal compatibility, reflecting the close genetic relationships and interfertility among these American Vitis species and with V. vinifera. Universal compatibility simplifies viticultural decision-making by eliminating compatibility as a selection criterion, allowing growers to choose rootstocks based solely on edaphic adaptation, vigor, and stress tolerance. Compatibility testing for these established rootstocks is no longer routinely performed.
breit verträgliche Unterlage
broadly compatible rootstock
fully compatible rootstock
uneingeschränkt kompatible Unterlage
universal compatibility rootstock
universell kompatible Unterlage
unrestricted compatibility rootstock
Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22 Hartmann HT et al. (2011) Hartmann and Kester's Plant Propagation 8th Ed, Pearson Whiting JR (2004) Grapevine rootstocks, in Viticulture Vol 1 Resources, Winetitles pp.: 167-188
Universal Compatibility Rootstock Role
The unknown parentage note for Zinfandel (syn. Primitivo, Crljenak Kaštelanski, Tribidrag) documents the progressive resolution of a historically enigmatic origin — SSR analysis by Sefc et al. (2000) and Maletic et al. (2004) established Zinfandel's identity with the Croatian cultivar Crljenak Kaštelanski found in a single old vineyard in Kaštela, and subsequent analysis identified Tribidrag as the original Croatian name, but the parentage of Tribidrag/Zinfandel itself remained unresolved despite screening against panels of over 1,500 cultivars including extensive Croatian, Montenegrin, and Italian accessions — one parent has been tentatively identified as Dobričić (a red cultivar from the island of Šolta) based on half-parentage analysis at 32 SSR loci, while the second parent remains unknown and is presumed to be an extinct or uncollected cultivar from the Dalmatian coastal region, with continued screening of newly discovered old-vine accessions from Croatian and Montenegrin collections ongoing.
A parentage information record that explicitly documents the absence of identified parents for a Vitis vinifera cultivar, specifying the extent of parentage investigation conducted (marker systems applied, candidate panel size, databases searched), any partial results (one parent identified, candidate parents excluded), and the probable reasons for the parentage remaining unresolved (parent cultivar extinct, not represented in available germplasm collections, or insufficient marker resolution), distinguishing cultivars whose parentage has been actively investigated without success from those that have not yet been subjected to molecular parentage analysis.
Unknown_Parentage_Note provides the explicit documentation of unresolved parentage that is essential for distinguishing "investigated but unresolved" from "not yet investigated" — a critical distinction for directing future research priorities and managing expectations about cultivar genealogical knowledge. The majority of the approximately 10,000 genetically distinct V. vinifera cultivars documented in global databases have unknown parentage, and for most of these no molecular parentage analysis has been attempted: the subset of cultivars that have been actively investigated without full parentage resolution (estimated at 200-300 cultivars as of 2023) represent cases where one or both parents are likely extinct, uncollected, or represented in germplasm collections under unrecognized identities. The progressive expansion of genotyped reference panels — from a few hundred cultivars in early studies to over 5,000 accessions in current databases — continues to resolve previously unknown parentages as newly genotyped accessions are matched against existing orphan cultivars, suggesting that many currently unknown parentages will eventually be resolved as germplasm collection and genotyping efforts extend to undersampled regions (the Caucasus, Central Asia, the Balkans, and Iberian landraces).
Abstammung-unbekannt-Eintrag
Unbekannte-Abstammung-Vermerk
orphan cultivar parentage note
parentage unknown record
ungeklärte Elternschafts-Dokumentation
unresolved parentage documentation
Maletic E et al. (2004) Zinfandel, Dobričić, and Plavac Mali — the genetic relationship among three cultivars of the Dalmatian coast of Croatia, American Journal of Enology and Viticulture 55: 174-180 Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC), Vitis 54: 155-157 Lacombe T et al. (2013) Large-scale parentage analysis in an extended set of grapevine cultivars, Theoretical and Applied Genetics 126: 401-414
Unknown Parentage Note
An upper canopy apical leaf at node 13 on a shoot of Vitis vinifera cv. Sangiovese in Chianti, Italy, positioned in the upper canopy zone 45 cm above median canopy height on a vertically trained VSP system, receiving approximately 78% of full sunlight (daily PPFD averaging 1250 μmol photons m⁻² s⁻¹) with minimal shading from sparse overlying foliage, measuring 13.8 cm length by 12.5 cm width representing 76% of mature leaf dimensions for this position, exhibiting medium-dark green coloration with chlorophyll content of 435 μg cm⁻², developing sun leaf anatomy with two approaching-mature palisade layers totaling 72 μm thickness and thick cuticle of 5.5 μm, demonstrating photosynthesis rate of 10.8 μmol CO₂ m⁻² s⁻¹ representing 78% of mature capacity and approaching carbon balance with daily photosynthetic production of 38 mg carbohydrate approximately equaling respiratory and minimal remaining growth costs, occurring at 3 weeks post-emergence during final expansion phase with maturity expected within 10-14 days, measured at shoot BBCH stage 81 (beginning of berry ripening) in early August, representing well-positioned upper canopy apical leaf with favorable light exposure and substantial functional contribution.
An apical leaf positioned in the upper canopy zone occupying elevated positions in the three-dimensional canopy architecture in Vitis vinifera, characterized by location in nodes representing the distal portion of shoots positioned above median canopy height, favorable light exposure typically receiving 60-90% of full sunlight due to elevated position with minimal shading from overlying structures, developmental stages ranging from late expansion through approaching maturity depending on shoot extension status with leaves 2-4 weeks post-emergence, intermediate to near-mature dimensions typically 60-85% of final mature size, medium to dark green coloration reflecting substantial chlorophyll accumulation, developing sun leaf anatomical characteristics including multiple palisade layers, thick cuticle, and high chlorophyll per unit area responding to high light exposure during development, increasing photosynthetic capacity typically 60-85% of mature leaf rates transitioning toward carbon balance or slight net source status, and functional role contributing meaningfully to shoot and whole-vine carbon economy particularly if shoot extension ceases allowing maturation completion. Upper canopy apical leaves benefit from favorable position receiving high light availability without the extreme youth and developmental immaturity of terminal positions, creating optimal conditions for rapid development toward functional maturity, high photosynthetic potential through sun leaf differentiation, and substantial contribution to canopy carbon gain particularly during mid to late season when these leaves represent the youngest mature or near-mature photosynthetic surfaces.
Upper canopy apical leaves occupy favorable positions combining apical zone characteristics including relative youth and developmental advancement potential with elevated canopy positions providing excellent light exposure. These leaves avoid both the extreme immaturity and vulnerability of terminal positions and the progressive shading that affects lower canopy zones, creating optimal conditions for rapid high-quality development. Sun leaf characteristics develop in response to high light during expansion, with elevated positions typically receiving 60-90% full sun supporting anatomical differentiation toward multiple palisade layers, thick cuticles, and high photosynthetic enzyme concentrations. Functional transition from sink to source occurs relatively rapidly in upper canopy positions due to favorable light supporting high photosynthetic rates that quickly exceed respiratory and growth costs. Upper canopy apical leaves often represent the most productive recent additions to canopy photosynthetic capacity, contributing substantially to vine carbon balance particularly during fruit ripening periods when their source function supports berry sugar accumulation. Position effects create systematic patterns with uppermost apical leaves showing youngest developmental stages while lower apical zone leaves show advancing maturity, creating functional gradients across the apical canopy zone.
elevated apical leaf
gut-exponiertes oberes Weinblatt
high-position young leaf
hoch-positioniertes junges Blatt
obere Kanopie apikales Blatt
well-exposed upper leaf
Based on Smart (1985) "Principles of grapevine canopy microclimate manipulation with implications for yield and quality" American Journal of Enology and Viticulture 36=230-239 Poni et al. (2009) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 60=397-407 Schultz (1996) "Leaf absorptance of visible radiation in Vitis vinifera L.= estimates of age and shade effects with a simple field method" Scientia Horticulturae 66=93-102 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 179-214.
Upper Canopy Apical Leaf
An upper lamina surface on a mature sun leaf at node 7 of Vitis vinifera cv. Syrah in the Rhône Valley, France, positioned facing south receiving approximately 85% of full sunlight with peak midday irradiance exceeding 2000 μmol photons m⁻² s⁻¹, exhibiting dark green coloration from underlying palisade mesophyll with chlorophyll concentration of 520 μg cm⁻², thick cuticle of 6.4 μm visible as glossy surface sheen reflecting incident light, epicuticular wax deposits evident from water droplet beading and rolling, relatively smooth texture with sparse trichomes (density 8 trichomes mm⁻² compared to 45 trichomes mm⁻² on lower surface), very low stomatal density of 12 stomata mm⁻² representing only 6% of total leaf stomatal population concentrated on lower surface, epidermal cells tightly packed with thick outer walls (4.2 μm) providing structural integrity and protection, surface temperature frequently 2-4°C above ambient during high radiation periods due to direct solar heating, showing no visible stress symptoms despite high light exposure indicating adequate photoprotection mechanisms, measured at BBCH stage 83 (berries developing color) in mid-August, representing typical upper surface characteristics adapted for high light exposure and environmental stress resistance.
The adaxial surface of the grapevine leaf blade facing upward toward incident light in Vitis vinifera, characterized by exposure to higher light intensities and direct solar radiation, thick cuticle deposition typically 4-7 μm providing protection against water loss and UV radiation, relatively smooth texture with reduced trichome density compared to lower surface, darker green coloration from high chlorophyll concentrations in underlying palisade mesophyll layers, low stomatal density typically 5-20 stomata mm⁻² representing only 3-8% of total leaf stomata, tight cell packing in underlying epidermis with relatively small cells and minimal intercellular spaces, and functional specialization for light capture and protection with cuticle and epidermis filtering and regulating light penetration to photosynthetic tissues. The upper lamina surface serves as the primary light-receiving interface, experiencing maximum solar radiation exposure including potentially damaging UV wavelengths, bearing the brunt of environmental stresses including heat, desiccation, and particulate deposition, exhibiting specialized adaptations including thick cuticle providing water retention and UV screening, epicuticular wax deposits creating water-repellent properties facilitating droplet runoff, and pigmentation patterns including anthocyanins in some cultivars providing additional photoprotection under high light or stress conditions.
Upper lamina surface is distinguished from lower surface by anatomical and functional specializations for light reception and environmental protection rather than gas exchange. The thick cuticle, low stomatal density, smooth texture, and tight cell packing reflect this specialization. Sun versus shade leaves show differential upper surface development, with sun leaves exhibiting thicker cuticles (5-7 μm), denser wax deposits, and occasionally anthocyanin pigmentation compared to shade leaves with thinner cuticles (3-4 μm) and reduced protective investments. The upper surface experiences maximum environmental exposure including intense solar radiation, temperature extremes, desiccating conditions, and atmospheric pollutant deposition, requiring robust protective mechanisms. Water repellency from cuticular waxes facilitates rapid shedding of rain, dew, and irrigation water, reducing disease infection opportunities and preventing light scattering by surface water films. Some cultivars show distinctive upper surface characteristics including glabrous (hairless) versus pubescent conditions, glossy versus matte finishes, or pigmentation patterns that contribute to ampelographic identification. Functional measurements typically focus on underlying tissues accessed through upper surface including chlorophyll fluorescence assessed from adaxial side and spectral reflectance measurements capturing pigment signatures.
Adaxialseite
Blattoberseite
adaxial surface
obere Blattfläche
top lamina surface
upper leaf surface
Based on Schultz (1996) "Leaf absorptance of visible radiation in Vitis vinifera L.= estimates of age and shade effects with a simple field method" Scientia Horticulturae 66=93-102 Patakas et al. (2003) "Differences in stomatal and photosynthetic responses of Vitis vinifera L. (cv. Sultana) leaves in response to progressive water stress" Vitis 42=113-115 Kolb et al. (2001) "Effects of natural intensities of visible and ultraviolet radiation on epidermal ultraviolet screening and photosynthesis in grape leaves" Plant Physiology 127=863-875 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 95-136.
Upper Lamina Surface
An upper lateral lobe representing the right upper lateral on a mature leaf at node 6 of Vitis vinifera cv. Cabernet Sauvignon in Bordeaux, France, positioned as the first lateral lobe below the apical lobe, vascularized by the first primary lateral vein branching from median vein at 62-degree angle at 3.2 cm from petiolar junction, measuring 5.2 cm maximum width and 5.5 cm length representing the largest lateral lobe on this leaf, separated from apical lobe by upper right lateral sinus of 3.0 cm depth creating well-defined boundary and from lower lateral lobe by middle right lateral sinus of 2.4 cm depth, lobe apex exhibiting acute configuration with approximately 52-degree angle, lobe margins showing slightly convex curvature with characteristic heterogeneous tooth pattern, lobe area of 22.8 cm² representing 19% of total blade area making it the second-largest lobe after the apical lobe, prominent development contributing substantially to overall leaf width of 15.8 cm (maximum width occurring at upper lateral lobe level), measured at BBCH stage 75 (berries pea-sized) in late June, representing typical upper lateral lobe morphology with substantial development characteristic of this deeply dissected cultivar.
A lateral lobe positioned in the upper intermediate zone between the apical lobe and lower lateral lobes in Vitis vinifera, characterized by position as the first pair of lateral lobes below the apical lobe, vascularization by the first pair of primary lateral veins branching from the median vein typically at 50-70 degree angles, larger size compared to lower lateral lobes generally representing the largest and most prominent lateral lobes, separation from the apical lobe by upper lateral sinuses and from lower lateral lobes by middle lateral sinuses, cultivar-specific characteristics including relative size often approaching or occasionally exceeding apical lobe dimensions, degree of development from weakly defined to prominently isolated, apex configurations, and overall prominence serving as diagnostic features. Upper lateral lobes represent the most prominent lateral lobe pair, developing as major blade divisions creating substantial lobe area typically 15-22% per lobe of total blade area, exhibiting pronounced development in deeply dissected cultivars contrasting with weak development in weakly lobed varieties, and contributing significantly to overall leaf width and outline shape determining maximum blade width often occurring at upper lateral lobe level.
Upper lateral lobes are distinguished from lower lateral lobes by superior position closer to apical lobe and typically larger size. The upper designation emphasizes position in the first lateral lobe pair below the apical lobe, corresponding to first-order lateral vein branching. Size superiority over lower laterals is typical but not universal, with some cultivars showing more balanced lateral lobe sizes. Upper lateral lobes often determine maximum blade width, with the widest blade dimension commonly occurring at or near upper lateral lobe level particularly in deeply dissected cultivars with prominent upper laterals. Separation from apical lobe through upper lateral sinuses shows high diagnostic importance, with deep upper sinuses creating prominent isolated upper laterals while shallow sinuses produce weakly separated configuration. Development ranges from prominent in deeply dissected cultivars (Cabernet Sauvignon, Syrah) to weak in poorly lobed varieties (some Gewürztraminer clones, Chardonnay). Bilateral symmetry is typical with left and right upper laterals showing similar size and shape, though subtle asymmetries occur. Functional contribution involves substantial area provision typically 30-40% combined area of both upper laterals, making them major photosynthetic contributors. Ampelographic assessment includes upper lateral size relative to apical lobe, degree of separation, apex characteristics, and overall prominence contributing to cultivar identification.
first lateral lobe
oberer Seitenlappen
primary side lobe
primärer Seitenlappen
superior lateral lobe
superiorer Laterallappen
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 90-135 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin This et al. (2006) "Development and characterization of a standard set of 15 microsatellite markers for grapevine genotyping and fingerprinting" Vitis 45=55-61 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 173-194.
Upper Lateral Lobe
An upper lateral secondary inflorescence at node 5 on a shoot of Vitis vinifera cv. Syrah in the Rhône Valley, France, measuring 15.2 cm length representing robust secondary development, positioned in the upper shoot region receiving approximately 82% of full sunlight with minimal shading from sparse overlying foliage, located 38 cm from shoot base and 32 cm from current shoot tip creating upper-intermediate position on this 10-node shoot bearing two inflorescences (primary at node 3, this secondary at node 5), exhibiting well-developed compound paniculate architecture with central rachis bearing 4 orders of branching, rachis diameter of 3.1 mm at base representing substantial construction, bearing approximately 480 flowers, showing dimensions approaching the primary inflorescence (16.8 cm, 650 flowers) representing only 9% length reduction and 26% flower number reduction indicating favorable development conditions, positioned in actively growing shoot region during inflorescence development receiving good resource allocation from vigorous vegetative growth, developmental stage at BBCH 65 (full bloom) showing minimal phenological delay compared to primary, upper position providing excellent air circulation reducing disease pressure, expected to contribute approximately 38% of shoot yield, measured in mid-June during peak bloom, representing well-developed upper lateral secondary with dimensions approaching primary due to favorable position.
A lateral secondary inflorescence positioned in the upper portion of the shoot reproductive zone in Vitis vinifera, characterized by position at node 5 or occasionally node 6 creating upper-intermediate location when multiple inflorescences occur, proximity to the shoot apex relative to other inflorescences on shoots with 2-3 total inflorescences, favorable light exposure typical of upper shoot positions receiving 70-90% of full sunlight, reduced shading from overlying canopy structures, position in actively extending shoot regions during inflorescence development, dimensions typically 12-16 cm representing good development for secondary inflorescences, moderate to good branching complexity showing 3-4 orders, and functional characteristics including good resource availability from active shoot growth supporting relatively robust secondary development. Upper lateral secondary inflorescences represent the well-positioned variant of secondary inflorescences, occurring in upper shoot zones with favorable light and resource conditions, exhibiting relatively good development approaching primary dimensions in some cases, and contributing substantially to shoot yield particularly when crop loads favor secondary retention.
Upper lateral secondary inflorescences represent the favorably positioned variant, distinguished from lower lateral secondaries by upper shoot location and typically superior development. The upper designation emphasizes the elevated position relative to other inflorescences creating favorable conditions. Position at node 5 on shoots bearing 2-3 total inflorescences creates upper-intermediate or upper location depending on total inflorescence number and shoot length. Light exposure advantages from upper positioning typically provide 70-90% full sun compared to 50-70% for lower positions, supporting robust photosynthetic resource generation during development. Active shoot growth association means upper secondaries develop during peak vegetative extension when resource availability is high, potentially enhancing development compared to secondaries on slower-growing shoots. Size approaching primary dimensions in favorable cases reflects the positional advantages, with some upper secondaries showing only 10-20% reduction from primaries compared to typical 30-40% reduction. Branching complexity often shows good development with 4 orders common approaching primary elaboration. Flower number may exceed 450-500 in well-developed examples approaching or occasionally exceeding small primaries. Phenological development often shows minimal delay with upper secondaries sometimes flowering nearly simultaneously with primaries. Yield contribution can be substantial at 35-45% particularly in high-fertility cultivars with robust secondary development. Management implications include potential for retention in balanced crop load scenarios where upper secondaries provide meaningful yield without excessive overcropping due to their substantial development.
elevated secondary inflorescence
erhöhte Zweitinfloreszenz
gut-beleuchtete laterale Sekundärrispe
obere laterale sekundäre Infloreszenz
upper position secondary
well-lit lateral secondary
Based on Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 259-295 Carmona et al. (2008) "Grapevine inflorescence and flower development" in Biology of the Grapevine, Cambridge University Press pp. 219-254 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 399-438.
Upper Lateral Secondary Inflorescence
An upper lateral sinus representing the right upper lateral sinus on a mature leaf at node 7 of Vitis vinifera cv. Syrah in the Rhône Valley, France, positioned between the apical lobe and right upper lateral lobe, measuring 3.4 cm depth from sinus base to the imaginary line connecting the apical and upper lateral lobe edges, width of 1.6 cm at the sinus opening, exhibiting characteristic U-shaped contour with rounded sinus base, sinus base positioned 4.8 cm from petiolar junction representing approximately 32% of distance to blade apex, left margin formed by the right edge of the apical lobe showing smooth convex curvature, right margin formed by the left edge of the upper lateral lobe showing corresponding convex curvature, sinus creating clear separation between apical and upper lateral lobes contributing to well-defined five-lobed architecture, sinus depth substantially greater than lower lateral sinus depth (2.2 cm) on the same leaf reflecting characteristic deep dissection in upper blade region, bilateral symmetry with left upper lateral sinus showing similar depth (3.2 cm) and configuration, measured at BBCH stage 81 (beginning of berry ripening) in early August, representing typical deep upper lateral sinus morphology characteristic of this deeply dissected cultivar.
The sinuses positioned between the apical lobe and upper lateral lobes in Vitis vinifera, characterized by location in the upper blade region separating the terminal apical lobe from the first pair of lateral lobes, bilateral occurrence as paired left and right upper lateral sinuses, variable depth typically 1.5-4.5 cm depending on cultivar lobe dissection patterns, U-shaped, V-shaped, or intermediate contours, width at opening ranging from 0.8-3 cm, role in determining the degree of apical lobe separation and prominence, contribution to overall blade dissection affecting leaf outline shape and perimeter, and cultivar-specific characteristics including depth, shape, and symmetry serving as important secondary diagnostic features. Upper lateral sinuses represent critical structural features determining apical lobe isolation and prominence, with deep upper lateral sinuses creating well-defined separated terminal lobes while shallow sinuses produce poorly differentiated apical regions, exhibiting genetic control creating cultivar-diagnostic patterns though with greater environmental sensitivity than petiolar sinuses, and contributing to overall lobe architecture characterizing cultivar morphology.
Upper lateral sinuses are distinguished from lower lateral sinuses by position between apical and upper lateral lobes rather than between lateral lobe pairs, and from petiolar sinuses by upper blade location. Depth shows wide variation from shallow (1-2 cm) in weakly lobed cultivars to deep (4-5 cm) in strongly dissected varieties, with depth determining degree of apical lobe isolation and prominence. Deep upper lateral sinuses create prominent well-separated terminal lobes characteristic of deeply dissected cultivars (Syrah, Mourvèdre) while shallow sinuses produce weakly differentiated apical regions in poorly lobed varieties (Chardonnay, some Gewürztraminer clones). Shape characteristics include U-shaped (rounded base, common) and V-shaped (acute base, less common) configurations. Bilateral symmetry is typical with left and right upper lateral sinuses showing similar depths and shapes, though subtle asymmetries occur. Genetic control is moderate to strong with upper lateral sinus depth showing cultivar-specific patterns, though environmental modification occurs with vigor and resource availability affecting final depth to some extent. Functional implications include contribution to apical lobe isolation enabling independent movement and affecting overall blade flexibility. Ampelographic utility is substantial as secondary diagnostic feature, with upper lateral sinus depth combined with petiolar sinus configuration and other features enabling cultivar identification, particularly useful for distinguishing among deeply versus weakly lobed cultivar groups.
apical-lateral sinus
apikale Lateralsinus
obere Seitenbucht
superior lateral sinus
superiore Lappeneinbuchtung
upper lobe sinus
Based on Galet (2000) "A Practical Ampelography= Grapevine Identification" Cornell University Press pp. 1-57 OIV (2009) "OIV Descriptor List for Grape Varieties and Vitis Species" 2nd Edition, Organisation Internationale de la Vigne et du Vin Chitwood et al. (2014) "Leaf asymmetry as a developmental constraint imposed by auxin-dependent phyllotactic patterning" The Plant Cell 26=609-621 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 137-172.
Upper Lateral Sinus
The upper lateral sinus description of Riesling documents moderately deep upper lateral sinuses (OIV 070:5) with an open V-shaped base (OIV 080:2) and no teeth at the sinus floor — the sinuses extend approximately 40-50% of the distance from margin to midvein, creating clearly defined but not dramatically separated lobes that contribute to Riesling's characteristic moderately dissected pentagonal leaf outline. The V-shaped sinus base (converging straight sides meeting at an acute angle) distinguishes Riesling from cultivars with U-shaped sinus bases (parallel sides meeting in a rounded curve, as in Müller-Thurgau) and from cultivars with lyre-shaped sinus bases (convex sides creating a wider opening than the V-shape, as in some Muscat types), providing additional discrimination within the moderately-deep-lobed cultivar group.
A sinus description component that records the morphological characteristics of the upper lateral sinuses — the indentations between the terminal (central) lobe and the two upper lateral lobes — documenting sinus depth (OIV 070), shape of the sinus base (OIV 080), and the presence of teeth within the sinus, with the upper lateral sinuses being the deepest lateral sinuses in most V. vinifera cultivars and a primary determinant of the overall leaf dissection pattern.
Upper_Lateral_Sinus_Description documents the sinuses that most strongly influence the visual impression of leaf dissection — because the upper lateral sinuses are typically the deepest sinuses in V. vinifera leaves, they create the primary division between the terminal lobe and the lateral portions of the blade, and their depth largely determines whether a leaf appears compact and entire or dramatically dissected. The shape of the sinus base provides additional cultivar-diagnostic information — OIV 080 distinguishes between V-shaped (sides converging to a point), U-shaped (sides parallel then curving to meet), and lyre-shaped (sides initially converging then diverging before meeting, creating a wine-glass profile) base forms that reflect differences in the developmental geometry of sinus formation: the sinus base shape is determined by the pattern of growth inhibition at the leaf margin during primordial development — sharp growth arrest produces V-shaped sinuses, gradual inhibition produces U-shaped sinuses, and complex growth dynamics with secondary expansion at the sinus floor produce the characteristic lyre shape. The presence of a tooth at the sinus base (a small projection of blade tissue at the deepest point of the sinus) is an additional character observed in some cultivars that provides fine-scale discrimination.
Hauptseitenbuchten-Morphologie
main lateral indentation morphology
obere Lateralsinus-Charakterisierung
obere Seitenbuchten-Beschreibung
superior sinus description
upper lateral sinus characterization
Upper Lateral Sinus Description
In Sauvignon Blanc vineyards of Marlborough, New Zealand, at BBCH 55-57, upper primary inflorescence branches (positions 13-18 from the rachis base among 18 total primary branches) measure only 1.5-2.8 cm in length and bear 1-2 secondary branches each, contributing merely 80-120 flowers to the total inflorescence complement of 350-480 flowers. Nebbiolo in Barolo exhibits particularly abbreviated upper primary branches of 0.8-2 cm with minimal or absent secondary branching, creating sharply tapered, cylindrical clusters. The pronounced size gradient from lower to upper primary branches in Riesling (Mosel Valley) creates the characteristic elegant, tapering cluster morphology valued for premium wine production.
An upper primary inflorescence branch is a primary inflorescence branch located in the apical third to half of the main inflorescence rachis, characterized by reduced length, diameter, and sub-branching complexity compared to lower and middle primary branches, contributing proportionally less to total flower number while determining the apical taper and overall length of the mature cluster. Upper primary branches occupy positions approximately 60-95% of the distance from the rachis base to apex (measured at BBCH 55), with the terminal-most primary branch typically positioned 2-8 mm below the rachis apex. These branches represent the latest-initiated primary branches during inflorescence primordium development within the dormant bud and consequently exhibit less advanced developmental elaboration at any given phenological stage. Branch length typically ranges from 0.5-3.5 cm (cultivar-dependent), with basal diameter often 0.5-1.2 mm, representing 25-50% reduction compared to lower primary branch dimensions. Upper primary branches characteristically bear fewer secondary branches (0-3 secondary branches per upper primary branch in most cultivars occasionally 4-5 in extremely loose-clustered types), with secondary branches themselves showing limited or absent tertiary subdivision. This reduced architectural elaboration results in upper primary branches bearing only 5-20 flowers each, representing 15-25% of total inflorescence flower number despite constituting 30-40% of primary branch count. The abbreviated, less subdivided character of upper primary branches creates the characteristic pointed or tapered apex of cylindrical and conical cluster morphologies, distinguishing cultivars with pronounced apical taper from those with more uniform branch length distribution. Vascular anatomy of upper primary branches shows smaller xylem vessel diameters (20-40 μm) compared to lower branches (35-65 μm), reflecting reduced resource demand for the limited flower complement. Upper branch development proceeds 2-5 days behind lower primary branches during BBCH 53-57, with some terminal upper branches still emerging from the rachis while basal lower branches have already completed secondary branch initiation. The reduced development of upper primary branches contributes to the natural density gradient in most clusters, with looser berry packing in apical regions facilitating air circulation and reducing disease pressure, though extremely reduced upper branch development can compromise cluster appearance and market value in table grape production.
Upper_Primary_Inflorescence_Branch is distinguished from Lower_Primary_Inflorescence_Branch by position (upper branches occupy the apical third of rachis length), reduced dimensions, delayed development, and reduced sub-branching. The boundary between 'upper' and 'lower' primary branches is necessarily somewhat arbitrary but follows the natural inflection point in the basipetal gradient of branch length and complexity, typically occurring at 55-65% of rachis length from base. These branches are less frequent targets for cultural manipulation but are important for cluster morphology description and for understanding resource allocation patterns within the inflorescence. Extreme reduction of upper primary branches can indicate previous-season stress during late stages of inflorescence primordium differentiation.
apical primary branch
apikaler Hauptast
oberer Hauptast der Infloreszenz
oberer Primärast
upper first-order branch
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 89-91 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 77-80 Galet, P. (2000). A Practical Ampelography= Grapevine Identification, pp. 45-52.
Upper Primary Inflorescence Branch
In Chardonnay vineyards of Chablis at BBCH 55-57, upper secondary branches on primary branches measure only 0.8-1.8 cm length and typically lack tertiary branches, contributing 5-12 flowers per upper secondary to the moderately compact clusters of 80-120 berries. Pinot Noir in Champagne exhibits similarly abbreviated upper secondary branches of 0.5-1.5 cm on all but basal primary branches, creating the characteristically compact, cylindrical cluster morphology essential for the region's sparkling wine production. The pronounced size reduction from lower to upper secondary branches on individual primary branches in Riesling contributes to the elegant, gradually tapering cluster form.
An upper secondary inflorescence branch is a secondary inflorescence branch that arises from the apical half of a primary inflorescence branch, characterized by reduced length, diameter, and tertiary sub-branching compared to lower secondary branches on the same primary branch, contributing proportionally less to the flower complement while determining the degree of apical elaboration of primary branches. Upper secondary branches occupy positions approximately 50-95% of the distance from the primary branch base to apex, with the terminal-most upper secondary typically positioned 0.5-3 mm below the primary branch apex. These branches represent the latest-initiated secondary branches during inflorescence primordium development and consequently exhibit less advanced developmental elaboration at any phenological stage. Branch length typically ranges from 0.3-2.5 cm depending on parent primary branch vigor and cultivar type, with basal diameter often 0.3-0.8 mm, representing 40-60% reduction compared to lower secondary branch dimensions on the same primary. Upper secondary branches characteristically bear fewer or no tertiary branches (0-2 tertiary branches per upper secondary, often completely absent in compact-clustered cultivars), with tertiary branch emergence delayed or suppressed during BBCH 55-57. This reduced architectural elaboration results in upper secondary branches bearing only 3-15 flowers each, representing 30-45% of the flower number contributed by their parent primary branch despite constituting 50-60% of the secondary branch count on that primary. The abbreviated, less subdivided character of upper secondary branches creates the characteristic pointed or tapered terminus of individual primary branches, contributing to the overall fineness and elegance of cluster morphology, particularly in cultivars valued for loose cluster architecture. Vascular anatomy of upper secondary branches shows reduced vessel diameters (18-35 μm) compared to lower secondaries (25-50 μm), reflecting decreased resource demand for the limited flower complement. Upper secondary branch development proceeds 1-4 days behind lower secondaries on the same primary during BBCH 53-57, with some terminal upper secondaries still emerging while basal lower secondaries have already initiated tertiary branches. The reduced development of upper secondary branches contributes to natural density gradients within individual primary branches, complementing the basipetal gradient of primary branch development along the main rachis to create two-dimensional density patterning across the entire inflorescence. In extremely compact-clustered cultivars, upper secondary branches may be completely absent from apical primary branches, while even in loose-clustered types, upper secondaries on upper primary branches show minimal elaboration.
Upper_Secondary_Inflorescence_Branch is distinguished from Lower_Secondary_Inflorescence_Branch by position on the parent primary branch (apical half versus basal half), reduced dimensions, delayed development, and reduced or absent tertiary subdivision. The boundary between 'upper' and 'lower' secondary branches follows the natural inflection point in the basipetal gradient of branch length and complexity along the primary branch axis, typically occurring at the midpoint of primary branch length. These branches contribute to overall cluster architecture primarily through their modest development, creating apical taper of primary branches and contributing to loose packing in apical cluster regions. Extreme reduction or absence of upper secondary branches can indicate previous-season stress during late stages of inflorescence differentiation or represent a fixed cultivar characteristic in compact-clustered types.
apical secondary branch
apikaler Seitenast zweiter Ordnung
distal second-order branch
oberer Sekundärast
oberer sekundärer Blütenstandsast
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 91-93 Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 78-81 Galet, P. (2000). A Practical Ampelography, pp. 48-51.
Upper Secondary Inflorescence Branch
The upper surface description of Tempranillo documents strongly blistered interveinal areas (OIV 075:7) with pronounced convex bulging between the main and secondary veins — the interveinal mesophyll expands more rapidly than the vein-constrained tissue, pushing the upper epidermis upward to create a deeply rugose surface with 2-4 mm elevation differences between vein troughs and interveinal crests, giving the leaf a characteristically crumpled, textured appearance. The adaxial surface is dark green with a matte finish (low glossiness, minimal surface wax layer), and shows no trichomes visible to the naked eye (the adaxial surface of most V. vinifera cultivars is glabrous or nearly so, in contrast to the typically pubescent abaxial surface), with the overall surface texture creating a visually and tactilely distinctive leaf that is immediately distinguishable from the flatter, smoother upper surfaces of Garnacha (OIV 075:3) and Graciano (OIV 075:3).
A leaf surface description component that records the textural characteristics of the adaxial (upper) surface of the fully expanded leaf blade, documenting the degree of blistering or convexity of the interveinal areas (OIV 075), the overall surface profile (flat, undulate, involute, or revolute), glossiness or mattness of the cuticle, and the presence or absence of surface waxiness, providing cultivar-diagnostic characters based on the epidermal and mesophyll architecture visible on the photosynthetically active upper leaf surface.
Upper_Surface_Description focuses on the adaxial leaf surface that faces the observer when examining leaves in situ on the vine. The dominant character of the upper surface is the degree of blistering (bullosity) — the three-dimensional surface topography created by differential growth between vein-associated and interveinal tissue during blade expansion. Blistering is scored on the OIV 1-9 scale from flat (state 1, the interveinal surface level with the veins) through moderate (state 5, gentle convexity) to very strong (state 9, deeply rugose surface with pronounced interveinal bulges): the blistering character is primarily genetically determined, with QTL contributions from chromosomes 1 and 14 (Chitwood et al. 2014), though environmental factors (particularly nitrogen status and water availability affecting cell expansion rates) can modulate blistering intensity by 1-2 OIV scale points. The glossiness or mattness of the upper surface reflects the cuticle thickness and surface microstructure — some cultivars (e.g., Chasselas, Sultana) have distinctly glossy upper surfaces with visible light reflection, while most wine grape cultivars show matte surfaces with minimal specular reflection.
Blattoberseiten-Beschreibung
adaxial surface characterization
adaxiale Oberflächen-Charakterisierung
obere Blattflächen-Morphologie
upper blade surface morphology
upper leaf texture
Upper Surface Description
An upper trunk basal shoot on Vitis vinifera cv. Pinot Noir arising from trunk at 62cm height (18cm below original bilateral cordon position at 80cm height), activated following bot canker girdling and killing cordons while leaving upper trunk viable, shoot emerged in early May (prompt activation indicating localized damage rather than systemic stress), developed to 185cm length by mid-July, 22 leaves, no clusters but vigorous balanced growth, representing ideal trunk basal recovery from advantageous upper trunk position, minimal additional vertical development needed (only 18cm elevation required to reach target cordon height), this shoot trained as temporary fruiting position during first season (producing 8 vegetative shoots, 2.1kg photosynthetic contribution, zero crop as expected), second season topped at 80cm and lateral trained developing bilateral cordons restoring original training system architecture, complete productive structure restoration achieved by third season versus 4-5 seasons typically required from lower trunk positions.
A trunk basal shoot originating from upper trunk positions, characterized by emergence from trunk regions within 20-50cm of head or cordon origin points, elevated accessibility facilitating management, advantageous positioning requiring minimal vertical training before lateral structure development possible, and function as preferred trunk renewal source, convenient damage recovery mechanism, or efficient architectural modification option when upper trunk latent buds activate providing optimal starting positions for structural development. Upper trunk basal shoots represent ideal trunk recovery scenario, arising from positions approaching normal head or cordon heights requiring minimal additional vertical development, providing accessible convenient training locations, offering rapid pathway to productive structure restoration (often 2-3 seasons versus 4-5 for lower positions), and enabling high-quality structural outcomes through favorable starting geometry and reduced complications compared to low-origin alternatives.
Upper trunk basal shoots represent optimal basal recovery outcomes, with their elevated origin positions providing numerous advantages including minimal vertical development requirements (often already at or near target structural heights), convenient accessibility (standing-position management without bending), rapid productive restoration (shorter training timelines to functional structures), favorable shoot geometry (appropriate angles and positions for cordon or arm development), and reduced complications (elevated positions avoiding ground proximity issues). Upper trunk basal activation typically indicates localized damage (disease cankers, freeze damage, mechanical injury affecting specific trunk zones) rather than systemic problems, with viable upper trunk tissue and selective bud activation suggesting targeted recovery response. Management utilizes upper trunk basal shoots as preferred renewal sources when available, implementing efficient training protocols capitalizing on advantageous starting positions, potentially achieving productive structure restoration in 2-3 seasons versus longer timelines from lower positions, with upper trunk origin enabling high-quality architectural outcomes approaching or matching original training system geometry.
elevated trunk renewal shoot
high trunk shoot
kopfnaher Stockausschlag
near-head trunk shoot
oberer Stamm-basaltrieb
Based on Mundy & Manning (2010) "Budbreak and shoot development in Vitis vinifera following severe trunk damage" Australian Journal of Grape and Wine Research 16=310-316 Sosnowski et al. (2007) "The influence of grapevine cultivar and isolate of Eutypa lata on wood and foliar symptoms" Plant Disease 91=924-931.
Upper Trunk Basal Shoot
An upper trunk latent bud located 15cm below the cordon on Vitis vinifera cv. Cabernet Franc, activated following Esca disease symptoms that killed the right cordon, producing a water shoot 165cm in length during the first season, trained horizontally along the trellis wire to replace the dead cordon, with 8 renewal spurs retained during first dormant pruning, establishing 16 fruiting shoots with moderate crop (1.2 kg per vine) in the second season, returning to normal production (2.8 kg per vine) by the third season post-activation.
A trunk latent bud located in the upper portion of the permanent trunk structure, typically within 20-40cm of the cordon or head, characterized by proximity to existing vascular connections supporting fruiting wood, relatively easy access for shoot positioning and training, moderate vigor potential due to established xylem pathway to root system, and higher likelihood of activation compared to lower trunk positions due to closer proximity to hormonal signals from canopy. Upper trunk latent buds are preferred for cordon or arm replacement following damage or disease, as resulting shoots can be positioned into fruiting positions with minimal structural modification and shorter time to return to production, typically requiring only 1-2 seasons to establish productive fruiting wood compared to 3-4 seasons for lower trunk regeneration.
Upper trunk latent buds offer advantages for vineyard recovery and trunk disease management programs, providing faster return to production than lower trunk options while maintaining original vine height and trellis compatibility. However, upper trunk retraining can perpetuate trunk disease inoculum if the pathogen remains active in underlying wood, sometimes necessitating more aggressive trunk renewal from lower positions despite longer recovery time. The decision between upper versus lower trunk renewal depends on disease distribution, vine age, vineyard economics, and grower patience for yield recovery, with upper trunk offering speed but potentially compromised long-term trunk health.
high trunk bud
hohe Adventivknospe
obere Stammknospe
upper trunk epicormic bud
Based on Rolshausen et al. (2010) "Evaluation of pruning wound susceptibility and protection against fungi associated with grapevine trunk diseases" American Journal of Enology and Viticulture 61=113-119 Mundy & Manning (2010) "Budbreak and shoot development in Vitis vinifera following severe trunk damage" Australian Journal of Grape and Wine Research 16=310-316 Weber et al. (2007) "Grapevine trunk diseases in California" Practical Winery & Vineyard Journal 29=34-46.
Upper Trunk Latent Bud
An upper trunk water shoot on Vitis vinifera cv. Grenache in Châteauneuf-du-Pape, France, emerging at 105 cm trunk height (15 cm below the bilateral cordon positioned at 120 cm), developing following severe hail damage to the southern cordon arm that necessitated its complete removal during summer pruning, growing 185 cm by September with 20 nodes, internode lengths averaging 12-14 cm, leaves of 195 cm² average area showing normal scion characteristics, positioned ideally for training as a replacement cordon arm during the following season, requiring only 15 cm upward extension to reach target cordon height, and showing favorable vascular connections to the main trunk suggesting good long-term viability for framework incorporation.
A trunk water shoot emerging from the upper trunk region between the trunk midpoint and the first permanent arm, cordon, or head position in Vitis vinifera, characterized by proximity to established framework vascular systems, moderate to strong vigor reflecting intermediate resource access between lower trunk and arm positions, and strategic importance for potential framework reconstruction or training system modification without complete trunk replacement. Upper trunk water shoots typically develop in response to localized damage or disease in adjacent cordon or arm structures, severe reduction in canopy size through winter pruning or environmental damage, or intentional stimulation through strategic scoring or notching techniques applied by viticulturists seeking framework renewal from optimal positions. These shoots occupy architectural positions well-suited for developing replacement cordons or arms when original structures suffer from progressive trunk diseases, physical damage, or age-related decline, allowing framework reconstruction while maintaining established trunk height and preserving years of structural development. Management considerations for upper trunk water shoots include evaluation of their potential role in long-term training system evolution, assessment of underlying causes triggering their emergence, and strategic decisions about retention for framework renewal versus removal to maintain existing architectural plans.
Upper trunk water shoots are distinguished from lower trunk water shoots by position above the trunk midpoint and proximity to established framework structures, and from arm water shoots by origin on the main trunk rather than on permanent branch structures. These shoots represent the most favorable positions for framework renewal when needed, requiring less vertical development than lower trunk shoots and offering clearer architectural integration into existing training systems. The decision to retain versus remove upper trunk water shoots depends on framework condition, vine age, disease pressure in existing structures, and long-term vineyard management plans. Shoots retained for framework renewal typically require 2-3 seasons of managed development to achieve full productive capacity.
cordon-level trunk sucker
head-proximal water shoot
kopfnaher Stammtrieb
kordonnaher Wassertrieb
oberer Stammwasserschoss
upper stem water shoot
Based on Winkler et al. (1974) "General Viticulture" University of California Press pp. 302-318 Galet (2000) "General Viticulture" (English translation), Oenoplurimedia pp. 267-285 Creasy & Creasy (2009) "Grapes= Crop Production Science in Horticulture" CABI Publishing pp. 112-128 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 319-342.
Upper Trunk Water Shoot
Documentation of Upright_Growth_Disposition at UC Davis characterized shoot architecture in cultivars recognized for strongly vertical growth habit. Reference cultivars 'Cabernet Sauvignon', 'Merlot', 'Petit Verdot', and 'Tannat' were grown on minimal trellis systems allowing free shoot expression, with detailed angular and mechanical measurements. Upright_Growth_Disposition cultivars showed consistent distinguishing characteristics. 'Cabernet Sauvignon' exhibited archetypal upright growth with mean shoot angle of 12° from vertical at 50 cm shoot length, maintaining 15° even at 100 cm length with shoots weighing 85 g, and shoot tips ascending to 8° demonstrating persistent gravitropic correction. Internode flexural stiffness averaged 2.8 N·mm² (highest quartile among cultivars tested). 'Petit Verdot' showed even stronger upright tendency (10° at 50 cm, stiffness 3.2 N·mm²). 'Merlot' displayed typical Upright_Growth_Disposition (14° at 50 cm) with moderate stiffness (2.4 N·mm²). In VSP training trials, Upright_Growth_Disposition cultivars required positioning labor of only 12-18 hours/ha/season (1-2 positioning passes), compared to 45-65 hours/ha for trailing cultivars. Canopy geometry analysis showed that upright cultivars achieved target VSP parameters ( 40% cluster exposure) with minimal intervention, while maintaining fruit zone height consistency (coefficient of variation 8% versus 22% for trailing types). Natural shoot spacing was more uniform in upright cultivars (CV 15% versus 28%), contributing to even canopy light distribution.
An Upright_Growth_Disposition is a Shoot_Orientation_Disposition that inheres in Vitis vinifera shoots and is realized through strong negative gravitropism producing shoots that grow predominantly vertically, maintaining angles of less than 20° from vertical throughout their length, with shoot tips ascending even as shoots elongate and gain weight. This disposition represents the upper extreme of shoot orientation, characteristic of cultivars with robust gravitropic response that maintains vertical shoot positioning with minimal or no mechanical support or positioning intervention. Upright-growing cultivars naturally conform to vertical shoot positioning (VSP) training systems, reducing or eliminating the labor requirement for shoot positioning while achieving optimal canopy geometry for light interception and fruit zone exposure. The mechanical basis of Upright_Growth_Disposition involves stiff internodes (high lignification and/or cell wall properties), strong gravitropic sensing and response, and sometimes shorter internodes that reduce the mechanical moment tending to bend shoots. Upright growth creates naturally well-organized canopies with good light penetration, predictable fruit zone location, and compatibility with mechanical harvesting. The disposition is advantageous for VSP systems, pergola, and other vertically-oriented training systems, but may be less suitable for sprawl or GDC systems that benefit from lateral shoot spread.
Upright_Growth_Disposition is_a Shoot_Orientation_Disposition representing vertical shoot growth (< 20° from vertical). Characterized by strong negative gravitropism and stiff internodes. Examples include 'Cabernet Sauvignon', 'Merlot', 'Petit Verdot', 'Tannat', 'Cabernet Franc'. Advantages include VSP compatibility, reduced positioning labor, uniform canopy geometry, and mechanical harvest suitability. The disposition creates naturally organized canopies with good light penetration. High internode stiffness is a contributing mechanical factor.
Aufrechte-Wuchs-Disposition
Stark-Aufrechter-Habitus
Vertikaler-Triebwuchs
erect growth
strongly upright habit
vertical shoot growth
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles, Adelaide Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review, American Journal of Enology and Viticulture 60:251-268 Dry PR (2000) Canopy management for fruitfulness, Australian Journal of Grape and Wine Research 6:109-115
Upright Growth Disposition
Upright growth suitability quality evaluation in Mosel Riesling clones compared natural shoot orientation and VSP management requirements across five clonal selections. Clone Gm 239 demonstrated superior upright growth suitability quality with mean shoot angle of 82° from horizontal at 40 days post-budbreak, requiring only 1.5 positioning passes to achieve 95% catch wire compliance and maintaining excellent vertical canopy organization through harvest with minimal shoot displacement even following late-season storms. Clone Gm 198 showed moderate upright growth suitability quality with mean angle of 68° and requirement for 2.8 positioning passes achieving 88% compliance. Clone Gm 110 exhibited poor upright growth suitability quality with mean angle of only 54° indicating semi-spreading habit, requiring 4.2 positioning passes with persistent catch wire escapes limiting compliance to 76% and motivating recommendation for alternative training approaches. Labor cost analysis demonstrated 45% higher positioning expense for the lowest versus highest upright growth suitability quality clones, establishing this quality as economically significant for VSP vineyard profitability.
Upright growth suitability quality is a growth habit compatibility quality that inheres in a grapevine cultivar or clone and represents the degree to which the genotype exhibits naturally vertical shoot orientation with strong negative gravitropic response that maintains upward growth direction throughout shoot development, conferring high compatibility with vertically shoot positioned training systems and related configurations that depend on upright shoot arrangement to create narrow, well-organized canopy walls with optimal light distribution and simplified management requirements. This quality directly determines the labor efficiency of vertical training systems by establishing whether shoots will self-position upright between catch wires with minimal intervention or require repeated manual positioning to achieve and maintain the vertical orientation essential for these widely-adopted training configurations. The ontological significance of upright growth suitability quality lies in its role as a key selection criterion for cultivars intended for vertical shoot positioned systems that dominate modern premium wine grape production, wherein high upright growth suitability enables the canopy organization and fruit exposure benefits of vertical training to be achieved efficiently while poor suitability can render vertical systems economically impractical due to excessive positioning labor. The physiological basis of upright growth habit involves strong negative gravitropic response in shoot tips that continuously reorients growth toward vertical regardless of shoot origin position, stiff shoot tissue with adequate mechanical strength to support upright growth without bending or lodging, and internode lengths and lateral development patterns that permit shoot placement within catch wire systems without excessive crowding or displacement. Cultivars with high upright growth suitability quality typically exhibit erect shoot tips from early development, maintenance of vertical orientation as shoots elongate, resistance to wind displacement and repositioning toward original vertical alignment following disturbance, and appropriate internode dimensions for catch wire spacing in standard trellis configurations. The assessment of upright growth suitability quality employs observation of natural shoot orientation in freely-growing conditions, quantification of shoot angle distributions, measurement of positioning labor requirements in vertical systems, and evaluation of canopy organization quality achieved with standardized management inputs.
Upright_Growth_Suitability_Quality is_a Growth_Habit_Compatibility_Quality, representing the specific compatibility of vertically-oriented growth habit with vertical shoot positioned and related training systems. This class addresses cultivars exhibiting naturally erect shoot development. Distinguished from Trailing_Growth_Suitability_Quality which addresses the opposite growth orientation. The quantitative metrics of shoot angle and positioning labor requirements provide objective assessment basis. Most relevant for VSP, single curtain, and related vertical training configurations that dominate premium wine grape production.
Aufrechtwuchseignung
Steilwuchsneigung
aufrechte Triebhaltung
erect shoot tendency
negative gravitropism strength
upright shoot orientation
vertical growth habit
vertikale Wuchstendenz
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 32-43 Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review. American Journal of Enology and Viticulture 60:251-268 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 85-112
Upright Growth Suitability Quality
A UV-exposed mature leaf at node 6 on an upper canopy shoot of Vitis vinifera cv. Malbec in Mendoza, Argentina, at 1200 meters elevation receiving very high UV radiation (UV-B fluence rate 1.8 W m⁻² at solar noon compared to 0.8 W m⁻² at sea level), measuring 10.5 cm length by 9.8 cm width, exhibiting pronounced bronze-purple coloration on adaxial surface from high anthocyanin concentrations (2.8 mg g⁻¹ fresh weight compared to 0.6 mg g⁻¹ in low-UV leaves), flavonoid content of 12.5 mg g⁻¹ dry weight in epidermal layers creating UV absorption of 92% of incident UV-B before reaching mesophyll, very thick cuticle of 7.8 μm with substantial epicuticular wax creating slight glaucous appearance and additional UV reflection, epidermal cell walls thickened to 4.5 μm (compared to 2.8 μm in low-UV leaves), palisade layers very compact with minimal intercellular air spaces limiting UV penetration, no visible UV damage symptoms (necrotic spots, bleaching, or malformation), maintained high photosynthetic function with light-saturated rate of 14.5 μmol CO₂ m⁻² s⁻¹ despite high UV stress, measured at BBCH stage 81 (beginning of berry ripening) in February (Southern Hemisphere summer), representing maximum UV protection mechanisms enabling sustained function under extreme UV conditions typical of high-altitude viticulture.
A sun-exposed mature leaf experiencing substantial ultraviolet radiation exposure in Vitis vinifera, characterized by enhanced protective mechanisms against UV damage including elevated concentrations of UV-absorbing compounds particularly flavonoids and hydroxycinnamic acids in epidermal and subepidermal cell layers creating UV screening that absorbs 80-95% of incident UV-B radiation before reaching photosynthetic mesophyll, very thick cuticle with substantial wax deposits reflecting and scattering UV radiation, increased epidermal cell wall thickness providing structural UV protection, elevated anthocyanin accumulation particularly in upper epidermis and palisade cells creating red to bronze pigmentation that absorbs UV and visible light, enhanced DNA repair mechanisms including photolyase enzymes that reverse UV-induced pyrimidine dimer formation, increased antioxidant concentrations including ascorbate, glutathione, and tocopherols neutralizing reactive oxygen species generated by UV stress, and anatomical features including increased palisade layer thickness and decreased intercellular air spaces that limit UV penetration into deep mesophyll layers. UV-exposed mature leaves develop primarily in high-altitude vineyards where thinner atmosphere provides less UV filtration, in low-latitude regions with high solar angles increasing UV intensity, in continental climates with clear skies and low atmospheric moisture reducing UV scattering, and in exterior canopy positions without shading by upper leaves or training structures.
UV-exposed mature leaves are distinguished from other sun leaves by specific adaptations to UV radiation stress, requiring assessment of UV-screening compounds, pigmentation patterns, and anatomical features providing UV protection. UV exposure correlates with but differs from general high light exposure, as UV represents only 3-5% of total solar radiation but has disproportionate damaging effects due to high photon energy. Geographic and altitudinal patterns strongly affect UV exposure, with approximately 10-12% UV increase per 1000 m elevation gain, and low-latitude high-altitude regions experiencing greatest UV stress. Cultivar variation exists in UV tolerance and protective capacity, with some varieties showing enhanced constitutive UV protection while others rely more on induced responses. UV protection comes at metabolic cost through resource allocation to protective compounds and repair mechanisms, potentially reducing resources available for growth and reproduction, but provides essential function in high-UV environments where unprotected leaves would suffer severe photoinhibition and tissue damage.
UV-exponiertes reifes Blatt
UV-protected sun leaf
high UV environment leaf
hochstrahlungs Weinblatt
ultraviolet-adapted leaf
ultraviolett-geschütztes Sonnenblatt
Based on Caldwell et al. (2007) "Terrestrial ecosystems, increased solar ultraviolet radiation, and interactions with other climate change factors" Photochemical & Photobiological Sciences 6=252-266 Kolb et al. (2001) "Effects of natural intensities of visible and ultraviolet radiation on epidermal ultraviolet screening and photosynthesis in grape leaves" Plant Physiology 127=863-875 Berli et al. (2010) "Abscisic acid is involved in the response of grape (Vitis vinifera L.) cv. Malbec leaf tissues to ultraviolet-B radiation by enhancing ultraviolet-absorbing compounds" Australian Journal of Grape and Wine Research 16=395-403 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 249-272.
UV Exposed Mature Leaf
Systematic evaluation of UV_Tolerance_Disposition at the University of Burgundy employed supplemental UV-B exposure (ambient + 8 kJ m⁻² d⁻¹ biologically effective UV-B for 21 days) across 20 cultivars, assessing damage indicators (DNA lesion frequency, lipid peroxidation, PSII efficiency) and protection mechanisms (epidermal UV absorbance, flavonoid content, DNA repair rate). UV_Tolerance_Disposition showed substantial cultivar variation. 'Malbec' (high-altitude Argentinian adaptation) exhibited strong tolerance with epidermal UV-B transmittance of only 8% (effective screening), leaf flavonoid content of 2.8 mg/g FW (quercetin glycosides dominant), DNA lesion frequency 65% lower than sensitive cultivars, and PSII efficiency (Fv/Fm) declining only 5% under UV treatment. 'Chasselas' (temperate Swiss origin) showed poor UV_Tolerance_Disposition with epidermal transmittance of 22%, flavonoid content of 1.1 mg/g, high DNA lesion accumulation, and 18% PSII decline. 'Gewürztraminer' displayed intermediate characteristics. Interestingly, 'Pinot Noir' showed moderate screening capacity but exceptional DNA repair, with cyclobutane pyrimidine dimer (CPD) removal rate of 78% within 4 hours of UV cessation (photoreactivation pathway) versus 45% in 'Chasselas'. Berry UV tolerance partially paralleled leaf tolerance but with distinct patterns—'Malbec' berries showed exceptional anthocyanin stability under UV while 'Pinot Noir' anthocyanins were more UV-labile despite adequate leaf protection. These findings demonstrate that UV_Tolerance_Disposition involves multiple mechanisms that vary independently among cultivars.
A UV_Tolerance_Disposition is an Abiotic_Stress_Tolerance_Disposition that inheres in a Vitis vinifera plant and is realized through physiological mechanisms that protect tissues from damage caused by ultraviolet radiation (UV-A, 315-400 nm UV-B, 280-315 nm) through a combination of UV screening, damage repair, and antioxidant defense. This disposition represents the capacity to tolerate UV exposure, important for viticulture as vineyards are often located in high-UV environments including high-altitude sites, Mediterranean climates with clear skies, and Southern Hemisphere locations with reduced ozone protection. UV radiation causes multiple types of damage including direct DNA lesions (cyclobutane pyrimidine dimers, 6-4 photoproducts), protein damage, lipid peroxidation through reactive oxygen species generation, and photosystem damage affecting photosynthetic function. UV exposure also triggers signaling pathways affecting gene expression, hormone levels, and secondary metabolism, with some UV effects being adaptive (inducing protective responses) rather than purely damaging. UV_Tolerance_Disposition encompasses UV screening capacity (accumulation of UV-absorbing compounds in the epidermis preventing penetration to sensitive underlying tissues), efficient DNA repair mechanisms (photoreactivation and nucleotide excision repair), enhanced antioxidant capacity (neutralizing UV-generated ROS), and tolerance of unavoidable damage at the cellular level. Grapevine shows substantial genotypic variation in UV tolerance, with cultivars adapted to high-altitude or low-latitude origins generally showing enhanced protection compared to those from more temperate regions.
UV_Tolerance_Disposition is_a Abiotic_Stress_Tolerance_Disposition. UV tolerance involves screening (preventing damage), repair (fixing damage), and tolerance (functioning despite damage). Child classes address Flavonoid_Accumulation_Disposition (screening) and DNA_Repair_Disposition (repair). UV exposure is increasing at many vineyard sites due to climate patterns and altitude shifts. Cultivars from high-altitude or low-latitude origins often show enhanced UV protection. UV effects on berry composition (flavonoids, aroma precursors) have quality implications beyond stress tolerance.
Solare-UV-Resistenz
UV-B resistance
UV-Toleranz-Disposition
Ultraviolett-Toleranz
solar UV protection
ultraviolet radiation tolerance
Frohnmeyer H, Staiger D (2003) Ultraviolet-B radiation-mediated stress responses in plants, Trends in Plant Science 8:135-141 Jansen MAK et al. (1998) Higher plants and UV-B radiation: balancing damage, repair and acclimation, Trends in Plant Science 3:131-135 Berli FJ et al. (2010) UV-B impairs growth and gas exchange in grapevines grown in high altitude, Physiologia Plantarum 138:146-155
UV Tolerance Disposition
Molecular and physiological characterization of Vacuolar_Sequestration_Disposition at CSIRO Adelaide combined tonoplast vesicle transport assays with gene expression analysis across 10 rootstocks under salt stress (50 mM NaCl, 21 days). Tonoplast vesicles isolated from root and leaf tissues were assayed for NHX-mediated ²²Na⁺/H⁺ exchange and CLC-mediated ³⁶Cl⁻ transport, while qRT-PCR quantified expression of VvNHX1, VvNHX2, VvCLCc, and VvCLCg. Vacuolar_Sequestration_Disposition showed significant rootstock variation. '1103 Paulsen' exhibited strong sequestration capacity with leaf tonoplast Na⁺/H⁺ exchange rate of 18.5 nmol mg⁻¹ protein min⁻¹ (3-fold higher than salt-sensitive 'Schwarzmann' at 6.2 nmol mg⁻¹ min⁻¹) and correspondingly high VvNHX1 expression (5.2-fold induction under salt versus 1.8-fold in 'Schwarzmann'). Chloride sequestration paralleled sodium patterns, with '1103 Paulsen' showing Cl⁻ transport of 12.4 nmol mg⁻¹ min⁻¹ versus 4.8 in 'Schwarzmann'. X-ray microanalysis confirmed functional significance, with high-sequestration '1103 Paulsen' maintaining cytoplasmic Na⁺ below 35 mM despite vacuolar accumulation to 290 mM, while low-sequestration 'Schwarzmann' showed cytoplasmic Na⁺ of 95 mM with vacuolar 180 mM. Critically, the cytoplasmic K⁺/Na⁺ ratio in '1103 Paulsen' remained at 6.5 (adequate for enzyme function) compared to only 1.8 in 'Schwarzmann' (below critical threshold). These findings demonstrate that Vacuolar_Sequestration_Disposition directly protects cytoplasmic metabolism through NHX and CLC transporter activity.
A Vacuolar_Sequestration_Disposition is an Ion_Compartmentalization_Disposition that inheres in Vitis cells and is realized through active transport of sodium and chloride ions from the cytoplasm into the vacuole, maintaining low cytoplasmic ion concentrations compatible with enzyme function while using accumulated vacuolar ions for osmotic adjustment. This disposition represents intracellular compartmentalization, the most immediate mechanism for protecting cytoplasmic metabolism from ion toxicity. The vacuole, occupying 80-90% of mature cell volume, provides a large capacity sink for ion sequestration, and vacuolar ion accumulation simultaneously contributes to osmotic adjustment by lowering vacuolar water potential. Sodium sequestration is mediated primarily by NHX (Na⁺/H⁺ antiporter) family transporters on the tonoplast (vacuolar membrane), which use the proton gradient generated by V-ATPase and V-PPase to drive sodium into the vacuole against its concentration gradient. Chloride sequestration involves CLC (Chloride Channel) family proteins, particularly those functioning as Cl⁻/H⁺ antiporters on the tonoplast. Effective Vacuolar_Sequestration_Disposition requires coordinated activity of these transporters with the proton pumps that energize them, and maintaining this compartmentalization is an ongoing energy cost as ions tend to leak back to the cytoplasm. Genotypic variation in sequestration capacity reflects differences in transporter expression levels, kinetic properties, and regulation under salt stress.
Vacuolar_Sequestration_Disposition is_a Ion_Compartmentalization_Disposition. Vacuolar sequestration protects cytoplasm while contributing to osmotic adjustment. NHX antiporters (Na⁺/H⁺) mediate sodium sequestration CLC channels/antiporters mediate chloride sequestration. V-ATPase and V-PPase provide energization through proton gradient. Sequestration capacity correlates with transporter expression levels. Maintaining compartmentalization requires ongoing energy expenditure. Cytoplasmic K⁺/Na⁺ ratio is the key functional outcome, with ratios > 1-5 required for enzyme function.
Intrazelluläre-Ionen-Kompartimentierung
Tonoplast-Sequestrierung
Vakuoläre-Sequestrierungs-Disposition
intracellular ion compartmentation
tonoplast sequestration
vacuolar ion storage
Apse MP et al. (1999) Salt tolerance conferred by overexpression of a vacuolar Na⁺/H⁺ antiport in Arabidopsis, Science 285:1256-1258 Bassil E, Blumwald E (2014) The ins and outs of intracellular ion homeostasis: NHX-type cation/H⁺ transporters, Current Opinion in Plant Biology 22:1-6 Teakle NL, Tyerman SD (2010) Mechanisms of Cl⁻ transport contributing to salt tolerance, Plant Cell and Environment 33:566-589
Vacuolar Sequestration Disposition
In a Rheingau Riesling vineyard, vacuolar sugar storage was characterized in late September Year N+1 (BBCH 87) when vacuolar hexose concentration measured 1580 mM combined (glucose 785 mM, fructose 795 mM, ratio 0.99), representing 23.4° Brix, vacuolar volume 2.8 nL per mesocarp cell occupying 96% of cell volume, berry fresh weight 1.65 g having expanded 95% from veraison onset, osmotic potential -2.85 MPa, titratable acidity 7.8 g/L, under cool German conditions (daily high 19-22°C, extended ripening period).
Vacuolar_Sugar_Storage represents the accumulation of glucose and fructose in mesocarp cell vacuoles creating the massive hexose reservoir that determines berry sweetness, defines wine alcohol potential, and provides osmotic driving force for berry expansion during ripening. This storage process intensifies during veraison through ripening (BBCH 83-89), beginning 2-5 days post-transport activation and continuing until harvest, and is characterized by vacuolar hexose concentrations increasing from pre-veraison 100-250 mM to veraison onset 400-700 mM to peak ripening 1400-2200 mM representing 8-12 fold enhancement, total berry sugar content rising from 12-25 g/L to 200-280 g/L (20-28° Brix), and vacuolar volume fraction expanding from 75-85% to 95-97% of mesocarp cell volume creating storage capacity 1.2-3.5 nL per cell accommodating this massive hexose accumulation. Molecular basis of vacuolar storage includes: tonoplast hexose transporters (possibly including VvHT isoforms localized to tonoplast) facilitating glucose and fructose transport from cytoplasm (60-100 mM) into vacuole (accumulating to 700-1100 mM each sugar, combined 1400-2200 mM), transport likely predominantly facilitated diffusion down concentration gradients maintained by continued plasma membrane import and metabolic consumption in cytoplasm preventing equilibration, potential role for H⁺-coupled secondary active transport though evidence suggests primarily facilitated mechanism, and vacuolar membrane integrity essential for maintaining compartmentation preventing hexose leakage back to cytoplasm. Vacuolar biophysics supporting sugar storage demonstrates: osmotic potential declining from -0.6 to -1.2 MPa (pre-veraison) to -2.4 to -3.6 MPa (peak storage) through hexose accumulation, turgor pressure maintained 0.3-0.6 MPa (compared to 0.6-1.0 MPa green berry) through balance of osmotic potential and cell wall mechanical resistance creating positive pressure sustaining cellular structure, water potential following osmotic potential declining from -0.4 to -0.8 MPa to -2.0 to -3.0 MPa creating strong gradient favoring water import from vine xylem (-0.5 to -1.2 MPa typical) driving osmotic berry expansion, and vacuolar pH maintained 2.8-3.6 throughout storage providing acidic environment important for organic acid balance, anthocyanin stability (in reds), and potential enzyme regulation. Glucose and fructose storage characteristics show: both hexoses accumulating approximately equimolar with glucose:fructose ratio 0.95-1.05 in vacuoles despite potential differential metabolism in cytoplasm, final concentrations typically 700-1100 mM each (combined 1400-2200 mM total hexoses) representing 25-40% w/v solution, spatial distribution relatively uniform within individual vacuoles though may show gradients across tissue with highest concentrations in central mesocarp declining toward periphery, and temporal accumulation following sigmoid pattern with slow initial phase, rapid exponential phase 3.0-5.0% Brix per week, and decelerating plateau 32°C impairs to 2.0-3.2% per week and cool <20°C reduces to 2.2-3.2% per week, water availability with adequate vine water status enabling osmotic berry expansion supporting storage while severe deficit (<-1.6 MPa) limiting expansion potentially constraining total storage capacity though concentration may increase, light operating through source photosynthesis providing sucrose substrate with adequate canopy illumination essential for sugar supply, and nutrient status particularly nitrogen and potassium affecting metabolic capacity for supporting storage processes. Cultivar genetic variation in storage capacity shows: high-Brix varieties (Gewürztraminer, Sémillon potential 26-30° Brix) achieving vacuolar concentrations 1800-2400 mM combined hexoses through enhanced transport and potentially larger vacuoles, moderate cultivars (Chardonnay, Cabernet Sauvignon 22-26° Brix) reaching 1400-1900 mM, and lower-sugar types (Pinot Noir cool climates 20-24° Brix) attaining 1200-1600 mM reflecting genetic differences in maximum storage capacity and accumulation duration. Storage stability and harvest window considerations show: vacuolar sugar relatively stable once accumulated with <5-10% losses over 2-4 weeks post-peak enabling flexible harvest timing in stable cultivars, environmental stresses (heat, water deficit extremes) potentially destabilizing storage causing 10-30% sugar losses through respiratory consumption or cellular disruption, and cultivar variation with stable types (Cabernet Sauvignon, Syrah) maintaining storage 4-6 weeks while unstable varieties (Pinot Noir warm climates) may lose 15-35% within 2-3 weeks post-peak requiring prompt harvest. Quality implications of vacuolar storage inclu final storage level determining wine alcohol potential with each 1° Brix approximately yielding 0.55-0.60% ethanol post-fermentation, sugar-acid balance critically affecting wine style with ratio 20-35:1 (Brix:titratable acidity g/L) typical for quality wines, uniformity of storage across berries and clusters determining harvest maturity consistency, and coordination of sugar storage with phenolic and flavor development defining optimal harvest window balancing multiple quality parameters. Physiological limits to storage inclu maximum vacuolar hexose concentrations ~2200-2600 mM approaching saturation potentially limiting further accumulation, membrane integrity possibly compromised at extreme osmotic potentials <-3.5 MPa, cell wall mechanical limits potentially constraining vacuolar expansion beyond certain size, and metabolic regulation through feedback mechanisms limiting accumulation as homeostatic set-points approached preventing excessive storage. Viticultural management targeting optimal storage involves: maintaining adequate vine water status supporting osmotic expansion without dilution from excess water, ensuring source photosynthesis through canopy health providing sucrose substrate, optimizing crop load 6-12 kg per vine variety-dependent enabling adequate storage without source limitation, and timing harvest to capture peak storage before degradation in unstable varieties or allowing extended hang time in stable cultivars for flavor development.
Vacuolar_Sugar_Storage is distinguished from Sugar_Uptake_Enhancement by representing compartmentalization rather than import, from Hexose_Transport_Initiation by focusing on storage rather than membrane transport, and from sugar accumulation onset by showing progressive storage (100-250 → 1400-2200 mM) rather than metabolic transition. Duration throughout ripening BBCH 83-89. Vacuolar volume fraction 95-97% at peak. Osmotic potential -2.4 to -3.6 MPa. Glucose:fructose ratio ~1.0. Vacuolar_Sugar_Storage represents culminating compartmentalization determining berry sweetness and wine alcohol potential.
Glukose-Fruktose-Sequestrierung
Hexose-vakuoläre Akkumulation
Tonoplast-vermittelte Lagerung
Zucker-Kompartimentierung
glucose-fructose sequestration
hexose vacuolar accumulation
sugar compartmentalization
tonoplast-mediated storage
Keller M (2020) The Science of Grapevines 3rd Ed pages 221-228 vacuolar sugar storage Fontes N, Gerós H, Delrot S (2011) Grape berry vacuole:a complex and heterogeneous membrane system specialized in the accumulation of solutes Amer J Enol Vitic 62:270-278 vacuolar storage characteristics Wada H, Matthews MA, Shackel KA (2008) Seasonal pattern of apoplastic solute accumulation and loss of cell turgor during ripening of Vitis vinifera fruit under field conditions Amer J Enol Vitic 59:173-180 storage dynamics and cell physiology.
Vacuolar Sugar Storage
Pinot Noir in Burgundy shows vacuole enlargement initiation during late July-early August Year N+1 at BBCH 73-75, with vacuoles expanding from 18% to 62% cell volume, tonoplast membrane synthesis, malate accumulation to 68 mM and tartrate to 42 mM, vacuole pH 3.2, osmotic potential -0.9 MPa, progressive fusion creating 1-2 large vacuoles per cell, under continental conditions of 22-27°C temperature, typical of this early-to-mid season cultivar developing vacuolar system
Vacuole enlargement initiation represents the osmotic compartment development phase during initial cell enlargement when small vacuoles in post-division cells expand through water influx and membrane synthesis, multiple vacuoles coalesce into fewer larger compartments, progressive vacuole growth from 10-30% to 50-70% cell volume creates dominant storage organelle, and vacuole establishes as major osmotic engine and solute storage depot driving cell expansion. This vacuolar development phase occurs during mid-late summer in mid-late July to early August in temperate regions Year N+1, proceeding over weeks 3-5 post-fertilization during initial enlargement, corresponding to BBCH 73-75 pea-sized to berry touch stages. Vacuole initiation mechanism involves small vacuoles 10-30% volume in post-division cells beginning expansion, tonoplast membrane synthesis accommodating growth, water influx into vacuoles driven by solute accumulation, vacuole fusion events creating larger compartments, progressive enlargement to 50-70% cell volume by initiation phase completion, and vacuole as expanding osmotic and storage compartment. Post-division vacuole characteristics show cells exiting division containing multiple small vacuoles 2-5 micrometers diameter, vacuoles derived from provacuoles or ER-originated compartments, total vacuole fraction 10-30% cell volume, and small vacuoles as starting point for enlargement. Vacuole expansion mechanisms show tonoplast membrane synthesis increasing vacuole surface area, lipid and protein synthesis for membrane expansion, water influx driven by accumulating solutes increasing vacuole volume, membrane synthesis coordinated with water uptake enabling growth, and integrated processes driving vacuole enlargement. Tonoplast membrane synthesis shows phospholipid biosynthesis producing membrane lipids, membrane protein synthesis including H+-ATPase proton pumps, transporter proteins enabling solute accumulation, aquaporin water channels facilitating water influx, and coordinated synthesis accommodating vacuole expansion. Tonoplast H+-ATPase function shows proton pump acidifying vacuole interior, ATP-dependent H+ transport creating pH gradient 2-3 units across tonoplast, acidification enabling secondary active transport of solutes, and pump as key regulator of vacuole function. Solute accumulation into vacuole shows organic acids particularly malate transported into vacuole, tartrate synthesis and accumulation in vacuole, potassium import balancing organic acid anions, sugar beginning accumulation though limited during initial phase, and solute accumulation creating osmotic potential driving water influx. Malate accumulation shows malic acid as primary accumulated organic acid during initiation, malate transporter-mediated import from cytoplasm to vacuole, vacuolar malate concentration reaching 50-100 mM, and malate as major osmolyte and contributor to acidity. Tartrate accumulation shows tartaric acid synthesis from ascorbate pathway in cytoplasm, tartrate transport into vacuole, tartrate relatively stable once accumulated, concentration 30-60 mM in vacuole, and tartrate as characteristic grape organic acid. Potassium accumulation shows K+ import through tonoplast K+/H+ antiporter, potassium concentration 80-150 mM in vacuole, K+ as major cation neutralizing organic acid negative charges, and potassium contributing to osmotic potential. Water influx into vacuole shows osmotic gradient from accumulated solutes driving water movement, water crossing tonoplast through aquaporin channels, water influx increasing vacuole volume, and water as primary volume contributor. Vacuole osmotic potential shows accumulated solutes creating highly negative osmotic potential -0.8 to -1.2 MPa within vacuole, osmotic potential more negative than cytoplasm driving water from cytoplasm into vacuole, and vacuole as osmotic engine driving cell water uptake and expansion. Vacuole volume increase shows typical expansion from 10-30% initial to 50-70% cell volume by initiation completion representing 3-7 fold vacuole volume increase, progressive enlargement paralleling cell expansion, and vacuole volume increase as major cell volume contributor. Vacuole fusion events show multiple small vacuoles fusing into fewer larger vacuoles, fusion mediated by SNARE proteins and fusion machinery, progressive fusion creating single or few large vacuoles per cell, and fusion consolidating vacuolar compartment. Vacuole number reduction shows cells transitioning from multiple small vacuoles to typically 1-3 larger vacuoles, fusion reducing compartment number while increasing individual size, and eventual dominance of single large central vacuole in mature cells. Vacuole as storage compartment shows vacuole storing accumulated organic acids, ions, later sugars during ripening, water comprising 85-90% vacuole mass, and vacuole as major storage depot. Vacuole pH characteristics show acidic pH 3.0-3.5 from organic acid accumulation and proton pump activity, pH maintained through continued pumping, acidic pH affecting enzyme activities and solute partitioning, and vacuole acidity as characteristic feature. Vacuole-cytoplasm relationships show vacuole occupying increasing cell volume fraction leaving cytoplasm as thin peripheral layer, cytoplasm maintaining metabolic functions in reduced volume, and spatial organization with vacuole central and cytoplasm peripheral. Cell volume increase from vacuole expansion shows vacuole enlargement contributing substantially to cell volume increase, typical 60-80% cell volume increase attributable to vacuole during initiation, and vacuole as dominant volume contributor. Environmental effects on vacuole initiation include temperature affecting membrane synthesis and solute accumulation rate, water availability affecting water influx and vacuole expansion, nutrient supply affecting solute accumulation capacity, and overall favorable conditions supporting vacuole development. Water effects show adequate water essential for vacuole expansion through water influx, water deficit limiting vacuole growth potentially reducing cell expansion, and water as critical resource. Temperature effects show warm conditions promoting faster membrane synthesis and vacuole development, cool conditions slowing initiation though not preventing, and temperature affecting development rate. Vacuole initiation assessment methods include transmission electron microscopy showing vacuole structure and size, confocal microscopy with vacuole-selective dyes, vacuole isolation and compositional analysis, and morphometric analysis quantifying vacuole fraction. The process represents second aspect of initial cell enlargement establishing osmotic compartment, following turgor-driven expansion onset and preceding cell wall loosening. Duration weeks 3-5 representing progressive vacuole development during initial enlargement. Regional timing shows vacuole enlargement initiation during mid-late July to early August across climates during initial enlargement. Viticultural significance includes representing development of major storage compartment, establishing osmotic engine driving expansion, demonstrating water and nutrient accumulation system, and creating compartment for subsequent sugar accumulation during ripening. Research applications include analyzing vacuole biogenesis and expansion mechanisms, investigating tonoplast transport and regulation, evaluating solute accumulation dynamics, quantifying vacuole contribution to cell volume, and assessing vacuole development as determinant of expansion capacity.
Vacuole_Enlargement_Initiation is second aspect of Initial_Cell_Enlargement, distinguished by osmotic compartment development. Represents expansion from 10-30% to 50-70% cell volume. Tonoplast synthesis, solute accumulation (malate, tartrate, K+), water influx. Osmotic potential -0.8 to -1.2 MPa. Vacuole fusion reducing number. pH 3.0-3.5. Contributes 60-80% cell volume increase. Duration weeks 3-5. Follows Turgor_Driven_Expansion_Onset. Precedes Cell_Wall_Loosening. Establishes osmotic engine and storage depot.
Vakuolen-Entwicklung
Vakuolen-Vergrößerungs-Initiierung
osmotic compartment formation
osmotisches Kompartment-Bildung
vacuolar expansion onset
vacuole development
vacuole growth initiation
vakuoläre Expansions-Beginn
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 444-447 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Vacuole Enlargement Initiation
Vapor pressure deficit response characterization comparing Mediterranean (Monastrell) and Atlantic (Albariño) cultivars reveals climate-adapted VPD sensitivities, with Monastrell showing high VPD sensitivity (m = 145 mmol/m²/s per ln(kPa), steep conductance decline protecting water status under high evaporative demand typical of hot inland climates) while Albariño shows low sensitivity (m = 65, maintaining conductance under VPD conditions rarely experienced in cool coastal origin), demonstrating how VPD response reflects adaptation to regional atmospheric conditions and informs cultivar selection for different viticultural climates.
Vapor pressure deficit response quality is a transpiration rate quality that represents the functional relationship between vine gas exchange and atmospheric vapor pressure deficit, characterizing how stomatal conductance, transpiration, and photosynthesis change as the driving force for evaporation varies with humidity conditions that differ among regions, through seasons, and within each day. Within the BFO framework, this quality addresses the atmospheric demand dimension of gas exchange regulation, distinguishing the VPD response from responses to other environmental factors while recognizing that VPD is a primary determinant of both transpiration rate and stomatal behavior, creating feedback loops between atmospheric conditions and vine water use that fundamentally shape productivity and water relations. The ontological significance lies in VPD response determining vine performance across the range of atmospheric humidity conditions experienced in viticulture, where the sensitivity of stomatal conductance to VPD establishes how effectively vines limit water loss under high evaporative demand, with implications for productivity under hot, dry conditions and for understanding regional differences in vine water use patterns. The physiological basis for VPD response involves both direct effects on transpiration (evaporative demand proportional to VPD at constant conductance) and indirect effects through stomatal regulation (stomata typically closing at high VPD through hydropassive and hydroactive mechanisms), with the net effect being that transpiration increases with VPD but often less than proportionally due to concurrent stomatal closure that partially compensates for increased driving force. The quantitative characterization of VPD response typically employs the relationship between stomatal conductance and VPD, often modeled as gs = gsref - m × ln(VPD) where gsref is reference conductance and m is the sensitivity parameter, with higher m indicating more pronounced stomatal closure at elevated VPD and more conservative water use. The cultivar variation in VPD response contributes to the isohydric-anisohydric spectrum, with isohydric cultivars showing strong VPD response (steep conductance decline maintaining water status) while anisohydric types show weaker response (continued transpiration allowing productivity but risking hydraulic damage). Understanding vapor pressure deficit response quality enables prediction of vine behavior under varying humidity conditions, cultivar matching to regional climates, and interpretation of diurnal and seasonal patterns in vine water use driven by atmospheric demand variation.
Vapor_Pressure_Deficit_Response_Quality addresses the functional relationship between gas exchange and atmospheric humidity/VPD. This quality determines vine behavior under varying evaporative demand conditions. The class supports climate adaptation assessment and cultivar-region matching through VPD response characterization.
Luftfeuchtigkeitsreaktion
VPD sensitivity
VPD-Antwort
VPD-Sensitivität
atmospheric demand response
atmosphärische Bedarfsreaktion
evaporative demand sensitivity
humidity response
Oren R et al. (1999) Survey and synthesis of intra- and interspecific variation in stomatal sensitivity to vapour pressure deficit. Plant, Cell & Environment 22(12): 1515-1526 Rogiers SY et al. (2012) Stomatal response of an anisohydric grapevine cultivar to evaporative demand, available soil moisture and abscisic acid. Tree Physiology 32(3): 249-261
Vapor Pressure Deficit Response Quality
Variegated coloration in Pinot Grigio berries represents a cultivar-specific characteristic where genetic mosaicism produces irregular sectors of pink-grey skin interspersed with green-yellow zones on individual berries, creating the distinctive appearance that gives this Pinot Noir mutation its name (grigio meaning grey) this inherent variegation, unlike the defect-associated variegation of other cultivars, does not indicate ripening problems and produces the copper-pink tinted wines characteristic of traditional Italian Pinot Grigio production styles from Alto Adige and Friuli.
Variegated_Coloration_Quality represents the color distribution condition where discrete zones, patches, or sectors of contrasting color appear across the surface of a grapevine organ, creating a discontinuous pattern of color variation. Variegation is distinguished from gradient coloration by the presence of relatively sharp boundaries between differently colored zones rather than continuous transitions. In grape berries, variegated coloration typically manifests as green or pale patches on otherwise colored fruit, resulting from localized failure of anthocyanin biosynthesis in specific skin regions. Contact zones between adjacent berries in compact clusters commonly display reduced coloration, creating variegated patterns when these pale areas contrast with the fully colored exposed surfaces. Sunburn damage can produce variegated patterns with necrotic brown patches on otherwise normal fruit. Certain viral infections, particularly leafroll virus, cause variegated leaf coloration with interveinal reddening while vein-adjacent tissue remains green, a diagnostic symptom pattern. In leaves, variegation may result from chimeral tissue structure where genetically distinct cell layers produce different pigmentation, from viral infection causing mosaic patterns, or from mineral deficiency creating interveinal chlorosis with green veins. The spatial scale of variegation ranges from fine speckling with millimeter-scale patches to broad sectoring with distinct organ regions showing different colors. Assessment of variegation documents the pattern type (spotted, sectored, mosaic), the size and shape of contrasting zones, and the proportion of organ surface displaying each color category.
Variegated_Coloration_Quality encompasses multiple pattern types unified by the presence of discrete contrasting color zones. The distinction between variegation (discrete patches) and gradient (continuous transition) may be ambiguous when transition zones exist between patches variegation is diagnosed when distinct patch boundaries are visible. Variegation may be a stable cultivar character (as in some ornamental or chimeral varieties), a symptom of disease or disorder, or a consequence of environmental heterogeneity during development. The underlying cause of variegation affects interpretation chimeral variegation is stable while environment-induced variegation may vary among organs and seasons.
discontinuous pigmentation
fleckige Farbverteilung
gefleckte Pigmentierung
gescheckte Färbung
mottled color
panaschierte Färbung
patchy coloration
sectored coloration
Boss PK and Thomas MR (2002) Association of Dwarfism and Floral Induction with a Grape 'Green Revolution' Mutation. Nature 416(6883): 847-850 Guidoni S et al. (2008) Allelic Variation in the VvMYBA1 and VvMYBA2 Genes and the Effect of Berry Color Variation in Cultivars of Vitis vinifera L. Australian Journal of Grape and Wine Research 14(2): 115-123
Variegated Coloration Quality
Varietal aroma development assessment in cool versus warm site Syrah reveals the terroir effect on varietal expression, with cool site showing moderate rotundone (150 ng/L, expressing the black pepper character distinctive to Syrah), preserved violet-contributing compounds, and restrained fruit intensity (varietal character emphasizing spice and floral aspects), while warm site shows reduced rotundone (40 ng/L, minimal pepper character), enhanced fruity esters from higher sugar at harvest, and dominant dark fruit character (varietal expression shifted toward jammy rather than peppery typicity), demonstrating how varietal aroma development produces site-specific expressions of cultivar identity that contribute to regional wine character.
Varietal aroma development quality is an aromatic maturity quality that represents the degree to which cultivar-characteristic aroma compounds have accumulated and achieved their typical expression profile, characterizing the development of the signature aromatic character that distinguishes grape varieties and defines the varietal typicity expected in wines, from the grassy character of Sauvignon Blanc to the floral notes of Gewürztraminer to the cassis of Cabernet Sauvignon. Within the BFO framework, this quality specifically addresses the varietal expression dimension of aromatic maturity, distinguishing the development of cultivar-specific character from generic aroma compound accumulation while recognizing that varietal typicity arises from specific compounds or compound ratios that vary among cultivars and must reach appropriate levels and proportions for authentic varietal expression. The ontological significance lies in varietal aroma development determining the authenticity and quality of varietal wine character, where each cultivar possesses signature compounds (or compound combinations) that must achieve sufficient concentration and appropriate ratios for the wine to express recognizable varietal character, with insufficient development resulting in neutral wines lacking varietal identity and overdevelopment potentially producing atypical or unbalanced aromatic profiles. The cultivar-specific compounds defining varietal character include monoterpenes for aromatic varieties (linalool and geraniol dominant in Muscat, α-terpineol in Gewürztraminer), methoxypyrazines for herbaceous varieties (IBMP providing green bell pepper in Sauvignon Blanc and Cabernet family), rotundone for spicy varieties (black pepper character in Syrah), volatile thiols for tropical varieties (4MMP and 3MH providing tropical fruit in Sauvignon Blanc), and damascenone contributing complexity across many varieties, with each compound showing variety-specific accumulation patterns. The environmental modulation of varietal aroma development shows substantial site and vintage effects on the expression of varietal character, with cool climates often favoring higher methoxypyrazine retention (more herbaceous character) while warm climates favor greater methoxypyrazine degradation and potentially enhanced thiol precursor accumulation (more tropical character in Sauvignon Blanc), demonstrating how the same variety expresses different aspects of its aromatic potential depending on growing conditions. The harvest timing effects on varietal character involve the progressive evolution of aroma profiles during ripening, where early harvest typically expresses green/herbaceous aspects of varietal character while later harvest shifts toward riper fruit expression, with optimal varietal typicity achieved at maturity levels that may vary by wine style objectives. Understanding varietal aroma development quality enables assessment of varietal expression potential, harvest timing for target varietal character, and recognition of the environmental and maturity factors affecting cultivar-specific aromatic profiles.
Varietal_Aroma_Development_Quality addresses the accumulation of cultivar-specific compounds determining varietal typicity. This quality involves signature compounds that must reach appropriate levels for authentic varietal expression. The class supports varietal wine quality assessment through cultivar-specific aroma development characterization.
Rebsortencharakter
Sortenaromenentwicklung
Sortenausdruck
cultivar aroma expression
signature aroma accumulation
varietal character development
Siebert TE et al. (2008) Development of a stable isotope dilution analysis for the quantification of sesquiterpene rotundone in red and white wines. Journal of Agricultural and Food Chemistry 56(10): 3745-3748 Dunlevy JD et al. (2009) A methyltransferase essential for the methoxypyrazine-derived flavour of wine. The Plant Journal 59(3): 404-412
Varietal Aroma Development Quality
Varietal authorization for Blaufränkisch in Austria classifies the cultivar as "Qualitätsrebsorte" (quality grape variety) under Annex 1 of the Austrian Weingesetz, authorized for quality wine production in all nine Austrian wine-growing regions (Niederösterreich, Burgenland, Steiermark, Wien, and their sub-regions), with specific DAC authorization in Mittelburgenland (as the exclusive red variety), Leithaberg (as a permitted variety alongside Pinot Noir), and Eisenberg (as the principal red variety) — the classification reflecting the cultivar's established position in Austrian viticulture and its particular significance for Burgenland red wine identity. Varietal authorization for Cabernet Sauvignon in Italy documents a progressive expansion — initially classified as an authorized non-traditional variety ("vitigno non tradizionale") in Tuscany following the experimental plantings of the 1970s-1980s (Sassicaia, Ornellaia), subsequently upgraded to recommended status in multiple regions, and now authorized as a permitted blending component in numerous DOC and DOCG regulations including Bolgheri DOC (where it may constitute up to 100%) and Chianti Classico DOCG (where it is permitted at up to 20%).
Varietal authorization is a regulatory information entity that documents the legal permission, classification, and conditions under which specific grape cultivars are authorized for commercial cultivation and wine production within defined administrative jurisdictions, encompassing the classification status of the cultivar within the national or regional variety catalogue (authorized, recommended, temporarily authorized, experimental, prohibited), the geographic scope of the authorization (nationwide, regional, appellation-specific), the product categories for which the cultivar is authorized (quality wine, geographical indication wine, varietal wine, table wine, rootstock use), and the conditions or restrictions attached to the authorization (maximum permitted yield, minimum planting density, mandatory training system), providing the primary regulatory gateway that determines which cultivars may legally be grown for commercial wine production in each viticultural jurisdiction.
Varietal authorization represents the intersection of viticultural science, agricultural policy, and cultural heritage protection — the decision to authorize or restrict specific cultivars in defined regions reflects competing policy objectives including quality assurance (restricting appellations to historically proven cultivars), agricultural modernization (permitting new cultivars with improved agronomic characteristics), climate adaptation (authorizing cultivars suited to changing climatic conditions), and cultural identity preservation (maintaining the traditional cultivar portfolios that define regional wine character). The EU framework for varietal authorization has evolved substantially — the pre-2008 system classified cultivars as "recommended" (incentivized for planting), "authorized" (permitted), or "temporarily authorized" (tolerated pending review) within each administrative region, while the post-2008 system simplified classification to a single authorized/non-authorized distinction at the member state level, with more specific restrictions delegated to appellation-level regulations. The practical consequence is that varietal authorization now operates at two levels — the national variety catalogue (listing all cultivars permitted for commercial wine production within the country) and the appellation specification (cahier des charges / Produktspezifikation) listing the cultivars permitted for wines carrying a specific PDO or PGI designation, with the latter typically being far more restrictive than the former. The climate change dimension of varietal authorization is increasingly prominent — multiple European appellations are undertaking experimental authorization of heat-tolerant and drought-resistant cultivars not traditionally grown in their regions (Bordeaux's 2021 authorization of six new varieties including Touriga Nacional and Marselan for experimental blending, INAO's progressive approach to "cépages d'adaptation"), creating tension between innovation and tradition in the authorization framework. The non-European approach to varietal authorization is fundamentally different — most New World wine jurisdictions (USA, Australia, Chile, Argentina, New Zealand, South Africa) do not restrict which Vitis vinifera cultivars may be planted, leaving varietal selection entirely to market forces and grower judgment, though appellation-specific rules may require certain cultivars for origin-labeled wines.
Rebsortenklassifikation
Sortengenehmigung
Sortenregistrierung
Sortenzulassung
cultivar authorization
cultivar classification status
varietal registration
variety approval
European Parliament (2013) Regulation (EU) No 1308/2013, Articles 81-110 Österreichisches Weingesetz (2009) §3 and Annex 1 (Qualitätsrebsorten) INAO (2021) Cahier des charges des appellations d'origine contrôlées van Leeuwen C et al. (2019) An update on the impact of climate change in viticulture and potential adaptations. Agronomy 9(9): 514
Varietal Authorization
Varietal labeling rule for EU wines requires a minimum of 85% of the named cultivar in the wine's composition for single-variety labeling (Article 50 of Regulation 2019/33), with the remaining 15% permitted from any other authorized variety — thus a wine labeled "Grüner Veltliner" must contain at least 85% Grüner Veltliner, though many Austrian producers use 100% for premium bottlings. When two cultivars are named (e.g., "Grüner Veltliner - Riesling"), both must be present and their combined total must reach 100%, listed in descending order of proportion, with the smaller component constituting at least 15%. Varietal labeling rule for US wines requires a minimum of 75% of the named cultivar for wines carrying a non-AVA appellation, and 75% for AVA-designated wines (27 CFR §4.23) — a notably lower threshold than the EU's 85%, meaning that a US wine labeled "Cabernet Sauvignon" may contain up to 25% of other varieties without disclosure. Oregon law is a notable exception within the US, requiring 90% minimum for most varietal wines (ORS 471.445), reflecting a deliberate policy of enhanced consumer transparency.
Varietal labeling rule is a label regulation entity that documents the legal conditions under which a grape cultivar name may appear on a wine label, specifying the minimum percentage of the named cultivar required in the wine's composition, the regulatory framework establishing this threshold, any exceptions or derogations for specific cultivar names or wine categories, the rules governing the declaration of multiple cultivars on a single label, and the relationship between varietal labeling and other label elements (appellation, quality tier, vintage), providing the regulatory framework that determines when and how cultivar identity — one of the most commercially significant label claims — may be communicated to the consumer.
Varietal labeling rules encode a fundamental tension between consumer information (the expectation that a wine labeled with a cultivar name predominantly consists of that cultivar) and winemaking flexibility (the traditional practice of blending small proportions of complementary varieties to enhance complexity, balance, or consistency). The 85% threshold adopted by the EU represents a compromise between the 100% purity that consumers might expect and the practical reality that many acclaimed wines traditionally incorporate small proportions of complementary cultivars — white Hermitage (Northern Rhône Syrah with up to 15% Marsanne/Roussanne) and Chianti Classico (predominantly Sangiovese with permitted additions of Canaiolo, Colorino, Cabernet, Merlot) would be precluded from varietal labeling under a 100% rule. The cross-jurisdictional threshold variation — 85% in the EU, 75% in the US (except Oregon at 90%), 85% in Australia and New Zealand, 75% in Chile and Argentina, 75% in South Africa — creates both consumer confusion and trade compliance complexity for internationally marketed wines, as the same blend proportions may qualify for varietal labeling in one jurisdiction but not another. The cultivar name protection dimension intersects with varietal labeling — certain cultivar names that are also geographic terms (Champagne, Burgundy, Chablis, Tokay) carry additional restrictions or have been the subject of international trade negotiations (the EU-US wine accord addressing "semi-generic" American use of European geographic terms as varietal or type designations). The interaction between varietal labeling and appellation requirements creates additional complexity — some appellations mandate varietal labeling (Weinviertel DAC requires declaration as Grüner Veltliner), some prohibit it (Châteauneuf-du-Pape traditionally uses only the appellation name without varietal specification), and some permit it optionally (many Italian DOCs allow but do not require varietal identification on the label).
Rebsortenangabenregel
Rebsortenbezeichnungsrecht
Sortenbezeichnungsregel
Sortendeklarationsvorschrift
Sortennennungsregel
cultivar labeling rule
grape variety labeling
varietal declaration rule
varietal percentage rule
variety naming rule
European Parliament (2019) Commission Delegated Regulation (EU) 2019/33, Articles 49-53 TTB (2023) 27 CFR §4.23 (varietal labeling) Oregon Revised Statutes §471.445 Wine Australia (2023) Label Integrity Program requirements
Varietal Labeling Rule
Varietal typicity assessment in international Pinot Noir competition applies variety-specific criteria including red fruit character (cherry, raspberry) dominance over dark fruit, earthy-forest floor notes present, absence of jammy or pruney character, and overall elegance rather than power, with wines from warm regions showing dark fruit and extraction rated as atypical despite technical quality, demonstrating how varietal typicity standards may conflict with climate influence and stylistic preference, creating tension between global variety definitions and legitimate regional expressions of the same variety.
Varietal typicity quality is an aroma typicity quality that represents the degree to which a wine's aromatic profile expresses the characteristic signature of its declared grape variety, characterizing the recognizability of variety-specific aromatic markers that enable variety identification and confirm varietal authenticity. Within the BFO framework, this quality specifically addresses conformity to varietal aromatic expectations, distinguishing variety-specific typicity from regional or stylistic typicity while recognizing that varietal character represents the genetic contribution to aromatic profile that persists across diverse growing conditions and winemaking approaches. The ontological significance lies in varietal typicity providing the primary basis for variety identification and varietal wine marketing, where typical varietal expression justifies variety labeling, commands variety-specific price premiums, and fulfills consumer expectations created by varietal wine communication, making varietal typicity a commercial necessity for varietally-labeled wines. The compositional basis for varietal typicity includes variety-specific impact compounds (methoxypyrazines defining Sauvignon Blanc herbaceous character, rotundone providing Syrah pepper, terpenes creating Muscat floral signature), variety-characteristic compound ratios (ester profiles distinguishing Pinot Noir from Cabernet Sauvignon fruitiness), and the overall aromatic gestalt that experienced tasters associate with specific varieties. The variety-specificity of aromatic markers varies substantially, with highly distinctive varieties (Gewürztraminer, Sauvignon Blanc, Muscat) showing unmistakable aromatic signatures while less distinctive varieties (Chardonnay, Merlot) express variety more through structure and subtle aromatic nuance than through striking impact compounds. The winemaking influence on varietal expression creates complexity, as certain practices (oak aging, malolactic fermentation, extended maceration) may either enhance varietal character or mask it beneath winemaking-derived aromas, creating wines that taste more of process than of grape. Understanding varietal typicity quality enables variety identification and authentication, marketing claim verification, and winemaking optimization for varietal expression versus processing character.
Varietal_Typicity_Quality specifically addresses conformity to grape variety aromatic expectations, representing the genetic contribution to wine character. This quality requires specification of the reference variety and acknowledgment that typicity standards may vary among markets, critics, and evaluation contexts. The class supports variety authentication while raising questions about legitimate stylistic diversity within variety categories.
cépage typicity
grape variety expression
varietal authenticity
varietal character
varietale Authentizität
Swiegers JH et al. (2005) Yeast and bacterial modulation of wine aroma and flavour. Australian Journal of Grape and Wine Research 11(2): 139-173 Parr WV et al. (2007) The distinctiveness of Sauvignon Blanc wine. Food Quality and Preference 18(6): 849-861
Varietal Typicity Quality
Cabernet Sauvignon vascular bundle mesocarp showing typical organization= 4 major dorsal bundles (positioned at approximately 90° intervals around berry circumference, each 150-280 μm diameter) running from pedicel attachment through mesocarp to stylar scar, with xylem tissue 60-120 μm diameter (vessels 28-45 μm diameter, non-functional at maturity with cavitated air-filled vessels) and phloem tissue 40-95 μm diameter (sieve elements 15-25 μm diameter, companion cells 8-15 μm, functional during ripening), plus approximately 45-85 secondary lateral bundles (30-95 μm diameter) branching from major bundles creating distribution network throughout mesocarp representing total vascular bundle mesocarp <2.5% of mesocarp volume but providing import pathway supporting entire berry composition development. Merlot vascular bundle mesocarp phloem import during ripening= phloem sieve elements conducting sucrose-rich sap (12-18% sucrose concentration in phloem sap) from source leaves through pedicel into berry vascular bundles, with sucrose unloading into vascular bundle mesocarp and surrounding tissues, invertase enzymes converting sucrose to glucose and fructose for storage in mesocarp cell vacuoles, creating progressive sugar accumulation from 14% at veraison to 24.5% at maturity through continuous phloem import and distribution via vascular bundle mesocarp network. Syrah vascular bundle mesocarp xylem-to-phloem transition at veraison= pre-veraison (BBCH 75-79, weeks 6-9) showing functional xylem conducting water and minerals with transpiration-driven flow through stomata (stomatal conductance 80-150 mmol H₂O m⁻² s⁻¹) veraison (BBCH 81, week 10) showing xylem cavitation and cessation of xylem import as stomata close and transpiration declines post-veraison (BBCH 81-89, weeks 10-18) showing phloem-exclusive import with xylem vessels air-filled and non-conducting, demonstrating developmental switch in vascular bundle mesocarp import pathway from xylem-dominant to phloem-exclusive at veraison transition. Chardonnay vascular bundle mesocarp distribution efficiency= vascular bundles representing 1.8-2.4% of mesocarp volume providing import supporting uniform sugar distribution throughout mesocarp, with sugar concentration showing <3% spatial variation (central mesocarp 22.2%, mid-radius 21.8%, peripheral 22.5%, all within analytical variation) despite concentrated import through small vascular bundle mesocarp fraction, demonstrating effective distribution from vascular import points throughout storage parenchyma tissue. Pinot Noir vascular bundle mesocarp anatomical variation= berry with 3 major dorsal bundles (less than typical 4-5) showing adequate import supporting normal composition development (23.8% sugars, 6.9 g/L acids at maturity, comparable to 4-bundle berries) berry with 5 bundles showing similar composition (24.1% sugars, 6.8 g/L acids), demonstrating vascular bundle number variation (3-5 major bundles typical) with functional redundancy as varying bundle numbers achieve similar import capacity and composition outcomes through compensatory changes in bundle size or import rates.
A vascular bundle mesocarp is the mesocarp tissue containing or immediately adjacent to vascular bundles that penetrate and ramify through the mesocarp, serving as import pathways distributing sugars, water, and minerals throughout the flesh, typically representing <2-3% of mesocarp volume but critically important for resource distribution supporting berry growth and composition development. Vascular bundle mesocarp organization comprises= (1) primary (major) vascular bundles representing main import pathways from pedicel through berry to stylar end, typically 3-5 major dorsal bundles running longitudinally through berry oriented 90-120° apart around berry circumference (2) secondary (lateral) vascular bundles branching from primary bundles and ramifying throughout mesocarp tissue creating distribution network (3) vascular bundle-adjacent mesocarp cells receiving direct import from phloem before distribution to more distant cells. Vascular bundle mesocarp vascular tissue structure= (1) xylem vessels 20-50 μm diameter comprising thick-walled dead cells providing water transport pathway, functional during early berry development (Phase I) but largely non-functional during ripening as vessels cavitate and cease conducting (2) phloem tissue comprising sieve elements (12-28 μm diameter thin-walled living cells conducting sugars and other solutes), companion cells (supporting sieve element function), and phloem parenchyma, remaining functional throughout ripening as primary import pathway (3) vascular cambium (when present) between xylem and phloem potentially producing additional vascular tissue during berry growth (4) bundle sheath cells surrounding vascular tissue creating boundary between conducting tissues and surrounding mesocarp parenchyma. Vascular bundle mesocarp solute unloading= (1) phloem import delivering sugars (primarily sucrose in phloem converted to glucose and fructose in berry), water, minerals, and other solutes to vascular bundle regions (2) symplastic unloading (cell-to-cell through plasmodesmata) or apoplastic unloading (into cell wall space) releasing solutes from phloem into surrounding tissues (3) distribution from vascular bundle-adjacent cells to distant mesocarp cells through cell-to-cell transport (short distances) or diffusion through apoplast (longer distances) (4) uptake by storage parenchyma cells through plasma membrane and vacuolar membrane transporters accumulating solutes in vacuoles. Vascular bundle mesocarp development= early development (Phase I) showing xylem-dominant import with functional xylem vessels conducting water and dissolved minerals supporting rapid cell division and growth transition at/before veraison from xylem to phloem dominance with xylem vessels cavitating (filling with air and ceasing function) and phloem becoming primary import pathway ripening (Phase II) showing phloem-exclusive import conducting sugars and water supporting berry expansion and composition development with non-functional xylem vessels remaining as structural elements. Vascular bundle mesocarp in berry composition distribution= vascular bundle distribution network creating relatively uniform sugar and acid distribution throughout mesocarp despite localized import through vascular bundles cells immediately adjacent to vascular bundles potentially showing slightly earlier composition changes during ripening as they receive import first before distribution to distant cells overall mesocarp composition showing <5% spatial variation in sugars and acids despite concentrated import through <3% of tissue volume containing vascular bundles, demonstrating effective distribution from vascular bundle mesocarp throughout tissue.
Vascular_Bundle_Mesocarp represents the tissue containing or adjacent to vascular import pathways and is distinguished from bulk storage mesocarp by presence of conducting tissues and import functions. Sub-classification into Primary and Secondary vascular bundle mesocarp distinguishes major import pathways from lateral distribution branches. Understanding vascular bundle mesocarp is essential for comprehending resource import mechanisms, for understanding xylem-to-phloem transition at veraison, and for detailed anatomical characterization of berry internal structure and resource distribution systems.
Gefäßbündel-Mesokarp
Leitgewebe-Fruchtfleisch
conducting tissue zone
vascular flesh tissue
vaskuläres Fruchtfleisch
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 190-198 Rogiers, S.Y. et al. (2006). Vascular function in berries. Australian Journal of Grape and Wine Research 12=203-214 Chatelet, D.S. et al. (2008). Xylem structure and connectivity in grapevine shoots. American Journal of Botany 95=1076-1085.
Vascular Bundle Mesocarp
Merlot vascular bundle termination in brush region at berry maturity (BBCH 89) shows 3 major terminations representing main pedicel bundle endpoints with xylem vessels diminishing from 35-40 μm diameter proximally to 15-20 μm at terminal elements, 8-12 secondary terminations from diverging traces extending 1-3 mm into basal berry tissues, and terminal phloem elements showing partial callose deposition indicating reduced transport activity
A vascular bundle termination is the ending region of a vascular bundle where xylem and phloem conducting elements conclude their transport pathway, occurring at the distal ends of vascular traces within the brush, receptacle, or berry tissues where vascular supply reaches its final destination. Vascular bundle terminations in grape berry brush and receptacle represent the anatomical endpoints where= pedicel vascular bundles diverge into multiple smaller traces entering the berry, individual vascular traces extend into berry tissues (exocarp, mesocarp, or toward seed regions) and ultimately terminate, xylem vessels and phloem sieve tubes end their continuous conducting pathways through specialized terminal elements or gradual diminishment. Vascular bundle termination anatomy includes= terminal xylem elements (final tracheary elements that may show reduced diameter, modified wall thickening, or blind endings), terminal phloem elements (final sieve tube elements and companion cells), transition zones where vascular elements gradually decrease in size and organization, potential connection to non-vascular cells (parenchyma or other cell types receiving vascular supply), and supporting cells (parenchyma, sclerenchyma) surrounding terminal vascular elements. Vascular bundle termination patterns in berry brush region include= major bundle terminations where main pedicel bundles end at receptacle level (2-4 main terminations per berry representing primary pedicel vascular supply), secondary trace terminations where smaller vascular branches extending into berry tissues conclude (6-15 secondary terminations distributed in receptacle and basal berry regions), and peripheral terminations where finest vascular traces end in berry peripheral tissues. Vascular bundle termination functional significance includes= endpoints for xylem water and mineral delivery (terminal xylem elements release water and solutes to surrounding tissues through pit connections or cell wall permeability), endpoints for phloem sugar and nutrient delivery (terminal phloem unloads sugars and nutrients to surrounding berry tissues through symplastic or apoplastic pathways), transition from vascular to non-vascular transport (from long-distance vascular transport to local cell-to-cell movement in berry tissues), and potential sites of vascular occlusion or blockage (terminations may be sites where xylem embolism or phloem callose deposition terminates further transport). Vascular bundle termination development during berry growth includes= early establishment during fruit set and early berry development (BBCH 71-75) when vascular architecture differentiates, maintenance of active terminations during berry growth phases (BBCH 75-85) with ongoing xylem and phloem delivery, and potential modification during late ripening (BBCH 85-89) including xylem occlusion, phloem callose deposition, or vascular disconnection. Vascular bundle termination variation includes= termination number (4-20 per berry depending on vascular bundle branching complexity), termination position (primarily in receptacle/brush region but some extending into berry basal tissues), termination type (abrupt endings versus gradual diminishment), and functionality status (active conducting terminations versus occluded non-functional terminations).
Vascular_Bundle_Termination emphasizes vascular pathway endings and is distinguished from Terminal_Vascular_Bundle (specific bundle endpoint structures) and End_Point_Vascular_Bundle (final conducting elements) by encompassing complete termination region anatomy and function. Critical for understanding berry vascular supply architecture, nutrient and water delivery endpoints, vascular transport cessation during ripening, and berry-pedicel vascular disconnection mechanisms.
Bundle endpoint
Gefäßbündel-Terminierung
Leitbündel-Endpunkt
Vascular ending
Vaskuläres Ende
Pratt, C. (1971). Reproductive anatomy in cultivated grapes - a review. American Journal of Enology and Viticulture, 22(2), 92-109. | Keller, M. (2020). The Science of Grapevines= Anatomy and Physiology (3rd ed.). Academic Press.
Vascular Bundle Termination
Syrah vascular cambium at 15 years post-planting shows cylindrical meristem 4-5 cell layers thick positioned 24-30 mm from bark surface, fusiform initials 420-520 μm length producing axial xylem and phloem elements, ray initials generating multiseriate rays 3-5 cells wide, cambium circumference 280-340 mm requiring continuous anticlinal divisions, spring reactivation late March triggered by rising auxin from expanding buds, active division April-September producing 3.2-4.8 mm xylem annual ring (comprising 88-91% of production) and 0.4-0.7 mm phloem increment, asymmetric division ratio approximately 7':1 xylem:phloem cells
Vascular cambium is the bifacial lateral meristem in Vitis vinifera trunk positioned between secondary xylem and secondary phloem, producing secondary xylem (wood) centripetally toward trunk center and secondary phloem (inner bark) centrifugally toward trunk surface, representing the primary growth-determining meristem responsible for trunk diameter increase and vascular tissue production. Vascular cambium cellular composition includes= fusiform initials (vertically elongated cambial cells 200-600 μm length, 15-30 μm width producing axial xylem and phloem elements including vessels, fibers, and axial parenchyma), ray initials (smaller approximately isodiametric cells 30-60 μm dimensions producing ray parenchyma), cambial initial cells maintaining indefinite division capacity (capable of continued division throughout vine life potentially spanning decades), immediate derivatives in differentiation (xylem mother cells differentiating into various xylem elements, phloem mother cells becoming sieve tubes, companion cells, fibers, and parenchyma), and cambial zone totaling 3-6 cell layers (initials plus recent derivatives in early differentiation stages). Vascular cambium location and organization includes= cylindrical sheath completely encircling trunk axis, positioned between wood (inward) and inner bark (outward), continuous from root cambium through trunk to branch cambium (maintaining vascular system integration), depth from bark surface varying by vine age (10-15 mm in young 3-5 year trunks, 20-30 mm in mature 12-20 year trunks, 35-60 mm in old 30-50 year trunks), and circumference increasing with trunk diameter (expanding from ~30 mm in year 1 to 250-350 mm at year 15, requiring cell division and ray initial multiplication to maintain tissue proportions). Vascular cambium division patterns include= periclinal divisions (parallel to trunk surface producing xylem mother cells inward and phloem mother cells outward), anticlinal divisions (perpendicular to surface increasing cambial circumference as trunk diameter expands), asymmetric production ratio (approximately 5-10 xylem cells produced per 1 phloem cell creating wood-dominated trunk structure), fusiform initial division producing axial elements (vessels, fibers, axial parenchyma in xylem, sieve tubes, companion cells, fibers, parenchyma in phloem), and ray initial division producing ray parenchyma (maintaining radial continuity between xylem and phloem). Vascular cambium seasonal activity includes= spring reactivation (late March-April in Northern Hemisphere temperate climates, synchronized with budbreak 0-2 weeks before bud opening), active division period (April-September with peak activity May-July during rapid shoot growth), progressive slow-down (August-September as shoot growth ceases and fruit ripens), fall cessation (September-October coinciding with leaf senescence and dormancy preparation), and winter dormancy (November-February with minimal metabolic activity and no cell division awaiting spring reactivation). Vascular cambium hormonal regulation includes= auxin (indole-3-acetic acid) primary activator (produced in expanding buds and young leaves, transported basipetally through phloem to cambium triggering division), cytokinin enhancing division (root-produced cytokinins transported in xylem sap promoting cambial activity), gibberellin modulating differentiation (affecting xylem element size and lignification patterns), ethylene potentially inhibiting cambial growth (elevated ethylene during stress potentially reducing division), and auxin-cytokinin ratio determining tissue type (high auxin/cytokinin ratio favoring xylem production, more balanced ratio producing phloem, ratio modulation controlling wood versus bark proportions). Vascular cambium annual production includes= secondary xylem increment (annual ring width 1.5-5 mm radial increase depending on vine vigor, site conditions, and stress levels), secondary phloem increment (inner bark increasing 0.2-0.8 mm radially per year, narrower than xylem due to asymmetric cambial production), xylem comprising 85-92% of total vascular cambium production by volume (reflecting strong centripetal bias in cell division and expansion), phloem comprising 8-15% of production, and cumulative effect (decades of cambial activity producing 40-100 mm wood radius and 3-8 mm living phloem depth in mature 15-25 year trunks). Vascular cambium environmental responses includes= water availability strongly affecting activity (drought stress reducing or stopping division, adequate water supporting vigorous cambial growth), temperature requirements (sustained temperatures >10-15°C necessary for division, optimal activity 20-28°C), nutrient availability affecting production rates (nitrogen promoting cambial division and cell expansion, phosphorus and potassium supporting metabolism), carbohydrate supply from photosynthesis (current photosynthate and stored reserves fueling energy-intensive cell division and wall synthesis), and mechanical stress potentially influencing activity (tension and compression forces from wind or weight possibly modulating cambial responses). Vascular cambium age-related changes include= fusiform initial lengthening (initials 200-300 μm in young cambium elongating to 400-600 μm in old cambium through apical intrusive growth), ray initial multiplication (new ray initials arising from fusiform initial divisions as cambium circumference increases maintaining ray density), cambial circumference expansion (requiring continuous anticlinal divisions and cell rearrangement), declining division rates in old vines (senescent cambium showing reduced activity producing narrow rings 0.5-2 mm compared to 3-6 mm in youth), and maintained division capacity (even ancient 80-100 year cambium retaining ability to produce annual increments albeit narrow ones). Vascular cambium significance in trunk function includes= producing conducting xylem (vessels transporting water and minerals from roots to canopy), producing conducting phloem (sieve tubes transporting photosynthate from leaves to roots and storage tissues), generating storage tissues (xylem and phloem parenchyma storing carbohydrate reserves), enabling trunk diameter increase (cambial activity accounting for >95% of trunk radial growth), and supporting vine longevity (perennial cambial activity enabling decades of continued vascular tissue production maintaining vine function).
Vascular_Cambium emphasizes bifacial meristem producing xylem and phloem and is distinguished from Trunk_Cambium (including cork cambium) and Cork_Cambium (producing periderm) by specific function producing vascular tissues. Critical for understanding trunk diameter increase, wood and bark production, seasonal growth dynamics, hormonal regulation of secondary growth, and vascular tissue renewal enabling long-term trunk function.
Bifacial cambium
Holzkambium
Secondary cambium
Sekundärkambium
Vaskuläres Kambium
Wood cambium
Vascular Cambium
Vascular connection establishment in grapevine grafts requires 21-42 days under optimal conditions, with xylem continuity typically preceding phloem connection by 7-14 days. Compatible unions show 80-95% of the graft interface occupied by functional vascular tissue by 8-12 weeks, with vessel diameter at the union reaching 60-80% of normal internode vessel size. Hydraulic conductance across mature compatible unions reaches 70-90% of ungrafted stem conductance. Incompatible combinations may show only 30-50% vascular coverage with discontinuous elements, reduced vessel diameter (40-60% of normal), and hydraulic conductance below 50% of ungrafted values. Phloem connectivity assessed by carboxyfluorescein transport shows bidirectional movement within 35-45 days in compatible unions.
A graft union formation disposition that is the inherited capacity of a Vitis vinifera graft combination to establish functional xylem and phloem connections across the graft interface through redifferentiation of callus cells into vascular elements, enabling water and nutrient transport from rootstock to scion and photosynthate transport from scion to rootstock.
Vascular connection represents the critical functional outcome of graft union formation, as inadequate vascular continuity limits water, nutrient, and photosynthate exchange regardless of callus development success. The process requires redifferentiation of callus cells along auxin gradients, with polar auxin transport establishing the directional pattern of vascular elements. Xylem connections form through transdifferentiation of callus cells into tracheary elements, while phloem connections require sieve element differentiation and companion cell association. Union anatomy assessment by sectioning provides definitive compatibility evaluation.
Gefäßverbindung
Leitbündelanschluss
vascular bridge formation
vascular differentiation capacity
vaskuläre Konnektivität
xylem-phloem reconnection
Melnyk CW (2017) Plant grafting: insights into tissue regeneration. Regeneration 4:3-14 Gambetta GA et al. (2017) Grapevine rootstocks and their role in managing salinity in Australian viticulture. Australian Journal of Grape and Wine Research 23:137-149
Vascular Connection Disposition
Cabernet Sauvignon vascular connection zone at véraison (BBCH 83) contains 3-4 longitudinal vascular bundles in pedicel with xylem vessels 25-40 μm diameter providing declining water transport (0.2-0.4 mg/berry/hour) and phloem sieve tubes providing increasing sugar import (0.8-1.5 mg sucrose/berry/hour), branching at receptacle into 8-12 vascular traces entering berry exocarp and extending into mesocarp
A vascular connection zone is the specialized region within the pedicel where vascular tissues (xylem and phloem) transition from the rachis vascular system into the pedicel and ultimately connect to the berry vascular network, serving as the critical conduit for water, mineral, and photosynthate transport supporting berry development and ripening. The vascular connection zone in Vitis vinifera pedicels represents a complex anatomical region where= rachis vascular bundles diverge to form pedicel traces (typically 2-4 vascular bundles entering each pedicel from rachis vascular network), pedicel vascular bundles run longitudinally through pedicel providing transport pathway, and pedicel vasculature branches and expands at berry attachment forming receptacle vascular network that connects to berry exocarp and extends into berry interior vascular system. Vascular connection zone anatomy includes= proximal region (rachis-pedicel junction) with vascular bundle divergence from rachis, mid-pedicel region with longitudinal vascular bundles embedded in parenchymatous cortex, distal region (pedicel-berry junction) with vascular expansion and branching forming receptacle, xylem tissue containing vessels (tracheary elements 15-50 μm diameter) for water and mineral transport, phloem tissue containing sieve tube elements and companion cells for sugar and nutrient transport, and supporting tissues (parenchyma, sclerenchyma providing structural support and storage). Vascular connection zone function varies by berry developmental phase= during Phase I berry growth (BBCH 73-77) xylem transport dominates with high water and mineral import supporting rapid berry cell division and expansion (xylem flow 0.1-0.5 mg water/berry/hour), during Phase II lag phase (BBCH 77-81) both xylem and phloem are active with balanced water import and beginning sugar accumulation, during Phase III ripening (BBCH 81-89) phloem increasingly dominates with high sugar import supporting rapid berry sugar accumulation (phloem sugar transport 0.5-2.0 mg/berry/hour) while xylem flow declines or reverses. Vascular connection zone structural characteristics influencing berry development include= vascular bundle number (more bundles provide greater transport capacity supporting larger berry size), xylem vessel diameter (larger vessels provide greater hydraulic conductance, 30-50 μm diameter vessels in large-berried cultivars versus 15-30 μm in small-berried cultivars), phloem sieve tube abundance (greater phloem capacity supports higher sugar accumulation rates during ripening), and vascular bundle positioning (symmetrical distribution provides uniform berry nutrition). Vascular connection zone modifications during ripening include= progressive xylem vessel embolism or blockage (reducing water transport capacity during Phase III), phloem sieve tube callose deposition (potentially regulating sugar transport), vascular tissue lignification (providing structural reinforcement), and potential vascular disruption in abscission zone region (contributing to berry hydraulic isolation at maturity). Cultivar variation in vascular connection zone includes= bundle number variation (2-5 bundles per pedicel), vessel diameter variation (15-50 μm), vascular network complexity at receptacle (simple versus highly branched), and vascular capacity relative to berry size (affecting berry expansion potential and sugar accumulation capacity).
Vascular_Connection_Zone emphasizes the complete vascular transport pathway and is distinguished from Primary_Vascular_Connection_Zone (main structural conduits) and Conducting_Vascular_Connection_Zone (active transport function) by encompassing entire vascular architecture from rachis divergence through berry entry. Critical for understanding berry size determination through vascular capacity, sugar accumulation patterns during ripening, water relations and berry expansion, and cultivar differences in berry development potential.
Gefäßverbindungszone
Leitbündel-Anschlussbereich
Pedicel vascular system
Vascular transition zone
Vaskuläre Übergangszone
Vascular Connection Zone
Investigation of Vector_Transmission_Resistance_Disposition at UC Davis compared vine mealybug (Planococcus ficus) population development and GLRaV-3 transmission efficiency across 8 cultivars in controlled greenhouse assays. Viruliferous mealybugs (10 per plant) were caged on healthy test vines, and vector populations were assessed weekly for 8 weeks while transmission was evaluated by RT-PCR at 12 weeks post-infestation. Significant cultivar variation in mealybug reproduction was observed, with 'Cabernet Sauvignon' supporting mean populations of 245 individuals per plant versus only 82 on 'Thompson Seedless' (67% reduction), suggesting differential Vector_Transmission_Resistance_Disposition. However, GLRaV-3 transmission rates did not differ significantly (75-90% across all cultivars), indicating that population suppression was insufficient to substantially reduce transmission probability. Analysis of leaf surface characteristics revealed that 'Thompson Seedless' possessed higher trichome density (32 vs 18 per mm²) and thicker epicuticular wax, potentially deterring mealybug settlement. Field trials showed similar trends but confirmed that natural vector pressure and transmission rates did not differ practically among cultivars under commercial conditions, limiting the current utility of vector-based resistance for leafroll management.
A Vector_Transmission_Resistance_Disposition is a Leafroll_Resistance_Disposition that inheres in a Vitis plant and is realized through host characteristics that prevent or limit Grapevine leafroll-associated virus transmission by mealybug (Pseudococcidae) and soft scale (Coccidae) insect vectors, primarily through resistance or non-preference affecting vector feeding behavior on aerial plant parts. This disposition represents indirect viral resistance operating by interrupting the vector-mediated transmission cycle rather than through direct antiviral mechanisms. GLRaV transmission occurs when viruliferous vectors feed on phloem tissues, introducing viral particles that colonize sieve elements. Unlike the soil-dwelling nematode vectors of GFLV that can be excluded through resistant rootstocks, leafroll vectors are mobile insects that feed on above-ground tissues including leaves, shoots, and clusters, necessitating scion-based resistance. Vector transmission resistance mechanisms may include leaf surface characteristics (trichome density, wax composition) affecting vector settling and probing behavior, phloem chemistry deterring sustained feeding, induced defenses triggered by vector feeding that reduce transmission efficiency, and antibiosis factors reducing vector survival or reproduction. Variation in mealybug population development among grapevine cultivars has been documented, with some cultivars supporting lower populations that could theoretically reduce transmission pressure. However, strong genetic resistance to leafroll vectors has not been clearly demonstrated in Vitis germplasm, and practical vector management relies primarily on insecticides, mating disruption, and biological control rather than host plant resistance.
Vector_Transmission_Resistance_Disposition is_a Leafroll_Resistance_Disposition. This class represents resistance to mealybug and soft scale insect vectors rather than to GLRaVs themselves. Unlike Xiphinema_Transmission_Resistance_Disposition for fanleaf where strong nematode resistance effectively controls disease, no strong resistance to leafroll vectors has been identified that provides practical disease control. The aerial feeding habit of leafroll vectors (versus soil-dwelling nematodes for fanleaf) means rootstock-based resistance is not applicable, making scion resistance the relevant target. Current leafroll vector management relies on cultural and chemical approaches rather than host resistance.
Blattroll-Vektor-Resistenz
Schmierläuse-Resistenz
Vektor-Übertragungs-Resistenz-Disposition
mealybug resistance disposition
scale insect transmission resistance
Daane KM et al. (2012) Biology and management of mealybugs in vineyards, Arthropod Management in Vineyards, Springer Almeida RPP et al. (2013) Ecology and management of grapevine leafroll disease, Frontiers in Microbiology 4:94 Bertin S et al. (2016) Transmission of Grapevine leafroll-associated virus-1 and -3 by soft scale and mealybug vectors, Proceedings of the International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG) 18:96-97
Vector Transmission Resistance Disposition
A vegetative active bud at node 2 on a winter-pruned cane of Vitis vinifera cv. Pinot Gris in late April, developing into a vigorous shoot with 8 fully expanded leaves, 3 partially expanded leaves, and 2 tendrils by mid-May, showing no inflorescence initials upon dissection, typical of basal node positions which exhibit low bud fruitfulness despite favorable growing conditions during primordium differentiation in the previous season.
An active bud containing exclusively or predominantly vegetative primordia that develops into a shoot bearing only leaves without inflorescence structures, characterized by rapid shoot elongation, successive leaf emergence at regular phyllotactic intervals, tendril formation at nodes opposite leaf positions, and absence of flower cluster development. Vegetative active buds typically arise from secondary or tertiary positions in compound buds following primary bud damage, from basal nodes with inherently low fruitfulness, from water shoots or suckers emerging from trunk or arm latent buds, or from lateral buds on current season growth, producing shoots that contribute to canopy leaf area and photosynthetic capacity but not directly to current season yield.
Vegetative active buds are essential for rebuilding canopy following winter pruning, frost damage, or other defoliation events, but excessive vegetative growth from these buds can create canopy management challenges including excessive shading, delayed fruit maturation, and increased disease pressure. The vegetative versus reproductive fate of developing buds is determined during inflorescence primordium initiation in the previous season (typically 6-8 weeks after budbreak), influenced by light exposure, temperature, vine water status, and nitrogen availability during the critical differentiation period.
Blattknospe
non-fruiting active bud
sterile active bud
vegetative austreibende Knospe
vegetative breaking bud
Based on Srinivasan & Mullins (1981) "Physiology of flowering in the grapevine - a review" American Journal of Enology and Viticulture 32=47-63 Vasconcelos et al. (2009) "The flowering process of Vitis vinifera= a review" American Journal of Enology and Viticulture 60=411-434 Lebon et al. (2008) "Sugars and flowering in the grapevine (Vitis vinifera L.)" Journal of Experimental Botany 59=2565-2578.
Vegetative Active Bud
A vegetative dominant mixed bud on Vitis vinifera cv. Cabernet Sauvignon in a young, vigorous vineyard block in Napa Valley, developing into a shoot with 22 fully expanded leaves, shoot length 185cm by mid-July, one small inflorescence bearing only 45 berries clustered near shoot base, strong lateral shoot development from multiple nodes, creating excessive canopy density requiring aggressive shoot thinning and leaf removal to maintain adequate light penetration and air circulation for disease management and fruit ripening.
A mixed active bud developing into a shoot with predominant vegetative growth relative to reproductive structures, characterized by vigorous shoot elongation, abundant leaf production (typically 15-25 leaves), long internodes, strong apical dominance, and proportionally small or limited inflorescence development (one small cluster or reduced cluster size). Vegetative dominant mixed buds typically arise from high-vigor vineyard sites with deep fertile soils, generous water and nitrogen availability, or unbalanced vines following light cropping or severe winter damage, producing shoots that contribute substantially to canopy leaf area and shading problems but provide limited fruit yield and often show reduced bud fruitfulness for the following season due to excessive shading during critical inflorescence initiation period.
Vegetative dominant mixed buds indicate imbalanced vines with excessive vigor, requiring management intervention to restore balance through strategies including increased crop load (lighter pruning, minimal cluster thinning), deficit irrigation during vegetative growth period, cover crop competition for water and nutrients, and aggressive canopy management (shoot positioning, hedging, leaf removal). Continued vegetative dominance over multiple seasons leads to progressive decline in bud fruitfulness as shaded canopy conditions during summer suppress inflorescence initiation, creating a negative feedback loop of increasing vegetative vigor and decreasing reproductive capacity.
vegetativ-dominante gemischte Knospe
vegetative-biased mixed bud
vigorous mixed bud
Based on Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Howell (2001) "Sustainable grape productivity and the growth-yield relationship= A review" American Journal of Enology and Viticulture 52=165-174 Naor et al. (2002) "Irrigation and crop load influence fruit size and water relations in field-grown 'Spadona' pear" Journal of the American Society for Horticultural Science 127=252-255.
Vegetative Dominant Mixed Bud
Monitoring Vegetative_Growth_Vigor_Quality in a Marlborough Sauvignon Blanc vineyard through weekly shoot length measurements reveals rapid growth of 4-5 cm/day during the pre-flowering period (BBCH 15-57), followed by growth cessation within two weeks post-fruit set under imposed water stress, indicating successful vigor regulation that concentrates vine resources into fruit development rather than continued vegetative expansion.
Vegetative_Growth_Vigor_Quality is a Vigor_Quality that inheres in a grapevine (Vitis vinifera L.) and represents the dynamic expression of vigor during active growth periods as manifested through real-time shoot extension, leaf production, and lateral development processes. This quality specifically captures the temporal dimension of vigor expression — the rate and intensity of vegetative growth as it occurs throughout the growing season rather than the accumulated biomass measured post-dormancy. In BFO terms, this quality reflects the vine's instantaneous growth state, changing continuously from dormancy release through shoot maturation, with characteristic phases of rapid early-season extension followed by growth cessation as the vine transitions toward reproductive development and dormancy preparation. The distinction between growth vigor and accumulated vigor expression is viticulturally significant because identical final biomass can result from dramatically different growth patterns, with implications for fruit quality and vine health. Rapid early-season growth followed by timely cessation indicates optimal vigor regulation, while extended late-season growth suggests imbalanced source-sink relationships, delayed fruit ripening, and incomplete shoot lignification that compromises winter hardiness. Shoot growth rate measurements during the rapid growth phase (BBCH 15-65) provide the most sensitive indicators of vine water and nutrient status, with growth cessation timing serving as a critical indicator of appropriate water stress imposition in regulated deficit irrigation programs. Cultivar differences in vegetative growth vigor are pronounced, with early-maturing varieties like Chasselas and Müller-Thurgau typically showing compact growth patterns with early cessation, while late-maturing varieties like Cabernet Sauvignon and Mourvèdre maintain growth potential deep into the season under favorable conditions. Viticultural interventions affecting vegetative growth vigor include irrigation timing and quantity, nitrogen fertilization, cover crop competition, rootstock selection, and pruning severity, with each factor influencing not only growth magnitude but also temporal distribution of vegetative activity throughout the season.
Vegetative_Growth_Vigor_Quality captures the process-oriented aspect of vigor, distinguishing it from Vigor_Expression_Quality which represents accumulated biomass outcomes. This class serves as parent for specific growth measurements including shoot growth rate, internode length, total shoot length, and lateral development — all assessable during the active growing season. The temporal sensitivity of these measurements makes them valuable for in-season management decisions, contrasting with pruning weight assessments that only become available post-dormancy.
Triebwachstumsstärke
Wachstumsstärke
active growth vigor
aktive Wuchskraft
growth vigor
seasonal growth vigor
vegetative Wuchskraft
vegetative growth intensity
Pellegrino A, Lebon E, Simonneau T, Wery J (2005) Towards a simple indicator of water stress in grapevine (Vitis vinifera L.) based on the differential sensitivities of vegetative growth components. Australian Journal of Grape and Wine Research 11:306-315 Schultz HR, Matthews MA (1988) Vegetative growth distribution during water deficits in Vitis vinifera L. Australian Journal of Plant Physiology 15:641-656
Vegetative Growth Vigor Quality
Research trials in Marlborough Sauvignon Blanc compared vegetative reproductive balance quality across irrigation treatments, finding that vines receiving 70% ETc replacement achieved optimal balance with yield-to-pruning-weight ratios of 8.2 and complete sugar accumulation to 23.5°Brix, while fully irrigated vines showed vegetative dominance with ratios of 4.1 and delayed ripening requiring additional hangtime, and severely restricted vines exhibited reproductive excess with ratios exceeding 15 but incomplete flavor development due to premature leaf senescence and limited photosynthetic capacity during the critical final ripening phase.
Vegetative reproductive balance quality is a vigor balance quality that inheres in a bearing grapevine organism and represents the specific proportional relationship between carbon allocation to vegetative growth structures and reproductive development as quantified through direct measurement of biomass partitioning between leaves, shoots, and fruit. This quality captures the fundamental either-or nature of resource allocation wherein photoassimilates directed toward shoot elongation and leaf expansion are unavailable for berry sugar accumulation, while conversely strong reproductive sink demand can limit vegetative development and carbohydrate reserve storage. The ontological basis of vegetative reproductive balance quality rests on the principle that grapevines, as perennial organisms with indeterminate growth habit, must continuously mediate the competing demands of current-season reproduction against vegetative growth supporting both immediate photosynthetic capacity and future productive potential through bud fruitfulness and reserve accumulation. Quantification of this balance typically employs biomass-based metrics including the classic Ravaz Index expressing yield mass relative to dormant pruning weight, with optimal values generally ranging from 5-10 depending on cultivar and production goals, though premium red wine production may target lower values around 4-6 to achieve concentrated fruit. The physiological mechanisms governing vegetative reproductive balance quality involve hormonal signaling networks wherein gibberellins and cytokinins promote vegetative growth while abscisic acid accumulation during fruit ripening contributes to growth cessation and reproductive prioritization. Source-sink relationships fundamentally determine balance outcomes, with developing clusters exerting powerful sink strength that can mobilize carbohydrates from distant leaves and even woody storage tissues when local supply proves insufficient. Environmental factors modulating this balance include water availability, where moderate deficit shifts allocation toward reproduction, nitrogen status, where excess promotes vegetative dominance, and temperature regimes affecting both photosynthetic rate and developmental timing. The temporal dynamics of vegetative reproductive balance quality follow characteristic seasonal patterns wherein spring growth is predominantly vegetative, transitioning through flowering and fruit set to increasingly reproductive dominance as berries accumulate sugars during ripening. Management interventions including shoot thinning, hedging, leaf removal, and crop load adjustment provide tools for manipulating this balance toward specific quality objectives, with the recognition that optimal balance represents a dynamic target varying with cultivar characteristics, site conditions, vintage variation, and intended wine style.
Vegetative_Reproductive_Balance_Quality is_a Vigor_Balance_Quality, representing the biomass-based assessment of growth-reproduction equilibrium. This class focuses on the direct measurement of carbon allocation through tissue weights rather than indirect indicators like canopy density. The Ravaz Index (yield/pruning weight) serves as the primary quantitative expression of this quality. Distinguished from Vine_Balance_Index_Quality which incorporates additional parameters beyond simple biomass ratios.
Biomasseverteilung
Kohlenstoffallokation
Wachstum-Reproduktion-Verhältnis
biomass allocation balance
carbon partitioning equilibrium
growth-reproduction balance
vegetativ-reproduktive Balance
vegetative-reproductive partitioning
Ravaz L (1911) L'effeuillage de la vigne. Annales de l'École Nationale d'Agriculture de Montpellier 11:216-244 Bravdo B et al. (1985) Effect of crop level and crop load on growth, yield, must and wine composition, and quality of Cabernet Sauvignon. American Journal of Enology and Viticulture 36:125-131 Kliewer WM, Dokoozlian NK (2005) Leaf area/crop weight ratios of grapevines: influence on fruit composition and wine quality. American Journal of Enology and Viticulture 56:170-181
Vegetative Reproductive Balance Quality
A vegetative shoot on Vitis vinifera cv. Merlot arising from node 2 (basal position with typical low fruitfulness), currently 125cm length in late June with 16 fully expanded leaves providing approximately 940 cm² photosynthetic leaf area, 3 tendrils, no inflorescences present upon examination, representing common vegetative shoot from low-fruitfulness basal node contributing substantial photosynthetic capacity (estimated 15-18% of total vine leaf area from this single shoot) supporting fruit maturation on reproductive shoots from mid-cane positions, managed through shoot positioning into trellis catch wires but retained rather than removed due to adequate canopy space and valuable photosynthetic contribution to whole-vine carbon balance.
A shoot bearing only vegetative structures without inflorescences, characterized by nodes bearing leaves and tendrils but no flower clusters, developmental origin from vegetative buds (typically basal positions, secondary or tertiary buds, latent buds, or water shoots), and function providing photosynthetic capacity and canopy volume without direct current-season reproductive contribution though supporting overall vine function and potentially providing fruiting positions for subsequent seasons. Vegetative shoots represent non-fruiting growth arising from various sources including low-fruitfulness basal buds, secondary/tertiary bud activation following primary damage, latent bud breakage on permanent wood, and deliberately retained renewal positions, contributing to vine leaf area, carbohydrate production, and architectural development while requiring management decisions regarding retention versus removal based on canopy density, vigor balance, and renewal wood requirements.
Vegetative shoots are distinguished from reproductive shoots (bearing inflorescences) and mixed shoots (bearing both vegetative structures and reproductive organs), with the vegetative designation indicating complete absence of current-season reproductive structures. However, vegetative shoots are not necessarily unproductive long-term, as buds differentiating on vegetative shoots may develop inflorescence primordia for following season depending on light exposure, timing, and vine conditions during critical June-August differentiation period. Management of vegetative shoots involves decisions balancing photosynthetic contribution (retention supporting whole-vine carbon status) against canopy density concerns (excessive vegetative shoots creating shading, disease pressure, delayed ripening), with typical strategies retaining well-positioned vegetative shoots with adequate spacing while removing crowded or poorly-positioned shoots regardless of vegetative versus reproductive status.
Blatttrieb
leaf-bearing shoot
non-fruiting shoot
sterile shoot
vegetativer Trieb
Based on Pratt (1974) "Vegetative anatomy of cultivated grapes - a review" American Journal of Enology and Viticulture 25=131-150 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 113-145 Srinivasan & Mullins (1981) "Physiology of flowering in the grapevine - a review" American Journal of Enology and Viticulture 32=47-63.
Vegetative Shoot
A veinlet network in an intercostal blade region on a mature leaf at node 7 of Vitis vinifera cv. Pinot Noir in Burgundy, France, forming dense reticulate mesh between adjacent tertiary veins spaced 1.2 mm apart, individual veinlet segments measuring 0.3-0.6 mm length creating irregular polygonal areoles (mesh openings) averaging 0.28 mm² area, veinlet diameter of approximately 0.08 mm barely visible under 10× magnification, containing single tracheid or very small xylem element (8-12 μm diameter) providing final water delivery, few phloem cells facilitating local photosynthate collection, veinlets extending to within 2-3 mesophyll cells (approximately 60-80 μm) of all intercostal regions ensuring no photosynthetic cells are more than 3-4 cell layers from nearest vascular element, creating total veinlet density of 10.2 mm per mm² representing approximately 75% of total venation length (including all vein orders), forming complete reticulate network isolating approximately 850 areoles per mm² blade area, each areole containing 15-25 mesophyll cells serviced by surrounding veinlets, creating maximum mesophyll-to-vein diffusion distance of approximately 80 μm enabling efficient gas and solute exchange, measured at BBCH stage 77 (berries beginning to touch) in mid-July, representing the fine-scale terminal vascular network providing ultimate hydraulic connection to all photosynthetic tissues.
The finest vascular elements in grapevine leaves forming the ultimate reticulate network in Vitis vinifera, characterized by very small diameter typically 0.05-0.15 mm at or near the limit of visibility without magnification, minimal vascular anatomy often consisting of single tracheids or small xylem elements and few phloem cells, fourth-order and higher hierarchical positions representing terminal branches of the venation system, irregular reticulate organization lacking strict hierarchical branching patterns and creating dense mesh network, very high density with typical total veinlet length of 8-12 mm per mm² blade area creating the dominant component of total venation length, short individual veinlet segments typically 0.2-0.8 mm creating fine mesh with areole (mesh opening) sizes of 0.1-0.5 mm², and critical functional roles including providing ultimate water delivery to all mesophyll cells minimizing diffusion distances, collecting photosynthates from mesophyll for export through higher-order veins, creating the final anatomical bottleneck determining maximum hydraulic capacity and photosynthetic rates, and forming areolar networks that isolate local damage preventing vascular dysfunction from spreading throughout the blade. Veinlets represent the terminal elaboration of the venation system, developing during final stages of leaf expansion filling all spaces between tertiary veins, exhibiting cultivar and environmental variation in density affecting blade hydraulic properties, and creating the fine-scale vascular architecture that ultimately determines leaf functional capacity through direct interaction with photosynthetic mesophyll cells.
Veinlets are distinguished from tertiary and higher-order lateral veins by very small diameter, irregular reticulate organization, and terminal position in the vascular hierarchy. The transition from the finest tertiary veins to veinlets is somewhat arbitrary, typically based on diameter thresholds (often 0.15 mm) and loss of strict hierarchical branching patterns. Veinlets constitute the majority of total vein length despite their small individual size, with typical contributions of 70-80% of total venation length per unit area. Functional importance is critical as veinlets create the final hydraulic connection to mesophyll cells, with veinlet density and distribution determining maximum diffusion distances and therefore maximum rates of transpiration and photosynthesis supportable before hydraulic limitation occurs. Sun versus shade leaves show veinlet density differences, with sun leaves typically developing higher densities creating shorter diffusion distances enabling higher maximum photosynthetic rates, while shade leaves show lower densities reflecting lower functional demands and resource limitations. Developmental timing involves late expansion phases with veinlet differentiation filling spaces between established higher-order veins. Anatomical simplification to single tracheids or minimal vascular elements represents minimum functional vascular capacity, with some veinlets possibly lacking phloem and serving purely xylem delivery roles. Areolar organization creates compartmentalization isolating small blade regions, potentially limiting damage spread from localized injuries or pathogen infections.
Feinader
fine veinlet
minor vein
terminal vein
terminale Vene
Äderchen
Based on Sack & Holbrook (2006) "Leaf hydraulics" Annual Review of Plant Biology 57=361-381 Brodribb et al. (2007) "Leaf hydraulic capacity in ferns, conifers and angiosperms= impacts on photosynthetic maxima" New Phytologist 165=839-846 Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 195-218.
Veinlet
Velvetiness quality in Pomerol Merlot-dominated blend (velvetiness rating 9/10) exemplifies the textural ideal for this appellation, where the variety's inherently soft tannins combined with clay soil concentration and careful élevage produce the quintessential velvety mouthfeel characterized by dense, plush contact and gentle grip reminiscent of fine velvet fabric, with the sumptuous texture supporting and enhancing the plum, truffle, and mineral flavors rather than competing for attention, demonstrating how velvetiness defines the Pomerol textural signature.
Velvetiness quality is a texture perception quality that represents the soft, plush, dense tactile impression reminiscent of velvet fabric, characterizing the rich, cushioned mouthfeel that combines smoothness with substantial body and gentle grip, creating the luxurious texture associated with full-bodied wines of exceptional quality. Within the BFO framework, this quality specifically addresses the plush-dense dimension of texture perception, distinguishing velvety character from silky (which lacks the density and body) or creamy (which emphasizes dairy-like richness over fabric-like plushness) while recognizing velvetiness as a textural ideal for full-bodied wine styles where opulent mouthfeel complements concentrated flavor. The ontological significance lies in velvetiness representing the textural pinnacle for concentrated, age-worthy wines, where the combination of smooth surface, dense body, and refined tannin grip creates the sumptuous, enveloping mouthfeel that defines the tactile expression of great wines from Pomerol, Barossa, Napa Valley, and other regions producing powerful, elegant styles. The perceptual characteristics of velvetiness include the sensation of dense, cushioned contact between wine and oral surfaces, smooth surface character without grain or friction, subtle underlying grip from refined tannins (like velvet's nap), and overall impression of luxurious richness that makes the wine feel substantial yet refined. The compositional requirements for velvetiness include elevated alcohol and extract (providing the necessary density), refined, well-integrated tannins (contributing gentle grip without harshness), adequate polysaccharides (smoothing texture), and the overall concentration that creates dense body while maintaining elegance through component balance. The variety and style associations of velvetiness center on full-bodied, age-worthy reds, with Merlot particularly noted for velvety potential (explaining Pomerol's textural reputation), while Cabernet Sauvignon, Shiraz, and other varieties achieve velvetiness when concentration and refinement combine successfully. Understanding velvetiness quality enables recognition of textural excellence in full-bodied styles, quality assessment incorporating the specific velvety character valued in concentrated wines, and aging evaluation where velvetiness development indicates successful maturation of tannic structure.
Velvetiness_Quality addresses the plush-dense dimension of texture perception, representing the textural ideal for full-bodied, concentrated wine styles. This quality combines smoothness with density and refined grip, distinguishing it from lighter silkiness or dairy-like creaminess. The class supports quality assessment in concentrated wines where velvety texture indicates successful achievement of power with elegance.
Samtigkeit
dense smoothness
luxurious texture
luxuriöse Textur
plush mouthfeel
plüschiges Mundgefühl
samtartige Textur
velvet-like texture
Velvetiness Quality
Venation conductivity assessment comparing sun and shade leaves in Merlot reveals anatomical acclimation affecting hydraulic capacity, with sun leaves showing vein density of 13.8 mm/mm² and venation conductivity contributing 65% of total Kleaf (high conductivity matching high transpiration demand), while shade leaves show vein density of 9.2 mm/mm² with venation contributing only 45% of lower total Kleaf (reduced vascular investment appropriate for lower demand), demonstrating how venation architecture acclimation matches hydraulic supply to expected transpiration environment.
Venation conductivity quality is a leaf hydraulic conductance quality that represents the water transport capacity of the leaf vein network from petiole insertion through midrib, secondary, and minor veins to the sites of water delivery to mesophyll tissues, characterizing the xylem-based component of leaf hydraulic conductance that distributes water throughout the leaf lamina. Within the BFO framework, this quality specifically addresses the vascular component of leaf water transport, distinguishing vein conductivity from mesophyll pathways while recognizing that the hierarchical venation network must supply water to all regions of the leaf lamina with appropriate redundancy and efficiency to support spatially distributed transpiration. The ontological significance lies in venation architecture determining both the efficiency and safety of leaf water distribution, where vein density, hierarchical organization, and connectivity affect both the conductivity available to supply transpiration and the vulnerability to localized failure that could strand leaf regions without water supply. The anatomical basis for venation conductivity involves the hierarchical branching pattern from large-diameter, high-conductivity midrib and secondary veins through progressively smaller tertiary and minor veins to the dense network of finest veins that deliver water near every mesophyll cell, with conductivity at each level determined by vessel dimensions and at the network level by branching architecture and connectivity. The vein density relationship with hydraulic capacity shows higher vein density (length per unit leaf area, typically 8-15 mm/mm² in grapevine) providing both higher conductivity and greater redundancy against local failure, with grapevine showing relatively high vein density consistent with its high transpiration capacity. The vulnerability of venation to embolism contributes to the decline in Kleaf under water stress, with evidence that leaf vein embolism occurs at less negative water potentials than stem embolism in many species including grapevine, consistent with hydraulic segmentation that sacrifices leaves to protect more costly stems. Understanding venation conductivity quality enables assessment of the vascular contribution to leaf water supply, interpretation of leaf anatomy effects on hydraulic function, and evaluation of the venation network's capacity to distribute water throughout the leaf lamina.
Venation_Conductivity_Quality addresses the xylem-based vein network component of leaf hydraulic conductance. This quality is determined by vein density and hierarchical architecture affecting water distribution. The class supports leaf anatomy interpretation and hydraulic capacity assessment through venation conductivity characterization.
Adernnetzleitfähigkeit
Blattgefäßleitfähigkeit
Kven
Venationsleitfähigkeit
leaf vascular conductance
vein conductivity
venation hydraulics
Sack L, Scoffoni C (2013) Leaf venation: structure, function, development, evolution, ecology and applications in the past, present and future. New Phytologist 198(4): 983-1000 Brodribb TJ et al. (2007) Leaf maximum photosynthetic rate and venation are linked by hydraulics. Plant Physiology 144(4): 1890-1898
Venation Conductivity Quality
Pinot Noir in Burgundy shows veraison during late August-early September Year N+1 at BBCH 83-85, with berry softening initiating with 12-22% firmness decline from 3.2 to 2.6 N, anthocyanin accumulation beginning 0.06-0.14 mg/g creating visible pink color at stylar end, sugar acceleration to 2.2% per week, malate declining from 0.85% to 0.72%, berry growth 14 mg per day, water uptake 0.48 g per berry per day, ABA spiking to 920 ng/g, veraison spreading over 8-12 days within cluster, occurring 9.5-10.5 weeks post-flowering at 1250-1350 GDD, under continental conditions of 22-27°C temperature, typical of this early-to-mid season cultivar beginning ripening transformation
Veraison represents the dramatic developmental transition phase during berry development when berry transforms from lag phase quiescence to active ripening state, characterized by berry softening as universal marker across all cultivars, color development in red cultivars from anthocyanin synthesis, sugar accumulation acceleration from enhanced phloem import, organic acid decline from respiration and metabolism, berry expansion resumption from water uptake, and coordinated physiological changes establishing ripening phase, occurring over 5-14 days within cluster with asynchronous berry transformation typically weeks 8-12 post-fertilization corresponding to BBCH 83-85. This transformation phase occurs during late summer to early autumn in late August-September in temperate regions Year N+1, representing lag-to-ripening transition marking onset of berry ripening. Veraison mechanism involves berry softening through cell wall modification particularly pectin degradation creating first measurable firmness decline 10-25%, color development in red cultivars through anthocyanin biosynthesis creating visible pink to red color, sugar accumulation acceleration from hexose transporter activation increasing import 2-5 fold, organic acid decline from enhanced malate respiration, berry expansion from increased water uptake driven by osmotic gradient from sugars, abscisic acid spike coordinating or triggering transformation, and integrated changes establishing ripening metabolism and growth. Berry softening characteristics shows firmness declining from pre-veraison maximum 2.5-4.5 N to initial softening 2.0-3.5 N representing 10-25% decline, progressive softening continuing through ripening, softening as universal veraison marker occurring in all cultivars, softening from cell wall modification particularly pectin degradation in middle lamella and cell walls, and softening as defining transformation. Color development in red cultivars shows anthocyanin biosynthesis pathway activating, anthocyanin accumulation beginning with 0.05-0.2 mg/g at veraison onset increasing rapidly, visible color change from green to pink or light red, color typically initiating at stylar end progressing toward pedicel, color development as red cultivar veraison marker, and color from phenolic pigment synthesis. White cultivar veraison shows white cultivars lacking anthocyanin synthesis, veraison marked by softening and possible yellowing from chlorophyll decline, softening as primary visible marker, and white cultivar transformation without color development. Sugar accumulation acceleration shows sugar import increasing from lag minimal 0.5-1.5% per week to veraison-ripening 1.5-3.5% per week, hexose transporters activated from accumulated protein, phloem unloading enhanced, sugars increasing from 12-17% at veraison to progressive accumulation, and sugar increase as ripening characteristic. Organic acid decline shows malate respiration increasing, malate declining from 0.7-1.0% at veraison onset, tartrate relatively stable, pH rising from 3.1-3.4 toward ripening values, and acid dynamics characteristic of ripening onset. Berry expansion resumption shows berry growth rate increasing from lag minimal <5 mg per day to veraison-ripening 10-30 mg per day, water uptake accelerating from <0.1 to 0.3-0.8 g per berry per day, osmotic gradient from sugars driving water influx, berry volume increasing 5-15% during veraison period, and expansion marking lag phase exit. Abscisic acid dynamics shows ABA spiking from lag levels 50-200 ng/g to veraison peak 500-2000 ng/g, ABA as veraison trigger or correlate, hormone coordinating transformation, ABA possibly regulating gene expression and metabolism, and ABA spike as hormonal signature. Metabolic transformation shows respiratory rate increasing from lag 1.0-1.5 μmol O₂ g⁻¹ h⁻¹ to ripening 1.5-2.5 μmol O₂ g⁻¹ h⁻¹, carbon flux redirected to ripening processes including anthocyanin and sugar metabolism, secondary metabolism intensified, and metabolic shift establishing ripening state. Gene expression changes shows ripening genes highly upregulated including anthocyanin pathway, sugar transporters, softening enzymes, ripening transcription factors activating target genes, and expression establishing ripening program. Veraison timing shows onset typically weeks 8-12 post-fertilization varying with cultivar and environment, early cultivars showing veraison 7-9 weeks, mid-season 9-11 weeks, late cultivars 11-14 weeks, and timing determining harvest date. Heat accumulation triggering shows growing degree days from flowering to veraison as major timing determinant, cultivar-specific GDD thresholds triggering veraison, warm conditions advancing veraison, cool conditions delaying, and temperature as primary timing factor. Veraison asynchrony shows veraison beginning in earliest berries while others remain in lag, typical within-cluster veraison spread 5-14 days, asynchrony creating heterogeneous berry populations, earliest berries at stylar cluster positions often, and asynchrony as characteristic pattern. Cluster veraison progression shows progressive transformation wave through cluster, earliest 10-20% berries showing veraison at onset, 50% veraison midpoint used as cluster veraison date, eventual 80-95% berries transformed, and progression over days to weeks. Berry appearance at veraison shows berries softening to touch, red cultivars showing pink to red color development, white cultivars possibly showing yellowing, berry size increasing, and appearance indicating transformation. Veraison assessment shows visual observation of color and softening, firmness measurement quantifying softening, sugar and acid analysis showing compositional changes, and measurements characterizing veraison. BBCH staging shows BBCH 81 late lag pre-veraison, BBCH 83 berries beginning to develop color in red or berries beginning to soften in white as veraison onset, BBCH 85 softening of berries as advanced veraison, and staging documenting progression. Environmental effects on veraison include temperature as dominant factor with GDD threshold triggering onset, water stress possibly advancing veraison through ABA, light possibly affecting anthocyanin synthesis, and environmental factors determining timing and quality. Temperature effects shows warm conditions advancing veraison 7-14 days, cool conditions delaying comparable period, and temperature affecting cluster uniformity. Water stress effects shows moderate deficit possibly advancing veraison 3-7 days through enhanced ABA, severe stress potentially disrupting development, and stress as timing factor. Veraison significance shows transition marking lag phase end and ripening onset, determining harvest timing from veraison date, establishing berry composition trajectory toward maturity, critical quality determinant, and major developmental milestone. Viticultural management shows veraison timing determining harvest planning, cluster thinning often at veraison, deficit irrigation possibly managing veraison timing, and management affecting veraison and quality. The process encompasses berry softening with initial and progressive phases, alongside color development in red cultivars covered separately, sugar and acid dynamics, and berry expansion. Duration veraison transformation over 5-14 days at cluster level with individual berry transformation over 2-5 days. Regional timing shows veraison during late August-September in temperate climates varying with cultivar and season. Viticultural significance includes representing critical developmental transition determining ripening onset and harvest date, establishing berry quality trajectory, demonstrating environmental sensitivity particularly temperature, and serving as major vineyard monitoring milestone. Research applications include analyzing veraison triggering mechanisms and regulation, investigating hormonal control particularly abscisic acid, evaluating environmental effects on timing and uniformity, quantifying transformation dynamics and asynchrony, and assessing veraison as determinant of ripening progression and harvest quality.
Veraison represents lag-to-ripening transition within Berry_Development. Characterized by berry softening (universal, 10-25% initial firmness decline), color development in red cultivars (anthocyanin 0.05-0.2 mg/g onset), sugar acceleration (1.5-3.5%/week), acid decline, berry expansion (10-30 mg/day), ABA spike (500-2000 ng/g). Duration 5-14 days cluster-level asynchrony. BBCH 83-85. Timing: GDD-triggered, weeks 8-12 post-fertilization. Subdivided into Berry_Softening (and other aspects). Critical developmental milestone determining harvest timing. Temperature dominant environmental factor.
Beeren-Transformation
Farbumschlag
Reifungsbeginn
Véraison
berry transformation
color change
ripening onset
veraison period
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 468-485 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Veraison
Merlot in Bordeaux shows veraison approach during early September Year N+1 at BBCH 81-83, with veraison imminent within 1-2 days, earliest 8-12% berries showing faint pink color from 0.04-0.06 mg/g anthocyanin at stylar end, firmness in earliest berries declining 8-12% from peak, growth rate increasing to 8-14 mg per day in transforming berries, abscisic acid at 880 ng/g approaching peak, under oceanic conditions of 20-26°C temperature, typical of this mid-season cultivar at transformation threshold
Veraison approach represents the imminent transformation phase during lag phase termination when veraison onset becomes imminent within 0-3 days, earliest berries possibly showing initial transformation signs, berry at developmental threshold ready for immediate transformation, abscisic acid at or approaching peak levels, and berry poised for definitive veraison onset, occurring during final 1-2 days of lag phase as transformation begins. This imminent veraison phase occurs during late summer to early autumn in early-mid September in temperate regions Year N+1, proceeding over final 1-2 days before or as veraison begins, corresponding to BBCH 81-83 transition period. Approach mechanism involves berry achieving complete veraison readiness, transformation threshold reached or crossed, earliest berries beginning visible transformation, abscisic acid peak levels triggering or correlating with onset, developmental competence fully established, and integrated readiness enabling immediate transformation. Transformation imminence shows veraison onset within 0-3 days in earliest berries, berry at or crossing transformation threshold, visible changes imminent or beginning, and imminence as approach defining characteristic. Developmental threshold shows berry achieving critical state enabling veraison, accumulated ripening machinery adequate for transformation, metabolic state supporting ripening, hormonal signals triggering or permitting progression, and threshold crossing enabling veraison. Earliest transformation signs shows first berries possibly showing initial color in red cultivars with trace anthocyanin 0.02-0.08 mg/g, first measurable softening with 5-15% firmness decline in earliest berries, and earliest signs indicating transformation beginning. Color onset in red cultivars shows anthocyanin synthesis initiating in earliest berry sectors, typical color onset at stylar end calyx scar region, color appearing as faint pink tinge, and color as earliest visible transformation marker. Anthocyanin synthesis onset shows anthocyanin biosynthesis pathway flux increasing dramatically, earliest accumulation 0.02-0.08 mg/g anthocyanins, synthesis rate beginning acceleration, and anthocyanin as color development mechanism. Softening onset shows earliest berries showing initial firmness decline, texture beginning change from hard to yielding, firmness declining 5-15% from pre-veraison maximum in earliest berries, and softening as earliest tactile transformation marker. Berry appearance shows earliest berries showing faint color in red cultivars or beginning softening in white cultivars, majority of berries still in late lag state, cluster showing beginning asynchronous transformation, and appearance indicating transformation onset. Abscisic acid peak approach shows ABA approaching or at maximum levels 500-2000 ng/g, hormone accumulation reaching plateau, ABA peak coinciding with veraison onset, and hormone as transformation signal. Abscisic acid timing shows ABA beginning increase 3-7 days pre-veraison, rapid accumulation during final days, peak levels 1-3 days surrounding veraison onset, and timing as hormonal trigger or correlate. Berry growth rate shows possible slight increase from lag minimal <5 mg per day toward 5-15 mg per day in earliest transforming berries, growth increase from beginning water uptake acceleration, and growth as possible approach indicator. Berry composition shows sugars 13-17%, acids 0.7-0.9%, pH 3.1-3.4, and composition at transformation starting point. Cluster heterogeneity shows earliest berries approaching or entering veraison, middle berries in late lag, latest berries in early-mid lag, and heterogeneity creating asynchronous transformation pattern. Veraison asynchrony shows typical veraison spread over 5-14 days across cluster, earliest berries transforming first, progressive transformation wave through cluster, and asynchrony as characteristic pattern. Environmental effects on approach include temperature determining approach timing with heat accumulation triggering transformation when threshold reached, water availability affecting transformation dynamics, stress possibly advancing approach through hormonal effects, and environmental factors determining veraison onset date. Temperature as trigger shows growing degree days accumulating to cultivar-specific threshold triggering veraison, heat accumulation as primary timing determinant, warm conditions advancing approach, and temperature effect as major viticultural factor. The process represents first aspect of lag phase termination emphasizing imminent transformation, preceding softening initiation and lag phase exit. Duration final 1-2 days representing imminent veraison. Regional timing shows veraison approach during early-mid September as transformation becomes imminent. Viticultural significance includes representing imminent veraison onset determining harvest date, demonstrating transformation readiness, enabling veraison prediction from approach signs, and establishing ripening commencement affecting quality and timing. Research applications include analyzing transformation thresholds and triggering, investigating abscisic acid dynamics and veraison onset, evaluating environmental effects on transformation timing, quantifying earliest transformation markers, and assessing veraison approach as predictor of onset timing and ripening progression.
Veraison_Approach is first aspect of Lag_Phase_Termination, distinguished by imminent transformation. Represents veraison within 0-3 days. Earliest berries showing initial color (0.02-0.08 mg/g anthocyanin) or softening (5-15% decline). ABA at or approaching peak 500-2000 ng/g. Growth rate possibly 5-15 mg/day. Developmental threshold reached. Duration 1-2 days. Precedes Softening_Initiation. Transformation imminent.
Transformations-Schwelle
Veraison-Annäherung
Veraison-Beginn-Annäherung
ripening commencement
transformation threshold
veraison onset approach
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 468-475 Coombe BG (1995) Adoption of a system for identifying grapevine growth stages, Aust J Grape Wine Res 1:104-110
Veraison Approach
Cabernet Sauvignon in Napa Valley shows veraison preparation during early September Year N+1 at BBCH 81, with anthocyanin biosynthesis genes maximally expressed CHS 88 fold, DFR 96 fold, UFGT 115 fold, sugar transporters VvHT1 28 fold, VvHT2 24 fold, softening enzymes polygalacturonase 26 fold, pectin methylesterase 22 fold, enzyme activities anthocyanin pathway 38%, sugar transport 42%, softening 32% eventual maxima, berry compositionally optimized at 15% sugars, 0.8% acids, pH 3.2, under warm climate conditions of 24-30°C temperature, typical of this late-season cultivar at veraison threshold
Veraison preparation represents the final ripening readiness phase during pre-veraison lag when ripening gene expression reaches near-maximal pre-transformation levels 30-80 fold increases, ripening enzymes accumulate to 30-50% eventual maximum activity enabling rapid function, metabolic pathways achieve full activation capacity, berry structural and compositional state optimizes for transformation, and comprehensive preparations position berry for immediate veraison onset when triggered, occurring during final days of pre-veraison lag. This final preparation phase occurs during late summer to early autumn in late August-early September in temperate regions Year N+1, proceeding over final 3-7 days before veraison, corresponding to BBCH 81 late pre-veraison period. Preparation mechanism involves ripening gene expression reaching activation-ready high levels, enzyme protein accumulation achieving functional threshold concentrations, metabolic pathway component assembly completing, berry compositional optimization for ripening initiation, developmental competence fully established, and integrated preparations creating transformation-ready state. Ripening gene expression maxima shows anthocyanin biosynthesis genes in red cultivars reaching 40-120 fold increases representing near-maximal pre-veraison levels, sugar transporter genes 15-35 fold preparing import machinery, softening enzyme genes 12-30 fold enabling texture transformation, volatile biosynthesis genes beginning expression, and comprehensive gene expression establishing ripening program. Anthocyanin pathway gene levels shows phenylalanine ammonia lyase PAL 30-70 fold, chalcone synthase CHS 40-100 fold, chalcone isomerase CHI 35-85 fold, flavanone 3-hydroxylase F3H 45-95 fold, dihydroflavonol reductase DFR 50-110 fold, anthocyanidin synthase ANS 45-100 fold, UDP-glucose flavonoid glucosyltransferase UFGT 60-140 fold, and complete pathway highly expressed. Anthocyanin pathway readiness shows all biosynthetic steps with high gene expression, enzyme proteins accumulated to functional levels, pathway flux capacity prepared for rapid anthocyanin synthesis, and pathway poised for color development at veraison. Sugar transporter gene levels shows hexose transporters VvHT1 18-38 fold, VvHT2 14-32 fold, VvHT3 possibly increasing, VvHT6 tonoplast transporter increasing, sucrose transporters VvSUC11 possibly 8-20 fold, VvSUC12 possibly increasing, and comprehensive transporter expression enabling maximal import. Sugar transport capacity shows plasma membrane hexose import capacity increased, tonoplast hexose storage capacity enhanced, phloem unloading efficiency prepared, and transport machinery ready for accelerated sugar accumulation. Softening enzyme gene levels shows polygalacturonase PG 15-35 fold, pectin methylesterase PME 12-28 fold, pectate lyase possibly 8-20 fold, β-galactosidase possibly 10-25 fold, expansin genes 8-22 fold, and softening machinery comprehensively expressed. Softening enzyme capacity shows pectin-degrading enzymes accumulated, cell wall modification capacity established, enzymes positioned for rapid texture change, and softening machinery ready for veraison activation. Flavor and aroma gene expression shows terpene synthase genes in aromatic cultivars beginning expression 5-15 fold, linalool synthase possibly 8-18 fold, volatile ester biosynthesis genes increasing, methoxypyrazine degradation genes possibly upregulating in some cultivars, and flavor machinery beginning assembly. Enzyme protein accumulation shows ripening enzymes accumulated to 30-50% eventual ripening maximum concentrations, enzyme activities at functional pre-activation levels, enzymes positioned in appropriate cellular compartments, and protein accumulation enabling rapid function at veraison. Enzyme activity thresholds shows anthocyanin biosynthesis enzymes 25-45% maximal activity, sugar transporters 30-50% maximal capacity, softening enzymes 20-40% maximal activity, and activity thresholds enabling veraison progression. Metabolic pathway assembly shows phenylpropanoid pathway components complete, sugar metabolism pathway enhanced, organic acid metabolism enzymes positioned for decline, and comprehensive pathway readiness. Phenylpropanoid pathway shows pathway from phenylalanine through to anthocyanins fully assembled, intermediate metabolite pools established, cofactor availability adequate, and pathway ready for high flux. Sugar metabolism preparation shows glycolysis and sugar storage pathways enhanced, vacuole hexose accumulation capacity increased, osmotic capacity for water uptake prepared, and sugar metabolism ready for ripening. Organic acid metabolism shows malate metabolism shifting toward decline, malic enzyme possibly upregulating, malate dehydrogenase patterns changing, and acid metabolism prepared for ripening reduction. Berry compositional optimization shows acids declining to 0.7-0.9% optimizing pH for enzyme function, sugars 13-16% providing accumulation base, pH 3.1-3.3 approaching optimal range for ripening enzymes, phenolic precursors accumulated providing substrates, and composition optimized for transformation. pH optimization shows pH rising toward 3.2-3.5 range optimal for many ripening enzymes, pH increase enabling enhanced enzyme activities, and pH as critical compositional parameter. Substrate availability shows phenolic precursors p-coumaric acid, caffeic acid accumulated, sugar substrates glucose, fructose available, UDP-glucose cofactor available, and substrates ready for biosynthesis. Berry structural preparation shows cell wall composition prepared for modification, membrane properties adjusted for altered transport, cellular organization ready for transformation, and structure optimized for ripening. Cell wall preparation shows pectin partially de-esterified enabling calcium cross-linking and subsequent degradation, hemicellulose and cellulose providing stable framework, wall poised for softening, and structure ready for modification. Membrane preparation shows transporter proteins inserted at high density, membrane lipid composition possibly adjusted, membrane properties enabling high flux, and membranes ready for ripening transport. Developmental competence establishment shows berry fully competent for veraison transformation, accumulated machinery sufficient for rapid progression, metabolic state enabling transformation, and competence as preparation outcome. Transformation-ready state shows berry poised at veraison threshold, minimal additional preparation required, berry responsive to veraison trigger, and state enabling immediate transformation onset. Berry appearance during preparation shows berry still predominantly green, possible trace color in earliest red cultivar berries, texture firm though possibly softening initiated, and appearance at pre-transformation state. Environmental effects on preparation include temperature affecting preparation completion rate, warm conditions accelerating final preparations, cool conditions extending preparation period, and temperature determining veraison timing. The process represents first aspect of pre-veraison lag emphasizing final ripening readiness, preceding metabolic reprogramming. Duration final 3-7 days representing preparation completion. Regional timing shows veraison preparation during late August-early September immediately before veraison onset. Viticultural significance includes representing final ripening readiness determining transformation capacity, establishing comprehensive ripening machinery enabling successful veraison, demonstrating preparation completion as veraison prerequisite, and determining veraison quality through adequate preparation. Research applications include analyzing ripening gene expression thresholds and readiness, investigating enzyme accumulation and activity requirements, evaluating metabolic pathway assembly and capacity, quantifying preparation completion markers, and assessing veraison preparation as determinant of transformation success and ripening quality.
Veraison_Preparation is first aspect of Pre_Veraison_Lag, distinguished by final ripening readiness. Represents gene expression 30-120 fold maxima (anthocyanin 40-140, transporters 15-38, softening 12-35 fold). Enzyme accumulation 30-50% ripening maxima. Complete pathway assembly. Compositional optimization (acids 0.7-0.9%, pH 3.1-3.3, sugars 13-16%). Structural preparation. Developmental competence established. Duration 3-7 days. Precedes Metabolic_Reprogramming. Transformation-ready state.
Reifungs-Kompetenz
Transformations-Vorbereitung
Veraison-Vorbereitung
final ripening readiness
finale Reifungs-Bereitschaft
ripening competence
transformation preparation
veraison machinery assembly
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 460-468 Coombe BG (1992) Research on development and ripening of the grape berry, Am J Enol Vitic 43:101-110
Veraison Preparation
Véraison synchrony assessment comparing cultivars under identical conditions reveals the genetic differences in ripening coordination, with Merlot showing high synchrony (85% of berries completing color change within 8-day window, minimal within-cluster variation, "red and green" period brief), while Grenache shows characteristically low synchrony (only 55% completion within 8-day window, substantial within-cluster variation with pink berries adjacent to green and fully colored berries persisting for 18+ days), with the inherent synchrony difference requiring cultivar-specific harvest strategies—single-date harvest suitable for Merlot while Grenache may require either extended hang time or selective harvesting to manage maturity heterogeneity.
Véraison synchrony quality is an intra-plant synchrony quality that represents the temporal coordination of véraison onset among different berries and clusters within a single grapevine, characterizing the uniformity of the ripening transition that determines the homogeneity of the berry population entering the maturation phase and strongly predicts the harvest-time uniformity that affects fruit quality and sorting requirements. Within the BFO framework, this quality specifically addresses the véraison dimension of intra-plant synchrony, distinguishing the ripening transition coordination from earlier flowering synchrony or subsequent ripening rate uniformity while recognizing that véraison synchrony is perhaps the most visible and consequential synchrony measure, as the color change (in red varieties) or softening transition makes synchrony directly observable and as véraison timing variation propagates directly to harvest-time maturity variation. The ontological significance lies in véraison synchrony determining the maturity uniformity available at any subsequent harvest date, where highly synchronized véraison produces berry populations that will achieve maturity targets simultaneously enabling single-harvest capture of optimal quality, while asynchronous véraison creates berry populations spanning a range of developmental ages that will require either extended hang time (allowing lagging berries to catch up), selective harvest (removing only mature fruit), or acceptance of maturity heterogeneity (harvesting mixed-maturity populations). The factors affecting véraison synchrony include the accumulated effects of earlier developmental synchrony (flowering synchrony propagating to véraison), cluster microclimate (shaded versus exposed clusters potentially differing in véraison timing), berry position within clusters (inner versus outer berries, shoulder versus tip), and water status during the pre-véraison period (water stress potentially accelerating véraison in some berries while delaying others). The cultivar-specific patterns of véraison synchrony show substantial variation, with some varieties (Merlot, Chardonnay) characteristically showing tight véraison synchrony, while others (Grenache, Zinfandel) inherently show extended "red and green" periods even under favorable conditions, representing genetic differences in developmental coordination that affect management requirements. Understanding véraison synchrony quality enables prediction of harvest-time maturity uniformity, assessment of sorting requirements based on expected berry population heterogeneity, and cultivar-appropriate expectations for achievable synchrony levels.
Véraison_Synchrony_Quality addresses the coordination of ripening transition timing affecting harvest-time maturity uniformity. This quality is highly visible in red cultivars and strongly predicts subsequent maturity homogeneity. The class supports harvest planning and sorting assessment through véraison synchrony characterization.
Farbumschlaguniformität
Reifebeginngleichmäßigkeit
Reifeübergangskoordination
Véraison-Synchronie
color change uniformity
maturation transition uniformity
ripening onset synchrony
véraison coordination
Coombe BG, McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6(2): 131-135 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 277-316
Veraison Synchrony Quality
The véraison timing description of Pinot Noir documents early véraison (OIV 303:3) — reaching 50% berry color change approximately 5-7 days before Chasselas in Burgundy, typically during the last week of July to the first week of August (July 25 - August 5 at Beaune), approximately 10-14 days before Cabernet Sauvignon at the same latitude. The early véraison of Pinot Noir initiates a ripening period of 35-45 days culminating in harvest during the first two weeks of September — this early ripening window positions Pinot Noir harvest before the autumn equinoctial weather deterioration that typically begins in mid-September in Burgundy, a climatic advantage that is partially responsible for the historical predominance of Pinot Noir in this northerly wine region (47°N) where later-ripening cultivars would face unacceptable harvest-period weather risk.
A phenological timing description component that records the date at which 50% of berries in the vineyard have undergone the color change and softening that marks the onset of ripening (E-L stage 35, Baggiolini stage M, BBCH 81-83) — the phenological event signaling the transition from the herbaceous growth phase to the maturation phase — scored according to OIV descriptor 303 on a scale from very early (state 1) to very late (state 9) relative to Chasselas, with véraison timing marking the beginning of the 35-55 day final ripening period during which sugar accumulates, acid declines, phenolic compounds develop, and aroma precursors are synthesized.
Veraison_Timing_Description documents the phenological transition that fundamentally transforms the berry from a photosynthetically active, acid-accumulating green organ to a sugar-importing, pigment-synthesizing ripening fruit — the most dramatic developmental shift in the grapevine annual cycle. Véraison is visible as the simultaneous onset of berry softening (loss of turgor-driven firmness as cell walls loosen and sugars begin accumulating osmotically), color change (in red cultivars, the activation of anthocyanin biosynthesis producing the green-to-red-to-blue transition in white cultivars, the shift from green to translucent yellow-green as chlorophyll degrades), and the cessation of organic acid accumulation (malic acid concentration begins declining through respiratory consumption): these coordinated changes are triggered by the hormonal transition from auxin-dominated (growth phase) to abscisic acid-dominated (ripening phase) signaling, with the ABA accumulation activating the transcription factors that drive anthocyanin biosynthesis, sugar transporter expression, and cell wall modification gene programs. The flowering-to-véraison interval (typically 45-65 days, 400-700 GDD base 10°C) is the most temperature-sensitive interphenological period — warm summers compress this interval, advancing véraison and extending the véraison-to-harvest ripening period, while cool summers delay véraison and compress the ripening period, with direct consequences for fruit composition at harvest.
Farbumschlag-Beschreibung
Reifebeginn-Zeitpunkt
Véraison-Dokumentation
color change timing record
ripening onset date
véraison period documentation
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Coombe BG and McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening, Australian Journal of Grape and Wine Research 6: 131-135 Jones GV and Davis RE (2000) Climate influences on grapevine phenology, American Journal of Enology and Viticulture 51: 249-261 Duchêne E and Schneider C (2005) Grapevine and climatic changes, Agronomy for Sustainable Development 25: 93-99
Veraison Timing Description
Veraison occurs 40-70 days after flowering across V. vinifera cultivars, with Chasselas reaching 50% color change approximately 45-50 days post-bloom and Mourvèdre requiring 65-75 days. The flowering-to-veraison interval requires 700-1100 GDD (base 10°C) depending on cultivar, with thermal time models predicting veraison onset within ±4-7 days accuracy. Veraison date in Pinot Noir at Burgundy reference sites averages August 10-15, advancing by approximately 2 days per decade since 1980. The physiological transition spans 7-21 days per vine from first to last berry depending on cultivar and cluster architecture.
A phenological timing disposition that is the inherited capacity of a Vitis vinifera plant to initiate the onset of ripening (veraison, BBCH 81-85) at a characteristic time relative to flowering and environmental conditions, marking the transition from berry growth phase to sugar accumulation and color development through coordinated hormonal, metabolic, and gene expression changes.
Veraison_Timing_Disposition serves as the parent class for early veraison, uniform veraison, and extended veraison categories. Veraison represents the most dramatic developmental transition in grape berry development, involving abscisic acid accumulation, ethylene signaling, sugar transporter activation, and anthocyanin biosynthesis initiation in colored cultivars. The timing of veraison critically affects the length of the ripening period and final fruit composition, with earlier veraison generally providing longer maturation periods in temperate climates.
Farbumschlagstermin
Reifebeginn
Véraison-Zeitpunkt
color change timing
ripening initiation trait
veraison onset timing
Coombe BG, McCarthy MG (2000) Dynamics of grape berry growth and physiology of ripening. Australian Journal of Grape and Wine Research 6:131-135 Kuhn N et al. (2014) Berry ripening: recently heard through the grapevine. Journal of Experimental Botany 65:4543-4559
Veraison Timing Disposition
A vertical cordon on Vitis vinifera cv. Trebbiano in tendone (overhead trellis) system in Puglia, Italy, extending 220cm vertically from trunk to overhead trellis structure at 200cm height, bearing 18 two-bud spurs distributed along vertical length at approximately 24cm intervals, shoots from spurs growing laterally and downward creating overhead canopy coverage, fruit hanging beneath canopy in shaded protected position, representing traditional vertical cordon architecture supporting overhead production system common in hot southern Italian regions, with vertical structure enabling efficient space utilization in high-density planting and creating favorable microclimate for fruit development through overhead shading while maintaining adequate photosynthetic capacity in extensive lateral canopy.
A cordon trained and maintained in vertical or near-vertical orientation, characterized by upward extension from trunk or head rather than lateral horizontal spread, fruiting spurs distributed along vertical axis at regular intervals, and suitability for specific training systems including pergola, tendone, and some traditional European configurations where vertical permanent structure supports lateral fruiting wood deployment rather than upward shoot growth from horizontal cordons. Vertical cordons represent alternative architectural approach to horizontal cordon dominance, employed in specific contexts including overhead canopy systems (Italian tendone, Argentine parral), high-density plantings where vertical structure conserves space, traditional systems in some regions, and situations where vertical permanent wood provides advantages for fruit exposure, canopy management, or harvest access compared to horizontal alternatives.
Vertical cordons represent minority configuration in modern viticulture, with specific applications where vertical architecture provides advantages over dominant horizontal cordon approach. Overhead trellis systems (tendone, parral, pergola) universally employ vertical cordons as structural columns supporting overhead fruiting framework, with vertical orientation essential for architectural function. Some high-density systems use vertical cordons to minimize lateral space requirements while maintaining adequate spur positions through vertical distribution. Traditional systems in some European regions employ vertical or angled cordons for historical reasons, fruit exposure objectives, or specific microclimate management. However, vertical cordons create challenges for mechanization (most equipment designed for horizontal cordon architecture), potentially uneven spur vigor (apical positions often showing greater vigor than basal), and different management approaches compared to conventional horizontal cordon systems.
ascending cordon
aufrechte Kordon
perpendicular cordon
upright cordon
vertikale Kordon
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Reynolds & Vanden Heuvel (2009) "Influence of grapevine training systems on vine growth and fruit composition= a review" American Journal of Enology and Viticulture 60=251-268 Intrieri & Poni (1995) "Integrated evolution of trellis training systems and machines to improve grape quality and vintage quality of mechanized Italian vineyards" American Journal of Enology and Viticulture 46=116-127.
Vertical Cordon
Madeleine Angevine exhibits very early ripening disposition, reaching 20°Brix approximately 30-38 days post-veraison with total season heat requirement of 900-1050 GDD (base 10°C) from budbreak. Siegerrebe and Ortega ripen in early-mid August in German wine regions, 6-8 weeks before late cultivars in the same vineyard. Pearl of Csaba, a table grape, achieves market maturity in late July in Mediterranean climates, commanding premium early-season prices. Very early cultivars typically produce wines of 10-11.5% alcohol with fresh, aromatic profiles suited to light wine styles or sparkling base production.
A ripening timing disposition that is the inherited capacity of a Vitis vinifera plant to reach harvest maturity very early in the season, typically within the first 10-15% of cultivar harvest dates for a region, requiring minimal heat accumulation from veraison to maturity and suited to cool climates or early-season fresh market windows.
Very early ripening disposition enables viticulture in marginal cool climates (southern England, Belgium, northern Germany) where heat accumulation is insufficient for later cultivars. The trait may associate with lower total phenolic development due to compressed ripening period, though aromatic compound synthesis can be favored by cooler maturation temperatures. Very early cultivars face harvest timing pressure, as rapid sugar accumulation can outpace flavor development, requiring careful monitoring to avoid overripeness.
Frühreife
first-harvest trait
sehr frühe Reife
sehr früher Reifetermin
ultra-early maturation
very precocious ripening
Skelton S (2014) Wine Growing in Great Britain, 2nd Ed, SP Skelton pp.: 89-112 Fraga H et al. (2016) Future scenarios for viticultural zoning in Europe: ensemble projections and uncertainties. International Journal of Biometeorology 60:1599-1613
Very Early Ripening Disposition
Mourvèdre exhibits very late ripening disposition, reaching 23-25°Brix approximately 65-80 days post-veraison with total season heat requirement of 1800-2100 GDD (base 10°C) from budbreak. Carignan and Graciano show similar very late ripening, completing harvest in late October to early November in Mediterranean climates. Very late ripening in Mourvèdre at Bandol (Provence) reference sites occurs at mean date of October 20-November 5, making it among the last cultivars harvested in French viticulture. Successful very late cultivar production requires Winkler Region III-V climates (>1670 GDD), limiting their range to Mediterranean, warm continental, and subtropical zones.
A ripening timing disposition that is the inherited capacity of a Vitis vinifera plant to reach harvest maturity very late in the season, typically within the final 10-15% of cultivar harvest dates for a region, requiring high heat accumulation and long growing seasons, restricting cultivation to warm or hot climates.
Very late ripening disposition evolved as an adaptation to hot, dry Mediterranean climates where extended ripening periods under declining autumn temperatures optimize flavor development. The trait becomes problematic in cool or short-season climates where incomplete ripening produces unbalanced wines with green, vegetal characteristics. Climate change is expanding the viable range for very late cultivars into previously marginal regions, though the compressed post-harvest recovery period may affect vine health and following-season fruitfulness in newly suitable areas.
Spätreife
final-harvest trait
sehr späte Reife
sehr später Erntetermin
ultra-late maturation
very tardy ripening
Huglin P, Schneider C (1998) Biologie et écologie de la vigne, 2nd Ed, Lavoisier pp.: 312-345 Anderson K et al. (2017) Which Winegrape Varieties are Grown Where? A Global Empirical Picture, University of Adelaide Press pp.: 45-78
Very Late Ripening Disposition
Gewürztraminer in cool vintages with poor fruit set exhibits very loose cluster disposition with 2-4 berries/cm rachis and OIV 204 scores of 1-3, leaving extensive visible rachis framework. Coulure-affected Merlot clusters may show very loose architecture with only 30-50 berries per cluster versus the normal 80-150. Intentionally loosened clusters through gibberellin treatment in Flame Seedless achieve 2-3 berries/cm for optimal berry sizing. Very loose clusters virtually eliminate Botrytis risk but may increase sunburn exposure on previously shaded berries and complicate mechanical harvest through berry shatter.
A cluster compactness disposition that is the inherited capacity of a Vitis vinifera plant to develop highly open clusters with widely spaced berries, extensive visible rachis structure, very low berry density (typically less than 4 berries/cm rachis), resulting from low fruit set, long pedicels, or extensive rachis branching.
Very loose cluster disposition may result from genetic factors (low fruit set cultivars, long pedicel genotypes) or environmental/management effects (coulure, gibberellin application). While advantageous for disease control, extremely loose clusters reduce yield, complicate harvest, and may indicate underlying reproductive problems. Some breeding programs target loose but not excessively sparse cluster architecture as an optimum. The PIWI cultivars often exhibit loose cluster architecture inherited from disease-resistant wild Vitis species.
schüttere Traube
sehr lockere Traube
sparse cluster disposition
stark aufgelockerte Traubenstruktur
straggly bunch
very open cluster
Töpfer R et al. (2011) New horizons for grapevine breeding. Fruit, Vegetable and Cereal Science and Biotechnology 5:79-100 Tello J, Ibáñez J (2014) Evaluation of indexes for the quantitative and objective estimation of grapevine bunch compactness. Vitis 53:9-16
Very Loose Cluster Disposition
Wild V. vinifera ssp. sylvestris staminate male flowers exhibit very short vestigial styles 0.12-0.24 mm connecting vestigial ovaries 0.22-0.38 mm to rudimentary stigmas 0.1-0.18 mm, with very short styles representing 25-38% of vestigial pistil length (0.48-0.72 mm total), requiring 20-30× magnification for accurate measurement and showing minimal transmitting tract differentiation in anatomical sections with poorly developed extracellular matrix and lack of distinct transmitting tract cell morphology, representing developmental remnants rather than functional reproductive structures. Cabernet Franc rare extreme functionally male flowers (<0.3% frequency) show very short styles 0.15-0.26 mm as part of maximally reduced pistils (total length 0.85-1.15 mm) including small ovaries 0.48-0.62 mm and minimal stigmas 0.18-0.28 mm, with very short styles showing extremely limited transmitting tract tissue and achieving <2% fruit set through combined effects of stigma non-receptivity, style inadequacy, and ovule abortion. Breeding populations segregating for sex determination produce staminate individuals with very short vestigial styles 0.14-0.22 mm that are barely distinguishable as separate structures between tiny ovaries and minute stigmas, with the very short style zone identifiable mainly through subtle narrowing of pistil diameter between the slightly wider ovary base and stigma apex rather than through obvious structural differentiation. Anatomical serial sectioning of staminate flower pistils reveals very short styles consisting of 8-15 layers of relatively undifferentiated parenchyma cells with minimal central cavity or extracellular matrix, contrasting sharply with hermaphrodite style sections showing 25-40 cell layers surrounding well-developed transmitting tract with abundant extracellular matrix, demonstrating the extreme developmental reduction characterizing very short styles.
A very short style is a short style at the extreme lower end of the length range, typically less than 0.3 mm at maturity, characteristic of staminate flowers with vestigial pistils and rare extreme functionally male flowers, representing styles so abbreviated that they may be barely distinguishable as separate structures between ovary and stigma, approaching the limit where style as a distinct pistil component becomes undefined. Very short styles are characterized by= (1) length typically 0.1-0.28 mm at anthesis, representing <40% of typical hermaphrodite style length and <30% of functionally female/pistillate style length (2) diameter often 0.08-0.18 mm creating extremely slender appearance (3) transmitting tract tissue minimally developed or poorly differentiated, with extracellular matrix production reduced or absent and specialized transmitting tract cells difficult to distinguish from surrounding parenchyma (4) connection between vestigial ovary (typically 0.2-0.5 mm) and rudimentary stigma (typically 0.1-0.25 mm) in staminate flower contexts creating extremely reduced pistil (total 0.4-0.8 mm) where very short style represents a barely distinguishable intermediate zone. Very short styles develop through extreme developmental suppression or arrest affecting pistil elongation, occurring in= (1) staminate flowers where strong male-determining genetic factors suppress female organ development, producing vestigial pistils with all components extremely reduced including very short styles (2) rare extreme functionally male flowers showing maximal pistil reduction beyond typical functionally male expression (3) developmental abnormalities in other flower types creating extreme style reduction through severe developmental disruption, though such cases are exceptionally rare. Very short styles represent near-complete style suppression while retaining minimal evidence of style presence as a pistil component, approaching but not quite reaching complete style absence which would create pistils lacking a distinct intermediate zone between ovary and stigma. The functional significance of very short styles is minimal as structures this reduced cannot support pollen tube growth effectively even if pollen were to germinate= transmitting tract tissue in very short styles shows poor differentiation lacking the elaborate extracellular matrix and specialized cellular organization characteristic of functional styles very short styles in staminate flowers are part of completely non-functional vestigial pistils that achieve <1% fruit set and represent developmental remnants rather than reproductive structures very short styles provide no reproductive advantage through reduced pollen tube transit distance as stigma non-receptivity and ovule absence/abortion prevent fertilization regardless of style dimensions. Very short styles are so abbreviated that they require magnification (typically 15-30×) for clear observation and measurement, making them relevant primarily for detailed anatomical studies rather than for field-based pistil assessments. The extreme shortening in very short styles may reflect threshold effects in developmental regulation, where style-promoting genetic or hormonal signals fall below levels necessary to induce significant style elongation, producing minimal residual style tissue.
Very_Short_Style represents the extreme lower end of style length range and is distinguished from Reduced_Short_Style (0.25-0.45 mm, moderate short style expression) by extreme abbreviation (<0.3 mm). Very short styles occur almost exclusively in staminate flowers with vestigial pistils and represent near-complete style suppression while retaining minimal vestigial structure. Very short styles have no reproductive function and are relevant primarily for developmental biology studies of organ suppression and for detailed characterization of staminate flower morphology. The very short style condition approaches the threshold where style ceases to be a distinguishable pistil component.
Miniatur-Stempelgriffel
extremely reduced style
sehr kurzer Griffel
vestigial style
vestigialer Stylus
Very Short Style
Vessel conductivity assessment comparing vigorous and moderate rootstocks reveals hydraulic strategy differences, with vigorous rootstock producing wide vessels (mean diameter 85 μm) achieving theoretical specific conductivity of 62 kg/m/s/MPa (high efficiency) but P50 of -1.4 MPa (moderate vulnerability), while moderate rootstock shows narrower vessels (55 μm), lower conductivity of 28 kg/m/s/MPa, but improved safety with P50 of -2.1 MPa, demonstrating the efficiency-safety trade-off in vessel design that affects rootstock hydraulic characteristics transferred to grafted scions.
Vessel conductivity quality is a stem hydraulic conductivity quality that represents the water transport capacity of individual xylem vessel elements and their interconnected networks, characterizing the fundamental conduits of long-distance water transport whose dimensions, connections, and integrity determine the bulk of stem hydraulic conductance in woody plants including grapevine. Within the BFO framework, this quality specifically addresses the conduit-level dimension of xylem transport, distinguishing the intrinsic conductivity of vessel networks from whole-stem conductivity (which integrates vessel properties with stem cross-sectional organization) while recognizing that vessel characteristics are the primary determinants of both transport capacity and vulnerability to dysfunction. The ontological significance lies in vessel properties establishing the fundamental trade-off between hydraulic efficiency and safety, where wider vessels provide dramatically higher conductivity (fourth-power scaling with diameter) but are generally more vulnerable to embolism, creating the efficiency-safety trade-off that shapes xylem evolution and determines hydraulic strategy across plant lineages. The anatomical determinants of vessel conductivity include vessel diameter (the dominant factor, with conductivity ∝ diameter⁴ according to Hagen-Poiseuille), vessel length (longer vessels reducing resistance from end-wall pit membranes), pit membrane properties (porosity affecting both inter-vessel flow and embolism vulnerability), and vessel network organization (connectivity and redundancy affecting the consequences of individual vessel failure). The measurement of vessel conductivity employs hydraulic methods (perfusing stem segments with water under pressure), anatomical approaches (calculating theoretical conductivity from vessel dimensions), and dye or particle perfusion (visualizing functional conducting area), with comparison of measured to theoretical conductivity indicating the degree of dysfunction from embolism or other factors. The environmental and developmental variation in vessel properties shows wider vessels in vigorous, well-watered conditions (maximizing conductivity) and narrower, safer vessels under stress or in peripheral stem positions (prioritizing embolism resistance), with this plasticity enabling some matching of hydraulic properties to experienced conditions. Understanding vessel conductivity quality enables assessment of the anatomical basis for stem hydraulic capacity, evaluation of the efficiency-safety trade-off in xylem design, and interpretation of how vessel characteristics affect drought tolerance through embolism vulnerability.
Vessel_Conductivity_Quality addresses the conduit-level determinants of xylem transport capacity and vulnerability. This quality reflects the efficiency-safety trade-off in vessel design that shapes drought tolerance. The class supports anatomical interpretation of hydraulic function through vessel conductivity and dimensional analysis.
Gefäßleitfähigkeit
Konduitkapazität
Xylemgefäßeffizienz
conduit conductivity
vessel hydraulic efficiency
xylem vessel capacity
Sperry JS et al. (2006) Size and function in conifer tracheids and angiosperm vessels. American Journal of Botany 93(10): 1490-1500 Brodersen CR et al. (2010) The dynamics of embolism repair in xylem. Plant Physiology 154(3): 1088-1095
Vessel Conductivity Quality
Anatomical characterization of Vessel_Diameter_Disposition at Stellenbosch University employed microscopic analysis of transverse stem sections across 15 cultivars to quantify vessel diameter distributions and relate anatomy to hydraulic function. Sections from standardized positions on current-year shoots were stained and imaged, with vessel diameters measured for 200+ vessels per cultivar. Additionally, theoretical conductivity (Kth) was calculated from vessel dimensions assuming Hagen-Poiseuille flow. Vessel_Diameter_Disposition showed significant cultivar variation. 'Shiraz' produced the widest vessels with mean diameter of 165 μm, maximum 285 μm, and high calculated Kth of 45 kg m⁻¹ MPa⁻¹ s⁻¹. 'Pinot Noir' showed intermediate dimensions (mean 138 μm, max 225 μm, Kth 28). 'Grenache' exhibited the narrowest vessels (mean 118 μm, max 195 μm, Kth 18) among major cultivars. Vessel frequency showed inverse pattern, with narrow-vesseled 'Grenache' compensating somewhat through higher vessel density (125 vessels mm⁻² versus 85 for 'Shiraz'). The efficiency-safety trade-off was confirmed by correlation between mean vessel diameter and P50 (r = 0.62), with large-vesseled cultivars showing lower cavitation resistance. Environmental plasticity was demonstrated by comparing irrigated versus deficit-irrigated vines of 'Cabernet Sauvignon', with water deficit during shoot development reducing mean vessel diameter by 18% (142 versus 173 μm) and improving P50 by 0.4 MPa, representing acclimatory adjustment of Vessel_Diameter_Disposition enhancing hydraulic safety at the cost of transport efficiency.
A Vessel_Diameter_Disposition is a Hydraulic_Architecture_Disposition that inheres in the xylem of a Vitis vinifera plant and is realized through the anatomical dimensions of xylem vessels, determining the trade-off between hydraulic conductance efficiency (favored by large vessels) and hydraulic safety (enhanced by small vessels). This disposition represents the anatomical foundation of xylem hydraulic properties, as vessel diameter has powerful effects on both water transport capacity (conductance scales with the fourth power of radius according to the Hagen-Poiseuille equation) and vulnerability to dysfunction (larger vessels tend to be more susceptible to cavitation and embolism spread). Grapevine is notable for producing exceptionally large vessels among woody plants, with maximum vessel diameters commonly exceeding 200 μm and sometimes reaching 400 μm in vigorous shoots, enabling the high transpiration rates that support rapid growth but creating inherent vulnerability to hydraulic failure. Vessel_Diameter_Disposition varies among cultivars and rootstocks, with genotypes producing larger vessels showing higher conductance under adequate water supply but potentially greater vulnerability to drought-induced conductivity loss. Vessel diameter is also plastic, with environmental factors including water availability during wood formation, temperature, and vigor affecting vessel dimensions, such that drought conditions during xylogenesis typically produce narrower, more numerous vessels than well-watered conditions.
Vessel_Diameter_Disposition is_a Hydraulic_Architecture_Disposition. Vessel diameter is the primary anatomical determinant of xylem conductance (scales with radius⁴). Grapevine produces exceptionally large vessels enabling high transport capacity but creating cavitation vulnerability. Vessel diameter varies among genotypes and shows environmental plasticity. Narrow vessels provide safety but reduce conductance plants often compensate through vessel frequency. The efficiency-safety trade-off in vessel dimensions underlies the broader Hydraulic_Architecture_Disposition.
Gefäßdurchmesser-Disposition
Vaskuläre-Anatomie
Xylem-Gefäßgröße
vascular anatomy
vessel size
xylem conduit diameter
Tyree MT, Zimmermann MH (2002) Xylem Structure and the Ascent of Sap, 2nd Edition, Springer-Verlag, Berlin Lovisolo C, Schubert A (1998) Effects of water stress on vessel size and xylem hydraulic conductivity in Vitis vinifera L., Journal of Experimental Botany 49:693-700 Hochberg U et al. (2017) Grapevine petioles are more sensitive to drought induced embolism than stems: evidence from in vivo MRI and microCT observations of hydraulic vulnerability segmentation, Plant Cell and Environment 40:1831-1842
Vessel Diameter Disposition
Wild V. vinifera ssp. sylvestris male plants in riparian forests of the Iberian Peninsula and southern France commonly exhibit vestigial pistil flowers with minute central structures 0.25-0.35 mm long appearing under 15× magnification as small undifferentiated tissue masses surrounded by five robust stamens producing 30,000-38,000 pollen grains per flower, achieving 0% fruit set, and functioning identically to male only flowers in pollen production and dispersal despite retention of minimal pistil remnants. Staminate individuals in breeding populations derived from wild dioecious × cultivated hermaphrodite crosses frequently show vestigial pistil flowers with pistil remnants 0.2-0.4 mm requiring careful microscopic examination for detection, creating classification challenges in early seedling evaluation but ultimately equivalent to male only types in fruit set failure and unsuitability for fruit production.
A vestigial pistil flower is a reduced pistil flower in which the pistil is reduced to an extremely small, barely recognizable remnant showing minimal organ differentiation, typically measuring 0.2-0.4 mm in total length and appearing as a small undifferentiated protuberance or cellular mass in the flower center with no clearly distinguishable ovary, style, or stigma components, representing near-maximum pistil suppression while retaining minimal morphological evidence of pistil developmental initiation. Vestigial pistil flowers are characterized by= (1) an extremely minute central structure <0.4 mm long occupying the position where a functional pistil would occur, often requiring magnification (10-20×) for clear visualization (2) pistil remnant showing minimal cellular differentiation with largely undifferentiated parenchyma cells lacking the specialized tissue types characteristic of functional pistils (no recognizable epidermis with stigmatic papillae, no transmitting tract tissue with specialized extracellular matrix, no ovule primordia or integuments) (3) pistil remnant often appearing as a simple dome or cone of cells without clear division into ovary, style, and stigma regions that characterize even rudimentary pistils (4) absence of any vascular tissue supplying the pistil remnant in many cases, or presence of only a minimal vascular strand terminating in undifferentiated tissue (5) well-developed stamens typical of staminate flowers (erect, 2.8-4.2 mm filaments, 0.6-0.95 mm anthers, 27,000-42,000 pollen grains per flower) completely dominating the flower morphology and overshadowing the minute pistil remnant. The vestigial pistil represents arrest of pistil development at extremely early stages, likely shortly after pistil primordium initiation and before significant organ elaboration, resulting in retention of minimal tissue representing the aborted developmental program but lacking any functional pistil characteristics. The size threshold distinguishing vestigial from rudimentary pistils is necessarily somewhat arbitrary but conventionally set at approximately 0.4 mm, with structures below this size generally showing minimal organ differentiation (vestigial) while structures above this size typically show recognizable though reduced organ components (rudimentary). Vestigial pistil flowers achieve absolute zero fruit set (0.0%) as the minute pistil remnant completely lacks all components necessary for reproduction= no receptive surface for pollen landing, no style for pollen tube growth, no ovary cavity, and no ovules for fertilization. The pistil remnant in vestigial pistil flowers appears to serve no reproductive function and likely represents developmental legacy tissue that has escaped complete elimination despite strong selection for pistil suppression in staminate flowers. The metabolic cost of maintaining vestigial pistils is minimal given their extremely small size (<1% of hermaphrodite pistil mass), likely explaining persistence despite non-functionality, as selection pressure for complete elimination may be weak when energetic savings would be trivial. Vestigial pistil flowers are common in wild dioecious V. vinifera ssp. sylvestris male populations, occurring in 30-50% of male individuals, with the remaining males split between male only flowers (complete pistil absence) and rudimentary pistil flowers (more obvious pistil remnants). The presence of vestigial pistils provides developmental genetic evidence that staminate flowers retain pistil developmental programming at the genetic level (pistil identity genes are presumably present and capable of initiating pistil primordia) but this programming is strongly suppressed by male-determining factors that prevent pistil elaboration beyond extremely early stages.
Vestigial_Pistil_Flower represents near-maximum pistil suppression while retaining minimal morphological traces and is distinguished from Male_Only_Flower (no recognizable pistil) by presence of minute tissue remnants and from Rudimentary_Pistil_Flower (recognizable organ components) by extreme size reduction and lack of organ differentiation. The boundary between vestigial and male only is subtle, as extremely minute vestigial remnants <0.2 mm can be easily overlooked, potentially leading to classification as male only when vestigial structures are actually present. The distinction between vestigial and rudimentary at approximately 0.4 mm reflects a natural inflection point where larger structures begin showing recognizable organ regions while smaller structures remain largely undifferentiated. Vestigial pistils provide evidence for developmental genetics studies of organ suppression, demonstrating that pistil developmental programs can be initiated but then arrested at very early stages through sex-determining genetic factors.
Blüte mit Pistillrudiment
Blüte mit Stempelrest
minimal-pistil flower
trace-pistil staminate flower
vestigiale Pistill Blüte
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 102-104 Caporali, E. et al. (2003). The arrest of development of abortive reproductive organs. Sexual Plant Reproduction 15=291-300 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644.
Vestigial Pistil Flower
Wild V. vinifera ssp. sylvestris female plants in Mediterranean riparian populations commonly exhibit vestigial stamen flowers with minute structures 0.25-0.4 mm long visible under 15× magnification as small undifferentiated tissue masses in stamen positions, surrounded by or adjacent to the prominent functional pistil (2.2-2.5 mm length, ovary 1.4-1.5 mm diameter), producing no pollen, achieving 30-36% fruit set through cross-pollination from nearby staminate males comparable to female only flowers despite vestigial stamen presence, and requiring careful examination to distinguish from female only types. Pistillate individuals in breeding populations from wild × cultivated crosses frequently show vestigial stamen flowers with stamen remnants 0.2-0.45 mm requiring microscopic examination for detection, functionally equivalent to female only flowers in absolute cross-pollination dependency and fruit set patterns but providing evidence of incomplete stamen suppression at the developmental level.
A vestigial stamen flower is a reduced stamen flower in which stamens are reduced to extremely small, barely recognizable remnants showing minimal organ differentiation, typically measuring 0.2-0.5 mm in total length and appearing as small undifferentiated protuberances or cellular masses in some or all of the five stamen positions with no clearly distinguishable filament or anther components, representing near-maximum stamen suppression while retaining minimal morphological evidence of stamen developmental initiation. Vestigial stamen flowers are characterized by= (1) extremely minute structures <0.5 mm long occupying some or all of the five positions where functional stamens would occur in hermaphrodite flowers, often requiring magnification (10-20×) for clear visualization (2) stamen remnants showing minimal cellular differentiation with largely undifferentiated parenchyma cells lacking the specialized tissue types characteristic of functional stamens (no recognizable epidermis with characteristic surface features, no vascular tissue, no anther wall layers, no pollen-producing tissue) (3) stamen remnants often appearing as simple protuberances or small cellular masses without clear division into filament and anther regions (4) complete absence of pollen production in all cases, as no functional anther tissue develops (5) well-developed functional pistil (2.1-2.6 mm length, ovary 1.3-1.6 mm diameter with 3-4 viable ovules, prominent receptive wet stigma) completely dominating the flower morphology. The vestigial stamen condition represents arrest of stamen development at extremely early stages, likely shortly after stamen primordium initiation and before significant organ elaboration, resulting in retention of minimal tissue representing the aborted developmental program but lacking any functional stamen characteristics. The size threshold distinguishing vestigial from rudimentary stamens is necessarily somewhat arbitrary but conventionally set at approximately 0.5 mm, with structures below this size generally showing minimal organ differentiation (vestigial) while structures above this size typically show recognizable though reduced filament and anther components (rudimentary). Vestigial stamen flowers achieve fruit set only through cross-pollination, with fruit set percentages (26-38% under good pollination conditions) comparable to female only flowers, as the minute stamen remnants produce no pollen and are incapable of any male reproductive function. The stamen remnants in vestigial stamen flowers appear to serve no reproductive function and likely represent developmental legacy tissue that has escaped complete elimination despite strong selection for stamen suppression in pistillate flowers. The metabolic cost of maintaining vestigial stamens is minimal given their extremely small size (<1% of hermaphrodite stamen mass), likely explaining persistence despite non-functionality. Vestigial stamen flowers are common in wild dioecious V. vinifera ssp. sylvestris female populations, occurring in 35-55% of female individuals, with the remaining females split between female only flowers (complete stamen absence) and rudimentary stamen flowers (more obvious stamen remnants). The presence of vestigial stamens provides developmental genetic evidence that pistillate flowers retain stamen developmental programming at the genetic level (stamen identity genes presumably present and capable of initiating stamen primordia) but this programming is strongly suppressed by female-determining factors that prevent stamen elaboration beyond extremely early stages.
Vestigial_Stamen_Flower represents near-maximum stamen suppression while retaining minimal morphological traces and is distinguished from Female_Only_Flower (no recognizable stamens) by presence of minute tissue remnants and from Rudimentary_Stamen_Flower (recognizable organ components) by extreme size reduction and lack of organ differentiation. The boundary between vestigial and female only is subtle, as extremely minute vestigial remnants <0.2 mm can be easily overlooked, potentially leading to classification as female only when vestigial structures are actually present. The distinction between vestigial and rudimentary at approximately 0.5 mm reflects a natural inflection point where larger structures begin showing recognizable filament and anther regions while smaller structures remain largely undifferentiated. Vestigial stamens provide valuable material for developmental genetics studies of organ suppression mechanisms, demonstrating that stamen developmental programs can be initiated but then arrested at very early stages through sex-determining genetic factors.
Blüte mit Staubblattrest
Blüte mit Staubblattrudiment
minimal-stamen flower
trace-stamen pistillate flower
vestigiale Staubblatt Blüte
Mullins, M.G., Bouquet, A., Williams, L.E. (1992). Biology of the Grapevine, pp. 103-105 Caporali, E. et al. (2003). The arrest of development of abortive reproductive organs. Sexual Plant Reproduction 15=291-300 Carbonneau, A. (1983). Stérilités mâle et femelle dans le genre Vitis. Agronomie 3=635-644.
Vestigial Stamen Flower
A vestigial tertiary reproductive bud at node 9 on Vitis vinifera cv. Gewürztraminer, dissected at 60x magnification revealing extremely small primordia (0.7mm) with what three expert observers disagreed about - two interpreting as purely vegetative with 3-4 leaf initials, one suggesting possible presence of 8-12 cells in unusual arrangement that might represent aborted inflorescence initial, with no consensus reached and practical significance zero since even if reproductive structure present, probability of successful activation, development, and fruit production under tertiary-requiring damage scenarios approaches zero, making the distinction between vestigial reproductive and purely vegetative tertiary buds academically interesting but operationally irrelevant.
A tertiary reproductive bud containing barely detectable traces of what might represent inflorescence primordial initiation, characterized by ambiguous structures that are difficult to distinguish from vegetative primordia under standard microscopic examination, questionable functional significance even if structures are truly reproductive in origin, and practical equivalence to purely vegetative tertiary buds in terms of developmental outcomes and crop contribution potential. Vestigial tertiary reproductive buds represent the absolute minimum of reproductive investment possible in compound bud architecture, possibly reflecting incomplete suppression of inflorescence differentiation pathways during tertiary bud formation or anatomical remnants of reproductive programs that failed to progress beyond initial cellular commitment, with their existence debated among researchers and their practical significance in viticulture essentially nonexistent.
Vestigial tertiary reproductive buds illustrate the challenges of definitive primordial classification at the extreme limits of bud development, where structures are so small and rudimentary that even experienced anatomists may disagree on interpretation. The difficulty distinguishes between incomplete vegetative primordia, failed reproductive initiation, and true if minimal reproductive structures raises questions about developmental thresholds and classification criteria. From practical viticulture perspective, the debate is moot - whether vestigial tertiaries contain traces of reproductive structures or not, their contribution to crop recovery following catastrophic damage is effectively zero, and management decisions treat all tertiary positions as non-fruiting regardless of anatomical minutiae. The category exists primarily for completeness in formal classification rather than practical vineyard application.
Spur-Tertiärknospe
marginale reproduktive Tertiärknospe
rudimentary tertiary inflorescence bud
trace reproductive tertiary bud
Vestigial Tertiary Reproductive Bud
Zinfandel grafted on St. George rootstock at 54 years post-grafting showing veteran old grafted rootstock trunk 13.2 cm diameter, trunk extending 29 cm from root crown to ancient graft union positioned 23 cm above soil, graft union completely invisible beneath 6.8 mm extremely thick weathered bark, veteran trunk bark showing very deep longitudinal fissures with nearly black coloration from extreme weathering, extensive lichen and moss covering 60% of bark surface, outer bark sloughing in plates, diameter growth 0.1 cm current season (minimal growth), cross-section showing 54 annual rings with recent 10-15 rings extremely narrow, extensive central heartwood 8.5 cm diameter with functional sapwood only 2.4 cm thickness, trunk structurally sound despite age, one old wound site showing limited compartmentalized decay, supporting sparse canopy 2.3 m shoot length, producing low but concentrated crop 3.8 kg with exceptional fruit quality characteristic of veteran old vine, vine showing clear aging but remaining viable
Veteran old grafted rootstock trunk is old grafted rootstock trunk in mid old phase typically years 40-60 post-grafting, characterized by many decades of stable integration, very thick heavily weathered bark showing extreme fissuring and weathering, large diameter typically 10-15 cm with notably slowed growth of 0.1-0.2 cm annually, extensive heartwood with significantly reduced sapwood, measurable but manageable vigor decline, and representing late but still functional productive life where trunk continues supporting production of potentially high-quality fruit though showing clear aging characteristics and potentially approaching eventual replacement consideration. Veteran old grafted rootstock trunk age and veteran status includes= spanning years 40-60 post-grafting typically (mid to late old trunk phase), representing very long-established vine (trunk having supported production through four to six decades), showing pronounced aging characteristics (clear indicators of advanced trunk age beyond early old phase), displaying measurable functional changes (noticeable reduction in growth rate and potential vigor compared to younger trunk), and marking veteran status (trunk qualifying as very old vine in most regional old vine definitions). Veteran old grafted rootstock trunk very old integration includes= showing 4-6 decade old graft union (callus tissue formed 40-60 years prior completely and permanently incorporated), displaying lifelong vascular continuity (xylem and phloem connections maintained for many decades though potentially with some age-related efficiency reduction), exhibiting absolutely stable graft union (bulge morphology completely static for decades with no changes in recent decades), showing completely invisible graft line (extremely thick veteran bark and decades-old tissue making grafting point identification impossible without extensive bark removal), and demonstrating exceptional proven durability (graft union having withstood 40-60 years of stress confirming remarkable long-term integration success). Veteran old grafted rootstock trunk pronounced physical characteristics includes= exhibiting large veteran diameter (10-15 cm typical range reflecting many decades of cumulative growth), showing very thick veteran bark (periderm 5-8 mm thickness representing decades of successive periderm formation), displaying extremely deep fissured patterns (longitudinal furrows reaching maximum depth with extensive weathering showing decades of exposure), revealing very dark heavily weathered coloration (nearly black bark in some areas showing extreme oxidation and environmental weathering), showing significant bark sloughing (outer bark separating in plates or strips due to age and weathering more pronounced than early old trunk), potentially showing bark damage areas (accumulated injury sites, dead bark patches, or exposed wood areas from decades of exposure and stress), and appearing impressively massive and ancient (veteran trunk conveying strong impression of great age and stability). Veteran old grafted rootstock trunk significantly slowed growth includes= showing notably reduced annual increase (0.1-0.2 cm typical veteran trunk growth significantly reduced from 0.3-0.6 cm mature trunk), displaying highly variable growth (annual increments inconsistent year to year with some years showing minimal or no measurable increase), exhibiting greatly reduced cambial vigor (vascular cambium activity substantially declined producing very narrow annual rings), forming many very narrow recent rings (recent growth rings in cross-section extremely narrow indicating severely slowed growth), and approaching potential growth cessation (some veteran trunks showing minimal annual diameter increase suggesting approaching end of active radial growth). Veteran old grafted rootstock trunk extensive heartwood formation includes= showing substantial heartwood core (extensive inner xylem converted to non-functional heartwood occupying majority of trunk cross-section), displaying markedly reduced functional sapwood (active water-conducting xylem limited to narrow outer ring typically outer 2-3 cm representing fraction of total trunk diameter), exhibiting large central heartwood (50-70% of trunk diameter converted to heartwood providing structural support but no transport function), showing clear heartwood-sapwood boundary (distinct color difference between dark central heartwood and lighter outer sapwood visible in cross-section), and maintaining minimally adequate transport (narrow sapwood ring sufficient for reduced canopy and crop of veteran vine though with limited reserve capacity). Veteran old grafted rootstock trunk measurable vigor decline includes= showing noticeable vegetative vigor reduction (shoot growth measurably reduced compared to mature trunk vigor with shorter weaker shoots), exhibiting reduced canopy density (leaf area noticeably reduced with sparser canopy than younger vine), displaying less vigorous shoot growth (annual shoot length substantially shorter than mature vine potentially 40-60% of mature vine shoot length), showing reduced reserve accumulation (carbohydrate storage notably reduced in aging trunk with limited capacity for stress recovery), and exhibiting potentially delayed phenology (bud break, bloom, or veraison potentially delayed compared to younger vines though pattern varying by site and cultivar). Veteran old grafted rootstock trunk altered production includes= generally showing reduced fruit production (veteran trunk typically supporting lower crop than mature vine potentially 50-70% of mature vine yield), often producing distinctive fruit quality (veteran vine fruit potentially showing concentrated flavors, balanced composition, and distinctive character valued in quality wine production), displaying smaller clusters and berries (natural yield reduction often manifested as reduced cluster and berry size contributing to concentration), exhibiting natural vine balance (reduced vigor and yield often resulting in excellent balance between canopy and crop), showing variable annual crops (year-to-year production potentially more variable in veteran vine than stable mature vine due to reduced reserves and stress sensitivity), and potentially demonstrating old vine terroir expression (veteran vine fruit potentially showing distinctive site characteristics and old vine typicity). Veteran old grafted rootstock trunk structural aging includes= generally maintaining structural integrity (most veteran trunks remaining mechanically adequate despite age), showing accumulated damage and wounds (decades of vineyard operations resulting in multiple scars, old pruning wounds, and injury sites), potentially exhibiting localized decay (some veteran trunks showing areas of wood rot particularly in old wound sites though often compartmentalized), displaying extensively healed old wounds (evidence of numerous past injuries successfully compartmentalized and occluded over many years), occasionally showing hollow areas (some veteran trunks developing limited internal hollows in central heartwood though usually maintaining adequate structural capacity), and requiring assessment (veteran trunk structural condition requiring evaluation to ensure continued stability and safety). Veteran old grafted rootstock trunk disease susceptibility includes= showing increased disease vulnerability (aging veteran trunk more susceptible to trunk diseases particularly in stressed or damaged areas), potentially exhibiting trunk diseases (Eutypa, Botryosphaeria, Esca, or other pathogens potentially affecting veteran trunk especially with inadequate management), displaying multiple old disease scars (evidence of past disease events managed and compartmentalized though potentially reducing functional tissue), showing reduced disease resistance (veteran trunk less able to mount vigorous defense responses compared to younger vine), requiring intensive disease monitoring (veteran trunk needing careful inspection and management to prevent disease progression threatening vine viability), and demonstrating resilience in well-managed vines (many veteran trunks remaining remarkably healthy despite decades of pathogen exposure when management appropriate). Veteran old grafted rootstock trunk management intensity includes= requiring careful assessment of viability (evaluating whether veteran trunk capable of continued production or approaching end of functional life), needing intensive disease and health monitoring (regular inspection critical for detecting problems before they threaten veteran vine survival), potentially benefiting from special cultural practices (modified irrigation, nutrition, or canopy management potentially helping maintain veteran vine health and productivity), considering trunk renewal or replacement decisions (determining whether to maintain veteran vine, implement renewal, or plan replacement), valuing fruit quality benefits (managing veteran vine to optimize old vine fruit quality potentially justifying continued cultivation despite reduced yields), and planning vineyard succession (developing long-term plan for veteran vineyard renewal or replanting as vines approach end of life). Veteran old grafted rootstock trunk as old vine designation includes= typically qualifying for old vine designation (veteran trunk age exceeding thresholds for old vine labeling in most regions and certification programs), representing valuable old vine vineyard (veteran vine plantings highly prized for fruit quality and old vine prestige), contributing significantly to vineyard heritage (veteran trunk representing living connection to vineyard history spanning multiple generations), demonstrating exceptional viticulture (maintaining healthy productive veteran vine through decades showing outstanding vineyard stewardship), potentially certified as heritage vines (some veteran vineyards eligible for historic vineyard designation or heritage vine certification), and commanding market premium (fruit and wine from veteran vines often commanding premium prices based on old vine reputation and quality). Veteran old grafted rootstock trunk diameter characteristics includes= typical veteran diameter 10-15 cm (mid to large old trunk size), vigorous rootstocks showing 14-20+ cm (1103P, 110R reaching very impressive dimensions in veteran status), moderate vigor rootstocks showing 10-14 cm (SO4, 3309C showing substantial veteran trunk), low vigor rootstocks showing 8-12 cm (Riparia types showing moderate veteran size even after decades), ancient largest veterans approaching 18-25 cm (oldest and most vigorous combinations reaching truly massive dimensions), and reflecting site and history (veteran trunk size reflecting decades of specific site conditions, rootstock genetics, and management history). Veteran old grafted rootstock trunk replacement and renewal considerations includes= weighing continued productivity against costs (evaluating whether veteran vine yield and quality justify continued maintenance versus replanting), assessing structural and health status (determining whether veteran trunk sound and healthy or showing problematic decay, disease, or declining health threatening viability), considering fruit quality premium (factoring whether old vine fruit quality and market value justify maintaining veteran vine despite challenges), evaluating trunk renewal feasibility (determining whether cordon renewal, head renewal, or other techniques could extend veteran vine productive life), planning replacement timing (if replacement decided, determining optimal schedule for replanting to minimize production loss and capture remaining veteran vine productivity), and valuing historical and cultural significance (considering vineyard heritage, old vine prestige, and historical value of veteran planting in decision-making). Veteran old grafted rootstock trunk in ancient vineyard context includes= characterizing ancient vineyard plantings (veteran trunks typical of rare ancient vineyards planted 40-60+ years ago), representing irreplaceable viticultural assets (veteran vine plantings impossible to replicate quickly given decades required to achieve veteran status), embodying vineyard history and tradition (veteran trunk representing living history connecting present viticulture to past generations), contributing to regional wine identity (veteran vineyards often fundamental to regional wine character and reputation), potentially designated as heritage or historic vineyard (some veteran vineyards protected or recognized for cultural and historical significance), and marking exceptional human achievement (maintaining productive veteran vine through many decades demonstrating remarkable dedication, skill, and stewardship). Veteran old grafted rootstock trunk resilience and adaptation includes= demonstrating long-term site adaptation (decades of growth showing how rootstock-scion combination suited and adapted to specific terroir), showing climate resilience (veteran trunk having survived and adapted to many years of climate variability including extreme events), exhibiting disease survival (veteran trunk demonstrating resistance or tolerance to endemic diseases encountered over decades), revealing management effects (veteran trunk characteristics reflecting cumulative effects of decades of specific cultural practices), and indicating sustainable viticulture (veteran trunk demonstrating grapevine potential for sustainable long-term production with appropriate care). Veteran old grafted rootstock trunk significance includes= representing exceptional vine longevity (4-6 decades of production demonstrating remarkable durability of grafted vine with appropriate management), showing advanced aging while maintaining function (veteran trunk demonstrating grapevine capacity to remain productive despite pronounced aging characteristics), embodying old vine mystique and quality (veteran trunk contributing to old vine reputation and perceived fruit quality benefits), requiring critical management decisions (veteran trunk forcing evaluation of continued value and appropriate management or replacement), marking rare achievement (veteran trunk representing comparatively rare accomplishment of maintaining productive vine for many decades), and demonstrating viticultural heritage (veteran trunk embodying accumulated knowledge, tradition, and connection to past generations of viticulturists).
Veteran_Old_Grafted_Rootstock_Trunk emphasizes mid to late old phase (years 40-60) and is distinguished from early Old_Grafted_Rootstock_Trunk (beginning old age 30-40 years) and Multi_Decade_Old_Grafted_Rootstock_Trunk (extremely ancient >60 years) by focus on pronounced advanced aging with measurable decline while maintaining production. Critical for understanding veteran vine characteristics, advanced aging indicators, old vine quality potential, management challenges, replacement considerations, and the veteran phase representing rare achievement of maintaining productive vine through many decades.
Langzeit-Veteranen-Pfropfstamm
Long-veteran grafted rootstock trunk
Sehr alter Wurzelstockstamm
Very old grafted rootstock trunk
Veteranen-verpfropfter Unterlagenstamm
Veteran Old Grafted Rootstock Trunk
Mourvèdre veteran old grafted scion trunk on 110R vigorous drought-tolerant rootstock in Bandol planted 1962 (64 years old) shows 295-340 mm scion diameter with 80% hollowed heartwood, graft union at 12 cm above soil with 128-142% swelling (increased significantly from prime maturity indicating age-related union stress), rootstock diameter 315-365 mm also extensively decayed, 15-22 mm functional scion sapwood with compromised union hydraulic conductance, 55-85 g combined depleted reserves, produces 1.2-2.5 kg exceptionally concentrated fruit from 4-10 sparse clusters, requires intensive care and structural support, survives in phylloxera-infested terroir due to rootstock resistance where own-rooted would be impossible
A veteran old grafted scion trunk is an old grafted scion trunk of exceptional age (typically 40-80+ years post-planting) that has survived decades beyond typical commercial grafted vineyard lifespan, characterized by very large scion diameter, advanced decay, severely declining vigor, and continued functionality primarily due to rootstock resilience supporting survival in conditions where own-rooted vines would have long since failed, representing exceptional longevity in grafted viticulture. Veteran old grafted scion trunk occurrence reflects= exceptional rootstock-scion compatibility (decades of successful integration without catastrophic incompatibility failure), rootstock benefits enabling survival (phylloxera resistance, drought tolerance, lime tolerance, or other rootstock adaptations critical for site longevity), intensive management maintaining veteran vines (careful pruning, disease management, structural support), optimal site conditions (deep soils, favorable climates supporting extended life), and deliberate preservation (heritage vineyards, genetic repositories, historical or cultural value motivating continued maintenance despite marginal economics). Veteran old grafted scion trunk anatomical characteristics include= very large scion diameter (typically 180-350 mm at 50-80 years on vigorous rootstocks, 150-280 mm on moderate, 120-240 mm on low-vigor), massive scion heartwood extensively decayed (central 70-90% showing wood decay, potential hollowing), minimal functional scion sapwood (10-20 mm radial depth providing limited water transport), very irregular growth rings (recent rings 0.2-2 mm, frequent missing rings), graft union showing age-related deterioration (union swelling may exceed 120-140% in some veteran combinations, callus tissue cracking or separating), thick rough scion bark (10-25 mm with deep fissures), and rootstock trunk also showing age (rootstock diameter 190-370 mm, similar decay and aging patterns). Veteran old grafted scion trunk graft union aging effects include= union potentially weakened by decades of stress (union swelling >125% common, mechanical strength declining), union vascular continuity potentially compromised (hydraulic conductance across union may decline 30-50% compared to prime maturity creating increasing bottleneck), union as major disease entry (high pathogen colonization of aging union tissues), union potential failure point (risk of union separation or catastrophic failure in very old combinations particularly with historical incompatible pairings), but survival indicating union adequacy (veteran status confirms decades of functional union despite aging). Veteran old grafted scion trunk physiological status shows= severely reduced water transport (limited sapwood and potential union bottleneck restricting hydraulic conductance), rootstock continuing to provide benefits (phylloxera resistance, drought tolerance, nutrient efficiency enabling survival despite scion decline), depleted combined reserves (scion and rootstock starch 40-100 g compared to 120-190 g in prime, insufficient for vigorous growth but adequate for survival), minimal vegetative growth (annual shoot growth 25-70 cm reflecting severe vigor decline), extended irregular phenology (budbreak, véraison highly delayed and asynchronous), and extreme stress sensitivity (despite rootstock benefits, veteran age creating vulnerability to drought, disease, environmental extremes). Veteran old grafted scion trunk productivity characteristics include= very low yields (typically 0.8-3.5 kg per vine, 15-40% of prime production, varying by rootstock class), sparse cluster number (2-15 clusters from limited fruitful shoots, rootstock vigor affecting upper range), very small clusters and berries (cluster 50-110 g, berry 8-13 mm diameter), extremely concentrated fruit (potentially exceptional quality with very high skin/pulp ratio, intense flavors, concentrated phenolics, rootstock quality effects still expressed at low intensities), and high vintage variation (productivity ranging from near-zero in stress years to moderate in favorable conditions). Veteran old grafted scion trunk rootstock significance in longevity includes= rootstock enabling survival impossible for own-rooted (phylloxera-infested soils supporting 50-80 year grafted vines where own-rooted would fail within 5-15 years post-infestation), rootstock stress tolerance supporting veteran status (drought-tolerant rootstocks like 110R, 140Ru, 1103P enabling survival in water-limited conditions, lime-tolerant rootstocks preventing chlorosis on calcareous soils), rootstock vigor potentially extending productive life (vigorous rootstocks maintaining minimal productivity 5-15 years longer than low-vigor equivalents), and rootstock as limiting factor (rootstock decline, root disease, or rootstock-soil mismatch potentially causing veteran vine death even if scion genetically capable of longer life). Veteran old grafted scion trunk viticulture significance includes= exceptional genetic value (veteran grafted vines representing rare heritage scion selections on proven rootstock combinations tested across decades), historical significance (vines planted 1940s-1980s representing viticultural heritage, particularly early grafting programs), potential exceptional quality (extremely concentrated fruit with rootstock-modulated terroir expression producing distinctive wines commanding premium pricing), research value (studying grafted vine extreme longevity, graft union aging, rootstock long-term effects), and preservation imperative (irreplaceable genetic and historical resources requiring specialized conservation). Veteran old grafted scion trunk management is highly specialized= minimal interventions avoiding stress (gentle pruning accounting for limited reserves), disease management accepting inevitable decay (focusing on preventing catastrophic collapse rather than cure), rootstock sucker vigilance (preventing rootstock canopy that would bypass declining scion), structural support for weakened trunks and unions (staking, propping preventing breakage), protective irrigation (minimal water during critical stress periods), and succession planning (propagating both scion and rootstock genetics before inevitable vine death, re-grafting selections onto appropriate rootstocks preserving combinations).
Veteran_Old_Grafted_Scion_Trunk emphasizes exceptional grafted longevity and is distinguished from Old_Grafted_Scion_Trunk (typical senescence) and Multi_Decade_Old_Grafted_Scion_Trunk (40-60 years) by extreme age 40-80+ years with rootstock critically enabling survival. Critical for understanding rootstock contributions to extreme vine longevity, heritage grafted vineyard preservation, rootstock selection impacts on multi-decade performance, and genetic conservation of rare scion-rootstock combinations.
Langlebiger Veredelungsstamm
Long-lived grafted scion
Sehr alter gepfropfter Stamm
Very old grafted trunk
Veteran-Aufpropfung
Veteran Old Grafted Scion Trunk
Chardonnay viable mature seeds from berries harvested at optimal ripeness (BBCH 87-88, 21°Brix) show 65-75% germination rates despite incomplete seed coat browning (70-85% brown coloration), tannin polymerization 55-65%, moisture content 38-42%
A viable mature seed is a mature seed of Vitis vinifera containing a living embryo capable of germination and seedling establishment following appropriate dormancy release treatments, representing seeds that have achieved sufficient developmental maturity to support successful propagation while potentially retaining some incomplete developmental characteristics. Viable mature seeds are defined primarily by functional germination capacity (typically 40-90% germination after stratification) rather than complete morphological or biochemical maturity. These seeds have completed critical developmental thresholds including= formation of viable embryo with differentiated cotyledons and radicle, sufficient endosperm accumulation to support germination and early seedling growth, development of basic desiccation tolerance, and acquisition of seed dormancy requiring cold stratification for release. However, viable mature seeds may exhibit incomplete development in some characteristics= partial seed coat lignification (60-90% brown coloration with some tan or green patches), intermediate tannin polymerization (50-70%), or slightly elevated moisture content (35-45%) compared to fully developed seeds. This category is particularly important for breeding programs and germplasm conservation, where the critical criterion is propagation capacity rather than complete maturity. In viticulture practice, viable mature seeds can occur in berries harvested at optimal wine grape maturity, representing seeds that achieved functional maturity without complete developmental completion. Seed viability assessment methods include= germination testing after stratification (90-120 days at 1-5°C), tetrazolium staining to assess embryo viability, X-ray imaging to detect embryo presence and size, and float testing (viable seeds typically sink in water while non-viable seeds float). The distinction between viable and fully developed mature seeds has implications for breeding efficiency= while fully developed seeds provide optimal germination rates and seedling vigor, viable seeds with incomplete development can still produce acceptable breeding populations, particularly when rare genotypes or limited seed availability necessitates use of sub-optimal material.
Viable_Mature_Seed emphasizes functional germination capacity and is distinguished from Fully_Developed_Mature_Seed (optimal complete maturation) by acceptance of incomplete developmental characteristics that do not prevent successful propagation. Critical for breeding program seed utilization decisions, germplasm conservation viability assessment, and understanding that harvest-maturity wine grapes may contain functionally viable but incompletely developed seeds.
Functional seed
Funktionsfähiger Traubensamen
Germinable seed
Keimfähiger Samen
Vermehrungsfähiger Kern
Bouquet, A., & Davis, H. P. (1989). Culture in vitro d'embryons immatures de vigne (Vitis vinifera L.) appliquée à la sélection de variétés de cuve sans pépins. Agronomie, 9(6), 565-574. | Ristic, R., & Iland, P. G. (2005). Relationships between seed and berry development of Vitis vinifera L. cv Shiraz= Developmental changes in seed morphology and phenolic composition. Australian Journal of Grape and Wine Research, 11(1), 43-58.
Viable Mature Seed
A 60-year-old Gamay vine in Beaujolais's granitic crus bears a Vieilles_Vignes_Role, sourced for a "Morgon Vieilles Vignes" bottling that commands 30-40% premium over the producer's standard Morgon based on the old vine designation. A 50-year-old Grenache vine in Châteauneuf-du-Pape serves in a Vieilles_Vignes_Role, contributing to a special "Cuvée Vieilles Vignes" that represents the estate's old-vine selection from its most venerable parcels. A 45-year-old Chenin Blanc vine in Vouvray bears a Vieilles_Vignes_Role, the designation distinguishing the producer's old-vine bottling from wines made from younger plantings.
An old vine role that inheres in a grapevine designated "Vieilles Vignes" (French: "old vines"), the traditional French term signaling advanced vine age and the quality associations attributed to old vine fruit. Vieilles Vignes designation is used on wine labels throughout France and French-influenced wine regions to indicate that grapes were sourced from old vines, though no legal definition specifies minimum age, leaving interpretation to individual producers. The term carries cultural weight in French winemaking tradition, where old vines are associated with lower yields, greater concentration, deeper terroir expression, and wines worthy of aging, justifying premium pricing and elevated critical regard. Vieilles Vignes status is typically claimed for vines exceeding 35-50 years, though some producers apply the term to younger vines while others reserve it for 60+ year plantings, creating market variability. The designation is particularly significant in regions where old vine survival is notable—Beaujolais (where phylloxera-resistant sandy granitic soils preserve old Gamay), Southern Rhône (ancient Grenache and Carignan bush vines), and Champagne (where old vine parcels are prized for reserve wines)—while less commonly emphasized in regions with shorter replanting cycles.
Vieilles_Vignes_Role captures the specifically French designation that has become internationally influential as a quality signifier. The term's lack of legal definition creates both flexibility (producers can self-determine qualification) and potential confusion (no guarantee of any specific minimum age), with market discipline through reputation and critical assessment partially constraining misuse. Vieilles Vignes labeling has increased substantially since the 1980s as producers recognized consumer willingness to pay premiums for old vine wines, with the designation now common across French appellations and adopted (in translation or directly) in other wine regions. The cultural meaning extends beyond objective age to encompass continuity with viticultural heritage, survival through historical challenges, and connection to traditional practices that modern replanting would sever. Critical evaluation of Vieilles Vignes claims requires producer-specific understanding the term is most meaningful from producers with documented old vine holdings and least meaningful when applied indiscriminately.
VV-Bezeichnung
Vieilles Vignes
alte Reben
old vines (French)
vieilles vignes designation
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Parker R (2005) The World's Greatest Wine Estates, Simon & Schuster Jefford A (2002) The New France, Mitchell Beazley Johnson H, Robinson J (2019) The World Atlas of Wine, 8th Edition, Mitchell Beazley
Vieilles Vignes Role
A Napa Valley Cabernet Sauvignon vineyard planted on deep alluvial soils with 110R rootstock exhibited poor vigor balance quality with Ravaz Index values of 3.2, indicating excessive vegetative growth relative to crop load. Implementation of regulated deficit irrigation reducing water application by 40% post-véraison combined with increased crop load through minimal cluster thinning improved vigor balance quality to Ravaz Index 7.5 over three seasons, with concurrent improvements in fruit phenolic concentration, reduced green character in resulting wines, and decreased powdery mildew pressure due to improved fruit zone light penetration.
Vigor balance quality is a viticultural quality that inheres in a bearing grapevine organism and represents the equilibrium state between vegetative growth capacity and reproductive output as manifested through the proportional allocation of photosynthetic resources between shoot extension, leaf area development, and fruit production. This quality reflects the fundamental physiological trade-off that governs grapevine productivity, wherein the vine must partition fixed carbon and mineral nutrients between competing sinks including actively growing shoot tips, expanding leaves, developing clusters, maturing berries, perennial storage organs, and root system maintenance. The ontological significance of vigor balance quality lies in its role as an integrative measure of whole-plant resource allocation that determines both current-season fruit quality and long-term vineyard sustainability. From a mechanistic perspective, vigor balance quality emerges from the interaction between genetically determined growth potential, environmental resource availability including water, nutrients, and light, and management-imposed constraints such as pruning severity, crop load adjustment, and canopy manipulation. Optimal vigor balance represents a cultivar-specific and site-specific equilibrium wherein sufficient leaf area exists to ripen the crop to target specifications while avoiding excessive vegetative growth that would compromise fruit exposure, increase disease pressure, and divert resources from berry development. The measurement of vigor balance quality requires assessment of both vegetative and reproductive parameters, with common approaches including the Ravaz Index relating yield to pruning weight, leaf area to fruit weight ratios, and visual assessment of shoot growth cessation timing relative to véraison. Balanced vines typically exhibit shoot growth cessation 2-3 weeks before véraison, moderate lateral development, and pruning weights of 0.3-0.6 kg per meter of canopy for vertical shoot positioned systems. Excessive vigor manifests as continued shoot growth through véraison, dense shading within the fruit zone, delayed and uneven ripening, herbaceous flavor development, and elevated disease incidence, while insufficient vigor produces premature growth cessation, leaf senescence before harvest, incomplete ripening, and reduced yields in subsequent seasons due to inadequate carbohydrate storage. The temporal dynamics of vigor balance quality reflect seasonal patterns wherein early-season vegetative dominance gradually shifts toward reproductive priority as fruit development creates increasingly powerful sink demand. Climate, rootstock selection, soil properties, and irrigation management fundamentally influence vigor balance quality, with warm climates and fertile soils promoting vegetative excess while water limitation and appropriate rootstock matching can optimize the vegetative-reproductive equilibrium for specific quality objectives.
Vigor_Balance_Quality serves as the parent class for specific balance assessment approaches including Vegetative_Reproductive_Balance_Quality, Vine_Balance_Index_Quality, and Point_Quadrat_Index_Quality. This class represents the overall equilibrium concept while child classes address particular measurement methodologies and balance dimensions. Distinguished from Vigor_Quality which addresses growth intensity per se, Vigor_Balance_Quality specifically concerns the proportional relationship between vegetative and reproductive function. Integration with Source_Sink_Balance_Quality reflects shared physiological mechanisms though different assessment perspectives.
Rebgleichgewicht
Wachstums-Ertrags-Balance
Wuchsbalance
growth-yield balance
vegetativ-reproduktives Gleichgewicht
vegetative-reproductive equilibrium
vine equilibrium state
vine vigor balance
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 12-31 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 327-345 Dry PR (2000) Canopy management for fruitfulness. Australian Journal of Grape and Wine Research 6:109-115
Vigor Balance Quality
Systematic quantification of Vigor_Conferred_To_Scion_Disposition at INRAE Bordeaux employed a factorial trial with 12 rootstocks grafted to a common scion ('Cabernet Sauvignon') grown under standardized conditions over 8 years, enabling isolation of rootstock effects on scion vigor. Measurements included scion pruning weight, shoot length, shoot number, trunk cross-sectional area, and leaf area per vine. Vigor_Conferred_To_Scion_Disposition showed wide variation among rootstocks. 'SO4' conferred highest vigor with scion pruning weight of 2.35 kg/vine (indexed as 145% relative to grand mean), shoot length of 195 cm, and trunk area of 12.8 cm². '110 Richter' and '140 Ruggeri' showed similarly high vigor conferral (138% and 142% respectively). 'Gravesac' conferred moderate vigor (102%). '101-14 Mgt' conferred substantially reduced vigor with pruning weight of 1.12 kg/vine (69% of mean), shoot length of 125 cm, and trunk area of 7.2 cm². '3309C' and 'Riparia Gloire' showed similar vigor-reducing effects (75% and 62% respectively). The effect was consistent across years (r > 0.90 for rootstock rankings between years) and when tested with alternative scions ('Merlot', 'Sauvignon Blanc'), confirming that Vigor_Conferred_To_Scion_Disposition is a stable rootstock characteristic. Correlation analysis revealed that vigor conferral was positively associated with root system size (r = 0.72), root hydraulic conductance (r = 0.68), and xylem cytokinin concentration (r = 0.58), indicating multiple contributing mechanisms. The range of vigor modulation (62-145% of mean) demonstrates that rootstock selection can nearly double or halve scion vigor expression.
A Vigor_Conferred_To_Scion_Disposition is a Vigor_Disposition that inheres in a Vitis rootstock and is realized through the rootstock's influence on the vegetative growth of the grafted scion, modifying scion shoot elongation, leaf area production, and overall biomass accumulation relative to what the scion would produce on its own roots or on a neutral rootstock. This disposition represents the rootstock's functional effect on scion vigor, integrating root system characteristics (size, distribution, efficiency) with physiological signaling (hormones, hydraulic conductance, nutrient supply) that together determine how the rootstock modulates scion growth expression. The vigor conferred to scion can range from substantial reduction (vigor-reducing rootstocks decreasing scion pruning weight by 30-50% relative to own-rooted vines) to substantial enhancement (vigor-enhancing rootstocks increasing pruning weight by 50-100%). This effect is relatively consistent across different scions—a vigor-reducing rootstock will reduce growth of both inherently vigorous and weak scions, though the absolute magnitude of reduction may differ. Vigor_Conferred_To_Scion_Disposition is only partially explained by Root_Vigor_Disposition (r ≈ 0.7), indicating that rootstock-scion signaling mechanisms contribute independently to vigor modulation. Mechanisms include regulation of cytokinin export from roots (promoting shoot growth), hydraulic conductance affecting water supply to shoots, and efficiency of nutrient uptake and translocation. Understanding Vigor_Conferred_To_Scion_Disposition enables strategic rootstock selection to achieve desired vine balance—pairing vigor-reducing rootstocks with vigorous scions or fertile sites, and vigor-enhancing rootstocks with weak scions or limiting sites.
Vigor_Conferred_To_Scion_Disposition is_a Vigor_Disposition representing rootstock influence on scion vigor. Effect ranges from ~60% to ~150% of neutral reference, enabling substantial vigor modulation through rootstock selection. Child classes include Vigor_Reducing_Rootstock_Disposition and Vigor_Enhancing_Rootstock_Disposition. Vigor conferral is partially but not fully explained by root system size (~70% correlation). Additional mechanisms include hydraulic conductance, cytokinin export, and nutrient supply efficiency. Vigor conferral is a stable rootstock characteristic consistent across scions and years.
Pfropfungs-Wuchskraft-Einfluss
Unterlage-Wuchskraft-Übertragung-Disposition
Unterlagen-Vigor-Effekt
grafting vigor influence
rootstock vigor effect
rootstock-conferred vigor
Ollat N et al. (2003) Grapevine rootstock effects on scion growth and mineral nutrition, Acta Horticulturae 617:47-54 Tandonnet JP et al. (2010) Long term effects of rootstock on vine performance and wine quality for Vitis vinifera cv. Cabernet Sauvignon, Australian Journal of Grape and Wine Research 16:298-306 Cookson SJ et al. (2013) Graft union formation in grapevine induces transcriptional changes related to cell wall modification, wounding, hormone signalling, and secondary metabolism, Journal of Experimental Botany 64:2997-3008
Vigor Conferred To Scion Disposition
Vigor data for Cabernet Sauvignon clone 337 at Rutherford, Napa Valley on 110R rootstock documents mean dormant pruning weight of 0.55 kg/m cordon (classified as moderate), average internode length of 6.5 cm, lateral shoot number of 2.8 per primary shoot, and leaf layer number (LLN) of 1.4 at véraison — an inherently balanced canopy structure requiring minimal hedging intervention. In contrast, clone 8 under identical conditions yielded pruning weight of 0.95 kg/m (high vigor), internode length of 8.2 cm, and LLN of 2.8, necessitating two hedging passes and aggressive lateral removal to achieve adequate fruit zone exposure. Vigor data for Riesling clone 198 at Geisenheim documents consistently low-moderate vigor (pruning weight 0.35-0.50 kg/m on 5C rootstock), supporting its recommendation for fertile loess soils where natural vigor control is essential for maintaining the Ravaz Index within the quality range of 6-8.
Vigor data is an agronomic performance data entity that records the vegetative growth intensity of a clonal line, quantified through shoot elongation rate, internode length, lateral shoot production, pruning weight per vine or per meter of cordon, trunk cross-sectional area increment, and canopy dimensional parameters (height, width, leaf layer number), measured across multiple vintages and expressed relative to evaluation site conditions and rootstock combination, characterizing the clone's inherent growth potential and its implications for training system selection, vine spacing, and canopy management requirements.
Vigor data is among the most management-relevant agronomic parameters because vine vigor directly determines canopy management labor requirements, which represent 30-50% of annual vineyard operating costs in premium production. The genetic component of vigor operates through differences in shoot apical meristem activity, hormone signaling (primarily gibberellin and cytokinin pathways), and carbon allocation patterns between vegetative and reproductive sinks. Pruning weight per meter of cordon is the most widely used and reproducible vigor metric, with generally accepted ranges of 0.3-0.6 kg/m for quality-oriented production, though optimal values vary with cultivar and climate. The clone-by-rootstock interaction for vigor is particularly strong and often non-additive — a low-vigor clone on a vigorous rootstock does not necessarily produce moderate total vigor, as rootstock growth promotion may overwhelm the scion's inherent restraint or, conversely, may fail to compensate for a weak scion. Remote sensing technologies including NDVI (Normalized Difference Vegetation Index) mapping from aerial or satellite imagery provide spatial vigor characterization that reveals within-block variability attributable to soil heterogeneity, rootstock variation, and localized pathogen effects, complementing the vine-level measurements from evaluation trials. Vigor trends over vine age should be documented, as some clones exhibit declining vigor with maturation (desirable for self-regulating canopy management) while others maintain juvenile vigor levels into advanced age.
Triebwachstumsdaten
Vitalitätsdaten
Wuchsstärkedaten
Wüchsigkeitsdaten
canopy vigor data
growth vigor data
vegetative Wachstumsdaten
vegetative growth data
vigor record
vine vigor assessment
Smart RE, Robinson MD (1991) Sunlight into Wine, Winetitles, pp.: 30-42 Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 195-225 Dry PR (2000) Canopy management for fruitfulness. Australian Journal of Grape and Wine Research 6(2): 109-115
Vigor Data
The vigor description of Mourvèdre documents high inherent vigor (OIV state 7) — under favorable growing conditions with adequate water and nutrition, Mourvèdre produces vigorous primary shoots of 1.5-2.5 m length with long internodes (80-110 mm), large leaves (blade length 180-220 mm), and strong lateral shoot development, generating a dense canopy that can exceed the optimal leaf area index of 3-5 m²/m² trellis by 50-100% without management intervention. This high vigor necessitates careful site selection (low-fertility soils, water-limiting conditions), moderate-vigor rootstocks (110R, 140Ru rather than the vigorous SO4 or 5BB), and aggressive summer canopy management (shoot thinning, hedging, leaf removal) to maintain the canopy openness required for fruit quality. The high vigor of Mourvèdre contrasts with the low vigor of Muscat à Petits Grains (OIV state 3, producing compact vines with short shoots and limited lateral development even under favorable conditions) and the medium vigor of Cabernet Sauvignon (OIV state 5, producing moderate growth that is readily managed on standard VSP trellising with bilateral cordon).
A growth habit description component that records the inherent vegetative vigor of the cultivar — the genetically determined potential for shoot elongation, leaf area production, and total biomass accumulation per vine — scored on a scale from very low (state 1) through low, medium, high, to very high (state 9), with vigor reflecting the cultivar-specific rates of cell division and expansion in shoot meristems that, modulated by rootstock, soil fertility, water availability, and crop load, produce the vine size and canopy density that dominate viticultural management decisions.
Vigor_Description documents the cultivar-specific growth potential that is the single most important factor determining the viticultural management intensity required — high-vigor cultivars demand more canopy management labor (shoot positioning, hedging, leaf removal, lateral removal) and more careful site-rootstock matching to achieve the balanced vine growth that produces high-quality fruit, while low-vigor cultivars may struggle to fill the allocated trellis space and produce adequate yields on vigorous sites. The genetic basis of vigor involves multiple interacting physiological systems — shoot apical meristem activity (determining the rate of new phytomer initiation), gibberellin-mediated internode elongation (determining the length of each phytomer), leaf expansion rate and final size (determining the photosynthetic source capacity), and root system architecture and hydraulic conductance (determining the water and nutrient supply capacity): cultivar differences in any of these systems contribute to the overall vigor phenotype, and the polygenic nature of vigor determination explains the continuous distribution of vigor across the V. vinifera cultivar spectrum rather than discrete vigor classes. The critical interaction between cultivar vigor and site vigor (determined by soil depth, water-holding capacity, fertility, and rootstock) defines the concept of vine balance — the ratio of vegetative growth to reproductive growth that, when optimized at approximately 1.0-1.5 kg pruning weight per kg fruit weight (the Ravaz index of 5-8), produces the moderate vine size, adequate but not excessive canopy density, and well-exposed fruit zone associated with high wine quality.
Triebstärke-Charakterisierung
Wüchsigkeits-Beschreibung
growth potential characterization
vegetative Wachstumskraft
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Edition Smart RE and Robinson M (1991) Sunlight into Wine, Winetitles Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Kliewer WM and Dokoozlian NK (2005) Leaf area/crop weight ratios of grapevines: influence on fruit composition and wine quality, American Journal of Enology and Viticulture 56: 170-181
Vigor Description
Systematic evaluation of Vigor_Disposition at INRAE Bordeaux characterized vegetative growth across 80 cultivars and 20 rootstocks in a factorial trial assessing both scion and rootstock contributions to vigor expression. Standardized measurements of pruning weight per meter of cordon, shoot length, shoot number per meter, and Ravaz index (yield/pruning weight ratio) were collected over five seasons. Vigor_Disposition showed wide variation among scion cultivars, with pruning weights ranging from 0.4 kg/m ('Gewürztraminer' on '3309C') to 2.8 kg/m ('Carignane' on '110R'). Rootstock effects were substantial, with the same scion showing up to 2-fold variation in pruning weight depending on rootstock—'Cabernet Sauvignon' produced 0.9 kg/m on '101-14 Mgt' (devigorating) versus 1.8 kg/m on '110 Richter' (vigorating). However, scion genotype explained 65% of vigor variation while rootstock explained 25% and scion × rootstock interaction 10%, confirming that Vigor_Disposition is primarily a scion characteristic modulated by rootstock. Classification of cultivars by Vigor_Disposition revealed that vigor correlated with geographic origin—Mediterranean cultivars ('Carignan', 'Grenache', 'Mourvèdre') showed higher vigor than continental European cultivars ('Pinot Noir', 'Riesling', 'Gewürztraminer'), potentially reflecting adaptation to longer growing seasons and higher evaporative demand. Vigor_Disposition was partially independent of yield potential (r = 0.45), indicating that some high-vigor cultivars channel growth into vegetative rather than reproductive structures.
A Vigor_Disposition is a Growth_And_Development_Disposition that inheres in a Vitis vinifera plant and is realized through the inherent capacity for vegetative growth, determining the rate and extent of shoot elongation, leaf area production, lateral branching, and overall biomass accumulation during the growing season. This disposition represents the fundamental growth intensity of the genotype, reflecting the integrated activity of shoot apical meristems, cambial growth, and the mobilization and allocation of resources (carbon, nutrients, water) toward vegetative structures. Vigor is expressed through multiple measurable parameters including shoot elongation rate (cm/day during active growth), final shoot length, internode length, leaf size, lateral shoot production, trunk diameter increment, and total pruning weight (the most commonly used integrative measure). High vigor indicates strong inherent growth capacity producing abundant vegetative biomass, while low vigor indicates restrained growth with limited biomass production. Vigor_Disposition has profound viticultural implications, as it determines training system suitability (high-vigor cultivars requiring extensive canopy division or large canopy systems), planting density (low-vigor cultivars tolerating closer spacing), canopy management intensity (high vigor requiring more hedging, shoot thinning, and leaf removal), and the vine's capacity to ripen fruit while supporting adequate leaf area. The vigor expressed in any season represents the interaction between Vigor_Disposition (genetic potential) and environmental/management factors, but genotypic vigor rankings remain relatively stable across conditions.
Vigor_Disposition is_a Growth_And_Development_Disposition. Vigor represents growth intensity/capacity, the most fundamental growth characteristic. Pruning weight is the standard integrative measure of vigor expression. Child classes include Shoot_Vigor_Disposition and potentially Root_Vigor_Disposition. Vigor is primarily scion-determined but significantly modulated by rootstock (25% of variance). High vigor creates canopy management challenges low vigor may limit fruit ripening capacity. Vigor interacts with but is partially independent of yield potential.
Vegetative-Wuchskraft
Wachstums-Intensität
Wuchsstärke-Disposition
growth capacity
growth intensity
vegetative vigor
Howell GS (2001) Sustainable grape productivity and the growth-yield relationship: a review, American Journal of Enology and Viticulture 52:165-174 Dry PR et al. (2010) Canopy management for fruitfulness, Australian Journal of Grape and Wine Research 16:47-52 Keller M et al. (2004) Warm spring temperatures induce persistent season-long changes in shoot development in grapevines, Annals of Botany 93:739-746
Vigor Disposition
Documentation of Vigor_Enhancing_Rootstock_Disposition at the Australian Wine Research Institute characterized the vigor-promoting effects of rootstocks recognized for scion growth enhancement. A trial with 18 rootstocks grafted to 'Shiraz' over 12 years quantified vigor parameters and identified mechanisms of vigor enhancement. Rootstocks classified as vigor-enhancing showed consistent characteristics. '110 Richter' exhibited strong Vigor_Enhancing_Rootstock_Disposition with scion pruning weight of 2.45 kg/vine (142% of trial mean), shoot length of 215 cm (138% of mean), and trunk cross-sectional area of 14.2 cm² (155% of mean). Root system excavation revealed total root length of 2,950 m with maximum depth of 2.8 m, indicating extensive Root_Vigor_Disposition as a primary mechanism. 'SO4' showed similar vigor enhancement (148% pruning weight) with somewhat shallower but dense root system (2,680 m, 1.9 m depth). '140 Ruggeri' displayed the strongest vigor enhancement (156% pruning weight) associated with the most extensive root development (3,280 m, 3.2 m depth). Physiological analysis revealed that Vigor_Enhancing_Rootstock_Disposition rootstocks showed 55% higher xylem sap cytokinin concentration and 45% higher root hydraulic conductance compared to vigor-reducing types. Canopy management requirements were substantially increased—vigor-enhancing rootstock vines required 3-4 hedging passes per season versus 1-2 for vigor-reducing, and fruit zone leaf layer number averaged 3.2 versus 1.8. However, drought resilience was substantially enhanced, with vigor-enhancing rootstock vines maintaining favorable water status 22 days longer during dry-down due to access to deeper soil water reserves.
A Vigor_Enhancing_Rootstock_Disposition is a Vigor_Conferred_To_Scion_Disposition that inheres in a Vitis rootstock and is realized through mechanisms that promote scion vegetative growth, increasing shoot elongation, leaf area production, and pruning weight relative to own-rooted vines or vines grafted to vigor-neutral rootstocks. This disposition represents rootstock genotypes that amplify scion vigor, typically increasing pruning weight by 30-60% compared to mean rootstock performance, achieved through extensive root systems, high hydraulic conductance, enhanced cytokinin export, and efficient nutrient acquisition. Vigor-enhancing rootstocks are strategically valuable for supporting inherently weak scions, overcoming growth limitations on poor or shallow soils, achieving adequate canopy development in challenging environments, and maximizing vine establishment vigor. The vigor enhancement effect is generally proportional across scions—a vigor-enhancing rootstock will increase growth of any scion, though inherently high-vigor scions may become excessively vigorous requiring intensive management. Vigor-enhancing rootstocks are typically characterized by V. berlandieri and V. rupestris parentage, which contribute extensive root systems and robust root-to-shoot signaling. The benefits of vigor enhancement may be offset by canopy management challenges on favorable sites, delayed fruit maturity due to vegetative competition, and potentially reduced fruit quality from excessive shading in dense canopies.
Vigor_Enhancing_Rootstock_Disposition is_a Vigor_Conferred_To_Scion_Disposition representing vigor-promoting rootstocks. Typical vigor enhancement is 30-60% above mean pruning weight. Examples include '110 Richter', '140 Ruggeri', '1103 Paulsen', 'SO4', 'Ramsey', '99 Richter'. V. berlandieri and V. rupestris parentage is characteristic. Mechanisms include extensive root system, high hydraulic conductance, enhanced cytokinin export. Advantages include support for weak scions, drought resilience, and performance on limiting sites. Trade-offs include canopy management challenges and potential quality reduction from excessive vigor. Strategic use with low-vigor scions or limiting sites.
Stärker-Wachsende-Unterlage
Wuchskraft-Förderndes-Unterlagen-Disposition
Wuchskraft-Steigernde-Unterlage
vigor-promoting rootstock
Walker RR et al. (2002) Rootstock effects on salt tolerance of irrigated field-grown grapevines, Australian Journal of Grape and Wine Research 8:146-155 Tandonnet JP et al. (2010) Long term effects of rootstock on vine performance and wine quality for Vitis vinifera cv. Cabernet Sauvignon, Australian Journal of Grape and Wine Research 16:298-306 Zhang L et al. (2016) The effect of rootstock on berry composition and aroma profile of Shiraz, American Journal of Enology and Viticulture 67:311-320
Vigor Enhancing Rootstock Disposition
End-of-season assessment of Vigor_Expression_Quality across a variable Cabernet Sauvignon vineyard reveals pruning weights ranging from 0.3 kg/vine on shallow, rocky soils to 1.8 kg/vine on deep alluvial deposits, guiding the implementation of variable-rate balanced pruning with 15 nodes retained per vine on low-vigor zones versus 40 nodes per vine on high-vigor zones to achieve uniform crop load to canopy ratios and consistent wine quality across the heterogeneous site.
Vigor_Expression_Quality is a Vigor_Quality that inheres in a grapevine (Vitis vinifera L.) and represents the accumulated biomass outcomes of vegetative growth as measured through structural dimensions and mass of woody tissue retained through the dormant season. This quality captures the integrated result of seasonal growth vigor — the permanent vegetative increment that remains after leaf fall and represents the vine's net carbon investment in structural growth during the preceding season. In BFO terms, Vigor_Expression_Quality exists as a mass-related and dimensional quality complex measurable during the dormant period, providing a retrospective assessment of vigor status that complements the real-time measurements captured by Vegetative_Growth_Vigor_Quality. The distinction between growth vigor and expressed vigor is viticulturally significant because the same growth rate can produce different biomass accumulation depending on growth duration, lateral development, and shoot maturation processes, while identical final biomass can result from dramatically different growth trajectories with distinct implications for fruit quality. Pruning weight measurement represents the standard viticultural assessment of vigor expression, capturing the total mass of one-year-old wood (canes) removed during winter pruning, with values varying from less than 300 g per vine in severely stressed or devigorated conditions to over 3 kg per vine in excessively vigorous situations. The relationship between vigor expression and fruit quality follows the principle of vine balance, with optimal wine quality typically associated with intermediate vigor expression levels that maintain adequate photosynthetic capacity without creating the dense canopies and competitive sink relationships that compromise fruit composition. Vigor expression measurements form the basis for pruning decisions in balanced pruning systems, where node retention is adjusted proportionally to pruning weight to maintain consistent crop load to canopy ratios across vines varying in vigor status.
Vigor_Expression_Quality serves as the parent class for biomass-based vigor assessments including cane diameter, pruning weight, and pruning weight per unit canopy length. These measurements share the characteristic of being available only post-dormancy, providing retrospective vigor assessment that guides subsequent season management decisions. The class relates to vine balance concepts through the Ravaz Index (yield/pruning weight) and to nutrient management through the understanding that pruning weight reflects season-long nutrient availability and uptake.
Biomasseproduktion
Holzertrag
Wuchsausdruck
accumulated vigor
akkumulierte Wuchskraft
biomass vigor
expressed vigor
structural vigor
Bravdo B, Hepner Y, Loinger C, Cohen S, Tabacman H (1985) Effect of crop level and crop load on growth, yield, must and wine composition, and quality of Cabernet Sauvignon. American Journal of Enology and Viticulture 36:125-131 Howell GS (2001) Sustainable grape productivity and the growth-yield relationship: a review. American Journal of Enology and Viticulture 52:165-174
Vigor Expression Quality
Comparison of vigor manipulation response quality across five red Bordeaux cultivars in Napa Valley evaluated lateral regrowth following standardized hedging at 15 nodes post-véraison. Cabernet Sauvignon exhibited the strongest response with mean 4.2 lateral shoots per hedged primary and 28 cm mean lateral length at harvest, while Merlot showed intermediate response with 2.8 laterals averaging 19 cm, and Cabernet Franc demonstrated lowest vigor manipulation response quality with 1.9 laterals averaging 12 cm, informing cultivar-specific recommendations for hedging timing and the potential need for follow-up lateral removal in high-response varieties.
Vigor manipulation response quality is a viticultural quality that inheres in a grapevine organism and represents the characteristic manner and magnitude of vegetative growth adjustment exhibited following deliberate management interventions designed to modify canopy development, shoot extension, or overall vine vigor through physical removal, restriction, or redirection of growing tissues. This quality captures the dynamic responsiveness of grapevines to canopy management operations and determines the effectiveness of vigor control strategies in achieving desired canopy architecture, fruit exposure, and vine balance outcomes. The ontological significance of vigor manipulation response quality lies in its role as a phenotypic expression of the interaction between genetically determined growth potential, current physiological status, environmental conditions, and the specific nature and timing of management interventions, fundamentally influencing the practical utility of various canopy management techniques across different production contexts. Grapevines exhibit characteristic compensatory growth responses to tissue removal that reflect the disruption of established source-sink relationships and hormonal balances, with the removal of actively growing shoot tips releasing apical dominance and stimulating lateral bud development while removal of mature leaf area can trigger mobilization of carbohydrate reserves and altered root-shoot allocation patterns. The magnitude and character of vigor manipulation responses vary systematically with cultivar genetics, wherein varieties with strong apical dominance and high intrinsic vigor such as Cabernet Sauvignon and Syrah typically exhibit more pronounced compensatory responses than lower-vigor cultivars, requiring repeated interventions to maintain canopy targets. Timing relative to phenological development critically modulates response characteristics, with early-season interventions during active vegetative growth provoking stronger compensatory responses than operations conducted after natural growth cessation approaching véraison. Environmental factors including water availability, nitrogen status, and temperature regime further condition vigor manipulation responses, with well-watered, high-fertility conditions amplifying regrowth potential while moderate stress can dampen compensatory responses and improve intervention persistence. The measurement of vigor manipulation response quality requires assessment of regrowth characteristics following intervention, including lateral shoot number and length, secondary growth rate, and the duration of response persistence before canopy stabilization. Understanding cultivar-specific and site-specific vigor manipulation response patterns enables optimization of intervention timing, intensity, and frequency to achieve efficient canopy management with minimal input requirements while avoiding excessive compensatory growth that can negate intervention benefits.
Vigor_Manipulation_Response_Quality serves as the parent class for specific intervention response types including Topping_Response_Quality, Hedging_Response_Quality, and Shoot_Thinning_Response_Quality. This class captures the general concept of vine responsiveness to vigor-modifying interventions while child classes address particular management operations. Distinguished from Vigor_Quality which addresses growth intensity per se, this class specifically concerns the dynamic response to deliberate manipulation. Part of the broader Management_Responsive_Quality domain within Viticultural_Quality.
Eingriffs-Antwortverhalten
Laubarbeitsreaktion
Wuchskontrollreaktion
Wuchsmanipulationsreaktion
growth intervention response
management responsiveness
vigor control response
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 64-85 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-325 Reynolds AG, Vanden Heuvel JE (2009) Influence of grapevine training systems on vine growth and fruit composition: a review. American Journal of Enology and Viticulture 60:251-268
Vigor Manipulation Response Quality
101-14 Millardet et de Grasset (V. riparia × V. rupestris) confers low vigor to Merlot in Bordeaux, reducing pruning weights to 0.3-0.5 kg/m compared to 0.6-0.9 kg/m on the high-vigor Gravesac (161-49C × 3309C), achieving Ravaz indices of 5-8 considered optimal for premium red wine production. In Napa Valley, St. George (V. rupestris) confers very high vigor to Cabernet Sauvignon with pruning weights reaching 1.0-1.5 kg/m, while Riparia Gloire (V. riparia) limits growth to 0.2-0.4 kg/m pruning weight under similar conditions.
A rootstock role that inheres in a grafted rootstock by virtue of its capacity to modulate the vegetative growth vigor of the scion through differential root system architecture, hydraulic conductance, nutrient uptake efficiency, and hormonal signaling. Vigor modification is quantified relative to own-rooted Vitis vinifera performance, with rootstocks classified along a continuum from strongly dwarfing (reducing scion pruning weight by 40-60%) to highly invigorating (increasing scion pruning weight by 30-50% or more). The vigor-modifying effect is mediated by root density (typically 0.5-4.0 km root length per m³ soil), root hydraulic conductance, cytokinin-to-ABA ratio in xylem sap, and total absorptive root surface area. Vigor modification fundamentally determines canopy size, fruit-to-vegetative growth balance, microclimate within the fruiting zone, and ultimately wine quality potential.
Vigor_Modifying_Rootstock_Role is subclassed into High_Vigor_Rootstock_Role, Moderate_Vigor_Rootstock_Role, Low_Vigor_Rootstock_Role, and Dwarfing_Rootstock_Role. Vigor modification is context-dependent and interacts with soil type, water availability, scion cultivar, and climate the same rootstock may confer different relative vigor under different conditions. This role captures the functional vigor effect, while the underlying physiological mechanisms are represented as dispositions in the Grapevine_Disposition hierarchy.
Wuchskraft modifizierende Unterlage
growth-regulating rootstock
vigor-controlling rootstock
vigor-modifying rootstock
wuchsregulierende Unterlage
wuchssteuernde Unterlage
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 285-310 Ollat N et al. (2003) Effects of rootstock genotype on vine vigor, must composition and wine quality of Vitis vinifera cv. Cabernet Sauvignon, OENO One 37(2): 69-76 Tandonnet JP et al. (2010) Long term effects of genetic and environmental factors on grapevine rootstock growth, Australian Journal of Grape and Wine Research 16(2): 327-339
Vigor Modifying Rootstock Role
A Napa Valley Cabernet Sauvignon vineyard on deep alluvial soils with 1103 Paulsen rootstock exhibits excessive Vigor_Quality with pruning weights exceeding 1.5 kg per meter of canopy, requiring aggressive shoot thinning, hedging, and leaf removal to achieve acceptable fruit exposure, while an adjacent block on shallow, well-drained hillside soils with 101-14 rootstock demonstrates naturally balanced vigor with pruning weights of 0.4 kg/m, minimal canopy management requirements, and consistently superior wine quality ratings.
Vigor_Quality is a Viticultural_Quality that inheres in a grapevine (Vitis vinifera L.) and represents the overall capacity and expression of vegetative growth potential as manifested through shoot extension, biomass accumulation, and developmental intensity throughout the growing season. This quality encompasses the fundamental biological drive toward vegetative expansion that characterizes each vine's inherent growth characteristics as modified by environmental conditions, cultural practices, and rootstock-scion interactions. In the BFO framework, Vigor_Quality exists as a specifically dependent continuant that cannot exist independently of its bearer vine, reflecting the integrated physiological state arising from carbohydrate reserves, hormonal regulation, water and nutrient availability, and genetic predisposition. The concept of vine vigor represents one of the most critical parameters in viticultural management, directly influencing the balance between vegetative and reproductive development that ultimately determines fruit quality and wine character. High-vigor vines characteristically produce excessive shoot growth, dense canopies with poor light penetration, increased disease pressure, delayed fruit ripening, and wines with herbaceous or vegetal characteristics, while appropriately managed vigor promotes optimal source-sink relationships, uniform fruit maturation, and balanced wine composition. Vigor expression varies dramatically across cultivars, with varieties like Cabernet Sauvignon and Syrah exhibiting inherently high vigor requiring careful management, whereas Pinot Noir and Grenache display moderate vigor more amenable to quality wine production without intensive canopy manipulation. Rootstock selection profoundly influences scion vigor, with high-vigor rootstocks such as 1103 Paulsen and 140 Ruggeri promoting vigorous growth suitable for challenging conditions, while devigorating rootstocks like 101-14 Millardet and Riparia Gloire restrict vegetative development, concentrating vine resources into fruit production. The terroir concept fundamentally incorporates vigor regulation, as premier vineyard sites typically impose natural vigor limitation through water stress, nutrient restriction, or soil constraints that channel vine energy toward reproductive rather than vegetative development. Precision viticulture increasingly employs vigor mapping through NDVI (Normalized Difference Vegetation Index) and other remote sensing technologies, enabling site-specific management interventions targeting vigor zones within heterogeneous vineyard blocks. Understanding and manipulating vine vigor represents perhaps the single most important viticultural skill, as appropriate vigor management underlies virtually every aspect of quality wine production from pruning decisions through harvest timing.
Vigor_Quality serves as the primary parent class for all vigor-related qualities in VVO, positioned under Viticultural_Quality to reflect its central importance in vineyard management and wine quality determination. The class encompasses both inherent genetic vigor potential and expressed vigor as modified by environmental and cultural factors, with child classes differentiating between growth process measurements (Vegetative_Growth_Vigor_Quality) and accumulated biomass indicators (Vigor_Expression_Quality). This distinction parallels the viticultural practice of assessing vigor both during the growing season through shoot growth observations and post-dormancy through pruning weight measurements.
Rebenwuchskraft
Triebigkeit
Wuchskraft
Wuchsstärke
grapevine vigor
vine vigor
vine vigour
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd ed. Academic Press, pp. 267-312 Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 12-35 Dry PR, Loveys BR (1998) Factors influencing grapevine vigour and the potential for control with partial rootzone drying. Australian Journal of Grape and Wine Research 4:140-148
Vigor Quality
Characterization of Vigor_Reducing_Rootstock_Disposition at Geisenheim University documented the vigor-limiting effects of rootstocks recognized for scion growth reduction. A trial with 15 rootstocks grafted to 'Riesling' over 10 years quantified vigor parameters and identified mechanisms of vigor reduction. Rootstocks classified as vigor-reducing showed consistent characteristics. '101-14 Mgt' exhibited archetypal Vigor_Reducing_Rootstock_Disposition with scion pruning weight of 0.68 kg/vine (65% of trial mean), shoot length of 118 cm (72% of mean), and trunk cross-sectional area increment of 1.2 cm²/year (58% of mean). Root system excavation revealed total root length of 1,180 m and maximum depth of 1.4 m, indicating limited Root_Vigor_Disposition as a primary mechanism. '3309C' showed similar vigor reduction (72% pruning weight, 75% shoot length) with intermediate root system size (1,520 m). 'Riparia Gloire' displayed the strongest vigor reduction (58% pruning weight) associated with minimal root development (980 m total length). Physiological analysis revealed that Vigor_Reducing_Rootstock_Disposition rootstocks showed 35% lower xylem sap cytokinin concentration and 28% lower root hydraulic conductance compared to vigor-enhancing types, implicating both hormonal and hydraulic mechanisms. Ravaz index analysis confirmed improved vine balance on vigor-reducing rootstocks (Ravaz 8.5-10.0 versus 5.5-7.0 on vigor-enhancing), indicating that growth reduction shifted allocation toward reproduction. However, drought sensitivity was elevated, with vigor-reducing rootstock vines showing stress symptoms 15 days earlier during controlled dry-down.
A Vigor_Reducing_Rootstock_Disposition is a Vigor_Conferred_To_Scion_Disposition that inheres in a Vitis rootstock and is realized through mechanisms that limit scion vegetative growth, reducing shoot elongation, leaf area production, and pruning weight relative to own-rooted vines or vines grafted to vigor-neutral rootstocks. This disposition represents rootstock genotypes that naturally constrain scion vigor, typically reducing pruning weight by 25-45% compared to mean rootstock performance, achieved through limited root system size, reduced hydraulic conductance, lower cytokinin export, or some combination of these factors. Vigor-reducing rootstocks are strategically valuable for managing inherently vigorous scions, controlling excess growth on fertile or well-watered sites, achieving balanced canopies without intensive management intervention, and enabling closer vine spacing in high-density planting systems. The vigor reduction effect is generally proportional across scions—a vigor-reducing rootstock will reduce growth of any scion, though inherently low-vigor scions may become excessively weak requiring management compensation. Vigor-reducing rootstocks are typically characterized by V. riparia parentage, which contributes compact root systems and lower root-to-shoot signaling intensity. The trade-off for vigor reduction is often increased sensitivity to drought and other stresses due to limited root system buffering capacity, and potentially reduced vine longevity due to smaller carbohydrate reserves.
Vigor_Reducing_Rootstock_Disposition is_a Vigor_Conferred_To_Scion_Disposition representing vigor-limiting rootstocks. Typical vigor reduction is 25-45% of pruning weight versus mean. Examples include '101-14 Mgt', '3309C', 'Riparia Gloire', '420A', '161-49C'. V. riparia parentage is characteristic. Mechanisms include limited root system, reduced hydraulic conductance, lower cytokinin export. Advantages include vigor control and improved vine balance. Trade-offs include increased drought sensitivity and potentially reduced longevity. Strategic use with vigorous scions or fertile sites.
Schwächer-Wachsende-Unterlage
Wuchskraft-Limitierende-Unterlage
Wuchskraft-Reduzierendes-Unterlagen-Disposition
vigor-limiting rootstock
Keller M et al. (2012) Interactive effects of rootstock and deficit irrigation on grapevine physiology and fruit quality, American Journal of Enology and Viticulture 63:303-312 Ollat N et al. (2003) Grapevine rootstock effects on scion growth and mineral nutrition, Acta Horticulturae 617:47-54 Harbertson JF, Keller M (2012) Rootstock effects on deficit-irrigated red winegrapes, American Journal of Enology and Viticulture 63:80-89
Vigor Reducing Rootstock Disposition
Analysis of vigor uniformity quality across a 25-hectare Coonawarra Cabernet Sauvignon vineyard using NDVI mapping revealed coefficient of variation of 18.7% with distinct low-vigor zones associated with shallow terra rossa over limestone and high-vigor zones corresponding to deeper soil profiles over clay subsoils. Implementation of variable-rate irrigation based on vigor zone delineation improved vigor uniformity quality to CV 11.2% over three seasons while simultaneously reducing total water application by 23% through avoiding over-irrigation of inherently low-vigor zones where additional water could not overcome soil depth limitations.
Vigor uniformity quality is a viticultural quality that inheres in a grapevine population or vineyard unit and represents the degree of consistency in vegetative growth characteristics across individual vines, vine sections, or spatial zones within the production area as manifested through variation in shoot length, cane diameter, leaf area, pruning weight, and canopy dimensions. This quality reflects the homogeneity of growing conditions and vine responses that determines the feasibility of uniform management practices and the consistency of fruit quality at harvest. The ontological significance of vigor uniformity quality lies in its role as a population-level or spatial property that emerges from the interaction between environmental heterogeneity, planting material variation, and historical management effects, fundamentally influencing both operational efficiency and product quality consistency. High vigor uniformity quality indicates that vines across the assessed unit respond similarly to environmental conditions and management inputs, enabling standardized practices including uniform irrigation scheduling, consistent canopy management timing, and single-pass mechanical operations without compromising individual vine optimization. Conversely, low vigor uniformity quality signals substantial spatial or inter-vine variation requiring either differential management approaches or acceptance of quality inconsistency in the resulting fruit. The causes of vigor non-uniformity encompass soil heterogeneity in depth, texture, water holding capacity, and nutrient status topographic variation affecting drainage, cold air accumulation, and solar exposure rootstock variation in mixed plantings vine age differences from replanting disease pressure variation including virus distribution and trunk disease incidence and irrigation system non-uniformity. Measurement of vigor uniformity quality employs statistical descriptors of variation including coefficient of variation, range, and spatial autocorrelation metrics applied to vigor indicators such as pruning weights, shoot counts, trunk circumference, or remotely sensed vegetation indices. The management implications of vigor uniformity quality extend beyond harvest logistics to encompass block design for differential management, harvest segregation decisions, replanting prioritization, and root cause diagnosis of vineyard underperformance. Precision viticulture technologies have transformed vigor uniformity quality assessment through enabling spatially continuous vigor mapping via aerial and satellite imagery, supporting identification of management zones with distinct uniformity characteristics within larger vineyard blocks.
Vigor_Uniformity_Quality serves as the parent class for specific uniformity assessment scales including Within_Vine_Vigor_Uniformity_Quality, Block_Vigor_Uniformity_Quality, and NDVI_Uniformity_Quality. This class addresses variation in vigor across vines or spatial units rather than the vigor level itself, which is captured by Vigor_Quality classes. The coefficient of variation serves as the standard statistical expression of this quality. Important relationships exist with Spatial_Yield_Uniformity_Quality given the correlation between vigor and yield patterns.
Rebwuchskonsistenz
Wuchsgleichmäßigkeit
Wuchshomogenität
growth uniformity
räumliche Wuchsvariation
spatial vigor variation
vine vigor consistency
vineyard vigor homogeneity
Bramley RGV, Hamilton RP (2004) Understanding variability in winegrape production systems 1. Within vineyard variation in yield over several vintages. Australian Journal of Grape and Wine Research 10:32-45 Taylor JA et al. (2010) Considerations on spatial crop load mapping. Australian Journal of Grape and Wine Research 16:144-155 Hall A et al. (2011) Optical remote sensing applications in viticulture: a review. Australian Journal of Grape and Wine Research 17:235-248
Vigor Uniformity Quality
A vigorous dominant apical shoot on Vitis vinifera cv. Grenache grafted on 110R rootstock in Châteauneuf-du-Pape, France, emerging from node 4 on a young trunk following frost damage to the terminal bud in April, achieving 240 cm total extension growth by September with 28 nodes, internode lengths averaging 10-12 cm in the mid-shoot region, individual leaves reaching 180 cm² area with deep green coloration indicating high chlorophyll content, stem diameter of 11 mm at node 15, and complete suppression of lateral shoot development along its entire length, requiring three successive topping interventions during the season to prevent excessive vegetative growth and redirect carbohydrate allocation toward root system development.
A dominant apical shoot exhibiting exceptionally strong growth characterized by rapid internode elongation rates exceeding 4-5 cm per day during peak growth periods, large leaf area production with individual leaves reaching 150-200 cm² or greater, thick stem diameter development typically 8-12 mm or more at mid-shoot positions, and very strong apical dominance effects completely suppressing lateral bud outgrowth along the entire shoot length in Vitis vinifera. Vigorous dominant apical shoots represent maximum expression of the vine's compensatory growth capacity following main shoot loss, typically arising when release from apical dominance coincides with high soil moisture availability, abundant nitrogen nutrition, favorable temperatures, and early-season timing that allows extended growth duration. These shoots can achieve total seasonal extension growth of 2-3 meters or more, producing 20-30 or more nodes with fully expanded leaves, and may create significant canopy management challenges through excessive vegetative growth that competes with reproductive development, creates dense shading in interior canopy zones, and complicates mechanization operations. However, vigorous dominant apical shoots are valuable in young vineyard establishment where rapid framework development is prioritized, and can be managed through strategic topping, lateral removal, or resource manipulation to redirect growth energy toward desired architectural outcomes.
Vigorous dominant apical shoots are distinguished from moderate or weak dominant apical shoots by quantitative growth parameters including internode length, leaf size, stem diameter, and growth rate, with thresholds varying by cultivar, rootstock, and environmental conditions. These shoots often indicate high soil fertility, particularly nitrogen availability, or vigorous rootstock influence, and their presence may signal need for adjusted fertilization programs or water management strategies. In production vineyards, vigorous dominant apical shoots emerging from established cordons or arms typically require removal to prevent canopy disorder, while in young vineyards they may be retained and managed to accelerate framework establishment. Excessive vigor can be modulated through summer pruning, shoot positioning, or induced water stress.
excessive dominant shoot
highly vigorous substitute shoot
hochwüchsiger Ersatztrieb
sehr kräftiger dominanter Seitentrieb
stark-dominanter Lateraltrieb
strong lateral-origin apical shoot
Based on Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Poni et al. (2013) "Grapevine quality= A multiple choice issue" Scientia Horticulturae 234=445-462 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 298-324.
Vigorous Dominant Apical Shoot
A vigorous primary vegetative shoot on Vitis vinifera cv. Cabernet Sauvignon from basal node 1 in young vigorous vineyard (third leaf), reached 215cm length by mid-July despite multiple hedging operations (topped at 180cm twice, regrew rapidly), currently 11mm diameter at base, 22 leaves with average area 185 cm² totaling approximately 4070 cm² leaf area from single shoot, strong apical dominance with minimal lateral development, representing excessive vegetative vigor from combination of young vine, fertile deep soil, generous irrigation, and light crop load (1.8 kg on entire vine), requiring removal during summer shoot thinning to reduce excessive canopy density and redirect resources toward fruit development and wood maturation.
A primary vegetative shoot exhibiting strong growth characteristics, characterized by rapid elongation rate (typically 4-8cm per day during active growth phase), extended final length (often 150-250cm or more), large internodes (8-15cm spacing), thick diameter (8-12mm at base), abundant large leaves (15-25 leaves, individual leaf areas 150-250 cm²), and high total photosynthetic capacity though without reproductive contribution, typical of high-vigor sites, young vines, excessive nitrogen availability, or reduced crop load situations creating resource surplus channeled into vegetative development. Vigorous primary vegetative shoots represent extreme vegetative expression, indicating imbalanced vine conditions with excessive growth resources relative to reproductive demand, requiring management intervention including shoot removal or retention with aggressive hedging, crop load adjustment in following seasons, nitrogen reduction, possible deficit irrigation, or other practices redirecting excess vigor toward productive balance.
Vigorous primary vegetative shoots indicate vine imbalance favoring vegetative over reproductive development, often problematic rather than beneficial despite high photosynthetic capacity, as excessive vigor creates shading (reducing fruit quality and bud fruitfulness), disease pressure (dense canopy with poor air circulation), delayed maturation (continued vegetative growth consuming resources needed for fruit ripening), and poor wood maturation (late-season growth preventing adequate lignification). Management strategies include immediate intervention (shoot removal or severe hedging), medium-term adjustments (increased crop load, reduced nitrogen and water), and long-term planning (rootstock or training system modifications if chronic vigor excess). However, some vigorous vegetative shoots may be valuable as renewal wood, with strong growth providing excellent material for training new cordons, arms, or replacement structures where architectural development is desired.
high-vigor non-fruiting shoot
starkwachsender Blatttrieb
strong primary vegetative shoot
wüchsiger primärer vegetativer Trieb
Based on Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 113-145 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Howell (2001) "Sustainable grape productivity and the growth-yield relationship= A review" American Journal of Enology and Viticulture 52=165-174.
Vigorous Primary Vegetative Shoot
A vigorous water shoot on Vitis vinifera cv. Malbec grafted on 110R rootstock in Mendoza, Argentina, emerging from a lower trunk latent bud following severe hail damage in November that destroyed 95% of the canopy, achieving 380 cm total extension growth by late March with 32 nodes, individual internodes reaching 18-22 cm in mid-shoot regions, leaves in the 280-320 cm² range with pale green coloration and soft texture indicating high water content, stem diameter of 16 mm at node 15, showing continued active growth in late summer when normal shoots had ceased elongation, requiring multiple topping interventions to prevent excessive resource diversion and late-season growth that would compromise winter cold hardiness in this continental climate region.
A water shoot exhibiting growth characteristics significantly exceeding typical water shoot vigor in Vitis vinifera, characterized by exceptional daily elongation rates of 8-12 cm or more during peak growth periods, very long internodes frequently reaching 15-25 cm length, extremely large leaves often exceeding 250-300 cm² area with light green coloration indicating high nitrogen content and active growth metabolism, thick succulent stems of 12-18 mm diameter or greater despite current-season origin, and complete absence of reproductive structures reflecting fully vegetative developmental programming. Vigorous water shoots represent maximum expression of the grapevine's compensatory growth capacity, typically developing under conditions combining severe canopy loss with abundant soil moisture, high nitrogen availability, warm temperatures, and early-season emergence timing that permits extended growth duration. These shoots can achieve total seasonal extension of 3-4 meters or more, producing 25-35 nodes or more with massive total leaf area that may equal or exceed the entire original canopy photosynthetic surface, creating severe resource competition with remaining productive structures and potentially compromising fruit ripening, wood maturation, and winter hardiness through continued late-season vegetative growth that prevents proper dormancy preparation. However, vigorous water shoots also provide rapid canopy reconstruction following catastrophic damage events and can be strategically managed for framework renewal in situations requiring urgent structural replacement.
Vigorous water shoots are distinguished from moderately vigorous or typical water shoots by quantitative growth parameters including internode length, leaf size, stem diameter, and total extension growth, with thresholds varying by cultivar and environmental conditions but generally representing the upper 15-20% of water shoot vigor distribution. These shoots often indicate excessive nitrogen fertilization, over-irrigation, or extremely vigorous rootstock influences in addition to the triggering damage or pruning events. Management of vigorous water shoots requires aggressive intervention including early topping to limit total growth, removal of competing shoots to reduce resource demands, or complete removal if framework position is inappropriate. Retention of vigorous water shoots for framework renewal requires multi-season vigor moderation strategies including crop load adjustment, controlled water stress, and nitrogen management.
excessive water shoot
extremely vigorous sucker
exzessiver Stammtrieb
hyper-vigorous epicormic shoot
hypervigorer Wassertrieb
sehr kräftiger Wasserschoss
Based on Winkler et al. (1974) "General Viticulture" University of California Press pp. 246-267 Kliewer & Dokoozlian (2005) "Leaf area/crop weight ratios of grapevines= influence on fruit composition and wine quality" American Journal of Enology and Viticulture 56=170-181 Dry (2000) "Canopy management for fruitfulness" Australian Journal of Grape and Wine Research 6=109-115 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 389-415.
Vigorous Water Shoot
A Pinot Noir vine in a non-classified parcel within the commune of Volnay bearing a village wine role produces fruit under the Volnay AOC at maximum yields of 40 hl/ha with minimum natural alcohol of 10.5%, labeled simply as "Volnay" without vineyard designation, typically priced at 25-50 €/bottle representing the commune's baseline quality expression, with approximately 60% of Volnay's 213 ha of vineyards classified at village level versus 35% Premier Cru and 0% Grand Cru.
An appellation-specific role that inheres in a grapevine situated within a commune-level appellation whose fruit qualifies for village-level designation without individual vineyard or cru classification, representing the standard communal production tier positioned below Premier Cru and above regional appellations in hierarchical classification systems.
Village_Wine_Role represents the communal baseline of hierarchical appellation systems, most clearly articulated in Burgundy where 44 commune-level appellations form the backbone of the regional classification. The role is analogous to the VDP.Ortswein tier in Germany, the Classico zone designation in Italian DOC systems, and the Cru Bourgeois level in Bordeaux: each system defines village-level production as the standard quality expression of a specific geographic commune. Village wines may originate from any qualifying parcel within the commune not classified at a higher tier, and may be blended across parcels within the communal boundary.
Dorflagen-Rolle
Gemeindewein-Rolle
Ortswein-Rolle
communal appellation role
commune wine role
village-level vine role
INAO (2023) Cahiers des charges appellations communales de Bourgogne VDP (2022) VDP.Klassifikation Statuten Robinson J and Harding J (2015) The Oxford Companion to Wine 4th Ed, Oxford University Press pp: 103-105
Village Wine Role
The Vin de Pays designation Vin de Pays d'Oc (now IGP Pays d'Oc) covers the entire Languedoc-Roussillon region and has become the most commercially successful Vin de Pays category with annual production exceeding 600 million bottles — the designation permits varietal labeling (e.g., Chardonnay Pays d'Oc) which is prohibited for most AOC wines, allows 58 authorized grape varieties including international cultivars absent from regional AOC specifications, and accepts maximum yields of 90 hl/ha, providing a regulatory framework that enabled the transformation of the Languedoc from bulk wine production to competitive varietal wine marketing in international export markets from the 1990s onward.
A regional wine designation within the French wine classification system that identifies a wine as originating from a defined Vin de Pays zone, now formally classified as a Protected Geographic Indication (IGP) under EU harmonization but retaining the traditional Vin de Pays terminology on labels as permitted by EU Regulation, representing the intermediate quality tier between Vin de France (without geographic indication) and AOC/AOP (Protected Designation of Origin).
Vin_De_Pays_Designation documents the French national designation system that pioneered the concept of intermediate geographic indication between table wine and full appellation status. Created in 1968 and formally regulated from 1973, the Vin de Pays system predates the EU PGI framework by two decades and provided the model for the European harmonization of geographic indication categories: the 2009 EU wine reform formally reclassified all Vin de Pays as IGP (Indication Géographique Protégée), but EU Regulation 1308/2013 Art. 112 permits continued use of the traditional Vin de Pays terminology on labels as an alternative to the harmonized IGP term. The Vin de Pays system operates at three geographic levels — departmental (e.g., Vin de Pays de l'Hérault), zonal (e.g., Vin de Pays des Coteaux de l'Ardèche), and regional (e.g., Vin de Pays d'Oc) — with the regional level dominating commercial production.
French IGP designation
VdP classification
VdP-Klassifikation
Vin de Pays
Vin-de-Pays-Bezeichnung
französische Landweinbezeichnung
EU Regulation 1308/2013 Art. 112 on traditional terms INAO (2023) Les indications géographiques protégées Decree of September 13, 1968 establishing the Vin de Pays category
Vin De Pays Designation
A 5-year-old Chardonnay vine bears a Vine_Age_Role as a young vine transitioning from establishment to production, with crop load still restricted to build vine structure while root system development continues. A 25-year-old Cabernet Sauvignon vine serves in a Vine_Age_Role as a mature, fully productive plant with optimized root-shoot balance producing consistent yields of site-characteristic fruit. A 100-year-old Zinfandel vine bears a Vine_Age_Role as a centenarian commanding premium pricing, its low yields and deep-rooted resilience producing intensely concentrated fruit despite—or because of—its advanced age.
A vineyard role that inheres in a grapevine by virtue of its chronological age since planting, with age determining physiological status, productive capacity, management requirements, and market positioning. Vine age roles recognize that grapevines are perennial crops with productive lifespans of 25-100+ years, passing through distinct developmental phases—establishment (years 1-3), young production (years 4-10), mature production (years 11-30), and old vine status (30+ years)—each with characteristic physiology and fruit quality implications. Age effects on vine performance are complex: young vines show vigorous but unbalanced growth mature vines achieve optimal balance between vegetative and reproductive development old vines often produce reduced yields but concentrated, terroir-expressive fruit valued for premium production. Age-related market positioning is significant, with "old vine" and similar designations commanding premium pricing in many markets based on perceived quality associations with vine age. The relationship between chronological age and physiological status varies with site conditions, variety, rootstock, and management history a 50-year-old vine on favorable terroir may be productively vigorous while a 20-year-old vine on marginal site may show senescence.
Vine_Age_Role is subclassed by age category—Young, Established, Mature, and Old—recognizing the distinct phases of grapevine productive life. Age categories are not rigidly defined boundaries vary with regional tradition, variety characteristics, and production context. The age-quality relationship is contested while old vine fruit often commands premiums, the mechanism (reduced yields, deep roots, physiological balance) rather than age per se likely drives quality differences. Vine age interacts with site quality—exceptional sites may support productive old vines while marginal sites force replanting at younger ages—making age a proxy for site suitability as much as an independent quality factor. The economics of vine age involve tradeoff between the investment in old vines (accumulated development, established root systems, potential premium pricing) and the opportunity cost of not replanting with improved material (better clones, more appropriate rootstocks, optimized training systems).
Pflanzalter
Rebenalter
Rebenreife
Weinbergsalter
planting age
vine age
vine maturity
vineyard age
Vaudour E et al. (2015) Vine age effect on terroir expression, in Piéri P (ed) Proceedings of International Terroir Congress, OENO One Bonada M et al. (2020) Vine age and grape composition, Australian Journal of Grape and Wine Research 26: 139-152 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, Academic Press
Vine Age Role
A precision viticulture study in Barossa Valley Shiraz employed vine balance index quality assessment combining Ravaz Index, leaf area index measured via LAI-2200, point quadrat canopy gaps percentage, and fruit zone photosynthetically active radiation to identify management zones within a 15-hectare block. Zones with optimal vine balance index quality (Ravaz 6-8, LAI 1.5-2.0, gaps >30%, fruit PAR >15% ambient) produced fruit with 15% higher anthocyanin concentration and more favorable sensory profiles compared to imbalanced zones requiring targeted intervention through differential irrigation, selective shoot thinning, and zone-specific harvest timing.
Vine balance index quality is a vigor balance quality that inheres in a bearing grapevine organism and represents an integrative assessment of vine equilibrium incorporating multiple vegetative and reproductive parameters beyond simple biomass ratios to provide comprehensive evaluation of growth-yield relationships and their implications for fruit quality and vineyard sustainability. This quality extends traditional balance concepts by considering the temporal dynamics of growth, the spatial distribution of vegetative and reproductive organs, and the functional relationships between canopy architecture and fruit development rather than solely endpoint biomass measurements. The ontological significance of vine balance index quality lies in its capacity to capture nuanced aspects of vine performance that single-parameter indices may miss, including the timing of growth cessation relative to fruit development, the adequacy of leaf area exposure for fruit ripening, and the distribution of crop load relative to canopy capacity across different vine positions. Various formulations of vine balance indices have been proposed, ranging from modifications of the Ravaz Index incorporating leaf area measurements to complex algorithms integrating yield components, pruning weights, canopy dimensions, and fruit composition parameters. The Smart-Robinson approach to balance assessment considers not only yield-to-pruning-weight ratios but also canopy density indicators, fruit exposure metrics, and growth cessation timing to provide multidimensional characterization of vine equilibrium status. Functional vine balance index quality recognizes that optimal balance varies systematically with production objectives, wherein high-yield table grape production may tolerate or require different balance parameters than premium wine grape production targeting fruit concentration and phenolic development. The measurement of vine balance index quality typically requires comprehensive data collection spanning the growing season, including dormant pruning weights, shoot counts and lengths at multiple developmental stages, leaf area estimates through direct measurement or modeling, yield component analysis, and potentially fruit composition assessment at harvest. Advanced approaches to vine balance index quality incorporate precision viticulture technologies including normalized difference vegetation index mapping, proximal canopy sensors, and yield monitors to enable spatially explicit assessment of balance variation within vineyard blocks, supporting site-specific management interventions to optimize equilibrium across heterogeneous growing conditions.
Vine_Balance_Index_Quality is_a Vigor_Balance_Quality, representing comprehensive multi-parameter approaches to balance assessment beyond single ratios. This class encompasses various integrative methodologies including the Smart-Robinson system, functional balance concepts, and precision viticulture approaches. Distinguished from Vegetative_Reproductive_Balance_Quality by incorporation of canopy architecture, light environment, and temporal dynamics rather than solely biomass partitioning.
Mehrfaktor-Balanceindex
Rebbalanceindex
comprehensive vine balance
integrated balance assessment
integrierter Gleichgewichtsindex
multifactor balance index
umfassende Rebbalance
vine equilibrium index
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 88-107 Bramley RGV (2010) Precision Viticulture: managing vineyard variability for improved quality outcomes. In: Reynolds AG (ed) Managing Wine Quality, Volume 1, Woodhead Publishing, pp. 445-480 Tardaguila J et al. (2011) Smart-Robinson comprehensive system for canopy characterization. American Journal of Enology and Viticulture 62:76-84
Vine Balance Index Quality
Vine longevity ranges from 25-40 years for short-lived cultivar-site combinations to over 150 years for exceptional old vine vineyards, with documented examples of productive vines exceeding 200 years in Australia (Barossa Shiraz), California (Zinfandel), and Europe (various cultivars). Longevity-promoting factors include inherent trunk disease resistance, appropriate rootstock selection (matching vigor to site), balanced crop load management, and absence of catastrophic stress events. Cultivars reputed for longevity include Zinfandel, Mourvèdre (Monastrell), Carignan, and Grenache, while Merlot and Sauvignon Blanc are often considered shorter-lived. Annual trunk diameter increment of 0.5-2.0 mm in mature vines gradually increases conductive capacity but also accumulates tyloses, wound tissue, and pathogen-damaged zones that eventually compromise function.
A longevity and resilience disposition that is the inherited capacity of a Vitis vinifera plant to maintain structural integrity, vascular function, and reproductive capacity over extended time periods, with genotype-specific maximum productive lifespan determined by trunk disease resistance, wood durability, and the balance between accumulating damage and regenerative capacity.
Vine_Longevity_Disposition serves as the parent class for structural integrity, decline resistance, and growth resumption traits. Longevity is determined by the race between damage accumulation (trunk diseases, mechanical injury, environmental stress) and the vine's regenerative capacity. The concept of "vine age" as a quality factor is embedded in wine marketing (vieilles vignes), though the mechanisms linking age to quality likely involve root system development, yield reduction, and vine balance rather than age per se.
Lebensdauer
Produktionsdauer
Rebstock-Langlebigkeit
long-term viability disposition
productive lifespan
vine persistence trait
Fontaine F et al. (2016) Grapevine trunk diseases: etiology and management. In: Maixner M, Eder J (eds) Phytoplasma, Methods in Molecular Biology, Springer pp.: 427-443 Smart RE (2014) Old vines - can we define the term for Barossa wines? Wine and Viticulture Journal 29:38-42
Vine Longevity Disposition
A Pinot Noir vine planted at 1.0 m × 1.0 m (10,000 vines/hectare) in Burgundy bears a Vine_Spacing_Role reflecting traditional high-density planting where close spacing limits individual vine vigor on potentially rich soils while maximizing per-hectare site utilization. A Cabernet Sauvignon vine at 2.4 m × 1.8 m (2,315 vines/hectare) in Napa Valley serves in a Vine_Spacing_Role optimized for mechanical operations and the vigorous growth enabled by fertile valley floor soils and irrigation. A Shiraz vine at 3.5 m × 2.0 m (1,430 vines/hectare) in Australian irrigated production bears a Vine_Spacing_Role suited to high-vigor conditions and divided canopy systems requiring wide inter-row spacing.
A vineyard role that inheres in a grapevine by virtue of its spatial positioning within the vineyard planting pattern, determined by the distances between vines within rows (vine spacing) and between rows (row spacing), which together define vine density and the area available to each vine. Vine spacing roles reflect fundamental vineyard design decisions that influence vine competition, root zone volume, canopy development potential, mechanization compatibility, and economic productivity throughout the vineyard's productive life. Spacing affects vine physiology through resource partitioning—closer spacing increases root zone competition for water and nutrients while distributing canopy over more vines, potentially reducing per-vine vigor but increasing per-hectare productivity. Traditional European vineyards often feature high-density planting (8,000-12,000 vines/hectare) developed before mechanization, while New World vineyards typically use wider spacing (1,500-4,000 vines/hectare) compatible with tractor operations and adapted to more vigorous growing conditions. The optimal spacing reflects variety vigor, site fertility, water availability, training system, mechanization requirements, and production philosophy, with quality-focused producers often favoring higher density for its perceived contribution to fruit concentration and terroir expression.
Vine_Spacing_Role is subclassed by density category—High, Medium, Low, and Wide—recognizing the distinct production contexts and management implications of different planting intensities. Spacing decisions are among the most consequential in vineyard establishment, as the permanent planting pattern determines options for the vineyard's 25-50+ year lifespan. Historical European high-density planting reflected pre-mechanization labor availability and the quality philosophy that individual vine stress from competition produces more concentrated fruit modern interpretation continues this tradition while acknowledging that optimal density depends on site-specific vigor potential. The trend toward higher density in quality-focused New World viticulture represents philosophical convergence with European models, though practical constraints (mechanization, vigor management) often limit achievable density. Replanting to different spacing requires complete vineyard redevelopment, making initial density decisions particularly significant.
Pflanzabstand
Rebabstand
Rebendichte
Stockdichte
plant spacing
planting density
vine density
vine spacing
Archer E, Strauss HC (1985) Effect of plant density on root distribution of three-year-old grafted 99 Richter grapevines, South African Journal of Enology and Viticulture 6: 25-30 Reynolds AG et al. (1994) Vine performance and fruit composition affected by vine spacing, American Journal of Enology and Viticulture 45: 444-459 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, Academic Press
Vine Spacing Role
Vine-to-vine yield variance assessment in aging Syrah vineyard reveals the trunk disease impact on population uniformity, with healthy vine population (n=50, no visible disease) showing low variance (mean 5.2 kg/vine, CV 18%, range 3.8-6.8 kg), while disease-affected population (n=50, variable Esca symptoms) shows high variance (mean 3.8 kg/vine, CV 52%, range 0-7.2 kg including dead and dying vines), with the variance difference indicating that average yield statistics poorly represent the diseased block where many vines produce normally while others fail entirely, requiring individual vine assessment rather than block-level management approaches.
Vine-to-vine yield variance quality is a yield consistency quality that represents the variation in production among individual vines within a vineyard or block, characterizing the plant-level heterogeneity that underlies spatial patterns and determines the uniformity of the vine population as a production unit. Within the BFO framework, this quality specifically addresses the individual plant dimension of yield variation, distinguishing vine-level differences from zone-level or whole-vineyard patterns while recognizing that vine-to-vine variance represents the fundamental unit of production heterogeneity from which larger-scale patterns emerge, with high variance indicating a diverse vine population producing widely different amounts while low variance indicates a uniform population where most vines produce similarly. The ontological significance lies in vine-to-vine variance affecting both practical management and production quality, where high variance creates harvest logistics challenges (difficulty estimating total production from sample counts), complicates vine balance assessment (average metrics may not represent actual vine condition), and may indicate underlying problems (disease, missing vines, rootstock failures) requiring individual vine attention, while low variance enables efficient sample-based estimation and uniform management application. The factors contributing to vine-to-vine variance include planting material uniformity (clonal versus seedling origin, rootstock consistency), establishment success (missing or weak vines from poor establishment), disease effects (individual vines affected by trunk diseases, virus infection), soil microvariation (localized differences affecting individual vine positions), and management history (individual vine responses to past damage, pruning variation). The measurement of vine-to-vine variance involves weighing fruit from individual vines (labor-intensive but direct) or estimating from cluster counts and weights on sample vines, with variance expressed as coefficient of variation (CV) enabling comparison across vineyards with different mean yields, with typical CVs ranging from 15-25% in uniform vineyards to 40-60% in heterogeneous populations. Understanding vine-to-vine yield variance quality enables assessment of population uniformity, identification of individual problem vines, and yield estimation precision based on the sampling intensity required for given variance levels.
Vine_To_Vine_Yield_Variance_Quality addresses the production variation among individual plants within vineyard populations. This quality reflects planting uniformity, vine health, and microsite variation. The class supports population assessment and sampling design through vine-to-vine variance characterization.
Einzelrebenvarianz
Einzelstockertragsunterschiede
Rebe-zu-Rebe-Ertragsvarianz
Rebpopulationsheterogenität
individual vine variation
plant-level yield variance
single-vine yield differences
vine population heterogeneity
Bramley RGV et al. (2011) Vineyard variability in Marlborough, New Zealand: characterising variation in vineyard performance and options for the implementation of Precision Viticulture. Australian Journal of Grape and Wine Research 17(1): 72-78 Hall A et al. (2011) Within-season temporal variation in correlations between vineyard canopy and winegrape composition and yield. Precision Agriculture 12(1): 103-117
Vine To Vine Yield Variance Quality
Vineyard block synchrony assessment using spatially distributed monitoring in a 2-hectare Chardonnay block reveals the uniformity level affecting management strategy, with block-wide véraison date CV of 5.2% (moderate synchrony, most vines within 4-day window but tails extending 10 days), ripening trajectory analysis showing that the 5.2% véraison CV amplifies to 8.8% Brix CV at target harvest date (differential ripening rates exacerbating initial spread), and spatial mapping revealing systematic west-to-east gradient (soil drainage variation), suggesting that either the block be managed as two zones (east and west halves) or harvested in two passes to capture optimal maturity in both subpopulations.
Vineyard block synchrony quality is an inter-plant synchrony quality that represents the developmental uniformity among vines within a defined vineyard management unit, characterizing the phenological coherence that determines whether the block functions as a homogeneous production unit suitable for uniform management and single-harvest treatment or requires subdivision into more uniform zones for optimized management. Within the BFO framework, this quality specifically addresses the block-level dimension of inter-plant synchrony, distinguishing uniformity within defined management units from broader cultivar or vineyard-wide synchrony while recognizing that block synchrony determines the practical efficiency of vineyard operations that are typically organized at the block level. The ontological significance lies in vineyard block synchrony determining the validity of block-level management decisions, where high block synchrony supports the assumption that treatments applied uniformly across the block will be appropriately timed for all vines, while low block synchrony indicates that uniform treatment represents a compromise among optimal timings for different vine subpopulations, potentially reducing treatment efficacy or increasing quality variation. The spatial patterns of block synchrony typically reflect underlying environmental gradients, with soil variation (texture, depth, water holding), topographic variation (elevation, slope, aspect), and microclimate variation (frost pockets, wind exposure, heat accumulation) creating systematic phenological patterns across blocks that may justify subdivision into more uniform management zones. The quantification of block synchrony involves measures of developmental spread such as the coefficient of variation in phenological dates among vines, the range from earliest to latest development, or the proportion of vines falling within a defined developmental window, with these metrics enabling comparison among blocks and tracking of synchrony changes across seasons. The management implications of block synchrony include the appropriateness of block-level versus zone-level decision-making, the efficiency gains achievable from high-synchrony blocks, and the potential quality improvements from subdividing low-synchrony blocks into more uniform management zones that enable optimized treatment timing. Understanding vineyard block synchrony quality enables evidence-based decisions about management unit definition, appropriate expectations for block-level quality uniformity, and identification of within-block variation patterns that might benefit from precision viticulture approaches.
Vineyard_Block_Synchrony_Quality addresses the developmental uniformity within defined management units determining treatment efficiency. This quality reflects spatial patterns from environmental variation within blocks. The class supports management zoning decisions through block synchrony characterization.
Blocksynchronie
Management-Einheit-Synchronie
management unit synchrony
räumliche Entwicklungsuniformität
spatial developmental uniformity
within-block coordination
Bramley RGV (2005) Understanding variability in winegrape production systems 2. Within vineyard variation in quality over several vintages. Australian Journal of Grape and Wine Research 11(1): 33-42 Hall A et al. (2011) Within-season temporal variation in correlations between vineyard canopy and winegrape composition and yield. Precision Agriculture 12(1): 103-117
Vineyard Block Synchrony Quality
The vineyard designation system reaches its most elaborate expression in Burgundy, where approximately 1,250 individually named climats (vineyard parcels) are officially delimited and classified — from village-level vineyards (e.g., Les Charmes in Meursault) through Premier Cru sites (e.g., Clos Saint-Jacques in Gevrey-Chambertin, 6.7 hectares) to Grand Cru monopoles (e.g., Romanée-Conti, 1.81 hectares owned solely by Domaine de la Romanée-Conti), each climat name appearing on the wine label as a precise geographic identifier enabling consumers and collectors to track terroir expression at the individual site level across vintages and producers.
A geographic indication that identifies a specific vineyard site, parcel, or named viticultural lieu-dit within a broader appellation or wine region, providing the finest level of geographic resolution in wine origin communication and associating the wine with the distinctive terroir characteristics of an individual site recognized by tradition, regulatory delimitation, or proprietor designation.
Vineyard_Designation captures the site-level naming systems that represent the finest geographic resolution in viticultural classification. The concept of named vineyard sites has deep historical roots — Burgundy's climats have been individually identified since the Cistercian monastic period (12th-13th centuries), Germany's Einzellagen were catalogued in the 1971 Wine Law, and Italy's Menzioni Geografiche Aggiuntive have been progressively delimited since 2010: the common principle is that individual vineyard sites possess distinctive and consistent terroir characteristics (soil, microclimate, aspect, elevation) that produce wines with recognizable site-specific qualities discernible to experienced tasters. Subclasses represent the principal national traditions of vineyard-level naming — French (lieu-dit, climat, cru), German (Lage), Austrian (Ried), Italian (contrada), and Spanish (pago) — each carrying distinct regulatory and cultural connotations.
Einzellagen-Kennung
Weinbergsbezeichnung
single-site indication
site designation
vineyard name
INAO (2023) Délimitation parcellaire des appellations d'origine Robinson J and Harding J (2015) The Oxford Companion to Wine 4th Ed, Oxford University Press EU Regulation 1308/2013 Art. 93 and Delegated Regulation 2019/33 Art. 55
Vineyard Designation
The vineyard location description for a parcel in the Wachau (Lower Austria) documents the hierarchical geographic identity — Country: Austria Federal State: Niederösterreich (Lower Austria) Wine Region: Wachau DAC Commune: Dürnstein Ried (named vineyard): Kellerberg Parcel: Grundstücksnummer 789/2 — this administrative location places the vineyard within the Austrian Districtus Austriae Controllatus (DAC) quality wine system, identifies it as within the Wachau protected designation of origin, and specifies the Ried Kellerberg as a classified vineyard site (Erste Lage in the Österreichische Traditionsweingüter classification, analogous to the Burgundian Premier Cru). The location description enables integration with the Austrian vineyard cadastral database (Weinbaukataster), which records the permitted cultivars, maximum yields, and quality classification for each registered parcel.
A site description component that records the administrative and geographic location of the vineyard — country, region, sub-region, commune or township, and named vineyard or parcel — providing the hierarchical geographic reference that links the vineyard to its applicable regulatory framework (appellation, designation of origin, geographic indication), positions it within the recognized viticultural regions of the world, and enables its identification within the landscape of local and regional terroir classifications.
Vineyard_Location_Description provides the administrative identity that connects the physical vineyard to the regulatory, economic, and cultural frameworks governing its production. The hierarchical structure of vineyard location — from country through region to individual parcel — mirrors the nested hierarchy of appellation and designation of origin systems that define the legal framework for wine labeling in every major wine-producing country: the European Union's PDO/PGI system, the French AOC/AOP hierarchy (from regional appellations like Bourgogne through communal appellations like Vosne-Romanée to individual vineyard designations like Grand Cru Romanée-Conti), the Italian DOC/DOCG system, the German Anbaugebiet/Bereich/Einzellage hierarchy, and the New World geographic indication systems (Australian GIs, American AVAs, South African WOs) all define wine identity and permitted production practices at progressively finer geographic scales. The vineyard location also determines the applicable viticultural regulations — permitted cultivars, planting density requirements, maximum yields, minimum ripeness levels, and authorized viticultural practices vary between appellations — making accurate location documentation essential for regulatory compliance and for the labeling claims that communicate geographic origin to consumers.
Reb-Lagebeschreibung
Weinberg-Lage-Beschreibung
administrative site identification
geographic location documentation
geographische Standortbeschreibung
vineyard address record
van Leeuwen C and Seguin G (2006) The concept of terroir in viticulture, Journal of Wine Research 17: 1-10 OIV (2010) Resolution OIV/VITI 333/2010 — Definition of vitivinicultural terroir European Union Regulation (EU) No 1308/2013 — Common organisation of the markets in agricultural products
Vineyard Location Description
A Syrah vine trained to bilateral cordon in a Barossa Valley vineyard bears a Vineyard_Role defined by its cordon training, spur pruning regime, and premium red wine production objective—its role determining canopy architecture, yield expectations, and harvest timing decisions. A Riesling vine in steep Mosel terracing serves in a Vineyard_Role shaped by single-stake training adapted to slope cultivation, manual labor requirements, and Prädikatswein quality targeting. A machine-harvested Chardonnay vine in an irrigated Central Valley vineyard bears a Vineyard_Role optimized for mechanical operations, high-volume production, and bulk wine or juice processing.
A grapevine role that inheres in a plant by virtue of its position, function, or management status within a vineyard production system, encompassing the diverse roles vines occupy as they are cultivated for fruit production under specific training systems, management regimes, and production objectives. Vineyard roles reflect the transformation of grapevines from biological organisms to managed agricultural units, with role assignment determining how vines are trained, pruned, irrigated, protected, and harvested within the vineyard context. Role categories span training system implementation (how vine architecture is shaped), production intensity (yield targets and quality positioning), spatial organization (row spacing, vine density, trellising), and temporal management (establishment, mature production, decline). Vineyard role assignment interacts with terroir expression—the same variety and clone may express differently under different training systems and management regimes, with vineyard role choices influencing fruit composition, wine character, and economic outcomes. Contemporary vineyard management increasingly optimizes role assignment through precision viticulture, matching training systems and management intensity to site-specific conditions for improved efficiency and quality.
Vineyard_Role is the parent class for the hierarchy of vine functions within production systems, subclassed by training system (Training_System_Role), production intensity, spatial organization, and other management dimensions. The vineyard role concept recognizes that viticulture imposes functional categories on biological variation—vines are assigned roles that determine their treatment, expectations, and evaluation criteria. Role assignment reflects accumulated knowledge about variety-site-management interactions, with traditional regional systems (Burgundy's guyot, Champagne's taille Chablis, Mosel's single-stake) representing centuries of optimization for local conditions. Contemporary viticulture increasingly recognizes that optimal role assignment varies within vineyards, with precision approaches matching management to spatial variation in soil, microclimate, and vine performance.
Bewirtschaftungsrolle
Produktionsrolle
Rebanlagenrolle
Weinbergsrolle
production role
vine management role
vineyard role
Jackson DI, Lombard PB (1993) Environmental and management practices affecting grape composition and wine quality, American Journal of Enology and Viticulture 44: 409-430 Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles Gladstones J (1992) Viticulture and Environment, Winetitles
Vineyard Role
Detailed profiling of Viniferin_Production_Disposition at the Julius Kühn-Institut characterized stilbene oligomerization dynamics across 18 cultivars following Plasmopara viticola inoculation. Leaf samples were analyzed by LC-MS at 24, 48, 72, and 96 hpi for resveratrol, piceid, ε-viniferin, δ-viniferin, and pterostilbene to track both phytoalexin production and metabolic conversion. Cultivars showed distinct profiles reflecting differential Viniferin_Production_Disposition. Resistant 'Regent' exhibited rapid conversion with ε-viniferin exceeding resveratrol by 48 hpi (32 μg/g versus 18 μg/g), reaching final ε-viniferin concentration of 58 μg/g by 72 hpi representing 65% of total stilbenes. Susceptible 'Müller-Thurgau' showed delayed, limited oligomerization with resveratrol remaining dominant through 96 hpi and ε-viniferin never exceeding 4 μg/g (22% of total stilbenes). The PIWI cultivar 'Solaris' displayed the strongest Viniferin_Production_Disposition with early resveratrol peak at 24 hpi followed by rapid conversion, achieving ε-viniferin-to-resveratrol ratio of 4.5 by 72 hpi. Correlation analysis revealed that ε-viniferin concentration at 48 hpi was a better predictor of resistance (r = -0.85 with sporulation) than total stilbenes (r = -0.68) or resveratrol alone (r = -0.55), confirming the superior defensive value of oligomeric stilbenes. Peroxidase activity assays showed that 'Solaris' leaf extracts converted exogenous resveratrol to ε-viniferin 3-fold faster than 'Müller-Thurgau' extracts, indicating that enhanced peroxidase activity contributes to strong Viniferin_Production_Disposition.
A Viniferin_Production_Disposition is a Phytoalexin_Production_Disposition that inheres in a Vitis vinifera plant and is realized through the capacity to synthesize and accumulate viniferins—oligomeric stilbenes formed by oxidative coupling of resveratrol units—which exhibit substantially higher antifungal activity than their monomeric precursor resveratrol. This disposition represents the production of the most potent antimicrobial stilbenes in grapevine defense, including ε-viniferin (resveratrol dimer), δ-viniferin (another dimeric form), α-viniferin (cyclic trimer), and higher oligomers, which together constitute the active phytoalexin arsenal in resistant responses to fungal pathogens. Viniferin formation occurs through the action of peroxidases that catalyze oxidative coupling of resveratrol, a process that can occur both enzymatically within plant tissues and chemically through pathogen-derived oxidases, creating a dynamic interplay between host and pathogen during infection. The antifungal activity of ε-viniferin exceeds that of resveratrol by 5-10 fold against major grapevine pathogens, with minimum inhibitory concentrations of 10-50 μg/mL, making viniferin accumulation particularly important for effective stilbene-based defense. Genotypes with strong Viniferin_Production_Disposition not only produce resveratrol rapidly but also efficiently convert it to viniferins, resulting in phytoalexin profiles dominated by the more potent oligomeric forms. The kinetics of viniferin production typically lag behind resveratrol accumulation by 12-24 hours, as viniferin formation depends on prior resveratrol synthesis.
Viniferin_Production_Disposition is_a Phytoalexin_Production_Disposition. Viniferins are formed by oxidative coupling of resveratrol, making their accumulation dependent on prior resveratrol synthesis but also requiring peroxidase activity for oligomerization. ε-Viniferin is typically the dominant and most studied viniferin form, with 5-10 fold higher antifungal activity than resveratrol. The ratio of viniferins to resveratrol may be as important as total stilbene concentration for effective defense. Some fungal pathogens produce laccases that can detoxify stilbenes, creating potential for pathogen counter-adaptation to phytoalexin-based resistance.
Oligomere-Stilben-Produktion
Resveratrol-Oligomerisierung
Viniferin-Produktions-Disposition
oligomeric stilbene production
resveratrol oligomerization
viniferin synthesis capacity
Pezet R et al. (2004) δ-Viniferin, a resveratrol dehydrodimer: one of the major stilbenes synthesized by grapevine leaves, Journal of Agricultural and Food Chemistry 51:5488-5492 Langcake P, Pryce RJ (1977) A new class of phytoalexins from grapevines, Experientia 33:151-152 Schnee S et al. (2013) Vitis vinifera canes, a source of stilbenoids against downy mildew, OENO One 47:259-268
Viniferin Production Disposition
Vintage declaration rule for EU wines requires a minimum of 85% of the wine to derive from grapes harvested in the declared vintage year (Regulation 2019/33, Article 49), permitting up to 15% from adjacent vintages to accommodate practical winemaking needs (topping up barrels, adjusting blend) — thus a wine labeled "2024" must contain at least 85% wine from the 2024 harvest. Vintage declaration rule for Champagne follows the general EU 85% rule for vintage-dated Champagne (millésimé), but the majority of Champagne production is non-vintage (NV), where the base wine from the current harvest is blended with reserve wines from multiple prior vintages to achieve house-style consistency — a practice explicitly permitted by the appellation specification and representing the most commercially significant application of multi-vintage blending in the wine world. Vintage declaration rule for Port follows specific Portuguese regulations — Vintage Port must be 100% from the declared year, Late Bottled Vintage (LBV) similarly 100%, while Tawny Port with an age indication (10, 20, 30, 40 years) uses a blend of multiple vintages with the age statement reflecting the average age of the blend components.
Vintage declaration rule is a label regulation entity that documents the legal conditions under which the harvest year may appear on a wine label, specifying the minimum percentage of grapes from the declared vintage year required in the wine's composition, the regulatory framework establishing this threshold, the treatment of multi-vintage blends (non-vintage wines, solera-aged wines, reserve blends), and any appellation-specific vintage requirements that exceed the baseline regulatory minimum, providing the temporal-origin framework that enables consumers to identify the harvest year and, by extension, to assess the wine's age, vintage quality reputation, and developmental stage.
Vintage declaration rules address the temporal dimension of wine identity — the harvest year being one of the most commercially significant label elements because it enables consumers to access vintage quality information (published vintage reports, critic scores by vintage), to assess wine age and developmental stage, and to make cellaring decisions informed by the vintage's reputation for aging potential. The 85% threshold adopted by most major wine jurisdictions represents a pragmatic compromise between vintage purity and winemaking necessity — the practice of topping up barrels during aging (to compensate for evaporation through the ullage) and of minor blending adjustments means that absolute single-vintage purity would exclude many traditional winemaking practices, while a threshold substantially below 85% would undermine the meaningfulness of the vintage declaration. The non-vintage tradition — most prominently embodied in Champagne NV cuvées but also practiced in Sherry solera blending, Tawny Port age-dated blends, and some sparkling wine and fortified wine traditions — represents a fundamentally different quality philosophy where consistency across vintages, achieved through skillful multi-vintage blending, is valued over single-vintage expression. The interaction between vintage declaration and appellation requirements creates additional regulatory layers — some appellations require vintage dating (most European PDO still wines carry mandatory vintage declaration), some explicitly permit non-vintage production (Champagne, Sherry), and some impose vintage-related aging minima (Barolo DOCG requiring wine from a single vintage with minimum aging periods calculated from the vintage date). The consumer information value of vintage declarations extends beyond the wine itself — vintage dates on library releases and auction lots provide provenance documentation that supports authenticity verification and enables price differentiation based on vintage reputation, making the vintage declaration one of the most economically consequential label elements for collectible wines.
Erntejahrbestimmung
Erntejahrdeklaration
Jahrgangsangabenregel
Jahrgangsbezeichnungsvorschrift
Vintagedeklarationsregel
harvest year declaration
millésime declaration rule
vintage labeling rule
vintage percentage rule
year of harvest requirement
European Parliament (2019) Commission Delegated Regulation (EU) 2019/33, Article 49 TTB (2023) 27 CFR §4.27 (vintage labeling) CIVC (2023) Cahier des charges de l'appellation d'origine contrôlée Champagne IVDP (2023) Regulamento das denominações de origem Porto e Douro
Vintage Declaration Rule
Vintage predictability quality comparison across major Pinot Noir regions evaluated the confidence available for advance planning based on historical patterns and early-season indicators. Burgundy demonstrated good vintage predictability quality with April budbreak date explaining 72% of variance in harvest date (r²=0.72), enabling 6-week advance scheduling with ±5 day accuracy once budbreak timing was established historical quality tier prediction from July weather patterns achieved 78% accuracy across 30 vintages. Central Otago showed moderate vintage predictability quality with greater inter-annual climate variation reducing budbreak-harvest correlation to r²=0.58 and expanding harvest date uncertainty to ±9 days quality tier prediction achieved only 62% accuracy from mid-season indicators. Oregon Willamette Valley exhibited lower vintage predictability quality attributed to the variable influence of autumn precipitation on harvest timing, with October rainfall introducing substantial unpredictability despite reasonable early-season forecasts, requiring flexible harvest capacity to accommodate vintage variation ranging from early-September to mid-October across the historical record.
Vintage predictability quality is a harvest timing indicators quality that inheres in a grapevine cultivar or clone within a specific growing region and represents the consistency and reliability with which seasonal ripening patterns, harvest timing, and resulting wine characteristics can be forecast from available information including historical data, early-season phenological observations, and weather pattern analysis, enabling advance planning of vintage operations and quality outcome expectations with confidence appropriate to the prediction accuracy achievable for the specific cultivar-region combination. This quality determines the strategic planning horizon for vintage operations and the residual uncertainty that must be accommodated through flexible response capacity, with highly predictable combinations enabling confident advance scheduling while unpredictable situations require substantial contingency planning and adaptive management approaches. The ontological significance of vintage predictability quality lies in its value for production planning across the entire vintage cycle, from pruning decisions influenced by expected crop load through harvest scheduling and winery preparation to marketing planning based on anticipated quality outcomes, with reliable prediction enabling efficient resource allocation while poor predictability creates planning uncertainty and potential misallocation. The biological basis of vintage predictability quality encompasses the consistency of climate patterns in the growing region determining whether seasons follow predictable trajectories or exhibit high inter-annual variation, the responsiveness of cultivar phenology to environmental signals enabling reliable prediction from early-season observations, the stability of ripening rates once established enabling forward projection from current status, and the consistency of cultivar response to environmental variation permitting inference from similar historical vintages. Regions with continental climates exhibiting high inter-annual variation typically show lower vintage predictability quality than maritime or Mediterranean regions with more consistent seasonal patterns, while cultivars with strong phenological responses to thermal accumulation exhibit higher predictability than those with complex multi-factor ripening control. Assessment of vintage predictability quality involves compilation of multi-decade records enabling characterization of vintage variation patterns, development and validation of prediction models using early-season indicators, and quantification of forecast accuracy under normal and extreme conditions.
Vintage_Predictability_Quality is_a Harvest_Timing_Indicators_Quality, representing the reliability of forecasting vintage characteristics from available information. This class addresses prediction confidence across the planning horizon rather than measurement precision at a point in time. Distinguished from Optimal_Harvest_Date_Quality by addressing forecast reliability rather than date consistency per se. Climate stability, cultivar phenological responsiveness, and historical data availability all influence predictability. Critical for strategic vintage planning and resource allocation across seasons.
Ernteergebnis-Prognostizierbarkeit
Jahrgangsprognose-Zuverlässigkeit
Jahrgangsvorhersagbarkeit
harvest outcome forecasting
saisonale Vorhersehbarkeit
seasonal predictability
vintage forecasting reliability
vintage planning confidence
Jones GV, Davis RE (2000) Climate influences on grapevine phenology, grape composition, and wine production and quality for Bordeaux, France. American Journal of Enology and Viticulture 51:249-261 Webb LB et al. (2012) Modelled impact of future climate change on the phenology of winegrapes in Australia. Australian Journal of Grape and Wine Research 18:20-31 van Leeuwen C et al. (2019) An update on the impact of climate change in viticulture and potential adaptations. Agronomy 9:514
Vintage Predictability Quality
Violet aroma profiling in high-altitude Mendoza Malbec reveals the varietal signature with violet rating of 8.0 on 10-point scale, correlating with β-ionone concentration of 1.8 μg/L (significantly above the 0.8 μg/L detection threshold), complemented by iris (4.5) and subtle lavender (3.2), with this violet dominance distinguishing Malbec's floral profile from Syrah where violet appears alongside pepper and savory notes, demonstrating how violet aroma intensity and context enable variety differentiation.
Violet aroma quality is a floral aroma quality that represents olfactory perception evoking Viola flower-associated scents including fresh violet, violet candy, iris root (orris), and powdery floral notes, characterizing the distinctive sweet-powdery aromatic impression particularly associated with certain red wine varieties and contributing elegance to their aromatic profile. Within the BFO framework, this quality specifically addresses the violet subcategory of floral aroma, distinguished by its strong association with red wines rather than whites and its derivation from carotenoid-degradation products rather than the monoterpenes responsible for most white wine floral character. The ontological significance lies in violet aroma serving as a key varietal and quality marker for red wines, where prominent violet character indicates varieties such as Malbec (whose violet note is considered definitional), Syrah, and Nebbiolo, and suggests quality and ripeness appropriate for elegant, complex wine expression. The compositional basis for violet aroma centers on β-ionone, a C13-norisoprenoid derived from carotenoid degradation that provides violet character at extremely low detection threshold (0.09 ng/L in water, approximately 0.8 μg/L in wine matrix), making trace concentrations capable of significant aromatic impact. Additional contributors include α-ionone (woody-violet) and certain phenylpropanoid-derived compounds contributing iris-like character in aged wines. The ripeness association of violet character shows increased expression with grape maturity as carotenoid precursor accumulation enables greater β-ionone production, though the compound's volatility means violet intensity may decrease with extended aging as the compound volatilizes or transforms. The color association between violet aroma and purple-hued red wines creates cross-modal enhancement where visual perception of purple color intensifies violet aroma perception, demonstrating sensory integration affecting quality assessment. Understanding violet aroma quality enables varietal identification for violet-associated red varieties, ripeness assessment based on violet development, and quality evaluation where violet elegance indicates successful maturation and winemaking.
Violet_Aroma_Quality addresses the Viola flower subcategory of floral aroma, encompassing fresh violet, violet candy, iris, and powdery floral notes. This quality distinguishes from other floral categories by its strong red wine association and norisoprenoid (rather than monoterpene) chemical origin. The extremely low detection threshold of β-ionone makes violet a potent aromatic contributor even at trace concentrations, and its association with specific varieties provides valuable identification information.
Irischarakter
Veilchenaroma
Veilchenduft
iris character
powdery floral scent
viola aroma
violet flower notes
violette Blütennote
Pineau B et al. (2007) Examples of the roles of specific wine aroma compounds in wine quality assessment. Australian Journal of Grape and Wine Research 13(2): 66-74 Sefton MA et al. (2011) Norisoprenoids in wine. Australian Journal of Grape and Wine Research 17(2): 15-20
Violet Aroma Quality
Viral infection indicator quality assessment in a California nursery certification program evaluated mother vines for symptoms suggesting viral infection requiring laboratory confirmation before propagation approval. Visual screening for viral infection indicator quality identified multiple symptom types across the evaluated selections. Grapevine leafroll-associated virus symptoms showed the characteristic downward rolling of leaf margins with interveinal reddening (red cultivars) or chlorosis (white cultivars) intensifying in late season. Grapevine fanleaf virus symptoms showed the diagnostic fan-shaped leaves with abnormal vein patterns, shortened internodes, and yellow mosaic discoloration. Grapevine red blotch virus symptoms presented as irregular red blotches on leaf blades distinct from the interveinal pattern of leafroll. The viral infection indicator quality screening identified 15% of submitted selections as potentially virus-infected based on foliar symptoms, with subsequent PCR and ELISA testing confirming viral presence in 12% and identifying symptomless infected vines in an additional 8% of selections showing no obvious viral infection indicator quality, demonstrating the necessity of laboratory confirmation beyond visual assessment for certification programs. The comprehensive viral infection indicator quality evaluation and laboratory testing enabled certification of 80% of selections as virus-tested, with infected materials rejected or directed to virus-elimination therapy before release.
Viral infection indicator quality is a pathogen infection indicator quality that inheres in a grapevine organism and represents the ensemble of symptoms and diagnostic markers associated with viral pathogen infection, including the characteristic leaf discolorations, deformations, growth abnormalities, and fruit quality reductions caused by the numerous viruses affecting grapevine, with viral diseases differing from fungal and bacterial diseases in their obligate intracellular replication, their transmission primarily through vegetative propagation and specific invertebrate vectors, and the complete absence of curative treatments making prevention through clean planting material the foundation of viral disease management. This quality captures the plant responses that enable recognition of viral infections, recognizing that grapevine is host to more than 80 identified viruses making it among the most virus-affected of all crop species, with viral diseases causing estimated global yield losses of 10-40% and quality reductions that substantially impact wine value even when yield effects are modest. The ontological significance of viral infection indicator quality lies in its role enabling disease identification for management decisions focused on preventing spread through propagation and vector control, since no treatments can eliminate viruses from infected vines, making symptom-based detection and laboratory confirmation essential for maintaining virus-free planting stock and managing viral disease pressure in established vineyards. The biological basis of viral infection indicators encompasses the disruption of normal plant physiology by viral replication and gene expression, including interference with photosynthesis causing chlorosis and discoloration, disruption of hormone signaling causing growth abnormalities and deformation, impairment of phloem function affecting carbohydrate transport and fruit development, and triggering of plant defense responses that may themselves cause visible symptoms. The latent nature of many grapevine viral infections complicates diagnosis, with some viruses causing clear symptoms only on certain cultivars, under specific environmental conditions, or when present in mixed infections with other viruses, necessitating laboratory testing for definitive diagnosis even when visual symptoms suggest viral etiology.
Viral_Infection_Indicator_Quality is_a Pathogen_Infection_Indicator_Quality, serving as parent class for Leafroll_Symptom_Quality, Fanleaf_Symptom_Quality, Red_Blotch_Symptom_Quality, and Fleck_Symptom_Quality which address specific viral diseases. Grapevine hosts >80 identified viruses making it exceptionally virus-affected. No curative treatments exist clean stock and vector management are only control strategies. Latent infections common, requiring laboratory testing beyond visual diagnosis. Transmission primarily through propagation and specific vectors (mealybugs, nematodes, leafhoppers).
Virose-Zeichen
Virusinfektions-Indikatoren
Viruskrankheits-Symptome
viral disease symptoms
viral pathogen markers
virale Pathogen-Marker
virosis symptoms
virus infection indicators
Martelli GP (2014) Directory of virus and virus-like diseases of the grapevine and their agents. Journal of Plant Pathology 96(1 Supplement):1-136 Naidu RA et al. (2015) Grapevine leafroll disease and associated viruses: a unique pathosystem. Annual Review of Phytopathology 53:613-634 Fuchs M (2020) Grapevine viruses: a multitude of diverse species with simple but overall poorly adopted management solutions in the vineyard. Journal of Plant Pathology 102:643-653
Viral Infection Indicator Quality
Screening for Viral_Resistance_Disposition at INRAE Colmar evaluated responses of 45 Vitis accessions to Grapevine fanleaf virus (GFLV) through controlled graft inoculation using infected rootstock material. Virus presence and titer were monitored by DAS-ELISA at 6, 12, and 24 months post-grafting. Susceptible V. vinifera 'Gewürztraminer' showed systemic infection in 100% of grafted plants within 6 months, with high viral titers (OD₄₀₅ > 2.0) and severe symptoms including leaf deformation, yellow mosaic, and 40-60% yield reduction. The V. rupestris × V. riparia hybrid 'Börner' exhibited strong Viral_Resistance_Disposition with only 15% of grafted plants testing positive at 24 months, low viral titers when detected (OD₄₀₅ 85% of healthy controls), demonstrating that Viral_Resistance_Disposition encompasses both resistance (limiting infection) and tolerance (limiting disease impact despite infection).
A Viral_Resistance_Disposition is a Pathogen_Resistance_Disposition that inheres in a Vitis vinifera plant or plant part and is realized through defense processes that limit infection, replication, systemic movement, or symptom expression by plant viruses affecting grapevine. This disposition represents the capacity to resist or tolerate the diverse viral pathogens that persistently infect grapevine, including nepoviruses (Grapevine fanleaf virus), ampeloviruses (Grapevine leafroll-associated viruses), vitiviruses, marafiviruses, and the recently characterized geminivirids (Grapevine red blotch virus). Unlike fungal and bacterial pathogens that can be controlled through chemical applications, viral infections are incurable once established, making resistance dispositions the primary long-term management strategy alongside use of certified virus-free planting material. Viral resistance mechanisms in plants include restriction of viral entry and initial replication, RNA silencing pathways degrading viral genomes, translation inhibition of viral proteins, impaired cell-to-cell movement through plasmodesmata, blocked systemic transport via phloem, and hypersensitive response eliminating infected cells. Additionally, resistance to viral vectors (nematodes for nepoviruses, mealybugs and soft scales for leafroll viruses) provides indirect protection by preventing virus acquisition and transmission. True genetic resistance to grapevine viruses is rare in Vitis vinifera, with most cultivars susceptible to infection but varying in tolerance (ability to maintain productivity despite infection). Wild Vitis species, particularly North American species co-evolved with endemic nepoviruses, provide sources of both direct viral resistance and vector resistance that can be introgressed into cultivated backgrounds through breeding.
Viral_Resistance_Disposition is_a Pathogen_Resistance_Disposition. This class encompasses resistance to all viral pathogens of grapevine, distinct from Fungal_Resistance_Disposition and Bacterial_Resistance_Disposition which address other pathogen kingdoms. Child classes are organized by viral disease complex: Fanleaf_Resistance_Disposition, Leafroll_Resistance_Disposition, Red_Blotch_Resistance_Disposition, and Fleck_Tolerance_Disposition. Viral resistance may operate directly against the virus or indirectly through vector resistance. The distinction between resistance (limiting infection) and tolerance (limiting impact despite infection) is particularly important for viral diseases where true immunity is rare.
Virale-Krankheitsresistenz
Virusresistenz-Disposition
Virustoleranz
grapevine virus immunity
viral disease tolerance
virus resistance disposition
Martelli GP (2017) An overview on grapevine viruses, viroids, and the diseases they cause, Grapevine Viruses: Molecular Biology, Diagnostics and Management, Springer Oliver JE, Fuchs M (2011) Tolerance and resistance to viruses and their vectors in Vitis sp.: a virologist's perspective of the literature, American Journal of Enology and Viticulture 62:438-451 Maree HJ et al. (2013) Grapevine leafroll-associated virus 3, Frontiers in Microbiology 4:82
Viral Resistance Disposition
A Chardonnay clone 96 vine in the UC Davis FPS screenhouse bears a Virus_Free_Mother_Plant_Role, having tested negative by biological indexing, ELISA, RT-PCR, and HTS for all 47 characterized grapevine viruses plus viroids, qualifying for nuclear stock status and serving as the ultimate source for all clone 96 certified material. A Cabernet Franc clone 214 at ENTAV serves in a Virus_Free_Mother_Plant_Role following thermotherapy and meristem culture that eliminated previously detected GLRaV-3, with subsequent testing confirming successful virus elimination. A Gewürztraminer selection from Alsace achieved Virus_Free_Mother_Plant_Role status after three years of comprehensive testing, enabling its registration as a new certified clone and entry into the French propagation system.
A mother plant role that inheres in a grapevine confirmed to be free from all regulated viral pathogens through comprehensive testing, representing the health status goal of certification programs and the prerequisite for highest-tier (Pre-Basic/Nuclear) classification. Virus-free mother plants have tested negative for all viruses on the applicable regulatory list through appropriate diagnostic methods (biological indexing for graft-transmissible agents, ELISA for serologically detectable viruses, PCR/RT-PCR for molecularly characterized pathogens, and increasingly HTS for comprehensive virome assessment). The virus-free designation reflects current testing capability and regulated pathogen definitions—a plant may harbor uncharacterized or non-regulated viruses while meeting "virus-free" criteria for certification purposes. Virus-free status is not permanent continuous monitoring and isolation are required to maintain freedom from reinfection through vectors (mealybugs, leafhoppers, nematodes) or contaminated equipment. The economic value of virus-free material is demonstrated by yield comparisons showing 20-60% advantages over infected plantings, with additional benefits in fruit quality, vine longevity, and reduced pathogen reservoir for vector transmission.
Virus_Free_Mother_Plant_Role represents the health status outcome that certification systems aim to achieve, providing the "clean" genetic resource from which healthy vineyards are established. The designation is necessarily relative to current detection capability and regulatory definitions the discovery of new viruses (e.g., Grapevine red blotch virus, characterized 2012) may retrospectively reveal that "virus-free" plants harbor previously undetected pathogens. The distinction between regulatory virus-free status (meeting certification requirements) and absolute virus-free status (harboring no viral genetic material) is increasingly relevant as HTS reveals ubiquitous low-level viral sequences of uncertain significance. Maintaining virus-free status requires ongoing vigilance—vector exclusion, equipment sanitation, isolation from potentially infected material—as reinfection is always possible and eliminates the investment in virus elimination.
disease-free vine
gesunde Rebe
krankheitsfreie Mutterpflanze
pathogen-free mother plant
pathogenfreier Mutterstock
virus-free vine
virusfreie Mutterpflanze
Golino DA et al. (2017) Foundation Plant Services, HortScience 52: 1190-1194 European Union (2020) Commission Implementing Directive 2020/177 Martelli GP (2017) Grapevine viruses overview, Springer pp.: 31-46 Al Rwahnih M et al. (2015) High-throughput sequencing for detection of grapevine pathogens, Phytopathology 105: 758-763
Virus Free Mother Plant Role
Virus status for Cabernet Sauvignon clone 337 (ENTAV) is certified as virus-free based on negative results for all eight viruses in the French certification panel: GFLV (Grapevine fanleaf virus), ArMV (Arabis mosaic virus), GFkV (Grapevine fleck virus), GLRaV-1 and GLRaV-3 (Grapevine leafroll-associated virus 1 and 3), GVA and GVB (Grapevine virus A and B), and GPGV (Grapevine Pinot gris virus), confirmed by both ELISA and multiplex RT-PCR at ENTAV-INRA Grau-du-Roi. Virus status for Zweigelt clone Zw-70 documents ELISA-negative results for GFLV, GLRaV-1, GLRaV-3, GVA, GFkV, and ArMV, with additional RT-PCR confirmation for GLRaV-2 and GRSPaV (Grapevine rupestris stem pitting-associated virus), the latter detected at low titer but classified as non-certification-relevant under Austrian phytosanitary regulations.
Virus status is a disease performance data entity that records the presence or absence of specific grapevine viruses and virus-like organisms in a clonal line, determined through a defined panel of diagnostic tests including enzyme-linked immunosorbent assay (ELISA), reverse transcription polymerase chain reaction (RT-PCR), biological indexing on indicator plants, and next-generation sequencing (NGS), documenting the sanitary classification of the clone within the certification framework and establishing the baseline health condition upon which all other performance data should be interpreted.
Virus status determination is the cornerstone of modern grapevine certification systems worldwide, reflecting the recognition that chronic viral infections reduce yield by 15-40%, compromise fruit quality (delayed ripening, reduced color, altered acidity), and shorten vineyard productive lifespan. The diagnostic methodology has evolved substantially — biological indexing on woody indicators (the historical gold standard, requiring 2-3 years) is increasingly supplemented or replaced by serological (ELISA, 1-2 days) and molecular (RT-PCR, real-time RT-PCR, 1-2 days) methods that offer greater sensitivity, specificity, and throughput. Next-generation sequencing (NGS/metagenomics) represents the emerging frontier, capable of detecting all known viruses and potentially novel agents in a single assay without prior knowledge of target sequences. The interpretation of virus status requires understanding that some viruses are near-ubiquitous in older vine populations (GRSPaV is present in >80% of tested accessions globally) while others are strictly excluded from certified material (GFLV, GLRaV-1, GLRaV-3). Mixed infections, where multiple viruses co-infect a single vine, create synergistic symptom expression more severe than any individual virus, making comprehensive panel testing essential. Virus status is not permanent — certified virus-free clones can become reinfected through vector transmission in commercial vineyards, necessitating periodic re-testing of foundation and mother blocks within certification maintenance programs.
Gesundheitsstatus
Virusbefund
Virusfreiheitsstatus
Virusstatus
sanitary status
sanitärer Status
viral health record
viral status
virus infection status
virus testing result
Martelli GP (2014) Directory of virus and virus-like diseases of the grapevine. Journal of Plant Pathology 96(1S): 1-136 Maree HJ et al. (2013) Grapevine leafroll-associated virus 3. Frontiers in Microbiology 4: 82 Mannini F, Digiaro M (2017) The effects of viruses and viral diseases on grapes and wine. In Grapevine Viruses: Molecular Biology, Diagnostics and Management, Springer, pp.: 453-482
Virus Status
A Sangiovese selection from a Tuscan vineyard bears a Virus_Tested_Mother_Plant_Role after undergoing ELISA screening for GFLV, GLRaV-1, GLRaV-2, GLRaV-3, GVA, and GVB—test results showing negative for GFLV and leafroll viruses but positive for GVA, disqualifying it from Certified status until virus elimination treatment but providing documented health information for management decisions. A Grüner Veltliner candidate clone serves in a Virus_Tested_Mother_Plant_Role during the evaluation phase of clonal selection, with comprehensive testing (biological indexing plus molecular assays) determining whether it advances to nuclear stock status or requires pathogen elimination treatment. A Riesling mother vine in a commercial nursery bears a Virus_Tested_Mother_Plant_Role at the CAC level, having tested negative for regulated pathogens within the past triennial testing cycle.
A mother plant role that inheres in a grapevine that has been subjected to laboratory testing for specified viral pathogens, with results documented regardless of outcome, distinguishing it from untested material of unknown health status. Virus-tested mother plants have undergone diagnostic assays (ELISA, PCR, RT-PCR, biological indexing, or high-throughput sequencing) for a defined panel of grapevine viruses, with testing results recorded and available for certification documentation. The "virus-tested" designation does not guarantee virus-free status—a plant may test positive for certain viruses while remaining negative for others—but confirms that health status has been actively assessed rather than assumed. Virus-tested material may be certified at various levels depending on test results plants negative for all regulated pathogens qualify for higher certification categories, while those positive for tolerated pathogens (depending on jurisdiction) may qualify for lower categories or require retesting after attempted elimination. The testing panel varies by certification scheme but typically includes economically significant pathogens (GFLV, GLRaV-1/-3, GVA, GVB, GFkV) causing demonstrated yield and quality impacts.
Virus_Tested_Mother_Plant_Role captures the fundamental distinction between tested and untested material that underlies all certification schemes. "Virus-tested" is a procedural designation (testing has occurred) rather than a health status (virus-free) the distinct Virus_Free_Mother_Plant_Role captures the outcome of negative test results. Testing frequency varies by certification tier and jurisdiction—annual for nuclear stock, triennial for foundation/Basic, and variable (often only at entry) for commercial categories. The evolution from single-pathogen indexing to multiplex molecular assays to untargeted HTS has expanded detection capability while raising questions about how to regulate newly discovered viruses with unknown economic significance. Testing documentation provides the traceability essential for certification and enables epidemiological investigation when pathogen outbreaks occur in nursery material.
assayed stock plant
getesteter Mutterstock
health-tested vine
tested mother plant
virus-tested vine
virusgeprüfte Rebe
virusgetestete Mutterpflanze
Maliogka VI et al. (2015) Grapevine virus diseases, APS Press Rowhani A et al. (2018) Grapevine virus testing and certification protocols, UC Davis FPS EPPO (2008) Certification schemes for pathogen-tested material of grapevine, EPPO Bulletin 38: 422-429 Martelli GP (2017) Grapevine viruses overview, Springer
Virus Tested Mother Plant Role
Virus testing status for Sauvignon Blanc clone 242 (ENTAV) documents comprehensive screening results: GFLV negative by DAS-ELISA and RT-PCR, ArMV negative by DAS-ELISA, GLRaV-1 negative by DAS-ELISA and real-time RT-PCR, GLRaV-3 negative by DAS-ELISA and real-time RT-PCR, GFkV negative by DAS-ELISA, GVA negative by DAS-ELISA, GVB negative by DAS-ELISA, and GPGV negative by RT-PCR, all conducted at ENTAV Le Grau-du-Roi laboratory (ISO 17025 accredited) in September 2022 using dormant cane material collected from foundation mother vine block B-242-F3. Virus testing status for Grüner Veltliner clone 17 records negative results across the Austrian mandatory panel (GFLV, ArMV, GLRaV-1, GLRaV-3, GVA, GFkV) by ELISA, with supplementary RT-PCR confirmation for GLRaV-2 and GRSPaV, the latter detected at low titer (Ct value 31.5) classified as non-relevant for certification under current Austrian regulations.
Virus testing status is a certification status entity that documents the comprehensive results of all diagnostic assays performed on a clonal line to determine its phytosanitary condition with respect to grapevine viruses, viroids, and phytoplasmas, including the specific pathogens tested, the diagnostic methods employed (serological, molecular, biological), the test results (positive, negative, inconclusive), the testing laboratory, and the date of testing, providing the analytical evidence upon which the health component of certification decisions is based.
Virus testing status documentation must capture sufficient methodological detail to enable retrospective evaluation as diagnostic technologies evolve and new viral threats emerge. The mandatory virus testing panel differs between certification authorities — the EU minimum under Directive 2005/43/EC requires testing for GFLV, ArMV, GLRaV-1, and GLRaV-3 for Pre-basic material, while national implementations frequently extend the panel to include GFkV, GVA, GVB, and increasingly GPGV and GRSPaV. The interpretation of testing results requires understanding assay sensitivity and specificity — ELISA detection limits are typically 1-10 ng/mL viral coat protein (suitable for high-titer infections but potentially missing low-titer or unevenly distributed viruses), while RT-PCR achieves sensitivity 100-1000× greater but is more susceptible to false positives from contamination. The seasonal timing of sample collection influences detection probability — leafroll viruses are most reliably detected from phloem tissue collected during late summer through leaf fall (August-November in the Northern Hemisphere), while GFLV detection is optimal in dormant cane tissue during winter. The emergence of next-generation sequencing (NGS/high-throughput sequencing) as a universal detection platform is progressively transforming virus testing from targeted pathogen-specific assays toward comprehensive virome characterization, though regulatory frameworks have not yet fully incorporated NGS-based results into certification requirements.
Gesundheitstestung
Virusscreeningergebnis
Virustestungsstatus
Virusuntersuchungsergebnis
diagnostic testing status
health testing documentation
phytosanitary testing record
phytosanitärer Testbefund
virus screening results
virus test results
Martelli GP (2014) Directory of virus and virus-like diseases of the grapevine. Journal of Plant Pathology 96(1S): 1-136 Mannini F, Digiaro M (2017) The effects of viruses and viral diseases on grapes and wine. In Grapevine Viruses, Springer, pp.: 453-482 Maree HJ et al. (2013) Grapevine leafroll-associated virus 3. Frontiers in Microbiology 4: 82
Virus Testing Status
Viscoelastic characterization of Muscat table grape berries using dynamic mechanical analysis reveals storage modulus of 0.85 MPa and loss modulus of 0.18 MPa at 1 Hz loading frequency, with the loss tangent (tan δ = 0.21) indicating significant viscous contribution to mechanical response, explaining why rapid squeezing during consumer assessment yields firmer perception than slow compression, and why prolonged holding during packaging causes progressive berry deformation not predicted by instantaneous elastic modulus alone.
Viscoelastic property quality is a physical quality that inheres in grapevine tissues and represents the combined viscous and elastic mechanical behavior where deformation response depends on both the magnitude and the time history of applied stress, exhibiting characteristics intermediate between purely elastic solids and purely viscous fluids. Within the BFO framework, this quality characterizes the time-dependent mechanical behavior arising from the complex polymer architecture of plant cell walls and the fluid-filled cellular structure, where immediate elastic response is followed by time-dependent viscous flow under sustained loading. The ontological significance lies in viscoelasticity determining how tissues respond to dynamic loading conditions encountered during growth, wind exposure, handling, and mastication, where the rate of force application affects the apparent stiffness and the duration of loading affects the extent of permanent deformation. Grapevine tissues exhibit viscoelastic behavior because their mechanical properties derive from hydrated polymer networks (cellulose, hemicellulose, pectin) that can rearrange under sustained stress, combined with cellular fluid that redistributes under pressure gradients, creating time-dependent deformation patterns not captured by purely elastic models. The viscoelastic response is characterized by phenomena including creep (continued deformation under constant stress), stress relaxation (declining stress under constant strain), and strain-rate dependence of apparent modulus (stiffer response to rapid loading). Temperature significantly affects viscoelastic behavior, with higher temperatures increasing the viscous component and reducing time constants for relaxation processes, while lower temperatures shift behavior toward more elastic response. Measurement approaches include dynamic mechanical analysis applying oscillatory loads to determine storage and loss moduli, creep testing under constant stress, stress relaxation testing under constant strain, and time-dependent recovery analysis following load removal. Understanding viscoelastic properties enables prediction of tissue behavior under realistic dynamic loading, interpretation of texture perception during consumption, and optimization of handling protocols to minimize damage from time-dependent mechanical effects.
Viscoelastic_Property_Quality serves as the parent class for specific viscoelastic phenomena (Creep_Behavior, Stress_Relaxation), establishing the ontological framework for time-dependent mechanical behavior. This quality distinguishes from purely elastic behavior where deformation depends only on current stress state. The class supports dynamic mechanical analysis interpretation, texture perception modeling, and handling damage prediction requiring consideration of loading rate and duration effects.
dynamic mechanical property
dynamisch-mechanische Eigenschaft
rheological property
rheologische Eigenschaft
time-dependent elasticity
viscous-elastic behavior
viskoelastische Eigenschaft
zeitabhängige Elastizität
Bourne MC (2002) Food Texture and Viscosity: Concept and Measurement, 2nd Ed. Academic Press, San Diego, pp. 182-213 Jackman RL, Stanley DW (1995) Perspectives in the textural evaluation of plant foods. Trends in Food Science & Technology 6(6): 187-194
Viscoelastic Property Quality
Viscosity appearance assessment comparing Mosel Riesling Kabinett (thin, rapidly draining film with minimal leg formation indicating low alcohol and light body) with Amarone della Valpolicella (thick, slowly draining film with pronounced, persistent legs indicating 15%+ alcohol and full body) demonstrates the visual body differentiation possible before tasting, with viscosity observation creating appropriate expectation for the delicate, refreshing palate of the former versus the rich, powerful palate of the latter.
Viscosity appearance quality is a visual appearance quality that represents the perceived thickness, flow resistance, and textural impression of wine observed through swirling behavior and drainage patterns on glass surfaces, characterizing the visual correlates of wine body that provide indirect information about alcohol content, glycerol concentration, and extract level. Within the BFO framework, this quality addresses the rheological dimension of visual assessment, distinguishing viscosity-related observations from color and clarity while recognizing that viscosity assessment provides body and concentration cues through purely visual examination without tactile contact. The ontological significance lies in viscosity appearance serving as a visual proxy for mouthfeel prediction, where observed flow characteristics create expectations for palate weight and texture that frame subsequent gustatory evaluation, though the correlation between visual viscosity and actual mouthfeel involves complex relationships among multiple compositional factors. The physical basis for viscosity appearance involves the resistance of wine to flow under gravity, determined primarily by alcohol content (increasing viscosity above approximately 12% through hydrogen bonding effects), glycerol concentration (contributing to oily texture), residual sugar (increasing density and viscosity), and extract level (dissolved solids adding to flow resistance), with these factors interacting to produce the observed flow behavior. The assessment methodology for viscosity involves swirling wine in the glass and observing drainage patterns, with thin, rapidly draining wines appearing watery while thick, slowly draining wines with pronounced leg formation appearing viscous and full-bodied. The interpretation limitations of viscosity assessment are significant, as alcohol contributes more to leg formation than glycerol despite common misconception, and visual viscosity correlates imperfectly with actual mouthfeel where acidity, tannin, and carbonation modify perceived body independent of true viscosity. Understanding viscosity appearance quality enables body prediction from visual cues, alcohol level estimation from leg behavior, and appropriate expectation setting for palate assessment while acknowledging the limitations of visual-to-tactile inference.
Viscosity_Appearance_Quality serves as the parent class for viscosity-related visual qualities (Legs, Body Visual Impression), addressing the rheological dimension of visual assessment. This quality provides body and concentration cues through observation of flow behavior, though the relationship between visual viscosity and actual mouthfeel is complex and imperfect. The class supports pre-tasting body prediction while acknowledging interpretation limitations.
Dickflüssigkeit
Fließverhalten
Viskositätserscheinung
flow characteristics
rheological quality
thickness appearance
visuelle Körperwahrnehmung
wine body visual
Pickering GJ et al. (1998) The influence of ethanol concentration on the perception of viscosity and density in white wine. Journal of Food Science 63(5): 845-848 Nurgel C, Pickering GJ (2005) Contribution of glycerol, ethanol and sugar to the perception of viscosity and density elicited by model white wines. Journal of Texture Studies 36(3): 303-323
Viscosity Appearance Quality
Viscosity description for Amarone della Valpolicella (2020 vintage, 16.0% alcohol, 5.5 g/L RS) records prominent, slow-moving, densely packed legs descending at approximately 2-3 mm/s after swirling, with a viscous, almost oily meniscus appearance — the full viscosity consistent with the wine's extreme alcohol concentration, elevated glycerol (12-15 g/L typical for Amarone from partially dried grapes), and residual sugar that collectively produce the characteristically dense, full-bodied mouthfeel. Viscosity description for Mosel Riesling Kabinett (2024 vintage, 8.5% alcohol, 45 g/L RS) documents thin, rapidly descending, widely spaced legs despite the significant residual sugar, as the low alcohol produces minimal Marangoni effect — demonstrating that the leg phenomenon is driven primarily by ethanol evaporation rather than sugar or glycerol content. Viscosity description for Grüner Veltliner Smaragd from the Wachau (2024 vintage, 13.5% alcohol) records medium-full legs of moderate speed and density, consistent with the moderate-to-high alcohol content.
Viscosity description is a visual description entity that documents the apparent viscosity and surface tension behavior of wine as visually assessed through the observation of flow patterns, meniscus formation, and the Marangoni effect (the formation, speed, and density of "legs" or "tears" descending from the wetted glass surface above the wine line), characterized on a scale from thin/watery (rapid, widely spaced, barely visible legs) through light, medium, and medium-full to full/viscous (slow, dense, prominent legs), providing a visual proxy for the wine's ethanol, glycerol, and residual sugar content that indicates body weight and confirms stylistic expectations established by the wine's classification and analytical parameters.
Viscosity description through the observation of wine legs or tears is one of the most widely practiced yet commonly misinterpreted visual assessment techniques in wine evaluation. The Marangoni effect that produces the leg phenomenon is driven primarily by the differential evaporation rate between ethanol and water at the thin film left on the glass surface above the wine line — ethanol evaporates more rapidly from this thin film, locally increasing surface tension (pure water has higher surface tension than ethanol-water mixtures), which draws additional wine upward along the glass until gravity causes the accumulated liquid to descend as visible rivulets or "legs." This physical mechanism means that leg formation correlates most strongly with alcohol content (the primary driver of the surface tension differential) rather than with sugar, glycerol, or "body" as commonly believed. The speed of leg descent is influenced by the viscosity of the film, which increases with glycerol concentration (contributing to slower, more visible legs) and residual sugar (increasing liquid viscosity), but these effects are secondary to the ethanol-driven Marangoni flow. The practical diagnostic value of viscosity observation in the tasting context is primarily confirmatory — viscosity assessment corroborates expectations established by the wine's declared alcohol content, known style, and region of origin. Discrepancies between expected and observed viscosity may indicate analytical issues (incorrect alcohol declaration), unexpected residual sugar, or high extract concentration. The description vocabulary progresses from "watery/thin" (wines below 10% alcohol) through "light" (10-11.5%), "medium" (11.5-13%), "medium-full" (13-14.5%) to "full/viscous" (above 14.5% or high RS), though these thresholds are approximate and influenced by other compositional factors. Temperature affects viscosity observation — cooler wines show reduced Marangoni effect due to lower evaporation rates, while warmer wines display more pronounced leg formation.
Schlierenbeschreibung
Tränenbildung
Viskositätsbeschreibung
Weinviskosität
body indication
flow behavior description
legs description
tears description
wine viscosity
Vuilleumier C et al. (1995) The Marangoni effect and wine tears. Physics of Fluids 7(10): 2291-2293 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 467-470 Waterhouse AL et al. (2016) Understanding Wine Chemistry, Wiley, pp.: 35-55 Nikolov AD et al. (2018) Marangoni effect, wine tears, and the ring stain. Current Opinion in Colloid and Interface Science 36: 1-11
Viscosity Description
Viscosity mouthfeel comparison between two full-bodied Chardonnays of equivalent alcohol reveals different textural profiles, with Burgundy Meursault showing silky, flowing viscosity (6/10) from balanced malo and moderate lees contact creating elegant coating that clears smoothly, while California counterpart shows oily, unctuous viscosity (8/10) from extended sur lie aging and higher glycerol creating more persistent coating that some perceive as rich complexity and others as excessive heaviness, demonstrating how viscosity quality distinguishes wines of similar body and contributes to style character.
Viscosity mouthfeel quality is a mouthfeel quality that represents the perceived resistance to flow and oral movement of wine, characterizing the coating, slippery, oily, or watery tactile impression arising from wine's rheological properties as they interact with oral surfaces during consumption. Within the BFO framework, this quality specifically addresses the flow resistance dimension of mouthfeel, distinguishing viscosity perception from body (overall weight) while recognizing that viscosity contributes to body impression but represents a distinct tactile quality related to how wine moves and coats in the mouth. The ontological significance lies in viscosity mouthfeel providing the textural nuance that distinguishes wines of similar body, where two wines may have equivalent weight but differ in whether they feel silky, oily, or watery, with viscosity quality affecting elegance perception and overall mouthfeel harmony. The perceptual basis for viscosity mouthfeel involves mechanoreceptor detection of shear resistance as wine moves across oral surfaces, with higher viscosity wines showing greater resistance to movement and more persistent coating while lower viscosity wines move easily and clear quickly from oral surfaces. The compositional determinants of viscosity include alcohol (increasing viscosity through hydrogen bonding, particularly above 12%), glycerol (contributing oily texture), residual sugar (adding viscosity proportional to concentration), polysaccharides (significantly enhancing smoothness and coating at low concentrations), and the combined effect of these factors creating the characteristic textural profile. The descriptive terminology for viscosity mouthfeel ranges from watery (minimal viscosity, rapid clearing) through flowing and silky (moderate viscosity, smooth movement) to oily, unctuous, and viscous (high viscosity, slow coating movement), with these terms capturing the qualitative character of flow behavior. Understanding viscosity mouthfeel quality enables textural characterization beyond simple body assessment, quality evaluation incorporating viscosity appropriateness for style, and appreciation of how flow properties contribute to overall mouthfeel elegance or heaviness.
Viscosity_Mouthfeel_Quality addresses the flow resistance dimension of oral tactile perception, distinct from overall body. This quality provides textural nuance distinguishing wines of similar weight, contributing to elegance or heaviness perception. The class supports detailed mouthfeel assessment beyond body category and appreciation of how compositional factors create different textural profiles.
Beschichtungsgefühl
Fließtextur
Viskositätsmundgefühl
coating texture
flow sensation
oral viscosity
orale Viskosität
wine slipperiness
Nurgel C, Pickering GJ (2005) Contribution of glycerol, ethanol and sugar to the perception of viscosity and density elicited by model white wines. Journal of Texture Studies 36(3): 303-323 Smith AK et al. (1996) The influence of ethanol on the temporal perception of sweetness, sourness, bitterness, and the flavour of white wine. Food Quality and Preference 7(3-4): 291-295
Viscosity Mouthfeel Quality
Syrah on Ramsey rootstock visible incompatibility zone at 7 years post-grafting shows 22-35 mm zone with pronounced observable symptoms= excessive swelling 165-185% diameter creating dramatic 4-7 cm radial bulge clearly visible from distance, extreme asymmetry with 2× greater diameter on one side creating lopsided kidney-shaped cross-section, bark showing multiple longitudinal cracks 2-5 mm wide and 8-15 cm long exposing underlying tissue, necrotic bark patches 3-6 cm diameter appearing dark brown-grey contrasting with normal bark, union showing 20° bend from straight trunk axis from differential growth, caliper measurements documenting swelling increasing from 135% year 3 to 185% year 7, visual symptoms enabling immediate identification as severely incompatible union requiring vine removal and replanting with compatible combination
Visible incompatibility zone is incompatibility zone manifesting observable external symptoms including excessive union swelling, bark abnormalities, structural deformities, or asymmetries enabling visual detection without dissection, representing incompatibility expression producing readily identifiable indicators distinguishing problematic unions from successfully healed grafts. Visible incompatibility characteristics includes= excessive swelling (union circumference >130-200% of adjacent trunk diameter creating pronounced bulge easily detected visually or by measurement), asymmetric enlargement (extreme eccentricity with union showing disproportionate growth on certain sides creating lopsided appearance), bark abnormalities (cracks, fissures, cankers, or necrotic zones visible on union surface indicating internal problems), structural discontinuities (visible steps or abrupt diameter changes at union rather than smooth transition), color differences (bark discoloration at union, darker or lighter zones suggesting tissue stress or necrosis), and deformation (twisted, bent, or irregular union form contrasting with relatively straight uniform trunk above and below). Visible incompatibility excessive swelling manifestation includes= moderate excessive swelling (130-150% diameter, readily noticeable but vine potentially functional though compromised), severe excessive swelling (150-200%+ diameter, creating dramatic bulge unmistakably indicating incompatibility), circumferential measurement (caliper measurements quantifying swelling comparing union to adjacent trunk sections), progressive enlargement (swelling increasing over years detected by repeated measurements), compensatory growth (excessive swelling potentially representing attempt to compensate for inadequate vascular connections through increased tissue production), and tissue composition (swelling comprising disorganized callus, abnormal vascular tissues rather than organized secondary growth). Visible incompatibility bark symptoms includes= cracking and fissuring (bark splitting longitudinally or circumferentially at union from internal pressure, tissue expansion, or necrosis), bark necrosis (dead bark patches appearing brown, black, or grey contrasting with healthy tan-brown bark), bark separation (peeling or lifting bark at union indicating underlying tissue death or poor adhesion), canker formation (localized dead areas with sunken or raised margins indicating tissue death and attempted wound healing), exudation (gum or sap oozing from union cracks or necrotic zones indicating tissue stress or pathogen presence), and abnormal texture (rough, warty, or irregular bark surface at union contrasting with normal smooth or moderately furrowed bark above and below). Visible incompatibility structural deformity includes= pronounced bulging (union protruding substantially 2-6 cm radially beyond normal trunk contour), abrupt diameter transitions (sharp changes in trunk diameter at union boundaries rather than gradual tapering), asymmetric form (union showing extreme one-sided development creating lopsided or kidney-shaped cross-section), bent or twisted union (junction showing deviation from straight trunk axis potentially from differential growth stresses), overgrowth patterns (scion or rootstock showing excessive tissue formation partially or completely covering graft interface creating characteristic morphology), and instability (union showing movement, flexing, or visible gaps under pressure indicating poor mechanical integration). Visible incompatibility color and appearance includes= bark discoloration (union bark showing different color than adjacent trunk, often darker grey-brown or lighter tan suggesting stress or abnormal tissue), discolored wood (exposed wood at cracks showing brown staining indicating tissue death or phenolic accumulation), contrasting tissue appearance (smooth shiny callus-like tissue visible at union contrasting with normal rough bark texture), and weathering patterns (union aging or deteriorating differently than normal bark showing distinctive appearance over time). Visible incompatibility detection methods includes= visual inspection (walking vineyard observing unions detecting excessive swelling, bark problems, deformities), caliper measurements (measuring union diameter comparing with trunk above and below quantifying swelling percentage), photographic documentation (images enabling comparison over time detecting progressive changes), trunk strength assessment (attempting to flex union detecting excessive movement or weakness indicating poor integration), and comparative assessment (comparing suspect unions with healthy grafts or different combinations identifying abnormal characteristics). Visible incompatibility early detection importance includes= enabling early intervention (detecting problems within first 2-3 years allowing culling and replanting before extensive vineyard establishment investment), guiding management decisions (visible symptoms indicating unions requiring special attention, support, or monitoring), facilitating quality control (nursery or vineyard managers identifying and removing severely incompatible unions improving stand uniformity), preventing vineyard establishment with poor unions (detecting problematic grafts before vineyard planting avoiding future decline or failure), and enabling compatibility assessment (visible symptoms providing field indicators for evaluating new scion-rootstock combinations). Visible incompatibility relationship to internal problems includes= external symptoms reflecting internal dysfunction (excessive swelling, bark necrosis indicating underlying vascular discontinuities, tissue necrosis), severity correlation (more pronounced visible symptoms generally indicating more severe internal incompatibility though correlation imperfect), some internal problems invisible (moderate incompatibility potentially showing minimal external symptoms despite significant vascular or mechanical issues), and dissection verification (cutting through visibly abnormal unions confirming internal necrosis, vascular problems, tissue disorganization). Visible incompatibility in different combinations and conditions includes= characteristic patterns by combination (some scion-rootstock pairs showing typical visible incompatibility symptoms if incompatible), environmental modulation (stress potentially making incompatibility more visible, exacerbating swelling or bark problems), age progression (visible symptoms often becoming more pronounced with vine age as incompatibility progresses), and variable expression (some unions showing dramatic visible symptoms while others with similar incompatibility showing subtler external signs). Visible incompatibility management responses includes= culling severely affected unions (removing vines with excessive swelling >150%, extensive bark necrosis, structural deformity within first 2-3 years), monitoring moderately affected unions (tracking unions with 130-145% swelling or minor bark problems assessing progression determining if acceptable or requiring removal), providing support (staking or tying unions showing structural weakness preventing breakage), protective measures (wrapping or painting unions with bark problems reducing stress, disease entry, environmental damage), and evaluation for replanting (assessing whether incompatibility severity justifies vineyard replanting with different compatible combination). Visible incompatibility comparison with normal union appearance includes= normal healed unions showing moderate smooth swelling 105-118% versus excessive irregular swelling >130% in incompatible, normal showing intact continuous bark versus cracked necrotic bark in incompatible, normal showing relatively symmetric form versus extreme asymmetry in incompatible, normal showing smooth transition versus abrupt steps in incompatible, and normal showing stable appearance over years versus progressive deterioration in incompatible. Visible incompatibility significance includes= enabling non-destructive assessment (visible symptoms allowing union evaluation without vine sacrifice or destructive sampling), facilitating vineyard management (visible indicators guiding decisions about union support, protection, monitoring, or removal), supporting compatibility evaluation (observable symptoms providing practical field criteria for assessing scion-rootstock combinations), and emphasizing proper selection (readily visible incompatibility consequences highlighting importance of using proven compatible combinations, proper grafting technique, quality control in nursery and vineyard establishment).
Visible_Incompatibility_Zone emphasizes observable external symptoms and is distinguished from Incompatibility_Zone (general failed integration), Delayed_Incompatibility_Zone (temporal emphasis), and internal incompatibility (requiring dissection) by focus on readily detected external indicators. Critical for understanding non-destructive incompatibility assessment, visual detection methods, external symptom interpretation, early problem identification, vineyard quality control, and visible symptoms as practical field indicators enabling union evaluation and management decisions without requiring destructive sampling or specialized diagnostics.
Apparent union failure
Erkennbare Abstoßung
External incompatibility symptoms
Observable incompatibility
Sichtbare Inkompatibilität
Äußerlich sichtbare Verwachsungs-Störung
Visible Incompatibility Zone
Visual appearance quality assessment in professional blind tasting of red wine examines clarity (brilliant, no haze), color intensity (deep), color hue (ruby with purple rim suggesting youth), and viscosity (medium-high legs suggesting elevated alcohol), enabling preliminary identification as young, full-bodied red before aromatic assessment begins, with these visual cues creating expectations of concentrated fruit and firm structure that frame interpretation of subsequent olfactory and taste impressions, demonstrating visual assessment's role in establishing the evaluative context for complete sensory analysis.
Visual appearance quality is a sensory quality that represents the optical properties of wine perceived through visual examination, encompassing clarity, color, viscosity, and effervescence attributes that provide the first sensory impression and offer diagnostic information about wine identity, condition, and quality before olfactory and gustatory assessment begins. Within the BFO framework, this quality addresses the visual dimension of wine sensory evaluation, distinguishing optically-perceived attributes from chemosensory qualities while recognizing that visual assessment creates expectations that influence subsequent olfactory and taste perception through cross-modal cognitive effects. The ontological significance lies in visual appearance providing immediate quality indicators accessible to all observers regardless of sensory training, where clarity suggests careful production, appropriate color indicates variety and age, and viscosity hints at body and alcohol level, making visual assessment the universal entry point to wine evaluation that shapes the interpretive framework for subsequent sensory analysis. The perceptual basis for visual assessment involves light interaction with wine in the glass, including transmission (determining transparency and color saturation), reflection (creating brilliance and surface character), and scattering (producing haziness when particles are present), with these optical phenomena enabling assessment of wine physical state and compositional properties without direct contact. The compositional determinants of visual appearance include phenolic compounds (anthocyanins and tannins determining color), suspended particles (affecting clarity), glycerol and alcohol content (influencing viscosity and legs), and dissolved carbon dioxide (creating effervescence), with visual properties thus providing indirect compositional information. The evaluation protocol for visual assessment involves standardized conditions including white background, appropriate lighting, tilted glass examination for color gradient assessment, and swirling for viscosity observation, with systematic visual examination preceding aromatic assessment in professional tasting protocols. Understanding visual appearance quality enables quality screening based on optical properties, variety and age estimation from color characteristics, and condition assessment detecting faults visible before aromatic evaluation.
Visual_Appearance_Quality serves as the parent class for visual assessment categories (Clarity, Color, Viscosity, Effervescence), establishing the framework for optically-perceived wine properties. This quality represents the first sensory impression in evaluation sequence, providing information that shapes expectations for subsequent assessment. The class addresses a distinct sensory modality (vision) from the chemosensory qualities (olfaction, gustation) that dominate wine evaluation.
Erscheinungsbild
Weinaussehen
optical appearance
optische Qualität
visual assessment
visual quality
visuelle Erscheinung
wine appearance
Jackson RS (2014) Wine Science: Principles and Applications, 4th Ed. Academic Press, San Diego, pp. 831-887 Peynaud E, Blouin J (1996) The Taste of Wine: The Art and Science of Wine Appreciation, 2nd Ed. John Wiley & Sons, New York, pp. 63-91
Visual Appearance Quality
Visual description for Barolo DOCG (2018 vintage, 6 years post-harvest) records a medium garnet-ruby core with pronounced orange-brick rim evolution extending 5-8 mm from the meniscus, star-bright clarity with no haze or particulate, and medium-high viscosity with slow, well-defined legs — the advanced rim color signaling the characteristic rapid oxidative color evolution of Nebbiolo anthocyanins (low total anthocyanin, predominantly non-acylated peonidin and cyanidin glucosides) that distinguishes aged Barolo from slower-evolving thick-skinned varieties. Visual description for Grüner Veltliner Federspiel from the Wachau (2024 vintage) documents pale straw-yellow with subtle green-gold reflections, brilliant clarity, and light-medium viscosity with fine, rapid legs, consistent with a youthful, medium-bodied dry white wine from a cool continental climate.
Visual description is a sensory description entity that documents the appearance characteristics of wine as assessed by human visual evaluation under standardized lighting conditions, encompassing color (hue, intensity, and rim variation), clarity (brilliance, turbidity, and particulate status), and viscosity (body indication through surface tension and glycerol-related flow behavior), observed by tilting a partially filled ISO tasting glass against a white background under diffuse daylight or D65 illuminant-equivalent artificial lighting, and providing the first sensory impression that establishes expectations for wine style, age, concentration, and potential quality.
Visual description provides the opening chapter of wine sensory assessment, establishing a perceptual framework that primes subsequent olfactory and gustatory evaluation — research demonstrates that visual cues systematically influence aroma and flavor perception through cross-modal cognitive interactions, making appearance evaluation both informative in its own right and consequential for the entire tasting experience. The standardization of visual assessment conditions is essential for reproducible evaluation — the ISO 3591 tasting glass (215 mL, tulip-shaped, transparent crystal) filled to approximately one-third capacity and tilted at 45° against a white background provides the standardized observation geometry, while consistent ambient lighting (daylight or D65 illuminant at 500-1000 lux, avoiding colored light sources) ensures color perception accuracy. The three principal dimensions of visual assessment — color, clarity, and viscosity — provide complementary information about wine composition, production history, and condition. Color reveals cultivar identity (Pinot Noir's distinctive pale ruby versus Syrah's opaque purple-black), age trajectory (the progressive shift from purple-blue through red-garnet to tawny-brown reflecting anthocyanin polymerization and oxidation), and concentration level (intensity correlating with skin extraction and vine yield). Clarity indicates filtration status, protein and tartrate stability, and potential microbial activity. Viscosity signals alcohol and glycerol content through the Marangoni effect (surface tension-driven flow producing the "legs" or "tears" on the glass surface). The visual assessment of sparkling wines adds the additional dimension of mousse quality — bubble size, persistence, bead formation, and crown foam development providing quality indicators that reflect the wine's protein and surfactant composition, CO₂ pressure, and glass surface nucleation site characteristics.
Aussehen
appearance description
optical description
optische Bewertung
visual evaluation
visuelle Beschreibung
visuelle Beurteilung
Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 461-485 Morrot G et al. (2001) The color of odors. Brain and Language 79(2): 309-320 ISO 3591:1977 Sensory analysis — Apparatus — Wine-tasting glass Parr WV et al. (2003) The nose knows: influence of colour on perception of wine aroma. Journal of Wine Research 14(2-3): 79-101
Visual Description
The viticultural practice description for Domaine de la Romanée-Conti's Romanée-Conti vineyard documents an integrated management system comprising organic certification (since 1985) and biodynamic practices (since 1997), Guyot simple training with 1 cane of 8 buds, planting density of 10,000 vines per hectare on 1.0 x 1.0 m spacing, selection massale propagation from pre-phylloxera-origin genetic material, manual debudding to 6-8 shoots per vine, systematic green harvest at véraison to limit yields to 20-25 hl/ha, hand harvesting with three successive passes (tries) selecting only optimally ripe clusters, and a detailed soil management program including biodynamic preparations 500 and 501 applied according to the Maria Thun astronomical calendar.
A grapevine information entity that documents the methods, techniques, protocols, and management decisions applied in the cultivation of Vitis vinifera, encompassing all recorded knowledge about vineyard establishment, canopy management, soil management, irrigation, pest and disease control, harvest operations, and other agronomic interventions that constitute the human component of terroir expression and wine quality determination.
Viticultural_Practice_Description is the umbrella class for all documented information about vine cultivation methods and vineyard management decisions. This class bridges the gap between the theoretical knowledge encoded in viticultural science and the practical implementation of that knowledge in specific vineyard contexts: the description of viticultural practices serves multiple purposes including regulatory compliance documentation (organic certification records, appellation production rule verification), precision viticulture decision support (linking management actions to vine response and wine quality outcomes), knowledge transfer between practitioners, and historical documentation of evolving management paradigms. Subclasses are organized by viticultural domain, with Training_System_Description as the first major subclass covering the architectural framework within which the vine is managed.
Anbautechnik-Beschreibung
Bewirtschaftungsdokumentation
Weinbaupraxis-Beschreibung
vineyard practice documentation
viticultural technique description
viticulture management record
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles Jackson RS (2020) Wine Science: Principles and Applications 5th Ed, Academic Press
Viticultural Practice Description
Viticultural quality assessment comparing management systems in Cabernet Sauvignon reveals the integrated performance differences affecting production outcomes, with conventional high-input system showing high yield quantity (12 t/ha) but moderate fruit quality (uneven ripening, dilute color), while precision-managed system shows moderate yield (8 t/ha) with high fruit quality (uniform ripening, concentrated composition), demonstrating how viticultural quality encompasses both quantity and quality dimensions that may require tradeoffs in management optimization depending on production objectives and market positioning.
Viticultural quality is a quality that represents the agronomic attributes and management-responsive characteristics of grapevine cultivation, characterizing the production parameters, vine performance indicators, and cultivation outcomes that determine the economic viability of grape growing and the raw material quality available for winemaking or other utilization. Within the BFO framework, this quality addresses the agricultural dimension of grapevine systems, distinguishing the cultivation-related attributes from morphological structures, physiological processes, or phenological events while recognizing that viticultural quality emerges from the interaction of genetics, environment, and management—the triad that determines what any vineyard actually produces and how well it produces it. The ontological significance lies in viticultural quality capturing the outcome measures that determine whether grape growing achieves its practical purposes, where yield quantity affects economic returns, yield quality affects product value, vine balance affects both current production and long-term vineyard sustainability, and the integration of these factors determines the overall success of viticultural enterprise. The scope of viticultural quality encompasses yield parameters (the quantity and components of grape production), balance indicators (the relationships among vegetative growth, reproductive output, and resource availability that affect vine sustainability), fruit composition (the chemical constituents affecting product quality), and management responsiveness (how effectively vines respond to cultural interventions), with these dimensions together characterizing the agronomic performance that management seeks to optimize. The interaction between viticultural quality and wine quality recognizes that viticultural decisions establish the raw material constraints within which winemaking operates, with superior viticultural quality providing the compositional foundation that enables rather than limits wine quality outcomes, while compromised viticultural quality creates compositional deficits that winemaking cannot fully overcome. Understanding viticultural quality enables assessment of vineyard performance, identification of factors limiting production or quality, and management optimization targeting the specific quality dimensions most relevant for production objectives.
Viticultural_Quality serves as the parent class for yield, balance, and management-responsive quality dimensions characterizing agronomic performance. This quality addresses the cultivation outcomes determining economic viability and raw material quality. The class supports vineyard performance assessment through viticultural quality characterization.
Kultivierungsqualität
Produktionsqualität
agronomic quality
agronomische Qualität
cultivation quality
production quality
vineyard performance
weinbauliche Qualität
Jackson DI, Lombard PB (1993) Environmental and management practices affecting grape composition and wine quality - a review. American Journal of Enology and Viticulture 44(4): 409-430 Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 145-230
Viticultural Quality
41B Millardet et de Grasset (V. vinifera Chasselas × V. berlandieri) maximizes lime tolerance (40% active lime) through substantial V. berlandieri contribution, supporting Chardonnay on Champagne chalk, though with reduced phylloxera resistance from the V. vinifera parent. SO4 (V. berlandieri × V. riparia) inherits lime tolerance (17-20% active lime) and late budburst from V. berlandieri, combined with improved rooting and cold hardiness from V. riparia. 110 Richter (V. berlandieri × V. rupestris) combines V. berlandieri's lime tolerance with V. rupestris's deep rooting, creating the archetypal Mediterranean rootstock tolerating both calcareous soils and drought.
A parentage-based rootstock role that inheres in a grafted rootstock by virtue of its genetic derivation from Vitis berlandieri Planchon (synonym V. cinerea var. helleri), the Spanish grape or winter grape native to the limestone hills and canyons of the Edwards Plateau in central Texas and adjacent Mexico. V. berlandieri contributes characteristic traits that are essential for viticulture on calcareous soils, including exceptional lime tolerance (tolerating 40-60% active limestone through efficient Fe³⁺-chelate reductase activity and rhizosphere acidification), good-to-excellent drought tolerance (deep rooting on permeable substrates, efficient water use), moderate vigor, and strong phylloxera resistance. V. berlandieri shows poor cold hardiness (surviving only to approximately -15°C), poor rooting from hardwood cuttings (rooting rates often below 30% without special treatment), and late budburst providing some spring frost protection. The species is the dominant genetic contributor to modern rootstock breeding, present in the parentage of approximately 70% of commercial rootstocks worldwide through the V. berlandieri × V. riparia (SO4, 5BB, 5C, etc.) and V. berlandieri × V. rupestris (110R, 99R, 1103P, 140Ru) hybrid groups.
Vitis_Berlandieri_Derived_Rootstock_Role represents the most important genetic contributor to modern rootstock breeding due to the unique lime tolerance of this species—a trait essentially absent in other American Vitis species. V. berlandieri evolved on the highly calcareous soils of the Texas Hill Country (Edwards Plateau limestone), developing iron-efficiency mechanisms unmatched in the genus. The species' poor rooting necessitated crossing with easier-rooting species (V. riparia, V. rupestris) for commercial propagation, creating the dominant rootstock hybrid groups. Climate change may increase lime chlorosis incidence in some regions, potentially expanding demand for berlandieri-derived rootstocks.
Berlandieri-Unterlage
Spanish grape derived rootstock
V. berlandieri derived rootstock
V.-berlandieri-stämmige Unterlage
berlandieri rootstock
von Vitis berlandieri abstammende Unterlage
winter grape derived rootstock
Heinitz CC et al. (2019) Crop wild relatives of grape, in North American Crop Wild Relatives Vol 2, Springer pp.: 329-351 Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22 Bavaresco L et al. (2003) Grapevine iron nutrition update, Journal International des Sciences de la Vigne et du Vin 37: 89-100 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia
Vitis Berlandieri Derived Rootstock Role
Ramsey (Salt Creek, V. champinii selection) provides exceptional nematode resistance (Rf <0.5 for Meloidogyne incognita) and salt tolerance (leaf Cl⁻ <0.25% at ECw 2.0 dS/m) supporting table grape production in the Coachella Valley, California, and wine grape production on nematode-infested sandy soils of the Barossa Valley, Australia. Dog Ridge (V. champinii selection) contributes nematode resistance genes to breeding programs at UC Davis, serving as the resistance donor for Freedom and Harmony. Freedom (1613C × Dog Ridge) combines V. champinii nematode resistance with improved cold hardiness and graft compatibility, widely used on nematode-infested sites in California's Central Coast and San Joaquin Valley.
A parentage-based rootstock role that inheres in a grafted rootstock by virtue of its genetic derivation from Vitis champinii Planchon, a natural hybrid species (V. mustangensis × V. rupestris) native to the limestone and sandy soils of central and southern Texas. V. champinii contributes characteristic traits that are uniquely valuable for challenging viticultural environments, including exceptional nematode resistance (particularly to root-knot nematodes Meloidogyne spp. and dagger nematodes Xiphinema spp.), high salt tolerance through efficient chloride and sodium exclusion, heat tolerance (thriving at soil temperatures exceeding 35°C), drought tolerance, and high vigor with extensive root systems. V. champinii shows moderate phylloxera resistance (lower than V. riparia or V. berlandieri), moderate lime tolerance (14-17% active lime), and poor cold hardiness limiting its use to warm climates. The species is represented commercially by Dog Ridge and Ramsey (Salt Creek) selections, and contributes to important hybrid rootstocks including Freedom (1613C × Dog Ridge) and Harmony (1613C × Dog Ridge).
Vitis_Champinii_Derived_Rootstock_Role captures the unique contribution of a species adapted to the hot, sandy, often saline soils of south-central Texas where nematode pressure is severe. V. champinii is the primary genetic source of broad-spectrum nematode resistance in grapevine rootstocks, a trait largely absent in V. riparia, V. rupestris, and V. berlandieri. The species' hybrid origin (V. mustangensis × V. rupestris) contributes to its complex trait combination. Climate change, water quality degradation, and soil fumigation phase-outs are increasing demand for champinii-derived rootstocks beyond traditional warm-climate applications.
Champin grape derived rootstock
Champinii-Unterlage
V. champinii derived rootstock
V.-champinii-stämmige Unterlage
champinii rootstock
von Vitis champinii abstammende Unterlage
Heinitz CC et al. (2019) Crop wild relatives of grape, in North American Crop Wild Relatives Vol 2, Springer pp.: 329-351 Walker MA et al. (2014) Rootstock breeding for resistance to nematodes and phylloxera, Acta Horticulturae 1045: 89-97 Cousins P, Walker MA (2002) Genetics of resistance to Meloidogyne incognita in crosses of grape rootstocks, Theoretical and Applied Genetics 105: 802-807 McKenry MV, Anwar SA (2006) Nematode and grape rootstock interactions, Journal of Nematology 38: 312-318
Vitis Champinii Derived Rootstock Role
Börner (V. riparia Geisenheim 183 × V. cinerea Arnold) demonstrates exceptional phylloxera resistance approaching immunity, with essentially no nodosity or tuberosity formation even under heavy infestation pressure, used in German viticulture for replanting phylloxera-devastated sites and in experimental high-value plantings where absolute phylloxera protection is prioritized. The V. cinerea parent contributes the extreme resistance level that exceeds typical V. riparia or V. rupestris resistance. Experimental V. cinerea hybrids at INRAE Colmar show promise for combining phylloxera immunity with improved Xiphinema index resistance for GFLV management in replant situations.
A parentage-based rootstock role that inheres in a grafted rootstock by virtue of its genetic derivation from Vitis cinerea Engelmann (sensu stricto), the sweet winter grape or graybark grape native to the floodplains and bottomlands of the central and southeastern United States from Illinois to Texas and east to Florida. V. cinerea contributes characteristic traits including very strong phylloxera resistance (approaching immunity in some accessions), moderate-to-high vigor, good waterlogging tolerance reflecting adaptation to periodically flooded bottomland habitats, and moderate lime tolerance. V. cinerea shows poor drought tolerance (requiring adequate moisture), intermediate cold hardiness, and difficult rooting from cuttings. The species has been less extensively utilized in commercial rootstock breeding than V. riparia, V. rupestris, or V. berlandieri, but contributes to important specialized rootstocks including Börner (V. riparia × V. cinerea) selected for exceptional phylloxera resistance, and is being incorporated into new rootstock breeding programs for its unique pest resistance profile.
Vitis_Cinerea_Derived_Rootstock_Role captures a relatively underutilized genetic resource that offers exceptional phylloxera resistance exceeding that of the more commonly used American species. V. cinerea var. helleri is now recognized as synonymous with V. berlandieri, so the V. cinerea designation in rootstock breeding refers specifically to V. cinerea sensu stricto (graybark grape). The species' adaptation to bottomland flooding contributes waterlogging tolerance absent in the upland-adapted V. berlandieri and V. rupestris. Breeding programs are increasingly exploring V. cinerea germplasm for resistance stacking and climate adaptation.
Cinerea-Unterlage
V. cinerea derived rootstock
V.-cinerea-stämmige Unterlage
cinerea rootstock
graybark grape derived rootstock
sweet winter grape derived rootstock
von Vitis cinerea abstammende Unterlage
Heinitz CC et al. (2019) Crop wild relatives of grape, in North American Crop Wild Relatives Vol 2, Springer pp.: 329-351 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Schmid J et al. (2009) Geisenheimer Rebsorten und Klone, Geisenheimer Berichte 67 Töpfer R et al. (2011) New horizons for grapevine breeding, Fruit, Vegetable and Cereal Science and Biotechnology 5: 79-100
Vitis Cinerea Derived Rootstock Role
1616 Couderc (V. solonis × V. riparia, where V. solonis may include V. longii germplasm) shows adaptation to sandy, seasonally wet soils in southwestern France, with the solonis/longii contribution providing tolerance of heavy, poorly drained substrates not suited to V. berlandieri × V. rupestris rootstocks. Historical rootstock selections from T.V. Munson's breeding program in Denison, Texas incorporated V. longii germplasm for adaptation to the sandy prairie soils of the southern Great Plains. Experimental V. longii hybrids have been evaluated at Texas A&M for adaptation to the challenging soil conditions of the Texas High Plains viticultural region.
A parentage-based rootstock role that inheres in a grafted rootstock by virtue of its genetic derivation from Vitis longii Prince (synonym V. solonis var. longii), a species or variety native to the sandy prairies and river bottomlands of Texas, Oklahoma, and adjacent states. V. longii contributes characteristic traits including moderate nematode tolerance, adaptation to sandy soils, moderate salt tolerance, and vigorous growth with extensive root systems. The species shows moderate phylloxera resistance (lower than V. riparia or V. berlandieri), poor lime tolerance (restricted to neutral or acidic soils), and limited cold hardiness. V. longii has been sparingly used in rootstock breeding, contributing primarily to some early American rootstock selections and experimental hybrids. Its taxonomic status is debated, with some authorities treating it as a variety of V. solonis (V. solonis var. longii) rather than a distinct species, reflecting morphological intergradation in natural populations.
Vitis_Longii_Derived_Rootstock_Role represents a minor contributor to commercial rootstock germplasm, included for completeness of the parentage classification. The species' adaptation to sandy prairies and moderate nematode tolerance offer traits potentially valuable for specialized applications, but V. longii has been largely overshadowed by V. champinii (superior nematode resistance) and V. berlandieri (superior lime tolerance) in breeding programs. Taxonomic uncertainty regarding V. longii versus V. solonis reflects the complex hybridization and introgression among wild Vitis species in the Texas-Oklahoma region. Germplasm preservation efforts are important as wild populations face habitat loss.
Long grape derived rootstock
Longii-Unterlage
V. longii derived rootstock
V.-longii-stämmige Unterlage
longii rootstock
von Vitis longii abstammende Unterlage
Heinitz CC et al. (2019) Crop wild relatives of grape, in North American Crop Wild Relatives Vol 2, Springer pp.: 329-351 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Munson TV (1909) Foundations of American Grape Culture, Orange Judd Company, New York Moore MO (1991) Classification and systematics of eastern North American Vitis, Sida 14: 339-367
Vitis Longii Derived Rootstock Role
Riparia Gloire de Montpellier (pure V. riparia selection) confers exceptional cold hardiness and low vigor to Pinot Noir in Burgundy's Côte d'Or and Oregon's Willamette Valley, but is restricted to low-lime acidic soils (active lime <9%). SO4 (V. berlandieri × V. riparia) inherits phylloxera resistance and moderate vigor from V. riparia while gaining lime tolerance from V. berlandieri, making it the dominant rootstock in German viticulture. 3309 Couderc (V. riparia × V. rupestris) combines V. riparia's cold hardiness and low vigor with improved rooting from V. rupestris, widely used in Burgundy and Champagne on low-lime sites.
A parentage-based rootstock role that inheres in a grafted rootstock by virtue of its genetic derivation from Vitis riparia Michaux, the riverbank grape native to the northeastern and north-central regions of North America from Quebec to Montana and south to Texas. V. riparia contributes characteristic traits including excellent cold hardiness (surviving midwinter temperatures to -35°C or below), strong phylloxera resistance through rapid wound periderm formation, low-to-moderate vigor with shallow but dense root systems, early growth cessation promoting timely autumn acclimation, efficient magnesium uptake, and restricted potassium translocation beneficial for must acidity. V. riparia-derived rootstocks are adapted to acidic, low-lime soils (typically tolerating only 6-11% active lime) and show poor drought tolerance due to shallow rooting. The species was among the first American vines introduced to Europe following the phylloxera crisis, with Riparia Gloire de Montpellier (selected 1880s) remaining in commercial use. V. riparia is a parent of numerous important rootstock crosses including the V. berlandieri × V. riparia group (SO4, 5BB, 5C, 125AA, 420A) and V. riparia × V. rupestris group (3309C, 101-14Mgt, Schwarzmann).
Vitis_Riparia_Derived_Rootstock_Role captures the genetic contribution of the most cold-hardy and earliest-maturing American Vitis species. V. riparia's natural range extending to 50°N latitude and its adaptation to short growing seasons make it essential for continental climate viticulture. The trade-off for cold hardiness is poor lime tolerance and drought sensitivity, addressed by crossing with V. berlandieri (lime tolerance) or V. rupestris (improved rooting depth). The V. riparia genome has been sequenced and serves as a reference for grapevine genomics research, facilitating marker-assisted selection for riparia-derived traits.
Riparia-Unterlage
V. riparia derived rootstock
V.-riparia-stämmige Unterlage
riparia rootstock
riverbank grape derived rootstock
von Vitis riparia abstammende Unterlage
Heinitz CC et al. (2019) Crop wild relatives of grape, in North American Crop Wild Relatives Vol 2, Springer pp.: 329-351 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22 Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10: 201-235
Vitis Riparia Derived Rootstock Role
St. George (Rupestris du Lot, pure V. rupestris selection) confers very high vigor and deep rooting to Cabernet Sauvignon in Napa Valley, with root penetration exceeding 4 m on deep alluvial soils, but is limited to low-lime sites (active lime <14%). 110 Richter (V. berlandieri × V. rupestris) inherits deep rooting and drought tolerance from V. rupestris while gaining lime tolerance from V. berlandieri, dominating Mediterranean viticulture. 3309 Couderc (V. riparia × V. rupestris) combines V. rupestris's improved rooting capacity with V. riparia's cold hardiness, showing rooting rates of 85-90% in nursery propagation compared to 60-70% for pure V. riparia selections.
A parentage-based rootstock role that inheres in a grafted rootstock by virtue of its genetic derivation from Vitis rupestris Scheele, the sand grape or rock grape native to the gravelly streambeds and rocky outcrops of the Ozark Plateau region of Missouri, Arkansas, Oklahoma, and Texas. V. rupestris contributes characteristic traits including strong phylloxera resistance, deep penetrating root systems (often exceeding 3-4 m depth on permeable substrates), good drought tolerance through hydraulic lift and deep water access, moderate-to-high vigor, easy propagation from hardwood cuttings (rooting rates typically 80-95%), and moderate lime tolerance (10-14% active lime). V. rupestris shows intermediate cold hardiness between V. riparia (superior) and V. berlandieri (inferior). The species contributed to early European rootstock selections including St. George (Rupestris du Lot), and is a parent of major rootstock crosses including V. berlandieri × V. rupestris (110R, 99R, 1103P, 140Ru) and V. riparia × V. rupestris (3309C, 101-14Mgt, Schwarzmann).
Vitis_Rupestris_Derived_Rootstock_Role captures the contribution of a species adapted to the rocky, well-drained, drought-prone habitats of the Ozark region. V. rupestris's deep rooting habit and tolerance of gravelly substrates made it invaluable for drought adaptation in Mediterranean climates when crossed with the lime-tolerant V. berlandieri. The species is now rare in the wild due to habitat loss, making germplasm conservation a priority. Pure V. rupestris rootstocks (St. George) were historically important but have been largely replaced by interspecific hybrids combining rupestris traits with lime tolerance or cold hardiness from other species.
Rupestris-Unterlage
V. rupestris derived rootstock
V.-rupestris-stämmige Unterlage
rock grape derived rootstock
rupestris rootstock
sand grape derived rootstock
von Vitis rupestris abstammende Unterlage
Heinitz CC et al. (2019) Crop wild relatives of grape, in North American Crop Wild Relatives Vol 2, Springer pp.: 329-351 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Ollat N et al. (2016) Grapevine rootstocks: origins and perspectives, Acta Horticulturae 1136: 11-22 Carbonneau A (1985) The early selection of grapevine rootstocks for resistance to drought conditions, American Journal of Enology and Viticulture 36: 195-198
Vitis Rupestris Derived Rootstock Role
1616 Couderc (V. solonis × V. riparia) was historically planted on heavy clay soils of the Bordeaux region where its V. solonis parentage confers tolerance of poorly drained, waterlogged conditions, though poor phylloxera resistance has limited its persistence. Solonis × Othello 1613 (complex hybrid including V. solonis) contributed to the development of Freedom and Harmony rootstocks through crossing with Dog Ridge, with the solonis background potentially contributing to wet-soil adaptation. Historical French rootstock trials documented V. solonis selections for the argilo-calcaire (clay-limestone) soils of the southern Charentes where pure V. berlandieri × V. rupestris crosses showed poor establishment on heavy clays.
A parentage-based rootstock role that inheres in a grafted rootstock by virtue of its genetic derivation from Vitis solonis Planchon ex Hance (including V. solonis var. solonis and potentially V. solonis var. longii), a species native to the creek banks, river bottoms, and low prairies of Texas and adjacent states. V. solonis contributes characteristic traits including tolerance of heavy, poorly drained clay soils, moderate waterlogging tolerance reflecting bottomland habitat adaptation, moderate vigor, and adaptation to neutral-to-alkaline substrates. The species shows poor phylloxera resistance (among the weakest of American Vitis species), poor drought tolerance (requiring adequate moisture), limited cold hardiness, and moderate-to-difficult rooting from cuttings. V. solonis was utilized in early French rootstock breeding, contributing to the 1616 Couderc and Solonis × Riparia 1616 selections designed for heavy clay soils, but has been largely superseded by more phylloxera-resistant species combinations in modern breeding.
Vitis_Solonis_Derived_Rootstock_Role represents a historically significant but now minor contributor to rootstock germplasm, largely superseded by species with superior phylloxera resistance. V. solonis's value lies primarily in adaptation to heavy, wet clay soils—a niche not well served by the dominant V. berlandieri × V. riparia or V. berlandieri × V. rupestris hybrid groups adapted to well-drained substrates. The species' weak phylloxera resistance means solonis-derived rootstocks are viable only in regions with low phylloxera pressure or in combination with more resistant species. Taxonomic relationships between V. solonis, V. longii, and V. acerifolia remain debated among systematists.
Solonis grape derived rootstock
Solonis-Unterlage
V. solonis derived rootstock
V.-solonis-stämmige Unterlage
solonis rootstock
von Vitis solonis abstammende Unterlage
Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Heinitz CC et al. (2019) Crop wild relatives of grape, in North American Crop Wild Relatives Vol 2, Springer pp.: 329-351 Pongrácz DP (1983) Rootstocks for Grape-vines, David Philip Publishers, Cape Town Viala P, Ravaz L (1903) American Vines: Their Adaptation, Culture, Grafting and Propagation, Coulet et Fils, Montpellier
Vitis Solonis Derived Rootstock Role
A mature Vitis vinifera cv. Pinot Noir vine trained in Guyot pruning in the Côte de Nuits, Burgundy, representing the complete organism whose morphological structures (root system, trunk, cordons, canes, shoots, leaves, clusters), phenological cycle (dormancy through harvest and senescence), measurable qualities (vigor, fertility, sugar content), dispositions (cold hardiness, disease resistance), roles (rootstock, scion), and associated information entities (cultivar names, BBCH codes, ampelographic descriptors) constitute the domain of the VVO.
Vitis_vinifera is a material entity (BFO_0000040 (https://www.ebi.ac.uk/ols4/ontologies/bfo/classes/http%253A%252F%252Fpurl.obolibrary.org%252Fobo%252FBFO_0000040)) representing the plant species Vitis vinifera L. (Vitaceae), the common grapevine, a perennial woody liana originating from the Near East and Mediterranean basin, domesticated approximately 6,000-8,000 years ago, and now the most widely cultivated fruit crop globally for wine, table grape, and raisin production. As the singular root class of the Vitis Vinifera Ontology (VVO), this entity serves as the taxonomic and ontological anchor from which all morphological structures, phenological processes, qualities, dispositions, roles, and information entities of the grapevine are organized. Vitis vinifera is distinguished from other Vitis species by its superior enological properties, high sugar accumulation capacity, complex aromatic compound synthesis, and extensive clonal diversity encompassing over 10,000 recognized cultivars worldwide.
Vitis_vinifera is the root class of VVO and the only class aligned directly with BFO_0000001 (entity). All six top-level categories of VVO are immediate subclasses of Vitis_vinifera, reflecting the ontological commitment that every class in VVO describes an aspect of this single species. VVO does not model other Vitis species (V. labrusca, V. riparia, V. rupestris, etc.) except insofar as they serve as rootstock partners in grafted vine systems, captured under Grapevine_Role. The taxonomic placement follows the APG IV classification: Order Vitales, Family Vitaceae, Genus Vitis, Subgenus Vitis, Species V. vinifera. The binomial authority is Linnaeus (1753) Species Plantarum. Lineage (full) is defined in NCBI Taxonomy Browser, taxonomy ID 29760 (https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?command=show&mode=node&id=29760&lvl=).
Edle Weinrebe
European grapevine
Europäische Weinrebe
Kulturrebe
Vitis vinifera L.
Weinstock
common grapevine
cultivated grapevine
wine grape
This OJ, Park Y-J, et al. (2006) Genetic characterization of grapevines collected from the southwestern Korean Peninsula via SSR markers, Vitis 45(3): 137-143 McGovern PE (2003) Ancient Wine: The Search for the Origins of Viniculture, Princeton University Press pp.: 1-45 Mullins MG, Bouquet A, Williams LE (1992) Biology of the Grapevine, Cambridge University Press pp.: 1-16 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 1-30 OIV (2017) Focus OIV: Distribution of the World's Grapevine Varieties, International Organisation of Vine and Wine
Vitis vinifera
The VIVC assigns Riesling Weiss (VIVC number 10077) as the prime name for the cultivar known in Australia as Rhine Riesling, in Italy as Riesling Renano, in Hungary as Rajnai Rizling, and in South Africa as Weisser Riesling, with the VIVC database linking 147 documented synonym entries to this single prime name and providing the unique VIVC variety number that serves as a persistent digital identifier enabling unambiguous cross-referencing of the cultivar across all partner databases including EURISCO, Genesys, and national germplasm information systems.
A prime name that is the accepted designation for a grapevine cultivar in the Vitis International Variety Catalogue (VIVC), the most comprehensive global database of grapevine cultivar nomenclature maintained by the Julius Kühn-Institut, serving as the de facto international scientific standard for cultivar name resolution and synonym cross-referencing.
VIVC_Prime_Name represents the scientifically authoritative cultivar naming standard, maintained at the Julius Kühn-Institut in Siebeldingen, Germany since 1986, documenting over 24,000 cultivar designations linked to approximately 6,000-8,000 genetically distinct accessions. The VIVC prime name is selected based on historical priority, international usage frequency, and taxonomic clarity, and may differ from both the OIV registered name and national official prime names: for example, VIVC uses Riesling Weiss (specifying berry color) while the OIV registers simply Riesling, and the German Bundessortenamt lists Weisser Riesling. The VIVC variety number provides a language-independent persistent identifier that resolves nomenclatural ambiguity more reliably than any name-based system.
VIVC accepted name
VIVC catalogue designation
VIVC variety name
VIVC-Katalogname
VIVC-Leitname
VIVC-Sortenbezeichnung
Maul E and Töpfer R (2015) Vitis International Variety Catalogue (VIVC): a cultivar database, Vitis 54: 155-157 VIVC (2024) https://www.vivc.de OIV (2013) OIV List of Vine Varieties and Their Synonyms
VIVC Prime Name
The VMC marker series contributed approximately 200 characterized microsatellite loci distributed across all 19 Vitis chromosomes, developed through an industry-academia consortium that pooled resources for large-scale SSR isolation and characterization — VMC markers such as VMC1b11, VMC4f3.1, VMCNG4b9, and VMC7f2 were incorporated into the high-density genetic maps (Doligez et al. 2006, Adam-Blondon et al. 2004) that provided the marker framework for the first comprehensive QTL studies of berry quality traits in grapevine. The European Vitis Database extended cultivar identification panel includes 11 VMC loci in addition to the 9 OIV standard markers, providing a 20-marker panel with cumulative probability of identity below 10⁻¹⁵ — sufficient to distinguish not only cultivars but also first-degree relatives and, in some cases, clones within a cultivar that share identical 9-locus standard profiles.
A microsatellite marker data record for the VMC (Vitis Microsatellite Consortium) series of SSR markers developed through an international collaborative project coordinated by Agrogene S.A. (Moissy-Cramayel, France), comprising the largest single collection of characterized Vitis microsatellite markers with over 200 loci developed through high-throughput genomic library screening, providing dense genome coverage used extensively in genetic mapping, QTL analysis, and extended cultivar identification panels requiring higher discrimination power than the standard 9-locus set.
VMC_Marker_Series documents the most extensive coordinated microsatellite development effort in grapevine genetics — a consortium-based approach that pooled the resources of multiple European research institutions and the private company Agrogene to generate a comprehensive SSR marker toolkit at a scale beyond the capacity of individual research groups. The VMC consortium model (active from approximately 1997 to 2005) addressed the practical limitation that SSR development through genomic library construction, colony screening, sequencing, primer design, and polymorphism testing is labor-intensive and expensive per marker: by centralizing the high-throughput steps and distributing characterized markers to consortium members, the VMC project achieved economies of scale that produced 10-fold more markers than any individual laboratory effort. The resulting marker density — approximately 10 characterized SSRs per chromosome on average — provided the foundation for saturated genetic mapping and the fine-scale QTL analyses that linked specific genomic regions to economically important traits including muscat aroma (chromosome 5), berry color (chromosome 2), seedlessness (chromosome 18), and disease resistance loci. While SNP markers have largely replaced SSRs for genetic mapping and QTL analysis since the 2010s, VMC markers remain in active use for cultivar identification in extended panels and for targeted genotyping applications where a small number of highly informative codominant markers is more practical than array-based SNP genotyping.
Agrogene SSR series
Agrogene-SSR-Serie
VMC SSR loci
VMC-Marker-Serie
Vitis Microsatellite Consortium markers
Vitis-Mikrosatelliten-Konsortium-Marker
Doligez A et al. (2006) New stable QTLs for berry weight do not colocalize with QTLs for seed traits in cultivated grapevine (Vitis vinifera L.), BMC Plant Biology 6: 12 Adam-Blondon AF et al. (2004) Genetic mapping of grapevine, Functional and Integrative Genomics 4: 196-211 Sefc KM et al. (2001) Microsatellite markers for grapevine, in Roubelakis-Angelakis KA (ed) Molecular Biology and Biotechnology of the Grapevine, Kluwer pp: 433-463
VMC Marker Series
Volatile acidity for Chardonnay from Meursault (2023 vintage) measures 0.38 g/L as acetic acid, within the normal range for barrel-fermented white Burgundy (0.25-0.50 g/L), with the moderate VA level reflecting controlled oxidative exposure during 14 months of barrel aging without indicating any spoilage activity. Volatile acidity for Barolo DOCG (2020 vintage) records 0.62 g/L after 36 months of extended oak aging, approaching but not exceeding the EU regulatory maximum of 1.20 g/L for red wines, with the elevated VA consistent with the extended oxidative maturation in large Slavonian botti that characterizes traditional Barolo production — a level perceived by many tasters as contributing positive aromatic lift rather than vinegar-like defect. Volatile acidity for Zweigelt from Neusiedlersee (2024 vintage, early-drinking style) documents 0.28 g/L, well below any spoilage concern and consistent with clean reductive winemaking in stainless steel.
Volatile acidity is a wine analysis data entity that records the concentration of steam-distillable organic acids in finished wine, predominantly acetic acid with minor contributions of formic, butyric, and propionic acids, determined by steam distillation followed by titration with sodium hydroxide (OIV reference method) or by enzymatic assay specific to acetic acid, expressed in grams per liter as acetic acid equivalents (g/L), and serving as the primary indicator of wine microbial health and spoilage status, as elevated volatile acidity signals uncontrolled oxidative bacterial metabolism by Acetobacter or excessive production by aberrant yeast or lactic acid bacteria, while trace levels contribute positively to aromatic complexity and flavor lift.
Volatile acidity occupies a unique position as both a wine quality indicator and a potential wine fault, with the boundary between positive contribution and spoilage defect determined by concentration, wine style, and cultural context. At low levels (0.2-0.5 g/L), volatile acidity contributes to the perception of aromatic lift, complexity, and vinosity — the ethyl acetate formed from acetic acid and ethanol at concentrations below 80-100 mg/L adds a pleasant fruity complexity, while acetic acid itself enhances the perception of fruit freshness. Above approximately 0.7-0.8 g/L (depending on wine style and individual threshold variation), volatile acidity becomes perceptible as a distinct vinegar-like sharpness that most tasters consider a defect, with the perception threshold further lowered by the synergistic effect of ethyl acetate at concentrations above 120-150 mg/L. The EU regulatory limits for volatile acidity are 1.08 g/L for white and rosé wines and 1.20 g/L for red wines (expressed as acetic acid), with higher limits permitted for wines with extended aging (1.50 g/L for some aged wines) and for certain sweet wine categories where botrytis-related VA is inherent to the production process. The sources of volatile acidity in wine include normal yeast fermentation (0.15-0.40 g/L produced by healthy Saccharomyces cerevisiae), malolactic fermentation (0.05-0.15 g/L additional from Oenococcus oeni), and spoilage organisms (Acetobacter species oxidizing ethanol to acetic acid under aerobic conditions, heterolactic bacteria producing acetic acid from sugar metabolism). The OIV steam distillation method captures all volatile acids and requires correction for free and bound sulfur dioxide (which co-distills), while the enzymatic assay using acetyl-CoA synthetase provides specific acetic acid quantification without sulfur dioxide interference, making it the preferred screening method for routine quality control.
Essiggehalt
Essigsäure
VA-Wert
acetic acid content
flüchtige Säure
flüchtige Säure des Weines
steam-distillable acidity
volatile acid
wine VA
OIV (2021) Compendium of International Methods of Analysis, Method OIV-MA-AS313-02 (volatile acidity) Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2, 2nd Ed, Wiley, pp.: 12-22 Waterhouse AL et al. (2016) Understanding Wine Chemistry, Wiley, pp.: 80-95 du Toit WJ, Lambrechts MG (2002) The enumeration and identification of acetic acid bacteria from South African red wine fermentations. International Journal of Food Microbiology 74(1-2): 57-64
Volatile Acidity
Volatile acidity aroma diagnosis in suspect Zinfandel barrel identifies vinegar-like character (intensity 5.5 on 10-point fault scale) with underlying nail polish notes (3.8), with analytical confirmation showing acetic acid at 1.1 g/L and ethyl acetate at 220 mg/L, both exceeding perception thresholds and approaching legal limits (typically 1.2-1.4 g/L depending on jurisdiction), with the barrel traced to incomplete filling allowing headspace oxygen that enabled Acetobacter growth on the wine surface, demonstrating the oxygen-dependent etiology and the importance of topping for VA prevention.
Volatile acidity aroma quality is an off-aroma quality that represents olfactory perception of acetic acid and ethyl acetate arising from aerobic microbial metabolism, characterizing the vinegar-like, nail polish remover, or solvent aromatic impression that indicates spoilage by acetic acid bacteria or certain aerobic yeast species. Within the BFO framework, this quality specifically addresses the volatile acidity fault category, distinguishing acetic-derived off-aromas from other fault types while recognizing that low levels of volatile acidity contribute to wine aromatic lift and complexity, with the fault designation applying only when concentrations exceed acceptable thresholds. The ontological significance lies in volatile acidity representing one of the most recognized and universally rejected wine faults, where vinegar character indicates fundamental spoilage that renders wine commercially unacceptable at elevated levels, with volatile acidity measurement (expressed as acetic acid equivalents) serving as a standard quality control parameter with legal maximum limits in most wine regions. The compositional basis for volatile acidity aroma includes acetic acid (providing vinegar smell, perception threshold approximately 0.7-0.8 g/L in wine), ethyl acetate (contributing nail polish, solvent character, threshold approximately 150-180 mg/L), and related volatile acids and esters formed through acetic acid bacteria metabolism of ethanol under aerobic conditions. The microbial origins of volatile acidity include Acetobacter and Gluconobacter species (acetic acid bacteria requiring oxygen), certain film-forming yeasts (Pichia, Candida species), and Brettanomyces under specific conditions, with contamination risk highest during pre-fermentation handling, sluggish fermentation, and post-fermentation storage where oxygen exposure enables aerobic metabolism. The remediation options for volatile acidity are limited, with reverse osmosis and ion exchange capable of reducing acetic acid but expensive and potentially stripping other wine components, making prevention through oxygen exclusion and microbial control more practical than correction. Understanding volatile acidity aroma quality enables fault identification from acetic-derived character, spoilage risk assessment based on vulnerable process stages, and quality control through volatile acidity monitoring and threshold enforcement.
Volatile_Acidity_Aroma_Quality addresses the acetic acid and ethyl acetate fault category, representing one of the most universally recognized wine defects. This quality has established analytical thresholds (VA expressed as g/L acetic acid) that serve regulatory and quality control functions. The class acknowledges that low-level volatile acidity contributes positively to aromatic lift, with fault designation applying only at elevated concentrations where vinegar character becomes apparent.
Acescence
Essigstich
VA fault
VA-Fehler
acetic character
acetification
vinegar aroma
Bartowsky EJ, Henschke PA (2008) Acetic acid bacteria spoilage of bottled red wine — a review. International Journal of Food Microbiology 125(1): 60-70 Drysdale GS, Fleet GH (1988) Acetic acid bacteria in winemaking: a review. American Journal of Enology and Viticulture 39(2): 143-154
Volatile Acidity Aroma Quality
Volatile compound profile for Pinot Noir from Volnay, Burgundy (2024 vintage, post-MLF) identifies 185 individual compounds by GC-MS full scan, with major contributors including ethyl acetate (62 mg/L), isoamyl alcohol (185 mg/L), 2-phenylethanol (42 mg/L, rose aroma), isoamyl acetate (3.8 mg/L, banana), ethyl hexanoate (0.85 mg/L, apple-pear), ethyl octanoate (0.65 mg/L, tropical fruit), diacetyl (2.2 mg/L from MLF, butter), and linalool (18 µg/L, floral, sub-threshold), with the overall profile characterized by a dominance of fermentation esters modulated by varietal β-damascenone (5.5 µg/L) and subtle whole-cluster derived eugenol (8 µg/L, spice). Volatile compound profile for Grüner Veltliner from Wachau Smaragd (2024 vintage) documents a clean fermentative base of isoamyl acetate (4.2 mg/L), ethyl butyrate (0.45 mg/L), 2-phenylethanol (28 mg/L), overlaid with cultivar-specific rotundone (below the 16 ng/L threshold but contributing to the background spiciness), 3MH (380 ng/L, grapefruit), and trace linalool (22 µg/L).
Volatile compound profile is an aromatic analysis data entity that records the comprehensive inventory of all identified volatile and semi-volatile organic compounds in grape must or wine beyond the specific terpene, norisoprenoid, methoxypyrazine, and thiol classes, determined by GC-MS full-scan analysis following headspace, SPME, or liquid-liquid extraction, typically encompassing higher alcohols (fusel alcohols), ethyl esters, acetate esters, fatty acids, aldehydes, ketones, lactones, volatile phenols, and sulfur compounds, expressed in milligrams per liter (mg/L) for major compounds and micrograms per liter (µg/L) for minor components, and providing the holistic chemical characterization of the complete volatile matrix that creates the composite aroma perception of a wine.
Volatile compound profile data provides the comprehensive chemical context within which individual aroma compound classes (terpenes, norisoprenoids, methoxypyrazines, thiols) exert their sensory effects, as wine aroma is perceived as a gestalt arising from the interaction of hundreds of volatile compounds rather than from any single chemical species. The approximately 600-1000 volatile compounds identified in wine to date arise from three sources — grape-derived (varietal aroma, representing cultivar identity), fermentation-derived (produced by Saccharomyces cerevisiae and malolactic bacteria, representing winemaking process), and aging-derived (formed through chemical reactions during maturation, representing wine evolution) — with the relative contribution of each source shifting as wine develops. Higher alcohols (predominantly isoamyl alcohol, isobutanol, 2-phenylethanol, produced through the Ehrlich pathway from amino acid catabolism) typically constitute the largest volatile fraction by mass (100-400 mg/L total), while esters (fruity character from ethyl esters of fatty acids and acetate esters of higher alcohols) contribute most to fruity aroma despite lower concentrations (1-50 mg/L total). The interaction effects between volatile compounds — synergy, suppression, and additive perception — mean that the sensory impact of a volatile profile cannot be predicted by simple summation of individual compound odor activities. Comprehensive volatile profiling has applications beyond aroma characterization, including authenticity verification (volatile fingerprinting for origin and vintage authentication), fault detection (specific marker compounds for Brettanomyces contamination, oxidation, reduction), and process optimization (monitoring fermentation-derived volatile production kinetics). The emergence of electronic nose (e-nose) technology and comprehensive GC×GC-TOF-MS platforms is progressively enabling rapid, high-resolution volatile profiling that was previously limited to specialized research laboratories.
Aromaprofil
Gesamtvolatilprofil
VOC profile
Volatilprofil
aroma volatile inventory
flüchtige Substanzen Profil
flüchtiges Verbindungsprofil
volatile composition data
volatile fingerprint
volatile profile
Robinson AL et al. (2014) Origins of grape and wine aroma. Part 2. Chemical and sensory analysis. American Journal of Enology and Viticulture 65(1): 25-42 Ferreira V et al. (2000) Quantitative determination of the odorants of young red wines from different grape varieties. Journal of the Science of Food and Agriculture 80(11): 1659-1667 Ebeler SE, Thorngate JH (2009) Wine chemistry and flavor. Journal of Agricultural and Food Chemistry 57(18): 8098-8108 Hazelwood LA et al. (2008) The Ehrlich pathway for fusel alcohol production. Applied and Environmental Microbiology 74(8): 2259-2266
Volatile Compound Profile
In Scheurebe berries in Rheinhessen, Germany during BBCH 83-89, volatile compound synthesis generates free monoterpenes increasing from 0.2 to 1.4 mg/L over 45 days, with linalool synthase activity reaching 28 nmol/min/g fresh weight producing free linalool of 0.55 mg/L, concurrent norisoprenoid synthesis generating free beta-damascenone of 3.2 µg/L and beta-ionone of 1.8 µg/L, while C6 aldehyde content declines from 95 to 28 µg/L hexanal as green character diminishes with advancing ripeness under cool continental conditions averaging 18°C.
Volatile compound synthesis encompasses the enzymatic biosynthesis of free, aromatic volatile compounds in Vitis vinifera berries during ripening, wherein primary and secondary metabolic pathways generate the diverse array of low-molecular-weight organic molecules possessing sufficient vapor pressure for olfactory detection, including monoterpenes, sesquiterpenes, C13-norisoprenoids, volatile thiols, esters, aldehydes, and alcohols that collectively constitute the immediately perceptible aromatic character of grape berries prior to any precursor hydrolysis or fermentation-derived modifications. This process involves multiple biosynthetic routes operating in distinct cellular compartments, with the methylerythritol phosphate pathway generating geranyl diphosphate precursor in plastids followed by terpene synthase conversion to monoterpene hydrocarbons and alcohols in cytosolic compartments, carotenoid cleavage dioxygenases generating norisoprenoid volatiles through oxidative degradation of carotenoid substrates in chromoplasts, and fatty acid metabolism producing C6 aldehydes and alcohols through lipoxygenase and hydroperoxide lyase sequential activities in various cellular locations. The temporal regulation of volatile compound synthesis shows compound-class-specific patterns, with monoterpene synthesis increasing progressively throughout post-veraison ripening as terpene synthase expression rises, norisoprenoid generation accelerating during mid-to-late ripening coincident with carotenoid degradation in coloring berries, and C6 volatile synthesis showing complex dynamics reflecting the balance between lipoxygenase pathway activity and aldehyde reduction or oxidation reactions that modify initially formed products. Cultivar genetics fundamentally determine volatile compound synthesis capacity through variety-specific expression of biosynthetic enzyme genes, with aromatic Muscat varieties expressing high levels of linalool synthase and geraniol synthase generating 10-100 fold higher monoterpene concentrations than neutral varieties, and Cabernet family varieties expressing characteristic O-methyltransferases producing methoxypyrazines distinctive to these cultivars. The partitioning between free volatile production and subsequent conjugation to precursor forms represents a critical regulatory point, with the balance between synthesis rates and glycosylation activities determining what fraction of produced volatiles remains in free, immediately aromatic form versus conjugated storage awaiting release during winemaking. Environmental modulation of volatile compound synthesis occurs through temperature effects on enzyme activities and substrate availability, light regulation of terpenoid and carotenoid pathway expression, and stress-responsive induction of secondary metabolism under water deficit or other environmental challenges that activate volatile biosynthetic pathways.
Volatile_Compound_Synthesis is parent class under Aroma_Development representing biosynthesis of free aromatic volatiles. Distinguishes from Aroma_Precursor_Formation which generates conjugated forms. Contains subclasses Free_Volatile_Production, Ester_Formation, and Aldehyde_Synthesis. Multiple pathways including MEP terpenoid, carotenoid cleavage, and fatty acid oxidation. Cultivar-specific enzyme expression determines synthesis capacity.
Volatil-Verbindungs-Synthese
aromatic volatile production
aromatische Volatil-Produktion
free aroma compound formation
free volatile biosynthesis
freie Aroma-Verbindungs-Bildung
freie Volatil-Biosynthese
volatile aroma synthesis
Martin DM et al (2010) Functional annotation of the grapevine terpene synthase gene family. BMC Plant Biology 10: 226 Kalua CM and Boss PK (2009) Evolution of volatile compounds during development of Cabernet Sauvignon grapes. Journal of Agricultural and Food Chemistry 57: 3818-3830 Keller M (2020) The Science of Grapevines 3rd Ed pp 285-295
Volatile Compound Synthesis
Berry volume determination in Pinot Noir destined for premium Burgundy production employs water displacement volumetry on 100-berry samples collected at weekly intervals from véraison through harvest, tracking the characteristic double-sigmoid growth curve from initial volumes of 0.4-0.6 mL per berry at véraison through final volumes of 1.0-1.4 mL at optimal harvest maturity, with smaller final volumes (below 1.0 mL) correlating with higher phenolic concentration and more concentrated wine character.
Volume_Quality represents the category of morphological qualities that quantify three-dimensional spatial occupation of grapevine organs. Volume measurements integrate all three spatial dimensions to characterize the total space enclosed within organ boundaries, providing essential data for understanding storage capacity, dilution effects, and relationships between organ size and physiological function. This category comprises organ-specific subclasses including Berry_Volume_Quality and Cluster_Volume_Quality that address the most agriculturally important volume measurements. Volume is the critical dimension for understanding concentration relationships in grape berries because juice and extractable compounds exist within the three-dimensional berry space. Larger berries dilute skin-derived compounds relative to juice volume, while smaller berries concentrate these compounds, fundamentally affecting wine composition and style. Volume measurements also apply to vegetative organs including trunk and root systems where volume correlates with storage capacity for carbohydrates and nutrients. Canopy volume characterizes the three-dimensional space occupied by foliage, relevant for spray application calibration and light interception modeling. Volume determination methods depend on organ characteristics: liquid displacement for dense, impermeable organs gas pycnometry for precise measurements geometric calculation from linear dimensions assuming standard shapes and three-dimensional scanning for irregular geometries. The relationship between volume and surface area (surface-to-volume ratio) decreases with increasing organ size, with important physiological consequences for exchange processes.
Volume_Quality is positioned parallel to Linear_Dimension_Quality, Area_Quality, and Mass_Quality under Size_Quality. Volume relates to but is distinct from mass, with the relationship mediated by density (mass per unit volume). For berries, volume can be estimated from linear dimensions assuming ellipsoidal geometry (V = 4/3 × π × a × b × c where a, b, c are semi-axes), or measured directly by displacement methods. Tissue-specific volumes (skin volume, pulp volume, seed volume) can be determined through dissection and separate measurement, providing data on berry compartment proportions. PATO_0000918 (volume) provides the upper-ontology reference.
Volumenmaß
Volumenqualität
dreidimensionale Ausdehnungsqualität
organ volume
räumliche Ausdehnung
spatial occupation
three-dimensional extent quality
volumetric quality
Coombe BG (1976) The Development of Fleshy Fruits. Annual Review of Plant Physiology 27(1): 207-228 Matthews MA and Nuzzo V (2007) Berry Size and Yield Paradigms on Grapes and Wines Quality. Acta Horticulturae 754: 423-436
Volume Quality
The VrZAG marker series published by Sefc et al. (1999) comprised approximately 25 microsatellite loci isolated from a Vitis riparia genomic library, of which VrZAG62 (chromosome 7, allele range 183-204 bp in V. vinifera, 15 alleles, PIC 0.80) and VrZAG79 (chromosome 5, allele range 237-261 bp, 18 alleles, PIC 0.83) were selected for the OIV standard panel based on their high polymorphism in V. vinifera despite their origin from an American Vitis species — this cross-species transferability is characteristic of Vitis microsatellites, where conserved flanking regions enable primer pairs designed from one species to amplify orthologous loci in other species within the genus, with the repeat regions themselves often showing even higher polymorphism in V. vinifera than in the source species due to the larger effective population size and greater genetic diversity of the cultivated grapevine gene pool.
A microsatellite marker data record for the VrZAG (Vitis riparia Züchtungsforschung Abteilung Genetik) series of SSR markers developed at the University of Natural Resources and Life Sciences (BOKU), Vienna, by Sefc, Regner, Glössl, and Steinkellner, originally isolated from Vitis riparia genomic libraries but demonstrating excellent cross-species transferability to V. vinifera, with VrZAG62 and VrZAG79 contributing 2 of the 9 markers in the OIV standard cultivar identification panel.
VrZAG_Marker_Series documents the contribution of Austrian grapevine genetics to the international SSR marker toolkit, notable for the deliberate development of markers from an American Vitis species (V. riparia) rather than from V. vinifera itself. This cross-species development strategy exploited the high conservation of SSR flanking sequences within the genus Vitis — primer pairs designed from V. riparia sequences amplify the orthologous loci in V. vinifera with success rates exceeding 80%, while the microsatellite repeat regions often show higher allelic diversity in V. vinifera than in the source species: this cross-species utility makes VrZAG markers particularly valuable for genotyping interspecific hybrids and rootstock cultivars (which contain V. riparia, V. berlandieri, and V. rupestris germplasm) where markers developed from V. vinifera may show reduced polymorphism or amplification failure. The VrZAG series also contributed markers used in the first comprehensive genetic maps of grapevine (Adam-Blondon et al. 2004, Doligez et al. 2006) that provided the framework for QTL mapping of berry quality traits, disease resistance, and phenological timing.
BOKU Vienna microsatellite markers
BOKU-Wien-Mikrosatelliten
Sefc-Steinkellner SSR series
Sefc-Steinkellner-SSR-Serie
VrZAG SSR loci
VrZAG-Marker-Serie
Sefc KM et al. (1999) Identification of microsatellite sequences in Vitis riparia and their applicability for genotyping of different Vitis species, Genome 42: 367-373 This P et al. (2004) Development of a standard set of microsatellite reference alleles, Theoretical and Applied Genetics 109: 1448-1458 Sefc KM et al. (2001) Microsatellite markers for grapevine, in Roubelakis-Angelakis KA (ed) Molecular Biology and Biotechnology of the Grapevine, Kluwer pp: 433-463
VrZAG Marker Series
VSP suitability quality comparison in Willamette Valley evaluated performance of five Pinot Noir clones under standardized vertical shoot positioning management. Dijon clone 115 demonstrated excellent VSP suitability quality with moderate vigor filling 1.2 m canopy height target, naturally upright growth requiring only 2.1 positioning passes, leaf layer number of 1.3 at véraison, and fruit zone PAR of 18% ambient achieving optimal phenolic development. Dijon 777 showed good VSP suitability quality with slightly higher vigor requiring 2.8 positioning passes but comparable canopy metrics. Pommard clone exhibited marginal VSP suitability quality with excessive vigor creating leaf layer number of 2.4 despite 3.5 positioning passes, compromised fruit exposure at 8% PAR, and elevated disease pressure requiring enhanced fungicide programs. Wadenswil clone displayed poor VSP suitability quality with sprawling growth habit and excessive lateral development creating persistent canopy management challenges that motivated evaluation of alternative training approaches for this clone in the regional context.
VSP suitability quality is a trellis adaptation quality that inheres in a grapevine cultivar or clone and represents the degree of compatibility with vertical shoot positioning training systems characterized by upright shoot arrangement within catch wires creating narrow, planar canopy walls with defined fruiting zones at cordon or cane height and vegetative growth trained vertically above, enabling efficient canopy management, optimal light distribution to fruit and renewal zones, and compatibility with mechanical harvesting and precision canopy management equipment. This quality determines whether a cultivar can achieve the organized canopy architecture, appropriate vigor balance, and fruit quality outcomes that justify the infrastructure investment and management intensity associated with VSP systems that dominate modern premium wine grape production worldwide. The ontological significance of VSP suitability quality lies in its role as a primary selection criterion for cultivars intended for contemporary commercial viticulture, wherein VSP has become the predominant training system for quality wine production due to its superior canopy organization, mechanization compatibility, and fruit exposure characteristics compared to traditional unstructured approaches. The biological requirements for high VSP suitability quality include moderately upright natural shoot orientation enabling efficient catch wire placement, appropriate vigor level that fills but does not exceed the canopy dimensions defined by standard VSP trellis configurations, internode lengths compatible with typical catch wire spacing of 20-30 cm, moderate lateral development that contributes to canopy fill without creating excessive density, and cluster architecture suitable for the exposed fruit zone environment created by vertical canopy organization. Cultivars with excessive vigor overwhelm VSP canopy capacity creating management challenges and compromised fruit quality, while insufficient vigor produces sparse canopies failing to achieve adequate light interception and yield potential. The assessment of VSP suitability quality involves evaluation of canopy organization achieved with standardized positioning inputs, quantification of canopy quality metrics including leaf layer number and gap frequency, measurement of fruit zone microclimate conditions, and comparison of fruit composition and wine quality outcomes to alternative training configurations. Regional viticulture has accumulated extensive empirical knowledge of cultivar VSP suitability supporting informed variety selection for VSP-based production systems.
VSP_Suitability_Quality_(Vertical_Shoot_Positioning) is_a Trellis_Adaptation_Quality, representing compatibility with the predominant modern training system for premium wine production. This class addresses the specific requirements of vertical shoot positioned canopies including upright growth habit, moderate vigor, and appropriate internode length. The target canopy metrics of LLN 1.0-1.5, gaps 20-40%, and fruit zone PAR 10-20% provide quantitative benchmarks for suitability assessment. Critical for cultivar selection in regions where VSP dominates commercial production.
Spaliererziehungs-Kompatibilität
Steillagentauglichkeit
VSP compatibility
VSP-Eignung
Vertikalerziehungseignung
upright canopy system fitness
vertical shoot positioning adaptation
vertical training suitability
Smart RE, Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management. Winetitles, Adelaide, pp. 32-43 Reynolds AG et al. (2004) Influence of training system and vine spacing on vine performance of Vitis vinifera L. cv. Riesling. American Journal of Enology and Viticulture 55:79-86 Vanden Heuvel JE et al. (2004) Influence of training system and shoot density on the light microclimate and fruit composition of Vitis vinifera L. cv. Riesling. American Journal of Enology and Viticulture 55:87-97
VSP Suitability Quality (Vertical Shoot Positioning)
Vulnerability to cavitation assessment in Syrah stems reveals the hydraulic safety margins determining drought tolerance limits, with vulnerability curve showing P50 of -1.9 MPa and P88 of -2.8 MPa, while typical midday stem water potentials during moderate deficit irrigation reach -1.4 MPa (operating with 0.5 MPa safety margin to P50), but severe drought conditions can reach -2.2 MPa (exceeding P50, causing >50% conductivity loss), demonstrating how cavitation vulnerability establishes the hydraulic limits that irrigation management must respect to avoid catastrophic transport system damage.
Vulnerability to cavitation quality is a hydraulic conductivity quality that represents the susceptibility of xylem conduits to air entry and embolism formation under water stress, characterizing the relationship between water potential and hydraulic conductivity loss that determines the drought tolerance of the water transport system and the risk of catastrophic hydraulic failure under severe stress conditions. Within the BFO framework, this quality addresses the failure mode dimension of hydraulic architecture, distinguishing vulnerability to dysfunction from baseline conductivity while recognizing that the water potential threshold for embolism, rather than maximum conductivity, often determines the functional limits of the hydraulic system under drought conditions. The ontological significance lies in cavitation vulnerability determining the ultimate hydraulic limitation under drought, where the progressive loss of functional conduits through air entry reduces the conductivity available to supply transpiration, potentially creating a positive feedback where declining conductivity causes further water potential decline that induces more embolism, risking runaway hydraulic failure if critical thresholds are exceeded. The physical basis for cavitation vulnerability involves air entry through pit membranes connecting adjacent vessels, with the pressure difference required to pull air through pit membrane pores determined by membrane porosity (larger pores allowing air entry at less negative pressures), such that the trade-off between pit conductivity and cavitation resistance creates the fundamental efficiency-safety trade-off in xylem design. The quantitative characterization of vulnerability employs vulnerability curves relating percent loss of hydraulic conductivity (PLC) to xylem water potential, with key parameters including P50 (water potential causing 50% conductivity loss), P88 (causing 88% loss, often considered effectively lethal), and the slope of the vulnerability curve (indicating how rapidly conductivity declines around threshold values). The recovery from cavitation through embolism repair is possible in some species and organs, with evidence for both positive pressure-driven refilling and other mechanisms, though the extent and conditions for recovery in grapevine remain subjects of active research. Understanding vulnerability to cavitation quality enables assessment of drought tolerance limits, prediction of hydraulic failure risk under severe stress, and evaluation of the safety margins between operating water potentials and dysfunction thresholds.
Vulnerability_To_Cavitation_Quality serves as the parent class for specific vulnerability parameters (P50, P88, Resistance). This quality determines drought tolerance limits through embolism susceptibility. The class supports drought management and stress tolerance assessment through vulnerability characterization.
Embolievulnerabilität
Kavitationsvulnerabilität
Xylemvulnerabilität
cavitation resistance
embolism vulnerability
hydraulische Sicherheit
xylem vulnerability
Choat B et al. (2012) Global convergence in the vulnerability of forests to drought. Nature 491(7426): 752-755 Brodersen CR, McElrone AJ (2013) Maintenance of xylem network transport capacity: a review of embolism repair in vascular plants. Frontiers in Plant Science 4: 108
Vulnerability To Cavitation Quality
The VVMD marker series contributed the landmark parentage discoveries that transformed understanding of cultivar origins — using VVMD5, VVMD7, VVMD25, VVMD27, VVMD28, and VVMD32 together with VVS2, VrZAG62, and VrZAG79, Bowers and Meredith (1997) demonstrated that Cabernet Sauvignon is the progeny of Cabernet Franc × Sauvignon Blanc, Bowers et al. (1999) identified Pinot and Gouais Blanc as parents of at least 16 major cultivars including Chardonnay, Gamay Noir, Aligoté, Auxerrois, and Melon, and subsequent studies using the same markers established that Syrah originated from Dureza × Mondeuse Blanche (Vouillamoz and Grando 2006) — the VVMD series markers were selected for the OIV panel because of their high polymorphism information content (PIC 0.74-0.87 across V. vinifera), reliable amplification, and the extensive allele frequency data available from the UC Davis germplasm collection of over 3,000 accessions.
A microsatellite marker data record for the VVMD (Vitis Vinifera Microsatellite Davis) series of SSR markers developed at the University of California, Davis by Bowers, Dangl, and Meredith, comprising dinucleotide repeat microsatellites isolated from Vitis vinifera genomic libraries that include 6 of the 9 markers in the OIV standard identification panel (VVMD5, VVMD7, VVMD25, VVMD27, VVMD28, VVMD32) and that have been instrumental in establishing the parentage relationships of major wine grape cultivars.
VVMD_Marker_Series documents the most influential set of microsatellite markers in grapevine genetics, both in terms of their contribution to the OIV standard panel (6 of 9 markers) and their role in the parentage revolution that transformed ampelography from a descriptive to a molecular science. The VVMD markers were developed in two phases — VVMD1-VVMD8 published by Bowers et al. (1996) from a dinucleotide-enriched genomic library of cv. Pinot Noir, and VVMD24-VVMD36 published by Bowers et al. (1999) from additional library screening: the naming convention VVMD (Vitis Vinifera Microsatellite Davis) reflects the species of origin and institution. The markers map to specific chromosomal positions in the PN40024 reference genome — VVMD5 on chromosome 16, VVMD7 on chromosome 7, VVMD25 on chromosome 11, VVMD27 on chromosome 5, VVMD28 on chromosome 3, VVMD32 on chromosome 4 — providing coverage of 6 of the 19 chromosomes within the 9-locus standard panel. The allele frequency distributions at VVMD loci have been extensively characterized in multiple populations representing all major viticultural regions, providing the statistical foundation for parentage likelihood calculations and population structure analysis.
Bowers-Meredith SSR series
Bowers-Meredith-SSR-Serie
UC Davis microsatellite markers
UC-Davis-Mikrosatelliten
VVMD SSR loci
VVMD-Marker-Serie
Bowers JE et al. (1996) Development and characterization of additional microsatellite DNA markers for grape, American Journal of Enology and Viticulture 47: 121-130 Bowers JE et al. (1999) Historical genetics: the parentage of Chardonnay, Gamay, and other wine grapes of northeastern France, Science 285: 1562-1565 Bowers JE and Meredith CP (1997) The parentage of a classic wine grape, Cabernet Sauvignon, Nature Genetics 16: 84-87 This P et al. (2004) Development of a standard set of microsatellite reference alleles, Theoretical and Applied Genetics 109: 1448-1458
VVMD Marker Series
The VVS marker series published by Thomas and Scott (1993) comprised VVS1 through VVS5, isolated from a Muscat Gordo Blanco (Muscat of Alexandria) genomic library enriched for (CT)n dinucleotide repeats — of these, VVS2 (located on chromosome 11, allele range 123-165 bp in V. vinifera, 22 alleles documented, PIC 0.84, expected heterozygosity 0.86) emerged as the single most informative marker in the OIV standard panel and was included as the first locus (OIV descriptor 801) due to its exceptional polymorphism, reliable amplification across all Vitis species and interspecific hybrids, and minimal null allele frequency (<0.02 in V. vinifera). VVS2 has been genotyped in more grapevine accessions than any other single molecular marker, with allele size data available for over 8,000 cultivars across the VIVC and European Vitis Database, making it the de facto anchor locus for cross-database genotype matching.
A microsatellite marker data record for the VVS (Vitis Vinifera Sydney) series of SSR markers developed at the Commonwealth Scientific and Industrial Research Organisation (CSIRO) Plant Industry, Adelaide and Sydney, by Thomas and Scott, comprising the earliest published Vitis microsatellite markers including VVS2 — the first locus in the OIV standard 9-marker identification panel and one of the most widely genotyped microsatellite loci in plant genetics.
VVS_Marker_Series documents the pioneering microsatellite development effort that initiated the molecular marker era in grapevine genetics. The 1993 publication by Thomas and Scott represented the first demonstration that microsatellite markers — already proven highly polymorphic in human genetics — could be developed for and applied to Vitis vinifera cultivar identification, opening the pathway for all subsequent SSR-based grapevine genotyping: the VVS markers were isolated using the then-novel approach of constructing genomic libraries enriched for specific repeat motifs through hybridization capture, a technique that dramatically increased the efficiency of microsatellite isolation compared to random library screening. Although the VVS series is numerically small (5 loci) compared to the later VVMD and VMC series, its historical significance is disproportionate — VVS2 became the most widely used single locus in grapevine molecular identification and its inclusion as the first marker in the OIV standard panel acknowledges both its technical performance and its foundational role in establishing SSR-based cultivar identification as the international standard.
CSIRO microsatellite markers
CSIRO-Mikrosatelliten
Thomas-Scott SSR series
Thomas-Scott-SSR-Serie
VVS SSR loci
VVS-Marker-Serie
Thomas MR and Scott NS (1993) Microsatellite repeats in grapevine reveal DNA polymorphisms when analysed as sequence-tagged sites (STSs), Theoretical and Applied Genetics 86: 985-990 This P et al. (2004) Development of a standard set of microsatellite reference alleles, Theoretical and Applied Genetics 109: 1448-1458 Sefc KM et al. (2001) Microsatellite markers for grapevine, in Roubelakis-Angelakis KA (ed) Molecular Biology and Biotechnology of the Grapevine, Kluwer pp: 433-463
VVS Marker Series
Véraison color quality monitoring in Napa Valley Cabernet Sauvignon employs twice-weekly cluster photography of tagged vines from 5% color change through completion, with image analysis software quantifying the progression from initial pink touch on sun-exposed berry shoulders (CIELAB a* transitioning from negative to positive values) through the characteristic mottled purple-and-green appearance of mid-véraison (40-60% berry coloration) to the uniform dark purple (a* greater than 20) that indicates véraison completion this detailed color staging enables prediction of harvest date at 45±5 days from 50% véraison with 90% accuracy across vintages, supporting premium wine production planning.
Véraison_Color_Quality represents the transitional coloration state of grape berries during the onset of ripening, when dramatic color change marks the shift from the green, photosynthetically active pre-véraison berry to the mature, sugar-accumulating ripening berry. Véraison (from French, meaning "change of color") is defined by this visible color transition in colored cultivars, though the underlying physiological changes occur in all cultivars including white varieties where color change is less visually dramatic. During véraison, chlorophyll degradation unmasks carotenoid pigments producing yellow-green to golden hues in white cultivars, while anthocyanin biosynthesis activation produces the progressive appearance of pink, red, and purple coloration in colored cultivars. The véraison color state is characteristically heterogeneous at both berry and cluster scales, with individual berries at different stages of transition and the overall cluster displaying a mixture of green, transitioning, and colored berries. This heterogeneity typically spans 7-14 days from first berry coloration to completion of visible color change across the cluster. Véraison color assessment employs visual estimation of the percentage of berries showing color change, categorical staging of individual berries (green, pink-touch, half-colored, fully colored), or instrumental measurement of color parameters on sample populations. The timing and duration of véraison color transition responds to temperature, with warmer conditions accelerating the process, and to crop load, with heavily cropped vines showing delayed and extended véraison periods. Véraison marks a critical phenological stage for vineyard management decisions including irrigation regime adjustments, crop load estimation, and harvest date projection.
Véraison_Color_Quality specifically captures the transitional color state during the véraison phenological stage, distinct from Mature_Color_Quality which represents the final ripening coloration. The French term véraison is standard in international viticulture terminology and is retained in the class name. For white cultivars, véraison color transition is less visually dramatic but can be assessed as the shift from bright green toward yellow-green or golden hues. The heterogeneity characteristic of véraison color (different berries at different stages) distinguishes this stage from both the uniform green of pre-véraison and the more uniform coloration of mature fruit. BBCH stage 81 corresponds to the beginning of véraison color change.
Reifebeginnfärbung
Umfärbungsstadium
Véraison-Färbung
color change onset
color turning stage
ripening color transition
véraison coloration
Coombe BG (1995) Growth Stages of the Grapevine: Adoption of a System for Identifying Grapevine Growth Stages. Australian Journal of Grape and Wine Research 1(2): 104-110 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 225-267
Véraison Color Quality
Warming sensation assessment comparing two Napa Valley Cabernet Sauvignons of identical 15% alcohol reveals integration differences affecting quality perception, with first wine showing well-integrated warmth (rating 5/10) where alcohol heat emerges gradually and blends with fruit concentration and tannin structure to create powerful but harmonious finish, while second wine displays aggressive heat (8/10) where alcohol jumps out as burning sensation on mid-palate and finish, detracting from fruit expression and creating harsh rather than powerful impression, demonstrating how warming sensation quality depends on integration as much as absolute alcohol level.
Warming sensation quality is a temperature perception quality that represents the perception of warmth or heat during wine consumption arising primarily from alcohol activation of heat-sensing receptors, characterizing the warming, heating, or potentially burning tactile impression that contributes to perceived body, finish length, and overall power in wines above moderate alcohol levels. Within the BFO framework, this quality specifically addresses the warm-hot dimension of thermal perception derived from alcohol's chemesthetic effect, distinguishing alcohol warmth from actual temperature warmth while recognizing that alcohol-induced heat represents one of the most significant mouthfeel contributors in modern full-bodied wines. The ontological significance lies in warming sensation serving as both a positive quality contributor (moderate warmth adding to body and finish) and a potential quality deficiency (excessive heat creating burning that overwhelms other sensations), with the boundary between beneficial warmth and detrimental heat depending on wine style, alcohol integration, and balance with other components. The physiological basis for alcohol warming involves ethanol activation of TRPV1 ion channels (the same receptors that detect capsaicin heat), with alcohol lowering the channel's activation threshold so that body temperature triggers the heat response, creating the sensation of warmth that increases with alcohol concentration, reaching burning perception at high concentrations or with poor integration. The compositional determinants of warming sensation include alcohol concentration (primary factor, with perception threshold around 10% and increasing intensity with higher levels), the rate of alcohol perception (rapid onset suggesting poor integration), and the modulation by other components (sweetness, glycerol, and body potentially masking alcohol heat while high acidity may enhance it). The quality implications of warming sensation depend on balance and style, with moderate warmth (appropriate for full-bodied styles) contributing positively to body and finish, while excessive heat indicates either over-alcoholic wine or poor integration that detracts from harmony. Understanding warming sensation quality enables alcohol balance assessment, style-appropriate warmth evaluation, and quality judgment distinguishing integrated warmth contributing positively from harsh heat indicating deficiency.
Warming_Sensation_Quality_(Alcohol) addresses the alcohol-derived warmth dimension of thermal perception, representing a significant mouthfeel contributor in contemporary wines often exceeding 14% alcohol. This quality may be positive (integrated warmth adding to body and finish) or negative (harsh heat indicating poor balance). The class supports alcohol integration assessment and quality evaluation distinguishing beneficial warmth from detrimental burning.
Alkoholwärme
Brennen
Wärmeempfindung
alcohol heat
burning sensation
ethanol warmth
heißer Abgang
hot finish
Green BG (1987) The sensitivity of the tongue to ethanol. Annals of the New York Academy of Sciences 510(1): 315-317 Nurgel C, Pickering GJ (2006) Modeling of sweetness and bitterness in ice wine. Journal of Food Quality 29(3): 256-268
Warming Sensation Quality (Alcohol)
Chardonnay showing water absorptive feeder roots in July, sap flow measurements showing vine transpiring 4.2 liters per day, isotope tracer (deuterium-labeled water) demonstrating feeder roots <2 mm providing 81% of total water uptake, water absorptive roots concentrated 15-40 cm depth where soil moisture adequate, aquaporin gene expression analysis showing high PIP1 and PIP2 levels in feeder roots, water absorptive feeder roots demonstrating critical hydraulic function supporting transpiration
Water absorptive feeder root is absorptive feeder root emphasizing water uptake function and hydraulic conductivity, characterized by aquaporin water channels facilitating rapid water movement across membranes, favorable geometry with high surface-to-volume ratio optimizing water absorption, thin cortex minimizing radial transport resistance, root hairs and mycorrhizae extending water uptake zone, and demonstrated capacity to supply vine transpirational demand through measured water flux, with water absorptive feeder roots showing functional characteristics including high hydraulic conductivity measured through root pressure or uptake experiments, aquaporin expression particularly PIP plasma membrane intrinsic proteins enabling rapid membrane water transport, responsive regulation with aquaporin activity adjusting to transpirational demand and soil water availability, diurnal uptake patterns with water absorption peaking during day correlating with transpiration, and critical importance providing 70-85% of total vine water uptake particularly from upper moist soil layers where water absorptive feeder roots concentrated. Water absorptive feeder root hydraulic characteristics includes= showing high conductivity (water absorptive roots showing hydraulic conductivity 10-50 fold higher per unit surface area than structural roots), exhibiting radial pathway (water moving radially from soil through epidermis, cortex, endodermis to xylem requiring passage through or around multiple cell layers), demonstrating aquaporin facilitation (water channel proteins increasing membrane permeability 5-20 fold above lipid bilayer baseline), revealing pathway components (water moving through apoplastic, symplastic, and transcellular pathways with relative contributions varying), and displaying resistance distribution (major hydraulic resistances occurring at endodermis and root-soil interface). Water absorptive feeder root aquaporin function includes= showing PIP expression (plasma membrane intrinsic proteins particularly PIP1 and PIP2 isoforms abundant in water absorptive root membranes), exhibiting gating regulation (aquaporins showing open-closed gating responding to pH, calcium, phosphorylation, and other signals), demonstrating isoform diversity (multiple aquaporin genes expressed potentially with different regulatory properties and localizations), revealing trafficking control (aquaporin abundance in membranes regulated through vesicle trafficking and degradation), and displaying functional importance (aquaporin activity critically affecting root hydraulic conductivity and water uptake capacity). Water absorptive feeder root geometric optimization includes= showing high surface area (fine diameter creating large surface area per unit volume maximizing soil contact for water uptake), exhibiting cylindrical form (cylindrical geometry optimizing surface area per volume for radial water flow), demonstrating thin cortex (minimal cortex thickness reducing radial transport distance from epidermis to xylem), revealing appropriate xylem (adequate xylem vessels for axial water transport though not extensive given small root size), and displaying branching architecture (intensive branching creating network thoroughly exploring soil water). Water absorptive feeder root surface enhancements includes= showing root hair contribution (abundant root hairs extending effective water uptake zone 5-20 fold increasing absorption), exhibiting mycorrhizal hyphae (external fungal hyphae potentially contributing to water uptake particularly under drought), demonstrating soil contact (close association between water absorptive root surface and soil particles minimizing interface resistance), revealing mucilage effects (root mucilage potentially affecting rhizosphere hydraulic properties and root-soil contact), and displaying soil water access (water absorptive roots accessing water in small pores through capillary contact). Water absorptive feeder root uptake dynamics includes= showing transpiration coupling (water uptake driven by shoot transpirational demand creating soil-plant-atmosphere continuum), exhibiting diurnal patterns (water absorption peaking during day when stomata open and transpiration high declining at night), demonstrating rapid response (water uptake quickly adjusting to changing transpirational demand or soil water availability), revealing threshold effects (water uptake declining sharply when soil water potential decreases below critical values), and displaying capacity limits (maximum water uptake limited by root hydraulic conductance and available soil water). Water absorptive feeder root drought response includes= showing vulnerability (water absorptive feeder roots highly sensitive to drought being among first roots to lose function or die), exhibiting aquaporin regulation (aquaporin expression or activity potentially declining under severe drought protecting against excessive water loss), demonstrating production decline (water absorptive root production decreasing under water deficit reducing resource allocation to vulnerable tissues), revealing mortality increase (water absorptive root death rates increasing under drought reducing functional population), and displaying recovery capacity (water absorptive roots rapidly regrowing when moisture restored enabling quick recovery). Water absorptive feeder root seasonal patterns includes= showing spring maximum (water absorptive root function peaking spring when soil moisture high and transpirational demand increasing), exhibiting summer decline (water uptake potentially limited summer by soil drying reducing absorptive root function), demonstrating fall recovery (water uptake recovering fall with rainfall and reduced transpiration), revealing winter minimum (water absorption minimal winter when transpiration very low and roots dormant), and displaying annual integration (cumulative annual water uptake through absorptive roots meeting vine transpirational water requirement). Water absorptive feeder root in vine water status includes= determining water availability (water absorptive root capacity limiting vine water uptake and leaf water potential), enabling transpiration support (adequate water absorptive roots necessary for maintaining stomatal conductance and photosynthesis), demonstrating stress sensitivity (water absorptive root dysfunction contributing to drought stress symptoms), revealing irrigation importance (irrigation maintaining water absorptive root function supporting consistent water supply), and displaying management implications (practices affecting water absorptive root distribution and function critically impacting vine water relations).
Water_Absorptive_Feeder_Root emphasizes water uptake and hydraulic function and is distinguished from nutrient absorption by focus on water transport. Critical for understanding aquaporins, hydraulic conductivity, radial pathway, transpiration coupling, drought sensitivity, and the specialized roots providing 70-85% of vine water uptake supporting transpiration.
Hydraulic absorptive root
Hydraulische Saugwurzel
Transpiration-supporting root
Transpiration-unterstützende Wurzel
Wasser-absorbierende Feinwurzel
Water-uptake root
Water Absorptive Feeder Root
Water activity quality monitoring in Tokaji Aszú production ensures the botrytized berries have achieved sufficient dehydration (aw 0.75-0.85) through noble rot desiccation before collection into puttonyos baskets, with water activity below 0.80 indicating the microbial stability required for extended pre-fermentation maceration (24-48 hours) that extracts the concentrated sugars, acids, and botrytis-derived glycerol without spoilage risk fresh berries at aw 0.97-0.99 would rapidly succumb to secondary fungal colonization and bacterial spoilage during equivalent maceration, making water activity reduction through noble rot essential to the traditional production process.
Water_Activity_Quality represents the chemical quality characterizing the thermodynamic availability of water in grape tissue, expressed as the ratio of water vapor pressure over the sample to that over pure water at the same temperature (aw = p/p₀). Water activity provides a more functionally relevant measure of water status than moisture content for predicting microbial growth, enzymatic reaction rates, and chemical stability, as it reflects the energy state of water rather than simply its quantity. In fresh grape berries at full hydration, water activity approaches 0.99, indicating nearly free water availability. As berries dehydrate or as solute concentration increases, water activity decreases, with values below 0.95 increasingly limiting microbial growth and below 0.85 substantially inhibiting most spoilage organisms. The relationship between moisture content and water activity follows a sorption isotherm specific to the tissue composition, with the high sugar content of grapes producing characteristic depression of water activity at given moisture levels. During raisining or appassimento, water activity decreases from approximately 0.98 in fresh grapes to 0.60-0.75 in dried products, conferring microbial stability through water limitation rather than chemical preservation. Water activity affects not only microbial growth but also non-enzymatic browning (Maillard reactions), lipid oxidation, and enzyme activity, all of which show water activity optima that influence quality during storage and processing. Measurement of water activity employs specialized instruments (chilled mirror dewpoint, capacitance sensors) that equilibrate sample and headspace to determine the equilibrium relative humidity corresponding to water activity. For winemaking applications, water activity of must and wine (typically 0.92-0.98) relates to fermentation behavior and microbial stability, with lower water activity from high sugar concentration potentially limiting yeast activity in very concentrated musts.
Water_Activity_Quality addresses the thermodynamically meaningful water parameter that better predicts biological and chemical behavior than moisture content alone. The water activity concept derives from food science where it enables prediction of shelf stability and microbial risk. The sorption isotherm relating moisture content to water activity is matrix-specific, meaning identical moisture contents in different materials produce different water activities. Water activity measurement requires specialized instrumentation distinct from moisture determination methods.
Wasseraktivität
Wasserverfügbarkeit
aw value
aw-Wert
equilibrium relative humidity quality
thermodynamic water status
thermodynamischer Wasserstatus
water availability
Labuza TP and Altunakar LN (2007) Water Activity Prediction and Moisture Sorption Isotherms. In: Barbosa-Cánovas GV et al. (eds) Water Activity in Foods: Fundamentals and Applications, Blackwell Publishing pp.: 109-154 Serratosa MP et al. (2008) Changes in Colour and Phenolic Compounds during the Raisining of Grape cv. Pedro Ximénez. Journal of Agricultural and Food Chemistry 56(8): 2810-2816
Water Activity Quality
The water application rate for a deficit-irrigated Merlot vineyard in the Columbia Valley (Washington State) specifies 15 liters per vine per event (equivalent to 5.6 mm/ha at 3,700 vines/ha) applied twice weekly during the peak demand period of January through February (Southern Hemisphere equivalent July-August), totaling approximately 120 liters per vine per week or 1,800-2,200 liters per vine per season (180-220 mm/ha/season) — this seasonal total represents approximately 45% of the estimated full ETc of 450-500 mm, consistent with the regulated deficit irrigation strategy targeting moderate water stress for premium red wine production in the semi-arid Columbia Valley where annual rainfall of 150-200 mm provides less than 40% of vine water requirements.
An irrigation protocol description that documents the volume of water applied per irrigation event or per time period, expressed as liters per vine per event, millimeters per hectare per event, cubic meters per hectare per season, or percentage of reference evapotranspiration replaced, specifying the quantitative water delivery parameter that determines the depth and lateral extent of soil wetting and the degree to which vine water demand is met or intentionally restricted.
Water_Application_Rate translates the conceptual irrigation strategy into operational water delivery quantities. The rate must be calibrated to the specific combination of soil type (water-holding capacity and infiltration rate), emitter output (typically 1-4 L/h for drip systems), irrigation duration, and vine spacing to achieve the target soil wetting pattern and vine water supply: sandy soils with low water-holding capacity (80-120 mm/m) require more frequent, smaller applications than clay or loam soils with higher capacity (180-250 mm/m) to maintain water within the root zone without drainage losses. Seasonal water application totals for deficit-irrigated premium wine grapes typically range from 100-250 mm in Mediterranean climates (California, southern France, southern Australia) to 300-500 mm in semi-arid continental climates (eastern Washington, Mendoza, central Spain) where atmospheric demand exceeds rainfall by larger margins. Appellation regulations in regions permitting irrigation may specify maximum seasonal water application rates as a quality control mechanism.
Beregnungsmenge-Spezifikation
Bewässerungsvolumen-Angabe
Wasserausbringungsmenge
application depth specification
irrigation volume specification
water delivery rate
Williams LE and Ayars JE (2005) Grapevine water use and the crop coefficient are linear functions of the shaded area measured beneath the canopy, Agricultural and Forest Meteorology 132: 201-211 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp: 395-410 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 168-195
Water Application Rate
Water content quality management in Amarone della Valpolicella production deliberately reduces berry water from 78-82% at harvest to 60-68% through three to four months of appassimento (controlled drying in fruttai), concentrating sugars from 22-26°Brix to 26-32°Brix and proportionally concentrating acids, phenolics, and flavor compounds this controlled dehydration, monitored through weight loss (30-40% target) and periodic sampling, transforms the Corvina, Corvinone, and Rondinella grapes into the intensely concentrated, raisined material yielding the powerful, complex wines that define the Amarone style.
Water_Content_Quality encompasses all chemical qualities that characterize the concentration, distribution, and thermodynamic state of water in grape berry tissue, capturing the fundamental solvent medium that affects berry composition, processing behavior, and wine yield. Water constitutes 75-85% of fresh grape berry mass, serving as the solvent for sugars, acids, phenolics, and other solutes while maintaining cell turgor and enabling metabolic processes. The water content of berries changes dynamically during ripening, with net water accumulation during the initial growth phase (Stage I), minimal change during the lag phase (Stage II), and complex water dynamics during the ripening phase (Stage III) when phloem-delivered water accompanies sugar import while xylem connectivity declines and transpirational loss continues. This category subdivides into Moisture_Content_Quality addressing the gravimetrically determined water fraction Water_Activity_Quality capturing the thermodynamic availability of water affecting microbial growth and chemical reaction rates Relative_Water_Content_Quality describing hydration status relative to full turgor and Juice_Yield_Quality characterizing the extractable liquid fraction relevant to processing. Water content profoundly affects must composition through concentration effects, as water loss concentrates all solutes proportionally while water gain dilutes them. Late-season dehydration, whether from extended hang time, noble rot infection, or deliberate drying (appassimento), can increase sugar concentration by 20-50% while proportionally concentrating acids, phenolics, and flavor compounds. Conversely, late-season rainfall causing berry rehydration dilutes must composition and can precipitate splitting in tight-skinned varieties. The balance between water loss (transpiration, cuticular permeation) and water gain (phloem import, rain absorption) determines final berry water content at harvest. Understanding water content quality enables prediction of juice yield, must composition, and optimal harvest timing for different wine styles ranging from light table wines to concentrated dessert wines.
Water_Content_Quality is positioned as a direct subclass of Chemical_Quality, recognizing water as a chemical compound whose content profoundly affects all other compositional parameters through concentration and dilution effects. The dynamic nature of berry water content during ripening and post-harvest distinguishes it from more stable compositional parameters. Water content qualities describe current hydration state distinct from water-related dispositions (e.g., dehydration susceptibility) and processes (e.g., transpiration, phloem import). The multiple subclasses reflect different measurement approaches and functional interpretations of water status.
Beerenwassergehalt
Feuchtigkeitsstatus
Hydratationsqualität
Wassergehalt
berry water content
hydration quality
moisture status
water composition quality
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 225-280 Coombe BG and McCarthy MG (2000) Dynamics of Grape Berry Growth and Physiology of Ripening. Australian Journal of Grape and Wine Research 6(2): 131-135
Water Content Quality
Water deficit severity quality classification in Napa Valley Cabernet Sauvignon established threshold values and quality associations for three severity categories using midday stem water potential as the diagnostic indicator. Mild water deficit severity quality was defined by stem water potential of -0.8 to -1.0 MPa, associated with modest stomatal closure maintaining 85% of unstressed photosynthesis, continued but reduced shoot growth, and subtle quality enhancement through improved vine balance with yield reduction of less than 10%. Moderate water deficit severity quality at -1.0 to -1.2 MPa showed pronounced stomatal closure reducing photosynthesis to 65% of potential, growth cessation by véraison, and significant quality enhancement including 25% higher anthocyanin concentration and improved tannin structure, with yield reduction of 15-25% considered acceptable for premium production. Severe water deficit severity quality below -1.2 MPa indicated stress exceeding beneficial ranges, with stomatal and non-stomatal limitations, premature leaf senescence, incomplete ripening, and quality compromise despite concentration effects, establishing -1.2 MPa as the threshold requiring irrigation intervention regardless of quality objectives.
Water deficit severity quality is a drought stress indicator quality that inheres in a grapevine organism and represents the categorization of water stress intensity into functionally distinct levels characterized by specific physiological responses, visual symptoms, and quality implications, enabling standardized communication about vine water status and providing the framework for irrigation management targeting specific severity categories to achieve production objectives ranging from stress avoidance for maximum yield through controlled moderate stress for quality enhancement. This quality provides the classification structure within which specific stress measurements are interpreted, recognizing that continuous water potential or other indicator values gain practical meaning when mapped to discrete severity categories with established physiological and quality associations. The ontological significance of water deficit severity quality lies in its role translating quantitative measurements into actionable management categories, enabling irrigation decisions based on target severity levels rather than requiring interpretation of raw physiological values for each decision point, and facilitating communication among researchers, advisors, and producers using shared severity terminology with consistent meaning. The biological basis of water deficit severity classification reflects the non-linear relationship between declining water availability and vine response, wherein distinct physiological thresholds separate qualitatively different response regimes rather than smooth continuous gradients across the full stress range. The transition from mild to moderate stress involves stomatal closure becoming sufficiently pronounced to reduce photosynthesis despite initial stomatal sensitivity serving to maintain carbon assimilation, while the transition from moderate to severe stress involves the onset of hydraulic dysfunction and non-stomatal limitations that persist beyond stress relief and may cause lasting damage. The quality implications of water deficit severity vary systematically across categories, with mild stress often enhancing quality through subtle vine balance improvements, moderate stress providing pronounced quality benefits through growth restriction and concentration effects for red wine production, and severe stress compromising quality through impaired ripening and stress-induced off-flavors. Assessment of water deficit severity quality involves measurement of water status indicators and classification according to established thresholds specific to the indicator employed, cultivar characteristics, and regional calibration reflecting local climate and soil conditions.
Water_Deficit_Severity_Quality is_a Drought_Stress_Indicator_Quality, serving as parent class for Mild_Water_Stress_Quality, Moderate_Water_Stress_Quality, and Severe_Water_Stress_Quality which define specific severity categories. This class addresses the classification framework while child classes capture individual category characteristics. Threshold values vary with indicator choice (predawn vs. midday, leaf vs. stem water potential), cultivar, and regional calibration. The non-linear quality response with optimal outcomes at moderate stress represents key viticultural principle. Critical for regulated deficit irrigation implementation targeting specific severity categories.
Dürreschwere-Klassifizierung
Stressintensitätskategorie
Wasserdefizit-Schweregrad
Wasserstress-Intensität
drought severity classification
stress intensity category
water deficit level
water stress intensity
Ojeda H et al. (2002) Influence of pre- and postveraison water deficit on synthesis and concentration of skin phenolic compounds during berry growth of Vitis vinifera cv. Shiraz. American Journal of Enology and Viticulture 53:261-267 Castellarin SD et al. (2007) Water deficits accelerate ripening and induce changes in gene expression regulating flavonoid biosynthesis in grape berries. Planta 227:101-112 Chaves MM et al. (2010) Grapevine under deficit irrigation: hints from physiological and molecular data. Annals of Botany 105:661-676
Water Deficit Severity Quality
Water potential monitoring in Napa Valley Cabernet Sauvignon using predawn and midday pressure chamber measurements guides deficit irrigation strategy, with target predawn values of -0.2 to -0.4 MPa indicating adequate soil water availability while midday values of -1.2 to -1.4 MPa induce moderate stress promoting flavor concentration without compromising photosynthesis, with irrigation triggered when predawn values decline below -0.5 MPa or midday values exceed -1.6 MPa to prevent excessive stress.
Water potential quality is a physical quality that inheres in grapevine tissues and represents the thermodynamic measure of water's free energy status relative to pure water at atmospheric pressure, characterizing the driving force for water movement through the soil-plant-atmosphere continuum where water flows spontaneously from regions of higher (less negative) to lower (more negative) water potential. Within the BFO framework, this quality serves as the master variable integrating osmotic, pressure, matric, and gravitational components that together determine the energy state of water in plant tissues, enabling prediction of water flux direction and magnitude throughout the vine hydraulic system. The ontological significance lies in water potential providing a unified framework for understanding water relations across scales from cellular to whole-plant to ecosystem, where the gradient in water potential drives the transpiration stream from soil through roots, stems, and leaves to atmosphere. Water potential in grapevines ranges from near zero in well-watered soil to -0.3 MPa in unstressed leaves during transpiration, declining to -1.5 to -2.5 MPa under moderate water stress and potentially reaching -3.0 to -4.0 MPa under severe drought before hydraulic failure. The relationship between water potential and physiological function defines critical thresholds for stomatal closure, photosynthetic decline, and hydraulic dysfunction that vary among cultivars reflecting their drought adaptation strategies. Measurement approaches include pressure chamber (Scholander bomb) for bulk tissue water potential, thermocouple psychrometry for precise laboratory determination, and leaf porometry combined with stomatal models for indirect estimation. The diurnal dynamics of water potential, with minimum values at midday when transpiration demand peaks and recovery toward predawn equilibrium with soil, provide the temporal framework for water stress assessment and irrigation scheduling. Understanding water potential quality enables mechanistic interpretation of vine water status, prediction of physiological responses to drought, and optimization of irrigation management for yield and quality objectives.
Water_Potential_Quality serves as the parent class for measurement-timing-specific water potential qualities (Predawn, Midday, Stem, Leaf), establishing the ontological framework for plant water status assessment. This quality equals the sum of component potentials (Ψ = Ψs + Ψp + Ψm + Ψg), with osmotic and pressure components dominant in most tissue contexts. The class supports irrigation scheduling, stress physiology research, and climate adaptation assessment through quantitative characterization of vine water status.
Gewebewasserpotential
Wasserpotential
Wasserzustand
hydraulic potential
hydraulisches Potential
tissue water energy
water status
Choné X et al. (2001) Stem water potential is a sensitive indicator of grapevine water status. Annals of Botany 87(4): 477-483 Williams LE, Araujo FJ (2002) Correlations among predawn leaf, midday leaf, and midday stem water potential and their correlations with other measures of soil and plant water status in Vitis vinifera. Journal of the American Society for Horticultural Science 127(3): 448-454
Water Potential Quality
Water relations parameters vary substantially among grapevine genotypes, with whole-vine hydraulic conductance ranging from 15-45 mmol H₂O/MPa/s across rootstock-scion combinations. Daily water flux through a mature vine reaches 20-50 L under high evaporative demand, requiring coordinated hydraulic capacity from roots through stems to leaves. Water potential gradients from soil (-0.1 to -1.5 MPa) through roots, stem, and leaves (midday values -0.8 to -1.8 MPa) drive transpirational flow along the resistance pathway. Rootstock genotype accounts for 40-60% of variation in scion water status under deficit irrigation, demonstrating strong genetic control of water relations.
A grapevine disposition that is the inherited capacity of a Vitis vinifera plant to acquire, transport, and regulate water movement through the soil-plant-atmosphere continuum, encompassing traits related to root water uptake, xylem hydraulic conductance, stomatal regulation, and the maintenance of favorable tissue water status under varying environmental conditions.
Water_Relations_Disposition serves as the top-level parent class for traits governing plant water economy. The category encompasses water acquisition (root uptake), long-distance transport (xylem conductance), and loss regulation (stomatal control, addressed separately under drought tolerance). Water relations traits critically determine vine productivity, fruit composition, and stress tolerance across the range of vineyard water environments from rain-fed to fully irrigated systems. The trait complex shows strong rootstock influence, making rootstock selection a primary tool for water relations management.
Wasserbeziehungen
Wasserhaushalt
hydraulic capacity
hydraulische Disposition
plant water relations trait
water management disposition
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 197-245 Lovisolo C et al. (2010) Grapevine adaptations to water stress: new perspectives about soil/plant interactions. Theoretical and Experimental Plant Physiology 22:1-15
Water Relations Disposition
A water shoot emerging from a latent bud on the trunk of Vitis vinifera cv. Sangiovese in Tuscany, Italy, at 60 cm above the graft union, developing in response to severe hail damage that defoliated 85% of the primary canopy in June, achieving 220 cm total growth by September with 22 nodes, internode lengths averaging 14-16 cm, individual leaves reaching 240 cm² area with light green coloration and reduced anthocyanin pigmentation compared to normal shoots, stem diameter of 13 mm at mid-shoot despite current-season origin, showing no inflorescence primordia when dissected in autumn, and requiring removal during dormant pruning to prevent permanent framework disruption.
A vigorous vegetative shoot arising from latent or adventitious buds on older woody structures including trunks, arms, and cordons in Vitis vinifera, characterized by exceptional growth rates often exceeding 5-8 cm per day, very long internodes typically 12-20 cm or greater, large leaves frequently surpassing 200 cm² area, thick succulent stems, and absence of reproductive structures, typically emerging in response to severe pruning, physical damage, pathogen infection, or imbalanced carbohydrate-to-nitrogen ratios favoring excessive vegetative growth. Water shoots represent the grapevine's emergency response mechanism for canopy rebuilding following catastrophic loss of primary photosynthetic surface area, developing from dormant buds that may have remained quiescent for multiple years along trunk and arm surfaces, and exhibiting juvenilized growth characteristics including reduced lignification, extended growth duration, poor cold hardiness, and low fruitfulness potential in subsequent seasons. These shoots create significant management challenges in production viticulture through resource diversion from fruit development, canopy disorder, increased disease pressure in dense growth zones, and mechanical harvest interference, but also provide opportunities for framework reconstruction following severe damage events or intentional trunk renewal strategies in aging vineyards.
Water shoots are distinguished from normal lateral shoots by their origin from latent or adventitious buds on woody structures rather than from current-season buds on green shoots, their exceptional vigor exceeding typical lateral shoot growth, and their juvenilized morphological characteristics. The term "water shoot" reflects historical observations that these shoots contain higher water content and lower dry matter percentage than normal shoots, though this is secondary to their developmental rather than compositional distinctiveness. Not all vigorous shoots from woody structures are water shoots - renewal shoots intentionally developed from selected positions represent managed growth rather than emergency response. Water shoots arising from below the graft union (rootstock suckers) require immediate removal to prevent rootstock dominance.
Adventivtrieb
Wasserschoss
Wassertrieb
basal shoot
epicormic shoot
sucker shoot
Based on Winkler et al. (1974) "General Viticulture" University of California Press pp. 228-245 Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 93-108 Lavee (1987) "Usefulness of growth regulators for controlling vine growth and improving grape quality in intensive vineyards" Acta Horticulturae 206=89-108 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 272-294.
Water Shoot
A water stress-induced summer dormant bud on non-irrigated Vitis vinifera cv. Grenache in the Southern Rhône Valley during July-August, with soil water deficit at -2.1 MPa, predawn leaf water potential of -0.9 MPa, complete cessation of shoot growth for four weeks, stem water potential declining to -1.4 MPa, and bud resuming growth within 12 days following 40mm rainfall event that restored predawn leaf water potential to -0.3 MPa.
A summer dormant bud that enters growth arrest specifically in response to severe soil water deficit and plant water stress (typically when predawn leaf water potential falls below -0.6 to -0.8 MPa in Vitis vinifera), characterized by hydraulic signaling-mediated cessation of cell expansion, ABA-regulated stomatal closure in associated leaves, prioritization of water allocation to reproductive structures over vegetative growth, and protective reduction in transpirational surface area through growth suspension. Water stress-induced dormancy represents a drought survival strategy allowing vines to maintain viability during extended dry periods while preserving meristematic potential for growth resumption upon water availability.
Water stress-induced summer dormancy is a central component of deficit irrigation strategies in viticulture, where controlled water stress is used to manage vegetative vigor and enhance fruit quality. The threshold for growth arrest varies among cultivars and rootstocks, with some showing isohydric behavior (tight stomatal control, early growth cessation) and others anisohydric behavior (continued transpiration, later growth arrest). Prolonged severe water stress can transition from reversible dormancy to irreversible damage including bud necrosis.
drought-induced dormant bud
hydraulically dormant bud
trockenstressinduzierte Ruheknospe
water deficit dormant bud
Based on Schultz (2003) "Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought" Plant, Cell & Environment 26=1393-1405 Gambetta et al. (2020) "The physiology of drought stress in grapevine= towards an integrative definition of drought tolerance" Journal of Experimental Botany 71=4658-4676 Charrier et al. (2018) "Evidence for hydraulic vulnerability segmentation and lack of xylem refilling under tension" Plant Physiology 172=1657-1668.
Water Stress Summer Dormant Bud
Whole-shoot hydraulic conductance in grapevines ranges from 8-25 mmol H₂O/m²/s/MPa across genotypes, with high-conductance cultivars such as Grenache and Syrah showing 2× the transport capacity of conservative types like Chardonnay. The transpiration stream moves through a series of hydraulic resistances, with roots contributing 30-50%, stems 20-40%, and leaves 20-40% of total resistance depending on growth stage and environmental conditions. Maximum sap flow velocities of 0.5-2.0 m/hour occur at midday under high vapor pressure deficit, with daily total transport of 20-50 L in mature vines. Xylem transport capacity must exceed maximum transpirational demand by a safety margin to prevent hydraulic failure from cavitation.
A water relations disposition that is the inherited capacity of a Vitis vinifera plant to conduct water through the xylem vascular system from roots to transpiring leaves, determined by xylem vessel anatomy (diameter, length, pit membrane properties), hydraulic pathway architecture, and the maintenance of xylem functionality under tension.
Water_Transport_Disposition serves as the parent class for xylem conductivity and hydraulic architecture traits. Long-distance water transport in the xylem occurs under negative pressure (tension), making the system vulnerable to cavitation and embolism under severe water stress. The balance between transport efficiency (large vessels, high conductance) and transport safety (small vessels, cavitation resistant) represents a fundamental hydraulic trade-off with genotype-specific resolution. Vine training and canopy management modify transport path length and resistance distribution.
Wassertransport
long-distance water movement
vascular hydraulic capacity
vaskuläre Transportkapazität
xylem water transport
Schultz HR (2003) Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought. Plant, Cell and Environment 26:1393-1405 Tyree MT, Zimmermann MH (2002) Xylem Structure and the Ascent of Sap, 2nd Ed, Springer pp.: 175-220
Water Transport Disposition
Water uptake capacity varies 2-3 fold among rootstocks, with deep-rooted types such as 110 Richter and 140 Ruggeri maintaining uptake from subsoil water sources inaccessible to shallow-rooted types like Riparia Gloire. Root water uptake follows a composite transport pathway involving apoplastic flow through cell walls and symplastic flow through membranes regulated by aquaporins. Maximum uptake rates of 0.5-1.5 mL H₂O/g root fresh weight/hour occur under high transpirational demand and adequate soil moisture. Uptake efficiency (water absorbed per unit root biomass) increases with fine root proportion, with rootstocks showing 20-60% fine root content differing correspondingly in uptake efficiency.
A water relations disposition that is the inherited capacity of a Vitis vinifera plant to absorb water from the soil solution through root system activity, determined by root surface area, root hydraulic conductivity, aquaporin expression, and the ability to maintain uptake across varying soil water potentials and temperatures.
Water_Uptake_Disposition serves as the parent class for root hydraulic conductivity and aquaporin activity traits. Water uptake represents the entry point of the soil-plant-atmosphere continuum, integrating soil water availability with root system architecture and membrane transport properties. Uptake limitation during soil drying triggers hydraulic and chemical (ABA) signals that coordinate stomatal closure with reduced water availability. Rootstock breeding for enhanced water uptake efficiency is a strategy for improving drought tolerance without sacrificing productivity.
Wasserakquisitions-Disposition
Wasseraufnahme
Wurzelwasserabsorption
root uptake trait
root water absorption
water acquisition capacity
Gambetta GA et al. (2012) Sugars and abscisic acid signaling orthologs are activated at the onset of ripening in grape. Planta 236:851-863 Lovisolo C, Schubert A (1998) Effects of water stress on vessel size and xylem hydraulic conductivity in Vitis vinifera L. Journal of Experimental Botany 49:693-700
Water Uptake Disposition
Water use efficiency in grapevines ranges from 3-8 g dry matter per kg H₂O transpired (whole-plant WUE) or 2-6 µmol CO₂/mmol H₂O at the leaf level (instantaneous WUE), with substantial genotypic variation. Intrinsic WUE (A/gs, the ratio of photosynthesis to stomatal conductance) ranges from 40-90 µmol CO₂/mol H₂O across cultivars, with Tempranillo and Cabernet Sauvignon showing high values (70-90) and Grenache showing lower values (45-60). WUE increases under moderate water stress as stomatal closure restricts transpiration proportionally more than photosynthesis (non-stomatal limitation). Seasonal WUE of 4-8 kg fruit/m³ water applied is typical for deficit-irrigated wine grapes versus 2-4 kg/m³ for fully irrigated table grapes.
A water relations disposition that is the inherited capacity of a Vitis vinifera plant to produce biomass or yield per unit of water transpired, integrating stomatal regulation of the CO₂/H₂O exchange ratio with mesophyll conductance, photosynthetic capacity, and respiratory losses to determine overall carbon gain relative to water expenditure.
Water_Use_Efficiency_Disposition serves as the parent class for carbon isotope discrimination and transpiration efficiency traits. This trait also appears under Drought_Tolerance_Disposition where the focus is stress-adaptive mechanisms here the emphasis is on general water economy under all conditions. WUE represents a key target for sustainable viticulture, as improving water productivity reduces irrigation requirements and environmental impact. The trait shows complex inheritance and genotype × environment interaction, complicating selection efforts.
WUE trait
Wasserproduktivität
transpiration efficiency
water productivity disposition
Medrano H et al. (2015) Improving water use efficiency of vineyards in semi-arid regions: a review. Agronomy for Sustainable Development 35:499-517 Tomás M et al. (2014) Variability of water use efficiency in grapevines. Environmental and Experimental Botany 103:148-157
Water Use Efficiency Disposition
Comprehensive evaluation of Waterlogging_Tolerance_Disposition at the University of Florida compared survival, growth, and physiological responses across 15 rootstocks subjected to complete soil waterlogging for 30 days during the growing season. Measurements included survival rate, shoot growth, leaf gas exchange, root morphology (adventitious root formation, aerenchyma presence), and root ATP concentration as indicator of energy status. Waterlogging_Tolerance_Disposition varied dramatically among rootstocks. 'Riparia Gloire' (V. riparia) exhibited exceptional tolerance with 100% survival, shoot growth continuing at 65% of control rate, stomatal conductance maintained at 55% of control through formation of adventitious roots (12.5 new roots per plant in aerated surface zone), visible aerenchyma in 45% of root cross-sectional area, and root ATP maintained at 48% of control levels. '1103 Paulsen' (V. berlandieri × V. rupestris) showed good tolerance (95% survival, 50% growth rate, 35% aerenchyma). In contrast, '140 Ruggeri' (V. berlandieri × V. rupestris, but from drier ecotype) displayed poor Waterlogging_Tolerance_Disposition with only 55% survival, growth cessation by day 10, minimal adventitious rooting (2.3 roots/plant), negligible aerenchyma (8% of cross-section), and root ATP collapsing to 12% of control. 'Schwarzmann' showed intermediate tolerance (80% survival, 40% growth). Notably, waterlogging tolerance showed weak correlation with drought tolerance (r = 0.25), indicating these represent distinct adaptive syndromes. These findings demonstrate that Waterlogging_Tolerance_Disposition involves coordinated anatomical and metabolic adaptations that vary substantially among rootstock genotypes.
A Waterlogging_Tolerance_Disposition is an Abiotic_Stress_Tolerance_Disposition that inheres in a Vitis plant and is realized through physiological, anatomical, and metabolic mechanisms that enable survival and continued function under soil flooding conditions where oxygen availability to roots is severely restricted, causing hypoxia or anoxia that would otherwise lead to root death, impaired water and nutrient uptake, and vine decline. This disposition represents the capacity to tolerate excess soil water, important for viticulture in regions with heavy clay soils, high water tables, or periodic flooding, and increasingly relevant as climate change intensifies precipitation events in many wine regions. Waterlogging stress results primarily from oxygen deprivation as soil pore spaces fill with water, limiting oxygen diffusion to roots (10,000× slower in water than air), and secondarily from accumulation of toxic reduced compounds (Fe²⁺, Mn²⁺, H₂S) in anaerobic soil and potentially from pathogen pressure in wet conditions. Root hypoxia impairs aerobic respiration, forcing reliance on inefficient fermentative pathways, depleting carbohydrate reserves, and disrupting ATP-dependent processes including ion uptake and phytohormone synthesis. Waterlogging_Tolerance_Disposition encompasses anatomical adaptations (adventitious root formation providing roots in aerated surface zones, aerenchyma development creating internal oxygen transport pathways), metabolic adaptations (efficient anaerobic metabolism, tolerance of fermentation end-products), and hormonal responses (ethylene-mediated adaptive responses). In grafted grapevines, rootstock genotype is the primary determinant of waterlogging tolerance, with species from riparian habitats (V. riparia, V. rupestris) generally showing superior tolerance compared to those from well-drained hillside environments.
Waterlogging_Tolerance_Disposition is_a Abiotic_Stress_Tolerance_Disposition. Waterlogging tolerance is primarily determined by rootstock genotype. The core problem is root oxygen deprivation, not excess water per se. Child classes address specific mechanisms including Adventitious_Root_Formation_Disposition, Aerenchyma_Formation_Disposition, and Anaerobic_Metabolism_Disposition. V. riparia and V. rupestris (riparian origins) generally show superior tolerance. Waterlogging tolerance is largely independent of drought tolerance, reflecting different adaptive origins.
Hypoxie-Toleranz
Staunässe-Toleranz-Disposition
excess water tolerance
flooding tolerance
hypoxia tolerance
Überflutungs-Toleranz
Kozlowski TT (1997) Responses of woody plants to flooding and salinity, Tree Physiology Monograph 1:1-29 Dat JF et al. (2004) Sensing and signalling during plant flooding, Plant Physiology and Biochemistry 42:273-282 Striegler RK et al. (1993) Influence of rootstock on the response of grapevines to flooding stress, American Journal of Enology and Viticulture 44:313-319
Waterlogging Tolerance Disposition
SO4 (V. berlandieri × V. riparia) demonstrates moderate-to-good waterlogging tolerance, surviving winter flooding of 3-4 weeks duration on heavy clay soils of the Loire Valley alluvial plains, supporting Cabernet Franc in Bourgueil and Chinon where periodic flooding occurs. Dog Ridge (V. champinii) shows substantial waterlogging tolerance derived from adaptation to seasonally flooded Texas river bottomlands, maintaining Flame Seedless productivity on poorly drained clay soils of the southern San Joaquin Valley. 1103 Paulsen (V. berlandieri × V. rupestris) demonstrates moderate waterlogging tolerance, sustaining Primitivo on the seasonally waterlogged clay plains of the Salento peninsula, Puglia, where winter rainfall creates temporary flooding.
An abiotic stress-tolerant rootstock role that inheres in a grafted rootstock by virtue of its capacity to maintain scion productivity under conditions of soil waterlogging, flooding, or impeded drainage that create root zone hypoxia or anoxia. Waterlogging-tolerant rootstocks sustain root function during oxygen deprivation through metabolic adaptations including enhanced glycolytic and fermentative capacity (ethanol and lactate production), formation of aerenchyma tissue providing internal oxygen transport, development of adventitious roots above the saturated zone, and tolerance of toxic reduced compounds (Fe²⁺, Mn²⁺, H₂S) accumulating in anaerobic soils. Tolerance is quantified by vine survival and productivity following flooding events of defined duration and timing, with tolerant rootstocks surviving 2-4 weeks of continuous flooding during the dormant season and 5-10 days during active growth, whereas sensitive rootstocks show mortality after 1-2 weeks dormant flooding or 2-3 days growing-season flooding.
Waterlogging_Tolerant_Rootstock_Role addresses an abiotic stress of increasing relevance under climate change scenarios with more frequent extreme precipitation events. Waterlogging tolerance often correlates with Phytophthora resistance because both stresses occur under wet soil conditions and Phytophthora zoospore activity increases with soil saturation. V. champinii and V. riparia parentage contribute waterlogging tolerance through different mechanisms (anatomical versus metabolic adaptations). Winter dormant-season flooding is generally better tolerated than growing-season flooding, when oxygen demand is higher and root damage more immediately impacts scion function.
flooding tolerant rootstock
hypoxia tolerant rootstock
nässetolerante Unterlage
staunässetolerante Unterlage
waterlogging tolerant rootstock
wet soil tolerant rootstock
überflutungstolerante Unterlage
Striegler RK, Howell GS (1991) The influence of rootstock on the cold hardiness of Seyval grapevines, Vitis 30: 1-11 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 330-335 Dry N (2007) Grapevine rootstocks: selection and management for South Australian vineyards, Lythrum Press Smithyman RP et al. (2001) Influence of rootstock on vine water relations and water use efficiency, American Journal of Enology and Viticulture 52: 324-330
Waterlogging Tolerant Rootstock Role
Cabernet Sauvignon waxy cuticle layer showing heavy bloom characteristics= total wax content 165-210 μg/cm² with well-developed epicuticular crystalloids 1.2-1.8 μm height visible by SEM as platelet structures creating prominent whitish bloom on dark purple-black berry surface easily visible to naked eye, providing water contact angle 110-118° (very hydrophobic), water vapor permeability 0.95×10⁻⁹ m s⁻¹ (low), transpiration 1.0 g H₂O m⁻² h⁻¹, with heavy bloom considered characteristic quality feature for this cultivar indicating proper berry development and environmental exposure. Concord table grape waxy cuticle layer showing extreme bloom= total wax 210-280 μg/cm² (among highest in Vitis) creating very heavy whitish-blue bloom on dark purple-black berries, with bloom considered essential quality characteristic for Concord market identity and consumer expectation, though bloom easily rubbed off during handling requiring gentle harvest and packing to maintain bloom appearance for retail presentation. Syrah waxy cuticle layer sun exposure effects= exposed berries showing 185-235 μg/cm² wax, prominent crystalloid structures, heavy visible bloom, contact angle 112-120° shaded berries showing 110-155 μg/cm² wax, less developed crystalloids, lighter bloom, contact angle 100-108°, demonstrating 45-70% wax content increase in exposed versus shaded berries with sunlight strongly inducing wax biosynthesis and deposition creating enhanced waxy cuticle layer in sun-exposed fruit. Merlot waxy cuticle layer cluster position variation= cluster exterior shoulder berries (maximum sun exposure) showing 195 μg/cm² wax, very heavy bloom cluster middle berries (moderate exposure) showing 145 μg/cm² wax, moderate bloom cluster interior tip berries (minimal exposure) showing 105 μg/cm² wax, light bloom, creating within-cluster gradient in waxy cuticle layer development from heavy exterior to light interior with bloom heterogeneity visible on harvested clusters. Thompson Seedless table grape bloom management= cultivar naturally producing moderate bloom (120-165 μg/cm² wax), with gentle machine harvesting maintaining 70-85% of original bloom creating acceptable appearance, versus rough handling removing 40-60% of bloom exposing green-yellow berry surface creating unattractive appearance for fresh market, demonstrating practical importance of waxy cuticle layer preservation during table grape harvest and handling for maintaining visual quality.
A waxy cuticle layer is a cuticle layer with substantial epicuticular wax deposition creating prominent waxy bloom appearance on berry surface, typically showing wax content >130-150 μg/cm² berry surface area with well-developed epicuticular wax crystalloid structures projecting 0.8-2 μm from surface, providing enhanced hydrophobicity (water contact angle typically >105-110°), reduced water permeability, and characteristic whitish coating particularly visible on dark-skinned cultivars where contrast with anthocyanin pigmentation creates distinct bloom. Waxy cuticle layer characteristics= (1) high total wax content typically 150-250+ μg/cm² (versus 50-120 μg/cm² in less waxy cuticles), with very-long-chain aliphatic compounds predominant (alkanes C29-C33, primary alcohols C26-C32) (2) prominent epicuticular wax crystalloids visible by scanning electron microscopy as platelet, tubular, or rod-shaped structures projecting from cuticle surface creating three-dimensional wax layer (3) thick cuticle overall typically 4.5-8 μm total thickness (versus 2.5-5 μm in less waxy cuticles) reflecting both cutin matrix thickness and abundant wax deposition (4) enhanced hydrophobicity with water contact angle 105-120° creating very water-repellent surface (5) visible waxy bloom appearance to naked eye as whitish, grayish, or bluish coating on berry surface, easily rubbed off with handling revealing underlying berry skin color. Waxy cuticle layer barrier enhancement= (1) reduced water vapor permeability typically 0.8-1.2×10⁻⁹ m s⁻¹ (versus 1.3-2×10⁻⁹ for less waxy cuticles) limiting berry transpiration to 0.8-1.3 g H₂O m⁻² h⁻¹ (2) enhanced resistance to pathogen penetration as thick wax layer requires cutinase enzyme production by fungi for penetration creating additional barrier (3) reduced wettability limiting water film formation on berry surface reducing favorable conditions for pathogen spore germination (4) potential UV protection through light scattering by wax crystalloid structures. Waxy cuticle layer development shows cultivar and environmental influences= genetic component with certain cultivars consistently producing heavy bloom (Cabernet Sauvignon, Syrah, Concord typically waxy) versus light bloom (some table grape cultivars, Pinot Noir moderate) sunlight exposure enhancing wax deposition through UV-B induction of wax biosynthetic genes with exposed berries accumulating 30-60% more wax than shaded berries water deficit potentially enhancing wax deposition as stress response developmental progression with wax accumulation continuing through ripening achieving maximum at maturity. Waxy cuticle layer in viticulture and wine production= bloom appearance as visual quality indicator for table grapes (uniform heavy bloom considered desirable for market appearance) bloom as indicator of berry handling (bloom easily rubbed off revealing whether berries handled gently) bloom influence on wine production minimal as waxes insoluble in aqueous or alcoholic media and not extracted during winemaking bloom removal through washing or handling exposing underlying skin affecting berry appearance and potentially water loss rates. Waxy cuticle layer variation among berry positions= cluster exterior berries often showing heavier bloom (higher wax content 180-240 μg/cm²) due to greater sun exposure and environmental stress cluster interior shaded berries showing lighter bloom (100-150 μg/cm²) due to reduced sun exposure and more protected environment within-cluster bloom variation creating visual heterogeneity in table grapes potentially affecting market appearance.
Waxy_Cuticle_Layer represents cuticles with heavy wax deposition (>130-150 μg/cm²) creating prominent bloom and is distinguished from less waxy cuticles and from Protective_Cuticle_Layer (which emphasizes barrier function over wax appearance). Waxy cuticle layer provides enhanced barrier properties and characteristic bloom appearance important for some table grape markets. Understanding waxy cuticle layer development and environmental modulation is relevant for managing berry quality through canopy management affecting sun exposure, and for table grape handling protocols preserving bloom appearance.
Blüte-Schicht
Starker Wachsüberzug
bloom layer
heavy wax coating
wachsige Cuticula-Schicht
Rogiers, S.Y. et al. (2004). Grape berry epicuticular wax and transpiration. American Journal of Enology and Viticulture 55=121-127 Rosenquist, J.K., Morrison, J.C. (1989). Cuticle and wax accumulation. American Journal of Enology and Viticulture 40=241-244.
Waxy Cuticle Layer
Waxy surface quality assessment in premium table grape production for Autumn Royal evaluates bloom integrity as a freshness indicator, with intact heavy bloom (contact angle above 120°, complete surface dulling) commanding premium prices because it demonstrates minimal handling from harvest through cold chain fruit showing bloom loss from mechanical contact exhibits shiny patches (contact angle below 90°) that signal rough handling and correlate with reduced storage life, making bloom preservation a primary objective in harvest logistics that mandates single-layer packaging, minimal transfer points, and foam-cushioned containers.
Waxy_Surface_Quality represents the surface texture condition where a grapevine organ is covered with epicuticular wax deposits that create the characteristic bloom or pruina visible on grape berries and certain leaf surfaces. The waxy bloom is a complex mixture of triterpenoids, primarily oleanolic acid, with smaller amounts of fatty alcohols, aldehydes, and alkanes that form crystalline microstructures on the cuticle surface. These wax crystals scatter light, producing the dusty, matte, or frosted appearance that distinguishes bloom-covered surfaces from the shiny appearance of wax-free surfaces. In grape berries, the bloom is a conspicuous quality character that develops progressively during berry growth, reaching maximum coverage during ripening, and serves multiple functional roles including reducing water loss, providing a hydrophobic barrier against rain and dew accumulation, limiting UV penetration, and potentially deterring certain pathogens through surface hydrophobicity. The intensity of bloom development varies among cultivars, with some varieties producing heavy bloom that completely dulls the surface while others develop lighter bloom revealing underlying skin color. Environmental factors including humidity, light exposure, and temperature modulate bloom development. Handling and abrasion readily remove bloom, making its presence an indicator of careful fruit handling in table grape marketing. Bloom assessment employs visual evaluation of the degree of surface dulling, water droplet contact angle measurement indicating surface hydrophobicity, and chemical quantification of epicuticular wax content.
Waxy_Surface_Quality specifically addresses the epicuticular wax layer (bloom/pruina) that characterizes berry surfaces and some leaf surfaces. The class name includes the parenthetical (bloom/pruina) following the uploaded structure. Bloom is distinct from the structural cuticle beneath it the cuticle proper provides the mechanical barrier while the epicuticular wax crystals provide the optical and hydrophobic properties. Bloom thickness and composition vary with cultivar, developmental stage, and environment. The term pruina derives from Latin for frost and describes the frosted appearance. OIV descriptors do not directly assess bloom intensity, though it contributes to perceived berry color intensity.
Bereifung
Pruina-Bedeckung
Wachsschichtqualität
bloom quality
epicuticular wax texture
pruina coverage
wachsige Oberflächenbeschaffenheit
waxy bloom character
Waxy Surface Quality
Characterization of Weak_Apical_Dominance_Disposition at the University of Bordeaux documented intermediate lateral development in cultivars with moderate branching tendency. Reference cultivars 'Cabernet Sauvignon', 'Cabernet Franc', 'Sauvignon Blanc', and 'Pinot Noir' were assessed under standardized conditions without lateral management. Weak_Apical_Dominance_Disposition cultivars showed consistent intermediate characteristics. 'Cabernet Sauvignon' exhibited typical weak apical dominance with mean of 7.8 lateral shoots per primary shoot of 175 cm, distributed across basal (40%), mid-shoot (45%), and distal (15%) nodes, with total lateral length of 95 cm (ratio 0.54). Lateral shoots averaged 12 cm length with some exceeding 25 cm, indicating incomplete but not absent suppression. 'Pinot Noir' showed similar patterns (6.5 laterals, 78 cm total) with notable node-position effect—laterals at nodes 8-12 were significantly longer than at basal or distal positions, suggesting a zone of reduced apical influence at intermediate distance from apex. Apex auxin concentration in Weak_Apical_Dominance_Disposition cultivars averaged 195 ng/g FW (near cultivar mean), intermediate between strong dominance (285 ng/g) and proliferation types (145 ng/g). Management trials confirmed that Weak_Apical_Dominance_Disposition cultivars benefited from targeted lateral management—removal of laterals at fruiting zone nodes improved cluster exposure without requiring the intensive whole-shoot lateral removal needed for proliferation types. Labor requirement averaged 45 hours/ha, intermediate between strong dominance (25 hours) and proliferation types (85 hours).
A Weak_Apical_Dominance_Disposition is a Branching_Pattern_Disposition that inheres in a Vitis vinifera plant and is realized through limited hormonal suppression of lateral bud outgrowth, resulting in moderate lateral shoot production from primary shoot nodes even while the apex remains intact and actively growing. This disposition represents intermediate correlative inhibition, characteristic of cultivars where the auxin signal from the shoot apex is insufficient to fully suppress lateral bud activation, particularly at nodes distant from the apex or under favorable growing conditions. Cultivars with Weak_Apical_Dominance_Disposition typically produce 6-10 lateral shoots per primary shoot, with laterals developing at both basal and mid-shoot positions while the apex continues primary growth. The viticultural implications are intermediate—more lateral management is required than for strong apical dominance types, but the naturally moderate branching does not create the extremely dense canopies characteristic of lateral proliferation types. Weak_Apical_Dominance_Disposition provides flexibility, as lateral development can be allowed to increase leaf area where desired or removed where canopy density is excessive. The disposition is associated with lower apex auxin concentrations and/or reduced auxin transport efficiency, allowing cytokinin-driven bud activation to proceed at a moderate level.
Weak_Apical_Dominance_Disposition is_a Branching_Pattern_Disposition representing moderate lateral development (6-10 laterals per primary shoot). Characterized by intermediate apex auxin and partial correlative inhibition. Examples include 'Cabernet Sauvignon', 'Cabernet Franc', 'Sauvignon Blanc', 'Pinot Noir', 'Chardonnay'. Laterals develop at basal and mid-shoot positions while apex remains active. Moderate management flexibility—laterals can be retained or removed as needed. Represents the most common branching pattern among commercial cultivars.
Intermediäre-Verzweigung
Mäßige-Apikalkontrolle
Schwache-Apikaldominanz-Disposition
intermediate branching
moderate apical control
partial lateral suppression
Lebon E et al. (2006) Shoot development in grapevine (Vitis vinifera L.) is affected by the modular branching pattern of the stem and intra- and inter-shoot trophic competition, Annals of Botany 98:1045-1056 Bennett T, Leyser O (2006) Something on the side: axillary meristems and plant development, Plant Molecular Biology 60:843-854 Dry PR (2000) Canopy management for fruitfulness, Australian Journal of Grape and Wine Research 6:109-115
Weak Apical Dominance Disposition
A weak primary vegetative shoot on Vitis vinifera cv. Riesling from node 2 in vineyard showing widespread weak growth symptoms, reached only 52cm length by late July with growth essentially ceased, 8 small leaves averaging 75 cm² (total only 600 cm² leaf area), 5mm diameter showing minimal secondary growth, chlorotic appearance suggesting nutritional deficiency or disease stress, representing problematic weak development investigated and attributed to combination of severe drought stress (no irrigation, rainfall 40% below normal), possible root disease (Armillaria suspected based on adjacent vine mortality), and nutrient deficiency (soil tests revealing severe potassium and magnesium depletion), triggering remedial interventions including supplemental irrigation, fertilization, and vineyard health assessment program.
A primary vegetative shoot exhibiting reduced growth characteristics, characterized by slow elongation (1-2cm per day or less), limited final length (typically 40-80cm), short internodes (3-6cm spacing), thin diameter (4-6mm at base), reduced leaf development (6-12 small leaves, individual areas 60-120 cm²), and minimal photosynthetic contribution, indicating vine stress, resource limitation, disease effects, or unfavorable growing conditions constraining normal shoot development despite primary bud origin. Weak primary vegetative shoots represent suboptimal vegetative expression, signaling problems requiring diagnosis and correction including inadequate water or nutrients, disease pressure (root diseases, trunk diseases, foliar pathogens), environmental stress (extreme temperatures, insufficient light), vine decline, or inappropriate cultural practices, with weak shoot development threatening current productivity and long-term vine sustainability.
Weak primary vegetative shoots serve as important diagnostic indicators, with their reduced growth signaling problems affecting whole-vine function and requiring investigation and correction. Causes include water stress (drought, irrigation system failure, poor soil water holding), nutrient deficiencies (nitrogen, potassium, magnesium, iron), diseases (Phytophthora root rot, Armillaria root disease, Eutypa dieback, virus infections), environmental extremes (heat stress, excessive cold, hail damage, herbicide drift), or vine decline syndromes (age-related senescence, cumulative stress). Management responses depend on cause diagnosis, ranging from irrigation supplementation (water stress) to fertilization (nutrient deficiency) to disease management (pathogen identification and control) to possible vine removal and replanting (severe decline, root disease). Weak vegetative shoots provide minimal photosynthetic contribution and unsuitable renewal wood, typically removed during canopy management to redirect limited resources toward stronger shoots and crop development.
kümmernder Blatttrieb
low-vigor non-fruiting shoot
schwacher primärer vegetativer Trieb
stunted primary vegetative shoot
Based on Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 113-145 Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide Pearson & Goheen (1988) "Compendium of Grape Diseases" APS Press, pp. 1-93.
Weak Primary Vegetative Shoot
A weak secondary vegetative shoot on Vitis vinifera cv. Pinot Noir from node 6, activated following severe spring frost (-4.1°C) with 94% primary bud mortality, achieved only 58cm length by late July with minimal subsequent growth, 8 small chlorotic leaves averaging 72 cm² totaling only 576 cm² leaf area, 4.5mm diameter showing poor lignification, representing severely compromised secondary development attributed to combination of catastrophic frost damage depleting vine reserves, secondary activation from nearly all positions creating excessive competition for limited resources, inadequate remaining crop (5% normal) providing insufficient sink demand for resource mobilization, and cool poor growing season following frost limiting photosynthesis and growth, this shoot providing minimal photosynthetic contribution and unsuitable for any productive use, retained only to maintain minimal canopy presence supporting basic vine survival and gradual reserve rebuilding for following season recovery.
A secondary vegetative shoot exhibiting reduced growth despite activation, characterized by limited elongation achieving only 40-75cm length, sparse leaf development with 6-10 small leaves providing 500-1100 cm² photosynthetic area, thin diameter (4-6mm at base), and minimal growth performance reflecting either inherent secondary weakness, unfavorable post-damage conditions, excessive remaining crop stress, or compound effects of severe damage plus inadequate recovery resources. Weak secondary vegetative shoots demonstrate minimal compensation from backup positions, providing limited photosynthetic replacement following primary loss, contributing marginally to canopy recovery, and indicating problematic recovery circumstances requiring assessment and potential intervention including crop reduction, supplemental irrigation or nutrition, or acceptance of poor vintage outcomes recognizing inadequate compensation capacity given damage severity and vine conditions.
Weak secondary vegetative shoots indicate compromised recovery following damage, with poor secondary development signaling inadequate compensation capacity threatening current season outcomes and potentially affecting subsequent seasons through poor reserve accumulation and inadequate bud differentiation. Causes include severe widespread damage (>80% primary loss creating overwhelming compensation demand), poor post-damage conditions (cool temperatures, drought, nutrient limitation), excessive remaining crop (partial primary survival maintaining sink demand beyond weak secondary capacity), or pre-existing vine stress (disease, previous-season stress, inadequate reserves). Management responses include realistic expectations (accepting poor vintage), possible crop reduction (removing remaining crop to support vegetative recovery), supplemental inputs (irrigation, foliar nutrition), and planning for multi-year recovery recognizing single-season compensation impossibility. Weak secondary shoots provide minimal current benefits and unsuitable renewal wood, typically removed during canopy management unless no stronger shoots available for minimal canopy maintenance.
kümmernder Ersatztrieb
poor compensatory shoot
schwacher sekundärer vegetativer Trieb
stunted secondary vegetative shoot
Weak Secondary Vegetative Shoot
Pinot Noir displays relatively weak tendril disposition with tendril elongation rates of 4-6 cm/day, delayed coiling response (2-4 hours post-contact), and attachment tensile strength of 5-10 N. Table grape cultivars selected for ease of harvest such as 'Thompson Seedless' often exhibit weak tendril traits, facilitating bunch removal. Some dwarfing rootstocks reduce scion tendril vigor by 20-35% compared to vigorous rootstocks.
A tendril development disposition that is the inherited capacity of a Vitis vinifera plant to produce tendrils with reduced biomass, slower elongation rate, delayed or diminished coiling response, and lower attachment strength, resulting in limited natural climbing ability.
Weak tendril disposition may be advantageous in highly managed vineyard systems where tendrils can interfere with mechanical harvesting or require additional labor for removal. The trait shows moderate heritability (h² ≈ 0.45) and correlates weakly with overall plant vigor (r = 0.38), indicating partial genetic independence. Reduced tendril strength in some cultivars may reflect selection pressure during domestication for ease of training.
geringe Rankenkraft
low tendril vigor
reduced tendril formation
reduzierte Rankenentwicklung
schwache Rankenbildung
weak climbing capacity
This OD, Martinez-Zapater JM (2016) Genetic and genomic approaches for adaptation of grapevine to climate change. In: Gerós H et al. (eds) Grapevine in a Changing Environment, Wiley pp.: 157-178 Dry PR et al. (2010) Canopy management for fruitfulness. Australian Journal of Grape and Wine Research 16:25-35
Weak Tendril Disposition
A weakly attached functional tendril at node 12 on a late-season shoot of Vitis vinifera cv. Viognier in the northern Rhône Valley, France, exhibiting three loose contact coils around a 10 mm diameter stake with coil diameters of 45-50 mm, showing only light tan coloration indicating minimal lignification five weeks after initial contact, supporting a 40 cm distal shoot section bearing four leaves totaling approximately 280 grams weight, showing visible coil slippage of 3-4 cm along the stake when shoot is moved by moderate wind, detaching completely during a thunderstorm with wind gusts of 55 km/h, measured at BBCH stage 85 (softening of berries) in late August, representing failed attachment that necessitated manual shoot repositioning and securing with twine to maintain vertical shoot positioning through harvest.
An attached functional tendril exhibiting marginal grip strength and limited mechanical stability in Vitis vinifera, characterized by loose coiling with contact coil diameters substantially exceeding optimal values relative to support structure diameter, incomplete or minimal lignification leaving tissues partially flexible and susceptible to deformation, few contact points generating low friction forces that may slip under loading, and demonstrated load-bearing capacity barely sufficient for current shoot weights, typically supporting 200-500 grams maximum before visible strain or risk of detachment. Weakly attached functional tendrils develop through suboptimal attachment conditions including interrupted or brief mechanosensory stimulation preventing complete coiling response, support structures with inappropriate dimensions either too thick for effective grasping or too thin creating minimal contact area, environmental stress during the critical attachment period including water deficit or temperature extremes limiting cell expansion and wall reinforcement, mechanical disturbances during coiling that disrupt optimal contact geometry, or late-season attachment when declining vine resources limit structural development. These tendrils provide minimal mechanical support, often sufficient only in calm conditions with light shoot weights, and represent vulnerable attachment points susceptible to failure during wind events, heavy rain, or fruit cluster development that increases mechanical loading. Weakly attached tendrils may undergo progressive deterioration through coil slippage, partial detachment, or tissue degradation, potentially transitioning to completely detached status during the growing season.
Weakly attached functional tendrils are distinguished from firmly attached tendrils by quantitative grip strength measurements (resistance to minimal detachment forces typically <500 grams), qualitative coiling assessment (loose geometry, incomplete lignification, visible tissue flexibility), and demonstrated performance failures under normal loading conditions. Weak attachment often correlates with late-season tendril development when declining carbohydrate availability limits structural investment, with tendril position on weak or stressed shoots, or with environmental conditions unfavorable for lignification. Management implications include recognition that vines relying primarily on weakly attached tendrils require supplemental support through positioning, tying, or training system adjustments to prevent canopy collapse. Weak attachments serve as indicators of vine stress conditions requiring diagnostic attention.
geringe-festigkeit Greifranke
insecure grip tendril
locker befestigte Ranke
low-strength attachment tendril
poorly attached tendril
schwach verankerte Ranke
Based on Jaffe & Galston (1968) "The physiology of tendrils" Annual Review of Plant Physiology 19=417-434 Silk & Hubbard (1991) "Axial forces and normal stresses in coiling tendrils" Biomechanics and Modeling in Mechanobiology 24=599-606 Darwin (1875) "The Movements and Habits of Climbing Plants" John Murray, London Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 219-236.
Weakly Attached Functional Tendril
Weather dependency quality assessment for Pinot Noir across Burgundy and Oregon evaluated the constraints imposed by autumn weather patterns on harvest scheduling. Burgundy Pinot Noir demonstrated high weather dependency quality with thin skin susceptibility to rain-induced splitting and Botrytis proliferation creating strong requirement for dry harvest conditions, with quality tier reduction averaging 15% for fruit harvested during or within 24 hours of precipitation events compared to dry-harvested fruit from equivalent vineyard sources. Oregon showed even higher weather dependency quality due to more frequent October precipitation, with quality variance between dry and wet harvest conditions exceeding 20% and leading to widespread adoption of covered canopy systems and weather-responsive harvest protocols. Comparative assessment of Cabernet Sauvignon showed substantially lower weather dependency quality with thick skin resisting splitting, loose clusters drying rapidly after rain, and robust tannin structure masking subtle quality effects from suboptimal harvest conditions, permitting more flexible scheduling that tolerated occasional rain events without significant quality penalty.
Weather dependency quality is a harvest condition quality that inheres in a grapevine cultivar or clone and represents the degree to which harvest operations and resulting fruit quality are constrained by meteorological conditions including precipitation, humidity, temperature extremes, and wind, determining the weather windows required for successful harvest and the consequences of proceeding under suboptimal conditions when scheduling pressures or impending weather deterioration necessitate compromise. This quality captures the practical reality that vintage operations must accommodate weather variability that cannot be controlled, with cultivars exhibiting high weather dependency requiring favorable conditions that may not coincide with optimal ripeness timing, while those with low dependency permit harvest across a broader range of conditions providing operational flexibility. The ontological significance of weather dependency quality lies in its role as a risk factor for vintage operations, wherein high-dependency cultivars concentrate harvest risk into brief favorable weather windows where delays can result in either quality loss from unfavorable harvest conditions or from waiting beyond optimal ripeness for weather improvement, creating difficult tradeoffs that low-dependency cultivars avoid through their tolerance for diverse conditions. The biological basis of weather dependency encompasses multiple condition-quality interactions including the effects of precipitation on berry dilution, surface moisture promoting pathogen development, and physical damage to clusters from hail or heavy rain temperature effects on fruit metabolism, oxidation rates, and aromatic compound stability during harvest humidity influences on surface moisture retention, disease pressure, and evaporative cooling during transport and wind effects on drying conditions, physical damage potential, and harvest operation efficiency. Cultivar characteristics affecting weather dependency include skin thickness influencing rain-induced berry splitting and dilution susceptibility, cluster architecture determining moisture retention and drying rate, disease resistance affecting tolerance for damp conditions, and compositional sensitivity to temperature during the harvest-to-processing interval. Assessment of weather dependency quality involves documentation of quality outcomes across varied harvest weather conditions, identification of condition thresholds beyond which quality compromise occurs, and development of weather-based harvest decision protocols tailored to specific cultivar requirements.
Weather_Dependency_Quality is_a Harvest_Condition_Quality, representing the sensitivity of harvest success to meteorological conditions. This class addresses the practical constraints imposed by weather on vintage operations. Precipitation effects through dilution and disease, temperature effects on metabolism and oxidation, and humidity effects on drying and spoilage all contribute. Distinguished from Morning_Harvest_Suitability_Quality and Night_Harvest_Suitability_Quality which address diurnal timing rather than weather conditions per se. Critical for harvest risk assessment and contingency planning.
Erntewetterempfindlichkeit
Wetterabhängigkeit
Witterungsabhängigkeit bei der Ernte
climate dependency during harvest
harvest weather sensitivity
meteorological harvest constraints
meteorologische Lesebeschränkungen
weather window requirements
van Leeuwen C, Darriet P (2016) The impact of climate change on viticulture and wine quality. Journal of Wine Economics 11:150-167 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 509-547 Gladstones J (1992) Viticulture and Environment. Winetitles, Adelaide, pp. 112-145
Weather Dependency Quality
Merlot exhibits pronounced weather-induced coulure disposition with fruit set declining from 35% at 25°C to only 12% when mean flowering temperature drops to 14°C. Gewürztraminer shows 50-60% flower loss during rainy bloom periods exceeding 5 consecutive days with precipitation. Cold-sensitive pollen tube growth (optimum 25-30°C) underlies weather-induced coulure, with elongation rates falling below the threshold for timely fertilization at temperatures under 15°C. Wind speeds exceeding 25 km/h during anthesis reduce fruit set by 15-25% through pollen dispersal and stigma desiccation.
A coulure susceptibility disposition that is the inherited capacity of a Vitis vinifera plant to exhibit flower or young berry abscission in response to adverse weather conditions during the flowering period, including low temperatures, excessive rainfall, high humidity, or persistent cloudiness that impair pollination and fertilization.
Weather-induced coulure results from direct effects on gametophyte function (pollen germination and tube growth) and indirect effects through reduced photosynthesis and carbohydrate supply to developing flowers. Rain during anthesis physically removes pollen from stigmas and dilutes stigmatic exudate. Cultivar differences in weather-induced coulure susceptibility correlate with pollen germination temperature optima and carpel development rate.
climate-induced shatter
klimainduzierter Blütenabwurf
temperaturbedingte Durchrieselung
temperature-sensitive coulure
weather-dependent poor set
witterungsbedingte Verrieselung
Ebadi A et al. (1996) Effect of short-term temperature and shading on fruit-set, seed and berry development in model vines of V. vinifera cvs Chardonnay and Shiraz. Australian Journal of Grape and Wine Research 2:2-9 Staudt G (1982) Pollen germination and pollen tube growth in vivo in Vitis and the dependence on temperature. Vitis 21:205-216
Weather Induced Coulure Disposition
In Riesling vines in the Rheingau, weather responsive dormancy during February and March creates year-to-year budbreak timing variation of 2-3 weeks depending on spring temperature patterns, with warm springs providing rapid heat accumulation and early budbreak in late March, cool springs delaying heat accumulation and budbreak until mid-April, and this weather responsiveness quantified through growing degree day tracking that enables budbreak prediction within approximately 5 days accuracy when initiated from late February.
Weather responsive dormancy in Vitis vinifera represents the dynamic relationship between bud status and ambient weather conditions during ecodormancy, whereby bud quiescence or activity responds continuously to prevailing meteorological conditions with growth activity increasing during favorable weather and decreasing during unfavorable weather. This responsiveness creates the characteristic weather-dependent variability in budbreak timing and progression observed during late winter and early spring, contrasting with the weather-independent dormancy of endodormancy where bud status is determined by internal factors regardless of external conditions. The physiological basis of weather responsiveness involves the absence of buffering internal regulatory mechanisms, leaving bud physiology directly coupled to environmental temperature through the fundamental temperature-dependence of metabolic reaction rates, with warming accelerating growth-related processes and cooling decelerating them in direct proportion to temperature effects on enzyme activities and cellular processes. The time-scale of weather responsiveness ranges from immediate biochemical responses to temperature changes affecting metabolic rates, through short-term responses over days affecting gene expression and protein accumulation, to longer-term developmental responses over weeks determining progress toward budbreak, with all these timescales contributing to the overall relationship between weather and bud status. The practical consequences of weather responsive dormancy for viticulture include the difficulty of predicting budbreak timing more than a few weeks in advance due to dependence on future weather patterns, the need for flexible frost protection strategies that can respond to weather-driven variation in bud developmental stage, and the potential for weather extremes to advance or delay budbreak outside normal timing windows. The quantitative relationship between weather and dormancy status is captured by growing degree day models that integrate temperature exposure over time, with accumulated heat units providing a measure of progress toward budbreak that accounts for the varying intensity of growth-promoting conditions experienced during different weather patterns. The transition from weather responsive dormancy to active growth represents the point where accumulated weather-driven progress has satisfied the thermal requirement for budbreak, typically occurring after 100-300 growing degree days have accumulated above the base temperature following completion of chilling requirement.
Weather_Responsive_Dormancy represents the dynamic weather-bud relationship within Conditional_Dormancy. Distinguished from weather-independent endodormancy status. Creates year-to-year budbreak timing variability of 2-4 weeks typically. Quantified through growing degree day accumulation models. Complicates budbreak prediction beyond short-term weather forecasts. Response timescales range from immediate metabolic effects to developmental progress over weeks. Base temperature typically 10°C with requirements of 100-300 GDD for budbreak.
climate-responsive dormancy
klimaabhängige Dormanz
meteorologically responsive quiescence
meteorologisch responsive Ruhe
weather-driven dormancy
wetterabhängige Dormanz
wettergesteuerte Dormanz
Parker AK et al (2011) General phenological model to characterise the timing of flowering and veraison of Vitis vinifera L. Australian Journal of Grape and Wine Research 17: 206-216 Keller M (2020) The Science of Grapevines 3rd Ed Chapter 2 Phenology and Growth Cycle Moncur MW et al (1989) Base temperatures for budbreak and leaf appearance of grapevines. American Journal of Enology and Viticulture 40: 21-26
Weather Responsive Dormancy
Trollinger clone 1 We bears a Weinsberg_Clone_Role supporting Württemberg's signature red variety, selected for local conditions and the light, fresh wine style traditional to the region—demonstrating Weinsberg's focus on regional variety needs. Lemberger (Blaufränkisch) clone 1 We serves in a Weinsberg_Clone_Role for Württemberg's important Lemberger production, with selection emphasizing the structured, age-worthy style valued in the region. Schwarzriesling (Pinot Meunier) clone 1 We bears a Weinsberg_Clone_Role for this regionally significant variety, with Württemberg representing Germany's primary Schwarzriesling production zone and Weinsberg providing adapted clone selections.
A German clone role that inheres specifically in clones selected and registered by the Staatliche Lehr- und Versuchsanstalt für Wein- und Obstbau Weinsberg (LVWO, State Institute for Viticulture and Pomology Weinsberg), located in Württemberg (southwestern Germany), with selections designated by the "We" suffix (e.g., Trollinger 1 We, Lemberger 1 We). Weinsberg clones reflect the institution's focus on Württemberg's distinctive variety portfolio, including regional specialties (Trollinger, Lemberger/Blaufränkisch, Schwarzriesling/Pinot Meunier) that are less emphasized by other German selection programs. LVWO Weinsberg combines clone selection with variety breeding, having developed new varieties (Dornfelder, Regent, Acolon) that have achieved national and international importance alongside its clone evaluation work. The institution serves Württemberg's substantial wine industry (Germany's fourth-largest wine region) while contributing regional variety clones to the national registry. Weinsberg's dual focus on traditional regional varieties and innovative new variety development distinguishes its program within the German institutional landscape.
Weinsberg_Clone_Role identifies LVWO selections with particular significance for Württemberg's distinctive variety portfolio. The region's relative isolation within German wine geography (separated from other major regions by terrain and tradition) created demand for locally adapted selections of varieties with limited importance elsewhere, which Weinsberg addresses through its regional focus. The institution's variety breeding success (Dornfelder becoming Germany's second most-planted red variety, Regent pioneering practical disease resistance) complements clone selection, with Weinsberg contributing both refined selections of existing varieties and entirely new variety options. The combination of clone selection, variety breeding, and educational mission creates an institution addressing multiple dimensions of regional viticultural advancement.
LVWO clone
LVWO-Klon
We clone
We-Klon
Weinsberg clone
Weinsberg selection
Weinsberg-Klon
Weinsberger Selektion
Württemberg clone
Württemberg-Klon
Staatliche Lehr- und Versuchsanstalt für Wein- und Obstbau Weinsberg (2020) Rebenzüchtung und Klonenselektion Bundessortenamt (2020) Beschreibende Sortenliste Reben Becker H (1985) German viticulture and breeding, American Journal of Enology and Viticulture 36: 58-63 Eibach R, Töpfer R (2015) Traditional grapevine breeding techniques, in Reynolds A (ed) Grapevine Breeding Programs for the Wine Industry, Elsevier pp.: 3-22
Weinsberg Clone Role
Sauvignon Blanc grafted on SO4 rootstock at 14 years post-grafting showing well established mature grafted rootstock trunk 6.2 cm diameter, trunk extending 25 cm from root crown to graft union positioned 20 cm above soil, graft union displaying stable 1.3 cm diameter bulge completely integrated with mature bark obscuring original graft line, trunk bark 2.8 mm thickness showing characteristic SO4 moderate longitudinal fissures and grey-brown coloration, trunk fully self-supporting, diameter growth 0.4 cm current season, 14 clear annual rings in cross-section, no disease or damage, supporting vigorous canopy 3.9 m total shoot length and 6.8 kg fruit load, vine in peak early mature production
Well established mature grafted rootstock trunk is mature grafted rootstock trunk in early to mid mature phase typically years 8-18 post-grafting, characterized by recently achieved full structural and physiological development, stable graft union with no additional callus formation, well-developed bark showing complete rootstock cultivar characteristics, diameter typically 5-8 cm, consistent annual growth of 0.3-0.5 cm, complete self-supporting capacity, and representing early optimal productive phase where trunk has achieved mature status and is providing reliable foundation for sustained high-quality production with potential for decades more productive life ahead. Well established mature grafted rootstock trunk age and phase includes= spanning years 8-18 post-grafting typically (early to mid mature trunk period), representing recently achieved maturity (trunk having transitioned from establishing to mature status within past 3-10 years), showing early prime productive phase (beginning of optimal production period with many decades of productive potential remaining), displaying fully developed but not yet aged characteristics (trunk exhibiting all mature features without old age indicators), and marking vigorous mature stage (trunk at height of structural and physiological capacity before any age-related decline). Well established mature grafted rootstock trunk structural characteristics includes= exhibiting early mature diameter range (5-8 cm typical for well established phase depending on rootstock vigor and cultivar), showing fully developed bark (periderm 2-4 mm thickness with complete expression of rootstock-specific texture and patterns), displaying vigorous bark appearance (bark showing mature characteristics but relatively fresh without extensive weathering or damage accumulation), revealing clear annual ring patterns (8-18 growth rings visible in cross-section showing consistent regular growth), and showing robust substantial structure (trunk appearing solid, healthy, and capable of supporting large productive vine for many years). Well established mature grafted rootstock trunk graft union stability includes= displaying completely integrated graft union (all callus tissue mature with no ongoing callus accumulation), showing stable bulge morphology (graft union swelling 0.8-2.0 cm diameter increase stabilized with no further expansion), exhibiting obscured graft line (thick bark and complete integration making original grafting point difficult to locate visually), demonstrating maximum mechanical integration (graft union strength equivalent to continuous non-grafted trunk), and showing no signs of incompatibility (absence of necrosis, cracking, excessive bulging, or other integration problems confirming successful long-term partnership). Well established mature grafted rootstock trunk growth patterns includes= showing consistent moderate annual increase (0.3-0.5 cm diameter growth typical in well established phase), displaying regular predictable growth (annual increments consistent year to year indicating stable healthy condition), exhibiting active cambial activity (vigorous vascular cambium producing xylem and phloem each growing season), forming clear distinct annual rings (each growing season creating well-defined ring visible in cross-section), and maintaining stable wood properties (consistent wood density, vessel characteristics, and mechanical properties reflecting healthy mature status). Well established mature grafted rootstock trunk vascular function includes= providing optimal water transport capacity (xylem area adequate for 50-100+ liters daily water conduction supporting full productive canopy), enabling complete nutrient distribution (phloem and xylem capacity sufficient for all mineral and photosynthate transport needs), showing maximum transport efficiency (well-developed vascular architecture minimizing resistance and maximizing flow rates), supporting consistent crop production (trunk capacity adequate for sustained annual fruit loads of 5-12+ kg depending on cultivar), and maintaining substantial reserves (significant carbohydrate storage in ray parenchyma supporting resilience and consistent performance). Well established mature grafted rootstock trunk in optimal production includes= supporting peak fruit quality (stable resource transport enabling consistent high-quality grape production), enabling reliable annual yields (adequate trunk capacity for commercial crop loads without stress), showing appropriate vigor balance (trunk size well matched to training system and production objectives), maintaining healthy productive canopy (trunk supporting optimal leaf area and photosynthetic capacity), demonstrating excellent vine balance (trunk capacity supporting equilibrium between growth, production, and reserves), and providing foundation for quality viticulture (robust trunk enabling superior grape production for wine or table use). Well established mature grafted rootstock trunk management advantages includes= requiring minimal special attention (well established trunk needing only standard vineyard maintenance practices), showing excellent disease resistance (mature healthy trunk displaying good resistance to cankers, crown gall, and decay), tolerating environmental stress well (substantial reserves and robust structure supporting recovery from drought, freeze, or heat events), responding predictably to management (consistent trunk performance enabling reliable vineyard planning and management), and providing stable foundation (reliable trunk characteristics year after year supporting consistent vineyard operations and crop quality). Well established mature grafted rootstock trunk rootstock expression includes= conferring full phylloxera protection (rootstock genetics providing complete resistance to root form protecting vine), expressing complete vigor control (rootstock variety fully manifesting characteristic low, moderate, or high vigor effects on scion), providing optimal site adaptation (rootstock traits including drought tolerance, lime tolerance, or nematode resistance fully expressed), showing characteristic bark features (rootstock cultivar-specific bark texture, color, and fissure patterns clearly identifiable), and demonstrating cultivar-specific performance (well established trunk displaying predictable rootstock variety characteristics affecting whole vine behavior). Well established mature grafted rootstock trunk physical appearance includes= displaying fresh mature bark (bark showing full mature characteristics but relatively unweathered and undamaged), showing clear fissure patterns (longitudinal furrows characteristic of rootstock variety well developed but not yet extremely deep or eroded), exhibiting healthy coloration (grey-brown, tan, or reddish tones typical of rootstock without discoloration or extensive weathering), revealing visible lenticels (gas exchange structures prominent on bark surface as horizontal lines or oval formations), and showing clean trunk surface (minimal lichen growth, moss accumulation, or damage compared to older aged trunks). Well established mature grafted rootstock trunk diameter characteristics includes= typical range 5-8 cm for years 8-18 (common dimensions across rootstock varieties in well established phase), vigorous rootstocks showing 6-9 cm (1103P, 110R showing upper range of well established dimensions), moderate vigor rootstocks showing 5-7 cm (SO4, 3309C typical for well established phase), low vigor rootstocks showing 4.5-6.5 cm (Riparia types showing smaller well established trunk), and continuing steady diameter increase (trunk still actively expanding though at stable mature rate). Well established mature grafted rootstock trunk cultivar characteristics includes= SO4 showing 5.5-7 cm diameter with moderately furrowed grey-brown bark, 3309C displaying 5-6.5 cm diameter with fine fissures and light grey bark, Teleki 5C exhibiting 5.5-7.5 cm diameter with moderate bark texture, 1103P showing 7-9 cm diameter with rough deeply furrowed bark, Riparia Gloire displaying 4.5-6 cm diameter with relatively smooth bark, 110 Richter exhibiting 6.5-8.5 cm diameter with very thick rough bark, and 101-14 showing 4.5-6 cm diameter with thin moderately fissured bark. Well established mature grafted rootstock trunk as vineyard asset includes= representing peak investment value (well established trunk marking period of maximum productive capacity relative to establishment costs), providing reliable consistent performance (trunk characteristics stable and predictable year to year supporting vineyard planning), showing excellent health status (minimal disease, damage, or stress indicators in well maintained well established trunk), demonstrating successful establishment (well established trunk confirming first 7-8 years' work succeeded in building solid foundation), and offering decades more productivity (well established trunk with potential for 30-60+ additional years of productive life). Well established mature grafted rootstock trunk monitoring and assessment includes= showing healthy bark condition (minimal cracks, cankers, or damage on trunk surface), displaying vigorous annual growth (consistent 0.3-0.5 cm diameter increase confirming healthy status), exhibiting stable graft union (no signs of incompatibility, cracking, or deterioration at grafting point), demonstrating appropriate vigor (trunk supporting balanced vine without excessive vegetative growth or inadequate vigor), and indicating optimal health (all trunk characteristics showing well established trunk performing as expected for age and cultivar). Well established mature grafted rootstock trunk transition characteristics includes= having recently completed establishment phase (transitioned from establishing to mature status within past 3-8 years), showing no signs of advancing age (absence of old trunk characteristics such as declining growth, heartwood formation, or declining vigor), displaying prime condition (trunk at peak of structural and physiological capacity), maintaining vigorous growth (continued active diameter increase at stable mature rate without decline), and representing early maturity (well established trunk in first half of mature phase with many years before old trunk transition). Well established mature grafted rootstock trunk production capacity includes= supporting full commercial crop loads (adequate capacity for consistent annual production without overload stress), enabling high fruit quality (stable resource supply supporting optimal berry development and composition), providing balanced vine performance (trunk capacity appropriate for vigorous growth and adequate fruit production), maintaining consistent year-to-year performance (stable trunk function reducing production variability), and demonstrating economic productivity (well established trunk supporting profitable grape production justifying vineyard investment). Well established mature grafted rootstock trunk as quality indicator includes= trunk condition reflecting vineyard management (well established trunk characteristics indicating successful establishment practices and ongoing care), diameter growth showing site suitability (consistent annual increase confirming appropriate rootstock-site matching), bark health indicating disease management (clean healthy bark showing effective vineyard pest and disease control), graft union integration confirming compatibility (stable well-integrated union validating rootstock-scion combination choice), and overall appearance revealing vineyard quality (robust healthy well established trunk characteristic of well-managed quality vineyard). Well established mature grafted rootstock trunk significance includes= representing optimal early mature phase (well established trunk marking entry into peak productive period with decades of potential ahead), demonstrating successful grafting and establishment (healthy well established trunk confirming first 7-8 years' management succeeded), providing foundation for sustained production (robust trunk enabling reliable annual cropping for many years), showing maximum productive potential (trunk at peak capacity before any age-related decline), and exemplifying quality viticulture (well established mature trunk characteristic of successful modern wine grape production).
Well_Established_Mature_Grafted_Rootstock_Trunk emphasizes early to mid mature phase (years 8-18) and is distinguished from Fully_Integrated_Mature_Grafted_Rootstock_Trunk (later mature phase) by focus on recently achieved maturity and early prime productive period. Critical for understanding early optimal production, peak capacity before aging, successful establishment confirmation, and the vigorous mature phase offering decades more productive potential.
Early mature grafted rootstock trunk
Frühreifer Wurzelstockstamm
Gut etablierter reifer verpfropfter Unterlagenstamm
Kürzlich gereifter Pfropfstamm
Prime young-mature grafted rootstock trunk
Recently matured grafted rootstock trunk
Well Established Mature Grafted Rootstock Trunk
Pinot Noir well established mature grafted scion trunk on 3309C low-vigor rootstock in Burgundy at 11 years post-planting shows 68-82 mm scion diameter with 11 annual rings (recent 3 years averaging 2.5-3.2 mm width appropriate for low-vigor rootstock), graft union at 8 cm above soil with stable 110-115% swelling indicating excellent compatibility, rootstock diameter 72-86 mm showing proportional development, scion sapwood 20-26 mm with rootstock-mediated drought tolerance, 155-185 g combined starch reserves (scion 55-68 g, rootstock 100-117 g), and peak productivity 4.2-5.8 kg per vine (78-98 hL/ha) with excellent fruit quality reflecting low-vigor rootstock concentration benefits
A well established mature grafted scion trunk is a mature grafted scion trunk at peak developmental status (typically years 8-16 post-planting) characterized by optimal scion-rootstock integration, maximum functional benefit from rootstock selection, peak carbohydrate storage capacity, and highest productivity potential for the specific scion-rootstock combination, representing the culmination of grafted vine development with rootstock advantages fully realized. Well established mature grafted scion trunk characteristics include= scion trunk diameter approaching cultivar-rootstock maximum (varying by combination= vigorous rootstocks 85-125 mm, moderate 70-105 mm, low-vigor 60-95 mm at year 10-12), graft union completely stable (diameter ratio 108-118% maintained without drift suggesting excellent compatibility), optimal scion sapwood cross-section (20-35 mm radial depth providing maximum rootstock-mediated water transport), scion heartwood occupying 30-50% of trunk (providing mechanical support while maintaining adequate sapwood), and combined scion-rootstock carbohydrate storage at maximum (140-210 g total= scion 50-75 g, rootstock 90-135 g providing optimal spring growth support and stress buffering). Well established mature grafted scion trunk anatomical optimization includes= maximum functional xylem vessels in scion sapwood (providing rootstock-modulated water transport 50-120 L/day depending on rootstock vigor and canopy size), optimal scion phloem cross-section (2-3 mm active phloem supporting efficient nutrient distribution), well-developed union vascular integration (xylem and phloem continuity across union approaching seamless connection in highly compatible combinations), mature union callus fully lignified (union mechanical strength at maximum reducing breakage risk to minimal levels), and rootstock trunk proportional development (rootstock diameter 95-115% of scion at equivalent positions indicating balanced growth). Well established mature grafted scion trunk physiological optimization includes= peak rootstock-mediated benefits fully expressed (drought tolerance, nutrient efficiency, vigor control, disease resistance at maximum levels for rootstock genotype), optimal scion hydraulic conductance per unit leaf area (rootstock providing adequate water supply without excessive oversizing or restrictive undersizing), maximum photosynthetic efficiency per unit crop (established scion-rootstock physiology supporting efficient carbon assimilation with rootstock-appropriate leaf area/crop balance), balanced rootstock-influenced hormone production (appropriate auxin, cytokinin, gibberellin levels supporting healthy development without excessive vigor or deficiency), and established stress response patterns (drought tolerance, cold hardiness, salinity resistance reflecting rootstock contribution at peak expression). Well established mature grafted scion trunk productivity characteristics include= maximum sustainable yield for combination (vigorous rootstocks 6-11 kg per vine, moderate 5-8 kg, low-vigor 3-6 kg, optimized for rootstock vigor potential), optimal fruit quality for combination (rootstock effects on berry composition, phenolics, aromatics fully expressed), minimal year-to-year variation in absence of extreme weather (stable productivity reflecting mature scion-rootstock homeostasis), excellent recovery capacity from moderate stress (robust combined reserves and rootstock stress tolerance supporting rapid recovery), and rootstock-appropriate crop load balance (vigorous rootstocks supporting higher yields without quality compromise, low-vigor rootstocks requiring crop limitation for quality maintenance). Well established mature grafted scion trunk represents optimal commercial status for grafted vineyard= peak economic return on grafted vineyard investment (maximum yield with quality appropriate for rootstock-scion combination), full realization of rootstock benefits (phylloxera resistance, soil adaptation, vigor control fully functional), lowest per-unit production costs (established vines requiring minimal remedial intervention with rootstock providing stress mitigation), greatest flexibility for crop management (adequate reserves and rootstock support enabling various crop strategies), and optimal terroir expression through rootstock lens (mature scion physiology expressing site characteristics as modulated by rootstock characteristics). Well established mature grafted scion trunk duration varies by rootstock and site= optimal sites with well-matched rootstocks may maintain well-established status 10-25 years (years 8-33 post-planting), mismatched rootstock-soil combinations showing briefer peak 6-15 years (years 8-23), and exceptional combinations with intensive management potentially extending peak 15-35 years (years 8-43).
Well_Established_Mature_Grafted_Scion_Trunk emphasizes peak developmental status with rootstock and is distinguished from Mature_Grafted_Scion_Trunk (broader productive phase) and Productive_Mature_Grafted_Scion_Trunk (sustained production emphasis) by focus on optimal scion-rootstock integration and maximum benefit realization. Critical for understanding peak grafted vineyard value period, optimal rootstock benefit timing, importance of rootstock-soil matching for peak performance, and economic analysis of grafted vineyard productive lifespan.
Höchstreifer Veredelungsstamm
Optimal entwickelter gepfropfter Stamm
Optimal maturity grafted scion
Peak grafted trunk development
Voll integrierte Aufpropfung
Well Established Mature Grafted Scion Trunk
A well lit upper canopy apical leaf at node 12 on a shoot of Vitis vinifera cv. Cabernet Sauvignon in Bordeaux, France, positioned in upper canopy zone on east-facing canopy receiving morning sun and filtered afternoon light totaling approximately 72% of full sunlight (daily PPFD averaging 1150 μmol photons m⁻² s⁻¹), measuring 13.5 cm length by 12.8 cm width representing typical apical zone dimensions, exhibiting thick lamina of 312 μm with two well-developed palisade layers totaling 95 μm and one partial third layer in some lamina regions, substantial cuticle of 5.8 μm providing good protection, high chlorophyll content of 495 μg cm⁻², moderate trichome coverage creating medium green-gray abaxial appearance, dark green adaxial surface without pronounced bronze coloration, demonstrating high photosynthesis rate of 12.8 μmol CO₂ m⁻² s⁻¹ representing excellent function without maximum extreme, moderate light compensation point of 48 μmol photons m⁻² s⁻¹ enabling positive carbon balance across broad light ranges, good photoprotective capacity maintaining PSII efficiency of 0.82, occurring at 3 weeks post-emergence during final expansion phase, measured at shoot BBCH stage 81 (beginning of berry ripening) in early August, representing optimal well-lit upper canopy conditions supporting high-quality leaf development and function.
An upper canopy apical leaf experiencing favorable but not extreme light exposure in elevated canopy positions in Vitis vinifera, characterized by location receiving 60-80% of full sunlight representing well-lit conditions without the intensity extremes of fully exposed positions, development under moderate to high light during expansion creating intermediate to strong sun leaf characteristics including thick lamina typically 280-340 μm with two to three palisade cell layers, substantial cuticle development of 4.5-6.5 μm, high chlorophyll concentrations of 450-550 μg cm⁻², moderate to high stomatal density, medium to dark green coloration, and physiological characteristics including high photosynthesis rates of 11-14 μmol CO₂ m⁻² s⁻¹, moderate light compensation points of 40-60 μmol photons m⁻² s⁻¹, and balanced photoprotective mechanisms providing adequate protection without the extreme resource investment required for maximum sun exposure tolerance. Well lit upper canopy leaves occupy favorable positions balancing excellent light availability supporting high photosynthetic capacity with avoidance of the most extreme light intensities that increase photoinhibition risk and require maximum investment in protective structures, creating leaves with high functional performance, good efficiency across moderate to high light ranges, and reliable contribution to canopy carbon gain without the vulnerability to extreme light stress affecting fully exposed positions.
Well lit upper canopy leaves represent optimal positions balancing light availability and stress exposure, distinguished from sun exposed leaves by moderate rather than extreme light intensities and from less-illuminated positions by favorable light supporting strong sun leaf development. The 60-80% full sun range represents the sweet spot for many grapevine leaves, providing adequate irradiance to saturate or nearly saturate photosynthesis while avoiding the highest intensities that increase photoinhib risks and require maximum protective investment. Position effects creating well-lit conditions include east or west-facing canopy orientations receiving direct sun during part of the photoperiod, upper canopy zones with some sparse shading from thin overlying foliage, or elevated interior positions receiving good diffuse light. Anatomical characteristics show strong sun leaf tendencies with multiple palisade layers and thick cuticles but typically less extreme than maximum sun exposure positions. Functional performance shows high photosynthetic rates with good efficiency, lower light compensation points than fully exposed leaves enabling function across broader light ranges, and reduced vulnerability to photoinhibition and stress. Well lit upper canopy leaves often provide optimal cost-benefit ratios, producing high photosynthetic returns without requiring maximum protective investments.
favorable exposure apical leaf
gut-beleuchtetes obere Kanopie Blatt
moderate high-light upper leaf
mäßig hochlicht oberes Blatt
optimal illumination upper leaf
optimal-belichtetes apikales Weinblatt
Based on Schultz (1996) "Leaf absorptance of visible radiation in Vitis vinifera L.= estimates of age and shade effects with a simple field method" Scientia Horticulturae 66=93-102 Poni et al. (2009) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 60=397-407 Flexas et al. (2002) "Drought-inhibition of photosynthesis in C₃ plants= stomatal and non-stomatal limitations revisited" Annals of Botany 89=183-189 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 259-294.
Well Lit Upper Canopy Leaf
A whip-and-tongue graft of Riesling clone 239 scion on Kober 5BB rootstock bears a Whip_Graft_Role, prepared by a skilled propagator making 3 cm diagonal cuts on both components with interlocking 1 cm tongue cuts, then wrapped tightly with budding tape before callusing—the technique preferred for this nursery's smaller production volume. A whip graft of Gewürztraminer on 101-14 Millardet serves in a Whip_Graft_Role for Alsatian nursery production, where traditional hand grafting maintains artisanal quality standards despite lower productivity than machine methods. A whip graft of heritage variety Trousseau onto 3309C bears a Whip_Graft_Role in a conservation propagation program, where the small quantities required do not justify machine setup and the operator's skill ensures high success with limited material.
A propagation material role that inheres in a grafted combination created using the whip graft or whip-and-tongue technique, where matching diagonal cuts (and optionally interlocking tongue cuts) on scion and rootstock create aligned cambial surfaces for union formation. Whip grafting is a traditional hand-grafting technique that remains important for specialized applications, small-scale production, and situations where omega machines are unavailable or impractical. The basic whip graft involves matching diagonal cuts (approximately 2.5-4 cm long at 20-30° angle) on scion base and rootstock apex the whip-and-tongue modification adds interlocking vertical cuts (tongues) that mechanically interlock the components and increase cambial contact area. Whip graft success requires skilled hand technique (matching cut angles and lengths), diameter compatibility (within 2-3 mm for adequate cambial alignment), and secure wrapping (budding tape, raffia, or parafilm) to maintain contact during callusing. The technique achieves success rates of 80-90% with skilled operators but requires substantially more time per graft than machine omega grafting.
Whip_Graft_Role captures the traditional hand-grafting technique that preceded machine omega grafting and remains relevant for specialized applications. The whip-and-tongue modification substantially improves on the simple whip graft by providing mechanical interlocking and increased cambial contact, achieving success rates approaching machine grafting when performed by skilled operators. The technique requires manual dexterity and practice (typically 6-12 months to achieve consistent quality) but enables grafting with minimal equipment, making it accessible for small producers, research applications, and situations where machine investment is unjustified. Whip grafting accommodates greater diameter variation than omega grafting—modest mismatches can be compensated by aligning one side's cambium—providing flexibility for working with variable-diameter material. The technique's continued importance reflects the diversity of grapevine propagation contexts, from large commercial nurseries (omega-dominated) through small specialty producers and research programs where hand grafting remains practical and appropriate.
Geißfußveredelung
Kopulation
Kopulationsveredelung
Zungenpfropfung
hand graft
splice graft
tongue graft
whip graft
whip-and-tongue graft
Hartmann HT et al. (2011) Plant Propagation 8th Ed, Pearson pp.: 450-452 Galet P (1998) Grape Varieties and Rootstock Varieties, Oenoplurimedia Pongrácz DP (1983) Rootstocks for Grape-vines, David Philip Publishers Walter B, Martelli GP (1996) Sanitary Selection of the Grapevine, INRA Editions
Whip Graft Role
Merlot vineyard showing white fine feeder roots in May, soil cores revealing abundant white roots 0.4-0.8 mm diameter comprising 58% of fine feeder root population, white roots showing characteristic fuzzy appearance from abundant root hairs, FDA viability staining showing 94% of white roots viable versus 45% of tan roots, isotope tracing showing white roots providing 78% of feeder root nitrogen uptake despite comprising 58% of population, white fine feeder roots demonstrating young active status with peak absorption capacity
White fine feeder root is fine feeder root showing characteristic white or cream coloration indicating youth and active function, characterized by white appearance resulting from lack of pigmentation in young tissues, recently formed status typically 1-6 weeks post-maturation representing early active phase, maximum physiological activity with peak metabolism and uptake, abundant root hair production creating white fuzzy coating, and serving as visual indicator of functional status and root system health with white root abundance reflecting favorable growing conditions and active root production, with white fine feeder roots showing physiological characteristics including recently completed cell differentiation and maturation, dense cytoplasm with high organelle content, thin unpigmented cell walls lacking phenolic compounds or melanin, active membrane systems with maximum transport capacity, developing arbuscular mycorrhizal colonization with actively forming arbuscules, and demonstrating that white coloration represents diagnostic feature indicating young active feeder roots at peak absorption capacity providing convenient visual assessment of functional root population without requiring elaborate testing. White fine feeder root coloration basis includes= showing white to cream color (characteristic appearance from unpigmented young tissues), exhibiting lack of phenolics (white roots lacking accumulated phenolic compounds that create tan or brown coloration in older roots), demonstrating thin walls (young thin cell walls appearing translucent contributing to white appearance), revealing dense cytoplasm (abundant cytoplasm in young cells potentially contributing to white opaque appearance through light scattering), and displaying root hair coverage (abundant white root hairs potentially enhancing overall white fuzzy appearance). White fine feeder root age and status includes= indicating recent formation (white color typically indicating roots 1-6 weeks post-maturation in early functional phase), showing active status (white roots generally functional and actively absorbing contrasting with tan aging or brown dying roots), exhibiting youth markers (white coloration correlating with other youth indicators including turgidity, abundant root hairs, dense cytoplasm), demonstrating peak function (white roots typically in peak functional period showing maximum uptake capacity), and revealing vitality (white appearance indicating healthy vital roots contrasting with compromised or stressed roots). White fine feeder root physiological characteristics includes= showing maximum metabolic rate (white roots exhibiting highest respiration and metabolic activity among feeder root population), exhibiting peak uptake capacity (white roots demonstrating highest absorption rates per unit surface area or length), demonstrating active growth potential (white roots potentially still capable of elongation if conditions favorable), revealing responsive capacity (white roots most responsive to environmental signals and plant demands), and displaying optimal function (white roots representing optimal functional state before aging-related declines begin). White fine feeder root root hair production includes= showing abundant production (white roots typically producing root hairs at maximum density), exhibiting active growth (root hair population on white roots including many hairs still elongating), demonstrating optimal function (root hairs on white roots at peak absorption capacity with intact membranes), revealing extensive coverage (white root hair zone potentially extending several centimeters behind tip), and displaying visual impact (abundant white root hairs creating characteristic fuzzy appearance enhancing white coloration). White fine feeder root as visual indicator includes= providing functional assessment (white root abundance offering convenient visual estimate of functional feeder root population), enabling field evaluation (white roots visible during soil examination providing immediate feedback on root system status), demonstrating condition indication (abundant white roots indicating favorable soil conditions and healthy root system), revealing seasonal patterns (white root proportion varying seasonally reflecting production and aging patterns), and displaying management response (white root abundance responding visibly to irrigation, fertilization, or other practices). White fine feeder root versus colored roots includes= contrasting with tan roots (white indicating young active versus tan indicating mature aging), showing difference from brown roots (white indicating living functional versus brown indicating dying or dead), exhibiting age correlation (white to tan to brown progression correlating with aging from young through mature to senescent), demonstrating functional implications (white roots providing majority of uptake versus tan contributing less and brown providing zero), and revealing population dynamics (white-tan-brown proportions indicating root population age structure and turnover). White fine feeder root environmental and management effects includes= showing production indication (abundant white roots indicating active root production responding to favorable conditions), exhibiting irrigation response (white root abundance increasing with adequate irrigation maintaining favorable moisture), demonstrating fertilization effects (white roots proliferating in response to fertilizer placement), revealing seasonal variation (white root proportion peaking spring and fall when conditions favorable for growth), and displaying stress effects (white root proportion declining under drought, heat, or other stresses as production decreases and mortality increases). White fine feeder root in research and assessment includes= enabling visual sorting (white color allowing rapid separation of young active roots from older roots during processing), providing vitality proxy (white root abundance serving as proxy for functional root population in field assessments), demonstrating photographic documentation (white roots visible in photographs allowing image-based root assessment), revealing temporal tracking (sequential observations documenting white root production and aging over time), and displaying practical utility (white color providing practical convenient indicator for growers and researchers assessing root system status).
White_Fine_Feeder_Root emphasizes characteristic white coloration indicating youth and is distinguished from tan or brown roots by focus on color as functional indicator. Critical for understanding visual assessment, youth indication, peak function, root hair abundance, vitality marker, and the white coloration providing convenient diagnostic feature indicating young active feeder roots at peak absorption capacity.
Fresh feeder root
Junge weiße Wurzel
Unpigmented fine root
Unpigmentierte Saugwurzel
Weiße Feinwurzel
Young white root
White Fine Feeder Root
White flower aroma assessment in Moscato d'Asti reveals the signature profile of orange blossom (8.8 on 10-point scale) and jasmine (7.5) from linalool concentration of 850 μg/L, complemented by honeysuckle (5.2) and subtle acacia (3.8), with the orange blossom dominance typical of Muscat Blanc à Petits Grains distinguishing it from Muscat of Alexandria where more musky, less pure floral character appears, demonstrating how white flower expression enables sub-variety identification within the Muscat family.
White flower aroma quality is a floral aroma quality that represents olfactory perception evoking white-petaled flower-associated scents including jasmine, honeysuckle, orange blossom, acacia, elderflower, and hawthorn notes, characterizing the delicate, often sweet, heady aromatic impression particularly associated with Muscat varieties and contributing elegance to diverse white wine styles. Within the BFO framework, this quality specifically addresses the white flower subcategory of floral aroma distinguished by its association with sweetness perception and aromatic intensity, recognizing that white flower notes typically evoke impressions of richness and opulence even in technically dry wines. The ontological significance lies in white flower aroma serving as the signature floral expression for Muscat family varieties where orange blossom and jasmine define varietal identity, while contributing complexity notes to non-aromatic varieties where elderflower in Sauvignon Blanc or acacia in Chardonnay indicate site and winemaking quality. The compositional basis for white flower aroma includes linalool (contributing jasmine and orange blossom character, present at high concentrations in Muscat), geraniol (adding rose-geranium notes that cross into white flower perception), nerol (orange blossom), and certain aromatic esters (methyl anthranilate in some labrusca hybrids providing distinctive grape-floral character). The perception intensity of white flower notes shows strong temperature dependence, with warmer serving temperature releasing more terpenes and enhancing floral perception, making serving protocol critical for full expression of white flower character in aromatic varieties. The ripeness association of white flower character typically shows increased floral intensity with grape maturity as terpene precursor accumulation proceeds, though excessive ripeness may shift floral character toward heavier, less elegant expression. Understanding white flower aroma quality enables varietal identification for Muscat family varieties, quality assessment based on floral elegance and intensity, and style characterization where white flower prominence distinguishes aromatic expressions.
White_Flower_Aroma_Quality addresses the white-petaled flower subcategory of floral aroma, encompassing jasmine, orange blossom, honeysuckle, elderflower, and related notes. This quality is particularly important for Muscat variety evaluation where white flower character defines varietal identity. The association with sweetness perception makes white flower notes relevant for style assessment even in dry wines where floral character creates impression of richness.
Holunderblütennote
Jasminduft
Orangenblütencharakter
Weißblütenaroma
elderflower scent
jasmine notes
orange blossom character
white blossom aroma
Selli S et al. (2006) Volatile composition of Muscat wine during fermentation. Journal of Food Engineering 74(4): 545-549 Fenoll J et al. (2009) Changes in the free and glycosidically-bound volatile compounds during ripening in five Muscat varieties. Food Chemistry 114(2): 420-428
White Flower Aroma Quality
Thompson Seedless (Sultana) bears a White_Table_Grape_Scion_Role as the world's most planted table grape variety, with small-medium seedless berries (2-4 g), neutral sweet flavor, green-to-amber color, and versatility for fresh, dried, and juice markets—California alone producing over 200,000 tonnes annually for fresh consumption. Sugraone (Superior Seedless) serves in a White_Table_Grape_Scion_Role as a premium large-berry white seedless, with berries of 6-8 g, yellow-green color, light muscat flavor, and crisp texture commanding premium early-season prices. Italia (Ideal, Muscat Italia) bears a White_Table_Grape_Scion_Role as the classic European seeded table grape, with very large berries (8-12 g), distinctive muscat aroma, golden-green color, and crisp texture defining quality table grapes in Mediterranean and South American markets.
A fresh consumption table grape role that inheres in a grafted scion cultivar by virtue of its white, green, or yellow berry skin coloration, lacking significant anthocyanin pigmentation. White table grape scions develop their characteristic green-to-golden coloration from chlorophyll (in green-skinned varieties) and carotenoids and flavonols (in yellow-golden varieties), with color typically shifting from green toward yellow-amber as ripening progresses and chlorophyll degrades. White/green table grapes traditionally dominated global markets before the rise of seedless red varieties, and remain the largest color category in many markets due to the success of Thompson Seedless (Sultana) and its descendants. Consumer perception associates white/green grapes with refreshing, crisp character, making them particularly popular in warm-weather markets. Major white table grape varieties include both seeded heritage cultivars (Italia, Muscat of Alexandria) and modern seedless varieties (Thompson Seedless, Sugraone, Autumn King, Princess, Sweet Globe) that represent the majority of production volume.
White_Table_Grape_Scion_Role encompasses the green-through-yellow color spectrum, with "white" being the traditional commercial designation regardless of actual hue. Thompson Seedless remains the global volume leader, though its relatively small berry size and neutral flavor have ceded premium market segments to larger-berried, flavor-distinctive newer varieties. Muscat-flavored white table grapes (Italia, Sugraone, Muscat of Alexandria) command premium prices in markets valuing aromatic character. Color uniformity is less challenging than for red grapes, though excessive sun exposure can cause browning/raisining while insufficient exposure yields pale, unattractive coloration. Breeding trends favor larger berries, enhanced crunch, and distinctive flavor profiles while maintaining the refreshing character consumers associate with white grapes.
green table grape
grüne Tafeltraube
weiße Esstraube
weiße Tafeltraube
white eating grape
white table grape
yellow table grape
Creasy GL, Creasy LL (2009) Grapes, CABI Christensen LP (2000) Raisin Production Manual, UC ANR Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press Fidelibus MW et al. (2017) California Table Grape Commission Production Guidelines
White Table Grape Scion Role
Chardonnay bears a White_Wine_Scion_Role globally as the most planted white wine variety, producing styles from lean, unoaked Chablis (12% alcohol, 7 g/L acidity) through rich, malolactic-fermented, barrel-aged Meursault. Riesling serves in a White_Wine_Scion_Role across Germany, Alsace, and Austria, contributing pronounced terpene aromatics (linalool, geraniol) and high natural acidity (8-12 g/L) enabling diverse expressions from bone-dry Grosses Gewächs to nobly sweet Trockenbeerenauslese. Sauvignon Blanc bears a White_Wine_Scion_Role in Loire, Bordeaux, and New Zealand, defined by volatile thiolic aromatics (3-mercaptohexanol, 3-mercaptohexyl acetate) at concentrations of 500-5000 ng/L creating distinctive grapefruit, passionfruit, and herbaceous character.
A wine production scion role that inheres in a grafted scion cultivar by virtue of its designation for white wine production, where juice is typically separated from skins before or shortly after crushing to prevent significant phenolic extraction, yielding wines ranging from water-white through pale straw to deep gold. White wine scions are characterized by green, yellow, or pink-skinned berries with colorless flesh, though the white wine role may also be fulfilled by red-skinned cultivars processed without skin contact (blanc de noirs). White wine scions are selected for aromatic compound accumulation (free and bound terpenes, thiols, esters, norisoprenoids) that express varietal character without masking by tannins, appropriate acidity profiles (typically 6-9 g/L total acidity at harvest) providing freshness and aging potential, and sugar accumulation sufficient for target alcohol levels (typically 11-14% v/v). Phenolic content in white wines is dramatically lower than reds (typically 200-500 mg/L versus 2000-4000 mg/L), with extraction limited to brief juice-skin contact during pressing. White wine production encompasses diverse styles from austere, mineral-driven wines to rich, oak-aged expressions.
White_Wine_Scion_Role is subclassed by quality tier (Premium, Bulk) and aromatic profile (Aromatic, Neutral), reflecting the sensory and commercial diversity within white wine production. The aromatic/neutral distinction is particularly important for white wines where varietal aroma compounds are not masked by phenolic extraction. Many white wine scions also serve in Sparkling_Wine_Scion_Role (Chardonnay, Pinot Blanc, Chenin Blanc) and some may produce rosé or orange wines through modified vinification. White wines account for approximately 45% of global still wine production, with Chardonnay, Sauvignon Blanc, and Pinot Grigio/Gris dominating international markets.
Weißweinrebe
Weißweinsorte
weiße Rebsorte
white grape variety
white wine cultivar
white wine variety
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Jackson RS (2014) Wine Science 4th Ed, Academic Press pp.: 350-420 Waterhouse AL et al. (2016) Understanding Wine Chemistry, Wiley Ribéreau-Gayon P et al. (2006) Handbook of Enology Vol 2, Wiley
White Wine Scion Role
The whole genome sequence of Cabernet Sauvignon (Chin et al. 2016) produced using Pacific Biosciences single-molecule real-time (SMRT) sequencing at 140× coverage represents the first diploid-phased assembly of a highly heterozygous plant genome — the two haplotypes (590 Mb total diploid assembly) were resolved into separate chromosome-scale pseudomolecules revealing 36,139 predicted protein-coding genes (approximately 6,000 more than the haploid PN40024 reference due to hemizygous gene content), extensive structural variation between parental haplotypes (Cabernet Franc and Sauvignon Blanc contributions distinguishable at the chromosomal level), and haplotype-specific transposable element insertions including the Gret1 retrotransposon in the VvMYBA1 locus present on one haplotype and absent on the other, explaining Cabernet Sauvignon's capacity for color-variant somatic mutation.
A genetic information entity that records the complete or near-complete nucleotide sequence of all three compartmental genomes (nuclear, chloroplast, and mitochondrial) of a specific Vitis vinifera accession, generated through shotgun sequencing, assembly, and annotation to produce a reference-quality or draft-quality genome that enables comprehensive analysis of gene content, structural variation, regulatory elements, repetitive sequences, and evolutionary features at the whole-genome scale.
Whole_Genome_Sequence represents the most comprehensive level of genetic information for a grapevine accession, providing the complete nucleotide template from which all other genetic information types (SSR profiles, SNP genotypes, gene sequences, regulatory elements) are derived. The progression of Vitis genome sequencing from the first reference genome (PN40024, Jaillon et al. 2007, Sanger sequencing, haploid-equivalent assembly) through second-generation sequencing-based resequencing projects (Liang et al. 2019, 472 accessions at 10-40× coverage) to third-generation long-read assemblies producing chromosome-scale, haplotype-resolved references (Chin et al. 2016 for Cabernet Sauvignon, Massonnet et al. 2020 for Chardonnay) reflects the rapid technological evolution that has made whole-genome sequencing increasingly accessible for individual cultivar characterization: the current cost of approximately 1,000-3,000 EUR per genome at 30× coverage enables routine whole-genome sequencing for breeding programs, clonal identification projects, and germplasm characterization initiatives. Pan-genome approaches that capture the full spectrum of genomic variation across multiple cultivars — including sequences absent from any single reference — are emerging as the next analytical framework for exploiting whole-genome data in grapevine improvement.
Genom-Referenzsequenz
Gesamtgenom-Sequenz
complete genome sequence
genomic reference sequence
vollständige Genomsequenz
whole genome assembly
Jaillon O et al. (2007) The grapevine genome sequence suggests ancestral hexaploidization in major angiosperm phyla, Nature 449: 463-467 Chin CS et al. (2016) Phased diploid genome assembly with single-molecule real-time sequencing, Nature Methods 13: 1050-1054 Liang Z et al. (2019) Whole-genome resequencing of 472 Vitis accessions, Nature Communications 10: 1190 Massonnet M et al. (2020) The genetic basis of sex determination in grapes, Nature Communications 11: 2902
Whole Genome Sequence
Whole-plant hydraulic conductance comparison of rootstock-scion combinations reveals the integration of component conductivities affecting vine water relations, with Cabernet Sauvignon on 110R showing Kplant of 4.8 mmol/m² leaf/s/MPa (high root conductance from drought-adapted rootstock), the same scion on SO4 showing 3.2 mmol/m² leaf/s/MPa (lower root conductance limiting whole-plant capacity), with the 50% difference in Kplant translating to 0.4 MPa difference in midday leaf water potential at similar transpiration rates, demonstrating how rootstock effects on whole-plant conductance affect scion water status and stress exposure.
Whole plant hydraulic conductance quality is a hydraulic conductivity quality that represents the integrated water transport capacity from soil through root, stem, and leaf pathways to evaporating surfaces, characterizing the total conductance of the soil-plant-atmosphere continuum that determines the relationship between soil water potential and leaf water potential at any given transpiration rate. Within the BFO framework, this quality addresses the integrated system-level hydraulic function, distinguishing whole-plant conductance (Kplant) from component conductivities while recognizing that Kplant emerges from the series and parallel arrangement of root, stem, and leaf pathways whose individual properties combine to determine overall water transport capacity. The ontological significance lies in whole-plant hydraulic conductance establishing the fundamental relationship between environmental water availability and plant water status, where Kplant determines the leaf water potential achieved at any transpiration rate according to E = Kplant × (Ψsoil - Ψleaf), making whole-plant conductance a key determinant of vine water relations performance across the range of soil moisture and evaporative demand conditions experienced in vineyards. The integration of component conductivities follows resistance principles where series resistances add (1/Ktotal = 1/Kroot + 1/Kstem + 1/Kleaf) and parallel resistances combine reciprocally, with the component contributing the largest resistance (lowest conductance) limiting whole-plant capacity and representing the primary target for improving hydraulic function. The dynamic nature of whole-plant hydraulic conductance reflects the regulation and vulnerability of component pathways, with Kplant typically declining through the day as leaf and possibly root conductances decrease, declining seasonally as cumulative stress affects system integrity, and potentially showing rapid changes from embolism events or aquaporin regulation. The rootstock and scion effects on whole-plant conductance are both significant in grafted grapevines, with rootstock genotype determining root conductance and often affecting scion xylem development, while scion cultivar determines leaf properties, creating the combined hydraulic architecture that affects vine water relations performance. Understanding whole-plant hydraulic conductance quality enables assessment of integrated vine water transport capacity, prediction of leaf water potential from soil water status and transpiration demand, and evaluation of rootstock-scion combinations for hydraulic performance under target environmental conditions.
Whole_Plant_Hydraulic_Conductance_Quality addresses the integrated system-level water transport capacity from soil to evaporating surfaces. This quality determines the relationship between soil water availability and plant water status. The class supports rootstock-scion evaluation and water relations modeling through whole-plant conductance characterization.
Ganzkörperleitfähigkeit
Gesamtpflanzenhydraulikleitfähigkeit
Kplant
SPAC conductance
SPAC-Leitfähigkeit
plant hydraulic conductance
whole-vine conductance
Tyree MT (2003) Hydraulic limits on tree performance: transpiration, carbon gain and growth of trees. Trees 17(2): 95-100 Alsina MM et al. (2011) Seasonal changes of whole root system conductance by a drought-tolerant grape root system. Journal of Experimental Botany 62(1): 99-109
Whole Plant Hydraulic Conductance Quality
Pinot Noir wide annual ring at year 3 post-planting shows 5.2-7.8 mm radial width reflecting vigorous establishment growth, expanded earlywood zone 2.5-3.8 mm with 3-4 rows of vessels 220-290 μm diameter providing substantial hydraulic capacity, latewood zone 2.7-4.0 mm with abundant fibers, ring representing pre-cropping vigor before first significant crop in year 4 when ring width declines to 2.8-4.2 mm
A wide annual ring is an annual ring characterized by increased radial width (typically 3.5-8 mm or more, compared to 1.5-4 mm for typical mature vine rings), indicating enhanced cambial activity and secondary xylem production during the formation year, reflecting favorable environmental conditions, high vine vigor, abundant resources, or reduced crop load promoting vegetative growth. Wide annual ring formation results from factors promoting cambial division and cell expansion= abundant water availability (optimal soil moisture supporting extended cell division and expansion periods), high nutrient availability (nitrogen, phosphorus, potassium promoting photosynthesis and growth), favorable temperature (warm growing seasons with long frost-free periods maximizing growth duration), low or absent crop load (absence of fruit allowing photosynthate allocation to vegetative growth, common in young pre-cropping vines or years following crop failure), vigorous rootstock effects (in grafted vines, vigorous rootstocks promoting enhanced scion growth), youth and establishment vigor (young vines in years 1-5 typically producing wide rings during rapid establishment), and optimal management (irrigation, nutrition, pest control supporting maximum growth potential). Wide annual ring anatomical characteristics include= increased radial width (3.5-8+ mm typical for wide rings compared to 1.5-4 mm in mature moderate-growth vines), expanded earlywood zone (large vessel band 1.5-4 mm radial width providing extensive hydraulic capacity), multiple rows of large earlywood vessels (2-5 vessel rows compared to 1-3 in narrow rings), maximum vessel diameter achieved (200-300 μm vessels approaching genetic maximum for cultivar), substantial latewood zone (1.5-4 mm providing mechanical strength and fiber content), optimal vessel density (35-55 vessels/mm² distributed through ring), and expanded parenchyma volume (more storage tissue providing carbohydrate reserve capacity). Wide annual ring vessel characteristics include= abundant large vessel production (numerous earlywood vessels providing high hydraulic conductance increment), maximum vessel diameter (optimal growing conditions enabling vessels to achieve genetic size potential), high vessel lumen area fraction (vessels occupying 40-55% of ring area maximizing transport capacity), long vessel elements (favorable growth allowing extended cell elongation producing 300-700 μm element length), and optimal vessel wall development (adequate time for secondary wall deposition producing strong functional vessels). Wide annual ring formation timing includes= extended growing season (early spring warmth initiating growth in March, late fall mild conditions extending growth through October), optimal early season (abundant April-June moisture and warmth promoting maximum earlywood production), favorable mid-season (July-August conditions supporting continued latewood formation), and lack of growth-limiting events (absence of mid-season drought, disease, or defoliation allowing uninterrupted cambial activity). Wide annual ring dendrochronological significance includes= indicating favorable years (wide rings marking wet years, optimal temperatures, or excellent growing conditions), enabling cross-dating (matching wide ring patterns across sites confirming synchronous favorable conditions), providing climate reconstruction (ring width correlating with precipitation surplus, warm temperatures, extended growing seasons), and revealing management effects (wide rings following reduced crop load, irrigation initiation, or fertility improvement). Wide annual ring pattern interpretation includes= single wide ring indicating exceptional year (unusually favorable conditions or reduced crop causing growth pulse), multiple consecutive wide rings indicating sustained favorable conditions (sequence of optimal years or young vine rapid establishment), progressively wider rings indicating improving conditions (soil amelioration, disease management improvement, or irrigation implementation), and wide inner rings indicating youth (young vines typically showing wide rings during establishment years 1-5 before cropping begins). Wide annual ring implications for vine physiology includes= substantial hydraulic conductance increment (wide ring adding significant water transport capacity), increased carbohydrate storage increment (abundant wood parenchyma providing enhanced reserve capacity), indication of excess vigor (particularly in mature vines, very wide rings suggesting excessive vegetative growth potentially compromising fruit quality), and resource allocation to wood (photosynthate invested in trunk growth rather than fruit or reserve storage potentially affecting productivity or stress resilience). Wide annual ring occurrence patterns vary= young vines consistently showing wide rings (years 1-5 typically 4-8 mm reflecting establishment vigor), vigorous sites occasionally showing wide rings (optimal water years on fertile soils producing enhanced growth), stressed sites rarely showing wide rings (even favorable years producing only moderate width on poor sites), and grafted vines on vigorous rootstocks showing wider rings (rootstock effect enhancing scion growth compared to own-rooted or low-vigor rootstock combinations). Wide annual ring in mature vines includes= potentially indicating excessive vigor (wine quality concerns in mature vines showing >5 mm rings suggesting need for vigor management), reflecting low crop load (suggesting need for crop load increase or improved fruit set), or indicating disease remission (wide ring following disease treatment or spontaneous remission showing recovery).
Wide_Annual_Ring emphasizes increased width favorable condition indicator and is distinguished from Annual_Ring (all rings) and Narrow_Annual_Ring (stress indicator) by increased dimensions reflecting optimal conditions or high vigor. Critical for understanding youth and vigor expression in wood, identifying favorable years in dendrochronology, assessing excessive vigor in mature vines, and interpreting establishment growth patterns in young vineyards.
Breiter Jahresring
Broad ring
Dicker Holzring
Favorable growth ring
Günstiger Wachstumsring
Thick ring
Wide Annual Ring
A wide petiolar sinus on a mature leaf at node 7 of Vitis vinifera cv. Sauvignon Blanc in Marlborough, New Zealand, exhibiting characteristic wide open V-shaped configuration with basal lobe margins separated by 3.2 cm at the widest point creating very substantial gap, sinus depth of 2.5 cm creating width-to-depth ratio of 1.28 indicating width exceeding depth, V-shaped contour with relatively acute sinus base angle of approximately 55 degrees, left and right basal lobe margins showing strongly divergent orientation with margins angled outward at approximately 75 degrees from median vein axis, petiole clearly visible through the wide gap for 4.2 cm above the node attachment extending well into the blade region, sinus margins formed by small divergent basal lobes measuring only 1.8 cm length each, wide petiolar sinus configuration serving as primary diagnostic feature for Sauvignon Blanc enabling confident identification and clear distinction from closed-sinus cultivars like Merlot or overlapping-sinus types like Pinot Noir, measured at BBCH stage 75 (berries pea-sized) in late June (Southern Hemisphere), representing very wide petiolar sinus typical of narrow-based cultivars with minimal basal lobe development.
A petiolar sinus exhibiting wide separation between basal lobe margins creating substantial gap at the petiole-blade junction in Vitis vinifera, characterized by opening width typically exceeding 2-3 cm measured at the widest point between basal lobe margins, clear visibility of the petiole for substantial length often 3-5 cm above the node attachment, basal lobe margins showing divergent orientation creating expanding rather than converging configuration, typically U-shaped or V-shaped contour with the gap width exceeding depth in many cases, occurrence in cultivars with small divergent basal lobes including Cabernet Sauvignon, Sauvignon Blanc, and many other narrow-based varieties, and functional implications providing maximum access to the petiole-blade junction and potentially reducing mechanical stress concentration through distributed load transfer across wider connection zone. Wide petiolar sinuses represent the open extreme of petiolar sinus configurations, developing through small divergent basal lobe patterns creating characteristic narrow-based leaf architecture, exhibiting high diagnostic reliability as cultivars with wide sinuses show consistent expression across environmental conditions, and serving as definitive identification features distinguishing open-sinus cultivar groups from closed or overlapping types.
Wide petiolar sinuses are distinguished from narrow, closed, or overlapping configurations by substantial gap width typically >2.5-3 cm, with exact thresholds somewhat arbitrary. The wide designation emphasizes the open accessible character creating maximum petiole visibility and blade base accessibility. Width often exceeds depth creating distinctive broad shallow sinus appearance contrasting with narrow deep configurations. Basal lobe characteristics determine sinus width, with small divergent basal lobes creating wide sinuses while large convergent lobes create narrow or closed sinuses. Shape shows variation from U-shaped (rounded base with parallel or diverging margins) to V-shaped (acute base with straight diverging margins), with V-shaped more common in very wide sinuses. Occurrence predominates in narrow-based cultivars showing weak basal lobe development, representing the majority of major commercial varieties including Cabernet Sauvignon, Sauvignon Blanc, Chardonnay, and Riesling. Genetic control is strong with wide sinus pattern showing consistent expression and high heritability. Environmental stability is high with sinus width showing minimal modification across vigor and resource conditions, making wide petiolar sinus a highly reliable diagnostic character. Functional implications include maximum petiole-blade junction accessibility potentially facilitating mechanical flexibility, though experimental support is limited. Ampelographic utility is exceptional with wide versus narrow/closed sinus serving as primary bifurcation in many identification keys, enabling immediate separation of major cultivar groups.
breite Basaleinbuchtung
broad basal sinus
gaping petiole sinus
offene Stielbucht
open petiolar sinus
weiter Petiolarsinus
Wide Petiolar Sinus
Merlot grafted on 1103P rootstock at 6 years post-grafting showing wide rootstock annual ring for year 3 measuring 0.94 cm radial width, wide ring reflecting vigorous juvenile growth phase with optimal conditions, wide ring structure showing extensive earlywood zone 0.62 cm width with abundant large vessels 140-175 micrometers diameter and substantial latewood zone 0.32 cm width providing mechanical strength, ring 3 followed by progressively narrowing rings years 4-6 (0.78, 0.64, 0.52 cm) showing expected juvenile-to-mature growth transition, wide ring adding estimated 18% increase to trunk water transport capacity in single year, management records confirming optimal irrigation and fertility during year 3, wide ring typical of vigorous 1103P rootstock in favorable conditions during early establishment phase
Wide rootstock annual ring is rootstock annual ring with increased radial width typically greater than 0.6 cm, characterized by extensive secondary xylem production during growing season reflecting vigorous cambial activity from favorable conditions including adequate water availability, optimal temperatures, sufficient nutrients, young vigorous trunk, inherently vigorous rootstock cultivar, or reduced competitive stress, showing expanded earlywood and latewood zones with abundant cells produced, increased vessel area significantly enhancing trunk water transport capacity, and serving as indicator of favorable growing conditions, high vigor, and optimal vine growth during formation year providing valuable dendrochronological information about favorable periods in trunk and vineyard history. Wide rootstock annual ring width definition includes= showing radial width greater than 0.6 cm (wide ring typically 0.6-1.2+ cm compared to typical 0.3-0.6 cm or narrow 0.1-0.3 cm rings), exhibiting expanded structure (substantial radial extent indicating vigorous xylem production during growing season), displaying prominent appearance (thick conspicuous band visible in cross-section much wider than adjacent rings), demonstrating robust growth (wide ring representing substantial diameter increase during that year), and showing relative wideness (ring considered wide in comparison to trunk's typical ring widths with interpretation depending on context). Wide rootstock annual ring formation causes includes= resulting from adequate water (abundant rainfall or optimal irrigation providing ample soil moisture supporting vigorous cambial activity), reflecting optimal nutrients (adequate nitrogen, phosphorus, and other nutrients enabling maximum growth), showing favorable temperatures (moderate temperatures and extended growing season allowing prolonged cambial activity), indicating juvenile vigor (young trunk years 1-10 naturally producing wide rings from rapid youthful growth phase), exhibiting high rootstock vigor (vigorous rootstock cultivar producing naturally wider rings than low-vigor cultivar under similar conditions), demonstrating reduced competition (minimal inter-vine competition, effective weed control, or optimal spacing providing resources for vigorous growth), and revealing management benefits (effective irrigation, fertilization, and cultural practices optimizing conditions for growth). Wide rootstock annual ring structural characteristics includes= showing extensive earlywood zone (broad spring wood formation with abundant large vessel production potentially 0.4-0.8 cm width providing maximum water transport capacity addition), exhibiting substantial latewood zone (considerable late season growth with thick latewood band potentially 0.2-0.4 cm width adding significant structural strength), displaying numerous cells (large number of xylem cells produced across ring indicating vigorous sustained cambial division activity throughout growing season), containing abundant large vessels (extensive production of large-diameter vessels particularly in earlywood significantly increasing trunk hydraulic conductivity), demonstrating full cell expansion (cells achieving maximum normal dimensions due to adequate water and resources during formation), and showing extended growth period (cambium remaining active throughout entire potential growing season producing maximum xylem before dormancy). Wide rootstock annual ring in trunk age context includes= common in young trunk (wide rings typical in young vigorous trunk years 1-10 reflecting rapid juvenile growth as normal healthy pattern), occasional in mature trunk (wide rings in mature trunk years 15-40 typically indicating particularly favorable year rather than consistent pattern), rare in old trunk (wide rings uncommon in aged trunk years 40+ except under exceptional conditions since age-related growth slowing typically limiting width), requiring age-adjusted interpretation (wide ring significance varying with trunk age with 0.8 cm expected in year 3 but remarkable in year 40), and demonstrating juvenile vigor (widest rings typically in earliest trunk years showing maximum growth rate during establishment phase). Wide rootstock annual ring favorable conditions signature includes= indicating wet year (wide ring typical signature of water-abundant growing season with optimal soil moisture throughout season), showing optimal irrigation (wide ring potentially indicating irrigation regime meeting vine water needs optimally), exhibiting favorable temperature patterns (wide ring reflecting moderate temperatures and extended frost-free growing season), demonstrating nutrient adequacy (wide ring indicating sufficient fertility supporting maximum cambial activity and xylem production), and revealing management success (wide ring potentially documenting effective cultural practices optimizing growth conditions). Wide rootstock annual ring and trunk capacity includes= substantially increasing trunk diameter (wide ring adding significant radial growth potentially 0.6-1.2 cm diameter increase substantially expanding trunk size that year), maximally increasing vessel area (wide ring producing abundant new vessels significantly enhancing trunk water transport capacity addition that year), optimally expanding functional capacity (wide ring adding maximum water transport, structural support, and storage capacity to trunk in single year), demonstrating vigorous capacity growth (wide ring showing trunk capacity expanding robustly potentially keeping pace with or exceeding canopy and crop expansion), and indicating adequate growth (persistent wide rings suggesting trunk adequately expanding to support vine needs assuming appropriate for age and cultivar). Wide rootstock annual ring in different rootstock cultivars includes= relative to cultivar vigor (wide defined relative to rootstock typical width with Riparia showing 0.5-0.6 cm considered wide while 1103P showing 0.5-0.6 cm possibly only moderate), showing cultivar-specific patterns (vigorous rootstocks naturally producing wider rings than low-vigor rootstocks requiring cultivar-appropriate interpretation), exhibiting different response magnitudes (some rootstock varieties producing particularly wide rings under optimal conditions while others showing more modest response), demonstrating genetic growth potential (wide ring showing rootstock variety maximum growth rate under favorable conditions), and revealing adaptation indicators (consistent wide rings potentially indicating good rootstock-site match with cultivar thriving under local conditions). Wide rootstock annual ring diagnostic value includes= indicating favorable year (isolated wide ring marking specific growing season with optimal conditions), revealing optimal period (series of consecutive wide rings indicating sustained favorable conditions from management, weather, or vine development), showing juvenile vigor (wide rings in young trunk indicating healthy vigorous establishment as expected normal pattern), demonstrating management success (wide rings potentially reflecting effective irrigation, fertility, or other cultural practices achieving growth objectives), enabling favorable event dating (wide rings associated with known favorable events such as wet years or management improvements providing temporal markers), and supporting vine evaluation (wide ring patterns informing assessment of vine health, vigor, and response to management informing future practices). Wide rootstock annual ring earlywood-latewood structure includes= showing extensive earlywood (wide ring typically featuring broad earlywood zone with abundant large vessel production substantially increasing hydraulic capacity), exhibiting substantial latewood (considerable latewood development providing significant mechanical strength addition), displaying balanced proportions (both earlywood and latewood well-developed with each contributing to ring width increase), demonstrating optimal vessel production (wide ring producing maximum number of large earlywood vessels significantly enhancing water transport capacity), and revealing favorable formation conditions (wide ring structure reflecting extended growing season with favorable conditions throughout spring, summer, and into fall). Wide rootstock annual ring in growth rate assessment includes= indicating rapid growth period (wide ring marking year of substantial trunk diameter expansion), showing high vigor year (wide ring suggesting elevated overall vine vigor during that growing season from favorable conditions), demonstrating growth potential (wide ring revealing maximum growth capacity when limiting factors removed), enabling growth rate quantification (wide ring width providing precise measurement of annual growth rate for favorable year), and supporting long-term analysis (patterns of wide and narrower rings revealing growth rate trends and condition variations over trunk life). Wide rootstock annual ring functional implications includes= substantially increasing water transport capacity (wide ring producing abundant vessels adding significantly to trunk hydraulic conductivity potentially 15-25% increase in single year), significantly enhancing structural support (considerable wood added providing major contribution to trunk mechanical strength and rigidity), substantially expanding carbohydrate storage (extensive ray parenchyma added in wide ring significantly increasing storage capacity), demonstrating optimal capacity expansion (wide ring showing trunk capacity increasing robustly supporting vine growth and productivity), and indicating functional adequacy (wide rings suggesting trunk capacity expanding appropriately with little constraint on vine development). Wide rootstock annual ring in vineyard management includes= indicating management success (wide rings potentially documenting effective practices achieving vigor and growth objectives), revealing favorable response (wide rings showing positive vine response to irrigation, fertility, or other management practices), supporting practice validation (consistent wide rings confirming management approach appropriate and effective), enabling treatment evaluation (comparing ring widths before and after management changes assessing intervention effectiveness such as irrigation installation or fertility program), and guiding future practices (wide ring patterns informing continuation or modification of successful management approaches). Wide rootstock annual ring in young vine context includes= expected in establishing trunk (wide rings normal in young trunk years 1-10 reflecting vigorous juvenile growth as healthy establishment pattern), indicating successful establishment (consistent wide rings confirming young vine thriving and establishing properly), demonstrating adequate resources (wide rings showing young vine receiving sufficient water, nutrients, and care for optimal development), revealing growth potential (wide rings in youth showing cultivar maximum growth capacity under favorable conditions), and supporting training system development (vigorous wide ring growth enabling rapid trunk diameter increase supporting training system establishment and first production). Wide rootstock annual ring multiple consecutive patterns includes= indicating sustained favorable conditions (series of wide rings suggesting persistent optimal circumstances from management, favorable weather cycles, or vine developmental stage), revealing optimal period (sequence of wide rings marking extended favorable period potentially multiple years of optimal growth), demonstrating consistent vigor (continued wide rings showing vine maintaining high vigor without decline or stress), suggesting optimal management (persistent wide rings potentially indicating sustained effective cultural practices maintaining favorable conditions), and documenting favorable climate period (consecutive wide rings potentially reflecting multi-year favorable climate pattern identifiable in regional climate records). Wide rootstock annual ring and dendrochronology includes= marking favorable years (wide rings corresponding to documented favorable years providing dendrochronological record of optimal conditions), indicating beneficial climate periods (sequences of wide rings revealing extended periods of favorable growing conditions), enabling cross-dating (matching wide ring patterns across multiple trunks or sites synchronizing chronologies and validating dating), supporting growth analysis (wide ring patterns revealing historical growth conditions and vine responses across decades), and documenting management history (wide rings potentially corresponding to known management changes such as irrigation installation or fertilization programs providing temporal markers). Wide rootstock annual ring measurement and analysis includes= measuring precise width (determining exact radial dimension typically 0.6-1.2+ cm for wide ring using microscope or digital imaging), comparing to trunk average (assessing wideness relative to trunk's typical ring width determining positive deviation from normal), analyzing in context (interpreting wide ring considering trunk age, rootstock variety, and known environmental or management history determining significance), examining anatomical details (studying earlywood-latewood structure, vessel characteristics, and cell dimensions in wide ring revealing favorable formation conditions), and integrating with records (correlating wide rings with weather data, management records, or known favorable periods determining causes). Wide rootstock annual ring vessel production includes= producing abundant large earlywood vessels (wide ring creating numerous large-diameter vessels 100-180 micrometers substantially increasing water transport capacity), generating extensive vessel network (dense vessel distribution throughout wide ring creating redundant parallel transport pathways), demonstrating optimal vessel development (vessels achieving maximum normal size and connectivity reflecting favorable conditions during differentiation), showing increased hydraulic conductivity per ring (wide ring adding disproportionately more transport capacity than narrow ring due to vessel number and size), and enabling capacity expansion (wide rings essential for trunk hydraulic capacity to keep pace with canopy and crop growth). Wide rootstock annual ring in trunk core sampling includes= prominent in increment cores (wide rings easily visible and identifiable in extracted core samples facilitating growth analysis), enabling rapid age determination (wide rings readily counted in cores though potentially requiring fewer rings counted if earliest years all wide), supporting detailed growth analysis (wide ring widths easily measured in cores revealing annual growth rates and favorable year identification), facilitating vigor assessment (wide ring patterns in cores documenting vigor history and growth trends without trunk destruction), and allowing temporal monitoring (cores extracted multiple years tracking whether wide ring patterns continuing, declining, or varying over time). Wide rootstock annual ring significance includes= indicating robust annual growth (wide ring representing substantial secondary xylem production and significant trunk diameter increase that year), revealing favorable conditions (wide ring documenting adequate water, optimal temperature, sufficient nutrients, or other favorable factors during formation year), demonstrating high vigor (wide ring showing vigorous cambial activity and rapid xylem production reflecting overall vine health and growth capacity), enabling favorable period identification (wide ring marking specific year of optimal conditions allowing temporal documentation of successful years), substantially expanding functional capacity (wide ring adding significant water transport, structural support, and storage capacity enhancing trunk capabilities), supporting vine potential assessment (wide ring patterns revealing maximum growth potential achievable under optimal conditions informing management expectations), and requiring contextual interpretation.
Wide_Rootstock_Annual_Ring emphasizes increased width greater than 0.6 cm and is distinguished from Rootstock_Annual_Ring (typical width) and Narrow_Rootstock_Annual_Ring (reduced width) by focus on vigorous growth reflecting favorable conditions, high vigor, or youth. Critical for understanding growth potential, favorable condition indicators, juvenile vigor patterns, capacity expansion, diagnostic value, management success markers, and the wide rings documenting optimal growth periods representing maximum xylem production during favorable years.
Breiter Unterlagen-Jahresring
Broad rootstock annual ring
Dicker Wurzelstock-Wachstumsring
Thick rootstock annual ring
Vigor ring
Vigoröser Ring
Wide Rootstock Annual Ring
A bush-trained Grenache vine at 1,100 vines/hectare (3.0 m × 3.0 m) in traditional Châteauneuf-du-Pape bears a Wide_Spacing_Role, the historic dryland spacing providing each vine with 9 m² of soil surface for water and nutrient access in the hot, dry Mediterranean climate. A Monastrell vine at 900 vines/hectare in Spanish Jumilla serves in a Wide_Spacing_Role reflecting centuries of adaptation to 300 mm annual rainfall, where closer spacing would create unsustainable competition for limited water. A Carignan vine at 1,400 vines/hectare in traditional Languedoc gobelet training bears a Wide_Spacing_Role, the extensive spacing enabling survival of ungrafted old vines on sandy phylloxera-resistant soils with minimal inputs.
A vine spacing role that inheres in a grapevine planted at very low density (typically <1,500 vines per hectare), with extensive spacing creating minimal inter-vine competition and maximum per-vine resources, characteristic of traditional dryland viticulture, extreme climate adaptation, or specific production systems emphasizing individual vine development. Wide spacing represents the extensive end of the planting density spectrum, historically common in Mediterranean dryland viticulture (parts of Spain, southern France, Mediterranean islands) where limited rainfall necessitated wide spacing to provide each vine with adequate water catchment area. Traditional bush-trained vineyards often feature wide spacing (2.5-3.5 m × 2.5-3.5 m, yielding 800-1,600 vines/hectare) that enables each self-supporting vine to develop its full canopy potential without inter-vine interference. Wide spacing is also associated with pergola systems (parral, tendone) where the overhead canopy architecture requires substantial space per vine for the spreading canopy to develop. Contemporary quality viticulture rarely establishes new wide-spaced vineyards, though heritage plantings with wide spacing persist and may be valued for their old-vine character and traditional production.
Wide_Spacing_Role captures the extreme extensive planting tradition, increasingly rare in new plantings but persisting in heritage vineyards valued for their old-vine character. The wide-spacing model reflects adaptation to resource-limited environments—each vine requires substantial soil volume to access adequate water and nutrients under dryland conditions—rather than a quality philosophy preference. Modern irrigation and fertigation have largely eliminated the resource constraints that necessitated wide spacing, enabling higher densities on sites that traditionally required extensive planting. Wide-spaced heritage vineyards are increasingly valued precisely for their rarity and traditional character, with the extensive spacing contributing to the low yields and vine individuality associated with old-vine quality. Preservation of wide-spaced vineyards represents cultural heritage conservation as much as viticultural choice, maintaining landscape aesthetics and production traditions that modernization would erase.
Trockenanbau-Abstand
dryland spacing
extensive Bepflanzung
extensive spacing
traditional spacing
traditioneller Abstand
weiter Abstand
wide spacing
Dry PR, Loveys BR (1998) Factors influencing grapevine vigour, Australian Journal of Grape and Wine Research 4: 140-149 Gladstones J (1992) Viticulture and Environment, Winetitles Huglin P, Schneider C (1998) Biologie et écologie de la vigne, Lavoisier
Wide Spacing Role
Grenache (loose-clustered selections) in Southern Rhône at BBCH 65 exhibits widely spaced loose flower clusters of 20-25 cm length and 15-20 cm width bearing 1000-1500 flowers distributed across 18-25 primary branches with 6-9 secondary and 2-4 tertiary branches, creating flower densities of 4-6 flowers/cm and filigree structures (OIV code 1) with essentially zero Botrytis at bloom but 20-25% shatter losses in windy conditions and 15-20% bird damage at harvest without netting. Mourvèdre in Bandol shows similarly extreme loose architecture (OIV code 1-2) with 800-1200 flowers on extensively branched inflorescences creating multi-lobed, compound clusters requiring 30-40% more harvest labor per ton due to low cluster weights (150-300 g) but achieving exceptional wine quality through perfect berry health and optimal phenolic ripening.
A widely spaced loose cluster is a loose flower cluster exhibiting very low flower density with flowers distributed on extensively branched axes with long internodes and extended pedicels, creating maximally open, lacy structures that represent the extreme of loose cluster architecture while potentially compromising pollination efficiency and creating challenges for uniform fruit set across the dispersed flower population. Widely spaced loose clusters exhibit flower densities typically below 5-8 flowers per cm of total inflorescence axis length, achieved through moderate-to-low flower initiation combined with maximal rachis and branch elongation, extensive secondary branching (5-10 secondary branches per primary), frequent tertiary branching (2-4 tertiary branches per secondary on basal primaries), and long pedicel length (6-12 mm). These clusters typically measure 15-25 cm in length and often 12-20 cm in maximum width at BBCH 65 while bearing 600-1500+ flowers, creating a dispersed three-dimensional flower distribution with extensive visible axis and air space dominating cluster volume. The extremely loose architecture results from genetic programming for maximal internode elongation during inflorescence development, combined with extensive hierarchical branching that creates the maximum spatial distribution of flowers observed in cultivated Vitis vinifera. Widely spaced loose flower clusters characteristically show compound shapes with pronounced lateral lobes, wings, or shoulders, often with width exceeding length (length=width ratios 0.8=1-1.2=1), creating a 'multi-winged' or 'filigree' appearance with flowers appearing as discrete elements within a branching framework rather than as a cohesive mass. Flower spacing within widely spaced loose clusters typically exceeds 8-15 mm between adjacent flower pedicels along any given axis, with individual flowers standing free and well-separated from neighbors. The extremely open flower arrangement creates a microclimate essentially identical to ambient conditions, with no humidity elevation within the cluster, instantaneous equilibration with ambient air after precipitation (drying time differences <30 minutes compared to loose clusters, <4 hours compared to compact clusters), and complete air circulation access to all flowers. This optimal microclimate essentially eliminates bloom-period Botrytis risk (<0.5% infection rate even in wet springs), allowing successful organic production with zero fungicide applications at flowering in most climates. However, the widely dispersed flower distribution creates significant challenges= (1) reduced pollination efficiency due to large distances between flowers reducing probability of pollen transfer, potentially lowering fruit set by 10-20% compared to moderately loose clusters (2) increased susceptibility to shatter during bloom, as unsheltered flower positions experience maximum wind exposure, potentially causing 15-30% flower abscission in windy conditions (3) increased difficulty in achieving uniform pollination across the extensive cluster volume, potentially creating significant berry size heterogeneity within clusters (4) maximum bird predation risk at maturity due to completely accessible berry positions. The transformation from widely spaced loose flower cluster to widely spaced loose berry cluster maintains extreme openness, with berry-berry contact typically absent even at full maturity, berries retaining perfect spherical morphology, harvest occurring as picking of individual berries rather than cluster cutting in some cases, and clusters appearing ornamental but providing minimal harvest efficiency (low weight per cluster, high picking time per unit yield).
Widely_Spaced_Loose_Cluster represents the extreme end of the loose spectrum and is distinguished from Open_Loose_Cluster by quantitative density (<5-8 flowers/cm versus 8-12 flowers/cm), maximal branching complexity (extensive tertiary branching versus moderate secondary branching), and specific advantages/disadvantages (eliminated disease risk versus compromised pollination and shatter susceptibility). This class corresponds specifically to OIV code 1 (maximum looseness) and represents significant viticultural trade-offs= exceptional disease resistance and sustainability but challenges in yield efficiency, pollination reliability, and bird management. The extremely loose architecture is characteristic of Mediterranean and warm-climate cultivars where Botrytis pressure historically drove selection for maximum cluster openness, but modern production in cooler climates with different pest complexes may favor moderately loose rather than extremely loose types.
extremely open flower bunch
sehr aufgelockerte Blütentraube
sehr lockere Blütentraube
very loose flower cluster
weit gestreute Blütentraube
Keller, M. (2020). The Science of Grapevines, 3rd ed., pp. 94-96 OIV (2009). Descriptor List for Grape Varieties and Vitis Species, OIV code 204 level 1 Galet, P. (2000). A Practical Ampelography, pp. 49-52 Intrieri, C., Poni, S. (1995). Integrated evolution of trellis training systems. American Journal of Enology and Viticulture 46=116-127.
Widely Spaced Loose Cluster
Leaf blade width in Chardonnay shows characteristic response to vine water status, with well-watered vines producing leaves of 150-180 mm width while deficit-irrigated vines (stem water potential below -1.2 MPa at midday) produce leaves of only 110-140 mm width, resulting in reduced total leaf area, lower whole-canopy photosynthesis, and modified fruit composition including higher phenolic concentration and more concentrated flavors.
Width_Quality represents the linear dimension measured perpendicular to the principal longitudinal axis of a grapevine organ, typically capturing the greatest transverse extent. Width measurements complement length data to characterize organ proportions, shape indices, and cross-sectional geometry. For leaves, width refers to the greatest distance across the blade perpendicular to the midrib, a key ampelographic character that distinguishes cultivars and responds to environmental conditions. Berry width, measured as the maximum equatorial diameter, determines fruit size perception and influences skin-to-pulp ratio. Cluster width characterizes bunch shape and compactness, affecting disease susceptibility and harvest logistics. Shoot and cane width (diameter) at standardized positions indicates vigor and carbohydrate storage capacity. Trunk width at defined heights tracks vine development and productive potential. Width measurements are particularly informative when expressed as ratios to length, yielding shape indices that describe organ geometry independently of absolute size. A length-to-width ratio near 1.0 indicates globose form, while higher ratios indicate elongation. Leaf blade width relative to length distinguishes orbicular, cuneiform, and reniform blade shapes. Environmental factors modulate width expression water deficit typically reduces width more than length in expanding organs, yielding higher length-to-width ratios. Temperature affects cell division patterns that establish organ width during early development. Standardized width measurements require definition of the measurement axis and reference landmarks appropriate to each organ type.
Width_Quality is a sibling class to Length_Quality under Linear_Dimension_Quality, distinguished by its perpendicular measurement axis. For organs with circular cross-section (internodes, roots), width equals diameter Diameter_Quality is used as the preferred term in such cases. The leaf blade width measurement specified in OIV descriptor 065 exemplifies standardized width assessment. Shape_Quality classes utilize width measurements as inputs for ratio-based shape descriptors. PATO_0000921 (width) provides upper-ontology alignment.
Breite
Breitenmessung
Querdimension
Transversalausdehnung
breadth
cross-dimension
lateral extent
transverse dimension
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 52-55 Maul E et al. (2012) Vitis International Variety Catalogue. Julius Kühn-Institut, Federal Research Centre for Cultivated Plants
Width Quality
Vitis riparia accessions from Minnesota natural populations bear Wild_Vitis_Genetic_Resource_Roles, with extreme cold hardiness (-40°C survival), early ripening, and phylloxera resistance traits valuable for both rootstock and cold-climate scion breeding. Muscadinia rotundifolia from southeastern US populations serves in a Wild_Vitis_Genetic_Resource_Role as the source of Run1 and Ren1 powdery mildew immunity—qualitatively different from V. vinifera partial resistance and central to contemporary PIWI breeding. Vitis amurensis accessions from Chinese and Russian collections bear Wild_Vitis_Genetic_Resource_Roles for cold hardiness introgression, with -40°C bud survival and Rpv10-12 downy mildew resistance loci contributing to continental climate variety development.
A genetic resource role that inheres in grapevine genotypes representing wild (uncultivated) Vitis species, maintained in natural populations, botanical collections, or germplasm repositories as sources of genetic diversity unavailable in cultivated V. vinifera. Wild Vitis species (approximately 60-70 species worldwide) evolved resistance mechanisms through millions of years of coevolution with pathogens and pests—resistance largely absent from V. vinifera which evolved in geographic isolation from these selective pressures. North American Vitis species (V. riparia, V. rupestris, V. berlandieri, V. cinerea, V. aestivalis, V. labrusca, V. rotundifolia/Muscadinia) provide the resistance genes underlying rootstock breeding and contemporary PIWI development, while Asian species (V. amurensis, V. coignetiae, V. davidii) contribute cold hardiness and additional resistance sources. Wild Vitis genetic resources face conservation challenges—habitat loss, climate change, and in some regions, hybridization with cultivated or feral vines threatening wild population integrity. In situ conservation (protected natural populations) complements ex situ preservation (germplasm repositories) to maintain wild Vitis diversity for current and future breeding access.
Wild_Vitis_Genetic_Resource_Role captures the fundamental diversity reservoir underlying all resistance breeding in grapevines. V. vinifera's susceptibility to phylloxera, powdery mildew, downy mildew, and Pierce's disease—pathogens that evolved with American Vitis species—necessitates wild species genetic contribution for effective resistance. Wild Vitis diversity is threatened by multiple factors: habitat fragmentation and destruction, potential climate change impacts on distribution, and gene flow from cultivated or feral grapes into wild populations. Conservation efforts include in situ protection (natural reserves, identified populations), ex situ collection (germplasm repositories maintaining living accessions), and increasingly, genomic characterization providing a molecular inventory of diversity. Access to wild Vitis resources is governed by both national regulations (collection permits, export restrictions) and international agreements (Convention on Biological Diversity, Nagoya Protocol) affecting breeding program access to genetic diversity.
Wildreben-Genressource
Wildvitis-Ressource
uncultivated Vitis
undomestizierte Vitis
wild Vitis resource
wild grape genetic resource
wild species germplasm
wilde Vitis-Arten
Riaz S et al. (2019) Genetic resources for grape improvement, Springer pp.: 25-51 Heinitz CC et al. (2019) Crop wild relatives of grape in North America, in Greene SL et al. (eds) North American Crop Wild Relatives Vol 2, Springer pp.: 329-351 This P et al. (2006) Historical origins of wine grapes, Trends in Genetics 22: 511-519 Wan Y et al. (2013) Wild grape genetic resources, PLoS ONE 8: e56093
Wild Vitis Genetic Resource Role
Wind data for the Salinas Valley (Monterey, California) documents persistent afternoon northwesterly winds averaging 20-30 km/h during June-September, channeled through the valley's funnel topography from Monterey Bay, depressing growing season temperatures by 3-5°C relative to inland sites and extending the Pinot Noir and Chardonnay ripening period by 2-4 weeks. The Mistral in the southern Rhône Valley (Châteauneuf-du-Pape) blows from the north-northwest at 40-60 km/h for 100-150 days per year, desiccating canopies and dramatically reducing Botrytis incidence while requiring gobelet training to withstand mechanical forces. Wind data for Burgenland, Austria records dominant northwesterly winds across the Neusiedlersee, promoting autumn fog formation critical for Botrytis development on Furmint and Welschriesling for Ausbruch and Trockenbeerenauslese production.
Wind data is a terroir information entity that documents the velocity, direction, frequency, turbulence, and seasonal patterns of atmospheric air movement at a viticultural site, serving as a determinant of canopy evapotranspiration rates, disease pressure modulation, mechanical stress on vine structures, and the mesoclimatic temperature regime through advective heat transport and cold air drainage dynamics.
Wind data integrates multiple measurement parameters — mean speed, gust frequency and intensity, directional constancy, diurnal and seasonal patterns, and vertical wind profiles — each carrying distinct viticultural implications. Wind speed below 5 km/h is generally insufficient to maintain boundary layer renewal at leaf surfaces, while sustained speeds above 30 km/h cause stomatal closure, physical damage to shoots and clusters, and excessive transpiration stress. The aerodynamic roughness of vineyard landscapes (determined by training system height, row orientation, and surrounding topography) modulates the effective wind speed experienced by individual vines. Windbreak design (natural or artificial) represents a key management response to excessive wind exposure, though the beneficial effects of moderate wind on disease suppression and canopy drying must be balanced against protection needs. Row orientation relative to prevailing wind direction influences both wind penetration into the canopy interior and the mechanical loading on trellis structures.
Winddaten
Windinformation
Windklimatologie
Windverhältnisse
anemometric data
anemometrische Daten
vineyard wind data
wind climate data
wind information
wind record
Gladstones J (2011) Wine, Terroir and Climate Change, Wakefield Press, pp.: 145-160 Smart RE, Robinson MD (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles, pp.: 23-30 Jackson RS (2020) Wine Science: Principles and Applications, 5th Ed, Academic Press, pp.: 72-85
Wind Data
Wind speed data for Casablanca Valley, Chile records mean growing season wind speeds of 12-18 km/h from the Pacific coast, with afternoon maxima of 25-35 km/h that induce partial stomatal closure in Sauvignon Blanc and Chardonnay, limiting photosynthetic rates but enhancing aromatic thiol precursor accumulation through moderate water stress. Coonawarra, South Australia experiences relatively calm conditions with mean speeds of 8-12 km/h, insufficient to cause vine stress but adequate for canopy boundary layer renewal. The Etesian winds (Meltemi) in Santorini, Greece generate sustained speeds of 30-50 km/h during July-August, necessitating the traditional kouloura (basket-trained) vine architecture that shields clusters within a low, ground-hugging canopy ring, protecting Assyrtiko berries from physical damage and excessive desiccation.
Wind speed data is a wind data entity that records the velocity of atmospheric air movement at a viticultural site, measured at standardized anemometer height (typically 2 m or 10 m above ground) and expressed in meters per second (m/s) or kilometers per hour (km/h), encompassing mean speed, maximum sustained speed, gust intensity, and temporal variability parameters that collectively characterize the mechanical and physiological wind loading experienced by grapevines.
Wind speed measurement at viticultural sites requires careful attention to sensor height, exposure, and averaging period, as standard meteorological stations at 10 m height may substantially overestimate wind conditions at vine canopy level (typically 1-2 m). The logarithmic wind profile allows conversion between measurement heights using surface roughness parameters specific to vineyard training systems. Key viticultural thresholds include 2-5 km/h (minimum for effective canopy drying), 15-25 km/h (optimal range for transpiration and disease suppression without stomatal closure), and >35 km/h (risk of mechanical damage to shoots, tendrils, and developing clusters). Gust speeds exceeding 60 km/h can cause catastrophic shoot breakage, trellis failure, and crop loss. Wind speed variability is equally important — diurnal patterns with calm mornings and windy afternoons create different physiological responses than constant moderate winds, as vines exhibit hydraulic adjustment during sustained wind events but are more vulnerable to sudden onset gusts.
Anemometerdaten
Windgeschwindigkeitsdaten
Windgeschwindigkeitsmessung
Windstärkedaten
anemometer data
wind force data
wind speed measurement
wind speed record
wind velocity data
Smart RE, Robinson MD (1991) Sunlight into Wine, Winetitles, pp.: 24-29 Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 65-72 Campbell-Clause JM (1998) Stomatal response of grapevines to wind. Australian Journal of Experimental Agriculture 38(1): 77-82
Wind Speed Data
Wine analysis data for Grüner Veltliner Smaragd from the Wachau (2024 vintage, post-bottling) records the complete analytical profile: alcohol 13.5% vol., residual sugar 2.8 g/L (dry), total acidity 5.8 g/L (as tartaric acid), volatile acidity 0.32 g/L (as acetic acid), free SO₂ 28 mg/L, total SO₂ 95 mg/L, and pH 3.22 — all parameters within the regulatory limits defined by the Austrian Weingesetz and EU Regulation 2019/934, and consistent with the Vinea Wachau Smaragd classification requirements. Wine analysis data for Cabernet Sauvignon from Rutherford, Napa Valley (2022 vintage, 24 months post-bottling) documents alcohol 14.6% vol., residual sugar 1.2 g/L, TA 5.5 g/L, VA 0.58 g/L, free SO₂ 32 mg/L, total SO₂ 118 mg/L, and pH 3.62 — establishing the wine within the typical compositional envelope for premium Napa Cabernet and confirming stability for extended cellar aging.
Wine analysis data is a wine specification entity that documents the standard analytical characterization of finished wine through the measurement of key compositional parameters required for regulatory compliance, quality control, labeling accuracy, and commercial transaction, encompassing alcohol content, residual sugar, total acidity, volatile acidity, free and total sulfur dioxide, and pH, determined through officially recognized analytical methods (OIV, AOAC, national reference methods) on the finished product prior to or following bottling, and providing the objective chemical identity card that defines the wine's regulatory category, stability status, and fundamental compositional profile.
Wine analysis data represents the regulatory and commercial minimum analytical characterization required for virtually all wines entering the market, with most wine-producing jurisdictions mandating laboratory analysis certificates for quality wine approval, export documentation, and label claim verification. The seven parameters documented in this hierarchy — alcohol, residual sugar, total acidity, volatile acidity, free SO₂, total SO₂, and pH — constitute the universally recognized core analytical panel, though many jurisdictions require additional determinations (extract, ash, sulfate, citric acid, sorbic acid, carbon dioxide pressure for sparkling wines) for specific wine categories. The distinction between wine analysis data (finished product characterization) and must analysis data (pre-fermentation characterization, documented in the Must_Analysis_Data hierarchy) is ontologically significant — must analysis informs production decisions, while wine analysis documents product compliance and quality outcomes. Several parameters appear in both hierarchies (Total_Acidity, pH) but carry different interpretive significance and reference ranges in must versus wine contexts — must TA reflects grape acidity for harvest decisions, while wine TA reflects the post-fermentation acid balance after MLF, precipitation, and any permitted adjustments. The analytical methods specified by the OIV Compendium of International Methods and adopted by the EU as reference methods provide the internationally harmonized framework for wine analysis, though national methods (AOAC in the USA, methods specified in the Weinverordnung in Austria and Germany) may differ in procedural details. The increasing deployment of FTIR spectroscopy (WineScan and similar instruments) enables rapid multi-parameter screening of all seven core parameters in a single 30-second measurement, though regulatory and trade dispute resolution still requires confirmation by the official reference methods.
Weinanalysedaten
Weinchemieanalyse
Weinuntersuchungsdaten
Weinzusammensetzungsdaten
analytisches Weinprofil
finished wine analysis
wine analytical profile
wine chemical analysis
wine composition data
wine laboratory data
OIV (2021) Compendium of International Methods of Analysis of Wines and Musts, International Organisation of Vine and Wine European Parliament (2019) Regulation (EU) 2019/934 supplementing Regulation (EU) No 1308/2013 as regards oenological practices Iland P et al. (2011) Chemical Analysis of Grapes and Wine, 2nd Ed, Patrick Iland Wine Promotions, pp.: 120-175
Wine Analysis Data
Thompson Seedless (Sultana, Sultanina) bears a Wine_And_Raisin_Dual_Purpose_Role as the archetypal dual-purpose variety, with California production divided among raisin (primary use at 22-24°Brix), table grape (early harvest at 16-18°Brix), and wine/juice (sorted-out fruit, excess production) markets—annual allocation responding to market prices, with growers shifting substantially between categories across seasons. Muscat of Alexandria serves in a Wine_And_Raisin_Dual_Purpose_Role for Muscatel raisin production (retaining distinctive aromatic character through drying) and Muscat wine production (sweet and dry styles) in Mediterranean, South African, and Australian production regions. Zante Currant (Black Corinth) bears a Wine_And_Raisin_Dual_Purpose_Role in Greek production, with primary currant production complemented by traditional sweet wine production (Vino di Corinto) from the same tiny, intensely flavored berries.
A multi-purpose scion role that inheres in a grafted scion cultivar by virtue of its commercially significant utilization for both wine production and dried grape (raisin) production, reflecting characteristics suitable for both fermented and dehydrated products. Wine-and-raisin dual-purpose scions must achieve sugar levels adequate for raisin production (22-26°Brix) while maintaining the acidity and varietal character valued in wine, with allocation between markets determined by harvest timing, vintage conditions, and relative prices. Early-harvested fruit (18-22°Brix) may be directed to wine or juice production, while fully mature fruit (22-26°Brix) can be dried for raisins or pressed for high-alcohol wines and distillation base. Thompson Seedless exemplifies this dual-purpose role, with California production annually divided between table, raisin, and wine/juice markets based on quality sorting and price signals. The wine-raisin combination reflects shared requirements for high sugar accumulation in warm, dry climates where both products concentrate successfully.
Wine_And_Raisin_Dual_Purpose_Role captures the shared warm-climate, high-sugar production system where varieties achieve concentrations suitable for either drying or fermentation into high-alcohol wines. The economics typically favor raisins when prices are strong (higher per-kilogram value), with wine production absorbing lower-grade fruit or surplus tonnage. Thompson Seedless dominates this category by volume, though its neutral character produces unremarkable wines suited primarily for brandy distillation or blending. More flavorful varieties like Muscat of Alexandria maintain genuine dual-purpose value where both raisin and wine products express distinctive varietal character. Climate conditions during harvest determine optimal allocation—rain during the raisin drying window may force fruit into wine production, while ideal drying weather maximizes raisin quality and returns.
Doppelnutzungs-Trockentraube
Wein- und Rosinentraube
Wein-Trocken-Traube
dual-purpose drying grape
wine and raisin grape
wine-raisin variety
Christensen LP (2000) Raisin Production Manual, UC ANR Creasy GL, Creasy LL (2009) Grapes, CABI Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane California Raisin Marketing Board (2020) Industry Statistics
Wine And Raisin Dual Purpose Role
Italia (Ideal, Muscat Italia) bears a Wine_And_Table_Dual_Purpose_Role in southern Europe, serving premium table grape markets (large berries, muscat aroma, crisp texture) while also producing aromatic white wines in Sicily, Puglia, and other Mediterranean regions where its Muscat heritage contributes distinctive terpene-rich character to both fresh and fermented products. Chasselas (Fendant, Gutedel) serves in a Wine_And_Table_Dual_Purpose_Role as Switzerland's historic dual-purpose variety—traditionally the finest Swiss table grape, now rare in fresh markets but remaining the foundation of Valais and Vaud white wine production (Fendant) with delicate, mineral character. Cardinal bears a Wine_And_Table_Dual_Purpose_Role in some Mediterranean regions, with early-ripening red grapes serving both fresh consumption (table grape) and early-drinking red wine production, though quality limitations restrict both uses to local markets.
A multi-purpose scion role that inheres in a grafted scion cultivar by virtue of its commercially significant utilization for both wine production and fresh table grape consumption, reflecting characteristics suitable for both fermented and fresh markets. Wine-and-table dual-purpose scions must satisfy divergent requirements—wine production favoring smaller berries (higher skin-to-juice ratio for phenolic extraction), concentrated flavors, and specific sugar-acid balance for fermentation, while table grape production favors larger berries, crisp texture, attractive appearance, and eating quality. Historically, many V. vinifera varieties served dual purposes before market specialization contemporary dual-purpose utilization persists where traditional preferences value particular varieties for both uses, where market infrastructure supports flexible allocation, or where breeding has achieved genuine excellence in both categories. Regional examples include Italia in southern Europe (premium table grape and aromatic wine), Chasselas in Switzerland (table grape tradition and Fendant wine), and Muscat varieties globally (table, wine, and other uses).
Wine_And_Table_Dual_Purpose_Role captures the intersection of two major grape utilization categories where specific varieties maintain commercial relevance in both markets. Modern market evolution has generally driven specialization—table grape breeding focuses on seedlessness, size, and shipping durability while wine grape breeding emphasizes phenolic concentration and terroir expression—reducing the overlap between categories. However, certain varieties (particularly Muscats and traditional European cultivars) retain genuine dual-purpose value where regional traditions support both uses. The economic logic differs between markets table grapes command higher per-kilogram prices but require more intensive production practices, while wine grapes generate value through processing and branding rather than fresh sales.
Doppelnutzungs-Sorte
Tafelwein-Traube
Wein- und Tafeltraube
dual-purpose wine grape
fresh and fermented variety
wine and table grape
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes, Ecco/Allen Lane Creasy GL, Creasy LL (2009) Grapes, CABI Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press Ferrara G et al. (2017) Italia table grape, ISHS Acta Horticulturae 1157: 365-370
Wine And Table Dual Purpose Role
Wine pH for Riesling from the Wachau Smaragd (2024 vintage) measures 3.22, an optimally low value for dry white wine that maximizes molecular SO₂ efficacy (3.8% of free SO₂ in molecular form), ensures brilliant color preservation, and provides an inherently hostile environment for spoilage organisms (Brettanomyces dekkera growth is strongly inhibited below pH 3.3). Wine pH for Châteauneuf-du-Pape (2022 vintage) records 3.78, near the upper boundary of acceptable range for red wine, where the high pH necessitates elevated free SO₂ levels (45 mg/L to achieve 0.5 mg/L molecular SO₂), increases susceptibility to Brettanomyces and lactic acid bacteria spoilage, and accelerates the rate of tannin-anthocyanin polymerization and browning reactions during aging. Wine pH for Blaufränkisch Reserve from Eisenberg DAC (2022 vintage) documents 3.35, within the optimal range (3.2-3.5) for age-worthy Austrian red wine, balancing adequate microbial protection with sufficiently soft acid perception for the 12+ year aging trajectory intended for this bottling.
Wine pH is a wine analysis data entity that records the hydrogen ion activity in finished wine as determined by potentiometric measurement using a calibrated glass electrode pH meter at 20°C, expressed on the logarithmic pH scale (dimensionless units), and representing the master variable of wine chemistry that governs the efficacy of sulfur dioxide protection, the equilibrium state of anthocyanin color expression, the activity and survival of spoilage microorganisms, the rate of chemical aging reactions, the perception of acid taste, and the overall chemical stability trajectory of the wine from bottling through decades of cellar aging.
Wine pH measured on the finished product carries both continuity with and important differences from must pH (documented in the Must_Analysis_Data hierarchy, Batch 66), as multiple intervening processes modify the acid-base equilibrium between harvest and bottling. Malolactic fermentation raises wine pH by 0.1-0.3 units through the conversion of the stronger diprotic malic acid to the weaker monoprotic lactic acid. Potassium bitartrate precipitation removes both potassium and tartrate ions from solution, with a net pH effect that depends on the relative amounts precipitated — typically a modest pH decrease of 0.05-0.15 units. Cold stabilization (chilling to -4°C to precipitate KHT before bottling) further modifies pH, while acid additions (permitted tartaric acid addition in warm regions) or deacidification (permitted calcium carbonate or potassium bicarbonate addition in cool regions) represent deliberate pH adjustments. The wine pH value integrates all these processes and represents the final acid-base state that will govern the wine's chemical evolution during storage and aging. The cascading effects of pH on wine chemistry cannot be overstated — a difference of 0.3 pH units (e.g., 3.3 versus 3.6) halves the molecular SO₂ fraction, doubles the rate of many oxidation reactions, shifts the anthocyanin equilibrium substantially toward colorless forms, and fundamentally alters the wine's microbial vulnerability profile. The climate change-driven trend toward higher wine pH (documented at 0.02-0.05 pH units per decade in many European regions) represents one of the most significant quality challenges for contemporary winemakers, as historically optimal pH ranges for traditional wine styles may become increasingly difficult to achieve without intervention. Measurement precision is critical — a calibration error of ±0.05 pH units propagates to a 12-15% error in molecular SO₂ calculation, potentially leading to inadequate microbial protection.
Wasserstoffionenaktivität des Weines
Wein pH-Wert
Wein-pH
pH of wine
pH-Messung des Weines
pH-Wert des Weines
wine acid-base status
wine acidity pH
wine hydrogen ion activity
wine pH value
Waterhouse AL et al. (2016) Understanding Wine Chemistry, Wiley, pp.: 15-35 OIV (2021) Compendium of International Methods of Analysis, Method OIV-MA-AS313-15 (pH) Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 2, 2nd Ed, Wiley, pp.: 3-15 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 330-345
Wine pH
Wine price data for Riesling illustrates the cultivar's exceptional quality-price range — from €3-5 for Mosel Qualitätswein at German retail, through €10-25 for Spätlese and Erste Lage bottlings, €30-80 for Grosse Lage Grand Cru-equivalent dry Riesling from top estates (Dönnhoff, Keller, Weil), to €200-1,500+ for Trockenbeerenauslese at auction (historic TBA from celebrated vintages reaching €5,000-15,000 per half-bottle at major auction houses) — the price spectrum reflecting both the breadth of Riesling production from entry-level to ultra-premium and the extraordinary aging capacity that sustains collector interest in library vintages. Wine price data for Blaufränkisch from Mittelburgenland DAC documents an evolving premium trajectory — entry-level DAC Classic at €8-12 retail, Reserve and single-vineyard bottlings at €18-40, and top cuvées from leading estates (Iby, Gesellmann, Heinrich) reaching €40-80 — a price structure that has expanded substantially over the past two decades as critical recognition and export market development have validated the cultivar's quality potential.
Wine price data is a market data entity that documents the prices at which finished wines are traded and sold across market channels, encompassing ex-cellar prices (the producer's selling price to the first commercial buyer), wholesale-distributor prices (trade channel pricing between winery and retailer), retail prices (consumer purchase prices at point of sale — wine shops, supermarkets, online retailers), and auction prices (realized prices at fine wine auctions for collectible and library wines), differentiated by cultivar, producer, appellation, vintage, quality tier, and market channel, derived from trade price lists, retail tracking databases, auction records, and commercial price aggregation platforms, providing the value realization data that quantifies the commercial return on viticultural and oenological investment.
Wine price data captures the commercial value realization at successive stages of the distribution chain — each price level reflecting not only the intrinsic quality of the wine but the accumulated value of branding, critical acclaim, appellation prestige, vintage reputation, and market positioning that transforms an agricultural product into a cultural-commercial object. The price formation mechanisms differ dramatically across market channels — ex-cellar prices are set by producers based on production costs, quality positioning, and market strategy, wholesale prices reflect the trade margin structure and competitive dynamics of the distribution sector, retail prices incorporate retailer margins and are influenced by shelf positioning, promotional activity, and competitive context, and auction prices are determined by collector demand for scarce, prestigious, or aged wines in a transparent bidding environment. The fine wine auction market provides the most visible and best-documented wine pricing data — major houses (Christie's, Sotheby's, Acker Merrall & Condit, Hart Davis Hart) publish detailed results for every lot, and the Liv-ex exchange provides continuous secondary market pricing for actively traded fine wines — but this data represents only the uppermost tier of the wine market, covering perhaps 1-2% of global wine production by volume. The vast majority of wine transactions occur at retail through channels ranging from specialized wine merchants through supermarket shelves to online platforms, with pricing data aggregated by commercial services (Wine-Searcher, Vivino) that provide increasingly comprehensive but still incomplete coverage of global retail pricing. The relationship between wine prices and quality ratings has been extensively studied — the correlation is positive but modest (typically r = 0.2-0.4 between critic scores and prices within a category), with substantial price variation unexplained by quality scores and attributable to brand reputation, appellation prestige, scarcity, and collector speculation.
Einzelhandelspreis
Flaschenpreis
Weinhandelswert
Weinpreisdaten
bottle price
retail price data
wine commercial value
wine pricing
wine valuation
Liv-ex (2023) Fine Wine Market Report and Indices Wine-Searcher (2023) Global Wine Price Database Ashenfelter O (2008) Predicting the quality and prices of Bordeaux wine. The Economic Journal 118(529): F174-F184 Storchmann K (2012) Wine economics. Journal of Wine Economics 7(1): 1-33
Wine Price Data
Chardonnay bears a Wine_Production_Scion_Role across its global distribution from Burgundy to California to Australia, selected for its neutral-to-buttery flavor profile, malolactic fermentation suitability, and adaptability to diverse winemaking styles from lean Chablis to opulent Napa. Tempranillo serves in a Wine_Production_Scion_Role throughout Iberia, providing the tannic structure, cherry-tobacco aromatics, and oak-aging capacity defining Rioja and Ribera del Duero. Sauvignon Blanc bears a Wine_Production_Scion_Role in Loire, Bordeaux, Marlborough, and globally, contributing the thiolic aromatics (grapefruit, passionfruit, boxwood) that define the variety's distinctive sensory profile.
A scion role that inheres in a grafted scion cultivar by virtue of its designation for wine production, where the harvested berries are crushed, fermented, and processed into wine. Wine production scions are selected and cultivated to optimize berry composition for vinification, including sugar accumulation sufficient for target alcohol levels (typically 11-15% v/v requiring 180-260 g/L sugar at harvest), appropriate organic acid concentrations (5-9 g/L total acidity) for microbial stability and sensory balance, phenolic compound profiles (anthocyanins, tannins, flavonols) contributing color, structure, and mouthfeel, and aroma precursor accumulation (terpenes, thiols, norisoprenoids, methoxypyrazines) defining varietal character. Wine production scions differ from table grape scions in prioritizing extractable compounds and flavor intensity over berry size, appearance, and textural qualities for fresh consumption. The category encompasses red, white, rosé, sparkling, fortified, and dessert wine production across the full spectrum from bulk commodity wines to ultra-premium collectibles.
Wine_Production_Scion_Role is the predominant scion role globally, with wine grape production accounting for approximately 55% of global grape production by volume (the remainder being table grapes, raisins, and juice). The role is subclassed by wine color/style (Red, White, Rosé, Sparkling, Fortified, Dessert, Orange) reflecting the distinct cultivar attributes and viticultural practices associated with each wine category. A single cultivar may bear multiple wine production sub-roles Grenache serves in Red_Wine, Rosé_Wine, and Fortified_Wine scion roles depending on regional tradition and winemaker intent. Wine production scion selection integrates cultivar-climate matching, terroir expression, market demand, and regulatory constraints (appellation rules often specify permitted varieties).
Keltersorte
Keltertraubensorte
Weinrebe
Weintraube
vinification variety
wine cultivar
wine grape scion
wine variety
Robinson J, Harding J, Vouillamoz J (2012) Wine Grapes: A Complete Guide to 1,368 Vine Varieties, Ecco/Allen Lane Jackson RS (2014) Wine Science: Principles and Applications 4th Ed, Academic Press Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp.: 343-430
Wine Production Scion Role
Wine specification for Cabernet Sauvignon clone 337 from Rutherford, Napa Valley encompasses still wine specification for a full-bodied, oak-aged red (14.0-14.8% alcohol, 24-28 months in French oak, 65% new), classified as Napa Valley AVA Cabernet Sauvignon, with secondary blending use in Meritage-style proprietary blends at 60-75% inclusion, and no sparkling, fortified, or sweet wine suitability documented for this clone-site combination. Wine specification for Muscat Blanc à Petits Grains from Rutherglen encompasses four distinct wine type applications from a single cultivar: still wine (dry Muscat, 12.5% alcohol), sparkling wine (Asti-method low-pressure frizzante), fortified wine (Rutherglen Muscat, spirit-fortified to 18% alcohol with extended oxidative aging), and sweet wine (late harvest, 11% alcohol with 80 g/L residual sugar) — demonstrating the exceptional oenological versatility of this cultivar across wine categories.
Wine specification is a grapevine information entity that documents the oenological end-use characterization of grape cultivars, clones, and vineyard sites in terms of the wine types, styles, and quality categories for which the fruit is suited or intended, encompassing wine style descriptions, technical winemaking parameters, regulatory classification eligibility, and the relationship between viticultural inputs and wine product outcomes, providing the teleological framework that connects grape production to the commercial wine products derived from the harvested fruit.
Wine specification represents the ontological bridge between the viticultural domain (grape production, cultivar characterization, site assessment) and the oenological domain (wine production, product definition, market classification), encoding the information about what types of wine can or should be made from specific grape material. This entity class is essential for VVO because grapevine ontology without reference to wine product outcomes would lack the teleological dimension that gives viticultural practices their purpose — varieties are planted, clones selected, and vineyards managed with specific wine products in mind, and the specification of these intended products provides the evaluative framework for assessing viticultural success. The wine specification hierarchy distinguishes between the major wine categories recognized by international trade and regulatory frameworks — still wines (the dominant category by volume and diversity), sparkling wines (requiring specific base wine characteristics), fortified wines (involving spirit addition and extended maturation), and sweet wines (encompassing late harvest, botrytized, ice wine, and dried grape styles) — each imposing distinct requirements on grape composition, cultivar suitability, and viticultural management. The relationship between grape material and wine specification is many-to-many — a single cultivar may produce multiple wine types (Riesling for dry, sweet, sparkling, and ice wine) while a single wine specification may accept multiple cultivars (Champagne permits Chardonnay, Pinot Noir, and Pinot Meunier). Wine specification data should reference applicable regulatory frameworks (EU wine categories, national appellation systems, geographical indication requirements) that constrain the permissible wine types from specific regions and cultivars.
Weinherstellungsspezifikation
Weinproduktbeschreibung
Weinspezifikation
Weintypbeschreibung
oenological specification
wine category data
wine product specification
wine production specification
wine type data
önologische Spezifikation
OIV (2022) International Code of Oenological Practices, International Organisation of Vine and Wine Robinson J, Harding J (2015) The Oxford Companion to Wine, 4th Ed, Oxford University Press, pp.: 813-818 Jackson RS (2020) Wine Science: Principles and Applications, 5th Ed, Academic Press, pp.: 1-45
Wine Specification
Wine style description for Riesling from the Mosel encompasses a remarkable stylistic spectrum from a single cultivar and region: Kabinett feinherb (light-bodied, 8-9% alcohol, 20-30 g/L residual sugar, unoaked, early-drinking freshness), Spätlese trocken (medium-bodied, 11.5-12.5% alcohol, dry with concentrated minerality), Auslese (medium-bodied, 8-9% alcohol, 60-90 g/L residual sugar, nobly sweet), and Sekt (traditional method sparkling, 12% alcohol, brut dosage, 3+ years on lees) — the stylistic range enabled by Riesling's uniquely high natural acidity that maintains structural balance across all sweetness levels. Wine style description for Blaufränkisch from Mittelburgenland DAC specifies a medium-to-full-bodied dry red wine (13.0-14.0% alcohol, ≤4 g/L residual sugar), aged 12-18 months in large Austrian oak casks (Stockinger or similar, 500-2500 L), with the DAC regulation mandating varietal character of dark fruit, fine-grained tannin, and spice without excessive new oak influence — exemplifying how appellation regulations codify wine style descriptions.
Wine style description is a wine specification entity that documents the sensory, compositional, and production-method characteristics defining the intended wine product category derived from specific grape material, encompassing the four principal wine types (still, sparkling, fortified, sweet) and their sub-classifications by color (white, rosé, red, orange), body (light, medium, full), residual sugar (dry, off-dry, medium-sweet, sweet), production method (traditional, tank, ancestral for sparkling oxidative, reductive for maturation), and aging regime (unoaked, oak-aged, extended cask, bottle-aged), providing the categorical framework that links viticultural fruit characteristics to the diversity of wine products in the global marketplace.
Wine style description provides the categorical vocabulary through which the diversity of wine products is organized, communicated, and regulated — a vocabulary that has evolved over centuries through the intersection of regional tradition, consumer preference, regulatory codification, and winemaking technology. The hierarchy of wine style categories is not purely taxonomic but reflects fundamentally different production processes — still wine production involves standard alcoholic fermentation, sparkling wine requires a secondary fermentation for carbonation, fortified wine involves spirit addition during or after fermentation, and sweet wine demands specific mechanisms for sugar concentration (late harvest, botrytis, desiccation, freezing) or arrested fermentation. Within each major category, the sub-classification by color, body, sweetness, and aging regime creates a multidimensional style space — for example, still white wine alone encompasses styles from crisp, unoaked Muscadet (light body, bone dry, no oak) through rich, barrel-fermented Meursault (full body, dry, 12+ months new oak) to unctuous Sauternes (full body, intensely sweet, 18+ months new oak), each requiring fundamentally different fruit characteristics and viticultural management. The relationship between grape composition and achievable wine style is constrained but not deterministic — high-acid, moderate-sugar grapes (cool-climate Riesling, Chenin Blanc) offer the widest stylistic range because the acid backbone supports both dry and sweet styles, while high-sugar, low-acid grapes (warm-climate Grenache, Mourvèdre) are largely restricted to dry or fortified styles. The global trend toward geographical indication and appellation systems (EU PDO/PGI, US AVA, Australian GI) increasingly codifies wine style descriptions into regulatory specifications that constrain permissible cultivars, production methods, and compositional parameters for wines bearing specific origin designations.
Stilklassifizierung
Weinkategorisierung
Weinstilbeschreibung
oenological style description
wine category description
wine product type
wine style classification
wine type description
Robinson J, Harding J (2015) The Oxford Companion to Wine, 4th Ed, Oxford University Press, pp.: 813-818 Jackson RS (2020) Wine Science, 5th Ed, Academic Press, pp.: 1-45 OIV (2022) International Code of Oenological Practices Halliday J, Johnson H (2014) The Art and Science of Wine, Mitchell Beazley, pp.: 200-250
Wine Style Description
Winemaking protocol for Pinot Noir from Gevrey-Chambertin (Premier Cru) specifies the complete vinification sequence: 100% destemming with 5% whole-cluster inclusion, 5-day cold soak at 8°C in open-top oak fermenters (25 hL), inoculation with RC212 yeast at 18°C, fermentation peak temperature 32°C with twice-daily punch-down, total skin contact 16-18 days, free-run and light press wine separated, malolactic fermentation in barrel (228 L Burgundy pièces, 30% new François Frères and Rousseau cooperage), aging 16 months on fine lees with quarterly racking, assemblage from 3 vineyard parcels in January, egg-white fining at 3 g/hL, unfined and unfiltered bottling in March. Winemaking protocol for Grüner Veltliner Federspiel from the Wachau documents whole-cluster pressing (pneumatic, maximum 1.8 bar), 12-hour cold settling at 10°C, racking of clear juice to stainless steel (80%) and neutral 2500 L Stockinger cask (20%), fermentation with indigenous yeast at 16-18°C over 3-4 weeks, aging 6 months on full lees with monthly bâtonnage, blending of steel and cask portions in April, cross-flow filtration, and bottling in May.
Winemaking protocol is a wine specification entity that documents the standardized or site-specific sequence of oenological operations, process parameters, and technical decisions applied to transform harvested grape material into finished wine, encompassing fermentation management, maceration regime, aging treatment, and blending strategy, with each protocol element specifying the method employed, the target parameters and acceptable ranges, the equipment and inputs used, and the decision criteria governing protocol modifications in response to vintage variation, providing the procedural framework that translates grape quality potential into realized wine quality.
Winemaking protocol documentation serves as the operational bridge between viticultural grape quality assessment and the realized quality of the finished wine, encoding the accumulated empirical knowledge and deliberate aesthetic choices that distinguish individual producers, regional traditions, and quality tiers. The protocol concept is inherently hierarchical — a complete winemaking protocol comprises sub-protocols for each major processing phase (fermentation, maceration, aging, blending), each of which contains multiple decision points where alternative approaches produce fundamentally different wine outcomes. The distinction between prescriptive protocols (detailed SOPs followed consistently across vintages, typical of large-scale commercial production) and adaptive protocols (framework guidelines modified in response to vintage conditions, typical of artisanal and premium production) reflects a fundamental philosophical divide in winemaking between consistency and terroir expression. Protocol documentation for VVO serves the specific purpose of linking cultivar and clone characteristics to optimal processing approaches — thick-skinned Cabernet Sauvignon tolerates extended maceration that would over-extract thin-skinned Pinot Noir, high-acid Riesling permits residual sugar retention that would unbalance low-acid Viognier, and aromatic Muscat requires reductive handling that is unnecessary for neutral Trebbiano. The increasing adoption of precision oenology — integrating real-time sensor data (dissolved oxygen, temperature profiles, density curves) with algorithmic decision support — is transforming protocol management from experience-based to data-driven, though the winemaker's interpretive judgment remains central to quality outcomes. Protocol documentation should capture both the standard operating procedure and the vintage-specific deviations that reflect the adaptive responses distinguishing great winemaking from mechanical production.
Herstellungsverfahren
Kellerprotokoll
Vinifikationsprotokoll
Weinbereitungsprotokoll
cellar protocol
oenological protocol
vinification protocol
wine production protocol
winemaking procedure
önologisches Protokoll
Ribéreau-Gayon P et al. (2006) Handbook of Enology Volume 1: The Microbiology of Wine and Vinifications, 2nd Ed, Wiley, pp.: 327-440 Boulton RB et al. (1999) Principles and Practices of Winemaking, Springer, pp.: 1-30 Rankine BC (2004) Making Good Wine: A Manual of Winemaking Practice for Australia and New Zealand, Pan Macmillan, pp.: 1-25
Winemaking Protocol
The wing description of Merlot documents predominantly one prominent wing (OIV 209:2) inserted at the first or second rachis node — the wing extends 30-50 mm laterally from the main rachis axis, bearing 15-30 berries on its own secondary branching system, and creating the characteristic shouldered appearance that contributes to Merlot's conical cluster outline. The wing is typically angled at 30-60 degrees from the rachis, producing a cluster silhouette intermediate between the wingless cylindrical cluster of Pinot Noir (OIV 209:1, no lateral branches sufficiently developed to constitute a wing) and the heavily winged clusters of Trebbiano Toscano (OIV 209:3, with 2-3 wings at successive proximal nodes creating a broad, pyramidal cluster that can appear to be composed of multiple sub-clusters joined at the peduncle region).
A cluster description component that records the presence, number, size, and insertion position of wings — the elongated primary lateral branches of the rachis that project laterally beyond the main cluster body, creating secondary sub-clusters attached to the main rachis at proximal node positions — according to OIV descriptor 209, scored from absent (state 1) through one wing present (state 2) to two or more wings (state 3), with wing development providing one of the most distinctive cultivar-specific cluster architecture characters.
Wing_Description documents a character that is one of the most immediately recognizable features of cluster architecture — the presence and size of wings fundamentally alters the cluster silhouette from the simple geometric forms (cylinder, cone) of wingless clusters to the more complex, branched forms that characterize winged cultivars. Wings develop from the most proximal primary lateral branches of the inflorescence — these basal branches, being the first to develop during inflorescence organogenesis and having the longest growth period, accumulate more flowers and elongate further than the more distal branches, producing the prominent lateral extensions that constitute wings in the mature cluster: the number and size of wings is therefore determined primarily by the inflorescence branching architecture established during bud organogenesis, making wing presence one of the more genetically stable cluster characters — a one-winged cultivar consistently produces one wing regardless of fruit set or berry size variation. Wings have practical significance for cluster management — prominent wings increase the effective cluster surface area (improving spray penetration and air circulation within the cluster), but can also create structural weakness at the wing insertion point where the wing may break from the rachis during transport or handling, and the wing berries may ripen at a different rate than the main cluster body due to their different position relative to the rachis vascular supply.
Beitraube-Beschreibung
Seitentraube-Charakterisierung
Traubenflügel-Beschreibung
cluster wing record
lateral branch description
shoulder branch characterization
Wing Description
Cabernet Sauvignon exhibits strong wing development disposition with 1-2 prominent shoulders per cluster, creating the characteristic winged-conical cluster shape described in ampelographic references. Wing weight in Cabernet Sauvignon averages 20-35% of total cluster weight, with shoulder berry counts of 30-80. Merlot shows variable wing development (0-2 shoulders) depending on clone and vigor. Thompson Seedless typically lacks wings (simple conical shape), while Flame Seedless may develop small shoulders. Wing presence increases effective cluster surface area, potentially improving spray coverage and air circulation compared to cylindrical clusters of equivalent weight.
A cluster architecture disposition that is the inherited capacity of a Vitis vinifera plant to develop prominent lateral branches (shoulders or wings) at the proximal end of the cluster, creating compound cluster shapes with one or more distinct secondary sub-clusters attached to the main rachis below the peduncle.
Wing development results from enhanced growth of the first one or two basal branches of the inflorescence rachis, which elongate and bear berries comparably to the main cluster axis. The trait shows moderate heritability and clone-specific expression. Wings may detach during mechanical harvest, requiring separation equipment adjustments. Ampelographic description systems classify wing development on scales of absent/present/prominent, with further characterization of wing size relative to main cluster body.
Flügelentwicklung
Schulterbildung
Traubenschulter-Disposition
bunch wing trait
cluster shoulder development
lateral branch disposition
OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin Galet P (2000) Dictionnaire encyclopédique des cépages, Hachette pp.: 45-52
Wing Development Disposition
A winter dormant bud at node 4 on a one-year cane of Vitis vinifera cv. Riesling in December in the Mosel Valley, Germany, exhibiting tightly closed brown bud scales covered with dense white trichomes, requiring approximately 900 chilling hours below 7°C for dormancy release, with dissection revealing complete inflorescence primordia containing 150-200 flower initials formed during the previous July.
A dormant bud that enters and maintains arrested development during winter months in response to decreasing photoperiod and temperatures, characterized by endodormancy requiring chilling hour accumulation (typically 800-1500 hours below 7°C depending on cultivar) for release, protected by multiple layers of modified scale leaves with dense trichomes and often waxy or resinous coatings. Winter dormant buds contain fully differentiated primordia formed during the previous growing season, with shoot and inflorescence structures complete but metabolically quiescent, awaiting satisfaction of chilling requirements and subsequent heat accumulation to trigger bud break in spring.
Distinguish from Summer_Dormant_Bud, which enters dormancy due to environmental stress rather than seasonal photoperiod and temperature cues. Winter dormancy depth varies significantly among cultivars, with early-budding varieties typically requiring fewer chilling hours than late-budding ones. The bud scales of winter dormant buds often show cultivar-specific coloration (brown, reddish, gray) and pubescence patterns useful for ampelographic identification.
Winterknospe
hibernating bud
overwintering bud
winter bud
winterruhende Knospe
Based on Lavee & May (1997) "Dormancy of grapevine buds - facts and speculation" Australian Journal of Grape and Wine Research 3=31-46 Fennell (2014) "Genomics and functional genomics of winter low-temperature tolerance in temperate fruit crops" Critical Reviews in Plant Sciences 33=125-140 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press, pp. 167-195.
Winter Dormant Bud
Winter freeze damage quality evaluation in Finger Lakes Riesling following a polar vortex event with minimum temperature of -28°C assessed damage severity across the differential hardiness hierarchy of dormant vine tissues. Bud assessment using the cross-section browning method revealed 65% primary bud mortality across the region, with variation from 40% mortality on well-acclimated vines in favorable sites to 90% mortality on marginally hardened vines or exposed positions, establishing the -28°C event as having exceeded primary bud hardiness (approximately -25°C LT50 for Riesling) across most of the region. Secondary bud survival was substantially higher at 80% regionally, consistent with their 3-4°C greater hardiness providing a recovery pathway through secondary shoot development. Cane phloem damage assessed through cambium browning showed 30% of canes with significant phloem injury that would limit spring sap flow and shoot development from surviving buds. Trunk damage was limited to 5% of vines, concentrated in young plantings and chronically stressed vines with depleted reserves, where freeze cracks and cambium death at the soil line indicated trunk temperatures had exceeded -25°C tolerance limits. The winter freeze damage quality assessment projected 40% yield reduction for the following season from the combined effects of primary bud loss (partially compensated by secondary bud survival), cane damage limiting spur positions, and the carbohydrate depletion from injury response and repair processes.
Winter freeze damage quality is a frost damage severity quality that inheres in a grapevine organism and represents the extent of injury to dormant buds, canes, trunks, and roots from severe cold exposure during the winter dormancy period when temperatures fall below the hardiness limits of vine tissues despite maximum cold acclimation, with damage severity ranging from loss of some primary buds through extensive cane and trunk injury to complete vine kill under the most extreme conditions. This quality addresses the survival-threatening cold stress experienced in continental and cold-climate viticultural regions where winter minimum temperatures may challenge even the cold hardiness of fully acclimated vines, representing a fundamentally different damage dynamic from growing-season frost damage by affecting dormant rather than active tissues. The ontological significance of winter freeze damage quality lies in its potential to cause permanent structural damage or vine death that affects productivity for multiple seasons or requires replanting, distinguishing it from growing-season frost damage that primarily affects current-season crop, and making winter hardiness a primary determinant of cultivar and site suitability in cold-climate viticulture. The biological basis of winter freeze damage encompasses the cellular injury occurring when temperatures fall below the threshold that dormant tissues can survive through supercooling or toleration of extracellular ice formation, with the specific damage threshold varying among tissue types and being influenced by the depth and duration of cold acclimation achieved before the cold event. The characteristic pattern of winter freeze damage reflects the differential hardiness among tissue types, with primary buds typically most sensitive (LT50 around -20 to -25°C for V. vinifera), secondary buds more hardy (2-4°C more tolerant than primaries), cane phloem showing intermediate hardiness, and cane xylem, trunk, and roots showing progressively greater cold tolerance under normal acclimation. The extent of damage from specific winter minimum temperatures depends critically on the acclimation status of vines, which may be compromised by warm autumn conditions preventing full hardening, midwinter warm spells causing deacclimation, or inadequate carbohydrate reserves limiting the metabolic support for cold acclimation processes.
Winter_Freeze_Damage_Quality is_a Frost_Damage_Severity_Quality, representing freeze injury to dormant tissues during the winter hardiness period. This class addresses the most severe cold stress potentially causing permanent vine damage or death. The tissue hardiness hierarchy (primary buds < secondary buds < cane phloem < cane xylem < trunk < roots) determines damage patterns at specific temperatures. LT50 (lethal temperature for 50% mortality) provides quantitative hardiness characterization. Acclimation status critically modulates actual damage thresholds. Distinguished from growing-season frost damage by affecting dormant versus active tissues. Critical for cold-climate cultivar and site selection.
Midwinter-Frostverletzung
Ruhephasen-Kälteschaden
Winterfrostschaden
Wintergefrierschaden
dormant tissue cold injury
freeze damage
midwinter freeze injury
winter kill
Fennell A (2004) Freezing tolerance and injury in grapevines. Journal of Crop Improvement 10:201-235 Mills LJ et al. (2006) Timing of low temperature exposure affects cold hardiness and cold deacclimation kinetics in grape. American Journal of Enology and Viticulture 57:20-26 Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection. Michigan State University Extension Bulletin E2930
Winter Freeze Damage Quality
The wire configuration description for a standard VSP trellis in the Rheingau specifies 5 wires arranged as follows — fruiting wire (2.5 mm high-tensile galvanized steel) at 0.70 m height carrying the Guyot cane or cordon, first pair of movable foliage wires (2.0 mm galvanized) at 1.00 m in clips permitting lateral repositioning for shoot tucking, second pair of movable foliage wires (2.0 mm galvanized) at 1.30 m, and a fixed top wire (2.5 mm galvanized) at 1.55 m for final canopy containment — total wire length per hectare approximately 25,000 m at 2.0 m row spacing, with fruiting wire tension maintained at 500-700 N by in-line ratchet strainers to prevent sagging under crop load.
A training system description that documents the number, height, gauge, material, tension, and functional assignment of individual wires within the trellis framework, specifying the spatial arrangement that defines the fruiting zone, foliage support zone, and canopy containment boundaries within which vine growth is directed and maintained.
Wire_Configuration_Description provides the detailed wire-level specification that translates training system concepts into the physical wire layout that vineyard workers interact with daily. The functional assignment of individual wires — fruiting wire (supporting the permanent cordon or annual cane), foliage wires (containing and positioning vegetative shoots), and containment wires (defining the outer canopy boundary) — determines the canopy microclimate and the efficiency of manual and mechanical management operations: the height of the fruiting wire relative to the ground affects frost exposure, heat stress, and worker ergonomics the spacing between foliage wires determines canopy density control and the use of movable versus fixed foliage wires determines the feasibility and timing of shoot positioning operations. Wire gauge selection balances strength (higher gauge for cordon support under crop load) against handling ease (lower gauge for movable foliage wires that must be manipulated during shoot tucking).
Drahtanordnungs-Spezifikation
Drahtbelegung-Dokumentation
Drahtkonfiguration-Beschreibung
trellis wire configuration
wire height and function description
wire layout specification
Smart RE and Robinson M (1991) Sunlight into Wine: A Handbook for Winegrape Canopy Management, Winetitles pp: 61-90 Dry PR et al. (2004) Viticulture Volume 2: Practices, 2nd Ed, Winetitles pp: 66-96
Wire Configuration Description
A wire-trained horizontal cordon on Vitis vinifera cv. Sauvignon Blanc in Marlborough, New Zealand, extending 86cm along 12-gauge high-tensile trellis wire positioned at 90cm height, cordon resting directly on wire with wire providing continuous support preventing sagging or displacement, spurs emerging from upper surface of cordon above wire, cordon diameter (28mm) now substantially exceeding wire diameter (2.8mm) but wire remaining embedded along bottom of cordon from years of growth around initial training wire, representing mature wire-trained architecture where cordon and wire function as integrated structural unit, with wire support enabling extended cordon length and heavy crop loads without structural failure risk, typical of modern commercial cordon training systems worldwide.
A horizontal cordon deliberately trained and maintained along a specific trellis wire, characterized by intimate association between cordon and wire with the wire providing continuous support along cordon length, cordon positioned directly on or immediately adjacent to wire facilitating structural stability and simplified training, and integration of cordon architecture with trellis infrastructure creating unified permanent framework for annual fruiting wood and canopy development. Wire-trained horizontal cordons represent standard modern cordon configuration where cordons follow and are supported by permanent trellis wires, contrasting with historical stake-trained or unsupported cordon approaches, with wire integration providing structural support reducing breakage risk, simplified establishment through clear training target, and stable long-term architecture maintaining consistent positioning as cordons age and increase in diameter.
Wire-trained horizontal cordons are essentially universal in modern viticulture, with the wire providing both training target during establishment (young cordons trained along wire) and permanent support during mature phase (cordons resting on and partially enveloping wire through secondary growth). The wire integration creates stable architecture resisting displacement from wind, crop weight, or mechanical operations, enables extended cordon lengths (wires can support 1+ meter cordons that would sag or break without support), and simplifies establishment (clear visual and physical target for training shoots during cordon development). The wire specifications (gauge, material, tension) must accommodate long-term cordon loads, with heavier wires (11-12 gauge high-tensile) preferred for substantial cordons bearing heavy crops, and proper tensioning essential to prevent wire stretch and cordon sagging over time as cumulative loads from multiple vines accumulate.
drahtgeführte Kordon
drahtgestützte Kordon
trellis-integrated cordon
wire-following cordon
wire-supported cordon
Based on Smart & Robinson (1991) "Sunlight into Wine= A Handbook for Winegrape Canopy Management" Winetitles, Adelaide White (2015) "Understanding Vineyard Soils" 2nd Edition, Oxford University Press, pp. 234-256 (trellis specifications) Intrieri et al. (2011) "Grapevine mechanization= Constraints and perspectives" Acta Horticulturae 978=471-478.
Wire Trained Horizontal Cordon
Detailed assessment of within vine vigor uniformity quality in bilateral cordon-trained Merlot examined shoot length variation across eight spur positions per cordon at 50 days post-budbreak. Vines affected by Eutypa dieback showed coefficient of variation of 42% with pronounced vigor depression in shoots distal to infected wood, while healthy vines exhibited CV of 16% with gradual proximal-to-distal vigor gradient. Remediation through retraining replacement trunks from basal suckers restored within vine vigor uniformity quality to CV 18% within three growing seasons, with concurrent improvement in fruit composition uniformity reflected in reduced berry-to-berry Brix variation at harvest.
Within vine vigor uniformity quality is a vigor uniformity quality that inheres in an individual grapevine organism and represents the consistency of vegetative growth characteristics across different positions and organs within a single vine including variation between shoots at different node positions, between arms or cordons, between renewal and permanent structure zones, and between canopy faces in divided training systems. This quality reflects the internal resource distribution patterns and developmental coordination that determine the homogeneity of fruit produced by individual vines and the effectiveness of standardized within-vine management practices. The ontological basis of within vine vigor uniformity quality rests on the recognition that individual grapevines exhibit substantial internal variation driven by factors including vascular architecture and sap flow patterns, positional effects along cordons and canes, light exposure gradients within canopies, and localized root function differences affecting specific vine sectors. Shoots originating from basal nodes typically exhibit greater vigor than those from distal positions along cordons due to vascular resistance gradients and proximity to perennial storage tissues, while cordon arms may develop differentially based on orientation, disease history, or winter damage affecting localized hydraulic conductivity. The measurement of within vine vigor uniformity quality requires assessment of variation among individual shoot lengths, node numbers, or lateral development across positions within representative vines, with coefficient of variation values below 15% indicating acceptable uniformity for standard management while higher values may warrant position-specific interventions. High within vine vigor uniformity quality facilitates uniform fruit ripening, simplified harvest decisions, and consistent responses to canopy management operations, while non-uniform vines may produce fruit with substantial variation in composition between positions requiring either selective harvesting or acceptance of blend heterogeneity. Training system selection significantly influences within vine vigor uniformity quality, with cane-pruned systems offering annual renewal opportunity that can improve uniformity compared to cordon systems where positional effects accumulate over years, though cordon systems with appropriate spur positioning and health maintenance can achieve excellent uniformity. Management interventions to improve within vine vigor uniformity quality include balanced pruning adjusting bud numbers to match vine capacity, selective shoot thinning removing weak or excessive shoots at specific positions, and trunk disease management preventing localized vascular compromise.
Within_Vine_Vigor_Uniformity_Quality is_a Vigor_Uniformity_Quality, representing the intra-plant scale of uniformity assessment. This class addresses variation among shoots, positions, and organs within individual vines rather than variation among vines within blocks. Positional effects along cordons, arm balance in bilateral systems, and canopy face differences in divided systems all contribute to this quality. Distinguished from Block_Vigor_Uniformity_Quality which addresses inter-vine and spatial variation patterns.
Einzelstock-Wuchsgleichmäßigkeit
innerpflanzliche Wuchsvariation
intra-vine vigor variation
positional vigor uniformity
positionsabhängige Wuchskonsistenz
single vine vigor homogeneity
within-plant growth consistency
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 85-112 Holzapfel BP et al. (2010) Manipulating the postharvest period and its impact on vine productivity of Semillon grapevines. American Journal of Enology and Viticulture 61:201-206 Trought MCT et al. (2008) Fruit zone leaf removal effects on canopy characteristics, fruit composition and vine carbohydrate status. New Zealand Winegrower 12:82-87
Within Vine Vigor Uniformity Quality
Merlot wood ray in 12-year trunk shows multiseriate configuration 3-5 cells wide, 15-30 cells high (600-1200 μm vertical extent), heterocellular composition with procumbent ray cells 60-100 μm radial length and marginal upright cells, living parenchyma in sapwood storing starch at 8-11% dry weight, rays occupying 10-14% of xylem volume, ray density 7-10 rays per mm tangential, ray-vessel pits facilitating lateral water movement, and rays extending radially through 8-12 annual rings
A wood ray (xylem ray) is a radially oriented band of parenchyma cells extending from vascular cambium through secondary xylem toward trunk center in Vitis vinifera, providing lateral transport of water and solutes between xylem and phloem, storing carbohydrate reserves, and enabling radial communication across annual rings. Wood rays develop from ray initials in vascular cambium= specialized cambial cells divide producing ray parenchyma cells rather than axial xylem elements, ray cells differentiate maintaining radial orientation (contrasting with axial vessel, fiber, and parenchyma orientation), rays extend through successive annual rings as cambium produces new xylem layers, ray height and width vary by ray type (uniseriate rays 1 cell wide, multiseriate rays 2-8 cells wide), and rays persist through sapwood maintaining living parenchyma (eventually dying in heartwood transition). Wood ray anatomy in Vitis vinifera includes= heterocellular composition (rays containing both procumbent cells - radially elongated, 40-120 μm radial length, and upright cells - vertically elongated, positioned at ray margins), multiseriate rays dominant (2-8 cells wide, typically 3-5 cells, compared to uniseriate rays 1 cell wide that are less common), ray height variable (5-40 cells high, 200-1500 μm vertical extent depending on ray class), ray cell walls with numerous pits (bordered or simple pits enabling solute exchange with adjacent vessels and axial parenchyma), and living parenchyma cells throughout sapwood (maintaining metabolic activity, dying in heartwood transition). Wood ray density and distribution includes= 6-12 rays per mm tangential (measured on tangential surface cutting across rays), rays occupying 8-18% of xylem volume (varying by wood age and growth conditions), rays distributed uniformly around circumference (no preferential radial sectors with higher or lower ray density), ray frequency approximately constant across rings (mature wood showing consistent ray density regardless of ring width), and rays crossing multiple annual rings (individual rays extending radially through 5-20+ rings depending on trunk age). Wood ray functions include= lateral transport of water (radial water movement from sapwood vessels to cambium and phloem through ray cells and pits), solute transport (sugars, minerals, hormones moving radially through ray parenchyma from phloem to xylem or vice versa), carbohydrate storage (ray parenchyma storing starch representing 15-30% of total trunk starch reserves with axial parenchyma providing remainder), radial communication (ray parenchyma potentially transmitting signals, hormones, or metabolites across rings), and embolism response (ray parenchyma potentially sensing vessel dysfunction and participating in limited refilling processes). Wood ray carbohydrate storage characteristics include= ray parenchyma accumulating starch particularly in outer sapwood (ray cells showing 6-12% dry weight starch concentration in active sapwood), ray storage contributing to total trunk reserves (rays providing 20-35% of sapwood starch despite being 8-18% of volume, due to high parenchyma density), seasonal starch cycling (accumulation post-harvest, mobilization spring budbreak, replenishment during growing season), and ray starch accessibility (radial orientation facilitating rapid loading from phloem during accumulation and mobilization to cambium and xylem during utilization). Wood ray vessel connection includes= ray-vessel pits (bordered or simple pits in vessel walls adjacent to rays enabling lateral water movement), pit density high on ray-vessel boundaries (40-80 pits per 100 μm² vessel wall adjacent to rays facilitating exchange), rays providing lateral water redistribution (enabling water bypass around embolized or damaged vessels), and rays connecting xylem hydraulic network (lateral connections through rays enhancing overall hydraulic integration). Wood ray development and aging includes= rays initiated at cambium from ray initial cells (maintaining radial files through successive divisions), rays maintaining similar width as they extend (multiseriate rays typically remaining 3-5 cells wide through radial extent), ray parenchyma living in sapwood (metabolically active cells with functioning organelles and membranes), and ray parenchyma dying in heartwood transition (8-15 years post-formation ray cells losing viability along with axial parenchyma). Wood ray anatomical variation includes= uniseriate rays (1 cell wide, 10-30% of total rays, shorter than multiseriate, 5-15 cells high), low multiseriate rays (2-3 cells wide, 20-35% of rays, intermediate height 10-25 cells), high multiseriate rays (4-8 cells wide, 35-50% of rays, tallest rays 20-40+ cells high), aggregate rays (appearing as very wide rays but actually multiple rays closely grouped), and fusiform rays (containing axial elements like vessels or tracheids embedded within ray parenchyma, uncommon in Vitis). Wood ray significance in trunk function includes= facilitating rapid radial transport (enabling efficient loading of photosynthate from phloem into xylem storage parenchyma during reserve accumulation), enabling mobilization distribution (radial movement of mobilized sugars from storage sites to cambium and developing buds), providing hydraulic integration (lateral water connections enhancing trunk hydraulic network resilience), contributing substantially to storage (rays providing 20-35% of trunk carbohydrate reserves), and potentially participating in wound response (ray parenchyma producing defense compounds and perhaps signaling danger to adjacent tissues).
Wood_Ray emphasizes radial parenchyma bands and is distinguished from specific ray types (Primary_Wood_Ray, Secondary_Wood_Ray, Radial_Wood_Ray) by encompassing all ray structures regardless of origin or characteristics. Critical for understanding radial transport in trunk, carbohydrate storage distribution, lateral hydraulic connections, and parenchyma-mediated trunk functions complementing axial vessel transport.
Holzstrahl
Medullary ray
Radial parenchyma
Radiales Parenchym
Xylem ray
Xylemstrahl
Pratt, C. (1974). Vegetative anatomy of cultivated grapes - a review. American Journal of Enology and Viticulture, 25(3), 131-150. | Carlquist, S. (2001). Comparative Wood Anatomy (2nd ed.). Springer.
Wood Ray
Merlot grafted on SO4 rootstock at 19 years showing woody secondary roots as first-order laterals from primary roots, largest woody secondary root 6.8 cm diameter containing extensive secondary xylem with 18 annual growth rings indicating formation in establishment year, woody root bark 2.4 mm thick with protective periderm, woody root extending 168 cm length at average depth 64 cm, woody root xylem conducting water from extensive fine root network, woody tissue showing dense starch staining in dormant winter sample indicating substantial carbohydrate storage, woody secondary root providing structural framework, transport capacity, and storage function
Woody secondary root is secondary root characterized by extensive secondary growth producing substantial woody tissue, emphasizing lignified structural development and wood-like characteristics, showing large diameter typically 2-8 cm reflecting extensive secondary xylem accumulation, demonstrating multiple annual growth rings potentially 5-20+ rings in mature woody secondary roots documenting years of growth, exhibiting protective bark with periderm formation similar to above-ground woody stems, and serving primarily structural framework, water and nutrient transport, and carbohydrate storage rather than direct absorption which occurs mainly in non-woody fine roots, with woody secondary roots typically being first-order laterals or occasionally large second-order branches representing lower orders in root hierarchy, persisting for many years or decades providing permanent structural skeleton, and demonstrating that like above-ground trunk and canes woody secondary roots undergo secondary growth producing wood tissue distinguishing woody perennial grapevine root architecture from herbaceous plants showing only primary root anatomy. Woody secondary root secondary growth characteristics includes= showing vascular cambium activity (meristematic layer dividing to produce secondary xylem inward and secondary phloem outward), producing extensive secondary xylem (annual wood increment accumulating progressively creating substantial woody cylinder comprising majority of root volume), exhibiting annual growth rings (woody secondary root xylem showing distinct rings analogous to trunk allowing age determination), forming secondary phloem (inner bark with sieve elements and companion cells conducting photosynthates though much smaller volume than xylem), and demonstrating progressive thickening (woody secondary root diameter increasing each year through secondary growth potentially 0.2-0.8 cm annual increment). Woody secondary root diameter and structure includes= showing large diameter (woody secondary roots typically 2-8 cm reflecting years of secondary growth accumulation), exhibiting substantial length (woody roots potentially 80-200 cm providing extensive structural framework), displaying woody consistency (tissue firm and dense analogous to branch wood distinguishing from soft fine roots), demonstrating mechanical strength (woody structure providing strength supporting extensive branching network and resisting mechanical forces), and revealing biomass significance (woody secondary roots containing large proportion of total root system biomass despite being numerically less abundant than fine roots). Woody secondary root bark development includes= forming protective periderm (cork cambium producing cork, phellogen, and phelloderm creating protective bark layer), showing bark thickness (woody root bark typically 1-4 mm developing progressively thicker bark with age), exhibiting color variation (woody root bark tan to brown reflecting cork oxidation and aging), demonstrating weathering resistance (bark protecting woody root from mechanical damage, desiccation, and pathogen entry), and revealing similarity to trunk bark (woody secondary root bark structure and function analogous to above-ground woody tissue bark). Woody secondary root xylem characteristics includes= showing vessel elements (woody xylem containing vessels for water transport with diameter typically 60-150 micrometers depending on position and cultivar), exhibiting fiber cells (thick-walled fibers providing mechanical support and strength in woody tissue), demonstrating parenchyma cells (ray and axial parenchyma enabling radial transport and providing storage capacity), displaying annual ring structure (earlywood with large vessels alternating with latewood with smaller vessels and thick fibers), and revealing functional specialization (woody xylem primarily for water transport and mechanical support rather than absorption). Woody secondary root storage function includes= accumulating substantial carbohydrate reserves (woody root large volume with abundant parenchyma and ray cells storing starch particularly during dormancy), showing seasonal storage cycles (reserves highest fall-winter declining spring as mobilized for growth and metabolism), providing major storage capacity (large woody secondary roots potentially storing 300-800+ grams dry weight carbohydrates per root system depending on size), supporting spring bud break (woody root reserves mobilized before leaves functional providing energy for initial shoot and root growth), and enabling stress recovery (woody root carbohydrates supporting vine recovery from freeze, disease, or other stress events). Woody secondary root transport function includes= conducting water from absorptive network (woody root xylem moving water from fine roots through woody secondary root to primary root and trunk), transporting nutrients basipetally (woody root xylem conveying minerals from absorption sites toward plant body), showing high hydraulic conductivity (large-diameter vessels in woody xylem providing efficient low-resistance water transport pathway), enabling phloem distribution (woody root phloem conducting photosynthates from shoots to fine roots and enabling storage in woody tissue), and demonstrating substantial transport capacity (large woody secondary roots capable of transporting 15-40+ liters daily from local root network depending on root size). Woody secondary root structural framework function includes= providing permanent skeleton (woody secondary roots forming stable structural framework supporting ephemeral fine root network), enabling root system architecture (woody root distribution and orientation determining overall root system spatial organization), bearing mechanical loads (woody roots supporting tension, compression, and bending forces from soil, fine roots, and above-ground structure), contributing to anchorage (woody secondary root network helping stabilize vine resisting uprooting and toppling forces), and demonstrating persistence (woody roots functioning for years to decades providing long-term stable framework unlike ephemeral fine roots). Woody secondary root and root system age includes= documenting root system development (annual rings in woody secondary roots recording years of growth providing chronological history), enabling age determination (counting rings in woody root cross-section revealing root age and establishing when initiated), revealing growth rate variation (woody root ring width documenting year-to-year variation in growth conditions and vine vigor), showing growth history (woody root ring patterns potentially correlating with vine management, climate, and stress history), and demonstrating longevity (woody secondary roots potentially persisting and growing throughout entire vine life from establishment through old age).
Woody_Secondary_Root emphasizes extensive secondary growth and lignified structure and is distinguished from fine non-woody secondary roots by focus on wood formation. Includes Lignified_Woody_Secondary_Root and Structural_Woody_Secondary_Root. Critical for understanding secondary growth, annual rings, bark development, carbohydrate storage, transport infrastructure, structural framework, and the wood-producing secondary roots forming permanent structural skeleton analogous to above-ground woody organs.
Holzige Sekundärwurzel
Lignified secondary root
Lignifizierte Seitenwurzel
Structural woody root
Verholzte Strukturwurzel
Wood-forming root
Woody Secondary Root
Syrah hardwood cutting showing wound callus adventitious roots, cutting base forming white callus tissue 5 days after preparation, callus expanding to 8 mm diameter by day 10, microscopic examination revealing root primordia organizing within callus 12-14 days, first roots emerging from callus surface 16 days, cutting producing 7 wound callus adventitious roots by day 24 all emerging from basal callus tissue, roots establishing vascular connection through callus to cutting vascular system enabling water and nutrient uptake supporting establishment
Wound callus adventitious root is callus-derived adventitious root arising from wound-induced callus tissue at injury sites, characterized by formation following mechanical damage such as cutting, pruning, or other trauma that triggers dedifferentiation and callus proliferation, showing developmental sequence of wounding, callus formation, and subsequent root organogenesis in callus, demonstrating particular importance at cutting bases where severing creates wound stimulating callus that becomes site of root formation, and serving wound healing and regeneration functions enabling damaged plants to produce new root systems compensating for root loss or enabling vegetative propagation through wounded cuttings, with wound callus roots showing typical characteristics including formation from wound-induced parenchymatous callus, dependence on wound signaling including ethylene and jasmonic acid in addition to auxin, emergence from callus surface rather than directly from parent tissue, and functional capacity once established though potentially delayed integration compared to direct tissue roots, and demonstrating that wound-induced rooting represents adaptive response allowing plants to recover from damage or enabling humans to exploit wounding for propagation purposes. Wound callus adventitious root wound response includes= showing injury detection (mechanical damage triggering cell signaling cascades), exhibiting dedifferentiation (cells near wound losing specialized identity reverting to pluripotent state), demonstrating cell proliferation (dedifferentiated cells dividing rapidly creating callus tissue mass), revealing wound signaling (ethylene, jasmonic acid, and other wound signals activating responsive genes), and displaying callus characterization (wound callus showing parenchymatous structure with meristematic capacity). Wound callus adventitious root at cutting bases includes= showing universal occurrence (virtually all stem cuttings forming wound callus at cut base as first response), exhibiting basal swelling (wound callus creating visible tissue swelling at cutting base within days), demonstrating root emergence site (callus providing primary site from which roots emerge in many cultivars), revealing callus-root correlation (extent of basal callus often correlating with subsequent root number particularly in difficult-to-root cultivars), and displaying propagation importance (successful cutting propagation often depending on appropriate balance of callus formation and root induction). Wound callus adventitious root hormonal dynamics includes= showing wound-induced auxin (wounding potentially stimulating local auxin accumulation or synthesis), exhibiting ethylene production (wound response generating ethylene which may interact with auxin affecting rooting), demonstrating jasmonic acid (wound signal potentially affecting callus formation and subsequent root organogenesis), revealing auxin application (exogenous auxin treatments compensating for or supplementing endogenous wound-related auxin), and displaying hormone timing (sequential hormone dynamics from wound response through callus formation to root induction). Wound callus adventitious root developmental timing includes= showing rapid callus formation (wound callus visible 3-7 days after cutting preparation or wounding), exhibiting delayed root formation (roots typically emerging from callus 10-20 days after wounding later than callus formation), demonstrating variable kinetics (timing varying with cultivar, hormone treatment, and environmental conditions), revealing cultivar patterns (easy-rooting cultivars showing faster callus-to-root transition than difficult types), and displaying environmental sensitivity (temperature, moisture affecting both callus formation rate and root induction timing). Wound callus adventitious root in difficult-to-root cultivars includes= showing predominant pathway (some difficult cultivars showing mainly callus-mediated rather than direct rooting), exhibiting callus abundance (difficult types often forming extensive callus with delayed or limited root formation), demonstrating protocol importance (managing callus-root balance critical for difficult cultivar propagation success), revealing treatment strategies (hormone concentrations and timing optimized to promote root formation from callus), and displaying research focus (difficult-to-root cultivars subject of research on promoting root induction from wound callus). Wound callus adventitious root vascular integration includes= showing delayed connection (wound callus roots requiring vascular differentiation within callus before connecting to parent), exhibiting callus vasculature (callus potentially developing vascular islands that roots connect to), demonstrating tortuous pathway (vascular connection from root through callus to parent potentially indirect), revealing functional lag (wound callus roots potentially showing delayed functional integration affecting early cutting establishment), and displaying eventual normalization (once integrated, wound callus roots functioning normally supporting plant growth). Wound callus adventitious root management includes= requiring appropriate wounding (cutting preparation creating wound of appropriate size and cleanliness), showing hormone treatment importance (auxin application promoting both callus formation and subsequent root induction), exhibiting moisture requirements (maintaining adequate moisture at wound site essential for callus formation and root development), demonstrating temperature control (optimal temperatures 20-25°C promoting callus-root progression), and displaying monitoring needs (tracking callus formation and root emergence informing propagation progress and success). Wound callus adventitious root practical significance includes= enabling cutting propagation (wound callus rooting fundamental to hardwood and softwood cutting propagation), providing regeneration mechanism (wound callus roots allowing plants to recover from root damage through new root formation), demonstrating healing process (wound callus contributing to overall wound healing beyond just root formation), revealing cultivar challenges (excessive callus without roots problematic in some cultivars requiring protocol optimization), and displaying commercial importance (understanding and managing wound callus rooting essential for commercial nursery propagation).
Wound_Callus_Adventitious_Root emphasizes wound-induced callus origin and is distinguished from other callus roots by focus on injury response. Critical for understanding wound response, cutting base rooting, callus-root progression, delayed integration, difficult cultivars, and the adventitious roots arising from wound-induced callus representing primary rooting mechanism in cutting propagation.
Cutting base root
Injury callus root
Schnittbasis-Wurzel
Verletzungs-induzierte Wurzel
Wound-induced callus root
Wund-Kallus-Adventivwurzel
Wound Callus Adventitious Root
Quantitative characterization of Wound_Compartmentalization_Disposition at Hochschule Geisenheim compared barrier zone formation in 8 cultivars following standardized pruning wound inoculation with Phaeomoniella chlamydospora. At 6 and 12 months post-inoculation, wood sections were harvested and analyzed for compartmentalization zone dimensions, histochemistry, and pathogen extent. 'Riesling' exhibited strong Wound_Compartmentalization_Disposition with barrier zone width of 2.8 mm (versus 0.9 mm in susceptible 'Müller-Thurgau'), barrier zone completeness of 94% (continuous arc around wound), and intense phloroglucinol-HCl staining indicating lignin deposition throughout the barrier. Pathogen colonization was restricted to 15 mm from the wound in 'Riesling' versus 48 mm in 'Müller-Thurgau' at 12 months. Biochemical analysis revealed 4.2-fold higher accumulation of trans-resveratrol and ε-viniferin in 'Riesling' barrier zone tissues. Importantly, barrier zone formation was most effective when wounds were made during vine dormancy (December-February), with spring pruning wounds showing 40% reduction in compartmentalization completeness across all cultivars, demonstrating the interaction between genetic disposition and cultural practices in determining wound compartmentalization efficacy.
A Wound_Compartmentalization_Disposition is an Esca_Resistance_Disposition that inheres in a Vitis vinifera plant and is realized through the formation of anatomical and chemical barriers that restrict the spread of wood-inhabiting pathogens from initial wound infection sites into healthy surrounding tissues. This disposition represents the grapevine's capacity to implement the CODIT (Compartmentalization of Decay in Trees) response, forming boundary zones that limit pathogen movement in radial, tangential, and axial directions within woody tissues. Effective wound compartmentalization involves multiple coordinated processes including rapid formation of wound periderm sealing the wound surface, deposition of suberin and lignin in cell walls of the barrier zone, accumulation of phenolic compounds (particularly stilbenes and flavonoids) with antimicrobial activity, and formation of dense tyloses occluding vessels to prevent longitudinal pathogen spread. The strength of compartmentalization responses varies substantially among grapevine cultivars and is influenced by wound timing, vine age and vigor, and environmental conditions. Cultivars with strong Wound_Compartmentalization_Disposition form narrow, dense barrier zones that effectively contain infections, while susceptible cultivars produce diffuse, incomplete barriers allowing pathogen escape and progressive colonization of functional xylem. This disposition is particularly important following pruning wounds, the primary infection courts for esca-associated fungi, and its enhancement through cultivar selection or cultural practices represents a key strategy for trunk disease management.
Wound_Compartmentalization_Disposition is_a Esca_Resistance_Disposition. This class represents the anatomical barrier formation component of esca resistance, mechanistically related to but distinct from Tylosis_Formation_Disposition which specifically addresses vessel occlusion. CODIT (Compartmentalization of Decay in Trees) principles apply to grapevines though the specific anatomical responses differ from forest trees. Compartmentalization effectiveness is strongly influenced by pruning timing, wound size, and vine physiological status, creating important interactions between genetic disposition and viticultural management.
Barrierezone-Bildung
CODIT response disposition
Wundabschottung
Wundkompartimentierungs-Disposition
decay compartmentalization
wound barrier formation
Shigo AL (1984) Compartmentalization: a conceptual framework for understanding how trees grow and defend themselves, Annual Review of Phytopathology 22:189-214 Pouzoulet J et al. (2014) Xylem vessel diameter affects the compartmentalization of the vascular pathogen Phaeomoniella chlamydospora in grapevine, Frontiers in Plant Science 5:253 Pierron RJG et al. (2016) Cork formation and anatomical responses of grapevine xylem to Phaeomoniella chlamydospora infection, FEMS Microbiology Letters 363:fnw025
Wound Compartmentalization Disposition
Evaluation of Wound_Healing_Efficiency_Disposition at the University of Minnesota compared wound closure dynamics among 10 cultivars and rootstocks with varying crown gall susceptibility under controlled conditions. Standardized wounds (5mm diameter, 3mm depth) were created on green shoots, and healing progression was monitored by measuring wound exudate volume, callus coverage percentage, and periderm formation at 3, 7, 14, and 21 days post-wounding. The cold-hardy hybrid 'Marquette' exhibited superior Wound_Healing_Efficiency_Disposition with 50% callus coverage by day 7 (versus 20% in 'Chardonnay'), complete wound sealing by day 14 (versus day 28 in 'Chardonnay'), and 65% lower cumulative wound exudate volume. When parallel wounds were inoculated with Agrobacterium vitis at 0, 3, 7, or 14 days post-wounding, 'Marquette' showed significantly reduced gall formation on wounds inoculated at 7 days (35% incidence versus 90% in 'Chardonnay') and no galls on wounds inoculated at 14 days (versus 45% in 'Chardonnay'), demonstrating that rapid wound healing effectively shortens the infection window. However, immediate post-wounding inoculation (day 0) resulted in equivalent gall formation (95%) across all genotypes, confirming that Wound_Healing_Efficiency_Disposition operates by reducing infection opportunity rather than through direct antibacterial mechanisms.
A Wound_Healing_Efficiency_Disposition is a Crown_Gall_Resistance_Disposition that inheres in a Vitis plant and is realized through rapid and effective wound closure processes that minimize the temporal window for Agrobacterium colonization of susceptible wound tissues and reduce the availability of wound-released phenolic compounds that induce bacterial virulence genes. This disposition represents a tissue-level resistance mechanism operating before bacterial transformation occurs, based on the principle that Agrobacterium requires access to actively dividing cells at fresh wound surfaces and that wound-released signals trigger bacterial attachment and T-DNA transfer. Effective wound healing involves rapid formation of wound periderm sealing the exposed surface, deposition of suberin and lignin in wound-adjacent cell walls, callus proliferation covering and protecting the wound, and reduced exudation of phenolic compounds (acetosyringone and related molecules) that activate Agrobacterium vir genes. Genotypes with enhanced Wound_Healing_Efficiency_Disposition would be expected to show faster wound closure, reduced wound exudate accumulation, and consequently shorter infection windows and lower transformation success rates. This resistance mechanism is particularly relevant for crown gall management during nursery propagation (graft union protection), mechanical harvesting (trunk wound protection), and winter injury recovery in cold climates. While variation in wound healing among grapevine genotypes has been observed, its specific contribution to crown gall resistance versus other wound-associated pathogens (trunk disease fungi) requires further research.
Wound_Healing_Efficiency_Disposition is_a Crown_Gall_Resistance_Disposition. This class represents tissue-level wound response affecting infection opportunity, distinct from Agrobacterium_Transformation_Resistance_Disposition which operates at the cellular/molecular level. Rapid wound healing provides broad protection against wound-invading pathogens including both Agrobacterium and trunk disease fungi, making this a potentially valuable multi-purpose resistance trait. Cold-hardy cultivars bred for winter survival may have been indirectly selected for enhanced wound healing due to recurrent winter injury pressure. The disposition operates by shortening the infection window rather than eliminating susceptibility.
Schnelle-Wundschließung
Wundheilungs-Effizienz-Disposition
Wundversiegelungs-Kapazität
rapid wound healing
wound closure efficiency
wound sealing capacity
Stover EW et al. (1997) Crown gall in grape: biology and control, Viticulture and Enology Research Center, Fresno State University Pierron RJG et al. (2016) Cork formation and anatomical responses of grapevine xylem to Phaeomoniella chlamydospora infection, FEMS Microbiology Letters 363:fnw025 Burr TJ, Otten L (1999) Crown gall of grape: biology and disease management, Annual Review of Phytopathology 37:53-80
Wound Healing Efficiency Disposition
Long-term field validation of Xiphinema_Index_Resistance_Disposition at the USDA-ARS research vineyard in Parlier, California, compared X. index population dynamics and GFLV transmission over 12 years across rootstocks planted in soil naturally infested with viruliferous nematodes. The susceptible rootstock '5C Teleki' (V. berlandieri × V. riparia) supported X. index populations averaging 1,250 per 250g soil, with 95% of grafted 'Thompson Seedless' scions becoming GFLV-positive within 4 years and exhibiting severe fanleaf symptoms. The resistant '039-16' (V. vinifera × M. rotundifolia BC₃) demonstrated exceptional Xiphinema_Index_Resistance_Disposition, suppressing X. index to 35 per 250g soil (97% reduction), with only 8% of scions becoming infected after 12 years and those showing low viral titers and minimal symptoms. Importantly, '039-16' resistance remained stable throughout the trial with no evidence of resistance-breaking nematode biotypes emerging. Root excavation revealed that '039-16' roots triggered rapid browning and cell death at X. index feeding sites, preventing sustained feeding and reproduction. This field performance validates that Xiphinema_Index_Resistance_Disposition provides durable, effective fanleaf management superior to any other available strategy in heavily infested sites.
A Xiphinema_Index_Resistance_Disposition is a Nematode_Resistance_Disposition that inheres in a Vitis plant root system and is realized through defense processes that limit feeding, reproduction, and population development of the dagger nematode Xiphinema index, an ectoparasitic nematode of paramount importance in viticulture as the primary vector of Grapevine fanleaf virus (GFLV). This disposition represents resistance to a large (3-4mm) nematode that feeds externally on root tips using a long odontostyle (feeding stylet) to penetrate cells and extract contents, causing direct root damage through feeding site destruction and, critically, transmitting GFLV between infected and healthy vines. X. index acquires GFLV particles during feeding on infected roots and retains virus in association with its foregut cuticle, remaining viruliferous for months to years and transmitting to healthy vines during subsequent feeding. Resistance mechanisms include root tip anatomical features impeding stylet penetration, rapid cell death at feeding sites preventing sustained nutrition, chemical factors in root tissues deterring feeding initiation or causing nematode mortality, and overall poor host suitability reducing nematode reproduction and population buildup. Xiphinema_Index_Resistance_Disposition serves dual functions:protecting root system integrity from feeding damage and, more importantly, interrupting fanleaf virus transmission by eliminating or dramatically reducing vector populations. Muscadinia rotundifolia and certain North American Vitis species (V. arizonica, V. candicans, V. rufotomentosa) provide strong X. index resistance that has been introgressed into rootstock breeding lines, offering the most effective long-term strategy for managing fanleaf disease in infested vineyards.
Xiphinema_Index_Resistance_Disposition is_a Nematode_Resistance_Disposition. This class specifically addresses resistance to X. index, the primary GFLV vector, distinct from general nematode resistance. The dual importance of X. index resistance for both direct root protection and fanleaf virus management makes it the highest priority nematode resistance trait for rootstock breeding in GFLV-endemic regions. The resistance mechanism appears similar to Root_Phylloxera_Resistance_Disposition, involving hypersensitive-like cell death at feeding sites. Resistance sources include M. rotundifolia and several North American Vitis species, with the XiR1 locus from M. rotundifolia being the best-characterized resistance gene.
Dolchnematoden-Resistenz
GFLV-Vektor-Resistenz
X. index resistance
Xiphinema-index-Resistenz-Disposition
dagger nematode resistance
fanleaf vector resistance
Walker MA et al. (1994) Resistance to Xiphinema index in Vitis and its hybrids, Vitis 33:21-26 Hwang CF et al. (2010) Fine mapping and characterization of XiR1, a locus for Xiphinema index resistance in grape, Theoretical and Applied Genetics 121:1749-1762 Esmenjaud D, Bouquet A (2009) Selection and application of resistant germplasm for grapevine nematodes management, Integrated Management of Fruit Crops and Forest Nematodes, Springer
Xiphinema Index Resistance Disposition
Muscadinia rotundifolia and its hybrids show near immunity to Xiphinema index, with nematode populations declining to undetectable levels on VR 043-43 (V. vinifera × M. rotundifolia) rootstocks in INRAE trials at Montpellier. O39-16 (M. rotundifolia × V. vinifera) demonstrates strong X. index resistance supporting Cabernet Sauvignon in GFLV-infested experimental vineyards in the Napa Valley, reducing fanleaf transmission by over 90% compared to susceptible 5C Teleki. Dog Ridge (V. champinii) shows moderate X. index resistance with nematode reproduction reduced by 60-70% relative to SO4 or Kober 5BB.
A nematode-resistance rootstock role that inheres in a grafted rootstock by virtue of its specific capacity to resist feeding by Xiphinema index Thorne and Allen, the dagger nematode that vectors Grapevine fanleaf virus (GFLV). Xiphinema index is a migratory ectoparasite that feeds on root tips using a long odontostyle (stylet length 120-130 µm), acquiring and transmitting GFLV during feeding bouts lasting 15-60 minutes. Resistant rootstocks deter X. index feeding through root tip anatomical features (thick-walled epidermal cells, dense trichome-like root hairs), production of repellent or toxic exudates, and rapid wound healing that limits stylet penetration. Resistance is assessed by nematode population density per gram of root (resistant genotypes supporting fewer than 10 nematodes/g versus 50-200/g on susceptible roots) and by GFLV transmission efficiency, which is reduced on resistant rootstocks due to shortened feeding duration.
Xiphinema_Index_Resistant_Rootstock_Role addresses both direct nematode damage (root tip destruction, reduced nutrient uptake) and the critical epidemiological role of X. index as the sole natural vector of GFLV, the most economically damaging viral disease of grapevine worldwide causing yield losses of 50-80% in severely affected vineyards. Muscadinia rotundifolia carries the strongest known resistance but presents graft compatibility challenges with V. vinifera scions. Ongoing breeding programs at INRAE, UC Davis, and CSIRO are developing M. rotundifolia-derived rootstocks with improved graft compatibility while retaining X. index resistance.
GFLV vector resistant rootstock
Rebvirosen-Vektor-resistente Unterlage
Xiphinema index resistant rootstock
Xiphinema-index-resistente Unterlage
fanleaf nematode resistant rootstock
gegen Xiphinema index resistente Unterlage
Bouquet A et al. (2000) Grapevine rootstocks with resistance to Xiphinema index, Acta Horticulturae 528: 521-528 Walker MA et al. (1994) Resistance to Xiphinema index in Vitis and Muscadinia species, Vitis 33: 75-78 Andret-Link P et al. (2004) Grapevine fanleaf virus: still a major threat to the grapevine industry, Journal of Plant Pathology 86: 183-195
Xiphinema Index Resistant Rootstock Role
Greenhouse evaluation of Xiphinema_Transmission_Resistance_Disposition at UC Davis quantified nematode feeding preference and population development on 20 rootstock genotypes in controlled pot assays. Vines were planted in soil inoculated with 500 X. index individuals per pot, and nematode populations were assessed after 6 months. Susceptible 'Freedom' (V. champinii derivative) supported vigorous nematode reproduction with final populations averaging 2,850 per pot (470% increase). The resistant selection 'O39-16' (V. vinifera × M. rotundifolia BC₃) showed strong Xiphinema_Transmission_Resistance_Disposition with final populations of only 85 per pot (83% decline from initial inoculum), indicating that nematodes failed to reproduce and gradually died. Microscopic examination of roots revealed that X. index probing attempts on 'O39-16' triggered rapid browning of contacted cells, with 78% of observed feeding attempts abandoned within 10 minutes compared to sustained feeding lasting >60 minutes on 'Freedom'. In field trials with GFLV-viruliferous X. index, 'O39-16' rootstocks remained 95% virus-free after 8 years while adjacent 'Freedom' rootstocks reached 100% infection within 4 years, demonstrating effective translation of nematode resistance into field-level fanleaf disease control.
A Xiphinema_Transmission_Resistance_Disposition is a Fanleaf_Resistance_Disposition that inheres in a Vitis plant and is realized through host characteristics that prevent or limit Grapevine fanleaf virus transmission by Xiphinema dagger nematodes, primarily through resistance to nematode feeding on roots. This disposition represents indirect viral resistance operating by interrupting the vector-mediated transmission cycle rather than through direct antiviral mechanisms. Xiphinema index, the primary GFLV vector, is an ectoparasitic nematode that feeds on root tips using a long odontostyle (feeding stylet) to penetrate epidermal cells and extract cell contents. The nematode acquires GFLV particles during feeding on infected roots and retains virus in association with its foregut lining, transmitting to healthy plants during subsequent feeding over periods of months to years. Resistance to X. index feeding in certain Vitis genotypes involves multiple mechanisms including root tip anatomical features impeding stylet penetration, biochemical factors deterring feeding initiation or continuation, hypersensitive response killing cells at feeding sites, and root regeneration capacity compensating for feeding damage. Muscadinia rotundifolia and certain North American Vitis species (V. arizonica, V. rufotomentosa, V. candicans) exhibit strong X. index resistance that dramatically reduces nematode populations and consequently limits virus spread. This resistance has been successfully introgressed into rootstock breeding lines, providing the most effective practical strategy for managing fanleaf disease in infested vineyards through disruption of the virus-vector relationship.
Xiphinema_Transmission_Resistance_Disposition is_a Fanleaf_Resistance_Disposition. This class represents resistance to the nematode vector rather than to GFLV itself, providing indirect but highly effective viral disease control. X. index resistance is the primary mechanism through which resistant rootstocks protect grafted V. vinifera scions from fanleaf infection. The resistance is distinct from general nematode resistance, as X. index has specific host preferences and resistance mechanisms may not transfer to other nematode species. Resistance sources include M. rotundifolia and several North American Vitis species.
Nematoden-Vektor-Resistenz
Xiphinema index resistance
Xiphinema-index-Resistenz
Xiphinema-Übertragungs-Resistenz-Disposition
nematode transmission resistance
vector-mediated GFLV resistance
Xiphinema Transmission Resistance Disposition
Quantitative characterization of Xylella_Multiplication_Resistance_Disposition at the USDA-ARS San Joaquin Valley Agricultural Sciences Center employed time-course bacterial population measurements following standardized needle inoculation with X. fastidiosa strain Temecula1 (10⁴ CFU delivered per inoculation point). Petiole samples were collected at 4, 8, 12, and 16 weeks post-inoculation from susceptible 'Chardonnay', resistant V. arizonica b43-17, and 20 interspecific breeding selections. 'Chardonnay' exhibited rapid bacterial multiplication with populations increasing from initial inoculum to 10⁵ CFU/g by week 4, 10⁷ CFU/g by week 8, and plateau at 10⁸ CFU/g by week 12, accompanied by progressive symptom development. V. arizonica b43-17 demonstrated exceptional Xylella_Multiplication_Resistance_Disposition with populations remaining below 10³ CFU/g through week 16, representing 100,000-fold suppression relative to 'Chardonnay'. Breeding selections showed continuous variation in multiplication resistance, with PdR1-positive selections maintaining populations of 10³-10⁵ CFU/g while PdR1-negative selections reached 10⁶-10⁸ CFU/g. Critically, a symptom severity threshold was identified at approximately 10⁶ CFU/g, below which vines remained asymptomatic regardless of genotype, demonstrating that Xylella_Multiplication_Resistance_Disposition sufficient to maintain subthreshold populations provides functional disease resistance.
A Xylella_Multiplication_Resistance_Disposition is a Pierces_Disease_Resistance_Disposition that inheres in a Vitis plant and is realized through host factors that specifically limit Xylella fastidiosa population growth within xylem vessels, resulting in bacterial titers orders of magnitude below those achieved in susceptible genotypes under equivalent inoculation conditions. This disposition represents the quantitative aspect of Pierce's disease resistance most directly correlated with disease outcomes, as symptom severity and vine mortality are strongly associated with bacterial population levels in planta. Mechanisms contributing to Xylella multiplication resistance may include unfavorable xylem sap composition (pH, ion concentrations, organic nutrient availability) limiting bacterial growth, antimicrobial compounds present in xylem fluid or vessel walls, rapid host recognition triggering localized defense responses, and restriction of bacterial spread between vessels limiting systemic colonization. The distinction between multiplication resistance and systemic spread resistance is experimentally important, as some host genotypes may allow local bacterial growth but prevent systemic colonization, while others may permit slow spread but maintain low populations throughout. Xylella_Multiplication_Resistance_Disposition is the primary measurable phenotype in Pierce's disease resistance screening programs, assessed through quantitative culture plating or qPCR of petiole or stem tissues at standardized times post-inoculation. Strong multiplication resistance is characteristic of PdR1-mediated resistance, but quantitative variation in multiplication rates exists even among genotypes lacking major resistance genes, providing potential for accumulating minor factors through breeding.
Xylella_Multiplication_Resistance_Disposition is_a Pierces_Disease_Resistance_Disposition. This class represents the quantitative phenotype of limiting bacterial populations, which can be conferred by major genes (PdR1) or potentially by accumulation of minor factors. Multiplication resistance is distinct from tolerance (Xylem_Occlusion_Tolerance_Disposition), which enables productivity maintenance despite higher bacterial populations. The 10⁶ CFU/g symptom threshold provides a practical resistance target for breeding programs. Multiplication resistance is measured by culture plating (CFU/g) or qPCR (genome copies/g), with strong correlation between methods.
Bakterien-Titer-Beschränkung
Xylella population limitation
Xylella-Populationskontrolle
Xylella-Vermehrungs-Resistenz-Disposition
bacterial titer restriction
multiplication rate resistance
Krivanek AF, Walker MA (2005) Vitis resistance to Pierce's disease is characterized by differential Xylella fastidiosa populations in stems and leaves, Phytopathology 95:44-52 Roper MC et al. (2007) Xylella fastidiosa requires polygalacturonase for colonization and pathogenicity in Vitis vinifera grapevines, Molecular Plant-Microbe Interactions 20:411-419 Lindow S (2019) Secrets of success: biological control of Pierce's disease by Paraburkholderia phytofirmans and Xylella fastidiosa, Phytopathology 109:5
Xylella Multiplication Resistance Disposition
Xylem-specific hydraulic conductivity (Ks) in grapevine stems ranges from 3-12 kg/m/s/MPa across genotypes, with vessel diameter being the primary determinant (conductivity scales with the fourth power of radius per Hagen-Poiseuille law). Syrah and Grenache show wide vessels (mean diameter 80-120 µm) with high Ks (8-12 kg/m/s/MPa), while Pinot Noir and Riesling have narrower vessels (60-90 µm) with lower Ks (4-7 kg/m/s/MPa). Leaf-specific conductivity (KL, normalized to supplied leaf area) of 2-8 × 10⁻⁴ kg/m/s/MPa determines the water potential gradient required to sustain transpiration. Seasonal conductivity loss from winter embolism ranges from 20-60% in cold climates, with spring refilling restoring 70-90% of original conductivity through positive root pressure.
A water transport disposition that is the inherited capacity of a Vitis vinifera plant to conduct water through xylem vessels with genotype-specific hydraulic efficiency, determined by vessel diameter, vessel density, vessel length, intervessel pit membrane properties, and the proportion of functional (non-embolized) conduits.
Xylem conductivity reflects the anatomical investment in water transport infrastructure, with higher conductivity enabling greater stomatal opening and photosynthesis but potentially increasing cavitation vulnerability. The Hagen-Poiseuille relationship means that small increases in mean vessel diameter yield disproportionate conductivity gains, creating selection pressure for large vessels constrained by cavitation risk. Petiole and shoot conductivity measurements provide accessible proxies for whole-vine hydraulic status. Ring-porous versus diffuse-porous anatomy affects seasonal conductivity patterns.
Xylem-Leitfähigkeit
stem hydraulic efficiency
vascular conductivity
vaskuläre Leitungseffizienz
xylem hydraulic conductance
Lovisolo C, Schubert A (1998) Effects of water stress on vessel size and xylem hydraulic conductivity in Vitis vinifera L. Journal of Experimental Botany 49:693-700 Brodersen CR et al. (2010) The dynamics of embolism repair in xylem: in vivo visualizations using high-resolution computed tomography. Plant Physiology 154:1088-1095
Xylem Conductivity Disposition
Xylem discoloration quality assessment in Okanagan Valley Merlot following a severe winter freeze (-28°C for 72 hours) documented wood damage patterns and their relationship to subsequent vine decline. Cross-sectional examination of cane and trunk samples revealed characteristic xylem discoloration quality patterns distinguishing freeze injury from other causes of wood browning. Freeze-damaged xylem showed xylem discoloration quality as irregular brown-to-black sectors distributed across the wood cross-section, concentrated in the outer growth rings representing the most recent 2-3 years of wood formation that showed greater freeze sensitivity than older, more heavily lignified inner wood. The percentage of xylem cross-sectional area showing discoloration was scored, with values ranging from 15% in mildly affected vines to 75% in severely damaged specimens. Follow-up assessment 18 months after the freeze event revealed that xylem discoloration quality correlated strongly with subsequent Botryosphaeria canker development, with 45% of vines showing greater than 50% xylem discoloration developing visible canker symptoms versus only 8% of vines with less than 25% initial discoloration. The xylem discoloration quality assessment established that freeze-damaged wood served as infection court for trunk disease pathogens, with the initial mechanical damage creating susceptibility to secondary biotic injury that produced progressive vine decline extending years beyond the original freeze event, emphasizing the importance of xylem damage evaluation for predicting long-term vineyard health consequences of winter injury.
Xylem discoloration quality is a cold damage symptom quality that inheres in grapevine woody tissues and represents the extent and pattern of abnormal coloration in the xylem (wood) visible in cross-sectioned canes and trunks following freezing injury, with discoloration patterns indicating ice damage to xylem vessels and surrounding parenchyma cells that may compromise water transport capacity and create entry points for vascular pathogens that colonize damaged tissue. This quality addresses damage to the water-conducting system that supports transpiration and photosynthesis, recognizing that xylem damage may not cause immediate vine mortality due to the redundancy of multiple xylem vessels but can progressively impair function as damaged vessels become non-functional through embolism or pathogen colonization. The ontological significance of xylem discoloration quality lies in its indication of structural wood damage that may have delayed consequences emerging over subsequent growing seasons, with freeze-damaged xylem potentially becoming colonized by vascular pathogens including Phomopsis and Botryosphaeria that exploit freeze wounds as entry points, leading to progressive canker development and structural decline that traces back to initial freeze injury. The biological basis of xylem discoloration encompasses several distinct damage mechanisms including direct ice damage causing cell death in ray parenchyma and vessel-associated cells, the formation of ice in vessel lumens causing mechanical damage to vessel walls, and the subsequent oxidation and pathogen colonization of damaged tissues producing visible discoloration. The characteristic appearance of freeze-damaged xylem varies with damage severity, ranging from subtle browning in scattered sectors indicating localized damage to extensive dark brown or black discoloration indicating severe widespread injury, with the pattern providing information about damage extent and the likely functional consequences. The radial pattern of xylem discoloration in cross-section may show annual ring effects where the most recent growth rings are more severely affected, reflecting the differential hardiness of xylem tissues of different ages, with younger, less lignified wood typically showing greater vulnerability than older, fully mature wood.
Xylem_Discoloration_Quality is_a Cold_Damage_Symptom_Quality, representing freeze injury to the xylem water-conducting tissue and associated parenchyma. This class addresses wood damage with delayed consequences through pathogen colonization. Irregular sectoral browning pattern distinguishes freeze damage from vascular disease. Outer growth rings (younger wood) show greater freeze sensitivity. Strong correlation with subsequent canker development demonstrates the pathogen-entry function of freeze wounds. Long-term vine decline may trace back to initial freeze injury through secondary infection. Distinguished from Phloem_Necrosis_Quality by focus on xylem rather than phloem.
Holzverfärbung
Xylem-Frostschaden
Xylemverfärbung
freeze damage to wood
vascular wood injury
vaskulärer Holzschaden
wood discoloration
xylem browning
Zabadal TJ et al. (2007) Winter injury to grapevines and methods of protection. Michigan State University Extension Bulletin E2930 Úrbez-Torres JR et al. (2014) Identification and pathogenicity of fungi associated with grapevine trunk diseases in British Columbia. Canadian Journal of Plant Pathology 36:358-376 Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 508-545
Xylem Discoloration Quality
Investigation of Xylem_Hardiness_Disposition at Washington State University employed differential thermal analysis (DTA) to detect low-temperature exotherms corresponding to intracellular freezing in xylem ray parenchyma across 18 cultivars at maximum midwinter hardiness. DTA of cane sections revealed distinct exotherm profiles, with high-temperature exotherms (HTEs) representing extracellular ice formation in apoplast and vessel lumens (non-lethal), and low-temperature exotherms (LTEs) representing lethal intracellular freezing in supercooled parenchyma. Xylem_Hardiness_Disposition, defined by LTE temperature, ranged from -27°C ('Merlot') to -42°C ('Valiant'), with xylem consistently the hardiest cane tissue. The xylem-phloem hardiness differential ranged from 2°C ('Frontenac') to 5°C ('Cabernet Franc'), indicating cultivar-specific vulnerability hierarchies. Anatomical analysis revealed that ray parenchyma cell size correlated with supercooling capacity, with smaller cells (mean diameter 12 μm in 'Frontenac') supercooling to lower temperatures than larger cells (22 μm in 'Merlot'), consistent with homogeneous ice nucleation theory predicting greater supercooling in smaller volumes. Following extreme cold exposure (-38°C) in field conditions, cultivars with Xylem_Hardiness_Disposition below this threshold showed spring symptoms including delayed budbreak, reduced xylem sap flow (measured by bleeding after pruning cuts), and impaired starch mobilization from ray parenchyma. These findings establish xylem ray parenchyma supercooling limits as the ultimate cold tolerance boundary in grapevine canes.
A Xylem_Hardiness_Disposition is a Cane_Cold_Hardiness_Disposition that inheres specifically in the xylem tissue of dormant grapevine canes and is realized through the freezing tolerance of xylem parenchyma cells (ray and axial parenchyma) and the structural integrity of vessel elements and fibers, determining the capacity for water transport and structural support of spring growth. This disposition represents the cold hardiness of the water-conducting and mechanical support tissue, with xylem function essential for supplying water to developing buds and shoots and maintaining cane structural integrity. Xylem exhibits a complex freezing response, with the non-living vessel elements and fibers tolerating ice formation in their lumens (indeed, winter embolism is common and resolved by spring root pressure), while living xylem parenchyma cells—particularly ray parenchyma containing stored starch—survive through deep supercooling, maintaining liquid water within cells to temperatures far below the equilibrium freezing point. Deep supercooling capacity in grapevine xylem parenchyma is limited by ice nucleation events, with lethal intracellular freezing occurring when supercooling limits are exceeded, detectable as low-temperature exotherms (LTEs) in differential thermal analysis. The supercooling limit of xylem ray parenchyma represents the absolute cold tolerance boundary for xylem function, typically the hardiest tissue in canes and exceeded only in the most extreme cold events. Xylem damage manifests as browning of ray parenchyma visible in cross-sections and can disrupt spring water transport and carbohydrate storage/mobilization functions.
Xylem_Hardiness_Disposition is_a Cane_Cold_Hardiness_Disposition. Xylem parenchyma survives freezing through deep supercooling, distinct from the freeze-tolerance mechanism of phloem. LTEs detected by DTA represent lethal intracellular freezing when supercooling limits are exceeded. Xylem is typically the hardiest cane tissue, so xylem damage indicates extreme cold events. Ray parenchyma stores starch critical for spring growth initiation, so xylem damage affects both water transport and carbohydrate mobilization.
Holzhärte
Xylem-Härte-Disposition
Xylemparenchym-Kältetoleranz
ray parenchyma cold resistance
wood parenchyma hardiness
xylem freezing tolerance
Pierquet P, Stushnoff C (1980) Relationship of low temperature exotherms to cold injury in Vitis riparia Michx., American Journal of Enology and Viticulture 31:1-6 George MF et al. (1974) Low temperature exotherms and woody plant distribution, HortScience 9:519-522 Fennell A (2004) Freezing tolerance and injury in grapevines, Journal of Crop Improvement 10:201-235
Xylem Hardiness Disposition
Direct assessment of Xylem_Loading_Restriction_Disposition at the Australian Centre for Plant Functional Genomics employed xylem sap collection and analysis combined with transporter gene expression profiling across 8 rootstocks under salinity (50 mM NaCl for 21 days). Xylem sap was collected by root pressurization, and chloride concentration was measured alongside root cortex chloride to calculate the loading ratio (xylem/cortex concentration). Xylem_Loading_Restriction_Disposition showed striking genotypic variation. '140 Ruggeri' demonstrated exceptional loading restriction with xylem sap chloride of only 3.2 mM despite root cortex concentration of 65 mM, yielding a loading ratio of 0.05 (only 5% of cortical chloride reaching xylem). 'Schwarzmann' showed minimal restriction with xylem sap chloride of 42 mM and cortex of 58 mM, yielding loading ratio of 0.72. Gene expression analysis revealed that '140 Ruggeri' roots showed 6-fold lower VvSLAH3 expression and 4-fold lower VvSLAH1 expression under salinity compared to 'Schwarzmann', directly implicating these anion channels in Xylem_Loading_Restriction_Disposition. Electrophysiological studies on isolated xylem parenchyma protoplasts confirmed lower chloride channel activity in '140 Ruggeri' cells (current density -15 pA/pF versus -52 pA/pF in 'Schwarzmann' at -120 mV). These findings establish that Xylem_Loading_Restriction_Disposition is the dominant mechanism underlying rootstock chloride exclusion differences, mediated by differential SLAH channel expression.
A Xylem_Loading_Restriction_Disposition is a Chloride_Exclusion_Disposition that inheres in Vitis root stele tissues and is realized through mechanisms that limit the transfer of chloride ions from root symplast into the xylem vessels, preventing chloride translocation to shoots even when root cortical chloride concentrations are elevated. This disposition represents the critical second barrier in chloride exclusion, operating at the root stele where chloride accumulated in cortical cells must cross xylem parenchyma membranes to enter the transpiration stream. Xylem loading is an active process requiring energy expenditure to move chloride against the concentration gradient from relatively low xylem sap concentrations into the high-volume transpiration flow. The molecular machinery for xylem chloride loading includes SLAH (Slow Anion Channel-Associated Homolog) family anion channels and potentially NPF (Nitrate transporter/Peptide transporter Family) members that facilitate chloride release into xylem vessels. Effective Xylem_Loading_Restriction_Disposition involves reduced expression, activity, or plasma membrane localization of these xylem-loading transporters in stelar parenchyma cells under salinity conditions. Xylem_Loading_Restriction_Disposition appears to be the primary site of genotypic variation in grapevine chloride exclusion, with excluding rootstocks showing dramatically lower xylem sap chloride concentrations than non-excluders at equivalent root cortical concentrations, indicating that root uptake differences alone cannot explain exclusion phenotypes.
Xylem_Loading_Restriction_Disposition is_a Chloride_Exclusion_Disposition. Xylem loading restriction is the primary site of genotypic variation in grapevine chloride exclusion. Loading ratio (xylem sap/root cortex concentration) quantifies restriction effectiveness. SLAH family anion channels are key molecular determinants, with excluder rootstocks showing reduced SLAH expression under salinity. Xylem loading is an active process requiring energy but protecting shoot tissues is energetically favorable compared to toxicity costs. Loading restriction can maintain low xylem chloride even when root cortical chloride is elevated.
Steläre-Chlorid-Retention
Wurzel-Spross-Chlorid-Kontrolle
Xylembeladungs-Restriktions-Disposition
root-to-shoot chloride restriction
stelar chloride retention
xylem chloride loading control
Henderson SW et al. (2014) Shoot chloride exclusion and salt tolerance in grapevine is associated with differential ion transporter expression in roots, BMC Plant Biology 14:273 Gilliham M, Tester M (2005) The regulation of anion loading to the maize root xylem, Plant Physiology 137:819-828 Teakle NL, Tyerman SD (2010) Mechanisms of Cl⁻ transport contributing to salt tolerance, Plant Cell and Environment 33:566-589
Xylem Loading Restriction Disposition
Investigation of Xylem_Occlusion_Tolerance_Disposition at Texas A&M University compared hydraulic responses to X. fastidiosa infection across 6 cultivars with varying disease phenotypes. Vines were mechanically inoculated and assessed at 16 weeks post-inoculation for bacterial population (qPCR), percent loss of hydraulic conductivity (PLC) measured by flow meter, and whole-vine transpiration in controlled environment chambers. Susceptible 'Chardonnay' (bacterial titer 10⁸ CFU/g) exhibited 78% PLC and transpiration reduced to 25% of healthy controls, with severe leaf scorch on all leaves. The interspecific hybrid 'Blanc du Bois' showed equivalent bacterial titer (10⁸ CFU/g) but demonstrated enhanced Xylem_Occlusion_Tolerance_Disposition with only 52% PLC and transpiration maintained at 55% of controls, with scorch limited to older leaves. Anatomical examination revealed that 'Blanc du Bois' possessed 40% higher native vessel density (185 vs 132 vessels/mm²) and smaller mean vessel diameter (38 vs 52 μm), providing greater hydraulic redundancy. When both cultivars were subjected to equivalent artificial flow restriction (mimicking 60% PLC), 'Blanc du Bois' maintained 70% of control transpiration versus 40% in 'Chardonnay', confirming intrinsic tolerance differences independent of bacterial effects. These findings suggest that Xylem_Occlusion_Tolerance_Disposition provides an additional resistance strategy complementing direct antibacterial mechanisms.
A Xylem_Occlusion_Tolerance_Disposition is a Pierces_Disease_Resistance_Disposition that inheres in a Vitis plant and is realized through the capacity to maintain adequate hydraulic conductivity and water transport despite partial occlusion of xylem vessels by Xylella fastidiosa biofilms, bacterial aggregates, and host-produced tyloses, thereby minimizing water stress symptoms and productivity losses even when bacterial infection is established. This disposition represents a tolerance mechanism distinct from resistance mechanisms that limit bacterial multiplication or spread, enabling functional compensation for partial vascular blockage. Pierce's disease symptoms including leaf scorch, premature defoliation, and vine decline result substantially from hydraulic failure as progressive vessel occlusion reduces water transport capacity below levels required to meet transpirational demand. Xylem_Occlusion_Tolerance_Disposition may involve anatomical features including high native vessel density providing hydraulic redundancy, smaller vessel diameters reducing individual vessel contribution to total conductivity, efficient inter-vessel connectivity enabling bypass of occluded elements, and enhanced root hydraulic conductance compensating for reduced stem conductivity. Additionally, tolerance may involve physiological adjustments including osmotic adjustment maintaining cell turgor at reduced water potentials, stomatal regulation balancing transpiration with reduced supply, and modified growth allocation prioritizing hydraulic function. The distinction between multiplication resistance and occlusion tolerance is experimentally important, as some cultivars maintain productivity despite substantial infection (tolerance), while others decline rapidly once infection thresholds are exceeded (low tolerance). Breeding for occlusion tolerance could complement multiplication resistance strategies, particularly for conditions where resistance is incomplete.
Xylem_Occlusion_Tolerance_Disposition is_a Pierces_Disease_Resistance_Disposition. This class represents tolerance to vascular dysfunction rather than resistance to bacterial infection, providing a complementary strategy to Xylella_Multiplication_Resistance_Disposition. Occlusion tolerance may enable functional viticulture even where infection cannot be completely prevented. The disposition overlaps conceptually with general drought tolerance, as both involve maintaining hydraulic function under water stress. Tylosis formation, while contributing to bacterial containment, paradoxically exacerbates occlusion thus, optimal defense may involve balanced tylosis response sufficient to limit spread without catastrophic conductivity loss.
Gefäßblockade-Toleranz
Hydraulik-Dysfunktion-Toleranz
Xylem-Okklusions-Toleranz-Disposition
hydraulic dysfunction tolerance
vascular blockage tolerance
vessel occlusion tolerance
McElrone AJ et al. (2003) Variation in xylem structure and function in stems and roots of trees to 20 m of depth, New Phytologist 163:507-517 Sun Q et al. (2013) Xylem occlusion in grapevine to limit the systemic spread of Pierce's disease, American Journal of Enology and Viticulture 64:153-159 Choat B et al. (2010) Vascular function in grape berries across development and its relevance to apparent hydraulic isolation, Plant Physiology 154:1546-1555
Xylem Occlusion Tolerance Disposition
Cabernet Sauvignon xylem vessel in earlywood of ring 8 shows 220-280 μm diameter, vessel element length 300-450 μm with simple perforation plates enabling continuous water column, lignified secondary walls with bordered pits (40-60 pits per 100 μm² wall area) enabling lateral connections to adjacent vessels and ray parenchyma, dead protoplasm creating hollow lumen for water transport, vessel contributing to ring-porous pattern with earlywood vessels providing 65-75% of total ring hydraulic conductance despite being only 35-45% of ring area
An xylem vessel is a tubular water-conducting structure in Vitis vinifera secondary xylem formed by vertical alignment of vessel elements (tracheary elements) with perforated end walls creating continuous tubes for long-distance water and mineral transport from roots to canopy, representing the principal hydraulic conduit in grapevine trunk wood. Xylem vessel structure includes= vertical series of vessel elements (individual tracheary elements 200-600 μm length aligned vertically), perforated end walls (perforation plates with large openings allowing axial water flow between elements with minimal resistance), lignified secondary cell walls (providing mechanical strength and preventing collapse under negative pressure), dead cellular contents at maturity (protoplasm degraded leaving hollow lumen for water flow), and lateral bordered pits (enabling water exchange with adjacent vessels and ray parenchyma). Xylem vessel diameter varies by wood type and position= earlywood vessels 120-300 μm diameter (large vessels formed early season providing maximum hydraulic conductance), latewood vessels 40-120 μm diameter (smaller vessels formed late season providing backup capacity and mechanical support), vessel diameter declining from outer to inner rings (recent rings showing larger maximum vessels than old heartwood rings), cultivar variation (some varieties characteristically producing larger or smaller vessels), and site effects (water-stressed sites potentially producing smaller vessels than well-watered sites). Xylem vessel length varies substantially= individual vessel elements 200-600 μm long (single cellular units comprising vessels), vessels formed by vertical element series (ranging from few centimeters to several meters total vessel length), Vitis characterized as long-vesseled (individual continuous vessels potentially reaching 1-10+ meters length in some trunk sections), end wall perforation enabling continuity (large openings in perforation plates providing low-resistance connections between elements), and vessel length affecting embolism spread (long vessels potentially enabling rapid embolism propagation versus short vessels providing compartmentalization). Xylem vessel hydraulic function includes= water transport from roots to leaves (transpirational water loss creating tension pulling water through vessel network), hydraulic conductance proportional to radius⁴ (Hagen-Poiseuille equation showing large vessels disproportionately important for total conductance), sap flow velocity 0.1-5 m/hour (depending on transpiration demand, vessel diameter, and trunk position), xylem sap composition (water plus dissolved minerals, small amount of organic compounds), and tension in water columns (negative pressure -0.5 to -3.0 MPa typical during transpiration pulling water upward). Xylem vessel embolism vulnerability includes= air seeding when tension exceeds vessel strength (water column breaking forming gas bubble blocking vessel), larger vessels more vulnerable (wider vessels having larger pit membrane area and experiencing higher tensions increasing cavitation probability), embolism spreading through pit connections (air bubble in embolized vessel potentially spreading to adjacent vessels through pit membranes), progressive summer embolism (increasing percentage of vessels becoming embolized through growing season under water stress), and catastrophic embolism in severe stress (majority of vessels embolizing in extreme drought causing hydraulic failure). Xylem vessel anatomical variations include= perforation plate types (simple perforation plates with single large opening dominant in Vitis, compared to scalariform plates with multiple bars found in some woody plants), pit types (bordered pits with overarching secondary wall and torus-margo structure in vessel-vessel pits, simpler pits in vessel-parenchyma boundaries), vessel grouping patterns (vessels solitary or in radial multiples of 2-4, tangential bands in earlywood), and vessel distribution (ring-porous pattern with earlywood vessel concentration contrasting latewood). Xylem vessel development and maturation includes= vessel elements differentiating from cambial fusiform initials (derivatives of cambial cells specializing into vessel precursors), secondary wall deposition (lignified cell wall formation with characteristic patterns leaving primary wall regions as pits), perforation plate formation (end wall degradation creating large openings), protoplasm degradation (programmed cell death removing cellular contents creating hollow lumen), and vessel maturation (completed dead hollow tubes ready for water transport within weeks of cambial production). Xylem vessel density and arrangement includes= vessel density 25-60 vessels/mm² (varying by wood position, ring width, and cultivar), vessels occupying 30-50% of xylem cross-sectional area (highest in earlywood, lower in latewood), vessels arranged in radial or tangential patterns (earlywood showing tangential vessel bands, latewood more scattered), and vessel-vessel connections through pit pairs (bordered pits enabling lateral water movement between adjacent vessels). Xylem vessel significance in vine physiology includes= determining hydraulic capacity (vessel number, diameter, and functionality controlling maximum water transport rate), limiting canopy size (vessel hydraulic conductance ultimately constraining supportable leaf area), vulnerability determining drought tolerance (embolism resistance affecting vine performance under water stress), vessel characteristics affecting vintage variation (summer embolism extent influencing vine water status and fruit development), and vessel dysfunction causing vine decline (progressive vessel loss from disease or aging reducing hydraulic capacity causing vigor decline).
Xylem_Vessel emphasizes complete tubular conducting structure and is distinguished from vessel elements (individual cellular units) and specific vessel types (Early_Wood_Xylem_Vessel, Late_Wood_Xylem_Vessel, Large_Diameter_Xylem_Vessel) by encompassing all water-conducting tubes regardless of position or dimensions. Critical for understanding trunk hydraulic capacity, water transport mechanisms, embolism vulnerability, drought stress physiology, and hydraulic limitations on vine size and productivity.
Conducting vessel
Tracheary element tube
Tracheen-Röhre
Wasserleitgefäß
Water vessel
Xylemgefäß
Xylem Vessel
YAN data for Pinot Noir from Carneros, Napa Valley (2023 vintage) records 155 mg N/L at 24.5° Brix, categorized as "low" against the recommended threshold of 250 mg N/L for the expected 14.5% alcohol fermentation, with a calculated nitrogen deficit of 95 mg N/L informing a staged supplementation strategy of 0.20 g/L DAP at 16° Brix and 0.10 g/L Fermaid-O at 10° Brix. YAN data for Riesling from the Pfalz (2024 vintage) documents 240 mg N/L at 85° Oechsle, classified as "adequate" for the projected 11.5% alcohol dry fermentation, with no supplementation planned and the favorable nitrogen status expected to support clean, reductive fermentation preserving primary terpene aromas. YAN data for Zweigelt from Carnuntum (2024 vintage) records 195 mg N/L at 18° KMW, marginally sufficient for the 12.5% alcohol target, with the winemaker opting for a small organic nitrogen supplement (0.15 g/L yeast autolysate) at the onset of stationary phase to ensure complete dryness.
YAN data is a nitrogen data entity that records the concentration of yeast assimilable nitrogen in grape must, calculated as the sum of ammonium nitrogen and primary amino nitrogen (excluding proline and hydroxyproline), determined by formol titration, enzymatic assay, or the combination of specific ammonium and α-amino nitrogen quantifications, expressed in milligrams of nitrogen per liter (mg N/L), and serving as the single most important fermentation nutrient metric for predicting fermentation performance, determining supplementation requirements, and managing the risk of problematic fermentation outcomes.
YAN has become the universally recognized shorthand metric for must fermentation nutrient status since its formalization in the 1990s, displacing earlier approaches based on total nitrogen or individual amino acid analysis. The formol titration method (Sörensen method) provides a rapid estimate of total α-amino nitrogen but requires correction for ammonium (which also reacts with formaldehyde) and is subject to interference from sulfur dioxide additions — many laboratories now prefer the combination of specific enzymatic ammonium assay plus NOPA (o-phthaldialdehyde/N-acetyl-L-cysteine) primary amino nitrogen assay for improved accuracy. The categorization of YAN levels into deficient (350 mg N/L) provides general guidance, but optimal levels are context-dependent — yeast strain nitrogen demand varies 2-fold across commercial Saccharomyces cerevisiae strains, and the desired aroma profile influences nitrogen management strategy. Research has demonstrated that the ratio of ammonium to amino acid nitrogen influences fermentation-derived flavor compound production — musts with high ammonium-to-PAN ratios tend to produce lower ester concentrations and higher higher-alcohol levels than amino acid-rich musts at equivalent total YAN. The timing of nitrogen availability during fermentation also matters — yeast preferentially consume ammonium before amino acids (nitrogen catabolite repression), and staged nitrogen additions during fermentation can enhance aroma complexity compared with single pre-fermentation supplementation. Vineyard management directly influences YAN through soil nitrogen availability, cover crop competition, irrigation-mediated nitrogen mobilization, and crop load effects on nitrogen partitioning to fruit.
YAN value
YAN-Wert
assimilable nitrogen content
assimilierbarer Stickstoff
available nitrogen
fermentable nitrogen
gärfähiger Stickstoff
hefeverwertbarer Stickstoff
verfügbarer Stickstoff
yeast assimilable nitrogen
Bell SJ, Henschke PA (2005) Implications of nitrogen nutrition for grapes, fermentation and wine. Australian Journal of Grape and Wine Research 11(3): 242-295 Dukes BC, Butzke CE (1998) Rapid determination of primary amino acids in grape juice using an o-phthaldialdehyde/N-acetyl-L-cysteine spectrophotometric assay. American Journal of Enology and Viticulture 49(2): 125-134 Ugliano M et al. (2007) Effect of nitrogen supplementation on hydrogen sulfide and other volatile sulfur compounds. Journal of Agricultural and Food Chemistry 55(15): 6231-6239
YAN Data
Yeast assimilable nitrogen quality management in Marlborough Sauvignon Blanc targets YAN levels of 250-350 mg N/L to support the complete, vigorous fermentation required for varietal thiol expression, with vineyard nitrogen protocols (60-80 kg N/ha applied in spring) calibrated to achieve target must YAN winemakers supplement YAN-deficient musts (below 200 mg N/L) with organic nitrogen sources (yeast hulls, inactivated yeast) in preference to DAP to provide amino acid nutrition supporting the complex yeast metabolism that maximizes thiol release from cysteine conjugate precursors.
Yeast_Assimilable_Nitrogen_Quality_(YAN) represents the chemical quality characterizing the concentration of nitrogen compounds directly available for uptake and metabolism by Saccharomyces cerevisiae during alcoholic fermentation, capturing the nutritionally functional nitrogen fraction that determines fermentation performance. YAN comprises two main components: ammonium nitrogen (NH₄⁺) from inorganic ammonium salts, and alpha-amino nitrogen from free amino acids excluding proline (which Saccharomyces cerevisiae cannot metabolize under anaerobic fermentation conditions). The YAN concept emerged from recognition that total nitrogen poorly predicts fermentation behavior because substantial nitrogen fractions (proteins, proline, nucleic acids) are inaccessible to wine yeast. YAN concentration in grape must typically ranges from 100-400 mg N/L, with values below 140 mg N/L considered deficient and likely to cause fermentation problems, 140-250 mg N/L adequate for most fermentation conditions, and above 300 mg N/L potentially excessive. The consequences of YAN deficiency include sluggish or stuck fermentation, hydrogen sulfide production (as yeast scavenge sulfur-containing amino acids), and reduced ester formation limiting fruity aroma development. Conversely, excessive YAN can produce elevated volatile acidity, increased ethyl carbamate risk (from arginine-derived precursors), and wasteful nitrogen that remains in wine providing microbial nutrients. Winemakers routinely measure YAN pre-fermentation to assess supplementation needs, with diammonium phosphate (DAP) and complex organic nitrogen products providing options for correcting deficiencies. The YAN requirement varies with fermentation conditions, increasing with must sugar content, low temperature, high alcohol tolerance requirements, and yeast strain demands.
Yeast_Assimilable_Nitrogen_Quality_(YAN) serves as the parent class for Ammonium_Content_Quality and Primary_Amino_Nitrogen_Quality, the two components summed to yield YAN. The parenthetical (YAN) follows the uploaded structure naming convention and provides the widely-used abbreviation. YAN measurement employs enzymatic methods (formol titration for amino nitrogen, enzymatic ammonium) or HPLC amino acid analysis summed appropriately. The proline exclusion from amino nitrogen calculation reflects its non-assimilability by Saccharomyces under anaerobic conditions, though some non-Saccharomyces yeasts can utilize proline aerobically.
Hefeverwertbarer Stickstoff
YAN content
YAN-Gehalt
assimilable nitrogen quality
assimilierbarer Stickstoffgehalt
fermentable nitrogen concentration
gärfähige Stickstoffkonzentration
yeast-available nitrogen
Bely M et al. (1990) Description of Alcoholic Fermentation Kinetics: Its Variability and Significance. American Journal of Enology and Viticulture 41(4): 319-324 Torrea D et al. (2011) Comparison of Inorganic and Organic Nitrogen Supplementation of Grape Juice—Effect on Volatile Composition and Aroma Profile of a Chardonnay Wine Fermented with Saccharomyces cerevisiae Yeast. Food Chemistry 127(3): 1072-1083
Yeast Assimilable Nitrogen Quality (YAN)
Yellow hue development in Sémillon berries provides critical harvest timing indicators for Sauternes botrytized wine production, with healthy fruit displaying pale green-yellow hue (CIELAB h° 95-105°) that intensifies to golden-yellow (h° 80-90°) as noble rot infection concentrates sugars and carotenoids the deep amber-yellow hue (h° 70-80°) of fully botrytized berries signals optimal concentration for harvest of the passerillé fruit that will produce the complex, honeyed character of great Sauternes vintages.
Yellow_Hue_Quality represents the chromatic character of grapevine tissues displaying coloration in the yellow portion of the visible spectrum, corresponding to wavelengths approximately 570-590 nm and CIELAB hue angles of approximately 70-100°. Yellow hue results primarily from carotenoid pigments, including beta-carotene, lutein, and xanthophylls, which absorb blue and some green light while reflecting yellow and red wavelengths. In grapevine, yellow hue appears prominently in several contexts: autumn leaf senescence when chlorophyll degradation unmasks carotenoids mature berries of yellow-fruited cultivars young shoot tips in some cultivars (captured in OIV descriptor 003) and flowers. The golden-yellow coloration of many white wine grapes at maturity represents carotenoid pigments in the berry skin, with cultivars ranging from pale green-yellow (Pinot Grigio) through golden yellow (Chardonnay) to deep amber-yellow (Petit Manseng). Yellow hue intensity increases during berry ripening as chlorophyll degrades and carotenoids become the dominant visible pigments. In leaves, yellow coloration signals senescence and nutrient remobilization, though premature yellowing indicates stress conditions including nitrogen deficiency, water deficit, viral infection, or root dysfunction. Yellow hue in young shoot tips provides an ampelographic character, with some cultivars displaying bronzy-yellow tips while others show green or reddish coloration. Assessment employs visual comparison to color standards or instrumental measurement of hue angle in the yellow spectral region.
Yellow_Hue_Quality represents the yellow region of the color spectrum and is particularly relevant for white (green-skinned) grape cultivars where yellow represents the ripe coloration. The boundary between green-yellow and pure yellow hues involves continuous variation categorical assignment requires reference to standardized color charts or hue angle ranges. Yellow hue should be distinguished from the yellowing associated with chlorosis (nutrient deficiency or disease), which typically involves loss of pigmentation intensity as well as hue shift. OIV descriptors for berry skin color include yellow-green and yellow categories.
Gelbton
Karotinoidfärbung
amber hue quality
carotenoid coloration
gelbe Färbung
golden hue
goldene Tönung
yellow color
Mendes-Pinto MM (2009) Carotenoid Breakdown Products the Norisoprenoids in Wine Aroma. Archives of Biochemistry and Biophysics 483(2): 236-245 OIV (2009) OIV Descriptor List for Grape Varieties and Vitis Species, 2nd Ed, Organisation Internationale de la Vigne et du Vin, Paris pp.: 83-87
Yellow Hue Quality
Yield component analysis of Pinot Noir clones in Carneros revealed that the 40% yield difference between clone 667 (7.2 t/ha) and clone 115 (4.3 t/ha) was attributable primarily to cluster weight (145 g vs. 88 g) rather than cluster number per vine (48 vs. 46), with clone 115's lower cluster weight driven by both fewer berries per cluster (62 vs. 95) and smaller berry size (0.98 g vs. 1.12 g), the latter contributing to its higher skin-to-pulp ratio and more concentrated phenolic extraction. Yield components for Grüner Veltliner clone 2 versus clone 17 at Langenlois documented identical cluster counts (22-24 per vine) but divergent cluster architecture — clone 2 producing larger, more compact clusters (210 g, compactness OIV 204 score 7) versus clone 17's smaller, looser clusters (165 g, OIV 204 score 5), with the latter's reduced compactness conferring significant Botrytis resistance advantage.
Yield components is a yield performance data entity that records the individual contributing factors whose product determines total fruit production of a clonal line, including the number of clusters per vine (fruitfulness), average cluster weight, number of berries per cluster, average berry weight, and the hierarchical subcomponents of each (buds per vine, shoots per bud, inflorescences per shoot, flowers per inflorescence, fruit set percentage, seeds per berry), enabling diagnostic decomposition of yield differences between clones into their underlying morphological and physiological determinants.
Yield component analysis provides the mechanistic understanding of why clones differ in productivity, enabling targeted management responses and informed clone selection for specific production objectives. The multiplicative yield equation (yield = vines/ha × clusters/vine × berries/cluster × berry weight) defines a hierarchy of components determined at successive phenological stages — bud fruitfulness is established during the previous season's flower initiation (BBCH 53-57 of the preceding year), cluster number is fixed at budburst, berries per cluster are determined by fruit set (BBCH 71), and berry weight develops through cell division and expansion phases (BBCH 71-85). This temporal separation means that different yield components respond to different environmental signals and management interventions. Clonal differences in yield components are often more stable across environments than total yield itself, because the compensatory relationships between components (e.g., fewer clusters with larger berries) buffer total yield variation. Berry weight deserves particular attention as a yield component because it simultaneously influences both yield and quality — smaller berries increase the skin-to-juice ratio, enhancing extraction of phenolics, anthocyanins, and flavor compounds, creating the well-established inverse relationship between berry size and wine concentration in red cultivars.
Erntestrukturanalyse
Ertragsfaktoren
Ertragskomponenten
Ertragskomponentenanalyse
Ertragsstruktur
crop components
crop composition data
production components
yield breakdown
yield component analysis
Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 235-260 Dry PR et al. (2010) Bunch architecture and the susceptibility of Vitis vinifera to Botrytis cinerea. In Proceedings of the 7th International Workshop on Grapevine Downy and Powdery Mildew, IOBC-WPRS Bulletin 67: 53-56 Houel C et al. (2015) Genetic variability of berry size in the grapevine. Australian Journal of Grape and Wine Research 21(1): 36-47
Yield Components
Yield consistency assessment comparing established and developing vineyard blocks in Merlot reveals the maturity effect on production reliability, with mature block (15 years old, uniform soil) showing high consistency (annual yield CV 12% over 8 vintages, spatial CV 15% across zones), while developing block (5 years old, variable soil) shows low consistency (annual CV 35%, spatial CV 42%), with the consistency difference affecting management approaches—mature block enabling fixed contracts and standard planning while developing block requires flexible arrangements and zone-specific interventions to address the variable production pattern.
Yield consistency quality is a yield quality that represents the reliability and predictability of grape production across time and space, characterizing the stability of yields from year to year and the uniformity of production across different vines and vineyard zones, with consistency affecting both economic planning and quality management through its determination of how reliably production targets can be achieved. Within the BFO framework, this quality addresses the reliability dimension of yield, distinguishing the variation patterns from the absolute yield level while recognizing that consistent yields enable more accurate forecasting, more efficient resource allocation, and more stable business operations than highly variable production, even if average yields are similar. The ontological significance lies in yield consistency determining the predictability of vineyard output that affects every aspect of production planning, where consistent yields enable confident harvest scheduling, winery capacity planning, contract fulfillment, and financial projections, while inconsistent yields create uncertainty requiring contingency planning, flexible arrangements, and acceptance of variable outcomes that complicate business management. The temporal dimension of consistency addresses year-to-year variation arising from weather differences, pest and disease pressure, and cumulative management effects, with some vineyards showing stable production across years while others fluctuate dramatically with vintage conditions. The spatial dimension addresses within-vineyard variation arising from soil heterogeneity, microclimate differences, and vine-to-vine variation in condition and productivity, with uniform vineyards producing similar yields across all zones while variable vineyards show substantial spatial yield patterns requiring zone-specific management. The factors affecting yield consistency include site uniformity (consistent soil and microclimate promoting uniform production), vine age and establishment (mature, well-established vines showing more stable yields), management consistency (uniform practices promoting uniform outcomes), and cultivar characteristics (some varieties inherently more stable than others). Understanding yield consistency quality enables assessment of production reliability, identification of factors creating variation that might be addressable, and appropriate planning that accounts for the actual consistency level of specific vineyards.
Yield_Consistency_Quality serves as the parent class for temporal and spatial yield stability dimensions. This quality addresses production reliability affecting planning and management efficiency. The class supports risk assessment and operational planning through yield consistency characterization.
Ernteberechenbarkeit
Ertragskonsistenz
Ertragsreliabilität
Produktionsstabilität
harvest predictability
production stability
yield reliability
yield uniformity
Bramley RGV (2005) Understanding variability in winegrape production systems 2. Within vineyard variation in quality over several vintages. Australian Journal of Grape and Wine Research 11(1): 33-42 White RE (2015) Understanding Vineyard Soils, 2nd Ed. Oxford University Press, Oxford, pp. 1-280
Yield Consistency Quality
Yield efficiency assessment comparing management approaches in Pinot Noir reveals the balance implications of different efficiency targets, with low-efficiency management (heavy crop thinning, Ravaz index 4) producing 4 t/ha with excessive vigor requiring summer hedging, optimal-efficiency management (moderate thinning, Ravaz index 7) producing 7 t/ha with balanced canopy and timely ripening, and high-efficiency management (minimal thinning, Ravaz index 12) producing 10 t/ha but with delayed ripening, poor color, and declining vine reserves suggesting unsustainable overcropping, demonstrating how yield efficiency assessment identifies the production level optimizing quality and sustainability.
Yield efficiency quality is a yield quality that represents the relationship between grape production and the vine resources supporting that production, characterizing how effectively vine capacity is converted into fruit yield and whether the balance between vegetative and reproductive growth supports sustainable, quality-focused production. Within the BFO framework, this quality addresses the efficiency dimension of yield, distinguishing the productivity relative to resources from absolute yield while recognizing that yield efficiency captures the vine balance concept central to quality viticulture—efficient vines fully utilizing their capacity for fruit production without exceeding sustainable limits, while inefficient vines either waste capacity on excessive vegetation or stress themselves through overproduction beyond what leaf area can support. The ontological significance lies in yield efficiency determining both current fruit quality and long-term vine sustainability, where appropriate efficiency achieves the quality benefits of balanced source-sink relationships (adequate photosynthate for full ripening, appropriate vigor for canopy management) while maintaining the carbohydrate reserves and structural investment needed for continued productivity, and where efficiency imbalance creates either immediate quality problems (overcropping stress) or future productivity problems (depleted reserves, declining vigor). The metrics for assessing yield efficiency include the Ravaz index (yield-to-pruning-weight ratio), leaf area-to-fruit ratio, and various source-sink indicators that compare productive output against capacity indicators, with optimal ranges for these metrics varying somewhat with variety, climate, and production objectives but generally showing clear zones of balanced efficiency versus over- or under-utilization. The management of yield efficiency involves matching crop load to vine capacity through pruning (setting potential bud number), cluster thinning (adjusting actual cluster load), and canopy management (optimizing leaf area for fruit support), with efficiency optimization targeting the balance point where yield is maximized within quality and sustainability constraints. Understanding yield efficiency quality enables assessment of vine balance, diagnosis of over- or under-cropping conditions, and management calibration targeting optimal efficiency for sustainable quality production.
Yield_Efficiency_Quality serves as the parent class for specific efficiency metrics (Ravaz Index, Leaf Area Ratio, Source-Sink Balance). This quality addresses productive output relative to vine capacity affecting quality and sustainability. The class supports vine balance optimization through yield efficiency characterization.
Ertrags-Kapazitäts-Verhältnis
Ertragseffizienz
Produktionseffizienz
Rebeneffizienz
production efficiency
productive efficiency
vine efficiency
yield-to-capacity ratio
Yield Efficiency Quality
Yield maintenance under infection quality evaluation for powdery mildew in Central Valley table grape production compared the relationship between Erysiphe necator infection severity and marketable yield across cultivars to identify production-stable genotypes for reduced-spray programs. Controlled studies established a range of powdery mildew severity levels (10%, 25%, 50% cluster coverage) across cultivars through differential fungicide protection, then assessed yield and quality outcomes relative to fully protected controls. Thompson Seedless showed superior yield maintenance under infection quality with 25% cluster powdery mildew severity causing only 12% reduction in marketable yield through a combination of berry compensation (unaffected berries enlarging to partially offset losses from severely affected berries) and surface scarring that reduced grade but not total tonnage. Crimson Seedless showed intermediate yield maintenance under infection quality with 22% yield reduction at 25% severity, while Flame Seedless demonstrated poor yield maintenance with 38% yield reduction at equivalent severity due to rachis infection causing cluster drop and poor compensation for berry losses. The yield maintenance under infection quality assessment established economic thresholds, with Thompson Seedless production remaining profitable at severity levels up to 30% where spray cost savings exceeded yield losses, while Flame Seedless required maintenance below 15% severity for acceptable economic outcomes. The yield maintenance under infection quality characterization informed cultivar-specific disease thresholds for integrated pest management programs, calibrating management intensity to the tolerance capacity of each cultivar rather than applying uniform thresholds across all cultivars.
Yield maintenance under infection quality is a disease tolerance quality that inheres in a grapevine organism and represents the capacity to sustain grape production at acceptable levels despite pathogen infection through compensatory growth, efficient resource reallocation, and maintenance of reproductive development, achieving yield outcomes superior to what infection severity would predict based on susceptible cultivar performance at equivalent disease levels. This quality captures the productivity dimension of tolerance, recognizing that from a commercial perspective the ability to produce acceptable crops despite disease pressure may be more valuable than resistance that fails under high pressure, particularly for diseases where complete control is difficult or economically impractical. The ontological significance of yield maintenance under infection quality lies in its direct economic relevance, where cultivars maintaining yield under infection provide production security in disease-endemic environments and may reduce dependence on intensive fungicide programs if acceptable yields can be achieved at infection levels that would devastate intolerant cultivars. The biological basis of yield maintenance under infection encompasses the physiological compensatory mechanisms including enhanced photosynthesis in unaffected leaf area, preferential allocation of carbohydrates to reproductive organs, maintenance of fruit set and berry development despite infection-induced stress, and the architectural features that provide redundant capacity for loss compensation. The assessment of yield maintenance under infection requires experimental designs establishing equivalent infection levels across cultivars followed by yield comparison, or field observations relating natural variation in infection level to yield outcomes within and across cultivars. The compensation mechanisms underlying yield maintenance may have limits, with compensation capacity exhausted at high infection levels such that tolerant cultivars show progressive yield loss as disease severity increases, distinguishing yield maintenance under moderate infection from immunity to yield loss.
Yield_Maintenance_Under_Infection_Quality is_a Disease_Tolerance_Quality, representing the capacity to sustain acceptable crop production despite infection through compensatory mechanisms. This class addresses the productivity dimension of tolerance with direct economic relevance. Compensation mechanisms include enhanced photosynthesis, preferential allocation to fruit, and architectural redundancy. Assessment requires yield comparison under equivalent infection levels across cultivars. Economic thresholds for disease management can be cultivar-specific based on tolerance capacity. Compensation capacity has limits tolerance does not imply immunity to yield loss at high severity.
Ertragserhaltung unter Infektion
Ertragsstabilität bei Krankheit
crop stability under disease
harvest maintenance
productive tolerance
produktive Toleranz
Ney B et al. (2013) Crop tolerance to diseases: concepts, measurement, and application. Phytopathology 103:1163-1173 Pagán I, García-Arenal F (2018) Tolerance to plant pathogens: theory and experimental evidence. International Journal of Molecular Sciences 19:810 Calonnec A et al. (2004) Effects of Uncinula necator on the yield and quality of grapes (Vitis vinifera) and wine. Plant Pathology 53:434-445
Yield Maintenance Under Infection Quality
Yield per hectare assessment comparing planting systems in Pinot Noir reveals the density-yield relationship affecting quality outcomes, with high-density planting (7,500 vines/ha at 2.5 kg/vine) producing 18.75 t/ha from many small-cropped vines showing uniform ripening and concentrated fruit, while low-density planting (2,000 vines/ha at 9.0 kg/vine) produces 18.0 t/ha from fewer heavily-cropped vines showing delayed ripening and diluted composition, demonstrating how similar per-hectare yields can result from different vine configurations with substantially different quality outcomes, making per-hectare yield alone insufficient for quality assessment without considering the underlying planting and per-vine yield patterns.
Yield per hectare quality is a quantity yield quality that represents the total mass of fruit harvested per unit land area, characterizing the area-based productivity that enables comparison among vineyards of different sizes, calculation of production economics on a land basis, and regulatory compliance with appellation yield limits that typically specify maximum tonnes per hectare. Within the BFO framework, this quality specifically addresses the land area dimension of yield, distinguishing area-based production from per-vine metrics while recognizing that yield per hectare integrates both per-vine productivity and planting density, such that the same per-hectare yield can result from many low-yielding vines or fewer high-yielding vines with different implications for vine balance and fruit quality. The ontological significance lies in yield per hectare being the standard metric for vineyard productivity comparison, economic analysis, and regulatory specification, where per-hectare yields determine gross production value per land unit, enable benchmarking against regional or appellation averages, and must comply with any yield restrictions imposed by quality designation systems (such as AOC limits in France typically ranging from 35-65 hL/ha depending on appellation). The calculation of yield per hectare involves multiplying yield per vine by planting density (vines per hectare), or dividing total vineyard harvest by area, with results expressed in tonnes per hectare (t/ha) or hectoliters per hectare (hL/ha, common in European regulation where 100 kg grapes ≈ 75 L juice/wine), with normal ranges from 3-6 t/ha for premium production through 8-15 t/ha for standard quality to 20-30 t/ha for high-yielding juice or table grape production. The planting density interaction with per-hectare yield shows that high-density plantings (5,000-10,000 vines/ha) can achieve similar per-hectare yields to low-density plantings (1,000-2,500 vines/ha) while maintaining lower per-vine yields, potentially improving vine balance and fruit quality at equivalent area productivity, though with higher establishment and management costs. Understanding yield per hectare quality enables vineyard productivity assessment, economic analysis of land utilization, and compliance management for appellation yield requirements.
Yield_Per_Hectare_Quality addresses the area-based productivity enabling economic and regulatory yield assessment. This quality integrates per-vine yield with planting density for land-unit production metrics. The class supports productivity benchmarking and appellation compliance through yield per hectare characterization.
Ertrag pro Hektar
Flächenertrag
Hektarertrag
Produktivität pro Hektar
area yield
hectare yield
production per hectare
vineyard productivity
Dry PR, Coombe BG (2004) Viticulture Volume 1: Resources, 2nd Ed. Winetitles, Adelaide, pp. 150-200 Reynolds AG (2010) Managing Wine Quality, Volume 1: Viticulture and Wine Quality. Woodhead Publishing, Cambridge, pp. 1-57
Yield Per Hectare Quality
Yield per vine assessment evaluating crop load in Zinfandel reveals the balance implications of per-vine production, with low-yielding vines (3.5 kg/vine, Ravaz index 4.5) showing excessive vegetative vigor and shading from under-utilization of capacity, balanced vines (6.0 kg/vine, Ravaz index 7.2) achieving target ripeness with appropriate vigor, and over-cropped vines (10.5 kg/vine, Ravaz index 13.5) showing delayed ripening, reduced reserves, and declining vigor over subsequent seasons, demonstrating how yield per vine assessment identifies the production level appropriate for sustainable vine balance.
Yield per vine quality is a quantity yield quality that represents the total mass of fruit harvested from individual grapevines, characterizing the per-plant productivity that provides the fundamental unit of yield measurement from which area-based yields are calculated and that reflects the integrated outcome of vine capacity, crop load, and seasonal growing conditions. Within the BFO framework, this quality specifically addresses the individual vine dimension of yield, distinguishing per-vine production from area-based metrics while recognizing that yield per vine is the direct measure of each plant's productivity and the scale at which many management decisions (pruning severity, cluster thinning) are implemented and evaluated. The ontological significance lies in yield per vine representing the balance between vine capacity (the carbohydrate production potential from leaf area) and reproductive load (the fruit sink demanding those carbohydrates), with optimal yield per vine achieving full utilization of vine capacity without exceeding it—excessive yield stressing the vine and compromising quality, while insufficient yield underutilizing capacity and sacrificing potential production. The calculation of yield per vine involves weighing the fruit harvested from each vine (or calculating from cluster count × average cluster weight), with results typically expressed in kilograms per vine (kg/vine), with normal ranges varying substantially with vine size, training system, and production objectives—from 1-3 kg/vine for low-yielding premium production to 15-25 kg/vine for high-capacity, high-yielding systems. The vine capacity relationship to appropriate yield is captured in the crop load concept, where yield per vine should be proportional to vine size (often assessed through pruning weight) to maintain vine balance, with the Ravaz index (yield/pruning weight ratio) providing a diagnostic tool with optimal ranges typically 5-10 for quality wine production, lower ratios indicating under-cropping, and higher ratios indicating overcropping stress. Understanding yield per vine quality enables assessment of individual vine productivity, evaluation of vine balance through crop load analysis, and management decisions targeting appropriate per-vine production for sustainable, quality-focused cultivation.
Yield_Per_Vine_Quality addresses the total fruit production per individual grapevine. This quality reflects vine capacity utilization and affects vine balance and sustainability. The class supports crop load management through yield per vine characterization.
Einzelstockproduktion
Ertrag pro Rebe
Pflanzenproduktivität
Rebstockertrag
individual vine crop
per-vine production
plant productivity
vine yield
Bravdo B et al. (1985) Effect of crop level on growth, yield and wine quality of a high yielding Carignane vineyard. American Journal of Enology and Viticulture 36(2): 125-131 Kliewer WM, Dokoozlian NK (2005) Leaf area/crop weight ratios of grapevines: influence on fruit composition and wine quality. American Journal of Enology and Viticulture 56(2): 170-181
Yield Per Vine Quality
Yield performance data for Pinot Noir clone 115 (Dijon) across 15 vintages in Oregon's Willamette Valley documents average production of 4.5-6.0 t/ha, approximately 25-30% below clone 777 (6.5-8.0 t/ha) under identical management, with the yield differential attributable primarily to smaller cluster size (85-110 g vs. 130-165 g) and higher millerandage incidence (15-25% shot berries). Yield performance data for Grüner Veltliner clone 17 at Klosterneuburg trial sites records 10-year mean production of 9.8 t/ha (CV 18%) with a yield-must weight regression coefficient of -0.85° KMW per additional t/ha, quantifying the quality-quantity trade-off essential for production target calibration.
Yield performance data is a performance data entity that documents the quantitative production capacity of a clonal line, encompassing total fruit weight per vine or per hectare, yield components (cluster number, cluster weight, berry number, berry weight), yield trends over vine age, and the relationship between yield level and fruit quality parameters, measured across multiple vintages in replicated evaluation trials and providing the primary agronomic characterization upon which clone productivity assessments and vineyard economic projections are based.
Yield performance data is the most frequently cited and economically consequential component of clone evaluation, as yield level directly determines vineyard revenue and production cost per unit of fruit. However, yield data requires careful contextual interpretation — absolute yield values are strongly influenced by vine age, spacing, training system, rootstock, irrigation regime, and crop load management decisions that may differ between evaluation trials and commercial practice. Standardized metrics such as yield per meter of cordon (kg/m) or Ravaz Index (yield-to-pruning-weight ratio, optimal range 5-10 for quality wine) provide more transferable comparisons than absolute yield per hectare. The temporal stability of yield across vintages is equally important to the mean yield level, as high-yielding but erratic clones create greater production and cash flow uncertainty than moderately yielding but consistent performers. Clone-by-rootstock interactions can significantly modify yield performance, necessitating evaluation on regionally relevant rootstock combinations rather than a single rootstock standard. The negative correlation between yield and quality, while generally observed, is not linear and exhibits clone-specific thresholds — some clones maintain quality metrics to substantially higher yield levels than others, a characteristic of considerable economic value.
Ernteertragsdaten
Ertragsdaten
Ertragsleistungsdaten
Klonertragsbewertung
Produktionsleistungsdaten
clone yield record
crop performance data
production performance
productivity data
yield evaluation data
Anderson MM et al. (2008) Viticultural evaluation of French and California Pinot Noir clones. American Journal of Enology and Viticulture 59(2): 188-193 Wolpert JA et al. (1994) Field performance of six Chardonnay clones in the Napa Valley. American Journal of Enology and Viticulture 45(4): 393-400 Keller M (2020) The Science of Grapevines, 3rd Ed, Academic Press, pp.: 235-260
Yield Performance Data
Yield potential assessment in Chardonnay tracking sequential potential losses reveals the determination stages affecting final yield, with bud fruitfulness providing initial potential of 2.1 clusters/bud (42 clusters from 20 retained buds), flower development reducing potential through some inflorescence abortion to 38 clusters, fruit set at 35% establishing berry potential of approximately 13,300 berries (38 clusters × 1000 flowers/cluster × 0.35 set), with subsequent berry development determining final yield of 4.8 kg/vine from this potential framework, demonstrating how yield potential is established through sequential stages each representing opportunity for potential loss.
Yield potential quality is a yield quality that represents the maximum production achievable given the reproductive capacity established during bud differentiation and flower development, characterizing the upper limit of yield set by bud fruitfulness, flower number, and fruit set success before subsequent berry development determines actual harvest weight. Within the BFO framework, this quality addresses the capacity dimension of yield, distinguishing the potential production from actual realized yield while recognizing that yield potential establishes the ceiling that subsequent development can approach but not exceed—every flower that fails to set and every bud that lacks fruitfulness represents permanently lost yield potential for that season. The ontological significance lies in yield potential determining the framework within which yield management operates, where high potential provides flexibility for crop adjustment (thinning to target level) while low potential constrains production regardless of subsequent conditions, with potential assessment enabling early-season yield prediction and identification of factors limiting reproductive capacity that might be addressed for future seasons. The components of yield potential include bud fruitfulness (clusters per bud, determined during bud differentiation the previous year), flower number per inflorescence (determined during late bud development), and fruit set percentage (determined during flowering by pollination success and post-fertilization berry retention), with each component representing a sequential gate through which potential must pass—failure at any stage permanently reducing potential for that season. The temporal determination of yield potential occurs across two seasons, with cluster initiation during summer of year N establishing the framework expressed in year N+1, and with conditions during both the initiation period and the expression period affecting whether potential is fully realized, partially achieved, or lost to various limitations. Understanding yield potential quality enables early-season yield forecasting, diagnosis of factors limiting reproductive success, and management targeting the specific stages where potential is established or lost.
Yield_Potential_Quality serves as the parent class for reproductive capacity components (Bud Fruitfulness, Fruit Set, Coulure, Millerandage). This quality addresses the maximum achievable yield established before berry development. The class supports yield forecasting and reproductive success assessment through potential characterization.
Ertragsdecke
Produktionspotential
maximum achievable yield
production potential
reproduktive Kapazität
yield ceiling
Yield Potential Quality
Yield quality assessment comparing crop levels in Pinot Noir reveals the quantity-quality tradeoff informing management decisions, with high yield (10 t/ha) showing adequate cluster and berry parameters but compromised fruit quality (22.5 Brix, dilute color, extended ripening), moderate yield (6 t/ha) showing optimized quality (24.5 Brix, concentrated color, timely ripening) with economically viable production, and very low yield (3 t/ha) showing exceptional quality (26 Brix, intense concentration) but questionable economic return, demonstrating how yield quality assessment must integrate quantity and quality dimensions to identify the optimal production level.
Yield quality is a viticultural quality that represents the production output characteristics of grapevine cultivation, characterizing both the quantity of grapes produced and the compositional quality of that production, recognizing that yield encompasses multiple dimensions including total weight, component analysis (clusters, berries), and the quality attributes that determine the value and suitability of the crop for intended uses. Within the BFO framework, this quality addresses the productivity dimension of viticultural performance, distinguishing yield outcomes from the vine condition or management inputs while recognizing that yield represents the primary economic output of viticulture and the measure by which vineyard productivity is assessed, with both quantity (how much is produced) and quality (how good is what is produced) contributing to overall yield value. The ontological significance lies in yield quality determining the economic viability of grape production, where yield quantity affects gross production value, yield quality affects per-unit value (price per tonne varying substantially with quality indicators), and the optimization of yield quality involves balancing quantity against quality to maximize total value under given market conditions. The yield-quality relationship in viticulture shows the characteristic inverse correlation where very high yields often compromise fruit quality (through resource dilution, delayed ripening, reduced concentration) while very low yields may achieve exceptional quality but insufficient quantity for economic viability, with the management challenge being to find the yield level that optimizes value by balancing these opposing tendencies. The components of yield include the structural elements (cluster number × cluster weight, or cluster number × berries per cluster × berry weight) that together determine total production, with each component under different genetic and environmental control and responsive to different management interventions, enabling targeted yield manipulation through component-specific approaches. Understanding yield quality enables assessment of vineyard productivity, analysis of yield components to identify limiting factors, and management optimization balancing yield quantity against quality for maximum production value.
Yield_Quality serves as the parent class for quantity and compositional yield dimensions. This quality addresses the productivity outcomes determining economic viability and raw material value. The class supports production optimization through yield quality characterization balancing quantity and quality.
Ertragsqualität
Produktionsleistung
Traubenertrag
crop yield
grape yield
harvest quantity
production output
Keller M (2015) The Science of Grapevines: Anatomy and Physiology, 2nd Ed. Academic Press, San Diego, pp. 145-230 Reynolds AG (2010) Managing Wine Quality, Volume 1: Viticulture and Wine Quality. Woodhead Publishing, Cambridge, pp. 1-57
Yield Quality
Yield stability analysis for Merlot clones in Bordeaux over 15 vintages (2005-2019) revealed that clone 181 exhibited exceptional stability (CV 14%, Finlay-Wilkinson b = 0.92) maintaining 7-9 t/ha even in climatically challenging years, while clone 348 showed higher mean yield (10.5 t/ha) but poor stability (CV 32%, b = 1.45) with production swinging from 5 t/ha in frost-affected 2017 to 14 t/ha in favorable 2018. Yield stability assessment of Riesling clones at Geisenheim demonstrated that clone 198 (CV 16%) outperformed clone 64 (CV 28%) in production consistency over 12 vintages despite similar mean yields of 9.5 and 10.2 t/ha respectively, with clone 64's instability traced to pronounced alternate bearing tendencies.
Yield stability is a yield performance data entity that records the consistency, regularity, and predictability of fruit production by a clonal line across vintages and sites, quantified through statistical measures of temporal variability including coefficient of variation (CV), regression stability parameters (Finlay-Wilkinson regression coefficient, deviation from regression), and ecovalence, characterizing the degree to which a clone maintains its expected yield level under varying climatic conditions and providing critical information for production planning and economic risk assessment.
Yield stability is a distinct performance attribute from mean yield, and the two may be negatively correlated — high-yielding clones frequently exhibit greater vintage-to-vintage variability because their larger crop loads make them more susceptible to resource limitation in challenging years. The statistical characterization of yield stability draws on quantitative genetics methodologies developed for annual crop breeding, adapted for perennial vine evaluation where individual vintage observations represent the environmental gradient. The Finlay-Wilkinson regression coefficient (b) provides interpretable stability classification: b = 1.0 indicates average responsiveness to environment, b 1.0 indicates environmental sensitivity. Alternate bearing (biennial oscillation between high and low yield) represents a specific form of yield instability common in some cultivars, driven by competition between reproductive and vegetative growth, and clonal selection can identify genotypes with reduced alternate bearing tendency. For economic modeling, yield stability directly influences cash flow predictability, grape contract reliability, and winery supply planning — making it a key criterion alongside mean yield in commercial clone selection decisions.
Ertragskonstanz
Ertragsregelmäßigkeit
Ertragssicherheit
Ertragsstabilität
Produktionszuverlässigkeit
crop regularity
vintage-to-vintage yield variation
yield consistency
Finlay KW, Wilkinson GN (1963) The analysis of adaptation in a plant-breeding programme. Australian Journal of Agricultural Research 14(6): 742-754 Mannini F (2004) Clonal selection in grapevines. Acta Horticulturae 652: 21-27 Rühl EH et al. (2004) Grapevine clonal selection, evaluation, and management. Acta Horticulturae 652: 29-34
Yield Stability
Ravaz Index assessment tracking vine balance over multiple seasons in Merlot reveals the feedback between cropping level and subsequent capacity, with Year 1 showing overcropped condition (Ravaz 14, poor ripening, reduced pruning weight following season), Year 2 adjusted through cluster thinning to Ravaz 8 (improved ripening, recovery of pruning weight), Year 3 maintaining Ravaz 7 (optimal balance, full ripening at 24.5 Brix, stable pruning weight indicating sustainable capacity), demonstrating how Ravaz Index monitoring enables crop load calibration that achieves and maintains optimal vine balance.
Yield-to-pruning-weight ratio quality, commonly known as the Ravaz Index, is a yield efficiency quality that represents the relationship between harvested fruit weight and the weight of dormant prunings removed the following winter, characterizing the balance between reproductive and vegetative growth that indicates whether vines are appropriately cropped relative to their capacity or are over- or under-cropped with consequences for fruit quality and vine sustainability. Within the BFO framework, this quality specifically addresses the yield-pruning relationship dimension of efficiency, distinguishing this widely-used balance indicator from other efficiency metrics while recognizing that the Ravaz Index provides a practical, easily-measured assessment of vine balance that integrates the full season's growth and production into a single ratio reflecting whether carbohydrate partitioning favored fruit, vegetation, or was appropriately balanced. The ontological significance lies in the Ravaz Index capturing the fundamental vine balance that determines both immediate quality outcomes and long-term productivity, where the ratio directly reflects how the vine allocated its photosynthetic output between reproductive (fruit yield) and vegetative (shoot growth producing next season's capacity) functions, with this allocation determining whether resources were adequate for full fruit ripening and whether sufficient vegetative growth maintains vine capacity. The interpretation of Ravaz Index values follows established guidelines with optimal ranges typically 5-10 for quality wine production—ratios below 5 indicating undercropping (excessive vegetation, wasted capacity, potential shading problems), ratios of 5-10 indicating balanced cropping (appropriate resource allocation, sustainable production), and ratios above 10 indicating overcropping (stressed vines, compromised ripening, depleted reserves threatening future productivity), though optimal values vary somewhat with variety and climate. The calculation methodology involves dividing total fruit yield per vine (kg) by total pruning weight per vine (kg), both measured on the same vines across consecutive measurements (harvest in autumn, pruning the following winter), with the ratio being dimensionless and thus comparable across different vine sizes, training systems, and production levels. Understanding Ravaz Index quality enables straightforward vine balance assessment, identification of crop load adjustment needs, and tracking of balance trends over time to ensure sustainable production.
Yield_To_Pruning_Weight_Ratio_Quality_(Ravaz_Index) addresses the classic vine balance metric relating production to vegetative growth. This quality provides practical assessment of appropriate crop load relative to vine capacity. The class supports crop load management through Ravaz Index characterization with established optimal ranges.
Balance-Index
Ertrag-Schnittgewicht-Verhältnis
Ertragsindex
Ravaz Index
Ravaz-Index
crop load index
yield-pruning ratio
Ravaz L (1911) L'effeuillage de la vigne. Annales de l'Ecole Nationale d'Agriculture de Montpellier 11: 216-244 Bravdo B et al. (1985) Effect of crop level on growth, yield and wine quality of a high yielding Carignane vineyard. American Journal of Enology and Viticulture 36(2): 125-131
Yield To Pruning Weight Ratio Quality (Ravaz Index)
Pinot Noir showing young active feeder roots in June, soil cores revealing feeder roots 0.5-1.2 mm with characteristic white coloration, roots 2-6 weeks post-maturation in young active phase, microscopic examination showing dense cytoplasm, abundant organelles, maximum root hair density 340 per cm, developing arbuscular mycorrhizal colonization 25%, isotope tracing showing young feeder roots providing 62% of total feeder root nitrogen uptake despite comprising only 38% of feeder root length, young active feeder roots demonstrating peak absorption efficiency
Young active feeder root is active feeder root in early functional phase typically 1-8 weeks post-maturation, characterized by recent completion of development and differentiation, vigorous metabolism with high respiratory rates, maximum root hair production and density, peak uptake capacity per unit surface area, and white coloration indicating youth and active growth, with young active feeder roots representing most efficient absorption phase in feeder root lifespan showing highest specific uptake rates, most responsive capacity, greatest proliferation potential, and representing primary target for maximizing vine resource acquisition, exhibiting developmental characteristics including recently completed cell differentiation and maturation, minimal or absent wall thickening or secondary modifications, abundant dense cytoplasm with high organelle content, maximum root hair density potentially 200-400+ hairs per centimeter, developing arbuscular mycorrhizal colonization with arbuscule formation actively proceeding, and demonstrating that young active phase represents peak feeder root performance period where absorption capacity maximized relative to construction and maintenance costs creating highest return on investment for vine resource allocation to root production. Young active feeder root age definition includes= showing recent maturation (young feeder roots typically 1-8 weeks post-maturation having completed differentiation), exhibiting early functional period (young status representing first portion of active functional lifespan), demonstrating pre-senescence state (young roots far from senescence onset maintaining maximum vitality), revealing variable duration (young period varying with root order, environmental conditions, growth rate), and displaying transition character (young state transitioning to mature active then senescent states). Young active feeder root morphological indicators includes= showing white coloration (young feeder roots characteristically white or cream colored distinct from tan mature or brown senescent roots), exhibiting turgid appearance (young roots fully turgid reflecting optimal water status and cellular function), demonstrating smooth surface (young root surface smooth before irregular aging changes develop), revealing abundant root hairs (young feeder roots showing maximum root hair density creating fuzzy appearance), and displaying uniform diameter (young roots maintaining consistent diameter without irregular swelling or constrictions). Young active feeder root physiological characteristics includes= showing high metabolic rate (young feeder roots exhibiting elevated respiration supporting growth and function), exhibiting maximum uptake capacity (young roots showing highest absorption rates per unit surface area or length), demonstrating rapid growth potential (young feeder roots capable of continued elongation if conditions favorable), revealing responsive capacity (young roots most responsive to environmental signals adjusting quickly to conditions), and displaying positive carbon balance (photosynthate allocation to young roots supporting net growth and function). Young active feeder root cellular features includes= showing dense cytoplasm (young root cells with abundant cytoplasm occupying large proportion of cell volume), exhibiting abundant organelles (high mitochondrial, ER, Golgi density supporting active metabolism), demonstrating thin walls (primary cell walls thin and flexible without secondary thickening), revealing large nuclei (nuclei prominent relative to cell size indicating active gene expression), and displaying minimal vacuolation (vacuoles present but smaller than in mature cells leaving more cytoplasmic space). Young active feeder root absorption performance includes= providing highest specific uptake (young feeder roots showing maximum uptake per unit surface area or biomass), exhibiting nutrient responsiveness (young roots quickly increasing uptake when nutrients available), demonstrating water acquisition (young roots contributing disproportionately to total water uptake given their efficiency), revealing preferential function (isotope studies often showing young roots contributing more than proportional to their abundance), and displaying efficiency advantage (young roots providing maximum absorption return per unit carbon investment). Young active feeder root root hair characteristics includes= showing maximum density (young feeder roots potentially producing 200-400+ root hairs per centimeter), exhibiting active growth (root hair population on young roots including hairs still elongating), demonstrating optimal function (young root hairs at peak absorption capacity with intact membranes and active transport), revealing extensive coverage (root hair zone on young roots potentially extending 2-5 cm behind tip), and displaying surface multiplication (young root hairs increasing effective absorption surface 10-25 fold). Young active feeder root mycorrhizal development includes= showing active colonization (young feeder roots being actively colonized by arbuscular mycorrhizal fungi), exhibiting arbuscule formation (developing arbuscules in cortex cells creating nutrient exchange interfaces), demonstrating hyphal growth (external mycorrhizal hyphae extending from young roots exploring surrounding soil), revealing establishment phase (mycorrhizal symbiosis establishing and developing toward mature functional state), and displaying benefit initiation (mycorrhizal enhancement of uptake beginning as arbuscules form). Young active feeder root in resource capture includes= determining proliferation success (young feeder root survival and continued function determining root system expansion), enabling patch exploitation (young roots rapidly proliferating in resource-rich patches capturing nutrients and water), demonstrating competitive ability (young vigorous feeder roots competing effectively for soil resources), revealing plasticity (young roots showing developmental plasticity responding to heterogeneous soil conditions), and displaying growth investment (vine allocating substantial photosynthate to young feeder root production and maintenance).
Young_Active_Feeder_Root emphasizes early functional phase and peak performance and is distinguished from mature active roots by focus on recent maturation and maximum capacity. Critical for understanding peak efficiency, white coloration, maximum root hairs, high metabolic rate, mycorrhizal development, and the recently matured feeder roots in early functional phase showing highest specific uptake capacity representing most efficient absorption period.
Early-stage active root
Jugendliche Absorptionswurzel
Junge aktive Feinwurzel
Juvenile absorptive root
Neu-funktionelle Saugwurzel
Newly functional feeder root
Young Active Feeder Root
Chardonnay grafted on SO4 rootstock at 2.5 years post-grafting showing young grafted rootstock trunk 3.2 cm diameter, trunk extending 21 cm from root crown to graft union positioned 17 cm above soil, graft union bulge 2.4 cm diameter with actively forming callus tissue, young trunk bark smooth light grey-brown with minimal fissuring and visible lenticels, diameter increased 0.9 cm current season indicating vigorous growth, trunk requiring continued stake support, no signs of disease or damage
Young grafted rootstock trunk is grafted rootstock trunk in early developmental stage spanning approximately first 3 years post-grafting, characterized by active establishment of graft union integration, rapid diameter increase as trunk builds structural capacity, smooth or minimally furrowed bark reflecting recent lignification, relatively small diameter typically 1.5-4.5 cm, and ongoing development of complete vascular continuity across graft union as rootstock and scion tissues progressively integrate into functional unit. Young grafted rootstock trunk age characteristics includes= spanning years 1-3 post-grafting (from initial grafting season through third growing season), representing early establishment phase (critical period of graft union formation and trunk development), showing rapid proportional growth (diameter increasing 0.4-1.2 cm annually compared to 0.3-0.6 cm in mature trunks), exhibiting relatively small absolute diameter (1.5-4.5 cm typical range at 3 years compared to 4-8 cm in mature trunks), and displaying juvenile wood properties (higher proportion of juvenile xylem with smaller vessels and less dense wood than mature trunk). Young grafted rootstock trunk graft union development includes= actively forming callus bridge (wound healing tissue continuously accumulating at graft interface during first 2-3 years), establishing vascular connections (xylem and phloem continuity progressively improving from incomplete initial connections to well-integrated transport pathways), showing prominent graft union bulge (callus tissue creating 1.5-3.0 cm diameter increase at graft union particularly visible in young stage), potentially displaying graft union scar (visible line marking original grafting point more prominent in young stage before bark thickening obscures it), and gradually stabilizing integration (graft union becoming mechanically and physiologically more robust with each growing season). Young grafted rootstock trunk bark characteristics includes= showing smooth or minimally developed bark (periderm formation just beginning with thin cork layer 0.3-1.2 mm thickness), displaying light coloration (young bark typically lighter tan or greenish-brown before weathering to grey-brown), exhibiting minimal fissuring (bark surface largely intact without deep longitudinal furrows characteristic of older trunks), revealing greenish undertones (chlorophyll potentially visible through thin bark in first-year trunks), and showing lenticels (gas exchange pores visible as small dots or horizontal lines on smooth young bark surface). Young grafted rootstock trunk structural development includes= building mechanical strength (rapidly increasing trunk diameter and wood density to support growing scion canopy), establishing stability (young trunk requiring staking support until achieving sufficient diameter and lignification for self-support typically by year 3), showing flexible properties (young wood more flexible and less brittle than mature wood allowing some bending without breaking), developing secondary xylem (active vascular cambium producing wood layers building trunk cross-sectional area), and forming initial annual rings (first 2-3 growth rings representing young trunk's early development visible in cross-section). Young grafted rootstock trunk vulnerability includes= susceptible to mechanical damage (thin bark and smaller diameter making young trunk more vulnerable to equipment injury, animal damage, or wind breakage), sensitive to freeze damage (limited cold hardiness in young trunk cambium and bark potentially leading to winter injury in severe cold), requiring protection from sunburn (young smooth bark vulnerable to southwest sun exposure potentially causing bark necrosis in hot climates), showing higher disease susceptibility (young trunk more vulnerable to crown gall infections at graft union and trunk cankers than mature trunks), and needing support (young trunk requiring stakes or trellising to maintain upright position and prevent wind damage until sufficient structural development). Young grafted rootstock trunk transport capacity includes= establishing water conduction (initially limited xylem cross-sectional area increasing as trunk diameter expands building transport capacity), developing nutrient transport pathways (phloem tissue increasing in cross-sectional area improving nutrient distribution between roots and scion), showing increasing flow rates (water transport capacity growing from 5-15 liters daily in year 1 to 20-40 liters daily by year 3 as xylem area expands), enabling resource distribution (improving capacity to transport photosynthates from scion leaves to root system for storage and root growth), and building reserves (young trunk beginning to accumulate carbohydrate reserves in ray parenchyma though storage capacity limited compared to mature trunk). Young grafted rootstock trunk management requirements includes= maintaining graft union protection (wrapping or painting young graft union in some climates preventing sunburn and freeze damage), ensuring adequate staking (supporting young trunk preventing wind damage until trunk achieves 3-4 cm diameter sufficient for stability), implementing trunk protection (wrapping, painting, or tubing young trunk in hot climates preventing sunburn damage), managing competing shoots (removing rootstock suckers below graft union that compete with establishing scion), providing optimal irrigation (regular water supply supporting rapid growth and graft union integration during establishment years), and monitoring for damage (regular inspection of vulnerable young trunk detecting mechanical injury, disease, or freeze damage early). Young grafted rootstock trunk growth patterns includes= showing rapid diameter increase (0.4-1.2 cm annual increment depending on rootstock vigor and growing conditions), displaying upward extension (young trunk elongating 2-8 cm first year primarily from graft union callus growth rather than intercalary elongation), exhibiting scion influence (scion vigor and canopy development affecting young trunk diameter growth through photosynthate supply), responding to training (young trunk development shaped by staking, pruning, and canopy management establishing permanent trunk form), and stabilizing by year 3 (growth rate beginning to moderate toward mature trunk growth patterns). Young grafted rootstock trunk in first-year grafts includes= showing initial callus formation (first weeks and months post-grafting critical for graft union establishment), establishing minimal bark (initially no periderm with just thin epidermis covering young trunk), displaying very small diameter (0.8-2.0 cm typical first-season range), requiring intensive protection (very vulnerable young trunk needing support, protection, and careful management), and beginning vascular integration (initial xylem and phloem connections forming across graft union though not yet fully functional). Young grafted rootstock trunk transition to mature status includes= diameter reaching 4-5 cm threshold (marking transition from young to mature trunk structural capacity), bark developing characteristic fissures (periderm formation creating textured bark typical of cultivar and trunk age), graft union integration stabilizing (callus formation largely complete with well-established vascular continuity), growth rate moderating (annual diameter increase declining from juvenile rapid growth to stable mature growth), and achieving structural independence (trunk no longer requiring stake support having achieved adequate strength and diameter). Young grafted rootstock trunk cultivar variation includes= vigorous rootstocks (1103P, 110R, 140Ru) showing rapid young trunk development potentially reaching 4 cm diameter by year 2, moderate vigor rootstocks (SO4, 3309C, Teleki 5C) showing intermediate young trunk growth reaching 3-4 cm by year 3, low vigor rootstocks (Riparia Gloire, 101-14) showing slower young trunk development potentially requiring 4 years to reach mature trunk dimensions, and rootstock-specific bark development (some rootstocks showing early bark texturing while others maintaining smooth bark longer). Young grafted rootstock trunk significance includes= representing critical establishment period (first 3 years determining long-term graft union success and vine foundation), requiring intensive management (young trunk vulnerability necessitating protection, support, and monitoring ensuring successful establishment), establishing long-term vine structure (young trunk development determining permanent trunk form and training system implementation), and marking transition phase (young grafted rootstock trunk representing progress from recently grafted vulnerable union to established integrated foundation).
Young_Grafted_Rootstock_Trunk emphasizes early developmental stage (years 1-3) and is distinguished from Mature_Grafted_Rootstock_Trunk (established integration and stable growth) by focus on active establishment phase. Critical for understanding graft union development, establishment requirements, vulnerability factors, management needs during critical early years, and the foundation-building phase of grafted vine development.
Early-stage grafted rootstock trunk
Establishing rootstock trunk
Jugendlicher Pfropfstamm
Junger verpfropfter Unterlagenstamm
Juvenile grafted rootstock trunk
Sich etablierender Wurzelstockstamm
Young Grafted Rootstock Trunk
Syrah young grafted scion trunk on 1103P vigorous rootstock at 3 years post-planting shows 32-42 mm scion diameter with 3 annual rings averaging 5.0-7.0 mm width, graft union at 10 cm above soil with stable 112-118% diameter swelling, rootstock diameter 38-48 mm below union, 50-75 g combined starch reserves (scion 18-28 g, rootstock 32-47 g), developing bilateral cordons at 65 cm height, and rootstock-enhanced vigor supporting year 3 light crop of 12-18 clusters
A young grafted scion trunk is a grafted scion trunk in early developmental phases (typically years 1-4 post-planting) characterized by rapid diameter increase, establishment of training architecture, graft union maturation, and transition from vulnerable establishment to approaching commercial productivity, representing the critical period when graft union stability and rootstock-scion compatibility are confirmed. Young grafted scion trunk development proceeds through characteristic phases= year 1 with graft union stabilization and initial scion shoot growth (scion trunk diameter 5-15 mm by dormancy, rootstock already established from nursery), year 2 with accelerated scion growth and architecture initiation (diameter 12-28 mm, rootstock-scion diameter ratio stabilizing), year 3 with continued rapid growth and beginning productivity (diameter 22-45 mm, approaching commercial architecture), and year 4 with approaching mature status (diameter 35-65 mm depending on rootstock vigor influence). Young grafted scion trunk anatomical characteristics include= active vascular cambium with wide annual growth rings (3-8 mm ring width in scion compared to 1-3 mm in mature grafted trunks), scion wood anatomy reflecting cultivar characteristics (ring-porous pattern, vessel diameter, parenchyma distribution specific to variety), graft union with maturing vascular connections (progressive xylem and phloem integration across graft interface), relatively thin scion bark (1-3 mm compared to 4-10 mm in mature trunks), and minimal heartwood formation (scion wood entirely functional sapwood conducting water). Young grafted scion trunk graft union maturation includes= progressive union swelling stabilization (diameter ratio between union and adjacent trunk reaching equilibrium typically 105-120%), vascular connection strengthening (increasing xylem and phloem continuity improving hydraulic conductance and nutrient transport across union), callus tissue lignification (initial soft callus becoming woody reducing breakage risk), union bark development (periderm formation protecting graft interface), and compatibility confirmation or incompatibility manifestation (successful combinations showing stable growth, incompatible combinations showing excessive swelling >130%, necrosis, or growth suppression). Young grafted scion trunk physiological characteristics include= rootstock vigor expression (scion growth rate, shoot length, leaf size reflecting rootstock influence), high growth rates with strong apical dominance (vigorous vegetative development establishing canopy architecture), rootstock-mediated water relations (rootstock characteristics affecting scion water status, drought response, hydraulic conductance), rootstock nutrient uptake patterns (rootstock genetics influencing scion mineral nutrition, pH tolerance, nutrient efficiency), and establishment of carbohydrate partitioning patterns (reserves accumulating in both scion trunk and rootstock, typically 15-25 g in scion trunk and 25-45 g in rootstock by year 3-4). Young grafted scion trunk management considerations include= training to desired architecture (cordon development, head training similar to own-rooted but accounting for union position), rootstock sucker removal (vigilant removal of shoots emerging from rootstock below union preventing rootstock canopy development), graft union protection (avoiding mechanical damage, freeze injury, pathogen infection at vulnerable union), crop load progression (typically no crop year 1, light crop 15-25 clusters years 2-3, approaching full crop year 4), and early incompatibility detection (monitoring for excessive union swelling, poor growth, leaf symptoms indicating incompatibility requiring replanting). Young grafted scion trunk rootstock influence on development includes= vigorous rootstocks (SO4, 110R, 1103P) accelerating scion establishment (reaching 45-70 mm diameter by year 4, enabling earlier commercial production), moderate rootstocks (5BB, 420A, 101-14Mgt) producing balanced development (35-55 mm diameter by year 4, standard establishment timeline), low-vigor rootstocks (Riparia Gloire, 3309C) slowing scion growth (30-50 mm diameter by year 4, potentially delaying full productivity to year 5-6). Young grafted scion trunk establishment challenges include= graft union failure (5-15% loss in first 2 years from incompatibility or poor grafting quality), union breakage from mechanical stress (wind, equipment contact particularly vulnerable during years 1-2), rootstock suckering competition (vigorous rootstock shoots diverting resources from scion if not removed), delayed productivity compared to own-rooted in some cases (graft union establishment potentially slowing initial growth), and environmental stress sensitivity (union as physiological bottleneck potentially increasing drought, heat, or cold sensitivity during establishment).
Young_Grafted_Scion_Trunk emphasizes early establishment with graft union and is distinguished from Young_Own_Rooted_Trunk (no union, different establishment dynamics) and Mature_Grafted_Scion_Trunk (established productive phase) by graft union maturation process and rootstock-influenced development. Critical for understanding grafted vine establishment timeline, union stability confirmation, rootstock vigor effects on early growth, and management specific to grafted vine establishment.
Establishing grafted trunk
Etablierender Veredelungsstamm
Junger gepfropfter Edelstamm
Juvenile grafted scion
Juveniler Aufpropfungsstamm
Young Grafted Scion Trunk
A young leaf at node 11 on an actively extending shoot of Vitis vinifera cv. Pinot Noir in Burgundy, France, measuring 6.5 cm length by 5.8 cm width representing approximately 45% of expected mature dimensions, exhibiting pale green coloration with bronze tinge from anthocyanin accumulation particularly along veins and margins, soft fragile texture with thin cuticle lacking glossy appearance of mature leaves, incompletely differentiated mesophyll showing single developing palisade layer of 35 μm and loosely organized spongy mesophyll, chlorophyll content of 180 μg cm⁻² (compared to 450 μg cm⁻² in mature leaves on same shoot), demonstrating photosynthesis rate of 5.2 μmol CO₂ m⁻² s⁻¹ representing 42% of mature leaf capacity, positioned 12 cm behind shoot apex in zone of active cell expansion, occurring 15 days post-emergence from apical meristem, measured at shoot BBCH stage 71 (fruit set) in early June though this distal leaf is still in early development, representing typical young leaf characteristics during mid-expansion phase requiring continued resource investment for approximately 2-3 additional weeks before achieving mature functional status.
A leaf in early developmental stages from initial primordium formation through expansion phases before achieving full maturity in Vitis vinifera, characterized by incomplete structural development with ongoing cell division and expansion processes, dimensions substantially below cultivar-typical mature size typically 20-80% of final dimensions depending on developmental stage, immature tissue organization with incompletely differentiated mesophyll layers, developing vascular systems showing incomplete secondary development, thin immature cuticle with partial or absent wax deposition, pale green to reddish coloration reflecting low chlorophyll concentrations and often high anthocyanin content providing photoprotection during vulnerable developmental stages, soft fragile texture from thin cell walls and high water content, and suboptimal photosynthetic capacity typically 30-70% of mature leaf rates reflecting incomplete chloroplast development and lower photosynthetic enzyme concentrations. Young leaves emerge continuously from shoot apical meristems throughout the growing season as shoots extend, with new leaf primordia initiated at approximately 2-4 day intervals creating the characteristic phyllotactic pattern, followed by 3-5 weeks of development through cell division, expansion, and differentiation phases before achieving mature status, consuming substantial resources particularly nitrogen for protein synthesis and carbohydrates for cell wall construction while contributing limited photosynthetic return until later developmental stages. The young leaf developmental period represents a critical vulnerability window with high susceptibility to environmental stresses including frost damage, heat stress, water deficit, pest attack particularly by mites and leafhoppers, and fungal diseases including downy and powdery mildew that preferentially infect young tender tissues.
Young leaves are distinguished from mature leaves by incomplete development across multiple parameters including size, tissue differentiation, physiological function, and structural characteristics, and from senescent leaves by active developmental processes rather than degradation. Classification as young requires assessment of multiple developmental indicators, as size alone is insufficient given variation in mature leaf dimensions. Young leaves pass through sequential developmental stages from emergence through expansion to maturation, with boundaries between stages somewhat arbitrary but useful for developmental characterization. Environmental conditions during young leaf development critically determine mature leaf characteristics through phenotypic plasticity, with light, temperature, water, and nutrient availability during the 3-5 week development period establishing sun versus shade leaf anatomy, final size, and functional capacity. Young leaves represent major resource sinks consuming carbohydrates and nitrogen while contributing limited photosynthetic return, creating temporary negative carbon balance at shoot level that must be supported by mature leaves or reserves until young leaves achieve net productivity.
developing leaf
immature leaf
junges Blatt
juvenile leaf
juveniles Blatt
sich entwickelndes Weinblatt
Based on Mullins et al. (1992) "Biology of the Grapevine" Cambridge University Press pp. 60-92 Palliotti & Cartechini (2000) "Developmental changes in gas exchange activity in flowers, berries, and tendrils of field-grown Cabernet Sauvignon" American Journal of Enology and Viticulture 51=317-323 Poni et al. (2009) "Effects of early defoliation on shoot photosynthesis, yield components, and grape composition" American Journal of Enology and Viticulture 60=397-407 Keller (2020) "The Science of Grapevines= Anatomy and Physiology" 3rd Edition, Academic Press pp. 56-94.
Young Leaf
The young leaf color of Sangiovese recorded at the third and fourth leaf position is OIV 051 state 3 (bronze-orange) — the adaxial surface displays a warm bronze-copper tone resulting from the combination of emerging chlorophyll (green) with substantial anthocyanin and flavonoid accumulation (red-orange), a pigmentation balance that persists for 10-14 days as the leaf expands before chlorophyll production dominates and the mature green color is established. This bronze-orange young leaf color distinguishes Sangiovese from the green-yellow young leaves of Trebbiano Toscano (OIV 051:1, minimal non-chlorophyll pigmentation), the strongly copper-red young leaves of Aglianico (OIV 051:4, intense anthocyanin expression), and the crimson young leaves of Tannat (OIV 051:5, maximum anthocyanin expression in young leaf tissue).
A young leaf description component that records the ground color and overlying pigmentation of the third and fourth unfolded leaves, scored according to OIV descriptor 051 (color of upper surface of the blade) on a scale from green-yellow (state 1) through yellow (state 2), bronze-orange (state 3), copper-red (state 4) to crimson-red (state 5), reflecting the balance between chlorophyll accumulation and anthocyanin/flavonoid pigmentation in expanding leaf tissue that is determined by cultivar genotype and modulated by light exposure.
Young_Leaf_Color documents a character that bridges the shoot tip anthocyanin assessment (OIV 003) and the mature leaf coloration, providing cultivar identification information during the developmental window when shoot tip characters are fading but mature leaf morphology is not yet fully expressed. The color of young leaves reflects the complex developmental regulation of pigment biosynthesis during leaf expansion — newly unfolded leaves must balance the need for photoprotection (provided by anthocyanins and UV-absorbing flavonoids that shield developing chloroplasts from excess light) against the need for photosynthetic capacity (requiring chlorophyll accumulation and chloroplast maturation): cultivars differ genetically in the intensity and duration of this photoprotective pigmentation phase, producing the range of young leaf colors from green-yellow (rapid chlorophyll accumulation, minimal photoprotective pigments) through bronze (balanced pigment production) to crimson-red (strong anthocyanin expression persisting well into the leaf expansion period). The bronze coloration is particularly diagnostic because it represents a specific pigment mixture that creates a visually distinctive appearance not achievable by any single pigment class.
Entwicklungsblatt-Pigmentierung
Jungblatt-Farbe
developing leaf color assessment
junge Blattoberseiten-Färbung
third-fourth leaf coloration
young leaf pigmentation record
Young Leaf Color
The young leaf description of Tempranillo recorded at E-L stage 12 (6-8 leaves unfolded) documents the third and fourth leaves as having a yellow-green ground color with strong bronze-red overtones on the adaxial surface (OIV 051:4), medium density of prostrate hairs on the abaxial surface between main veins (OIV 053:5), sparse erect hairs on the abaxial surface between main veins (OIV 054:3), and a distinctly bullate (blistered) surface texture already visible at this developmental stage — this combination of strong bronze coloration with moderate pubescence is characteristic of Tempranillo and distinguishes it from the similarly bronze-tinted but more densely pubescent Grenache (OIV 053:7) and the green-yellow, minimally bronze Graciano (OIV 051:2) during the spring observation period.
An ampelographic description component that documents the morphological characteristics of the third and fourth unfolded leaves counted from the shoot tip — the standardized observation position for young leaf descriptors in the OIV system — recording color, hairiness, and distinctive pigmentation features during the active growth phase (E-L stages 9-15) that provide cultivar-diagnostic characters complementing the shoot tip observations and bridging the developmental transition from juvenile to mature leaf morphology.
Young_Leaf_Description captures the developmental stage intermediate between the shoot tip (OIV 001-004) and the mature leaf (OIV 065-094), documenting characters of the third and fourth leaves that are still expanding but have progressed beyond the primordial stage. This observation position was selected by the OIV because the third and fourth leaves represent a reproducible developmental stage — old enough to display cultivar-specific color and hairiness patterns clearly, but young enough that environmental modification of these characters remains minimal compared to fully mature leaves: the standardization of observation position (specifically the third and fourth leaves, not the second or fifth) is critical for comparability because young leaf characters change rapidly along the developmental gradient from tip to base, with each successive leaf differing measurably in pigmentation intensity, trichome density, and surface texture. The young leaf descriptors (OIV 051-056) provide information not captured by the shoot tip descriptors — particularly the bronze coloration phenotype and the surface hairiness patterns that become visible only as the leaf blade expands sufficiently to display its trichome architecture.
Jungblatt-Beschreibung
developing leaf characterization
junge Laubblatt-Morphologie
sich entwickelndes Blatt-Charakterisierung
third-fourth leaf description
young foliage morphology
Young Leaf Description
The young leaf hairiness of Viognier recorded at the third and fourth leaf position documents very dense prostrate hairs on the abaxial surface between main veins (OIV 053:9) combined with sparse erect hairs (OIV 054:3) — the dense prostrate trichome mat creates a white-felted lower leaf surface visible to the naked eye, a character so pronounced in Viognier that it remains clearly evident even on mature leaves (unusual for most cultivars where young leaf pubescence diminishes substantially during maturation). This trichome combination — very dense prostrate with sparse erect — distinguishes Viognier from Roussanne (dense prostrate with medium erect, OIV 053:7/054:5) and Marsanne (medium prostrate with sparse erect, OIV 053:5/054:3) among the major white Rhône cultivars.
A young leaf description component that records the type and density of trichomes on the abaxial (lower) surface of the third and fourth unfolded leaves, distinguishing between prostrate hairs (OIV 053, woolly/appressed trichomes forming a mat-like covering) and erect hairs (OIV 054, upright bristle-like trichomes) scored independently on 1-9 density scales, providing trichome architecture data at the young leaf developmental stage that complements the shoot tip hairiness assessment and predicts the mature leaf pubescence pattern.
Young_Leaf_Hairiness captures the trichome phenotype at a developmental stage when pubescence is typically at or near its maximum expression — young expanding leaves generally carry denser trichome coverage than mature leaves because the trichomes are formed during early leaf development and become progressively dispersed as the leaf blade expands, reducing the apparent density per unit area. Scoring hairiness at the standardized third-fourth leaf position therefore provides the highest-contrast observations for cultivar discrimination: the independent scoring of prostrate and erect trichome types on separate scales is essential because these trichome types are developmentally distinct (originating from different epidermal cell populations), genetically independently controlled, and diagnostically complementary — some cultivars (e.g., Merlot) show high prostrate but low erect density, others (e.g., Sangiovese) show moderate prostrate with medium erect, and still others (e.g., Chasselas) show low density of both types, creating a two-dimensional trichome character space that provides substantially greater cultivar discrimination than either dimension alone.
Entwicklungsblatt-Pubeszenz
Jungblatt-Behaarung
developing leaf trichome assessment
junge Blattunterseiten-Trichome
third-fourth leaf hairiness
young leaf pubescence record
Young Leaf Hairiness
Syrah young own-rooted trunk at 3 years post-planting in California vineyard shows 28-38 mm diameter with 3 annual growth rings averaging 4.5-6.5 mm width, active cambium producing vigorous secondary growth, 2-3 mm bark thickness, 45-65 g stored starch reserves, and developing cordon arms at 60 cm trunk height supporting light crop of 2-4 clusters per shoot
A young own-rooted trunk is an own-rooted trunk in early developmental phases (typically years 1-4 post-planting) characterized by rapid diameter increase, establishment of secondary growth, development of training architecture, and transition from juvenile to productive maturity, representing the critical establishment period before regular commercial fruit production begins. Young own-rooted trunk development proceeds through characteristic phases= year 1 with initial root establishment and modest shoot growth (trunk diameter 5-15 mm by dormancy), year 2 with accelerated vegetative growth and trunk thickening (diameter 12-25 mm), year 3 with continued rapid growth and beginning fruit production (diameter 20-40 mm), and year 4 with approaching mature architecture and increasing productivity (diameter 30-60 mm depending on cultivar vigor). Young own-rooted trunk anatomical characteristics include= active vascular cambium with wide annual growth rings (3-8 mm ring width compared to 1-3 mm in mature trunks), high proportion of earlywood with large vessels (supporting rapid expansion and high water transport capacity), relatively thin bark (1-3 mm compared to 4-10 mm in mature trunks), minimal heartwood formation (most wood remains sapwood conducting water), and absence of significant trunk pathogen colonization (young healthy wood without decay or disease symptoms). Young own-rooted trunk physiological characteristics include= high growth rates with strong apical dominance, vigorous vegetative development with large leaf area production, high photosynthetic capacity supporting rapid biomass accumulation (net photosynthesis 12-20 μmol CO₂/m²/s in optimal conditions), high carbohydrate partitioning to vegetative growth rather than fruit (fruit production deliberately limited or prevented in years 1-2), and establishment of perennial carbohydrate reserves (starch accumulation in trunk wood parenchyma increasing from 10-20 g/vine in year 1 to 40-80 g/vine by year 4). Young own-rooted trunk management considerations include= training to desired architecture (vertical shoot positioning, cordon development, head training), selective shoot retention and positioning, fruit load management (typically no crop year 1, light crop years 2-3, approaching full crop year 4), irrigation to support establishment (particularly critical in first growing season), protection from environmental stress (frost, drought, excessive heat during vulnerable establishment period), and pest and disease management (young vigorous growth attractive to certain pests, disease prevention establishing healthy trunk foundation). Young own-rooted trunk cultivation targets include= achieving target trunk diameter for training system (30-50 mm for most systems by year 3-4), establishing permanent cordon or head positions, developing adequate carbohydrate reserves to support cropping (50-80 g starch per vine by year 3-4), creating strong trunk architecture resistant to wind damage or mechanical stress, and avoiding trunk diseases during establishment (preventing Eutypa or Esca infection of pruning wounds). Young own-rooted trunk variation by cultivar includes= high-vigor varieties (Grenache, Mourvèdre) showing faster trunk development reaching 40-60 mm diameter by year 3, moderate-vigor varieties (Cabernet Sauvignon, Merlot) reaching 25-40 mm by year 3, low-vigor varieties (Pinot Noir, Chardonnay) reaching 20-35 mm by year 3, and environmental effects (fertile soils, adequate water promoting faster development than stressed conditions).
Young_Own_Rooted_Trunk emphasizes early establishment phase and is distinguished from Mature_Own_Rooted_Trunk (established productive phase) and Old_Own_Rooted_Trunk (declining phase) by rapid growth, architecture development, and transition to productivity. Critical for understanding vineyard establishment timeline and costs, early training and pruning decisions, optimal crop load progression, and foundation for long-term vine health and productivity.
Establishing trunk
Etablierender Eigenwurzelstamm
Junger wurzelechter Stamm
Juvenile own-rooted trunk
Juveniler ungepfropfter Stamm
Young Own Rooted Trunk
The young shoot description of Nebbiolo recorded at E-L stage 9-12 (4-6 leaves unfolded) documents green internodes with distinct red-violet anthocyanin stripes on the dorsal (sun-exposed) surface (OIV 006:3), nodes with intensified anthocyanin accumulation forming a conspicuous ring (OIV 007:5), a semi-erect to horizontal growth habit, and the characteristic tendril pattern of two consecutive tendrils followed by one tendril-free node (2-0-2-0 pattern typical of V. vinifera) — the combination of striped anthocyanin pattern on green ground color with strongly pigmented nodes is highly distinctive for Nebbiolo and separates it from other Piedmontese cultivars including Barbera (uniformly green internodes, OIV 006:1) and Dolcetto (diffuse anthocyanin without distinct striping, OIV 006:2) during the spring observation period.
A shoot tip description component that documents the morphological characteristics of the young shoot from the first to the fourth or fifth unfolded leaf, recording internode color, node pigmentation, tendril distribution pattern, and the developmental gradient from the shoot tip through the expanding leaves that provides cultivar-diagnostic features during the active growth phase from budburst through early shoot elongation (E-L stages 4-12).
Young_Shoot_Description captures the morphological features of the elongating shoot that complement the shoot tip characters observed earlier in the season, extending the ampelographic observation window from budburst through the first 6-8 weeks of growth. The young shoot provides several diagnostically important characters not available from the shoot tip alone — particularly internode color and anthocyanin distribution pattern (OIV 006, scored as green, green with red stripes, or red, providing a three-category character with high discrimination), and node pigmentation intensity (OIV 007, varying from absent through moderate to very strong): these characters reflect the interaction between constitutive anthocyanin biosynthesis capacity (genetically determined) and light exposure (environmentally influenced), with the anthocyanin distribution pattern being more genetically stable than the total intensity. The tendril distribution pattern — consistently two consecutive tendrils alternating with one tendril-free node in V. vinifera versus continuous tendrils in some wild Vitis species — serves more as a species-level diagnostic than a cultivar-level character, but variations in tendril size and branching can contribute to cultivar distinction within V. vinifera.
Jungtrieb-Beschreibung
developing shoot characterization
early shoot morphology
junge Internodien-Beschreibung
sich entwickelnder Trieb-Charakterisierung
young internode description
Young Shoot Description
A 2-year-old Pinot Noir vine bears a Young_Vine_Role with first fruiting shoots emerging but crop removed entirely to direct energy toward root and trunk development—the training stakes and grow tubes still in place guiding establishment. A 4-year-old Cabernet Sauvignon vine serves in a Young_Vine_Role with first partial crop allowed (50% of mature target), bilateral cordons now established but spur positions still developing toward the full complement expected at maturity. A replanted vine in an established vineyard block bears a Young_Vine_Role requiring individual attention (irrigation, pest protection, training) distinct from the mature management regime applied to surrounding vines.
A vine age role that inheres in a grapevine during its early post-planting years (typically years 1-5), characterized by root system development, canopy establishment, training system implementation, and gradual transition toward full production. Young vines prioritize vegetative development over fruit production, with crop load typically restricted during the first 2-3 years to channel resources toward root and permanent structure establishment. The young vine phase involves active training—trunk development, cordon or head formation, establishment of spur positions—that shapes the vine architecture for its productive lifespan. Young vine management balances the desire for early fruit production (accelerating return on vineyard investment) against the need for adequate establishment (ensuring long-term vine health and productivity). Root system development during the young vine phase is particularly critical, as the root architecture established in early years influences water and nutrient acquisition capacity throughout the vine's life. Young vine fruit, when produced, often shows character differences from mature vine fruit—higher acidity, less concentrated flavors, less complex wines—reflecting the vine's immature physiology and limited root system.
Young_Vine_Role captures the establishment phase requiring intensive management investment to create productive vine architecture. The transition from young to established status varies with variety vigor, site conditions, and management intensity fast-establishing combinations may reach productive maturity by year 4-5 while challenging situations may extend the young vine phase to year 6-8. Young vine identification in mixed-age blocks is important for appropriate management—young vines may require different irrigation, nutrition, and pest management than mature neighbors. The economics of vineyard establishment depend heavily on the young vine period duration and productivity techniques accelerating establishment (potted vines, aggressive early training, optimized nutrition) can improve investment returns by shortening the period before full production. Young vine fruit, when included in production, is often segregated or blended rather than used for premium bottlings.
Junganlagen-Rebe
Jungrebe
establishing vine
junge Rebe
juvenile vine
neu gepflanzte Rebe
newly planted vine
young vine
Keller M (2015) The Science of Grapevines, Academic Press Smart RE, Robinson M (1991) Sunlight into Wine, Winetitles Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press
Young Vine Role
Young's modulus measurement of Riesling rachis tissue using uniaxial tension testing reveals values of 45±8 MPa in green rachis at véraison declining to 28±6 MPa at harvest maturity, with the 38% modulus reduction correlating with increased rachis flexibility that reduces mechanical damage during harvest handling but also increases susceptibility to bunch stem necrosis when lignification fails to progress normally, illustrating the balance between mechanical properties and physiological function.
Young's modulus quality is an elastic modulus quality that represents the uniaxial stiffness of tissue defined as the ratio of tensile or compressive stress to longitudinal strain in one-dimensional loading, characterizing the resistance to stretching or compression along a single axis that is fundamental to mechanical testing and engineering analysis of biological materials. Within the BFO framework, this quality specifically characterizes the elastic property measured in standard uniaxial mechanical testing geometry, distinguishing from bulk modulus measured under volumetric loading and providing the mechanical parameter most directly comparable to materials science and engineering data. The ontological significance lies in Young's modulus representing the classical definition of elastic modulus (E = σ/ε) that enables prediction of tissue deformation under uniaxial loads, calculation of stresses from observed strains, and comparison of plant tissue mechanical properties with synthetic materials and other biological tissues. Young's modulus in grape berry skin typically ranges from 1-10 MPa depending on ripeness and measurement direction, while woody stem tissues show values of 100-1000 MPa reflecting the stiffening effect of lignification, demonstrating the wide range of mechanical properties achieved through differential cell wall composition and tissue architecture. The anisotropic structure of plant tissues means that Young's modulus often differs depending on the direction of loading relative to tissue organization, with longitudinal modulus (parallel to vascular elements or cellulose microfibrils) typically exceeding transverse modulus in organized tissues. Measurement employs uniaxial tensile or compression testing with force and displacement sensors, calculating stress from force and cross-sectional area and strain from displacement relative to initial gauge length. The strain rate dependence of viscoelastic plant tissues means that measured Young's modulus varies with testing speed, requiring standardization for meaningful comparison across studies. Understanding Young's modulus quality enables engineering analysis of tissue mechanics, prediction of failure under specified loading, and comparison of plant tissue mechanical properties with design requirements for applications ranging from harvest machinery to food texture engineering.
Youngs_Modulus_Quality provides the uniaxial elastic modulus from standard mechanical testing, distinguishing from Bulk_Elastic_Modulus_Quality derived from pressure-volume analysis. This quality is direction-dependent in anisotropic tissues and strain-rate-dependent in viscoelastic tissues, requiring specification of testing conditions for meaningful comparison. The class enables interface with engineering analysis methods and materials science literature using standardized mechanical parameters.
Dehnungsmodul
Young-Modul
Zugmodul
longitudinal elastic modulus
tensile modulus
uniaxial elastic modulus
Niklas KJ (1992) Plant Biomechanics: An Engineering Approach to Plant Form and Function. University of Chicago Press, Chicago, pp. 123-178 Burgert I (2006) Exploring the micromechanical design of plant cell walls. American Journal of Botany 93(10): 1391-1401
Youngs Modulus Quality
Zinc content quality management in South Australian Shiraz addresses the widespread zinc deficiency affecting sandy, low-pH soils of the Riverland and Barossa regions, with foliar zinc sulfate applications (2-3 kg/ha ZnSO₄) at flowering correcting the "little leaf" and poor fruit set that otherwise limit yield and quality adequate zinc nutrition (must concentration 1-3 mg/L) supports the alcohol dehydrogenase activity essential for clean fermentation of high-Baumé Shiraz, preventing the sluggish fermentations and residual sugar problems that can affect zinc-deficient fruit.
Zinc_Content_Quality represents the chemical quality characterizing the concentration of zinc (Zn²⁺) in grape berry tissue, capturing the essential micronutrient central to enzyme activation, protein synthesis, and hormone regulation in grapevines. Zinc in grape juice typically ranges from 0.5-5 mg/L, with variation reflecting cultivar, soil zinc availability, and the soil chemistry factors affecting zinc uptake. As a cofactor for over 300 enzymes including carbonic anhydrase, alcohol dehydrogenase, and superoxide dismutase, zinc participates in carbon dioxide fixation, fermentation biochemistry, and antioxidant defense. Zinc is also essential for auxin synthesis, making it critical for cell elongation and fruit development. Zinc deficiency produces distinctive symptoms in grapevines including stunted shoots with shortened internodes, small leaves with distorted margins (little leaf), and poor fruit set with straggly, shot-berry clusters. Soil zinc availability decreases with increasing pH, creating deficiency risk in calcareous soils, while zinc binds strongly to organic matter, potentially limiting availability in high-organic soils despite adequate total zinc content. Sandy soils with low cation exchange capacity are prone to zinc leaching and deficiency. In winemaking, zinc affects fermentation through its role in alcohol dehydrogenase and other yeast enzymes, with adequate zinc nutrition supporting healthy fermentation while zinc-deficient musts may show sluggish fermentation. Wine zinc content is typically lower than must due to precipitation and yeast uptake, with normal levels (less than 5 mg/L) not causing quality concerns. Zinc analysis employs atomic absorption spectroscopy or ICP-OES with appropriate sample preparation to prevent contamination from zinc-containing laboratory materials.
Zinc_Content_Quality addresses a micronutrient with distinctive deficiency symptoms (little leaf, poor fruit set) that enable visual diagnosis. The enzyme cofactor role of zinc spans both plant metabolism and fermentation biochemistry, linking vineyard zinc status to winemaking outcomes. Zinc availability in soil depends on pH, organic matter, and phosphorus status, with high phosphorus potentially inducing zinc deficiency through antagonistic interactions. Contamination risk during sampling and analysis requires careful protocol to prevent zinc introduction from galvanized materials.
Beerenzinkqualität
Zinkgehalt
Zinkkonzentration
Zn content
Zn-Gehalt
berry zinc quality
zinc concentration
zinc level
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Ed, Academic Press pp.: 341-385 Christensen P (2005) Foliar Fertilization in Vine Mineral Nutrient Management Programs. In: Christensen LP and Smart DR (eds) Proceedings of the Soil Environment and Vine Mineral Nutrition Symposium, American Society for Enology and Viticulture pp.: 83-90
Zinc Content Quality
Zinc deficiency symptom quality diagnosis in San Joaquin Valley Thompson Seedless identified Zn limitation as the cause of poor fruit set and characteristic foliar symptoms on a high-pH soil with history of heavy phosphorus fertilization. Visual zinc deficiency symptom quality assessment revealed the pathognomonic little leaf syndrome with affected shoots showing abnormally small leaves (blade area 25-40 cm² versus normal 100-120 cm²), shortened petioles, and asymmetric blade development with unequal lobes. Additional zinc deficiency symptom quality features included shortened internodes creating bunchy shoot tips, interveinal mottling with irregular light green to yellow patches between major veins, and wide spacing between marginal serrations giving leaves a "sawtooth" appearance distinct from the fine serrations of normal leaves. Fruit assessment revealed zinc deficiency symptom quality manifesting as millerandage with 35-45% of berries in affected clusters remaining as small seedless shot berries while others developed normally, creating the "hens and chickens" cluster appearance that reduced both yield and table grape marketability. Tissue analysis confirmed zinc deficiency symptom quality diagnosis with young leaf Zn at 11 ppm (severely deficient, threshold 20 ppm) and bloom petiole Zn at 14 ppm (deficient threshold 26 ppm), with soil Zn at only 0.4 ppm DTPA-extractable despite adequate total Zn indicating pH-limited availability. The P:Zn ratio in tissue was elevated at 450 versus optimal below 200, indicating P-Zn antagonism from historical high P fertilization contributing to zinc deficiency symptom quality. Corrective foliar ZnSO₄ application at 0.5% concentration (three applications from budbreak through bloom) resolved symptoms within one season and improved fruit set to normal levels the following year.
Zinc deficiency symptom quality is a micronutrient deficiency symptom quality that inheres in a grapevine organism and represents the characteristic visual symptoms resulting from inadequate zinc availability for its numerous enzymatic and regulatory functions, including the pathognomonic "little leaf" symptom where young leaves emerge abnormally small with shortened petioles, asymmetric blade development, and widely spaced serrations, along with shortened internodes creating a rosette or bunchy growth habit, poor fruit set leading to millerandage (shot berries), and interveinal mottling or chlorosis on affected leaves. This quality captures the grapevine response to zinc limitation, recognizing that zinc deficiency produces among the most distinctive and readily identifiable nutrient deficiency symptoms in viticulture, with the little leaf syndrome virtually pathognomonic for zinc limitation. The ontological significance of zinc deficiency symptom quality lies in both its diagnostic distinctiveness enabling confident field identification and its significant impact on fruit production through the millerandage that can reduce yields substantially while also affecting cluster appearance and uniformity important for table grape production. The biological basis of zinc deficiency symptoms encompasses the essential role of zinc in carbonic anhydrase affecting CO₂ fixation, in RNA polymerase and numerous transcription factors affecting gene expression, in auxin metabolism where zinc is required for tryptophan synthesis as the auxin precursor, and in membrane integrity through zinc's role in protecting sulfhydryl groups from oxidation. The little leaf symptom reflects disrupted auxin metabolism, with inadequate auxin production limiting cell expansion during leaf development, while the millerandage results from impaired fertilization and fruit set when zinc deficiency affects pollen development and pollen tube growth. The soil conditions predisposing to zinc deficiency include high pH (particularly above 7.5) reducing zinc solubility, high phosphorus availability that can induce zinc deficiency through P-Zn antagonism, and sandy low-organic-matter soils with limited zinc retention, making zinc deficiency common on calcareous soils where it often co-occurs with iron deficiency.
Zinc_Deficiency_Symptom_Quality is_a Micronutrient_Deficiency_Symptom_Quality, serving as parent class for Little_Leaf_Quality and Millerandage_Induction_Quality_(Zn) which address specific zinc deficiency manifestations. This class addresses the overall zinc deficiency syndrome while child classes capture particular symptom types. "Little leaf" is virtually pathognomonic for Zn deficiency. Millerandage (shot berries) causes significant yield and quality impact. Immobile nutrient showing symptoms on youngest tissues. High pH and P-Zn antagonism are primary predisposing factors. Petiole Zn threshold ~26 ppm at bloom young leaf ~20 ppm.
Niedrig-Zink-Symptome
Zinkmangel-Symptome
Zinkstress-Indikatoren
Zn deficiency symptoms
Zn-Mangel-Zeichen
low zinc symptoms
zinc limitation signs
zinc stress indicators
Keller M (2020) The Science of Grapevines: Anatomy and Physiology, 3rd Edition. Academic Press, pp. 297-345 Christensen P (2000) Nutrient deficiency symptoms in grapevines. University of California ANR Publication Swietlik D (1999) Zinc nutrition in horticultural crops. Horticultural Reviews 23:109-178
Zinc Deficiency Symptom Quality
Zinc uptake efficiency varies 2-3 fold among rootstocks, with 1103 Paulsen and 110 Richter showing high efficiency (maintaining petiole Zn above 30 ppm) compared to 101-14 Mgt and SO4 (petiole Zn 15-25 ppm) under identical low-Zn soil conditions. Zinc deficiency symptoms include shortened internodes (little leaf), mottled interveinal chlorosis, and poor fruit set with millerandage, appearing when petiole Zn falls below 15-20 ppm at bloom. VvZIP family members encode zinc transporters with root expression increasing 3-5 fold under Zn limitation. Zinc availability declines sharply above soil pH 7.0, with uptake reduced by 50-70% between pH 6.5 and 8.0 under equivalent total soil Zn.
A micronutrient uptake disposition that is the inherited capacity of a Vitis vinifera plant to absorb zinc ions (Zn²⁺) from the soil solution through ZIP family transporters, with genotype-specific efficiency affecting zinc-dependent enzyme function, auxin metabolism, and reproductive development.
Zinc deficiency is among the most common micronutrient disorders in viticulture, affecting both vegetative growth and reproductive success. The characteristic little leaf symptom results from impaired auxin synthesis (Zn is required for tryptophan-dependent IAA biosynthesis). Zinc uptake efficiency is particularly important on high-pH soils and in high-phosphorus conditions that inhibit Zn absorption. Foliar Zn application before bloom effectively supplements root uptake for reproductive development.
Zinkaneignungsvermögen
Zinkaufnahme
Zn uptake capacity
Zn-Akquisition
zinc acquisition
zinc foraging trait
Christensen LP (1980) Timing of zinc foliar sprays: II. Effects on size, sweetness, and bunch rot of Thompson Seedless grapes. American Journal of Enology and Viticulture 31:256-260 Sinclair AH, Edwards AC (2008) Micronutrient deficiency problems in agricultural crops in Europe. In: Alloway BJ (ed) Micronutrient Deficiencies in Global Crop Production, Springer pp.: 225-244
Zinc Uptake Disposition
Zinc uptake efficiency assessment in vineyard with marginal soil zinc (DTPA-Zn 0.6 mg/kg) reveals the factors affecting acquisition, with mycorrhizal vines showing adequate leaf Zn of 28 mg/kg (fungal hyphae extending zinc acquisition zone), non-mycorrhizal vines in fumigated soil showing deficient Zn of 14 mg/kg (limited to direct root uptake), and high-phosphorus treatment further reducing leaf Zn to 11 mg/kg (P-Zn antagonism), demonstrating how zinc uptake efficiency depends on both root transport capacity and the biotic and chemical soil environment affecting zinc availability at root surfaces.
Zinc uptake efficiency quality is a micronutrient uptake efficiency quality that represents the capacity of grapevine roots to acquire zinc from soil, characterizing the effectiveness of Zn²⁺ absorption that determines vine zinc status affecting enzyme function, auxin metabolism, protein synthesis, and membrane integrity with implications for fruit set, berry development, and wine quality. Within the BFO framework, this quality specifically addresses the zinc component of micronutrient uptake, distinguishing zinc acquisition from other trace elements while recognizing that zinc deficiency causes distinctive symptoms including small leaves with shortened internodes, poor fruit set, and reduced berry size that can significantly impact yield and quality. The ontological significance lies in zinc uptake efficiency determining access to this essential cofactor for over 300 enzymes including carbonic anhydrase, alcohol dehydrogenase, and RNA polymerase, where adequate zinc is required for auxin synthesis (tryptophan pathway), protein synthesis, and membrane stability, with deficiency causing the characteristic "little leaf" symptom from impaired auxin function and reduced fruit set from pollen tube abnormalities. The mechanistic basis for zinc uptake involves ZIP (Zinc-Iron Permease) family transporters that mediate Zn²⁺ absorption from soil solution, with high-affinity uptake systems operating at the low Zn²⁺ concentrations typical of deficient soils, and with transporter expression upregulated under zinc deficiency to maximize acquisition capacity. The soil factors affecting zinc availability include pH (availability declining sharply above pH 6.5 from precipitation and adsorption), phosphorus status (high P can induce zinc deficiency through antagonistic interactions), organic matter (both chelating Zn and providing mineralizable zinc), and soil texture (sandy soils more prone to zinc deficiency from leaching), creating site-specific zinc availability patterns. The mycorrhizal contribution to zinc uptake can be substantial, as arbuscular mycorrhizal hyphae access soil volume beyond the zinc depletion zone around roots, with mycorrhizal vines often showing improved zinc nutrition particularly in low-zinc soils. Understanding zinc uptake efficiency quality enables assessment of zinc deficiency risk, management of soil and foliar zinc applications, and appreciation of mycorrhizal contributions to zinc nutrition.
Zinc_Uptake_Efficiency_Quality addresses the acquisition of this essential enzyme cofactor affecting auxin metabolism and fruit development. This quality is particularly affected by soil pH, phosphorus status, and mycorrhizal associations. The class supports zinc nutrition management through uptake efficiency characterization identifying deficiency risk factors.
ZIP transporter activity
ZIP-Transporteraktivität
Zinkaufnahmeeffizienz
Zinkversorgung
Zn uptake efficiency
zinc foraging
Sinclair SA, Krämer U (2012) The zinc homeostasis network of land plants. Biochimica et Biophysica Acta 1823(9): 1553-1567 Marschner H (2012) Marschner's Mineral Nutrition of Higher Plants, 3rd Ed. Academic Press, London, pp. 212-229
Zinc Uptake Efficiency Quality
Pinot Noir in Burgundy shows zygote development initiation during late June-early July Year N+1 at BBCH 69, with fertilized zygote undergoing metabolic activation and zygotic genome activation over 18-32 hours post-fertilization, developmental polarity establishing apical-basal axis, DNA replication completing preparing for division, under continental conditions of 22-28°C temperature, typical of this early-to-mid season cultivar completing activation 28-52 hours post-pollination preparing for first embryonic division
Zygote development initiation represents the early activated diploid cell phase immediately following egg fertilization when zygote undergoes metabolic activation, establishes developmental polarity, initiates zygotic gene expression, and prepares for first mitotic division, establishing functional unicellular embryo capable of embryogenesis. This critical activation phase occurs during zygote formation in late June-early July in temperate regions Year N+1, proceeding over 12-36 hours following fertilization before first division, corresponding to BBCH 69 fertilization completion stages. Development initiation involves coordinated processes including sustained metabolic activation from fertilization-triggered changes, developmental polarity establishment creating apical-basal axis, zygotic genome activation initiating embryo-specific gene expression, cell growth and cytoplasmic reorganization preparing division, and cell cycle progression through G1 and S phases toward mitosis. Metabolic activation characteristics show elevated respiratory rate 2-3 fold above unfertilized egg consuming oxygen and producing ATP, increased protein synthesis from maternal mRNAs and newly transcribed zygotic genes, carbohydrate metabolism intensifying mobilizing stored reserves, biosynthetic pathways activating producing membranes and structural components, and overall transition from quiescent to actively growing state. Respiratory activation shows mitochondrial oxidative phosphorylation intensifying, oxygen consumption increasing substantially measurable through respirometry, ATP production supporting biosynthesis and cellular processes, and metabolic rate elevation sustained throughout pre-division period. Protein synthesis shows translation from maternal mRNAs accumulated during oogenesis providing initial developmental proteins, ribosome activation and polysome formation increasing translation capacity, progressive transition to zygotic mRNA translation as genome activates, and protein accumulation supporting growth and division preparation. Zygotic genome activation shows transition from maternal to embryonic genetic control, initial zygotic transcription occurring within hours of fertilization, progressive increase in zygotic gene expression over pre-division period, embryo-specific genes activating including developmental regulators and cell cycle controllers, and genome activation establishing embryonic developmental program. Transcriptional activation timing shows early zygotic genes expressed within 2-6 hours post-fertilization, progressive activation of additional genes over 12-24 hours, and major transition to predominantly zygotic control by first division, though timing varies among species and specific genes. Developmental gene expression shows transcription factors regulating embryo patterning, cell cycle regulators controlling division, metabolic enzymes supporting growth, and signaling molecules coordinating development. Chromatin remodeling shows modifications enabling transcriptional activation, histone modifications marking active versus repressed genomic regions, DNA methylation changes affecting gene expression patterns, and epigenetic regulation establishing developmental programs. Developmental polarity establishment shows asymmetric distribution of developmental regulators creating apical-basal axis, cytoplasmic gradients determining positional information, differential localization of transcription factors and mRNAs, and polarity determining embryo organization in subsequent development. Polarity axis orientation shows typical apical pole toward micropyle and basal pole toward chalaza corresponding to embryo proper and suspensor regions respectively, axis established during early post-fertilization period possibly influenced by sperm entry site or inherent egg organization, and axis maintained through subsequent divisions determining embryo structure. Molecular asymmetry shows differential distribution of key developmental regulators including transcription factors affecting cell fate, localized mRNAs encoding developmental proteins, and protein gradients establishing positional information along apical-basal axis. Cytoplasmic reorganization shows organelle repositioning with mitochondria, ER, and Golgi distributing asymmetrically, cytoskeletal rearrangement creating structural polarity, and cytoplasmic streaming possibly contributing to molecular redistribution. Cell growth during initiation shows zygote enlargement through water uptake and biosynthesis, volume increase up to 1.5-2 fold before division, cell wall expansion accommodating growth, and resource accumulation from endosperm nutritive support. Cell wall modification shows loosening enabling expansion, new wall synthesis accommodating growth, and wall properties maintaining structural integrity while allowing enlargement. Nucleus characteristics show diploid nucleus with decondensed interphase chromatin, active transcription with RNA polymerase activity, nucleolus reformation and ribosomal RNA synthesis, and nuclear growth correlating with cell enlargement. Cell cycle progression shows G1 phase with growth and preparation for DNA replication, S phase entry with DNA synthesis initiating typically 12-24 hours post-fertilization, DNA replication proceeding doubling genome from 2n to 4n DNA content, G2 phase with continued growth and mitotic preparation, and progression toward M phase and first division. DNA replication dynamics show replication origin firing initiating synthesis, replication fork progression along chromosomes, coordinated replication ensuring complete genome duplication, and S phase completion within 6-12 hours enabling progression to G2. Mitotic preparation shows chromatin condensation beginning in late G2, centrosome duplication preparing spindle formation, nuclear envelope breakdown approaching as M phase nears, and cytological changes indicating imminent division. Environmental requirements during initiation include optimal temperature 22-28°C supporting normal metabolic and developmental processes, adequate nutrient supply from surrounding endosperm providing resources for growth and synthesis, appropriate osmotic environment maintaining cellular hydration, and overall favorable conditions during critical 12-36 hour initiation period. Temperature effects on initiation show warm conditions promoting rapid progression through initiation with accelerated metabolism and gene expression, cool conditions slowing processes potentially extending initiation period, and temperature as major factor affecting developmental rate. Endosperm support shows nutritive tissue providing amino acids, sugars, lipids, and other resources, signaling molecules possibly coordinating zygote-endosperm development, and essential support for normal zygote growth and division preparation. Failed development initiation causes include genetic defects preventing normal activation or gene expression, inadequate metabolic activation failing to support growth, abnormal polarity establishment disrupting organization, insufficient endosperm support limiting resources, or environmental stress disrupting developmental processes. Development initiation success rate shows high success in normal fertilized zygotes with majority successfully activating and progressing toward division, genetic or environmental problems causing minority failures, and successful initiation essential for subsequent embryogenesis. The process represents first aspect of zygote formation emphasizing activation and preparation, preceding embryo genesis start marking first division. Duration typically 12-36 hours from fertilization to division onset. Cultivar variation may include activation timing, developmental rate, and gene expression patterns. Regional timing shows zygote development initiation during late June-early July across all climates within day following fertilization. Viticultural significance includes representing critical activation establishing embryo developmental program, requiring successful metabolic and genetic activation, demonstrating temperature sensitivity affecting developmental timing, and establishing initiation success as prerequisite for embryogenesis and seed formation. Research applications include analyzing metabolic activation mechanisms and kinetics, investigating zygotic genome activation and transcriptional programs, evaluating polarity establishment and molecular asymmetry, quantifying cell cycle progression and division preparation, and assessing developmental initiation as determinant of embryo development outcomes.
Zygote_Development_Initiation is first aspect of Zygote_Formation, distinguished by activation and preparation phase. Represents metabolic activation, genome activation, polarity establishment. Duration 12-36 hours post-fertilization. Involves transition from maternal to zygotic genetic control. Temperature affects developmental rate. Establishes embryonic program. Preceded by Egg_Fertilization. Followed by Embryo_Genesis_Start as division approaches. Critical phase determining embryo developmental competence.
Entwicklungsprogramm-Initiierung
Zygoten-Aktivierung
Zygoten-Entwicklungs-Initiierung
developmental program initiation
early zygote development
frühe Zygoten-Entwicklung
post-fertilization activation
zygote activation
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 398-400 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 302-304
Zygote Development Initiation
Pinot Noir in Burgundy shows zygote division during late June-early July Year N+1, with diploid zygote 2n=38 dividing 32-52 hours post-fertilization at 22-26°C, asymmetric division producing apical cell 24-36 micrometers and basal cell 46-58 micrometers, apical cell specified for embryo proper fate, basal cell forming suspensor, two-cell proembryo establishing apical-basal axis, under continental conditions initiating embryogenesis, typical of this early-to-mid season cultivar beginning seed development
Zygote division represents the first embryonic cell division phase during embryo development when diploid zygote 2n=38 undergoes asymmetric division producing two-cell proembryo with distinct apical and basal cells, establishing apical-basal body axis, creating developmental foundation with apical cell forming embryo proper and basal cell forming suspensor, and initiating multicellular embryogenesis, occurring 24-72 hours post-fertilization depending on temperature as developmental transition from single-cell to two-cell stage. This first division phase occurs during early summer in late June-early July in temperate regions Year N+1, representing zygote to embryo transition 1-3 days post-fertilization. Division mechanism involves zygote activation and cell cycle progression following fertilization, DNA replication doubling chromosome content from 2n to 4n, mitotic apparatus formation with spindle positioning determining division plane, asymmetric cytokinesis producing unequal daughter cells, apical cell specification for embryo fate, basal cell specification for suspensor fate, and division establishing developmental program. Zygote developmental state shows zygote formed from sperm-egg fusion creating diploid 2n=38 single cell, zygote polarized with apical-basal axis from egg polarity or post-fertilization establishment, zygote cell typically 40-70 micrometers diameter, metabolically active preparing for division, and zygote as embryogenesis starting point. Zygote activation shows fertilization triggering metabolic activation, calcium signaling initiating developmental program, zygotic genome activation with embryo-specific genes beginning expression, transition from maternal to embryonic genetic control, and activation establishing autonomous development. Zygote cell cycle shows G1 phase with growth and preparation, S phase DNA replication creating 4n chromosome content, G2 phase with mitotic preparation and spindle assembly, M phase mitosis with chromosome segregation, and cytokinesis completing division. Cell cycle timing shows division occurring 24-72 hours post-fertilization depending on temperature, warm conditions 25-28°C enabling division 24-36 hours, cool conditions 18-22°C extending to 48-72 hours, and temperature as major timing determinant. DNA replication shows semiconservative replication duplicating 475 Mb grapevine genome, replication producing sister chromatids, chromosome content doubling to 4n, and replication as division prerequisite. Mitotic spindle formation shows centrosomes nucleating microtubules, bipolar spindle forming with poles at cell ends, kinetochores on chromosomes attaching to spindle microtubules, and spindle providing chromosome segregation machinery. Spindle positioning shows spindle axis orientation determining division plane, asymmetric positioning creating unequal division, spindle position regulated by cortical cues and cytoskeletal forces, and positioning as asymmetry mechanism. Asymmetric division shows unequal cytokinesis producing larger basal cell 40-60 micrometers and smaller apical cell 20-40 micrometers, asymmetry from unequal spindle positioning and cytokinesis plane, asymmetric distribution of developmental regulators possibly influencing fate, and asymmetry establishing cell fate difference. Cytokinesis characteristics shows phragmoplast forming at division plane, cell plate synthesizing from center outward, new cell wall forming between daughter cells, and cytokinesis completing division creating two-cell proembryo. Two-cell proembryo characteristics shows apical cell smaller typically 20-40 micrometers diameter positioned toward micropyle, basal cell larger 40-60 micrometers positioned toward chalaza, cells separated by new wall, and proembryo as two-cell embryo stage. Apical cell fate shows apical cell destined to form embryo proper including all seedling structures, cell undergoing subsequent divisions creating embryo, cell containing embryo developmental program, and apical fate as primary embryo pathway. Basal cell fate shows basal cell forming suspensor supporting structure, suspensor typically 3-7 cells anchoring embryo in endosperm, suspensor possibly transferring nutrients though function debated in grapevine, suspensor degenerating during later development, and basal fate as support pathway. Apical-basal axis establishment shows division creating primary body axis, axis orientation from micropyle apical to chalaza basal, axis persisting through development into seedling root-shoot axis, and axis as fundamental body plan feature. Developmental asymmetry shows unequal cells having different developmental fates, asymmetry as mechanism for creating embryo organization, asymmetric division common in plant embryogenesis, and asymmetry establishing pattern. Cell fate determination shows molecular mechanisms specifying apical versus basal fate possibly involving transcription factors and signaling molecules though poorly characterized in grapevine, fate determination early in development, and determination creating distinct pathways. First division significance shows division as embryogenesis initiation, transition from single-cell to multicellular organism, establishment of body axis and cell fates, and division as critical developmental event. Environmental effects on division include temperature affecting division timing with warm conditions accelerating and cool conditions delaying within 24-72 hour range, temperature as major factor, and division occurring across temperature ranges enabling development. Division timing variation shows cultivar and individual variation in timing, typical range 24-72 hours covering temperature and genetic effects, and timing determining subsequent development pace. Failed division shows occasional zygotes failing to divide from developmental abnormalities, failure causing ovule abortion, and failure as early developmental loss. The process represents first phase of embryo development establishing two-cell proembryo, preceding embryo morphogenesis. Duration represents single division event occurring over hours at 24-72 hours post-fertilization. Regional timing shows zygote division during late June-early July as embryogenesis initiates. Viticultural significance includes representing embryogenesis initiation establishing multicellular embryo, creating apical-basal axis and cell fate asymmetry, determining subsequent embryo development, and marking reproductive success transition from fertilization to development. Research applications include analyzing zygote activation and developmental initiation, investigating asymmetric division mechanisms and cell fate determination, evaluating environmental effects on division timing, quantifying division characteristics and outcomes, and assessing zygote division as embryogenesis foundation.
Zygote_Division is first phase of Embryo_Development, distinguished by first asymmetric division. Represents single division event 24-72 hours post-fertilization (temperature-dependent). Produces apical cell 20-40 μm (embryo proper fate) and basal cell 40-60 μm (suspensor fate). Establishes apical-basal axis. Creates two-cell proembryo. Followed by Embryo_Morphogenesis. Critical embryogenesis initiation. Temperature 25-28°C optimal (24-36h), cool 18-22°C extends (48-72h).
Proembryo-Bildung
Zygoten-Teilung
initial embryo division
proembryo formation
Pratt C (1971) Reproductive anatomy in cultivated grapes, Am J Enol Vitic 22:92-109 Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 438-440
Zygote Division
Cabernet Sauvignon in Napa Valley shows zygote formation during late June-early July Year N+1 at BBCH 69, with diploid zygote developing over 30-60 hours post-fertilization, developmental polarity establishing, first mitotic division occurring 36-52 hours post-fertilization producing two-cell embryo, under warm climate conditions of 24-29°C temperature, typical of this late-season cultivar completing fertilization 28-42 hours post-pollination and initiating embryogenesis 60-90 hours post-pollination
Zygote formation represents the diploid single-cell embryo establishment phase following egg fertilization when sperm-egg fusion creates diploid nucleus, egg activation initiates developmental program, and first zygotic division preparations proceed, establishing unicellular embryo that will develop into mature plant through embryogenesis. This critical embryo initiation phase occurs during late fertilization in mid-late June in temperate regions Year N+1, proceeding over hours to days following egg fertilization approximately 24-72 hours after initial pollination, corresponding to BBCH 69 fertilization completion stages transitioning to early berry development. Formation process involves diploid zygote nucleus establishment through sperm-egg nuclear fusion, zygote activation triggered by calcium signaling and metabolic changes, developmental polarity establishment determining future embryo organization, cell cycle reentry preparing for first mitotic division, and zygote growth and preparation for cleavage creating two-cell embryo initiating embryogenesis. Zygote characteristics show single diploid cell containing complete genetic information from both parents, 2n=38 chromosomes in Vitis vinifera representing fusion of two 1n=19 gametes, large cell typically 40-80 micrometers diameter containing abundant cytoplasm and organelles, developmental totipotency with capacity to generate entire plant, and position within embryo sac at micropylar pole adjacent to penetrated synergid. Diploid nucleus structure shows nuclear envelope surrounding combined maternal and paternal chromatin, interphase chromatin organization with chromosomes decondensed, active transcription initiating zygotic gene expression, and nucleus containing complete genomic complement enabling embryo development. Cytoplasmic organization shows abundant ribosomes for protein synthesis, mitochondria providing energy for metabolism and division, endoplasmic reticulum and Golgi apparatus supporting secretory functions, stored reserves including lipids and proteins providing resources for early development, and maternally inherited organelles including mitochondria and plastids transmitting through egg. Zygote activation continuation shows sustained calcium signaling maintaining active state, metabolic activation with respiration and biosynthesis intensifying, protein synthesis from maternal mRNAs accumulated during oogenesis, transition to zygotic genome activation expressing embryo developmental genes, and cell cycle progression preparing for division. Developmental polarity establishment shows asymmetric cytoplasmic organization creating apical-basal axis, differential distribution of developmental regulators establishing positional information, molecular gradients determining cell fate in first division, and polarity determining embryo proper versus suspensor formation in subsequent cleavage. Apical-basal axis determination shows polarity established during or shortly after fertilization, axis orientation typically with apical pole toward micropyle and basal pole toward chalaza, and axis determining embryo structural organization through subsequent development. Cell fate determinants show asymmetric distribution of transcription factors and signaling molecules, localized mRNAs and proteins creating developmental gradients, and molecular asymmetry determining whether daughter cells develop into embryo proper or suspensor in first division. Cell cycle reentry shows exit from meiotic metaphase II arrest completing egg meiosis, G1 phase entry following fertilization, S phase initiation with DNA replication preparing for division, and progression through cell cycle toward first mitotic division typically 24-48 hours post-fertilization. DNA replication shows chromosome duplication during S phase, genome doubling from 2n to 4n DNA content preparing for mitosis, replication fork progression through entire genome, and completion establishing readiness for chromosome segregation in division. Mitotic preparation shows chromatin condensation into visible chromosomes, nuclear envelope breakdown releasing chromosomes into cytoplasm, spindle formation with microtubule arrays establishing chromosome segregation machinery, and progression toward metaphase alignment and anaphase separation. First division timing shows variable timing among species and individuals with typical first cleavage 24-48 hours post-fertilization in favorable conditions, temperature-dependent timing with warm conditions accelerating and cool conditions delaying, and division establishing two-cell embryo marking definitive embryogenesis initiation. Zygote growth before division shows cell enlargement through water uptake and biosynthesis, volume increase up to 1.5-2 fold before cleavage, and resource accumulation supporting subsequent divisions. Environmental effects on zygote formation include temperature critically affecting developmental rate with optimal 22-28°C supporting normal progression, nutrient availability from endosperm supporting growth and metabolism, appropriate hydration maintaining cellular function, and overall favorable conditions during first days post-fertilization critical for normal embryo initiation. Temperature sensitivity shows warm conditions promoting rapid development with first division within 24-36 hours, cool conditions substantially delaying division to 48-72 hours or more, excessive heat potentially disrupting development, and temperature as major factor affecting embryogenesis timing. Osmotic environment shows surrounding endosperm providing appropriate osmotic conditions, proper balance essential for zygote hydration and growth, and osmotic stress potentially disrupting development. Zygote developmental competence shows inherent capacity to develop into mature embryo, competence depending on genetic integrity and proper fertilization, developmental abnormalities from chromosomal aberrations or genetic defects causing embryo abortion, and majority of normal zygotes successfully initiating embryogenesis under favorable conditions. Failed zygote formation causes include abnormal fertilization producing aneuploid or polyploid zygotes with developmental incompetence, genetic defects preventing normal development, inadequate endosperm support failing to provide necessary resources, environmental stress during critical formation period disrupting development, or developmental arrest from unknown causes. Zygote to embryo transition shows first division as definitive transition from unicellular zygote to multicellular embryo, progressive cell divisions elaborating embryo structure, and continuing development through globular, heart, torpedo, and mature embryo stages over subsequent weeks. The process shows two distinguishable aspects progressing from zygote development initiation establishing diploid cell and activation, through embryo genesis start marking first division and embryogenesis. Duration typically spans 24-72 hours from fertilization to first division depending on conditions. Cultivar variation in zygote formation may include division timing, developmental rate, and embryo developmental characteristics. Regional timing shows zygote formation during late June-early July across all climates 1-3 days following fertilization as embryo development initiates. Viticultural significance includes representing unicellular embryo stage initiating seed development, requiring successful fertilization and activation, demonstrating environmental sensitivity affecting developmental timing, establishing zygote formation as determinant of embryo and seed viability, and marking transition from fertilization to embryogenesis phase. Research applications include analyzing zygote activation mechanisms and developmental programs, investigating polarity establishment and cell fate determination, evaluating environmental effects on zygote development and division timing, quantifying zygote to embryo transition dynamics, and assessing zygote formation as determinant of seed development outcomes.
Zygote_Formation represents diploid unicellular embryo stage following Egg_Fertilization. Subdivided into Zygote_Development_Initiation establishing activated diploid cell and Embryo_Genesis_Start marking first division. Duration 24-72 hours from fertilization to first cleavage. Involves developmental activation, polarity establishment, cell cycle reentry. Temperature strongly affects timing. First division marks transition to multicellular embryo. Essential stage for seed development. Determines embryo developmental potential.
Zygoten-Entwicklung
Zygoten-Formation
befruchtete Eizell-Entwicklung
diploid cell formation
einzelliger Embryo
fertilized egg development
unicellular embryo
zygote development
Keller M (2020) The Science of Grapevines 3rd Ed, Academic Press pp. 398-402 Mullins MG et al. (1992) Biology of the Grapevine, Cambridge University Press pp. 302-305
Zygote Formation